Haberlandt and Nemec ("Ber. d. Deutschen bot. Gesellschaft", XVIII. 1900. See F. Darwin, Presidential Address to Section K, British Association, 1904.) published independently and simultaneously a theory of the mechanism by which plants are orientated in relation to gravitation. And here again we find an arrangement identical in principle with that by which certain animals recognise the vertical, namely the pressure of free particles on the irritable wall of a cavity. In the higher plants, Nemec and Haberlandt believe that special loose and freely movable starch-grains play the part of the otoliths or statoliths of the crustacea, while the protoplasm lining the cells in which they are contained corresponds to the sensitive membrane lining the otocyst of the animal. What is of special interest in our present connection is that according to this ingenious theory (The original conception was due to Noll ("Heterogene Induction", Leipzig, 1892), but his view differed in essential points from those here given.) the sense of verticality in a plant is a form of contact-irritability. The vertical position is distinguished from the horizontal by the fact that, in the latter case, the loose starch-grains rest on the lateral walls of the cells instead of on the terminal walls as occurs in the normal upright position. It should be added that the statolith theory is still sub judice; personally I cannot doubt that it is in the main a satisfactory explanation of the facts.

With regard to the RAPIDITY of the reaction of tendrils, Darwin records ("Climbing Plants", page 155. Others have observed movement after about 6".) that a Passion-Flower tendril moved distinctly within 25 seconds of stimulation. It was this fact, more than any other, that made him doubt the current explanation, viz. that the movement is due to unequal growth on the two sides of the tendril. The interesting work of Fitting (Pringsheim's "Jahrb." XXXVIII. 1903, page 545.) has shown, however, that the primary cause is not (as Darwin supposed) contraction on the concave, but an astonishingly rapid increase in growth-rate on the convex side.

On the last page of "Climbing Plants" Darwin wrote: "It has often been vaguely asserted that plants are distinguished from animals by not having the power of movement. It should rather be said that plants acquire and display this power only when it is of some advantage to them."

He gradually came to realise the vividness and variety of vegetable life, and that a plant like an animal has capacities of behaving in different ways under different circumstances, in a manner that may be compared to the instinctive movements of animals. This point of view is expressed in well-known passages in the "Power of Movement". ("The Power of Movement in Plants", 1880, pages 571-3.) "It is impossible not to be struck with the resemblance between the... movements of plants and many of the actions performed unconsciously by the lower animals." And again, "It is hardly an exaggeration to say that the tip of the radicle... having the power of directing the movements of the adjoining parts, acts like the brain of one of the lower animals; the brain being seated within the anterior end of the body, receiving impressions from the sense-organs, and directing the several movements."

The conception of a region of perception distinct from a region of movement is perhaps the most fruitful outcome of his work on the movements of plants. But many years before its publication, viz. in 1861, he had made out the wonderful fact that in the Orchid Catasetum ("Life and Letters", III. page 268.) the projecting organs or antennae are sensitive to a touch, and transmit an influence "for more than one inch INSTANTANEOUSLY," which leads to the explosion or violent ejection of the pollinia. And as we have already seen a similar transmission of a stimulus was discovered by him in Sundew in 1860, so that in 1862 he could write to Hooker ("Life and Letters", III. page 321.): "I cannot avoid the conclusion, that Drosera possesses matter at least in some degree analogous in constitution and function to nervous matter." I propose in what follows to give some account of the observations on the transmission of stimuli given in the "Power of Movement". It is impossible within the space at my command to give anything like a complete account of the matter, and I must necessarily omit all mention of much interesting work. One well-known experiment consisted in putting opaque caps on the tips of seedling grasses (e.g. oat and canary-grass) and then exposing them to light from one side. The difference, in the amount of curvature towards the light, between the blinded and unblinded specimens, was so great that it was concluded that the light-sensitiveness resided exclusively in the tip. The experiment undoubtedly proves that the sensitiveness is much greater in the tip than elsewhere, and that there is a transmission of stimulus from the tip to the region of curvature. But Rothert (Rothert, Cohn's "Beitrage", VII. 1894.) has conclusively proved that the basal part where the curvature occurs is also DIRECTLY sensitive to light. He has shown, however, that in other grasses (Setaria, Panicum) the cotyledon is the only part which is sensitive, while the hypocotyl, where the movement occurs, is not directly sensitive.

It was however the question of the localisation of the gravitational sense in the tip of the seedling root or radicle that aroused most attention, and it was on this question that a controversy arose which has continued to the present day.

The experiment on which Darwin's conclusion was based consisted simply in cutting off the tip, and then comparing the behaviour of roots so treated with that of normal specimens. An uninjured root when placed horizontally regains the vertical by means of a sharp downward curve; not so a decapitated root which continues to grow more or less horizontally. It was argued that this depends on the loss of an organ specialised for the perception of gravity, and residing in the tip of the root; and the experiment (together with certain important variants) was claimed as evidence of the existence of such an organ.

It was at once objected that the amputation of the tip might check curvature by interfering with longitudinal growth, on the distribution of which curvature depends. This objection was met by showing that an injury, e.g. splitting the root longitudinally (See F. Darwin, "Linnean Soc. Journal (Bot)." XIX. 1882, page 218.), which does not remove the tip, but seriously checks growth, does not prevent geotropism. This was of some interest in another and more general way, in showing that curvature and longitudinal growth must be placed in different categories as regards the conditions on which they depend.

Another objection of a much more serious kind was that the amputation of the tip acts as a shock. It was shown by Rothert (See his excellent summary of the subject in "Flora" 1894 (Erganzungsband), page 199.) that the removal of a small part of the cotyledon of Setaria prevents the plant curving towards the light, and here there is no question of removing the sense-organ since the greater part of the sensitive cotyledon is intact. In view of this result it was impossible to rely on the amputations performed on roots as above described.

At this juncture a new and brilliant method originated in Pfeffer's laboratory. (See Pfeffer, "Annals of Botany", VIII. 1894, page 317, and Czapek, Pringsheim's "Jahrb." XXVII. 1895, page 243.) Pfeffer and Czapek showed that it is possible to bend the root of a lupine so that, for instance, the supposed sense-organ at the tip is vertical while the motile region is horizontal. If the motile region is directly sensitive to gravity the root ought to curve downwards, but this did not occur: on the contrary it continued to grow horizontally. This is precisely what should happen if Darwin's theory is the right one: for if the tip is kept vertical, the sense-organ is in its normal position and receives no stimulus from gravitation, and therefore can obviously transmit none to the region of curvature. Unfortunately this method did not convince the botanical world because some of those who repeated Czapek's experiment failed to get his results.

Czapek ("Berichte d. Deutsch. bot. Ges." XV. 1897, page 516, and numerous subsequent papers. English readers should consult Czapek in the "Annals of Botany", XIX. 1905, page 75.) has devised another interesting method which throws light on the problem. He shows that roots, which have been placed in a horizontal position and have therefore been geotropically stimulated, can be distinguished by a chemical test from vertical, i.e. unstimulated roots. The chemical change in the root can be detected before any curvature has occurred and must therefore be a symptom of stimulation, not of movement. It is particularly interesting to find that the change in the root, on which Czapek's test depends, takes place in the tip, i.e. in the region which Darwin held to be the centre for gravitational sensitiveness.

In 1899 I devised a method (F. Darwin, "Annals of Botany", XIII. 1899, page 567.) by which I sought to prove that the cotyledon of Setaria is not only the organ for light-perception, but also for gravitation. If a seedling is supported horizontally by pushing the apical part (cotyledon) into a horizontal tube, the cotyledon will, according to my supposition, be stimulated gravitationally and a stimulus will be transmitted to the basal part of the stem (hypocotyl) causing it to bend. But this curvature merely raises the basal end of the seedling, the sensitive cotyledon remains horizontal, imprisoned in its tube; it will therefore be continually stimulated and will continue to transmit influences to the bending region, which should therefore curl up into a helix or corkscrew-like form,—and this is precisely what occurred.

I have referred to this work principally because the same method was applied to roots by Massart (Massart, "Mem. Couronnes Acad. R. Belg." LXII. 1902.) and myself (F. Darwin, "Linnean Soc. Journ." XXXV. 1902, page 266.) with a similar though less striking result. Although these researches confirmed Darwin's work on roots, much stress cannot be laid on them as there are several objections to them, and they are not easily repeated.

The method which—as far as we can judge at present—seems likely to solve the problem of the root-tip is most ingenious and is due to Piccard. (Pringsheim's "Jahrb." XL. 1904, page 94.)

Andrew Knight's celebrated experiment showed that roots react to centrifugal force precisely as they do to gravity. So that if a bean root is fixed to a wheel revolving rapidly on a horizontal axis, it tends to curve away from the centre in the line of a radius of the wheel. In ordinary demonstrations of Knight's experiment the seed is generally fixed so that the root is at right angles to a radius, and as far as convenient from the centre of rotation. Piccard's experiment is arranged differently. (A seed is depicted below a horizontal dotted line AA, projecting a root upwards.) The root is oblique to the axis of rotation, and the extreme tip projects beyond that axis. Line AA represents the axis of rotation, T is the tip of the root just above the line AA, and B is the region just below line AA in which curvature takes place. If the motile region B is directly sensitive to gravitation (and is the only part which is sensitive) the root will curve (down and away from the vertical) away from the axis of rotation, just as in Knight's experiment. But if the tip T is alone sensitive to gravitation the result will be exactly reversed, the stimulus originating in T and conveyed to B will produce curvature (up towards the vertical). We may think of the line AA as a plane dividing two worlds. In the lower one gravity is of the earthly type and is shown by bodies falling and roots curving downwards: in the upper world bodies fall upwards and roots curve in the same direction. The seedling is in the lower world, but its tip containing the supposed sense-organ is in the strange world where roots curve upwards. By observing whether the root bends up or down we can decide whether the impulse to bend originates in the tip or in the motile region.

Piccard's results showed that both curvatures occurred and he concluded that the sensitive region is not confined to the tip. (Czapek (Pringsheim's "Jahrb." XXXV. 1900, page 362) had previously given reasons for believing that, in the root, there is no sharp line of separation between the regions of perception and movement.)

Haberlandt (Pringsheim's "Jahrb." XLV. 1908, page 575.) has recently repeated the experiment with the advantage of better apparatus and more experience in dealing with plants, and has found as Piccard did that both the tip and the curving region are sensitive to gravity, but with the important addition that the sensitiveness of the tip is much greater than that of the motile region. The case is in fact similar to that of the oat and canary-grass. In both instances my father and I were wrong in assuming that the sensitiveness is confined to the tip, yet there is a concentration of irritability in that region and transmission of stimulus is as true for geotropism as it is for heliotropism. Thus after nearly thirty years the controversy of the root-tip has apparently ended somewhat after the fashion of the quarrels at the "Rainbow" in "Silas Marner"—"you're both right and you're both wrong." But the "brain-function" of the root-tip at which eminent people laughed in early days turns out to be an important part of the truth. (By using Piccard's method I have succeeded in showing that the gravitational sensitiveness of the cotyledon of Sorghum is certainly much greater than the sensitiveness of the hypocotyl—if indeed any such sensitiveness exists. See Wiesner's "Festschrift", Vienna, 1908.)

Another observation of Darwin's has given rise to much controversy. ("Power of Movement", page 133.) If a minute piece of card is fixed obliquely to the tip of a root some influence is transmitted to the region of curvature and the root bends away from the side to which the card was attached. It was thought at the time that this proved the root-tip to be sensitive to contact, but this is not necessarily the case. It seems possible that the curvature is a reaction to the injury caused by the alcoholic solution of shellac with which the cards were cemented to the tip. This agrees with the fact given in the "Power of Movement" that injuring the root-tip on one side, by cutting or burning it, induced a similar curvature. On the other hand it was shown that curvature could be produced in roots by cementing cards, not to the naked surface of the root-tip, but to pieces of gold-beaters skin applied to the root; gold-beaters skin being by itself almost without effect. But it must be allowed that, as regards touch, it is not clear how the addition of shellac and card can increase the degree of contact. There is however some evidence that very close contact from a solid body, such as a curved fragment of glass, produces curvature: and this may conceivably be the explanation of the effect of gold-beaters skin covered with shellac. But on the whole it is perhaps safer to classify the shellac experiments with the results of undoubted injury rather than with those of contact.

Another subject on which a good deal of labour was expended is the sleep of leaves, or as Darwin called it their NYCTITROPIC movement. He showed for the first time how widely spread this phenomenon is, and attempted to give an explanation of the use to the plant of the power of sleeping. His theory was that by becoming more or less vertical at night the leaves escape the chilling effect of radiation. Our method of testing this view was to fix some of the leaves of a sleeping plant so that they remained horizontal at night and therefore fully exposed to radiation, while their fellows were partly protected by assuming the nocturnal position. The experiments showed clearly that the horizontal leaves were more injured than the sleeping, i.e. more or less vertical, ones. It may be objected that the danger from cold is very slight in warm countries where sleeping plants abound. But it is quite possible that a lowering of the temperature which produces no visible injury may nevertheless be hurtful by checking the nutritive processes (e.g. translocation of carbohydrates), which go on at night. Stahl ("Bot. Zeitung", 1897, page 81.) however has ingeniously suggested that the exposure of the leaves to radiation is not DIRECTLY hurtful because it lowers the temperature of the leaf, but INDIRECTLY because it leads to the deposition of dew on the leaf-surface. He gives reasons for believing that dew-covered leaves are unable to transpire efficiently, and that the absorption of mineral food-material is correspondingly checked. Stahl's theory is in no way destructive of Darwin's, and it is possible that nyctitropic leaves are adapted to avoid the indirect as well as the direct results of cooling by radiation.

In what has been said I have attempted to give an idea of some of the discoveries brought before the world in the "Power of Movement" (In 1881 Professor Wiesner published his "Das Bewegungsvermogen der Pflanzen", a book devoted to the criticism of "The Power of Movement in Plants". A letter to Wiesner, published in "Life and Letters", III. page 336, shows Darwin's warm appreciation of his critic's work, and of the spirit in which it is written.) and of the subsequent history of the problems. We must now pass on to a consideration of the central thesis of the book,—the relation of circumnutation to the adaptive curvatures of plants.

Darwin's view is plainly stated on pages 3-4 of the "Power of Movement". Speaking of circumnutation he says, "In this universally present movement we have the basis or groundwork for the acquirement, according to the requirements of the plant, of the most diversified movements." He then points out that curvatures such as those towards the light or towards the centre of the earth can be shown to be exaggerations of circumnutation in the given directions. He finally points out that the difficulty of conceiving how the capacities of bending in definite directions were acquired is diminished by his conception. "We know that there is always movement in progress, and its amplitude, or direction, or both, have only to be modified for the good of the plant in relation with internal or external stimuli."

It may at once be allowed that the view here given has not been accepted by physiologists. The bare fact that circumnutation is a general property of plants (other than climbing species) is not generally rejected. But the botanical world is no nearer to believing in the theory of reaction built on it.

If we compare the movements of plants with those of the lower animals we find a certain resemblance between the two. According to Jennings (H.S. Jennings, "The Behavior of the Lower Animals". Columbia U. Press, N.Y. 1906.) a Paramoecium constantly tends to swerve towards the aboral side of its body owing to certain peculiarities in the set and power of its cilia. But the tendency to swim in a circle, thus produced, is neutralised by the rotation of the creature about its longitudinal axis. Thus the direction of the swerves IN RELATION TO THE PATH of the organism is always changing, with the result that the creature moves in what approximates to a straight line, being however actually a spiral about the general line of progress. This method of motion is strikingly like the circumnutation of a plant, the apex of which also describes a spiral about the general line of growth. A rooted plant obviously cannot rotate on its axis, but the regular series of curvatures of which its growth consists correspond to the aberrations of Paramoecium distributed regularly about its course by means of rotation. (In my address to the Biological Section of the British Association at Cardiff (1891) I have attempted to show the connection between circumnutation and RECTIPETALITY, i.e. the innate capacity of growing in a straight line.) Just as a plant changes its direction of growth by an exaggeration of one of the curvature-elements of which circumnutation consists, so does a Paramoecium change its course by the accentuation of one of the deviations of which its path is built. Jennings has shown that the infusoria, etc., react to stimuli by what is known as the "method of trial." If an organism swims into a region where the temperature is too high or where an injurious substance is present, it changes its course. It then moves forward again, and if it is fortunate enough to escape the influence, it continues to swim in the given direction. If however its change of direction leads it further into the heated or poisonous region it repeats the movement until it emerges from its difficulties. Jennings finds in the movements of the lower organisms an analogue with what is known as pain in conscious organisms. There is certainly this much resemblance that a number of quite different sub-injurious agencies produce in the lower organisms a form of reaction by the help of which they, in a partly fortuitous way, escape from the threatening element in their environment. The higher animals are stimulated in a parallel manner to vague and originally purposeless movements, one of which removes the discomfort under which they suffer, and the organism finally learns to perform the appropriate movement without going through the tentative series of actions.

I am tempted to recognise in circumnutation a similar groundwork of tentative movements out of which the adaptive ones were originally selected by a process rudely representative of learning by experience.

It is, however, simpler to confine ourselves to the assumption that those plants have survived which have acquired through unknown causes the power of reacting in appropriate ways to the external stimuli of light, gravity, etc. It is quite possible to conceive this occurring in plants which have no power of circumnutating—and, as already pointed out, physiologists do as a fact neglect circumnutation as a factor in the evolution of movements. Whatever may be the fate of Darwin's theory of circumnutation there is no doubt that the research he carried out in support of, and by the light of, this hypothesis has had a powerful influence in guiding the modern theories of the behaviour of plants. Pfeffer ("The Physiology of Plants", Eng. Tr. III. page 11.), who more than any one man has impressed on the world a rational view of the reactions of plants, has acknowledged in generous words the great value of Darwin's work in the same direction. The older view was that, for instance, curvature towards the light is the direct mechanical result of the difference of illumination on the lighted and shaded surfaces of the plant. This has been proved to be an incorrect explanation of the fact, and Darwin by his work on the transmission of stimuli has greatly contributed to the current belief that stimuli act indirectly. Thus we now believe that in a root and a stem the mechanism for the perception of gravitation is identical, but the resulting movements are different because the motor-irritabilities are dissimilar in the two cases. We must come back, in fact, to Darwin's comparison of plants to animals. In both there is perceptive machinery by which they are made delicately alive to their environment, in both the existing survivors are those whose internal constitution has enabled them to respond in a beneficial way to the disturbance originating in their sense-organs.



XX. THE BIOLOGY OF FLOWERS. By K. Goebel, Ph.D.

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 mass 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 participate 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 documents 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 principally 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 sexual 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 sexual 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 assumption, 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 astonishing 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 illustration 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 assist 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 assistance 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 flourishing 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 title 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 constituted 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 illustration 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 assumed 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 sexual 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—assuredly the best proof of the acumen 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 assent, 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 circumstances, 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 astonishing 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 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.



XXI. MENTAL FACTORS IN EVOLUTION. By C. Lloyd Morgan, LL.D., F.R.S.

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.