Saprolegnia, Achlya, Pythiopsis, Dictyuchus, Aplanes. Motile zoospores
which escape from the zoosporangium are present except in Aplanes.
The sexual reproduction shows all transitions between forms which
are normally sexual, like the Peronosporaceae, to forms in which
no antheridium is developed and the oospheres develop parthenogenetically.
The oogonia, unlike the Peronosporaceae, contain more
than one oosphere. Klebs has shown that the development of
zoosporangia or of oogonia and pollinodia respectively in Saprolegnia
is dependent on the external conditions; so long as a continued
stream of suitable food-material is ensured the mycelium grows on
without forming reproductive organs, but directly the supplies of
nitrogenous and carbonaceous food fall below a certain degree of
concentration sporangia are developed. Further reduction of the
supplies of food effects the formation of oogonia. This explains the
sequence of events in the case of a Saprolegnia-mycelium radiating
from a dead fly in water. Those parts nearest the fly and best
supplied develop barren hyphae only; in a zone at the periphery,
where the products of putrefaction dissolved in the water form a
dilute but easily accessible supply, the zoosporangia are developed
in abundance; oogonia, however, are only formed in the depths of
this radiating mycelium, where the supplies of available food
materials are least abundant.
Chytridineae.—These parasitic and minute, chiefly aquatic, forms may be looked upon as degenerate Oomycetes, since a sexual process and feeble unicellular mycelium occur in some; or they may be regarded as series of primitive forms leading up to higher members. There is no means of deciding the question. They are usually included in Oomycetes, but their simple structure, minute size, usually uniciliate zoospores, and their negative characters would justify their retention as a separate group. It contains less than 200 species, chiefly parasitic on or in algae and other water-plants or animals, of various kinds, or in other fungi, seedlings, pollen and higher plants. They are often devoid of hyphae, or put forth fine protoplasmic filaments into the cells of their hosts. After absorbing the cell-contents of the latter, which it does in a few hours or days, the fungus puts out a sporangium, the contents of which break up into numerous minute swarm-spores, usually one-ciliate, rarely two-ciliate. Any one of these soon comes to rest on a host-cell, and either pierces it and empties its contents into its cavity, where the further development occurs (Olpidium), or merely sends in delicate protoplasmic filaments (Rhizophydium) or a short hyphal tube of, at most, two or three cells, which acts as a haustorium, the further development taking place outside the cell-wall of the host (Chytridium). In some cases resting spores are formed inside the host (Chytridium), and give rise to zoosporangia on germination. In a few species a sexual process is described, consisting in the conjugation of similar cells (Zygochytrium) or the union of two dissimilar ones (Polyphagus). In the development of distinct antheridial and oogonial cells the allied Ancylistineae show close alliances to Pythium and the Oomycetes. On the other hand, the uniciliate zoospores of Polyphagus have slightly amoeboid movements, and in this and the pseudopodium-like nature of the protoplasmic processes, such forms suggest resemblances to the Myxomycetes. Opinions differ as to whether the Chytridineae are degraded or primitive forms, and the group still needs critical revision. Many new forms will doubtless be discovered, as they are rarely collected on account of their minuteness. Some forms cause damping off of seedlings—e.g. Olpidium Brassicae; others discoloured spots and even tumour-like swellings—e.g. Synchytium Scabiosae, S. Succisae, Urophlyctis, &c., on higher plants. Analogies have been pointed out between Chytridiaceae and unicellular algae, such as Chlorosphaeraceae, Protococcaceae, “Palmellaceae,” &c., some of which are parasitic, and suggestions may be entertained as to possible origin from such algae.
The Zygomycetes, of which about 200 species are described, are especially important from a theoretical standpoint, since they furnished the series whence Brefeld derived the vast majority of the fungi. They are characterized especially by the zygospores, but the asexual organs (sporangia) exhibit interesting series of changes, beginning with the typical sporangium of Mucor containing numerous endospores, passing to cases where, as in Thamnidium, these are accompanied with more numerous small sporangia (sporangioles) containing few spores, and thence to Chaetocladium and Piptocephalis, where the sporangioles form but one spore and fall and germinate as a whole; that is to say, the monosporous sporangium has become a conidium, and Brefeld regarded these and similar series of changes as explaining the relation of ascus to conidium in higher fungi. According to his view, the ascus is in effect the sporangium with several spores, the conidium the sporangiole with but one spore, and that not loose but fused with the sporangiole wall. On this basis, with other interesting morphological comparisons, Brefeld erected his hypothesis, now untenable, that the Ascomycetes and Basidiomycetes diverge from the Zygomycetes, the former having particularly specialized the ascus (sporangial) mode of reproduction, the latter having specialized the conidial (indehiscent one-spored sporangiole) mode. In addition to sporangia and the conidial spores referred to, some Mucorini show a peculiar mode of vegetative reproduction by means of gemmae or chlamydospores—i.e. short segments of the hyphae become stored with fatty reserves and act as spores. The gemmae formed on submerged Mucors may bud like a yeast, and even bring about alcoholic fermentation in a saccharine solution.
|
From Strasburger’s Lehrbuch der Botanik, by permission of Gustav Fischer.
|
| Fig. 6.—Mucor Mucedo. Different stages in the formation and germination of the zygospore. (After Brefeld, 1-4. 5 from v. Tavel, Pilze.) 1, Two conjugating branches in contact. 2, Septation of the conjugating cells (a) from the suspensors (b).
3, More advanced stage, the conjugating cells (a) are still distinct from one another; the warty thickenings of their walls have commenced to form.
4, Ripe zygospore (b) between the suspensors (a).
5, Germinating zygospore with a germ-tube bearing a sporangium.
|
The segments of the hyphae in this group usually contain several nuclei. At the time of sporangial formation the protoplasm with numerous nuclei streams into the swollen end of the sporangiophore and there becomes cut off by a cell-wall to form the sporangium. The protoplasm then becomes cut up by a series of clefts into a number of smaller and smaller pieces which are unicellular in Pilobolus, multicellular in Sporodinia. These then become surrounded by a cell-wall and form the spores. This mode of spore-formation is totally different from that in the ascus; hence one of the difficulties of the acceptance of Brefeld’s view of the homology of ascus and sporangium. The cytology of zygospore-formation is not known in detail; the so-called gametes which fuse are multinucleate and are no doubt of the nature of gametangia. The fate of these nuclei is doubtful, probably they fuse in pairs (fig. 6).
Blakeslee has lately made some very important observations of the Zygomycetes. It is well known that while in some forms, e.g. Spordinia, zygospores are easily obtained, in others, e.g. most species of Mucor, they are very erratic in their appearance. This has now been explained by Blakeslee, who finds that the Mucorinae can be divided into two groups, termed homothallic and heterothallic respectively. In the first group zygospores can arise by the union of branches from the same mycelium and so can be produced by the growth from a single spore; this group includes Spordinia grandis, Spinellus fusiger, some species of Mucor, &c. The majority of forms, however, fall into the heterothallic group, in which the association of branches from two mycelia different in nature is necessary for the formation of zygospores. These structures cannot then be produced from the product of a single spore nor even from the thalli derived from any two spores. The two kinds of thalli Blakeslee considers to have a differentiation of the nature of sex and he distinguishes them as (+) and (−) forms; the former being usually distinguished by a somewhat greater luxuriance of growth.
The classification of the Mucorini depends on the prevalence and characters of the conidia, and of the sporangia and zygospores—e.g. the presence or absence of a columella in the former, the formation of an investment round the latter. Most genera are saprophytes, but some—Chaetocladium, Piptocephalis—are parasites on other Mucorini, and one or two are associated casually with the rotting of tomatoes and other fruits, bulbs, &c., the fleshy parts of which are rapidly destroyed if once the hyphae gain entrance. Even more important is the question of mycosis in man and other animals, referred to species of Mucor, and investigated by Lucet and Costantin. Klebs has concluded that transpiration is the important factor in determining the formation of sporangia, while zygote-development depends on totally different conditions; these results have been called in question by Falck.
The Entomophthoraceae contain three genera, Empusa, Entomophthora and Basidiobolus. The two first genera consist of forms which are parasitic on insects. Empusa Muscae causes the well-known epidemic in house-flies during the autumn; the dead, affected flies are often found attached to the window surrounded by a white halo of conidia. B. ranarum is found in the alimentary canal of the frog and growing on its excrement. In these three genera the conidia are cast off with a jerk somewhat in the same way as the sporangium of Pilobolus.
