morphological peculiarities underlying the utmost diversity of superficial characters.
All Hydrozoa, in the first place, exhibit the three structural features distinctive of the Coelentera (q.v.). (1) The body is built up of two layers only, an external protective and sensory layer, the ectoderm, and an internal digestive layer, the endoderm. (2) The body contains but a single internal cavity, the coelenteron or gastrovascular space, which may be greatly ramified, but is not shut off into cavities distinct from the central digestive space. (3) The generative cells are produced in either the ectoderm or endoderm, and not in a third layer arising in the embryo, distinct from the two primary layers; in other words, there is no mesoderm or coelom.
To these three characters the Hydrozoa add a fourth which is distinctive of the subdivision of the Coelenterata termed the Cnidaria; that is to say, they always possess peculiar stinging organs known as nettle-cells, or nematocysts (Cnidae), each produced in a cell forming an integral part of the animal’s tissues. The Hydrozoa are thus shown to belong to the group of Coelenterata Cnidaria, and it remains to consider more fully their distinctive features, and in particular those which mark them off from the other main division of the Cnidaria, the Anthozoa (q.v.), comprising the corals and sea-anemones.
The great diversity, to which reference has already been made, in the form and structure of the Hydrozoa is due to two principal causes. In the first place, we find in this group two distinct types of person or individual, the polyp and the medusa (qq.v.), each capable of a wide range of variations; and when both polyp and medusa occur in the life-cycle of the same species, as is frequently the case, the result is an alternation of generations of a type peculiarly characteristic of the class. In the second place, the power of non-sexual reproduction by budding is practically of universal occurrence among the Hydrozoa, and by the buds failing to separate from the parent stock, colonies are produced, more or less complicated in structure and often of great size. We find that polyps may either bud other polyps or may produce medusae, and that medusae may bud medusae, though never, apparently, polyps. Hence we have a primary subdivision of the colonies of Hydrozoa into those produced by budding of polyps and those produced by budding of medusae. The former may contain polyp-persons and medusa-persons, either one kind alone or both kinds combined; the latter will contain only medusa-persons variously modified.
The morphology of the Hydrozoa reduces itself, therefore, to a consideration of the morphology of the polyp, of the medusa and of the colony. Putting aside the last-named, for a detailed account of which see Hydromedusae, we can best deal with the peculiarities of the polyp and medusa from a developmental point of view.
In the development of the Hydrozoa, and indeed of the Cnidaria generally, the egg usually gives rise to an oval larva which swims about by means of a coating of cilia on the surface of the body. This very characteristic larva is termed a planula, but though very uniform externally, the planulae of different species, or of the same species at different periods, do not always represent the same stage of embryonic development internally. On examining more minutely the course of the development, it is found that the ovum goes through the usual process of cleavage, always total and regular in this group, and so gives rise to a hollow sphere or ovoid with the wall composed of a single layer of cells, and containing a spacious cavity, the blastocoele or segmentation-cavity. This is the blastula stage occurring universally in all Metazoa, probably representing an ancestral Protozoan colony in phylogeny. Next the blastula gives rise to an internal mass of cells (fig. 1, hy) which come from the wall either by immigration (fig. 1, A) or by splitting off (delamination). The formation of an inner cell-mass converts the single-layered blastula (monoblastula) into a double-layered embryo (diblastula) which may be termed a parenchymula, since at first the inner cell-mass forms an irregular parenchyma which may entirely fill up and obliterate the segmentation cavity (fig. 1, B). At a later stage, however, the cells of the inner mass arrange themselves in a definite layer surrounding an internal cavity (fig. 1, C, al), which soon acquires an opening to the exterior at one pole, and so forms the characteristic embryonic stage of all Enterozoa known as the gastrula (fig. 2). In this stage the body is composed of two layers, ectoderm (d) externally, and endoderm (c) internally, surrounding a central cavity, the archenteron (b), which communicates with the exterior by a pore (a), the blastopore.
From Balfour, after Kowalewsky.
Fig. 1.—Formation of the Diblastula of Eucope (one of the
Calyptoblastic Hydromedusae) by immigration. A, B, C, three
successive stages. ep, Ectoderm; hy, endoderm; al, enteric cavity.
Thus a planula larva may be a blastula, or but slightly advanced beyond this stage, or it may be (and most usually is) a parenchymula; or in some cases (Scyphomedusae) it may be a gastrula. It should be added that the process of development, the gastrulation as it is termed, may be shortened by the immigration of cells taking place at one pole only, and in a connected layer with orderly arrangement, so that the gastrula stage is reached at once from the blastula without any intervening parenchymula stage. This is a process of gastrulation by invagination which is found in all animals above the Coelenterata, but which is very rare in the Cnidaria, and is known only in the Scyphomedusae amongst the Hydrozoa.
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| From Gegenbaur’s Elements of Comparative Anatomy. |
| Fig. 2.—Diagram of a Diblastula. a, Blastopore. |
After the gastrula stage, which is found as a developmental stage in all Enterozoa, the embryo of the Hydrozoa proceeds to develop characters which are peculiar to the Coelenterata only. Round the blastopore hollow outgrowths, variable in number, arise by the evagination of the entire body-wall, both ectoderm and endoderm. Each outgrowth contains a prolongation of the archenteric cavity (compare figs. 2 and 3, A). In this way is formed a ring of tentacles, the most characteristic organs of the Cnidaria. They surround a region which is termed the peristome, and which contains in the centre the blastopore, which becomes the adult mouth. The archenteron becomes the gastrovascular system or coelenteron. Between the ectoderm and endoderm a gelatinous supporting layer, termed the mesogloea, makes its appearance. The gastrula has now become an actinula, which may be termed the distinctive larva of the Cnidaria, and doubtless represents in a transitory manner the common ancestor of the group. In no case known, however, does the actinula become the adult, sexually mature individual, but always undergoes further modifications, whereby it develops into either a polyp or a medusa.
To become a polyp, the actinula (fig. 3, A) becomes attached to some firm object by the pole farthest from the mouth, and its growth preponderates in the direction of the principal axis, that is to say, the axis passing through the mouth (fig. 3, a-b). As a result the body becomes columnar in form (fig. 3, B), and without further change passes into the characteristic polyp-form (see Polyp).
Fig. 3.—Diagram showing the change of the Actinula (A) into a
Polyp (B); a-b, principal (vertical) axis; c-d, horizontal axis. The
endoderm is shaded, the ectoderm is left clear.
It is convenient to distinguish two types of polyp by the names hydropolyp and anthopolyp, characteristic of the Hydrozoa and
