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MR. MARTINEAU ON EVOLUTION.
[First published in The Contemporary Review, for June, 1872.]
The article by Mr. Martineau, in the April number of the Contemporary Review, on "The Place of Mind in Nature, and Intuition of Man," recalled to me a partially-formed intention to deal with the chief criticisms which have from time to time been made on the general doctrine set forth in First Principles; since, though not avowedly directed against propositions asserted or implied in that work, Mr. Martineau's reasoning tells against them by implication. The fulfilment of this intention I should, however, have continued to postpone, had I not learned that the arguments of Mr. Martineau are supposed by many to be conclusive, and that, in the absence of replies, it will be assumed that no replies can be made. It seems desirable, therefore, to notice these arguments at once—especially as the essential ones may, I think, be effectually dealt with in a comparatively small space.
* * * * *
The first definite objection which Mr. Martineau raises is, that the hypothesis of General Evolution is powerless to account even for the simpler orders of facts in the absence of numerous different substances. He argues that were matter all of one kind, no such phenomena as chemical changes would be possible; and that, "in order to start the world on its chemical career, you must enlarge its capital and present it with an outfit of heterogeneous constituents. Try, therefore, the effect of such a gift; fling into the pre-existing cauldron the whole list of recognized elementary substances, and give leave to their affinities to work." The intended implication obviously is, that there must exist the separately-created elements before evolution can begin.
Here, however, Mr. Martineau makes an assumption which few, if any, chemists will commit themselves to, and which many will distinctly deny. There are no "recognized elementary substances," if the expression means substances known to be elementary. What chemists, for convenience, call elementary substances, are merely substances which they have thus far failed to decompose; but, bearing in mind past experiences, they do not dare to say that they are absolutely undecomposable. Water was taken to be an element for more than two thousand years, and then was proved to be a compound; and, until Davy brought a galvanic current to bear upon them, the alkalies and the earths were supposed to be elements. So little true is it that "recognized elementary substances" are supposed to be absolutely elementary, that there has been much speculation among chemists respecting the process of compounding and recompounding by which they have been formed out of some ultimate substance—some chemists having supposed the atom of hydrogen to be the unit of composition, but others having contended that the atomic weights of the so-called elements are not thus interpretable. If I remember rightly, Sir John Herschel was one, among others, who, some five-and-twenty years ago, threw out suggestions respecting a system of compounding that might explain these relations of the atomic weights.
What was at that time a suspicion has now become practically a certainty. Spectrum-analysis yields results wholly irreconcilable with the assumption that the conventionally-named simple substances are really simple. Each yields a spectrum having lines varying in number from two to eighty or more, every one of which implies the intercepting of ethereal undulations of a certain order by something oscillating in unison or in harmony with them. Were iron absolutely elementary, it is not conceivable that its atom could intercept ethereal undulations of eighty different orders. Though it does not follow that its molecule contains as many separate atoms as there are lines in its spectrum, it must clearly be a complex molecule. The evidence thus gained points to the conclusion that, out of some primordial units, the so-called elements arise by compounding and recompounding; just as by the compounding and recompounding of so-called elements there arise oxides, and acids, and salts.
And this hypothesis is entirely in harmony with the phenomena of allotropy. Various substances, conventionally distinguished as simple, have several forms under which they present quite different properties. The semi-transparent, colourless, extremely active substance called phosphorus may be so changed as to become opaque, dark red, and inert. Like changes are known to occur in some gaseous, non-metallic elements, as oxygen; and also in metallic elements, as antimony. These total changes of properties, brought about without any changes to be called chemical, are interpretable only as due to molecular rearrangements; and, by showing that difference of property is producible by difference of arrangement, they support the inference otherwise to be drawn, that the properties of different elements result from differences of arrangement arising by the compounding and recompounding of ultimate homogeneous units.
Thus Mr. Martineau's objection, which at best would imply a turning of our ignorance of the nature of elements into positive knowledge that they are simple, is, in fact, to be met by two sets of evidences, which imply that they are compound.
* * * * *
Mr. Martineau next alleges that a fatal difficulty is put in the way of the General Doctrine of Evolution by the existence of a chasm between the living and the not-living. He says:—"But with all your enlargement of data, turn them as you will, at the end of every passage which they explore, the door of life is closed against them still." Here again our ignorance is employed to play the part of knowledge. The fact that we do not know distinctly how an alleged transition has taken place, is transformed into the assumption that no transition has taken place. We have, in a more general shape, the argument which until lately was thought conclusive—the argument that because the genesis of each species of creature had not been explained, therefore each species must have been separately created.
Merely noting this, however, I go on to remark that scientific discovery is day by day narrowing the chasm, or, to vary Mr. Martineau's metaphor, "opening the door." Not many years since, it was held as certain that the chemical compounds distinguished as organic could not be formed artificially. Now, more than a thousand organic compounds have been formed artificially. Chemists have discovered the art of building them up from the simpler to the more complex, and do not doubt that they will eventually produce the most complex. Moreover, the phenomena attending isomeric change give a clue to those movements which are the only indications we have of life in its lowest forms. In various colloidal substances, including the albuminoid, isomeric change is accompanied by contraction or expansion, and consequent motion; and, in such primordial types as the Protogenes of Haeckel, which do not differ in appearance from minute portions of albumen, the observed motions are comprehensible as accompanying isomeric changes caused by variations in surrounding physical actions. The probability of this interpretation will be seen on remembering the evidence we have that, in the higher organisms, many functions are essentially effected by isomeric changes from one to another of the multitudinous forms which protein assumes.
Thus the reply to this objection is, first, that there is going on from both sides a narrowing of the chasm supposed to be impassable; and, secondly, that, even were the chasm not in course of being filled up, we should no more be justified in therefore assuming a supernatural commencement of life, than Kepler was justified in assuming that there were guiding-spirits to keep the planets in their orbits, because he could not see how else they were to be kept in their orbits.
* * * * *
The third definite objection made by Mr. Martineau is of kindred nature. The Hypothesis of Evolution is, he thinks, met by the insurmountable difficulty that plant life and animal life are absolutely distinct. "You cannot," he says, "take a single step toward the deduction of sensation and thought: neither at the upper limit do the highest plants (the exogens) transcend themselves and overbalance into animal existence; nor at the lower, grope as you may among the sea-weeds and sponges, can you persuade the sporules of the one to develop into the other."
This is an extremely unfortunate objection to raise. For, though there are no transitions from vegetal to animal life at the places Mr. Martineau names, where, indeed, no biologist would look for them; yet the connexion between the two great kingdoms of living things is so complete that separation is now regarded as impossible. For a long time naturalists endeavored to frame definitions such as would, the one include all plants and exclude all animals, and the other include all animals and exclude all plants. But they have been so repeatedly foiled in the attempt that they have given it up. There is no chemical distinction which holds; there is no structural distinction which holds; there is no functional distinction which holds; there is no distinction as to mode of existence which holds. Large groups of the simpler animals contain chlorophyll, and decompose carbonic acid under the influence of light, as plants do. Large groups of the simpler plants, as you may observe in the diatoms from any stagnant pool, are no less actively locomotive than the minute creatures classed as animals seen along with them. Nay, among these lowest types of living things, it is common for the life to be now predominantly animal and presently to become predominantly vegetal. The very name zoospores, given to germs of algae, which for a while swim about actively by means of cilia, and presently settling down grow into plant-forms, is given because of this conspicuous community of nature. So complete is this community of nature that for some time past many naturalists have wished to establish for these lowest types a sub-kingdom, intermediate between the animal and the vegetal: the reason against this course being, however, that the difficulty crops up afresh at any assumed places where this intermediate sub-kingdom may be supposed to join the other two.
Thus the assumption on which Mr. Martineau proceeds is diametrically opposed to the conviction of naturalists in general.
* * * * *
Though I do not perceive that it is specifically stated, there appears to be tacitly implied a fourth difficulty of allied kind—the difficulty that there is no possibility of transition from life of the simplest kind to mind. Mr. Martineau says, indeed, that there can be "with only vital resources, as in the vegetable world, no beginning of mind:" apparently leaving it to be inferred that in the animal world the resources are such as to make the "beginning of mind" comprehensible. If, however, instead of leaving it a latent inference, he had distinctly asserted a chasm between mind and bodily life, for which there is certainly quite as much reason as for asserting a chasm between animal life and vegetal life, the difficulties in his way would have been no less insuperable.
For those lowest forms of irritability in the animal kingdom which, I suppose, Mr. Martineau refers to as the "beginning of mind," are not distinguishable from the irritability which plants display: they in no greater degree imply consciousness. If the sudden folding of a sensitive-plant's leaf when touched, or the spreading out of the stamens in a wild-cistus when gently brushed, is to be considered a vital action of a purely physical kind; then so too must be considered the equally slow contraction of a polype's tentacles. And yet, from this simple motion of an animal of low type, we may pass by insensible stages through ever-complicating forms of actions, with their accompanying signs of feeling and intelligence, until we reach the highest.
Even apart from the evidence derived from the ascending grades of animals up from zoophytes, as they are significantly named, it needs only to observe the evolution of a single animal to see that there does not exist any break or chasm between the life which shows no mind and the life which shows mind. The yelk of an egg which the cook has just broken, not only yields no sign of mind, but yields no sign of life. It does not respond to a stimulus as much even as many plants do. Had the egg, instead of being broken by the cook, been left under the hen for a certain time, the yelk would have passed by infinitesimal gradations through a series of forms ending in the chick; and by similarly infinitesimal gradations would have arisen those functions which end in the chick breaking its shell; and which, when it gets out, show themselves in running about, distinguishing and picking up food, and squeaking if hurt. When did the feeling begin? and how did there come into existence that power of perception which the chick's actions show? Should it be objected that the chick's actions are mainly automatic, I will not dwell on the fact that, though they are largely so, the chick manifestly has feeling and therefore consciousness; but I will accept the objection, and propose that instead we take the human being. The course of development before birth is just of the same general kind; and similarly, at a certain stage, begins to be accompanied by reflex movements. At birth there is displayed an amount of mind certainly not greater than that of the chick: there is no power of running from danger—no power of distinguishing and picking up food. If we say the chick is unintelligent, we must certainly say the infant is unintelligent. And yet from the unintelligence of the infant to the intelligence of the adult, there is an advance by steps so small that on no day is the amount of mind shown, appreciably different from that shown on preceding and succeeding days.
Thus the tacit assumption that there exists a break, is not simply gratuitous, but is negatived by the most obvious facts.
* * * * *
Certain of the words and phrases used in explaining that particular part of the Doctrine of Evolution which deals with the origin of species, are commented upon by Mr. Martineau as having implications justifying his view. Let us consider his comments.
He says that competition is not an "original power, which can of itself do anything;" further, that "it cannot act except in the presence of some possibility of a better or worse;" and that this "possibility of a better or worse" implies a "world pre-arranged for progress," "a directing Will intent upon the good." Had Mr. Martineau looked more closely into the matter, he would have found that, though the words and phrases he quotes are used for convenience, the conceptions they imply are not at all essential to the doctrine. Under its rigorously-scientific form, the doctrine is expressible in purely-physical terms, which neither imply competition nor imply better and worse.[37]
Beyond this indirect mistake there is a direct mistake. Mr. Martineau speaks of the "survivorship of the better," as though that were the statement of the law; and then adds that the alleged result cannot be inferred "except on the assumption that whatever is better is stronger too." But the words he here uses are his own words, not the words of those he opposes. The law is the survival of the fittest. Probably, in substituting "better" for "fittest," Mr. Martineau did not suppose that he was changing the meaning; though I dare say he perceived that the meaning of the word "fittest" did not suit his argument so well. Had he examined the facts, he would have found that the law is not the survival of the "better" or the "stronger," if we give to those words any thing like their ordinary meanings. It is the survival of those which are constitutionally fittest to thrive under the conditions in which they are placed; and very often that which, humanly speaking, is inferiority, causes the survival. Superiority, whether in size, strength, activity, or sagacity, is, other things equal, at the cost of diminished fertility; and where the life led by a species does not demand these higher attributes, the species profits by decrease of them, and accompanying increase of fertility. This is the reason why there occur so many cases of retrograde metamorphosis—this is the reason why parasites, internal and external, are so commonly degraded forms of higher types. Survival of the "better" does not cover these cases, though survival of the "fittest" does; and as I am responsible for the phrase, I suppose I am competent to say that the word "fittest" was chosen for this reason. When it is remembered that these cases outnumber all others—that there are more species of parasites than there are species of all other animals put together—it will be seen that the expression "survivorship of the better" is wholly inappropriate, and the argument Mr. Martineau bases upon it quite untenable. Indeed, if, in place of those adjustments of the human sense-organs, which he so eloquently describes as implying pre-arrangement, Mr. Martineau had described the countless elaborate appliances which enable parasites to torture animals immeasurably superior to them, and which, from his point of view, no less imply pre-arrangement, I think the notes of admiration which end his descriptions would not have seemed to him so appropriate.
One more word there is from the intrinsic meaning of which Mr. Martineau deduces what appears a powerful argument—the word Evolution itself. He says:—
"It means, to unfold from within; and it is taken from the history of the seed or embryo of living natures. And what is the seed but a casket of pre-arranged futurities, with its whole contents prospective, settled to be what they are by reference to ends still in the distance?"
Now, this criticism would have been very much to the point did the word Evolution truly express the process it names. If this process, as scientifically defined, really involved that conception which the word evolution was originally designed to convey, the implications would be those Mr. Martineau alleges. But, unfortunately for him, the word, having been in possession of the field before the process was understood, has been adopted merely because displacing it by another word seemed impracticable. And this adoption of it has been joined with a caution against misunderstandings arising from its unfitness. Here is a part of the caution:—"Evolution has other meanings, some of which are incongruous with, and some even directly opposed to, the meaning here given to it.... The antithetical word, Involution, would much more truly express the nature of the process; and would, indeed, describe better the secondary characters of the process which we shall have to deal with presently."[38] So that the meanings which the word involves, and which Mr. Martineau regards as fatal to the hypothesis, are already repudiated as not belonging to the hypothesis.
* * * * *
And now, having dealt with the essential objections raised by Mr. Martineau to the Hypothesis of Evolution as it is presented under that purely scientific form which generalizes the process of things, firstly as observed and secondly as inferred from certain ultimate principles, let me go on to examine that form of the Hypothesis which he propounds—Evolution as determined by Mind and Will—Evolution as pre-arranged by a Divine Actor. For Mr. Martineau apparently abandons the primitive theory of creation by "fiat of Almighty Will", and also the theory of creation by manufacture—by "a contriving and adapting power," and seems to believe in evolution: requiring only that "an originating Mind" shall be taken as its antecedent. Let us ask, first, in what relation Mr. Martineau conceives the "originating Mind" to stand to the evolving Universe. From some passages it is inferable that he considers the "presence of mind" to be everywhere needful. He says:—
"It is impossible to work the theory of Evolution upwards from the bottom. If all force is to be conceived as One, its type must be looked for in the highest and all-comprehending term; and Mind must be conceived as there, and as divesting itself of some speciality at each step of its descent to a lower stratum of law, till represented at the base under the guise of simple Dynamics."
This seems to be an unmistakable assertion that, wherever Evolution is going on, Mind is then and there behind it. At the close of the argument, however, a quite different conception is implied. Mr. Martineau says:—
"If the Divine Idea will not retire at the bidding of our speculative science, but retains its place, it is natural to ask, What is its relation to the series of so-called Forces in the world? But the question is too large and deep to be answered here. Let it suffice to say, that there need not be any overruling of these forces by the Will of God, so that the supernatural should disturb the natural; or any supplementing of them, so that He should fill up their deficiencies. Rather is His thought related to them as, in man, the mental force is related to all below it."
It would take too much space to deal fully with the various questions which this last passage raises. There is the question—Whence come these "Forces," spoken of as separate from the "Will of God"—did they pre-exist? Then what becomes of the Divine Power? Do they exist by the Divine Will? Then what kind of nature is that by which they act apart from the Divine Will? Again, there is the question—How do these deputy-forces co-operate in each particular phenomenon, if the presiding Will is not there present to control them? Either an organ which develops into fitness for its function, develops by the co-operation of these forces under the direction of Mind then present, or it so develops in the absence of Mind. If it develops in the absence of Mind, the hypothesis is given up; and if the "originating Mind" is required to be then and there present, we must suppose a particular providence to be present in each particular organ of each particular creature throughout the universe. Once more there is the question—If "His thought is related to them [these Forces] as, in Man, the mental force is related to all below it," how can "His thought" be regarded as the cause of Evolution? In man the mental force is related to the forces below it neither as a creator of them nor as a regulator of them, save in a very limited way: the greater part of the forces present in man, both structural and functional, defy the mental force absolutely. Nay, more, it needs but to injure a nerve to see that the power of the mental force over the physical forces is dependent on conditions which are themselves physical; and one who takes morphia in mistake for magnesia, discovers that the power of the physical forces over the mental is unconditioned by any thing mental.
Not dwelling on these questions, however, I will merely draw attention to the entire incongruity of this conception with the previous conception which I have quoted. Assuming that, when the choice is pressed on him, Mr. Martineau will choose the first, which alone has any thing like defensibility, let us go on to ask how far Evolution is made more comprehensible by postulating Mind, universally immanent, as its cause.
In metaphysical controversy, many of the propositions propounded and accepted as quite believable, are absolutely inconceivable. There is a perpetual confusing of actual ideas with what are nothing but pseud-ideas. No distinction is made between propositions that contain real thoughts, and propositions that are only the forms of thoughts. A thinkable proposition is one of which the two terms can be brought together in consciousness under the relation said to exist between them. But very often, when the subject of a proposition has been thought of as something known, and when the predicate has been thought of as something known, and when the relation alleged between them has been thought of as a known relation, it is supposed that the proposition itself has been thought. The thinking separately of the elements of a proposition is mistaken for the thinking of them in the combination which the proposition affirms. And hence it continually happens that propositions which cannot be rendered into thought at all, are supposed to be not only thought but believed. The proposition that Evolution is caused by Mind is one of this nature. The two terms are separately intelligible; but they can be regarded in the relation of effect and cause only so long as no attempt is made to put them together in this relation.
The only thing which any one knows as Mind is the series of his own states of consciousness; and if he thinks of any mind other than his own, he can think of it only in terms derived from his own. If I am asked to frame a notion of Mind divested of all those structural traits under which alone I am conscious of mind in myself, I cannot do it. I know nothing of thought save as carried on in ideas originally traceable to the effects wrought by objects and forces on me. A mental act is an unintelligible phrase if I am not to regard it as an act in which states of consciousness are severally known as like other states in the series that has gone by, and in which the relations between them are severally known as like past relations in the series. If, then, I have to conceive Evolution as caused by an "originating Mind," I must conceive this Mind as having attributes akin to those of the only mind I know, and without which I cannot conceive Mind at all.
I will not dwell on the many incongruities hence resulting, by asking how the "originating Mind" is to be thought of as having states produced by things objective to it; as discriminating among these states, and classing them as like and unlike; and as preferring one objective result to another. I will simply ask—What happens if we ascribe to the "originating Mind" the character absolutely essential to the conception of Mind, that it consists of a series of states of consciousness? Put a series of states of consciousness as cause, and the evolving Universe as effect, and then endeavor to see the last as flowing from the first. I find it possible to imagine in some dim way a series of states of consciousness serving as antecedent to any one of the movements I see going on; for my own states of consciousness are often indirectly the antecedents to such movements. But how if I attempt to think of such a series as antecedent to all actions throughout the Universe—to the motions of the multitudinous stars through space, to the revolutions of all their planets round them, to the gyrations of all these planets on their axes, to the infinitely-multiplied physical processes going on in each of these suns and planets? I cannot think of a single series of states of consciousness as causing even the relatively small group of actions going on over the Earth's surface. I cannot think of it even as antecedent to all the various winds and the dissolving clouds they bear, to the currents of all the rivers, and the grinding actions of all the glaciers; still less can I think of it as antecedent to the infinity of processes simultaneously going on in all the plants that cover the globe, from scattered polar lichens to crowded tropical palms, and in all the millions of quadrupeds that roam among them, and the millions of millions of insects that buzz about them. Even to a single small set of these multitudinous terrestrial changes, I cannot conceive as antecedent a single series of states of consciousness—cannot, for instance, think of it as causing the hundred thousand breakers that are at this instant curling over on the shores of England. How, then, is it possible for me to conceive an "originating Mind," which I must represent to myself as a single series of states of consciousness, working the infinitely-multiplied sets of changes simultaneously going on in worlds too numerous to count, dispersed throughout a space that baffles imagination?
If, to account for this infinitude of physical changes everywhere going on, "Mind must be conceived as there" "under the guise of simple Dynamics," then the reply is that, to be so conceived, Mind must be divested of all attributes by which it is distinguished; and that, when thus divested of its distinguishing attributes, the conception disappears—the word Mind stands for a blank. If Mr. Martineau takes refuge in the entirely different and, as it seems to me, incongruous hypothesis of something like a plurality of minds—if he accepts, as he seems to do, the doctrine that you cannot explain Evolution "unless among your primordial elements you scatter already the germs of Mind as well as the inferior elements"—if the insuperable difficulties I have just pointed out are to be met by assuming a local series of states of consciousness for each phenomenon, then we are obviously carried back to something like the alleged fetichistic notion, with the difference only, that the assumed spiritual agencies are indefinitely multiplied.
Clearly, therefore, the proposition that an "originating Mind" is the cause of Evolution, is a proposition that can be entertained so long only as no attempt is made to unite in thought its two terms in the alleged relation. That it should be accepted as a matter of faith, may be a defensible position, provided good cause is shown why it should be so accepted; but that it should be accepted as a matter of understanding—as a statement making the order of the universe comprehensible—is a quite indefensible position.
* * * * *
Here let me guard myself against a misinterpretation very likely to be put upon the foregoing arguments; especially by those who have read the Essay to which they reply. The statements of that Essay carry the implication that all who adhere to the hypothesis it combats, imagine they have solved the mystery of things when they have shown the processes of Evolution to be naturally caused. Mr. Martineau tacitly represents them as believing that, when every thing has been interpreted in terms of Matter and Motion, nothing remains to be explained. This, however, is by no means the fact. The Doctrine of Evolution, under its purely scientific form, does not involve Materialism, though its opponents persistently represent it as doing so. Indeed, among adherents of it who are friends of mine, there are those who speak of the Materialism of Buechner and his school, with a contempt certainly not less than that felt by Mr. Martineau. To show how anti-materialistic my own view is, I may, perhaps, without impropriety, quote some out of many passages which I have written on the question elsewhere:
"Hence though of the two it seems easier to translate so-called Matter into so-called Spirit, than to translate so-called Spirit into so-called Matter (which latter is, indeed, wholly impossible); yet no translation can carry us beyond our symbols."[39]
And again:
"See then our predicament. We can think of Matter only in terms of Mind. We can think of Mind only in terms of Matter. When we have pushed our explorations of the first to the uttermost limit, we are referred to the second for a final answer; and, when we have got the final answer of the second, we are referred back to the first for an interpretation of it. We find the value of x in terms of y; then we find the value of y in terms of x; and so on we may continue forever without coming nearer to a solution. The antithesis of subject and object, never to be transcended while consciousness lasts, renders impossible all knowledge of that Ultimate Reality in which subject and object are united."[40]
It is thus, I think, manifest that the difference between Mr. Martineau's view and the view he opposes is by no means so wide as he makes it appear; and further, it seems to me that such difference as exists is rather the reverse of that indicated by his exposition. Briefly expressed, the difference is that, where he thinks there is no mystery, the doctrine he combats recognizes a mystery. Speaking for myself only, I may say that, agreeing entirely with Mr. Martineau in repudiating the materialistic interpretation as utterly futile, I differ from him simply in this, that while he says he has found another interpretation, I confess that I cannot find any interpretation; while he holds that he can understand the Power which is manifested in things, I feel obliged to admit, after many failures, that I cannot understand it. So that, in presence of the transcendent problem which the universe presents, Mr. Martineau regards the human intellect as capable, and I as incapable. This contrast does not appear to me of the kind which his Essay tacitly asserts. If there is such a thing as the "pride of Science," it is obviously exceeded by the pride of Theology. I fail to perceive humility in the belief that the human mind is able to comprehend that which is behind appearances; and I do not see how piety is especially exemplified in the assertion that the Universe contains no mode of existence higher in Nature than that which is present to us in consciousness. On the contrary, I think it quite a defensible proposition that humility is better shown by a confession of incompetence to grasp in thought the Cause of all things; and that the religious sentiment may find its highest sphere in the belief that the Ultimate Power is no more representable in terms of human consciousness than human consciousness is representable in terms of a plant's functions.
FOOTNOTES:
[Footnote 37: Principles of Biology, Sec.Sec. 159-168.]
[Footnote 38: First Principles, second edition, Sec. 97.]
[Footnote 39: Principles of Psychology, second edition, vol. i., Sec. 63.]
[Footnote 40: Ibid., Sec. 272.]
THE FACTORS OF ORGANIC EVOLUTION.
[First published in The Nineteenth Century, for April and May, 1886.]
I.
Within the recollection of men now in middle life, opinion concerning the derivation of animals and plants was in a chaotic state. Among the unthinking there was tacit belief in creation by miracle, which formed an essential part of the creed of Christendom; and among the thinking there were two parties, each of which held an indefensible hypothesis. Immensely the larger of these parties, including nearly all whose scientific culture gave weight to their judgments, though not accepting literally the theologically-orthodox doctrine, made a compromise between that doctrine and the doctrines which geologists had established; while opposed to them were some, mostly having no authority in science, who held a doctrine which was heterodox both theologically and scientifically. Professor Huxley, in his lecture on "The Coming of Age of the Origin of Species," remarks concerning the first of these parties as follows:—
"One-and-twenty years ago, in spite of the work commenced by Hutton and continued with rare skill and patience by Lyell, the dominant view of the past history of the earth was catastrophic. Great and sudden physical revolutions, wholesale creations and extinctions of living beings, were the ordinary machinery of the geological epic brought into fashion by the misapplied genius of Cuvier. It was gravely maintained and taught that the end of every geological epoch was signalised by a cataclysm, by which every living being on the globe was swept away, to be replaced by a brand-new creation when the world returned to quiescence. A scheme of nature which appeared to be modelled on the likeness of a succession of rubbers of whist, at the end of each of which the players upset the table and called for a new pack, did not seem to shock anybody.
I may be wrong, but I doubt if, at the present time, there is a single responsible representative of these opinions left. The progress of scientific geology has elevated the fundament principle of uniformitarianism, that the explanation of the past is to be sought in the study of the present, into the position of an axiom; and the wild speculations of the catastrophists, to which we all listened with respect a quarter of a century ago, would hardly find a single patient hearer at the present day."
Of the party above referred to as not satisfied with this conception described by Professor Huxley, there were two classes. The great majority were admirers of the Vestiges of the Natural History of Creation—a work which, while it sought to show that organic evolution has taken place, contended that the cause of organic evolution, is "an impulse" supernaturally "imparted to the forms of life, advancing them, ... through grades of organization." Being nearly all very inadequately acquainted with the facts, those who accepted the view set forth in the Vestiges were ridiculed by the well-instructed for being satisfied with evidence, much of which was either invalid or easily cancelled by counter-evidence, and at the same time they exposed themselves to the ridicule of the more philosophical for being content with a supposed explanation which was in reality no explanation: the alleged "impulse" to advance giving us no more help in understanding the facts than does Nature's alleged "abhorrence of a vacuum" help us to understand the ascent of water in a pump. The remnant, forming the second of these classes, was very small. While rejecting this mere verbal solution, which both Dr. Erasmus Darwin and Lamarck had shadowed forth in other language, there were some few who, rejecting also the hypothesis indicated by both Dr. Darwin and Lamarck, that the promptings of desires or wants produced growths of the parts subserving them, accepted the single vera causa assigned by these writers—the modification of structures resulting from modification of functions. They recognized as the sole process in organic development, the adaptation of parts and powers consequent on the effects of use and disuse—that continual moulding and re-moulding of organisms to suit their circumstances, which is brought about by direct converse with such circumstances.
But while this cause accepted by these few is a true cause, since unquestionably during the life of the individual organism changes of function produce changes of structure; and while it is a tenable hypothesis that changes of structure so produced are inheritable; yet it was manifest to those not prepossessed, that this cause cannot with reason be assigned for the greater part of the facts. Though in plants there are some characters which may not irrationally be ascribed to the direct effects of modified functions consequent on modified circumstances, yet the majority of the traits presented by plants are not to be thus explained. It is impossible that the thorns by which a briar is in large measure defended against browsing animals, can have been developed and moulded by the continuous exercise of their protective actions; for in the first place, the great majority of the thorns are never touched at all, and, in the second place, we have no ground whatever for supposing that those which are touched are thereby made to grow, and to take those shapes which render them efficient. Plants which are rendered uneatable by the thick woolly coatings of their leaves, cannot have had these coatings produced by any process of reaction against the action of enemies; for there is no imaginable reason why, if one part of a plant is eaten, the rest should thereafter begin to develop the hairs on its surface. By what direct effect of function on structure, can the shell of a nut have been evolved? Or how can those seeds which contain essential oils, rendering them unpalatable to birds, have been made to secrete such essential oils by these actions of birds which they restrain? Or how can the delicate plumes borne by some seeds, and giving the wind power to waft them to new stations, be due to any immediate influences of surrounding conditions? Clearly in these and in countless other cases, change of structure cannot have been directly caused by change of function. So is it with animals to a large extent, if not to the same extent. Though we have proof that by rough usage the dermal layer may be so excited as to produce a greatly thickened epidermal layer, sometimes quite horny; and though it is a feasible hypothesis that an effect of this kind persistently produced may be inherited; yet no such cause can explain the carapace of the turtle, the armour of the armadillo, or the imbricated covering of the manis. The skins of these animals are no more exposed to habitual hard usage than are those of animals covered by hair. The strange excrescences which distinguish the heads of the hornbills, cannot possibly have arisen from any reaction against the action of surrounding forces; for even were they clearly protective, there is no reason to suppose that the heads of these birds need protection more than the heads of other birds. If, led by the evidence that in animals the amount of covering is in some cases affected by the degree of exposure, it were admitted as imaginable that the development of feathers from preceding dermal growths had resulted from that extra nutrition caused by extra superficial circulation, we should still be without explanation of the structure of a feather. Nor should we have any clue to the specialities of feathers—the crests of various birds, the tails sometimes so enormous, the curiously placed plumes of the bird of paradise, &c., &c. Still more obviously impossible is it to explain as due to use or disuse the colours of animals. No direct adaptation to function could have produced the blue protuberances on a mandril's face, or the striped hide of a tiger, or the gorgeous plumage of a kingfisher, or the eyes in a peacock's tail, or the multitudinous patterns of insects' wings. One single case, that of a deer's horns, might alone have sufficed to show how insufficient was the assigned cause. During their growth, a deer's horns are not used at all; and when, having been cleared of the dead skin and dried-up blood-vessels covering them, they are ready for use, they are nerveless and non-vascular, and hence are incapable of undergoing any changes of structure consequent on changes of function.
Of these few then, who rejected the belief described by Professor Huxley, and who, espousing the belief in a continuous evolution, had to account for this evolution, it must be said that though the cause assigned was a true cause, yet, even admitting that it operated through successive generations, it left unexplained the greater part of the facts. Having been myself one of these few, I look back with surprise at the way in which the facts which were congruous with the espoused view monopolized consciousness and kept out the facts which were incongruous with it—conspicuous though many of them were. The misjudgment was not unnatural. Finding it impossible to accept any doctrine which implied a breach in the uniform course of natural causation, and, by implication, accepting as unquestionable the origin and development of all organic forms by accumulated modifications naturally caused, that which appeared to explain certain classes of these modifications, was supposed to be capable of explaining the rest: the tendency being to assume that these would eventually be similarly accounted for, though it was not clear how.
Returning from this parenthetic remark, we are concerned here chiefly to remember that, as said at the outset, there existed thirty years ago, no tenable theory about the genesis of living things. Of the two alternative beliefs, neither would bear critical examination.
* * * * *
Out of this dead lock we were released—in large measure, though not I believe entirely—by the Origin of Species. That work brought into view a further factor; or rather, such factor, recognized as in operation by here and there an observer (as pointed out by Mr. Darwin in his introduction to the second edition), was by him for the first time seen to have played so immense a part in the genesis of plants and animals.
Though laying myself open to the charge of telling a thrice-told tale, I feel obliged here to indicate briefly the several great classes of facts which Mr. Darwin's hypothesis explains; because otherwise that which follows would scarcely be understood. And I feel the less hesitation in doing this because the hypothesis which it replaced, not very widely known at any time, has of late so completely dropped into the background, that the majority of readers are scarcely aware of its existence, and do not therefore understand the relation between Mr. Darwin's successful interpretation and the preceding unsuccessful attempt at interpretation. Of these classes of facts, four chief ones may be here distinguished.
In the first place, such adjustments as those exemplified above are made comprehensible. Though it is inconceivable that a structure like that of the pitcher-plant could have been produced by accumulated effects of function on structure; yet it is conceivable that successive selections of favourable variations might have produced it; and the like holds of the no less remarkable appliance of the Venus's Fly-trap, or the still more astonishing one of that water-plant by which infant-fish are captured. Though it is impossible to imagine how, by direct influence of increased use, such dermal appendages as a porcupine's quills could have been developed; yet, profiting as the members of a species otherwise defenceless might do by the stiffness of their hairs, rendering them unpleasant morsels to eat, it is a feasible supposition that from successive survivals of individuals thus defended in the greatest degrees, and the consequent growth in successive generations of hairs into bristles, bristles into spines, spines into quills (for all these are homologous), this change could have arisen. In like manner, the odd inflatable bag of the bladder-nosed seal, the curious fishing-rod with its worm-like appendage carried on the head of the lophius or angler, the spurs on the wings of certain birds, the weapons of the sword-fish and saw-fish, the wattles of fowls, and numberless such peculiar structures, though by no possibility explicable as due to effects of use or disuse, are explicable as resulting from natural selection operating in one or other way.
In the second place, while showing us how there have arisen countless modifications in the forms, structures, and colours of each part, Mr. Darwin has shown us how, by the establishment of favourable variations, there may arise new parts. Though the first step in the production of horns on the heads of various herbivorous animals, may have been the growth of callosities consequent on the habit of butting—such callosities thus functionally initiated being afterwards developed in the most advantageous ways by selection; yet no explanation can be thus given of the sudden appearance of a duplicate set of horns, as occasionally happens in sheep: an addition which, where it proved beneficial, might readily be made a permanent trait by natural selection. Again, the modifications which follow use and disuse can by no possibility account for changes in the numbers of vertebrae; but after recognizing spontaneous, or rather fortuitous, variation as a factor, we can see that where an additional vertebra hence resulting (as in some pigeons) proves beneficial, survival of the fittest may make it a constant character; and there may, by further like additions, be produced extremely long strings of vertebrae, such as snakes show us. Similarly with the mammary glands. It is not an unreasonable supposition that by the effects of greater or less function, inherited through successive generations, these may be enlarged or diminished in size; but it is out of the question to allege such a cause for changes in their numbers. There is no imaginable explanation of these save the establishment by inheritance of spontaneous variations, such as are known to occur in the human race.
So too, in the third place, with certain alterations in the connexions of parts. According to the greater or smaller demands made on this or that limb, the muscles moving it may be augmented or diminished in bulk; and, if there is inheritance of changes so wrought, the limb may, in course of generations, be rendered larger or smaller. But changes in the arrangements or attachments of muscles cannot be thus accounted for. It is found, especially at the extremities, that the relations of tendons to bones and to one another are not always the same. Variations in their modes of connexion may occasionally prove advantageous, and may thus become established. Here again, then, we have a class of structural changes to which Mr. Darwin's hypothesis gives us the key, and to which there is no other key.
Once more there are the phenomena of mimicry. Perhaps in a more striking way than any others, these show how traits which seem inexplicable are explicable as due to the more frequent survival of individuals that have varied in favourable ways. We are enabled to understand such marvellous simulations as those of the leaf-insect, those of beetles which "resemble glittering dew-drops upon the leaves;" those of caterpillars which, when asleep, stretch themselves out so as to look like twigs. And we are shown how there have arisen still more astonishing imitations—those of one insect by another. As Mr. Bates has proved, there are cases in which a species of butterfly, rendered so unpalatable to insectivorous birds by its disagreeable taste that they will not catch it, is simulated in its colours and markings by a species which is structurally quite different—so simulated that even a practised entomologist is liable to be deceived: the explanation being that an original slight resemblance, leading to occasional mistakes on the part of birds, was increased generation after generation by the more frequent escape of the most-like individuals, until the likeness became thus great.
But now, recognizing in full this process brought into clear view by Mr. Darwin, and traced out by him with so much care and skill, can we conclude that, taken alone, it accounts for organic evolution? Has the natural selection of favourable variations been the sole factor? On critically examining the evidence, we shall find reason to think that it by no means explains all that has to be explained. Omitting for the present any consideration of a factor which may be distinguished as primordial, it may be contended that the above-named factor alleged by Dr. Erasmus Darwin and by Lamarck, must be recognized as a co-operator. Utterly inadequate to explain the major part of the facts as is the hypothesis of the inheritance of functionally-produced modifications, yet there is a minor part of the facts, very extensive though less, which must be ascribed to this cause.
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When discussing the question more than twenty years ago (Principles of Biology, Sec. 166), I instanced the decreased size of the jaws in the civilized races of mankind, as a change not accounted for by the natural selection of favourable variations; since no one of the decrements by which, in thousands of years, this reduction has been effected, could have given to an individual in which it occurred, such advantage as would cause his survival, either through diminished cost of local nutrition or diminished weight to be carried. I did not then exclude, as I might have done, two other imaginable causes. It may be said that there is some organic correlation between increased size of brain and decreased size of jaw: Camper's doctrine of the facial angle being referred to in proof. But this argument may be met by pointing to the many examples of small-jawed people who are also small-brained, and by citing not infrequent cases of individuals remarkable for their mental powers, and at the same time distinguished by jaws not less than the average but greater. Again, if sexual selection be named as a possible cause, there is the reply that, even supposing such slight diminution of jaw as took place in a single generation to have been an attraction, yet the other incentives to choice on the part of men have been too many and great to allow this one to weigh in an adequate degree; while, during the greater portion of the period, choice on the part of women has scarcely operated: in earlier times they were stolen or bought, and in later times mostly coerced by parents. Thus, reconsideration of the facts does not show me the invalidity of the conclusion drawn, that this decrease in size of jaw can have had no other cause than continued inheritance of those diminutions consequent on diminutions of function, implied by the use of selected and well-prepared food. Here, however, my chief purpose is to add an instance showing, even more clearly, the connexion between change of function and change of structure. This instance, allied in nature to the other, is presented by those varieties, or rather sub-varieties, of dogs, which, having been household pets, and habitually fed on soft food, have not been called on to use their jaws in tearing and crunching, and have been but rarely allowed to use them in catching prey and in fighting. No inference can be drawn from the sizes of the jaws themselves, which, in these dogs, have probably been shortened mainly by selection. To get direct proof of the decrease of the muscles concerned in closing the jaws or biting, would require a series of observations very difficult to make. But it is not difficult to get indirect proof of this decrease by looking at the bony structures with which these muscles are connected. Examination of the skulls of sundry indoor dogs contained in the Museum of the College of Surgeons, proves the relative smallness of such parts. The only pug-dog's skull is that of an individual not perfectly adult; and though its traits are quite to the point they cannot with safety be taken as evidence. The skull of a toy-terrier has much restricted areas of insertion for the temporal muscles; has weak zygomatic arches; and has extremely small attachments for the masseter muscles. Still more significant is the evidence furnished by the skull of a King Charles's spaniel, which, if we allow three years to a generation, and bear in mind that the variety must have existed before Charles the Second's reign, we may assume belongs to something approaching to the hundredth generation of these household pets. The relative breadth between the outer surfaces of the zygomatic arches is conspicuously small; the narrowness of the temporal fossae is also striking; the zygomata are very slender; the temporal muscles have left no marks whatever, either by limiting lines or by the character of the surfaces covered; and the places of attachment for the masseter muscles are very feebly developed. At the Museum of Natural History, among skulls of dogs there is one which, though unnamed, is shown by its small size and by its teeth, to have belonged to one variety or other of lap-dogs, and which has the same traits in an equal degree with the skull just described. Here, then, we have two if not three kinds of dogs which, similarly leading protected and pampered lives, show that in the course of generations the parts concerned in clenching the jaws have dwindled. To what cause must this decrease be ascribed? Certainly not to artificial selection; for most of the modifications named make no appreciable external signs: the width across the zygomata could alone be perceived. Neither can natural selection have had anything to do with it; for even were there any struggle for existence among such dogs, it cannot be contended that any advantage in the struggle could be gained by an individual in which a decrease took place. Economy of nutrition, too, is excluded. Abundantly fed as such dogs are, the constitutional tendency is to find places where excess of absorbed nutriment may be conveniently deposited, rather than to find places where some cutting down of the supplies is practicable. Nor again can there be alleged a possible correlation between these diminutions and that shortening of the jaws which has probably resulted from selection; for in the bull-dog, which has also relatively short jaws, these structures concerned in closing them are unusually large. Thus there remains as the only conceivable cause, the diminution of size which results from diminished use. The dwindling of a little-exercised part has, by inheritance, been made more and more marked in successive generations.
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Difficulties of another class may next be exemplified—those which present themselves when we ask how there can be effected by the selection of favourable variations, such changes of structure as adapt an organism to some useful action in which many different parts co-operate. None can fail to see how a simple part may, in course of generations, be greatly enlarged, if each enlargement furthers, in some decided way, maintenance of the species. It is easy to understand, too, how a complex part, as an entire limb, may be increased as a whole by the simultaneous due increase of its co-operative parts; since if, while it is growing, the channels of supply bring to the limb an unusual quantity of blood, there will naturally result a proportionately greater size of all its components—bones, muscles, arteries, veins, &c. But though in cases like this, the co-operative parts forming some large complex part may be expected to vary together, nothing implies that they necessarily do so; and we have proof that in various cases, even when closely united, they do not do so. An example is furnished by those blind crabs named in the Origin of Species which inhabit certain dark caves of Kentucky, and which, though they have lost their eyes, have not lost the foot-stalks which carried their eyes. In describing the varieties which have been produced by pigeon-fanciers, Mr. Darwin notes the fact that along with changes in length of beak produced by selection, there have not gone proportionate changes in length of tongue. Take again the case of teeth and jaws. In mankind these have not varied together. During civilization the jaws have decreased, but the teeth have not decreased in proportion; and hence that prevalent crowding of them, often remedied in childhood by extraction of some, and in other cases causing that imperfect development which is followed by early decay. But the absence of proportionate variation in co-operative parts that are close together, and are even bound up in the same mass, is best seen in those varieties of dogs named above as illustrating the inherited effects of disuse. We see in them, as we see in the human race, that diminution in the jaws has not been accompanied by corresponding diminution in the teeth. In the catalogue of the College of Surgeons Museum, there is appended to the entry which identifies a Blenheim Spaniel's skull, the words—"the teeth are closely crowded together," and to the entry concerning the skull of a King Charles's Spaniel the words—"the teeth are closely packed, p. 3, is placed quite transversely to the axis of the skull." It is further noteworthy that in a case where there is no diminished use of the jaws, but where they have been shortened by selection, a like want of concomitant variation is manifested: the case being that of the bull-dog, in the upper jaw of which also, "the premolars ... are excessively crowded, and placed obliquely or even transversely to the long axis of the skull."[41]
If, then, in cases where we can test it, we find no concomitant variation in co-operative parts that are near together—if we do not find it in parts which, though belonging to different tissues, are so closely united as teeth and jaws—if we do not find it even when the co-operative parts are not only closely united, but are formed out of the same tissue, like the crab's eye and its peduncle; what shall we say of co-operative parts which, besides being composed of different tissues, are remote from one another? Not only are we forbidden to assume that they vary together, but we are warranted in asserting that they can have no tendency to vary together. And what are the implications in cases where increase of a structure can be of no service unless there is concomitant increase in many distant structures, which have to join it in performing the action for which it is useful?
As far back as 1864 (Principles of Biology, Sec. 166) I named in illustration an animal carrying heavy horns—the extinct Irish elk; and indicated the many changes in bones, muscles, blood-vessels, nerves, composing the fore-part of the body, which would be required to make an increment of size in such horns advantageous. Here let me take another instance—that of the giraffe: an instance which I take partly because, in the sixth edition of the Origin of Species, issued in 1872, Mr. Darwin has referred to this animal when effectually disposing of certain arguments urged against his hypothesis. He there says:—
"In order that an animal should acquire some structure specially and largely developed, it is almost indispensable that several other parts should be modified and co-adapted. Although every part of the body varies slightly, it does not follow that the necessary parts should always vary in the right direction and to the right degree" (p. 179).
And in the summary of the chapter, he remarks concerning the adjustments in the same quadruped, that "the prolonged use of all the parts together with inheritance will have aided in an important manner in their co-ordination" (p. 199): a remark probably having reference chiefly to the increased massiveness of the lower part of the neck; the increased size and strength of the thorax required to bear the additional burden; and the increased strength of the fore-legs required to carry the greater weight of both. But now I think that further consideration suggests the belief that the entailed modifications are much more numerous and remote than at first appears; and that the greater part of these are such as cannot be ascribed in any degree to the selection of favourable variations, but must be ascribed exclusively to the inherited effects of changed functions. Whoever has seen a giraffe gallop will long remember the sight as a ludicrous one. The reason for the strangeness of the motions is obvious. Though the fore limbs and the hind limbs differ so much in length, yet in galloping they have to keep pace—must take equal strides. The result is that at each stride, the angle which the hind limbs describe round their centre of motion is much larger than the angle described by the fore limbs. And beyond this, as an aid in equalizing the strides, the hind part of the back is at each stride bent very much downwards and forwards. Hence the hind-quarters appear to be doing nearly all the work. Now a moment's observation shows that the bones and muscles composing the hind-quarters of the giraffe, perform actions differing in one or other way and degree, from the actions performed by the homologous bones and muscles in a mammal of ordinary proportions, and from those in the ancestral mammal which gave origin to the giraffe. Each further stage of that growth which produced the large fore-quarters and neck, entailed some adapted change in sundry of the numerous parts composing the hind-quarters; since any failure in the adjustment of their respective strengths would entail some defect in speed and consequent loss of life when chased. It needs but to remember how, when continuing to walk with a blistered foot, the taking of steps in such a modified way as to diminish pressure on the sore point, soon produces aching of muscles which are called into unusual action, to see that over-straining of any one of the muscles of the giraffe's hind-quarters might quickly incapacitate the animal when putting out all its powers to escape; and to be a few yards behind others would cause death. Hence if we are debarred from assuming that co-operative parts vary together even when adjacent and closely united—if we are still more debarred from assuming that with increased length of fore-legs or of neck, there will go an appropriate change in any one muscle or bone in the hind-quarters; how entirely out of the question it is to assume that there will simultaneously take place the appropriate changes in all those many components of the hind-quarters which severally require re-adjustment. It is useless to reply that an increment of length in the fore-legs or neck might be retained and transmitted to posterity, waiting an appropriate variation in a particular bone or muscle in the hind-quarters, which, being made, would allow of a further increment. For besides the fact that until this secondary variation occurred the primary variation would be a disadvantage often fatal; and besides the fact that before such an appropriate secondary variation might be expected in the course of generations to occur, the primary variation would have died out; there is the fact that the appropriate variation of one bone or muscle in the hind-quarters would be useless without appropriate variations of all the rest—some in this way and some in that—a number of appropriate variations which it is impossible to suppose.
Nor is this all. Far more numerous appropriate variations would be indirectly necessitated. The immense change in the ratio of fore-quarters to hind-quarters would make requisite a corresponding change of ratio in the appliances carrying on the nutrition of the two. The entire vascular system, arterial and veinous, would have to undergo successive unbuildings and rebuildings to make its channels everywhere adequate to the local requirements; since any want of adjustment in the blood-supply in this or that set of muscles, would entail incapacity, failure of speed, and loss of life. Moreover the nerves supplying the various sets of muscles would have to be proportionately changed; as well as the central nervous tracts from which they issued. Can we suppose that all these appropriate changes, too, would be step by step simultaneously made by fortunate spontaneous variations, occurring along with all the other fortunate spontaneous variations? Considering how immense must be the number of these required changes, added to the changes above enumerated, the chances against any adequate re-adjustments fortuitously arising must be infinity to one.
If the effects of use and disuse of parts are inheritable, then any change in the fore parts of the giraffe which affects the action of the hind limbs and back, will simultaneously cause, by the greater or less exercise of it, a re-moulding of each component in the hind limbs and back in a way adapted to the new demands; and generation after generation the entire structure of the hind-quarters will be progressively fitted to the changed structure of the fore-quarters: all the appliances for nutrition and innervation being at the same time progressively fitted to both. But in the absence of this inheritance of functionally-produced modifications, there is no seeing how the required re-adjustments can be made.
* * * * *
Yet a third class of difficulties stands in the way of the belief that the natural selection of useful variations is the sole factor of organic evolution. This class of difficulties, already pointed out in Sec. 166 of the Principles of Biology, I cannot more clearly set forth than in the words there used. Hence I may perhaps be excused for here quoting them.
"Where the life is comparatively simple, or where surrounding circumstances render some one function supremely important, the survival of the fittest may readily bring about the appropriate structural change, without any aid from the transmission of functionally-acquired modifications. But in proportion as the life grows complex—in proportion as a healthy existence cannot be secured by a large endowment of some one power, but demands many powers; in the same proportion do there arise obstacles to the increase of any particular power, by 'the preservation of favoured races in the struggle for life.' As fast as the faculties are multiplied, so fast does it become possible for the several members of a species to have various kinds of superiorities over one another. While one saves its life by higher speed, another does the like by clearer vision, another by keener scent, another by quicker hearing, another by greater strength, another by unusual power of enduring cold or hunger, another by special sagacity, another by special timidity, another by special courage; and others by other bodily and mental attributes. Now it is unquestionably true that, other things equal, each of these attributes, giving its possessor an extra chance of life, is likely to be transmitted to posterity. But there seems no reason to suppose that it will be increased in subsequent generations by natural selection. That it may be thus increased, the individuals not possessing more than average endowments of it, must be more frequently killed off than individuals highly endowed with it; and this can happen only when the attribute is one of greater importance, for the time being, than most of the other attributes. If those members of the species which have but ordinary shares of it, nevertheless survive by virtue of other superiorities which they severally possess; then it is not easy to see how this particular attribute can be developed by natural selection in subsequent generations. The probability seems rather to be, that by gamogenesis, this extra endowment will, on the average, be diminished in posterity—just serving in the long run to compensate the deficient endowments of other individuals, whose special powers lie in other directions; and so to keep up the normal structure of the species. The working out of the process is here somewhat difficult to follow; but it appears to me that as fast as the number of bodily and mental faculties increases, and as fast as the maintenance of life comes to depend less on the amount of any one, and more on the combined action of all; so fast does the production of specialities of character by natural selection alone, become difficult. Particularly does this seem to be so with a species so multitudinous in its powers as mankind; and above all does it seem to be so with such of the human powers as have but minor shares in aiding the struggle for life—the aesthetic faculties, for example."
Dwelling for a moment on this last illustration of the class of difficulties described, let us ask how we are to interpret the development of the musical faculty. I will not enlarge on the family antecedents of the great composers. I will merely suggest the inquiry whether the greater powers possessed by Beethoven and Mozart, by Weber and Rossini, than by their fathers, were not due in larger measure to the inherited effects of daily exercise of the musical faculty by their fathers, than to inheritance, with increase, of spontaneous variations; and whether the diffused musical powers of the Bach clan, culminating in those of Johann Sebastian, did not result in part from constant practice; but I will raise the more general question—How came there that endowment of musical faculty which characterizes modern Europeans at large, as compared with their remote ancestors. The monotonous chants of low savages cannot be said to show any melodic inspiration; and it is not evident that an individual savage who had a little more musical perception than the rest, would derive any such advantage in the maintenance of life as would secure the spread of his superiority by inheritance of the variation. And then what are we to say of harmony? We cannot suppose that the appreciation of this, which is relatively modern, can have arisen by descent from the men in whom successive variations increased the appreciation of it—the composers and musical performers; for on the whole, these have been men whose worldly prosperity was not such as enabled them to rear many children inheriting their special traits. Even if we count the illegitimate ones, the survivors of these added to the survivors of the legitimate ones, can hardly be held to have yielded more than average numbers of descendants; and those who inherited their special traits have not often been thereby so aided in the struggle for existence as to further the spread of such traits. Rather the tendency seems to have been the reverse.
Since the above passage was written, I have found in the second volume of Animals and Plants under Domestication, a remark made by Mr. Darwin, practically implying that among creatures which depend for their lives on the efficiency of numerous powers, the increase of any one by the natural selection of a variation is necessarily difficult. Here it is.
"Finally, as indefinite and almost illimitable variability is the usual result of domestication and cultivation, with the same part or organ varying in different individuals in different or even in directly opposite ways; and as the same variation, if strongly pronounced, usually recurs only after long intervals of time, any particular variation would generally be lost by crossing, reversion, and the accidental destruction of the varying individuals, unless carefully preserved by man."—Vol. ii, 292.
Remembering that mankind, subject as they are to this domestication and cultivation, are not, like domesticated animals, under an agency which picks out and preserves particular variations; it results that there must usually be among them, under the influence of natural selection alone, a continual disappearance of any useful variations of particular faculties which may arise. Only in cases of variations which are specially preservative, as for example, great cunning during a relatively barbarous state, can we expect increase from natural selection alone. We cannot suppose that minor traits, exemplified among others by the aesthetic perceptions, can have been evolved by natural selection. But if there is inheritance of functionally-produced modifications of structure, evolution of such minor traits is no longer inexplicable.
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Two remarks made by Mr. Darwin have implications from which the same general conclusion must, I think, be drawn. Speaking of the variability of animals and plants under domestication, he says:—
"Changes of any kind in the conditions of life, even extremely slight changes, often suffice to cause variability.... Animals and plants continue to be variable for an immense period after their first domestication; ... In the course of time they can be habituated to certain changes, so as to become less variable; ... There is good evidence that the power of changed conditions accumulates; so that two, three, or more generations must be exposed to new conditions before any effect is visible.... Some variations are induced by the direct action of the surrounding conditions on the whole organization, or on certain parts alone, and other variations are induced indirectly through the reproductive system being affected in the same manner as is so common with organic beings when removed from their natural conditions."—(Animals and Plants under Domestication, vol. ii, 270.)
There are to be recognized two modes of this effect produced by changed conditions on the reproductive system, and consequently on offspring. Simple arrest of development is one. But beyond the variations of offspring arising from imperfectly developed reproductive systems in parents—variations which must be ordinarily in the nature of imperfections—there are others due to a changed balance of functions caused by changed conditions. The fact noted by Mr. Darwin in the above passage, "that the power of changed conditions accumulates; so that two, three, or more generations must be exposed to new conditions before any effect is visible," implies that during these generations there is going on some change of constitution consequent on the changed proportions and relations of the functions. I will not dwell on the implication, which seems tolerably clear, that this change must consist of such modifications of organs as adapt them to their changed functions; and that if the influence of changed conditions "accumulates," it must be through the inheritance of such modifications. Nor will I press the question—What is the nature of the effect registered in the reproductive elements, and which is subsequently manifested by variations?—Is it an effect entirely irrelevant to the new requirements of the variety?—Or is it an effect which makes the variety less fit for the new requirements?—Or is it an effect which makes it more fit for the new requirements? But not pressing these questions, it suffices to point out the necessary implication that changed functions of organs do, in some way or other, register themselves in changed proclivities of the reproductive elements. In face of these facts it cannot be denied that the modified action of a part produces an inheritable effect—be the nature of that effect what it may.
The second of the remarks above adverted to as made by Mr. Darwin, is contained in his sections dealing with correlated variations. In the Origin of Species, p. 114, he says—
"The whole organization is so tied together during its growth and development, that when slight variations in any one part occur, and are accumulated through natural selection, other parts become modified."
And a parallel statement contained in Animals and Plants under Domestication, vol. ii, p. 320, runs thus—
"Correlated variation is an important subject for us; for when one part is modified through continued selection, either by man or under nature, other parts of the organization will be unavoidably modified. From this correlation it apparently follows that, with our domesticated animals and plants, varieties rarely or never differ from each other by some single character alone."
By what process does a changed part modify other parts? By modifying their functions in some way or degree, seems the necessary answer. It is indeed, imaginable, that where the part changed is some dermal appendage which, becoming larger, has abstracted more of the needful material from the general stock, the effect may consist simply in diminishing the amount of this material available for other dermal appendages, leading to diminution of some or all of them, and may fail to affect in appreciable ways the rest of the organism: save perhaps the blood-vessels near the enlarged appendage. But where the part is an active one—a limb, or viscus, or any organ which constantly demands blood, produces waste matter, secretes, or absorbs—then all the other active organs become implicated in the change. The functions performed by them have to constitute a moving equilibrium; and the function of one cannot, by alteration of the structure performing it, be modified in degree or kind, without modifying the functions of the rest—some appreciably and others inappreciably, according to the directness or indirectness of their relations. Of such inter-dependent changes, the normal ones are naturally inconspicuous; but those which are partially or completely abnormal, sufficiently carry home the general truth. Thus, unusual cerebral excitement affects the excretion through the kidneys in quantity or quality or both. Strong emotions of disagreeable kinds check or arrest the flow of bile. A considerable obstacle to the circulation offered by some important structure in a diseased or disordered state, throwing more strain upon the heart, causes hypertrophy of its muscular walls; and this change which is, so far as concerns the primary evil, a remedial one, often entails mischiefs in other organs. "Apoplexy and palsy, in a scarcely credible number of cases, are directly dependent on hypertrophic enlargement of the heart." And in other cases, asthma, dropsy, and epilepsy are caused. Now if a result of this inter-dependence as seen in the individual organism, is that a local modification of one part produces, by changing their functions, correlative modifications of other parts, then the question here to be put is—Are these correlative modifications, when of a kind falling within normal limits, inheritable or not. If they are inheritable, then the fact stated by Mr. Darwin that "when one part is modified through continued selection," "other parts of the organization will be unavoidably modified" is perfectly intelligible: these entailed secondary modifications are transmitted pari passu with the successive modifications produced by selection. But what if they are not inheritable? Then these secondary modifications caused in the individual, not being transmitted to descendants, the descendants must commence life with organizations out of balance, and with each increment of change in the part affected by selection, their organizations must get more out of balance—must have a larger and larger amounts of re-organization to be made during their lives. Hence the constitution of the variety must become more and more unworkable.
The only imaginable alternative is that the re-adjustments are effected in course of time by natural selection. But, in the first place, as we find no proof of concomitant variation among directly co-operative parts which are closely united, there cannot be assumed any concomitant variation among parts which are both indirectly co-operative and far from one another. And, in the second place, before all the many required re-adjustments could be made, the variety would die out from defective constitution. Even were there no such difficulty, we should still have to entertain a strange group of propositions, which would stand as follows:—1. Change in one part entails, by reaction on the organism, changes, in other parts, the functions of which are necessarily changed. 2. Such changes worked in the individual, affect, in some way, the reproductive elements: these being found to evolve unusual structures when the constitutional balance has been continuously disturbed. 3. But the changes in the reproductive elements thus caused, are not such as represent these functionally-produced changes: the modifications conveyed to offspring are irrelevant to these various modifications functionally produced in the organs of the parents. 4. Nevertheless, while the balance of functions cannot be re-established through inheritance of the effects of disturbed functions on structures, wrought throughout the individual organism; it can be re-established by the inheritance of fortuitous variations which occur in all the affected organs without reference to these changes of function.
Now without saying that acceptance of this group of propositions is impossible, we may certainly say that it is not easy.
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"But where are the direct proofs that inheritance of functionally-produced modifications takes place?" is a question which will be put by those who have committed themselves to the current exclusive interpretation. "Grant that there are difficulties; still, before the transmitted effects of use and disuse can be legitimately assigned in explanation of them, we must have good evidence that the effects of use and disuse are transmitted."
Before dealing directly with this demurrer, let me deal with it indirectly, by pointing out that the lack of recognized evidence may be accounted for without assuming that there is not plenty of it. Inattention and reluctant attention lead to the ignoring of facts which really exist in abundance; as is well illustrated in the case of pre-historic implements. Biassed by the current belief that no traces of man were to be found on the Earth's surface, save in certain superficial formations of very recent date, geologists and anthropologists not only neglected to seek such traces, but for a long time continued to pooh-pooh those who said they had found them. When M. Boucher de Perthes at length succeeded in drawing the eyes of scientific men to the flint implements discovered by him in the quarternary deposits of the Somme valley; and when geologists and anthropologists had thus been convinced that evidences of human existence were to be found in formations of considerable age, and thereafter began to search for them; they found plenty of them all over the world. Or again, to take an instance closely germane to the matter, we may recall the fact that the contemptuous attitude towards the hypothesis of organic evolution which naturalists in general maintained before the publication of Mr. Darwin's work, prevented them from seeing the multitudinous facts by which it is supported. Similarly, it is very possible that their alienation from the belief that there is a transmission of those changes of structure which are produced by changes of action, makes naturalists slight the evidence which supports that belief and refuse to occupy themselves in seeking further evidence.
If it be asked how it happens that there have been recorded multitudinous instances of variations fortuitously arising and re-appearing in offspring, while there have not been recorded instances of the transmission of changes functionally produced, there are three replies. The first is that changes of the one class are many of them conspicuous, while those of the other class are nearly all inconspicuous. If a child is born with six fingers, the anomaly is not simply obvious but so startling as to attract much notice; and if this child, growing up, has six-fingered descendents, everybody in the locality hears of it. A pigeon with specially-coloured feathers, or one distinguished by a broadened and upraised tail, or by a protuberance of the neck, draws attention by its oddness; and if in its young the trait is repeated, occasionally with increase, the fact is remarked, and there follows the thought of establishing the peculiarity by selection. A lamb disabled from leaping by the shortness of its legs, could not fail to be observed; and the fact that its offspring were similarly short-legged, and had a consequent inability to get over fences, would inevitably become widely known. Similarly with plants. That this flower had an extra number of petals, that that was unusually symmetrical, and that another differed considerably in colour from the average of its kind, would be easily seen by an observant gardener; and the suspicion that such anomalies are inheritable having arisen, experiments leading to further proofs that they are so, would frequently be made. But it is not thus with functionally-produced modifications. The seats of these are in nearly all cases the muscular, osseous, and nervous systems, and the viscera—parts which are either entirely hidden or greatly obscured. Modification in a nervous centre is inaccessible to vision; bones may be considerably altered in size or shape without attention being drawn to them; and, covered with thick coats as are most of the animals open to continuous observation, the increases or decreases in muscles must be great before they become externally perceptible.
A further important difference between the two inquiries is that to ascertain whether a fortuitous variation is inheritable, needs merely a little attention to the selection of individuals and the observation of offspring; while to ascertain whether there is inheritance of a functionally-produced modification, it is requisite to make arrangements which demand the greater or smaller exercise of some part or parts; and it is difficult in many cases to find such arrangements, troublesome to maintain them even for one generation, and still more through successive generations.
Nor is this all. There exist stimuli to inquiry in the one case which do not exist in the other. The money-interest and the interest of the fancier, acting now separately and now together, have prompted multitudinous individuals to make experiments which have brought out clear evidence that fortuitous variations are inherited. The cattle-breeders who profit by producing certain shapes and qualities; the keepers of pet animals who take pride in the perfections of those they have bred; the florists, professional and amateur, who obtain new varieties and take prizes; form a body of men who furnish naturalists with countless of the required proofs. But there is no such body of men, led either by pecuniary interest or the interest of a hobby, to ascertain by experiments whether the effects of use and disuse are inheritable.
Thus, then, there are amply sufficient reasons why there is a great deal of direct evidence in the one case and but little in the other: such little being that which comes out incidentally. Let us look at what there is of it.
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Considerable weight attaches to a fact which Brown-Sequard discovered, quite by accident, in the course of his researches. He found that certain artificially-produced lesions of the nervous system, so small even as a section of the sciatic nerve, left, after healing, an increasing excitability which ended in liability to epilepsy; and there afterwards came out the unlooked-for result that the offspring of guinea-pigs which had thus acquired an epileptic habit such that a pinch on the neck would produce a fit, inherited an epileptic habit of like kind. It has, indeed, been since alleged that guinea pigs tend to epilepsy, and that phenomena of the kind described, occur where there have been no antecedents like those in Brown-Sequard's case. But considering the improbability that the phenomena observed by him happened to be nothing more than phenomena which occasionally arise naturally, we may, until there is good proof to the contrary, assign some value to his results.
Evidence not of this directly experimental kind, but nevertheless of considerable weight, is furnished by other nervous disorders. There is proof enough that insanity admits of being induced by circumstances which, in one or other way, derange the nervous functions—excesses of this or that kind; and no one questions the accepted belief that insanity is inheritable. Is it alleged that the insanity which is inheritable is that which spontaneously arises, and that the insanity which follows some chronic perversion of functions is not inheritable? This does not seem a very reasonable allegation; and until some warrant for it is forthcoming, we may fairly assume that there is here a further support for belief in the transmission of functionally-produced changes.
Moreover, I find among physicians the belief that nervous disorders of a less severe kind are inheritable. Men who have prostrated their nervous systems by prolonged overwork or in some other way, have children more or less prone to nervousness. It matters not what may be the form of inheritance—whether it be of a brain in some way imperfect, or of a deficient blood-supply; it is in any case the inheritance of functionally-modified structures.
Verification of the reasons above given for the paucity of this direct evidence, is yielded by contemplation of it; for it is observable that the cases named are cases which, from one or other cause, have thrust themselves on observation. They justify the suspicion that it is not because such cases are rare that many of them cannot be cited; but simply because they are mostly unobtrusive, and to be found only by that deliberate search which nobody makes. I say nobody, but I am wrong. Successful search has been made by one whose competence as an observer is beyond question, and whose testimony is less liable than that of all others to any bias towards the conclusion that such inheritance takes place. I refer to the author of the Origin of Species.
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Now-a-days most naturalists are more Darwinian than Mr. Darwin himself. I do not mean that their beliefs in organic evolution are more decided; though I shall be supposed to mean this by the mass of readers, who identify Mr. Darwin's great contribution to the theory of organic evolution, with the theory of organic evolution itself, and even with the theory of evolution at large. But I mean that the particular factor which he first recognized as having played so immense a part in organic evolution, has come to be regarded by his followers as the sole factor, though it was not so regarded by him. It is true that he apparently rejected altogether the causal agencies alleged by earlier inquirers. In the Historical Sketch prefixed to the later editions of his Origin of Species (p. xiv, note), he writes:—"It is curious how largely my grandfather, Dr. Erasmus Darwin, anticipated the views and erroneous grounds of opinion of Lamarck in his 'Zoonomia' (vol. i, pp. 500-510), published in 1794." And since, among the views thus referred to, was the view that changes of structure in organisms arise by the inheritance of functionally-produced changes, Mr. Darwin seems, by the above sentence, to have implied his disbelief in such inheritance. But he did not mean to imply this; for his belief in it as a cause of evolution, if not an important cause, is proved by many passages in his works. In the first chapter of the Origin of Species (p. 8 of the sixth edition), he says respecting the inherited effects of habit, that "with animals the increased use or disuse of parts has had a more marked influence;" and he gives as instances the changed relative weights of the wing bones and leg bones of the wild duck and the domestic duck, "the great and inherited development of the udders in cows and goats," and the drooping ears of various domestic animals. Here are other passages taken from the latest edition of the work.
"I think there can be no doubt that use in our domestic animals has strengthened and enlarged certain parts, and disuse diminished them; and that such modifications are inherited" (p. 108). [And on the following pages he gives five further examples of such effects.] "Habit in producing constitutional peculiarities and use in strengthening and disuse in weakening and diminishing organs, appear in many cases to have been potent in their effects" (p. 131). "When discussing special cases, Mr. Mivart passes over the effects of the increased use and disuse of parts, which I have always maintained to be highly important, and have treated in my 'Variation under Domestication' at greater length than, as I believe, any other writer" (p. 176). "Disuse, on the other hand, will account for the less developed condition of the whole inferior half of the body, including the lateral fins" (p. 188). "I may give another instance of a structure which apparently owes its origin exclusively to use or habit" (p. 188). "It appears probable that disuse has been the main agent in rendering organs rudimentary" (pp. 400-401). "On the whole, we may conclude that habit, or use and disuse, have, in some cases, played a considerable part in the modification of the constitution and structure; but that the effects have often been largely combined with, and sometimes overmastered by, the natural selection of innate variations" (p. 114).
In his subsequent work, The Variation of Animals and Plants under Domestication, where he goes into full detail, Mr. Darwin gives more numerous illustrations of the inherited effects of use and disuse. The following are some of the cases, quoted from volume i of the first edition.
Treating of domesticated rabbits, he says:—"the want of exercise has apparently modified the proportional length of the limbs in comparison with the body" (p. 116). "We thus see that the most important and complicated organ [the brain] in the whole organization is subject to the law of decrease in size from disuse" (p. 129). He remarks that in birds of the oceanic islands "not persecuted by any enemies, the reduction of their wings has probably been caused by gradual disuse." After comparing one of these, the water-hen of Tristan d'Acunha, with the European water-hen, and showing that all the bones concerned in flight are smaller, he adds—"Hence in the skeleton of this natural species nearly the same changes have occurred, only carried a little further, as with our domestic ducks, and in this latter case I presume no one will dispute that they have resulted from the lessened use of the wings and the increased use of the legs" (pp. 286-7). "As with other long-domesticated animals, the instincts of the silk-moth have suffered. The caterpillars, when placed on a mulberry-tree, often commit the strange mistake of devouring the base of the leaf on which they are feeding, and consequently fall down; but they are capable, according to M. Robinet, of again crawling up the trunk. Even this capacity sometimes fails, for M. Martins placed some caterpillars on a tree, and those which fell were not able to remount and perished of hunger; they were even incapable of passing from leaf to leaf" (p. 304).
Here are some instances of like meaning from volume ii.
"In many cases there is reason to believe that the lessened use of various organs has affected the corresponding parts in the offspring. But there is no good evidence that this ever follows in the course of a single generation.... Our domestic fowls, ducks, and geese have almost lost, not only in the individual but in the race, their power of flight; for we do not see a chicken, when frightened, take flight like a young pheasant.... With domestic pigeons, the length of the sternum, the prominence of its crest, the length of the scapulae and furcula, the length of the wings as measured from tip to tip of the radius, are all reduced relatively to the same parts in the wild pigeon." [After detailing kindred diminutions in fowls and ducks, Mr. Darwin adds] "The decreased weight and size of the bones, in the foregoing cases, is probably the indirect result of the reaction of the weakened muscles on the bones" (pp. 297-8). "Nathusius has shown that, with the improved races of the pig, the shortened legs and snout, the form of the articular condyles of the occiput, and the position of the jaws with the upper canine teeth projecting in a most anomalous manner in front of the lower canines, may be attributed to these parts not having been fully exercised.... These modifications of structure, which are all strictly inherited, characterise several improved breeds, so that they cannot have been derived from any single domestic or wild stock. With respect to cattle, Professor Tanner has remarked that the lungs and liver in the improved breeds 'are found to be considerably reduced in size when compared with those possessed by animals having perfect liberty;' ... The cause of the reduced lungs in highly-bred animals which take little exercise is obvious" (pp. 299-300). [And on pp. 301, 302 and 303, he gives facts showing the effects of use and disuse in changing, among domestic animals, the characters of the ears, the lengths of the intestines, and, in various ways, the natures of the instincts.] |
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