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Journal Article

Morphology of the Genus Actinomyces. II

Charles Drechsler
Botanical Gazette
Vol. 67, No. 2 (Feb., 1919), pp. 147-168
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Page Count: 30

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Topics: Mycelium, Hyphae, Spores, Actinomyces, Fructification, Sporulation, Species, Septum, Branching, Fertility
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1. The vegetative thallus of Actinomyces consists of a mycelium composed of profusely branching hyphae, the terminal growing portions of which are densely filled with protoplasm. Toward the center of the thallus the vacuoles increase in size and may be associated with the presence of metachromatic granules, the latter having nothing in common with bacterial endospores or "micrococci," for which they were mistaken by early observers. 2. The vegetative mycelium attains an extent incomparably greater than the branching figures recorded for bacteria of the acid-fast group, and the hyphae lack the uniformity in diameter generally characteristic of the Schizomycetes. 3. The aerial mycelium produced on suitable substrata by most species occurs usually in the form of a mat of discrete fructifications; but in other species these fructifications are frequently combined to form numerous and peculiar erect Isarioid sporodochia. 4. In any case each individual fructification represents a well characterized sporogenous apparatus, consisting of a sterile axial filament bearing branches in an open racemose or dense capitate arrangement. The primary branches may function directly as sporogenous hyphae, or may proliferate branches of the second and of higher orders, sporogenesis in the latter case being confined to the terminal elements, the hyphal portions below the points of attachment of branches remaining sterile. 5. Two tendencies in the development of fructifications are recognizable: one leading to an erect drendroidal type, in which successively proliferated fertile elements undergo processes of sporogenesis in continuous sequence; and the other leading to a prostrate racemose type, in which sporogenesis is delayed in the older branches until the younger branches have also attained their final extension. The majority of species show these tendencies combined in different ways. 6. The sporogenous hyphae of most species are coiled in peculiar spirals, sometimes resembling the spores of the hyphomycetous genus Helicoon. These spirals exhibit pronounced specific characteristics in the number, diameter, and obliquity of their turns, and especially in the direction of rotation (whether dextrorse or sinistrorse). 7. Sporogenesis, where it can be followed, begins at the tips of the fertile branches and proceeds basipetally. In the larger number of species the process involves the insertion of septa which, in certain cases, are relatively very massive, and in others so thin as to be barely discernible. The disposition of these septa, while the delimited spores undergo maturation processes, varies with the species: (1) they may remain more or less unaltered; (2) they may suffer a median split, the two resulting halves being then separated as the result of the longitudinal contraction of the young spores, leaving alternate portions of hyphal walls completely evacuated; or (3) they may first become considerably constricted and subsequently converted into non-stainable isthmuses connecting the mature spores. The apparent absence of septa in the sporogenous hyphae of other forms is perhaps attributable to optical difficulties. 8. Granules are readily differentiated in the spores of many species which possess the staining properties and uniformity of size characteristic of nuclei; they generally occur singly, but in the larger spores of a few forms two are often found occupying diagonally opposite positions. 9. As in the vegetative thallus, metachromatic granules occur in the aerial mycelium, being very rarely found in the spores or sporogenous hyphae, but becoming very abundant in degenerate sterile hyphae. 10. The older axial filaments of some species show marked distensions which, in extreme cases, result in figures simulating Leptomitus. These arise as local distensions at the points of attachment of the more extensive lateral sporogenous processes. Cuneate modifications of the sterile axial filaments below the origins of branches also occur. 11. Curious spherical structures appear regularly in some forms, both in the sterile axial hyphae, where they may contain either a median septum or a number of peripheral metachromatic granules, and in the sporogenous hyphae, where they are associated with the regularly spaced septa. 12. The spores germinate readily in suitable solutions, producing 1-4 germ tubes, the approximate number being more or less characteristic of the species. 13. Owing to the absence of any well defined bacterial characteristics, the writer is of the opinion that the view that Actinomyces represents a transition between the Hyphomycetes and the Schizomycetes, as well as the phylogenetic corollary based upon it, may safely be abandoned. If mere size is to be regarded as important, it would appear to be equally profitable to look for bacterial affinities in some ascomycetous and sphaeropsideaceous forms, the hyphae of which are similarly very minute. It is doubtful whether far-reaching taxonomic generalizations can be based on the "acid-fast" staining reaction, especially as this reaction has not played a very important role in mycological research. There seems to be no adequate reason why the genus should not be classed in an unqualified manner with the Hyphomycetes, as a mucedineous group with tendencies toward an erect Isarioid habit.