Chapter II: Preface (2)

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 descendants, 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.

* * * * *

Considerable weight attaches to a fact which Brown-Séquard 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-Séquard'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_.

* * * * *

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. 11 of the first edition), he says respecting the inherited effects of habit, that “with animals the increased use or disuse of parts has had a 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 scapulæ 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.]

But Mr. Darwin's admission, or rather his assertion, that the inheritance of functionally-produced modifications has been a factor in organic evolution, is made clear not by these passages alone and by kindred ones. It is made clearer still by a passage in the preface to the second edition of his _Descent of Man_. He there protests against that current version of his views in which this factor makes no appearance. The passage is as follows.

“I may take this opportunity of remarking that my critics frequently
assume that I attribute all changes of corporeal structure and
mental power exclusively to the natural selection of such variations
as are often called spontaneous; whereas, even in the first edition
of the 'Origin of Species,' I distinctly stated that great weight
must be attributed to the inherited effects of use and disuse, with
respect both to the body and mind.”

Nor is this all. There is evidence that Mr. Darwin's belief in the efficiency of this factor, became stronger as he grew older and accumulated more evidence. The first of the extracts above given, taken from the sixth edition of the _Origin of Species_, runs thus:—

“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.”

Now on turning to the first edition, p. 134, it will be found that instead of the words—“I think there can be no doubt,” the words originally used were—“I think there can be _little_ doubt.” That this deliberate erasure of a qualifying word and substitution of a word implying unqualified belief, was due to a more decided recognition of a factor originally under-estimated, is clearly implied by the wording of the above-quoted passage from the preface to the _Descent of Man_; where he says that “_even_ in the first edition of the 'Origin of Species,'” &c.: the implication being that much more in subsequent editions, and subsequent works, had he insisted on this factor. The change thus indicated is especially significant as having occurred at a time of life when the natural tendency is towards fixity of opinion.

During that earlier period when he was discovering the multitudinous cases in which his own hypothesis afforded solutions, and simultaneously observing how utterly futile in these multitudinous cases was the hypothesis propounded by his grandfather and Lamarck, Mr. Darwin was, not unnaturally, almost betrayed into the belief that the one is all-sufficient and the other inoperative. But in the mind of one so candid and ever open to more evidence, there naturally came a reaction. The inheritance of functionally-produced modifications, which, judging by the passage quoted above concerning the views of these earlier enquirers, would seem to have been at one time denied, but which as we have seen was always to some extent recognized, came to be recognized more and more, and deliberately included as a factor of importance.

* * * * *

Of this reaction displayed in the later writings of Mr. Darwin, let us now ask—Has it not to be carried further? Was the share in organic evolution which Mr. Darwin latterly assigned to the transmission of modifications caused by use and disuse, its due share? Consideration of the groups of evidences given above, will, I think, lead us to believe that its share has been much larger than he supposed even in his later days.

There is first the implication yielded by extensive classes of phenomena which remain inexplicable in the absence of this factor. If, as we see, co-operative parts do not vary together, even when few and close together, and may not therefore be assumed to do so when many and remote, we cannot account for those innumerable changes in organization which are implied when, for advantageous use of some modified part, many other parts which join it in action have to be modified.

Further, as increasing complexity of structure, accompanying increasing complexity of life, implies increasing number of faculties, of which each one conduces to preservation of self or descendants; and as the various individuals of a species, severally requiring something like the normal amounts of all these, may individually profit, here by an unusual amount of one, and there by an unusual amount of another; it follows that as the number of faculties becomes greater, it becomes more difficult for any one to be further developed by natural selection. Only where increase of some one is _predominantly_ advantageous does the means seem adequate to the end. Especially in the case of powers which do not subserve self-preservation in appreciable degrees, does development by natural selection appear impracticable.

It is a fact recognized by Mr. Darwin, that where, by selection through successive generations, a part has been increased or decreased, its reaction upon other parts entails changes in them. This reaction is effected through the changes of function involved. If the changes of structure produced by such changes of function, are inheritable, then the re-adjustment of parts throughout the organism, taking place generation after generation, maintains an approximate balance; but if not, then generation after generation the organism must get more and more out of gear, and tend to become unworkable.

Further, as it is proved that change in the balance of functions registers its effects on the reproductive elements, we have to choose between the alternatives that the registered effects are irrelevant to the particular modifications which the organism has undergone, or that they are such as tend to produce repetitions of these modifications. The last of these alternatives makes the facts comprehensible; but the first of them not only leaves us with several unsolved problems, but is incongruous with the general truth that by reproduction, ancestral traits, down to minute details, are transmitted.

Though, in the absence of pecuniary interests and the interests in hobbies, no such special experiments as those which have established the inheritance of fortuitous variations have been made to ascertain whether functionally-produced modifications are inherited; yet certain apparent instances of such inheritance have forced themselves on observation without being sought for. In addition to other indications of a less conspicuous kind, is the one I have given above—the fact that the apparatus for tearing and mastication has decreased with decrease of its function, alike in civilized man and in some varieties of dogs which lead protected and pampered lives. Of the numerous cases named by Mr. Darwin, it is observable that they are yielded not by one class of parts only, but by most if not all classes—by the dermal system, the muscular system, the osseous system, the nervous system, the viscera; and that among parts liable to be functionally modified, the most numerous observed cases of inheritance are furnished by those which admit of preservation and easy comparison—the bones: these cases, moreover, being specially significant as showing how, in sundry unallied species, parallel changes of structure have occurred along with parallel changes of habit.

What, then, shall we say of the general implication? Are we to stop short with the admission that inheritance of functionally-produced modifications takes place only in cases in which there is evidence of it? May we properly assume that these many instances of changes of structure caused by changes of function, occurring in various tissues and various organs, are merely special and exceptional instances having no general significance? Shall we suppose that though the evidence which already exists has come to light without aid from a body of inquirers, there would be no great increase were due attention devoted to the collection of evidence? This is, I think, not a reasonable supposition. To me the _ensemble_ of the facts suggests the belief, scarcely to be resisted, that the inheritance of functionally-produced modifications takes place universally. Looking at physiological phenomena as conforming to physical principles, it is difficult to conceive that a changed play of organic forces which in many cases of different kinds produces an inherited change of structure, does not do this in all cases. The implication, very strong I think, is that the action of every organ produces on it a reaction which, usually not altering its rate of nutrition, sometimes leaves it with diminished nutrition consequent on diminished action, and at other times increases its nutrition in proportion to its increased action; that while generating a modified _consensus_ of functions and of structures, the activities are at the same time impressing this modified _consensus_ on the sperm-cells and germ-cells whence future individuals are to be produced; and that in ways mostly too small to be identified, but occasionally in more conspicuous ways and in the course of generations, the resulting modifications of one or other kind show themselves. Further, it seems to me that as there are certain extensive classes of phenomena which are inexplicable if we assume the inheritance of fortuitous variations to be the sole factor, but which become at once explicable if we admit the inheritance of functionally-produced changes, we are justified in concluding that this inheritance of functionally-produced changes has been not simply a co-operating factor in organic evolution, but has been a co-operating factor without which organic evolution, in its higher forms at any rate, could never have taken place.

Be this or be it not a warrantable conclusion, there is, I think, good reason for a provisional acceptance of the hypothesis that the effects of use and disuse are inheritable; and for a methodic pursuit of inquiries with the view of either establishing it or disproving it. It seems scarcely reasonable to accept without clear demonstration, the belief that while a trivial difference of structure arising spontaneously is transmissible, a massive difference of structure, maintained generation after generation by change of function, leaves no trace in posterity. Considering that unquestionably the modification of structure by function is a _vera causa_, in so far as concerns the individual; and considering the number of facts which so competent an observer as Mr. Darwin regarded as evidence that transmission of such modifications takes place in particular cases; the hypothesis that such transmission takes place in conformity with a general law, holding of all active structures, should, I think, be regarded as at least a good working hypothesis.

* * * * *

But now supposing the broad conclusion above drawn to be granted—supposing all to agree that from the beginning, along with inheritance of useful variations fortuitously arising, there has been inheritance of effects produced by use and disuse; do there remain no classes of organic phenomena unaccounted for? To this question I think it must be replied that there do remain classes of organic phenomena unaccounted for. It may, I believe, be shown that certain cardinal traits of animals and plants at large are still unexplained; and that a further factor must be recognized. To show this, however, will require another paper.

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Footnote 1:

It is probable that this shortening has resulted not directly but
indirectly, from the selection of individuals which were noted for
tenacity of hold; for the bull-dog's peculiarity in this respect seems
due to relative shortness of the upper jaw, giving the underhung
structure which, involving retreat of the nostrils, enables the dog to
continue breathing while holding.

II.

Ask a plumber who is repairing your pump, how the water is raised in it, and he replies—“By suction.” Recalling the ability which he has to suck up water into his mouth through a tube, he is certain that he understands the pump's action. To inquire what he means by suction, seems to him absurd. He says you know as well as he does, what he means; and he cannot see that there is any need for asking how it happens that the water rises in the tube when he strains his mouth in a particular way. To the question why the pump, acting by suction, will not make the water rise above 32 feet, and practically not so much, he can give no answer; but this does not shake his confidence in his explanation.

On the other hand an inquirer who insists on knowing what suction is, may obtain from the physicist answers which give him clear ideas, not only about it but about many other things. He learns that on ourselves and all things around, there is an atmospheric pressure amounting to about 15 pounds on the square inch: 15 pounds being the average weight of a column of air having a square inch for its base and extending upwards from the sea-level to the limit of the Earth's atmosphere. He is made to observe that when he puts one end of a tube into water and the other end into his mouth, and then draws back his tongue, so leaving a vacant space, two things happen. One is that the pressure of air outside his cheeks, no longer balanced by an equal pressure of air inside, thrusts his cheeks inwards; and the other is that the pressure of air on the surface of the water, no longer balanced by an equal pressure of air within the tube and his mouth (into which part of the air from the tube has gone) the water is forced up the tube in consequence of the unequal pressure. Once understanding thus the nature of the so-called suction, he sees how it happens that when the plunger of the pump is raised and relieves from atmospheric pressure the water below it, the atmospheric pressure on the water in the well, not being balanced by that on the water in the tube, forces the water higher up the tube, so that it follows the plunger. And now he sees why the water cannot be raised beyond the theoretic limit of 32 feet: a limit made much lower in practice by imperfections in the apparatus. For if, simplifying the conception, he supposes the tube of the pump to be a square inch in section, then the atmospheric pressure of 15 pounds per square inch on the water in the well, can raise the water in the tube to such height only that the entire column of it weighs 15 pounds. Having been thus enlightened about the pump's action, the action of a barometer becomes intelligible. He perceives how, under the conditions established, the weight of the column of mercury balances that of an atmospheric column of equal diameter; and how, as the weight of the atmospheric column varies, there is a corresponding variation in the weight of the mercurial column,—shown by change of height. Moreover, having previously supposed that he understood the ascent of a balloon when he ascribed it to relative lightness, he now sees that he did not truly understand it. For he did not recognize it as a result of that upward pressure caused by the difference between the weight of the mass formed by the gas in the balloon _plus_ the cylindrical column of air extending above it to the limit of the atmosphere, and the weight of a similar cylindrical column of air extending down to the under surface of the balloon: this difference of weight causing an equivalent upward pressure on the under surface.

Why do I introduce these familiar truths so entirely irrelevant to my subject? I do it to show, in the first place, the contrast between a vague conception of a cause and a distinct conception of it; or rather, the contrast between that conception of a cause which results when it is simply classed with some other or others which familiarity makes us think we understand, and that conception of a cause which results when it is represented in terms of definite physical forces admitting of measurement. And I do it to show, in the second place, that when we insist on resolving a verbally-intelligible cause into its actual factors, we get not only a clear solution of the problem before us, but we find that the way is opened to solutions of sundry other problems. While we rest satisfied with unanalyzed causes, we may be sure both that we do not rightly comprehend the production of the particular effects ascribed to them, and that we overlook other effects which would be revealed to us by contemplation of the causes as analyzed. Especially must this be so where the causation is complex. Hence we may infer that the phenomena presented by the development of species, are not likely to be truly conceived unless we keep in view the concrete agencies at work. Let us look closely at the facts to be dealt with.

* * * * *

The growth of a thing is effected by the joint operation of certain forces on certain materials; and when it dwindles, there is either a lack of some materials, or the forces co-operate in a way different from that which produces growth. If a structure has varied, the implication is that the processes which built it up were made unlike the parallel processes in other cases, by the greater or less amount of some one or more of the matters or actions concerned. Where there is unusual fertility, the play of vital activities is thereby shown to have deviated from the ordinary play of vital activities; and conversely, if there is infertility. If the germs, or ova, or seed, or offspring partially developed, survive more or survive less, it is either because their molar or molecular structures are unlike the average ones, or because they are affected in unlike ways by surrounding agencies. When life is prolonged, the fact implies that the combination of actions, visible and invisible, constituting life, retains its equilibrium longer than usual in presence of environing forces which tend to destroy its equilibrium. That is to say, growth, variation, survival, death, if they are to be reduced to the forms in which physical science can recognize them, must be expressed as effects of agencies definitely conceived—mechanical forces, light, heat, chemical affinity, &c.

This general conclusion brings with it the thought that the phrases employed in discussing organic evolution, though convenient and indeed needful, are liable to mislead us by veiling the actual agencies. That which really goes on in every organism is the working together of component parts in ways conducing to the continuance of their combined actions, in presence of things and actions outside; some of which tend to subserve, and others to destroy, the combination. The matters and forces in these two groups, are the sole causes properly so called. The words “natural selection,” do not express a cause in the physical sense. They express a mode of co-operation among causes—or rather, to speak strictly, they express an effect of this mode of co-operation. The idea they convey seems perfectly intelligible. Natural selection having been compared with artificial selection, and the analogy pointed out, there apparently remains no indefiniteness: the inconvenience being, however, that the definiteness is of a wrong kind. The tacitly implied Nature which selects, is not an embodied agency analogous to the man who selects artificially; and the selection is not the picking out of an individual fixed on, but the overthrowing of many individuals by agencies which one successfully resists, and hence continues to live and multiply. Mr. Darwin was conscious of these misleading implications. In the introduction to his _Animals and Plants under Domestication_ (p. 6) he says:—

“For brevity sake I sometimes speak of natural selection as an
intelligent power; ... I have, also, often personified the word
Nature; for I have found it difficult to avoid this ambiguity; but I
mean by nature only the aggregate action and product of many natural
laws,—and by laws only the ascertained sequence of events.”

But while he thus clearly saw, and distinctly asserted, that the factors of organic evolution are the concrete actions, inner and outer, to which every organism is subject, Mr. Darwin, by habitually using the convenient figure of speech, was, I think, prevented from recognizing so fully as he would otherwise have done, certain fundamental consequences of these actions.

Though it does not personalize the cause, and does not assimilate its mode of working to a human mode of working, kindred objections may be urged against the expression to which I was led when seeking to present the phenomena in literal terms rather than metaphorical terms—the survival of the fittest;[2] for in a vague way the first word, and in a clear way the second word, calls up an anthropocentric idea. The thought of survival inevitably suggests the human view of certain sets of phenomena, rather than that character which they have simply as groups of changes. If, asking what we really know of a plant, we exclude all the ideas associated with the words life and death, we find that the sole facts known to us are that there go on in the plant certain inter-dependent processes, in presence of certain aiding and hindering influences outside of it; and that in some cases a difference of structure or a favourable set of circumstances, allows these inter-dependent processes to go on for longer periods than in other cases. Again, in the working together of those many actions, internal and external, which determine the lives or deaths of organisms, we see nothing to which the words fitness and unfitness are applicable in the physical sense. If a key fits a lock, or a glove a hand, the relation of the things to one another is presentable to the perceptions. No approach to fitness of this kind is made by an organism which continues to live under certain conditions. Neither the organic structures themselves, nor their individual movements, nor those combined movements of certain among them which constitute conduct, are related in any analogous way to the things and actions in the environment. Evidently the word fittest, as thus used, is a figure of speech; suggesting the fact that amid surrounding actions, an organism characterized by the word has either a greater ability than others of its kind to maintain the equilibrium of its vital activities, or else has so much greater a power of multiplication that though not longer lived than they, it continues to live in posterity more persistently. And indeed, as we here see, the word fittest has to cover cases in which there may be less ability than usual to survive individually, but in which the defect is more than made good by higher degrees of fertility.

I have elaborated this criticism with the intention of emphasizing the need for studying the changes which have gone on, and are ever going on, in organic bodies, from an exclusively physical point of view. On contemplating the facts from this point of view, we become aware that, besides those special effects of the co-operating forces which eventuate in the longer survival of one individual than of others, and in the consequent increase through generations, of some trait which furthered its survival, many other effects are being wrought on each and all of the individuals. Bodies of every class and quality, inorganic as well as organic, are from instant to instant subject to the influences in their environments; are from instant to instant being changed by these in ways that are mostly inconspicuous; and are in course of time changed by them in conspicuous ways. Living things in common with dead things, are, I say, being thus perpetually acted upon and modified; and the changes hence resulting, constitute an all-important part of those undergone in the course of organic evolution. I do not mean to imply that changes of this class pass entirely unrecognized; for, as we shall see, Mr. Darwin takes cognizance of certain secondary and special ones. But the effects which are not taken into account, are those primary and universal effects which give certain fundamental characters to all organisms. Contemplation of an analogy will best prepare the way for appreciation of them, and of the relation they bear to those which at present monopolize attention.

An observant rambler along shores, will, here and there, note places where the sea has deposited things more or less similar, and separated them from dissimilar things—will see shingle parted from sand; larger stones sorted from smaller stones; and will occasionally discover deposits of shells more or less worn by being rolled about. Sometimes the pebbles or boulders composing the shingle at one end of a bay, he will find much larger than those at the other: intermediate sizes, having small average differences, occupying the space between the extremes. An example occurs, if I remember rightly, some mile or two to the west of Tenby; but the most remarkable and well-known example is that afforded by the Chesil bank. Here, along a shore some sixteen miles long, there is a gradual increase in the sizes of the stones; which, being at one end but mere pebbles, are at the other end immense boulders. In this case, then, the breakers and the undertow have effected a selection—have at each place left behind those stones which were too large to be readily moved, while taking away others small enough to be moved easily. But now, if we contemplate exclusively this selective action of the sea, we overlook certain important effects which the sea simultaneously works. While the stones have been differently acted upon in so far that some have been left here and some carried there; they have been similarly acted upon in two allied, but distinguishable, ways. By perpetually rolling them about and knocking them one against another, the waves have so broken off their most prominent parts as to produce in all of them more or less rounded forms; and then, further, the mutual friction of the stones simultaneously caused, has smoothed their surfaces. That is to say in general terms, the actions of environing agencies, so far as they have operated indiscriminately, have produced in the stones a certain unity of character; at the same time that they have, by their differential effects, separated them: the larger ones having withstood certain violent actions which the smaller ones could not withstand.

Similarly with other assemblages of objects which are alike in their primary traits but unlike in their secondary traits. When simultaneously exposed to the same set of actions, some of these actions, rising to a certain intensity, may be expected to work on particular members of the assemblage changes which they cannot work in those which are markedly unlike; while others of the actions will work in all of them similar changes, because of the uniform relations between these actions and certain attributes common to all members of the assemblage. Hence it is inferable that on living organisms, which form an assemblage of this kind, and are unceasingly exposed in common to the agencies composing their inorganic environments, there must be wrought two such sets of effects. There will result a universal likeness among them consequent on the likeness of their respective relations to the matters and forces around; and there will result, in some cases, the differences due to the differential effects of these matters and forces, and in other cases, the changes which, being life-sustaining or life-destroying, eventuate in certain natural selections.

I have, above, made a passing reference to the fact that Mr. Darwin did not fail to take account of some among these effects directly produced on organisms by surrounding inorganic agencies. Here are extracts from the sixth edition of the _Origin of Species_ showing this.

“It is very difficult to decide how far changed conditions, such as
of climate, food, &c., have acted in a definite manner. There is
reason to believe that in the course of time the effects have been
greater than can be proved by clear evidence.... Mr. Gould believes
that birds of the same species are more brightly coloured under a
clear atmosphere, than when living near the coast or on islands; and
Wollaston is convinced that residence near the sea affects the
colours of insects. Moquin-Tandon gives a list of plants which, when
growing near the sea-shore, have their leaves in some degree fleshy,
though not elsewhere fleshy” (pp. 106-7). “Some observers are
convinced that a damp climate affects the growth of the hair, and
that with the hair the horns are correlated” (p. 159).

In his subsequent work, _Animals and Plants under Domestication_, Mr. Darwin still more clearly recognizes these causes of change in organization. A chapter is devoted to the subject. After premising that “the direct action of the conditions of life, whether leading to definite or indefinite results, is a totally distinct consideration from the effects of natural selection;” he goes on to say that changed conditions of life “have acted so definitely and powerfully on the organisation of our domesticated productions, that they have sufficed to form new sub-varieties or races, without the aid of selection by man or of natural selection.” Of his examples here are two.

“I have given in detail in the ninth chapter the most remarkable
case known to me, namely, that in Germany several varieties of maize
brought from the hotter parts of America were transformed in the
course of only two or three generations.” (Vol. ii, p. 277.) [And in
this ninth chapter concerning these and other such instances he says
“some of the foregoing differences would certainly be considered of
specific value with plants in a state of nature.” (Vol. i, p. 321.)]
“Mr. Meehan, in a remarkable paper, compares twenty-nine kinds of
American trees, belonging to various orders, with their nearest
European allies, all grown in close proximity in the same garden and
under as nearly as possible the same conditions.” And then
enumerating six traits in which the American forms all of them
differ in like ways from their allied European forms, Mr. Darwin
thinks there is no choice but to conclude that these “have been
definitely caused by the long-continued action of the different
climate of the two continents on the trees.” (Vol. ii, pp. 281-2.)

But the fact we have to note is that while Mr. Darwin thus took account of special effects due to special amounts and combinations of agencies in the environment, he did not take account of the far more important effects due to the general and constant operation of these agencies.[3] If a difference between the quantities of a force which acts on two organisms, otherwise alike and otherwise similarly conditioned, produces some difference between them; then, by implication, this force produces in both of them effects which they show in common. The inequality between two things cannot have a value unless the things themselves have values. Similarly if, in two cases, some unlikeness of proportion among the surrounding inorganic agencies to which two plants or two animals are exposed, is followed by some unlikeness in the changes wrought on them; then it follows that these several agencies taken separately, work changes in both of them. Hence we must infer that organisms have certain structural characters in common, which are consequent on the action of the medium in which they exist: using the word medium in a comprehensive sense, as including all physical forces falling upon them as well as matters bathing them. And we may conclude that from the primary characters thus produced there must result secondary characters.

Before going on to observe those general traits of organisms due to the general action of the inorganic environment upon them, I feel tempted to enlarge on the effects produced by each of the several matters and forces constituting the environment. I should like to do this not only to give a clear preliminary conception of the ways in which all organisms are affected by these universally-present agents, but also to show that, in the first place, these agents modify inorganic bodies as well as organic bodies, and that, in the second place, the organic are far more modifiable by them than the inorganic. But to avoid undue suspension of the argument, I content myself with saying that when the respective effects of gravitation, heat, light, &c, are studied, as well as the respective effects, physical and chemical, of the matters forming the media, water and air, it will be found that while more or less operative on all bodies, each modifies organic bodies to an extent immensely greater than the extent to which it modifies inorganic bodies.

* * * * *

Here, not discriminating among the special effects which these various forces and matters in the environment produce on both classes of bodies, let us consider their combined effects, and ask—What is the most general trait of such effects?

Obviously the most general trait is the greater amount of change wrought on the outer surface than on the inner mass. In so far as the matters of which the medium is composed come into play, the unavoidable implication is that they act more on the parts directly exposed to them than on the parts sheltered from them. And in so far as the forces pervading the medium come into play, it is manifest that, excluding gravity, which affects outer and inner parts indiscriminately, the outer parts have to bear larger shares of their actions. If it is a question of heat, then the exterior must lose it or gain it faster than the interior; and in a medium which is now warmer and now colder, the two must habitually differ in temperature to some extent—at least where the size is considerable. If it is a question of light, then in all but absolutely transparent masses, the outer parts must undergo more of any change producible by it than the inner parts—supposing other things equal; by which I mean, supposing the case is not complicated by any such convexities of the outer surface as produce internal concentrations of rays. Hence then, speaking generally, the necessity is that the primary and almost universal effect of the converse between the body and its medium, is to differentiate its outside from its inside. I say almost universal, because where the body is both mechanically and chemically stable, like, for instance, a quartz crystal, the medium may fail to work either inner or outer change.

Of illustrations among inorganic bodies, a convenient one is supplied by an old cannon-ball that has been long lying exposed. A coating of rust, formed of flakes within flakes, incloses it; and this thickens year by year, until, perhaps, it reaches a stage at which its exterior loses as much by rain and wind as its interior gains by further oxidation of the iron. Most mineral masses—pebbles, boulders, rocks—if they show any effect of the environment at all, show it only by that disintegration of surface which follows the freezing of absorbed water: an effect which, though mechanical rather than chemical, equally illustrates the general truth. Occasionally a “rocking-stone” is thus produced. There are formed successive layers relatively friable in texture, each of which, thickest at the most exposed parts, and being presently lost by weathering, leaves the contained mass in a shape more rounded than before; until, resting on its convex under-surface, it is easily moved. But of all instances perhaps the most remarkable is one to be seen on the west bank of the Nile at Philæ, where a ridge of granite 100 feet high, has had its outer parts reduced in course of time to a collection of boulder-shaped masses, varying from say a yard in diameter to six or eight feet, each one of which shows in progress an exfoliation of successively-formed shells of decomposed granite: most of the masses having portions of such shells partially detached.

If, now, inorganic masses, relatively so stable in composition, thus have their outer parts differentiated from their inner parts, what must we say of organic masses, characterized by such extreme chemical instability?—instability so great that their essential material is named protein, to indicate the readiness with which it passes from one isomeric form to another. Clearly the necessary inference is that this effect of the medium must be wrought inevitably and promptly, wherever the relation of outer and inner has become settled: a qualification for which the need will be seen hereafter.

* * * * *

Beginning with the earliest and most minute kinds of living things, we necessarily encounter difficulties in getting direct evidence; since, of the countless species now existing, all have been subject during millions upon millions of years to the evolutionary process, and have had their primary traits complicated and obscured by those endless secondary traits which the natural selection of favourable variations has produced. Among protophytes it needs but to think of the multitudinous varieties of diatoms and desmids, with their elaborately-constructed coverings; or of the definite methods of growth and multiplication among such simple _Algæ_ as the _Conjugatæ_; to see that most of their distinctive characters are due to inherited constitutions, which have been slowly moulded by survival of the fittest to this or that mode of life. To disentangle such parts of their developmental changes as are due to the action of the medium, is therefore hardly possible. We can hope only to get a general conception of it by contemplating the totality of the facts.