The Evolution of Man, V.1., Ernst Haeckel [ebook pdf reader for pc .TXT] 📗

ch axial rod or chorda, sh chorda-sheath, bh ventral wall, g hind and v fore root of the spinal nerves, a = af = am amniotic fold, p body-cavity or coeloma, df gut-fibre layer, ao primitive aortas, sa secondary aorta, vc cardinal veins, d = dd gut-gland layer, dr gastric groove. In Figure 1.143 the larger part of the right half, in Figure 1.144 the larger part of the left half, of the section is omitted. Of the yelk-sac or remainder of the embryonic vesicle only a small piece of the wall is indicated below.)

We find in the construction of the ventral wall precisely the same processes as in the formation of the dorsal wall (Figure 1.137 B, Figure 1.144 hp, Figure 1.146 bh). It is formed on the flat embryonic shield of the amniotes from the upper plates of the parietal zone. The right and left parietal plates bend downwards towards each other, and grow round the gut in the same way as the gut itself closes. The outer part of the lateral plates forms the ventral wall or the lower wall of the body, the two lateral plates bending considerably on the inner side of the amniotic fold, and growing towards each other from right and left. While the alimentary canal is closing, the body-wall also closes on all sides. Hence the ventral wall, which encloses the whole ventral cavity below, consists of two parts, two lateral plates that bend towards each other. These approach each other all along, and at last meet at the navel. We ought, therefore, really to distinguish two navels, an inner and an outer one. The internal or intestinal navel is the definitive point of the closing of the gut wall, which puts an end to the open communication between the ventral cavity and the cavity of the yelk-sac (Figure 1.105). The external navel in the skin is the definitive point of the closing of the ventral wall; this is visible in the developed body as a small depression.

(FIGURE 1.147. Median longitudinal section of the embryo of a chick (fifth day of incubation), seen from the right side (head to the right, tail to the left). Dorsal body dark, with convex outline. d gut, o mouth, a anus, l lungs, h liver, g mesentery, v auricle of the heart, k ventricle of the heart, b arch of the arteries, t aorta, c yelk-sac, m vitelline (yelk) duct, u allantois, r pedicle (stalk) of the allantois, n amnion, w amniotic cavity, s serous membrane. (From Baer.))

With the formation of the internal navel and the closing of the alimentary canal is connected the formation of two cavities, which we call the capital and the pelvic sections of the visceral cavity. As the embryonic shield lies flat on the wall of the embryonic vesicle at first, and only gradually separates from it, its fore and hind ends are independent in the beginning; on the other hand, the middle part of the ventral surface is connected with the yelk-sac by means of the vitelline or umbilical duct (Figure 1.147 m). This leads to a notable curving of the dorsal surface; the head-end bends downwards towards the breast and the tail-end towards the belly. We see this very clearly in the excellent old diagrammatic illustration given by Baer (Figure 1.147), a median longitudinal section of the embryo of the chick, in which the dorsal body or episoma is deeply shaded. The embryo seems to be trying to roll up, like a hedgehog protecting itself from its pursuers. This pronounced curve of the back is due to the more rapid growth of the convex dorsal surface, and is directly connected with the severance of the embryo from the yelk-sac. The further bending of the embryo leads to the formation of the "head-cavity" of the gut (Figure 1.148 above D) and a similar one at the tail, known as its "pelvic cavity."

As a result of these processes the embryo attains a shape that may be compared to a wooden shoe, or, better still, to an overturned canoe. Imagine a canoe or boat with both ends rounded and a small covering before and behind; if this canoe is turned upside down, so that the curved keel is uppermost, we have a fair picture of the canoe-shaped embryo (Figure 1.147). The upturned convex keel corresponds to the middle line of the back; the small chamber underneath the fore-deck represents the capital cavity, and the small chamber under the rear-deck the pelvic chamber of the gut (cf. Figure 1.140).

The embryo now, as it were, presses into the outer surface of the embryonic vesicle with its free ends, while it moves away from it with its middle part. As a result of this change the yelk-sac becomes henceforth only a pouch-like outer appendage at the middle of the ventral wall. The ventral appendage, growing smaller and smaller, is afterwards called the umbilical (navel) vesicle. The cavity of the yelk-sac or umbilical vesicle communicates with the corresponding visceral cavity by a wide opening, which gradually contracts into a narrow and long canal, the vitelline (yelk) duct (ductus vitellinus, Figure 1.147 m). Hence, if we were to imagine ourselves in the cavity of the yelk-sac, we could get from it through the yelk-duct into the middle and still wide open part of the alimentary canal. If we were to go forward from there into the head-part of the embryo, we should reach the capital cavity of the gut, the fore-end of which is closed up.

The reader will ask: "Where are the mouth and the anus?" These are not at first present in the embryo. The whole of the primitive gut-cavity is completely closed, and is merely connected in the middle by the vitelline duct with the equally closed cavity of the embryonic vesicle (Figure 1.140). The two later apertures of the alimentary canal--the anus and the mouth--are secondary constructions, formed from the outer skin. In the horn-plate, at the spot where the mouth is found subsequently, a pit-like depression is formed, and this grows deeper and deeper, pushing towards the blind fore-end of the capital cavity; this is the mouth-pit. In the same way, at the spot in the outer skin where the anus is afterwards situated a pit-shaped depression appears, grows deeper and deeper, and approaches the blind hind-end of the pelvic cavity; this is the anus-pit. In the end these pits touch with their deepest and innermost points the two blind ends of the primitive alimentary canal, so that they are now only separated from them by thin membranous partitions. This membrane finally disappears, and henceforth the alimentary canal opens in front at the mouth and in the rear by the anus (Figures 1.141 and 1.147). Hence at first, if we penetrate into these pits from without, we find a partition cutting them off from the cavity of the alimentary canal, which gradually disappears. The formation of mouth and anus is secondary in all the vertebrates.

(FIGURE 1.148. Longitudinal section of the fore half of a chick-embryo at the end of the first day of incubation (seen from the left side). k head-plates, ch chorda. Above it is the blind fore-end of the ventral tube (m); below it the capital cavity of the gut. d gut-gland layer, df gut-fibre layer, h horn plate, hh cavity of the heart, hk heart-capsule, ks head-sheath, kk head-capsule. (From Remak.))

During the important processes which lead to the formation of the navel, and of the intestinal wall and ventral wall, we find a number of other interesting changes taking place in the embryonic shield of the amniotes. These relate chiefly to the prorenal ducts and the first blood-vessels. The prorenal (primitive kidney) ducts, which at first lie quite flat under the horn-plate or epiderm (Figure 1.93 ung), soon back towards each other in consequence of special growth movements (Figures 1.143 to 1.145 ung). They depart more and more from their point of origin, and approach the gut-gland layer. In the end they lie deep in the interior, on either side of the mesentery, underneath the chorda, (Figure 1.145 ung). At the same time, the two primitive aortas change their position (cf. Figures 1.138 to 1.145 ao); they travel inwards underneath the chorda, and there coalesce at last to form a single secondary aorta, which is found under the rudimentary vertebral column (Figure 1.145 ao). The cardinal veins, the first venous blood-vessels, also back towards each other, and eventually unite immediately above the rudimentary kidneys (Figures 1.145 vc, 152 cav). In the same spot, at the inner side of the fore-kidneys, we soon see the first trace of the sexual organs. The most important part of this apparatus (apart from all its appendages) is the ovary in the female and the testicle in the male. Both develop from a small part of the cell-lining of the body-cavity, at the spot where the skin-fibre layer and gut-fibre layer touch. The connection of this embryonic gland with the prorenal ducts, which lie close to it and assume most important relations to it, is only secondary.

(FIGURE 1.149. Longitudinal section of a human embryo of the fourth week, one-fifth of an inch long, magnified fifteen times. Showing: bend of skull, yelk-sac, umbilical cord, terminal gut, rudimentary kidneys, mesoderm, head-gut (with gill-clefts), primitive lungs, liver, stomach, pancreas, mesentery, primitive kidneys, allantoic duct, rectum. (From Kollmann.)

FIGURE 1.150. Transverse section of a human embryo of fourteen days. mr medullary tube, ch chorda. vu umbilical vein, mt myotome, mp middle plate, ug prorenal duct, lh body-cavity, e ectoderm, bh ventral skin, hf skin-fibre layer, df gut-fibre layer. (From Kollmann.)

FIGURE 1.151. Transverse section of a shark-embryo (or young selachius). mr medullary tube, ch chorda, a aorta, d gut, vp principal (or subintestinal) vein, mt myotome, mm muscular mass of the provertebra, mp middle plate, ug prorenal duct, lh body-cavity, e ectoderm of the rudimentary extremities, mz mesenchymic cells, z point where the myotome and nephrotome separate. (From H.E. Ziegler.)

FIGURE 1.152. Transverse section of a duck-embryo with twenty-four primitive segments. (From Balfour.) From a dorsal lateral joint of the medullary tube (spc) the spinal ganglia (spg) grow out between it and the horn-plate. ch chorda, ao double aorta, hy gut-gland layer, sp gut-fibre layer, with blood-vessels in section, ms muscle plate, in the dorsal wall of the myocoel (episomite). Below the cardinal vein (cav) is the prorenal duct (wd) and a segmental prorenal canal (st). The skin-fibre layer of the body-wall (so) is continued in the amniotic fold (am). Between the four secondary germinal layers and the structures formed from them there is formed embryonic connective matter with stellate cells and vascular structures (Hertwig's "mesenchym").)

CHAPTER XIV(14. THE ARTICULATION OF THE BODY.*)

 

(* The term articulation is used in this chapter to denote both "segmentation" and "articulation" in the ordinary sense.--Translator.)

The vertebrate stem, to which our race belongs as one of the latest and most advanced outcomes of the natural development of life, is rightly placed at the head of the animal kingdom. This privilege must be accorded to it, not only because man does in point of fact soar far above all other animals, and has been lifted to the position of "lord of creation"; but also because the vertebrate organism far surpasses all the other animal-stems