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138
[ORGANIZATION
HYDROMEDUSAE


access of water to the contents; when the cnidocil is stimulated it sets in action a mechanism or perhaps a series of chemical changes by which the plug is dissolved or removed; as a result water penetrates into the capsule and causes its contents to swell, with the result that the thread is everted violently. A second point of dispute concerns the spot at which the poison is lodged. Iwanzov believes it to be contained within the thread itself before discharge, and to be introduced into the tissues of the prey by the eversion of the thread. A third point of dispute is whether the nematocysts are formed in situ, or whether the cnidoblasts migrate with them to the region where they are most needed; the fact that in Hydra, for example, there are no interstitial cells in the tentacles, where nematocysts are very abundant, is certainly in favour of the view that the cnidoblasts migrate on to the tentacles from the body, and that like the genital cells the cnidoblasts are wandering cells.

EB1911 Hydromedusae - structure of Nematocysts and their mode of working.jpg

Fig. 7.—Diagrams to show the structure of Nematocysts and their mode of working. (After Iwanzov.)

a, Undischarged nematocyst.
b, Commencing discharge.
c, Discharge complete.
cn, Cnidocil.
N, Nucleus of cnidoblast.
o.c Outer capsule.
x, Plug closing the opening of the outer capsule.
i.c., Inner capsule, continuous with the wall of the filament, f.
b, Barbs.

The muscular tissue consists primarily of processes from the bases of the epithelial cells, processes which are contractile in nature and may be distinctly striated. A further stage in evolution is that the muscle-cells lose their connexion with the epithelium and come to lie entirely beneath it, forming a sub-epithelial contractile layer, developed chiefly in the tentacles of the polyp. The evolution of the ganglion-cells, is probably similar; an epithelial cell develops processes of nervous nature from the base, which come into connexion with the bases of the sensory cells, with the muscular cells, and with the similar processes of other nerve-cells; next the nerve-cell loses its connexion with the outer epithelium and becomes a sub-epithelial ganglion-cell which is closely connected with the muscular layer, conveying stimuli from the sensory cells to the contractile elements. The ganglion-cells of Hydromedusae are generally very small. In the polyp the nervous tissue is always in the form of a scattered plexus, never concentrated to form a definite nervous system as in the medusa.

EB1911 Hydromedusae - Cells from the axis of the tentacle of a Medusa.jpg

From Gegenbaur’s Elements of Comparative Anatomy.

Fig. 8.—Vacuolated Endoderm Cells of cartilaginous consistence from the axis of the tentacle of a Medusa (Cunina).

The endoderm of the polyp is typically a flagellated epithelium of large cells (fig. 6), from the bases of which arise contractile muscular processes lying in the plane of the transverse section of the body. In different parts of the coelenteron the endoderm may be of three principal types—(1) digestive endoderm, the primitive type, with cells of large size and considerably vacuolated, found in the hydranth; some of these cells may become special glandular cells, without flagella or contractile processes; (2) circulatory endoderm, without vacuoles and without basal contractile processes, found in the hydrorhiza and hydrocaulus; (3) supporting endoderm (fig. 8), seen in solid tentacles as a row of cubical vacuolated cells, occupying the axis of the tentacle, greatly resembling notochordal tissue, particularly that of Amphioxus at a certain stage of development; as a fourth variety of endodermal cells excretory cells should perhaps be reckoned, as seen in the pores in the foot of Hydra and elsewhere (cf. C. Chun, Hydrozoa [1], pp. 314, 315).

EB1911 Hydromedusae - Colony of Hydractinia echinata on a Whelk.jpg

From Allman’s Gymnoblastic Hydroids, by permission of the Council of the Ray Society.

Fig. 9.—Colony of Hydractinia echinata, growing on the Shell of a Whelk. Natural size.

The mesogloea in the hydropolyp is a thin elastic layer, in which may be lodged the muscular fibres and ganglion cells mentioned above, but which never contains any connective tissue or skeletogenous cells or any other kind of special mesogloeal corpuscles.

EB1911 Hydromedusae - Polyps from a Colony of Hydractinia.jpg

From Allman’s Gymnoblastic Hydroids, by permission of the Council of the Ray Society.

Fig. 10.—Polyps from a Colony of Hydractinia, magnified. dz, dactylozoid; gz, gastrozoid; b, blastostyle; gon, gonophores; rh, hydrorhiza.

2. The Polyp-colony.—All known hydropolyps possess the power of reproduction by budding, and the buds produced may become either polyps or medusae. The buds may all become detached after a time and give rise to separate and independent individuals, as in the common Hydra, in which only polyp-individuals are produced and sexual elements are developed upon the polyps themselves; or, on the other hand, the polyp-individuals produced by budding may remain permanently in connexion with the parent polyp, in which case sexual elements are never developed on polyp-individuals but only on medusa-individuals, and a true colony is formed. Thus the typical hydroid colony starts from a “founder” polyp, which in the vast majority of cases is fixed, but which may be floating, as in Nemopsis, Pelagohydra, &c. The founder-polyp usually produces by budding polyp-individuals, and these in their turn produce other buds. The polyps are all non-sexual individuals whose function is purely nutritive. After a time the polyps, or certain of them, produce by budding medusa-individuals, which sooner or later develop sexual elements; in some cases, however, the founder-polyp remains solitary, that is to say, does not produce polyp-buds, but only medusa-buds, from the first (Corymorpha, fig. 3, Myriothela, &c.). In primitive forms the medusa-individuals are set free before reaching sexual maturity and do not contribute anything to the colony. In other cases, however, the medusa-individuals become sexually mature while still attached to the parent polyp, and are then not set free at all, but become appanages of the hydroid colony and undergo degenerative changes leading to reduction and even to complete obliteration of their original medusan structure. In this way the hydroid colony becomes composed of two portions of different function, the nutritive “trophosome,” composed of non-sexual polyps, and the reproductive “gonosome,” composed of sexual medusa-individuals, which never exercise a nutritive function while attached to the colony. As a general rule polyp-buds are produced from the hydrorhiza and hydrocaulus, while medusa-buds are formed on the hydranth. In some cases, however, medusa-buds are formed on the hydrorhiza, as in Hydrocorallines.

In such a colony of connected individuals, the exact limits of the separate “persons” are not always clearly marked out. Hence it is necessary to distinguish between, first, the “zooids,” indicated in the case of the polyps by the hydranths, each with mouth and tentacles; and, secondly, the “coenosarc,” or common flesh, which cannot be assigned more to one individual than another, but consists of a more or less complicated network of tubes, corresponding to the hydrocaulus and hydrorhiza of the primitive independent polyp-individual. The coenosarc constitutes a system by which the digestive cavity of any one polyp is put into communication with that of any other individual either of the trophosome or gonosome. In this manner the food absorbed by one individual contributes to the welfare of the whole colony, and the coenosarc has the