The Principles of Biology, Volume 1 (of 2)

"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 forehead, and the dark bars across the back part of the legs."¹⁰⁶

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 to the first husband? To these questions nothing like satisfactory replies seem forthcoming; and, in the absence of replies, scepticism, if not disbelief, may be held reasonable.

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 French and English breeds of sheep. The writer of the translated paper, M. Malingie-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 75 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 those 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 type in the ascendant… All the lambs produced strikingly resembled each other, and even Englishmen took them for animals of their own country."

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 in 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 obvious among most 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 obtained the volume, and there found what I expected. It was contained in a paper communicated by Dr. Wollaston from Daniel Giles, Esq., concerning his "sow and her produce," which said that —

"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 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 embryo, some of their germ-plasm passes into the mass of somatic cells constituting the parental body, and becomes a permanent component of it. Further, they necessitate the inference that this introduced germ-plasm, everywhere diffused, is some of it included in the reproductive cells subsequently formed. And if we thus get a demonstration that the somewhat different units of a foreign germ-plasm permeating the organism, permeate also the subsequently formed reproductive cells, and affect the structures of the individuals arising from them, the implication is that the like happens with those native units which have been made somewhat different by modified functions: there must be a tendency to inheritance of acquired characters.

One more step only has to be taken. It remains to ask what is the flaw in the assumption with which Weismann's theory sets out. If, as we see, the conclusions drawn from it do not correspond to the facts, then, either the reasoning is invalid, or the original postulate is untrue. Leaving aside all questions concerning the reasoning, it will suffice here to show the untruth of the postulate. Had his work been written during the early years of the cell-doctrine, the supposition that the multiplying cells of which the Metazoa and Metaphyta are composed, become completely separate, could not have been met by a reasonable scepticism; but now, not only is scepticism justifiable, but denial is called for. Some dozen years ago it was discovered that in many cases vegetal cells are connected with one another by threads of protoplasm – threads which unite the internal protoplasm of one cell with the internal protoplasms of cells around It is as though the pseudopodia of imprisoned rhizopods were fused with the pseudopodia of adjacent imprisoned rhizopods. We cannot reasonably suppose that the continuous network of protoplasm thus constituted has been produced after the cells have become adult. These protoplasmic connections must have survived the process of fission. The implication is that the cells forming the embryo-plant retained their protoplasmic connections while they multiplied, and that such connections continued throughout all subsequent multiplications – an implication which has, I believe, been established by researches upon germinating palm-seeds. But now we come to a verifying series of facts which the cell-structures of animals in their early stages present. In his Monograph of the Development of Peripatus Capensis, Mr. Adam Sedgwick, F.R.S., Reader in Animal Morphology at Cambridge, writes as follows: —

"All the cells of the ovum, ectodermal as well as endodermal, are connected together by a fine protoplasmic reticulum." (p. 41)

"The continuity of the various cells of the segmenting ovum is primary, and not secondary; i. e., in the cleavage the segments do not completely separate from one another. But are we justified in speaking of cells at all in this case? The fully segmented ovum is a syncytium, and there are not and have not been at any stage cell limits." (p. 41)

"It is becoming more and more clear every day that the cells composing the tissues of animals are not isolated units, but that they are connected with one another. I need only refer to the connection known to exist between connective tissue cells, cartilage cells, epithelial cells, &c. And not only may the cells of one tissue be continuous with each other, but they may also be continuous with the cells of other tissues." (pp. 47-8)

"Finally, if the protoplasm of the body is primitively a syncytium, and the ovum until maturity a part of that syncytium, the separation of the generative products does not differ essentially from the internal gemmation of a Protozoon, and the inheritance by the offspring of peculiarities first appearing in the parent, though not explained, is rendered less mysterious; for the protoplasm of the whole body being continuous, change in the molecular constitution of any part of it would naturally be expected to spread, in time, through the whole mass." (p. 49)

Mr. Sedgwick's subsequent investigations confirm these conclusions. In a letter of December 27, 1892, passages which he allows me to publish run as follows: —

"All the embryological studies that I have made since that to which you refer confirm me more and more in the view that the connections between the cells of adults are not secondary connections, but primary, dating from the time when the embryo was a unicellular structure… My own investigations on this subject have been confined to the Arthropoda, Elasmobranchii, and Aves. I have thoroughly examined the development of at least one kind of each of these groups, and I have never been able to detect a stage in which the cells were not continuous with each other; and I have studied innumerable stages from the beginning of cleavage onwards."

So that the alleged independence of the reproductive cells does not exist. The soma– to use Weismann's name for the aggregate of cells forming the body – is, in the words of Mr. Sedgwick, "a continuous mass of vacuolated protoplasm;" and the reproductive cells are nothing more than portions of it separated some little time before they are required to perform their functions.

Thus the theory of Weismann is doubly disproved. Inductively we are shown that there does take place that communication of characters from the somatic cells to the reproductive cells, which he says cannot take place; and deductively we are shown that this communication is a natural sequence of connections between the two which he ignores; his various conclusions are deduced from a postulate which is untrue.

* * * * *

From the title of this essay, and from much of its contents, nine readers out of ten will infer that it is directed against the views of Mr. Darwin. They will be astonished on being told that, contrariwise, it is directed against the views of those who, in a considerable measure, dissent from Mr. Darwin. For the inheritance of acquired characters, which it is now the fashion in the biological world to deny, was, by Mr. Darwin, fully recognized and often insisted on. Such of the foregoing arguments as touch Mr. Darwin's views, simply imply that the cause of evolution which at first he thought unimportant, but the importance of which he increasingly perceived as he grew older, is more important than he admitted, even at the last. The neo-Darwinists, however, do not admit this cause at all.

Let it not be supposed that this explanation implies any disapproval of the dissentients, considered as such. Seeing how little regard for authority I have myself usually shown, it would be absurd in me to reflect in any degree upon those who have rejected certain of Mr. Darwin's teachings, for reasons which they have held sufficient. But while their independence of thought is to be applauded rather than blamed, it is, I think, to be regretted that they have not guarded themselves against a long-standing bias. It is a common trait of human nature to seek some excuse when found in the wrong. Invaded self-esteem sets up a defence, and anything is made to serve. Thus it happened that when geologists and biologists, previously holding that all kinds of organisms arose by special creations, surrendered to the battery opened upon them by The Origin of Species, they sought to minimise their irrationality by pointing to irrationality on the other side. "Well, at any rate, Lamarck was in the wrong." "It is clear that we were right in rejecting his doctrine." And so, by duly emphasizing the fact that he overlooked "Natural Selection" as the chief cause, and by showing how erroneous were some of his interpretations, they succeeded in mitigating the sense of their own error. It is true their creed was that at successive periods in the Earth's history, old Floras and Faunas had been abolished and others introduced; just as though, to use Professor Huxley's figure, the table had been now and again kicked over and a new pack of cards brought out. And it is true that Lamarck, while he rejected this absurd creed, assigned for the facts reasons some of which are absurd. But in consequence of the feeling described, his defensible belief was forgotten and only his indefensible ones remembered. This one-sided estimate has become traditional; so that there is now often shown a subdued contempt for those who suppose that there can be any truth in the reasonings of a man whose general conception was partly sense, at a time when the general conceptions of his contemporaries were wholly nonsense. Hence results unfair treatment – hence result the different dealings with the views of Lamarck and of Weismann.

"Where are the facts proving the inheritance of acquired characters?" ask those who deny it. Well, in the first place, there might be asked the counter-question – Where are the facts which disprove it? Surely if not only the general structures of organisms, but also many of the modifications arising in them, are inheritable, the natural implication is that all modifications are inheritable; and if any say that the inheritableness is limited to those arising in a certain way, the onus lies on them of proving that those otherwise arising are not inheritable.¹⁰⁷ Leaving this counter-question aside, however, it will suffice if we ask another counter-question. It is asserted that the dwindling of organs from disuse is due to the successive survivals in posterity of individuals in which the organs have varied in the direction of decrease. Where now are the facts supporting this assertion? Not one has been assigned or can be assigned. Not a single case can be named in which panmixia is a proved cause of diminution. Even had the deductive argument for panmixia been as valid as we have found it to be invalid, there would still have been required, in pursuance of scientific method, some verifying inductive evidence. Yet, though not a shred of such evidence has been given, the doctrine is accepted with acclamation, and adopted as part of current biological theory. Articles are written and letters published in which it is assumed that this mere speculation, justified by not a tittle of proof, displaces large conclusions previously drawn. And then, passing into the outer world, this unsupported belief affects opinions there too; so that we have recently had a Right Honourable lecturer who, taking for granted its truth, represents the inheritance of acquired characters as an exploded hypothesis, and proceeds to give revised views of human affairs.

Finally, there comes the reply that there are facts proving the inheritance of acquired characters. All those assigned by Mr. Darwin, together with others such, remain outstanding when we find that the interpretation by panmixia is untenable. Indeed, even had that hypothesis been tenable, it would have been inapplicable to these cases; since in domestic animals, artificially fed and often overfed, the supposed advantage from economy cannot be shown to tell; and since, in these cases, individuals are not naturally selected during the struggle for life, in which certain traits are advantageous, but are artificially selected by man without regard to such traits. Should it be urged that the assigned facts are not numerous, it may be replied that there are no persons whose occupations and amusements incidentally bring out such facts; and that they are probably as numerous as those which would have been available for Mr. Darwin's hypothesis, had there been no breeders and fanciers and gardeners who, in pursuit of their profits and hobbies, furnished him with evidence. It may be added that the required facts are not likely to be numerous, if biologists refuse to seek for them.

See, then, how the case stands. Natural selection, or survival of the fittest, is almost exclusively operative throughout the vegetal world and throughout the lower animal world, characterized by relative passivity. But with the ascent to higher types of animals, its effects are in increasing degrees involved with those produced by inheritance of acquired characters; until, in animals of complex structures, inheritance of acquired characters becomes an important, if not the chief, cause of evolution. We have seen that natural selection cannot work any changes in organisms save such as conduce in considerable degrees, directly or indirectly, to the multiplication of the stirp; whence failure to account for various changes ascribed to it. And we have seen that it yields no explanation of the co-adaptation of co-operative parts, even when the co-operation is relatively simple, and still less when it is complex. On the other hand, we see that if, along with the transmission of generic and specific structures, there tend to be transmitted modifications arising in a certain way, there is a strong a priori probability that there tend to be transmitted modifications arising in all ways. We have a number of facts confirming this inference, and showing that acquired characters are inherited – as large a number as can be expected, considering the difficulty of observing them and the absence of search. And then to these facts may be added the facts with which this essay set out, concerning the distribution of tactual discriminativeness. While we saw that these are inexplicable by survival of the fittest, we saw that they are clearly explicable as resulting from the inheritance of acquired characters. And here let it be added that this conclusion is conspicuously warranted by one of the methods of inductive logic, known as the method of concomitant variations. For throughout the whole series of gradations in perceptive power, we saw that the amount of the effect is proportionate to the amount of the alleged cause.

II

Apart from those more special theories of Professor Weismann I lately dealt with, the wide acceptance of which by the biological world greatly surprises me, there are certain more general theories of his – fundamental theories – the acceptance of which surprises me still more. Of the two on which rests the vast superstructure of his speculations, the first concerns the distinction between the reproductive elements of each organism and the non-reproductive elements. He says: —

"Let us now consider how it happened that the multicellular animals and plants, which arose from unicellular forms of life, came to lose this power of living for ever.

"The answer to this question is closely bound up with the principle of division of labour which appeared among multicellular organisms at a very early stage…

"The first multicellular organism was probably a cluster of similar cells, but these units soon lost their original homogeneity. As the result of mere relative position, some of the cells were especially fitted to provide for the nutrition of the colony, while others undertook the work of reproduction." (Essays upon Heredity, i, p. 27)

Here, then, we have the great principle of the division of labour, which is the principle of all organization, taken as primarily illustrated in the division between the reproductive cells and the non-reproductive or somatic cells – the cells devoted to the continuance of the species, and the cells which subserve the life of the individual. And the early separation of reproductive cells from somatic cells, is alleged on the ground that this primary division of labour is that which arises between elements devoted to species-life and elements devoted to individual life. Let us not be content with words but look at the facts.

When Milne-Edwards first used the phrase "physiological division of labour," he was obviously led to do so by perceiving the analogy between the division of labour in a society, as described by political economists, and the division of labour in an organism. Every one who reads has been familiarized with the first as illustrated in the early stages, when men were warriors while the cultivation and drudgery were done by slaves and women; and as illustrated in the later stages, when not only are agriculture and manufactures carried on by separate classes, but agriculture is carried on by landlords, farmers, and labourers, while manufactures, multitudinous in their kinds, severally involve the actions of capitalists, overseers, workers, &c., and while the great function of distribution is carried on by wholesale and retail dealers in different commodities. Meanwhile students of biology, led by Milne-Edwards's phrase, have come to recognize a parallel arrangement in a living creature; shown, primarily, in the devoting of the outer parts to the general business of obtaining food and escaping from enemies, while the inner parts are devoted to the utilization of food, and supporting themselves and the outer parts; and shown, secondarily, by the subdivision of these great functions into those of various limbs and senses in the one case, and in the other case into those of organs for digestion, respiration, circulation, excretion, &c. But now let us ask what is the essential nature of this division of labour. In both cases it is an exchange of services– an arrangement under which, while one part devotes itself to one kind of action and yields benefits to all the rest, all the rest, jointly and severally performing their special actions, yield benefits to it in exchange. Otherwise described, it is a system of mutual dependence: A depends for its welfare upon B, C, and D; B upon A, C, and D; and so with the rest: all depend upon each and each upon all. Now let us apply this true conception of the division of labour, to that which Professor Weismann calls a division of labour. Where is the exchange of services between somatic cells and reproductive cells? There is none. The somatic cells render great services to the reproductive cells, by furnishing them with materials for growth and multiplication; but the reproductive cells render no services at all to the somatic cells. If we look for the mutual dependence we look in vain. We find entire dependence on the one side and none on the other. Between the parts devoted to individual life and the part devoted to species-life, there is no division of labour whatever. The individual works for the species; but the species works not for the individual. Whether at the stage when the species is represented by reproductive cells, or at the stage when it is represented by eggs, or at the stage when it is represented by young, the parent does everything for it, and it does nothing for the parent. The essential part of the conception is gone: there is no giving and receiving, no exchange, no mutuality.

But now suppose we pass over this fallacious interpretation, and grant Professor Weismann his fundamental assumption and his fundamental corollary. Suppose we grant that because the primary division of labour is that between somatic cells and reproductive cells, these two groups are the first to be differentiated. Having granted this corollary, let us compare it with the facts. As the alleged primary division of labour is universal, so the alleged primary differentiation should be universal too. Let us see whether it is so. Already, in the paragraph from which I have quoted above, a crack in the doctrine is admitted: it is said that "this differentiation was not at first absolute, and indeed it is not always so to-day." And then, on turning to page 74, we find that the crack has become a chasm. Of the reproductive cells it is stated that – "In Vertebrata they do not become distinct from the other cells of the body until the embryo is completely formed." That is to say, in this large and most important division of the animal kingdom, the implied universal law does not hold. Much more than this is confessed. Lower down the page we read – "There may be in fact cases in which such separation does not take place until after the animal is completely formed, and others, as I believe that I have shown, in which it first arises one or more generations later, viz., in the buds produced by the parent."