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- The Evolution of Man, V.2 - 40/63 -


was formed by gradual degeneration and differentiation from the many-toed fish-fin (Figure 2.341). The reduction of the radii to four was accompanied by a further differentiation of the fin-rod, its transverse segmentation into upper and lower halves, and the formation of the zone of the limb, which is composed originally of three limbs before and behind in the higher Vertebrates. The simple arch of the original shoulder-zone divides on each side into an upper (dorsal) piece, the shoulder-blade (scapula), and a lower (ventral) piece; the anterior part of the latter forms the primitive clavicle (procoracoideum), and the posterior part the coracoideum. In the same way the simple arch of the pelvic zone breaks up into an upper (dorsal) piece, the iliac-bone (os ilium), and a lower (ventral) piece; the anterior part of the latter forms the pubic bone (os pubis), and the posterior the ischial bone (os ischii).

There is also a complete agreement between the fore and hind limb in the stem or shaft. The first section of the stem is supported by a single strong bone--the humerus in the fore, the femur in the hind limb. The second section contains two bones: in front the radius (r) and ulna (u), behind the tibia and fibula. (Cf. the skeletons in Figures 2.260, 2.265, 2.270, 2.278 to 2.282, and 2.348.) The succeeding numerous small bones of the wrist (carpus) and ankle (tarsus) are also similarly arranged in the fore and hind extremities, and so are the five bones of the middle-hand (metacarpus) and middle-foot (metatarsus). Finally, it is the same with the toes themselves, which have a similar characteristic composition from a series of bony pieces before and behind. We find a complete parallel in all the parts of the fore leg and the hind leg.

When we thus learn from comparative anatomy that the skeleton of the human limbs is composed of just the same bones, put together in the same way, as the skeleton in the four higher classes of Vertebrates, we may at once infer a common descent of them from a single stem-form. This stem-form was the earliest amphibian that had five toes on each foot. It is particularly the outer parts of the limbs that have been modified by adaptation to different conditions. We need only recall the immense variations they offer within the mammal class. We have the slender legs of the deer and the strong springing legs of the kangaroo, the climbing feet of the sloth and the digging feet of the mole, the fins of the whale and the wings of the bat. It will readily be granted that these organs of locomotion differ as much in regard to size, shape, and special function as can be conceived. Nevertheless, the bony skeleton is substantially the same in every case. In the different limbs we always find the same characteristic bones in essentially the same rigidly hereditary connection; this is as splendid a proof of the theory of evolution as comparative anatomy can discover in any organ of the body. It is true that the skeleton of the limbs of the various mammals undergoes many distortions and degenerations besides the special adaptations (Figure 2.342). Thus we find the first finger or the thumb atrophied in the fore-foot (or hand) of the dog (II). It has entirely disappeared in the pig (III) and tapir (V). In the ruminants (such as the ox, IV) the second and fifth toes are also atrophied, and only the third and fourth are well developed (VI, 3). Nevertheless, all these different fore-feet, as well as the hand of the ape (Figure 2.340) and of man (Figure 2.341), were originally developed from a common pentadactyle stem-form. This is proved by the rudiments of the degenerated toes, and by the similarity of the arrangement of the wrist-bones in all the pentanomes (Figure 2.342 a to p).

If we candidly compare the bony skeleton of the human arm and hand with that of the nearest anthropoid apes, we find an almost perfect identity. This is especially true of the chimpanzee. In regard to the proportions of the various parts, the lowest living races of men (the Veddahs of Ceylon, Figure 2.344) are midway between the chimpanzee (Figure 2.343) and the European (Figure 2.345). More considerable are the differences in structure and the proportions of the various parts between the different genera of anthropoid apes (Figures 2.278 to 2.282); and still greater is the morphological distance between these and the lowest apes (the Cynopitheca). Here, again, impartial and thorough anatomic comparison confirms the accuracy of Huxley's pithecometra principle (Chapter 1.15).

The complete unity of structure which is thus revealed by the comparative anatomy of the limbs is fully confirmed by their embryology. However different the extremities of the four-footed Craniotes may be in their adult state, they all develop from the same rudimentary structure. In every case the first trace of the limb in the embryo is a very simple protuberance that grows out of the side of the hyposoma. These simple structures develop directly into fins in the fishes and Dipneusts by differentiation of their cells. In the higher classes of Vertebrates each of the four takes the shape in its further growth of a leaf with a stalk, the inner half becoming narrower and thicker and the outer half broader and thinner. The inner half (the stalk of the leaf) then divides into two sections--the upper and lower parts of the limb. Afterwards four shallow indentations are formed at the free edge of the leaf, and gradually deepen; these are the intervals between the five toes (Figure 1.174). The toes soon make their appearance. But at first all five toes, both of fore and hind feet, are connected by a thin membrane like a swimming-web; they remind us of the original shaping of the foot as a paddling fin. The further development of the limbs from this rudimentary structure takes place in the same way in all the Vertebrates according to the laws of heredity.

The embryonic development of the muscles, or ACTIVE organs of locomotion, is not less interesting than that of the skeleton, or PASSIVE organs. But the comparative anatomy and ontogeny of the muscular system are much more difficult and inaccessible, and consequently have hitherto been less studied. We can therefore only draw some general phylogenetic conclusions therefrom.

It is incontestable that the musculature of the Vertebrates has been evolved from that of lower Invertebrates; and among these we have to consider especially the unarticulated Vermalia. They have a simple cutaneous muscular layer, developing from the mesoderm. This was afterwards replaced by a pair of internal lateral muscles, that developed from the middle wall of the coelom-pouches; we still find the first rudiments of the muscles arising from the muscle-plate of these in the embryos of all the Vertebrates (cf. Figures 1.124, 1.158 to 1.160, 2.222 to 2.224 mp). In the unarticulated stem-forms of the Chordonia, which we have called the Prochordonia, the two coelom-pouches, and therefore also the muscle-plates of their walls, were not yet segmented. A great advance was made in the articulation of them, as we have followed it step by step in the Amphioxus (Figures 1.124 and 1.158). This segmentation of the muscles was the momentous historical process with which vertebration, and the development of the vertebrate stem, began. The articulation of the skeleton came after this segmentation of the muscular system, and the two entered into very close correlation.

The episomites or dorsal coelom-pouches of the Acrania, Cyclostomes, and Selachii (Figure 1.161 h) first develop from their inner or median wall (from the cell-layer that lies directly on the skeletal plate [sk] and the medullary tube [nr]) a strong muscle-plate (mp). By dorsal growth (w) it also reaches the external wall of the coelom-pouches, and proceeds from the dorsal to the ventral wall. From these segmental muscle-plates, which are chiefly concerned in the segmentation of the Vertebrates, proceed the lateral muscles of the stem, as we find in the simplest form in the Amphioxus (Figure 2.210). By the formation of a horizontal frontal septum they divide on each side into an upper and lower series of myotomes, dorsal and ventral lateral muscles. This is seen with typical regularity in the transverse section of the tail of a fish (Figure 2.346). From these earlier lateral muscles of the trunk develop the greater part of the subsequent muscles of the trunk, and also the much later "muscular buds" of the limbs.* (* The ontogeny of the muscles is mostly cenogenetic. The greater part of the muscles of the head (or the visceral muscles) belong originally to the hyposoma of the vertebrate organism, and develop from the wall of the hyposomites or ventral coelom-pouches. This also applies originally to the primary muscles of the limbs, as these too belong phylogenetically to the hyposoma. (Cf. Chapter 1.14))

CHAPTER 2.27. THE EVOLUTION OF THE ALIMENTARY SYSTEM.

The chief of the vegetal organs of the human frame, to the evolution of which we now turn our attention, is the alimentary canal. The gut is the oldest of all the organs of the metazoic body, and it leads us back to the earliest age of the formation of organs--to the first section of the Laurentian period. As we have already seen, the result of the first division of labour among the homogeneous cells of the earliest multicellular animal body was the formation of an alimentary cavity. The first duty and first need of every organism is self-preservation. This is met by the functions of the nutrition and the covering of the body. When, therefore, in the primitive globular Blastaea the homogeneous cells began to effect a division of labour, they had first to meet this twofold need. One half were converted into alimentary cells and enclosed a digestive cavity, the gut. The other half became covering cells, and formed an envelope round the alimentary tube and the whole body. Thus arose the primary germinal layers--the inner, alimentary, or vegetal layer, and the outer, covering, or animal layer. (Cf. Chapter 2.19.)

When we try to construct an animal frame of the simplest conceivable type, that has some such primitive alimentary canal and the two primary layers constituting its wall, we inevitably come to the very remarkable embryonic form of the gastrula, which we have found with extraordinary persistence throughout the whole range of animals, with the exception of the unicellulars--in the Sponges, Cnidaria, Platodes, Vermalia, Molluscs, Articulates, Echinoderms, Tunicates, and Vertebrates. In all these stems the gastrula recurs in the same very simple form. It is certainly a remarkable fact that the gastrula is found in various animals as a larva-stage in their individual development, and that this gastrula, though much disguised by cenogenetic modifications, has everywhere essentially the same palingenetic structure (Figures 1.30 to 1.35). The elaborate alimentary canal of the higher animals develops ontogenetically from the same simple primitive gut of the gastrula.

This gastraea theory is now accepted by nearly all zoologists. It was first supported and partly modified by Professor Ray-Lankester; he proposed three years afterwards (in his essay on the development of the Molluscs, 1875) to give the name of archenteron to the primitive gut and blastoporus to the primitive mouth.

Before we follow the development of the human alimentary canal in detail, it is necessary to say a word about the general features of its composition in the fully-developed man. The mature alimentary canal in man is constructed in all its main features like that of all the higher mammals, and particularly resembles that of the Catarrhines, the narrow-nosed apes of the Old World. The entrance into it, the mouth, is armed with thirty-two teeth, fixed in rows in the upper and lower jaws. As we have seen, our dentition is exactly the same as that of the Catarrhines, and differs from that of all other animals (Chapter 2.23). Above the mouth-cavity is the double nasal cavity; they are separated by the palate-wall. But we saw that this separation is not there from the first, and that originally there is a common mouth-nasal cavity in the embryo; and this is only divided


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