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140
[ORGANIZATION
HYDROMEDUSAE


most one; and the axis of the system is a false axis composed of portions of each of the consecutive polyps. In this method of budding there are two types. In one, the biserial type (fig. 15), the polyps produce buds right and left alternately, so that the hydranths are arranged in a zigzag fashion, forming a “scorpioid cyme,” as in Obelia and Sertularia. In the other, the uniserial type (fig. 16), the buds are formed always on the same side, forming a “helicoid cyme,” as in Hydrallmania, according to H. Driesch, in which, however, the primitively uniserial arrangement becomes masked later by secondary torsions of the hydranths.

EB1911 Hydromedusae - Diagram of sympodial budding, biserial type.jpg

Fig. 15.—Diagram of sympodial budding, biserial type, shown in five stages (1-5). F, founder-polyp; 1, 2, 3, 4, 5, 6, succession of polyps budded from the founder; a, b, c, second series of polyps budded from the founder; a³, b³, series budded from 3.

In a colony formed by sympodial budding, a polyp always produces first a bud, which contributes to the system to which it belongs, i.e. continues the stem or branch of which its parent forms a part. The polyp may then form a second bud, which becomes the starting point of a new system, the beginning, that is, of a new branch; and even a third bud, starting yet another system, may be produced from the same polyp. Hence the colonies of Calyptoblastea may be complexly branched, and the budding may be biserial throughout, uniserial throughout, or partly one, partly the other. Thus in Plumularidae (figs. 17, 18) there is formed a main stem by biserial budding; each polyp on the main stem forms a second bud, which usually forms a side branch or pinnule by uniserial budding. In this way are formed the familiar feathery colonies of Plumularia, in which the pinnules are all in one plane, while in the allied Antennularia the pinnules are arranged in whorls round the main biserial stem. The pinnules never branch again, since in the uniserial mode of budding a polyp never forms a second polyp-bud. On the other hand, a polyp on the main stem may form a second bud which, instead of forming a pinnule by uniserial budding, produces by biserial budding a branch, from which pinnules arise as from the main stem (fig. 18—3, 6). Or a polyp on the main stem, after having budded a second time to form a pinnule, may give rise to a third bud, which starts a new biserial system, from which uniserial pinnules arise as from the main stem—type of Aglaophenia (fig. 19). The laws of budding in hydroids have been worked out in an interesting manner by H. Driesch [13], to whose memoirs the reader must be referred for further details.

EB1911 Hydromedusae - Diagram of sympodial budding, uniserial type.jpg

Fig. 16.—Diagram of sympodial budding, uniserial type, shown in four stages (1-4). F, founder-polyp; 1, 2, 3, succession of polyps budded from the founder.

Individualization of Polyp-Colonies.—As in other cases where animal colonies are formed by organic union of separate individuals, there is ever a tendency for the polyp-colony as a whole to act as a single individual, and for the members to become subordinated to the needs of the colony and to undergo specialization for particular functions, with the result that they simulate organs and their individuality becomes masked to a greater or less degree. Perhaps the earliest of such specializations is connected with the reproductive function. Whereas primitively any polyp in a colony may produce medusa-buds, in many hydroid colonies medusae are budded only by certain polyps termed blastostyles (fig. 10, b). At first not differing in any way from other polyps (fig. 5), the blastostyles gradually lose their nutritive function and the organs connected with it; the mouth and tentacles disappear, and the blastostyle obtains the nutriment necessary for its activity by way of the coenosarc. In the Calyptoblastea, where the polyps are protected by special capsules of the perisarc, the gonothecae enclosing the blastostyles differ from the hydrothecae protecting the hydranths (fig. 54).

EB1911 Hydromedusae - Diagram of sympodial budding, simple unbranched Plumularia-type.jpg

Fig. 17.—Diagram of sympodial budding, simple unbranched Plumularia-type. F, founder; 1-8, main axis formed by biserial budding from founder; a-e, pinnule formed by uniserial budding from founder; a¹-d¹, branch formed by similar budding from 1; a²-d² from 2, and so forth.

In other colonies the two functions of the nutritive polyp, namely, capture and digestion of food, may be shared between different polyps (fig. 10). One class of polyps, the dactylozoids (dz), lose their mouth and stomach, and become elongated and tentacle-like, showing great activity of movement. Another class, the gastrozoids (gz), have the tentacles reduced or absent, but have the mouth and stomach enlarged. The dactylozoids capture food, and pass it on to the gastrozoids, which swallow and digest it.

EB1911 Hydromedusae - Diagram showing method of branching in the Plumularia-type.jpg

Fig. 18.—Diagram showing method of branching in the Plumularia-type; compare with fig. 17. Polyps 3 and 6, instead of producing uniserial pinnules, have produced biserial branches (3¹, 3², 3³, 3⁴; 6¹-6³), which give off uniserial branches in their turn.

Besides the three types of individual above mentioned, there are other appendages of hydroid colonies, of which the individuality is doubtful. Such are the “guard-polyps” (machopolyps) of Plumularidae, which are often regarded as individuals of the nature of dactylozoids, but from a study of the mode of budding in this hydroid family Driesch concluded that the guard-polyps were not true polyp-individuals, although each is enclosed in a small protecting cup of the perisarc, known as a nematophore. Again, the spines arising from the basal crust of Podocoryne have been interpreted by some authors as reduced polyps.

EB1911 Hydromedusae - Diagram showing method of branching in the Aglaophenia-type.jpg

Fig. 19.—Diagram showing method of branching in the Aglaophenia-type. Polyp 7 has produced as its first bud, 8; as its second bud, a⁷, which starts a uniserial pinnule; and as a third bud I⁷, which starts a biserial branch (II⁷-VI⁷) that repeats the structure of the main stem and gives off pinnules. The main stem is indicated by-·-·-·, the new stem by ······.

3. The Medusa.—In the Hydromedusae the medusa-individual occurs, as already stated, in one of two conditions, either as an independent organism leading a true life in the open seas, or as a subordinate individuality in the hydroid colony, from which it is never set free; it then becomes a mere reproductive appendage or gonophore, losing successively its organs of sense, locomotion and nutrition, until its medusoid nature and organization become scarcely recognizable. Hence it is convenient to consider the morphology of the medusa from these two aspects.

(a) The Medusa as an Independent Organism.—The general structure and characteristics of the medusa are described elsewhere (see articles Hydrozoa and Medusa), and it is only necessary here to deal with the peculiarities of the Hydromedusa.

As regards habit of life the vast majority of Hydromedusae are