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560
INFUSORIA

organs of most varied kinds: tail-springs, cirrhi for crawling and darting, cilia and membranellae for continuous swimming in the open or gliding over surfaces or waltzing on the substratum (Trichodina, fig. iii. 8) or for eddying in wild turns through the water (Strombidium, Tintinnus, Halteria). Their forms offer a most interesting variety, and the flexibility of many adds to their easy grace of movement, especially where the front of the body is produced and elongated like the neck of a swan (Amphileptus, fig. iii. 5; Lacrymaria).

EB1911 Infusoria - Colpidium colpoda - changes during conjugation.jpg
From Lankester’s Treatise on Zoology.

Fig. v.—Diagram 1 illustrating changes during conjugation of Colpidium colpoda. (From Hickson, after Maupas.)

M, Old meganucleus undergoing disintegration.

m, Micronucleus.

N, migratory, and

S, Stationary pairing-nucleus.

M′, M′, the new meganuclei, and

m′, The new micronuclei in the products of the first fission of each of the exconjugates; the continuous vertical line indicates period of fusion, its cessation, separation; dotted lines indicate fission; the spaces lettered 1-7 successive stages in the process; the clear circles indicate functionless nuclei which degenerate.

The cytoplasm is very highly differentiated: especially the ectoplasm or ectosarc. This has always a distinct elastic “pellicle” or limiting layer, in a few cases hard, or even with local hardenings that affect the disposition of a coat of mail (Coleps) or a pair of valves (Dysteria); but is usually only marked into a rhomboidal network by intersecting depressions, with the cilia occupying the centres of the areas or meshes defined. The cytoplasm within is distinctly alveolated, and frequently contains tubular alveoli running along the length of the animal. Between these are dense fibrous thickenings, which from their double refraction, from their arrangement, and from their shortening in contracted animals are regarded as of muscular function and termed “myonemes.” Other threads running alongside of these, and not shortening but becoming wavy in the general contraction have been described in a few species as “neuronemes” and as possessing a nervous, conducting character. On this level, too, lie the dot-like granules at the bases of the cilia, which form definite groups in the case of such organs as are composed of fused cilia; in the deeper part of the ectoplasm the vacuoles or alveoli are more numerous, and reserve granules are also found; here too exist the canals, sometimes developed into a complex network, which open into the contractile vacuole.

The cilia themselves have a stiffer basal part, probably strengthened by an axial rod, and a distal flexible lash; when cilia are united by the outer plasmatic layer, they form (1) “Cirrhi,” stiff and either hook-like and pointed at the end, or brush-like, with a frayed apex; (2) membranelles, flattened organs composed of a number of cilia fused side by side, sometimes on a single row, sometimes on two rows approximated at either end so as to form a narrow oval, the membranelle thus being hollow; (3) the oral “undulating membrane,” merely a very elongated membranelle whose base may extend over a length nearly equal to the length of the animal; such membranes are present in the mouth oral depression and pharynx of all but Gymnostomaceae, and aid in ingestion; a second or third may be present, and behave like active lips; (4) in Peritrichaceae the cilia of the peristomial wreath are united below into a continuous undulating membrane, forming a spiral of more than one turn, and fray out distally into a fringe; (5) the dorsal cilia of Hypotrichaceae are slender and motionless, probably sensory.

Embedded in the ectosarc of many Ciliates are trichocysts, little elongated sacs at right angles to the surface, with a fine hair-like process projecting. On irritation these elongate into strong prominent threads, often with a more or less barb-like head, and may be ejected altogether from the body. Those over the surface of the body appear to be protective; but in the Gymnostomaceae specially strong ones surround the mouth. They can be injected into the prey pursued, and appear to have a distinctly poisonous effect on it. They are combined also into defensive batteries in the Gymnostome Loxophyllum. They are absent from most Heterotrichaceae and Hypotrichaceae, and from Peritrichaceae, except for a zone round the collar of the peristome.

The openings of the body are the mouth, absent in a few parasital species (Opalinopsis, fig. i. 1, 2), the anus and the pore of the contractile vacuole. The mouth is easily recognizable; in the most primitive forms of the Gymnostomaceae and some other groups, it is terminal, but it passes further and further back in more modified species, thereby defining a ventral, and correspondingly a dorsal surface; it usually lies on the left side. The anus is usually only visible during excretion, though its position is permanent; in a few genera it is always visible (e.g. Nyctotherus, fig. i. 16). The pore of the contractile vacuole might be described in the same terms.

The endoplasm has also an alveolar structure, and contains besides large food-vacuoles or digestive vacuoles, and shows movements of rotation within the ectoplasm, from which, however, it is not usually distinctly bounded. In Ophryoscolex and Didinium (fig. i. 13) a permanent cavity traverses it from mouth to anus.

EB1911 Infusoria - motile reaction of Paramecium.jpg
From Calkins’ Protozoa, by permission of the Macmillan Company, N. Y.

Fig. vi.—Diagrammatic view of behaviour of the motile reaction of Paramecium after meeting a mechanical obstruction at A. (From G. N. Calkins after H. S. Jennings.) For clearness and simplicity the normal motion is supposed to be straight instead of spiral.

Ingestion of food is of the same character in all the Hymenostomata. The ciliary current drives a powerful stream into the mouth, which impinges against the endosarc, carrying with it the food particles; these adhere and accumulate to form a pellet, which ultimately is pushed by an apparently sudden action into the substance of the endosarc which closes behind it (fig. ii. 2). In some of the Aspirotrichaceae accessory undulating membranes play the part of lips, and there is a closer approximation to true deglutition. The mouth is rarely terminal, more frequently at the bottom of a depression, the “vestibule,” which may be prolonged into a slender canal, sometimes called the “pharynx” or “oral tube,” ciliated as well as provided with a membrane, and extending deep down into the body in many Peritrichaceae.

In Spirostomaceae the “adoral wreath” of membranelles encloses more or less completely an anterior part of the body, the “peristome,” within which lies the vestibule. This area may be depressed, truncate, convex or produced into a short obconical disk or into one or more lobes, or finally form a funnel, or a twisted spiral like a paper cone. In most Peritrichaceae a collar-like rim surrounds the peristome, and marks out a gutter from which the vestibule opens; the peristome can be retracted, and the collar close over it. This rim forms a deep permanent spiral funnel in Spirochona (fig. iii. 10).

Movements of Ciliata.—H. S. Jennings has made a very detailed study of these movements, which resemble those of most minute free-swimming organisms. The following account applies practically to all active “Infusoria” in the widest sense.

EB1911 Infusoria - mode of progression of Paramecium.jpg

Fig. vii.—Diagram of a mode of progression of a Ciliate like Paramecium; m, mouth and pharynx; the straight line A, B, represents the axis of progression described by the posterior end, and the spiral line the curve described by the anterior end; the clear circles are the contractile vacuoles on the dorsal side.

The position of the free-swimming Infusoria, like that of Rotifers and other small swimming animals, is with the front end of the body inclined outward to the axis of advance, constantly changing its azimuth while preserving its angle constant or nearly so; if advance were ignored the body would thus rotate so as to trace out a cone, with the hinder end at the apex, and the front describing the base. On any irritation, (1) the motion is arrested, (2) the animal reverses its cilia and swims backwards, (3) it swerves outwards away from the axis so as to make a larger angle with it, and (4) then swims forwards along a new axis of progression, to which it is inclined at the same angle as to the previous axis (figs. vi., vii.). In this way it alters its axis of progression when it finds itself under conditions of stimulation. Thus a Paramecium coming into a region relatively too cold, too hot, or too poor in CO2 or in nutriment, alters its direction of swimming; in this way individuals come to assemble in crowds where food is abundant, or even where there is a slight excess of CO2. This reaction may lead to fatal results; if a solution of corrosive sublimate (Mercuric chloride) diffuses towards the hinder end of the animal faster than it progresses, the stimulus affecting the hinder end first, the axis of progression is altered so as to bring the