The Evolution of Man Volume Ii Part 12

(FIGURE 2.293. The human brain, seen from the left. (From H. Meyer.) The furrows of the cerebrum are indicated by thick, and those of the cerebellum by finer lines. Under the latter we can see the medulla oblongata. f1 to f2 frontal convolutions, C central convolutions, S fissure of Sylvius, T temporal furrow, Pa parietal lobes, An angular gyrus, Po parieto-occipital fissure.)

When we examine the embryology of the human nervous system, we must start from the important fact, which we have already seen, that the first structure of it in man and all the higher Vertebrates is the simple medullary tube, and that this separates from the outer germinal layer in the middle line of the sole-shaped embryonic shield. As the reader will remember, the straight medullary furrow first appears in the middle of the sandal-shaped embryonic shield. At each side of it the parallel borders curve over in the form of dorsal or medullary swellings. These bend together with their free borders, and thus form the closed medullary tube (Figures 1.133 to 1.137). At first this tube lies directly underneath the h.o.r.n.y plate; but it afterwards travels inwards, the upper edges of the provertebral plates growing together between the h.o.r.n.y plate and the tube, joining above the latter, and forming a completely closed ca.n.a.l. As Gegenbaur very properly observes, "this gradual imbedding in the inner part of the body is a process acquired with the progressive differentiation and the higher potentiality that this secures; by this process the organ of greater value to the organism is buried within the frame." (Cf. Figures 1.143 to 1.146).

(FIGURES 2.294 TO 2.296. Central marrow of the human embryo from the seventh week, 4/5 inch long. (From Kolliker.)

FIGURE 2.294. The brain from above, v fore brain, z intermediate brain, m middle brain, h hind brain, n after brain.

FIGURE 2.295. The brain with the uppermost part of the cord, from the left.

FIGURE 2.296. Back view of the whole embryo: brain and spinal cord exposed.)

In the Cyclostoma--a stage above the Acrania--the fore end of the cylindrical medullary tube begins early to expand into a pear-shaped vesicle; this is the first outline of an independent brain. In this way the central marrow of the Vertebrates divides clearly into its two chief sections, brain and spinal cord. The simple vesicular form of the brain, which persists for some time in the Cyclostoma, is found also at first in all the higher Vertebrates (Figure 1.153 hb). But in these it soon pa.s.ses away, the one vesicle being divided into several successive parts by transverse constrictions. There are first two of these constrictions, dividing the brain into three consecutive vesicles (fore brain, middle brain, and hind brain, Figure 1.154 v, m, h). Then the first and third are sub-divided by fresh constrictions, and thus we get five successive sections (Figure 1.155).

In all the Craniotes, from the Cyclostoma up to man, the same parts develop from these five original cerebral vesicles, though in very different ways. The first vesicle, the fore brain (Figure 1.155 v), forms by far the largest part of the cerebrum--namely, the large hemispheres, the olfactory lobes, the corpora striata, the callosum, and the fornix. From the second vesicle, the intermediate brain (z), originate especially the optic thalami, the other parts that surround the third cerebral ventricle, and the infundibulum and pineal gland.

The third vesicle, the middle brain (m), produces the corpora quadrigemina and the aqueduct of Sylvius. From the fourth vesicle, the hind brain (h), develops the greater part of the cerebellum--namely, the vermis and the two small hemispheres. Finally, the fifth vesicle, the after brain (n), forms the medulla oblongata, with the quadrangular pit (the floor of the fourth ventricle), the pyramids, olivary bodies, etc.

We must certainly regard it as a comparative-anatomical and ontogenetic fact of the greatest significance that in all the Craniotes, from the lowest Cyclostomes and fishes up to the apes and man, the brain develops in just the same way in the embryo. The first rudiment of it is always a simple vesicular enlargement of the fore end of the medullary tube. In every case, first three, then five, vesicles develop from this bulb, and the permanent brain with all its complex anatomic structures, of so great a variety in the various cla.s.ses of Vertebrates, is formed from the five primitive vesicles.

When we compare the mature brain of a fish, an amphibian, a reptile, a bird, and a mammal, it seems incredible that we can trace the various parts of these organs, that differ so much internally and externally, to common types. Yet all these different Craniote brains have started with the same rudimentary structure. To convince ourselves of this we have only to compare the corresponding stages of development of the embryos of these different animals.

(FIGURE 2.297. Head of a chick embryo (hatched fifty-eight hours), from the back, magnified forty times. (From Mihalkovics.) vw anterior wall of the fore brain. vh its ventricle. au optic vesicles, mh middle brain, kh hind brain, nh after brain, hz heart (seen from below), vw vitelline veins, us primitive segment, rm spinal cord.)

This comparison is extremely instructive. If we extend it through the whole series of the Craniotes, we soon discover this interesting fact: In the Cyclostomes (the Myxinoida and Petromyzonta), which we have recognised as the lowest and earliest Craniotes, the whole brain remains throughout life at a very low stage, which is very brief and pa.s.sing in the embryos of the higher Craniotes; they retain the five original sections of the brain unchanged. In the fishes we find an essential and considerable modification of the five vesicles; it is clearly the brain of the Selachii in the first place, and subsequently the brain of the Ganoids, from which the brain of the rest of the fishes on the one hand and of the Dipneusts and Amphibia, and through these of the higher Vertebrates, on the other hand, must be derived.

In the fishes and Amphibia (Figure 2.300) there is a preponderant development of the middle brain, and also the after brain, the first, second, and fourth sections remaining very primitive. It is just the reverse in the higher Vertebrates, in which the first and third sections, the cerebrum and cerebellum, are exceptionally developed; while the middle brain and after brain remain small. The corpora quadrigemina are mostly covered by the cerebrum, and the oblongata by the cerebellum. But we find a number of stages of development within the higher Vertebrates themselves. From the Amphibia upwards the brain (and with it the psychic life) develops in two different directions; one of these is followed by the reptiles and birds, and the other by the mammals. The development of the first section, the fore brain, is particularly characteristic of the mammals. It is only in them that the cerebrum becomes so large as to cover all the other parts of the brain (Figures 2.293 and 2.301 to 2.304).

There are also notable variations in the relative position of the cerebral vesicles. In the lower Craniotes they lie originally almost in the same plane. When we examine the brain laterally, we can cut through all five vesicles with a straight line. But in the Amniotes there is a considerable curve in the brain along with the bending of the head and neck; the whole of the upper dorsal surface of the brain develops much more than the under ventral surface. This causes a curve, so that the parts come to lie as follows: The fore brain is right in front and below, the intermediate brain a little higher, and the middle brain highest of all; the hind brain lies a little lower, and the after brain lower still. We find this only in the Amniotes--the reptiles, birds, and mammals.

(FIGURE 2.298. Brain of three craniote embryos in vertical section. A of a shark (Heptarchus), B of a serpent (Coluber), C of a goat (Capra). a fore brain, b intermediate brain, c middle brain, d hind brain, e after brain, s primitive cleft. (From Gegenbaur.)

FIGURE 2.299. Brain of a shark (Scyllium), back view. g fore-brain, h olfactory lobes, which send the large olfactory nerves to the nasal capsule (o), d intermediate brain, b middle brain; behind this the insignificant structure of the hind brain, a after brain. (From Gegenbaur.)

FIGURE 2.300. Brain and spinal cord of the frog. A from the dorsal, B from the ventral side. a olfactory lobes before the (b) fore brain, i infundibulum at the base of the intermediate brain, c middle brain, d hind brain, s quadrangular pit in the after brain, m spinal cord (very short in the frog), m apostrophe roots of the spinal nerves, t terminal fibres of the spinal cord. (From Gegenbaur.)

FIGURE 2.301. Brain of an ox-embryo, two inches in length. (From Mihalkovics, magnified three times.) Left view; the lateral wall of the left hemisphere has been removed, st corpora striata, ml Monro-foramen, ag arterial plexus, ah Ammon's horn, mh middle brain, kh cerebellum. dv roof of the fourth ventricle, bb pons Varolii, na medulla oblongata.)

Thus, while the brain of the mammals agrees a good deal in general growth with that of the birds and reptiles, there are some striking differences between the two. In the Sauropsids (birds and reptiles) the middle brain and the middle part of the hind brain are well developed. In the mammals these parts do not grow, and the fore-brain develops so much that it overlies the other vesicles. As it continues to grow towards the rear, it at last covers the whole of the rest of the brain, and also encloses the middle parts from the sides (Figures 2.301 to 2.303). This process is of great importance, because the fore brain is the organ of the higher psychic life, and in it those functions of the nerve-cells are discharged which we sum up in the word "soul." The highest achievements of the animal body--the wonderful manifestations of consciousness and the complex molecular processes of thought--have their seat in the fore brain. We can remove the large hemispheres, piece by piece, from the mammal without killing it, and we then see how the higher functions of consciousness, thought, will, and sensation, are gradually destroyed, and in the end completely extinguished. If the animal is fed artificially, it may be kept alive for a long time, as the destruction of the psychic organs by no means involves the extinction of the faculties of digestion, respiration, circulation, urination--in a word, the vegetative functions. It is only conscious sensation, voluntary movement, thought, and the combination of various higher psychic functions that are affected.

(FIGURE 2.302. Brain of a human embryo, twelve weeks old. (From Mihalkovics, natural size.) Seen from behind and above. ms mantle-furrow, mh corpora quadrigemina (middle brain), vs anterior medullary ala, kh cerebellum, vv fourth ventricle, na medulla oblongata.)

The fore brain, the organ of these functions, only attains this high level of development in the more advanced Placentals, and thus we have the simple explanation of the intellectual superiority of the higher mammals. The soul of most of the lower Placentals is not much above that of the reptiles, but among the higher Placentals we find an uninterrupted gradation of mental power up to the apes and man. In harmony with this we find an astonishing variation in the degree of development of their fore brain, not only qualitatively, but also quant.i.tatively. The ma.s.s and weight of the brain are much greater in modern mammals, and the differentiation of its various parts more important, than in their extinct Tertiary ancestors. This can be shown paleontologically in any particular order. The brains of the living ungulates are (relatively to the size of the body) four to six times (in the highest groups even eight times) as large as those of their earlier Tertiary ancestors, the well-preserved skulls of which enable us to determine the size and weight of the brain.

(FIGURE 2.303. Brain of a human embryo, twenty-four weeks old, halved in the median plane: right hemisphere seen from inside. (From Mihalkovics, natural size.) rn olfactory nerve. tr funnel of the intermediate brain, vc anterior commissure, ml Monro-foramen, gw fornix, ds transparent sheath, bl corpus callosum, br fissure at its border, hs occipital fissure, zh cuneus, sf occipital transverse fissure, zb pineal gland, mh corpora quadrigemina, kh cerebellum.

In the lower mammals the surface of the cerebral hemispheres is quite smooth and level, as in the rabbit (Figure 2.304). Moreover, the fore brain remains so small that it does not cover the middle brain. At a stage higher the middle brain is covered, but the hind brain remains free. Finally, in the apes and man, the latter also is covered by the fore brain. We can trace a similar gradual development in the fissures and convolutions that are found on the surface of the cerebrum of the higher mammals (Figures 2.292 and 2.293). If we compare different groups of mammals in regard to these fissures and convolutions, we find that their development proceeds step by step with the advance of mental life.

Of late years great attention has been paid to this special branch of cerebral anatomy, and very striking individual differences have been detected within the limits of the human race. In all human beings of special gifts and high intelligence the convolutions and fissures are much more developed than in the average man; and they are more developed in the latter than in idiots and others of low mental capacity. There is a similar gradation among the mammals in the internal structure of the fore brain. In particular the corpus callosum, that unites the two cerebral hemispheres, is only developed in the Placentals. Other structures--for instance, in the lateral ventricles--that seem at first to be peculiar to man, are also found in the higher apes, and these alone. It was long thought that man had certain distinctive organs in his cerebrum which were not found in any other animal. But careful examination has discovered that this is not the case, but that the characteristic features of the human brain are found in a rudimentary form in the lower apes, and are more or less fully developed in the higher apes. Huxley has convincingly shown, in his Man's Place in Nature (1863), that the differences in the formation of the brain within the ape-group const.i.tute a deeper gulf between the lower and higher apes than between the higher apes and man.

The comparative anatomy and physiology of the brain of the higher and lower mammals are very instructive, and give important information in connection with the chief questions of psychology.

(FIGURE 2.304. Brain of the rabbit. A from the dorsal, B from the ventral side, lo olfactory lobes, I fore brain, h hypophysis at the base of the intermediate brain, III middle brain, IV hind brain, V after brain, 2 optic nerve, 3 oculo-motor nerve, 5 to 8 cerebral nerves. In A the roof of the right hemisphere (I) is removed, so that we can see the corpora striata in the lateral ventricle. (From Gegenbaur.))

The central marrow (brain and spinal cord) develops from the medullary tube in man just as in all the other mammals, and the same applies to the conducting marrow or "peripheral nervous system." It consists of the SENSORY nerves, which conduct centripetally the impressions from the skin and the sense-organs to the central marrow, and of the MOTOR nerves, which convey centrifugally the movements of the will from the central marrow to the muscles. All these peripheral nerves grow out of the medullary tube (Figure 1.171), and are, like it, products of the skin-sense layer.

The complete agreement in the structure and development of the psychic organs which we find between man and the highest mammals, and which can only be explained by their common origin, is of profound importance in the monistic psychology. This is only seen in its full light when we compare these morphological facts with the corresponding physiological phenomena, and remember that every psychic action requires the complete and normal condition of the correlative brain structure for its full and normal exercise. The very complex molecular movements inside the neural cells, which we describe comprehensively as "the life of the soul," can no more exist in the vertebrate, and therefore in man, without their organs than the circulation without the heart and blood. And as the central marrow develops in man from the same medullary tube as that of the other vertebrates, and as man shares the characteristic structure of his cerebrum (the organ of thought) with the anthropoid apes, his psychic life also must have the same origin as theirs.

If we appreciate the full weight of these morphological and physiological facts, and put a proper phylogenetic interpretation on the observations of embryology, we see that the older idea of the personal immortality of the human soul is scientifically untenable.

Death puts an end, in man as in any other vertebrate, to the physiological function of the cerebral neurona, the countless microscopic ganglionic cells, the collective activity of which is known as "the soul." I have shown this fully in the eleventh chapter of my Riddle of the Universe.

CHAPTER 2.25. EVOLUTION OF THE SENSE-ORGANS.

The sense-organs are indubitably among the most important and interesting parts of the human body; they are the organs by means of which we obtain our knowledge of objects in the surrounding world.

Nihil est in intellectu quod non prius fuerit in sensu. They are the first sources of the life of the soul. There is no other part of the body in which we discover such elaborate anatomical structures, co-operating with a definite purpose; and there is no other organ in which the wonderful and purposive structure seems so clearly to compel us to admit a Creator and a preconceived plan. Hence we find special efforts made by dualists to draw our attention here to the "wisdom of the Creator" and the design visible in his works. As a matter of fact, you will discover, on mature reflection, that on this theory the Creator is at bottom only playing the part of a clever mechanic or watch-maker; all these familiar teleological ideas of Creator and creation are based, in the long run, on a similar childlike anthropomorphism.

However, we must grant that at the first glance the teleological theory seems to give the simplest and most satisfactory explanation of these purposive structures. If we merely examine the structure and functions of the most advanced sense-organs, it seems impossible to explain them without postulating a creative act. Yet evolution shows us quite clearly that this popular idea is totally wrong. With its a.s.sistance we discover that the purposive and remarkable sense-organs were developed, like all other organs, without any preconceived design--developed by the same mechanical process of natural selection, the same constant correlation of adaptation and heredity, by which the other purposive structures in the animal frame were slowly and gradually brought forth in the struggle for life.

Like most other Vertebrates, man has six sensory organs, which serve for eight different cla.s.ses of sensations. The skin serves for sensations of pressure and temperature. This is the oldest, lowest, and vaguest of the sense-organs; it is distributed over the surface of the body. The other sensory activities are localised. The s.e.xual sense is bound up with the skin of the external s.e.xual organs, the sense of taste with the mucous lining of the mouth (tongue and palate), and the sense of smell with the mucous lining of the nasal cavity. For the two most advanced and most highly differentiated sensory functions there are special and very elaborate mechanical structures--the eye for the sense of sight, and the ear for the sense of hearing and s.p.a.ce (equilibrium).

Comparative anatomy and physiology teach us that there are no differentiated sense-organs in the lower animals; all their sensations are received by the surface of the skin. The undifferentiated skin-layer or ectoderm of the Gastraea is the simple stratum of cells from which the differentiated sense-organs of all the Metazoa (including the Vertebrates) have been evolved. Starting from the a.s.sumption that necessarily only the superficial parts of the body, which are in direct touch with the outer world, could be concerned in the origin of sensations, we can see at once that the sense-organs also must have arisen there. This is really the case. The chief part of all the sense-organs originates from the skin-sense layer, partly directly from the h.o.r.n.y plate, partly from the brain, the foremost part, of the medullary tube, after it has separated from the h.o.r.n.y plate. If we compare the embryonic development of the various sense-organs, we see that they all make their appearance in the simplest conceivable form; the wonderful contrivances that make the higher sense-organs among the most remarkable and elaborate structures in the body develop only gradually. In the phylogenetic explanation of them comparative anatomy and ontogeny achieve their greatest triumphs.

But at first all the sense-organs are merely parts of the skin in which sensory nerves expand. These nerves themselves were originally of a h.o.m.ogeneous character. The different functions or specific energies of the differentiated sense-nerves were only gradually developed by division of labour. At the same time, their simple terminal expansions in the skin were converted into extremely complex organs.

The great instructiveness of these historical facts in connection with the life of the soul is not difficult to see. The whole philosophy of the future will be transformed as soon as psychology takes cognisance of these genetic phenomena and makes them the basis of its speculations. When we examine impartially the manuals of psychology that have been published by the most distinguished speculative philosophers and are still widely distributed, we are astonished at the naivete with which the authors raise their airy metaphysical speculations, regardless of the momentous embryological facts that completely refute them. Yet the science of evolution, in conjunction with the great advance of the comparative anatomy and physiology of the sense-organs, provides the one sound empirical basis of a natural psychology.

(FIGURE 2.305. Head of a shark (Scyllium), from the ventral side. m mouth, o olfactory pits, r nasal groove, n nasal fold in natural position, n apostrophe nasal fold drawn up. (The dots are openings of the mucous ca.n.a.ls.) (From Gegenbaur.))

In respect of the terminal expansions of the sensory nerves, we can distribute the human sense-organs in three groups, which correspond to three stages of development. The first group comprises those organs the nerves of which spread out quite simply in the free surface of the skin itself (organs of the sense of pressure, warmth, and s.e.x). In the second group the nerves spread out in the mucous coat of cavities which are at first depressions in or inv.a.g.i.n.ations of the skin (organs of the sense of smell and taste). The third group is formed of the very elaborate organs, the nerves of which spread out in an internal vesicle, separated from the skin (organs of the sense of sight, hearing, and s.p.a.ce).

(FIGURES 2.306 AND 2.307. Head of a chick embryo, three days old: 2.306 front view, 2.307 from the right. n rudimentary nose (olfactory pits), l rudimentary eyes (optic pits), g rudimentary ear (auscultory pit), v fore brain, gl eye-cleft, o process of upper jaw, u process of lower jaw of the first gill-arch.

FIGURE 2.308. Head of a chick embryo, four days old, from below. n nasal pit, o upper-jaw process of the first gill-arch, u lower-jaw process of same, k double apostrophe second gill-arch, sp choroid fissure of eye, s gullet.

FIGURES 2.309 AND 2.310. Heads of chick embryos: 2.309 from the end of the fourth, 2.310 from the beginning of the fifth week. Letters as in Figure 2.308, except: in inner, an outer, nasal process, nf nasal furrow, st frontal process, m mouth. (From Kolliker.) Figures 2.306 to 2.310 are magnified to the same extent.)

There is little to be said of the development of the lower sense-organs. We have already considered (Chapter 2.24) the organ of touch and temperature in the skin. I need only add that in the corium of man and all the higher Vertebrates countless microscopic sense-organs develop, but the precise relation of these to the sensations of pressure or resistance, of warmth and cold, has not yet been explained. Organs of this kind, in or on which sensory cutaneous nerves terminate, are the "tactile corpuscles" (or the Pacinian corpuscles) and end-bulbs. We find similar corpuscles in the organs of the s.e.xual sense, the male p.e.n.i.s and the female c.l.i.toris; they are processes of the skin, the development of which we will consider later (together with the rest of the s.e.xual parts, Chapter 2.29). The evolution of the organ of taste, the tongue and palate, will also be treated later, together with that of the alimentary ca.n.a.l to which these parts belong (Chapter 2.27). I will only point out for the present that the mucous coat of the tongue and palate, in which the gustatory nerve ends, originates from a part of the outer skin. As we have seen, the whole of the mouth-cavity is formed, not as a part of the gut-tube proper, but as a pit-like fold in the outer skin (Chapter 1.13). Its mucous lining is therefore formed, not from the visceral, but from the cutaneous layer, and the taste-cells at the surface of the tongue and palate are not products of the gut-fibre layer, but of the skin-sense layer.

This applies also to the mucous lining of the olfactory organ, the nose. However, the development of this organ is much more interesting.

Although the nose seems superficially to be simple and single, it really consists, in man and all other Gnathostomes, of two completely separated halves, the right and left cavities. They are divided by a vertical part.i.tion, so that the right nostril leads into the right cavity alone and the left nostril into the left cavity. They open internally (and separately) by the posterior nasal apertures into the pharynx, so that we can get direct into the gullet through the nasal pa.s.sages without touching the mouth. This is the way the air usually pa.s.ses in respiration; the mouth being closed, it goes through the nose into the gullet, and through the larynx and bronchial tubes into the lungs. The nasal cavities are separated from the mouth by the horizontal bony palate, to which is attached behind (as a dependent process) the soft palate with the uvula. In the upper and hinder parts of the nasal cavities the olfactory nerve, the first pair of cerebral nerves, expands in the mucous coat which clothes them. The terminal branches of it spread partly over the septum (part.i.tion), partly on the side walls of the internal cavities, to which are attached the turbinated bones. These bones are much more developed in many of the higher mammals than in man, but there are three of them in all mammals. The sensation of smell arises by the pa.s.sage of a current of air containing odorous matter over the mucous lining of the cavities, and stimulating the olfactory cells of the nerve-endings.

Man has all the features which distinguish the olfactory organ of the mammals from that of the lower Vertebrates. In all essential points the human nose entirely resembles that of the Catarrhine apes, some of which have quite a human external nose (compare the face of the long-nosed apes). However, the first structure of the olfactory organ in the human embryo gives no indication of the future ample proportions of our catarrhine nose. It has the form in which we find it permanently in the fishes--a couple of simple depressions in the skin at the outer surface of the head. We find these blind olfactory pits in all the fishes; sometimes they lie near the eyes, sometimes more forward at the point of the muzzle, sometimes lower down, near the mouth (Figure 2.249).

(FIGURE 2.311. Frontal section of the mouth and throat of a human embryo, neck half-inch long. "Invented" by Wilhelm His. The vertical section (in the frontal plane, from left to right) is so constructed that we see the nasal pits in the upper third of the figure and the eyes at the sides: in the middle third the primitive gullet with the gill-clefts (gill-arches in section); in the lower third the pectoral cavity with the bronchial tubes and the rudimentary lungs.)

This first rudimentary structure of the double nose is the same in all the Gnathostomes; it has no connection with the primitive mouth. But even in a section of the fishes a connection of this kind begins to make its appearance, a furrow in the surface of the skin running from each side of the nasal pit to the nearest corner of the mouth. This furrow, the nasal groove or furrow (Figure 2.305 r), is very important. In many of the sharks, such as the Scyllium, a special process of the frontal skin, the nasal fold or internal nasal process, is formed internally over the groove (n, n apostrophe). In contrast to this the outer edge of the furrow rises in an "external nasal process." As the two processes meet and coalesce over the nasal groove in the Dipneusts and Amphibia, it is converted into a ca.n.a.l, the nasal ca.n.a.l. Henceforth we can penetrate from the external pits through the nasal ca.n.a.ls direct into the mouth, which has been formed quite independently. In the Dipneusts and the lower Amphibia the internal aperture of the nasal ca.n.a.ls lies in front (behind the lips); in the higher Amphibia it is right behind. Finally, in the three higher cla.s.ses of Vertebrates the primary mouth-cavity is divided by the formation of the horizontal palate-roof into two distinct cavities--the upper (secondary) nasal cavity and the lower (secondary) mouth-cavity. The nasal cavity in turn is divided by the construction of the vertical septum into two halves--right and left.

(FIGURE 2.312. Diagrammatic section of the mouth-nose cavity. While the palate-plates (p) divide the original mouth-cavity into the lower secondary mouth (m) and the upper nasal cavity, the latter in turn is divided by the vertical part.i.tion (e) into two halves (n, n). (From Gegenbaur.))

Comparative anatomy shows us to-day, in the series of the double-nosed Vertebrates, from the fishes up to man, all the different stages in the development of the nose, which the advanced olfactory organ of the higher mammals has pa.s.sed through at various periods in the course of its phylogeny. It first appears in the embryo of man and the higher Vertebrates, in which the double fish-nose persists throughout life.

At an early stage, before there is any trace of the characteristic human face, a pair of small pits are formed in the head over the original mouth-cavity; these were first discovered by Baer, and rightly called the "olfactory pits" (Figures 2.306 n and 2.307 n).

These primitive nasal pits are quite separate from the rudimentary mouth, which also originates as a pit-like depression in the skin, in front of the blind fore end of the gut. Both the pair of nasal pits and the single mouth-pit (Figure 2.310 m) are clothed with the h.o.r.n.y plate. The original separation of the former from the latter is, however, presently abolished, a process forming above the mouth-pit--the "frontal process" (Figure 2.309 st). Its outer edge rises to the right and left in the shape of two lateral processes; these are the inner nasal processes or folds (in). Opposite to these a parallel ridge is formed on either side between the eye and the nasal pit; these are the outer nasal processes (an). Thus between the inner and outer nasal processes a groove-like depression is formed on either side, which leads from the nasal pit towards the mouth-pit (m); this groove is, as the reader will guess, the same nasal furrow or groove that we have already seen in the shark (Figure 2.305 r). As the parallel edges of the inner and outer nasal processes bend towards each other and join above the nasal groove, this is converted into a tube, the primitive nasal ca.n.a.l. Hence the nose of man and all the other Amniotes consists at this embryonic stage of a couple of narrow tubes, the nasal ca.n.a.ls, which lead from the outer surface of the forehead into the rudimentary mouth. This transitory condition resembles that in which we find the nose permanently in the Dipneusts and Amphibia.