Essays Upon Heredity and Kindred Biological Problems

In this sense I trust that the sign-post or compass which I offer may be accepted. Even though it should be its fate to be replaced by a better one at a later period, it will have fulfilled its object if it enables science to advance for even a short distance.

APPENDICES

Appendix I. Further considerations which oppose Nägeli’s

explanation of transformation as due to internal causes199

When I describe Nägeli’s theory of transformation as due to active causes lying within the organism, as a phyletic force of transformation, I do not mean to imply that it is one of those mysterious principles which, according to some writers, constitute the unconscious cause which directs the transformation of species. Nägeli’s idioplasm, which changes from within itself, is conceived as a thoroughly scientific, mechanically operating principle. This cause is undoubtedly capable of theoretical conception: the only question is whether it has any real existence. According to Nägeli, the growing organic substance, the idioplasm, not only represents a perpetuum mobile rendered possible as long as its substance continually receives from without the matter and force which are necessary for continuous growth, but it also represents a perpetuum variabile due to the action of internal causes200. But this is just the doubtful point, viz., whether the structure of the idioplasm itself compels it to change gradually during the course of its growth, or whether it is not rather the external conditions which compel the ever slightly varying idioplasm to change in a certain direction by the summation of small differences. It has been shown above that we do not gain anything by adopting Nägeli’s theory, because the main problem which organic nature offers for our solution, viz. adaptation, remains unsolved. Hence this theory does not explain the phenomena of nature, and I believe that there are also certain facts which are directly antagonistic to it.

If the idioplasm really possessed the power of spontaneous variability ascribed to it by Nägeli; if, as a result of its own growth, it were compelled to undergo gradual changes, and thus to produce new species, we should expect that the duration of species, genera, orders, &c. would be of approximately equal length respectively, at least in forms of equal structural complexity. The time required by the idioplasm to undergo such changes as would constitute transformation into a new species ought to be always the same at equal heights in the scale of organization, that is, with equal complexity in the molecular structure of the idioplasm. It appears to me to be a necessary consequence of Nägeli’s theory that the causes of transformation lie solely in this molecular structure of the idioplasm. If nothing more than a certain amount of growth, and consequently a certain period of time during which the organism reproduces itself with a certain intensity, is required to produce a change in the idioplasm, then we must conclude that the alteration in the latter must take place when this certain amount of growth has been reached, or after this certain period has elapsed. In other words, the time during which a species exists—from its origin as a modification of some older species, until its own transformation into a new one—must be the same in species with the same degree of organization. But the facts are very far from supporting this consequence of Nägeli’s theory. The duration of species is excessively variable: many arise and perish within the limits of a single geological formation, while others may be restricted to a very small part of a formation; others again may last through several formations. It must be admitted that we cannot estimate the exact position of extinct species in the scale of organization, and the differences may therefore depend upon differences of organization: or they may be explained by the supposition that certain species may have become incapable of transformation, and might, under favourable conditions, continue to exist for an indefinite period. But this reply would introduce a new hypothesis in direct antagonism to Nägeli’s theory, which assumes that the variability of idioplasm takes place as the consequence of mere growth, and necessarily depends upon molecular structure. Nägeli himself asserts that the essential substance (idioplasm) of the descendants of the earliest forms of life is in a state of perpetual change, which would continue even if the series of successive generations were indefinitely prolonged201. Hence there can be no rest in the process of change which the idioplasm must undergo; and this is as true of each single species as it is of the organic world taken as a whole. We could, perhaps, find shelter in the insufficiency of our geological knowledge, but the number of ascertained facts is too great for this to be possible. Thus it is well known that the genus Nautilus has lasted from Silurian times, through all the three geological periods, up to the present day: while all its Silurian allies (Orthoceras, Gomphoceras, Goniatites, &c.) became extinct at a comparatively early period.

A keen and clever controversialist might still bring forward many objections against such an argument. I do not therefore place too much dependence upon the geological facts by themselves, as a disproof of the self-variability of Nägeli’s idioplasm; for it must be admitted that the facts are not sufficiently complete for this purpose. For instance, in the case of Nautilus it might be argued that we do not know anything about the fossil Cephalopods of pre-Silurian times, and that it is therefore possible that the above-mentioned allies of Nautilus may have existed previously for as long a period as that through which Nautilus has lived in post-Silurian time. However this may be, it will be at least conceded that the geological facts do not lend any support to Nägeli’s theory, for we can see no trace of even an approximately regular succession of forms.

Appendix II. Nägeli’s explanation of adaptation202

In order to explain adaptation Nägeli assumes that, under certain circumstances, external influences may cause slight permanent changes in the idioplasm. If then such influences act continually in the same direction during long periods of time, the changes in the idioplasm may increase to a perceptible amount, i. e. to a degree which makes itself felt in visible external characters203. But such changes alone could not be considered as adaptations, for the essential character of an adaptation is that it must be a purposeful change. Nägeli, however, brings forward the fact that external stimuli often produce their chief effects at that very part of the organism to which the stimuli themselves were applied. ‘If the results are detrimental, the organism attempts to defend itself against the stimulus: a confluence of nutrient fluid takes place towards the part upon which the stimulus has acted, and new tissues are formed which restore the integrity of the organism by replacing the lost structures as far as possible. Thus in plants the healthy tissues begin to grow actively around the seat of an injury, tending to close it up, and to afford protection by impenetrable layers of cork.’ Purposeful reactions of this kind are certainly common in the organic world, occurring in animals as well as in plants. Thus in the human body an injury causes a rapid growth of the surrounding tissues, which leads to the closing-up of the wound; while in the Salamander even the amputated leg or tail is replaced by growth. An extreme example of these purposeful reactions is afforded by the tree-frog (Hyla), which is of a light-green colour when seated upon a light-green leaf, but becomes dark brown when transferred to dark surroundings. Hence this animal adapts itself to the colour of its environment, and thus gains protection from its enemies.

Admitting this capability on the part of organisms to react under certain stimuli in a purposeful manner, the question remains whether such a power is a primitive original quality belonging to the essential nature of each organism. The power of changing the colour of the skin in correspondence with that of the surroundings is not very common in the animal kingdom. In the frog this power depends upon a highly complex reflex mechanism. Certain chromatophores in the skin are connected with nerves204 which pass to the brain and are there brought into relation, by means of nerve-cells, with the nervous centres of the organ of vision. The relation is of such a kind that strong light falling upon the retina constitutes a stimulus for the production of an impulse, which is conducted, along the previously mentioned motor nerves, from the brain to the chromatophores, thus determining the contraction of these latter and the consequent appearance of a light-coloured skin. When the strong stimulus (of light) ceases, the chromatophores expand again, and the skin becomes dark. That the chromatophores do not themselves react upon the direct stimulus of light was proved by Lister205, who showed that blind frogs do not possess the power of altering their colour in correspondence with that of their environment. It is quite obvious that in this case we are not dealing with a primary, but with a secondarily produced character; and it has yet to be proved that all the purposeful reactions mentioned by Nägeli are not similarly secondary characters or adaptations, and thus very far from being primitive qualities of the organic substance of the forms in which they occur.

I do not by any means doubt that some of the reactions witnessed in organisms do not depend upon adaptation, but such reactions are not usually purposeful. Curiously enough, Nägeli mentions the formation of galls in plants among his instances of purposeful reactions under external stimuli. I think, however, that it can hardly be maintained that the galls are of any use to the plant: on the contrary, they may even be very injurious to it. The gall is only useful to the insect which it protects and supplies with food. The recent and most excellent investigations of Adler206 and of Beyerinck207 have shown that the puncture made by the Cynips in depositing its eggs is not the stimulus which produces the gall, as was formerly believed to be the case, but that such a stimulus is provided by the larva which developes from the egg. The presence of this small, actively moving, foreign body stimulates the tissue of the plant in a definite manner, always producing a result which is advantageous to the larva and not to the plant. It would be to the advantage of the latter if it killed the intruding larva, either enclosing it by woody tissue devoid of nourishment, or poisoning it by some acrid secretion, or simply crushing it by the active growth of the surrounding tissues. But nothing of the kind occurs: in fact an active growth of cells (forming the so-called ‘Blastem’ of Beyerinck) takes place around the embryo, while it is still enclosed in the egg-capsule; but the growth is not such as to crush the embryo, which remains free in the cavity, the so-called larval chamber, which is formed around it. It would be out of place to discuss here the question as to how we can conceive that the plant is thus compelled to produce a growth which is at any rate indifferent and may be injurious to it; and which, moreover, is exactly adapted to the needs of its insect-enemy. But it is at all events obvious that this cannot be an example of a self-protecting reaction under a stimulus, and that therefore an organism does not always respond to external stimuli in a manner useful to itself.

But even if we could accept the suggestion that the purposeful reaction of an organism under stimulation is a primary and not a secondarily produced character, such a principle would by no means suffice for the explanation of existing adaptations. Nägeli attempts to explain certain selected cases of adaptation as the direct results of external stimuli. He looks upon the thick hairy coat of mammals in arctic regions, and the winter covering of animals in temperate regions, as a direct reaction of the skin under the influence of cold. He considers that the horns, claws, and tusks of animals have arisen directly as reactions under stimuli applied to certain parts of the surface of the body in attack and defence208. This interpretation is similar to that offered by Lamarck at the beginning of this century. At first sight such a suggestion appears to be plausible, for the acquisition of a thick hairy covering by the mammals of temperate regions is actually contemporaneous with the cold season of the year. But the question arises as to whether the production of a larger number of hairs at the beginning of winter is not merely another instance of a secondary character, like the assumption of a green colour by the tree-frog under the stimulus exerted by strong light.

In the case of the hairy coat it is only necessary to produce a larger number of structures such as had existed previously; but how can it have been possible for the petals of flowers, with their peculiar and complex forms, to have been developed from stamens as a direct result of the insects which visit them in order to obtain pollen and nectar? How could the creeping of these insects and the small punctures made by them constitute stimuli for the production of an increased rate of growth? And how is it possible in any way to explain, by mere increase in growth, the origin of a structure in which each part has its own distinct meaning and plays a peculiar part in attracting insects and in the process of cross-fertilization effected by them? Even if the manifold peculiarities of form could be explained in this way, how can such an explanation possibly hold for the colours of flowers? How could the white colour of flowers which open at night be explained as the direct result of the creeping of insects? How can the suggestion of such a cause offer any interpretation of the fact that flowers which open by day are tinted with various colours, or of the fact that there is often a bright or highly coloured spot which shows the way to the hidden nectary?

There are, moreover, a large number of very striking adaptations in form and colour, for which no stimulus acting directly upon the organism can be found. Can we imagine that the green caterpillar209, plant-bug, or grasshopper, sitting among green surroundings, is thus exposed to a stimulus which directly produces the green colour in the skin? Can the walking-stick insect, which resembles a brown twig, be subject to a transforming stimulus by sitting on such branches or by looking at them? Or again, if we consider the phenomena of mimicry, how can one species of butterfly, by flying about with another species, exercise upon the latter such an influence as to render it similar to the first in appearance? In many cases of mimicry, the mimicked and the mimicking species do not even live in the same place, as we see in the moths, flies, and beetles which resemble in appearance the much-dreaded wasps.

The interpretation of adaptation is the weak part of Nägeli’s theory, and it is somewhat remarkable that so acute a thinker should not have perceived this himself. One very nearly gains the impression that Nägeli does not wish to understand the theory of natural selection. He says, for instance, in speaking of the mutual adaptation observable between the proboscis, the so-called ‘tongue’ of butterflies, and flowers with tubular corolla210:—‘Among the most remarkable and commonest adaptations observable in the forms of flowers, are the corollas with long tubes considered in relation to the long “tongues” of insects, which suck the nectar from the bottom of the long narrow tubes, and at the same time effect the cross-fertilization of the plant. Both these arrangements have been gradually developed to their present degree of complexity—the long-tubed corollas from those without tubes, and from those with short ones, the long “tongues” from short ones. Undoubtedly both have been developed at the same rate so that the length of both sets of structures has always remained the same.’

No objection can be raised against these statements, but Nägeli goes on to say:—‘But how can such a process of development be explained by the theory of natural selection, for at each stage in the process the adaptation was invariably complete. The tube of the corolla and the “tongue” must have reached, for instance, at a certain time, a length of 5 or 10 mm. If now the tube of the corolla became longer in some plants, such an alteration would have been disadvantageous because the insects would be no longer able to obtain food from them, and would therefore visit flowers with shorter tubes. Hence, according to the theory of natural selection, the longer tubes ought to have disappeared. If on the other hand the “tongue” became longer in some insects, such a change would be superfluous and should have been given up, according to the same theory, as unnecessary structural waste. The simultaneous change in the two structures must, according to the theory of natural selection, be due to the same principle as that by which Münchhausen pulled himself out of a bog by means of his own pig-tail.’

But, according to the theory of natural selection, the case appears in a very different light from that in which it is put by Nägeli. The flower and the insect do not compete for the greater length of their respective organs: all through the gradual process, the flower is the first to lengthen its corolla and the butterfly follows. Their relation is not like that between a certain species of animal and another which serves as its prey, where each strives to be the quicker, so that the speed of both is increased to the greatest possible extent in the course of generations. Nor do they stand in the same relation as that obtaining between an insectivorous bird and a certain species of butterfly which forms its principal food; in such a case two totally different characters may be continually increased up to their highest point, e.g. in the butterfly similarity to the dead and fallen leaves among which it seeks protection when pursued, in the bird keenness of sight. As long as the latter quality is still capable of increase, so long will it still be advantageous to any individual butterfly to resemble the leaf a little more completely than other individuals of the same species; for it will thus be capable of escaping those birds which possess a rather keener sight than others. On the other hand, a bird with rather keener sight will have the greatest chance of catching the better protected butterflies. It is only in this way that we can explain the constant production of such extraordinary similarities between insects and leaves or other parts of plants. At every stage of growth both the insect and its pursuer are completely adapted to each other; i.e. they are so far protected and so far successful respectively, as is necessary to prevent that gradual decrease in the average number of individuals which would lead to the extermination of the species211. But the fact that there is complete adaptation at each stage does not prevent the two species from increasing those qualities of protection and of pursuit upon which they respectively depend. So far from this being the case, they would be necessarily compelled to gradually increase these qualities so long as the physical possibility of improvement remained on both sides. As long as some birds possessed a rather keener sight than those which previously existed, so long would those butterflies possess an advantage in which the resemblance to leaf-veining was more distinct than in others. But from the moment at which the maximum keenness of eyesight attainable had been reached, at which therefore all butterflies resembled leaves so completely that even the birds with the keenest eyesight might fail to detect them when at rest,—from this very point any further improvement in the similarity to leaves would cease, because the advantage to be gained from any such improvement would cease at the same time.

Such reciprocal intensification of adaptive characters appears to me to have been one of the most important factors in the transformation of species: it must have persisted through long series of species during phylogeny: it must have affected the most diverse parts and characters in the most diverse groups of organisms.

In certain large butterflies of the Indian and African forests—Kallima paralecta, K. inachis, and K. albofasciata—it has been frequently pointed out that the deceptive resemblance to a leaf is so striking that an observer who has received no hint upon the subject believes that he sees a leaf, even when he is looking at the butterfly very closely. The similarity is nevertheless incomplete; for out of sixteen specimens in the collections at Amsterdam and Leyden, I could not find a single one which had more than two lateral veins on one side of the mid-rib of the supposed leaf, or more than three upon the other side; while about six or seven veins should have been present on each side. But from two to three lateral veins are amply sufficient to produce a high degree of resemblance; in fact so much so that it is a matter for wonder as to how it has been possible for such a relatively perfect copy to have been produced; or how the sight of birds can have become so highly developed that while flying rapidly they could perceive the vein-like markings; or to state the case more accurately, that they could detect those individuals with a less number of veins than others. It is possible that the process of increase in resemblance is still proceeding in the species of the genus Kallima; at all events, I was struck by the rather strong individual differences in the markings of the supposed leaf.

On the other hand, the cause of the increase in length of the tubular corolla and of the butterfly’s ‘tongue,’ lies neither in the flower nor in the butterfly, but it is to be found in those other insects which visit the flower and steal its honey without being of any assistance in cross-fertilization. It may be stated shortly, that non-tubular corollas, with the honey freely exposed—for it must be assumed the ancestral form was of this kind—gradually developed into corollas with the honey deeply concealed. The whole process was presumably first started by the flower, for the gradual withdrawal of the honey to greater depths conferred the advantage of protection from rain (Hermann Müller), while larger quantities of honey could be stored up, and this would also increase the number of insects visiting the flower and render their visits more certain. As soon as this withdrawal occurred, the mouth-parts of insects began to be subjected to a selective process whereby these organs in some of them were lengthened at the same rate as that at which the honey was withdrawn. When once the process had begun, its continuance was ensured, for as soon as flower-frequenting insects were divided into two groups with short and with long mouth-parts respectively, a further increase in the length of the corolla-tube necessarily took place in all those flowers which were especially benefited by the assured visits of a relatively small number of species of insects, viz., those flowers in which cross-fertilization was more certainly performed in this way than by the uncertain visits of a great variety of species. This would imply that a still further increase in length would take place, for it is obvious that the cross-fertilization of any flower would be more certainly performed by an insect when the number of species of plants visited by it became less; and hence the cross-fertilization would be rendered most certain when the insect became completely adapted—in size, form, character of its surface, and the manner in which it obtained the honey—to the peculiarities of the flower. Those insects which obtain honey from a great variety of flowers are sure to waste a great part of the pollen by carrying it to the flowers of many different species, while insects which can only obtain honey from a few species of plants must necessarily visit many flowers of the same species one after the other, and they would therefore more generally distribute the pollen in an effective manner.

Hence the tube of the corolla, and the ‘tongue’ of the butterfly which brings about fertilization, would have continued to increase in length as long as it remained advantageous for the flower to exclude other less useful visitors, and as long as it was advantageous for the butterfly to secure the sole possession of the flower. Hence there is no competition between the flower and the butterfly which fertilizes it, but between these two on the one side, and the other would-be visitors of the flower on the other. Further details as to the advantages which the flower gains by excluding all other visitors, and the butterfly by being the only visitor of the flower, and also as to the manifold and elaborate mutual adaptations between insects and flowers, and as to the advantages and disadvantages which follow from the concealment of the honey—will be found in Hermann Müller’s212 work on the fertilization of flowers, in which all these subjects are minutely discussed, and are clearly explained in a most admirable manner.