My father's interest in plants was of two kinds, which may be roughly distinguished as EVOLUTIONARY and PHYSIOLOGICAL. Thus in his purely evolutionary work, for instance in "The Origin of Species" and in his book on "Variation under Domestication", plants as well as animals served as material for his generalisations. He was largely dependent on the work of others for the facts used in the evolutionary work, and despised himself for belonging to the "blessed gang" of compilers. And he correspondingly rejoiced in the employment of his wonderful power of observation in the physiological problems which occupied so much of his later life. But inasmuch as he felt evolution to be his life's work, he regarded himself as something of an idler in observing climbing plants, insectivorous plants, orchids, etc. In this physiological work he was to a large extent urged on by his passionate desire to understand the machinery of all living things. But though it is true that he worked at physiological problems in the naturalist's spirit of curiosity, yet there was always present to him the bearing of his facts on the problem of evolution. His interests, physiological and evolutionary, were indeed so interwoven that they cannot be sharply separated. Thus his original interest in the fertilisation of flowers was evolutionary. "I was led" ("Life and Letters", I. page 90.), he says, "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." In the same way the value of his experimental work on heterostyled plants crystalised out in his mind into the conclusion that the product of illegitimate unions are equivalent to hybrids—a conclusion of the greatest interest from an evolutionary point of view. And again his work "Cross and Self Fertilisation" may be condensed to a point of view of great importance in reference to the meaning and origin of sexual reproduction. (See Professor Goebel's article in the present volume.)
The whole of his physiological work may be looked at as an illustration of the potency of his theory as an "instrument for the extension of the realm of natural knowledge." (Huxley in Darwin's "Life and Letters." II. page 204.)
His doctrine of natural selection gave, as is well known, an impulse to the investigation of the use of organs—and thus created the great school of what is known in Germany as Biology—a department of science for which no English word exists except the rather vague term Natural History. This was especially the case in floral biology, and it is interesting to see with what hesitation he at first expressed the value of his book on Orchids ("Life and Letters", III. page 254.), "It will perhaps serve to illustrate how Natural History may be worked under the belief of the modification of species" (1861). And in 1862 he speaks (Loc. cit.) more definitely of the relation of his work to natural selection: "I can show the meaning of some of the apparently meaningless ridges (and) horns; who will now venture to say that this or that structure is useless?" It is the fashion now to minimise the value of this class of work, and we even find it said by a modern writer that to inquire into the ends subserved by organs is not a scientific problem. Those who take this view surely forget that the structure of all living things is, as a whole, adaptive, and that a knowledge of how the present forms come to be what they are includes a knowledge of why they survived. They forget that the SUMMATION of variations on which divergence depends is under the rule of the environment considered as a selective force. They forget that the scientific study of the interdependence of organisms is only possible through a knowledge of the machinery of the units. And that, therefore, the investigation of such widely interesting subjects as extinction and distribution must include a knowledge of function. It is only those who follow this line of work who get to see the importance of minute points of structure and understand as my father did even in 1842, as shown in his sketch of the "Origin" (Now being prepared for publication.), that every grain of sand counts for something in the balance. Much that is confidently stated about the uselessness of different organs would never have been written if the naturalist spirit were commoner nowadays. This spirit is strikingly shown in my father's work on the movements of plants. The circumstance that botanists had not, as a class, realised the interest of the subject accounts for the fact that he was able to gather such a rich harvest of results from such a familiar object as a twining plant. The subject had been investigated by H. von Mohl, Palm, and Dutrochet, but they failed not only to master the problem but (which here concerns us) to give the absorbing interest of Darwin's book to what they discovered.
His work on climbing plants was his first sustained piece of work on the physiology of movement, and he remarks in 1864: "This has been new sort of work for me." ("Life and Letters", III. page 315. He had, however, made a beginning on the movements of Drosera.) He goes on to remark with something of surprise, "I have been pleased to find what a capital guide for observations a full conviction of the change of species is."
It was this point of view that enabled him to develop a broad conception of the power of climbing as an adaptation by means of which plants are enabled to reach the light. Instead of being compelled to construct a stem of sufficient strength to stand alone, they succeed in the struggle by making use of other plants as supports. He showed that the great class of tendril- and root-climbers which do not depend on twining round a pole, like a scarlet-runner, but on attaching themselves as they grow upwards, effect an economy. Thus a Phaseolus has to manufacture a stem three feet in length to reach a height of two feet above the ground, whereas a pea "which had ascended to the same height by the aid of its tendrils, was but little longer than the height reached." ("Climbing Plants" (2nd edition 1875), page 193.)
Thus he was led on to the belief that TWINING is the more ancient form of climbing, and that tendril-climbers have been developed from twiners. In accordance with this view we find LEAF-CLIMBERS, which may be looked on as incipient tendril-bearers, occurring in the same genera with simple twiners. (Loc. cit. page 195.) He called attention to the case of Maurandia semperflorens in which the young flower-stalks revolve spontaneously and are sensitive to a touch, but neither of these qualities is of any perceptible value to the species. This forced him to believe that in other young plants the rudiments of the faculty needed for twining would be found—a prophecy which he made good in his "Power of Movement" many years later.
In "Climbing Plants" he did little more than point out the remarkable fact that the habit of climbing is widely scattered through the vegetable kingdom. Thus climbers are to be found in 35 out of the 59 Phanerogamic Alliances of Lindley, so that "the conclusion is forced on our minds that the capacity of revolving (If a twining plant, e.g. a hop, is observed before it has begun to ascend a pole, it will be noticed that, owing to the curvature of the stem, the tip is not vertical but hangs over in a roughly horizontal position. If such a shoot is watched it will be found that if, for instance, it points to the north at a given hour, it will be found after a short interval pointing north-east, then east, and after about two hours it will once more be looking northward. The curvature of the stem depends on one side growing quicker than the opposite side, and the revolving movement, i.e. circumnutation, depends on the region of quickest growth creeping gradually round the stem from south through west to south again. Other plants, e.g. Phaseolus, revolve in the opposite direction.), on which most climbers depend, is inherent, though undeveloped, in almost every plant in the vegetable kingdom." ("Climbing Plants", page 205.)
In the "Origin" (Edition I. page 427, Edition VI. page 374.) Darwin speaks of the "apparent paradox, that the very same characters are analogical when one class or order is compared with another, but give true affinities when the members of the same class or order are compared one with another." In this way we might perhaps say that the climbing of an ivy and a hop are analogical; the resemblance depending on the adaptive result rather than on community of blood; whereas the relation between a leaf-climber and a true tendril-bearer reveals descent. This particular resemblance was one in which my father took especial delight. He has described an interesting case occurring in the Fumariaceae. ("Climbing Plants", page 195.) "The terminal leaflets of the leaf-climbing Fumaria officinalis are not smaller than the other leaflets; those of the leaf-climbing Adlumia cirrhosa are greatly reduced; those of Corydalis claviculata (a plant which may be indifferently called a leaf-climber or a tendril-bearer) are either reduced to microscopical dimensions or have their blades wholly aborted, so that this plant is actually in a state of transition; and finally in the Dicentra the tendrils are perfectly characterized."
It is a remarkable fact that the quality which, broadly speaking, forms the basis of the climbing habit (namely revolving nutation, otherwise known as circumnutation) subserves two distinct ends. One of these is the finding of a support, and this is common to twiners and tendrils. Here the value ends as far as tendril-climbers are concerned, but in twiners Darwin believed that the act of climbing round a support is a continuation of the revolving movement (circumnutation). If we imagine a man swinging a rope round his head and if we suppose the rope to strike a vertical post, the free end will twine round it. This may serve as a rough model of twining as explained in the "Movements and Habits of Climbing Plants". It is on these points—the nature of revolving nutation and the mechanism of twining—that modern physiologists differ from Darwin. (See the discussion in Pfeffer's "The Physiology of Plants" Eng. Tr. (Oxford, 1906), III. page 34, where the literature is given. Also Jost, "Vorlesungen uber Pflanzenphysiologie", page 562, Jena, 1904.)
Their criticism originated in observations made on a revolving shoot which is removed from the action of gravity by keeping the plant slowly rotating about a horizontal axis by means of the instrument known as a klinostat. Under these conditions circumnutation becomes irregular or ceases altogether. When the same experiment is made with a plant which has twined spirally up a stick, the process of climbing is checked and the last few turns become loosened or actually untwisted. From this it has been argued that Darwin was wrong in his description of circumnutation as an automatic change in the region of quickest growth. When the free end of a revolving shoot points towards the north there is no doubt that the south side has been elongating more than the north; after a time it is plain from the shoot hanging over to the east that the west side of the plant has grown most, and so on. This rhythmic change of the position of the region of greatest growth Darwin ascribes to an unknown internal regulating power. Some modern physiologists, however, attempt to explain the revolving movement as due to a particular form of sensitiveness to gravitation which it is not necessary to discuss in detail in this place. It is sufficient for my purpose to point out that Darwin's explanation of circumnutation is not universally accepted. Personally I believe that circumnutation is automatic—is primarily due to internal stimuli. It is however in some way connected with gravitational sensitiveness, since the movement normally occurs round a vertical line. It is not unnatural that, when the plant has no external stimulus by which the vertical can be recognised, the revolving movement should be upset.
Very much the same may be said of the act of twining, namely that most physiologists refuse to accept Darwin's view (above referred to) that twining is the direct result of circumnutation. Everyone must allow that the two phenomena are in some way connected, since a plant which circumnutates clockwise, i.e. with the sun, twines in the same direction, and vice versa. It must also be granted that geotropism has a bearing on the problem, since all plants twine upwards, and cannot twine along a horizontal support. But how these two factors are combined, and whether any (and if so what) other factors contribute, we cannot say. If we give up Darwin's explanation, we must at the same time say with Pfeffer that "the causes of twining are... unknown." ("The Physiology of Plants", Eng. Tr. (Oxford, 1906), III. page 37.)
Let us leave this difficult question and consider some other points made out in the progress of the work on climbing plants. One result of what he called his "niggling" ("Life and Letters", III. page 312.) work on tendrils was the discovery of the delicacy of their sense of touch, and the rapidity of their movement. Thus in a passion-flower tendril, a bit of platinum wire weighing 1.2 mg. produced curvature ("Climbing Plants", page 171.), as did a loop of cotton weighing 2 mg. Pfeffer ("Untersuchungen a.d. Bot. Inst. z. Tubingen", Bd. I. 1881-85, page 506.), however, subsequently found much greater sensitiveness: thus the tendril of Sicyos angulatus reacted to 0.00025 mg., but this only occurred when the delicate rider of cottonwool fibre was disturbed by the wind. The same author expanded and explained in a most interesting way the meaning of Darwin's observation that tendrils are not stimulated to movement by drops of water resting on them. Pfeffer showed that DIRTY water containing minute particles of clay in suspension acts as a stimulus. He also showed that gelatine acts like pure water; if a smooth glass rod is coated with a 10 per cent solution of gelatine and is then applied to a tendril, no movement occurs in spite of the fact that the gelatine is solid when cold. Pfeffer ("Physiology", Eng. Tr. III. page 52. Pfeffer has pointed out the resemblance between the contact irritability of plants and the human sense of touch. Our skin is not sensitive to uniform pressure such as is produced when the finger is dipped into mercury (Tubingen "Untersuchungen", I. page 504.) generalises the result in the statement that the tendril has a special form of irritability and only reacts to "differences of pressure or variations of pressure in contiguous... regions." Darwin was especially interested in such cases of specialised irritability. For instance in May, 1864, he wrote to Asa Gray ("Life and Letters", III. page 314.) describing the tendrils of Bignonia capreolata, which "abhor a simple stick, do not much relish rough bark, but delight in wool or moss." He received, from Gray, information as to the natural habitat of the species, and finally concluded that the tendrils "are specially adapted to climb trees clothed with lichens, mosses, or other such productions." ("Climbing Plants", page 102.)
Tendrils were not the only instance discovered by Darwin of delicacy of touch in plants. In 1860 he had already begun to observe Sundew (Drosera), and was full of astonishment at its behaviour. He wrote to Sir Joseph Hooker ("Life and Letters", III. page 319.): "I have been working like a madman at Drosera. Here is a fact for you which is certain as you stand where you are, though you won't believe it, that a bit of hair 1/78000 of one grain in weight placed on gland, will cause ONE of the gland-bearing hairs of Drosera to curve inwards." Here again Pfeffer (Pfeffer in "Untersuchungen a. d. Bot. Inst. z. Tubingen", I. page 491.) has, as in so many cases, added important facts to my father's observations. He showed that if the leaf of Drosera is entirely freed from such vibrations as would reach it if observed on an ordinary table, it does not react to small weights, so that in fact it was the vibration of the minute fragment of hair on the gland that produced movement. We may fancifully see an adaptation to the capture of insects—to the dancing of a gnat's foot on the sensitive surface.
Darwin was fond of telling how when he demonstrated the sensitiveness of Drosera to Mr Huxley and (I think) to Sir John Burdon Sanderson, he could perceive (in spite of their courtesy) that they thought the whole thing a delusion. And the story ended with his triumph when Mr Huxley cried out, "It IS moving."
Darwin's work on tendrils has led to some interesting investigations on the mechanisms by which plants perceive stimuli. Thus Pfeffer (Tubingen "Untersuchungen" I. page 524.) showed that certain epidermic cells occurring in tendrils are probably organs of touch. In these cells the protoplasm burrows as it were into cavities in the thickness of the external cell-walls and thus comes close to the surface, being separated from an object touching the tendril merely by a very thin layer of cell-wall substance. Haberlandt ("Physiologische Pflanzenanatomie", Edition III. Leipzig, 1904. "Sinnesorgane im Pflanzenreich", Leipzig, 1901, and other publications.) has greatly extended our knowledge of vegetable structure in relation to mechanical stimulation. He defines a sense-organ as a contrivance by which the DEFORMATION or forcible change of form in the protoplasm—on which mechanical stimulation depends—is rendered rapid and considerable in amplitude ("Sinnesorgane", page 10). He has shown that in certain papillose and bristle-like contrivances, plants possess such sense-organs; and moreover that these contrivances show a remarkable similarity to corresponding sense-organs in animals.
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.
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.)