This page contains affiliate links. As Amazon Associates we earn from qualifying purchases.
  • 1909
Buy it on Amazon FREE Audible 30 days

tells us, in the most emphatic manner, that she abhors perpetual self- fertilisation…For may we not further infer as probable, in accordance with the belief of the vast majority of the breeders of our domestic productions, that marriage between near relations is likewise in some way injurious, that some unknown great good is derived from the union of individuals which have been kept distinct for many generations?” (Ibid., page 359.)

This view was supported by observations on plants of other families, e.g. Papilionaceae; it could, however, in the absence of experimental proof, be regarded only as a “working hypothesis.”

All adaptations to cross-pollination might also be of use simply because they made pollination possible when for any reason self-pollination had become difficult or impossible. Cross-pollination would, therefore, be of use, not as such, but merely as a means of pollination in general; it would to some extent serve as a remedy for a method unsuitable in itself, such as a modification standing in the way of self-pollination, and on the other hand as a means of increasing the chance of pollination in the case of flowers in which self-pollination was possible, but which might, in accidental circumstances, be prevented. It was, therefore, very important to obtain experimental proof of the conclusion to which Darwin was led by the belief of the majority of breeders and by the evidence of the widespread occurrence of cross-pollination and of the remarkable adaptations thereto.

This was supplied by the researches which are described in the two other works named above. The researches on which the conclusions rest had, in part at least, been previously published in separate papers: this is the case as regards the heterostyled plants. The discoveries which Darwin made in the course of his investigations of these plants belong to the most brilliant in biological science.

The case of Primula is now well known. C.K. Sprengel and others were familiar with the remarkable fact that different individuals of the European species of Primula bear differently constructed flowers; some plants possess flowers in which the styles project beyond the stamens attached to the corolla-tube (long-styled form), while in others the stamens are inserted above the stigma which is borne on a short style (short-styled form). It has been shown by Breitenbach that both forms of flower may occur on the same plant, though this happens very rarely. An analogous case is occasionally met with in hybrids, which bear flowers of different colour on the same plant (e.g. Dianthus caryophyllus). Darwin showed that the external differences are correlated with others in the structure of the stigma and in the nature of the pollen. The long-styled flowers have a spherical stigma provided with large stigmatic papillae; the pollen grains are oblong and smaller than those of the short-styled flowers. The number of the seeds produced is smaller and the ovules larger, probably also fewer in number. The short-styled flowers have a smooth compressed stigma and a corolla of somewhat different form; they produce a greater number of seeds.

These different forms of flowers were regarded as merely a case of variation, until Darwin showed “that these heterostyled plants are adapted for reciprocal fertilisation; so that the two or three forms, though all are hermaphrodites, are related to one another almost like the males and females of ordinary unisexual animals.” (“Forms of Flowers” (1st edition), page 2.) We have here an example of hermaphrodite flowers which are sexually different. There are essential differences in the manner in which fertilisation occurs. This may be effected in four different ways; there are two legitimate and two illegitimate types of fertilisation. The fertilisation is legitimate if pollen from the long-styled flowers reaches the stigma of the short-styled form or if pollen of the short-styled flowers is brought to the stigma of the long-styled flower, that is the organs of the same length of the two different kinds of flower react on one another. Illegitimate fertilisation is represented by the two kinds of self-fertilisation, also by cross-fertilisation, in which the pollen of the long-styled form reaches the stigma of the same type of flower and, similarly, by cross-pollination in the case of the short-styled flowers.

The applicability of the terms legitimate and illegitimate depends, on the one hand, upon the fact that insects which visit the different forms of flowers pollinate them in the manner suggested; the pollen of the short- styled flowers adhere to that part of the insect’s body which touches the stigma of the long-styled flower and vice versa. On the other hand, it is based also on the fact that experiment shows that artificial pollination produces a very different result according as this is legitimate or illegitimate; only the legitimate union ensures complete fertility, the plants thus produced being stronger than those which are produced illegitimately.

If we take 100 as the number of flowers which produce seeds as the result of legitimate fertilisation, we obtain the following numbers from illegitimate fertilisation:

Primula officinalis (P. veris) (Cowslip) … 69 Primula elatior (Oxlip) ……………….. 27 Primula acaulis (P. vulgaris) (Primrose) … 60

Further, the plants produced by the illegitimate method of fertilisation showed, e.g. in P. officinalis, a decrease in fertility in later generations, sterile pollen and in the open a feebler growth. (Under very favourable conditions (in a greenhouse) the fertility of the plants of the fourth generation increases–a point, which in view of various theoretical questions, deserves further investigation.) They behave in fact precisely in the same way as hybrids between species of different genera. This result is important, “for we thus learn that the difficulty in sexually uniting two organic forms and the sterility of their offspring, afford no sure criterion of so-called specific distinctness” (“Forms of Flowers”, page 242): the relative or absolute sterility of the illegitimate unions and that of their illegitimate descendants depend exclusively on the nature of the sexual elements and on their inability to combine in a particular manner. This functional difference of sexual cells is characteristic of the behaviour of hybrids as of the illegitimate unions of heterostyled plants. The agreement becomes even closer if we regard the Primula plants bearing different forms of flowers not as belonging to a systematic entity or “species,” but as including several elementary species. The legitimately produced plants are thus true hybrids (When Darwin wrote in reference to the different forms of heterostyled plants, “which all belong to the same species as certainly as do the two sexes of the same species” (“Cross and Self fertilisation”, page 466), he adopted the term species in a comprehensive sense. The recent researches of Bateson and Gregory (“On the inheritance of Heterostylism in Primula”; “Proc. Roy. Soc.” Ser. B, Vol. LXXVI. 1905, page 581) appear to me also to support the view that the results of illegitimate crossing of heterostyled Primulas correspond with those of hybridisation. The fact that legitimate pollen effects fertilisation, even if illegitimate pollen reaches the stigma a short time previously, also points to this conclusion. Self-pollination in the case of the short-styled form, for example, is not excluded. In spite of this, the numerical proportion of the two forms obtained in the open remains approximately the same as when the pollination was exclusively legitimate, presumably because legitimate pollen is prepotent.), with which their behaviour in other respects, as Darwin showed, presents so close an agreement. This view receives support also from the fact that descendants of a flower fertilised illegitimately by pollen from another plant with the same form of flower belong, with few exceptions, to the same type as that of their parents. The two forms of flower, however, behave differently in this respect. Among 162 seedlings of the long-styled illegitimately pollinated plants of Primula officinalis, including five generations, there were 156 long-styled and only six short-styled forms, while as the result of legitimate fertilisation nearly half of the offspring were long-styled and half short-styled. The short-styled illegitimately pollinated form gave five long-styled and nine short-styled; the cause of this difference requires further explanation. The significance of heterostyly, whether or not we now regard it as an arrangement for the normal production of hybrids, is comprehensively expressed by Darwin: “We may feel sure that plants have been rendered heterostyled to ensure cross-fertilisation, for we now know that a cross between the distinct individuals of the same species is highly important for the vigour and fertility of the offspring.” (“Forms of Flowers”, page 258.) If we remember how important the interpretation of heterostyly has become in all general problems as, for example, those connected with the conditions of the formation of hybrids, a fact which was formerly overlooked, we can appreciate how Darwin was able to say in his autobiography: “I do not think anything in my scientific life has given me so much satisfaction as making out the meaning of the structure of these plants.” (“Life and Letters”, Vol. I. page 91.)

The remarkable conditions represented in plants with three kinds of flowers, such as Lythrum and Oxalis, agree in essentials with those in Primula. These cannot be considered in detail here; it need only be noted that the investigation of these cases was still more laborious. In order to establish the relative fertility of the different unions in Lythrum salicaria 223 different fertilisations were made, each flower being deprived of its male organs and then dusted with the appropriate pollen.

In the book containing the account of heterostyled plants other species are dealt with which, in addition to flowers opening normally (chasmogamous), also possess flowers which remain closed but are capable of producing fruit. These cleistogamous flowers afford a striking example of habitual self-pollination, and H. von Mohl drew special attention to them as such shortly after the appearance of Darwin’s Orchid book. If it were only a question of producing seed in the simplest way, cleistogamous flowers would be the most conveniently constructed. The corolla and frequently other parts of the flower are reduced; the development of the seed may, therefore, be accomplished with a smaller expenditure of building material than in chasmogamous flowers; there is also no loss of pollen, and thus a smaller amount suffices for fertilisation.

Almost all these plants, as Darwin pointed out, have also chasmogamous flowers which render cross-fertilisation possible. His view that cleistogamous flowers are derived from originally chasmogamous flowers has been confirmed by more recent researches. Conditions of nutrition in the broader sense are the factors which determine whether chasmogamous or cleistogamous flowers are produced, assuming, of course, that the plants in question have the power of developing both forms of flower. The former may fail to appear for some time, but are eventually developed under favourable conditions of nourishment. The belief of many authors that there are plants with only cleistogamous flowers cannot therefore be accepted as authoritative without thorough experimental proof, as we are concerned with extra-european plants for which it is often difficult to provide appropriate conditions in cultivation.

Darwin sees in cleistogamous flowers an adaptation to a good supply of seeds with a small expenditure of material, while chasmogamous flowers of the same species are usually cross-fertilised and “their offspring will thus be invigorated, as we may infer from a wide-spread analogy.” (“Forms of Flowers” (1st edition), page 341.) Direct proof in support of this has hitherto been supplied in a few cases only; we shall often find that the example set by Darwin in solving such problems as these by laborious experiment has unfortunately been little imitated.

Another chapter of this book treats of the distribution of the sexes in polygamous, dioecious, and gyno-dioecious plants (the last term, now in common use, we owe to Darwin). It contains a number of important facts and discussions and has inspired the experimental researches of Correns and others.

The most important of Darwin’s work on floral biology is, however, that on cross and self-fertilisation, chiefly because it states the results of experimental investigations extending over many years. Only such experiments, as we have pointed out, could determine whether cross- fertilisation is in itself beneficial, and self-fertilisation on the other hand injurious; a conclusion which a merely comparative examination of pollination-mechanisms renders in the highest degree probable. Later floral biologists have unfortunately almost entirely confined themselves to observations on floral mechanisms. But there is little more to be gained by this kind of work than an assumption long ago made by C.K. Sprengel that “very many flowers have the sexes separate and probably at least as many hermaphrodite flowers are dichogamous; it would thus appear that Nature was unwilling that any flower should be fertilised by its own pollen.”

It was an accidental observation which inspired Darwin’s experiments on the effect of cross and self-fertilisation. Plants of Linaria vulgaris were grown in two adjacent beds; in the one were plants produced by cross- fertilisation, that is, from seeds obtained after fertilisation by pollen of another plant of the same species; in the other grew plants produced by self-fertilisation, that is from seed produced as the result of pollination of the same flower. The first were obviously superior to the latter.

Darwin was surprised by this observation, as he had expected a prejudicial influence of self-fertilisation to manifest itself after a series of generations: “I always supposed until lately that no evil effects would be visible until after several generations of self-fertilisation, but now I see that one generation sometimes suffices and the existence of dimorphic plants and all the wonderful contrivances of orchids are quite intelligible to me.” (“More Letters”, Vol. II. page 373.)

The observations on Linaria and the investigations of the results of legitimate and illegitimate fertilisation in heterostyled plants were apparently the beginning of a long series of experiments. These were concerned with plants of different families and led to results which are of fundamental importance for a true explanation of sexual reproduction.

The experiments were so arranged that plants were shielded from insect- visits by a net. Some flowers were then pollinated with their own pollen, others with pollen from another plant of the same species. The seeds were germinated on moist sand; two seedlings of the same age, one from a cross and the other from a self-fertilised flower, were selected and planted on opposite sides of the same pot. They grew therefore under identical external conditions; it was thus possible to compare their peculiarities such as height, weight, fruiting capacity, etc. In other cases the seedlings were placed near to one another in the open and in this way their capacity of resisting unfavourable external conditions was tested. The experiments were in some cases continued to the tenth generation and the flowers were crossed in different ways. We see, therefore, that this book also represents an enormous amount of most careful and patient original work.

The general result obtained is that plants produced as the result of cross- fertilisation are superior, in the majority of cases, to those produced as the result of self-fertilisation, in height, resistance to external injurious influences, and in seed-production.

Ipomoea purpurea may be quoted as an example. If we express the result of cross-fertilisation by 100, we obtain the following numbers for the fertilised plants.

Generation. Height. Number of seeds.

1 100 : 76 100 : 64
2 100 : 79 –
3 100 : 68 100 : 94
4 100 : 86 100 : 94
5 100 : 75 100 : 89
6 100 : 72 –
7 100 : 81 –
8 100 : 85 –
9 100 : 79 100 : 26 (Number of capsules) 10 100 : 54 –

Taking the average, the ratio as regards growth is 100:77. The considerable superiority of the crossed plants is apparent in the first generation and is not increased in the following generations; but there is some fluctuation about the average ratio. The numbers representing the fertility of crossed and self-fertilised plants are more difficult to compare with accuracy; the superiority of the crossed plants is chiefly explained by the fact that they produce a much larger number of capsules, not because there are on the average more seeds in each capsule. The ratio of the capsules was, e.g. in the third generation, 100:38, that of the seeds in the capsules 100:94. It is also especially noteworthy that in the self-fertilised plants the anthers were smaller and contained a smaller amount of pollen, and in the eighth generation the reduced fertility showed itself in a form which is often found in hybrids, that is the first flowers were sterile. (Complete sterility was not found in any of the plants investigated by Darwin. Others appear to be more sensitive; Cluer found Zea Mais “almost sterile” after three generations of self-fertilisation. (Cf. Fruwirth, “Die Zuchtung der Landwirtschaftlichen Kulturpflanzen”, Berlin, 1904, II. page 6.)

The superiority of crossed individuals is not exhibited in the same way in all plants. For example in Eschscholzia californica the crossed seedlings do not exceed the self-fertilised in height and vigour, but the crossing considerably increases the plant’s capacity for flower-production, and the seedlings from such a mother-plant are more fertile.

The conception implied by the term crossing requires a closer analysis. As in the majority of plants, a large number of flowers are in bloom at the same time on one and the same plant, it follows that insects visiting the flowers often carry pollen from one flower to another of the same stock. Has this method, which is spoken of as Geitonogamy, the same influence as crossing with pollen from another plant? The results of Darwin’s experiments with different plants (Ipomoea purpurea, Digitalis purpurea, Mimulus luteus, Pelargonium, Origanum) were not in complete agreement; but on the whole they pointed to the conclusion that Geitonogamy shows no superiority over self-fertilisation (Autogamy). (Similarly crossing in the case of flowers of Pelargonium zonale, which belong to plants raised from cuttings from the same parent, shows no superiority over self- fertilisation.) Darwin, however, considered it possible that this may sometimes be the case. “The sexual elements in the flowers on the same plant can rarely have been differentiated, though this is possible, as flower-buds are in one sense distinct individuals, sometimes varying and differing from one another in structure or constitution.” (“Cross and Self fertilisation” (1st edition), page 444.)

As regards the importance of this question from the point of view of the significance of cross-fertilisation in general, it may be noted that later observers have definitely discovered a difference between the results of autogamy and geitonogamy. Gilley and Fruwirth found that in Brassica Napus, the length and weight of the fruits as also the total weight of the seeds in a single fruit were less in the case of autogamy than in geitonogamy. With Sinapis alba a better crop of seeds was obtained after geitonogamy, and in the Sugar Beet the average weight of a fruit in the case of a self-fertilised plant was 0.009 gr., from geitonogamy 0.012 gr., and on cross-fertilisation 0.013 gr.

On the whole, however, the results of geitonogamy show that the favourable effects of cross-fertilisation do not depend simply on the fact that the pollen of one flower is conveyed to the stigma of another. But the plants which are crossed must in some way be different. If plants of Ipomoea purpurea (and Mimulus luteus) which have been self-fertilised for seven generations and grown under the same conditions of cultivation are crossed together, the plants so crossed would not be superior to the self- fertilised; on the other hand crossing with a fresh stock at once proves very advantageous. The favourable effect of crossing is only apparent, therefore, if the parent plants are grown under different conditions or if they belong to different varieties. “It is really wonderful what an effect pollen from a distinct seedling plant, which has been exposed to different conditions of life, has on the offspring in comparison with pollen from the same flower or from a distinct individual, but which has been long subjected to the same conditions. The subject bears on the very principle of life, which seems almost to require changes in the conditions.” (“More Letters”, Vol. II. page 406.)

The fertility–measured by the number or weight of the seeds produced by an equal number of plants–noticed under different conditions of fertilisation may be quoted in illustration.

On crossing On crossing On self- with a fresh plants of the fertilisation stock same stock
Mimuleus luteus
(First and ninth generation) 100 4 3

Eschscholzia californica
(second generation) 100 45 40

Dianthus caryophyllus
(third and fourth generation) 100 45 33

Petunia violacea 100 54 46

Crossing under very similar conditions shows, therefore, that the difference between the sexual cells is smaller and thus the result of crossing is only slightly superior to that given by self-fertilisation. Is, then, the favourable result of crossing with a foreign stock to be attributed to the fact that this belongs to another systematic entity or to the fact that the plants, though belonging to the same entity were exposed to different conditions? This is a point on which further researches must be taken into account, especially since the analysis of the systematic entities has been much more thorough than formerly. (In the case of garden plants, as Darwin to a large extent claimed, it is not easy to say whether two individuals really belong to the same variety, as they are usually of hybrid origin. In some instances (Petunia, Iberis) the fresh stock employed by Darwin possessed flowers differing in colour from those of the plant crossed with it.) We know that most of Linneaus’s species are compound species, frequently consisting of a very large number of smaller or elementary species formerly included under the comprehensive term varieties. Hybridisation has in most cases affected our garden and cultivated plants so that they do not represent pure species but a mixture of species.

But this consideration has no essential bearing on Darwin’s point of view, according to which the nature of the sexual cells is influenced by external conditions. Even individuals growing close to one another are only apparently exposed to identical conditions. Their sexual cells may therefore be differently influenced and thus give favourable results on crossing, as “the benefits which so generally follow from a cross between two plants apparently depend on the two differing somewhat in constitution or character.” As a matter of fact we are familiar with a large number of cases in which the condition of the reproductive organs is influenced by external conditions. Darwin has himself demonstrated this for self-sterile plants, that is plants in which self-fertilisation produces no result. This self-sterility is affected by climatic conditions: thus in Brazil Eschscholzia californica is absolutely sterile to the pollen of its own flowers; the descendants of Brazilian plants in Darwin’s cultures were partially self-fertile in one generation and in a second generation still more so. If one has any doubt in this case whether it is a question of the condition of the style and stigma, which possibly prevents the entrance of the pollen-tube or even its development, rather than that of the actual sexual cells, in other cases there is no doubt that an influence is exerted on the latter.

Janczewski (Janczewski, “Sur les antheres steriles des Groseilliers”, “Bull. de l’acad. des sciences de Cracovie”, June, 1908.) has recently shown that species of Ribes cultivated under unnatural conditions frequently produce a mixed (i.e. partly useless) or completely sterile pollen, precisely as happens with hybrids. There are, therefore, substantial reasons for the conclusion that conditions of life exert an influence on the sexual cells. “Thus the proposition that the benefit from cross-fertilisation depends on the plants which are crossed having been subjected during previous generations to somewhat different conditions, or to their having varied from some unknown cause as if they had been thus subjected, is securely fortified on all sides.” (“Cross and Self fertilisation” (1st edition), page 444.)

We thus obtain an insight into the significance of sexuality. If an occasional and slight alteration in the conditions under which plants and animals live is beneficial (Reasons for this are given by Darwin in “Variation under Domestication” (2nd edition), Vol. II. page 127.), crossing between organisms which have been exposed to different conditions becomes still more advantageous. The entire constitution is in this way influenced from the beginning, at a time when the whole organisation is in a highly plastic state. The total life-energy, so to speak, is increased, a gain which is not produced by asexual reproduction or by the union of sexual cells of plants which have lived under the same or only slightly different conditions. All the wonderful arrangements for cross- fertilisation now appear to be useful adaptations. Darwin was, however, far from giving undue prominence to this point of view, though this has been to some extent done by others. He particularly emphasised the following consideration:–“But we should always keep in mind that two somewhat opposed ends have to be gained; the first and more important one being the production of seeds by any means, and the second, cross- fertilisation.” (“Cross and Self fertilisation” (1st edition), page 371.) Just as in some orchids and cleistogamic flowers self-pollination regularly occurs, so it may also occur in other cases. Darwin showed that Pisum sativum and Lathyrus odoratus belong to plants in which self-pollination is regularly effected, and that this accounts for the constancy of certain sorts of these plants, while a variety of form is produced by crossing. Indeed among his culture plants were some which derived no benefit from crossing. Thus in the sixth self-fertilised generation of his Ipomoea cultures the “Hero” made its appearance, a form slightly exceeding its crossed companion in height; this was in the highest degree self-fertile and handed on its characteristics to both children and grandchildren. Similar forms were found in Mimulus luteus and Nicotiana (In Pisum sativum also the crossing of two individuals of the same variety produced no advantage; Darwin attributed this to the fact that the plants had for several generations been self-fertilised and in each generation cultivated under almost the same conditions. Tschermak (“Ueber kunstliche Kreuzung an Pisum sativum”) afterwards recorded the same result; but he found on crossing different varieties that usually there was no superiority as regards height over the products of self-fertilisation, while Darwin found a greater height represented by the ratios 100:75 and 100:60.), types which, after self-fertilisation, have an enhanced power of seed-production and of attaining a greater height than the plants of the corresponding generation which are crossed together and self-fertilised and grown under the same conditions. “Some observations made on other plants lead me to suspect that self-fertilisation is in some respects beneficial; although the benefit thus derived is as a rule very small compared with that from a cross with a distinct plant.” (“Cross and Self fertilisation”, page 350.) We are as ignorant of the reason why plants behave differently when crossed and self-fertilised as we are in regard to the nature of the differentiation of the sexual cells, which determines whether a union of the sexual cells will prove favourable or unfavourable.

It is impossible to discuss the different results of cross-fertilisation; one point must, however, be emphasised, because Darwin attached considerable importance to it. It is inevitable that pollen of different kinds must reach the stigma. It was known that pollen of the same “species” is dominant over the pollen of another species, that, in other words, it is prepotent. Even if the pollen of the same species reaches the stigma rather later than that of another species, the latter does not effect fertilisation.

Darwin showed that the fertilising power of the pollen of another variety or of another individual is greater than that of the plant’s own pollen. (“Cross and Self fertilisation”, page 391.) This has been demonstrated in the case of Mimulus luteus (for the fixed white-flowering variety) and Iberis umbellata with pollen of another variety, and observations on cultivated plants, such as cabbage, horseradish, etc. gave similar results. It is, however, especially remarkable that pollen of another individual of the same variety may be prepotent over the plant’s own pollen. This results from the superiority of plants crossed in this manner over self- fertilised plants. “Scarcely any result from my experiments has surprised me so much as this of the prepotency of pollen from a distinct individual over each plant’s own pollen, as proved by the greater constitutional vigour of the crossed seedlings.” (Ibid. page 397.) Similarly, in self- fertile plants the flowers of which have not been deprived of the male organs, pollen brought to the stigma by the wind or by insects from another plant effects fertilisation, even if the plant’s own pollen has reached the stigma somewhat earlier.

Have the results of his experimental investigations modified the point of view from which Darwin entered on his researches, or not? In the first place the question is, whether or not the opinion expressed in the Orchid book that there is “Something injurious” connected with self-fertilisation, has been confirmed. We can, at all events, affirm that Darwin adhered in essentials to his original position; but self-fertilisation afterwards assumed a greater importance than it formerly possessed. Darwin emphasised the fact that “the difference between the self-fertilised and crossed plants raised by me cannot be attributed to the superiority of the crossed, but to the inferiority of the self-fertilised seedlings, due to the injurious effects of self-fertilisation.” (Ibid. page 437.) But he had no doubt that in favourable circumstances self-fertilised plants were able to persist for several generations without crossing. An occasional crossing appears to be useful but not indispensable in all cases; its sporadic occurrence in plants in which self-pollination habitually occurs is not excluded. Self-fertilisation is for the most part relatively and not absolutely injurious and always better than no fertilisation. “Nature abhors perpetual self-fertilisation” (It is incorrect to say, as a writer has lately said, that the aphorism expressed by Darwin in 1859 and 1862, “Nature abhors perpetual self-fertilisation,” is not repeated in his later works. The sentence is repeated in “Cross and Self fertilisation” (page 8), with the addition, “If the word perpetual had been omitted, the aphorism would have been false. As it stands, I believe that it is true, though perhaps rather too strongly expressed.”) is, however, a pregnant expression of the fact that cross-fertilisation is exceedingly widespread and has been shown in the majority of cases to be beneficial, and that in those plants in which we find self-pollination regularly occurring cross- pollination may occasionally take place.

An attempt has been made to express in brief the main results of Darwin’s work on the biology of flowers. We have seen that his object was to elucidate important general questions, particularly the question of the significance of sexual reproduction.

It remains to consider what influence his work has had on botanical science. That this influence has been very considerable, is shown by a glance at the literature on the biology of flowers published since Darwin wrote. Before the book on orchids was published there was nothing but the old and almost forgotten works of Kolreuter and Sprengel with the exception of a few scattered references. Darwin’s investigations gave the first stimulus to the development of an extensive literature on floral biology. In Knuth’s “Handbuch der Blutenbiologie” (“Handbook of Flower Pollination”, Oxford, 1906) as many as 3792 papers on this subject are enumerated as having been published before January 1, 1904. These describe not only the different mechanisms of flowers, but deal also with a series of remarkable adaptations in the pollinating insects. As a fertilising rain quickly calls into existence the most varied assortment of plants on a barren steppe, so activity now reigns in a field which men formerly left deserted. This development of the biology of flowers is of importance not only on theoretical grounds but also from a practical point of view. The rational breeding of plants is possible only if the flower-biology of the plants in question (i.e. the question of the possibility of self-pollination, self- sterility, etc.) is accurately known. And it is also essential for plant- breeders that they should have “the power of fixing each fleeting variety of colour, if they will fertilise the flowers of the desired kind with their own pollen for half-a-dozen generations, and grow the seedlings under the same conditions.” (“Cross and Self fertilisation” (1st edition), page 460.)

But the influence of Darwin on floral biology was not confined to the development of this branch of Botany. Darwin’s activity in this domain has brought about (as Asa Gray correctly pointed out) the revival of teleology in Botany and Zoology. Attempts were now made to determine, not only in the case of flowers but also in vegetative organs, in what relation the form and function of organs stand to one another and to what extent their morphological characters exhibit adaptation to environment. A branch of Botany, which has since been called Ecology (not a very happy term) has been stimulated to vigorous growth by floral biology.

While the influence of the work on the biology of flowers was extraordinarily great, it could not fail to elicit opinions at variance with Darwin’s conclusions. The opposition was based partly on reasons valueless as counterarguments, partly on problems which have still to be solved; to some extent also on that tendency against teleological conceptions which has recently become current. This opposing trend of thought is due to the fact that many biologists are content with teleological explanations, unsupported by proof; it is also closely connected with the fact that many authors estimate the importance of natural selection less highly than Darwin did. We may describe the objections which are based on the widespread occurrence of self- fertilisation and geitonogamy as of little importance. Darwin did not deny the occurrence of self-fertilisation, even for a long series of generations; his law states only that “Nature abhors PERPETUAL self- fertilisation.” (It is impossible (as has been attempted) to express Darwin’s point of view in a single sentence, such as H. Muller’s statement of the “Knight-Darwin law.” The conditions of life in organisms are so various and complex that laws, such as are formulated in physics and chemistry, can hardly be conceived.) An exception to this rule would therefore occur only in the case of plants in which the possibility of cross-pollination is excluded. Some of the plants with cleistogamous flowers might afford examples of such cases. We have already seen, however, that such a case has not as yet been shown to occur. Burck believed that he had found an instance in certain tropical plants (Anonaceae, Myrmecodia) of the complete exclusion of cross-fertilisation. The flowers of these plants, in which, however,–in contrast to the cleistogamous flowers–the corolla is well developed, remain closed and fruit is produced.

Loew (E. Loew, “Bemerkungen zu Burck…”, “Biolog. Centralbl.” XXVI. (1906).) has shown that cases occur in which cross-fertilisation may be effected even in these “cleistopetalous” flowers: humming birds visit the permanently closed flowers of certain species of Nidularium and transport the pollen. The fact that the formation of hybrids may occur as the result of this shows that pollination may be accomplished.

The existence of plants for which self-pollination is of greater importance than it is for others is by no means contradictory to Darwin’s view. Self- fertilisation is, for example, of greater importance for annuals than for perennials as without it seeds might fail to be produced. Even in the case of annual plants with small inconspicuous flowers in which self- fertilisation usually occurs, such as Senecio vulgaris, Capsella bursa- pastoris and Stellaria media, A. Bateson (Anna Bateson, “The effects of cross-fertilisation on inconspicuous flowers”, “Annals of Botany”, Vol. I. 1888, page 255.) found that cross-fertilisation gave a beneficial result, although only in a slight degree. If the favourable effects of sexual reproduction, according to Darwin’s view, are correlated with change of environment, it is quite possible that this is of less importance in plants which die after ripening their seeds (“hapaxanthic”) and which in any case constantly change their situation. Objections which are based on the proof of the prevalence of self-fertilisation are not, therefore, pertinent. At first sight another point of view, which has been more recently urged, appears to have more weight.

W. Burck (Burck, “Darwin’s Kreuzeungsgesetz…”, “Biol. Centralbl”. XXVIII. 1908, page 177.) has expressed the opinion that the beneficial results of cross-fertilisation demonstrated by Darwin concern only hybrid plants. These alone become weaker by self-pollination; while pure species derive no advantage from crossing and no disadvantage from self-fertilisation. It is certain that some of the plants used by Darwin were of hybrid origin. (It is questionable if this was always the case.) This is evident from his statements, which are models of clearness and precision; he says that his Ipomoea plants “were probably the offspring of a cross.” (“Cross and Self fertilisation” (1st edition), page 55.) The fixed forms of this plant, such as Hero, which was produced by self-fertilisation, and a form of Mimulus with white flowers spotted with red probably resulted from splitting of the hybrids. It is true that the phenomena observed in self- pollination, e.g. in Ipomoea, agree with those which are often noticed in hybrids; Darwin himself drew attention to this.

Let us next call to mind some of the peculiarities connected with hybridisation. We know that hybrids are often characterized by their large size, rapidity of growth, earlier production of flowers, wealth of flower- production and a longer life; hybrids, if crossed with one of the two parent forms, are usually more fertile than when they are crossed together or with another hybrid. But the characters which hybrids exhibit on self- fertilisation are rather variable. The following instance may be quoted from Gartner: “There are many hybrids which retain the self-fertility of the first generation during the second and later generations, but very often in a less degree; a considerable number, however, become sterile.” But the hybrid varieties may be more fertile in the second generation than in the first, and in some hybrids the fertility with their own pollen increases in the second, third, and following generations. (K.F. Gartner, “Versuche uber die Bastarderzeugung”, Stuttgart, 1849, page 149.) As yet it is impossible to lay down rules of general application for the self- fertility of hybrids. That the beneficial influence of crossing with a fresh stock rests on the same ground–a union of sexual cells possessing somewhat different characters–as the fact that many hybrids are distinguished by greater luxuriance, wealth of flowers, etc. corresponds entirely with Darwin’s conclusions. It seems to me to follow clearly from his investigations that there is no essential difference between cross- fertilisation and hybridisation. The heterostyled plants are normally dependent on a process corresponding to hybridisation. The view that specifically distinct species could at best produce sterile hybrids was always opposed by Darwin. But if the good results of crossing were EXCLUSIVELY dependent on the fact that we are concerned with hybrids, there must then be a demonstration of two distinct things. First, that crossing with a fresh stock belonging to the same systematic entity or to the same hybrid, but cultivated for a considerable time under different conditions, shows no superiority over self-fertilisation, and that in pure species crossing gives no better results than self-pollination. If this were the case, we should be better able to understand why in one plant crossing is advantageous while in others, such as Darwin’s Hero and the forms of Mimulus and Nicotiana no advantage is gained; these would then be pure species. But such a proof has not been supplied; the inference drawn from cleistogamous and cleistopetalous plants is not supported by evidence, and the experiments on geitonogamy and on the advantage of cross-fertilisation in species which are usually self-fertilised are opposed to this view. There are still but few researches on this point; Darwin found that in Ononis minutissima, which produces cleistogamous as well as self-fertile chasmogamous flowers, the crossed and self-fertilised capsules produced seed in the proportion of 100:65 and that the average bore the proportion 100:86. Facts previously mentioned are also applicable to this case. Further, it is certain that the self-sterility exhibited by many plants has nothing to do with hybridisation. Between self-sterility and reduced fertility as the result of self-fertilisation there is probably no fundamental difference.

It is certain that so difficult a problem as that of the significance of sexual reproduction requires much more investigation. Darwin was anything but dogmatic and always ready to alter an opinion when it was not based on definite proof: he wrote, “But the veil of secrecy is as yet far from lifted; nor will it be, until we can say why it is beneficial that the sexual elements should be differentiated to a certain extent, and why, if the differentiation be carried still further, injury follows.” He has also shown us the way along which to follow up this problem; it is that of carefully planned and exact experimental research. It may be that eventually many things will be viewed in a different light, but Darwin’s investigations will always form the foundation of Floral Biology on which the future may continue to build.



In developing his conception of organic evolution Charles Darwin was of necessity brought into contact with some of the problems of mental evolution. In “The Origin of Species” he devoted a chapter to “the diversities of instinct and of the other mental faculties in animals of the same class.” (“Origin of Species” (6th edition), page 205.) When he passed to the detailed consideration of “The Descent of Man”, it was part of his object to show “that there is no fundamental difference between man and the higher mammals in their mental faculties.” (“Descent of Man” (2nd edition 1888), Vol. I. page 99; Popular edition page 99.) “If no organic being excepting man,” he said, “had possessed any mental power, or if his powers had been of a wholly different nature from those of the lower animals, then we should never have been able to convince ourselves that our high faculties had been gradually developed.” (Ibid. page 99.) In his discussion of “The Expression of the Emotions” it was important for his purpose “fully to recognise that actions readily become associated with other actions and with various states of the mind.” (“The Expression of the Emotions” (2nd edition), page 32.) His hypothesis of sexual selection is largely dependent upon the exercise of choice on the part of the female and her preference for “not only the more attractive but at the same time the more vigorous and victorious males.” (“Descent of Man”, Vol. II. page 435.) Mental processes and physiological processes were for Darwin closely correlated; and he accepted the conclusion “that the nervous system not only regulates most of the existing functions of the body, but has indirectly influenced the progressive development of various bodily structures and of certain mental qualities.” (Ibid. pages 437, 438.)

Throughout his treatment, mental evolution was for Darwin incidental to and contributory to organic evolution. For specialised research in comparative and genetic psychology, as an independent field of investigation, he had neither the time nor the requisite training. None the less his writings and the spirit of his work have exercised a profound influence on this department of evolutionary thought. And, for those who follow Darwin’s lead, mental evolution is still in a measure subservient to organic evolution. Mental processes are the accompaniments or concomitants of the functional activity of specially differentiated parts of the organism. They are in some way dependent on physiological and physical conditions. But though they are not physical in their nature, and though it is difficult or impossible to conceive that they are physical in their origin, they are, for Darwin and his followers, factors in the evolutionary process in its physical or organic aspect. By the physiologist within his special and well-defined universe of discourse they may be properly regarded as epiphenomena; but by the naturalist in his more catholic survey of nature they cannot be so regarded, and were not so regarded by Darwin. Intelligence has contributed to evolution of which it is in a sense a product.

The facts of observation or of inference which Darwin accepted are these: Conscious experience accompanies some of the modes of animal behaviour; it is concomitant with certain physiological processes; these processes are the outcome of development in the individual and evolution in the race; the accompanying mental processes undergo a like development. Into the subtle philosophical questions which arise out of the naive acceptance of such a creed it was not Darwin’s province to enter; “I have nothing to do,” he said (“Origin of Species” (6th edition), page 205.), “with the origin of the mental powers, any more than I have with that of life itself.” He dealt with the natural history of organisms, including not only their structure but their modes of behaviour; with the natural history of the states of consciousness which accompany some of their actions; and with the relation of behaviour to experience. We will endeavour to follow Darwin in his modesty and candour in making no pretence to give ultimate explanations. But we must note one of the implications of this self- denying ordinance of science. Development and evolution imply continuity. For Darwin and his followers the continuity is organic through physical heredity. Apart from speculative hypothesis, legitimate enough in its proper place but here out of court, we know nothing of continuity of mental evolution as such: consciousness appears afresh in each succeeding generation. Hence it is that for those who follow Darwin’s lead, mental evolution is and must ever be, within his universe of discourse, subservient to organic evolution. Only in so far as conscious experience, or its neural correlate, effects some changes in organic structure can it influence the course of heredity; and conversely only in so far as changes in organic structure are transmitted through heredity, is mental evolution rendered possible. Such is the logical outcome of Darwin’s teaching.

Those who abide by the cardinal results of this teaching are bound to regard all behaviour as the expression of the functional activities of the living tissues of the organism, and all conscious experience as correlated with such activities. For the purposes of scientific treatment, mental processes are one mode of expression of the same changes of which the physiological processes accompanying behaviour are another mode of expression. This is simply accepted as a fact which others may seek to explain. The behaviour itself is the adaptive application of the energies of the organism; it is called forth by some form of presentation or stimulation brought to bear on the organism by the environment. This presentation is always an individual or personal matter. But in order that the organism may be fitted to respond to the presentation of the environment it must have undergone in some way a suitable preparation. According to the theory of evolution this preparation is primarily racial and is transmitted through heredity. Darwin’s main thesis was that the method of preparation is predominantly by natural selection. Subordinate to racial preparation, and always dependent thereon, is individual or personal preparation through some kind of acquisition; of which the guidance of behaviour through individually won experience is a typical example. We here introduce the mental factor because the facts seem to justify the inference. Thus there are some modes of behaviour which are wholly and solely dependent upon inherited racial preparation; there are other modes of behaviour which are also dependent, in part at least, on individual preparation. In the former case the behaviour is adaptive on the first occurrence of the appropriate presentation; in the latter case accommodation to circumstances is only reached after a greater or less amount of acquired organic modification of structure, often accompanied (as we assume) in the higher animals by acquired experience. Logically and biologically the two classes of behaviour are clearly distinguishable: but the analysis of complex cases of behaviour where the two factors cooperate, is difficult and requires careful and critical study of life-history.

The foundations of the mental life are laid in the conscious experience that accompanies those modes of behaviour, dependent entirely on racial preparation, which may broadly be described as instinctive. In the eighth chapter of “The Origin of Species” Darwin says (“Origin of Species” (6th edition), page 205.), “I will not attempt any definition of instinct…Every one understands what is meant, when it is said that instinct impels the cuckoo to migrate and to lay her eggs in other birds’ nests. An action, which we ourselves require experience to enable us to perform, when performed by an animal, more especially by a very young one, without experience, and when performed by many individuals in the same way, without their knowing for what purpose it is performed, is usually said to be instinctive.” And in the summary at the close of the chapter he says (“Origin of Species” (6th edition), page 233.), “I have endeavoured briefly to show that the mental qualities of our domestic animals vary, and that the variations are inherited. Still more briefly I have attempted to show that instincts vary slightly in a state of nature. No one will dispute that instincts are of the highest importance to each animal. Therefore there is no real difficulty, under changing conditions of life, in natural selection accumulating to any extent slight modifications of instinct which are in any way useful. In many cases habit or use and disuse have probably come into play.”

Into the details of Darwin’s treatment there is neither space nor need to enter. There are some ambiguous passages; but it may be said that for him, as for his followers to-day, instinctive behaviour is wholly the result of racial preparation transmitted through organic heredity. For the performance of the instinctive act no individual preparation under the guidance of personal experience is necessary. It is true that Darwin quotes with approval Huber’s saying that “a little dose of judgment or reason often comes into play, even with animals low in the scale of nature.” (Ibid. page 205.) But we may fairly interpret his meaning to be that in behaviour, which is commonly called instinctive, some element of intelligent guidance is often combined. If this be conceded the strictly instinctive performance (or part of the performance) is the outcome of heredity and due to the direct transmission of parental or ancestral aptitudes. Hence the instinctive response as such depends entirely on how the nervous mechanism has been built up through heredity; while intelligent behaviour, or the intelligent factor in behaviour, depends also on how the nervous mechanism has been modified and moulded by use during its development and concurrently with the growth of individual experience in the customary situations of daily life. Of course it is essential to the Darwinian thesis that what Sir E. Ray Lankester has termed “educability,” not less than instinct, is hereditary. But it is also essential to the understanding of this thesis that the differentiae of the hereditary factors should be clearly grasped.

For Darwin there were two modes of racial preparation, (1) natural selection, and (2) the establishment of individually acquired habit. He showed that instincts are subject to hereditary variation; he saw that instincts are also subject to modification through acquisition in the course of individual life. He believed that not only the variations but also, to some extent, the modifications are inherited. He therefore held that some instincts (the greater number) are due to natural selection but that others (less numerous) are due, or partly due, to the inheritance of acquired habits. The latter involve Lamarckian inheritance, which of late years has been the centre of so much controversy. It is noteworthy however that Darwin laid especial emphasis on the fact that many of the most typical and also the most complex instincts–those of neuter insects–do not admit of such an interpretation. “I am surprised,” he says (“Origin of Species” (6th edition), page 233.), “that no one has hitherto advanced this demonstrative case of neuter insects, against the well-known doctrine of inherited habit, as advanced by Lamarck.” None the less Darwin admitted this doctrine as supplementary to that which was more distinctively his own–for example in the case of the instincts of domesticated animals. Still, even in such cases, “it may be doubted,” he says (Ibid. pages 210, 211.), “whether any one would have thought of training a dog to point, had not some one dog naturally shown a tendency in this line…so that habit and some degree of selection have probably concurred in civilising by inheritance our dogs.” But in the interpretation of the instincts of domesticated animals, a more recently suggested hypothesis, that of organic selection (Independently suggested, on somewhat different lines, by Profs. J. Mark Baldwin, Henry F. Osborn and the writer.), may be helpful. According to this hypothesis any intelligent modification of behaviour which is subject to selection is probably coincident in direction with an inherited tendency to behave in this fashion. Hence in such behaviour there are two factors: (1) an incipient variation in the line of such behaviour, and (2) an acquired modification by which the behaviour is carried further along the same line. Under natural selection those organisms in which the two factors cooperate are likely to survive. Under artificial selection they are deliberately chosen out from among the rest.

Organic selection has been termed a compromise between the more strictly Darwinian and the Lamarckian principles of interpretation. But it is not in any sense a compromise. The principle of interpretation of that which is instinctive and hereditary is wholly Darwinian. It is true that some of the facts of observation relied upon by Lamarckians are introduced. For Lamarckians however the modifications which are admittedly factors in survival, are regarded as the parents of inherited variations; for believers in organic selection they are only the foster parents or nurses. It is because organic selection is the direct outcome of and a natural extension of Darwin’s cardinal thesis that some reference to it here is justifiable. The matter may be put with the utmost brevity as follows. (1) Variations (V) occur, some of which are in the direction of increased adaptation (+), others in the direction of decreased adaptation (-). (2) Acquired modifications (M) also occur. Some of these are in the direction of increased accommodation to circumstances (+), while others are in the direction of diminished accommodation (-). Four major combinations are

(a) + V with + M,
(b) + V with – M,
(c) – V with + M,
(d) – V with – M.

Of these (d) must inevitably be eliminated while (a) are selected. The predominant survival of (a) entails the survival of the adaptive variations which are inherited. The contributory acquisitions (+M) are not inherited; but they are none the less factors in determining the survival of the coincident variations. It is surely abundantly clear that this is Darwinism and has no tincture of Lamarck’s essential principle, the inheritance of acquired characters.

Whether Darwin himself would have accepted this interpretation of some at least of the evidence put forward by Lamarckians is unfortunately a matter of conjecture. The fact remains that in his interpretation of instinct and in allied questions he accepted the inheritance of individually acquired modifications of behaviour and structure.

Darwin was chiefly concerned with instinct from the biological rather than from the psychological point of view. Indeed it must be confessed that, from the latter standpoint, his conception of instinct as a “mental faculty” which “impels” an animal to the performance of certain actions, scarcely affords a satisfactory basis for genetic treatment. To carry out the spirit of Darwin’s teaching it is necessary to link more closely biological and psychological evolution. The first step towards this is to interpret the phenomena of instinctive behaviour in terms of stimulation and response. It may be well to take a particular case. Swimming on the part of a duckling is, from the biological point of view, a typical example of instinctive behaviour. Gently lower a recently hatched bird into water: coordinated movements of the limbs follow in rhythmical sequence. The behaviour is new to the individual though it is no doubt closely related to that of walking, which is no less instinctive. There is a group of stimuli afforded by the “presentation” which results from partial immersion: upon this there follows as a complex response an application of the functional activities in swimming; the sequence of adaptive application on the appropriate presentation is determined by racial preparation. We know, it is true, but little of the physiological details of what takes place in the central nervous system; but in broad outline the nature of the organic mechanism and the manner of its functioning may at least be provisionally conjectured in the present state of physiological knowledge. Similarly in the case of the pecking of newly-hatched chicks; there is a visual presentation, there is probably a cooperating group of stimuli from the alimentary tract in need of food, there is an adaptive application of the activities in a definite mode of behaviour. Like data are afforded in a great number of cases of instinctive procedure, sometimes occurring very early in life, not infrequently deferred until the organism is more fully developed, but all of them dependent upon racial preparation. No doubt there is some range of variation in the behaviour, just such variation as the theory of natural selection demands. But there can be no question that the higher animals inherit a bodily organisation and a nervous system, the functional working of which gives rise to those inherited modes of behaviour which are termed instinctive.

It is to be noted that the term “instinctive” is here employed in the adjectival form as a descriptive heading under which may be grouped many and various modes of behaviour due to racial preparation. We speak of these as inherited; but in strictness what is transmitted through heredity is the complex of anatomical and physiological conditions under which, in appropriate circumstances, the organism so behaves. So far the term “instinctive” has a restricted biological connotation in terms of behaviour. But the connecting link between biological evolution and psychological evolution is to be sought,–as Darwin fully realised,–in the phenomena of instinct, broadly considered. The term “instinctive” has also a psychological connotation. What is that connotation?

Let us take the case of the swimming duckling or the pecking chick, and fix our attention on the first instinctive performance. Grant that just as there is, strictly speaking, no inherited behaviour, but only the conditions which render such behaviour under appropriate circumstances possible; so too there is no inherited experience, but only the conditions which render such experience possible; then the cerebral conditions in both cases are the same. The biological behaviour-complex, including the total stimulation and the total response with the intervening or resultant processes in the sensorium, is accompanied by an experience-complex including the initial stimulation-consciousness and resulting response- consciousness. In the experience-complex are comprised data which in psychological analysis are grouped under the headings of cognition, affective tone and conation. But the complex is probably experienced as an unanalysed whole. If then we use the term “instinctive” so as to comprise all congenital modes of behaviour which contribute to experience, we are in a position to grasp the view that the net result in consciousness constitutes what we may term the primary tissue of experience. To the development of this experience each instinctive act contributes. The nature and manner of organisation of this primary tissue of experience are dependent on inherited biological aptitudes; but they are from the outset onwards subject to secondary development dependent on acquired aptitudes. Biological values are supplemented by psychological values in terms of satisfaction or the reverse.

In our study of instinct we have to select some particular phase of animal behaviour and isolate it so far as is possible from the life of which it is a part. But the animal is a going concern, restlessly active in many ways. Many instinctive performances, as Darwin pointed out (“Origin of Species” (6th edition), page 206.), are serial in their nature. But the whole of active life is a serial and coordinated business. The particular instinctive performance is only an episode in a life-history, and every mode of behaviour is more or less closely correlated with other modes. This coordination of behaviour is accompanied by a correlation of the modes of primary experience. We may classify the instinctive modes of behaviour and their accompanying modes of instinctive experience under as many heads as may be convenient for our purposes of interpretation, and label them instincts of self-preservation, of pugnacity, of acquisition, the reproductive instincts, the parental instincts, and so forth. An instinct, in this sense of the term (for example the parental instinct), may be described as a specialised part of the primary tissue of experience differentiated in relation to some definite biological end. Under such an instinct will fall a large number of particular and often well-defined modes of behaviour, each with its own peculiar mode of experience.

It is no doubt exceedingly difficult as a matter of observation and of inference securely based thereon to distinguish what is primary from what is in part due to secondary acquisition–a fact which Darwin fully appreciated. Animals are educable in different degrees; but where they are educable they begin to profit by experience from the first. Only, therefore, on the occasion of the first instinctive act of a given type can the experience gained be weighed as WHOLLY primary; all subsequent performance is liable to be in some degree, sometimes more, sometimes less, modified by the acquired disposition which the initial behaviour engenders. But the early stages of acquisition are always along the lines predetermined by instinctive differentiation. It is the task of comparative psychology to distinguish the primary tissue of experience from its secondary and acquired modifications. We cannot follow up the matter in further detail. It must here suffice to suggest that this conception of instinct as a primary form of experience lends itself better to natural history treatment than Darwin’s conception of an impelling force, and that it is in line with the main trend of Darwin’s thought.

In a characteristic work,–characteristic in wealth of detail, in closeness and fidelity of observation, in breadth of outlook, in candour and modesty,–Darwin dealt with “The Expression of the Emotions in Man and Animals”. Sir Charles Bell in his “Anatomy of Expression” had contended that many of man’s facial muscles had been specially created for the sole purpose of being instrumental in the expression of his emotions. Darwin claimed that a natural explanation, consistent with the doctrine of evolution, could in many cases be given and would in other cases be afforded by an extension of the principles he advocated. “No doubt,” he said (“Expression of the Emotions”, page 13. The passage is here somewhat condensed.), “as long as man and all other animals are viewed as independent creations, an effectual stop is put to our natural desire to investigate as far as possible the causes of Expression. By this doctrine, anything and everything can be equally well explained…With mankind, some expressions…can hardly be understood, except on the belief that man once existed in a much lower and animal-like condition. The community of certain expressions in distinct though allied species…is rendered somewhat more intelligible, if we believe in their descent from a common progenitor. He who admits on general grounds that the structure and habits of all animals have been gradually evolved, will look at the whole subject of Expression in a new and interesting light.”

Darwin relied on three principles of explanation. “The first of these principles is, that movements which are serviceable in gratifying some desire, or in relieving some sensation, if often repeated, become so habitual that they are performed, whether or not of any service, whenever the same desire or sensation is felt, even in a very weak degree.” (Ibid. page 368.) The modes of expression which fall under this head have become instinctive through the hereditary transmission of acquired habit. “As far as we can judge, only a few expressive movements are learnt by each individual; that is, were consciously and voluntarily performed during the early years of life for some definite object, or in imitation of others, and then became habitual. The far greater number of the movements of expression, and all the more important ones, are innate or inherited; and such cannot be said to depend on the will of the individual. Nevertheless, all those included under our first principle were at first voluntarily performed for a definite object,–namely, to escape some danger, to relieve some distress, or to gratify some desire.” (Ibid. pages 373, 374.)

“Our second principle is that of antithesis. The habit of voluntarily performing opposite movements under opposite impulses has become firmly established in us by the practice of our whole lives. Hence, if certain actions have been regularly performed, in accordance with our first principle, under a certain frame of mind, there will be a strong and involuntary tendency to the performance of directly opposite actions, whether or not these are of any use, under the excitement of an opposite frame of mind.” (“Expression of the Emotions”, page 368.) This principle of antithesis has not been widely accepted. Nor is Darwin’s own position easy to grasp.

“Our third principle,” he says (Ibid. page 369.), “is the direct action of the excited nervous system on the body, independently of the will, and independently, in large part, of habit. Experience shows that nerve-force is generated and set free whenever the cerebro-spinal system is excited. The direction which this nerve-force follows is necessarily determined by the lines of connection between the nerve-cells, with each other and with various parts of the body.”

Lack of space prevents our following up the details of Darwin’s treatment of expression. Whether we accept or do not accept his three principles of explanation we must regard his work as a masterpiece of descriptive analysis, packed full of observations possessing lasting value. For a further development of the subject it is essential that the instinctive factors in expression should be more fully distinguished from those which are individually acquired–a difficult task–and that the instinctive factors should be rediscussed in the light of modern doctrines of heredity, with a view to determining whether Lamarckian inheritance, on which Darwin so largely relied, is necessary for an interpretation of the facts.

The whole subject as Darwin realised is very complex. Even the term “expression” has a certain amount of ambiguity. When the emotion is in full flood the animal fights, flees, or faints. Is this full-tide effect to be regarded as expression; or are we to restrict the term to the premonitory or residual effects–the bared canine when the fighting mood is being roused, the ruffled fur when reminiscent representations of the object inducing anger cross the mind? Broadly considered both should be included. The activity of premonitory expression as a means of communication was recognised by Darwin; he might, perhaps, have emphasised it more strongly in dealing with the lower animals. Man so largely relies on a special means of communication, that of language, that he sometimes fails to realise that for animals with their keen powers of perception, and dependent as they are on such means of communication, the more strictly biological means of expression are full of subtle suggestiveness. Many modes of expression, otherwise useless, are signs of behaviour that may be anticipated,–signs which stimulate the appropriate attitude of response. This would not, however, serve to account for the utility of the organic accompaniments–heart-affection, respiratory changes, vaso-motor effects and so forth, together with heightened muscular tone,–on all of which Darwin lays stress (“Expression of the Emotions”, pages 65 ff.) under his third principle. The biological value of all this is, however, of great importance, though Darwin was hardly in a position to take it fully into account.

Having regard to the instinctive and hereditary factors of emotional expression we may ask whether Darwin’s third principle does not alone suffice as an explanation. Whether we admit or reject Lamarckian inheritance it would appear that all hereditary expression must be due to pre-established connections within the central nervous system and to a transmitted provision for coordinated response under the appropriate stimulation. If this be so, Darwin’s first and second principles are subordinate and ancillary to the third, an expression, so far as it is instinctive or hereditary, being “the direct result of the constitution of the nervous system.”

Darwin accepted the emotions themselves as hereditary or acquired states of mind and devoted his attention to their expression. But these emotions themselves are genetic products and as such dependent on organic conditions. It remained, therefore, for psychologists who accepted evolution and sought to build on biological foundations to trace the genesis of these modes of animal and human experience. The subject has been independently developed by Professors Lange and James (Cf. William James, “Principles of Psychology”, Vol. II. Chap. XXV, London, 1890.); and some modification of their view is regarded by many evolutionists as affording the best explanation of the facts. We must fix our attention on the lower emotions, such as anger or fear, and on their first occurrence in the life of the individual organism. It is a matter of observation that if a group of young birds which have been hatched in an incubator are frightened by an appropriate presentation, auditory or visual, they instinctively respond in special ways. If we speak of this response as the expression, we find that there are many factors. There are certain visible modes of behaviour, crouching at once, scattering and then crouching, remaining motionless, the braced muscles sustaining an attitude of arrest, and so forth. There are also certain visceral or organic effects, such as affections of the heart and respiration. These can be readily observed by taking the young bird in the hand. Other effects cannot be readily observed; vaso-motor changes, affections of the alimentary canal, the skin and so forth. Now the essence of the James-Lange view, as applied to these congenital effects, is that though we are justified in speaking of them as effects of the stimulation, we are not justified, without further evidence, in speaking of them as effects of the emotional state. May it not rather be that the emotion as a primary mode of experience is the concomitant of the net result of the organic situation–the initial presentation, the instinctive mode of behaviour, the visceral disturbances? According to this interpretation the primary tissue of experience of the emotional order, felt as an unanalysed complex, is generated by the stimulation of the sensorium by afferent or incoming physiological impulses from the special senses, from the organs concerned in the responsive behaviour, from the viscera and vaso-motor system.

Some psychologists, however, contend that the emotional experience is generated in the sensorium prior to, and not subsequent to, the behaviour- response and the visceral disturbances. It is a direct and not an indirect outcome of the presentation to the special senses. Be this as it may, there is a growing tendency to bring into the closest possible relation, or even to identify, instinct and emotion in their primary genesis. The central core of all such interpretations is that instinctive behaviour and experience, its emotional accompaniments, and its expression, are but different aspects of the outcome of the same organic occurrences. Such emotions are, therefore, only a distinguishable aspect of the primary tissue of experience and exhibit a like differentiation. Here again a biological foundation is laid for a psychological doctrine of the mental development of the individual.

The intimate relation between emotion as a psychological mode of experience and expression as a group of organic conditions has an important bearing on biological interpretation. The emotion, as the psychological accompaniment of orderly disturbances in the central nervous system profoundly influences behaviour and often renders it more vigorous and more effective. The utility of the emotions in the struggle for existence can, therefore, scarcely be over-estimated. Just as keenness of perception has survival- value; just as it is obviously subject to variation; just as it must be enhanced under natural selection, whether individually acquired increments are inherited or not; and just as its value lies not only in this or that special perceptive act but in its importance for life as a whole; so the vigorous effectiveness of activity has survival-value; it is subject to variation; it must be enhanced under natural selection; and its importance lies not only in particular modes of behaviour but in its value for life as a whole. If emotion and its expression as a congenital endowment are but different aspects of the same biological occurrence; and if this is a powerful supplement to vigour effectiveness and persistency of behaviour, it must on Darwin’s principles be subject to natural selection.

If we include under the expression of the emotions not only the premonitory symptoms of the initial phases of the organic and mental state, not only the signs or conditions of half-tide emotion, but the full-tide manifestation of an emotion which dominates the situation, we are naturally led on to the consideration of many of the phenomena which are discussed under the head of sexual selection. The subject is difficult and complex, and it was treated by Darwin with all the strength he could summon to the task. It can only be dealt with here from a special point of view–that which may serve to illustrate the influence of certain mental factors on the course of evolution. From this point of view too much stress can scarcely be laid on the dominance of emotion during the period of courtship and pairing in the more highly organised animals. It is a period of maximum vigour, maximum activity, and, correlated with special modes of behaviour and special organic and visceral accompaniments, a period also of maximum emotional excitement. The combats of males, their dances and aerial evolutions, their elaborate behaviour and display, or the flood of song in birds, are emotional expressions which are at any rate coincident in time with sexual periodicity. From the combat of the males there follows on Darwin’s principles the elimination of those which are deficient in bodily vigour, deficient in special structures, offensive or protective, which contribute to success, deficient in the emotional supplement of which persistent and whole-hearted fighting is the expression, and deficient in alertness and skill which are the outcome of the psychological development of the powers of perception. Few biologists question that we have here a mode of selection of much importance, though its influence on psychological evolution often fails to receive its due emphasis. Mr Wallace (“Darwinism”, pages 282, 283, London, 1889.) regards it as “a form of natural selection”; “to it,” he says, “we must impute the development of the exceptional strength, size, and activity of the male, together with the possession of special offensive and defensive weapons, and of all other characters which arise from the development of these or are correlated with them.” So far there is little disagreement among the followers of Darwin– for Mr Wallace, with fine magnanimity, has always preferred to be ranked as such, notwithstanding his right, on which a smaller man would have constantly insisted, to the claim of independent originator of the doctrine of natural selection. So far with regard to sexual selection Darwin and Mr Wallace are agreed; so far and no farther. For Darwin, says Mr Wallace (Ibid. page 283.), “has extended the principle into a totally different field of action, which has none of that character of constancy and of inevitable result that attaches to natural selection, including male rivalry; for by far the larger portion of the phenomena, which he endeavours to explain by the direct action of sexual selection, can only be so explained on the hypothesis that the immediate agency is female choice or preference. It is to this that he imputes the origin of all secondary sexual characters other than weapons of offence and defence…In this extension of sexual selection to include the action of female choice or preference, and in the attempt to give to that choice such wide-reaching effects, I am unable to follow him more than a very little way.”

Into the details of Mr Wallace’s criticisms it is impossible to enter here. We cannot discuss either the mode of origin of the variations in structure which have rendered secondary sexual characters possible or the modes of selection other than sexual which have rendered them, within narrow limits, specifically constant. Mendelism and mutation theories may have something to say on the subject when these theories have been more fully correlated with the basal principles of selection. It is noteworthy that Mr Wallace says (“Darwinism”, pages 283, 284.): “Besides the acquisition of weapons by the male for the purpose of fighting with other males, there are some other sexual characters which may have been produced by natural selection. Such are the various sounds and odours which are peculiar to the male, and which serve as a call to the female or as an indication of his presence. These are evidently a valuable addition to the means of recognition of the two sexes, and are a further indication that the pairing season has arrived; and the production, intensification, and differentiation of these sounds and odours are clearly within the power of natural selection. The same remark will apply to the peculiar calls of birds, and even to the singing of the males.” Why the same remark should not apply to their colours and adornments is not obvious. What is obvious is that “means of recognition” and “indication that the pairing season has arrived” are dependent on the perceptive powers of the female who recognises and for whom the indication has meaning. The hypothesis of female preference, stripped of the aesthetic surplusage which is psychologically both unnecessary and unproven, is really only different in degree from that which Mr Wallace admits in principle when he says that it is probable that the female is pleased or excited by the display.

Let us for our present purpose leave on one side and regard as sub judice the question whether the specific details of secondary sexual characters are the outcome of female choice. For us the question is whether certain psychological accompaniments of the pairing situation have influenced the course of evolution and whether these psychological accompaniments are themselves the outcome of evolution. As a matter of observation, specially differentiated modes of behaviour, often very elaborate, frequently requiring highly developed skill, and apparently highly charged with emotional tone, are the precursors of pairing. They are generally confined to the males, whose fierce combats during the period of sexual activity are part of the emotional manifestation. It is inconceivable that they have no biological meaning; and it is difficult to conceive that they have any other biological end than to evoke in the generally more passive female the pairing impulse. They are based on instinctive foundations ingrained in the nervous constitution through natural (or may we not say sexual?) selection in virtue of their profound utility. They are called into play by a specialised presentation such as the sight or the scent of the female at, or a little in advance of, a critical period of the physiological rhythm. There is no necessity that the male should have any knowledge of the end to which his strenuous activity leads up. In presence of the female there is an elaborate application of all the energies of behaviour, just because ages of racial preparation have made him biologically and emotionally what he is–a functionally sexual male that must dance or sing or go through hereditary movements of display, when the appropriate stimulation comes. Of course after the first successful courtship his future behaviour will be in some degree modified by his previous experience. No doubt during his first courtship he is gaining the primary data of a peculiarly rich experience, instinctive and emotional. But the biological foundations of the behaviour of courtship are laid in the hereditary coordinations. It would seem that in some cases, not indeed in all, but perhaps especially in those cases in which secondary sexual behaviour is most highly evolved,–correlative with the ardour of the male is a certain amount of reluctance in the female. The pairing act on her part only takes place after prolonged stimulation, for affording which the behaviour of male courtship is the requisite presentation. The most vigorous, defiant and mettlesome male is preferred just because he alone affords a contributory stimulation adequate to evoke the pairing impulse with its attendant emotional tone.

It is true that this places female preference or choice on a much lower psychological plane than Darwin in some passages seems to contemplate where, for example, he says that the female appreciates the display of the male and places to her credit a taste for the beautiful. But Darwin himself distinctly states (“Descent of Man” (2nd edition), Vol. II. pages 136, 137; (Popular edition), pages 642, 643.) that “it is not probable that she consciously deliberates; but she is most excited or attracted by the most beautiful, or melodious, or gallant males.” The view here put forward, which has been developed by Prof. Groos (“The Play of Animals”, page 244, London, 1898.), therefore seems to have Darwin’s own sanction. The phenomena are not only biological; there are psychological elements as well. One can hardly suppose that the female is unconscious of the male’s presence; the final yielding must surely be accompanied by heightened emotional tone. Whether we call it choice or not is merely a matter of definition of terms. The behaviour is in part determined by supplementary psychological values. Prof. Groos regards the coyness of females as “a most efficient means of preventing the too early and too frequent yielding to the sexual impulse.” (Ibid. page 283.) Be that as it may, it is, in any case, if we grant the facts, a means through which male sexual behaviour with all its biological and psychological implications, is raised to a level otherwise perhaps unattainable by natural means, while in the female it affords opportunities for the development in the individual and evolution in the race of what we may follow Darwin in calling appreciation, if we empty this word of the aesthetic implications which have gathered round it in the mental life of man.

Regarded from this standpoint sexual selection, broadly considered, has probably been of great importance. The psychological accompaniments of the pairing situation have profoundly influenced the course of biological evolution and are themselves the outcome of that evolution.

Darwin makes only passing reference to those modes of behaviour in animals which go by the name of play. “Nothing,” he says (“Descent of Man”, Vol. II. page 60; (Popular edition), page 566.), “is more common than for animals to take pleasure in practising whatever instinct they follow at other times for some real good.” This is one of the very numerous cases in which a hint of the master has served to stimulate research in his disciples. It was left to Prof. Groos to develop this subject on evolutionary lines and to elaborate in a masterly manner Darwin’s suggestion. “The utility of play,” he says (“The Play of Animals”, page 76.), “is incalculable. This utility consists in the practice and exercise it affords for some of the more important duties of life,”–that is to say, for the performance of activities which will in adult life be essential to survival. He urges (Ibid. page 75.) that “the play of young animals has its origin in the fact that certain very important instincts appear at a time when the animal does not seriously need them.” It is, however, questionable whether any instincts appear at a time when they are not needed. And it is questionable whether the instinctive and emotional attitude of the play-fight, to take one example, can be identified with those which accompany fighting in earnest, though no doubt they are closely related and have some common factors. It is probable that play, as preparatory behaviour, differs in biological detail (as it almost certainly does in emotional attributes) from the earnest of after-life and that it has been evolved through differentiation and integration of the primary tissue of experience, as a preparation through which certain essential modes of skill may be acquired–those animals in which the preparatory play-propensity was not inherited in due force and requisite amount being subsequently eliminated in the struggle for existence. In any case there is little question that Prof. Groos is right in basing the play-propensity on instinctive foundations. (“The Play of Animals” page 24.) None the less, as he contends, the essential biological value of play is that it is a means of training the educable nerve-tissue, of developing that part of the brain which is modified by experience and which thus acquires new characters, of elaborating the secondary tissue of experience on the predetermined lines of instinctive differentiation and thus furthering the psychological activities which are included under the comprehensive term “intelligent.”

In “The Descent of Man” Darwin dealt at some length with intelligence and the higher mental faculties. (“Descent of Man” (1st edition), Chapters II, III, V; (2nd edition), Chapters III, IV, V.) His object, he says, is to show that there is no fundamental difference between man and the higher mammals in their mental faculties; that these faculties are variable and the variations tend to be inherited; and that under natural selection beneficial variations of all kinds will have been preserved and injurious ones eliminated.

Darwin was too good an observer and too honest a man to minimise the “enormous difference” between the level of mental attainment of civilised man and that reached by any animal. His contention was that the difference, great as it is, is one of degree and not of kind. He realised that, in the development of the mental faculties of man, new factors in evolution have supervened–factors which play but a subordinate and subsidiary part in animal intelligence. Intercommunication by means of language, approbation and blame, and all that arises out of reflective thought, are but foreshadowed in the mental life of animals. Still he contends that these may be explained on the doctrine of evolution. He urges (Ibid. Vol. I. pages 70, 71; (Popular edition), pages 70, 71.)” that man is variable in body and mind; and that the variations are induced, either directly or indirectly, by the same general causes, and obey the same general laws, as with the lower animals.” He correlates mental development with the evolution of the brain. (Ibid. page 81.) “As the various mental faculties gradually developed themselves, the brain would almost certainly become larger. No one, I presume, doubts that the large proportion which the size of man’s brain bears to his body, compared to the same proportion in the gorilla or orang, is closely connected with his higher mental powers.” “With respect to the lower animals,” he says (“Descent of Man” (Popular edition), page 82.), “M.E. Lartet (“Comptes Rendus des Sciences”, June 1, 1868.), by comparing the crania of tertiary and recent mammals belonging to the same groups, has come to the remarkable conclusion that the brain is generally larger and the convolutions are more complex in the more recent form.”

Sir E. Ray Lankester has sought to express in the simplest terms the implications of the increase in size of the cerebrum. “In what,” he asks, “does the advantage of a larger cerebral mass consist?” “Man,” he replies “is born with fewer ready-made tricks of the nerve-centres–these performances of an inherited nervous mechanism so often called by the ill- defined term ‘instincts’–than are the monkeys or any other animal. Correlated with the absence of inherited ready-made mechanism, man has a greater capacity of developing in the course of his individual growth similar nervous mechanisms (similar to but not identical with those of ‘instinct’) than any other animal…The power of being educated– ‘educability’ as we may term it–is what man possesses in excess as compared with the apes. I think we are justified in forming the hypothesis that it is this ‘educability’ which is the correlative of the increased size of the cerebrum.” There has been natural selection of the more educable animals, for “the character which we describe as ‘educability’ can be transmitted, it is a congenital character. But the RESULTS of education can NOT be transmitted. In each generation they have to be acquired afresh, and with increased ‘educability’ they are more readily acquired and a larger variety of them…The fact is that there is no community between the mechanisms of instinct and the mechanisms of intelligence, and that the latter are later in the history of the evolution of the brain than the former and can only develop in proportion as the former become feeble and defective.” (“Nature”, Vol. LXI. pages 624, 625 (1900).)

In this statement we have a good example of the further development of views which Darwin foreshadowed but did not thoroughly work out. It states the biological case clearly and tersely. Plasticity of behaviour in special accommodation to special circumstances is of survival value; it depends upon acquired characters; it is correlated with increase in size and complexity of the cerebrum; under natural selection therefore the larger and more complex cerebrum as the organ of plastic behaviour has been the outcome of natural selection. We have thus the biological foundations for a further development of genetic psychology.

There are diversities of opinion, as Darwin showed, with regard to the range of instinct in man and the higher animals as contrasted with lower types. Darwin himself said (“Descent of Man”, Vol. I. page 100.) that “Man, perhaps, has somewhat fewer instincts than those possessed by the animals which come next to him in the series.” On the other hand, Prof. Wm. James says (“Principles of Psychology,” Vol. II. page 289.) that man is probably the animal with most instincts. The true position is that man and the higher animals have fewer complete and self-sufficing instincts than those which stand lower in the scale of mental evolution, but that they have an equally large or perhaps larger mass of instinctive raw material which may furnish the stuff to be elaborated by intelligent processes. There is, perhaps, a greater abundance of the primary tissue of experience to be refashioned and integrated by secondary modification; there is probably the same differentiation in relation to the determining biological ends, but there is at the outset less differentiation of the particular and specific modes of behaviour. The specialised instinctive performances and their concomitant experience-complexes are at the outset more indefinite. Only through acquired connections, correlated with experience, do they become definitely organised.

The full working-out of the delicate and subtle relationship of instinct and educability–that is, of the hereditary and the acquired factors in the mental life–is the task which lies before genetic and comparative psychology. They interact throughout the whole of life, and their interactions are very complex. No one can read the chapters of “The Descent of Man” which Darwin devotes to a consideration of the mental characters of man and animals without noticing, on the one hand, how sedulous he is in his search for hereditary foundations, and, on the other hand, how fully he realises the importance of acquired habits of mind. The fact that educability itself has innate tendencies–is in fact a partially differentiated educability–renders the unravelling of the factors of mental progress all the more difficult.

In his comparison of the mental powers of men and animals it was essential that Darwin should lay stress on points of similarity rather than on points of difference. Seeking to establish a doctrine of evolution, with its basal concept of continuity of process and community of character, he was bound to render clear and to emphasise the contention that the difference in mind between man and the higher animals, great as it is, is one of degree and not of kind. To this end Darwin not only recorded a large number of valuable observations of his own, and collected a considerable body of information from reliable sources, he presented the whole subject in a new light and showed that a natural history of mind might be written and that this method of study offered a wide and rich field for investigation. Of course those who regarded the study of mind only as a branch of metaphysics smiled at the philosophical ineptitude of the mere man of science. But the investigation, on natural history lines, has been prosecuted with a large measure of success. Much indeed still remains to be done; for special training is required, and the workers are still few. Promise for the future is however afforded by the fact that investigation is prosecuted on experimental lines and that something like organised methods of research are taking form. There is now but little reliance on casual observations recorded by those who have not undergone the necessary discipline in these methods. There is also some change of emphasis in formulating conclusions. Now that the general evolutionary thesis is fully and freely accepted by those who carry on such researches, more stress is laid on the differentiation of the stages of evolutionary advance than on the fact of their underlying community of nature. The conceptual intelligence which is especially characteristic of the higher mental procedure of man is more firmly distinguished from the perceptual intelligence which he shares with the lower animals–distinguished now as a higher product of evolution, no longer as differing in origin or different in kind. Some progress has been made, on the one hand in rendering an account of intelligent profiting by experience under the guidance of pleasure and pain in the perceptual field, on lines predetermined by instinctive differentiation for biological ends, and on the other hand in elucidating the method of conceptual thought employed, for example, by the investigator himself in interpreting the perceptual experience of the lower animals.

Thus there is a growing tendency to realise more fully that there are two orders of educability–first an educability of the perceptual intelligence based on the biological foundation of instinct, and secondly an educability of the conceptual intelligence which refashions and rearranges the data afforded by previous inheritance and acquisition. It is in relation to this second and higher order of educability that the cerebrum of man shows so large an increase of mass and a yet larger increase of effective surface through its rich convolutions. It is through educability of this order that the human child is brought intellectually and affectively into touch with the ideal constructions by means of which man has endeavoured, with more or less success, to reach an interpretation of nature, and to guide the course of the further evolution of his race–ideal constructions which form part of man’s environment.

It formed no part of Darwin’s purpose to consider, save in broad outline, the methods, or to discuss in any fulness of detail the results of the process by which a differentiation of the mental faculties of man from those of the lower animals has been brought about–a differentiation the existence of which he again and again acknowledges. His purpose was rather to show that, notwithstanding this differentiation, there is basal community in kind. This must be remembered in considering his treatment of the biological foundations on which man’s systems of ethics are built. He definitely stated that he approached the subject “exclusively from the side of natural history.” (“Descent of Man”, Vol. I. page 149.) His general conclusion is that the moral sense is fundamentally identical with the social instincts, which have been developed for the good of the community; and he suggests that the concept which thus enables us to interpret the biological ground-plan of morals also enables us to frame a rational ideal of the moral end. “As the social instincts,” he says (Ibid. page 185.), “both of man and the lower animals have no doubt been developed by nearly the same steps, it would be advisable, if found practicable, to use the same definition in both cases, and to take as the standard of morality, the general good or welfare of the community, rather than the general happiness.” But the kind of community for the good of which the social instincts of animals and primitive men were biologically developed may be different from that which is the product of civilisation, as Darwin no doubt realised. Darwin’s contention was that conscience is a social instinct and has been evolved because it is useful to the tribe in the struggle for existence against other tribes. On the other hand, J.S. Mill urged that the moral feelings are not innate but acquired, and Bain held the same view, believing that the moral sense is acquired by each individual during his life-time. Darwin, who notes (Ibid. page 150 (footnote).) their opinion with his usual candour, adds that “on the general theory of evolution this is at least extremely improbable. It is impossible to enter into the question here: much turns on the exact connotation of the terms “conscience” and “moral sense,” and on the meaning we attach to the statement that the moral sense is fundamentally identical with the social instincts.

Presumably the majority of those who approach the subjects discussed in the third, fourth and fifth chapters of “The Descent of Man” in the full conviction that mental phenomena, not less than organic phenomena, have a natural genesis, would, without hesitation, admit that the intellectual and moral systems of civilised man are ideal constructions, the products of conceptual thought, and that as such they are, in their developed form, acquired. The moral sentiments are the emotional analogues of highly developed concepts. This does not however imply that they are outside the range of natural history treatment. Even though it may be desirable to differentiate the moral conduct of men from the social behaviour of animals (to which some such term as “pre-moral” or “quasi-moral” may be applied), still the fact remains that, as Darwin showed, there is abundant evidence of the occurrence of such social behaviour–social behaviour which, even granted that it is in large part intelligently acquired, and is itself so far a product of educability, is of survival value. It makes for that integration without which no social group could hold together and escape elimination. Furthermore, even if we grant that such behaviour is intelligently acquired, that is to say arises through the modification of hereditary instincts and emotions, the fact remains that only through these instinctive and emotional data is afforded the primary tissue of the experience which is susceptible of such modification.

Darwin sought to show, and succeeded in showing, that for the intellectual and moral life there are instinctive foundations which a biological treatment alone can disclose. It is true that he did not in all cases analytically distinguish the foundations from the superstructure. Even to- day we are scarcely in a position to do so adequately. But his treatment was of great value in giving an impetus to further research. This value indeed can scarcely be overestimated. And when the natural history of the mental operations shall have been written, the cardinal fact will stand forth, that the instinctive and emotional foundations are the outcome of biological evolution and have been ingrained in the race through natural selection. We shall more clearly realise that educability itself is a product of natural selection, though the specific results acquired through cerebral modifications are not transmitted through heredity. It will, perhaps, also be realised that the instinctive foundations of social behaviour are, for us, somewhat out of date and have undergone but little change throughout the progress of civilisation, because natural selection has long since ceased to be the dominant factor in human progress. The history of human progress has been mainly the history of man’s higher educability, the products of which he has projected on to his environment. This educability remains on the average what it was a dozen generations ago; but the thought-woven tapestry of his surroundings is refashioned and improved by each succeeding generation. Few men have in greater measure enriched the thought-environment with which it is the aim of education to bring educable human beings into vital contact, than has Charles Darwin. His special field of work was the wide province of biology; but he did much to help us realise that mental factors have contributed to organic evolution and that in man, the highest product of Evolution, they have reached a position of unquestioned supremacy.


Professor of Philosophy in the University of Copenhagen.


It is difficult to draw a sharp line between philosophy and natural science. The naturalist who introduces a new principle, or demonstrates a fact which throws a new light on existence, not only renders an important service to philosophy but is himself a philosopher in the broader sense of the word. The aim of philosophy in the stricter sense is to attain points of view from which the fundamental phenomena and the principles of the special sciences can be seen in their relative importance and connection. But philosophy in this stricter sense has always been influenced by philosophy in the broader sense. Greek philosophy came under the influence of logic and mathematics, modern philosophy under the influence of natural science. The name of Charles Darwin stands with those of Galileo, Newton, and Robert Mayer–names which denote new problems and great alterations in our conception of the universe.

First of all we must lay stress on Darwin’s own personality. His deep love of truth, his indefatigable inquiry, his wide horizon, and his steady self- criticism make him a scientific model, even if his results and theories should eventually come to possess mainly an historical interest. In the intellectual domain the primary object is to reach high summits from which wide surveys are possible, to reach them toiling honestly upwards by way of experience, and then not to turn dizzy when a summit is gained. Darwinians have sometimes turned dizzy, but Darwin never. He saw from the first the great importance of his hypothesis, not only because of its solution of the old problem as to the value of the concept of species, not only because of the grand picture of natural evolution which it unrolls, but also because of the life and inspiration its method would impart to the study of comparative anatomy, of instinct and of heredity, and finally because of the influence it would exert on the whole conception of existence. He wrote in his note-book in the year 1837: “My theory would give zest to recent and fossil comparative anatomy; it would lead to the study of instinct, heredity, and mind-heredity, whole (of) metaphysics.” (“Life and Letters of Charles Darwin”, Vol. I. page 8.)

We can distinguish four main points in which Darwin’s investigations possess philosophical importance.

The evolution hypothesis is much older than Darwin; it is, indeed, one of the oldest guessings of human thought. In the eighteenth century it was put forward by Diderot and Lamettrie and suggested by Kant (1786). As we shall see later, it was held also by several philosophers in the first half of the nineteenth century. In his preface to “The Origin of Species”, Darwin mentions the naturalists who were his forerunners. But he has set forth the hypothesis of evolution in so energetic and thorough a manner that it perforce attracts the attention of all thoughtful men in a much higher degree than it did before the publication of the “Origin”.

And further, the importance of his teaching rests on the fact that he, much more than his predecessors, even than Lamarck, sought a foundation for his hypothesis in definite facts. Modern science began by demanding–with Kepler and Newton–evidence of verae causae; this demand Darwin industriously set himself to satisfy–hence the wealth of material which he collected by his observations and his experiments. He not only revived an old hypothesis, but he saw the necessity of verifying it by facts. Whether the special cause on which he founded the explanation of the origin of species–Natural Selection–is sufficient, is now a subject of discussion. He himself had some doubt in regard to this question, and the criticisms which are directed against his hypothesis hit Darwinism rather than Darwin. In his indefatigable search for empirical evidence he is a model even for his antagonists: he has compelled them to approach the problems of life along other lines than those which were formerly followed.

Whether the special cause to which Darwin appealed is sufficient or not, at least to it is probably due the greater part of the influence which he has exerted on the general trend of thought. “Struggle for existence” and “natural selection” are principles which have been applied, more or less, in every department of thought. Recent research, it is true, has discovered greater empirical discontinuity–leaps, “mutations”–whereas Darwin believed in the importance of small variations slowly accumulated. It has also been shown by the experimental method, which in recent biological work has succeeded Darwin’s more historical method, that types once constituted possess great permanence, the fluctuations being restricted within clearly defined boundaries. The problem has become more precise, both as to variation and as to heredity. The inner conditions of life have in both respects shown a greater independence than Darwin had supposed in his theory, though he always admitted that the cause of variation was to him a great enigma, “a most perplexing problem,” and that the struggle for life could only occur where variation existed. But, at any rate, it was of the greatest importance that Darwin gave a living impression of the struggle for life which is everywhere going on, and to which even the highest forms of existence must be amenable. The philosophical importance of these ideas does not stand or fall with the answer to the question, whether natural selection is a sufficient explanation of the origin of species or not: it has an independent, positive value for everyone who will observe life and reality with an unbiassed mind.

In accentuating the struggle for life Darwin stands as a characteristically English thinker: he continues a train of ideas which Hobbes and Malthus had already begun. Moreover in his critical views as to the conception of species he had English forerunners; in the middle ages Occam and Duns Scotus, in the eighteenth century Berkeley and Hume. In his moral philosophy, as we shall see later, he is an adherent of the school which is represented by Hutcheson, Hume and Adam Smith. Because he is no philosopher in the stricter sense of the term, it is of great interest to see that his attitude of mind is that of the great thinkers of his nation.

In considering Darwin’s influence on philosophy we will begin with an examination of the attitude of philosophy to the conception of evolution at the time when “The Origin of Species” appeared. We will then examine the effects which the theory of evolution, and especially the idea of the struggle for life, has had, and naturally must have, on the discussion of philosophical problems.


When “The Origin of Species” appeared fifty years ago Romantic speculation, Schelling’s and Hegel’s philosophy, still reigned on the continent, while in England Positivism, the philosophy of Comte and Stuart Mill, represented the most important trend of thought. German speculation had much to say on evolution, it even pretended to be a philosophy of evolution. But then the word “evolution” was to be taken in an ideal, not in a real, sense. To speculative thought the forms and types of nature formed a system of ideas, within which any form could lead us by continuous transitions to any other. It was a classificatory system which was regarded as a divine world of thought or images, within which metamorphoses could go on–a condition comparable with that in the mind of the poet when one image follows another with imperceptible changes. Goethe’s ideas of evolution, as expressed in his “Metamorphosen der Pflanzen und der Thiere”, belong to this category; it is, therefore, incorrect to call him a forerunner of Darwin. Schelling and Hegel held the same idea; Hegel expressly rejected the conception of a real evolution in time as coarse and materialistic. “Nature,” he says, “is to be considered as a SYSTEM OF STAGES, the one necessarily arising from the other, and being the nearest truth of that from which it proceeds; but not in such a way that the one is NATURALLY generated by the other; on the contrary (their connection lies) in the inner idea which is the ground of nature. The METAMORPHOSIS can be ascribed only to the notion as such, because it alone is evolution…It has been a clumsy idea in the older as well as in the newer philosophy of nature, to regard the transformation and the transition from one natural form and sphere to a higher as an outward and actual production.” (“Encyclopaedie der philosophischen Wissenschaften” (4th edition), Berlin, 1845, paragraph 249.)

The only one of the philosophers of Romanticism who believed in a real, historical evolution, a real production of new species, was Oken. (“Lehrbuch der Naturphilosophie”, Jena, 1809.) Danish philosophers, such as Treschow (1812) and Sibbern (1846), have also broached the idea of an historical evolution of all living beings from the lowest to the highest. Schopenhauer’s philosophy has a more realistic character than that of Schelling’s and Hegel’s, his diametrical opposites, though he also belongs to the romantic school of thought. His philosophical and psychological views were greatly influenced by French naturalists and philosophers, especially by Cabanis and Lamarck. He praises the “ever memorable Lamarck,” because he laid so much stress on the “will to live.” But he repudiates as a “wonderful error” the idea that the organs of animals should have reached their present perfection through a development in time, during the course of innumerable generations. It was, he said, a consequence of the low standard of contemporary French philosophy, that Lamarck came to the idea of the construction of living beings in time through succession! (“Ueber den Willen in der Natur” (2nd edition), Frankfurt a. M., 1854, pages 41-43.)

The positivistic stream of thought was not more in favour of a real evolution than was the Romantic school. Its aim was to adhere to positive facts: it looked with suspicion on far-reaching speculation. Comte laid great stress on the discontinuity found between the different kingdoms of nature, as well as within each single kingdom. As he regarded as unscientific every attempt to reduce the number of physical forces, so he rejected entirely the hypothesis of Lamarck concerning the evolution of species; the idea of species would in his eyes absolutely lose its importance if a transition from species to species under the influence of conditions of life were admitted. His disciples (Littre, Robin) continued to direct against Darwin the polemics which their master had employed against Lamarck. Stuart Mill, who, in the theory of knowledge, represented the empirical or positivistic movement in philosophy–like his English forerunners from Locke to Hume–founded his theory of knowledge and morals on the experience of the single individual. He sympathised with the theory of the original likeness of all individuals and derived their differences, on which he practically and theoretically laid much stress, from the influence both of experience and education, and, generally, of physical and social causes. He admitted an individual evolution, and, in the human species, an evolution based on social progress; but no physiological evolution of species. He was afraid that the hypothesis of heredity would carry us back to the old theory of “innate” ideas.

Darwin was more empirical than Comte and Mill; experience disclosed to him a deeper continuity than they could find; closer than before the nature and fate of the single individual were shown to be interwoven in the great web binding the life of the species with nature as a whole. And the continuity which so many idealistic philosophers could find only in the world of thought, he showed to be present in the world of reality.


Darwin’s energetic renewal of the old idea of evolution had its chief importance in strengthening the conviction of this real continuity in the world, of continuity in the series of form and events. It was a great support for all those who were prepared to base their conception of life on scientific grounds. Together with the recently discovered law of the conservation of energy, it helped to produce the great realistic movement which characterises the last third of the nineteenth century. After the decline of the Romantic movement people wished to have firmer ground under their feet and reality now asserted itself in a more emphatic manner than in the period of Romanticism. It was easy for Hegel to proclaim that “the real” was “the rational,” and that “the rational” was “the real”: reality itself existed for him only in the interpretation of ideal reason, and if there was anything which could not be merged in the higher unity of thought, then it was only an example of the “impotence of nature to hold to the idea.” But now concepts are to be founded on nature and not on any system of categories too confidently deduced a priori. The new devotion to nature had its recompense in itself, because the new points of view made us see that nature could indeed “hold to ideas,” though perhaps not to those which we had cogitated beforehand.

A most important question for philosophers to answer was whether the new views were compatible with an idealistic conception of life and existence. Some proclaimed that we have now no need of any philosophy beyond the principles of the conservation of matter and energy and the principle of natural evolution: existence should and could be definitely and completely explained by the laws of material nature. But abler thinkers saw that the thing was not so simple. They were prepared to give the new views their just place and to examine what alterations the old views must undergo in order to be brought into harmony with the new data.

The realistic character of Darwin’s theory was shown not only in the idea of natural continuity, but also, and not least, in the idea of the cause whereby organic life advances step by step. This idea–the idea of the struggle for life–implied that nothing could persist, if it had no power to maintain itself under the given conditions. Inner value alone does not decide. Idealism was here put to its hardest trial. In continuous evolution it could perhaps still find an analogy to the inner evolution of ideas in the mind; but in the demand for power in order to struggle with outward conditions Realism seemed to announce itself in its most brutal form. Every form of Idealism had to ask itself seriously how it was going to “struggle for life” with this new Realism.

We will now give a short account of the position which leading thinkers in different countries have taken up in regard to this question.

I. Herbert Spencer was the philosopher whose mind was best prepared by his own previous thinking to admit the theory of Darwin to a place in his conception of the world. His criticism of the arguments which had been put forward against the hypothesis of Lamarck, showed that Spencer, as a young man, was an adherent to the evolution idea. In his “Social Statics” (1850) he applied this idea to human life and moral civilisation. In 1852 he wrote an essay on “The Development Hypothesis”, in which he definitely stated his belief that the differentiation of species, like the differentiation within a single organism, was the result of development. In the first edition of his “Psychology” (1855) he took a step which put him in opposition to the older English school (from Locke to Mill): he acknowledged “innate ideas” so far as to admit the tendency of acquired habits to be inherited in the course of generations, so that the nature and functions of the individual are only to be understood through its connection with the life of the species. In 1857, in his essay on “Progress”, he propounded the law of differentiation as a general law of evolution, verified by examples from all regions of experience, the evolution of species being only one of these examples. On the effect which the appearance of “The Origin of Species” had on his mind he writes in his “Autobiography”: “Up to that time…I held that the sole cause of organic evolution is the inheritance of functionally-produced modifications. The “Origin of Species” made it clear to me that I was wrong, and that the larger part of the facts cannot be due to any such cause…To have the theory of organic evolution justified was of course to get further support for that theory of evolution at large with which…all my conceptions were bound up.” (Spencer, “Autobiography”, Vol. II. page 50, London, 1904.) Instead of the metaphorical expression “natural selection,” Spencer introduced the term “survival of the fittest,” which found favour with Darwin as well as with Wallace.

In working out his ideas of evolution, Spencer found that differentiation was not the only form of evolution. In its simplest form evolution is mainly a concentration, previously scattered elements being integrated and losing independent movement. Differentiation is only forthcoming when minor wholes arise within a greater whole. And the highest form of evolution is reached when there is a harmony between concentration and differentiation, a harmony which Spencer calls equilibration and which he defines as a moving equilibrium. At the same time this definition enables him to illustrate the expression “survival of the fittest.” “Every living organism exhibits such a moving equilibrium–a balanced set of functions constituting its life; and the overthrow of this balanced set of functions or moving equilibrium is what we call death. Some individuals in a species are so constituted that their moving equilibria are less easily overthrown than those of other individuals; and these are the fittest which survive, or, in Mr Darwin’s language, they are the select which nature preserves.” (Ibid. page 100.) Not only in the domain of organic life, but in all domains, the summit of evolution is, according to Spencer, characterised by such a harmony–by a moving equilibrium.

Spencer’s analysis of the concept of evolution, based on a great variety of examples, has made this concept clearer and more definite than before. It contains the three elements; integration, differentiation and equilibration. It is true that a concept which is to be valid for all domains of experience must have an abstract character, and between the several domains there is, strictly speaking, only a relation of analogy. So there is only analogy between psychical and physical evolution. But this is no serious objection, because general concepts do not express more than analogies between the phenomena which they represent. Spencer takes his leading terms from the material world in defining evolution (in the simplest form) as integration of matter and dissipation of movement; but as he–not always quite consistently (Cf. my letter to him, 1876, now printed in Duncan’s “Life and Letters of Herbert Spencer”, page 178, London, 1908.)–assumed a correspondence of mind and matter, he could very well give these terms an indirect importance for psychical evolution. Spencer has always, in my opinion with full right, repudiated the ascription of materialism. He is no more a materialist than Spinoza. In his “Principles of Psychology” (paragraph 63) he expressed himself very clearly: “Though it seems easier to translate so-called matter into so-called spirit, than to translate so-called spirit into so-called matter–which latter is indeed wholly impossible–yet no translation can carry us beyond our symbols.” These words lead us naturally to a group of thinkers whose starting-point was psychical evolution. But we have still one aspect of Spencer’s philosophy to mention.

Spencer founded his “laws of evolution” on an inductive basis, but he was convinced that they could be deduced from the law of the conservation of energy. Such a deduction is, perhaps, possible for the more elementary forms of evolution, integration and differentiation; but it is not possible for the highest form, the equilibration, which is a harmony of integration and differentiation. Spencer can no more deduce the necessity for the eventual appearance of “moving equilibria” of harmonious totalities than Hegel could guarantee the “higher unities” in which all contradictions should be reconciled. In Spencer’s hands the theory of evolution acquired a more decidedly optimistic character than in Darwin’s; but I shall deal later with the relation of Darwin’s hypothesis to the opposition of optimism and pessimism.

II. While the starting-point of Spencer was biological or cosmological, psychical evolution being conceived as in analogy with physical, a group of eminent thinkers–in Germany Wundt, in France Fouillee, in Italy Ardigo– took, each in his own manner, their starting-point in psychical evolution as an original fact and as a type of all evolution, the hypothesis of Darwin coming in as a corroboration and as a special example. They maintain the continuity of evolution; they find this character most prominent in psychical evolution, and this is for them a motive to demand a corresponding continuity in the material, especially in the organic domain.

To Wundt and Fouillee the concept of will is prominent. They see the type of all evolution in the transformation of the life of will from blind impulse to conscious choice; the theories of Lamarck and Darwin are used to support the view that there is in nature a tendency to evolution in steady reciprocity with external conditions. The struggle for life is here only a secondary fact. Its apparent prominence is explained by the circumstance that the influence of external conditions is easily made out, while inner conditions can be verified only through their effects. For Ardigo the evolution of thought was the starting-point and the type: in the evolution of a scientific hypothesis we see a progress from the indefinite (indistinto) to the definite (distinto), and this is a characteristic of all evolution, as Ardigo has pointed out in a series of works. The opposition between indistinto and distinto corresponds to Spencer’s opposition between homogeneity and heterogeneity. The hypothesis of the origin of differences of species from more simple forms is a special example of the general law of evolution.

In the views of Wundt and Fouillee we find the fundamental idea of idealism: psychical phenomena as expressions of the innermost nature of existence. They differ from the older Idealism in the great stress which they lay on evolution as a real, historical process which is going on through steady conflict with external conditions. The Romantic dread of reality is broken. It is beyond doubt that Darwin’s emphasis on the struggle for life as a necessary condition of evolution has been a very important factor in carrying philosophy back to reality from the heaven of pure ideas. The philosophy of Ardigo, on the other side, appears more as a continuation and deepening of positivism, though the Italian thinker arrived at his point of view independently of French-English positivism. The idea of continuous evolution is here maintained in opposition to Comte’s and Mill’s philosophy of discontinuity. From Wundt and Fouillee Ardigo differs in conceiving psychical evolution not as an immediate revelation of the innermost nature of existence, but only as a single, though the most accessible example, of evolution.

III. To the French philosophers Boutroux and Bergson, evolution proper is continuous and qualitative, while outer experience and physical science give us fragments only, sporadic processes and mechanical combinations. To Bergson, in his recent work “L’Evolution Creatrice”, evolution consists in an elan de vie which to our fragmentary observation and analytic reflexion appears as broken into a manifold of elements and processes. The concept of matter in its scientific form is the result of this breaking asunder, essential for all scientific reflexion. In these conceptions the strongest opposition between inner and outer conditions of evolution is expressed: in the domain of internal conditions spontaneous development of qualitative forms–in the domain of external conditions discontinuity and mechanical