The Origin of Species by Means of Natural Selection

In searching for the gradations through which an organ in any species has been perfected, we ought to look exclusively to its lineal progenitors; but this is scarcely ever possible, and we are forced to look to other species and genera of the same group, that is to the collateral descendants from the same parent-form, in order to see what gradations are possible, and for the chance of some gradations having been transmitted in an unaltered or little altered condition. But the state of the same organ in distinct classes may incidentally throw light on the steps by which it has been perfected.

The simplest organ which can be called an eye consists of an optic nerve, surrounded by pigment-cells and covered by translucent skin, but without any lens or other refractive body. We may, however, according to M. Jourdain, descend even a step lower and find aggregates of pigment-cells, apparently serving as organs of vision, without any nerves, and resting merely on sarcodic tissue. Eyes of the above simple nature are not capable of distinct vision, and serve only to distinguish light from darkness. In certain star-fishes, small depressions in the layer of pigment which surrounds the nerve are filled, as described by the author just quoted, with transparent gelatinous matter, projecting with a convex surface, like the cornea in the higher animals. He suggests that this serves not to form an image, but only to concentrate the luminous rays and render their perception more easy. In this concentration of the rays we gain the first and by far the most important step towards the formation of a true, picture-forming eye; for we have only to place the naked extremity of the optic nerve, which in some of the lower animals lies deeply buried in the body, and in some near the surface, at the right distance from the concentrating apparatus, and an image will be formed on it.

In the great class of the Articulata, we may start from an optic nerve simply coated with pigment, the latter sometimes forming a sort of pupil, but destitute of lens or other optical contrivance. With insects it is now known that the numerous facets on the cornea of their great compound eyes form true lenses, and that the cones include curiously modified nervous filaments. But these organs in the Articulata are so much diversified that Muller formerly made three main classes with seven subdivisions, besides a fourth main class of aggregated simple eyes.

When we reflect on these facts, here given much too briefly, with respect to the wide, diversified, and graduated range of structure in the eyes of the lower animals; and when we bear in mind how small the number of all living forms must be in comparison with those which have become extinct, the difficulty ceases to be very great in believing that natural selection may have converted the simple apparatus of an optic nerve, coated with pigment and invested by transparent membrane, into an optical instrument as perfect as is possessed by any member of the Articulata class.

He who will go thus far, ought not to hesitate to go one step further, if he finds on finishing this volume that large bodies of facts, otherwise inexplicable, can be explained by the theory of modification through natural selection; he ought to admit that a structure even as perfect as an eagle's eye might thus be formed, although in this case he does not know the transitional states. It has been objected that in order to modify the eye and still preserve it as a perfect instrument, many changes would have to be effected simultaneously, which, it is assumed, could not be done through natural selection; but as I have attempted to show in my work on the variation of domestic animals, it is not necessary to suppose that the modifications were all simultaneous, if they were extremely slight and gradual. Different kinds of modification would, also, serve for the same general purpose: as Mr. Wallace has remarked, "If a lens has too short or too long a focus, it may be amended either by an alteration of curvature, or an alteration of density; if the curvature be irregular, and the rays do not converge to a point, then any increased regularity of curvature will be an improvement. So the contraction of the iris and the muscular movements of the eye are neither of them essential to vision, but only improvements which might have been added and perfected at any stage of the construction of the instrument." Within the highest division of the animal kingdom, namely, the Vertebrata, we can start from an eye so simple, that it consists, as in the lancelet, of a little sack of transparent skin, furnished with a nerve and lined with pigment, but destitute of any other apparatus. In fishes and reptiles, as Owen has remarked, "The range of gradation of dioptric structures is very great." It is a significant fact that even in man, according to the high authority of Virchow, the beautiful crystalline lens is formed in the embryo by an accumulation of epidermic cells, lying in a sack-like fold of the skin; and the vitreous body is formed from embryonic subcutaneous tissue. To arrive, however, at a just conclusion regarding the formation of the eye, with all its marvellous yet not absolutely perfect characters, it is indispensable that the reason should conquer the imagination; but I have felt the difficulty far to keenly to be surprised at others hesitating to extend the principle of natural selection to so startling a length.

It is scarcely possible to avoid comparing the eye with a telescope. We know that this instrument has been perfected by the long-continued efforts of the highest human intellects; and we naturally infer that the eye has been formed by a somewhat analogous process. But may not this inference be presumptuous? Have we any right to assume that the Creator works by intellectual powers like those of man? If we must compare the eye to an optical instrument, we ought in imagination to take a thick layer of transparent tissue, with spaces filled with fluid, and with a nerve sensitive to light beneath, and then suppose every part of this layer to be continually changing slowly in density, so as to separate into layers of different densities and thicknesses, placed at different distances from each other, and with the surfaces of each layer slowly changing in form. Further we must suppose that there is a power, represented by natural selection or the survival of the fittest, always intently watching each slight alteration in the transparent layers; and carefully preserving each which, under varied circumstances, in any way or degree, tends to produce a distincter image. We must suppose each new state of the instrument to be multiplied by the million; each to be preserved until a better is produced, and then the old ones to be all destroyed. In living bodies, variation will cause the slight alteration, generation will multiply them almost infinitely, and natural selection will pick out with unerring skill each improvement. Let this process go on for millions of years; and during each year on millions of individuals of many kinds; and may we not believe that a living optical instrument might thus be formed as superior to one of glass, as the works of the Creator are to those of man?

MODES Of TRANSITION.

If it could be demonstrated that any complex organ existed, which could not possibly have been formed by numerous, successive, slight modifications, my theory would absolutely break down. But I can find out no such case. No doubt many organs exist of which we do not know the transitional grades, more especially if we look to much-isolated species, around which, according to the theory, there has been much extinction. Or again, if we take an organ common to all the members of a class, for in this latter case the organ must have been originally formed at a remote period, since which all the many members of the class have been developed; and in order to discover the early transitional grades through which the organ has passed, we should have to look to very ancient ancestral forms, long since become extinct.

We should be extremely cautious in concluding that an organ could not have been formed by transitional gradations of some kind. Numerous cases could be given among the lower animals of the same organ performing at the same time wholly distinct functions; thus in the larva of the dragon-fly and in the fish Cobites the alimentary canal respires, digests, and excretes. In the Hydra, the animal may be turned inside out, and the exterior surface will then digest and the stomach respire. In such cases natural selection might specialise, if any advantage were thus gained, the whole or part of an organ, which had previously performed two functions, for one function alone, and thus by insensible steps greatly change its nature. Many plants are known which regularly produce at the same time differently constructed flowers; and if such plants were to produce one kind alone, a great change would be effected with comparative suddenness in the character of the species. It is, however, probable that the two sorts of flowers borne by the same plant were originally differentiated by finely graduated steps, which may still be followed in some few cases.

Again, two distinct organs, or the same organ under two very different forms, may simultaneously perform in the same individual the same function, and this is an extremely important means of transition: to give one instance – there are fish with gills or branchiae that breathe the air dissolved in the water, at the same time that they breathe free air in their swim-bladders, this latter organ being divided by highly vascular partitions and having a ductus pneumaticus for the supply of air. To give another instance from the vegetable kingdom: plants climb by three distinct means, by spirally twining, by clasping a support with their sensitive tendrils, and by the emission of aerial rootlets; these three means are usually found in distinct groups, but some few species exhibit two of the means, or even all three, combined in the same individual. In all such cases one of the two organs might readily be modified and perfected so as to perform all the work, being aided during the progress of modification by the other organ; and then this other organ might be modified for some other and quite distinct purpose, or be wholly obliterated.

The illustration of the swim-bladder in fishes is a good one, because it shows us clearly the highly important fact that an organ originally constructed for one purpose, namely flotation, may be converted into one for a widely different purpose, namely respiration. The swim-bladder has, also, been worked in as an accessory to the auditory organs of certain fishes. All physiologists admit that the swim-bladder is homologous, or "ideally similar" in position and structure with the lungs of the higher vertebrate animals: hence there is no reason to doubt that the swim-bladder has actually been converted into lungs, or an organ used exclusively for respiration.

According to this view it may be inferred that all vertebrate animals with true lungs are descended by ordinary generation from an ancient and unknown prototype which was furnished with a floating apparatus or swim-bladder. We can thus, as I infer from Professor Owen's interesting description of these parts, understand the strange fact that every particle of food and drink which we swallow has to pass over the orifice of the trachea, with some risk of falling into the lungs, notwithstanding the beautiful contrivance by which the glottis is closed. In the higher Vertebrata the branchiae have wholly disappeared – but in the embryo the slits on the sides of the neck and the loop-like course of the arteries still mark their former position. But it is conceivable that the now utterly lost branchiae might have been gradually worked in by natural selection for some distinct purpose: for instance, Landois has shown that the wings of insects are developed from the trachea; it is therefore highly probable that in this great class organs which once served for respiration have been actually converted into organs for flight.

In considering transitions of organs, it is so important to bear in mind the probability of conversion from one function to another, that I will give another instance. Pedunculated cirripedes have two minute folds of skin, called by me the ovigerous frena, which serve, through the means of a sticky secretion, to retain the eggs until they are hatched within the sack. These cirripedes have no branchiae, the whole surface of the body and of the sack, together with the small frena, serving for respiration. The Balanidae or sessile cirripedes, on the other hand, have no ovigerous frena, the eggs lying loose at the bottom of the sack, within the well-enclosed shell; but they have, in the same relative position with the frena, large, much-folded membranes, which freely communicate with the circulatory lacunae of the sack and body, and which have been considered by all naturalists to act as branchiae. Now I think no one will dispute that the ovigerous frena in the one family are strictly homologous with the branchiae of the other family; indeed, they graduate into each other. Therefore it need not be doubted that the two little folds of skin, which originally served as ovigerous frena, but which, likewise, very slightly aided in the act of respiration, have been gradually converted by natural selection into branchiae, simply through an increase in their size and the obliteration of their adhesive glands. If all pedunculated cirripedes had become extinct, and they have suffered far more extinction than have sessile cirripedes, who would ever have imagined that the branchiae in this latter family had originally existed as organs for preventing the ova from being washed out of the sack?

There is another possible mode of transition, namely, through the acceleration or retardation of the period of reproduction. This has lately been insisted on by Professor Cope and others in the United States. It is now known that some animals are capable of reproduction at a very early age, before they have acquired their perfect characters; and if this power became thoroughly well developed in a species, it seems probable that the adult stage of development would sooner or later be lost; and in this case, especially if the larva differed much from the mature form, the character of the species would be greatly changed and degraded. Again, not a few animals, after arriving at maturity, go on changing in character during nearly their whole lives. With mammals, for instance, the form of the skull is often much altered with age, of which Dr. Murie has given some striking instances with seals. Every one knows how the horns of stags become more and more branched, and the plumes of some birds become more finely developed, as they grow older. Professor Cope states that the teeth of certain lizards change much in shape with advancing years. With crustaceans not only many trivial, but some important parts assume a new character, as recorded by Fritz Muller, after maturity. In all such cases – and many could be given – if the age for reproduction were retarded, the character of the species, at least in its adult state, would be modified; nor is it improbable that the previous and earlier stages of development would in some cases be hurried through and finally lost. Whether species have often or ever been modified through this comparatively sudden mode of transition, I can form no opinion; but if this has occurred, it is probable that the differences between the young and the mature, and between the mature and the old, were primordially acquired by graduated steps.

SPECIAL DIFFICULTIES OF THE THEORY OF NATURAL SELECTION.

Although we must be extremely cautious in concluding that any organ could not have been produced by successive, small, transitional gradations, yet undoubtedly serious cases of difficulty occur.

One of the most serious is that of neuter insects, which are often differently constructed from either the males or fertile females; but this case will be treated of in the next chapter. The electric organs of fishes offer another case of special difficulty; for it is impossible to conceive by what steps these wondrous organs have been produced. But this is not surprising, for we do not even know of what use they are. In the gymnotus and torpedo they no doubt serve as powerful means of defence, and perhaps for securing prey; yet in the ray, as observed by Matteucci, an analogous organ in the tail manifests but little electricity, even when the animal is greatly irritated; so little that it can hardly be of any use for the above purposes. Moreover, in the ray, besides the organ just referred to, there is, as Dr. R. McDonnell has shown, another organ near the head, not known to be electrical, but which appears to be the real homologue of the electric battery in the torpedo. It is generally admitted that there exists between these organs and ordinary muscle a close analogy, in intimate structure, in the distribution of the nerves, and in the manner in which they are acted on by various reagents. It should, also, be especially observed that muscular contraction is accompanied by an electrical discharge; and, as Dr. Radcliffe insists, "in the electrical apparatus of the torpedo during rest, there would seem to be a charge in every respect like that which is met with in muscle and nerve during the rest, and the discharge of the torpedo, instead of being peculiar, may be only another form of the discharge which attends upon the action of muscle and motor nerve." Beyond this we cannot at present go in the way of explanation; but as we know so little about the uses of these organs, and as we know nothing about the habits and structure of the progenitors of the existing electric fishes, it would be extremely bold to maintain that no serviceable transitions are possible by which these organs might have been gradually developed.

These organs appear at first to offer another and far more serious difficulty; for they occur in about a dozen kinds of fish, of which several are widely remote in their affinities. When the same organ is found in several members of the same class, especially if in members having very different habits of life, we may generally attribute its presence to inheritance from a common ancestor; and its absence in some of the members to loss through disuse or natural selection. So that, if the electric organs had been inherited from some one ancient progenitor, we might have expected that all electric fishes would have been specially related to each other; but this is far from the case. Nor does geology at all lead to the belief that most fishes formerly possessed electric organs, which their modified descendants have now lost. But when we look at the subject more closely, we find in the several fishes provided with electric organs, that these are situated in different parts of the body, that they differ in construction, as in the arrangement of the plates, and, according to Pacini, in the process or means by which the electricity is excited – and lastly, in being supplied with nerves proceeding from different sources, and this is perhaps the most important of all the differences. Hence in the several fishes furnished with electric organs, these cannot be considered as homologous, but only as analogous in function. Consequently there is no reason to suppose that they have been inherited from a common progenitor; for had this been the case they would have closely resembled each other in all respects. Thus the difficulty of an organ, apparently the same, arising in several remotely allied species, disappears, leaving only the lesser yet still great difficulty: namely, by what graduated steps these organs have been developed in each separate group of fishes.

The luminous organs which occur in a few insects, belonging to widely different families, and which are situated in different parts of the body, offer, under our present state of ignorance, a difficulty almost exactly parallel with that of the electric organs. Other similar cases could be given; for instance in plants, the very curious contrivance of a mass of pollen-grains, borne on a foot-stalk with an adhesive gland, is apparently the same in Orchis and Asclepias, genera almost as remote as is possible among flowering plants; but here again the parts are not homologous. In all cases of beings, far removed from each other in the scale of organisation, which are furnished with similar and peculiar organs, it will be found that although the general appearance and function of the organs may be the same, yet fundamental differences between them can always be detected. For instance, the eyes of Cephalopods or cuttle-fish and of vertebrate animals appear wonderfully alike; and in such widely sundered groups no part of this resemblance can be due to inheritance from a common progenitor. Mr. Mivart has advanced this case as one of special difficulty, but I am unable to see the force of his argument. An organ for vision must be formed of transparent tissue, and must include some sort of lens for throwing an image at the back of a darkened chamber. Beyond this superficial resemblance, there is hardly any real similarity between the eyes of cuttle-fish and vertebrates, as may be seen by consulting Hensen's admirable memoir on these organs in the Cephalopoda. It is impossible for me here to enter on details, but I may specify a few of the points of difference. The crystalline lens in the higher cuttle-fish consists of two parts, placed one behind the other like two lenses, both having a very different structure and disposition to what occurs in the vertebrata. The retina is wholly different, with an actual inversion of the elemental parts, and with a large nervous ganglion included within the membranes of the eye. The relations of the muscles are as different as it is possible to conceive, and so in other points. Hence it is not a little difficult to decide how far even the same terms ought to be employed in describing the eyes of the Cephalopoda and Vertebrata. It is, of course, open to any one to deny that the eye in either case could have been developed through the natural selection of successive slight variations; but if this be admitted in the one case it is clearly possible in the other; and fundamental differences of structure in the visual organs of two groups might have been anticipated, in accordance with this view of their manner of formation. As two men have sometimes independently hit on the same invention, so in the several foregoing cases it appears that natural selection, working for the good of each being, and taking advantage of all favourable variations, has produced similar organs, as far as function is concerned, in distinct organic beings, which owe none of their structure in common to inheritance from a common progenitor.

Fritz Muller, in order to test the conclusions arrived at in this volume, has followed out with much care a nearly similar line of argument. Several families of crustaceans include a few species, possessing an air-breathing apparatus and fitted to live out of the water. In two of these families, which were more especially examined by Muller, and which are nearly related to each other, the species agree most closely in all important characters: namely in their sense organs, circulating systems, in the position of the tufts of hair within their complex stomachs, and lastly in the whole structure of the water-breathing branchiae, even to the microscopical hooks by which they are cleansed. Hence it might have been expected that in the few species belonging to both families which live on the land, the equally important air-breathing apparatus would have been the same; for why should this one apparatus, given for the same purpose, have been made to differ, while all the other important organs were closely similar, or rather, identical.

Fritz Muller argues that this close similarity in so many points of structure must, in accordance with the views advanced by me, be accounted for by inheritance from a common progenitor. But as the vast majority of the species in the above two families, as well as most other crustaceans, are aquatic in their habits, it is improbable in the highest degree that their common progenitor should have been adapted for breathing air. Muller was thus led carefully to examine the apparatus in the air-breathing species; and he found it to differ in each in several important points, as in the position of the orifices, in the manner in which they are opened and closed, and in some accessory details. Now such differences are intelligible, and might even have been expected, on the supposition that species belonging to distinct families had slowly become adapted to live more and more out of water, and to breathe the air. For these species, from belonging to distinct families, would have differed to a certain extent, and in accordance with the principle that the nature of each variation depends on two factors, viz., the nature of the organism and that of the surrounding conditions, their variability assuredly would not have been exactly the same. Consequently natural selection would have had different materials or variations to work on, in order to arrive at the same functional result; and the structures thus acquired would almost necessarily have differed. On the hypothesis of separate acts of creation the whole case remains unintelligible. This line of argument seems to have had great weight in leading Fritz Muller to accept the views maintained by me in this volume.

Another distinguished zoologist, the late Professor Claparede, has argued in the same manner, and has arrived at the same result. He shows that there are parasitic mites (Acaridae), belonging to distinct sub-families and families, which are furnished with hair-claspers. These organs must have been independently developed, as they could not have been inherited from a common progenitor; and in the several groups they are formed by the modification of the fore legs, of the hind legs, of the maxillae or lips, and of appendages on the under side of the hind part of the body.

In the foregoing cases, we see the same end gained and the same function performed, in beings not at all or only remotely allied, by organs in appearance, though not in development, closely similar. On the other hand, it is a common rule throughout nature that the same end should be gained, even sometimes in the case of closely related beings, by the most diversified means. How differently constructed is the feathered wing of a bird and the membrane-covered wing of a bat; and still more so the four wings of a butterfly, the two wings of a fly, and the two wings with the elytra of a beetle. Bivalve shells are made to open and shut, but on what a number of patterns is the hinge constructed, from the long row of neatly interlocking teeth in a Nucula to the simple ligament of a Mussel! Seeds are disseminated by their minuteness, by their capsule being converted into a light balloon-like envelope, by being embedded in pulp or flesh, formed of the most diverse parts, and rendered nutritious, as well as conspicuously coloured, so as to attract and be devoured by birds, by having hooks and grapnels of many kinds and serrated awns, so as to adhere to the fur of quadrupeds, and by being furnished with wings and plumes, as different in shape as they are elegant in structure, so as to be wafted by every breeze. I will give one other instance: for this subject of the same end being gained by the most diversified means well deserves attention. Some authors maintain that organic beings have been formed in many ways for the sake of mere variety, almost like toys in a shop, but such a view of nature is incredible. With plants having separated sexes, and with those in which, though hermaphrodites, the pollen does not spontaneously fall on the stigma, some aid is necessary for their fertilisation. With several kinds this is effected by the pollen-grains, which are light and incoherent, being blown by the wind through mere chance on to the stigma; and this is the simplest plan which can well be conceived. An almost equally simple, though very different plan occurs in many plants in which a symmetrical flower secretes a few drops of nectar, and is consequently visited by insects; and these carry the pollen from the anthers to the stigma.

From this simple stage we may pass through an inexhaustible number of contrivances, all for the same purpose and effected in essentially the same manner, but entailing changes in every part of the flower. The nectar may be stored in variously shaped receptacles, with the stamens and pistils modified in many ways, sometimes forming trap-like contrivances, and sometimes capable of neatly adapted movements through irritability or elasticity. From such structures we may advance till we come to such a case of extraordinary adaptation as that lately described by Dr. Cruger in the Coryanthes. This orchid has part of its labellum or lower lip hollowed out into a great bucket, into which drops of almost pure water continually fall from two secreting horns which stand above it; and when the bucket is half-full, the water overflows by a spout on one side. The basal part of the labellum stands over the bucket, and is itself hollowed out into a sort of chamber with two lateral entrances; within this chamber there are curious fleshy ridges. The most ingenious man, if he had not witnessed what takes place, could never have imagined what purpose all these parts serve. But Dr. Cruger saw crowds of large humble-bees visiting the gigantic flowers of this orchid, not in order to suck nectar, but to gnaw off the ridges within the chamber above the bucket; in doing this they frequently pushed each other into the bucket, and their wings being thus wetted they could not fly away, but were compelled to crawl out through the passage formed by the spout or overflow. Dr. Cruger saw a "continual procession" of bees thus crawling out of their involuntary bath. The passage is narrow, and is roofed over by the column, so that a bee, in forcing its way out, first rubs its back against the viscid stigma and then against the viscid glands of the pollen-masses. The pollen-masses are thus glued to the back of the bee which first happens to crawl out through the passage of a lately expanded flower, and are thus carried away. Dr. Cruger sent me a flower in spirits of wine, with a bee which he had killed before it had quite crawled out, with a pollen-mass still fastened to its back. When the bee, thus provided, flies to another flower, or to the same flower a second time, and is pushed by its comrades into the bucket and then crawls out by the passage, the pollen-mass necessarily comes first into contact with the viscid stigma, and adheres to it, and the flower is fertilised. Now at last we see the full use of every part of the flower, of the water-secreting horns of the bucket half-full of water, which prevents the bees from flying away, and forces them to crawl out through the spout, and rub against the properly placed viscid pollen-masses and the viscid stigma.