Chapter X. "on the Imperfection of the Geological Record," and Chapter

XI. "On the Geological Succession of Organic Beings"; there is, however, little about plants in these chapters. At the present time the truth of Evolution is no longer seriously disputed, though there are writers, like Reinke, who insist, and rightly so, that the doctrine is still only a belief, rather than an established fact of science. (J. Reinke, "Kritische Abstammungslehre", "Wiesner-Festschrift", page 11, Vienna, 1908.) Evidently, then, however little the Theory of Descent may be questioned in our own day, it is desirable to assure ourselves how the case stands, and in particular how far the evidence from fossil plants has grown stronger with time.

As regards direct evidence for the derivation of one species from another, there has probably been little advance since Darwin wrote, at least so we must infer from the emphasis laid on the discontinuity of successive fossil species by great systematic authorities like Grand'Eury and Zeiller in their most recent writings. We must either adopt the mutationist views of those authors (referred to in the last section of this essay) or must still rely on Darwin's explanation of the absence of numerous intermediate varieties. The attempts which have been made to trace, in the Tertiary rocks, the evolution of recent species, cannot, owing to the imperfect character of the evidence, be regarded as wholly satisfactory.

When we come to groups of a somewhat higher order we have an interesting history of the evolution of a recent family in the work, not yet completed, of Kidston and Gwynne-Vaughan on the fossil Osmundaceae. ("Trans. Royal Soc. Edinburgh", Vol. 45, Part III. 1907, Vol. 46, Part II. 1908, Vol. 46, Part III. 1909.) The authors are able, mainly on anatomical evidence, to trace back this now limited group of Ferns, through the Tertiary and Mesozoic to the Permian, and to show, with great probability, how their structure has been derived from that of early Palaeozoic types.

The history of the Ginkgoaceae, now represented only by the isolated maidenhair tree, scarcely known in a wild state, offers another striking example of a family which can be traced with certainty to the older Mesozoic and perhaps further back still. (See Seward and Gowan, "The Maidenhair Tree (Gingko biloba)", "Annals of Botany", Vol. XIV. 1900, page 109; also A. Sprecher "Le Ginkgo biloba", L., Geneva, 1907.)

On the wider question of the derivation of the great groups of plants, a very considerable advance has been made, and, so far as the higher plants are concerned, we are now able to form a far better conception than before of the probable course of evolution. This is a matter of phylogeny, and the facts will be considered under that head; our immediate point is that the new knowledge of the relations between the classes of plants in question materially strengthens the case for the theory of descent. The discoveries of the last few years throw light especially on the relation of the Angiosperms to the Gymnosperms, on that of the Seed-plants generally to the Ferns, and on the interrelations between the various classes of the higher Cryptogams.

That the fossil record has not done still more for Evolution is due to the fact that it begins too late--a point on which Darwin laid stress ("Origin of Species" (6th edition), page 286.) and which has more recently been elaborated by Poulton. ("Essays on Evolution", pages 46 et seq., Oxford, 1908.) An immense proportion of the whole evolutionary history lies behind the lowest fossiliferous rocks, and the case is worse for plants than for animals, as the record for the former begins, for all practical purposes, much higher up in the rocks.

It may be well here to call attention to a question, often overlooked, which has lately been revived by Reinke. (Reinke, loc. cit. page 13.) As all admit, we know nothing of the origin of life; consequently, for all we can tell, it is as probable that life began, on this planet, with many living things, as with one. If the first organic beings were many, they may have been heterogeneous, or at least exposed to different conditions, from their origin; in either case there would have been a number of distinct series from the beginning, and if so we should not be justified in assuming that all organisms are related to one another. There may conceivably be several of the original lines of descent still surviving, or represented among extinct forms--to reverse the remark of a distinguished botanist, there may be several Vegetable Kingdoms! However improbable this may sound, the possibility is one to be borne in mind.

That all VASCULAR plants really belong to one stock seems certain, and here the palaeontological record has materially strengthened the case for a monophyletic history. The Bryophyta are not likely to be absolutely distinct, for their sexual organs, and the stomata of the Mosses strongly suggest community of descent with the higher plants; if this be so it no doubt establishes a certain presumption in favour of a common origin for plants generally, for the gap between "Mosses and Ferns" has been regarded as the widest in the Vegetable Kingdom. The direct evidence of consanguinity is however much weaker when we come to the Algae, and it is conceivable (even if improbable) that the higher plants may have had a distinct ancestry (now wholly lost) from the beginning. The question had been raised in Darwin's time, and he referred to it in these words: "No doubt it is possible, as Mr G.H. Lewes has urged, that at the first commencement of life many different forms were evolved; but if so, we may conclude that only a very few have left modified descendants." ("Origin of Species", page 425.) This question, though it deserves attention, does not immediately affect the subject of the palaeontological record of plants, for there can be no reasonable doubt as to the interrelationship of those groups on which the record at present throws light.

The past history of plants by no means shows a regular progression from the simple to the complex, but often the contrary. This apparent anomaly is due to two causes.

1. The palaeobotanical record is essentially the story of the successive ascendancy of a series of dominant families, each of which attained its maximum, in organisation as well as in extent, and then sank into comparative obscurity, giving place to other families, which under new conditions were better able to take a leading place. As each family ran its downward course, either its members underwent an actual reduction in structure as they became relegated to herbaceous or perhaps aquatic life (this may have happened with the Horsetails and with Isoetes if derived from Lepidodendreae), or the higher branches of the family were crowded out altogether and only the "poor relations" were able to maintain their position by evading the competition of the ascendant races; this is also illustrated by the history of the Lycopod phylum. In either case there would result a lowering of the type of organisation within the group.

2. The course of real progress is often from the complex to the simple. If, as we shall find some grounds for believing, the Angiosperms came from a type with a flower resembling in its complexity that of Mesozoic "Cycads," almost the whole evolution of the flower in the highest plants has been a process of reduction. The stamen, in particular, has undoubtedly become extremely simplified during evolution; in the most primitive known seed-plants it was a highly compound leaf or pinna; its reduction has gone on in the Conifers and modern Cycads, as well as in the Angiosperms, though in different ways and to a varying extent.

The seed offers another striking example; the Palaeozoic seeds (if we leave the seed-like organs of certain Lycopods out of consideration) were always, so far as we know, highly complex structures, with an elaborate vascular system, a pollen-chamber, and often a much-differentiated testa. In the present day such seeds exist only in a few Gymnosperms which retain their ancient characters--in all the higher Spermophytes the structure is very much simplified, and this holds good even in the Coniferae, where there is no countervailing complication of ovary and stigma.

Reduction, in fact, is not always, or even generally, the same thing as degeneration. Simplification of parts is one of the most usual means of advance for the organism as a whole. A large proportion of the higher plants are microphyllous in comparison with the highly megaphyllous fern-like forms from which they appear to have been derived.

Darwin treated the general question of advance in organisation with much caution, saying: "The geological record... does not extend far enough back, to show with unmistakeable clearness that within the known history of the world organisation has largely advanced." ("Origin of Species", page 308.) Further on (Ibid. page 309.) he gives two standards by which advance may be measured: "We ought not solely to compare the highest members of a class at any two periods... but we ought to compare all the members, high and low, at the two periods." Judged by either standard the Horsetails and Club Mosses of the Carboniferous were higher than those of our own day, and the same is true of the Mesozoic Cycads. There is a general advance in the succession of classes, but not within each class.

Darwin's argument that "the inhabitants of the world at each successive period in its history have beaten their predecessors in the race for life, and are, in so far, higher in the scale" ("Origin of Species", page 315.) is unanswerable, but we must remember that "higher in the scale" only means "better adapted to the existing conditions." Darwin points out (Ibid. page 279.) that species have remained unchanged for long periods, probably longer than the periods of modification, and only underwent change when the conditions of their life were altered. Higher organisation, judged by the test of success, is thus purely relative to the changing conditions, a fact of which we have a striking illustration in the sudden incoming of the Angiosperms with all their wonderful floral adaptations to fertilisation by the higher families of Insects.

II. PHYLOGENY.

The question of phylogeny is really inseparable from that of the truth of the doctrine of evolution, for we cannot have historical evidence that evolution has actually taken place without at the same time having evidence of the course it has followed.

As already pointed out, the progress hitherto made has been rather in the way of joining up the great classes of plants than in tracing the descent of particular species or genera of the recent flora. There appears to be a difference in this respect from the Animal record, which tells us so much about the descent of living species, such as the elephant or the horse. The reason for this difference is no doubt to be found in the fact that the later part of the palaeontological record is the most satisfactory in the case of animals and the least so in the case of plants. The Tertiary plant-remains, in the great majority of instances, are impressions of leaves, the conclusions to be drawn from which are highly precarious; until the whole subject of Angiospermous palaeobotany has been reinvestigated, it would be rash to venture on any statements as to the descent of the families of Dicotyledons or Monocotyledons.

Our attention will be concentrated on the following questions, all relating to the phylogeny of main groups of plants: i. The Origin of the Angiosperms. ii. The Origin of the Seed-plants. iii. The Origin of the different classes of the Higher Cryptogamia.

i. THE ORIGIN OF THE ANGIOSPERMS.

The first of these questions has long been the great crux of botanical phylogeny, and until quite recently no light had been thrown upon the difficulty. The Angiosperms are the Flowering Plants, par excellence, and form, beyond comparison, the dominant sub-kingdom in the flora of our own age, including, apart from a few Conifers and Ferns, all the most familiar plants of our fields and gardens, and practically all plants of service to man. All recent work has tended to separate the Angiosperms more widely from the other seed-plants now living, the Gymnosperms. Vast as is the range of organisation presented by the great modern sub-kingdom, embracing forms adapted to every environment, there is yet a marked uniformity in certain points of structure, as in the development of the embryo-sac and its contents, the pollination through the intervention of a stigma, the strange phenomenon of double fertilisation (One sperm fertilising the egg, while the other unites with the embryo-sac nucleus, itself the product of a nuclear fusion, to give rise to a nutritive tissue, the endosperm.), the structure of the stamens, and the arrangement of the parts of the flower. All these points are common to Monocotyledons and Dicotyledons, and separate the Angiosperms collectively from all other plants.

In geological history the Angiosperms first appear in the Lower Cretaceous, and by Upper Cretaceous times had already swamped all other vegetation and seized the dominant position which they still hold. Thus they are isolated structurally from the rest of the Vegetable Kingdom, while historically they suddenly appear, almost in full force, and apparently without intermediaries with other groups. To quote Darwin's vigorous words: "The rapid development, as far as we can judge, of all the higher plants within recent geological times is an abominable mystery." ("More Letters of Charles Darwin", Vol. II. page 20, letter to J.D. Hooker, 1879.) A couple of years later he made a bold suggestion (which he only called an "idle thought") to meet this difficulty. He says: "I have been so astonished at the apparently sudden coming in of the higher phanerogams, that I have sometimes fancied that development might have slowly gone on for an immense period in some isolated continent or large island, perhaps near the South Pole." (Ibid, page 26, letter to Hooker, 1881.) This idea of an Angiospermous invasion from some lost southern land has sometimes been revived since, but has not, so far as the writer is aware, been supported by evidence. Light on the problem has come from a different direction.

The immense development of plants with the habit of Cycads, during the Mesozoic Period up to the Lower Cretaceous, has long been known. The existing Order Cycadaceae is a small family, with 9 genera and perhaps 100 species, occurring in the tropical and sub-tropical zones of both the Old and New World, but nowhere forming a dominant feature in the vegetation. Some few attain the stature of small trees, while in the majority the stem is short, though often living to a great age. The large pinnate or rarely bipinnate leaves give the Cycads a superficial resemblance in habit to Palms. Recent Cycads are dioecious; throughout the family the male fructification is in the form of a cone, each scale of the cone representing a stamen, and bearing on its lower surface numerous pollen-sacs, grouped in sori like the sporangia of Ferns. In all the genera, except Cycas itself, the female fructifications are likewise cones, each carpel bearing two ovules on its margin. In Cycas, however, no female cone is produced, but the leaf-like carpels, bearing from two to six ovules each, are borne directly on the main stem of the plant in rosettes alternating with those of the ordinary leaves--the most primitive arrangement known in any living seed-plant. The whole Order is relatively primitive, as shown most strikingly in its cryptogamic mode of fertilisation, by means of spermatozoids, which it shares with the maidenhair tree alone, among recent seed-plants.

In all the older Mesozoic rocks, from the Trias to the Lower Cretaceous, plants of the Cycad class (Cycadophyta, to use Nathorst's comprehensive name) are extraordinarily abundant in all parts of the world; in fact they were almost as prominent in the flora of those ages as the Dicotyledons are in that of our own day. In habit and to a great extent in anatomy, the Mesozoic Cycadophyta for the most part much resemble the recent Cycadaceae. But, strange to say, it is only in the rarest cases that the fructification has proved to be of the simple type characteristic of the recent family; the vast majority of the abundant fertile specimens yielded by the Mesozoic rocks possess a type of reproductive apparatus far more elaborate than anything known in Cycadaceae or other Gymnosperms. The predominant Mesozoic family, characterised by this advanced reproductive organisation, is known as the Bennettiteae; in habit these plants resembled the more stunted Cycads of the recent flora, but differed from them in the presence of numerous lateral fructifications, like large buds, borne on the stem among the crowded bases of the leaves. The organisation of these fructifications was first worked out on European specimens by Carruthers, Solms-Laubach, Lignier and others, but these observers had only more or less ripe fruits to deal with; the complete structure of the flower has only been elucidated within the last few years by the researches of Wieland on the magnificent American material, derived from the Upper Jurassic and Lower Cretaceous beds of Maryland, Dakota and Wyoming. (G.R. Wieland, "American Fossil Cycads", Carnegie Institution, Washington, 1906.) The word "flower" is used deliberately, for reasons which will be apparent from the following brief description, based on Wieland's observations.

The fructification is attached to the stem by a thick stalk, which, in its upper part, bears a large number of spirally arranged bracts, forming collectively a kind of perianth and completely enclosing the essential organs of reproduction. The latter consist of a whorl of stamens, of extremely elaborate structure, surrounding a central cone or receptacle bearing numerous ovules. The stamens resemble the fertile fronds of a fern; they are of a compound, pinnate form, and bear very large numbers of pollen-sacs, each of which is itself a compound structure consisting of a number of compartments in which the pollen was formed. In their lower part the stamens are fused together by their stalks, like the "monadelphous" stamens of a mallow. The numerous ovules borne on the central receptacle are stalked, and are intermixed with sterile scales; the latter are expanded at their outer ends, which are united to form a kind of pericarp or ovary-wall, only interrupted by the protruding micropyles of the ovules. There is thus an approach to the closed pistil of an Angiosperm, but it is evident that the ovules received the pollen directly. The whole fructification is of large size; in the case of Cycadeoidea dacotensis, one of the species investigated by Wieland, the total length, in the bud condition, is about 12 cm., half of which belongs to the peduncle.

The general arrangement of the organs is manifestly the same as in a typical Angiospermous flower, with a central pistil, a surrounding whorl of stamens and an enveloping perianth; there is, as we have seen, some approach to the closed ovary of an Angiosperm; another point, first discovered nearly 20 years ago by Solms-Laubach in his investigation of a British species, is that the seed was practically "exalbuminous," its cavity being filled by the large, dicotyledonous embryo, whereas in all known Gymnosperms a large part of the sac is occupied by a nutritive tissue, the prothallus or endosperm; here also we have a condition only met with elsewhere among the higher Flowering Plants.

Taking all the characters into account, the indications of affinity between the Mesozoic Cycadophyta and the Angiosperms appear extremely significant, as was recognised by Wieland when he first discovered the hermaphrodite nature of the Bennettitean flower. The Angiosperm with which he specially compared the fossil type was the Tulip tree (Liriodendron) and certainly there is a remarkable analogy with the Magnoliaceous flowers, and with those of related orders such as Ranunculaceae and the Water-lilies. It cannot, of course, be maintained that the Bennettiteae, or any other Mesozoic Cycadophyta at present known, were on the direct line of descent of the Angiosperms, for there are some important points of difference, as, for example, in the great complexity of the stamens, and in the fact that the ovary-wall or pericarp was not formed by the carpels themselves, but by the accompanying sterile scale-leaves. Botanists, since the discovery of the bisexual flowers of the Bennettiteae, have expressed different views as to the nearness of their relation to the higher Flowering Plants, but the points of agreement are so many that it is difficult to resist the conviction that a real relation exists, and that the ancestry of the Angiosperms, so long shrouded in complete obscurity, is to be sought among the great plexus of Cycad-like plants which dominated the flora of the world in Mesozoic times. (On this subject see, in addition to Wieland's great work above cited, F.W. Oliver, "Pteridosperms and Angiosperms", "New Phytologist", Vol. V. 1906; D.H. Scott, "The Flowering Plants of the Mesozoic Age in the Light of Recent Discoveries", "Journal R. Microscop. Soc." 1907, and especially E.A.N. Arber and J. Parkin, "On the Origin of Angiosperms", "Journal Linn. Soc." (Bot.) Vol. XXXVIII. page 29, 1907.)

The great complexity of the Bennettitean flower, the earliest known fructification to which the word "flower" can be applied without forcing the sense, renders it probable, as Wieland and others have pointed out, that the evolution of the flower in Angiosperms has consisted essentially in a process of reduction, and that the simplest forms of flower are not to be regarded as the most primitive. The older morphologists generally took the view that such simple flowers were to be explained as reductions from a more perfect type, and this opinion, though abandoned by many later writers, appears likely to be true when we consider the elaboration of floral structure attained among the Mesozoic Cycadophyta, which preceded the Angiosperms in evolution.

If, as now seems probable, the Angiosperms were derived from ancestors allied to the Cycads, it would naturally follow that the Dicotyledons were first evolved, for their structure has most in common with that of the Cycadophyta. We should then have to regard the Monocotyledons as a side-line, diverging probably at a very early stage from the main dicotyledonous stock, a view which many botanists have maintained, of late, on other grounds. (See especially Ethel Sargant, "The Reconstruction of a Race of Primitive Angiosperms", "Annals of Botany", Vol. XXII. page 121, 1908.) So far, however, as the palaeontological record shows, the Monocotyledons were little if at all later in their appearance than the Dicotyledons, though always subordinate in numbers. The typical and beautifully preserved Palm-wood from Cretaceous rocks is striking evidence of the early evolution of a characteristic monocotyledonous family. It must be admitted that the whole question of the evolution of Monocotyledons remains to be solved.

Accepting, provisionally, the theory of the cycadophytic origin of Angiosperms, it is interesting to see to what further conclusions we are led. The Bennettiteae, at any rate, were still at the gymnospermous level as regards their pollination, for the exposed micropyles of the ovules were in a position to receive the pollen directly, without the intervention of a stigma. It is thus indicated that the Angiosperms sprang from a gymnospermous source, and that the two great phyla of Seed-plants have not been distinct from the first, though no doubt the great majority of known Gymnosperms, especially the Coniferae, represent branch-lines of their own.

The stamens of the Bennettiteae are arranged precisely as in an angiospermous flower, but in form and structure they are like the fertile fronds of a Fern, in fact the compound pollen-sacs, or synangia as they are technically called, almost exactly agree with the spore-sacs of a particular family of Ferns--the Marattiaceae, a limited group, now mainly tropical, which was probably more prominent in the later Palaeozoic times than at present. The scaly hairs, or ramenta, which clothe every part of the plant, are also like those of Ferns.

It is not likely that the characters in which the Bennettiteae resemble the Ferns came to them directly from ancestors belonging to that class; an extensive group of Seed-plants, the Pteridospermeae, existed in Palaeozoic times and bear evident marks of affinity with the Fern phylum. The fern-like characters so remarkably persistent in the highly organised Cycadophyta of the Mesozoic were in all likelihood derived through the Pteridosperms, plants which show an unmistakable approach to the cycadophytic type.

The family Bennettiteae thus presents an extraordinary association of characters, exhibiting, side by side, features which belong to the Angiosperms, the Gymnosperms and the Ferns.

ii. ORIGIN OF SEED-PLANTS.

The general relation of the gymnospermous Seed-plants to the Higher Cryptogamia was cleared up, independently of fossil evidence, by the brilliant researches of Hofmeister, dating from the middle of the past century. (W. Hofmeister, "On the Germination, Development and Fructification of the Higher Cryptogamia", Ray Society, London, 1862. The original German treatise appeared in 1851.) He showed that "the embryo-sac of the Coniferae may be looked upon as a spore remaining enclosed in its sporangium; the prothallium which it forms does not come to the light." (Ibid. page 438.) He thus determined the homologies on the female side. Recognising, as some previous observers had already done, that the microspores of those Cryptogams in which two kinds of spore are developed, are equivalent to the pollen-grains of the higher plants, he further pointed out that fertilisation "in the Rhizocarpeae and Selaginellae takes place by free spermatozoa, and in the Coniferae by a pollen-tube, in the interior of which spermatozoa are probably formed"--a remarkable instance of prescience, for though spermatozoids have not been found in the Conifers proper, they were demonstrated in the allied groups Cycadaceae and Ginkgo, in 1896, by the Japanese botanists Ikeno and Hirase. A new link was thus established between the Gymnosperms and the Cryptogams.

It remained uncertain, however, from which line of Cryptogams the gymnospermous Seed-plants had sprung. The great point of morphological comparison was the presence of two kinds of spore, and this was known to occur in the recent Lycopods and Water-ferns (Rhizocarpeae) and was also found in fossil representatives of the third phylum, that of the Horsetails. As a matter of fact all the three great Cryptogamic classes have found champions to maintain their claim to the ancestry of the Seed-plants, and in every case fossil evidence was called in. For a long time the Lycopods were the favourites, while the Ferns found the least support. The writer remembers, however, in the year 1881, hearing the late Prof. Sachs maintain, in a lecture to his class, that the descent of the Cycads could be traced, not merely from Ferns, but from a definite family of Ferns, the Marattiaceae, a view which, though in a somewhat crude form, anticipated more modern ideas.

Williamson appears to have been the first to recognise the presence, in the Carboniferous flora, of plants combining the characters of Ferns and Cycads. (See especially his "Organisation of the Fossil Plants of the Coal-Measures", Part XIII. "Phil. Trans. Royal Soc." 1887 B. page 299.) This conclusion was first reached in the case of the genera Heterangium and Lyginodendron, plants, which with a wholly fern-like habit, were found to unite an anatomical structure holding the balance between that of Ferns and Cycads, Heterangium inclining more to the former and Lyginodendron to the latter. Later researches placed Williamson's original suggestion on a firmer basis, and clearly proved the intermediate nature of these genera, and of a number of others, so far as their vegetative organs were concerned. This stage in our knowledge was marked by the institution of the class Cycadofilices by Potonie in 1897.

Nothing, however, was known of the organs of reproduction of the Cycadofilices, until F.W. Oliver, in 1903, identified a fossil seed, Lagenostoma Lomaxi, as belonging to Lyginodendron, the identification depending, in the first instance, on the recognition of an identical form of gland, of very characteristic structure, on the vegetative organs of Lyginodendron and on the cupule enveloping the seed. This evidence was supported by the discovery of a close anatomical agreement in other respects, as well as by constant association between the seed and the plant. (F.W. Oliver and D.H. Scott, "On the Structure of the Palaeozoic Seed, Lagenostoma Lomaxi, etc." "Phil. Trans. Royal Soc." Vol. 197 B. 1904.) The structure of the seed of Lyginodendron, proved to be of the same general type as that of the Cycads, as shown especially by the presence of a pollen-chamber or special cavity for the reception of the pollen-grains, an organ only known in the Cycads and Ginkgo among recent plants.

Within a few months after the discovery of the seed of Lyginodendron, Kidston found the large, nut-like seed of a Neuropteris, another fern-like Carboniferous plant, in actual connection with the pinnules of the frond, and since then seeds have been observed on the frond in species of Aneimites and Pecopteris, and a vast body of evidence, direct or indirect, has accumulated, showing that a large proportion of the Palaeozoic plants formerly classed as Ferns were in reality reproduced by seeds of the same type as those of recent Cycadaceae. (A summary of the evidence will be found in the writer's article "On the present position of Palaeozoic Botany", "Progressus Rei Botanicae", 1907, page 139, and "Studies in Fossil Botany", Vol. II. (2nd edition) London, 1909.) At the same time, the anatomical structure, where it is open to investigation, confirms the suggestion given by the habit, and shows that these early seed-bearing plants had a real affinity with Ferns. This conclusion received strong corroboration when Kidston, in 1905, discovered the male organs of Lyginodendron, and showed that they were identical with a fructification of the genus Crossotheca, hitherto regarded as belonging to Marattiaceous Ferns. (Kidston, "On the Microsporangia of the Pteridospermeae, etc." "Phil. Trans. Royal Soc." Vol. 198, B. 1906.)

The general conclusion which follows from the various observations alluded to, is that in Palaeozoic times there was a great body of plants (including, as it appears, a large majority of the fossils previously regarded as Ferns) which had attained the rank of Spermophyta, bearing seeds of a Cycadean type on fronds scarcely differing from the vegetative foliage, and in other respects, namely anatomy, habit and the structure of the pollen-bearing organs, retaining many of the characters of Ferns. From this extensive class of plants, to which the name Pteridospermeae has been given, it can scarcely be doubted that the abundant Cycadophyta, of the succeeding Mesozoic period, were derived. This conclusion is of far-reaching significance, for we have already found reason to think that the Angiosperms themselves sprang, in later times, from the Cycadophytic stock; it thus appears that the Fern-phylum, taken in a broad sense, ultimately represents the source from which the main line of descent of the Phanerogams took its rise.

It must further be borne in mind that in the Palaeozoic period there existed another group of seed-bearing plants, the Cordaiteae, far more advanced than the Pteridospermeae, and in many respects approaching the Coniferae, which themselves begin to appear in the latest Palaeozoic rocks. The Cordaiteae, while wholly different in habit from the contemporary fern-like Seed-plants, show unmistakable signs of a common origin with them. Not only is there a whole series of forms connecting the anatomical structure of the Cordaiteae with that of the Lyginodendreae among Pteridosperms, but a still more important point is that the seeds of the Cordaiteae, which have long been known, are of the same Cycadean type as those of the Pteridosperms, so that it is not always possible, as yet, to discriminate between the seeds of the two groups. These facts indicate that the same fern-like stock which gave rise to the Cycadophyta and through them, as appears probable, to the Angiosperms, was also the source of the Cordaiteae, which in their turn show manifest affinity with some at least of the Coniferae. Unless the latter are an artificial group, a view which does not commend itself to the writer, it would appear probable that the Gymnosperms generally, as well as the Angiosperms, were derived from an ancient race of Cryptogams, most nearly related to the Ferns. (Some botanists, however, believe that the Coniferae, or some of them, are probably more nearly related to the Lycopods. See Seward and Ford, "The Araucarieae, Recent and Extinct", "Phil. Trans. Royal Soc." Vol. 198 B. 1906.)

It may be mentioned here that the small gymnospermous group Gnetales (including the extraordinary West African plant Welwitschia) which were formerly regarded by some authorities as akin to the Equisetales, have recently been referred, on better grounds, to a common origin with the Angiosperms, from the Mesozoic Cycadophyta.

The tendency, therefore, of modern work on the palaeontological record of the Seed-plants has been to exalt the importance of the Fern-phylum, which, on present evidence, appears to be that from which the great majority, possibly the whole, of the Spermophyta have been derived.

One word of caution, however, is necessary. The Seed-plants are of enormous antiquity; both the Pteridosperms and the more highly organised family Cordaiteae, go back as far in geological history (namely to the Devonian) as the Ferns themselves or any other Vascular Cryptogams. It must therefore be understood that in speaking of the derivation of the Spermophyta from the Fern-phylum, we refer to that phylum at a very early stage, probably earlier than the most ancient period to which our record of land-plants extends. The affinity between the oldest Seed-plants and the Ferns, in the widest sense, seems established, but the common stock from which they actually arose is still unknown; though no doubt nearer to the Ferns than to any other group, it must have differed widely from the Ferns as we now know them, or perhaps even from any which the fossil record has yet revealed to us.

iii. THE ORIGIN OF THE HIGHER CRYPTOGAMIA.

The Sub-kingdom of the higher Spore-plants, the Cryptogamia possessing a vascular system, was more prominent in early geological periods than at present. It is true that the dominance of the Pteridophyta in Palaeozoic times has been much exaggerated owing to the assumption that everything which looked like a Fern really was a Fern. But, allowing for the fact, now established, that most of the Palaeozoic fern-like plants were already Spermophyta, there remains a vast mass of Cryptogamic forms of that period, and the familiar statement that they formed the main constituent of the Coal-forests still holds good. The three classes, Ferns (Filicales), Horsetails (Equisetales) and Club-mosses (Lycopodiales), under which we now group the Vascular Cryptogams, all extend back in geological history as far as we have any record of the flora of the land; in the Palaeozoic, however, a fourth class, the Sphenophyllales, was present.

As regards the early history of the Ferns, which are of special interest from their relation to the Seed-plants, it is impossible to speak quite positively, owing to the difficulty of discriminating between true fossil Ferns and the Pteridosperms which so closely simulated them. The difficulty especially affects the question of the position of Marattiaceous Ferns in the Palaeozoic Floras. This family, now so restricted, was until recently believed to have been one of the most important groups of Palaeozoic plants, especially during later Carboniferous and Permian times. Evidence both from anatomy and from sporangial characters appeared to establish this conclusion. Of late, however, doubts have arisen, owing to the discovery that some supposed members of the Marattiaceae bore seeds, and that a form of fructification previously referred to that family (Crossotheca) was really the pollen-bearing apparatus of a Pteridosperm (Lyginodendron). The question presents much difficulty; though it seems certain that our ideas of the extent of the family in Palaeozoic times will have to be restricted, there is still a decided balance of evidence in favour of the view that a considerable body of Marattiaceous Ferns actually existed. The plants in question were of large size (often arborescent) and highly organised--they represent, in fact, one of the highest developments of the Fern-stock, rather than a primitive type of the class.

There was, however, in the Palaeozoic period, a considerable group of comparatively simple Ferns (for which Arber has proposed the collective name Primofilices); the best known of these are referred to the family Botryopterideae, consisting of plants of small or moderate dimensions, with, on the whole, a simple anatomical structure, in certain cases actually simpler than that of any recent Ferns. On the other hand the sporangia of these plants were usually borne on special fertile fronds, a mark of rather high differentiation. This group goes back to the Devonian and includes some of the earliest types of Fern with which we are acquainted. It is probable that the Primofilices (though not the particular family Botryopterideae) represent the stock from which the various families of modern Ferns, already developed in the Mesozoic period, may have sprung.

None of the early Ferns show any clear approach to other classes of Vascular Cryptogams; so far as the fossil record affords any evidence, Ferns have always been plants with relatively large and usually compound leaves. There is no indication of their derivation from a microphyllous ancestry, though, as we shall see, there is some slight evidence for the converse hypothesis. Whatever the origin of the Ferns may have been it is hidden in the older rocks.

It has, however, been held that certain other Cryptogamic phyla had a common origin with the Ferns. The Equisetales are at present a well-defined group; even in the rich Palaeozoic floras the habit, anatomy and reproductive characters usually render the members of this class unmistakable, in spite of the great development and stature which they then attained. It is interesting, however, to find that in the oldest known representatives of the Equisetales the leaves were highly developed and dichotomously divided, thus differing greatly from the mere scale-leaves of the recent Horsetails, or even from the simple linear leaves of the later Calamites. The early members of the class, in their forked leaves, and in anatomical characters, show an approximation to the Sphenophyllales, which are chiefly represented by the large genus Sphenophyllum, ranging through the Palaeozoic from the Middle Devonian onwards. These were plants with rather slender, ribbed stems, bearing whorls of wedge-shaped or deeply forked leaves, six being the typical number in each whorl. From their weak habit it has been conjectured, with much probability, that they may have been climbing plants, like the scrambling Bedstraws of our hedgerows. The anatomy of the stem is simple and root-like; the cones are remarkable for the fact that each scale or sporophyll is a double structure, consisting of a lower, usually sterile lobe and one or more upper lobes bearing the sporangia; in one species both parts of the sporophyll were fertile. Sphenophyllum was evidently much specialised; the only other known genus is based on an isolated cone, Cheirostrobus, of Lower Carboniferous age, with an extraordinarily complex structure. In this genus especially, but also in the entire group, there is an evident relation to the Equisetales; hence it is of great interest that Nathorst has described, from the Devonian of Bear Island in the Arctic regions, a new genus Pseudobornia, consisting of large plants, remarkable for their highly compound leaves which, when found detached, were taken for the fronds of a Fern. The whorled arrangement of the leaves, and the habit of the plant, suggest affinities either with the Equisetales or the Sphenophyllales; Nathorst makes the genus the type of a new class, the Pseudoborniales. (A.G. Nathorst, "Zur Oberdevonischen Flora der Baren-Insel", "Kongl. Svenska Vetenskaps-Akademiens Handlingar" Bd. 36, No. 3, Stockholm, 1902.)

The available data, though still very fragmentary, certainly suggest that both Equisetales and Sphenophyllales may have sprung from a common stock having certain fern-like characters. On the other hand the Sphenophylls, and especially the peculiar genus Cheirostrobus, have in their anatomy a good deal in common with the Lycopods, and of late years they have been regarded as the derivatives of a stock common to that class and the Equisetales. At any rate the characters of the Sphenophyllales and of the new group Pseudoborniales suggest the existence, at a very early period, of a synthetic race of plants, combining the characters of various phyla of the Vascular Cryptogams. It may further be mentioned that the Psilotaceae, an isolated epiphytic family hitherto referred to the Lycopods, have been regarded by several recent authors as the last survivors of the Sphenophyllales, which they resemble both in their anatomy and in the position of their sporangia.

The Lycopods, so far as their early history is known, are remarkable rather for their high development in Palaeozoic times than for any indications of a more primitive ancestry. In the recent Flora, two of the four living genera (Excluding Psilotaceae.) (Selaginella and Isoetes) have spores of two kinds, while the other two (Lycopodium and Phylloglossum) are homosporous. Curiously enough, no certain instance of a homosporous Palaeozoic Lycopod has yet been discovered, though well-preserved fructifications are numerous. Wherever the facts have been definitely ascertained, we find two kinds of spore, differentiated quite as sharply as in any living members of the group. Some of the Palaeozoic Lycopods, in fact, went further, and produced bodies of the nature of seeds, some of which were actually regarded, for many years, as the seeds of Gymnosperms. This specially advanced form of fructification goes back at least as far as the Lower Carboniferous, while the oldest known genus of Lycopods, Bothrodendron, which is found in the Devonian, though not seed-bearing, was typically heterosporous, if we may judge from the Coal-measure species. No doubt homosporous Lycopods existed, but the great prevalence of the higher mode of reproduction in days which to us appear ancient, shows how long a course of evolution must have already been passed through before the oldest known members of the group came into being. The other characters of the Palaeozoic Lycopods tell the same tale; most of them attained the stature of trees, with a corresponding elaboration of anatomical structure, and even the herbaceous forms show no special simplicity. It appears from recent work that herbaceous Lycopods, indistinguishable from our recent Selaginellas, already existed in the time of the Coal-measures, while one herbaceous form (Miadesmia) is known to have borne seeds.

The utmost that can be said for primitiveness of character in Palaeozoic Lycopods is that the anatomy of the stem, in its primary ground-plan, as distinguished from its secondary growth, was simpler than that of most Lycopodiums and Selaginellas at the present day. There are also some peculiarities in the underground organs (Stigmaria) which suggest the possibility of a somewhat imperfect differentiation between root and stem, but precisely parallel difficulties are met with in the case of the living Selaginellas, and in some degree in species of Lycopodium.

In spite of their high development in past ages the Lycopods, recent and fossil, constitute, on the whole, a homogeneous group, and there is little at present to connect them with other phyla. Anatomically some relation to the Sphenophylls is indicated, and perhaps the recent Psilotaceae give some support to this connection, for while their nearest alliance appears to be with the Sphenophylls, they approach the Lycopods in anatomy, habit, and mode of branching.

The typically microphyllous character of the Lycopods, and the simple relation between sporangium and sporophyll which obtains throughout the class, have led various botanists to regard them as the most primitive phylum of the Vascular Cryptogams. There is nothing in the fossil record to disprove this view, but neither is there anything to support it, for this class so far as we know is no more ancient than the megaphyllous Cryptogams, and its earliest representatives show no special simplicity. If the indications of affinity with Sphenophylls are of any value the Lycopods are open to suspicion of reduction from a megaphyllous ancestry, but there is no direct palaeontological evidence for such a history.

The general conclusions to which we are led by a consideration of the fossil record of the Vascular Cryptogams are still very hypothetical, but may be provisionally stated as follows:

The Ferns go back to the earliest known period. In Mesozoic times practically all the existing families had appeared; in the Palaeozoic the class was less extensive than formerly believed, a majority of the supposed Ferns of that age having proved to be seed-bearing plants. The oldest authentic representatives of the Ferns were megaphyllous plants, broadly speaking, of the same type as those of later epochs, though differing much in detail. As far back as the record extends they show no sign of becoming merged with other phyla in any synthetic group.

The Equisetales likewise have a long history, and manifestly attained their greatest development in Palaeozoic times. Their oldest forms show an approach to the extinct class Sphenophyllales, which connects them to some extent, by anatomical characters, with the Lycopods. At the same time the oldest Equisetales show a somewhat megaphyllous character, which was more marked in the Devonian Pseudoborniales. Some remote affinity with the Ferns (which has also been upheld on other grounds) may thus be indicated. It is possible that in the Sphenophyllales we may have the much-modified representatives of a very ancient synthetic group.

The Lycopods likewise attained their maximum in the Palaeozoic, and show, on the whole, a greater elaboration of structure in their early forms than at any later period, while at the same time maintaining a considerable degree of uniformity in morphological characters throughout their history. The Sphenophyllales are the only other class with which they show any relation; if such a connection existed, the common point of origin must lie exceedingly far back.

The fossil record, as at present known, cannot, in the nature of things, throw any direct light on what is perhaps the most disputed question in the morphology of plants--the origin of the alternating generations of the higher Cryptogams and the Spermophyta. At the earliest period to which terrestrial plants have been traced back all the groups of Vascular Cryptogams were in a highly advanced stage of evolution, while innumerable Seed-plants--presumably the descendants of Cryptogamic ancestors--were already flourishing. On the other hand we know practically nothing of Palaeozoic Bryophyta, and the evidence even for their existence at that period cannot be termed conclusive. While there are thus no palaeontological grounds for the hypothesis that the Vascular plants came of a Bryophytic stock, the question of their actual origin remains unsolved.

III. NATURAL SELECTION.

Hitherto we have considered the palaeontological record of plants in relation to Evolution. The question remains, whether the record throws any light on the theory of which Darwin and Wallace were the authors--that of Natural Selection. The subject is clearly one which must be investigated by other methods than those of the palaeontologist; still there are certain important points involved, on which the palaeontological record appears to bear.

One of these points is the supposed distinction between morphological and adaptive characters, on which Nageli, in particular, laid so much stress. The question is a difficult one; it was discussed by Darwin ("Origin of Species" (6th edition), pages 170-176.), who, while showing that the apparent distinction is in part to be explained by our imperfect knowledge of function, recognised the existence of important morphological characters which are not adaptations. The following passage expresses his conclusion. "Thus, as I am inclined to believe, morphological differences, which we consider as important--such as the arrangement of the leaves, the divisions of the flower or of the ovarium, the position of the ovules, etc.--first appeared in many cases as fluctuating variations, which sooner or later became constant through the nature of the organism and of the surrounding conditions, as well as through the inter-crossing of distinct individuals, but not through natural selection; for as these morphological characters do not affect the welfare of the species, any slight deviations in them could not have been governed or accumulated through this latter agency." (Ibid. page 176.)

This is a sufficiently liberal concession; Nageli, however, went much further when he said: "I do not know among plants a morphological modification which can be explained on utilitarian principles." (See "More Letters", Vol. II. page 375 (footnote).) If this were true the field of Natural Selection would be so seriously restricted, as to leave the theory only a very limited importance.

It can be shown, as the writer believes, that many typical "morphological characters," on which the distinction between great classes of plants is based, were adaptive in origin, and even that their constancy is due to their functional importance. Only one or two cases will be mentioned, where the fossil evidence affects the question.

The pollen-tube is one of the most important morphological characters of the Spermophyta as now existing--in fact the name Siphonogama is used by Engler in his classification, as expressing a peculiarly constant character of the Seed-plants. Yet the pollen-tube is a manifest adaptation, following on the adoption of the seed-habit, and serving first to bring the spermatozoids with greater precision to their goal, and ultimately to relieve them of the necessity for independent movement. The pollen-tube is constant because it has proved to be indispensable.

In the Palaeozoic Seed-plants there are a number of instances in which the pollen-grains, contained in the pollen-chamber of a seed, are so beautifully preserved that the presence of a group of cells within the grain can be demonstrated; sometimes we can even see how the cell-walls broke down to emit the sperms, and quite lately it is said that the sperms themselves have been recognised. (F.W. Oliver, "On Physostoma elegans, an archaic type of seed from the Palaeozoic Rocks", "Annals of Botany", January, 1909. See also the earlier papers there cited.) In no case, however, is there as yet any satisfactory evidence for the formation of a pollen-tube; it is probable that in these early Seed-plants the pollen-grains remained at about the evolutionary level of the microspores in Pilularia or Selaginella, and discharged their spermatozoids directly, leaving them to find their own way to the female cells. It thus appears that there were once Spermophyta without pollen-tubes. The pollen-tube method ultimately prevailed, becoming a constant "morphological character," for no other reason than because, under the new conditions, it provided a more perfect mechanism for the accomplishment of the act of fertilisation. We have still, in the Cycads and Ginkgo, the transitional case, where the tube remains short, serves mainly as an anchor and water-reservoir, but yet is able, by its slight growth, to give the spermatozoids a "lift" in the right direction. In other Seed-plants the sperms are mere passengers, carried all the way by the pollen-tube; this fact has alone rendered the Angiospermous method of fertilisation through a stigma possible.

We may next take the seed itself--the very type of a morphological character. Our fossil record does not go far enough back to tell us the origin of the seed in the Cycadophyta and Pteridosperms (the main line of its development) but some interesting sidelights may be obtained from the Lycopod phylum. In two Palaeozoic genera, as we have seen, seed-like organs are known to have been developed, resembling true seeds in the presence of an integument and of a single functional embryo-sac, as well as in some other points. We will call these organs "seeds" for the sake of shortness. In one genus (Lepidocarpon) the seeds were borne on a cone indistinguishable from that of the ordinary cryptogamic Lepidodendreae, the typical Lycopods of the period, while the seed itself retained much of the detailed structure of the sporangium of that family. In the second genus, Miadesmia, the seed-bearing plant was herbaceous, and much like a recent Selaginella. (See Margaret Benson, "Miadesmia membranacea, a new Palaeozoic Lycopod with a seed-like structure", "Phil. Trans. Royal Soc. Vol." 199, B. 1908.) The seeds of the two genera are differently constructed, and evidently had an independent origin. Here, then, we have seeds arising casually, as it were, at different points among plants which otherwise retain all the characters of their cryptogamic fellows; the seed is not yet a morphological character of importance. To suppose that in these isolated cases the seed sprang into being in obedience to a Law of Advance ("Vervollkommungsprincip"), from which other contemporary Lycopods were exempt, involves us in unnecessary mysticism. On the other hand it is not difficult to see how these seeds may have arisen, as adaptive structures, under the influence of Natural Selection. The seed-like structure afforded protection to the prothallus, and may have enabled the embryo to be launched on the world in greater security. There was further, as we may suppose, a gain in certainty of fertilisation. As the writer has pointed out elsewhere, the chances against the necessary association of the small male with the large female spores must have been enormously great when the cones were borne high up on tall trees. The same difficulty may have existed in the case of the herbaceous Miadesmia, if, as Miss Benson conjectures, it was an epiphyte. One way of solving the problem was for pollination to take place while the megaspore was still on the parent plant, and this is just what the formation of an ovule or seed was likely to secure.

The seeds of the Pteridosperms, unlike those of the Lycopod stock, have not yet been found in statu nascendi--in all known cases they were already highly developed organs and far removed from the cryptogamic sporangium. But in two respects we find that these seeds, or some of them, had not yet realised their possibilities. In the seed of Lyginodendron and other cases the micropyle, or orifice of the integument, was not the passage through which the pollen entered; the open neck of the pollen-chamber protruded through the micropyle and itself received the pollen. We have met with an analogous case, at a more advanced stage of evolution, in the Bennettiteae, where the wall of the gynaecium, though otherwise closed, did not provide a stigma to catch the pollen, but allowed the micropyles of the ovules to protrude and receive the pollen in the old gymnospermous fashion. The integument in the one case and the pistil in the other had not yet assumed all the functions to which the organ ultimately became adapted. Again, no Palaeozoic seed has yet been found to contain an embryo, though the preservation is often good enough for it to have been recognised if present. It is probable that the nursing of the embryo had not yet come to be one of the functions of the seed, and that the whole embryonic development was relegated to the germination stage.

In these two points, the reception of the pollen by the micropyle and the nursing of the embryo, it appears that many Palaeozoic seeds were imperfect, as compared with the typical seeds of later times. As evolution went on, one function was superadded on another, and it appears impossible to resist the conclusion that the whole differentiation of the seed was a process of adaptation, and consequently governed by Natural Selection, just as much as the specialisation of the rostellum in an Orchid, or of the pappus in a Composite.

Did space allow, other examples might be added. We may venture to maintain that the glimpses which the fossil record allows us into early stages in the evolution of organs now of high systematic importance, by no means justify the belief in any essential distinction between morphological and adaptive characters.

Another point, closely connected with Darwin's theory, on which the fossil history of plants has been supposed to have some bearing, is the question of Mutation, as opposed to indefinite variation. Arber and Parkin, in their interesting memoir on the Origin of Angiosperms, have suggested calling in Mutation to explain the apparently sudden transition from the cycadean to the angiospermous type of foliage, in late Mesozoic times, though they express themselves with much caution, and point out "a distinct danger that Mutation may become the last resort of the phylogenetically destitute"!

The distinguished French palaeobotanists, Grand'Eury (C. Grand'Eury, "Sur les mutations de quelques Plantes fossiles du Terrain houiller". "Comptes Rendus", CXLII. page 25, 1906.) and Zeiller (R. Zeiller "Les Vegetaux fossiles et leurs Enchainements", "Revue du Mois", III. February, 1907.), are of opinion, to quote the words of the latter writer, that the facts of fossil Botany are in agreement with the sudden appearance of new forms, differing by marked characters from those that have given them birth; he adds that these results give more amplitude to this idea of Mutation, extending it to groups of a higher order, and even revealing the existence of discontinuous series between the successive terms of which we yet recognise bonds of filiation. (Loc. cit. page 23.)

If Zeiller's opinion should be confirmed, it would no doubt be a serious blow to the Darwinian theory. As Darwin said: "Under a scientific point of view, and as leading to further investigation, but little advantage is gained by believing that new forms are suddenly developed in an inexplicable manner from old and widely different forms, over the old belief in the creation of species from the dust of the earth." ("Origin of Species", page 424.)

It most however be pointed out, that such mutations as Zeiller, and to some extent Arber and Parkin, appear to have in view, bridging the gulf between different Orders and Classes, bear no relation to any mutations which have been actually observed, such as the comparatively small changes, of sub-specific value, described by De Vries in the type-case of Oenothera Lamarckiana. The results of palaeobotanical research have undoubtedly tended to fill up gaps in the Natural System of plants--that many such gaps still persist is not surprising; their presence may well serve as an incentive to further research but does not, as it seems to the writer, justify the assumption of changes in the past, wholly without analogy among living organisms.

As regards the succession of species, there are no greater authorities than Grand'Eury and Zeiller, and great weight must be attached to their opinion that the evidence from continuous deposits favours a somewhat sudden change from one specific form to another. At the same time it will be well to bear in mind that the subject of the "absence of numerous intermediate varieties in any single formation" was fully discussed by Darwin. ("Origin of Species", pages 275-282, and page 312.); the explanation which he gave may go a long way to account for the facts which recent writers have regarded as favouring the theory of saltatory mutation.

The rapid sketch given in the present essay can do no more than call attention to a few salient points, in which the palaeontological records of plants has an evident bearing on the Darwinian theory. At the present day the whole subject of palaeobotany is a study in evolution, and derives its chief inspiration from the ideas of Darwin and Wallace. In return it contributes something to the verification of their teaching; the recent progress of the subject, in spite of the immense difficulties which still remain, has added fresh force to Darwin's statement that "the great leading facts in palaeontology agree admirably with the theory of descent with modification through variation and natural selection." (Ibid. page 313.)

XIII. THE INFLUENCE OF ENVIRONMENT ON THE FORMS OF PLANTS. By Georg Klebs, PH.D.

Professor of Botany in the University of Heidelberg.

The dependence of plants on their environment became the object of scientific research when the phenomena of life were first investigated and physiology took its place as a special branch of science. This occurred in the course of the eighteenth century as the result of the pioneer work of Hales, Duhamel, Ingenhousz, Senebier and others. In the nineteenth century, particularly in the second half, physiology experienced an unprecedented development in that it began to concern itself with the experimental study of nutrition and growth, and with the phenomena associated with stimulus and movement; on the other hand, physiology neglected phenomena connected with the production of form, a department of knowledge which was the province of morphology, a purely descriptive science. It was in the middle of the last century that the growth of comparative morphology and the study of phases of development reached their highest point.

The forms of plants appeared to be the expression of their inscrutable inner nature; the stages passed through in the development of the individual were regarded as the outcome of purely internal and hidden laws. The feasibility of experimental inquiry seemed therefore remote. Meanwhile, the recognition of the great importance of such a causal morphology emerged from the researches of the physiologists of that time, more especially from those of Hofmeister (Hofmeister, "Allgemeine Morphologie", Leipzig, 1868, page 579.), and afterwards from the work of Sachs. (Sachs, "Stoff und Form der Pflanzenorgane", Vol. I. 1880; Vol. II. 1882. "Gesammelte Abhandlungen uber Pflanzen-Physiologie", II. Leipzig, 1893.) Hofmeister, in speaking of this line of inquiry, described it as "the most pressing and immediate aim of the investigator to discover to what extent external forces acting on the organism are of importance in determining its form." This advance was the outcome of the influence of that potent force in biology which was created by Darwin's "Origin of Species" (1859).

The significance of the splendid conception of the transformation of species was first recognised and discussed by Lamarck (1809); as an explanation of transformation he at once seized upon the idea--an intelligible view--that the external world is the determining factor. Lamarck (Lamarck, "Philosophie zoologique", pages 223-227. Paris, 1809.) endeavoured, more especially, to demonstrate from the behaviour of plants that changes in environment induce change in form which eventually leads to the production of new species. In the case of animals, Lamarck adopted the teleological view that alterations in the environment first lead to alterations in the needs of the organisms, which, as the result of a kind of conscious effort of will, induce useful modifications and even the development of new organs. His work has not exercised any influence on the progress of science: Darwin himself confessed in regard to Lamarck's work--"I got not a fact or idea from it." ("Life and Letters", Vol. II. page 215.)

On a mass of incomparably richer and more essential data Darwin based his view of the descent of organisms and gained for it general acceptance; as an explanation of modification he elaborated the ingeniously conceived selection theory. The question of special interest in this connection, namely what is the importance of the influence of the environment, Darwin always answered with some hesitation and caution, indeed with a certain amount of indecision.

The fundamental principle underlying his theory is that of general variability as a whole, the nature and extent of which, especially in cultivated organisms, are fully dealt with in his well-known book. (Darwin, "The variation of Animals and Plants under domestication", 2 vols., edition 1, 1868; edition 2, 1875; popular edition 1905.) In regard to the question as to the cause of variability Darwin adopts a consistently mechanical view. He says: "These several considerations alone render it probable that variability of every kind is directly or indirectly caused by changed conditions of life. Or, to put the case under another point of view, if it were possible to expose all the individuals of a species during many generations to absolutely uniform conditions of life, there would be no variability." ("The variation of Animals and Plants" (2nd edition), Vol. II. page 242.) Darwin did not draw further conclusions from this general principle.

Variations produced in organisms by the environment are distinguished by Darwin as "the definite" and "the indefinite." (Ibid. II. page 260. See also "Origin of Species" (6th edition), page 6.) The first occur "when all or nearly all the offspring of an individual exposed to certain conditions during several generations are modified in the same manner." Indefinite variation is much more general and a more important factor in the production of new species; as a result of this, single individuals are distinguished from one another by "slight" differences, first in one then in another character. There may also occur, though this is very rare, more marked modifications, "variations which seem to us in our ignorance to arise spontaneously." ("Origin of Species" (6th edition), page 421.) The selection theory demands the further postulate that such changes, "whether extremely slight or strongly marked," are inherited. Darwin was no nearer to an experimental proof of this assumption than to the discovery of the actual cause of variability. It was not until the later years of his life that Darwin was occupied with the "perplexing problem... what causes almost every cultivated plant to vary" ("Life and Letters", Vol. III. page 342.): he began to make experiments on the influence of the soil, but these were soon given up.

In the course of the violent controversy which was the outcome of Darwin's work the fundamental principles of his teaching were not advanced by any decisive observations. Among the supporters and opponents, Nageli (Nageli, "Theorie der Abstammungslehre", Munich, 1884; cf. Chapter III.) was one of the few who sought to obtain proofs by experimental methods. His extensive cultural experiments with alpine Hieracia led him to form the opinion that the changes which are induced by an alteration in the food-supply, in climate or in habitat, are not inherited and are therefore of no importance from the point of view of the production of species. And yet Nageli did attribute an important influence to the external world; he believed that adaptations of plants arise as reactions to continuous stimuli, which supply a need and are therefore useful. These opinions, which recall the teleological aspect of Lamarckism, are entirely unsupported by proof. While other far-reaching attempts at an explanation of the theory of descent were formulated both in Nageli's time and afterwards, some in support of, others in opposition to Darwin, the necessity of investigating, from different standpoints, the underlying causes, variability and heredity, was more and more realised. To this category belong the statistical investigations undertaken by Quetelet and Galton, the researches into hybridisation, to which an impetus was given by the re-discovery of the Mendelian law of segregation, as also by the culture experiments on mutating species following the work of de Vries, and lastly the consideration of the question how far variation and heredity are governed by external influences. These latter problems, which are concerned in general with the causes of form-production and form-modification, may be treated in a short summary which falls under two heads, one having reference to the conditions of form-production in single species, the other being concerned with the conditions governing the transformation of species.

I. THE INFLUENCE OF EXTERNAL CONDITIONS ON FORM-PRODUCTION IN SINGLE SPECIES.

The members of plants, which we express by the terms stem, leaf, flower, etc. are capable of modification within certain limits; since Lamarck's time this power of modification has been brought more or less into relation with the environment. We are concerned not only with the question of experimental demonstration of this relationship, but, more generally, with an examination of the origin of forms, the sequences of stages in development that are governed by recognisable causes. We have to consider the general problem; to study the conditions of all typical as well as of atypic forms, in other words, to found a physiology of form.

If we survey the endless variety of plant-forms and consider the highly complex and still little known processes in the interior of cells, and if we remember that the whole of this branch of investigation came into existence only a few decades ago, we are able to grasp the fact that a satisfactory explanation of the factors determining form cannot be discovered all at once. The goal is still far away. We are not concerned now with the controversial question, whether, on the whole, the fundamental processes in the development of form can be recognised by physiological means. A belief in the possibility of this can in any case do no harm. What we may and must attempt is this--to discover points of attack on one side or another, which may enable us by means of experimental methods to come into closer touch with these elusive and difficult problems. While we are forced to admit that there is at present much that is insoluble there remains an inexhaustible supply of problems capable of solution.

The object of our investigations is the species; but as regards the question, what is a species, science of to-day takes up a position different from that of Darwin. For him it was the Linnean species which illustrates variation: we now know, thanks to the work of Jordan, de Bary, and particularly to that of de Vries (de Vries, "Die Mutationstheorie", Leipzig, 1901, Vol. I. page 33.), that the Linnean species consists of a large or small number of entities, elementary species. In experimental investigation it is essential that observations be made on a pure species, or, as Johannsen (Johannsen, "Ueber Erblichkeit in Populationen und reinen Linien", Jena, 1903.) says, on a pure "line." What has long been recognised as necessary in the investigation of fungi, bacteria and algae must also be insisted on in the case of flowering plants; we must start with a single individual which is reproduced vegetatively or by strict self-fertilisation. In dioecious plants we must aim at the reproduction of brothers and sisters.

We may at the outset take it for granted that a pure species remains the same under similar external conditions; it varies as these vary. IT IS CHARACTERISTIC OF A SPECIES THAT IT ALWAYS EXHIBITS A CONSTANT RELATION TO A PARTICULAR ENVIRONMENT. In the case of two different species, e.g. the hay and anthrax bacilli or two varieties of Campanula with blue and white flowers respectively, a similar environment produces a constant difference. The cause of this is a mystery.

According to the modern standpoint, the living cell is a complex chemico-physical system which is regarded as a dynamical system of equilibrium, a conception suggested by Herbert Spencer and which has acquired a constantly increasing importance in the light of modern developments in physical chemistry. The various chemical compounds, proteids, carbohydrates, fats, the whole series of different ferments, etc. occur in the cell in a definite physical arrangement. The two systems of two species must as a matter of fact possess a constant difference, which it is necessary to define by a special term. We say, therefore, that the SPECIFIC STRUCTURE is different.

By way of illustrating this provisionally, we may assume that the proteids of the two species possess a constant chemical difference. This conception of specific structure is specially important in its bearing on a further treatment of the subject. In the original cell, eventually also in every cell of a plant, the characters which afterwards become apparent must exist somewhere; they are integral parts of the capabilities or potentialities of specific structure. Thus not only the characters which are exhibited under ordinary conditions in nature, but also many others which become apparent only under special conditions (In this connection I leave out of account, as before, the idea of material carriers of heredity which since the publication of Darwin's Pangenesis hypothesis has been frequently suggested. See my remarks in "Variationen der Bluten", "Pringsheim's Jahrb. Wiss. Bot." 1905, page 298; also Detto, "Biol. Centralbl." 1907, page 81, "Die Erklarbarkeit der Ontogenese durch materielle Anlagen".), are to be included as such potentialities in cells; the conception of specific structure includes the WHOLE OF THE POTENTIALITIES OF A SPECIES; specific structure comprises that which we must always assume without being able to explain it.

A relatively simple substance, such as oxalate of lime, is known under a great number of different crystalline forms belonging to different systems (Compare Kohl's work on "Anatomisch-phys. Untersuchungen uber Kalksalze", etc. Marburg, 1889.); these may occur as single crystals, concretions or as concentric sphaerites. The power to assume this variety of form is in some way inherent in the molecular structure, though we cannot, even in this case, explain the necessary connection between structure and crystalline form. These potentialities can only become operative under the influence of external conditions; their stimulation into activity depends on the degree of concentration of the various solutions, on the nature of the particular calcium salt, on the acid or alkaline reactions. Broadly speaking, the plant cell behaves in a similar way. The manifestation of each form, which is inherent as a potentiality in the specific structure, is ultimately to be referred to external conditions.

An insight into this connection is, however, rendered exceedingly difficult, often quite impossible, because the environment never directly calls into action the potentialities. Its influence is exerted on what we may call the inner world of the organism, the importance of which increases with the degree of differentiation. The production of form in every plant depends upon processes in the interior of the cells, and the nature of these determines which among the possible characters is to be brought to light. In no single case are we acquainted with the internal process responsible for the production of a particular form. All possible factors may play a part, such as osmotic pressure, permeability of the protoplasm, the degree of concentration of the various chemical substances, etc.; all these factors should be included in the category of INTERNAL CONDITIONS. This inner world appears the more hidden from our ken because it is always represented by a certain definite state, whether we are dealing with a single cell or with a small group of cells. These have been produced from pre-existing cells and they in turn from others; the problem is constantly pushed back through a succession of generations until it becomes identified with that of the origin of species.

A way, however, is opened for investigation; experience teaches us that this inner world is not a constant factor: on the contrary, it appears to be very variable. The dependence of VARIABLE INTERNAL on VARIABLE EXTERNAL conditions gives us the key with which research may open the door. In the lower plants this dependence is at once apparent, each cell is directly subject to external influences. In the higher plants with their different organs, these influences were transmitted to cells in course of development along exceedingly complex lines. In the case of the growing-point of a bud, which is capable of producing a complete plant, direct influences play a much less important part than those exerted through other organs, particularly through the roots and leaves, which are essential in nutrition. These correlations, as we may call them, are of the greatest importance as aids to an understanding of form-production. When a bud is produced on a particular part of a plant, it undergoes definite internal modifications induced by the influence of other organs, the activity of which is governed by the environment, and as the result of this it develops along a certain direction; it may, for example, become a flower. The particular direction of development is determined before the rudiment is differentiated and is exerted so strongly that further development ensues without interruption, even though the external conditions vary considerably and exert a positively inimical influence: this produces the impression that development proceeds entirely independently of the outer world. The widespread belief that such independence exists is very premature and at all events unproven.

The state of the young rudiment is the outcome of previous influences of the external world communicated through other organs. Experiments show that in certain cases, if the efficiency of roots and leaves as organs concerned with nutrition is interfered with, the production of flowers is affected, and their characters, which are normally very constant, undergo far-reaching modifications. To find the right moment at which to make the necessary alteration in the environment is indeed difficult and in many cases not yet possible. This is especially the case with fertilised eggs, which in a higher degree than buds have acquired, through parental influences, an apparently fixed internal organisation, and this seems to have pre-determined their development. It is, however, highly probable that it will be possible, by influencing the parents, to alter the internal organisation and to switch off development on to other lines.

Having made these general observations I will now cite a few of the many facts at our disposal, in order to illustrate the methods and aim of the experimental methods of research. As a matter of convenience I will deal separately with modification of development and with modification of single organs.

I. EFFECT OF ENVIRONMENT UPON THE COURSE OF DEVELOPMENT.

Every plant, whether an alga or a flowering plant passes, under natural conditions, through a series of developmental stages characteristic of each species, and these consist in a regular sequence of definite forms. It is impossible to form an opinion from mere observation and description as to what inner changes are essential for the production of the several forms. We must endeavour to influence the inner factors by known external conditions in such a way that the individual stages in development are separately controlled and the order of their sequence determined at will by experimental treatment. Such control over the course of development may be gained with special certainty in the case of the lower organisms.

With these it is practicable to control the principal conditions of cultivation and to vary them in various ways. By this means it has been demonstrated that each developmental stage depends upon special external conditions, and in cases where our knowledge is sufficient, a particular stage may be obtained at will. In the Green Algae (See Klebs, "Die Bedingung der Fortpflanzung... ", Jena, 1896; also "Jahrb. fur Wiss. Bot." 1898 and 1900; "Probleme der Entwickelung, III." "Biol. Centralbl." 1904, page 452.), as in the case of Fungi, we may classify the stages of development into purely vegetative growth (growth, cell-division, branching), asexual reproduction (formation of zoospores, conidia) and sexual processes (formation of male and female sexual organs). By modifying the external conditions it is possible to induce algae or fungi (Vaucheria, Saprolegnia) to grow continuously for several years or, in the course of a few days, to die after an enormous production of asexual or sexual cells. In some instances even an almost complete stoppage of growth may be caused, reproductive cells being scarcely formed before the organism is again compelled to resort to reproduction. Thus the sequence of the different stages in development can be modified as we may desire.

The result of a more thorough investigation of the determining conditions appears to produce at first sight a confused impression of all sorts of possibilities. Even closely allied species exhibit differences in regard to the connection between their development and external conditions. It is especially noteworthy that the same form in development may be produced as the result of very different alterations in the environment. At the same time we can undoubtedly detect a certain unity in the multiplicity of the individual phenomena.

If we compare the factors essential for the different stages in development, we see that the question always resolves itself into one of modification of similar conditions common to all life-processes. We should rather have inferred that there exist specific external stimuli for each developmental stage, for instance, certain chemical agencies. Experiments hitherto made support the conclusion that QUANTITATIVE alterations in the general conditions of life produce different types of development. An alga or a fungus grows so long as all the conditions of nutrition remain at a certain optimum for growth. In order to bring about asexual reproduction, e.g. the formation of zoospores, it is sometimes necessary to increase the degree of intensity of external factors; sometimes, on the other hand, these must be reduced in intensity. In the case of many algae a decrease in light-intensity or in the amount of salts in the culture solution, or in the temperature, induces asexual reproduction, while in others, on the contrary, an increase in regard to each of these factors is required to produce the same result. This holds good for the quantitative variations which induce sexual reproduction in algae. The controlling factor is found to be a reduction in the supply of nutritive salts and the exposure of the plants to prolonged illumination or, better still, an increase in the intensity of the light, the efficiency of illumination depending on the consequent formation of organic substances such as carbohydrates.

The quantitative alterations of external conditions may be spoken of as releasing stimuli. They produce, in the complex equilibrium of the cell, quantitative modifications in the arrangement and distribution of mass, by means of which other chemical processes are at once set in motion, and finally a new condition of equilibrium is attained. But the commonly expressed view that the environment can as a rule act only as a releasing agent is incorrect, because it overlooks an essential point. The power of a cell to receive stimuli is only acquired as the result of previous nutrition, which has produced a definite condition of concentration of different substances. Quantities are in this case the determining factors. The distribution of quantities is especially important in the sexual reproduction of algae, for which a vigorous production of the materials formed during carbon-assimilation appears to be essential.

In the Flowering plants, on the other hand, for reasons already mentioned, the whole problem is more complicated. Investigations on changes in the course of development of fertilised eggs have hitherto been unsuccessful; the difficulty of influencing egg-cells deeply immersed in tissue constitutes a serious obstacle. Other parts of plants are, however, convenient objects of experiment; e.g. the growing apices of buds which serve as cuttings for reproductive purposes, or buds on tubers, runners, rhizomes, etc. A growing apex consists of cells capable of division in which, as in egg-cells, a complete series of latent possibilities of development is embodied. Which of these possibilities becomes effective depends upon the action of the outer world transmitted by organs concerned with nutrition.

Of the different stages which a flowering plant passes through in the course of its development we will deal only with one in order to show that, in spite of its great complexity, the problem is, in essentials, equally open to attack in the higher plants and in the simplest organisms. The most important stage in the life of a flowering plant is the transition from purely vegetative growth to sexual reproduction--that is, the production of flowers. In certain cases it can be demonstrated that there is no internal cause, dependent simply on the specific structure, which compels a plant to produce its flowers after a definite period of vegetative growth. (Klebs, "Willkurliche Entwickelungsanderungen", Jena 1903; see also "Probleme der Entwickelung", I. II. "Centralbl." 1904.)

One extreme case, that of exceptionally early flowering, has been observed in nature and more often in cultivation. A number of plants under certain conditions are able to flower soon after germination. (Cf. numerous records of this kind by Diels, "Jugendformen und Bluten", Berlin, 1906.) This shortening of the period of development is exhibited in the most striking form in trees, as in the oak (Mobius, "Beitrage zur Lehre von der Fortpflanzung", Jena, 1897, page 89.), flowering seedlings of which have been observed from one to three years old, whereas normally the tree does not flower until it is sixty or eighty years old.

Another extreme case is represented by prolonged vegetative growth leading to the complete suppression of flower-production. This result may be obtained with several plants, such as Glechoma, the sugar beet, Digitalis, and others, if they are kept during the winter in a warm, damp atmosphere, and in rich soil; in the following spring or summer they fail to flower. (Klebs, "Willkurliche Aenderungen", etc. Jena, 1903, page 130.) Theoretically, however, experiments are of greater importance in which the production of flowers is inhibited by very favourable conditions of nutrition (Klebs, "Ueber kunstliche Metamorphosen", Stuttgart, 1906, page 115) ("Abh. Naturf. Ges. Halle", XXV.) occurring at the normal flowering period. Even in the case of plants of Sempervivum several years old, which, as is shown by control experiments on precisely similar plants, are on the point of flowering, flowering is rendered impossible if they are forced to very vigorous growth by an abundant supply of water and salts in the spring. Flowering, however, occurs, if such plants are cultivated in relatively dry sandy soil and in the presence of strong light. Careful researches into the conditions of growth have led, in the cases Sempervivum, to the following results: (1) With a strong light and vigorous carbon-assimilation a considerably increased supply of water and nutritive salts produces active vegetative growth. (2) With a vigorous carbon-assimilation in strong light, and a decrease in the supply of water and salts active flower-production is induced. (3) If an average supply of water and salts is given both processes are possible; the intensity of carbon-assimilation determines which of the two is manifested. A diminution in the production of organic substances, particularly of carbohydrates, induces vegetative growth. This can be effected by culture in feeble light or in light deprived of the yellow-red rays: on the other hand, flower-production follows an increase in light-intensity. These results are essentially in agreement with well-known observations on cultivated plants, according to which, the application of much moisture, after a plentiful supply of manure composed of inorganic salts, hinders the flower-production of many vegetables, while a decrease in the supply of water and salts favours flowering.

ii. INFLUENCE OF THE ENVIRONMENT ON THE FORM OF SINGLE ORGANS. (A considerable number of observations bearing on this question are given by Goebel in his "Experimentelle Morphologie der Pflanzen", Leipzig, 1908. It is not possible to deal here with the alteration in anatomical structure; cf. Kuster, "Pathologische Pflanzenanatomie", Jena, 1903.)

If we look closely into the development of a flowering plant, we notice that in a given species differently formed organs occur in definite positions. In a potato plant colourless runners are formed from the base of the main stem which grow underground and produce tubers at their tips: from a higher level foliage shoots arise nearer the apex. External appearances suggest that both the place of origin and the form of these organs were predetermined in the egg-cell or in the tuber. But it was shown experimentally by the well-known investigator Knight (Knight, "Selection from the Physiological and Horticultural Papers", London, 1841.) that tubers may be developed on the aerial stem in place of foliage shoots. These observations were considerably extended by Vochting. (Vochting, "Ueber die Bildung der Knollen", Cassel, 1887; see also "Bot. Zeit." 1902, 87.) In one kind of potato, germinating tubers were induced to form foliage shoots under the influence of a higher temperature; at a lower temperature they formed tuber-bearing shoots. Many other examples of the conversion of foliage-shoots into runners and rhizomes, or vice versa, have been described by Goebel and others. As in the asexual reproduction of algae quantitative alteration in the amount of moisture, light, temperature, etc. determines whether this or that form of shoot is produced. If the primordia of these organs are exposed to altered conditions of nutrition at a sufficiently early stage a complete substitution of one organ for another is effected. If the rudiment has reached a certain stage in development before it is exposed to these influences, extraordinary intermediate forms are obtained, bearing the characters of both organs.

The study of regeneration following injury is of greater importance as regards the problem of the development and place of origin of organs. (Reference may be made to the full summary of results given by Goebel in his "Experimentelle Morphologie", Leipzig and Berlin, 1908, Section IV.) Only in relatively very rare cases is there a complete re-formation of the injured organ itself, as e.g. in the growing-apex. Much more commonly injury leads to the development of complementary formations, it may be the rejuvenescence of a hitherto dormant rudiment, or it may be the formation of such ab initio. In all organs, stems, roots, leaves, as well as inflorescences, this kind of regeneration, which occurs in a great variety of ways according to the species, may be observed on detached pieces of the plant. Cases are also known, such, for example, as the leaves of many plants which readily form roots but not shoots, where a complete regeneration does not occur.

The widely spread power of reacting to wounding affords a very valuable means of inducing a fresh development of buds and roots on places where they do not occur in normal circumstances. Injury creates special conditions, but little is known as yet in regard to alterations directly produced in this way. Where the injury consists in the separation of an organ from its normal connections, the factors concerned are more comprehensible. A detached leaf, e.g., is at once cut off from a supply of water and salts, and is deprived of the means of getting rid of organic substances which it produces; the result is a considerable alteration in the degree of concentration. No experimental investigation on these lines has yet been made. Our ignorance has often led to the view that we are dealing with a force whose specific quality is the restitution of the parts lost by operation; the proof, therefore, that in certain cases a similar production of new roots or buds may be induced without previous injury and simply by a change in external conditions assumes an importance. (Klebs, "Willkurliche Entwickelung", page 100; also, "Probleme der Entwickelung", "Biol. Centralbl." 1904, page 610.)

A specially striking phenomenon of regeneration, exhibited also by uninjured plants, is afforded by polarity, which was discovered by Vochting. (See the classic work of Vochting, "Ueber Organbildung im Pflanzenreich", I. Bonn, 1888; also "Bot. Zeit." 1906, page 101; cf. Goebel, "Experimentelle Morphologie", Leipzig and Berlin, 1908, Section V, Polaritat.) It is found, for example, that roots are formed from the base of a detached piece of stem and shoots from the apex. Within the limits of this essay it is impossible to go into this difficult question; it is, however, important from the point of view of our general survey to emphasise the fact that the physiological distinctions between base and apex of pieces of stem are only of a quantitative kind, that is, they consist in the inhibition of certain phenomena or in favouring them. As a matter of fact roots may be produced from the apices of willows and cuttings of other plants; the distinction is thus obliterated under the influence of environment. The fixed polarity of cuttings from full grown stems cannot be destroyed; it is the expression of previous development. Vochting speaks of polarity as a fixed inherited character. This is an unconvincing conclusion, as nothing can be deduced from our present knowledge as to the causes which led up to polarity. We know that the fertilised egg, like the embryo, is fixed at one end by which it hangs freely in the embryo-sac and afterwards in the endosperm. From the first, therefore, the two ends have different natures, and these are revealed in the differentiation into root-apex and stem-apex. A definite direction in the flow of food-substances is correlated with this arrangement, and this eventually leads to a polarity in the tissues. This view requires experimental proof, which in the case of the egg-cells of flowering plants hardly appears possible; but it derives considerable support from the fact that in herbaceous plants, e.g. Sempervivum (Klebs, "Variationen der Bluten", "Jahrb. Wiss. Bot." 1905, page 260.), rosettes or flower-shoots are formed in response to external conditions at the base, in the middle, or at the apex of the stem, so that polarity as it occurs under normal conditions cannot be the result of unalterable hereditary factors. On the other hand, the lower plants should furnish decisive evidence on this question, and the experiments of Stahl, Winkler, Kniep, and others indicate the right method of attacking the problem.

The relation of leaf-form to environment has often been investigated and is well known. The leaves of bog and water plants (Cf.Goebel, loc. cit. chapter II.; also Gluck, "Untersuchungen uber Wasser- und Sumpfgewachse", Jena, Vols. I.-II. 1905-06.) afford the most striking examples of modifications: according as they are grown in water, moist or dry air, the form of the species characteristic of the particular habitat is produced, since the stems are also modified. To the same group of phenomena belongs the modification of the forms of leaves and stems in plants on transplantation from the plains to the mountains (Bonnier, "Recherches sur l'Anatomie experimentale des Vegetaux", Corbeil, 1895.) or vice versa. Such variations are by no means isolated examples. All plants exhibit a definite alteration in form as the result of prolonged cultivation in moist or dry air, in strong or feeble light, or in darkness, or in salt solutions of different composition and strength.

Every individual which is exposed to definite combinations of external factors exhibits eventually the same type of modification. This is the type of variation which Darwin termed "definite." It is easy to realise that indefinite or fluctuating variations belong essentially to the same class of phenomena; both are reactions to changes in environment. In the production of individual variations two different influences undoubtedly cooperate. One set of variations is caused by different external conditions, during the production, either of sexual cells or of vegetative primordia; another set is the result of varying external conditions during the development of the embryo into an adult plant. The two sets of influences cannot as yet be sharply differentiated. If, for purposes of vegetative reproduction, we select pieces of the same parent-plant of a pure species, the second type of variation predominates. Individual fluctuations depend essentially in such cases on small variations in environment during development.

These relations must be borne in mind if we wish to understand the results of statistical methods. Since the work of Quetelet, Galton, and others the statistical examination of individual differences in animals and plants has become a special science, which is primarily based on the consideration that the application of the theory of probability renders possible mathematical statement and control of the results. The facts show that any character, size of leaf, length of stem, the number of members in a flower, etc. do not vary haphazard but in a very regular manner. In most cases it is found that there is a value which occurs most commonly, the average or medium value, from which the larger and smaller deviations, the so-called plus and minus variations fall away in a continuous series and end in a limiting value. In the simpler cases a falling off occurs equally on both sides of the curve; the curve constructed from such data agrees very closely with the Gaussian curve of error. In more complicated cases irregular curves of different kinds are obtained which may be calculated on certain suppositions.

The regular fluctuations about a mean according to the rule of probability is often attributed to some law underlying variability. (de Vries, "Mutationstheorie", Vol. I. page 35, Leipzig, 1901.) But there is no such law which compels a plant to vary in a particular manner. Every experimental investigation shows, as we have already remarked, that the fluctuation of characters depends on fluctuation in the external factors. The applicability of the method of probability follows from the fact that the numerous individuals of a species are influenced by a limited number of variable conditions. (Klebs, "Willkurl. Ent." Jena, 1903, page 141.) As each of these conditions includes within certain limits all possible values and exhibits all possible combinations, it follows that, according to the rules of probability, there must be a mean value, about which the larger and smaller deviations are distributed. Any character will be found to have the mean value which corresponds with that combination of determining factors which occurs most frequently. Deviations towards plus and minus values will be correspondingly produced by rarer conditions.

A conclusion of fundamental importance may be drawn from this conception, which is, to a certain extent, supported by experimental investigation. (Klebs, "Studien uber Variation", "Arch. fur Entw." 1907.) There is no normal curve for a particular CHARACTER, there is only a curve for the varying combinations of conditions occurring in nature or under cultivation. Under other conditions entirely different curves may be obtained with other variants as a mean value. If, for example, under ordinary conditions the number 10 is the most frequent variant for the stamens of Sedum spectabile, in special circumstances (red light) this is replaced by the number 5. The more accurately we know the conditions for a particular form or number, and are able to reproduce it by experiment, the nearer we are to achieving our aim of rendering a particular variation impossible or of making it dominant.

In addition to the individual variations of a species, more pronounced fluctuations occur relatively rarely and sporadically which are spoken of as "single variations," or if specially striking as abnormalities or monstrosities. These forms have long attracted the attention of morphologists; a large number of observations of this kind are given in the handbooks of Masters (Masters, "Vegetable Teratology", London, 1869.) and Penzig (Penzig, "Pflanzen-Teratologie", Vols I. and II. Genua, 1890-94.) These variations, which used to be regarded as curiosities, have now assumed considerable importance in connection with the causes of form-development. They also possess special interest in relation to the question of heredity, a subject which does not at present concern us, as such deviations from normal development undoubtedly arise as individual variations induced by the influence of environment.

Abnormal developments of all kinds in stems, leaves, and flowers, may be produced by parasites, insects, or fungi. They may also be induced by injury, as Blaringhem (Blaringhem, "Mutation et traumatismes", Paris, 1907.) has more particularly demonstrated, which, by cutting away the leading shoots of branches in an early stage of development, caused fasciation, torsion, anomalous flowers, etc. The experiments of Blaringhem point to the probability that disturbances in the conditions of food-supply consequent on injury are the cause of the production of monstrosities. This is certainly the case in my experiments with species of Sempervivum (Klebs, "Kunstliche Metamorphosen", Stuttgart, 1906.); individuals, which at first formed normal flowers, produced a great variety of abnormalities as the result of changes in nutrition, we may call to mind the fact that the formation of inflorescences occurs normally when a vigorous production of organic compounds, such as starch, sugar, etc. follows a diminution in the supply of mineral salts. On the other hand, the development of inflorescences is entirely suppressed if, at a suitable moment before the actual foundations have been laid, water and mineral salts are supplied to the roots. If, during the week when the inflorescence has just been laid down and is growing very slowly, the supply of water and salts is increased, the internal conditions of the cells are essentially changed. At a later stage, after the elongation of the inflorescence, rosettes of leaves are produced instead of flowers, and structures intermediate between the two kinds of organs; a number of peculiar plant-forms are thus obtained (Cf. Lotsy, "Vorlesungen uber Deszendenztheorien", Vol. II. pl. 3, Jena, 1908.) Abnormalities in the greatest variety are produced in flowers by varying the time at which the stimulus is applied, and by the cooperation of other factors such as temperature, darkness, etc. In number and arrangement the several floral members vary within wide limits; sepals, petals, stamens, and carpels are altered in form and colour, a transformation of stamens to carpels and from carpels to stamens occurs in varying degrees. The majority of the deviations observed had not previously been seen either under natural conditions or in cultivation; they were first brought to light through the influence of external factors.

Such transformations of flowers become apparent at a time, which is separated by about two months from the period at which the particular cause began to act. There is, therefore, no close connection between the appearance of the modifications and the external conditions which prevail at the moment. When we are ignorant of the causes which are operative so long before the results are seen, we gain the impression that such variations as occur are spontaneous or autonomous expressions of the inner nature of the plant. It is much more likely that, as in Sempervivum, they were originally produced by an external stimulus which had previously reached the sexual cells or the young embryo. In any case abnormalities of this kind appear to be of a special type as compared with ordinary fluctuating variations. Darwin pointed out this difference; Bateson (Bateson, "Materials for the study of Variation", London, 1894, page 5.) has attempted to make the distinction sharper, at the same time emphasising its importance in heredity.

Bateson applies the term CONTINUOUS to small variations connected with one another by transitional stages, while those which are more striking and characterised from the first by a certain completeness, he names DISCONTINUOUS. He drew attention to a great difficulty which stands in the way of Lamarck's hypothesis, as also of Darwin's view. "According to both theories, specific diversity of form is consequent upon diversity of environment, and diversity of environment is thus the ultimate measure of diversity of specific form. Here then we meet the difficulty that diverse environments often shade into each other insensibly and form a continuous series, whereas the Specific Forms of life which are subject to them on the whole form a Discontinuous Series." This difficulty is, however, not of fundamental importance as well authenticated facts have been adduced showing that by alteration of the environment discontinuous variations, such as alterations in the number and form of members of a flower, may be produced. We can as yet no more explain how this happens than we can explain the existence of continuous variations. We can only assert that both kinds of variation arise in response to quantitative alterations in external conditions. The question as to which kind of variation is produced depends on the greater or less degree of alteration; it is correlated with the state of the particular cells at the moment.

In this short sketch it is only possible to deal superficially with a small part of the subject. It has been clearly shown that in view of the general dependence of development on the factors of the environment a number of problems are ready for experimental treatment. One must, however, not forget that the science of the physiology of form has not progressed beyond its initial stages. Just now our first duty is to demonstrate the dependence on external factors in as many forms of plants as possible, in order to obtain a more thorough control of all the different plant-forms. The problem is not only to produce at will (and independently of their normal mode of life) forms which occur in nature, but also to stimulate into operation potentialities which necessarily lie dormant under the conditions which prevail in nature. The constitution of a species is much richer in possibilities of development than would appear to be the case under normal conditions. It remains for man to stimulate into activity all the potentialities.

But the control of plant-form is only a preliminary step--the foundation stones on which to erect a coherent scientific structure. We must discover what are the internal processes in the cell produced by external factors, which as a necessary consequence result in the appearance of a definite form. We are here brought into contact with the most obscure problem of life. Progress can only be made pari passu with progress in physics and chemistry, and with the growth of our knowledge of nutrition, growth, etc.

Let us take one of the simplest cases--an alteration in form. A cylindrical cell of the alga Stigeoclonium assumes, as Livingstone (Livingstone, "On the nature of the stimulus which causes the change of form, etc." "Botanical Gazette", XXX. 1900; also XXXII. 1901.) has shown, a spherical form when the osmotic pressure of the culture fluid is increased; or a spore of Mucor, which, in a sugar solution grows into a branched filament, in the presence of a small quantity of acid (hydrogen ions) becomes a comparatively large sphere. (Ritter, "Ueber Kugelhefe, etc." "Ber. bot. Gesell." Berlin, XXV. page 255, 1907.) In both cases there has undoubtedly been an alteration in the osmotic pressure of the cell-sap, but this does not suffice to explain the alteration in form, since the unknown alterations, which are induced in the protoplasm, must in their turn influence the cell-membrane. In the case of the very much more complex alterations in form, such as we encounter in the course of development of plants, there do not appear to be any clues which lead us to a deeper insight into the phenomena. Nevertheless we continue the attempt, seeking with the help of any available hypothesis for points of attack, which may enable us to acquire a more complete mastery of physiological methods. To quote a single example; I may put the question, what internal changes produce a transition from vegetative growth to sexual reproduction?

The facts, which are as clearly established from the lower as for the higher plants, teach us that quantitative alteration in the environment produces such a transition. This suggests the conclusion that quantitative internal changes in the cells, and with them disturbances in the degree of concentration, are induced, through which the chemical reactions are led in the direction of sexual reproduction. An increase in the production of organic substances in the presence of light, chiefly of the carbohydrates, with a simultaneous decrease in the amount of inorganic salts and water, are the cause of the disturbance and at the same time of the alteration in the direction of development. Possibly indeed mineral salts as such are not in question, but only in the form of other organic combinations, particularly proteid material, so that we are concerned with an alteration in the relation of the carbohydrates and proteids. The difficulties of such researches are very great because the methods are not yet sufficiently exact to demonstrate the frequently small quantitative differences in chemical composition. Questions relating to the enzymes, which are of the greatest importance in all these life-processes, are especially complicated. In any case it is the necessary result of such an hypothesis that we must employ chemical methods of investigation in dealing with problems connected with the physiology of form.

II. INFLUENCE OF ENVIRONMENT ON THE TRANSFORMATION OF SPECIES.

The study of the physiology of form-development in a pure species has already yielded results and makes slow but sure progress. The physiology of the possibility of the transformation of one species into another is based, as yet, rather on pious hope than on accomplished fact. From the first it appeared to be hopeless to investigate physiologically the origin of Linnean species and at the same time that of the natural system, an aim which Darwin had before him in his enduring work. The historical sequence of events, of which an organism is the expression, can only be treated hypothetically with the help of facts supplied by comparative morphology, the history of development, geographical distribution, and palaeontology. (See Lotsy, "Vorlesungen" (Jena, I. 1906, II. 1908), for summary of the facts.) A glance at the controversy which is going on today in regard to different hypotheses shows that the same material may lead different investigators to form entirely different opinions. Our ultimate aim is to find a solution of the problem as to the cause of the origin of species. Indeed such attempts are now being made: they are justified by the fact that under cultivation new and permanent strains are produced; the fundamental importance of this was first grasped by Darwin. New points of view in regard to these lines of inquiry have been adopted by H. de Vries who has succeeded in obtaining from Oenothera Lamarckiana a number of constant "elementary" species. Even if it is demonstrated that he was simply dealing with the complex splitting up of a hybrid (Bateson, "Reports to the Evolution Committee of the Royal Society", London, 1902; cf. also Lotsy, "Vorlesungen", Vol. I. page 234.), the facts adduced in no sense lose their very great value.

We must look at the problem in its simplest form; we find it in every case where a new race differs essentially from the original type in a single character only; for example, in the colour of the flowers or in the petalody of the stamens (doubling of flowers). In this connection we must keep in view the fact that every visible character in a plant is the resultant of the cooperation of specific structure, with its various potentialities, and the influence of the environment. We know, that in a pure species all characters vary, that a blue-flowering Campanula or a red Sempervivum can be converted by experiment into white-flowering forms, that a transformation of stamens into petals may be caused by fungi or by the influence of changed conditions of nutrition, or that plants in dry and poor soil become dwarfed. But so far as the experiments justify a conclusion, it would appear that such alterations are not inherited by the offspring. Like all other variations they appear only so long as special conditions prevail in the surroundings.

It has been shown that the case is quite different as regards the white-flowering, double or dwarf races, because these retain their characters when cultivated under practically identical conditions, and side by side with the blue, single-flowering or tall races. The problem may therefore be stated thus: how can a character, which appears in the one case only under the strictly limited conditions of the experiment, in other cases become apparent under the very much wider conditions of ordinary cultivation? If a character appears, in these circumstances, in the case of all individuals, we then speak of constant races. In such simple cases the essential point is not the creation of a new character but rather an ALTERATION OF THIS CHARACTER IN ACCORDANCE WITH THE ENVIRONMENT. In the examples mentioned the modified character in the simple varieties (or a number of characters in elementary species) appears more or less suddenly and is constant in the above sense. The result is what de Vries has termed a Mutation. In this connection we must bear in mind the fact that no difference, recognisable externally, need exist between individual variation and mutation. Even the most minute quantitative difference between two plants may be of specific value if it is preserved under similar external conditions during many successive generations. We do not know how this happens. We may state the problem in other terms; by saying that the specific structure must be altered. It is possible, to some extent, to explain this sudden alteration, if we regard it as a chemical alteration of structure either in the specific qualities of the proteids or of the unknown carriers of life. In the case of many organic compounds their morphological characters (the physical condition, crystalline form, etc.) are at once changed by alteration of atomic relations or by incorporation of new radicals. (For instance ethylchloride (C2H5Cl) is a gas at 21 deg C., ethylenechloride (C2H4Cl2) a fluid boiling at 84 deg C., beta trichlorethane (C2H3Cl3) a fluid boiling at 113 deg C., perchlorethane (C2Cl6) a crystalline substance. Klebs, ("Willkurliche Entwickelungsanderungen" page 158.) Much more important, however, would be an answer to the question, whether an individual variation can be converted experimentally into an inherited character--a mutation in de Vries's sense.

In all circumstances we may recognise as a guiding principle the assumption adopted by Lamarck, Darwin, and many others, that the inheritance of any one character, or in more general terms, the transformation of one species into another, is, in the last instance, to be referred to a change in the environment. From a causal-mechanical point of view it is not a priori conceivable that one species can ever become changed into another so long as external conditions remain constant. The inner structure of a species must be essentially altered by external influences. Two methods of experimental research may be adopted, the effect of crossing distinct species and, secondly, the effect of definite factors of the environment.

The subject of hybridisation is dealt with in another part of this essay. It is enough to refer here to the most important fact, that as the result of combinations of characters of different species new and constant forms are produced. Further, Tschermack, Bateson and others have demonstrated the possibility that hitherto unknown inheritable characters may be produced by hybridisation.

The other method of producing constant races by the influence of special external conditions has often been employed. The sporeless races of Bacteria and Yeasts (Cf. Detto, "Die Theorie der direkten Anpassung... ", pages 98 et seq., Jena, 1904; see also Lotsy, "Vorlesungen", II. pages 636 et seq., where other similar cases are described.) are well known, in which an internal alteration of the cells is induced by the influence of poison or higher temperature, so that the power of producing spores even under normal conditions appears to be lost. A similar state of things is found in some races which under certain definite conditions lose their colour or their virulence. Among the phanerogams the investigations of Schubler on cereals afford parallel cases, in which the influence of a northern climate produces individuals which ripen their seeds early; these seeds produce plants which seed early in southern countries. Analogous results were obtained by Cieslar in his experiments; seeds of conifers from the Alps when planted in the plains produced plants of slow growth and small diameter.

All these observations are of considerable interest theoretically; they show that the action of environment certainly induces such internal changes, and that these are transmitted to the next generation. But as regards the main question, whether constant races may be obtained by this means, the experiments cannot as yet supply a definite answer. In phanerogams, the influence very soon dies out in succeeding generations; in the case of bacteria, in which it is only a question of the loss of a character it is relatively easy for this to reappear. It is not impossible, that in all such cases there is a material hanging-on of certain internal conditions, in consequence of which the modification of the character persists for a time in the descendants, although the original external conditions are no longer present.

Thus a slow dying-out of the effect of a stimulus was seen in my experiments on Veronica chamaedrys. (Klebs, "Kunstliche Metamorphosen", Stuttgart, 1906, page 132.) During the cultivation of an artificially modified inflorescence I obtained a race showing modifications in different directions, among which twisting was especially conspicuous. This plant, however, does not behave as the twisted race of Dipsacus isolated by de Vries (de Vries, "Mutationstheorie", Vol. II. Leipzig, 1903, page 573.), which produced each year a definite percentage of twisted individuals. In the vegetative reproduction of this Veronica the torsion appeared in the first, also in the second and third year, but with diminishing intensity. In spite of good cultivation this character has apparently now disappeared; it disappeared still more quickly in seedlings. In another character of the same Veronica chamaedrys the influence of the environment was stronger. The transformation of the inflorescences to foliage-shoots formed the starting-point; it occurred only under narrowly defined conditions, namely on cultivation as a cutting in moist air and on removal of all other leaf-buds. In the majority (7/10) of the plants obtained from the transformed shoots, the modification appeared in the following year without any interference. Of the three plants which were under observation several years the first lost the character in a short time, while the two others still retain it, after vegetative propagation, in varying degrees. The same character occurs also in some of the seedlings; but anything approaching a constant race has not been produced.

Another means of producing new races has been attempted by Blaringhem. (Blaringhem, "Mutation et Traumatisme", Paris, 1907.) On removing at an early stage the main shoots of different plants he observed various abnormalities in the newly formed basal shoots. From the seeds of such plants he obtained races, a large percentage of which exhibited these abnormalities. Starting from a male Maize plant with a fasciated inflorescence, on which a proportion of the flowers had become male, a new race was bred in which hermaphrodite flowers were frequently produced. In the same way Blaringhem obtained, among other similar results, a race of barley with branched ears. These races, however, behaved in essentials like those which have been demonstrated by de Vries to be inconstant, e.g. Trifolium pratense quinquefolium and others. The abnormality appears in a proportion of the individuals and only under very special conditions. It must be remembered too that Blaringhem worked with old cultivated plants, which from the first had been disposed to split into a great variety of races. It is possible, but difficult to prove, that injury contributed to this result.

A third method has been adopted by MacDougal (MacDougal, "Heredity and Origin of species", "Monist", 1906; "Report of department of botanical research", "Fifth Year-book of the Carnegie Institution of Washington", page 119, 1907.) who injected strong (10 percent) sugar solution or weak solutions of calcium nitrate and zinc sulphate into young carpels of different plants. From the seeds of a plant of Raimannia odorata the carpels of which had been thus treated he obtained several plants distinguished from the parent-forms by the absence of hairs and by distinct forms of leaves. Further examination showed that he had here to do with a new elementary species. MacDougal also obtained a more or less distinct mutant of Oenothera biennis. We cannot as yet form an opinion as to how far the effect is due to the wound or to the injection of fluid as such, or to its chemical properties. This, however, is not so essential as to decide whether the mutant stands in any relation to the influence of external factors. It is at any rate very important that this kind of investigation should be carried further.

If it could be shown that new and inherited races were obtained by MacDougal's method, it would be safe to conclude that the same end might be gained by altering the conditions of the food-stuff conducted to the sexual cells. New races or elementary species, however, arise without wounding or injection. This at once raises the much discussed question, how far garden-cultivation has led to the creation of new races? Contrary to the opinion expressed by Darwin and others, de Vries ("Mutationstheorie", Vol. I. pages 412 et seq.) tried to show that garden-races have been produced only from spontaneous types which occur in a wild state or from sub-races, which the breeder has accidentally discovered but not originated. In a small number of cases only has de Vries adduced definite proof. On the other side we have the work of Korschinsky (Korschinsky, "Heterogenesis und Evolution", "Flora", 1901.) which shows that whole series of garden-races have made their appearance only after years of cultivation. In the majority of races we are entirely ignorant of their origin.

It is, however, a fact that if a plant is removed from natural conditions into cultivation, a well-marked variation occurs. The well-known plant-breeder L. de Vilmorin (L. de Vilmorin, "Notices sur l'amelioration des plantes", Paris, 1886, page 36.), speaking from his own experience, states that a plant is induced to "affoler," that is to exhibit all possible variations from which the breeder may make a further selection only after cultivation for several generations. The effect of cultivation was particularly striking in Veronica chamaedrys (Klebs, "Kunstliche Metamorphosen", Stuttgart, 1906, page 152.) which, in spite of its wide distribution in nature, varies very little. After a few years of cultivation this "good" and constant species becomes highly variable. The specimens on which the experiments were made were three modified inflorescence cuttings, the parent-plants of which certainly exhibited no striking abnormalities. In a short time many hitherto latent potentialities became apparent, so that characters, never previously observed, or at least very rarely, were exhibited, such as scattered leaf-arrangement, torsion, terminal or branched inflorescences, the conversion of the inflorescence into foliage-shoots, every conceivable alteration in the colour of flowers, the assumption of a green colour by parts of the flowers, the proliferation of flowers.

All this points to some disturbance in the species resulting from methods of cultivation. It has, however, not yet been possible to produce constant races with any one of these modified characters. But variations appeared among the seedlings, some of which, e.g. yellow variegation, were not inheritable, while others have proved constant. This holds good, so far as we know at present, for a small rose-coloured form which is to be reckoned as a mutation. Thus the prospect of producing new races by cultivation appears to be full of promise.

So long as the view is held that good nourishment, i.e. a plentiful supply of water and salts, constitutes the essential characteristic of garden-cultivation, we can hardly conceive that new mutations can be thus produced. But perhaps the view here put forward in regard to the production of form throws new light on this puzzling problem.

Good manuring is in the highest degree favourable to vegetative growth, but is in no way equally favourable to the formation of flowers. The constantly repeated expression, good or favourable nourishment, is not only vague but misleading, because circumstances favourable to growth differ from those which promote reproduction; for the production of every form there are certain favourable conditions of nourishment, which may be defined for each species. Experience shows that, within definite and often very wide limits, it does not depend upon the ABSOLUTE AMOUNT of the various food substances, but upon their respective degrees of concentration. As we have already stated, the production of flowers follows a relative increase in the amount of carbohydrates formed in the presence of light, as compared with the inorganic salts on which the formation of albuminous substances depends. (Klebs, "Kunstliche Metamorphosen", page 117.) The various modifications of flowers are due to the fact that a relatively too strong solution of salts is supplied to the rudiments of these organs. As a general rule every plant form depends upon a certain relation between the different chemical substances in the cells and is modified by an alteration of that relation.

During long cultivation under conditions which vary in very different degrees, such as moisture, the amount of salts, light intensity, temperature, oxygen, it is possible that sudden and special disturbances in the relations of the cell substances have a directive influence on the inner organisation of the sexual cells, so that not only inconstant but also constant varieties will be formed.

Definite proof in support of this view has not yet been furnished, and we must admit that the question as to the cause of heredity remains, fundamentally, as far from solution as it was in Darwin's time. As the result of the work of many investigators, particularly de Vries, the problem is constantly becoming clearer and more definite. The penetration into this most difficult and therefore most interesting problem of life and the creation by experiment of new races or elementary species are no longer beyond the region of possibility.

XIV. EXPERIMENTAL STUDY OF THE INFLUENCE OF ENVIRONMENT ON ANIMALS. By Jacques Loeb, M.D. Professor of Physiology in the University of California.

I. INTRODUCTORY REMARKS.

What the biologist calls the natural environment of an animal is from a physical point of view a rather rigid combination of definite forces. It is obvious that by a purposeful and systematic variation of these and by the application of other forces in the laboratory, results must be obtainable which do not appear in the natural environment. This is the reasoning underlying the modern development of the study of the effects of environment upon animal life. It was perhaps not the least important of Darwin's services to science that the boldness of his conceptions gave to the experimental biologist courage to enter upon the attempt of controlling at will the life-phenomena of animals, and of bringing about effects which cannot be expected in Nature.

The systematic physico-chemical analysis of the effect of outside forces upon the form and reactions of animals is also our only means of unravelling the mechanism of heredity beyond the scope of the Mendelian law. The manner in which a germ-cell can force upon the adult certain characters will not be understood until we succeed in varying and controlling hereditary characteristics; and this can only be accomplished on the basis of a systematic study of the effects of chemical and physical forces upon living matter.

Owing to limitation of space this sketch is necessarily very incomplete, and it must not be inferred that studies which are not mentioned here were considered to be of minor importance. All the writer could hope to do was to bring together a few instances of the experimental analysis of the effect of environment, which indicate the nature and extent of our control over life-phenomena and which also have some relation to the work of Darwin. In the selection of these instances preference is given to those problems which are not too technical for the general reader.

The forces, the influence of which we shall discuss, are in succession chemical agencies, temperature, light, and gravitation. We shall also treat separately the effect of these forces upon form and instinctive reactions.

II. THE EFFECTS OF CHEMICAL AGENCIES.

(a) HETEROGENEOUS HYBRIDISATION.

It was held until recently that hybridisation is not possible except between closely related species and that even among these a successful hybridisation cannot always be counted upon. This view was well supported by experience. It is, for instance, well known that the majority of marine animals lay their unfertilised eggs in the ocean and that the males shed their sperm also into the sea-water. The numerical excess of the spermatozoa over the ova in the sea-water is the only guarantee that the eggs are fertilised, for the spermatozoa are carried to the eggs by chance and are not attracted by the latter. This statement is the result of numerous experiments by various authors, and is contrary to common belief. As a rule all or the majority of individuals of a species in a given region spawn on the same day, and when this occurs the sea-water constitutes a veritable suspension of sperm. It has been shown by experiment that in fresh sea-water the sperm may live and retain its fertilising power for several days. It is thus unavoidable that at certain periods more than one kind of spermatozoon is suspended in the sea-water and it is a matter of surprise that the most heterogeneous hybridisations do not constantly occur. The reason for this becomes obvious if we bring together mature eggs and equally mature and active sperm of a different family. When this is done no egg is, as a rule, fertilised. The eggs of a sea-urchin can be fertilised by sperm of their own species, or, though in smaller numbers, by the sperm of other species of sea-urchins, but not by the sperm of other groups of echinoderms, e.g. starfish, brittle-stars, holothurians or crinoids, and still less by the sperm of more distant groups of animals. The consensus of opinion seemed to be that the spermatozoon must enter the egg through a narrow opening or canal, the so-called micropyle, and that the micropyle allowed only the spermatozoa of the same or of a closely related species to enter the egg.

It seemed to the writer that the cause of this limitation of hybridisation might be of another kind and that by a change in the constitution of the sea-water it might be possible to bring about heterogenous hybridisations, which in normal sea-water are impossible. This assumption proved correct. Sea-water has a faintly alkaline reaction (in terms of the physical chemist its concentration of hydroxyl ions is about (10 to the power minus six)N at Pacific Grove, California, and about (10 to the power minus 5)N at Woods Hole, Massachusetts). If we slightly raise the alkalinity of the sea-water by adding to it a small but definite quantity of sodium hydroxide or some other alkali, the eggs of the sea-urchin can be fertilised with the sperm of widely different groups of animals, possibly with the sperm of any marine animal which sheds it into the ocean. In 1903 it was shown that if we add from about 0.5 to 0.8 cubic centimetre N/10 sodium hydroxide to 50 cubic centimetres of sea-water, the eggs of Strongylocentrotus purpuratus (a sea-urchin which is found on the coast of California) can be fertilised in large quantities by the sperm of various kinds of starfish, brittle-stars and holothurians; while in normal sea-water or with less sodium hydroxide not a single egg of the same female could be fertilised with the starfish sperm which proved effective in the hyper-alkaline sea-water. The sperm of the various forms of starfish was not equally effective for these hybridisations; the sperm of Asterias ochracea and A. capitata gave the best results, since it was possible to fertilise 50 per cent or more of the sea-urchin eggs, while the sperm of Pycnopodia and Asterina fertilised only 2 per cent of the same eggs.

Godlewski used the same method for the hybridisation of the sea-urchin eggs with the sperm of a crinoid (Antedon rosacea). Kupelwieser afterwards obtained results which seemed to indicate the possibility of fertilising the eggs of Strongylocentrotus with the sperm of a mollusc (Mytilus.) Recently, the writer succeeded in fertilising the eggs of Strongylocentrotus franciscanus with the sperm of a mollusc--Chlorostoma. This result could only be obtained in sea-water the alkalinity of which had been increased (through the addition of 0.8 cubic centimetre N/10 sodium hydroxide to 50 cubic centimetres of sea-water). We thus see that by increasing the alkalinity of the sea-water it is possible to effect heterogeneous hybridisations which are at present impossible in the natural environment of these animals.

It is, however, conceivable that in former periods of the earth's history such heterogeneous hybridisations were possible. It is known that in solutions like sea-water the degree of alkalinity must increase when the amount of carbon-dioxide in the atmosphere is diminished. If it be true, as Arrhenius assumes, that the Ice age was caused or preceded by a diminution in the amount of carbon-dioxide in the air, such a diminution must also have resulted in an increase of the alkalinity of the sea-water, and one result of such an increase must have been to render possible heterogeneous hybridisations in the ocean which in the present state of alkalinity are practically excluded.

But granted that such hybridisations were possible, would they have influenced the character of the fauna? In other words, are the hybrids between sea-urchin and starfish, or better still, between sea-urchin and mollusc, capable of development, and if so, what is their character? The first experiment made it appear doubtful whether these heterogeneous hybrids could live. The sea-urchin eggs which were fertilised in the laboratory by the spermatozoa of the starfish, as a rule, died earlier than those of the pure breeds. But more recent results indicate that this was due merely to deficiencies in the technique of the earlier experiments. The writer has recently obtained hybrid larvae between the sea-urchin egg and the sperm of a mollusc (Chlorostoma) which, in the laboratory, developed as well and lived as long as the pure breeds of the sea-urchin, and there was nothing to indicate any difference in the vitality of the two breeds.

So far as the question of heredity is concerned, all the experiments on heterogeneous hybridisation of the egg of the sea-urchin with the sperm of starfish, brittle-stars, crinoids and molluscs, have led to the same result, namely, that the larvae have purely maternal characteristics and differ in no way from the pure breed of the form from which the egg is taken. By way of illustration it may be said that the larvae of the sea-urchin reach on the third day or earlier (according to species and temperature) the so-called pluteus stage, in which they possess a typical skeleton; while neither the larvae of the starfish nor those of the mollusc form a skeleton at the corresponding stage. It was, therefore, a matter of some interest to find out whether or not the larvae produced by the fertilisation of the sea-urchin egg with the sperm of starfish or mollusc would form the normal and typical pluteus skeleton. This was invariably the case in the experiments of Godlewski, Kupelwieser, Hagedoorn, and the writer. These hybrid larvae were exclusively maternal in character.

It might be argued that in the case of heterogeneous hybridisation the sperm-nucleus does not fuse with the egg-nucleus, and that, therefore, the spermatozoon cannot transmit its hereditary substances to the larvae. But these objections are refuted by Godlewski's experiments, in which he showed definitely that if the egg of the sea-urchin is fertilised with the sperm of a crinoid the fusion of the egg-nucleus and sperm-nucleus takes place in the normal way. It remains for further experiments to decide what the character of the adult hybrids would be.

(b). ARTIFICIAL PARTHENOGENESIS.

Possibly in no other field of Biology has our ability to control life-phenomena by outside conditions been proved to such an extent as in the domain of fertilisation. The reader knows that the eggs of the overwhelming majority of animals cannot develop unless a spermatozoon enters them. In this case a living agency is the cause of development and the problem arises whether it is possible to accomplish the same result through the application of well-known physico-chemical agencies. This is, indeed, true, and during the last ten years living larvae have been produced by chemical agencies from the unfertilised eggs of sea-urchins, starfish, holothurians and a number of annelids and molluscs; in fact this holds true in regard to the eggs of practically all forms of animals with which such experiments have been tried long enough. In each form the method of procedure is somewhat different and a long series of experiments is often required before the successful method is found.

The facts of Artificial Parthenogenesis, as the chemical fertilisation of the egg is called, have, perhaps, some bearing on the problem of evolution. If we wish to form a mental image of the process of evolution we have to reckon with the possibility that parthenogenetic propagation may have preceded sexual reproduction. This suggests also the possibility that at that period outside forces may have supplied the conditions for the development of the egg which at present the spermatozoon has to supply. For this, if for no other reason, a brief consideration of the means of artificial parthenogenesis may be of interest to the student of evolution.

It seemed necessary in these experiments to imitate as completely as possible by chemical agencies the effects of the spermatozoon upon the egg. When a spermatozoon enters the egg of a sea-urchin or certain starfish or annelids, the immediate effect is a characteristic change of the surface of the egg, namely the formation of the so-called membrane of fertilisation. The writer found that we can produce this membrane in the unfertilised egg by certain acids, especially the monobasic acids of the fatty series, e.g. formic, acetic, propionic, butyric, etc. Carbon-dioxide is also very efficient in this direction. It was also found that the higher acids are more efficient than the lower ones, and it is possible that the spermatozoon induces membrane-formation by carrying into the egg a higher fatty acid, namely oleic acid or one of its salts or esters.

The physico-chemical process which underlies the formation of the membrane seems to be the cause of the development of the egg. In all cases in which the unfertilised egg has been treated in such a way as to cause it to form a membrane it begins to develop. For the eggs of certain animals membrane-formation is all that is required to induce a complete development of the unfertilised egg, e.g. in the starfish and certain annelids. For the eggs of other animals a second treatment is necessary, presumably to overcome some of the injurious effects of acid treatment. Thus the unfertilised eggs of the sea-urchin Strongylocentrotus purpuratus of the Californian coast begin to develop when membrane-formation has been induced by treatment with a fatty acid, e.g. butyric acid; but the development soon ceases and the eggs perish in the early stages of segmentation, or after the first nuclear division. But if we treat the same eggs, after membrane-formation, for from 35 to 55 minutes (at 15 deg C.) with sea-water the concentration (osmotic pressure) of which has been raised through the addition of a definite amount of some salt or sugar, the eggs will segment and develop normally, when transferred back to normal sea-water. If care is taken, practically all the eggs can be caused to develop into plutei, the majority of which may be perfectly normal and may live as long as larvae produced from eggs fertilised with sperm.

It is obvious that the sea-urchin egg is injured in the process of membrane-formation and that the subsequent treatment with a hypertonic solution only acts as a remedy. The nature of this injury became clear when it was discovered that all the agencies which cause haemolysis, i.e. the destruction of the red blood corpuscles, also cause membrane-formation in unfertilised eggs, e.g. fatty acids or ether, alcohols or chloroform, etc., or saponin, solanin, digitalin, bile salts and alkali. It thus happens that the phenomena of artificial parthenogenesis are linked together with the phenomena of haemolysis which at present play so important a role in the study of immunity. The difference between cytolysis (or haemolysis) and fertilisation seems to be this, that the latter is caused by a superficial or slight cytolysis of the egg, while if the cytolytic agencies have time to act on the whole egg the latter is completely destroyed. If we put unfertilised eggs of a sea-urchin into sea-water which contains a trace of saponin we notice that, after a few minutes, all the eggs form the typical membrane of fertilisation. If the eggs are then taken out of the saponin solution, freed from all traces of saponin by repeated washing in normal sea-water, and transferred to the hypertonic sea-water for from 35 to 55 minutes, they develop into larvae. If, however, they are left in the sea-water containing the saponin they undergo, a few minutes after membrane-formation, the disintegration known in pathology as CYTOLYSIS. Membrane-formation is, therefore, caused by a superficial or incomplete cytolysis. The writer believes that the subsequent treatment of the egg with hypertonic sea-water is needed only to overcome the destructive effects of this partial cytolysis. The full reasons for this belief cannot be given in a short essay.

Many pathologists assume that haemolysis or cytolysis is due to a liquefaction of certain fatty or fat-like compounds, the so-called lipoids, in the cell. If this view is correct, it would be necessary to ascribe the fertilisation of the egg to the same process.

The analogy between haemolysis and fertilisation throws, possibly, some light on a curious observation. It is well known that the blood corpuscles, as a rule, undergo cytolysis if injected into the blood of an animal which belongs to a different family. The writer found last year that the blood of mammals, e.g. the rabbit, pig, and cattle, causes the egg of Strongylocentrotus to form a typical fertilisation-membrane. If such eggs are afterwards treated for a short period with hypertonic sea-water they develop into normal larvae (plutei). Some substance contained in the blood causes, presumably, a superficial cytolysis of the egg and thus starts its development.

We can also cause the development of the sea-urchin egg without membrane-formation. The early experiments of the writer were done in this way and many experimenters still use such methods. It is probable that in this case the mechanism of fertilisation is essentially the same as in the case where the membrane-formation is brought about, with this difference only, that the cytolytic effect is less when no fertilisation-membrane is formed. This inference is corroborated by observations on the fertilisation of the sea-urchin egg with ox blood. It very frequently happens that not all of the eggs form membranes in this process. Those eggs which form membranes begin to develop, but perish if they are not treated with hypertonic sea-water. Some of the other eggs, however, which do not form membranes, develop directly into normal larvae without any treatment with hypertonic sea-water, provided they are exposed to the blood for only a few minutes. Presumably some blood enters the eggs and causes the cytolytic effects in a less degree than is necessary for membrane-formation, but in a sufficient degree to cause their development. The slightness of the cytolytic effect allows the egg to develop without treatment with hypertonic sea-water.

Since the entrance of the spermatozoon causes that degree of cytolysis which leads to membrane-formation, it is probable that, in addition to the cytolytic or membrane-forming substance (presumably a higher fatty acid), it carries another substance into the egg which counteracts the deleterious cytolytic effects underlying membrane-formation.

The question may be raised whether the larvae produced by artificial parthenogenesis can reach the mature stage. This question may be answered in the affirmative, since Delage has succeeded in raising several parthenogenetic sea-urchin larvae beyond the metamorphosis into the adult stage and since in all the experiments made by the writer the parthenogenetic plutei lived as long as the plutei produced from fertilised eggs.

(c). ON THE PRODUCTION OF TWINS FROM ONE EGG THROUGH A CHANGE IN THE CHEMICAL CONSTITUTION OF THE SEA-WATER.

The reader is probably familiar with the fact that there exist two different types of human twins. In the one type the twins differ as much as two children of the same parents born at different periods; they may or may not have the same sex. In the second type the twins have invariably the same sex and resemble each other most closely. Twins of the latter type are produced from the same egg, while twins of the former type are produced from two different eggs.

The experiments of Driesch and others have taught us that twins originate from one egg in this manner, namely, that the first two cells into which the egg divides after fertilisation become separated from each other. This separation can be brought about by a change in the chemical constitution of the sea-water. Herbst observed that if the fertilised eggs of the sea-urchin are put into sea-water which is freed from calcium, the cells into which the egg divides have a tendency to fall apart. Driesch afterwards noticed that eggs of the sea-urchin treated with sea-water which is free from lime have a tendency to give rise to twins. The writer has recently found that twins can be produced not only by the absence of lime, but also through the absence of sodium or of potassium; in other words, through the absence of one or two of the three important metals in the sea-water. There is, however, a second condition, namely, that the solution used for the production of twins must have a neutral or at least not an alkaline reaction.

The procedure for the production of twins in the sea-urchin egg consists simply in this:--the eggs are fertilised as usual in normal sea-water and then, after repeated washing in a neutral solution of sodium chloride (of the concentration of the sea-water), are placed in a neutral mixture of potassium chloride and calcium chloride, or of sodium chloride and potassium chloride, or of sodium chloride and calcium chloride, or of sodium chloride and magnesium chloride. The eggs must remain in this solution until half an hour or an hour after they have reached the two-cell stage. They are then transferred into normal sea-water and allowed to develop. From 50 to 90 per cent of the eggs of Strongylocentrotus purpuratus treated in this manner may develop into twins. These twins may remain separate or grow partially together and form double monsters, or heal together so completely that only slight or even no imperfections indicate that the individual started its career as a pair of twins. It is also possible to control the tendency of such twins to grow together by a change in the constitution of the sea-water. If we use as a twin-producing solution a mixture of sodium, magnesium and potassium chlorides (in the proportion in which these salts exist in the sea-water) the tendency of the twins to grow together is much more pronounced than if we use simply a mixture of sodium chloride and magnesium chloride.

The mechanism of the origin of twins, as the result of altering the composition of the sea-water, is revealed by observation of the first segmentation of the egg in these solutions. This cell-division is modified in a way which leads to a separation of the first two cells. If the egg is afterwards transferred back into normal sea-water, each of these two cells develops into an independent embryo. Since normal sea-water contains all three metals, sodium, calcium, and potassium, and since it has besides an alkaline reaction, we perceive the reason why twins are not normally produced from one egg. These experiments suggest the possibility of a chemical cause for the origin of twins from one egg or of double monstrosities in mammals. If, for some reason, the liquids which surround the human egg a short time before and after the first cell-division are slightly acid, and at the same time lacking in one of the three important metals, the conditions for the separation of the first two cells and the formation of identical twins are provided.

In conclusion it may be pointed out that the reverse result, namely, the fusion of normally double organs, can also be brought about experimentally through a change in the chemical constitution of the sea-water. Stockard succeeded in causing the eyes of fish embryos (Fundulus heteroclitus) to fuse into a single cyclopean eye through the addition of magnesium chloride to the sea-water. When he added about 6 grams of magnesium chloride to 100 cubic centimetres of sea-water and placed the fertilised eggs in the mixture, about 50 per cent of the eggs gave rise to one-eyed embryos. "When the embryos were studied the one-eyed condition was found to result from the union or fusion of the 'anlagen' of the two eyes. Cases were observed which showed various degrees in this fusion; it appeared as though the optic vessels were formed too far forward and ventral, so that their antero-ventro-median surfaces fused. This produces one large optic cup, which in all cases gives more or less evidence of its double nature." (Stockard, "Archiv f. Entwickelungsmechanik", Vol. 23, page 249, 1907.)

We have confined ourselves to a discussion of rather simple effects of the change in the constitution of the sea-water upon development. It is a priori obvious, however, that an unlimited number of pathological variations might be produced by a variation in the concentration and constitution of the sea-water, and experience confirms this statement. As an example we may mention the abnormalities observed by Herbst in the development of sea-urchins through the addition of lithium to sea-water. It is, however, as yet impossible to connect in a rational way the effects produced in this and similar cases with the cause which produced them; and it is also impossible to define in a simple way the character of the change produced.

III. THE INFLUENCE OF TEMPERATURE.

(a) THE INFLUENCE OF TEMPERATURE UPON THE DENSITY OF PELAGIC ORGANISMS AND THE DURATION OF LIFE.

It has often been noticed by explorers who have had a chance to compare the faunas in different climates that in polar seas such species as thrive at all in those regions occur, as a rule, in much greater density than they do in the moderate or warmer regions of the ocean. This refers to those members of the fauna which live at or near the surface, since they alone lend themselves to a statistical comparison. In his account of the Valdivia expedition, Chun (Chun, "Aus den Tiefen des Weltmeeres", page 225, Jena, 1903.) calls especial attention to this quantitative difference in the surface fauna and flora of different regions. "In the icy water of the Antarctic, the temperature of which is below 0 deg C., we find an astonishingly rich animal and plant life. The same condition with which we are familiar in the Arctic seas is repeated here, namely, that the quantity of plankton material exceeds that of the temperate and warm seas." And again, in regard to the pelagic fauna in the region of the Kerguelen Islands, he states: "The ocean is alive with transparent jelly fish, Ctenophores (Bolina and Callianira) and of Siphonophore colonies of the genus Agalma."

The paradoxical character of this general observation lies in the fact that a low temperature retards development, and hence should be expected to have the opposite effect from that mentioned by Chun. Recent investigations have led to the result that life-phenomena are affected by temperature in the same sense as the velocity of chemical reactions. In the case of the latter van't Hoff had shown that a decrease in temperature by 10 degrees reduces their velocity to one half or less, and the same has been found for the influence of temperature on the velocity of physiological processes. Thus Snyder and T.B. Robertson found that the rate of heartbeat in the tortoise and in Daphnia is reduced to about one-half if the temperature is lowered 10 deg C., and Maxwell, Keith Lucas, and Snyder found the same influence of temperature for the rate with which an impulse travels in the nerve. Peter observed that the rate of development in a sea-urchin's egg is reduced to less than one-half if the temperature (within certain limits) is reduced by 10 degrees. The same effect of temperature upon the rate of development holds for the egg of the frog, as Cohen and Peter calculated from the experiments of O. Hertwig. The writer found the same temperature-coefficient for the rate of maturation of the egg of a mollusc (Lottia).

All these facts prove that the velocity of development of animal life in Arctic regions, where the temperature is near the freezing point of water, must be from two to three times smaller than in regions where the temperature of the ocean is about 10 deg C. and from four to nine times smaller than in seas the temperature of which is about 20 deg C. It is, therefore, exactly the reverse of what we should expect when authors state that the density of organisms at or near the surface of the ocean in polar regions is greater than in more temperate regions.

The writer believes that this paradox finds its explanation in experiments which he has recently made on the influence of temperature on the duration of life of cold-blooded marine animals. The experiments were made on the fertilised and unfertilised eggs of the sea-urchin, and yielded the result that for the lowering of temperature by 1 deg C. the duration of life was about doubled. Lowering the temperature by 10 degrees therefore prolongs the life of the organism 2 to the power 10, i.e. over a thousand times, and a lowering by 20 degrees prolongs it about one million times. Since this prolongation of life is far in excess of the retardation of development through a lowering of temperature, it is obvious that, in spite of the retardation of development in Arctic seas, animal life must be denser there than in temperate or tropical seas. The excessive increase of the duration of life at the poles will necessitate the simultaneous existence of more successive generations of the same species in these regions than in the temperate or tropical regions.

The writer is inclined to believe that these results have some bearing upon a problem which plays an important role in theories of evolution, namely, the cause of natural death. It has been stated that the processes of differentiation and development lead also to the natural death of the individual. If we express this in chemical terms it means that the chemical processes which underlie development also determine natural death. Physical chemistry has taught us to identify two chemical processes even if only certain of their features are known. One of these means of identification is the temperature coefficient. When two chemical processes are identical, their velocity must be reduced by the same amount if the temperature is lowered to the same extent. The temperature coefficient for the duration of life of cold-blooded organisms seems, however, to differ enormously from the temperature coefficient for their rate of development. For a difference in temperature of 10 deg C. the duration of life is altered five hundred times as much as the rate of development; and, for a change of 20 deg C., it is altered more than a hundred thousand times as much. From this we may conclude that, at least for the sea-urchin eggs and embryo, the chemical processes which determine natural death are certainly not identical with the processes which underlie their development. T.B. Robertson has also arrived at the conclusion, for quite different reasons, that the process of senile decay is essentially different from that of growth and development.

(b) CHANGES IN THE COLOUR OF BUTTERFLIES PRODUCED THROUGH THE INFLUENCE OF TEMPERATURE.

The experiments of Dorfmeister, Weismann, Merrifield, Standfuss, and Fischer, on seasonal dimorphism and the aberration of colour in butterflies have so often been discussed in biological literature that a short reference to them will suffice. By seasonal dimorphism is meant the fact that species may appear at different seasons of the year in a somewhat different form or colour. Vanessa prorsa is the summer form, Vanessa levana the winter form of the same species. By keeping the pupae of Vanessa prorsa several weeks at a temperature of from 0 deg to 1 deg Weismann succeeded in obtaining from the summer chrysalids specimens which resembled the winter variety, Vanessa levana.

If we wish to get a clear understanding of the causes of variation in the colour and pattern of butterflies, we must direct our attention to the experiments of Fischer, who worked with more extreme temperatures than his predecessors, and found that almost identical aberrations of colour could be produced by both extremely high and extremely low temperatures. This can be clearly seen from the following tabulated results of his observations. At the head of each column the reader finds the temperature to which Fischer submitted the pupae, and in the vertical column below are found the varieties that were produced. In the vertical column A are given the normal forms:

(Temperatures in deg C.)

0 to -20 0 to +10 A. +35 to +37 +36 to +41 +42 to +46 (Normal forms)

ichnusoides polaris urticae ichnusa polaris ichnusoides (nigrita) (nigrita)

antigone fischeri io - fischeri antigone (iokaste) (iokaste)

testudo dixeyi polychloros erythromelas dixeyi testudo

hygiaea artemis antiopa epione artemis hygiaea

elymi wiskotti cardui - wiskotti elymi

klymene merrifieldi atalanta - merrifieldi klymene

weismanni porima prorsa - porima weismanni

The reader will notice that the aberrations produced at a very low temperature (from 0 to -20 deg C.) are absolutely identical with the aberrations produced by exposing the pupae to extremely high temperatures (42 to 46 deg C.). Moreover the aberrations produced by a moderately low temperature (from 0 to 10 deg C.) are identical with the aberrations produced by a moderately high temperature (36 to 41 deg C.)

From these observations Fischer concludes that it is erroneous to speak of a specific effect of high and of low temperatures, but that there must be a common cause for the aberration found at the high as well as at the low temperature limits. This cause he seems to find in the inhibiting effects of extreme temperatures upon development.

If we try to analyse such results as Fischer's from a physico-chemical point of view, we must realise that what we call life consists of a series of chemical reactions, which are connected in a catenary way; inasmuch as one reaction or group of reactions (a) (e.g. hydrolyses) causes or furnishes the material for a second reaction or group of reactions (b) (e.g. oxydations). We know that the temperature coefficient for physiological processes varies slightly at various parts of the scale; as a rule it is higher near 0 and lower near 30 deg. But we know also that the temperature coefficients do not vary equally from the various physiological processes. It is, therefore, to be expected that the temperature coefficients for the group of reactions of the type (a) will not be identical through the whole scale with the temperature coefficients for the reactions of the type (b). If therefore a certain substance is formed at the normal temperature of the animal in such quantities as are needed for the catenary reaction (b), it is not to be expected that this same perfect balance will be maintained for extremely high or extremely low temperatures; it is more probable that one group of reactions will exceed the other and thus produce aberrant chemical effects, which may underlie the colour aberrations observed by Fischer and other experimenters.

It is important to notice that Fischer was also able to produce aberrations through the application of narcotics. Wolfgang Ostwald has produced experimentally, through variation of temperature, dimorphism of form in Daphnia. Lack of space precludes an account of these important experiments, as of so many others.

IV. THE EFFECTS OF LIGHT.

At the present day nobody seriously questions the statement that the action of light upon organisms is primarily one of a chemical character. While this chemical action is of the utmost importance for organisms, the nutrition of which depends upon the action of chlorophyll, it becomes of less importance for organisms devoid of chlorophyll. Nevertheless, we find animals in which the formation of organs by regeneration is not possible unless they are exposed to light. An observation made by the writer on the regeneration of polyps in a hydroid, Eudendrium racemosum, at Woods Hole, may be mentioned as an instance of this. If the stem of this hydroid, which is usually covered with polyps, is put into an aquarium the polyps soon fall off. If the stems are kept in an aquarium where light strikes them during the day, a regeneration of numerous polyps takes place in a few days. If, however, the stems of Eudendrium are kept permanently in the dark, no polyps are formed even after an interval of some weeks; but they are formed in a few days after the same stems have been transferred from the dark to the light. Diffused daylight suffices for this effect. Goldfarb, who repeated these experiments, states that an exposure of comparatively short duration is sufficient for this effect, it is possible that the light favours the formation of substances which are a prerequisite for the origin of polyps and their growth.

Of much greater significance than this observation are the facts which show that a large number of animals assume, to some extent, the colour of the ground on which they are placed. Pouchet found through experiments upon crustaceans and fish that this influence of the ground on the colour of animals is produced through the medium of the eyes. If the eyes are removed or the animals made blind in another way these phenomena cease. The second general fact found by Pouchet was that the variation in the colour of the animal is brought about through an action of the nerves on the pigment-cells of the skin; the nerve-action being induced through the agency of the eye.

The mechanism and the conditions for the change in colouration were made clear through the beautiful investigations of Keeble and Gamble, on the colour-change in crustaceans. According to these authors the pigment-cells can, as a rule, be considered as consisting of a central body from which a system of more or less complicated ramifications or processes spreads out in all directions. As a rule, the centre of the cell contains one or more different pigments which under the influence of nerves can spread out separately or together into the ramifications. These phenomena of spreading and retraction of the pigments into or from the ramifications of the pigment-cells form on the whole the basis for the colour changes under the influence of environment. Thus Keeble and Gamble observed that Macromysis flexuosa appears transparent and colourless or grey on sandy ground. On a dark ground their colour becomes darker. These animals have two pigments in their chromatophores, a brown pigment and a whitish or yellow pigment; the former is much more plentiful than the latter. When the animal appears transparent all the pigment is contained in the centre of the cells, while the ramifications are free from pigment. When the animal appears brown both pigments are spread out into the ramifications. In the condition of maximal spreading the animals appear black.

This is a comparatively simple case. Much more complicated conditions were found by Keeble and Gamble in other crustaceans, e.g. in Hippolyte cranchii, but the influence of the surroundings upon the colouration of this form was also satisfactorily analysed by these authors.

While many animals show transitory changes in colour under the influence of their surroundings, in a few cases permanent changes can be produced. The best examples of this are those which were observed by Poulton in the chrysalids of various butterflies, especially the small tortoise-shell. These experiments are so well known that a short reference to them will suffice. Poulton (Poulton, E.B., "Colours of Animals" (The International Scientific Series), London, 1890, page 121.) found that in gilt or white surroundings the pupae became light coloured and there was often an immense development of the golden spots, "so that in many cases the whole surface of the pupae glittered with an apparent metallic lustre. So remarkable was the appearance that a physicist to whom I showed the chrysalids, suggested that I had played a trick and had covered them with goldleaf." When black surroundings were used "the pupae were as a rule extremely dark, with only the smallest trace, and often no trace at all, of the golden spots which are so conspicuous in the lighter form." The susceptibility of the animal to this influence of its surroundings was found to be greatest during a definite period when the caterpillar undergoes the metamorphosis into the chrysalis stage. As far as the writer is aware, no physico-chemical explanation, except possibly Wiener's suggestion of colour-photography by mechanical colour adaptation, has ever been offered for the results of the type of those observed by Poulton.

V. EFFECTS OF GRAVITATION.

(a) EXPERIMENTS ON THE EGG OF THE FROG.

Gravitation can only indirectly affect life-phenomena; namely, when we have in a cell two different non-miscible liquids (or a liquid and a solid) of different specific gravity, so that a change in the position of the cell or the organ may give results which can be traced to a change in the position of the two substances. This is very nicely illustrated by the frog's egg, which has two layers of very viscous protoplasm one of which is black and one white. The dark one occupies normally the upper position in the egg and may therefore be assumed to possess a smaller specific gravity than the white substance. When the egg is turned with the white pole upwards a tendency of the white protoplasm to flow down again manifests itself. It is, however, possible to prevent or retard this rotation of the highly viscous protoplasm, by compressing the eggs between horizontal glass plates. Such compression experiments may lead to rather interesting results, as O. Schultze first pointed out. Pflueger had already shown that the first plane of division in a fertilised frog's egg is vertical and Roux established the fact that the first plane of division is identical with the plane of symmetry of the later embryo. Schultze found that if the frog's egg is turned upside down at the time of its first division and kept in this abnormal position, through compression between two glass plates for about 20 hours, a small number of eggs may give rise to twins. It is possible, in this case, that the tendency of the black part of the egg to rotate upwards along the surface of the egg leads to a separation of its first cells, such a separation leading to the formation of twins.

T.H. Morgan made an interesting additional observation. He destroyed one half of the egg after the first segmentation and found that the half which remained alive gave rise to only one half of an embryo, thus confirming an older observation of Roux. When, however, Morgan put the egg upside down after the destruction of one of the first two cells, and compressed the eggs between two glass plates, the surviving half of the egg gave rise to a perfect embryo of half size (and not to a half embryo of normal size as before.) Obviously in this case the tendency of the protoplasm to flow back to its normal position was partially successful and led to a partial or complete separation of the living from the dead half; whereby the former was enabled to form a whole embryo, which, of course, possessed only half the size of an embryo originating from a whole egg.

(b) EXPERIMENTS ON HYDROIDS.

A striking influence of gravitation can be observed in a hydroid, Antennularia antennina, from the bay of Naples. This hydroid consists of a long straight main stem which grows vertically upwards and which has at regular intervals very fine and short bristle-like lateral branches, on the upper side of which the polyps grow. The main stem is negatively geotropic, i.e. its apex continues to grow vertically upwards when we put it obliquely into the aquarium, while the roots grow vertically downwards. The writer observed that when the stem is put horizontally into the water the short lateral branches on the lower side give rise to an altogether different kind of organ, namely, to roots, and these roots grow indefinitely in length and attach themselves to solid bodies; while if the stem had remained in its normal position no further growth would have occurred in the lateral branches. From the upper side of the horizontal stem new stems grow out, mostly directly from the original stem, occasionally also from the short lateral branches. It is thus possible to force upon this hydroid an arrangement of organs which is altogether different from the hereditary arrangement. The writer had called the change in the hereditary arrangement of organs or the transformation of organs by external forces HETEROMORPHOSIS. We cannot now go any further into this subject, which should, however, prove of interest in relation to the problem of heredity.

If it is correct to apply inferences drawn from the observation on the frog's egg to the behaviour of Antennularia, one might conclude that the cells of Antennularia also contain non-miscible substances of different specific gravity, and that wherever the specifically lighter substance comes in contact with the sea-water (or gets near the surface of the cell) the growth of a stem is favoured; while contact with the sea-water of the specifically heavier of the substances, will favour the formation of roots.

VI. THE EXPERIMENTAL CONTROL OF ANIMAL INSTINCTS.

(a) EXPERIMENTS ON THE MECHANISM OF HELIOTROPIC REACTIONS IN ANIMALS.

Since the instinctive reactions of animals are as hereditary as their morphological character, a discussion of experiments on the physico-chemical character of the instinctive reactions of animals should not be entirely omitted from this sketch. It is obvious that such experiments must begin with the simplest type of instincts, if they are expected to lead to any results; and it is also obvious that only such animals must be selected for this purpose, the reactions of which are not complicated by associative memory, or, as it may preferably be termed, associative hysteresis.

The simplest type of instincts is represented by the purposeful motions of animals to or from a source of energy, e.g. light; and it is with some of these that we intend to deal here. When we expose winged aphides (after they have flown away from the plant), or young caterpillars of Porthesia chrysorrhoea (when they are aroused from their winter sleep) or marine or freshwater copepods and many other animals, to diffused daylight falling in from a window, we notice a tendency among these animals to move towards the source of light. If the animals are naturally sensitive, or if they are rendered sensitive through the agencies which we shall mention later, and if the light is strong enough, they move towards the source of light in as straight a line as the imperfections and peculiarities of their locomotor apparatus will permit. It is also obvious that we are here dealing with a forced reaction in which the animals have no more choice in the direction of their motion than have the iron filings in their arrangement in a magnetic field. This can be proved very nicely in the case of starving caterpillars of Porthesia. The writer put such caterpillars into a glass tube the axis of which was at right angles to the plane of the window: the caterpillars went to the window side of the tube and remained there, even if leaves of their food-plant were put into the tube directly behind them. Under such conditions the animals actually died from starvation, the light preventing them from turning to the food, which they eagerly ate when the light allowed them to do so. One cannot say that these animals, which we call positively helioptropic, are attracted by the light, since it can be shown that they go towards the source of the light even if in so doing they move from places of a higher to places of a lower degree of illumination.

The writer has advanced the following theory of these instinctive reactions. Animals of the type of those mentioned are automatically orientated by the light in such a way that symmetrical elements of their retina (or skin) are struck by the rays of light at the same angle. In this case the intensity of light is the same for both retinae or symmetrical parts of the skin.

This automatic orientation is determined by two factors, first a peculiar photo-sensitiveness of the retina (or skin), and second a peculiar nervous connection between the retina and the muscular apparatus. In symmetrically built heliotropic animals in which the symmetrical muscles participate equally in locomotion, the symmetrical muscles work with equal energy as long as the photo-chemical processes in both eyes are identical. If, however, one eye is struck by stronger light than the other, the symmetrical muscles will work unequally and in positively heliotropic animals those muscles will work with greater energy which bring the plane of symmetry back into the direction of the rays of light and the head towards the source of light. As soon as both eyes are struck by the rays of light at the same angle, there is no more reason for the animal to deviate from this direction and it will move in a straight line. All this holds good on the supposition that the animals are exposed to only one source of light and are very sensitive to light.

Additional proof for the correctness of this theory was furnished through the experiments of G.H. Parker and S.J. Holmes. The former worked on a butterfly, Vanessa antiope, the latter on other arthropods. All the animals were in a marked degree positively heliotropic. These authors found that if one cornea is blackened in such an animal, it moves continually in a circle when it is exposed to a source of light, and in these motions the eye which is not covered with paint is directed towards the centre of the circle. The animal behaves, therefore, as if the darkened eye were in the shade.

(b) THE PRODUCTION OF POSITIVE HELIOTROPISM BY ACIDS AND OTHER MEANS AND THE PERIODIC DEPTH-MIGRATIONS OF PELAGIC ANIMALS.

When we observe a dense mass of copepods collected from a freshwater pond, we notice that some have a tendency to go to the light while others go in the opposite direction and many, if not the majority, are indifferent to light. It is an easy matter to make the negatively heliotropic or the indifferent copepods almost instantly positively heliotropic by adding a small but definite amount of carbon-dioxide in the form of carbonated water to the water in which the animals are contained. If the animals are contained in 50 cubic centimetres of water it suffices to add from three to six cubic centimetres of carbonated water to make all the copepods energetically positively heliotropic. This heliotropism lasts about half an hour (probably until all the carbon-dioxide has again diffused into the air.) Similar results may be obtained with any other acid.

The same experiments may be made with another freshwater crustacean, namely Daphnia, with this difference, however, that it is as a rule necessary to lower the temperature of the water also. If the water containing the Daphniae is cooled and at the same time carbon-dioxide added, the animals which were before indifferent to light now become most strikingly positively heliotropic. Marine copepods can be made positively heliotropic by the lowering of the temperature alone, or by a sudden increase in the concentration of the sea-water.

These data have a bearing upon the depth-migrations of pelagic animals, as was pointed out years ago by Theo. T. Groom and the writer. It is well known that many animals living near the surface of the ocean or freshwater lakes, have a tendency to migrate upwards towards evening and downwards in the morning and during the day. These periodic motions are determined to a large extent, if not exclusively, by the heliotropism of these animals. Since the consumption of carbon-dioxide by the green plants ceases towards evening, the tension of this gas in the water must rise and this must have the effect of inducing positive heliotropism or increasing its intensity. At the same time the temperature of the water near the surface is lowered and this also increases the positive heliotropism in the organisms.

The faint light from the sky is sufficient to cause animals which are in a high degree positively heliotropic to move vertically upwards towards the light, as experiments with such pelagic animals, e.g. copepods, have shown. When, in the morning, the absorption of carbon-dioxide by the green algae begins again and the temperature of the water rises, the animals lose their positive heliotropism, and slowly sink down or become negatively heliotropic and migrate actively downwards.

These experiments have also a bearing upon the problem of the inheritance of instincts. The character which is transmitted in this case is not the tendency to migrate periodically upwards and downwards, but the positive heliotropism. The tendency to migrate is the outcome of the fact that periodically varying external conditions induce a periodic change in the sense and intensity of the heliotropism of these animals. It is of course immaterial for the result, whether the carbon-dioxide or any other acid diffuse into the animal from the outside or whether they are produced inside in the tissue cells of the animals. Davenport and Cannon found that Daphniae, which at the beginning of the experiment, react sluggishly to light react much more quickly after they have been made to go to the light a few times. The writer is inclined to attribute this result to the effect of acids, e.g. carbon-dioxide, produced in the animals themselves in consequence of their motion. A similar effect of the acids was shown by A.D. Waller in the case of the response of nerve to stimuli.

The writer observed many years ago that winged male and female ants are positively helioptropic and that their heliotropic sensitiveness increases and reaches its maximum towards the period of nuptial flight. Since the workers show no heliotropism it looks as if an internal secretion from the sexual glands were the cause of their heliotropic sensitiveness. V. Kellogg has observed that bees also become intensely positively heliotropic at the period of their wedding flight, in fact so much so that by letting light fall into the observation hive from above, the bees are prevented from leaving the hive through the exit at the lower end.

We notice also the reverse phenomenon, namely, that chemical changes produced in the animal destroy its heliotropism. The caterpillars of Porthesia chrysorrhoea are very strongly positively heliotropic when they are first aroused from their winter sleep. This heliotropic sensitiveness lasts only as long as they are not fed. If they are kept permanently without food they remain permanently positively heliotropic until they die from starvation. It is to be inferred that as soon as these animals take up food, a substance or substances are formed in their bodies which diminish or annihilate their heliotropic sensitiveness.

The heliotropism of animals is identical with the heliotropism of plants. The writer has shown that the experiments on the effect of acids on the heliotropism of copepods can be repeated with the same result in Volvox. It is therefore erroneous to try to explain these heliotropic reactions of animals on the basis of peculiarities (e.g. vision) which are not found in plants.

We may briefly discuss the question of the transmission through the sex cells of such instincts as are based upon heliotropism. This problem reduces itself simply to that of the method whereby the gametes transmit heliotropism to the larvae or to the adult. The writer has expressed the idea that all that is necessary for this transmission is the presence in the eyes (or in the skin) of the animal of a photo-sensitive substance. For the transmission of this the gametes need not contain anything more than a catalyser or ferment for the synthesis of the photo-sensitive substance in the body of the animal. What has been said in regard to animal heliotropism might, if space permitted, be extended, mutatis mutandis, to geotropism and stereotropism.

(c) THE TROPIC REACTIONS OF CERTAIN TISSUE-CELLS AND THE MORPHOGENETIC EFFECTS OF THESE REACTIONS.

Since plant-cells show heliotropic reactions identical with those of animals, it is not surprising that certain tissue-cells also show reactions which belong to the class of tropisms. These reactions of tissue-cells are of special interest by reason of their bearing upon the inheritance of morphological characters. An example of this is found in the tiger-like marking of the yolk-sac of the embryo of Fundulus and in the marking of the young fish itself. The writer found that the former is entirely, and the latter at least in part, due to the creeping of the chromatophores upon the blood-vessels. The chromatophores are at first scattered irregularly over the yolk-sac and show their characteristic ramifications. There is at that time no definite relation between blood-vessels and chromatophores. As soon as a ramification of a chromatophore comes in contact with a blood-vessel the whole mass of the chromatophore creeps gradually on the blood-vessel and forms a complete sheath around the vessel, until finally all the chromatophores form a sheath around the vessels and no more pigment cells are found in the meshes between the vessels. Nobody who has not actually watched the process of the creeping of the chromatophores upon the blood-vessels would anticipate that the tiger-like colouration of the yolk-sac in the later stages of the development was brought about in this way. Similar facts can be observed in regard to the first marking of the embryo itself. The writer is inclined to believe that we are here dealing with a case of chemotropism, and that the oxygen of the blood may be the cause of the spreading of the chromatophores around the blood-vessels. Certain observations seem to indicate the possibility that in the adult the chromatophores have, in some forms at least, a more rigid structure and are prevented from acting in the way indicated. It seems to the writer that such observations as those made on Fundulus might simplify the problem of the hereditary transmission of certain markings.

Driesch has found that a tropism underlies the arrangement of the skeleton in the pluteus larvae of the sea-urchin. The position of this skeleton is predetermined by the arrangement of the mesenchyme cells, and Driesch has shown that these cells migrate actively to the place of their destination, possibly led there under the influence of certain chemical substances. When Driesch scattered these cells mechanically before their migration, they nevertheless reached their destination.

In the developing eggs of insects the nuclei, together with some cytoplasm, migrate to the periphery of the egg. Herbst pointed out that this might be a case of chemotropism, caused by the oxygen surrounding the egg. The writer has expressed the opinion that the formation of the blastula may be caused generally by a tropic reaction of the blastomeres, the latter being forced by an outside influence to creep to the surface of the egg.

These examples may suffice to indicate that the arrangement of definite groups of cells and the morphological effects resulting therefrom may be determined by forces lying outside the cells. Since these forces are ubiquitous and constant it appears as if we were dealing exclusively with the influence of a gamete; while in reality all that it is necessary for the gamete to transmit is a certain form of irritability.

(d) FACTORS WHICH DETERMINE PLACE AND TIME FOR THE DEPOSITION OF EGGS.

For the preservation of species the instinct of animals to lay their eggs in places in which the young larvae find their food and can develop is of paramount importance. A simple example of this instinct is the fact that the common fly lays its eggs on putrid material which serves as food for the young larvae. When a piece of meat and of fat of the same animal are placed side by side, the fly will deposit its eggs upon the meat on which the larvae can grow, and not upon the fat, on which they would starve. Here we are dealing with the effect of a volatile nitrogenous substance which reflexly causes the peristaltic motions for the laying of the egg in the female fly.

Kammerer has investigated the conditions for the laying of eggs in two forms of salamanders, e.g. Salamandra atra and S. maculosa. In both forms the eggs are fertilised in the body and begin to develop in the uterus. Since there is room only for a few larvae in the uterus, a large number of eggs perish and this number is the greater the longer the period of gestation. It thus happens that when the animals retain their eggs a long time, very few young ones are born; and these are in a rather advanced stage of development, owing to the long time which elapsed since they were fertilised. When the animal lays its eggs comparatively soon after copulation, many eggs (from 12 to 72) are produced and the larvae are of course in an early stage of development. In the early stage the larvae possess gills and can therefore live in water, while in later stages they have no gills and breathe through their lungs. Kammerer showed that both forms of Salamandra can be induced to lay their eggs early or late, according to the physical conditions surrounding them. If they are kept in water or in proximity to water and in a moist atmosphere they have a tendency to lay their eggs earlier and a comparatively high temperature enhances the tendency to shorten the period of gestation. If the salamanders are kept in comparative dryness they show a tendency to lay their eggs rather late and a low temperature enhances this tendency.

Since Salamandra atra is found in rather dry alpine regions with a relatively low temperature and Salamandra maculosa in lower regions with plenty of water and a higher temperature, the fact that S. atra bears young which are already developed and beyond the stage of aquatic life, while S. maculosa bears young ones in an earlier stage, has been termed adaptation. Kammerer's experiments, however, show that we are dealing with the direct effects of definite outside forces. While we may speak of adaptation when all or some of the variables which determine a reaction are unknown, it is obviously in the interest of further scientific progress to connect cause and effect directly whenever our knowledge allows us to do so.

VII. CONCLUDING REMARKS.

The discovery of De Vries, that new species may arise by mutation and the wide if not universal applicability of Mendel's Law to phenomena of heredity, as shown especially by Bateson and his pupils, must, for the time being, if not permanently, serve as a basis for theories of evolution. These discoveries place before the experimental biologist the definite task of producing mutations by physico-chemical means. It is true that certain authors claim to have succeeded in this, but the writer wishes to apologise to these authors for his inability to convince himself of the validity of their claims at the present moment. He thinks that only continued breeding of these apparent mutants through several generations can afford convincing evidence that we are here dealing with mutants rather than with merely pathological variations.

What was said in regard to the production of new species by physico-chemical means may be repeated with still more justification in regard to the second problem of transformation, namely the making of living from inanimate matter. The purely morphological imitations of bacteria or cells which physicists have now and then proclaimed as artificially produced living beings; or the plays on words by which, e.g. the regeneration of broken crystals and the regeneration of lost limbs by a crustacean were declared identical, will not appeal to the biologist. We know that growth and development in animals and plants are determined by definite although complicated series of catenary chemical reactions, which result in the synthesis of a DEFINITE compound or group of compounds, namely, NUCLEINS.

The nucleins have the peculiarity of acting as ferments or enzymes for their own synthesis. Thus a given type of nucleus will continue to synthesise other nuclein of its own kind. This determines the continuity of a species; since each species has, probably, its own specific nuclein or nuclear material. But it also shows us that whoever claims to have succeeded in making living matter from inanimate will have to prove that he has succeeded in producing nuclein material which acts as a ferment for its own synthesis and thus reproduces itself. Nobody has thus far succeeded in this, although nothing warrants us in taking it for granted that this task is beyond the power of science.

XV. THE VALUE OF COLOUR IN THE STRUGGLE FOR LIFE. By E.B. Poulton.

Hope Professor of Zoology in the University of Oxford.

INTRODUCTION.

The following pages have been written almost entirely from the historical stand-point. Their principal object has been to give some account of the impressions produced on the mind of Darwin and his great compeer Wallace by various difficult problems suggested by the colours of living nature. In order to render the brief summary of Darwin's thoughts and opinions on the subject in any way complete, it was found necessary to say again much that has often been said before. No attempt has been made to display as a whole the vast contribution of Wallace; but certain of its features are incidentally revealed in passages quoted from Darwin's letters. It is assumed that the reader is familiar with the well-known theories of Protective Resemblance, Warning Colours, and Mimicry both Batesian and Mullerian. It would have been superfluous to explain these on the present occasion; for a far more detailed account than could have been attempted in these pages has recently appeared. (Poulton, "Essays on Evolution" Oxford, 1908, pages 293-382.) Among the older records I have made a point of bringing together the principal observations scattered through the note-books and collections of W.J. Burchell. These have never hitherto found a place in any memoir dealing with the significance of the colours of animals.

INCIDENTAL COLOURS.

Darwin fully recognised that the colours of living beings are not necessarily of value as colours, but that they may be an incidental result of chemical or physical structure. Thus he wrote to T. Meehan, Oct. 9, 1874: "I am glad that you are attending to the colours of dioecious flowers; but it is well to remember that their colours may be as unimportant to them as those of a gall, or, indeed, as the colour of an amethyst or ruby is to these gems." ("More Letters of Charles Darwin", Vol. I. pages 354, 355. See also the admirable account of incidental colours in "Descent of Man" (2nd edition), 1874, pages 261, 262.)

Incidental colours remain as available assets of the organism ready to be turned to account by natural selection. It is a probable speculation that all pigmentary colours were originally incidental; but now and for immense periods of time the visible tints of animals have been modified and arranged so as to assist in the struggle with other organisms or in courtship. The dominant colouring of plants, on the other hand, is an essential element in the paramount physiological activity of chlorophyll. In exceptional instances, however, the shapes and visible colours of plants may be modified in order to promote concealment.

TELEOLOGY AND ADAPTATION.

In the department of Biology which forms the subject of this essay, the adaptation of means to an end is probably more evident than in any other; and it is therefore of interest to compare, in a brief introductory section, the older with the newer teleological views.

The distinctive feature of Natural Selection as contrasted with other attempts to explain the process of Evolution is the part played by the struggle for existence. All naturalists in all ages must have known something of the operations of "Nature red in tooth and claw"; but it was left for this great theory to suggest that vast extermination is a necessary condition of progress, and even of maintaining the ground already gained.

Realising that fitness is the outcome of this fierce struggle, thus turned to account for the first time, we are sometimes led to associate the recognition of adaptation itself too exclusively with Natural Selection. Adaptation had been studied with the warmest enthusiasm nearly forty years before this great theory was given to the scientific world, and it is difficult now to realise the impetus which the works of Paley gave to the study of Natural History. That they did inspire the naturalists of the early part of the last century is clearly shown in the following passages.

In the year 1824 the Ashmolean Museum at Oxford was intrusted to the care of J.S. Duncan of New College. He was succeeded in this office by his brother, P.B. Duncan, of the same College, author of a History of the Museum, which shows very clearly the influence of Paley upon the study of nature, and the dominant position given to his teachings: "Happily at this time (1824) a taste for the study of natural history had been excited in the University by Dr Paley's very interesting work on Natural Theology, and the very popular lectures of Dr Kidd on Comparative Anatomy, and Dr Buckland on Geology." In the arrangement of the contents of the Museum the illustration of Paley's work was given the foremost place by J.S. Duncan: "The first division proposes to familiarize the eye to those relations of all natural objects which form the basis of argument in Dr Paley's Natural Theology; to induce a mental habit of associating the view of natural phenomena with the conviction that they are the media of Divine manifestation; and by such association to give proper dignity to every branch of natural science." (From "History and Arrangement of the Ashmolean Museum" by P.B. Duncan: see pages vi, vii of "A Catalogue of the Ashmolean Museum", Oxford, 1836.)

The great naturalist, W.J. Burchell, in his classical work shows the same recognition of adaptation in nature at a still earlier date. Upon the subject of collections he wrote ("Travels in the Interior of Southern Africa", London, Vol. I. 1822, page 505. The references to Burchell's observations in the present essay are adapted from the author's article in "Report of the British and South African Associations", 1905, Vol. III. pages 57-110.): "It must not be supposed that these charms (the pleasures of Nature) are produced by the mere discovery of new objects: it is the harmony with which they have been adapted by the Creator to each other, and to the situations in which they are found, which delights the observer in countries where Art has not yet introduced her discords." The remainder of the passage is so admirable that I venture to quote it: "To him who is satisfied with amassing collections of curious objects, simply for the pleasure of possessing them, such objects can afford, at best, but a childish gratification, faint and fleeting; while he who extends his view beyond the narrow field of nomenclature, beholds a boundless expanse, the exploring of which is worthy of the philosopher, and of the best talents of a reasonable being."

On September 14, 1811, Burchell was at Zand Valley (Vlei), or Sand Pool, a few miles south-west of the site of Prieska, on the Orange River. Here he found a Mesembryanthemum (M. turbiniforme, now M. truncatum) and also a "Gryllus" (Acridian), closely resembling the pebbles with which their locality was strewn. He says of both of these, "The intention of Nature, in these instances, seems to have been the same as when she gave to the Chameleon the power of accommodating its color, in a certain degree, to that of the object nearest to it, in order to compensate for the deficiency of its locomotive powers. By their form and colour, this insect may pass unobserved by those birds, which otherwise would soon extirpate a species so little able to elude its pursuers, and this juicy little Mesembryanthemum may generally escape the notice of cattle and wild animals." (Loc. cit. pages 310, 311. See Sir William Thiselton-Dyer "Morphological Notes", XI.; "Protective Adaptations", I.; "Annals of Botany", Vol. XX. page 124. In plates VII., VIII. and IX. accompanying this article the author represents the species observed by Burchell, together with others in which analogous adaptations exist. He writes: "Burchell was clearly on the track on which Darwin reached the goal. But the time had not come for emancipation from the old teleology. This, however, in no respect detracts from the merit or value of his work. For, as Huxley has pointed out ("Life and Letters of Thomas Henry Huxley", London, 1900, I. page 457), the facts of the old teleology are immediately transferable to Darwinism, which simply supplies them with a natural in place of a supernatural explanation.") Burchell here seems to miss, at least in part, the meaning of the relationship between the quiescence of the Acridian and its cryptic colouring. Quiescence is an essential element in the protective resemblance to a stone--probably even more indispensable than the details of the form and colouring. Although Burchell appears to overlook this point he fully recognised the community between protection by concealment and more aggressive modes of defence; for, in the passage of which a part is quoted above, he specially refers to some earlier remarks on page 226 of his Vol. I. We here find that even when the oxen were resting by the Juk rivier (Yoke river), on July 19, 1811, Burchell observed "Geranium spinosum, with a fleshy stem and large white flowers...; and a succulent species of Pelargonium... so defended by the old panicles, grown to hard woody thorns, that no cattle could browze upon it." He goes on to say, "In this arid country, where every juicy vegetable would soon be eaten up by the wild animals, the Great Creating Power, with all-provident wisdom, has given to such plants either an acrid or poisonous juice, or sharp thorns, to preserve the species from annihilation... " All these modes of defence, especially adapted to a desert environment, have since been generally recognised, and it is very interesting to place beside Burchell's statement the following passage from a letter written by Darwin, Aug. 7, 1868, to G.H. Lewes; "That Natural Selection would tend to produce the most formidable thorns will be admitted by every one who has observed the distribution in South America and Africa (vide Livingstone) of thorn-bearing plants, for they always appear where the bushes grow isolated and are exposed to the attacks of mammals. Even in England it has been noticed that all spine-bearing and sting-bearing plants are palatable to quadrupeds, when the thorns are crushed." ("More Letters", I. page 308.)

ADAPTATION AND NATURAL SELECTION.

I have preferred to show the influence of the older teleology upon Natural History by quotations from a single great and insufficiently appreciated naturalist. It might have been seen equally well in the pages of Kirby and Spence and those of many other writers. If the older naturalists who thought and spoke with Burchell of "the intention of Nature" and the adaptation of beings "to each other, and to the situations in which they are found," could have conceived the possibility of evolution, they must have been led, as Darwin was, by the same considerations to Natural Selection. This was impossible for them, because the philosophy which they followed contemplated the phenomena of adaptation as part of a static immutable system. Darwin, convinced that the system is dynamic and mutable, was prevented by these very phenomena from accepting anything short of the crowning interpretation offered by Natural Selection. ("I had always been much struck by such adaptations (e.g. woodpecker and tree-frog for climbing, seeds for dispersal), and until these could be explained it seemed to me almost useless to endeavour to prove by indirect evidence that species have been modified." "Autobiography" in "Life and Letters of Charles Darwin", Vol. I. page 82. The same thought is repeated again and again in Darwin's letters to his friends. It is forcibly urged in the Introduction to the "Origin" (1859), page 3.) And the birth of Darwin's unalterable conviction that adaptation is of dominant importance in the organic world,--a conviction confirmed and ever again confirmed by his experience as a naturalist--may probably be traced to the influence of the great theologian. Thus Darwin, speaking of his Undergraduate days, tells us in his "Autobiography" that the logic of Paley's "Evidences of Christianity" and "Moral Philosophy" gave him as much delight as did Euclid.

"The careful study of these works, without attempting to learn any part by rote, was the only part of the academical course which, as I then felt and as I still believe, was of the least use to me in the education of my mind. I did not at that time trouble myself about Paley's premises; and taking these on trust, I was charmed and convinced by the long line of argumentation." ("Life and Letters", I. page 47.)

When Darwin came to write the "Origin" he quoted in relation to Natural Selection one of Paley's conclusions. "No organ will be formed, as Paley has remarked, for the purpose of causing pain or for doing an injury to its possessor." ("Origin of Species" (1st edition) 1859, page 201.)

The study of adaptation always had for Darwin, as it has for many, a peculiar charm. His words, written Nov. 28, 1880, to Sir W. Thiselton-Dyer, are by no means inapplicable to-day: "Many of the Germans are very contemptuous about making out use of organs; but they may sneer the souls out of their bodies, and I for one shall think it the most interesting part of natural history." ("More Letters" II. page 428.)

PROTECTIVE AND AGGRESSIVE RESEMBLANCE: PROCRYPTIC AND ANTICRYPTIC COLOURING.

Colouring for the purpose of concealment is sometimes included under the head Mimicry, a classification adopted by H.W. Bates in his classical paper. Such an arrangement is inconvenient, and I have followed Wallace in keeping the two categories distinct.

The visible colours of animals are far more commonly adapted for Protective Resemblance than for any other purpose. The concealment of animals by their colours, shapes and attitudes, must have been well known from the period at which human beings first began to take an intelligent interest in Nature. An interesting early record is that of Samuel Felton, who (Dec. 2, 1763) figured and gave some account of an Acridian (Phyllotettix) from Jamaica. Of this insect he says "THE THORAX is like a leaf that is raised perpendicularly from the body." ("Phil. Trans. Roy. Soc." Vol. LIV. Tab. VI. page 55.)

Both Protective and Aggressive Resemblances were appreciated and clearly explained by Erasmus Darwin in 1794: "The colours of many animals seem adapted to their purposes of concealing themselves either to avoid danger, or to spring upon their prey." ("Zoonomia", Vol. I. page 509, London, 1794.)

Protective Resemblance of a very marked and beautiful kind is found in certain plants, inhabitants of desert areas. Examples observed by Burchell almost exactly a hundred years ago have already been mentioned. In addition to the resemblance to stones Burchell observed, although he did not publish the fact, a South African plant concealed by its likeness to the dung of birds. (Sir William Thiselton-Dyer has suggested the same method of concealment ("Annals of Botany", Vol. XX. page 123). Referring to Anacampseros papyracea, figured on plate IX., the author says of its adaptive resemblance: "At the risk of suggesting one perhaps somewhat far-fetched, I must confess that the aspect of the plant always calls to my mind the dejecta of some bird, and the more so owing to the whitening of the branches towards the tips" (loc. cit. page 126). The student of insects, who is so familiar with this very form of protective resemblance in larvae, and even perfect insects, will not be inclined to consider the suggestion far-fetched.) The observation is recorded in one of the manuscript journals kept by the great explorer during his journey. I owe the opportunity of studying it to the kindness of Mr Francis A. Burchell of the Rhodes University College, Grahamstown. The following account is given under the date July 5, 1812, when Burchell was at the Makkwarin River, about half-way between the Kuruman River and Litakun the old capital of the Bachapins (Bechuanas): "I found a curious little Crassula (not in flower) so snow white, that I should never has (have) distinguished it from the white limestones... It was an inch high and a little branchy,... and was at first mistaken for the dung of birds of the passerine order. I have often had occasion to remark that in stony place(s) there grow many small succulent plants and abound insects (chiefly Grylli) which have exactly the same colour as the ground and must for ever escape observation unless a person sit on the ground and observe very attentively."

The cryptic resemblances of animals impressed Darwin and Wallace in very different degrees, probably in part due to the fact that Wallace's tropical experiences were so largely derived from the insect world, in part to the importance assigned by Darwin to Sexual Selection "a subject which had always greatly interested me," as he says in his "Autobiography", ("Life and Letters", Vol. I. page 94.) There is no reference to Cryptic Resemblance in Darwin's section of the Joint Essay, although he gives an excellent short account of Sexual Selection (see page 295). Wallace's section on the other hand contains the following statement: "Even the peculiar colours of many animals, especially insects, so closely resembling the soil or the leaves or the trunks on which they habitually reside, are explained on the same principle; for though in the course of ages varieties of many tints may have occurred, YET THOSE RACES HAVING COLOURS BEST ADAPTED TO CONCEALMENT FROM THEIR ENEMIES WOULD INEVITABLY SURVIVE THE LONGEST." ("Journ. Proc. Linn. Soc." Vol. III. 1859, page 61. The italics are Wallace's.)

It would occupy too much space to attempt any discussion of the difference between the views of these two naturalists, but it is clear that Darwin, although fully believing in the efficiency of protective resemblance and replying to St George Mivart's contention that Natural Selection was incompetent to produce it ("Origin" (6th edition) London, 1872, pages 181, 182; see also page 66.), never entirely agreed with Wallace's estimate of its importance. Thus the following extract from a letter to Sir Joseph Hooker, May 21, 1868, refers to Wallace: "I find I must (and I always distrust myself when I differ from him) separate rather widely from him all about birds' nests and protection; he is riding that hobby to death." ("More Letters", I. page 304.) It is clear from the account given in "The Descent of Man", (London, 1874, pages 452-458. See also "Life and Letters", III. pages 123-125, and "More Letters", II. pages 59-63, 72-74, 76-78, 84-90, 92, 93.), that the divergence was due to the fact that Darwin ascribed more importance to Sexual Selection than did Wallace, and Wallace more importance to Protective Resemblance than Darwin. Thus Darwin wrote to Wallace, Oct. 12 and 13, 1867: "By the way, I cannot but think that you push protection too far in some cases, as with the stripes on the tiger." ("More Letters", I. page 283.) Here too Darwin was preferring the explanation offered by Sexual Selection ("Descent of Man" (2nd edition) 1874, pages 545, 546.), a preference which, considering the relation of the colouring of the lion and tiger to their respective environments, few naturalists will be found to share. It is also shown that Darwin contemplated the possibility of cryptic colours such as those of Patagonian animals being due to sexual selection influenced by the aspect of surrounding nature.

Nearly a year later Darwin in his letter of May 5, 1868?, expressed his agreement with Wallace's views: "Expect that I should put sexual selection as an equal, or perhaps as even a more important agent in giving colour than Natural Selection for protection." ("More Letters", II. pages 77, 78.) The conclusion expressed in the above quoted passage is opposed by the extraordinary development of Protective Resemblance in the immature stages of animals, especially insects.

It must not be supposed, however, that Darwin ascribed an unimportant role to Cryptic Resemblances, and as observations accumulated he came to recognise their efficiency in fresh groups of the animal kingdom. Thus he wrote to Wallace, May 5, 1867: "Haeckel has recently well shown that the transparency and absence of colour in the lower oceanic animals, belonging to the most different classes, may be well accounted for on the principle of protection." ("More Letters", II. page 62. See also "Descent of Man", page 261.) Darwin also admitted the justice of Professor E.S. Morse's contention that the shells of molluscs are often adaptively coloured. ("More Letters", II. page 95.) But he looked upon cryptic colouring and also mimicry as more especially Wallace's departments, and sent to him and to Professor Meldola observations and notes bearing upon these subjects. Thus the following letter given to me by Dr A.R. Wallace and now, by kind permission, published for the first time, accompanied a photograph of the chrysalis of Papilio sarpedon choredon, Feld., suspended from a leaf of its food-plant:

July 9th, Down, Beckenham, Kent.

My Dear Wallace,

Dr G. Krefft has sent me the enclosed from Sydney. A nurseryman saw a caterpillar feeding on a plant and covered the whole up, but when he searched for the cocoon (pupa), was long before he could find it, so good was its imitation in colour and form to the leaf to which it was attached. I hope that the world goes well with you. Do not trouble yourself by acknowledging this.

Ever yours

Ch. Darwin.

Another deeply interesting letter of Darwin's bearing upon protective resemblance, has only recently been shown to me by my friend Professor E.B. Wilson, the great American Cytologist. With his kind consent and that of Mr Francis Darwin, this letter, written four months before Darwin's death on April 19, 1882, is reproduced here (The letter is addressed: "Edmund B. Wilson, Esq., Assistant in Biology, John Hopkins University, Baltimore Md, U. States."):

December 21, 1881.

Dear Sir,

I thank you much for having taken so much trouble in describing fully your interesting and curious case of mimickry.

I am in the habit of looking through many scientific Journals, and though my memory is now not nearly so good as it was, I feel pretty sure that no such case as yours has been described (amongst the nudibranch) molluscs. You perhaps know the case of a fish allied to Hippocampus, (described some years ago by Dr Gunther in "Proc. Zoolog. Socy.") which clings by its tail to sea-weeds, and is covered with waving filaments so as itself to look like a piece of the same sea-weed. The parallelism between your and Dr Gunther's case makes both of them the more interesting; considering how far a fish and a mollusc stand apart. It would be difficult for anyone to explain such cases by the direct action of the environment.--I am glad that you intend to make further observations on this mollusc, and I hope that you will give a figure and if possible a coloured figure.

With all good wishes from an old brother naturalist,

I remain, Dear Sir,

Yours faithfully,

Charles Darwin.

Professor E.B. Wilson has kindly given the following account of the circumstances under which he had written to Darwin: "The case to which Darwin's letter refers is that of the nudibranch mollusc Scyllaea, which lives on the floating Sargassum and shows a really astonishing resemblance to the plant, having leaf-shaped processes very closely similar to the fronds of the sea-weed both in shape and in colour. The concealment of the animal may be judged from the fact that we found the animal quite by accident on a piece of Sargassum that had been in a glass jar in the laboratory for some time and had been closely examined in the search for hydroids and the like without disclosing the presence upon it of two large specimens of the Scyllaea (the animal, as I recall it, is about two inches long). It was first detected by its movements alone, by someone (I think a casual visitor to the laboratory) who was looking closely at the Sargassum and exclaimed 'Why, the sea-weed is moving its leaves'! We found the example in the summer of 1880 or 1881 at Beaufort, N.C., where the Johns Hopkins laboratory was located for the time being. It must have been seen by many others, before or since.

"I wrote and sent to Darwin a short description of the case at the suggestion of Brooks, with whom I was at the time a student. I was, of course, entirely unknown to Darwin (or to anyone else) and to me the principal interest of Darwin's letter is the evidence that it gives of his extraordinary kindness and friendliness towards an obscure youngster who had of course absolutely no claim upon his time or attention. The little incident made an indelible impression upon my memory and taught me a lesson that was worth learning."

VARIABLE PROTECTIVE RESEMBLANCE.

The wonderful power of rapid colour adjustment possessed by the cuttle-fish was observed by Darwin in 1832 at St Jago, Cape de Verd Islands, the first place visited during the voyage of the "Beagle". From Rio he wrote to Henslow, giving the following account of his observations, May 18, 1832: "I took several specimens of an Octopus which possessed a most marvellous power of changing its colours, equalling any chameleon, and evidently accommodating the changes to the colour of the ground which it passed over. Yellowish green, dark brown, and red, were the prevailing colours; this fact appears to be new, as far as I can find out." ("Life and Letters", I. pages 235, 236. See also Darwin's "Journal of Researches", 1876, pages 6-8, where a far more detailed account is given together with a reference to "Encycl. of Anat. and Physiol.")

Darwin was well aware of the power of individual colour adjustment, now known to be possessed by large numbers of lepidopterous pupae and larvae. An excellent example was brought to his notice by C.V. Riley ("More Letters" II, pages 385, 386.), while the most striking of the early results obtained with the pupae of butterflies--those of Mrs M.E. Barber upon Papilio nireus--was communicated by him to the Entomological Society of London. ("Trans. Ent. Soc. Lond." 1874, page 519. See also "More Letters", II. page 403.)

It is also necessary to direct attention to C.W. Beebe's ("Zoologica: N.Y. Zool. Soc." Vol. I. No. 1, Sept. 25, 1907: "Geographic variation in birds with especial reference to the effects of humidity".) recent discovery that the pigmentation of the plumage of certain birds is increased by confinement in a superhumid atmosphere. In Scardafella inca, on which the most complete series of experiments was made, the changes took place only at the moults, whether normal and annual or artificially induced at shorter periods. There was a corresponding increase in the choroidal pigment of the eye. At a certain advanced stage of feather pigmentation a brilliant iridescent bronze or green tint made its appearance on those areas where iridescence most often occurs in allied genera. Thus in birds no less than in insects, characters previously regarded as of taxonomic value, can be evoked or withheld by the forces of the environment.

WARNING OR APOSEMATIC COLOURS.

From Darwin's description of the colours and habits it is evident that he observed, in 1833, an excellent example of warning colouring in a little South American toad (Phryniscus nigricans). He described it in a letter to Henslow, written from Monte Video, Nov. 24, 1832: "As for one little toad, I hope it may be new, that it may be christened 'diabolicus.' Milton must allude to this very individual when he talks of 'squat like a toad'; its colours are by Werner ("Nomenclature of Colours", 1821) ink black, vermilion red and buff orange." ("More Letters", I. page 12.) In the "Journal of Researches" (1876, page 97.) its colours are described as follows: "If we imagine, first, that it had been steeped in the blackest ink, and then, when dry, allowed to crawl over a board, freshly painted with the brightest vermilion, so as to colour the soles of its feet and parts of its stomach, a good idea of its appearance will be gained." "Instead of being nocturnal in its habits, as other toads are, and living in damp obscure recesses, it crawls during the heat of the day about the dry sand-hillocks and arid plains,... " The appearance and habits recall T. Belt's well-known description of the conspicuous little Nicaraguan frog which he found to be distasteful to a duck. ("The Naturalist in Nicaragua" (2nd edition) London, 1888, page 321.)

The recognition of the Warning Colours of caterpillars is due in the first instance to Darwin, who, reflecting on Sexual Selection, was puzzled by the splendid colours of sexually immature organisms. He applied to Wallace "who has an innate genius for solving difficulties." ("Descent of Man", page 325. On this and the following page an excellent account of the discovery will be found, as well as in Wallace's "Natural Selection", London, 1875, pages 117-122.) Darwin's original letter exists ("Life and Letters", III. pages 93, 94.), and in it we are told that he had taken the advice given by Bates: "You had better ask Wallace." After some consideration Wallace replied that he believed the colours of conspicuous caterpillars and perfect insects were a warning of distastefulness and that such forms would be refused by birds. Darwin's reply ("Life and Letters", III. pages 94, 95.) is extremely interesting both for its enthusiasm at the brilliancy of the hypothesis and its caution in acceptance without full confirmation:

"Bates was quite right; you are the man to apply to in a difficulty. I never heard anything more ingenious than your suggestion, and I hope you may be able to prove it true. That is a splendid fact about the white moths (A single white moth which was rejected by young turkeys, while other moths were greedily devoured: "Natural Selection", 1875, page 78.); it warms one's very blood to see a theory thus almost proved to be true."

Two years later the hypothesis was proved to hold for caterpillars of many kinds by J. Jenner Weir and A.G. Butler, whose observations have since been abundantly confirmed by many naturalists. Darwin wrote to Weir, May 13, 1869: "Your verification of Wallace's suggestion seems to me to amount to quite a discovery." ("More Letters", II. page 71 (footnote).)

RECOGNITION OR EPISEMATIC CHARACTERS.

This principle does not appear to have been in any way foreseen by Darwin, although he draws special attention to several elements of pattern which would now be interpreted by many naturalists as epismes. He believed that the markings in question interfered with the cryptic effect, and came to the conclusion that, even when common to both sexes, they "are the result of sexual selection primarily applied to the male." ("Descent of Man", page 544.) The most familiar of all recognition characters was carefully explained by him, although here too explained as an ornamental feature now equally transmitted to both sexes: "The hare on her form is a familiar instance of concealment through colour; yet this principle partly fails in a closely-allied species, the rabbit, for when running to its burrow, it is made conspicuous to the sportsman, and no doubt to all beasts of prey, by its upturned white tail." ("Descent of Man", page 542.)

The analogous episematic use of the bright colours of flowers to attract insects for effecting cross-fertilisation and of fruits to attract vertebrates for effecting dispersal is very clearly explained in the "Origin". (Edition 1872, page 161. For a good example of Darwin's caution in dealing with exceptions see the allusion to brightly coloured fruit in "More Letters", II. page 348.)

It is not, at this point, necessary to treat sematic characters at any greater length. They will form the subject of a large part of the following section, where the models of Batesian (Pseudaposematic) mimicry are considered as well as the Mullerian (Synaposematic) combinations of Warning Colours.

MIMICRY,--BATESIAN OR PSEUDAPOSEMATIC, MULLERIAN OR SYNAPOSEMATIC.

The existence of superficial resemblances between animals of various degrees of affinity must have been observed for hundreds of years. Among the early examples, the best known to me have been found in the manuscript note-books and collections of W.J. Burchell, the great traveller in Africa (1810-15) and Brazil (1825-30). The most interesting of his records on this subject are brought together in the following paragraphs.

Conspicuous among well-defended insects are the dark steely or iridescent greenish blue fossorial wasps or sand-wasps, Sphex and the allied genera. Many Longicorn beetles mimic these in colour, slender shape of body and limbs, rapid movements, and the readiness with which they take to flight. On Dec. 21, 1812, Burchell captured one such beetle (Promeces viridis) at Kosi Fountain on the journey from the source of the Kuruman River to Klaarwater. It is correctly placed among the Longicorns in his catalogue, but opposite to its number is the comment "Sphex! totus purpureus."

In our own country the black-and-yellow colouring of many stinging insects, especially the ordinary wasps, affords perhaps the commonest model for mimicry. It is reproduced with more or less accuracy on moths, flies and beetles. Among the latter it is again a Longicorn which offers one of the best-known, although by no means one of the most perfect, examples. The appearance of the well-known "wasp-beetle" (Clytus arietis) in the living state is sufficiently suggestive to prevent the great majority of people from touching it. In Burchell's Brazilian collection there is a nearly allied species (Neoclytus curvatus) which appears to be somewhat less wasp-like than the British beetle. The specimen bears the number "1188," and the date March 27, 1827, when Burchell was collecting in the neighbourhood of San Paulo. Turning to the corresponding number in the Brazilian note-book we find this record: "It runs rapidly like an ichneumon or wasp, of which it has the appearance."

The formidable, well-defended ants are as freely mimicked by other insects as the sand-wasps, ordinary wasps and bees. Thus on February 17, 1901, Guy A.K. Marshall captured, near Salisbury, Mashonaland, three similar species of ants (Hymenoptera) with a bug (Hemiptera) and a Locustid (Orthoptera), the two latter mimicking the former. All the insects, seven in number, were caught on a single plant, a small bushy vetch. ("Trans. Ent. Soc. Lond." 1902, page 535, plate XIX. figs. 53-59.)

This is an interesting recent example from South Africa, and large numbers of others might be added--the observations of many naturalists in many lands; but nearly all of them known since that general awakening of interest in the subject which was inspired by the great hypotheses of H.W. Bates and Fritz Muller. We find, however, that Burchell had more than once recorded the mimetic resemblance to ants. An extremely ant-like bug (the larva of a species of Alydus) in his Brazilian collection is labelled "1141," with the date December 8, 1826, when Burchell was at the Rio das Pedras, Cubatao, near Santos. In the note-book the record is as follows: "1141 Cimex. I collected this for a Formica."

Some of the chief mimics of ants are the active little hunting spiders belonging to the family Attidae. Examples have been brought forward during many recent years, especially by my friends Dr and Mrs Peckham, of Milwaukee, the great authorities on this group of Araneae. Here too we find an observation of the mimetic resemblance recorded by Burchell, and one which adds in the most interesting manner to our knowledge of the subject. A fragment, all that is now left, of an Attid spider, captured on June 30, 1828, at Goyaz, Brazil, bears the following note, in this case on the specimen and not in the note-book: "Black... runs and seems like an ant with large extended jaws." My friend Mr R.I. Pocock, to whom I have submitted the specimen, tells me that it is not one of the group of species hitherto regarded as ant-like, and he adds, "It is most interesting that Burchell should have noticed the resemblance to an ant in its movements. This suggests that the perfect imitation in shape, as well as in movement, seen in many species was started in forms of an appropriate size and colour by the mimicry of movement alone." Up to the present time Burchell is the only naturalist who has observed an example which still exhibits this ancestral stage in the evolution of mimetic likeness.

Following the teachings of his day, Burchell was driven to believe that it was part of the fixed and inexorable scheme of things that these strange superficial resemblances existed. Thus, when he found other examples of Hemipterous mimics, including one (Luteva macrophthalma) with "exactly the manners of a Mantis," he added the sentence, "In the genus Cimex (Linn.) are to be found the outward resemblances of insects of many other genera and orders" (February 15, 1829). Of another Brazilian bug, which is not to be found in his collection, and cannot therefore be precisely identified, he wrote: "Cimex... Nature seems to have intended it to imitate a Sphex, both in colour and the rapid palpitating and movement of the antennae" (November 15, 1826). At the same time it is impossible not to feel the conviction that Burchell felt the advantage of a likeness to stinging insects and to aggressive ants, just as he recognised the benefits conferred on desert plants by spines and by concealment. Such an interpretation of mimicry was perfectly consistent with the theological doctrines of his day. (See Kirby and Spence, "An Introduction to Entomology" (1st edition), London, Vol. II. 1817, page 223.)

The last note I have selected from Burchell's manuscript refers to one of the chief mimics of the highly protected Lycid beetles. The whole assemblage of African insects with a Lycoid colouring forms a most important combination and one which has an interesting bearing upon the theories of Bates and Fritz Muller. This most wonderful set of mimetic forms, described in 1902 by Guy A.K. Marshall, is composed of flower-haunting beetles belonging to the family Lycidae, and the heterogeneous group of varied insects which mimic their conspicuous and simple scheme of colouring. The Lycid beetles, forming the centre or "models" of the whole company, are orange-brown in front for about two-thirds of the exposed surface, black behind for the remaining third. They are undoubtedly protected by qualities which make them excessively unpalatable to the bulk of insect-eating animals. Some experimental proof of this has been obtained by Mr Guy Marshall. What are the forms which surround them? According to the hypothesis of Bates they would be, at any rate mainly, palatable hard-pressed insects which only hold their own in the struggle for life by a fraudulent imitation of the trade-mark of the successful and powerful Lycidae. According to Fritz Muller's hypothesis we should expect that the mimickers would be highly protected, successful and abundant species, which (metaphorically speaking) have found it to their advantage to possess an advertisement, a danger-signal, in common with each other, and in common with the beetles in the centre of the group.

How far does the constitution of this wonderful combination--the largest and most complicated as yet known in all the world--convey to us the idea of mimicry working along the lines supposed by Bates or those suggested by Muller? Figures 1 to 52 of Mr Marshall's coloured plate ("Trans. Ent. Soc. Lond." 1902, plate XVIII. See also page 517, where the group is analysed.) represent a set of forty-two or forty-three species or forms of insects captured in Mashonaland, and all except two in the neighbourhood of Salisbury. The combination includes six species of Lycidae; nine beetles of five groups all specially protected by nauseous qualities, Telephoridae, Melyridae, Phytophaga, Lagriidae, Cantharidae; six Longicorn beetles; one Coprid beetle; eight stinging Hymenoptera; three or four parasitic Hymenoptera (Braconidae, a group much mimicked and shown by some experiments to be distasteful); five bugs (Hemiptera, a largely unpalatable group); three moths (Arctiidae and Zygaenidae, distasteful families); one fly. In fact the whole combination, except perhaps one Phytophagous, one Coprid and the Longicorn beetles, and the fly, fall under the hypothesis of Muller and not under that of Bates. And it is very doubtful whether these exceptions will be sustained: indeed the suspicion of unpalatability already besets the Longicorns and is always on the heels,--I should say the hind tarsi--of a Phytophagous beetle.

This most remarkable group which illustrates so well the problem of mimicry and the alternative hypotheses proposed for its solution, was, as I have said, first described in 1902. Among the most perfect of the mimetic resemblances in it is that between the Longicorn beetle, Amphidesmus analis, and the Lycidae. It was with the utmost astonishment and pleasure that I found this very resemblance had almost certainly been observed by Burchell. A specimen of the Amphidesmus exists in his collection and it bears "651." Turning to the same number in the African Catalogue we find that the beetle is correctly placed among the Longicorns, that it was captured at Uitenhage on Nov. 18, 1813, and that it was found associated with Lycid beetles in flowers ("consocians cum Lycis 78-87 in floribus"). Looking up Nos. 78-87 in the collection and catalogue, three species of Lycidae are found, all captured on Nov. 18, 1813, at Uitenhage. Burchell recognised the wide difference in affinity, shown by the distance between the respective numbers; for his catalogue is arranged to represent relationships. He observed, what students of mimicry are only just beginning to note and record, the coincidence between model and mimic in time and space and in habits. We are justified in concluding that he observed the close superficial likeness although he does not in this case expressly allude to it.

One of the most interesting among the early observations of superficial resemblance between forms remote in the scale of classification was made by Darwin himself, as described in the following passage from his letter to Henslow, written from Monte Video, Aug. 15, 1832: "Amongst the lower animals nothing has so much interested me as finding two species of elegantly coloured true Planaria inhabiting the dewy forest! The false relation they bear to snails is the most extraordinary thing of the kind I have ever seen." ("More Letters", I. page 9.)

Many years later, in 1867, he wrote to Fritz Muller suggesting that the resemblance of a soberly coloured British Planarian to a slug might be due to mimicry. ("Life and Letters", III. page 71.)

The most interesting copy of Bates's classical memoir on Mimicry ("Contributions to an Insect Fauna of the Amazon Valley". "Trans. Linn. Soc." Vol. XXIII. 1862, page 495.), read before the Linnean Society in 1861, is that given by him to the man who has done most to support and extend the theory. My kind friend has given that copy to me; it bears the inscription:

"Mr A.R. Wallace from his old travelling companion the Author."

Only a year and a half after the publication of the "Origin", we find that Darwin wrote to Bates on the subject which was to provide such striking evidence of the truth of Natural Selection: "I am glad to hear that you have specially attended to 'mimetic' analogies--a most curious subject; I hope you publish on it. I have for a long time wished to know whether what Dr Collingwood asserts is true--that the most striking cases generally occur between insects inhabiting the same country." (The letter is dated April 4, 1861. "More Letters", I. page 183.)

The next letter, written about six months later, reveals the remarkable fact that the illustrious naturalist who had anticipated Edward Forbes in the explanation of arctic forms on alpine heights ("I was forestalled in only one important point, which my vanity has always made me regret, namely, the explanation by means of the Glacial period of the presence of the same species of plants and of some few animals on distant mountain summits and in the arctic regions. This view pleased me so much that I wrote it out in extenso, and I believe that it was read by Hooker some years before E. Forbes published his celebrated memoir on the subject. In the very few points in which we differed, I still think that I was in the right. I have never, of course, alluded in print to my having independently worked out this view." "Autobiography, Life and Letters", I. page 88.), had also anticipated H.W. Bates in the theory of Mimicry: "What a capital paper yours will be on mimetic resemblances! You will make quite a new subject of it. I had thought of such cases as a difficulty; and once, when corresponding with Dr Collingwood, I thought of your explanation; but I drove it from my mind, for I felt that I had not knowledge to judge one way or the other." (The letter is dated Sept. 25, 1861: "More Letters", I. page 197.)

Bates read his paper before the Linnean Society, Nov. 21, 1861, and Darwin's impressions on hearing it were conveyed in a letter to the author dated Dec. 3: "Under a general point of view, I am quite convinced (Hooker and Huxley took the same view some months ago) that a philosophic view of nature can solely be driven into naturalists by treating special subjects as you have done. Under a special point of view, I think you have solved one of the most perplexing problems which could be given to solve." ("Life and Letters", II. page 378.) The memoir appeared in the following year, and after reading it Darwin wrote as follows, Nov. 20, 1862: "... In my opinion it is one of the most remarkable and admirable papers I ever read in my life... I am rejoiced that I passed over the whole subject in the "Origin", for I should have made a precious mess of it. You have most clearly stated and solved a wonderful problem... Your paper is too good to be largely appreciated by the mob of naturalists without souls; but, rely on it, that it will have LASTING value, and I cordially congratulate you on your first great work. You will find, I should think, that Wallace will fully appreciate it." ("Life and Letters", II. pages 391-393.) Four days later, Nov. 24, Darwin wrote to Hooker on the same subject: "I have now finished his paper...' it seems to me admirable. To my mind the act of segregation of varieties into species was never so plainly brought forward, and there are heaps of capital miscellaneous observations." ("More Letters", I. page 214.)

Darwin was here referring to the tendency of similar varieties of the same species to pair together, and on Nov. 25 he wrote to Bates asking for fuller information on this subject. ("More Letters", I. page 215. See also parts of Darwin's letter to Bates in "Life and Letters", II. page 392.) If Bates's opinion were well founded, sexual selection would bear a most important part in the establishment of such species. (See Poulton, "Essays on Evolution", 1908, pages 65, 85-88.) It must be admitted, however, that the evidence is as yet quite insufficient to establish this conclusion. It is interesting to observe how Darwin at once fixed on the part of Bates's memoir which seemed to bear upon sexual selection. A review of Bates's theory of Mimicry was contributed by Darwin to the "Natural History Review" (New Ser. Vol. III. 1863, page 219.) and an account of it is to be found in the "Origin" (Edition 1872, pages 375-378.) and in "The Descent of Man". (Edition 1874, pages 323-325.)

Darwin continually writes of the value of hypothesis as the inspiration of inquiry. We find an example in his letter to Bates, Nov. 22, 1860: "I have an old belief that a good observer really means a good theorist, and I fully expect to find your observations most valuable." ("More Letters", I. page 176.) Darwin's letter refers to many problems upon which Bates had theorised and observed, but as regards Mimicry itself the hypothesis was thought out after the return of the letter from the Amazons, when he no longer had the opportunity of testing it by the observation of living Nature. It is by no means improbable that, had he been able to apply this test, Bates would have recognised that his division of butterfly resemblances into two classes,--one due to the theory of mimicry, the other to the influence of local conditions,--could not be sustained.

Fritz Muller's contributions to the problem of Mimicry were all made in S.E. Brazil, and numbers of them were communicated, with other observations on natural history, to Darwin, and by him sent to Professor R. Meldola who published many of the facts. Darwin's letters to Meldola (Poulton, "Charles Darwin and the theory of Natural Selection", London, 1896, pages 199-218.) contain abundant proofs of his interest in Muller's work upon Mimicry. One deeply interesting letter (Loc. cit. pages 201, 202.) dated Jan. 23, 1872, proves that Fritz Muller before he originated the theory of Common Warning Colours (Synaposematic Resemblance or Mullerian Mimicry), which will ever be associated with his name, had conceived the idea of the production of mimetic likeness by sexual selection.

Darwin's letter to Meldola shows that he was by no means inclined to dismiss the suggestion as worthless, although he considered it daring. "You will also see in this letter a strange speculation, which I should not dare to publish, about the appreciation of certain colours being developed in those species which frequently behold other forms similarly ornamented. I do not feel at all sure that this view is as incredible as it may at first appear. Similar ideas have passed through my mind when considering the dull colours of all the organisms which inhabit dull-coloured regions, such as Patagonia and the Galapagos Is." A little later, on April 5, he wrote to Professor August Weismann on the same subject: "It may be suspected that even the habit of viewing differently coloured surrounding objects would influence their taste, and Fritz Muller even goes so far as to believe that the sight of gaudy butterflies might influence the taste of distinct species." ("Life and Letters", III. page 157.)

This remarkable suggestion affords interesting evidence that F. Muller was not satisfied with the sufficiency of Bates's theory. Nor is this surprising when we think of the numbers of abundant conspicuous butterflies which he saw exhibiting mimetic likenesses. The common instances in his locality, and indeed everywhere in tropical America, were anything but the hard-pressed struggling forms assumed by the theory of Bates. They belonged to the groups which were themselves mimicked by other butterflies. Fritz Muller's suggestion also shows that he did not accept Bates's alternative explanation of a superficial likeness between models themselves, based on some unknown influence of local physico-chemical forces. At the same time Muller's own suggestion was subject to this apparently fatal objection, that the sexual selection he invoked would tend to produce resemblances in the males rather than the females, while it is well known that when the sexes differ the females are almost invariably more perfectly mimetic than the males and in a high proportion of cases are mimetic while the males are non-mimetic.

The difficulty was met several years later by Fritz Muller's well-known theory, published in 1879 ("Kosmos", May 1879, page 100.), and immediately translated by Meldola and brought before the Entomological Society. ("Proc. Ent. Soc. Lond." 1879, page xx.) Darwin's letter to Meldola dated June 6, 1879, shows "that the first introduction of this new and most suggestive hypothesis into this country was due to the direct influence of Darwin himself, who brought it before the notice of the one man who was likely to appreciate it at its true value and to find the means for its presentation to English naturalists." ("Charles Darwin and the Theory of Natural Selection", page 214.) Of the hypothesis itself Darwin wrote "F. Muller's view of the mutual protection was quite new to me." (Ibid. page 213.) The hypothesis of Mullerian mimicry was at first strongly opposed. Bates himself could never make up his mind to accept it. As the Fellows were walking out of the meeting at which Professor Meldola explained the hypothesis, an eminent entomologist, now deceased, was heard to say to Bates: "It's a case of save me from my friends!" The new ideas encountered and still encounter to a great extent the difficulty that the theory of Bates had so completely penetrated the literature of natural history. The present writer has observed that naturalists who have not thoroughly absorbed the older hypothesis are usually far more impressed by the newer one than are those whose allegiance has already been rendered. The acceptance of Natural Selection itself was at first hindered by similar causes, as Darwin clearly recognised: "If you argue about the non-acceptance of Natural Selection, it seems to me a very striking fact that the Newtonian theory of gravitation, which seems to every one now so certain and plain, was rejected by a man so extraordinarily able as Leibnitz. The truth will not penetrate a preoccupied mind." (To Sir J. Hooker, July 28, 1868, "More Letters", I. page 305. See also the letter to A.R. Wallace, April 30, 1868, in "More Letters" II. page 77, lines 6-8 from top.)

There are many naturalists, especially students of insects, who appear to entertain an inveterate hostility to any theory of mimicry. Some of them are eager investigators in the fascinating field of geographical distribution, so essential for the study of Mimicry itself. The changes of pattern undergone by a species of Erebia as we follow it over different parts of the mountain ranges of Europe is indeed a most interesting inquiry, but not more so than the differences between e.g. the Acraea johnstoni of S.E. Rhodesia and of Kilimanjaro. A naturalist who is interested by the Erebia should be equally interested by the Acraea; and so he would be if the student of mimicry did not also record that the characteristics which distinguish the northern from the southern individuals of the African species correspond with the presence, in the north but not in the south, of certain entirely different butterflies. That this additional information should so greatly weaken, in certain minds, the appeal of a favourite study, is a psychological problem of no little interest. This curious antagonism is I believe confined to a few students of insects. Those naturalists who, standing rather farther off, are able to see the bearings of the subject more clearly, will usually admit the general support yielded by an ever-growing mass of observations to the theories of Mimicry propounded by H.W. Bates and Fritz Muller. In like manner natural selection itself was in the early days often best understood and most readily accepted by those who were not naturalists. Thus Darwin wrote to D.T. Ansted, Oct. 27, 1860: "I am often in despair in making the generality of NATURALISTS even comprehend me. Intelligent men who are not naturalists and have not a bigoted idea of the term species, show more clearness of mind." ("More Letters", I. page 175.)

Even before the "Origin" appeared Darwin anticipated the first results upon the mind of naturalists. He wrote to Asa Gray, Dec. 21, 1859: "I have made up my mind to be well abused; but I think it of importance that my notions should be read by intelligent men, accustomed to scientific argument, though NOT naturalists. It may seem absurd, but I think such men will drag after them those naturalists who have too firmly fixed in their heads that a species is an entity." ("Life and Letters" II. page 245.)

Mimicry was not only one of the first great departments of zoological knowledge to be studied under the inspiration of natural Selection, it is still and will always remain one of the most interesting and important of subjects in relation to this theory as well as to evolution. In mimicry we investigate the effect of environment in its simplest form: we trace the effects of the pattern of a single species upon that of another far removed from it in the scale of classification. When there is reason to believe that the model is an invader from another region and has only recently become an element in the environment of the species native to its second home, the problem gains a special interest and fascination. Although we are chiefly dealing with the fleeting and changeable element of colour we expect to find and we do find evidence of a comparatively rapid evolution. The invasion of a fresh model is for certain species an unusually sudden change in the forces of the environment and in some instances we have grounds for the belief that the mimetic response has not been long delayed.

MIMICRY AND SEX.

Ever since Wallace's classical memoir on mimicry in the Malayan Swallowtail butterflies, those naturalists who have written on the subject have followed his interpretation of the marked prevalence of mimetic resemblance in the female sex as compared with the male. They have believed with Wallace that the greater dangers of the female, with slower flight and often alighting for oviposition, have been in part met by the high development of this special mode of protection. The fact cannot be doubted. It is extremely common for a non-mimetic male to be accompanied by a beautifully mimetic female and often by two or three different forms of female, each mimicking a different model. The male of a polymorphic mimetic female is, in fact, usually non-mimetic (e.g. Papilio dardanus = merope), or if a mimic (e.g. the Nymphaline genus Euripus), resembles a very different model. On the other hand a non-mimetic female accompanied by a mimetic male is excessively rare. An example is afforded by the Oriental Nymphaline, Cethosia, in which the males of some species are rough mimics of the brown Danaines. In some of the orb-weaving spiders the males mimic ants, while the much larger females are non-mimetic. When both sexes mimic, it is very common in butterflies and is also known in moths, for the females to be better and often far better mimics than the males.

Although still believing that Wallace's hypothesis in large part accounts for the facts briefly summarised above, the present writer has recently been led to doubt whether it offers a complete explanation. Mimicry in the male, even though less beneficial to the species than mimicry in the female, would still surely be advantageous. Why then is it so often entirely restricted to the female? While the attempt to find an answer to this question was haunting me, I re-read a letter written by Darwin to Wallace, April 15, 1868, containing the following sentences: "When female butterflies are more brilliant than their males you believe that they have in most cases, or in all cases, been rendered brilliant so as to mimic some other species, and thus escape danger. But can you account for the males not having been rendered equally brilliant and equally protected? Although it may be most for the welfare of the species that the female should be protected, yet it would be some advantage, certainly no disadvantage, for the unfortunate male to enjoy an equal immunity from danger. For my part, I should say that the female alone had happened to vary in the right manner, and that the beneficial variations had been transmitted to the same sex alone. Believing in this, I can see no improbability (but from analogy of domestic animals a strong probability) that variations leading to beauty must often have occurred in the males alone, and been transmitted to that sex alone. Thus I should account in many cases for the greater beauty of the male over the female, without the need of the protective principle." ("More Letters", II. pages 73, 74. On the same subject--"the gay-coloured females of Pieris" (Perrhybris (Mylothris) pyrrha of Brazil), Darwin wrote to Wallace, May 5, 1868, as follows: "I believe I quite follow you in believing that the colours are wholly due to mimicry; and I further believe that the male is not brilliant from not having received through inheritance colour from the female, and from not himself having varied; in short, that he has not been influenced by selection." It should be noted that the male of this species does exhibit a mimetic pattern on the under surface. "More Letters" II. page 78.)

The consideration of the facts of mimicry thus led Darwin to the conclusion that the female happens to vary in the right manner more commonly than the male, while the secondary sexual characters of males supported the conviction "that from some unknown cause such characters (viz. new characters arising in one sex and transmitted to it alone) apparently appear oftener in the male than in the female." (Letter from Darwin to Wallace, May 5, 1867, "More Letters", II. Page 61.)

Comparing these conflicting arguments we are led to believe that the first is the stronger. Mimicry in the male would be no disadvantage but an advantage, and when it appears would be and is taken advantage of by selection. The secondary sexual characters of males would be no advantage but a disadvantage to females, and, as Wallace thinks, are withheld from this sex by selection. It is indeed possible that mimicry has been hindered and often prevented from passing to the males by sexual selection. We know that Darwin was much impressed ("Descent of Man", page 325.) by Thomas Belt's daring and brilliant suggestion that the white patches which exist, although ordinarily concealed, on the wings of mimetic males of certain Pierinae (Dismorphia), have been preserved by preferential mating. He supposed this result to have been brought about by the females exhibiting a deep-seated preference for males that displayed the chief ancestral colour, inherited from periods before any mimetic pattern had been evolved in the species. But it has always appeared to me that Belt's deeply interesting suggestion requires much solid evidence and repeated confirmation before it can be accepted as a valid interpretation of the facts. In the present state of our knowledge, at any rate of insects and especially of Lepidoptera, it is probable that the female is more apt to vary than the male and that an important element in the interpretation of prevalent female mimicry is provided by this fact.

In order adequately to discuss the question of mimicry and sex it would be necessary to analyse the whole of the facts, so far as they are known in butterflies. On the present occasion it is only possible to state the inferences which have been drawn from general impressions,--inferences which it is believed will be sustained by future inquiry.

(1) Mimicry may occasionally arise in one sex because the differences which distinguish it from the other sex happen to be such as to afford a starting-point for the resemblance. Here the male is at no disadvantage as compared with the female, and the rarity of mimicry in the male alone (e.g. Cethosia) is evidence that the great predominance of female mimicry is not to be thus explained.

(2) The tendency of the female to dimorphism and polymorphism has been of great importance in determining this predominance. Thus if the female appear in two different forms and the male in only one it will be twice as probable that she will happen to possess a sufficient foundation for the evolution of mimicry.

(3) The appearance of melanic or partially melanic forms in the female has been of very great service, providing as it does a change of ground-colour. Thus the mimicry of the black generally red-marked American "Aristolochia swallowtails" (Pharmacophagus) by the females of Papilio swallowtails was probably begun in this way.

(4) It is probably incorrect to assume with Haase that mimicry always arose in the female and was later acquired by the male. Both sexes of the third section of swallowtails (Cosmodesmus) mimic Pharmacophagus in America, far more perfectly than do the females of Papilio. But this is not due to Cosmodesmus presenting us with a later stage of history begun in Papilio; for in Africa Cosmodesmus is still mimetic (of Danainae) in both sexes although the resemblances attained are imperfect, while many African species of Papilio have non-mimetic males with beautifully mimetic females. The explanation is probably to be sought in the fact that the females of Papilio are more variable and more often tend to become dimorphic than those of Cosmodesmus, while the latter group has more often happened to possess a sufficient foundation for the origin of the resemblance in patterns which, from the start, were common to male and female.

(5) In very variable species with sexes alike, mimicry can be rapidly evolved in both sexes out of very small beginnings. Thus the reddish marks which are common in many individuals of Limenitis arthemis were almost certainly the starting-point for the evolution of the beautifully mimetic L. archippus. Nevertheless in such cases, although there is no reason to suspect any greater variability, the female is commonly a somewhat better mimic than the male and often a very much better mimic. Wallace's principle seems here to supply the obvious interpretation.

(6) When the difference between the patterns of the model and presumed ancestor of the mimic is very great, the female is often alone mimetic; when the difference is comparatively small, both sexes are commonly mimetic. The Nymphaline genus Hypolimnas is a good example. In Hypolimnas itself the females mimic Danainae with patterns very different from those preserved by the non-mimetic males: in the sub-genus Euralia, both sexes resemble the black and white Ethiopian Danaines with patterns not very dissimilar from that which we infer to have existed in the non-mimetic ancestor.

(7) Although a melanic form or other large variation may be of the utmost importance in facilitating the start of a mimetic likeness, it is impossible to explain the evolution of any detailed resemblance in this manner. And even the large colour variation itself may well be the expression of a minute and "continuous" change in the chemical and physical constitution of pigments.

SEXUAL SELECTION (EPIGAMIC CHARACTERS).

We do not know the date at which the idea of Sexual Selection arose in Darwin's mind, but it was probably not many years after the sudden flash of insight which, in October 1838, gave to him the theory of Natural Selection. An excellent account of Sexual Selection occupies the concluding paragraph of Part I. of Darwin's Section of the Joint Essay on Natural Selection, read July 1st, 1858, before the Linnean Society. ("Journ. Proc. Linn. Soc." Vol. III. 1859, page 50.) The principles are so clearly and sufficiently stated in these brief sentences that it is appropriate to quote the whole: "Besides this natural means of selection, by which those individuals are preserved, whether in their egg, or larval, or mature state, which are best adapted to the place they fill in nature, there is a second agency at work in most unisexual animals, tending to produce the same effect, namely, the struggle of the males for the females. These struggles are generally decided by the law of battle, but in the case of birds, apparently, by the charms of their song, by their beauty or their power of courtship, as in the dancing rock-thrush of Guiana. The most vigorous and healthy males, implying perfect adaptation, must generally gain the victory in their contests. This kind of selection, however, is less rigorous than the other; it does not require the death of the less successful, but gives to them fewer descendants. The struggle falls, moreover, at a time of year when food is generally abundant, and perhaps the effect chiefly produced would be the modification of the secondary sexual characters, which are not related to the power of obtaining food, or to defence from enemies, but to fighting with or rivalling other males. The result of this struggle amongst the males may be compared in some respects to that produced by those agriculturists who pay less attention to the careful selection of all their young animals, and more to the occasional use of a choice mate."

A full exposition of Sexual Selection appeared in the "The Descent of Man" in 1871, and in the greatly augmented second edition, in 1874. It has been remarked that the two subjects, "The Descent of Man and Selection in Relation to Sex", seem to fuse somewhat imperfectly into the single work of which they form the title. The reason for their association is clearly shown in a letter to Wallace, dated May 28, 1864: "... I suspect that a sort of sexual selection has been the most powerful means of changing the races of man." ("More Letters", II. page 33.)

Darwin, as we know from his Autobiography ("Life and Letters", I. page 94.), was always greatly interested in this hypothesis, and it has been shown in the preceding pages that he was inclined to look favourably upon it as an interpretation of many appearances usually explained by Natural Selection. Hence Sexual Selection, incidentally discussed in other sections of the present essay, need not be considered at any length, in the section specially allotted to it.

Although so interested in the subject and notwithstanding his conviction that the hypothesis was sound, Darwin was quite aware that it was probably the most vulnerable part of the "Origin". Thus he wrote to H.W. Bates, April 4, 1861: "If I had to cut up myself in a review I would have (worried?) and quizzed sexual selection; therefore, though I am fully convinced that it is largely true, you may imagine how pleased I am at what you say on your belief." ("More Letters", I. page 183.)

The existence of sound-producing organs in the males of insects was, Darwin considered, the strongest evidence in favour of the operation of sexual selection in this group. ("Life and Letters", III. pages 94, 138.) Such a conclusion has received strong support in recent years by the numerous careful observations of Dr F.A. Dixey ("Proc. Ent. Soc. Lond." 1904, page lvi; 1905, pages xxxvii, liv; 1906, page ii.) and Dr G.B. Longstaff ("Proc. Ent. Soc. Lond." 1905, page xxxv; "Trans. Ent. Soc. Lond." 1905, page 136; 1908, page 607.) on the scents of male butterflies. The experience of these naturalists abundantly confirms and extends the account given by Fritz Muller ("Jen. Zeit." Vol. XI. 1877, page 99; "Trans. Ent. Soc. Lond." 1878, page 211.) of the scents of certain Brazilian butterflies. It is a remarkable fact that the apparently epigamic scents of male butterflies should be pleasing to man while the apparently aposematic scents in both sexes of species with warning colours should be displeasing to him. But the former is far more surprising than the latter. It is not perhaps astonishing that a scent which is ex hypothesi unpleasant to an insect-eating Vertebrate should be displeasing to the human sense; but it is certainly wonderful that an odour which is ex hypothesi agreeable to a female butterfly should also be agreeable to man.

Entirely new light upon the seasonal appearance of epigamic characters is shed by the recent researches of C.W. Beebe ("The American Naturalist", Vol. XLII. No. 493, Jan. 1908, page 34.), who caused the scarlet tanager (Piranga erythromelas) and the bobolink (Dolichonyx oryzivorus) to retain their breeding plumage through the whole year by means of fattening food, dim illumination, and reduced activity. Gradual restoration to the light and the addition of meal-worms to the diet invariably brought back the spring song, even in the middle of winter. A sudden alteration of temperature, either higher or lower, caused the birds nearly to stop feeding, and one tanager lost weight rapidly and in two weeks moulted into the olive-green winter plumage. After a year, and at the beginning of the normal breeding season, "individual tanagers and bobolinks were gradually brought under normal conditions and activities," and in every case moulted from nuptial plumage to nuptial plumage. "The dull colours of the winter season had been skipped." The author justly claims to have established "that the sequence of plumage in these birds is not in any way predestined through inheritance..., but that it may be interrupted by certain factors in the environmental complex."

XVI. GEOGRAPHICAL DISTRIBUTION OF PLANTS. By Sir William Thiselton-Dyer, K.C.M.G., C.I.E. Sc.D., F.R.S.

The publication of "The Origin of Species" placed the study of Botanical Geography on an entirely new basis. It is only necessary to study the monumental "Geographie Botanique raisonnee" of Alphonse De Candolle, published four years earlier (1855), to realise how profound and far-reaching was the change. After a masterly and exhaustive discussion of all available data De Candolle in his final conclusions could only arrive at a deadlock. It is sufficient to quote a few sentences:--

"L'opinion de Lamarck est aujourd'hui abandonee par tous les naturalistes qui ont etudie sagement les modifications possibles des etres organises...

"Et si l'on s'ecarte des exagerations de Lamarck, si l'on suppose un premier type de chaque genre, de chaque famille tout au moins, on se trouve encore a l'egard de l'origine de ces types en presence de la grande question de la creation.

"Le seul parti a prendre est donc d'envisager les etres organises comme existant depuis certaines epoques, avec leurs qualites particulieres." (Vol. II. page 1107.)

Reviewing the position fourteen years afterwards, Bentham remarked:--"These views, generally received by the great majority of naturalists at the time De Candolle wrote, and still maintained by a few, must, if adhered to, check all further enquiry into any connection of facts with causes," and he added, "there is little doubt but that if De Candolle were to revise his work, he would follow the example of so many other eminent naturalists, and... insist that the present geographical distribution of plants was in most instances a derivative one, altered from a very different former distribution." ("Pres. Addr." (1869) "Proc. Linn. Soc." 1868-69, page lxviii.)

Writing to Asa Gray in 1856, Darwin gave a brief preliminary account of his ideas as to the origin of species, and said that geographical distribution must be one of the tests of their validity. ("Life and Letters", II. page 78.) What is of supreme interest is that it was also their starting-point. He tells us:--"When I visited, during the voyage of H.M.S. "Beagle", the Galapagos Archipelago,... I fancied myself brought near to the very act of creation. I often asked myself how these many peculiar animals and plants had been produced: the simplest answer seemed to be that the inhabitants of the several islands had descended from each other, undergoing modification in the course of their descent." ("The Variation of Animals and Plants" (2nd edition), 1890, I. pages 9, 10.) We need not be surprised then, that in writing in 1845 to Sir Joseph Hooker, he speaks of "that grand subject, that almost keystone of the laws of creation, Geographical Distribution." ("Life and Letters", I. page 336.)

Yet De Candolle was, as Bentham saw, unconsciously feeling his way, like Lyell, towards evolution, without being able to grasp it. They both strove to explain phenomena by means of agencies which they saw actually at work. If De Candolle gave up the ultimate problem as insoluble:--"La creation ou premiere formation des etres organises echappe, par sa nature et par son anciennete, a nos moyens d'observation" (Loc. cit. page 1106.), he steadily endeavoured to minimise its scope. At least half of his great work is devoted to the researches by which he extricated himself from a belief in species having had a multiple origin, the view which had been held by successive naturalists from Gmelin to Agassiz. To account for the obvious fact that species constantly occupy dissevered areas, De Candolle made a minute study of their means of transport. This was found to dispose of the vast majority of cases, and the remainder he accounted for by geographical change. (Loc. cit. page 1116.)

But Darwin strenuously objected to invoking geographical change as a solution of every difficulty. He had apparently long satisfied himself as to the "permanence of continents and great oceans." Dana, he tells us "was, I believe, the first man who maintained" this ("Life and Letters", III. page 247. Dana says:--"The continents and oceans had their general outline or form defined in earliest time," "Manual of Geology", revised edition. Philadelphia, 1869, page 732. I have no access to an earlier edition.), but he had himself probably arrived at it independently. Modern physical research tends to confirm it. The earth's centre of gravity, as pointed out by Pratt from the existence of the Pacific Ocean, does not coincide with its centre of figure, and it has been conjectured that the Pacific Ocean dates its origin from the separation of the moon from the earth.

The conjecture appears to be unnecessary. Love shows that "the force that keeps the Pacific Ocean on one side of the earth is gravity, directed more towards the centre of gravity than the centre of the figure." ("Report of the 77th Meeting of the British Association" (Leicester, 1907), London, 1908, page 431.) I can only summarise the conclusions of a technical but masterly discussion. "The broad general features of the distribution of continent and ocean can be regarded as the consequences of simple causes of a dynamical character," and finally, "As regards the contour of the great ocean basins, we seem to be justified in saying that the earth is approximately an oblate spheroid, but more nearly an ellipsoid with three unequal axes, having its surface furrowed according to the formula for a certain spherical harmonic of the third degree" (Ibid. page 436.), and he shows that this furrowed surface must be produced "if the density is greater in one hemispheroid than in the other, so that the position of the centre of gravity is eccentric." (Ibid. page 431.) Such a modelling of the earth's surface can only be referred to a primitive period of plasticity. If the furrows account for the great ocean basins, the disposition of the continents seems equally to follow. Sir George Darwin has pointed out that they necessarily "arise from a supposed primitive viscosity or plasticity of the earth's mass. For during this course of evolution the earth's mass must have suffered a screwing motion, so that the polar regions have travelled a little from west to east relatively to the equator. This affords a possible explanation of the north and south trend of our great continents." ("Encycl. Brit." (9th edition), Vol. XXIII. "Tides", page 379.)

It would be trespassing on the province of the geologist to pursue the subject at any length. But as Wallace ("Island Life" (2nd edition), 1895, page 103.), who has admirably vindicated Darwin's position, points out, the "question of the permanence of our continents... lies at the root of all our inquiries into the great changes of the earth and its inhabitants." But he proceeds: "The very same evidence which has been adduced to prove the GENERAL stability and permanence of our continental areas also goes to prove that they have been subjected to wonderful and repeated changes in DETAIL." (Loc. cit. page 101.) Darwin of course would have admitted this, for with a happy expression he insisted to Lyell (1856) that "the skeletons, at least, of our continents are ancient." ("More Letters", II. page 135.) It is impossible not to admire the courage and tenacity with which he carried on the conflict single-handed. But he failed to convince Lyell. For we still find him maintaining in the last edition of the "Principles": "Continents therefore, although permanent for whole geological epochs, shift their positions entirely in the course of ages." (Lyell's "Principles of Geology" (11th edition), London, 1872, I. page 258.)

Evidence, however, steadily accumulates in Darwin's support. His position still remains inexpugnable that it is not permissible to invoke geographical change to explain difficulties in distribution without valid geological and physical support. Writing to Mellard Reade, who in 1878 had said, "While believing that the ocean-depths are of enormous age, it is impossible to reject other evidences that they have once been land," he pointed out "the statement from the 'Challenger' that all sediment is deposited within one or two hundred miles from the shores." ("More Letters", II. page 146.) The following year Sir Archibald Geikie ("Geographical Evolution", "Proc. R. Geogr. Soc." 1879, page 427.) informed the Royal Geographical Society that "No part of the results obtained by the 'Challenger' expedition has a profounder interest for geologists and geographers than the proof which they furnish that the floor of the ocean basins has no real analogy among the sedimentary formations which form most of the framework of the land."

Nor has Darwin's earlier argument ever been upset. "The fact which I pointed out many years ago, that all oceanic islands are volcanic (except St Paul's, and now that is viewed by some as the nucleus of an ancient volcano), seem to me a strong argument that no continent ever occupied the great oceans." ("More Letters", II. page 146.)

Dr Guppy, who devoted several years to geological and botanical investigations in the Pacific, found himself forced to similar conclusions. "It may be at once observed," he says, "that my belief in the general principle that islands have always been islands has not been shaken," and he entirely rejects "the hypothesis of a Pacific continent." He comes back, in full view of the problems on the spot, to the position from which, as has been seen, Darwin started: "If the distribution of a particular group of plants or animals does not seem to accord with the present arrangement of the land, it is by far the safest plan, even after exhausting all likely modes of explanation, not to invoke the intervention of geographical changes; and I scarcely think that our knowledge of any one group of organisms is ever sufficiently precise to justify a recourse to hypothetical alterations in the present relations of land and sea." ("Observations of a Naturalist in the Pacific between 1896 and 1899", London, 1903, I. page 380.) Wallace clinches the matter when he finds "almost the whole of the vast areas of the Atlantic, Pacific, Indian, and Southern Oceans, without a solitary relic of the great islands or continents supposed to have sunk beneath their waves." ("Island Life", page 105.)

Writing to Wallace (1876), Darwin warmly approves the former's "protest against sinking imaginary continents in a quite reckless manner, as was stated by Forbes, followed, alas, by Hooker, and caricatured by Wollaston and (Andrew) Murray." ("Life and Letters", III. page 230.) The transport question thus became of enormously enhanced importance. We need not be surprised then at his writing to Lyell in 1856:--"I cannot avoid thinking that Forbes's 'Atlantis' was an ill-service to science, as checking a close study of means of dissemination" (Ibid. II. page 78.), and Darwin spared no pains to extend our knowledge of them. He implores Hooker, ten years later, to "admit how little is known on the subject," and summarises with some satisfaction what he had himself achieved:--"Remember how recently you and others thought that salt water would soon kill seeds... Remember that no one knew that seeds would remain for many hours in the crops of birds and retain their vitality; that fish eat seeds, and that when the fish are devoured by birds the seeds can germinate, etc. Remember that every year many birds are blown to Madeira and to the Bermudas. Remember that dust is blown 1000 miles across the Atlantic." ("More Letters", I. page 483.)

It has always been the fashion to minimise Darwin's conclusions, and these have not escaped objection. The advocatus diaboli has a useful function in science. But in attacking Darwin his brief is generally found to be founded on a slender basis of facts. Thus Winge and Knud Andersen have examined many thousands of migratory birds and found "that their crops and stomachs were always empty. They never observed any seeds adhering to the feathers, beaks or feet of the birds." (R.F. Scharff, "European Animals", page 64, London, 1907.) The most considerable investigation of the problem of Plant Dispersal since Darwin is that of Guppy. He gives a striking illustration of how easily an observer may be led into error by relying on negative evidence.

"When Ekstam published, in 1895, the results of his observations on the plants of Nova Zembla, he observed that he possessed no data to show whether swimming and wading birds fed on berries; and he attached all importance to dispersal by winds. On subsequently visiting Spitzbergen he must have been at first inclined, therefore, to the opinion of Nathorst, who, having found only a solitary species of bird (a snow-sparrow) in that region, naturally concluded that birds had been of no importance as agents in the plant-stocking. However, Ekstam's opportunities were greater, and he tells us that in the craws of six specimens of Lagopus hyperboreus shot in Spitzbergen in August he found represented almost 25 per cent. of the usual phanerogamic flora of that region in the form of fruits, seeds, bulbils, flower-buds, leaf-buds, etc... "

"The result of Ekstam's observations in Spitzbergen was to lead him to attach a very considerable importance in plant dispersal to the agency of birds; and when in explanation of the Scandinavian elements in the Spitzbergen flora he had to choose between a former land connection and the agency of birds, he preferred the bird." (Guppy, op. cit. II. pages 511, 512.)

Darwin objected to "continental extensions" on geological grounds, but he also objected to Lyell that they do not "account for all the phenomena of distribution on islands" ("Life and Letters", II. page 77.), such for example as the absence of Acacias and Banksias in New Zealand. He agreed with De Candolle that "it is poor work putting together the merely POSSIBLE means of distribution." But he also agreed with him that they were the only practicable door of escape from multiple origins. If they would not work then "every one who believes in single centres will have to admit continental extensions" (Ibid. II. page 82.), and that he regarded as a mere counsel of despair:--"to make continents, as easily as a cook does pancakes." (Ibid. II. page 74.)

The question of multiple origins however presented itself in another shape where the solution was much more difficult. The problem, as stated by Darwin, is this:--"The identity of many plants and animals, on mountain-summits, separated from each other by hundreds of miles of lowlands... without the apparent possibility of their having migrated from one point to the other." He continues, "even as long ago as 1747, such facts led Gmelin to conclude that the same species must have been independently created at several distinct points; and we might have remained in this same belief, had not Agassiz and others called vivid attention to the Glacial period, which affords... a simple explanation of the facts." ("Origin of Species" (6th edition) page 330.)

The "simple explanation" was substantially given by E. Forbes in 1846. It is scarcely too much to say that it belongs to the same class of fertile and far-reaching ideas as "natural selection" itself. It is an extraordinary instance, if one were wanted at all, of Darwin's magnanimity and intense modesty that though he had arrived at the theory himself, he acquiesced in Forbes receiving the well-merited credit. "I have never," he says, "of course alluded in print to my having independently worked out this view." But he would have been more than human if he had not added:--"I was forestalled in... one important point, which my vanity has always made me regret." ("Life and Letters", I. page 88.)

Darwin, however, by applying the theory to trans-tropical migration, went far beyond Forbes. The first enunciation to this is apparently contained in a letter to Asa Gray in 1858. The whole is too long to quote, but the pith is contained in one paragraph. "There is a considerable body of geological evidence that during the Glacial epoch the whole world was colder; I inferred that,... from erratic boulder phenomena carefully observed by me on both the east and west coast of South America. Now I am so bold as to believe that at the height of the Glacial epoch, AND WHEN ALL TROPICAL PRODUCTIONS MUST HAVE BEEN CONSIDERABLY DISTRESSED, several temperate forms slowly travelled into the heart of the Tropics, and even reached the southern hemisphere; and some few southern forms penetrated in a reverse direction northward." ("Life and Letters", II. page 136.) Here again it is clear that though he credits Agassiz with having called vivid attention to the Glacial period, he had himself much earlier grasped the idea of periods of refrigeration.

Putting aside the fact, which has only been made known to us since Darwin's death, that he had anticipated Forbes, it is clear that he gave the theory a generality of which the latter had no conception. This is pointed out by Hooker in his classical paper "On the Distribution of Arctic Plants" (1860). "The theory of a southern migration of northern types being due to the cold epochs preceding and during the glacial, originated, I believe, with the late Edward Forbes; the extended one, of the trans-tropical migration, is Mr Darwin's." ("Linn. Trans." XXIII. page 253. The attempt appears to have been made to claim for Heer priority in what I may term for short the arctic-alpine theory (Scharff, "European Animals", page 128). I find no suggestion of his having hit upon it in his correspondence with Darwin or Hooker. Nor am I aware of any reference to his having done so in his later publications. I am indebted to his biographer, Professor Schroter, of Zurich, for an examination of his earlier papers with an equally negative result.) Assuming that local races have derived from a common ancestor, Hooker's great paper placed the fact of the migration on an impregnable basis. And, as he pointed out, Darwin has shown that "such an explanation meets the difficulty of accounting for the restriction of so many American and Asiatic arctic types to their own peculiar longitudinal zones, and for what is a far greater difficulty, the representation of the same arctic genera by most closely allied species in different longitudes."

The facts of botanical geography were vital to Darwin's argument. He had to show that they admitted of explanation without assuming multiple origins for species, which would be fatal to the theory of Descent. He had therefore to strengthen and extend De Candolle's work as to means of transport. He refused to supplement them by hypothetical geographical changes for which there was no independent evidence: this was simply to attempt to explain ignotum per ignotius. He found a real and, as it has turned out, a far-reaching solution in climatic change due to cosmical causes which compelled the migration of species as a condition of their existence. The logical force of the argument consists in dispensing with any violent assumption, and in showing that the principle of descent is adequate to explain the ascertained facts.

It does not, I think, detract from the merit of Darwin's conclusions that the tendency of modern research has been to show that the effects of the Glacial period were less simple, more localised and less general than he perhaps supposed. He admitted that "equatorial refrigeration... must have been small." ("More Letters", I. page 177.) It may prove possible to dispense with it altogether. One cannot but regret that as he wrote to Bates:--"the sketch in the 'Origin' gives a very meagre account of my fuller MS. essay on this subject." (Loc. cit.) Wallace fully accepted "the effect of the Glacial epoch in bringing about the present distribution of Alpine and Arctic plants in the NORTHERN HEMISPHERE," but rejected "the lowering of the temperature of the tropical regions during the Glacial period" in order to account for their presence in the SOUTHERN hemisphere. ("More Letters", II. page 25 (footnote 1).) The divergence however does not lie very deep. Wallace attaches more importance to ordinary means of transport. "If plants can pass in considerable numbers and variety over wide seas and oceans, it must be yet more easy for them to traverse continuous areas of land, wherever mountain-chains offer suitable stations." ("Island Life" (2nd edition), London, 1895, page 512.) And he argues that such periodical changes of climate, of which the Glacial period may be taken as a type, would facilitate if not stimulate the process. (Loc. cit. page 518.)

It is interesting to remark that Darwin drew from the facts of plant distribution one of his most ingenious arguments in support of this theory. (See "More Letters", I. page 424.) He tells us, "I was led to anticipate that the species of the larger genera in each country would oftener present varieties, than the species of the smaller genera." ("Origin", page 44.) He argues "where, if we may use the expression, the manufactory of species has been active, we ought generally to find the manufactory still in action." (Ibid. page 45.) This proved to be the case. But the labour imposed upon him in the study was immense. He tabulated local floras "belting the whole northern hemisphere" ("More Letters", I. page 107.), besides voluminous works such as De Candolle's "Prodromus". The results scarcely fill a couple of pages. This is a good illustration of the enormous pains which he took to base any statement on a secure foundation of evidence, and for this the world, till the publication of his letters, could not do him justice. He was a great admirer of Herbert Spencer, whose "prodigality of original thought" astonished him. "But," he says, "the reflection constantly recurred to me that each suggestion, to be of real value to service, would require years of work." (Ibid. II. page 235.)

At last the ground was cleared and we are led to the final conclusion. "If the difficulties be not insuperable in admitting that in the long course of time all the individuals of the same species belonging to the same genus, have proceeded from some one source; then all the grand leading facts of geographical distribution are explicable on the theory of migration, together with subsequent modification and the multiplication of new forms." ("Origin", page 360.) In this single sentence Darwin has stated a theory which, as his son F. Darwin has said with justice, has "revolutionized botanical geography." ("The Botanical Work of Darwin", "Ann. Bot." 1899, page xi.) It explains how physical barriers separate and form botanical regions; how allied species become concentrated in the same areas; how, under similar physical conditions, plants may be essentially dissimilar, showing that descent and not the surroundings is the controlling factor; how insular floras have acquired their peculiarities; in short how the most various and apparently uncorrelated problems fall easily and inevitably into line.

The argument from plant distribution was in fact irresistible. A proof, if one were wanted, was the immediate conversion of what Hooker called "the stern keen intellect" ("More Letters", I. page 134.) of Bentham, by general consent the leading botanical systematist at the time. It is a striking historical fact that a paper of his own had been set down for reading at the Linnean Society on the same day as Darwin's, but had to give way. In this he advocated the fixity of species. He withdrew it after hearing Darwin's. We can hardly realise now the momentous effect on the scientific thought of the day of the announcement of the new theory. Years afterwards (1882) Bentham, notwithstanding his habitual restraint, could not write of it without emotion. "I was forced, however reluctantly, to give up my long-cherished convictions, the results of much labour and study." The revelation came without preparation. Darwin, he wrote, "never made any communications to me in relation to his views and labours." But, he adds, "I... fully adopted his theories and conclusions, notwithstanding the severe pain and disappointment they at first occasioned me." ("Life and Letters", II. page 294.) Scientific history can have few incidents more worthy. I do not know what is most striking in the story, the pathos or the moral dignity of Bentham's attitude.

Darwin necessarily restricted himself in the "Origin" to establishing the general principles which would account for the facts of distribution, as a part of his larger argument, without attempting to illustrate them in particular cases. This he appears to have contemplated doing in a separate work. But writing to Hooker in 1868 he said:--"I shall to the day of my death keep up my full interest in Geographical Distribution, but I doubt whether I shall ever have strength to come in any fuller detail than in the "Origin" to this grand subject." ("More Letters", II. page 7.) This must be always a matter for regret. But we may gather some indication of his later speculations from the letters, the careful publication of which by F. Darwin has rendered a service to science, the value of which it is difficult to exaggerate. They admit us to the workshop, where we see a great theory, as it were, in the making. The later ideas that they contain were not it is true public property at the time. But they were communicated to the leading biologists of the day and indirectly have had a large influence.

If Darwin laid the foundation, the present fabric of Botanical Geography must be credited to Hooker. It was a happy partnership. The far-seeing, generalising power of the one was supplied with data and checked in conclusions by the vast detailed knowledge of the other. It may be permitted to quote Darwin's generous acknowledgment when writing the "Origin":--"I never did pick any one's pocket, but whilst writing my present chapter I keep on feeling (even when differing most from you) just as if I were stealing from you, so much do I owe to your writings and conversation, so much more than mere acknowledgements show." ("Life and Letters", II. page 148 (footnote).) Fourteen years before he had written to Hooker: "I know I shall live to see you the first authority in Europe on... Geographical Distribution." (Ibid. I. page 336.) We owe it to Hooker that no one now undertakes the flora of a country without indicating the range of the species it contains. Bentham tells us: "After De Candolle, independently of the great works of Darwin... the first important addition to the science of geographical botany was that made by Hooker in his "Introductory Essay to the Flora of Tasmania", which, though contemporaneous only with the "Origin of Species", was drawn up with a general knowledge of his friend's observations and views." (Pres. Addr. (1869), "Proc. Linn. Soc." 1868-69, page lxxiv.) It cannot be doubted that this and the great memoir on the "Distribution of Arctic Plants" were only less epoch-making than the "Origin" itself, and must have supplied a powerful support to the general theory of organic evolution.

Darwin always asserted his "entire ignorance of Botany." ("More Letters", I. page 400.) But this was only part of his constant half-humorous self-depreciation. He had been a pupil of Henslow, and it is evident that he had a good working knowledge of systematic botany. He could find his way about in the literature and always cites the names of plants with scrupulous accuracy. It was because he felt the want of such a work for his own researches that he urged the preparation of the "Index Kewensis", and undertook to defray the expense. It has been thought singular that he should have been elected a "correspondant" of the Academie des Sciences in the section of Botany, but it is not surprising that his work in Geographical Botany made the botanists anxious to claim him. His heart went with them. "It has always pleased me," he tells us, "to exalt plants in the scale of organised beings." ("Life and Letters", I. page 98.) And he declares that he finds "any proposition more easily tested in botanical works (Ibid. II. page 99.) than in zoological."

In the "Introductory Essay" Hooker dwelt on the "continuous current of vegetation from Scandinavia to Tasmania" ("Introductory Essay to the Flora of Tasmania", London, 1859. Reprinted from the "Botany of the Antarctic Expedition", Part III., "Flora of Tasmania", Vol I. page ciii.), but finds little evidence of one in the reverse direction. "In the New World, Arctic, Scandinavian, and North American genera and species are continuously extended from the north to the south temperate and even Antarctic zones; but scarcely one Antarctic species, or even genus advances north beyond the Gulf of Mexico" (page civ.). Hooker considered that this negatived "the idea that the Southern and Northern Floras have had common origin within comparatively modern geological epochs." (Loc. cit.) This is no doubt a correct conclusion. But it is difficult to explain on Darwin's view alone, of alternating cold in the two hemispheres, the preponderant migration from the north to the south. He suggests, therefore, that it "is due to the greater extent of land in the north and to the northern forms... having... been advanced through natural selection and competition to a higher stage of perfection or dominating power." ("Origin of Species" (6th edition), page 340; cf. also "Life and Letters", II. page 142.) The present state of the Flora of New Zealand affords a striking illustration of the correctness of this view. It is poor in species, numbering only some 1400, of which three-fourths are endemic. They seem however quite unable to resist the invasion of new comers and already 600 species of foreign origin have succeeded in establishing themselves.

If we accept the general configuration of the earth's surface as permanent a continuous and progressive dispersal of species from the centre to the circumference, i.e. southwards, seems inevitable. If an observer were placed above a point in St George's Channel from which one half of the globe was visible he would see the greatest possible quantity of land spread out in a sort of stellate figure. The maritime supremacy of the English race has perhaps flowed from the central position of its home. That such a disposition would facilitate a centrifugal migration of land organisms is at any rate obvious, and fluctuating conditions of climate operating from the pole would supply an effective means of propulsion. As these became more rigorous animals at any rate would move southwards to escape them. It would be equally the case with plants if no insuperable obstacle interposed. This implies a mobility in plants, notwithstanding what we know of means of transport which is at first sight paradoxical. Bentham has stated this in a striking way: "Fixed and immovable as is the individual plant, there is no class in which the race is endowed with greater facilities for the widest dispersion... Plants cast away their offspring in a dormant state, ready to be carried to any distance by those external agencies which we may deem fortuitous, but without which many a race might perish from the exhaustion of the limited spot of soil in which it is rooted." (Pres. Addr.(1869), "Proc. Linn. Soc." 1868-69, pages lxvi, lxvii.)

I have quoted this passage from Bentham because it emphasises a point which Darwin for his purpose did not find it necessary to dwell upon, though he no doubt assumed it. Dispersal to a distance is, so to speak, an accidental incident in the life of a species. Lepidium Draba, a native of South-eastern Europe, owes its prevalence in the Isle of Thanet to the disastrous Walcheren expedition; the straw-stuffing of the mattresses of the fever-stricken soldiers who were landed there was used by a farmer for manure. Sir Joseph Hooker ("Royal Institution Lecture", April 12, 1878.) tells us that landing on Lord Auckland's Island, which was uninhabited, "the first evidence I met with of its having been previously visited by man was the English chickweed; and this I traced to a mound that marked the grave of a British sailor, and that was covered with the plant, doubtless the offspring of seed that had adhered to the spade or mattock with which the grave had been dug."

Some migration from the spot where the individuals of a species have germinated is an essential provision against extinction. Their descendants otherwise would be liable to suppression by more vigorous competitors. But they would eventually be extinguished inevitably, as pointed out by Bentham, by the exhaustion of at any rate some one necessary constituent of the soil. Gilbert showed by actual analysis that the production of a "fairy ring" is simply due to the using up by the fungi of the available nitrogen in the enclosed area which continually enlarges as they seek a fresh supply on the outside margin. Anyone who cultivates a garden can easily verify the fact that every plant has some adaptation for varying degrees of seed-dispersal. It cannot be doubted that slow but persistent terrestrial migration has played an enormous part in bringing about existing plant-distribution, or that climatic changes would intensify the effect because they would force the abandonment of a former area and the occupation of a new one. We are compelled to admit that as an incident of the Glacial period a whole flora may have moved down and up a mountain side, while only some of its constituent species would be able to take advantage of means of long-distance transport.

I have dwelt on the importance of what I may call short-distance dispersal as a necessary condition of plant life, because I think it suggests the solution of a difficulty which leads Guppy to a conclusion with which I am unable to agree. But the work which he has done taken as a whole appears to me so admirable that I do so with the utmost respect. He points out, as Bentham had already done, that long-distance dispersal is fortuitous. And being so it cannot have been provided for by previous adaptation. He says (Guppy, op. cit. II. page 99.): "It is not conceivable that an organism can be adapted to conditions outside its environment." To this we must agree; but, it may be asked, do the general means of plant dispersal violate so obvious a principle? He proceeds: "The great variety of the modes of dispersal of seeds is in itself an indication that the dispersing agencies avail themselves in a hap-hazard fashion of characters and capacities that have been developed in other connections." (Loc. cit. page 102.) "Their utility in these respects is an accident in the plant's life." (Loc. cit. page 100.) He attributes this utility to a "determining agency," an influence which constantly reappears in various shapes in the literature of Evolution and is ultra-scientific in the sense that it bars the way to the search for material causes. He goes so far as to doubt whether fleshy fruits are an adaptation for the dispersal of their contained seeds. (Loc. cit. page 102.) Writing as I am from a hillside which is covered by hawthorn bushes sown by birds, I confess I can feel little doubt on the subject myself. The essential fact which Guppy brings out is that long-distance unlike short-distance dispersal is not universal and purposeful, but selective and in that sense accidental. But it is not difficult to see how under favouring conditions one must merge into the other.

Guppy has raised one novel point which can only be briefly referred to but which is of extreme interest. There are grounds for thinking that flowers and insects have mutually reacted upon one another in their evolution. Guppy suggests that something of the same kind may be true of birds. I must content myself with the quotation of a single sentence. "With the secular drying of the globe and the consequent differentiation of climate is to be connected the suspension to a great extent of the agency of birds as plant dispersers in later ages, not only in the Pacific Islands but all over the tropics. The changes of climate, birds and plants have gone on together, the range of the bird being controlled by the climate, and the distribution of the plant being largely dependent on the bird." (Loc.cit. II. page 221.)

Darwin was clearly prepared to go further than Hooker in accounting for the southern flora by dispersion from the north. Thus he says: "We must, I suppose, admit that every yard of land has been successively covered with a beech-forest between the Caucasus and Japan." ("More Letters", II. page 9.) Hooker accounted for the dissevered condition of the southern flora by geographical change, but this Darwin could not admit. He suggested to Hooker that the Australian and Cape floras might have had a point of connection through Abyssinia (Ibid. I. page 447.), an idea which was promptly snuffed out. Similarly he remarked to Bentham (1869): "I suppose you think that the Restiaceae, Proteaceae, etc., etc. once extended over the whole world, leaving fragments in the south." (Ibid. I. page 380.) Eventually he conjectured "that there must have been a Tertiary Antarctic continent, from which various forms radiated to the southern extremities of our present continents." ("Life and Letters", III. page 231.) But characteristically he could not admit any land connections and trusted to "floating ice for transporting seed." ("More Letters", I. page 116.) I am far from saying that this theory is not deserving of serious attention, though there seems to be no positive evidence to support it, and it immediately raises the difficulty how did such a continent come to be stocked?

We must, however, agree with Hooker that the common origin of the northern and southern floras must be referred to a remote past. That Darwin had this in his mind at the time of the publication of the "Origin" is clear from a letter to Hooker. "The view which I should have looked at as perhaps most probable (though it hardly differs from yours) is that the whole world during the Secondary ages was inhabited by marsupials, araucarias (Mem.--Fossil wood of this nature in South America), Banksia, etc.; and that these were supplanted and exterminated in the greater area of the north, but were left alive in the south." (Ibid. I. page 453.) Remembering that Araucaria, unlike Banksia, belongs to the earlier Jurassic not to the angiospermous flora, this view is a germinal idea of the widest generality.

The extraordinary congestion in species of the peninsulas of the Old World points to the long-continued action of a migration southwards. Each is in fact a cul-de-sac into which they have poured and from which there is no escape. On the other hand the high degree of specialisation in the southern floras and the little power the species possess of holding their own in competition or in adaptation to new conditions point to long-continued isolation. "An island... will prevent free immigration and competition, hence a greater number of ancient forms will survive." (Ibid. I. page 481.) But variability is itself subject to variation. The nemesis of a high degree of protected specialisation is the loss of adaptability. (See Lyell, "The Geological Evidences of the Antiquity of Man", London, 1863, page 446.) It is probable that many elements of the southern flora are doomed: there is, for example, reason to think that the singular Stapelieae of S. Africa are a disappearing group. The tree Lobelias which linger in the mountains of Central Africa, in Tropical America and in the Sandwich Islands have the aspect of extreme antiquity. I may add a further striking illustration from Professor Seward: "The tall, graceful fronds of Matonia pectinata, forming miniature forests on the slopes of Mount Ophir and other districts in the Malay Peninsula in association with Dipteris conjugata and Dipteris lobbiana, represent a phase of Mesozoic life which survives 'Like a dim picture of the drowned past.'" ("Report of the 73rd Meeting of the British Assoc." (Southport, 1903), London, 1904, page 844.)

The Matonineae are ferns with an unusually complex vascular system and were abundant "in the northern hemisphere during the earlier part of the Mesozoic era."

It was fortunate for science that Wallace took up the task which his colleague had abandoned. Writing to him on the publication of his "Geographical Distribution of Animals" Darwin said: "I feel sure that you have laid a broad and safe foundation for all future work on Distribution. How interesting it will be to see hereafter plants treated in strict relation to your views." ("More Letters", II. page 12.) This hope was fulfilled in "Island Life". I may quote a passage from it which admirably summarises the contrast between the northern and the southern floras.

"Instead of the enormous northern area, in which highly organised and dominant groups of plants have been developed gifted with great colonising and aggressive powers, we have in the south three comparatively small and detached areas, in which rich floras have been developed with SPECIAL adaptations to soil, climate, and organic environment, but comparatively impotent and inferior beyond their own domain." (Wallace, "Island Life", pages 527, 528.)

It will be noticed that in the summary I have attempted to give of the history of the subject, efforts have been concentrated on bringing into relation the temperate floras of the northern and southern hemispheres, but no account has been taken of the rich tropical vegetation which belts the world and little to account for the original starting-point of existing vegetation generally. It must be remembered on the one hand that our detailed knowledge of the floras of the tropics is still very incomplete and far inferior to that of temperate regions; on the other hand palaeontological discoveries have put the problem in an entirely new light. Well might Darwin, writing to Heer in 1875, say: "Many as have been the wonderful discoveries in Geology during the last half-century, I think none have exceeded in interest your results with respect to the plants which formerly existed in the arctic regions." ("More Letters", II. page 240.)

As early as 1848 Debey had described from the Upper Cretaceous rocks of Aix-la-Chapelle Flowering plants of as high a degree of development as those now existing. The fact was commented upon by Hooker ("Introd. Essay to the Flora of Tasmania", page xx.), but its full significance seems to have been scarcely appreciated. For it implied not merely that their evolution must have taken place but the foundations of existing distribution must have been laid in a preceding age. We now know from the discoveries of the last fifty years that the remains of the Neocomian flora occur over an area extending through 30 deg of latitude. The conclusion is irresistible that within this was its centre of distribution and probably of origin.

Darwin was immensely impressed with the outburst on the world of a fully fledged angiospermous vegetation. He warmly approved the brilliant theory of Saporta that this happened "as soon (as) flower-frequenting insects were developed and favoured intercrossing." ("More Letters", II. page 21.) Writing to him in 1877 he says: "Your idea that dicotyledonous plants were not developed in force until sucking insects had been evolved seems to me a splendid one. I am surprised that the idea never occurred to me, but this is always the case when one first hears a new and simple explanation of some mysterious phenomenon." ("Life and Letters", III. page 285. Substantially the same idea had occurred earlier to F.W.A. Miquel. Remarking that "sucking insects (Haustellata)... perform in nature the important duty of maintaining the existence of the vegetable kingdom, at least as far as the higher orders are concerned," he points our that "the appearance in great numbers of haustellate insects occurs at and after the Cretaceous epoch, when the plants with pollen and closed carpels (Angiosperms) are found, and acquire little by little the preponderance in the vegetable kingdom." "Archives Neerlandaises", III. (1868). English translation in "Journ. of Bot." 1869, page 101.)

Even with this help the abruptness still remains an almost insoluble problem, though a forecast of floral structure is now recognised in some Jurassic and Lower Cretaceous plants. But the gap between this and the structural complexity and diversity of angiosperms is enormous. Darwin thought that the evolution might have been accomplished during a period of prolonged isolation. Writing to Hooker (1881) he says: "Nothing is more extraordinary in the history of the Vegetable Kingdom, as it seems to me, than the APPARENTLY very sudden or abrupt development of the higher plants. I have sometimes speculated whether there did not exist somewhere during long ages an extremely isolated continent, perhaps near the South Pole." ("Life and Letters", III. page 248.)

The present trend of evidence is, however, all in favour of a northern origin for flowering plants, and we can only appeal to the imperfection of the geological record as a last resource to extricate us from the difficulty of tracing the process. But Darwin's instinct that at some time or other the southern hemisphere had played an important part in the evolution of the vegetable kingdom did not mislead him. Nothing probably would have given him greater satisfaction than the masterly summary in which Seward has brought together the evidence for the origin of the Glossopteris flora in Gondwana land.

"A vast continental area, of which remnants are preserved in Australia, South Africa and South America... A tract of enormous extent occupying an area, part of which has since given place to a southern ocean, while detached masses persist as portions of more modern continents, which have enabled us to read in their fossil plants and ice-scratched boulders the records of a lost continent, in which the Mesozoic vegetation of the northern continent had its birth." ("Encycl. Brit." (10th edition 1902), Vol. XXXI. ("Palaeobotany; Mesozoic"), page 422.) Darwin would probably have demurred on physical grounds to the extent of the continent, and preferred to account for the transoceanic distribution of its flora by the same means which must have accomplished it on land.

It must in fairness be added that Guppy's later views give some support to the conjectural existence of the "lost continent." "The distribution of the genus Dammara" (Agathis) led him to modify his earlier conclusions. He tells us:--"In my volume on the geology of Vanua Levu it was shown that the Tertiary period was an age of submergence in the Western Pacific, and a disbelief in any previous continental condition was expressed. My later view is more in accordance with that of Wichmann, who, on geological grounds, contended that the islands of the Western Pacific were in a continental condition during the Palaeozoic and Mesozoic periods, and that their submergence and subsequent emergence took place in Tertiary times." (Guppy, op. cit. II. page 304.)

The weight of the geological evidence I am unable to scrutinise. But though I must admit the possibility of some unconscious bias in my own mind on the subject, I am impressed with the fact that the known distribution of the Glossopteris flora in the southern hemisphere is precisely paralleled by that of Proteaceae and Restiaceae in it at the present time. It is not unreasonable to suppose that both phenomena, so similar, may admit of the same explanation. I confess it would not surprise me if fresh discoveries in the distribution of the Glossopteris flora were to point to the possibility of its also having migrated southwards from a centre of origin in the northern hemisphere.

Darwin, however, remained sceptical "about the travelling of plants from the north EXCEPT DURING THE TERTIARY PERIOD." But he added, "such speculations seem to me hardly scientific, seeing how little we know of the old floras." ("Life and Letters", III. page 247.) That in later geological times the south has been the grave of the weakened offspring of the aggressive north can hardly be doubted. But if we look to the Glossopteris flora for the ancestry of Angiosperms during the Secondary period, Darwin's prevision might be justified, though he has given us no clue as to how he arrived at it.

It may be true that technically Darwin was not a botanist. But in two pages of the "Origin" he has given us a masterly explanation of "the relationship, with very little identity, between the productions of North America and Europe." (Pages 333, 334.) He showed that this could be accounted for by their migration southwards from a common area, and he told Wallace that he "doubted much whether the now called Palaearctic and Neartic regions ought to be separated." ("Life and Letters", III. page 230.) Catkin-bearing deciduous trees had long been seen to justify Darwin's doubt: oaks, chestnuts, beeches, hazels, hornbeams, birches, alders, willows and poplars are common both to the Old and New World. Newton found that the separate regions could not be sustained for birds, and he is now usually followed in uniting them as the Holartic. One feels inclined to say in reading the two pages, as Lord Kelvin did to a correspondent who asked for some further development of one of his papers, It is all there. We have only to apply the principle to previous geological ages to understand why the flora of the Southern United States preserves a Cretaceous facies. Applying it still further we can understand why, when the northern hemisphere gradually cooled through the Tertiary period, the plants of the Eocene "suggest a comparison of the climate and forests with those of the Malay Archipelago and Tropical America." (Clement Reid, "Encycl. Brit." (10th edition), Vol. XXXI. ("Palaeobotany; Tertiary"), page 435.) Writing to Asa Gray in 1856 with respect to the United States flora, Darwin said that "nothing has surprised me more than the greater generic and specific affinity with East Asia than with West America." ("More Letters", I. page 434.) The recent discoveries of a Tulip tree and a Sassafras in China afford fresh illustrations. A few years later Asa Gray found the explanation in both areas being centres of preservation of the Cretaceous flora from a common origin. It is interesting to note that the paper in which this was enunciated at once established his reputation.

In Europe the latitudinal range of the great mountain chains gave the Miocene flora no chance of escape during the Glacial period, and the Mediterranean appears to have equally intercepted the flow of alpine plants to the Atlas. (John Ball in Appendix G, page 438, in "Journal of a Tour in Morocco and the Great Atlas", J.D. Hooker and J. Ball, London, 1878.) In Southern Europe the myrtle, the laurel, the fig and the dwarf-palm are the sole representatives of as many great tropical families. Another great tropical family, the Gesneraceae has left single representatives from the Pyrenees to the Balkans; and in the former a diminutive yam still lingers. These are only illustrations of the evidence which constantly accumulates and which finds no rational explanation except that which Darwin has given to it.

The theory of southward migration is the key to the interpretation of the geographical distribution of plants. It derived enormous support from the researches of Heer and has now become an accepted commonplace. Saporta in 1888 described the vegetable kingdom as "emigrant pour suivre une direction determinee et marcher du nord au sud, a la recherche de regions et de stations plus favorables, mieux appropriees aux adaptations acquises, a meme que la temperature terrestre perd ses conditions premieres." ("Origine Paleontologique des arbres", Paris, 1888, page 28.) If, as is so often the case, the theory now seems to be a priori inevitable, the historian of science will not omit to record that the first germ sprang from the brain of Darwin.

In attempting this sketch of Darwin's influence on Geographical Distribution, I have found it impossible to treat it from an external point of view. His interest in it was unflagging; all I could say became necessarily a record of that interest and could not be detached from it. He was in more or less intimate touch with everyone who was working at it. In reading the letters we move amongst great names. With an extraordinary charm of persuasive correspondence he was constantly suggesting, criticising and stimulating. It is hardly an exaggeration to say that from the quiet of his study at Down he was founding and directing a wide-world school.

POSTSCRIPTUM.

Since this essay was put in type Dr Ernst's striking account of the "New Flora of the Volcanic Island of Krakatau" (Cambridge, 1909.) has reached me. All botanists must feel a debt of gratitude to Prof. Seward for his admirable translation of a memoir which in its original form is practically unprocurable and to the liberality of the Cambridge University Press for its publication. In the preceding pages I have traced the laborious research by which the methods of Plant Dispersal were established by Darwin. In the island of Krakatau nature has supplied a crucial experiment which, if it had occurred earlier, would have at once secured conviction of their efficiency. A quarter of a century ago every trace of organic life in the island was "destroyed and buried under a thick covering of glowing stones." Now, it is "again covered with a mantle of green, the growth being in places so luxuriant that it is necessary to cut one's way laboriously through the vegetation." (Op. cit. page 4.) Ernst traces minutely how this has been brought about by the combined action of wind, birds and sea currents, as means of transport. The process will continue, and he concludes:--"At last after a long interval the vegetation on the desolated island will again acquire that wealth of variety and luxuriance which we see in the fullest development which Nature has reached in the primaeval forest in the tropics." (Op. cit. page 72.) The possibility of such a result revealed itself to the insight of Darwin with little encouragement or support from contemporary opinion.

One of the most remarkable facts established by Ernst is that this has not been accomplished by the transport of seeds alone. "Tree stems and branches played an important part in the colonisation of Krakatau by plants and animals. Large piles of floating trees, stems, branches and bamboos are met with everywhere on the beach above high-water mark and often carried a considerable distance inland. Some of the animals on the island, such as the fat Iguana (Varanus salvator) which suns itself in the beds of streams, may have travelled on floating wood, possibly also the ancestors of the numerous ants, but certainly plants." (Op. cit. page 56.) Darwin actually had a prevision of this. Writing to Hooker he says:--"Would it not be a prodigy if an unstocked island did not in the course of ages receive colonists from coasts whence the currents flow, trees are drifted and birds are driven by gales?" ("More Letters", I. page 483.) And ten years earlier:--"I must believe in the... whole plant or branch being washed into the sea; with floods and slips and earthquakes; this must continually be happening." ("Life and Letters", II. pages 56, 57.) If we give to "continually" a cosmic measure, can the fact be doubted? All this, in the light of our present knowledge, is too obvious to us to admit of discussion. But it seems to me nothing less than pathetic to see how in the teeth of the obsession as to continental extension, Darwin fought single-handed for what we now know to be the truth.

Guppy's heart failed him when he had to deal with the isolated case of Agathis which alone seemed inexplicable by known means of transport. But when we remember that it is a relic of the pre-Angiospermous flora, and is of Araucarian ancestry, it cannot be said that the impossibility, in so prolonged a history, of the bodily transference of cone-bearing branches or even of trees, compels us as a last resort to fall back on continental extension to account for its existing distribution.

When Darwin was in the Galapagos Archipelago, he tells us that he fancied himself "brought near to the very act of creation." He saw how new species might arise from a common stock. Krakatau shows us an earlier stage and how by simple agencies, continually at work, that stock might be supplied. It also shows us how the mixed and casual elements of a new colony enter into competition for the ground and become mutually adjusted. The study of Plant Distribution from a Darwinian standpoint has opened up a new field of research in Ecology. The means of transport supply the materials for a flora, but their ultimate fate depends on their equipment for the "struggle for existence." The whole subject can no longer be regarded as a mere statistical inquiry which has seemed doubtless to many of somewhat arid interest. The fate of every element of the earth's vegetation has sooner or later depended on its ability to travel and to hold its own under new conditions. And the means by which it has secured success is an each case a biological problem which demands and will reward the most attentive study. This is the lesson which Darwin has bequeathed to us. It is summed up in the concluding paragraph of the "Origin" ("Origin of Species" (6th edition), page 429.):--"It is interesting to contemplate a tangled bank, clothed with many plants of many kinds, with birds singing on the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and dependent upon each other in so complex a manner, have all been produced by laws acting around us."

XVII. GEOGRAPHICAL DISTRIBUTION OF ANIMALS. By Hans Gadow, M.A., Ph.D., F.R.S.

Strickland Curator and Lecturer on Zoology in the University of Cambridge.

The first general ideas about geographical distribution may be found in some of the brilliant speculations contained in Buffon's "Histoire Naturelle". The first special treatise on the subject was however written in 1777 by E.A.W. Zimmermann, Professor of Natural Science at Brunswick, whose large volume, "Specimen Zoologiae Geographicae Quadrupedum"..., deals in a statistical way with the mammals; important features of the large accompanying map of the world are the ranges of mountains and the names of hundreds of genera indicating their geographical range. In a second work he laid special stress on domesticated animals with reference to the spreading of the various races of Mankind.

In the following year appeared the "Philosophia Entomologica" by J.C. Fabricius, who was the first to divide the world into eight regions. In 1803 G.R. Treviranus ("Biologie oder Philosophie der lebenden Natur", Vol. II. Gottingen, 1803.) devoted a long chapter of his great work on "Biologie" to a philosophical and coherent treatment of the distribution of the whole animal kingdom. Remarkable progress was made in 1810 by F. Tiedemann ("Anatomie und Naturgeschichte der Vogel". Heidelberg, 1810.) of Heidelberg. Few, if any, of the many subsequent Ornithologists seem to have appreciated, or known of, the ingenious way in which Tiedemann marshalled his statistics in order to arrive at general conclusions. There are, for instance, long lists of birds arranged in accordance with their occurrence in one or more continents: by correlating the distribution of the birds with their food he concludes "that the countries of the East Indian flora have no vegetable feeders in common with America," and "that it is probably due to the great peculiarity of the African flora that Africa has few phytophagous kinds in common with other countries, whilst zoophagous birds have a far more independent, often cosmopolitan, distribution." There are also remarkable chapters on the influence of environment, distribution, and migration, upon the structure of the Birds! In short, this anatomist dealt with some of the fundamental causes of distribution.

Whilst Tiedemann restricted himself to Birds, A. Desmoulins in 1822 wrote a short but most suggestive paper on the Vertebrata, omitting the birds; he combated the view recently proposed by the entomologist Latreille that temperature was the main factor in distribution. Some of his ten main conclusions show a peculiar mixture of evolutionary ideas coupled with the conception of the stability of species: whilst each species must have started from but one creative centre, there may be several "analogous centres of creation" so far as genera and families are concerned. Countries with different faunas, but lying within the same climatic zones, are proof of the effective and permanent existence of barriers preventing an exchange between the original creative centres.

The first book dealing with the "geography and classification" of the whole animal kingdom was written by W. Swainson ("A Treatise on the Geography and Classification of Animals", Lardner's "Cabinet Cyclopaedia" London, 1835.) in 1835. He saw in the five races of Man the clue to the mapping of the world into as many "true zoological divisions," and he reconciled the five continents with his mystical quinary circles.

Lyell's "Principles of Geology" should have marked a new epoch, since in his "Elements" he treats of the past history of the globe and the distribution of animals in time, and in his "Principles" of their distribution in space in connection with the actual changes undergone by the surface of the world. But as the sub-title of his great work "Modern changes of the Earth and its inhabitants" indicates, he restricted himself to comparatively minor changes, and, emphatically believing in the permanency of the great oceans, his numerous and careful interpretations of the effect of the geological changes upon the dispersal of animals did after all advance the problem but little.

Hitherto the marine faunas had been neglected. This was remedied by E. Forbes, who established nine homozoic zones, based mainly on the study of the mollusca, the determining factors being to a great extent the isotherms of the sea, whilst the 25 provinces were given by the configuration of the land. He was followed by J.D. Dana, who, taking principally the Crustacea as a basis, and as leading factors the mean temperatures of the coldest and of the warmest months, established five latitudinal zones. By using these as divisors into an American, Afro-European, Oriental, Arctic and Antarctic realm, most of which were limited by an eastern and western land-boundary, he arrived at about threescore provinces.

In 1853 appeared L.K. Schmarda's ("Die geographische Verbreitung der Thiere", Wien, 1853.) two volumes, embracing the whole subject. Various centres of creation being, according to him, still traceable, he formed the hypothesis that these centres were originally islands, which later became enlarged and joined together to form the great continents, so that the original faunas could overlap and mix whilst still remaining pure at their respective centres. After devoting many chapters to the possible physical causes and modes of dispersal, he divided the land into 21 realms which he shortly characterises, e.g. Australia as the only country inhabited by marsupials, monotremes and meliphagous birds. Ten main marine divisions were diagnosed in a similar way. Although some of these realms were not badly selected from the point of view of being applicable to more than one class of animals, they were obviously too numerous for general purposes, and this drawback was overcome, in 1857, by P.L. Sclater. ("On the general Geographical Distribution of the members of the class Aves", "Proc. Linn. Soc." (Zoology II. 1858, pages 130-145.)) Starting with the idea, that "each species must have been created within and over the geographical area, which it now occupies," he concluded "that the most natural primary ontological divisions of the Earth's surface" were those six regions, which since their adoption by Wallace in his epoch-making work, have become classical. Broadly speaking, these six regions are equivalent to the great masses of land; they are convenient terms for geographical facts, especially since the Palaearctic region expresses the unity of Europe with the bulk of Asia. Sclater further brigaded the regions of the Old World as Palaeogaea and the two Americas as Neogaea, a fundamental mistake, justifiable to a certain extent only since he based his regions mainly upon the present distribution of the Passerine birds.

Unfortunately these six regions are not of equal value. The Indian countries and the Ethiopian region (Africa south of the Sahara) are obviously nothing but the tropical, southern continuations or appendages of one greater complex. Further, the great eastern mass of land is so intimately connected with North America that this continent has much more in common with Europe and Asia than with South America. Therefore, instead of dividing the world longitudinally as Sclater had done, Huxley, in 1868 ("On the classification and distribution of the Alectoromorphae and Heteromorphae", "Proc. Zool. Soc." 1868, page 294.), gave weighty reasons for dividing it transversely. Accordingly he established two primary divisions, Arctogaea or the North world in a wider sense, comprising Sclater's Indian, African, Palaearctic and Neartic regions; and Notogaea, the Southern world, which he divided into (1) Austro-Columbia (an unfortunate substitute for the neotropical region), (2) Australasia, and (3) New Zealand, the number of big regions thus being reduced to three but for the separation of New Zealand upon rather negative characters. Sclater was the first to accept these four great regions and showed, in 1874 ("The geographical distribution of Mammals", "Manchester Science Lectures", 1874.), that they were well borne out by the present distribution of the Mammals.

Although applicable to various other groups of animals, for instance to the tailless Amphibia and to Birds (Huxley himself had been led to found his two fundamental divisions on the distribution of the Gallinaceous birds), the combination of South America with Australia was gradually found to be too sweeping a measure. The obvious and satisfactory solution was provided by W.T. Blanford (Anniversary address (Geological Society, 1889), "Proc. Geol. Soc." 1889-90, page 67; "Quart. Journ." XLVI 1890.), who in 1890 recognised three main divisions, namely Australian, South American, and the rest, for which the already existing terms (although used partly in a new sense, as proposed by an anonymous writer in "Natural Science", III. page 289) "Notogaea," "Neogaea" and "Arctogaea" have been gladly accepted by a number of English writers.

After this historical survey of the search for larger and largest or fundamental centres of animal creation, which resulted in the mapping of the world into zoological regions and realms of after all doubtful value, we have to return to the year 1858. The eleventh and twelfth chapters of "The Origin of Species" (1859), dealing with "Geographical Distribution," are based upon a great amount of observation, experiment and reading. As Darwin's main problem was the origin of species, nature's way of making species by gradual changes from others previously existing, he had to dispose of the view, held universally, of the independent creation of each species and at the same time to insist upon a single centre of creation for each species; and in order to emphasise his main point, the theory of descent, he had to disallow convergent, or as they were then called, analogous forms. To appreciate the difficulty of his position we have to take the standpoint of fifty years ago, when the immutability of the species was an axiom and each was supposed to have been created within or over the geographical area which it now occupies. If he once admitted that a species could arise from many individuals instead of from one pair, there was no way of shutting the door against the possibility that these individuals may have been so numerous that they occupied a very large district, even so large that it had become as discontinuous as the distribution of many a species actually is. Such a concession would at once be taken as an admission of multiple, independent, origin instead of descent in Darwin's sense.

For the so-called multiple, independently repeated creation of species as an explanation of their very wide and often quite discontinuous distribution, he substituted colonisation from the nearest and readiest source together with subsequent modification and better adaptation to their new home.

He was the first seriously to call attention to the many accidental means, "which more properly should be called occasional means of distribution," especially to oceanic islands. His specific, even individual, centres of creation made migrations all the more necessary, but their extent was sadly baulked by the prevailing dogma of the permanency of the oceans. Any number of small changes ("many islands having existed as halting places, of which not a wreck now remains" ("The Origin of Species" (1st edition), page 396.).) were conceded freely, but few, if any, great enough to permit migration of truly terrestrial creatures. The only means of getting across the gaps was by the principle of the "flotsam and jetsam," a theory which Darwin took over from Lyell and further elaborated so as to make it applicable to many kinds of plants and animals, but sadly deficient, often grotesque, in the case of most terrestrial creatures.

Another very fertile source was Darwin's strong insistence upon the great influence which the last glacial epoch must have had upon the distribution of animals and plants. Why was the migration of northern creatures southwards of far-reaching and most significant importance? More northerners have established themselves in southern lands than vice versa, because there is such a great mass of land in the north and greater continents imply greater intensity of selection. "The productions of real islands have everywhere largely yielded to continental forms." (Ibid. page 380.)... "The Alpine forms have almost everywhere largely yielded to the more dominant forms generated in the larger areas and more efficient workshops of the North."

Let us now pass in rapid survey the influence of the publication of "The Origin of Species" upon the study of Geographical Distribution in its wider sense.

Hitherto the following thought ran through the minds of most writers: Wherever we examine two or more widely separated countries their respective faunas are very different, but where two faunas can come into contact with each other, they intermingle. Consequently these faunas represent centres of creation, whence the component creatures have spread peripherally so far as existing boundaries allowed them to do so. This is of course the fundamental idea of "regions." There is not one of the numerous writers who considered the possibility that these intermediate belts might represent not a mixture of species but transitional forms, the result of changes undergone by the most peripheral migrants in adaptation to their new surroundings. The usual standpoint was also that of Pucheran ("Note sur l'equateur zoologique", "Rev. et Mag. de Zoologie", 1855; also several other papers, ibid. 1865, 1866, and 1867.) in 1855. But what a change within the next ten years! Pucheran explains the agreement in coloration between the desert and its fauna as "une harmonie post-etablie"; the Sahara, formerly a marine basin, was peopled by immigrants from the neighbouring countries, and these new animals adapted themselves to the new environment. He also discusses, among other similar questions, the Isthmus of Panama with regard to its having once been a strait. From the same author may be quoted the following passage as a strong proof of the new influence: "By the radiation of the contemporaneous faunas, each from one centre, whence as the various parts of the world successively were formed and became habitable, they spread and became modified according to the local physical conditions."

The "multiple" origin of each species as advocated by Sclater and Murray, although giving the species a broader basis, suffered from the same difficulties. There was only one alternative to the old orthodox view of independent creation, namely the bold acceptance of land-connections to an extent for which geological and palaeontological science was not yet ripe. Those who shrank from either view, gave up the problem as mysterious and beyond the human intellect. This was the expressed opinion of men like Swainson, Lyell and Humboldt. Only Darwin had the courage to say that the problem was not insoluble. If we admit "that in the long course of time the individuals of the same species, and likewise of allied species, have proceeded from some one source; then I think all the grand leading facts of geographical distribution are explicable on the theory of migration... together with subsequent modification and the multiplication of new forms." We can thus understand how it is that in some countries the inhabitants "are linked to the extinct beings which formerly inhabited the same continent." We can see why two areas, having nearly the same physical conditions, should often be inhabited by very different forms of life,... and "we can see why in two areas, however distant from each other, there should be a correlation, in the presence of identical species... and of distinct but representative species." ("The Origin of Species" (1st edition), pages 408, 409.)

Darwin's reluctance to assume great geological changes, such as a land-connection of Europe with North America, is easily explained by the fact that he restricted himself to the distribution of the present and comparatively recent species. "I do not believe that it will ever be proved that within the recent period continents which are now quite separate, have been continuously, or almost continuously, united with each other, and with the many existing oceanic islands." (Ibid. page 357.) Again, "believing... that our continents have long remained in nearly the same relative position, though subjected to large, but partial oscillations of level," that means to say within the period of existing species, or "within the recent period." (Ibid. page. 370.) The difficulty was to a great extent one of his own making. Whilst almost everybody else believed in the immutability of the species, which implies an enormous age, logically since the dawn of creation, to him the actually existing species as the latest results of evolution, were necessarily something very new, so young that only the very latest of the geological epochs could have affected them. It has since come to our knowledge that a great number of terrestrial "recent" species, even those of the higher classes of Vertebrates, date much farther back than had been thought possible. Many of them reach well into the Miocene, a time since which the world seems to have assumed the main outlines of the present continents.

In the year 1866 appeared A. Murray's work on the "Geographical Distribution of Mammals", a book which has perhaps received less recognition than it deserves. His treatment of the general introductory questions marks a considerable advance of our problem, although, and partly because, he did not entirely agree with Darwin's views as laid down in the first edition of "The Origin of Species", which after all was the great impulse given to Murray's work. Like Forbes he did not shrink from assuming enormous changes in the configuration of the continents and oceans because the theory of descent, with its necessary postulate of great migrations, required them. He stated, for instance, "that a Miocene Atlantis sufficiently explains the common distribution of animals and plants in Europe and America up to the glacial epoch." And next he considers how, and by what changes, the rehabilitation and distribution of these lands themselves were effected subsequent to that period. Further, he deserves credit for having cleared up a misunderstanding of the idea of specific centres of creation. Whilst for instance Schmarda assumed without hesitation that the same species, if occurring at places separated by great distances, or apparently insurmountable barriers, had been there created independently (multiple centres), Lyell and Darwin held that each species had only one single centre, and with this view most of us agree, but their starting point was to them represented by one individual, or rather one single pair. According to Murray, on the other hand, this centre of a species is formed by all the individuals of a species, all of which equally undergo those changes which new conditions may impose upon them. In this respect a new species has a multiple origin, but this in a sense very different from that which was upheld by L. Agassiz. As Murray himself puts it: "To my multiple origin, communication and direct derivation is essential. The species is compounded of many influences brought together through many individuals, and distilled by Nature into one species; and, being once established it may roam and spread wherever it finds the conditions of life not materially different from those of its original centre." (Murray, "The Geographical Distribution of Mammals", page 14. London, 1866.) This declaration fairly agrees with more modern views, and it must be borne in mind that the application of the single-centre principle to the genera, families and larger groups in the search for descent inevitably leads to one creative centre for the whole animal kingdom, a condition as unwarrantable as the myth of Adam and Eve being the first representatives of Mankind.

It looks as if it had required almost ten years for "The Origin of Species" to show its full effect, since the year 1868 marks the publication of Haeckel's "Naturliche Schoepfungsgeschichte" in addition to other great works. The terms "Oecology" (the relation of organisms to their environment) and "Chorology" (their distribution in space) had been given us in his "Generelle Morphologie" in 1866. The fourteenth chapter of the "History of Creation" is devoted to the distribution of organisms, their chorology, with the emphatic assertion that "not until Darwin can chorology be spoken of as a separate science, since he supplied the acting causes for the elucidation of the hitherto accumulated mass of facts." A map (a "hypothetical sketch") shows the monophyletic origin and the routes of distribution of Man.

Natural Selection may be all-mighty, all-sufficient, but it requires time, so much that the countless aeons required for the evolution of the present fauna were soon felt to be one of the most serious drawbacks of the theory. Therefore every help to ease and shorten this process should have been welcomed. In 1868 M. Wagner (The first to formulate clearly the fundamental idea of a theory of migration and its importance in the origin of new species was L. von Buch, who in his "Physikalische Beschreibung der Canarischen Inseln", written in 1825, wrote as follows: "Upon the continents the individuals of the genera by spreading far, form, through differences of the locality, food and soil, varieties which finally become constant as new species, since owing to the distances they could never be crossed with other varieties and thus be brought back to the main type. Next they may again, perhaps upon different roads, return to the old home where they find the old type likewise changed, both having become so different that they can interbreed no longer. Not so upon islands, where the individuals shut up in narrow valleys or within narrow districts, can always meet one another and thereby destroy every new attempt towards the fixing of a new variety." Clearly von Buch explains here why island types remain fixed, and why these types themselves have become so different from their continental congeners.--Actually von Buch is aware of a most important point, the difference in the process of development which exists between a new species b, which is the result of an ancestral species a having itself changed into b and thereby vanished itself, and a new species c which arose through separation out of the same ancestral a, which itself persists as such unaltered. Von Buch's prophetic view seems to have escaped Lyell's and even Wagner's notice.) came to the rescue with his "Darwin'sche Theorie und das Migrations-Gesetz der Organismen". (Leipzig, 1868.) He shows that migration, i.e. change of locality, implies new environmental conditions (never mind whether these be new stimuli to variation, or only acting as their selectors or censors), and moreover secures separation from the original stock and thus eliminates or lessens the reactionary dangers of panmixia. Darwin accepted Wagner's theory as "advantageous." Through the heated polemics of the more ardent selectionists Wagner's theory came to grow into an alternative instead of a help to the theory of selectional evolution. Separation is now rightly considered a most important factor by modern students of geographical distribution.

For the same year, 1868, we have to mention Huxley, whose Arctogaea and Notogaea are nothing less than the reconstructed main masses of land of the Mesozoic period. Beyond doubt the configuration of land at that remote period has left recognisable traces in the present continents, but whether they can account for the distribution of such a much later group as the Gallinaceous birds is more than questionable. In any case he took for his text a large natural group of birds, cosmopolitan as a whole, but with a striking distribution. The Peristeropodes, or pigeon-footed division, are restricted to the Australian and Neotropical regions, in distinction to the Alectoropodes (with the hallux inserted at a level above the front toes) which inhabit the whole of the Arctogaea, only a few members having spread into the South World. Further, as Asia alone has its Pheasants and allies, so is Africa characterised by its Guinea-fowls and relations, America has the Turkey as an endemic genus, and the Grouse tribe in a wider sense has its centre in the holarctic region: a splendid object lesson of descent, world-wide spreading and subsequent differentiation. Huxley, by the way, was the first--at least in private talk--to state that it will be for the morphologist, the well-trained anatomist, to give the casting vote in questions of geographical distribution, since he alone can determine whether we have to deal with homologous, or analogous, convergent, representative forms.

It seems late to introduce Wallace's name in 1876, the year of the publication of his standard work. ("The Geographical Distribution of Animals", 2 vols. London, 1876.) We cannot do better than quote the author's own words, expressing the hope that his "book should bear a similar relation to the eleventh and twelfth chapters of the "Origin of Species" as Darwin's "Animals and Plants under Domestication" does to the first chapter of that work," and to add that he has amply succeeded. Pleading for a few primary centres he accepts Sclater's six regions and does not follow Huxley's courageous changes which Sclater himself had accepted in 1874. Holding the view of the permanence of the oceans he accounts for the colonisation of outlying islands by further elaborating the views of Lyell and Darwin, especially in his fascinating "Island Life", with remarkable chapters on the Ice Age, Climate and Time and other fundamental factors. His method of arriving at the degree of relationship of the faunas of the various regions is eminently statistical. Long lists of genera determine by their numbers the affinity and hence the source of colonisation. In order to make sure of his material he performed the laborious task of evolving a new classification of the host of Passerine birds. This statistical method has been followed by many authors, who, relying more upon quantity than quality, have obscured the fact that the key to the present distribution lies in the past changes of the earth's surface. However, with Wallace begins the modern study of the geographical distribution of animals and the sudden interest taken in this subject by an ever widening circle of enthusiasts far beyond the professional brotherhood.

A considerable literature has since grown up, almost bewildering in its range, diversity of aims and style of procedure. It is a chaos, with many paths leading into the maze, but as yet very few take us to a position commanding a view of the whole intricate terrain with its impenetrable tangle and pitfalls.

One line of research, not initiated but greatly influenced by Wallace's works, became so prominent as to almost constitute a period which may be characterised as that of the search by specialists for either the justification or the amending of his regions. As class after class of animals was brought up to reveal the secret of the true regions, some authors saw in their different results nothing but the faultiness of previously established regions; others looked upon eventual agreements as their final corroboration, especially when for instance such diverse groups as mammals and scorpions could, with some ingenuity, be made to harmonise. But the obvious result of all these efforts was the growing knowledge that almost every class seemed to follow principles of its own. The regions tallied neither in extent nor in numbers, although most of them gravitated more and more towards three centres, namely Australia, South America and the rest of the world. Still zoologists persisted in the search, and the various modes and capabilities of dispersal of the respective groups were thought sufficient explanation of the divergent results in trying to bring the mapping of the world under one scheme.

Contemporary literature is full of devices for the mechanical dispersal of animals. Marine currents, warm and cold, were favoured all the more since they showed the probable original homes of the creatures in question. If these could not stand sea-water, they floated upon logs or icebergs, or they were blown across by storms; fishes were lifted over barriers by waterspouts, and there is on record even an hypothetical land tortoise, full of eggs, which colonised an oceanic island after a perilous sea voyage upon a tree trunk. Accidents will happen, and beyond doubt many freaks of discontinuous distribution have to be accounted for by some such means. But whilst sufficient for the scanty settlers of true oceanic islands, they cannot be held seriously to account for the rich fauna of a large continent, over which palaeontology shows us that the immigrants have passed like waves. It should also be borne in mind that there is a great difference between flotsam and jetsam. A current is an extension of the same medium and the animals in it may suffer no change during even a long voyage, since they may be brought from one litoral to another where they will still be in the same or but slightly altered environment. But the jetsam is in the position of a passenger who has been carried off by the wrong train. Almost every year some American land birds arrive at our western coasts and none of them have gained a permanent footing although such visits must have taken place since prehistoric times. It was therefore argued that only those groups of animals should be used for locating and defining regions which were absolutely bound to the soil. This method likewise gave results not reconcilable with each other, even when the distribution of fossils was taken into account, but it pointed to the absolute necessity of searching for former land-connections regardless of their extent and the present depths to which they may have sunk.

That the key to the present distribution lies in the past had been felt long ago, but at last it was appreciated that the various classes of animals and plants have appeared in successive geological epochs and also at many places remote from each other. The key to the distribution of any group lies in the configuration of land and water of that epoch in which it made its first appearance. Although this sounds like a platitude, it has frequently been ignored. If, for argument's sake, Amphibia were evolved somewhere upon the great southern land-mass of Carboniferous times (supposed by some to have stretched from South America across Africa to Australia), the distribution of this developing class must have proceeded upon lines altogether different from that of the mammals which dated perhaps from lower Triassic times, when the old south continental belt was already broken up. The broad lines of this distribution could never coincide with that of the other, older class, no matter whether the original mammalian centre was in the Afro-Indian, Australian, or Brazilian portion. If all the various groups of animals had come into existence at the same time and at the same place, then it would be possible, with sufficient geological data, to construct a map showing the generalised results applicable to the whole animal kingdom. But the premises are wrong. Whatever regions we may seek to establish applicable to all classes, we are necessarily mixing up several principles, namely geological, historical, i.e. evolutionary, with present day statistical facts. We might as well attempt one compound picture representing a chick's growth into an adult bird and a child's growth into manhood.

In short there are no general regions, not even for each class separately, unless this class be one which is confined to a comparatively short geological period. Most of the great classes have far too long a history and have evolved many successive main groups. Let us take the mammals. Marsupials live now in Australia and in both Americas, because they already existed in Mesozoic times; Ungulata existed at one time or other all over the world except in Australia, because they are post-Cretaceous; Insectivores, although as old as any Placentalia, are cosmopolitan excepting South America and Australia; Stags and Bears, as examples of comparatively recent Arctogaeans, are found everywhere with the exception of Ethiopia and Australia. Each of these groups teaches a valuable historical lesson, but when these are combined into the establishment of a few mammalian "realms," they mean nothing but statistical majorities. If there is one at all, Australia is such a realm backed against the rest of the world, but as certainly it is not a mammalian creative centre!

Well then, if the idea of generally applicable regions is a mare's nest, as was the search for the Holy Grail, what is the object of the study of geographical distribution? It is nothing less than the history of the evolution of life in space and time in the widest sense. The attempt to account for the present distribution of any group of organisms involves the aid of every branch of science. It bids fair to become a history of the world. It started in a mild, statistical way, restricting itself to the present fauna and flora and to the present configuration of land and water. Next came Oceanography concerned with the depths of the seas, their currents and temperatures; then enquiries into climatic changes, culminating in irreconcilable astronomical hypotheses as to glacial epochs; theories about changes of the level of the seas, mainly from the point of view of the physicist and astronomer. Then came more and more to the front the importance of the geological record, hand in hand with the palaeontological data and the search for the natural affinities, the genetic system of the organisms. Now and then it almost seems as if the biologists had done their share by supplying the problems and that the physicists and geologists would settle them, but in reality it is not so. The biologists not only set the problems, they alone can check the offered solutions. The mere fact of palms having flourished in Miocene Spitzbergen led to an hypothetical shifting of the axis of the world rather than to the assumption, by way of explanation, that the palms themselves might have changed their nature. One of the most valuable aids in geological research, often the only means for reconstructing the face of the earth in by-gone periods, is afforded by fossils, but only the morphologist can pronounce as to their trustworthiness as witnesses, because of the danger of mistaking analogous for homologous forms. This difficulty applies equally to living groups, and it is so important that a few instances may not be amiss.

There is undeniable similarity between the faunas of Madagascar and South America. This was supported by the Centetidae and Dendrobatidae, two entire "families," as also by other facts. The value of the Insectivores, Solenodon in Cuba, Centetes in Madagascar, has been much lessened by their recognition as an extremely ancient group and as a case of convergence, but if they are no longer put into the same family, this amendment is really to a great extent due to their widely discontinuous distribution. The only systematic difference of the Dendrobatidae from the Ranidae is the absence of teeth, morphologically a very unimportant character, and it is now agreed, on the strength of their distribution, that these little arboreal, conspicuously coloured frogs, Dendrobates in South America, Mantella in Madagascar, do not form a natural group, although a third genus, Cardioglossa in West Africa, seems also to belong to them. If these creatures lived all on the same continent, we should unhesitatingly look upon them as forming a well-defined, natural little group. On the other hand the Aglossa, with their three very divergent genera, namely Pipa in South America, Xenopus and Hymenochirus in Africa, are so well characterised as one ancient group that we use their distribution unhesitatingly as a hint of a former connection between the two continents. We are indeed arguing in vicious circles. The Ratitae as such are absolutely worthless since they are a most heterogeneous assembly, and there are untold groups, of the artificiality of which many a zoo-geographer had not the slightest suspicion when he took his statistical material, the genera and families, from some systematic catalogues or similar lists. A lamentable instance is that of certain flightless Rails, recently extinct or sub-fossil, on the isalnds of Mauritius, Rodriguez and Chatham. Being flightless they have been used in support of a former huge Antarctic continent, instead of ruling them out of court as Rails which, each in its island, have lost the power of flight, a process which must have taken place so recently that it is difficult, upon morphological grounds, to justify their separation into Aphanapteryx in Mauritius, Erythromachus in Rodriguez and Diaphorapteryx on Chatham Island. Morphologically they may well form but one genus, since they have sprung from the same stock and have developed upon the same lines; they are therefore monogenetic: but since we know that they have become what they are independently of each other (now unlike any other Rails), they are polygenetic and therefore could not form one genus in the old Darwinian sense. Further, they are not a case of convergence, since their ancestry is not divergent but leads into the same stratum.

THE RECONSTRUCTION OF THE GEOGRAPHY OF SUCCESSIVE EPOCHS.

A promising method is the study by the specialist of a large, widely distributed group of animals from an evolutionary point of view. Good examples of this method are afforded by A.E. Ortmann's ("The geographical distribution of Freshwater Decapods and its bearing upon ancient geography", "Proc. Amer. Phil. Soc." Vol. 41, 1902.) exhaustive paper and by A.W. Grabau's "Phylogeny of Fusus and its Allies" ("Smithsonian Misc. Coll." 44, 1904.) After many important groups of animals have been treated in this way--as yet sparingly attempted--the results as to hypothetical land-connections etc. are sure to be corrective and supplementary, and their problems will be solved, since they are not imaginary.

The same problems are attacked, in the reverse way, by starting with the whole fauna of a country and thence, so to speak, letting the research radiate. Some groups will be considered as autochthonous, others as immigrants, and the directions followed by them will be inquired into; the search may lead far and in various directions, and by comparison of results, by making compound maps, certain routes will assume definite shape, and if they lead across straits and seas they are warrants to search for land-connections in the past. (A fair sample of this method is C.H. Eigenmann's "The Freshwater Fishes of South and Middle America", "Popular Science Monthly", Vol. 68, 1906.) There are now not a few maps purporting to show the outlines of land and water at various epochs. Many of these attempts do not tally with each other, owing to the lamentable deficiencies of geological and fossil data, but the bolder the hypothetical outlines are drawn, the better, and this is preferable to the insertion of bays and similar detail which give such maps a fallacious look of certainty where none exists. Moreover it must be borne in mind that, when we draw a broad continental belt across an ocean, this belt need never have existed in its entirety at any one time. The features of dispersal, intended to be explained by it, would be accomplished just as well by an unknown number of islands which have joined into larger complexes while elsewhere they subsided again: like pontoon-bridges which may be opened anywhere, or like a series of superimposed dissolving views of land and sea-scapes. Hence the reconstructed maps of Europe, the only continent tolerably known, show a considerable number of islands in puzzling changes, while elsewhere, e.g. in Asia, we have to be satisfied with sweeping generalisations.

At present about half-a-dozen big connections are engaging our attention, leaving as comparatively settled the extent and the duration of such minor "bridges" as that between Africa and Madagascar, Tasmania and Australia, the Antilles and Central America, Europe and North Africa. (Not a few of those who are fascinated by, and satisfied with, the statistical aspect of distribution still have a strong dislike to the use of "bridges" if these lead over deep seas, and they get over present discontinuous occurrences by a former "universal or sub-universal distribution" of their groups.) This is indeed an easy method of cutting the knot, but in reality they shunt the question only a stage or two back, never troubling to explain how their groups managed to attain to that sub-universal range; or do they still suppose that the whole world was originally one paradise where everything lived side by side, until sin and strife and glacial epochs left nothing but scattered survivors?

The permanence of the great ocean-basins had become a dogma since it was found that a universal elevation of the land to the extent of 100 fathoms would produce but little changes, and when it was shown that even the 1000 fathom-line followed the great masses of land rather closely, and still leaving the great basins (although transgression of the sea to the same extent would change the map of the world beyond recognition), by general consent one mile was allowed as the utmost speculative limit of subsidence. Naturally two or three miles, the average depth of the oceans, seems enormous, and yet such a difference in level is as nothing in comparison with the size of the Earth. On a clay model globe ten feet in diameter an ocean bed three miles deep would scarcely be detected, and the highest mountains would be smaller than the unavoidable grains in the glazed surface of our model. There are but few countries which have not be submerged at some time or other.

CONNECTION OF SOUTH EASTERN ASIA WITH AUSTRALIA. Neumayr's Sino-Australian continent during mid-Mesozoic times was probably a much changing Archipelago, with final separations subsequent to the Cretaceous period. Henceforth Australasia was left to its own fate, but for a possible connection with the antarctic continent.

AFRICA, MADAGASCAR, INDIA. The "Lemuria" of Sclater and Haeckel cannot have been more than a broad bridge in Jurassic times; whether it was ever available for the Lemurs themselves must depend upon the time of its duration, the more recent the better, but it is difficult to show that it lasted into the Miocene.

AFRICA AND SOUTH AMERICA. Since the opposite coasts show an entire absence of marine fossils and deposits during the Mesozoic period, whilst further north and south such are known to exist and are mostly identical on either side, Neumayr suggested the existence of a great Afro-Son American mass of land during the Jurassic epoch. Such land is almost a necessity and is supported by many facts; it would easily explain the distribution of numerous groups of terrestrial creatures. Moreover to the north of this hypothetical land, somewhere across from the Antilles and Guiana to North Africa and South Western Europe, existed an almost identical fauna of Corals and Molluscs, indicating either a coast-line or a series of islands interrupted by shallow seas, just as one would expect if, and when, a Brazil-Ethiopian mass of land were breaking up. Lastly from Central America to the Mediterranean stretches one of the Tertiary tectonic lines of the geologists. Here also the great question is how long this continent lasted. Apparently the South Atlantic began to encroach from the south so that by the later Cretaceous epoch the land was reduced to a comparatively narrow Brazil-West Africa, remnants of which persisted certainly into the early Tertiary, until the South Atlantic joined across the equator with the Atlantic portion of the "Thetys," leaving what remained of South America isolated from the rest of the world.

ANTARCTIC CONNECTIONS. Patagonia and Argentina seem to have joined Antartica during the Cretaceous epoch, and this South Georgian bridge had broken down again by mid-Tertiary times when South America became consolidated. The Antarctic continent, presuming that it existed, seems also to have been joined, by way of Tasmania, with Australia, also during the Cretaceous epoch, and it is assumed that the great Australia-Antarctic-Patagonian land was severed first to the south of Tasmania and then at the South Georgian bridge. No connection, and this is important, is indicated between Antarctica and either Africa or Madagascar.

So far we have followed what may be called the vicissitudes of the great Permo-Carboniferous Gondwana land in its fullest imaginary extent, an enormous equatorial and south temperate belt from South America to Africa, South India and Australia, which seems to have provided the foundation of the present Southern continents, two of which temporarily joined Antarctica, of which however we know nothing except that it exists now.

Let us next consider the Arctic and periarctic lands. Unfortunately very little is known about the region within the arctic circle. If it was all land, or more likely great changing archipelagoes, faunistic exchange between North America, Europe and Siberia would present no difficulties, but there is one connection which engages much attention, namely a land where now lies the North temperate and Northern part of the Atlantic ocean. How far south did it ever extend and what is the latest date of a direct practicable communication, say from North Western Europe to Greenland? Connections, perhaps often interrupted, e.g. between Greenland and Labrador, at another time between Greenland and Scandinavia, seem to have existed at least since the Permo-Carboniferous epoch. If they existed also in late Cretaceous and in Tertiary times, they would of course easily explain exchanges which we know to have repeatedly taken place between America and Europe, but they are not proved thereby, since most of these exchanges can almost as easily have occurred across the polar regions, and others still more easily by repeated junction of Siberia with Alaska.

Let us now describe a hypothetical case based on the supposition of connecting bridges. Not to work in a circle, we select an important group which has not served as a basis for the reconstruction of bridges; and it must be a group which we feel justified in assuming to be old enough to have availed itself of ancient land-connections.

The occurrence of one species of Peripatus in the whole of Australia, Tasmania and New Zealand (the latter being joined to Australia by way of New Britain in Cretaceous times but not later) puts the genus back into this epoch, no unsatisfactory assumption to the morphologist. The apparent absence of Peripatus in Madagascar indicates that it did not come from the east into Africa, that it was neither Afro-Indian, nor Afro-Australian; nor can it have started in South America. We therefore assume as its creative centre Australia or Malaya in the Cretaceous epoch, whence its occurrence in Sumatra, Malay Peninsula, New Britain, New Zealand and Australia is easily explained. Then extension across Antarctica to Patagonia and Chile, whence it could spread into the rest of South America as this became consolidated in early Tertiary times. For getting to the Antilles and into Mexico it would have to wait until the Miocene, but long before that time it could arrive in Africa, there surviving as a Congolese and a Cape species. This story is unsupported by a single fossil. Peripatus may have been "sub-universal" all over greater Gondwana land in Carboniferous times, and then its absence from Madagascar would be difficult to explain, but the migrations suggested above amount to little considering that the distance from Tasmania to South America could be covered in far less time than that represented by the whole of the Eocene epoch alone.

There is yet another field, essentially the domain of geographical distribution, the cultivation of which promises fair to throw much light upon Nature's way of making species. This is the study of the organisms with regard to their environment. Instead of revealing pedigrees or of showing how and when the creatures got to a certain locality, it investigates how they behaved to meet the ever changing conditions of their habitats. There is a facies, characteristic of, and often peculiar to, the fauna of tropical moist forests, another of deserts, of high mountains, of underground life and so forth; these same facies are stamped upon whole associations of animals and plants, although these may be--and in widely separated countries generally are--drawn from totally different families of their respective orders. It does not go to the root of the matter to say that these facies have been brought about by the extermination of all the others which did not happen to fit into their particular environment. One might almost say that tropical moist forests must have arboreal frogs and that these are made out of whatever suitable material happened to be available; in Australia and South America Hylidae, in Africa Ranidae, since there Hylas are absent. The deserts must have lizards capable of standing the glare, the great changes of temperature, of running over or burrowing into the loose sand. When as in America Iguanids are available, some of these are thus modified, while in Africa and Asia the Agamids are drawn upon. Both in the Damara and in the Transcaspian deserts, a Gecko has been turned into a runner upon sand!

We cannot assume that at various epochs deserts, and at others moist forests were continuous all over the world. The different facies and associations were developed at various times and places. Are we to suppose that, wherever tropical forests came into existence, amongst the stock of humivagous lizards were always some which presented those nascent variations which made them keep step with the similarly nascent forests, the overwhelming rest being eliminated? This principle would imply that the same stratum of lizards always had variations ready to fit any changed environment, forests and deserts, rocks and swamps. The study of Ecology indicates a different procedure, a great, almost boundless plasticity of the organism, not in the sense of an exuberant moulding force, but of a readiness to be moulded, and of this the "variations" are the visible outcome. In most cases identical facies are produced by heterogeneous convergences and these may seem to be but superficial, affecting only what some authors are pleased to call the physiological characters; but environment presumably affects first those parts by which the organism comes into contact with it most directly, and if the internal structures remain unchanged, it is not because these are less easily modified but because they are not directly affected. When they are affected, they too change deeply enough.

That the plasticity should react so quickly--indeed this very quickness seems to have initiated our mistaking the variations called forth for something performed--and to the point, is itself the outcome of the long training which protoplasm has undergone since its creation.

In Nature's workshop he does not succeed who has ready an arsenal of tools for every conceivable emergency, but he who can make a tool at the spur of the moment. The ordeal of the practical test is Charles Darwin's glorious conception of Natural Selection.

XVIII. DARWIN AND GEOLOGY. By J.W. Judd, C.B., LL.D., F.R.S.

(Mr Francis Darwin has related how his father occasionally came up from Down to spend a few days with his brother Erasmus in London, and, after his brother's death, with his daughter, Mrs Litchfield. On these occasions, it was his habit to arrange meetings with Huxley, to talk over zoological questions, with Hooker, to discuss botanical problems, and with Lyell to hold conversations on geology. After the death of Lyell, Darwin, knowing my close intimacy with his friend during his later years, used to ask me to meet him when he came to town, and "talk geology." The "talks" took place sometimes at Jermyn Street Museum, at other times in the Royal College of Science, South Kensington; but more frequently, after having lunch with him, at his brother's or his daughter's house. On several occasions, however, I had the pleasure of visiting him at Down. In the postscript of a letter (of April 15, 1880) arranging one of these visits, he writes: "Since poor, dear Lyell's death, I rarely have the pleasure of geological talk with anyone.")

In one of the very interesting conversations which I had with Charles Darwin during the last seven years of his life, he asked me in a very pointed manner if I were able to recall the circumstances, accidental or otherwise, which had led me to devote myself to geological studies. He informed me that he was making similar inquiries of other friends, and I gathered from what he said that he contemplated at that time a study of the causes producing SCIENTIFIC BIAS in individual minds. I have no means of knowing how far this project ever assumed anything like concrete form, but certain it is that Darwin himself often indulged in the processes of mental introspection and analysis; and he has thus fortunately left us--in his fragments of autobiography and in his correspondence--the materials from which may be reconstructed a fairly complete history of his own mental development.

There are two perfectly distinct inquiries which we have to undertake in connection with the development of Darwin's ideas on the subject of evolution:

FIRST. How, when, and under what conditions was Darwin led to a conviction that species were not immutable, but were derived from pre-existing forms?

SECONDLY. By what lines of reasoning and research was he brought to regard "natural selection" as a vera causa in the process of evolution?

It is the first of these inquiries which specially interests the geologist; though geology undoubtedly played a part--and by no means an insignificant part--in respect to the second inquiry.

When, indeed, the history comes to be written of that great revolution of thought in the nineteenth century, by which the doctrine of evolution, from being the dream of poets and visionaries, gradually grew to be the accepted creed of naturalists, the paramount influence exerted by the infant science of geology--and especially that resulting from the publication of Lyell's epoch-making work, the "Principles of Geology"--cannot fail to be regarded as one of the leading factors. Herbert Spencer in his "Autobiography" bears testimony to the effect produced on his mind by the recently published "Principles", when, at the age of twenty, he had already begun to speculate on the subject of evolution (Herbert Spencer's "Autobiography", London, 1904, Vol. I. pages 175-177.); and Alfred Russel Wallace is scarcely less emphatic concerning the part played by Lyell's teaching in his scientific education. (See "My Life; a record of Events and Opinions", London, 1905, Vol. I. page 355, etc. Also his review of Lyell's "Principles" in "Quarterly Review" (Vol. 126), 1869, pages 359-394. See also "The Darwin-Wallace Celebration by the Linnean Society" (1909), page 118.) Huxley wrote in 1887 "I owe more than I can tell to the careful study of the "Principles of Geology" in my young days." ("Science and Pseudo Science"; "Collected Essays", London, 1902, Vol. V. page 101.) As for Charles Darwin, he never tired--either in his published writings, his private correspondence or his most intimate conversations--of ascribing the awakening of his enthusiasm and the direction of his energies towards the elucidation of the problem of development to the "Principles of Geology" and the personal influence of its author. Huxley has well expressed what the author of the "Origin of Species" so constantly insisted upon, in the statements "Darwin's greatest work is the outcome of the unflinching application to Biology of the leading idea and the method applied in the "Principles" to Geology ("Proc. Roy. Soc." Vol. XLIV. (1888), page viii.; "Collected Essays" II. page 268, 1902.), and "Lyell, for others, as for myself, was the chief agent in smoothing the road for Darwin." ("Life and Letters of Charles Darwin" II. page 190.)

We propose therefore to consider, first, what Darwin owed to geology and its cultivators, and in the second place how he was able in the end so fully to pay a great debt which he never failed to acknowledge. Thanks to the invaluable materials contained in the "Life and Letters of Charles Darwin" (3 vols.) published by Mr Francis Darwin in 1887; and to "More Letters of Charles Darwin" (2 vols.) issued by the same author, in conjunction with Professor A.C. Seward, in 1903, we are permitted to follow the various movements in Darwin's mind, and are able to record the story almost entirely in his own words. (The first of these works is indicated in the following pages by the letters "L.L."; the second by "M.L.")

From the point of view of the geologist, Darwin's life naturally divides itself into four periods. In the first, covering twenty-two years, various influences were at work militating, now for and now against, his adoption of a geological career; in the second period--the five memorable years of the voyage of the "Beagle"--the ardent sportsman with some natural-history tastes, gradually became the most enthusiastic and enlightened of geologists; in the third period, lasting ten years, the valuable geological recruit devoted nearly all his energies and time to geological study and discussion and to preparing for publication the numerous observations made by him during the voyage; the fourth period, which covers the latter half of his life, found Darwin gradually drawn more and more from geological to biological studies, though always retaining the deepest interest in the progress and fortunes of his "old love." But geologists gladly recognise the fact that Darwin immeasurably better served their science by this biological work, than he could possibly have done by confining himself to purely geological questions.

From his earliest childhood, Darwin was a collector, though up to the time when, at eight years of age, he went to a preparatory school, seals, franks and similar trifles appear to have been the only objects of his quest. But a stone, which one of his schoolfellows at that time gave to him, seems to have attracted his attention and set him seeking for pebbles and minerals; as the result of this newly acquired taste, he says (writing in 1838) "I distinctly recollect the desire I had of being able to know something about every pebble in front of the hall door--it was my earliest and only geological aspiration at that time." ("M.L." I. page 3.) He further suspects that while at Mr Case's school "I do not remember any mental pursuits except those of collecting stones," etc... "I was born a naturalist." ("M.L." I. page 4.)

The court-yard in front of the hall door at the Mount House, Darwin's birthplace and the home of his childhood, is surrounded by beds or rockeries on which lie a number of pebbles. Some of these pebbles (in quite recent times as I am informed) have been collected to form a "cobbled" space in front of the gate in the outer wall, which fronts the hall door; and a similar "cobbled area," there is reason to believe, may have existed in Darwin's childhood before the door itself. The pebbles, which were obtained from a neighbouring gravel-pit, being derived from the glacial drift, exhibit very striking differences in colour and form. It was probably this circumstance which awakened in the child his love of observation and speculation. It is certainly remarkable that "aspirations" of the kind should have arisen in the mind of a child of 9 or 10!

When he went to Shrewsbury School, he relates "I continued collecting minerals with much zeal, but quite unscientifically,--all that I cared about was a new-NAMED mineral, and I hardly attempted to classify them." ("L.L." I. page 34.)

There has stood from very early times in Darwin's native town of Shrewsbury, a very notable boulder which has probably marked a boundary and is known as the "Bell-stone"--giving its name to a house and street. Darwin tells us in his "Autobiography" that while he was at Shrewsbury School at the age of 13 or 14 "an old Mr Cotton in Shropshire, who knew a good deal about rocks" pointed out to me "... the 'bell-stone'; he told me that there was no rock of the same kind nearer than Cumberland or Scotland, and he solemnly assured me that the world would come to an end before anyone would be able to explain how this stone came where it now lay"! Darwin adds "This produced a deep impression on me, and I meditated over this wonderful stone." ("L.L." I. page 41.)

The "bell-stone" has now, owing to the necessities of building, been removed a short distance from its original site, and is carefully preserved within the walls of a bank. It is a block of irregular shape 3 feet long and 2 feet wide, and about 1 foot thick, weighing probably not less than one-third of a ton. By the courtesy of the directors of the National Provincial Bank of England, I have been able to make a minute examination of it, and Professors Bonney and Watts, with Mr Harker and Mr Fearnsides have given me their valuable assistance. The rock is a much altered andesite and was probably derived from the Arenig district in North Wales, or possibly from a point nearer the Welsh Border. (I am greatly indebted to the Managers of the Bank at Shrewsbury for kind assistance in the examination of this interesting memorial: and Mr H.T. Beddoes, the Curator of the Shrewsbury Museum, has given me some archaeological information concerning the stone. Mr Richard Cotton was a good local naturalist, a Fellow both of the Geological and Linnean Societies; and to the officers of these societies I am indebted for information concerning him. He died in 1839, and although he does not appear to have published any scientific papers, he did far more for science by influencing the career of the school boy!) It was of course brought to where Shrewsbury now stands by the agency of a glacier--as Darwin afterwards learnt.

We can well believe from the perusal of these reminiscences that, at this time, Darwin's mind was, as he himself says, "prepared for a philosophical treatment of the subject" of Geology. ("L.L." I. page 41.) When at the age of 16, however, he was entered as a medical student at Edinburgh University, he not only did not get any encouragement of his scientific tastes, but was positively repelled by the ordinary instruction given there. Dr Hope's lectures on Chemistry, it is true, interested the boy, who with his brother Erasmus had made a laboratory in the toolhouse, and was nicknamed "Gas" by his schoolfellows, while undergoing solemn and public reprimand from Dr Butler at Shrewsbury School for thus wasting his time. ("L.L." I. page 35.) But most of the other Edinburgh lectures were "intolerably dull," "as dull as the professors" themselves, "something fearful to remember." In after life the memory of these lectures was like a nightmare to him. He speaks in 1840 of Jameson's lectures as something "I... for my sins experienced!" ("L.L." I. page 340.) Darwin especially signalises these lectures on Geology and Zoology, which he attended in his second year, as being worst of all "incredibly dull. The sole effect they produced on me was the determination never so long as I lived to read a book on Geology, or in any way to study the science!" ("L.L." I. page 41.)

The misfortune was that Edinburgh at that time had become the cockpit in which the barren conflict between "Neptunism" and "Plutonism" was being waged with blind fury and theological bitterness. Jameson and his pupils, on the one hand, and the friends and disciples of Hutton, on the other, went to the wildest extremes in opposing each other's peculiar tenets. Darwin tells us that he actually heard Jameson "in a field lecture at Salisbury Craigs, discoursing on a trap-dyke, with amygdaloidal margins and the strata indurated on each side, with volcanic rocks all around us, say that it was a fissure filled with sediment from above, adding with a sneer that there were men who maintained that it had been injected from beneath in a molten condition." ("L.L." I. pages 41-42.) "When I think of this lecture," added Darwin, "I do not wonder that I determined never to attend to Geology." (This was written in 1876 and Darwin had in the summer of 1839 revisited and carefully studied the locality ("L.L." I. page 290.) It is probable that most of Jameson's teaching was of the same controversial and unilluminating character as this field-lecture at Salisbury Craigs.

There can be no doubt that, while at Edinburgh, Darwin must have become acquainted with the doctrines of the Huttonian School. Though so young, he mixed freely with the scientific society of the city, Macgillivray, Grant, Leonard Horner, Coldstream, Ainsworth and others being among his acquaintances, while he attended and even read papers at the local scientific societies. It is to be feared, however, that what Darwin would hear most of, as characteristic of the Huttonian teaching, would be assertions that chalk-flints were intrusions of molten silica, that fossil wood and other petrifactions had been impregnated with fused materials, that heat--but never water--was always the agent by which the induration and crystallisation of rock-materials (even siliceous conglomerate, limestone and rock-salt) had been effected! These extravagant "anti-Wernerian" views the young student might well regard as not one whit less absurd and repellant than the doctrine of the "aqueous precipitation" of basalt. There is no evidence that Darwin, even if he ever heard of them, was in any way impressed, in his early career, by the suggestive passages in Hutton and Playfair, to which Lyell afterwards called attention, and which foreshadowed the main principles of Uniformitarianism.

As a matter of fact, I believe that the influence of Hutton and Playfair in the development of a philosophical theory of geology has been very greatly exaggerated by later writers on the subject. Just as Wells and Matthew anticipated the views of Darwin on Natural Selection, but without producing any real influence on the course of biological thought, so Hutton and Playfair adumbrated doctrines which only became the basis of vivifying theory in the hands of Lyell. Alfred Russel Wallace has very justly remarked that when Lyell wrote the "Principles of Geology", "the doctrines of Hutton and Playfair, so much in advance of their age, seemed to be utterly forgotten." ("Quarterly Review", Vol. CXXVI. (1869), page 363.) In proof of this it is only necessary to point to the works of the great masters of English geology, who preceded Lyell, in which the works of Hutton and his followers are scarcely ever mentioned. This is true even of the "Researches in Theoretical Geology" and the other works of the sagacious De la Beche. (Of the strength and persistence of the prejudice felt against Lyell's views by his contemporaries, I had a striking illustration some little time after Lyell's death. One of the old geologists who in the early years of the century had done really good work in connection with the Geological Society expressed a hope that I was not "one of those who had been carried away by poor Lyell's fads." My surprise was indeed great when further conversation showed me that the whole of the "Principles" were included in the "fads"!) Darwin himself possessed a copy of Playfair's "Illustrations of the Huttonian Theory", and occasionally quotes it; but I have met with only one reference to Hutton, and that a somewhat enigmatical one, in all Darwin's writings. In a letter to Lyell in 1841, when his mind was much exercised concerning glacial questions, he says "What a grand new feature all this ice work is in Geology! How old Hutton would have stared!" ("M.L." II. page 149.)

As a consequence of the influences brought to bear on his mind during his two years' residence in Edinburgh, Darwin, who had entered that University with strong geological aspirations, left it and proceeded to Cambridge with a pronounced distaste for the whole subject. The result of this was that, during his career as an under-graduate, he neglected all the opportunities for geological study. During that important period of life, when he was between eighteen and twenty years of age, Darwin spent his time in riding, shooting and beetle-hunting, pursuits which were undoubtedly an admirable preparation for his future work as an explorer; but in none of his letters of this period does he even mention geology. He says, however, "I was so sickened with lectures at Edinburgh that I did not even attend Sedgwick's eloquent and interesting lectures." ("L.L." I. page 48.)

It was only after passing his examination, and when he went up to spend two extra terms at Cambridge, that geology again began to attract his attention. The reading of Sir John Herschel's "Introduction to the Study of Natural Philosophy", and of Humboldt's "Personal Narrative", a copy of which last had been given to him by his good friend and mentor Henslow, roused his dormant enthusiasm for science, and awakened in his mind a passionate desire for travel. And it was from Henslow, whom he had accompanied in his excursions, but without imbibing any marked taste, at that time, for botany, that the advice came to think of and to "begin the study of geology." ("L.L." I. page 56.) This was in 1831, and in the summer vacation of that year we find him back again at Shrewsbury "working like a tiger" at geology and endeavouring to make a map and section of Shropshire--work which he says was not "as easy as I expected." ("L.L." I. page 189.) No better field for geological studies could possibly be found than Darwin's native county.

Writing to Henslow at this time, and referring to a form of the instrument devised by his friend, Darwin says: "I am very glad to say I think the clinometer will answer admirably. I put all the tables in my bedroom at every conceivable angle and direction. I will venture to say that I have measured them as accurately as any geologist going could do." But he adds: "I have been working at so many things that I have not got on much with geology. I suspect the first expedition I take, clinometer and hammer in hand, will send me back very little wiser and a good deal more puzzled than when I started." ("L.L." I. page 189.) Valuable aid was, however, at hand, for at this time Sedgwick, to whom Darwin had been introduced by the ever-helpful Henslow, was making one of his expeditions into Wales, and consented to accept the young student as his companion during the geological tour. ("L.L." I. page 56.) We find Darwin looking forward to this privilege with the keenest interest. ("L.L." I. page 189.)

When at the beginning of August (1831), Sedgwick arrived at his father's house in Shrewsbury, where he spent a night, Darwin began to receive his first and only instruction as a field-geologist. The journey they took together led them through Llangollen, Conway, Bangor, and Capel Curig, at which latter place they parted after spending many hours in examining the rocks at Cwm Idwal with extreme care, seeking for fossils but without success. Sedgwick's mode of instruction was admirable--he from time to time sent the pupil off on a line parallel to his own, "telling me to bring back specimens of the rocks and to mark the stratification on a map." ("L.L." I. page 57.) On his return to Shrewsbury, Darwin wrote to Henslow, "My trip with Sedgwick answered most perfectly," ("L.L." I. page 195.), and in the following year he wrote again from South America to the same friend, "Tell Professor Sedgwick he does not know how much I am indebted to him for the Welsh expedition; it has given me an interest in Geology which I would not give up for any consideration. I do not think I ever spent a more delightful three weeks than pounding the north-west mountains." ("L.L." I. pages 237-8.)

It would be a mistake, however, to suppose that at this time Darwin had acquired anything like the affection for geological study, which he afterwards developed. After parting with Sedgwick, he walked in a straight line by compass and map across the mountains to Barmouth to visit a reading party there, but taking care to return to Shropshire before September 1st, in order to be ready for the shooting. For as he candidly tells us, "I should have thought myself mad to give up the first days of partridge-shooting for geology or any other science!" ("L.L." I. page 58.)

Any regret we may be disposed to feel that Darwin did not use his opportunities at Edinburgh and Cambridge to obtain systematic and practical instruction in mineralogy and geology, will be mitigated, however, when we reflect on the danger which he would run of being indoctrinated with the crude "catastrophic" views of geology, which were at that time prevalent in all the centres of learning.

Writing to Henslow in the summer of 1831, Darwin says "As yet I have only indulged in hypotheses, but they are such powerful ones that I suppose, if they were put into action but for one day, the world would come to an end." ("L.L." I. page 189.)

May we not read in this passage an indication that the self-taught geologist had, even at this early stage, begun to feel a distrust for the prevalent catastrophism, and that his mind was becoming a field in which the seeds which Lyell was afterwards to sow would "fall on good ground"?

The second period of Darwin's geological career--the five years spent by him on board the "Beagle"--was the one in which by far the most important stage in his mental development was accomplished. He left England a healthy, vigorous and enthusiastic collector; he returned five years later with unique experiences, the germs of great ideas, and a knowledge which placed him at once in the foremost ranks of the geologists of that day. Huxley has well said that "Darwin found on board the "Beagle" that which neither the pedagogues of Shrewsbury, nor the professoriate of Edinburgh, nor the tutors of Cambridge had managed to give him." ("Proc. Roy. Soc." Vol. XLIV. (1888), page IX.) Darwin himself wrote, referring to the date at which the voyage was expected to begin: "My second life will then commence, and it shall be as a birthday for the rest of my life." ("L.L." I. page 214.); and looking back on the voyage after forty years, he wrote; "The voyage of the 'Beagle' has been by far the most important event in my life, and has determined my whole career;... I have always felt that I owe to the voyage the first real training or education of my mind; I was led to attend closely to several branches of natural history, and thus my powers of observation were improved, though they were always fairly developed." ("L.L." I. page 61.)

Referring to these general studies in natural history, however, Darwin adds a very significant remark: "The investigation of the geology of the places visited was far more important, as reasoning here comes into play. On first examining a new district nothing can appear more hopeless than the chaos of rocks; but by recording the stratification and nature of the rocks and fossils at many points, always reasoning and predicting what will be found elsewhere, light soon begins to dawn on the district, and the structure of the whole becomes more or less intelligible." ("L.L." I. page 62.)

The famous voyage began amid doubts, discouragements and disappointments. Fearful of heart-disease, sad at parting from home and friends, depressed by sea-sickness, the young explorer, after being twice driven back by baffling winds, reached the great object of his ambition, the island of Teneriffe, only to find that, owing to quarantine regulations, landing was out of the question.

But soon this inauspicious opening of the voyage was forgotten. Henslow had advised his pupil to take with him the first volume of Lyell's "Principles of Geology", then just published--but cautioned him (as nearly all the leaders in geological science at that day would certainly have done) "on no account to accept the views therein advocated." ("L.L." I. page 73.) It is probable that the days of waiting, discomfort and sea-sickness at the beginning of the voyage were relieved by the reading of this volume. For he says that when he landed, three weeks after setting sail from Plymouth, in St Jago, the largest of the Cape de Verde Islands, the volume had already been "studied attentively; and the book was of the highest service to me in many ways... " His first original geological work, he declares, "showed me clearly the wonderful superiority of Lyell's manner of treating geology, compared with that of any other author, whose works I had with me or ever afterwards read." ("L.L." I. page 62.)

At St Jago Darwin first experienced the joy of making new discoveries, and his delight was unbounded. Writing to his father he says, "Geologising in a volcanic country is most delightful; besides the interest attached to itself, it leads you into most beautiful and retired spots." ("L.L." I. page 228.) To Henslow he wrote of St Jago: "Here we spent three most delightful weeks... St Jago is singularly barren, and produces few plants or insects, so that my hammer was my usual companion, and in its company most delightful hours I spent." "The geology was pre-eminently interesting, and I believe quite new; there are some facts on a large scale of upraised coast (which is an excellent epoch for all the volcanic rocks to date from), that would interest Mr Lyell." ("L.L." I. page 235.) After more than forty years the memory of this, his first geological work, seems as fresh as ever, and he wrote in 1876, "The geology of St Jago is very striking, yet simple: a stream of lava formerly flowed over the bed of the sea, formed of triturated recent shells and corals, which it has baked into a hard white rock. Since then the whole island has been upheaved. But the line of white rock revealed to me a new and important fact, namely, that there had been afterwards subsidence round the craters, which had since been in action, and had poured forth lava." ("L.L." I. page 65.)

It was at this time, probably, that Darwin made his first attempt at drawing a sketch-map and section to illustrate the observations he had made (see his "Volcanic Islands", pages 1 and 9). His first important geological discovery, that of the subsidence of strata around volcanic vents (which has since been confirmed by Mr Heaphy in New Zealand and other authors) awakened an intense enthusiasm, and he writes: "It then first dawned on me that I might perhaps write a book on the geology of the various countries visited, and this made me thrill with delight. That was a memorable hour to me, and how distinctly I can call to mind the low cliff of lava beneath which I rested, with the sun glaring hot, a few strange desert plants growing near, and with living corals in the tidal pools at my feet." ("L.L." I. page 66.)

But it was when the "Beagle", after touching at St Paul's rock and Tristan d'Acunha (for a sufficient time only to collect specimens), reached the shores of South America, that Darwin's real work began; and he was able, while the marine surveys were in progress, to make many extensive journeys on land. His letters at this time show that geology had become his chief delight, and such exclamations as "Geology carries the day," "I find in Geology a never failing interest," etc. abound in his correspondence.

Darwin's time was divided between the study of the great deposits of red mud--the Pampean formation--with its interesting fossil bones and shells affording proofs of slow and constant movements of the land, and the underlying masses of metamorphic and plutonic rocks. Writing to Henslow in March, 1834, he says: "I am quite charmed with Geology, but, like the wise animal between two bundles of hay, I do not know which to like best; the old crystalline groups of rocks, or the softer and fossiliferous beds. When puzzling about stratification, etc., I feel inclined to cry 'a fig for your big oysters, and your bigger megatheriums.' But then when digging out some fine bones, I wonder how any man can tire his arms with hammering granite." ("L.L." I. page 249.) We are told by Darwin that he loved to reason about and attempt to predict the nature of the rocks in each new district before he arrived at it.

This love of guessing as to the geology of a district he was about to visit is amusingly expressed by him in a letter (of May, 1832) to his cousin and old college-friend, Fox. After alluding to the beetles he had been collecting--a taste his friend had in common with himself--he writes of geology that "It is like the pleasure of gambling. Speculating on first arriving, what the rocks may be, I often mentally cry out 3 to 1 tertiary against primitive; but the latter have hitherto won all the bets." ("L.L." I. page 233.)

Not the least important of the educational results of the voyage to Darwin was the acquirement by him of those habits of industry and method which enabled him in after life to accomplish so much--in spite of constant failures of health. From the outset, he daily undertook and resolutely accomplished, in spite of sea-sickness and other distractions, four important tasks. In the first place he regularly wrote up the pages of his Journal, in which, paying great attention to literary style and composition, he recorded only matters that would be of general interest, such as remarks on scenery and vegetation, on the peculiarities and habits of animals, and on the characters, avocations and political institutions of the various races of men with whom he was brought in contact. It was the freshness of these observations that gave his "Narrative" so much charm. Only in those cases in which his ideas had become fully crystallised, did he attempt to deal with scientific matters in this journal. His second task was to write in voluminous note-books facts concerning animals and plants, collected on sea or land, which could not be well made out from specimens preserved in spirit; but he tells us that, owing to want of skill in dissecting and drawing, much of the time spent in this work was entirely thrown away, "a great pile of MS. which I made during the voyage has proved almost useless." ("L.L." I. page 62.) Huxley confirmed this judgment on his biological work, declaring that "all his zeal and industry resulted, for the most part, in a vast accumulation of useless manuscript." ("Proc. Roy. Soc." Vol. XLIV. (1888), page IX.) Darwin's third task was of a very different character and of infinitely greater value. It consisted in writing notes of his journeys on land--the notes being devoted to the geology of the districts visited by him. These formed the basis, not only of a number of geological papers published on his return, but also of the three important volumes forming "The Geology of the voyage of the 'Beagle'". On July 24th, 1834, when little more than half of the voyage had been completed, Darwin wrote to Henslow, "My notes are becoming bulky. I have about 600 small quarto pages full; about half of this is Geology." ("M.L." I. page 14.) The last, and certainly not the least important of all his duties, consisted in numbering, cataloguing, and packing his specimens for despatch to Henslow, who had undertaken the care of them. In his letters he often expresses the greatest solicitude lest the value of these specimens should be impaired by the removal of the numbers corresponding to his manuscript lists. Science owes much to Henslow's patient care of the collections sent to him by Darwin. The latter wrote in Henslow's biography, "During the five years' voyage, he regularly corresponded with me and guided my efforts; he received, opened, and took care of all the specimens sent home in many large boxes." ("Life of Henslow", by L. Jenyns (Blomefield), London, 1862, page 53.)

Darwin's geological specimens are now very appropriately lodged for the most part in the Sedgwick Museum, Cambridge, his original Catalogue with subsequent annotations being preserved with them. From an examination of these catalogues and specimens we are able to form a fair notion of the work done by Darwin in his little cabin in the "Beagle", in the intervals between his land journeys.

Besides writing up his notes, it is evident that he was able to accomplish a considerable amount of study of his specimens, before they were packed up for despatch to Henslow. Besides hand-magnifiers and a microscope, Darwin had an equipment for blowpipe-analysis, a contact-goniometer and magnet; and these were in constant use by him. His small library of reference (now included in the Collection of books placed by Mr F. Darwin in the Botany School at Cambridge ("Catalogue of the Library of Charles Darwin now in the Botany School, Cambridge". Compiled by H.W. Rutherford; with an introduction by Francis Darwin. Cambridge, 1908.)) appears to have been admirably selected, and in all probability contained (in addition to a good many works relating to South America) a fair number of excellent books of reference. Among those relating to mineralogy, he possessed the manuals of Phillips, Alexander Brongniart, Beudant, von Kobell and Jameson: all the "Cristallographie" of Brochant de Villers and, for blowpipe work, Dr Children's translation of the book of Berzelius on the subject. In addition to these, he had Henry's "Experimental Chemistry" and Ure's "Dictionary" (of Chemistry). A work, he evidently often employed, was P. Syme's book on "Werner's Nomenclature of Colours"; while, for Petrology, he used Macculloch's "Geological Classification of Rocks". How diligently and well he employed his instruments and books is shown by the valuable observations recorded in the annotated Catalogues drawn up on board ship.

These catalogues have on the right-hand pages numbers and descriptions of the specimens, and on the opposite pages notes on the specimens--the result of experiments made at the time and written in a very small hand. Of the subsequently made pencil notes, I shall have to speak later. (I am greatly indebted to my friend Mr A. Harker, F.R.S., for his assistance in examining these specimens and catalogues. He has also arranged the specimens in the Sedgwick Museum, so as to make reference to them easy. The specimens from Ascension and a few others are however in the Museum at Jermyn Street.)

It is a question of great interest to determine the period and the occasion of Darwin's first awakening to the great problem of the transmutation of species. He tells us himself that his grandfather's "Zoonomia" had been read by him "but without producing any effect," and that his friend Grant's rhapsodies on Lamarck and his views on evolution only gave rise to "astonishment." ("L.L." I. page 38.)

Huxley, who had probably never seen the privately printed volume of letters to Henslow, expressed the opinion that Darwin could not have perceived the important bearing of his discovery of bones in the Pampean Formation, until they had been studied in England, and their analogies pronounced upon by competent comparative anatomists. And this seemed to be confirmed by Darwin's own entry in his pocket-book for 1837, "In July opened first notebook on Transmutation of Species. Had been greatly struck from about the month of previous March on character of South American fossils... " ("L.L." I. page 276.)

The second volume of Lyell's "Principles of Geology" was published in January, 1832, and Darwin's copy (like that of the other two volumes, in a sadly dilapidated condition from constant use) has in it the inscription, "Charles Darwin, Monte Video. Nov. 1832." As everyone knows, Darwin in dedicating the second edition of his Journal of the Voyage to Lyell declared, "the chief part of whatever scientific merit this journal and the other works of the author may possess, has been derived from studying the well-known and admirable 'Principles of Geology'".

In the first chapter of this second volume of the "Principles", Lyell insists on the importance of the species question to the geologist, but goes on to point out the difficulty of accepting the only serious attempt at a transmutation theory which had up to that time appeared--that of Lamarck. In subsequent chapters he discusses the questions of the modification and variability of species, of hybridity, and of the geographical distribution of plants and animals. He then gives vivid pictures of the struggle for existence, ever going on between various species, and of the causes which lead to their extinction--not by overwhelming catastrophes, but by the silent and almost unobserved action of natural causes. This leads him to consider theories with regard to the introduction of new species, and, rejecting the fanciful notions of "centres or foci of creation," he argues strongly in favour of the view, as most reconcileable with observed facts, that "each species may have had its origin in a single pair, or individual, where an individual was sufficient, and species may have been created in succession at such times and in such places as to enable them to multiply and endure for an appointed period, and occupy an appointed space on the globe." ("Principles of Geology", Vol. II. (1st edition 1832), page 124. We now know, as has been so well pointed out by Huxley, that Lyell, as early as 1827, was prepared to accept the doctrine of the transmutation of species. In that year he wrote to Mantell, "What changes species may really undergo! How impossible will it be to distinguish and lay down a line, beyond which some of the so-called extinct species may have never passed into recent ones" (Lyell's "Life and Letters" Vol. I. page 168). To Sir John Herschel in 1836, he wrote, "In regard to the origination of new species, I am very glad to find that you think it probable that it may be carried on through the intervention of intermediate causes. I left this rather to be inferred, not thinking it worth while to offend a certain class of persons by embodying in words what would only be a speculation" (Ibid. page 467). He expressed the same views to Whewell in 1837 (Ibid. Vol. II. page 5.), and to Sedgwick (Ibid. Vol. II. page 36) to whom he says, of "the theory, that the creation of new species is going on at the present day"--"I really entertain it," but "I have studiously avoided laying the doctrine down dogmatically as capable of proof" (see Huxley in "L.L." II. pages 190-195.))

After pointing out how impossible it would be for a naturalist to prove that a newly DISCOVERED species was really newly CREATED (Mr F. Darwin has pointed out that his father (like Lyell) often used the term "Creation" in speaking of the origin of new species ("L.L." II. chapter 1.)), Lyell argued that no satisfactory evidence OF THE WAY in which these new forms were created, had as yet been discovered, but that he entertained the hope of a possible solution of the problem being found in the study of the geological record.

It is not difficult, in reading these chapters of Lyell's great work, to realise what an effect they would have on the mind of Darwin, as new facts were collected and fresh observations concerning extinct and recent forms were made in his travels. We are not surprised to find him writing home, "I am become a zealous disciple of Mr Lyell's views, as known in his admirable book. Geologising in South America, I am tempted to carry parts to a greater extent even than he does." ("L.L." I. page 263.)

Lyell's anticipation that the study of the geological record might afford a clue to the discovery of how new species originate was remarkably fulfilled, within a few months, by Darwin's discovery of fossil bones in the red Pampean mud.

It is very true that, as Huxley remarked, Darwin's knowledge of comparative anatomy must have been, at that time, slight; but that he recognised the remarkable resemblances between the extinct and existing mammals of South America is proved beyond all question by a passage in his letter to Henslow, written November 24th, 1832: "I have been very lucky with fossil bones; I have fragments of at least six distinct animals... I found a large surface of osseous polygonal plates... Immediately I saw them I thought they must belong to an enormous armadillo, living species of which genus are so abundant here," and he goes on to say that he has "the lower jaw of some large animal which, from the molar teeth, I should think belonged to the Edentata." ("M.L." I. pages 11, 12. See "Extracts of Letters addressed to Prof. Henslow by C. Darwin" (1835), page 7.)

Having found this important clue, Darwin followed it up with characteristic perseverance. In his quest for more fossil bones he was indefatigable. Mr Francis Darwin tells us, "I have often heard him speak of the despair with which he had to break off the projecting extremity of a huge, partly excavated bone, when the boat waiting for him would wait no longer." ("L.L." I. page 276 (footnote).) Writing to Haeckel in 1864, Darwin says: "I shall never forget my astonishment when I dug out a gigantic piece of armour, like that of the living armadillo." (Haeckel, "History of Creation", Vol. I. page 134, London, 1876.)

In a letter to Henslow in 1834 Darwin says: "I have just got scent of some fossil bones... what they may be I do not know, but if gold or galloping will get them they shall be mine." ("M.L." I. page 15.)

Darwin also showed his sense of the importance of the discovery of these bones by his solicitude about their safe arrival and custody. From the Falkland Isles (March, 1834), he writes to Henslow: "I have been alarmed by your expression 'cleaning all the bones' as I am afraid the printed numbers will be lost: the reason I am so anxious they should not be, is, that a part were found in a gravel with recent shells, but others in a very different bed. Now with these latter there were bones of an Agouti, a genus of animals, I believe, peculiar to America, and it would be curious to prove that some one of the genus co-existed with the Megatherium: such and many other points depend on the numbers being carefully preserved." ("Extracts from Letters etc.", pages 13-14.) In the abstract of the notes read to the Geological Society in 1835, we read: "In the gravel of Patagonia he (Darwin) also found many bones of the Megatherium and of five or six other species of quadrupeds, among which he has detected the bones of a species of Agouti. He also met with several examples of the polygonal plates, etc." ("Proc. Geol. Soc." Vol. II. pages 211-212.)

Darwin's own recollections entirely bear out the conclusion that he fully recognised, WHILE IN SOUTH AMERICA, the wonderful significance of the resemblances between the extinct and recent mammalian faunas. He wrote in his "Autobiography": "During the voyage of the 'Beagle' I had been deeply impressed by discovering in the Pampean formation great fossil animals covered with armour like that on the existing armadillos." ("L.L." I. page 82.)

The impression made on Darwin's mind by the discovery of these fossil bones, was doubtless deepened as, in his progress southward from Brazil to Patagonia, he found similar species of Edentate animals everywhere replacing one another among the living forms, while, whenever fossils occurred, they also were seen to belong to the same remarkable group of animals. (While Darwin was making these observations in South America, a similar generalisation to that at which he arrived was being reached, quite independently and almost simultaneously, with respect to the fossil and recent mammals of Australia. In the year 1831, Clift gave to Jameson a list of bones occurring in the caves and breccias of Australia, and in publishing this list the latter referred to the fact that the forms belonged to marsupials, similar to those of the existing Australian fauna. But he also stated that, as a skull had been identified (doubtless erroneously) as having belonged to a hippopotamus, other mammals than marsupials must have spread over the island in late Tertiary times. It is not necessary to point out that this paper was quite unknown to Darwin while in South America. Lyell first noticed it in the third edition of his "Principles", which was published in May, 1834 (see "Edinb. New Phil. Journ." Vol. X. (1831), pages 394-6, and Lyell's "Principles" (3rd edition), Vol. III. page 421). Darwin referred to this discovery in 1839 (see his "Journal", page 210.))

That the passage in Darwin's pocket-book for 1837 can only refer to an AWAKENING of Darwin's interest in the subject--probably resulting from a sight of the bones when they were being unpacked--I think there cannot be the smallest doubt; AND WE MAY THEREFORE CONFIDENTLY FIX UPON NOVEMBER, 1832, AS THE DATE AT WHICH DARWIN COMMENCED THAT LONG SERIES OF OBSERVATIONS AND REASONINGS WHICH EVENTUALLY CULMINATED IN THE PREPARATION OF THE "ORIGIN OF SPECIES". Equally certain is it, that it was his geological work that led Darwin into those paths of research which in the end conducted him to his great discoveries. I quite agree with the view expressed by Mr F. Darwin and Professor Seward, that Darwin, like Lyell, "thought it 'almost useless' to try to prove the truth of evolution until the cause of change was discovered" ("M.L." I. page 38.), and that possibly he may at times have vacillated in his opinions, but I believe there is evidence that, from the date mentioned, the "species question" was always more or less present in Darwin's mind. (Although we admit with Huxley that Darwin's training in comparative anatomy was very small, yet it may be remembered that he was a medical student for two years, and, if he hated the lectures, he enjoyed the society of naturalists. He had with him in the little "Beagle" library a fair number of zoological books, including works on Osteology by Cuvier, Desmarest and Lesson, as well as two French Encyclopaedias of Natural History. As a sportsman, he would obtain specimens of recent mammals in South America, and would thus have opportunities of studying their teeth and general anatomy. Keen observer, as he undoubtedly was, we need not then be surprised that he was able to make out the resemblances between the recent and fossil forms.)

It is clear that, as time went on, Darwin became more and more absorbed in his geological work. One very significant fact was that the once ardent sportsman, when he found that shooting the necessary game and zoological specimens interfered with his work with the hammer, gave up his gun to his servant. ("L.L." I. page 63.) There is clear evidence that Darwin gradually became aware how futile were his attempts to add to zoological knowledge by dissection and drawing, while he felt ever increasing satisfaction with his geological work.

The voyage fortunately extended to a much longer period (five years) than the two originally intended, but after being absent nearly three years, Darwin wrote to his sister in November, 1834, "Hurrah! hurrah! it is fixed that the 'Beagle' shall not go one mile south of Cape Tres Montes (about 200 miles south of Chiloe), and from that point to Valparaiso will be finished in about five months. We shall examine the Chonos Archipelago, entirely unknown, and the curious inland sea behind Chiloe. For me it is glorious. Cape Tres Montes is the most southern point where there is much geological interest, as there the modern beds end. The Captain then talks of crossing the Pacific; but I think we shall persuade him to finish the coast of Peru, where the climate is delightful, the country hideously sterile, but abounding with the highest interest to the geologist... I have long been grieved and most sorry at the interminable length of the voyage (though I never would have quitted it)... I could not make up my mind to return. I could not give up all the geological castles in the air I had been building up for the last two years." ("L.L." I. pages 257-58.)

In April, 1835, he wrote to another sister: "I returned a week ago from my excursion across the Andes to Mendoza. Since leaving England I have never made so successful a journey... how deeply I have enjoyed it; it was something more than enjoyment; I cannot express the delight which I felt at such a famous winding-up of all my geology in South America. I literally could hardly sleep at nights for thinking over my day's work. The scenery was so new, and so majestic; everything at an elevation of 12,000 feet bears so different an aspect from that in the lower country... To a geologist, also, there are such manifest proofs of excessive violence; the strata of the highest pinnacles are tossed about like the crust of a broken pie." ("L.L." I. pages 259-60.)

Darwin anticipated with intense pleasure his visit to the Galapagos Islands. On July 12th, 1835, he wrote to Henslow: "In a few days' time the "Beagle" will sail for the Galapagos Islands. I look forward with joy and interest to this, both as being somewhat nearer to England and for the sake of having a good look at an active volcano. Although we have seen lava in abundance, I have never yet beheld the crater." ("M.L." I. page 26.) He could little anticipate, as he wrote these lines, the important aid in the solution of the "species question" that would ever after make his visit to the Galapagos Islands so memorable. In 1832, as we have seen, the great discovery of the relations of living to extinct mammals in the same area had dawned upon his mind; in 1835 he was to find a second key for opening up the great mystery, by recognising the variations of similar types in adjoining islands among the Galapagos.

The final chapter in the second volume of the "Principles" had aroused in Darwin's mind a desire to study coral-reefs, which was gratified during his voyage across the Pacific and Indian Oceans. His theory on the subject was suggested about the end of 1834 or the beginning of 1835, as he himself tells us, before he had seen a coral-reef, and resulted from his work during two years in which he had "been incessantly attending to the effects on the shores of South America of the intermittent elevation of the land, together with denudation and the deposition of sediment." ("L.L." I. page 70.)

On arriving at the Cape of Good Hope in July, 1836, Darwin was greatly gratified by hearing that Sedgwick had spoken to his father in high terms of praise concerning the work done by him in South America. Referring to the news from home, when he reached Bahia once more, on the return voyage (August, 1836), he says: "The desert, volcanic rocks, and wild sea of Ascension... suddenly wore a pleasing aspect, and I set to work with a good-will at my old work of Geology." ("L.L." I. page 265.) Writing fifty years later, he says: "I clambered over the mountains of Ascension with a bounding step and made the volcanic rocks resound under my geological hammer!" ("L.L." I. page 66.)

That his determination was now fixed to devote his own labours to the task of working out the geological results of the voyage, and that he was prepared to leave to more practised hands the study of his biological collections, is clear from the letters he sent home at this time. From St Helena he wrote to Henslow asking that he would propose him as a Fellow of the Geological Society; and his Certificate, in Henslow's handwriting, is dated September 8th, 1836, being signed from personal knowledge by Henslow and Sedgwick. He was proposed on November 2nd and elected November 30th, being formally admitted to the Society by Lyell, who was then President, on January 4th, 1837, on which date he also read his first paper. Darwin did not become a Fellow of the Linnean Society till eighteen years later (in 1854).

An estimate of the value and importance of Darwin's geological discoveries during the voyage of the "Beagle" can best be made when considering the various memoirs and books in which the author described them. He was too cautious to allow himself to write his first impressions in his Journal, and wisely waited till he could study his specimens under better conditions and with help from others on his return. The extracts published from his correspondence with Henslow and others, while he was still abroad, showed, nevertheless, how great was the mass of observation, how suggestive and pregnant with results were the reasonings of the young geologist.

Two sets of these extracts from Darwin's letters to Henslow were printed while he was still abroad. The first of these was the series of "Geological Notes made during a survey of the East and West Coasts of South America, in the years 1832, 1833, 1834 and 1835, with an account of a transverse section of the Cordilleras of the Andes between Valparaiso and Mendoza". Professor Sedgwick, who read these notes to the Geological Society on November 18th, 1835, stated that "they were extracted from a series of letters (addressed to Professor Henslow), containing a great mass of information connected with almost every branch of natural history," and that he (Sedgwick) had made a selection of the remarks which he thought would be more especially interesting to the Geological Society. An abstract of three pages was published in the "Proceedings of the Geological Society" (Vol. II. pages 210-12.), but so unknown was the author at this time that he was described as F. Darwin, Esq., of St John's College, Cambridge! Almost simultaneously (on November 16th, 1835) a second set of extracts from these letters--this time of a general character--were read to the Philosophical Society at Cambridge, and these excited so much interest that they were privately printed in pamphlet form for circulation among the members.

Many expeditions and "scientific missions" have been despatched to various parts of the world since the return of the "Beagle" in 1836, but it is doubtful whether any, even the most richly endowed of them, has brought back such stores of new information and fresh discoveries as did that little "ten-gun brig"--certainly no cabin or laboratory was the birth-place of ideas of such fruitful character as was that narrow end of a chart-room, where the solitary naturalist could climb into his hammock and indulge in meditation.

The third and most active portion of Darwin's career as a geologist was the period which followed his return to England at the end of 1836. His immediate admission to the Geological Society, at the beginning of 1837, coincided with an important crisis in the history of geological science.

The band of enthusiasts who nearly thirty years before had inaugurated the Geological Society--weary of the fruitless conflicts between "Neptunists" and "Plutonists"--had determined to eschew theory and confine their labours to the collection of facts, their publications to the careful record of observations. Greenough, the actual founder of the Society, was an ardent Wernerian, and nearly all his fellow-workers had come, more or less directly, under the Wernerian teaching. Macculloch alone gave valuable support to the Huttonian doctrines, so far as they related to the influence of igneous activity--but the most important portion of the now celebrated "Theory of the Earth"--that dealing with the competency of existing agencies to account for changes in past geological times--was ignored by all alike. Macculloch's influence on the development of geology, which might have had far-reaching effects, was to a great extent neutralised by his peculiarities of mind and temper; and, after a stormy and troublous career, he retired from the society in 1832. In all the writings of the great pioneers in English geology, Hutton and his splendid generalisation are scarcely ever referred to. The great doctrines of Uniformitarianism, which he had foreshadowed, were completely ignored, and only his extravagances of "anti-Wernerianism" seem to have been remembered.

When between 1830 and 1832, Lyell, taking up the almost forgotten ideas of Hutton, von Hoff and Prevost, published that bold challenge to the Catastrophists--the "Principles of Geology"--he was met with the strongest opposition, not only from the outside world, which was amused by his "absurdities" and shocked by his "impiety"--but not less from his fellow-workers and friends in the Geological Society. For Lyell's numerous original observations, and his diligent collection of facts his contemporaries had nothing but admiration, and they cheerfully admitted him to the highest offices in the society, but they met his reasonings on geological theory with vehement opposition and his conclusions with coldness and contempt.

There is, indeed, a very striking parallelism between the reception of the "Principles of Geology" by Lyell's contemporaries and the manner in which the "Origin of Species" was met a quarter of a century later, as is so vividly described by Huxley. ("L.L." II. pages 179-204.) Among Lyell's fellow-geologists, two only--G. Poulett Scrope and John Herschel (Both Lyell and Darwin fully realised the value of the support of these two friends. Scrope in his appreciative reviews of the "Principles" justly pointed out what was the weakest point, the inadequate recognition of sub-aerial as compared with marine denudation. Darwin also admitted that Scrope had to a great extent forestalled him in his theory of Foliation. Herschel from the first insisted that the leading idea of the "Principles" must be applied to organic as well as to inorganic nature and must explain the appearance of new species (see Lyell's "Life and Letters", Vol. I. page 467). Darwin tells us that Herschel's "Introduction to the Study of Natural Philosophy" with Humboldt's "Personal Narrative" "stirred up in me a burning zeal" in his undergraduate days. I once heard Lyell exclaim with fervour "If ever there was a heaven-born genius it was John Herschel!")--declared themselves from the first his strong supporters. Scrope in two luminous articles in the "Quarterly Review" did for Lyell what Huxley accomplished for Darwin in his famous review in the "Times"; but Scrope unfortunately was at that time immersed in the stormy sea of politics, and devoted his great powers of exposition to the preparation of fugitive pamphlets. Herschel, like Scrope, was unable to support Lyell at the Geological Society, owing to his absence on the important astronomical mission to the Cape.

It thus came about that, in the frequent conflicts of opinion within the walls of the Geological Society, Lyell had to bear the brunt of battle for Uniformitarianism quite alone, and it is to be feared that he found himself sadly overmatched when opposed by the eloquence of Sedgwick, the sarcasm of Buckland, and the dead weight of incredulity on the part of Greenough, Conybeare, Murchison and other members of the band of pioneer workers. As time went on there is evidence that the opposition of De la Beche and Whewell somewhat relaxed; the brilliant "Paddy" Fitton (as his friends called him) was sometimes found in alliance with Lyell, but was characteristically apt to turn his weapon, as occasion served, on friend or foe alike; the amiable John Phillips "sat upon the fence." Only when a new generation arose--including Jukes, Ramsay, Forbes and Hooker--did Lyell find his teachings received with anything like favour.

We can well understand, then, how Lyell would welcome such a recruit as young Darwin--a man who had declared himself more Lyellian than Lyell, and who brought to his support facts and observations gleaned from so wide a field.

The first meeting of Lyell and Darwin was characteristic of the two men. Darwin at once explained to Lyell that, with respect to the origin of coral-reefs, he had arrived at views directly opposed to those published by "his master." To give up his own theory, cost Lyell, as he told Herschel, a "pang at first," but he was at once convinced of the immeasurable superiority of Darwin's theory. I have heard members of Lyell's family tell of the state of wild excitement and sustained enthusiasm, which lasted for days with Lyell after this interview, and his letters to Herschel, Whewell and others show his pleasure at the new light thrown upon the subject and his impatience to have the matter laid before the Geological Society.

Writing forty years afterwards, Darwin, speaking of the time of the return of the "Beagle", says: "I saw a great deal of Lyell. One of his chief characteristics was his sympathy with the work of others, and I was as much astonished as delighted at the interest which he showed when, on my return to England, I explained to him my views on coral-reefs. This encouraged me greatly, and his advice and example had much influence on me." ("L.L." I. page 68.) Darwin further states that he saw more of Lyell at this time than of any other scientific man, and at his request sent his first communication to the Geological Society. ("L.L." I. page 67.)

"Mr Lonsdale" (the able curator of the Geological Society), Darwin wrote to Henslow, "with whom I had much interesting conversation," "gave me a most cordial reception," and he adds, "If I was not much more inclined for geology than the other branches of Natural History, I am sure Mr Lyell's and Lonsdale's kindness ought to fix me. You cannot conceive anything more thoroughly good-natured than the heart-and-soul manner in which he put himself in my place and thought what would be best to do." ("L.L." I. page 275.)

Within a few days of Darwin's arrival in London we find Lyell writing to Owen as follows:

"Mrs Lyell and I expect a few friends here on Saturday next, 29th (October), to an early tea party at eight o'clock, and it will give us great pleasure if you can join it. Among others you will meet Mr Charles Darwin, whom I believe you have seen, just returned from South America, where he has laboured for zoologists as well as for hammer-bearers. I have also asked your friend Broderip." ("The Life of Richard Owen", London, 1894, Vol. I. page 102.) It would probably be on this occasion that the services of Owen were secured for the work on the fossil bones sent home by Darwin.

On November 2nd, we find Lyell introducing Darwin as his guest at the Geological Society Club; on December 14th, Lyell and Stokes proposed Darwin as a member of the Club; between that date and May 3rd of the following year, when his election to the Club took place, he was several times dining as a guest.

On January 4th, 1837, as we have already seen, Darwin was formally admitted to the Geological Society, and on the same evening he read his first paper (I have already pointed out that the notes read at the Geological Society on Nov. 18, 1835 were extracts made by Sedgwick from letters sent to Henslow, and not a paper sent home for publication by Darwin.) before the Society, "Observations of proofs of recent elevation on the coast of Chili, made during the Survey of H.M.S. "Beagle", commanded by Captain FitzRoy, R.N." By C. Darwin, F.G.S. This paper was preceded by one on the same subject by Mr A. Caldcleugh, and the reading of a letter and other communications from the Foreign Office also relating to the earthquakes in Chili.

At the meeting of the Council of the Geological Society on February 1st, Darwin was nominated as a member of the new Council, and he was elected on February 17th.

The meeting of the Geological Society on April 19th was devoted to the reading by Owen of his paper on Toxodon, perhaps the most remarkable of the fossil mammals found by Darwin in South America; and at the next meeting, on May 3rd, Darwin himself read "A Sketch of the Deposits containing extinct Mammalia in the neighbourhood of the Plata". The next following meeting, on May 17th, was devoted to Darwin's Coral-reef paper, entitled "On certain areas of elevation and subsidence in the Pacific and Indian Oceans, as deduced from the study of Coral Formations". Neither of these three early papers of Darwin were published in the Transactions of the Geological Society, but the minutes of the Council show that they were "withdrawn by the author by permission of the Council."

Darwin's activity during this session led to some rather alarming effects upon his health, and he was induced to take a holiday in Staffordshire and the Isle of Wight. He was not idle, however, for a remark of his uncle, Mr Wedgwood, led him to make those interesting observations on the work done by earthworms, that resulted in his preparing a short memoir on the subject, and this paper, "On the Formation of Mould", was read at the Society on November 1st, 1837, being the first of Darwin's papers published in full; it appeared in Vol. V. of the "Geological Transactions", pages 505-510.

During this session, Darwin attended nearly all the Council meetings, and took such an active part in the work of the Society that it is not surprising to find that he was now requested to accept the position of Secretary. After some hesitation, in which he urged his inexperience and want of knowledge of foreign languages, he consented to accept the appointment. ("L.L." I. page 285.)

At the anniversary meeting on February 16th, 1838, the Wollaston Medal was given to Owen in recognition of his services in describing the fossil mammals sent home by Darwin. In his address, the President, Professor Whewell, dwelt at length on the great value of the papers which Darwin had laid before the Society during the preceding session.

On March 7th, Darwin read before the Society the most important perhaps of all his geological papers, "On the Connexion of certain Volcanic Phenomena in South America, and on the Formation of Mountain-Chains and Volcanoes as the effect of Continental Elevations". In this paper he boldly attacked the tenets of the Catastrophists. It is evident that Darwin at this time, taking advantage of the temporary improvement in his health, was throwing himself into the breach of Uniformitarianism with the greatest ardour. Lyell wrote to Sedgwick on April 21st, 1837, "Darwin is a glorious addition to any society of geologists, and is working hard and making way, both in his book and in our discussions." ("The Life and Letters of the Reverend Adam Sedgwick", Vol. I. page 484, Cambridge, 1890.)

We have unfortunately few records of the animated debates which took place at this time between the old and new schools of geologists. I have often heard Lyell tell how Lockhart would bring down a party of friends from the Athenaeum Club to Somerset House on Geological nights, not, as he carefully explained, that "he cared for geology, but because he liked to while the fellows fight." But it fortunately happens that a few days after this last of Darwin's great field-days, at the Geological Society, Lyell, in a friendly letter to his father-in-law, Leonard Horner, wrote a very lively account of the proceedings while his impressions were still fresh; and this gives us an excellent idea of the character of these discussions.

Neither Sedgwick nor Buckland were present on this occasion, but we can imagine how they would have chastised their two "erring pupils"--more in sorrow than in anger--had they been there. Greenough, too, was absent--possibly unwilling to countenance even by his presence such outrageous doctrines.

Darwin, after describing the great earthquakes which he had experienced in South America, and the evidence of their connection with volcanic outbursts, proceeded to show that earthquakes originated in fractures, gradually formed in the earth's crust, and were accompanied by movements of the land on either side of the fracture. In conclusion he boldly advanced the view "that continental elevations, and the action of volcanoes, are phenomena now in progress, caused by some great but slow change in the interior of the earth; and, therefore, that it might be anticipated, that the formation of mountain chains is likewise in progress: and at a rate which may be judged of by either actions, but most clearly by the growth of volcanoes." ("Proc. Geol. Soc." Vol. II. pages 654-60.)

Lyell's account ("Life, Letters and Journals of Sir Charles Lyell, Bart.", edited by his sister-in-law, Mrs Lyell, Vol. II. pages 40, 41 (Letter to Leonard Horner, 1838), 2 vols. London, 1881.) of the discussion was as follows: "In support of my heretical notions," Darwin "opened upon De la Beche, Phillips and others his whole battery of the earthquakes and volcanoes of the Andes, and argued that spaces at least a thousand miles long were simultaneously subject to earthquakes and volcanic eruptions, and that the elevation of the Pampas, Patagonia, etc., all depended on a common cause; also that the greater the contortions of strata in a mountain chain, the smaller must have been each separate and individual movement of that long series which was necessary to upheave the chain. Had they been more violent, he contended that the subterraneous fluid matter would have gushed out and overflowed, and the strata would have been blown up and annihilated. (It is interesting to compare this with what Darwin wrote to Henslow seven years earlier.) He therefore introduces a cooling of one small underground injection, and then the pumping in of other lava, or porphyry, or granite, into the previously consolidated and first-formed mass of igneous rock. (Ideas somewhat similar to this suggestion have recently been revived by Dr See ("Proc. Am. Phil. Soc." Vol. XLVII. 1908, page 262.).) When he had done his description of the reiterated strokes of his volcanic pump, De la Beche gave us a long oration about the impossibility of strata of the Alps, etc., remaining flexible for such a time as they must have done, if they were to be tilted, convoluted, or overturned by gradual small shoves. He never, however, explained his theory of original flexibility, and therefore I am as unable as ever to comprehend why flexiblility is a quality so limited in time.

"Phillips then got up and pronounced a panegyric upon the "Principles of Geology", and although he still differed, thought the actual cause doctrine had been so well put, that it had advanced the science and formed a date or era, and that for centuries the two opposite doctrines would divide geologists, some contending for greater pristine forces, others satisfied, like Lyell and Darwin, with the same intensity as nature now employs.

"Fitton quizzed Phillips a little for the warmth of his eulogy, saying that he (Fitton) and others, who had Mr Lyell always with them, were in the habit of admiring and quarrelling with him every day, as one might do with a sister or cousin, whom one would only kiss and embrace fervently after a long absence. This seemed to be Mr Phillips' case, coming up occasionally from the provinces. Fitton then finished this drollery by charging me with not having done justice to Hutton, who he said was for gradual elevation.

"I replied, that most of the critics had attacked me for overrating Hutton, and that Playfair understood him as I did.

"Whewell concluded by considering Hopkins' mathematical calculations, to which Darwin had often referred. He also said that we ought not to try and make out what Hutton would have taught and thought, if he had known the facts which we now know."

It may be necessary to point out, in explanation of the above narrative, that while it was perfectly clear from Hutton's rather obscure and involved writings that he advocated slow and gradual change on the earth's surface, his frequent references to violent action and earthquakes led many--including Playfair, Lyell and Whewell--to believe that he held the changes going on in the earth's interior to be of a catastrophic nature. Fitton, however, maintained that Hutton was consistently uniformitarian. Before the idea of the actual "flowing" of solid bodies under intense pressure had been grasped by geologists, De la Beche, like Playfair before him, maintained that the bending and folding of rocks must have been effected before their complete consolidation.

In concluding his account of this memorable discussion, Lyell adds: "I was much struck with the different tone in which my gradual causes was treated by all, even including De la Beche, from that which they experienced in the same room four years ago, when Buckland, De la Beche(?), Sedgwick, Whewell, and some others treated them with as much ridicule as was consistent with politeness in my presence."

This important paper was, in spite of its theoretical character, published in full in the "Transactions of the Geological Society" (Ser. 2, Vol. V. pages 601-630). It did not however appear till 1840, and possibly some changes may have been made in it during the long interval between reading and printing. During the year 1839, Darwin continued his regular attendance at the Council meetings, but there is no record of any discussions in which he may have taken part, and he contributed no papers himself to the Society. At the beginning of 1840, he was re-elected for the third time as Secretary, but the results of failing health are indicated by the circumstance that, only at one meeting early in the session, was he able to attend the Council. At the beginning of the next session (Feb. 1841) Bunbury succeeded him as Secretary, Darwin still remaining on the Council. It may be regarded as a striking indication of the esteem in which he was held by his fellow geologists, that Darwin remained on the Council for 14 consecutive years down to 1849, though his attendances were in some years very few. In 1843 and 1844 he was a Vice-president, but after his retirement at the beginning of 1850, he never again accepted re-nomination. He continued, however, to contribute papers to the Society, as we shall see, down to the end of 1862.

Although Darwin early became a member of the Geological Dining Club, it is to be feared that he scarcely found himself in a congenial atmosphere at those somewhat hilarious gatherings, where the hardy wielders of the hammer not only drank port--and plenty of it--but wound up their meal with a mixture of Scotch ale and soda water, a drink which, as reminiscent of the "field," was regarded as especially appropriate to geologists. Even after the meetings, which followed the dinners, they reassembled for suppers, at which geological dainties, like "pterodactyle pie" figured in the bill of fare, and fines of bumpers were inflicted on those who talked the "ologies."

After being present at a fair number of meetings in 1837 and 1838, Darwin's attendances at the Club fell off to two in 1839, and by 1841 he had ceased to be a member. In a letter to Lyell on Dec. 2nd, 1841, Leonard Horner wrote that the day before "At the Council, I had the satisfaction of seeing Darwin again in his place and looking well. He tried the last evening meeting, but found it too much, but I hope before the end of the season he will find himself equal to that also. I hail Darwin's recovery as a vast gain to science." Darwin's probably last attendance, this time as a guest, was in 1851, when Horner again wrote to Lyell, "Charles Darwin was at the Geological Society's Club yesterday, where he had not been for ten years--remarkably well, and grown quite stout." ("Memoirs of Leonard Horner" (privately printed), Vol. II. pages 39 and 195.)

It may be interesting to note that at the somewhat less lively dining Club--the Philosophical--in the founding of which his friends Lyell and Hooker had taken so active a part, Darwin found himself more at home, and he was a frequent attendant--in spite of his residence being at Down--from 1853 to 1864. He even made contributions on scientific questions after these dinners. In a letter to Hooker he states that he was deeply interested in the reforms of the Royal Society, which the Club was founded to promote. He says also that he had arranged to come to town every Club day "and then my head, I think, will allow me on an average to go to every other meeting. But it is grievous how often any change knocks me up." ("L.L." II. pages 42, 43.)

Of the years 1837 and 1838 Darwin himself says they were "the most active ones which I ever spent, though I was occasionally unwell, and so lost some time... I also went a little into society." ("L.L." I. pages 67, 68.) But of the four years from 1839 to 1842 he has to confess sadly "I did less scientific work, though I worked as hard as I could, than during any other equal length of time in my life. This was owing to frequently recurring unwellness, and to one long and serious illness." ("L.L." I. page 69.)

Darwin's work at the Geological Society did not by any means engage the whole of his energies, during the active years 1837 and 1838. In June of the latter year, leaving town in somewhat bad health, he found himself at Edinburgh again, and engaged in examining the Salisbury Craigs, in a very different spirit to that excited by Jameson's discourse. ("L.L." I. page 290.) Proceeding to the Highlands he then had eight days of hard work at the famous "Parallel Roads of Glen Roy", being favoured with glorious weather.

He says of the writing of the paper on the subject--the only memoir contributed by Darwin to the Royal Society, to which he had been recently elected--that it was "one of the most difficult and instructive tasks I was ever engaged on." The paper extends to 40 quarto pages and is illustrated by two plates. Though it is full of the records of careful observation and acute reasoning, yet the theory of marine beaches which he propounded was, as he candidly admitted in after years ("M.L." II page 188.), altogether wrong. The alternative lake-theory he found himself unable to accept at the time, for he could not understand how barriers could be formed at successive levels across the valleys; and until the following year, when the existence of great glaciers in the district was proved by the researches of Agassiz, Buckland and others, the difficulty appeared to him an insuperable one. Although Darwin said of this paper in after years that it "was a great failure and I am ashamed of it"--yet he retained his interest in the question ever afterwards, and he says "my error has been a good lesson to me never to trust in science to the principle of exclusion." ("M.L." II. pages 171-93.)

Although Darwin had not realised in 1838 that large parts of the British Islands had been occupied by great glaciers, he had by no means failed while in South America to recognise the importance of ice-action. His observations, as recorded in his Journal, on glaciers coming down to the sea-level, on the west coast of South America, in a latitude corresponding to a much lower one than that of the British Islands, profoundly interested geologists; and the same work contains many valuable notes on the boulders and unstratified beds in South America in which they were included.

But in 1840 Agassiz read his startling paper on the evidence of the former existence of glaciers in the British Islands, and this was followed by Buckland's memoir on the same subject. On April 14, 1841, Darwin contributed to the Geological Society his important paper "On the Distribution of Erratic Boulders and the Contemporaneous Unstratified Deposits of South America", a paper full of suggestiveness for those studying the glacial deposits of this country. It was published in the "Transactions" in 1842.

The description of traces of glacial action in North Wales, by Buckland, appears to have greatly excited the interest of Darwin. With Sedgwick he had, in 1831, worked at the stratigraphy of that district, but neither of them had noticed the very interesting surface features. ("L.L." I. page 58.) Darwin was able to make a journey to North Wales in June, 1842 (alas! it was his last effort in field-geology) and as a result he published his most able and convincing paper on the subject in the September number of the "Philosophical Magazine" for 1842. Thus the mystery of the bell-stone was at last solved and Darwin, writing many years afterwards, said "I felt the keenest delight when I first read of the action of icebergs in transporting boulders, and I gloried in the progress of Geology." ("L.L." I. page 41.) To the "Geographical Journal" he had sent in 1839 a note "On a Rock seen on an Iceberg in 16 deg S. Latitude." For the subject of ice-action, indeed, Darwin retained the greatest interest to the end of his life. ("M.L." II. pages 148-71.)

In 1846, Darwin read two papers to the Geological Society "On the dust which falls on vessels in the Atlantic, and On the Geology of the Falkland Islands"; in 1848 he contributed a note on the transport of boulders from lower to higher levels; and in 1862 another note on the thickness of the Pampean formation, as shown by recent borings at Buenos Ayres. An account of the "British Fossil Lepadidae" read in 1850, was withdrawn by him.

At the end of 1836 Darwin had settled himself in lodgings in Fitzwilliam Street, Cambridge, and devoted three months to the work of unpacking his specimens and studying his collection of rocks. The pencilled notes on the Manuscript Catalogue in the Sedgwick Museum enable us to realise his mode of work, and the diligence with which it was carried on. The letters M and H, indicate the assistance he received from time to time from Professor Miller, the crystallographer, and from his friend Henslow. Miller not only measured many of the crystals submitted to him, but evidently taught Darwin to use the reflecting goniometer himself with considerable success. The "book of measurements" in which the records were kept, appears to have been lost, but the pencilled notes in the catalogue show how thoroughly the work was done. The letter R attached to some of the numbers in the catalogue evidently refers to the fact that they were submitted to Mr Trenham Reeks (who analysed some of his specimens) at the Geological Survey quarters in Craig's Court. This was at a later date when Darwin was writing the "Volcanic Islands" and "South America".

It was about the month of March, 1837, that Darwin completed this work upon his rocks, and also the unpacking and distribution of his fossil bones and other specimens. We have seen that November, 1832, must certainly be regarded as the date when he FIRST realised the important fact that the fossil mammals of the Pampean formation were all closely related to the existing forms in South America; while October, 1835, was, as undoubtedly, the date when the study of the birds and other forms of life in the several islands of the Galapagos Islands gave him his SECOND impulse towards abandoning the prevalent view of the immutability of species. When then in his pocket-book for 1837 Darwin wrote the often quoted passage: "In July opened first note-book on Transmutation of Species. Had been greatly struck from about the month of previous March on character of South American fossils, and species on Galapagos Archipelago. These facts (especially latter), origin of all my views" ("L.L." I. page 276.), it is clear that he must refer, not to his first inception of the idea of evolution, but to the flood of recollections, the reawakening of his interest in the subject, which could not fail to result from the sight of his specimens and the reference to his notes.

Except during the summer vacation, when he was visiting his father and uncle, and with the latter making his first observations upon the work of earthworms, Darwin was busy with his arrangements for the publication of the five volumes of the "Zoology of the 'Beagle'" and in getting the necessary financial aid from the government for the preparation of the plates. He was at the same time preparing his "Journal" for publication. During the years 1837 to 1843, Darwin worked intermittently on the volumes of Zoology, all of which he edited, while he wrote introductions to those by Owen and Waterhouse and supplied notes to the others.

Although Darwin says of his Journal that the preparation of the book "was not hard work, as my MS. Journal had been written with care." Yet from the time that he settled at 36, Great Marlborough Street in March, 1837, to the following November he was occupied with this book. He tells us that the account of his scientific observations was added at this time. The work was not published till March, 1839, when it appeared as the third volume of the "Narrative of the Surveying Voyages of H.M. Ships 'Adventure' and 'Beagle' between the years 1826 and 1836". The book was probably a long time in the press, for there are no less than 20 pages of addenda in small print. Even in this, its first form, the work is remarkable for its freshness and charm, and excited a great amount of attention and interest. In addition to matters treated of in greater detail in his other works, there are many geological notes of extreme value in this volume, such as his account of lightning tubes, of the organisms found in dust, and of the obsidian bombs of Australia.

Having thus got out of hand a number of preliminary duties, Darwin was ready to set to work upon the three volumes which were designed by him to constitute "The Geology of the Voyage of the 'Beagle'". The first of these was to be on "The Structure and Distribution of Coral-reefs". He commenced the writing of the book on October 5, 1838, and the last proof was corrected on May 6, 1842. Allowing for the frequent interruptions through illness, Darwin estimated that it cost him twenty months of hard work.

Darwin has related how his theory of Coral-reefs which was begun in a more "deductive spirit" than any of his other work, for in 1834 or 1835 it "was thought out on the west coast of South America, before I had seen a true coral-reef." ("L.L." I. page 70.) The final chapter in Lyell's second volume of the "Principles" was devoted to the subject of Coral-reefs, and a theory was suggested to account for the peculiar phenomena of "atolls." Darwin at once saw the difficulty of accepting the view that the numerous and diverse atolls all represent submerged volcanic craters. His own work had for two years been devoted to the evidence of land movements over great areas in South America, and thus he was led to announce his theory of subsidence to account for barrier and encircling reefs as well as atolls.

Fortunately, during his voyage across the Pacific and Indian Oceans, in his visit to Australia and his twelve days' hard work at Keeling Island, he had opportunities for putting his theory to the test of observation.

On his return to England, Darwin appears to have been greatly surprised at the amount of interest that his new theory excited. Urged by Lyell, he read to the Geological Society a paper on the subject, as we have seen, with as little delay as possible, but this paper was "withdrawn by permission of the Council." An abstract of three pages however appeared in the "Proceedings of the Geological Society". (Vol. II. pages 552-554 (May 31, 1837).) A full account of the observations and the theory was given in the "Journal" (1839) in the 40 pages devoted to Keeling Island in particular and to Coral formations generally. ("Journal" (1st edition), pages 439-69.)

It will be readily understood what an amount of labour the book on Coral reefs cost Darwin when we reflect on the number of charts, sailing directions, narratives of voyages and other works which, with the friendly assistance of the authorities at the Admiralty, he had to consult before he could draw up his sketch of the nature and distribution of the reefs, and this was necessary before the theory, in all its important bearings, could be clearly enunciated. Very pleasing is it to read how Darwin, although arriving at a different conclusion to Lyell, shows, by quoting a very suggestive passage in the "Principles" (1st edition Vol. II. page 296.), how the latter only just missed the true solution. This passage is cited, both in the "Journal" and the volume on Coral-reefs. Lyell, as we have seen, received the new theory not merely ungrudgingly, but with the utmost enthusiasm.

In 1849 Darwin was gratified by receiving the support of Dana, after his prolonged investigation in connection with the U.S. Exploring Expedition ("M.L." II. pages 226-8.), and in 1874 he prepared a second edition of his book, in which some objections which had been raised to the theory were answered. A third edition, edited by Professor Bonney, appeared in 1880, and a fourth (a reprint of the first edition, with introduction by myself) in 1890.

Although Professor Semper, in his account of the Pelew Islands, had suggested difficulties in the acceptance of Darwin's theory, it was not till after the return of the "Challenger" expedition in 1875 that a rival theory was propounded, and somewhat heated discussions were raised as to the respective merits of the two theories. While geologists have, nearly without exception, strongly supported Darwin's views, the notes of dissent have come almost entirely from zoologists. At the height of the controversy unfounded charges of unfairness were made against Darwin's supporters and the authorities of the Geological Society, but this unpleasant subject has been disposed of, once for all, by Huxley. ("Essays upon some Controverted Questions", London, 1892, pages 314-328 and 623-625.)

Darwin's final and very characteristic utterance on the coral-reef controversy is found in a letter which he wrote to Professor Alexander Agassiz, May 5th, 1881: less than a year before his death: "If I am wrong, the sooner I am knocked on the head and annihilated so much the better. It still seems to me a marvellous thing that there should not have been much, and long-continued, subsidence in the beds of the great oceans. I wish that some doubly rich millionaire would take it into his head to have borings made in some of the Pacific and Indian atolls, and bring home cores for slicing from a depth of 500 or 600 feet." ("L.L." III. page 184.)

Though the "doubly rich millionaire" has not been forthcoming, the energy, in England, of Professor Sollas, and in New South Wales of Professor Anderson Stuart served to set on foot a project, which, aided at first by the British Association for the Advancement of Science, and afterwards taken up jointly by the Royal Society, the New South Wales Government, and the Admiralty, has led to the most definite and conclusive results.

The Committee appointed by the Royal Society to carry out the undertaking included representatives of all the views that had been put forward on the subject. The place for the experiment was, with the consent of every member of the Committee, selected by the late Admiral Sir W.J. Wharton--who was not himself an adherent of Darwin's views--and no one has ventured to suggest that his selection, the splendid atoll of Funafuti, was not a most judicious one.

By the pluck and perseverance of Professor Sollas in the preliminary expedition, and of Professor T. Edgeworth David and his pupils, in subsequent investigations of the island, the rather difficult piece of work was brought to a highly satisfactory conclusion. The New South Wales Government lent boring apparatus and workmen, and the Admiralty carried the expedition to its destination in a surveying ship which, under Captain (now Admiral) A. Mostyn Field, made the most complete survey of the atoll and its surrounding seas that has ever been undertaken in the case of a coral formation.

After some failures and many interruptions, the boring was carried to the depth of 1114 feet, and the cores obtained were sent to England. Here the examination of the materials was fortunately undertaken by a zoologist of the highest repute, Dr G.J. Hinde--who has a wide experience in the study of organisms by sections--and he was aided at all points by specialists in the British Museum of Natural History and by other naturalists. Nor were the chemical and other problems neglected.

The verdict arrived at, after this most exhaustive study of a series of cores obtained from depths twice as great as that thought necessary by Darwin, was as follows:--"The whole of the cores are found to be built up of those organisms which are seen forming coral-reefs near the surface of the ocean--many of them evidently in situ; and not the slightest indication could be detected, by chemical or microscopic means, which suggested the proximity of non-calcareous rocks, even in the lowest portions brought up."

But this was not all. Professor David succeeded in obtaining the aid of a very skilful engineer from Australia, while the Admiralty allowed Commander F.C.D. Sturdee to take a surveying ship into the lagoon for further investigations. By very ingenious methods, and with great perseverance, two borings were put down in the midst of the lagoon to the depth of nearly 200 feet. The bottom of the lagoon, at the depth of 101 1/2 feet from sea-level, was found to be covered with remains of the calcareous, green sea-weed Halimeda, mingled with many foraminifera; but at a depth of 163 feet from the surface of the lagoon the boring tools encountered great masses of coral, which were proved from the fragments brought up to belong to species that live within AT MOST 120 feet from the surface of the ocean, as admitted by all zoologists. ("The Atoll of Funafuti; Report of the Coral Reef Committee of the Royal Society", London, 1904.)

Darwin's theory, as is well known, is based on the fact that the temperature of the ocean at any considerable depth does not permit of the existence and luxuriant growth of the organisms that form the reefs. He himself estimated this limit of depth to be from 120 to 130 feet; Dana, as an extreme, 150 feet; while the recent very prolonged and successful investigations of Professor Alexander Agassiz in the Pacific and Indian Oceans lead him also to assign a limiting depth of 150 feet; the EFFECTIVE, REEF-FORMING CORALS, however, flourishing at a much smaller depth. Mr Stanley Gardiner gives for the most important reef-forming corals depths between 30 and 90 feet, while a few are found as low as 120 feet or even 180 feet.

It will thus be seen that the verdict of Funafuti is clearly and unmistakeably in favour of Darwin's theory. It is true that some zoologists find a difficulty in realising a slow sinking of parts of the ocean floor, and have suggested new and alternative explanations: but geologists generally, accepting the proofs of slow upheaval in some areas--as shown by the admirable researches of Alexander Agassiz--consider that it is absolutely necessary to admit that this elevation is balanced by subsidence in other areas. If atolls and barrier-reefs did not exist we should indeed be at a great loss to frame a theory to account for their absence.

After finishing his book on Coral-reefs, Darwin made his summer excursion to North Wales, and prepared his important memoir on the glaciers of that district: but by October (1842) we find him fairly settled at work upon the second volume of his "Geology of the 'Beagle'--Geological Observations on the Volcanic Islands, visited during the Voyage of H.M.S. 'Beagle'". The whole of the year 1843 was devoted to this work, but he tells his friend Fox that he could "manage only a couple of hours per day, and that not very regularly." ("L.L." I. page 321.) Darwin's work on the various volcanic islands examined by him had given him the most intense pleasure, but the work of writing the book by the aid of his notes and specimens he found "uphill work," especially as he feared the book would not be read, "even by geologists." (Loc. cit.)

As a matter of fact the work is full of the most interesting observations and valuable suggestions, and the three editions (or reprints) which have appeared have proved a most valuable addition to geological literature. It is not necessary to refer to the novel and often very striking discoveries described in this well-known work. The subsidence beneath volcanic vents, the enormous denudation of volcanic cones reducing them to "basal wrecks," the effects of solfatarric action and the formation of various minerals in the cavities of rocks--all of these subjects find admirable illustration from his graphic descriptions. One of the most important discussions in this volume is that dealing with the "lamination" of lavas as especially well seen in the rocks of Ascension. Like Scrope, Darwin recognised the close analogy between the structure of these rocks and those of metamorphic origin--a subject which he followed out in the volume "Geological Observations on South America".

Of course in these days, since the application of the microscope to the study of rocks in thin sections, Darwin's nomenclature and descriptions of the petrological characters of the lavas appear to us somewhat crude. But it happened that the "Challenger" visited most of the volcanic islands described by Darwin, and the specimens brought home were examined by the eminent petrologist Professor Renard. Renard was so struck with the work done by Darwin, under disadvantageous conditions, that he undertook a translation of Darwin's work into French, and I cannot better indicate the manner in which the book is regarded by geologists than by quoting a passage from Renard's preface. Referring to his own work in studying the rocks brought home by the "Challenger" (Renard's descriptions of these rocks are contained in the "Challenger Reports". Mr Harker is supplementing these descriptions by a series of petrological memoirs on Darwin's specimens, the first of which appeared in the "Geological Magazine" for March, 1907.), he says:

"Je dus, en me livrant a ces recherches, suivre ligne par ligne les divers chapitres des "Observations geologiques" consacrees aux iles de l'Atlantique, oblige que j'etais de comparer d'une maniere suivie les resultats auxquels j'etais conduit avec ceux de Darwin, qui servaient de controle a mes constatations. Je ne tardai pas a eprouver une vive admiration pour ce chercheur qui, sans autre appareil que la loupe, sans autre reaction que quelques essais pyrognostiques, plus rarement quelques mesures au goniometre, parvenait a discerner la nature des agregats mineralogiques les plue complexes et les plus varies. Ce coup d'oeil qui savait embrasser de si vastes horizons, penetre ici profondement tous les details lithologiques. Avec quelle surete et quelle exactitude la structure et la composition des roches ne sont'elles pas determinees, l'origne de ces masses minerales deduite et confirmee par l'etude comparee des manifestations volcaniques d'autres regions; avec quelle science les relations entre les faits qu'il decouvre et ceux signales ailleurs par ses devanciers ne sont'elles pas etablies, et comme voici ebranlees les hypotheses regnantes, admises sans preuves, celles, par exemple, des crateres de soulevement et de la differenciation radicale des phenomenes plutoniques et volcaniques! Ce qui acheve de donner a ce livre un incomparable merite, ce sont les idees nouvelles qui s'y trouvent en germe et jetees la comme au hasard ainsi qu'un superflu d'abondance intellectuelle inepuisable." ("Observations Geologiques sur les Iles Volcaniques... ", Paris, 1902, pages vi., vii.)

While engaged in his study of banded lavas, Darwin was struck with the analogy of their structure with that of glacier ice, and a note on the subject, in the form of a letter addressed to Professor J.D. Forbes, was published in the "Proceedings of the Royal Society of Edinburgh". (Vol. II. (1844-5), pages 17, 18.)

From April, 1832, to September, 1835, Darwin had been occupied in examining the coast or making inland journeys in the interior of the South American continent. Thus while eighteen months were devoted, at the beginning and end of the voyage to the study of volcanic islands and coral-reefs, no less than three and a half years were given to South American geology. The heavy task of dealing with the notes and specimens accumulated during that long period was left by Darwin to the last. Finishing the "Volcanic Islands" on February 14th, 1844, he, in July of the same year, commenced the preparation of two important works which engaged him till near the end of the year 1846. The first was his "Geological Observations on South America", the second a recast of his "Journal", published under the short title of "A Naturalist's Voyage round the World".

The first of these works contains an immense amount of information collected by the author under great difficulties and not unfrequently at considerable risk to life and health. No sooner had Darwin landed in South America than two sets of phenomena powerfully arrested his attention. The first of these was the occurrence of great masses of red mud containing bones and shells, which afforded striking evidence that the whole continent had shared in a series of slow and gradual but often interrupted movements. The second related to the great masses of crystalline rocks which, underlying the muds, cover so great a part of the continent. Darwin, almost as soon as he landed, was struck by the circumstance that the direction, as shown by his compass, of the prominent features of these great crystalline rock-masses--their cleavage, master-joints, foliation and pegmatite veins--was the same as the orientation described by Humboldt (whose works he had so carefully studied) on the west of the same great continent.

The first five chapters of the book on South America were devoted to formations of recent date and to the evidence collected on the east and west coasts of the continent in regard to those grand earth-movements, some of which could be shown to have been accompanied by earthquake-shocks. The fossil bones, which had given him the first hint concerning the mutability of species, had by this time been studied and described by comparative anatomists, and Darwin was able to elaborate much more fully the important conclusion that the existing fauna of South America has a close analogy with that of the period immediately preceding our own.

The remaining three chapters of the book dealt with the metamorphic and plutonic rocks, and in them Darwin announced his important conclusions concerning the relations of cleavage and foliation, and on the close analogy of the latter structure with the banding found in rock-masses of igneous origin. With respect to the first of these conclusions, he received the powerful support of Daniel Sharpe, who in the years 1852 and 1854 published two papers on the structure of the Scottish Highlands, supplying striking confirmation of the correctness of Darwin's views. Although Darwin's and Sharpe's conclusions were contested by Murchison and other geologists, they are now universally accepted. In his theory concerning the origin of foliation, Darwin had been to some extent anticipated by Scrope, but he supplied many facts and illustrations leading to the gradual acceptance of a doctrine which, when first enunciated, was treated with neglect, if not with contempt.

The whole of this volume on South American geology is crowded with the records of patient observations and suggestions of the greatest value; but, as Darwin himself saw, it was a book for the working geologist and "caviare to the general." Its author, indeed, frequently expressed his sense of the "dryness" of the book; he even says "I long hesitated whether I would publish it or not," and he wrote to Leonard Horner "I am astonished that you should have had the courage to go right through my book." ("M.L." II. page 221.)

Fortunately the second book, on which Darwin was engaged at this time, was of a very different character. His "Journal", almost as he had written it on board ship, with facts and observations fresh in his mind, had been published in 1839 and attracted much attention. In 1845, he says, "I took much pains in correcting a new edition," and the work which was commenced in April, 1845, was not finished till August of that year. The volume contains a history of the voyage with "a sketch of those observations in Natural History and Geology, which I think will possess some interest for the general reader." It is not necessary to speak of the merits of this scientific classic. It became a great favourite with the general public--having passed through many editions--it was, moreover, translated into a number of different languages. Darwin was much gratified by these evidences of popularity, and naively remarks in his "Autobiography", "The success of this my first literary child tickles my vanity more than that of any of my other books" ("L.L." I. page 80.)--and this was written after the "Origin of Species" had become famous!

In Darwin's letters there are many evidences that his labours during these ten years devoted to the working out of the geological results of the voyage often made many demands on his patience and indomitable courage. Most geologists have experience of the contrast between the pleasures felt when wielding the hammer in the field, and the duller labour of plying the pen in the study. But in Darwin's case, innumerable interruptions from sickness and other causes, and the oft-deferred hope of reaching the end of his task were not the only causes operating to make the work irksome. The great project, which was destined to become the crowning achievement of his life, was now gradually assuming more definite shape, and absorbing more of his time and energies.

Nevertheless, during all this period, Darwin so far regarded his geological pursuits as his PROPER "work," that attention to other matters was always spoken of by him as "indulging in idleness." If at the end of this period the world had sustained the great misfortune of losing Darwin by death before the age of forty--and several times that event seemed only too probable--he might have been remembered only as a very able geologist of most advanced views, and a traveller who had written a scientific narrative of more than ordinary excellence!

The completion of the "Geology of the 'Beagle'" and the preparation of a revised narrative of the voyage mark the termination of that period of fifteen years of Darwin's life during which geological studies were his principal occupation. Henceforth, though his interest in geological questions remained ever keen, biological problems engaged more and more of his attention to the partial exclusion of geology.

The eight years from October, 1846, to October, 1854, were mainly devoted to the preparation of his two important monographs on the recent and fossil Cirripedia. Apart from the value of his description of the fossil forms, this work of Darwin's had an important influence on the progress of geological science. Up to that time a practice had prevailed for the student of a particular geological formation to take up the description of the plant and animal remains in it--often without having anything more than a rudimentary knowledge of the living forms corresponding to them. Darwin in his monograph gave a very admirable illustration of the enormous advantage to be gained--alike for biology and geology--by undertaking the study of the living and fossil forms of a natural group of organisms in connection with one another. Of the advantage of these eight years of work to Darwin himself, in preparing for the great task lying before him, Huxley has expressed a very strong opinion indeed. ("L.L." II. pages 247-48.)

But during these eight years of "species work," Darwin found opportunities for not a few excursions into the field of geology. He occasionally attended the Geological Society, and, as we have already seen, read several papers there during this period. His friend, Dr Hooker, then acting as botanist to the Geological Survey, was engaged in studying the Carboniferous flora, and many discussions on Palaezoic plants and on the origin of coal took place at this period. On this last subject he felt the deepest interest and told Hooker, "I shall never rest easy in Down churchyard without the problem be solved by some one before I die." ("M.L." I. pages 63, 64.)

As at all times, conversations and letters with Lyell on every branch of geological science continued with unabated vigour, and in spite of the absorbing character of the work on the Cirripedes, time was found for all. In 1849 his friend Herschel induced him to supply a chapter of forty pages on Geology to the Admiralty "Manual of Scientific Inquiry" which he was editing. This is Darwin's single contribution to books of an "educational" kind. It is remarkable for its clearness and simplicity and attention to minute details. It may be read by the student of Darwin's life with much interest, for the directions he gives to an explorer are without doubt those which he, as a self-taught geologist, proved to be serviceable during his life on the "Beagle".

On the completion of the Cirripede volumes, in 1854, Darwin was able to grapple with the immense pile of MS. notes which he had accumulated on the species question. The first sketch of 35 pages (1842), had been enlarged in 1844 into one of 230 pages ([The first draft of the "Origin" is being prepared for Press by Mr Francis Darwin and will be published by the Cambridge University Press this year (1909). A.C.S.]); but in 1856 was commenced the work (never to be completed) which was designed on a scale three or four times more extensive than that on which the "Origin of Species" was in the end written.

In drawing up those two masterly chapters of the "Origin", "On the Imperfection of the Geological Record," and "On the Geological Succession of Organic Beings", Darwin had need of all the experience and knowledge he had been gathering during thirty years, the first half of which had been almost wholly devoted to geological study. The most enlightened geologists of the day found much that was new, and still more that was startling from the manner of its presentation, in these wonderful essays. Of Darwin's own sense of the importance of the geological evidence in any presentation of his theory a striking proof will be found in a passage of the touching letter to his wife, enjoining the publication of his sketch of 1844. "In case of my sudden death," he wrote, "... the editor must be a geologist as well as a naturalist." ("L.L." II. pages 16, 17.)

In spite of the numerous and valuable palaeontological discoveries made since the publication of "The Origin of Species", the importance of the first of these two geological chapters is as great as ever. It still remains true that "Those who believe that the geological record is in any degree perfect, will at once reject the theory"--as indeed they must reject any theory of evolution. The striking passage with which Darwin concludes this chapter--in which he compares the record of the rocks to the much mutilated volumes of a human history--remains as apt an illustration as it did when first written.

And the second geological chapter, on the Succession of Organic Beings--though it has been strengthened in a thousand ways, by the discoveries concerning the pedigrees of the horse, the elephant and many other aberrant types, though new light has been thrown even on the origin of great groups like the mammals, and the gymnosperms, though not a few fresh links have been discovered in the chains of evidence, concerning the order of appearance of new forms of life--we would not wish to have re-written. Only the same line of argument could be adopted, though with innumerable fresh illustrations. Those who reject the reasonings of this chapter, neither would they be persuaded if a long and complete succession of "ancestral forms" could rise from the dead and pass in procession before them.

Among the geological discussions, which so frequently occupied Darwin's attention during the later years of his life, there was one concerning which his attitude seemed somewhat remarkable--I allude to his views on "the permanence of Continents and Ocean-basins." In a letter to Mr Mellard Reade, written at the end of 1880, he wrote: "On the whole, I lean to the side that the continents have since Cambrian times occupied approximately their present positions. But, as I have said, the question seems a difficult one, and the more it is discussed the better." ("M.L." II. page 147.) Since this was written, the important contribution to the subject by the late Dr W.T. Blanford (himself, like Darwin, a naturalist and geologist) has appeared in an address to the Geological Society in 1890; and many discoveries, like that of Dr Woolnough in Fiji, have led to considerable qualifications of the generalisation that all the islands in the great ocean are wholly of volcanic or coral origin.

I remember once expressing surprise to Darwin that, after the views which he had originated concerning the existence of areas of elevation and others of subsidence in the Pacific Ocean, and in face of the admitted difficulty of accounting for the distribution of certain terrestrial animals and plants, if the land and sea areas had been permanent in position, he still maintained that theory. Looking at me with a whimsical smile, he said: "I have seen many of my old friends make fools of themselves, by putting forward new theoretical views or revising old ones, AFTER THEY WERE SIXTY YEARS OF AGE; so, long ago, I determined that on reaching that age I would write nothing more of a speculative character."

Though Darwin's letters and conversations on geology during these later years were the chief manifestations of the interest he preserved in his "old love," as he continued to call it, yet in the sunset of that active life a gleam of the old enthusiasm for geology broke forth once more. There can be no doubt that Darwin's inability to occupy himself with field-work proved an insuperable difficulty to any attempt on his part to resume active geological research. But, as is shown by the series of charming volumes on plant-life, Darwin had found compensation in making patient and persevering experiment take the place of enterprising and exact observation; and there was one direction in which he could indulge the "old love" by employment of the new faculty.

We have seen that the earliest memoir written by Darwin, which was published in full, was a paper "On the Formation of Mould" which was read at the Geological Society on November 1st, 1837, but did not appear in the "Transactions" of the Society till 1840, where it occupied four and a half quarto pages, including some supplementary matter, obtained later, and a woodcut. This little paper was confined to observations made in his uncle's fields in Staffordshire, where burnt clay, cinders, and sand were found to be buried under a layer of black earth, evidently brought from below by earthworms, and to a recital of similar facts from Scotland obtained through the agency of Lyell. The subsequent history of Darwin's work on this question affords a striking example of the tenacity of purpose with which he continued his enquiries on any subject that interested him.

In 1842, as soon as he was settled at Down, he began a series of observations on a foot-path and in his fields, that continued with intermissions during his whole life, and he extended his enquiries from time to time to the neighbouring parks of Knole and Holwood. In 1844 we find him making a communication to the "Gardener's Chronicle" on the subject. About 1870, his attention to the question was stimulated by the circumstance that his niece (Miss L. Wedgwood) undertook to collect and weigh the worm-casts thrown up, during a whole year, on measured squares selected for the purpose, at Leith Hill Place. He also obtained information from Professor Ramsay concerning observations made by him on a pavement near his house in 1871. Darwin at this time began to realise the great importance of the action of worms to the archaeologist. At an earlier date he appears to have obtained some information concerning articles found buried on the battle-field of Shrewsbury, and the old Roman town of Uriconium, near his early home; between 1871 and 1878 Mr (afterwards Lord) Farrer carried on a series of investigations at the Roman Villa discovered on his land at Abinger; Darwin's son William examined for his father the evidence at Beaulieu Abbey, Brading, Stonehenge and other localities in the neighbourhood of his home; his sons Francis and Horace were enlisted to make similar enquiries at Chideock and Silchester; while Francis Galton contributed facts noticed in his walks in Hyde Park. By correspondence with Fritz Muller and Dr Ernst, Darwin obtained information concerning the worm-casts found in South America; from Dr Kreft those of Australia; and from Mr Scott and Dr (afterwards Sir George) King, those of India; the last-named correspondent also supplied him with much valuable information obtained in the South of Europe. Help too was obtained from the memoirs on Earthworms published by Perrier in 1874 and van Hensen in 1877, while Professor Ray Lankester supplied important facts with regard to their anatomy.

When therefore the series of interesting monographs on plant-life had been completed, Darwin set to work in bringing the information that he had gradually accumulated during forty-four years to bear on the subject of his early paper. He also utilised the skill and ingenuity he had acquired in botanical work to aid in the elucidation of many of the difficulties that presented themselves. I well remember a visit which I paid to Down at this period. At the side of the little study stood flower-pots containing earth with worms, and, without interrupting our conversation, Darwin would from time to time lift the glass plate covering a pot to watch what was going on. Occasionally, with a humorous smile, he would murmur something about a book in another room, and slip away; returning shortly, without the book but with unmistakeable signs of having visited the snuff-jar outside. After working about a year at the worms, he was able at the end of 1881 to publish the charming little book--"The Formation of Vegetable Mould through the Action of Worms, with Observations on their Habits". This was the last of his books, and its reception by reviewers and the public alike afforded the patient old worker no little gratification. Darwin's scientific career, which had begun with geological research, most appropriately ended with a return to it.

It has been impossible to sketch the origin and influence of Darwin's geological work without, at almost every step, referring to the part played by Lyell and the "Principles of Geology". Haeckel, in the chapters on Lyell and Darwin in his "History of Creation", and Huxley in his striking essay "On the Reception of the Origin of Species" ("L.L." II. pages 179-204.) have both strongly insisted on the fact that the "Origin" of Darwin was a necessary corollary to the "Principles" of Lyell.

It is true that, in an earlier essay, Huxley had spoken of the doctrine of Uniformitarianism as being, in a certain sense, opposed to that of Evolution (Huxley's Address to the Geological Society, 1869. "Collected Essays", Vol. VIII. page 305, London, 1896.); but in his later years he took up a very different and more logical position, and maintained that "Consistent uniformitarianism postulates evolution as much in the organic as in the inorganic world. The origin of a new species by other than ordinary agencies would be a vastly greater 'catastrophe' than any of those which Lyell success fully eliminated from sober geological speculation." ("L.L." II. page 190.)

Huxley's admiration for the "Principles of Geology", and his conviction of the greatness of the revolution of thought brought about by Lyell, was almost as marked as in the case of Darwin himself. (See his Essay on "Science and Pseudo Science". "Collected Essays", Vol. V. page 90, London, 1902.) He felt, however, as many others have done, that in one respect the very success of Lyell's masterpiece has been the reason why its originality and influence have not been so fully recognised as they deserved to be. Written as the book was before its author had arrived at the age of thirty, no less than eleven editions of the "Principles" were called for in his lifetime. With the most scrupulous care, Lyell, devoting all his time and energies to the task of collecting and sifting all evidence bearing on the subjects of his work, revised and re-revised it; and as in each edition, eliminations, modifications, corrections, and additions were made, the book, while it increased in value as a storehouse of facts, lost much of its freshness, vigour and charm as a piece of connected reasoning.

Darwin undoubtedly realised this when he wrote concerning the "Principles", "the first edition, my old true love, which I never deserted for the later editions." ("M.L." II. page 222.) Huxley once told me that when, in later life, he read the first edition, he was both surprised and delighted, feeling as if it were a new book to him. (I have before me a letter which illustrates this feeling on Huxley's part. He had lamented to me that he did not possess a copy of the first edition of the "Principles", when, shortly afterwards, I picked up a dilapidated copy on a bookstall; this I had bound and sent to my old teacher and colleague. His reply is characteristic:

October 8, 1884.

My Dear Judd,

You could not have made me a more agreeable present than the copy of the first edition of Lyell, which I find on my table. I have never been able to meet with the book, and your copy is, as the old woman said of her Bible, "the best of books in the best of bindings."

Ever yours sincerely,

T.H. Huxley.

(I cannot refrain from relating an incident which very strikingly exemplifies the affection for one another felt by Lyell and Huxley. In his last illness, when confined to his bed, Lyell heard that Huxley was to lecture at the Royal Institution on the "Results of the 'Challenger' expedition": he begged me to attend the lecture and bring him an account of it. Happening to mention this to Huxley, he at once undertook to go to Lyell in my place, and he did so on the morning following his lecture. I shall never forget the look of gratitude on the face of the invalid when he told me, shortly afterwards, how Huxley had sat by his bedside and "repeated the whole lecture to him.")

Darwin's generous nature seems often to have made him experience a fear lest he should do less than justice to his "dear old master," and to the influence that the "Principles of Geology" had in moulding his mind. In 1845 he wrote to Lyell, "I have long wished, not so much for your sake, as for my own feelings of honesty, to acknowledge more plainly than by mere reference, how much I geologically owe you. Those authors, however, who like you, educate people's minds as well as teach them special facts, can never, I should think, have full justice done them except by posterity, for the mind thus insensibly improved can hardly perceive its own upward ascent." ("L.L." I. pages 337-8.) In another letter, to Leonard Horner, he says: "I always feel as if my books came half out of Lyell's brain, and that I never acknowledge this sufficiently." ("M.L." II. page 117.) Darwin's own most favourite book, the "Narrative of the Voyage", was dedicated to Lyell in glowing terms; and in the "Origin of Species" he wrote of "Lyell's grand work on the "Principles of Geology", which the future historian will recognise as having produced a revolution in Natural Science." "What glorious good that work has done" he fervently exclaims on another occasion. ("L.L." I. page 342.)

To the very end of his life, as all who were in the habit of talking with Darwin can testify, this sense of his indebtedness to Lyell remained with him. In his "Autobiography", written in 1876, the year after Lyell's death, he spoke in the warmest terms of the value to him of the "Principles" while on the voyage and of the aid afforded to him by Lyell on his return to England. ("L.L." I. page 62.) But the year before his own death, Darwin felt constrained to return to the subject and to place on record a final appreciation--one as honourable to the writer as it is to his lost friend:

"I saw more of Lyell than of any other man, both before and after my marriage. His mind was characterised, as it appeared to me, by clearness, caution, sound judgment, and a good deal of originality. When I made any remark to him on Geology, he never rested until he saw the whole case clearly, and often made me see it more clearly than I had done before. He would advance all possible objections to my suggestion, and even after these were exhausted would remain long dubious. A second characteristic was his hearty sympathy with the work of other scientific men... His delight in science was ardent, and he felt the keenest interest in the future progress of mankind. He was very kind-hearted... His candour was highly remarkable. He exhibited this by becoming a convert to the Descent theory, though he had gained much fame by opposing Lamarck's views, and this after he had grown old."

"THE SCIENCE OF GEOLOGY IS ENORMOUSLY INDEBTED TO LYELL--MORE SO, AS I BELIEVE, THAN TO ANY OTHER MAN WHO EVER LIVED." ("L.L." I. pages 71-2 (the italics are mine.))

Those who knew Lyell intimately will recognise the truth of the portrait drawn by his dearest friend, and I believe that posterity will endorse Darwin's deliberate verdict concerning the value of his labours.

It was my own good fortune, to be brought into close contact with these two great men during the later years of their life, and I may perhaps be permitted to put on record the impressions made upon me during friendly intercourse with both.

In some respects, there was an extraordinary resemblance in their modes and habits of thought, between Lyell and Darwin; and this likeness was also seen in their modesty, their deference to the opinion of younger men, their enthusiasm for science, their freedom from petty jealousies and their righteous indignation for what was mean and unworthy in others. But yet there was a difference. Both Lyell and Darwin were cautious, but perhaps Lyell carried his caution to the verge of timidity. I think Darwin possessed, and Lyell lacked, what I can only describe by the theological term, "faith--the substance of things hoped for, the evidence of things not seen." Both had been constrained to feel that the immutability of species could not be maintained. Both, too, recognised the fact that it would be useless to proclaim this conviction, unless prepared with a satisfactory alternative to what Huxley called "the Miltonic hypothesis." But Darwin's conviction was so far vital and operative that it sustained him while working unceasingly for twenty-two years in collecting evidence bearing on the question, till at last he was in the position of being able to justify that conviction to others.

And yet Lyell's attitude--and that of Hooker, which was very similar--proved of inestimable service to science, as Darwin often acknowledged. One of the greatest merits of the "Origin of Species" is that so many difficulties and objections are anticipated and fairly met; and this was to a great extent the result of the persistent and very candid--if always friendly--criticism of Lyell and Hooker.

I think the divergence of mental attitude in Lyell and Darwin must be attributed to a difference in temperament, the evidence of which sometimes appears in a very striking manner in their correspondence. Thus in 1838, while they were in the thick of the fight with the Catastrophists of the Geological Society, Lyell wrote characteristically: "I really find, when bringing up my Preliminary Essays in "Principles" to the science of the present day, so far as I know it, that the great outline, and even most of the details, stand so uninjured, and in many cases they are so much strengthened by new discoveries, especially by yours, that we may begin to hope that the great principles there insisted on will stand the test of new discoveries." (Lyell's "Life, Letters and Journals", Vol. II. page 44.) To which the more youthful and impetuous Darwin replies: "BEGIN TO HOPE: why the POSSIBILITY of a doubt has never crossed my mind for many a day. This may be very unphilosophical, but my geological salvation is staked on it... it makes me quite indignant that you should talk of HOPING." ("L.L." I. page 296.)

It was not only Darwin's "geological salvation" that was at stake, when he surrendered himself to his enthusiasm for an idea. To his firm faith in the doctrine of continuity we owe the "Origin of Species"; and while Darwin became the "Paul" of evolution, Lyell long remained the "doubting Thomas."

Many must have felt like H.C. Watson when he wrote: "How could Sir C. Lyell... for thirty years read, write, and think, on the subject of species AND THEIR SUCCESSION, and yet constantly look down the wrong road!" ("L.L." II. page 227.) Huxley attributed this hesitation of Lyell to his "profound antipathy" to the doctrine of the "pithecoid origin of man." ("L.L." II. page 193.) Without denying that this had considerable influence (and those who knew Lyell and his great devotion to his wife and her memory, are aware that he and she felt much stronger convictions concerning such subjects as the immortality of the soul than Darwin was able to confess to) yet I think Darwin had divined the real characteristics of his friend's mind, when he wrote: "He would advance all possible objections... AND EVEN AFTER THESE WERE EXHAUSTED, WOULD REMAIN LONG DUBIOUS."

Very touching indeed was the friendship maintained to the end between these two leaders of thought--free as their intercourse was from any smallest trace of self-seeking or jealousy. When in 1874 I spent some time with Lyell in his Forfarshire home, a communication from Darwin was always an event which made a "red-letter day," as Lyell used to say; and he gave me many indications in his conversation of how strongly he relied upon the opinion of Darwin--more indeed than on the judgment of any other man--this confidence not being confined to questions of science, but extending to those of morals, politics, and religion.

I have heard those who knew Lyell only slightly, speak of his manners as cold and reserved. His complete absorption in his scientific work, coupled with extreme short-sightedness, almost in the end amounting to blindness, may have permitted those having but a casual acquaintance with him to accept such a view. But those privileged to know him intimately recognised the nobleness of his character and can realise the justice and force of Hooker's words when he heard of his death: "My loved, my best friend, for well nigh forty years of my life. The most generous sharer of my own and my family's hopes, joys and sorrows, whose affection for me was truly that of a father and brother combined."

But the strongest of all testimonies to the grandeur of Lyell's character is the lifelong devotion to him of such a man as Darwin. Before the two met, we find Darwin constantly writing of facts and observations that he thinks "will interest Mr Lyell"; and when they came together the mutual esteem rapidly ripened into the warmest affection. Both having the advantage of a moderate independence, permitting of an entire devotion of their lives to scientific research, they had much in common, and the elder man--who had already achieved both scientific and literary distinction--was able to give good advice and friendly help to the younger one. The warmth of their friendship comes out very strikingly in their correspondence. When Darwin first conceived the idea of writing a book on the "species question," soon after his return from the voyage, it was "by following the example of Lyell in Geology" that he hoped to succeed ("L.L." I. page 83.); when in 1844, Darwin had finished his first sketch of the work, and, fearing that his life might not be spared to complete his great undertaking, committed the care of it in a touching letter to his wife, it was his friend Lyell whom he named as her adviser and the possible editor of the book ("L.L." II. pages 17-18.); it was Lyell who, in 1856, induced Darwin to lay the foundations of a treatise ("L.L." I. page 84.) for which the author himself selected the "Principles" as his model; and when the dilemma arose from the receipt of Wallace's essay, it was to Lyell jointly with Hooker that Darwin turned, not in vain, for advice and help.

During the later years of his life, I never heard Darwin allude to his lost friend--and he did so very often--without coupling his name with some term of affection. For a brief period, it is true, Lyell's excessive caution when the "Origin" was published, seemed to try even the patience of Darwin; but when "the master" was at last able to declare himself fully convinced, he was the occasion of more rejoicing on the part of Darwin, than any other convert to his views. The latter was never tired of talking of Lyell's "magnanimity" and asserted that, "To have maintained in the position of a master, one side of a question for thirty years, and then deliberately give it up, is a fact to which I much doubt whether the records of science offer a parallel." ("L.L." II. pages 229-30.)

Of Darwin himself, I can safely affirm that I never knew anyone who had met him, even for the briefest period, who was not charmed by his personality. Who could forget the hearty hand-grip at meeting, the gentle and lingering pressure of the palm at parting, and above all that winning smile which transformed his countenance--so as to make portraits, and even photographs, seem ever afterwards unsatisfying! Looking back, one is indeed tempted to forget the profoundness of the philosopher, in recollection of the loveableness of the man.

XIX. DARWIN'S WORK ON THE MOVEMENTS OF PLANTS. By Francis Darwin,

Honorary Fellow of Christ's College, Cambridge.

My father's interest in plants was of two kinds, which may be roughly distinguished as EVOLUTIONARY and PHYSIOLOGICAL. Thus in his purely evolutionary work, for instance in "The Origin of Species" and in his book on "Variation under Domestication", plants as well as animals served as material for his generalisations. He was largely dependent on the work of others for the facts used in the evolutionary work, and despised himself for belonging to the "blessed gang" of compilers. And he correspondingly rejoiced in the employment of his wonderful power of observation in the physiological problems which occupied so much of his later life. But inasmuch as he felt evolution to be his life's work, he regarded himself as something of an idler in observing climbing plants, insectivorous plants, orchids, etc. In this physiological work he was to a large extent urged on by his passionate desire to understand the machinery of all living things. But though it is true that he worked at physiological problems in the naturalist's spirit of curiosity, yet there was always present to him the bearing of his facts on the problem of evolution. His interests, physiological and evolutionary, were indeed so interwoven that they cannot be sharply separated. Thus his original interest in the fertilisation of flowers was evolutionary. "I was led" ("Life and Letters", I. page 90.), he says, "to attend to the cross-fertilisation of flowers by the aid of insects, from having come to the conclusion in my speculations on the origin of species, that crossing played an important part in keeping specific forms constant." In the same way the value of his experimental work on heterostyled plants crystalised out in his mind into the conclusion that the product of illegitimate unions are equivalent to hybrids--a conclusion of the greatest interest from an evolutionary point of view. And again his work "Cross and Self Fertilisation" may be condensed to a point of view of great importance in reference to the meaning and origin of sexual reproduction. (See Professor Goebel's article in the present volume.)

The whole of his physiological work may be looked at as an illustration of the potency of his theory as an "instrument for the extension of the realm of natural knowledge." (Huxley in Darwin's "Life and Letters." II. page 204.)

His doctrine of natural selection gave, as is well known, an impulse to the investigation of the use of organs--and thus created the great school of what is known in Germany as Biology--a department of science for which no English word exists except the rather vague term Natural History. This was especially the case in floral biology, and it is interesting to see with what hesitation he at first expressed the value of his book on Orchids ("Life and Letters", III. page 254.), "It will perhaps serve to illustrate how Natural History may be worked under the belief of the modification of species" (1861). And in 1862 he speaks (Loc. cit.) more definitely of the relation of his work to natural selection: "I can show the meaning of some of the apparently meaningless ridges (and) horns; who will now venture to say that this or that structure is useless?" It is the fashion now to minimise the value of this class of work, and we even find it said by a modern writer that to inquire into the ends subserved by organs is not a scientific problem. Those who take this view surely forget that the structure of all living things is, as a whole, adaptive, and that a knowledge of how the present forms come to be what they are includes a knowledge of why they survived. They forget that the SUMMATION of variations on which divergence depends is under the rule of the environment considered as a selective force. They forget that the scientific study of the interdependence of organisms is only possible through a knowledge of the machinery of the units. And that, therefore, the investigation of such widely interesting subjects as extinction and distribution must include a knowledge of function. It is only those who follow this line of work who get to see the importance of minute points of structure and understand as my father did even in 1842, as shown in his sketch of the "Origin" (Now being prepared for publication.), that every grain of sand counts for something in the balance. Much that is confidently stated about the uselessness of different organs would never have been written if the naturalist spirit were commoner nowadays. This spirit is strikingly shown in my father's work on the movements of plants. The circumstance that botanists had not, as a class, realised the interest of the subject accounts for the fact that he was able to gather such a rich harvest of results from such a familiar object as a twining plant. The subject had been investigated by H. von Mohl, Palm, and Dutrochet, but they failed not only to master the problem but (which here concerns us) to give the absorbing interest of Darwin's book to what they discovered.

His work on climbing plants was his first sustained piece of work on the physiology of movement, and he remarks in 1864: "This has been new sort of work for me." ("Life and Letters", III. page 315. He had, however, made a beginning on the movements of Drosera.) He goes on to remark with something of surprise, "I have been pleased to find what a capital guide for observations a full conviction of the change of species is."

It was this point of view that enabled him to develop a broad conception of the power of climbing as an adaptation by means of which plants are enabled to reach the light. Instead of being compelled to construct a stem of sufficient strength to stand alone, they succeed in the struggle by making use of other plants as supports. He showed that the great class of tendril- and root-climbers which do not depend on twining round a pole, like a scarlet-runner, but on attaching themselves as they grow upwards, effect an economy. Thus a Phaseolus has to manufacture a stem three feet in length to reach a height of two feet above the ground, whereas a pea "which had ascended to the same height by the aid of its tendrils, was but little longer than the height reached." ("Climbing Plants" (2nd edition 1875), page 193.)

Thus he was led on to the belief that TWINING is the more ancient form of climbing, and that tendril-climbers have been developed from twiners. In accordance with this view we find LEAF-CLIMBERS, which may be looked on as incipient tendril-bearers, occurring in the same genera with simple twiners. (Loc. cit. page 195.) He called attention to the case of Maurandia semperflorens in which the young flower-stalks revolve spontaneously and are sensitive to a touch, but neither of these qualities is of any perceptible value to the species. This forced him to believe that in other young plants the rudiments of the faculty needed for twining would be found--a prophecy which he made good in his "Power of Movement" many years later.

In "Climbing Plants" he did little more than point out the remarkable fact that the habit of climbing is widely scattered through the vegetable kingdom. Thus climbers are to be found in 35 out of the 59 Phanerogamic Alliances of Lindley, so that "the conclusion is forced on our minds that the capacity of revolving (If a twining plant, e.g. a hop, is observed before it has begun to ascend a pole, it will be noticed that, owing to the curvature of the stem, the tip is not vertical but hangs over in a roughly horizontal position. If such a shoot is watched it will be found that if, for instance, it points to the north at a given hour, it will be found after a short interval pointing north-east, then east, and after about two hours it will once more be looking northward. The curvature of the stem depends on one side growing quicker than the opposite side, and the revolving movement, i.e. circumnutation, depends on the region of quickest growth creeping gradually round the stem from south through west to south again. Other plants, e.g. Phaseolus, revolve in the opposite direction.), on which most climbers depend, is inherent, though undeveloped, in almost every plant in the vegetable kingdom." ("Climbing Plants", page 205.)

In the "Origin" (Edition I. page 427, Edition VI. page 374.) Darwin speaks of the "apparent paradox, that the very same characters are analogical when one class or order is compared with another, but give true affinities when the members of the same class or order are compared one with another." In this way we might perhaps say that the climbing of an ivy and a hop are analogical; the resemblance depending on the adaptive result rather than on community of blood; whereas the relation between a leaf-climber and a true tendril-bearer reveals descent. This particular resemblance was one in which my father took especial delight. He has described an interesting case occurring in the Fumariaceae. ("Climbing Plants", page 195.) "The terminal leaflets of the leaf-climbing Fumaria officinalis are not smaller than the other leaflets; those of the leaf-climbing Adlumia cirrhosa are greatly reduced; those of Corydalis claviculata (a plant which may be indifferently called a leaf-climber or a tendril-bearer) are either reduced to microscopical dimensions or have their blades wholly aborted, so that this plant is actually in a state of transition; and finally in the Dicentra the tendrils are perfectly characterized."

It is a remarkable fact that the quality which, broadly speaking, forms the basis of the climbing habit (namely revolving nutation, otherwise known as circumnutation) subserves two distinct ends. One of these is the finding of a support, and this is common to twiners and tendrils. Here the value ends as far as tendril-climbers are concerned, but in twiners Darwin believed that the act of climbing round a support is a continuation of the revolving movement (circumnutation). If we imagine a man swinging a rope round his head and if we suppose the rope to strike a vertical post, the free end will twine round it. This may serve as a rough model of twining as explained in the "Movements and Habits of Climbing Plants". It is on these points--the nature of revolving nutation and the mechanism of twining--that modern physiologists differ from Darwin. (See the discussion in Pfeffer's "The Physiology of Plants" Eng. Tr. (Oxford, 1906), III. page 34, where the literature is given. Also Jost, "Vorlesungen uber Pflanzenphysiologie", page 562, Jena, 1904.)

Their criticism originated in observations made on a revolving shoot which is removed from the action of gravity by keeping the plant slowly rotating about a horizontal axis by means of the instrument known as a klinostat. Under these conditions circumnutation becomes irregular or ceases altogether. When the same experiment is made with a plant which has twined spirally up a stick, the process of climbing is checked and the last few turns become loosened or actually untwisted. From this it has been argued that Darwin was wrong in his description of circumnutation as an automatic change in the region of quickest growth. When the free end of a revolving shoot points towards the north there is no doubt that the south side has been elongating more than the north; after a time it is plain from the shoot hanging over to the east that the west side of the plant has grown most, and so on. This rhythmic change of the position of the region of greatest growth Darwin ascribes to an unknown internal regulating power. Some modern physiologists, however, attempt to explain the revolving movement as due to a particular form of sensitiveness to gravitation which it is not necessary to discuss in detail in this place. It is sufficient for my purpose to point out that Darwin's explanation of circumnutation is not universally accepted. Personally I believe that circumnutation is automatic--is primarily due to internal stimuli. It is however in some way connected with gravitational sensitiveness, since the movement normally occurs round a vertical line. It is not unnatural that, when the plant has no external stimulus by which the vertical can be recognised, the revolving movement should be upset.

Very much the same may be said of the act of twining, namely that most physiologists refuse to accept Darwin's view (above referred to) that twining is the direct result of circumnutation. Everyone must allow that the two phenomena are in some way connected, since a plant which circumnutates clockwise, i.e. with the sun, twines in the same direction, and vice versa. It must also be granted that geotropism has a bearing on the problem, since all plants twine upwards, and cannot twine along a horizontal support. But how these two factors are combined, and whether any (and if so what) other factors contribute, we cannot say. If we give up Darwin's explanation, we must at the same time say with Pfeffer that "the causes of twining are... unknown." ("The Physiology of Plants", Eng. Tr. (Oxford, 1906), III. page 37.)

Let us leave this difficult question and consider some other points made out in the progress of the work on climbing plants. One result of what he called his "niggling" ("Life and Letters", III. page 312.) work on tendrils was the discovery of the delicacy of their sense of touch, and the rapidity of their movement. Thus in a passion-flower tendril, a bit of platinum wire weighing 1.2 mg. produced curvature ("Climbing Plants", page 171.), as did a loop of cotton weighing 2 mg. Pfeffer ("Untersuchungen a.d. Bot. Inst. z. Tubingen", Bd. I. 1881-85, page 506.), however, subsequently found much greater sensitiveness: thus the tendril of Sicyos angulatus reacted to 0.00025 mg., but this only occurred when the delicate rider of cottonwool fibre was disturbed by the wind. The same author expanded and explained in a most interesting way the meaning of Darwin's observation that tendrils are not stimulated to movement by drops of water resting on them. Pfeffer showed that DIRTY water containing minute particles of clay in suspension acts as a stimulus. He also showed that gelatine acts like pure water; if a smooth glass rod is coated with a 10 per cent solution of gelatine and is then applied to a tendril, no movement occurs in spite of the fact that the gelatine is solid when cold. Pfeffer ("Physiology", Eng. Tr. III. page 52. Pfeffer has pointed out the resemblance between the contact irritability of plants and the human sense of touch. Our skin is not sensitive to uniform pressure such as is produced when the finger is dipped into mercury (Tubingen "Untersuchungen", I. page 504.) generalises the result in the statement that the tendril has a special form of irritability and only reacts to "differences of pressure or variations of pressure in contiguous... regions." Darwin was especially interested in such cases of specialised irritability. For instance in May, 1864, he wrote to Asa Gray ("Life and Letters", III. page 314.) describing the tendrils of Bignonia capreolata, which "abhor a simple stick, do not much relish rough bark, but delight in wool or moss." He received, from Gray, information as to the natural habitat of the species, and finally concluded that the tendrils "are specially adapted to climb trees clothed with lichens, mosses, or other such productions." ("Climbing Plants", page 102.)

Tendrils were not the only instance discovered by Darwin of delicacy of touch in plants. In 1860 he had already begun to observe Sundew (Drosera), and was full of astonishment at its behaviour. He wrote to Sir Joseph Hooker ("Life and Letters", III. page 319.): "I have been working like a madman at Drosera. Here is a fact for you which is certain as you stand where you are, though you won't believe it, that a bit of hair 1/78000 of one grain in weight placed on gland, will cause ONE of the gland-bearing hairs of Drosera to curve inwards." Here again Pfeffer (Pfeffer in "Untersuchungen a. d. Bot. Inst. z. Tubingen", I. page 491.) has, as in so many cases, added important facts to my father's observations. He showed that if the leaf of Drosera is entirely freed from such vibrations as would reach it if observed on an ordinary table, it does not react to small weights, so that in fact it was the vibration of the minute fragment of hair on the gland that produced movement. We may fancifully see an adaptation to the capture of insects--to the dancing of a gnat's foot on the sensitive surface.

Darwin was fond of telling how when he demonstrated the sensitiveness of Drosera to Mr Huxley and (I think) to Sir John Burdon Sanderson, he could perceive (in spite of their courtesy) that they thought the whole thing a delusion. And the story ended with his triumph when Mr Huxley cried out, "It IS moving."

Darwin's work on tendrils has led to some interesting investigations on the mechanisms by which plants perceive stimuli. Thus Pfeffer (Tubingen "Untersuchungen" I. page 524.) showed that certain epidermic cells occurring in tendrils are probably organs of touch. In these cells the protoplasm burrows as it were into cavities in the thickness of the external cell-walls and thus comes close to the surface, being separated from an object touching the tendril merely by a very thin layer of cell-wall substance. Haberlandt ("Physiologische Pflanzenanatomie", Edition III. Leipzig, 1904. "Sinnesorgane im Pflanzenreich", Leipzig, 1901, and other publications.) has greatly extended our knowledge of vegetable structure in relation to mechanical stimulation. He defines a sense-organ as a contrivance by which the DEFORMATION or forcible change of form in the protoplasm--on which mechanical stimulation depends--is rendered rapid and considerable in amplitude ("Sinnesorgane", page 10). He has shown that in certain papillose and bristle-like contrivances, plants possess such sense-organs; and moreover that these contrivances show a remarkable similarity to corresponding sense-organs in animals.

Haberlandt and Nemec ("Ber. d. Deutschen bot. Gesellschaft", XVIII. 1900. See F. Darwin, Presidential Address to Section K, British Association, 1904.) published independently and simultaneously a theory of the mechanism by which plants are orientated in relation to gravitation. And here again we find an arrangement identical in principle with that by which certain animals recognise the vertical, namely the pressure of free particles on the irritable wall of a cavity. In the higher plants, Nemec and Haberlandt believe that special loose and freely movable starch-grains play the part of the otoliths or statoliths of the crustacea, while the protoplasm lining the cells in which they are contained corresponds to the sensitive membrane lining the otocyst of the animal. What is of special interest in our present connection is that according to this ingenious theory (The original conception was due to Noll ("Heterogene Induction", Leipzig, 1892), but his view differed in essential points from those here given.) the sense of verticality in a plant is a form of contact-irritability. The vertical position is distinguished from the horizontal by the fact that, in the latter case, the loose starch-grains rest on the lateral walls of the cells instead of on the terminal walls as occurs in the normal upright position. It should be added that the statolith theory is still sub judice; personally I cannot doubt that it is in the main a satisfactory explanation of the facts.

With regard to the RAPIDITY of the reaction of tendrils, Darwin records ("Climbing Plants", page 155. Others have observed movement after about 6".) that a Passion-Flower tendril moved distinctly within 25 seconds of stimulation. It was this fact, more than any other, that made him doubt the current explanation, viz. that the movement is due to unequal growth on the two sides of the tendril. The interesting work of Fitting (Pringsheim's "Jahrb." XXXVIII. 1903, page 545.) has shown, however, that the primary cause is not (as Darwin supposed) contraction on the concave, but an astonishingly rapid increase in growth-rate on the convex side.

On the last page of "Climbing Plants" Darwin wrote: "It has often been vaguely asserted that plants are distinguished from animals by not having the power of movement. It should rather be said that plants acquire and display this power only when it is of some advantage to them."

He gradually came to realise the vividness and variety of vegetable life, and that a plant like an animal has capacities of behaving in different ways under different circumstances, in a manner that may be compared to the instinctive movements of animals. This point of view is expressed in well-known passages in the "Power of Movement". ("The Power of Movement in Plants", 1880, pages 571-3.) "It is impossible not to be struck with the resemblance between the... movements of plants and many of the actions performed unconsciously by the lower animals." And again, "It is hardly an exaggeration to say that the tip of the radicle... having the power of directing the movements of the adjoining parts, acts like the brain of one of the lower animals; the brain being seated within the anterior end of the body, receiving impressions from the sense-organs, and directing the several movements."

The conception of a region of perception distinct from a region of movement is perhaps the most fruitful outcome of his work on the movements of plants. But many years before its publication, viz. in 1861, he had made out the wonderful fact that in the Orchid Catasetum ("Life and Letters", III. page 268.) the projecting organs or antennae are sensitive to a touch, and transmit an influence "for more than one inch INSTANTANEOUSLY," which leads to the explosion or violent ejection of the pollinia. And as we have already seen a similar transmission of a stimulus was discovered by him in Sundew in 1860, so that in 1862 he could write to Hooker ("Life and Letters", III. page 321.): "I cannot avoid the conclusion, that Drosera possesses matter at least in some degree analogous in constitution and function to nervous matter." I propose in what follows to give some account of the observations on the transmission of stimuli given in the "Power of Movement". It is impossible within the space at my command to give anything like a complete account of the matter, and I must necessarily omit all mention of much interesting work. One well-known experiment consisted in putting opaque caps on the tips of seedling grasses (e.g. oat and canary-grass) and then exposing them to light from one side. The difference, in the amount of curvature towards the light, between the blinded and unblinded specimens, was so great that it was concluded that the light-sensitiveness resided exclusively in the tip. The experiment undoubtedly proves that the sensitiveness is much greater in the tip than elsewhere, and that there is a transmission of stimulus from the tip to the region of curvature. But Rothert (Rothert, Cohn's "Beitrage", VII. 1894.) has conclusively proved that the basal part where the curvature occurs is also DIRECTLY sensitive to light. He has shown, however, that in other grasses (Setaria, Panicum) the cotyledon is the only part which is sensitive, while the hypocotyl, where the movement occurs, is not directly sensitive.

It was however the question of the localisation of the gravitational sense in the tip of the seedling root or radicle that aroused most attention, and it was on this question that a controversy arose which has continued to the present day.

The experiment on which Darwin's conclusion was based consisted simply in cutting off the tip, and then comparing the behaviour of roots so treated with that of normal specimens. An uninjured root when placed horizontally regains the vertical by means of a sharp downward curve; not so a decapitated root which continues to grow more or less horizontally. It was argued that this depends on the loss of an organ specialised for the perception of gravity, and residing in the tip of the root; and the experiment (together with certain important variants) was claimed as evidence of the existence of such an organ.

It was at once objected that the amputation of the tip might check curvature by interfering with longitudinal growth, on the distribution of which curvature depends. This objection was met by showing that an injury, e.g. splitting the root longitudinally (See F. Darwin, "Linnean Soc. Journal (Bot)." XIX. 1882, page 218.), which does not remove the tip, but seriously checks growth, does not prevent geotropism. This was of some interest in another and more general way, in showing that curvature and longitudinal growth must be placed in different categories as regards the conditions on which they depend.

Another objection of a much more serious kind was that the amputation of the tip acts as a shock. It was shown by Rothert (See his excellent summary of the subject in "Flora" 1894 (Erganzungsband), page 199.) that the removal of a small part of the cotyledon of Setaria prevents the plant curving towards the light, and here there is no question of removing the sense-organ since the greater part of the sensitive cotyledon is intact. In view of this result it was impossible to rely on the amputations performed on roots as above described.

At this juncture a new and brilliant method originated in Pfeffer's laboratory. (See Pfeffer, "Annals of Botany", VIII. 1894, page 317, and Czapek, Pringsheim's "Jahrb." XXVII. 1895, page 243.) Pfeffer and Czapek showed that it is possible to bend the root of a lupine so that, for instance, the supposed sense-organ at the tip is vertical while the motile region is horizontal. If the motile region is directly sensitive to gravity the root ought to curve downwards, but this did not occur: on the contrary it continued to grow horizontally. This is precisely what should happen if Darwin's theory is the right one: for if the tip is kept vertical, the sense-organ is in its normal position and receives no stimulus from gravitation, and therefore can obviously transmit none to the region of curvature. Unfortunately this method did not convince the botanical world because some of those who repeated Czapek's experiment failed to get his results.

Czapek ("Berichte d. Deutsch. bot. Ges." XV. 1897, page 516, and numerous subsequent papers. English readers should consult Czapek in the "Annals of Botany", XIX. 1905, page 75.) has devised another interesting method which throws light on the problem. He shows that roots, which have been placed in a horizontal position and have therefore been geotropically stimulated, can be distinguished by a chemical test from vertical, i.e. unstimulated roots. The chemical change in the root can be detected before any curvature has occurred and must therefore be a symptom of stimulation, not of movement. It is particularly interesting to find that the change in the root, on which Czapek's test depends, takes place in the tip, i.e. in the region which Darwin held to be the centre for gravitational sensitiveness.

In 1899 I devised a method (F. Darwin, "Annals of Botany", XIII. 1899, page 567.) by which I sought to prove that the cotyledon of Setaria is not only the organ for light-perception, but also for gravitation. If a seedling is supported horizontally by pushing the apical part (cotyledon) into a horizontal tube, the cotyledon will, according to my supposition, be stimulated gravitationally and a stimulus will be transmitted to the basal part of the stem (hypocotyl) causing it to bend. But this curvature merely raises the basal end of the seedling, the sensitive cotyledon remains horizontal, imprisoned in its tube; it will therefore be continually stimulated and will continue to transmit influences to the bending region, which should therefore curl up into a helix or corkscrew-like form,--and this is precisely what occurred.

I have referred to this work principally because the same method was applied to roots by Massart (Massart, "Mem. Couronnes Acad. R. Belg." LXII. 1902.) and myself (F. Darwin, "Linnean Soc. Journ." XXXV. 1902, page 266.) with a similar though less striking result. Although these researches confirmed Darwin's work on roots, much stress cannot be laid on them as there are several objections to them, and they are not easily repeated.

The method which--as far as we can judge at present--seems likely to solve the problem of the root-tip is most ingenious and is due to Piccard. (Pringsheim's "Jahrb." XL. 1904, page 94.)

Andrew Knight's celebrated experiment showed that roots react to centrifugal force precisely as they do to gravity. So that if a bean root is fixed to a wheel revolving rapidly on a horizontal axis, it tends to curve away from the centre in the line of a radius of the wheel. In ordinary demonstrations of Knight's experiment the seed is generally fixed so that the root is at right angles to a radius, and as far as convenient from the centre of rotation. Piccard's experiment is arranged differently. (A seed is depicted below a horizontal dotted line AA, projecting a root upwards.) The root is oblique to the axis of rotation, and the extreme tip projects beyond that axis. Line AA represents the axis of rotation, T is the tip of the root just above the line AA, and B is the region just below line AA in which curvature takes place. If the motile region B is directly sensitive to gravitation (and is the only part which is sensitive) the root will curve (down and away from the vertical) away from the axis of rotation, just as in Knight's experiment. But if the tip T is alone sensitive to gravitation the result will be exactly reversed, the stimulus originating in T and conveyed to B will produce curvature (up towards the vertical). We may think of the line AA as a plane dividing two worlds. In the lower one gravity is of the earthly type and is shown by bodies falling and roots curving downwards: in the upper world bodies fall upwards and roots curve in the same direction. The seedling is in the lower world, but its tip containing the supposed sense-organ is in the strange world where roots curve upwards. By observing whether the root bends up or down we can decide whether the impulse to bend originates in the tip or in the motile region.

Piccard's results showed that both curvatures occurred and he concluded that the sensitive region is not confined to the tip. (Czapek (Pringsheim's "Jahrb." XXXV. 1900, page 362) had previously given reasons for believing that, in the root, there is no sharp line of separation between the regions of perception and movement.)

Haberlandt (Pringsheim's "Jahrb." XLV. 1908, page 575.) has recently repeated the experiment with the advantage of better apparatus and more experience in dealing with plants, and has found as Piccard did that both the tip and the curving region are sensitive to gravity, but with the important addition that the sensitiveness of the tip is much greater than that of the motile region. The case is in fact similar to that of the oat and canary-grass. In both instances my father and I were wrong in assuming that the sensitiveness is confined to the tip, yet there is a concentration of irritability in that region and transmission of stimulus is as true for geotropism as it is for heliotropism. Thus after nearly thirty years the controversy of the root-tip has apparently ended somewhat after the fashion of the quarrels at the "Rainbow" in "Silas Marner"--"you're both right and you're both wrong." But the "brain-function" of the root-tip at which eminent people laughed in early days turns out to be an important part of the truth. (By using Piccard's method I have succeeded in showing that the gravitational sensitiveness of the cotyledon of Sorghum is certainly much greater than the sensitiveness of the hypocotyl--if indeed any such sensitiveness exists. See Wiesner's "Festschrift", Vienna, 1908.)

Another observation of Darwin's has given rise to much controversy. ("Power of Movement", page 133.) If a minute piece of card is fixed obliquely to the tip of a root some influence is transmitted to the region of curvature and the root bends away from the side to which the card was attached. It was thought at the time that this proved the root-tip to be sensitive to contact, but this is not necessarily the case. It seems possible that the curvature is a reaction to the injury caused by the alcoholic solution of shellac with which the cards were cemented to the tip. This agrees with the fact given in the "Power of Movement" that injuring the root-tip on one side, by cutting or burning it, induced a similar curvature. On the other hand it was shown that curvature could be produced in roots by cementing cards, not to the naked surface of the root-tip, but to pieces of gold-beaters skin applied to the root; gold-beaters skin being by itself almost without effect. But it must be allowed that, as regards touch, it is not clear how the addition of shellac and card can increase the degree of contact. There is however some evidence that very close contact from a solid body, such as a curved fragment of glass, produces curvature: and this may conceivably be the explanation of the effect of gold-beaters skin covered with shellac. But on the whole it is perhaps safer to classify the shellac experiments with the results of undoubted injury rather than with those of contact.

Another subject on which a good deal of labour was expended is the sleep of leaves, or as Darwin called it their NYCTITROPIC movement. He showed for the first time how widely spread this phenomenon is, and attempted to give an explanation of the use to the plant of the power of sleeping. His theory was that by becoming more or less vertical at night the leaves escape the chilling effect of radiation. Our method of testing this view was to fix some of the leaves of a sleeping plant so that they remained horizontal at night and therefore fully exposed to radiation, while their fellows were partly protected by assuming the nocturnal position. The experiments showed clearly that the horizontal leaves were more injured than the sleeping, i.e. more or less vertical, ones. It may be objected that the danger from cold is very slight in warm countries where sleeping plants abound. But it is quite possible that a lowering of the temperature which produces no visible injury may nevertheless be hurtful by checking the nutritive processes (e.g. translocation of carbohydrates), which go on at night. Stahl ("Bot. Zeitung", 1897, page 81.) however has ingeniously suggested that the exposure of the leaves to radiation is not DIRECTLY hurtful because it lowers the temperature of the leaf, but INDIRECTLY because it leads to the deposition of dew on the leaf-surface. He gives reasons for believing that dew-covered leaves are unable to transpire efficiently, and that the absorption of mineral food-material is correspondingly checked. Stahl's theory is in no way destructive of Darwin's, and it is possible that nyctitropic leaves are adapted to avoid the indirect as well as the direct results of cooling by radiation.

In what has been said I have attempted to give an idea of some of the discoveries brought before the world in the "Power of Movement" (In 1881 Professor Wiesner published his "Das Bewegungsvermogen der Pflanzen", a book devoted to the criticism of "The Power of Movement in Plants". A letter to Wiesner, published in "Life and Letters", III. page 336, shows Darwin's warm appreciation of his critic's work, and of the spirit in which it is written.) and of the subsequent history of the problems. We must now pass on to a consideration of the central thesis of the book,--the relation of circumnutation to the adaptive curvatures of plants.

Darwin's view is plainly stated on pages 3-4 of the "Power of Movement". Speaking of circumnutation he says, "In this universally present movement we have the basis or groundwork for the acquirement, according to the requirements of the plant, of the most diversified movements." He then points out that curvatures such as those towards the light or towards the centre of the earth can be shown to be exaggerations of circumnutation in the given directions. He finally points out that the difficulty of conceiving how the capacities of bending in definite directions were acquired is diminished by his conception. "We know that there is always movement in progress, and its amplitude, or direction, or both, have only to be modified for the good of the plant in relation with internal or external stimuli."

It may at once be allowed that the view here given has not been accepted by physiologists. The bare fact that circumnutation is a general property of plants (other than climbing species) is not generally rejected. But the botanical world is no nearer to believing in the theory of reaction built on it.

If we compare the movements of plants with those of the lower animals we find a certain resemblance between the two. According to Jennings (H.S. Jennings, "The Behavior of the Lower Animals". Columbia U. Press, N.Y. 1906.) a Paramoecium constantly tends to swerve towards the aboral side of its body owing to certain peculiarities in the set and power of its cilia. But the tendency to swim in a circle, thus produced, is neutralised by the rotation of the creature about its longitudinal axis. Thus the direction of the swerves IN RELATION TO THE PATH of the organism is always changing, with the result that the creature moves in what approximates to a straight line, being however actually a spiral about the general line of progress. This method of motion is strikingly like the circumnutation of a plant, the apex of which also describes a spiral about the general line of growth. A rooted plant obviously cannot rotate on its axis, but the regular series of curvatures of which its growth consists correspond to the aberrations of Paramoecium distributed regularly about its course by means of rotation. (In my address to the Biological Section of the British Association at Cardiff (1891) I have attempted to show the connection between circumnutation and RECTIPETALITY, i.e. the innate capacity of growing in a straight line.) Just as a plant changes its direction of growth by an exaggeration of one of the curvature-elements of which circumnutation consists, so does a Paramoecium change its course by the accentuation of one of the deviations of which its path is built. Jennings has shown that the infusoria, etc., react to stimuli by what is known as the "method of trial." If an organism swims into a region where the temperature is too high or where an injurious substance is present, it changes its course. It then moves forward again, and if it is fortunate enough to escape the influence, it continues to swim in the given direction. If however its change of direction leads it further into the heated or poisonous region it repeats the movement until it emerges from its difficulties. Jennings finds in the movements of the lower organisms an analogue with what is known as pain in conscious organisms. There is certainly this much resemblance that a number of quite different sub-injurious agencies produce in the lower organisms a form of reaction by the help of which they, in a partly fortuitous way, escape from the threatening element in their environment. The higher animals are stimulated in a parallel manner to vague and originally purposeless movements, one of which removes the discomfort under which they suffer, and the organism finally learns to perform the appropriate movement without going through the tentative series of actions.

I am tempted to recognise in circumnutation a similar groundwork of tentative movements out of which the adaptive ones were originally selected by a process rudely representative of learning by experience.

It is, however, simpler to confine ourselves to the assumption that those plants have survived which have acquired through unknown causes the power of reacting in appropriate ways to the external stimuli of light, gravity, etc. It is quite possible to conceive this occurring in plants which have no power of circumnutating--and, as already pointed out, physiologists do as a fact neglect circumnutation as a factor in the evolution of movements. Whatever may be the fate of Darwin's theory of circumnutation there is no doubt that the research he carried out in support of, and by the light of, this hypothesis has had a powerful influence in guiding the modern theories of the behaviour of plants. Pfeffer ("The Physiology of Plants", Eng. Tr. III. page 11.), who more than any one man has impressed on the world a rational view of the reactions of plants, has acknowledged in generous words the great value of Darwin's work in the same direction. The older view was that, for instance, curvature towards the light is the direct mechanical result of the difference of illumination on the lighted and shaded surfaces of the plant. This has been proved to be an incorrect explanation of the fact, and Darwin by his work on the transmission of stimuli has greatly contributed to the current belief that stimuli act indirectly. Thus we now believe that in a root and a stem the mechanism for the perception of gravitation is identical, but the resulting movements are different because the motor-irritabilities are dissimilar in the two cases. We must come back, in fact, to Darwin's comparison of plants to animals. In both there is perceptive machinery by which they are made delicately alive to their environment, in both the existing survivors are those whose internal constitution has enabled them to respond in a beneficial way to the disturbance originating in their sense-organs.

XX. THE BIOLOGY OF FLOWERS. By K. Goebel, Ph.D.

Professor of Botany in the University of Munich.

There is scarcely any subject to which Darwin devoted so much time and work as to his researches into the biology of flowers, or, in other words, to the consideration of the question to what extent the structural and physiological characters of flowers are correlated with their function of producing fruits and seeds. We know from his own words what fascination these studies possessed for him. We repeatedly find, for example, in his letters expressions such as this:--"Nothing in my life has ever interested me more than the fertilisation of such plants as Primula and Lythrum, or again Anacamptis or Listera." ("More Letters of Charles Darwin", Vol. II. page 419.)

Expressions of this kind coming from a man whose theories exerted an epoch-making influence, would be unintelligible if his researches into the biology of flowers had been concerned only with records of isolated facts, however interesting these might be. We may at once take it for granted that the investigations were undertaken with the view of following up important problems of general interest, problems which are briefly dealt with in this essay.

Darwin published the results of his researches in several papers and in three larger works, (i) "On the various contrivances by which British and Foreign Orchids are fertilised by insects" (First edition, London, 1862; second edition, 1877; popular edition, 1904.) (ii) "The effects of Cross and Self fertilisation in the vegetable kingdom" (First edition, 1876; second edition, 1878). (iii) "The different forms of Flowers on plants of the same species" (First edition, 1877; second edition, 1880).

Although the influence of his work is considered later, we may here point out that it was almost without a parallel; not only does it include a mass of purely scientific observations, but it awakened interest in very wide circles, as is shown by the fact that we find the results of Darwin's investigations in floral biology universally quoted in school books; they are even willingly accepted by those who, as regards other questions, are opposed to Darwin's views.

The works which we have mentioned are, however, not only of special interest because of the facts they contribute, but because of the MANNER in which the facts are expressed. A superficial reader seeking merely for catch-words will, for instance, probably find the book on cross and self-fertilisation rather dry because of the numerous details which it contains: it is, indeed, not easy to compress into a few words the general conclusions of this volume. But on closer examination, we cannot be sufficiently grateful to the author for the exactness and objectivity with which he enables us to participate in the scheme of his researches. He never tries to persuade us, but only to convince us that his conclusions are based on facts; he always gives prominence to such facts as appear to be in opposition to his opinions,--a feature of his work in accordance with a maxim which he laid down:--"It is a golden rule, which I try to follow, to put every fact which is opposed to one's preconceived opinion in the strongest light." ("More Letters", Vol. II. page 324.)

The result of this method of presentation is that the works mentioned above represent a collection of most valuable documents even for those who feel impelled to draw from the data other conclusions than those of the author. Each investigation is the outcome of a definite question, a "preconceived opinion," which is either supported by the facts or must be abandoned. "How odd it is that anyone should not see that all observation must be for or against some view if it is to be of any service!" (Ibid. Vol. I. page 195.)

The points of view which Darwin had before him were principally the following. In the first place the proof that a large number of the peculiarities in the structure of flowers are not useless, but of the greatest significance in pollination must be of considerable importance for the interpretation of adaptations; "The use of each trifling detail of structure is far from a barren search to those who believe in natural selection." ("Fertilisation of Orchids" (1st edition), page 351; (2nd edition 1904) page 286.) Further, if these structural relations are shown to be useful, they may have been acquired because from the many variations which have occurred along different lines, those have been preserved by natural selection "which are beneficial to the organism under the complex and ever-varying conditions of life." (Ibid. page 351.) But in the case of flowers there is not only the question of adaptation to fertilisation to be considered. Darwin, indeed, soon formed the opinion which he has expressed in the following sentence,--"From my own observations on plants, guided to a certain extent by the experience of the breeders of animals, I became convinced many years ago that it is a general law of nature that flowers are adapted to be crossed, at least occasionally, by pollen from a distinct plant." ("Cross and Self fertilisation" (1st edition), page 6.)

The experience of animal breeders pointed to the conclusion that continual in-breeding is injurious. If this is correct, it raises the question whether the same conclusion holds for plants. As most flowers are hermaphrodite, plants afford much more favourable material than animals for an experimental solution of the question, what results follow from the union of nearly related sexual cells as compared with those obtained by the introduction of new blood. The answer to this question must, moreover, possess the greatest significance for the correct understanding of sexual reproduction in general.

We see, therefore, that the problems which Darwin had before him in his researches into the biology of flowers were of the greatest importance, and at the same time that the point of view from which he attacked the problems was essentially a teleological one.

We may next inquire in what condition he found the biology of flowers at the time of his first researches, which were undertaken about the year 1838. In his autobiography he writes,--"During the summer of 1839, and, I believe, during the previous summer, I was led to attend to the cross-fertilisation of flowers by the aid of insects, from having come to the conclusion in my speculations on the origin of species, that crossing played an important part in keeping specific forms constant." ("The Life and Letters of Charles Darwin", Vol. I. page 90, London, 1888.) In 1841 he became acquainted with Sprengel's work: his researches into the biology of flowers were thus continued for about forty years.

It is obvious that there could only be a biology of flowers after it had been demonstrated that the formation of seeds and fruit in the flower is dependent on pollination and subsequent fertilisation. This proof was supplied at the end of the seventeenth century by R.J. Camerarius (1665-1721). He showed that normally seeds and fruits are developed only when the pollen reaches the stigma. The manner in which this happens was first thoroughly investigated by J.G. Kolreuter (1733-1806 (Kolreuter, "Vorlaufige Nachricht von einigen das Geschlecht der Planzen betreffenden Versuchen und Beobachtungen", Leipzig, 1761; with three supplements, 1763-66. Also, "Mem. de l'acad. St Petersbourg", Vol. XV. 1809.)), the same observer to whom we owe the earliest experiments in hybridisation of real scientific interest. Kolreuter mentioned that pollen may be carried from one flower to another partly by wind and partly by insects. But he held the view, and that was, indeed, the natural assumption, that self-fertilisation usually occurs in a flower, in other words that the pollen of a flower reaches the stigma of the same flower. He demonstrated, however, certain cases in which cross-pollination occurs, that is in which the pollen of another flower of the same species is conveyed to the stigma. He was familiar with the phenomenon, exhibited by numerous flowers, to which Sprengel afterwards applied the term Dichogamy, expressing the fact that the anthers and stigmas of a flower often ripen at different times, a peculiarity which is now recognised as one of the commonest means of ensuring cross-pollination.

With far greater thoroughness and with astonishing power of observation C.K. Sprengel (1750-1816) investigated the conditions of pollination of flowers. Darwin was introduced by that eminent botanist Robert Brown to Sprengel's then but little appreciated work,--"Das entdeckte Geheimniss der Natur im Bau und in der Befruchtung der Blumen" (Berlin, 1793); this is by no means the least service to Botany rendered by Robert Brown.

Sprengel proceeded from a naive teleological point of view. He firmly believed "that the wise Author of nature had not created a single hair without a definite purpose." He succeeded in demonstrating a number of beautiful adaptations in flowers for ensuring pollination; but his work exercised but little influence on his contemporaries and indeed for a long time after his death. It was through Darwin that Sprengel's work first achieved a well deserved though belated fame. Even such botanists as concerned themselves with researches into the biology of flowers appear to have formerly attached much less value to Sprengel's work than it has received since Darwin's time. In illustration of this we may quote C.F. Gartner whose name is rightly held in the highest esteem as that of one of the most eminent hybridologists. In his work "Versuche und Beobachtungen uder die Befruchtungsorgane der vollkommeneren Gewachse und uber die naturliche und kunstliche Befruchtung durch den eigenen Pollen" he also deals with flower-pollination. He recognised the action of the wind, but he believed, in spite of the fact that he both knew and quoted Kolreuter and Sprengel, that while insects assist pollination, they do so only occasionally, and he held that insects are responsible for the conveyance of pollen; thorough investigations would show "that a very small proportion of the plants included in this category require this assistance in their native habitat." (Gartner, "Versucher und Beobachtungen... ", page 335, Stuttgart, 1844.) In the majority of plants self-pollination occurs.

Seeing that even investigators who had worked for several decades at fertilisation-phenomena had not advanced the biology of flowers beyond the initial stage, we cannot be surprised that other botanists followed to even a less extent the lines laid down by Kolreuter and Sprengel. This was in part the result of Sprengel's supernatural teleology and in part due to the fact that his book appeared at a time when other lines of inquiry exerted a dominating influence.

At the hands of Linnaeus systematic botany reached a vigorous development, and at the beginning of the nineteenth century the anatomy and physiology of plants grew from small beginnings to a flourishing branch of science. Those who concerned themselves with flowers endeavoured to investigate their development and structure or the most minute phenomena connected with fertilisation and the formation of the embryo. No room was left for the extension of the biology of flowers on the lines marked out by Kolreuter and Sprengel. Darwin was the first to give new life and a deeper significance to this subject, chiefly because he took as his starting-point the above-mentioned problems, the importance of which is at once admitted by all naturalists.

The further development of floral biology by Darwin is in the first place closely connected with the book on the fertilisation of Orchids. It is noteworthy that the title includes the sentence,--"and on the good effects of intercrossing."

The purpose of the book is clearly stated in the introduction:--"The object of the following work is to show that the contrivances by which Orchids are fertilised, are as varied and almost as perfect as any of the most beautiful adaptations in the animal kingdom; and, secondly, to show that these contrivances have for their main object the fertilisation of each flower by the pollen of another flower." ("Fertilisation of Orchids", page 1.) Orchids constituted a particularly suitable family for such researches. Their flowers exhibit a striking wealth of forms; the question, therefore, whether the great variety in floral structure bears any relation to fertilisation (In the older botanical literature the word fertilisation is usually employed in cases where POLLINATION is really in question: as Darwin used it in this sense it is so used here.) must in this case possess special interest.

Darwin succeeded in showing that in most of the orchids examined self-fertilisation is either an impossibility, or, under natural conditions, occurs only exceptionally. On the other hand these plants present a series of extraordinarily beautiful and remarkable adaptations which ensure the transference of pollen by insects from one flower to another. It is impossible to describe adequately in a few words the wealth of facts contained in the Orchid book. A few examples may, however, be quoted in illustration of the delicacy of the observations and of the perspicuity employed in interpreting the facts.

The majority of orchids differ from other seed plants (with the exception of the Asclepiads) in having no dust-like pollen. The pollen, or more correctly, the pollen-tetrads, remain fastened together as club-shaped pollinia usually borne on a slender pedicel. At the base of the pedicel is a small viscid disc by which the pollinium is attached to the head or proboscis of one of the insects which visit the flower. Darwin demonstrated that in Orchis and other flowers the pedicel of the pollinium, after its removal from the anther, undergoes a curving movement. If the pollinium was originally vertical, after a time it assumed a horizontal position. In the latter position, if the insect visited another flower, the pollinium would exactly hit the sticky stigmatic surface and thus effect fertilisation. The relation between the behaviour of the viscid disc and the secretion of nectar by the flower is especially remarkable. The flowers possess a spur which in some species (e.g. Gymnadenia conopsea, Platanthera bifolia, etc.) contains honey (nectar), which serves as an attractive bait for insects, but in others (e.g. our native species of Orchis) the spur is empty. Darwin held the opinion, confirmed by later investigations, that in the case of flowers without honey the insects must penetrate the wall of the nectarless spurs in order to obtain a nectar-like substance. The glands behave differently in the nectar-bearing and in the nectarless flowers. In the former they are so sticky that they at once adhere to the body of the insect; in the nectarless flowers firm adherence only occurs after the viscid disc has hardened. It is, therefore, adaptively of value that the insects should be detained longer in the nectarless flowers (by having to bore into the spur),--than in flowers in which the nectar is freely exposed. "If this relation, on the one hand, between the viscid matter requiring some little time to set hard, and the nectar being so lodged that moths are delayed in getting it; and, on the other hand, between the viscid matter being at first as viscid as ever it will become, and the nectar lying all ready for rapid suction, be accidental, it is a fortunate accident for the plant. If not accidental, and I cannot believe it to be accidental, what a singular case of adaptation!" ("Fertilisation of Orchids" (1st edition), page 53.)

Among exotic orchids Catasetum is particularly remarkable. One and the same species bears different forms of flowers. The species known as Catasetum tridentatum has pollinia with very large viscid discs; on touching one of the two filaments (antennae) which occur on the gynostemium of the flower the pollinia are shot out to a fairly long distance (as far as 1 metre) and in such manner that they alight on the back of the insect, where they are held. The antennae have, moreover, acquired an importance, from the point of view of the physiology of stimulation, as stimulus-perceiving organs. Darwin had shown that it is only a touch on the antennae that causes the explosion, while contact, blows, wounding, etc. on other places produce no effect. This form of flower proved to be the male. The second form, formerly regarded as a distinct species and named Monachanthus viridis, is shown to be the female flower. The anthers have only rudimentary pollinia and do not open; there are no antennae, but on the other hand numerous seeds are produced. Another type of flower, known as Myanthus barbatus, was regarded by Darwin as a third form: this was afterwards recognised by Rolfe (Rolfe, R.A. "On the sexual forms of Catasetum with special reference to the researches of Darwin and others," "Journ. Linn. Soc." Vol. XXVII. (Botany), 1891, pages 206-225.) as the male flower of another species, Catasetum barbatum Link, an identification in accordance with the discovery made by Cruger in Trinidad that it always remains sterile.

Darwin had noticed that the flowers of Catasetum do not secrete nectar, and he conjectured that in place of it the insects gnaw a tissue in the cavity of the labellum which has a "slightly sweet, pleasant and nutritious taste." This conjecture as well as other conclusions drawn by Darwin from Catasetum have been confirmed by Cruger--assuredly the best proof of the acumen with which the wonderful floral structure of this "most remarkable of the Orchids" was interpretated far from its native habitat.

As is shown by what we have said about Catasetum, other problems in addition to those concerned with fertilisation are dealt with in the Orchid book. This is especially the case in regard to flower morphology. The scope of flower morphology cannot be more clearly and better expressed than by these words: "He will see how curiously a flower may be moulded out of many separate organs--how perfect the cohesion of primordially distinct parts may become,--how organs may be used for purposes widely different from their proper function,--how other organs may be entirely suppressed, or leave mere useless emblems of their former existence." ("Fertilisation of Orchids", page 289.)

In attempting, from this point of view, to refer the floral structure of orchids to their original form, Darwin employed a much more thorough method than that of Robert Brown and others. The result of this was the production of a considerable literature, especially in France, along the lines suggested by Darwin's work. This is the so-called anatomical method, which seeks to draw conclusions as to the morphology of the flower from the course of the vascular bundles in the several parts. (He wrote in one of his letters, "... the destiny of the whole human race is as nothing to the course of vessels of orchids" ("More Letters", Vol. II. page 275.) Although the interpretation of the orchid flower given by Darwin has not proved satisfactory in one particular point--the composition of the labellum--the general results have received universal assent, namely "that all Orchids owe what they have in common to descent from some monocotyledonous plant, which, like so many other plants of the same division, possessed fifteen organs arranged alternately three within three in five whorls." ("Fertilisation of Orchids" (1st edition), page 307.) The alterations which their original form has undergone have persisted so far as they were found to be of use.

We see also that the remarkable adaptations of which we have given some examples are directed towards cross-fertilisation. In only a few of the orchids investigated by Darwin--other similar cases have since been described--was self-fertilisation found to occur regularly or usually. The former is the case in the Bee Ophrys (Ophrys apifera), the mechanism of which greatly surprised Darwin. He once remarked to a friend that one of the things that made him wish to live a few thousand years was his desire to see the extinction of the Bee Ophrys, an end to which he believed its self-fertilising habit was leading. ("Life and Letters", Vol. III. page 276 (footnote).) But, he wrote, "the safest conclusion, as it seems to me, is, that under certain unknown circumstances, and perhaps at very long intervals of time, one individual of the Bee Ophrys is crossed by another." ("Fertilisation of Orchids" page 71.)

If, on the one hand, we remember how much more sure self-fertilisation would be than cross-fertilisation, and, on the other hand, if we call to mind the numerous contrivances for cross-fertilisation, the conclusion is naturally reached that "it is an astonishing fact that self-fertilisation should not have been an habitual occurrence. It apparently demonstrates to us that there must be something injurious in the process. Nature thus tells us, in the most emphatic manner, that she abhors perpetual self-fertilisation... For may we not further infer as probable, in accordance with the belief of the vast majority of the breeders of our domestic productions, that marriage between near relations is likewise in some way injurious, that some unknown great good is derived from the union of individuals which have been kept distinct for many generations?" (Ibid., page 359.)

This view was supported by observations on plants of other families, e.g. Papilionaceae; it could, however, in the absence of experimental proof, be regarded only as a "working hypothesis."

All adaptations to cross-pollination might also be of use simply because they made pollination possible when for any reason self-pollination had become difficult or impossible. Cross-pollination would, therefore, be of use, not as such, but merely as a means of pollination in general; it would to some extent serve as a remedy for a method unsuitable in itself, such as a modification standing in the way of self-pollination, and on the other hand as a means of increasing the chance of pollination in the case of flowers in which self-pollination was possible, but which might, in accidental circumstances, be prevented. It was, therefore, very important to obtain experimental proof of the conclusion to which Darwin was led by the belief of the majority of breeders and by the evidence of the widespread occurrence of cross-pollination and of the remarkable adaptations thereto.

This was supplied by the researches which are described in the two other works named above. The researches on which the conclusions rest had, in part at least, been previously published in separate papers: this is the case as regards the heterostyled plants. The discoveries which Darwin made in the course of his investigations of these plants belong to the most brilliant in biological science.

The case of Primula is now well known. C.K. Sprengel and others were familiar with the remarkable fact that different individuals of the European species of Primula bear differently constructed flowers; some plants possess flowers in which the styles project beyond the stamens attached to the corolla-tube (long-styled form), while in others the stamens are inserted above the stigma which is borne on a short style (short-styled form). It has been shown by Breitenbach that both forms of flower may occur on the same plant, though this happens very rarely. An analogous case is occasionally met with in hybrids, which bear flowers of different colour on the same plant (e.g. Dianthus caryophyllus). Darwin showed that the external differences are correlated with others in the structure of the stigma and in the nature of the pollen. The long-styled flowers have a spherical stigma provided with large stigmatic papillae; the pollen grains are oblong and smaller than those of the short-styled flowers. The number of the seeds produced is smaller and the ovules larger, probably also fewer in number. The short-styled flowers have a smooth compressed stigma and a corolla of somewhat different form; they produce a greater number of seeds.

These different forms of flowers were regarded as merely a case of variation, until Darwin showed "that these heterostyled plants are adapted for reciprocal fertilisation; so that the two or three forms, though all are hermaphrodites, are related to one another almost like the males and females of ordinary unisexual animals." ("Forms of Flowers" (1st edition), page 2.) We have here an example of hermaphrodite flowers which are sexually different. There are essential differences in the manner in which fertilisation occurs. This may be effected in four different ways; there are two legitimate and two illegitimate types of fertilisation. The fertilisation is legitimate if pollen from the long-styled flowers reaches the stigma of the short-styled form or if pollen of the short-styled flowers is brought to the stigma of the long-styled flower, that is the organs of the same length of the two different kinds of flower react on one another. Illegitimate fertilisation is represented by the two kinds of self-fertilisation, also by cross-fertilisation, in which the pollen of the long-styled form reaches the stigma of the same type of flower and, similarly, by cross-pollination in the case of the short-styled flowers.

The applicability of the terms legitimate and illegitimate depends, on the one hand, upon the fact that insects which visit the different forms of flowers pollinate them in the manner suggested; the pollen of the short-styled flowers adhere to that part of the insect's body which touches the stigma of the long-styled flower and vice versa. On the other hand, it is based also on the fact that experiment shows that artificial pollination produces a very different result according as this is legitimate or illegitimate; only the legitimate union ensures complete fertility, the plants thus produced being stronger than those which are produced illegitimately.

If we take 100 as the number of flowers which produce seeds as the result of legitimate fertilisation, we obtain the following numbers from illegitimate fertilisation:

Primula officinalis (P. veris) (Cowslip)... 69 Primula elatior (Oxlip).................... 27 Primula acaulis (P. vulgaris) (Primrose)... 60

Further, the plants produced by the illegitimate method of fertilisation showed, e.g. in P. officinalis, a decrease in fertility in later generations, sterile pollen and in the open a feebler growth. (Under very favourable conditions (in a greenhouse) the fertility of the plants of the fourth generation increases--a point, which in view of various theoretical questions, deserves further investigation.) They behave in fact precisely in the same way as hybrids between species of different genera. This result is important, "for we thus learn that the difficulty in sexually uniting two organic forms and the sterility of their offspring, afford no sure criterion of so-called specific distinctness" ("Forms of Flowers", page 242): the relative or absolute sterility of the illegitimate unions and that of their illegitimate descendants depend exclusively on the nature of the sexual elements and on their inability to combine in a particular manner. This functional difference of sexual cells is characteristic of the behaviour of hybrids as of the illegitimate unions of heterostyled plants. The agreement becomes even closer if we regard the Primula plants bearing different forms of flowers not as belonging to a systematic entity or "species," but as including several elementary species. The legitimately produced plants are thus true hybrids (When Darwin wrote in reference to the different forms of heterostyled plants, "which all belong to the same species as certainly as do the two sexes of the same species" ("Cross and Self fertilisation", page 466), he adopted the term species in a comprehensive sense. The recent researches of Bateson and Gregory ("On the inheritance of Heterostylism in Primula"; "Proc. Roy. Soc." Ser. B, Vol. LXXVI. 1905, page 581) appear to me also to support the view that the results of illegitimate crossing of heterostyled Primulas correspond with those of hybridisation. The fact that legitimate pollen effects fertilisation, even if illegitimate pollen reaches the stigma a short time previously, also points to this conclusion. Self-pollination in the case of the short-styled form, for example, is not excluded. In spite of this, the numerical proportion of the two forms obtained in the open remains approximately the same as when the pollination was exclusively legitimate, presumably because legitimate pollen is prepotent.), with which their behaviour in other respects, as Darwin showed, presents so close an agreement. This view receives support also from the fact that descendants of a flower fertilised illegitimately by pollen from another plant with the same form of flower belong, with few exceptions, to the same type as that of their parents. The two forms of flower, however, behave differently in this respect. Among 162 seedlings of the long-styled illegitimately pollinated plants of Primula officinalis, including five generations, there were 156 long-styled and only six short-styled forms, while as the result of legitimate fertilisation nearly half of the offspring were long-styled and half short-styled. The short-styled illegitimately pollinated form gave five long-styled and nine short-styled; the cause of this difference requires further explanation. The significance of heterostyly, whether or not we now regard it as an arrangement for the normal production of hybrids, is comprehensively expressed by Darwin: "We may feel sure that plants have been rendered heterostyled to ensure cross-fertilisation, for we now know that a cross between the distinct individuals of the same species is highly important for the vigour and fertility of the offspring." ("Forms of Flowers", page 258.) If we remember how important the interpretation of heterostyly has become in all general problems as, for example, those connected with the conditions of the formation of hybrids, a fact which was formerly overlooked, we can appreciate how Darwin was able to say in his autobiography: "I do not think anything in my scientific life has given me so much satisfaction as making out the meaning of the structure of these plants." ("Life and Letters", Vol. I. page 91.)

The remarkable conditions represented in plants with three kinds of flowers, such as Lythrum and Oxalis, agree in essentials with those in Primula. These cannot be considered in detail here; it need only be noted that the investigation of these cases was still more laborious. In order to establish the relative fertility of the different unions in Lythrum salicaria 223 different fertilisations were made, each flower being deprived of its male organs and then dusted with the appropriate pollen.

In the book containing the account of heterostyled plants other species are dealt with which, in addition to flowers opening normally (chasmogamous), also possess flowers which remain closed but are capable of producing fruit. These cleistogamous flowers afford a striking example of habitual self-pollination, and H. von Mohl drew special attention to them as such shortly after the appearance of Darwin's Orchid book. If it were only a question of producing seed in the simplest way, cleistogamous flowers would be the most conveniently constructed. The corolla and frequently other parts of the flower are reduced; the development of the seed may, therefore, be accomplished with a smaller expenditure of building material than in chasmogamous flowers; there is also no loss of pollen, and thus a smaller amount suffices for fertilisation.

Almost all these plants, as Darwin pointed out, have also chasmogamous flowers which render cross-fertilisation possible. His view that cleistogamous flowers are derived from originally chasmogamous flowers has been confirmed by more recent researches. Conditions of nutrition in the broader sense are the factors which determine whether chasmogamous or cleistogamous flowers are produced, assuming, of course, that the plants in question have the power of developing both forms of flower. The former may fail to appear for some time, but are eventually developed under favourable conditions of nourishment. The belief of many authors that there are plants with only cleistogamous flowers cannot therefore be accepted as authoritative without thorough experimental proof, as we are concerned with extra-european plants for which it is often difficult to provide appropriate conditions in cultivation.

Darwin sees in cleistogamous flowers an adaptation to a good supply of seeds with a small expenditure of material, while chasmogamous flowers of the same species are usually cross-fertilised and "their offspring will thus be invigorated, as we may infer from a wide-spread analogy." ("Forms of Flowers" (1st edition), page 341.) Direct proof in support of this has hitherto been supplied in a few cases only; we shall often find that the example set by Darwin in solving such problems as these by laborious experiment has unfortunately been little imitated.

Another chapter of this book treats of the distribution of the sexes in polygamous, dioecious, and gyno-dioecious plants (the last term, now in common use, we owe to Darwin). It contains a number of important facts and discussions and has inspired the experimental researches of Correns and others.

The most important of Darwin's work on floral biology is, however, that on cross and self-fertilisation, chiefly because it states the results of experimental investigations extending over many years. Only such experiments, as we have pointed out, could determine whether cross-fertilisation is in itself beneficial, and self-fertilisation on the other hand injurious; a conclusion which a merely comparative examination of pollination-mechanisms renders in the highest degree probable. Later floral biologists have unfortunately almost entirely confined themselves to observations on floral mechanisms. But there is little more to be gained by this kind of work than an assumption long ago made by C.K. Sprengel that "very many flowers have the sexes separate and probably at least as many hermaphrodite flowers are dichogamous; it would thus appear that Nature was unwilling that any flower should be fertilised by its own pollen."

It was an accidental observation which inspired Darwin's experiments on the effect of cross and self-fertilisation. Plants of Linaria vulgaris were grown in two adjacent beds; in the one were plants produced by cross-fertilisation, that is, from seeds obtained after fertilisation by pollen of another plant of the same species; in the other grew plants produced by self-fertilisation, that is from seed produced as the result of pollination of the same flower. The first were obviously superior to the latter.

Darwin was surprised by this observation, as he had expected a prejudicial influence of self-fertilisation to manifest itself after a series of generations: "I always supposed until lately that no evil effects would be visible until after several generations of self-fertilisation, but now I see that one generation sometimes suffices and the existence of dimorphic plants and all the wonderful contrivances of orchids are quite intelligible to me." ("More Letters", Vol. II. page 373.)

The observations on Linaria and the investigations of the results of legitimate and illegitimate fertilisation in heterostyled plants were apparently the beginning of a long series of experiments. These were concerned with plants of different families and led to results which are of fundamental importance for a true explanation of sexual reproduction.

The experiments were so arranged that plants were shielded from insect-visits by a net. Some flowers were then pollinated with their own pollen, others with pollen from another plant of the same species. The seeds were germinated on moist sand; two seedlings of the same age, one from a cross and the other from a self-fertilised flower, were selected and planted on opposite sides of the same pot. They grew therefore under identical external conditions; it was thus possible to compare their peculiarities such as height, weight, fruiting capacity, etc. In other cases the seedlings were placed near to one another in the open and in this way their capacity of resisting unfavourable external conditions was tested. The experiments were in some cases continued to the tenth generation and the flowers were crossed in different ways. We see, therefore, that this book also represents an enormous amount of most careful and patient original work.

The general result obtained is that plants produced as the result of cross-fertilisation are superior, in the majority of cases, to those produced as the result of self-fertilisation, in height, resistance to external injurious influences, and in seed-production.

Ipomoea purpurea may be quoted as an example. If we express the result of cross-fertilisation by 100, we obtain the following numbers for the fertilised plants.

Generation. Height. Number of seeds.

1 100: 76 100: 64 2 100: 79 - 3 100: 68 100: 94 4 100: 86 100: 94 5 100: 75 100: 89 6 100: 72 - 7 100: 81 - 8 100: 85 - 9 100: 79 100: 26 (Number of capsules) 10 100: 54 -

Taking the average, the ratio as regards growth is 100:77. The considerable superiority of the crossed plants is apparent in the first generation and is not increased in the following generations; but there is some fluctuation about the average ratio. The numbers representing the fertility of crossed and self-fertilised plants are more difficult to compare with accuracy; the superiority of the crossed plants is chiefly explained by the fact that they produce a much larger number of capsules, not because there are on the average more seeds in each capsule. The ratio of the capsules was, e.g. in the third generation, 100:38, that of the seeds in the capsules 100:94. It is also especially noteworthy that in the self-fertilised plants the anthers were smaller and contained a smaller amount of pollen, and in the eighth generation the reduced fertility showed itself in a form which is often found in hybrids, that is the first flowers were sterile. (Complete sterility was not found in any of the plants investigated by Darwin. Others appear to be more sensitive; Cluer found Zea Mais "almost sterile" after three generations of self-fertilisation. (Cf. Fruwirth, "Die Zuchtung der Landwirtschaftlichen Kulturpflanzen", Berlin, 1904, II. page 6.))

The superiority of crossed individuals is not exhibited in the same way in all plants. For example in Eschscholzia californica the crossed seedlings do not exceed the self-fertilised in height and vigour, but the crossing considerably increases the plant's capacity for flower-production, and the seedlings from such a mother-plant are more fertile.

The conception implied by the term crossing requires a closer analysis. As in the majority of plants, a large number of flowers are in bloom at the same time on one and the same plant, it follows that insects visiting the flowers often carry pollen from one flower to another of the same stock. Has this method, which is spoken of as Geitonogamy, the same influence as crossing with pollen from another plant? The results of Darwin's experiments with different plants (Ipomoea purpurea, Digitalis purpurea, Mimulus luteus, Pelargonium, Origanum) were not in complete agreement; but on the whole they pointed to the conclusion that Geitonogamy shows no superiority over self-fertilisation (Autogamy). (Similarly crossing in the case of flowers of Pelargonium zonale, which belong to plants raised from cuttings from the same parent, shows no superiority over self-fertilisation.) Darwin, however, considered it possible that this may sometimes be the case. "The sexual elements in the flowers on the same plant can rarely have been differentiated, though this is possible, as flower-buds are in one sense distinct individuals, sometimes varying and differing from one another in structure or constitution." ("Cross and Self fertilisation" (1st edition), page 444.)

As regards the importance of this question from the point of view of the significance of cross-fertilisation in general, it may be noted that later observers have definitely discovered a difference between the results of autogamy and geitonogamy. Gilley and Fruwirth found that in Brassica Napus, the length and weight of the fruits as also the total weight of the seeds in a single fruit were less in the case of autogamy than in geitonogamy. With Sinapis alba a better crop of seeds was obtained after geitonogamy, and in the Sugar Beet the average weight of a fruit in the case of a self-fertilised plant was 0.009 gr., from geitonogamy 0.012 gr., and on cross-fertilisation 0.013 gr.

On the whole, however, the results of geitonogamy show that the favourable effects of cross-fertilisation do not depend simply on the fact that the pollen of one flower is conveyed to the stigma of another. But the plants which are crossed must in some way be different. If plants of Ipomoea purpurea (and Mimulus luteus) which have been self-fertilised for seven generations and grown under the same conditions of cultivation are crossed together, the plants so crossed would not be superior to the self-fertilised; on the other hand crossing with a fresh stock at once proves very advantageous. The favourable effect of crossing is only apparent, therefore, if the parent plants are grown under different conditions or if they belong to different varieties. "It is really wonderful what an effect pollen from a distinct seedling plant, which has been exposed to different conditions of life, has on the offspring in comparison with pollen from the same flower or from a distinct individual, but which has been long subjected to the same conditions. The subject bears on the very principle of life, which seems almost to require changes in the conditions." ("More Letters", Vol. II. page 406.)

The fertility--measured by the number or weight of the seeds produced by an equal number of plants--noticed under different conditions of fertilisation may be quoted in illustration.

On crossing On crossing On self- with a fresh plants of the fertilisation stock same stock Mimuleus luteus (First and ninth generation) 100 4 3

Eschscholzia californica (second generation) 100 45 40

Dianthus caryophyllus (third and fourth generation) 100 45 33

Petunia violacea 100 54 46

Crossing under very similar conditions shows, therefore, that the difference between the sexual cells is smaller and thus the result of crossing is only slightly superior to that given by self-fertilisation. Is, then, the favourable result of crossing with a foreign stock to be attributed to the fact that this belongs to another systematic entity or to the fact that the plants, though belonging to the same entity were exposed to different conditions? This is a point on which further researches must be taken into account, especially since the analysis of the systematic entities has been much more thorough than formerly. (In the case of garden plants, as Darwin to a large extent claimed, it is not easy to say whether two individuals really belong to the same variety, as they are usually of hybrid origin. In some instances (Petunia, Iberis) the fresh stock employed by Darwin possessed flowers differing in colour from those of the plant crossed with it.) We know that most of Linneaus's species are compound species, frequently consisting of a very large number of smaller or elementary species formerly included under the comprehensive term varieties. Hybridisation has in most cases affected our garden and cultivated plants so that they do not represent pure species but a mixture of species.

But this consideration has no essential bearing on Darwin's point of view, according to which the nature of the sexual cells is influenced by external conditions. Even individuals growing close to one another are only apparently exposed to identical conditions. Their sexual cells may therefore be differently influenced and thus give favourable results on crossing, as "the benefits which so generally follow from a cross between two plants apparently depend on the two differing somewhat in constitution or character." As a matter of fact we are familiar with a large number of cases in which the condition of the reproductive organs is influenced by external conditions. Darwin has himself demonstrated this for self-sterile plants, that is plants in which self-fertilisation produces no result. This self-sterility is affected by climatic conditions: thus in Brazil Eschscholzia californica is absolutely sterile to the pollen of its own flowers; the descendants of Brazilian plants in Darwin's cultures were partially self-fertile in one generation and in a second generation still more so. If one has any doubt in this case whether it is a question of the condition of the style and stigma, which possibly prevents the entrance of the pollen-tube or even its development, rather than that of the actual sexual cells, in other cases there is no doubt that an influence is exerted on the latter.

Janczewski (Janczewski, "Sur les antheres steriles des Groseilliers", "Bull. de l'acad. des sciences de Cracovie", June, 1908.) has recently shown that species of Ribes cultivated under unnatural conditions frequently produce a mixed (i.e. partly useless) or completely sterile pollen, precisely as happens with hybrids. There are, therefore, substantial reasons for the conclusion that conditions of life exert an influence on the sexual cells. "Thus the proposition that the benefit from cross-fertilisation depends on the plants which are crossed having been subjected during previous generations to somewhat different conditions, or to their having varied from some unknown cause as if they had been thus subjected, is securely fortified on all sides." ("Cross and Self fertilisation" (1st edition), page 444.)

We thus obtain an insight into the significance of sexuality. If an occasional and slight alteration in the conditions under which plants and animals live is beneficial (Reasons for this are given by Darwin in "Variation under Domestication" (2nd edition), Vol. II. page 127.), crossing between organisms which have been exposed to different conditions becomes still more advantageous. The entire constitution is in this way influenced from the beginning, at a time when the whole organisation is in a highly plastic state. The total life-energy, so to speak, is increased, a gain which is not produced by asexual reproduction or by the union of sexual cells of plants which have lived under the same or only slightly different conditions. All the wonderful arrangements for cross-fertilisation now appear to be useful adaptations. Darwin was, however, far from giving undue prominence to this point of view, though this has been to some extent done by others. He particularly emphasised the following consideration:--"But we should always keep in mind that two somewhat opposed ends have to be gained; the first and more important one being the production of seeds by any means, and the second, cross-fertilisation." ("Cross and Self fertilisation" (1st edition), page 371.) Just as in some orchids and cleistogamic flowers self-pollination regularly occurs, so it may also occur in other cases. Darwin showed that Pisum sativum and Lathyrus odoratus belong to plants in which self-pollination is regularly effected, and that this accounts for the constancy of certain sorts of these plants, while a variety of form is produced by crossing. Indeed among his culture plants were some which derived no benefit from crossing. Thus in the sixth self-fertilised generation of his Ipomoea cultures the "Hero" made its appearance, a form slightly exceeding its crossed companion in height; this was in the highest degree self-fertile and handed on its characteristics to both children and grandchildren. Similar forms were found in Mimulus luteus and Nicotiana (In Pisum sativum also the crossing of two individuals of the same variety produced no advantage; Darwin attributed this to the fact that the plants had for several generations been self-fertilised and in each generation cultivated under almost the same conditions. Tschermak ("Ueber kunstliche Kreuzung an Pisum sativum") afterwards recorded the same result; but he found on crossing different varieties that usually there was no superiority as regards height over the products of self-fertilisation, while Darwin found a greater height represented by the ratios 100:75 and 100:60.), types which, after self-fertilisation, have an enhanced power of seed-production and of attaining a greater height than the plants of the corresponding generation which are crossed together and self-fertilised and grown under the same conditions. "Some observations made on other plants lead me to suspect that self-fertilisation is in some respects beneficial; although the benefit thus derived is as a rule very small compared with that from a cross with a distinct plant." ("Cross and Self fertilisation", page 350.) We are as ignorant of the reason why plants behave differently when crossed and self-fertilised as we are in regard to the nature of the differentiation of the sexual cells, which determines whether a union of the sexual cells will prove favourable or unfavourable.

It is impossible to discuss the different results of cross-fertilisation; one point must, however, be emphasised, because Darwin attached considerable importance to it. It is inevitable that pollen of different kinds must reach the stigma. It was known that pollen of the same "species" is dominant over the pollen of another species, that, in other words, it is prepotent. Even if the pollen of the same species reaches the stigma rather later than that of another species, the latter does not effect fertilisation.

Darwin showed that the fertilising power of the pollen of another variety or of another individual is greater than that of the plant's own pollen. ("Cross and Self fertilisation", page 391.) This has been demonstrated in the case of Mimulus luteus (for the fixed white-flowering variety) and Iberis umbellata with pollen of another variety, and observations on cultivated plants, such as cabbage, horseradish, etc. gave similar results. It is, however, especially remarkable that pollen of another individual of the same variety may be prepotent over the plant's own pollen. This results from the superiority of plants crossed in this manner over self-fertilised plants. "Scarcely any result from my experiments has surprised me so much as this of the prepotency of pollen from a distinct individual over each plant's own pollen, as proved by the greater constitutional vigour of the crossed seedlings." (Ibid. page 397.) Similarly, in self-fertile plants the flowers of which have not been deprived of the male organs, pollen brought to the stigma by the wind or by insects from another plant effects fertilisation, even if the plant's own pollen has reached the stigma somewhat earlier.

Have the results of his experimental investigations modified the point of view from which Darwin entered on his researches, or not? In the first place the question is, whether or not the opinion expressed in the Orchid book that there is "Something injurious" connected with self-fertilisation, has been confirmed. We can, at all events, affirm that Darwin adhered in essentials to his original position; but self-fertilisation afterwards assumed a greater importance than it formerly possessed. Darwin emphasised the fact that "the difference between the self-fertilised and crossed plants raised by me cannot be attributed to the superiority of the crossed, but to the inferiority of the self-fertilised seedlings, due to the injurious effects of self-fertilisation." (Ibid. page 437.) But he had no doubt that in favourable circumstances self-fertilised plants were able to persist for several generations without crossing. An occasional crossing appears to be useful but not indispensable in all cases; its sporadic occurrence in plants in which self-pollination habitually occurs is not excluded. Self-fertilisation is for the most part relatively and not absolutely injurious and always better than no fertilisation. "Nature abhors perpetual self-fertilisation" (It is incorrect to say, as a writer has lately said, that the aphorism expressed by Darwin in 1859 and 1862, "Nature abhors perpetual self-fertilisation," is not repeated in his later works. The sentence is repeated in "Cross and Self fertilisation" (page 8), with the addition, "If the word perpetual had been omitted, the aphorism would have been false. As it stands, I believe that it is true, though perhaps rather too strongly expressed.") is, however, a pregnant expression of the fact that cross-fertilisation is exceedingly widespread and has been shown in the majority of cases to be beneficial, and that in those plants in which we find self-pollination regularly occurring cross-pollination may occasionally take place.

An attempt has been made to express in brief the main results of Darwin's work on the biology of flowers. We have seen that his object was to elucidate important general questions, particularly the question of the significance of sexual reproduction.

It remains to consider what influence his work has had on botanical science. That this influence has been very considerable, is shown by a glance at the literature on the biology of flowers published since Darwin wrote. Before the book on orchids was published there was nothing but the old and almost forgotten works of Kolreuter and Sprengel with the exception of a few scattered references. Darwin's investigations gave the first stimulus to the development of an extensive literature on floral biology. In Knuth's "Handbuch der Blutenbiologie" ("Handbook of Flower Pollination", Oxford, 1906) as many as 3792 papers on this subject are enumerated as having been published before January 1, 1904. These describe not only the different mechanisms of flowers, but deal also with a series of remarkable adaptations in the pollinating insects. As a fertilising rain quickly calls into existence the most varied assortment of plants on a barren steppe, so activity now reigns in a field which men formerly left deserted. This development of the biology of flowers is of importance not only on theoretical grounds but also from a practical point of view. The rational breeding of plants is possible only if the flower-biology of the plants in question (i.e. the question of the possibility of self-pollination, self-sterility, etc.) is accurately known. And it is also essential for plant-breeders that they should have "the power of fixing each fleeting variety of colour, if they will fertilise the flowers of the desired kind with their own pollen for half-a-dozen generations, and grow the seedlings under the same conditions." ("Cross and Self fertilisation" (1st edition), page 460.)

But the influence of Darwin on floral biology was not confined to the development of this branch of Botany. Darwin's activity in this domain has brought about (as Asa Gray correctly pointed out) the revival of teleology in Botany and Zoology. Attempts were now made to determine, not only in the case of flowers but also in vegetative organs, in what relation the form and function of organs stand to one another and to what extent their morphological characters exhibit adaptation to environment. A branch of Botany, which has since been called Ecology (not a very happy term) has been stimulated to vigorous growth by floral biology.

While the influence of the work on the biology of flowers was extraordinarily great, it could not fail to elicit opinions at variance with Darwin's conclusions. The opposition was based partly on reasons valueless as counterarguments, partly on problems which have still to be solved; to some extent also on that tendency against teleological conceptions which has recently become current. This opposing trend of thought is due to the fact that many biologists are content with teleological explanations, unsupported by proof; it is also closely connected with the fact that many authors estimate the importance of natural selection less highly than Darwin did. We may describe the objections which are based on the widespread occurrence of self-fertilisation and geitonogamy as of little importance. Darwin did not deny the occurrence of self-fertilisation, even for a long series of generations; his law states only that "Nature abhors PERPETUAL self-fertilisation." (It is impossible (as has been attempted) to express Darwin's point of view in a single sentence, such as H. Muller's statement of the "Knight-Darwin law." The conditions of life in organisms are so various and complex that laws, such as are formulated in physics and chemistry, can hardly be conceived.) An exception to this rule would therefore occur only in the case of plants in which the possibility of cross-pollination is excluded. Some of the plants with cleistogamous flowers might afford examples of such cases. We have already seen, however, that such a case has not as yet been shown to occur. Burck believed that he had found an instance in certain tropical plants (Anonaceae, Myrmecodia) of the complete exclusion of cross-fertilisation. The flowers of these plants, in which, however,--in contrast to the cleistogamous flowers--the corolla is well developed, remain closed and fruit is produced.

Loew (E. Loew, "Bemerkungen zu Burck... ", "Biolog. Centralbl." XXVI. (1906).) has shown that cases occur in which cross-fertilisation may be effected even in these "cleistopetalous" flowers: humming birds visit the permanently closed flowers of certain species of Nidularium and transport the pollen. The fact that the formation of hybrids may occur as the result of this shows that pollination may be accomplished.

The existence of plants for which self-pollination is of greater importance than it is for others is by no means contradictory to Darwin's view. Self-fertilisation is, for example, of greater importance for annuals than for perennials as without it seeds might fail to be produced. Even in the case of annual plants with small inconspicuous flowers in which self-fertilisation usually occurs, such as Senecio vulgaris, Capsella bursa-pastoris and Stellaria media, A. Bateson (Anna Bateson, "The effects of cross-fertilisation on inconspicuous flowers", "Annals of Botany", Vol. I. 1888, page 255.) found that cross-fertilisation gave a beneficial result, although only in a slight degree. If the favourable effects of sexual reproduction, according to Darwin's view, are correlated with change of environment, it is quite possible that this is of less importance in plants which die after ripening their seeds ("hapaxanthic") and which in any case constantly change their situation. Objections which are based on the proof of the prevalence of self-fertilisation are not, therefore, pertinent. At first sight another point of view, which has been more recently urged, appears to have more weight.

W. Burck (Burck, "Darwin's Kreuzeungsgesetz... ", "Biol. Centralbl". XXVIII. 1908, page 177.) has expressed the opinion that the beneficial results of cross-fertilisation demonstrated by Darwin concern only hybrid plants. These alone become weaker by self-pollination; while pure species derive no advantage from crossing and no disadvantage from self-fertilisation. It is certain that some of the plants used by Darwin were of hybrid origin. (It is questionable if this was always the case.) This is evident from his statements, which are models of clearness and precision; he says that his Ipomoea plants "were probably the offspring of a cross." ("Cross and Self fertilisation" (1st edition), page 55.) The fixed forms of this plant, such as Hero, which was produced by self-fertilisation, and a form of Mimulus with white flowers spotted with red probably resulted from splitting of the hybrids. It is true that the phenomena observed in self-pollination, e.g. in Ipomoea, agree with those which are often noticed in hybrids; Darwin himself drew attention to this.

Let us next call to mind some of the peculiarities connected with hybridisation. We know that hybrids are often characterized by their large size, rapidity of growth, earlier production of flowers, wealth of flower-production and a longer life; hybrids, if crossed with one of the two parent forms, are usually more fertile than when they are crossed together or with another hybrid. But the characters which hybrids exhibit on self-fertilisation are rather variable. The following instance may be quoted from Gartner: "There are many hybrids which retain the self-fertility of the first generation during the second and later generations, but very often in a less degree; a considerable number, however, become sterile." But the hybrid varieties may be more fertile in the second generation than in the first, and in some hybrids the fertility with their own pollen increases in the second, third, and following generations. (K.F. Gartner, "Versuche uber die Bastarderzeugung", Stuttgart, 1849, page 149.) As yet it is impossible to lay down rules of general application for the self-fertility of hybrids. That the beneficial influence of crossing with a fresh stock rests on the same ground--a union of sexual cells possessing somewhat different characters--as the fact that many hybrids are distinguished by greater luxuriance, wealth of flowers, etc. corresponds entirely with Darwin's conclusions. It seems to me to follow clearly from his investigations that there is no essential difference between cross-fertilisation and hybridisation. The heterostyled plants are normally dependent on a process corresponding to hybridisation. The view that specifically distinct species could at best produce sterile hybrids was always opposed by Darwin. But if the good results of crossing were EXCLUSIVELY dependent on the fact that we are concerned with hybrids, there must then be a demonstration of two distinct things. First, that crossing with a fresh stock belonging to the same systematic entity or to the same hybrid, but cultivated for a considerable time under different conditions, shows no superiority over self-fertilisation, and that in pure species crossing gives no better results than self-pollination. If this were the case, we should be better able to understand why in one plant crossing is advantageous while in others, such as Darwin's Hero and the forms of Mimulus and Nicotiana no advantage is gained; these would then be pure species. But such a proof has not been supplied; the inference drawn from cleistogamous and cleistopetalous plants is not supported by evidence, and the experiments on geitonogamy and on the advantage of cross-fertilisation in species which are usually self-fertilised are opposed to this view. There are still but few researches on this point; Darwin found that in Ononis minutissima, which produces cleistogamous as well as self-fertile chasmogamous flowers, the crossed and self-fertilised capsules produced seed in the proportion of 100:65 and that the average bore the proportion 100:86. Facts previously mentioned are also applicable to this case. Further, it is certain that the self-sterility exhibited by many plants has nothing to do with hybridisation. Between self-sterility and reduced fertility as the result of self-fertilisation there is probably no fundamental difference.

It is certain that so difficult a problem as that of the significance of sexual reproduction requires much more investigation. Darwin was anything but dogmatic and always ready to alter an opinion when it was not based on definite proof: he wrote, "But the veil of secrecy is as yet far from lifted; nor will it be, until we can say why it is beneficial that the sexual elements should be differentiated to a certain extent, and why, if the differentiation be carried still further, injury follows." He has also shown us the way along which to follow up this problem; it is that of carefully planned and exact experimental research. It may be that eventually many things will be viewed in a different light, but Darwin's investigations will always form the foundation of Floral Biology on which the future may continue to build.

XXI. MENTAL FACTORS IN EVOLUTION. By C. Lloyd Morgan, LL.D., F.R.S.

In developing his conception of organic evolution Charles Darwin was of necessity brought into contact with some of the problems of mental evolution. In "The Origin of Species" he devoted a chapter to "the diversities of instinct and of the other mental faculties in animals of the same class." ("Origin of Species" (6th edition), page 205.) When he passed to the detailed consideration of "The Descent of Man", it was part of his object to show "that there is no fundamental difference between man and the higher mammals in their mental faculties." ("Descent of Man" (2nd edition 1888), Vol. I. page 99; Popular edition page 99.) "If no organic being excepting man," he said, "had possessed any mental power, or if his powers had been of a wholly different nature from those of the lower animals, then we should never have been able to convince ourselves that our high faculties had been gradually developed." (Ibid. page 99.) In his discussion of "The Expression of the Emotions" it was important for his purpose "fully to recognise that actions readily become associated with other actions and with various states of the mind." ("The Expression of the Emotions" (2nd edition), page 32.) His hypothesis of sexual selection is largely dependent upon the exercise of choice on the part of the female and her preference for "not only the more attractive but at the same time the more vigorous and victorious males." ("Descent of Man", Vol. II. page 435.) Mental processes and physiological processes were for Darwin closely correlated; and he accepted the conclusion "that the nervous system not only regulates most of the existing functions of the body, but has indirectly influenced the progressive development of various bodily structures and of certain mental qualities." (Ibid. pages 437, 438.)

Throughout his treatment, mental evolution was for Darwin incidental to and contributory to organic evolution. For specialised research in comparative and genetic psychology, as an independent field of investigation, he had neither the time nor the requisite training. None the less his writings and the spirit of his work have exercised a profound influence on this department of evolutionary thought. And, for those who follow Darwin's lead, mental evolution is still in a measure subservient to organic evolution. Mental processes are the accompaniments or concomitants of the functional activity of specially differentiated parts of the organism. They are in some way dependent on physiological and physical conditions. But though they are not physical in their nature, and though it is difficult or impossible to conceive that they are physical in their origin, they are, for Darwin and his followers, factors in the evolutionary process in its physical or organic aspect. By the physiologist within his special and well-defined universe of discourse they may be properly regarded as epiphenomena; but by the naturalist in his more catholic survey of nature they cannot be so regarded, and were not so regarded by Darwin. Intelligence has contributed to evolution of which it is in a sense a product.

The facts of observation or of inference which Darwin accepted are these: Conscious experience accompanies some of the modes of animal behaviour; it is concomitant with certain physiological processes; these processes are the outcome of development in the individual and evolution in the race; the accompanying mental processes undergo a like development. Into the subtle philosophical questions which arise out of the naive acceptance of such a creed it was not Darwin's province to enter; "I have nothing to do," he said ("Origin of Species" (6th edition), page 205.), "with the origin of the mental powers, any more than I have with that of life itself." He dealt with the natural history of organisms, including not only their structure but their modes of behaviour; with the natural history of the states of consciousness which accompany some of their actions; and with the relation of behaviour to experience. We will endeavour to follow Darwin in his modesty and candour in making no pretence to give ultimate explanations. But we must note one of the implications of this self-denying ordinance of science. Development and evolution imply continuity. For Darwin and his followers the continuity is organic through physical heredity. Apart from speculative hypothesis, legitimate enough in its proper place but here out of court, we know nothing of continuity of mental evolution as such: consciousness appears afresh in each succeeding generation. Hence it is that for those who follow Darwin's lead, mental evolution is and must ever be, within his universe of discourse, subservient to organic evolution. Only in so far as conscious experience, or its neural correlate, effects some changes in organic structure can it influence the course of heredity; and conversely only in so far as changes in organic structure are transmitted through heredity, is mental evolution rendered possible. Such is the logical outcome of Darwin's teaching.

Those who abide by the cardinal results of this teaching are bound to regard all behaviour as the expression of the functional activities of the living tissues of the organism, and all conscious experience as correlated with such activities. For the purposes of scientific treatment, mental processes are one mode of expression of the same changes of which the physiological processes accompanying behaviour are another mode of expression. This is simply accepted as a fact which others may seek to explain. The behaviour itself is the adaptive application of the energies of the organism; it is called forth by some form of presentation or stimulation brought to bear on the organism by the environment. This presentation is always an individual or personal matter. But in order that the organism may be fitted to respond to the presentation of the environment it must have undergone in some way a suitable preparation. According to the theory of evolution this preparation is primarily racial and is transmitted through heredity. Darwin's main thesis was that the method of preparation is predominantly by natural selection. Subordinate to racial preparation, and always dependent thereon, is individual or personal preparation through some kind of acquisition; of which the guidance of behaviour through individually won experience is a typical example. We here introduce the mental factor because the facts seem to justify the inference. Thus there are some modes of behaviour which are wholly and solely dependent upon inherited racial preparation; there are other modes of behaviour which are also dependent, in part at least, on individual preparation. In the former case the behaviour is adaptive on the first occurrence of the appropriate presentation; in the latter case accommodation to circumstances is only reached after a greater or less amount of acquired organic modification of structure, often accompanied (as we assume) in the higher animals by acquired experience. Logically and biologically the two classes of behaviour are clearly distinguishable: but the analysis of complex cases of behaviour where the two factors cooperate, is difficult and requires careful and critical study of life-history.

The foundations of the mental life are laid in the conscious experience that accompanies those modes of behaviour, dependent entirely on racial preparation, which may broadly be described as instinctive. In the eighth chapter of "The Origin of Species" Darwin says ("Origin of Species" (6th edition), page 205.), "I will not attempt any definition of instinct... Every one understands what is meant, when it is said that instinct impels the cuckoo to migrate and to lay her eggs in other birds' nests. An action, which we ourselves require experience to enable us to perform, when performed by an animal, more especially by a very young one, without experience, and when performed by many individuals in the same way, without their knowing for what purpose it is performed, is usually said to be instinctive." And in the summary at the close of the chapter he says ("Origin of Species" (6th edition), page 233.), "I have endeavoured briefly to show that the mental qualities of our domestic animals vary, and that the variations are inherited. Still more briefly I have attempted to show that instincts vary slightly in a state of nature. No one will dispute that instincts are of the highest importance to each animal. Therefore there is no real difficulty, under changing conditions of life, in natural selection accumulating to any extent slight modifications of instinct which are in any way useful. In many cases habit or use and disuse have probably come into play."

Into the details of Darwin's treatment there is neither space nor need to enter. There are some ambiguous passages; but it may be said that for him, as for his followers to-day, instinctive behaviour is wholly the result of racial preparation transmitted through organic heredity. For the performance of the instinctive act no individual preparation under the guidance of personal experience is necessary. It is true that Darwin quotes with approval Huber's saying that "a little dose of judgment or reason often comes into play, even with animals low in the scale of nature." (Ibid. page 205.) But we may fairly interpret his meaning to be that in behaviour, which is commonly called instinctive, some element of intelligent guidance is often combined. If this be conceded the strictly instinctive performance (or part of the performance) is the outcome of heredity and due to the direct transmission of parental or ancestral aptitudes. Hence the instinctive response as such depends entirely on how the nervous mechanism has been built up through heredity; while intelligent behaviour, or the intelligent factor in behaviour, depends also on how the nervous mechanism has been modified and moulded by use during its development and concurrently with the growth of individual experience in the customary situations of daily life. Of course it is essential to the Darwinian thesis that what Sir E. Ray Lankester has termed "educability," not less than instinct, is hereditary. But it is also essential to the understanding of this thesis that the differentiae of the hereditary factors should be clearly grasped.

For Darwin there were two modes of racial preparation, (1) natural selection, and (2) the establishment of individually acquired habit. He showed that instincts are subject to hereditary variation; he saw that instincts are also subject to modification through acquisition in the course of individual life. He believed that not only the variations but also, to some extent, the modifications are inherited. He therefore held that some instincts (the greater number) are due to natural selection but that others (less numerous) are due, or partly due, to the inheritance of acquired habits. The latter involve Lamarckian inheritance, which of late years has been the centre of so much controversy. It is noteworthy however that Darwin laid especial emphasis on the fact that many of the most typical and also the most complex instincts--those of neuter insects--do not admit of such an interpretation. "I am surprised," he says ("Origin of Species" (6th edition), page 233.), "that no one has hitherto advanced this demonstrative case of neuter insects, against the well-known doctrine of inherited habit, as advanced by Lamarck." None the less Darwin admitted this doctrine as supplementary to that which was more distinctively his own--for example in the case of the instincts of domesticated animals. Still, even in such cases, "it may be doubted," he says (Ibid. pages 210, 211.), "whether any one would have thought of training a dog to point, had not some one dog naturally shown a tendency in this line... so that habit and some degree of selection have probably concurred in civilising by inheritance our dogs." But in the interpretation of the instincts of domesticated animals, a more recently suggested hypothesis, that of organic selection (Independently suggested, on somewhat different lines, by Profs. J. Mark Baldwin, Henry F. Osborn and the writer.), may be helpful. According to this hypothesis any intelligent modification of behaviour which is subject to selection is probably coincident in direction with an inherited tendency to behave in this fashion. Hence in such behaviour there are two factors: (1) an incipient variation in the line of such behaviour, and (2) an acquired modification by which the behaviour is carried further along the same line. Under natural selection those organisms in which the two factors cooperate are likely to survive. Under artificial selection they are deliberately chosen out from among the rest.

Organic selection has been termed a compromise between the more strictly Darwinian and the Lamarckian principles of interpretation. But it is not in any sense a compromise. The principle of interpretation of that which is instinctive and hereditary is wholly Darwinian. It is true that some of the facts of observation relied upon by Lamarckians are introduced. For Lamarckians however the modifications which are admittedly factors in survival, are regarded as the parents of inherited variations; for believers in organic selection they are only the foster parents or nurses. It is because organic selection is the direct outcome of and a natural extension of Darwin's cardinal thesis that some reference to it here is justifiable. The matter may be put with the utmost brevity as follows. (1) Variations (V) occur, some of which are in the direction of increased adaptation (+), others in the direction of decreased adaptation (-). (2) Acquired modifications (M) also occur. Some of these are in the direction of increased accommodation to circumstances (+), while others are in the direction of diminished accommodation (-). Four major combinations are

(a) + V with + M, (b) + V with - M, (c) - V with + M, (d) - V with - M.

Of these (d) must inevitably be eliminated while (a) are selected. The predominant survival of (a) entails the survival of the adaptive variations which are inherited. The contributory acquisitions (+M) are not inherited; but they are none the less factors in determining the survival of the coincident variations. It is surely abundantly clear that this is Darwinism and has no tincture of Lamarck's essential principle, the inheritance of acquired characters.

Whether Darwin himself would have accepted this interpretation of some at least of the evidence put forward by Lamarckians is unfortunately a matter of conjecture. The fact remains that in his interpretation of instinct and in allied questions he accepted the inheritance of individually acquired modifications of behaviour and structure.

Darwin was chiefly concerned with instinct from the biological rather than from the psychological point of view. Indeed it must be confessed that, from the latter standpoint, his conception of instinct as a "mental faculty" which "impels" an animal to the performance of certain actions, scarcely affords a satisfactory basis for genetic treatment. To carry out the spirit of Darwin's teaching it is necessary to link more closely biological and psychological evolution. The first step towards this is to interpret the phenomena of instinctive behaviour in terms of stimulation and response. It may be well to take a particular case. Swimming on the part of a duckling is, from the biological point of view, a typical example of instinctive behaviour. Gently lower a recently hatched bird into water: coordinated movements of the limbs follow in rhythmical sequence. The behaviour is new to the individual though it is no doubt closely related to that of walking, which is no less instinctive. There is a group of stimuli afforded by the "presentation" which results from partial immersion: upon this there follows as a complex response an application of the functional activities in swimming; the sequence of adaptive application on the appropriate presentation is determined by racial preparation. We know, it is true, but little of the physiological details of what takes place in the central nervous system; but in broad outline the nature of the organic mechanism and the manner of its functioning may at least be provisionally conjectured in the present state of physiological knowledge. Similarly in the case of the pecking of newly-hatched chicks; there is a visual presentation, there is probably a cooperating group of stimuli from the alimentary tract in need of food, there is an adaptive application of the activities in a definite mode of behaviour. Like data are afforded in a great number of cases of instinctive procedure, sometimes occurring very early in life, not infrequently deferred until the organism is more fully developed, but all of them dependent upon racial preparation. No doubt there is some range of variation in the behaviour, just such variation as the theory of natural selection demands. But there can be no question that the higher animals inherit a bodily organisation and a nervous system, the functional working of which gives rise to those inherited modes of behaviour which are termed instinctive.

It is to be noted that the term "instinctive" is here employed in the adjectival form as a descriptive heading under which may be grouped many and various modes of behaviour due to racial preparation. We speak of these as inherited; but in strictness what is transmitted through heredity is the complex of anatomical and physiological conditions under which, in appropriate circumstances, the organism so behaves. So far the term "instinctive" has a restricted biological connotation in terms of behaviour. But the connecting link between biological evolution and psychological evolution is to be sought,--as Darwin fully realised,--in the phenomena of instinct, broadly considered. The term "instinctive" has also a psychological connotation. What is that connotation?

Let us take the case of the swimming duckling or the pecking chick, and fix our attention on the first instinctive performance. Grant that just as there is, strictly speaking, no inherited behaviour, but only the conditions which render such behaviour under appropriate circumstances possible; so too there is no inherited experience, but only the conditions which render such experience possible; then the cerebral conditions in both cases are the same. The biological behaviour-complex, including the total stimulation and the total response with the intervening or resultant processes in the sensorium, is accompanied by an experience-complex including the initial stimulation-consciousness and resulting response-consciousness. In the experience-complex are comprised data which in psychological analysis are grouped under the headings of cognition, affective tone and conation. But the complex is probably experienced as an unanalysed whole. If then we use the term "instinctive" so as to comprise all congenital modes of behaviour which contribute to experience, we are in a position to grasp the view that the net result in consciousness constitutes what we may term the primary tissue of experience. To the development of this experience each instinctive act contributes. The nature and manner of organisation of this primary tissue of experience are dependent on inherited biological aptitudes; but they are from the outset onwards subject to secondary development dependent on acquired aptitudes. Biological values are supplemented by psychological values in terms of satisfaction or the reverse.

In our study of instinct we have to select some particular phase of animal behaviour and isolate it so far as is possible from the life of which it is a part. But the animal is a going concern, restlessly active in many ways. Many instinctive performances, as Darwin pointed out ("Origin of Species" (6th edition), page 206.), are serial in their nature. But the whole of active life is a serial and coordinated business. The particular instinctive performance is only an episode in a life-history, and every mode of behaviour is more or less closely correlated with other modes. This coordination of behaviour is accompanied by a correlation of the modes of primary experience. We may classify the instinctive modes of behaviour and their accompanying modes of instinctive experience under as many heads as may be convenient for our purposes of interpretation, and label them instincts of self-preservation, of pugnacity, of acquisition, the reproductive instincts, the parental instincts, and so forth. An instinct, in this sense of the term (for example the parental instinct), may be described as a specialised part of the primary tissue of experience differentiated in relation to some definite biological end. Under such an instinct will fall a large number of particular and often well-defined modes of behaviour, each with its own peculiar mode of experience.

It is no doubt exceedingly difficult as a matter of observation and of inference securely based thereon to distinguish what is primary from what is in part due to secondary acquisition--a fact which Darwin fully appreciated. Animals are educable in different degrees; but where they are educable they begin to profit by experience from the first. Only, therefore, on the occasion of the first instinctive act of a given type can the experience gained be weighed as WHOLLY primary; all subsequent performance is liable to be in some degree, sometimes more, sometimes less, modified by the acquired disposition which the initial behaviour engenders. But the early stages of acquisition are always along the lines predetermined by instinctive differentiation. It is the task of comparative psychology to distinguish the primary tissue of experience from its secondary and acquired modifications. We cannot follow up the matter in further detail. It must here suffice to suggest that this conception of instinct as a primary form of experience lends itself better to natural history treatment than Darwin's conception of an impelling force, and that it is in line with the main trend of Darwin's thought.

In a characteristic work,--characteristic in wealth of detail, in closeness and fidelity of observation, in breadth of outlook, in candour and modesty,--Darwin dealt with "The Expression of the Emotions in Man and Animals". Sir Charles Bell in his "Anatomy of Expression" had contended that many of man's facial muscles had been specially created for the sole purpose of being instrumental in the expression of his emotions. Darwin claimed that a natural explanation, consistent with the doctrine of evolution, could in many cases be given and would in other cases be afforded by an extension of the principles he advocated. "No doubt," he said ("Expression of the Emotions", page 13. The passage is here somewhat condensed.), "as long as man and all other animals are viewed as independent creations, an effectual stop is put to our natural desire to investigate as far as possible the causes of Expression. By this doctrine, anything and everything can be equally well explained... With mankind, some expressions... can hardly be understood, except on the belief that man once existed in a much lower and animal-like condition. The community of certain expressions in distinct though allied species... is rendered somewhat more intelligible, if we believe in their descent from a common progenitor. He who admits on general grounds that the structure and habits of all animals have been gradually evolved, will look at the whole subject of Expression in a new and interesting light."

Darwin relied on three principles of explanation. "The first of these principles is, that movements which are serviceable in gratifying some desire, or in relieving some sensation, if often repeated, become so habitual that they are performed, whether or not of any service, whenever the same desire or sensation is felt, even in a very weak degree." (Ibid. page 368.) The modes of expression which fall under this head have become instinctive through the hereditary transmission of acquired habit. "As far as we can judge, only a few expressive movements are learnt by each individual; that is, were consciously and voluntarily performed during the early years of life for some definite object, or in imitation of others, and then became habitual. The far greater number of the movements of expression, and all the more important ones, are innate or inherited; and such cannot be said to depend on the will of the individual. Nevertheless, all those included under our first principle were at first voluntarily performed for a definite object,--namely, to escape some danger, to relieve some distress, or to gratify some desire." (Ibid. pages 373, 374.)

"Our second principle is that of antithesis. The habit of voluntarily performing opposite movements under opposite impulses has become firmly established in us by the practice of our whole lives. Hence, if certain actions have been regularly performed, in accordance with our first principle, under a certain frame of mind, there will be a strong and involuntary tendency to the performance of directly opposite actions, whether or not these are of any use, under the excitement of an opposite frame of mind." ("Expression of the Emotions", page 368.) This principle of antithesis has not been widely accepted. Nor is Darwin's own position easy to grasp.

"Our third principle," he says (Ibid. page 369.), "is the direct action of the excited nervous system on the body, independently of the will, and independently, in large part, of habit. Experience shows that nerve-force is generated and set free whenever the cerebro-spinal system is excited. The direction which this nerve-force follows is necessarily determined by the lines of connection between the nerve-cells, with each other and with various parts of the body."

Lack of space prevents our following up the details of Darwin's treatment of expression. Whether we accept or do not accept his three principles of explanation we must regard his work as a masterpiece of descriptive analysis, packed full of observations possessing lasting value. For a further development of the subject it is essential that the instinctive factors in expression should be more fully distinguished from those which are individually acquired--a difficult task--and that the instinctive factors should be rediscussed in the light of modern doctrines of heredity, with a view to determining whether Lamarckian inheritance, on which Darwin so largely relied, is necessary for an interpretation of the facts.

The whole subject as Darwin realised is very complex. Even the term "expression" has a certain amount of ambiguity. When the emotion is in full flood the animal fights, flees, or faints. Is this full-tide effect to be regarded as expression; or are we to restrict the term to the premonitory or residual effects--the bared canine when the fighting mood is being roused, the ruffled fur when reminiscent representations of the object inducing anger cross the mind? Broadly considered both should be included. The activity of premonitory expression as a means of communication was recognised by Darwin; he might, perhaps, have emphasised it more strongly in dealing with the lower animals. Man so largely relies on a special means of communication, that of language, that he sometimes fails to realise that for animals with their keen powers of perception, and dependent as they are on such means of communication, the more strictly biological means of expression are full of subtle suggestiveness. Many modes of expression, otherwise useless, are signs of behaviour that may be anticipated,--signs which stimulate the appropriate attitude of response. This would not, however, serve to account for the utility of the organic accompaniments--heart-affection, respiratory changes, vaso-motor effects and so forth, together with heightened muscular tone,--on all of which Darwin lays stress ("Expression of the Emotions", pages 65 ff.) under his third principle. The biological value of all this is, however, of great importance, though Darwin was hardly in a position to take it fully into account.

Having regard to the instinctive and hereditary factors of emotional expression we may ask whether Darwin's third principle does not alone suffice as an explanation. Whether we admit or reject Lamarckian inheritance it would appear that all hereditary expression must be due to pre-established connections within the central nervous system and to a transmitted provision for coordinated response under the appropriate stimulation. If this be so, Darwin's first and second principles are subordinate and ancillary to the third, an expression, so far as it is instinctive or hereditary, being "the direct result of the constitution of the nervous system."

Darwin accepted the emotions themselves as hereditary or acquired states of mind and devoted his attention to their expression. But these emotions themselves are genetic products and as such dependent on organic conditions. It remained, therefore, for psychologists who accepted evolution and sought to build on biological foundations to trace the genesis of these modes of animal and human experience. The subject has been independently developed by Professors Lange and James (Cf. William James, "Principles of Psychology", Vol. II. Chap. XXV, London, 1890.); and some modification of their view is regarded by many evolutionists as affording the best explanation of the facts. We must fix our attention on the lower emotions, such as anger or fear, and on their first occurrence in the life of the individual organism. It is a matter of observation that if a group of young birds which have been hatched in an incubator are frightened by an appropriate presentation, auditory or visual, they instinctively respond in special ways. If we speak of this response as the expression, we find that there are many factors. There are certain visible modes of behaviour, crouching at once, scattering and then crouching, remaining motionless, the braced muscles sustaining an attitude of arrest, and so forth. There are also certain visceral or organic effects, such as affections of the heart and respiration. These can be readily observed by taking the young bird in the hand. Other effects cannot be readily observed; vaso-motor changes, affections of the alimentary canal, the skin and so forth. Now the essence of the James-Lange view, as applied to these congenital effects, is that though we are justified in speaking of them as effects of the stimulation, we are not justified, without further evidence, in speaking of them as effects of the emotional state. May it not rather be that the emotion as a primary mode of experience is the concomitant of the net result of the organic situation--the initial presentation, the instinctive mode of behaviour, the visceral disturbances? According to this interpretation the primary tissue of experience of the emotional order, felt as an unanalysed complex, is generated by the stimulation of the sensorium by afferent or incoming physiological impulses from the special senses, from the organs concerned in the responsive behaviour, from the viscera and vaso-motor system.

Some psychologists, however, contend that the emotional experience is generated in the sensorium prior to, and not subsequent to, the behaviour-response and the visceral disturbances. It is a direct and not an indirect outcome of the presentation to the special senses. Be this as it may, there is a growing tendency to bring into the closest possible relation, or even to identify, instinct and emotion in their primary genesis. The central core of all such interpretations is that instinctive behaviour and experience, its emotional accompaniments, and its expression, are but different aspects of the outcome of the same organic occurrences. Such emotions are, therefore, only a distinguishable aspect of the primary tissue of experience and exhibit a like differentiation. Here again a biological foundation is laid for a psychological doctrine of the mental development of the individual.

The intimate relation between emotion as a psychological mode of experience and expression as a group of organic conditions has an important bearing on biological interpretation. The emotion, as the psychological accompaniment of orderly disturbances in the central nervous system profoundly influences behaviour and often renders it more vigorous and more effective. The utility of the emotions in the struggle for existence can, therefore, scarcely be over-estimated. Just as keenness of perception has survival-value; just as it is obviously subject to variation; just as it must be enhanced under natural selection, whether individually acquired increments are inherited or not; and just as its value lies not only in this or that special perceptive act but in its importance for life as a whole; so the vigorous effectiveness of activity has survival-value; it is subject to variation; it must be enhanced under natural selection; and its importance lies not only in particular modes of behaviour but in its value for life as a whole. If emotion and its expression as a congenital endowment are but different aspects of the same biological occurrence; and if this is a powerful supplement to vigour effectiveness and persistency of behaviour, it must on Darwin's principles be subject to natural selection.

If we include under the expression of the emotions not only the premonitory symptoms of the initial phases of the organic and mental state, not only the signs or conditions of half-tide emotion, but the full-tide manifestation of an emotion which dominates the situation, we are naturally led on to the consideration of many of the phenomena which are discussed under the head of sexual selection. The subject is difficult and complex, and it was treated by Darwin with all the strength he could summon to the task. It can only be dealt with here from a special point of view--that which may serve to illustrate the influence of certain mental factors on the course of evolution. From this point of view too much stress can scarcely be laid on the dominance of emotion during the period of courtship and pairing in the more highly organised animals. It is a period of maximum vigour, maximum activity, and, correlated with special modes of behaviour and special organic and visceral accompaniments, a period also of maximum emotional excitement. The combats of males, their dances and aerial evolutions, their elaborate behaviour and display, or the flood of song in birds, are emotional expressions which are at any rate coincident in time with sexual periodicity. From the combat of the males there follows on Darwin's principles the elimination of those which are deficient in bodily vigour, deficient in special structures, offensive or protective, which contribute to success, deficient in the emotional supplement of which persistent and whole-hearted fighting is the expression, and deficient in alertness and skill which are the outcome of the psychological development of the powers of perception. Few biologists question that we have here a mode of selection of much importance, though its influence on psychological evolution often fails to receive its due emphasis. Mr Wallace ("Darwinism", pages 282, 283, London, 1889.) regards it as "a form of natural selection"; "to it," he says, "we must impute the development of the exceptional strength, size, and activity of the male, together with the possession of special offensive and defensive weapons, and of all other characters which arise from the development of these or are correlated with them." So far there is little disagreement among the followers of Darwin--for Mr Wallace, with fine magnanimity, has always preferred to be ranked as such, notwithstanding his right, on which a smaller man would have constantly insisted, to the claim of independent originator of the doctrine of natural selection. So far with regard to sexual selection Darwin and Mr Wallace are agreed; so far and no farther. For Darwin, says Mr Wallace (Ibid. page 283.), "has extended the principle into a totally different field of action, which has none of that character of constancy and of inevitable result that attaches to natural selection, including male rivalry; for by far the larger portion of the phenomena, which he endeavours to explain by the direct action of sexual selection, can only be so explained on the hypothesis that the immediate agency is female choice or preference. It is to this that he imputes the origin of all secondary sexual characters other than weapons of offence and defence... In this extension of sexual selection to include the action of female choice or preference, and in the attempt to give to that choice such wide-reaching effects, I am unable to follow him more than a very little way."

Into the details of Mr Wallace's criticisms it is impossible to enter here. We cannot discuss either the mode of origin of the variations in structure which have rendered secondary sexual characters possible or the modes of selection other than sexual which have rendered them, within narrow limits, specifically constant. Mendelism and mutation theories may have something to say on the subject when these theories have been more fully correlated with the basal principles of selection. It is noteworthy that Mr Wallace says ("Darwinism", pages 283, 284.): "Besides the acquisition of weapons by the male for the purpose of fighting with other males, there are some other sexual characters which may have been produced by natural selection. Such are the various sounds and odours which are peculiar to the male, and which serve as a call to the female or as an indication of his presence. These are evidently a valuable addition to the means of recognition of the two sexes, and are a further indication that the pairing season has arrived; and the production, intensification, and differentiation of these sounds and odours are clearly within the power of natural selection. The same remark will apply to the peculiar calls of birds, and even to the singing of the males." Why the same remark should not apply to their colours and adornments is not obvious. What is obvious is that "means of recognition" and "indication that the pairing season has arrived" are dependent on the perceptive powers of the female who recognises and for whom the indication has meaning. The hypothesis of female preference, stripped of the aesthetic surplusage which is psychologically both unnecessary and unproven, is really only different in degree from that which Mr Wallace admits in principle when he says that it is probable that the female is pleased or excited by the display.

Let us for our present purpose leave on one side and regard as sub judice the question whether the specific details of secondary sexual characters are the outcome of female choice. For us the question is whether certain psychological accompaniments of the pairing situation have influenced the course of evolution and whether these psychological accompaniments are themselves the outcome of evolution. As a matter of observation, specially differentiated modes of behaviour, often very elaborate, frequently requiring highly developed skill, and apparently highly charged with emotional tone, are the precursors of pairing. They are generally confined to the males, whose fierce combats during the period of sexual activity are part of the emotional manifestation. It is inconceivable that they have no biological meaning; and it is difficult to conceive that they have any other biological end than to evoke in the generally more passive female the pairing impulse. They are based on instinctive foundations ingrained in the nervous constitution through natural (or may we not say sexual?) selection in virtue of their profound utility. They are called into play by a specialised presentation such as the sight or the scent of the female at, or a little in advance of, a critical period of the physiological rhythm. There is no necessity that the male should have any knowledge of the end to which his strenuous activity leads up. In presence of the female there is an elaborate application of all the energies of behaviour, just because ages of racial preparation have made him biologically and emotionally what he is--a functionally sexual male that must dance or sing or go through hereditary movements of display, when the appropriate stimulation comes. Of course after the first successful courtship his future behaviour will be in some degree modified by his previous experience. No doubt during his first courtship he is gaining the primary data of a peculiarly rich experience, instinctive and emotional. But the biological foundations of the behaviour of courtship are laid in the hereditary coordinations. It would seem that in some cases, not indeed in all, but perhaps especially in those cases in which secondary sexual behaviour is most highly evolved,--correlative with the ardour of the male is a certain amount of reluctance in the female. The pairing act on her part only takes place after prolonged stimulation, for affording which the behaviour of male courtship is the requisite presentation. The most vigorous, defiant and mettlesome male is preferred just because he alone affords a contributory stimulation adequate to evoke the pairing impulse with its attendant emotional tone.

It is true that this places female preference or choice on a much lower psychological plane than Darwin in some passages seems to contemplate where, for example, he says that the female appreciates the display of the male and places to her credit a taste for the beautiful. But Darwin himself distinctly states ("Descent of Man" (2nd edition), Vol. II. pages 136, 137; (Popular edition), pages 642, 643.) that "it is not probable that she consciously deliberates; but she is most excited or attracted by the most beautiful, or melodious, or gallant males." The view here put forward, which has been developed by Prof. Groos ("The Play of Animals", page 244, London, 1898.), therefore seems to have Darwin's own sanction. The phenomena are not only biological; there are psychological elements as well. One can hardly suppose that the female is unconscious of the male's presence; the final yielding must surely be accompanied by heightened emotional tone. Whether we call it choice or not is merely a matter of definition of terms. The behaviour is in part determined by supplementary psychological values. Prof. Groos regards the coyness of females as "a most efficient means of preventing the too early and too frequent yielding to the sexual impulse." (Ibid. page 283.) Be that as it may, it is, in any case, if we grant the facts, a means through which male sexual behaviour with all its biological and psychological implications, is raised to a level otherwise perhaps unattainable by natural means, while in the female it affords opportunities for the development in the individual and evolution in the race of what we may follow Darwin in calling appreciation, if we empty this word of the aesthetic implications which have gathered round it in the mental life of man.

Regarded from this standpoint sexual selection, broadly considered, has probably been of great importance. The psychological accompaniments of the pairing situation have profoundly influenced the course of biological evolution and are themselves the outcome of that evolution.

Darwin makes only passing reference to those modes of behaviour in animals which go by the name of play. "Nothing," he says ("Descent of Man", Vol. II. page 60; (Popular edition), page 566.), "is more common than for animals to take pleasure in practising whatever instinct they follow at other times for some real good." This is one of the very numerous cases in which a hint of the master has served to stimulate research in his disciples. It was left to Prof. Groos to develop this subject on evolutionary lines and to elaborate in a masterly manner Darwin's suggestion. "The utility of play," he says ("The Play of Animals", page 76.), "is incalculable. This utility consists in the practice and exercise it affords for some of the more important duties of life,"--that is to say, for the performance of activities which will in adult life be essential to survival. He urges (Ibid. page 75.) that "the play of young animals has its origin in the fact that certain very important instincts appear at a time when the animal does not seriously need them." It is, however, questionable whether any instincts appear at a time when they are not needed. And it is questionable whether the instinctive and emotional attitude of the play-fight, to take one example, can be identified with those which accompany fighting in earnest, though no doubt they are closely related and have some common factors. It is probable that play, as preparatory behaviour, differs in biological detail (as it almost certainly does in emotional attributes) from the earnest of after-life and that it has been evolved through differentiation and integration of the primary tissue of experience, as a preparation through which certain essential modes of skill may be acquired--those animals in which the preparatory play-propensity was not inherited in due force and requisite amount being subsequently eliminated in the struggle for existence. In any case there is little question that Prof. Groos is right in basing the play-propensity on instinctive foundations. ("The Play of Animals" page 24.) None the less, as he contends, the essential biological value of play is that it is a means of training the educable nerve-tissue, of developing that part of the brain which is modified by experience and which thus acquires new characters, of elaborating the secondary tissue of experience on the predetermined lines of instinctive differentiation and thus furthering the psychological activities which are included under the comprehensive term "intelligent."

In "The Descent of Man" Darwin dealt at some length with intelligence and the higher mental faculties. ("Descent of Man" (1st edition), Chapters II, III, V; (2nd edition), Chapters III, IV, V.) His object, he says, is to show that there is no fundamental difference between man and the higher mammals in their mental faculties; that these faculties are variable and the variations tend to be inherited; and that under natural selection beneficial variations of all kinds will have been preserved and injurious ones eliminated.

Darwin was too good an observer and too honest a man to minimise the "enormous difference" between the level of mental attainment of civilised man and that reached by any animal. His contention was that the difference, great as it is, is one of degree and not of kind. He realised that, in the development of the mental faculties of man, new factors in evolution have supervened--factors which play but a subordinate and subsidiary part in animal intelligence. Intercommunication by means of language, approbation and blame, and all that arises out of reflective thought, are but foreshadowed in the mental life of animals. Still he contends that these may be explained on the doctrine of evolution. He urges (Ibid. Vol. I. pages 70, 71; (Popular edition), pages 70, 71.)" that man is variable in body and mind; and that the variations are induced, either directly or indirectly, by the same general causes, and obey the same general laws, as with the lower animals." He correlates mental development with the evolution of the brain. (Ibid. page 81.) "As the various mental faculties gradually developed themselves, the brain would almost certainly become larger. No one, I presume, doubts that the large proportion which the size of man's brain bears to his body, compared to the same proportion in the gorilla or orang, is closely connected with his higher mental powers." "With respect to the lower animals," he says ("Descent of Man" (Popular edition), page 82.), "M.E. Lartet ("Comptes Rendus des Sciences", June 1, 1868.), by comparing the crania of tertiary and recent mammals belonging to the same groups, has come to the remarkable conclusion that the brain is generally larger and the convolutions are more complex in the more recent form."

Sir E. Ray Lankester has sought to express in the simplest terms the implications of the increase in size of the cerebrum. "In what," he asks, "does the advantage of a larger cerebral mass consist?" "Man," he replies "is born with fewer ready-made tricks of the nerve-centres--these performances of an inherited nervous mechanism so often called by the ill-defined term 'instincts'--than are the monkeys or any other animal. Correlated with the absence of inherited ready-made mechanism, man has a greater capacity of developing in the course of his individual growth similar nervous mechanisms (similar to but not identical with those of 'instinct') than any other animal... The power of being educated--'educability' as we may term it--is what man possesses in excess as compared with the apes. I think we are justified in forming the hypothesis that it is this 'educability' which is the correlative of the increased size of the cerebrum." There has been natural selection of the more educable animals, for "the character which we describe as 'educability' can be transmitted, it is a congenital character. But the RESULTS of education can NOT be transmitted. In each generation they have to be acquired afresh, and with increased 'educability' they are more readily acquired and a larger variety of them... The fact is that there is no community between the mechanisms of instinct and the mechanisms of intelligence, and that the latter are later in the history of the evolution of the brain than the former and can only develop in proportion as the former become feeble and defective." ("Nature", Vol. LXI. pages 624, 625 (1900).)

In this statement we have a good example of the further development of views which Darwin foreshadowed but did not thoroughly work out. It states the biological case clearly and tersely. Plasticity of behaviour in special accommodation to special circumstances is of survival value; it depends upon acquired characters; it is correlated with increase in size and complexity of the cerebrum; under natural selection therefore the larger and more complex cerebrum as the organ of plastic behaviour has been the outcome of natural selection. We have thus the biological foundations for a further development of genetic psychology.

There are diversities of opinion, as Darwin showed, with regard to the range of instinct in man and the higher animals as contrasted with lower types. Darwin himself said ("Descent of Man", Vol. I. page 100.) that "Man, perhaps, has somewhat fewer instincts than those possessed by the animals which come next to him in the series." On the other hand, Prof. Wm. James says ("Principles of Psychology," Vol. II. page 289.) that man is probably the animal with most instincts. The true position is that man and the higher animals have fewer complete and self-sufficing instincts than those which stand lower in the scale of mental evolution, but that they have an equally large or perhaps larger mass of instinctive raw material which may furnish the stuff to be elaborated by intelligent processes. There is, perhaps, a greater abundance of the primary tissue of experience to be refashioned and integrated by secondary modification; there is probably the same differentiation in relation to the determining biological ends, but there is at the outset less differentiation of the particular and specific modes of behaviour. The specialised instinctive performances and their concomitant experience-complexes are at the outset more indefinite. Only through acquired connections, correlated with experience, do they become definitely organised.

The full working-out of the delicate and subtle relationship of instinct and educability--that is, of the hereditary and the acquired factors in the mental life--is the task which lies before genetic and comparative psychology. They interact throughout the whole of life, and their interactions are very complex. No one can read the chapters of "The Descent of Man" which Darwin devotes to a consideration of the mental characters of man and animals without noticing, on the one hand, how sedulous he is in his search for hereditary foundations, and, on the other hand, how fully he realises the importance of acquired habits of mind. The fact that educability itself has innate tendencies--is in fact a partially differentiated educability--renders the unravelling of the factors of mental progress all the more difficult.

In his comparison of the mental powers of men and animals it was essential that Darwin should lay stress on points of similarity rather than on points of difference. Seeking to establish a doctrine of evolution, with its basal concept of continuity of process and community of character, he was bound to render clear and to emphasise the contention that the difference in mind between man and the higher animals, great as it is, is one of degree and not of kind. To this end Darwin not only recorded a large number of valuable observations of his own, and collected a considerable body of information from reliable sources, he presented the whole subject in a new light and showed that a natural history of mind might be written and that this method of study offered a wide and rich field for investigation. Of course those who regarded the study of mind only as a branch of metaphysics smiled at the philosophical ineptitude of the mere man of science. But the investigation, on natural history lines, has been prosecuted with a large measure of success. Much indeed still remains to be done; for special training is required, and the workers are still few. Promise for the future is however afforded by the fact that investigation is prosecuted on experimental lines and that something like organised methods of research are taking form. There is now but little reliance on casual observations recorded by those who have not undergone the necessary discipline in these methods. There is also some change of emphasis in formulating conclusions. Now that the general evolutionary thesis is fully and freely accepted by those who carry on such researches, more stress is laid on the differentiation of the stages of evolutionary advance than on the fact of their underlying community of nature. The conceptual intelligence which is especially characteristic of the higher mental procedure of man is more firmly distinguished from the perceptual intelligence which he shares with the lower animals--distinguished now as a higher product of evolution, no longer as differing in origin or different in kind. Some progress has been made, on the one hand in rendering an account of intelligent profiting by experience under the guidance of pleasure and pain in the perceptual field, on lines predetermined by instinctive differentiation for biological ends, and on the other hand in elucidating the method of conceptual thought employed, for example, by the investigator himself in interpreting the perceptual experience of the lower animals.

Thus there is a growing tendency to realise more fully that there are two orders of educability--first an educability of the perceptual intelligence based on the biological foundation of instinct, and secondly an educability of the conceptual intelligence which refashions and rearranges the data afforded by previous inheritance and acquisition. It is in relation to this second and higher order of educability that the cerebrum of man shows so large an increase of mass and a yet larger increase of effective surface through its rich convolutions. It is through educability of this order that the human child is brought intellectually and affectively into touch with the ideal constructions by means of which man has endeavoured, with more or less success, to reach an interpretation of nature, and to guide the course of the further evolution of his race--ideal constructions which form part of man's environment.

It formed no part of Darwin's purpose to consider, save in broad outline, the methods, or to discuss in any fulness of detail the results of the process by which a differentiation of the mental faculties of man from those of the lower animals has been brought about--a differentiation the existence of which he again and again acknowledges. His purpose was rather to show that, notwithstanding this differentiation, there is basal community in kind. This must be remembered in considering his treatment of the biological foundations on which man's systems of ethics are built. He definitely stated that he approached the subject "exclusively from the side of natural history." ("Descent of Man", Vol. I. page 149.) His general conclusion is that the moral sense is fundamentally identical with the social instincts, which have been developed for the good of the community; and he suggests that the concept which thus enables us to interpret the biological ground-plan of morals also enables us to frame a rational ideal of the moral end. "As the social instincts," he says (Ibid. page 185.), "both of man and the lower animals have no doubt been developed by nearly the same steps, it would be advisable, if found practicable, to use the same definition in both cases, and to take as the standard of morality, the general good or welfare of the community, rather than the general happiness." But the kind of community for the good of which the social instincts of animals and primitive men were biologically developed may be different from that which is the product of civilisation, as Darwin no doubt realised. Darwin's contention was that conscience is a social instinct and has been evolved because it is useful to the tribe in the struggle for existence against other tribes. On the other hand, J.S. Mill urged that the moral feelings are not innate but acquired, and Bain held the same view, believing that the moral sense is acquired by each individual during his life-time. Darwin, who notes (Ibid. page 150 (footnote).) their opinion with his usual candour, adds that "on the general theory of evolution this is at least extremely improbable. It is impossible to enter into the question here: much turns on the exact connotation of the terms "conscience" and "moral sense," and on the meaning we attach to the statement that the moral sense is fundamentally identical with the social instincts."

Presumably the majority of those who approach the subjects discussed in the third, fourth and fifth chapters of "The Descent of Man" in the full conviction that mental phenomena, not less than organic phenomena, have a natural genesis, would, without hesitation, admit that the intellectual and moral systems of civilised man are ideal constructions, the products of conceptual thought, and that as such they are, in their developed form, acquired. The moral sentiments are the emotional analogues of highly developed concepts. This does not however imply that they are outside the range of natural history treatment. Even though it may be desirable to differentiate the moral conduct of men from the social behaviour of animals (to which some such term as "pre-moral" or "quasi-moral" may be applied), still the fact remains that, as Darwin showed, there is abundant evidence of the occurrence of such social behaviour--social behaviour which, even granted that it is in large part intelligently acquired, and is itself so far a product of educability, is of survival value. It makes for that integration without which no social group could hold together and escape elimination. Furthermore, even if we grant that such behaviour is intelligently acquired, that is to say arises through the modification of hereditary instincts and emotions, the fact remains that only through these instinctive and emotional data is afforded the primary tissue of the experience which is susceptible of such modification.

Darwin sought to show, and succeeded in showing, that for the intellectual and moral life there are instinctive foundations which a biological treatment alone can disclose. It is true that he did not in all cases analytically distinguish the foundations from the superstructure. Even to-day we are scarcely in a position to do so adequately. But his treatment was of great value in giving an impetus to further research. This value indeed can scarcely be overestimated. And when the natural history of the mental operations shall have been written, the cardinal fact will stand forth, that the instinctive and emotional foundations are the outcome of biological evolution and have been ingrained in the race through natural selection. We shall more clearly realise that educability itself is a product of natural selection, though the specific results acquired through cerebral modifications are not transmitted through heredity. It will, perhaps, also be realised that the instinctive foundations of social behaviour are, for us, somewhat out of date and have undergone but little change throughout the progress of civilisation, because natural selection has long since ceased to be the dominant factor in human progress. The history of human progress has been mainly the history of man's higher educability, the products of which he has projected on to his environment. This educability remains on the average what it was a dozen generations ago; but the thought-woven tapestry of his surroundings is refashioned and improved by each succeeding generation. Few men have in greater measure enriched the thought-environment with which it is the aim of education to bring educable human beings into vital contact, than has Charles Darwin. His special field of work was the wide province of biology; but he did much to help us realise that mental factors have contributed to organic evolution and that in man, the highest product of Evolution, they have reached a position of unquestioned supremacy.

XXII. THE INFLUENCE OF THE CONCEPTION OF EVOLUTION ON MODERN PHILOSOPHY. By H. Hoffding.

Professor of Philosophy in the University of Copenhagen.

It is difficult to draw a sharp line between philosophy and natural science. The naturalist who introduces a new principle, or demonstrates a fact which throws a new light on existence, not only renders an important service to philosophy but is himself a philosopher in the broader sense of the word. The aim of philosophy in the stricter sense is to attain points of view from which the fundamental phenomena and the principles of the special sciences can be seen in their relative importance and connection. But philosophy in this stricter sense has always been influenced by philosophy in the broader sense. Greek philosophy came under the influence of logic and mathematics, modern philosophy under the influence of natural science. The name of Charles Darwin stands with those of Galileo, Newton, and Robert Mayer--names which denote new problems and great alterations in our conception of the universe.

First of all we must lay stress on Darwin's own personality. His deep love of truth, his indefatigable inquiry, his wide horizon, and his steady self-criticism make him a scientific model, even if his results and theories should eventually come to possess mainly an historical interest. In the intellectual domain the primary object is to reach high summits from which wide surveys are possible, to reach them toiling honestly upwards by way of experience, and then not to turn dizzy when a summit is gained. Darwinians have sometimes turned dizzy, but Darwin never. He saw from the first the great importance of his hypothesis, not only because of its solution of the old problem as to the value of the concept of species, not only because of the grand picture of natural evolution which it unrolls, but also because of the life and inspiration its method would impart to the study of comparative anatomy, of instinct and of heredity, and finally because of the influence it would exert on the whole conception of existence. He wrote in his note-book in the year 1837: "My theory would give zest to recent and fossil comparative anatomy; it would lead to the study of instinct, heredity, and mind-heredity, whole (of) metaphysics." ("Life and Letters of Charles Darwin", Vol. I. page 8.)

We can distinguish four main points in which Darwin's investigations possess philosophical importance.

The evolution hypothesis is much older than Darwin; it is, indeed, one of the oldest guessings of human thought. In the eighteenth century it was put forward by Diderot and Lamettrie and suggested by Kant (1786). As we shall see later, it was held also by several philosophers in the first half of the nineteenth century. In his preface to "The Origin of Species", Darwin mentions the naturalists who were his forerunners. But he has set forth the hypothesis of evolution in so energetic and thorough a manner that it perforce attracts the attention of all thoughtful men in a much higher degree than it did before the publication of the "Origin".

And further, the importance of his teaching rests on the fact that he, much more than his predecessors, even than Lamarck, sought a foundation for his hypothesis in definite facts. Modern science began by demanding--with Kepler and Newton--evidence of verae causae; this demand Darwin industriously set himself to satisfy--hence the wealth of material which he collected by his observations and his experiments. He not only revived an old hypothesis, but he saw the necessity of verifying it by facts. Whether the special cause on which he founded the explanation of the origin of species--Natural Selection--is sufficient, is now a subject of discussion. He himself had some doubt in regard to this question, and the criticisms which are directed against his hypothesis hit Darwinism rather than Darwin. In his indefatigable search for empirical evidence he is a model even for his antagonists: he has compelled them to approach the problems of life along other lines than those which were formerly followed.

Whether the special cause to which Darwin appealed is sufficient or not, at least to it is probably due the greater part of the influence which he has exerted on the general trend of thought. "Struggle for existence" and "natural selection" are principles which have been applied, more or less, in every department of thought. Recent research, it is true, has discovered greater empirical discontinuity--leaps, "mutations"--whereas Darwin believed in the importance of small variations slowly accumulated. It has also been shown by the experimental method, which in recent biological work has succeeded Darwin's more historical method, that types once constituted possess great permanence, the fluctuations being restricted within clearly defined boundaries. The problem has become more precise, both as to variation and as to heredity. The inner conditions of life have in both respects shown a greater independence than Darwin had supposed in his theory, though he always admitted that the cause of variation was to him a great enigma, "a most perplexing problem," and that the struggle for life could only occur where variation existed. But, at any rate, it was of the greatest importance that Darwin gave a living impression of the struggle for life which is everywhere going on, and to which even the highest forms of existence must be amenable. The philosophical importance of these ideas does not stand or fall with the answer to the question, whether natural selection is a sufficient explanation of the origin of species or not: it has an independent, positive value for everyone who will observe life and reality with an unbiassed mind.

In accentuating the struggle for life Darwin stands as a characteristically English thinker: he continues a train of ideas which Hobbes and Malthus had already begun. Moreover in his critical views as to the conception of species he had English forerunners; in the middle ages Occam and Duns Scotus, in the eighteenth century Berkeley and Hume. In his moral philosophy, as we shall see later, he is an adherent of the school which is represented by Hutcheson, Hume and Adam Smith. Because he is no philosopher in the stricter sense of the term, it is of great interest to see that his attitude of mind is that of the great thinkers of his nation.

In considering Darwin's influence on philosophy we will begin with an examination of the attitude of philosophy to the conception of evolution at the time when "The Origin of Species" appeared. We will then examine the effects which the theory of evolution, and especially the idea of the struggle for life, has had, and naturally must have, on the discussion of philosophical problems.

II.

When "The Origin of Species" appeared fifty years ago Romantic speculation, Schelling's and Hegel's philosophy, still reigned on the continent, while in England Positivism, the philosophy of Comte and Stuart Mill, represented the most important trend of thought. German speculation had much to say on evolution, it even pretended to be a philosophy of evolution. But then the word "evolution" was to be taken in an ideal, not in a real, sense. To speculative thought the forms and types of nature formed a system of ideas, within which any form could lead us by continuous transitions to any other. It was a classificatory system which was regarded as a divine world of thought or images, within which metamorphoses could go on--a condition comparable with that in the mind of the poet when one image follows another with imperceptible changes. Goethe's ideas of evolution, as expressed in his "Metamorphosen der Pflanzen und der Thiere", belong to this category; it is, therefore, incorrect to call him a forerunner of Darwin. Schelling and Hegel held the same idea; Hegel expressly rejected the conception of a real evolution in time as coarse and materialistic. "Nature," he says, "is to be considered as a SYSTEM OF STAGES, the one necessarily arising from the other, and being the nearest truth of that from which it proceeds; but not in such a way that the one is NATURALLY generated by the other; on the contrary (their connection lies) in the inner idea which is the ground of nature. The METAMORPHOSIS can be ascribed only to the notion as such, because it alone is evolution... It has been a clumsy idea in the older as well as in the newer philosophy of nature, to regard the transformation and the transition from one natural form and sphere to a higher as an outward and actual production." ("Encyclopaedie der philosophischen Wissenschaften" (4th edition), Berlin, 1845, paragraph 249.)

The only one of the philosophers of Romanticism who believed in a real, historical evolution, a real production of new species, was Oken. ("Lehrbuch der Naturphilosophie", Jena, 1809.) Danish philosophers, such as Treschow (1812) and Sibbern (1846), have also broached the idea of an historical evolution of all living beings from the lowest to the highest. Schopenhauer's philosophy has a more realistic character than that of Schelling's and Hegel's, his diametrical opposites, though he also belongs to the romantic school of thought. His philosophical and psychological views were greatly influenced by French naturalists and philosophers, especially by Cabanis and Lamarck. He praises the "ever memorable Lamarck," because he laid so much stress on the "will to live." But he repudiates as a "wonderful error" the idea that the organs of animals should have reached their present perfection through a development in time, during the course of innumerable generations. It was, he said, a consequence of the low standard of contemporary French philosophy, that Lamarck came to the idea of the construction of living beings in time through succession! ("Ueber den Willen in der Natur" (2nd edition), Frankfurt a. M., 1854, pages 41-43.)

The positivistic stream of thought was not more in favour of a real evolution than was the Romantic school. Its aim was to adhere to positive facts: it looked with suspicion on far-reaching speculation. Comte laid great stress on the discontinuity found between the different kingdoms of nature, as well as within each single kingdom. As he regarded as unscientific every attempt to reduce the number of physical forces, so he rejected entirely the hypothesis of Lamarck concerning the evolution of species; the idea of species would in his eyes absolutely lose its importance if a transition from species to species under the influence of conditions of life were admitted. His disciples (Littre, Robin) continued to direct against Darwin the polemics which their master had employed against Lamarck. Stuart Mill, who, in the theory of knowledge, represented the empirical or positivistic movement in philosophy--like his English forerunners from Locke to Hume--founded his theory of knowledge and morals on the experience of the single individual. He sympathised with the theory of the original likeness of all individuals and derived their differences, on which he practically and theoretically laid much stress, from the influence both of experience and education, and, generally, of physical and social causes. He admitted an individual evolution, and, in the human species, an evolution based on social progress; but no physiological evolution of species. He was afraid that the hypothesis of heredity would carry us back to the old theory of "innate" ideas.

Darwin was more empirical than Comte and Mill; experience disclosed to him a deeper continuity than they could find; closer than before the nature and fate of the single individual were shown to be interwoven in the great web binding the life of the species with nature as a whole. And the continuity which so many idealistic philosophers could find only in the world of thought, he showed to be present in the world of reality.

III.

Darwin's energetic renewal of the old idea of evolution had its chief importance in strengthening the conviction of this real continuity in the world, of continuity in the series of form and events. It was a great support for all those who were prepared to base their conception of life on scientific grounds. Together with the recently discovered law of the conservation of energy, it helped to produce the great realistic movement which characterises the last third of the nineteenth century. After the decline of the Romantic movement people wished to have firmer ground under their feet and reality now asserted itself in a more emphatic manner than in the period of Romanticism. It was easy for Hegel to proclaim that "the real" was "the rational," and that "the rational" was "the real": reality itself existed for him only in the interpretation of ideal reason, and if there was anything which could not be merged in the higher unity of thought, then it was only an example of the "impotence of nature to hold to the idea." But now concepts are to be founded on nature and not on any system of categories too confidently deduced a priori. The new devotion to nature had its recompense in itself, because the new points of view made us see that nature could indeed "hold to ideas," though perhaps not to those which we had cogitated beforehand.

A most important question for philosophers to answer was whether the new views were compatible with an idealistic conception of life and existence. Some proclaimed that we have now no need of any philosophy beyond the principles of the conservation of matter and energy and the principle of natural evolution: existence should and could be definitely and completely explained by the laws of material nature. But abler thinkers saw that the thing was not so simple. They were prepared to give the new views their just place and to examine what alterations the old views must undergo in order to be brought into harmony with the new data.

The realistic character of Darwin's theory was shown not only in the idea of natural continuity, but also, and not least, in the idea of the cause whereby organic life advances step by step. This idea--the idea of the struggle for life--implied that nothing could persist, if it had no power to maintain itself under the given conditions. Inner value alone does not decide. Idealism was here put to its hardest trial. In continuous evolution it could perhaps still find an analogy to the inner evolution of ideas in the mind; but in the demand for power in order to struggle with outward conditions Realism seemed to announce itself in its most brutal form. Every form of Idealism had to ask itself seriously how it was going to "struggle for life" with this new Realism.

We will now give a short account of the position which leading thinkers in different countries have taken up in regard to this question.

I. Herbert Spencer was the philosopher whose mind was best prepared by his own previous thinking to admit the theory of Darwin to a place in his conception of the world. His criticism of the arguments which had been put forward against the hypothesis of Lamarck, showed that Spencer, as a young man, was an adherent to the evolution idea. In his "Social Statics" (1850) he applied this idea to human life and moral civilisation. In 1852 he wrote an essay on "The Development Hypothesis", in which he definitely stated his belief that the differentiation of species, like the differentiation within a single organism, was the result of development. In the first edition of his "Psychology" (1855) he took a step which put him in opposition to the older English school (from Locke to Mill): he acknowledged "innate ideas" so far as to admit the tendency of acquired habits to be inherited in the course of generations, so that the nature and functions of the individual are only to be understood through its connection with the life of the species. In 1857, in his essay on "Progress", he propounded the law of differentiation as a general law of evolution, verified by examples from all regions of experience, the evolution of species being only one of these examples. On the effect which the appearance of "The Origin of Species" had on his mind he writes in his "Autobiography": "Up to that time... I held that the sole cause of organic evolution is the inheritance of functionally-produced modifications. The "Origin of Species" made it clear to me that I was wrong, and that the larger part of the facts cannot be due to any such cause... To have the theory of organic evolution justified was of course to get further support for that theory of evolution at large with which... all my conceptions were bound up." (Spencer, "Autobiography", Vol. II. page 50, London, 1904.) Instead of the metaphorical expression "natural selection," Spencer introduced the term "survival of the fittest," which found favour with Darwin as well as with Wallace.

In working out his ideas of evolution, Spencer found that differentiation was not the only form of evolution. In its simplest form evolution is mainly a concentration, previously scattered elements being integrated and losing independent movement. Differentiation is only forthcoming when minor wholes arise within a greater whole. And the highest form of evolution is reached when there is a harmony between concentration and differentiation, a harmony which Spencer calls equilibration and which he defines as a moving equilibrium. At the same time this definition enables him to illustrate the expression "survival of the fittest." "Every living organism exhibits such a moving equilibrium--a balanced set of functions constituting its life; and the overthrow of this balanced set of functions or moving equilibrium is what we call death. Some individuals in a species are so constituted that their moving equilibria are less easily overthrown than those of other individuals; and these are the fittest which survive, or, in Mr Darwin's language, they are the select which nature preserves." (Ibid. page 100.) Not only in the domain of organic life, but in all domains, the summit of evolution is, according to Spencer, characterised by such a harmony--by a moving equilibrium.

Spencer's analysis of the concept of evolution, based on a great variety of examples, has made this concept clearer and more definite than before. It contains the three elements; integration, differentiation and equilibration. It is true that a concept which is to be valid for all domains of experience must have an abstract character, and between the several domains there is, strictly speaking, only a relation of analogy. So there is only analogy between psychical and physical evolution. But this is no serious objection, because general concepts do not express more than analogies between the phenomena which they represent. Spencer takes his leading terms from the material world in defining evolution (in the simplest form) as integration of matter and dissipation of movement; but as he--not always quite consistently (Cf. my letter to him, 1876, now printed in Duncan's "Life and Letters of Herbert Spencer", page 178, London, 1908.)--assumed a correspondence of mind and matter, he could very well give these terms an indirect importance for psychical evolution. Spencer has always, in my opinion with full right, repudiated the ascription of materialism. He is no more a materialist than Spinoza. In his "Principles of Psychology" (paragraph 63) he expressed himself very clearly: "Though it seems easier to translate so-called matter into so-called spirit, than to translate so-called spirit into so-called matter--which latter is indeed wholly impossible--yet no translation can carry us beyond our symbols." These words lead us naturally to a group of thinkers whose starting-point was psychical evolution. But we have still one aspect of Spencer's philosophy to mention.

Spencer founded his "laws of evolution" on an inductive basis, but he was convinced that they could be deduced from the law of the conservation of energy. Such a deduction is, perhaps, possible for the more elementary forms of evolution, integration and differentiation; but it is not possible for the highest form, the equilibration, which is a harmony of integration and differentiation. Spencer can no more deduce the necessity for the eventual appearance of "moving equilibria" of harmonious totalities than Hegel could guarantee the "higher unities" in which all contradictions should be reconciled. In Spencer's hands the theory of evolution acquired a more decidedly optimistic character than in Darwin's; but I shall deal later with the relation of Darwin's hypothesis to the opposition of optimism and pessimism.

II. While the starting-point of Spencer was biological or cosmological, psychical evolution being conceived as in analogy with physical, a group of eminent thinkers--in Germany Wundt, in France Fouillee, in Italy Ardigo--took, each in his own manner, their starting-point in psychical evolution as an original fact and as a type of all evolution, the hypothesis of Darwin coming in as a corroboration and as a special example. They maintain the continuity of evolution; they find this character most prominent in psychical evolution, and this is for them a motive to demand a corresponding continuity in the material, especially in the organic domain.

To Wundt and Fouillee the concept of will is prominent. They see the type of all evolution in the transformation of the life of will from blind impulse to conscious choice; the theories of Lamarck and Darwin are used to support the view that there is in nature a tendency to evolution in steady reciprocity with external conditions. The struggle for life is here only a secondary fact. Its apparent prominence is explained by the circumstance that the influence of external conditions is easily made out, while inner conditions can be verified only through their effects. For Ardigo the evolution of thought was the starting-point and the type: in the evolution of a scientific hypothesis we see a progress from the indefinite (indistinto) to the definite (distinto), and this is a characteristic of all evolution, as Ardigo has pointed out in a series of works. The opposition between indistinto and distinto corresponds to Spencer's opposition between homogeneity and heterogeneity. The hypothesis of the origin of differences of species from more simple forms is a special example of the general law of evolution.

In the views of Wundt and Fouillee we find the fundamental idea of idealism: psychical phenomena as expressions of the innermost nature of existence. They differ from the older Idealism in the great stress which they lay on evolution as a real, historical process which is going on through steady conflict with external conditions. The Romantic dread of reality is broken. It is beyond doubt that Darwin's emphasis on the struggle for life as a necessary condition of evolution has been a very important factor in carrying philosophy back to reality from the heaven of pure ideas. The philosophy of Ardigo, on the other side, appears more as a continuation and deepening of positivism, though the Italian thinker arrived at his point of view independently of French-English positivism. The idea of continuous evolution is here maintained in opposition to Comte's and Mill's philosophy of discontinuity. From Wundt and Fouillee Ardigo differs in conceiving psychical evolution not as an immediate revelation of the innermost nature of existence, but only as a single, though the most accessible example, of evolution.

III. To the French philosophers Boutroux and Bergson, evolution proper is continuous and qualitative, while outer experience and physical science give us fragments only, sporadic processes and mechanical combinations. To Bergson, in his recent work "L'Evolution Creatrice", evolution consists in an elan de vie which to our fragmentary observation and analytic reflexion appears as broken into a manifold of elements and processes. The concept of matter in its scientific form is the result of this breaking asunder, essential for all scientific reflexion. In these conceptions the strongest opposition between inner and outer conditions of evolution is expressed: in the domain of internal conditions spontaneous development of qualitative forms--in the domain of external conditions discontinuity and mechanical combination.

We see, then, that the theory of evolution has influenced philosophy in a variety of forms. It has made idealistic thinkers revise their relation to the real world; it has led positivistic thinkers to find a closer connection between the facts on which they based their views; it has made us all open our eyes for new possibilities to arise through the prima facie inexplicable "spontaneous" variations which are the condition of all evolution. This last point is one of peculiar interest. Deeper than speculative philosophy and mechanical science saw in the days of their triumph, we catch sight of new streams, whose sources and laws we have still to discover. Most sharply does this appear in the theory of mutation, which is only a stronger accentuation of a main point in Darwinism. It is interesting to see that an analogous problem comes into the foreground in physics through the discovery of radioactive phenomena, and in psychology through the assumption of psychical new formations (as held by Boutroux, William James and Bergson). From this side, Darwin's ideas, as well as the analogous ideas in other domains, incite us to renewed examination of our first principles, their rationality and their value. On the other hand, his theory of the struggle for existence challenges us to examine the conditions and discuss the outlook as to the persistence of human life and society and of the values that belong to them. It is not enough to hope (or fear?) the rising of new forms; we have also to investigate the possibility of upholding the forms and ideals which have hitherto been the bases of human life. Darwin has here given his age the most earnest and most impressive lesson. This side of Darwin's theory is of peculiar interest to some special philosophical problems to which I now pass.

IV.

Among philosophical problems the problem of knowledge has in the last century occupied a foremost place. It is natural, then, to ask how Darwin and the hypothesis whose most eminent representative he is, stand to this problem.

Darwin started an hypothesis. But every hypothesis is won by inference from certain presuppositions, and every inference is based on the general principles of human thought. The evolution hypothesis presupposes, then, human thought and its principles. And not only the abstract logical principles are thus presupposed. The evolution hypothesis purports to be not only a formal arrangement of phenomena, but to express also the law of a real process. It supposes, then, that the real data--all that in our knowledge which we do not produce ourselves, but which we in the main simply receive--are subjected to laws which are at least analogous to the logical relations of our thoughts; in other words, it assumes the validity of the principle of causality. If organic species could arise without cause there would be no use in framing hypotheses. Only if we assume the principle of causality, is there a problem to solve.

Though Darwinism has had a great influence on philosophy considered as a striving after a scientific view of the world, yet here is a point of view--the epistemological--where philosophy is not only independent but reaches beyond any result of natural science. Perhaps it will be said: the powers and functions of organic beings only persist (perhaps also only arise) when they correspond sufficiently to the conditions under which the struggle of life is to go on. Human thought itself is, then, a variation (or a mutation) which has been able to persist and to survive. Is not, then, the problem of knowledge solved by the evolution hypothesis? Spencer had given an affirmative answer to this question before the appearance of "The Origin of Species". For the individual, he said, there is an a priori, original, basis (or Anlage) for all mental life; but in the species all powers have developed in reciprocity with external conditions. Knowledge is here considered from the practical point of view, as a weapon in the struggle for life, as an "organon" which has been continuously in use for generations. In recent years the economic or pragmatic epistemology, as developed by Avenarius and Mach in Germany, and by James in America, points in the same direction. Science, it is said, only maintains those principles and presuppositions which are necessary to the simplest and clearest orientation in the world of experience. All assumptions which cannot be applied to experience and to practical work, will successively be eliminated.

In these views a striking and important application is made of the idea of struggle for life to the development of human thought. Thought must, as all other things in the world, struggle for life. But this whole consideration belongs to psychology, not to the theory of knowledge (epistemology), which is concerned only with the validity of knowledge, not with its historical origin. Every hypothesis to explain the origin of knowledge must submit to cross-examination by the theory of knowledge, because it works with the fundamental forms and principles of human thought. We cannot go further back than these forms and principles, which it is the aim of epistemology to ascertain and for which no further reason can be given. (The present writer, many years ago, in his "Psychology" (Copenhagen, 1882; English translation London, 1891), criticised the evolutionistic treatment of the problem of knowledge from the Kantian point of view.)

But there is another side of the problem which is, perhaps, of more importance and which epistemology generally overlooks. If new variations can arise, not only in organic but perhaps also in inorganic nature, new tasks are placed before the human mind. The question is, then, if it has forms in which there is room for the new matter? We are here touching a possibility which the great master of epistemology did not bring to light. Kant supposed confidently that no other matter of knowledge could stream forth from the dark source which he called "the thing-in-itself," than such as could be synthesised in our existing forms of knowledge. He mentions the possibility of other forms than the human, and warns us against the dogmatic assumption that the human conception of existence should be absolutely adequate. But he seems to be quite sure that the thing-in-itself works constantly, and consequently always gives us only what our powers can master. This assumption was a consequence of Kant's rationalistic tendency, but one for which no warrant can be given. Evolutionism and systematism are opposing tendencies which can never be absolutely harmonised one with the other. Evolution may at any time break some form which the system-monger regards as finally established. Darwin himself felt a great difference in looking at variation as an evolutionist and as a systematist. When he was working at his evolution theory, he was very glad to find variations; but they were a hindrance to him when he worked as a systematist, in preparing his work on Cirripedia. He says in a letter: "I had thought the same parts of the same species more resemble (than they do anyhow in Cirripedia) objects cast in the same mould. Systematic work would be easy were it not for this confounded variation, which, however, is pleasant to me as a speculatist, though odious to me as a systematist." ("Life and Letters", Vol. II. page 37.) He could indeed be angry with variations even as an evolutionist; but then only because he could not explain them, not because he could not classify them. "If, as I must think, external conditions produce little DIRECT effect, what the devil determines each particular variation?" (Ibid. page 232.) What Darwin experienced in his particular domain holds good of all knowledge. All knowledge is systematic, in so far as it strives to put phenomena in quite definite relations, one to another. But the systematisation can never be complete. And here Darwin has contributed much to widen the world for us. He has shown us forces and tendencies in nature which make absolute systems impossible, at the same time that they give us new objects and problems. There is still a place for what Lessing called "the unceasing striving after truth," while "absolute truth" (in the sense of a closed system) is unattainable so long as life and experience are going on.

There is here a special remark to be made. As we have seen above, recent research has shown that natural selection or struggle for life is no explanation of variations. Hugo de Vries distinguishes between partial and embryonal variations, or between variations and mutations, only the last-named being heritable, and therefore of importance for the origin of new species. But the existence of variations is not only of interest for the problem of the origin of species; it has also a more general interest. An individual does not lose its importance for knowledge, because its qualities are not heritable. On the contrary, in higher beings at least, individual peculiarities will become more and more independent objects of interest. Knowledge takes account of the biographies not only of species, but also of individuals: it seeks to find the law of development of the single individual. (The new science of Ecology occupies an intermediate position between the biography of species and the biography of individuals. Compare "Congress of Arts and Science", St Louis, Vol. V. 1906 (the Reports of Drude and Robinson) and the work of my colleague E. Warming.) As Leibniz said long ago, individuality consists in the law of the changes of a being. "La loi du changement fait l'individualite de chaque substance." Here is a world which is almost new for science, which till now has mainly occupied itself with general laws and forms. But these are ultimately only means to understand the individual phenomena, in whose nature and history a manifold of laws and forms always cooperate. The importance of this remark will appear in the sequel.

To many people the Darwinian theory of natural selection or struggle for existence seemed to change the whole conception of life, and particularly all the conditions on which the validity of ethical ideas depends. If only that has persistence which can be adapted to a given condition, what will then be the fate of our ideals, of our standards of good and evil? Blind force seems to reign, and the only thing that counts seems to be the most heedless use of power. Darwinism, it was said, has proclaimed brutality. No other difference seems permanent save that between the sound, powerful and happy on the one side, the sick, feeble and unhappy on the other; and every attempt to alleviate this difference seems to lead to general enervation. Some of those who interpreted Darwinism in this manner felt an aesthetic delight in contemplating the heedlessness and energy of the great struggle for existence and anticipated the realisation of a higher human type as the outcome of it: so Nietzsche and his followers. Others recognising the same consequences in Darwinism regarded these as one of the strongest objections against it; so Duhring and Kropotkin (in his earlier works).

This interpretation of Darwinism was frequent in the interval between the two main works of Darwin--"The Origin of Species" and "The Descent of Man". But even during this interval it was evident to an attentive reader that Darwin himself did not found his standard of good and evil on the features of the life of nature he had emphasised so strongly. He did not justify the ways along which nature reached its ends; he only pointed them out. The "real" was not to him, as to Hegel, one with the "rational." Darwin has, indeed, by his whole conception of nature, rendered a great service to ethics in making the difference between the life of nature and the ethical life appear in so strong a light. The ethical problem could now be stated in a sharper form than before. But this was not the first time that the idea of the struggle for life was put in relation to the ethical problem. In the seventeenth century Thomas Hobbes gave the first impulse to the whole modern discussion of ethical principles in his theory of bellum omnium contra omnes. Men, he taught, are in the state of nature enemies one of another, and they live either in fright or in the glory of power. But it was not the opinion of Hobbes that this made ethics impossible. On the contrary, he found a standard for virtue and vice in the fact that some qualities and actions have a tendency to bring us out of the state of war and to secure peace, while other qualities have a contrary tendency. In the eighteenth century even Immanuel Kant's ideal ethics had--so far as can be seen--a similar origin. Shortly before the foundation of his definitive ethics, Kant wrote his "Idee zu einer allgemeinen Weltgeschichte" (1784), where--in a way which reminds us of Hobbes, and is prophetic of Darwin--he describes the forward-driving power of struggle in the human world. It is here as with the struggle of the trees for light and air, through which they compete with one another in height. Anxiety about war can only be allayed by an ordinance which gives everyone his full liberty under acknowledgment of the equal liberty of others. And such ordinance and acknowledgment are also attributes of the content of the moral law, as Kant proclaimed it in the year after the publication of his essay (1785) (Cf. my "History of Modern Philosophy" (English translation London, 1900), I. pages 76-79.) Kant really came to his ethics by the way of evolution, though he afterwards disavowed it. Similarly the same line of thought may be traced in Hegel though it has been disguised in the form of speculative dialectics. ("Herrschaft und Knechtschaft", "Phanomenologie des Geistes", IV. A., Leiden, 1907.) And in Schopenhauer's theory of the blind will to live and its abrogation by the ethical feeling, which is founded on universal sympathy, we have a more individualistic form of the same idea.

It was, then, not entirely a foreign point of view which Darwin introduced into ethical thought, even if we take no account of the poetical character of the word "struggle" and of the more direct adaptation, through the use and non-use of power, which Darwin also emphasised. In "The Descent of Man" he has devoted a special chapter ("The Descent of Man", Vol. I. Ch. iii.) to a discussion of the origin of the ethical consciousness. The characteristic expression of this consciousness he found, just as Kant did, in the idea of "ought"; it was the origin of this new idea which should be explained. His hypothesis was that the ethical "ought" has its origin in the social and parental instincts, which, as well as other instincts (e.g. the instinct of self-preservation), lie deeper than pleasure and pain. In many species, not least in the human species, these instincts are fostered by natural selection; and when the powers of memory and comparison are developed, so that single acts can be valued according to the claims of the deep social instinct, then consciousness of duty and remorse are possible. Blind instinct has developed to conscious ethical will.

As already stated, Darwin, as a moral philosopher belongs to the school that was founded by Shaftesbury, and was afterwards represented by Hutcheson, Hume, Adam Smith, Comte and Spencer. His merit is, first, that he has given this tendency of thought a biological foundation, and that he has stamped on it a doughty character in showing that ethical ideas and sentiments, rightly conceived, are forces which are at work in the struggle for life.

There are still many questions to solve. Not only does the ethical development within the human species contain features still unexplained (The works of Westermarck and Hobhouse throw new light on many of these features.); but we are confronted by the great problem whether after all a genetic historical theory can be of decisive importance here. To every consequent ethical consciousness there is a standard of value, a primordial value which determines the single ethical judgments as their last presupposition, and the "rightness" of this basis, the "value" of this value can as little be discussed as the "rationality" of our logical principles. There is here revealed a possibility of ethical scepticism which evolutionistic ethics (as well as intuitive or rationalistic ethics) has overlooked. No demonstration can show that the results of the ethical development are definitive and universal. We meet here again with the important opposition of systematisation and evolution. There will, I think, always be an open question here, though comparative ethics, of which we have so far only the first attempts, can do much to throw light on it.

It would carry us too far to discuss all the philosophical works on ethics, which have been influenced directly or indirectly by evolutionism. I may, however, here refer to the book of C.M. Williams, "A Review of the Systems of Ethics founded on the Theory of Evolution" (New York and London, 1893.), in which, besides Darwin, the following authors are reviewed: Wallace, Haeckel, Spencer, Fiske, Rolph, Barratt, Stephen, Carneri, Hoffding, Gizycki, Alexander, Ree. As works which criticise evolutionistic ethics from an intuitive point of view and in an instructive way, may be cited: Guyau "La morale anglaise contemporaine" (Paris, 1879.), and Sorley, "Ethics of Naturalism". I will only mention some interesting contributions to ethical discussion which can be found in Darwinism besides the idea of struggle for life.

The attention which Darwin has directed to variations has opened our eyes to the differences in human nature as well as in nature generally. There is here a fact of great importance for ethical thought, no matter from what ultimate premiss it starts. Only from a very abstract point of view can different individuals be treated in the same manner. The most eminent ethical thinkers, men such as Jeremy Bentham and Immanuel Kant, who discussed ethical questions from very opposite standpoints, agreed in regarding all men as equal in respect of ethical endowment. In regard to Bentham, Leslie Stephen remarks: "He is determined to be thoroughly empirical, to take men as he found them. But his utilitarianism supposed that men's views of happiness and utility were uniform and clear, and that all that was wanted was to show them the means by which their ends could be reached." ("English literature and society in the eighteenth century", London, 1904, page 187.) And Kant supposed that every man would find the "categorical imperative" in his consciousness, when he came to sober reflexion, and that all would have the same qualifications to follow it. But if continual variations, great or small, are going on in human nature, it is the duty of ethics to make allowance for them, both in making claims, and in valuing what is done. A new set of ethical problems have their origin here. (Cf. my paper, "The law of relativity in Ethics," "International Journal of Ethics", Vol. I. 1891, pages 37-62.) It is an interesting fact that Stuart Mill's book "On Liberty" appeared in the same year as "The Origin of Species". Though Mill agreed with Bentham about the original equality of all men's endowments, he regarded individual differences as a necessary result of physical and social influences, and he claimed that free play shall be allowed to differences of character so far as is possible without injury to other men. It is a condition of individual and social progress that a man's mode of action should be determined by his own character and not by tradition and custom, nor by abstract rules. This view was to be corroborated by the theory of Darwin.

But here we have reached a point of view from which the criticism, which in recent years has often been directed against Darwin--that small variations are of no importance in the struggle for life--is of no weight. From an ethical standpoint, and particularly from the ethical standpoint of Darwin himself, it is a duty to foster individual differences that can be valuable, even though they can neither be of service for physical preservation nor be physically inherited. The distinction between variation and mutation is here without importance. It is quite natural that biologists should be particularly interested in such variations as can be inherited and produce new species. But in the human world there is not only a physical, but also a mental and social heredity. When an ideal human character has taken form, then there is shaped a type, which through imitation and influence can become an important factor in subsequent development, even if it cannot form a species in the biological sense of the word. Spiritually strong men often succumb in the physical struggle for life; but they can nevertheless be victorious through the typical influence they exert, perhaps on very distant generations, if the remembrance of them is kept alive, be it in legendary or in historical form. Their very failure can show that a type has taken form which is maintained at all risks, a standard of life which is adhered to in spite of the strongest opposition. The question "to be or not to be" can be put from very different levels of being: it has too often been considered a consequence of Darwinism that this question is only to be put from the lowest level. When a stage is reached, where ideal (ethical, intellectual, aesthetic) interests are concerned, the struggle for life is a struggle for the preservation of this stage. The giving up of a higher standard of life is a sort of death; for there is not only a physical, there is also a spiritual, death.

VI.

The Socratic character of Darwin's mind appears in his wariness in drawing the last consequences of his doctrine, in contrast both with the audacious theories of so many of his followers and with the consequences which his antagonists were busy in drawing. Though he, as we have seen, saw from the beginning that his hypothesis would occasion "a whole of metaphysics," he was himself very reserved as to the ultimate questions, and his answers to such questions were extorted from him.

As to the question of optimism and pessimism, Darwin held that though pain and suffering were very often the ways by which animals were led to pursue that course of action which is most beneficial to the species, yet pleasurable feelings were the most habitual guides. "We see this in the pleasure from exertion, even occasionally from great exertion of the body or mind, in the pleasure of our daily meals, and especially in the pleasure derived from sociability, and from loving our families." But there was to him so much suffering in the world that it was a strong argument against the existence of an intelligent First Cause. ("Life and Letters" Vol. I. page 310.)

It seems to me that Darwin was not so clear on another question, that of the relation between improvement and adaptation. He wrote to Lyell: "When you contrast natural selection and 'improvement,' you seem always to overlook... that every step in the natural selection of each species implies improvement in that species IN RELATION TO ITS CONDITION OF LIFE... Improvement implies, I suppose, EACH FORM OBTAINING MANY PARTS OR ORGANS, all excellently adapted for their functions." "All this," he adds, "seems to me quite compatible with certain forms fitted for simple conditions, remaining unaltered, or being degraded." (Ibid. Vol. II. page 177.) But the great question is, if the conditions of life will in the long run favour "improvement" in the sense of differentiation (or harmony of differentiation and integration). Many beings are best adapted to their conditions of life if they have few organs and few necessities. Pessimism would not only be the consequence, if suffering outweighed happiness, but also if the most elementary forms of happiness were predominant, or if there were a tendency to reduce the standard of life to the simplest possible, the contentment of inertia or stable equilibrium. There are animals which are very highly differentiated and active in their young state, but later lose their complex organisation and concentrate themselves on the one function of nutrition. In the human world analogies to this sort of adaptation are not wanting. Young "idealists" very often end as old "Philistines." Adaptation and progress are not the same.

Another question of great importance in respect to human evolution is, whether there will be always a possibility for the existence of an impulse to progress, an impulse to make great claims on life, to be active and to alter the conditions of life instead of adapting to them in a passive manner. Many people do not develop because they have too few necessities, and because they have no power to imagine other conditions of life than those under which they live. In his remarks on "the pleasure from exertion" Darwin has a point of contact with the practical idealism of former times--with the ideas of Lessing and Goethe, of Condorcet and Fichte. The continual striving which was the condition of salvation to Faust's soul, is also the condition of salvation to mankind. There is a holy fire which we ought to keep burning, if adaptation is really to be improvement. If, as I have tried to show in my "Philosophy of Religion", the innermost core of all religion is faith in the persistence of value in the world, and if the highest values express themselves in the cry "Excelsior!" then the capital point is, that this cry should always be heard and followed. We have here a corollary of the theory of evolution in its application to human life.

Darwin declared himself an agnostic, not only because he could not harmonise the large amount of suffering in the world with the idea of a God as its first cause, but also because he "was aware that if we admit a first cause, the mind still craves to know whence it came and how it arose." ("Life and Letters", Vol. I. page 306.) He saw, as Kant had seen before him and expressed in his "Kritik der Urtheilskraft", that we cannot accept either of the only two possibilities which we are able to conceive: chance (or brute force) and design. Neither mechanism nor teleology can give an absolute answer to ultimate questions. The universe, and especially the organic life in it, can neither be explained as a mere combination of absolute elements nor as the effect of a constructing thought. Darwin concluded, as Kant, and before him Spinoza, that the oppositions and distinctions which our experience presents, cannot safely be regarded as valid for existence in itself. And, with Kant and Fichte, he found his stronghold in the conviction that man has something to do, even if he cannot solve all enigmas. "The safest conclusion seems to me that the whole subject is beyond the scope of man's intellect; but man can do his duty." (Ibid. page 307.)

Is this the last word of human thought? Does not the possibility, that man can do his duty, suppose that the conditions of life allow of continuous ethical striving, so that there is a certain harmony between cosmic order and human ideals? Darwin himself has shown how the consciousness of duty can arise as a natural result of evolution. Moreover there are lines of evolution which have their end in ethical idealism, in a kingdom of values, which must struggle for life as all things in the world must do, but a kingdom which has its firm foundation in reality.

XXIII. DARWINISM AND SOCIOLOGY. By C. Bougle.

Professor of Social Philosophy in the University of Toulouse and Deputy-Professor at the Sorbonne, Paris.

How has our conception of social phenomena, and of their history, been affected by Darwin's conception of Nature and the laws of its transformations? To what extent and in what particular respects have the discoveries and hypotheses of the author of "The Origin of Species" aided the efforts of those who have sought to construct a science of society?

To such a question it is certainly not easy to give any brief or precise answer. We find traces of Darwinism almost everywhere. Sociological systems differing widely from each other have laid claim to its authority; while, on the other hand, its influence has often made itself felt only in combination with other influences. The Darwinian thread is worked into a hundred patterns along with other threads.

To deal with the problem, we must, it seems, first of all distinguish the more general conclusions in regard to the evolution of living beings, which are the outcome of Darwinism, from the particular explanations it offers of the ways and means by which that evolution is effected. That is to say, we must, as far as possible, estimate separately the influence of Darwin as an evolutionist and Darwin as a selectionist.

The nineteenth century, said Cournot, has witnessed a mighty effort to "reintegrer l'homme dans la nature." From divers quarters there has been a methodical reaction against the persistent dualism of the Cartesian tradition, which was itself the unconscious heir of the Christian tradition. Even the philosophy of the eighteenth century, materialistic as were for the most part the tendencies of its leaders, seemed to revere man as a being apart, concerning whom laws might be formulated a priori. To bring him down from his pedestal there was needed the marked predominance of positive researches wherein no account was taken of the "pride of man." There can be no doubt that Darwin has done much to familiarise us with this attitude. Take for instance the first part of "The Descent of Man": it is an accumulation of typical facts, all tending to diminish the distance between us and our brothers, the lower animals. One might say that the naturalist had here taken as his motto, "Whosoever shall exalt himself shall be abased; and he that shall humble himself shall be exalted." Homologous structures, the survival in man of certain organs of animals, the rudiments in the animal of certain human faculties, a multitude of facts of this sort, led Darwin to the conclusion that there is no ground for supposing that the "king of the universe" is exempt from universal laws. Thus belief in the imperium in imperio has been, as it were, whittled away by the progress of the naturalistic spirit, itself continually strengthened by the conquests of the natural sciences. The tendency may, indeed, drag the social sciences into overstrained analogies, such, for instance, as the assimilation of societies to organisms. But it will, at least, have had the merit of helping sociology to shake off the pre-conception that the groups formed by men are artificial, and that history is completely at the mercy of chance. Some years before the appearance of "The Origin of Species", Auguste Comte had pointed out the importance, as regards the unification of positive knowledge, of the conviction that the social world, the last refuge of spiritualism, is itself subject to determininism. It cannot be doubted that the movement of thought which Darwin's discoveries promoted contributed to the spread of this conviction, by breaking down the traditional barrier which cut man off from Nature.

But Nature, according to modern naturalists, is no immutable thing: it is rather perpetual movement, continual progression. Their discoveries batter a breach directly into the Aristotelian notion of species; they refuse to see in the animal world a collection of immutable types, distinct from all eternity, and corresponding, as Cuvier said, to so many particular thoughts of the Creator. Darwin especially congratulated himself upon having been able to deal this doctrine the coup de grace: immutability is, he says, his chief enemy; and he is concerned to show--therein following up Lyell's work--that everything in the organic world, as in the inorganic, is explained by insensible but incessant transformations. "Nature makes no leaps"--"Nature knows no gaps": these two dicta form, as it were, the two landmarks between which Darwin's idea of transformation is worked out. That is to say, the development of Darwinism is calculated to further the application of the philosophy of Becoming to the study of human institutions.

The progress of the natural sciences thus brings unexpected reinforcements to the revolution which the progress of historical discipline had begun. The first attempt to constitute an actual science of social phenomena--that, namely, of the economists--had resulted in laws which were called natural, and which were believed to be eternal and universal, valid for all times and all places. But this perpetuality, brother, as Knies said, of the immutability of the old zoology, did not long hold out against the ever swelling tide of the historical movement. Knowledge of the transformations that had taken place in language, of the early phases of the family, of religion, of property, had all favoured the revival of the Heraclitean view: panta rei. As to the categories of political economy, it was soon to be recognised, as by Lassalle, that they too are only historical. The philosophy of history, moreover, gave expression under various forms to the same tendency. Hegel declares that "all that is real is rational," but at the same time he shows that all that is real is ephemeral, and that for history there is nothing fixed beneath the sun. It is this sense of universal evolution that Darwin came with fresh authority to enlarge. It was in the name of biological facts themselves that he taught us to see only slow metamorphoses in the history of institutions, and to be always on the outlook for survivals side by side with rudimentary forms. Anyone who reads "Primitive Culture", by Tylor,--a writer closely connected with Darwin--will be able to estimate the services which these cardinal ideas were to render to the social sciences when the age of comparative research had succeeded to that of a priori construction.

Let us note, moreover, that the philosophy of Becoming in passing through the Darwinian biology became, as it were, filtered: it got rid of those traces of finalism, which, under different forms, it had preserved through all the systems of German Romanticism. Even in Herbert Spencer, it has been plausibly argued, one can detect something of that sort of mystic confidence in forces spontaneously directing life, which forms the very essence of those systems. But Darwin's observations were precisely calculated to render such an hypothesis futile. At first people may have failed to see this; and we call to mind the ponderous sarcasms of Flourens when he objected to the theory of Natural Selection that it attributed to nature a power of free choice. "Nature endowed with will! That was the final error of last century; but the nineteenth no longer deals in personifications." (P. Flourens, "Examen du Livre de M. Darwin sur l'Origine des Especes", page 53, Paris, 1864. See also Huxley, "Criticisms on the 'Origin of Species'", "Collected Essays", Vol. II, page 102, London, 1902.) In fact Darwin himself put his readers on their guard against the metaphors he was obliged to use. The processes by which he explains the survival of the fittest are far from affording any indication of the design of some transcendent breeder. Nor, if we look closely, do they even imply immanent effort in the animal; the sorting out can be brought about mechanically, simply by the action of the environment. In this connection Huxley could with good reason maintain that Darwin's originality consisted in showing how harmonies which hitherto had been taken to imply the agency of intelligence and will could be explained without any such intervention. So, when later on, objective sociology declares that, even when social phenomena are in question, all finalist preconceptions must be distrusted if a science is to be constituted, it is to Darwin that its thanks are due; he had long been clearing paths for it which lay well away from the old familiar road trodden by so many theories of evolution.

This anti-finalist doctrine, when fully worked out, was, moreover, calculated to aid in the needful dissociation of two notions: that of evolution and that of progress. In application to society these had long been confounded; and, as a consequence, the general idea seemed to be that only one type of evolution was here possible. Do we not detect such a view in Comte's sociology, and perhaps even in Herbert Spencer's? Whoever, indeed, assumes an end for evolution is naturally inclined to think that only one road leads to that end. But those whose minds the Darwinian theory has enlightened are aware that the transformations of living beings depend primarily upon their conditions, and that it is these conditions which are the agents of selection from among individual variations. Hence, it immediately follows that transformations are not necessarily improvements. Here, Darwin's thought hesitated. Logically his theory proves, as Ray Lankester pointed out, that the struggle for existence may have as its outcome degeneration as well as amelioration: evolution may be regressive as well as progressive. Then, too--and this is especially to be borne in mind--each species takes its good where it finds it, seeks its own path and survives as best it can. Apply this notion to society and you arrive at the theory of multilinear evolution. Divergencies will no longer surprise you. You will be forewarned not to apply to all civilisations the same measure of progress, and you will recognise that types of evolution may differ just as social species themselves differ. Have we not here one of the conceptions which mark off sociology proper from the old philosophy of history?

But if we are to estimate the influence of Darwinism upon sociological conceptions, we must not dwell only upon the way in which Darwin impressed the general notion of evolution upon the minds of thinkers. We must go into details. We must consider the influence of the particular theories by which he explained the mechanism of this evolution. The name of the author of "The Origin of Species" has been especially attached, as everyone knows, to the doctrines of "natural selection" and of "struggle for existence," completed by the notion of "individual variation." These doctrines were turned to account by very different schools of social philosophy. Pessimistic and optimistic, aristocratic and democratic, individualistic and socialistic systems were to war with each other for years by casting scraps of Darwinism at each other's heads.

It was the spectacle of human contrivance that suggested to Darwin his conception of natural selection. It was in studying the methods of pigeon breeders that he divined the processes by which nature, in the absence of design, obtains analogous results in the differentiation of types. As soon as the importance of artificial selection in the transformation of species of animals was understood, reflection naturally turned to the human species, and the question arose, How far do men observe, in connection with themselves, those laws of which they make practical application in the case of animals? Here we come upon one of the ideas which guided the researches of Galton, Darwin's cousin. The author of "Inquiries into Human Faculty and its Development" ("Inquiries into Human Faculty", pages 1, 2, 3 sq., London, 1883.), has often expressed his surprise that, considering all the precautions taken, for example, in the breeding of horses, none whatever are taken in the breeding of the human species. It seems to be forgotten that the species suffers when the "fittest" are not able to perpetuate their type. Ritchie, in his "Darwinism and Politics" ("Darwinism and Politics" pages 9, 22, London, 1889.) reminds us of Darwin's remark that the institution of the peerage might be defended on the ground that peers, owing to the prestige they enjoy, are enabled to select as wives "the most beautiful and charming women out of the lower ranks." ("Life and Letters of Charles Darwin", II. page 385.) But, says Galton, it is as often as not "heiresses" that they pick out, and birth statistics seem to show that these are either less robust or less fecund than others. The truth is that considerations continue to preside over marriage which are entirely foreign to the improvement of type, much as this is a condition of general progress. Hence the importance of completing Odin's and De Candolle's statistics which are designed to show how characters are incorporated in organisms, how they are transmitted, how lost, and according to what law eugenic elements depart from the mean or return to it.

But thinkers do not always content themselves with undertaking merely the minute researches which the idea of Selection suggests. They are eager to defend this or that thesis. In the name of this idea certain social anthropologists have recast the conception of the process of civilisation, and have affirmed that Social Selection generally works against the trend of Natural Selection. Vacher de Lapouge--following up an observation by Broca on the point--enumerates the various institutions, or customs, such as the celibacy of priests and military conscription, which cause elimination or sterilisation of the bearers of certain superior qualities, intellectual or physical. In a more general way he attacks the democratic movement, a movement, as P. Bourget says, which is "anti-physical" and contrary to the natural laws of progress; though it has been inspired "by the dreams of that most visionary of all centuries, the eighteenth." (V. de Lapouge, "Les Selections sociales", page 259, Paris, 1896.) The "Equality" which levels down and mixes (justly condemned, he holds, by the Comte de Gobineau), prevents the aristocracy of the blond dolichocephales from holding the position and playing the part which, in the interests of all, should belong to them. Otto Ammon, in his "Natural Selection in Man", and in "The Social Order and its Natural Bases" ("Die naturliche Auslese beim Menschen", Jena, 1893; "Die Gesellschaftsordnung und ihre naturlichen Grundlagen". "Entwurf einer Sozialanthropologie", Jena, 1896.), defended analogous doctrines in Germany; setting the curve representing frequency of talent over against that of income, he attempted to show that all democratic measures which aim at promoting the rise in the social scale of the talented are useless, if not dangerous; that they only increase the panmixia, to the great detriment of the species and of society.

Among the aristocratic theories which Darwinism has thus inspired we must reckon that of Nietzsche. It is well known that in order to complete his philosophy he added biological studies to his philological; and more than once in his remarks upon the "Wille zur Macht" he definitely alludes to Darwin; though it must be confessed that it is generally in order to proclaim the in sufficiency of the processes by which Darwin seeks to explain the genesis of species. Nevertheless, Nietzsche's mind is completely possessed by an ideal of Selection. He, too, has a horror of panmixia. The naturalists' conception of "the fittest" is joined by him to that of the "hero" of romance to furnish a basis for his doctrine of the Superman. Let us hasten to add, moreover, that at the very moment when support was being sought in the theory of Selection for the various forms of the aristocratic doctrine, those same forms were being battered down on another side by means of that very theory. Attention was drawn to the fact that by virtue of the laws which Darwin himself had discovered isolation leads to etiolation. There is a risk that the privilege which withdraws the privileged elements of Society from competition will cause them to degenerate. In fact, Jacoby in his "Studies in Selection, in connexion with Heredity in Man", ("Etudes sur la Selection dans ses rapports avec l'heredite chez l'homme", Paris, page 481, 1881.), concludes that "sterility, mental debility, premature death and, finally, the extinction of the stock were not specially and exclusively the fate of sovereign dynasties; all privileged classes, all families in exclusively elevated positions share the fate of reigning families, although in a minor degree and in direct proportion to the loftiness of their social standing. From the mass of human beings spring individuals, families, races, which tend to raise themselves above the common level; painfully they climb the rugged heights, attain the summits of power, of wealth, of intelligence, of talent, and then, no sooner are they there than they topple down and disappear in gulfs of mental and physical degeneracy." The demographical researches of Hansen ("Die drei Bevolkerungsstufen", Munich, 1889.) (following up and completing Dumont's) tended, indeed, to show that urban as well as feudal aristocracies, burgher classes as well as noble castes, were liable to become effete. Hence it might well be concluded that the democratic movement, operating as it does to break down class barriers, was promoting instead of impeding human selection.

So we see that, according to the point of view, very different conclusions have been drawn from the application of the Darwinian idea of Selection to human society. Darwin's other central idea, closely bound up with this, that, namely, of the "struggle for existence" also has been diversely utilised. But discussion has chiefly centered upon its signification. And while some endeavour to extend its application to everything, we find others trying to limit its range. The conception of a "struggle for existence" has in the present day been taken up into the social sciences from natural science, and adopted. But originally it descended from social science to natural. Darwin's law is, as he himself said, only Malthus' law generalised and extended to the animal world: a growing disproportion between the supply of food and the number of the living is the fatal order whence arises the necessity of universal struggle, a struggle which, to the great advantage of the species, allows only the best equipped individuals to survive. Nature is regarded by Huxley as an immense arena where all living beings are gladiators. ("Evolution and Ethics", page 200; "Collected Essays", Vol. IX, London, 1894.)

Such a generalisation was well adapted to feed the stream of pessimistic thought; and it furnished to the apologists of war, in particular, new arguments, weighted with all the authority which in these days attaches to scientific deliverances. If people no longer say, as Bonald did, and Moltke after him, that war is a providential fact, they yet lay stress on the point that it is a natural fact. To the peace party Dragomirov's objection is urged that its attempts are contrary to the fundamental laws of nature, and that no sea wall can hold against breakers that come with such gathered force.

But in yet another quarter Darwinism was represented as opposed to philanthropic intervention. The defenders of the orthodox political economy found in it support for their tenets. Since in the organic world universal struggle is the condition of progress, it seemed obvious that free competition must be allowed to reign unchecked in the economic world. Attempts to curb it were in the highest degree imprudent. The spirit of Liberalism here seemed in conformity with the trend of nature: in this respect, at least, contemporary naturalism, offspring of the discoveries of the nineteenth century, brought reinforcements to the individualist doctrine, begotten of the speculations of the eighteenth: but only, it appeared, to turn mankind away for ever from humanitarian dreams. Would those whom such conclusions repelled be content to oppose to nature's imperatives only the protests of the heart? There were some who declared, like Brunetiere, that the laws in question, valid though they might be for the animal kingdom, were not applicable to the human. And so a return was made to the classic dualism. This indeed seems to be the line that Huxley took, when, for instance, he opposed to the cosmic process an ethical process which was its reverse.

But the number of thinkers whom this antithesis does not satisfy grows daily. Although the pessimism which claims authorisation from Darwin's doctrines is repugnant to them, they still are unable to accept the dualism which leaves a gulf between man and nature. And their endeavour is to link the two by showing that while Darwin's laws obtain in both kingdoms, the conditions of their application are not the same: their forms, and, consequently, their results, vary with the varying mediums in which the struggle of living beings takes place, with the means these beings have at disposal, with the ends even which they propose to themselves.

Here we have the explanation of the fact that among determined opponents of war partisans of the "struggle for existence" can be found: there are disciples of Darwin in the peace party. Novicow, for example, admits the "combat universel" of which Le Dantec ("Les Luttes entre Societies humaines et leurs phases successives", Paris, 1893,) speaks; but he remarks that at different stages of evolution, at different stages of life the same weapons are not necessarily employed. Struggles of brute force, armed hand to hand conflicts, may have been a necessity in the early phases of human societies. Nowadays, although competition may remain inevitable and indispensable, it can assume milder forms. Economic rivalries, struggles between intellectual influences, suffice to stimulate progress: the processes which these admit are, in the actual state of civilisation, the only ones which attain their end without waste, the only ones logical. From one end to the other of the ladder of life, struggle is the order of the day; but more and more as the higher rungs are reached, it takes on characters which are proportionately more "humane."

Reflections of this kind permit the introduction into the economic order of limitations to the doctrine of "laisser faire, laisser passer." This appeals, it is said, to the example of nature where creatures, left to themselves, struggle without truce and without mercy; but the fact is forgotten that upon industrial battlefields the conditions are different. The competitors here are not left simply to their natural energies: they are variously handicapped. A rich store of artificial resources exists in which some participate and others do not. The sides then are unequal; and as a consequence the result of the struggle is falsified. "In the animal world," said De Laveleye ("Le socialisme contemporain", page 384 (6th edition), Paris, 1891.), criticising Spencer, "the fate of each creature is determined by its individual qualities; whereas in civilised societies a man may obtain the highest position and the most beautiful wife because he is rich and well-born, although he may be ugly, idle or improvident; and then it is he who will perpetuate the species. The wealthy man, ill constituted, incapable, sickly, enjoys his riches and establishes his stock under the protection of the laws." Haycraft in England and Jentsch in Germany have strongly emphasised these "anomalies," which nevertheless are the rule. That is to say that even from a Darwinian point of view all social reforms can readily be justified which aim at diminishing, as Wallace said, inequalities at the start.

But we can go further still. Whence comes the idea that all measures inspired by the sentiment of solidarity are contrary to Nature's trend? Observe her carefully, and she will not give lessons only in individualism. Side by side with the struggle for existence do we not find in operation what Lanessan calls "association for existence." Long ago, Espinas had drawn attention to "societies of animals," temporary or permanent, and to the kind of morality that arose in them. Since then, naturalists have often insisted upon the importance of various forms of symbiosis. Kropotkin in "Mutual Aid" has chosen to enumerate many examples of altruism furnished by animals to mankind. Geddes and Thomson went so far as to maintain that "Each of the greater steps of progress is in fact associated with an increased measure of subordination of individual competition to reproductive or social ends, and of interspecific competition to co-operative association." (Geddes and Thomson, "The Evolution of Sex", page 311, London, 1889.) Experience shows, according to Geddes, that the types which are fittest to surmount great obstacles are not so much those who engage in the fiercest competitive struggle for existence, as those who contrive to temper it. From all these observations there resulted, along with a limitation of Darwinian pessimism, some encouragement for the aspirations of the collectivists.

And Darwin himself would, doubtless, have subscribed to these rectifications. He never insisted, like his rival, Wallace, upon the necessity of the solitary struggle of creatures in a state of nature, each for himself and against all. On the contrary, in "The Descent of Man", he pointed out the serviceableness of the social instincts, and corroborated Bagehot's statements when the latter, applying laws of physics to politics, showed the great advantage societies derived from intercourse and communion. Again, the theory of sexual evolution which makes the evolution of types depend increasingly upon preferences, judgments, mental factors, surely offers something to qualify what seems hard and brutal in the theory of natural selection.

But, as often happens with disciples, the Darwinians had out-Darwined Darwin. The extravagancies of social Darwinism provoked a useful reaction; and thus people were led to seek, even in the animal kingdom, for facts of solidarity which would serve to justify humane effort.

On quite another line, however, an attempt has been made to connect socialist tendencies with Darwinian principles. Marx and Darwin have been confronted; and writers have undertaken to show that the work of the German philosopher fell readily into line with that of the English naturalist and was a development of it. Such has been the endeavour of Ferri in Italy and of Woltmann in Germany, not to mention others. The founders of "scientific socialism" had, moreover, themselves thought of this reconciliation. They make more than one allusion to Darwin in works which appeared after 1859. And sometimes they use his theory to define by contrast their own ideal. They remark that the capitalist system, by giving free course to individual competition, ends indeed in a bellum omnium contra omnes; and they make it clear that Darwinism, thus understood, is as repugnant to them as to Duhring.

But it is at the scientific and not at the moral point of view that they place themselves when they connect their economic history with Darwin's work. Thanks to this unifying hypothesis, they claim to have constructed--as Marx does in his preface to "Das Kapital"--a veritable natural history of social evolution. Engels speaks in praise of his friend Marx as having discovered the true mainspring of history hidden under the veil of idealism and sentimentalism, and as having proclaimed in the primum vivere the inevitableness of the struggle for existence. Marx himself, in "Das Kapital", indicated another analogy when he dwelt upon the importance of a general technology for the explanation of this psychology:--a history of tools which would be to social organs what Darwinism is to the organs of animal species. And the very importance they attach to tools, to apparatus, to machines, abundantly proves that neither Marx nor Engels were likely to forget the special characters which mark off the human world from the animal. The former always remains to a great extent an artificial world. Inventions change the face of its institutions. New modes of production revolutionise not only modes of government, but modes even of collective thought. Therefore it is that the evolution of society is controlled by laws special to it, of which the spectacle of nature offers no suggestion.

If, however, even in this special sphere, it can still be urged that the evolution of the material conditions of society is in accord with Darwin's theory, it is because the influence of the methods of production is itself to be explained by the incessant strife of the various classes with each other. So that in the end Marx, like Darwin, finds the source of all progress in struggle. Both are grandsons of Heraclitus:--polemos pater panton. It sometimes happens, in these days, that the doctrine of revolutionary socialism is contrasted as rude and healthy with what may seem to be the enervating tendency of "solidarist" philanthropy: the apologists of the doctrine then pride themselves above all upon their faithfulness to Darwinian principles.

So far we have been mainly concerned to show the use that social philosophies have made of the Darwinian laws for practical purposes: in order to orientate society towards their ideals each school tries to show that the authority of natural science is on its side. But even in the most objective of theories, those which systematically make abstraction of all political tendencies in order to study the social reality in itself, traces of Darwinism are readily to be found.

Let us take for example Durkheim's theory of Division of Labour ("De la Division du Travail social", Paris, 1893.) The conclusions he derives from it are that whenever professional specialisation causes multiplication of distinct branches of activity, we get organic solidarity--implying differences--substituted for mechanical solidarity, based upon likenesses. The umbilical cord, as Marx said, which connects the individual consciousness with the collective consciousness is cut. The personality becomes more and more emancipated. But on what does this phenomenon, so big with consequences, itself depend? The author goes to social morphology for the answer: it is, he says, the growing density of population which brings with it this increasing differentiation of activities. But, again, why? Because the greater density, in thrusting men up against each other, augments the intensity of their competition for the means of existence; and for the problems which society thus has to face differentiation of functions presents itself as the gentlest solution.

Here one sees that the writer borrows directly from Darwin. Competition is at its maximum between similars, Darwin had declared; different species, not laying claim to the same food, could more easily coexist. Here lay the explanation of the fact that upon the same oak hundreds of different insects might be found. Other things being equal, the same applies to society. He who finds some unadopted speciality possesses a means of his own for getting a living. It is by this division of their manifold tasks that men contrive not to crush each other. Here we obviously have a Darwinian law serving as intermediary in the explanation of that progress of division of labour which itself explains so much in the social evolution.

And we might take another example, at the other end of the series of sociological systems. G. Tarde is a sociologist with the most pronounced anti-naturalistic views. He has attempted to show that all application of the laws of natural science to society is misleading. In his "Opposition Universelle" he has directly combatted all forms of sociological Darwinism. According to him the idea that the evolution of society can be traced on the same plan as the evolution of species is chimerical. Social evolution is at the mercy of all kinds of inventions, which by virtue of the laws of imitation modify, through individual to individual, through neighbourhood to neighbourhood, the general state of those beliefs and desires which are the only "quantities" whose variation matters to the sociologist. But, it may be rejoined, that however psychical the forces may be, they are none the less subject to Darwinian laws. They compete with each other; they struggle for the mastery of minds. Between types of ideas, as between organic forms, selection operates. And though it may be that these types are ushered into the arena by unexpected discoveries, we yet recognise in the psychological accidents, which Tarde places at the base of everything, near relatives of those small accidental variations upon which Darwin builds. Thus, accepting Tarde's own representations, it is quite possible to express in Darwinian terms, with the necessary transpositions, one of the most idealistic sociologies that have ever been constructed.

These few examples suffice. They enable us to estimate the extent of the field of influence of Darwinism. It affects sociology not only through the agency of its advocates but through that of its opponents. The questionings to which it has given rise have proved no less fruitful than the solutions it has suggested. In short, few doctrines, in the history of social philosophy, will have produced on their passage a finer outcrop of ideas.

XXIV. THE INFLUENCE OF DARWIN UPON RELIGIOUS THOUGHT. By P.N. Waggett, M.A., S.S.J.E.

The object of this paper is first to point out certain elements of the Darwinian influence upon Religious thought, and then to show reason for the conclusion that it has been, from a Christian point of view, satisfactory. I shall not proceed further to urge that the Christian apologetic in relation to biology has been successful. A variety of opinions may be held on this question, without disturbing the conclusion that the movements of readjustment have been beneficial to those who remain Christians, and this by making them more Christian and not only more liberal. The theologians may sometimes have retreated, but there has been an advance of theology. I know that this account incurs the charge of optimism. It is not the worst that could be made. The influence has been limited in personal range, unequal, even divergent, in operation, and accompanied by the appearance of waste and mischievous products. The estimate which follows requires for due balance a full development of many qualifying considerations. For this I lack space, but I must at least distinguish my view from the popular one that our difficulties about religion and natural science have come to an end.

Concerning the older questions about origins--the origin of the world, of species, of man, of reason, conscience, religion--a large measure of understanding has been reached by some thoughtful men. But meanwhile new questions have arisen, questions about conduct, regarding both the reality of morals and the rule of right action for individuals and societies. And these problems, still far from solution, may also be traced to the influence of Darwin. For they arise from the renewed attention to heredity, brought about by the search for the causes of variation, without which the study of the selection of variations has no sufficient basis.

Even the existing understanding about origins is very far from universal. On these points there were always thoughtful men who denied the necessity of conflict, and there are still thoughtful men who deny the possibility of a truce.

It must further be remembered that the earlier discussion now, as I hope to show, producing favourable results, created also for a time grave damage, not only in the disturbance of faith and the loss of men--a loss not repaired by a change in the currents of debate--but in what I believe to be a still more serious respect. I mean the introduction of a habit of facile and untested hypothesis in religious as in other departments of thought.

Darwin is not responsible for this, but he is in part the cause of it. Great ideas are dangerous guests in narrow minds; and thus it has happened that Darwin--the most patient of scientific workers, in whom hypothesis waited upon research, or if it provisionally outstepped it did so only with the most scrupulously careful acknowledgment--has led smaller and less conscientious men in natural science, in history, and in theology to an over-eager confidence in probable conjecture and a loose grip upon the facts of experience. It is not too much to say that in many quarters the age of materialism was the least matter-of-fact age conceivable, and the age of science the age which showed least of the patient temper of inquiry.

I have indicated, as shortly as I could, some losses and dangers which in a balanced account of Darwin's influence would be discussed at length.

One other loss must be mentioned. It is a defect in our thought which, in some quarters, has by itself almost cancelled all the advantages secured. I mean the exaggerated emphasis on uniformity or continuity; the unwillingness to rest any part of faith or of our practical expectation upon anything that from any point of view can be called exceptional. The high degree of success reached by naturalists in tracing, or reasonably conjecturing, the small beginnings of great differences, has led the inconsiderate to believe that anything may in time become anything else.

It is true that this exaggeration of the belief in uniformity has produced in turn its own perilous reaction. From refusing to believe whatever can be called exceptional, some have come to believe whatever can be called wonderful.

But, on the whole, the discontinuous or highly various character of experience received for many years too little deliberate attention. The conception of uniformity which is a necessity of scientific description has been taken for the substance of history. We have accepted a postulate of scientific method as if it were a conclusion of scientific demonstration. In the name of a generalisation which, however just on the lines of a particular method, is the prize of a difficult exploit of reflexion, we have discarded the direct impressions of experience; or, perhaps it is more true to say, we have used for the criticism of alleged experiences a doctrine of uniformity which is only valid in the region of abstract science. For every science depends for its advance upon limitation of attention, upon the selection out of the whole content of consciousness of that part or aspect which is measurable by the method of the science. Accordingly there is a science of life which rightly displays the unity underlying all its manifestations. But there is another view of life, equally valid, and practically sometimes more important, which recognises the immediate and lasting effect of crisis, difference, and revolution. Our ardour for the demonstration of uniformity of process and of minute continuous change needs to be balanced by a recognition of the catastrophic element in experience, and also by a recognition of the exceptional significance for us of events which may be perfectly regular from an impersonal point of view.

An exorbitant jealousy of miracle, revelation, and ultimate moral distinctions has been imported from evolutionary science into religious thought. And it has been a damaging influence, because it has taken men's attention from facts, and fixed them upon theories.

II.

With this acknowledgment of important drawbacks, requiring many words for their proper description, I proceed to indicate certain results of Darwin's doctrine which I believe to be in the long run wholly beneficial to Christian thought. These are:

The encouragement in theology of that evolutionary method of observation and study, which has shaped all modern research:

The recoil of Christian apologetics towards the ground of religious experience, a recoil produced by the pressure of scientific criticism upon other supports of faith:

The restatement, or the recovery of ancient forms of statement, of the doctrines of Creation and of divine Design in Nature, consequent upon the discussion of evolution and of natural selection as its guiding factor.

(1) The first of these is quite possibly the most important of all. It was well defined in a notable paper read by Dr Gore, now Bishop of Birmingham, to the Church Congress at Shrewsbury in 1896. We have learnt a new caution both in ascribing and in denying significance to items of evidence, in utterance or in event. There has been, as in art, a study of values, which secures perspective and solidity in our representation of facts. On the one hand, a given utterance or event cannot be drawn into evidence as if all items were of equal consequence, like sovereigns in a bag. The question whence and whither must be asked, and the particular thing measured as part of a series. Thus measured it is not less truly important, but it may be important in a lower degree. On the other hand, and for exactly the same reason, nothing that is real is unimportant. The "failures" are not mere mistakes. We see them, in St Augustine's words, as "scholar's faults which men praise in hope of fruit."

We cannot safely trace the origin of the evolutionistic method to the influence of natural science. The view is tenable that theology led the way. Probably this is a case of alternate and reciprocal debt. Quite certainly the evolutionist method in theology, in Christian history, and in the estimate of scripture, has received vast reinforcement from biology, in which evolution has been the ever present and ever victorious conception.

(2) The second effect named is the new willingness of Christian thinkers to take definite account of religious experience. This is related to Darwin through the general pressure upon religious faith of scientific criticism. The great advance of our knowledge of organisms has been an important element in the general advance of science. It has acted, by the varied requirements of the theory of organisms, upon all other branches of natural inquiry, and it held for a long time that leading place in public attention which is now occupied by speculative physics. Consequently it contributed largely to our present estimation of science as the supreme judge in all matters of inquiry (F.R. Tennant: "The Being of God in the light of Physical Science", in "Essays on some theological questions of the day". London, 1905.), to the supposed destruction of mystery and the disparagement of metaphysic which marked the last age, as well as to the just recommendation of scientific method in branches of learning where the direct acquisitions of natural science had no place.

Besides this, the new application of the idea of law and mechanical regularity to the organic world seemed to rob faith of a kind of refuge. The romantics had, as Berthelot ("Evolutionisme et Platonisme", pages 45, 46, 47. Paris, 1908.) shows, appealed to life to redress the judgments drawn from mechanism. Now, in Spencer, evolution gave us a vitalist mechanic or mechanical vitalism, and the appeal seemed cut off. We may return to this point later when we consider evolution; at present I only endeavour to indicate that general pressure of scientific criticism which drove men of faith to seek the grounds of reassurance in a science of their own; in a method of experiment, of observation, of hypothesis checked by known facts. It is impossible for me to do more than glance across the threshold of this subject. But it is necessary to say that the method is in an elementary stage of revival. The imposing success that belongs to natural science is absent: we fall short of the unchallengeable unanimity of the Biologists on fundamentals. The experimental method with its sure repetitions cannot be applied to our subject-matter. But we have something like the observational method of palaeontology and geographical distribution; and in biology there are still men who think that the large examination of varieties by way of geography and the search of strata is as truly scientific, uses as genuinely the logical method of difference, and is as fruitful in sure conclusions as the quasi-chemical analysis of Mendelian laboratory work, of which last I desire to express my humble admiration. Religion also has its observational work in the larger and possibly more arduous manner.

But the scientific work in religion makes its way through difficulties and dangers. We are far from having found the formula of its combination with the historical elements of our apologetic. It is exposed, therefore, to a damaging fire not only from unspiritualist psychology and pathology but also from the side of scholastic dogma. It is hard to admit on equal terms a partner to the old undivided rule of books and learning. With Charles Lamb, we cry in some distress, "must knowledge come to me, if it come at all, by some awkward experiment of intuition, and no longer by this familiar process of reading?" ("Essays of Elia", "New Year's Eve", page 41; Ainger's edition. London, 1899.) and we are answered that the old process has an imperishable value, only we have not yet made clear its connection with other contributions. And all the work is young, liable to be drawn into unprofitable excursions, side-tracked by self-deceit and pretence; and it fatally attracts, like the older mysticism, the curiosity and the expository powers of those least in sympathy with it, ready writers who, with all the air of extended research, have been content with narrow grounds for induction. There is a danger, besides, which accompanies even the most genuine work of this science and must be provided against by all its serious students. I mean the danger of unbalanced introspection both for individuals and for societies; of a preoccupation comparable to our modern social preoccupation with bodily health; of reflection upon mental states not accompanied by exercise and growth of the mental powers; the danger of contemplating will and neglecting work, of analysing conviction and not criticising evidence.

Still, in spite of dangers and mistakes, the work remains full of hopeful indications, and, in the best examples (Such an example is given in Baron F. von Hugel's recently finished book, the result of thirty years' research: "The Mystical Element of Religion, as studied in Saint Catherine of Genoa and her Friends". London, 1908.), it is truly scientific in its determination to know the very truth, to tell what we think, not what we think we ought to think. (G. Tyrrell, in "Mediaevalism", has a chapter which is full of the important MORAL element in a scientific attitude. "The only infallible guardian of truth is the spirit of truthfulness." "Mediaevalism" page 182, London, 1908.), truly scientific in its employment of hypothesis and verification, and in growing conviction of the reality of its subject-matter through the repeated victories of a mastery which advances, like science, in the Baconian road of obedience. It is reasonable to hope that progress in this respect will be more rapid and sure when religious study enlists more men affected by scientific desire and endowed with scientific capacity.

The class of investigating minds is a small one, possibly even smaller than that of reflecting minds. Very few persons at any period are able to find out anything whatever. There are few observers, few discoverers, few who even wish to discover truth. In how many societies the problems of philology which face every person who speaks English are left unattempted! And if the inquiring or the successfully inquiring class of minds is small, much smaller, of course, is the class of those possessing the scientific aptitude in an eminent degree. During the last age this most distinguished class was to a very great extent absorbed in the study of phenomena, a study which had fallen into arrears. For we stood possessed, in rudiment, of means of observation, means for travelling and acquisition, qualifying men for a larger knowledge than had yet been attempted. These were now to be directed with new accuracy and ardour upon the fabric and behaviour of the world of sense. Our debt to the great masters in physical science who overtook and almost out-stripped the task cannot be measured; and, under the honourable leadership of Ruskin, we may all well do penance if we have failed "in the respect due to their great powers of thought, or in the admiration due to the far scope of their discovery." ("Queen of the Air", Preface, page vii. London, 1906.) With what miraculous mental energy and divine good fortune--as Romans said of their soldiers--did our men of curiosity face the apparently impenetrable mysteries of nature! And how natural it was that immense accessions of knowledge, unrelated to the spiritual facts of life, should discredit Christian faith, by the apparent superiority of the new work to the feeble and unprogressive knowledge of Christian believers! The day is coming when men of this mental character and rank, of this curiosity, this energy and this good fortune in investigation, will be employed in opening mysteries of a spiritual nature. They will silence with masterful witness the over-confident denials of naturalism. They will be in danger of the widespread recognition which thirty years ago accompanied every utterance of Huxley, Tyndall, Spencer. They will contribute, in spite of adulation, to the advance of sober religious and moral science.

And this result will be due to Darwin, first because by raising the dignity of natural science, he encouraged the development of the scientific mind; secondly because he gave to religious students the example of patient and ardent investigation; and thirdly because by the pressure of naturalistic criticism the religious have been driven to ascertain the causes of their own convictions, a work in which they were not without the sympathy of men of science. (The scientific rank of its writer justifies the insertion of the following letter from the late Sir John Burdon-Sanderson to me. In the lecture referred to I had described the methods of Professor Moseley in teaching Biology as affording a suggestion of the scientific treatment of religion.)

Oxford, April 30, 1902.

Dear Sir,

I feel that I must express to you my thanks for the discourse which I had the pleasure of listening to yesterday afternoon.

I do not mean to say that I was able to follow all that you said as to the identity of Method in the two fields of Science and Religion, but I recognise that the "mysticism" of which you spoke gives us the only way by which the two fields can be brought into relation.

Among much that was memorable, nothing interested me more than what you said of Moseley.

No one, I am sure, knew better than you the value of his teaching and in what that value consisted.

Yours faithfully

J. Burdon-Sanderson. 31-2.)

In leaving the subject of scientific religious inquiry, I will only add that I do not believe it receives any important help--and certainly it suffers incidentally much damaging interruption--from the study of abnormal manifestations or abnormal conditions of personality.

(3) Both of the above effects seem to me of high, perhaps the very highest, importance to faith and to thought. But, under the third head, I name two which are more directly traceable to the personal work of Darwin, and more definitely characteristic of the age in which his influence was paramount: viz. the influence of the two conceptions of evolution and natural selection upon the doctrine of creation and of design respectively.

It is impossible here, though it is necessary for a complete sketch of the matter, to distinguish the different elements and channels of this Darwinian influence; in Darwin's own writings, in the vigorous polemic of Huxley, and strangely enough, but very actually for popular thought, in the teaching of the definitely anti-Darwinian evolutionist Spencer.

Under the head of the directly and purely Darwinian elements I should class as preeminent the work of Wallace and of Bates; for no two sets of facts have done more to fix in ordinary intelligent minds a belief in organic evolution and in natural selection as its guiding factor than the facts of geographical distribution and of protective colour and mimicry. The facts of geology were difficult to grasp and the public and theologians heard more often of the imperfection than of the extent of the geological record. The witness of embryology, depending to a great extent upon microscopic work, was and is beyond the appreciation of persons occupied in fields of work other than biology.

III.

From the influence in religion of scientific modes of thought we pass to the influence of particular biological conceptions. The former effect comes by way of analogy, example, encouragement and challenge; inspiring or provoking kindred or similar modes of thought in the field of theology; the latter by a collision of opinions upon matters of fact or conjecture which seem to concern both science and religion.

In the case of Darwinism the story of this collision is familiar, and falls under the heads of evolution and natural selection, the doctrine of descent with modification, and the doctrine of its guidance or determination by the struggle for existence between related varieties. These doctrines, though associated and interdependent, and in popular thought not only combined but confused, must be considered separately. It is true that the ancient doctrine of Evolution, in spite of the ingenuity and ardour of Lamarck, remained a dream tantalising the intellectual ambition of naturalists, until the day when Darwin made it conceivable by suggesting the machinery of its guidance. And, further, the idea of natural selection has so effectively opened the door of research and stimulated observation in a score of principal directions that, even if the Darwinian explanation became one day much less convincing than, in spite of recent criticism, it now is, yet its passing, supposing it to pass, would leave the doctrine of Evolution immeasurably and permanently strengthened. For in the interests of the theory of selection, "Fur Darwin," as Muller wrote, facts have been collected which remain in any case evidence of the reality of descent with modification.

But still, though thus united in the modern history of convictions, though united and confused in the collision of biological and traditional opinion, yet evolution and natural selection must be separated in theological no less than in biological estimation. Evolution seemed inconsistent with Creation; natural selection with Providence and Divine design.

Discussion was maintained about these points for many years and with much dark heat. It ranged over many particular topics and engaged minds different in tone, in quality, and in accomplishment. There was at most times a degree of misconception. Some naturalists attributed to theologians in general a poverty of thought which belonged really to men of a particular temper or training. The "timid theism" discerned in Darwin by so cautious a theologian as Liddon (H.P. Liddon, "The Recovery of S. Thomas"; a sermon preached in St Paul's, London, on April 23rd, 1882 (the Sunday after Darwin's death).) was supposed by many biologists to be the necessary foundation of an honest Christianity. It was really more characteristic of devout NATURALISTS like Philip Henry Gosse, than of religious believers as such. (Dr Pusey ("Unscience not Science adverse to Faith" 1878) writes: "The questions as to 'species,' of what variations the animal world is capable, whether the species be more or fewer, whether accidental variations may become hereditary... and the like, naturally fall under the province of science. In all these questions Mr Darwin's careful observations gained for him a deserved approbation and confidence.") The study of theologians more considerable and even more typically conservative than Liddon does not confirm the description of religious intolerance given in good faith, but in serious ignorance, by a disputant so acute, so observant and so candid as Huxley. Something hid from each other's knowledge the devoted pilgrims in two great ways of thought. The truth may be, that naturalists took their view of what creation was from Christian men of science who naturally looked in their own special studies for the supports and illustrations of their religious belief. Of almost every laborious student it may be said "Hic ab arte sua non recessit." And both the believing and the denying naturalists, confining habitual attention to a part of experience, are apt to affirm and deny with trenchant vigour and something of a narrow clearness "Qui respiciunt ad pauca, de facili pronunciant." (Aristotle, in Bacon, quoted by Newman in his "Idea of a University", page 78. London, 1873.)

Newman says of some secular teachers that "they persuade the world of what is false by urging upon it what is true." Of some early opponents of Darwin it might be said by a candid friend that, in all sincerity of devotion to truth, they tried to persuade the world of what is true by urging upon it what is false. If naturalists took their version of orthodoxy from amateurs in theology, some conservative Christians, instead of learning what evolution meant to its regular exponents, took their view of it from celebrated persons, not of the front rank in theology or in thought, but eager to take account of public movements and able to arrest public attention.

Cleverness and eloquence on both sides certainly had their share in producing the very great and general disturbance of men's minds in the early days of Darwinian teaching. But by far the greater part of that disturbance was due to the practical novelty and the profound importance of the teaching itself, and to the fact that the controversy about evolution quickly became much more public than any controversy of equal seriousness had been for many generations.

We must not think lightly of that great disturbance because it has, in some real sense, done its work, and because it is impossible in days of more coolness and light, to recover a full sense of its very real difficulties.

Those who would know them better should add to the calm records of Darwin ("Life and Letters" and "More Letters of Charles Darwin".) and to the story of Huxley's impassioned championship, all that they can learn of George Romanes. ("Life and Letters", London, 1896. "Thoughts on Religion", London, 1895. "Candid Examination of Theism", London, 1878.) For his life was absorbed in this very struggle and reproduced its stages. It began in a certain assured simplicity of biblical interpretation; it went on, through the glories and adventures of a paladin in Darwin's train, to the darkness and dismay of a man who saw all his most cherished beliefs rendered, as he thought, incredible. ("Never in the history of man has so terrific a calamity befallen the race as that which all who look may now (viz. in consequence of the scientific victory of Darwin) behold advancing as a deluge, black with destruction, resistless in might, uprooting our most cherished hopes, engulphing our most precious creed, and burying our highest life in mindless destruction."--"A Candid Examination of Theism", page 51.) He lived to find the freer faith for which process and purpose are not irreconcilable, but necessary to one another. His development, scientific, intellectual and moral, was itself of high significance; and its record is of unique value to our own generation, so near the age of that doubt and yet so far from it; certainly still much in need of the caution and courage by which past endurance prepares men for new emergencies. We have little enough reason to be sure that in the discussions awaiting us we shall do as well as our predecessors in theirs. Remembering their endurance of mental pain, their ardour in mental labour, the heroic temper and the high sincerity of controversialists on either side, we may well speak of our fathers in such words of modesty and self-judgment as Drayton used when he sang the victors of Agincourt. The progress of biblical study, in the departments of Introduction and Exegesis, resulting in the recovery of a point of view anciently tolerated if not prevalent, has altered some of the conditions of that discussion. In the years near 1858, the witness of Scripture was adduced both by Christian advocates and their critics as if unmistakeably irreconcilable with Evolution.

Huxley ("Science and Christian Tradition". London, 1904.) found the path of the blameless naturalist everywhere blocked by "Moses": the believer in revelation was generally held to be forced to a choice between revealed cosmogony and the scientific account of origins. It is not clear how far the change in Biblical interpretation is due to natural science, and how far to the vital movements of theological study which have been quite independent of the controversy about species. It belongs to a general renewal of Christian movement, the recovery of a heritage. "Special Creation"--really a biological rather than a theological conception,--seems in its rigid form to have been a recent element even in English biblical orthodoxy.

The Middle Ages had no suspicion that religious faith forbad inquiry into the natural origination of the different forms of life. Bartholomaeus Anglicus, an English Franciscan of the thirteenth century, was a mutationist in his way, as Aristotle, "the Philosopher" of the Christian Schoolmen, had been in his. So late as the seventeenth century, as we learn not only from early proceedings of the Royal Society, but from a writer so homely and so regularly pious as Walton, the variation of species and "spontaneous" generations had no theological bearing, except as instances of that various wonder of the world which in devout minds is food for devotion.

It was in the eighteenth century that the harder statement took shape. Something in the preciseness of that age, its exaltation of law, its cold passion for a stable and measured universe, its cold denial, its cold affirmation of the power of God, a God of ice, is the occasion of that rigidity of religious thought about the living world which Darwin by accident challenged, or rather by one of those movements of genius which, Goethe ("No productiveness of the highest kind... is in the power of anyone."--"Conversations of Goethe with Eckermann and Soret". London, 1850.) declares, are "elevated above all earthly control."

If religious thought in the eighteenth century was aimed at a fixed and nearly finite world of spirit, it followed in all these respects the secular and critical lead. ("La philosophie reformatrice du XVIIIe siecle" (Berthelot, "Evolutionisme et Platonisme", Paris, 1908, page 45.) ramenait la nature et la societe a des mecanismes que la pensee reflechie peut concevoir et recomposer." In fact, religion in a mechanical age is condemned if it takes any but a mechanical tone. Butler's thought was too moving, too vital, too evolutionary, for the sceptics of his time. In a rationalist, encyclopaedic period, religion also must give hard outline to its facts, it must be able to display its secret to any sensible man in the language used by all sensible men. Milton's prophetic genius furnished the eighteenth century, out of the depth of the passionate age before it, with the theological tone it was to need. In spite of the austere magnificence of his devotion, he gives to smaller souls a dangerous lead. The rigidity of Scripture exegesis belonged to this stately but imperfectly sensitive mode of thought. It passed away with the influence of the older rationalists whose precise denials matched the precise and limited affirmations of the static orthodoxy.

I shall, then, leave the specially biblical aspect of the debate--interesting as it is and even useful, as in Huxley's correspondence with the Duke of Argyll and others in 1892 ("Times", 1892, passim.)--in order to consider without complication the permanent elements of Christian thought brought into question by the teaching of evolution.

Such permanent elements are the doctrine of God as Creator of the universe, and the doctrine of man as spiritual and unique. Upon both the doctrine of evolution seemed to fall with crushing force.

With regard to Man I leave out, acknowledging a grave omission, the doctrine of the Fall and of Sin. And I do so because these have not yet, as I believe, been adequately treated: here the fruitful reaction to the stimulus of evolution is yet to come. The doctrine of sin, indeed, falls principally within the scope of that discussion which has followed or displaced the Darwinian; and without it the Fall cannot be usefully considered. For the question about the Fall is a question not merely of origins, but of the interpretation of moral facts whose moral reality must first be established.

I confine myself therefore to Creation and the dignity of man.

The meaning of evolution, in the most general terms, is that the differentiation of forms is not essentially separate from their behaviour and use; that if these are within the scope of study, that is also; that the world has taken the form we see by movements not unlike those we now see in progress; that what may be called proximate origins are continuous in the way of force and matter, continuous in the way of life, with actual occurrences and actual characteristics. All this has no revolutionary bearing upon the question of ultimate origins. The whole is a statement about process. It says nothing to metaphysicians about cause. It simply brings within the scope of observation or conjecture that series of changes which has given their special characters to the different parts of the world we see. In particular, evolutionary science aspires to the discovery of the process or order of the appearance of life itself: if it were to achieve its aim it could say nothing of the cause of this or indeed of the most familiar occurrences. We should have become spectators or convinced historians of an event which, in respect of its cause and ultimate meaning, would be still impenetrable.

With regard to the origin of species, supposing life already established, biological science has the well founded hopes and the measure of success with which we are all familiar. All this has, it would seem, little chance of collision with a consistent theism, a doctrine which has its own difficulties unconnected with any particular view of order or process. But when it was stated that species had arisen by processes through which new species were still being made, evolutionism came into collision with a statement, traditionally religious, that species were formed and fixed once for all and long ago.

What is the theological import of such a statement when it is regarded as essential to belief in God? Simply that God's activity, with respect to the formation of living creatures, ceased at some point in past time.

"God rested" is made the touchstone of orthodoxy. And when, under the pressure of the evidences, we found ourselves obliged to acknowledge and assert the present and persistent power of God, in the maintenance and in the continued formation of "types," what happened was the abolition of a time-limit. We were forced only to a bolder claim, to a theistic language less halting, more consistent, more thorough in its own line, as well as better qualified to assimilate and modify such schemes as Von Hartmann's philosophy of the unconscious--a philosophy, by the way, quite intolerant of a merely mechanical evolution. (See Von Hartmann's "Wahrheit und Irrthum in Darwinismus". Berlin, 1875.)

Here was not the retrenchment of an extravagant assertion, but the expansion of one which was faltering and inadequate. The traditional statement did not need paring down so as to pass the meshes of a new and exacting criticism. It was itself a net meant to surround and enclose experience; and we must increase its size and close its mesh to hold newly disclosed facts of life. The world, which had seemed a fixed picture or model, gained first perspective and then solidity and movement. We had a glimpse of organic HISTORY; and Christian thought became more living and more assured as it met the larger view of life.

However unsatisfactory the new attitude might be to our critics, to Christians the reform was positive. What was discarded was a limitation, a negation. The movement was essentially conservative, even actually reconstructive. For the language disused was a language inconsistent with the definitions of orthodoxy; it set bounds to the infinite, and by implication withdrew from the creative rule all such processes as could be brought within the descriptions of research. It ascribed fixity and finality to that "creature" in which an apostle taught us to recognise the birth-struggles of an unexhausted progress. It tended to banish mystery from the world we see, and to confine it to a remote first age.

In the reformed, the restored, language of religion, Creation became again not a link in a rational series to complete a circle of the sciences, but the mysterious and permanent relation between the infinite and the finite, between the moving changes we know in part, and the Power, after the fashion of that observation, unknown, which is itself "unmoved all motion's source." (Hymn of the Church-- Rerum Deus tenax vigor, Immotus in te permanens.)

With regard to man it is hardly necessary, even were it possible, to illustrate the application of this bolder faith. When the record of his high extraction fell under dispute, we were driven to a contemplation of the whole of his life, rather than of a part and that part out of sight. We remembered again, out of Aristotle, that the result of a process interprets its beginnings. We were obliged to read the title of such dignity as we may claim, in results and still more in aspirations.

Some men still measure the value of great present facts in life--reason and virtue and sacrifice--by what a self-disparaged reason can collect of the meaner rudiments of these noble gifts. Mr Balfour has admirably displayed the discrepancy, in this view, between the alleged origin and the alleged authority of reason. Such an argument ought to be used not to discredit the confident reason, but to illuminate and dignify its dark beginnings, and to show that at every step in the long course of growth a Power was at work which is not included in any term or in all the terms of the series.

I submit that the more men know of actual Christian teaching, its fidelity to the past, and its sincerity in face of discovery, the more certainly they will judge that the stimulus of the doctrine of evolution has produced in the long run vigour as well as flexibility in the doctrine of Creation and of man.

I pass from Evolution in general to Natural Selection.

The character in religious language which I have for short called mechanical was not absent in the argument from design as stated before Darwin. It seemed to have reference to a world conceived as fixed. It pointed, not to the plastic capacity and energy of living matter, but to the fixed adaptation of this and that organ to an unchanging place or function.

Mr Hobhouse has given us the valuable phrase "a niche of organic opportunity." Such a phrase would have borne a different sense in non-evolutionary thought. In that thought, the opportunity was an opportunity for the Creative Power, and Design appeared in the preparation of the organism to fit the niche. The idea of the niche and its occupant growing together from simpler to more complex mutual adjustment was unwelcome to this teleology. If the adaptation was traced to the influence, through competition, of the environment, the old teleology lost an illustration and a proof. For the cogency of the proof in every instance depended upon the absence of explanation. Where the process of adaptation was discerned, the evidence of Purpose or Design was weak. It was strong only when the natural antecedents were not discovered, strongest when they could be declared undiscoverable.

Paley's favourite word is "Contrivance"; and for him contrivance is most certain where production is most obscure. He points out the physiological advantage of the valvulae conniventes to man, and the advantage for teleology of the fact that they cannot have been formed by "action and pressure." What is not due to pressure may be attributed to design, and when a "mechanical" process more subtle than pressure was suggested, the case for design was so far weakened. The cumulative proof from the multitude of instances began to disappear when, in selection, a natural sequence was suggested in which all the adaptations might be reached by the motive power of life, and especially when, as in Darwin's teaching, there was full recognition of the reactions of life to the stimulus of circumstance. "The organism fits the niche," said the teleologist, "because the Creator formed it so as to fit." "The organism fits the niche," said the naturalist, "because unless it fitted it could not exist." "It was fitted to survive," said the theologian. "It survives because it fits," said the selectionist. The two forms of statement are not incompatible; but the new statement, by provision of an ideally universal explanation of process, was hostile to a doctrine of purpose which relied upon evidences always exceptional however numerous. Science persistently presses on to find the universal machinery of adaptation in this planet; and whether this be found in selection, or in direct-effect, or in vital reactions resulting in large changes, or in a combination of these and other factors, it must always be opposed to the conception of a Divine Power here and there but not everywhere active.

For science, the Divine must be constant, operative everywhere and in every quality and power, in environment and in organism, in stimulus and in reaction, in variation and in struggle, in hereditary equilibrium, and in "the unstable state of species"; equally present on both sides of every strain, in all pressures and in all resistances, in short in the general wonder of life and the world. And this is exactly what the Divine Power must be for religious faith.

The point I wish once more to make is that the necessary readjustment of teleology, so as to make it depend upon the contemplation of the whole instead of a part, is advantageous quite as much to theology as to science. For the older view failed in courage. Here again our theism was not sufficiently theistic.

Where results seemed inevitable, it dared not claim them as God-given. In the argument from Design it spoke not of God in the sense of theology, but of a Contriver, immensely, not infinitely wise and good, working within a world, the scene, rather than the ever dependent outcome, of His Wisdom; working in such emergencies and opportunities as occurred, by forces not altogether within His control, towards an end beyond Himself. It gave us, instead of the awful reverence due to the Cause of all substance and form, all love and wisdom, a dangerously detached appreciation of an ingenuity and benevolence meritorious in aim and often surprisingly successful in contrivance.

The old teleology was more useful to science than to religion, and the design-naturalists ought to be gratefully remembered by Biologists. Their search for evidences led them to an eager study of adaptations and of minute forms, a study such as we have now an incentive to in the theory of Natural Selection. One hardly meets with the same ardour in microscopical research until we come to modern workers. But the argument from Design was never of great importance to faith. Still, to rid it of this character was worth all the stress and anxiety of the gallant old war. If Darwin had done nothing else for us, we are to-day deeply in his debt for this. The world is not less venerable to us now, not less eloquent of the causing mind, rather much more eloquent and sacred. But our wonder is not that "the underjaw of the swine works under the ground" or in any or all of those particular adaptations which Paley collected with so much skill, but that a purpose transcending, though resembling, our own purposes, is everywhere manifest; that what we live in is a whole, mutually sustaining, eventful and beautiful, where the "dead" forces feed the energies of life, and life sustains a stranger existence, able in some real measure to contemplate the whole, of which, mechanically considered, it is a minor product and a rare ingredient. Here, again, the change was altogether positive. It was not the escape of a vessel in a storm with loss of spars and rigging, not a shortening of sail to save the masts and make a port of refuge. It was rather the emergence from narrow channels to an open sea. We had propelled the great ship, finding purchase here and there for slow and uncertain movement. Now, in deep water, we spread large canvas to a favouring breeze.

The scattered traces of design might be forgotten or obliterated. But the broad impression of Order became plainer when seen at due distance and in sufficient range of effect, and the evidence of love and wisdom in the universe could be trusted more securely for the loss of the particular calculation of their machinery.

Many other topics of faith are affected by modern biology. In some of these we have learnt at present only a wise caution, a wise uncertainty. We stand before the newly unfolded spectacle of suffering, silenced; with faith not scientifically reassured but still holding fast certain other clues of conviction. In many important topics we are at a loss. But in others, and among them those I have mentioned, we have passed beyond this negative state and find faith positively strengthened and more fully expressed.

We have gained also a language and a habit of thought more fit for the great and dark problems that remain, less liable to damaging conflicts, equipped for more rapid assimilation of knowledge. And by this change biology itself is a gainer. For, relieved of fruitless encounters with popular religion, it may advance with surer aim along the path of really scientific life-study which was reopened for modern men by the publication of "The Origin of Species".

Charles Darwin regretted that, in following science, he had not done "more direct good" ("Life and Letters", Vol. III. page 359.) to his fellow-creatures. He has, in fact, rendered substantial service to interests bound up with the daily conduct and hopes of common men; for his work has led to improvements in the preaching of the Christian faith.

XXV. THE INFLUENCE OF DARWINISM ON THE STUDY OF RELIGIONS. By Jane Ellen Harrison.

Hon. D.Litt. (Durham), Hon. LL.D. (Aberdeen), Staff Lecturer and sometime Fellow of Newnham College, Cambridge.

Corresponding member of the German Archaeological Institute.

The title of my paper might well have been "the creation by Darwinism of the scientific study of Religions," but that I feared to mar my tribute to a great name by any shadow of exaggeration. Before the publication of "The Origin of Species" and "The Descent of Man", even in the eighteenth century, isolated thinkers, notably Hume and Herder, had conjectured that the orthodox beliefs of their own day were developments from the cruder superstitions of the past. These were however only particular speculations of individual sceptics. Religion was not yet generally regarded as a proper subject for scientific study, with facts to be collected and theories to be deduced. A Congress of Religions such as that recently held at Oxford would have savoured of impiety.

In the brief space allotted me I can attempt only two things; first, and very briefly, I shall try to indicate the normal attitude towards religion in the early part of the last century; second, and in more detail, I shall try to make clear what is the outlook of advanced thinkers to-day. (To be accurate I ought to add "in Europe." I advisedly omit from consideration the whole immense field of Oriental mysticism, because it has remained practically untouched by the influence of Darwinism.) From this second inquiry it will, I hope, be abundantly manifest that it is the doctrine of evolution that has made this outlook possible and even necessary.

The ultimate and unchallenged presupposition of the old view was that religion was a DOCTRINE, a body of supposed truths. It was in fact what we should now call Theology, and what the ancients called Mythology. Ritual was scarcely considered at all, and, when considered, it was held to be a form in which beliefs, already defined and fixed as dogma, found a natural mode of expression. This, it will be later shown, is a profound error or rather a most misleading half-truth. Creeds, doctrines, theology and the like are only a part, and at first the least important part, of religion.

Further, and the fact is important, this DOGMA, thus supposed to be the essential content of the "true" religion, was a teleological scheme complete and unalterable, which had been revealed to man once and for all by a highly anthropomorphic God, whose existence was assumed. The duty of man towards this revelation was to accept its doctrines and obey its precepts. The notion that this revelation had grown bit by bit out of man's consciousness and that his business was to better it would have seemed rank blasphemy. Religion, so conceived, left no place for development. "The Truth" might be learnt, but never critically examined; being thus avowedly complete and final, it was doomed to stagnation.

The details of this supposed revelation seem almost too naive for enumeration. As Hume observed, "popular theology has a positive appetite for absurdity." It is sufficient to recall that "revelation" included such items as the Creation (It is interesting to note that the very word "Creator" has nowadays almost passed into the region of mythology. Instead we have "L'Evolution Creatrice".) of the world out of nothing in six days; the making of Eve from one of Adam's ribs; the Temptation by a talking snake; the confusion of tongues at the tower of Babel; the doctrine of Original Sin; a scheme of salvation which demanded the Virgin Birth, Vicarious Atonement, and the Resurrection of the material body. The scheme was unfolded in an infallible Book, or, for one section of Christians, guarded by the tradition of an infallible Church, and on the acceptance or refusal of this scheme depended an eternity of weal or woe. There is not one of these doctrines that has not now been recast, softened down, mysticised, allegorised into something more conformable with modern thinking. It is hard for the present generation, unless their breeding has been singularly archaic, to realise that these amazing doctrines were literally held and believed to constitute the very essence of religion; to doubt them was a moral delinquency.

It had not, however, escaped the notice of travellers and missionaries that savages carried on some sort of practices that seemed to be religious, and believed in some sort of spirits or demons. Hence, beyond the confines illuminated by revealed truth, a vague region was assigned to NATURAL Religion. The original revelation had been kept intact only by one chosen people, the Jews, by them to be handed on to Christianity. Outside the borders of this Goshen the world had sunk into the darkness of Egypt. Where analogies between savage cults and the Christian religions were observed, they were explained as degradations; the heathen had somehow wilfully "lost the light." Our business was not to study but, exclusively, to convert them, to root out superstition and carry the torch of revelation to "Souls in heathen darkness lying." To us nowadays it is a commonplace of anthropological research that we must seek for the beginnings of religion in the religions of primitive peoples, but in the last century the orthodox mind was convinced that it possessed a complete and luminous ready-made revelation; the study of what was held to be a mere degradation seemed idle and superfluous.

But, it may be asked, if, to the orthodox, revealed religion was sacrosanct and savage religion a thing beneath consideration, why did not the sceptics show a more liberal spirit, and pursue to their logical issue the conjectures they had individually hazarded? The reason is simple and significant. The sceptics too had not worked free from the presupposition that the essence of religion is dogma. Their intellectualism, expressive of the whole eighteenth century, was probably in England strengthened by the Protestant doctrine of an infallible Book. Hume undoubtedly confused religion with dogmatic theology. The attention of orthodox and sceptics alike was focussed on the truth or falsity of certain propositions. Only a few minds of rare quality were able dimly to conceive that religion might be a necessary step in the evolution of human thought.

It is not a little interesting to note that Darwin, who was leader and intellectual king of his generation, was also in this matter to some extent its child. His attitude towards religion is stated clearly, in