The mesoblast, so far as is known, gives rise, in addition to the parts already mentioned, to the excretory organs, generative organs, vascular system, etc.
Excretory organs. There are found in the embryo of Nephelis and Hirudo certain remarkable provisional excretory organs the origin and history of which is not yet fully made out. In Nephelis they appear as one (according to Robin, No. [364]), or (according to Bütschli, No. [359]) as two successive pairs of convoluted tubes on the dorsal side of the embryo, which are stated by the latter author to develop from the scattered mesoblast cells underneath the skin. At their fullest development they extend, according to Robin, from close to the head to near the ventral sucker. Each of them is U-shaped, with the open end forwards, each limb of the U being formed by two tubes united in front. No external opening has been clearly made out. Semper believed that the tubes were continuous with the three posterior vitelline cells, but this has been shewn not to be the case. Fürbringer[148] is inclined from his own researches to believe that they open laterally. They contain a clear fluid.
In Hirudo, Leuckart (No. [362]) has described three similar pairs of organs the structure of which he has fully elucidated. They are situated in the posterior part of the body, and each of them commences with an enlargement from which a convoluted tube is continued for some distance backwards; it then turns forwards again and afterwards bends upon itself to open to the exterior. The anterior part is broken up into a kind of labyrinthic network.
The true segmental organs are found in a certain number of the segments and are stated (Whitman) to develop from groups of mesoblast cells. Their origin requires however further investigation.
A double row of colossal cells on each side of the body has been described in Clepsine by Whitman as derived from the mesoblastic plates. These cells ([fig. 58] B), which he calls segment-cells, lie opposite the walls of the septa. The inner row is stated to be connected with the segmental organs. Their eventual history is unknown, but they are conjectured by Whitman to be the mother cells of the testes.
The alimentary tract. This is formed primitively of two parts—the epiblastic stomodæum—forming mouth, pharynx, and œsophagus, and the hypoblastic mesenteron. The anus is formed very late as a simple perforation immediately dorsal to the posterior sucker.
In Clepsine, where there is an epibolic gastrula, the rudiment of the mesenteron is at first formed of the three vitelline spheres, from the surface of which a true hypoblastic layer enclosing a central yolk mass becomes differentiated, as already described. The mesenteric sack so formed is constricted by the growth of the mesoblastic septa into a series of lobes, while the posterior part forms a narrow and at first very short tube opening by the anus.
The lobed region forms the sacculated stomach of the adult. The sacculations of the stomach by their mode of origin necessarily correspond with the segments. In the adult however the anterior lobe is really double and has two divisions for the two segments it fills, while the posterior lobe, which, as is well known, extends backwards parallel with the rectum, is composed of five segmental sacculations. In connection with the stomodæum a protrusible pharynx is developed.
In Hirudo and Nephelis the mesenteron has from the first a sack-like form. The cells which compose the sack give rise to a secondary deposit of food-yolk. The further changes are practically the same as in Clepsine. In Hirudo the posterior sacculation of the stomach is primitively unpaired. The jaws are formed at about the same time as the eyes as protuberances on the wall of the oral cavity.
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