Chapter 9

During this period nevertheless one distinct advance was made, that with which Weismann's name is prominently connected. In Darwin's genetic scheme the hereditary transmission of parental experience and its consequences played a considerable role. Exactly how great that role was supposed to be, he with his habitual caution refrained from specifying, for the sufficient reason that he did not know. Nevertheless much of the process of Evolution, especially that by which organs have become degenerate and rudimentary, was certainly attributed by Darwin to such inheritance, though since belief in the inheritance of acquired characters fell into dispute, the fact has been a good deal overlooked. The _Origin_ without "use and disuse" would be a materially different book. A certain vacillation is discernible in Darwin's utterances on this question, and the fact gave to the astute Butler an opportunity for his most telling attack. The discussion which best illustrates the genetic views of the period arose in regard to the production of the rudimentary condition of the wings of many beetles in the Madeira group of islands, and by comparing passages from the _Origin_[64] Butler convicts Darwin of saying first that this condition was in the main the result of Selection, with disuse aiding, and in another place that the main cause of degeneration was disuse, but that Selection had aided. To Darwin however I think the point would have seemed one of dialetics merely. To him the one paramount purpose was to show that somehow an Evolution by means of Variation and Heredity might have brought about the facts observed, and whether they had come to pass in the one way or the other was a matter of subordinate concern.

To us moderns the question at issue has a diminished significance. For over all such debates a change has been brought by Weismann's challenge for evidence that use and disuse have any transmitted effects at all. Hitherto the transmission of many acquired characteristics had seemed to most naturalists so obvious as not to call for demonstration.[65] Weismann's demand for facts in support of the main proposition revealed at once that none having real cogency could be produced. The time-honoured examples were easily shown to be capable of different explanations. A few certainly remain which cannot be so summarily dismissed, but--though it is manifestly impossible here to do justice to such a subject--I think no one will dispute that these residual and doubtful phenomena, whatever be their true nature, are not of a kind to help us much in the interpretation of any of those complex cases of adaptation which on the hypothesis of unguided Natural Selection are especially difficult to understand. Use and disuse were invoked expressly to help us over these hard places; but whatever changes can be induced in offspring by direct treatment of the parents, they are not of a kind to encourage hope of real assistance from that quarter. It is not to be denied that through the collapse of this second line of argument the Selection hypothesis has had to take an increased and perilous burden. Various ways of meeting the difficulty have been proposed, but these mostly resolve themselves into improbable attempts to expand or magnify the powers of Natural Selection.

Weismann's interpellation, though negative in purpose, has had a lasting and beneficial effect, for through his thorough demolition of the old loose and distracting notions of inherited experience, the ground has been cleared for the construction of a true knowledge of heredity based on experimental fact.

In another way he made a contribution of a more positive character, for his elaborate speculations as to the genetic meaning of cytological appearances have led to a minute investigation of the visible phenomena occurring in those cell-divisions by which germ-cells arise. Though the particular views he advocated have very largely proved incompatible with the observed facts of heredity, yet we must acknowledge that it was chiefly through the stimulus of Weismann's ideas that those advances in cytology were made; and though the doctrine of the continuity of germ-plasm cannot be maintained in the form originally propounded, it is in the main true and illuminating.[66] Nevertheless in the present state of knowledge we are still as a rule quite unable to connect cytological appearances with any genetic consequences and save in one respect (obviously of extreme importance--to be spoken of later) the two sets of phenomena might, for all we can see, be entirely distinct.

I cannot avoid attaching importance to this want of connection between the nuclear phenomena and the features of bodily organisation. All attempts to investigate Heredity by cytological means lie under the disadvantage that it is the nuclear changes which can alone be effectively observed. Important as they must surely be, I have never been persuaded that the rest of the cell counts for nothing. What we know of the behaviour and variability of chromosomes seems in my opinion quite incompatible with the belief that they alone govern form, and are the sole agents responsible in heredity.[67]

If, then, progress was to be made in Genetics, work of a different kind was required. To learn the laws of Heredity and Variation there is no other way than that which Darwin himself followed, the direct examination of the phenomena. A beginning could be made by collecting fortuitous observations of this class, which have often thrown a suggestive light, but such evidence can be at best but superficial and some more penetrating instrument of research is required. This can only be provided by actual experiments in breeding.

The truth of these general considerations was becoming gradually clear to many of us when in 1900 Mendel's work was rediscovered. Segregation, a phenomenon of the utmost novelty, was thus revealed. From that moment not only in the problem of the origin of species, but in all the great problems of biology a new era began. So unexpected was the discovery that many naturalists were convinced it was untrue, and at once proclaimed Mendel's conclusions as either altogether mistaken, or if true, of very limited application. Many fantastic notions about the workings of Heredity had been asserted as general principles before: this was probably only another fancy of the same class.

Nevertheless those who had a preliminary acquaintance with the facts of Variation were not wholly unprepared for some such revelation. The essential deduction from the discovery of segregation was that the characters of living things are dependent on the presence of definite elements or factors, which are treated as units in the processes of Heredity. These factors can thus be recombined in various ways. They act sometimes separately, and sometimes they interact in conduction with each other, producing their various effects. All this indicates a definiteness and specific order in heredity, and therefore in variation. This order cannot by the nature of the case be dependent on Natural Selection for its existence, but must be a consequence of the fundamental chemical and physical nature of living things. The study of Variation had from the first shown that an orderliness of this kind was present. The bodies and the properties of livings things are cosmic, not chaotic. No matter how low in the scale we go, never do we find the slightest hint of a diminution in that all-pervading orderliness, nor can we conceive an organism existing for a moment in any other state. Moreover not only does this order prevail in normal forms, but again and again it is to be seen in newly-sprung varieties, which by general consent cannot have been subjected to a prolonged Selection. The discovery of Mendelian elements admirably coincided with and at once gave a rationale of these facts. Genetic Variation is then primarily the consequence of additions to, or omissions from, the stock of elements which the species contains. The further investigation of the species-problem must thus proceed by the analytical method which breeding experiments provide.

In the nine years which have elapsed since Mendel's clue became generally known, progress has been rapid. We now understand the process by which a polymorphic race maintains its polymorphism. When a family consists of dissimilar members, given the numerical proportions in which these members are occurring, we can represent their composition symbolically and state what types can be transmitted by the various members. The difficulty of the "swamping effects of inter-crossing" is practically at an end. Even the famous puzzle of sex-limited inheritance is solved, at all events in its more regular manifestations, and we know now how it is brought about that the normal sisters of a colour-blind man can transmit the colour-blindness while his normal brothers cannot transmit it.

We are still only on the fringe of the inquiry. It can be seen extending and ramifying in many directions. To enumerate these here would be impossible. A whole new range of possibilities is being brought into view by study of the inter-relations between the simple factors. By following up the evidence as to segregation, indications have been obtained which can only be interpreted as meaning that when many factors are being simultaneously redistributed among the germ-cells, certain of them exert what must be described as a repulsion upon other factors. We cannot surmise whither this discovery may lead.

In the new light all the old problems wear a fresh aspect. Upon the question of the nature of Sex, for example, the bearing of Mendelian evidence is close. Elsewhere I have shown that from several sets of parallel experiments the conclusion is almost forced upon us that, in the types investigated, of the two sexes the female is to be regarded as heterozygous in sex, containing one unpaired dominant element, while the male is similarly homozygous in the absence of that element.[68] It is not a little remarkable that on this point--which is the only one where observations of the nuclear processes of gameto-genesis have yet been brought into relation with the visible characteristics of the organisms themselves--there should be diametrical opposition between the results of breeding experiments and those derived from cytology.

Those who have followed the researches of the American school will be aware that, after it had been found in certain insects that the spermatozoa were of two kinds according as they contained or did not contain the accessory chromosome, E. B. Wilson succeeded in proving that the sperms possessing this accessory body were destined to form _females_ on fertilisation, while sperms without it form males, the eggs being apparently indifferent. Perhaps the most striking of all this series of observations is that lately made by T. H. Morgan,[69] since confirmed by von Baehr, that in a Phylloxeran two kinds of spermatids are formed, respectively with and without an accessory (in this case, _double_) chromosome. Of these, only those possessing the accessory body become functional spermatozoa, the others degenerating. We have thus an elucidation of the puzzling fact that in these forms fertilisation results in the formation of _females_ only. How the males are formed--for of course males are eventually produced by the parthenogenetic females--we do not know.

If the accessory body is really to be regarded as bearing the factor for femaleness, then in Mendelian terms female is DD and male is DR. The eggs are indifferent and the spermatozoa are each male, _or_ female. But according to the evidence derived from a study of the sex-limited descent of certain features in other animals the conclusion seems equally clear that in them female must be regarded as DR and male as RR. The eggs are thus each either male or female and the spermatozoa are indifferent. How this contradictory evidence is to be reconciled we do not yet know. The breeding work concerns fowls, canaries, and the Currant moth (_Abraxas grossulariata_). The accessory chromosome has been now observed in most of the great divisions of insects,[70] except, as it happens, Lepidoptera. At first sight it seems difficult to suppose that a feature apparently so fundamental as sex should be differently constituted in different animals, but that seems at present the least improbable inference. I mention these two groups of facts as illustrating the nature and methods of modern genetic work. We must proceed by minute and specific analytical investigation. Wherever we look we find traces of the operation of precise and specific rules.

In the light of present knowledge it is evident that before we can attack the Species-problem with any hope of success there are vast arrears to be made up. He would be a bold man who would now assert that there was no sense in which the term Species might not have a strict and concrete meaning in contradistinction to the term Variety. We have been taught to regard the difference between species and variety as one of degree. I think it unlikely that this conclusion will bear the test of further research. To Darwin the question, What is a variation? presented no difficulties. Any difference between parent and offspring was a variation. Now we have to be more precise. First we must, as de Vries has shown, distinguish real, genetic, variation from _fluctuational_ variations, due to environmental and other accidents, which cannot be transmitted. Having excluded these sources of error the variations observed must be expressed in terms of the factors to which they are due before their significance can be understood. For example, numbers of the variations seen under domestication, and not a few witnessed in nature, are simply the consequence of some ingredient being in an unknown way omitted from the composition of the varying individual. The variation may on the contrary be due to the addition of some new element, but to prove that it is so is by no means an easy matter. Casual observation is useless, for though these latter variations will always be dominants, yet many dominant characteristics may arise from another cause, namely the meeting of complementary factors, and special study of each case in two generations at least is needed before these two phenomena can be distinguished.

When such considerations are fully appreciated it will be realised that medleys of most dissimilar occurrences are all confused together under the term Variation. One of the first objects of genetic analysis is to disentangle this mass of confusion.

To those who have made no study of heredity it sometimes appears that the question of the effect of conditions in causing variation is one which we should immediately investigate, but a little thought will show that before any critical inquiry into such possibilities can be attempted, a knowledge of the working of heredity under conditions as far as possible uniform must be obtained. At the time when Darwin was writing, if a plant brought into cultivation gave off an albino variety, such an event was without hesitation ascribed to the change of life. Now we see that albino _gametes_, germs, that is to say, which are destitute of the pigment-forming factor, may have been originally produced by individuals standing an indefinite number of generations back in the ancestry of the actual albino, and it is indeed almost certain that the variation to which the appearance of the albino is due cannot have taken place in a generation later than that of the grandparents. It is true that when a new _dominant_ appears we should feel greater confidence that we were witnessing the original variation, but such events are of extreme rarity, and no such case has come under the notice of an experimenter in modern times, as far as I am aware. That they must have appeared is clear enough. Nothing corresponding to the Brown-breasted Game fowl is known wild, yet that colour is a most definite dominant, and at some moment since _Gallus bankiva_ was domesticated, the element on which that special colour depends must have at least once been formed in the germ-cell of a fowl; but we need harder evidence than any which has yet been produced before we can declare that this novelty came through over-feeding, or change of climate, or any other disturbance consequent on domestication. When we reflect on the intricacies of genetic problems as we must now conceive them there come moments when we feel almost thankful that the Mendelian principles were unknown to Darwin. The time called for a bold pronouncement, and he made it, to our lasting profit and delight. With fuller knowledge we pass once more into a period of cautious expectation and reserve.

In every arduous enterprise it is pleasanter to look back at difficulties overcome than forward to those which still seem insurmountable, but in the next stage there is nothing to be stained by disguising the fact that the attributes of living things are not what we used to suppose. If they are more complex in the sense that the properties they display are throughout so regular[71] that the Selection of minute random variations is an unacceptable account of the origin of their diversity, yet by virtue of that very regularity the problem is limited in scope and thus simplified.

To begin with, we must relegate Selection to its proper place. Selection permits the viable to continue and decides that the non-viable shall perish; just as the temperature of our atmosphere decides that no liquid carbon shall be found on the face of the earth: but we do not suppose that the form of the diamond has been gradually achieved by a process of Selection. So again, as the course of descent branches in the successive generations, Selection determines along which branch Evolution shall proceed, but it does not decide what novelties that branch shall bring forth. "_La Nature contient le fonds de toutes ces variétés, mais le hazard ou l'art les mettent en oeuvre_," as Maupertuis most truly said.

Not till knowledge of the genetic properties of organisms has attained to far greater completeness can evolutionary speculations have more than a suggestive value. By genetic experiment, cytology and physiological chemistry aiding, we may hope to acquire such knowledge. In 1872 Nathusius wrote:[72] "Das Gesetz der Vererbung ist noch nicht erkannt; der Apfel ist noch nicht vom Baum der Erkenntniss gefallen, welcher, der Sage nach, Newton auf den rechten Weg zur Ergründung der Gravitationsgesetze führte." We cannot pretend that the words are not still true, but in Mendelian analysis the seeds of that apple-tree at last are sown.

If we were asked what discovery would do most to forward our inquiry, what one bit of knowledge would more than any other illuminate the problem, I think we may give the answer without hesitation. The greatest advance that we can foresee will be made when it is found possible to connect the geometrical phenomena of development with the chemical. The geometrical symmetry of living things is the key to a knowledge of their regularity, and the forces which cause it. In the symmetry of the dividing cell the basis of that resemblance we call Heredity is contained. To imitate the morphological phenomena of life we have to devise a system which can divide. It must be able to divide, and to segment as--grossly--a vibrating plate or rod does, or as an icicle can do as it becomes ribbed in a continuous stream of water; but with this distinction, that the distribution of chemical differences and properties must simultaneously be decided and disposed in orderly relation to the pattern of the segmentation. Even if a model which would do this could be constructed it might prove to be a useful beginning.

This may be looking too far ahead. If we had to choose some one piece of more proximate knowledge which we would more especially like to acquire, I suppose we should ask for the secret of interracial sterility. Nothing has yet been discovered to remove the grave difficulty, by which Huxley in particular was so much oppressed, that among the many varieties produced under domestication--which we all regard as analogous to the species seen in nature--no clear case of interracial sterility has been demonstrated. The phenomenon is probably the only one to which the domesticated products seem to afford no parallel. No solution of the difficulty can be offered which has positive value, but it is perhaps worth considering the facts in the light of modern ideas. It should be observed that we are not discussing incompatibility of two species to produce offspring (a totally distinct phenomenon), but the sterility of the offspring which many of them do produce.

When two species, both perfectly fertile severally, produce on crossing a sterile progeny, there is a presumption that the sterility is due to the development in the hybrid of some substance which can only be formed by the meeting of two complementary factors. That some such account is correct in essence may be inferred from the well-known observation that if the hybrid is not totally sterile but only partially so, and thus is able to form some good germ-cells which develop into new individuals, the sterility of these daughter-individuals is sensibly reduced or may be entirely absent. The fertility once re-established, the sterility does not return in the later progeny, a fact strongly suggestive of segregation. Now if the sterility of the cross-bred be really the consequence of the meeting of two complementary factors, we see that the phenomenon could only be produced among the divergent offspring of one species by the acquisition of at least _two_ new factors; for if the acquisition of a single factor caused sterility the line would then end. Moreover each factor must be separately acquired by distinct individuals, for if both were present together, the possessors would by hypothesis be sterile. And in order to imitate the case of species each of these factors must be acquired by distinct breeds. The factors need not, and probably would not, produce any other perceptible effects; they might, like the colour-factors present in white flowers, make no difference in the form or other characters. Not till the cross was actually made between the two complementary individuals would either factor come into play, and the effects even then might be unobserved until an attempt was made to breed from the cross-bred.

Next, if the factors responsible for sterility were acquired, they would in all probability be peculiar to certain individuals and would not readily be distributed to the whole breed. Any member of the breed also into which _both_ the factors were introduced would drop out of the pedigree by virtue of its sterility. Hence the evidence that the various domesticated breeds say of dogs or fowls can when mated together produce fertile offspring, is beside the mark. The real question is, Do they ever produce sterile offspring? I think the evidence is clearly that sometimes they do, oftener perhaps than is commonly supposed. These suggestions are quite amenable to experimental tests. The most obvious way to begin is to get a pair of parents which are known to have had any sterile offspring, and to find the proportions in which these steriles were produced. If, as I anticipate, these proportions are found to be definite, the rest is simple.