THE VASCULAR SYSTEM, THE LYMPHATICS, AND THE BLOOD-GLANDS
The Cyclostomata and Fishes possess a closed vascular system, consisting of a heart, arteries, capillaries, and veins, the whole forming a continuous series of blood-containing channels provided with definite limiting walls, through which the blood is propelled in a constant direction by the rhythmical contractions of the heart. In the course of the circulation the blood flows from the heart through a single large trunk, the ventral aorta, to the capillaries of the gills. From the gills the arterialised blood is collected into a large dorsally-situated vessel, the dorsal aorta, and thence is distributed through a system of arteries to the capillaries of the various organs of the body. Finally, the blood is collected from the capillaries and returned to the heart by the veins.
Although in most instances the organs of the body are supplied with arterialised blood conveyed to them by arteries, there are nevertheless cases in which an organ may receive venous blood by a vein in addition to arterial blood supplied by an artery. For example, the capillaries of the liver not only receive blood from the hepatic artery, but also venous blood by a large vein (hepatic portal vein), formed by the union of a number of smaller veins by which venous blood is collected from the capillaries of the stomach, intestine, spleen, and pancreas. In this and similar instances, where a vein formed by the union of the capillaries of an organ, or of a series of organs, instead of uniting with other veins and proceeding towards the heart, becomes continuous with a second set of capillaries in some other organ, a "portal" system is said to be formed, and in the particular example of the liver it is termed the "hepatic portal" system. A similar, or "renal portal," system also exists in connexion with the kidneys in the majority of Fishes.
There is little doubt that, primarily, the vascular system of Vertebrate animals consisted of a dorsal artery (dorsal aorta), running along the median dorsal line of the alimentary canal, and a ventral or subintestinal vein similarly related to the ventral surface of the digestive tube. The two vessels were connected by a series of pairs of lateral branches, which had their origins from the dorsal vessel, and, by their subdivision, formed a capillary network in the walls of the alimentary canal. From these networks paired veins issued and opened into the subintestinal vein. The simplicity of this primitive arrangement was somewhat disturbed in the region of the pharynx by the development of gill-clefts, in the walls of which the blood circulated for respiratory purposes from the ventral to the dorsal vessel; and also by the development of a hepatic portal circulation in connexion with the liver. In the latter instance the subintestinal vein entered the liver and subdivided into capillaries in the substance of that organ, the corresponding efferent vessel, or hepatic vein, becoming continuous with the anterior or pharyngeal section of the subintestinal vein, or, as it is usually termed, the ventral aorta. In this low grade of vascular system, which is perhaps most completely retained in Amphioxus, the circulation of the blood was probably effected by the wave-like contractions of more or fewer of the larger vessels; but subsequently a definite chambered heart was developed at the origin of the ventral aorta.
Of Fishes in general it may be said that the primitive dorsal and ventral vessels are present in the embryo, and for a time retain their original relations and physiological importance. To a very unequal extent they may also be retained in the adult, where, however, they co-exist with numerous other vessels, which the increasing differentiation of the body has called into existence. Thus, at a later period of embryonic life, the subintestinal vein becomes somewhat fragmentary. Its caudal section (caudal vein) ceases to be continuous with the precaudal portion, and the blood collected from the muscles and other structures of the tail is conveyed to the heart by a pair of posterior cardinal veins, which are either directly continuous with the caudal vein, or indirectly through the intervention of a renal portal system in the kidneys. The precaudal portion of the subintestinal vein is represented by a vein which runs forwards in the intestinal wall, and is one of the minor affluents of the hepatic portal vein, while its prehepatic section is represented in succession by the hepatic vein, the heart, and the ventral aorta. Of the additional veins which supplement these remnants of a primitively continuous subintestinal vein, the largest and most constant are (a) the posterior cardinal veins which, commencing in the kidneys and receiving the blood from those organs, pass forwards to the heart; (b) a pair of anterior cardinal veins, formed by the union of smaller veins from the head, including the brain, and passing backwards towards the heart. At the level of the latter organ each anterior cardinal vein joins the posterior cardinal of the same side of the body to form a short but wide transverse vessel, the Cuvierian duct or precaval vein, which opens into the hindermost of the cavities of the heart, viz. the sinus venosus; (c) a pair of inferior jugular veins by which the nutrient blood of the branchial apparatus is returned to the right and left Cuvierian ducts. In addition to these principal veins there may also be a pair of lateral veins collecting the blood from the lateral walls of the trunk, and also opening into the Cuvierian ducts; and subclavian and femoral veins from the pectoral and pelvic fins.
On the other hand, the primitive dorsal vessel (dorsal aorta), retains not only its original position and relations, but also its primary function as the main channel for the distribution of arterialised blood to all parts of the body. The system of lateral and probably segmentally arranged vessels, by which the dorsal and subintestinal vessels were connected in the primitive Vertebrata, have undergone considerable modification in all existing Fishes, but nevertheless retain much of their original disposition and relations in the pharyngeal region of the alimentary canal, where they are represented by the afferent and efferent vessels of the gills.
A more detailed account of the condition of the vascular system in the Cyclostomata and Fishes will now be given.
The Venous System.—The Cyclostomata,[[359]] as might be expected, exhibit a more primitive condition of the venous system in certain features than is the case in any other group. The precaudal portion of the subintestinal vein retains much of its original importance and runs in the rudimentary intestinal spiral valve as far as the liver, where it becomes the hepatic portal vein. From the liver the blood is collected into a single hepatic vein, and by it is conveyed to the sinus venosus. The caudal section of the subintestinal vein, now known as the caudal vein, bifurcates near the anus, and its two branches become directly continuous with the right and left posterior cardinals, without forming a renal portal system. In their forward course to the heart the posterior cardinals are situated directly beneath the notochord, and after receiving the blood from the kidneys and gonads, and from the numerous pairs of segmental veins of the body-wall, join the corresponding anterior cardinal veins, and form on each side a short transverse Cuvierian duct which opens into the sinus venosus. There is also a pair of inferior jugular veins which, however, unite opposite the fifth pair of gill-sacs to form a single trunk; this vessel is continued backwards, externally to the medio-ventral cartilage of the branchial basket, and finally opens directly into the sinus venosus.
In Elasmobranchs (e.g. Mustelus antarcticus)[[360]] the caudal vein (Fig. 186) lies in the haemal canal of the caudal portion of the vertebral column. On reaching the kidneys the vein divides into two renal portal veins, which, however, are not directly continuous with the posterior cardinal veins as in the Cyclostomata, but, on the contrary, after receiving the posterior segmental and oviducal veins, become continuous with the capillaries of the kidneys.
From the latter organs the blood is collected by a series of renal veins, and by them conveyed to the posterior cardinals, and thence to the Cuvierian ducts. In the adult, therefore, there is a well-developed renal portal system, but it is worthy of note, nevertheless, that this system is developed comparatively late in embryonic life, and that at an earlier stage the caudal vein is directly continuous with the two posterior cardinals, precisely as is the case in the Cyclostomata throughout life. The posterior cardinal veins are situated in the dorsal wall of the coelom (Fig. 187), beneath the vertebral column. For the hinder portion of their extent they are embedded in the kidneys (Fig. 186); and in this region the two veins are in close relation in the median line, and here and there freely communicate with each other. More anteriorly, they enlarge so much that they present the appearance of cavernous sinuses. In addition to the anterior segmental and oviducal veins, the posterior cardinals receive the spermatic or the ovarian vein from the male or female gonad.