Darwin and Modern Science Part 34

Professor of Botany in the University of Munich.

There is scarcely any subject to which Darwin devoted so much time and work as to his researches into the biology of flowers, or, in other words, to the consideration of the question to what extent the structural and physiological characters of flowers are correlated with their function of producing fruits and seeds. We know from his own words what fascination these studies possessed for him. We repeatedly find, for example, in his letters expressions such as this:--"Nothing in my life has ever interested me more than the fertilisation of such plants as Primula and Lythrum, or again Anacamptis or Listera." ("More Letters of Charles Darwin", Vol. II. page 419.)

Expressions of this kind coming from a man whose theories exerted an epoch-making influence, would be unintelligible if his researches into the biology of flowers had been concerned only with records of isolated facts, however interesting these might be. We may at once take it for granted that the investigations were undertaken with the view of following up important problems of general interest, problems which are briefly dealt with in this essay.

Darwin published the results of his researches in several papers and in three larger works, (i) "On the various contrivances by which British and Foreign Orchids are fertilised by insects" (First edition, London, 1862; second edition, 1877; popular edition, 1904.) (ii) "The effects of Cross and Self fertilisation in the vegetable kingdom" (First edition, 1876; second edition, 1878). (iii) "The different forms of Flowers on plants of the same species" (First edition, 1877; second edition, 1880).

Although the influence of his work is considered later, we may here point out that it was almost without a parallel; not only does it include a ma.s.s of purely scientific observations, but it awakened interest in very wide circles, as is shown by the fact that we find the results of Darwin's investigations in floral biology universally quoted in school books; they are even willingly accepted by those who, as regards other questions, are opposed to Darwin's views.

The works which we have mentioned are, however, not only of special interest because of the facts they contribute, but because of the MANNER in which the facts are expressed. A superficial reader seeking merely for catch-words will, for instance, probably find the book on cross and self-fertilisation rather dry because of the numerous details which it contains: it is, indeed, not easy to compress into a few words the general conclusions of this volume. But on closer examination, we cannot be sufficiently grateful to the author for the exactness and objectivity with which he enables us to partic.i.p.ate in the scheme of his researches.

He never tries to persuade us, but only to convince us that his conclusions are based on facts; he always gives prominence to such facts as appear to be in opposition to his opinions,--a feature of his work in accordance with a maxim which he laid down:--"It is a golden rule, which I try to follow, to put every fact which is opposed to one's preconceived opinion in the strongest light." ("More Letters", Vol. II.

page 324.)

The result of this method of presentation is that the works mentioned above represent a collection of most valuable doc.u.ments even for those who feel impelled to draw from the data other conclusions than those of the author. Each investigation is the outcome of a definite question, a "preconceived opinion," which is either supported by the facts or must be abandoned. "How odd it is that anyone should not see that all observation must be for or against some view if it is to be of any service!" (Ibid. Vol. I. page 195.)

The points of view which Darwin had before him were princ.i.p.ally the following. In the first place the proof that a large number of the peculiarities in the structure of flowers are not useless, but of the greatest significance in pollination must be of considerable importance for the interpretation of adaptations; "The use of each trifling detail of structure is far from a barren search to those who believe in natural selection." ("Fertilisation of Orchids" (1st edition), page 351; (2nd edition 1904) page 286.) Further, if these structural relations are shown to be useful, they may have been acquired because from the many variations which have occurred along different lines, those have been preserved by natural selection "which are beneficial to the organism under the complex and ever-varying conditions of life." (Ibid. page 351.) But in the case of flowers there is not only the question of adaptation to fertilisation to be considered. Darwin, indeed, soon formed the opinion which he has expressed in the following sentence,--"From my own observations on plants, guided to a certain extent by the experience of the breeders of animals, I became convinced many years ago that it is a general law of nature that flowers are adapted to be crossed, at least occasionally, by pollen from a distinct plant." ("Cross and Self fertilisation" (1st edition), page 6.)

The experience of animal breeders pointed to the conclusion that continual in-breeding is injurious. If this is correct, it raises the question whether the same conclusion holds for plants. As most flowers are hermaphrodite, plants afford much more favourable material than animals for an experimental solution of the question, what results follow from the union of nearly related s.e.xual cells as compared with those obtained by the introduction of new blood. The answer to this question must, moreover, possess the greatest significance for the correct understanding of s.e.xual reproduction in general.

We see, therefore, that the problems which Darwin had before him in his researches into the biology of flowers were of the greatest importance, and at the same time that the point of view from which he attacked the problems was essentially a teleological one.

We may next inquire in what condition he found the biology of flowers at the time of his first researches, which were undertaken about the year 1838. In his autobiography he writes,--"During the summer of 1839, and, I believe, during the previous summer, I was led to attend to the cross-fertilisation of flowers by the aid of insects, from having come to the conclusion in my speculations on the origin of species, that crossing played an important part in keeping specific forms constant."

("The Life and Letters of Charles Darwin", Vol. I. page 90, London, 1888.) In 1841 he became acquainted with Sprengel's work: his researches into the biology of flowers were thus continued for about forty years.

It is obvious that there could only be a biology of flowers after it had been demonstrated that the formation of seeds and fruit in the flower is dependent on pollination and subsequent fertilisation. This proof was supplied at the end of the seventeenth century by R.J. Camerarius (1665-1721). He showed that normally seeds and fruits are developed only when the pollen reaches the stigma. The manner in which this happens was first thoroughly investigated by J.G. Kolreuter (1733-1806 (Kolreuter, "Vorlaufige Nachricht von einigen das Geschlecht der Planzen betreffenden Versuchen und Beobachtungen", Leipzig, 1761; with three supplements, 1763-66. Also, "Mem. de l'acad. St Petersbourg", Vol. XV.

1809.)), the same observer to whom we owe the earliest experiments in hybridisation of real scientific interest. Kolreuter mentioned that pollen may be carried from one flower to another partly by wind and partly by insects. But he held the view, and that was, indeed, the natural a.s.sumption, that self-fertilisation usually occurs in a flower, in other words that the pollen of a flower reaches the stigma of the same flower. He demonstrated, however, certain cases in which cross-pollination occurs, that is in which the pollen of another flower of the same species is conveyed to the stigma. He was familiar with the phenomenon, exhibited by numerous flowers, to which Sprengel afterwards applied the term Dichogamy, expressing the fact that the anthers and stigmas of a flower often ripen at different times, a peculiarity which is now recognised as one of the commonest means of ensuring cross-pollination.

With far greater thoroughness and with astonis.h.i.+ng power of observation C.K. Sprengel (1750-1816) investigated the conditions of pollination of flowers. Darwin was introduced by that eminent botanist Robert Brown to Sprengel's then but little appreciated work,--"Das entdeckte Geheimniss der Natur im Bau und in der Befruchtung der Blumen" (Berlin, 1793); this is by no means the least service to Botany rendered by Robert Brown.

Sprengel proceeded from a naive teleological point of view. He firmly believed "that the wise Author of nature had not created a single hair without a definite purpose." He succeeded in demonstrating a number of beautiful adaptations in flowers for ensuring pollination; but his work exercised but little influence on his contemporaries and indeed for a long time after his death. It was through Darwin that Sprengel's work first achieved a well deserved though belated fame. Even such botanists as concerned themselves with researches into the biology of flowers appear to have formerly attached much less value to Sprengel's work than it has received since Darwin's time. In ill.u.s.tration of this we may quote C.F. Gartner whose name is rightly held in the highest esteem as that of one of the most eminent hybridologists. In his work "Versuche und Beobachtungen uder die Befruchtungsorgane der vollkommeneren Gewachse und uber die naturliche und kunstliche Befruchtung durch den eigenen Pollen" he also deals with flower-pollination. He recognised the action of the wind, but he believed, in spite of the fact that he both knew and quoted Kolreuter and Sprengel, that while insects a.s.sist pollination, they do so only occasionally, and he held that insects are responsible for the conveyance of pollen; thorough investigations would show "that a very small proportion of the plants included in this category require this a.s.sistance in their native habitat." (Gartner, "Versucher und Beobachtungen... ", page 335, Stuttgart, 1844.) In the majority of plants self-pollination occurs.

Seeing that even investigators who had worked for several decades at fertilisation-phenomena had not advanced the biology of flowers beyond the initial stage, we cannot be surprised that other botanists followed to even a less extent the lines laid down by Kolreuter and Sprengel.

This was in part the result of Sprengel's supernatural teleology and in part due to the fact that his book appeared at a time when other lines of inquiry exerted a dominating influence.

At the hands of Linnaeus systematic botany reached a vigorous development, and at the beginning of the nineteenth century the anatomy and physiology of plants grew from small beginnings to a flouris.h.i.+ng branch of science. Those who concerned themselves with flowers endeavoured to investigate their development and structure or the most minute phenomena connected with fertilisation and the formation of the embryo. No room was left for the extension of the biology of flowers on the lines marked out by Kolreuter and Sprengel. Darwin was the first to give new life and a deeper significance to this subject, chiefly because he took as his starting-point the above-mentioned problems, the importance of which is at once admitted by all naturalists.

The further development of floral biology by Darwin is in the first place closely connected with the book on the fertilisation of Orchids.

It is noteworthy that the t.i.tle includes the sentence,--"and on the good effects of intercrossing."

The purpose of the book is clearly stated in the introduction:--"The object of the following work is to show that the contrivances by which Orchids are fertilised, are as varied and almost as perfect as any of the most beautiful adaptations in the animal kingdom; and, secondly, to show that these contrivances have for their main object the fertilisation of each flower by the pollen of another flower."

("Fertilisation of Orchids", page 1.) Orchids const.i.tuted a particularly suitable family for such researches. Their flowers exhibit a striking wealth of forms; the question, therefore, whether the great variety in floral structure bears any relation to fertilisation (In the older botanical literature the word fertilisation is usually employed in cases where POLLINATION is really in question: as Darwin used it in this sense it is so used here.) must in this case possess special interest.

Darwin succeeded in showing that in most of the orchids examined self-fertilisation is either an impossibility, or, under natural conditions, occurs only exceptionally. On the other hand these plants present a series of extraordinarily beautiful and remarkable adaptations which ensure the transference of pollen by insects from one flower to another. It is impossible to describe adequately in a few words the wealth of facts contained in the Orchid book. A few examples may, however, be quoted in ill.u.s.tration of the delicacy of the observations and of the perspicuity employed in interpreting the facts.

The majority of orchids differ from other seed plants (with the exception of the Asclepiads) in having no dust-like pollen. The pollen, or more correctly, the pollen-tetrads, remain fastened together as club-shaped pollinia usually borne on a slender pedicel. At the base of the pedicel is a small viscid disc by which the pollinium is attached to the head or proboscis of one of the insects which visit the flower.

Darwin demonstrated that in Orchis and other flowers the pedicel of the pollinium, after its removal from the anther, undergoes a curving movement. If the pollinium was originally vertical, after a time it a.s.sumed a horizontal position. In the latter position, if the insect visited another flower, the pollinium would exactly hit the sticky stigmatic surface and thus effect fertilisation. The relation between the behaviour of the viscid disc and the secretion of nectar by the flower is especially remarkable. The flowers possess a spur which in some species (e.g. Gymnadenia conopsea, Platanthera bifolia, etc.) contains honey (nectar), which serves as an attractive bait for insects, but in others (e.g. our native species of Orchis) the spur is empty.

Darwin held the opinion, confirmed by later investigations, that in the case of flowers without honey the insects must penetrate the wall of the nectarless spurs in order to obtain a nectar-like substance. The glands behave differently in the nectar-bearing and in the nectarless flowers.

In the former they are so sticky that they at once adhere to the body of the insect; in the nectarless flowers firm adherence only occurs after the viscid disc has hardened. It is, therefore, adaptively of value that the insects should be detained longer in the nectarless flowers (by having to bore into the spur),--than in flowers in which the nectar is freely exposed. "If this relation, on the one hand, between the viscid matter requiring some little time to set hard, and the nectar being so lodged that moths are delayed in getting it; and, on the other hand, between the viscid matter being at first as viscid as ever it will become, and the nectar lying all ready for rapid suction, be accidental, it is a fortunate accident for the plant. If not accidental, and I cannot believe it to be accidental, what a singular case of adaptation!"

("Fertilisation of Orchids" (1st edition), page 53.)

Among exotic orchids Catasetum is particularly remarkable. One and the same species bears different forms of flowers. The species known as Catasetum tridentatum has pollinia with very large viscid discs; on touching one of the two filaments (antennae) which occur on the gynostemium of the flower the pollinia are shot out to a fairly long distance (as far as 1 metre) and in such manner that they alight on the back of the insect, where they are held. The antennae have, moreover, acquired an importance, from the point of view of the physiology of stimulation, as stimulus-perceiving organs. Darwin had shown that it is only a touch on the antennae that causes the explosion, while contact, blows, wounding, etc. on other places produce no effect. This form of flower proved to be the male. The second form, formerly regarded as a distinct species and named Monachanthus viridis, is shown to be the female flower. The anthers have only rudimentary pollinia and do not open; there are no antennae, but on the other hand numerous seeds are produced. Another type of flower, known as Myanthus barbatus, was regarded by Darwin as a third form: this was afterwards recognised by Rolfe (Rolfe, R.A. "On the s.e.xual forms of Catasetum with special reference to the researches of Darwin and others," "Journ. Linn. Soc."

Vol. XXVII. (Botany), 1891, pages 206-225.) as the male flower of another species, Catasetum barbatum Link, an identification in accordance with the discovery made by Cruger in Trinidad that it always remains sterile.

Darwin had noticed that the flowers of Catasetum do not secrete nectar, and he conjectured that in place of it the insects gnaw a tissue in the cavity of the labellum which has a "slightly sweet, pleasant and nutritious taste." This conjecture as well as other conclusions drawn by Darwin from Catasetum have been confirmed by Cruger--a.s.suredly the best proof of the ac.u.men with which the wonderful floral structure of this "most remarkable of the Orchids" was interpretated far from its native habitat.

As is shown by what we have said about Catasetum, other problems in addition to those concerned with fertilisation are dealt with in the Orchid book. This is especially the case in regard to flower morphology.

The scope of flower morphology cannot be more clearly and better expressed than by these words: "He will see how curiously a flower may be moulded out of many separate organs--how perfect the cohesion of primordially distinct parts may become,--how organs may be used for purposes widely different from their proper function,--how other organs may be entirely suppressed, or leave mere useless emblems of their former existence." ("Fertilisation of Orchids", page 289.)

In attempting, from this point of view, to refer the floral structure of orchids to their original form, Darwin employed a much more thorough method than that of Robert Brown and others. The result of this was the production of a considerable literature, especially in France, along the lines suggested by Darwin's work. This is the so-called anatomical method, which seeks to draw conclusions as to the morphology of the flower from the course of the vascular bundles in the several parts. (He wrote in one of his letters, "... the destiny of the whole human race is as nothing to the course of vessels of orchids" ("More Letters", Vol.

II. page 275.) Although the interpretation of the orchid flower given by Darwin has not proved satisfactory in one particular point--the composition of the labellum--the general results have received universal a.s.sent, namely "that all Orchids owe what they have in common to descent from some monocotyledonous plant, which, like so many other plants of the same division, possessed fifteen organs arranged alternately three within three in five whorls." ("Fertilisation of Orchids" (1st edition), page 307.) The alterations which their original form has undergone have persisted so far as they were found to be of use.

We see also that the remarkable adaptations of which we have given some examples are directed towards cross-fertilisation. In only a few of the orchids investigated by Darwin--other similar cases have since been described--was self-fertilisation found to occur regularly or usually.

The former is the case in the Bee Ophrys (Ophrys apifera), the mechanism of which greatly surprised Darwin. He once remarked to a friend that one of the things that made him wish to live a few thousand years was his desire to see the extinction of the Bee Ophrys, an end to which he believed its self-fertilising habit was leading. ("Life and Letters", Vol. III. page 276 (footnote).) But, he wrote, "the safest conclusion, as it seems to me, is, that under certain unknown circ.u.mstances, and perhaps at very long intervals of time, one individual of the Bee Ophrys is crossed by another." ("Fertilisation of Orchids" page 71.)

If, on the one hand, we remember how much more sure self-fertilisation would be than cross-fertilisation, and, on the other hand, if we call to mind the numerous contrivances for cross-fertilisation, the conclusion is naturally reached that "it is an astonis.h.i.+ng fact that self-fertilisation should not have been an habitual occurrence. It apparently demonstrates to us that there must be something injurious in the process. Nature thus 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 circ.u.mstances, 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 a.n.a.logous 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 unis.e.xual animals." ("Forms of Flowers" (1st edition), page 2.) We have here an example of hermaphrodite flowers which are s.e.xually 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 s.e.xually 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 s.e.xual elements and on their inability to combine in a particular manner. This functional difference of s.e.xual 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 ent.i.ty 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 s.e.xes 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, a.s.suming, 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 a.n.a.logy."