Chapter M: J. Berkeley (5)

"As I have said, the Agaric of the olive-tree, which is itself very yellow, reflects a strong brilliant light, and remains endowed with this remarkable faculty whilst it grows, or, at least, while it appears to preserve an active life, and remains fresh. The phosphorescence is at first, and more ordinarily, recognizable at the surface of the hymenium. I have seen a great number of young fungi which were very phosphorescent in the gills, but not in any other part. In another case, and amongst more aged fungi, the hymenium of which had ceased to give light, the stipe, on the contrary, threw out a brilliant glare. Habitually, the phosphorescence is distributed in an unequal manner upon the stipe, and the same upon the gills. Although the stipe is luminous at its surface, it is not always necessarily so in its interior substance, if one bruises it, but this substance frequently becomes phosphorescent after contact with the air. Thus, I had irregularly split and slit a large stipe in its length, and I found the whole flesh obscure, whilst on the exterior were some luminous places. I roughly joined the lacerated parts, and the following evening, on observing them anew, I found them all flashing a bright light. At another time, I had with a scalpel split vertically many fungi in order to hasten their dessication; the evening of the same day, the surface of all these cuts was phosphorescent, but in many of these pieces of fungi the luminosity was limited to the cut surface which remained exposed to the air; the flesh beneath was unchanged.

"I have seen a stipe opened and lacerated irregularly, the whole of the flesh of which remained phosphorescent during three consecutive evenings, but the brightness diminished in intensity from the exterior to the interior, so that on the third day it did not issue from the inner part of the stipe. The phosphorescence of the gills is in no way modified at first by immersing the fungus in water; when they have been immersed they are as bright as in the air, but the fungi which I left immersed until the next evening lost all their phosphorescence, and communicated to the water an already sensible yellow tint; alcohol put upon the phosphorescent gills did not at once completely obliterate the light, but visibly enfeebled it. As to the spores, which are white, I have found many times very dense coats of them thrown down on porcelain plates, but I have never seen them phosphorescent.

"As to the observation made by Delile that the Agaric of the olive does not shine during the day when placed in total darkness, I think that it could not have been repeated. From what I have said of the phosphorescence of _A. olearius_, one naturally concludes that there does not exist any necessary relation between this phenomenon and the fructification of the fungus; the luminous brightness of the hymenium shows, says Delile, 'the greater activity of the reproductive organs,' but it is not in consequence of its reproductive functions, which may be judged only as an accessory phenomenon, the cause of which is independent of, and more general than these functions, since all the parts of the fungus, its entire substance, throws forth at one time, or at successive times, light. From these experiments Tulasne infers that the same agents, oxygen, water, and warmth, are perfectly necessary to the production of phosphorescence as much in living organized beings as in those which have ceased to live. In either case, the luminous phenomena accompany a chemical reaction which consists principally in a combination of the organized matter with the oxygen of the air; that is to say, in its combustion, and in the discharge of carbonic acid which thus shows itself."

We have quoted at considerable length from these observations of Tulasne on the Agaric of the olive, as they serve very much to illustrate similar manifestations in other species, which doubtless resemble each other in their main features.

Mr. Gardner has graphically described his first acquaintance in Brazil with the phosphorescent species which now bears his name. It was encountered on a dark night of December, while passing through the streets of Villa de Natividate. Some boys were amusing themselves with some luminous object, which at first he supposed to be a kind of large fire-fly, but on making inquiry he found it to be a beautiful phosphorescent Agaric, which he was told grew abundantly in the neighbourhood on the decaying fronds of a dwarf palm. The whole plant gives out at night a bright light somewhat similar to that emitted by the larger fire-flies, having a pale greenish hue. From this circumstance, and from growing on a palm, it was called by the inhabitants "flor de coco."[C]

The number of recognized phosphorescent species of _Agaricus_ is not large, although two or three others may be enumerated in addition to those cited by Tulasne. Of these, _Agaricus lampas_, and some others, are found in Australia.[D] In addition to the _Agaricus noctileucus_, discovered by Gaudichaud, and the _Agaricus igneus_ of Rumphius, found in Amboyna, Dr. Hooker speaks of the phenomenon as common in Sikkim, but he seems never to have been able to ascertain with what species it was associated.

Dr. Cuthbert Collingwood has communicated some further information relative to the luminosity of a species of _Agaricus_ in Borneo (supposed to be _A. Gardneri_), in which he says, "The night being dark, the fungi could be very distinctly seen, though not at any great distance, shining with a soft pale greenish light. Here and there spots of much more intense light were visible, and these proved to be very young and minute specimens. The older specimens may more properly be described as possessing a greenish luminous glow, like the glow of the electric discharge, which, however, was quite sufficient to define its shape, and, when closely examined, the chief details of its form and appearance. The luminosity did not impart itself to the hand, and did not appear to be affected by the separation from the root on which it grew, at least not for some hours. I think it probable that the mycelium of this fungus is also luminous, for, upon turning up the ground in search of small luminous worms, minute spots of light were observed, which could not be referred to any particular object or body when brought to the light and examined, and were probably due to some minute portions of its mycelium."[E] The same writer also adds, "Mr. Hugh Low has assured me that he saw the jungle all in a blaze of light (by which he could see to read) as, some years ago, he was riding across the island by the jungle road; and that this luminosity was produced by an Agaric."

Similar experiences were detailed by Mr. James Drummond in a letter from Swan River, in which two species of Agaric are concerned. They grew on the stumps of trees, and had nothing remarkable in their appearance by day, but by night emitted a most curious light, such as the writer never saw described in any book. One species was found growing on the stump of a _Banksia_ in Western Australia. The stump was at the time surrounded by water. It was on a dark night, when passing, that the curious light was first observed. When the fungus was laid on a newspaper, it emitted by night a phosphorescent light, enabling persons to read the words around it, and it continued to do so for several nights with gradually decreasing intensity as the plant dried up. In the other instance, which occurred some years after, the author, during one of his botanical trips, was struck by the appearance of a large Agaric, measuring sixteen inches in diameter, and weighing about five pounds. This specimen was hung up to dry in the sitting-room, and on passing through the apartment in the dark it was observed to give out the same remarkable light. The luminous property continued, though gradually diminishing, for four or five nights, when it ceased on the plant becoming dry. "We called some of the natives," he adds, "and showed them this fungus when emitting light, and the poor creatures cried out 'chinga,' their name for a spirit, and seemed much afraid of it."[F]

Although the examples already cited are those of species of Agaric, luminosity is not by any means wholly confined to that genus. Mr. Worthington Smith has recorded his experiences of some specimens of the common _Polyporus annosus_ which were found on some timbers in the Cardiff coal mines. He remarks that the colliers are well acquainted with phosphorescent fungi, and the men state that sufficient light is given "to see their hands by." The specimens of _Polyporus_ were so luminous that they could be seen in the dark at a distance of twenty yards. He observes further, that he has met with specimens of _Polyporus sulfureus_ which were phosphorescent. Some of the fungi found in mines, which emit light familiar to the miners, belong to the incomplete genus _Rhizomorpha_, of which Humboldt amongst others gives a glowing account. Tulasne has also investigated this phenomenon in connection with the common _Rhizomorpha subterranea_, Pers. This species extends underneath the soil in long strings, in the neighbourhood of old tree stumps, those of the oak especially, which are becoming rotten, and upon these it is fixed by one of its branches. These are cylindrical, very flexible, branching, and clothed with a hard bark, encrusting and fragile, at first smooth and brown, becoming later very rough and black. The interior tissue, at first whitish, afterwards of a more or less deep brown colour, is formed of extremely long parallel filaments from .0035 to .015 _mm._ in diameter.

On the evening of the day when I received the specimens,[G] he writes, the temperature being about 22° Cent., all the young branches brightened with an uniform phosphoric light the whole of their length; it was the same with the surface of some of the older branches, the greater number of which were still brilliant in some parts, and only on their surface. I split and lacerated many of these twigs, but their internal substance remained dull. The next evening, on the contrary, this substance, having been exposed to contact with the air, exhibited at its surface the same brightness as the bark of the branches. I made this observation upon the old stalks as well as upon the young ones. Prolonged friction of the luminous surfaces reduced the brightness and dried them to a certain degree, but did not leave on the fingers any phosphorescent matter. These parts continued with the same luminous intensity after holding them in the mouth so as to moisten them with saliva; plunged into water, held to the flame of a candle so that the heat they acquired was very appreciable to the touch, they still emitted in the dark a feeble light; it was the same after being held in water heated to 30° C.; but putting them in water bearing a temperature of 55° C. extinguished them entirely. They are equally extinguished if held in the mouth until they catch the temperature; perhaps, still, it might be attributed less to the heat which is communicated to them than to the deficiency of sufficient oxygen, because I have seen some stalks, having become dull in the mouth, recover after a few instants a little of their phosphorescence. A young stalk which had been split lengthwise, and the internal substance of which was very phosphorescent, could imbibe olive oil many times and yet continue for a long time to give a feeble light. By preserving these _Rhizomorphæ_ in an adequate state of humidity, I have been able for many evenings to renew the examination of their phosphorescence; the commencement of dessication, long before they really perish, deprives them of the faculty of giving light. Those which had been dried for more than a month, when plunged into water, commenced to vegetate anew and send forth numerous branches in a few days; but I could only discover phosphorescence at the surface of these new formations, or very rarely in their immediate neighbourhood, the mother stalks appearing to have lost by dessication their luminous properties, and did not recover them on being recalled to life. These observations prove that what Schmitz has written was not true, that all parts of these fungi were seldom phosphorescent.

The luminous phenomenon in question is without doubt more complicated than it appears, and the causes to which we attribute it are certainly powerfully modified by the general character of the objects in which they reside. Most of the German botanists give this explanation, others suppose that it forms at first or during its continuance a special matter, in which the luminous property resides; this matter, which is said to be mucilaginous in the luminous wood, appears to be in the _Rhizomorpha_ only a kind of chemical combination between the membrane and some gummy substance which they contain. Notwithstanding this opinion, I am assured that all external mucous matter was completely absent from the _Agaricus olearius_, and I neither discovered it upon the branches of _Rhizomorpha subterranea_ nor upon the dead leaves which I have seen phosphorescent; in all these objects the luminous surfaces were nothing else than their proper tissue.

It may be remarked here that the so-called species of _Rhizomorpha_ are imperfect fungi, being entirely devoid of fructification, consisting in fact only of a vegetative system--a sort of compact mycelium--(probably of species of _Xylaria_) with some affinity to _Sclerotium_.

Recently an extraordinary instance of luminosity was recorded as occurring in our own country.[H] "A quantity of wood had been purchased in a neighbouring parish, which was dragged up a very steep hill to its destination. Amongst them was a log of larch or spruce, it is not quite certain which, 24 feet long and a foot in diameter. Some young friends happened to pass up the hill at night, and were surprised to find the road scattered with luminous patches, which, when more closely examined, proved to be portions of bark or little fragments of wood. Following the track, they came to a blaze of white light which was perfectly surprising. On examination, it appeared that the whole of the inside of the bark of the log was covered with a white byssoid mycelium of a peculiarly strong smell, but unfortunately in such a state that the perfect form could not be ascertained. This was luminous, but the light was by no means so bright as in those parts of the wood where the spawn had penetrated more deeply, and where it was so intense that the roughest treatment scarcely seemed to check it. If any attempt was made to rub off the luminous matter it only shone the more brightly, and when wrapped up in five folds of paper the light penetrated through all the folds on either side as brightly as if the specimen was exposed; when, again, the specimens were placed in the pocket, the pocket when opened was a mass of light. The luminosity had now been going on for three days. Unfortunately we did not see it ourselves till the third day, when it had, possibly from a change in the state of electricity, been somewhat impaired; but it was still most interesting, and we have merely recorded what we observed ourselves. It was almost possible to read the time on the face of a watch even in its less luminous condition. We do not for a moment suppose that the mycelium is essentially luminous, but are rather inclined to believe that a peculiar concurrence of climatic conditions is necessary for the production of the phenomenon, which is certainly one of great rarity. Observers as we have been of fungi in their native haunts for fifty years, it has never fallen to our lot to witness a similar case before, though Prof. Churchill Babington once sent us specimens of luminous wood, which had, however, lost their luminosity before they arrived. It should be observed that the parts of the wood which were most luminous were not only deeply penetrated by the more delicate parts of the mycelium, but were those which were most decomposed. It is probable, therefore, that this fact is an element in the case as well as the presence of fungoid matter."

In all cases of phosphorescence recorded, the light emitted is described as of the same character, varying only in intensity. It answers well to the name applied to it, as it seems remarkably similar to the light emitted by some living insects and other animal organisms, as well as to that evolved, under favourable conditions, by dead animal matter--a pale bluish light, resembling that emitted by phosphorus as seen in a dark room.

Another phenomenon worthy of note is the change of colour which the bruised or cut surface of some fungi undergo. Most prominent amongst these are certain poisonous species of _Boletus_, such, for instance, as _Boletus luridus_, and some others, which, on being bruised, cut, or divided, exhibit an intense, and in some cases vivid, blue. At times this change is so instantaneous that before the two freshly-cut portions of a _Boletus_ can be separated, it has already commenced, and proceeds rapidly till the depth of intensity has been gained. This blue colour is so universally confined to dangerous species that it is given as a caution that all species which exhibit a blue colour when cut or bruised, should on no account be eaten. The degree of intensity varies considerably according to the condition of the species. For example, _Boletus cærulescens_ is sometimes only very slightly, if at all, tinged with blue when cut, though, as the name implies, the peculiar phenomenon is generally highly developed. It cannot be said that this change of colour has as yet been fully investigated. One writer some time since suggested, if he did not affirm, that the colour was due to the presence of aniline, others have contented themselves with the affirmation that it was a rapid oxidization and chemical change, consequent upon exposure of the surfaces to the air. Archdeacon Robinson examined this phenomenon in different gases, and arrived at the conclusion that the change depends on an alteration of molecular arrangement.[I]

One of the best of the edible species of _Lactarius_, known as _Lactarius deliciosus_, changes, wherever cut or bruised, to a dull livid green. This fungus is filled with an orange milky fluid, which becomes green on exposure to the air, and it is consequently the juice which oxidizes on exposure. Some varieties more than others of the cultivated mushroom become brownish on being cut, and a similar change we have observed, though not recorded, in other species.

The presence of a milky juice in certain fungi has been alluded to. This is by no means confined to the genus _Lactarius_, in which such juice is universal, sometimes white, sometimes yellow, and sometimes colourless. In Agarics, especially in the subgenus _Mycena_, the gills and stem are replete with a milky juice. Also in some species of _Peziza_, as for instance in _Peziza succosa_, B., sometimes found growing on the ground in gardens, and in _Peziza saniosa_, Schrad., also a terrestrial species, the same phenomenon occurs. To this might be added such species as _Stereum spadiceum_, Fr., and _Stereum sanguinolentum_, Fr., both of which become discoloured and bleeding when bruised, while _Corticium lactescens_ distils a watery milk.

Fungi in general have not a good repute for pleasant odours, and yet it must be conceded that they are not by any means devoid of odour, sometimes peculiar, often strong, and occasionally very offensive. There is a peculiar odour common to a great many forms, which has come to be called a fungoid odour; it is the faint smell of a long-closed damp cellar, an odour of mouldiness and decay, which often arises from a process of eremocausis. But there are other, stronger, and equally distinct odours, which, when once inhaled, are never to be forgotten. Amongst these is the fetid odour of the common stinkhorn, which is intensified in the more beautiful and curious _Clathrus_. It is very probable that, after all, the odour of the _Phallus_ would not be so unpleasant if it were not so strong. It is not difficult to imagine, when one encounters a slight sniff borne on a passing breeze, that there is the element of something not by any means unpleasant about the odour when so diluted; yet it must be confessed that when carried in a vasculum, in a close carriage, or railway car, or exposed in a close room, there is no scruple about pronouncing the odour intensely fetid. The experience of more than one artist, who has attempted the delineation of _Clathrus_ from the life, is to the effect that the odour is unbearable even by an enthusiastic artist determined on making a sketch.

Perhaps one of the most fetid of fungi is _Thelephora palmata_. Some specimens were on one occasion taken by Mr. Berkeley into his bedroom at Aboyne, when, after an hour or two, he was horrified at finding the scent far worse than that of any dissecting room. He was anxious to save the specimens, but the scent was so powerful that it was quite intolerable till he had wrapped them in twelve thick folds of the strongest brown paper. The scent of _Thelephora fastidiosa_ is bad enough, but, like that of _Coprinus picaceus_, it is probably derived from the imbibition of the ordure on which it is developed. There needs no stronger evidence that the scent must not only be powerful, but unpleasant, when an artist is compelled, before a rough sketch is more than half finished, to throw it away, and seek relief in the open air. A great number of edible Agarics have the peculiar odour of fresh meal, but two species, _Agaricus odorus_ and _Agaricus fragrans_, have a pleasant anise-like odour. In two or three species of tough _Hydnum_, there is a strong persistent odour somewhat like melilot or woodruffe, which does not pass away after the specimen has been dried for years. In some species of _Marasmius_, there is a decidedly strong odour of garlic, and in one species of _Hygrophorus_, such a resemblance to that of the larva of the goat moth, that it bears the name of _Hygrophorus cossus_. Most of the fleshy forms exhale a strong nitrous odour during decay, but the most powerful we remember to have experienced was developed by a very large specimen of _Choiromyces meandriformis_, a gigantic subterranean species of the truffle kind, and this specimen was four inches in diameter when found, and then partially decayed. It was a most peculiar, but strong and unpleasantly pungent nitrous odour, such as we never remember to have met with in any other substance. _Peziza venosa_ is remarkable when fresh for a strong scent like that of aquafortis.

Of colour, fungi exhibit an almost endless variety, from white, through ochraceous, to all tints of brown until nearly black, or through sulphury yellow to reds of all shades, deepening into crimson, or passing by vinous tints into purplish black. These are the predominating gradations, but there are occasional blues and mineral greens, passing into olive, but no pure or chlorophyllous green. The nearest approach to the latter is found in the hymenium of some _Boleti_. Some of the Agarics exhibit bright colours, but the larger number of bright-coloured species occur in the genus _Peziza_. Nothing can be more elegant than the orange cups of _Peziza aurantia_, the glowing crimson of _Peziza coccinea_, the bright scarlet of _Peziza rutilans_, the snowy whiteness of _Peziza nivea_, the delicate yellow of _Peziza theleboloides_, or the velvety brown of _Peziza repanda_. Amongst Agarics, the most noble _Agaricus muscarius_, with its warty crimson pileus, is scarcely eclipsed by the continental orange _Agaricus cæsarius_. The amethystine variety of _Agaricus laccatus_ is so common and yet so attractive; whilst some forms and species _Russula_ are gems of brilliant colouring. The golden tufts of more than one species of _Clavaria_ are exceedingly attractive, and the delicate pink of immature _Lycogala epidendrum_ is sure to command admiration. The minute forms which require the microscope, as much to exhibit their colour as their structure, are not wanting in rich and delicate tints, so that the colour-student would find much to charm him, and good practice for his pencil in these much despised examples of low life.

Amongst phenomena might be cursorily mentioned the peculiar sarcodioid mycelium of _Myxogastres_, the development of amoeboid forms from their spores, and the extraordinary rapidity of growth, as the well-known instance of the _Reticularia_ which Schweinitz observed running over iron a few hours after it had been red hot. Mr. Berkeley has observed that the creamy mycelium of _Lycogala_ will not revive after it has become dry for a few hours, though so active before.

[A] M. J. Berkeley, "Introduction to Cryptogamic Botany," p. 265.

[B] Tulasne, "Sur la Phosphorescence des Champignons," in "Ann. des
Sci. Nat." (1848), vol. ix, p. 338.

[C] In "Hooker's Journal of Botany" (1840), vol. ii. p. 426.

[D] Berkeley, "Introduction to Crypt. Bot." t. 265.

[E] Dr. Collingwood, in "Journal of Linnæan Society (Botany)," vol.
x. p. 469.

[F] In "Hooker's Journal of Botany" for April, 1842.

[G] Tulasne, "Sur la Phosphorescence," in "Ann. des Sci. Nat."
(1848), vol ix. p. 340, &c.

[H] Rev. M. J. Berkeley, in "Gardener's Chronicle" for 1872, p.
1258.

[I] Berkeley, "Introduction to Crypt. Bot." p. 266.

VI.

THE SPORE AND ITS DISSEMINATION.

A work of this character would hardly be deemed complete without some reference to the above subject, which has moreover a relation to some of the questions discussed, and particularly of spore diffusion in the atmosphere. The largest spore is microscopic, and the smallest known scarcely visible under a magnifying power of 360 diameters. Taking into account the large number of species of fungi, probably scarcely less numerous than all the flowering plants, and the immense number of spores which some of the individuals produce, they must be exceedingly plentiful and widely diffused, though from their minuteness not easy to be discerned. It has been attempted to estimate the number of spores which might be produced by one single plant of _Lycoperdon_, but the number so far exceeds that which the mind is accustomed to contemplate that it seems scarcely possible to realize their profusion. Recent microscopic examinations of the common atmosphere[A] show the large quantity of spores that are continually suspended. In these investigations it was found that spores and similar cells were of constant occurrence, and were generally present in considerable numbers. That the majority of the cells were living, and ready to undergo development on meeting with suitable conditions, was very manifest, as in those cases in which preparations were retained under observation for any length of time, germination rapidly took place in many of the cells. In few instances did any development take place, beyond the formation of networks of mycelium, or masses of toruloid cells, but, in one or two, distinct sporules were developed on the filaments arising from some of the larger septate spores; and in a few others, _Penicillium_ and _Aspergillus_ produced their characteristic heads of fructification. With regard to the precise nature of the spores, and other cells present in various instances, little can be said, as, unless their development were to be carefully followed out through all its stages, it is impossible to refer them to their correct species or even genera. The greater number of them are apparently referable to the old orders of fungi, _Sphæronemei_, _Melanconei_, _Torulacei_, _Dematiei_ and _Mucedines_, while some probably belonged to the _Pucciniæi_ and _Cæomacei_.

Hence it is demonstrated that a large number of the spores of fungi are constantly present in the atmosphere, which is confirmed by the fact that whenever a suitable pabulum is exposed it is taken possession of by floating spores, and soon converted into a forest of fungoid vegetation. It is admitted that the spores of such common moulds as _Aspergillus_ and _Penicillium_ are so widely diffused, that it is almost impossible to exclude them from closed vessels, or the most carefully guarded preparations. Special contrivances for the dispersion of the spores in the different groups follow a few general types, and it is only rarely that we meet with any method that is confined only to a species or genus. Some of the more significant forms of spores may be illustrated, with their modes of dissemination.

BASIDIOSPORES is a term which we may employ here to designate all spores borne at the tips of such supports as are found in the _Hymenomycetes_ and _Gasteromycetes_, to which the name of basidia has been given. In fact, under this section we may include all the spores of those two orders, although we may be ignorant of the precise mode in which the fruit of most of the _Myxogastres_ is developed. Guarding ourselves at the outset against any misinterpretation as to the use of this term, which, in fact, we employ simply to designate the fruit of _Hymenomycetes_, we may have excuse in our desire to limit special terms as much as possible. In the _Agaricini_ the spores are plentiful, and are distributed over the hymenium or gill plates, the surface of which is studded with basidia, each of which normally terminates with four short, erect, delicate, thread-like processes, each of which is surmounted by a spore. These spores are colourless or coloured, and it is upon this fact that primary divisions in the genus _Agaricus_ are based, inasmuch as colour in the spores appears to be a permanent feature. In white-spored species the spores are white in all the individuals, not mutable as the colour of the pileus, or the corolla in phanerogamic plants. So also with the pink spored, rusty spored, black spored, and others. This may serve to explain why colour, which is so little relied upon in classification amongst the higher plants, should be introduced as an element of classification in one of the largest genera of fungi.

There are considerable differences in size and form amongst the spores of the _Agaricini_, although at first globose; when mature they are globose, oval, oblong, elliptic, fusiform, and either smooth or tuberculated, often maintaining in the different genera or subgenera one particular characteristic, or typical form. It is unnecessary here to particularize all the modifications which the form and colour of the spores undergo in different species, as this has already been alluded to. The spores in the _Polyporei_, _Hydnei_, &c., are less variable, of a similar character, as in all the _Hymenomycetes_, except perhaps the _Tremellini_.

When an Agaric is mature, if the stem is cut off close to the gills, and the pileus inverted, with the gills downwards on a sheet of black paper (one of the pale-spored species is best for this purpose), and left for a few hours, or all night, in that position, the paper will be found imprinted in the morning with a likeness of the under side of the pileus with its radiating gills, the spores having been thrown down upon the paper in such profusion, from the hymenium, and in greater numbers from the opposed surfaces of the gills. This little experiment will be instructive in two or three points. It will illustrate the facility with which the spores are disseminated, the immense number in which they are produced, and the adaptability of the gill structure to the economy of space, and the development of the largest number of basidiospores from a given surface. The tubes or pores in _Polyporei_, the spines in _Hydnei_, are modifications of the same principles, producing a like result.

In the _Gasteromycetes_ the spores are produced in many cases, probably in most, if not all, at the tips of sporophores; but the hymenium, instead of being exposed, as in the _Hymenomycetes_, is enclosed within an outer peridium or sac, which is sometimes double. The majority of these spores are globose in form, some of them extremely minute, variously coloured, often dark, nearly black, and either externally smooth or echinulate. In some genera, as _Enerthenema_, _Badhamia_, &c., a definite number of spores are at first enclosed in delicate cysts, but these are exceptions to the general rule: this also is the case in at least one species of _Hymenogaster_. As the spores approach maturity, it may be observed in such genera as _Stemonitis_, _Arcyria_, _Diachea_, _Dictydium_, _Cribraria_, _Trichia_, &c., that they are accompanied by a sort of reticulated skeleton of threads, which remain permanent, and served in earlier stages, doubtless, as supports for the spores; being, in fact, the skeleton of the hymenium. It has been suggested that the spiral character of the threads in _Trichia_ calls to mind the elaters in the _Hepaticæ_, and like them may, by elasticity, aid in the dispersion of the spores. There is nothing known, however, which will warrant this view. When the spores are mature, the peridium ruptures either by an external orifice, as in _Geaster_, _Lycoperdon_, &c., or by an irregular opening, and the light, minute, delicate, spores are disseminated by the slightest breath of air. Specimens of _Geaster_ and _Bovista_ are easily separated from the spot on which they grew; when rolling from place to place, the spores are deposited over a large surface. In the _Phalloidei_ the spores are involved in a slimy mucus which would prevent their diffusion in such a manner. This gelatinous substance has nevertheless a peculiar attraction for insects, and it is not altogether romantic to believe that in sucking up the fetid slime, they also imbibe the spores and transfer them from place to place, so that even amongst fungi insects aid in the dissemination of species. Whether or not the _Myxogastres_ should be included here is matter of opinion, since the mode in which the spores are developed is but little known; analogy with the _Trichogastres_ in other points alone leading to the conclusion that they may produce basidiospores. The slender, elastic stems which support the peridia in many species are undoubted aids to the dissemination of the spores.[B]

Under the name of STYLOSPORES may be classed those spores which in some orders of _Coniomycetes_ are produced at the apex of short threads, either enclosed in a perithecium, or seated upon a kind of stroma. These are exceedingly variable, sometimes large, and multiseptate, at other times minute, resembling spermatia. In such genera as are chiefly epiphytal, in _Septoria_, _Phyllosticta_, and their allies, the minute spores are enclosed within membranaceous perithecia, and when mature these are ejected from the orifice at the apex, or are exposed by the breaking off of the upper portion of the perithecia. In _Diplodia_ and _Hendersonia_ the spores are larger, mostly coloured, often very fine in the latter genus, and multiseptate, escaping from the perithecia by a terminal pore. Probably the species are only pycnidia of _Sphæriacei_, but that is of no consequence in relation to our present inquiry. Of stylospores which deserve mention on account of their singularity of form, we may note those of _Dilophospora graminis_, which are straight, and have two or three hair-like appendages at each extremity. In _Discosia_ there is a single oblique bristle at each end, or at the side of the septate spores, whilst in _Neottiospora_ a tuft of delicate hairs is found at one extremity only. The appendages in _Dinemasporium_ are similar to those of _Discosia_. The spores in _Prosthemium_ may be said in some sort to resemble compound _Hendersonia_, being fusiform and multiseptate, often united at the base in a stellate manner. In this genus, as in _Darluca_, _Cytispora_, and the most of those belonging to the _Melanconiei_, the spores when mature are expelled from the orifice of the perithecium or spurious perithecium, either in the form of tendrils, or in a pasty mass. In these instances the spores are more or less involved in gelatine, and when expelled lie spread over the matrix, around the orifice; their ultimate diffusion being due to moisture washing them over other parts of the same tree, since it is probable that their natural area of dissemination is not large, the higher plants, of which they are mostly conditions, being developed on the same branches. More must be known of the relations between _Melanconium_ and Tulasne's sphæriaceous genus _Melanconis_ before we can appreciate entirely the advantage to _Melanconium_ and some other genera, that the wide diffusion of their spores should be checked by involving them in mucus, or their being agglutinated to the surface of the matrix, only to be softened and diffused by rain. The spores in many species amongst the _Melanconiei_ are remarkably fine; those of _Stegonosporium_ have the endochrome partite and cellular. In _Stilbospora_ and _Coryneum_ the spores are multiseptate, large, and mostly coloured. In _Asterosporium_ the spores are stellate, whilst in _Pestalozzia_ they are septate, with a permanent peduncle, and crested above with two or three hyaline appendages.

The _Torulacei_ externally, and to the naked eye, are very similar to the black moulds, and the mode of dissemination will be alike in both. The spores are chiefly compound, at first resembling septate threads, and at length breaking up into joints, each joint of which possesses the function of a spore. In some instances the threads are connate, side by side, as in _Torula hysterioides_, and in _Speira_, being concentrically arranged in laminæ in the latter genus. The structure in _Sporochisma_ is very peculiar, the joints breaking up within an external tube or membrane. The spores in _Sporidesmium_ appear to consist of irregular masses of cells, agglomerated into a kind of compound spore. Most of the species become pulverulent, and the spores are easily diffused through the air like an impalpable dust. They form a sort of link between the stylospores of one section of the _Coniomycetes_, and the pseudospores of the parasitical section.

PSEUDOSPORE is, perhaps, the most fitting name which can be applied to the so-called spores of the parasitical _Coniomycetes_. Their peculiar germination, and the production of reproductive bodies on the germ tubes, prove their analogy to some extent with the prothallus of other cryptogams, and necessitate the use of some term to distinguish them from such spores as are reproductive without the intervention of a promycelium. The differences between these pseudospores in the several genera are confined in some instances to their septation, in others to their mode of development. In the _Æcidiacei_ the pseudospores are more or less globose, produced in chains within an external cellular peridium. In the _Cæomacei_ they are simple, sometimes produced in chains, and sometimes free, with or without a caduceous peduncle. In the _Ustilaginei_ they are simple, dark coloured, and occasionally attached in subglobose masses, as in _Urocystis_ and _Thecaphora_, which, are more or less compact. In the _Pucciniæi_ the distinctive features of the genera are based upon the more or less complex nature of the pseudospores, which are bilocular in _Puccinia_, trilocular in _Triphragmium_, multilocular in _Phragmidium_, &c. In the curious genus _Podisoma_ the septate pseudospores are involved in a gelatinous element. The diffusion of these fruits is more or less complete according to their compact or pulverulent nature. In some species of _Puccinia_ the sori are so compact that they remain attached to the leaves long after they are dead and fallen. In the genus _Melampsora_, the wedge-shaped winter-pseudospores are not perfected until after the dead leaves have for a long time remained and almost rotted on the ground. It is probable that their ultimate diffusion is only accomplished by the rotting and disintegration of the matrix. In the _Cæomacei_, _Ustilaginei_, and _Æcidiacei_ the pseudospores are pulverulent, as in some species of _Puccinia_, and are easily diffused by the motion of the leaves in the wind, or the contact of passing bodies. Their diffusion in the atmosphere seems to be much less than in the case of the _Hyphomycetes_. By what means such a species as _Puccinia malvacearum_, which has very compact sori, has become within so short a period diffused over such a wide area, is a problem which in the present state of our knowledge must remain unsolved. It may be through minute and plentiful secondary spores.

SPERMATIA are very minute delicate bodies found associated with many of the epiphyllous _Coniomycetes_, and it has been supposed are produced in conjunction with some of the _Sphæriacei_, but their real function is at present obscure, and the name is applied rather upon conjecture than knowledge. It is by no means improbable that spermatia do exist extensively amongst fungi, but we must wait in patience for the history of their relationship.

TRICHOSPORES might be applied better, perhaps, than _conidia_ to the spores which are produced on the threads of the _Hyphomycetes_. Some of them are known to be the conidia of higher plants; but as this is by no means the case with all, it would be assuming too much to give the name of conidia to the whole. By whatever name they may be called, the spores of the _Hyphomycetes_ are of quite a different type from any yet mentioned, approximating, perhaps, most closely to the basidiospores of the _Hymenomycetes_ in some, and _Gasteromycetes_ in others; as, for instance, in the _Sepedoniei_ and the _Trichodermacei_. The form of the spores and their size differ materially, as well as the manner in which they are produced on the threads. In many they are very minute and profuse, but larger and less plentiful in the _Dematiei_ than in the _Mucedines_. The spores of some species of _Helminthosporium_ are large and multiseptate, calling to mind the spores of the _Melanconiei_. Others are very curious, being stellate in _Triposporium_, circinate in _Helicoma_ and _Helicocoryne_, angular in _Gonatosporium_, and ciliate in _Menispora ciliata_. Some are produced singly and some in chains, and in some the threads are nearly obsolete. In _Peronospora_, it has been demonstrated that certain species produce minute zoospores from the so-called spores. The dissemination of the minute spores of the _Mucedines_ through the air is undoubted; rain also certainly assists not only in the dispersion of the spores in this as in other groups, but also in the production of zoospores which require moisture for that purpose. The form of the threads, and the mode of attachment of the spores, is far more variable amongst the _Mucedines_ than the form of the spores, but the latter are in all instances so slightly attached to their supports as to be dissevered by the least motion. This aids also in the diffusion of the spores through the atmosphere.

SPORANGIA are produced in the _Physomycetes_ usually on the tips or branches of delicate threads, and these when mature dehisce and set free the minute sporidia. These are so small and uniform in their character that they require but a passing mention. The method of diffusion agrees much with that of the _Mucedines_, the walls of the sporangia being usually so thin and delicate as to be easily ruptured. Other modes of fructification prevail in some species by the production of cysts, which are the result of conjugation of the threads. These bodies are for the most part furnished with thicker and more resistant walls, and the diffusion of their contents will be regulated by other circumstances than those which influence the dispersion of the minute sporidia from the terminal cysts. Probably they are more perennial in their character, and are assimilated more to the oogonia of _Cystopus_ and _Peronospora_, being rather of the nature of resting spores, inasmuch as the same threads usually bear the terminal fruits.

THECASPORES is a term which may be applied generally to all sporidia produced in asci, but these are in turn so innumerable and variable that it will be necessary to treat of some of the groups individually. The _Thecaspores_, for instance, of the _Tuberacei_ offer several features whereby they may be distinguished from other thecaspores. The asci in which these sporidia are generated mostly partake of a broadly saccate, ovate form. The number of sporidia contained in an individual ascus is usually less than in the majority of the _Ascomycetes_, and the sporidia approximate more nearly to the globose form. Usually, also, they are comparatively large. Many have been figured by Corda[C] and Tulasne.[D] Three types of spores may be said to prevail in the _Tuberacei_: the smooth spored, the warted or spinulose, and the areolate. The first of these may be represented by the _Stephensia bombycina_, in which the globose sporidia are quite smooth and colourless. The warted sporidia may be observed in _Genea verrucosa_, the spinulose in _Tuber nitidum_, and the areolate are present in _Tuber æstivum_ and _Tuber excavatum_, in which the epispore is divided into polygonal alveoli, bounded by thin, membranaceous, prominent partitions. This form of sporidium is very beautiful. In all no special provision is made for the dissemination of the sporidia, as, from their subterranean habit, none would be available save the ultimate dissolution of the external integuments. As they are greedily devoured by several animals, it is possible that they may be dispersed through the excrements.

In the _Perisporiacei_ the perithecium has no proper orifice, or ostiolum, for the discharge of the mature sporidia, which are usually small, and are disseminated by the irregular rupture of the somewhat fragile conceptacles. The asci are usually more or less saccate, and the sporidia approximate to a globose form. The asci are often very diffluent. In _Perisporium vulgare_ the ovate brown sporidia are at first, and for some time, attached together in fours in a concatenate or beaded manner. In some species of _Erysiphei_ the conceptacle encloses but a single sporangium, in others several, which are attached together at the base. In some species the sporangia contain two, in others four, in others eight, and in others numerous sporidia. In _Chætomium_ the asci are cylindrical, and in most cases the coloured sporidia are lemon-shaped. When the conceptacles are fully matured, it is commonly the case that the asci are absorbed and the sporidia are free in the interior of the conceptacles.

Of the fleshy _Discomycetes_ the genus _Peziza_ may be taken as the type. If the structure which prevails in this genus be brought to mind, it will be remembered that the hymenium lines an expanded cup, and that the asci are packed together, side by side, with their apices outwards, and their bases attached to a substratum of cells which form the inner layer of the receptacle. The sporidia are usually eight in each ascus, either arranged in single or double rows, or irregularly grouped together. The asci are produced in succession; the later, pressing themselves upwards between those previously developed, cause the rupture of the mature asci at the apex and the ejection of the sporidia with considerable force. When a large _Peziza_ is observed for a time a whitish cloud will be seen to rise suddenly from the surface of the disc, which is repeated again and again whenever the specimen is moved. This cloud consists of sporidia ejected simultaneously from several asci. Sometimes the ejected sporidia lie like frost on the surface of the disc. Theories have been devised to account for this sudden extrusion of the sporidia, in _Ascobolus_, and a few species of _Peziza_, of the asci also, the most feasible one being the successive growth of the asci; contraction of the cup may also assist, as well as some other less potent causes. It may be remarked here that the sporidia in _Peziza_ and _Helotium_ are mostly colourless, whilst in _Ascobolus_ they pass through pink to violet, or dark brown, and the epispore, which is of a waxy nature, becomes fissured in a more or less reticulated manner.

The sporidia in _Hysterium_ proper are usually coloured, often multiseptate, sometimes fenestrate, and occasionally of considerable size. There is no evidence that the sporidia are ever excluded in the same manner as in _Peziza_, the lips closing over the disc so much as to prevent this. The diffusion of the sporidia probably depends on the dissolution of the asci, and hence they will not be widely dispersed, unless, perhaps, by the action of rain.

In _Tympanis_, asci of two kinds have been observed in some species; one kind containing an indefinite number of very minute bodies resembling spermatia, and the other octosporous, containing sporidia of the usual type.

The _Sphæriacei_ include an almost infinite variety in the form and character of the sporidia. Some of these are indefinite in the number contained in an ascus, although the majority are eight, and a few less. In the genera _Torrubia_ and _Hypocrea_ the structure differs somewhat from other groups, inasmuch as in the former the long thread-like sporidia break up into short joints, and in the latter the ascus contains sixteen subglobose or subquadrate sporidia. Other species contain linear sporidia, which are often the length of the ascus, and may either be simple or septate. In _Sphæria ulnaspora_ the sporidia are abruptly bent at the second joint. Shorter fusiform sporidia are by no means uncommon, varying in the number of septa, and in constriction at the joints in different species. Elliptic or ovate sporidia are common, as are those of the peculiar form which may be termed sausage-shaped. These are either hyaline or coloured of some shade of brown. Coloured sporidia of this kind are common in _Xylaria_ and _Hypoxylon_, as well as in certain species of the section _Superficiales_. Coloured sporidia are often large and beautiful: they are mostly of an elongated, elliptical form, or fusiform. As noteworthy may be mentioned the sporidia of _Melanconis lanciformis_, those of _Valsa profusa_, and some species of _Massaria_, the latter being at first invested with a hyaline coat. Some coloured sporidia have hyaline appendages at each extremity, as in _Melanconis Berkeleii_, and an allied species, _Melanconis bicornis_, from the United States, also some dung _Sphæriæ_, as _S. fimiseda_, included under the proposed genus _Sordaria_.[E] Hyaline sporidia occasionally exhibit a delicate bristle-like appendage at each extremity, as in the _Valsa thelebola_, or with two additional cilia at the central constriction, as in _Valsa taleola_. A peculiar form of sporidium is present in certain species of _Sphæria_ found on dung, for which the generic name of _Sporormia_ has been proposed, in which the sporidium (as in _Perisporium vulgare_) consists of four coloured ovate joints, which ultimately separate. Multiseptate fenestrate sporidia are not uncommon in _Cucurbitaria_ and _Pleospora_, as well as in _Valsa fenestrata_ and some other species. In the North American _Sphæria putaminum_ the sporidia are extraordinarily large.

The dissemination of the sporidia may, from identity of structure in the perithecium, be deemed to follow a like method in all. When mature, they are in a great measure expelled from the mouth of the perithecia, as is evident in species with large dark sporidia, such as exist in the genera _Hypoxylon_, _Melanconis_, and _Massaria_. In these genera the sporidia, on maturity, may be observed blackening the matrix round the mouths of the perithecia. As moisture has an evident effect in producing an expulsion of sporidia by swelling the gelatinous nucleus, it may be assumed that this is one of the causes of expulsion, and therefore of aids to dissemination. When _Sphæriæ_ are submitted to extra moisture, either by placing the twig which bears them on damp sand, or dipping one end in a vessel of water, the sporidia will exude and form a gelatinous bead at the orifice. There may be other methods, and possibly the successive production of new asci may also be one, and the increase in bulk by growth of the sporidia another; but of this the evidence is scanty.

Finally, OOGONIA may be mentioned as occurring in such genera as _Peronospora_ amongst moulds, _Cystopus_ amongst Uredines, and the _Saprolegniaceæ_ amongst the _Physomycetes_. The zoospores being furnished with vibratile cilia, are for some time active, and need only water in which to disseminate themselves, and this is furnished by rain.

We have briefly indicated the characteristics of some of the more important types of spores to be found in fungi, and some of the modes by which it is known, or presumed, that their dissemination takes place. In this summary we have been compelled to rest content with suggestions, since an exhaustive essay would have occupied considerable space. The variability in the fruit of fungi, in so far as we have failed to demonstrate, will be found exhibited in the illustrated works devoted more especially to the minute species.[F]

[A] Cunningham, in "Ninth Annual Report of the Sanitary Commissioner
with the Government of India." Calcutta, 1872.

[B] See "Corda Icones," tab. 2.

[C] Corda, "Icones Fungorum," vol. vi. Prague.

[D] Tulasne, "Fungi Hypogæi." Paris.

[E] Winter, "Die Deutschen Sordarien" (1873).

[F] Corda, "Icones Fungorum," 6 vols. (1837-1842); Sturm,
"Deutschlands Flora," Pilze (1841); Tulasne, "Selecta Fungorum
Carpologia;" Bischoff, "Kryptogamenkunde" (1860); Corda,
"Anleitung zum Studium der Mykologie" (1842); Fresenius,
"Beiträge zur Mykologie" (1850); Nees Ton Esenbeck, "Das System
der Pilze" (1816); Bonorden, "Handbuch der Allgemeinen
Mykologie" (1851).

VII.

GERMINATION AND GROWTH.

In describing the structure of these organisms in a previous chapter, the modes of germination and growth from the spores have been purposely excluded and reserved for the present. It may be assumed that the reader, having followed us to this point, is prepared for our observations by some knowledge of the chief features of structure in the principal groups, and of the main distinctions in the classification, or at least sufficient to obviate any repetition here. In very many species it is by no means difficult to induce germination of the spores, whilst in others success is by no means certain.