Form and Function Part 36

_Formative Reize in der thierischen Ontogenese_, Leipzig, 1901.

[506] "La Morphologie dynamique," No. i. of the _Collection de Morphologie dynamique_, Paris, 1911.

[507] "Forme, Puissance et Stabilite des Poissons," No.

iv. of the _Collection_, Paris, 1912.

CHAPTER XIX

SAMUEL BUTLER AND THE MEMORY THEORIES OF HEREDITY

We have laid stress upon the distinction established by Roux between the two stages of development--the automatic and the functional--because of the light which it seems to throw upon the phylogenetic relation of form to function. We have pointed out, too, the paramount role that function plays in Roux's theories of development and heredity, and we have brought out the close kins.h.i.+p existing between his theory and that of Lamarck. For Roux, as for Lamarck, the function creates the organ, and it is only after long generations that the organ appears before the function.

It so happened that just about the time when Roux's papers were beginning to appear a brilliant attempt was made by Samuel Butler to revive and complete the Lamarckian doctrine.

A man of singular freshness and openness of mind, combining in an extraordinary degree extreme intellectual subtlety with a childlike simplicity of outlook, Butler was one of the most fascinating figures of the 19th century. He was not a professional biologist, and much of his biological work is, for that reason, imperfect. But he brought to bear upon the central problems of biology an unbia.s.sed and powerful intelligence, and his att.i.tude to these problems, just because it is that of a cultivated layman, is singularly illuminating.

He was not well acquainted with biological literature; he seems to have hit upon the main ideas of his theory of life and habit in complete independence of Lamarck, and only later to have become aware that Lamarck had in a measure forestalled him. He puts this very beautifully in the following pa.s.sage from his chief biological work _Life and Habit_ (1877[508]):--"I admit that when I began to write upon my subject I did not seriously believe in it. I saw, as it were, a pebble upon the ground, with a sheen that pleased me; taking it up, I turned it over and over for my amus.e.m.e.nt, and found it always grow brighter and brighter the more I examined it. At length I became fascinated, and gave loose rein to self-illusion. The aspect of the world changed; the trifle which I had picked up idly had proved to be a talisman of inestimable value, and had opened a door through which I caught glimpses of a strange and interesting transformation. Then came one who told me that the stone was not mine, but that it had been dropped by Lamarck, to whom it belonged rightfully, but who had lost it; whereon I said I cared not who was the owner, if only I might use it and enjoy it. Now, therefore, having polished it with what art and care one who is no jeweller could bestow upon it, I return it, as best I may, to its possessor" (p. 306). In one of his later works, however, Butler made up for his first neglect of his predecessors by giving what is undeniably the best account in English literature of the work of Buffon, Lamarck, and Erasmus Darwin--in his _Evolution, Old and New_ (1879). Many of his facts he took from Charles Darwin, whose theory of natural selection he bitterly opposed, in the two books just mentioned and in _Unconscious Memory_ (1880) and _Luck or Cunning_ (1887).

Butler's main thesis is that living things are active, intelligent agents, personally continuous with all their ancestors, possessing an intense but unconscious memory of all that their ancestors did and suffered, and moving through habit from the spontaneity of striving to the automatism of remembrance.

The primary cause of all variation in structure is the active response of the organism to needs experienced by it, and the indispensable link between the outer world and the creature itself is that same "sense of need" upon which Lamarck insisted. "According to Lamarck, genera and species have been evolved, in the main, by exactly the same process as that by which human inventions and civilisations are now progressing; and this involves that intelligence, ingenuity, heroism, and all the elements of romance, should have had the main share in the development of every herb and living creature around us" (_Life and Habit_, p. 253).

Variations are indubitably the raw material of evolution--"The question is as to the origin and character of these variations. We say they mainly originate in a creature through a sense of its needs, and vary through the varying surroundings which will cause those needs to vary, and through the opening-up of new desires in many creatures, as the consequence of the gratification of old ones; they depend greatly on differences of individual capacity and temperament; they are communicated, and in the course of time transmitted, as what we call hereditary habits or structures, though these are only, in truth, intense and epitomised memories of how certain creatures liked to deal with protoplasm" (p. 267).

Butler's theory then is essentially a bold and enlightened Lamarckism, completed and rounded off by the conception that heredity too is a psychological process, of the same nature as memory.

In seeking to establish a close a.n.a.logy between memory and heredity Butler starts out from the fact of common experience, that actions which on their first performance require the conscious exercise of will and intelligence, and are then carried out with difficulty and hesitation, gradually through long-continued practice come to be performed easily and automatically, without the conscious exercise of intelligence or will.

He tries to show that this is a general law--that knowledge and will become intense and perfect only when through long-continued exercise they become automatic and unconscious--and he applies this conception to the elucidation of development.

Developmental processes, especially the early ones (of Roux's first stage) are automatic and unconscious, and yet imply the possession by the embryo of a wonderfully perfect knowledge of the processes to be gone through, and an a.s.sured power of will and judgment. Is it conceivable, says Butler, that the embryo can do all these things without knowing how to do them, and without having done them before?

"Shall we say ... that a baby of a day old sucks (which involves the whole principle of the pump, and hence a profound practical knowledge of the laws of pneumatics and hydrostatics), digests, oxygenises its blood (millions of years before Sir Humphrey Davy discovered oxygen), sees and hears--all most difficult and complicated operations, involving a knowledge of the facts concerning optics and acoustics, compared with which the discoveries of Newton sink into utter insignificance? Shall we say that a baby can do all these things at once, doing them so well and so regularly, without being even able to direct its attention to them, and without mistake, and at the same time not know how to do them, and never have done them before?" (p. 54). a.s.suredly not.

The only possible explanation is that the embryo's ancestors have done these things so often, throughout so many millions of generations, that the embryo's knowledge of how to do them has become unconscious and automatic by reason of this age-long practice. This implies that there is in a very real sense actual personal continuity between the embryo and all its ancestors, so that their experiences are his, their memory also his. "We must suppose the continuity of life and sameness between living beings, whether plants or animals, to be far closer than we have hitherto believed; so that the experience of one person is not enjoyed by his successor, so much as that the successor is _bona fide_ but a part of the life of his progenitor, imbued with all his memories, profiting by all his experiences--which are, in fact, his own--and only unconscious of the extent of his own memories and experiences owing to their vastness and already infinite repet.i.tions" (p. 50). It is very suggestive in this connection, he continues--"I. That we are _most conscious of, and have most control over_, such habits as speech, the upright position, the arts and sciences, which are acquisitions peculiar to the human race, always acquired after birth, and not common to ourselves and any ancestor who had not become entirely human.

"II. That we are _less conscious of, and have less control over_, eating and drinking, swallowing, breathing, seeing and hearing, which were acquisitions of our prehuman ancestry, and for which we had provided ourselves with all the necessary apparatus before we saw light, but which are, geologically speaking, recent, or comparatively recent.

"III. That we are _most unconscious of, and have least control over_, our digestion and circulation, which belonged even to our invertebrate ancestry, and which are habits, geologically speaking, of extreme antiquity.... Does it not seem as though the older and more confirmed the habit, the more unquestioning the act of volition, till, in the case of the oldest habits, the practice of succeeding existences has so formulated the procedure, that, on being once committed to such and such a line beyond a certain point, the subsequent course is so clear as to be open to no further doubt, to admit of no alternative, till the very power of questioning is gone, and even the consciousness of volition"

(pp. 51-2).

The hypothesis then, that heredity and development are due to unconscious memory, finds much to support it--"the self-development of each new life in succeeding generations--the various stages through which it pa.s.ses (as it would appear, at first sight, without rhyme or reason), the manner in which it prepares structures of the most surpa.s.sing intricacy and delicacy, for which it has no use at the time when it prepares them, and the many elaborate instincts which it exhibits immediately on, and indeed before, birth--all point in the direction of habit and memory, as the only causes which could produce them" (p. 125). The hypothesis explains, for instance, the fact of recapitulation:--"Why should the embryo of any animal go through so many stages--embryological allusions to forefathers of a widely different type? And why, again, should the germs of the same kind of creature always go through the same stages? If the germ of any animal now living is, in its simplest state, but part of the personal ident.i.ty of one of the original germs of all life whatsoever, and hence, if any now living organism must be considered without quibble as being itself millions of years old, and as imbued with an intense though unconscious memory of all that it has done sufficiently often to have made a permanent impression; if this be so, we can answer the above questions perfectly well. The creature goes through so many intermediate stages between its earliest state as life at all, and its latest development, for the simplest of all reasons, namely, because this is the road by which it has always. .h.i.therto travelled to its present differentiation; this is the road it knows, and into every turn and up or down of which it has been guided by the force of circ.u.mstances and the balance of considerations" (pp. 125-6).

The hypothesis explains also the way in which the orderly succession of stages in embryogeny is brought about, for we can readily understand that the embryo will not remember any stage until it has pa.s.sed through the stage immediately preceding it. "Each step of normal development will lead the impregnated ovum up to, and remind it of, its next ordinary course of action, in the same way as we, when we recite a well-known pa.s.sage, are led up to each successive sentence by the sentence which has immediately preceded it.... Though the ovum immediately after impregnation is instinct with all the memories of both parents, not one of these memories can normally become active till both the ovum itself and its surroundings are sufficiently like what they respectively were, when the occurrence now to be remembered last took place. The memory will then immediately return, and the creature will do as it did on the last occasion that it was in like case as now. This ensures that similarity of order shall be preserved in all the stages of development in successive generations" (pp. 297-8).

Abnormal conditions of development will cause the embryo to pause and hesitate, as if at a loss what to do, having no ancestral experience to guide it. Abnormalities of development represent the embryo's attempt to make the best of an unexpected situation. Or, as Butler puts it, "When ... events are happening to it which, if it has the kind of memory we are attributing to it, would baffle that memory, or which have rarely or never been included in the category of its recollections, _it acts precisely as a creature acts_ _when its recollection is disturbed, or when it is required to do something which it has never done before_" (p.

132). "It is certainly noteworthy that the embryo is never at a loss, unless something happens to it which has not usually happened to its forefathers, and which in the nature of things it cannot remember" (p.

132).

Butler's teleological conception of organic evolution was of course completely antagonistic to the naturalistic conceptions current in his time. In one of his later books he repeats Paley's arguments in favour of design, and to the question, "Where, then, is your designer of beasts and birds, of fishes, and of plants?" he replies: "Our answer is simple enough; it is that we can and do point to a living tangible person with flesh, blood, eyes, nose, ears, organs, senses, dimensions, who did of his own cunning, after infinite proof of every kind of hazard and experiment, scheme out and fas.h.i.+on each organ of the human body. This is the person whom we claim as the designer and artificer of that body, and he is the one of all others the best fitted for the task by his antecedents, and his practical knowledge of the requirements of the case--for he is man himself. Not man, the individual of any given generation, but man in the entirety of his existence from the dawn of life onwards to the present moment" (_Evolution, Old and New_, p. 30, 1879).

Butler's theory of life and habit remained only a sketch, and he was perhaps not fully aware of its philosophical implications. Since Butler's time, a new complexion has been put upon biological philosophy by the profound speculations of Bergson.

But it is not impossible that the future development of biological thought will follow some such lines as those which he tentatively laid down.

Butler was not the first to suggest that there is a close connection between heredity and memory--it is a thought likely to occur to any unprejudiced thinker. The first enunciation of it which attracted general attention was that contained in Hering's famous lecture "On Memory as a general Function of organised Matter."[509] Butler was not aware of Hering's work when he published his _Life and Habit_, but in _Unconscious Memory_ (1880) he gave full credit to Hering as the first discoverer, and supplied an admirable translation of Hering's lecture.

As far as the a.s.similation of heredity to memory is concerned Hering and Butler have much in common, but Hering did not share Butler's Lamarckian and vitalistic views, preferring to hold fast, for the practical purposes of physiology at all events, to the general accepted theory of the parallelism between psychical and physical processes. He was inclined to regard memory in the ordinary sense as a function of the brain, and memory in general as a function of all organised matter.

Speaking of the psychical life, he says, "Thus the cause which produces the unity of all single phenomena of consciousness must be looked for in unconscious life. As we know nothing of this except what we learn from our investigations of matter, and since in a purely empirical consideration, matter and the unconscious must be regarded as identical, the physiologist may justly define memory in a wider sense to be a faculty of the brain, the results of which to a great extent belong to both consciousness and unconsciousness."[510] Hering's views were supported by Haeckel.[511]

In 1893 an American, H. F. Orr,[512] tried to work out a theory of development and heredity based upon the fundamental idea "that the property which is the basis of bodily development in organisms is the same property which we recognise as the basis of psychic activity and psychic development." He tried also to explain the recapitulation of phylogeny by ontogeny as due to habit.

The neo-Lamarckian school of American palaeontologists were also in sympathy with the memory idea, and this was expressed most clearly perhaps by Cope.[513]

In 1904 appeared the work on this subject which has attracted the most attention--R. Semon's _Die Mneme_.[514] This was an elaborate treatment of the question from the materialistic point of view, the main a.s.sumption of Semon's theory being that the action of a stimulus upon the organism leaves a more or less permanent material trace or "engramm," of such a nature as to modify the subsequent action of the organism.

Applied to the explanation of heredity and development, Semon's theory comes to very much the same as Weismann's, with engramms subst.i.tuted for determinants, but it has the great advantage of allowing for the transmission of acquired characters. The application of the concept of stimulus is valuable and suggestive, but it seems to us that the memory theory of heredity can be properly utilised only by adopting a frankly Lamarckian and vitalistic standpoint, and this standpoint Semon expressly combats. As Ward[515] points out in his illuminating lecture on heredity and memory--"Records or memoranda alone are not memory, for they presuppose it. _They_ may consist of physical traces, but memory, even when called 'unconscious,' suggests mind; for, as we have seen, the automatic character implied by this term 'unconscious' presupposes foregone experience.... The mnemic theory then, if it is to be worth anything, seems to me clearly to require not merely physical records or 'engrams,' but living experience or tradition. The mnemic theory will work for those who can accept a monadistic or pampsychist interpretation of the beings that make up the world, who believe with Spinoza and Leibniz that 'all individual things are animated albeit in divers degree'" (pp. 55-6).

Perhaps the best and most ingenious treatment of memory and heredity from a physical standpoint is that offered by E. Rignano in his book, _Sur la transmissibilite des caracteres acquis_.[516] Rignano seeks to construct a physico-chemical "model" which will explain both heredity and memory.

His system, which is based more firmly upon the facts of experimental embryology than Semon's, postulates the existence of "specific nervous acc.u.mulators." The essential hypothesis set up is that every functional stimulus is transformed into specific vital energy, and deposits in the nucleus of the cell a specific substance which is capable of discharging, in an inverse direction, the nervous current which has formed it, as soon as the dynamical equilibrium of the organism is restored to the state in which it was when the original stimulus acted upon it. These specific nuclear substances, different for each cell, are acc.u.mulated also in the nuclei of the germinal substance, const.i.tuting what Rignano calls the central zone of development. That is to say, each functional adaptation changes slightly the dynamical equilibrium of the organism, and this change in the system of distribution of the nervous currents leads to the deposit in the central zone of development of a new specific substance. In the development of the next individual this new specific element enters into activity, and reproduces the nervous current which has formed it, as soon as the organism reaches the same conditions of dynamical equilibrium as those obtaining when the stimulus acted on the parent.

Development can thus be regarded as consisting of a number of stages, at each of which new specific elements enter automatically into play and lead the embryo from that stage to the stage succeeding. The germinal substance on this theory of Rignano's is to be regarded as being composed of a large number of specific elements, originally formed as a result of each new functional adaptation, but now forming part of the hereditary equipment.

The theory represents an advance upon the more static conceptions of Semon. It owes much to Roux's influence.

In this country, the mnemic theories have been championed particularly by M. Hartog[517] and Sir Francis Darwin.[518]

[508] The quotations are taken from the 1910 reprint, London, Fifield.

[509] _Ueber das Gedachtnis als eine allgemeine Funktion der organisierten Materie_, Wien, 1870.

[510] Eng. trans, in E. Hering, _Memory_, p. 9, Chicago and London, 1913.

[511] _Die Perigenesis der Plastidule_, Jena, 1875.

[512] _A Theory of Development and Heredity_, New York, 1893.

[513] _The Primary Factors of Organic Evolution_, Chicago, 1896.

[514] _Die Mneme als erhaltendes Prinzip im Wechsel des organischen Geschehens_, Leipzig, 1904; 2nd ed., 1908.

[515] _Heredity and Memory_, Cambridge, 1913.

[516] Paris, 1906. Also in Italian and German. Eng. trans.

by B. C. ,H. Harvey, Chicago, 1911.

[517] See _Problems of Life and Reproduction_, London, 1913.