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

(b the three basal pieces of the fin: mt metapterygium, rudiment of the humerus, ms mesopterygium, p propterygium.) (From Gegenbaur.))

The bony skull (cranium), the head-part of the secondary axial skeleton, develops in just the same way as the vertebral column. The skull forms a bony envelope for the brain, just as the vertebral canal does for the spinal cord; and as the brain is only a peculiarly differentiated part of the head, while the spinal cord represents the longer trunk-section of the originally homogeneous medullary tube, we shall expect to find that the osseous coat of the one is a special modification of the osseous envelope of the other. When we examine the adult human skull in itself (Figure 2.332), it is difficult to conceive how it can be merely the modified fore part of the vertebral column. It is an elaborate and extensive bony structure, composed of no less than twenty bones of different shapes and sizes. Seven of them form the spacious shell that surrounds the brain, in which we distinguish the solid ventral base below and the curved dorsal vault above. The other thirteen bones form the facial skull, which is especially the bony envelope of the higher sense-organs, and at the same time encloses the entrance of the alimentary canal. The lower jaw is articulated at the base of the skull (usually regarded as the XXI cranial bone). Behind the lower jaw we find the hyoid bone at the root of the tongue, also formed from the gill-arches, and a part of the lower arches that have developed as "head-ribs" from the ventral side of the base of the cranium.

Although the fully-developed skull of the higher Vertebrates, with its peculiar shape, its enormous size, and its complex composition, seems to have nothing in common with the ordinary vertebrae, nevertheless even the older comparative anatomists came to recognise at the end of the eighteenth century that it is really nothing else originally than a series of modified vertebrae. When Goethe in 1790 "picked up the skull of a slain victim from the sand of the Jewish cemetery at Venice, he noticed at once that the bones of the face also could be traced to vertebrae (like the three hind-most cranial vertebrae)." And when Oken (without knowing anything of Goethe's discovery) found at Ilenstein, "a fine bleached skull of a hind, the thought flashed across him like lightning: 'It is a vertebral column.'"

(FIGURE 2.339. Skeleton of the fore leg of an amphibian. h upper-arm (humerus), ru lower arm (r radius, u ulna), rcicu apostrophe, wrist-bones of first series (r radiale, i intermedium, c centrale, u apostrophe ulnare). 1, 2, 3, 4, 5 wrist-bones of the second series. (From Gegenbaur.)

FIGURE 2.340. Skeleton of gorilla's hand. (From Huxley.)

FIGURE 2.341. Skeleton of human hand, back. (From Meyer.))

This famous vertebral theory of the skull has interested the most distinguished zoologists for more than a century: the chief representatives of comparative anatomy have devoted their highest powers to the solution of the problem, and the interest has spread far beyond their circle. But it was not until 1872 that it was happily solved, after seven years' labour, by the comparative anatomist who surpassed all other experts of this science in the second half of the nineteenth century by the richness of his empirical knowledge and the acuteness and depth of his philosophic speculations. Carl Gegenbaur has shown, in his classic Studies of the Comparative Anatomy of the Vertebrates (third section), that we find the most solid foundation for the vertebral theory of the skull in the head-skeleton of the Selachii. Earlier anatomists had wrongly started from the mammal skull, and had compared the several bones that compose it with the several parts of the vertebra (Figure 2.333) they thought they could prove in this way that the fully-formed mammal skull was made of from three to six vertebrae.

The older theory was refuted by simple and obvious facts, which were first pointed out by Huxley. Nevertheless, the fundamental idea of it--the belief that the skull is formed from the head-part of the perichordal axial skeleton, just as the brain is from the simple medullary tube, by differentiation and modification--remained. The work now was to discover the proper way of supplying this philosophic theory with an empirical foundation, and it was reserved for Gegenbaur to achieve this. He first opened out the phylogenetic path which here, as in all morphological questions, leads most confidently to the goal. He showed that the primitive fishes (Figures 2.249 to 2.251), the ancestors of all the Gnathostomes, still preserve permanently in the form of their skull the structure out of which the transformed skull of the higher Vertebrates, including man, has been evolved. He further showed that the branchial arches of the Selachii prove that their skull originally consisted of a large number of (at least nine or ten) provertebrae, and that the cerebral nerves that proceed from the base of the brain entirely confirm this. These cerebral nerves are (with the exception of the first and second pair, the olfactory and optic nerves) merely modifications of spinal nerves, and are essentially similar to them in their peripheral expansion. The comparative anatomy of these cerebral nerves, their origin and their expansion, furnishes one of the strongest arguments for the new vertebral theory of the skull.

(FIGURE 2.342. Skeleton of the hand or fore foot of six mammals. I man, II dog, III pig, IV ox, V tapir, VI horse. r radius, u ulna, a scaphoideum, b lunare, a triquetrum, d trapezium, e trapezoid, f capitatum, g hamatum, p pisiforme. 1 thumb, 2 index finger, 3 middle finger, 4 ring finger, 5 little finger. (From Gegenbaur.))

We have not space here to go into the details of Gegenbaur's theory of the skull. I must be content to refer the reader to the great work I have mentioned, in which it is thoroughly established from the empirico-philosophical point of view. He has also given a comprehensive and up-to-date treatment of the subject in his Comparative Anatomy of the Vertebrates (1898). Gegenbaur indicates as original "cranial ribs," or "lower arches of the cranial vertebrae," at each side of the head of the Selachii (Figure 2.334), the following pairs of arches: I and II, two lip-cartilages, the anterior (a) of which is composed of an upper piece only, the posterior (bc) from an upper and lower piece; III, the maxillary arches, also consisting of two pieces on each side--the primitive upper jaw (os palato-quadratum, o) and the primitive lower jaw (u); IV, the hyaloid bone (II); finally, V to X, six branchial arches in the narrower sense (III to VIII). From the anatomic features of these nine to ten cranial ribs or "lower vertebral arches" and the cranial nerves that spread over them, it is clear that the apparently simple cartilaginous primitive skull of the Selachii was originally formed from so many (at least nine) somites or provertebrae. The blending of these primitive segments into a single capsule is, however, so ancient that, in virtue of the law of curtailed heredity, the original division seems to have disappeared; in the embryonic development it is very difficult to detect it in isolated traces, and in some respects quite impossible. It is claimed that several (three to six) traces of provertebrae have been discovered in the anterior (pre-chordal) part of the Selachii-skull; this would bring up the number of cranial somites to twelve or sixteen, or even more.

(FIGURES 2.343 TO 2.345. Arm and hand of three anthropoids.

FIGURE 2.343. Chimpanzee (Anthropithecus niger).

FIGURE 2.344. Veddah of Ceylon (Homo veddalis).

FIGURE 2.345. European (Homo mediterraneus). (From Paul and Fritz Sarasin.))

In the primitive skull of man (Figure 2.323) and the higher Vertebrates, which has been evolved from that of the Selachii, five consecutive sections are discoverable at a certain early period of development, and one might be induced to trace these to five primitive vertebrae; but these sections are due entirely to adaptation to the five primitive cerebral vesicles, and correspond, like these, to a large number of metamera. That we have in the primitive skull of the mammals a greatly modified and transformed organ, and not at all a primitive formation, is clear from the circumstance that its original soft membranous form only assumes the cartilaginous character for the most part at the base and the sides, and remains membranous at the roof. At this part the bones of the subsequent osseous skull develop as external coverings over the membranous structure, without an intermediate cartilaginous stage, as there is at the base of the skull. Thus a large part of the cranial bones develop originally as covering bones from the corium, and only secondarily come into close touch with the primitive skull (Figure 2.333). We have previously seen how this very rudimentary beginning of the skull in man is formed ontogenetically from the "head-plates," and thus the fore end of the chorda is enclosed in the base of the skull. (Cf. Figs 1.145 and

Chapters

1.13 and 1.14.)

The phylogeny of the skull has made great progress during the last three decades through the joint attainments of comparative anatomy, ontogeny, and paleontology. By the judicious and comprehensive application of the phylogenetic method (in the sense of Gegenbaur) we have found the key to the great and important problems that arise from the thorough comparative study of the skull. Another school of research, the school of what is called "exact craniology" (in the sense of Virchow), has, meantime, made fruitless efforts to obtain this result. We may gratefully acknowledge all that this descriptive school has done in the way of accurately describing the various forms and measurements of the human skull, as compared with those of other mammals. But the vast empirical material that it has accumulated in its extensive literature is mere dead and sterile erudition until it is vivified and illumined by phylogenetic speculation.

Virchow confined himself to the most careful analysis of large numbers of human skulls and those of anthropoid mammals. He saw only the differences between them, and sought to express these in figures.

Without adducing a single solid reason, or offering any alternative explanation, he rejected evolution as an unproved hypothesis. He played a most unfortunate part in the controversy as to the significance of the fossil human skulls of Spy and Neanderthal, and the comparison of them with the skull of the Pithecanthropus (Figure 2.283). All the interesting features of these skulls that clearly indicated the transition from the anthropoid to the man were declared by Virchow to be chance pathological variations. He said that the roof of the skull of Pithecanthropus (Figure 2.335, 3) must have belonged to an ape, because so pronounced an orbital stricture (the horizontal constriction between the outer edge of the eye-orbit and the temples) is not found in any human being. Immediately afterwards Nehring showed in the skull of a Brazilian Indian (Figure 2.335, 2), found in the Sambaquis of Santos, that this stricture can be even deeper in man than in many of the apes. It is very instructive in this connection to compare the roofs of the skulls (seen from above) of different primates. I have, therefore, arranged nine such skulls in Figure 2.335, and reduced them to a common size.

(FIGURE 2.346. Transverse section of a fish's tail (from the tunny). (From Johannes Muller.) a upper (dorsal) lateral muscles, a apostrophe, b apostrophe lower (ventral) lateral muscles, d vertebral bodies, b sections of incomplete conical mantle, B attachment lines of the inter-muscular ligaments (from the side).)

We turn now to the branchial arches, which were regarded even by the earlier natural philosophers as "head-ribs." (Cf. Figures 1.167 to 1.170). Of the four original gill-arches of the mammals the first lies between the primitive mouth and the first gill-cleft. From the base of this arch is formed the upper-jaw process, which joins with the inner and outer nasal processes on each side, in the manner we have previously explained, and forms the chief parts of the skeleton of the upper jaw (palate bone, pterygoid