Fig. 51.—Carchesium polypinum. Scheme of the path taken by the ingested food in digestion and expulsion of the excreta. The food enters through the pharynx and is transported downward (small circles), where it is stored in the concavity of the sausage-shaped meganucleus (the latter is recognised by its containing darker bodies). It remains here for some time at rest (small crosses). Then it passes upward upon the other side (dots) and returns to the middle of the cell, where it undergoes solution. The excreta are removed to the outside, through the vestibule and cell mouth. The black line with arrows indicates the direction of the path. (From Verworn, after Greenwood.)
We owe to Miss Greenwood[[163]] a full account of the formation and changes of the food-vacuoles in Carchesium polypinum. The vacuole passes steadily along the endosarc for a certain time after its sudden admission into it, and then enters on a phase of quiescence. A little later the contents of the vacuole aggregate together in the centre of the vacuole, where they are surrounded by a zone of clear liquid; this takes place in the hollow of the meganucleus, in this species horseshoe-shaped. The vacuole then slowly passes on towards the peristome, lying deep in the endosarc, and the fluid peripheral zone is absorbed. For some time no change is shown in the food-material itself: this is the stage of "storage." Eventually a fresh zone of liquid, the true digestive vacuole, forms again round the food-pellet, and this contains a peptic juice, of acid reaction. The contents, so far as they are capable of being digested, liquefy and disappear. Ultimately the solid particles in their vacuole reach the anal area of the vestibule, and pass into it, to be swept away by the overflow of the food-current. The anus is seated on a transverse ridge about a third down the tube, the remaining two-thirds being the true pharynx.
Fission is usually transverse; but is oblique in the conical Heterotrichaceae, and longitudinal in the Peritrichaceae. It involves the peristome, of which one of the two sisters receives the greater, the other the lesser part; each regenerates what is missing. When there are two contractile vacuoles, as in Paramecium, either sister receives one, and has to form another; where there is a canal or reservoir divided at fission, an extension of this serves to give rise to a new vacuole in that sister which does not retain the old one. In some cases the fission is so unequal as to have the character of budding (Spirochona). We have described above (p. [144]) the relations of the nuclear apparatus in fission.
Several of the Ciliata divide only when encysted, and then the divisions are in close succession, forming a brood of four, rarely more. This is well seen in the common Colpoda cucullus. In the majority, however, encystment is resorted to only as a means of protection against drought, etc., or for quiet rest after a full meal (Lacrymaria).
Maupas[[164]] has made a very full study of the life-cycles of the Ciliata. He cultivated them under the usual conditions for microscopic study, i.e. on a slide under a thin glass cover supported by bristles to avoid pressure, preserved in a special moist chamber; and examined them at regular intervals.
Fig. 52.—Paramecium caudatum, stages in conjugation. gul, Gullet; mg.nu, meganucleus; Mg.nu, reconstructed meganucleus; mi.nu, micronucleus; Mi.nu, reconstructed micronucleus; o, mouth. (From Parker and Haswell, after Hertwig.)
The animals collect at that zone where the conditions of aeration are most suitable, usually just within the edge of the cover, and when well supplied with food are rather sluggish, not swimming far, so that they are easily studied and counted. When well supplied with appropriate food they undergo binary fission at frequent intervals, dividing as often as five times in the twenty-four hours at a temperature of 65-69° F. (Glaucoma scintillans), so that in this period a single individual has resolved itself into a posterity of 32; but such a rapid increase is exceptional. At a minimum and a maximum temperature multiplication is arrested, the optimum lying midway. If the food-supply is cut off, encystment occurs in those species capable of the process; but when there is a mixture of members of different broods of the same species, subject to the limitations that we shall learn, conjugation ensues. Under the conditions of Maupas' investigations he found a limit to the possibilities of continuous fissions, even when interrupted by occasional encystment. The individuals of a series ultimately dwindle in size, their ciliary apparatus is reduced, and their nuclear apparatus degenerates. Thus the ultimate members of a fission-cycle show a progressive decay, notably in the nuclear apparatus, which Maupas has aptly compared to "senility" or "old age" in the Metazoan. If by the timely mixture of broods conjugation be induced, these senile degenerations do not occur.[[165]] In Stylonychia mytilus the produce of a being after conjugation died of senility after 336 fissions; in Leucophrys after 660.
Save in the Peritrichaceae (p. [151]) conjugation takes place between similar mates, either of the general character and size of the species, or reduced by fissions, in rapid succession, induced by the same conditions as those of mating. The two mates approach, lying parallel and with their oral faces or their sides (Stentor) together, and partially fuse thereby; though no passage of cytoplasm is seen it is probable that there is some interchange or mixture.[[166]]
