The Evolution of Man, vol 2, Ernst Haeckel [online e reader txt] 📗

vesicle, filled with fluid, and containing otoliths, with the acoustic nerve expanding on its wall. But, while the auditory vesicle is usually of a simple round or oval shape in the Invertebrates, it has in the Vertebrates a special and curious structure, the labyrinth. This thin-membraned labyrinth is enclosed in a bony capsule of the same shape, the osseous labyrinth (Figure 2.321), and this lies in the middle of the petrous bone of the skull. The labyrinth is divided into two vesicles in all the Gnathostomes. The larger one is called the utriculus, and has three arched appendages, called the “semi-circular canals” (c, d, e). The smaller vesicle is called the sacculus, and is connected with a peculiar appendage, with (in man and the higher mammals) a spiral form something like a snail’s shell, and therefore called the cochlea (= snail, b). On the thin wall of this delicate labyrinth the acoustic nerve, which comes from the after-brain, spreads out in most elaborate fashion. It divides into two main branches—a cochlear nerve (for the cochlea) and a vestibular nerve (for the rest of the labyrinth). The former seems to have more to do with the quality, the latter with the quantity, of the acoustic sensations. Through the cochlear nerves we learn the height and timbre, through the vestibular nerves the intensity, of tones.

(FIGURE 2.321. The bony labyrinth of the human ear (left side). a vestibulum, b cochlea, c upper canal, d posterior canal, e outer canal, f oval fenestra, g round fenestra. (From Meyer.)

FIGURE 2.322. Development of the auscultory labyrinth of the chick, in five successive stages (A to E). (Vertical transverse sections of the skull.) fl auscultory pits, lv auscultory vesicles, lr labyrinthic appendage, c rudimentary cochlea, csp posterior canal, cse external canal, jv jugular vein. (From Reissner.))

The first structure of this highly elaborate organ is very simple in the embryo of man and all the other Craniotes; it is a pit-like depression in the skin. At the back part of the head at both sides, near the after brain, a small thickening of the horny plate is formed at the upper end of the second gill-cleft (Figure 2.322 A fl). This sinks into a sort of pit, and severs from the epidermis, just as the lens of the eye does. In this way is formed at each side, directly under the horny plate of the back part of the head, a small vesicle filled with fluid, the primitive auscultory vesicle, or the primary labyrinth. As it separates from its source, the horny plate, and presses inwards and backwards into the skull, it changes from round to pear-shaped (Figures 2.322 B lv and 2.323 o). The outer part of it is lengthened into a thin stem, which at first still opens outwards by a narrow canal. This is the labyrinthic appendage (Figure 2.322 lr). In the lower Vertebrates it develops into a special cavity filled with calcareous crystals, which remains open permanently in some of the primitive fishes, and opens outwards in the upper part of the skull. But in the mammals the labyrinthic appendage degenerates. In these it has only a phylogenetic interest as a rudimentary organ, with no actual physiological significance. The useless relic of it passes through the wall of the petrous bone in the shape of a narrow canal, and is called the vestibular aqueduct.

It is only the inner and lower bulbous part of the separated auscultory vesicle that develops into the highly complex and differentiated structure that is afterwards known as the secondary labyrinth. This vesicle divides at an early stage into an upper and larger and a lower and smaller section. From the one we get the utriculus with the semi-circular canals; from the other the sacculus and the cochlea (Figure 2.320 c). The canals are formed in the shape of simple pouch-like involutions of the utricle (cse and csp). The edges join together in the middle part of each fold, and separate from the utricle, the two ends remaining in open connection with its cavity. All the Gnathostomes have these three canals like man, whereas among the Cyclostomes the lampreys have only two and the hag-fishes only one. The very complex structure of the cochlea, one of the most elaborate and wonderful outcomes of adaptation in the mammal body, develops originally in very simple fashion as a flask-like projection from the sacculus. As Hasse and Retzius have pointed out, we find the successive ontogenetic stages of its growth represented permanently in the series of the higher Vertebrates. The cochlea is wanting even in the Monotremes, and is restricted to the rest of the mammals and man.

The auditory nerve, or eighth cerebral nerve, expands with one branch in the cochlea, and with the other in the remaining parts of the labyrinth. This nerve is, as Gegenbaur has shown, the sensory dorsal branch of a cerebro-spinal nerve, the motor ventral branch of which acts for the muscles of the face (nervus facialis). It has therefore originated phylogenetically from an ordinary cutaneous nerve, and so is of quite different origin from the optic and olfactory nerves, which both represent direct outgrowths of the brain. In this respect the auscultory organ is essentially different from the organs of sight and smell. 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