The Variation of Animals and Plants under Domestication — Volume 2

In several distinct breeds of the pigeon and fowl, the legs and the two outer toes are heavily feathered, so that in the trumpeter pigeon they appear like little wings. In the feather-legged bantam the "boots" or feathers, which grow from the outside of the leg and generally from the two outer toes, have, according to the excellent authority of Mr. Hewitt (25/6. 'The Poultry Book' by W.B. Tegetmeier 1866 page 250.), been seen to exceed the wing-feathers in length, and in one case were actually nine and a half inches long! As Mr. Blyth has remarked to me, these leg-feathers resemble the primary wing- feathers, and are totally unlike the fine down which naturally grows on the legs of some birds, such as grouse and owls. Hence it may be suspected that excess of food has first given redundancy to the plumage, and then that the law of homologous variation has led to the development of feathers on the legs, in a position corresponding with those on the wing, namely, on the outside of the tarsi and toes. I am strengthened in this belief by the following curious case of correlation, which for a long time seemed to me utterly inexplicable, namely, that in pigeons of any breed, if the legs are feathered, the two outer toes are partially connected by skin. These two outer toes correspond with our third and fourth toes. (25/7. Naturalists differ with respect to the homologies of the digits of birds; but several uphold the view above advanced. See on this subject Dr. E.S. Morse in 'Annals of the Lyceum of Nat. Hist. of New York' volume 10 1872 page 16.) Now, in the wing of the pigeon or of any other bird, the first and fifth digits are aborted; the second is rudimentary and carries the so-called "bastard-wing;" whilst the third and fourth digits are completely united and enclosed by skin, together forming the extremity of the wing. So that in feather-footed pigeons, not only does the exterior surface support a row of long feathers, like wing-feathers, but the very same digits which in the wing are completely united by skin become partially united by skin in the feet; and thus by the law of the correlated variation of homologous parts we can understand the curious connection of feathered legs and membrane between the two outer toes.

Andrew Knight (24/8. A. Walker on 'Intermarriage' 1838 page 160.) has remarked that the face or head and the limbs usually vary together in general proportions. Compare, for instance, the limbs of a dray and race horse, or of a greyhound and mastiff. What a monster a greyhound would appear with the head of a mastiff! The modern bulldog, however, has fine limbs, but this is a recently-selected character. From the measurements given in the sixth chapter, we see that in several breeds of the pigeon the length of the beak and the size of the feet are correlated. The view which, as before explained, seems the most probable is, that disuse in all cases tends to diminish the feet, the beak becoming at the same time shorter through correlation; but that in some few breeds in which length of beak has been a selected point, the feet, notwithstanding disuse, have increased in size through correlation. In the following case some kind of correlation is seen to exist between the feet and beak: several specimens have been sent to Mr. Bartlett at different times, as hybrids between ducks and fowls, and I have seen one; these were, as might be expected, ordinary ducks in a semi-monstrous condition, and in all of them the swimming-web between the toes was quite deficient or much reduced, and in all the beak was narrow and ill-shaped.

With the increased length of the beak in pigeons, not only the tongue increases in length, but likewise the orifice of the nostrils. But the increased length of the orifice of the nostrils perhaps stands in closer correlation with the development of the corrugated skin or wattle at the base of the beak, for when there is much wattle round the eyes, the eyelids are greatly increased or even doubled in length.

There is apparently some correlation even in colour between the head and the extremities. Thus with horses a large white star or blaze on the forehead is generally accompanied by white feet. (25/9. 'The Farrier and Naturalist' volume 1 1828 page 456. A gentleman who has attended to this point, tells me that about three-fourths of white-faced horses have white legs.) With white rabbits and cattle, dark marks often co-exist on the tips of the ears and on the feet. In black and tan dogs of different breeds, tan-coloured spots over the eyes and tan-coloured feet almost invariably go together. These latter cases of connected colouring may be due either to reversion or to analogous variation, — subjects to which I shall hereafter return, — but this does not necessarily determine the question of their original correlation. Mr. H.W. Jackson informs me that he has observed many hundred white-footed cats, and he finds that all are more or less conspicuously marked with white on the front of the neck or chest.

The lopping forwards and downwards of the immense ears of fancy rabbits seems partly due to the disuse of the muscles, and partly to the weight and length of the ears, which have been increased by selection during many generations. Now, with the increased size and changed direction of the ears not only has the bony auditory meatus become changed in outline, direction, and greatly in size, but the whole skull has been slightly modified. This could be clearly seen in "half-lops" — that is, in rabbits with only one ear lopping forward — for the opposite sides of their skulls were not strictly symmetrical. This seems to me a curious instance of correlation, between hard bones and organs so soft and flexible, as well as so unimportant under a physiological point of view, as the external ears. The result no doubt is largely due to mere mechanical action, that is, to the weight of the ears, on the same principle that the skull of a human infant is easily modified by pressure.

The skin and the appendages of hair, feathers, hoofs, horns, and teeth, are homologous over the whole body. Every one knows that the colour of the skin and that of the hair usually vary together; so that Virgil advises the shepherd to look whether the mouth and tongue of the ram are black, lest the lambs should not be purely white. The colour of the skin and hair, and the odour emitted by the glands of the skin, are said (25/10. Godron 'Sur l'Espece' tome 2 page 217.) to be connected, even in the same race of men. Generally the hair varies in the same way all over the body in length, fineness, and curliness. The same rule holds good with feathers, as we see with the laced and frizzled breeds both of fowls and pigeons. In the common cock the feathers on the neck and loins are always of a particular shape, called hackles: now in the Polish breed, both sexes are characterised by a tuft of feathers on the head, and through correlation these feathers in the male always assume the form of hackles. The wing and tail-feathers, though arising from parts not homologous, vary in length together; so that long or short winged pigeons generally have long or short tails. The case of the Jacobin-pigeon is more curious, for the wing and tail feathers are remarkably long; and this apparently has arisen in correlation with the elongated and reversed feathers on the back of the neck, which form the hood.

The hoofs and hair are homologous appendages; and a careful observer, namely Azara (25/11. 'Quadrupedes du Paraguay' tome 2 page 333.), states that in Paraguay horses of various colours are often born with their hair curled and twisted like that on the head of a negro. This peculiarity is strongly inherited. But what is remarkable is that the hoofs of these horses "are absolutely like those of a mule." The hair also of their manes and tails is invariably much shorter than usual, being only from four to twelve inches in length; so that curliness and shortness of the hair are here, as with the negro, apparently correlated.

With respect to the horns of sheep, Youatt (25/12. 'On Sheep' page 142.) remarks that "multiplicity of horns is not found in any breed of much value; it is generally accompanied by great length and coarseness of the fleece." Several tropical breeds of sheep which are clothed with hair instead of wool, have horns almost like those of a goat. Sturm (25/13. 'Ueber Racen, Kreuzungen' etc. 1825 s. 24.) expressly declares that in different races the more the wool is curled the more the horns are spirally twisted. We have seen in the third chapter, where other analogous facts have been given, that the parent of the Mauchamp breed, so famous for its fleece, had peculiarly shaped horns. The inhabitants of Angora assert (25/14. Quoted from Conolly in 'The Indian Field' February 1859 volume 2 page 266.) that "only the white goats which have horns wear the fleece in the long curly locks that are so much admired; those which are not horned having a comparatively close coat." From these cases we may infer that the hair or wool and the horns tend to vary in a correlated manner. (25/15. In the third chapter I have said that "the hair and horns are so closely related to each other, that they are apt to vary together." Dr. Wilckens ("Darwin's Theorie" 'Jahrbuch der Deutschen Viehzucht' 1866 1. Heft) translates my words into "lang- und grobhaarige Thiere sollen geneigter sein, lange und viele Horner zu bekommen" and he then justly disputes this proposition; but what I have really said, in accordance with the authorities just quoted, may, I think, be trusted.) Those who have tried hydropathy are aware that the frequent application of cold water stimulates the skin; and whatever stimulates the skin tends to increase the growth of the hair, as is well shown in the abnormal growth of hair near old inflamed surfaces. Now, Professor Low (25/16. 'Domesticated Animals of the British Islands' pages 307, 368. Dr. Wilckens argues ('Landwirth. Wochenblatt' Nr. 10 1869) to the same effect with respect to domestic animals in Germany.) is convinced that with the different races of British cattle thick skin and long hair depend on the humidity of the climate which they inhabit. We can thus see how a humid climate might act on the horns — in the first place directly on the skin and hair, and secondly by correlation on the horns. The presence or absence of horns, moreover, both in the case of sheep and cattle, acts, as will presently be shown, by some sort of correlation on the skull.

With respect to hair and teeth, Mr. Yarrell (25/17. 'Proceedings Zoolog. Soc.' 1833 page 113.) found many of the teeth deficient in three hairless "Egyptian dogs," and in a hairless terrier. The incisors, canines, and the premolars suffered most, but in one case all the teeth, except the large tubercular molar on each side, were deficient. With man several striking cases have been recorded (25/18. Sedgwick 'Brit. and Foreign Medico-Chirurg. Review' April 1863 page 453.) of inherited baldness with inherited deficiency, either complete or partial, of the teeth. I may give an analogous case, communicated to me by Mr. W. Wedderburn, of a Hindoo family in Scinde, in which ten men, in the course of four generations, were furnished, in both jaws taken together, with only four small and weak incisor teeth and with eight posterior molars. The men thus affected have very little hair on the body, and become bald early in life. They also suffer much during hot weather from excessive dryness of the skin. It is remarkable that no instance has occurred of a daughter being thus affected; and this fact reminds us how much more liable men are in England to become bald than women. Though the daughters in the above family are never affected, they transmit the tendency to their sons; and no case has occurred of a son transmitting it to his sons. The affection thus appears only in alternate generations, or after longer intervals. There is a similar connection between hair and teeth, according to Mr. Sedgwick, in those rare cases in which the hair has been renewed in old age, for this has "usually been accompanied by a renewal of the teeth." I have remarked in a former part of this volume that the great reduction in the size of the tusks in domestic boars probably stands in close relation with their diminished bristles, due to a certain amount of protection; and that the reappearance of the tusks in boars, which have become feral and are fully exposed to the weather, probably depends on the reappearance of the bristles. I may add, though not strictly connected with our present point, that an agriculturist (25/19. 'Gardener's Chronicle' 1849 page 205.) asserts that "pigs with little hair on their bodies are most liable to lose their tails, showing a weakness of the tegumental structure. It may be prevented by crossing with a more hairy breed."

In the previous cases deficient hair, and teeth deficient in number or size, are apparently connected. In the following cases abnormally redundant hair, and teeth either deficient or redundant, are likewise connected. Mr. Crawfurd (25/20. 'Embassy to the Court of Ava' volume 1 page 320.) saw at the Burmese Court a man, thirty years old, with his whole body, except the hands and feet, covered with straight silky hair, which on the shoulders and spine was five inches in length. At birth the ears alone were covered. He did not arrive at puberty, or shed his milk teeth, until twenty years old; and at this period he acquired five teeth in the upper jaw, namely, four incisors and one canine, and four incisor teeth in the lower jaw; all the teeth were small. This man had a daughter who was born with hair within her ears; and the hair soon extended over her body. When Captain Yule (25/21. 'Narrative of a Mission to the Court of Ava in 1855' page 94.) visited the Court, he found this girl grown up; and she presented a strange appearance with even her nose densely covered with soft hair. Like her father, she was furnished with incisor teeth alone. The King had with difficulty bribed a man to marry her, and of her two children, one, a boy fourteen months old, had hair growing out of his ears, with a beard and moustache. This strange peculiarity has, therefore, been inherited for three generations, with the molar teeth deficient in the grandfather and mother; whether these teeth would likewise fail in the infant could not then be told.

A parallel case of a man fifty-five years old, and of his son, with their faces covered with hair, has recently occurred in Russia. Dr. Alex. Brandt has sent me an account of this case, together with specimens of the extremely fine hair from the cheeks. The man is deficient in teeth, possessing only four incisors in the lower and two in the upper jaw. His son, about three years old, has no teeth except four lower incisors. The case, as Dr. Brandt remarks in his letter, no doubt is due to an arrest of development in the hair and teeth. We here see how independent of the ordinary conditions of existence such arrests must be, for the lives of a Russian peasant and of a native of Burmah are as different as possible. (25/22. I owe to the kindness of M. Chauman, of St. Petersburg, excellent photographs of this man and his son, both of whom have since been exhibited in Paris and London.)

Here is another and somewhat different case communicated to me by Mr. Wallace on the authority of Dr. Purland, a dentist: Julia Pastrana, a Spanish dancer, was a remarkably fine woman, but she had a thick masculine beard and a hairy forehead; she was photographed, and her stuffed skin was exhibited as a show; but what concerns us is, that she had in both the upper and lower jaw an irregular double set of teeth, one row being placed within the other, of which Dr. Purland took a cast. From the redundancy of teeth her mouth projected, and her face had a gorilla-like appearance. These cases and those of the hairless dogs forcibly call to mind the fact, that the two orders of mammals — namely, the Edentata and Cetacea — which are the most abnormal in their dermal covering, are likewise the most abnormal either by deficiency or redundancy of teeth.

The organs of sight and hearing are generally admitted to be homologous with one another and with various dermal appendages; hence these parts are liable to be abnormally affected in conjunction. Mr. White Cowper says "that in all cases of double microphthalmia brought under his notice he has at the same time met with defective development of the dental system." Certain forms of blindness seem to be associated with the colour of the hair; a man with black hair and a woman with light-coloured hair, both of sound constitution, married and had nine children, all of whom were born blind; of these children, five "with dark hair and brown iris were afflicted with amaurosis; the four others, with light-coloured hair and blue iris, had amaurosis and cataract conjoined." Several cases could be given, showing that some relation exists between various affections of the eyes and ears; thus Liebreich states that out of 241 deaf-mutes in Berlin, no less than fourteen suffered from the rare disease called pigmentary retinitis. Mr. White Cowper and Dr. Earle have remarked that inability to distinguish different colours, or colour-blindness, "is often associated with a corresponding inability to distinguish musical sounds." (25/23. These statements are taken from Mr. Sedgwick in the 'Medico-Chirurg. Review' July 1861 page 198; April 1863 pages 455 and 458. Liebreich is quoted by Professor Devay in his 'Mariages Consanguins' 1862 page 116.)

Here is a more curious case: white cats, if they have blue eyes, are almost always deaf. I formerly thought that the rule was invariable, but I have heard of a few authentic exceptions. The first two notices were published in 1829 and relate to English and Persian cats: of the latter, the Rev. W.T. Bree possessed a female, and he states, "that of the offspring produced at one and the same birth, such as, like the mother, were entirely white (with blue eyes) were, like her, invariably deaf; while those that had the least speck of colour on their fur, as invariably possessed the usual faculty of hearing." (25/24. Loudon's 'Mag. of Nat. Hist.' volume 1 1829 pages 66, 178. See also Dr. P. Lucas 'L'Hered. Nat.' tome 1 page 428 on the inheritance of deafness in cats. Mr. Lawson Tait states ('Nature' 1873 page 323) that only male cats are thus affected; but this must be a hasty generalisation. The first case recorded in England by Mr. Bree related to a female, and Mr. Fox informs me that he has bred kittens from a white female with blue eyes, which was completely deaf; he has also observed other females in the same condition.) The Rev. W. Darwin Fox informs me that he has seen more than a dozen instances of this correlation in English, Persian, and Danish cats; but he adds "that, if one eye, as I have several times observed, be not blue, the cat hears. On the other hand, I have never seen a white cat with eyes of the common colour that was deaf." In France Dr. Sichel (25/25. 'Annales des Sc. Nat.' Zoolog. 3rd series 1847 tome 8 page 239.) has observed during twenty years similar facts; he adds the remarkable case of the iris beginning, at the end of four months, to grow dark-coloured, and then the cat first began to hear.

This case of correlation in cats has struck many persons as marvellous. There is nothing unusual in the relation between blue eyes and white fur; and we have already seen that the organs of sight and hearing are often simultaneously affected. In the present instance the cause probably lies in a slight arrest of development in the nervous system in connection with the sense-organs. Kittens during the first nine days, whilst their eyes are closed, appear to be completely deaf; I have made a great clanging noise with a poker and shovel close to their heads, both when they were asleep and awake, without producing any effect. The trial must not be made by shouting close to their ears, for they are, even when asleep, extremely sensitive to a breath of air. Now, as long as the eyes continue closed, the iris is no doubt blue, for in all the kittens which I have seen this colour remains for some time after the eyelids open. Hence, if we suppose the development of the organs of sight and hearing to be arrested at the stage of the closed eyelids, the eyes would remain permanently blue and the ears would be incapable of perceiving sound; and we should thus understand this curious case. As, however, the colour of the fur is determined long before birth, and as the blueness of the eyes and the whiteness of the fur are obviously connected, we must believe that some primary cause acts at a much earlier period.

The instances of correlated variability hitherto given have been chiefly drawn from the animal kingdom, and we will now turn to plants. Leaves, sepals, petals, stamens, and pistils are all homologous. In double flowers we see that the stamens and pistils vary in the same manner, and assume the form and colour of the petals. In the double columbine (Aquilegia vulgaris), the successive whorls of stamens are converted into cornucopias, which are enclosed within one another and resemble the true petals. In hose-in-hose flowers the sepals mock the petals. In some cases the flowers and leaves vary together in tint: in all the varieties of the common pea, which have purple flowers, a purple mark may be seen on the stipules.

M. Faivre states that with the varieties of Primula sinensis the colour of the flower is evidently correlated with the colour of the under side of the leaves; and he adds that the varieties with fimbriated flowers almost always have voluminous, balloon-like calyces. (25/26. 'Revue des Cours Scientifiques' June 5, 1869 page 430.) With other plants the leaves and fruit or seeds vary together in colour, as in a curious pale-leaved variety of the sycamore, which has recently been described in France (25/27. 'Gardener's Chronicle' 1864 page 1202.), and as in the purple-leaved hazel, in which the leaves, the husk of the nut, and the pellicle round the kernel are all coloured purple. (25/28. Verlot gives several other instances 'Des Varietes' 1865 page 72.) Pomologists can predict to a certain extent, from the size and appearance of the leaves of their seedlings, the probable nature of the fruit; for, as Van Mons remarks (25/29. 'Arbres Fruitiers' 1836 tome 2 pages 204, 226.) variations in the leaves are generally accompanied by some modification in the flower, and consequently in the fruit. In the Serpent melon, which has a narrow tortuous fruit above a yard in length, the stem of the plant, the peduncle of the female flower, and the middle lobe of the leaf, are all elongated in a remarkable manner. On the other hand, several varieties of Cucurbita, which have dwarfed stems, all produce, as Naudin remarks, leaves of the same peculiar shape. Mr. G. Maw informs me that all the varieties of the scarlet Pelargoniums which have contracted or imperfect leaves have contracted flowers: the difference between "Brilliant" and its parent "Tom Thumb" is a good instance of this. It may be suspected that the curious case described by Risso (25/30. 'Annales du Museum' tome 20 page 188.), of a variety of the Orange which produces on the young shoots rounded leaves with winged petioles, and afterwards elongated leaves on long but wingless petioles, is connected with the remarkable change in form and nature which the fruit undergoes during its development.

In the following instance we have the colour and the form of the petals apparently correlated, and both dependent on the nature of the season. An observer, skilled in the subject, writes (25/31. 'Gardener's Chronicle' 1843 page 877.), "I noticed, during the year 1842, that every Dahlia of which the colour had any tendency to scarlet, was deeply notched — indeed, to so great an extent as to give the petals the appearance of a saw; the indentures were, in some instances, more than a quarter of an inch deep." Again, Dahlias which have their petals tipped with a different colour from the rest of the flower are very inconstant, and during certain years some, or even all the flowers, become uniformly coloured; and it has been observed with several varieties (25/32. Ibid 1845 page 102.) that when this happens the petals grow much elongated and lose their proper shape. This, however, may be due to reversion, both in colour and form, to the aboriginal species.

In this discussion on correlation, we have hitherto treated of cases in which we can partly understand the bond of connection; but I will now give cases in which we cannot even conjecture, or can only very obscurely see, the nature of the bond. Isidore Geoffroy Saint-Hilaire, in his work on Monstrosities, insists (25/33. 'Hist. des Anomalies' tome 3 page 402. See also M. Camille Dareste 'Recherches sur les Conditions' etc. 1863 pages 16, 48.), "que certaines anomalies coexistent rarement entr'elles, d'autres frequemment, d'autres enfin presque constamment, malgre la difference tres-grande de leur nature, et quoiqu'elles puissent paraitre COMPLETEMENT INDEPENDANTES les unes des autres." We see something analogous in certain diseases: thus in a rare affection of the renal capsules (of which the functions are unknown), the skin becomes bronzed; and in hereditary syphilis, as I hear from Sir J. Paget, both the milk and the second teeth assume a peculiar and characteristic form. Professor Rolleston, also, informs me that the incisor teeth are sometimes furnished with a vascular rim in correlation with intra-pulmonary deposition of tubercles. In other cases of phthisis and of cyanosis the nails and finger- ends become clubbed like acorns. I believe that no explanation has been offered of these and of many other cases of correlated disease.

What can be more curious and less intelligible than the fact previously given, on the authority of Mr. Tegetmeier, that young pigeons of all breeds, which when mature have white, yellow, silver-blue, or dun-coloured plumage, come out of the egg almost naked; whereas pigeons of other colours when first born are clothed with plenty of down? White Pea-fowls, as has been observed both in England and France (25/34. Rev. E.S. Dixon 'Ornamental Poultry' 1848 page 111; Isidore Geoffroy 'Hist. Anomalies' tome 1 page 211.), and as I have myself seen, are inferior in size to the common coloured kind; and this cannot be accounted for by the belief that albinism is always accompanied by constitutional weakness; for white or albino moles are generally larger than the common kind.

To turn to more important characters: the niata cattle of the Pampas are remarkable from their short foreheads, upturned muzzles, and curved lower jaws. In the skull the nasal and premaxillary bones are much shortened, the maxillaries are excluded from any junction with the nasals, and all the bones are slightly modified, even to the plane of the occiput. From the analogous case of the dog, hereafter to be given, it is probable that the shortening of the nasal and adjoining bones is the proximate cause of the other modifications in the skull, including the upward curvature of the lower jaw, though we cannot follow out the steps by which these changes have been effected.

Polish fowls have a large tuft of feathers on their heads; and their skulls are perforated by numerous holes, so that a pin can be driven into the brain without touching any bone. That this deficiency of bone is in some way connected with the tuft of feathers is clear from tufted ducks and geese likewise having perforated skulls. The case would probably be considered by some authors as one of balancement or compensation. In the chapter on Fowls, I have shown that with Polish fowls the tuft of feathers was probably at first small; by continued selection it became larger, and then rested on a fibrous mass; and finally, as it became still larger, the skull itself became more and more protuberant until it acquired its present extraordinary structure. Through correlation with the protuberance of the skull, the shape and even the relative connection of the premaxillary and nasal bones, the shape of the orifice of the nostrils, the breadth of the frontal bone, the shape of the post-lateral processes of the frontal and squamosal bones, and the direction of the bony cavity of the ear, have all been modified. The internal configuration of the skull and the whole shape of the brain have likewise been altered in a truly marvellous manner.

After this case of the Polish fowl it would be superfluous to do more than refer to the details previously given on the manner in which the changed form of the comb has affected the skull, in various breeds of the fowl, causing by correlation crests, protuberances, and depressions on its surface.

With our cattle and sheep the horns stand in close connection with the size of the skull, and with the shape of the frontal bones; thus Cline (25/35. 'On the Breeding of Domestic Animals' 1829 page 6.) found that the skull of a horned ram weighed five times as much as that of a hornless ram of the same age. When cattle become hornless, the frontal bones are "materially diminished in breadth towards the poll;" and the cavities between the bony plates "are not so deep, nor do they extend beyond the frontals." (25/36. Youatt on 'Cattle' 1834 page 283.)

It may be well here to pause and observe how the effects of correlated variability, of the increased use of parts, and of the accumulation of so- called spontaneous variations through natural selection, are in many cases inextricably commingled. We may borrow an illustration from Mr. Herbert Spencer, who remarks that, when the Irish elk acquired its gigantic horns, weighing above one hundred pounds, numerous co-ordinated changes of structure would have been indispensable, — namely, a thickened skull to carry the horns; strengthened cervical vertebrae, with strengthened ligaments; enlarged dorsal vertebrae to support the neck, with powerful fore-legs and feet; all these parts being supplied with proper muscles, blood-vessels, and nerves. How then could these admirably co-ordinated modifications of structure have been acquired? According to the doctrine which I maintain, the horns of the male elk were slowly gained through sexual selection, — that is, by the best-armed males conquering the worse-armed, and leaving a greater number of descendants. But it is not at all necessary that the several parts of the body should have simultaneously varied. Each stag presents individual characteristics, and in the same district those which had slightly heavier horns, or stronger necks, or stronger bodies, or were the most courageous, would secure the greater number of does, and consequently have a greater number of offspring. The offspring would inherit, in a greater or less degree, these same qualities, would occasionally intercross with one another, or with other individuals varying in some favourable manner; and of their offspring, those which were the best endowed in any respect would continue multiplying; and so onwards, always progressing, sometimes in one direction, and sometimes in another, towards the excellently co-ordinated structure of the male elk. To make this clear, let us reflect on the probable steps, as shown in the twentieth chapter, by which our race and dray horses have arrived at their present state of excellence; if we could view the whole series of intermediate forms between one of these animals and an early unimproved progenitor, we should behold a vast number of animals, not equally improved in each generation throughout their entire structure, but sometimes a little more in one point, and sometimes in another, yet on the whole gradually approaching in character to our present race or dray horses, which are so admirably fitted in the one case for fleetness and in the other for draught.

Although natural selection would thus (25/37. Mr. Herbert Spencer 'Principles of Biology' 1864 volume 1 pages 452, 468 takes a different view; and in one place remarks: "We have seen reason to think that, as fast as essential faculties multiply, and as fast as the number of organs that co-operate in any given function increases, indirect equilibration through natural selection becomes less and less capable of producing specific adaptations; and remains fully capable only of maintaining the general fitness of constitution to conditions." This view that natural selection can do little in modifying the higher animals surprises me, seeing that man's selection has undoubtedly effected much with our domesticated quadrupeds and birds.) tend to give to the male elk its present structure, yet it is probable that the inherited effects of use, and of the mutual action of part on part, have been equally or more important. As the horns gradually increased in weight the muscles of the neck, with the bones to which they are attached, would increase in size and strength; and these parts would react on the body and legs. Nor must we overlook the fact that certain parts of the skull and the extremities would, judging by analogy, tend from the first to vary in a correlated manner. The increased weight of the horns would also act directly on the skull, in the same manner as when one bone is removed in the leg of a dog, the other bone, which has to carry the whole weight of the body, increases in thickness. But from the fact given with respect to horned and hornless cattle, it is probable that the horns and skull would immediately act on each other through the principle of correlation. Lastly, the growth and subsequent wear and tear of the augmented muscles and bones would require an increased supply of blood, and consequently increased supply of food; and this again would require increased powers of mastication, digestion, respiration, and excretion.

COLOUR AS CORRELATED WITH CONSTITUTIONAL PECULIARITIES.

It is an old belief that with man there is a connection between complexions and constitution; and I find that some of the best authorities believe in this to the present day. (25/38. Dr. Prosper Lucas apparently disbelieves in any such connection; 'L'Hered. Nat.' tome 2 pages 88-94.) Thus Dr. Beddoe by his tables shows (25/39. 'British Medical Journal' 1862 page 433.) that a relation exists between liability to consumption and the colour of the hair, eyes, and skin. It has been affirmed (25/40. Boudin 'Geograph. Medicale' tome 1 page 406.) that, in the French army which invaded Russia, soldiers having a dark complexion from the southern parts of Europe, withstood the intense cold better than those with lighter complexions from the north; but no doubt such statements are liable to error.

In the second chapter on Selection I have given several cases proving that with animals and plants differences in colour are correlated with constitutional differences, as shown by greater or less immunity from certain diseases, from the attacks of parasitic plants and animals, from scorching by the sun, and from the action of certain poisons. When all the individuals of any one variety possess an immunity of this nature, we do not know that it stands in any sort of correlation with their colour; but when several similarly coloured varieties of the same species are thus characterised, whilst other coloured varieties are not thus favoured, we must believe in the existence of a correlation of this kind. Thus, in the United States purple- fruited plums of many kinds are far more affected by a certain disease than green or yellow-fruited varieties. On the other hand, yellow-fleshed peaches of various kinds suffer from another disease much more than the white-fleshed varieties. In the Mauritius red sugar-canes are much less affected by a particular disease than the white canes. White onions and verbenas are the most liable to mildew; and in Spain the green-fruited grapes suffered from the vine-disease more than other coloured varieties. Dark-coloured pelargoniums and verbenas are more scorched by the sun than varieties of other colours. Red wheats are believed to be hardier than white; and red-flowered hyacinths were more injured during one particular winter in Holland than other coloured varieties. With animals, white terriers suffer most from the distemper, white chickens from a parasitic worm in their tracheae, white pigs from scorching by the sun, and white cattle from flies; but the caterpillars of the silk-moth which yield white cocoons suffered in France less from the deadly parasitic fungus than those producing yellow silk.

The cases of immunity from the action of certain vegetable poisons, in connexion with colour, are more interesting, and are at present wholly inexplicable. I have already given a remarkable instance, on the authority of Professor Wyman, of all the hogs, excepting those of a black colour, suffering severely in Virginia from eating the root of the Lachnanthes tinctoria. According to Spinola and others (25/41. This fact and the following cases, when not stated to the contrary, are taken from a very curious paper by Prof. Heusinger in 'Wochenschrift fur Heilkunde' May 1846 s. 277. Settegast 'Die Thierzucht' 1868 page 39 says that white or white-spotted sheep suffer like pigs, or even die from eating buckwheat; whilst black or dark-woolled individuals are not in the least affected.), buckwheat (Po1ygonum fagopyrum), when in flower, is highly injurious to white or white-spotted pigs, if they are exposed to the heat of the sun, but is quite innocuous to black pigs. According to two accounts, the Hypericum crispum in Sicily is poisonous to white sheep alone; their heads swell, their wool falls off, and they often die; but this plant, according to Lecce, is poisonous only when it grows in swamps; nor is this improbable, as we know how readily the poisonous principle in plants is influenced by the conditions under which they grow.

Three accounts have been published in Eastern Prussia, of white and white- spotted horses being greatly injured by eating mildewed and honeydewed vetches; every spot of skin bearing white hairs becoming inflamed and gangrenous. The Rev. J. Rodwell informs me that his father turned out about fifteen cart-horses into a field of tares which in parts swarmed with black aphides, and which no doubt were honeydewed, and probably mildewed; the horses, with two exceptions, were chestnuts and bays with white marks on their faces and pasterns, and the white parts alone swelled and became angry scabs. The two bay horses with no white marks entirely escaped all injury. In Guernsey, when horses eat fool's parsley (Aethusa cynapium) they are sometimes violently purged; and this plant "has a peculiar effect on the nose and lips, causing deep cracks and ulcers, particularly on horses with white muzzles." (25/42. Mr. Mogford in the 'Veterinarian' quoted in 'The Field' January 22, 1861 page 545.) With cattle, independently of the action of any poison, cases have been published by Youatt and Erdt of cutaneous diseases with much constitutional disturbance (in one instance after exposure to a hot sun) affecting every single point which bore a white hair, but completely passing over other parts of the body. Similar cases have been observed with horses. (25/43. 'Edinburgh Veterinary Journal' October 1860 page 347.)

We thus see that not only do those parts of the skin which bear white hair differ in a remarkable manner from those bearing hair of any other colour, but that some great constitutional difference must be correlated with the colour of the hair; for in the above-mentioned cases, vegetable poisons caused fever, swelling of the head, as well as other symptoms, and even death, to all the white, or white-spotted animals.

CHAPTER 2.XXVI

LAWS OF VARIATION, continued. — SUMMARY.

THE FUSION OF HOMOLOGOUS PARTS. THE VARIABILITY OF MULTIPLE AND HOMOLOGOUS PARTS. COMPENSATION OF GROWTH. MECHANICAL PRESSURE. RELATIVE POSITION OF FLOWERS WITH RESPECT TO THE AXIS, AND OF SEEDS IN THE OVARY, AS INDUCING VARIATION. ANALOGOUS OR PARALLEL VARIETIES. SUMMARY OF THE THREE LAST CHAPTERS.

THE FUSION OF HOMOLOGOUS PARTS.

Geoffroy Saint-Hilaire formerly propounded what he called la loi de l'affinite de soi pour soi, which has been discussed and illustrated by his son, Isidore, with respect to monsters in the animal kingdom (26/1. 'Hist. des Anomalies' 1832 tome 1 pages 22, 537-556; tome 3 page 462.), and by Moquin-Tandon, with respect to monstrous plants.

This law seems to imply that homologous parts actually attract one another and then unite. No doubt there are many wonderful cases, in which such parts become intimately fused together. This is perhaps best seen in monsters with two heads, which are united, summit to summit, or face to face, or Janus-like, back to back, or obliquely side to side. In one instance of two heads united almost face to face, but a little obliquely, four ears were developed, and on one side a perfect face, which was manifestly formed by the fusion of two half-faces. Whenever two bodies or two heads are united, each bone, muscle, vessel, and nerve on the line of junction appears as if it had sought out its fellow, and had become completely fused with it. Lereboullet (26/2. 'Comptes Rendus' 1855 pages 855, 1039.), who carefully studied the development of double monsters in fishes, observed in fifteen instances the steps by which two heads gradually became united into one. In all such cases it is now thought by the greater number of capable judges that the homologous parts do not attract each other, but that in the words of Mr. Lowne (26/3. 'Catalogue of the Teratological Series in the Museum of the R. Coll. of Surgeons' 1872 page 16.): "As union takes place before the differentiation of distinct organs occurs, these are formed in continuity with each other." He adds that organs already differentiated probably in no case become united to homologous ones. M. Dareste does not speak (26/4. 'Archives de Zoolog. Exper.' January 1874 page 78.) quite decisively against the law of soi pour soi, but concludes by saying, "On se rend parfaitement compte de la formation des monstres, si l'on admet que les embryons qui se soudent appartiennent a un meme oeuf; qu'ils s'unissent en meme temps qu'ils se forment, et que la soudure ne se produit que pendant la premiere periode de la vie embryonnaire, celle ou les organes ne sont encore constitues que par des blastemes homogenes."

By whatever means the abnormal fusion of homologous parts is effected, such cases throw light on the frequent presence of organs which are double during an embryonic period (and throughout life in other and lower members of the same class) but which afterwards unite by a normal process into a single medial organ. In the vegetable kingdom Moquin-Tandon (26/5. 'Teratologie Veg.' 1841 livre 3.) gives a long list of cases, showing how frequently homologous parts, such as leaves, petals, stamens, and pistils, flowers, and aggregates of homologous parts, such as buds, as well as fruit, become blended, both normally and abnormally, with perfect symmetry into one another.

THE VARIABILITY OF MULTIPLE AND HOMOLOGOUS PARTS.

Isidore Geoffroy (26/6. 'Hist. des Anomalies' tome 3 pages 4, 5, 6.) insists that, when any part or organ is repeated many times in the same animal, it is particularly liable to vary both in number and structure. With respect to number, the proposition may, I think, be considered as fully established; but the evidence is chiefly derived from organic beings living under their natural conditions, with which we are not here concerned. Whenever such parts as the vertebrae or teeth, the rays in the fins of fishes, or the feathers in the tails of birds, or petals, stamens, pistils, or seeds, are very numerous, the number is generally variable. With respect to the structure of multiple parts, the evidence of variability is not so decisive; but the fact, as far as it may be trusted, probably depends on multiple parts being of less physiological importance than single parts; consequently their structure has been less rigorously guarded by natural selection.

COMPENSATION OF GROWTH, OR BALANCEMENT.

This law, as applied to natural species, was propounded by Goethe and Geoffroy Saint-Hilaire at nearly the same time. It implies that, when much organised matter is used in building up some one part, other parts are starved and become reduced. Several authors, especially botanists, believe in this law; others reject it. As far as I can judge, it occasionally holds good; but its importance has probably been exaggerated. It is scarcely possible to distinguish between the supposed effects of such compensation, and the effects of long-continued selection which may lead to the augmentation of one part, and simultaneously to the diminution of another. Anyhow, there can be no doubt that an organ may be greatly increased without any corresponding diminution of an adjoining part. To recur to our former illustration of the Irish elk, it may be asked what part has suffered in consequence of the immense development of the horns?

It has already been observed that the struggle for existence does not bear hard on our domesticated productions, and consequently the principle of economy of growth will seldom come into play, so that we ought not to expect to find with them frequent evidence of compensation. We have, however, some such cases. Moquin-Tandon describes a monstrous bean (26/7. 'Teratologie Veg.' page 156. See also my book on 'The Movements and Habits of Climbing Plants' 2nd edition 1875 page 202.), in which the stipules were enormously developed, and the leaflets apparently in consequence completely aborted; this case is interesting, as it represents the natural condition of Lathyrus aphaca, with its stipules of great size, and its leaves reduced to mere threads, which act as tendrils. De Candolle (26/8. 'Memoires du Museum' etc. tome 8 page 178.) has remarked that the varieties of Raphanus sativus which have small roots yield numerous seed containing much oil, whilst those with large roots are not productive in oil; and so it is with Brassica asperifolia. The varieties of Cucurbita pepo which bear large fruit yield a small crop, according to Naudin; whilst those producing small fruit yield a vast number. Lastly, I have endeavoured to show in the eighteenth chapter that with many cultivated plants unnatural treatment checks the full and proper action of the reproductive organs, and they are thus rendered more or less sterile; consequently, in the way of compensation, the fruit becomes greatly enlarged, and, in double flowers, the petals are greatly increased in number.

With animals, it has been found difficult to produce cows which yield much milk, and are afterwards capable of fattening well. With fowls which have large top-knots and beards the comb and wattles are generally much reduced in size; though there are exceptions to this rule. Perhaps the entire absence of the oil-gland in fantail pigeons may be connected with the great development of their tails.

MECHANICAL PRESSURE AS A CAUSE OF MODIFICATIONS.

In some few cases there is reason to believe that mere mechanical pressure has affected certain structures. Vrolik and Weber (26/9. Prichard 'Phys. Hist. of Mankind' 1851 volume 1 page 324.) maintain that the shape of the human head is influenced by the shape of the mother's pelvis. The kidneys in different birds differ much in form, and St. Ange (26/10. 'Annales des Sc. Nat.' 1st series tome 19 page 327.) believes that this is determined by the form of the pelvis, which again, no doubt, stands in close relation with their power of locomotion. In snakes, the viscera are curiously displaced, in comparison with their position in other vertebrates; and this has been attributed by some authors to the elongation of their bodies; but here, as in so many previous cases, it is impossible to disentangle a direct result of this kind from that consequent on natural selection. Godron has argued (26/11. 'Comptes Rendus' December 1864 page 1039.) that the abortion of the spur on the inner side of the flowers in Corydalis, is caused by the buds at a very early period of growth whilst underground being closely pressed against one another and against the stem. Some botanists believe that the singular difference in the shape both of the seed and corolla, in the interior and exterior florets in certain Compositous and Umbelliferous plants, is due to the pressure to which the inner florets are subjected; but this conclusion is doubtful.

The facts just given do not relate to domesticated productions, and therefore do not strictly concern us. But here is a more appropriate case: H. Muller (26/12. "Ueber fotale Rachites" 'Wurzburger Medicin. Zeitschrift' 1860 b. 1 s. 265.) has shown that in shortfaced races of the dog some of the molar teeth are placed in a slightly different position to that which they occupy in other dogs, especially in those having elongated muzzles; and as he remarks, any inherited change in the arrangement of the teeth deserves notice, considering their classificatory importance. This difference in position is due to the shortening of certain facial bones and the consequent want of space; and the shortening results from a peculiar and abnormal state of the embryonal cartilages of the bones.

[RELATIVE POSITION OF FLOWERS WITH RESPECT TO THE AXIS, AND OF SEEDS IN THE OVARY, AS INDUCING VARIATION.

In the thirteenth chapter various peloric flowers were described, and their production was shown to be due either to arrested development, or to reversion to a primordial condition. Moquin-Tandon has remarked that the flowers which stand on the summit of the main stem or of a lateral branch are more liable to become peloric than those on the sides (26/13. 'Teratologie Veg.' page 192.); and he adduces, amongst other instances, that of Teucrium campanulatum. In another Labiate plant grown by me, viz., the Galeobdolon luteum, the peloric flowers were always produced on the summit of the stem, where flowers are not usually borne. In Pelargonium, a SINGLE flower in the truss is frequently peloric, and when this occurs I have during several years invariably observed it to be the central flower. This is of such frequent occurrence that one observer (26/14. 'Journal of Horticulture' July 2, 1861 page 253.) gives the names of ten varieties flowering at the same time, in every one of which the central flower was peloric. Occasionally more than one flower in the truss is peloric, and then of course the additional ones must be lateral. These flowers are interesting as showing how the whole structure is correlated. In the common Pelargonium the upper sepal is produced into a nectary which coheres with the flower-peduncle; the two upper petals differ a little in shape from the three lower ones, and are marked with dark shades of colour; the stamens are graduated in length and upturned. In the peloric flowers, the nectary aborts; all the petals become alike both in shape and colour; the stamens are generally reduced in number and become straight, so that the whole flower resembles that of the allied genus Erodium. The correlation between these changes is well shown when one of the two upper petals alone loses its dark mark, for in this case the nectary does not entirely abort, but is usually much reduced in length. (26/15. It would be worth trial to fertilise with the same pollen the central and lateral flowers of the pelargonium, or of other highly cultivated plants, protecting them of course from insects: then to sow the seed separately, and observe whether the one or the other lot of seedlings varied the most.)