XXII. OUR FIVE-TOED ANCESTORS
With the phylogenetic study of the four higher classes of Vertebrates, which must now engage our attention, we reach much firmer ground and more light in the construction of our genealogy than we have, perhaps, enjoyed up to the present. In the first place, we owe a number of very valuable data to the very interesting class of Vertebrates that come next to the Dipneusts and have been developed from them—the Amphibia. To this group belong the salamander, the frog, and the toad. In earlier days all the reptiles were, on the example of Linne, classed with the Amphibia (lizards, serpents, crocodiles, and tortoises). But the reptiles are much more advanced than the Amphibia, and are nearer to the birds in the chief points of their structure. The true Amphibia are nearer to the Dipneusta and the fishes; they are also much older than the reptiles. There were plenty of highly-developed (and sometimes large) Amphibia during the Carboniferous period; but the earliest reptiles are only found in the Permian period. It is probable that the Amphibia were evolved even earlier—during the Devonian period—from the Dipneusta. The extinct Amphibia of which we have fossil remains from that remote period (very numerous especially in the Triassic strata) were distinguished for a graceful scaly coat or a powerful bony armour on the skin (like the crocodile), whereas the living amphibia have usually a smooth and slippery skin.
The earliest of these armoured Amphibia (Phractamphibia) form the order of Stegocephala (“roof-headed”) (Fig. 260). It is among these, and not among the actual Amphibia, that we must look for the forms that are directly related to the genealogy of our race, and are the ancestors of the three higher classes of Vertebrates. But even the existing Amphibia have such important relations to us in their anatomic structure, and especially their embryonic development, that we may say: Between the Dipneusts and the Amniotes there was a series of extinct intermediate forms which we should certainly class with the Amphibia if we had them before us. In their whole organisation even the actual Amphibia seem to be an instructive transitional group. In the important respects of respiration and circulation they approach very closely to the Dipneusta, though in other respects they are far superior to them.
This is particularly true of the development of their limbs or extremities. In them we find these for the first time as five-toed feet. The thorough investigations of Gegenbaur have shown that the fish’s fins, of which very erroneous opinions were formerly held, are many-toed feet. The various cartilaginous or bony radii that are found in large numbers in each fin correspond to the fingers or toes of the higher Vertebrates. The several joints of each fin-radius correspond to the various parts of the toe. Even in the Dipneusta the fin is of the same construction as in the fishes; it was afterwards gradually evolved into the five-toed form, which we first encounter in the Amphibia. This reduction of the number of the toes to six, and then to five, probably took place in the second half of the Devonian period—at the latest, in the subsequent Carboniferous period—in those Dipneusta which we regard as the ancestors of the Amphibia. We have several fossil remains of five-toed Amphibia from this period. There are numbers of fossil impressions of them in the Triassic of Thuringia (Chirotherium).
The fact that the toes number five is of great importance, because they have clearly been transmitted from the Amphibia to all the higher Vertebrates. Man entirely resembles his amphibian ancestors in this respect, and indeed in the whole structure of the bony skeleton of his five-toed extremities. A careful comparison of the skeleton of the frog with our own is enough to show this. It is well known that this hereditary number of the toes has assumed a very great practical importance from remote times; on it our whole system of enumeration (the decimal system applied to measurement of time, mass, weight, etc.) is based. There is absolutely no reason why there should be five toes in the fore and hind feet in the lowest Amphibia, the reptiles, and the higher Vertebrates, unless we ascribe it to inheritance from a common stem-form. Heredity alone can explain it. It is true that we find less than five toes in many of the Amphibia and of the higher Vertebrates. But in all these cases we can prove that some of the toes atrophied, and were in time lost altogether.
Fig.260. Fossil amphibian from the Permian, found in the Plauen terrain near Dresden (Branchiosaurus amblystomus). Fig. 260—Fossil amphibian from the Permian, found in the Plauen terrain near Dresden (Branchiosaurus amblystomus). (From Credner.) A skeleton of a young larva. B larva, restored, with gills. C the adult form.)
The causes of this evolution of the five-toed foot from the many-toed fin in the amphibian ancestor must be sought in adaptation to the entire change of function that the limbs experienced in passing from an exclusively aquatic to a partly terrestrial life. The many-toed fin had been used almost solely for motion in the water; it had now also to support the body in creeping on the solid ground. This led to a modification both of the skeleton and the muscles of the limbs. The number of the fin-radii was gradually reduced, and sank finally to five. But these five remaining radii became much stronger. The soft cartilaginous radii became bony rods. The rest of the skeleton was similarly strengthened. Thus from the one-armed lever of the many-toed fish-fin arose the improved many-armed lever system of the five-toed amphibian limbs. The movements of the body gained in variety as well as in strength. The various parts of the skeletal system and correlated muscular system began to differentiate more and more. In view of the close correlation of the muscular and nervous systems, this also made great advance in structure and function. Hence we find, as a matter of fact, that the brain is much more developed in the higher Amphibia than in the fishes, the Dipneusta, and the lower Amphibia.
Fig.261. Larva of the Spotted Salamander (Salamandra maculata), seen from the ventral side. Fig. 261—Larva of the Spotted Salamander (Salamandra maculata), seen from the ventral side. In the centre a yelk-sac still hangs from the gut. The external gills are gracefully ramified. The two pairs of legs are still very small.
The first advance in organisation that was occasioned by the adoption of life on land was naturally the construction of an organ for breathing air—a lung. This was formed directly from the floating-bladder inherited from the fishes. At first its function was insignificant beside that of the gills, the older organ for water-respiration. Hence we find in the lowest Amphibia, the gilled Amphibia, that, like the Dipneusta, they pass the greater part of their life in the water, and breathe water through gills. They only come to the surface at brief intervals, or creep on to the land, and then breathe air by their lungs. But some of the tailed Amphibia—the salamanders—remain entirely in the water when they are young, and afterwards spend most of their time on land. In the adult state they only breathe air through lungs. The same applies to the most advanced of the Amphibia, the Batrachia (frogs and toads); some of them have entirely lost the gill-bearing larva form.[30] This is also the case with certain small, serpentine Amphibia, the Cæcilia (which live in the ground like earth-worms).
[30] The tree-frog of Martinique (Hylades martinicensis) loses the gills on the seventh, and the tail and yelk-sac on the eighth, day of fœtal life. On the ninth or tenth day after fecundation the frog emerges from the egg.
The great interest of the natural history of the Amphibia consists especially in their intermediate position between the lower and higher Vertebrates. The lower Amphibia approach very closely to the Dipneusta in their whole organisation, live mainly in the water, and breathe by gills; but the higher Amphibia are just as close to the Amniotes, live mainly on land, and breathe by lungs. But in their younger state the latter resemble the former, and only reach the higher stage by a complete metamorphosis. The embryonic development of most of the higher Amphibia still faithfully reproduces the stem-history of the whole class, and the various stages of the advance that was made by the lower Vertebrates in passing from aquatic to terrestrial life during the Devonian or the Carboniferous period are repeated in the spring by every frog that develops from an egg in our ponds.
Fig.262. Larva of the common grass-frog (Rana temporaria), or “tadpole.” Fig. 262—Larva of the common grass-frog (Rana temporaria), or “tadpole.” m mouth, n a pair of suckers for fastening on to stones, d skin-fold from which the gill-cover develops; behind it the gill-clefts, from which the branching gills (k) protrude, s tail-muscles, f cutaneous fin-fringe of the tail.
The common frog leaves the egg in the shape of a larva, like the tailed salamander (Fig. 261), and this is altogether different from the mature frog (Fig. 262). The short trunk ends in a long tail, with the form and structure of a fish’s tail (s). There are no limbs at first. The respiration is exclusively branchial, first through external (k) and then internal gills. In harmony with this the heart has the same structure as in the fish, and consists of two sections—an atrium that receives the venous blood from the body, and a ventricle that forces it through the arteries into the gills.
We find the larvæ of the frog (or tadpoles, Gyrini) in great numbers in our ponds every spring in this fish-form, using their muscular tails in swimming, just like the fishes and young Ascidia. When they have reached a certain size, the remarkable metamorphosis from the fish-form to the frog begins. A blind sac grows out of the gullet, and expands into a couple of spacious sacs: these are the lungs. The simple chamber of the heart is divided into two sections by the development of a partition, and there are at the same time considerable changes in the structure of the chief arteries. Previously all the blood went from the auricle through the aortic arches into the gills, but now only part of it goes to the gills, the other part passing to the lungs through the new-formed pulmonary artery. From this point arterial blood returns to the left auricle of the heart, while the venous blood gathers in the right auricle. As both auricles open into a single ventricle, this contains mixed blood. The dipneust form has now succeeded to the fish-form. In the further course of the metamorphosis the gills and the branchial vessels entirely disappear, and the respiration becomes exclusively pulmonary. Later, the long swimming tail is lost, and the frog now hops to the land with the legs that have grown meantime.
This remarkable metamorphosis of the Amphibia is very instructive in connection with our human genealogy, and is particularly interesting from the fact that the various groups of actual Amphibia have remained at different stages of their stem-history, in harmony with the biogenetic law. We have first of all a very low order of Amphibia—the Sozobranchia (“gilled-amphibia”), which retain their gills throughout life, like the fishes. In a second order of the salamanders the gills are lost in the metamorphosis, and when fully grown they have only pulmonary respiration. Some of the tailed Amphibia still retain the gill-clefts in the side of the neck, though they have lost the gills themselves (Menopoma). If we force the larvæ of our salamanders (Fig. 261) and tritons to remain in the water, and prevent them from reaching the land, we can in favourable circumstances make them retain their gills. In this fish-like condition they reach sexual maturity, and remain throughout life at the lower stage of the gilled Amphibia.
fish-like axolotl (Siredon pisciformis). It was formerly regarded as a permanent gilled amphibian persisting throughout life at the fish-stage. But some of the hundreds of these animals that are kept in the Botanical Garden at Paris got on to the land for some reason or other, lost their gills, and changed into a form closely resembling the salamander (Amblystoma). Other species of the genus became sexually mature for the first time in this condition. This has been regarded as an astounding phenomenon, although every common frog and salamander repeats the metamorphosis in the spring. The whole change from the aquatic and gill-breathing animal to the terrestrial lung-breathing form may be followed step by step in this case. But what we see here in the development of the individual has happened to the whole class in the course of its stem-history.
Fig.263. Fossil mailed amphibian, from the Bohemian Carboniferous (Seeleya). Fig. 263—Fossil mailed amphibian, from the Bohemian Carboniferous (Seeleya). (From Fritsch.) The scaly coat is retained on the left.
The metamorphosis goes farther in a third order of Amphibia, the Batrachia or Anura, than in the salamander. To this belong the various kinds of toads, ringed snakes, water-frogs, tree-frogs, etc. These lose, not only the gills, but also (sooner or later) the tail, during metamorphosis.
The ontogenetic loss of the gills and the tail in the frog and toad can only be explained on the assumption that they are descended from long-tailed Amphibia of the salamander type. This is also clear from the comparative anatomy of the two groups. This remarkable metamorphosis is, however, also interesting because it throws a certain light on the phylogeny of the tail-less apes and man. Their ancestors also had long tails and gills like the gilled Amphibia, as the tail and the gill-arches of the human embryo clearly show.
For comparative anatomical and ontogenetic reasons, we must not seek these amphibian ancestors of ours—as one would be inclined to do, perhaps—among the tail-less Batrachia, but among the tailed lower Amphibia.
The vertebrate form that comes next to the Amphibia in the series of our ancestors is a lizard-like animal, the earlier existence of which can be confidently deduced from the facts of comparative anatomy and ontogeny. The living Hatteria of New Zealand (Fig. 264) and the extinct Rhyncocephala of the Permian period (Fig. 265) are closely related to this important stem-form; we may call them the Protamniotes, or Primitive Amniotes. All the Vertebrates above the Amphibia—or the three classes of reptiles, birds, and mammals—differ so much in their whole organisation from all the lower Vertebrates we have yet considered, and have so great a resemblance to each other, that we put them all together in a single group with the title of Amniotes. In these three classes alone we find the remarkable embryonic membrane, already mentioned, which we called the amnion; a cenogenetic adaptation that we may regard as a result of the sinking of the growing embryo into the yelk-sac.
All the Amniotes known to us—all reptiles, birds, and mammals (including man)—agree in so many important points of internal structure and development that their descent from a common ancestor can be affirmed with tolerable certainty. If the evidence of comparative anatomy and ontogeny is ever entirely beyond suspicion, it is certainly the case here. All the peculiarities that accompany and follow the formation of the amnion, and that we have learned in our consideration of human embryology; all the peculiarities in the development of the organs which we will presently follow in detail; finally, all the principal special features of the internal structure of the full-grown Amniotes—prove so clearly the common origin of all the Amniotes from single extinct stem-form that it is difficult to entertain the idea of their evolution from several independent stems. This unknown common stem-form is our primitive Amniote (Protamnion). In outward appearance it was probably something between the salamander and the lizard.
It is very probable that some part of the Permian period was the age of the origin of the Protamniotes. This follows from the fact that the Amphibia are not fully developed until the Carboniferous period, and that the first fossil reptiles (Palæhatteria, Homœosaurus, Proterosaurus) are found towards the close of the Permian period. Among the important changes of the vertebrate organisation that marked the rise of the first Amniotes from salamandrine Amphibia during this period the following three are especially noteworthy: the entire disappearance of the water-breathing gills and the conversion of the gill-arches into other organs, the formation of the allantois or primitive urinary sac, and the development of the amnion.
One of the most salient characteristics of the Amniotes is the complete loss of the gills. All Amniotes, even if living in water (such as sea-serpents and whales), breathe air through lungs, never water through gills. All the Amphibia (with very rare exceptions) retain their gills for some time when young, and have for a time (if not permanently) branchial respiration; but after these there is no question of branchial respiration. The Protamniote itself must have entirely abandoned water-breathing. Nevertheless, the gill-arches are preserved by heredity, and develop into totally different (in part rudimentary) organs—various parts of the bone of the tongue, the frame of the jaws, the organ of hearing, etc. But we do not find in the embryos of the Amniotes any trace of gill-leaves, or of real respiratory organs on the gill-arches.
With this complete abandonment of the gills is probably connected the formation of another organ, to which we have already referred in embryology—namely, the allantois or primitive urinary sac (cf. p. 166). It is very probable that the urinary bladder of the Dipneusts is the first structure of the allantois. We find in these a urinary bladder that proceeds from the lower wall of the hind end of the gut, and serves as receptacle for the renal secretions. This organ has been transmitted to the Amphibia, as we can see in the frog.
The formation of the amnion and the allantois and the complete disappearance of the gills are the chief characteristics that distinguish the Amniotes from the lower Vertebrates we have hitherto considered. To these we may add several subordinate features that are transmitted to all the Amniotes, and are found in these only. One striking embryonic character of the Amniotes is the great curve of the head and neck in the embryo. We also find an advance in the structure of several of the internal organs of the Amniotes which raises them above the highest of the anamnia. In particular, a partition is formed in the simple ventricle of the heart, dividing into right and left chambers. In connection with the complete metamorphosis of the gill-arches we find a further development of the auscultory organs. Also, there is a great advance in the structure of the brain, skeleton, muscular system, and other parts. Finally, one of the most important changes is the reconstruction of the kidneys. In all the earlier Vertebrates we have found the primitive kidneys as excretory organs, and these appear at an early stage in the embryos of all the higher Vertebrates up to man. But in the Amniotes these primitive kidneys cease to act at an early stage of embryonic life, and their function is taken up by the permanent or secondary kidneys, which develop from the terminal section of the prorenal ducts.
Taking all these peculiarities of the Amniotes together, it is impossible to doubt that all the animals of this group—all reptiles, birds, and mammals—have a common origin, and form a single blood-related stem. Our own race belongs to this stem. Man is, in every feature of his organisation and embryonic development, a true Amniote, and has descended from the Protamniote with all the other Amniotes. Though they appeared at the end (possibly even in the middle) of the Paleozoic age, the Amniotes only reached their full development during the Mesozoic age. The birds and mammals made their first appearance during this period. Even the reptiles show their greatest growth at this time, so that it is called “the reptile age.” The extinct Protamniote, the ancestor of the whole group, belongs in its whole organisation to the reptile class.
Fig.264. The lizard (Hatteria punctata = Sphenodon punctatus) of New Zealand. Fig. 264—The lizard (Hatteria punctata = Sphenodon punctatus) of New Zealand. The sole surviving proreptile. (From Brehm.)
The genealogical tree of the amniote group is clearly indicated in its chief lines by their paleontology, comparative anatomy, and ontogeny. The group succeeding the Protamniote divided into two branches. The branch that will claim our whole interest is the class of the Mammals. The other branch, which developed in a totally different direction, and only comes in contact with the Mammals at its root, is the combined group of the reptiles and birds; these two classes may, with Huxley, be conveniently grouped together as the Sauropsida. Their common stem-form is an extinct lizard-like reptile of the order of the Rhyncocephalia. From this have been developed in various directions the serpents, crocodiles, tortoises, etc.—in a word, all the members of the reptile class. But the remarkable class of the birds has also been evolved directly from a branch of the reptile group, as is now established beyond question. The embryos of the reptiles and birds are identical until a very late stage, and have an astonishing resemblance even later. Their whole structure agrees so much that no anatomist now questions the descent of the birds from the reptiles. On the other hand, the mammal line has descended from the group of the Sauromammalia, a different branch of the Proreptilia. It is connected at its deepest roots with the reptile line, but it then diverges completely from it and follows a distinctive development. Man is the highest outcome of this class, the “crown of creation.” The hypothesis that the three higher Vertebrate classes represent a single Amniote-stem, and that the common root of this stem is to be found in the amphibian class, is now generally admitted.
Fig.265. Homoeosaurus pulchellus, a Jurassic proreptile from Kehlheim. Fig. 265—Homœosaurus pulchellus, a Jurassic proreptile from Kehlheim. (From Zittel.)
The instructive group of the Permian Tocosauria, the common root from which the divergent stems of the Sauropsids and mammals have issued, merits our particular attention as the stem-group of all the Amniotes. Fortunately a living representative of this extinct ancestral group has been preserved to our day; this is the remarkable lizard of New Zealand, Hatteria punctata (Fig. 264). Externally it differs little from the ordinary lizard; but in many important points of internal structure, especially in the primitive construction of the vertebral column, the skull, and the limbs, it occupies a much lower position, and approaches its amphibian ancestors, the Stegocephala. Hence Hatteria is the phylogenetically oldest of all living reptiles, an isolated survivor from the Permian period, closely resembling the common ancestor of the Amniotes. It must differ so little from this extinct form, our hypothetical Protamniote, that we put it next to the Proreptilia. The remarkable Permian Palæhatteria, that Credner discovered in the Plauen terrain at Dresden in 1888, belongs to the same group (Fig. 266). The Jurassic genus Homœosaurus (Fig. 265), of which well-preserved skeletons are found in the Solenhofen schists, is perhaps still more closely related to them.
Unfortunately, the numerous fossil remains of Permian and Triassic Tocosauria that we have found in the last two decades are, for the most part, very imperfectly preserved. Very often we can make only precarious inferences from these skeletal fragments as to the anatomic characters of the soft parts that went with the bony skeleton of the extinct Tocosauria. Hence it has not yet been possible to arrange these important fossils with any confidence in the ancestral series that descend from the Protamniotes to the Sauropsids on the one side and the Mammals on the other. Opinions are particularly divided as to the place in classification and the phylogenetic significance of the remarkable Theromorpha. Cope gives this name to a very interesting and extensive group of extinct terrestrial reptiles, of which we have only fossil remains from the Permian and Triassic strata. Forty years ago some of these Therosauria (fresh-water animals) were described by Owen as Anomodontia. But during the last twenty years the distinguished American paleontologists, Cope and Osborn, have greatly increased our knowledge of them, and have claimed that the stem-forms of the Mammals must be sought in this order. As a matter of fact, the Theromorpha are nearer to the Mammals in the chief points of structure than any other reptiles. This is especially true of the Thereodontia, to which the Pureosauria and Pelycosauria belong (Fig. 267). The whole structure of their pelvis and hind-feet has attained the same form as in the Monotremes, the lowest Mammals. The formation of the scapula and the quadrate bone shows an approach to the Mammals such as we find in no other group of reptiles. The teeth also are already divided into incisors, canines, and molars. Nevertheless, it is very doubtful whether the Theromorpha really are in the ancestral line of the Sauromammals, or lead direct from the Tocosauria to the earliest Mammals. Other experts on this group believe that it is an independent legion of the reptiles, connected, perhaps, at its lowest root, with the Sauromammals, but developed quite independently of the Mammals—though parallel to them in many ways.
One of the most important of the zoological facts that we rely on in our investigation of the genealogy of the human race is the position of man in the Mammal class. However different the views of zoologists may have been as to this position in detail, and as to his relations to the apes, no scientist has ever doubted that man is a true mammal in his whole organisation and development. Linné drew attention to this fact in the first edition of his famous Systema Naturæ (1735). As will be seen in any museum of anatomy or any manual of comparative anatomy; the human frame has all the characteristics that are common to the Mammals and distinguish them conspicuously from all other animals.
Fig.266. Skull of a Permian lizard (Palaehatteria longicaudata). Fig. 266—Skull of a Permian lizard (Palæhatteria longicaudata). (From Credner.) n nasal bone, pf frontal bone, l lachrymal bone, po postorbital bone, sq covering bone, i cheek-bone, vo vomer, im inter-maxillary.
If we examine this undoubted fact from the point of view of phylogeny, in the light of the theory of descent, it follows at once that man is of a common stem with all the other Mammals, and comes from the same root as they. But the various features in which the Mammals agree and by which they are distinguished are of such a character as to make a polyphyletic hypothesis quite inadmissible. It is impossible to entertain the idea that all the living and extinct Mammals come from a number of separate roots. If we accept the general theory of evolution, we are bound to admit the monophyletic hypothesis of the descent of all the Mammals (including man) from a single mammalian stem-form. We may call this long-extinct root-form and its earliest descendants (a few genera of one family) “primitive mammals” or “stem-mammals” (Promammalia). As we have already seen, this root-form developed from the primitive Proreptile stem in a totally different direction from the birds, and soon separated from the main stem of the reptiles. The differences between the Mammals and the reptiles and birds are so important and characteristic that we can assume with complete confidence this division of the vertebrate stem at the commencement of the development of the Amniotes. The reptiles and birds, which we group together as the Sauropsids, generally agree in the characteristic structure of the skull and brain, and this is notably different from that of the Mammals. In most of the reptiles and birds the skull is connected with the first cervical vertebra (the atlas) by a single, and in the Mammals (and Amphibia) by a double, condyle at the back of the head. In the former the lower jaw is composed of several pieces, and connected with the skull so that it can move by a special maxillary bone (the quadratum); in the Mammals the lower jaw consists of one pair of bony pieces, which articulate directly with the temporal bone. Further, in the Sauropsids the skin is clothed with scales or feathers; in the Mammals with hair. The red blood-cells of the former have a nucleus; those of the latter have not. In fine, two quite characteristic features of the Mammals, which distinguish them not only from the birds and reptiles, but from all other animals, are the possession of a complete diaphragm and of mammary glands that produce the milk for the nutrition of the young. It is only in the Mammals that the diaphragm forms a transverse partition of the body-cavity, completely separating the pectoral from the abdominal cavity. It is only in the mammals that the mother suckles its young, and this rightly gives the name to the whole class (mamma = breast).
Fig.267. Skull of a Triassic theromorphum (Galesaurus planiceps), from the Karoo formation in South Africa. Fig. 267—Skull of a Triassic theromorphum (Galesaurus planiceps), from the Karoo formation in South Africa. (From Owen.) a from the right, b from below, c from above, d tricuspid tooth. N nostrils, Na nasal bone, Mx upper jaw, Prf prefrontal, Fr frontal bone, A eye-pits, S temple-pits. Pa Parietal eye, Bo joint at back of head, Pt pterygoid-bone, Md lower jaw.
From these pregnant facts of comparative anatomy and ontogeny it follows absolutely that the whole of the Mammals belong to a single natural stem, which branched off at an early date from the reptile-root. It follows further with the same absolute certainty that the human race is also a branch of this stem. Man shares all the characteristics I have described with all the Mammals, and differs in them from all other animals. Finally, from these facts we deduce with the same confidence those advances in the vertebrate organisation by which one branch of the Sauromammals was converted into the stem-form of the Mammals. Of these advances the chief were: (1) The characteristic modification of the skull and the brain; (2) the development of a hairy coat; (3) the complete formation of the diaphragm; and (4) the construction of the mammary glands and adaptation to suckling. Other important changes of structure proceeded step by step with these.
The epoch at which these important advances were made, and the foundation of the Mammal class was laid, may be put with great probability in the first section of the Mesozoic or secondary age—the Triassic period. The oldest fossil remains of mammals that we know were found in strata that belong to the earliest Triassic period—the upper Kueper. One of the earliest forms is the genus Dromatherium, from the North American Triassic (Fig. 268). Their teeth still strikingly recall those of the Pelycosauria. Hence we may assume that this small and probably insectivorous mammal belonged to the stem-group of the Promammals. We do not find any positive trace of the third and most advanced division of the Mammals—the Placentals. These (including man) are much younger, and we do not find indisputable fossil remains of them until the Cenozoic age, or the Tertiary period. This paleontological fact is very important, because it fully harmonises with the evolutionary succession of the Mammal orders that is deduced from their comparative anatomy and ontogeny.
The latter science teaches us that the whole Mammal class divides into three main groups or sub-classes, which correspond to three successive phylogenetic stages. These three stages, which also represent three important stages in our human genealogy, were first distinguished in 1816 by the eminent French zoologist, Blainville, and received the names of Ornithodelphia, Didelphia, and Monodelphia, according to the construction of the female organs (delphys = uterus or womb). Huxley afterwards gave them the names of Prototheria, Metatheria, and Epitheria. But the three sub-classes differ so widely from each other, not only in the construction of the sexual organs, but in many other respects also, that we may confidently draw up the following important phylogenetic thesis: The Monodelphia or Placentals descend from the Didelphia or Marsupials; and the latter, in turn, are descended from the Monotremes or Ornithodelphia.
Thus we must regard as the twenty-first stage in our genealogical tree the earliest and lowest chief group of the Mammals—the sub-class of the Monotremes (“cloaca-animals,” Ornithodelphia, or Prototheria, Figs. 269 and 270). They take their name from the cloaca which they share with all the lower Vertebrates. This cloaca is the common outlet for the passage of the excrements, the urine, and the sexual products. The urinary ducts and sexual canals open into the hindmost part of the gut, while in all the other Mammals they are separated from the rectum and anus. The latter have a special uro-genital outlet (porus urogenitalis). The bladder also opens into the cloaca in the Monotremes, and, indeed, apart from the two urinary ducts; in all the other Mammals the latter open directly into the bladder. It was proved by Haacke and Caldwell in 1884 that the Monotremes lay large eggs like the reptiles, while all the other Mammals are viviparous. In 1894 Richard Semon further proved that these large eggs, rich in food-yelk, have a partial segmentation and discoid gastrulation, as I had hypothetically assumed in 1879; here again they resemble their reptilian ancestors. The construction of the mammary gland is also peculiar in the Monotremes. In them the glands have no teats for the young animal to suck, but there is a special part of the breast pierced with holes like a sieve, from which the milk issues, and the young Monotreme must lick it off. Further, the brain of the Monotremes is very little advanced. It is feebler than that of any of the other Mammals. The fore-brain or cerebrum, in particular, is so small that it does not cover the cerebellum. In the skeleton (Fig. 270) the formation of the scapula among other parts is curious; it is quite different from that of the other Mammals, and rather agrees with that of the reptiles and Amphibia. Like these, the Monotremes have a strongly developed caracoideum. From these and other less prominent characteristics it follows absolutely that the Monotremes occupy the lowest place among the Mammals, and represent a transitional group between the Tocosauria and the rest of the Mammals. All these remarkable reptilian characters must have been possessed by the stem-form of the whole mammal class, the Promammal of the Triassic period, and have been inherited from the Proreptiles.
Fig.268. Lower jaw of a Primitive Mammal or Promammal (Dromatherium silvestre) from the North American Triassic. Fig. 268—Lower jaw of a Primitive Mammal or Promammal (Dromatherium silvestre) from the North American Triassic. i incisors, c canine, p premolars, m molars. (From Döderlein.)
During the Triassic and Jurassic periods the sub-class of the Monotremes was represented by a number of different stem-mammals. Numerous fossil remains of them have lately been discovered in the Mesozoic strata of Europe, Africa, and America. To-day there are only two surviving specimens of the group, which we place together in the family of the duck-bills, Ornithostoma. They are confined to Australia and the neighbouring island of Van Diemen’s Land (or Tasmania); they become scarcer every year, and will soon, like their blood-relatives, be counted among the extinct animals. One form lives in the rivers, and builds subterraneous dwellings on the banks; this is the Ornithorhyncus paradoxus, with webbed feet, a thick soft fur, and broad flat jaws, which look very much like the bill of a duck (Figs. 269, 270). The other form, the land duck-bill, or spiny ant-eater (Echidna hystrix), is very much like the anteaters in its habits and the peculiar construction of its thin snout and very long tongue; it is covered with needles, and can roll itself up like a hedgehog. A cognate form (Parechidna Bruyni) has lately been found in New Guinea.
These modern Ornithostoma are the scattered survivors of the vast Mesozoic group of Monotremes; hence they have the same interest in connection with the stem history of the Mammals as the living stem-reptiles (Hatteria) for that of the reptiles, and the isolated Acrania (Amphioxus) for the phylogeny of the Vertebrate stem.
The Australian duck-bills are distinguished externally by a toothless bird-like beak or snout. This absence of real bony teeth is a late result of adaptation, as in the toothless Placentals (Edentata, armadillos and ant-eaters). The extinct Monotremes, to which the Promammalia belonged, must have had developed teeth, inherited from the reptiles. Lately small rudiments of real molars have been discovered in the young of the Ornithorhyncus, which has horny plates in the jaws instead of real teeth.
Fig. 269. The Ornithorhyncus or Duck-mole. (Ornithorhyncus paradoxus). Fig. 269—The Ornithorhyncus or Duck-mole. (Ornithorhyncus paradoxus).
Fig. 270. Skeleton of the Ornithorhyncus. Fig. 270—Skeleton of the Ornithorhyncus.
The living Ornithostoma and the stem-forms of the Marsupials (or Didelphia) must be regarded as two widely diverging lines from the Promammals. This second sub-class of the Mammals is very interesting as a perfect intermediate stage between the other two. While the Marsupials retain a great part of the characteristics of the Monotremes, they have also acquired some of the chief features of the Placentals. Some features are also peculiar to the Marsupials, such as the construction of the male and female sexual organs and the form of the lower jaw. The Marsupials are distinguished by a peculiar hook-like bony process that bends from the corner of the lower jaw and points inwards. As most of the Placentals have not this process, we can, with some probability, recognise the Marsupial from this feature alone. Most of the mammal remains that we have from the Jurassic and Cretaceous deposits are merely lower jaws, and most of the jaws found in the Jurassic deposits at Stonesfield and Purbeck have the peculiar hook-like process that characterises the lower jaw of the Marsupial. On the strength of this paleontological fact, we may suppose that they belonged to Marsupials. Placentals do not seem to have existed at the middle of the Mesozoic age—not until towards its close (in the Cretaceous period). At all events, we have no fossil remains of indubitable Placentals from that period.
The existing Marsupials, of which the plant-eating kangaroo and the carnivorous opossum (Fig. 272) are the best known, differ a good deal in structure, shape, and size, and correspond in many respects to the various orders of Placentals. Most of them live in Australia, and a small part of the Australian and East Malayan islands. There is now not a single living Marsupial on the mainland of Europe, Asia, or Africa. It was very different during the Mesozoic and even during the Cenozoic age. The sedimentary deposits of these periods contain a great number and variety of marsupial remains, sometimes of a colossal size, in various parts of the earth, and even in Europe. We may infer from this that the existing Marsupials are the remnant of an extensive earlier group that was distributed all over the earth. It had to give way in the struggle for life to the more powerful Placentals during the Tertiary period. The survivors of the group were able to keep alive in Australia and South America because the one was completely separated from the other parts of the earth during the whole of the Tertiary period, and the other during the greater part of it.
Fig.271. Lower jaw of a Promammal (Dryolestes priscus), from the Jurassic of the Felsen strata. Fig. 271—Lower jaw of a Promammal (Dryolestes priscus), from the Jurassic of the Felsen strata. (From Marsh.)
From the comparative anatomy and ontogeny of the existing Marsupials we may draw very interesting conclusions as to their intermediate position between the earlier Monotremes and the later Placentals. The defective development of the brain (especially the cerebrum), the possession of marsupial bones, and the simple construction of the allantois (without any placenta as yet) were inherited by the Marsupials, with many other features, from the Monotremes, and preserved. On the other hand, they have lost the independent bone (caracoideum) at the shoulder-blade. But we have a more important advance in the disappearance of the cloaca; the rectum and anus are separated by a partition from the uro-genital opening (sinus urogenitalis). Moreover, all the Marsupials have teats on the mammary glands, at which the new-born animal sucks. The teats pass into the cavity of a pouch or pocket on the ventral side of the mother, and this is supported by a couple of marsupial bones. The young are born in a very imperfect condition, and carried by the mother for some time longer in her pouch, until they are fully developed (Fig. 272). In the giant kangaroo, which is as tall as a man, the embryo only develops for a month in the uterus, is then born in a very imperfect state, and finishes its growth in the mother’s pouch (marsupium); it remains in this about nine months, and at first hangs continually on to the teat of the mammary gland.
From these and other characteristics (especially the peculiar construction of the internal and external sexual organs in male and female) it is clear that we must conceive the whole sub-class of the Marsupials as one stem group, which has been developed from the Promammalia. From one branch of these Marsupials (possibly from more than one) the stem-forms of the higher Mammals, the Placentals, were afterwards evolved. Of the existing forms of the Marsupials, which have undergone various modifications through adaptation to different environments, the family of the opossums (Didelphida or Pedimana) seems to be the oldest and nearest to the common stem-form of the whole class. To this family belong the crab-eating opossum of Brazil (Fig. 272) and the opossum of Virginia, on the embryology of which Selenka has given us a valuable work (cf. Figs. 63–67 and 131–135). These Didelphida climb trees like the apes, grasping the branches with their hand-shaped hind feet. We may conclude from this that the stem-forms of the Primates, which we must regard as the earliest Lemurs, were evolved directly from the opossum. We must not forget, however, that the conversion of the five-toed foot into a prehensile hand is polyphyletic. By the same adaptation to climbing trees the habit of grasping their branches with the feet has in many different cases brought about that opposition of the thumb or great toe to the other toes which makes the hand prehensile. We see this in the climbing lizards (chameleon), the birds, and the tree-dwelling mammals of various orders.
Fig.272. The crab-eating Opossum (Philander cancrivorus). The female has three young in the pouch. Fig. 272—The crab-eating Opossum (Philander cancrivorus). The female has three young in the pouch. (From Brehm.)
Some zoologists have lately advanced the opposite opinion, that the Marsupials represent a completely independent sub-class of the Mammals, with no direct relation to the Placentals, and developing independently of them from the Monotremes. But this opinion is untenable if we examine carefully the whole organisation of the three sub-classes, and do not lay the chief stress on incidental features and secondary adaptations (such as the formation of the marsupium). It is then clear that the Marsupials—viviparous Mammals without placenta—are a necessary transition from the oviparous Monotremes to the higher Placentals with chorion-villi. In this sense the Marsupial class certainly contains some of man’s ancestors.
XXIII. OUR APE ANCESTORS
The long series of animal forms which we must regard as the ancestors of our race has been confined within narrower and narrower circles as our phylogenetic inquiry has progressed. The great majority of known animals do not fall in the line of our ancestry, and even within the vertebrate stem only a small number are found to do so. In the most advanced class of the stem, the mammals, there are only a few families that belong directly to our genealogical tree. The most important of these are the apes and their predecessors, the half-apes, and the earliest Placentals (Prochoriata).
The Placentals (also called Choriata, Monodelphia, Eutheria or Epitheria) are distinguished from the lower mammals we have just considered, the Monotremes and Marsupials, by a number of striking peculiarities. Man has all these distinctive features; that is a very significant fact. We may, on the ground of the most careful comparative-anatomical and ontogenetic research, formulate the thesis: “Man is in every respect a true Placental.” He has all the characteristics of structure and development that distinguish the Placentals from the two lower divisions of the mammals, and, in fact, from all other animals. Among these characteristics we must especially notice the more advanced development of the brain. The fore-brain or cerebrum especially is much more developed in them than in the lower animals. The corpus callosum, which forms a sort of wide bridge connecting the two hemispheres of the cerebrum, is only fully formed in the Placentals; it is very rudimentary in the Marsupials and Monotremes. It is true that the lowest Placentals are not far removed from the Marsupials in cerebral development; but within the placental group we can trace an unbroken gradation of progressive development of the brain, rising gradually from this lowest stage up to the elaborate psychic organ of the apes and man. The human soul—a physiological function of the brain—is in reality only a more advanced ape-soul.
The mammary glands of the Placentals are provided with teats like those of the Marsupials; but we never find in the Placentals the pouch in which the latter carry and suckle their young. Nor have they the marsupial bones in the ventral wall at the anterior border of the pelvis, which the Marsupials have in common with the Monotremes, and which are formed by a partial ossification of the sinews of the inner oblique abdominal muscle. There are merely a few insignificant remnants of them in some of the Carnivora. The Placentals are also generally without the hook-shaped process at the angle of the lower jaw which is found in the Marsupials.
However, the feature that characterises the Placentals above all others, and that has given its name to the whole sub-class, is the formation of the placenta. We have already considered the formation and significance of this remarkable embryonic organ when we traced the development of the chorion and the allantois in the human embryo (pp.165–9) The urinary sac or the allantois, the curious vesicle that grows out of the hind part of the gut, has essentially the same structure and function in the human embryo as in that of all the other Amniotes (cf. Figs. 194–6). There is a quite secondary difference, on which great stress has wrongly been laid, in the fact that in man and the higher apes the original cavity of the allantois quickly degenerates, and the rudiment of it sticks out as a solid projection from the primitive gut. The thin wall of the allantois consists of the same two layers or membranes as the wall of the gut—the gut-gland layer within and the gut-fibre layer without. In the gut-fibre layer of the allantois there are large blood-vessels, which serve for the nutrition, and especially the respiration, of the embryo—the umbilical vessels (p. 170).In the reptiles and birds the allantois enlarges into a spacious sac, which encloses the embryo with the amnion, and does not combine with the outer fœtal membrane (the chorion). This is the case also with the lowest mammals, the oviparous Monotremes and most of the Marsupials. It is only in some of the later Marsupials (Peramelida) and all the Placentals that the allantois develops into the distinctive and remarkable structure that we call the placenta.
Fig.273. Foetal membranes of the human embryo (diagrammatic). Fig. 273—Fœtal membranes of the human embryo (diagrammatic). m the thick muscular wall of the womb. plu placenta [the inner layer (plu′) of which penetrates into the chorion-villi (chz) with its processes]. chf tufted, chl smooth chorion. a amnion, ah amniotic cavity, as amniotic sheath of the umbilical cord (which passes under into the navel of the embryo—not given here), dg vitelline duct, ds yelk sac, dv, dr decidua (vera and reflexa). The uterine cavity (uh) opens below into the vagina and above on the right into an oviduct (t). (From Kölliker.)
The placenta is formed by the branches of the blood-vessels in the wall of the allantois growing into the hollow ectodermic tufts (villi) of the chorion, which run into corresponding depressions in the mucous membrane of the womb. The latter also is richly permeated with blood-vessels which bring the mother’s blood to the embryo. As the partition in the villi between the maternal blood-vessels and those of the fœtus is extremely thin, there is a direct exchange of fluid between the two, and this is of the greatest importance in the nutrition of the young mammal. It is true that the maternal vessels do not entirely pass into the fœtal vessels, so that the two kinds of blood are simply mixed. But the partition between them is so thin that the nutritive fluid easily transudes through it. By means of this transudation or diosmosis the exchange of fluids takes place without difficulty. The larger the embryo is in the placentals, and the longer it remains in the womb, the more necessary it is to have special structures to meet its great consumption of food.
In this respect there is a very conspicuous difference between the lower and higher mammals. In the Marsupials, in which the embryo is only a comparatively short time in the womb and is born in a very immature condition, the vascular arrangements in the yelk-sac and the allantois suffice for its nutrition, as we find them in the Monotremes, birds, and reptiles. But in the Placentals, where gestation lasts a long time, and the embryo reaches its full development under the protection of its enveloping membranes, there has to be a new mechanism for the direct supply of a large quantity of food, and this is admirably met by the formation of the placenta.
Branches of the blood-vessels penetrate into the chorion-villi from within, starting from the gut-fibre layer of the allantois, and bringing the blood of the fœtus through the umbilical vessels (Fig. 273 chz). On the other hand, a thick network of blood-vessels develops in the mucous membrane that clothes the inner surface of the womb, especially in the region of the depressions into which the chorion-villi penetrate (plu). This network of arteries contains maternal blood, brought by the uterine vessels. As the connective tissue between the enlarged capillaries of the uterus disappears, wide cavities filled with maternal blood appear, and into these the chorion-villi of the embryo penetrate. The sum of these vessels of both kinds, that are so intimately correlated at this point, together with the connective and enveloping tissue, is the placenta. The placenta consists, therefore, properly speaking, of two different though intimately connected parts—the fœtal placenta (Fig. 273 chz) within and the maternal or uterine placenta (plu) without. The latter is made up of the mucous coat of the uterus and its blood-vessels, the former of the tufted chorion and the umbilical vessels of the embryo (cf. Fig. 196).
Fig.274. Skull of a fossil lemur (Adapis parisiensis,), from the Miocene at Quercy. Fig. 274—Skull of a fossil lemur (Adapis parisiensis), from the Miocene at Quercy. A lateral view from the right. B lower jaw, C lower molar, i incisors, c canines, p premolars, m molars.
The manner in which these two kinds of vessels combine in the placenta, and the structure, form, and size of it, differ a good deal in the various Placentals; to some extent they give us valuable data for the natural classification, and therefore the phylogeny, of the whole of this sub-class. On the ground of these differences we divide it into two principal sections; the lower Placentals or Indecidua, and the higher Placentals or Deciduata.
To the Indecidua belong three important groups of mammals: the Lemurs (Prosimiæ), the Ungulates (tapirs, horses, pigs, ruminants, etc.), and the Cetacea (dolphins and whales). In these Indecidua the villi are distributed over the whole surface of the chorion (or its greater part) either singly or in groups. They are only loosely connected with the mucous coat of the uterus, so that the whole fœtal membrane with its villi can be easily withdrawn from the uterine depressions like a hand from a glove. There is no real coalescence of the two placentas at any part of the surface of contact. Hence at birth the fœtal placenta alone comes away; the uterine placenta is not torn away with it.
The formation of the placenta is very different in the second and higher section of the Placentals, the Deciduata. Here again the whole surface of the chorion is thickly covered with the villi in the beginning. But they afterwards disappear from one part of the surface, and grow proportionately thicker on the other part. We thus get a differentiation between the smooth chorion (chorion laeve, Fig. 273 chl) and the thickly-tufted chorion (chorion frondosum, Fig. 273 chf). The former has only a few small villi or none at all; the latter is thickly covered with large and well-developed villi; this alone now constitutes the placenta. In the great majority of the Deciduata the placenta has the same shape as in man >(Figs. 197 and 200)—namely a thick, circular disk like a cake; so we find in the Insectivora, Chiroptera, Rodents, and Apes. This discoplacenta lies on one side of the chorion. But in the Sarcotheria (both the Carnivora and the seals, Pinnipedia) and in the elephant and several other Deciduates we find a zonoplacenta; in these the rich mass of villi runs like a girdle round the middle of the ellipsoid chorion, the two poles of it being free from them.
Still more characteristic of the Deciduates is the peculiar and very intimate connection between the chorion frondosum and the corresponding part of the mucous coat of the womb, which we must regard as a real coalescence of the two. The villi of the chorion push their branches into the blood-filled tissues of the coat of the uterus, and the vessels of each loop together so intimately that it is no longer possible to separate the fœtal from the maternal placenta; they form henceforth a compact and apparently simple placenta. In consequence of this coalescence, a whole piece of the lining of the womb comes away at birth with the fœtal membrane that is interlaced with it. This piece is called the “falling-away” membrane (decidua). It is also called the serous (spongy) membrane, because it is pierced like a sieve or sponge. All the higher Placentals that have this decidua are classed together as the “Deciduates.” The tearing away of the decidua at birth naturally causes the mother to lose a quantity of blood, which does not happen in the Indecidua. The last part of the uterine coat has to be repaired by a new growth after birth in the Deciduates. (Cf. Figs. 199, 200, pp. 168–70.)
Fig.275. The Slender Lori (Stenops gracilis) of Ceylon, a tail-less lemur. Fig. 275—The Slender Lori (Stenops gracilis) of Ceylon, a tail-less lemur.
In the various orders of the Deciduates, the placenta differs considerably both in outer form and internal structure. The extensive investigations of the last ten years have shown that there is more variation in these respects among the higher mammals than was formerly supposed. The physiological work of this important embryonic organ, the nutrition of the fœtus during its long sojourn in the womb, is accomplished in the various groups of the Placentals by very different and sometimes very elaborate structures. They have lately been fully described by Hans Strahl.
The phylogeny of the placenta has become more intelligible from the fact that we have found a number of transitional forms of it. Some of the Marsupials (Perameles) have the beginning of a placenta. In some of the Lemurs (Tarsius) a discoid placenta with decidua is developed.
While these important results of comparative embryology have been throwing further light on the close blood-relationship of man and the anthropoid apes in the last few years (p. 172), the great advance of paleontology has at the same time been affording us a deeper insight into the stem-history of the Placental group. In the seventh chapter of my Systematic Phylogeny of the Vertebrates I advanced the hypothesis that the Placentals form a single stem with many branches, which has been evolved from an older group of the Marsupials (Prodidelphia). The four great legions of the Placentals—Rodents, Ungulates, Carnassia, and Primates—are sharply separated to-day by important features of organisation. But if we consider their extinct ancestors of the Tertiary period, the differences gradually disappear, the deeper we go in the Cenozoic deposits; in the end we find that they vanish altogether. The primitive stem-forms of the Rodents (Esthonychida), the Ungulates (Chondylarthra), the Carnassia (Ictopsida), and the Primates (Lemuravida) are so closely related at the beginning of the Tertiary period that we might group them together as different families of one order, the Proplacentals (Mallotheria or Prochoriata).
Hence the great majority of the Placentals have no direct and close relationship to man, but only the legion of the Primates. This is now generally divided into three orders—the half-apes (Prosimiæ), apes (Simiæ), and man (Anthropi). The lemurs or half-apes are the stem-group, descending from the older Mallotheria of the Cretaceous period. From them the apes were evolved in the Tertiary period, and man was formed from these towards its close.
The Lemurs (Prosimiæ) have few living representatives. But they are very interesting, and are the last survivors of a once extensive group. We find many fossil remains of them in the older Tertiary deposits of Europe and North America, in the Eocene and Miocene. We distinguish two sub-orders, the fossil Lemuravida and the modern Lemurogona. The earliest and most primitive forms of the Lemuravida are the Pachylemurs (Hypopsodina); they come next to the earliest Placentals (Prochoriata), and have the typical full dentition, with forty-four teeth (3.1.4.3. over 3.1.4.3.). The Necrolemurs (Adapida, Fig. 274) have only forty teeth, and have lost an incisor in each jaw (2.1.4.3. over 2.1.4.3.). The dentition is still further reduced in the Lemurogona (Autolemures), which usually have only thirty-six teeth (2.1.3.3. over 2.1.3.3.). These living survivors are scattered far over the southern part of the Old World. Most of the species live in Madagascar, some in the Sunda Islands, others on the mainland of Asia and Africa. They are gloomy and melancholic animals; they live a quiet life, climbing trees, and eating fruit and insects. They are of different kinds. Some are closely related to the Marsupials (especially the opossum). Others (Macrotarsi) are nearer to the Insectivora, others again (Chiromys) to the Rodents. Some of the lemurs (Brachytarsi) approach closely to the true apes. The numerous fossil remains of half-apes and apes that have been recently found in the Tertiary deposits justify us in thinking that man’s ancestors were represented by several different species during this long period. Some of these were almost as big as men, such as the diluvial lemurogonon Megaladapis of Madagascar.
Fig.276. The white-nosed ape (Cercopithecus petaurista). Fig. 276—The white-nosed ape (Cercopithecus petaurista).
Next to the lemurs come the true apes (Simiæ), the twenty-sixth stage in our ancestry. It has been beyond question for some time now that the apes approach nearest to man in every respect of all the animals. Just as the lowest apes come close to the lemurs, so the highest come next to man. When we carefully study the comparative anatomy of the apes and man, we can trace a gradual and uninterrupted advance in the organisation of the ape up to the purely human frame, and, after impartial examination of the “ape problem” that has been discussed of late years with such passionate interest, we come infallibly to the important conclusion, first formulated by Huxley in 1863: “Whatever systems of organs we take, the comparison of their modifications in the series of apes leads to the same result: that the anatomic differences that separate man from the gorilla and chimpanzee are not as great as those that separate the gorilla from the lower apes.” Translated into phylogenetic language, this “pithecometra-law,” formulated in such masterly fashion by Huxley, is quite equivalent to the popular saying: “Man is descended from the apes.”
In the very first exposition of his profound natural classification (1735) Linné placed the anthropoid mammals at the head of the animal kingdom, with three genera: man, the ape, and the sloth. He afterwards called them the “Primates”—the “lords” of the animal world; he then also separated the lemur from the true ape, and rejected the sloth. Later zoologists divided the order of Primates. First the Gottingen anatomist, Blumenbach, founded a special order for man, which he called Bimana (“two-handed”); in a second order he united the apes and lemurs under the name of Quadrumana (“four-handed”); and a third order was formed of the distantly-related Chiroptera (bats, etc.). The separation of the Bimana and Quadrumana was retained by Cuvier and most of the subsequent zoologists. It seems to be extremely important, but, as a matter of fact, it is totally wrong. This was first shown in 1863 by Huxley, in his famous Man’s Place in Nature. On the strength of careful comparative anatomical research he proved that the apes are just as truly “two-handed” as man; or, if we prefer to reverse it, that man is as truly four-handed as the ape. He showed convincingly that the ideas of hand and foot had been wrongly defined, and had been improperly based on physiological instead of morphological grounds. The circumstance that we oppose the thumb to the other four fingers in our hand, and so can grasp things, seemed to be a special distinction of the hand in contrast to the foot, in which the corresponding great toe cannot be opposed in this way to the others. But the apes can grasp with the hind-foot as well as the fore, and so were regarded as quadrumanous. However, the inability to grasp that we find in the foot of civilised man is a consequence of the habit of clothing it with tight coverings for thousands of years. Many of the bare-footed lower races of men, especially among the negroes, use the foot very freely in the same way as the hand. As a result of early habit and continued practice, they can grasp with the foot (in climbing trees, for instance) just as well as with the hand. Even new-born infants of our own race can grasp very strongly with the great toe, and hold a spoon with it as firmly as with the hand. Hence the physiological distinction between hand and foot can neither be pressed very far, nor has it a scientific basis. We must look to morphological characters.
Fig.277. The drill-baboon (Cynocephalus leucophaeus) (From Brehm.) Fig. 277—The drill-baboon (Cynocephalus leucophæus). (From Brehm.)
As a matter of fact, it is possible to draw such a sharp morphological distinction—a distinction based on anatomic structure—between the fore and hind extremity. In the formation both of the bony skeleton and of the muscles that are connected with the hand and foot before and behind there are material and constant differences; and these are found both in man and the ape. For instance, the number and arrangement of the smaller bones of the hand and foot are quite different. There are similar constant differences in the muscles. The hind extremity always has three muscles (a short flexor muscle, a short extensor muscle, and a long calf-muscle) that are not found in the fore extremity. The arrangement of the muscles also is different before and behind. These characteristic differences between the fore and hind extremities are found in man as well as the ape. There can be no doubt, therefore, that the ape’s foot deserves that name just as much as the human foot does, and that all true apes are just as “bimanous” as man. The common distinction of the apes as “quadrumanous” is altogether wrong morphologically.
But it may be asked whether, quite apart from this, we can find any other features that distinguish man more sharply from the ape than the various species of apes are distinguished from each other. Huxley gave so complete and demonstrative a reply to this question that the opposition still raised on many sides is absolutely without foundation. On the ground of careful comparative anatomical research, Huxley proved that in all morphological respects the differences between the highest and lowest apes are greater than the corresponding differences between the highest apes and man. He thus restored Linné’s order of the Primates (excluding the bats), and divided it into three sub-orders, the first composed of the half-apes (Lemuridæ), the second of the true apes (Simiadæ), the third of men (Anthropidæ).
But, as we wish to proceed quite consistently and impartially on the laws of systematic logic, we may, on the strength of Huxley’s own law, go a good deal farther in this division. We are justified in going at least one important step farther, and assigning man his natural place within one of the sections of the order of apes. All the features that characterise this group of apes are found in man, and not found in the other apes. We do not seem to be justified, therefore, in founding for man a special order distinct from the apes.
The order of the true apes (Simiæ or Pitheca)—excluding the lemurs—has long been divided into two principal groups, which also differ in their geographical distribution. One group (Hesperopitheca, or western apes) live in America. The other group, to which man belongs, are the Eopitheca or eastern apes; they are found in Asia and Africa, and were formerly in Europe. All the eastern apes agree with man in the features that are chiefly used in zoological classification to distinguish between the two simian groups, especially in the dentition. The objection might be raised that the teeth are too subordinate an organ physiologically for us to lay stress on them in so important a question. But there is a good reason for it; it is with perfect justice that zoologists have for more than a century paid particular attention to the teeth in the systematic division and arrangement of the orders of mammals. The number, form, and arrangement of the teeth are much more faithfully inherited in the various orders than most other characters.
Hence the form of dentition in man is very important. In the fully developed condition we have thirty-two teeth; of these eight are incisors, four canine, and twenty molars. The eight incisors, in the middle of the jaws, have certain characteristic differences above and below. In the upper jaw the inner incisors are larger than the outer; in the lower jaw the inner are the smaller. Next to these, at each side of both jaws, is a canine (or “eye tooth”), which is larger than the incisors. Sometimes it is very prominent in man, as it is in most apes and many of the other mammals, and forms a sort of tusk. Next to this there are five molars above and below on each side, the first two of which (the “pre-molars”) are small, have only one root, and are included in the change of teeth; the three back ones are much larger, have two roots, and only come with the second teeth. The apes of the Old World, or all the living or fossil apes of Asia, Africa, and Europe, have the same dentition as man.
Figs. 278 to 282. Skeletons of a man and the four anthropoid apes. Fig. 278. Gibbon (Hylobates). Fig. 279. Orang (Satyrus). Fig. 280. Chimpanzee (Anthropithecus). Fig. 281. Gorilla (Gorilla). Fig. 282. Man (Homo). Fig. 278–282—Skeletons of a man and the four anthropoid apes. (Fig. 278, Gibbon; Fig. 279, Orang; Fig. 280, Chimpanzee; Fig. 281, Gorilla; Fig. 282, Man. (From Huxley.) Cf. Figs. 203–209.
On the other hand, all the American apes have an additional pre-molar in each half of the jaw. They have six molars above and below on each side, or thirty-six teeth altogether. This characteristic difference between the eastern and western apes has been so faithfully inherited that it is very instructive for us. It is true that there seems to be an exception in the case of a small family of South American apes. The small silky apes (Arctopitheca or Hapalidæ), which include the tamarin (Midas) and the brush-monkey (Jacchus), have only five molars in each half of the jaw (instead of six), and so seem to be nearer to the eastern apes. But it is found, on closer examination, that they have three premolars, like all the western apes, and that only the last molar has been lost. Hence the apparent exception really confirms the above distinction.
Of the other features in which the two groups of apes differ, the structure of the nose is particularly instructive and conspicuous. All the eastern apes have the same type of nose as man—a comparatively narrow partition between the two halves, so that the nostrils run downwards. In some of them the nose protrudes as far as in man, and has the same characteristic structure. We have already alluded to the curious long-nosed apes, which have a long, finely-curved nose. Most of the eastern apes have, it is true, rather flat noses, like, for instance, the white-nosed monkey (Fig. 276); but the nasal partition is thin and narrow in them all. The American apes have a different type of nose. The partition is very broad and thick at the bottom, and the wings of the nostrils are not developed, so that they point outwards instead of downwards. This difference in the form of the nose is so constantly inherited in both groups that the apes of the New World are called “flat-nosed” (Platyrrhinæ), and those of the Old World “narrow-nosed” (Catarrhinæ). The bony passage of the ear (at the bottom of which is the tympanum) is short and wide in all the Platyrrhines, but long and narrow in all the Catarrhines; and in man this difference also is significant.
This division of the apes into Platyrrhines and Catarrhines, on the ground of the above hereditary features, is now generally admitted in zoology, and receives strong support from the geographical distribution of the two groups in the east and west. It follows at once, as regards the phylogeny of the apes, that two divergent lines proceeded from the common stem-form of the ape-order in the early Tertiary period, one of which spread over the Old, the other over the New, World. It is certain that all the Platyrrhines come of one stock, and also all the Catarrhines; but the former are phylogenetically older, and must be regarded as the stem-group of the latter.
What can we deduce from this with regard to our own genealogy? Man has just the same characters, the same form of dentition, auditory passage, and nose, as all the Catarrhines; in this he radically differs from the Platyrrhines. We are thus forced to assign him a position among the eastern apes in the order of Primates, or at least place him alongside of them. But it follows that man is a direct blood relative of the apes of the Old World, and can be traced to a common stem-form together with all the Catarrhines. In his whole organisation and in his origin man is a true Catarrhine; he originated in the Old World from an unknown, extinct group of the eastern apes. The apes of the New World, or the Platyrrhines, form a divergent branch of our genealogical tree, and this is only distantly related at its root to the human race. We must assume, of course, that the earliest Eocene apes had the full dentition of the Platyrrhines; hence we may regard this stem-group as a special stage (the twenty-sixth) in our ancestry, and deduce from it (as the twenty-seventh stage) the earliest Catarrhines.
We have now reduced the circle of our nearest relatives to the small and comparatively scanty group that is represented by the sub-order of the Catarrhines; and we are in a position to answer the question of man’s place in this sub-order, and say whether we can deduce anything further from this position as to our immediate ancestors. In answering this question the comprehensive and able studies that Huxley gives of the comparative anatomy of man and the various Catarrhines in his Man’s Place in Nature are of great assistance to us. It is quite clear from these that the differences between man and the highest Catarrhines (gorilla, chimpanzee, and orang) are in every respect slighter than the corresponding differences between the highest and the lowest Catarrhines (white-nosed monkey, macaco, baboon, etc.). In fact, within the small group of the tail-less anthropoid apes the differences between the various genera are not less than the differences between them and man. This is seen by a glance at the skeletons that Huxley has put together (Figs. 278–282). Whether we take the skull or the vertebral column or the ribs or the fore or hind limbs, or whether we extend the comparison to the muscles, blood-vessels, brain, placenta, etc., we always reach the same result on impartial examination—that man is not more different from the other Catarrhines than the extreme forms of them (for instance, the gorilla and baboon) differ from each other. We may now, therefore, complete the Huxleian law we have already quoted with the following thesis: “Whatever system of organs we take, a comparison of their modifications in the series of Catarrhines always leads to the same conclusion; the anatomic differences that separate man from the most advanced Catarrhines (orang, gorilla, chimpanzee) are not as great as those that separate the latter from the lowest Catarrhines (white-nosed monkey, macaco, baboon).”
We must, therefore, consider the descent of man from other Catarrhines to be fully proved. Whatever further information on the comparative anatomy and ontogeny of the living Catarrhines we may obtain in the future, it cannot possibly disturb this conclusion. Naturally, our Catarrhine ancestors must have passed through a long series of different forms before the human type was produced. The chief advances that effected this “creation of man,” or his differentiation from the nearest related Catarrhines, were: the adoption of the erect posture and the consequent greater differentiation of the fore and hind limbs, the evolution of articulate speech and its organ, the larynx, and the further development of the brain and its function, the soul; sexual selection had a great influence in this, as Darwin showed in his famous work.
With an eye to these advances we can distinguish at least four important stages in our simian ancestry, which represent prominent points in the historical process of the making of man. We may take, after the Lemurs, the earliest and lowest Platyrrhines of South America, with thirty-six teeth, as the twenty-sixth stage of our genealogy; they were developed from the Lemurs by a peculiar modification of the brain, teeth, nose, and fingers. From these Eocene stem-apes were formed the earliest Catarrhines or eastern apes, with the human dentition (thirty-two teeth), by modification of the nose, lengthening of the bony channel of the ear, and the loss of four pre-molars. These oldest stem-forms of the whole Catarrhine group were still thickly coated with hair, and had long tails—baboons (Cynopitheca) or tailed apes (Menocerca, Fig. 276). They lived during the Tertiary period, and are found fossilised in the Miocene. Of the actual tailed apes perhaps the nearest to them are the Semnopitheci.
If we take these Semnopitheci as the twenty-seventh stage in our ancestry, we may put next to them, as the twenty-eighth, the tail-less anthropoid apes. This name is given to the most advanced and man-like of the existing Catarrhines. They were developed from the other Catarrhines by losing the tail and part of the hair, and by a higher development of the brain, which found expression in the enormous growth of the skull. Of this remarkable family there are only a few genera to-day, and we have already dealt with them (Chapter XV)—the gibbon (Hylobates, Fig. 203) and orang (Satyrus, Figs. 204, 205) in South-Eastern Asia and the Archipelago; and the chimpanzee (Anthropithecus, Figs. 206, 207) and gorilla (Gorilla, Fig. 208) in Equatorial Africa.
The great interest that every thoughtful man takes in these nearest relatives of ours has found expression recently in a fairly large literature. The most distinguished of these works for impartial treatment of the question of affinity is Robert Hartmann’s little work on The Anthropoid Apes. Hartmann divides the primate order into two families: (1) Primarii (man and the anthropoid apes); and (2) Simianæ (true apes, Catarrhines and Platyrrhines). Professor Klaatsch, of Heidelberg, has advanced a different view in his interesting and richly illustrated work on The Origin and Development of the Human Race. This is a substantial supplement to my Anthropogeny, in so far as it gives the chief results of modern research on the early history of man and civilisation. But when Klaatsch declares the descent of man from the apes to be “irrational, narrow-minded, and false,” in the belief that we are thinking of some living species of ape, we must remind him that no competent scientist has ever held so narrow a view. All of us look merely—in the sense of Lamarck and Darwin—to the original unity (admitted by Klaatsch) of the primate stem. This common descent of all the Primates (men, apes, and lemurs) from one primitive stem-form, from which the most far-reaching conclusions follow for the whole of anthropology and philosophy, is admitted by Klaatsch as well as by myself and all other competent zoologists who accept the theory of evolution in general. He says explicitly (p. 172): “The three anthropoid apes—gorilla, chimpanzee, and orang—seem to be branches from a common root, and this was not far from that of the gibbon and man.” That is in the main the opinion that I have maintained (especially against Virchow) in a number of works ever since 1866. The hypothetical common ancestor of all the Primates, which must have lived in the earliest Tertiary period (more probably in the Cretaceous), was called by me Archiprimus; Klaatsch now calls it Primatoid. Dubois has proposed the appropriate name of Prothylobates for the common and much younger stem-form of the anthropomorpha (man and the anthropoid apes). The actual Hylobates is nearer to it than the other three existing anthropoids. None of these can be said to be absolutely the most man-like. The gorilla comes next to man in the structure of the hand and foot, the chimpanzee in the chief features of the skull, the orang in brain development, and the gibbon in the formation of the chest. None of these existing anthropoid apes is among the direct ancestors of our race; they are scattered survivors of an ancient branch of the Catarrhines, from which the human race developed in a particular direction.
Fig.283. Skull of the fossil ape-man of Java (Pithecanthropus erectus), restored by Eugen Dubois. Fig. 283—Skull of the fossil ape-man of Java (Pithecanthropus erectus), restored by Eugen Dubois.
Although man is directly connected with this anthropoid family and originates from it, we may assign an important intermediate form between the Prothylobates and him (the twenty-ninth stage in our ancestry), the ape-men (Pithecanthropi). I gave this name in the History of Creation to the “speechless primitive men” (Alali), which were men in the ordinary sense as far as the general structure is concerned (especially in the differentiation of the limbs), but lacked one of the chief human characteristics, articulate speech and the higher intelligence that goes with it, and so had a less developed brain. The phylogenetic hypothesis of the organisation of this “ape-man” which I then advanced was brilliantly confirmed twenty-four years afterwards by the famous discovery of the fossil Pithecanthropus erectus by Eugen Dubois (then military surgeon in Java, afterwards professor at Amsterdam). In 1892 he found at Trinil, in the residency of Madiun in Java, in Pliocene deposits, certain remains of a large and very man-like ape (roof of the skull, femur, and teeth), which he described as “an erect ape-man” and a survivor of a “stem-form of man” (Fig. 283). Naturally, the Pithecanthropus excited the liveliest interest, as the long-sought transitional form between man and the ape: we seemed to have found “the missing link.” There were very interesting scientific discussions of it at the last three International Congresses of Zoology (Leyden, 1895, Cambridge, 1898, and Berlin, 1901). I took an active part in the discussion at Cambridge, and may refer the reader to the paper I read there on “The Present Position of Our Knowledge of the Origin of Man” (translated by Dr. Gadow with the title of The Last Link).
An extensive and valuable literature has grown up in the last ten years on the Pithecanthropus and the pithecoid theory connected with it. A number of distinguished anthropologists, anatomists, paleontologists, and phylogenists have taken part in the controversy, and made use of the important data furnished by the new science of pre-historic research. Hermann Klaatsch has given a good summary of them, with many fine illustrations, in the above-mentioned work. I refer the reader to it as a valuable supplement to the present work, especially as I cannot go any further here into these anthropological and pre-historic questions. I will only repeat that I think he is wrong in the attitude of hostility that he affects to take up with regard to my own views on the descent of man from the apes.
The most powerful opponent of the pithecoid theory—and the theory of evolution in general—during the last thirty years (until his death in September, 1902) was the famous Berlin anatomist, Rudolf Virchow. In the speeches which he delivered every year at various congresses and meetings on this question, he was never tired of attacking the hated “ape theory.” His constant categorical position was: “It is quite certain that man does not descend from the ape or any other animal.” This has been repeated incessantly by opponents of the theory, especially theologians and philosophers. In the inaugural speech that he delivered in 1894 at the Anthropological Congress at Vienna, he said that “man might just as well have descended from a sheep or an elephant as from an ape.” Absurd expressions like this only show that the famous pathological anatomist, who did so much for medicine in the establishment of cellular pathology, had not the requisite attainments in comparative anatomy and ontogeny, systematic zoology and paleontology, for sound judgment in the province of anthropology. The Strassburg anatomist, Gustav Schwalbe, deserved great praise for having the moral courage to oppose this dogmatic and ungrounded teaching of Virchow, and showing its untenability. The recent admirable works of Schwalbe on the Pithecanthropus, the earliest races of men, and the Neanderthal skull (1897–1901) will supply any candid and judicious reader with the empirical material with which he can convince himself of the baselessness of the erroneous dogmas of Virchow and his clerical friends (J. Ranke, J. Bumüller, etc.).
As the Pithecanthropus walked erect, and his brain (judging from the capacity of his skull, Fig. 283) was midway between the lowest men and the anthropoid apes, we must assume that the next great step in the advance from the Pithecanthropus to man was the further development of human speech and reason.
Comparative philology has recently shown that human speech is polyphyletic in origin; that we must distinguish several (probably many) different primitive tongues that were developed independently. The evolution of language also teaches us (both from its ontogeny in the child and its phylogeny in the race) that human speech proper was only gradually developed after the rest of the body had attained its characteristic form. It is probable that language was not evolved until after the dispersal of the various species and races of men, and this probably took place at the commencement of the Quaternary or Diluvial period. The speechless ape-men or Alali certainly existed towards the end of the Tertiary period, during the Pliocene, possibly even the Miocene, period.
The third, and last, stage of our animal ancestry is the true or speaking man (Homo), who was gradually evolved from the preceding stage by the advance of animal language into articulate human speech. As to the time and place of this real “creation of man” we can only express tentative opinions. It was probably during the Diluvial period in the hotter zone of the Old World, either on the mainland in tropical Africa or Asia or on an earlier continent (Lemuria—now sunk below the waves of the Indian Ocean), which stretched from East Africa (Madagascar, Abyssinia) to East Asia (Sunda Islands, Further India). I have given fully in my History of Creation, (chapter xxviii) the weighty reasons for claiming this descent of man from the anthropoid eastern apes, and shown how we may conceive the spread of the various races from this “Paradise” over the whole earth. I have also dealt fully with the relations of the various races and species of men to each other.
SYNOPSIS OF THE CHIEF SECTIONS OF OUR STEM-HISTORY
First Stage: The Protists
Man’s ancestors are unicellular protozoa, originally unnucleated Monera like the Chromacea, structureless green particles of plasm; afterwards real nucleated cells (first plasmodomous Protophyta, like the Palmella; then plasmophagous Protozoa, like the Amœba).
Second Stage: The Blastæads
Man’s ancestors are round cœnobia or colonies of Protozoa; they consist of a close association of many homogeneous cells, and thus are individuals of the second order. They resemble the round cell-communities of the Magospheræ and Volvocina, equivalent to the ontogenetic blastula: hollow globules, the wall of which consists of a single layer of ciliated cells (blastoderm).
Third Stage: The Gastræads
Man’s ancestors are Gastræads, like the simplest of the actual Metazoa (Prophysema, Olynthus, Hydra, Pemmatodiscus). Their body consists merely of a primitive gut, the wall of which is made up of the two primary germinal layers.
Fourth Stage: The Platodes
Man’s ancestors have substantially the organisation of simple Platodes (at first like the cryptocœlic Platodaria, later like the rhabdocœlic Turbellaria). The leaf-shaped bilateral-symmetrical body has only one gut-opening, and develops the first trace of a nervous centre from the ectoderm in the middle line of the back (Figs. 239, 240).
Fifth Stage: The Vermalia
Man’s ancestors have substantially the organisation of unarticulated Vermalia, at first Gastrotricha (Ichthydina), afterwards Frontonia (Nemertina, Enteropneusta). Four secondary germinal layers develop, two middle layers arising between the limiting layers (cœloma). The dorsal ectoderm forms the vertical plate, acroganglion (Fig. 243).
Sixth Stage: The Prochordonia
Man’s ancestors have substantially the organisation of a simple unarticulated Chordonium (Copelata and Ascidia-larvæ). The unsegmented chorda develops between the dorsal medullary tube and the ventral gut-tube. The simple cœlom-pouches divide by a frontal septum into two on each side; the dorsal pouch (episomite) forms a muscle-plate; the ventral pouch (hyposomite) forms a gonad. Head-gut with gill-clefts.
Seventh Stage: The Acrania
Man’s ancestors are skull-less Vertebrates, like the Amphioxus. The body is a series of metamera, as several of the primitive segments are developed. The head contains in the ventral half the branchial gut, the trunk the hepatic gut. The medullary tube is still simple. No skull, jaws, or limbs.
Eighth Stage: The Cyclostoma
Man’s ancestors are jaw-less Craniotes (like the Myxinoida and Petromyzonta). The number of metamera increases. The fore-end of the medullary tube expands into a vesicle and forms the brain, which soon divides into five cerebral vesicles. In the sides of it appear the three higher sense-organs: nose, eyes, and auditory vesicles. No jaws, limbs, or floating bladder.
Ninth Stage: The Ichthyoda
Man’s ancestors are fish-like Craniotes: (1) Primitive fishes (Selachii); (2) plated fishes (Ganoida); (3) amphibian fishes (Dipneusta); (4) mailed amphibia (Stegocephala). The ancestors of this series develop two pairs of limbs: a pair of fore (breast-fins) and of hind (belly-fins) legs. The gill-arches are formed between the gill-clefts: the first pair form the maxillary arches (the upper and lower jaws). The floating bladder (lung) and pancreas grow out of the gut.
Tenth Stage: The Amniotes
Man’s ancestors are Amniotes or gill-less Vertebrates: (1) Primitive Amniotes (Proreptilia); (2) Sauromammals; (3) Primitive Mammals (Monotremes); (4) Marsupials; (5) Lemurs (Prosimiæ); (6) Western apes (Platyrrhinæ); (7) Eastern apes (Catarrhinæ): at first tailed Cynopitheca; then tail-less anthropoids; later speechless ape-men (Alali); finally speaking man. The ancestors of these Amniotes develop an amnion and allantois, and gradually assume the mammal, and finally the specifically human, form.
XXIV. EVOLUTION OF THE NERVOUS SYSTEM
The previous chapters have taught us how the human body as a whole develops from the first simple rudiment, a single layer of cells. The whole human race owes its origin, like the individual man, to a simple cell. The unicellular stem-form of the race is reproduced daily in the unicellular embryonic stage of the individual. We have now to consider in detail the evolution of the various parts that make up the human frame. I must, naturally, confine myself to the most general and principal outlines; to make a special study of the evolution of each organ and tissue is both beyond the scope of this work, and probably beyond the anatomic capacity of most of my readers to appreciate. In tracing the evolution of the various organs we shall follow the method that has hitherto guided us, except that we shall now have to consider the ontogeny and phylogeny of the organs together. We have seen, in studying the evolution of the body as a whole, that phylogeny casts a light over the darker paths of ontogeny, and that we should be almost unable to find our way in it without the aid of the former. We shall have the same experience in the study of the organs in detail, and I shall be compelled to give simultaneously their ontogenetic and phylogenetic origin. The more we go into the details of organic development, and the more closely we follow the rise of the various parts, the more we see the inseparable connection of embryology and stem-history. The ontogeny of the organs can only be understood in the light of their phylogeny, just as we found of the embryology of the whole body. Each embryonic form is determined by a corresponding stem-form. This is true of details as well as of the whole.
We will consider first the animal and then the vegetal systems of organs of the body. The first group consists of the psychic and the motor apparatus. To the former belong the skin, the nervous system, and the sense-organs. The motor apparatus is composed of the passive and the active organs of movement (the skeleton and the muscles). The second or vegetal group consists of the nutritive and the reproductive apparatus. To the nutritive apparatus belong the alimentary canal with all its appendages, the vascular system, and the renal (kidney) system. The reproductive apparatus comprises the different organs of sex (embryonic glands, sexual ducts, and copulative organs).
As we know from previous chapters (XI–XIII), the animal systems of organs (the organs of sensation and presentation) develop for the most part out of the outer primary germ-layer, or the cutaneous (skin) layer. On the other hand, the vegetal systems of organs arise for the most part from the inner primary germ-layer, the visceral layer. It is true that this antithesis of the animal and vegetal spheres of the body in man and all the higher animals is by no means rigid; several parts of the animal apparatus (for instance, the greater part of the muscles) are formed from cells that come originally from the entoderm; and a great part of the vegetative apparatus (for instance, the mouth-cavity and the gonoducts) are composed of cells that come from the ectoderm.
In the more advanced animal body there is so much interlacing and displacement of the various parts that it is often very difficult to indicate the sources of them. But, broadly speaking, we may take it as a positive and important fact that in man and the higher animals the chief part of the animal organs comes from the ectoderm, and the greater part of the vegetative organs from the entoderm. It was for this reason that Carl Ernst von Baer called the one the animal and the other the vegetative layer (see p. 16).
The solid foundation of this important thesis is the gastrula, the most instructive embryonic form in the animal world, which we still find in the same shape in the most diverse classes of animals. This form points demonstrably to a common stem-form of all the Metazoa, the Gastræa; in this long-extinct stem-form the whole body consisted throughout life of the two primary germinal layers, as is now the case temporarily in the gastrula; in the Gastræa the simple cutaneous (skin) layer actually represented all the animal organs and functions, and the simple visceral (gut) layer all the vegetal organs and functions. This is the case with the modern Gastræads (Fig. 233); and it is also the case potentially with the gastrula.
We shall easily see that the gastræa theory is thus able to throw a good deal of light, both morphologically and physiologically, on some of the chief features of embryonic development, if we take up first the consideration of the chief element in the animal sphere, the psychic apparatus or sensorium and its evolution. This apparatus consists of two very different parts, which seem at first to have very little connection with each other—the outer skin, with all its hairs, nails, sweat-glands, etc., and the nervous system. The latter comprises the central nervous system (brain and spinal cord), the peripheral, cerebral, and spinal nerves, and the sense-organs. In the fully-formed vertebrate body these two chief elements of the sensorium lie far apart, the skin being external to, and the central nervous system in the very centre of, the body. The one is only connected with the other by a section of the peripheral nervous system and the sense-organs. Nevertheless, as we know from human embryology, the medullary tube is formed from the cutaneous layer. The organs that discharge the most advanced functions of the animal body—the organs of the soul, or of psychic life—develop from the external skin. This is a perfectly natural and necessary process. If we reflect on the historical evolution of the psychic and sensory functions, we are forced to conclude that the cells which accomplish them must originally have been located on the outer surface of the body. Only elementary organs in this superficial position could directly receive the influences of the environment. Afterwards, under the influence of natural selection, the cellular group in the skin which was specifically “sensitive” withdrew into the inner and more protected part of the body, and formed there the foundation of a central nervous organ. As a result of increased differentiation, the skin and the central nervous system became further and further separated, and in the end the two were only permanently connected by the afferent peripheral sensory nerves.
Fig.244. The human skin in vertical section. Fig. 284—The human skin in vertical section (from Ecker), highly magnified, a horny layer of the epidermis, b mucous layer of the epidermis, c papillæ of the corium, d blood-vessels of same, ef ducts of the sweat-glands (g), h fat-glands in the corium, i nerve, passing into a tactile corpuscle above.
The observations of the comparative anatomist are in complete accord with this view. He tells us that large numbers of the lower animals have no nervous system, though they exercise the functions of sensation and will like the higher animals. In the unicellular Protozoa, which do not form germinal layers, there is, of course, neither nervous system nor skin. But in the second division of the animal kingdom also, the Metazoa, there is at first no nervous system. Its functions are represented by the simple cell-layer of the ectoderm, which the lower Metazoa have inherited from the Gastræa (Fig. 30 e). We find this in the lowest Zoophytes—the Gastræads, Physemaria, and Sponges (Figs. 233–238). The lowest Cnidaria (the hydroid polyps) also are little superior to the Gastræads in structure. Their vegetative functions are accomplished by the simple visceral layer, and their animal functions by the simple cutaneous layer. In these cases the simple cell-layer of the ectoderm is at once skin, locomotive apparatus, and nervous system.
Fig.285. Epidermic cells of a human embryo of two months. Fig. 285—Epidermic cells of a human embryo of two months. (From Kölliker.)
When we come to the higher Metazoa, in which the sensory functions and their organs are more advanced, we find a division of labour among the ectodermic cells. Groups of sensitive nerve cells separate from the ordinary epidermic cells; they retire into the more protected tissue of the mesodermic under-skin, and form special neural ganglia there. Even in the Platodes, especially the Turbellaria, we find an independent nervous system, which has separated from the outer skin. This is the “upper pharyngeal ganglion,” or acroganglion, situated above the gullet (Fig. 241 g).From this rudimentary structure has been developed the elaborate central nervous system of the higher animals. In some of the higher worms, such as the earth-worm, the first rudiment of the central nervous system (Fig. 74 n) is a local thickening of the skin-sense layer (hs), which afterwards separates altogether from the horny plate. In the earliest Platodes (Cryptocœla) and Vermalia (Gastrotricha) the acroganglion remains in the epidermis. But the medullary tube of the Vertebrates originates in the same way. Our embryology has taught us that this first structure of the central nervous system also develops originally from the outer germinal layer.
Let us now examine more closely the evolution of the human skin, with its various appendages, the hairs and glands. This external covering has, physiologically, a double and important part to play. It is, in the first place, the common integument that covers the whole surface of the body, and forms a protective envelope for the other organs. As such it also effects a certain exchange of matter between the body and the surrounding atmosphere (exhalation, perspiration). In the second place, it is the earliest and original sense organ, the common organ of feeling that experiences the sensation of the temperature of the environment and the pressure or resistance of bodies that come into contact.
The human skin (like that of all the higher animals) is composed of two layers, the outer and the inner or underlying skin. The outer skin or epidermis, consists of simple ectodermic cells, and contains no blood-vessels (Fig. 284 a, b). It develops from the outer germinal layer, or skin-sense layer. The underlying skin (corium or hypodermis) consists chiefly of connective tissue, contains numerous blood-vessels and nerves, and has a totally different origin. It comes from the outermost parietal stratum of the middle germinal layer, or the skin-fibre layer. The corium is much thicker than the epidermis. In its deeper strata (the subcutis) there are clusters of fat-cells (Fig. 284 h). Its uppermost stratum (the cutis proper, or the papillary stratum) forms, over almost the whole surface of the body, a number of conical microscopic papillæ (something like warts), which push into the overlying epidermis (c). These tactile or sensory particles contain the finest sensory organs of the skin, the touch corpuscles. Others contain merely end-loops of the blood-vessels that nourish the skin (c, d). The various parts of the corium arise by division of labour from the originally homogeneous cells of the cutis-plate, the outermost lamina of the mesodermic skin-fibre layer (Fig. 145 hpr, and Figs. 161, 162 cp).
In the same way, all the parts and appendages of the epidermis develop by differentiation from the homogeneous cells of this horny plate (Fig. 285). At an early stage the simple cellular layer of this horny plate divides into two. The inner and softer stratum (Fig. 284 b) is known as the mucous stratum, the outer and harder (a) as the horny (corneous) stratum. This horny layer is being constantly used up and rubbed away at the surface; new layers of cells grow up in their place out of the underlying mucous stratum. At first the epidermis is a simple covering of the surface of the body. Afterwards various appendages develop from it, some internally, others externally. The internal appendages are the cutaneous glands—sweat, fat, etc. The external appendages are the hairs and nails.
The cutaneous glands are originally merely solid cone-shaped growths of the epidermis, which sink into the underlying corium (Fig. 286 1). Afterwards a canal (2, 3) is formed inside them, either by the softening and dissolution of the central cells or by the secretion of fluid internally. Some of the glands, such as the sudoriferous, do not ramify (Fig. 284 efg). These glands, which secrete the perspiration, are very long, and have a spiral coil at the end, but they never ramify; so also the wax-glands of the ears. Most of the other cutaneous glands give out buds and ramify; thus, for instance, the lachrymal glands of the upper eye-lid that secrete tears (Fig. 286), and the sebaceous glands which secrete the fat in the skin and generally open into the hair-follicles. Sudoriferous and sebaceous glands are found only in mammals. But we find lachrymal glands in all the three classes of Amniotes—reptiles, birds, and mammals. They are wanting in the lower aquatic vertebrates.
Fig.286. Rudimentary lachrymal glands from a human embryo of four months. Fig. 286—Rudimentary lachrymal glands from a human embryo of four months. (From Kölliker.) 1 earliest structure, in the shape of a simple solid cone, 2 and 3 more advanced structures, ramifying and hollowing out. a solid buds, e cellular coat of the hollow buds, f structure of the fibrous envelope, which afterwards forms the corium about the glands.
The mammary glands (Figs. 287, 288) are very remarkable; they are found in all mammals, and in these alone. They secrete the milk for the feeding of the new-born mammal. In spite of their unusual size these structures are nothing more than large sebaceous glands in the skin. The milk is formed by the liquefaction of the fatty milk-cells inside the branching mammary-gland tubes (Fig. 287 c), in the same way as the skin-grease or hair-fat, by the solution of fatty cells inside the sebaceous glands. The outlets of the mammary glands enlarge and form sac-like mammary ducts (b); these narrow again (a), and open in the teats or nipples of the breast by sixteen to twenty-four fine apertures. The first structure of this large and elaborate gland is a very simple cone in the epidermis, which penetrates into the corium and ramifies. In the new-born infant it consists of twelve to eighteen radiating lobes (Fig. 288). These gradually ramify, their ducts become hollow and larger, and rich masses of fat accumulate between the lobes. Thus is formed the prominent female breast (mamma), on the top of which rises the teat or nipple (mammilla). The latter is only developed later on, when the mammary gland is fully-formed; and this ontogenetic phenomenon is extremely interesting, because the earlier mammals (the stem-forms of the whole class) have no teats. In them the milk comes out through a flat portion of the ventral skin that is pierced like a sieve, as we still find in the lowest living mammals, the oviparous Monotremes of Australia. The young animal licks the milk from the mother instead of sucking it. In many of the lower mammals we find a number of milk-glands at different parts of the ventral surface. In the human female there is usually only one pair of glands, at the breast; and it is the same with the apes, bats, elephants, and several other mammals. Sometimes, however, we find two successive pairs of glands (or even more) in the human female. Some women have four or five pairs of breasts, like pigs and hedgehogs (Fig. 103). This polymastism points back to an older stem-form. We often find these accessory breasts in the male also (Fig. 103 D). Sometimes, moreover, the normal mammary glands are fully developed and can suckle in the male; but as a rule they are merely rudimentary organs without functions in the male. We have already (Chapter XI) dealt with this remarkable and interesting instance of atavism.
While the cutaneous glands are inner growths of the epidermis, the appendages which we call hairs and nails are external local growths in it. The nails (Ungues) which form important protective structures on the back of the most sensitive parts of our limbs, the tips of the fingers and toes, are horny growths of the epidermis, which we share with the apes. The lower mammals usually have claws instead of them; the ungulates, hoofs. The stem-form of the mammals certainly had claws; we find them in a rudimentary form even in the salamander. The horny claws are highly developed in most of the reptiles (Fig. 264), and the mammals have inherited them from the earliest representatives of this class, the stem-reptiles (Tocosauria). Like the hoofs (ungulæ) of the Ungulates, the nails of apes and men have been evolved from the claws of the older mammals. In the human embryo the first rudiment of the nails is found (between the horny and the mucous stratum of the epidermis) in the fourth month. But their edges do not penetrate through until the end of the sixth month.
Fig.287. The female breast (mamma) in vertical section. Fig. 287—The female breast (mamma) in vertical section. c racemose glandular lobes, b enlarged milk-ducts, a narrower outlets, which open into the nipple. (From H. Meyer.)
The most interesting and important appendages of the epidermis are the hairs; on account of their peculiar composition and origin we must regard them as highly characteristic of the whole mammalian class. It is true that we also find hairs in many of the lower animals, such as insects and worms. But these hairs, like the hairs of plants, are thread-like appendages of the surface, and differ entirely from the hairs of the mammals in the details of their structure and development.
The embryology of the hairs is known in all its details, but there are two different views as to their phylogeny. On the older view the hairs of the mammals are equivalent or homologous to the feathers of the bird or the horny scales of the reptile. As we deduce all three classes of Amniotes from a common stem-group, we must assume that these Permian stem-reptiles had a complete scaly coat, inherited from their Carboniferous ancestors, the mailed amphibia (Stegocephala); the bony scales of their corium were covered with horny scales. In passing from aquatic to terrestrial life the horny scales were further developed, and the bony scales degenerated in most of the reptiles. As regards the bird’s feathers, it is certain that they are modifications of the horny scales of their reptilian ancestors. But it is otherwise with the hairs of the mammals. In their case the hypothesis has lately been advanced on the strength of very extensive research, especially by Friedrich Maurer, that they have been evolved from the cutaneous sense-organs of amphibian ancestors by modification of functions; the epidermic structure is very similar in both in its embryonic rudiments. This modern view, which had the support of the greatest expert on the vertebrates, Carl Gegenbaur, can be harmonised with the older theory to an extent, in the sense that both formations, scales and hairs, were very closely connected originally. Probably the conical budding of the skin-sense layer grew up under the protection of the horny scale, and became an organ of touch subsequently by the cornification of the hairs; many hairs are still sensory organs (tactile hairs on the muzzle and cheeks of many mammals: pubic hairs).
This middle position of the genetic connection of scales and hairs was advanced in my Systematic Phylogeny of the Vertebrates (p. 433). It is confirmed by the similar arrangement of the two cutaneous formations. As Maurer pointed out, the hairs, as well as the cutaneous sense-organs and the scales, are at first arranged in regular longitudinal series, and they afterwards break into alternate groups. In the embryo of a bear two inches long, which I owe to the kindness of Herr von Schmertzing (of Arva Varallia, Hungary), the back is covered with sixteen to twenty alternating longitudinal rows of scaly protuberances (Fig. 289). They are at the same time arranged in regular transverse rows, which converge at an acute angle from both sides towards the middle of the back. The tip of the scale-like wart is turned inwards. Between these larger hard scales (or groups of hairs) we find numbers of rudimentary smaller hairs.
The human embryo is, as a rule, entirely clothed with a thick coat of fine wool during the last three or four weeks of gestation. This embryonic woollen coat (Lanugo) generally disappears in part during the last weeks of fœtal life but in any case, as a rule, it is lost immediately after birth, and is replaced by the thinner coat of the permanent hair. These permanent hairs grow out of hair-follicles, which are formed from the root-sheaths of the disappearing wool-fibres. The embryonic wool-coat usually, in the case of the human embryo, covers the whole body, with the exception of the palms of the hands and soles of the feet. These parts are always bare, as in the case of apes and of most other mammals. Sometimes the wool-coat of the embryo has a striking effect, by its colour, on the later permanent hair-coat. Hence it happens occasionally, for instance, among our Indo-Germanic races, that children of blond parents seem—to the dismay of the latter—to be covered at birth with a dark brown or even a black woolly coat. Not until this has disappeared do we see the permanent blond hair which the child has inherited. Sometimes the darker coat remains for weeks, and even months, after birth. This remarkable woolly coat of the human embryo is a legacy from the apes, our ancient long-haired ancestors.
Fig.288. Mammary gland of a new-born infant. Fig. 288—Mammary gland of a new-born infant, a original central gland, b small and c large buds of same. (From Langer.)
It is not less noteworthy that many of the higher apes approach man in the thinness of the hair on various parts of the body. With most of the apes, especially the higher Catarrhines (or narrow-nosed apes), the face is mostly, or entirely, bare, or at least it has hair no longer or thicker than that of man. In their case, too, the back of the head is usually provided with a thicker growth of hair; this is lacking, however, in the case of the bald-headed chimpanzee (Anthropithecus calvus). The males of many species of apes have a considerable beard on the cheeks and chin; this sign of the masculine sex has been acquired by sexual selection. Many species of apes have a very thin covering of hair on the breast and the upper side of the limbs—much thinner than on the back or the under side of the limbs. On the other hand, we are often astonished to find tufts of hair on the shoulders, back, and extremities of members of our Indo-Germanic and of the Semitic races. Exceptional hair on the face, as on the whole body, is hereditary in certain families of hairy men. The quantity and the quality of the hair on head and chin are also conspicuously transmitted in families. These extraordinary variations in the total and partial hairy coat of the body, which are so noticeable, not only in comparing different races of men, but also in comparing different families of the same race, can only be explained on the assumption that in man the hairy coat is, on the whole, a rudimentary organ, a useless inheritance from the more thickly-coated apes. In this man resembles the elephant, rhinoceros, hippopotamus, whale, and other mammals of various orders, which have also, almost entirely or for the most part, lost their hairy coats by adaptation.
The particular process of adaptation by which man lost the growth of hair on most parts of his body, and retained or augmented it at some points, was most probably sexual selection. As Darwin luminously showed in his Descent of Man, sexual selection has been very active in this respect. As the male anthropoid apes chose the females with the least hair, and the females favoured the males with the finest growths on chin and head, the general coating of the body gradually degenerated, and the hair of the beard and head was more strongly developed. The growth of hair at other parts of the body (arm-pit, pubic region) was also probably due to sexual selection. Moreover, changes of climate, or habits, and other adaptations unknown to us, may have assisted the disappearance of the hairy coat.
Fig.289. Embryo of a bear (Ursus arctos). Fig. 289—Embryo of a bear (Ursus arctos). A seen from ventral side, B from the left.
The fact that our coat of hair is inherited directly from the anthropoid apes is proved in an interesting way, according to Darwin, by the direction of the rudimentary hairs on our arms, which cannot be explained in any other way. Both on the upper and the lower part of the arm they point towards the elbow. Here they meet at an obtuse angle. This curious arrangement is found only in the anthropoid apes—gorilla, chimpanzee, orang, and several species of gibbons—besides man (Figs. 203, 207). In other species of gibbon the hairs are pointed towards the hand both in the upper and lower arm, as in the rest of the mammals. We can easily explain this remarkable peculiarity of the anthropoids and man on the theory that our common ancestors were accustomed (as the anthropoid apes are to-day) to place their hands over their heads, or across a branch above their heads, during rain. In this position, the fact that the hairs point downwards helps the rain to run off. Thus the direction of the hair on the lower part of our arm reminds us to-day of that useful custom of our anthropoid ancestors.
The nervous system in man and all the other Vertebrates is, when fully formed, an extremely complex apparatus, that we may compare, in anatomic structure and physiological function, with an extensive telegraphic system. The chief station of the system is the central marrow or central nervous system, the innumerable ganglionic cells or neurona (Fig. 9)of which are connected by branching processes with each other and with numbers of very fine conducting wires. The latter are the peripheral and ubiquitous nerve-fibres; with their terminal apparatus, the sense-organs, etc., they constitute the conducting marrow or peripheral nervous system. Some of them—the sensory nerve-fibres—conduct the impressions from the skin and other sense-organs to the central marrow; others—the motor nerve-fibres—convey the commands of the will to the muscles.
The central nervous system or central marrow (medulla centralis) is the real organ of psychic action in the narrower sense. However we conceive the intimate connection of this organ and its functions, it is certain that its characteristic actions, which we call sensation, will, and thought, are inseparably dependent on the normal development of the material organ in man and all the higher animals. We must, therefore, pay particular attention to the evolution of the latter. As it can give us most important information regarding the nature of the “soul,” it should be full of interest. If the central marrow develops in just the same way in the human embryo as in the embryo of the other mammals, the evolution of the human psychic organ from the central organ of the other mammals, and through them from the lower vertebrates, must be beyond question. No one can doubt the momentous bearing of these embryonic phenomena.
CaFig.290. Human embryo, three months old, from the dorsal side: brain and spinal cord exposed. Fig. 291. Central marrow of a human embryo, four months old, from the back.nyon Fig. 290—Human embryo, three months old, from the dorsal side: brain and spinal cord exposed. (From Kölliker.) h cerebral hemispheres (fore brain), m corpora quadrigemina (middle brain), c cerebellum (hind brain): under the latter is the triangular medulla oblongata (after brain). Fig. 291—Central marrow of a human embryo, four months old, from the back. (From Kölliker.) h large hemispheres, v quadrigemina, c cerebellum, mo medulla oblongata: underneath it the spinal cord.
In order to understand them fully we must first say a word or two of the general form and the anatomic composition of the mature human central marrow. Like the central nervous system of all the other Craniotes, it consists of two parts, the head-marrow or brain (medulla capitis or encephalon) and the spinal-marrow (medulla spinalis or notomyelon). The one is enclosed in the bony skull, the other in the bony vertebral column. Twelve pairs of cerebral nerves proceed from the brain, and thirty-one pairs of spinal nerves from the spinal cord, to the rest of the body (Fig. 171). On general anatomic investigation the spinal marrow is found to be a cylindrical cord, with a spindle-shaped bulb both in the region of the neck above (at the last cervical vertebra) and the region of the loins (at the first lumbar vertebra) below (Fig. 291). At the cervical bulb the strong nerves of the upper limbs, and at the lumbar bulb those of the lower limbs, proceed from the spinal cord. Above, the latter passes into the brain through the medulla oblongata (Fig. 291 mo). The spinal cord seems to be a thick mass of nervous matter, but it has a narrow canal at its axis, which passes into the further cerebral ventricles above, and is filled, like these, with a clear fluid.
The brain is a large nerve-mass, occupying the greater part of the skull, of most elaborate structure. On general examination it divides into two parts, the cerebrum and cerebellum. The cerebrum lies in front and above, and has the familiar characteristic convolutions and furrows on its surface (Figs. 292, 293). On the upper side it is divided by a deep longitudinal fissure into two halves, the cerebral hemispheres; these are connected by the corpus callosum. The large cerebrum is separated from the small cerebellum by a deep transverse furrow. The latter lies behind and below, and has also numbers of furrows, but much finer and more regular, with convolutions between, at its surface. The cerebellum also is divided by a longitudinal fissure into two halves, the “small hemispheres”; these are connected by a worm-shaped piece, the vermis cerebelli, above, and by the broad pons Varolii below (Fig. 292 VI).
Fig.292. The human brain, seen from below. Fig. 292—The human brain, seen from below. (From H. Meyer.) Above (in front) is the cerebrum with its extensive branching furrows; below (behind) the cerebellum with its narrow parallel furrows. The Roman numbers indicate the roots of the twelve pairs of cerebral nerves in a series towards the rear.
But comparative anatomy and ontogeny teach us that in man and all the other Craniotes the brain is at first composed, not of these two, but of three, and afterwards five, consecutive parts. These are found in just the same form—as five consecutive vesicles—in the embryo of all the Craniotes, from the Cyclostoma and fishes to man. But, however much they agree in their rudimentary condition, they differ considerably afterwards. In man and the higher mammals the first of these ventricles, the cerebrum, grows so much that in its mature condition it is by far the largest and heaviest part of the brain. To it belong not only the large hemispheres, but also the corpus callosum that unites them, the olfactory lobes, from which the olfactory nerves start, and most of the structures that are found at the roof and bottom of the large lateral ventricles inside the two hemispheres, such as the corpora striata. On the other hand, the optic thalami, which lie between the latter, belong to the second division, which develops from the “intermediate brain ”; to the same section belong the single third cerebral ventricle and the structures that are known as the corpora geniculata, the infundibulum, and the pineal gland. Behind these parts we find, between the cerebrum and cerebellum, a small ganglion composed of two prominences, which is called the corpus quadrigeminum on account of a superficial transverse fissure cutting across (Figs. 290 m and 291 v). Although this quadrigeminum is very insignificant in man and the higher mammals, it forms a special third section, greatly developed in the lower vertebrates, the “middle brain.” The fourth section is the “hind-brain” or little brain (cerebellum) in the narrower sense, with the single median part, the vermis, and the pair of lateral parts, the “small hemispheres” (Fig. 291 c). Finally, we have the fifth and last section, the medulla oblongata (Fig. 291 mo), which contains the single fourth cerebral cavity and the contiguous parts (pyramids, olivary bodies, corpora restiformia). The medulla oblongata passes straight into the medulla spinalis (spinal cord). The narrow central canal of the spinal cord continues above into the quadrangular fourth cerebral cavity of the medulla oblongata, the floor of which is the quadrangular depression. From here a narrow duct, called “the aqueduct of Sylvius,” passes through the corpus quadrigeminum to the third cerebral ventricle, which lies between the two optic thalami; and this in turn is connected with the pairs of lateral ventricles which lie to the right and left in the large hemispheres. Thus all the cavities of the central marrow are directly interconnected. All these parts of the brain have an infinitely complex structure in detail, but we cannot go into this. Although it is much more elaborate in man and the higher Vertebrates than in the lower classes, it develops in them all from the same rudimentary structure, the five simple cerebral vesicles of the embryonic brain.
But before we consider the development of the complicated structure of the brain from this simple series of vesicles, let us glance for a moment at the lower animals, which have no brain. Even in the skull-less vertebrate, the Amphioxus, we find no independent brain, as we have seen. The whole central marrow is merely a simple cylindrical cord which runs the length of the body, and ends equally simply at both extremities—a plain medullary tube. All that we can discover is a small vesicular bulb at the foremost part of the tube, a degenerate rudiment of a primitive brain. We meet the same simple medullary tube in the first structure of the ascidia larva, in the same characteristic position, above the chorda. On closer examination we find here also a small vesicular swelling at the fore end of the tube, the first trace of a differentiation of it into brain and spinal cord. It is probable that this differentiation was more advanced in the extinct Provertebrates, and the brain-bulb more pronounced (Figs. 98–102). The brain is phylogenetically older than the spinal cord, as the trunk was not developed until after the head. If we consider the undeniable affinity of the Ascidiæ to the Vermalia, and remember that we can trace all the Chordonia to lower Vermalia, it seems probable that the simple central marrow of the former is equivalent to the simple nervous ganglion, which lies above the gullet in the lower worms, and has long been known as the “upper pharyngeal ganglion” (ganglion pharyngeum superius); it would be better to call it the primitive or vertical brain (acroganglion).
Probably this upper pharyngeal ganglion of the lower worms is the structure from which the complex central marrow of the higher animals has been evolved. The medullary tube of the Chordonia has been formed by the lengthening of the vertical brain on the dorsal side. In all the other animals the central nervous system has been developed in a totally different way from the upper pharyngeal ganglion; in the Articulates, especially, a pharyngeal ring, with ventral marrow, has been added. The Molluscs also have a pharyngeal ring, but it is not found in the Vertebrates. In these the central marrow has been prolonged down the dorsal side; in the Articulates down the ventral side. This fact proves of itself that there is no direct relationship between the Vertebrates and the Articulates. The unfortunate attempts to derive the dorsal marrow of the former from the ventral marrow of the latter have totally failed (cf. p. 219).
Fig.293. The human brain, seen from the left. Fig. 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–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 (Figs. 133–137). At first this tube lies directly underneath the horny plate; but it afterwards travels inwards, the upper edges of the provertebral plates growing together between the horny plate and the tube, joining above the latter, and forming a completely closed canal. 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. Figs. 143–146).
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 (Fig. 153 hb). But in these it soon passes 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, Fig. 154 v, m, h). Then the first and third are sub-divided by fresh constrictions, and thus we get five successive sections (Fig. 155).
Fig.294. Central marrow of the human embryo from the seventh week, 4/5 inch long. Fig. 294. The brain from above. Fig. 295. The brain with the uppermost part of the cord, from the left. Fig. 296. Back view of the whole embryo: brain and spinal cord exposed. Fig. 294–296—Central marrow of the human embryo from the seventh week, 4/5 inch long. (From Kölliker.) Fig. 294. The brain from above, v fore brain, z intermediate brain, m middle brain, h hind brain, n after brain. Fig. 2955. The brain with the uppermost part of the cord, from the left. Fig. 296. Back view of the whole embryo: brain and spinal cord exposed.
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 (Fig. 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 classes 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.
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 passing 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 (Fig. 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 (Figs. 293, 301–304).
Fig.297. Head of a chick embryo (hatched fifty-eight hours), from the back. Fig. 297—Head of a chick embryo (hatched fifty-eight hours), from the back. (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.
Fig.298. Brain of three craniote embryos in vertical section. Fig. 299. Brain of a shark (Scyllium), back view. Fig. 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.) Fig. 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.)
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.
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 (Figs. 301–303). 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.
Fig.300. Brain and spinal cord of the frog. Fig. 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′ roots of the spinal nerves, t terminal fibres of the spinal cord. (From Gegenbaur.)
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 quantitatively. The mass 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.
Fig.301. Brain of an ox-embryo, two inches in length. Fig. 301—Brain of an ox-embryo, two inches in length. (From Mihalkovics.) 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.
Fig. 302. Brain of a human embryo, twelve weeks old. Fig. 302—Brain of a human embryo, twelve weeks old. (From Mihalkovics.) 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.
In the lower mammals the surface of the cerebral hemispheres is quite smooth and level, as in the rabbit (Fig. 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 (Figs. 292, 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.
Fig.303. Brain of a human embryo, twenty-four weeks old, halved in the median plane: right hemisphere seen from inside. Fig. 303—Brain of a human embryo, twenty-four weeks old, halved in the median plane: right hemisphere seen from inside. (From Mihalkovics.) 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.
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 constitute a deeper gulf between the lower and higher apes than between the higher apes and man.
Fig.304. Brain of the rabbit. Fig. 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, 5–8 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 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.
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 (Fig. 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.
XXV. 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 assistance 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 classes 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 sexual sense is bound up with the skin of the external sexual 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 space (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 Gastræa 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 assumption 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 horny plate, partly from the brain, the foremost part, of the medullary tube, after it has separated from the horny 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 homogeneous 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.
Fig.305. Head of a shark (Scyllium), from the ventral side. Fig. 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′ nasal fold drawn up. (The dots are openings of the mucous canals.) (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 sex). In the second group the nerves spread out in the mucous coat of cavities which are at first depressions in or invaginations 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 space).
There is little to be said of the development of the lower sense-organs. We have already considered (p. 268) 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 sexual sense, the male penis and the female clitoris; they are processes of the skin, the development of which we will consider later (together with the rest of the sexual parts, Chapter XXIX). The evolution of the organ of taste, the tongue and palate, will also be treated later, together with that of the alimentary canal to which these parts belong (Chapter XXVII). 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 (p. 139). 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.
Figs. 306 and 307. Head of a chick embryo, three days old. Fig. 308. Head of a chick embryo, four days old, from below. Figs. 309 and 310. Heads of chick embryos: 309 from the end of the fourth, 310 from the beginning of the fifth week. Fig. 306 and 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.
Fig. 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″ second gill-arch, sp choroid fissure of eye, s gullet.
Fig. 309 and 310—Heads of chick embryos: 309 from the end of the fourth, 310 from the beginning of the fifth week. Letters as in Fig. 308, except: in inner, an outer, nasal process, nf nasal furrow, st frontal process, m mouth. (From Kölliker.).
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 partition, 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 passages without touching the mouth. This is the way the air usually passes 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 (partition), 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 passage 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 (Fig. 249).
Fig.311. Frontal section of the mouth and throat of a human embryo, neck half-inch long. Fig. 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 (Fig. 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″). 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 canal, the nasal canal. Henceforth we can penetrate from the external pits through the nasal canals direct into the mouth, which has been formed quite independently. In the Dipneusts and the lower Amphibia the internal aperture of the nasal canals lies in front (behind the lips); in the higher Amphibia it is right behind. Finally, in the three higher classes 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.
Fig.312. Diagrammatic section of the mouth-nose cavity. Fig. 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 partition (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 passed 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” (Figs. 306 n, 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 (Fig. 310 m) are clothed with the horny plate. The original separation of the former from the latter is, however, presently abolished, a process forming above the mouth-pit—the “frontal process” (Fig. 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 (Fig. 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 canal. Hence the nose of man and all the other Amniotes consists at this embryonic stage of a couple of narrow tubes, the nasal canals, 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.
A cone-shaped structure, which grows from below towards the lower ends of the two nasal processes and joins with them, plays an important part in the conversion of the open nasal groove into the closed canal. This is the upper-jaw process (Figs. 306–310 o). Below the mouth-pit are the gill-arches, which are separated by the gill-clefts. The first of these gill-arches, and the most important for our purpose, which we may call the maxillary (jaw) arch, forms the skeleton of the jaws. Above at the basis a small process grows out of this first gill-arch; this is the upper-jaw process. The first gill-arch itself develops a cartilage at one of its inner sides, the “Meckel cartilage” (named after its discoverer), on the outer surface of which the lower jaw is formed (Figs. 306–310 u). The upper-jaw process forms the chief part of the skeleton of that jaw, the palate bone, and the pterygoid bone. On its outer side is afterwards formed the upper-jaw bone, in the narrower sense, while the middle part of the skeleton of the upper jaw, the intermaxillary, develops from the foremost part of the frontal process.
The two upper-jaw processes are of great importance in the further development of the face. From them is formed, growing into the primitive mouth-cavity, the important horizontal partition (the palate) that divides the former into two distinct cavities. The upper cavity, into which the nasal canals open, now develops into the nasal cavity, the air-passage and the organ of smell. The lower cavity forms the permanent secondary mouth (Fig. 312 m), the food-passage and the organ of taste. Both the upper and lower cavities open behind into the gullet (pharynx). The hard palate that separates them is formed by the joining of two lateral halves, the horizontal plates of the two upper-jaw processes, or the palate-plates (p). When these do not, sometimes, completely join in the middle, a longitudinal cleft remains, through which we can penetrate from the mouth straight into the nasal cavity. This is the malformation known as “wolf’s throat.” “Hare-lip” is the lesser form of the same defect. At the same time as the horizontal partition of the hard palate a vertical partition is formed by which the single nasal cavity is divided into two sections—a right and left half (Fig. 312 n, n).
Figs. 313 and 314. Upper part of the body of a human embryo, two-thirds of an inch long, of the sixth week; Fig. 313 from the left, Fig. 314 from the front. Figs. 313 and 314—Upper part of the body of a human embryo, two-thirds of an inch long, of the sixth week; Fig. 313 from the left, Fig. 314 from the front. The origin of the nose and the upper lip from two lateral and originally separate halves can be clearly seen. Nose and upper lip are large in proportion to the rest of the face, and especially to the lower lip. (From Kollmann.)
The double nose has now acquired the characteristic form that man shares with the other mammals. Its further development is easy to follow; it consists of the formation of the inner and outer processes of the walls of the two cavities. The external nose is not formed until long after all these essential parts of the internal organ of smell. The first traces of it in the human embryo are found about the middle of the second month (Figs. 313–316). As can be seen in any human embryo during the first month, there is at first no trace of the external nose. It only develops afterwards from the foremost nasal part of the primitive skull, growing forwards from behind. The characteristic human nose is formed very late. Much stress is at times laid on this organ as an exclusive privilege of man. But there are apes that have similar noses, such as the long-nosed ape.
The evolution of the eye is not less interesting and instructive than that of the nose. Although this noblest of the sensory organs is one of the most elaborate and purposive on account of its optic perfection and remarkable structure, it nevertheless develops, without preconceived design, from a simple process of the outer germinal layer. The fully-formed human eye is a round capsule, the eye-ball (Fig. 317). This lies in the bony cavity of the skull, surrounded by protective fat and motor muscles. The greater part of it is taken up with a semi-fluid, transparent gelatinous substance, the corpus vitreum. The crystalline lens is fitted into the anterior surface of the ball (Fig. 317 l). It is a lenticular, bi-convex, transparent body, the most important of the refractive media in the eye. Of this group we have, besides the corpus vitreum and the lens, the watery fluid (humor aqueus) that is found in front of the lens (at the letter m in Fig. 317). These three transparent refractive media, by which the rays of light that enter the eye are broken up and re-focussed, are enclosed in a solid round capsule, composed of several different coats, something like the concentric layers of an onion. The outermost and thickest of these envelopes is the white sclerotic coat of the eye. It consists of tough white connective tissue. In front of the lens a circular, strongly-curved, transparent plate is fitted into the sclerotic, like the glass of a watch—the cornea (b). At its outer surface the cornea is covered with a very thin layer of the epidermis; this is known as the conjunctiva. It goes from the cornea over the inner surface of the eye-lids, the upper and lower folds which we draw over the eye in closing it. At the inner corner of the eye we have a rudimentary organ in the shape of the relic of a third (inner) eye-lid, which is greatly developed, as “nictitating (winking) membrane,” in the lower Vertebrates (p. 5). Underneath the upper eye-lid are the lachrymal glands, the product of which, the lachrymal fluid, keeps the outer surface of the eye smooth and clean.
Fig.315. Face of a human embryo, seven weeks old. Fig. 315—Face of a human embryo, seven weeks old. (From Kollmann.) Joining of the nasal processes (e outer, i inner) with the upper-jaw process (o), n nasal wall, a ear-opening.
Immediately under the sclerotic we find a very delicate, dark-red membrane, very rich in blood-vessels—the choroid coat—and inside this the retina (o), the expansion of the optic nerve (i). The latter is the second cerebral nerve. It proceeds from the optic thalami (the second cerebral vesicle) to the eye; penetrates its outer envelopes, and then spreads out like a net between the choroid and the corpus vitreum. Between the retina and the choroid there is a very delicate membrane, which is usually (but wrongly) associated with the latter. This is the black pigment-membrane (n). It consists of a single stratum of graceful, hexagonal, regularly-joined cells, full of granules of black colouring matter. This pigment membrane clothes, not only the inner surface of the choroid proper, but also the hind surface of its anterior muscular continuation, which covers the edge of the lens in front as a circular membrane, and arrests the rays of light at the sides. This is the well-known iris of the eye (h), coloured differently in different individuals (blue, grey, brown, etc.); it forms the anterior border of the choroid. The circular opening that is left in the middle is the pupil, through which the rays of light penetrate into the eye. At the point where the iris leaves the anterior border of the choroid proper the latter is very thick, and forms a delicate crown of folds (g), which surrounds the edge of the lens with about seventy large and many smaller rays (corona ciliaris.)
Fig.316. Face of a human embryo, eight weeks old. Fig. 316—Face of a human embryo, eight weeks old. (From Ecker.)
At a very early stage a couple of pear-shaped vesicles develop from the foremost part of the first cerebral vesicle in the embryo of man and the other Craniotes (Figs. 155 a, 297 au). These growths are the primary optic vesicles. They are at first directed outwards and forwards, but presently grow downward, so that, after the complete separation of the five cerebral vesicles, they lie at the base of the intermediate brain. The inner cavities of these pear-shaped vesicles, which soon attain a considerable size, are openly connected with the ventricle of the intermediate brain by their hollow stems. They are covered externally by the epidermis.
At the point where this comes into direct contact with the most curved part of the primary optic vesicle there is a thickening (l) and also a depression (o) of the horny plate (Fig. 318, I). This pit, which we may call the lens-pit, is converted into a closed sac, the thick- walled lens-vesicle (2, l), the thick edges of the pit joining together above it. In the same way in which the medullary tube separates from the outer germinal layer, we now see this lens-sac sever itself entirely from the horny plate (h), its source of origin. The hollow of the sac is afterwards filled with the cells of its thick walls, and thus we get the solid crystalline lens. This is, therefore, a purely epidermic structure. Together with the lens the small underlying piece of corium-plate also separates from the skin.
As the lens separates from the corneous plate and grows inwards, it necessarily hollows out the contiguous primary optic vesicle (Fig. 318, 1–3). This is done in just the same way as the invagination of the blastula, which gives rise to the gastrula in the amphioxus (Fig. 38 C–F). In both cases the hollowing of the closed vesicle on one side goes so far that at last the inner, folded part touches the outer, not folded part, and the cavity disappears. As in the gastrula the first part is converted into the entoderm and the latter into the ectoderm, so in the invagination of the primary optic vesicle the retina (r) is formed from the first (inner) part, and the black pigment membrane (u) from the latter (outer, non-invaginated) part. The hollow stem of the primary optic vesicle is converted into the optic nerve. The lens (l), which has so important a part in this process, lies at first directly on the invaginated part, or the retina (r). But they soon separate, a new structure, the corpus vitreum (gl), growing between them. While the lenticular sac is being detached and is causing the invagination of the primary optic vesicle, another invagination is taking place from below; this proceeds from the superficial part of the skin-fibre layer—the corium of the head. Behind and under the lens a last-shaped process rises from the cutis-plate (Fig. 319 g), hollows out the cup-shaped optic vesicle from below, and presses between the lens (l) and the retina (i). In this way the optic vesicle acquires the form of a hood.
Fig.317. The human eye in section. Fig. 317—The human eye in section. a sclerotic coat, b cornea, c conjunctiva, d circular veins of the iris, e choroid coat, f ciliary muscle, g corona ciliaris, h iris, i optic nerve, k anterior border of the retina, l crystalline lens, m inner covering of the cornea (aqueous membrane), n pigment membrane, o retina, p Petit’s canal, q yellow spot of the retina. (From Helmholtz.)
Finally, a complete fibrous envelope, the fibrous capsule of the eye-ball, is formed about the secondary optic vesicle and its stem (the secondary optic nerve). It originates from the part of the head-plates which immediately encloses the eye. This fibrous envelope takes the form of a closed round vesicle, surrounding the whole of the ball and pushing between the lens and the horny plate at its outer side. The round wall of the capsule soon divides into two different membranes by surface-cleavage. The inner membrane becomes the choroid or vascular coat, and in front the ciliary corona and iris. The outer membrane is converted into the white protective or sclerotic coat—in front, the transparent cornea. The eye is now formed in all its essential parts. The further development—the complicated differentiation and composition of the various parts—is a matter of detail.
The chief point in this remarkable evolution of the eye is the circumstance that the optic nerve, the retina, and the pigment membrane originate really from a part of the brain—an outgrowth of the intermediate brain—while the lens, the chief refractive body, develops from the outer skin. From the skin—the horny plate—also arises the delicate conjunctiva, which afterwards covers the outer surface of the eyeball. The lachrymal glands are ramified growths from the conjunctiva (Fig. 286). All these important parts of the eye are products of the outer germinal layer. The remaining parts—the corpus vitreum (with the vascular capsule of the lens), the choroid (with the iris), and the sclerotic (with the cornea)—are formed from the middle germinal layer.
Fig.318. Eye of the chick embryo in longitudinal section (1. from an embryo sixty-five hours old; 2. from a somewhat older embryo; 3. from an embryo four days old). Fig. 318—Eye of the chick embryo in longitudinal section (1. from an embryo sixty-five hours old; 2. from a somewhat older embryo; 3. from an embryo four days old). h horny plate, o lens-pit, l lens (in 1. still part of the epidermis, in 2. and 3. separated from it), x thickening of the horny plate at the point where the lens has severed itself, gl corpus vitreum, r retina, u pigment membrane. (From Remak.)
The outer protection of the eye, the eye-lids, are merely folds of the skin, which are formed in the third month of human embryonic life. In the fourth month the upper eye-lid reaches the lower, and the eye remains covered with them until birth. As a rule, they open wide shortly before birth (sometimes only after birth). Our craniote ancestors had a third eye-lid, the nictitating membrane, which was drawn over the eye from its inner angle. It is still found in many of the Selachii and Amniotes. In the apes and man it has degenerated, and there is now only a small relic of it at the inner corner of the eye, the semi-lunar fold, a useless rudimentary organ (cf. p. 32). The apes and man have also lost the Harderian gland that opened under the nictitating membrane; we find this in the rest of the mammals, and the birds, reptiles, and amphibia.
The peculiar embryonic development of the vertebrate eye does not enable us to draw any definite conclusions as to its obscure phylogeny; it is clearly cenogenetic to a great extent, or obscured by the reduction and curtailment of its original features. It is probable that many of the earlier stages of its phylogeny have disappeared without leaving a trace. It can only be said positively that the peculiar ontogeny of the complicated optic apparatus in man follows just the same laws as in all the other Vertebrates. Their eye is a part of the fore brain, which has grown forward towards the skin, not an original cutaneous sense-organ, as in the Invertebrates.
Fig.319. Horizontal transverse section of the eye of a human embryo, four weeks old. Fig. 319—Horizontal transverse section of the eye of a human embryo, four weeks old. (From Kölliker.) t lens (the dark wall of which is as thick as the diameter of the central cavity), g corpus vitreum (connected by a stem, g, with the corium), v vascular loop (pressing behind the lens inside the corpus vitreum by means of this stem g), i retina (inner thicker, invaginated layer of the primary optic vesicle), a pigment membrane (outer, thin, non-invaginated layer of same), h space between retina and pigment membrane (remainder of the cavity of the primary optic vesicle).
The vertebrate ear resembles the eye and nose in many important respects, but is different in others, in its development. The auscultory organ in the fully-developed man is like that of the other mammals, and especially the apes, in the main features. As in them, it consists of two chief parts—an apparatus for conducting sound (external and middle ear) and an apparatus for the sensation of sound (internal ear). The external ear opens in the shell at the side of the head (Fig. 320 a). From this point the external passage (b), about an inch in length, leads into the head. The inner end of it is closed by the tympanum, a vertical, but not quite upright, thin membrane of an oval shape (c). This tympanum separates the external passage from the tympanic cavity (d). This is a small cavity, filled with air, in the temporal bone; it is connected with the mouth by a special tube. This tube is rather longer, but much narrower, than the outer passage, leads inwards obliquely from the anterior wall of the tympanic cavity, and opens in the throat below, behind the nasal openings. It is called the Eustachian tube (e); it serves to equalise the pressure of the air within the tympanic cavity and the outer atmosphere that enters by the external passage. Both the Eustachian tube and the tympanic cavity are lined with a thin mucous coat, which is a direct continuation of the mucous lining of the throat. Inside the tympanic cavity there are three small bones which are known (from their shape) as the hammer, anvil, and stirrup (Fig. 320, f, g, h). The hammer (f) is the outermost, next to the tympanum. The anvil (g) fits between the other two, above and inside the hammer. The stirrup (h) lies inside the anvil, and touches with its base the outer wall of the internal ear, or auscultory vesicle. All these parts of the external and middle ear belong to the apparatus for conducting sound. Their chief task is to convey the waves of sound through the thick wall of the head to the inner-lying auscultory vesicle. They are not found at all in the fishes. In these the waves of sound are conveyed directly by the wall of the head to the auscultory vesicle.
Fig.320. The human ear (left ear, seen from the front). Fig. 320—The human ear (left ear, seen from the front), a shell of ear, b external passage, c tympanum, d tympanic cavity, e Eustachian tube, f, g, h the three bones of the ear (f hammer, g anvil, h stirrup), i utricle, k the three semi-circular canals, l the sacculus, m cochlea, n auscultory nerve.
Fig. 321. The bony labyrinth of the human ear (left side). Fig. 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.)
The internal apparatus for the sensation of sound, which receives the waves of sound from the conducting apparatus, consists in man and all other mammals of a closed auscultory vesicle filled with fluid and an auditory nerve, the ends of which expand over the wall of this vesicle. The vibrations of the sound-waves are conveyed by these media to the nerve-endings. In the labyrinthic water that fills the auscultory vesicle there are small stones at the points of entry of the acoustic nerves, which are composed of groups of microscopic calcareous crystals (otoliths). The auscultory organ of most of the Invertebrates has substantially the same composition. It usually consists of a closed 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 (Fig. 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.
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 (Fig. 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 (Figs. 322 B lv, 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 (Fig. 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.
Fig.322. Development of the auscultory labyrinth of the chick, in five successive stages (A to E). Fig. 322—Development of the auscultory labyrinth of the chick, in five successive stages (A–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.)
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 (Fig. 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.
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”).
Fig.323. Primitive skull of the human embryo, four weeks old, vertical section, left half seen internally. Fig. 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 Kölliker.)
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.
Fig.324. The rudimentary muscles of the ear in the human skull. Fig. 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 (Figs. 313, 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 (Fig. 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.
XXVI. 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.
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.
Fig.325. The human skeleton from the right. Fig. 326. The human skeleton. Front. Fig. 325—The human skeleton. From the right. Fig. 326—The human skeleton. Front.
Fig.327. The human vertebral column (standing upright, from the right side). Fig. 327—The human vertebral column (standing upright, from the right side). (From H. Meyer.)
Fig.328. A piece of the axial rod (chorda dorsalis), from a sheep embryo. Fig. 328—A piece of the axial rod (chorda dorsalis), from a sheep embryo. a cuticular sheath, b cells. (From Kölliker.)
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 (Fig. 325, the human skeleton seen from the right; Fig. 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.
The vertebral column (Fig. 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 vertebræ 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 vertebræ 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 vertebræ, twelve dorsal vertebræ, five lumbar vertebræ, five sacral vertebræ, and four to six caudal vertebræ. The uppermost, or those next to the skull, are the cervical vertebræ (Fig. 327); they have a hole in each of the lateral processes. There are seven of these vertebræ 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 or hedgehog. This constant number, which has few exceptions (due to adaptation), is a strong proof of the common descent of the mammals; it can only be explained by faithful heredity from a common stem-form, a primitive mammal with seven cervical vertebræ. If each species had been created separately, it would have been better to have given the long-necked mammals more, and the short-necked animals less, cervical vertebræ. Next to these come the dorsal (or pectoral) vertebræ, which number twelve to thirteen (usually twelve) in man and most of the other mammals. Each dorsal vertebra (Fig. 165) has at the side, connected by joints, a couple of ribs, long bony arches that lie in and protect the wall of the chest. The twelve pairs of ribs, together with the connecting intercostal muscles and the sternum, which joins the ends of the right and left ribs in front, form the chest (thorax). In this strong and elastic frame are the lungs, and between them the heart. Next to the dorsal vertebræ comes a short but stronger section of the column, formed of five large vertebræ. These are the lumbar vertebræ (Fig. 166); they have no ribs and no holes in the transverse processes. To these succeeds the sacral bone, which is fitted between the two halves of the pelvic zone. The sacrum is formed of five vertebræ, completely blended together. Finally, we have at the end a small rudimentary caudal column, the coccyx. This consists of a varying number (usually four, more rarely three, or five or six) of small degenerated vertebræ, and is a useless rudimentary organ with no actual physiological significance. Morphologically, however, it is of great interest as an irrefragable proof of the descent of man and the anthropoids from long-tailed apes. On no other theory can we explain the existence of this rudimentary tail. In the earlier stages of development the tail of the human embryo protrudes considerably. It afterwards atrophies; but the relic of the atrophied caudal vertebræ and of the rudimentary muscles that once moved it remains permanently. Sometimes, in fact, the external tail is preserved. The older anatomists say that the tail is usually one vertebra longer in the human female than in the male (or four against five); Steinbach says it is the reverse.
Fig.329. Three dorsal vertebræ, from a human embryo, eight weeks old, in lateral longitudinal section. Fig. 329—Three dorsal vertebræ, from a human embryo, eight weeks old, in lateral longitudinal section. v cartilaginous vertebral body, li inter-vertebral disks, ch chorda. (From Kölliker.)
Fig.330. A dorsal vertebra of the same embryo, in lateral transverse section. Fig. 330—A dorsal vertebra of the same embryo, in lateral transverse section. cv cartilaginous vertebral body, ch chorda, pr transverse process, a vertebral arch (upper arch), c upper end of the rib (lower arch). (From Kölliker.)
In the human vertebral column there are usually thirty-three vertebræ. It is interesting to find, however, that the number often changes, one or two vertebræ dropping out or an additional one appearing. Often, also, a mobile rib is formed at the last cervical or the first lumbar vertebra, so that there are then thirteen dorsal vertebræ, besides six cervical and four lumbar. In this way the contiguous vertebræ of the various sections of the column may take each other’s places.
In order to understand the embryology of the human vertebral column we must first carefully consider the shape and connection of the vertebræ. Each vertebra has, in general, the shape of a seal-ring (Figs. 164–166). The thicker portion, which is turned towards the ventral side, is called the body of the vertebra, and forms a short osseous disk; the thinner part forms a semi-circular arch, the vertebral arch, and is turned towards the back. The arches of the successive vertebræ are connected by thin intercrural ligaments in such a way that the cavity they collectively enclose represents a long canal. In this vertebral canal we find the trunk part of the central nervous system, the spinal cord. Its head part, the brain, is enclosed by the skull, and the skull itself is merely the uppermost part of the vertebral column, distinctively modified. The base or ventral side of the vesicular cranial capsule corresponds originally to a number of developed vertebral bodies; its vault or dorsal side to their combined upper vertebral arches.
While the solid, massive bodies of the vertebræ represent the real central axis of the skeleton, the dorsal arches serve to protect the central marrow they enclose. But similar arches develop on the ventral side for the protection of the viscera in the breast and belly. These lower or ventral vertebral arches, proceeding from the ventral side of the vertebral bodies, form, in many of the lower Vertebrates, a canal in which the large blood-vessels are enclosed on the lower surface of the vertebral column (aorta and caudal vein). In the higher Vertebrates the majority of these vertebral arches are lost or become rudimentary. But at the thoracic section of the column they develop into independent strong osseous arches, the ribs (costæ). In reality the ribs are merely large and independent lower vertebral arches, which have lost their original connection with the vertebral bodies.
Fig.331. Intervertebral disk of a new-born infant, transverse section. Fig. 331—Intervertebral disk of a new-born infant, transverse section. a rest of the chorda. (From Kölliker.)
If we turn from this anatomic survey of the composition of the column to the question of its development, I may refer the reader to earlier pages with regard to the first and most important points (pp. 145–148). It will be remembered that in the human embryo and that of the other vertebrates we find at first, instead of the segmented column, only a simple unarticulated cartilaginous rod. This solid but flexible and elastic rod is the axial rod (or the chorda dorsalis). In the lowest Vertebrate, the Amphioxus, it retains this simple form throughout life, and permanently represents the whole internal skeleton (Fig. 210 i). In the Tunicates, also, the nearest Invertebrate relatives of the Vertebrates, we meet the same chorda—transitorily in the passing larva tail of the Ascidia, permanently in the Copelata (Fig. 225 c). Undoubtedly both the Tunicates and Acrania have inherited the chorda from a common unsegmented stem-form; and these ancient, long-extinct ancestors of all the chordonia are our hypothetical Prochordonia.
Long before there is any trace of the skull, limbs, etc., in the embryo of man or any of the higher Vertebrates—at the early stage in which the whole body is merely a sole-shaped embryonic shield—there appears in the middle line of the shield, directly under the medullary furrow, the simple chorda. (Cf. Figs. 131–135 ch). It follows the long axis of the body in the shape of a cylindrical axial rod of elastic but firm composition, equally pointed at both ends. In every case the chorda originates from the dorsal wall of the primitive gut; the cells that compose it (Fig. 328 b) belong to the entoderm (Figs. 216–221). At an early stage the chorda develops a transparent structureless sheath, which is secreted from its cells (Fig. 328 a). This chordalemma is often called the “inner chorda-sheath,” and must not be confused with the real external sheath, the mesoblastic perichorda.
Fig. 332. Human skull. Fig. 332—Human skull.
But this unsegmented primary axial skeleton is soon replaced by the segmented secondary axial skeleton, which we know as the vertebral column. The provertebral plates (Fig. 124 s) differentiate from the innermost, median part of the visceral layer of the cœlom-pouches at each side of the chorda. As they grow round the chorda and enclose it they form the skeleton plate or skeletogenetic layer—that is to say, the skeleton-forming stratum of cells, which provides the mobile foundation of the permanent vertebral column and skull (scleroblast). In the head-half of the embryo the skeletal plate remains a continuous, simple, undivided layer of tissue, and presently enlarges into a thin-walled capsule enclosing the brain, the primordial skull. In the trunk-half the provertebral plate divides into a number of homogeneous, cubical, successive pieces; these are the several primitive vertebræ. They are not numerous at first, but soon increase as the embryo grows longer (Figs. 153–155).
Fig. 333. Skull of a new-born child. Fig. 333—Skull of a new-born child. (From Kollmann.) Above, in the three bones of the roof of the skull, we see the lines that radiate from the central points of ossification; in front, the frontal bone; behind, the occipital bone; between the two the large parietal bone, p. s the scurf bone, w mastoid fontanelle, f petrous bone, t tympanic bone, l lateral part, b bulla, j cheek-bone, a large wing of cuneiform bone, k fontanelle of cuneiform bone.
In all the Craniotes the soft, indifferent cells of the mesoderm, which originally compose the skeletal plate, are afterwards converted for the most part into cartilaginous cells, and these secrete a firm and elastic intercellular substance between them, and form cartilaginous tissue. Like most of the other parts of the skeleton, the membranous rudiments of the vertebræ soon pass into a cartilaginous state, and in the higher Vertebrates this is afterwards replaced by the hard osseous tissue with its characteristic stellate cells (Fig. 6). The primary axial skeleton remains a simple chorda throughout life in the Acrania, the Cyclostomes, and the lowest fishes. In most of the other Vertebrates the chorda is more or less replaced by the cartilaginous tissue of the secondary perichorda that grows round it. In the lower Craniotes (especially the fishes) a more or less considerable part of the chorda is preserved in the bodies of the vertebræ. In the mammals it disappears for the most part. By the end of the second month in the human embryo the chorda is merely a slender thread, running through the axis of the thick, cartilaginous vertebral column (Figs. 182 ch, 329 ch). In the cartilaginous vertebral bodies themselves, which afterwards ossify, the slender remnant of the chorda presently disappears (Fig. 330 ch). But in the elastic inter-vertebral disks, which develop from the skeletal plate between each pair of vertebral bodies (Fig. 329 li), a relic of the chorda remains permanently. In the new-born child there is a large pear-shaped cavity in each intervertebral disk, filled with a gelatinous mass of cells (Fig. 331 a).
Fig.334. Head-skeleton of a primitive fish. Fig. 334—Head-skeleton of a primitive fish. n nasal pit, eth cribriform bone region, orb orbit of eye, la wall of auscultory labyrinth, occ occipital region of primitive skull, cv vertebral column, a fore, bc hind-lip cartilage, o primitive upper jaw (palato-quadratum), u primitive lower jaw, II hyaloid bone, III–VIII first to sixth branchial arches. (From Gegenbaur.)
Though less sharply defined, this gelatinous nucleus of the elastic cartilaginous disks persists throughout life in the mammals, but in the birds and most reptiles the last trace of the chorda disappears. In the subsequent ossification of the cartilaginous vertebra the first deposit of bony matter (“first osseous nucleus”) takes place in the vertebral body immediately round the remainder of the chorda, and soon displaces it altogether. Then there is a special osseous nucleus formed in each half of the vertebral arch. The ossification does not reach the point at which the three nuclei are joined until after birth. In the first year the two osseous halves of the arches unite; but it is much later—in the second to the eighth year—
that they connect with the osseous vertebral bodies.
Fig.345. Roofs of the skulls of nine Primates (Cattarrhines), seen from above and reduced to a common size. Fig. 335—Roofs of the skulls of nine Primates (Cattarrhines), seen from above and reduced to a common size. 1 European, 2 Brazilian, 3 Pithecanthropus, 4 Gorilla, 5 Chimpanzee, 6 Orang, 7 Gibbon, 8 Tailed ape, 9 Baboon.
The bony skull (cranium), the head-part of the secondary axial skeleton, develops in just the same way as the vertebral column. The skull forms a bony envelope for the brain, just as the vertebral canal does for the spinal cord; and as the brain is only a peculiarly differentiated part of the head, while the spinal cord represents the longer trunk-section of the originally homogeneous medullary tube, we shall expect to find that the osseous coat of the one is a special modification of the osseous envelope of the other. When we examine the adult human skull in itself (Fig. 332), it is difficult to conceive how it can be merely the modified fore part of the vertebral column. It is an elaborate and extensive bony structure, composed of no less than twenty bones of different shapes and sizes. Seven of them form the spacious shell that surrounds the brain, in which we distinguish the solid ventral base below and the curved dorsal vault above. The other thirteen bones form the facial skull, which is especially the bony envelope of the higher sense-organs, and at the same time encloses the entrance of the alimentary canal. The lower jaw is articulated at the base of the skull (usually regarded as the XXI cranial bone). Behind the lower jaw we find the hyoid bone at the root of the tongue, also formed from the gill-arches, and a part of the lower arches that have developed as “head-ribs” from the ventral side of the base of the cranium.
Fig.336. Skeleton of the breast-fin of Ceratodus (biserial feathered skeleton). Fig. 337. Skeleton of the breast-fin of an early Selachius (Acanthias). Fig. 338. Skeleton of the breast-fin of a young Selachius. Fig. 336—Skeleton of the breast-fin of Ceratodus (biserial feathered skeleton). A, B, cartilaginous series of the fin-stem. rr cartilaginous fin-radii. (From Gunther.) Fig. 337—Skeleton of the breast-fin of an early Selachius (Acanthias). The radii of the median fin-border (B) have disappeared for the most part; a few only (R) are left. R, R, radii of the lateral fin-border, mt metapterygium, ms mesopterygium, p propterygium. (From Gegenbaur.) Fig. 338—Skeleton of the breast-fin of a young Selachius. The radii of the median fin-border have wholly disappeared. The shaded part on the right is the section that persists in the five-fingered hand of the higher Vertebrates. (b the three basal pieces of the fin: mt metapterygium, rudiment of the humerus, ms mesopterygium, p propterygium.) (From Gegenbaur.)
Although the fully-developed skull of the higher Vertebrates, with its peculiar shape, its enormous size, and its complex composition, seems to have nothing in common with the ordinary vertebræ, nevertheless even the older comparative anatomists came to recognise at the end of the eighteenth century that it is really nothing else originally than a series of modified vertebræ. When Goethe in 1790 “picked up the skull of a slain victim from the sand of the Jewish cemetery at Venice, he noticed at once that the bones of the face also could be traced to vertebræ (like the three hind-most cranial vertebræ).” And when Oken (without knowing anything of Goethe’s discovery) found at Ilenstein, “a fine bleached skull of a hind, the thought flashed across him like lightning: ‘It is a vertebral column.’”
This famous vertebral theory of the skull has interested the most distinguished zoologists for more than a century: the chief representatives of comparative anatomy have devoted their highest powers to the solution of the problem, and the interest has spread far beyond their circle. But it was not until 1872 that it was happily solved, after seven years’ labour, by the comparative anatomist who surpassed all other experts of this science in the second half of the nineteenth century by the richness of his empirical knowledge and the acuteness and depth of his philosophic speculations. Carl Gegenbaur has shown, in his classic Studies of the Comparative Anatomy of the Vertebrates (third section), that we find the most solid foundation for the vertebral theory of the skull in the head-skeleton of the Selachii. Earlier anatomists had wrongly started from the mammal skull, and had compared the several bones that compose it with the several parts of the vertebra (Fig. 333) they thought they could prove in this way that the fully-formed mammal skull was made of from three to six vertebræ.
Fig.339. Skeleton of the fore leg of an amphibian. Fig. 340. Skeleton of gorilla’s hand. Fig. 341. Skeleton of human hand, back. Fig. 339—Skeleton of the fore leg of an amphibian. h upper-arm (humerus), ru lower arm (r radius, u ulna), rcicu′, wrist-bones of first series (r radiale, i intermedium, c centrale, u′ ulnare). 1, 2, 3, 4, 5 wrist-bones of the second series. (From Gegenbaur.) Fig. 340—Skeleton of gorilla’s hand. (From Huxley.) Fig. 341—Skeleton of human hand, back. (From Meyer.)
The older theory was refuted by simple and obvious facts, which were first pointed out by Huxley. Nevertheless, the fundamental idea of it—the belief that the skull is formed from the head-part of the perichordal axial skeleton, just as the brain is from the simple medullary tube, by differentiation and modification—remained. The work now was to discover the proper way of supplying this philosophic theory with an empirical foundation, and it was reserved for Gegenbaur to achieve this. He first opened out the phylogenetic path which here, as in all morphological questions, leads most confidently to the goal. He showed that the primitive fishes (Figs. 249–251), the ancestors of all the Gnathostomes, still preserve permanently in the form of their skull the structure out of which the transformed skull of the higher Vertebrates, including man, has been evolved. He further showed that the branchial arches of the Selachii prove that their skull originally consisted of a large number of (at least nine or ten) provertebræ, and that the cerebral nerves that proceed from the base of the brain entirely confirm this. These cerebral nerves are (with the exception of the first and second pair, the olfactory and optic nerves) merely modifications of spinal nerves, and are essentially similar to them in their peripheral expansion. The comparative anatomy of these cerebral nerves, their origin and their expansion, furnishes one of the strongest arguments for the new vertebral theory of the skull.
Fig.342. Skeleton of the hand or fore foot of six mammals. I man, II dog, III pig, IV ox, V tapir, VI horse. Fig. 342—Skeleton of the hand or fore foot of six mammals. I man, II dog, III pig, IV ox, V tapir, VI horse. r radius, u ulna, a scaphoideum, b lunare, a triquetrum, d trapezium, e trapezoid, f capitatum, g hamatum, p pisiforme. 1 thumb, 2 index finger, 3 middle finger, 4 ring finger, 5 little finger. (From Gegenbaur.)
We have not space here to go into the details of Gegenbaur’s theory of the skull. I must be content to refer the reader to the great work I have mentioned, in which it is thoroughly established from the empirico-philosophical point of view. He has also given a comprehensive and up-to-date treatment of the subject in his Comparative Anatomy of the Vertebrates (1898). Gegenbaur indicates as original “cranial ribs,” or “lower arches of the cranial vertebræ,” at each side of the head of the Selachii (Fig. 334), the following pairs of arches: I and II, two lip-cartilages, the anterior (a) of which is composed of an upper piece only, the posterior (bc) from an upper and lower piece; III, the maxillary arches, also consisting of two pieces on each side—the primitive upper jaw (os palato-quadratum, o) and the primitive lower jaw (u); IV, the hyaloid bone (II); finally, V–X, six branchial arches in the narrower sense (III–VIII). From the anatomic features of these nine to ten cranial ribs or “lower vertebral arches” and the cranial nerves that spread over them, it is clear that the apparently simple cartilaginous primitive skull of the Selachii was originally formed from so many (at least nine) somites or provertebræ. The blending of these primitive segments into a single capsule is, however, so ancient that, in virtue of the law of curtailed heredity, the original division seems to have disappeared; in the embryonic development it is very difficult to detect it in isolated traces, and in some respects quite impossible. It is claimed that several (three to six) traces of provertebræ have been discovered in the anterior (pre-chordal) part of the Selachii-skull; this would bring up the number of cranial somites to twelve or sixteen, or even more.
In the primitive skull of man (Fig. 323) and the higher Vertebrates, which has been evolved from that of the Selachii, five consecutive sections are discoverable at a certain early period of development, and one might be induced to trace these to five primitive vertebræ; but these sections are due entirely to adaptation to
the five primitive cerebral vesicles, and correspond, like these, to a large number of metamera. That we have in the primitive skull of the mammals a greatly modified and transformed organ, and not at all a primitive formation, is clear from the circumstance that its original soft membranous form only assumes the cartilaginous character for the most part at the base and the sides, and remains membranous at the roof. At this part the bones of the subsequent osseous skull develop as external coverings over the membranous structure, without an intermediate cartilaginous stage, as there is at the base of the skull. Thus a large part of the cranial bones develop originally as covering bones from the corium, and only secondarily come into close touch with the primitive skull (Fig. 333). We have previously seen how this very rudimentary beginning of the skull in man is formed ontogenetically from the “head-plates,” and thus the fore end of the chorda is enclosed in the base of the skull. (Cf. Fig. 145 and pp. 138, 144, and 149.)
Fig.343-345. Arm and hand of three anthropoids. Fig. 343. Chimpanzee (Anthropithecus niger). Fig. 344. Veddah of Ceylon (Homo veddalis). Fig. 345. European (Homo mediterraneus). Figs. 343–345—Arm and hand of three anthropoids. Fig. 343—Chimpanzee (Anthropithecus niger). Fig. 344—Veddah of Ceylon (Homo veddalis). Fig. 345—European (Homo mediterraneus). (From Paul and Fritz Sarasin.)
The phylogeny of the skull has made great progress during the last three decades through the joint attainments of comparative anatomy, ontogeny, and paleontology. By the judicious and comprehensive application of the phylogenetic method (in the sense of Gegenbaur) we have found the key to the great and important problems that arise from the thorough comparative study of the skull. Another school of research, the school of what is called “exact craniology” (in the sense of Virchow), has, meantime, made fruitless efforts to obtain this result. We may gratefully acknowledge all that this descriptive school has done in the way of accurately describing the various forms and measurements of the human skull, as compared with those of other mammals. But the vast empirical material that it has accumulated in its extensive literature is mere dead and sterile erudition until it is vivified and illumined by phylogenetic speculation.
Virchow confined himself to the most careful analysis of large numbers of human skulls and those of anthropoid mammals. He saw only the differences between them, and sought to express these in figures.
Fig.346. Transverse section of a fish’s tail (from the tunny). Fig. 346—Transverse section of a fish’s tail (from the tunny). (From Johannes Müller.) a upper (dorsal) lateral muscles, a′, b′ lower (ventral) lateral muscles, d vertebral bodies, b sections of incomplete conical mantle, B attachment lines of the inter-muscular ligaments (from the side).
Without adducing a single solid reason, or offering any alternative explanation, he rejected evolution as an unproved hypothesis. He played a most unfortunate part in the controversy as to the significance of the fossil human skulls of Spy and Neanderthal, and the comparison of them with the skull of the Pithecanthropus (Fig. 283). All the interesting features of these skulls that clearly indicated the transition from the anthropoid to the man were declared by Virchow to be chance pathological variations. He said that the roof of the skull of Pithecanthropus (Fig. 335, 3) must have belonged to an ape, because so pronounced an orbital stricture (the horizontal constriction between the outer edge of the eye-orbit and the temples) is not found in any human being. Immediately afterwards Nehring showed in the skull of a Brazilian Indian (Fig. 335, 2), found in the Sambaquis of Santos, that this stricture can be even deeper in man than in many of the apes. It is very instructive in this connection to compare the roofs of the skulls (seen from above) of different primates. I have, therefore, arranged nine such skulls in Fig. 335, and reduced them to a common size.
We turn now to the branchial arches, which were regarded even by the earlier natural philosophers as “head-ribs.” (Cf. Figs. 167–170). Of the four original gill-arches of the mammals the first lies between the primitive mouth and the first gill-cleft. From the base of this arch is formed the upper-jaw process, which joins with the inner and outer nasal processes on each side, in the manner we have previously explained, and forms the chief parts of the skeleton of the upper jaw (palate bone, pterygoid bone, etc.) (Cf. p. 284.) 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. Figs. 169, 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 (Figs. 210, 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 (Figs. 248–259). 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 (Figs. 263–265).
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 (Fig. 257). Both the breast-fin and the belly-fin are flat oval paddles, in which we find a biserial cartilaginous skeleton (Fig. 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.[31]
[31] 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 (Fig. 248), Ganoids (Fig. 253), and Dipneusts (Fig. 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 (Fig. 337). We thus get the uniserial fin, which has been transmitted from the Selachii to the rest of the fishes (Fig. 338).
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.[32]
[32] 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 Fig. 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 Fig. 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 (Fig. 339 r and u) and humerus (h) of the higher Vertebrates.
Fig.347. Human skeleton. Fig. 348. Skeleton of the giant gorilla. Fig. 347—Human skeleton. (Cf. Figure 326.) Fig. 348—Skeleton of the giant gorilla. (Cf. Figure 209.)
In this way the five-toed foot of the Amphibia, which we first meet in the Carboniferous Stegocephala (Fig. 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 (Fig. 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 Figs. 260, 265, 270, 278–282, and 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 (Fig. 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 (Fig. 340) and of man (Fig. 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 (Fig. 342 a–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, Fig. 344) are midway between the chimpanzee (Fig. 343) and the European (Fig. 345). More considerable are the differences in structure and the proportions of the various parts between the different genera of anthropoid apes (Figs. 278–282); 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 p. 171.
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 (Fig. 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 cœlom-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. Figs. 124, 158–160, 222–224 mp). In the unarticulated stem-forms of the Chordonia, which we have called the Prochordonia, the two cœlom-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 (Figs. 124, 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 cœlom-pouches of the Acrania, Cyclostomes, and Selachii (Fig. 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 cœlom-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 (Fig. 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 (Fig. 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.[33]
[33] 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 cœlom-pouches. This also applies originally to the primary muscles of the limbs, as these too belong phylogenetically to the hyposoma. (Cf. Chapter XIV.)