B. Higher Fungi.—Now that Brefeld’s view of the origin of these forms from the Zygomycetes has been overthrown, the relationship of the higher and lower forms of fungi is left in obscurity. The term Eumycetes is sometimes applied to this group to distinguish them from the Phycomycetes, but as the same name is also applied to the fungi as a whole to differentiate them from the Mycetozoa and Bacteria, the term had best be dropped. The Higher Fungi fall into three groups: the Ustilaginales, of doubtful position, and the two very sharply marked groups Basidiales and Ascomycetes.
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From Vine’s Students’ Text Book of Botany, by permission of Swan Sonnenschein & Co.
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| Fig. 7.—Germinating resting-gonidia. A, of Ustilago receptaculorum; B, of Tilletia Caries.
sp, The gonidium. d, The sporidia: in B the sporidia have coalesced in pairs at v.
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I. Ustilaginales.—This includes two families Ustilaginaceae (smuts) and Tilletiaceae (bunts). The bunts and smuts which damage our grain and fodder plants comprise about 400 species of internal parasites, found in all countries on herbaceous plants, and especially on Monocotyledons. They are remarkable for their dark spores developed in gall-like excrescences on the leaves, stems, &c., or in the fruits of the host. The discovery of the yeast-conidia of these fungi, and their thorough investigation by Brefeld, have thrown new lights on the group, as also have the results elucidating the nature of the ordinary dark spores—smuts, bunt, &c.—which by their mode of origin and development are chlamydospores. When the latter germinate a slender “promycelium” is put out; in Ustilago and its allies this is transversely septate, and bears lateral conidia (sporidia); in Tilletia and its allies non-septate, and bears a terminal tuft of conidia (sporidia) (fig. 7). Brefeld regarded the promycelium as a kind of basidium, bearing lateral or terminal conidia (comparable to basidiospores), but since the number of basidiospores is not fixed, and the basidium has not yet assumed very definite morphological characters, Brefeld termed the group Hemibasidii, and regarded them as a half-way stage in the evolution of the true Basidiomycetes from Phycomycetes, the Tilletia type leading to the true basidium (Autobasidium), the Ustilago type to the protobasidium, with lateral spores; but this view is based on very poor evidence, so that it is best to place these forms as a separate group, the Ustilaginales. The yeast-conidia, which bud off from the conidia or their resulting mycelium when sown in nutrient solutions, are developed in successive crops by budding exactly as in the yeast plant, but they cannot ferment sugar solutions. It is the rapid spread of these yeast-conidia in manure and soil waters which makes it so difficult to get rid of smuts, &c., in the fields, and they, like the ordinary conidia, readily infect the seedling wheat, oats, barley or other cereals. Infection in these cases occurs in the seedling at the place where root and shoot meet, and the infecting hypha having entered the plant goes on living in it and growing up with it as if it had no parasitic action at all. When the flowers form, however, the mycelium sends hyphae into the young ovaries and rapidly replaces the stores of sugar and starch, &c., which would have gone to make the grain, by the soot-like mass of spores so well known as smut, &c. These spores adhere to the grain, and unless destroyed, by “steeping” or other treatment, are sown with it, and again produce sporidia and yeast-conidia which infect the seedlings. In other species the infection occurs through the style of the flower, but the fungus after reaching the ovule develops no further during that year but remains dormant in the embryo of the seed. On germination, however, the fungus behaves in the same way as one which has entered in the seedling stage. The cytology of these forms is very little known; Dangeard states that there is a fusion of two nuclei in the chlamydospore, but this requires confirmation. Apart from this observation there is no other trace of sexuality in the group.
II. Ascomycetes.—This, except in the case of a few of the simpler forms, is a very sharply marked group characterized by a special type of sporangium, the ascus. In the development of the ascus we find two nuclei at the base which fuse together to form the single nucleus of the young ascus. The single nucleus divides by three successive divisions to form eight nuclei lying free in the protoplasm of the ascus. Then by a special method, described first by Harper, a mass of protoplasm is cut out round each nucleus; thus eight uninucleate ascospores are formed by free-cell formation. The protoplasm remaining over is termed epiplasm and often contains glycogen (fig. 8). In some cases nuclear division is carried further before spore-formation occurs, and the number of spores is then 16, 32 and 64, &c.; in a few cases the number of spores is less than eight by abortion of some of the eight nuclei. The ascus is thus one of the most sharply characterized structures among the fungi.
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From Strasburger’s Lehrbuch der Botanik, by permission of Gustav Fischer.
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| Fig. 8.—Development of the Ascus. | |
| A–C, | Pyronema confluens. (After Harper.) |
| D, | Young ascus of Boudiera with eight spores. (After Claussen.) |
In some forms we find definite male and female sexual organs (Sphaerotheca, Pyronema, &c.), in others the antheridium is abortive or absent, but the ascogonium (oogonium) is still present and the female nuclei fuse in pairs (Lachnea stercorea, Humaria granulata, Ascobolus furfuraceus); while in other forms ascogonium and antheridium are both absent and fusion occurs between vegetative nuclei (Humaria rutilans, and probably the majority of other forms). In other cases the sexual fusion is apparently absent altogether, as in Exoascus. In the first case (fig. 9) we have a true sexual process, while in the second and third cases we have a reduced sexual process in which the fusion of other nuclei has replaced the fusion of the normal male and female nuclei. It is to be noted that all the forms exhibit the fusion of nuclei in the ascus, so that those with the normal or reduced sexual process described above have two nuclear fusions in their life-history. The advantage or significance of the second (ascus) fusion is not clearly understood.
The group of the Hemiasci was founded by Brefeld to include forms which were supposed to be a connecting link between Phycomycetes and Ascomycetes. As mentioned before, the connexion between these two groups is very doubtful, and the derivation of the ascus from an ordinary sporangium of the Zygomycetes cannot be accepted. The majority of the forms which were formerly included in this group have been shown to be either true Phycomycetes (like Ascoidea) or true Ascomycetes (like Thelebolus). Eremascus and Dipodascus, which are often placed among the Hemiasci, possibly do not belong to the Ascomycetes series at all.
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From Strasburger’s Lehrbuch der Botanik, by permission of Gustav Fischer.
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Fig. 9.—Sphaerotheca Castagnei. Fertilization and Development of the Perithecium. (After Harper.) 1, Oogonium (og) with the antheridial branch (az) applied to its surface.
2, Separation of antheridium (an).
3, Passage of the antheridial nucleus towards that of the oogonium.
4, Union of the nuclei.
5, Fertilized oogonium surrounded by two layers of hyphae derived from the stalk-cell (st).
6, The multicellular ascogonium derived by division from the oogonium; the terminal cell with the two nuclei (as) gives rise to the ascus.
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Exoascaceae are a small group of doubtful extent here used to include Exoascus, Taphrina, Ascorticium and Endomyces. The mycelium is very much reduced in extent. The asci are borne directly on the mycelium and are therefore fully exposed, being devoid from the beginning of any investment. The Taphrineae, which include Exoascus and Taphrina, are important parasites—e.g. pocket-plums and witches’ brooms on birches, &c., are due to their action (fig. 10). Exoascus and Ascorticium present interesting parallels to Exobasidium and Corticium among the Basidiomycetes.
Saccharomycetaceae include the well-known yeasts which belong mainly to the genus Saccharomyces. They are characterized by their unicellular nature, their power of rapid budding, their capacity for fermenting various sugars, and their power of forming endogenous
