Now I do not wish to conceal from you that many good zoologists believe that the vertebrate is descended from annelids; but for this and other reasons such a descent appears to me very improbable. It would seem far more natural to derive the vertebrate from some free swimming form like the schematic worm, whose largest nerve-cord lay on the dorsal surface because its branches ran to heavy muscles much used in swimming. Later the other nerve-cords degenerated, for such a degeneration of nerve-cords is not at all impossible or improbable. "No thoroughfare" is often written across paths previously followed by blood or nervous impulses, when other paths have been found more economical or effective.
But where did the notochord come from? I do not know. It always forms in the embryo out of the entoderm or layer which becomes the lining of the intestine. Now this is a very peculiar origin for cartilage, and the notochord is a very strange cartilage even if we have not made a mistake in calling it cartilage at all. My best guess would be that it is simply a thickened portion of the upper median surface of the intestine to keep the "b.a.l.l.s" of digesting nutriment or other hard particles in the intestine from "grinding"
against the nerve-cord as they are crowded along in the process of digestion. Once started its elasticity would be a great aid in swimming.
Professor Brooks has called attention to the fact that the higher a group stands in development, the longer its ancestors have maintained a swimming life. Thus we have noticed that the sponges were the first to settle; then a little later the ma.s.s of the coelenterates followed their example. But the etenophora, the nearest relatives of bilateral animals, have remained free swimming.
Then the flat worms and mollusks took to a creeping mode of life, while the annelids and vertebrates still swam. Then the annelids settled to the bottom and crept, and all their descendants remained creeping forms. The vertebrates alone remained swimming, and probably neither they nor their descendants ever crept until they emerged on the land, or as amphibia were preparing for land life. If this be true, it is a fact worthy of our most careful consideration. The swimming life would appear to be neither as easy nor as economical as the creeping. It is certainly hard to believe that food would not have been obtained with less effort and in greater abundance at the bottom than in the water above. The swimming life gave rise to higher and stronger forms; but did its maintenance give immediate advantage in the struggle for existence?
This is an exceedingly interesting and important question, and demands most careful consideration. But we shall be better prepared to answer it in a future lecture.
The period of development of mollusks, articulates, and vertebrates, is really one. They developed to a certain extent contemporaneously.
The development of vertebrates was slow, and they were the last to appear on the stage of geological history.
You must all have noticed that development, during this period, takes on a much more hopeful form than during that described in the last chapter. Then digestion and reproduction were dominant. Now muscle is of the greatest importance. If this fails of development, as in mollusks, the group is doomed to degeneration or at best stagnation. But we have seen the dawn of a still higher function. In insects and vertebrates the brain is becoming of importance, and absorbing more and more material. This is the promise of something vastly higher and better. Better sense-organs are appearing, fitted to aid in a wider perception of more distant objects. The vertebrate has discovered the right path; though a long journey still lies before it. The night is far spent, the day is at hand.
CHAPTER IV
VERTEBRATES: BACKBONE AND BRAIN
In tracing man"s ancestry from fish upward we ought properly to describe three or four fish, an amphibian, a reptile, and then take up the series of mammalian ancestors. But we have not sufficient time for so extended a study, and a simpler method may answer our purpose fairly well. Let us fix our attention on the few organs which still show the capacity of marked development, and follow each one of these rapidly in its upward course.
We must remember that there are changes in the vegetative organs.
The digestive and excretory systems improve. But this improvement is not for the sake of these vegetative functions. Brain and muscle demand vastly more fuel, and produce vastly more waste which must be removed. At almost the close of the series the reproductive system undergoes a modification which is almost revolutionary in its results. But we shall find that this modification is necessitated by the smaller amount of material which can be spared for this function; not by its increasing importance, still less its dominance for its own worth. The vertebrate is like an old Roman; everything is subordinated to mental and physical power. He is the world conqueror.
The important changes from fish upward affect the following organs: 1. The skeleton. A light, solid framework must be developed for the body. 2. The appendages start as fins, and end as the legs and arms of man. 3. The circulatory and respiratory systems developed so as to carry with the utmost rapidity and certainty fuel and oxygen to the muscular and nervous high-pressure engines. Or, to change the figure, they are the roads along which supplies and munitions can be carried to the army suddenly mobilized at any point on the frontier.
4. Above all, the brain, especially the cerebrum, the crown and goal of vertebrate structure. The improvement is now practically altogether in the animal organs of locomotion and thought. Still, among these animal organs, the lower systems will lead in point of time. The brain must to a certain extent wait for the skeleton.
1. The skeleton. The axial skeleton consists, in the lowest fish, of the notochord, a cylindrical unsegmented rod of cartilage running nearly the length of the body. This is surrounded by a sheath of connective tissue, at first merely membranous, later becoming cartilaginous or gristly. Pieces of cartilage extend upward over the spinal marrow, and downward around the great aortic artery, forming the neural and haemal arches. These unite with the ma.s.ses of cartilage surrounding the notochord to form cartilaginous vertebrae, which may be stiffened by an infiltration of carbonate of lime. The vertebral column of sharks has reached this stage. Then the cartilaginous vertebrae ossify and form a true backbone. I have described the process as if it were very simple. But only the student of comparative osteology can have any conception of the number of experiments which were tried in different groups before the definite mode of forming a bony vertebra was attained. At the same time the skull was developing in a somewhat similar manner. But the skull is far more complex in origin and undergoes far more numerous and important changes than the simpler vertebral column.
Into its history we have no time to enter.
And what shall we say of bone itself as a mere material or tissue, with its admirable lightness, compactness, and flawlessness. And every bone in our body is a triumph of engineering architecture. No engineer could better recognize the direction of strain and stress, and arrange his rods and columns, arches and b.u.t.tresses, to suitably meet them, than these problems are solved in the long bone of our thigh. And they must be lengthened while the child is leaping upon them. An engineer is justly proud if he can rebuild or lengthen a bridge without delaying the pa.s.sage of a single train. But what would he say if you asked him to rebuild a locomotive, while it was running even twenty miles an hour? And yet a similar problem had to be solved in our bodies.
But the vertebral column is not perfected by fish. The vertebrae with few exceptions are hollow in front and behind, biconcave; and between each two vertebrae there is a large cavity still occupied by the notochord. Thus these vertebrae join one another by their edges, like two shallow wine-gla.s.ses placed rim to rim. Only gradually is the notochord crowded out so that the vertebrae join by their whole adjacent surfaces. Even in highest forms, for the sake of mobility, they are united by washer-like disks of cartilage. Biconcave vertebrae persisted through the oldest amphibia, reptiles, and birds. But finally a firm backbone and skull were attained.
2. The appendages. Of these we can say but little. The fish has oar-like fins, attached to the body by a joint, but themselves unjointed. By the amphibia legs, with the same regions as our own and with five toes, have already appeared. The development of the leg out of the fin is one of the most difficult and least understood problems of vertebrate comparative anatomy. The legs are at first weak and scarcely capable of supporting the body. Only gradually do they strengthen into the fore- and hind-legs of mammals, or into the legs and wings of birds and old flying reptiles.
3. Changes in the circulatory and respiratory systems. The fish lives altogether in the water and breathes by gills, but the dipnoi among fishes breathes by lungs as well as gills. As long as respiration takes place by gills alone, the circulation is simple; the blood flows from the heart to the gills, and thence directly all over the body; the oxygenated blood from the gills does not return directly to the heart. But the blood from the lungs does return to the heart; and there at first mixes in the ventricle with the impure blood which has returned from the rest of the body. Gradually a part.i.tion arises in the ventricle, dividing it into a right and left half. Thus the two circulations of the venous blood to the lungs, and of the oxygenated blood over the body, are more and more separated until, in higher reptiles, they become entirely distinct.
As the animal came on land and breathed the air, more completely oxygenated blood was carried to the organs, and their activity was greatly heightened. As more and more heat was produced by the combustion in muscular and nervous tissues, and less was lost by conduction, the temperature of the body rose, and in birds and mammals becomes constant several degrees above the highest summer temperature of the surrounding air.
The changes in the brain affect mainly the large and small brain.
The cerebellum increases with the greater locomotive powers of the animal. But its development is evidently limited. The large brain, or cerebrum, is in fish hardly as heavy as the mid-brain; in amphibia the reverse is true. In higher recent reptiles the cerebrum would somewhat outweigh all the other portions of the brain put together. In mammals it extends upward and backward, has already in lower forms overspread the mid-brain, and is beginning to cover the small brain. But this was not so in the earliest mammals. Here the cerebrum was small, more like that of reptiles. But during the tertiary period the large brain began to increase with marvellous rapidity. It was very late in arriving at the period of rapid development, but it kept on after all the other organs of the body had settled down into comparative rest, perhaps retrogression.
We have given thus a rapid sketch in outline of the changes in the most characteristic systems between fish and mammals. Some of the changes which took place in mammals were along the same lines, but one at least is so new and unexpected that this highest cla.s.s demands more careful and detailed examination.
The mammal is a vertebrate. Hence all its organs are at their best.
But mammals stand, all things considered, at the head of vertebrates. The skeleton is firm and compact. The muscles are beautifully moulded and fitted to the skeleton so as to produce the greatest effect with the least ma.s.s and weight of tissue. The sense-organs are keen, and the eye and ear especially delicate, and fitted for perception at long range. Yet in all these respects they are surpa.s.sed by birds. As a mere anatomical machine the bird always seems to me superior to the mammal. It is not easy to see why it failed, as it has, to reach the goal of possibility of indefinite development and dominance in the animal world. Why he stopped short of the higher brain development I cannot tell. The fact remains that the mammal is pre-eminent in brain power, and that this gave him the supremacy.
But mammals came very late to the throne, and the probability of their ever gaining it must for ages have appeared very doubtful.
They seem to have been a fairly old group with a very slow early development. Reptiles especially, and even birds, were far more precocious than these slower and weaker forms which crept along the earth. But reptiles and birds, like many other precocious children, soon reached the limit of their development. They had muscle, the mammal brain and nerve; the mammal had the staying power and the future. Bitter and discouraging must have been the struggle of these feeble early mammals with their larger, swifter, and more powerful, reptilian relatives. And yet, perhaps, by this very struggle the mammal was trained to shrewdness and endurance.
The primitive mammals laid eggs like reptiles or birds. Only two genera, echidna and platypus, survive to bear witness of these old oviparous groups, and these only in New Zealand. These retain several old reptilian characteristics. Their lower position is shown also by the fact that the temperature of their bodies is, at least, ten degrees Fahrenheit below that of higher mammals. One of these carries the egg in a pouch on the ventral surface; the other, living largely in water, deposits its eggs in a nest in a burrow in the side of the bank of the stream.
After these came the marsupials. In these the eggs develop in a sort of uterus; but there is no placenta, in the sense of an organic connection between the embryo and the uterus of the mother. The young are at birth exceedingly small and feeble. The adult giant Kangaroo weighs over one hundred pounds; the young are at birth not as large as your thumb. They are placed by the mother in a marsupial pouch on her ventral surface, and here nourished till able to care for themselves.
Pardon a moment"s digression. The marsupials, except the opossum, are confined to Australia, and the oviparous mammals, or monotremes, to New Zealand. Formerly the marsupials, at least, ranged all over Europe and Asia, for we have indisputable evidence in their fossil remains. But they have survived only in this isolated area, and here apparently only because their isolation preserved them from the compet.i.tion with higher forms. If the Australian continent had not been thus early cut off from all the rest of the world, the only trace of both these lower groups would have been the opossum in America and certain peculiarities in the development of the egg in higher mammals. This shows us how much weight should be a.s.signed to the formerly popular argument of the "missing links." The wonder is not that so many links are missing, but that any of these primitive forms have come down to us. For we see here another proof of the fearful extermination of lower forms during the progress of life on the globe. It seems as if the intermediate forms were less common among these most recent animals than among the older types. This may not be true, for it is not easy to compare the gap between two mammals with that between two worms or insects, and mistakes are very easily made. But it seems as if extermination had done its work more ruthlessly among these highest forms than among their humbler and lower ancestors. I would not lay much weight on such an opinion; but, if true, it has a meaning and is worthy of study.
In higher, true, placental mammals the period of pregnancy is much longer, and the young are born in a far higher stage of development, or rather, growth. The stage of growth at which the young are born differs markedly in different groups. A new-born kitten is a much feebler, less developed being than a new-born calf. An embryonic appendage, the allantois, used in reptiles and birds for respiration, has here been turned to another purpose. It lays itself against the walls of the uterus, uterine projections interlock with those which it puts forth, and the blood of the mother circulates through a host of capillaries separated from those of the blood system of the embryo only by the thinnest membrane. This is the placenta, developed, in part from the allantois of the embryo, in part from the uterus of the mother. It is not a new organ, but an old one turned to better and fuller use. In these closely a.s.sociated systems of blood-vessels, nutriment and oxygen diffuse from the blood of the mother into that of the embryo, and thus rapid growth is a.s.sured. The importance and far-reaching effect of this new modification in the old reproductive system cannot be over-estimated. The internal intra-uterine development of the young, and the mammalian habit of suckling them, far more than any other factors, have made man what he is. Some explanation must be sought for such a fact.
We have already seen that any animal devotes to reproduction the balance between income and expenditure of nutriment. Now, the digestive system is here well developed, and the income is large.
But we have already noticed that, as animals grow larger, the ratio between the digestive surface and the ma.s.s to be supported grows continually smaller. On account of size alone the mammal has but a small balance. But the amount of expenditure is proportional to the ma.s.s and activity of the muscular and nervous systems. And the mammal is, and from the beginning had to be, an exceedingly active, energetic, and nervous animal. The income has increased, but the expenses have far outrun the increase. The mammal can devote but little to reproduction.
Moreover, it requires a large amount of material to form a mammalian egg, such as that of the monotreme. It requires indefinitely more nutriment to build a mammal than a worm, for the former is not only larger and more perfect at birth; it is also vastly more complicated. The embryonic journey has, so to speak, lengthened out immensely. One monotreme egg represents more economy and saving than a thousand eggs of a worm. Moreover, where the individuals are longer lived and the generations follow one another at longer intervals, the number of favorable variations and the possibility of conformity to environment through these is greatly lessened. In such a group it is of the utmost importance that every egg should develop; the destruction of a single one is a real and important loss to the species. It is not enough to produce such an egg; it must be most scrupulously guarded. Even the egg of the platypus is deposited in a nest in a hole in the bank, and the female Echidna carries the egg in a marsupial pouch until it develops.
Notice further that among certain species of fish, amphibia, and reptiles, the females carry the eggs in the body until the embryos or young are fairly developed. Viviparous forms are unknown by birds, probably because this mode of development is incompatible with flight, their dominant characteristic. Putting these facts together, what more probable than that certain primitive egg-laying mammals should have carried the eggs as long as possible in the uterus. The embryo under these conditions would be better nourished by a secretion of the uterine glands than by a very large amount of yolk. The yolk would diminish and the egg decrease in size, and thus the marsupial mode of development would have resulted. And, given the marsupial mode of development and an embryo possessing an allantois, it is almost a physiological necessity that in some forms at least a placenta should develop. That the placenta has resulted from some such process of evolution is proven by its different stages of development in different orders of mammals. And even the feeblest attachment of the allantois of the embryo to the wall of the uterus would be of the greatest advantage to the species.
This is not the whole explanation; other factors still undiscovered were undoubtedly concerned. But even this shows us that the internal development of the young and the habit of suckling them was a logical result of mammalian structure and position. The grand results of this change we shall trace farther on.
The changes from the lower true mammals to the apes are of great interest, but we can notice only one or two of the more important.
The prosimii, or "half apes," including the lemurs, are nearly all arboreal forms. Perhaps they were driven to this life by their more powerful compet.i.tors. The arboreal life developed the fingers and toes, and most of these end, not with a claw, but with a nail. The little group has much diversity of structure, and at present finds its home mainly in Madagascar; though in earlier times apparently occurring all over the globe. The brain is more highly developed than in the average mammal, but far inferior to that of the apes.
They have a fairly opposable thumb.
The highest mammals are the primates. Their characteristics are the following: Fingers and toes all armed with nails, the eyes comparatively near together and fully enclosed in a bony case. The cerebrum with well-developed furrows covers the other portions of the brain. There is but one pair of milk-glands, and these on the breast. The differences between hand and foot become most strongly marked by the "anthropoid" apes. These have become accustomed to an upright gait in their climbing; hence the feet are used for supporting the body and the hands for grasping. Both thumb and great toe are opposable; but the foot is a true foot, and the hand a true hand, in anatomical structure. The face, hands, and feet have mainly lost the covering of hair. They have no tail, or rather its rudiments are concealed beneath the skin. These include the gibbon, the orang, the gorilla, and the chimpanzee.
We can sum up the few attainments of mammals in a line. The lower forms attained the placental mode of embryonic development; the higher attained upright gait, hands and feet, and a great increase of brain. Anatomically considered these were but trifles, but the addition of these trifles revolutionized life on the globe. The princ.i.p.al anatomical differences between man and the anthropoid ape are the following: Man is a strictly erect animal. The foot of the ape is less fitted for walking on the ground, where he usually "goes on all fours." The skull is almost balanced on the condyles by which it articulates with the neck, and has but slight tendency to tip forward. The facial portion, nose and jaws, is less developed and retracted beneath the larger cranium or brain-case. This has greatly changed the appearance of the head. Protruding jaws and chin, even when combined with large cranium and brain, always give man the appearance of brutality and low intelligence.
The pelvis is broad and comparatively shallow. The legs, especially the thighs, are long. The foot is long and strong, and rests its lower surface, not merely the outer margin as in apes, on the ground. The elastic arch of the instep must be excepted in the above description, and adds lightness and swiftness to his otherwise slow gait. The great toe is short and generally not opposable. The muscles of the leg are heavy and the knee-joint has a very broad articulating surface. But the great result of man"s erect posture is that the hand is set free from the work of locomotion, and has become a delicate tactile and tool-using organ. The importance of this change we cannot over-estimate. The hand was the servant of the brain for trying all experiments. Had not our arboreal ancestors developed the hand for us we could never have invented tools nor used them if invented. And its reflex influence in developing the brain has been enormous. The arm is shorter and the hand smaller.
The brain is absolutely and relatively large, and its surface greatly convoluted. This gives place for a large amount of "gray matter," whose functions are perception, thought, and will. For this gray matter forms a layer on the outside of the brain.
Thus, even anatomically, man differs from the anthropoid apes. His whole structure is moulded to and by the higher mental powers, so that he is the "Anthropos" of the old Greek philosophers, the being who "turns his face upward." Yet in all these anatomical respects some of the apes differ less from him than from the lower apes or "half apes." And every one of these can easily be explained as the result of progressive development and modification. Whoever will deny the possibility or probability of man"s development from some lower form must argue on psychological, not on anatomical, grounds; and it grows clearer every day that even the former but poorly justify such a denial.
But it is interesting to note that no one ape most closely approaches man in all anatomical respects. Thus among the anthropoids the orang is perhaps most similar to man in cerebral structure, the chimpanzee in form of skull, the gorilla in feet and hands. No evolutionist would claim that any existing ape represents the ancestor of man. The anthropoids represent very probably the culmination of at least three distinct lines of development. But we must remember that in early tertiary times apes occurred all over Europe, and probably Asia, many degrees farther north than now. In those days, as later, the fauna and flora of northern climates were superior in vigor and height of development to that of Africa or Australia. It is thus, to say the least, not at all improbable that there existed in those times apes considerably, if not far, superior to any surviving forms. Whether the palaeontologist will find for us remains of such anthropoids is still to be seen.
But you will naturally ask, "Is there not, after all, a vast difference between the brain of man and that of the ape?" Let us examine this question as fully as our very brief time will allow.
Considerable emphasis used to be laid on the facial angle between a line drawn parallel to the base of the skull and one obliquely vertical touching the teeth and most prominent portion of the forehead. Now this angle is in man very large--from seventy-five to eighty-five degrees, or even more, and rarely falling below sixty-five degrees. But this angle depends largely on the protrusion of the jaws, and varies greatly in species of animals showing much the same grade of intelligence. In some not especially intelligent South American monkeys the facial angle amounts to about sixty-five degrees. In this respect the skull of a chimpanzee reminds us of a human skull of small cranial capacity and large jaws, in which the cranium has been pressed back and the jaws crowded forward and slightly upward.
The weight of the brain in proportion to that of the body has been considered as of great importance, and within certain limits this is undoubtedly correct. Thus, according to Leuret, the weight of the brain is to that of the whole body: In fish, 1:5,668; in reptiles, 1:1,320; in birds, 1:212; in mammals, 1:186. These figures give the averages of large numbers of observations and have a certain amount of value. But within the same cla.s.s the ratio varies extraordinarily. Thus the weight of the brain is to that of the whole body: In the elephant, 1:500; in the largest dogs, 1:305; in the cat, 1:156; in the rat, 1:76; in the chimpanzee, 1:50; in man, 1:36; in the field-mouse, 1:31; in the goldfinch, 1:24.
From this series it is evident that the relative weight of the brain is no index of the intelligence of the animal. Indeed if the brain were purely an organ of mind, there is no reason that it should be any larger in an elephant than in a mouse, provided they had the same mental capacity. As animals grow larger the weight of the brain, relatively to that of the body, decreases, and considering the size of man it is remarkable that it should form so large a fraction of his weight. Still the fraction in the chimpanzee is not so much smaller. It is still possible that this fraction is above the normal for the chimpanzee, for some of the observations may have been taken on animals which had died of consumption or some other wasting disease. I have not been able to find whether this possibility of error has been scrupulously avoided.
A fair idea of the size of the brain may be obtained by measuring the cranial capacity. This varies in man from almost one-hundred cubic inches to less than seventy. In the gorilla its average is perhaps thirty, in the orang and chimpanzee rather less, about twenty-eight. This is certainly a vast difference, especially when we remember that the gorilla far exceeds man in weight.
Le Bon tells us that of a series of skulls forty-five per cent, of the Australian had a cranial capacity of 1,200 to 1,300 c.c., while 46.7 per cent. of modern Parisian skulls showed a capacity of between 1,500 and 1,600 c.c. The skull of the gorilla contains about five hundred and seventy cubic centimetres. Broca found that the cranial capacity of 115 Parisian skulls, of probably the higher cla.s.ses from the twelfth century, averaged about 1,426 cubic centimetres, while ninety of those of the poorer cla.s.ses of the nineteenth century averaged about 1,484. His observations seemed to prove that there has been a steady increase in Parisian cranial capacity from the twelfth to the nineteenth century.
Turning to the actual weight of the brain, that of Cuvier weighed 64.5 ounces, and a few cases of weights exceeding 65 ounces have been recorded. The lowest limit of weight in a normal human brain has not yet been accurately determined. From 34 to 31 ounces have been a.s.signed by different writers. The brain of a Bush woman was computed by Marshall at 31.5 ounces, and weights of even 31 ounces have been recorded without any note to show that the possessors were especially lacking in intelligence. As Professor Huxley says in his "Man"s Place in Nature," a little book which I cannot too highly recommend to you all, "It may be doubted whether a healthy human adult brain ever weighed less than 31 or 32 ounces, or that the heaviest gorilla brain has ever exceeded 20 ounces. The difference in weight of brain between the highest and the lowest men is far greater, both relatively and absolutely, than that between the lowest man and the highest ape. The latter, as has been seen, is represented by 12 ounces of cerebral substance absolutely, or by 32:20 relatively. But as the largest recorded human brain weighed between 65 and 66 ounces, the former difference is represented by 33 ounces absolutely, or by 65:32 relatively."
But there is another characteristic of the brain which seems to bear a close relation to the degree of intelligence. The surface of the human brain is not smooth but covered with convolutions, with alternating grooves or sulci, which vastly increase its surface and thus make room for more gray matter. Says Gratiolett: "On comparing a series of human and simian brains we are immediately struck with the a.n.a.logy exhibited in the cerebral forms in all these creatures.