The Venous System.
The venous system, as it is found in the embryos of Fishes, consists in its earliest condition of a single large trunk, which traverses the splanchnic mesoblast investing the part of the alimentary tract behind the heart. This trunk is directly continuous in front with the heart, and underlies the alimentary canal through both its præanal and postanal sections. It is shown in section in [fig. 367], v, and may be called the subintestinal vein. This vein has been found in the embryos of Teleostei, Ganoidei, Elasmobranchii and Cyclostomata, and runs parallel to the dorsal aorta above, into which it is sometimes continued behind (Teleostei, Ganoidei, etc.).
In Elasmobranch embryos the subintestinal vein terminates, as may be gathered from sections ([fig. 368], v.cau), shortly before the end of the tail. The same series of sections also shews that at the cloaca, where the gut enlarges and comes in contact with the skin, this vein bifurcates, the two branches uniting into a single vein both in front of and behind the cloaca.
In most Fishes the anterior part of this vein atrophies, the caudal section alone remaining, but the anterior section of it persists in the fold of the intestine in Petromyzon, and also remains in the spiral valve of some Elasmobranchii. In Amphioxus, moreover, it forms, as in the embryos of higher types, the main venous trunk, though even here it is usually broken up into two or three parallel vessels.
It no doubt represents one of the primitive longitudinal trunks of the vermiform ancestors of the Chordata. The heart and the branchial artery constitute a specially modified anterior continuation of this vein. The dilated portal sinus of Myxine is probably also part of it; and if this is really rhythmically contractile[231] the fact would be interesting as shewing that this quality, which is now localised in the heart, was once probably common to the subintestinal vessel for its whole length.
Fig. 367. Section through the trunk of a Scyllium embryo slightly younger than 28 F.
sp.c. spinal canal; W. white matter of spinal cord; pr. posterior nerve-roots; ch. notochord; x. subnotochordal rod; ao. aorta; mp. muscle plate; mp´. inner layer of muscle-plate already converted into muscles; Vr. rudiment of vertebral body; st. segmental tube; sd. segmental duct; sp.v. spiral valve; v. subintestinal vein; p.o. primitive generative cells.
On the development of the cardinal veins (to be described below) considerable changes are effected in the subintestinal vein. Its postanal section, which is known in the adult as the caudal vein, unites with the cardinal veins. On this junction being effected retrogressive changes take place in the præanal section of the original subintestinal vessel. It breaks up in front into a number of smaller vessels, the most important of which is a special vein, which lies in the fold of the spiral valve, and which is more conspicuous in some Elasmobranchii than in Scyllium, in which the development of the vessel has been mainly studied. The lesser of the two branches connecting it round the cloaca with the caudal vein first vanishes, and then the larger; and the two posterior cardinals are left as the sole forward continuations of the caudal vein. The latter then becomes prolonged forwards, so that the two cardinals open into it some little distance in front of the hind end of the kidneys. By these changes, and by the disappearance of the postanal section of the gut, the caudal vein is made to appear as a supraintestinal and not, as it really is, a subintestinal vessel.
From the subintestinal vein there is given off a branch which supplies the yolk-sack. This leaves the subintestinal vein close to the liver. The liver, on its development, embraces the subintestinal vein, which then breaks up into a capillary system in the liver, the main part of its blood coming at this period from the yolk-sack.