Part 22

A good illustration of dimorphism in one of the commonest of moulds is given by De Bary in a paper from which we have already quoted. He writes thus:--In every household there is a frequent unbidden guest, which appears particularly on preserved fruits, viz., the mould which is called Aspergillus glaucus. It shows itself to the naked eye as a woolly floccy crust over the substance, first purely white, then gradually covered with little fine glaucous, or dark green dusty heads. More minute microscopical examination shows that the fungus consists of richly ramified fine filaments, which are partly disseminated in the substratum, and partly raised obliquely over it. They have a cylindrical form with rounded ends, and are divided into long outstretched members, each of which possesses the property which legitimatizes it as a vesicle in the ordinary sense of the word; it contains, enclosed within a delicate structureless wall, those bodies which bear the appearance of a finely granulated mucous substance, which is designated by the name of protoplasm, and which either equally fills the cells, or the older the cell the more it is filled with watery cavities called vacuoles.

All parts are at first colourless. The increase in the length of the filaments takes place through the preponderating growth near their points; these continually push forward, and, at a short distance from them, successive new partitions rise up, but at a greater distance, the growth in the length ceases. This kind of growth is called point growth. The twigs and branches spring up as lateral dilatations of the principal filament, which, once designed, enlarges according to the point growth. This point growth of every branch is, to a certain extent, unlimited. The filaments in and on the substratum are the first existing members of the fungus; they continue so long as it vegetates. As the parts which absorb nourishment from and consume the substance, they are called the mycelium. Nearly every fungus possesses a mycelium, which, without regard to the specific difference of form and size, especially shows the described nature in its construction and growth.

The superficial threads of the mycelium produce other filaments beside those numerous branches which have been described, and which are the fruit thread (carpophore) or conidia thread. These are on an average thicker than the mycelium threads, and only exceptionally ramified or furnished with partitions; they rise almost perpendicularly into the air, and attain a length of, on an average, half a millimetre, or one-fiftieth of an inch, but they seldom become longer, and then their growth is at an end. Their free upper end swells in a rounded manner, and from this is produced, on the whole of its upper part, rayed divergent protuberances, which attain an oval form, and a length almost equal to their radius, or, in weaker specimens, the diameter of the rounded head. The rayed divergent protuberances are the direct producers and bearers of the propagating cells, spores, or conidia, and are called sterigmata. Every sterigma at first produces at its point a little round protuberance, which, with a strong narrow basis, rests upon the sterigma. These are filled with protoplasm, swell more and more, and, after some time, separate themselves by a partition from the sterigma into independent cells, spores, or conidia.

The formation of the first spore takes place at the same end of the sterigma, and in the same manner a second follows, then a third, and so on; every one which springs up later pushes its predecessor in the direction of the axis of the sterigma in the same degree in which it grows itself; every successive spore formed from a sterigma remains for a time in a row with one another. Consequently every sterigma bears on its apex a chain of spores, which are so much the older, the farther they stand from the sterigma. The number of the links in a chain of spores reaches in normal specimens to ten or more. All sterigmata spring up at the same time, and keep pace with one another in the formation of the spores. Every spore grows for a time, according to its construction, and at last separates itself from its neighbours. The mass of dismembered spores forms that fine glaucous hue which is mentioned above. The spores, therefore, are articulated in rows, one after the other, from the ends of the sterigmata. The ripe spore, or conidium, is a cell of a round or broadly oval form, filled with a colourless protoplasm, and, if observed separately, is found to be provided with a brownish, finely verruculose, dotted wall.

The same mycelium which forms the pedicel for the conidia when it is near the end of its development, forms by normal vegetation a second kind of fructification. It begins as delicate thin little branches, which are not to be distinguished by the naked eye, and which mostly in four or six turns, after a quickly terminated growth, wind their ends like a corkscrew. (Fig. 102.) The sinuations decrease in width more and more, till they at last reach close to one another, and the whole end changes from the form of a corkscrew into that of a hollow screw. In and on that screw-like body, a change of a complicated kind takes place, which is a productive process. In consequence of this, from the screw body a globose receptacle is formed, consisting of a thin wall of delicate cells, and a closely entwined row of cells surrounded by this dense mass (d). By the enlargement of all these parts the round body grows so much, that by the time it is ripe it is visible to the naked eye. The outer surface of the wall assumes a compactness and a bright yellow colour; the greater part of the cells of the inner mass become asci for the formation of sporidia, while they free themselves from the reciprocal union, take a broad oval form, and each one produces within its inner space eight sporidia (e). These soon entirely fill the ascus. When they are quite ripe, the wall of the conceptacle becomes brittle, and from irregular fissures, arising easily from contact, the colourless round sporidia are liberated.

The pedicels of both kinds of fruit are formed from the same mycelium in the order just described. If we examine attentively, we can often see both springing up close to one another from the same filament of a mycelium. This is not very easy in the close interlacing of the stalks of a mass of fungi in consequence of their delicacy and fragility. Before their connection was known, the conceptacles and the conidia pedicels were considered as organs of two very different species of fungi. The conceptacles were called Eurotium herbariorum, and the conidia bearers were called Aspergillus glaucus.

Allied to Eurotium is the group of Erysiphei, in which well-authenticated polymorphy prevails. These fungi are developed on the green parts of growing plants, and at first consist of a white mouldy stratum, composed of delicate mycelium, on which erect threads are produced, which break up into subglobose joints or conidia. The species on grass was named Oidium monilioides before its relationship was known, but undoubtedly this is only the conidia of Erysiphe graminis. In like manner the vine disease (Oidium Tuckeri) is most probably only the conidia of a species of Erysiphe, of which the perfect condition has not yet been discovered. On roses the old Oidium leucoconium is but the conidia of Sphærotheca pannosa, and so of other species. The Erysiphe which ultimately appears on the same mycelium consists of globose perithecia, externally furnished with thread-like appendages, and internally with asci containing sporidia. In this genus there are no less than five different forms of fruit, the multiform threads on the mycelium, already alluded to as forms of Oidium, the asci contained in the sporangia, which is the proper fruit of the Erysiphe, larger stylospores which are produced in other sporangia, the smaller stylospores which are generated in the pycnidia, and separate sporules which are sometimes formed in the joints of the necklaces of the conidia. These forms are figured in the "Introduction to Cryptogamic Botany" from Sphærotheca Castagnei, which is the hop mildew. The vine disease, hop mildew, and rose mildew, are the most destructive species of this group, and the constant annoyance of cultivators.

When first describing an allied fungus found on old paper, and named Ascotricha chartarum, the Rev. M. J. Berkeley called attention to the presence of globose conidia attached to the threads which surround the conceptacles, and this occurred as long since as 1838. In a recent species of Chætomium found on old sacking, Chætomium griseum, Cooke, we have found tufts in all respects similar externally to the Chætomium, but no perithecium was formed, naked conidia being developed apparently at the base of the coloured threads. In Chætomium funicolum, Cooke, a black mould was also found which may possibly prove to be its conidia, but at present there is no direct evidence.

The brothers Tulasne have made us acquainted with a greater number of instances amongst the Sphæriacei in which multiple organs of reproduction prevail. Very often old and decaying individuals belonging to species of Boletus will be found filled, and their entire substance internally replaced, by the threads and multitudinous spores of a golden yellow parasite, to which the name of Sepedonium chrysospermum has been given. According to Tulasne, this is merely a condition of a sphæriaceous fungus belonging to his genus Hypomyces.

The same observers also first demonstrated that Trichoderma viride, P., was but the conidia-bearing stage of Hypocrea rufa, P., another sphæriaceous fungus. The ascigerous stroma of the latter is indeed frequently associated in a very close manner with the cushions of the pretended Trichoderma, or in other cases the same stroma will give rise to a different apparatus of conidia, of which the principal elements are acicular filaments, which are short, upright, and almost simple, and which give rise to small oval conidia which are solitary on the tips of the threads. Therefore this Hypocrea will possess two different kinds of conidia, as is the case in many species of Hypomyces.

A most familiar instance of dualism will be found in Nectria cinnabarina, of which the conidia form is one of the most common of fungi, forming little reddish nodules on all kinds of dead twigs.

Almost any small currant twig which has been lying on the ground in a damp situation will afford an opportunity of studying this phenomenon. The whole surface of the twig will be covered from end to end with little bright pink prominences, bursting through the bark at regular distances, scarcely a quarter of an inch apart. Towards one end of the twig probably the prominences will be of a deeper, richer colour, like powdered cinnabar. The naked eye is sufficient to detect some difference between the two kinds of pustules, and where the two merge into each other specks of cinnabar will be visible on the pink projections. By removing the bark it will be seen that the pink bodies have a sort of paler stem, which spreads above into a somewhat globose head, covered with a delicate mealy bloom. At the base it penetrates to the inner bark, and from it the threads of mycelium branch in all directions, confined, however, to the bark, and not entering the woody tissues beneath. The head, placed under examination, will be found to consist of delicate parallel threads compacted together to form the stem and head. Some of these threads are simple, others are branched, bearing here and there upon them delicate little bodies, which are readily detached, and which form the mealy bloom which covers the surface. These are the conidia, little slender cylindrical bodies, rounded at the ends.

Passing to the other bodies, which are of a deeper colour, it will soon be discovered that, instead of being simple rounded heads, each tubercle is composed of numerous smaller, nearly globose bodies, closely packed together, often compressed, all united to a base closely resembling the base of the other tubercles. If for a moment we look at one of the tubercles near the spot where the crimson tubercles seem to merge into the pink, we shall not only find them particoloured, but that the red points are the identical globose little heads just observed in clusters. This will lead to the suspicion, which can afterwards be verified, that the red heads are really produced on the stem or stroma of the pink tubercles.

A section of one of the red tubercles will show us how much the internal structure differs. The little subglobose bodies which spring from a common stroma or stem are hollow shells or capsules, externally granular, internally filled with a gelatinous nucleus. They are, indeed, the perithecia of a sphæriaceous fungus of the genus Nectria, and the gelatinous nucleus contains the fructification. Still further examination will show that this fructification consists of cylindrical asci, each enclosing eight elliptical sporidia, closely packed together, and mixed with slender threads called paraphyses.

Here, then, we have undoubted evidence of Nectria cinnabarina, with its fruit, produced in asci growing from the stroma or stem, and in intimate relationship with what was formerly named Tubercularia vulgaris. A fungus with two forms of fruit, one proper to the pink, or Tubercularia form, with naked slender conidia, the other proper to the mature fungus, enclosed in asci, and generated within the walls of a perithecium. Instances of this kind are now known to be far from uncommon, although they cannot always, or often, be so clearly and distinctly traced as in the illustration which we have selected.

It is not uncommon for the conidia of the Sphæria to partake of the characteristics of a mould, and then the perithecia are developed amongst the conidial threads. A recently recorded instance of this relates to Sphæria Epochnii, B. and Br., the conidia form of which was long known before the Sphæria related to it was discovered, under the name of Epochnium fungorum. The Epochnium forms a thin stratum, which overruns various species of Corticium. The conidia are at first uniseptate. The perithecia of the Sphæria are at first pale bottle-green, crowded in the centre of the Epochnium, then black green granulated, sometimes depressed at the summit, with a minute pore. The sporidia are strongly constricted in the centre, at first uniseptate, with two nuclei in each division.

Another Sphæria in which the association is undoubted is the Sphæria aquila, Fr., which is almost always found nestling in a woolly brown subiculum, for the most part composed of barren brown jointed threads. These threads, however, produce, under favourable conditions, mostly before the perfection of the perithecia, minute subglobose conidia, and in this state constitute what formerly bore the name of Sporotrichum fuscum, Link., but now recognized as the conidia of Sphæria aquila.

In Sphæria nidulans, Schw., a North American species, we have more than once found the dark brown subiculum bearing large triseptate conidia, having all the characters of the genus Helminthosporium. In Sphæria pilosa, P., Messrs. Berkeley and Broome have observed oblong conidia, rather irregular in outline, terminating the hairs of the perithecium. The same authors have also figured the curious pentagonal conidia springing from flexuous threads accompanying Sphæria felina, Fckl., and also the threads resembling those of a Cladotrichum with the angular conidia of Sphæria cupulifera, B. and Br. A most remarkable example is also given by the Brothers Tulasne in Pleospora polytricha, in which the conidia-bearing threads not only surround, but grow upon the perithecia, and are crowned by fascicles of septate conidia.

Instances of this kind have now become so numerous that only a few can be cited as examples of the rest. It is not at all improbable that the majority of what are now classed together as species under the genus of black moulds, Helminthosporium, will at some not very distant period be traced as the conidia of different species of ascomycetous fungi. The same fate may also await other allied genera, but until this association is established, they must keep the rank and position which has been assigned to them.

Another form of dualism, differing somewhat in character from the foregoing, finds illustration in the sphæriaceous genus Melanconis, of Tulasne, in which the free spores are still called conidia, though in most instances produced in a sort of spurious conceptaculum, or borne on short threads from a kind of cushion-shaped stroma. In the Melanconis stilbostoma, there are three forms, one of slender minute bodies, oozing out in the form of yellow tendrils, which may be spermatia, formerly called Nemaspora crocea. Then there are the oval brown or olive brown conidia, which are at first covered, then oozing out in a black pasty mass, formerly Melanconium bicolor, and finally the sporidia in asci of Sphæria stilbostoma, Fries. In Melanconis Berkeleii, Tul., the conidia are quadrilocular, previously known as Stilbospora macrosperma, B. and Br. In a closely-allied species from North America, Melanconis bicornis, Cooke, the appendiculate sporidia are similar, and the conidia would also appear to partake of the character of Stilbospora. We may remark here that we have seen a brown mould, probably an undescribed species of Dematiei, growing in definite patches around the openings in birch bark caused by the crumpent ostiola of the perithecia of Melanconis stilbostoma, from the United States.

In Melanconis lanciformis, Tul., there are, it would appear, four forms of fruit. One of these consists of conidia, characterized by Corda as Coryneum disciforme. Stylospores, which are also figured by Corda under the name of Coniothecium betulinum; pycnidia, first discovered by Berkeley and Broome, and named by them Hendersonia polycystis; and the ascophorous fruits which constituted the Sphæria lanciformis of Fries. Mr. Currey indicated Hendersonia polycystis, B. and Br., as a form of fruit of this species in a communication to the Royal Society in 1857. He says this plant grows upon birch, and is in perfection in very moist weather, when it may be recognized by the large black soft gelatinous protuberances on the bark, formed by spores escaping and depositing themselves upon and about the apex of the perithecium. This I suspect to be an abnormal state of a well-known Sphæria (S. lanciformis), which grows upon birch, and upon birch only.

We might multiply, almost indefinitely, instances amongst the Sphæriacei, but have already given sufficient for illustration, and will therefore proceed briefly to notice some instances amongst the Discomycetes, which also bear their complete or perfect fruit in asci.

The beautiful purple stipitate cups of Bulgaria sarcoides, which may be seen flourishing in the autumn on old rotten wood, are often accompanied by club-shaped bodies of the same colour; or earlier in the season these clavate bodies may be found alone, and at one time bore the name of Tremella sarcoides. The upper part of these clubs disseminate a great abundance of straight and very slender spermatia. Earlier than this they are covered with globose conidia. The fully-matured Bulgaria develops on its hymenium clavate delicate asci, each enclosing eight elongated hyaline sporidia, so that we have three forms of fruit belonging to the same fungus, viz. conidia and spermatia in the Tremella stage, and sporidia contained in asci in the mature condition. The same phenomena occur with Bulgaria purpurea, a larger species with different fruit, long confounded with Bulgaria sarcoides.

On the dead stems of nettles it is very common to meet with small orange tubercles, not much larger than a pin's head, which yield at this stage a profusion of slender linear bodies, produced on delicate branched threads, and at one time bore the name of Dacrymyces Urticæ, but which are now acknowledged to be only a condition of a little tremelloid Peziza of the same size and colour, which might be mistaken for it, if not examined with the microscope, but in which there are distinct asci and sporidia. Both forms together are now regarded as the same fungus, under the name of Peziza fusarioides, B.

The other series of phenomena grouped together under the name of polymorphism relate to forms which are removed from each other, so that the mycelium is not identical, or, more usually, produced on different plants. The first instance of this kind to which we shall make reference is one of particular interest, as illustrative of the old popular creed, that berberry bushes near corn-fields produced mildewed corn. There is a village in Norfolk, not far from Great Yarmouth, called "Mildew Rollesby," because of its unenviable notoriety in days past for mildewed corn, produced, it was said, by the berberry bushes, which were cut down, and then mildew disappeared from the corn-fields, so that Rollesby no longer merited its sobriquet. It has already been shown that the corn-mildew (Puccinia graminis) is dimorphous, having a one-celled fruit (Trichobasis), as well as a two-celled fruit (Puccinia). The fungus which attacks the berberry is a species of cluster-cup (Æcidium berberidis), in which little cup-like peridia, containing bright orange pseudospores, are produced in tufts or clusters on the green leaves, together with their spermogonia.

De Bary's observations on this association of forms were published in 1865. In view of the popular belief, he determined to sow the spores of Puccinia graminis on the leaves of the berberry. For this purpose he selected the septate resting spores from Poa pratensis and Triticum repens. Having caused the spores to germinate in a moist atmosphere, he placed fragments of the leaves on which they had developed their secondary spores on young but full-grown berberry leaves, under the same atmospheric conditions. In from twenty-four to forty-eight hours a quantity of the germinating threads had bored through the walls and penetrated amongst the subjacent cells. This took place both on the upper and under surface of the leaves. Since, in former experiments, it appeared that the spores would penetrate only in those cases where the plant was adapted to develop the parasite, the connection between P. graminis and Æcid. berberidis seemed more than ever probable. In about ten days the spermogonia appeared. After a time the cut leaves began to decay, so that the fungus never got beyond the spermogonoid stage. Some three-year-old seedlings were then taken, and the germinating resting spores applied as before. The plants were kept under a bell-glass from twenty-four to forty-eight hours, and then exposed to the air like other plants. From the sixth to the tenth day, yellow spots appeared, with single spermogonia; from the ninth to the twelfth, spermogonia appeared in numbers on either surface; and, a few days later, on the under surface of the leaves, the cylindrical sporangia of the Æcidium made their appearance, exactly as in the normally developed parasite, except that they were longer, from being protected from external agents. The younger the leaves, the more rapid was the development of the parasite, and sometimes, in the younger leaves, the luxuriance was far greater than in free nature. Similar plants, to the number of two hundred, were observed in the nursery, and though some of them had Æcidium pustules, not one fresh pustule was produced; while two placed under similar circumstances, but without the application of any resting spores, remained all the summer free from Æcidium. It seems, then, indubitable so far that Æcidium berberidis does spring from the spores of Puccinia graminis.

It has, however, to be remarked that De Bary was not equally successful in producing the Puccinia from the spores of the Æcidium. In many cases the spores do not germinate when placed on glass, and they do not preserve their power of germinating very long. He reverts then to the evidence of experiments instituted by agriculturists. Bönninghausen remarked, in 1818, that wheat, rye, and barley which were sown in the neighbourhood of a berberry bush covered with Æcidium contracted rust immediately after the maturation of the spores of the Æcidia. The rust was most abundant where the wind carried the spores. The following year the same observations were repeated; the spores of the Æcidium were collected, and applied to some healthy plants of rye. After five or six days these plants were affected with rust, while the remainder of the crop was sound. In 1863 some winter rye was sown round a berberry bush, which in the following year was infested with Æcidium, which was mature in the middle of May, when the rye was completely covered with rust. Of the wild grasses near the bush, Triticum repens was most affected. The distant plants of rye were free from rust.

The spores of the Æcidium would not germinate on berberry leaves; the berberry Æcidium could not therefore spring from the previous Æcidium. The uredospores of Puccinia graminis on germinating penetrate into the parenchym of the grass on which they are sown; but on berberry leaves, if the tips of the threads enter for a short distance into the stomates their growth at once ceases, and the leaves remain free from parasites.