Essays: Scientific, Political, and Speculative, Volume I

"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.

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.