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144
[ORGANIZATION
HYDROMEDUSAE


body. The distal portion of the vitreous body may project from the cavity of the cup, forming a non-cellular lens as in Lizzia (fig. 28). Beyond this simple condition the visual organs of the Hydromedusae do not advance, and are far from reaching the wonderful development of the eyes of Scyphomedusae (Charybdaea).

EB1911 Hydromedusae - statocyst of Geryonia (Carmarina hastata).jpg

After O. and R. Hertwig, Nervensystem und Sinnesorgane der Medusen, by permission of F. C. W. Vogel.

Fig. 37.—Section of statocyst of Geryonia (Carmarina hastata).

st.c Statocyst containing the minute tentaculocyst.
nr₁, Ex-umbral nerve-ring.
nr₂, Sub-umbral nerve-ring.
ex, Ex-umbral ectoderm.
sub, Sub-umbral ectoderm.
c.c, Circular canal.
v, Velum.

Besides the ordinary type of ocellus just described, there is found in one genus (Tiaropsis) a type of ocellus in which the visual elements are inverted, and have their cones turned away from the light, as in the human retina (fig. 30). In this case the pigment-cells are endodermal, forming a cup of pigment in which the visual cones are embedded. A similar ocellus is formed in Aurelia among the Scyphomedusae (q.v.).

Other sense organs of Hydromedusae are the so-called sense-clubs or cordyli found in a few Leptomedusae, especially in those genera in which otocysts are inconspicuous or absent (fig. 39). Each cordylus is a tentacle-like structure with an endodermal axis containing an axial cavity which may be continuous with the ring-canal, or may be partially occluded. Externally the cordylus is covered, by very flattened ectoderm, and bears no otoliths or sense-cells, but the base of the club rests upon the ex-umbral nerve-ring. Brooks regards these organs as sensory, serving for the sense of balance, and representing a primitive stage of the tentaculocysts of Trachylinae; Linko, on the other hand, finding no nerve-elements connected with them, regards them as digestive (?) in function.

The sense-organs of the two fresh-water medusae Limnocodium and Limnocnida are peculiar and of rather doubtful nature (see E. T. Browne [10]).

EB1911 Hydromedusae - Ocellus of Lizzia koellikeri.jpg

Fig. 38.—Ocellus of Lizzia koellikeri. oc, Pigmented ectodermal cells; l, lens. (After Hertwig.)

The endoderm of the medusa shows the same general types of structure as in the polyp, described above. We can distinguish (1) digestive endoderm, in the stomach, often with special glandular elements; (2) circulatory endoderm, in the radial and ring-canals; (3) supporting endoderm in the axes of the tentacles and in the endoderm-lamella; the latter is primitively a double layer of cells, produced by concrescence of the ex-umbral and sub-umbral layers of the coelenteron, but it is usually found as a single layer of flattened cells (fig. 40); in Geryonia, however, it remains double, and the centripetal canals arise by parting of the two layers; (4) excretory endoderm, lining pores at the margin of the umbrella, occurring in certain Leptomedusae as so-called “marginal tubercles,” opening, on the one hand, into the ring-canal and, on the other hand, to the exterior by “marginal funnels,” which debouch into the sub-umbral cavity above the velum. As has been described above, the endoderm may also contribute to the sense-organs, but such contributions are always of an accessory nature, for instance, concrement-cells in the otocysts, pigment in the ocelli, and never of sensory nature, sense-cells being in all cases ectodermal.

The reproductive cells may be regarded as belonging primarily to neither ectoderm nor endoderm, though lodged in the ectoderm in all Hydromedusae. As described for the polyp, they are wandering cells capable of extensive migrations before reaching the particular spot at which they ripen. In the Hydromedusae they usually, if not invariably, ripen in the ectoderm, but in the neighbourhood of the main sources of nutriment, that is to say, not far from the stomach. Hence the gonads are found on the manubrium in Anthomedusae generally; on the base of the manubrium, or under the gastral pouches, or in both these situations (Octorchidae), or under the radial canals, in Trachomedusae; under the gastral pouches or radial canals, in Narcomedusae. When ripe, the germ-cells are dehisced directly to the exterior.

EB1911 Hydromedusae - a Cordylus of Laodice.jpg

After W. K. Brooks, Journal of Morphology, x., by permission of Ginn & Co.

Fig. 39.—Section of a Cordylus of Laodice.

c.c Circular canal.
v, Velum.
t, Tentacle.
c, Cordylus, composed of flattened ectoderm ec covering a large-celled endodermal axis en.

Hydromedusae are of separate sexes, the only known exception being Amphogona apsteini, one of the Trachomedusae (Browne [9]). Moreover, all the medusae budded from a given hydroid colony are either male or female, so that even the non-sexual polyp must be considered to have a latent sex. (In Hydra, on the other hand, the individual is usually hermaphrodite.) The medusa always reproduces itself sexually, and in some cases non-sexually also. The non-sexual reproduction takes the form of fission, budding or sporogony, the details of which are described below. Buds may be produced from the manubrium, radial canals, ring-canal, or tentacle-bases, or from an aboral stolon (Narcomedusae). In all cases only medusa-buds are produced, never polyp-buds.

The mesogloea of the medusa is largely developed and of great thickness in the umbrella. The sub-epithelial tissues, i.e. the nervous and muscular cells, are lodged in the mesogloea, but in Hydromedusae it never contains tissue-cells or mesogloeal corpuscles.

(b) The Medusae as a Subordinate Individuality.—It has been shown above that polyps are budded only from polyps and that the medusae may be budded either from polyps or from medusae. In any case the daughter-individuals produced from the buds may be imagined as remaining attached to the parent and forming a colony of individuals in organic connexion with one another, and thus three possible cases arise. The first case gives a colony entirely composed of polyps, as in many Hydroidea. The second case gives a colony partly composed of polyp-individuals, partly of medusa-individuals, a possibility also realized in many colonies of Hydroidea. The third case gives a colony entirely composed of medusa-individuals, a possibility perhaps realized in the Siphonophora, which will be discussed in dealing with this group.

EB1911 Hydromedusae - Portions of Sections through the Disks of Medusae Lizzia and Aurelia.jpg

Fig. 40.—Portions of Sections through the Disk of Medusae—the upper one of Lizzia, the lower of Aurelia. (After Hertwig.)

el, Endoderm lamella.
m, Muscular processes of the ectoderm-cells in cross section.
d, Ectoderm.
en Endoderm lining the enteric cavity.
e, Wandering endoderm cells of the gelatinous substance.

The first step towards the formation of a mixed hydroid colony is undoubtedly a hastening of the sexual maturity of the medusa-individual. Normally the medusae are liberated in quite an immature state; they swim away, feed, grow and become adult mature individuals. From the bionomical point of view, the medusa is to be considered as a means of spreading the species, supplementing the deficiencies of the sessile polyp. It may be, however, that increased reproductiveness becomes of greater importance to the species than wide diffusion; such a condition will be brought about if the medusae mature quickly and are either set free in a mature condition or remain in the shelter of the polyp-colony, protected from risks of a free life in the open sea. In this way the medusa sinks from an independent personality to an organ of the polyp-colony, becoming a so-called medusoid gonophore, or bearer of the reproductive organs, and losing gradually all organs necessary for an independent existence, namely those of sense, locomotion and nutrition.

In some cases both free medusae and gonophores may be produced from the same hydroid colony. This is the case in Syncoryne mirabilis (Allman [1], p. 278) and in Campanularia volubilis; in the latter, free medusae are produced in summer, gonophores in winter (Duplessis [14]). Again in Pennaria, the male medusae are set free