The Evolution of Man, V.2 by Ernst Haeckel (comprehension books .TXT) 📕

(FIGURE 2.363. Head of a fish-embryo, with rudimentary vascular system, from the left. dc Cuvier's duct (juncture of the anterior and posterior principal veins), sv venous sinus (enlarged end of Cuvier's duct), a auricle, v ventricle, abr trunk of branchial artery, s gill-clefts (arterial arches between), ad aorta, c carotid artery, n nasal pit. (From Gegenbaur.)

FIGURE 2.364. The five arterial arches of the Craniotes (1 to 5) in their original disposition, a arterial cone or bulb, a double apostrophe aorta-trunk, c carotid artery (foremost continuation of the roots of the aorta). (From Rathke.)

FIGURE 2.365. The five arterial arches of the birds; the lighter parts of the structure disappear; only the shaded parts remain. Letters as in Figure 2.364. s subclavian arteries, p pulmonary artery, p apostrophe branches of same, c apostrophe outer carotid, c double apostrophe inner carotid. (From Rathke.)

FIGURE 2.366. The five arterial arches of mammals; letters as in Figure 2.365. v vertebral artery, b Botall's duct (open in the embryo, closed afterwards). (From Rathke.))

As the new direction became permanent in the earlier Prochordonia, which gave rise to the Vertebrate stem, the two vessels that proceed from either end of the tubular heart acquired a fixed function. The foremost section of the ventral vessel henceforth always conveys blood from the heart, and so acts as an artery; the hind section of the same vessel brings the blood from the body to the heart, and so becomes a vein. In view of their relation to the two sections of the gut, we may call the latter the intestinal vein and the former the branchial artery. The blood contained in both vessels, and also in the heart, is venous or carbonised blood--i.e. rich in carbonic acid; on the other hand, the blood that passes from the gills into the dorsal vessel is provided with fresh oxygen--arterial or oxydised blood. The finest branches of the arteries and veins pass into each other in the tissues by means of a network of very fine, ventral, hair-like vessels, or capillaries (Figure 2.359).

When we turn from the Tunicates to the closely-related Amphioxus we are astonished at first to find an apparent retrogression in the formation of the vascular system. As we have seen, the Amphioxus has no real heart; its colourless blood is driven along in its vascular system by the principal vessel itself, which contracts regularly in its whole length (cf. Figure 2.210). A dorsal vessel that lies above the gut (aorta) receives the arterial blood from the gills and drives it into the body. Returning from here, the venous blood gathers in a ventral vessel under the gut (intestinal vein), and goes back to the gills. A number of branchial vascular arches, which effect respiration and rise in the wall of the branchial gut from belly to back, absorb oxygen from the water and give off carbonic acid; they connect the ventral with the dorsal vessel. As the same section of the ventral vessel, which also forms the heart in the Craniotes, has developed in the Ascidia into a simple tubular heart, we may regard the absence of this in the Amphioxus as a result of degeneration, a return in this case to the earlier form of the vascular system, as we find it in many of the worms. We may assume that the Acrania that really belong to our ancestral series did not share this retrogression, but inherited the one-chambered heart of the Prochordonia, and transmitted it directly to the earliest Craniotes (cf. the ideal Primitive Vertebrate, Prospondylus, Figures 1.98 to 1.102).

(FIGURES 2.367 TO 2.370. Metamorphosis of the five arterial arches in the human embryo (diagram from Rathke). la arterial cone, 1, 2, 3, 4, 5 first to fifth pair of arteries, ad trunk of aorta, aw roots of aorta. In Figure 2.367 only three, in Figure 2.368 all five, of the aortic arches are given (the dotted ones only are developed). In Figure 2.369 the first two pairs have disappeared again. In Figure 2.370 the permanent trunks of the artery are shown; the dotted parts disappear, s subclavian artery, v vertebral, ax axillary, c carotid (c apostrophe outer, c double apostrophe inner carotid), p pulmonary.)

The further phylogenetic evolution of the vascular system is revealed to us by the comparative anatomy of the Craniotes. At the lowest stage of this group, in the Cyclostomes, we find for the first time the differentiation of the vasorium into two sections: a system of blood-vessels proper, which convey the RED blood about the body, and a system of lymphatic vessels, which absorb the colourless lymph from the tissues and convey it to the blood. The lymphatics that absorb from the gut and pour into the blood-stream the milky food-fluid formed by digestion are distinguished by the special name of "chyle-vessels." While the chyle is white on account of its high proportion of fatty particles, the lymph proper is colourless. Both chyle and lymph contain the colourless amoeboid cells (leucocytes, Figure 1.12) that we also find distributed in the blood as colourless blood-cells (or "white corpuscles"); but the blood also contains a much larger quantity of red cells, and these give its characteristic colour to the blood of the Craniotes (rhodocytes, Figure 2.358). The distinction between lymph, chyle, and blood-vessels which is found in all the Craniotes may be regarded as an outcome of division of labour between various sections of our originally simple vascular system. In the Gnathostomes the spleen makes its first appearance, an organ rich in blood, the chief function of which is the extensive formation of new colourless and red cells. It is not found in the Acrania and Cyclostomes, or any of the Invertebrates. It has been transmitted from the earliest fishes to all the Craniotes.

The heart also, the central organ of circulation in all the Craniotes, shows an advance in structure in the Cyclostomes. The simple, spindle-shaped heart-tube, found in the same form in the embryo of all the Craniotes, is divided into two sections or chambers in the Cyclostomes, and these are separated by a pair of valves. The hind section, the auricle, receives the venous blood from the body and passes it on to the anterior section, the ventricle. From this it is driven through the trunk of the branchial artery (the foremost section of the ventral vessel or principal vein) into the gills.

In the Selachii an arterial cone is developed from the foremost end of the ventricle, as a special division, cut off by valves. It passes into the enlarged base of the trunk of the branchial artery (Figure 2.363 abr). On each side 5 to 7 arteries proceed from it. These rise between the gill-clefts (s) on the gill-arches, surround the gullet, and unite above into a common trunk-aorta, the continuation of which over the gut corresponds to the dorsal vessel of the worms. As the curved arteries on the gill-arches spread into a network of respiratory capillaries, they contain venous blood in their lower part (as arches of the branchial artery) and arterial blood in the upper part (as arches of the aorta). The junctures of the various aortic arches on the right and left are called the roots of the aorta. Of an originally large number of aortic arches there remain at first six, then (owing to degeneration of the fifth arch) only five, pairs; and from these five pairs (Figure 2.364) the chief parts of the arterial system develop in all the higher Vertebrates.

(FIGURE 2.371. Heart of a rabbit-embryo, from behind, a vitelline veins, b auricles of the heart, c atrium, d ventricle, e arterial bulb, f base of the three pairs of arterial arches. (From Bischoff.)

FIGURE 2.372. Heart of the same embryo (Figure 2.371), from the front. v vitelline veins, a auricle, ca auricular canal, l left ventricle, r right ventricle, ta arterial bulb. (From Bischoff.))

The appearance of the lungs and the atmospheric respiration connected therewith, which we first meet in the Dipneusts, is the next important step in vascular evolution. In the Dipneusts the auricle of the heart is divided by an incomplete partition into two halves. Only the right auricle now receives the venous blood from the veins of the body. The left auricle receives the arterial blood from the pulmonary veins. The two auricles have a common opening into the simple ventricle, where the two kinds of blood mix, and are driven through the arterial cone or bulb into the arterial arches. From the last arterial arches the pulmonary arteries arise (Figure 2.365 p). These force a part of the mixed blood into the lungs, the other part of it going through the aorta into the body.

From the Dipneusts upwards we now trace a progressive development of the vascular system, which ends finally with the loss of branchial respiration and a complete separation of the two halves of the circulation. In the Amphibia the partition between the two auricles is complete. In their earlier stages, as tadpoles (Figure 2.262), they have still the branchial respiration and the circulation of the fishes, and their heart contains venous blood alone. Afterwards the lungs and pulmonary vessels are developed, and henceforth the ventricle of the heart contains mixed blood. In the reptiles the ventricle and its arterial cone begin to divide into two halves by a longitudinal partition, and this partition becomes complete in the higher reptiles and birds on the one hand, and the stem-forms of the mammals on the other. Henceforth, the right half of the heart contains only venous, and the left half only arterial, blood, as we find in all birds and mammals. The right auricle receives its carbonised or venous blood from the veins of the body, and the right ventricle drives it through the pulmonary arteries into the lungs. From here the blood returns, as oxydised or arterial blood, through the pulmonary veins to the left auricle, and is forced by the left ventricle into the arteries of the body. Between the pulmonary arteries and veins is the capillary system of the small or pulmonary circulation. Between the body-arteries and veins is the capillary system of the large or body-circulation. It is only in the two highest classes of Vertebrates--the birds and mammals--that we find a complete division of the circulations. Moreover, this complete separation has been developed quite independently in the two classes, as the dissimilar formation of the aortas shows of itself. In the birds the RIGHT half of the fourth arterial arch has become the permanent arch (Figure 2.365). In the mammals this has been developed from the LEFT half of the same fourth arch (Figure 2.366).

(FIGURE 2.373. Heart and head of a dog-embryo, from the front, a fore brain, b eyes, c middle brain, d primitive lower jaw, e primitive upper jaw, f gill-arches, g right auricle, h left auricle, i left ventricle, k right ventricle. (From Bischoff.)

FIGURE 2.374. Heart of the same dog-embryo, from behind. a inosculation of the vitelline veins, b left auricle, c right auricle, d auricle, e auricular canal, f left ventricle, g right ventricle, h arterial bulb, (From Bischoff)

FIGURE 2.375. Heart of a human embryo, four weeks old; 1. front view, 2. back view, 3. opened, and upper half of the atrium removed. a apostrophe left auricle, a double apostrophe right auricle, v apostrophe left ventricle, v double apostrophe right ventricle, ao arterial bulb, c superior vena cava (cd right, cs left), s rudiment of the interventricular wall. (From Kolliker.)

FIGURE 2.376. Heart of a human embryo, six weeks old, front view. r right ventricle, t left ventricle, s furrow between ventricles, ta arterial bulb, af furrow on its surface; to right and left are the two large auricles. (From Ecker.)

FIGURE 2.377. Heart of a human embryo, eight weeks old, back view. a apostrophe left auricle, a double apostrophe right auricle, v