The Principles of Biology Vol. I/Chapter II.8

§ 80. Already, in the last two chapters, the law of hereditary transmission has been tacitly assumed; as, indeed, it unavoidably is in all such discussions. Understood in its entirety, the law is that each plant or animal, if it reproduces, gives origin to others like itself: the likeness consisting, not so much in the repetition of individual traits as in the assumption of the same general structure. This truth has been rendered so familiar by daily illustration as almost to have lost its significance. That wheat produces wheat—that existing oxen have descended from ancestral oxen—that every unfolding organism eventually takes the form of the class, order, genus, and species from which it sprang; is a fact which, by force of repetition, has acquired in our minds almost the aspect of a necessity. It is in this, however, that Heredity is principally displayed: the manifestations of it commonly referred to being quite subordinate. And, as thus understood, Heredity is universal. The various instances of heterogenesis lately contemplated seem, indeed, to be at variance with this assertion. But they are not really so. Though the recurrence of like forms is, in these instances, not direct but cyclical, still, the like forms do recur; and, when taken together, the group of forms produced during one of the cycles is as much like the groups produced in preceding cycles, as the single individual arising by homogenesis is like ancestral individuals.

While, however, the general truth that organisms of a given type uniformly descend from organisms of the same type, is so well established by infinite illustrations as to have assumed the character of an axiom; it is not universally admitted that non-typical peculiarities are inherited. Many entertain a vague belief that the law of Heredity applies only to main characters of structure and not to details; or, at any rate, that though it applies to such details as constitute differences of species, it does not apply to smaller details. The circumstance that the tendency to repetition is in a slight degree qualified by the tendency to variation (which, as we shall hereafter see, is but an indirect result of the tendency to repetition), leads some to doubt whether Heredity is unlimited. A careful weighing of the evidence, however, and a due allowance for the influences by which the minuter manifestations of Heredity are obscured, may remove this scepticism.

First in order of importance comes the fact that not only are there uniformly transmitted from an organism to its offspring, those traits of structure which distinguish the class, order, genus, and species; but also those which distinguish the variety. We have numerous cases, among both plants and animals, where, by natural or artificial conditions, there have been produced divergent modifications of the same species; and abundant proof exists that the members of any one sub-species habitually transmit their distinctive peculiarities to their descendants. Agriculturists and gardeners can furnish unquestionable illustrations. Several varieties of wheat are known, of which each reproduces itself. Since the potato was introduced into England there have been formed from it a number of sub-species; some of them differing greatly in their forms, sizes, qualities, and periods of ripening. Of peas, also, the like may be said. And the case of the cabbage-tribe is often cited as showing the permanent establishment of races which have diverged widely from a common stock. Among fruits and flowers the multiplication of kinds, and the continuance of each kind with certainty by agamogenesis, and to some extent by gamogenesis, might be exemplified without end. From all sides evidence may be gathered showing a like persistence of varieties among animals. We have our distinct breeds of sheep, our distinct breeds of cattle, our distinct breeds of horses: each breed maintaining its characteristics. The many sorts of dogs which, if we accept the physiological test, we must consider as all of one species, show us in a marked manner the hereditary transmission of small differences—each sort, when kept pure, reproducing itself not only in size, form, colour, and quality of hair, but also in disposition and speciality of intelligence. Poultry, too, have their permanently-established races. And the Isle of Man sends us a tail-less kind of cat. Even in the absence of other evidence, that which ethnology furnishes would suffice. Grant them to be derived from one stock, and the varieties of man yield proof upon proof that non-specific traits of structure are bequeathed from generation to generation. Or grant only their derivation from several stocks, and we still have, between races descended from a common stock, distinctions which prove the inheritance of minor peculiarities. Besides seeing the Negroes continue to produce Negroes, copper-coloured men to produce men of a copper colour, and the fair-skinned races to perpetuate their fair skins—besides seeing that the broad-faced and flat-nosed Calmuck begets children with broad faces and flat noses, while the Jew bequeaths to his offspring the features which have so long characterized Jews; we see that those small unlikenesses which distinguish more nearly-allied varieties of men, are maintained from generation to generation. In Germany, the ordinary shape of skull is appreciably different from that common in Britain: near akin though the Germans are to the British. The average Italian face continues to be unlike the faces of northern nations. The French character is now, as it was centuries ago, contrasted in sundry respects with the characters of neighbouring peoples. Nay, even between races so closely allied as the Scotch Celts, the Welsh Celts, and the Irish Celts, appreciable differences of form and nature have become established.

The fact that sub-species and sub-sub-species thus exemplify the general law of inheritance which shows itself in the perpetuation of ordinal, generic, and species peculiarities, is strong reason for the belief that this general lay is unlimited in its application. This has the support of still more special evidences. They are divisible into two classes. In the one come cases where congenital peculiarities, not traceable to any obvious causes, are bequeathed to descendants. In the other come cases where the peculiarities thus bequeathed are not congenital, but have resulted from changes of functions during the lives of the individuals bequeathing them. We will consider first the cases that come in the first class.

§ 81. Note at the outset the character of the chief testimony. Excluding those inductions that have been so fully verified as to rank with exact science, there are no inductions so trustworthy as those which have undergone the mercantile test. When we have thousands of men whose profit or loss depends on the truth of their inferences from perpetually-repeated observations; and when we find that their inferences, handed down from generation to generation, have generated an unshakable conviction; we may accept it without hesitation. In breeders of animals we have such a class, led by such experiences, and entertaining such a conviction—the conviction that minor peculiarities of organization are inherited as well as major peculiarities. Hence the immense prices given for successful racers, bulls of superior forms, sheep that have certain desired peculiarities. Hence the careful record of pedigrees of high-bred horses and sporting dogs. Hence the care taken to avoid intermixture with inferior stocks. As quoted by Mr. Darwin, Youatt says the principle of selection "enables the agriculturist not only to modify the character of his flock but to change it altogether." Lord Somerville, speaking of what breeders have done for sheep, says:—"It would seem that they have chalked upon a wall a form perfect in itself and then given it existence." That most skilful breeder, Sir John Sebright, used to say, with respect to pigeons, that "he would produce any given feather in three years, but it would take him six years to obtain head and beak." In all which statements the tacit assertion is, that individual traits are bequeathed from generation to generation, and may be so perpetuated and increased as to become permanent distinctions.

Of special instances there are many besides that of the often-cited Otto-breed of sheep, descended from a single short-legged lamb, and that of the six-fingered Gratio Kelleia, who transmitted his peculiarity, in different degrees, to several of his children and to some of his grandchildren. In a paper contributed to the Edinburgh New Philosophical Journal for July, 1863, Dr. (now Sir John) Struthers gives cases of hereditary digital variations. Esther P——, who had six fingers on one hand, bequeathed this malformation along some lines of her descendants for two, three, and four generations. A—— S—— inherited an extra digit on each hand and each foot from his father; and C—— G——, who also had six fingers and six toes, had an aunt and a grandmother similarly formed. A collection of evidence published by Mr. Sedgwick in the Medico-Chirurgical Review for April and for July, 1863, in two articles on "The Influence of Sex in limiting Hereditary Transmission," includes the following cases:—Augustin Duforet, a pastry-cook of Douai, who had but two instead of three phalanges to all his fingers and toes, inherited this malformation from his grandfather and father, and had it in common with an uncle and numerous cousins. An account has been given by Dr. Lepine, of a man with only three fingers on each hand and four toes on each foot, and whose grandfather and son exhibited the like anomaly. Béchet describes Victoire Barré as a woman who, like her father and sister, had but one developed finger on each hand and but two toes on each foot, and whose monstrosity re-appeared in two daughters. And there is a case where the absence of two distal phalanges on the hands was traced for two generations. The various recorded instances in which there has been transmission from one generation to another, of webbed-fingers, of webbed-toes, of hare-lip, of congenital luxation of the thigh, of absent patellæ, of club-foot, &c., would occupy more space than can here be spared. Defects in the organs of sense are also not unfrequently inherited. Four sisters, their mother, and grandmother, are described by Duval as similarly affected by cataract. Prosper Lucas details an example of amaurosis affecting the females of a family for three generations. Duval, Graffe, Dufon, and others testify to like cases coming under their observation. Deafness, too, is occasionally transmitted from parent to child. There are deaf-mutes whose imperfections have been derived from ancestors; and malformations of the external ears have also been perpetuated in offspring. Of transmitted peculiarities of the skin and its appendages, many cases have been noted. One is that of a family remarkable for enormous black eyebrows; another that of a family in which every member had a lock of hair of a lighter colour than the rest on the top of the head; and there are also instances of congenital baldness being hereditary. From one of our leading sculptors I learn that his wife has a flat mole under the foot near the little toe, and one of her sons has the same. Entire absence of teeth, absence of particular teeth, and anomalous arrangements of teeth, are recorded as traits that have descended to children. And we have evidence that soundness and unsoundness of teeth are transmissible.

The inheritance of tendencies to such diseases as gout, consumption, and insanity is universally admitted. Among the less-common diseases of which the descent has been observed, are ichthyosis, leprosy, pityriasis, sebaceous tumours, plica polonica, dipsomania, somnambulism, catalepsy, epilepsy, asthma, apoplexy, elephantiasis. General nervousness displayed by parents almost always re-appears in their children. Even a bias towards suicide appears to be sometimes hereditary.

§ 82. To prove the transmission of those structural peculiarities which have resulted from functional peculiarities, is, for several reasons, comparatively difficult. Changes produced in the sizes of parts by changes in their amounts of action, are mostly unobtrusive. A muscle which has increased in bulk is usually so obscured by natural or artificial clothing, that unless the alteration is extreme it passes without remark. Such nervous developments as are possible in the course of a single life, cannot be seen externally. Visceral modifications of a normal kind are observable but obscurely, or not at all. And if the changes of structure worked in individuals by changes in their habits are thus difficult to trace, still more difficult to trace must be the transmission of them: further hidden, as this is, by the influences of other individuals who are often otherwise modified by other habits. Moreover, such specialities of structure as are due to specialities of function, are usually entangled with specialities of structure which are, or may be, due to selection, natural or artificial. In most cases it is impossible to say that a structural peculiarity which seems to have arisen in offspring from a functional peculiarity in a parent, is wholly independent of some congenital peculiarity of structure in the parent, whence this functional peculiarity arose. We are restricted to cases with which natural or artificial selection can have had nothing to do, and such cases are difficult to find. Some, however, may be noted.

A species of plant that has been transferred from one soil or climate to another, frequently undergoes what botanists call "change of habit"—a change which, without affecting its specific characters, is yet conspicuous. In its new locality the species is distinguished by leaves that are much larger or much smaller, or differently shaped, or more fleshy; or instead of being as before comparatively smooth, it becomes hairy; or its stem becomes woody instead of being herbaceous; or its branches, no longer growing upwards, assume a drooping character. Now these "changes of habit" are clearly determined by functional changes. Occurring, as they do, in many individuals which have undergone the same transportation, they cannot be classed as "spontaneous variations." They are modifications of structure consequent on modifications of function that have been produced by modifications in the actions of external forces. And as these modifications re-appear in succeeding generations, we have, in them, examples of functionally-established variations that are hereditarily transmitted.

Evidence of analogous changes in animals is difficult to disentangle. Only among domesticated kinds have we any opportunity of tracing the results of altered habits; and here, in nearly all cases, artificial selection has obscured them. Still, there are some facts which seem to the point. Mr. Darwin, while ascribing almost wholly to "natural selection" the production of those modifications which eventuate in differences of species, nevertheless admits the effects of use and disuse. He says—"I find in the domestic duck that the bones of the wing weigh less and the bones of the leg more, in proportion to the whole skeleton, than do the same bones in the wild duck; and I presume that this change may be safely attributed to the domestic duck flying much less, and walking more, than its wild parent. The great and inherited development of the udders in cows and goats in countries where they are habitually milked, in comparison with the state of these organs in other countries, is another instance of the effect of use. Not a single domestic animal can be named which has not in some country drooping ears; and the view suggested by some authors, that the drooping is due to the disuse of the muscles of the ear, from the animals not being much alarmed by danger, seems probable." Again—"The eyes of moles and of some burrowing rodents are rudimentary in size, and in some cases are quite covered up by skin and fur. This state of the eyes is probably due to gradual reduction from disuse, but aided perhaps by natural selection." ... "It is well known that several animals belonging to the most different classes, which inhabit the caves of Styria and of Kentucky, are blind. In some of the crabs the footstalk of the eye remains, though the eye is gone; the stand for the telescope is there, though the telescope with its glasses has been lost. As it is difficult to imagine that eyes, though useless, could be in any way injurious to animals living in darkness, I attribute their loss wholly to disuse." The direct inheritance of an acquired peculiarity is sometimes observable. Mr. Lewes gives a case. He "had a puppy taken from its mother at six weeks old, who, although never taught 'to beg' (an accomplishment his mother had been taught), spontaneously took to begging for everything he wanted when about seven or eight months old: he would beg for food, beg to be let out of the room, and one day was found opposite a rabbit hutch begging for rabbits." Instances are on record, too, of sporting dogs which spontaneously adopted in the field, certain modes of behaviour which their parents had learnt.

But the best examples of inherited modifications produced by modifications of function, occur in mankind. To no other cause can be ascribed the rapid metamorphoses undergone by the British races when placed in new conditions. In the United States the descendants of the immigrant Irish lose their Celtic aspect, and become Americanized. This cannot be ascribed to mixture, since the feeling with which Irish are regarded by Americans prevents any considerable amount of intermarriage. Equally marked is the case of the immigrant Germans who, though they keep very much apart, rapidly assume the prevailing type. To say that "spontaneous variation" increased by natural selection, can have produced this effect, is going too far. Peoples so numerous cannot have been supplanted in the course of two or three generations by varieties springing from them. Hence the implication is that physical and social conditions have wrought modifications of function and structure, which offspring have inherited and increased. Similarly with special cases. In the Cyclopædia of Practical Medicine, Vol. II., p. 419, Dr. Brown states that he "has in many instances observed in the case of individuals whose complexion and general appearance has been modified by residence in hot climates, that children born to them subsequently to such residence, have resembled them rather in their acquired than primary mien."

Some visible modifications of organs caused by changes in their functions, may be noted. That large hands are inherited by those whose ancestors led laborious lives, and that those descended from ancestors unused to manual labour commonly have small hands, are established opinions. It seems very unlikely that in the absence of any such connexion, the size of the hand should have come to be generally regarded as some index of extraction. That there exists a like relation between habitual use of the feet and largeness of the feet, we have strong evidence in the customs of the Chinese. The torturing practice of artificially arresting the growth of the feet, could never have become established among the ladies of China, had they not seen that a small foot was significant of superior rank—that is of a luxurious life—that is of a life without bodily labour. There is evidence, too, that modifications of the eyes, caused by particular uses of the eyes, are inherited. Short sight appears to be uncommon among peasants; but it is frequent among classes who use their eyes much for reading and writing, and is often congenital. Still more marked is this relation in Germany. There, the educated are notoriously studious, and judging from the numbers of young Germans who wear spectacles, there is reason to think that congenital myopia is very frequent among them.

Some of the best illustrations of functional heredity, are furnished by mental characteristics. Certain powers which mankind have gained in the course of civilization cannot, I think, be accounted for without admitting the inheritance of acquired modifications. The musical faculty is one of these. To say that "natural selection" has developed it by preserving the most musically endowed, seems an inadequate explanation. Even now that the development and prevalence of the faculty have made music an occupation by which the most musical can get sustenance and bring up families; it is very questionable whether, taking the musical career as a whole, it has any advantage over other careers in the struggle for existence and multiplication. Still more if we look back to those early stages through which the faculty must have passed before definite perception of melody was arrived at, we fail to see how those possessing the rudimentary faculty in a somewhat greater degree than the rest, would thereby be enabled the better to maintain themselves and their children. There is no explanation but that the habitual association of certain cadences of speech with certain emotions, has slowly established in the race an organized and inherited connection between such cadences and such emotions; that the combination of such cadences, more or less idealized, which constitutes melody, has all along had a meaning in the average mind, only because of the meaning which cadences had acquired in the average mind; and that by the continual hearing and practice of melody there has been gained and transmitted an increasing musical sensibility. Confirmation of this view may be drawn from individual cases. Grant that among a people endowed with musical faculty to a certain degree, spontaneous variation will occasionally produce men possessing it in a higher degree; it cannot be granted that spontaneous variation accounts for the frequent production, by such highly-endowed men, of men still more highly endowed. On the average, the children of marriages with others not similarly endowed, will be less distinguished rather than more distinguished. The most that can be expected is that this unusual amount of faculty shall re-appear in the next generation undiminished. How then shall we explain cases like those of Bach, Mozart, and Beethoven, all of them sons of men having unusual musical powers who were constantly exercising those powers, and who greatly excelled their fathers in their musical powers? What shall we say to the facts that Haydn was the son of an organist, that Hummel was born to a music master, and that Weber's father was a distinguished violinist? The occurrence of so many cases in one nation within a short period of time, cannot rationally be ascribed to the coincidence of "spontaneous variations." It can be ascribed to nothing but inherited developments of structure caused by augmentations of function.

But the clearest proof that structural alterations caused by alterations of function are inherited, occurs when the alterations are morbid. I had originally named in this place the results of M. Brown-Sequard's experiments on guinea-pigs, showing that those which had been artificially made epileptic had offspring which were epileptic; and I name them again though his inference is by many rejected. For, as exemplified a few pages back, strong evidence is often disregarded for trivial reasons by those who dislike the conclusion drawn. Just naming this evidence and its possible invalidity, let me pass to some results of experiences recently set forth by Dr. Savage, President of the Neurological Society. In an essay on "Heredity and Neurosis" published in Brain, Parts LXXVII, LXXVIII, 1897, he says:—"We recognise the transmission of a tendency to develop gout, and we recognise that the disease produced by the individual himself differs little from that which may have been inherited." [That is, acquired gout may be transmitted as constitutional gout.] "I have seen several patients whose history I have been able to examine carefully, in whom mental tricks have been transmitted from one generation to another." In the "musical prodigies" descending from musical parents, "there seemed to be a transmission of a greatly increased aptitude or tendency which is all one is contending for." "Though there is, in my opinion, power to transmit acquired peculiarities, yet the tendency is to transmit a predisposition." (pp. 19-21.) And an authority on nervous diseases who is second to none—Dr. Hughlings Jackson—takes the same view. The liability to consumption shown by children of consumptive parents, which no one doubts, shows us the same thing. It is admitted that consumption may be produced by conditions very unfavourable to life; and unless it is held that the disease so produced differs from the disease when inherited, the conclusion must be that here, too, there is a transmission of functionally-produced organic changes. This holds true whether the production of tubercle is due to innate defect or whether it is due to the invasion of a bacillus. For in this last case the consumptive diathesis must be regarded as a state of body more than usually liable to invasion by the bacillus, and this is the same when acquired as when transmitted.

§ 83. Two modified manifestations of Heredity remain to be noticed. The one is the re-appearance in offspring of traits not borne by the parents, but borne by the grandparents or by remoter ancestors. The other is the limitation of Heredity by sex—the restriction of transmitted peculiarities to offspring of the same sex as the parent possessing them.

Atavism, which is the name given to the recurrence of ancestral traits, is proved by many and varied facts. In the picture-galleries of old families, and on the monumental brasses in the adjacent churches, are often seen types of feature which are still, from time to time, repeated in members of these families. It is a matter of common remark that some constitutional diseases, such as gout and insanity, after missing a generation, will show themselves in the next. Dr. Struthers, in his above-quoted paper "On Variation in the Number of Fingers and Toes, and in the Phalanges in Man," gives cases of malformations common to grandparent and grandchild, but of which the parent had no trace. M. Girou (as quoted by Mr. Sedgwick) says—"One is often surprised to see lambs black, or spotted with black, born of ewes and rams with white wool, but if one takes the trouble to go back to the origin of this phenomena, it is found in the ancestors." Instances still more remarkable, in which the remoteness of the ancestors copied is very great, are given by Mr. Darwin. He points out that in crosses between varieties of the pigeon, there will sometimes re-appear the plumage of the original rock-pigeon, from which these varieties descended; and he thinks the faint zebra-like markings occasionally traceable in horses have probably a like meaning.

The other modified manifestation of heredity above referred to is the limitation of heredity by sex. In Mr. Sedgwick's essays, already named, will be found evidence implying that there exists some such tendency to limitation, which does or does not show itself distinctly according to the nature of the organic modification to be conveyed. On joining to the evidence he gives certain bodies of allied evidence we shall, I think, find the inconsistences comprehensible.

Beyond the familiar facts that in ourselves, along with the essential organs of sex there go minor structures and traits distinctive of sex, such as the beard and the voice in man, we have numerous cases in which, along with different sex-organs there go general differences, sometimes immense and often conspicuous. We have those in which (as in sundry parasites) the male is extremely small compared with the female; we have those in which the male is winged and the female wingless; we have those, as among birds, in which the plumage of males contrasts strongly with that of females; and among butterflies we have kindred instances in which the wings of the two sexes are wholly unlike—some, indeed, in which there is not simply dimorphism but polymorphism: two kinds of females both differing from the male. How shall we range these facts with the ordinary facts of inheritance? Without difficulty if heredity results from the proclivity which the component units contained in a germ-cell or a sperm-cell have to arrange themselves into a structure like that of the structure from which they were derived. For the obvious corollary is that where there is gamogenesis there will result partly concurring and partly conflicting proclivities. In the fertilized germ we have two groups of physiological units, slightly different in their structures. These slightly-different units severally multiply at the expense of the nutriment supplied to the unfolding germ—each kind moulding this nutriment into units of its own type. Throughout the process of development the two kinds of units, mainly agreeing in their proclivities and in the form which they tend to build themselves into, but having minor differences, work in unison to produce an organism of the species from which they were derived, but work in antagonism to produce copies of their respective parent-organisms. And hence ultimately results an organism in which traits of the one are mixed with traits of the other; and in which, according to the predominance of one or other group of units, one or other sex with all its concomitants is produced.

If so, it becomes comprehensible that with the predominance of either group, and the production of the same sex as that of the parent whence it was derived, there will go the repetition not only of the minor sex-traits of that parent but also of any peculiarities he or she possessed, such as monstrosities. Since the two groups are nearly balanced, and since inheritance is never an average of the two parents but a mixture of traits of the one with traits of the other, it is not difficult to see why there should be some irregularity in the transmission of these monstrosities and constitutional tendencies, though they are most frequently transmitted only to those of the same sex.

§ 84. Unawares in the last paragraph there has been taken for granted the truth of that suggestion concerning Heredity ventured in § 66. Anything like a positive explanation is not to be expected in the present stage of Biology, if at all. We can look for nothing beyond a simplification of the problem; and a reduction of it to the same category with certain other problems which also admit of hypothetical solutions only. If an hypothesis which sundry widespread phenomena have already thrust upon us, can be shown to render the phenomena of Heredity more intelligible than they at present seem, we shall have reason to entertain it. The applicability of any method of interpretation to two different but allied classes of facts, is evidence of its truth.

The power which many animals display of reproducing lost parts, we saw to be inexplicable except on the assumption that the units of which any organism is built have a tendency to arrange themselves into the shape of that organism ( § 65). This power is sufficiently remarkable in cases where a lost limb or tail is replaced, but it is still more remarkable in cases where, as among some annelids, the pieces into which an individual is cut severally complete themselves by developing heads and tails, or in cases like that of the Holothuria, which having, when alarmed, ejected its viscera, reproduces them. Such facts compel us to admit that the components of an organism have a proclivity towards a special structure—that the adult organism when mutilated exhibits that same proclivity which is exhibited by the young organism in the course of its normal development. As before said, we may, for want of a better name, figuratively call this power organic polarity: meaning by this phrase nothing more than the observed tendency towards a special arrangement. And such facts as those presented by the fragments of a Hydra, and by fragments of leaves from which complete plants are produced, oblige us to recognize this proclivity as existing throughout the tissues in general—nay, in the case of the Begonia phyllomaniaca, obliges us to recognize this proclivity as existing in the physiological units contained in each undifferentiated cell. Quite in harmony with this conclusion, are certain implications since noticed, respecting the characters of sperm-cells and germ-cells. We saw sundry reasons for rejecting the supposition that these are highly-specialized cells and for accepting the opposite supposition, that they are cells differing from others rather in being unspecialized. And here the assumption to which we seem driven by the ensemble of the evidence, is, that sperm-cells and germ-cells are essentially nothing more than vehicles in which are contained small groups of the physiological units in a fit state for obeying their proclivity towards the structural arrangement of the species they belong to.

If the likeness of offspring to parents is thus determined, it becomes manifest, à priori, that besides the transmission of generic and specific peculiarities, there will be a transmission of those individual peculiarities which, arising without assignable causes, are classed as "spontaneous." For if the assumption of a special arrangement of parts by an organism, is due to the proclivity of its physiological units towards that arrangement; then the assumption of an arrangement of parts slightly different from that of the species, implies physiological units slightly unlike those of the species; and these slightly-unlike physiological units, communicated through the medium of sperm-cell or germ-cell, will tend, in the offspring, to build themselves into a structure similarly diverging from the average of the species.

But it is not equally manifest that, on this hypothesis, alterations of structure caused by alterations of function must be transmitted to offspring. It is not obvious that change in the form of a part, caused by changed action, involves such change in the physiological units throughout the organism that these, when groups of them are thrown off in the shape of reproductive centres, will unfold into organisms that have this part similarly changed in form. Indeed, when treating of Adaptation ( § 69), we saw that an organ modified by increase or decrease of function, can but slowly re-act on the system at large, so as to bring about those correlative changes required to produce a new equilibrium; and yet only when such new equilibrium has been established, can we expect it to be fully expressed in the modified physiological units of which the organism is built—only then can we count on a complete transfer of the modification to descendants. Nevertheless, that changes of structure caused by changes of action must also be transmitted, however obscurely, appears to be a deduction from first principles—or if not a specific deduction, still, a general implication. For if an organism A, has, by any peculiar habit or condition of life, been modified into the form A′, it follows that all the functions of A′, reproductive function included, must be in some degree different from the functions of A. An organism being a combination of rhythmically-acting parts in moving equilibrium, the action and structure of any one part cannot be altered without causing alterations of action and structure in all the rest; just as no member of the Solar System could be modified in motion or mass, without producing rearrangements throughout the whole Solar System. And if the organism A, when changed to A′, must be changed in all its functions; then the offspring of A′ cannot be the same as they would have been had it retained the form A. That the change in the offspring must, other things equal, be in the same direction as the change in the parent, appears implied by the fact that the change propagated throughout the parental system is a change towards a new state of equilibrium—a change tending to bring the actions of all organs, reproductive included, into harmony with these new actions. Or, bringing the question to its ultimate and simplest form, we may say that as, on the one hand, physiological units will, because of their special polarities, build themselves into an organism of a special structure; so, on the other hand, if the structure of this organism is modified by modified function, it will impress some corresponding modification on the structures and polarities of its units. The units and the aggregate must act and re-act on each other. If nothing prevents, the units will mould the aggregate into a form in equilibrium with their pre-existing polarities. If, contrariwise, the aggregate is made by incident actions to take a new form, its forces must tend to re-mould the units into harmony with this new form. And to say that the physiological units are in any degree so re-moulded as to bring their polar forces towards equilibrium with the forces of the modified aggregate, is to say that when separated in the shape of reproductive centres, these units will tend to build themselves up into an aggregate modified in the same direction.

—A large amount of additional evidence supporting the belief that functionally produced modifications are inherited, will be found in Appendix B.