The Project Gutenberg eBook, The Whence and the Whither of Man, by John Mason Tyler
THE WHENCE AND THE
WHITHER OF MAN
A BRIEF HISTORY OF HIS ORIGIN AND DEVELOPMENT
THROUGH CONFORMITY TO ENVIRONMENT
Being the Morse Lectures of 1895
BY
JOHN M. TYLER
PROFESSOR OF BIOLOGY, AMHERST COLLEGE
New York
Charles Scribner's Sons
1896
Morse Lectures
1893—THE PLACE OF CHRIST IN
MODERN THEOLOGY. By Rev. A.M.
Fairbairn, D.D. 8vo, $2.50
1894—THE RELIGIONS OF JAPAN. By Rev.
William Elliot Griffis, D.D.
12mo, $2.00.
1895—THE WHENCE AND THE WHITHER OF
MAN. By Professor John M. Tyler.
12mo, $1.75.
TABLE OF CONTENTS
| [INTRODUCTION] | [ix] |
| [CHAPTER I] | |
| THE PROBLEM: THE MODE OF ITS SOLUTION | [1] |
| The question. — The two theories of man's origin. — The argumentpurely historical. — Means of tracing man's ancestry andhistory. — Classification. — Ontogenesis and Phylogenesis. | |
| [CHAPTER II] | |
| PROTOZOA TO WORMS: CELLS, TISSUES, AND ORGANS | [32] |
| Amœba: Its anatomy and physiology. — Development of the cell. — Hydra: The development of digestive and reproductive organs, and of tissues. — Forms intermediate between amœba and hydra: Magosphæra, volvox. — Embryonic development. — Turbellaria: Appearance of a body wall, of ganglion, and nerve-cords. | |
| [CHAPTER III] | |
| WORMS TO VERTEBRATES: SKELETON AND HEAD | [55] |
| Worms and the development of organs. — Mollusks: The externalprotective skeleton leads to degeneration or stagnation. — Annelidsand arthropods: The external locomotive skeleton leadsto temporary rapid advance, but fails of the goal. — Itsdisadvantages. — Vertebrates: The internal locomotive skeleton leadsto backbone and brain. — Reasons for their dominance. — The primitivevertebrate. | |
| [CHAPTER IV] | |
| VERTEBRATES: BACKBONE AND BRAIN | [81] |
| The advance of vertebrates from fish through amphibia and reptilesto mammals. — The development of skeleton, appendages, circulatoryand respiratory systems, and brain. — Mammals: The oviparousmonotremata. — Marsupials. — Placental mammals. — Development of theplacenta. — Primates. — Arboreal life and the development of thehand. — Comparison of man with the highest apes. — Recapitulation ofthe history of man's origin and development. — The sequence ofdominant functions. | |
| [CHAPTER V] | |
| THE HISTORY OF MENTAL DEVELOPMENT AND ITS SEQUENCE OF FUNCTIONS | [113] |
| Mode of investigation. — Intellect. — Sense-perceptions. — Association. — Inference and understanding. — Rational intelligence. — Modes of mentalor nervous action. — Reflex action, unconscious and comparativelymechanical. — Instinctive action: The actor is conscious, but guidedby heredity. — Intelligent action. — The actor is conscious, guided byintelligence resulting from experience or observation. — The willstimulated by motives. — Appetites. — Fear and other prudentialconsiderations. — Care for young and love of mates. — The dawn ofunselfishness. — Motives furnished by the rational intelligence:Truth, right, duty. — Recapitulation: The will, stimulated by everhigher motives, is finally to be dominated by unselfishness and loveof truth and righteousness. — These rouse the only inappeasablehunger, and are capable of indefinite development. — Strength ofthese motives. — Their complete dominance the goal of human development. | |
| [CHAPTER VI] | |
| NATURAL SELECTION AND ENVIRONMENT | [152] |
| The reversal of the sequence of functions leads to extermination,degeneration, or, rarely, to stagnation. — Natural selection becomesmore unsparing as we go higher. — Extinction. — Severity of thestruggle for life. — Environment one. — But lower animals come intovital relation with but a small part of it. — It consists of a myriadof forces, which, as acting on a given form, may be considered asone grand resultant. — Environment is thus a power making at firstfor digestion and reproduction, then for muscular strength andactivity, then for shrewdness, finally for unselfishness andrighteousness. — An ultimate "power, not ourselves, making forrighteousness," a personality. — Our knowledge of this personalitymay be valid, even though very incomplete. — Religion. — Conformity tothe spiritual in or behind environment is likeness to God. — Theconservative tendency in evolution. | |
| [CHAPTER VII] | |
| CONFORMITY TO ENVIRONMENT | [177] |
| Human environment. — The development of the family as the school ofman's training. — The family as the school of unselfishness andobedience. — The family as the basis of social life. — Society as anaid to conformity to environment by increasing intelligence andtraining conscience. — Mental and moral heredity. — Personalmagnetism. — Man's search for a king. — The essence ofChristianity. — Conformity to environment gives future supremacy, butoften at the cost of present hardship. — Conformity as obedience tothe laws of our being. — Environment best understood through thestudy of the human mind. — Productiveness and prospectiveness ofvital capital. — Faith. | |
| [CHAPTER VIII] | |
| MAN | [210] |
| Composed of atoms and molecules, hence subject to chemical andphysical laws. — As a living being. — As an animal. — As avertebrate. — As a mammal. — As a social being. — As a personal andmoral being. — The conflict between the higher and the lower inman. — As a religious being. — As hero. — He has not yetattained. — Future man. — He will utilize all his powers, dulysubordinating the lower to the higher. — The triumph of the commonpeople. | |
| [CHAPTER IX] | |
| THE TEACHINGS OF THE BIBLE | [241] |
| Subject of the Bible. — Man: Body, intellect, heart. — God:Law, sin, and penalty. — God manifested in Christ. — Salvation, the divinelife permeating man — Faith. — Prayer. — Hope. — The Church. — Thebattle. — The victory. — The crown. | |
| [CHAPTER X] | |
| PRESENT ASPECTS OF THE THEORY OF EVOLUTION | [278] |
| The struggle for existence. — Natural selection. — Correlation oforgans. — Fortuitous variation. — Origin of the fittest. — Nägeli'stheory: Initial tendency supreme. — Weismann and the Neo-Darwinians:Natural selection omnipotent. — The Neo-Lamarckians. — Comparison ofthe Neo-Darwinian and the Neo-Lamarckian views. — "Individuality" thecontrolling power throughout the life of the organism. — Transmissionof special effects of use and disuse. — Summary. | |
| CHART SHOWING SEQUENCE OF ATTAINMENTS AND OF DOMINANT FUNCTIONS | [309] |
| PHYLOGENETIC CHART OF THE ANIMAL KINGDOM | [310] |
| INDEX | [311] |
CHAPTERS: [Introduction], [I], [II], [III], [IV], [V], [VI], [VII], [VIII], [IX], [X], [Index]
FIGURES: [1], [2], [3], [4], [5], [6], [7], [8], [9], [10]
INTRODUCTION
In the year 1865 Professor Samuel Finley Breese Morse, to whom the world is indebted for the application of the principles of electro-magnetism to telegraphy, gave the sum of ten thousand dollars to Union Theological Seminary to found a lectureship in memory of his father, the Rev. Jedediah Morse, D.D., theologian, geographer, and gazetteer. The subject of the lectures was to have to do with "The relations of the Bible to any of the sciences." The ten chapters of this book correspond to ten lectures, eight of which were delivered as Morse Lectures at Union Theological Seminary during the early spring of 1895. The first nine chapters appear in form and substance as they were given in the lectures, except that Chapters VI. and VII. were condensed in one lecture. Chapter X. is new, and I have not hesitated to add a few paragraphs wherever the argument seemed especially to demand further evidence or illustration.
One of my friends, reading the title of these lectures, said: "Of man's origin you know nothing, of his future you know less." I fear that many share his opinion, although they might not express it so emphatically.
It would seem, therefore, to be in order to show that science is now competent to deal with this question; not that she can give a final and conclusive answer, but that we can reach results which are probably in the main correct. We may grant very cheerfully that we can attain no demonstration; the most that we can claim for our results will be a high degree of probability. If our conclusions are very probably correct, we shall do well to act according to them; for all our actions in life are suited to meet the emergencies of a probable but uncertain course of events.
We take for granted the probable truth of the theory of evolution as stated by Mr. Darwin, and that it applies to man as really as to any lower animal. At the same time it concerns our argument but little whether natural selection is "omnipotent" or of only secondary importance in evolution, as long as it is a real factor, or which theory of heredity or variation is the more probable.
If man has been evolved from simple living substance protoplasm, by a process of evolution, it will some day be possible to write a history of that process. But have we yet sufficient knowledge to justify such an attempt?
Before the history of any period can be written its events must have been accurately chronicled. Biological history can be written only when the successive stages of development and the attainments of each stage have been clearly perceived. In other words, the first prerequisite would seem to be a genealogical[1] tree of the animal kingdom. The means of tracing this genealogical tree are given in the first chapter, and the results in the second, third, and fourth chapters of this book.
Now, for some of the ancestral stages of man's development a very high degree of probability can be claimed. One of man's earliest ancestors was almost certainly a unicellular animal. A little later he very probably passed through a gastræa stage. He traversed fish, amphibian, and reptilian grades. The oviparous monotreme and the marsupial almost certainly represent lower mammalian ancestral stages. But what kind of fish, what species of amphibian, what form of reptiles most closely resembles the old ancestor? How did each of these ancestors look? I do not know. It looks as if our ancestral tree were entirely uncertain and we were left without any foundation for history or argument.
But the history of the development of anatomical details, however important and desirable, is not the only history which can be written, nor is it essential. It would be interesting to know the size of brain, girth of chest, average stature, and the features of the ancient Greeks and Romans. But this is not the most important part of their history, nor is it essential. The great question is, What did they contribute to human progress?
Even if we cannot accurately portray the anatomical details of a single ancestral stage, can we perhaps discover what function governed its life and was the aim of its existence? Did it live to eat, or to move, or to think? If we cannot tell exactly how it looked, can we tell what it lived for and what it contributed to the evolution of man?
Now, the sequence of dominant functions or aims in life can be traced with far more ease and safety, not to say certainty, than one of anatomical details. The latter characterize small groups, genera, families, or classes; while the dominant function characterizes all animals of a given grade, even those which through degeneration have reverted to this grade.
Even if I cannot trace the exact path which leads to the mountain-top, I may almost with certainty affirm that it leads from meadow and pasture through forest to bare rock, and thence over snow and ice to the summit; for each of these forms a zone encircling the mountain. Very similarly I find that, whatever genealogical tree I adopt, one sequence in the dominance of functions characterizes them all; digestion is dominant before locomotion and locomotion before thought.
And it is hardly less than a physiological necessity that it should be so. The plant can and does exist, living almost purely for digestion and reproduction, and the same is true of the lowest and most primitive animals. A muscular system cannot develop and do its work until some sort of a digestive system has arisen to furnish nutriment, any more than a steam-engine can run without fuel. And a brain is of no use until muscle and sense-organs have appeared.
This sequence of dominant functions,[2] of physiological dynasties, would seem therefore to be a fact. And our series of forms described in the second, third, and fourth chapters is merely a concrete illustration showing how this sequence may have been evolved. The substitution of other terms in the anatomical series there described—amœba, volvox, etc.—would not affect this result. By a change in the form of our history we have eliminated to a large extent the sources of uncertainty and error. And the dominant function of a group throws no little light on the details of its anatomy.
If we can be satisfied that ever higher functions have risen to dominance in the successive stages of animal and human development, if we can further be convinced that the sequence is irreversible, we shall be convinced that future man will be more and more completely controlled by the very highest powers or aims to which this sequence points. Otherwise we must disbelieve the continuity of history. But the germs of the future are always concealed in the history of the present. Hence—pardon the reiteration—if we can once trace this sequence of dominant functions, whose evolution has filled past ages, we can safely foretell something at least of man's future development.
The argument and method is therefore purely historical. Here and there we will try to find why and how things had to be so. But all such digressions are of small account compared with the fact that things were or are thus and so. And a mistaken explanation will not invalidate the facts of history.
The subject of our history is the development, not of a single human race nor of the movements of a century, but the development of animal life through ages. And even if our attempts to decipher a few pages here and there in the volumes of this vast biological history are not as successful as we could hope, we must not allow ourselves to be discouraged from future efforts. Even if our translation is here and there at fault, we must never forget the existence of the history. Some of the worst errors of biologists are due to their having forgotten that in the lower stages the germs of the higher must be present, even though invisible to any microscope. Our study of the worm is inadequate and likely to mislead us, unless we remember that a worm was the ancestor of man. And a biologist who can tell us nothing about man is neglecting his fairest field.
Conversely history and social science will rest on a firmer basis when their students recognize that many human laws and institutions are heirlooms, the attainments, or direct results of attainments, of animals far below man. We are just beginning to recognize that the study of zoölogy is an essential prerequisite to, and firm foundation for, that of history, social science, philosophy, and theology, just as really as for medicine. An adequate knowledge of any history demands more than the study of its last page. The zoölogist has been remiss in not claiming his birthright, and in this respect has sadly failed to follow the path pointed out by Mr. Darwin.
For palæontology, zoölogy, history, social and political science, and philosophy are really only parts of one great science, of biology in the widest sense, in distinction from the narrower sense in which it is now used to include zoölogy and botany. They form an organic unity in which no one part can be adequately understood without reference to the others. You know nothing of even a constellation, if you have studied only one of its stars. Much less can the study of a single organ or function give an adequate idea of the human body.
Only when we have attained a biological history can we have any satisfactory conception of environment. As we look about us in the world, environment often seems to us to be a chaos of forces aiding or destroying good and bad, fit and unfit, alike.
But our history of animal and human progress shows us successive stages, each a little higher than the preceding, and surviving, for a time at least, because more completely conformed to environment. If this be true, and it must be true unless our theory of evolution be false, higher forms are more completely conformed to their environment than lower; and man has attained the most complete conformity of all. Our biological history is therefore a record of the results of successive efforts, each attaining a little more complete conformity than the preceding. From such a history we ought to be able to draw certain valid deductions concerning the general character and laws of our environment, to discover the direction in which its forces are urging us, and how man can more completely conform to it.
If man is a product of evolution, his mental and moral, just as really as his physical, development must be the result of such a conformity. The study of environment from this standpoint should throw some light on the validity of our moral and religious creeds and theories. It would seem, therefore, not only justifiable, but imperative to attempt such a study.
Our argument is not directly concerned with modern theories of heredity, or variation, or with the "omnipotence" or secondary importance of natural selection. And yet Nägeli, and especially Weismann, have had so marked an influence on modern thought that we cannot afford to neglect their theories. We will briefly notice these in the closing chapter.
FOOTNOTES:
[1] See Phylogenetic Chart, p. [310].
[2] See condensed Chart of Development, etc., p. [309].
CHAPTER I
THE PROBLEM: THE MODE OF ITS SOLUTION
The story of a human life can be told in very few words. A youth of golden dreams and visions; a few years of struggle or of neglected opportunities; then retrospect and the end.
"We come like water, and like wind we go."
But how few of the visions are realized. Faust sums up the whole of life in the twice-repeated word versagen, renounce, and history tells a similar story. Terah died in Haran; Abraham obtained but a grave in the land promised him and his children; Jacob, cheated in marriage, bitterly disappointed in his children, died in exile, leaving his descendants to become slaves in the land of Egypt; and Moses, their heroic deliverer, died in the mountains of Moab in sight of the land which he was forbidden to enter. You may answer that it is no injury that the promise is too large, the vision too grand, to be fulfilled in the span of a single life, but must become the heritage of a race. But what has been the history of Abraham's descendants? A death-grapple for existence, captivity, and dispersion. Their national existence has long been lost.
Was there ever a nation of grander promise than Greece or Rome? But Greece died of premature old age, and Rome of rottenness begotten of sin. But each of them, you will say, left a priceless heritage to the immortal race. But if Greece and Rome and a host of older nations, of which History has often forgotten the very name, have failed and died, can anything but ultimate failure await the race? Is human history to prove a story told by an idiot, or does it "signify" something? Is the great march of humanity, which Carlyle so vividly depicts, "from the inane to the inane, or from God to God?"
This is the sphinx question put to every thinking man, and on his answer hangs his life. For according to that answer, he will either flinch and turn back, or expend every drop of blood and grain of power in urging on the march.
To this question the Bible gives a clear and emphatic answer. "God created man in his own image," and then, as if men might refuse to believe so astounding a statement, it is repeated, "in the image of God created he him." When, and by what mode or process, man was created we are not told. His origin is condensed almost into a line, his present and future occupy all the rest of the book. Whence we came is important only in so far as it teaches us humility and yet assures us that we may be Godlike because we are His handiwork and children, "heirs of God and joint heirs with Christ of a heavenly inheritance."
Now has Science any answer to this vital question? Perhaps. But this much is certain; it can foretell the future only from the past. Its answer to the question whither must be an inference from its knowledge as to whence we have come. The Bible looks mainly at the present and future; Science must at least begin with the study of the past. The deciphering of man's past history is the great aim of Biology, and ultimately of all Science. For the question of Man's past is only a part of a greater question, the origin of all living species.
We may say broadly that concerning the origin of species two theories, and only two, seem possible. The first theory is that every species is the result of an act of immediate creation. And every true species, however slightly it may differ from its nearest relative, represents such a creative act, and once created is practically unchangeable. This is the theory of immutability of species. According to the second theory all higher, probably all present existing, species are only mediately the result of a creative act. The first living germ, whenever and however created, was infused with power to give birth to higher species. Of these and their descendants some would continue to advance, others would degenerate. Each theory demands equally for its ultimate explanation a creative act; the second as much as, if not more than, the first. According to the first theory the creative power has been distributed over a series of acts, according to the second theory it has been concentrated in one primal creation. The second is the theory of the mutability of species, or, in general, of evolution, but not necessarily of Darwinism alone.
The first theory is considered by many the more attractive and hopeful. Now a theory need not be attractive, nor at first sight appear hopeful, provided only it is true. But let me call your attention to certain conclusions which, as it appears to me, are necessarily involved in it. Its central thought is the practical immutability of species. Each one of these lives its little span of time, for species are usually comparatively short-lived, grows possibly a very little better or worse, and dies. Its progress has added nothing to the total of life; its degeneration harmed no one, hardly even itself; it was doomed from the start. Progress there has been, in a sense. The Creator has placed ever higher forms on the globe. But all the progress lies in the gaps and distances between successive forms, not in any advance made, or victory won, by the species or individual. The most "aspiring ape," if ever there was such a being, remains but an ape. He must comfort himself with the thought that, while he and his descendants can never gain an inch, the gap between himself and the next higher form shall be far greater than that between himself and the lowest monkey.
And if this has been the history of thousands of other species, why should it not be true of man also? Who can wonder that many who accept this theory doubt whether the world is growing any better, or whether even man will ever be higher and better than he now is? Would it not be contrary to the whole course of past history, if you can properly call such a record a history, if he could advance at all? Now I have no wish to misrepresent this or any honestly accepted theory, but it appears to me essentially hopeless, a record not of the progress of life on the globe, but of a succession of stagnations, of deaths. I can never understand why some very good and intelligent people still think that the theory of the immediate creation of each species does more honor to the Creator and his creation than the theory of evolution. Evolution is a process, not a force. The power of the Creator is equally demanded in both cases; only it is differently distributed. And evolution is the very highest proof of the wisdom and skill of the Creator. It elevates our views of the living beings, must it not give a higher conception of Him who formed them?
The plant in its first stages shows no trace of flowers, but of leaves only. Later a branch or twig, similar in structure to all the rest, shortens. The cells and tissues which in other twigs turn into green leaves here become the petals and other organs of the rose or violet. Let us suppose for a moment that every rose and violet required a special act of immediate creation, would the springtime be as wonderful as now? Would the rose or violet be any more beautiful, or are they any less flowers because developed out of that which might have remained a common branch? The plant at least is glorified by the power to give rise to such beauty. And is not the creation of the seed of a violet or rose something infinitely grander than the decking of a flowerless plant with newly created roses? The attainment of the highest and most diversified beauty and utility with the fewest and simplest means is always the sign of what we call in man "creative" genius. Is not the same true of God? I think you all feel the force of the argument here.
There were at one time no flowering plants. The time came at last for their appearance. Which is the higher, grander mode of producing them, immediate creation of every flowering species, or development of the flower out of the green leaves of some old club moss or similar form? The latter seems to me at least by far the higher mode. And to have created a ground-pine which could give rise to a rose seems far more difficult and greater than to have created both separately. It requires more genius, so to speak. It gives us a far higher opinion of the ground-pine; does it disgrace the rose? We can look dispassionately at plants. The rose is still and always a rose, and the oak an oak, whatever its origin. And I believe that we shall all readily admit that evolution is here a theory which does the highest honor to the wisdom and power of the Creator. What if the animal kingdom is continually blossoming in ever higher forms? Does not the same reasoning hold true, only with added force? I firmly believe that we should all unhesitatingly answer, yes, could we but be assured that all men would everywhere and always believe that we, men, were the results of an immediate creative act.
But why do we so strenuously object to the application to ourselves of the theory of evolution? One or two reasons are easily seen. We have all of us a great deal of innate snobbery, we would rather have been born great than to have won greatness by the most heroic struggle. But is man any less a man for having arisen from something lower, and being in a fair way to become something higher? Certainly not, unless I am less a man for having once been a baby. It is only when I am unusually cross and irritable that I object to being reminded of my infancy. But a young child does not like to be reminded of it. He is afraid that some one will take him for a baby still. And the snob is always desperately afraid that some one will fail to notice what a high-born gentleman he is.
Now man can relapse into something lower than a brute; the only genuine brute is a degenerate man. And we all recognize the strength of tendencies urging us downward. Is not this the often unrecognized kern of our eagerness for some mark or stamp that shall prove to all that we are no apes, but men? It is not the pure gold that needs the "guinea stamp." If we are men, and as we become men, we shall cease to fear the theory of evolution. Now this is not the only, or perhaps the greatest, objection which men feel or speak against the theory. But I must believe that it has more weight with us than we are willing to admit.
But some say that the theory of immediate creation and immutability of species is the more natural and has always been accepted, while the theory of evolution is new and very likely to be as short-lived as many another theory which has for a time fascinated men only to be forgotten or ridiculed.
But the idea of evolution is as old as Hindu philosophy. The old Ionic natural philosophers were all evolutionists. So Aristophanes, quoting from these or Hesiod concerning the origin of things, says: "Chaos was and Night, and Erebus black, and wide Tartarus. No earth, nor air nor sky was yet; when, in the vast bosom of Erebus (or chaotic darkness) winged Night brought forth first of all the egg, from which in after revolving periods sprang Eros (Love) the much desired, glittering with golden wings; and Eros again, in union with Chaos, produced the brood of the human race." Here the formative process is a birth, not a creation; it is evolution pure and simple. "According to the ancient view," says Professor Lewis, "the present world was a growth; it was born, it came from something antecedent, not merely as a cause but as its seed, embryo or principium. Plato's world was a 'zoon,' a living thing, a natural production."
Furthermore, to the ancient writers of the Bible the idea of origin by birth from some antecedent form—and this is the essential idea of evolution—was perfectly natural. They speak of the "generations of the heavens and the earth" as of the "generations" of the patriarchs. The first book of the Bible is still called Genesis, the book of births. The writer of the ninetieth Psalm says, "Before the mountains were born, or ever thou hadst brought to birth the earth and the world." And what satisfactory meaning can you give to the words, "Let the earth bring forth," and "the earth brought forth," in immediate proximity to the words, "and God made," unless while the ultimate source was God's creative power, the immediate process of formation was one of evolution.
The Bible is big and broad enough to include both ideas, the human mind is prone to overestimate the one or the other. Traces, at least, of a similar mode of thought persisted by the Greek Fathers of the Church, and disappeared, if ever, with the predominance of Latin theology. To the oriental the idea of evolution is natural. The earth is to him no inert, resistant clod; she brings forth of herself.
But our ancestors lived on a barren soil beneath a forbidding sky. They were frozen in winter and parched in summer. Nature was to them no kind foster-mother, but a cruel stepmother, training them by stern discipline to battle with her and the world. They peopled the earth with gnomes and cobolds and giants, and their nymphs were the Valkyre. Their God was Thor, of the thunderbolt and hammer, and who yet lived in continual dread of the hostile powers of Nature. A Norse prophet or prophetess standing beside Elijah at Horeb would have bowed down before the earthquake or the fire; the oriental waited for the "still small voice." And we are heirs to a Latin theology grafted on to the Thor-worship of our pagan ancestors. The idea of a Nature producing beneficently and kindly at the word of a loving God is foreign to all our inherited modes of thought. And our views of the heart of Nature are about as correct as those of our ancestors were of God. A little more of oriental tendencies of thought would harm neither our theology nor our life.
What, then, is the biblical idea of Nature? God speaks to the earth, in the first chapter of Genesis, and the earth responds by "giving birth" to mountains and living beings. It is evidently no mere lifeless, inert clod, but pulsating with life and responsive to the divine commands. While yet a chaos it had been brooded over by the Divine Spirit. It is like the great "wheels within wheels," with rings full of eyes round about, which Ezekiel saw in his vision by the river Chebar. "When the living creatures went, the wheels went by them; and when the living creatures were lifted up from the earth, the wheels were lifted up. Whithersoever the spirit was to go, they went, thither was their spirit to go; and the wheels were lifted up over against them: for the spirit of the living creatures (or of life) was in the wheels." And above the living creatures was the firmament and the throne of God. So Nature may be material, but it is material interpenetrated by the divine; if you call it a fabric, the woof may be material but the warp is God. This view contains all the truth of materialism and pantheism, and vastly more than they, and it avoids their errors and omissions.
To the old metaphysical hypothesis of evolution Mr. Darwin gave a scientific basis. It had always been admitted that species were capable of slight variation and that this divergence might become hereditary and thus perhaps give rise to a variety of the parent species. But it was denied that the variation could go on increasing indefinitely, it seemed soon to reach a limit and stop. Early in the present century Lamarck had attempted to prove that by the use and disuse of organs through a series of generations a great divergence might arise resulting in new species. But the theory was crude, capable at best of but limited application, and fell before the arguments and authority of Cuvier. The times were not ripe for such a theory. Some fifty years later, Mr. Darwin called attention to the struggle for existence as a means of aggregating these slight modifications in a divergence sufficient to produce new species, genera, or families. His argument may be very briefly stated as follows:
1. There is in Nature a law of heredity; like begets like.
2. The offspring is never exactly like the parent; and the members of the second generation differ more or less from one another. This is especially noticeable in domesticated plants and animals, but no less true of wild forms. If the parent is not exactly like the other members of the species, some of its descendants will inherit its peculiarities enhanced, others diminished.
3. Every species tends to increase in geometrical progression. But most species actually increase in number very slowly, if at all. Now and then some insect or weed escapes from its enemies, comes under favorable food conditions, and multiplies with such rapidity that it threatens to ravage the country. But as it multiplies it furnishes an abundance of food for the enemies which devour it, or of food and place for the parasites in and upon it; and they increase with at least equal rapidity. Hence while the vanguard increases prodigiously in numbers, because it has outrun these enemies, the rear is continually slaughtered. And thus these plagues seem in successive generations to march across the continent.
And yet even they give but a faint idea of the reproductive powers of plants and animals. The female fish produces often many thousands, sometimes hundreds of thousands of eggs. Insects generally from a hundred to a thousand. Even birds, slowly as they increase, produce in a lifetime probably at least from twelve to twenty eggs. Now let us suppose that all these eggs developed, and all the birds lived out their normal period of life, and reproduced at the same rate. After not many centuries there would not be standing room on the globe for the descendants of a single pair.
Again, of the one hundred eggs of an insect let us suppose that only sixty develop into the first larval, caterpillar, stage. Of these sixty, the number of members of the species remaining constant, only two will survive. The other fifty-eight die—of starvation, parasites, or other enemies, or from inclement weather. Now which two of all shall survive? Those naturally best able to escape their enemies or to resist unfavorable influences; in a word, those best suited to their conditions, or, to use Mr. Darwin's words, "conformed to their environment."
Now if any individual has varied so as to possess some peculiarity which enables it even in slight degree to better escape its enemies or to resist unfavorable conditions, those of its descendants who inherit most markedly this peculiar quality or variation will be the most likely to escape, those without it to perish. If a form varies unfavorably, becomes for instance more conspicuous to its enemies, it will almost certainly perish. Thus favorable variations tend to increase and become more marked from generation to generation.
Now it has always been known that breeders could produce a race of markedly peculiar form or characteristics by selecting the individuals possessing this quality in the highest degree and breeding only from these. The breeder depends upon heredity, variation, and his selection of the individuals from which to breed. Similarly in nature new species have arisen through heredity, variation, and a selection according to the laws of nature of those varying in conformity with their environment. And this Mr. Darwin called natural, in contrast with the breeder's artificial, "selection," arising from the "struggle for existence," and resulting in what Mr. Spencer has called the "survival of the fittest."
Let us take a single illustration. Many of the species of beetles on oceanic islands have very rudimentary wings, or none at all, and yet their nearest relatives are winged forms on some neighboring continent. Mr. Darwin would explain the origin of these evidently distinct wingless species as follows: They are descended from winged ancestors blown or otherwise transported thither from the neighboring continent. But beetles are slow and clumsy fliers, and on these wind-swept islands those which flew most would be blown out to sea and drowned. Those which flew the least, and these would include the individuals with more poorly developed wings, would survive. There would thus be a survival in every generation of a larger proportion of those having the poorest wings, and destruction of those whose wings were strong, or whose habits most active. We have here a natural selection which must in time produce a species with rudimentary or aborted wings, just as surely as a human breeder, by artificial selection can produce such an animal as a pug or a poodle. These, like sin, are a human device; nature should not be held responsible for them.
But you may urge that the variation which would take place in a single generation would be, as a rule, too slight to be of any practical value to the animal, and could not be fostered by natural selection until greatly enhanced by some other means. Let us think a moment. If ten ordinary men run in a foot-race, the two foremost may lead by several feet. But if the number of runners be continually increased the finish will be ever closer until finally but an atom more wind or muscle or pluck would make all the difference between winning and losing the prize.
Similarly the million or more young of any species of insect in a given area may be said to run a race of which the prize is life, and the losing of which means literally death. The competition is inconceivably severe. How indefinitely slight will be the difference between the poorest of the 2,000 or 20,000 survivors and the best of the more than 900,000 which perish. The very slightest favorable variation may make all the difference between life and sure death. And yet these indefinitely slight variations continued and aggregated through ages would foot up an immense total divergence. The chalk cliffs of England have been built up of microscopic shells.
I have tried to give you very briefly a sketch of the essential points of Mr. Darwin's theory of evolution. But you should all read that marvel of patience, industry, clear insight, close reasoning, and grand honesty, the "Origin of Species." I have no time to give the arguments in its favor or to attempt to meet the objections which may arise in your minds. I ask you to believe only this much; that the theory is accepted with practical unanimity by scientific men because it, and it alone, furnishes an explanation for the facts which they discover in their daily work. And this is the strongest proof of the truth of any accepted theory.
Inasmuch as it is accepted by all scientists and largely by the public, it is certainly worth your while to know whether it has any bearing on the great moral and religious questions which you are considering. And in these lectures I shall take for granted, what some scientists still doubt, that man also is a product of evolution. For the weight of evidence in favor of this view is constantly increasing, and seems already to strongly preponderate. Also I wish in these lectures to grant all that the most ardent evolutionist can possibly claim. Not that I would lower man's position, but I have a continually increasing respect for the so-called "lower animals."
Now if the theory of evolution be true, and really only on this condition, life has had a history; and human history began ages before man's actual appearance on the globe, just as American history began to be fashioned by Anglo-Saxons, Danes, and Normans before they set foot even in England. We study history mainly to deduce its laws; and that knowing them we may from the past forecast the future, prepare for its emergencies, and avoid or wisely meet, its dangers. And we rely on these laws of history because they are the embodiment of ages of human experience.
Whatever be our system of philosophy we all practically rely on past experience and observation. Fire burns and water drowns. This we know, and this knowledge governs our daily lives, whatever be our theories, or even our ignorance, of the laws of heat and respiration. Now human history is the embodiment of the experience of the race; and we study it in the full confidence that, if we can deduce its laws, we can rely on racial experience certainly as safely as on that of the individual. Furthermore, if we can discover certain great movements or currents of human action or progress moving steadily on through past centuries, we have full confidence that these movements will continue in the future. The study of history should make us seers.
But the line of human progress is like a mountain road, veering and twisting, and often appearing to turn back upon itself, and having many by-roads, which lead us astray. If we know but a few miles of it we cannot tell whether it leads north or south or due west. But if from any mountain-top we can gain a clear bird's-eye view of its whole course, we easily distinguish the main road, its turns become quite insignificant, we see that it leads as directly as any engineering skill could locate it through the mountains to the fertile plains and rich harvests beyond.
Now our knowledge of the history of man covers so brief a period that we can scarcely more than hazard a guess as to the trend of human progress. Many of the most promising social movements are like by-roads which, at first less steep and difficult, end sooner or later against impassable obstacles. And even if there be a main line of march, advance seems to alternate with retreat, progress with retrogression. To illustrate further, the great waves rush onward only to fall back again, and we can hardly tell whether the tide is flowing or ebbing.
Yet already certain tendencies appear fairly clear. Governments tend to become democratic, if we define democracy as "any form of government in which the will of the people finds sovereign expression." The tendency of society seems to be toward furnishing all its members equality of opportunity to make the most of their natural endowments. But if we are convinced that these statements express even vaguely the tendency of human development in all its past history, we are confident that these tendencies will continue in the future for a period somewhat proportional to their time of growth in the past. If we are wise, we try to make our own lives and actions, and those of our fellows, conform to and advance them. Otherwise our lives will be thrown away.
But if the theory of evolution be true, human history is only the last page of the one history of all life. If we are to gain any adequate, true, extensive view of human progress, we must read more than this. We must take into account the history of man when he was not yet man. And if we believe in the future continuance of tendencies of a few centuries' growth, we shall rest assured of the permanence of tendencies which have grown and strengthened through the ages.
Our confidence in the results of historical study is therefore proportioned to the extent and thoroughness of the experience which they record, and to the time during which these laws can be proven to have held good. If I can make it even fairly probable that these laws, on obedience to which human progress and success seem to depend, are merely quoted from a grander code applicable to all life in all times, your confidence in them will be even greater. I trust I can prove to you that the animal kingdom has not drifted aimlessly at the mercy of every wind and tide and current of circumstance. I hope to show that along one line it has from the beginning through the ages held a steady course straight onward, and that deviation from this course has always led to failure or degeneration. From so vast a history we may hope to deduce some of the great laws of true success in life. Furthermore, if along this central line, at the head of which man stands, there always has been progress, we cannot doubt that future progress will be as certain, and perhaps far more rapid. In all the struggle of life we shall have the sure hope of success and victory; if not for ourselves still for those who shall come after us. "We are saved by hope." And we may be confident that this hope will never make us ashamed.
Finally, even from our present knowledge of the past progress of life we shall hope to catch hints at least that man's only path to his destined goal is the straight and narrow road pointed out in the Bible. If in this we are even fairly successful we shall find a relation and bond between the Bible and Science worthy of all consideration. And this is the only agreement which can ever satisfy us.
If I wished to bring before you a view of the development of man, I should best choose individuals or families from various periods of human history from the earliest times down to the present. I should try to tell you how they looked and lived. But if anyone should attempt to condense into three lectures such a history of even one line of the human race, you would probably think him insane. Even if he succeeded in giving a fairly clear view of the different stages, the successive stages would be so remote from one another, such vast changes would necessarily remain unnoticed or unexplained that you would hardly believe that they could have any genetic relation or belong to one developmental series.
But the history which I must attempt to condense for you is measured by ages, and the successive terms of the series will be indefinitely more remote from each other than the life and thoughts of Lincoln or Washington from those of our most primitive Aryan ancestor or of the rudest savage of the Stone Age. The series must appear exceedingly disconnected. Systems of organs will apparently spring suddenly into existence, and we shall have no time to trace their origin or earlier development. Even if we had an abundance of time many gaps would still remain; for the forms, which according to our theory must have occupied their place, have long since disappeared and left no trace nor sign. We have generally no conception at all of the amount of extermination and degeneration which have taken place in past ages.
I grant frankly that I do not believe that the forms which I have selected represent exactly the ancestors of man. They have all been more or less modified. I claim only that in the balance and relative development of their organic systems—muscular, digestive, nervous, etc.—they give us a very fair idea of what our ancestor at each stage must have been. But it is on this balance and relative development of the different systems, that is, whether an animal is more reproductive, digestive, or nervous, that my argument will in the main be based.
But if the older ancestors have so generally disappeared, and their surviving relatives have been so greatly modified, how can we make even a shrewd guess at the ancestry of higher forms? The genealogy of the animal kingdom has been really the study of centuries, although the earlier zoölogists did not know that this was to be the result of their labors. The first work of the naturalist was necessarily to classify the plants and animals which he found, and catalogue and tabulate them so that they might be easily recognized, and that later discovered forms might readily find a place in the system. Hypotheses and theories were looked upon with suspicion. "Even Linnæus," says Romanes, "was express in his limitations of true scientific work in natural history to the collecting and arranging of species of plants and animals." The question, "What is it?" came first; then, "How did it come to be what it is?" We are just awakening to the question, "Why this progressive system of forms, and what does it all mean?"
Let us experiment a little in forming our own classification of a few vertebrates. We see a bat flying through the air. We mistake it for a bird. But a glance at it shows that it is a mammal. It is covered with hair. It has fore and hind legs. Its wings are membranes stretched between the fingers and along the sides of the body. It has teeth. It suckles its young. In all these respects it differs from birds. It differs from mammals only in its wings. But we remember that flying squirrels have a membrane stretching along the sides of the body and serving as a parachute, though not as wings. We naturally consider the wings as a sort of after-thought superinduced on the mammalian structure. We do not hesitate to call it a mammal.
The whale makes us more trouble; it certainly looks remarkably like a fish. But the fin of its tail is horizontal, not vertical. Its front flippers differ altogether from the corresponding fins of fish; their bones are the same as those occurring in the forelegs of mammals, only shorter and more crowded together. Later we find that it has lungs, and a heart with four chambers instead of only two, as in fish. The vertebræ of its backbone are not biconcave, but flat in front and behind. And, finally, we discover that it suckles its young. It, too, is in all its deep-seated characteristics a mammal. It is fish-like only in characteristics which it might easily have acquired in adaptation to its aquatic life. And there are other aquatic mammals, like the seals, in which these characteristics are much less marked. Their adaptation has evidently not gone so far.
Now the first attempts resulted in artificial classifications, much like our grouping of bats with birds and whales with fish. All animals, like coral animals and starfishes, whose similar parts were arranged in lines radiating from a centre, were united as radiates, however much they might differ in internal structure and grade of organization. But this radiate structure proved again to be largely a matter of adaptation.
Practically all animals having a heavy calcareous shell were grouped with the snails and oysters as mollusks. But the barnacle did not fit well with other mollusks. Its shell was entirely different. It had several pairs of legs; and no mollusk has legs. The barnacle is evidently a sessile crab or better crustacean. Its molluscan characteristics were only skin-deep, evidently an adaptation to a mode of life like that of mollusks. The old artificial systems were based too much on merely external characteristics, the results of adaptation. When the internal anatomy had been thoroughly studied their groups had to be rearranged.
Reptiles and amphibia were at first united in one class because of their resemblance in external form. Our common salamanders look so much like lizards that they generally pass by this name. But the young salamander, like all amphibia, breathes by gills, its skeleton differs greatly from, and is far weaker than, that of the lizard, and there are important differences in the circulatory and other systems. Moreover, practically all amphibia differ from all reptiles in these respects. Evidently the fact that the alligator and many snakes and turtles (of which neither the young nor the embryos ever breathe by gills) live almost entirely in the water, is no better reason for classifying these with amphibia than to call a whale a fish, and not a mammal, because of its form and aquatic life.
When the comparative anatomy of fish, amphibia, and reptiles had been carefully studied it was evident that the amphibia stood far nearer the fish in general structure, while the higher reptiles closely approached birds. Then it was noticed that our common fish formed a fairly well-defined group, but that the ganoids, including the sturgeons, gar-pikes, and some others, had at least traces of amphibian characteristics. Such generalized forms, with the characteristics of the class less sharply marked, were usually by common consent placed at the bottom of the class. And this suited well their general structure, while in particular characteristics they were often more highly organized than higher groups of the same class.
The palæontologist found that the oldest fossil forms belonged to these generalized groups, and that more highly specialized forms—that is, those in which the special class distinctions were more sharply and universally marked—were of later geological origin. Thus the oldest fish were most like our present ganoids and sharks, though differing much from both. Our common teleost fish, like perch and cod, appeared much later. The oldest bird, the archæopteryx, had a long tail like that of a lizard, and teeth; and thus stood in many respects almost midway between birds and reptiles. And most of the earliest forms were "comprehensive," uniting the characteristics of two or more later groups. Thus as the classification became more natural, based on a careful comparison of the whole anatomy of the animals, its order was found to coincide in general with that of geological succession.
Then the zoölogist began to ask and investigate how the animal grew in the egg and attained its definite form. And this study of embryology brought to light many new and interesting facts. Agassiz especially emphasized and maintained the universality of the fact that there was a remarkable parallelism between embryos of later forms and adults of old or fossil groups. The embryos of higher forms, he said, pass through and beyond certain stages of structure, which are permanent in lower and older members of the same group.
You remember that the fin on the tail of a fish is as a rule bilobed. Now the backbone of a perch or cod ends at a point in the end of the tail opposite the angle between the two lobes, without extending out into either of them. In the shark it extends almost to the end of the upper lobe. Now we have seen that sharks and ganoids are older than cod. In the embryo of the cod or perch the backbone has, at an early stage, the same position as in the shark or ganoid; only at a later stage does it attain its definite position.
So Agassiz says the young lepidosteus (a ganoid fish), long after it is hatched, exhibits in the form of its tail characters thus far known only among the fossil fishes of the Devonian period. The embryology of turtles throws light upon the fossil chelonians. It is already known that the embryonic changes of frogs and toads coincide with what is known of their succession in past ages. The characteristics of extinct genera of mammals exhibit everywhere indications that their living representatives in early life resemble them more than they do their own parents. A minute comparison of a young elephant with any mastodon will show this most fully, not only in the peculiarities of their teeth, but even in the proportion of their limbs, their toes, etc. It may therefore be considered as a general fact that the phases of development of all living animals correspond to the order of succession of their extinct representatives in past geological times. The above statements are quoted almost word for word from Professor Agassiz's "Essay on Classification." The larvæ of barnacles and other more degraded parasitic crustacea are almost exactly like those of Crustacea in general. The embryos of birds have a long tail containing almost or quite as many vertebræ as that of archæopteryx. But most of these never reach their full development but are absorbed into the pelvis, or into the "ploughshare" bone supporting the tail feathers. Thus older forms may be said to have retained throughout life a condition only embryonic in their higher relatives. And the natural classification gave the order not only of geological succession but also of stages of embryonic development. Thus the system of classification improved continually, although more and more intermediate forms, like archæopteryx, were discovered, and certain aberrant groups could find no permanent resting-place.
But why should the generalized comprehensive forms stand at the bottom rather than the top of the systematic arrangement of their classes? Why should the system of classification coincide with the order of geologic occurrence, and this with the series of embryonic stages? Above all, why should the embryos of bird and perch form their tails by such a roundabout method? Why should the embryo of the bird have the tail of a lizard? No one could give any satisfactory explanation, although the facts were undoubted.
Mr. Darwin's theory was the one impulse needed to crystallize these disconnected facts into one comprehensible whole. The connecting link was everywhere common descent, difference was due to the continual variation and divergence of their ancestors. The classification, which all were seeking, was really the ancestral tree of the animal kingdom. Forms more generalized should be placed lower down on the ancestral tree, and must have had an earlier geological occurrence because they represented more nearly the ancestors of the higher. But this explains also the facts of embryonic development.
According to Mr. Darwin's theory all the species of higher animals have developed from unicellular ancestors. It had long been known that all higher forms start in life as single cells, egg and spermatozoon. And these, fused in the process of fertilization, form still a single cell. And when this single cell proceeds through successive embryonic stages to develop into an adult individual it naturally, through force of hereditary habit, so to speak, treads the same path which its ancestors followed from the unicellular condition to their present point of development. Thus higher forms should be expected to show traces of their early ancestry in their embryonic life. Older and lower adult forms should represent persistent embryonic stages of higher. It could not well be otherwise.
But the path which the embryo has to follow from the egg to the adult form is continually lengthening as life advances ever higher. From egg to sponge is, comparatively speaking, but a step; it is a long march from the egg to the earthworm; and the vertebrate embryo makes a vast journey. But embryonic life is and must remain short. Hence in higher forms the ancestral stages will often be slurred over and very incompletely represented. And the embryo may, and often does, shorten the path by "short-cuts" impossible to its original ancestor. Still it will in general hold true, and may be recognized as a law of vast importance, that any individual during his embryonic life repeats very briefly the different stages through which his ancestors have passed in their development since the beginning of life. Or, briefly stated, ontogenesis, or the embryonic development of the individual, is a brief recapitulation of phylogenesis, or the ancestral development of the phylum or group.
The illustration and proof of this law is the work of the embryologist. We have time to draw only one or two illustrations from the embryonic development of birds. We have already seen that the embryonic bird has the long tail of his reptilian ancestor. In early embryonic life it has gill-slits leading from the pharynx to the outside of the neck like those through which the water passes in the respiration of fish. The Eustachian tube and the canal of the external ear of man, separated only by the "drum," are nothing but such an old persistent gill-slit. No gills ever develop in these, but the great arteries run to them, and indeed to all parts of the embryo, on almost precisely the same general plan as in the adult fish. Only later is the definite avian circulation gradually acquired.
This law is even more strikingly illustrated in the embryonic development of the vertebral column and skull, if we had time to trace their development. And the development of the excretory system points to an ancestor far more primitive than even the fish. Our embryonic development is one of the very strongest evidences of our lowly origin.
Thus we have three sources of information for the study of animal genealogy. First, the comparative anatomy of all the different groups of animals; second, their comparative embryology; and third, their palæontological history. Each source has its difficulties or defects. But taken all together they give us a genealogical tree which is in the main points correct, though here and there very defective and doubtful in detail. The points in which we are left most in doubt in regard to each ancestor are its modes of life and locomotion, and body form. But these may temporarily vary considerably without affecting to any great extent the general plan of structure and the line of development of the most important deep-seated organs.
I have chosen a line composed of forms taken from the comparative anatomical series. All such present existing forms have probably been modified during the lapse of ages. But I shall try to tell you when they have diverged noticeably from the structure of the primitive ancestor of the corresponding stage. It is much safer for us to study concrete, actual forms than imaginary ones, however real may have been the former existence of the latter. And, after all, their lateral divergence is of small account compared with the great upward and onward march of life, to the right and left of which they have remained stationary or retrograded somewhat, like the tribes which remained on the other side of Jordan and never entered the Promised Land.
To recapitulate: Our question is the Whence and the Whither of man. To this question the Bible gives a clear and definite answer. Can Science also give an answer, and is this in the main in accord with the answer of Scripture? Science can answer the question only by the historical method of tracing the history of life in the past and observing the goal toward which it tends. If the evolution theory be true, the record of human achievement and progress forms only one short chapter in the history of the ages. If from the records of man's little span of life on the globe we can deduce laws of history on whose truth we can rely, with how much greater confidence and certainty may we rely on laws which have governed all life since its earliest appearance?—always provided that such can be found.
Our first effort must therefore be to trace the great line of development through a few of its most characteristic stages from the simplest living beings up to man. This will be our work in the three succeeding lectures. And to these I must ask you to bring a large store of patience. Anatomical details are at best dry and uninteresting. But these dry facts of anatomy form the foundation on which all our arguments and hopes must rest.
But if you will think long and carefully even of anatomical facts, you will see in and behind them something more and grander than they. You will catch glimpses of the divinity of Nature. Most of us travel threescore years and ten stone-blind in a world of marvellous beauty. Why does the artist see so much more in every fence-corner and on every hill-side than we, set face to face with the grandest landscapes? Primarily, I believe, because he is sympathetic, and looks on Nature as a comrade as near and dear as any human sister and companion. As Professor Huxley has said, "they get on rarely together." She speaks to the artist; to us she is dumb, and ought to be, for we are boorishly careless of her and her teachings.
Nature, to be known, must be loved. And though you have all the knowledge of a von Humboldt, and do not love her, you will never understand her or her teachings. You will go through life with her, and yet parted from her as by an adamantine wall.
I do not suppose that the author of the book of Job had ever studied geology, or mineralogy, or biology, but read him, and see whether this old prince of scientific heroes had loved, and understood, and caught the spirit of Nature. And what a grand, free spirit it was, and what a giant it made of him. I do not believe that Paul ever had a special course of anatomy or botany. But if he had not pondered long and lovingly on the structure of his body, and the germination of the seed, he never could have written the twelfth and fifteenth chapters of the first letter to the Corinthians. And time fails to speak of David and all the writers of the Psalms, and of those heroic souls misnamed the "Minor" Prophets.
Study the teachings of our Lord. How he must have considered the lilies of the field, and that such a tiny seed as that of the mustard could have produced so great an herb, and noticed and thought on the thorns and the tares and the wheat, and watched the sparrows, and pondered and wondered how the birds were fed. All his teaching was drawn from Nature. And all the study in the world could never have taught him what he knew, if it had not been a loving and appreciative study.
There is one strange and interesting passage in John's Gospel, xv. 1: "I am the true vine." My father used to tell us that the Greek word αληθινη, rendered true, is usually employed of the genuine in distinction from the counterfeit, the reality in distinction from the shadow and image. Is not this perhaps the clew to our Lord's use of natural imagery? Nature was always the presentation to his senses of the divine thought and purpose. He studied the words of the ancient Scripture, he found the same words and teachings clearly and concretely embodied in the processes of Nature. The interpretation of the Parable of the Sower was no mere play of fancy to him; it was the genuine and fundamental truth, deeper and more real than the existence of the sower, the soil, and the seed. The spiritual truth was the substance; the tangible soil and seed really only the shadow. And thus all Nature was to him divine.
We all of us need to offer the prayer of the blind man, "Lord, that our eyes may be opened." Let us learn, too, from the old heathen giant, Antæus, who, after every defeat and fall, rose strengthened and vivified from contact with his mother Earth. You will experience in life many a desperate struggle, many a hard fall. There is at such times nothing in the world so strengthening, healing, and life-giving as the thoughts and encouragements which Nature pours into the hearts and minds of her loving disciples. She will set you on your feet again, infused with new life, filled with an unconquerable spirit, with unfaltering courage, and an iron will to fight once more and win. In every battle her inspiring words will ring in your ears, and she will never fail you. We may not see her deepest realities, her rarest treasures of thought and wisdom; but if we will listen lovingly for her voice, we may be assured that she will speak to us many a word of cheer and encouragement, of warning and exhortation. For, to paraphrase the language of the nineteenth Psalm, "She has no speech nor language, her voice is not heard. But her rule is gone out throughout all the earth, and her words to the end of the world."
CHAPTER II
PROTOZOA TO WORMS: CELLS, TISSUES, AND ORGANS
The first and lowest form in our ancestral series is the amœba, a little fresh-water animal from 1/500 to 1/1000 of an inch in diameter. Under the microscope it looks like a little drop of mucilage. This semifluid, mucilaginous substance is the Protoplasm. Its outer portion is clear and transparent, its inner more granular. In the inner portion is a little spheroidal body, the nucleus. This is certainly of great importance in the life of the animal; but just what it does, or what is its relation to the surrounding protoplasm we do not yet know. There is also a little cavity around which the protoplasm has drawn back, and on which it will soon close in again, so that it pulsates like a heart. It is continually taking in water from the body, or the outside, and driving it out again, and thus aids in respiration and excretion. The animal has no organs in the proper sense of the word, and yet it has the rudiments of all the functions which we possess.
A little projection of the outer, clearer layer of protoplasm, a pseudopodium, appears; into this the whole animal may flow and thus advance a step, or the projection may be withdrawn. And this power of change of form is a lower grade of the contractility of our muscular cells. Prick it with a needle and it contracts. It recognizes its food even at a microscopic distance; it appears therefore to feel and perceive. Perhaps we might say that it has a mind and will of its own. It is safer to say that it is irritable, that is, it reacts to stimuli too feeble to be regarded as the cause of its reaction. It engulfs microscopic plants, and digests them in the internal protoplasm by the aid of an acid secretion. It breathes oxygen, and excretes carbonic acid and urea, through its whole body surface. Its mode of gaining the energy which it manifests is therefore apparently like our own, by combustion of food material.
1. AMŒBA PROTEUS. HERTWIG, FROM LEIDY.
ek, ectosarc; en, endosarc; N, food particles; n, nucleus; cv, contractile vesicle.
It grows and reaches a certain size, then constricts itself in the middle and divides into two. The old amœba has divided into two young ones, and there is no parent left to die, and death, except by violence, does not occur. But this absence of death in other rather distant relatives of the amœba, and probably in the amœba itself, holds true only provided that, after a series of self-divisions, reproduction takes place after another mode. Two rather small and weak individuals fuse together in one animal of renewed vigor, which soon divides into two larger and stronger descendants. We have here evidently a process corresponding to the fertilization of the egg in higher animals; yet there is no egg, spermatozoon, or sex.
It is a little mass of protoplasm containing a nucleus, and corresponds, therefore, to one of the cells, most closely to the egg-cell or spermatozoon of higher animals. If every living being is descended from a single cell, the fertilized egg, it is not hard to believe that all higher animals are descended from an ancestor having the general structure or lack of structure of the amœba.
But is the amœba really structureless? Probably it has an exceedingly complex structure, but our microscopes and technique are still too imperfect to show more than traces of it. Says Hertwig: "Protoplasm is not a single chemical substance, however complicated, but a mixture of many substances, which we must picture to ourselves as finest particles united in a wonderfully complicated structure." Truly protoplasm is, to borrow Mephistopheles' expression concerning blood, a "quite peculiar juice." And the complexity of the nucleus is far more evident than that of the protoplasm. Is protoplasm itself the result of a long development? If so, out of what and how did it develop? We cannot even guess. But the beginning of life may, apparently must, have been indefinitely farther back than the simplest now existing form. The study of the amœba cannot fail to raise a host of questions in the mind of any thoughtful man.
As we have here the animal reduced, so to speak, to lowest terms, it may be well to examine a little more closely into its physiology and compare it briefly with our own.
The amœba eats food as we do, but the food is digested directly in the internal protoplasm instead of in a stomach; and once digested it diffuses to all parts of the cell; here it is built up into compounds of a more complex structure, and forms an integral part of the animal body. The dead food particle has been transformed into living protoplasm, the continually repeated miracle of life. But it does not remain long in this condition. In contact with the oxygen from the air it is soon oxidized, burned up to furnish the energy necessary for the motion and irritability of the body. We are all of us low-temperature engines. The digestive function exists in all animals merely to bring the food into a soluble, diffusible form, so that it can pass to all parts of the body and be used for fuel or growth. In our body a circulatory system is necessary to carry food and oxygen to the cells and to remove their waste. For most of our cells lie at a distance from the stomach, lungs, and kidney. But in a small animal the circulatory system is often unnecessary and fails. Breathing and excretion take place through the whole surface of the body. The body of the frog is devoid of scales, so that the blood is separated from the surrounding water only by a thin membrane, and it breathes and excretes to a certain extent in the same way.
But another factor has to be considered. If we double each dimension of our amœba, we shall increase its surface four times, its mass eight-fold. Now the power of absorbing oxygen and excreting waste is evidently proportional to the excretory and respiratory surface, and much the same is true of digestion. But the amount of oxygen required, and of waste to be removed is proportional to the mass; for every particle of protoplasm requires food and oxygen, and produces waste. The particles of protoplasm in our new, larger amœba can therefore receive only half as much oxygen as before, and rid themselves of their waste only half as fast. There is danger of what in our bodies would be called suffocation and blood-poisoning. The amœba having attained a certain size meets this emergency by dividing into two small individuals, the division is a physical adaptation. But the many-celled animal cannot do this; it must keep its cells together. It gains the additional surface by folding and plaiting. And the complicated internal structure of higher animals is in its last analysis such a folding and plaiting in order to maintain the proper ratio between the exposed surface of the cells and their mass. And each cell in our bodies lives in one sense its own individual life, only bathed in the lymph and receiving from it its food and oxygen instead of taking it from the water.
But in another sense the cells of our body live an entirely different life, for they form a community. Division of labor has taken place between them, they are interdependent, correlated with one another, subject therefore to the laws of the whole community or organism. There are many respects in which it is impossible to compare Robinson Crusoe with a workman in a huge watch factory; yet they are both men.
Both the amœba and we live in the closest relation to our environment, and conformity to it is evidently necessary: life has been defined as the adjustment of internal relations to external conditions. We continually take food, use it for energy and growth, and return the simpler waste compounds. We are all of us, as Professor Huxley has said, "whirlpools on the surface of Nature;" when the whirl of exchange of particles ceases we die. We have seen that the fusion of two amœbæ results in a new rejuvenated individual. Why is a mixture of two protoplasms better than one? We can frame hypotheses; we know nothing about it. What of the mind of the amœba? A host of questions throng upon us and we can answer no one of them. All the great questions concerning life confront us here in the lowest term of the animal series, and appear as insoluble as in the highest.
Our second ancestral form is also a fresh-water animal, the hydra. This is a little, vase-shaped animal, which usually lives attached to grass-stems or sticks, but has the power to free itself and hang on the surface of the water or to slowly creep on the bottom. The mouth is at the top of the vase, and the simple, undivided cavity within the vase is the digestive cavity. Around the mouth is a ring of from four to ten hollow tentacles, whose cavities communicate freely underneath with the digestive cavity. Not only is food taken in at the mouth, but indigestible material is thrown out here. The animal may thus be compared to a nearly cylindrical sack with a circle of tubes attached to it above. The body consists of two layers of cells, the ectoderm on the outside and the entoderm lining the digestive cavity. Between these two is a structureless, elastic membrane, which tends to keep the body moderately expanded.
The food is captured by the tentacles; but digestion takes place only partially in the digestive cavity, for each surrounding cell engulfs small particles of food and digests them within itself. The entodermal cells behave in this respect much like a colony of amœbæ. The cells of both layers have at their bases long muscular fibrils, those of the ectodermal cells running longitudinally, those of the entoderm transversely. The animal can thus contract its body in both directions, or, if the body contain water and the transverse muscles are contracted, the pressure of the water lengthens the body and tends to extend the tentacles.
On the outside of the elastic membrane, just beneath the ectoderm, is a plexus or cobweb of nervous cells and fibrils. As in every nervous system, three elements are here to be found. 1. An afferent or sensory nerve-fibril, which under adequate stimulus is set in vibration by some cell of the epidermis or ectoderm, which is therefore called a sensory cell. 2. A central or ganglion cell, which receives the sensory impulse, translates it into consciousness, and is the seat of whatever powers of perception, thought, or will the animal possesses. This also gives rise to the efferent or motor impulses, which are conveyed by (3) a motor fibril to the corresponding muscle, exciting its contraction. But there are also nerve-fibrils connecting the different ganglion cells, so that they may act in unison. In the higher animals we shall find these central or ganglion cells condensed in one or a few masses or ganglia. But here they are scattered over the whole surface of the elastic supporting membrane.
The reproductive organs for the production of eggs and spermatozoa form little protuberances on the outside of the body below the tentacles. But hydra reproduces mostly by budding; new individuals growing out of the side of the old one, like branches from the trunk of a tree, but afterward breaking free and leading an independent life. There are special forms of cells besides those described; nettle cells for capturing food, interstitial cells, etc., but these do not concern us.
The distance from the single-celled amœba to hydra is vast, probably really greater than that between any other successive terms of our series. It may therefore be useful to consider one or two intermediate forms and the parallel embryonic stages of higher animals, and to see how the higher many-celled animal originates from the unicellular stage.
The amœba is an illustration of a great kingdom of similar, practically unicellular forms, which have played no unimportant part in the geological history of the globe. These are the protozoa. They include, first of all, the foraminifera, which usually have shells composed of carbonate of lime. These shells, settling to the bottom of the ocean, have accumulated in vast beds, and when compacted and raised above the surface, form chalk, limestone, or marble, according to the degree and mode of their hardening.
The protozoa include also the flagellata, a great, very poorly defined mass of forms occupying the boundary between the plant and animal kingdoms. They are usually unicellular, and their protoplasm is surrounded by a thin, structureless membrane. This prevents their putting out pseudopodia as organs of motion. Instead of these they have at one end of the ovoid or pear-shaped body a long, whiplash-like process or thread, a flagellum, and by swinging this they propel themselves through the water. These flagellata seem to have a rather marked tendency to form colonies. The first individual gives rise to others by division. But the division is not complete; the new individuals remain connected by the undivided rear end of the body. And such a colony may come to contain a large number of individuals.
2. MAGOSPHÆRA PLANULA. LANG, FROM HAECKEL.
Such a colony is represented by magosphæra. This is a microscopic globular form, discovered by Professor Haeckel on the coast of Norway. It consists of a large number of conical or pear-shaped individual cells, whose apices are turned toward the centre of the sphere. The cells are cemented together by a mucilaginous substance. Around their exposed larger ends, which form the surface of the sphere, are rows of flagella, by whose united action the colony rolls through the water. After a time each individual absorbs its flagella, the colony is broken up, the different individuals settle to the bottom, and each gives rise by division to a new colony. This group of cells may be considered as a colony or as an individual. Each term is defensible.
Volvox is also a spheroidal organism, composed often of a very large number of flagellated cells. But it differs from magosphæra in certain important respects. In the first place its cells have chlorophyl, the green coloring matter of plants. It lives therefore on unorganized fluid nourishment, carbon dioxide, nitrates, etc. It is a plant. But certain characteristics render it probable that it once lived on solid food and was therefore an animal. For where almost the sole difference between plants and animals is in the fluid or solid character of their food, a change from the one form into the other is not as difficult or improbable as one might naturally think. And plants and animals are here so near together, and travelling by roads so nearly parallel, that, even if volvox never was an animal, it might still serve very well to illustrate a stage through which animals must have passed.
The cells of volvox do not form a solid mass, but have arranged themselves in a single layer on the outer surface of the sphere. For a time, under favorable circumstances, volvox reproduces very much like magosphæra, and each cell can give rise to a new, many-celled individual. But after a time, especially under unfavorable circumstances, a new mode of reproduction appears. Certain cells withdraw from the outer layer into the interior of the colony. Here they are nourished by the other cells and develop into true reproductive elements, eggs and spermatozoa. Fertilization, that is, the union of egg and spermatozoon, or mainly of their nuclei, takes place; and the fertilized egg develops into a new organism. But the other cells, which have been all the time nourishing these, seem now to lack nutriment, strength, or vitality to give rise to a new colony. They die.
We find thus in volvox division of labor and corresponding difference of structure or differentiation; certain cells retain the power of fusing with other corresponding cells, and thus of rejuvenescence and of giving rise to a new organism. And these cells, forming a series through all generations, are evidently immortal like the protozoa. Natural death cannot touch them. These are the reproductive cells. The other cells nourish and transport them and carry on the work of excretion and respiration. These latter correspond practically to our whole body. We call them somatic cells. In volvox they are entirely subservient to, and exist for, the reproductive cells, and die when they have completed their service of these. The body is here only a vehicle for ova. Furthermore, in volvox there has arisen such an interdependence of cells that we can no longer speak of it as a colony. The colony has become an individual by division of labor and the resulting differentiation in structure.
But hydra gives us but a poor idea of the cœlenterata, to which kingdom it belongs. The higher cœlenterata have nearly or quite all the tissues of higher animals—muscular, connective, glandular, etc. And by tissues we mean groups of cells modified in form and structure for the performance of a special work or function. The protozoa developed the cell for all time to come, the cœlenterata developed the tissues which still compose our bodies. But they had them mainly in a diffuse form. A sort of digestive and reproductive system they did possess. But the work of arranging these tissues and condensing them into compact organs was to be done by the next higher group, the worms.
Let us now take a glance at certain stages of embryonic development which correspond to these earliest ancestral forms. We should expect some such correspondence from the fact already stated that the embryonic development of the individual is a brief recapitulation of the ancestral development of the species or larger group. The egg of the lowest vertebrate, amphioxus, shows these changes in a simple and apparently primitive form.
3. IMMATURE EGG-SHELL FROM OVARY OF ECHINODERM. HATSCHEK, FROM HERTWIG.
The fertilized egg of any animal consists of a single cell, a little mass of protoplasm containing a nucleus and surrounded by a structureless membrane. The egg is globular. The nucleus undergoes certain very peculiar, still but little understood, changes and divides into two. The protoplasm also soon divides into two masses clustering each around its own nucleus. The plane of division will be marked around the outside by a circular furrow, but the cells will still remain united by a large part of the membrane which bounds their adjacent, newly formed, internal faces.
Let us suppose that the egg lay so that the first plane of division was vertical and extending north and south. Each cell or half of the egg will divide into two precisely as before. The new plane of division will be vertical, but extending east and west. Each plane passes through the centre of the egg, and the four cells are of the same form and size, like much-rounded quarters of an orange. The third plane will lie horizontal or equatorial, and will divide each of these quarters into an upper and lower octant. The cells keep on dividing rapidly, the eight form sixteen, then thirty-two, etc. The sharp angle by which the cells met at the centre has become rounded off, and has left a little space, the segmentation cavity, filled with fluid in the middle of the embryo. The cells continue to press or be crowded away from the centre and form a layer one cell deep on the surface of the sphere.
This embryo, resembling a hollow rubber ball filled with fluid, is called a blastosphere. It corresponds in structure with the fully developed volvox, except, of course, in lacking reproductive cells.
4. GASTRULA. HATSCHEK, FROM HERTWIG.
Outer layer is the ectoderm; inner layer, the entoderm; internal cavity, the archenteron; mouth of cavity, blastopore.
If the rubber ball has a hole in it so that I can squeeze out the water, I can thrust the one-half into the other, and change the ball into a double-walled cup. A similar change takes place in the embryo. The cells of the lower half of the blastosphere are slightly larger than those of the upper half. This lower hemisphere flattens and then thrusts itself, or is invaginated, into the upper hemisphere of smaller cells and forms its lining. This cup-shaped embryo is called the gastrula. The cup deepens somewhat and becomes ovoid. Take a boiled egg, make a hole in the smaller end and remove the yolk, and you have a passable model of a gastrula. The shell corresponds to the ectoderm or outer layer of smaller cells; the layer of "white" represents the entoderm or lining of larger cells. The space occupied by the yolk corresponds to the archenteron or primitive digestive cavity; and the opening at the end to the primitive mouth or blastopore. Ectoderm and entoderm unite around the mouth. Both the blastosphere and gastrula often swim freely by flagella.
You can hardly have failed to notice how closely the gastrula corresponds to a hydra, and many facts lead us to believe that the still earlier ancestor of the hydra was free swimming, and that the tentacles are a later development correlated with its adult sessile life. Yet we must not forget that the hydra is even now not quite sessile, it moves somewhat. And our ancestor was almost certainly a free swimming gastræa, or hypothetical form corresponding in form and structure to the gastrula. The ancestor of man never settled down lazily into a sessile life.
But how is an adult worm or vertebrate formed out of such a gastrula? To answer this would require a course of lectures on embryology. But certain changes interest us. Between the ectoderm and entoderm of the gastrula, in the space occupied by the supporting membrane of hydra, a new layer of cells, the mesoderm, appears. This has been produced by the rapid growth and reproduction of certain cells of the entoderm which have migrated, so to speak, into this new position. In higher forms it becomes of continually greater importance, until finally nearly all the organs of the body develop from it. In our bodies only the lining of the mid-intestine and of its glands has arisen from the entoderm. And only the epidermis, or outer layer of our skin, and the nervous system and parts of our sense-organs have arisen from the ectoderm. But our mid-intestine is still the greatly elongated archenteron of the gastrula.
We may therefore compare the hydra or gastrula to a little portion of the lining of the human mid-intestine covered with a little flake of epidermis. This much the hydra has attained. But our bones and muscles and blood-vessels all come from the mesoderm by folding, plaiting, and channelling, and division of labor resulting in differentiation of structure. Of all true mesodermal structures the hydra has actually none, but in the ectodermal and entodermal cells he has the potentiality of them all. We must now try to discover how these potentialities became actualities in higher forms.
The third stage in our ancestral series is the turbellarian. This is a little, flat, oval worm, varying greatly in size in different species, and found both in fresh and salt water. Some would deny that this worm belonged in our series at all. But, while doubtless considerably modified, it has still retained many characteristics almost certainly possessed by our primitive bilateral ancestor. The different parts of hydra were arranged like those of most flowers, around one main vertical axis; it was thus radiate in structure, having neither front nor rear, right nor left side. But our little turbellaria, while still without a head, has one end which goes first and can be called the front end. The upper or dorsal surface is usually more colored with pigment cells than the lower or ventral surface, on which is the mouth. It has also a right and left side. It is thus bilateral.
The gastræa swam by cilia, little eyelash-like processes which urge the animal forward like a myriad of microscopic oars. In our bodies they are sometimes used to keep up a current, e.g., to remove foreign particles from the lungs. The turbellaria is still covered with cilia, probably an inheritance from the gastræa; for, while in smaller forms they may still be the principal means of locomotion, in larger ones the muscles are beginning to assume this function and the animal moves by writhing. The bilateral symmetry has arisen in connection with this mode of locomotion and is thus a mark of important progress.
In the turbellaria we find for the first time a true body-wall distinct from underlying organs. The outer layer of this is a ciliated epithelium or layer of cells. Under this an elastic membrane may occur. Then come true body muscles, running transversely, longitudinally and dorso-ventrally. Between the external transverse and the internal longitudinal layers we often find two muscular layers whose fibres run diagonally. The body is well provided with muscles, but their arrangement is still far from economical or effective.
5. TURBELLARIAN. LANG.
va and ha, front and rear branches of gastro-vascular cavity; ph, pharynx. The dark oval with fine branches represents the nervous system.
Within the body-wall is the parenchym. This is a spongy mass of connectile tissue in which the other organs are embedded. The mouth lies in the middle, or near the front of the ventral surface. The intestine varies in form, but is provided with its own layers of longitudinal and transverse muscles, and usually has paired pouches extending out from it into the body parenchym. These seem to distribute the dissolved nutriment; hence the whole cavity is still often called a gastro-vascular cavity as serving both digestion and circulation. There is no anal opening, but indigestible material is still cast out through the mouth.
The animal can gain sufficient oxygen to supply its muscles and nerves, which are the principal seats of combustion, through the external surface. It has, therefore, no special respiratory organs. But the waste matter of the muscles cannot escape so easily, for these are becoming deeper seated. Hence we find an excretory system consisting of two tubes with many branches in the parenchym, and discharging at the rear end of the body. This again is a sign that the muscles are becoming more important, for the excretory system is needed mainly to remove their waste. These tubes maybe only greatly enlarged glands of the skin.
The nervous system consists of a plexus of fibres and cells, the cells originating impulses and the fibres conveying them. But this much was present in hydra also. Here the front end of the body goes foremost and is continually coming in contact with new conditions. Here the lookout for food and danger must be kept. Hence, as a result of constant exercise, or selection, or both, the nerve-plexus has thickened at this point into a little compact mass of cells and fibres called a ganglion. And because this ganglion throughout higher forms usually lies over the œsophagus, it is called the supra-œsophogeal ganglion. This is the first faint and dim prophecy of a brain, and it sends its nerves to the front end of the body. But there run from it to the rear end of the body four to eight nerve-cords, consisting of bundles of nerve-threads like our nerves, but overlaid with a coating of ganglion cells capable of originating impulses. These cords are, therefore, like the plexus from which they have condensed, both nerves and centres; differentiation has not gone so far as at the front of the body. Sense organs are still very rudimentary. Special cells of the skin have been modified into neuro-epithelial cells, having sensory hairs protruding from them and nerve-fibrils running from their bases.
In a very few turbellaria we find otolith vesicles. These are little sacks in the skin, lined with neuro-epithelial cells and having in the middle a little concretion of carbonate of lime hung on rather a stiffer hair, like a clapper in a bell. Such organs serve in higher animals as organs of hearing, for the sensory hairs are set in vibration by the sound-waves. It is quite as probable that they here serve as organs for feeling the slightest vibrations in the surrounding water, and thus giving warning of approaching food or danger. The animal has also eyes, and these may be very numerous. They are not able to form images of external objects, but only of perceiving light and the direction of its source. A little group of these eyes lies directly over the brain, near the front end of the body; the others are distributed around the front or nearly the whole margin of the body.
6. CROSS-SECTION OF TURBELLARIAN. HATSCHEK, FROM JIJIMA.
e, external skin; rm, lateral muscles; la and li, longitudinal muscles; mdv, dorso-ventral muscles; pa, parenchyma; h, testicle; ov, oviduct; dt, yolk-gland; n, ventral nerve; i, gastro-vascular cavity.
The turbellaria, doubtless, have the sense of smell, although we can discover no special olfactory organ. This sense would seem to be as old as protoplasm itself.
This distribution of the eyes around a large portion of the margin, and certain other characteristics of the adult structure and of the embryonic development, are very interesting, as giving hints of the development of the turbellaria from some radiate ancestor. The mouth is in a most unfavorable position, in or near the middle of the body, rarely at the front end, as the animal has to swim over its food before it can grasp it. The animal only slowly rids itself of old disadvantageous form and structure and adapts itself completely to a higher mode of life.
By far the most highly developed system in the body is the reproductive. It is doubtful whether any animal, except, perhaps, the mollusk, has as complicated and highly developed reproductive organs. By markedly higher forms they certainly grow simpler.
And here we must notice certain general considerations. We found that reproduction in the amœba could be defined as growth beyond the limit normal to the individual. This form of growth benefits especially the species. The needs and expenses of the individual will therefore first be met and then the balance be devoted to reproduction. Now the income of the animal is proportional to its surface, its expense to its mass, and activity. And the ratio of surface to mass is most favorable in the smallest animals.[3] Hence, smaller animals, as a rule, increase faster than larger ones; and this is only one illustration of the fact that great size in an animal is anything but an unmixed advantage to its possessor. But muscles and nerves are the most expensive systems; here most of the food is burned up. Hence energetic animals have a small balance remaining. Now the turbellarian is small and sluggish, with a fair digestive system. With a great amount of nutriment at its disposal the reproductive system came rapidly to a high development, and relatively to other organs stands higher than it almost ever will again.
It is only fair to state that good authorities hold that so primitive an animal could not originally have had so highly developed a system, and that this characteristic must be acquired, not ancestral.
That certain portions of it may be later developments may be not only possible but probable. But anyone who has carefully studied the different groups of worms, will, I think, readily grant that in the stage of these flat worms reproduction was the dominant function, which had most nearly attained its possible height of development. From this time on the muscular and nervous systems were to claim an ever-increasing share of the nutriment, and the balance for reproduction is to grow smaller.
At the close of this lecture I wish to describe very briefly a hypothetical form. It no longer exists; perhaps it never did. But many facts of embryology and comparative anatomy point to such a form as a very possible ancestor of all forms higher than flat worms, viz., mollusks, arthropods, and vertebrates.
It was probably rather long and cylindrical, resembling a small and short earthworm in shape. The skin may have been much like that of turbellaria. Within this the muscles run in only two-directions—longitudinally and transversely. Between these and the intestine is a cavity—the perivisceral cavity—like that of our own bodies, but filled with a nutritive fluid like our lymph. This cavity seems to have developed by the expansion and cutting off of the paired lateral outgrowths of the digestive system of some old flat worm. But other modes of development are quite possible. The intestine has now an anal opening at or near the rear end of the body. The food moves only from front to rear, and reaches each part always in a certain condition. Digestion proper and absorption have been distributed to different cells, and the work is better done. Three portions can be readily distinguished: fore-intestine with the mouth, mid-intestine, as the seat of digestion and absorption, and hind-intestine, or rectum, with the anal opening. The front and hind-intestine are lined with infolded outer skin.
The nervous system consists of a supra-œsophageal ganglion with four posterior nerve-cords—one dorsal, two lateral, and one (or perhaps two) ventral. There were probably also remains of the old plexus, but this is fast disappearing. The excretory system consists of a pair of tubes discharging through the sides of the body-wall, and having each a ciliated, funnel-shaped opening in the perivisceral cavity. These have received the name of nephridia. Through these also the eggs and spermatozoa are discharged. The reproductive organs are modified patches of the peritoneum, or lining of the perivisceral cavity.
The number of muscles or muscular layers has been reduced in this animal. But such a reduction in the number of like parts in any animal is a sign of progress. And the longitudinal muscles have increased in size and strength, and the animal moves by writhing. Such a worm has the general plan of the body of the higher forms fairly well, though rudely, sketched. Many improvements will come, and details be added. But the rudiments of the trunk of even our own bodies are already visible. Head, in any proper sense of the term, and skeleton are still lacking; they remain to be developed.
And yet, taking the most hopeful view possible concerning the animal kingdom, its prospects of attaining anything very lofty seem at this point poor. Its highest representative is a headless trunk, without skeleton or legs. It has no brain in any proper sense of the word, its sense-organs are feeble; it moves by writhing. Its life is devoted to digestion and reproduction. Whatever higher organs it has are subsidiary to these lower functions. And yet it has taken ages on ages to develop this much. If this is the highest visible result of ages on ages of development, what hope is there for the future? Can such a thing be the ancestor of a thinking, moral, religious person, like man? "That is not first which is spiritual, but that which is natural (animal, sensuous); and afterward that which is spiritual." First, in order of time, must come the body, and then the mind and spirit shall be enthroned in it. The little knot of nervous material which forms the supra-œsophageal ganglion is so small that it might easily escape our notice; but it is the promise of an infinite future. The atom of nervous power shall increase until it subdues and dominates the whole mass.
FOOTNOTES:
CHAPTER III
WORMS TO VERTEBRATES: SKELETON AND HEAD
In tracing the genealogy of any American family it is often difficult or impossible to say whether a certain branch is descended from John Oldworthy or his cousin or second cousin. In the latter cases to find the common ancestor we must go back to the grandfather or great-grandfather. The same difficulty, but greatly enhanced, meets us when we try to make a genealogical tree of the animal kingdom. Thus it seems altogether probable that all higher forms are descended from an ancestor of the same general structure and grade of organization as the turbellaria, although probably free swimming, and hence with somewhat different form and development, especially of the muscular system. It seems to me altogether probable that all, except possibly Mollusca, are descended from a common ancestor closely resembling the schematic worm last described. Some would, however, maintain that they diverged rather earlier than even the turbellaria; others after the schematic worm, if such ever existed. As far as our argument is concerned it makes little difference which of these views we adopt.
From our turbellaria, or possibly from some even more primitive ancestor, many lines diverged. And this was to be expected. The cœlenterata, as we saw in hydra, had developed rude digestive and reproductive systems. The higher groups of this kingdom had developed all, or nearly all, the tissues used in building the bodies of higher animals—muscular, reproductive, connectile, glandular, nervous, etc. But these are mostly very diffuse. The muscular fibrils of a jelly-fish are mostly isolated or parallel in bands, rarely in compact well-defined bundles. The tissues have generally not yet been moulded into compact masses of definite form. There are as yet very few structures to which we can give the name of organs. To form organs and group them in a body of compact definite form was the work pre-eminently of worms. The material for the building was ready, but the architecture of the bilateral animal was not even sketched. And different worms were their own architects, untrammelled by convention or heredity, hence they built very different, sometimes almost fantastic, structures.
We must remember, too, the great age of this group. They are present in highly modified forms in the very oldest palæozoic strata, and probably therefore came into existence as the first traces of continental areas were beginning to rise above the primeval ocean. They are literally "older than the hills." They were exposed to a host of rapidly changing conditions, very different in different areas. This prepares us for the fact that the worms represent a stage in animal life corresponding fairly well to the Tower of Babel in biblical history. The animal kingdom seems almost to explode into a host of fragments. Our genealogical tree fairly bristles with branches, but the branches do not seem to form any regular whorls or spirals. Few of them have developed into more than feeble growths. They now contain generally but few species. Many of them are largely or entirely parasitic, and in connection with this mode of life have undergone modifications and degeneration which make it exceedingly difficult to decipher their descent or relationships.
Four of these branches have reached great prominence in numbers and importance. One or two others were formerly equally numerous and have since become almost extinct; so the brachiopoda, which have been almost entirely replaced by mollusks. The same may very possibly be true of others. For of the amount of extinction of larger groups we have generally but an exceedingly faint conception. Indeed in this respect the worms have been well compared to the relics which fill the shelves of one of our grandmother's china-closets.
The four great branches are the echinoderms, mollusks, articulates, and vertebrates. The echinoderms, including starfishes, sea-urchins, and others straggled early from the great army. We know as yet almost nothing of their history; when deciphered it will be as strange as any romance. The vertebrates are of course the most important line, as including the ancestors of man. But we must take a little glance at mollusks, including our clams, snails, and cuttle-fishes; and at the articulates, including annelids and culminating in insects. The molluscan and articulate lines, though divergent, are of great importance to us as throwing a certain amount of light on vertebrate development; and still more as showing how a certain line of development may seem, and at first really be, advantageous, and still lead to degeneration, or at best to but partial success.
When we compare the forms which represent fairly well the direction of development of these three lines, a snail or a clam with an insect and a fish, we find clearly, I think, that the fundamental anatomical difference lies in the skeleton; and that this resulted from, and almost irrevocably fixed, certain habits of life.
We may picture to ourselves the primitive ancestor of mollusks as a worm having the short and broad form of the turbellaria, but much thicker or deeper vertically. A fuller description can be found in the "Encyclopædia Britannica," Art., Mollusca. It was hemi-ovoid in form. It had apparently the perivisceral cavity and nephridia of the schematic worm, and a circulatory system. In this latter respect it stood higher than any form which we have yet studied. Its nervous system also was rather more advanced. It had apparently already taken to a creeping mode of life and the muscles of its ventral surface were strongly developed, while its exposed and far less muscular dorsal surface was protected by a cap-like shell covering the most important internal organs. But the integument of the whole dorsal surface was, as is not uncommon in invertebrates, hardening by the deposition of carbonate of lime in the integument. And this in time increased to such an extent as to replace the primitive, probably horny, shell.
Into the anatomy of this animal or of its descendants we have no time to enter, for here we must be very brief. We have already noticed that the most important viscera were lodged safely under the shell. And as these increased in size or were crowded upward by the muscles of the creeping disk, their portion of the body grew upward in the form of a "visceral hump." Apparently the animal could not increase much in length and retain the advantage of the protection of the shell; and the shell was the dominating structure. It had entered upon a defensive campaign. Motion, slow at the outset, became more difficult, and the protection of the shell therefore all the more necessary. The shell increased in size and weight and motion became almost impossible. The snail represents the average result of the experiment. It can crawl, but that is about all; it is neither swift nor energetic. Even the earthworm can outcrawl it. It has feelers and eyes, and is thus better provided with sense-organs than almost any worm. It has a supra-œsophageal ganglion of fair size.
The clams and oysters show even more clearly what we might call the logical results of molluscan structure. They increased the shell until it formed two heavy "valves" hanging down on each side of the body and completely enclosing it. They became almost sessile, living generally buried in the mud and gaining their food, consisting mostly of minute particles of organic matter, by means of currents created by cilia covering the large curtain-like gills. Their muscular system disappeared except in the ploughshare-shaped "foot" used mostly for burrowing, and in the muscles for closing the shell. That portion of the body which corresponds to the head of the snail practically aborted with nearly all the sense-organs. The nervous system degenerated and became reduced to a rudiment. They had given up locomotion, had withdrawn, so to speak, from the world; all the sense they needed was just enough to distinguish the particles of food as they swept past the mouth in the current of water. They have an abundance of food, and "wax fat." The clam is so completely protected by his shell and the mud that he has little to fear from enemies. They have increased and multiplied and filled the mud. "Requiescat in pace."
But zoölogy has its tragedies as well as human history. Let us turn to the development of a third molluscan line terminating in the cuttle-fishes. The ancestors of these cephalopods, although still possessed of a shell and a high visceral hump, regained the swimming life. First, apparently, by means of fins, and then by a simple but very effective use of a current of water, they acquired an often rapid locomotion. The highest forms gave up the purely defensive campaign, developed a powerful beak, led a life like that of the old Norse pirates, and were for a time the rulers and terrors of the sea. With their more rapid locomotion the supra-œsophageal ganglion reached a higher degree of development, and it was served by sense-organs of great efficiency. They reduced the external shell, and succeeded, in the highest forms, of almost ridding themselves of this burden and encumbrance. Traces of it remain in the squids, but transformed into an internal quill-like, supporting, not defensive, skeleton. They have retraced the downward steps of their ancestors as far as they could. And the high development of their supra-œsophageal ganglion and sense-organs, and their powerful jaws and arms, or tentacles, show to what good purpose they have struggled. But the struggle was in vain, as far as the supremacy of the animal kingdom was concerned. Their ancestors had taken a course which rendered it impossible for their descendants to reach the goal. Their progress became ever slower. They were entirely and hopelessly beaten by the vertebrates. They struggled hard, but too late.
The history of mollusks is full of interest. They show clearly how intimately nervous development is connected with the use of the locomotive organs. The snail crept, and slightly increased its nervous system and sense-organs. The clam almost lost them in connection with its stationary life. The cephalopods were exceedingly active, developed, therefore, keen sense-organs and a very large and complicated supra-œsophagal ganglion, which we might almost call a brain.
The articulate series consists of two groups of animals. The higher group includes the crabs, spiders, thousand-legs, and finally the insects, and forms the kingdom of arthropoda. The lower members are still usually reckoned as worms, and are included under the annelids. Of these our common earthworm is a good example, and near them belong the leeches. But the marine annelids, of which nereis, or a clam-worm, is a good example, are more typical. They are often quite large, a foot or even more in length. They are composed of many, often several hundred, rings or segments. Between these the body-wall is thin, so that the segments move easily upon each other, and thus the animal can creep or writhe.
7. EUNICE LIMOSA (ANNELID). LANG, FROM EHLERS.
Front and hind end seen from dorsal surface. fa, fp, fc, feelers; a, eye; k, gill; p, parapodia; ac, anal cirri.
These segments are very much alike except the first two and the last. If we examine one from the middle of the body we shall find its structure very much like that of our schematic worm. Outside we find a very thin, horny cuticle, secreted by the layer of cells just beneath it, the hypodermis. Beneath the skin we find a thin layer of transverse muscles, and then four heavy bands of longitudinal muscles. These latter have been grouped in the four quadrants, a much more effective arrangement than the cylindrical layer of the schematic worm. Furthermore, the animal has on each segment a pair of fin-like projections, stiffened with bristles, the parapodia. These are moved by special muscles and form effective organs of creeping.
Within the muscles is the perivisceral cavity, and in its central axis the intestine, segmented like the body-wall. The reproductive organs are formed from patches of the lining of the perivisceral cavity, and the reproductive elements, when fully developed, fall into the perivisceral fluid and are carried out by nephridia, just such as we found in the schematic worm. Beside the perivisceral cavity and its fluid there is a special circulatory system. This consists mainly of one long tube above the intestine and a second below, with often several smaller parallel tubes. Transverse vessels run from these to all parts of the body. The dorsal tube pulsates and thus acts as a heart. The surface of the body no longer suffices to gather oxygen, hence we find special feathery gills on the parapodia. But these gills are merely expanded portions of the body wall, arranged so as to offer the greatest possible amount of surface where the capillaries of the blood system can be almost immediately in contact with the surrounding water.
The nervous system consists of a large supra-œsophageal ganglion in the first segment; then of a chain of ganglia, one to each segment, on the ventral side of the body. With one ganglion in each segment there is far more controlling, perceptive, ganglionic material than in lower worms. Furthermore the supra-œsophageal ganglion is relieved of a large part of the direct control of the muscles of each segment, and is becoming more a centre of control and perception for the body as a whole. It is more like our brain, commander-in-chief, the other ganglia constituting its staff. The sense-organs have improved greatly. There are tentacles and otolith vesicles as very delicate organs of feeling, or possibly of hearing also.
But the annelids were probably the first animals to develop an eye capable of forming an image of external objects. The importance of this organ in the pursuit of food or the escape from enemies can scarcely be over-estimated. The lining of the mouth and pharynx can be protruded as a proboscis, and drawn back by powerful muscles, and is armed with two or more horny claws. Eyes and claws gave them a great advantage over their not quite blind but really visionless and comparatively defenceless neighbors, and they must have wrought terrible extinction of lower and older forms. But while we cannot over-estimate the importance of these eyes, we can easily exaggerate their perfectness. They were of short range, fitted for seeing objects only a few inches distant, and the image was very imperfect in detail. But the plan or fundamental scheme of these eyes is correct and capable of indefinitely greater development than the organs of touch or smell, perhaps greater even than the otolith vesicle.
8. CROSS-SECTION OF BODY SEGMENT OF ANNELID. LANG.
dp and vp, dorsal and ventral halves of parapodia; b and ac, bristles; k, gill; dc and vc, feelers; rm, lateral muscles; lm, longitudinal muscles; vd, dorsal blood-vessel; vo, ventral blood-vessel; bm, ventral ganglion; ov, ovary; tr, opening of nephridium in the perivisceral cavity; np, tubular portion of nephridium. The circles containing dots represent eggs floating in the perivisceral fluid.
And the reflex influence of the eye on the brain was the greatest advantage of all. Hitherto with feeble muscles and sense-organs it has hardly paid the animal to devote more material to building a larger brain. It was better to build more muscle. But now with stronger muscles at its command, and better sense-organs to report to it, every grain of added brain material is beginning to be worth ten devoted to muscle. The muscular system will still continue to develop, but the brain has begun an almost endless march of progress. The eye becomes of continually increasing advantage and importance because it has a capable brain to use it; and brain is a more and more profitable investment, because it is served by an ever-improving eye.
The annelid had hit upon a most advantageous line of development, which led ultimately to the insect. The study of the insect will show us clearly the advantages and defects of the annelid plan. First of all, the insect, like the mollusk, has an external skeleton. But the skeleton of the mollusk was purely protective, a hindrance to locomotion. That of the insect is still somewhat protective, but is mainly, almost purely, locomotive. It is never allowed to become so heavy as to interfere with locomotion. In the second place, the insect has three body regions, having each its own special functions or work. And one of these is a head. The annelid had two anterior segments differing from those of the rest of the body; these may, perhaps, be considered as the foreshadowings of a structure not yet realized; they can only by courtesy be called a head. Thirdly, the insect has legs. The annelid had fin-like parapodia, approaching the legs of insects about as closely as the fins of a fish approach the legs of a mammal. The reproductive and digestive systems, while somewhat improved, are not very markedly higher than those of annelids. The excretory system has more work to perform and reaches a rather higher development.
9. MYRMELEO FORMICARIUS. ANT-LION. HERTWIG, FROM SCHMARDA.
1, adult; 2, larva; 3, cocoon.
But in these organs there is no great or striking change; the time for marked and rapid development of the digestive and reproductive systems has gone by. Material can be more profitably invested in brain or muscle. Air is carried to all parts of the body by a special system of air-sacks and tubes. This is a very advantageous structure for small animals with an external skeleton. In very large animals, or where the skeleton is internal, it would hardly be practicable; the risk of compression of the tubes at some point, and of thus cutting off the air-supply of some portion of the body, would be altogether too great.
The circulatory system is very poor. It consists practically only of a heart, which drives the blood in an irregular circulation between the other organs of the body much as with a syringe you might keep up a system of currents in a bowl of water. But the rapidity of the flow of the blood in our bodies is mainly to furnish a supply of oxygen to the organs. A tea-spoonful of blood can carry a fair amount of dissolved solid nutriment like sugar, it can carry at each round but a very little gas like oxygen. Hence the blood must make its rounds rapidly, carrying but a little oxygen at each circuit. But in the insect the blood conveys only the dissolved solid nutriment, the food; hence a comparatively irregular circulation answers all purposes.
The skeleton is a thickening of the horny cuticle of the annelid on the surface of each segment. The horny cylinder surrounding each segment is composed of several pieces, and on the abdomen these are united by flexible, infolded membranes. This allows the increase in the size of the segment corresponding to the varying size of the digestive and reproductive systems. In this part of the body the skeletal ring of each segment is joined to that of the segments before and behind it in the same manner. But in other parts of the body we shall find the skeletal pieces of each segment and the rings of successive segments fused in one plate of mail. The legs are the parapodia of annelids carried to a vastly higher development. They are slender and jointed, and yet often very powerful. A large portion of the muscular system of the body is attached to these appendages.
But the insect has also jaws. The annelid had teeth or claws attached to the proboscis. But true jaws are something quite different. They always develop by modifying some other organ. In the insect they are modified legs. This is shown first by their embryonic development. But the king- or horseshoe-crab has still no true jaws, but uses the upper joints of its legs for chewing. There are primitively three pairs of jaws of various forms for the different kinds of food of different species or higher groups. But some of them may disappear and the others be greatly modified into awls for piercing, or a tube for sucking honey. Into the wonderful transformations of these modified legs we cannot enter.
The muscles are no longer arranged to form a sack as in annelids. Transverse muscles, running parallel to the unyielding plates of chitin or horn could accomplish nothing. They have largely disappeared. The work of locomotion has been transferred from the trunk to the legs.
The abdomen of the insect is as clearly composed of distinct segments as the body of the annelid. Of these there are perhaps typically eleven. The thorax is composed of three segments, distinct in the lowest forms, fused in the highest. This fusion of segments in the thorax of the highest forms furnishes a very firm framework for the attachment of wings and muscles. These wings are a new development, and how they arose is still a question. But they give the insect the capability of exceedingly rapid locomotion.
The three pairs of jaws, modified legs, in the rear half of the head show that this portion is composed of three segments. For only one pair of legs is ever developed on a single segment. Embryology has shown that the portion of the head in front of the mouth is also composed of three segments. Possibly between the præ- and post-oral portions still another segment should be included, making a total of seven in the head. The head has thus been formed by drawing forward segments from the trunk, and fusing them successively with the first or primitive head segment. This is difficult to conceive of in the fully developed insect, where the boundary between head and thorax is very sharp. But the ancestors of insects looked more like thousand-legs or centipedes, and here head and thorax are much less distinct. But in the annelid the mouth is on the second segment; here it is on the fourth. It has evidently travelled backward. That the mouth of an animal can migrate seems at first impossible, but if we had time to examine the embryology of annelids and insects, it would no longer appear inconceivable or improbable. And its backward migration brought it among the legs which were grasping and chewing the food. And in vertebrates the mouth has changed its position, though not in exactly the same way. Our present mouth is probably not at all the mouth of the primitive ancestor of vertebrates. Thus in the insect three segments have fused around the mouth, and three, possibly four, in front of it. This makes a head worthy of the name. The ganglia of the three post-oral segments, which bear the jaws, have fused in one compound ganglion innervating the mouth and jaws. Those of the three præ-oral segments have fused to form a brain. Eyes are well developed, giving images sometimes accurate in detail, sometimes very rude. Ears are not uncommon. The sense of smell is often keen.
Perhaps the greatest advance of the insect is its adaptation to land life. This gives it a larger supply of oxygen than any aquatic animal could ever obtain. This itself stimulates every function, and all the work of the body goes on more energetically. Then the heat produced is conducted off far less rapidly than in aquatic forms. Water is a good conductor of heat, and nearly all aquatic animals are cold-blooded. The few which are warm-blooded are protected by a thick layer of non-conducting fat. In all land animals, even when cold-blooded, the work of the different systems is aided by the longer retention of the heat in the body.
Let us recapitulate. The schematic worm had a body composed of two concentric tubes. The outer was composed of the muscles of the body covered by the protective integument. The inner tube was the alimentary canal with its special muscles. Between these two was the perivisceral cavity, filled with nutritive fluid, lymph, and furnishing a safe lodging-place for the more delicate viscera. It represented fairly the trunk of higher animals.
The annelid added segmentation, and thus greater freedom of motion by the parapodia. But the segments were still practically alike. In the insect division of labor took place, that is, each group of segments was allotted its own special work; and these groups of segments were modified in structure to best suit the performance of this part of the work of the body. The abdomen was least modified and its eleven segments were devoted to digestion, reproduction, and excretion—the old vegetative functions. Three segments were united in the thorax; all their energy was turned to locomotion, and the insect became thus an exceedingly active, swift animal. The third body-region, the head, includes six segments, of which three surrounded the mouth and furnished the jaws, while two more were crowded or drawn forward in order that their ganglia might be added to the old supraœsophageal ganglion and form a brain. It is interesting to note that a form, peripatus, still exists which stands almost midway between annelids and insects and has only four segments in the head. The formation of the head was thus a gradual process, one segment being added after another.
In the turbellaria the dominant functions were digestion and reproduction, and their organs composed almost the whole body. Here only eleven segments at most are devoted to these functions, and nine in head and thorax to locomotion and brain. Head and thorax have increased steadily in importance, while the abdomen has decreased as steadily in number of segments. And the brain is increasing thus rapidly because there are now muscles and sense-organs of sufficient power to make such a brain of value. And this brain perceives not only objects and qualities, but invisible relations between these, and this is an advance amounting to a revolution. It remembers, and uses its recollections. It is capable of learning a little by experience and observation. The A, B, C of thinking was probably learned long before the insect's time, and the bee shows a fair amount of intelligence.
The line of development which the insect followed was comparatively easy and its course probably rapid. Certain crustacea, aquatic arthropoda, are among the oldest fossils, and it is possible that insects lived on the land before the first fish swam in the sea. They had fine structure and powers; and yet during the later geologic periods they have scarcely advanced a step, and are now apparently at a standstill. They ran splendidly for a time, and then fell out of the race. What hindered and stopped them?
One vital defect in their whole plan of organization is evident. The external skeleton is admirably suited to animals of small size, but only to these. In larger animals living on land it would have to be made so heavy as to be unwieldy and no longer economical. Their mode of breathing also is fitted only for animals of small size having an external skeleton. Whatever may be our explanation the fact remains that insects are always small. This is in itself a disadvantage. Very small animals cannot keep up a constant high temperature unless the surrounding air is warm, for their radiating surface is too large in comparison with their heat-producing mass. At the first approach of even cool weather they become chilled and sluggish, and must hibernate or die. They are conformed to but a limited range of environment in temperature.
But small size is, as a rule, accompanied by an even greater disadvantage. It seems to be almost always correlated with short life. Why this is so, or how, we do not know. There are exceptions; a crow lives as long as a man; or would, if allowed to. But, as a rule, the length of an animal's days is roughly proportional to the size of its body. And the insect is, as a rule, very short-lived. It lives for a few days or weeks, or even months, but rarely outlasts the year. It has time to learn but little by experience. The same experience must be passed, the same emergency arise and be met, over and over again during the lifetime of the same individual if the animal is to learn thereby. And intelligence is based upon experience. Hence insects can and do possess but a low grade of intelligence. But instinct is in many cases habit fixed by heredity and improved by selection. The rapid recurrence of successive generations was exceedingly favorable to the development of instincts, but very unfavorable to intelligence. Insects are instinctive, the highest vertebrates intelligent. The future can never belong to a tiny animal governed by instincts. Mollusks and insects have both failed to reach the goal; another plan of structure than theirs must be sought if the animal kingdom is to have a future.
The future belonged to the vertebrate. To begin with less characteristic organs the digestive system is much like that of the annelid or schematic worm, but with greatly increased glandular and absorptive surfaces. The present mouth of nearly all vertebrates is probably not primitive. It is almost certainly one of the gill-slits of some old ancestor of fish, such as now are used to discharge the water which is used for respiration. The jaws are modified branchial arches or the cartilaginous or bony rods which in our present fish support the fringe of gills. These have formed a pair of exceedingly effective and powerful jaws. The reproductive system holds still to the old type and shows little if any improvement. The excretory organs, kidneys, are composed primitively of nephridial tubes like those of the schematic worm or annelid, but immensely increased in number, modified, and improved in certain very important particulars. The muscles in simplest forms are composed of heavy longitudinal bands, especially developed toward the dorsal surface of the body to the right and left of the axial skeleton. Locomotion was produced by lashing the tail right and left, as still in fish. There is improvement in all these organs, except perhaps the reproductive, but nothing very new or striking. The great improvement from this time on was not to be sought in the vegetative organs, or even directly to any great extent in muscles.
The new and characteristic organ was not the vertebral column, or series of vertebræ, or backbone, from which the kingdom has derived its name. This was a later production. The primitive skeleton was the notochord, still appearing in the embryos of all vertebrates and persisting throughout life in fish. This is an elastic rod of cartilage, lying just beneath the spinal marrow or nerve-cord, which runs backward from the brain. The nerve-centres are therefore here all dorsal, and the notochord or skeleton lies between these and the digestive or alimentary canal. The skeleton of the clam or snail is purely protective and a hindrance to locomotion. That of the insect is almost purely locomotive, but external, that of the vertebrate purely locomotive and internal. It does not lie outside even of the nervous system, although this system especially required, and was worthy of, protection. It does not protect even the brain; the skull of vertebrates is an after-thought. It is almost the deepest seated of all organs. But lying in the central axis of the body it furnishes the very best possible attachment for muscles. Around this primitive notochord was a layer of connectile tissue which later gave rise to the vertebræ forming our backbone.
10. CROSS-SECTION OF AXIAL SKELETON OF PETROMYZON. HERTWIG, FROM HIEDERSHEIM.
SS, skeletogenous layer; Ob, Ub, dorsal and ventral processes of SS; C, notochord; Cs, sheath of notochord; Ee, elastic external layer of sheath; F, fatty tissue; M, spinal marrow; P, sheath of M.
The nervous system on the dorsal surface of the notochord consists of the brain in the head and the spinal marrow running down the back. The brain of all except the very lowest vertebrates consists of four portions: 1. The cerebrum, or cerebral lobes, or simply "forebrain," the seat of consciousness, thought, and will, and from which no nerves proceed. Whether the primitive vertebrate had any cerebrum is still uncertain. 2. The mid-brain, which sends nerves to the eyes, and in this respect reminds us of the brain of insects. Its anterior portion appears from embryology to be very primitive. 3. The small brain, or cerebellum, which in all higher forms is the centre for co-ordination of the motions of the body. 4. The medulla, which controls especially the internal organs. The spinal marrow, or that portion of the nervous system which lies outside of the head, is at the same time a great nerve-trunk and a centre for reflex action of the muscles of the body. But the development of these distinct portions and the division of labor between them must have been a long and gradual process.
We have every reason to believe that here, as in insects, the head has been formed by annexation of segments from the rump and the fusion of their nervous matter with that of the brain. But here, instead of only three segments, from nine to fourteen have been fused in the head to furnish the material for the brain. Notochord and backbone may be the most striking and apparent characteristic of vertebrates, but their predominant characteristic is brain. On this system they lavished material, giving it from three to four times as much as any lower or earlier group had done. They very early set apart the cerebral lobes to be the commander-in-chief and centre of control for all other nerve-centres. To this all report, and from it all directly or indirectly receive orders. It can say to every other organ in the body, "Starve that I may live." It is the seat of thought and will. The other portions of the brain report to it what they have gathered of vision or sound; it explains the vision or song or parable. It is relieved as far as possible from all lower and routine work that it may think and remember and govern. The vertebrate built for mind, not neglecting the body.
Every trait of vertebrates is a promise of a great future. Its internal skeleton gives it the possibility of large size. This gave it in time the victory in the struggle with its competitors, as to whether it should eat or be eaten. It is vigorous and powerful, for all its organs are at the best. It gives the possibility of later, on land, becoming warm-blooded, i.e., of maintaining a constant high temperature. It is thus resistant to climate and hardship. In time its descendants will face the arctic winter as well as the heat of the tropics.
But it has started on the road which leads to mind. The greater size is correlated with longer life. The lessons of experience come to it over and over again, and it can and must learn them. It is the intelligent, remembering, thinking type. The insect had begun to peer into the world of invisible and intangible relations, the vertebrate will some day see them. This much is prophecied in his very structure. He must be heir to an indefinite future.
You have probably noticed that the vertebrate differs greatly from all his predecessors. The gulf between him and them is indeed wide and deep. His origin and ancestry are yet far from certain. But an attempt to decipher his past history, though it may lead to no sure conclusions, will yet be of use to us. Practically all aquatic vertebrates lead a swimming life, neither sessile nor creeping. The embryonic development of our appendages leads to the same conclusion. We must never forget that the embryonic development of the individual recapitulates briefly the history of the development of the race. Now the legs and arms, or fore- and hind-legs, of higher vertebrates and the corresponding paired fins of fish develop in the embryo as portions of a long ridge extending from front to rear of the side of the body.
This justifies the inference that the primitive vertebrate ancestor had a pair of long fins running along the sides of the body, but bending slightly downward toward the rear so as to meet one another and continue as a single caudal fin behind the anal opening. Such fins, like the feathers of an arrow, could be useful only to keep the animal "on an even keel" as it was forced through the water by the lateral sweeps of the tail. They would have been useless for creeping.
But there is another piece of evidence that he was a free swimming form. All vertebrates breathe by gills or lungs, and these are modified portions of the digestive system, of the walls of the œsophagus, from which even the lung is an embryonic outgrowth. Now practically all invertebrates breathe through modified portions of the integument or outer surface of the body, and their gills are merely expansions of this. In the annelid they are projections of the parapodia, in the mollusk expansions of the skin, where the foot or creeping sole joins the body. Why did the vertebrate take a new and strange, and, at first sight, disadvantageous mode of breathing? There must have been some good reason for this. The most natural explanation would seem to be that he had no projections on his outer surface which could develop into gills, and farther, that he could not afford to have any. Now projections on the lower portion of the sides of the body would be an advantage in creeping, but a hindrance in any such mode of swimming as we have described, or indeed in any mode of writhing through the water.
Furthermore, if he lived, not a creeping life on the bottom, but swimming in the water above, he would have to live almost entirely on microscopic animals and embryos; and these would be most easily captured by a current of water brought in at the mouth. The whole branchial apparatus in its simplest forms would seem to be an apparatus for sifting out the microscopic particles of food and only later a purely respiratory apparatus. Moreover, we have seen that the parapodia of annelids naturally point to the development of an external skeleton, for their muscles are already a part of the external body-wall and attached to the already existing horny cuticle. The logical goal of their development was the insect.
Now I do not wish to conceal from you that many good zoölogists 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 "balls" 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 mass of the cœlenterates 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 hæmal arches. These unite with the masses of cartilage surrounding the notochord to form cartilaginous vertebræ, which may be stiffened by an infiltration of carbonate of lime. The vertebral column of sharks has reached this stage. Then the cartilaginous vertebræ 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 buttresses, 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 passage 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 vertebræ with few exceptions are hollow in front and behind, biconcave; and between each two vertebræ there is a large cavity still occupied by the notochord. Thus these vertebræ join one another by their edges, like two shallow wine-glasses placed rim to rim. Only gradually is the notochord crowded out so that the vertebræ 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 vertebræ 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 partition 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 class 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 mass 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 surpassed 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 competition 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 assigned 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 associated systems of blood-vessels, nutriment and oxygen diffuse from the blood of the mother into that of the embryo, and thus rapid growth is assured. 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 mass 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 mass 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 competitors. 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 principal 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 palæontologist 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 class 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 classes from the twelfth century, averaged about 1,426 cubic centimetres, while ninety of those of the poorer classes 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 assigned 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 analogy exhibited in the cerebral forms in all these creatures. There is a cerebral form peculiar to man and the apes; and so in the cerebral convolutions, wherever they appear, there is a general unity of arrangement, a plan, the type of which is common to all these creatures." Professor Huxley says: "It is most remarkable that, as soon as all the principal sulci appear, the pattern according to which they are arranged is identical with the corresponding sulci in man. The surface of the brain of the monkey exhibits a sort of skeleton map of man's, and in the man-like apes the details become more and more filled in, until it is only in minor characters that the chimpanzee's or orang's brain can be structurally distinguished from man's."