prolongation of the ectosarc containing a bundle of myonemes, so that by the contractions of the bundle the stalk is pulled down into a corkscrew spiral, and on the relaxation of the muscle the elasticity of the hollow stalk straightens it out.
On fission the stalk may become branched, as the solid one of Epistylis and Opercularia (fig. iii. 20); and the myoneme also in the tubular stem of Zoothaminum; or the branch-myoneme for the one offspring may be inserted laterally on that for the other in Carchesium (fig. iii. 18). In several tubicolous Peritrichaceae there is some arrangement for closing their tubes. In Thuricola (fig. iii. 25-26) there is a valve which opens by the pressure of the animal on its protrusion, and closes automatically by elasticity on retraction. In Lagenophrys the animal adheres to the cup a little below the opening, so that its withdrawal closes the cup: at the adherent part the body mass is hardened, and so differentiated as to suggest the frame of the mouth of a purse. In Pyxicola (fig. iii. 21-22) the animal bears some way down the body a hardened shield (“operculum”) which closes the mouth of the shell on retraction.
1, Surface view of Paramecium, showing the disposition of the cilia in longitudinal rows.
2, a, mega-; b, micronucleus; c, junction of ecto- and endosarc; D, pellicle; E, endosarc; f, cilia (much too numerous and crowded); g, trichocysts; g′, same with thread; h, discharged; i, pharynx, its undulating membrane not shown; k, food granules collecting into a bolus; l, m, n, o, food vacuoles, their contents being digested as they pass in the endosarc along the path indicated by the arrows.
3, Outline showing contractile vacuoles in commencing diastole, surrounded by five afferent canals.
4-7 Successive stages of diastole of contractile vacuole.
The cytoplasm of the Infusoria is very susceptible to injuries; and when cut or torn, unless the pellicle contracts rapidly to enclose the wounded surface, the substance of the body swells up, becoming frothy, with bubbles which rapidly enlarge and finally burst; the cell thus disintegrates, leaving only a few granules to mark where it was. This phenomenon, observed by Dujardin, is called “diffluence.” The contractile vacuole appears to be one of the means by which diffluence is avoided in cells with no strong wall to resist the absorption of water in excess; for after growing in size for some time, its walls contract suddenly, and its contents are expelled to the outside by a pore, which is, like the anus, usually invisible, but permanent in position. The contractile vacuole may be single or multiple; it may receive the contents of a canal, or of a system of canals, which only become visible at the moment of the contraction of the vacuole (fig. ii. 4-7), giving liquid time to accumulate in them, or when the vacuole is acting sluggishly or imperfectly, as in the approach of asphyxia (fig. ii. 3). Besides this function, since the system passes a large quantity of water from without through the substance of the cell, it must needs act as a means of respiration and excretion. In all Peritrichaceae it opens to the vestibule, and in some of them it discharges through an intervening reservoir, curiously recalling the arrangements in the Flagellate Euglenaceae.
The nuclear apparatus consists of two parts, the meganucleus, and the micronucleus or micronuclei (fig. iii. 17d, iv. 1). The meganucleus alone regarded and described as “the nucleus” by older observers is always single, subject to a few reservations. It is most frequently oval, and then is indented by the micronucleus; but it may be lobed, the lobes lying far apart and connected by a slender bridge or moniliform, or horseshoe-shaped (Peritrichaceae). It often contains darker inclusions, like nucleoles.
It has been shown, more especially by Gruber, that many Ciliata are multinucleate, and do not possess merely a single meganucleus and a micronucleus. In Oxytricha the nuclei are large and numerous (about forty), scattered through the protoplasm, whilst in other cases the nucleus is so finely divided as to appear like a powder diffused uniformly through the medullary protoplasm (Trachelocerca). Carmine staining, after treatment with absolute alcohol, has led to this remarkable discovery. The condition described by Foettinger in his Opalinopsis (fig. i. 1, 2) is an example of this pulverization of the nucleus. The condition of pulverization had led in some cases to a total failure to detect any nucleus in the living animal, and it was only by the use of reagents that the actual state of the case was revealed. Before fission, whatever be its habitual character, it condenses, becomes oval, and divides by constriction; and though it usually is then fibrillated, only in a few cases does it approach the typical mitotic condition. The micronucleus described by older writers as the “nucleolus” or “paranucleus” (“endoplastule” of Huxley), may be single or multiple. When the meganucleus is bilobed there are always two micronuclei, and at least one is found next to every enlargement of the moniliform meganucleus. In the fission of the Infusoria, every micronucleus divides by a true mitotic process, during which, however, its wall remains intact. From their relative sizes the meganucleus would appear to discharge during cell-life, exclusively, the functions of the nucleus in ordinary cells. Since in conjugation, however, the meganucleus degenerates and is in great part either digested or excreted as waste matter, while the new nuclear apparatus in both exconjugates arises, as we shall see, from a conjugation-nucleus of exclusively micronuclear origin, we infer that the micronucleus has for its function the carrying on of the nuclear functions of the race from one fission cycle to the next from which the meganucleus is excluded.
Fission is the ordinary mode of reproduction in the Infusoria, and is usually transverse, but oblique in Stentor, &., as in Flagellata, longitudinal in Peritrichaceae; in some cases it is always more or less unequal owing to the differentiation of the body, and consequently it must be followed by a regeneration of the missing organs in either daughter-cell. In some cases it becomes very uneven, affording every transition to budding, which process assumes especial importance in the Suctoria. Multiple fission (brood-formation or sporulation) is exceptional in Infusoria, and when it occurs the broods rarely exceed four or eight—another difference from Flagellata. The nuclear processes during conjugation suggest the phylogenetic loss of a process of multiple fission into active gametes. As noted, in fission the meganucleus divides by direct constriction; each micronucleus by a mode of mitosis. The process of fission is subject in its activity to the influences of nutrition and temperature, slackening as the food supply becomes inadequate or as the temperature recedes from the optimum for the process. Moreover, if the descendants of a single animal be raised, it is found that the rapidity of fission, other conditions being the same, varies periodically, undergoing periods of depression, which may be followed by either (1) spontaneous recovery, (2) recovery under stimulating food, (3) recovery through conjugation, or (4) the death of the cycle, which would have ensued if 2 or 3 had been omitted at an earlier stage, but which ultimately seems inevitable, even the induction of conjugation failing to restore it. These physiological conditions were first studied by E. Maupas, librarian to the city of Algiers, in his pioneering work in the later ’eighties, and have been confirmed and extended by later observers, among whom we may especially cite G. N. Calkins.
Syngamy, usually termed conjugation or “karyogamy,” is of exceptional character in the majority of this group—the Peritrichaceae alone evincing an approximation to the usual typical process of the permanent fusion of two cells (pairing-cells or gametes), cytoplasm to cytoplasm, nucleus to nucleus, to form a new cell (coupled cell, zygote).
This process was elucidated by E. Maupas in 1889, and his results, eagerly questioned and repeatedly tested, have been confirmed in every fact and in every generalization of importance.
Previously all that had been definitely made out was that under certain undetermined conditions a fit of pairing two and two occurred among the animals of the same species in a culture or in a locality in the open; that after a union prolonged over hours, and sometimes even days, the mates separated; that during the union the meganucleus underwent changes of a degenerative character; and that the micronucleus underwent repeated divisions, and that from the offspring of the micronuclei the new nuclear apparatus was evolved for each mate. Maupas discovered the biological conditions leading to conjugation: (1) the presence of individuals belonging to distinct stocks; (2) their belonging to a generation sufficiently removed from previous conjugation, but not too far removed therefrom; (3) a deficiency of food. He also showed that during conjugation a “migratory” nucleus, the offspring of the divisions of the micronucleus, passes from either mate to the other, while its sister nucleus remains “stationary”; and that reciprocal fusion of the migratory nucleus of the one mate with the stationary nucleus of the other takes place to form a zygote nucleus in either mate; and that from these zygote nuclei in each by division, at least two nuclei are formed, the one of which enlarges to form a meganucleus, while the other remains small as the first micronucleus of the new reorganized animal, which now separates as an “exconjugate” (fig. iv). Moreover, if pairing be prevented, or be not induced, the individuals produced by successive fissions become gradually weaker, their nuclear apparatus degenerates, and finally they cannot be induced