ON TELEGONY. A widely different view, however, is taken by Mr. Herbert Spencer with regard to the theoretical interpretation of telegony. This, indeed, is precisely the opposite view to the one which is given in the text. For while I agree with Professor Weismann in holding that the facts of telegony (supposing them to be facts) are as compatible with the theory of germ-plasm as with that of gemmules, “physiological units,” or any other theory which postulates a centripetal flow of the carriers of heredity from somatic-cells to germ-cells, Mr. Spencer is of the opinion that these facts are destructive of any theory which postulates a continuity in the substance of heredity—i.e., a centrifugal flow of the carriers of heredity. And, unquestionably, Mr. Spencer’s view is the prevalent one. Therefore, seeing that his opinion is not only of weight per se, but is shared by the scientific world in general, I will here transcribe a somewhat lengthy discussion which I have recently held with him upon the subject. In the Contemporary Review for March, Mr. Spencer wrote as follows:— We pass now to evidence not much known in the world at large, but widely known in the biological world, though known in so incomplete a manner as to be undervalued in it. Indeed, when I name it probably many will vent a mental pooh-pooh. The fact to which I refer is one of which record is preserved I tried to breed from the male quagga and a young chestnut mare of seven-eighths Arabian blood, and which had never been bred from; the result was the production of a female hybrid, now five years old, and bearing, both in her form and in her colour, very decided indications of her mixed origin. I subsequently parted with the seven-eighths Arabian mare to Sir Gore Ouseley, who has bred from her by a very fine black Arabian horse. I yesterday morning examined the produce, namely, a two-year-old filly and a year-old colt. They have the character of the Arabian breed as decidedly as can be expected, where fifteen-sixteenths of the blood are Arabian; and they are fine specimens of that breed; but both in their colour and in the hair of their manes they have a striking resemblance to the quagga. Their colour is bay, marked more or less like the quagga in a darker tint. Both are distinguished by the dark line along the ridge of the back, the dark stripes across the fore-hand, and the dark bars across the back part of the legs Lord Morton then names sundry further correspondences. Dr. Wollaston, at that time President of the Royal Society, who had seen the animals, testified to the correctness of his description, and, as shown by his remarks, entertained no doubt about the alleged facts. But good reason for doubt may be assigned. There naturally arises the question—How does it happen that parallel results are not observed in other cases? If in any progeny certain traits not belonging to the sire, but belonging to a sire of preceding progeny, are reproduced, how is it that such anomalously-inherited traits are not observed in domestic animals, and indeed in mankind? How is it that the children of a widow by a second husband do not bear traceable resemblances of the first husband? To these questions nothing like satisfactory replies seem forthcoming; There is an explanation, however. Forty years ago I made acquaintance with a fact which impressed me by its significant implications; and has, for this reason I suppose, remained in my memory. It is set forth in the Journal of the Royal Agricultural Society, vol. xiv. (1853), pp. 214 et seq., and concerns certain results of crossing English and French breeds of sheep. The writer of the translated paper, M. MalingiÉ-Nouel, Director of the Agricultural School of La Charmoise, states that when the French breeds of sheep (in which were included “the mongrel Merinos”) were crossed with an English breed, “the lambs present the following results. Most of them resemble the mother more than the father; some show no trace of the father.” Joining the admission respecting the mongrels with the facts subsequently stated, it is tolerably clear that the cases in which the lambs bore no traces of the father were cases in which the mother was of pure breed. Speaking of the results of these crossings in the second generation “having seventy-five per cent. of English blood,” M. Nouel says:—“The lambs thrive, wear a beautiful appearance, and complete the joy of the breeder.... No sooner are the lambs weaned than their strength, their vigour, and their beauty begin to decay.... At last the constitution gives way ... he remains stunted for life”: the constitution being thus proved unstable or unadapted to the requirements. How, then, did M. Nouel succeed in obtaining a desirable combination of a fine English breed with the relatively poor French breeds? He took an animal from “flocks originally sprung from a mixture of the two distinct races that are established in these two provinces [Berry and La Sologne],” and these he “united with animals of another mixed breed, ... which blended the Tourangelle and native Merino blood of” La Beauce and Touraine, and obtained a mixture of all four races “without decided character, without fixity, ... but possessing the advantage of being used to our climate and management.” Putting one of these “mixed-blood ewes to a pure New-Kent ram ... one obtains a lamb containing fifty-hundredths of the purest and most ancient English blood, with twelve and a-half hundredths of four different French races, which are individually lost in the preponderance of English blood, and disappear almost entirely, leaving the improving M. Nouel goes on to remark that when this derived breed was bred with itself, the marks of the French breeds were lost. “Some slight traces could be detected by experts, but these soon disappeared.” Thus we get proof that relatively pure constitutions predominate in progeny over much mixed constitutions. The reason is not difficult to see. Every organism tends to become adapted to its conditions of life; and all the structures of a species, accustomed through multitudinous generations to the climate, food, and various influences of its locality, are moulded into harmonious co-operation favourable to life in that locality: the result being that in the development of each young individual, the tendencies conspire to produce the fit organization. It is otherwise when the species is removed to a habitat of different character, or when it is of mixed breed. In the one case its organs, partially out of harmony with the requirements of its new life, become partially out of harmony with one another; since, while one influence, say of climate, is but little changed, another influence, say of food, is much changed; and, consequently, the perturbed relations of the organs interfere with their original stable equilibrium. Still more in the other case is there a disturbance of equilibrium. In a mongrel the constitution derived from each source repeats itself as far as possible. Hence a conflict of tendencies to evolve two structures more or less unlike. The tendencies do not harmoniously conspire; but produce partially incongruous sets of organs. And evidently where the breed is one in which there are united the traits of various lines of ancestry, there results an organization so full of small incongruities of structure and action, that it has a much-diminished power of maintaining its balance; and while it cannot withstand so well adverse influences, it cannot so well hold its own in the offspring. Concerning parents of pure and mixed breeds respectively, severally tending to reproduce their own structures in progeny, we may therefore say, figuratively, that the house divided against itself cannot withstand the house of which the members are in concord. Now if this is shown to be the case with breeds the purest of which have been adapted to their habitats and modes of life during some few hundred years only, what shall we say when the question is of a breed which has had a constant mode of life in the same locality for ten thousand years or more, like the quagga? In this the stability of constitution must be such as no domestic animal can approach. Relatively stable as may have been the constitutions of Lord Morton’s horses, as compared with the constitutions of ordinary horses, yet, since Arab horses, even in their native country, have probably in the course of successive conquests and migrations of tribes become more or less mixed, and since they have been subject to the conditions of domestic life, differing much from the conditions of their original wild life, and since the English breed has undergone the perturbing effects of change from the climate and food of the East to the climate and food of the West, the organizations of the horse and mare in question could have had nothing like that perfect balance produced in the quagga by a hundred centuries of harmonious co-operation. Hence the result. And hence at the same time the interpretation of the fact that analogous phenomena are not perceived among domestic animals, or among ourselves; since both have relatively mixed, and generally extremely mixed, constitutions, which, as we see in ourselves, have been made generation after generation, not by the formation of a mean between two parents, but by the jumbling of traits of the one with traits of the other, until there exist no such conspiring tendencies among the parts as cause repetition of combined details of structure in posterity. Expectation that scepticism might be felt respecting this alleged anomaly presented by the quagga-marked foal, had led me to think over the matter; and I had reached this interpretation before sending to the College of Surgeons Museum (being unable to go myself) to obtain the particulars and refer to the records. When there was brought to me a copy of the account as set forth in the “Philosophical Transactions,” it was joined with the information that there existed an appended account of pigs, in which a parallel fact had been observed. To my immediate inquiry—“Was the male a wild pig?”—there came the reply: “I did not observe.” Of course I forthwith she was one of a well-known black and white breed of Mr. Western, the Member for Essex. About ten years since I put her to a boar of the wild breed, and of a deep chestnut colour, which I had just received from Hatfield House, and which was soon afterwards drowned by accident. The pigs produced (which were her first litter) partook in appearance of both boar and sow, but in some the chestnut colour of the boar strongly prevailed. The sow was afterwards put to a boar of Mr. Western’s breed (the wild boar having been long dead). The produce was a litter of pigs some of which, we observed with much surprise, to be stained and clearly marked with the chestnut colour which had prevailed in the former litter. Mr. Giles adds that in a second litter of pigs, the father of which was of Mr. Western’s breed, he and his bailiff believe there was a recurrence, in some, of the chestnut colour, but admits that their “recollection is much less perfect than I wish it to be.” He also adds that, in the course of many years’ experience, he had never known the least appearance of the chestnut colour in Mr. Western’s breed. What are the probabilities that these two anomalous results should have arisen, under these exceptional conditions, as a matter of chance? Evidently the probabilities against such a coincidence are enormous. The testimony is in both cases so good that, even apart from the coincidence, it would be unreasonable to reject it; but the coincidence makes acceptance of it imperative. There is mutual verification, at the same time that there is a joint interpretation yielded of the strange phenomenon, and of its non-occurrence under ordinary circumstances. And now, in the presence of these facts, what are we to say? Simply that they are fatal to Weismann’s hypothesis. They show that there is none of the alleged independence of the reproductive cells; but that the two sets of cells are in close communion. They prove that while the reproductive cells multiply and arrange themselves during the evolution of the My reply to this appeared in the April issue of the Contemporary Review, as follows:— Influence on Progeny of a Previous Sire. This is the last of the arguments which Mr. Spencer advances against the position of Professor Weismann. Alluding to the case of Lord Morton’s mare, he represents that the phenomenon which it serves so well to illustrate—viz., the influence of a previous sire on the progeny of another by the same dam—is hopelessly at variance with the theory of germ-plasm. I cannot quite gather the explanation which he would give of this phenomenon, further than that in some way or another it betokens an immediate influence of the hereditary material of the male on the body-tissues (“somatic cells”) of the female. And this is the view which is taken of the phenomenon by the Lamarckians in general. Yet, if we consider all that such an explanation involves, we shall find that it is a highly complex explanation, for it involves the following chain of hypotheses:—The first impregnation affects many, if not all, the somatic tissues of the mother by the germinal matter of the father; these tissues, in their turn, react on the maturing ova; this action and reaction is such that when one of the ova is afterwards fertilized by a different sire, the resulting offspring more Thirdly, is it not literally inconceivable that this Neo-Lamarckian explanation can be the true one? Can it be seriously contemplated that there is any such mechanism as the explanation must needs assume? If it is difficult to accept such a machinery as is supposed by the theory of pangenesis, whereby every cell in the body casts off “gemmules,” which are the carriers of heredity from their respective tissues to the germinal elements, what are we to say of such a machinery as the following:—A machinery which distributes through the body of a female gemmules from the disintegrated spermatozoa of her mate; which distributes them selectively, so that they shall all eventually lodge in those tissue-cells of the female which Such, it seems to me, is a fair statement of the whole case up to date. But I think it may be apposite now to publish the main results of an inquiry on which I have been engaged for the last three years. First as to the facts. The investigations have been pursued on three different lines: (1) I raised discussions on the subject in the principal breeders’ and fanciers’ journals of this country, and also of America. (2) I entered into private correspondence with contributors of the largest experience, and also with professional and amateur breeders, fanciers, &c., who addressed me directly on the subject. (3) I started experiments with the varieties which these inquiries indicated as most likely to yield positive results. At present nothing need be said with regard to these experiments, because they are not sufficiently matured. But it is desirable to state the general upshot of the correspondence. The principal result is to show that the phenomenon is of much less frequent occurrence than is generally supposed. Indeed, it is so rare that I doubt whether it takes place in more than one or two per cent. of cases. I must add, however, that nearly all my professional correspondents would deem this an absurdly low estimate. Most of them are quite persuaded that it is of frequent occurrence, many of them regard it as a general rule, while some of them go so far as to make a point of always putting a mare, a bitch, &c. to a good pedigree male in her first season, so that her subsequent progenies may be benefited by his influence, even though they be engendered by inferior sires. But I am certain that these estimates must be largely discounted in view of merely accidental resemblances, and still more on account of the prevalent belief upon the subject, which, where unquestioningly entertained, prevents anything like a critical estimate being formed. But that the phenomenon does occur in some small percentage of cases there can be no reasonable doubt—as a result, I mean, of analysing the hundreds of cases which have now been submitted to me, especially with regard to dogs. One thoroughly well observed case occurring among pedigree animals is worth any number of slipshod statements, when precedent belief, inefficient isolation, exaggeration of memory, and so forth, have to be allowed for. On the present occasion space does not admit of giving such special instances, so I must ask it to be taken for granted that my evidence is enough to prove the fact of a previous sire asserting his influence on a subsequent progeny, although this fact is one of comparatively rare occurrence. It may be added that I have failed to find any good evidence of its ever occurring at all in the case of man. For although I have met with an alleged instance of a white woman, who, after having borne children to a negro husband, had a second family to a white one, in which some negro characteristics appeared, I have not been able to meet with any corroboration of this instance. I have made inquiries among medical men in the Southern States of America, where in the days of slavery it was frequently the custom that young negresses should bear their first children to their masters, and their subsequent children to negro husbands; but it never seems to have been observed, according to my correspondents, that these subsequent children were other than pure negroes. Such, however, was not the same case as the one above mentioned, but a reciprocal case; and this may have made a difference. If any reader should happen to know of another instance where a negro was the first husband, I hope he will inform me as to the result. It has hitherto puzzled me why the phenomenon in question, since it does certainly occur in some cases, should occur so rarely as the above inquiries prove. But I think that Mr. Spencer’s suggestion on this point is a valuable one, as it seems to present an excellent promise of solving the puzzle. This suggestion, it will be remembered, is that when the first sire is of a relatively stable and also of a markedly different ancestral stock from the dam—e.g., of a different species, as in the case of Lord Morton’s mare—there will be most likelihood of his impressing his ancestral characters on the progeny of the So much, then, for the facts. As regards their interpretation, it certainly seems to me that the one which I have supposed to be given by Weismann is less difficult of acceptance than the one which is given by the Lamarckians, as we have seen above. But it also seems to me that the latter explanation is not the only one available under the Lamarckian hypothesis. For, even under this hypothesis, there is no need to assume that the influence of the first sire is exerted on all the somatic tissues of the mother, and that these again reflect this influence on the ovum which is afterwards fertilized by the second sire. A mechanism that could effect all this may well be deemed impossible. But a much simpler explanation can be furnished by the Neo-Lamarckians, on lines similar to those upon which I have supposed that Weismann’s explanation would run. For, on their common supposition that the substance of heredity is particulate, it matters not in the present connexion whether we suppose the particles to be ids or gemmules. Indeed, it is more in accordance with the hypothetical endowments of the latter than of the former, that they should be capable of penetrating the coats of an ovum, if they can survive the disintegration of their containing spermatozoÖn. Nevertheless, thus far it does not seem to me that any theory belonging to the family of pangenesis can gain any advantage over the theory of germ-plasm, The advantage which any theory of gemmules seeks to gain over the theory of germ-plasm by an appeal to the fact in question, consists in supposing that the influence of the previous sire is exercised in the first instance on the somatic cells of the female. For this would prove that the germinal elements of the male are capable of communicating their hereditary qualities, not only by mixing with the germinal elements of the female (as in ordinary fertilization) but also by direct contact with the general tissues of the female. And this again would prove that the fundamental postulate of the theory of germ-plasm is erroneous—i.e., the postulate of the continuity of germ-plasm, or of its perpetual restriction to a “sphere” of its own. This, as all who are acquainted with the literature of the subject will at once perceive, would be a serious blow to the whole Weismannian system. But, as we have seen, the current Lamarckian interpretation of the fact in question involves the supposition of a physiological machinery so inconceivably complex that instead of serving to corroborate the theory of gemmules (or of physiological units) it would go to render that theory incredible If, however, we turn to plants, we find a considerable number of facts which unquestionably demonstrate the only point which this interpretation has been adduced to suggest. For these facts show that, in not a few cases, the germinal matter of pollen-grains is capable of asserting its influence beyond the ovules to the somatic tissues of the ovary, and even to the flower-stalk of the mother plant. Here, then, we have simple and conclusive evidence of the material of heredity exercising a direct influence on somatic tissues. How this well-known fact is to be met by the theory of germ-plasm is a question which does not seem to have thus far engaged the attention of Professor Weismann, or of any of his followers. For particulars touching this phenomenon, so highly important in its relation to the theory of germ-plasm, I cannot do better than refer to the eleventh chapter of Darwin’s work on the “Variation of Animals and Plants under Domestication.” Again, in the Contemporary Review for May, Mr. Spencer wrote:— In the essay to which this is a postscript, conclusions were drawn from the remarkable case of the horse and quagga there narrated, along with an analogous case observed among pigs. These conclusions have since been confirmed. I am much indebted to a distinguished correspondent who has drawn my attention to verifying facts furnished by the offspring of whites and negroes in the United States. Referring to information given him many years ago, he says:—“It was to the effect that the children of white women by a white father had been repeatedly observed to show traces of black blood, in cases when the woman had previous connexion with [i. e., a child by] a negro.” At the time I received this information, an American was visiting me; and, on being appealed to, answered that in the United States there was an established belief to this effect. Not wishing, however, to depend upon hearsay, I at once wrote to America to make inquiries. Professor Cope of Philadelphia has written to friends in the South, but has not yet sent me the That cases of the kind should not be frequently seen in the North, especially nowadays, is of course to be expected. The first of the above quotations refers to facts observed in the South during slavery days; and, even then, the implied conditions were naturally very infrequent. Dr. W. J. Youmans of New York has, on my behalf, interviewed several medical professors, who, though they have not themselves met with instances, say that the alleged result, described above, “is generally accepted as a fact.” But he gives me what I think must be regarded as authoritative testimony. It is a quotation from the standard work of Professor Austin Flint, and runs as follows:— A peculiar and, it seems to me, an inexplicable fact is, that previous pregnancies have an influence upon offspring. This is well known to breeders of animals. If pine-blooded mares or bitches have been once covered by an inferior male, in subsequent fecundations the young are likely to partake of the character of the first male, even if they be afterwards bred with males of unimpeachable pedigree. What the mechanism of the influence of the first conception is, it is impossible to say; but the fact is incontestable. The same influence is observed in the human subject. A woman may have, by a second husband, children who resemble a former husband, and this is particularly well marked in certain instances by the colour of the hair and eyes. A white woman who has had children by a negro may subsequently bear children to a white man, these children presenting some of the unmistakable peculiarities of the negro race Dr. Youmans called on Professor Flint, who remembered “investigating the subject at the time his larger work was written [the above is from an abridgment], and said that he had never heard the statement questioned.” Some days before I received this letter and its contained It is a well-known and admitted fact that if a bitch has two litters by two different dogs, the character of the first father is sure to be perpetuated in any litters she may afterwards have, no matter how pure-bred a dog may be the begetter. After citing this testimony, Mr. Fookes goes on to give illustrations known to himself. A friend of mine near this had a very valuable Dachshund bitch, which most unfortunately had a litter by a stray sheep-dog. The next year her owner sent her on a visit to a pure Dachshund dog, but the produce took quite as much of the first father as the second, and the next year he sent her to another Dachshund with the same result. Another case:—A friend of mine in Devizes had a litter of puppies, unsought for, by a setter from a favourite pointer bitch, and after this she never bred any true pointers, no matter of what the paternity was. These further evidences, to which Mr. Fookes has since added others, render the general conclusion incontestable. Coming from remote places, from those who have no theory to support, and who are some of them astonished by the unexpected phenomena, the agreement dissipates all doubt. In four kinds of mammals, widely divergent in their natures—man, horse, dog, and pig—we have this same seemingly anomalous kind of heredity made visible under analogous conditions. We must take it as a demonstrated fact that, during gestation, traits of constitution inherited from the father produce effects upon the constitution of the mother; and that these communicated effects are transmitted by her to subsequent offspring. We are supplied with an absolute disproof of Professor Weismann’s doctrine that the reproductive cells are independent of, and uninfluenced by, the somatic cells; and there disappears absolutely the alleged obstacle to the transmission of acquired characters.... There is one other passage in Dr. Romanes’ criticism—that concerning the influence of a previous sire on progeny—which calls for comment. He sets down what he supposes Weismann will say in response to my argument. “First, he may question the fact.” Well, after the additional evidence given above, I think he is not likely to do that; unless, indeed, it be that along with readiness to base conclusions on things “it is easy to imagine” there goes reluctance to accept testimony which it is difficult to doubt. Second, he is supposed to reply that “the germ-plasm of the first sire has in some way or another become partly commingled with that of the immature ova”; and Dr. Romanes goes on to describe how there may be millions of spermatozoa and “thousands of millions” of their contained “ids” around the ovaries, to which these secondary effects are due. But, on the one hand, he does not explain why in such case each subsequent ovum, as it becomes matured, is not fertilized by the sperm-cells present, or their contained germ-plasm, rendering all subsequent fecundations needless; and, on the other hand, he does not explain why, if this does not happen, the potency of this remaining germ-plasm is nevertheless such as to affect not only the next succeeding offspring, but all subsequent offspring. The irreconcilability of these two implications would, I think, sufficiently dispose of the supposition, even had we not daily multitudinous proof that the surface of a mammalian ovarium is not a sperm-atheca. The third difficulty Dr. Romanes urges is the inconceivability of the process by which the germ-plasm of a preceding male parent affects the constitution of the female and her subsequent offspring. In response, I have to ask why he piles up a mountain of difficulties based on the assumption that Mr. Darwin’s explanation of heredity by “Pangenesis” is the only available explanation preceding that of Weismann? and why he presents these difficulties to me more especially, deliberately ignoring my own hypothesis of physiological units? It cannot be that he is ignorant of this hypothesis, since the work in which it is variously set forth (“Principles of Biology,” §§ 66-97) is one with which he is well acquainted: witness his “Scientific Evidences of Organic Evolution”; and he has had recent reminders of it in Weismann’s “Germ-plasm,” where it is To this I responded, in the Contemporary Review for June:— With regard to the influence of a previous sire, I ventured in my article to show that, even supposing it to be a fact, the phenomena concerned would not constitute any valid evidence against Weismann’s theory of germ-plasm, and, of course, still less would “they prove that while the reproductive cells multiply and arrange themselves during the evolution of the embryo, some of their germ-plasm passes into the mass of somatic cells constituting the parental body, and becomes a permanent component of it,” with the result that the phenomena in question “are simply fatal to Weismann’s hypothesis.” For a much simpler and more probable explanation is to be found in supposing that the unused germ-plasm of the first sire may survive the disintegration of its containing spermatozoa in the Fallopian tubes of the female, and thus gain access to the hitherto unripe ova directly, instead of first having to affect the whole maternal organism, and then being reflected from it to them. I showed, at some length, how immensely complex the mechanism of any such process would necessarily have to be; and for the purposes of exposition I employed the terminology of Darwin’s theory of Pangenesis. Mr. Spencer now says: “In response, I have to ask why he [I] piles up a mountain of difficulties based on the assumption that Mr. Darwin’s explanation of heredity by ‘Pangenesis’ is the only available explanation preceding that of Weismann? and why he presents these difficulties to me more expecially, deliberately ignoring my own hypothesis of physiological units?” Now my answer to this is very simple. I do not hold a brief for Weismann. On the contrary, I am in large measure an opponent of his In conclusion, I would like to take this opportunity of explaining that my former article was written in Madeira, where I did not receive a copy of Weismann’s most recent work, entitled The Germ-plasm, until the Contemporary Review for April was being printed off. Thus, I was not then aware that in this work Professor Weismann had fully anticipated several Lastly, in the same issue of the Contemporary Review, Mr. Spencer explained:— Mr. Darwin’s hypothesis of Pangenesis implies not only that the reproductive cell must contain numerous kinds of gemmules derived from different organs, but that the numbers of these gemmules must bear to one another something like the proportions which the originating organs bear to one another in size. The conception involves many different kinds, whose numbers are in many different proportions, and I supposed the difficulty alleged was, that for the influence of a previous sire to be communicated from the growing foetus to the mother would imply not only the transfer of the various kinds of gemmules derived from him, but also maintenance of their numerical proportions, and that again these gemmules, diffused throughout the maternal system, would have to be transferred in these proportions to the subsequently formed ova. No such difficulties arise if the units conveying hereditary characters are of one kind only. From this it is apparent that Mr. Spencer has misunderstood “the difficulty alleged,” and that the desired explanation is not yet forthcoming. I did not say anything about “kinds” or “proportions” of the carriers of heredity; my difficulty is to conceive of any mechanism whereby these carriers can first directly influence the somatic-cells of the mother, and then indirectly reflect this influence upon her germ-cells. Also, I cannot see any obvious necessity for the intervention of the “embryo” in the process.
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