An opening appears in the test some time after the larva is fixed, leading into the mouth, which then becomes functional. The branchial sack at the same time undergoes important changes. In the larva it is provided with only two ciliated slits, which open into the, at this stage, paired atrial cavity ([fig. 10]).
The openings of the atrial cavity at first are shut off from communication with the exterior by the test, but not long after the larva becomes fixed, two perforations are formed in the test, which lead into the openings of the two atrial cavities. At the same time the atrial cavities dilate so as gradually to embrace the whole branchial sack to which their inner walls attach themselves. Shortly after this the branchial clefts rapidly increase in number[9].
The increase of the branchial clefts is somewhat complicated. Between the two primitive clefts two new ones appear, and then a third appears behind the last cleft. In the interval between each branchial cleft is placed a vascular branchial vessel ([fig. 8] VI. bb). Soon a great number of clefts become added in a row on each side of the branchial sack. These clefts are small ciliated openings placed transversely with reference to the long axis of the branchial sack, but only occupying a small part of the breadth of each side. The intervals dorsal and ventral to them are soon filled by series of fresh rows of slits, separated from each other by longitudinal bars. Each side of the branchial sack becomes in this way perforated by a number of small openings arranged in rows, and separated by transverse and longitudinal bars. The whole structure forms the commencement of the branchial basketwork of the adult; the arrangement of which differs considerably in structure and origin from the simple system of branchial clefts of normal vertebrate types. At the junction of the transverse and longitudinal bars papillæ are formed projecting into the lumen of the branchial sack.
Fig. 11. Diagram of a very young Ascidian. (From Lankester.)
After the above changes are far advanced towards completion, the openings of the two atrial sacks gradually approximate in the dorsal line, and finally coalesce to form the single atrial opening of the adult. The two atrial cavities at the same time coalesce dorsally to form a single cavity, which is continuous round the branchial sack, except along the ventral line where the endostyle is present. The atrial cavity, from its mode of origin as a pair of epiblastic involutions[10], is clearly a structure of the same nature as the branchial or atrial cavity of Amphioxus; and has nothing whatever to do with the true body cavity.
It has already been stated that the anus opens into the original left atrial cavity; when the two cavities coalesce the anus opens into the atrial cavity in the median dorsal line.
Two of the most obscure points in the development are the origin of the mesoblast in the trunk, and of the body cavity. Of the former subject we know next to nothing, though it seems that the cells resulting from the atrophy of the tail are employed in the nutrition of the mesoblastic structures of the trunk.
The body cavity in the adult is well developed in the region of the intestine, where it forms a wide cavity lined by an epithelioid mesoblastic layer. In the region of the branchial sack it is reduced to the vascular channels in the walls of the sack.