Active organs of Locomotion. Muscles, their Properties, Arrangement, Mode of Action, etc.—If time and space had permitted, I would have considered it my duty to describe, more or less fully, the muscular arrangements of all the animals whose movements I propose to analyse. This is the more desirable, as the movements exhibited by animals of the higher types are directly referable to changes occurring in their muscular system. As, however, I could not hope to overtake this task within the limits prescribed for the present work, I shall content myself by merely stating the properties of muscles; the manner in which muscles act; and the manner in which they are grouped, with a view to moving the osseous levers which constitute the bony framework or skeleton of the animals to be considered. Hitherto, and by common consent, it has been believed that whereas a flexor muscle is situated on one aspect of a limb, and its corresponding extensor on the other aspect, these two muscles must be opposed to and antagonize each other. This belief is founded on what I regard as an erroneous assumption, viz., that muscles have only the power of shortening, and that when one muscle, say the flexor, shortens, it must drag out and forcibly elongate the corresponding extensor, and the converse. This would be a mere waste of power. Nature never works against herself. There are good grounds for believing, as I have stated elsewhere,14 that there is no such thing as antagonism in muscular movements; the several muscles known as flexors and extensors; abductors and adductors; pronators and supinators, being simply correlated. Muscles, when they act, operate upon bones or something extraneous to themselves, and not upon each other. The muscles are folded round the extremities and trunks of animals with a view to operating in masses. For this purpose they are arranged in cycles, there being what are equivalent to extensor and flexor cycles, abductor and adductor cycles, and pronator and supinator cycles. Within these muscular cycles the bones, or extraneous substances to be moved, are placed, and when one side of a cycle shortens, the other side elongates. Muscles are therefore endowed with a centripetal and centrifugal action. These cycles are placed at every degree of obliquity and even at right angles to each other, but they are so disposed in the bodies and limbs of animals that they always operate consentaneously and in harmony. Vide fig. 5, p. 25.

There are in animals very few simple movements, i.e. movements occurring in one plane and produced by the action of two muscles. Locomotion is for the most part produced by the consentaneous action of a great number of muscles; these or their fibres pursuing a variety of directions. This is particularly true of the movements of the extremities in walking, swimming, and flying.

Muscles are divided into the voluntary, the involuntary, and the mixed, according as the will of the animal can wholly, partly, or in no way control their movements. The voluntary muscles are principally concerned in the locomotion of animals. They are the power which moves the several orders of levers into which the skeleton of an animal resolves itself.

The movements of the voluntary and involuntary muscles are essentially wave-like in character, i.e. they spread from certain centres, according to a fixed order, and in given directions. In the extremities of animals the centripetal or converging muscular wave on one side of the bone to be moved, is accompanied by a corresponding centrifugal or diverging wave on the other side; the bone or bones by this arrangement being perfectly under control and moved to a hair’s-breadth. The centripetal or converging, and the centrifugal or diverging waves of force are, as already indicated, correlated.15 Similar remarks may be made regarding the different parts of the body of the serpent when creeping, of the body of the fish when swimming, of the wing of the bird when flying, and of our own extremities when walking. In all those cases the moving parts are thrown into curves or waves definitely correlated.

It may be broadly stated, that in every case locomotion is the result of the opening and closing of opposite sides of muscular cycles. By the closing or shortening, say of the flexor halves of the cycles, and the opening or elongation of the extensor halves, the angles formed by the osseous levers are diminished; by the closing or shortening of the extensor halves of the cycles, and the opening or elongation of the flexor halves, the angles formed by the osseous levers are increased. This alternate diminution and increase of the angles formed by the osseous levers produce the movements of walking, swimming, and flying. The muscular cycles of the trunk and extremities are so disposed with regard to the bones or osseous levers, that they in every case produce a maximum result with a minimum of power. The origins and insertions of the muscles, the direction of the muscles and the distribution of the muscular fibres insure, that if power is lost in moving a lever, speed is gained, there being an apparent but never a real loss. The variety and extent of movement is secured by the obliquity of the muscular fibres to their tendons; by the obliquity of the tendons to the bones they are to move; and by the proximity of the attachment of the muscles to the several joints. As muscles are capable of shortening and elongating nearly a fourth of their length, they readily produce the precise kind and degree of motion required in any particular case.16

The force of muscles, according to the experiments of Schwann, increases with their length, and vice versa. It is a curious circumstance, and worthy the attention of those interested in homologies, that the voluntary muscles of the superior and inferior extremities, and more especially of the trunk, are arranged in longitudinal, transverse, and oblique spiral lines, and in layers or strata precisely as in the ventricles of the heart and hollow muscles generally.17 If, consequently, I eliminate the element of bone from these several regions, I reproduce a typical hollow muscle; and what is still more remarkable, if I compare the bones removed (say the bones of the anterior extremity of a quadruped or bird) with the cast obtained from the cavity of a hollow muscle (say the left ventricle of the heart of the mammal), I find that the bones and the cast are twisted upon themselves, and form elegant screws, the threads or ridges of which run in the same direction. This affords a proof that the involuntary hollow muscles supply the type or pattern on which the voluntary muscles are formed. Fig. 6 represents the bones of the wing of the bird; fig. 7 the bones of the 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 system—should 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 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.18 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 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 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 ball, 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.19

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 extremities of quadrupeds, bipeds, etc. in walking; in the movements of the tails and fins of fishes, whales, etc. in swimming; and in the movements of the wings of insects, bats, and birds in flying. The straight and oblique muscles are usually found together, and co-operate in producing the movements in question; the amount of rotation in a part always increasing as the oblique muscles preponderate. The combination of ball-and-socket and hinge-joints, with their concomitant oblique and longitudinal muscular cycles (the former occurring in their most perfect forms where the extremities are united to the trunk, the latter in the extremities themselves), enable the animal to present, when necessary, an extensive resisting surface the one instant, and a greatly diminished and a comparatively non-resisting one the next. This arrangement secures the subtlety and nicety of motion demanded by the several media at different stages of progression.

The travelling surfaces of Animals modified and adapted to the medium on or in which they move.—In those land animals which take to the water occasionally, the feet, as a rule, are furnished with membranous expansions extending between the toes. Of such the Otter (fig. 12), Ornithorhynchus (fig. 11), Seal (fig. 14), Crocodile, Sea-Bear (fig. 37, p. 76), Walrus, Frog (fig. 13), and Triton, may be cited. The crocodile and triton, in addition to the membranous expansion occurring between the toes, are supplied with a powerful swimming-tail, which adds very materially to the surface engaged in natation. Those animals, one and all, walk awkwardly, it always happening that when the extremities are modified to operate upon two essentially different media (as, for instance, the land and water), the maximum of speed is attained in neither. For this reason those animals which swim the best, walk, as a rule, with the greatest difficulty, and vice versâ, as the movements of the auk and seal in and out of the water amply testify.

In addition to those land animals which run and swim, there are some which precipitate themselves, parachute-fashion, from immense heights, and others which even fly. In these the membranous expansions are greatly increased, the ribs affording the necessary support in the Dragon or Flying Lizard (fig. 15), the anterior and posterior extremities and tail, in the Flying Lemur (fig. 16) and Bat (fig. 17, p. 36).

Although no lizard is at present known to fly, there can be little doubt that the extinct Pterodactyles (which, according to Professor Huxley, are intermediate between the lizards and crocodiles) were possessed of this power. The bat is interesting as being the only mammal at present endowed with wings sufficiently large to enable it to fly.20 It affords an extreme example of modification for a special purpose,—its attenuated body, dwarfed posterior, and greatly elongated anterior extremities, with their enormous fingers and outspreading membranes, completely unfitting it for terrestrial progression. It is instructive as showing that flight may be attained, without the aid of hollow bones and air-sacs, by purely muscular efforts, and by the mere diminution and increase of a continuous membrane.

As the flying lizard, flying lemur, and bat (figs. 15, 16, and 17, pp. 35 and 36), connect terrestrial progression with aërial progression, so the auk, penguin (fig. 46, p. 91), and flying-fish (fig. 51, p. 98), connect progression in the water with progression in the air. The travelling surfaces of these anomalous creatures run the movements peculiar to the three highways of nature into each other, and bridge over, as it were, the gaps which naturally exist between locomotion on the land, in the water, and in the air.