Part 5

The lower animals devoid of backbones, and therefore called invertebrates, are not so well-known except to the student of comparative anatomy, because they are not so often met with, and because they are usually very small or microscopic; but in many respects their importance to the evolutionist surpasses that of the vertebrates. Their structural plans are far more varied, and they range more widely from higher and relatively complicated organisms to the unitary one-celled animals. A knowledge of some of them is essential for our present purpose, which is to learn how sure is the basis for the principles of relationship and how complete is the structural evidence of evolution.

Worms are represented in the minds of most people by the common earthworm or sandworm. The body in either case is made up of a series of segments or joints which agree closely throughout the animal in external appearance and in internal constitution. A section of the digestive tract, a pair of nerve centers, two funnel-like tubes for excretion, and similar blood vessels occur in each portion.

Precisely similar features are displayed by the crustacea, which seem to be so different. Every one is familiar with the appearance of lobsters and crabs. Even in these animals the body is composed of segments, but these are not like one another, nor are they freely movable throughout the body. Five are fused in all crustacea to make a head; in lower members of the order the eight succeeding segments are free, but in the lobster they are joined together and united with the head. The hinder part of this animal is a long abdomen whose segments remain more primitive and independent. But in a crab, the whole plan has been modified by the shortening and broadening of the head-thorax, and by the reduction of the abdomen, which is also turned under the anterior part of the body. The internal organic systems are constructed upon a worm plan with modifications. Nearly every one of the segments bears one pair of appendages, which can be referred by their forked nature to the two-parted, oarlike flaps of sandworms, but the appendages of crustacea have departed from their prototypes in functional respects and in details of structure. They are variously feelers, jaws, legs, pincers, and swimming paddles, evolved to serve different purposes, just as the limbs of the vertebrates we have described have become variously arms, wings, flippers and paddles in apes, bats, seals, and whales.

Butterflies, beetles, bees, and grasshoppers seem at first sight to be entirely different, even though they agree in being more or less segmented. But all of them have heads with four pairs of appendages of the same essential plan, middle thoracic regions of three segments more or less united, bearing three pairs of legs and usually two pairs of wings, while the hinder part is a freely jointed abdomen without real limbs. In these respects the countless varieties of insects agree so that they also like crustacea of various kinds seem to have been derived from wormlike animals with more simply segmented bodies. Indeed spiders and scorpions and their relatives of the group arachnida prove for similar reasons to be derivatives of the same original stock, and own cousins of the insects.

In nearly every one of the invertebrate branches we find representatives which interest us chiefly because they appear to have reached their present condition by retrograde evolution. Barnacles are really crustacea, but they have lost their eyes as well as some other structures that are most useful in animals with a free existence, because they have adopted a fixed mode of life, which has also brought about the loss of the original freely jointed character of the body. A tapeworm as an example of internal parasites is an extremely degenerate form which lacks a digestive tract, because this is superfluous in an animal which lives bathed in the nutrient fluids of its host. Comparing it in other respects with other low wormlike creatures, it appears to be a relative of peculiar simple worms with complete organization and independence of life. All these degenerate forms enlarge our conception of adaptation by adding the essential point that progress is not always the result of evolution. Indeed we have learned this in the case of vestigial and rudimentary structures of higher forms like whales, and now we find that entire animals may degenerate as a result of changes no less adaptive than progressive modifications.

Passing by other invertebrate groups made up of species arranged like higher animals in smaller and larger branches according to their degree of fundamental similarity, we arrive at a place in the scale occupied by two-layer animals without the highly developed and clearly differentiated organic systems of the forms above. The fresh-water animal Hydra exemplifies the creatures of this level, where also we find sea-anemones and the soft polyps which form corals and coral reefs by their combined skeletons. Hydra is an animal to which we must return again and again as we study one or another aspect of organic evolution. In general form it is a hollow cylinder closed at one end, by which it attaches itself, while at the upper end, surrounded by a group of tentacles, is the mouth which leads to the central cavity. The wall of this simple body is composed of two layers of cells, between which there is a gelatinous layer rarely invaded by cells. The inner layer lines the central space into which food organisms are thrust by the tentacles, and it is concerned primarily with digestion. The outer layer comprises cells for protection and sensation primarily. Cells of both layers have muscular prolongations which by their operation enable the whole animal to change its form and to move from one place to another.

It may seem that such an animal is totally unlike any of the higher and more complex types. In certain respects, however, it is identical with the other forms inasmuch as it performs all of the eight biological tasks demanded by nature. It is also similar in so far as its inner layer, like the innermost sheet of cells in higher forms, is concerned with problems of taking and preparing food, while the protective outer layer resembles in function the outermost covering of all animals higher in the scale. Beyond these a still more fundamental agreement is found in its cellular composition.

At the lower end of the animal scale are organisms which consist of one cell and nothing more. Amoeba, to which we must refer again and again, is an example of this group which possesses an overwhelming importance to the comparative student because the origins of all the characteristics of animals higher in the scale are to be found within it. Amoeba itself is a naked mass of protoplasm, about 1/100 of an inch in diameter, enclosing a nucleus. Its form is not constant during activity, for fingerlike processes called pseudopodia are pushed out tentatively in many directions to be followed as circumstances direct by the materials of the whole cell body. Other protozoa differ in possessing constant forms, or in having constant vibratile processes, or shells of some kind, while in still other cases like individuals combine to make colonies which are more or less definite and permanent. Here at the very foot of the organic scale are found animals which seem to be entirely different from those above. Upon examination they, like Hydra, prove to be the same as regards the number and kind of functions they perform, but in structural regards their evolutionary relation to all higher animals is indicated solely by the fact that they are cells composed of protoplasm. Nevertheless the principle which states that resemblance means consanguinity still holds true, for cellular constitution is a unique possession of things of the living world,--something which demonstrates the common origin of all living things just as truly as the "cat-ness" of our first series of examples reveals for a smaller group the significance of likeness and the nature of the basic law of comparative anatomy.

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Employing a figure of speech, we have climbed down the animal tree from the higher regions where the mammals belong. Having reached the very foot of the trunk we are in a position to review and summarize the evidences which we have discovered all about us as we have descended. The various examples we have mentioned and the groups to which they belong clearly occupy different places in the scale which begins with the protozoa and extends upward to the most complicated and differentiated animals. Hydra takes its place above the protozoa for obvious structural reasons; worms belong to a still higher zone, surpassed by the more complex jointed animals like crustacea and insects. Far above these are the vertebrates, among which we have already demonstrated the occurrence of different grades of organization, from the fish up to the higher amphibia and reptiles, and beyond in two directions to the diverging birds and mammals. The basic characteristics of every group in a high position may be traced back to some one or another of the divisions at a lower level, so that the general sequence of the structural levels from low to high becomes intelligible as the order of their evolution.

To my mind the rudimentary and vestigial structures of animals are in themselves proof positive of a natural history of change. The few illustrations can be reinforced by countless examples offered by every group of living animals. If such structures have not evolved naturally by degenerating from more efficient counterparts in ancestors of earlier times, and if they have been specially created, they are utterly meaningless and their very existence is unreasonable. If common sense is to be employed, they demonstrate evolution.

Everywhere throughout the whole series animals place themselves in a treelike arrangement, for in their respective levels they occur like leaves at the ends of the lines of descent which have led up to them and which are comparable to the branches and limbs arising from the trunk of a tree. Thus the major and minor divisions of animals do not follow in the order of the rungs of a ladder, even though they must be assigned to different levels according to the complexity of their construction. The summary given above, namely, that the occurrence of lower and higher levels reveals an order of evolution, is amplified and not contradicted by the statement that the species of animals are group in a treelike arrangement. It is the task of the evolutionist, provided with all the facts of comparative anatomy and dealing only with the various species as separate leaves, so to speak, to reconstruct the now invisible but not unreal twigs and branches and limbs of the animal tree, and to show how they have diverged at one time or another as they have grown and spread to produce the species of the present day. This he may do in so far as he may find sufficient materials to enable him to employ the methods of comparative anatomy and the great natural principle established by this method--that essential likeness means consanguinity.

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No evidence of evolution could be more significant and interesting than the results provided by the comparative study of development. In the first place it is an obvious fact that every living thing changes in the course of its life-history, and if as an adult it occupies a high place in the animal scale, its embryological transformation is more elaborate and intricate than in the case of a lower form. Every one knows that organisms do develop, and yet I believe that few appreciate the tremendous significance of the mere fact that this is true, while still fewer are aware that the peculiar and characteristic early stages through which an animal passes in becoming an adult are even more striking than the fact of development itself. We shall learn something of these earlier conditions in the development of some of our most familiar animals, but at the outset nothing can be more important than an appreciation of the first great lesson of this department of natural history--namely that organic transformation is real and natural. We do not need to employ the methods of formal logic to know that in growing up a human infant undergoes the changes of childhood and adolescence, that kittens become cats, and that an oak tree is produced by an acorn, for we know these things directly by observing them. It is natural for development to take place under normal conditions, and if it does not, then something has interfered with nature. Inasmuch as "growing up" is accomplished by the alteration of an organic mechanism with one structure into an individual with a changed plan of body, it is in essence the actual process of evolution which the comparative study of grown animals of to-day demonstrates in the way we have learned. The study of animal structure discovers the process of evolution because the most reasonable interpretation of the similarities and minor differences exhibited everywhere by the various groups of animals is that descent with adaptive and divergent modification has taken place; the result is reached by inference, it is true, but by scientific and logical inference. With development it is otherwise. No reasoning is necessary to tell us that organic transformation is a real and a natural process. We see it everywhere about us and we ourselves have come to be what we are by a natural history of change. Can we consistently deny that it is possible for a species to alter in the long course of time when a few brief weeks are sufficient for the new-laid egg of the fowl to develop into a fledgling? Many indeed strain at the gnat of the longer process in the past when without hesitation they recognize the real and obvious fact of individual development in a brief period.

I have said that development is a "natural" process. We employ this word for the familiar and everyday occurrence or thing; it does not imply that everything is known about the object or phenomenon, because science knows that complete and final knowledge is impossible. We say that it is natural for rain to fall to the earth, and we speak of the law of gravitation according to which this takes place as a natural principle, but it may not have occurred to many to inquire what makes rain fall and why do masses of matter everywhere behave toward one another in the consistent manner described by the law in question. Sunshine is natural, but we do not know why light travels as it does from the sun to the earth, and this is another question which, like the inquiry into the ultimate cause of the familiar and natural phenomenon of gravitation, has not yet been answered. But it is still regarded as natural for the rain to fall and for the sun to shine. In the same way does science view development, denoting it natural because it is an ordinary everyday matter. And we are under no more obligation to postulate supernatural control for the changing forms in the life-history of a chick or a cat than we need to assume that gravitation and the radiation of light demand immediate supernatural direction. The embryology of no form is fully understood or described or explained, but no intelligent person would be willing to assert that because complete knowledge is lacking, it is unnatural for organic transformation to take place during growth. Whatever may be the ultimate origin and nature of the directing powers behind gravitation and development and other phenomena, we have no concern with such matters because they cannot be handled by scientific methods and one belief about them is on the same plane with any other. Our task is to deal with the everyday phenomena of life and the production of living species.

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It is not necessary to go far afield to find an animal which will introduce us to the general principles of embryology. In the present instance as in the case of comparative anatomy almost any form will disclose the meaning of development, for animate nature is uniform and consistent in its methods of operation throughout its wide range. We shall begin with the familiar frog which every one knows is a product of a tadpole; passing on to the chick we will learn more facts that will enable us to formulate the main principle of comparative embryology in definite terms; we will then be prepared to extend our survey so as to include somewhat less familiar facts and animals that are even more significant than the first illustrations.

If we should visit a woodland pond in early spring, we would find somewhere among the leaves and sticks in the water large masses of a clear jellylike consistency enclosing hundreds of little black spheres about an eighth of an inch in diameter. These are the egg masses and eggs of a common frog. Watching them day by day we see the small one-celled egg spheres divide into more and more numerous portions which are the daughter-cells, destined to form by their products the many varied tissues and organs of the developing larva and adult frog. After three or four days the egg changes from its globular form into an oval or elliptical mass, and from one end of this a small knob projects to become a flattened waving tail a few days later. On the sides of the larger anterior portion shallow grooves make their appearance and soon break through from the throat or pharynx to the exterior as gill-slits. Shortly afterwards the little embryo wriggles out of its encasing coat of jelly, develops a mouth, and begins its independent existence as a small tadpole, with eyes, nasal and auditory organs, and all other parts that are necessary for a free life. Thus the one-celled egg has transformed into something that it was not at first, and in doing this it has proved the possibility and the reality of organic reconstruction.

The tadpole breathes by means of its gills, and it is at first entirely devoid of the lungs which the adult frog possesses and uses. When we speak of the larval respiratory organs as gills we imply that they are like the organs of a fish which have the same name; they are truly like those of fishes, for the blood-vessels which go to them are essentially the same as in the lower types and they are supported by simple skeletal rods like the gill-bars of the fish. In a word, they are the same things.

The animal feeds and grows during the months of its first summer, and hibernates the following winter; with the warmth of spring it revives and proceeds further along the course of its development. Near the base of the tail two minute legs grow out from the hinder part of the body, and while these are enlarging two front legs make their appearance a little behind the gills. The tadpole now rises more frequently to the surface where it takes small mouthfuls of air. Meanwhile great changes are effected inside the body where the various systems of fishlike organs become remodeled into amphibian structures. A sac is formed from the wall of the esophagus, and this enlarges and divides to form the two simple lungs. The legs increase in size, the tail dwindles more and more, the gills close up, and soon the animal hops out on land as a complete young frog. From this time on it breathes by means of its lungs instead of gills, even though it returns to the water to escape its foes, to seek its prey, and to hibernate in the mud of the lake bed during the winter months.

All these changes are familiar and natural, but until science places them and similar facts in their proper relations their significance is lost to us. The tadpole is essentially a fish in its general structure and mode of life, even though its heritage is such that it can develop into a higher animal. When it does become a frog it proves beyond a doubt that there is no impassable barrier between fishes and amphibia. Our earlier comparison of the structures of these two classes of vertebrates led to the conclusion that the latter had evolved from antecedents like the former, and had thus followed them upon the earth; now that sequence seems to have some connection with the method by which a tadpole, obviously not a fish but nevertheless actually fishlike, changes into a frog, a member of a higher class of vertebrates. This method is employed by developing frogs apparently because it follows the ancestral order of events, and because, so to speak, the only way a frog knows how to become a frog is to develop from an egg first into a fishlike tadpole and then to alter itself as its ancestors did during their evolution in the past. We begin to see, then, that in addition to the impressive fact of development itself, the mode of organic transformation is far more conclusive evidence of evolution, because it reveals an order of events which parallels the order established by comparative anatomy as the evolutionary sequence.

However it is well to review some of the changes by which a chick comes into existence before attempting to comprehend fully the fundamental principle of development that the tadpole's history discloses to us. The egg of a common fowl is certainly not a chick. Within the calcareous shell are two delicate membranes that enclose the white or albumen; within this, swung by two thickened cords of the albumen, is the yellow yolk ball enclosed by a proper membrane of its own. In the earliest condition, even before the albumen and the shell are added and before the egg is laid, on one side of the yolk-mass there is a tiny protoplasmic spot which is at first a single cell and nothing more. The hen's egg is relatively enormous, but nevertheless, like that of the frog, it starts upon its course of development as a single unitary biological element--a cell. During the earliest subsequent hours the first cell divides again and again to form a small disk upon the surface of the yolk. Soon the cells along the middle line of this small sheet become rearranged to make an obvious streak or band, and about this line a simple tube is constructed which is destined to become the future brain and spinal cord. The whole disk continues to enlarge by further division of its constituent elements so that it encloses more and more of the yolk mass, but the little chick itself is made out of the cells along the central line of the original plate, from which it folds at the sides and in front and behind so as to lie somewhat above and apart from the flatter enclosing cell layers which partly surround the yolk.

At the sides of the primitive nerve-tube small blocks of cells arise to develop into primitive muscles and other structures. As nourishment is brought to the embryo from the surrounding layers enclosing the nutrient yolk, one system after another takes its shape and builds its several parts into organs which can be recognized as elementary structures of a chick. Among the more interesting ones are small clefts or slits formed in the side walls of the rudimentary throat or pharynx. Blood-vessels go forward from the simple heart to run up through the intervening bars exactly as in the tadpole and the fish. In brief, the young chick possesses a series of gill-slits, for these structures are the same in essential plan and relations as the clefts of tadpoles and fishes. Does this mean that even birds have descended from gill-breathing ancestors? Science answers in the affirmative, because evolution gives the only reasonable explanation of such facts as these. The case seems different from that of the frog, because gills are used by the tadpole, but gill-slits and gill-bars can have no conceivable value for the chick as organs concerned with the purification of the blood. None the less, if the transition from a gilled tadpole to the adult with lungs means an evolution of amphibia from fishlike ancestors, then the change of a chick embryo with gill-clefts into the fledgling without them is most reasonably interpreted as proof that birds as well as amphibia have had ancestors as simple as fishes.

As development progresses four small pads make their appearance; two of these lie on either side of the body back of the head and the other two arise near the posterior end. They are far from being wings and legs, but as day follows day they become molded into somewhat similar limbs, as much alike in general plan as the four legs of a lizard; subsequently the ones at the front change into real wings and the hinder ones become legs. Meanwhile the internal organs slowly transform from fishlike structures into things that display the characteristics of reptilian counterparts, and only later do they become truly avian. Last of all the finishing touches are made, and the whole creature becomes a particular kind of a bird which picks its way out of the shell and shifts for itself as a chick.

Only a few of the countless details have been mentioned which demonstrate the resemblance of the successive stages first to fishes, and later to amphibia and reptiles. We have a wide choice of materials, but even the foregoing brief list of illustrations shows that the order in which the stages follow is the one which comparative anatomy independently proves to be the order of the evolution of fishes, amphibia, reptiles, and birds. Why, now, should it be necessary for a developing bird to follow this order? The answer has been found in the immense array of embryological facts that investigators have verified and classified, that all tell the same story. It is, that birds have arisen by evolution from ancestors which were really as simple as the members of these lower classes. It seems then that the only way a bird of to-day can become itself is to traverse the path along which its progenitors had progressed in evolution. Stating its conclusions precisely, science formulates the principle in the following words: individual development is a brief resume of the history of the species in past times, or, more technically, ontogeny recapitulates phylogeny. To be sure, the full history is not reviewed in detail, for the chick embryo does not actually swim in water and breathe by means of gills. Only a condensed account of evolution of its kind is presented by an embryo during its development; as Huxley and Haeckel have put it, whole lines and paragraphs and even pages are left out; many false passages of a later date are inserted as the result of peculiar larval and embryonic needs and adjustments. But in its major statements and as a general outline, the account is a trustworthy natural document submitted as evidence that higher species of to-day have evolved from ancestors which must have been like some of the present lower animals.

Coming now to the mammalia, it might seem that we have reached forms so highly developed that they would not exhibit the same kind of developmental history, but would have their own mode of growing up. This is not so, for like the adult fish, the larval tadpole, and the embryo chick, an embryo of a cat or a man is at one time constructed with a series of gill-clefts and with blood-vessels and skeletal supports of fishlike nature that are everywhere associated with gills. The embryos of wildcats and dogs, rabbits and rats, pigs, deer, and sheep, and of all other mammalia, possess similar structures. Thus they all pass through a stage which is found also in the development of reptiles, birds, and amphibia,--a stage which corresponds to the fish throughout its life. Unless these facts mean that the great classes of vertebrates have originated together from the same or closely similar ancestors, they are unintelligible; for we cannot see why a cat or a chick should have to be essentially fishlike at any time unless this is so. Comparative anatomy states as we have learned that the amphibia as a class have evolved from and have out-developed the fishes, that reptiles have progressed still higher, and that birds and mammals have originated from reptilian ancestors along roads that have diverged beyond the immediate parent class. Because the members of each class have to pass along the same path trodden by their many varied ancestors, although at express speed, as it were, the similarity of the earliest stages in their development is explained, for during these periods they are traversing a path over which their ancestors passed together.

The places where the developing embryos depart from the common mode show where the several divisions took leave of one another in their evolution,--a point that comes out with great clearness when the facts of mammalian development are broadly compared. The embryos of carnivora and rodents and hoofed animals are alike in their earlier development, and their agreement means a community of origin. At a certain point the cat and dog depart from the common mode, but they remain alike up to a far later stage than the one in which they are similar to the embryos of rats and sheep. The rat and squirrel and rabbit, on their part, remain together until long after they take leave of the carnivora and ungulates; while the sheep and cattle and pigs have their own branch line, which they follow in company after leaving the embryos of the other orders. The reasons for these facts seem to be that the members of the three orders exemplified have evolved from the same stock, which accounts for their embryonic similarity for a long time after they collectively come to differ from amphibia and reptiles, while the members in each order became differentiated only later, wherefore their embryonic paths coincide for a longer period. Thus the degree of adult resemblance which indicates the closeness of relationship corresponds with the degree of embryonic agreement; that is, the cat and dog are much alike and their modes of development are essentially the same to the latest stages, while the cat and horse agree only during the earliest and middle stages, and their lines diverge before those of the cat and dog on the one hand, or those of the horse and pig on the other.

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