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

Transverse section of shark-embryos (through the region of the kidneys). (From Wijhe and Hertwig.) In Figure 1.162 the dorsal segment-cavities (h) are already separated from the body-cavity (lh), but they are connected a little earlier (Figure 1.161), nr neural tube, ch chorda, sch subchordal string, ao aorta, sk skeletal-plate, mp muscle-plate, cp cutis-plate, w connection of latter (growth-zone), vn primitive kidneys, ug prorenal duct, uk prorenal canals, us point where they are cut off, tr prorenal funnel, mk middle germ-layer (mk1 parietal, mk2 visceral), ik inner germ-layer (gut-gland layer).)

This typical articulation of the two coelom-sacs begins very early in the lancelet, before they are yet severed from the primitive gut, so that at first each segment-cavity (us) still communicates by a narrow opening with the gut, like an intestinal gland. But this opening soon closes by complete severance, proceeding regularly backwards. The closed segments then extend more, so that their upper half grows upwards like a fold between the ectoderm (ak) and neural tube (n), and the lower half between the ectoderm and alimentary canal (ch; Figure 1.82 d, left half of the figure). Afterwards the two halves completely separate, a lateral longitudinal fold cutting between them (mk, right half of Figure 1.82). The dorsal segments (sd) provide the muscles of the trunk the whole length of the body (1.159): this cavity afterwards disappears. On the other hand, the ventral parts give rise, from their uppermost section, to the pronephridia or primitive-kidney canals, and from the lower to the segmental rudiments of the sexual glands or gonads. The partitions of the muscular dorsal pieces (myotomes) remain, and determine the permanent articulation of the vertebrate organism. But the partitions of the large ventral pieces (gonotomes) become thinner, and afterwards disappear in part, so that their cavities run together to form the metacoel, or the simple permanent body-cavity.

The articulation proceeds in substantially the same way in the other vertebrates, the craniota, starting from the coelom-pouches. But whereas in the former case there is first a transverse division of the coelom-sacs (by vertical folds) and then the dorso-ventral division, the procedure is reversed in the craniota; in their case each of the long coelom-pouches first divides into a dorsal (primitive segment plates) and a ventral (lateral plates) section by a lateral longitudinal fold. Only the former are then broken up into primitive segments by the subsequent vertical folds; while the latter (segmented for a time in the amphioxus) remain undivided, and, by the divergence of their parietal and visceral plates, form a body-cavity that is unified from the first. In this case, again, it is clear that we must regard the features of the younger craniota as cenogenetically modified processes that can be traced palingenetically to the older acrania.

We have an interesting intermediate stage between the acrania and the fishes in these and many other respects in the cyclostoma (the hag and the lamprey, cf.

Chapter 2.

21).

(FIGURE 1.163. Frontal (or horizontal-longitudinal) section of a triton-embryo with three pairs of primitive segments. ch chorda, us primitive segments, ush their cavity, ak horn plate.)

Among the fishes the selachii, or primitive fishes, yield the most important information on these and many other phylogenetic questions (Figures 1.161 and 1.162). The careful studies of Ruckert, Van Wijhe, H.E. Ziegler, and others, have given us most valuable results. The products of the middle germinal layer are partly clear in these cases at the period when the dorsal primitive segment cavities (or myocoels, h) are still connected with the ventral body-cavity (lh; Figure 1.161). In Figure 1.162, a somewhat older embryo, these cavities are separated. The outer or lateral wall of the dorsal segment yields the cutis-plate (cp), the foundation of the connective corium. From its inner or median wall are developed the muscle-plate (mp, the rudiment of the trunk-muscles) and the skeletal plate, the formative matter of the vertebral column (sk).

In the amphibia, also, especially the water-salamander (Triton), we can observe very clearly the articulation of the coelom-pouches and the rise of the primitive segments from their dorsal half (cf. Figure 1.91, A, B, C). A horizontal longitudinal section of the salamander-embryo (Figure 1.163) shows very clearly the series of pairs of these vesicular dorsal segments, which have been cut off on each side from the ventral side-plates, and lie to the right and left of the chorda.

(FIGURE 1.164. The third cervical vertebra (human).

FIGURE 1.165. The sixth dorsal vertebra (human).

FIGURE 1.166. The second lumbar vertebra (human).)

The metamerism of the amniotes agrees in all essential points with that of the three lower classes of vertebrates we have considered; but it varies considerably in detail, in consequence of cenogenetic disturbances that are due in the first place (like the degeneration of the coelom-pouches) to the large development of the food-yelk. As the pressure of this seems to force the two middle layers together from the start, and as the solid structure of the mesoderm apparently belies the original hollow character of the sacs, the two sections of the mesoderm, which are at that time divided by the lateral fold--the dorsal segment-plates and ventral side-plates--have the appearance at first of solid layers of cells (Figures 1.94 to 1.97). And when the articulation of the somites begins in the sole-shaped embryonic shield, and a couple of protovertebrae are developed in succession, constantly increasing in number towards the rear, these cube-shaped somites (formerly called protovertebrae, or primitive vertebrae) have the appearance of solid dice, made up of mesodermic cells (Figure 1.93). Nevertheless, there is for a time a ventral cavity, or provertebral cavity, even in these solid "protovertebrae" (Figure 1.143 uwh). This vesicular condition of the provertebra is of the greatest phylogenetic interest; we must, according to the coelom theory, regard it as an hereditary reproduction of the hollow dorsal somites of the amphioxus (Figures 1.156 to 1.160) and the lower vertebrates (Figures 1.161 to 1.163). This rudimentary "provertebral cavity" has no physiological significance whatever in the amniote-embryo; it soon disappears, being filled up with cells of the muscular plate.

(FIGURE 1.167. Head of a shark embryo (Pristiurus), one-third of an inch long, magnified twenty times. (From Parker.) Seen from the ventral side.)

The innermost median part of the primitive segment plates, which lies immediately on the chorda (Figure 1.145 ch) and the medullary tube (m), forms the vertebral column in all the higher vertebrates (it is wanting in the lowest); hence it may be called the skeleton plate. In each of the provertebrae it is called the "sclerotome" (in opposition to the outlying muscular plate, the "myotome"). From the phylogenetic point of view the myotomes are much older than the sclerotomes. The lower or ventral part of each sclerotome (the inner and lower edge of the cube-shaped provertebra) divides into two plates, which grow round the chorda, and thus form the foundation of the body of the vertebra (wh). The upper plate presses between the chorda and the medullary tube, the lower between the chorda and the alimentary canal (Figure 1.137 C). As the plates of two opposite provertebral pieces unite from the right and left, a circular sheath is formed round this part of the chorda. From this develops the BODY of a vertebra--that is to say, the massive lower or ventral half of the bony ring, which is called the "vertebra" proper and surrounds the medullary tube (Figures 1.164 to 1.166). The upper or dorsal half of this bony ring, the vertebral arch (Figure 1.145 wb), arises in just the same way from the upper part of the skeletal plate, and therefore from the inner and upper edge of the cube-shaped primitive vertebra. As the upper edges of two opposing somites grow together over the medullary tube from right and left, the vertebra-arch becomes closed.

The whole of the secondary vertebra, which is thus formed from the union of the skeletal plates of two provertebral pieces and encloses a part of the chorda in its body, consists at first of a rather soft mass of cells; this afterwards passes into a firmer, cartilaginous stage, and finally into a third, permanent, bony stage. These three stages can generally be distinguished in the greater part of the skeleton of the higher vertebrates; at first most parts of the skeleton are soft, tender, and membranous; they then become cartilaginous in the course of their development, and finally bony.

(FIGURES 1.168 AND 1.169. Head of a chick embryo, of the third day. Figure 1.168 from the front, Figure 1.169 from the right. n rudimentary nose (olfactory pit), l rudimentary eye (optic pit, lens-cavity), g rudimentary ear (auditory pit), v fore-brain, gl eye-cleft. Of the three pairs of gill-arches the first has passed into a process of the upper jaw (o) and of the lower jaw (u). (From Kolliker.))

At the head part of the embryo in the amniotes there is not generally a cleavage of the middle germinal layer into provertebral and lateral plates, but the dorsal and ventral somites are blended from the first, and form what are called the "head-plates" (Figure 1.148 k). From these are formed the skull, the bony case of the brain, and the muscles and corium of the body. The skull develops in the same way as the membranous vertebral column. The right and left halves of the head curve over the cerebral vesicle, enclose the foremost part of the chorda below, and thus finally form a simple, soft, membranous capsule about the brain. This is afterwards converted into a cartilaginous primitive skull, such as we find permanently in many of the fishes. Much later this cartilaginous skull becomes the permanent bony skull with its various parts. The bony skull in man and all the other amniotes is more highly differentiated and modified than that of the lower vertebrates, the amphibia and fishes. But as the one has arisen phylogenetically from the other, we must assume that in the former no less than the latter the skull was originally formed from the sclerotomes of a number of (at least nine) head-somites.

While the articulation of the vertebrate body is always obvious in the episoma or dorsal body, and is clearly expressed in the segmentation of the muscular plates and vertebrae, it is more latent in the hyposoma or ventral body. Nevertheless, the hyposomites of the vegetal half of the body are not less important than the episomites of the animal half. The segmentation in the ventral cavity affects the following principal systems of organs: 1, the gonads or sex-glands (gonotomes); 2, the nephridia or kidneys (nephrotomes); and 3, the head-gut with its gill-clefts (branchiotomes).

(FIGURE 1.170. Head of a dog embryo, seen from the front. a the two lateral halves of the foremost cerebral vesicle, b rudimentary eye, c middle cerebral vesicle, de first pair of gill-arches (e upper-jaw process, d lower-jaw process), f, f apostrophe, f double apostrophe, second, third, and fourth pairs of gill-arches, g h i k heart (g right, h left auricle; i left, k right ventricle), l origin of the aorta with three pairs of arches, which go to the gill-arches. (From Bischoff.))

The metamerism of the hyposoma is less conspicuous because in all the craniotes the cavities of the ventral segments, in the walls of which the sexual products are developed, have long since coalesced, and formed a single large body-cavity, owing to the disappearance of the partition. This cenogenetic process is so old that the cavity seems to be unsegmented from the first in all the craniotes, and the rudiment of the gonads also is almost always unsegmented. It is the more interesting to learn that, according to the important discovery of Ruckert, this sexual structure is at first segmental even in the actual selachii, and the several gonotomes only blend into a simple sexual gland on either side secondarily.

(FIGURE 1.171. Human embryo of the fourth week (twenty-six days old), one-fourth of an inch in length magnified twenty times, showing: point of development of the hind-leg, umbilical cord (underneath it the tail, bent upwards), trigeminal nerve V Trigeminus, optic-muscle nerve III Oculo-motorius, rolling muscle nerve IV Trochlearis, rudiment of ear (labyrinthic