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

The acoustic nerve is formed from ectodermic cells of the hind brain, and develops from the nervous structure that appears at its dorsal limit. On the other hand, all the membranous, cartilaginous, and osseous coverings of the labyrinth are formed from the mesodermic head-plates.

(FIGURE 2.323. Primitive skull of the human embryo, four weeks old, vertical section, left half seen internally. v, z, m, h, n the five pits of the cranial cavity, in which the five cerebral vesicles lie (fore, intermediate, middle, hind, and after brains), o pear-shaped primary auscultory vesicle (appearing through), a eye (appearing through), no optic nerve, p canal of the hypophysis, t central prominence of the skull. (From Kolliker.))

The apparatus for conducting sound which we find in the external and middle ear of mammals develops quite separately from the apparatus for the sensation of sound. It is both phylogenetically and ontogenetically an independent secondary formation, a later accession to the primary internal ear. Nevertheless, its development is not less interesting, and is explained with the same ease by comparative anatomy. In all the fishes and in the lowest Vertebrates there is no special apparatus for conducting sound, no external or middle ear; they have only a labyrinth, an internal ear, which lies within the skull. They are without the tympanum and tympanic cavity, and all its appendages. From many observations made in the last few decades it seems that many of the fishes (if not all) cannot distinguish tones; their labyrinth seems to be chiefly (if not exclusively) an organ for the sense of space (or equilibrium). If it is destroyed, the fishes lose their balance and fall. In the opinion of recent physiologists this applies also to many of the Invertebrates (including the nearer ancestors of the Vertebrates). The round vesicles which are considered to be their auscultory vesicles, and which contain an otolith, are supposed to be merely organs of the sense of space ("static vesicles or statocysts").

The middle ear makes its first appearance in the amphibian class, where we find a tympanum, tympanic cavity, and Eustachian tube; these animals, and all terrestrial Vertebrates, certainly have the faculty of hearing. All these essential parts of the middle ear originate from the first gill-cleft and its surrounding part; in the Selachii this remains throughout life an open squirting-hole, and lies between the first and second gill-arch. In the embryo of the higher Vertebrates it closes up in the centre, and thus forms the tympanic membrane. The outlying remainder of the first gill-cleft is the rudiment of the external meatus. From its inner part we get the tympanic cavity, and, further inward still, the Eustachian tube. Connected with this is the development of the three bones of the mammal ear from the first two gill-arches; the hammer and anvil are formed from the first, the stirrup from the upper end of the second, gill-arch.

(FIGURE 2.324. The rudimentary muscles of the ear in the human skull. a raising muscle (M. attollens), b drawing muscle (M. attrahens), c withdrawing muscle (M. retrahens), d large muscle of the helix (M. helicis major), e small muscle of the helix (M. helicis minor), f muscle of the angle of the ear (M. tragicus), g anti-angular muscle (M. antitragicus). (From H. Meyer.))

Finally, the shell (pinna or concha) and external meatus (passage to the tympanum) of the outer ear are developed in a very simple fashion from the skin that borders the external aperture of the first gill-cleft. The shell rises in the shape of a circular fold of the skin, in which cartilage and muscles are afterwards formed (Figures 2.313 and 2.315). This organ is only found in the mammalian class. It is very rudimentary in the lowest section, the Monotremes. In the others it is found at very different stages of development, and sometimes of degeneration. It is degenerate in most of the aquatic mammals. The majority of them have lost it altogether--for instance, the walruses and whales and most of the seals. On the other hand, the pinna is well developed in the great majority of the Marsupials and Placentals; it receives and collects the waves of sound, and is equipped with a very elaborate muscular apparatus, by means of which the pinna can be turned freely in any direction and its shape be altered. It is well known how readily domestic animals--horses, cows, dogs, hares, etc.--point their ears and move them in different directions. Most of the apes do the same, and our earlier ape ancestors were also able to do it. But our later simian ancestors, which we have in common with the anthropoid apes, abandoned the use of these muscles, and they gradually became rudimentary and useless. However, we possess them still (Figure 2.324). In fact, some men can still move their ears a little backward and forward by means of the drawing and withdrawing muscles (b and c); with practice this faculty can be much improved. But no man can now lift up his ears by the raising muscle (a), or change the shape of them by the small inner muscles (d, e, f, g). These muscles were very useful to our ancestors, but are of no consequence to us. This applies to most of the anthropoid apes as well.

We also share with the higher anthropoid apes (gorilla, chimpanzee, and orang) the characteristic form of the human outer ear, especially the folded border, the helix and the lobe. The lower apes have pointed ears, without folded border or lobe, like the other mammals. But Darwin has shown that at the upper part of the folded border there is in many men a small pointed process, which most of us do not possess. In some individuals this process is well developed. It can only be explained as the relic of the original point of the ear, which has been turned inwards in consequence of the curving of the edge. If we compare the pinna of man and the various apes in this respect, we find that they present a connected series of degenerate structures. In the common catarrhine ancestors of the anthropoids and man the degeneration set in with the folding together of the pinna. This brought about the helix of the ear, in which we find the significant angle which represents the relic of the salient point of the ear in our earlier simian ancestors. Here again, therefore, comparative anatomy enables us to trace with certainty the human ear to the similar, but more developed, organ of the lower mammals. At the same time, comparative physiology shows that it was a more or less useful implement in the latter, but it is quite useless in the anthropoids and man. The conducting of the sound has scarcely been affected by the loss of the pinna. We have also in this the explanation of the extraordinary variety in the shape and size of the shell of the ear in different men; in this it resembles other rudimentary organs.

CHAPTER XI(26. EVOLUTION OF THE ORGANS OF MOVEMENT.)

 

The peculiar structure of the locomotive apparatus is one of the features that are most distinctive of the vertebrate stem. The chief part of this apparatus is formed, as in all the higher animals, by the active organs of movement, the muscles; in consequence of their contractility they have the power to draw up and shorten themselves. This effects the movement of the various parts of the body, and thus the whole body is conveyed from place to place. But the arrangement of these muscles and their relation to the solid skeleton are different in the Vertebrates from the Invertebrates.

(FIGURE 2.325. The human skeleton. From the right.

FIGURE 2.326. The human skeleton. Front.)

In most of the lower animals, especially the Platodes and Vermalia, we find that the muscles form a simple, thin layer of flesh immediately underneath the skin. This muscular layer is very closely connected with the skin itself; it is the same in the Mollusc stem. Even in the large division of the Articulates, the classes of crabs, spiders, myriapods, and insects, we find a similar feature, with the difference that in this case the skin forms a solid armour--a rigid cutaneous skeleton made of chitine (and often also of carbonate of lime). This external chitine coat undergoes a very elaborate articulation both on the trunk and the limbs of the Articulates, and in consequence the muscular system also, the contractile fibres of which are attached inside the chitine tubes, is highly articulated. The Vertebrates form a direct contrast to this. In these alone a solid internal skeleton is developed, of cartilage or bone, to which the muscles are attached. This bony skeleton is a complex lever apparatus, or PASSIVE apparatus of movement. Its rigid parts, the arms of the levers, or the bones, are brought together by the actively mobile muscles, as if by drawing-ropes. This admirable locomotorium, especially its solid central axis, the vertebral column, is a special feature of the Vertebrates, and has given the name to the group.

(FIGURE 2.327. The human vertebral column (standing upright, from the right side). (From H. Meyer.))

In order to get a clear idea of the chief features of the development of the human skeleton, we must first examine its composition in the adult frame (Figure 2.325, the human skeleton seen from the right; Figure 2.326, front view of the whole skeleton). As in other mammals, we distinguish first between the axial or dorsal skeleton and the skeleton of the limbs. The axial skeleton consists of the vertebral column (the skeleton of the trunk) and the skull (skeleton of the head); the latter is a peculiarly modified part of the former. As appendages of the vertebral column we have the ribs, and of the skull we have the hyoid bone, the lower jaw, and the other products of the gill-arches.

The skeleton of the limbs or extremities is composed of two groups of parts--the skeleton of the extremities proper and the zone-skeleton, which connects these with the vertebral column. The zone-skeleton of the arms (or fore legs) is the shoulder-zone; the zone-skeleton of the legs (or hind legs) is the pelvic zone.

(FIGURE 2.328. A piece of the axial rod (chorda dorsalis), from a sheep embryo. a cuticular sheath, b cells. (From Kolliker.))

The vertebral column (Figure 2.327) in man is composed of thirty-three to thirty-five ring-shaped bones in a continuous series (above each other, in man's upright position). These vertebrae are separated from each other by elastic ligaments, and at the same time connected by joints, so that the whole column forms a firm and solid, but flexible and elastic, axial skeleton, moving freely in all directions. The vertebrae differ in shape and connection at the various parts of the trunk, and we distinguish the following groups in the series, beginning at the top: Seven cervical vertebrae, twelve dorsal vertebrae, five lumbar vertebrae, five sacral vertebrae, and four to six caudal vertebrae. The uppermost, or those next to the skull, are the cervical vertebrae (Figure 2.327); they have a hole in each of the lateral processes. There are seven of these vertebrae in man and almost all the other mammals, even if the neck is as long as that of the camel or giraffe, or as short as that of the mole