base to extremity, as in Clava (fig. 5); (2) capitate, i.e. knobbed
at the extremity, as in Coryne (see Allman, loc. cit. pl. iv.); (3)
branched, a rare form in the polyp, but seen in Cladocoryne (see
Allman, loc. cit. p. 380, fig. 82). Sometimes more than one type of
form is found in the same
polyp; in Pennaria and
Stauridium (fig. 2) the upper
whorls are capitate, the lower
filiform. Finally, as regards
structure, the tentacles may
retain their primitive hollow
nature, or become solid by
obliteration of the axial
cavity.
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Fig. 4.—Diagram of Tubularia indivisa. A single hydriform person a bearing a stalk carrying numerous degenerate medusiform persons or sporosacs b. (After Allman.) |
The hypostome of the hydropolyp may be small, or, on the other hand, as in Eudendrium (Allman, loc. cit. pls. xiii., xiv.), large and trumpet-shaped. In the curious polyp Myriothela the body of the polyp is differentiated into nutritive and reproductive portions.
Histology.—The ectoderm of the hydropolyp is chiefly sensory, contractile and protective in function. It may also be glandular in places. It consists of two regions, an external epithelial layer and a more internal sub-epithelial layer.
The epithelial layer consists of (1) so-called “indifferent” cells secreting the perisarc or cuticle and modified to form glandular cells in places; for example, the adhesive cells in the foot. (2) Sensory cells, which may be fairly numerous in places, especially on the tentacles, but which occur always scattered and isolated, never aggregated to form sense-organs as in the medusa. (3) Contractile or myo-epithelial cells, with the cell prolonged at the base into a contractile muscle-fibre (fig. 6, B). In the hydropolyp the ectodermal muscle-fibres are always directed longitudinally. Belonging primarily to the epithelial layer, the muscular cells may become secondarily sub-epithelial.
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From Allman’s Gymnoblastic Hydroids, by permission of the Council of the Ray Society. |
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Fig. 5.—Colonies of Clava. A, Clava squamata, magnified. B, C. multicornis, natural size; p, polyp; gon, gonophores; rh, hydrorhiza. |
The sub-epithelial layer consists primarily of the so-called interstitial cells, lodged between the narrowed basal portions of the epithelial cells. From them are developed two distinct types of histological elements; the genital cells and the cnidoblasts or mother-cells of the nematocysts. The sub-epithelial layer thus primarily constituted may be recruited by immigration from without of other elements, more especially by nervous (ganglion) cells and muscle-cells derived from the epithelial layer. In its fullest development, therefore, the sub-epithelial layer consists of four classes of cell-elements.
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Fig. 6 A.—Portion of the body-wall of Hydra, showing ectoderm cells above, separated by “structureless lamella” from three flagellate endoderm cells below. The latter are vacuolated, and contain each a nucleus and several dark granules. In the middle ectoderm cell are seen a nucleus and three nematocysts, with trigger hairs projecting beyond the cuticle. A large nematocyst, with everted thread, is seen in the right-hand ectodermal cell. (After F. E. Schulze.) |
The genital cells are simple wandering cells (archaeocytes), at first minute and without any specially distinctive features, until they begin to develop into germ-cells. According to Wulfert [60] the primitive germ-cells of Gonothyraea can be distinguished soon after the fixation of the planula, appearing amongst the interstitial cells of the ectoderm. The germ-cells are capable of extensive migrations, not only in the body of the same polyp, but also from parent to bud through many non-sexual generations of polyps in a colony (A. Weismann [58]).
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Fig. 6 B.—Epidermo-muscular cells of Hydra. m, muscular-fibre processes. (After Kleinenberg, from Gegenbaur.) |
The cnidoblasts are the mother-cells of the nematocysts, each cell producing one nematocyst in its interior. The complete nematocyst (fig. 7) is a spherical or oval capsule containing a hollow thread, usually barbed, coiled in its interior. The capsule has a double wall, an outer one (o.c.), tough and rigid in nature, and an inner one (i.c.) of more flexible consistence. The outer wall of the capsule is incomplete at one pole, leaving an aperture through which the thread is discharged. The inner membrane is continuous with the wall of the hollow thread at a spot immediately below the aperture in the outer wall, so that the thread itself (f) is simply a hollow prolongation of the wall of the inner capsule inverted and pushed into its cavity. The entire nematocyst is enclosed in the cnidoblast which formed it. When the nematocyst is completely developed, the cnidoblast passes outwards so as to occupy a superficial position in the ectoderm, and a delicate protoplasmic process of sensory nature, termed the cnidocil (cn) projects from the cnidoblast like a fine hair or cilium. Many points in the development and mechanism of the nematocyst are disputed, but it is tolerably certain (1) that the cnidocil is of sensory nature, and that stimulation, by contact with prey or in other ways, causes a reflex discharge of the nematocyst; (2) that the discharge is an explosive change whereby the in-turned thread is suddenly everted and turned inside out, being thus shot through the opening in the outer wall of the capsule, and forced violently into the tissues of the prey, or, it may be, of an enemy; (3) that the thread inflicts not merely a mechanical wound, but instils an irritant poison, numbing and paralysing in its action. The points most in dispute are, first, how the explosive discharge is brought about, whether by pressure exerted external to the capsule (i.e. by contraction of the cnidoblast) or by internal pressure. N. Iwanzov [27] has brought forward strong grounds for the latter view, pointing out that the cnidoblast has no contractile mechanism and that measurements show discharged capsules to be on the average slightly larger than undischarged ones. He believes that the capsule contains a substance which swells very rapidly when brought into contact with water, and that in the undischarged condition the capsule has its opening closed by a plug of protoplasm (x, fig. 7) which prevents
