detached portion of the scaphoid bone. In a very young chimpanzee this
bone is undoubtedly superficially indented with two transverse furrows,
but the three segments display only one uniform development of bone.
The distinct formation of _os centrale_, and its occasional appearance
in man, testify that it has an independent existence. Rosenberg
holds that this bone is not merely the _os centrale_ of mammals, but
that it is homologous with the two _ossa centralia_ of the fossil
_Enaliosauria_. It has become abortive in proportion to the reduction
in size which has taken place.[52] There would be no great difficulty
in tracing back this bone to remote types of vertebrate animals,
even as far as the _Urodela_ (Wiedersheim) of Eastern Asia.[53] The
persistence of this bone in man must be regarded as a reversion, not as
an arrest, of development.
[Illustration:
Fig. 42.--Skeleton of human hand, back view. _a_, Scaphoid bone.
_b_, Semi-lunar bone. _c_, Cuneiform bone. _d_, Pisiform bone.
_e_, Trapezium. _f_, Trapezoid bone. _g_, Os magnum. _h_,
Unciform bone. _l-l′_, Metacarpal bones. _m-m′_ and _nn_,
Phalanges.
]
On the femur of several mammals, especially in the horse, ass,
rhinoceros, and tapir, and more slightly indicated in the carnivora
and other families, there is, in addition to the two great and small
trochanters, a third, termed by Waldeyer _trochanter tertius_.[54]
Such a formation, low, blunt, and generally placed at the top of the
outer ridge of the superior bifurcation of the _linea aspera_, may
be observed in human skeletons of all races, but is either absent in
anthropoids or only faintly indicated. Virchow justly regards its
presence as theromorphic, but not as a characteristic of savage or
lower races.[55]
The human tibia displays in some instances a compression or lateral
flattening of its shaft or centre-piece, so that its transverse
diameter is quite out of proportion to its depth. Such a tibia is
termed sword-bladed, or platycnemic. Bones of this form have been
chiefly discovered in ancient deposits, as, for instance, at Gibraltar,
at Perthi-Chwareu, in Wiltshire, in Lozère, at Clichy, at Saint-Suzanne
(Sarthe), and especially at Cro-Magnon (Fig. 43), Janischwek, etc.
A similar formation has also been observed among men belonging to
cultured races, both of ancient and modern times. Virchow, for example,
discovered such bones in Transcaucasia (of the third and fourth century
of the Christian era) and at Hanai-Tepe in Troas. All the large
schools of anatomy in Europe contain specimens of tibiæ, which are to
some extent platycnemic. These are also observed in the skeletons of
primitive peoples of our time, as for example in the Negritos, Kanakas,
and other African races. While some scientific men regard these bones
as the result of an unhealthy condition, and the effect of rachitis,
others more justly ascribe them to a vigorous exercise of the muscles
in a one-sided direction. The idea expressed by Busk and others, that
the platycnemic tibiæ discovered in ancient sites of Europe have
belonged to a degraded race diffused over the whole continent, is
contradicted by the wide diffusion of this characteristic, even in
modern times. And it is doubtful whether platycnemy is absolutely
restricted to the lower races. At Janischewek, Virchow found an
extremely platycnemic tibia, exhumed from a kujawish grave of the Stone
Age, which belonged to a skull remarkable for its unusual beauty and
size, so that, taken by itself, the impression which it gave to an
anatomist was that of a highly organized race.[56]
[Illustration:
Fig. 43. Section through a platycnemic tibia from Cro-Magnon.
Fig. 44. Section through the tibia of a male gorilla.
Fig. 45. Section through the tibia of a male chimpanzee.
]
It is important to remark that platycnemy has been regarded as a
pithecoid structure, and for this reason the attempt has been made
to establish the degraded position of those peoples which are most
remarkable for platycnemy. But, as Boyd-Dawkins has already observed,
although the tibiæ of the gorilla and the chimpanzee are to some
extent platycnemic, they are much less so than the platycnemic bones
of the human skeleton. The tibia of a male gorilla in the College of
Surgeons Museum has an index width of 68·1, that of a female of 65·0,
while the index of the chimpanzee’s tibia is 61·1, which is about the
average of the tibias of Perthichwareu. It is unnecessary to indicate
the other marked distinctions between the tibiæ of men and apes; if
platycnemy is to be regarded as genetic, it must be admitted that man
has in this particular far exceeded apes.[57] Neither the gorilla,
the chimpanzee, the orang-utan, nor even the baboon possesses a tibia
which is flattened in its upper or middle part. In all these apes the
middle of the bone is more or less rounded, almost as if it had been
rounded by a turning-lathe. According to my experience, the degree of
platycnemy in anthropoids is subject to certain variations. It appears
to me to be least marked in the aged male gorilla (Fig. 41), and in
the gibbon (_Hylobates agilis_, _syndactylus_), in which latter animal
the transverse section of the tibia represents an almost equilateral
triangle. The platycnemy was more marked in an almost adult female
gorilla, still more decided in an aged male chimpanzee, which came
from the river Kiulu, and again in an aged female chimpanzee. On the
other hand, the centre of the shaft of the tibia in another aged male
chimpanzee which came from Loango, was rounded, and not platycnemic. In
the tibia of an adult orang-utan which I examined, the platycnemy was
very marked. But I agree with Boyd-Dawkins in never having met with an
anthropoid in which the platycnemy is so considerable as it is, for
instance, in the Cro-Magnon tibia, and in another found at Troy.
[Illustration:
Fig. 46.--Skeleton of the human foot, seen from above. _a_,
Astragalus. _b_, Os calcis. _c_, Scaphoid bone. _d_, _e_, _f_,
Cuneiform bones. _g_, Cuboid bone. _h_, Metatarsal bones. _ii_,
Phalanges.
]
If we give a cursory glance at the lower limbs of apes, we see that all
the same characteristics are present in their tarsus that we find in
the human tarsus. In each case there is an astragalus, an _os calcis_,
a scaphoid bone, three cuneiform bones, and a cuboid bone. There are
undoubtedly several peculiarities in which the tarsus differs from
the corresponding part of the human foot. The first metatarsal bone
is joined to the first cuneiform bone by an articular facet which
extends from the back to the sole of the foot. This joint plays a part
resembling that of the thumb of the human hand (see Figs. 20 and 46).
In Huxley’s opinion, the hinder limbs of the gorilla terminate in
a true foot, with a very movable great toe. It is undoubtedly a
prehensile foot, but in no sense a hand. It is a foot which does not
differ from the human foot in any essential characteristics, but only
in relative circumstances, in the degree of flexibility, and in the
subordinate arrangements of its parts. Huxley adds that it must not
be supposed that he wishes to undervalue differences which, however,
he does not regard as fundamental. They are important enough of
their kind, since in any case the structure of the foot is in close
correlation with the other parts of the organism. Although it cannot be
doubted that the increased division of labour in man, which relegates
the function of support entirely to the legs and feet, is a significant
advance in structure; yet, regarded as a whole from the anatomical
point of view, the points of agreement between the human foot and
that of the gorilla are much more striking and significant than their
differences.
The differences in the foot of the orang are still greater; in the very
long toes and short tarsus, the short great toe and the removal of
the heel from the ground, in the great obliquity of the joints which
connect the foot with the shank-bones, and in the absence of a long
flexor muscle to move the great toe, the orang’s foot differs still
more from that of the gorilla than the latter differs from the human
foot. In some of the lower apes the hands and feet are still further
removed from those of the gorilla than in the case of the orang. In the
American apes the thumb can no longer be opposed; in the ateles it is
reduced to a mere rudiment, covered with skin; in the sahius it is bent
forwards and provided with a curved claw like the other fingers. In all
these cases there is no doubt that the hand differs more from that of
the gorilla than the gorilla’s hand differs from that of man.[58]
Flower remarks that the chief distinction between the foot of a man
and an ape consists in the fact that the latter is transformed into a
prehensile organ. The tarsal and metatarsal bones, and the phalanges
are of the same number in both orders, and in the same relative
position, only in the foot of the ape the facet for articulation of
the first cuneiform bone with the great toe is saddle-shaped, and
obliquely directed towards the inner or tibial side of the foot. Thus,
the great toe is separated from the others, and so placed, that when
it is bent, it is directed downwards towards the sole, and is opposed
to the other toes, much more opposed to them than is the case with the
thumb of the human hand.[59] Owen also speaks of the characteristic
transformation of the great toe of an ape’s foot into a thumb, opposed
to the other toes, and adapted for grasping.[60]
[Illustration: Fig. 47.--Coaita (_Ateles paniscus_).]
K. E. von Bär does not agree with Huxley in considering that there is
less difference between man and the gorilla than that which exists
between different species of apes. “There are,” Von Bär remarks,
“differences of various kinds among apes. In some the thumb is only
a stump; in others, as in the orang-utan, the fingers of the hinder
extremities are so long and curved that they cannot be extended on flat
ground; in many of the smaller apes this member is still more like a
hand than in the larger species, and the fingers can be easily spread
out on the ground. In this case the foot is of a much blunter form, and
is more flexible, so that the sole, which is properly turned inward,
can lie flat on the ground. The heavier the body of the animal, the
more sharply cut the structure of the foot must be, so that it does
not admit of the free movements which are possible in the hand. But
all these are only modifications of a climbing foot, or prehensile
member--that is, of a hand, not modifications of a foot resting firmly
on the ground and supporting the whole weight of the body.
“It must not be forgotten that the structure of the skeleton is subject
to mechanical laws, which may be traced through the whole series of
the animal world. This is readily apparent when we turn to the human
structure.
“The human foot rests for the greater part of its length on the
ground, that is to say, with the heel and centre of the foot, which
form together a firm arch. The tarsus consists of the astragalus, and
also of the _os calcis_, which in man form a very prominent part,
taking a backward and downward direction, and of five other bones.
The metatarsus consists of five bones, on which the five toes are
inserted. In man these metatarsal bones are considerably longer than
the separate phalanges. Thus, the arch on which man is supported in
an erect position extends from the heel to the extremities of the
metatarsal bones. The several bones are slightly movable, but they are
so firmly connected that they can diverge but little from each other,
unless muscular power is exerted. In order to press the toes upon the
ground, it is again necessary to exert the muscles. The arched instep
has this advantage, that the foot can take a better hold of the slight
inequalities of the ground. In a profile view of the skeleton of a
human foot, the shortness of the toes, in comparison with the length of
the arched instep, is very apparent. In any natural position, even when
man is not walking or standing, the sole of the foot is not turned
inwards, but downwards.... The toes of the gorilla take the form of
a hand, since the great toe stands separate like a thumb, while the
other toes are turned outwards. In the gorilla the tarsus is short,
and the heel is bent inwards. The several bones of the human foot are
undoubtedly present in the hind hand of a gorilla, but the organ is
changed into a prehensile organ or hand. The conditions are the same as
in the parts of the mouth in insects which in some cases form movable
mandibles, while in others they are attenuated into a proboscis. When
it is asserted that apes are not quadrumanous, it is as if we were to
say that flies have no proboscis, but attenuated mandibles.”[61]
All apes, including anthropoids, occasionally make use of their hinder
extremities in order to snatch at objects. They also grasp with them in
climbing. On such occasions, when they wish to secure the fruit they
have seized from the voracity of their fellows, they take it between
the toes of one hinder extremity, in order to be able to get away more
quickly by means of the other, and by the use of both hands.
From what we have said, it will be seen how difficult it is to
reconcile the views of different observers with respect to the fitting
term to be given to the hinder extremities of apes. Against those who
uphold the designation of _hind hands_ we must oppose the anatomical
structure, and also the fact that a true hand ought to possess the
power of rotation in a degree which exists in the fore, but not in the
hind, extremities of apes. On this account I have already adopted, as
more suitable and equally distinctive, the term of _prehensile foot_
for this member.[62] I agree with Haeckel in rejecting the common
designation of apes as four-handed or quadrumanous.
The bands or ligaments which connect the different parts of the
anthropoid skeleton together, and convert the detached elements into
a movable machinery, do not on the whole differ much from the same
structure in man. A detailed account of these ligaments would, for
several reasons, be out of place in this work, and I shall only mention
a few special and more interesting distinctions. Such, for example,
is the uncommon strength of the _ligamentum nuchæ_ in the gorilla,
which is quite in harmony with the great development of the spinous
processes of the upper cervical vertebræ, and with the flattening of
the squamous occipital portion. Since the sacral vertebræ are deeply
inserted between the high ilia, the ilio-lumbar ligaments (_ligamenta
iliolumbalia_) and the sacro-iliac ligaments (_ligamenta iliosacralia_)
are of considerable size. In agreement with the projection in a
downward direction of the high, narrow ischial bones, the sacro-sciatic
ligaments which extend between these and the sacrum are very long
in the chimpanzee. Although in this case the ischial spine is only
represented by a roughness of the bone, yet there is on either side
between this and the sacrum a powerful lesser sacro-sciatic ligament
(_ligamentum spinoso-sacrum_).
The well-known anatomist, J. F. Meckel, has asserted that the
depression in the head of the femur (_fovea capitis_), which serves for
the insertion of the round ligament (_ligamentum teres_), is absent in
the chimpanzee and orang, and he adds that it is also absent in the
gibbon. In a skeleton of a young chimpanzee which had not shed its
milk-teeth, and of which the ligaments were also preserved, Welcker
found a fully developed round ligament inserted almost in the centre of
the head of the femur. This agrees in every particular with the same
formation in man. On the other hand, no trace of a round ligament was
to be found in the hip-joint of a young orang-utan. The cartilaginous
envelope of the head of the femur was smooth throughout, without any
indication of a place for inserting the ligament. Welcker again found
no such depression in the femur of an aged male orang-utan, nor was
there any trace of it in another aged male orang, designated as _Simia
Morio_. Welcker believes that he has established the fact that the
round ligament is wanting in the orang-utan, but that it is present in
the gorilla, chimpanzee, and gibbon. The same naturalist remarks that,
although we may certainly assume that the round ligament is absent
wherever there is no depression in the head of the femur, yet the
existence of such a depression in the acetabulum (_fovea acetabuli_)
is not enough to prove that a round ligament was inserted in it. The
innominate bones of an adult orang-utan were examined by Welcker, and
displayed a small, but well-defined depression, as if destined for
a receptacle for this ligament,[63] running from the cotyloid notch
down to the bottom of the acetabulum, between the two horns of the
semilunar-shaped articular cartilage.
In a subsequent paper, Welcker states that the absence of the round
ligament in the orang-utan, and its presence in the chimpanzee, had
been previously established by Camper and Owen.[64] In three specimens
of orangs which he had obtained immediately after death, Owen found
that the round ligament was imperfectly developed on both sides. The
chimpanzee differs from the orang in possessing a depression on the
head of the femur. In the gorilla, as Owen observes, this depression
has almost the same depth and relative position as in man. At Welcker’s
request, Professor Dippel ascertained the presence of the depression
in the femur of a gorilla skeleton which is preserved in the natural
history collection at Darmstadt. St. George Mivart saw the skeleton
of an orang in which the femur was marked with a slight but plainly
indicated depression, just where the round ligament is usually
attached. Welcker thinks it probable that in some specimens of the
gorilla the round ligament is only slightly developed, and that in
others it is altogether wanting. On several femurs of gorillas, this
naturalist observed only doubtful traces of the depression in question.
Duvernoy found the round ligament fully developed in the gorilla and
chimpanzee. Vrolik failed to find it in the orang-utan, but ascertained
its presence in the chimpanzee. Gratiolet and Alix saw that it was
fully developed in _Troglodytes Aubryi_.
In addition to these somewhat conflicting assertions, I have myself
observed, in the gorilla innominate and femur bones examined by me,
more or less distinct indications of the depression which receives
the round ligament. The ligament itself has been preserved with the
body of a gorilla. The same remark applies to the skeletons and
bodies of chimpanzees. In the case of the skeleton of an orang,
slight indications of a depression were observed on the head of the
left femur, and these indications were absent in the femurs of other
specimens. In a large orang-utan which died in the Berlin Aquarium,
only short, filamentous tufts of streaky fibres were apparent in the
right acetabulum, and these were intermingled singly or in groups
with the cartilaginous cells, somewhat resembling the cartilaginous
corpuscles of the synovial membrane. From these facts we may conclude
that the round ligament is generally but not invariably present in
the gorilla and chimpanzee, and that it is altogether absent in the
orang-utan. In the gibbon it is present in the majority of cases.
I have myself observed it in _Hylobates agilis_, _leuciscus_, and
_syndactylus_. Owen asserts that the unsteady gait of the orang is
partly due to the absence of this ligament, but the truth of this
surmise is rendered doubtful by the fact that the ligament is not
unfrequently absent in other anthropoids. Moreover, the gait of all
these arboreal and climbing animals is extremely ungainly.
The muscular system of anthropoid apes is very interesting. I must
necessarily refrain from giving a detailed account of it, and will only
mention some points in connection with this organic system, and their
relation to corresponding points in the muscular system of man. I rely
partly on the researches of others, and partly on my own. The amount of
material which has been collected up to this time is, unfortunately,
too scanty to enable us to draw satisfactory conclusions in all cases.
We are often unable to decide whether the conditions presented to us
in the case of anthropoids are normal or exceptional. Nor are the
statistics of muscular variations in the human subject by any means
firmly established. My own labours in this direction are not yet
concluded. The assertions on the subject which have been published to
the world and accepted as authoritative have already been shown to be
to some extent untrustworthy. Even the little which I am now able to
produce may not altogether stand the test of subsequent research. Brühl
justly remarks that in no department of anatomy more than in that which
treats of the muscles, is it more essential that we should not decide
whether a form is normal or exceptional until it has been repeatedly
examined.[65]
[Illustration:
Fig. 48.--Muscles of the head and face of a European. 1,
1′, Occipito-frontalis. 2, 3, Orbicularis palpebrarum. 4,
Pyramidalis nasi. 5, Levator labii superioris alæque nasi. 6,
Compressor naris. 7, Levator labii superioris. 7′, Zygomaticus
minor. 8, Levator anguli oris. 8′, Zygomaticus major. 9,
Orbicularis oris. 9′, Levator menti. 9″, Depressor labii
inferioris. 10, Depressor anguli oris. 11, Masseter. 12,
13, Risorius and the buccinator by which it is covered. 15,
Trapezius. 16, Attrahens. 17, 19, Attollens. 20, Retrahens
aurem. 21, Sterno-mastoid. 22, Splenius. A. Tendinous
aponeurosis. C. Malar bone (the parotis is removed). F. Skin of
neck.
]
The cranial muscles of anthropoids are formed like those of men,
except in a few unimportant particulars (comp. Figs. 48 and 50). I have
not observed in anthropoids the muscular fibres which in man branch
out from the orbicular muscle of the eye, and overlap the cheeks and
temples, and which are considerably developed in the head of a Monjalo
negro which was dissected by me (Fig. 49, ~3~, ~3′~). In apes that
portion of the orbicular muscle which covers the supra-orbital ridge
is very marked. There is generally a considerable layer of muscle on
the nose and upper lip. I have dissected it in detail in anthropoid and
other apes, including those of America; _i.e._ the zygomatic muscles,
the levator labii superioris, and the levator labii superioris alæque
nasi. This has also been done by Duvernoy, Alix, and Gratiolet, in the
case of anthropoids dissected by them, as well as by Macalister and
Bischoff.
[Illustration:
Fig. 49.--Head-muscles of a Monjalese negro. 1, 2,
Occipito-frontalis. 3, 3′, Orbicularis palpebrarum. 4,
Pyramidalis nasi. 4′, Levator labii superioris. 6, Levator
labii superioris alæque nasi. 6′, Compressor naris. 7′, Levator
anguli oris. 8, 8′, Zygomatici major et minor. 9, Orbicularis
oris. 9′, Levator menti. 9″, Depressor labii inferioris. 9‴,
Depressor anguli oris. 11, Masseter. 13, Buccinator. 14,
Platysma. 15, Trapezius. 17, 18, Attollens and attrahens
aurem. 19, Embedded temporal muscle. 20, Retrahens aurem. 21,
Sterno-mastoid. 22, Deeply set muscles of neck. A, Tendinous
aponeurosis. C, Zygoma. E, Parotis. *, Stensonian duct.
]
Bischoff was only able to identify a wide zygomatic muscle in the
orang with the small zygomatic in man. In the orang, the gibbon, and
the baboon, as well as in _Innus sinicus_ and _Ateles_, I myself was
quite able to trace a division into a large and small zygomatic. In the
gorilla dissected by me the levator labii superioris alæque nasi was
very wide (Fig. 50, ~6~). In the case of a gorilla, Ehlers dissected
the small zygomatic muscle, together with the levator labii superioris
alæque nasi, in the manner introduced by Henle as a single square
muscle of the upper lip (_Musculus quadratus labii superioris_). In the
gorilla I observed a levator alæque nasi, together with the already
mentioned levator labii superioris; but I failed to find any separate
levator labii superioris. The very wide cartilage of the nose is
occupied by a considerable amount of muscular tissue. All these muscles
are present in the orang, but they are of small size and separated
into detached bundles. The pyramidalis nasi may be traced in every
instance, especially in the gorilla (Fig. 50, ~4~) and in the orang. It
is not so strongly developed in the chimpanzee and gibbon, but is not
absent in these apes, nor in those which are not anthropoid, such as
the baboon, and ateles, or climbing ape.
[Illustration:
Fig. 50.--Head-muscles of gorilla presented in Fig. 3. 1,
2, Occipito-frontalis. 3, 3′, Orbicularis palpebrarum. 4,
Pyramidalis nasi. 5, Levator alæ nasi. 6, Levator labii
superioris alæque nasi. 7, Zygomaticus minor. 7′, Levator
anguli oris. 8, Zygomaticus major. 9, 9′, Orbicularis oris.
10, Risorius. 11, 16, Masseter. 1′, Buccinator. 12, Depressor
anguli oris. 13, Buccinator. 14, Platysma. 15, Trapezius. 17,
Temporal. 18, 19, 20, Attrahens, attollens, and retrahens
aurem. 21, Lesser muscle of helix. A, Tendinous aponeurosis. B,
Cartilage of nostril. C, Zygoma. D, External ear. *, Stensonian
duct.
]
I myself follow the original division of the muscles into those which
belong to the nostril and upper lip, in accordance with the principles
of Duchenne, Darwin, Gamba,[66] and others, and I do so the more
readily, since it is impossible not to perceive the manifold and lively
mimetic action which takes place in this particular region of an ape’s
head. The distinct action of the levator labii superioris alæque nasi,
the dilation of the nostrils, the function of a strongly developed
levator anguli oris, are especially characteristic of the gorilla; but
they are also perceptible in the chimpanzee and gibbon. The orang’s
face is the least mobile. I observed that in the gorilla the risorius
was very long, branching slightly in the fore-part of the corner of the
mouth, and behind into three distinct wide bundles. The lowest bundle
covered the platysma myoides, but could not be regarded as part of
the latter. In one chimpanzee I found that the risorius was slightly
developed, and in other animals of that species I failed to trace it
at all. Alix and Gratiolet represent the Aubry chimpanzee (Plate ix.
Fig. 1, ~15~) with the risorius strongly developed. I have not observed
this formation either in the orang or the gibbon, but it was apparent
in one of the ateles (_Ateles leucophthalmos_). In this case the muscle
covered the platysma myoides and Stenson’s duct, _i.e._ the duct
leading out of the parotid gland (Fig. 50, *).
For some time I was disposed to regard the risorius of this ape as only
a radiation of the platysma myoides, but my opinion upon this point is
again shaken.
In the gorilla a faint depressor anguli oris and an equally faint
depressor labii inferioris may be observed, the latter partly covered
by the large and predominant orbicularis oris (Fig. 50). In the
chimpanzee the two depressors are plainly apparent, and in the gibbon
the one first named was at any rate developed. The platysma myoides,
the depressors just mentioned, and the crescent-shaped orbiculares
are in this animal in close connection with each other. Froriep’s
suggestion becomes ever more probable, that these muscles of the lower
lip owe their origin to the intersection of the opposite portions of
the skin-muscles of the neck which overlap the face. The buccinator
muscle in anthropoids resembles on the whole that of man, and in
both cases is pierced by Stenson’s duct (Fig. 50). The form of the
masseter muscle is common to both (see Fig. 50, ~11, 16~). In the
external ear of anthropoids there is an attrahens, attollens, retrahens
(Fig. 50). Compared with that of a white man, and still more with
that of a negro (see Figs. 48, ~19~, and 49, ~17~), the attollens
is only slightly developed. The muscles attached to the cartilages
of the ear are extremely scanty or partially wanting, which is also
sometimes the case with man. The muscles of the helix are most strongly
marked in the gorilla (see, for example, Fig. 50, ~21~). Tiedemann,
Bischoff’s brother-in-law, carefully observed two living chimpanzees
in Philadelphia for six months without detecting any movement of the
ears. My own observation confirms his assertion and the remarks of
Darwin, which I have already quoted, to the effect that anthropoids are
incapable of moving their ears. I know of no individual exceptions.
This is the more remarkable since some men have retained the power of
voluntarily moving their ears, and the same power is also found in some
species of apes, such as the sea-cats, baboons, macacas, and magots.
It will not here be out of place to say something of the
characteristics, previously mentioned, of the physiognomical expression
of anthropoid apes. Thus, for example, when the gorilla is agitated, he
can move the skin of his head and bristle the hair which covers this
region. The chimpanzee can also move the skin of the head, but with no
very apparent bristling of the hair. The large male orang, which was in
the Berlin Aquarium in 1876, bristled his hair and the skin of his head
when he was much enraged. It is known that in some instances man also
possesses this power.
I have already spoken of the expression of the eyes of these animals. I
will only add that when anthropoids of every species are in great pain
or seriously ill, the expression of their eyes is often most affecting.
The forehead of these animals is frequently marked by transverse
furrows, and especially, as Darwin justly observes, when they raise
their eyebrows. The same great observer considers that the countenances
of anthropoids are, in comparison with those of men, generally
inexpressive, and indeed, chiefly in consequence of the fact that they
do not wrinkle the forehead when they are excited. The wrinkling of
the forehead, which is one of the most significant forms of expression
in man, is due to the action of the corrugatores supercilii, by which
the eyebrows are drawn down and closer to each other, so as to form
vertical folds on the forehead. It has been asserted that the orang
and chimpanzee possess these muscles, but they seem to be rarely
exercised--at any rate, to any remarkable extent.[67] When Darwin
brought a chimpanzee out of his dark chamber into bright sunshine, he
only once observed a slight wrinkling of the forehead. When the same
observer tickled the nose of a chimpanzee with a straw, its face was
slightly wrinkled, and faint vertical furrows appeared between the
eyebrows.[68] Darwin never observed any wrinkling of the forehead in
an orang. I myself have observed a contraction of that region of the
brows which is covered with bristly hairs, and a wrinkling of the skin
which covers the bridge of the nose in the gorilla and the chimpanzee,
and have illustrated this expression by a drawing.
Darwin goes on to say that when a young chimpanzee is tickled, to
which, as in the case of children, their armpits are peculiarly
sensitive, he generally utters a chuckling or laughing sound, although
sometimes the laugh is silent. The corners of the mouth are then drawn
back, and this sometimes causes the eyelids to be slightly wrinkled.
This wrinkling, which is so characteristic of the human laugh, is still
more apparent in some of the other apes. In the chimpanzee the teeth of
the upper jaw are not exposed when he utters this laughing sound, and
in this respect he differs from man. Darwin further observes that when
the tickled young orang ceases to laugh, an expression passes over his
face, which, according to Wallace, may be called a smile. Darwin has
observed something similar in the chimpanzee.[69]
My own observation confirms what has been said of the chuckling
of a tickled chimpanzee. When Dr. Hermes, the director of the
Berlin Aquarium, played with the chimpanzee which was kept in that
establishment, a contortion of the corner of the mouth, resembling
a somewhat sardonic smile, at once appeared. No specimen displayed
this smile with so much effect as the lively Augustus, who delighted
visitors by his inexhaustible humour in 1879. The gorilla, of which
an illustration is given in Fig. 3, also drew down the corner of his
mouth when he was pleased, by means of the muscular system which we
have just described.
When the gorilla is provoked, he displays both rows of teeth, and opens
his mouth to utter sounds of fury, while making ready to fight. It is
well known that anthropoids are able to pout and project their lips;
and Darwin says that they do this, not only when they are slightly
teased, and are sullen or disappointed, but also when anything occurs
to make them uneasy.
I have often observed in chimpanzees a slight wrinkling of the region
of the nasal cartilage, and even a vibration in a lateral and upward
direction. In any case, the muscles which we have described as acting
on the nose and upper lip are exercised.
The platysma myoides, which extends in man from the lower row of teeth
to just below the clavicle, occupies about the same area in the gibbon
and in other apes (Fig. 50). In the chimpanzee, however, this muscle
extends as high as the zygomatic arch, or even higher. In the gorilla
also I observed that this part extends comparatively high on the face.
In chimpanzees, orangs, and gibbons the upper fibres of this muscle
seem to form the risorius. In one case the platysma myoides sent forth
a fasciculus, about 18 mm. in width, to the beginning of the lower
temporal ridges. In the gorilla I saw that the uppermost fibres of the
platysma myoides were partly covered by the risorius (Fig. 50, ~10~).
From the corresponding muscle in the orang the lower fibres tend far
backward, and are in connection with the deltoid muscle covering a
segment of the capsular ligament. This muscle wrinkles the skin of
the neck, and helps to draw down the lower jaw. In cases in which it
extends far in an upward direction, as in those we have cited, it
affects the lateral extension of the middle and lower skin on the faces
of these animals, as well as the grinning contortion of the corner of
the mouth. It may also have to do with the grumbling sound issuing from
the throat-pouch, which is uttered by the animal when agitated, as he
rapidly opens and closes his mouth.
The strong sterno-cleido-mastoid muscle found in these animals, and
especially in the orang and gibbon, can be divided without difficulty
into a sternal and clavicular portion. The two portions diverge from
each other in a downward direction. As Bischoff justly states, a
muscle not hitherto observed in man may be traced in all four species
of anthropoids, a muscle which extends from the external part of the
clavicle to the transverse process of the first cervical vertebra.
Bischoff has called it the musculus omocervicalis. It is found in other
apes, although the site of its origin varies, sometimes occurring on
the spine of the scapula. Our Munich anatomist differs from Huxley in
regarding this muscle as “a brilliant proof of the relation of all apes
with each other.” I give this assertion without further comment.
The muscles which extend between the head, sternum, and clavicle,
together with the muscles of the acromion process of the scapula, make
an external covering to the throat-pouch, which I shall describe
presently. The pectoralis major of the gorilla, as well as that of man,
divides into two portions, one attached to the clavicle, the other
to the cartilages of the true ribs. The former is divided from the
deltoid by a wide interval, filled with connective tissue and fat. But
both portions of the pectoralis major are divided by a tolerably wide
space, into which, in Bischoff’s opinion, the throat-pouch is inserted.
This, however, I do not believe, since that organ would be compressed
and strangulated between the two portions of muscle whenever they were
exercised. It may, however, be supposed that room for an enlargement
of the throat-pouch when the animal is bellowing is afforded by the
existence of these spaces. Bischoff is right in the assertion that
the clavicular portion of the pectoralis major is wanting in the
orang-utan. The upper part of this muscle springs directly from the
sternum. The lower sternal ribs give origin to the pectoralis minor.
The chimpanzee and gibbon display clearly in this muscle the separation
we have mentioned into a clavicular and a sternal portion.
The structure of the pectoralis minor in these apes is full of
interest. In the gorilla it divides into an upper portion of firmer
tissue, less easily separable into digitations, which arises from the
third to the fifth ribs, and a lower portion, separable into three
digitations, of which the upper segment laps considerably over the
lower segment of the upper portion. In the chimpanzee an upper portion
of less firm texture extends from the second to the fourth, and a
lower with three digitations from the fourth to the seventh ribs. This
second lower portion is sometimes absent. I have seen the upper portion
attached to the coracoid process of the scapula, and the lower portion
to the ridge of the greater tuberosity of the humerus. In the orang
an upper portion, separable into three digitations, extends from the
second to the fifth ribs, and is attached to the coracoid process. A
lower portion, also separable into three digitations, extends from the
fifth to the seventh ribs, and is also attached either to the greater
tuberosity of the humerus or to its edge; this latter portion projects
below over the pectoralis major. In the gibbon (_Hylobates albimanus_),
the upper portion starts from the second, the lower from the third to
the fifth ribs. It may here be remarked that the pectoralis minor is in
man also sometimes separable into digitations, which may be connected
both with the coracoid process and with the capsular ligament of the
shoulder-joint. In anthropoids the tendon of insertion of this muscle
is remarkably slender.
According to Duvernoy, in the gorilla a fibrous, hood-like fascia
covers the whole region of the occiput and neck. In adult males
this fascia is 20 mm. in thickness. In a female dissected by me
the rudiments of a similar hood-like cervical fascia were present.
Duvernoy is justified in supposing that this is not yet developed in
the young gorilla, and that a layer of connective tissue and fat is
substituted for it. In a young gorilla I saw the trapezius divided
into distinct bundles of flesh by layers of fat (Fig. 50, ~15~). The
fascia corresponds to the great development of the trapezius, and the
same characteristic development exists in other anthropoids. The adult
male gorilla displays a powerful _ligamentum nuchæ_ in connection
with the long spinous processes of the cervical vertebræ, as well as
powerful inter-spinales muscles, spinales colli, and semi-spinales
colli and dorsi. The great development of the spinous processes of
the dorsal vertebræ of gorillas (Fig. 17), and also chimpanzees and
orangs, involve the development of powerful semi-spinales, as well
as of strong, fourfold spinales and inter-spinales muscles. The
whole of the fleshy formation of the neck of an adult male gorilla
which is covered by the trapezius is very voluminous, and especially
the splenius capitis and colli, the long cervical muscle (_Musculus
longissimus cervicis_), and the long head-muscle (_Musculus longissimus
capitis_), which have also been regarded by me as parts of the long
spinal extensor, and finally the oblique and vertical muscles at the
back of the head. With Chappuy, I am disposed to regard the latter as
modifications of the spinales and inter-spinales.
The levator anguli scapulæ is divided in anthropoids as in man. The
subclavius is slender, except in the gorilla, and in the latter animal
it sends a tendon obliquely to the coracoid process.
In all anthropoids the deltoid is strongly developed. In the gorilla
it projects forwards and outwards in order to attach itself to the
humerus, almost in its centre. Here it is separated from the brachialis
anticus in a manner with which we are only imperfectly acquainted. It
extends nearly as far in the gibbon and orang, while in the chimpanzee
its attachment is higher up. Bischoff observes, and it was previously
suggested by Vrolik, that in the chimpanzee the coraco-brachialis
muscle possesses at its origin a moderately large second portion, which
tends downwards over the lesser tuberosity of the humerus, and adheres
to its edge. But I have seen both portions of the muscle in question
attached to the coracoid process of the scapula in apes of this
species. In the gorilla, orang, and gibbon the position of this muscle
corresponds to that in man.
Chapman and Bischoff speak of a muscle common to all apes which starts
from the tendinous attachment of the _latissimus dorsi_ on the edge
of the lesser tuberosity of the humerus, and tends downwards on the
inner side of the humerus, and to this muscle they give the name
_latissimo-condyloideus_. Bischoff goes on to say that this muscle goes
in some cases into the fascia which covers the biceps; and in others,
as in the baboon, it is attached to the inner inter-muscular septum and
to the internal condyle of the humerus. In the gibbon it only extends
as far as the centre of the humerus, but in the orang it reaches to the
condyle, where it is pierced by the ulnar nerve. Bischoff adds that
this formation is wanting in man.
This structure is indeed remarkable in anthropoids. The muscle starts
in a lateral direction from the insertion point of the _latissimus
dorsi_. In the gorilla alone I observed that it started from the
coracoid process of the scapula, together with the two portions
of the pectoralis minor; it was connected for a space with the
coraco-brachialis, and finally it was attached, in the upper part of
the lower third of the humerus, to the inter-muscular septum which
is found between the brachialis anticus and the triceps. In the
chimpanzee, on the other hand, it has its origin in the _latissimus
dorsi_, and divides into an anterior and posterior portion; the former
is attached to the inner condyle of the humerus, while the latter is
connected either with the middle or inner head of the triceps. In the
orang the same division of this muscle may occur. In one of these
animals I observed an anterior portion, very thin and semi-membranous,
attached by an extremely slight tendon to the coracoid process of the
shoulder-blade, while the hind portion issued from the _latissimus
dorsi_. They were both in connection with the triceps and brachialis
anticus. In other instances the muscle consisted only of the posterior
portion, issuing from the _latissimus dorsi_. In the white-handed
gibbon, the muscle issued from the region in which the tendons of the
_latissimus dorsi_ and of the teres major are united, and was inserted
into the fascia which is found between the bicipital and the brachialis
anterior. This attachment may also occur in the centre of the shaft of
the humerus. Chapman and Chudzinsky have observed anomalous instances
of this formation in coloured races.[70]
It is well known that in man the biceps is inserted into the tuberosity
of the radius by means of a flattened round tendon. This tendon,
however, opposite the bend of the elbow, gives off a broad expansion,
which passes into the fascia of the forearm, and is termed _Aponeurosis
bicipitis_. In the gorilla this aponeurosis is carried on as strong
fibrous bundles of the fascia of the forearm into the palmar fascia. In
the gibbon the short head of the muscle does not always start from the
lesser tuberosity of the humerus, nor from the tendon of the pectoralis
major (Huxley), but sometimes from the edge of the lesser tuberosity,
which is here connected with the _latissimus dorsi_, as well as with
the sub-scapularis, the brachialis anticus, which is more to the side,
and with the triceps. In the gibbon, as Bischoff justly observes, the
supinator longus only reaches as far as the centre of the radius,
instead of extending to the styloid process of that bone, as it does in
other anthropoids, and in man.
The palmaris longus is wanting in the gorilla, but not in other
anthropoids. The long flexor muscles of the fingers and the lumbricales
resemble those of man (Figs. 51, 52). The flexor longus pollicis is
absent in the gorilla. Duvernoy considers that it is replaced by a
tendon of the long flexor of the fore-finger, but I have been unable
to verify the existence of this tendon. The same muscle is also absent
in the chimpanzee and the orang, but it may be traced in _Hylobates
albimanus_. Chapman states that in the gorilla the pronator radii teres
only sends forth one head,[71] but I have found it to be bicipital
in animals of this species. The lower or hinder head issues, as in
man, from the coronoid process of the ulna. Both in the gorilla and in
the chimpanzee it extends far in a downwards direction on the radius
(Fig. 52). The flexor carpi radialis starts in the chimpanzee with one
head from the inner condyle of the humerus, and with the other from
the radius. Bischoff describes the structure of the long abductor of
the thumb in the orang, the baboon, the _pithecia_, and the _hapale_
as resembling that of man. But in the gorilla, the chimpanzee, and
the macaca the tendon divides into two parts. Nor does one tendon
belong, as in man, to a short extensor of the thumb, but the latter is
wholly absent, and the division of the tendon only implies a continued
division of the attachment to the trapezium, as well as to the
metacarpal bones of the thumb. This division of the tendon also occurs
in the gorilla, which likewise possesses a short extensor of the thumb.
In this point, again, apes display a greater likeness to one another
than to man.
[Illustration:
Fig. 51.--Palmar muscles of man. _a_, Ligaments of wrist,
especially the anterior ligament. _c_, _c′_, Sheathing
ligaments. _d_, _e_, _f_, Oblique fibres of the ligaments
of the sheath of the flexor tendons. 1, 2, Tendons flexor
sublimis, and of the flexor profundus muscles of the fingers.
3, The reciprocal perforation of these tendons. 4, Continuation
of the tendons of the flexor profundus of the fingers. 5,
Tendon of the flexor longus pollicis. 6, Abductor pollicis. 7,
8, 9, Flexor brevis, adductor, and opponens pollicis. 10, 11,
12, Flexor brevis, abductor, and opponens minimi digiti. 13,
Lumbricales. 14, First dorsal inter-osseous muscle.
]
According to my own researches, the long abductor of the thumb in
anthropoids forms a muscle not more considerable than one in proximity
with it, of which the origin and more central direction recall the
short extensor of the human thumb. In all four species I found that
the abductor had two tendons, and was attached to the trapezium.
The muscle in its vicinity is inserted above the base of the first
metacarpal bone. I have not been able to discover an extra extensor of
the thumb in the gorilla. The question now arises what we should think
of the second muscle, which is found in these animals in the vicinity
of the abductor. In my opinion, it may be confidently accepted as a
short extensor of the thumb, since it always effects an extension of
the metacarpal bone of that member, and in this act of extension it
is supported by the long extensor which acts upon the phalanges. It
must be remembered that the comparatively short thumbs of anthropoids
have not to be employed in so many different ways as the human thumb,
and that we cannot therefore be surprised that the development of the
short extensor is less complete. A special extensor muscle of the index
finger is either altogether absent in the gorilla or very slightly
developed, while it is very apparent in _Hylobates albimanus_ (~6~,
Fig. 53). In the chimpanzee this muscle sends a tendon to the middle
finger. In the orang there is one extensor common to the four fingers.
In the gibbon’s hand, this, as well as the other extensor and flexor
muscles, is remarkable for its excessive slenderness. The manifold
connections of the extensor tendons with each other are an interesting
peculiarity (Fig. 53).
[Illustration:
Fig. 52.--Palmar muscles of gorilla. _a_, Anterior ligament.
_b_, Remains of the skin of palm, here covered with a very
sinewy skin. _c_, _f_, Oblique fibres of the ligaments of the
sheath of the flexor tendons of fingers. 1, 2, Flexor tendons.
3, Spaces between the heads of the flexor brevis pollicis,
whence in man the tendon of the flexor longus pollicis issues
(comp. Fig. 51, ~5~). 4, 3, 3′, 5, Abductor, flexor brevis,
abductor pollicis. 6, 7, 8, Opponens, flexor brevis, abductor,
minimi digiti. 9, Dumbricales. 10, Supinator longus. 12, Flexor
sublimis digitorum. 13, Flexor minimi digiti. 14, Flexor carpi
ulnaris.
]
In the chimpanzee I observed a superficial flexor, common to the
fingers, and enlarged in the region of the third and little fingers.
A superficial flexor, belonging to the index finger, started from the
inner condyle of the humerus, and from the back of the inter-muscular
septum. The deep finger-flexor was attached to the four fingers. In
the orang the first of these flexors forms a two-tendoned belly for
the index finger, as well as one for the other three fingers. The deep
flexor only displayed two bellies. In the gibbon, on the other hand,
the superficial flexor displays four bellies.
[Illustration:
Fig. 53.--Muscular system of the back of a gibbon’s hand. 1, The
extensor carpi radialis longior and brevior. 2, Abductor longus
pollicis. 3, Extensor primi internodii pollicis. 4, Extensor
secundi internodii pollicis. 5, Extensor communis digitorum. 6,
Extensor indicis. 7, Extensor minimi digiti. 8, Extensor carpi
ulnaris. 9, First dorsal inter-osseous muscle. 10, Continuation
of the same to index finger. 11, 12, The other inter-osseous
muscles of this region. A, The posterior annular ligament.
]
In the carpus of the chimpanzee there is, so far at least as my
experience goes, a so-called sesamoid bone. It is in this instance in
connection with the scaphoid and trapezium bones, just where the fibres
of the anterior and posterior ligaments of the wrist pass into each
other. In the chimpanzee the tendon of the long abductor muscle of the
thumb sends some fibres into this sesamoid bone, while the other fibres
of the tendon of this muscle, which divides into several strips, are
inserted in the trapezium bone, and a few also in the base of the first
metacarpal bone.
The short flexor muscle of the thumb, of which Bischoff has denied the
existence, is certainly present in these animals. In the chimpanzee
the lower fibres of the short abductor muscle of the thumb have their
origin in the sesamoid bone. The middle fibres, of the same muscle
issue from the strips of ligament attached to the sesamoid bone. On the
other hand, the upper part of the muscle has its origin in the anterior
annular ligament. In the orang, the lower fibres of the short abductor
of the thumb likewise have their origin in the sesamoid bone, while
the central fibres again start from the anterior annular ligament.
The upper fibres are strong, and are inserted into the base of the
first metacarpal bone. In a dissection of the orang the flexor longus
pollicis sent a thin, tendinous expansion on to the bone. This sesamoid
bone is also found in the gorilla, although Duvernoy and Rosenberg do
not appear to be aware of its existence.[72]
In the palm of the gorilla’s hand there is a short abductor, a short
bicipital flexor, an opponens, and an abductor of the thumb. The
longer belly of the short flexor extending in a more radial direction,
and in connection with the opponens, is only slightly developed. In
the muscular system of a gorilla’s little finger we may observe an
abductor, a short flexor, and an opponens. The palm of the chimpanzee
displays a short abductor, an opponens, a short bicipital flexor,
and an adductor of the thumb; also an abductor, a short flexor, and
an opponens of the little finger. In the orang I observed a short
abductor, a short flexor with two bellies, an opponens, and an adductor
of the thumb. In addition to the short flexor of the thumb, Langer and
Bischoff describe another short, independent muscle, representing the
long flexor, and attached to the second phalanx, but I have not myself
ascertained the existence of this muscle. The same anatomists mention
an adductor between the third metacarpal bone and the first joint of
the thumb, and another between the second metacarpal bone and the
second joint of the thumb, passing on into the extensor tendon. I am
myself convinced of the existence of a twofold adductor, but not of the
fact that the tendon of one of the muscles (termed by Langer the second
opponens) passes on into the extensor tendon. In the little finger of
the orang there is an abductor, a short flexor, and an opponens. In
the gibbon there is a short abductor, a faintly indicated opponens, a
short bicipital flexor, and an adductor of the thumb. In _Hylobates
albimanus_ this adductor divides into four or five portions, which
are attached to the whole of the first metacarpal bone. In the little
finger there is an abductor, a short flexor, and an opponens. In the
same animal the first inter-osseous muscle is attached by one portion
to the second metacarpal bone, by the other to the base of the second
phalanx of the index finger (Fig. 53, ~9~, ~10~).
Bischoff has described the muscles which Halford terms _Contrahentes
digitorum_ (contractors of the digits), which lie deep in the palm of
the hands and feet of the chimpanzee and gibbon, the mandril, baboon,
and other apes.[73] They rest upon the inter-osseous muscles, and are
covered by the tendons of the long flexors of the digits, as well as
by the lumbricales muscles. I have been unable to trace these _Musculi
contrahentes_ in the gorilla. In a female chimpanzee I observed a
_Musculus contrahens_ for the fourth, and another for the fifth finger,
and the same for the fourth and fifth toes. In the orang I observed a
_Musculus contrahens_ for the fourth, and one for the fifth fingers,
and two faintly indicated _Contrahentes_ for the fourth and fifth toes.
Similar muscles of the second, fourth, and fifth fingers, and of the
fourth and fifth toes, may be observed in the white-handed gibbon.
In correspondence with the height of the pelvic bones, the gluteus
maximus of these animals only displays a moderate width in comparison
with its length. The tendon which attaches it to the femur extends
low down, almost as far as the knee-joint. The gluteus medius and
minimus are also long, in correspondence with this structure of the
pelvis, although they are attached to the large trochanter, and to
the posterior inter-trochanteric line. The climbing muscle (_Musculus
scansorius_), which extends between the hip-bone and the condyles of
the femur, was discovered by Troill in the chimpanzee, and by Bischoff
in the orang, and is described by them as strongly made; it appears
to be absent in the gorilla and the gibbon. The pyriformis generally
forms portions of the neighbouring muscles. The tensor vaginæ femoris,
which is strong and wide in most anthropoids, is either greatly reduced
or altogether absent in the orang. The sartorius is not, as in man,
attached to the inner surface of the tibia, just below the internal
tuberosity, but it is inserted much lower down on this surface. In
the gorilla it has a tricipital attachment, one to the deep fascia of
the thigh, and two others to the internal border of the tibia. In the
chimpanzee and the gibbon the muscle extends equally low down. In the
orang it does not go so far, but the _gracilis_ and _semi-tendinosus_
are in the same relative position. The biceps of the femur is very
apparent in the orang; its long head divides in two parts, of which the
lower is inserted in the fibula, and is here united with the short head.
Bischoff at first denied the existence of the plantaris in the
chimpanzee, and Brühl had previously done the same, but it is as
normally present in that animal as in man, in whom also it is sometimes
absent. I, however, as well as other observers, have failed to discover
it in the gorilla, orang, and gibbon. The popliteus is developed in
every instance. The tibio-fibular muscle (_Musculus peroneotibialis_),
covered by the popliteus, of which the existence was ascertained by
Gruber, has not been observed by me in any of the anthropoids, with the
exception of the chimpanzee. But it was very apparent in a red sea-cat
monkey (_Cercopithecus ruber_).
The gastrocnemius, which is easily separable into two heads, and the
peroneal muscles have not the same relative width in anthropoids and
man, since in the former case the calf of the lower limb is small, and
it lacks the pleasing roundness which characterizes this part of the
human structure. These muscles, especially in the orang and gibbon,
appear to take a lateral direction. The Tendo Achillis is present,
but it has not the prominent development in height and width which
we observe in man. The long extensor, flexor, and tibial muscles are
in all cases fully developed. The peroneus tertius, as it is termed,
although it should only be regarded as a part of the extensor longus
digitorum, is absent in anthropoids.[74] I myself am not disposed, with
Huxley, Bischoff, and others, to regard this muscle as an abductor.
Brühl perceived in a chimpanzee a fourth rudimentary peroneal muscle
(_Musculus peroneus intermedius_), extending between the peroneus and
the little toe, a muscle sometimes found in man, and which I have
myself only observed in one adult chimpanzee. In the gorilla and the
chimpanzee the extensor longus digitorum passes through a remarkably
strong transverse ligament, formed of fibrous cartilage, which covers
the tarsus. It acts upon the four outer toes (Fig. 55). Brühl has
described the characteristic contraction and extension of the tendons
of the long and short extensors of the toes in the chimpanzee, but I
have myself found some difficulty in producing this action. In Fig. 55
I have endeavoured to represent this condition in the most natural way.
The extensor proprius pollicis is in all cases developed. The extensor
brevis digitorum produces a large, oblique belly for the great toe
(Fig. 55). In the gorilla there is for the great toe an abductor, a
bicipital flexor, an adductor, and an opponens (comp. Fig. 54).
From the extensor brevis digitorum the belly for the great toe rises
with a certain independence. On the right foot of a chimpanzee I
observed a fifth belly of this muscle, going to the little toe (Fig.
55). As my illustration is taken from this specimen, I have represented
the foot with, or in spite of, this interesting anomaly, which, as we
know, sometimes occurs in man.
[Illustration:
Fig. 54.--Muscular system of the human foot. 1, Tibialis anticus
and extensor proprius pollicis. 2, Extensor longus digitorum.
3, Tendon of peroneus tertius. 4, 5, Peroneus longus and
peroneus brevis. 4′, 5′, Tendons of the same. 6, 7, Tendons of
the extensor longus and extensor brevis digitorum.
]
The flexor brevis digitorum displays perforated tendons, belonging
to the second and third toes. The flexor longus digitorum displays
perforated tendons for the fourth and fifth toes. The flexor longus
pollicis divides into two tendons, one of which goes into the toe
itself, while the other is connected with the flexor longus digitorum,
and displays perforated tendons for the third and fourth toes, while
the perforated tendons of the second and fifth toes have their origin
in the other flexor.
[Illustration:
Fig. 55.--Muscles on the upper side of chimpanzee’s foot. 1.
Tibialis anticus muscle. 2, Extensor proprius pollicis. 3,
Extensor communis digitorum. 4, 5, Peroneus brevis and peroneus
longus. 6, Tendon Achilles. 7, Extensor brevis digitorum. 8,
Slip of the same for great toe. 9, First dorsal inter-osseous
muscle. 10, Adductor pollicis. 11, Abductor minimi digiti.
]
In the gorilla the lumbricales muscles of the foot are powerful. The
first inter-osseous muscle is likewise well developed and bicipital.
There is a short flexor and an abductor for the little toe. I have not
yet been able to assure myself of the existence of an opponens for that
toe. In the chimpanzee the muscular system of the great and little
toe does not essentially differ from that which we have described in
the gorilla. The flexor brevis digitorum forms the perforated tendons
of the second and third toes. The flexor longus digitorum provides
the fourth and fifth toes with perforated, and the second and fifth
toes with perforating, tendons, while those which belong to the third
and fourth toes have their origin in the flexor longus pollicis. As
in the gorilla, the latter muscle produces a fibrous investment for
the tendons of the flexor longus digitorum. In the orang there is an
abductor of the great toe, a very slightly developed opponens, a short
bicipital flexor, and an adductor. One of the long flexors of the toes
appears to represent the flexor longus pollicis in man. It provides
the second and fifth toes with perforating tendons, while those of the
third and fourth toes have their origin in the other flexor longus
digitorum. There is no long flexor tendon on the great toe. The
perforated tendons in this case generally belong to the short flexor
muscle. In addition to the perforated tendons of the fourth toe, there
is the long flexor already described.
In a gibbon’s great toe I observed an abductor, a short bicipital
flexor, and a slightly developed opponens, to which a wide fan-shaped
adductor is attached. The first dorsal inter-osseous muscle is, as in
the same animal’s hand (Fig. 53), attached to the first phalanx of the
second toe. The flexor longus digitorum provides the third and fourth
toes with perforating tendons, and also gives off a tendon for the
great toe. On the little toe there is a remarkably slender perforating
tendon. While the first of the two long flexors represents the human
flexor longus pollicis, the flexor longus digitorum is in this instance
limited to the little toe. In the gibbon, as well as in the orang, the
gorilla, and the chimpanzee, the two muscles are connected together
by an aponeurosis. It may be here mentioned that in the human foot
the flexor longus pollicis occasionally gives off a flexor for the
second and even for the third toes. In the gibbon, as Bischoff justly
observes, a muscle covers the flexor longus digitorum, which is still
undivided, but already enlarged. From this muscle perforated tendons
issue for the third and fourth toes. The second toe is provided with
such a tendon from the flexor brevis digitorum. The muscle we have
mentioned seems to represent the Quadratus plantæ, which is often
developed in the other anthropoids, although only to a slight extent.
With respect to the muscles of the small toe of the orang and gibbon,
I need only say that in the latter species the opponens seems to be
absent (Fig. 55).
It will be seen from the foregoing account that, in spite of several
apparently important peculiarities, in spite of great and manifold
variations which are established, even although our authorities do
not always agree together, the muscular system of anthropoids is on
the whole very like that of man. It displays, especially in the lower
limbs, peculiarities of structure which render them capable of walking
in an upright position, and others again which they have in common with
the lower animals, but on the whole the anthropoid characteristics of
the muscular system of these animals are predominant.
The digestive system of anthropoids likewise affords interesting points
of comparison. The cavity of the mouth is, as we have seen, bordered
by large and flexible lips. The mucous membrane of the mouth and the
gums are flesh-coloured; they assume a darker colour in older animals,
and are then sometimes marked with spots of a bluish or brownish grey.
Ehlers describes, as a peculiarity in the mucous membrane of the mouths
of the gorilla and chimpanzee, that there are what he calls buccal
folds, which pass on both sides from the fore surface of the upper and
lower jaw into the mucous membrane of the cheeks, and are of the height
of the canine teeth.[75] I have myself only observed these folds in the
gorilla, of which an illustration is given in Fig. 3, and not in any
other specimen. I have observed scarcely any indications of these folds
in other anthropoids, and then only of such a doubtful nature that I am
not disposed to regard the circumstance as of any special significance.
A small band on the upper and lower lips, sometimes only slightly
developed, but always perceptible, is present in all anthropoids.
The tongue is small, and not provided at its base with several
great concave follicles as in man; these are at least only faintly
represented, and not easy to observe. Around them there rise pock-like,
tufted warts, very close together, which in an aged gorilla are apt to
become hard and horny. These are also prominent between the follicles
of the tonsils. The circumvallate papillæ of the tongue are less
numerous than in man, and often, especially in the chimpanzee, they
take the form of a cross, or of the letter T, or in the gorilla of a V.
The uvula and palate present no special variation from the human
type. On the hard palate there are a number of folds, or rather
swellings, which extend laterally from the central suture of the
palate, towards the row of teeth in the upper jaw; these are sometimes
simple, sometimes complex, and vary in their details in individual
cases. They are particularly marked in the adult chimpanzee, and
are also very apparent in the gibbon, and they are arranged with a
somewhat ornamental regularity. These inequalities are not altogether
insignificant in the human palate, but this subject has not been much
studied since Gegenbaur directed the attention of scientific men to
them, and special light has been thrown upon it by Bischoff and Ehlers,
as far as anthropoids are concerned.
The teeth afford us important material for comparison. In the case of
anthropoids the formula for the teeth of the slender-nosed or Old-World
apes (_Catarrhina_) will generally apply: _i_ 2/2 _c_ 1/1 _p_ 2/2 _m_
3/3. The following is the formula for the milk-teeth: _i_ 2/2 _c_ 1/1
_m_ 2/2. Magitot and Giglioli[76] have shown that the milk-teeth are
cut in the same order as those of man--first, the lower; second, the
upper incisor teeth; third, the front pre-molars; fourth, the back
pre-molars; fifth, the canine teeth. According to the same authors, the
permanent teeth are cut in the following order:--first, the first molar
teeth; second, the lower, and then the upper incisor teeth; third, the
pre-molars; fourth, the canine teeth; fifth, the second molar teeth;
sixth, the third molar teeth. In the skull of a male gorilla, Giglioli
found that the permanent canine teeth were cut almost simultaneously
with the third molar teeth, and after the appearance of the second
molar teeth. The cutting of the canine teeth appears to be a longer
process than that of the other teeth.
In anthropoids the structure of the permanent teeth varies with the
species, and even with the sex. In the gorilla the two upper central
incisor teeth are wide, chisel-shaped, and much larger than the pair of
lateral incisors. The four lower incisor teeth are of about the size
of the upper lateral incisors, and, like these, are chisel-shaped, but
not so wide. The powerful upper canine teeth of an aged male are curved
in their lower part, both outwards and inwards. Their form is that of
a three-sided, cuneiform pyramid. The anterior surface is rounded, and
near its inner edge a deep furrow may be observed, extending from the
neck of the tooth almost to its point. The outer and inner sides of
the tooth meet in a sharp angle, somewhat convex in front, and level
or slightly concave behind. The inner side is concave, and furnished,
nearly in the centre, with a deep longitudinal farrow. The lower canine
teeth of an aged male are shorter than the upper, curved on their
upper and outer surface, and somewhat behind. Their form is also that
of a three-sided pyramid, rounded in front. The longitudinal furrow
which traverses their inner segment is much shorter than that on the
upper tooth. The outer side is somewhat convex, and at the same time
somewhat retreating, and is provided on its posterior segment with
two longitudinal furrows, or more rarely with one, reaching from the
neck to about the centre of the tooth. The inner side, like that of
the upper teeth, is somewhat concave. The lower canine teeth project
like pillars over the upper ones (Figs. 15, 16). The canine teeth of
a young male gorilla are less sharp in their angles, although they
already present the form of a three-sided pyramid. The canine teeth of
the adult female gorilla are much smaller than those of the adult male,
and are laterally more compressed. The three-sided pyramidal form is
only slightly marked. The outer surface is convex and furnished with
a scarcely apparent central longitudinal ridge. On the inner surface,
or that which is turned to the cavity of the mouth, there are from two
to three longitudinal furrows reaching from the neck to the centre
of the tooth. The lower teeth are of a three-sided, pyramidal form,
presenting an interior, posterior, and inner superficies.
The pre-molars of an aged male gorilla are wide, and are furnished
with a large outer, and a smaller inner, cusp. The three four-cusped
upper molars display a more regular and symmetrical arrangement of
their cusps than is the case with the female, in which the position of
the cusps is rather variable. Except for the difference of size, the
relative conditions of these teeth are the same in male and female.
The first pointed lower pre-molars are in the male of the form of a
four-sided pyramid, convex on the anterior and outer surface, flat on
the side directed to the cavity of the mouth, and marked with furrows
on the posterior surface. The small second and lower pre-molars have
two anterior and one posterior cusp. The last is generally worn away
at an early age. Each molar tooth has two outer and two inner cusps,
opposite to each other, and one posterior cusp. We cannot here fail to
notice the likeness to the conditions of the human teeth, a likeness
which is still more striking in the female.
In the chimpanzee, also, the upper central incisor teeth are broadly
chisel-shaped, while the upper and lower lateral incisors are
smaller. In the male there is often a considerable gap between these
and the canine teeth. The latter present the form of a three-sided
pyramid, of which the anterior edge is blunt and tends outwards,
while the posterior angle is sharp, scooped out in its upper third,
and terminating at the base of the crown in a posterior cusp. The
pre-molars have an external and an inner cusp; the molars have two
external and two inner cusps, connected with each other by their
enamel. The lower canine teeth of these animals are likewise of
the shape of a three-sided pyramid, of which the anterior angle is
very blunt, while the inner and posterior angles are sharply cut.
The anterior surface is not grooved like the upper canine teeth.
The lateral angle is much rounded. The back teeth plainly display
the posterior fifth cusp, which may also be observed in man. In the
orang-utan the characteristics of the upper incisors are such as we
have described in the case of other anthropoids. The upper canine
teeth are shaped like a three-sided pyramid, and are furnished with a
longitudinal furrow on the anterior side. A similar furrow is found on
the posterior superficies of the lower canine teeth. The back teeth
display no special characteristics when compared with those of other
anthropoids.
The canine teeth of these anthropoids are much worn down by age
on their posterior surface. Deep transverse grooves of varying
size characterize the teeth of anthropoids, owing to the unequal
distribution of the coating of enamel. These are developed with their
advancing growth. In addition to these incised furrows, longitudinal
marks, with raised edges, also appear, and especially on the anterior
surface of the incisor teeth.
In the gibbon the anterior surface of the incisor teeth is smooth; in
this animal the upper central incisor teeth are the largest, while the
lower central incisors are the smallest. The long and strong upper
canine teeth, which are laterally compressed, display a sharp posterior
angle, and an anterior and inner longitudinal furrow.
It has sometimes been said that the grooves found on the external
contour of the back teeth of anthropoids, extending to their roots,
constitute a not unimportant distinction between their structure and
that of the human teeth, in which the grooves do not extend to the
roots. But the corresponding human teeth do sometimes exhibit very
deep and extensive furrows. I cannot, therefore, ascribe any peculiar
significance to this assumed distinction. The development of the canine
teeth, like those of beasts of prey, seems to me much more important. A
supernumerary back tooth may sometimes be observed both in man and in
anthropoids, including also the gibbon.[77]
The stomach and intestines of these animals present only a few striking
differences from the same organs in man. The length of the intestines
varies in man as well as in anthropoids. I have only observed the
_valvulæ conniventes_ to be somewhat clearly developed in the gorilla
and the orang. The cæcum of these apes is long, broad, placed with the
power of free movement in the peritoneum, and furnished, especially in
the case of the orang, with a large, very long, and spirally coiled
vermiform appendix.
The liver is divided into two principal lobes, but in the orang
this division is not very clearly marked. I have not myself observed
a subdivision of these lobes, occurring on their edges, which is
mentioned by Bolau and Auzoux in the case of the gorilla. Bischoff
notices in the gorilla the absence of the H-shaped arrangement of the
fissures on the under surface of the liver, so noticeable in man; and
the same remark applies to other species of anthropoids. Moreover, the
fissures on this part of the liver are not incised on the substance
with the same uniform depth. The gall-bladder of the gorilla and the
orang is not remarkable for its size; in the chimpanzee I found that
this organ is large and twisted, and it is also large in the gibbon.
The spleen is elongated in the gorilla, chimpanzee, and gibbon, shorter
and wider in the orang. On its left contour it is uniformly bevelled
off. There is nothing in the pancreas which calls for remark.
The larynx of anthropoids possesses on the whole a structure resembling
that of man. This is especially the case at the entrance to that organ.
The anterior and specially vocal portion of the glottis is short, about
as long as the respiratory portion. In the chimpanzee there is a deep
cavity in the body of the hyoid bone. In the gorilla, chimpanzee, and
orang the throat-pouches or air-sacs correspond to Morgagni’s sacs.
These are the thin-skinned elastic sacs, closely united with their
surroundings by connective tissue. The right laryngeal sac appears
to be of larger diameter than the left. According to Duvernoy’s and
Ehlers’ accurate account only the upper portion of this organ occurs
in the gorilla. In that animal, and in the orang, a lower projection is
displayed, extending behind the sterno-mastoid as far as the shoulder,
and another extending to the pectoralis major muscle. In the chimpanzee
only the posterior segment is developed. It has been asserted that
in several cases there is found a single, irregular laryngeal sac,
communicating with the two Morgagni sacs, but I agree with Ehlers in
thinking this improbable. In such instances it seems likely that, owing
to the great want of symmetry in this organ, one of the sacs has been
overlooked. In an aged orang the throat-pouches, fastened together by
connective tissue, and covered by the external skin of the throat,
hang down slackly and heavily over the middle of the breast (see Fig.
9). According to Sandifort, the siamang is the only one of the gibbons
which displays a single throat-pouch; while Broca asserts that it
has two detached sacs, placed close to the larynx.[78] The halves of
the thyroid cartilage are generally connected with each other by an
intermediate piece.
The trachea of anthropoids generally includes from sixteen to eighteen
cartilaginous rings, but in the siamang there are twenty-one. They
ramify into branches which are, as a rule, wider on the right than on
the left side.[79] There is a further lateral ramification on the
right side, situated above the artery. Huxley and Ehlers hold that the
lungs of a gorilla are cleft like those of the human organism, the
right divided into three, and the left into two lobes. I have myself
observed this type, and in one instance I found three lobes on the
left. In the chimpanzee I saw that the right lung was divided into
three, and the left into two lobes. Bischoff observed an instance of
a chimpanzee which had four lobes on the right and two on the left
side. In an orang dissected by me I found only one lobe on each side,
with thin, slightly indented notches on the anterior edges of the
right lobe, and two on the left, and there was at the same time a
strongly marked indentation between the lobes. The lungs of a gibbon
are described as having four lobes on the right, and only one or two
on the left. I myself have examined a gibbon in which there were three
lobes on the right, and two on the left. It appears that there are not
unimportant individual variations of this structure in every species of
anthropoids; and indeed, human lungs are by no means exempt from them.
The male sexual organs correspond on the whole with the form and
arrangement of these organs in man. I must not omit to mention that the
penis of the swine-snouted baboon, and of other dog-headed apes, is
much more like the penis in man than is the case with anthropoids, with
the exception of the gorilla. In the last-named animal the scrotum is
short and tightly stretched. The right testicle is a little higher than
the left, and is divided from it by a wide raphé. The internal female
organs are also like those of the human organism, with only slight
variations. Bischoff is correct in the assertion that the external lips
of the pudendum and the mons veneris are almost wholly absent. Bolau,
Ehlers, and Hermes have ascertained that there is a menstruation which
occurs periodically, at any rate in the case of the chimpanzee, and the
other species cannot be exempt from the process. At such times there
is a blush and enlargement of the external parts, and a profusion of
the external lips of the pudendum, which are at other times scarcely
apparent. The nymphæ and the clitoris are of considerable size and
importance. There is often an excessive enlargement and reddening
of these parts, as well as of the posterior callosities in the
chimpanzee, and also in the baboon and macaca, during the period of
sexual excitement.
[Illustration:
Fig. 56.--The brain of an orang, seen from the side (Vogt, from
Gratiolet). F, Frontal lobe. P, Parietal lobe. O, Occipital
lobe. R, Fissure of Rolando. S, Fissure of Sylvius. C,
Cerebellum.
]
_Nervous system._--In this part of the organism we are especially
interested in the structure of the brain. Bastian justly remarks,
with reference to the brain of apes, that this family possesses many
cerebral characteristics in common, by which their close connection
with each other may be verified. Distinct stages of development have
been observed, which, however, cannot be classified in a consecutive
series. Starting from the brains of lemurs, which do not greatly
differ from those of rodents, we can advance by means of very distinct
transition forms to the more highly developed cerebral hemispheres
of the large anthropoid apes, the chimpanzee, the gorilla, and
orang-utan.[80]
[Illustration:
Fig. 57.--Brain of the chimpanzee, seen from above. The upper
part of the right hemisphere is removed so as to lay bare
the lateral ventricle (Vogt, from Marshall). L, Longitudinal
fissure (other indications the same as in Fig. 56). _c s_, The
corpus striatum in anterior cornu of the ventricle. _c a_,
Hippocampus major in descending cornu. _h m_, Hippocampus minor
in posterior cornu.
]
Very opposite views prevail among anatomists with regard to the
question which species of anthropoids possesses the most highly
developed brain. Some regard the chimpanzee’s brain as the simplest,
and that of the orang as the most highly developed. In all these apes
the lateral halves of the cerebrum, always divided from each other
by a deep longitudinal fissure, overlap the cerebellum as far as a
minute posterior segment. In this respect I find the brain of the
gorilla a little behind the other anthropoids. Up to this time, I have
only observed the projection of the cerebellum through the cerebrum
in the case of an orang[81] (see also Fig. 56). Retzius asserts that
the cerebellum of Lapps is incompletely covered, while the covering
is generally complete in the case of Slav and Tartar races. In German
and Latin races the cerebrum overlaps the cerebellum. In Mongolian,
Indian, and Negro races the covering appears to be generally imperfect.
[Illustration:
Fig. 58.--Brain of gorilla, side view (from Bolau and Pansch).
I., Frontal lobe. II., Fissure of Rolando. III., Parietal lobe.
IV., Temporal lobe. C, Cerebellum. _f s_, Fissure of Sylvius.
_s c_, External fissure parieto-occipital.
]
While the ground form of the gorilla brain approximates to a long
oval, and in this respect resembles the human brain, the brain of
chimpanzees and orangs is of a round-oval form. This is especially the
case with the chimpanzee (Fig. 57). In my opinion, the gorilla brain
is distinguished from that of the chimpanzee, but not from that of the
orang, by its very complex convolutions (Fig. 56).
[Illustration:
Fig. 59.--Brain of orang, seen from above (Duncan, from a
specimen in the Museum of Royal College of Surgeons). F,
frontal lobe. O, Occipital lobe.
]
In the gorilla, chimpanzee, and orang, the island of Reil in
the fissure of Sylvius is generally--at least, according to my
experience--overlapped by the operculum, although there are instances
in which this is not the case. In these three anthropoids, as Bastian
justly observes, the fissure of Sylvius is much less horizontal than
in man, and occupies a position more like that which it takes in the
black sea-cat monkey, the wanderers, and other macacas. In the gorilla
its direction is more horizontal than in the two other species of
anthropoids. The central fissure, termed fissure of Rolando, is very
marked, especially in the chimpanzee (Fig. 57 R); but it may also be
easily traced in other species of anthropoids (Fig. 58, II., 56, R).
The so-called simian fissure between the parietal and occipital lobes
of the cerebrum (Meynart’s elongated external occipital fissure),
presented in Fig. 58 _s c_, is very marked in the chimpanzee (Fig. 57,
_d_). The frontal lobes of the gorilla brain are high, while those of
the chimpanzee are short and low. It is said that those of the orang,
which are high and short, terminate in a beak-shaped curvature, but
this is not invariably the case.
[Illustration:
Fig. 60.--Longitudinal section of a gorilla’s brain (Bola and
Pansch). _s.cm_, Colloso marginal fissure. _f, p_, Internal
parieto-occipital fissure. _f, c_, Calcarine fissure, the
posterior part of the hippocampal fissure.
]
In the anthropoids we have been considering, and also in several of
the lower species of apes, there are three other fissures of less
importance in addition to those we have mentioned, namely, the fissure
parallel to the fissure of Sylvius, and placed behind it, the _corpus
callosum_ fissure, placed immediately above the _corpus callosum_ on
the inner side of the hemisphere of the cerebrum, and the calcarine
fissure (_Fissura calcarina_) (Fig. 60). The latter ends near the point
of junction of the inner and lower surfaces of the posterior division
of the hemisphere. The upper temporal convolution, termed by several
anatomists _Gyrus supramarginalis_, is said by Gratiolet to be absent
in anthropoids; but Rolleston, Bastian, and myself have all found it
well developed[82] (Fig. 56, orang, and Fig. 58, gorilla).
Bischoff asserts that the third frontal convolution (Broca’s
convolution) is very slightly developed in the chimpanzee, orang,
and gibbon. “Its great development in men,” Gewährsmann writes,
“constitutes one of the most marked distinctions between the brains
of apes and of men.”[83] In most of the other species of apes this
convolution is altogether absent, but Pansch is justified in the
assertion that it is fully developed in anthropoids. I cannot wholly
agree with Pansch in his analysis; but I must accept his statement
on this point (see the orang, Fig. 59). Gratiolet remarks that the
so-called annectant gyri (_plis de passage_) which serve as a covering
or _operculum_ for the posterior lobes in apes, are only superficially
apparent in man. In the chimpanzee the upper of those convolutions
is absent, while it is large in the orang, and likewise large and
undulated in man. In the orang the second annectant gyrus is covered,
but this covering is absent in man.[84]
In considering the inner structure of the brain of these animals, we
are first struck by the shortness of the _corpus callosum_. The soft
and thick anterior commissure of the third cerebral ventricle and the
thin posterior commissure have also been justly noted. In the lateral
ventricles more of the characteristics described in the human brain are
absent. The four eminences resemble those of man; nor does the fourth
cerebral ventricle present any remarkable differences of form. Neither
does the base or lower surface of the brain display any important
deviation from the human type. The transverse section of the nerves at
their intersection appears to me, however, to be somewhat more oval
than is the case in man.
There has recently been an attempt to recognize a pithecoid character,
or atavism, in microcephalic men, the smallness of whose heads is
allied with a greater or less degree of idiocy. A pithecoid structure
of the brain has also been traced in several individuals who are not
microcephalous, but subject to pathological affections. We will first
consider those who belong to the latter category. Krause examined
the brain of an ape-like boy aged seven years and a half, which, as
the author remarks, approximated in structure to the pithecoid type,
although without displaying microcephalic characteristics. The two
cerebral hemispheres were wanting in symmetry; they diverged from each
other in the region where the parieto-occipital fissure occurs on the
left cerebral hemisphere, and they formed an edge which curved outward
and backward so that the cerebellum remained uncovered. On the lower
surface of the frontal lobes there was a strongly marked ethmoidal
prominence. Neither of the fissures of Sylvius were closed, the left
less so than the right; the operculum was only slightly developed;
and the island of Reil and its fissures were almost uncovered. This
formation is almost the same as that of the brain of anthropoids.
The two central fissures of Rolando were close together, or less
deeply impressed on the edge of the hemispheres than is normally the
case, and forming no joint angle. Large and deeply marked pre-central
fissures seemed to represent the central fissures. The intra-parietal
fissures, diverging outwardly further than in man, received the
parieto-occipital fissure, a structure in conformity with the typical
brain of apes. The transverse occipital fissure became in this case a
deep fissure like the simian fissure, crossing the occipital lobes,
and almost completely dividing them from the parietal lobes. The
so-called _Fissura calcarina_, to which we have referred above, had
its origin on the upper surface of the occipital lobe, then joined
the parieto-occipital fissure, and went directly into the hippocampal
fissure (_Fissura hippocampi_) on its right side. This abnormal
structure is also in conformity with the typical brain of apes. The
first occipital convolution is divided from the upper parietal lobes by
the parieto-occipital fissure. Gratiolet asserts that this formation
occurs in many species of apes. The upper temporal convolution was
remarkably reduced on both sides, possessing only an average width of 5
mm. This characteristic reminded Krause of the brain of the chimpanzee.
In that animal the upper temporal convolution is always reduced.
Krause therefore asks whether some human brains may not possess the
typical structure of apes without being microcephalic. The brain we
have described scarcely differed from the normal weight; it possessed
all the convolutions and fissures, and indeed, the convolutions
were perhaps more numerous than in the normal structure, yet it was
different in every respect, and approximated in its whole structure to
the simian rather than to the human type. Krause adds that if the brain
had been placed before him without any intimation of its origin, he
should have been quite justified in concluding that it belonged to an
anthropoid ape, which stood somewhat nearer to man than the chimpanzee.
It is an unquestionable fact that some human beings, whether children
or adults, who are endowed with a defective bodily structure, and who
are affected with more or less pronounced physical incapacity and
mental weakness, by their appearance, ungainly tricks, and helpless
and aimless motions, impress us in the most forcible way with their
resemblance to apes. Different degrees of idiocy affect individuals of
limited intellect, and remind us of an absolutely brutish condition.
Krause describes the “ape-like” boy of seven and a half years old,
whom he had examined, as cheerful and inclined to play and dance, but
as passionate when he was teased. The child was very supple, fond of
climbing, and with great strength in his arms and hands, of which
the latter had a horny appearance, reminding him of the hands of a
chimpanzee. He could sit on the ground with his legs wide apart. His
gait was uncertain, and he was apt to tumble, falling with his knees
bent forward and his legs doubled under him; he was fond of hopping,
and at such times looked still more like an ape. The great toes of
both feet were at an angle to the foot, and thus gave the impression
of a prehensile foot. At first Krause supposed that this deviation was
produced by the child’s endeavour to supply a broader basis of support
for his uncertain gait; but he subsequently changed his opinion, since
he did not find the same peculiarity in other children of diseased
brain, as, for instance, in those suffering from water on the brain.
The boy could say very little, only papa and mamma, and it was long
before he could pronounce these words in two syllables; for the most
part, he only uttered a sound resembling a grunt. He imitated the
barking of a dog, with the sound of rolling _r_’s. He often stamped
his feet and clapped his hands together, making a grunting noise as
Krause had observed in the case of gorillas and chimpanzees. The boy
was smaller than other children of his age, and had weak eyes; his head
was sore, and his forehead narrow. His imitative tendency was strongly
marked, and his whole nature and all his movements strikingly resembled
those of apes. He had been much neglected by his parents.[85]
When I was a student at Berlin I had the opportunity of observing
a similar being of twelve years old, in what was at that time the
Weinbergswege, near the Rosenthaler Gate. This was a boy with a large
head, a low retreating forehead, glazed eyes, a morose expression, a
thin neck, prominent belly, crooked legs, large hands and feet. The
boy was of a slouching appearance, and his gait was unsteady: saliva
often dribbled from his wide mouth; and as he walked he held on to the
furniture, walls, etc., and often he fell powerless on his side, and
so remained in a crouching position. It seemed to give him peculiar
pleasure to creep on his hands and knees, and at such times he would
stamp with the closed fingers of one or the other hand upon the
ground, as if in triumph. This habit, his gait, and the gurgling sound
which was all that the boy could utter, constituted the points of his
resemblance to apes. All the other conditions of life were those of a
being whose mental and physical growth was arrested, and who, although
not epileptic, was to a certain extent idiotic. I am ignorant what
afterwards became of him.
In the course of a discussion on the instance adduced by Krause,
Virchow asks whether the psychological conditions of such a brain
are indeed simian. He is convinced that whoever has studied the
microcephalic child Margaret Becker (of Bürgel, Hanau) will find that
psychologically she had nothing in common with an ape. In her case
all the positive faculties and qualities of the ape were wanting;
the simian psychology was altogether absent, and there was only the
psychology of an imperfectly developed and deficient young child. Every
characteristic was human. Virchoff had the child in his room for hours
together during a period of two months, and was constantly occupied
about her, without observing anything in her nature which reminded
him even remotely of the psychological conditions of apes. She was a
degraded specimen of humanity, differing in no respect from the human
type.[86]
I also examined Margaret Becker, as well as another microcephalic
girl, who was in the Berlin Asylum in the years 1868 and 1869. With
respect to the former and more animated being, I have nothing essential
to add to the information published by Virchow. Ida X----, the other
individual whom I examined at Berlin, was at the time of my researches
aged thirteen years and five months. Her figure was slightly made and
well proportioned, while her profile reminded me to a modified extent
of that of the microcephalic Aztec, and also of the heads represented
in ancient sculpture of Mayapan, Palenque, and Copan. I must not omit
to say that Ida had light blue eyes and fair, glossy hair. She was
altogether impassive; could only utter the syllables _da-da_; and
once betrayed a slight sign of displeasure when the cold metal of the
measuring-rod was placed against the inner side of her thigh, for the
sake of obtaining the dimensions of the different parts of her body.
Virchow’s information respecting Esther Jacobwitz, of Waschahel, is
also extremely interesting. She was a microcephalic girl of the age of
fourteen, and a Hungarian Jew by race.[87] Virchow remarks that, in
his opinion, all Esther’s most striking characteristics presented the
strongest contrast to those of apes, since only negative traits have
hitherto been established, while all which characterizes the positive
development of the psychical life of apes was absent in this case. The
same remark applies to Ida X----. Virchow goes on to say that there
was undoubtedly something brute-like in the defects in question, but
that in order to reproduce the animal in its actual form and nature, so
as to show that the microcephalic child was really theromorphic, the
positive side of animal life must to some extent be presented to us,
and this was absolutely wanting.
Virchow also had the opportunity of examining a pair of twin children,
one of whom was quite normally developed, while the other (Karl
R----) was microcephalic. This was a very significant case, since two
individuals of the same birth were under consideration, so that the
question could be asked with greater confidence--Is this atavism, or a
morbid condition? From this point of view, it was of special interest
to establish the fact that the microcephalic child had, in fact,
displayed positive signs of a morbid condition.[88]
When I go through the accounts collected by C. Vogt of the lives
of well-known microcephalic beings,[89] I can find nothing which
specifically reminds me of the actions and habits of apes, although
we have an intimate acquaintance with their ways. These individuals
give the general impression of human beings whose bodily and mental
development has been arrested. According to Virchow’s experience,
all the cerebral disturbances are concentrated in the cerebrum in
these microcephalous cases. The anterior portions of the cerebrum are
affected to the greatest, and the posterior to the least, extent. Those
parts which are developed latest suffer the most, while those which are
the first to be developed generally escape disturbance.[90]
Klebs, Schaaffhausen, and others have sought to show that the mothers
of microcephalic children have suffered from severe pains of the uterus
during pregnancy. All scientific men consider that spasms of the uterus
distinctly affect the development of the brain of the offspring. Flesch
thinks it possible that these spasms of the uterus may have something
to do with the origin of microcephaly.[91] But he also asks whether
this morbid condition of the uterus may not have been produced by a
previously diseased condition of the offspring. This observer is,
moreover, still more inclined to make the influence of the father
responsible for the occurrence of microcephaly. In view of the fact
that there is much reason to suppose there has been a compression of
the uterus, and in default of any better suggestion, Flesch feels
justified in looking for a compression which has perhaps resulted
from some growth on the ovary. Hence ensues a disturbance, probably
inflammatory, of the organ of nutrition.[92]
Aeby also regards microcephaly, not as an expression of atavism, but
as the result of a morbid degeneration. “Microcephalic subjects do
not point back to the milestone which man left behind him in hoar
antiquity, and it is not through them that the chasm between man and
animals can be bridged over, nor even rendered less wide.”
Virchow’s researches led to the following conclusions, which we must
here subjoin:--1. There is no species of apes which presents that
precise configuration which is found in a microcephalic brain. 2.
Psychology offers the strongest arguments against men-apes. 3. The
instinctive side of psychical activity, which is almost wholly absent
in microcephalic subjects, is very prominent in anthropoids as well as
in other animals.[93]
In addition to these remarks, it may also be observed that among savage
races the medicine-men, shamans, sorcerers, rain-doctors, etc., often
assume ape-like attitudes in the contortions, leaps, dances, and other
gestures which are inseparable from their trade. Owing to their state
of excitement, in which they are not always mentally responsible for
their acts, this imitation may be often partly or wholly unconscious.
It is very common among the inspired Arabs termed Haschasch, who,
sometimes as dervishes, sometimes as poets or beast-tamers, roam
through the country and extend their wanderings from the interior
of Africa to the latticed gates of Dolma Bakhtsche. To them belong
also the dancing mendicant monks of Islam, who display their ape-like
gesture in the market-places and streets of Bokhara, as well as in the
other chief cities of Central Asia. In this case, indeed, many gestures
are conventional, and even adopted as the means of stimulating the
proposed effects, but at the same time they impress us with the idea
that a man under such conditions of life and work involuntarily adopts
the gestures of anthropoids. When we see a Zikr, an Islamite rite of
worship, accompanied by obligatory howls and contortions of body, we
are tempted to imagine ourselves in the midst of a troop of wild apes.
And the illusion is still stronger if the performers in the Zikr are
black fakirs, dressed as warriors.
The peripheral nervous system of anthropoids has not, up to this
time, been analyzed with the completeness we could wish. As far as
the observations of Vrolik, Gratiolet, and Alix go, together with my
personal experience in this department, no marked distinction can be
established between the structure of these organs in anthropoids and
those of the nervous system in man.
H. von Ihering has studied the relation of the nervous lumbo-sacral
plexus to the vertebral column of men and animals, and has come to
the conclusion that there is the most complete agreement between men
and animals with respect to the relations of the vertebral column to
the peripheral nervous system. According to this author, man, from
the anatomical point of view, stands so completely within the class
of anthropoids, that the attempt to assign to him any other place in
zoology is open to the charge of being biassed by considerations which
have nothing to do with facts.[94]
The organs of the senses in anthropoids do not present any noteworthy
points of difference from these organs in man. I have written, but not
yet published, a treatise on the eyes of these animals, showing their
general agreement with the conditions of the human eye. On the skin
of the fingers and toes of anthropoids developed corpuscles may be
detected which are connected with the sense of touch.
The vascular system of anthropoids has not up to this time been studied
in any exhaustive manner. The heart strongly resembles that organ in
man. In the gorilla, the chimpanzee, and the orang the great arterial
branches have the same relative conditions as in the human organism.
A common origin from one branch of the subclavian artery, and of the
right and left carotid arteries, often occurs in the orang and with
a certain constancy in the gibbon, so far as we can judge from the
researches which have been made up to this time. But we know that this
form of deviation from the common type is not altogether rare in man.
Bischoff and others have justly maintained that the resemblance to man
which is found in these animals in the arrangement of the heart and
larger blood-vessels appears to be connected with their mode of life.
For although their habits are arboreal, this very fact implies that
they are for the most part in an upright position.
The division of the femoral arteries displays a somewhat interesting
deviation from the normal human type. High up near the femoral arch
an artery, accompanied by veins and a large nerve, diverges from the
femoral artery, which extends, together with its accompanying parts,
as far as the back of the foot. In the gorilla this branch pierces the
sartorius.