Studies in the Theory of Descent, Volume II

In addition to this assumption, which is certainly but little worthy of consideration, there is perhaps one other remaining, viz., that all or many Pierides and other species of butterflies possessed the same tendency to a Heliconoid variation and were always everywhere striving to develop this type, but succeeded only where they accidentally coincided in time and place with the model, the “tendency” being thus furthered by natural selection. But the facts negative this assumption, since such imitative variations have never been observed to a perceptible extent in other species.125

All variations which are demonstrably useful can be similarly dealt with if their origin is explained by variational tendencies.

We perceive that the objection which Von Hartmann brings against heredity is only valid on the ground that this process affords no security for the preservation of variations which occur singly. That heredity itself is a mechanical process is not directly disputed; it is simply assumed that new characters can be transferred by inheritance only when they are produced by the metaphysical “developmental principle,” and not when they arise “accidentally.” This critic does not therefore direct his attack against heredity, but rather against the mechanical origin of variability.

Von Hartmann might have said here that a reference of the phenomenon of heredity to purely mechanical causes, i. e. a mechanical theory of heredity, is up to the present time wanting. That he has not done so proves on the one hand that he despised the dialectical art, but, on the other hand, that he himself has not overlooked the subserviency of the total phenomenon to law, and that he grants the possibility of finding a mechanical explanation therefor. If, in fact, the power of inheritance does not depend upon mechanical principles, I know not what organic processes we are entitled to regard as mechanical, since they are all dependent in essence upon heredity, with which process they are at one, and from which they cannot be thought of as isolated. Haeckel correctly designates reproduction as surplus individual growth, and accordingly refers the phenomena of heredity to those of growth. Conversely, growth may also be designated reproduction, since it depends upon a continuous process of multiplication of the cells composing the organism, from the germ-cell to the innumerable congeries of variously differentiated cells of the highly developed animal body. Who can fail to see that these two processes, the reproduction of the germ-cell and its offspring in the economy of the individual, and the reproduction of individuals and species in the economy of the organic world, show an exact and by no means simply superficial analogy?126 But whoso grants this must also conceive both processes to depend upon the same cause – he cannot assume for the one a causal power and for the other a directive principle. If nutrition and cell-multiplication are purely mechanical processes, so also is heredity. Although it has not yet been possible to demonstrate the mechanism of this phenomenon, it can nevertheless be seen broadly that by means of a minimum of living organic matter (e. g. the protoplasm of the sperm and germ-cell) certain motions are transferred, and these can be regarded as directions of development, as I have already briefly laid down in a former work.127 The power of organisms to transmit their properties to their offspring appears to me to be only conceivable in such a manner “that the germ of the organism by its chemico-physical composition together with its molecular structure, has communicated to it a fixed direction of development – the same direction of development as that originally possessed by the parental organism…” (loc. cit. p. 24). This is confessedly nothing more than a hint, and we do not learn therefrom the means by which developmental direction can be possibly transferred to another organism.

Recently Haeckel, that indefatigable pioneer to whom we are indebted for such a rich store of new ideas, has attempted to bridge over this gap in his essay on “The Perigenesis of the Plastidule,” Berlin, 1876. The basic idea, that heredity depends upon the transference of motion, and variability upon a change of this motion, completely corresponds with the conviction gained in the province of physical science, that “all laws must finally be merged in laws of motion” (Helmholtz128). I hold this view to be the more completely justifiable – although certainly not in the remotest degree as proved – because I formerly designated the acquired individual variations as the “diversion of the inherited direction of development.” Haeckel’s hypothesis in so far accomplishes more than Darwin’s pangenesis, in which a transference of matter, and not of a species of motion peculiar to this matter, is assumed. But although the germ of a mechanical theory of heredity may be contained in Haeckel’s hypothesis, this nevertheless appears to me to be somewhat remote from completely solving the problem. It brings well into prominence one portion of the process of inheritance; under the image of a molecular motion of the plastidule, which motion is modifiable by external influences, we can well understand the fact of a change gradually taking place in the course of generations. On the other hand, the assumption of consciousness in the plastidule, – however admissible philosophically – although only as a formula, scarcely furnishes any deeper knowledge. In the light of a theory, detailed instances which were formerly obscure should become comprehensible. I fail to see, however, how the various forms of atavism, e. g. the reversions which so commonly occur by crossing different races, become more comprehensible by assuming consciousness in the plastidule. If in both parents the plastidule long ago acquired different molecular motions, why, in its rencounters in the germ, does it recollect past times and reassume the older and long abandoned motion? That it does acquire the latter is indeed a fact if we once refer the directional development of the individual to molecular motion of the plastidule; the wherefore does not appear to me, however, to become clearer by assuming consciousness in the plastidule. A mechanical theory of heredity must rather be able to show that the plastidule movements of the male and female germ-cells, in their rencounter in the case of the crossing of widely divergent forms, become mutually modified in such a manner that the motion of the common ancestral form must occur as the resultant. To such demonstration there is however as yet a long step. Haeckel himself moreover points out that his hypothesis is by no means a “mechanical theory of heredity,” but only an introduction to this theory, which he hopes “will be capable of being elevated to the rank of a genetic molecular theory” (loc. cit. p. 17). But although we must also confess with the critic of the “Philosophy of the Unconscious,” that “the facts of heredity have hitherto defied every scientific explanation,”129 this furnishes us with no excuse for flying to a metaphysical explanation, “which is here certainly least able to satisfy the inability to understand the connection arising from natural laws.”

It is not to be wondered at that Von Hartmann, on the ground of the “Unconscious” on which he takes his stand, speaks of the law of correlation as an unconscious acknowledgment of a “non-mechanical universal principle on the side of Darwinism.” By “correlation” he understands something quite different to the idea which we attach to this expression. He supposes that “Darwinism sees itself compelled to acknowledge through empirical facts the uniform correlation of characters pertaining to the specific type; but it thereby contradicts its mechanical principles of explanation, all of which amount to the same thing as conceiving the type as a mosaic, chequered, superficial, and accidental aggregate of characters, which have been singly acquired, contemporaneously or successively, by selection or habit.” I do not believe, however, that any such conception has ever been admitted either by Darwin or any one else. The admission that not all, but only every deep-seated physiological detailed modification, is or may be bound up with a system of correlated changes, indeed implies that we on our side also acknowledge an internal harmony of parts – an equilibrium, as I have above expressed it.

But does this include the admission of a teleological principle, or exclude a mechanical explanation? Do we thereby acknowledge a “specific type” in the sense of an inseparably connected complex of characters, none of which can be taken away without all the others becoming modified? Does such a view agree generally with the empirical facts?

Neither of these views appears to me to represent the case.

I will first answer the second question. On all possible sides the earlier view of the absolute nature of species is contradicted; there is no boundary between species and varieties. But when Von Hartmann assumes that by the transformation of one species “into another” the “whole uniformly connected complex must become changed,” he falls back into the old doctrine of the absolute nature of species, which is sharply contradicted by multitudes of facts. We not unfrequently observe varieties which differ from the parent-form by only a single character, whilst others show numerous differences, and again others may be seen in which the differences predominate. This last deviation would then be designated by many systematists as a new species, but not so by others.

The “specific type” is thus indeed a kind of mosaic-work, but it is a structure to which all the single characters – the stones of the mosaic – belong and build up one harmonious whole, and not a meaningless confusion. Some of the stones or groups of stones can be taken away and replaced by others differently coloured without the structure being thereby necessarily distorted, i. e. destroyed as a structure; but the larger the stones which are exchanged the more necessary will corrections in the other parts of the structure become, in order that the harmony of the whole may be preserved.

Still more weighty than those insensible transitions which in various groups of animals so frequently connect species with species, appear to me, however, the facts made known in the second essay of the second part of this volume, which prove that the two forms in which one species appears can change entirely independently of one another. The caterpillar changes and becomes a new variety or even species (according to the form-value of the change), whilst the butterfly remains unaltered. How could this occur if some other law than that of physiological equilibrium linked together the parts or characters and permitted them to become severed? Must not the two stages become changed with and through one another, like the parts of one body, since they first together constitute the specific type? Is not the fact of this not happening a proof that the whole “uniformly connected complex” of the specific type is not bound and held together by a metaphysical principle, but simply by natural laws?

Now when Von Hartmann comprises the relations of different species to one another under the idea of correlation, such for instance as the relation of dependence in which orchidaceous flowers stand with respect to the insects which visit them, he completely abandons the scientific conception which should be associated with this expression, and compares together two heterogeneous things which have nothing in common excepting that they are both considered by him as a result of the “Unconscious.” The consequence which is then deduced from this correlation of his own construction, viz., that an organic law of correlation is only another expression for a “law of organic development” in the sense of a metaphysical power, obviously cannot be admitted.

By correlation we understand nothing more than the dependence of one part of the organism upon the others and the mutual inter-relations of these parts, which depend entirely upon a “physiological relation of dependence,” as Von Hartmann himself has correctly designated it. Herein is evidently comprised the total morphology of the organism – the structure as a whole, the length, thickness and weight of the single parts, as well as the histological structure of the tissues, since upon all these depends the performance of the single parts. But when, under correlation, Von Hartmann comprises “also a morphological, systematic, inter-action of all the elements of the organism with reference both to the typical ground-plan of the organization as well as to the microscopic anatomical structure of the tissues,” he drags into the idea something foreign to it, not on the ground of facts, but actually in opposition to them, and supported only by a supposed “innate developmental principle” which “is not of a mechanical nature.”

The living organism has already been often compared with a crystal, and the comparison is, mutatis mutandis, justifiable. As in the growing crystal the single molecules cannot become joined together at pleasure, but only in a fixed manner, so are the parts of an organism governed in their respective distribution. In the crystal where nothing but homogeneous parts become grouped together their resulting combination is likewise homogeneous, and it is obvious that they offer but very little possibility of modification, so that the governing laws thus appear restricted and immutable. In the organism, whether regarded microscopically or macroscopically, various parts become combined, and these therefore offer numerous possibilities of modification, so that the governing laws are more complex, and appear less restricted and unchangeable. In neither instance do we know the final causes which always lead to a given state of equilibrium; in the case of a crystal it has not occurred to anybody to ascribe the harmonious disposition of the parts to a teleological power; why then should we assume such a force in the organism, and thus discontinue the attempt, which has already been commenced, to refer to its natural causes that harmony of parts which is here certainly present and equally conformable to law?

On these grounds the assertion that the theory of selection is not an attempt at a “mechanical” explanation of organic development appears to me to be incorrect. Variability and heredity, as well as correlation, admit of being conceived as purely mechanical, and must be thus regarded so long as no more cogent reasons can be adduced for believing that some force other than physico-chemical lies concealed therein.

But we certainly cannot remain at the purely empirical conception as laid down by Darwin in his admirable work on the “Origin of Species.” If the theory of selection is to furnish a method of mechanical explanation, it is essential that its factors should be formulated in a precise mechanical sense. But as soon as we attempt to do this it is seen that, in the first enthusiasm over the newly discovered principle of selection, the one factor of transformation contained in this principle itself has been unduly pushed into the background, to make way for the other more apparent and better known factors.

I have for many years insisted that the first, and perhaps most important, or in any case the most indispensable, factor in every transformation, is the physical nature of the organism itself.130

It would be an error to believe that it is entirely the external conditions which determine what changes shall appear in a given species; the nature of these changes depends essentially upon the physical constitution of the species itself, and a modification actually arising can obviously be only regarded as the resultant of this constitution and of the external influences acting thereon.

But if an essential or perhaps even a preponderating share in determining new characters is to be undoubtedly ascribed to the organism itself, for a mechanical representation of organic developmental processes everything depends upon our being able to conceive this most important factor in a definite theoretical manner, and to comprise under one common point of view its apparently contradictory manifestations of constancy and variability.

Now every change of considerable extent is certainly considered by Darwin to be the direct or indirect consequence of external actions; but indirect action always presupposes a certain small variability (individual variability), without which larger modifications cannot be brought about. Empirically this small amount of variability is doubtless present, but the question arises, upon what does it depend? Can it be conceived as arising mechanically, or is it perhaps just at this point that the metaphysical principle steps in and offers those minute variations which make possible that course of development which, according to this view, is immutably pre-determined? It is certainly the absence of a theoretical definition of variability which always leaves open a door for smuggling in a teleological power. A mechanical explanation of variability must form the basis of this side of the theory of selection.

This explanation is not difficult to find. All dissimilarities of organisms must depend upon the individuals having been affected by dissimilar external influences during the course of the development of organic nature. If we ascribe to the organism the power of giving rise by multiplication only to exact copies of itself, or, more correctly, the power of transmitting unaltered to its successors the motion of its own course of development, each “individual variation” must depend upon the power of the organism to react upon external influences, i. e. to respond by changes of form and of function, and consequently to modify its original (inherited) developmental direction.

It has sometimes been insisted upon, that the “individuals of the same species” or the offspring of one mother cannot be absolutely equal, because, from the commencement of their existence, they have been subjected to dissimilar actions of the environment. But this implies that by perfectly equal influences they would become equal, i. e. it supposes that variability is not inseparably bound up with the essence of the organism, but is only the consequence of developmental tendencies which are in themselves equal being unequally influenced. As a matter of fact the first germs of an individual certainly cannot be supposed to be perfectly equal, because the individual differences of the ancestors must be contained therein in different degrees according to their constitution, and we should have to go back to the primordial organism of the earth in order to find a perfectly homogeneous root, a tabula rasa from which the descendants would commence their development. Whether such a homogeneous root ever existed is however doubtful; it is much more probable that numerous organisms first arose spontaneously,131 and these cannot be presumed to have been absolutely equal, since the conditions under which they came into life cannot have been perfectly identical. Let us, however, for the sake of simplicity assume a single primordial organism; the first generation which took its rise from this by reproduction could only have possessed such individual differences as were produced by the action of dissimilar external influences. But the third generation, together with self-acquired, would also have shown inherited, dissimilarities, and in each succeeding generation the number of tendencies to individual difference imparted to the germ by heredity must have increased to a certain degree, so that it may be said that all germs, from their first origination, bear in themselves a tendency to show individual peculiarities, and would develop these even if they should not be again affected by dissimilar influences. This is obviously the case, since the youngest egg-cells in the ovary of an animal are, as can be demonstrated, always exposed to unequal external conditions with respect to nutrition and pressure.132 Hence, if it were possible that two germs were exactly equal with respect to the direction of development imparted to them by heredity, they would nevertheless furnish two incongruent individuals; and if, conversely, it were possible that two individuals could be exposed to absolutely the same external influences from the formation of the embryo, these also could not be identical, because the individual differences of the ancestors would entail small differences, even in asexual reproduction, in the direction of development transmitted to the egg. The differences between individuals of similar origin thus finally depend entirely upon the dissimilarity of external influences – on the one side upon those which divert the development of the progenitors, and on the other side upon those which divert the individual itself from its course, i. e. from the developmental direction transmitted hereditarily. Although I thus essentially agree with Darwin and Haeckel in so far as these authors refer the “universal individual dissimilarity” to dissimilar external actions, I differ from Darwin in this, that I do not see an essential distinction between the direct and indirect production of individual differences, if by the latter is meant only the unequal influencing of the germ in the parental organism. Haeckel is certainly correct in referring the “primitive differences of the germs produced by the parents” to the inequalities of nutrition to which the single germs must inevitably have been exposed in the parent organism; but another dissimilarity of the germs must evidently be added – a dissimilarity which has nothing to do with unequal nutrition, but which depends upon unequal inheritance of the individual differences of the ancestors, a source of dissimilarity which must arise to a greater extent in sexual than in asexual reproduction. Just as in sexual propagation there occurs a blending of the characters (or more precisely, developmental directions) of two contemporaneous individuals in one germ, so in every mode of reproduction there meet together in the same germ the characters of a whole succession of individuals (the ancestral series), of which the most remote certainly make themselves but seldom felt in a marked degree.

The fact of individual variability can in this way be well understood; the living organism contains in itself no principle of variability – it is the statical element in the developmental processes of the organic world, and would always reproduce exact copies of itself if the inequality of the external influences did not affect the developmental course of each new individual; these influences are therefore the dynamical elements of the process.

From this conception of variability two important empirically established facts can be theoretically deduced, viz. the limitability of variation with respect to quality, which has already been previously mentioned, and the origination of transformations by the direct action of external conditions of life.

If the differences in individuals of the same origin depend upon the action of unequal influences, variation itself is nothing else than the reaction of the organism to a definite external inciting cause, the quality of the variation being determined by the quality of the inciting cause and by that of the organism. In the cases of individual variation hitherto considered, the quality of the organism is equal but that of the inciting cause is unequal, and in this way there arise minute differences in organisms of an equal physical constitution – variations of a different quality.

The same result, viz., different qualities of variation, may also arise in a reverse manner by organisms of a different physical nature being affected by equal external influences. The response of the organism to the cause inciting change would be different according to its nature, or, in other words, organisms of different natures react differently when affected by equal modifying influences. The physical nature of the organism plays the chief part with respect to the quality of the variations; each specific organism can thus give rise to extremely numerous, but not to all conceivable, variations; that is, only to such variations as are made possible by its physical composition. From this it follows further that the possibilities of variation in two species are more widely different, the wider they diverge in physical constitution (including bodily morphology) – that a cycle of variation is peculiar to every species. In this manner we are led to the knowledge that there must certainly exist a “fixed direction of variation,” but not in the sense of Askenasy and Von Hartmann, as the result of an unknown internal principle of development, but as the necessary, i. e. mechanical, consequence of the unequal physical nature of the species, which must respond even to the same inciting cause by unequal variations.