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 term " diblastula " was subsequently adopted in England for the gastrula of Haeckel.
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.
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 Coelen- a terata only.
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.
Thus the development of the two types of individual seen in the Hydrozoa may be summarized as follows: - Egg Free Blastula "Planula" Parenchymula Stage I Gastrula Actinula 1 Polyp Medusa 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.
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.
The gastrula is formed in this case by invagination.
After the formation of the gastrula by epibole the larva becomes enclosed by an ectodermic test covering the whole of the original surface of the body, including the shell-gland, and leaving only a small opening at the posterior end in which the stomodaeum and proctodaeum are formed.
The gastrula is in these cases said to be formed by epibole.
Gastrula phase (optical section).
B, The Gastrula has become a Trochosphere by the development of the ciliated ring vr (optical section).
An interesting feature of the development of Chaetognaths is that, as in some insects, the cells destined to form the reproductive organs are differentiated at a very early period, being apparent even in the gastrula stage.
The four principal phases in the development are: (I) Blastula, (2) Gastrula, (3) Flagellate Embryo, (4) Larva.
Be noted with regard to the gastrula, in which it seems to differ from the gastrulae of invertebrata.
After invagination is completed, the embryo begins to elongate, the blastopore becomes narrower, and the dorsal wall of the gastrula loses its convexity, and becomes flattened to form the dorsal plate, the outer layer of which is the primordium of the neurochord and the inner layer the primordium of the notochord.
While still within the egg-membrane the epiblastic cells become flagellated, and the gastrula rotates within the membrane.
Io, B) in the gastrula stage, and the orifice of invagination or blastopore, which persists, is situated at the hinder pole.
F, nerve the gastrula stage.
The gastrula thus formed has a large blastopore, which is at first posterior but afterwards gradually moves towards the anterior end of the ventral surface.
Cleavage leads to the formation of an epibolic gastrula and ciliated embryo which hatches as a free-swimming larva remarkably like that of a Polychaete worm (D).
Regarding the Echinoderms as a whole in the light of the foregoing account, we may give the following analytic summary of the characters that distinguish them from other coelomate animals: They live in salt or brackish water; a primitive bilateral symmetry is still manifest in the right and left divisions of the coelom; the middle coelomic cavities are primitively transformed into two hydrocoels communicating with the exterior indirectly through a duct or ducts of the anterior coelom; stereom, composed of crystalline carbonate of lime, is, with few exceptions, deposited by special amoebocytes in the meshes of a mesodermal stroma, chiefly in the integument; reproductive cells are derived from the endothelium, apparently of the anterior coelom; total segmentation of the ovum produces a coeloblastula and gastrula by invagination; mesenchyme is formed in the segmentation cavity by migration of cells, chiefly from the hypoblast.
After the formation of an invaginate gastrula the larval form is rapidly acquired.
The segmentation is peculiar, and leads to the formation of a solid gastrula, consisting of a cortex of ectoderm nuclei surrounding a central endodermal mass, which is exposed at one point - the blastopore.
0 A A, Gastrula stage, ventral view, showing blastopore.
B, Older gastrula stage, ventral view, showing elongated blastopore and primitive streak.
If the three principal organ-systems of the medusa, namely mouth, tentacles and umbrella, be considered in the light of phylogeny, it is evident that the manubrium bearing the mouth must be the oldest, as representing a common property of all the Coelentera, even of the gastrula embryo of all Enterozoa.