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Fungi: Their Nature and Uses · M. C. Cooke — chapter 5 of 45 · ~6,587 words · public domain

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HYPHOMYCETES.--The predominant feature in the structure of this order has already been intimated to consist in the development of the vegetative system under the form of simple or branched threads, on which the fruit is generated. The common name of mould is applied to them perhaps more generally than to other groups, although the term is too vague, and has been too vaguely applied to be of much service in giving an idea of the characteristics of this order. Leaving the smaller groups, and confining ourselves to the Dematiei and the Mucedines, we shall obtain some notion of the prevalent structure. In the former the threads are more or less carbonized, in the latter nearly colourless. One of the largest genera in Dematiei is Helminthosporium. It appears on decaying herbaceous plants, and on old wood, forming effused black velvety patches. The mycelium, of coloured jointed threads, overlays and penetrates the matrix; from this arise erect, rigid, and usually jointed threads, of a dark brown, nearly black colour at the base, but paler towards the apex. In most cases these threads have an externally cortical layer, which imparts rigidity; usually from the apex, but sometimes laterally, the spores are produced. Although sometimes colourless, these are most commonly of some shade of brown, more or less elongated, and divided transversely by few or many septa. In Helminthosporium Smithii, the spores much exceed the dimensions of the threads; in other species they are smaller. In Dendryphium, the threads and spores are very similar, except that the threads are branched at their apex, and the spores are often produced one at the end of another in a short chain. In Septosporium again, the threads and spores are similar, but the spores are pedicellate, and attached at or near the base; whilst in Acrothecium, with similar threads and spores, the latter are clustered together at the apex of the threads. In Triposporium, the threads are similar, but the spores are tri-radiate; and in Helicoma, the spores are twisted spirally. Thus, we might pass through all the genera to illustrate this chief feature of coloured, septate, rather rigid, and mostly erect threads, bearing at some point spores, which in most instances are elongated, coloured, and septate.

MUCEDINES.--Here, on the other hand, the threads, if coloured at all, are still delicate, more flexuous, with much thinner walls, and never invested with an external cortical layer. One of the most important and highly developed genera is Peronospora, the members of which are parasitic upon and destructive of living vegetables. It is to this genus that the mould of the too famous potato disease belongs. Professor De Bary has done more than any other mycologist in the investigation and elucidation of this genus; and his monograph is a masterpiece in its way. He was, however, preceded by Mr. Berkeley, and more especially by Dr. Montagne, by many years in elucidation of the structure of the flocci and conidia in a number of species. In this genus, there is a delicate mycelium, which penetrates the intercellular passages of living plants, giving rise to erect branched threads, which bear at the tips of their ultimate ramuli, sub-globose, ovate, or elliptic spores, or, as De Bary terms them--conidia. Deeply seated on the mycelium, within the substance of the foster plant, other reproductive bodies, called oogonia, originate. These are spherical, more or less warted and brownish, the contents of which become differentiated into vivacious zoospores, capable, when expelled, of moving in water by the aid of vibratile cilia. A similar structure has already been indicated in Cystopus, otherwise it is rare in fungi, if the Saprolegniei be excluded. In Botrytis and in Polyactis, the flocci and spores are similar, but the branches of the threads are shorter and more compact, and the septa are more common and numerous; the oogonia also are absent. De Bary has selected Polyactis cinerea, as it occurs on dead vine leaves, to illustrate his views of the dualism which he believes himself to have discovered in this species. "It spreads its mycelium in the tissue which is becoming brown," he writes, "and this shows at first essentially the same construction and growth as that of the mycelium filaments of Aspergillus." On the mycelium soon appear, besides those which are spread over the tissue of the leaves, strong, thick, mostly fasciculate branches, which stand close to one another, breaking forth from the leaf and rising up perpendicularly, the conidia-bearers. They grow about 1 mm. long, divide themselves, by successively rising partitions, into some prominent cylindrical linked cells, and then their growth is ended, and the upper cell produces near its point three to six branches almost standing rectangularly. Of these the under ones are the longest, and they again shoot forth from under their ends one or more still shorter little branches. The nearer they are to the top, the shorter are the branches, and less divided; the upper ones are quite branchless, and their length scarcely exceeds the breadth of the principal stem. Thus a system of branches appears, upon which, on a small scale, a bunch of grapes is represented. All the twigs soon end their growth; they all separate their inner space from the principal stem, by means of a cross partition placed close to it. All the ends, and also that of the principal stem, swell about the same time something like a bladder, and on the upper free half of each swelling appear again, simultaneously, several fine protuberances, close together, which quickly grow to little oval bladders filled with protoplasm, and resting on their bearers with a sub-sessile, pedicellate, narrow basis, and which at length separate themselves through a partition as in Aspergillus. The detached cells are the conidia of our fungus; only one is formed on each stalk. When the formation is completed in the whole of the panicle, the little branches which compose it are deprived of their protoplasm in favour of the conidia; it is the same with the under end of the principal stem, the limits of which are marked by a cross partition. The delicate wall of these parts shrinks up until it is unrecognizable; all the conidia of the panicle approach one another to form an irregular grape-like bunch, which rests loosely on the bearer, and from which it easily falls away as dust. If they be brought into water they fall off immediately; only the empty, shrivelled, delicate skins are to be found on the branch which bore them, and the places on which they are fixed to the principal stem clearly appear as round circumscribed hilums, generally rather arched towards the exterior. The development of the main stem is not ended here. It remains solid and filled with protoplasm as far as the portion which forms the end through its conidia. Its end, which is to be found among these pieces, becomes pointed after the ripening of the first panicle, pushes the end of the shrivelled member on one side, and grows to the same length as the height of one or two panicles, and then remains still, to form a second panicle similar to the first. This is later equally perfoliated as the first, then a third follows, and thus a large number of panicles are produced after and over one another on the same stem. In perfect specimens, every perfoliated panicle hangs loosely to its original place on the surface of the stem, until by shaking or the access of water to it, it falls immediately into the single conidia, or the remains of branches, and the already-mentioned oval hilums are left behind. Naturally, the stem becomes longer by every perfoliation; in luxuriant specimens the length can reach that of some lines. Its partition is already, by the ripening of the first panicle from the beginning of its foundation, strong and brown; it is only colourless at the end which is extending, and in all new formations. During all these changes the filament remains either unbranched, except as regards the transient panicles, or it sends out here and there, at the perfoliated spots, especially from the lower ones, one or two strong branches, standing opposite one another and resembling the principal stem.

The mycelium, which grows so exuberantly in the leaf, often brings forth many other productions, which are called sclerotia, and are, according to their nature, a thick bulbous tissue of mycelium filaments. Their formation begins with the profuse ramification of the mycelium threads in some place or other; generally, but not always, in the veins of the leaf; the intertwining twigs form an uninterrupted cavity, in which is often enclosed the shrivelling tissue of the leaf. The whole body swells to a greater thickness than that of the leaf, and protrudes on the surface like a thickened spot. Its form varies from circular to fusiform; its size is also very unequal, ranging between a few lines and about half a millimetre in its largest diameter. At first it is colourless, but afterwards its outer layers of cells become round, of a brown or black colour, and it is surrounded by a black rind, consisting of round cells, which separate it from the neighbouring tissue. The tissue within the rind remains colourless; it is an entangled uninterrupted tissue of fungus filaments, which gradually obtain very solid, hard, cartilaginous coats. The sclerotium, which ripens as the rind becomes black, loosens itself easily from the place of its formation, and remains preserved after the latter is decayed.

The sclerotia are, here as in many other fungi, biennial organs, designed to begin a new vegetation after a state of apparent quietude, and to send forth special fruit-bearers. They may in this respect be compared to the bulbs and perennial roots of under shrubs. The usual time for the development of the sclerotia is late in the autumn, after the fall of the vine leaves. As long as the frost does not set in, new ones continually spring up, and each one attains to ripeness in a few days. If frost appears, it can lie dry a whole year, without losing its power of development. This latter commences when the sclerotium is brought into contact with damp ground during the usual temperature of our warmer seasons. If this occur soon, at the latest some weeks after it is ripe, new vegetation grows very quickly, generally after a few days; in several parts the colourless filaments of the inner tissue begin to send out clusters of strong branches, which, breaking through the black rind, stretch themselves up perpendicularly towards the surface, separate from one another, and then take all the characteristics of the conidia-bearers. Many such clusters can be produced on one sclerotium, so that soon the greater part of the surface is covered by filamentous conidia-bearers with their panicles. The colourless tissue of the sclerotium disappears in the same degree as the conidia-bearers grow, and at last the black rind remains behind empty and shrivelled. If we bring, after many months, for the first time, the ripe sclerotium, in damp ground, in summer or autumn, after it has ripened, the further development takes place more slowly, and in an essentially different form. It is true that from the inner tissue numerous filamentous branches shoot forth at the cost of this growing fascicle, and break through the black rind, but its filaments remain strongly bound, in an almost parallel situation, to a cylindrical cord, which for a time lengthens itself and spreads out its free end to a flat plate-like disc. This is always formed of strongly united threads, ramifications of the cylindrical cord. On the free upper surface of the disc, the filaments shoot forth innumerable branches, which, growing to the same height, thick and parallel with one another, cover the before-named disc. Some remain narrow and cylindrical, are very numerous, and produce fine hairs (paraphyses); others, also very numerous, take the form of club-like ampulla cells, and each one forms in its interior eight free swimming oval spores. Those ampulla cells are sporidiiferous asci. After the spores have become ripe, the free point of the utricle bursts, and the spores are scattered to a great distance by a mechanism which we will not here further describe. New ampullas push themselves between those which are ripening and withering; a disc can, under favourable circumstances, always form new asci for weeks at a time. The number of the already described utricle-bearers is different, according to the size of the sclerotium; smaller specimens usually produce only one, larger two to four. The size is regulated by that of the sclerotia, and ranges, in full-grown specimens, between one and more millimetres for the length of the stalk, and a half to three (seldom more) millimetres for the breadth of the disc. For some time the conidia form, belonging to the Mucedines, has been known as Botrytis cinerea (or Polyactis cinerea). The compact mycelium, or sclerotium, as an imperfect fungus, bore the name of Sclerotium echinatum, whilst to the perfect and cup-like form has been given the name of Peziza Fuckeliana. We have reproduced De Bary's life-history of this mould here, as an illustration of structure in the Mucedines, but hereafter we shall have to write of similar transformations when treating of polymorphism.

The form of the threads, and the form and disposition of the spores, vary according to the genera of which this order is composed. In Oidium the mostly simple threads break up into joints. Many of the former species are now recognized as conditions of Erysiphe. In Aspergillus, the threads are simple and erect, with a globose head, around which are clustered chains of simple spores. In Penicillium, the lower portion of the threads is simple, but they are shortly branched at the apex, the branches being terminated by necklaces of minute spores. In Dactylium, the threads are branched, but the spores are collected in clusters usually, and are moreover septate. In other genera similar distinctions prevail. These two groups of black moulds and white moulds are the noblest, and contain the largest number of genera and species amongst the Hyphomycetes. There is, however, the small group of Isariacei, in which the threads are compacted, and a semblance of such hymenomycetal forms as Clavaria and Pterula is the result, but it is doubtful if this group contains many autonomous species. In another small group, the Stilbacei, there is a composite character in the head, or receptacle, and in the stem when the latter is present. Many of these, again, as Tubercularia, Volutella, Fusarium, &c., contain doubtful species. In Sepedoniei and Trichodermacei, the threads are reduced to a minimum, and the spores are such a distinctive element that through these groups the Hyphomycetes are linked with the Coniomycetes. These groups, however, are not of sufficient size or importance to demand from us, in a work of this character, anything more than the passing allusion which we have given to them.

We come now to consider the structure in the Sporidiifera, in which the fructifying corpuscles or germs, whether called spores or sporidia, are generated within certain privileged cysts, usually in definite numbers. In systematic works, these are included under two orders, the Physomycetes and the Ascomycetes. The former of these consists of cyst-bearing moulds, and from their nearest affinity to the foregoing will occupy the first place.

PHYSOMYCETES include, especially amongst the Mucorini, many most interesting and instructive species for study, which even very lately have occupied the attention of continental mycologists. Most of these phenomena are associated more or less with reproduction, and as such will have to be adverted to again, but there are points in the structure which can best be alluded to here. Again taking Professor de Bary's researches as our guide, we will illustrate this by the common Mucor mucedo: If we bring quite fresh horse-dung into a damp confined atmosphere, for example, under a bell-glass, there appears on its surface, after a few days, an immense white mildew. Upright strong filaments of the breadth of a hair raise themselves over the surface, each of them soon shows at its point a round little head, which gradually becomes black, and a closer examination shows us that in all principal points it perfectly agrees with the sporangia of other species. Each of these white filaments is a sporangia-bearer. They spring from a mycelium which is spread in the dung, and appear singly upon it. Certain peculiarities in the form of the sporangium, and the little long cylindrical spores, which, when examined separately, are quite flat and colourless, are characteristic of the species. If the latter be sown in a suitable medium, for example, in a solution of sugar, they swell, and shoot forth germinating utricles, which quickly grow to mycelia, which bear sporangia. This is easily produced on the most various organic bodies, and Mucor mucedo is therefore found spontaneously on every substratum which is capable of nourishing mildew, but on the above-named the most perfect and exuberant specimens are generally to be found. The sporangia-bearers are at first always branchless and without partitions. After the sporangium is ripe, cross partitions in irregular order and number often appear in the inner space, and on the upper surface branches of different number and size, each of which forms a sporangium at its point. The sporangia which are formed later are often very similar, but sometimes very different, to those which first appeared, because their partition is very thick and does not fall to pieces when it is ripe, but irregularly breaks off, or remains entire, enclosing the spores, and at last falls to the ground, when the fungus withers. The cross partition which separates the sporangia from its bearers is in those which are first formed (which are always relatively thicker sporangia) very strongly convex, while those which follow later are often smaller, and in little weak specimens much less arched, and sometimes quite straight. After a few days, similar filaments generally show themselves on the dung between the sporangia-bearers, which appear to the naked eye to be provided with delicate white frills. Where such an one is to be found, two to four rectangular expanding little branches spring up to the same height round the filament. Each of these, after a short and simple process, branch out into a furcated form; the furcations being made in such a manner that the ends of the branch at last so stand together that their surface forms a ball. Finally, each of the ends of a branch swells to a little round sporangium, which is limited by a partition (called sporangiolum, to distinguish it from the larger ones), in which some, generally four, spores are formed in the manner already known. When the sporangiola are alone, they have such a peculiar appearance, with their richly-branched bearers, that they can be taken for something quite different to the organs of the Mucor mucedo, and were formerly not considered to belong to it. That they really belong to the Mucor is shown by the principal filament which it bears, not always, but very often, ending with a large sporangium, which is characteristic of the Mucor mucedo; it is still more evident if we sow the spores of the sporangiolum, for, as it germinates, a mycelium is developed, which, near a simple bearer, can form large sporangia, and those form sporangiola, the first always considerably preponderating in number, and very often exclusively. If we examine a large number of specimens, we find every possible middle form between the simple or less branched sporangia-bearers and the typical sporangiola frills; and we arrive at last at the conclusion simply to place the latter among the varieties of form which the sporangia-bearer of the Mucor mucedo shows, like every other typical organic form within certain limits. On the other hand, propagation organs, differing from those of the sporangia and their products, belong to Mucor mucedo, which may be termed conidia. On the dung (they are rare on any other substance) these appear at the same time, or generally somewhat later, than the sporangia-bearers, and are not unlike those to the naked eye. In a more accurate examination, they appear different; a thicker, partition-less filament rises up and divides itself, generally three-forked, at the length of one millimetre, into several series of branchlets. The forked branches of the last series bear under their points, which are mostly capillary, short erect little ramuli, and these, with which the ends of the principal branches articulate on their somewhat broad tops, several spores and conidia, near one another; about fifteen to twenty are formed at the end of each little ramulus. The peculiarities and variations which so often appear in the ramification need not be discussed here. After the articulation of the conidia, their bearers sink together by degrees, and are quite destroyed. The ripe conidia are round like a ball, their surface is scarcely coloured, and almost wholly smooth. These conidioid forms were at first described as a separate species under the name of Botrytis Jonesii. How, then, do they belong to the Mucor? That they appear gregariously is as little proof of an original relation to one another, here as elsewhere. Attempts to prove that the conidia and sporangia-bearers originate on one and the same mycelium filament may possibly hereafter succeed. Till now this has not been the case, and he who has ever tried to disentangle the mass of filaments which exuberantly covers the substratum of a Mucor vegetation, which has reached so far as to form conidia, will not be surprised that all attempts have hitherto proved abortive. The suspicion of the connection founded on the gregariously springing up, and external resemblance, is fully justified, if we sow the conidia in a suitable medium, for example, in a solution of sugar. They here germinate and produce a mycelium which exactly resembles that of the Mucor mucedo, and, above all, they produce in profusion the typical sporangia of the same on its bearers. The latter are till now alone reproductions of conidia-bearers, and have never been observed on mycelia which have grown out of conidia.

These phenomena of development appear in the Mucor when it dwells on a damp substance, which must naturally contain the necessary nourishment for it, and is exposed to the atmospheric air. Its mycelium represents at first strong branched utricles without partitions; the branches are of the higher order, mostly divided into rich and very fine-pointed ramuli. In old mycelium, and also in the sporangia-bearers, the contents of which are mostly used for the formation of spores, and the substratum of which is exhausted for our fungus, short stationary pieces, filled with protoplasm, are very often formed into cells through partitions in order to produce spores, that is, grow to a new fruitful mycelium. These cells are called gemmules, brooding cells, and resemble such vegetable buds and sprouts of foliaceous plants which remain capable of development after the organs of vegetation are dead, in order to grow, under suitable circumstances, to new vegetating plants, as, for example, the bulbs of onions, &c.

If we bring a vegetating mycelium of Mucor mucedo into a medium which contains the necessary nourishment for it, but excluded from the free air, the formation of sporangia takes place very sparingly or not at all, but that of gemmules is very abundant. Single interstitial pieces of the ramuli, or even whole systems of branches, are quite filled with a rich greasy protoplasm; the short pieces and ends are bound by partitions which form particular, often tun-like or globular cells; the longer ones are changed, through the formation of cross partitions, into chains of similar cells; the latter often attain by degrees strong, thick walls, and their greasy contents often pass into innumerable drops of a very regular globular form and of equal size. Similar appearances show themselves after the sowing of spores, which are capable of germinating in the medium already described, from which the air is excluded. Either short germinating utricles shoot forth, which soon form themselves into rows of gemmules, or the spores swell to large round bladders filled with protoplasm, and shoot forth on various parts of their surface innumerable protuberances, which, fixing themselves with a narrow basis, soon become round vesiculate cells, and on which the same sprouts which caused their production are repeated, formations which remind us of the fungus of fermentation called globular yeast. Among all the known forms of gemmules we find a variety which are intermediate, all of which show, when brought into a normal condition of development, the same proportion, and the same germination, as those we first described.

We have detailed rather at length the structure and development of one of the most common of the Mucors, which will serve as an illustration of the order. Other distinctions there may be which are of more interest as defining the limits of genera, except such as may be noticed when we come to write more specially of reproduction.

ASCOMYCETES.--Passing now to the Ascomycetes, which are especially rich in genera and species, we must first, and but superficially, allude to Tuberacei, an order of sporidiiferous fungi of subterranean habit, and rather peculiar structure. In this order an external stratum of cells forms a kind of perithecium, which is more or less developed in different genera. This encloses the hymenium, which is sinuous, contorted, and twisted, often forming lacunæ. The hymenium in some genera consists of elongated, nearly cylindrical asci, enclosing a definite number of sporidia; in the true truffles and their immediate allies, the asci are broad sacs, containing very large and beautiful, often coloured, sporidia. These latter have either a smooth, warted, spinulose, or lacunose epispore, and, as will be seen from the figures in Tulasne's Monograph, or those in the last volume of Corda's great work, are attractive microscopical objects. In some cases, it is not difficult to detect paraphyses, but in others they would seem to be entirely absent. A comparatively large number have been discovered and recorded in Great Britain, but of those none are more suitable for study of general structure than the ordinary truffle of the markets.

The structure of the remaining Ascomycetes can be studied under two groups, i.e., the fleshy Ascomycetes, or, as they have been termed, the Discomycetes, and the hard, or carbonaceous Ascomycetes, sometimes called the Pyrenomycetes. Neither of these names gives an accurate idea of the distinctions between the two groups, in the former of which the discoid form is not universal, and the latter contains somewhat fleshy forms. But in the Discomycetes the hymenium soon becomes more or less exposed, and in the latter it is enclosed in a perithecium. The Discomycetes are of two kinds, the pileate and the cup-shaped. Of the pileate such a genus as Gyromitra or Helvella is, in a certain sense, analogous to the Agarics amongst Hymenomycetes, with a superior instead of an inferior hymenium, and enclosed, not naked, spores. Again, Geoglossum is somewhat analogous to Clavaria. Amongst the cup-shaped, Peziza is an Ascomycetous Cyphella. But these are perhaps more fanciful than real analogies.

Recently Boudier has examined one group of the cup-shaped Discomycetes, the Ascobolei, and, by making a somewhat free use of his Memoir, we may arrive at a general idea of the structure in the cupulate Discomycetes. They present themselves at first under the form of a small rounded globule, and almost entirely cellular. This small globule, the commencement of the receptacle, is not long in increasing, preserving its rounded form up to the development of the asci. At this period, under the influence of the rapid growth of these organs, it soon produces at its summit a fissure of the external membrane, which becomes a more marked depression in the marginate species. The receptacle thus formed increases rapidly, becomes plane, more convex, or more or less undulated at the margin, if at all of large size. Fixed to the place where it is generated by some more or less abundant mycelioid filaments, the receptacle becomes somewhat cup-shaped and either stipitate or sessile, composed of the receptacle proper and the hymenium.

The receptacle proper comprehends the subhymenial tissue, the parenchyma, and the external membrane. The subhymenial tissue is composed of small compact cells, forming generally a more coloured and dense stratum, the superior cells of which give rise to the asci and paraphyses. The parenchyma is seated beneath this, and is generally of interlaced filaments, of a looser consistency than the preceding, united by intermediate cellules. The external membrane, which envelopes the parenchyma, and limits the hymenium, differs from the preceding by the cells often being polyhedric, sometimes transverse, and united together, and sometimes separable. Externally it is sometimes smooth, and sometimes granular or hairy.

The hymenium is, however, the most, important part, consisting of (1) the paraphyses, (2) the asci, and sometimes (3) an investing mucilage. The asci are always present, the paraphyses are sometimes rare, and the mucilage in many cases seems to be entirely wanting.

The paraphyses, which are formed at the first commencement of the receptacle, are at first very short, but soon elongate, and become wholly developed before the appearance of the asci. They are linear, sometimes branched and sometimes simple, often more or less thickened at their tips; almost always they contain within them some oleaginous granules, either coloured or colourless. Their special function seems still somewhat obscure, and Boudier suggests that they may be excitatory organs for the dehiscence of the asci. However this may be, some mycologists are of opinion that, at least in some of the Ascomycetes, the paraphyses are abortive asci, or, at any rate, that abortive asci mixed with the paraphyses cannot be distinguished from them.

The mucilage forms itself almost at the same time as the paraphyses, and previous to the formation of the asci. This substance appears as a colourless or yellowish mucilage, which envelopes the paraphyses and asci, and so covers the hymenium with a shining coat.

The asci appear first at the base of the paraphyses, under the form of oblong cells, filled with colourless protoplasm. By rapid growth, they soon attain a considerable size and fulness, the protoplasm being gradually absorbed by the sporidia, the first indication of which is always the central nucleus. The mucilage also partly disappears, and the asci, attaining their maturity, become quite distinct, each enclosing its sporidia. But before they take their complete growth they detach themselves from the subhymenial tissue, and being attenuated towards their base, are forced upwards by pressure of the younger asci, to, and in some instances beyond, the upper surface of the disc. This phenomenon commences during the night, and continues during the night and all the morning. It attains its height at mid-day, and it is then that the slightest breath of air, the slightest movement, suffices to cause dehiscence, which is generally followed by a scarcely perceptible contractile motion of the receptacle.

There is manifestly a succession in formation and maturity of the asci in a receptacle. In the true Ascobolei, in which the sporidia are coloured, this may be more distinctly seen. At first some thin projecting points appear upon the disc, the next day they are more numerous, and become more and more so on following days, so as to render the disc almost covered with raised black or crystalline points; these afterwards diminish day by day, until they ultimately cease. The asci, after separation from the subhymenial tissue, continue to lengthen, or it may be that their elasticity permits of extension, during expulsion. Boudier considers that an amount of elasticity is certain, because he has seen an ascus arrive at maturity, eject its spores, and then make a sharp and considerable movement of retraction, then the ascus returned again, immediately towards its previous limits, always with a reduction in the number of its contained sporidia.

The dehiscence of the asci takes place in the Ascobolei, in some species of Peziza, Morchella, Helvella, and Verpa, by means of an apical operculum, and in other Pezizæ, Helotium, Geoglossum, Leotia, Mitrula, &c., by a fissure of the ascus. This operculum may be the more readily seen when the ascus is coloured by a drop of tincture of iodine.

The sporidia are usually four or eight, or some multiple of that number, in each ascus, rarely four, most commonly eight. At a fixed time the protoplasm, which at first filled the asci, disappears or is absorbed in a mucilaginous matter, which occupies its place, in the midst of which is a small nucleus, which is the rudiment of the first spore; other spores are formed consecutively, and then the substance separates into as many sections as there are sporidia. From this period each sporidium seems to have a separate existence. All have a nucleus, which is scarcely visible, often slightly granular, but which is quite distinct from the oleaginous sporidioles so frequent amongst the Discomycetes, and which are sometimes called by the same name. The sporidia are at first a little smaller than when mature, and are surrounded by mucilage. After this period the sporidia lose their nebulous granulations, whilst still preserving their nucleus; their outlines are distinct, and, amongst the true Ascobolei, commence acquiring a rosy colour, the first intimation of maturity. This colour manifests itself rapidly, accumulating exclusively upon the epispore, which becomes of a deep rose, then violet, and finally violet blue, so deep as sometimes to appear quite black. There are some modifications in this coloration, since, in some species, it passes from a vinous red to grey, then to black, or from rose-violet to brown.

The epispore acquires a waxy consistence by this pigmentation, so that it may be detached in granules. It is to this particular consistency of the epispore that the cracks so frequent in the coloured sporidia of Ascobolus are due, through contraction of the epispore. As they approach maturity, the sporidia accumulate towards the apex of the asci, and finally escape in the manner already indicated.

In all essential particulars there is a great similarity in the structure of the other Discomycetes, especially in their reproductive system. In most of them coloured sporidia are rare. In some the receptacle is pileate, clavate, or inflated, whilst in Stictis it is very much reduced, and in the lowest form of all, Ascomyces, it is entirely absent. In the Phacidiacei, the structure is very similar to that of the Elvellacei, whilst the Hysteriacei, with greater affinities with the latter, still tend towards the Pyrenomycetes by the more horny nature of the receptacle, and the greater tendency of the hymenium to remain closed, at least when dry. In some species of Hysterium, the sporidia are remarkably fine. M. Duby has subjected this group to examination, and M. Tulasne partly so.

SPHÆRIACEI.--In this group there is considerable variation, within certain limits. It contains an immense number of species, and these are daily being augmented. The general feature in all is the presence of a perithecium, which contains and encloses the hymenium, and at length opening by a pore or ostiolum at the apex. In some the perithecia are simple, in others compound; in some immersed in a stroma, in others free; in some fleshy or waxy, in others carbonaceous, and in others membranaceous. But in all there is this important difference from the Ascomycetes we have already had under consideration, that the hymenium is never exposed. The perithecium consists usually of an external layer of cellular structure, which is either smooth or hairy, usually blackish, and an internal stratum of less compact cells, which give rise to the hymenium.

As in the Discomycetes, the hymenium consists of asci, paraphyses, and mucilage, but the whole forms a less compact and more gelatinous mass within the perithecium. The formation and growth of the asci and sporidia differ little from what we have described, and when mature the asci dehisce, and the sporidia alone are ejected from the ostiolum. We are not aware that operculate asci have yet been detected. It has been shown in some instances, and suspected in others, that certain moulds, formerly classed with Mucedines and Dematiei, especially in the genus Helminthosporium, bear the conidia of species of Sphæria, so that this may be regarded as one form of fruit.

Perithecia, very similar externally to those of Sphæria, but containing spores borne on slender pedicels and not enclosed in asci, have had their relations to certain species of Sphæria indicated, and these are no longer regarded so much as species of Hendersonia or Diplodia as the pycnidia of Sphæria. Other and more minute perithecia, containing minute, slender stylospores in great numbers, formerly classed with Aposphæria, Phoma, &c., but are now recognized as spermogonia containing the spermatia of Sphæriæ. How these influence each other, when and under what circumstances the spermatia are instrumental in impregnation of the sporidia, is still matter of mystery. It is clear, however, that in all these conidia, macrospores, microspores, and some spermatia, or by whatever names they may be called, there exists a power of germination. Tulasne has indicated in some instances five or six forms of fruit as belonging to one fungus, of which the highest and most perfect condition is a species of Sphæria.

PERISPORIACEI.--Except in the perithecia rupturing irregularly, and not dehiscing by a pore, some of the genera in this group differ little in structure from the Sphæriacei. On the other hand, the Erysiphei present important and very interesting features. They occur chiefly on the green parts of growing plants. At first there is a more or less profuse white mycelium. This gives rise to chains of conidia (Oidium), and afterwards small sphæroid projections appear at certain points on the mycelium. These enlarge, take an orange colour, ultimately passing into brown, and then nearly black. Externally these perithecia are usually furnished with long, spreading, intertwined, or branching appendages, sometimes beautifully branched or hooked at their tips. In the interior of the receptacles, pear-shaped or ovate asci are formed in clusters, attached together at the base, and containing two or more hyaline sporidia. Other forms of fruit have also been observed on the same mycelium. In an exotic genus, Meliola, the fulcra, or appendages, as well as the mycelium, are black, otherwise it is very analogous to such a genus of Erysiphei as Microsphæria. In Chætomium, the perithecia bristle with rigid, dark-coloured hairs, and the sporidia are coloured. Our limits, however, will not permit of further elucidation of the complex and varied structure to be found amongst fungi.

A curious case occurred some years since at Bury St. Edmunds, which may be mentioned here in connection with the development of these nodules. Two children had died under suspicious circumstances, and an examination of the body of the latter after exhumation was made, a report having arisen that the child died after eating mushrooms. As certain white nodules appeared on the inner surface of the intestines, it was at once hastily concluded that the spores of the mushroom had germinated, and that the nodules were infant mushrooms. This appeared to one of us so strange, that application was made for specimens, which were kindly forwarded, and a cursory glance was enough to convince us that they were not fungoid. An examination under the microscope further confirmed the diagnosis, and the application of nitric acid showed that the nodules were merely due to chalk mixture, which had been given to the child for the diarrhetic symptoms under which he succumbed.

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