Especially noteworthy in the germinal budding of Margellium is the formation of the entocodon, as in the vegetative budding of the indirect type.
5. Sporogony.—This method of reproduction has been described by E. Metchnikoff in Cunina and allied genera. In individuals either of the male or female sex, germ-cells which are quite undifferentiated and neutral in character, become amoeboid, and wander into the endoderm. They divide each into two sister-cells, one of which—the spore—becomes enveloped by the other. The spore-cell multiplies by division, while the enveloping cell is nutrient and protective. The spore cell gives rise to a “spore-larva,” which is set free in the coelenteron and grows into a medusa. Whether sporogony occurs also in the polyp or not remains to be proved.
6. Sexual Reproduction and Embryology.—The ovum of Hydromedusae is usually one of a large number of oögonia, and grows at the expense of its sister-cells. No regular follicle is formed, but the oöcyte absorbs nutriment from the remaining oögonia. In Hydra the oöcyte is a large amoeboid cell, which sends out pseudopodia amongst the oögonia and absorbs nutriment from them. When the oöcyte is full grown, the residual oögonia die off and disintegrate.
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| Fig. 47.—Budding from the Ectoderm (germinal epithelium) inMargellium. (After C. Chun.) | |
A, The epithelium becomes two-layered. B, The lower layer forms a solid mass of cells, which (C) becomes a vesicle, the future endoderm, containing the coelenteric cavity (coel), while the outer layer furnishes the future ectoderm. D, E, F, a thickening of the ectoderm on the distal side of the bud forms an entocodon (Gc). | G,H, Formation of the medusae. s.c, Sub-umbral cavity. r.c, Radial canal. st, Stomach, which in H acquires a secondary communication with the digestive cavity of the mother. cc, Circular canal. v, Velum. t, Tentacle. |
The spermatogenesis and maturation and fertilization of the germ-cells present nothing out of the common and need not be described here. These processes have been studied in detail by A. Brauer [2] for Hydra.
The general course of the development is described in the article [Hydrozoa]. We may distinguish the following series of stages: (1) ovum; (2) cleavage, leading to formation of a blastula; (3) formation of an inner mass or parenchyma, the future endoderm, by immigration or delamination, leading to the so-called parenchymula-stage; (4) formation of an archenteric cavity, the future coelenteron, by a splitting of the internal parenchyma, and of a blastopore, the future mouth, by perforation at one pole, leading to the gastrula-stage; (5) the outgrowth of tentacles round the mouth (blastopore), leading to the actinula-stage; and (6) the actinula becomes the polyp or medusa in the manner described elsewhere (see articles [Hydrozoa], [Polyp] and [Medusa]). This is the full, ideal development, which is always contracted or shortened to a greater or less extent. If the embryo is set free as a free-swimming, so-called planula-larva, in the blastula, parenchymula, or gastrula stage, then a free actinula stage is not found; if, on the other hand, a free actinula occurs, then there is no free planula stage.
The cleavage of the ovum follows two types, both seen in Tubularia (Brauer [3]). In the first, a cleavage follows each nuclear division; in the second, the nuclei multiply by division a number of times, and then the ovum divides into as many blastomeres as there are nuclei present. The result of cleavage in all cases is a typical blastula, which when set free becomes oval and develops a flagellum to each cell, but when not set free, it remains spherical in form and has no flagella.
The germ-layer formation is always by immigration or delamination, never by invagination. When the blastula is oval and free-swimming the inner mass is formed by unipolar immigration from the hinder pole. When the blastula is spherical and not set free, the germ-layer formation is always multipolar, either by immigration or by delamination, i.e. by tangential division of the cells of the blastoderm, as in Geryonia, or by a mixture of immigration and delamination, as in Hydra, Tubularia, &c. The blastopore is formed as a secondary perforation at one spot, in free-swimming forms at the hinder pole. Formation of archenteron and blastopore may, however, be deferred till a later stage (actinula or after).
The actinula stage is usually suppressed or not set free, but it is seen in Tubularia (fig. 48), where it is ambulatory, in Gonionemus (Trachomedusae), and in Cunina (Narcomedusae), where it is parasitic.