Hydrozoa

Hydrozoa, one of the most widely spread and prolific groups of aquatic animals. They are for the most part marine in habitat, but a familiar fresh-water form is the common Hydra of ponds and ditches, which gives origin to the name of the class. The Hydrozoa comprise the hydroids, so abundant on all shores, most of which resemble vegetable organisms to the unassisted eye; the hydrocorallines, which, as their name implies, have a massive stony skeleton and resemble corals; the jelly-fishes so called; and the Siphonophora, of which the species best known by repute is the so-called “Portuguese man-of-war” (Physalia), dreaded by sailors on account of its terrible stinging powers.

In external form and appearance the Hydrozoa exhibit such striking differences that there would seem at first sight to be little in common between the more divergent members of the group. Nevertheless there is no other class in the animal kingdom with better marked characteristics, or with more uniform 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.  
  
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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.  
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From Gegenbaur’s Elements of Comparative Anatomy.  
Fig. 2.—Diagram of a Diblastula.  
  
a, Blastopore.

b, Archenteric cavity.

c, Endoderm.

d, Ectoderm.  
  
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.

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).  
  
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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.  
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Fig. 4.—Diagram showing the change of the Actinula into a Medusa. A, Vertical section of the actinula; a-b and c-d as in fig. 3, B, transitional stage, showing preponderating growth in the horizontal plane. C, C′, D, D′, two types of medusa organization; C and D are composite sections, showing a radius (R) on one side, an interradius (IR) on the other; C’ and D’ are plans; the mouth and manubrium are indicated at the centre, leading into the gastral cavity subdivided by the four areas of concrescence in each interradius (IR). t, tentacle; g.p, gastric pouch; r.c, radial canal not present in C and C′; c.c, circular or ring-canal; e.l, endoderm-lamella formed by concrescence. For a more detailed diagram of medusa-structure see article Medusa.  
  
It is convenient to distinguish two types of polyp by the names hydro polyp and anthopolyp, characteristic of the Hydrozoa and Anthozoa respectively. In the hydropolyp the body is typically elongated, the height of the column being far greater than the diameter. The peristome is relatively small and the mouth is generally raised on a projecting spout or hypostome. The ectoderm loses entirely the ciliation which it had in the planula and actinula stages and commonly secretes on its external surface a protective or supporting investment, the perisarc. Contrasting with this, the anthopolyp is generally of squat form, the diameter often exceeding the height; the peristome is wide, a hypostome is lacking, and the ectoderm, or so much of it as is exposed, i.e. not covered by secretion of skeletal or other investment, retains its ciliation throughout life. The internal structural differences are even more characteristic. In the hydropolyp the blastopore of the embryo forms the adult mouth situated at the extremity of the hypostome, and the ectoderm and endoderm meet at this point. In the anthopolyp the blastopore is carried inwards by an in-pushing of the body-wall of the region of the peristome, so that the adult mouth is an opening leading into a short ectodermal oesophagus or stomodaeum, at the bottom of which is the blastopore. Further, in the hydropolyp the digestive cavity either remains simple and undivided and circular in transverse section, or may show ridges projecting internally, which in this case are formed of endoderm alone, without any participation of the mesogloea. In the anthopolyp, on the other hand, the digestive cavity is always subdivided by so-called mesenteries, in-growths of the endoderm containing vertical lamellae of mesogloea (see Anthozoa). In short, the hydropolyp is characterized by a more simple type of organization than the anthopolyp, and is in most respects less modified from the actinula type of structure.

Returning now to the actinula, this form may, as already stated, develop into a medusa, a type of individual found only in the Hydrozoa, as here understood. To become a medusa, the actinula grows scarcely at all in the direction of the principal axis, but greatly along a plane at right angles to it. Thus the body becomes umbrella-shaped, the concave side representing the peristome, and the convex side the column, of the polyp. Hence the tentacles are found at the edge of the umbrella, and the hypostome forms usually a projecting tube, with the mouth at the extremity, forming the manubrium or handle of the umbrella. The medusa has a pronounced radial symmetry, and the positions of the primary tentacles, usually four in number, mark out the so-called radii, alternating with which are four interradii. The ectoderm retains its ciliation only in the sensory organs. The mesogloea becomes enormously increased in quantity (hence the popular name “jelly-fish”), and in correlation with this the endoderm-layer lining the coelenteron becomes pressed together in the interradial areas and undergoes concrescence, forming a more or less complicated gastrovascular system (see Medusa). It is sufficient to state here that the medusa is usually a free-swimming animal, floating mouth downwards on the open seas, but in some cases it may be attached by its aboral pole, like a polyp, to some firm basis, either temporarily or permanently.

Thus the development of the two types of individual seen in the Hydrozoa may be summarized as follows:—

This development, though probably representing the primitive sequence of events, is never actually found in its full extent, but is always abbreviated by omission or elimination of one or more of the stages. We have already seen that the parenchymula stage is passed over when the gastrulation is of the invaginate type. On the other hand, the parenchymula may develop directly into the actinula or even into the polyp, with suppression of the intervening steps. Great apparent differences may also be brought about by variations in the period at which the embryo is set free as a larva, and since two free-swimming stages, planula and actinula, are unnecessary, one or other of them is always suppressed. A good example of this is seen in two common genera of British hydroids, Cordylophora and Tabularia. In Cordylophora the embryo is set free at the parenchymula stage as a planula which fixes itself and develops into a polyp, both gastrula and actinula stages being suppressed. In Tubularia, on the other hand, the parenchymula develops into an actinula within the maternal tissues, and is then set free, creeps about for a time, and after fixing itself, changes into a polyp; hence in this case the planula-stage, as a free larva, is entirely suppressed.

The Hydrozoa may be defined, therefore, as Cnidaria in which two types of individual, the polyp and the medusa, may be present, each type developed along divergent lines from the primitive actinula form. The polyp (hydropolyp) is of simple structure and never has an ectodernal oesophagus or mesenteries.1 The general ectoderm loses its cilia, which persist only in the sensory cells, and it frequently secretes external protective or supporting structures. An internal mesogloeal skeleton is not found.

The class is divisible into two main divisions or sub-classes, Hydromedusae and Scyphomedusae, of which definitions and detailed systematic accounts will be found under these headings.

General Works on Hydrozoa.—C. Chun, “Coelenterata (Hohlthiere),” Bronn’s Klassen und Ordnungen des Thier-Reichs ii. 2 (1889 et seq.); Y. Delage, and E. Hérouard, Traité de zoologie concrète, ii. part 2, Les Coelentérés (1901); G. H. Fowler, “The Hydromedusae and Scyphomedusae” in E. R. Lankester’s Treatise on Zoology, ii. chapters iv. and v. (1900); S. J. Hickson, “Coelenterata and Ctenophora,” Cambridge Natural History, i. chapters x.-xv. (1906).

(E. A. M.)

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1 See further under Scyphomedusae.

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