be adduced as evidence in the investigation of the origin of species. There is no doubt that they may have much in common as regards their manner of origin, and that the origin of species, once understood, may lead to a better understanding of the monstrosities. But the reverse is not true, at least not as regards the main lines of development. Here, it is clear, monstrosities cannot have played a part of any significance.

Reversions, or atavistic changes, would seem to give a better support to the theory of descent through modifications. These have been of paramount importance on many lines of evolution of the animal as well as of the vegetable kingdom. It is often assumed that monocotyledons are descended from some lower group of dicotyledons, probably allied to that which includes the buttercup family. On this view the monocotyledons must be assumed to have lost the cambium and all its influence on secondary growth, the differentiation of the flower into calyx and corolla, the second cotyledon or seed-leaf and several other characters. Losses of characters such as these may have been the result of abrupt changes, but this does not prove that the characters themselves have been produced with equal suddenness. On the contrary, Darwin shows very convincingly that a modification may well be developed by a series of steps, and afterwards suddenly disappear. Many monstrosities, such as those represented by twisted stems, furnish direct proofs in support of this view, since they are produced by the loss of one character and this loss implies secondary changes in a large number of other organs and qualities.

Darwin criticises in detail the hypothesis of great and abrupt changes and comes to the conclusion that it does not give even a shadow of an explanation of the origin of species. It is as improbable as it is unnecessary.

Sports and spontaneous variations must now be considered. It is well known that they have produced a large number of fine horticultural varieties. The cut-leaved maple and many other trees and shrubs with split leaves are known to have been produced at a single step; this is true in the case of the single-leaf strawberry plant and of the laciniate variety of the greater celandine: many white flowers, white or yellow berries and numerous other forms had a similar origin. But changes such as these do not come under the head of adaptations, as they consist for the most part in the loss of some quality or organ belonging to the species from which they were derived. Darwin thinks it impossible to attribute to this cause the innumerable structures, which are so well adapted to the habits of life of each species. At the present time we should say that such adaptations require progressive modifications, which are additions to the stock of qualities already possessed by the ancestors, and cannot, therefore, be explained on the ground of a supposed analogy with sports, which are for the most part of a retrogressive nature.

Excluding all these more or less sudden changes, there remains a long series of gradations of variability, but all of these are not assumed by Darwin to be equally fit for the production of new species. In the first place, he disregards all mere temporary variations, such as size, albinism, etc.; further, he points out that very many species have almost certainly been produced by steps, not greater, and probably not very much smaller, than those separating closely related varieties. For varieties are only small species. Next comes the question of polymorphic species: their occurrence seems to have been a source of much doubt and difficulty in Darwin's mind, although at present it forms one of the main supports of the prevailing explanation of the origin of new species. Darwin simply states that this kind of variability seems to be of a peculiar nature; since polymorphic species are now in a stable condition their occurrence gives no clue as to the mode of origin of new species. Polymorphic species are the expression of the result of previous variability acting on a large scale; but they now simply consist of more or less numerous elementary species, which, as far as we know, do not at present exhibit a larger degree of variability than any other more uniform species. The vernal whitlow-grass (Draba verna) and the wild pansy are the best known examples; both have spread over almost the whole of Europe and are split up into hundreds of elementary forms. These sub-species show no signs of any extraordinary degree of variability, when cultivated under conditions necessary for the exclusion of inter-crossing. Hooker has shown, in the case of some ferns distributed over still wider areas, that the extinction of some of the intermediate forms in such groups would suffice to justify the elevation of the remaining types to the rank of distinct species. Polymorphic species may now be regarded as the link which unites ordinary variability with the historical production of species. But it does not appear that they had this significance for Darwin; and, in fact, they exhibit no phenomena which could explain the processes by which one species has been derived from another. By thus narrowing the limits of the species-producing variability Darwin was led to regard small deviations as the source from which natural selection derives material upon which to act. But even these are not all of the same type, and Darwin was well aware of the fact.

It should here be pointed out that in order to be selected, a change must first have been produced. This proposition, which now seems self-evident, has, however, been a source of much difference of opinion among Darwin's followers. The opinion that natural selection produces changes in useful directions has prevailed for a long time. In other words, it was assumed that natural selection, by the simple means of singling out, could induce small and useful changes to increase and to reach any desired degree of deviation from the original type. In my opinion this view was never actually held by Darwin. It is in contradiction with the acknowledged aim of all his work,--the explanation of the origin of species by means of natural forces and phenomena only. Natural selection acts as a sieve; it does not single out the best variations, but it simply destroys the larger number of those which are, from some cause or another, unfit for their present environment. In this way it keeps the strains up to the required standard, and, in special circumstances, may even improve them.

Returning to the variations which afford the material for the sieving- action of natural selection, we may distinguish two main kinds. It is true that the distinction between these was not clear at the time of Darwin, and that he was unable to draw a sharp line between them. Nevertheless, in many cases, he was able to separate them, and he often discussed the question which of the two would be the real source of the differentiation of species. Certain variations constantly occur, especially such as are connected with size, weight, colour, etc. They are usually too small for natural selection to act upon, having hardly any influence in the struggle for life: others are more rare, occurring only from time to time, perhaps once or twice in a century, perhaps even only once in a thousand years. Moreover, these are of another type, not simply affecting size, number or weight, but bringing about something new, which may be useful or not. Whenever the variation is useful natural selection will take hold of it and preserve it; in other cases the variation may either persist or disappear.

In his criticism of miscellaneous objections brought forward against the theory of natural selection after the publication of the first edition of "The Origin of Species", Darwin stated his view on this point very clearly:--"The doctrine of natural selection or the survival of the fittest, which implies that when variations or individual differences of a beneficial nature happen to arise, these will be preserved." ("Origin of Species" (6th edition), page 169, 1882.) In this sentence the words "HAPPEN TO ARISE" appear to me of prominent significance. They are evidently due to the same general conception which prevailed in Darwin's Pangenesis hypothesis. (Cf. de Vries, "Intracellulare Pangenesis", page 73, Jena, 1889, and "Die Mutationstheorie", I. page 63. Leipzig, 1901.)

A distinction is indicated between ordinary fluctuations which are always present, and such variations as "happen to arise" from time to time. ((I think it right to point out that the interpretation of this passage from the "Origin" by Professor de Vries is not accepted as correct either by Mr Francis Darwin or by myself. We do not believe that Darwin intended to draw any distinction between TWO TYPES of variation; the words "when variations or individual differences of a beneficial nature happen to arise" are not in our opinion meant to imply a distinction between ordinary fluctuations and variations which "happen to arise," but we believe that "or" is here used in the sense of ALIAS. With the permission of Professor de Vries, the following extract is quoted from a letter in which he replied to the objection raised to his reading of the passage in question:

"As to your remarks on the passage on page 6, I agree that it is now impossible to see clearly how far Darwin went in his distinction of the different kinds of variability. Distinctions were only dimly guessed at by him. But in our endeavour to arrive at a true conception of his view I think that the chapter on Pangenesis should be our leading guide, and that we should try to interpret the more difficult passages by that chapter. A careful and often repeated study of the Pangenesis hypothesis has convinced me that Darwin, when he wrote that chapter, was well aware that ordinary variability has nothing to do with evolution, but that other kinds of variation were necessary. In some chapters he comes nearer to a clear distinction than in others. To my mind the expression 'happen to arise' is the sharpest indication of his inclining in this direction. I am quite convinced that numerous expressions in his book become much clearer when looked at in this way."

The statement in this passage that "Darwin was well aware that ordinary variability has nothing to do with evolution, but that other kinds of variation were necessary" is contradicted by many passages in the "Origin". A.C.S.)) The latter afford the material for natural selection to act upon on the broad lines of organic development, but the first do not. Fortuitous variations are the species-producing kind, which the theory requires; continuous fluctuations constitute, in this respect, a useless type.

Of late, the study of variability has returned to the recognition of this distinction. Darwin's variations, which from time to time happen to arise, are MUTATIONS, the opposite type being commonly designed fluctuations. A large mass of facts, collected during the last few decades, has confirmed this view, which in Darwin's time could only be expressed with much reserve, and everyone knows that Darwin was always very careful in statements of this kind.

From the same chapter I may here cite the following paragraph: "Thus as I am inclined to believe, morphological differences,...such as the arrangement of the leaves, the divisions of the flower or of the ovarium, the position of the ovules, etc.--first appeared in many cases as fluctuating variations, which sooner or later became constant through the nature of the organism and of the surrounding conditions...but NOT THROUGH NATURAL SELECTION (The italics are mine (H. de V.).); for as these morphological characters do not affect the welfare of the species, any slight deviation in them could not have been governed or accumulated through this latter agency." ("Origin of Species" (6th edition), page 176.) We thus see that in Darwin's opinion, all small variations had not the same importance. In favourable circumstances some could become constant, but others could not.

Since the appearance of the first edition of "The Origin of Species" fluctuating variability has been thoroughly studied by Quetelet. He discovered the law, which governs all phenomena of organic life falling under this head. It is a very simple law, and states that individual variations follow the laws of probability. He proved it, in the first place, for the size of the human body, using the measurements published for Belgian recruits; he then extended it to various other measurements of parts of the body, and finally concluded that it must be of universal validity for all organic beings. It must hold true for all characters in man, physical as well as intellectual and moral qualities; it must hold true for the plant kingdom as well as for the animal kingdom; in short, it must include the whole living world.

Quetelet's law may be most easily studied in those cases where the variability relates to measure, number and weight, and a vast number of facts have since confirmed its exactness and its validity for all kinds of organisms, organs and qualities. But if we examine it more closely, we find that it includes just those minute variations, which, as Darwin repeatedly pointed out, have often no significance for the origin of species. In the phenomena, described by Quetelet's law nothing "happens to arise"; all is governed by the common law, which states that small deviations from the mean type are frequent, but that larger aberrations are rare, the rarer as they are larger. Any degree of variation will be found to occur, if only the number of individuals studied is large enough: it is even possible to calculate before hand, how many specimens must be compared in order to find a previously fixed degree of deviation.

The variations, which from time to time happen to appear, are evidently not governed by this law. They cannot, as yet, be produced at will: no sowings of thousands or even of millions of plants will induce them, although by such means the chance of their occurring will obviously be increased. But they are known to occur, and to occur suddenly and abruptly. They have been observed especially in horticulture, where they are ranged in the large and ill-defined group called sports. Korschinsky has collected all the evidence which horticultural literature affords on this point. (S. Korschinsky, "Heterogenesis und Evolution", "Flora", Vol. LXXXIX. pages 240-363, 1901.) Several cases of the first appearance of a horticultural novelty have been recorded: this has always happened in the same way; it appeared suddenly and unexpectedly without any definite relation to previously existing variability. Dwarf types are one of the commonest and most favourite varieties of flowering plants; they are not originated by a repeated selection of the smallest specimens, but appear at once, without intermediates and without any previous indication. In many instances they are only about half the height of the original type, thus constituting obvious novelties. So it is in other cases described by Korschinsky: these sports or mutations are now recognised to be the main source of varieties of horticultural plants.

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.)

It is probable that nutrition plays as important a part among the external causes of mutability as it does among those of fluctuating variability. Observations in support of this view, however, are too scanty to allow of a definite judgment. Darwin assumed an accumulative influence of external causes in the case of the production of new varieties or species. The accumulation might be limited to the life-time of a single individual, or embrace that of two or more generations. In the end a degree of instability in the equilibrium of one or more characters might be attained, great enough for a character to give way under a small shock produced by changed conditions of life. The character would then be thrown over from the old state of equilibrium into a new one.

Characters which happen to be in this state of unstable equilibrium are called mutable. They may be either latent or active, being in the former case derived from old active ones or produced as new ones (by the process, designated premutation). They may be inherited in this mutable condition during a long series of generations. I have shown that in the case of the evening primrose of Lamarck this state of mutability must have existed for at least half a century, for this species was introduced from Texas into England about the year 1860, and since then all the strains derived from its first distribution over the several countries of Europe show the same phenomena in producing new forms. The production of the dwarf evening primrose, or Oenothera nanella, is assumed to be due to one of the factors, which determines the tall stature of the parent form, becoming latent; this would, therefore, afford an example of retrogressive mutation. Most of the other types of my new mutants, on the other hand, seem to be due to progressive mutability.

The external causes of this curious period of mutability are as yet wholly unknown and can hardly be guessed at, since the origin of the Oenothera Lamarckiana is veiled in mystery. The seeds, introduced into England about 1860, were said to have come from Texas, but whether from wild or from cultivated plants we do not know. Nor has the species been recorded as having been observed in the wild condition. This, however, is nothing peculiar. The European types of Oenothera biennis and O. muricata are in the same condition. The first is said to have been introduced from Virginia, and the second from Canada, but both probably from plants cultivated in the gardens of these countries. Whether the same elementary species are still growing on those spots is unknown, mainly because the different sub-species of the species mentioned have not been systematically studied and distinguished.

The origin of new species, which is in part the effect of mutability, is, however, due mainly to natural selection. Mutability provides the new characters and new elementary species. Natural selection, on the other hand, decides what is to live and what to die. Mutability seems to be free, and not restricted to previously determined lines. Selection, however, may take place along the same main lines in the course of long geological epochs, thus directing the development of large branches of the animal and vegetable kingdom. In natural selection it is evident that nutrition and environment are the main factors. But it is probable that, while nutrition may be one of the main causes of mutability, environment may play the chief part in the decisions ascribed to natural selection. Relations to neighbouring plants and to injurious or useful animals, have been considered the most important determining factors ever since the time when Darwin pointed out their prevailing influence.