Chapter XLVII: Appendix: B (3)

And had Professor Huxley taken the actual weight, one-twelfth of a grain, the quintillion of _Aphides_ would evidently far outweigh the whole human population of the globe: five billions of tons being the weight, as brought out by my own calculation! Of course I do not cite this in proof of the extent to which multiplication of somatic cells, descending from a single ovum, may go; because it will be contended, with some reason, that each of the sexless _Aphides_, viviparously produced, arose by fission of a cell which had descended from the original reproductive cell. I cite it merely to show that when the cell-products of a fertilized ovum are perpetually divided and subdivided into small groups, distributed over an unlimited nutritive area, so that they can get materials for growth at no cost, and expend nothing appreciable in motion or maintenance of temperature, cell-production may go on without limit. For the agamic multiplication of _Aphides_ has been shown to continue for four years, and to all appearance would be ceaseless were the temperature and supply of food continued without break. But now let us pass to analogous illustrations of cause and consequence, open to no criticism of the kind just indicated. They are furnished by various kinds of _Entozoa_, of which take the _Trematoda_, infesting molluscs and fishes. Of one of them we read:--"_Gyrodactylus_ multiplies agamically by the development of a young Trematode within the body, as a sort of internal bud. A second generation appears within the first, and even a third within the second, before the young _Gyrodactylus_ is born."[112] And the drawings of Steenstrup, in his _Alternation of Generations_, show us, among creatures of this group, a sexless individual the whole interior of which is transformed into smaller sexless individuals, which severally, before or after their emergence, undergo similar transformations--a multiplication of somatic cells without any sign of reproductive cells. Under what circumstances do such modes of agamic multiplication, variously modified among parasites, occur? They occur where there is no expenditure whatever in motion or maintenance of temperature, and where nutriment surrounds the body on all sides. Other instances are furnished by groups in which, though the nutriment is not abundant, the cost of living is almost unappreciable. Among the _Coelenterata_ there are the Hydroid Polyps, simple and compound; and among the _Mollusca_ we have various types of Ascidians, fixed and floating, _Botryllidæ_ and _Salpæ_.

But now from these low animals in which sexless reproduction, and continued multiplication of somatic cells, is common, and one class of which is named "zoophytes," because its form of life simulates that of plants, let us pass to plants themselves. In these there is no expenditure in effort, there is no expenditure in maintaining temperature, and the food, some of it supplied by the earth, is the rest of it supplied by a medium which everywhere bathes the outer surface: the utilization of its contained material being effected _gratis_ by the Sun's rays. Just as was to be expected, we here find that agamogenesis may go on without end. Numerous plants and trees are propagated to an unlimited extent by cuttings and buds; and we have sundry plants which cannot be otherwise propagated. The most familiar are the double roses of our gardens: these do not seed, and yet have been distributed everywhere by grafts and buds. Hothouses furnish many cases, as I learn from an authority second to none. Of "the whole host of tropical orchids, for instance, not one per cent. has ever seeded, and some have been a century under cultivation." Again, we have the _Acorus calamus_, "that has hardly been known to seed anywhere, though it is found wild all over the north temperate hemisphere." And then there is the conspicuous and conclusive case of _Eloidea Canadensis_ (alias _Anacharis_,) introduced no one knows how (probably with timber), and first observed in 1847, in several places; and which, having since spread over nearly all England, now everywhere infests ponds, canals, and slow rivers. The plant is dioecious, and only the female exists here. Beyond all question, therefore, this vast progeny of the first slip or fragment introduced, sufficient to cover many square miles were it put together, is constituted entirely of somatic cells. Hence, as far as we can judge, these somatic cells are immortal in the sense given to the word by Professor Weismann; and the evidence that they are so is immeasurably stronger than the evidence which leads him to assert immortality for the fissiparously-multiplying _Protozoa_. This endless multiplication of somatic cells has been going on under the eyes of numerous observers for forty odd years. What observer has watched for forty years to see whether the fissiparous multiplication of _Protozoa_ does not cease? What observer has watched for one year, or one month, or one week?[113]

Even were not Professor Weismann's theory disposed of by this evidence, it might be disposed of by a critical examination of his own evidence, using his own tests. Clearly, if we are to measure relative mortalities, we must assume the conditions to be the same and must use the same measure. Let us do this with some appropriate animal--say Man, as the most open to observation. The mortality of the somatic cells constituting the mass of the human body, is, according to Professor Weismann, shown by the decline and final cessation of cell-multiplication in its various organs. Suppose we apply this test to all the organs: not to those only in which there continually arise bile-cells, epithelium-cells, &c., but to those also in which there arise reproductive cells. What do we find? That the multiplication of these last comes to an end long before the multiplication of the first. In a healthy woman, the cells which constitute the various active tissues of the body, continue to grow and multiply for many years after germ-cells have died out. If similarly measured, then, these cells of the last class prove to be more mortal than those of the first. But Professor Weismann uses a different measure for the two classes of cells. Passing over the illegitimacy of this proceeding, let us accept his other mode of measurement, and see what comes of it. As described by him, absence of death among the _Protozoa_ is implied by that unceasing division and subdivision of which they are said to be capable. Fission continued without end, is the definition of the immortality he speaks of. Apply this conception to the reproductive cells in a _Metazoon_. That the immense majority of them do not multiply without end, we have already seen: with very rare exceptions they die and disappear without result, and they cease their multiplication while the body as a whole still lives. But what of those extremely exceptional ones which, as being actually instrumental to the maintenance of the species, are alone contemplated by Professor Weismann? Do these continue their fissiparous multiplications without end? By no means. The condition under which alone they preserve a qualified form of existence, is that, instead of one becoming two, two become one. A member of series A and a member of series B, coalesce; and so lose their individualities. Now, obviously, if the immortality of a series is shown if its members divide and subdivide perpetually, then the opposite of immortality is shown when, instead of division, there is union. Each series ends, and there is initiated a new series, differing more or less from both. Thus the assertion that the reproductive cells are immortal, can be defended only by changing the conception of immortality otherwise implied.

Even apart from these last criticisms, however, we have clear disproof of the alleged inherent difference between the two classes of cells. Among animals, the multiplication of somatic cells is brought to an end by sundry restraining conditions; but in various plants, where these restraining conditions are absent, the multiplication is unlimited. It may, indeed, be said that the alleged distinction should be reversed; since the fissiparous multiplication of reproductive cells is necessarily interrupted from time to time by coalescence, while that of the somatic cells may go on for a century without being interrupted.

* * * * *

In the essay to which this is a postscript, conclusions were drawn from the remarkable case of the horse and the 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 connection 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 results. Professor Marsh, the distinguished palæontologist, of Yale, New Haven, who is also collecting evidence, sends a preliminary letter in which he says:--"I do not myself know of such a case, but have heard many statements that make their existence probable. One instance, in Connecticut, is vouched for so strongly by an acquaintance of mine, that I have good reason to believe it to be authentic."

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 pure-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."[114]

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 quotation, the remembrance of a remark I heard many years ago concerning dogs, led to the inquiry whether they furnished analogous evidence. It occurred to me that a friend who is frequently appointed judge of animals at agricultural shows, Mr. Fookes, of Fairfield, Pewsey, Wiltshire, might know something about the matter. A letter to him brought various confirmatory statements. From one "who had bred dogs for many years" he learnt that--

"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."

[Since the publication of this article additional evidences have come to
hand. One is from the late Prof. Riley, State Entomologist at Washington,
who says that telegony is an "established principle among well-educated
farmers" in the United States, and who gives me a case in horse-breeding
to which he was himself witness.

Mr. W. P. Smith, writing from Stoughton Grange, Guildford, but giving the
results of his experiences in America, says that "the fact of a previous
conception influencing subsequent offspring was so far recognised among
American cattle-breeders" that it was proposed to raise the rank of any
heifer that had borne a first calf by a thoroughbred bull, and though
this resolution when brought before one of the chief societies was not
carried, yet on all sides it was admitted that previous conceptions had
effects of the kind alleged. Mr. Smith in another letter says:--"When I
had a large mule and horse ranche in America I noticed that the foals of
mares by horse stallions had a mulish appearance in those cases where the
mare had previously given birth to a mule foal. Common heifers who have
had calves by a thoroughbred bull are apt thereafter to have well-bred
calves even from the veriest scrubs."

Yet another very interesting piece of evidence is furnished by Mr. W.
Sedgwick, M.R.C.S., in an article on "The Influence of Heredity in
Disease," published in the _British Medical Journal_ for Feb. 22, 1896,
pp. 460-2. It concerns the transmission of a malformation known among
medical men as hypospadias. Referring to a man belonging to a family in
which this defect prevailed, he writes:--"The widow of the man from whom
these three generations of hypospadians were descended married again,
after an interval of eighteen months; and in this instance the second
husband was not only free from the defect, but there was no history of it
in his family. By this second marriage she had four hypospadiac sons and
four hypospadiac grandsons; whilst there were seven grandsons and three
great-grandsons who were not malformed."]

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.

* * * * *

Notwithstanding experiences showing the futility of controversy for the establishment of truth, I am tempted here to answer opponents at some length. But even could the editor allow me the needful space, I should be compelled, both by lack of time and by ill-health, to be brief. I must content myself with noticing a few points which most nearly concern me.

Referring to my argument respecting tactual discriminativeness, Mr. Wallace thinks that I--

"afford a glaring example of taking the unessential in place of the
essential, and drawing conclusions from a partial and altogether
insufficient survey of the phenomena. For this 'tactual
discriminativeness,' which is alone dealt with by Mr. Spencer, forms the
least important, and probably only an incidental portion of the great
vital phenomenon of skin-sensitiveness, which is at once the watchman and
the shield of the organism against imminent external dangers."
(_Fortnightly Review_, April, 1893, p. 497)

Here Mr. Wallace assumes it to be self-evident that skin-sensitiveness is due to natural selection, and assumes that this must be admitted by me. He supposes it is only the unequal distribution of skin-discriminativeness which I contend is not thus accounted for. But I deny that either the general sensitiveness or the special sensitiveness results from natural selection; and I have years ago justified the first disbelief as I have recently the second. In "The Factors of Organic Evolution" (_Essays_, 454-8), I have given various reasons for inferring that the genesis of the nervous system cannot be due to survival of the fittest; but that it is due to the direct effects of converse between the surface and the environment; and that thus only is to be explained the strange fact that the nervous centres are originally superficial, and migrate inwards during development. These conclusions I have, in the essay Mr. Wallace criticizes, upheld by the evidence which blind boys and skilled compositors furnish; proving, as this does, that increased nervous development is peripherally initiated. Mr. Wallace's belief that skin-sensitiveness arose by natural selection, is unsupported by a single fact. He assumes that it _must_ have been so produced because it is all-important to self-preservation. My belief that it is directly initiated by converse with the environment, is supported by facts; and I have given proof that the assigned cause is now in operation. Am I called upon to abandon my own supported belief and accept Mr. Wallace's unsupported belief? I think not.

Referring to my argument concerning blind cave-animals, Professor Lankester, in _Nature_ of February 23, 1893, writes:--

"Mr. Spencer shows that the saving of ponderable material in the
suppression of an eye is but a small economy: he loses sight of the fact,
however, that possibly, or even probably, the saving to the organism in
the reduction of an eye to a rudimentary state is not to be measured by
mere bulk, but by the non-expenditure of special materials and special
activities which are concerned in the production of an organ so peculiar
and elaborate as is the vertebrate eye."

It seems to me that a supposition is here made to do duty as a fact; and that I might with equal propriety say that "possibly, or even probably," the vertebrate eye is physiologically cheap: its optical part, constituting nearly its whole bulk, consisting of a low order of tissue. There is, indeed, strong reason for considering it physiologically cheap. If any one remembers how relatively enormous are the eyes of a fish just out of the egg--a pair of eyes with a body and head attached; and if he then remembers that every egg contains material for such a pair of eyes; he will see that eye-material constitutes a very considerable part of the fish's roe; and that, since the female fish provides this quantity every year, it cannot be expensive. My argument against Weismann is strengthened rather than weakened by contemplation of these facts.

Professor Lankester asks my attention to a hypothesis of his own, published in the _Encyclopædia Britannica_, concerning the production of blind cave-animals. He thinks it can--

"be fully explained by natural selection acting on congenital fortuitous
variations. Many animals are thus born with distorted or defective eyes
whose parents have not had their eyes submitted to any peculiar
conditions. Supposing a number of some species of Arthropod or Fish to be
swept into a cavern or to be carried from less to greater depths in the
sea, those individuals with perfect eyes would follow the glimmer of
light and eventually escape to the outer air or the shallower depths,
leaving behind those with imperfect eyes to breed in the dark place. A
natural selection would thus be effected" in successive generations.

First of all, I demur to the words "many animals." Under the abnormal conditions of domestication, congenitally defective eyes may be not very uncommon; but their occurrence under natural conditions is, I fancy, extremely rare. Supposing, however, that in a shoal of young fish, there occur some with eyes seriously defective. What will happen? Vision is all-important to the young fish, both for obtaining food and for escaping from enemies. This is implied by the immense development of eyes just referred to; and the obvious conclusion to be drawn is that the partially blind would disappear. Considering that out of the enormous number of young fish hatched with perfect eyes, not one in a hundred reaches maturity, what chance of surviving would there be for those with imperfect eyes? Inevitably they would be starved or be snapped up. Hence the chances that a matured or partially matured semi-blind fish, or rather two such, male and female, would be swept into a cave and left behind are extremely remote. Still more remote must the chances be in the case of cray-fish. Sheltering themselves as these do under stones, in crevices, and in burrows which they make in the banks, and able quickly to anchor themselves to weeds or sticks by their claws, it seems scarcely supposable that any of them could be carried into a cave by a flood. What, then, is the probability that there will be two nearly blind ones, and that these will be thus carried? Then, after this first extreme improbability, there comes a second, which we may, I think, rather call an impossibility. How would it be possible for creatures subject to so violent a change of habitat to survive? Surely death would quickly follow the subjection to such utterly unlike conditions and modes of life. The existence of these blind cave-animals can be accounted for only by supposing that their remote ancestors began making excursions into the cave, and, finding it profitable, extended them, generation after generation, further in: undergoing the required adaptations little by little.[115]

Between Dr. Romanes and myself the first difference concerns the interpretation of "Panmixia." Clearer conceptions of these matters would be reached if, instead of thinking in abstract terms, the physiological processes concerned were brought into the foreground. Beyond the production of changes in the sizes of parts by the selection of fortuitously-arising variations, I can see but one other cause for the production of them--the competition among the parts for nutriment. This has the effect that active parts are well-supplied and grow, while inactive parts are ill-supplied and dwindle.[116] This competition is the cause of "economy of growth"; this is the cause of decrease from disuse; and this is the only conceivable cause of that decrease which Dr. Romanes contends follows the cessation of selection. The three things are aspects of the same thing. And now, before leaving this question, let me remark on the strange proposition which has to be defended by those who deny the dwindling of organs from disuse. Their proposition amounts to this:--that for a hundred generations an inactive organ may be partially denuded of blood all through life, and yet in the hundredth generation will be produced of just the same size as in the first!

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 cases 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 proofs that the surface of a mammalian ovarium is not a spermatheca. The third reply 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 repeatedly referred to. Why, then, does he assume that I abandon my own hypothesis and adopt that of Darwin; thereby entangling myself in difficulties which my own hypothesis avoids? If, as I have argued, the germ-plasm consists of substantially similar units (having only those minute differences expressive of individual and ancestral differences of structure), none of the complicated requirements which Dr. Romanes emphasizes exist; and the alleged inconceivability disappears.

Here I must end: not intending to say more, unless for some very urgent reason; and leaving others to carry on the discussion. I have, indeed, been led to suspend for a short time my proper work, only by consciousness of the transcendent importance of the question at issue. As I have before contended, a right answer to the question whether acquired characters are or are not inherited, underlies right beliefs, not only in Biology and Psychology, but also in Education, Ethics, and Politics.

III.

As a species of literature, controversy is characterised by a terrible fertility. Each proposition becomes the parent of half a dozen; so that a few replies and rejoinders produce an unmanageable population of issues, old and new, which end in being a nuisance to everybody. Remembering this, I shall refrain from dealing with all the points of Professor Weismann's answer. I must limit myself to a part; and that there may be no suspicion of a selection convenient to myself, I will take those contained in his first article.

Before dealing with his special arguments, let me say something about the general mode of argument which Professor Weismann adopts.

The title of his article is "The All-Sufficiency of Natural Selection."[117] Very soon, however, as on p. 322, we come to the admission, which he has himself italicised, "that _it is really very difficult to imagine this process of natural selection in its details_; and to this day it is impossible to demonstrate it in any one point." Elsewhere, as on pp. 327 and 336 _à propos_ of other cases, there are like admissions. But now if the sufficiency of an assigned cause cannot in any case be demonstrated, and if it is "really very difficult to imagine" in what way it has produced its alleged effects, what becomes of the "all-sufficiency" of the cause? How can its all-sufficiency be alleged when its action can neither be demonstrated nor easily imagined? Evidently to fit Professor Weismann's argument the title of the article should have been "The Doubtful Sufficiency of Natural Selection."

Observe, again, how entirely opposite are the ways in which he treats his own interpretation and the antagonist interpretation. He takes the problem presented by certain beautifully adapted structures on the anterior legs of "very many insects," which they use for cleansing their antennæ. These, he argues, cannot have resulted from the inheritance of acquired characters; since any supposed changes produced by function would be changes in the chitinous exo-skeleton, which, being a dead substance, cannot have had its changes transmitted. He then proceeds, very candidly, to point out the extreme difficulties which lie in the way of supposing these structures to have resulted from natural selection: admitting that an opponent might "say that it was absurd" to assume that the successive small variations implied were severally life-saving in their effects. Nevertheless, he holds it unquestionable that natural selection has been the cause. See then the difference. The supposition that the apparatus has been produced by the inheritance of acquired characters is rejected _because_ it presents insuperable difficulties. But the supposition that the apparatus has been produced by natural selection is accepted, _though_ it presents insuperable difficulties. If this mode of reasoning is allowable, no fair comparison between diverse hypotheses can be made.

With these remarks on Professor Weismann's method at large, let me now pass to the particular arguments he uses, taking them _seriatim_.

* * * * *

The first case he deals with is that of the progressive degradation of the human little toe. This he considers a good test case; and he proceeds to discuss an assigned cause--the inherited and accumulated effects of boot-pressure. Without much difficulty he shows that this interpretation is inadequate; since fusion of the phalanges, which constitutes in part the progressive degradation, is found among peoples who go barefoot, and has been found also in Egyptian mummies. Having thus disposed of Mr. Buckman's interpretation, Professor Weismann forthwith concludes that the ascription of this anatomical change to the inheritance of acquired characters is disposed of, and assumes, as the only other possible interpretation, a dwindling "through panmixia": "the hereditary degeneration of the little toe is thus quite simply explained from my standpoint."

It is surprising that Professor Weismann should not have seen that there is an explanation against which his criticism does not tell. If we go back to the genesis of the human type from some lower type of _primates_, we see that while the little toe has ceased to be of any use for climbing purposes, it has not come into any considerable use for walking and running. A glance at the feet of the sub-human _primates_ in general, shows that the inner digits are, as compared with those of men, quite small, have no such relative length and massiveness as the human great toes. Leaving out the question of cause, it is manifest that the great toes have been immensely developed, since there took place the change from arboreal habits to terrestrial habits. A study of the mechanics of walking shows why this has happened. Stability requires that the "line of direction" (the vertical line let fall from the centre of gravity) shall fall within the base, and, in walking, shall be brought at each step within the area of support, or so near it that any tendency to fall may be checked at the next step. A necessary result is that if, at each step, the chief stress of support is thrown on the outer side of the foot, the body must be swayed so that the "line of direction" may fall within the outer side of the foot, or close to it; and when the next step is taken it must be similarly swayed in an opposite way, so that the outer side of the other foot may bear the weight. That is to say, the body must oscillate from side to side, or waddle. The movements of a duck when walking or running show what happens when the points of support are wide apart. Clearly this kind of movement conflicts with efficient locomotion. There is a waste of muscular energy in making these lateral movements, and they are at variance with the forward movement. We may infer, then, that the developing man profited by throwing the stress as much as possible on the inner sides of the feet; and was especially led to do this when going fast, which enabled him to abridge the oscillations: as indeed we now see in a drunken man. Thus there was thrown a continually increasing stress upon the inner digits as they progressively developed from the effects of use; until now that the inner digits, so large compared with the others, bear the greater part of the weight, and being relatively near one another, render needless any marked swayings from side to side. But what has meanwhile happened to the outer digits? Evidently as fast as the great toes have come more and more into play and developed, the little toes have gone more and more out of play and have been dwindling for--how long shall we say?--perhaps a hundred thousand years.

So far, then, am I from feeling that Professor Weismann has here raised a difficulty in the way of the doctrine I hold, that I feel indebted to him for having drawn attention to a very strong evidence in its support. This modification in the form of the foot, which has occurred since arboreal habits have given place to terrestrial habits, shows the effects of use and disuse simultaneously. The inner digits have increased by use while the outer digits have decreased by disuse.

* * * * *

Saying that he will not "pause to refute other apparent proofs of the transmission of acquired characters," Professor Weismann proceeds to deal with the argument which, with various illustrations, I have several times urged--the argument that the natural selection of fortuitously-arising variations cannot account for the adjustment of co-operative parts. Very clearly and very fairly he summarises this argument as used in _The Principles of Biology_ in 1864. Admitting that in this case there are "enormous difficulties" in the way of any other interpretation than the inheritance of acquired characters, Professor Weismann before proceeding to assault this "last bulwark of the Lamarckian principle," premises that the inheritance of acquired characters cannot be a cause of change because inactive as well as active parts degenerate when they cease to be of use: instancing the "skin and skin-armature of crabs and insects." On this I may remark in the first place that an argument derived from degeneracy of passive structures scarcely meets the case of development of active structures; and I may remark in the second place that I have never dreamt of denying the efficiency of natural selection as a cause of degeneracy in passive structures when the degeneracy is such as aids the prosperity of the stirp.

Making this parenthetical reply to his parenthetical criticism I pass to his discussion of this particular argument which he undertakes to dispose of.

His _cheval de bataille_ is furnished him by the social insects--not a fresh one, however, as might be supposed from the way in which he mounts it. From time to time it has carried other riders, who have couched their lances with fatal effects as they supposed. But I hope to show that no one of them has unhorsed an antagonist, and that Professor Weismann fails to do this just as completely as his predecessors. I am, indeed, not sorry that he has afforded me the opportunity of criticising the general discussion concerning the peculiarities of these interesting creatures, which it has often seemed to me sets out with illegitimate assumptions. The supposition always is that the specialities of structures and instincts in the unlike classes of their communities, have arisen during the period in which the communities have existed in something like their present forms. This cannot be. It is doubtless true that association without differentiations of classes may pre-exist for co-operative purposes, as among wolves, and as among various insects which swarm under certain circumstances. Hence we may suppose that there arise in some cases permanent swarms--that survival of the fittest will establish these constant swarms where they are advantageous. But admitting this, we have also to admit a gradual rise of the associated state out of the solitary state. Wasps and bees present us with gradations. If, then, we are to understand how the organized societies have arisen, either out of the solitary state or out of undifferentiated swarms, we must assume that the differences of structure and instinct among the members of them arose little by little, as the social organization arose little by little. Fortunately we are able to trace the greater part of the process in the annually-formed communities of the common wasp; and we shall recognize in it an all-important factor (ignored by Professor Weismann) to which the phenomena, or at any rate the greater part of them, are due.

But before describing the wasp's annual history, let me set down certain observations made when, as a boy, I was given to angling, and, in July or August, sometimes used for bait "wasp-grubs," as they were called. After having had two or three days the combs or "cakes" of these, full of unfed larvæ in all stages of growth, I often saw some of them devouring the edges of their cells to satisfy their appetites; and saw others, probably the most advanced in growth, which were spinning the little covering caps to their cells, in preparation for assuming the pupa state. It is to be inferred that if, after a certain stage of growth has been reached, the food-supply becomes inadequate or is stopped altogether, the larva undergoes its transformation prematurely; and, as we shall presently see, this premature transformation has several natural sequences.

Let us return now to the wasp's family history. In the spring, a queen-wasp or mother-wasp which has survived the winter, begins to make a small nest containing four or more cells in which she lays eggs, and as fast as she builds additional cells, she lays an egg in each. Presently, to these activities, is added the feeding of the larvæ: one result being that the multiplication of larvæ involves a restriction of the food that can be given to each. If we suppose that the mother-wasp rears no more larvæ than she can fully feed, there will result queens or mothers like herself, relatively few in number. But if we suppose that, laying more numerous eggs she produces more larvæ than she can fully feed, the result will be that when these have reached a certain stage of growth, inadequate supply of food will be followed by premature retirement and transformation into pupæ. What will be the characters of the developed insects? The first effect of arrested nutrition will be smaller size. This we find. A second effect will be defective development of parts that are latest formed and least important for the survival of the individual. Hence we may look for arrested development of the reproductive organs--non-essential to individual life. And this expectation is in accord with what we see in animal development at large; for (passing over entirely sexless individuals) we see that though the reproductive organs may be marked out early in the course of development, they are not made fit for action until after the structures for carrying on individual life are nearly complete. The implication is, then, that an inadequately-fed and small larva will become a sterile imago. Having noted this, let us pass to a remarkable concomitant. In the course of development, organs are formed not alone in the order of their original succession, but partly in the order of importance and the share they have to take in adult activities--a change of order called by Haeckel "heterochrony." Hence the fact that we often see the maternal instinct precede the sexual instinct. Every little girl with her doll shows us that the one may become alive while the other remains dormant. In the case of wasps, then, premature arrest of development may result in incompleteness of the sexual traits, along with completeness of the maternal traits. What happens? Leave out the laying of eggs, and the energies of the mother-wasp are spent wholly in building cells and feeding larvæ, and the worker-wasp forthwith begins to spend its life in building cells and feeding larvæ. Thus interpreting the facts, we have no occasion to assume any constitutional difference between the eggs of worker-wasps and the eggs of queens; and that, their eggs are not different we see, first, in the fact that occasionally the worker-wasp is fertile and lays drone-producing eggs, and we see secondly that (if in this respect they are like the bees, of which, however, we have no proof) the larva of a worker-wasp can be changed into the larva of a queen-wasp by special feeding. But be this as it may, we have good evidence that the feeding determines everything. Says Dr. Ormerod, in his _British Social Wasps_:--

"When the swarm is strong and food plentiful ... the well fed larvæ
develop into females, full, large, and overflowing with fat. There are
all gradations of size, from the large fat female to the smallest
worker.... The larger the wasp, the larger and better developed, as the
rule, are the female organs, in all their details. In the largest wasps,
which are to be the queens of another year, the ovaries differ to all
appearances in nothing but their size from those of the larger worker
wasps.... Small feeble swarms produce few or no perfect females; but in
large strong swarms they are found by the score." (pp. 248-9)

To this evidence add the further evidence that queens and workers pass through certain parallel stages in respect of their maternal activities. At first the queen, besides laying eggs, builds cells and feeds larvæ, but after a time ceases to build cells, and feeds larvæ only, and eventually doing neither one nor the other, only lays eggs, and is supplied with food by the workers. So it is in part with the workers. While the members of each successive brood, when in full vigour, build cells and feed larvæ, by-and-by they cease to build cells, and only feed larvæ: the maternal activities and instincts undergo analogous changes. In this case, then, we are not obliged to assume that only by a process of natural selection can the differences of structure and instinct between queens and workers be produced. The only way in which natural selection here comes into play is in the better survival of the families of those queens which made as many cells, and laid as many eggs, as resulted in the best number of half-fed larvæ, producing workers; since by a rapid multiplication of workers the family is advantaged, and the ultimate production of more queens surviving into the next year insured.

The differentiation of classes does not go far among the wasps, because the cycle of processes is limited to a year, or rather to the few months of the summer. It goes further among the hive-bees, which, by storing food, survive from one year into the next. Unlike the queen-wasp, the queen-bee neither builds cells nor gathers food, but is fed by the workers: egg laying has become her sole business. On the other hand the workers, occupied exclusively in building and nursing, have the reproductive organs more dwarfed than they are in wasps. Still we see that the worker-bee occasionally lays drone-producing eggs, and that, by giving extra nutriment and the required extra space, a worker-larva can be developed into a queen-larva. In respect to the leading traits, therefore, the same interpretation holds. Doubtless there are subsidiary instincts which are apparently not thus interpretable. But before it can be assumed that an interpretation of another kind is necessary, it must be shown that these instincts cannot be traced back to those pre-social types and semi-social types which must have preceded the social types we now see. For unquestionably existing bees must have brought with them from the pre-social state an extensive endowment of instincts, and, acquiring other instincts during the unorganized social state, must have brought these into the present organized social state. It is clear, for instance, that the cell-building instinct in all its elaboration was mainly developed in the pre-social stage; for the transition from species building solitary cells to those building combs is traceable. We are similarly enabled to account for swarming as being an inheritance from remote ancestral types. For just in the same way that, with under-feeding of larvæ, there result individuals with imperfectly developed reproductive systems, so there will result individuals with imperfect sexual instincts; and just as the imperfect reproductive system partially operates upon occasion, so will the imperfect sexual instinct. Whence it will result that on the event which causes a queen to undertake a nuptial flight which is effectual, the workers may take abortive nuptial flights: so causing a swarm.

And here, before going further, let us note an instructive class of facts related to the class of facts above set forth. Summing up, in a chapter on "The Determination of Sex," an induction from many cases, Professor Geddes and Mr. Thompson remark that "such conditions as deficient or abnormal food," and others causing "preponderance of waste over repair ... tend to result in production of males;" while "abundant and rich nutrition" and other conditions which "favour constructive processes ... result in the production of females."[118] Among such evidences of this as immediately concern us, are these:--J. H. Fabre found that in the nests of _Osmia tricornis_, eggs at the bottom, first laid, and accompanied by much food, produced females, while those at the top, last laid, and accompanied by one-half or one-third the quantity of food, produced males,[119] Huber's observations on egg-laying by the honey-bee, show that in the normal course of things, the queen lays eggs of workers for eleven months, and only then lays eggs of drones: that is, when declining nutrition or exhaustion has set in. Further, we have the above-named fact, shown by wasps and bees, that when workers lay eggs these produce drones only.[120] Special evidence, harmonizing with general evidence, thus proves that among the social insects the sex is determined by degree of nutrition while the egg is being formed. See then how congruous this evidence is with the conclusion above drawn; for it is proved that after an egg, predetermined as a female, has been laid, the character of the produced insect as a perfect female or imperfect female is determined by the nutrition of the larva. _That is, one set of differences in structures and instincts is determined by nutrition before the egg is laid, and a further set of differences in structures and instincts is determined by nutrition after the egg is laid._

We come now to the extreme case--that of the ants. Is it not probable that the process of differentiation has been similar? There are sundry reasons for thinking so. With ants as with wasps and bees--the workers occasionally lay eggs; and an ant-community can, like a bee-community, when need be, produce queens out of worker-larvæ: presumably in the same manner by extra feeding. But here we have to add special evidence of great significance. For observe that the very facts concerning ants, which Professor Weismann names as exemplifying the formation of the worker type by selection, serve, as in the case of wasps, to exemplify its formation by arrested nutrition. He says that in several species the egg-tubes in the ovaries show progressive decrease in number; and this, like the different degrees of arrest in the ovaries of the worker-wasps, indicates arrest of larva-feeding at different stages. He gives cases showing that, in different degrees, the eyes of workers are less developed in the number of their facets than those of the perfect insects; and he also refers to the wings of workers as not being developed: remarking, however, that the rudiments of their wings show that the ancestral forms had wings. Are not these traits also results of arrested nutrition? Generally among insects the larvæ are either blind or have but rudimentary eyes; that is to say, visual organs are among the latest organs to arise in the genesis of the perfect organism. Hence early arrest of nutrition will stop formation of these, while various more ancient structures have become tolerably complete. Similarly with wings. Wings are late organs in insect phylogeny, and therefore will be among those most likely to abort where development is prematurely arrested. And both these traits will, for the same reason, naturally go along with arrested development of the reproductive system. Even more significant, however, is some evidence assigned by Mr. Darwin respecting the caste-gradations among the driver ants of West Africa. He says:--

"But the most important fact for us is, that, though the workers can be
grouped into castes of different sizes, yet they graduate insensibly into
each other, as does the widely-different structure of their jaws."[121]

"Graduate insensibly," he says; implying that there are very numerous intermediate forms. This is exactly what is to be expected if arrest of nutrition be the cause; for unless the ants have definite measures, enabling them to stop feeding at just the same stages, it must happen that the stoppage of feeding will be indefinite; and that, therefore, there will be all gradations between the extreme forms--"insensible gradations," both in size and in jaw-structure.

In contrast with this interpretation, consider now that of Professor Weismann. From whichever of the two possible suppositions he sets out, the result is equally fatal. If he is consistent, he must say that each of these intermediate forms of workers must have its special set of "determinants," causing its special set of modifications of organs; for he cannot assume that while perfect females and the extreme types of workers have their different sets of determinants, the intermediate types of workers have not. Hence we are introduced to the strange conclusion that besides the markedly-distinguished sets of determinants there must be, to produce these intermediate forms, many other sets slightly distinguished from one another--a score or more kinds of germ-plasm in addition to the four chief kinds. Next comes an introduction to the still stranger conclusion, that these numerous kinds of germ-plasm, producing these numerous intermediate forms, are not simply needless but injurious--produce forms not well fitted for either of the functions discharged by the extreme forms: the implication being that natural selection has originated these disadvantageous forms! If to escape from this necessity for suicide, Professor Weismann accepts the inference that the differences among these numerous intermediate forms are caused by arrested feeding of the larvæ at different stages, then he is bound to admit that the differences between the extreme forms, and between these and perfect females, are similarly caused. But if he does this, what becomes of his hypothesis that the several castes are constitutionally distinct, and result from the operation of natural selection? Observe, too, that his theory does not even allow him to make this choice; for we have clear proof that unlikenesses among the forms of the same species cannot be determined this way or that way by differences of nutrition. English greyhounds and Scotch greyhounds do not differ from one another so much as do the Amazon-workers from the inferior workers, or the workers from the queens. But no matter how a pregnant Scotch greyhound is fed, or her pups after they are born, they cannot be changed into English greyhounds: the different germ-plasms assert themselves spite of all treatment. But in these social insects the different structures of queens and workers _are_ determinable by differences of feeling. Therefore the production of their various castes does not result from the natural selection of varying germ-plasm.