Studies in the Theory of Descent, Volume II

The facts, as far as we know them, agree very well with this conclusion. Allied species vary in a similar manner, whilst species which are more distantly related vary in a different manner, even when acted upon by the same external influences. Thus, in the first part of these “Studies” I have remarked that many butterflies under the influence of a warm climate acquire an almost black coloration (Polyommatus Phlæas), whilst on the other hand others become lighter (Papilio Podalirius).

We can thus understand why always certain courses of development are followed, a fact which cannot be completely explained by the nature of the conditions of life which induce the variations. But as soon as we clearly perceive that the quality of the changes essentially depends upon the physical nature of the organism itself, we arrive at the conclusion that species of widely diverging constitutions must give rise to different variations, whilst those of allied constitutions would produce similar variations. But definite courses of development are thus traced out, and we perceive that from any point of the organic developmental series, it is impossible that any other point can be attained at pleasure. Variation in a definite direction thus by no means necessitates the acknowledgment of a metaphysical developmental principle, but can be well conceived as the mechanical result of the physical constitution of the organism.

The manner in which the dissimilar physical constitution of organisms must arise can also be easily shown, although the first commencement of the whole developmental series, i. e. the oldest living forms must be assumed to have been almost homogeneous in their physical constitution. The quality of the variation is, as said before, not merely the product of the physical constitution, but the resultant of this and of the quality of the changing external conditions. Thus from the first “species” there proceeded, through the dissimilar influence of external conditions of life, several new “species,” and as this took place the former physical nature of the organism at the same time became changed, necessitating also a new mode of reacting upon external influences, i. e. another direction of variation. The difference from the primary “species” must certainly be conceived as having been very minute, but it must have increased with each new transformation, and must have proceeded exactly parallel with the degree of physical change connected with each transformation. Thus, hand in hand with the modifications, the power of modification, or mode of reaction of the organism to changing influences, must have continually become re-modified, and we finally obtain an endless number of differently constituted living forms, of which the variational tendencies are different in exact proportion to their physical divergence, so that nearly allied forms respond similarly, and widely divergent forms very differently, to the same inciting causes.

Individual variation arises, as I have attempted to show, by each individual having been continually affected by different, and indeed by constantly changing, influences. Let us, however, imagine on the contrary, that a large group of individuals is affected by the same influences – in fact by such influences as the remaining individuals of the species are not exposed to: this group of individuals would then vary in a nearly similar manner, since both factors of variation, viz. the external influence and the physical constitution, are equal or nearly so. Such local variations would first become prominent when the same external influence had acted upon a series of generations, and the minima of variation produced in the individual by the once-exerted action of the cause inciting change had become augmented by heredity. Transformations of some importance (up to the form-value of species) can thus arise simply by the direct action of the environment, in the same way as that in which individual differences are produced – only the latter fluctuate from generation to generation, since the inciting influences continually change; whilst, in the former, the constant external cause inciting modification always reproduces the same variation, so that an accumulation of the latter can take place. Climatic varieties can be thus explained.

A more efficacious augmentation of the variations arising in the single individual is certainly brought about by the indirect action of the environment upon the organism. It is not here my intention to explain once more the processes of natural selection; I mention this only in order to point out that in these cases transformation depends upon a double action of the environment, since the latter first induces small deviations in the organism by direct action, and then accumulates by selection the variations thus produced.

By regarding variability in this manner – by considering each variation as the reaction of the organism to an external action, as a diversion of the inherited developmental direction, it follows that without a change in the environment no advance in the development of organic forms can take place. If we imagine that from any period in the earth’s history the conditions of life remain completely unchanged, the species present on the earth at this period would not, according to our view, undergo any further modification. Herein is clearly expressed the difference of this view from that other one according to which the inciting principle of modification is not in the environment, but lies in the organism itself in the form of a phyletic vital force.

I cannot here refrain from once more returning to the old (ontogenetic) vital force of the natural philosophers, since the parallel between this and its younger sister, the “phyletic vital force” which appears in so many disguises, is indeed striking. Were the inciting principle of the development of the individual actually an independent vital force acting within the organism, the birth and growth of the individual would be able to take place without the continuous encroachment of the environment, such as occurs in nutrition and respiration. Now this is known to be impossible, so that those who support the existence of such a force, if any still exist, would be driven to the obscure idea of a co-operation between the designing power and the influences of the environment, just in the same manner as such a co-operation is at present postulated by the defenders of the phyletic vital force. I shall further on take the opportunity of pointing out that this last idea is quite untenable; with respect to the (ontogenetic) vital force any clearer proof cannot well be adduced, but it will be admitted that the confused notion of the co-operation and inter-action of teleological and causal powers is, from our point of view, opposed to those very simple and clear ideas which are in harmony with the views on phyletic development. As in racial development each change of the organic type is entirely dependent upon the action of the environment upon the organism, so in the development of the individual, the totality of the phenomena of the personal life must depend upon similar actions. Physiology, as is known, herein entirely supports our view, since this shows that without the continual alternating action of the environment and of the organism there can be no life, and that vital phenomena are nothing but the reactions of the organism to the influences of the environment.

It will be immediately perceived how exactly the processes of phyletic and of ontogenetic development coincide, not merely in their external phenomena but in their nature, if we trace the consequences of the existing knowledge of the structure of the animal body. Although we may not entirely agree with Haeckel’s doctrine of individuality in its details, its correctness must on the whole be conceded, since it cannot be disputed that the notion of individuality is a relative one, and that several categories of morphological individuals exist, which appear not only singly as physiological individuals, i. e. as independent living beings of lowest grade, but which can also combine to form beings of a higher order.

But if we admit this, we should see with Haeckel nothing but reproduction in the origination of a high organism from a single cell, the egg; this reproduction being at the same time combined with various differentiations of the offspring, i. e. with adaptations of the latter to various conditions of life. Not even in the fact that the tissues and organs of a single physiological individual stand in great dependence upon one another through physical causes,133 is there any striking difference between this view and the phyletic composition of the animal (and vegetable) kingdom out of physiological individuals (Haeckel’s “Bionten”), since contemporaneous animals (individuals and species) are known to influence one another in the most active manner.

Now if we further consider that the same units (cells) which, by their reproduction and division of labour, at present compose the body of the highest organism, must at one time have constituted as independent beings the beginning of the whole of organic creation, and that consequently the same processes (division of cells) which now lead to the formation of a mammal, at that time led only to a long series of different independent beings, it will be admitted that both developmental series must depend upon the same inciting powers, and that with reference to the causes of the phenomena it is not possible that any great gap can exist between ontogeny and phylogeny, i. e. between the life-phenomena of the individual and those of the type. According to our view both depend upon that co-operation of the same material physical forces which admits of being briefly summarized as the reaction of organized living matter to influences of the environment.

Our opponents either cannot boast of such harmony in their conception of nature, or else they must, together with the phyletic vital force, re-admit into their theory the old ontogenetic vital force. I know not indeed why they should not do so. Whoever inclines to the view that organic nature is governed not merely by causal, but at the same time by teleological, forces, may admit that the latter are as effective as inciting causes of individual, as they are of phyletic, development. According to my idea they are even bound to admit this, since it cannot be perceived why the adaptations of the ontogeny should not depend upon the same metaphysical principle assumed for each individual, as the adaptations of the phylogeny; the latter are indeed only brought about by the former. I believe therefore that the vital force (ontogenetic) of the ancients stands or falls with the modern (phyletic) vital force. We must admit both or neither, since they both rest on the same basis, and are supported or opposed by the same arguments. Whoever feels justified in setting up a metaphysical principle where complete proof that known forces are sufficient for the explanation of the phenomena has not yet been adduced, must do the same with respect to individual, as he does to phyletic, development, since this proof is in both cases very far from being complete, and still contains large and numerous gaps.134

The theoretical conception of variation as the reaction of the organism to external influences has also not yet been experimentally shown to be correct. Our experiments are still too coarse as compared with the fine distinctions which separate one individual from another; and the difficulty of obtaining clear results is greatly increased by the circumstance that a portion of the individual deviations always depends upon heredity, so that it is frequently not only difficult, but absolutely impossible, to separate those which are inherited from those which are acquired. Still further are we removed from being able to refer variation to its final mechanical causes, i. e. from a mechanical theory of reproduction, which would bring within the range of mathematical calculation both the phenomena of stability (heredity) and of change (variability).

But although sufficient proofs of the correctness of the views here advocated cannot at present be adduced, these views are not contradicted by any known facts – they are, on the contrary, supported by many facts which they in turn make comprehensible (local forms, different cycles of variation in heterogeneous species). These views are finally completely justified by their furnishing the only possible theoretical formulation of variability on which a mechanical conception of organic development can be based. That such a conception is not only admissible, but is unavoidable, at least to the naturalist, I have already attempted to prove.

II. Mechanism and Teleology

In the third volume of his smaller works Karl Ernst von Baer submits the theory of selection to a most searching examination. Without actually calling in question its scientific admissibility, he believes that this theory is dependent upon its satisfying one condition, viz. that it should connect the teleological with the mechanical principle.

“The Darwinian hypothesis, as stated by its supporters, always ends in denying to the processes of nature any relation to a future, i. e. any relation of aim or design. Since such relations appear to me quite evident,” &c. And further: – “If the scientific correctness of the Darwinian hypothesis is to be admitted, it must accommodate itself to this universal striving after a purpose. If it cannot do this we should have to deny its value.”

These words appear almost equivalent to passing a sentence of doom upon the theory of selection and the mechanical conception of nature, for how can one and the same process be effected simultaneously by necessity and by designing powers? The one excludes the other, and we must – so it appears – take our stand either on one side or the other.

Nevertheless we cannot set aside Von Baer’s proposition without further examination simply because it is apparently incapable of being fulfilled, since it contains a truth which should not be overlooked, even by those who uphold the mechanical theory of nature. It is the same truth which is also made use of by the philosophical opponents of this theory, viz. that the universe as a whole cannot be conceived as having arisen from blind necessity – that the endless harmony revealed in every nook and corner by all the phenomena of organic and of inorganic nature cannot possibly be regarded as the work of chance, but rather as the result of a “vast designed process of development.” It is also quite correct when, in reply to the supposed objection that the mechanical theory of nature is not concerned with chances but with necessities, Von Baer answers that the operations of a series of necessities which “are not connected together” can only be termed accidents in their opposing relations. He illustrates this by instancing a target. If I hit the latter by a well-aimed shot, nobody would explain this as the result of an accident, but if “a horseman is riding along a gravelly road past this target, and one of the pebbles thrown up by the hoof of the galloping horse hits the mark, this would be termed an accident of extremely rare occurrence. My target was not the mark for the pebble, therefore the hit was purely accidental, although the projection of the stone in this precise direction with the velocity which it had acquired, was sufficiently explained by the kick given by the horse. But the hit was accidental because the kick of the galloping horse, although it necessarily projected the pebble, had no relation at all to my target. For the same reason we must regard the universe as an immense accident if the forces which move it are not designedly regulated – the more immense because it is not a single motion of projection that acts here, but a large number of heterogeneous powers, i. e. a large number of variously acting necessities which are, as a whole, devoid of purpose, but which nevertheless accomplish this purpose, not only at any single moment, but constantly. A truly admirable series of desirable accidents!”135

The same idea is expressed, although in a very different manner, by Von Hartmann, in the concluding chapter of his work already quoted. He thinks that “design is a necessary and certain consequence of the mechanical laws of nature.” “Were the mechanism of natural laws not teleological there would be no mechanically regulated laws, but a weak chaos of obstinate and capricious powers. Not until the causality of the laws of inorganic nature had superseded the expression “dead” nature, and had shown itself as the mainspring of life and of a conformability to design visible on all sides, did it deserve the name of mechanical lawfulness; just as a complication of wheels and machinery made by man, which move in some definite manner with respect to one another, only acquires the name of a mechanism or of a machine when the immanent teleology of the combination and of the various movements of the parts is revealed.”136

Against the correctness of the idea underlying these statements scarcely anything can in my opinion be said. The harmony of the universe and of that portion of it which we designate organic nature, cannot be explained by chance, i. e. without a common ground for co-operating necessities; by the side of mere mechanism it is impossible not to acknowledge a teleological principle – the only question is, in what manner can we conceive this as acting without abandoning the purely mechanical conception of nature?

This is obviously effected if, with Von Baer and Von Hartmann, we permit the metaphysical principle to interrupt the course of the mechanism of nature, and if we consider both the former and the latter to work together with equal power. Von Hartmann expressly makes such an admission under the name of an “internal principle of development,” to which he attributes such an important share that one cannot understand why it should have any need for the employment of causal powers, and why it does not simply do everything itself. Von Baer expresses himself much less decisively, and even in many places insists upon the purely mechanical connection of organic natural phenomena; but that with him also the idea of interruption by a metaphysical principle is present, is principally shown by his assuming, at least partly, the per saltum development of species. This necessarily involves an actively internal power of development.

Although I have already brought forward many arguments against the existence of such a power, and although in refuting it every form of development by directive powers is at the same time overthrown, it nevertheless appears to me not to be superfluous in such a deeply important question to show that a per saltum development, and especially the so-called heterogeneous generation, is inconceivable, not only on the ground of the arguments formerly employed against the phyletic vital force in general, but quite independently of these.

In the first place it must be said that the positive basis of this hypothesis is insecure. Cases of sudden transformation of the whole organism with subsequent inheritance are as yet quite unknown. It has been shown that the occasional transformation of the Axolotl must most probably be regarded in a different light. Another case, taken for heterogeneous generation, viz. the budding of twelve-rayed Medusæ in the gastric cavity of an eight-rayed species, has lately been shown by Franz Eilhard Schulze137 to be a kind of parasitism or commensalism. The buds of the Cuninæ do not spring, as was supposed, from the Geryonia, but are developed from a Cunina egg. But even if we recall here the cases of alternation of generation and heterogenesis, this would not be of any value by way of proof; it would only be thus indicated how one might picture to oneself a sudden transformation. That in alternation of generation, or generally, in every mode of cyclical reproduction, we have not to deal with the abandonment of one type of organization and the transition to some other, is proved by the continual return to the type of departure – by the cyclical character of the entire transformation. That two quite heterogeneous types can belong to one cycle of development is, however, capable of a far better and more correct explanation than would be given by the supporters of per saltum development. If we trace cyclical reproduction to the adaptation of different developmental stages or generations to deviating conditions of life, we thus not only explain the exact and often striking agreement between form and mode of life – we not only bridge over the gap between metamorphosis and alternation of generation, but we can also understand how, within one and the same family of Hydrozoa, species can occur with or without alternation of generation, and further how other species can exist in which the alternation of generation (the production of free Medusæ) is limited to the one sex; we can understand in general how one continuous series of forms may lead from the simple sexual organ of the Polypes to the independent and free swimming sexual form of the Medusæ, and how hand in hand with this the simple reproduction becomes gradually cyclical. It is just these intermediate steps between the two kinds of reproduction that make quite untenable the idea that the heterogeneous forms in cyclical propagation arise through so-called “heterogeneous generation,” i. e. through sudden per saltum transformation. It is excusable if philosophers to whom these facts are strange, or who have to take the trouble of working them up, should adduce alternation of generation as an instance of “heterogeneous generation,” but by naturalists this should be once and for ever abandoned.

All other facts which have hitherto been referred to “heterogeneous generation” are still less explicable as such, inasmuch as they always relate to changes in single parts of an organism, such as the sudden change of fruit or flower in cultivated plants. The notion of per saltum development, however, demands a total transformation – it comprises (as Von Hartmann quite correctly and logically admits) the idea of a fixed specific type which can only be re-modelled as a whole, and cannot become modified piecemeal. It must further be added, that the observed variations which have arisen abruptly in single parts are not as a rule inherited:138 fruit-trees are only propagated by grafting, i. e. by perpetuating the individual, and not by ordinary reproduction by seeds. Now, if we nowhere see sudden variations of large amount perpetuated by heredity, whilst we everywhere observe small variations which can all be inherited, must it not be concluded that per saltum modification is not the means which Nature employs in transforming species, but that an accumulation of small variations takes place, these leading in time to large differences? Is it logical to reject the latter conclusion because our period of observation is too brief to enable us to directly follow long series of accumulations, whilst per saltum variation is admitted, although unsupported by a single observation? As long as there remains any prospect of tracing large deviations to the continually observed phenomenon of small variations, I believe we have no right to resort to the purely hypothetical explanation afforded by per saltum variations.

But the hypothesis of “heterogeneous generation” is not only without a basis of facts – it can also be directly shown to be untenable. Since the operation of an internal power of transformation does not explain adaptation to the conditions of life, the claims of natural selection to explain these transformations must be admitted; but the co-operation of a phyletic vital force and natural selection is inconceivable if we imagine the modifications to occur per saltum.

The supposed “heterogeneous generation” is always illustrated by the example of alternation of generation. The origination of a new animal form is thus conceived to take place in the same manner as we now see, in the cyclical reproduction of the Medusæ, free swimming, bell-shaped Medusoids, produced from fixed polypites, or Cercariæ from Trematode worms by internal budding; in brief, it is imagined that one animal form suddenly gives rise to another widely deviating form by purely internal causes. Now on this theory it would be an unavoidable postulate, that by such a process of per saltum development there arises not merely a new type of some species, but at the same time individuals capable of living and of persisting under, and fitted to, given conditions of life. But every naturalist who has attempted to completely explain the relation between structure and mode of life knows that even the small differences which separate one species from another, always comprise a number of minute structural deviations which are related to well defined conditions of life – he knows that in every species of animal the whole structure is adapted in the most exact manner in every detail to special conditions of life. It is not an exaggeration when I say in every detail, since the so-called “purely morphological parts” could not be other than they are without causing changes in other parts which exercise a definite function. I will not indeed assert that in the most closely related species all the parts of the body must in some manner differ from one another, if only to a small extent; it seems to me not improbable, however, that an exact comparison would very frequently give this result. That animals which are so widely removed in their morphological relations as Medusæ and Polypes, or Trematoda and their “nurses,” are differently constructed in each of their parts can, however, be stated with certainty.