In other higher vertebrates, also, the kidneys develop (though very differently formed later on) from similar structures, which have been secondarily derived from the segmental pronephridia of the acrania. The parts of the mesoderm at which the first traces of them are found are usually called the middle or mesenteric plates. As the first traces of the gonads make their appearance in the lining of these middle plates nearer inward (or the middle) from the inner funnels of the nephro-canals, it is better to count this part of the mesoderm with the hyposoma.
The chief and oldest organ of the vertebrate hyposoma, the alimentary canal, is generally described as an unsegmented organ. But we could just as well say that it is the oldest of all the segmented organs of the vertebrate; the double row of the coelom-pouches grows out of the dorsal wall of the gut, on either side of the chorda. In the brief period during which these segmental coelom-pouches are still openly connected with the gut, they look just like a double chain of segmented visceral glands. But apart from this, we have originally in all vertebrates an important articulation of the fore-gut, that is wanting in the lower gut, the segmentation of the branchial (gill) gut.
(FIGURE 1.172. Transverse section of the shoulder and fore-limb (wing) of a chick-embryo of the fourth day, magnified about twenty times. Beside the medullary tube we can see on each side three clear streaks in the dark dorsal wall, which advance into the rudimentary fore-limb or wing (e). The uppermost of them is the muscular plate; the middle is the hind and the lowest the fore root of a spinal nerve. Under the chorda in the middle is the single aorta, at each side of it a cardinal vein, and below these the primitive kidneys. The gut is almost closed. The ventral wall advances into the amnion, which encloses the embryo. (From Remak.)
FIGURE 1.173. Transverse section of the pelvic region and hind legs of a chick-embryo of the fourth day, magnified about forty times. h horn-plate, w medullary tube, n canal of the tube, u primitive kidneys, x chorda, e hind legs, b allantoic canal in the ventral wall, t aorta, v cardinal veins, a gut, d gut-gland layer, f gut-fibre layer, g embryonic epithelium, r dorsal muscles, c body-cavity or coeloma. (From Waldeyer.))
The gill-clefts, which originally in the older acrania pierced the wall of the fore-gut, and the gill-arches that separated them, were presumably also segmental, and distributed among the various metamera of the chain, like the gonads in the after-gut and the nephridia. In the amphioxus, too, they are still segmentally formed. Probably there was a division of labour of the hyposomites in the older (and long extinct) acrania, in such wise that those of the fore-gut took over the function of breathing and those of the after-gut that of reproduction. The former developed into gill-pouches, the latter into sex-pouches. There may have been primitive kidneys in both. Though the gills have lost their function in the higher animals, certain parts of them have been generally maintained in the embryo by a tenacious heredity. At a very early stage we notice in the embryo of man and the other amniotes, at each side of the head, the remarkable and important structures which we call the gill-arches and gill-clefts (Figures 1.167 to 1.170 f). They belong to the characteristic and inalienable organs of the amniote-embryo, and are found always in the same spot and with the same arrangement and structure. There are formed to the right and left in the lateral wall of the fore-gut cavity, in its foremost part, first a pair and then several pairs of sac-shaped inlets, that pierce the whole thickness of the lateral wall of the head. They are thus converted into clefts, through which one can penetrate freely from without into the gullet. The wall thickens between these branchial folds, and changes into an arch-like or sickle-shaped piece—the gill, or gullet-arch. In this the muscles and skeletal parts of the branchial gut separate; a blood-vessel arch rises afterwards on their inner side (Figure 1.98 ka). The number of the branchial arches and the clefts that alternate with them is four or five on each side in the higher vertebrates (Figure 1.170 d, f, f apostrophe, f double apostrophe). In some of the fishes (selachii) and in the cyclostoma we find six or seven of them permanently.
These remarkable structures had originally the function of respiratory organs—gills. In the fishes the water that serves for breathing, and is taken in at the mouth, still always passes out by the branchial clefts at the sides of the gullet. In the higher vertebrates they afterwards disappear. The branchial arches are converted partly into the jaws, partly into the bones of the tongue and the ear. From the first gill-cleft is formed the tympanic cavity of the ear.
There are few parts of the vertebrate organism that, like the outer covering or integument of the body, are not subject to metamerism. The outer skin (epidermis) is unsegmented from the first, and proceeds from the continuous horny plate. Moreover, the underlying cutis is also not metamerous, although it develops from the segmental structure of the cutis-plates (Figures 1.161 and 1.162 cp). The vertebrates are strikingly and profoundly different from the articulates in these respects also.
Further, most of the vertebrates still have a number of unarticulated organs, which have arisen locally, by adaptation of particular parts of the body to certain special functions. Of this character are the sense-organs in the episoma, and the limbs, the heart, the spleen, and the large visceral glands—lungs, liver, pancreas, etc.—in the hyposoma. The heart is originally only a local spindle-shaped enlargement of the large ventral blood-vessel or principal vein, at the point where the subintestinal passes into the branchial artery, at the limit of the head and trunk (Figures 1.170 and 1.171). The three higher sense-organs—nose, eye, and ear—were originally developed in the same form in all the craniotes, as three pairs of small depressions in the skin at the side of the head.
The organ of smell, the nose, has the appearance of a pair of small pits above the mouth-aperture, in front of the head (Figure 1.169 n). The organ of sight, the eye, is found at the side of the head, also in the shape of a depression (Figures 1.169 l and 1.170 b), to which corresponds a large outgrowth of the foremost cerebral vesicle on each side. Farther behind, at each side of the head, there is a third depression, the first trace of the organ of hearing (Figure 1.169 g). As yet we can see nothing of the later elaborate structure of these organs, nor of the characteristic build of the face.
(FIGURE 1.174. Development of the lizard's legs (Lacerta agilis), with special relation to their blood-vessels. 1, 3, 5, 7, 9, 11 right fore-leg; 13, 15 left fore-leg; 2, 4, 6, 8, 10, 12 right hind-leg; 14, 16 left hind-leg; SRV lateral veins of the trunk, VU umbilical vein. (From F. Hochstetter.))