APPENDIX G.
THE INHERITANCE OF FUNCTIONALLY-CAUSED MODIFICATIONS.
In Part II, Chapter XA, I have confessed that the process by which a structure changed by use or disuse affects the sperm-cells or germ-cells whence arise descendants, is unimaginable: without, however, inferring that therefore such a process does not exist. With others it seems different. Some three years ago the following expression of opinion came to me from a zoological expert:—
“Many zoologists—most of us here at Cambridge—are intensely opposed to the doctrine of the inheritability of acquired variations. Even assuming that the developmental power of a germ is determined by its molecular structure (and I for one would question this—Driesch and his school when they find that they can squeeze a developing egg into all sorts of shapes without altering the final result, that one blastomere in an egg which has divided into 8 is still able to reproduce a whole embryo—question it also), we still fail to conceive any means by which, for instance, a change in the development of a muscle or nerve can effect a corresponding change in that part of the germ which is destined to produce a corresponding part in the descendant.”
Here it will be observed that belief in the inheritance of structural effects wrought by use and disuse, is rejected because of inability “to conceive any means” by which the modifications produced in an organ can effect a correlated modification in the germ of a descendant: failure to conceive is the test. The implication is that some alternative hypothesis is accepted because the correlating of a variation in an organ with a corresponding germ-variation is effected by a means which is conceivable. This is the hypothesis of Weismann. Concerning its conceivability I have, in the chapter just named, already written as follows:—
“If we follow Prof. Weismann we are led into an astounding supposition. He admits that every variable part must have a special determinant, and that this results in the assumption of over two hundred thousand for the four wings of a butterfly. Let us ask what must happen in the case of a peacock’s feather. On looking at the eye near its end, we see that the minute processes on the edge of each lateral thread must have been in some way exactly adjusted, in colour and position, so as to fall into line with the processes on adjacent threads: otherwise the symmetrical arrangement of coloured rings would be impossible. Each of these processes, then, being an independent variable, must have had its particular determinant. Now there are about 300 threads on the shaft of a large feather, and each of them bears on the average 1,600 processes, making for the whole feather 480,000 of these processes. For one feather alone there must have been 480,000 determinants, and for the whole tail many millions. And these, along with the determinants for the detailed parts of all the other feathers, and for the variable components of all organs forming the body at large, must have been contained in the microscopic head of a spermatozoon!” [And each of them must, throughout all the complex developmental processes, have preserved the ability to find its way to the exact place where it was wanted!]
If my Cambridge correspondent is able to conceive this process implied by the hypothesis of Weismann, I can only say that he has an enviable power of imagination.
But now comes the strange fact that an impossibility of thought implied by Weismann’s hypothesis does not cause rejection of it, but yet is urged as a reason for rejecting an alternative hypothesis which does not imply it. One objector cannot conceive that “a change in the development of a muscle or nerve can effect a corresponding change in that part of the germ which is destined to produce a corresponding part in the descendant”; and another objector says it is “very hard to believe” that a functionally-changed organ will so affect spermatozoa and ova that “one particular part of them will be so altered that the organisms which grow up from them will be able to present the same modification on the application of a different stimulus.” It is tacitly assumed by both that, as in the hypothesis of Weismann so in the counter-hypothesis, a particular part of the germ-plasm gives origin to a particular part of the developed organism. But nothing of the kind is implied. The nature of the counter-hypothesis (at any rate as held by me) is entirely misapprehended. Anyone who turns back to the chapters in the first volume where the conception of physiological units (or constitutional units) was set forth, or who re-reads the foregoing appendix, will see that there is altogether excluded any idea of correlation between certain parts of the germ and certain parts of the resulting organism. The units are supposed to be all alike, and during the progressive embryological changes local groups of them are supposed to take on different forms and structures under the combined forces, general and local, brought to bear on them. This conception is necessitated by all the evidence. The fact disclosed by the experiments of Driesch, Wilson, and Chabry, that from fractions of an ovum structures may be obtained like that obtained from the whole ovum, only smaller, necessitates it. The fact that any sufficiently large fragment of a polyp or planarian, no matter from what part of the body taken, will develop into a complete polyp or planarian necessitates it. The fact that from an undifferentiated portion of a plant, even so small as a scale, a complete plant may arise necessitates it. And it is necessitated by the fact that among plants, roots are produced by imbedded shoots and shoots by roots, as well as by the fact that low animals, such as hydroids, if deprived of both head and root, will develop a head from the root part and a root from the head part, if their respective conditions are inverted. All this evidence shows conclusively that the component units of each species, whether existing in the germ or in the developed organism, are, when not yet differentiated by local conditions, all alike, and that the notion of special parts of the germ-plasm correlated with special parts of the resulting organism, is entirely alien to the hypothesis.
“But how do the units of a modified organ affect the units of the germ in such wise that these produce an inherited modification of the organ?” will be asked. This difficulty has been dealt with in §§ 97d, 97e, where the analogy between the social organism and the individual organism has been brought in aid: serving, if not to furnish a conception, yet to furnish an adumbration. Regarding citizens as the units of an unfolding society, say a colony, it was pointed out that the nature they inherit from a mother-society gives them a proclivity towards a society of like structure, the traits of which are progressively assumed as the colony grows sufficiently large to make them possible. At the same time it was pointed out that while the influence of the entire aggregate on the individuals is seen in this forming of them into a society of the inherited type, the influences of local circumstances, and of individuals on one another, in each group, make them differentiate into appropriate social structures, taking on fit occupations and industries: the implication being that in virtue of their inherited natures they all have partial capacities for the various activities they undertake; so that an immigrant clerk sets up a tavern, a compositor takes to carpentering, and a university man rides after cattle or is employed on a sheep farm. Evidence was given in that place, as in the above paragraph, that the constitutional units of an organism similarly have all of them potentialities for taking on this or that structure and mode of action which local conditions determine. It was further argued that as citizens are continually being remoulded by their society into congruity with it, and, if circumstances change them, tend to remould their society; so in the individual organism, there is this reciprocal action of the whole on the units and of the units on the whole. Hence it was inferred that the modified units in any modified part tend to diffuse modifications like their own through the units at large: being aided by the circulation of protoplasm, as suggested in §§ 54d and 97f. And it was urged that, however inconceivably complex such a process may be, yet it seems not incredible when we recognise the probability that an organism is more or less permeable to undulations propagated by its molecules: Rontgen rays giving warrant. If such units throughout the tissues may take in and send out ethereal waves which bring it into rhythmical relations with others of its kind and tend to produce congruity, it becomes, if not conceivable still supposable, that throughout the circulating protoplasm there goes on a continual harmonization of its components—a moulding of each by all and of all by each. Should it be said that such a process is too marvellous to be reasonably assumed, the reply is that it is not more marvellous than heredity itself, which, were it not familiar to us, would be thought incredible.
But as I have said in the place referred to—“At last then we are obliged to admit that the actual organizing process transcends conception. It is not enough to say that we cannot know it; we must say that we cannot even conceive it:” can only conceive the possibility of a suggested interpretation.
Hence we have to rely upon evidences of other kinds. Among these, some which I think dispose absolutely of the fashionable hypothesis while they harmonize with the opposed hypothesis, have now to be named. That their implication should not have been generally recognized would have seemed to me incomprehensible were it not that I have myself only now observed this implication. The facts are these:—
“Verlot mentions a gardener who could distinguish 150 kinds of camellia, when not in flower; and it has been positively asserted that the famous old Dutch florist Voorhelm, who kept above 1,200 varieties of the hyacinth, was hardly ever deceived in knowing each variety by the bulb alone. Hence we must conclude that the bulbs of the hyacinth and the branches and leaves of the camellia, though appearing to an unpractised eye absolutely undistinguishable, yet really differ.” (Darwin, Variation of Animals and Plants, &c., vol. ii, p. 251.)
More recently testimony to like effect has been given by Dr. Maxwell Masters, and has already been quoted by me in a note to § 286 in illustration of another truth. He says concerning such variations:—
“To the untrained eye, the primordial differences noted are often very slight; even the botanist, unless his attention be specially directed to the matter, fails to see minute differences which are perceptible enough to the raiser or his workmen.... These apparently trifling morphological differences are often associated with physiological variations which render some varieties, say of wheat, much better enabled to resist mildew and disease generally than others. Some, again, prove to be better adapted for certain soils or for some climates than others; some are less liable to injury from predatory birds than others, and so on.”
In his Vegetable Teratology, p. 493, Dr. Masters names another fact having a like implication—the fact that among seedling stocks which have not yet flowered, those which will produce double flowers are distinguishable. He says:—
“This separation of the single from the double-flowered plants, M. Chatié tells us is not so difficult as might be supposed. The single stocks, he explains, have deep green leaves (glabrous in certain species), rounded at the top, the heart being in the form of a shuttlecock, and the plant stout and thick-set in its general aspect, while the plants yielding double flowers have very long leaves of a light green colour, hairy and curled at the edges, the heart consisting of whitish leaves, curved so that they enclose it completely.”
What is the general truth implied? Clearly that there exists no such thing as an independent local variation. Some marked change in the form or colour of a flower or a fruit draws attention; and, being a change which interests the florist or gardener, pecuniarily or otherwise, not only draws attention but usually monopolizes attention: the natural impression produced being that this variation stands there by itself—is without relation to variations elsewhere. But now it turns out that there are concomitant variations all over the plant. Even in underground bulbs certain appreciable differences go along with certain conspicuous differences in the flowers. And if along with a striking change in a flower which the florist contemplates, there go changes all over the plant not obvious to careless observers but visible to him, we must infer that there are everywhere minute differences which even the florist cannot perceive: the whole constitution of the plant has diverged in some measure from the constitutions of kindred plants. Every local variation implies a change pervading the entire organism, manifested in concomitant variations everywhere else.
If so, what becomes of the hypothesis of determinants—the hypothesis that there is a special element in the germ-plasm which results in a special local modification in the adult organism? That there are no facts supporting it has been all along manifest; but now it is manifest that the facts directly contradict it.
At the same time it may be remarked that while the facts are wholly incongruous with the hypothesis of determinants and its accompanying elaborate speculation, they are not incongruous with the alternative hypothesis. Impossible though it may be to imagine the natures of those ultimate units peculiar to each species, which have proclivities towards the particular form of organization characterizing it, yet that a change of structure arising in one part of the organism is accompanied by multitudinous changes of structure in other parts of the organism, is not only congruous with the belief that there exist such constitutional units, but yields it distinct support. For if, as above argued, a conspicuous local variation is not the result of any modification of units special to the locality, but is the result of a modification of the units at large, then it must happen that such modification must have its effects on all other parts of the organism; so that there cannot fail to result all those small concomitant variations above indicated.
May we not also say that it becomes less incomprehensible that structural changes caused by use and disuse are inherited? If, as we see, a local variation spontaneously arising is accompanied by multitudinous other local variations, implying a necessary correlation between each local variation and the general constitution of the organism; then it may be argued that if a marked change of function in an organ causes increase or decrease of it, this general correlation implies that there must be a reciprocal reaction between the part and the whole, tending to re-establish their congruity. The constitution at large will in so far be changed, and along with its change will go corresponding changes in the sperm-cells and germ-cells.
Finally let me add, not another argument, but another fact of observation, of the kind which opponents demand, but which, when they are from time to time furnished, are severally pooh-poohed as not enough. Each of them is spoken of as a solitary fact and slighted as inadequate; and when by and by another is named, this is treated in the same way; so that the facts which if brought together would be recognized as sufficient are never brought together. That to which I refer is set forth in a pamphlet by M. Leo Errera, Professor at the University of Brussels, entitled “Hérédite d’un Caractère acquis chez un Champignon pluricellulaire;” being an account of experiments of Dr. Hunger, at the Botanical Institute in Brussels. First enumerating various instances of adaptations to climate, as those of plants which, fitted to northern regions, preserve their constitutional rapidity of growth and seeding when brought south, and do this for several generations, he goes on to detail the culture-experiments of M. Hunger, and sums up the results of these in the following words:—
“On déduit de là que:
“1o Les conidies d’Aspergillus niger sont adaptées à la concentration du milieu où a vécu l’individu qui les porte; cet effet est encore plus marqué après deux générations passées dans un milieu donné (Expér. I et II);
“2o II s’agit d’une véritable adaptation et non pas simplement d’un accroissement de vigueur chez les conidies provenant des liquides concentrés, car ces mêmes conidies germent moins rapidement et donnent des plantes moins vigoureuses que les conidies normales lorsqu’on les sème de nouveau sur le milieu-type: en s’adaptant aux liquides concentrés, elles se sont désadaptées du liquide normal (Expér. III);
“3o Une génération passée sur le liquide normal n’efface pas l’influence d’une ou de deux générations antérieures passées sur une liquide plus concentré (Expér. IV).
“Tous ces résultats concordent: ils montrent une légère, mais incontestable transmission héréditaire de l’adaptation au milieu.”