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anterior extremity of the elephant; and fig. 8 the cast or mould of the cavity of the left ventricle of the heart of the deer.
It has been the almost invariable custom in teaching anatomy, and such parts of physiology as pertain to animal movements, to place much emphasis upon the configuration of the bony skeleton as a whole, and the conformation of its several articular surfaces in particular. This is very natural, as the osseous system stands the wear and tear of time, while all around it is in a great measure perishable. It is the link which binds extinct forms to living ones, and we naturally venerate and love what is enduring. It is no marvel that Oken, Goethe, Owen, and others should have attempted such splendid generalizations with regard to the osseous systemshould have proved with such cogency of argument that the head is an expanded vertebra. The bony skeleton is a miracle of design very wonderful and very beautiful in its way. But when all has been said, the fact remains that the skeleton, when it exists, forms only an adjunct of locomotion and motion generally. All the really essential movements of an animal occur in its soft parts. The osseous system is therefore to be regarded as secondary in importance to the muscular, of which it may be considered a differentiation. Instead of regarding the muscles as adapted to the bones, the bones ought to be regarded as adapted to the muscles. Bones have no power either of originating or perpetuating motion. This begins and terminates in the muscles. Nor must it be overlooked, that bone makes its appearance comparatively late in the scale of being; that innumerable creatures exist in which no trace either of an external or internal skeleton is to be found ; that these creatures move freely about, digest, circulate their nutritious juices and blood when present, multiply, and perform all the functions incident to life. While the skeleton is to be found in only a certain proportion of the animals existing on our globe, the soft parts are to be met
" On the Muscular Arrangements of the Bladder and Prostate, and the manner in which the Ureters and Urethra are closed,” by the Author.Philosophical Transactions, 1867.
« On the Muscular Tunics in the Stomach of Man and other Mammalia,” by the Author.–Proceedings Royal Society of London, 1867.
with in all; and this appears to me an all-sufficient reason for attaching great importance to the movements of soft parts, such as protoplasm, jelly masses, involuntary and voluntary muscles, etc. As the muscles of vertebrates are accurately applied to each other, and to the bones, while the bones are rigid, unyielding, and incapable of motion, it follows that the osseous system acts as a break or boundary to the muscular one,—and hence the arbitrary division of muscles into extensors and flexors, pronators and supinators, abductors and adductors. This division although convenient is calculated to mislead. The most highly organized animal is strictly speaking to be regarded as a living mass whose parts (hard, soft, and
Fig. 9. —The Superficial Muscles in the Horse, (after Bagg). otherwise) are accurately adapted to each other, every part reciprocating with scrupulous exactitude, and rendering it difficult to determine where motion begins and where it terminates. Fig. 9 shows the more superficial of the muscular masses which move the bones or osseous levers of the horse, as seen in the walk, trot, gallop, etc. A careful examination of these carneous masses or muscles will show that they run
1 Lectures “ On the Physiology of the Circulation in Plants, in the Lower Animals, and in Man," by the Author.- Edinburgh Medical Journal for September 1872.
longitudinally, transversely, and obliquely, the longitudinal and transverse muscles crossing each other at nearly right angles, the oblique ones tending to cross at various angles, as in the letter X. The crossing is seen to most advantage in the deep muscles.
In order to understand the twisting which occurs to a greater or less extent in the bodies and extremities (when present) of all vertebrated animals, it is necessary to reduce the bony and muscular systems to their simplest expression. If motion is desired in a dorsal, ventral, or lateral direction only, a dorsal and ventral or a right and left lateral set of longitudinal muscles acting upon straight bones articulated by an ordinary ball-and-socket joint will suffice. In this case the dorsal, ventral, and right and left lateral muscles form muscular cycles ; contraction or shortening on the one aspect of the cycle being accompanied by relaxation or elongation on the other, the bones and joints forming as it were the diameters of the cycles, and oscillating in a backward, forward, or lateral direction in proportion to the degree and direction of the muscular movements. Here the motion is confined to two planes intersecting each other at right angles. When, however, the muscular system becomes more highly differentiated, both as regards the number of the muscles employed, and the variety of the directions pursued by them, the bones and joints also become more complicated. Under these circumstances, the bones, as a rule, are twisted upon themselves, and their articular surfaces present various degrees of spirality to meet the requirements of the muscular system. Between the straight longitudinal muscles, therefore, arranged in dorsal and ventral, and right and left lateral sets, and those which run in a more or less transverse direction, and between the simple joint whose motion is confined to one plane and the ball-and-socket joints whose movements are universal, every degree of obliquity is found in the direction of the muscles, and every possible modification in the disposition of the articular surfaces. In the fish the muscles are for the most part arranged in dorsal, ventral, and lateral sets, which run longitudinally; and, as a result, the movements of the trunk, particularly towards the tail, are from side to side and sinuous. As, however, oblique fibres are also present, and the tendons of the longitudinal muscles in some instances cross obliquely towards the tail, the fish has also the power of tilting or twisting its trunk (particularly the lower half) as well as the caudal fin. In a mackerel which I examined, the oblique muscles were represented by the four lateral masses occurring between the dorsal, ventral, and lateral longitudinal muscles-two of these being found on either side of the fish, and corresponding to the myocommas or “grand muscle latéral” of Cuvier. The muscular system of the fish would therefore seem to be arranged on a fourfold plan,—there being four sets of longitudinal muscles, and a corresponding number of slightly oblique and oblique muscles, the oblique muscles being spiral in their nature and tending to cross or intersect at various angles, an arrest of the intersection, as it appears to me, giving rise to the myocommas and to that concentric arrangement of their constituent parts so evident on transverse section. This tendency of the muscular fibres to cross each other at various degrees of obliquity may also be traced in several parts of the human body, as, for instance, in the deltoid muscle of the arm and the deep muscles of the leg. Numerous other examples of penniform muscles might be adduced. Although the fibres of the myocommas have a more or less longitudinal direction, the myocommas themselves pursue an oblique spiral course from before backwards and from within outwards, i.e. from the spine towards the periphery, where they receive slightly oblique fibres from the longitudinal dorsal, ventral, and lateral muscles. As the spiral oblique myocommas and the oblique fibres from the longitudinal muscles act directly and indirectly upon the spines of the vertebræ, and the vertebræ themselves to which they are specially adapted, and as both sets of oblique fibres are geared by interdigitation to the fourfold set of longitudinal muscles, the lateral, sinuous, and rotatory movements of the body and tail of the fish are readily accounted for. The spinal column of the fish facilitates the lateral sinuous twisting movements of the tail and trunk, from the fact that the vertebræ composing it are united to each other by a series of modified universal joints—the vertebræ supplying the cup
shaped depressions or sockets, the intervertebral substance, the prominence or ball.
The same may be said of the general arrangement of the muscles in the trunk and tail of the Cetacea, the principal muscles in this case being distributed, not on the sides, but on the dorsal and ventral aspects. The lashing of the tail in the whales is consequently from above downwards or vertically, instead of from side to side. The spinal column is jointed as in the fish, with this difference, that the vertebræ (especially towards the tail) form the rounded prominences or hall, the meniscus or cup-shaped intervertebral plates the receptacles or socket.
When limbs are present, the spine may be regarded as being ideally divided, the spiral movements, under these circumstances, being thrown upon the extremities by typical ball-and-socket joints occurring at the shoulders and pelvis. This is peculiarly the case in the seal, where the spirally sinuous movements of the spine are transferred directly to the posterior extremities.
The extremities, when present, are provided with their own muscular cycles of extensor and flexor, abductor and adductor, pronator and supinator muscles,—these running longitudinally and at various degrees of obliquity, and enveloping the hard parts according to their direction—the bones being twisted upon themselves and furnished with articular surfaces which reflect the movements of the muscular cycles, whether these occur in straight lines anteriorly, posteriorly, or laterally, or in oblique lines in intermediate situations. The straight and oblique muscles are principally brought into play in the movements of the extremi
1 That the movements of the extremities primarily emanate from the spine is rendered probable by the remarkable powers possessed by serpents. “It is true,” writes Professor Owen (op. cit. p. 261), “ that the serpent has no limbs, yet it can outclimb the monkey, outswim the fish, outleap the jerboa, and, suddenly loosing the close coils of its crouching spiral, it can spring into the air and seize the bird upon the wing." .... “The serpent has neither hands nor talons, yet it can outwrestle the athlete, and crush the tiger in the embrace of its ponderous overlapping folds.” The peculiar endowments, wbich accompany the possession of extremities, it appears to me, present, themselves in an undeveloped or latent form in the trunk of the reptile.