Fig. 31.—Skeleton of the Superior Limb of a Bird (Vulture): Left Side, External Surface.
1, Humerus; 2, radius; 3, ulna; 4, radial bone of the carpus; 5, ulnar bone of the carpus; 6, first metacarpal; 7, second metacarpal; 8, third metacarpal; 9, first digit, the homologue of the thumb; 10, first phalanx of the second digit; 11, second phalanx of the second digit; 12, third digit.
The hand, in birds, is directed obliquely downwards and backwards (Fig. 31). For the better understanding of its position in relation to the forearm, we should remember that this latter, as we have described (p. 44), directed obliquely upwards and forwards, has the radius placed above the ulna; the hand being oblique in the opposite direction and placed under the forearm is, by this arrangement, inclined towards the ulnar border of the latter.
Fig. 32.—Superior Limb of the Human Being, the Different Segments being placed in the Attitude which the Corresponding Parts occupy in Birds: Left Side, External Surface.
For the rest, in order to be able to distinguish readily the corresponding parts in the hand of a bird and that of a man, we merely have to place the human forearm obliquely, in a direction upwards and forwards (Fig. 32), the radius being above; this position we can obtain by semi-pronation; then, to incline strongly the hand downwards and backwards, moving the ulnar border of the hand towards the ulna; the thumb is then anterior, the little finger posterior, and the palm of the hand is turned towards the trunk.
The carpus in birds is formed by two bones only, with which the skeleton of the forearm articulates. That which is in contact with the radius is called the radial bone of the carpus; and that with which the ulna articulates is named the ulnar bone.
The metacarpus is formed of three bones; the first, which is very short, is fused at its superior extremity with the adjoining one; this latter and the third, both longer than the first, but of unequal size, are fused at their extremities. The metacarpal, which articulates with the radial bone of the carpus, is larger than the one which is in line with the ulna. To the metacarpus succeed three digits, of which the central is the longest, and is formed of two phalanges; the other two are formed each by a small, stylet-shaped bone. The middle finger, situated on the prolongation of the metacarpal, which articulates with the radial bone of the carpus, has its first phalanx large and flattened transversely; this phalanx seems to have been formed by the union of two bones of unequal development; the second phalanx is styloid in form. As to the other two fingers, they are placed, one in front and the other behind; the first, which articulates with the short metacarpal, fused at its upper end with the principal bone of the metacarpus, in position represents the thumb. The other, which is the third finger, articulates with the inferior extremity of the thinnest bone of the metacarpus; it is sometimes closely united to the corresponding border of the first phalanx of the large—that is to say, of the median—digit.
Plantigrades: Bear (Fig. 33).—The scapula of the bear approaches in shape to a trapezium, of which the angles have been rounded off. The anterior border (cervical) is strongly convex in the part next the glenoid cavity. The junction of the superior (spinal) and the cervical border forms almost a right angle, the summit of which corresponds to the origin of the spine. At its posterior angle there is a prominence, directed downwards, the surface of which is hollowed and is separated from the infraspinous fossa by a crest, so that at this level a third fossa is added to the infraspinous one. The neck of the scapula is but slightly marked. The acromion is prominent, and projects a little beyond the glenoid cavity.
Fig. 33.—Skeleton of the Bear: Left Lateral Surface.
1. Cranium; 2, face; 3, atlas; 4, axis; 5, seventh cervical vertebra; 6, first dorsal vertebra; 7, fourteenth and last dorsal vertebra; 8, lumbar vertebræ; 9, sacrum; 10, coccygeal vertebræ; 11, sternum; 12, ninth and last sternal rib; 13, costal cartilages; 14, acromion process; 15, third fossa on the external surface of the scapula; 16, great tuberosity of the humerus; 17, musculo-spiral groove; 18, epicondyle; 19, radius; 20, ulna; 21, olecranon process; 22, carpus; 23, pisiform; 24, metacarpus; 25, phalanges; 26, ilium, external fossa; 27, pubis; 28, tuberosity of the ischium; 29, obturator foramen; 30, great trochanter of the femur; 31, condyles of the femur; 32, patella, or knee-cap; 33, anterior tuberosity of the tibia; 34, fibula; 35, tarsus; 36, calcaneum, or heel-bone; 37, metatarsus; 38, phalanges.
The clavicle is rudimentary, but, as an example of the complete development of this bone in plantigrade quadrupeds, we may cite the marmoset.
The humerus is furnished at its superior extremity with a large tuberosity, wide, and situated in front of the head of the bone; the effect of this is that the bicipital groove is internal. As in man, the great tuberosity does not reach so high as the humeral head, but it approaches more nearly to that level. The deltoid impression is very extensive, and descends pretty far down on the body of the bone. The epitrochlea is prominent; the epicondyle is surmounted by a well-marked crest, curved and flexuous in outline.
The articular surface, which is in contact with the radius, is not a regularly formed condyle; it is a little flattened on its anterior surface, and presents at this level a slight depression which corresponds to a small eminence on the anterior aspect of the superior extremity of the radius. The surface which articulates with the ulna, viewed on its anterior aspect, has the shape of a slightly-marked trochlea; except at the level of the internal lip, which, as in man, descends lower than the surface for articulation with the radius (condyle). Behind, the trochlea is more clearly defined.
The bear possesses a considerable power of rotation of the radius; the bones of the forearm are joined only at their extremities, while in the remainder of their extent they are widely separated. The ulna terminates below in a head and a styloid process; these articulate with the two last bones of the first row of the carpus—viz., the cuneiform and pisiform. The bones of the carpus are seven in number, the scaphoid and the semilunar being fused together.
The metacarpals, five in number, differ very little from one another in regard to length, though they increase in size from the first to the fifth; this may be demonstrated by looking at the palmar surface of the hand. It is the reverse of that which we find in man, for the fifth metacarpal is the thickest of all, and the first is the most slender.
At the level of each metacarpo-phalangeal articulation are two sesamoid bones.
The third digit is the longest. The terminal phalanges present two very different portions: one, the anterior, is curved and pointed; it serves to support the nail, whose shape it assumes; the other, posterior, forms a sort of sheath into which the base of the nail is received.
The inferior portion of the posterior surface of this latter part articulates with the second phalanx in the case of each of the last four digits, but with the first phalanx in the case of the thumb.
Fig. 34.—Skeleton of the Dog: Left Lateral Surface.
1, Cranium; 2, face; 3, atlas; 4, axis; 5, seventh cervical vertebra; 6, thirteenth and last dorsal vertebra; 7, lumbar vertebræ; 8, sacrum; 9, coccygeal vertebræ; 10, anterior extremity of the sternum; 11, xiphoid appendix; 12, ninth and last sternal rib; 13, costal cartilages; 14, spinal border of the scapula; 15, supraspinous fossa of the scapula; 16, infraspinous fossa of the scapula; 17, great tuberosity of the humerus; 18, deltoid impression; 19, musculo-spiral groove; 20, olecranon process; 21, radius; 22, carpus; 23, pisiform; 24, metacarpus; 25, sesamoid bones; 26, phalanges; 27, ilium, iliac crest; 28, pubis; 29, tuberosity of the ischium; 30, great trochanter of the femur; 31, patella, or knee-cap; 32, anterior tuberosity of the tibia; 33, fibula; 34, tarsus; 35, calcaneum, or heel-bone; 36, metatarsus; 37, sesamoid bones; 38, phalanges.
Digitigrades: Cat, Dog (Fig. 34).—In these animals the anterior (cervical) border of the scapula is convex; the posterior (axillary) border is straight or slightly concave. The supraspinous and infraspinous fossæ are of equal extent (Figs. 35 and 36). The neck is short. The spine of the scapula becomes more and more prominent towards its inferior extremity, where it ends in a twisted and inflexed portion, which represents the acromion process; this process terminates at the level of the glenoid cavity. The coracoid process is represented by a small tubercle, slightly curved inwards; this tubercle is situated above the glenoid cavity, at the inferior part of the cervical border.
Fig. 35.—Scapula of the Dog: Left Side, External Surface.
1, Posterior or axillary border; 2, superior or spinal border; 3, anterior or cervical border; 4, spine of scapula; 5, coracoid process; AA′, length of spinal border.
Fig. 36.—Left Scapula of the Cat: External Surface.
1, Posterior or axillary border; 2, superior or spinal border; 3, anterior or cervical border; 4, spine of the scapula; 5, coracoid process; AA′, length of the spinal border.
In the dog, the posterior angle, formed by the junction of the axillary and the superior (spinal) borders, is obtuse; the spine rises perpendicularly from the surface of the bone. The width of the scapula, measured at the level of the spinal border (from A to A′, Fig. 35), equals about half the length of the spine. We must, however, make an exception for the turnspit dog, in which the superior border equals three-fourths of that length. The scapula is, in this case, of a more compact type; it is broader, but shorter. In the cat, the anterior outline of the scapula, formed by the union of the cervical border and the corresponding half of the spinal, is more convex; the posterior angle is not obtuse, as in the dog. The spine is bent slightly downwards and backwards; before terminating in the acromion process it presents a triangular projection, the apex of which is directed downwards. The tubercle which represents the coracoid process is curved inwards more strongly than that of the dog, thus resembling more closely the appearance of this process in the human being.
All proportions considered, the scapula of the cat is broader than that of the dog; its width, measured along the length of its spinal border (from A to A′, Fig. 36), equals three-fourths of the length of the spine.
The clavicle is rudimentary; it is, however, better developed in the cat than in the dog. The clavicle of the cat is represented by a small, elongated bone, curved in outline, the convexity being turned forward; it is united to the acromion and the sternum by ligamentous fibres; that of the dog is merely a scale-like osseous plate situated on the posterior surface of a muscle of this region (see Figs. 16 and 17).
The humerus is long and twisted in the shape of an S. The inferior articular surface has the form of a simple pulley, for the condyle is very slightly marked. The internal part of this articular surface descends lower than the external; this condition resembles that found in the human being, where the inner lip of the trochlea is lower than the condyle.
In the dog, the olecranon fossa communicates with the coronoid by an opening.
In the cat, there is a supra-epitrochlear canal (see Fig. 19), but no olecranon perforation.
The bones of the forearm articulate at their extremities. The body of the radius is united to the body of the ulna by a short, thick, interosseous ligament; the fibres of this ligament, though short, do not prevent the production of some movements at the articulations of the bones.
The radius so crosses the ulna that above, it is in front and external to the latter, while below, it is internal. This bone is flattened from front to back, and slightly convex anteriorly. Its superior extremity is formed, externally, of a portion which represents the head of the radius in man; internally, by another portion which represents half of the coronoid process of the ulna, which, in the human being, belongs exclusively to the latter (see p. 39, the encroachment of the radius on the ulna). This extremity is surrounded with a vertical articular surface which is placed in contact with a small cavity which is hollowed out on the ulna (the lesser sigmoid cavity); and presents at its superior aspect a surface which articulates with the inferior extremity of the humerus. The shaft of the bone has on its internal border rugosities analogous to the imprint of the pronator radii teres of the human skeleton; these rugosities, indeed, give insertion to a muscle of the same function, and bearing the same name. The inferior extremity, broader than the superior, is hollowed on its external aspect by a small cavity which receives the inferior extremity of the ulna; its inferior surface (concave) articulates with the carpus; its anterior surface (the homologue of the posterior surface of the corresponding extremity of the human radius) presents grooves which serve for the passage of the tendons of the muscles which pass from the forearm to the back of the hand. (For the names of the muscles whose tendons pass in these grooves, see Fig. 29.)
The ulna is furnished at its superior extremity with an olecranon process, which is more prominent than that of the human ulna; this process is compressed laterally, and its internal surface is hollowed; there we also find a great sigmoid cavity, and a coronoid process situated at the internal part of the anterior surface, a process which, as we have previously shown, it shares with the radius.
The shaft of the bone, prismatic and triangular, diminishes in thickness as it approaches the lower extremity, which articulates with the corresponding extremity of the radius. In the dog, the ulna terminates inferiorly in a blunt point, without enlargement, analogous to the head of the human ulna; in the cat, by a head which is prolonged into a styloid process, by which it articulates with a portion of the carpus.
The carpus consists of seven bones—three in the superior row and four in the inferior. In the superior row the scaphoid and semilunar bones are fused together. The pisiform is elongated and expanded at its two extremities; it forms a prominence which, directed backwards, projects beyond the level of the other bones of this region.
The metacarpal bones are five in number; they are enumerated from within outwards; they articulate with the carpus and with each other. The inferior extremity of each metacarpal bone presents the form of a condyle in front; and is divided behind so as to form two lateral condyles, which are separated by a median crest; on these posterior condyles are applied two small sesamoid bones. The metacarpal bone of the thumb is very short; the third and fourth are the longest. The metacarpus, as a whole, is directed vertically.
Fig. 37.—Skeleton of the Finger of a Felide (Lion): Left Side, Internal Surface.
1, Metacarpus; 2, sesamoid bones; 3, first phalanx; 4, second phalanx; 5, third phalanx; 6, gutter for the reception of the base of the nail; 7, prominent osseous crest formed to lodge in the concavity of the nail.
The phalanges are three in number for each finger, except the thumb, which has but two. The first phalanx, directed almost horizontally forwards, is the longest; the second is directed downwards and forwards; the third consists of two portions: a posterior part, which forms a sort of sheath into which the base of the nail is received; and an anterior, conical in form, and curved in crochet shape, which forms a support for the nail (Fig. 37).
The third and fourth digits are the longest; the second and fifth are of equal length; the thumb is the shortest; it does not touch the ground, and does not even reach the articulation of the metacarpal bone and first phalanx of the second finger.
In the cat, the metacarpal bone of the thumb, although shorter than any of the others, is quite as thick. The third digit is a little longer than either the second or fourth. In animals of this genus, the claws, in the condition of repose, are retracted, and removed from the ground; this prevents their being worn, and thus preserves their sharpness. At such times the third phalanx is received into a groove which is found on the external surface of the second phalanx. In the dog, the claws are not tractile.
Fig. 38.—Skeleton of the Pig: Left Lateral Surface.
1, Cranium; 2, face; 3, atlas; 4, axis; 5, seventh cervical vertebra; 6, first dorsal vertebra; 7, fourteenth and last dorsal vertebra; 8, lumbar vertebræ; 9, sacrum; 10, coccygeal vertebræ; 11, anterior extremity of the sternum; 12, xiphoid appendix; 13, seventh and last sternal rib; 14, costal cartilage; 15, cartilage of prolongation of the scapula; 16, great tuberosity of the humerus; 17, olecranon process; 18, radius; 19, ulna; 20, pisiform; 21, metacarpus; 22, phalanges of the two great toes; 23, phalanges of the external toe; 24, ilium; 25, pubis; 26, tuberosity of the ischium; 27, great trochanter; 28, knee-cap; 29, anterior tuberosity of the tibia; 30, fibula; 31, tarsus; 32, calcaneum; 33, metatarsus; 34, phalanges of the two great toes; 35, phalanges of the external toe.
Unguligrades: Pig (Fig. 38).—The scapula is markedly narrowed in the region above the glenoid cavity. The spine is atrophied at both its extremities, so that at its inferior part we do not find the acromion process. In its middle portion the spine is prominent, and presents a triangular process which turns backwards, overlapping a part of the infraspinous fossa; this latter is much larger than the supraspinous. The spinal border is surmounted by the cartilage of prolongation, the superior margin of which is convex; this cartilage extends posteriorly beyond the posterior (axillary) border of the bone.
The small tuberosity of the superior extremity of the humerus is but slightly developed; the great tuberosity, on the contrary, is very large. The bicipital groove is situated internal to this. The deltoid impression is scarcely marked.
The forearm is short, directed obliquely downwards and inwards, thus forming with the hand an angle, of which the apex is directed inwards. The two bones of the forearm are strongly bound to one another by an interosseous ligament, which is formed of very short fibres. The radius appropriates, at its superior extremity, the coronoid process of the ulna. The latter is, notwithstanding, well developed in the rest of its extent; it has a flattened shaft which almost completely overlaps the posterior surface of the radius; its inferior extremity reaches to the carpus.
The carpus is formed of eight bones—four in the superior row, and four in the inferior. The third bone of the superior row (cuneiform) is more in contact with the ulna than with the radius.
There are but four metacarpal bones; there is no metacarpal of the thumb. The two median metacarpal bones are the longest; they are those which correspond to the digits which alone touch the ground. The internal digit and the external one are thin and short; they are functionless, as a rule, taking no part in supporting the limbs on the ground. Notwithstanding this, they are formed, as the other digits, of a number of phalanges, which give them the semblance of perfect digits. (We shall soon see that in certain animals there exist digits which, being incomplete with regard to the numbers of their constituent bones, more accurately merit the name of imperfect digits.)
The third phalanges are each enclosed in a horny hoof, to which the name of onglon has been given.
We have already drawn attention to the smaller lateral digits, and noted the general fact that they do not come in contact with the ground. It is necessary to modify this statement by adding that under certain conditions they give a slight amount of support; for example, when the individual is the subject of excessive obesity, the limbs yield under the weight, and the nails of the lateral digits may touch the ground.
A similar fact may be noticed in pigs of ordinary bulk at the moment when, during walking, each of the fore-limbs commences to bear the weight—that is to say, when it is directed obliquely downwards and forwards; then all the digits are in contact with the ground.
Fig. 39.—Skeleton of the Ox: Left Lateral Surface.
1, Cranium; 2, face; 3, atlas; 4, axis; 5, seventh cervical vertebra; 6, first dorsal vertebra; 7, thirteenth and last dorsal vertebra; 8, lumbar vertebræ; 9, sacrum; 10, coccygeal vertebræ; 11, sternum; 12, xiphoid appendix; 13, eighth and last sternal rib; 14, costal cartilages; 15, spine of scapula; 16, cartilage of prolongation of the scapula; 17, great tuberosity of the humerus; 18, musculo-spiral groove; 19, olecranon process; 20, radius; 21, carpus; 22, pisiform; 23, metacarpus; 24, rudimentary metacarpal; 25, sesamoid bones; 26, first phalanges; 27, second phalanges; 28, third phalanges; 29, anterior iliac spine; 30, pubis; 31, tuberosity of the ischium; 32, great trochanter; 33, supracondyloid fossa of the femur; 34, patella, or knee-cap; 35, anterior tuberosity of the tibia; 36, fibula; 37, coronoid tarsal bone; 38, tarsus; 39, calcaneum; 40, metatarsus; 41, rudimentary metatarsus; 42, sesamoid bones; 43, first phalanges; 44, second phalanges; 45, third phalanges.
Unguligrades (Ungulates): Sheep, Ox (Fig. 39).—The scapula, which is of elongated form, is very narrow in the vicinity of the glenoid cavity. The spine, which becomes more and more salient towards its inferior part, terminates abruptly in a border, which, forming an acute angle with the crest, produces a projection which represents the acromion process—a very rudimentary acromion, for it does not reach the level of the glenoid cavity. The supraspinous fossa is much smaller than the infraspinous; it hardly equals one-third the extent of the latter. The anterior border, thin and convex in its superior portion, is concave in the rest of its extent; the posterior border is thick and slightly concave; the spinal border is surmounted by the cartilage of prolongation. In the ox the spine of the scapula, in its middle portion, is flexed a little backwards on the infraspinous fossa.
The great tuberosity of the humerus is highly developed; its summit, very prominent, is flexed over the bicipital groove; a prominence of the small tuberosity also bends over the groove, with the result that at this level the latter is converted into a sort of canal. At the inferior extremity the condyle, although not large, is recognisable; for it is separated from the trochlea by a depression in form of a groove. In contrast to the condition found in man, the condyle descends to a level a little below that of the internal lip of the trochlea. (For the arrangement of the epicondyle and the epitrochlea, see p. 30.) In the sheep, the deltoid impression is but slightly marked; in the ox, it is more evident.
The forearm is directed obliquely downwards and inwards, so as to form, with the hand, an angle of which the apex is internal; this angular outline of the knee (wrist) is so characteristic of ruminants that the corresponding region of the horse, when salient inwards, receives the name of ox-knee. The radius bears the coronoid process, and the larger part of the articular surface which comes in contact with the inferior extremity of the humerus; the condyle and the trochlea articulate with the radius in front; while behind, the trochlea articulates with that part of the sigmoid cavity which belongs to the ulna. The posterior surface of the shaft of the radius is flattened; its anterior surface is slightly convex. The inferior extremity articulates with the carpus by a surface which is directed obliquely downwards and inwards. The shaft of the ulna is very slender, and fused in its middle third with the body of the radius; it terminates below, at the level of the external part of the inferior extremity of the radius, by a slightly expanded portion which, fused with this latter, forms the articular surface for the carpal bones.
In the ox the forearm is short; in the sheep it is proportionally longer.
The bones of the carpus are six in number—four in the upper row, and two in the lower; they form an irregular cuboid mass which contributes to the formation of the region known as the knee in ruminants, as in the horse; we have already remarked that the name ‘wrist’ would be more accurate. The anterior surface in its foremost part is vertical, and is slightly convex from side to side. At its posterior and external part the pisiform bone forms a prominence.
The metacarpus consists of two bones only—one, well developed, which is known as the principal metacarpal, or the canon bone (this is the name given to the region in the hoofed animals); and a rudimentary one, which is situated at the superior and external aspect of the preceding metacarpal. Sometimes there is found a third metacarpal at the internal aspect; but, when present, it is but very slightly developed.
The principal metacarpal consists of two metacarpals fused together; on this account the bone is longitudinally marked in the median line by a slight depression which marks the junction of the two bones of which it is formed. In some ruminants (certain species of chevrotains) the coalescence does not take place, and the two metacarpals remain separate.
The anterior surface of the principal metacarpal is convex transversely; its posterior surface is flattened. The superior extremity of this bone articulates by two facets with the two bones of the inferior row of the carpus; on the internal part of the anterior surface of this extremity is found a tubercle. The inferior extremity is divided into two parts by a fissure or notch; each part is articular, and consists of two separate condyles, which are separated from each other by an antero-posterior crest; on each side of this crest, and behind, are found two sesamoid bones. As for the external rudimentary metacarpal bone, it is nothing more than a small, short tongue of bone; which, in goats and sheep, is often absent.
The division of the inferior extremity of the principal metacarpal into two parts is correlated with the two perfect digits which give the foot of the ruminant its forked appearance. Each digit consists of three phalanges, which are directed obliquely downwards and forwards; further, these phalanges are inclined a little outwards from the axis of the limb, so that the two digits diverge from each other as they descend.
The first phalanx, which is the longest, articulates superiorly with the principal metacarpal; its inferior extremity terminates in a trochlea, and the lip of this, which is situated towards the axis of the limb, descends lower than that of the opposite side; this arrangement is correlated with the divergent direction of the digits. The second phalanx has its superior extremity moulded on the trochlea which terminates the extremity of the first; its inferior extremity is articular, and elongated from before backwards. On the posterior surface of this extremity is found a sesamoid bone.
With regard to the third phalanx, it presents the form of a triangular pyramid, and displays a postero-superior concave surface with which the second phalanx articulates; an anterior, convex surface, which terminates in a point on its anterior part; and an internal surface, which is flattened. The third phalanx of each digit is contained in a hoof (onglon).
There is also found in ruminants two imperfect rudimentary digits, which are represented by two small bones situated behind the articulation of the metacarpal and the digits which we have just been studying. These rudimentary digits are each enveloped in a layer of horn; they constitute the spurs. The two digits of the ruminants represent the third and fourth fingers of the human hand; the two lateral digits, greatly atrophied, are the homologues of the second and fifth fingers; the thumb is not present.
It is the same as regards the metacarpal bones, which form, by their union, the principal metacarpal; the external represents the fourth metacarpal, and the internal the third. It is to the latter that the tubercle, of which we have already made mention, belongs; and with the signification of which, because it gives attachment to a muscle, we shall concern ourselves in the section on myology (see Radial Muscles).
Fig. 40.—Skeleton of the Horse: Left Lateral Surface.
1, Cranium; 2, face; 3, atlas; 4, axis; 5, seventh cervical vertebra; 6, first dorsal vertebra; 7, eighteenth and last dorsal vertebra; 8, lumbar vertebræ; 9, sacrum; 10, coccygeal vertebræ; 11, sternum; 12, xiphoid appendix; 13, eighteenth and last sternal rib; 14, costal cartilage; 15, scapula; 16, cartilage of extension; 17, great tuberosity of the humerus; 18, deltoid crest; 19, olecranon process; 20, radius; 21, carpus; 22, pisiform; 23, principal metacarpal; 24, metacarpal, external rudimentary; 25, large sesamoids; 26, first phalanx; 27, second phalanx; 28, third phalanx; 29, ilium, showing external iliac fossa; 30, pubis; 31, tuberosity of the ischium; 32, great trochanter; 33, infratrochanteric crest, or third trochanter; 34, supracondyloid fossa of the femur; 35, knee-cap; 36, anterior tuberosity of the tibia; 37, the fibula; 38, tarsus astragalus; 39, calcaneum; 40, principal metatarsal; 41, rudimentary external metatarsal; 42, large sesamoids; 43, first phalanx; 44, second phalanx; 45, third phalanx.
To face p. 64.
Unguligrades: Horse (Fig. 40).—The scapula is narrow, compared with that of the animals we have just been considering. The anterior border is convex in its superior portion, and concave in its inferior; the posterior border is slightly hollowed out. The supraspinous fossa is less in extent than the infraspinous; but the difference is less than that between the same fossæ in the ox and the sheep; in the ox, as we have already indicated, the proportion is one-third; in the horse, one-half. The spine, which disappears at the extremities, is rough and thick in its middle third, there forming a kind of tuberosity—tuberosity of the spine. Above and in front of the glenoid cavity is found a strong process consisting of a rugous base, and a summit which is directed inwards. This forms a kind of hook curved towards the inside; it represents the coracoid process. The scapula is surmounted by the cartilage of prolongation, of which the superior border, which is thin and curved, is parallel to the superior border of the prominence of the withers; the cartilage forms, consequently, the lateral surface of this region. The cartilage of prolongation undergoes ossification in old horses. The humerus is short; the bicipital groove, situated on the anterior surface of the superior extremity, separates the greater tuberosity from the lesser, and is divided into two parts by a median ridge; it is this portion of the humerus which forms the prominence known as the point of the shoulder, or point of the arm. The deltoid impression well deserves the name of tuberosity which has been given to it, for it is very prominent; the musculo-spiral groove is very deep.
At the inferior extremity, the trochlea is large; the portion corresponding to the condyle of the humerus in man is, in proportion to the latter, of small extent. The olecranon fossa is deep. The epicondyle and the epitrochlea are somewhat different from those of the human bone. In the latter, the epitrochlea is salient towards the inner side, causing an increased transverse diameter of the inferior extremity of the humerus. In the horse—it is the same in ruminants—this tuberosity projects backwards, folds on itself in forming the internal boundary of the olecranon cavity, and exceeds in diameter, in the antero-posterior direction, the prominence of the epicondyle, which presents a nearly similar arrangement. This latter has, however, a part which, projecting externally, is situated at the inferior part of a crest, that forms the posterior boundary of the musculo-spiral groove. The result is that, contrary to the condition found in the human being, the epicondyle is more prominent transversely than the epitrochlea, but this latter is more salient on the posterior aspect. The epitrochlea and the epicondyle offer a larger surface for the origin of muscles of the forearm than the same prominences in the human bone do for the analogous muscles of the same region.
Some veterinary anatomists have given to the inferior and external articular surface of the humerus the name of trochlea; and to the internal one, that of condyle. On this account they designate the external prominence as the epitrochlea, and the internal one as the epicondyle. In addition to the fact that this point of view is not legitimate, it produces inevitable confusion when comparing the parts with those of the human humerus, and this confusion exists, not alone in describing the bone, but also in the description of the muscular attachments, and in the comparison of the muscles of the forearm of quadrupeds with the corresponding muscles in the human species.
The radius is placed in front of the ulna; its body, slightly convex forwards, has the anterior surface convex transversely, and the posterior surface plane in the same direction. It is to the external part of this latter that the ulna is applied, which is completely fused with the radius.
The superior extremity of the radius is a little larger than the inferior. Its superior aspect, concavo-convex, moulded on the inferior articular surface of the humerus, presents internally two cavities, which receive the lips of the trochlea, and, externally, another, smaller, cavity, which receives the condyle. The radius articulates with the trochlea and the condyle, having appropriated a portion of the ulna, as is proved by the presence of the coronoid process, which belongs to the former. This superior extremity presents, internally, a tuberosity into which the biceps is inserted; this is the bicipital tuberosity; and on the other side is another tuberosity, which is a little more prominent than the preceding.
The inferior extremity, which is flattened from before backwards, is furrowed on its anterior surface by grooves for the passage of muscles (the names of the muscles whose tendons pass in these grooves have already been given on p. 43). It articulates at the lower end with the superior row of the carpus, and it terminates laterally in tuberosities: one, external, on which is found a groove for the tendon of the lateral extensor of the phalanges, the homologue of the special extensor of the little finger; the other, internal, is a little more prominent than the one we have just described. These tuberosities are visible under the skin which covers the superior and lateral parts of the region known as the knee; but which, we again repeat, is no other than the wrist.
The ulna has a triangular shaft, situated at the posterior surface of the radius, with which it is fused. It disappears completely at the level of the inferior third of the forearm. Occasionally, in some horses, the ulna is abnormally long, in the form of a slender tongue of bone; and extends to the neighbourhood of the external tuberosity of the inferior extremity of the radius (see Fig. 79, p. 196). Its superior extremity is chiefly represented by the olecranon process, which is voluminous in bulk, and forms the projection known as the point of the elbow. This process is flattened laterally; its internal surface is excavated; the anterior surface, which is concave, forms a part of the great sigmoid cavity; the remainder of the cavity is formed by the radius.
In the ass, the ulna is a little longer than in the horse—that is to say, it descends lower; and the radius is a little more convex anteriorly.
The carpal bones are seven in number—four in the superior row, and three in the inferior. The trapezium is wanting in the latter. Sometimes, however, in certain varieties of horses the trapezium is developed, but then it is no more than a very small osseous nodule. The pisiform bone, situated at the external part of the first row of bone, is prominent posteriorly. It is of rounder form and flattened from without inwards. It articulates with the trapezium and the radius. It presents, on its external surface, a groove for the passage of the tendon of the posterior ulnar muscle, which is named by veterinary anatomists the external flexor of the metacarpus.
The carpus, as a whole, is of an irregularly cuboid shape; its anterior surface, slightly convex from side to side, forms the skeleton of the region of the knee (wrist). The metacarpus is formed of three bones: the principal metacarpal and the two rudimentary ones.
The principal metacarpal, which forms the region of the canon, is directed vertically; its anterior surface is slightly convex transversely. This surface is covered by a number of tendons, which slightly alter its appearance; so that it is the principal base of this part of the fore-limb. Its posterior surface is flattened. The superior extremity of this metacarpal presents plane surfaces, variously inclined, with which the bones of the inferior row of the carpus articulate. On the anterior surface, and a little to the inner side, is found a tuberosity, which is destined for the insertion of the anterior extensor of the metacarpus, the homologue of the radial muscles. The inferior extremity is formed by two condyles, an internal and an external; between which is found a median crest.
This extremity, the superior extremity of the first phalanx, which articulates with it, together with two sesamoid bones—the great sesamoids—which are situated on its posterior surface, collectively form the region which from its rounded outlines is called the ball.
With regard to the rudimentary metacarpals, external and internal, to which some authors give the name of fibulæ, they are applied to the sides of the posterior surface of the principal metacarpal. They are elongated bones, of which the superior extremity, which is a little thickened, is called the head; the lateral bones of the second row of the carpus partly rest on the heads of these. They become more slender as they descend, and terminate opposite the inferior fourth of the principal metacarpal. Each ends in a slight swelling, to which the name button has been given. The internal one is the better developed.
The rudimentary metacarpals are vestiges of atrophied digits, as will be explained further on.
The single finger of the horse consists of three phalanges. The first phalanx, which is directed obliquely downwards and forwards, corresponds to the constricted region situated below the ‘ball,’ and known as the pastern. It is flattened from before backwards; its anterior surface is convex transversely, while the posterior surface is plane. Its superior extremity is moulded on the inferior extremity of the principal metacarpal, and its inferior extremity, which is smaller, presents a trochlea with which the second phalanx articulates. This is also directed downwards and forwards, and is shorter. It corresponds to the region which, situated between the pastern and the hoof, is known as the cornet.
The third phalanx, situated entirely within the hoof, has the same direction as the first and second. It is large and broad, and presents three surfaces separated by well-marked angular borders (see Fig. 96). The anterior surface is oblique downwards and forwards; it is convex transversely. The inferior surface is slightly hollowed, and is in relation with the sole, or plantar surface of the hoof.
The superior surface, which is articular, is divided by a median ridge into two lateral cavities, which correspond to the trochlea on the inferior surface of the lower extremity of the second phalanx. The inferior border corresponds in shape with the hoof. The superior border presents in its median part a projection, the pyramidal eminence, which prolongs at this level the anterior surface of the bone. Finally, the posterior border, which is concave, is in contact with a sesamoid bone, the lesser sesamoid, which increases the superior articular surface behind, and is also in contact with the second phalanx.
As we have just seen, the horse possesses but one digit. In the ancestors of the animal—that is, in the prehistoric species which are now extinct (orohippus, miohippus, protohippus, or hipparion)—the number of digits was larger; this fact conclusively proves that the rudimentary metacarpals of the existing horse are vestiges of digits which have disappeared through want of use. In the first of those ancestors—orohippus—there were four digits; all save the first, the thumb, being then developed. In the others of the series there existed but three digits. It must, however, be noted that in those animals it is always the digit which corresponds to the middle finger of the pentedactyl hand that is longest. In other less ancient species the lateral fingers are reduced to the condition of mere splints of bone. It follows from what has been said that the digit which persists in the equine species should be considered as the third finger, and that the rudimentary metacarpals represent lateral digits considerably atrophied.
This disappearance of the lateral digits cannot excite surprise when we consider the functions of the organs. Becoming useless, they must undergo gradual atrophy from want of use.
There undoubtedly is, in this former existence of supplementary digits in the horse, something analogous to what we still find in the pig; where the two principal digits are accompanied by two shorter ones, which very probably, from their infrequent use, are destined to disappear in a more or less distant future.
As a supplement to the study of the anterior limbs which we have just finished, it appears necessary to give some indications of the relative proportions of certain of the segments which form these limbs in the plantigrades, the digitigrades, and the ungulates.
First, we would remark that, in following this order of classification, the scapula becomes less and less narrow, and assumes a form more and more elongated. In order to convince ourselves of this, it will be sufficient to study the bone first in man, then in the bear, the cat, dog, ox, and finally in the horse.
As to the proportions of length, which are those we should chiefly study, we shall commence with the comparison of the forearm and arm—that is to say, the radius and the humerus. The radius is found to be longer in proportion to the humerus, as the number of digits is smaller, and the hand loses more and more the functions of an organ of prehension. In man, the radius is shorter than the humerus; in the horse, on the contrary, it is longer.
To give an idea of this proportion, we shall employ what is known as the antibrachial index. This index gives the relation which exists between the length of the forearm and that of the humerus; the length of this latter, whatever may be the actual measurement, is represented by a fixed figure, the number 100. A very simple arithmetical operation gives the proportion—
| forearm × 100 | , the quotient obtained furnishes the index. |
| humerus |
The index is less than 100 if the forearm is shorter than the bone of the arm. The index is more than 100 if, on the contrary, the forearm is longer.
In man, the radius is shorter than the humerus; indeed, in adult individuals of the white race the average index is 74.
In the bear, the length of the radius approaches closely to that of the humerus; the index is about 90. In the skeleton of a bear in the anatomical museum of the École des Beaux-Arts, the humerus is 33 centimetres in length, and the radius 30 centimetres.
In the cat, the radius is very little shorter than the humerus. In the dog they are equal. The antibrachial index of the latter is, accordingly, 100.
In the horse, the radius is longer than the humerus; the index is therefore above 100. Thus, in the skeleton of the horse which we have in the museum of the École des Beaux-Arts, the index is 113—length of humerus, 29 centimetres; length of radius, 33 centimetres. In other skeletons which we have measured we found: in one, 108—humerus, 34 centimetres; radius, 37 centimetres; in another, 116—humerus, 25 centimetres; radius, 29 centimetres.
The metacarpal bone undergoes, relatively to the humerus, a proportional elongation, analogous to that of the forearm.
In man, the length of the metacarpus is contained about 51⁄2 times in that of the humerus; in the bear, it is contained 4 times; in the dog, 21⁄2 times; in the horse, 11⁄3 times only.
It is well known that the proportions vary according to race, and that what we have here given are but the general indications.
The knowledge of human arthrology which we presume the reader to have previously acquired makes it unnecessary for us to enter into numerous details regarding the configuration of the articular osseous surfaces and the disposition of the fibrous bands that retain them in position. Accordingly, in the description which follows, and also in that of the articulations of the posterior limbs, we shall occupy ourselves but very briefly with the details above referred to, so as to devote ourselves especially to the indication of the movements—that is to say, of that which, while easily comprehended on recollection of former studies, presents the greatest interest from the artistic standpoint in these studies in comparative anatomy.
The Scapulo-Humeral Articulation.—The head of the humerus and the glenoid cavity of the scapula being in contact, the two bones are bound together by a rather loose articular capsule, which is strengthened by the muscles of this region which fulfil the function of active ligaments.
This articulation, so movable in every direction in the human species, is not so much so in quadrupeds; the arm in the latter, as also the shoulder, being kept in contact with the lateral region of the thorax by the numerous muscles which surround it.
Of the movements performed by the humerus, flexion and extension are the most extensive; those of abduction and adduction are much less so.
It is necessary, before proceeding further, to determine what the two principal movements which we have just mentioned really are, viz., flexion and extension.
We know that in man the displacements of the humerus which take place in the antero-posterior direction are known as movement or projection forwards, and movement or projection backwards, respectively. We do not say that the humerus is flexed or extended, because, in reality, on account of the position which the skeleton of the shoulder occupies, it is not able to flex or place itself on the line of prolongation of the scapula with which it articulates.
In quadrupeds it is not so. The humerus and the scapula are contained in almost the same vertical plane; and the bone of the arm can take, in relation to the latter, the positions characteristic of flexion and extension—that is, of approach to the scapula and removal from it.
What makes the meanings of these terms a little confusing is that, in human anatomy, some authors consider the backward movement of the humerus as extension, and the forward movement as flexion; in order to be able to compare these movements to those that the femur executes in relation to the pelvis.
Now, in our opinion, the indication of this correspondence is not absolutely necessary; since it ceases to be exact if we wished, from the point of view of the direction given to other segments of the skeleton, to establish the same relation between the elbow and the articulation of the knee.
It is therefore indispensable, when discussing quadrupeds, to discontinue these terms, in order the more readily to recognise that: in flexion the inferior extremity of the humerus is directed backwards; in extension, on the contrary, it is directed forwards. In the first case the humerus approaches the scapula; in the second, on the contrary, it moves away from it.
These movements, which take place during walking, are executed in the following manner: When one of the anterior limbs is at the end of that stage of progression which is called support (see p. 289, Displacements of the Limbs)—that is to say, during the time that the foot remains in contact with the ground, whilst the trunk is moving forward—the direction of this limb becomes more and more oblique downwards and backwards. At a certain moment the limb is raised from the ground, to be carried forwards, in order to be again pressed on the ground, and recommence a new resting stage. In these different phases the humerus is flexed. But at the moment that the limb, when carried forwards, is about to resume its contact with the ground it becomes directed obliquely downwards and forwards; then the humerus is in the position of extension.
During these movements of the humerus, there exists an essential factor—that is, the scapular balance. (It is the same as what occurs in man when he balances his arm in the antero-posterior plane.) When the humerus is flexed, the scapula moves in such a way that the superior portion projects forwards; when it is extended, the scapula, on the other hand, is inclined more backwards. But it is necessary to add that, during these displacements, the scapulo-humeral angle varies; it tends to close during the flexion of the humerus, and becomes more open during extension.