1911 Encyclopædia Britannica/Coelentera
COELENTERA, a group or grade of the animal kingdom, the zoological importance of which has risen considerably since the time (1887) of the publication of the first article under that heading in the Ency. Brit. (9th edit.), even though their numbers have been reduced by the elevation of the Sponges or Porifera to the rank of an independent Phylum under the title Parazoa (W. J. Sollas, 1884). For the Coelentera thus restricted, the term Enterocoela, in contrast to Coelomocoela (the old Coelomata), was suggested by E. R. Lankester (1900).
From the more complex colonial Protozoa the Coelentera are readily separated by their possession of two distinct sets of cells, with diverse functions, arranged in two definite layers,—a condition found in no Protozoan. The old criterion by which they and other Metazoa were once distinguished from Protozoa, namely, the differentiation of large and small sexual cells from each other and from the remaining cells of the body, has been broken down by the discovery of numerous cases of such differentiation among Protozoa. The Coelentera, as contrasted with other Metazoa (but not Parazoa), consist of two layers of cells only, an outer layer or ectoderm, an inner layer or endoderm. They have hence been described as Diploblastica. In the remaining Metazoa certain cells are budded off at an early stage of development from one or both of the two original layers, to form later a third layer, the mesoderm, which lies between the ectoderm and endoderm; such forms have therefore received the name Triploblastica. At the same time it is necessary to observe that it is by no means certain that the mesoderm found in various groups of Metazoa is a similar or homologous formation in all cases. A second essential difference between Coelentera and other Metazoa (except Parazoa) is that in the former all spaces in the interior of the body are referable to a single cavity of endodermal origin, the “gastro-vascular cavity,” often termed the coelenteron: the spaces are always originally continuous with one another, and are in almost every case permanently so. This single cavity and its lining serve apparently for all those functions (digestion, excretion, circulation and often reproduction) which in more complex organisms are distributed among various cavities of independent and often very diverse origin.
In the Coelentera the ectoderm and endoderm are set apart from one another at a very early period in the life-history; generally either by delamination or invagination, processes described in the article Embryology. Between these two cell-layers a mesogloea (G. C. Bourne, 1887) is always intercalated as a secretion by one or both of them; this is a gelatinoid, primitively structureless lamella, which in the first instance serves merely as a basal support for the cells. In many cases, as, for example, in the Medusae or jelly-fish, the mesogloea may be so thick as to constitute the chief part of the body in bulk and weight. The ectoderm rarely consists of more than one layer of cells: these are divisible by structure and function into nervous, muscular and secretory cells, supported by interstitial cells. The endoderm is generally also an epithelium one cell in thickness, the cells being digestive, secretory and sometimes muscular. Reproductive sexual cells may be found in either of these two layers, according to the class and sub-class in question. The mesogloea is in itself an inert non-cellular secretion, but the immigration of muscular and other cells into its substance, from both ectoderm and endoderm, gives it in many cases a strong resemblance to the mesoderm of Triploblastica,—a resemblance which, while probably superficial, may yet serve to indicate the path of evolution of the mesoderm.
The Coelentera may thus be briefly defined as Metazoa which exhibit two embryonic cell-layers only,—the ectoderm and endoderm,—their body-cavities being referable to a single cavity or coelenteron in the endoderm. Their position in the animal kingdom and their main subdivisions may be expressed in the following table:—
In the above-given classification, the Scyphomedusae, formerly included with the Hydromedusae as Hydrozoa, are placed nearer the Anthozoa. The reasons for this may be stated briefly.
The Hydromedusae are distinguished from the Scyphozoa chiefly by negative characters; they have no stomodaeum, that is, no ingrowth of ectoderm at the mouth to form an oesophagus; they have no mesenteries (radiating partitions) which incompletely subdivide the coelenteron; and they have no concentration of digestive cells into special organs. Their ectodermal muscles are mainly longitudinal, their endodermal muscles are circularly arranged on the body-wall. Their sexual cells are (probably in all cases) produced from the ectoderm, and lie in those radii which are first accentuated in development. They typically present two structural forms, the non-sexual hydroid and the sexual medusoid; in such a case there is an alternation of generations (metagenesis), the hydroid giving rise to the medusoid by a sexual gemmation, the medusoid bearing sexual cells which develop into a hydroid. In some other cases medusoid develops directly from medusoid (hypogenesis), whether by sexual cells or by gemmation. The medusoids have a muscular velum of ectoderm and mesogloea only.
The Scyphozoa have the following features in common:—They typically exhibit an ectodermal stomodaeum; partitions or mesenteries project into their coelenteron from the body-wall, and on these are generally concentrated digestive cells (to form mesenterial filaments, phacellae or gastric filaments, &c.); the external musculature of the body-wall is circular (except in Cerianthus); the internal, longitudinal; and the sexual cells probably always arise in the endoderm.
The Scyphomedusae, like the Hydromedusae, typically present a metagenesis, the non-sexual scyphistomoid (corresponding to the hydroid) alternating with the sexual medusoid. In other cases the medusoid is hypogenetic, medusoid producing medusoid. The sexual cells of the medusoid lie in the endoderm on interradii, that is, on the second set of radii accentuated in the course of development. The medusoids have no true velum; in some cases a structure more or less resembling this organ, termed a velarium, is present, permeated by endodermal canals.
The Anthozoa differ from the Scyphomedusae in having no medusoid form; they all more or less resemble a sea-anemone, and may be termed actinioid. They are (with rare exceptions, probably secondarily acquired) hypogenetic, the offspring resembling the parent, and both being sexual. The sexual cells are borne on the mesenteries in positions irrespective of obvious developmental radii.
The Ctenophora are so aberrant in structure that it has been proposed to separate them from the Coelentera altogether: they are, however, theoretically deducible from an ancestor common to other Coelentera, but their extreme specialization precludes the idea of any close relationship with the rest.
As regards the other three groups, however, it is easy to conceive of them as derived from an ancestor, represented to-day to some extent by the planula-larva, which was Coelenterate in so far as it was composed of an ectoderm and endoderm, and had an internal digestive cavity (I. of the table).
At the point of divergence between Scyphozoa and Hydromedusae (II. of the table of hypothetical descent), we may conceive of its descendant as tentaculate, capable of either floating (swimming) or fixation at will like Lucernaria to-day; and exhibiting incipient differentiation of myoepithelial cells (formerly termed neuro-muscular cells). At the parting of the ways which led, on the one hand, to modern Scyphomedusae, on the other to Anthozoa (III.), it is probable that the common ancestor was marked by incipient mesenteries and by the limitation of the sexual cells to endoderm. The lines of descent—II. to Hydromedusae, and III. to Scyphomedusae—represent periods during which the hypothetical ancestors II. and III., capable of either locomotion or fixation at will, were either differentiated into alternating generations of fixed sterile nutritive hydroids (scyphistomoids) and locomotor sexual medusoids, or abandoned the power of fixation in hypogenetic cases. During the period represented by the line of descent—III. to Anthozoa—this group abandoned its power of adult locomotion by swimming. During these periods were also attained those less important structural characters which these three groups present to-day. (G. H. Fo.)