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I. DIFFERENT KINDS OF VARIABILITY.

Before Darwin, little was known concerning the phenomena of variability. The fact, that hardly two leaves on a tree were exactly the same, could not escape observation: small deviations of the same kind were met with everywhere, among individuals as well as among the organs of the same plant. Larger aberrations, spoken of as monstrosities, were for a long time regarded as lying outside the range of ordinary phenomena. A special branch of inquiry, that of Teratology, was devoted to them, but it constituted a science by itself, sometimes connected with morphology, but having scarcely any bearing on the processes of evolution and heredity.

Darwin was the first to take a broad survey of the whole range of variations in the animal and vegetable kingdoms. His theory of Natural Selection is based on the fact of variability. In order that this foundation should be as strong as possible he collected all the facts, scattered in the literature of his time, and tried to arrange them in a scientific way. He succeeded in showing that variations may be grouped along a line of almost continuous gradations, beginning with simple differences in size and ending with monstrosities. He was struck by the fact that, as a rule, the smaller the deviations, the more frequently they appear, very abrupt breaks in characters being of rare occurrence.

Among these numerous degrees of variability Darwin was always on the look out for those which might, with the greatest probability, be considered as affording material for natural selection to act upon in the development of new species. Neither of the extremes complied with his conceptions. He often pointed out, that there are a good many small fluctuations, which in this respect must be absolutely useless. On the other hand, he strongly combated the belief, that great changes would be necessary to explain the origin of species. Some authors had propounded the idea that highly adapted organs, e.g. the wings of a bird, could not have been developed in any other way than by a comparatively sudden modification of a well defined and important kind. Such a conception would allow of great breaks or discontinuity in the evolution of highly differentiated animals and plants, shortening the time for the evolution of the whole organic kingdom and getting over numerous difficulties inherent in the theory of slow and gradual progress. It would, moreover, account for the genetic relation of the larger groups of both animals and plants. It would, in a word, undoubtedly afford an easy means of simplifying the problem of descent with modification.

Darwin, however, considered such hypotheses as hardly belonging to the domain of science; they belong, he said, to the realm of miracles. That species have a capacity for change is admitted by all evolutionists; but there is no need to invoke modifications other than those represented by ordinary variability. It is well known that in artificial selection this tendency to vary has given rise to numerous distinct races, and there is no reason for denying that it can do the same in nature, by the aid of natural selection. On both lines an advance may be expected with equal probability.

His main argument, however, is that the most striking and most highly adapted modifications may be acquired by successive variations. Each of these may be slight, and they may affect different organs, gradually adapting them to the same purpose. The direction of the adaptations will be determined by the needs in the struggle for life, and natural selection will simply exclude all such changes as occur on opposite or deviating lines. In this way, it is not variability itself which is called upon to explain beautiful adaptations, but it is quite sufficient to suppose that natural selection has operated during long periods in the same way. Eventually, all the acquired characters, being transmitted together, would appear to us, as if they had all been simultaneously developed.

Correlations must play a large part in such special evolutions: when one part is modified, so will be other parts. The distribution of nourishment will come in as one of the causes, the reactions of different organs to the same external influences as another. But no doubt the more effective cause is that of the internal correlations, which, however, are still but dimly understood. Darwin repeatedly laid great stress on this view, although a definite proof of its correctness could not be given in his time. Such proof requires the direct observation of a mutation, and it should be stated here that even the first observations made in this direction have clearly confirmed Darwin's ideas. The new evening primroses which have sprung in my garden from the old form of Oenothera Lamarckiana, and which have evidently been derived from it, in each case, by a single mutation, do not differ from their parent species in one character only, but in almost all their organs and qualities. Oenothera gigas, for example, has stouter stems and denser foliage; the leaves are larger and broader; its thick flower-buds produce gigantic flowers, but only small fruits with large seeds. Correlative changes of this kind are seen in all my new forms, and they lend support to the view that in the gradual development of highly adapted structures, analogous correlations may have played a large part. They easily explain large deviations from an original type, without requiring the assumption of too many steps.

Monstrosities, as their name implies, are widely different in character from natural species; they cannot, therefore, 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.

From this discussion of the main causes of variability we may derive the proposition that the study of every phenomenon in the field of heredity, of variability, and of the origin of new species will have to be considered from two standpoints; on one hand we have the internal causes, on the other the external ones. Sometimes the first are more easily detected, in other cases the latter are more accessible to investigation. But the complete elucidation of any phenomenon of life must always combine the study of the influence of internal with that of external causes.

III. POLYMORPHIC VARIABILITY IN CEREALS.

One of the propositions of Darwin's theory of the struggle for life maintains that the largest amount of life can be supported on any area, by great diversification or divergence in the structure and constitution of its inhabitants. Every meadow and every forest affords a proof of this thesis. The numerical proportion of the different species of the flora is always changing according to external influences. Thus, in a given meadow, some species will flower abundantly in one year and then almost disappear, until, after a series of years, circumstances allow them again to multiply rapidly. Other species, which have taken their places, will then become rare. It follows from this principle, that notwithstanding the constantly changing conditions, a suitable selection from the constituents of a meadow will ensure a continued high production. But, although the principle is quite clear, artificial selection has, as yet, done very little towards reaching a really high standard.

The same holds good for cereals. In ordinary circumstances a field will give a greater yield, if the crop grown consists of a number of sufficiently differing types. Hence it happens that almost all older varieties of wheat are mixtures of more or less diverging forms. In the same variety the numerical composition will vary from year to year, and in oats this may, in bad years, go so far as to destroy more than half of the harvest, the wind-oats (Avena fatua), which scatter their grain to the winds as soon as it ripens, increasing so rapidly that they assume the dominant place. A severe winter, a cold spring and other extreme conditions of life will destroy one form more completely than another, and it is evident that great changes in the numerical composition of the mixture may thus be brought about.

This mixed condition of the common varieties of cereals was well known to Darwin. For him it constituted one of the many types of variability. It is of that peculiar nature to which, in describing other groups, he applies the term polymorphy. It does not imply that the single constituents of the varieties are at present really changing their characters. On the other hand, it does not exclude the possibility of such changes. It simply states that observation shows the existence of different forms; how these have originated is a question which it does not deal with. In his well-known discussion of the variability of cereals, Darwin is mainly concerned with the question, whether under cultivation they have undergone great changes or only small ones. The decision ultimately depends on the question, how many forms have originally been taken into cultivation. Assuming five or six initial species, the variability must be assumed to have been very large, but on the assumption that there were between ten and fifteen types, the necessary range of variability is obviously much smaller. But in regard to this point, we are of course entirely without historical data.

Few of the varieties of wheat show conspicuous differences, although their number is great. If we compare the differentiating characters of the smaller types of cereals with those of ordinary wild species, even within the same genus or family, they are obviously much less marked. All these small characters, however, are strictly inherited, and this fact makes it very probable that the less obvious constituents of the mixtures in ordinary fields must be constant and pure as long as they do not intercross. Natural crossing is in most cereals a phenomenon of rare occurrence, common enough to admit of the production of all possible hybrid combinations, but requiring the lapse of a long series of years to reach its full effect.

Darwin laid great stress on this high amount of variability in the plants of the same variety, and illustrated it by the experience of Colonel Le Couteur ("On the Varieties, Properties, and Classification of Wheat", Jersey, 1837.) on his farm on the isle of Jersey, who cultivated upwards of 150 varieties of wheat, which he claimed were as pure as those of any other agriculturalist. But Professor La Gasca of Madrid, who visited him, drew attention to aberrant ears, and pointed out, that some of them might be better yielders than the majority of plants in the crop, whilst others might be poor types. Thence he concluded that the isolation of the better ones might be a means of increasing his crops. Le Couteur seems to have considered the constancy of such smaller types after isolation as absolutely probable, since he did not even discuss the possibility of their being variable or of their yielding a changeable or mixed progeny. This curious fact proves that he considered the types, discovered in his fields by La Gasca to be of the same kind as his other varieties, which until that time he had relied upon as being pure and uniform. Thus we see, that for him, the variability of cereals was what we now call polymorphy. He looked through his fields for useful aberrations, and collected twenty-three new types of wheat. He was, moreover, clear about one point, which, on being rediscovered after half a century, has become the starting-point for the new Swedish principle of selecting agricultural plants. It was the principle of single-ear sowing, instead of mixing the grains of all the selected ears together. By sowing each ear on a separate plot he intended not only to multiply them, but also to compare their value. This comparison ultimately led him to the choice of some few valuable sorts, one of which, the "Bellevue de Talavera," still holds its place among the prominent sorts of wheat cultivated in France. This variety seems to be really a uniform type, a quality very useful under favourable conditions of cultivation, but which seems to have destroyed its capacity for further improvement by selection.

The principle of single-ear sowing, with a view to obtain pure and uniform strains without further selection, has, until a few years ago, been almost entirely lost sight of. Only a very few agriculturists have applied it: among these are Patrick Shirreff ("Die Verbesserung der Getreide-Arten", translated by R. Hesse, Halle, 1880.) in Scotland and Willet M. Hays ("Wheat, varieties, breeding, cultivation", Univ. Minnesota, Agricultural Experimental Station, Bull. no. 62, 1899.) in Minnesota. Patrick Shirreff observed the fact, that in large fields of cereals, single plants may from time to time be found with larger ears, which justify the expectation of a far greater yield. In the course of about twenty-five years he isolated in this way two varieties of wheat and two of oats. He simply multiplied them as fast as possible, without any selection, and put them on the market.

Hays was struck by the fact that the yield of wheat in Minnesota was far beneath that in the neighbouring States. The local varieties were Fife and Blue Stem. They gave him, on inspection, some better specimens, "phenomenal yielders" as he called them. These were simply isolated and propagated, and, after comparison with the parent-variety and with some other selected strains of less value, were judged to be of sufficient importance to be tested by cultivation all over the State of Minnesota. They have since almost supplanted the original types, at least in most parts of the State, with the result that the total yield of wheat in Minnesota is said to have been increased by about a million dollars yearly.

Definite progress in the method of single-ear sowing has, however, been made only recently. It had been foreshadowed by Patrick Shirreff, who after the production of the four varieties already mentioned, tried to carry out his work on a larger scale, by including numerous minor deviations from the main type. He found by doing so that the chances of obtaining a better form were sufficiently increased to justify the trial. But it was Nilsson who discovered the almost inexhaustible polymorphy of cereals and other agricultural crops and made it the starting-point for a new and entirely trustworthy method of the highest utility. By this means he has produced during the last fifteen years a number of new and valuable races, which have already supplanted the old types on numerous farms in Sweden and which are now being introduced on a large scale into Germany and other European countries.

It is now twenty years since the station at Svalof was founded. During the first period of its work, embracing about five years, selection was practised on the principle which was then generally used in Germany. In order to improve a race a sample of the best ears was carefully selected from the best fields of the variety. These ears were considered as representatives of the type under cultivation, and it was assumed that by sowing their grains on a small plot a family could be obtained, which could afterwards be improved by a continuous selection. Differences between the collected ears were either not observed or disregarded. At Svalof this method of selection was practised on a far larger scale than on any German farm, and the result was, broadly speaking, the same. This may be stated in the following words: improvement in a few cases, failure in all the others. Some few varieties could be improved and yielded excellent new types, some of which have since been introduced into Swedish agriculture and are now prominent races in the southern and middle parts of the country. But the station had definite aims, and among them was the improvement of the Chevalier barley. This, in Middle Sweden, is a fine brewer's barley, but liable to failure during unfavourable summers on account of its slender stems. It was selected with a view of giving it stiffer stems, but in spite of all the care and work bestowed upon it no satisfactory result was obtained.

This experience, combined with a number of analogous failures, could not fail to throw doubt upon the whole method. It was evident that good results were only exceptions, and that in most cases the principle was not one that could be relied upon. The exceptions might be due to unknown causes, and not to the validity of the method; it became therefore of much more interest to search for the causes than to continue the work along these lines.

In the year 1892 a number of different varieties of cereals were cultivated on a large scale and a selection was again made from them. About two hundred samples of ears were chosen, each apparently constituting a different type. Their seeds were sown on separate plots and manured and treated as much as possible in the same manner. The plots were small and arranged in rows so as to facilitate the comparison of allied types. During the whole period of growth and during the ripening of the ears the plots were carefully studied and compared: they were harvested separately; ears and kernels were counted and weighed, and notes were made concerning layering, rust and other cereal pests.

The result of this experiment was, in the main, no distinct improvement. Nilsson was especially struck by the fact that the plots, which should represent distinct types, were far from uniform. Many of them were as multiform as the fields from which the parent-ears were taken. Others showed variability in a less degree, but in almost all of them it was clear that a pure race had not been obtained. The experiment was a fair one, inasmuch as it demonstrated the polymorphic variability of cereals beyond all doubt and in a degree hitherto unsuspected; but from the standpoint of the selectionist it was a failure. Fortunately there were, however, one or two exceptions. A few lots showed a perfect uniformity in regard to all the stalks and ears: these were small families. This fact suggested the idea that each might have been derived from a single ear. During the selection in the previous summer, Nilsson had tried to find as many ears as possible of each new type which he recognised in his fields. But the variability of his crops was so great, that he was rarely able to include more than two or three ears in the same group, and, in a few cases, he found only one representative of the supposed type. It might, therefore, be possible that those small uniform plots were the direct progeny of ears, the grains of which had not been mixed with those from other ears before sowing. Exact records had, of course, been kept of the chosen samples, and the number of ears had been noted in each case. It was, therefore, possible to answer the question and it was found that those plots alone were uniform on which the kernels of one single ear only had been sown. Nilsson concluded that the mixture of two or more ears in a single sowing might be the cause of the lack of uniformity in the progeny. Apparently similar ears might be different in their progeny.

Once discovered, this fact was elevated to the rank of a leading principle and tested on as large a scale as possible. The fields were again carefully investigated and every single ear, which showed a distinct divergence from the main type in one character or another, was selected. A thousand samples were chosen, but this time each sample consisted of one ear only. Next year, the result corresponded to the expectation. Uniformity prevailed almost everywhere; only a few lots showed a discrepancy, which might be ascribed to the accidental selection of hybrid ears. It was now clear that the progeny of single ears was, as a rule, pure, whereas that of mixed ears was impure. The single-ear selection or single-ear sowing, which had fallen into discredit in Germany and elsewhere in Europe, was rediscovered. It proved to be the only trustworthy principle of selection. Once isolated, such single-parent races are constant from seed and remain true to their type. No further selection is needed; they have simply to be multiplied and their real value tested.

Patrick Shirreff, in his early experiments, Le Couteur, Hays and others had observed the rare occurrence of exceptionally good yielders and the value of their isolation to the agriculturist. The possibility of error in the choice of such striking specimens and the necessity of judging their value by their progeny were also known to these investigators, but they had not the slightest idea of all the possibilities suggested by their principle. Nilsson, who is a botanist as well as an agriculturist, discovered that, besides these exceptionably good yielders, every variety of a cereal consists of hundreds of different types, which find the best conditions for success when grown together, but which, after isolation, prove to be constant. Their preference for mixed growth is so definite, that once isolated, their claims on manure and treatment are found to be much higher than those of the original mixed variety. Moreover, the greatest care is necessary to enable them to retain their purity, and as soon as they are left to themselves they begin to deteriorate through accidental crosses and admixtures and rapidly return to the mixed condition.

Reverting now to Darwin's discussion of the variability of cereals, we may conclude that subsequent investigation has proved it to be exactly of the kind which he describes. The only difference is that in reality it reaches a degree, quite unexpected by Darwin and his contemporaries. But it is polymorphic variability in the strictest sense of the word. How the single constituents of a variety originate we do not see. We may assume, and there can hardly be a doubt about the truth of the assumption, that a new character, once produced, will slowly but surely be combined through accidental crosses with a large number of previously existing types, and so will tend to double the number of the constituents of the variety. But whether it first appears suddenly or whether it is only slowly evolved we cannot determine. It would, of course, be impossible to observe either process in such a mixture. Only cultures of pure races, of single-parent races as we have called them, can afford an opportunity for this kind of observation. In the fields of Svalof new and unexpected qualities have recently been seen, from time to time, to appear suddenly. These characters are as distinct as the older ones and appear to be constant from the moment of their origin.

Darwin has repeatedly insisted that man does not cause variability. He simply selects the variations given to him by the hand of nature. He may repeat this process in order to accumulate different new characters in the same family, thus producing varieties of a higher order. This process of accumulation would, if continued for a longer time, lead to the augmentation of the slight differences characteristic of varieties into the greater differences characteristic of species and genera. It is in this way that horticultural and agricultural experience contribute to the problem of the conversion of varieties into species, and to the explanation of the admirable adaptations of each organism to its complex conditions of life. In the long run new forms, distinguished from their allies by quite a number of new characters, would, by the extermination of the older intermediates, become distinct species.

Thus we see that the theory of the origin of species by means of natural selection is quite independent of the question, how the variations to be selected arise. They may arise slowly, from simple fluctuations, or suddenly, by mutations; in both cases natural selection will take hold of them, will multiply them if they are beneficial, and in the course of time accumulate them, so as to produce that great diversity of organic life, which we so highly admire.

Darwin has left the decision of this difficult and obviously subordinate point to his followers. But in his Pangenesis hypothesis he has given us the clue for a close study and ultimate elucidation of the subject under discussion.