As already stated, I do not pretend that the production of horticultural novelties is the prototype of the origin of new species in nature. I assume that they are, as a rule, derived from the parent species by the loss of some organ or quality, whereas the main lines of the evolution of the animal and vegetable kingdom are of course determined by progressive changes. Darwin himself has often pointed out this difference. But the saltatory origin of horticultural novelties is as yet the simplest parallel for natural mutations, since it relates to forms and phenomena, best known to the general student of evolution.
The point which I wish to insist upon is this. The difference between small and ever present fluctuations and rare and more sudden variations was clear to Darwin, although the facts known at his time were too meagre to enable a sharp line to be drawn between these two great classes of variability. Since Darwin's time evidence, which proves the correctness of his view, has accumulated with increasing rapidity. Fluctuations constitute one type; they are never absent and follow the law of chance, but they do not afford the material from which to build new species. Mutations, on the other hand, only happen to occur from time to time. They do not necessarily produce greater changes than fluctuations, but such as may become, or rather are from their very nature, constant. It is this constancy which is the mark of specific characters, and on this basis every new specific character may be assumed to have arisen by mutation.
Some authors have tried to show that the theory of mutation is opposed to Darwin's views. But this is erroneous. On the contrary, it is in fullest harmony with the great principle laid down by Darwin. In order to be acted upon by that complex of environmental forces, which Darwin has called natural selection, the changes must obviously first be there. The manner in which they are produced is of secondary importance and has hardly any bearing on the theory of descent with modification. ("Life and Letters" II. 125.)
A critical survey of all the facts of variability of plants in nature as well as under cultivation has led me to the conviction, that Darwin was right in stating that those rare beneficial variations, which from time to time happen to arise—the now so-called mutations—are the real source of progress in the whole realm of the organic world.
II. EXTERNAL AND INTERNAL CAUSES OF VARIABILITY.
All phenomena of animal and plant life are governed by two sets of causes; one of these is external, the other internal. As a rule the internal causes determine the nature of a phenomenon—what an organism can do and what it cannot do. The external causes, on the other hand, decide when a certain variation will occur, and to what extent its features may be developed.
As a very clear and wholly typical instance I cite the cocks-combs (Celosia). This race is distinguished from allied forms by its faculty of producing the well-known broad and much twisted combs. Every single individual possesses this power, but all individuals do not exhibit it in its most complete form. In some cases this faculty may not be exhibited at the top of the main stem, although developed in lateral branches: in others it begins too late for full development. Much depends upon nourishment and cultivation, but almost always the horticulturist has to single out the best individuals and to reject those which do not come up to the standard.
The internal causes are of a historical nature. The external ones may be defined as nourishment and environment. In some cases nutrition is the main factor, as, for instance, in fluctuating variability, but in natural selection environment usually plays the larger part.
The internal or historical causes are constant during the life-time of a species, using the term species in its most limited sense, as designating the so-called elementary species or the units out of which the ordinary species are built up. These historical causes are simply the specific characters, since in the origin of a species one or more of these must have been changed, thus producing the characters of the new type. These changes must, of course, also be due partly to internal and partly to external causes.
In contrast to these changes of the internal causes, the ordinary variability which is exhibited during the life-time of a species is called fluctuating variability. The name mutations or mutating variability is then given to the changes in the specific characters. It is desirable to consider these two main divisions of variability separately.
In the case of fluctuations the internal causes, as well as the external ones, are often apparent. The specific characters may be designated as the mean about which the observed forms vary. Almost every character may be developed to a greater or a less degree, but the variations of the single characters producing a small deviation from the mean are usually the commonest. The limits of these fluctuations may be called wide or narrow, according to the way we look at them, but in numerous cases the extreme on the favoured side hardly surpasses double the value of that on the other side. The degree of this development, for every individual and for every organ, is dependent mainly on nutrition. Better nourishment or an increased supply of food produces a higher development; only it is not always easy to determine which direction is the fuller and which is the poorer one. The differences among individuals grown from different seeds are described as examples of individual variability, but those which may be observed on the same plant, or on cuttings, bulbs or roots derived from one individual are referred to as cases of partial variability. Partial variability, therefore, determines the differences among the flowers, fruits, leaves or branches of one individual: in the main, it follows the same laws as individual variability, but the position of a branch on a plant also determines its strength, and the part it may take in the nourishment of the whole. Composite flowers and umbels therefore have, as a rule, fewer rays on weak branches than on the strong main ones. The number of carpels in the fruits of poppies becomes very small on the weak lateral branches, which are produced towards the autumn, as well as on crowded, and therefore on weakened individuals. Double flowers follow the same rule, and numerous other instances could easily be adduced.
Mutating variability occurs along three main lines. Either a character may disappear, or, as we now say, become latent; or a latent character may reappear, reproducing thereby a character which was once prominent in more or less remote ancestors. The third and most interesting case is that of the production of quite new characters which never existed in the ancestors. Upon this progressive mutability the main development of the animal and vegetable kingdom evidently depends. In contrast to this, the two other cases are called retrogressive and degressive mutability. In nature retrogressive mutability plays a large part; in agriculture and in horticulture it gives rise to numerous varieties, which have in the past been preserved, either on account of their usefulness or beauty, or simply as fancy-types. In fact the possession of numbers of varieties may be considered as the main character of domesticated animals and cultivated plants.
In the case of retrogressive and degressive mutability the internal cause is at once apparent, for it is this which causes the disappearance or reappearance of some character. With progressive mutations the case is not so simple, since the new character must first be produced and then displayed. These two processes are theoretically different, but they may occur together or after long intervals. The production of the new character I call premutation, and the displaying mutation. Both of course must have their external as well as their internal causes, as I have repeatedly pointed out in my work on the Mutation Theory. ("Die Mutationstheorie", 2 vols., Leipzig, 1901.)
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