The Descent of Man, and Selection in Relation to Sex
In some of the lower Quadrumana, in the Lemuridae and Carnivora, as well as in many marsupials, there is a passage near the lower end of the humerus, called the supra-condyloid foramen, through which the great nerve of the fore limb and often the great artery pass. Now in the humerus of man, there is generally a trace of this passage, which is sometimes fairly well developed, being formed by a depending hook-like process of bone, completed by a band of ligament. Dr. Struthers (49. With respect to inheritance, see Dr. Struthers in the 'Lancet,' Feb. 15, 1873, and another important paper, ibid. Jan. 24, 1863, p. 83. Dr. Knox, as I am informed, was the first anatomist who drew attention to this peculiar structure in man; see his 'Great Artists and Anatomists,' p. 63. See also an important memoir on this process by Dr. Gruber, in the 'Bulletin de l'Acad. Imp. de St. Petersbourg,' tom. xii. 1867, p. 448.), who has closely attended to the subject, has now shewn that this peculiarity is sometimes inherited, as it has occurred in a father, and in no less than four out of his seven children. When present, the great nerve invariably passes through it; and this clearly indicates that it is the homologue and rudiment of the supra-condyloid foramen of the lower animals. Prof. Turner estimates, as he informs me, that it occurs in about one per cent. of recent skeletons. But if the occasional development of this structure in man is, as seems probable, due to reversion, it is a return to a very ancient state of things, because in the higher Quadrumana it is absent.
There is another foramen or perforation in the humerus, occasionally present in man, which may be called the inter-condyloid. This occurs, but not constantly, in various anthropoid and other apes (50. Mr. St. George Mivart, 'Transactions Phil. Soc.' 1867, p. 310.), and likewise in many of the lower animals. It is remarkable that this perforation seems to have been present in man much more frequently during ancient times than recently. Mr. Busk (51. "On the Caves of Gibraltar," 'Transactions of the International Congress of Prehistoric Archaeology,' Third Session, 1869, p. 159. Prof. Wyman has lately shewn (Fourth Annual Report, Peabody Museum, 1871, p. 20), that this perforation is present in thirty-one per cent. of some human remains from ancient mounds in the Western United States, and in Florida. It frequently occurs in the negro.) has collected the following evidence on this head: Prof. Broca "noticed the perforation in four and a half per cent. of the arm-bones collected in the 'Cimetière du Sud,' at Paris; and in the Grotto of Orrony, the contents of which are referred to the Bronze period, as many as eight humeri out of thirty-two were perforated; but this extraordinary proportion, he thinks, might be due to the cavern having been a sort of 'family vault.' Again, M. Dupont found thirty per cent. of perforated bones in the caves of the Valley of the Lesse, belonging to the Reindeer period; whilst M. Leguay, in a sort of dolmen at Argenteuil, observed twenty-five per cent. to be perforated; and M. Pruner-Bey found twenty-six per cent. in the same condition in bones from Vaureal. Nor should it be left unnoticed that M. Pruner-Bey states that this condition is common in Guanche skeletons." It is an interesting fact that ancient races, in this and several other cases, more frequently present structures which resemble those of the lower animals than do the modern. One chief cause seems to be that the ancient races stand somewhat nearer in the long line of descent to their remote animal-like progenitors.
In man, the os coccyx, together with certain other vertebrae hereafter to be described, though functionless as a tail, plainly represent this part in other vertebrate animals. At an early embryonic period it is free, and projects beyond the lower extremities; as may be seen in the drawing (Fig. 1.) of a human embryo. Even after birth it has been known, in certain rare and anomalous cases (52. Quatrefages has lately collected the evidence on this subject. 'Revue des Cours Scientifiques,' 1867-1868, p. 625. In 1840 Fleischmann exhibited a human foetus bearing a free tail, which, as is not always the case, included vertebral bodies; and this tail was critically examined by the many anatomists present at the meeting of naturalists at Erlangen (see Marshall in Niederlandischen Archiv für Zoologie, December 1871).), to form a small external rudiment of a tail. The os coccyx is short, usually including only four vertebrae, all anchylosed together: and these are in a rudimentary condition, for they consist, with the exception of the basal one, of the centrum alone. (53. Owen, 'On the Nature of Limbs,' 1849, p. 114.) They are furnished with some small muscles; one of which, as I am informed by Prof. Turner, has been expressly described by Theile as a rudimentary repetition of the extensor of the tail, a muscle which is so largely developed in many mammals.
The spinal cord in man extends only as far downwards as the last dorsal or first lumbar vertebra; but a thread-like structure (the filum terminale) runs down the axis of the sacral part of the spinal canal, and even along the back of the coccygeal bones. The upper part of this filament, as Prof. Turner informs me, is undoubtedly homologous with the spinal cord; but the lower part apparently consists merely of the pia mater, or vascular investing membrane. Even in this case the os coccyx may be said to possess a vestige of so important a structure as the spinal cord, though no longer enclosed within a bony canal. The following fact, for which I am also indebted to Prof. Turner, shews how closely the os coccyx corresponds with the true tail in the lower animals: Luschka has recently discovered at the extremity of the coccygeal bones a very peculiar convoluted body, which is continuous with the middle sacral artery; and this discovery led Krause and Meyer to examine the tail of a monkey (Macacus), and of a cat, in both of which they found a similarly convoluted body, though not at the extremity.
The reproductive system offers various rudimentary structures; but these differ in one important respect from the foregoing cases. Here we are not concerned with the vestige of a part which does not belong to the species in an efficient state, but with a part efficient in the one sex, and represented in the other by a mere rudiment. Nevertheless, the occurrence of such rudiments is as difficult to explain, on the belief of the separate creation of each species, as in the foregoing cases. Hereafter I shall have to recur to these rudiments, and shall shew that their presence generally depends merely on inheritance, that is, on parts acquired by one sex having been partially transmitted to the other. I will in this place only give some instances of such rudiments. It is well known that in the males of all mammals, including man, rudimentary mammae exist. These in several instances have become well developed, and have yielded a copious supply of milk. Their essential identity in the two sexes is likewise shewn by their occasional sympathetic enlargement in both during an attack of the measles. The vesicula prostatica, which has been observed in many male mammals, is now universally acknowledged to be the homologue of the female uterus, together with the connected passage. It is impossible to read Leuckart's able description of this organ, and his reasoning, without admitting the justness of his conclusion. This is especially clear in the case of those mammals in which the true female uterus bifurcates, for in the males of these the vesicula likewise bifurcates. (54. Leuckart, in Todd's 'Cyclopaedia of Anatomy' 1849-52, vol. iv. p. 1415. In man this organ is only from three to six lines in length, but, like so many other rudimentary parts, it is variable in development as well as in other characters.) Some other rudimentary structures belonging to the reproductive system might have been here adduced. (55. See, on this subject, Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 675, 676, 706.)
The bearing of the three great classes of facts now given is unmistakeable. But it would be superfluous fully to recapitulate the line of argument given in detail in my 'Origin of Species.' The homological construction of the whole frame in the members of the same class is intelligible, if we admit their descent from a common progenitor, together with their subsequent adaptation to diversified conditions. On any other view, the similarity of pattern between the hand of a man or monkey, the foot of a horse, the flipper of a seal, the wing of a bat, etc., is utterly inexplicable. (56. Prof. Bianconi, in a recently published work, illustrated by admirable engravings ('La Théorie Darwinienne et la création dite indépendante,' 1874), endeavours to shew that homological structures, in the above and other cases, can be fully explained on mechanical principles, in accordance with their uses. No one has shewn so well, how admirably such structures are adapted for their final purpose; and this adaptation can, as I believe, be explained through natural selection. In considering the wing of a bat, he brings forward (p. 218) what appears to me (to use Auguste Comte's words) a mere metaphysical principle, namely, the preservation "in its integrity of the mammalian nature of the animal." In only a few cases does he discuss rudiments, and then only those parts which are partially rudimentary, such as the little hoofs of the pig and ox, which do not touch the ground; these he shews clearly to be of service to the animal. It is unfortunate that he did not consider such cases as the minute teeth, which never cut through the jaw in the ox, or the mammae of male quadrupeds, or the wings of certain beetles, existing under the soldered wing-covers, or the vestiges of the pistil and stamens in various flowers, and many other such cases. Although I greatly admire Prof. Bianconi's work, yet the belief now held by most naturalists seems to me left unshaken, that homological structures are inexplicable on the principle of mere adaptation.) It is no scientific explanation to assert that they have all been formed on the same ideal plan. With respect to development, we can clearly understand, on the principle of variations supervening at a rather late embryonic period, and being inherited at a corresponding period, how it is that the embryos of wonderfully different forms should still retain, more or less perfectly, the structure of their common progenitor. No other explanation has ever been given of the marvellous fact that the embryos of a man, dog, seal, bat, reptile, etc., can at first hardly be distinguished from each other. In order to understand the existence of rudimentary organs, we have only to suppose that a former progenitor possessed the parts in question in a perfect state, and that under changed habits of life they became greatly reduced, either from simple disuse, or through the natural selection of those individuals which were least encumbered with a superfluous part, aided by the other means previously indicated.
Thus we can understand how it has come to pass that man and all other vertebrate animals have been constructed on the same general model, why they pass through the same early stages of development, and why they retain certain rudiments in common. Consequently we ought frankly to admit their community of descent: to take any other view, is to admit that our own structure, and that of all the animals around us, is a mere snare laid to entrap our judgment. This conclusion is greatly strengthened, if we look to the members of the whole animal series, and consider the evidence derived from their affinities or classification, their geographical distribution and geological succession. It is only our natural prejudice, and that arrogance which made our forefathers declare that they were descended from demi-gods, which leads us to demur to this conclusion. But the time will before long come, when it will be thought wonderful that naturalists, who were well acquainted with the comparative structure and development of man, and other mammals, should have believed that each was the work of a separate act of creation.
CHAPTER II
ON THE MANNER OF DEVELOPMENT OF MAN FROM SOME LOWER FORMVariability of body and mind in man – Inheritance – Causes of variability – Laws of variation the same in man as in the lower animals – Direct action of the conditions of life – Effects of the increased use and disuse of parts – Arrested development – Reversion – Correlated variation – Rate of increase – Checks to increase – Natural selection – Man the most dominant animal in the world – Importance of his corporeal structure – The causes which have led to his becoming erect – Consequent changes of structure – Decrease in size of the canine teeth – Increased size and altered shape of the skull – Nakedness – Absence of a tail – Defenceless condition of man.
It is manifest that man is now subject to much variability. No two individuals of the same race are quite alike. We may compare millions of faces, and each will be distinct. There is an equally great amount of diversity in the proportions and dimensions of the various parts of the body; the length of the legs being one of the most variable points. (1. 'Investigations in Military and Anthropological Statistics of American Soldiers,' by B.A. Gould, 1869, p. 256.) Although in some quarters of the world an elongated skull, and in other quarters a short skull prevails, yet there is great diversity of shape even within the limits of the same race, as with the aborigines of America and South Australia – the latter a race "probably as pure and homogeneous in blood, customs, and language as any in existence" – and even with the inhabitants of so confined an area as the Sandwich Islands. (2. With respect to the "Cranial forms of the American aborigines," see Dr. Aitken Meigs in 'Proc. Acad. Nat. Sci.' Philadelphia, May 1868. On the Australians, see Huxley, in Lyell's 'Antiquity of Man,' 1863, p. 87. On the Sandwich Islanders, Prof. J. Wyman, 'Observations on Crania,' Boston, 1868, p. 18.) An eminent dentist assures me that there is nearly as much diversity in the teeth as in the features. The chief arteries so frequently run in abnormal courses, that it has been found useful for surgical purposes to calculate from 1040 corpses how often each course prevails. (3. 'Anatomy of the Arteries,' by R. Quain. Preface, vol. i. 1844.) The muscles are eminently variable: thus those of the foot were found by Prof. Turner (4. 'Transactions of the Royal Society of Edinburgh,' vol. xxiv. pp. 175, 189.) not to be strictly alike in any two out of fifty bodies; and in some the deviations were considerable. He adds, that the power of performing the appropriate movements must have been modified in accordance with the several deviations. Mr. J. Wood has recorded (5. 'Proceedings Royal Society,' 1867, p. 544; also 1868, pp. 483, 524. There is a previous paper, 1866, p. 229.) the occurrence of 295 muscular variations in thirty-six subjects, and in another set of the same number no less than 558 variations, those occurring on both sides of the body being only reckoned as one. In the last set, not one body out of the thirty-six was "found totally wanting in departures from the standard descriptions of the muscular system given in anatomical text books." A single body presented the extraordinary number of twenty-five distinct abnormalities. The same muscle sometimes varies in many ways: thus Prof. Macalister describes (6. 'Proc. R. Irish Academy,' vol. x. 1868, p. 141.) no less than twenty distinct variations in the palmaris accessorius.
The famous old anatomist, Wolff (7. 'Act. Acad. St. Petersburg,' 1778, part ii. p. 217.), insists that the internal viscera are more variable than the external parts: Nulla particula est quae non aliter et aliter in aliis se habeat hominibus. He has even written a treatise on the choice of typical examples of the viscera for representation. A discussion on the beau-ideal of the liver, lungs, kidneys, etc., as of the human face divine, sounds strange in our ears.
The variability or diversity of the mental faculties in men of the same race, not to mention the greater differences between the men of distinct races, is so notorious that not a word need here be said. So it is with the lower animals. All who have had charge of menageries admit this fact, and we see it plainly in our dogs and other domestic animals. Brehm especially insists that each individual monkey of those which he kept tame in Africa had its own peculiar disposition and temper: he mentions one baboon remarkable for its high intelligence; and the keepers in the Zoological Gardens pointed out to me a monkey, belonging to the New World division, equally remarkable for intelligence. Rengger, also, insists on the diversity in the various mental characters of the monkeys of the same species which he kept in Paraguay; and this diversity, as he adds, is partly innate, and partly the result of the manner in which they have been treated or educated. (8. Brehm, 'Thierleben,' B. i. ss. 58, 87. Rengger, 'Säugethiere von Paraguay,' s. 57.)
I have elsewhere (9. 'Variation of Animals and Plants under Domestication,' vol. ii. chap. xii.) so fully discussed the subject of Inheritance, that I need here add hardly anything. A greater number of facts have been collected with respect to the transmission of the most trifling, as well as of the most important characters in man, than in any of the lower animals; though the facts are copious enough with respect to the latter. So in regard to mental qualities, their transmission is manifest in our dogs, horses, and other domestic animals. Besides special tastes and habits, general intelligence, courage, bad and good temper, etc., are certainly transmitted. With man we see similar facts in almost every family; and we now know, through the admirable labours of Mr. Galton (10. 'Hereditary Genius: an Inquiry into its Laws and Consequences,' 1869.), that genius which implies a wonderfully complex combination of high faculties, tends to be inherited; and, on the other hand, it is too certain that insanity and deteriorated mental powers likewise run in families.
With respect to the causes of variability, we are in all cases very ignorant; but we can see that in man as in the lower animals, they stand in some relation to the conditions to which each species has been exposed, during several generations. Domesticated animals vary more than those in a state of nature; and this is apparently due to the diversified and changing nature of the conditions to which they have been subjected. In this respect the different races of man resemble domesticated animals, and so do the individuals of the same race, when inhabiting a very wide area, like that of America. We see the influence of diversified conditions in the more civilised nations; for the members belonging to different grades of rank, and following different occupations, present a greater range of character than do the members of barbarous nations. But the uniformity of savages has often been exaggerated, and in some cases can hardly be said to exist. (11. Mr. Bates remarks ('The Naturalist on the Amazons,' 1863, vol. ii p. 159), with respect to the Indians of the same South American tribe, "no two of them were at all similar in the shape of the head; one man had an oval visage with fine features, and another was quite Mongolian in breadth and prominence of cheek, spread of nostrils, and obliquity of eyes.") It is, nevertheless, an error to speak of man, even if we look only to the conditions to which he has been exposed, as "far more domesticated" (12. Blumenbach, 'Treatises on Anthropology.' Eng. translat., 1865, p. 205.) than any other animal. Some savage races, such as the Australians, are not exposed to more diversified conditions than are many species which have a wide range. In another and much more important respect, man differs widely from any strictly domesticated animal; for his breeding has never long been controlled, either by methodical or unconscious selection. No race or body of men has been so completely subjugated by other men, as that certain individuals should be preserved, and thus unconsciously selected, from somehow excelling in utility to their masters. Nor have certain male and female individuals been intentionally picked out and matched, except in the well-known case of the Prussian grenadiers; and in this case man obeyed, as might have been expected, the law of methodical selection; for it is asserted that many tall men were reared in the villages inhabited by the grenadiers and their tall wives. In Sparta, also, a form of selection was followed, for it was enacted that all children should be examined shortly after birth; the well-formed and vigorous being preserved, the others left to perish. (13. Mitford's 'History of Greece,' vol. i. p. 282. It appears also from a passage in Xenophon's 'Memorabilia,' B. ii. 4 (to which my attention has been called by the Rev. J.N. Hoare), that it was a well recognised principle with the Greeks, that men ought to select their wives with a view to the health and vigour of their children. The Grecian poet, Theognis, who lived 550 B.C., clearly saw how important selection, if carefully applied, would be for the improvement of mankind. He saw, likewise, that wealth often checks the proper action of sexual selection. He thus writes:
"With kine and horses, Kurnus! we proceedBy reasonable rules, and choose a breedFor profit and increase, at any price:Of a sound stock, without defect or vice.But, in the daily matches that we make,The price is everything: for money's sake,Men marry: women are in marriage givenThe churl or ruffian, that in wealth has thriven,May match his offspring with the proudest race:Thus everything is mix'd, noble and base!If then in outward manner, form, and mind,You find us a degraded, motley kind,Wonder no more, my friend! the cause is plain,And to lament the consequence is vain."(The Works of J. Hookham Frere, vol. ii. 1872, p. 334.))
If we consider all the races of man as forming a single species, his range is enormous; but some separate races, as the Americans and Polynesians, have very wide ranges. It is a well-known law that widely-ranging species are much more variable than species with restricted ranges; and the variability of man may with more truth be compared with that of widely- ranging species, than with that of domesticated animals.
Not only does variability appear to be induced in man and the lower animals by the same general causes, but in both the same parts of the body are affected in a closely analogous manner. This has been proved in such full detail by Godron and Quatrefages, that I need here only refer to their works. (14. Godron, 'De l'Espèce,' 1859, tom. ii. livre 3. Quatrefages, 'Unité de l'Espèce Humaine,' 1861. Also Lectures on Anthropology, given in the 'Revue des Cours Scientifiques,' 1866-1868.) Monstrosities, which graduate into slight variations, are likewise so similar in man and the lower animals, that the same classification and the same terms can be used for both, as has been shewn by Isidore Geoffroy St. – Hilaire. (15. 'Hist. Gen. et Part. des Anomalies de l'Organisation,' in three volumes, tom. i. 1832.) In my work on the variation of domestic animals, I have attempted to arrange in a rude fashion the laws of variation under the following heads: – The direct and definite action of changed conditions, as exhibited by all or nearly all the individuals of the same species, varying in the same manner under the same circumstances. The effects of the long- continued use or disuse of parts. The cohesion of homologous parts. The variability of multiple parts. Compensation of growth; but of this law I have found no good instance in the case of man. The effects of the mechanical pressure of one part on another; as of the pelvis on the cranium of the infant in the womb. Arrests of development, leading to the diminution or suppression of parts. The reappearance of long-lost characters through reversion. And lastly, correlated variation. All these so-called laws apply equally to man and the lower animals; and most of them even to plants. It would be superfluous here to discuss all of them (16. I have fully discussed these laws in my 'Variation of Animals and Plants under Domestication,' vol. ii. chap. xxii. and xxiii. M. J.P. Durand has lately (1868) published a valuable essay, 'De l'Influence des Milieux,' etc. He lays much stress, in the case of plants, on the nature of the soil.); but several are so important, that they must be treated at considerable length.
THE DIRECT AND DEFINITE ACTION OF CHANGED CONDITIONSThis is a most perplexing subject. It cannot be denied that changed conditions produce some, and occasionally a considerable effect, on organisms of all kinds; and it seems at first probable that if sufficient time were allowed this would be the invariable result. But I have failed to obtain clear evidence in favour of this conclusion; and valid reasons may be urged on the other side, at least as far as the innumerable structures are concerned, which are adapted for special ends. There can, however, be no doubt that changed conditions induce an almost indefinite amount of fluctuating variability, by which the whole organisation is rendered in some degree plastic.