The Origin of Species by Means of Natural Selection

Although the belief that an organ so perfect as the eye could have been formed by natural selection, is enough to stagger any one; yet in the case of any organ, if we know of a long series of gradations in complexity, each good for its possessor, then under changing conditions of life, there is no logical impossibility in the acquirement of any conceivable degree of perfection through natural selection. In the cases in which we know of no intermediate or transitional states, we should be extremely cautious in concluding that none can have existed, for the metamorphoses of many organs show what wonderful changes in function are at least possible. For instance, a swim-bladder has apparently been converted into an air-breathing lung. The same organ having performed simultaneously very different functions, and then having been in part or in whole specialised for one function; and two distinct organs having performed at the same time the same function, the one having been perfected whilst aided by the other, must often have largely facilitated transitions.

We have seen that in two beings widely remote from each other in the natural scale, organs serving for the same purpose and in external appearance closely similar may have been separately and independently formed; but when such organs are closely examined, essential differences in their structure can almost always be detected; and this naturally follows from the principle of natural selection. On the other hand, the common rule throughout nature is infinite diversity of structure for gaining the same end; and this again naturally follows from the same great principle.

In many cases we are far too ignorant to be enabled to assert that a part or organ is so unimportant for the welfare of a species, that modifications in its structure could not have been slowly accumulated by means of natural selection. In many other cases, modifications are probably the direct result of the laws of variation or of growth, independently of any good having been thus gained. But even such structures have often, as we may feel assured, been subsequently taken advantage of, and still further modified, for the good of species under new conditions of life. We may, also, believe that a part formerly of high importance has frequently been retained (as the tail of an aquatic animal by its terrestrial descendants), though it has become of such small importance that it could not, in its present state, have been acquired by means of natural selection.

Natural selection can produce nothing in one species for the exclusive good or injury of another; though it may well produce parts, organs, and excretions highly useful or even indispensable, or highly injurious to another species, but in all cases at the same time useful to the possessor. In each well-stocked country natural selection acts through the competition of the inhabitants and consequently leads to success in the battle for life, only in accordance with the standard of that particular country. Hence the inhabitants of one country, generally the smaller one, often yield to the inhabitants of another and generally the larger country. For in the larger country there will have existed more individuals, and more diversified forms, and the competition will have been severer, and thus the standard of perfection will have been rendered higher. Natural selection will not necessarily lead to absolute perfection; nor, as far as we can judge by our limited faculties, can absolute perfection be everywhere predicated.

On the theory of natural selection we can clearly understand the full meaning of that old canon in natural history, "Natura non facit saltum." This canon, if we look to the present inhabitants alone of the world, is not strictly correct; but if we include all those of past times, whether known or unknown, it must on this theory be strictly true.

It is generally acknowledged that all organic beings have been formed on two great laws – Unity of Type, and the Conditions of Existence. By unity of type is meant that fundamental agreement in structure which we see in organic beings of the same class, and which is quite independent of their habits of life. On my theory, unity of type is explained by unity of descent. The expression of conditions of existence, so often insisted on by the illustrious Cuvier, is fully embraced by the principle of natural selection. For natural selection acts by either now adapting the varying parts of each being to its organic and inorganic conditions of life; or by having adapted them during past periods of time: the adaptations being aided in many cases by the increased use or disuse of parts, being affected by the direct action of external conditions of life, and subjected in all cases to the several laws of growth and variation. Hence, in fact, the law of the Conditions of Existence is the higher law; as it includes, through the inheritance of former variations and adaptations, that of Unity of Type.

CHAPTER VII. MISCELLANEOUS OBJECTIONS TO THE THEORY OF NATURAL SELECTION

Longevity – Modifications not necessarily simultaneous – Modifications apparently of no direct service – Progressive development – Characters of small functional importance, the most constant – Supposed incompetence of natural selection to account for the incipient stages of useful structures – Causes which interfere with the acquisition through natural selection of useful structures – Gradations of structure with changed functions – Widely different organs in members of the same class, developed from one and the same source – Reasons for disbelieving in great and abrupt modifications.

I will devote this chapter to the consideration of various miscellaneous objections which have been advanced against my views, as some of the previous discussions may thus be made clearer; but it would be useless to discuss all of them, as many have been made by writers who have not taken the trouble to understand the subject. Thus a distinguished German naturalist has asserted that the weakest part of my theory is, that I consider all organic beings as imperfect: what I have really said is, that all are not as perfect as they might have been in relation to their conditions; and this is shown to be the case by so many native forms in many quarters of the world having yielded their places to intruding foreigners. Nor can organic beings, even if they were at any one time perfectly adapted to their conditions of life, have remained so, when their conditions changed, unless they themselves likewise changed; and no one will dispute that the physical conditions of each country, as well as the number and kinds of its inhabitants, have undergone many mutations.

A critic has lately insisted, with some parade of mathematical accuracy, that longevity is a great advantage to all species, so that he who believes in natural selection "must arrange his genealogical tree" in such a manner that all the descendants have longer lives than their progenitors! Cannot our critics conceive that a biennial plant or one of the lower animals might range into a cold climate and perish there every winter; and yet, owing to advantages gained through natural selection, survive from year to year by means of its seeds or ova? Mr. E. Ray Lankester has recently discussed this subject, and he concludes, as far as its extreme complexity allows him to form a judgment, that longevity is generally related to the standard of each species in the scale of organisation, as well as to the amount of expenditure in reproduction and in general activity. And these conditions have, it is probable, been largely determined through natural selection.

It has been argued that, as none of the animals and plants of Egypt, of which we know anything, have changed during the last three or four thousand years, so probably have none in any part of the world. But, as Mr. G.H. Lewes has remarked, this line of argument proves too much, for the ancient domestic races figured on the Egyptian monuments, or embalmed, are closely similar or even identical with those now living; yet all naturalists admit that such races have been produced through the modification of their original types. The many animals which have remained unchanged since the commencement of the glacial period, would have been an incomparably stronger case, for these have been exposed to great changes of climate and have migrated over great distances; whereas, in Egypt, during the last several thousand years, the conditions of life, as far as we know, have remained absolutely uniform. The fact of little or no modification having been effected since the glacial period, would have been of some avail against those who believe in an innate and necessary law of development, but is powerless against the doctrine of natural selection or the survival of the fittest, which implies that when variations or individual differences of a beneficial nature happen to arise, these will be preserved; but this will be effected only under certain favourable circumstances.

The celebrated palaeontologist, Bronn, at the close of his German translation of this work, asks how, on the principle of natural selection, can a variety live side by side with the parent species? If both have become fitted for slightly different habits of life or conditions, they might live together; and if we lay on one side polymorphic species, in which the variability seems to be of a peculiar nature, and all mere temporary variations, such as size, albinism, etc., the more permanent varieties are generally found, as far as I can discover, inhabiting distinct stations, such as high land or low land, dry or moist districts. Moreover, in the case of animals which wander much about and cross freely, their varieties seem to be generally confined to distinct regions.

Bronn also insists that distinct species never differ from each other in single characters, but in many parts; and he asks, how it always comes that many parts of the organisation should have been modified at the same time through variation and natural selection? But there is no necessity for supposing that all the parts of any being have been simultaneously modified. The most striking modifications, excellently adapted for some purpose, might, as was formerly remarked, be acquired by successive variations, if slight, first in one part and then in another; and as they would be transmitted all together, they would appear to us as if they had been simultaneously developed. The best answer, however, to the above objection is afforded by those domestic races which have been modified, chiefly through man's power of selection, for some special purpose. Look at the race and dray-horse, or at the greyhound and mastiff. Their whole frames, and even their mental characteristics, have been modified; but if we could trace each step in the history of their transformation – and the latter steps can be traced – we should not see great and simultaneous changes, but first one part and then another slightly modified and improved. Even when selection has been applied by man to some one character alone – of which our cultivated plants offer the best instances – it will invariably be found that although this one part, whether it be the flower, fruit, or leaves, has been greatly changed, almost all the other parts have been slightly modified. This may be attributed partly to the principle of correlated growth, and partly to so-called spontaneous variation.

A much more serious objection has been urged by Bronn, and recently by Broca, namely, that many characters appear to be of no service whatever to their possessors, and therefore cannot have been influenced through natural selection. Bronn adduces the length of the ears and tails in the different species of hares and mice – the complex folds of enamel in the teeth of many animals, and a multitude of analogous cases. With respect to plants, this subject has been discussed by Nageli in an admirable essay. He admits that natural selection has effected much, but he insists that the families of plants differ chiefly from each other in morphological characters, which appear to be quite unimportant for the welfare of the species. He consequently believes in an innate tendency towards progressive and more perfect development. He specifies the arrangement of the cells in the tissues, and of the leaves on the axis, as cases in which natural selection could not have acted. To these may be added the numerical divisions in the parts of the flower, the position of the ovules, the shape of the seed, when not of any use for dissemination, etc.

There is much force in the above objection. Nevertheless, we ought, in the first place, to be extremely cautious in pretending to decide what structures now are, or have formerly been, of use to each species. In the second place, it should always be borne in mind that when one part is modified, so will be other parts, through certain dimly seen causes, such as an increased or diminished flow of nutriment to a part, mutual pressure, an early developed part affecting one subsequently developed, and so forth – as well as through other causes which lead to the many mysterious cases of correlation, which we do not in the least understand. These agencies may be all grouped together, for the sake of brevity, under the expression of the laws of growth. In the third place, we have to allow for the direct and definite action of changed conditions of life, and for so-called spontaneous variations, in which the nature of the conditions apparently plays a quite subordinate part. Bud-variations, such as the appearance of a moss-rose on a common rose, or of a nectarine on a peach-tree, offer good instances of spontaneous variations; but even in these cases, if we bear in mind the power of a minute drop of poison in producing complex galls, we ought not to feel too sure that the above variations are not the effect of some local change in the nature of the sap, due to some change in the conditions. There must be some efficient cause for each slight individual difference, as well as for more strongly marked variations which occasionally arise; and if the unknown cause were to act persistently, it is almost certain that all the individuals of the species would be similarly modified.

In the earlier editions of this work I underrated, as it now seems probable, the frequency and importance of modifications due to spontaneous variability. But it is impossible to attribute to this cause the innumerable structures which are so well adapted to the habits of life of each species. I can no more believe in this than that the well-adapted form of a race-horse or greyhound, which before the principle of selection by man was well understood, excited so much surprise in the minds of the older naturalists, can thus be explained.

It may be worth while to illustrate some of the foregoing remarks. With respect to the assumed inutility of various parts and organs, it is hardly necessary to observe that even in the higher and best-known animals many structures exist, which are so highly developed that no one doubts that they are of importance, yet their use has not been, or has only recently been, ascertained. As Bronn gives the length of the ears and tail in the several species of mice as instances, though trifling ones, of differences in structure which can be of no special use, I may mention that, according to Dr. Schobl, the external ears of the common mouse are supplied in an extraordinary manner with nerves, so that they no doubt serve as tactile organs; hence the length of the ears can hardly be quite unimportant. We shall, also, presently see that the tail is a highly useful prehensile organ to some of the species; and its use would be much influence by its length.

With respect to plants, to which on account of Nageli's essay I shall confine myself in the following remarks, it will be admitted that the flowers of the orchids present a multitude of curious structures, which a few years ago would have been considered as mere morphological differences without any special function; but they are now known to be of the highest importance for the fertilisation of the species through the aid of insects, and have probably been gained through natural selection. No one until lately would have imagined that in dimorphic and trimorphic plants the different lengths of the stamens and pistils, and their arrangement, could have been of any service, but now we know this to be the case.

In certain whole groups of plants the ovules stand erect, and in others they are suspended; and within the same ovarium of some few plants, one ovule holds the former and a second ovule the latter position. These positions seem at first purely morphological, or of no physiological signification; but Dr. Hooker informs me that within the same ovarium the upper ovules alone in some cases, and in others the lower ones alone are fertilised; and he suggests that this probably depends on the direction in which the pollen-tubes enter the ovarium. If so, the position of the ovules, even when one is erect and the other suspended within the same ovarium, would follow the selection of any slight deviations in position which favoured their fertilisation, and the production of seed.

Several plants belonging to distinct orders habitually produce flowers of two kinds – the one open, of the ordinary structure, the other closed and imperfect. These two kinds of flowers sometimes differ wonderfully in structure, yet may be seen to graduate into each other on the same plant. The ordinary and open flowers can be intercrossed; and the benefits which certainly are derived from this process are thus secured. The closed and imperfect flowers are, however, manifestly of high importance, as they yield with the utmost safety a large stock of seed, with the expenditure of wonderfully little pollen. The two kinds of flowers often differ much, as just stated, in structure. The petals in the imperfect flowers almost always consist of mere rudiments, and the pollen-grains are reduced in diameter. In Ononis columnae five of the alternate stamens are rudimentary; and in some species of Viola three stamens are in this state, two retaining their proper function, but being of very small size. In six out of thirty of the closed flowers in an Indian violet (name unknown, for the plants have never produced with me perfect flowers), the sepals are reduced from the normal number of five to three. In one section of the Malpighiaceae the closed flowers, according to A. de Jussieu, are still further modified, for the five stamens which stand opposite to the sepals are all aborted, a sixth stamen standing opposite to a petal being alone developed; and this stamen is not present in the ordinary flowers of this species; the style is aborted; and the ovaria are reduced from three to two. Now although natural selection may well have had the power to prevent some of the flowers from expanding, and to reduce the amount of pollen, when rendered by the closure of the flowers superfluous, yet hardly any of the above special modifications can have been thus determined, but must have followed from the laws of growth, including the functional inactivity of parts, during the progress of the reduction of the pollen and the closure of the flowers.

It is so necessary to appreciate the important effects of the laws of growth, that I will give some additional cases of another kind, namely of differences in the same part or organ, due to differences in relative position on the same plant. In the Spanish chestnut, and in certain fir-trees, the angles of divergence of the leaves differ, according to Schacht, in the nearly horizontal and in the upright branches. In the common rue and some other plants, one flower, usually the central or terminal one, opens first, and has five sepals and petals, and five divisions to the ovarium; while all the other flowers on the plant are tetramerous. In the British Adoxa the uppermost flower generally has two calyx-lobes with the other organs tetramerous, while the surrounding flowers generally have three calyx-lobes with the other organs pentamerous. In many Compositae and Umbelliferae (and in some other plants) the circumferential flowers have their corollas much more developed than those of the centre; and this seems often connected with the abortion of the reproductive organs. It is a more curious fact, previously referred to, that the achenes or seeds of the circumference and centre sometimes differ greatly in form, colour and other characters. In Carthamus and some other Compositae the central achenes alone are furnished with a pappus; and in Hyoseris the same head yields achenes of three different forms. In certain Umbelliferae the exterior seeds, according to Tausch, are orthospermous, and the central one coelospermous, and this is a character which was considered by De Candolle to be in other species of the highest systematic importance. Professor Braun mentions a Fumariaceous genus, in which the flowers in the lower part of the spike bear oval, ribbed, one-seeded nutlets; and in the upper part of the spike, lanceolate, two-valved and two-seeded siliques. In these several cases, with the exception of that of the well-developed ray-florets, which are of service in making the flowers conspicuous to insects, natural selection cannot, as far as we can judge, have come into play, or only in a quite subordinate manner. All these modifications follow from the relative position and inter-action of the parts; and it can hardly be doubted that if all the flowers and leaves on the same plant had been subjected to the same external and internal condition, as are the flowers and leaves in certain positions, all would have been modified in the same manner.

In numerous other cases we find modifications of structure, which are considered by botanists to be generally of a highly important nature, affecting only some of the flowers on the same plant, or occurring on distinct plants, which grow close together under the same conditions. As these variations seem of no special use to the plants, they cannot have been influenced by natural selection. Of their cause we are quite ignorant; we cannot even attribute them, as in the last class of cases, to any proximate agency, such as relative position. I will give only a few instances. It is so common to observe on the same plant, flowers indifferently tetramerous, pentamerous, etc., that I need not give examples; but as numerical variations are comparatively rare when the parts are few, I may mention that, according to De Candolle, the flowers of Papaver bracteatum offer either two sepals with four petals (which is the common type with poppies), or three sepals with six petals. The manner in which the petals are folded in the bud is in most groups a very constant morphological character; but Professor Asa Gray states that with some species of Mimulus, the aestivation is almost as frequently that of the Rhinanthideae as of the Antirrhinideae, to which latter tribe the genus belongs. Aug. St. Hilaire gives the following cases: the genus Zanthoxylon belongs to a division of the Rutaceae with a single ovary, but in some species flowers may be found on the same plant, and even in the same panicle, with either one or two ovaries. In Helianthemum the capsule has been described as unilocular or tri-locular; and in H. mutabile, "Une lame PLUS OU MOINS LARGE, s'etend entre le pericarpe et le placenta." In the flowers of Saponaria officinalis Dr. Masters has observed instances of both marginal and free central placentation. Lastly, St. Hilaire found towards the southern extreme of the range of Gomphia oleaeformis two forms which he did not at first doubt were distinct species, but he subsequently saw them growing on the same bush; and he then adds, "Voila donc dans un meme individu des loges et un style qui se rattachent tantot a un axe verticale et tantot a un gynobase."

We thus see that with plants many morphological changes may be attributed to the laws of growth and the inter-action of parts, independently of natural selection. But with respect to Nageli's doctrine of an innate tendency towards perfection or progressive development, can it be said in the case of these strongly pronounced variations, that the plants have been caught in the act of progressing towards a higher state of development? On the contrary, I should infer from the mere fact of the parts in question differing or varying greatly on the same plant, that such modifications were of extremely small importance to the plants themselves, of whatever importance they may generally be to us for our classifications. The acquisition of a useless part can hardly be said to raise an organism in the natural scale; and in the case of the imperfect, closed flowers, above described, if any new principle has to be invoked, it must be one of retrogression rather than of progression; and so it must be with many parasitic and degraded animals. We are ignorant of the exciting cause of the above specified modifications; but if the unknown cause were to act almost uniformly for a length of time, we may infer that the result would be almost uniform; and in this case all the individuals of the species would be modified in the same manner.