The Evolution of Man, V.2 by Ernst Haeckel (comprehension books .TXT) 📕

Chapter 2.

25.) The remainder of the first branchial arch, which is now called, by way of contrast, the "upper-jaw process," forms from its base two of the ear-ossicles (hammer and anvil), and as to the rest is converted into a long strip of cartilage that is known, after its discoverer, as "Meckel's cartilage," or the promandibula. At the outer surface of the latter is formed from the cellular matter of the corium, as covering or accessory bone, the permanent bony lower jaw. From the first part or base of the second branchial arch we get, in the mammals, the third ossicle of the ear, the stirrup; and from the succeeding parts we get (in this order) the muscle of the stirrup, the styloid process of the temporal bone, the styloid-hyoid ligament, and the little horn of the hyoid bone. The third branchial arch is only cartilaginous at the foremost part, and here the body of the hyoid bone and its larger horn are formed at each side by the junction of its two halves. The fourth branchial arch is only found transitorily in the mammal embryo as a rudimentary organ, and does not develop special parts; and there is no trace in the embryo of the higher Vertebrates of the posterior branchial arches (fifth and sixth pair), which are permanent in the Selachii. They have been lost long ago. Moreover, the four gill-clefts of the human embryo are only interesting as rudimentary organs, and they soon close up and disappear. The first alone (between the first and second branchial arches) has any permanent significance; from it are developed the tympanic cavity and the Eustachian tube. (Cf. Figures 1.169 and 2.320.)

It was Carl Gegenbaur again who solved the difficult problem of tracing the skeleton of the limbs of the Vertebrates to a common type. Few parts of the vertebrate body have undergone such infinitely varied modifications in regard to size, shape, and adaptation of structure as the limbs or extremities; yet we are in a position to reduce them all to the same hereditary standard. We may generally distinguish three groups among the Vertebrates in relation to the formation of their limbs. The lowest and earliest Vertebrates, the Acrania and Cyclostomes, had, like their invertebrate ancestors, no pairs of limbs, as we see in the Amphioxus and the Cyclostomes to-day (Figures 2.210 and 2.247). The second group is formed of the two classes of the true fishes and the Dipneusts; here there are always two pairs of limbs at first, in the shape of many-toed fins--one pair of breast-fins or fore legs, and one pair of belly-fins or hind legs (Figures 2.248 to 2.259). The third group comprises the four higher classes of Vertebrates--the amphibia, reptiles, birds, and mammals; in these quadrupeds there are at first the same two pairs of limbs, but in the shape of five-toed feet. Frequently we find less than five toes, and sometimes the feet are wholly atrophied (as in the serpents). But the original stem-form of the group had five toes or fingers before and behind (Figures 2.263 to 2.265).

The true primitive form of the pairs of limbs, such as they were found in the primitive fishes of the Silurian period, is preserved for us in the Australian dipneust, the remarkable Ceratodus (Figure 2.257). Both the breast-fin and the belly-fin are flat oval paddles, in which we find a biserial cartilaginous skeleton (Figure 2.336). This consists, firstly, of a much segmented fin-rod or "stem" (A, B), which runs through the fin from base to tip; and secondly of a double row of thin articulated fin-radii (r, r), which are attached to both sides of the fin-rod, like the feathers of a feathered leaf. This primitive fin, which Gegenbaur first recognised, is attached to the vertebral column by a simple zone in the shape of a cartilaginous arch. It has probably originated from the branchial arches.* (* While Gegenbaur derives the fins from two pairs of posterior separated branchial arches, Balfour holds that they have been developed from segments of a pair of originally continuous lateral fins or folds of the skin.)

We find the same biserial primitive fin more or less preserved in the fossilised remains of the earliest Selachii (Figure 2.248), Ganoids (Figure 2.253), and Dipneusts (Figure 2.256). It is also found in modified form in some of the actual sharks and pikes. But in the majority of the Selachii it has already degenerated to the extent that the radii on one side of the fin-rod have been partly or entirely lost, and are retained only on the other (Figure 2.337). We thus get the uniserial fin, which has been transmitted from the Selachii to the rest of the fishes (Figure 2.338).

(FIGURE 2.347. Human skeleton. (Cf. Figure 2.326.)

FIGURE 2.348. Skeleton of the giant gorilla. (Cf. Figure 1.209.))

Gegenbaur has shown how the five-toed leg of the Amphibia, that has been inherited by the three classes of Amniotes, was evolved from the uniserial fish-fin.* (* The limb of the four higher classes of Vertebrates is now explained in the sense that the original fin-rod passes along its outer (ulnar or fibular) side, and ends in the fifth toe. It was formerly believed to go along the inner (radial or tibial) side, and end in the first toe, as Figure 2.339 shows.) In the dipneust ancestors of the Amphibia the radii gradually atrophy, and are lost, for the most part, on the other side of the fin-rod as well (the lighter cartilages in Figure 2.338). Only the four lowest radii (shaded in the illustration) are preserved; and these are the four inner toes of the foot (first to fourth). The little or fifth toe is developed from the lower end of the fin-rod. From the middle and upper part of the fin-rod was developed the long stem of the limb--the important radius and ulna (Figure 2.339 r and u) and humerus (h) of the higher Vertebrates.

In this way the five-toed foot of the Amphibia, which we first meet in the Carboniferous Stegocephala (Figure 2.260), and which was inherited from them by the reptiles on one side and the mammals on the other, 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.