The existence of some sort of sexual reproduction in Fungi has long been suspected, although in earlier instances upon insufficient grounds; but of late years observations have multiplied and facts accumulated which leave no doubt of its existence. If the SaprolegniÆ are left out of the question as disputed Fungi, there still remain a number of well authenticated instances of the phenomena of copulation, and many other facts which indicate some sort of sexual relationship. The precise manner in which those minute bodies, so common amongst the SphÆronemei, which we prefer to call stylospores, perform their functions is still to a great extent a mystery; yet it is no longer doubted that certain species of AposphÆria, Phoma, Septoria, &c., are only conditions of some species of SphÆria, often developed and matured in close proximity to them on the same host. In Æcidium, Roestelia, &c., spermogonia are produced plentifully on or near the same spots on which the fructification appears, either simultaneously or at a later period. We know not whether any importance is to be attached to the Fig. 95. In Phycomyces the organs of reproduction have been subjected to close examination by Van Tieghem, The germination of the zygospores in this species has not as yet been observed, but it is probably the same or very similar to that observed in other species of Mucor. In these the rough tuberculate epispore splits on one side, and its internal coat elongates itself and protrudes as a tube filled with protoplasm and oil globules, terminating in an ordinary sporangium. Usually the amount of nutriment contained in the zygospore is exhausted by the formation of the terminal sporangium, according to Brefeld; Fig. 96. De Bary has given a precise account of the formation of the zygospore in another of the Mucors, Rhizopus nigricans, in which he says that the filaments which conjugate are solid rampant tubes, which are branched without order and confusedly intermingled. Where two of these filaments meet each of them pushes towards the other an appendage which is at first cylindrical and of the same diameter. From the first these two processes are applied firmly one to the other by their extremities; they increase in size, become clavate, and constitute together a fusiform body placed across the two conjugated filaments. Between the two halves of this body there exists no constant difference of size; often they are both perfectly equal. In each Similar conjugation with like results also takes place in Syzygites megalocarpus. In this species the germination of the zygospores has been observed. If, after a certain time of repose, these bodies are placed on a moist substratum, they emit a germ-like tube, which, without originating a proper mycelium, develops at the expense of the nutritive material stored in the zygospore into a carpophore or fruit bearer, which is many times dichotomously branched, bearing terminal sporangia characteristic of the species. It has already been remarked by us that the Saprolegnei are claimed by some authors as AlgÆ, whilst we are more disposed to regard them as closely allied to the Mucors, and as they exhibit in themselves strong evidence in support of the existence of sexual reproduction, we cannot forbear giving a summary of what has been observed by De Bary and others in this very interesting and singular group of plants, to which M. Cornu has recently dedicated an exhaustive monograph. In Saprolegnia monoica, and others, the female organs consist of oogonia—that is to say, of cells which are at first globose and rich in plastic matter, which most generally terminate short branches of the mycelium, and which are rarely seen in an interstitial position. The constitutive membrane of the adult oogonia is reabsorbed in a great many points, and is there pierced with rounded holes. At the same time the plasma is divided into a larger or smaller number of distinct portions, which are rounded into little spheres, and separate from the Fig. 97. During the formation of the oogonia there arise from its pedicel or from neighbouring filaments slight cylindrical curved branches, sometimes turned round the support of the oogonia, and which all tend towards this organ. Their superior extremity is intimately applied to its wall, then ceases to be elongated, becomes slightly inflated, and is limited below by a partition; it is then an oblong cell, slightly curved, filled with protoplasm, and intimately applied to the oogonia—in fact, an antheridium or organ of the male sex. Each oogonium possesses one or several antheridia. Towards the time when the gonospheres are formed it may be observed that each antheridium sends to the interior of the oogonia one or several tubular processes, which have crossed its side wall, and which open at their extremity in order to discharge their contents. These, while they are flowing out, present some very agile corpuscles, and which, considering their resemblance to those in Vaucheria, to which the name of spermatozoids are applied, ought to be considered as the fecundating corpuscles. After the evacuation of the antheridia the gonospheres are found to be covered with cellulose; they then constitute so many oospores, with solid walls. De Bary considers that, bearing in mind analogous phenomena observed in Vaucheria, and the direct observations of Pringsheim, In Achlya dioica the antheridium is cylindrical, the plasma which it encloses is divided into particles, which attain nearly Amongst obscure and doubtful bodies are those described by Pringsheim, which have their origin in thick filaments or tubes, similar to those which form the zoosporangia, and represent so many distinct little masses of plasma within an homogeneous parietal ganglion. The contour of these plastic masses is soon delineated in a more precise manner. We see in their interior some homogeneous granules, which are at first globose, then oval, and finally travel to the enlarged and ampullÆform extremity of the generating tube. There they become rounded or oval cells covered with cellulose, and emit from their surface one or several cylindrical processes, which elongate towards the wall of the conceptacle, and pierce it, without, however, ever projecting very far beyond it. At the same time the lacunose protoplasm of each cell becomes divided into a number of corpuscles, which escape by the open extremity of the cylindrical neck. They resemble in their organization and agility the spermatozoids of Achlya dioica. They soon become motionless in water, and do not germinate. During the development of these organs, the protoplasm of the utricle which contains them offers at first completely normal characteristics, and disappears entirely by degrees as they increase. De Bary and Pringsheim believe that these organs constitute the antheridia of the species of Saprolegnia to which they belong. The oospores of the SaprolegniÆ, when arrived at maturity, possess a tolerably thick double integument, consisting of an epispore and an endospore. After a considerable time of repose De Bary has claimed for the oogonia in Cystopus and Peronospora a kind of fecundation which deserves mention here. Fig. 98. The branches of the mycelium which do not bear oogonia apply their obtuse extremities against the growing oogonia; this extremity swells, and, by a transverse partition, separates itself from the supporting tube. It is the antheridium, or male organ, which is formed by this process; it takes the form of an obliquely clavate or obovate cellule, which is always considerably smaller than the oogonium, and adheres to its walls by a plane or convex area. The slightly thickened membrane of the antheridia encloses protoplasm which is finely granular. It is seldom that more than one antheridium applies itself to an oogonium. The two organs having together achieved their development, the large granules contained in the oogonium accumulate at its centre to group themselves under the form of an irregular globule deprived of a proper membrane, and surrounded by a bed of almost homogeneous protoplasm. This globule is the gonosphere, or reproductive sphere, which, through the means of Fig. 99. Considering the great resemblance of these organs with the sexual organs of the SaprolegniÆ, which are closely allied to the AlgÆ, and of which the sexuality has been proved, De Bary adds, we have no doubt whatever that the phenomena just described represent an act of fecundation, and that the tube pushed out by the antheridium should be regarded as a fecundating tube. It is remarkable that amongst these fungi the tube projected by the antheridium effects fecundation only by contact. Its extremity never opens, and we never find antherozoids; on the contrary, the antheridium presents, up to the maturity of the oospore, the appearance which it presented at the moment of fecundation. The primitive membrane of the oospore, at first very thin, soon acquires a more sensible thickness, and becomes surrounded by an external layer (epospore), which is formed at the expense of the protoplasm of the periphery. This disappears in proportion as the epispore attains maturity, and finally there only remains a quantity of granules, suspended in a transparent watery fluid. At the period of maturity, the epispore is a slightly thickened, resistant membrane, of a yellowish-brown colour, and finely punctate. The surface is almost always provided with brownish warts, which are large and obtuse, sometimes isolated, and sometimes confluent, forming irregular crests. These warts are composed of cellulose, which reagents colour of a deep blue, whilst the membrane which bears them preserves its primitive colour. Amongst the principal observers of certain phenomena of copulation in cells formed in the earliest stages of the Discomycetes are Professor de Bary, Tulasne observes that this “scolecite” or ringed body can be readily isolated in Ascobolus furfuraceus. When the young receptacles In Peziza melanoloma, A. and S., the same observer succeeded still better in his searches after the scolecite, which he remarks is in this species most certainly a lateral branch of the filaments of the mycelium. This branch is isolated, simple, or forked at a short distance from its base, and in diameter generally exceeding that of the filament which bears it. This branch is soon arcuate or bent, and often elongated in describing a spiral, the irregular turns of which are lax or compressed. At the same time its interior, at first continuous, becomes divided by transverse septa into eight or ten or more cells. Sometimes this special branch terminates in a crozier shape, which is involved in the bent part of another crozier which terminates a neighbouring filament. In other cases the growing branch is connected, by its extremity, with that of a hooked branch. These contacts, however, did not appear to Tulasne to be so much normal as accidental. But of the importance of the ringed body, or “scolecite,” there was no room for doubt, as being the certain and habitual rudiment of the fertile cup. In fact, inferior cells are produced from the flexuous filaments which creep about its surface, cover and surround it on all sides, while joining themselves to each other. At first continuous, then septate, these cells by their union constitute a cellular tissue, which increases little by little until the scolecite is so closely enveloped that only its superior extremity can be seen. These cellular masses attain a considerable volume before the hymenium begins to show itself in a depression of their summit. So long as their smallness permits of their being seen in the field of the microscope, it can be determined that they adhere to a single filament of the mycelium by the base of the scolecite which remains naked. Fig. 100. Although Tulasne could not satisfy himself of the presence of any act of copulation in Ascobolus furfuraceus, or Peziza Fig. 100a. Certain phenomena concerned in the development of the Erysiphei belong also to this connection. The mycelium of Erysiphe cichoracearum, like that of other species, consists of branched filaments, crossed in all directions, which adhere as they climb to the epidermis of the plant on which the fungus lives as a parasite. The perithecia are engendered where two filaments cross each other. These swell slightly at this point, and each emits a process which imitates a nascent branch, and remains upright on the surface of the epidermis. The process originating from the inferior filament soon acquires an oval form and a diameter double that of the filament; then it becomes isolated from it by a septum, and constitutes a distinct cell, which De Bary During this time the oocyst enlarges and divides, without its being possible precisely to determine the way in which it happens, into a central cell and an outer layer, ordinarily simple, of smaller cells, contiguous to the general enveloping wall. The central cell becomes the single ascus, which is characteristic of the species, and the layer which surrounds it constitutes the inner wall of its perithecium. The only changes afterwards observed are the increase in size of the perithecium, the production of the root-like filaments which proceed from its outer wall, the brown tint which it assumes, and finally the formation of the sporidia in the ascus. The antheridium remains for a long time recognizable without undergoing any essential modification, but the dark colour of the perithecium soon hides it from the observer’s eye. De Bary thinks that he is authorized in assuming the probability that the conceptacles and organs of fructification of others of the Ascomycetes, including the Discomycetes and the Tuberacei, are the results of sexual generation. Certain phenomena which have been observed amongst the Coniomycetes are cited as examples of sexual association. Amongst these may be named the conjugation of the slender spores of the first generation, produced on the germinating Fig. 101. Finally, the spermogonia must be regarded as in some occult manner, which as yet has baffled detection, influencing the perfection of sporidia Several of the species of Hysterium also possess spermogonia, notably H. Fraxini, which may be distinguished from the ascigerous perithecia with which they are associated by their smaller size and flask-like shape. From these the spermatia are expelled long before the maturity of the spores. In Hypoderma virgultorum, H. commune, and H. scirpinum, the spermogonia are small depressed black capsules, which contain an abundance of minute spermatia. These were formerly regarded as distinct species, under the name of Leptostroma. In Stictis ocellata a great number of the tubercles do not pass into the perfect state Again, there is a very common fungus which forms black discoid spots on dead holly leaves, called Ceuthospora phacidioides, figured by Greville in his “Scottish Cryptogamic Flora,” which expels a profusion of minute stylospores; but later in the season, instead of these, we find the asci and sporidia of Phacidium ilicis, so that the two are forms and conditions the one of the other. In Tympanis conspersa the spermogonia are much more commonly met with than the complete fruit. There is a great external resemblance in them to the ascigerous cups, but there is no evidence that they are ever transformed into such. The perfect sporidia are also very minute and numerous, being contained in asci borne in cups, which usually surround the spermogonia. In several species of Dermatea the stylospores and spermatia co-exist, but they are disseminated before the appearance of the ascigerous receptacles, yet they are produced upon a common stroma not unlike that of Tubercularia. In its early stage the common and well-known Bulgaria inquinans, which when mature looks like a black Peziza, is a little tubercle, the whole mass of which is divided into ramified lobes, the extremities of which become, towards the surface of the tubercle, receptacles from whence escape waves of spermatia which are colourless, or stylospores mixed with them which are larger and nearly black. Amongst the SphÆriacei numerous instances might be cited of minute stylosporous bodies in consort with, or preceding, the ascigerous receptacles. A very familiar example may be found at the base of old nettle stems in what has been named AposphÆria acuta, but which truly are only the stylospores of the SphÆria coniformis, the perithecia of which flourish in company or in close proximity to them. Most of these bodies are so minute, delicate, and hyaline that the difficulties in the way Professor de Bary cautiously refrains from accepting spermatia other than as doubtful or at least uncertain sexual bodies. Many organs placed at first amongst spermatia have been recognized by M. Tulasne as being themselves susceptible of germination, and consequently ought to take their place among legitimate spores. Then it must be considered that very many spores can only germinate under certain conditions. It is, therefore, for the present a doubtful question whether there exist really any spermatia incapable of germination, or if the default of germination of these corpuscles does not rather depend on the experiments hitherto attempted not having included the conditions required by the phenomena. Moreover, as yet no trace has been discovered of the female organs which are specially fecundated by the spermatia. Finally, there exist in the Ascomycetes certain organs of reproduction, diverse spore-bearing apparatus, pycnidia, and others, which, like the spermogonia, usually precede ascophorous fruits. The real nature of the spermogonia and spermatia should therefore be regarded as, at present, very uncertain; as regards, however, the spermatia which have never been seen to germinate, perhaps it is as well not to absolutely reject the first opinion formed concerning them, or M. Tulasne has devoted a chapter to the spermogonia of the Uredines in his memoir, to which we have already alluded. Œersted, in “Verhandl der KÖnig. DÄn. Gesell. Der Wissensch,” 1st January, 1865; De Bary, “Handbuch der Physiol. Botanik” (1866), p. 172; “Annales des Sci. Nat.” (5me sÉr.), vol. v. (1866), p. 366. De Bary, “Morphologie und Physiologie der Pilze,” cap. 5, p. 160; “Ann. des Sci. Nat.” (1866), p. 343. De Bary, in “Annales des Sciences Naturelles” (5me sÉr.), vol. v. (1866), p. 343; Hoffmeister’s “Handbook” (Fungi), cap. v. p. 155. Tulasne, “On the Phenomena of Copulation in certain Fungi,” in “Ann. des Sci. Nat.” (1866), p. 211. Berkeley, in “Journ. Hort. Soc.” vol ii. p. 107; Tulasne, “Ann. d. Sc. Nat.” (4me sÉr.), vol. ii. tab. 12. Tulasne, “New Researches on the Reproductive Apparatus of Fungi;” “Comptes Rendus,” vol. xxxv. (1852), p. 841. |