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Rainer Maria Rilke - "Die Aufzeichnungen des Malte Laurid Brigge"

Honore de Balzac - "A Prince of Bohemia"

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The Descent of Man and Selection in Relation to Sex

C >> Charles Darwin >> The Descent of Man and Selection in Relation to Sex




The celebrated sculptor, Mr. Woolner, informs me of one little peculiarity
in the external ear, which he has often observed both in men and women, and
of which he perceived the full significance. His attention was first
called to the subject whilst at work on his figure of Puck, to which he had
given pointed ears. He was thus led to examine the ears of various
monkeys, and subsequently more carefully those of man. The peculiarity
consists in a little blunt point, projecting from the inwardly folded
margin, or helix. When present, it is developed at birth, and, according
to Prof. Ludwig Meyer, more frequently in man than in woman. Mr. Woolner
made an exact model of one such case, and sent me the accompanying drawing.
(Fig. 2). These points not only project inwards towards the centre of the
ear, but often a little outwards from its plane, so as to be visible when
the head is viewed from directly in front or behind. They are variable in
size, and somewhat in position, standing either a little higher or lower;
and they sometimes occur on one ear and not on the other. They are not
confined to mankind, for I observed a case in one of the spider-monkeys
(Ateles beelzebuth) in our Zoological Gardens; and Mr. E. Ray Lankester
informs me of another case in a chimpanzee in the gardens at Hamburg. The
helix obviously consists of the extreme margin of the ear folded inwards;
and this folding appears to be in some manner connected with the whole
external ear being permanently pressed backwards. In many monkeys, which
do not stand high in the order, as baboons and some species of macacus (32.
See also some remarks, and the drawings of the ears of the Lemuroidea, in
Messrs. Murie and Mivart's excellent paper in 'Transactions of the
Zoological Society,' vol. vii. 1869, pp. 6 and 90.), the upper portion of
the ear is slightly pointed, and the margin is not at all folded inwards;
but if the margin were to be thus folded, a slight point would necessarily
project inwards towards the centre, and probably a little outwards from the
plane of the ear; and this I believe to be their origin in many cases. On
the other hand, Prof. L. Meyer, in an able paper recently published (33.
'Uber das Darwin'sche Spitzohr,' Archiv fur Path. Anat. und Phys., 1871, p.
485.), maintains that the whole case is one of mere variability; and that
the projections are not real ones, but are due to the internal cartilage on
each side of the points not having been fully developed. I am quite ready
to admit that this is the correct explanation in many instances, as in
those figured by Prof. Meyer, in which there are several minute points, or
the whole margin is sinuous. I have myself seen, through the kindness of
Dr. L. Down, the ear of a microcephalous idiot, on which there is a
projection on the outside of the helix, and not on the inward folded edge,
so that this point can have no relation to a former apex of the ear.
Nevertheless in some cases, my original view, that the points are vestiges
of the tips of formerly erect and pointed ears, still seems to me probable.
I think so from the frequency of their occurrence, and from the general
correspondence in position with that of the tip of a pointed ear. In one
case, of which a photograph has been sent me, the projection is so large,
that supposing, in accordance with Prof. Meyer's view, the ear to be made
perfect by the equal development of the cartilage throughout the whole
extent of the margin, it would have covered fully one-third of the whole
ear. Two cases have been communicated to me, one in North America, and the
other in England, in which the upper margin is not at all folded inwards,
but is pointed, so that it closely resembles the pointed ear of an ordinary
quadruped in outline. In one of these cases, which was that of a young
child, the father compared the ear with the drawing which I have given (34.
'The Expression of the Emotions,' p. 136.) of the ear of a monkey, the
Cynopithecus niger, and says that their outlines are closely similar. If,
in these two cases, the margin had been folded inwards in the normal
manner, an inward projection must have been formed. I may add that in two
other cases the outline still remains somewhat pointed, although the margin
of the upper part of the ear is normally folded inwards--in one of them,
however, very narrowly. [Fig.3. Foetus of an Orang(?). Exact copy of a
photograph, shewing the form of the ear at this early age.] The following
woodcut (No. 3) is an accurate copy of a photograph of the foetus of an
orang (kindly sent me by Dr. Nitsche), in which it may be seen how
different the pointed outline of the ear is at this period from its adult
condition, when it bears a close general resemblance to that of man. It is
evident that the folding over of the tip of such an ear, unless it changed
greatly during its further development, would give rise to a point
projecting inwards. On the whole, it still seems to me probable that the
points in question are in some cases, both in man and apes, vestiges of a
former condition.

The nictitating membrane, or third eyelid, with its accessory muscles and
other structures, is especially well developed in birds, and is of much
functional importance to them, as it can be rapidly drawn across the whole
eye-ball. It is found in some reptiles and amphibians, and in certain
fishes, as in sharks. It is fairly well developed in the two lower
divisions of the mammalian series, namely, in the monotremata and
marsupials, and in some few of the higher mammals, as in the walrus. But
in man, the quadrumana, and most other mammals, it exists, as is admitted
by all anatomists, as a mere rudiment, called the semilunar fold. (35.
Muller's 'Elements of Physiology,' Eng. translat. 1842, vol. ii. p. 1117.
Owen, 'Anatomy of Vertebrates,' vol. iii. p. 260; ibid. on the Walrus,
'Proceedings of the Zoological Society,' November 8, 1854. See also R.
Knox, 'Great Artists and Anatomists,' p. 106. This rudiment apparently is
somewhat larger in Negroes and Australians than in Europeans, see Carl
Vogt, 'Lectures on Man,' Eng. translat. p. 129.)

The sense of smell is of the highest importance to the greater number of
mammals--to some, as the ruminants, in warning them of danger; to others,
as the Carnivora, in finding their prey; to others, again, as the wild
boar, for both purposes combined. But the sense of smell is of extremely
slight service, if any, even to the dark coloured races of men, in whom it
is much more highly developed than in the white and civilised races. (36.
The account given by Humboldt of the power of smell possessed by the
natives of South America is well known, and has been confirmed by others.
M. Houzeau ('Etudes sur les Facultes Mentales,' etc., tom. i. 1872, p. 91)
asserts that he repeatedly made experiments, and proved that Negroes and
Indians could recognise persons in the dark by their odour. Dr. W. Ogle
has made some curious observations on the connection between the power of
smell and the colouring matter of the mucous membrane of the olfactory
region as well as of the skin of the body. I have, therefore, spoken in
the text of the dark-coloured races having a finer sense of smell than the
white races. See his paper, 'Medico-Chirurgical Transactions,' London,
vol. liii. 1870, p. 276.) Nevertheless it does not warn them of danger,
nor guide them to their food; nor does it prevent the Esquimaux from
sleeping in the most fetid atmosphere, nor many savages from eating
half-putrid meat. In Europeans the power differs greatly in different
individuals, as I am assured by an eminent naturalist who possesses this
sense highly developed, and who has attended to the subject. Those who
believe in the principle of gradual evolution, will not readily admit that
the sense of smell in its present state was originally acquired by man, as
he now exists. He inherits the power in an enfeebled and so far
rudimentary condition, from some early progenitor, to whom it was highly
serviceable, and by whom it was continually used. In those animals which
have this sense highly developed, such as dogs and horses, the recollection
of persons and of places is strongly associated with their odour; and we
can thus perhaps understand how it is, as Dr. Maudsley has truly remarked
(37. 'The Physiology and Pathology of Mind,' 2nd ed. 1868, p. 134.), that
the sense of smell in man "is singularly effective in recalling vividly the
ideas and images of forgotten scenes and places."

Man differs conspicuously from all the other primates in being almost
naked. But a few short straggling hairs are found over the greater part of
the body in the man, and fine down on that of the woman. The different
races differ much in hairiness; and in the individuals of the same race the
hairs are highly variable, not only in abundance, but likewise in position:
thus in some Europeans the shoulders are quite naked, whilst in others they
bear thick tufts of hair. (38. Eschricht, Uber die Richtung der Haare am
menschlichen Korper, Muller's 'Archiv fur Anat. und Phys.' 1837, s. 47. I
shall often have to refer to this very curious paper.) There can be little
doubt that the hairs thus scattered over the body are the rudiments of the
uniform hairy coat of the lower animals. This view is rendered all the
more probable, as it is known that fine, short, and pale-coloured hairs on
the limbs and other parts of the body, occasionally become developed into
"thickset, long, and rather coarse dark hairs," when abnormally nourished
near old-standing inflamed surfaces. (39. Paget, 'Lectures on Surgical
Pathology,' 1853, vol. i. p. 71.)

I am informed by Sir James Paget that often several members of a family
have a few hairs in their eyebrows much longer than the others; so that
even this slight peculiarity seems to be inherited. These hairs, too, seem
to have their representatives; for in the chimpanzee, and in certain
species of Macacus, there are scattered hairs of considerable length rising
from the naked skin above the eyes, and corresponding to our eyebrows;
similar long hairs project from the hairy covering of the superciliary
ridges in some baboons.

The fine wool-like hair, or so-called lanugo, with which the human foetus
during the sixth month is thickly covered, offers a more curious case. It
is first developed, during the fifth month, on the eyebrows and face, and
especially round the mouth, where it is much longer than that on the head.
A moustache of this kind was observed by Eschricht (40. Eschricht, ibid.
s. 40, 47.) on a female foetus; but this is not so surprising a
circumstance as it may at first appear, for the two sexes generally
resemble each other in all external characters during an early period of
growth. The direction and arrangement of the hairs on all parts of the
foetal body are the same as in the adult, but are subject to much
variability. The whole surface, including even the forehead and ears, is
thus thickly clothed; but it is a significant fact that the palms of the
hands and the soles of the feet are quite naked, like the inferior surfaces
of all four extremities in most of the lower animals. As this can hardly
be an accidental coincidence, the woolly covering of the foetus probably
represents the first permanent coat of hair in those mammals which are born
hairy. Three or four cases have been recorded of persons born with their
whole bodies and faces thickly covered with fine long hairs; and this
strange condition is strongly inherited, and is correlated with an abnormal
condition of the teeth. (41. See my 'Variation of Animals and Plants
under Domestication,' vol. ii. p. 327. Prof. Alex. Brandt has recently
sent me an additional case of a father and son, born in Russia, with these
peculiarities. I have received drawings of both from Paris.) Prof. Alex.
Brandt informs me that he has compared the hair from the face of a man thus
characterised, aged thirty-five, with the lanugo of a foetus, and finds it
quite similar in texture; therefore, as he remarks, the case may be
attributed to an arrest of development in the hair, together with its
continued growth. Many delicate children, as I have been assured by a
surgeon to a hospital for children, have their backs covered by rather long
silky hairs; and such cases probably come under the same head.

It appears as if the posterior molar or wisdom-teeth were tending to become
rudimentary in the more civilised races of man. These teeth are rather
smaller than the other molars, as is likewise the case with the
corresponding teeth in the chimpanzee and orang; and they have only two
separate fangs. They do not cut through the gums till about the
seventeenth year, and I have been assured that they are much more liable to
decay, and are earlier lost than the other teeth; but this is denied by
some eminent dentists. They are also much more liable to vary, both in
structure and in the period of their development, than the other teeth.
(42. Dr. Webb, 'Teeth in Man and the Anthropoid Apes,' as quoted by Dr. C.
Carter Blake in Anthropological Review, July 1867, p. 299.) In the
Melanian races, on the other hand, the wisdom-teeth are usually furnished
with three separate fangs, and are generally sound; they also differ from
the other molars in size, less than in the Caucasian races. (43. Owen,
'Anatomy of Vertebrates,' vol. iii. pp. 320, 321, and 325.) Prof.
Schaaffhausen accounts for this difference between the races by "the
posterior dental portion of the jaw being always shortened" in those that
are civilised (44. 'On the Primitive Form of the Skull,' Eng. translat.,
in 'Anthropological Review,' Oct. 1868, p. 426), and this shortening may, I
presume, be attributed to civilised men habitually feeding on soft, cooked
food, and thus using their jaws less. I am informed by Mr. Brace that it
is becoming quite a common practice in the United States to remove some of
the molar teeth of children, as the jaw does not grow large enough for the
perfect development of the normal number. (45. Prof. Montegazza writes to
me from Florence, that he has lately been studying the last molar teeth in
the different races of man, and has come to the same conclusion as that
given in my text, viz., that in the higher or civilised races they are on
the road towards atrophy or elimination.)

With respect to the alimentary canal, I have met with an account of only a
single rudiment, namely the vermiform appendage of the caecum. The caecum
is a branch or diverticulum of the intestine, ending in a cul-de-sac, and
is extremely long in many of the lower vegetable-feeding mammals. In the
marsupial koala it is actually more than thrice as long as the whole body.
(46. Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 416, 434, 441.) It is
sometimes produced into a long gradually-tapering point, and is sometimes
constricted in parts. It appears as if, in consequence of changed diet or
habits, the caecum had become much shortened in various animals, the
vermiform appendage being left as a rudiment of the shortened part. That
this appendage is a rudiment, we may infer from its small size, and from
the evidence which Prof. Canestrini (47. 'Annuario della Soc. d. Nat.'
Modena, 1867, p. 94.) has collected of its variability in man. It is
occasionally quite absent, or again is largely developed. The passage is
sometimes completely closed for half or two-thirds of its length, with the
terminal part consisting of a flattened solid expansion. In the orang this
appendage is long and convoluted: in man it arises from the end of the
short caecum, and is commonly from four to five inches in length, being
only about the third of an inch in diameter. Not only is it useless, but
it is sometimes the cause of death, of which fact I have lately heard two
instances: this is due to small hard bodies, such as seeds, entering the
passage, and causing inflammation. (48. M. C. Martins ("De l'Unite
Organique," in 'Revue des Deux Mondes,' June 15, 1862, p. 16) and Haeckel
('Generelle Morphologie,' B. ii. s. 278), have both remarked on the
singular fact of this rudiment sometimes causing death.)

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 'Cimetiere 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 fur 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 Theorie Darwinienne et la creation
dite independante,' 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.

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