It is well known that de Vries himself considered fluctuating variations and mutations as something quite different. The former he considered as nothing new, only as augmentations or diminutions of something previously existing; and he regarded fluctuations as due to the action of the environment, following in their distribution the laws of chance.[30] Mutations, on the other hand, were something quite new. Now future analysis of variability will not, we think, bear out the validity of this distinction. It is far more likely that a fluctuation is a variation which is the result of some causes the action of which is variable. (We are regarding variability now as subject to “causation” in the physical sense, for only by so regarding it can we attempt its analysis). As a rule this process results in a fluctuation, but if its extent, or degree of operation, exceeds a certain “critical value” a mutation is produced. We may, following the example of the physicists, illustrate this by a “model.”
Fig. 23.
This model is a modification of Galton’s illustration of the degrees of stability of a species. It is a disc of wood rolling on its periphery. We divide it into sectors, and the arcs ab, cd, ef, and gh have all the same radius, 10, 20, 30, and 40. Then we flatten the sectors bc, de, fg, and ha, so that their radii are greater than are those of the other arcs. Now let us cause the disc to roll about the point 8 as a centre. It will oscillate backwards and forwards about a mean position 8. Let us think of these oscillations as fluctuations.
Suppose, however, that we cause the disc to roll a little more violently, so that it oscillates until either of the points 3 or 4 are perpendicularly beneath the centre O. In either of these positions the disc is in a condition of “unstable equilibrium,” and an infinitesimal increase in the extent of an oscillation will cause it to roll beyond the points 3 or 4. But if it does pass either of these critical points it will begin to oscillate about either of the new centres 5 or 7, thus rolling on one of the arcs, ha or de. This assumption of a new condition of stability we may compare with the formation of a mutation.
All this is merely a conceptual physical model of a process about which we know nothing at all. It is meant to illustrate the view that the organisation of a plant or animal is not something absolutely fixed and invariable. The organism in respect of each recognisable and measurable character oscillates about a point of stability, that is to say exhibits fluctuating variations about the mean value of this character. If the stability of the organisation is upset, so that it oscillates, or fluctuates about a new centre, that is, if the variations deviate in either direction from a new “type” or mean, a mutation has been established. A mutation is not, therefore, necessarily a large departure from “normality.” It is not necessarily a “discontinuous variation,” nor a “sport” nor a “freak.” It is essentially a shifting of the mean position about which the variations exhibited by the organism fluctuate.
Such a mutation will, in general, involve the creation of an “elementary species.” We have considered only one character, say stature, in the above discussion, but it generally happens that the assumption of a new centre of stability involves all the characters of the mutating organism. An elementary species therefore differs a little in respect of all its characters from the species from which it arose, or from the other elementary species near which it is situated. This is what we do usually find in the cases of the “races,” or “local varieties,” of any one common species of plant or animal. That we do not recognise that most, or perhaps all, of the species known to systematic biology are really composed of such local races is merely because such results involve an amount of close investigation such as has not generally been possible except in the few cases studied with the object of proving such variability; or in the case of those species which are studied with great attention to detail because of their economic importance. Thus the herrings of North European seas can be divided into such races, and it is possible for a person possessing great familiarity with these fishes to identify the various races or elementary species—that is, to name the locality from which the fish were taken—by considering the characteristics in respect of which the herrings of one part of the sea differ from those of other parts.
The term “variety” has rather a different connotation in systematic biology from that which is included by the term “elementary species.” The meaning of the latter is simple and clear. Two or more elementary species are assemblages of organisms, in each of which assemblages the mean positions about which the various characters fluctuate is different. The term “variety” cannot so easily be defined. The progeny of two different species (in the sense of the term as it is usually applied by systematists) may be called a hybrid variety of one or other of the parent species. In the case of the ordinary species of zoology such a hybrid would, in general, be infertile, or if it did produce offspring these would be infertile. In the case of ordinarily bred offspring from parents of the same species a large deviation from the parental characters might be a malformation, or the result of some irregularity of development. An “atavistic” variation we may regard as the reappearance of some character present in a more or less remote ancestor. Thus dogfishes and skates are no doubt descended from some elasmobranch fish which possessed an anterior dorsal fin. This fin persists in the dog-fishes, but has been lost in the skates and rays. Yet it may appear in the latter fishes as an atavistic variation.
In a variety (following de Vries’ analysis) a character which disappears is not really lost: it is only suppressed, and it still exists in a latent form. Some flowers are coloured, for instance, but there may be varieties in the species to which they belong in which the flowers are colourless. It may not be quite correct, in the physical sense, to say that the colour has been lost, but we may put it in this way. These flowers are then coloured and colourless varieties of the same species. Colour or lack of colour is not, however, fixed in the variety, for the individual plant bearing colourless flowers also bears in its organisation the potentiality of producing coloured flowers. The petals of a flower may be smooth or covered with hairs, and in the same stock both of these varieties may occur. But we must not speak of the presence or absence of hairs as constituting a difference of kind: the smooth-petalled flowers might be regarded as containing the epidermal rudiments of hairs. So also coloured and colourless flowers may be regarded as containing the same kinds of pigment, but these pigments are mixed in different proportions. Such a view enables us to look upon these contrasting characters in the same way as we look upon fluctuating variations, that is, as quantitative differences in the value of the same character.
Such a suppression of a character is not really a loss. An organism belonging to an elementary species in which, say, monochromatic flowers are usually produced may produce flowers which are striped. The progeny of the plant may still produce monochromatic flowers, but we must think of it as also possessing the potentiality of producing striped flowers. In the terminology of Mendelism the characters are dominant and recessive ones.