Fig. 15.—Vertical and Transverse Section, at the Plane of the Shoulders, of the Thorax of a Dog (Diagrammatic Figure).
1, Outline of the thorax at the level of the third dorsal vertebra; 2, 2, scapula; 3, spinal border of the scapula; 4, contour of the skin.
In quadrupeds whose scapula, on the contrary, is wanting in the cartilage of prolongation (in the clawed,[7] such as the cat and dog), the superior border of the scapula is visible, especially when the animal is resting on its fore-limbs, particularly when it crouches; at such a time the skin is markedly raised by that border; and the spinous processes of the vertebræ, beyond which it projects, occupy the bottom of a fossa (Fig. 15). The internal surface of the scapula is turned towards the ribs; it is known, as in man (in whom this surface is anterior), as the subscapular fossa.
Fig. 16.—Left Clavicle of the Cat: Superior Surface (Natural Size).
1, Internal extremity; 2, external extremity.
Its external surface is divided into two parts by the spine of the scapula; which, in some animals, terminates inferiorly in a flat and clearly distinct process, the homologue of the acromion process of the human scapula. The two regions separated by the spine are known as the supraspinous fossa and the infraspinous fossa. The supraspinous fossa is anterior to the spine, and the infraspinous is posterior to it. The surfaces of the scapula are, in quadrupeds, flatter than in the human being, and in particular the subscapular fossa, which is also less concave. Some authors attribute this to the lesser curvature of the ribs in quadrupeds. A few words will suffice to prove that there must be another reason. The scapula is not in immediate contact with the ribs; the subscapular fossa is not moulded on them. Besides, the form of the scapula is, as in other parts of the skeleton, dependent on the disposition of muscles, and the development of these latter is correlated to the extent and energy of the movements which the individual is able or required to execute. But the movements which those muscles produce (more especially the rotation of the humerus) are, in quadrupeds, less extensive than in the human being; and, consequently, the muscles which produce them are, proportionally, less strongly developed. The inferior angle (superior and external in man), situated at the junction of the cervical and axillary borders, presents the glenoid cavity, which, looking downwards, receives the articular surface of the superior extremity of the bone of the arm—that is to say, the head of the humerus. Above this cavity, on the lower part of the cervical border, is situated a tubercle which reminds us of the coracoid process of the human scapula. The region occupied by the glenoid cavity is separated from the body of the bone by a constriction—the neck of the scapula.
Fig. 18.—Skeleton of the Shoulder of a Bird (Vulture): Antero-External View of the Left Side.
1, Left clavicle; 2, inferior portion of the right clavicle, forming by its ankylosis with that of the other side the fourchette; 3, coracoid bone; 4, scapula; 5, articular surface for humerus; 6, superior half of the sternum; 7, keel of sternum; 8, spinous process of the dorsal vertebræ; 9, superior ribs; 10, process of one of these ribs; 11, inferior ribs.
In birds the scapula is elongated in a direction parallel to the vertebral column, and very narrow in the opposite (Fig. 18): it is also flat, and has no spine. Its coracoid process is represented by a peculiar bone—the coracoidean or coracoid bone—which we shall describe later on when we come to the study of the clavicle and of the anterior region of the shoulder (see p. 26).
The Clavicle.—The clavicle is found only in the human being, and in animals whose anterior limbs, possessing great freedom of movement in all directions, require that the scapula should possess a point of support which, while affording this, can be displaced with it, or draw it in certain directions. Now, this point of support is furnished by the clavicle.
In animals possessed of hoofs (ungulates), such as the sheep, ox, and horse, the clavicle does not exist. Indeed, in them the freedom of movement of the anterior limbs is limited; they move by projection in the forward and backward directions only; they merely fulfil the functions of giving support to and carrying about the body. The clavicle is rudimentary in the cat and the dog; in the cat it is a small, elongated bone (Fig. 16), 2 centimetres in length, thin and curved, connected with the sternum and the scapula by ligamentous bundles. In the dog it is represented by a small osseous plate only (Fig. 17), which is not connected with any of the neighbouring bones.
It is on the deep surface of a muscle which passes from the head and neck to the humerus (mastoido-humeral, a muscle common to the arm, neck, and head) in which this rudimentary bone is found to be developed.
The clavicle exists in perfect state in mammals which use their limbs for digging, grasping, or flying; the insectivora (hedgehog, mole) and some rodents (squirrel, woodchuck) are provided with it.
The cheiroptera (bats) possess an extremely well-developed clavicle, on account of the varied movements which their thoracic limbs execute.
This formation of the shoulder which favours flight in the bat is even more remarkable in birds. In these latter (Fig. 18) the clavicles, fused together by their lower extremities, form one bone, having the shape of the letter V or U, which is known as the fourchette; this bone, acting as a true spring, keeps the shoulders apart, and prevents their approximation during the energetic movements which flight necessitates.
In birds whose power of flight is strong, the two limbs of this bone are widely separated and thick, and the fourchette is U-shaped. Those whose flight is awkward and but slightly energetic have the limbs of the fourchette slender; they unite at a more acute angle, and the bone is V shaped.
Furthermore, a bone named the coracoid joins the scapula to the sternum; this bone, often fused with the scapula, where it contributes to the formation of the glenoid cavity, represents in birds the coracoid process of the human scapula. If we fancy this process directed inwards, and sufficiently lengthened to join the sternum, we shall have an idea of the disposition of the bone we are now discussing, and the reasons for which the name has been chosen by which it is designated. The coracoid bone, like the fourchette which it reinforces, offers to the wings a degree of support proportionate to the efforts developed by those limbs; for this reason it is thick and solid in birds of powerful flight.
The superior extremity of each branch of the fourchette, at the level of its junction with the coracoid and the scapula, bounds, with these latter, a foramen which gives passage to the tendon of the elevator muscle of the wing, or small pectoral. The importance of the fourchette being, as we have seen, in proportion to the movements of flying, it is easy to understand that the bone is not found in the ostrich.
A single bone, the humerus, forms the skeleton of this portion of the thoracic limb.
The Humerus.—The bone of the arm is, in quadrupeds, inclined from above downwards and from before backwards.
It is, with relation to other regions, short in proportion as the metacarpus is elongated, and as the number of digits is lessened. In the horse, for example, whose metacarpus is long, and in which but one digit is apparent, the humerus is very short. The slight development in length of the humerus explains its close application to the side of the animal as far as the elbow.
In animals in which the humerus is longer, the bone is slightly free, as well as the elbow, at its inferior extremity. Later on we will return to the consideration of this peculiarity and of the proportions of the humerus, after we have studied the other parts of the fore-limbs.
The humerus in quadrupeds is inflected like the letter S; in man this general form is less accentuated, the humerus being almost straight. On its body, which appears twisted on its own axis, we find the musculo-spiral groove,[8] which crosses the external surface, and is very deep in some animals. Above this groove, and on the external surface, there exists a rough surface which is the impression of the deltoid. In some species this rugosity is very prominent, and is called the tuberosity of the deltoid; it is prolonged downwards by a border which forms the anterior crest of the musculo-spiral groove and limits this latter in front. The external border of the bone, or posterior crest of the groove, limits it behind.
[8] It would be going outside our province to discuss whether the humerus is really twisted on its axis. This question, often discussed, has been solved in some recent works in the following manner: the humerus has undergone torsion at the level of its superior extremity, and not at the level of its body; this does not authorize us further to accord any definite sense to the denomination ‘groove of torsion’ (musculo-spiral groove). That which we must especially remember in connection with this fact, is, as we shall afterwards see, the difference of direction which the articular head presents according as the torsion has been more or less considerable: because this is established, according to the same order, in man and in quadrupeds.
The superior extremity is enlarged, and remarkable in three portions which it presents; these are: an articular surface and two tuberosities.
The articular surface, or head of the humerus, smooth and round, is in contact with the glenoid cavity of the scapula. This head in the human skeleton is directed upwards and inwards; in quadrupeds its direction is upwards and backwards. The inferior extremity, having in both one and the other its long axis directed transversely, and the point of the elbow looking backwards in all, the result is that the head of the humerus is not situated vertically above the same regions; in the first, it is almost directly above the internal part of this extremity; in the latter, it is situated above its posterior surface, or the point of the elbow in the complete skeleton. This difference of direction is correlated with the position of the scapula, the glenoid cavity of which, as we have already seen, is in man turned outwards, whereas in quadrupeds it looks downwards. In the latter case the scapula consequently rests on the head of the humerus; and this position is most favourable for the performance of the functions which the anterior limbs have to fulfil in these latter.
Of the tuberosities of the head of the humerus, one is situated on the external aspect—it is the great tuberosity, or trochiter; the other is placed internally—it is the small tuberosity, or trochin. The great tuberosity is divided into three parts—summit, convexity, and crest; these different parts give insertion to the muscles of the shoulder. We recollect that the facets (anterior, middle, and posterior) of the great tuberosity of the humerus in man give attachment to the muscles of the same region. The head of the humerus in the human body projects above the tuberosities. We shall see afterwards, when dealing with some special quadrupeds, that in some of these, on the other hand, the tuberosities are on a higher level than the articular head of the bone. Between the two tuberosities is the bicipital groove.
In man, the superior extremity of the humerus, although covered by the deltoid, reveals its presence by elevating the corresponding portion of the latter. In quadrupeds, the anterior part of this extremity, although similarly covered by muscular bundles, produces a prominence under the skin. This prominence is situated at the summit of the angle formed by the opposing directions of the scapula and the bone of the arm, and constitutes what is known by the name of the point of the shoulder, or of the point of the arm.
The inferior extremity, transversely enlarged, presents an undulating articular surface, which reminds us of the trochlea and the condyle of the human humerus; on which, however, the condyle is more sharply defined from the trochlea.
In the human skeleton, the internal lip of the trochlea descends lower than the external; and also lower than the condyle. In the bear, the cat, and the dog, it is the same. In the ox and the sheep, the condyle is lower than the trochlea, but only very little lower. In the horse the arrangement is still the same, but a little more accentuated.
On the lateral parts of this extremity we find: internally, a prominence, the epitrochlea; and, externally, another, the epicondyle. It is from this latter that the crest arises, which, passing upwards, forms the posterior limit of the groove of torsion.
The two prominences, which we have just described from a general point of view, present special arrangements which it is necessary to point out. When we examine the form of the outline of the inferior extremity of the humerus in man, the bear, the cat, the dog, the ox, and the horse, we find in following this order that the extremity tends to become narrow transversely, and that the epicondyle and the epitrochlea are less and less prominent on the external and internal aspects respectively. These two processes, indeed, project backwards; the epitrochlea always remaining more developed than the epicondyle. Because of this projection backwards, the cavity situated on the posterior surface of the inferior extremity, the olecranon fossa, is very deep, more so than in the humerus of man. Its borders being thus formed by the two processes, are very prominent. In front we find the coronoid fossa, which is less deep than that of which we have just spoken.
Fig. 19.—Inferior Extremity of the Left Humerus of a Felide (Lion).
1, Epitrochlea; 2, supra-epitrochlear foramen.
There exists in some mammals an osseous canal, situated above the epitrochlea, and known as the supratrochlear canal (Fig. 19). It is bounded by a plate of bone which at its middle portion is detached from the shaft of the humerus, and blends with the latter at both its extremities. The brachial artery and median nerve pass through the foramen.
Fig. 20.—Inferior Extremity of the Left Human Humerus, showing the Presence of a Supratrochlear Process.
1, Epitrochlea; 2, supra-epitrochlear process.
A similar condition is sometimes found, as an abnormality, in man, which presents itself under the following aspect (Fig. 20): an osseous prominence more or less long, in the shape of a crochet-needle—supra-epitrochlear process—situated 5 or 6 centimetres above the epitrochlea; the summit of this process gives attachment to a fibrous band, which is inserted by its other end into the epitrochlea and the internal intermuscular aponeurosis. The fibro-osseous ring thus formed gives passage to the brachial artery and the median nerve, or in case of a premature division of this artery to the ulnar branch of the same.[9]
[9] For further details of this anomaly, see Testut, ‘The Epitrochlear Process in Man’ (International Journal of Anatomy and Physiology, 1889); A. Nicolas, ‘New Studies on the Supratrochlear Process in Man’ (Review of Biology of the North of France, t. iii., 1890-1891).
There is also found in some mammals a perforation of the thin plate of bone which, in others, separates the olecranon fossa from the coronoid. This perforation is sometimes found as an abnormality in the human humerus.
Fig. 21.—Skeleton of a Bird (Vulture): Left Surface.
1, Cranium; 2, face; 3, cervical vertebræ; 4, spinous processes of the dorsal vertebræ; 5, coccygeal vertebræ; 6, sternum; 7, keel; 8, superior ribs; 9, inferior ribs; 10, clavicle; 11, coracoid bone (for the details of the skeleton of the shoulder, see Fig. 18); 12, humerus; 13, radius; 14, ulna; 15, carpus; 16, hand (for details of the skeleton of this region, see Fig. 31); 17, ilium; 18, ischium; 19, pubis (for the details of the pelvis, see Fig. 46); 20, femur; 21, tibia; 22, fibula; 23, osseous nodule, which some anatomists think represents the calcaneum; it is the sole vestige of the tarsus; 24, metatarsus; 25, foot; 26, first toe (for the details of the skeleton of the foot, see Fig. 48).
As does the sternum and the skeleton of the shoulder, the humerus of birds presents differences correlated to the functions which the thoracic limbs are destined to fulfil. Lying on the side of the thorax, directed obliquely downwards and backwards (Fig. 21), it is proportionately longer in individuals of powerful flight than in those which fly less or not at all. In the vulture it projects beyond the posterior part of the pelvis; in the cock it does not even reach the anterior border of the same. To these differences in length are added differences in volume and in the development of the processes which serve for muscular attachment, which are more considerable in birds of powerful flight.
The humerus is so placed that the radial border, external in man and quadrupeds, looks upwards, with the result that the surface of the bone of the arm, which in these latter is anterior, in the former looks outwards. The humeral head, which is turned forwards and a little inwards, is convex and elongated in the vertical direction. Behind and above this head is found a crest for the insertion of muscles. It is the same for the region below, where there is a tuberosity whose inferior surface presents a pretty large opening which looks inwards to a fossa from the floor of which a number of minute openings communicate with the interior of the bone. This is the pneumatic foramen of the humerus.
It is of interest to remember in connection with this subject that in birds, in keeping with the conditions of flight, every system of organs is adapted to diminish the weight of the body. We particularly draw attention to the osseous framework, the structure of which is such that the weight of the animal is greatly lessened. This condition is secured by the pneumaticity. The bone consists of a cover of compact tissue, which, instead of enclosing marrow, is hollowed out by cavities which contain air, and communicate with special pouches, the air-sacs, which are appendages of the lungs.[10]
[10] The presence of air in the bones does not seem to be always associated with the power of flight; as a matter of fact, we find air spaces in the bones of some birds which do not fly (E. J. Marey, ‘The Flight of Birds,’ Paris, 1890, p. 51).
The antibrachial extremity of the humerus is flattened from without inwards. It terminates in two articular surfaces, which articulate with the radius and ulna.
The olecranon process of the ulna being slightly developed, it follows that the olecranon fossa is not large; neither is the coronoid.
We now proceed to the study of the two regions of the fore-limbs which present the greatest variety in regard to the number of bones and also in regard to form and proportions. These two regions are the forearm and the hand.
It is first of all necessary to say that in man, when the fore-limb hangs beside the body, and the dorsum of the hand looks backwards, the two bones of the forearm are parallel, and that this position is known by the name of supination. It is also necessary to remember that there is another attitude, in which the radius, crossing the ulna, and carrying the hand with it, displaces the latter in such a way that the palmar surface looks backwards. This second position is known as pronation.
Let us now suppose that a man wishes to walk in the attitude of a quadruped. It will be necessary, in order that his upper limbs, being for the moment anterior ones, may act as members of support, to place the forearm in pronation, in order that, as is more normal, the hands may rest on the ground by their palmar surfaces. In this position the radius, being rotated on its own axis at its upper extremity and around the ulna in the rest of its extent, shall have its inferior extremity situated on the inner side of the corresponding extremity of the latter.
Such is the situation of the bones of the forearm and the attitude of the hand in quadrupeds. In short, quadrupeds have their anterior members in the position of pronation.
Fig. 22.—The Human Hand resting for its Whole Extent on its Palmar Surface: Left Side, External Surface.
The individual whom we have just supposed placed in the attitude of a quadruped would be able to maintain this position by pressing on the ground more or less extensive portions of his hands; the whole palm of the hand may be applied to the ground (Fig. 22); or the fingers only—that is to say, the phalanges (Fig. 23); or the extremities of the fingers only—that is to say, the third phalanges (Fig. 24). This last position, which is certainly difficult to maintain, should here be regarded rather as theoretical.
We shall meet with each of these modes of support in certain groups of animals. Thus, the bear, badger, and the majority of rodents, have the paws applied to the ground by the whole extent of the palmar surface of the hand, from the wrist to the tips of the fingers. They are therefore called plantigrade, from the analogy, in this case, of the palm of the hand to the plantar surface, or sole of the foot.
Fig. 24.—The Human Hand resting on the Tips of some of its Third Phalanges: Left Side, External View.
In others, such as the lion, tiger, panther, cat, wolf, and fox, the support is made no longer on the whole extent of the palmar surface, but on the corresponding surface of the fingers only—the metacarpus is turned back, and, consequently, the wrist—that is to say, the carpus—is removed from the ground. These are the digitigrades.
Lastly, the ruminants (sheep, oxen, deer, etc.), and also the pig, ass, and horse, rest on the third phalanx only. In them not only is the metacarpus turned back, but also the two first phalanges. The wrist is very far removed from the ground. In these animals, the third phalanx is enclosed in a case of horn, a nail (the hoof), and because the support of the limb is on that nail, the name of unguligrades has been given them. Nevertheless, as the point of support is on the third phalanx, which is also known by the name of phalangette, we are of opinion that, in order to specify definitely, although they walk on their fingers, as do the digitigrades, the support is provided not by the whole extent of those appendages, they might receive the name of phalangettigrades.
It is necessary among the ruminants to make an exception of the camel and the llama, which are digitigrades.
Just in proportion as the hand is raised from the ground, as we have just seen in passing from the plantigrades to the digitigrades and unguligrades, the number of bones of that region diminishes, the bones of the forearm coalesce, and the ulna tends to disappear; the hand becomes less and less suitable for grasping, climbing, or digging, so as to form an organ exclusively adapted for walking and supporting the body.
Thus, the bear (plantigrade) has five digits, and the power of performing the movements of supination and pronation. Indeed, we know with what facility this animal is able to move his paws in every direction, and climb a tree by grasping it with his fore-limbs. It is well known, however, that no animal except the ape can perform the movements of rotation of the radius around the ulna with the same facility as man; and that none possesses the same degree of suppleness, extent, and variety of movements of the forearm and hand.
In the digitigrades there is one finger which is but slightly developed, and which is always removed from the ground—that is, the thumb: there is also a little less mobility of the radius around the ulna.
In the ungulates the limbs are simply required to perform the movements of walking, and form veritable columns of support, which become the more solid as they are less divided. The bones of the forearm are fused together; there is therefore no possibility of rotation of the radius around the ulna. The metacarpus is reduced to a single piece, which in the horse constitutes what is known as the canon. The number of digits becomes diminished, so that in ruminants there are not more than two, and in the horse but one. We should, however, add that, up to the present, we have taken into account only perfect digits, those that rest on the ground. We shall see further on that there exist supplementary digits, but that they are only slightly developed, and are represented in some cases by mere osseous spurs; it is this fact that has permitted us to ignore them in the general study which we have just made.
Because, as we have already said, the unguligrades have the inferior extremity of the digit encased in a horny sheath, which forms the hoof of the horse and the corresponding structures (onglons) in the ox, those animals have been placed in a special group, which is based on that peculiarity—that is, the group of ungulate mammals.
The plantigrades and digitigrades, of which the paws have their surfaces of support strengthened by an epidermic sole and fatty pads, have the free extremities of the third phalanges covered on their dorsal surface by nails or claws; hence they are named unguiculate mammals.
The bat and birds have the bones of the forearm so arranged that the radius cannot rotate around the ulna. This is necessary in order that during flight, when the wing is being lowered, the radius and hand shall not be able to turn; for, if such rotation took place, each stroke of the wing would place it in a vertical position, which would occasion a loss of resistance incompatible with the effect to be obtained.
The skeleton of the forearm in quadrupeds is vertical in direction; consequently, it forms with the arm an angle open anteriorly; this is well seen on examining the lateral surface. If we examine it on its anterior surface, we find a slight obliquity directed downwards and inwards. In animals in which the bones of the forearm are separate—that is to say, susceptible of supination and pronation—we find a more close resemblance to those of the human skeleton. The ulna, the superior extremity of which always projects beyond that of the radius, has a shaft which gradually narrows from above downwards. Its inferior extremity is terminated by a round head in those animals in which the ulna is fully developed; in others, as it is atrophied, it ends in a thin, long process.
The ulna presents at its superior extremity a posterior process, the olecranon, which forms the point of the elbow. We find on the anterior surface of the same, another process, the coronoid.
Fig. 25.—Superior Extremity of the Bones of the Human Forearm: Left Side, Superior Surface.
1, Radius; 2, ulna; 3, olecranon process; 4, coronoid process.
Fig. 26.—Superior Extremity of the Bones of the Forearm of the Dog: Left Limb, Superior Surface.
1, Radius; 2, ulna; 3, olecranon process; 4, coronoid process.
It is necessary to dwell on the relations of these parts. In man the head of the radius is situated at the anterior part of the external surface of the superior extremity of the ulna (Fig. 25); indeed, the small sigmoid cavity with which the head articulates is situated on the outer side of the coronoid process, and this apophysis is placed in front. In the plantigrades and digitigrades the head of the radius is placed still more forward, so much so that it is situated almost in front of the superior extremity of the ulna (Fig. 26). In the unguligrades it is placed directly in front of this latter (Fig. 27).
Fig. 27.—Superior Extremity of the Bones of the Forearm of the Horse: Left Limb, Superior Surface.
1, Radius; 2, ulna; 3, olecranon process; 4, coronoid process.
Further, the displacement of the radius is made at the expense of the superior extremity of the neighbouring bone; the radius appears to appropriate more and more the parts which in man belong exclusively to the ulna—for example, the coronoid process. In the plantigrades and the digitigrades half of the process still belongs to the ulna and the remainder to the radius. In the ungulates—the horse, for example—the coronoid process belongs to the radius; the ulna, situated behind the latter, is correspondingly diminished in size.
In brief, when we study this region of the skeleton in plantigrades, then in digitigrades, and finally in unguligrades, we find a kind of progressive absorption of one of the two bones (ulna) by the other (radius), which thus becomes the more developed.
It is easy to explain this partial disappearance of the ulna. When the forearm is capable of performing the movements of pronation and supination, the ulna is completely developed, for it is in its small sigmoid cavity that the head of the radius revolves, and it is around its inferior extremity, the head, that the corresponding extremity of the radius turns. But when the movements of rotation of the forearm do not exist, the inferior extremity of the ulna becomes functionally useless and disappears. As to its rôle in the movements of the region of the wrist, that is nil, for we may remember—we will observe it again when we come to treat of the articulations—that the hand articulates with the radius alone (radio-carpal articulation); this is the reason that, when the forearm possesses the fullest mobility, the hand follows the movements which that bone makes around the ulna.
It is not so with the articulation at the elbow-joint; there it is the ulna, which, with the humerus, forms the essential parts (humero-ulnar articulation); its olecranon process limits the movement of extension of the forearm. It is for this reason that, even in those quadrupeds in which the ulna is atrophied, the olecranon process presents a relatively considerable degree of development.
We know that on the posterior surface of the inferior extremity of the bones of the human forearm are grooves in which pass the tendons of the posterior and external muscles which, belonging to this region, are directed for insertion towards the hand.
Fig. 28.—Inferior Extremity of the Bones of the Forearm of a Man: Left Side, Posterior Surface, Position of Supination.
1, Radius; 2, ulna; A, groove for the long abductor and short extensor muscles of the thumb; B, groove for the radial muscles; C, groove for the long extensor of the thumb; D, groove for the special extensor of the index finger and of the common extensor of the fingers; E, groove for the proper extensor of the little finger; F, groove for the posterior ulna.
In animals, because of the movement of rotation of the radius, the surface of this bone, which is anterior, corresponds to the posterior surface of the same in man. (To possess a clear conception of this, it is necessary to remember that, in this latter, the bones of the forearm are always described as in the position of supination; they are thus represented in Fig. 28. The direction of the surfaces of the radius is the reverse of that in animals, since the latter have the radius always in a state of pronation.)
Fig. 29.—Inferior Extremity of the Bones of the Forearm of the Dog: Left Side, Anterior Surface, Normal Position—that is, the Position of Pronation.
1, Radius; 2, ulna; A, groove for the long abductor and for the short extensor of the thumb; B, groove for the radials; D, groove for the long extensor of the thumb, the special extensor of the index-finger, and the common extensor of the fingers; E, groove for the special extensor of the little finger.
Consequently it is on the anterior surface of the bone that we find the grooves concerning which it is necessary to give some details. Regarding them in passing from the radius towards the ulna, those grooves give passage to the tendons of the muscles whose names occupy the columns on p. 43. The letters which are referred to each serve to define their order, and to facilitate reference to Figs. 28, 29, and 30.
Fig. 30.—Inferior Extremity of the Bone of the Forearm of the Horse: Left Side, Anterior Surface.
1, Radius; A, groove for the long abductor and the short extensor of the thumb; B, groove for the radials; D, groove for the common extensor of the digits; E, groove for the special extensor of the little finger.
We should mention that the groove E is situated, both in man and in the dog, at the level of the inferior radio-ulnar articulation; but that in the horse, as the ulna does not exist at that level, the groove is situated on the external surface of the inferior extremity of the radius. It is necessary to add that, in some horses, the ulna is, nevertheless, represented in this region by a tongue-like process of bone; and in such cases the groove is situated in front of this process, at the level of the line of coalescence, which there represents the articulation.
| Man. | Dog. | Horse. | |||
| A. | Long abductor of the thumb, and short extensor of the thumb. | A. | Long abductor of the thumb and short extensor of the thumb united as one muscle, the oblique extensor of the metacarpus.[11] | A. | Oblique extensor of the metacarpus, the homologue of the long abductor of the thumb and the short extensor of the thumb, united as one muscle. |
| B. | First and second external radials (extensor carpi radialis longior and brevior). | B. | The two radials blended superiorly, distinct inferiorly; this is the anterior extensor of the metacarpus. | B. | The radials represented by a single muscle, the anterior extensor of the metacarpus. |
| C. | Long extensor of the thumb. | C. | Long extensor of the thumb and special extensor of the index finger united superiorly. These muscles pass in the following groove. | C. | The long extensor of the thumb and the special extensor of the index are absent. |
| D. | Special extensor of the index finger and the common extensor of the fingers. | D. | Common extensor of the digits and the two preceding muscles. | D. | Anterior extensor of the phalanges, the homologue of the common extensor of the digits. |
| E. | Special extensor of the little finger. | E. | Extensor of the third, fourth, and fifth digits, or the lateral extensor of the digits, the homologue of the special extensor of the little finger. | E. | Lateral extensor of the phalanges, the homologue of the special extensor of the little finger. |
| F. | The posterior ulnar (extensor carpi ulnaris). | There does not exist on the forearm a groove for the posterior ulnar muscle, or external flexor of the metacarpus. | |||
[11] The words printed in italics are the names used in veterinary anatomy.
It is also useful to note, with reference to the groove F, in which passes, in man, the tendon of the posterior ulnar muscle, that, when the forearm is in pronation, the radius alone being displaced, we can only see this groove on the surface which looks backwards; and that it is then separated from the groove which contains the tendon of the special extensor of the little finger by an interval equal to the thickness of the head of the ulna.[12] When the forearm is supinated, the two grooves are found, on the other hand, one beside the other: and the tendons which they contain are very naturally in contact.
[12] Édouard Cuyer, ‘Shape of the Region of the Wrist in Supination and Pronation’ (Bulletin de la Société d’Anthropologie, Paris, 1888).
In birds the forearm is flexed on the arm, and the latter being directed downwards and backwards, the former is, consequently, directed upwards and forwards. Further, because of the position of the humerus, which, as we mentioned on p. 32, has its inferior extremity so turned that the surface which is anterior in man becomes external, the radius, instead of being outside the ulna, is placed above it. This latter is larger than the radius, but its olecranon process is very slightly developed.
The hand in animals, as in man, is formed of three parts—the carpus, metacarpus, and fingers. In man, the forearm and the hand being described in the position of supination; the bones of the carpus are named in passing from the most external to the most internal—that is to say, from that which corresponds to the radial side of the forearm to that which corresponds to the ulnar side. In animals in which, as we know, but it is not unprofitable to repeat, the hand is in pronation, the radial side of the forearm being placed inside, we enumerate the carpal bones in counting the most internal as the first; this is the only method which permits us, in taking our point of departure from the human skeleton as our standard, to recognise the homologies of the bones of the carpal region.
These bones, eight in number, are arranged in two transverse rows, of which one, the first, is superior or anti-brachial; the other, the second, is inferior or metacarpal. Each of these rows contains four bones. Considered in the order we have indicated above—that is to say, proceeding from the radial to the ulnar side—they are thus named: scaphoid, semilunar, cuneiform, and pisiform, in the first row; trapezium, trapezoid, os magnum, and unciform, in the second. The number of these bones is not the same in all animals on account of the coalescence or absence of some. In each row the bones are placed side by side, with the exception of the pisiform, which being placed on the palmar surface of the cuneiform, produces a small projection in man, but a very pronounced one in quadrupeds.
The pisiform is called the hooked bone in some veterinary anatomies. If we consider the hook which it forms, we may recognise that the name is appropriate; but from the point of view of comparison with the human carpus, the name is unfortunate, for it creates confusion between the true pisiform (the fourth bone in the upper row), and the last bone in the lower row, which is the veritable unciform bone. We do not here seek for similarity of form, but homology of regions; and it is only by using the same names to denote the same things that we can succeed in determining such homology.
Taken as a whole, the bones of the carpus form a mass which, by its superior border, articulates with the bones of the forearm, and by its inferior border is in relation with the metacarpal region. Its dorsal surface (anterior in quadrupeds) is slightly convex; its palmar surface (posterior in quadrupeds) is excavated, and forms a groove in which pass the tendons of the flexors of the fingers. This last, in man, has the appearance of a gutter, because of the prominences caused by the projection of the internal and external bones beyond their fellows.
In quadrupeds the palmar groove is especially determined by the pisiform bone, of which we have just mentioned the great development.
The region occupied by the carpus, in the unguligrades, is known as the knee; it would have been more appropriately named had it been called the wrist.
The number of the metacarpal bones in mammals never exceeds five, but it often falls below it; the same is true for the digits. The first are generally equal in number to the latter; an exception is met with in ruminants, whose two metacarpals coalescing soon after birth, form but one bone; this, the canon bone, articulates with two digits.
The number of metacarpals and digits diminishes in proportion as the limbs cease to be organs of prehension, and become more exclusively organs of support and locomotion.
The number of phalanges is two for the thumb and three for each of the other digits; except in the cetaceans, in which they are more numerous.
In the bat, the metacarpals and phalanges are very long, and form the skeleton of the wing; these phalanges are not furnished with nails; the thumb, which is very short, is alone provided with one (Fig. 8).
With regard to the relative dimensions of the bones of the metacarpus, it is necessary to remember that, in the human being, the second metacarpal is the longest; then, in the order of decrease, come the third, fourth, fifth, and first. In quadrupeds we shall also find differences in length (see the chapter relating to the anterior limbs in certain animals), but the order of decrease is not always that which we have just mentioned.
In man the articular surface, situated at the inferior extremity of each of the metacarpals, is rounded, and is called the head. This allows the first phalanx, which is in relation with that surface, to be displaced in every direction; indeed, this phalanx can not only be flexed and extended, but it can also be moved laterally; this latter movement allows of the fingers being separated and drawn together.
In quadrupeds which can only perform the movements of flexion and extension of the digits—for example, the horse—the inferior extremity of the metacarpal has not a rounded head of a regular outline; it is marked by a prominent median crest, directed from before backwards, so that the articular surfaces, which fit more exactly, form a sort of hinge which allows of backward and forward movements only, and permits no lateral displacement. In man, at the level of the inferior extremity of the first metacarpal, in the vicinity of the articulation of this bone with the first phalanx of the thumb, we find two sesamoid bones—small bones developed in the fibrous tissue which surrounds the articulation. We also meet with such structures, but more rarely, at the level of the corresponding articulation of the index and auricular digits; and, more rarely still, at those of the middle and ring fingers. In quadrupeds, these bones are normally developed, and we shall see afterwards that in some animals, as they reach a considerable size, they are able to influence the external outlines; we shall see this, for example, in the horse.