Page:EB1911 - Volume 22.djvu/497

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The protoplasmic body of the Protozoa is frequently differentiated into two zones or regions: a more external, termed the ectoplasm or ectosarc, and a more internal, termed the endoplasm or endosarc. The ectosarc is distinguished by being more clear and hyaline in appearance, and more tough and viscid in consistence; the endoplasm, on the other hand, is more granular and opaque, and of a more fluid nature. The ectoplasm is the protective layer of the body, and is also the portion most concerned in movement, in excretion, and perhaps also in sensation and in functions similar to those performed by the nervous systems of higher animals. The endoplasm, on the other hand, is the chief seat of digestive and reproductive functions.

As the protective layer of the body, the ectoplasm forms the envelopes or membranes which invest the surface of the body, and which are differentiations of the outermost layer of the ectoplasm. Thus in most Flagellata the ectoplasm is represented only by the more or less firm outer covering or periplast. Even when such envelopes are absent, however, the ectoplasm can still be seen to exert a protective function; as, for instance, in those Myxosporidia which are parasitic in the gall-bladders or urinary bladders of their hosts, and which can resist the action of the juices in which they live so long as the ectoplasm is intact, but succumb to the action of the medium if the ectoplasm be injured. In many Infusoria the ectoplasm contains special organs of offence termed trichocysts, each a minute ovoid body from which, on stimulation, a thread is shot out, in a manner similar to the nematocysts of Coelenterata. Similar organs are seen also in the spores of Myxosporidia, as the so-called polar capsules; but in this case the organs are not specially ectoplasmic, and appear to serve for adhesion and attachment, rather than for offence.

The connexion of the ectoplasm with movement is seen in the simplest forms, such as Amoeba, by the fact that all pseudopodia arise from it in the first instance. In forms with a definite cuticle, on the other hand, the ectoplasm usually contains contractile fibres or myonemes, forming, as it were, the muscular system of the organism. The dependence of the motility of the animal upon the development of the ectoplasm is well seen in Gregarines, in which other organs of locomotion are absent; in forms endowed with active powers of locomotion a distinct ectoplasmic layer is present below the cuticle; in those Gregarines incapable of active movement, on the other hand, the ectoplasm is absent or scarcely recognizable.

From the ectoplasm arise the special organs of locomotion, which, when present, take the form of pseudopodia, flagella or cilia. Pseudopodia, as already explained, are temporary protoplasmic organs which can be extruded or retracted at any point; they fall naturally into two principal types, between which, however, transitions are to be found: first, slender, filamentous or filose pseudopodia, composed of ectoplasm alone, which may remain separate from one another, or may anastomose to form networks, and are then termed reticulose; secondly, thick, blunt, so-called lobose pseudopodia, which are composed of ectoplasm with a core of endoplasm, and never form networks. In forms showing active locomotor powers the pseudopodia are usually more lobose in type; filose pseudopodia, on the other hand, are more adapted for the function of capturing food.

Flagella are long, slender, vibratile filaments, generally few in number when present, and usually placed at the pole of the body which is anterior in progression. Each flagellum performs peculiar lashing movements which cause the body, if free, to be dragged along after the flagellum in jerks or leaps; if, however, the body be fixed, the action of the flagellum or flagella causes a current towards it, by which means the animal obtains its food-supply. A flagellum which is anterior in movement has been distinguished by Lankester by the convenient term tractellum; sometimes, however, the flagellum is posterior in movement and acts as a propeller, like the tail of a fish; for this type Lankester has proposed the term pulsellum. The flagellum appears to arise in all cases from a distinct basal granule, and in some cases, as in the genus Trypanosoma, there is a portion of the nuclear apparatus set apart as a distinct kinetic nucleus, with the function, apparently, of governing the activities of the flagellum.

Cilia are minute, hair-like extensions of the ectoplasm, which pierce the cuticle and form typically a furry covering to the body. Though perhaps primitively derived from flagella, cilia, in their usual form, are distinguished from flagella by being of smaller size, by being present, as a rule, in much greater numbers, and above all by the character of their movements. In the place of the complicated lashing movements of the flagella, each cilium performs a simple stroke in one direction, becoming first bowed on one side, by an act of contraction, and then straightened out again when relaxed. The movements of the cilia are co-ordinated and they act in concert, though not absolutely in unison, each one contracting just before or after its neighbour, so that waves of movement pass over a ciliated surface in a given direction, similar to what may be seen in a cornfield when the wind is blowing over it. Primitively coating the whole surface of the body evenly, the cilia may become modified and specialized in various ways, which cannot be described in detail here (see Infusoria).

Besides the organs of locomotion already mentioned, there may be present so-called undulating membranes, in the form of thin sheets of ectoplasm which are capable of performing sinuous, undulating movements by their inherent contractility. In some cases distinct contractile threads or myonemes have been described in these membranes. Undulating membranes appear to be formed either by the fusion together of a row of cilia, side by side, or by the attachment of a flagellum to the body by means of an ectoplasmic web, in which case the flagellum forms the free edge of the membrane, as in the genus Trypanosoma.

Returning to the ectoplasm, the excretory function exerted by this layer is seen by the formation in it of the peculiar contractile vacuoles found in most free-living Protozoa. A contractile vacuole is a spherical drop of watery fluid which makes its appearance periodically at some particular spot near the surface of the animal’s body, or, if more than one such vacuole is present, at several definite and constant places. Each vacuole grows to a certain size, and when it has reached the limit of its growth it discharges its contents to the exterior by a sudden and rapid contraction. There is, apparently, in most if not in all cases, a definite pore through which the contractile vacuole empties itself to the exterior. On account of the relatively large size which the contractile vacuole attains it bulges inwards beyond the limits of the ectoplasm and comes to lie chiefly in the endoplasm, to which it is sometimes, but erroneously, ascribed. In the most highly differentiated Protozoa, for instance, the Ciliata, the ectoplasm contains an apparatus of excretory channels, situated in its deeper layers, and forming as it were a drainage-system, from which the contractile vacuoles are fed. The fluid discharged by the contractile vacuoles appears to be chiefly water which has been absorbed at the surface of the protoplasmic body, and which has filtered through the protoplasm, taking up the soluble waste nitrogenous products of the metabolism and the gaseous products of respiration; hence the contractile vacuoles may be compared in a general way to the urinary and respiratory organs of the Metazoa.

One of the first consequences of the parasitic habit of life is the disappearance of the contractile vacuoles, which are hardly ever found in truly parasitic Protozoa, that is to say, in forms which live in the interior of other animals and nourish themselves at their expense. They are also very frequently absent in marine forms.

Mechanisms of a nervous nature are very seldom found in Protozoa, but in some Ciliata special tactile bristles are found, and it is possible that flagella, and perhaps even pseudopodia, may be sometimes tactile rather than locomotor in function. Pigment-spots, apparently sensitive to light, may also occur in some Flagellata.

The endoplasm, as already stated, is the chief seat of nutritive and reproductive processes. In many Flagellata the ectoplasm