The Evolution of Man, V.2 - Ernst Haeckel (little bear else holmelund minarik TXT) 📗

or hedgehog. This constant number, which has few exceptions (due to adaptation), is a strong proof of the common descent of the mammals; it can only be explained by faithful heredity from a common stem-form, a primitive mammal with seven cervical vertebrae. If each species had been created separately, it would have been better to have given the long-necked mammals more, and the short-necked animals less, cervical vertebrae. Next to these come the dorsal (or pectoral) vertebrae, which number twelve to thirteen (usually twelve) in man and most of the other mammals. Each dorsal vertebra (Figure 1.165) has at the side, connected by joints, a couple of ribs, long bony arches that lie in and protect the wall of the chest. The twelve pairs of ribs, together with the connecting intercostal muscles and the sternum, which joins the ends of the right and left ribs in front, form the chest (thorax). In this strong and elastic frame are the lungs, and between them the heart. Next to the dorsal vertebrae comes a short but stronger section of the column, formed of five large vertebrae. These are the lumbar vertebrae (Figure 1.166); they have no ribs and no holes in the transverse processes. To these succeeds the sacral bone, which is fitted between the two halves of the pelvic zone. The sacrum is formed of five vertebrae, completely blended together. Finally, we have at the end a small rudimentary caudal column, the coccyx. This consists of a varying number (usually four, more rarely three, or five or six) of small degenerated vertebrae, and is a useless rudimentary organ with no actual physiological significance. Morphologically, however, it is of great interest as an irrefragable proof of the descent of man and the anthropoids from long-tailed apes. On no other theory can we explain the existence of this rudimentary tail. In the earlier stages of development the tail of the human embryo protrudes considerably. It afterwards atrophies; but the relic of the atrophied caudal vertebrae and of the rudimentary muscles that once moved it remains permanently. Sometimes, in fact, the external tail is preserved. The older anatomists say that the tail is usually one vertebra longer in the human female than in the male (or four against five); Steinbach says it is the reverse.

(FIGURE 2.329. Three dorsal vertebrae, from a human embryo, eight weeks old, in lateral longitudinal section. v cartilaginous vertebral body, li inter-vertebral disks, ch chorda. (From Kolliker.)

(FIGURE 2.330. A dorsal vertebra of the same embryo, in lateral transverse section. cv cartilaginous vertebral body, ch chorda, pr transverse process, a vertebral arch (upper arch), c upper end of the rib (lower arch). (From Kolliker.))

In the human vertebral column there are usually thirty-three vertebrae. It is interesting to find, however, that the number often changes, one or two vertebrae dropping out or an additional one appearing. Often, also, a mobile rib is formed at the last cervical or the first lumbar vertebra, so that there are then thirteen dorsal vertebrae, besides six cervical and four lumbar. In this way the contiguous vertebrae of the various sections of the column may take each other's places.

In order to understand the embryology of the human vertebral column we must first carefully consider the shape and connection of the vertebrae. Each vertebra has, in general, the shape of a seal-ring (Figures 1.164 to 1.166). The thicker portion, which is turned towards the ventral side, is called the body of the vertebra, and forms a short osseous disk; the thinner part forms a semi-circular arch, the vertebral arch, and is turned towards the back. The arches of the successive vertebrae are connected by thin intercrural ligaments in such a way that the cavity they collectively enclose represents a long canal. In this vertebral canal we find the trunk part of the central nervous system, the spinal cord. Its head part, the brain, is enclosed by the skull, and the skull itself is merely the uppermost part of the vertebral column, distinctively modified. The base or ventral side of the vesicular cranial capsule corresponds originally to a number of developed vertebral bodies; its vault or dorsal side to their combined upper vertebral arches.

(FIGURE 2.331. Intervertebral disk of a new-born infant, transverse section. a rest of the chorda. (From Kolliker.))

While the solid, massive bodies of the vertebrae represent the real central axis of the skeleton, the dorsal arches serve to protect the central marrow they enclose. But similar arches develop on the ventral side for the protection of the viscera in the breast and belly. These lower or ventral vertebral arches, proceeding from the ventral side of the vertebral bodies, form, in many of the lower Vertebrates, a canal in which the large blood-vessels are enclosed on the lower surface of the vertebral column (aorta and caudal vein). In the higher Vertebrates the majority of these vertebral arches are lost or become rudimentary. But at the thoracic section of the column they develop into independent strong osseous arches, the ribs (costae). In reality the ribs are merely large and independent lower vertebral arches, which have lost their original connection with the vertebral bodies.

If we turn from this anatomic survey of the composition of the column to the question of its development, I may refer the reader to earlier pages with regard to the first and most important points (

Chapter 1.

14). It will be remembered that in the human embryo and that of the other vertebrates we find at first, instead of the segmented column, only a simple unarticulated cartilaginous rod. This solid but flexible and elastic rod is the axial rod (or the chorda dorsalis). In the lowest Vertebrate, the Amphioxus, it retains this simple form throughout life, and permanently represents the whole internal skeleton (Figure 2.210 i). In the Tunicates, also, the nearest Invertebrate relatives of the Vertebrates, we meet the same chorda--transitorily in the passing larva tail of the Ascidia, permanently in the Copelata (Figure 2.225 c). Undoubtedly both the Tunicates and Acrania have inherited the chorda from a common unsegmented stem-form; and these ancient, long-extinct ancestors of all the chordonia are our hypothetical Prochordonia.

Long before there is any trace of the skull, limbs, etc., in the embryo of man or any of the higher Vertebrates--at the early stage in which the whole body is merely a sole-shaped embryonic shield--there appears in the middle line of the shield, directly under the medullary furrow, the simple chorda. (Cf. Figures 1.131 to 1.135 ch). It follows the long axis of the body in the shape of a cylindrical axial rod of elastic but firm composition, equally pointed at both ends. In every case the chorda originates from the dorsal wall of the primitive gut; the cells that compose it (Figure 2.328 b) belong to the entoderm (Figures 2.216 to 2.221). At an early stage the chorda develops a transparent structureless sheath, which is secreted from its cells (Figure 2.328 a). This chordalemma is often called the "inner chorda-sheath," and must not be confused with the real external sheath, the mesoblastic perichorda.

(FIGURE 2.332. Human skull.

FIGURE 2.333. Skull of a new-born child. (From Kollmann.) Above, in the three bones of the roof of the skull, we see the lines that radiate from the central points of ossification; in front, the frontal bone; behind, the occipital bone; between the two the large parietal bone, p. s the scurf bone, w mastoid fontanelle, f petrous bone, t tympanic bone, l lateral part, b bulla, j cheek-bone, a large wing of cuneiform bone, k fontanelle of cuneiform bone.)

But this unsegmented primary axial skeleton is soon replaced by the segmented secondary axial skeleton, which we know as the vertebral column. The provertebral plates (Figure 1.124 s) differentiate from the innermost, median part of the visceral layer of the coelom-pouches at each side of the chorda. As they grow round the chorda and enclose it they form the skeleton plate or skeletogenetic layer--that is to say, the skeleton-forming stratum of cells, which provides the mobile foundation of the permanent vertebral column and skull (scleroblast). In the head-half of the embryo the skeletal plate remains a continuous, simple, undivided layer of tissue, and presently enlarges into a thin-walled capsule enclosing the brain, the primordial skull. In the trunk-half the provertebral plate divides into a number of homogeneous, cubical, successive pieces; these are the several primitive vertebrae. They are not numerous at first, but soon increase as the embryo grows longer (Figures 1.153 to 1.155).

(FIGURE 2.334. Head-skeleton of a primitive fish, n nasal pit, eth cribriform bone region, orb orbit of eye, la wall of auscultory labyrinth, occ occipital region of primitive skull, cv vertebral column, a fore, bc hind-lip cartilage, o primitive upper jaw (palato-quadratum), u primitive lower jaw, II hyaloid bone, III to VIII first to sixth branchial arches. (From Gegenbaur.)

FIGURE 2.335. Roofs of the skulls of nine Primates (Cattarrhines), seen from above and reduced to a common size. 1 European, 2 Brazilian, 3 Pithecanthropus, 4 Gorilla, 5 Chimpanzee, 6 Orang, 7 Gibbon, 8 Tailed ape, 9 Baboon.)

In all the Craniotes the soft, indifferent cells of the mesoderm, which originally compose the skeletal plate, are afterwards converted for the most part into cartilaginous cells, and these secrete a firm and elastic intercellular substance between them, and form cartilaginous tissue. Like most of the other parts of the skeleton, the membranous rudiments of the vertebrae soon pass into a cartilaginous state, and in the higher Vertebrates this is afterwards replaced by the hard osseous tissue with its characteristic stellate cells (Figure 1.6). The primary axial skeleton remains a simple chorda throughout life in the Acrania, the Cyclostomes, and the lowest fishes. In most of the other Vertebrates the chorda is more or less replaced by the cartilaginous tissue of the secondary perichorda that grows round it. In the lower Craniotes (especially the fishes) a more or less considerable part of the chorda is preserved in the bodies of the vertebrae. In the mammals it disappears for the most part. By the end of the second month in the human embryo the chorda is merely a slender thread, running through the axis of the thick, cartilaginous vertebral column (Figures 1.182 ch and 2.329 ch). In the cartilaginous vertebral bodies themselves, which afterwards ossify, the slender remnant of the chorda presently disappears (Figure 2.330 ch). But in the elastic inter-vertebral disks, which develop from the skeletal plate between each pair of vertebral bodies (Figure 2.329 li), a relic of the chorda remains permanently. In the new-born child there is a large pear-shaped cavity in each intervertebral disk, filled with a gelatinous mass of cells (Figure 2.331 a). Though less sharply defined, this gelatinous nucleus of the elastic cartilaginous disks persists throughout life in the mammals, but in the birds and most reptiles the last trace of the chorda disappears. In the subsequent ossification of the cartilaginous vertebra the first deposit of bony matter ("first osseous nucleus") takes place in the vertebral body immediately round the remainder of the chorda, and soon displaces it altogether. Then there is a special osseous nucleus formed in each half of the vertebral arch. The ossification does not reach the point at which the three nuclei are joined until after birth. In the first year the two osseous halves of the arches unite; but it is much later--in the second to the eighth year--that they connect with the osseous vertebral bodies.

(FIGURE 2.336. Skeleton of the breast-fin of Ceratodus (biserial feathered skeleton). A, B, cartilaginous series of the fin-stem. rr cartilaginous fin-radii. (From Gunther.)

FIGURE 2.337. Skeleton of the breast-fin of an early Selachius (Acanthias). The radii of the median fin-border (B) have disappeared for the most part; a few only (R) are left. R, R, radii of the lateral fin-border, mt metapterygium, ms mesopterygium, p propterygium. (From Gegenbaur.)

FIGURE 2.338. Skeleton of the breast-fin of a young Selachius. The radii of the median fin-border have wholly disappeared. The shaded part on the right is the section that persists in the five-fingered hand of the higher Vertebrates.