The Descent of Man, and Selection in Relation to Sex

In the United States, above 1,000,000 soldiers, who served in the late war, were measured, and the States in which they were born and reared were recorded. (17. 'Investigations in Military and Anthrop. Statistics,' etc., 1869, by B.A. Gould, pp. 93, 107, 126, 131, 134.) From this astonishing number of observations it is proved that local influences of some kind act directly on stature; and we further learn that "the State where the physical growth has in great measure taken place, and the State of birth, which indicates the ancestry, seem to exert a marked influence on the stature." For instance, it is established, "that residence in the Western States, during the years of growth, tends to produce increase of stature." On the other hand, it is certain that with sailors, their life delays growth, as shewn "by the great difference between the statures of soldiers and sailors at the ages of seventeen and eighteen years." Mr. B.A. Gould endeavoured to ascertain the nature of the influences which thus act on stature; but he arrived only at negative results, namely that they did not relate to climate, the elevation of the land, soil, nor even "in any controlling degree" to the abundance or the need of the comforts of life. This latter conclusion is directly opposed to that arrived at by Villerme, from the statistics of the height of the conscripts in different parts of France. When we compare the differences in stature between the Polynesian chiefs and the lower orders within the same islands, or between the inhabitants of the fertile volcanic and low barren coral islands of the same ocean (18. For the Polynesians, see Prichard's 'Physical History of Mankind,' vol. v. 1847, pp. 145, 283. Also Godron, 'De l'Espèce,' tom. ii. p. 289. There is also a remarkable difference in appearance between the closely-allied Hindoos inhabiting the Upper Ganges and Bengal; see Elphinstone's 'History of India,' vol. i. p. 324.) or again between the Fuegians on the eastern and western shores of their country, where the means of subsistence are very different, it is scarcely possible to avoid the conclusion that better food and greater comfort do influence stature. But the preceding statements shew how difficult it is to arrive at any precise result. Dr. Beddoe has lately proved that, with the inhabitants of Britain, residence in towns and certain occupations have a deteriorating influence on height; and he infers that the result is to a certain extent inherited, as is likewise the case in the United States. Dr. Beddoe further believes that wherever a "race attains its maximum of physical development, it rises highest in energy and moral vigour." (19. 'Memoirs, Anthropological Society,' vol. iii. 1867-69, pp. 561, 565, 567.)

Whether external conditions produce any other direct effect on man is not known. It might have been expected that differences of climate would have had a marked influence, inasmuch as the lungs and kidneys are brought into activity under a low temperature, and the liver and skin under a high one. (20. Dr. Brakenridge, 'Theory of Diathesis,' 'Medical Times,' June 19 and July 17, 1869.) It was formerly thought that the colour of the skin and the character of the hair were determined by light or heat; and although it can hardly be denied that some effect is thus produced, almost all observers now agree that the effect has been very small, even after exposure during many ages. But this subject will be more properly discussed when we treat of the different races of mankind. With our domestic animals there are grounds for believing that cold and damp directly affect the growth of the hair; but I have not met with any evidence on this head in the case of man.

EFFECTS OF THE INCREASED USE AND DISUSE OF PARTS

It is well known that use strengthens the muscles in the individual, and complete disuse, or the destruction of the proper nerve, weakens them. When the eye is destroyed, the optic nerve often becomes atrophied. When an artery is tied, the lateral channels increase not only in diameter, but in the thickness and strength of their coats. When one kidney ceases to act from disease, the other increases in size, and does double work. Bones increase not only in thickness, but in length, from carrying a greater weight. (21. I have given authorities for these several statements in my 'Variation of Animals and Plants under Domestication,' vol. ii. pp. 297- 300. Dr. Jaeger, "Über das Langenwachsthum der Knochen," 'Jenäischen Zeitschrift,' B. v. Heft. i.) Different occupations, habitually followed, lead to changed proportions in various parts of the body. Thus it was ascertained by the United States Commission (22. 'Investigations,' etc., by B.A. Gould, 1869, p. 288.) that the legs of the sailors employed in the late war were longer by 0.217 of an inch than those of the soldiers, though the sailors were on an average shorter men; whilst their arms were shorter by 1.09 of an inch, and therefore, out of proportion, shorter in relation to their lesser height. This shortness of the arms is apparently due to their greater use, and is an unexpected result: but sailors chiefly use their arms in pulling, and not in supporting weights. With sailors, the girth of the neck and the depth of the instep are greater, whilst the circumference of the chest, waist, and hips is less, than in soldiers.

Whether the several foregoing modifications would become hereditary, if the same habits of life were followed during many generations, is not known, but it is probable. Rengger (23. 'Säugethiere von Paraguay,' 1830, s. 4.) attributes the thin legs and thick arms of the Payaguas Indians to successive generations having passed nearly their whole lives in canoes, with their lower extremities motionless. Other writers have come to a similar conclusion in analogous cases. According to Cranz (24. 'History of Greenland,' Eng. translat., 1767, vol. i. p. 230.), who lived for a long time with the Esquimaux, "the natives believe that ingenuity and dexterity in seal-catching (their highest art and virtue) is hereditary; there is really something in it, for the son of a celebrated seal-catcher will distinguish himself, though he lost his father in childhood." But in this case it is mental aptitude, quite as much as bodily structure, which appears to be inherited. It is asserted that the hands of English labourers are at birth larger than those of the gentry. (25. 'Intermarriage,' by Alex. Walker, 1838, p. 377.) From the correlation which exists, at least in some cases (26. 'The Variation of Animals under Domestication,' vol. i. p. 173.), between the development of the extremities and of the jaws, it is possible that in those classes which do not labour much with their hands and feet, the jaws would be reduced in size from this cause. That they are generally smaller in refined and civilised men than in hard-working men or savages, is certain. But with savages, as Mr. Herbert Spencer (27. 'Principles of Biology,' vol. i. p. 455.) has remarked, the greater use of the jaws in chewing coarse, uncooked food, would act in a direct manner on the masticatory muscles, and on the bones to which they are attached. In infants, long before birth, the skin on the soles of the feet is thicker than on any other part of the body; (28. Paget, 'Lectures on Surgical Pathology,' vol. ii, 1853, p. 209.) and it can hardly be doubted that this is due to the inherited effects of pressure during a long series of generations.

It is familiar to every one that watchmakers and engravers are liable to be short-sighted, whilst men living much out of doors, and especially savages, are generally long-sighted. (29. It is a singular and unexpected fact that sailors are inferior to landsmen in their mean distance of distinct vision. Dr. B.A. Gould ('Sanitary Memoirs of the War of the Rebellion,' 1869, p. 530), has proved this to be the case; and he accounts for it by the ordinary range of vision in sailors being "restricted to the length of the vessel and the height of the masts.") Short-sight and long-sight certainly tend to be inherited. (30. 'The Variation of Animals under Domestication,' vol. i. p. 8.) The inferiority of Europeans, in comparison with savages, in eyesight and in the other senses, is no doubt the accumulated and transmitted effect of lessened use during many generations; for Rengger (31. 'Säugethiere von Paraguay,' s. 8, 10. I have had good opportunities for observing the extraordinary power of eyesight in the Fuegians. See also Lawrence ('Lectures on Physiology,' etc., 1822, p. 404) on this same subject. M. Giraud-Teulon has recently collected ('Revue des Cours Scientifiques,' 1870, p. 625) a large and valuable body of evidence proving that the cause of short-sight, "C'est le travail assidu, de près.") states that he has repeatedly observed Europeans, who had been brought up and spent their whole lives with the wild Indians, who nevertheless did not equal them in the sharpness of their senses. The same naturalist observes that the cavities in the skull for the reception of the several sense- organs are larger in the American aborigines than in Europeans; and this probably indicates a corresponding difference in the dimensions of the organs themselves. Blumenbach has also remarked on the large size of the nasal cavities in the skulls of the American aborigines, and connects this fact with their remarkably acute power of smell. The Mongolians of the plains of northern Asia, according to Pallas, have wonderfully perfect senses; and Prichard believes that the great breadth of their skulls across the zygomas follows from their highly-developed sense organs. (32. Prichard, 'Physical History of Mankind,' on the authority of Blumenbach, vol. i. 1851, p. 311; for the statement by Pallas, vol. iv. 1844, p. 407.)

The Quechua Indians inhabit the lofty plateaux of Peru; and Alcide d'Orbigny states (33. Quoted by Prichard, 'Researches into the Physical History of Mankind,' vol. v. p. 463.) that, from continually breathing a highly rarefied atmosphere, they have acquired chests and lungs of extraordinary dimensions. The cells, also, of the lungs are larger and more numerous than in Europeans. These observations have been doubted, but Mr. D. Forbes carefully measured many Aymaras, an allied race, living at the height of between 10,000 and 15,000 feet; and he informs me (34. Mr. Forbes' valuable paper is now published in the 'Journal of the Ethnological Society of London,' new series, vol. ii. 1870, p.193.) that they differ conspicuously from the men of all other races seen by him in the circumference and length of their bodies. In his table of measurements, the stature of each man is taken at 1000, and the other measurements are reduced to this standard. It is here seen that the extended arms of the Aymaras are shorter than those of Europeans, and much shorter than those of Negroes. The legs are likewise shorter; and they present this remarkable peculiarity, that in every Aymara measured, the femur is actually shorter than the tibia. On an average, the length of the femur to that of the tibia is as 211 to 252; whilst in two Europeans, measured at the same time, the femora to the tibiae were as 244 to 230; and in three Negroes as 258 to 241. The humerus is likewise shorter relatively to the forearm. This shortening of that part of the limb which is nearest to the body, appears to be, as suggested to me by Mr. Forbes, a case of compensation in relation with the greatly increased length of the trunk. The Aymaras present some other singular points of structure, for instance, the very small projection of the heel.

These men are so thoroughly acclimatised to their cold and lofty abode, that when formerly carried down by the Spaniards to the low eastern plains, and when now tempted down by high wages to the gold-washings, they suffer a frightful rate of mortality. Nevertheless Mr. Forbes found a few pure families which had survived during two generations: and he observed that they still inherited their characteristic peculiarities. But it was manifest, even without measurement, that these peculiarities had all decreased; and on measurement, their bodies were found not to be so much elongated as those of the men on the high plateau; whilst their femora had become somewhat lengthened, as had their tibiae, although in a less degree. The actual measurements may be seen by consulting Mr. Forbes's memoir. From these observations, there can, I think, be no doubt that residence during many generations at a great elevation tends, both directly and indirectly, to induce inherited modifications in the proportions of the body. (35. Dr. Wilckens ('Landwirthschaft. Wochenblatt,' No. 10, 1869) has lately published an interesting essay shewing how domestic animals, which live in mountainous regions, have their frames modified.)

Although man may not have been much modified during the latter stages of his existence through the increased or decreased use of parts, the facts now given shew that his liability in this respect has not been lost; and we positively know that the same law holds good with the lower animals. Consequently we may infer that when at a remote epoch the progenitors of man were in a transitional state, and were changing from quadrupeds into bipeds, natural selection would probably have been greatly aided by the inherited effects of the increased or diminished use of the different parts of the body.

ARRESTS OF DEVELOPMENT

There is a difference between arrested development and arrested growth, for parts in the former state continue to grow whilst still retaining their early condition. Various monstrosities come under this head; and some, as a cleft palate, are known to be occasionally inherited. It will suffice for our purpose to refer to the arrested brain-development of microcephalous idiots, as described in Vogt's memoir. (36. 'Mémoire sur les Microcephales,' 1867, pp. 50, 125, 169, 171, 184-198.) Their skulls are smaller, and the convolutions of the brain are less complex than in normal men. The frontal sinus, or the projection over the eye-brows, is largely developed, and the jaws are prognathous to an "effrayant" degree; so that these idiots somewhat resemble the lower types of mankind. Their intelligence, and most of their mental faculties, are extremely feeble. They cannot acquire the power of speech, and are wholly incapable of prolonged attention, but are much given to imitation. They are strong and remarkably active, continually gambolling and jumping about, and making grimaces. They often ascend stairs on all-fours; and are curiously fond of climbing up furniture or trees. We are thus reminded of the delight shewn by almost all boys in climbing trees; and this again reminds us how lambs and kids, originally alpine animals, delight to frisk on any hillock, however small. Idiots also resemble the lower animals in some other respects; thus several cases are recorded of their carefully smelling every mouthful of food before eating it. One idiot is described as often using his mouth in aid of his hands, whilst hunting for lice. They are often filthy in their habits, and have no sense of decency; and several cases have been published of their bodies being remarkably hairy. (37. Prof. Laycock sums up the character of brute-like idiots by calling them "theroid;" 'Journal of Mental Science,' July 1863. Dr. Scott ('The Deaf and Dumb,' 2nd ed. 1870, p. 10) has often observed the imbecile smelling their food. See, on this same subject, and on the hairiness of idiots, Dr. Maudsley, 'Body and Mind,' 1870, pp. 46-51. Pinel has also given a striking case of hairiness in an idiot.)

REVERSION

Many of the cases to be here given, might have been introduced under the last heading. When a structure is arrested in its development, but still continues growing, until it closely resembles a corresponding structure in some lower and adult member of the same group, it may in one sense be considered as a case of reversion. The lower members in a group give us some idea how the common progenitor was probably constructed; and it is hardly credible that a complex part, arrested at an early phase of embryonic development, should go on growing so as ultimately to perform its proper function, unless it had acquired such power during some earlier state of existence, when the present exceptional or arrested structure was normal. The simple brain of a microcephalous idiot, in as far as it resembles that of an ape, may in this sense be said to offer a case of reversion. (38. In my 'Variation of Animals under Domestication' (vol. ii. p. 57), I attributed the not very rare cases of supernumerary mammae in women to reversion. I was led to this as a probable conclusion, by the additional mammae being generally placed symmetrically on the breast; and more especially from one case, in which a single efficient mamma occurred in the inguinal region of a woman, the daughter of another woman with supernumerary mammae. But I now find (see, for instance, Prof. Preyer, 'Der Kampf um das Dasein,' 1869, s. 45) that mammae erraticae, occur in other situations, as on the back, in the armpit, and on the thigh; the mammae in this latter instance having given so much milk that the child was thus nourished. The probability that the additional mammae are due to reversion is thus much weakened; nevertheless, it still seems to me probable, because two pairs are often found symmetrically on the breast; and of this I myself have received information in several cases. It is well known that some Lemurs normally have two pairs of mammae on the breast. Five cases have been recorded of the presence of more than a pair of mammae (of course rudimentary) in the male sex of mankind; see 'Journal of Anat. and Physiology,' 1872, p. 56, for a case given by Dr. Handyside, in which two brothers exhibited this peculiarity; see also a paper by Dr. Bartels, in 'Reichert's and du Bois-Reymond's Archiv.,' 1872, p. 304. In one of the cases alluded to by Dr. Bartels, a man bore five mammae, one being medial and placed above the navel; Meckel von Hemsbach thinks that this latter case is illustrated by a medial mamma occurring in certain Cheiroptera. On the whole, we may well doubt if additional mammae would ever have been developed in both sexes of mankind, had not his early progenitors been provided with more than a single pair.

In the above work (vol. ii. p. 12), I also attributed, though with much hesitation, the frequent cases of polydactylism in men and various animals to reversion. I was partly led to this through Prof. Owen's statement, that some of the Ichthyopterygia possess more than five digits, and therefore, as I supposed, had retained a primordial condition; but Prof. Gegenbaur ('Jenaischen Zeitschrift,' B. v. Heft 3, s. 341), disputes Owen's conclusion. On the other hand, according to the opinion lately advanced by Dr. Gunther, on the paddle of Ceratodus, which is provided with articulated bony rays on both sides of a central chain of bones, there seems no great difficulty in admitting that six or more digits on one side, or on both sides, might reappear through reversion. I am informed by Dr. Zouteveen that there is a case on record of a man having twenty-four fingers and twenty-four toes! I was chiefly led to the conclusion that the presence of supernumerary digits might be due to reversion from the fact that such digits, not only are strongly inherited, but, as I then believed, had the power of regrowth after amputation, like the normal digits of the lower vertebrata. But I have explained in the second edition of my Variation under Domestication why I now place little reliance on the recorded cases of such regrowth. Nevertheless it deserves notice, inasmuch as arrested development and reversion are intimately related processes; that various structures in an embryonic or arrested condition, such as a cleft palate, bifid uterus, etc., are frequently accompanied by polydactylism. This has been strongly insisted on by Meckel and Isidore Geoffroy St. – Hilaire. But at present it is the safest course to give up altogether the idea that there is any relation between the development of supernumerary digits and reversion to some lowly organised progenitor of man.) There are other cases which come more strictly under our present head of reversion. Certain structures, regularly occurring in the lower members of the group to which man belongs, occasionally make their appearance in him, though not found in the normal human embryo; or, if normally present in the human embryo, they become abnormally developed, although in a manner which is normal in the lower members of the group. These remarks will be rendered clearer by the following illustrations.

In various mammals the uterus graduates from a double organ with two distinct orifices and two passages, as in the marsupials, into a single organ, which is in no way double except from having a slight internal fold, as in the higher apes and man. The rodents exhibit a perfect series of gradations between these two extreme states. In all mammals the uterus is developed from two simple primitive tubes, the inferior portions of which form the cornua; and it is in the words of Dr. Farre, "by the coalescence of the two cornua at their lower extremities that the body of the uterus is formed in man; while in those animals in which no middle portion or body exists, the cornua remain ununited. As the development of the uterus proceeds, the two cornua become gradually shorter, until at length they are lost, or, as it were, absorbed into the body of the uterus." The angles of the uterus are still produced into cornua, even in animals as high up in the scale as the lower apes and lemurs.

Now in women, anomalous cases are not very infrequent, in which the mature uterus is furnished with cornua, or is partially divided into two organs; and such cases, according to Owen, repeat "the grade of concentrative development," attained by certain rodents. Here perhaps we have an instance of a simple arrest of embryonic development, with subsequent growth and perfect functional development; for either side of the partially double uterus is capable of performing the proper office of gestation. In other and rarer cases, two distinct uterine cavities are formed, each having its proper orifice and passage. (39. See Dr. A. Farre's well-known article in the 'Cyclopaedia of Anatomy and Physiology,' vol. v. 1859, p. 642. Owen, 'Anatomy of Vertebrates,' vol. iii. 1868, p. 687. Professor Turner, in 'Edinburgh Medical Journal,' February, 1865.) No such stage is passed through during the ordinary development of the embryo; and it is difficult to believe, though perhaps not impossible, that the two simple, minute, primitive tubes should know how (if such an expression may be used) to grow into two distinct uteri, each with a well-constructed orifice and passage, and each furnished with numerous muscles, nerves, glands and vessels, if they had not formerly passed through a similar course of development, as in the case of existing marsupials. No one will pretend that so perfect a structure as the abnormal double uterus in woman could be the result of mere chance. But the principle of reversion, by which a long-lost structure is called back into existence, might serve as the guide for its full development, even after the lapse of an enormous interval of time.

Professor Canestrini, after discussing the foregoing and various analogous cases, arrives at the same conclusion as that just given. He adduces another instance, in the case of the malar bone (40. 'Annuario della Soc. dei Naturalisti,' Modena, 1867, p. 83. Prof. Canestrini gives extracts on this subject from various authorities. Laurillard remarks, that as he has found a complete similarity in the form, proportions, and connection of the two malar bones in several human subjects and in certain apes, he cannot consider this disposition of the parts as simply accidental. Another paper on this same anomaly has been published by Dr. Saviotti in the 'Gazzetta delle Cliniche,' Turin, 1871, where he says that traces of the division may be detected in about two per cent. of adult skulls; he also remarks that it more frequently occurs in prognathous skulls, not of the Aryan race, than in others. See also G. Delorenzi on the same subject; 'Tre nuovi casi d'anomalia dell' osso malare,' Torino, 1872. Also, E. Morselli, 'Sopra una rara anomalia dell' osso malare,' Modena, 1872. Still more recently Gruber has written a pamphlet on the division of this bone. I give these references because a reviewer, without any grounds or scruples, has thrown doubts on my statements.), which, in some of the Quadrumana and other mammals, normally consists of two portions. This is its condition in the human foetus when two months old; and through arrested development, it sometimes remains thus in man when adult, more especially in the lower prognathous races. Hence Canestrini concludes that some ancient progenitor of man must have had this bone normally divided into two portions, which afterwards became fused together. In man the frontal bone consists of a single piece, but in the embryo, and in children, and in almost all the lower mammals, it consists of two pieces separated by a distinct suture. This suture occasionally persists more or less distinctly in man after maturity; and more frequently in ancient than in recent crania, especially, as Canestrini has observed, in those exhumed from the Drift, and belonging to the brachycephalic type. Here again he comes to the same conclusion as in the analogous case of the malar bones. In this, and other instances presently to be given, the cause of ancient races approaching the lower animals in certain characters more frequently than do the modern races, appears to be, that the latter stand at a somewhat greater distance in the long line of descent from their early semi-human progenitors.