Some of the larvæ become fixed, while others coalesce together and form a large mass, the fate of which has not been further studied. An invagination of epiblast takes place at the free end of the fixed larva, which gives rise to the so-called gastric cavity, i.e. the axial portion of the general enteric cavity, which would appear to be in reality a kind of stomodæum. Around the gastric cavity the hypoblast forms eight mesenteries, the chambers between which are filled with the homogeneous material which occupied the centre of the ovum in the previous stage. It is to be presumed, though not stated, that by an absorption of the blind end of the stomodæal invagination the gastric chamber is placed in free communication with the spaces between the mesenteries[81]. During the next stage the young Alcyonium also acquires eight tentacles, which arise as hollow papillæ opening into the eight mesenteric chambers. By this stage also the matter filling up the mesenteric chambers is nearly absorbed.
Between the epiblast and hypoblast there is formed an homogeneous membrane, which penetrates in between the two layers of hypoblast which form the mesenteries. On the outer side of this membrane, and therefore presumably derived from the epiblast, is a layer of connective-tissue cells, which eventually gives rise to the abundant gelatinous tissue (cœnenchyma) in which the skeletal elements are deposited. In Sympodium coralloides Kowalevsky (No. [168]) has shewn still more completely the derivation of the stellate mesoblast cells from the epiblast. He finds that the calcareous spicula develop in these cells as in the mesoblast cells of sponges. The branched gastrovascular canals in this tissue are outgrowths of the primitive enteric cavity. A layer of circular muscles is formed at a late period from the epiblast, but the longitudinal muscles of the mesenteries on the inner side of the homogeneous membrane are regarded by Kowalevsky as hypoblastic.
A ciliated planula with delaminated hypoblast is also found in Gorgonia and Corallium rubrum. In the former genus at the time when the larva becomes fixed, the hypoblast is formed of two strata, an outer one of columnar cells, and an inner one of round ciliated cells lining a central enteric cavity. The inner layer is believed by Kowalevsky to become eventually absorbed and to be homologous with the inner granular mass of Alcyonium.
Zoantharia. Amongst the Zoantharia several forms have been investigated by Kowalevsky ([147]) and Lacaze Duthiers ([170]), of which some are stated by the former author to pass through an invaginate gastrula stage, while in other instances the hypoblast is probably formed by delamination.
To the first group belongs an edible form of Sea Anemone found near Messina, Cerianthus, and perhaps also Caryophyllium. In the first of these segmentation results in the formation of a blastosphere. A normal invagination obliterating the segmentation cavity then ensues, and the blastopore narrows to form the mouth. The borders of the mouth bend inwards and so give rise to the gastric cavity (stomodæum) which as in the Alcyonidæ is lined by epiblast. Simultaneously with the formation of the mouth there appear the two first mesenteries.
In Cerianthus the segmentation is unequal, the early stages are the same as in the Actinia just described, but the hypoblast cells give rise to a mass of fatty material filling up the enteric cavity, which becomes eventually absorbed.
In the majority of the Zoantharia so far investigated, including species of Actinia, Sagartia, Bunodes, Astroides, Astræa, the segmentation, which is often unequal[82] and not accompanied by the formation of a segmentation cavity, results in a solid two-layered ciliated planula. In these forms the impregnation takes place in the ovary, and the early stages of development are passed through in the maternal tissues.
One end of the planula becomes somewhat oval and develops a special bunch of cilia. At the other end a shallow depression appears, which becomes deeper and forms an involution lined by epiblast. This involution is the stomodæum, and becomes the so-called gastric cavity. The true enteric cavity lined by hypoblast is for some time filled with yolk material. The larva always swims with the aboral end directed forwards.
Between the two embryonic layers a homogeneous membrane is formed, similar to that already described in the Alcyonidæ.
The further development of the larvæ especially concerns the formation of mesenteries, tentacles and calcareous skeleton. With reference to this subject the observations of Lacaze Duthiers are especially valuable and striking.