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,[301] 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.[302] 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.
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.