the teleutospores, each contain two nuclei when young, which
fuse as the spores mature. In young asci a similar fusion of two
nuclei occurs, and also in basidia, in each case the nucleus of the
ascus or of the basidium resulting from the fusion subsequently
giving rise by division to the nuclei of the ascospores and basidiospores
respectively. The significance of these fusions will be discussed
under the various groups. Nuclear division is usually
accompanied by all the essential features of karyokinesis.
Spores.—No agreement has ever been arrived at regarding the consistent use of the term spore. This is apparently owing to the facts that too much has been attempted in the definition, and that differences arise according as we aim at a morphological or a physiological definition. Physiologically, any cell or group of cells separated off from a hypha or unicellular fungus, and capable of itself growing out—germinating—to reproduce the fungus, is a spore; but it is evident that so wide a definition does not exclude the ordinary vegetative cells of sprouting fungi, such as yeasts, or small sclerotium like cell-aggregates of forms like Coniothecium. Morphologically considered, spores are marked by peculiarities of form, size, colour, place of origin, definiteness in number, mode of preparation, and so forth, such that they can be distinguished more or less sharply from the hyphae which produce them. The only physiological peculiarity exhibited in common by all spores is that they germinate and initiate the production of a new fungus-plant. Whether a spore results from the sexual union of two similar gametes (zygospore) or from the fertilization of an egg-cell by the protoplasm of a male organ (oospore); or is developed asexually as a motile (zoospore) or a quiescent body cut off from a hypha (conidium) or developed along its course (oidium or chlamydospore), or in its protoplasm (endospore), are matters of importance which have their uses in the classification and terminology of spores, though in many respects they are largely of academic interest.
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Fig. 2.—Peronospora parasitica (De Bary). Conidiophore with conidia. |
Klebs has attempted to divide spores into three categories as follows: (1) kinospores, arising by relatively simple cell-divisions and subserving rapid dissemination and propagation, e.g. zoospores, conidia, endogonidia, stylospores, &c.; (2) paulospores, due to simple rearrangement of cell-contents, and subserving the persistence of the fungus through periods of exigency, e.g. gemmae, chlamydospores, resting-cells, cysts, &c.; (3) carpospores, produced by a more or less complex formative process, often in special fructifications, and subserving either or both multiplication and persistence, e.g. zygospores, oospores, brand-spores, aecidiospores, ascospores, basidiospores, &c. Little or nothing is gained by these definitions, however, which are especially physiological. In practice these various kinds of spores of fungi receive further special names in the separate groups, and names, moreover, which will appear, to those unacquainted with the history, to have been given without any consistency or regard to general principles; nevertheless, for ordinary purposes these names are far more useful in most cases, owing to their descriptive character, than the proposed new names, which have been only partially accepted.
Sporophores.—In some of the simpler fungi the spores are not borne on or in hyphae which can be distinguished from the vegetative parts or mycelium, but in the vast majority of cases the sporogenous hyphae either ascend free into the air or radiate into the surrounding water as distinct branches, or are grouped into special columns, cushions, layers or complex masses obviously different in colour, consistency, shape and other characters from the parts which gather up and assimilate the food-materials. The term “receptacle” sometimes applied to these spore-bearing hyphae is better replaced by sporophore. The sporophore is obsolete when the spore-bearing hyphae are not sharply distinct from the mycelium, simple when the constituent hyphae are isolated, and compound when the latter are conjoined. The chief distinctive characters of the sporogenous hyphae are their orientation, usually vertical; their limited apical growth; their peculiar branching, form, colour, contents, consistency; and their spore-production. According to the characters of the last, we might theoretically divide them into conidiophores, sporangiophores, gametophores, oidiophores, &c.; but since the two latter rarely occur, and more than one kind of spore or spore-case may occur on a sporophore, it is impossible to carry such a scheme fully into practice.
A simple sporophore may be merely a single short hypha, the end of which stops growing and becomes cut off as a conidium by the formation of a septum, which then splits and allows the conidium to fall. More generally the hypha below the septum grows forwards again, and repeats this process several times before the terminal conidium falls, and so a chain of conidia results, the oldest of which terminates the series (Erysiphe); when the primary branch has thus formed a basipetal series, branches may arise from below and again repeat this process, thus forming a tuft (Penicillium). Or the primary hypha may first swell at its apex, and put forth a series of short peg-like branches (sterigmata) from the increased surface thus provided, each of which develops a similar basipetal chain of conidia (Aspergillus), and various combinations of these processes result in the development of numerous varieties of exquisitely branched sporophores of this type (Botrytis, Botryosporium, Verticillium, &c.).
Fig. 3.—Cystopus candidus.
| A. | a, | Conidia. |
| b, | Conidiophores. | |
| c, | Conidium emitting zoospores. | |
| d, | Free zoospore. | |
| B. | og, | Oogonium. |
| os, | Oosphere. | |
| an, | Antheridium. | |
| C. | Formation of zoospores by oospores. | |
| z, | Free zoospores. (After De Bary.) |
A second type is developed as follows: the primary hypha forms a septum below its apex as before, and the terminal conidium, thus abstricted, puts out a branch at its apex, which starts as a mere point and rapidly swells to a second conidium; this repeats the process, and so on, so that we now have a chain of conidia developed in acropetal succession, the oldest being below, and, as in Penicillium, &c., branches put forth lower down may repeat the process (Hormodendron). In all these cases we may speak of simple conidiophores. The simple sporophore does not necessarily terminate in conidia, however. In Mucor, for example, the end of the primary hypha swells into a spheroidal head (sporangium), the protoplasm of which undergoes segmentation into more or less numerous globular masses, each of which secretes an enveloping cell-wall and becomes a spore (endospore), and branched systems of sporangia may arise as before (Thamnidium). Such may be termed sporangiophores. In Sporodinia the branches give rise also to short branches, which meet and fuse their contents to form zygospores. In Peronospora, Saprolegnia, &c., the ends of the branches swell up into sporangia, which develop zoospores in their interior (zoosporangia), or their contents become oospheres, which may be fertilized by the contents of other branches (antheridia) and so form egg-cases (oogonia). Since in such cases the sporophore bears sexual cells, they may be conveniently termed gametophores.
Compound sporophores arise when any of the branched or unbranched types of spore-bearing hyphae described above ascend into the air in consort, and are more or less crowded into definite layers, cushions, columns or other complex masses. The same laws apply to the individual hyphae and their branches as to simple sporophores, and as long as the conidia, sporangia, gametes, &c., are borne on their external surfaces, it is quite consistent to speak of these as compound sporophores, &c., in the sense described, however complex they may become. Among the simplest cases are the sheet-like aggregates of sporogenous hyphae in Puccinia, Uromyces, &c., or of basidia in Exobasidium, Corticium, &c., or of asci in Exoascus, Ascocorticium, &c. In the former, where the layer is small, it is often termed a sorus, but where, as in the latter, the sporogenous layer is extensive, and spread out more or less sheet-like on the supporting tissues, it is more frequently termed a hymenium. Another simple case is that of the columnar aggregates of sporogenous hyphae in forms like Stilbum, Coremium, &c. These lead
