In its plasmodial state, as has been said, the slime-mould affects damp or moist situations, and during warm weather in such places spreads over all moist surfaces, creeps through the interstices of the rotting bark, spreads between the cells, between the growth-layers of the wood, runs in corded vein-like nets between the wood and bark, and finds in all these cases nutrition in the products of organic decomposition. Such a plasmodium may be divided, and so long as suitable surroundings are maintained, each part will manifest all the properties of the whole. Parts of the same plasmodium will even coalesce again. If a piece of plasmodium-bearing wood be brought indoors, be protected from desiccation by aid of a moist dark chamber, not too warm (70° F.), the organism seems to suffer little if any injury, but will continue for days or weeks to manifest all the phenomena of living matter. Thus, under such circumstances, the plasmodium will constantly change shape and position, can be induced to spread over a plate of moist glass, and so be transferred to the stage of a microscope, there to exhibit in the richest and most interesting and abundant fashion the streaming protoplasmic currents. As just indicated, the plasmodia follow moisture, creep from one moist substance to another, especially follow nutritive substrata. They seem also to secure in some way exclusive possession. I have never seen them interfered with by hyphæ or enemies of any sort, nor do they seem to interfere with one another. Plasmodia of two common species, Hemitrichia clavata and H. vesparium are often side by side on the same substratum, but do not mix, and their perfected fruits presently stand erect side by side, each with its own characteristics, entirely unaffected by the presence of the other. On the other hand, it is probable that some of the forms which, judged by their different fructifications, and by this alone, are to us distinct, may be more closely related than we suspect, and puzzling phases which show the distinctive marks supposed to characterize different species are no doubt sometimes to be explained on the theory of plasmodial crossing; they are hybrids.
Under certain conditions, low temperature, lack of moisture, the plasmodium may pass into a resting phase, when it masses itself in heaps and may become quite dry in lumps of considerable size, and so await the return of favorable conditions when former activity is quickly resumed. Sometimes the larger plasmodia pass into the resting phase by undergoing a very peculiar change of structure. In ordinary circumstances the abundant free nuclei demonstrable in the plasmodium afford the only evidence of cellular organization. In passing now into the condition of rest, the whole protoplasmic mass separates simultaneously into numerous definite polyhedral or parenchymatous cells, each with a well-developed cellulose wall.[4] When the conditions essential to activity are restored, the walls disappear, the cellulose is resorbed, and the plasmodium resumes its usual habit and structure.
The plasmodial phase of the slime-mould, like the hyphal phase of the fungus, may continue a long time; for months, possibly for years. The reason for making the latter statement will presently appear. But however long or short the plasmodial phase continue, the time of fruit, the reproductive phase, at length arrives. When this time comes, induced partly by a certain maturity in the organism itself, partly no doubt by the trend of external conditions, the plasmodium no longer as before evades the light, but pushes to the surface, and appears usually in some elevated or exposed position, the upper side of the log, the top of the stump, the upper surface of its habitat, whatever that may be; or even leaves its nutrient base entirely and finds lodging on some neighboring object. In such emergency the stems and leaves of flowering plants are often made to serve, and even fruits and flowers afford convenient resting places. The object now to be attained is not the formation of fruit alone, but likewise its speedy desiccation and the prompt dispersal of the perfected spores. Nothing can be more interesting than to watch the slime-mould as its plasmodium accomplishes this its last migration. If hitherto its habitat has been the soft interior of a rotten log, it now begins to ooze out in all directions, to well up through the crevices of the bark as if pushed by some energy acting in the rear, to stream down upon the ground, to flow in a hundred tiny streams over all the region round about, to climb all stems, ascend all branches, to the height of many inches, all to pass suddenly as if by magic charm into one widespread, dusty field of flying spores. Or, to be more exact, whatever the position ultimately assumed, the plasmodium soon becomes quiescent, takes on definite and ultimate shape, which varies greatly, almost for each species. Thus it may simply form a flat, cake-like mass, aethalium, internally divided into an indefinite number of ill-defined spore cases, sporangia; or the plasmodium may take the form of a simple net, plasmodiocarp, whose cords stand out like swollen veins, whose meshes vary both in form and size; or more commonly the whole protoplasmic mass breaks up into little spheroidal heaps which may be sessile directly on the substratum, or may be lifted on tiny stems, stipitate, which may rest in turn upon a common sheet-like film, or more or less continuous net, spreading beneath them all, the hypothallus. In any case, each differentiated portion of the plasmodium, portion poorly or well defined, elongate, net-like, spheroidal, elliptical, or of whatever shape, becomes at length a sporangium, spore-case, receptacle for the development and temporary preservation of the spores.[5]
The slime-moulds were formerly classed with the gasteromycetous fungi, puff-balls, and in description of their fruiting phase the terms applicable to the description of a puff-ball are still employed, although it will be understood that the structures described are not in the two cases homologous; analogous only. The sporangium of the slime-mould exhibits usually a distinct peridium, or outer limiting wall, which is at first continuous, enclosing the spores and their attendant machinery, but at length ruptures, irregularly as a rule, and so suffers the contents to escape. The peridium may be double, varies in texture, color, persistence, and so forth, as will be more fully set forth in the several specific descriptions. The peridium blends with the hypothallus below when such structure is recognizable, either directly, when the sporangium is sessile, or by the intervention of a stipe. The stipe may be hollow, may contain coloring matter of some sort, or may even contain peculiar spore-like cells or spores; is often furrowed, and in some cases shows a disposition to unite or blend with the stalks of neighboring sporangia. In many cases the stipe is continued upward, more or less definitely into the cavity of the sporangium, and there forms the columella, sometimes simple and rounded, like the analogous structure in the Mucores, sometimes as in Comatricha, branching again and again in wonderful richness and complexity.
Each sporangium is at maturity filled with numerous unicellular spores. These are usually spherical, sometimes flattened at various points by mutual contact; they are of various colors, more commonly yellow or violet brown, are sometimes smooth (?), but generally roughened either by the presence of minute warts, or spines, or by the occurence of more or less strongly elevated bands dividing reticulately the entire surface. The spores are in all cases small 3–20 µ, and reveal their surface characters only under the most excellent lenses.
Associated with the spores in the sporangium occurs the capillitium. This consists of most delicate thread-or hair-like elements, offering great variety both in form and structure. The threads composing the capillitium are not to be regarded, even when free, as cells, nor even of cellular origin; probably, as would appear from the researches of Strasburger and Harper, all forms of capillitial threads arise in connection with vacuoles in the protoplasmic mass. "Whether the thread is hollow or solid, simple or branched, free or connected with the peridium or a columella,—these are entirely secondary conditions, depending on the extent and form of the vacuoles."[6] They may occur singly or be combined into a net, they may be terete or flat, attached to the peridial wall or free, simple or adorned with bands or spires and knobs in every variety, uniform or profusely knotted and thickened at intervals, and burdened with calcic particles. In many cases, the capillitium contributes materially to the dispersal of the spores; in others, it doubtless contributes mechanically to the support of the peridial wall, and renders so far persistent the delicate sporangium. For more exact description the reader is again referred to the specific delineations which follow.
The transition from phase to phase requires, as intimated, no great length of time. Tilmadoche polycephala completed the transition from vegetative to fruiting phase in less than twelve hours.
The germination of the spores ensues closely upon their dispersal or maturity and is unique in many respects.[7] The wall of the spore is ruptured and the protoplasmic content escapes as a zoöspore indistinguishable so far from an amœba, or from the zoöspore of our chytridiaceous fungi. This amœboid zoöspore is without cell-wall, changes its outline, and moves slowly by creeping or flowing from point to point. At this stage many of the spores assume each a flagellate cilium, and so acquire power of more rapid locomotion. The zoöspores, whether ciliate or not, thus enjoy independent existence and are capable of continuing such existence for some time, assimilating, growing, and even reproducing themselves by simple fission, over and over again. This takes place, of course, only in the presence of suitable nutrient media.
Nevertheless the spores of many species germinate quickly simply in water, and a drop suspended in the form of the ordinary drop-culture on a cover-glass affords ample opportunity. In the course of time, usually not more than two or three days, the swarm spores cease their activity, lose their cilia, and come to rest, exhibiting at most nothing more than the slow amœboid movement already referred to. In the course of two or three days more, in favorable cases, the little spores begin to assemble and flow together; at first into small aggregations, then larger, until at length all have blended in one creeping protoplasmic mass to form thus once again the plasmodium, or plasmodial phase with which the round began. Small plasmodia may generally be thus obtained artificially from drop-cultures. Such, however, in the experience of the writer, are with difficulty kept alive. Hay infusions, infusions of rotten wood, etc., may sometimes for a time give excellent results.
The spores of Didymium crustaceum were sown upon a heap of leaves in autumn. An abundant display of the same species followed in the next June; but, of course, the intervening phases were not observed. The most satisfactory studies are obtained by plasmodia carefully brought in directly from the field. A plasmodium that appeared suddenly and passed to fruit on agar in a petri dish offers a valuable suggestion for further research.