The Descent of Man and Selection in Relation to Sex
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Charles Darwin >> The Descent of Man and Selection in Relation to Sex
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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. 'Memoires 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.
Various other anomalies in man, more or less analogous to the foregoing,
have been advanced by different authors, as cases of reversion; but these
seem not a little doubtful, for we have to descend extremely low in the
mammalian series, before we find such structures normally present. (41. A
whole series of cases is given by Isidore Geoffroy St.-Hilaire, 'Hist. des
Anomalies,' tom, iii, p. 437. A reviewer ('Journal of Anatomy and
Physiology,' 1871, p. 366) blames me much for not having discussed the
numerous cases, which have been recorded, of various parts arrested in
their development. He says that, according to my theory, "every transient
condition of an organ, during its development, is not only a means to an
end, but once was an end in itself." This does not seem to me necessarily
to hold good. Why should not variations occur during an early period of
development, having no relation to reversion; yet such variations might be
preserved and accumulated, if in any way serviceable, for instance, in
shortening and simplifying the course of development? And again, why
should not injurious abnormalities, such as atrophied or hypertrophied
parts, which have no relation to a former state of existence, occur at an
early period, as well as during maturity?)
In man, the canine teeth are perfectly efficient instruments for
mastication. But their true canine character, as Owen (42. 'Anatomy of
Vertebrates,' vol. iii. 1868, p. 323.) remarks, "is indicated by the
conical form of the crown, which terminates in an obtuse point, is convex
outward and flat or sub-concave within, at the base of which surface there
is a feeble prominence. The conical form is best expressed in the Melanian
races, especially the Australian. The canine is more deeply implanted, and
by a stronger fang than the incisors." Nevertheless, this tooth no longer
serves man as a special weapon for tearing his enemies or prey; it may,
therefore, as far as its proper function is concerned, be considered as
rudimentary. In every large collection of human skulls some may be found,
as Haeckel (43. 'Generelle Morphologie,' 1866, B. ii. s. clv.) observes,
with the canine teeth projecting considerably beyond the others in the same
manner as in the anthropomorphous apes, but in a less degree. In these
cases, open spaces between the teeth in the one jaw are left for the
reception of the canines of the opposite jaw. An inter-space of this kind
in a Kaffir skull, figured by Wagner, is surprisingly wide. (44. Carl
Vogt's 'Lectures on Man,' Eng. translat., 1864, p. 151.) Considering how
few are the ancient skulls which have been examined, compared to recent
skulls, it is an interesting fact that in at least three cases the canines
project largely; and in the Naulette jaw they are spoken of as enormous.
(45. C. Carter Blake, on a jaw from La Naulette, 'Anthropological Review,'
1867, p. 295. Schaaffhausen, ibid. 1868, p. 426.)
Of the anthropomorphous apes the males alone have their canines fully
developed; but in the female gorilla, and in a less degree in the female
orang, these teeth project considerably beyond the others; therefore the
fact, of which I have been assured, that women sometimes have considerably
projecting canines, is no serious objection to the belief that their
occasional great development in man is a case of reversion to an ape-like
progenitor. He who rejects with scorn the belief that the shape of his own
canines, and their occasional great development in other men, are due to
our early forefathers having been provided with these formidable weapons,
will probably reveal, by sneering, the line of his descent. For though he
no longer intends, nor has the power, to use these teeth as weapons, he
will unconsciously retract his "snarling muscles" (thus named by Sir C.
Bell) (46. The Anatomy of Expression, 1844, pp. 110, 131.), so as to
expose them ready for action, like a dog prepared to fight.
Many muscles are occasionally developed in man, which are proper to the
Quadrumana or other mammals. Professor Vlacovich (47. Quoted by Prof.
Canestrini in the 'Annuario della Soc. dei Naturalisti,' 1867, p. 90.)
examined forty male subjects, and found a muscle, called by him the ischio-
pubic, in nineteen of them; in three others there was a ligament which
represented this muscle; and in the remaining eighteen no trace of it. In
only two out of thirty female subjects was this muscle developed on both
sides, but in three others the rudimentary ligament was present. This
muscle, therefore, appears to be much more common in the male than in the
female sex; and on the belief in the descent of man from some lower form,
the fact is intelligible; for it has been detected in several of the lower
animals, and in all of these it serves exclusively to aid the male in the
act of reproduction.
Mr. J. Wood, in his valuable series of papers (48. These papers deserve
careful study by any one who desires to learn how frequently our muscles
vary, and in varying come to resemble those of the Quadrumana. The
following references relate to the few points touched on in my text:
'Proc. Royal Soc.' vol. xiv. 1865, pp. 379-384; vol. xv. 1866, pp. 241,
242; vol. xv. 1867, p. 544; vol. xvi. 1868, p. 524. I may here add that
Dr. Murie and Mr. St. George Mivart have shewn in their Memoir on the
Lemuroidea ('Transactions, Zoological Society,' vol. vii. 1869, p. 96), how
extraordinarily variable some of the muscles are in these animals, the
lowest members of the Primates. Gradations, also, in the muscles leading
to structures found in animals still lower in the scale, are numerous in
the Lemuroidea.), has minutely described a vast number of muscular
variations in man, which resemble normal structures in the lower animals.
The muscles which closely resemble those regularly present in our nearest
allies, the Quadrumana, are too numerous to be here even specified. In a
single male subject, having a strong bodily frame, and well-formed skull,
no less than seven muscular variations were observed, all of which plainly
represented muscles proper to various kinds of apes. This man, for
instance, had on both sides of his neck a true and powerful "levator
claviculae," such as is found in all kinds of apes, and which is said to
occur in about one out of sixty human subjects. (49. See also Prof.
Macalister in 'Proceedings, Royal Irish Academy,' vol. x. 1868, p. 124.)
Again, this man had "a special abductor of the metatarsal bone of the fifth
digit, such as Professor Huxley and Mr. Flower have shewn to exist
uniformly in the higher and lower apes." I will give only two additional
cases; the acromio-basilar muscle is found in all mammals below man, and
seems to be correlated with a quadrupedal gait, (50. Mr. Champneys in
'Journal of Anatomy and Physiology,' Nov. 1871, p. 178.) and it occurs in
about one out of sixty human subjects. In the lower extremities Mr.
Bradley (51. Ibid. May 1872, p. 421.) found an abductor ossis metatarsi
quinti in both feet of man; this muscle had not up to that time been
recorded in mankind, but is always present in the anthropomorphous apes.
The muscles of the hands and arms--parts which are so eminently
characteristic of man--are extremely liable to vary, so as to resemble the
corresponding muscles in the lower animals. (52. Prof. Macalister (ibid.
p. 121) has tabulated his observations, and finds that muscular
abnormalities are most frequent in the fore-arms, secondly, in the face,
thirdly, in the foot, etc.) Such resemblances are either perfect or
imperfect; yet in the latter case they are manifestly of a transitional
nature. Certain variations are more common in man, and others in woman,
without our being able to assign any reason. Mr. Wood, after describing
numerous variations, makes the following pregnant remark. "Notable
departures from the ordinary type of the muscular structures run in grooves
or directions, which must be taken to indicate some unknown factor, of much
importance to a comprehensive knowledge of general and scientific anatomy."
(53. The Rev. Dr. Haughton, after giving ('Proc. R. Irish Academy,' June
27, 1864, p. 715) a remarkable case of variation in the human flexor
pollicis longus, adds, "This remarkable example shews that man may
sometimes possess the arrangement of tendons of thumb and fingers
characteristic of the macaque; but whether such a case should be regarded
as a macaque passing upwards into a man, or a man passing downwards into a
macaque, or as a congenital freak of nature, I cannot undertake to say."
It is satisfactory to hear so capable an anatomist, and so embittered an
opponent of evolutionism, admitting even the possibility of either of his
first propositions. Prof. Macalister has also described ('Proceedings
Royal Irish Academy,' vol. x. 1864, p. 138) variations in the flexor
pollicis longus, remarkable from their relations to the same muscle in the
Quadrumana.)
That this unknown factor is reversion to a former state of existence may be
admitted as in the highest degree probable. (54. Since the first edition
of this book appeared, Mr. Wood has published another memoir in the
Philosophical Transactions, 1870, p. 83, on the varieties of the muscles of
the human neck, shoulder, and chest. He here shews how extremely variable
these muscles are, and how often and how closely the variations resemble
the normal muscles of the lower animals. He sums up by remarking, "It will
be enough for my purpose if I have succeeded in shewing the more important
forms which, when occurring as varieties in the human subject, tend to
exhibit in a sufficiently marked manner what may be considered as proofs
and examples of the Darwinian principle of reversion, or law of
inheritance, in this department of anatomical science.") It is quite
incredible that a man should through mere accident abnormally resemble
certain apes in no less than seven of his muscles, if there had been no
genetic connection between them. On the other hand, if man is descended
from some ape-like creature, no valid reason can be assigned why certain
muscles should not suddenly reappear after an interval of many thousand
generations, in the same manner as with horses, asses, and mules, dark-
coloured stripes suddenly reappear on the legs, and shoulders, after an
interval of hundreds, or more probably of thousands of generations.
These various cases of reversion are so closely related to those of
rudimentary organs given in the first chapter, that many of them might have
been indifferently introduced either there or here. Thus a human uterus
furnished with cornua may be said to represent, in a rudimentary condition,
the same organ in its normal state in certain mammals. Some parts which
are rudimentary in man, as the os coccyx in both sexes, and the mammae in
the male sex, are always present; whilst others, such as the supracondyloid
foramen, only occasionally appear, and therefore might have been introduced
under the head of reversion. These several reversionary structures, as
well as the strictly rudimentary ones, reveal the descent of man from some
lower form in an unmistakable manner.
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