Animal Locomotion; or, walking, swimming, and flying With a dissertation on aëronautics
Part 6
In examining figures 21, 22, and 23, the reader will do well to remember that the _near_ fore and hind feet of a horse are the _left_ fore and hind feet; the _off_ fore and hind feet being the _right_ fore and hind feet. The terms _near_ and _off_ are technical expressions, and apply to the left and right sides of the animal. Another point to be attended to in examining the figures in question, is the relation which exists between the fore and hind feet of the near and off sides of the body. In slow walking the near hind foot is planted behind the imprint made by the near fore foot. In rapid walking, on the contrary, the near hind foot is planted from six to twelve or more inches in advance of the imprint made by the near fore foot (fig. 21 represents the distance as eleven inches). In the trot the near hind foot is planted from twelve to eighteen or more inches in advance of the imprint made by the near fore foot (fig. 22 represents the distance as nineteen inches). In the gallop the near hind foot is planted 100 or more inches in advance of the imprint made by the near fore foot (fig. 23 represents the distance as 110-1/2 inches). The distance by which the near hind foot passes the near fore foot in rapid walking, trotting, and galloping, increases in a progressive ratio, and is due in a principal measure to the velocity or momentum acquired by the mass of the horse in rapid motion; the body of the animal carrying forward and planting the limbs at greater relative distances in the trot than in the rapid walk, and in the gallop than in the trot. I have chosen to speak of the near hind and near fore feet, but similar remarks may of course be made of the off hind and off fore feet.
“At fig. 23, which represents the gallop, the distance between two successive impressions produced, say by the near fore foot, is eighteen feet one inch and a half. Midway between these two impressions is the mark of the near hind foot, which therefore subdivides the space into nine feet and six-eighths of an inch, but each of these is again subdivided into two halves by the impressions produced by the off fore and off hind feet. It is thus seen that the horse’s body instead of being propelled through the air by bounds or leaps even when going at the highest attainable speed, acts on a system of levers, the mean distance between the points of resistance of which is four feet six inches. The exact length of stride, of course, only applies to that of the particular horse observed, and the rate of speed at which he is going. In the case of any one animal, the greater the speed the longer is the individual stride. In progression, the body moves before a limb is raised from the ground, as is most readily seen when the horse is beginning its slowest action, as in traction.”[27]
[27] Gamgee in Journal of Anatomy and Physiology, vol. iii. pp. 375, 376.
At fig. 22, which represents the trot, the stride is ten feet one inch. At fig. 21, which represents the walk, it is only five feet five inches. The speed acquired, Mr. Gamgee points out, determines the length of stride; the length of stride being the effect and evidence of speed and not the cause of it. The momentum acquired in the gallop, as already explained, greatly accelerates speed.
“In contemplating length of strides, with reference to the fulcra, allowance has to be made for the length of the feet, which is to be deducted from that of the strides, because the apex, or toe of the horse’s hind foot forms the fulcrum in one instant, and the heel of the fore foot in the next, and _vice versâ_. This phenomenon is very obvious in the action of the human foot, and is remarkable also for the range of leverage thus afforded in some of the fleetest quadrupeds, of different species. In the hare, for instance, between the point of its hock and the termination of its extended digits, there is a space of upwards of six inches of extent of leverage and variation of fulcrum, and in the fore limb from the _carpus_ to the toe-nails (whose function in progression is not to be underrated) upwards of three inches of leverage are found, being about ten inches for each lateral biped, and the double of that for the action of all four feet. Viewed in this way the stride is not really so long as would be supposed if merely estimated from the space between the footprints.
“Many interesting remarks might be made on the length of the stride of various animals; the full movement of the greyhound is, for instance, upwards of sixteen feet; that of the hare at least equal; whilst that of the Newfoundland dog is a little over nine feet.”[27]
_Locomotion of the Ostrich._--Birds have been divided by naturalists into eight orders:--the _Natatores_, or Swimming Birds; the _Grallatores_, or Wading Birds; the _Cursores_, or Running Birds; the _Scansores_, or Climbers; the _Rasores_, or Scrapers; the _Columbæ_, or Doves; the _Passeres_; and the _Raptores_, or Birds of Prey.
The first five orders have been classified according to their habits and modes of progression. The _Natatores_ I shall consider when I come to speak of swimming as a form of locomotion, and as there is nothing in the movements of the wading, scraping, and climbing birds,[28] or in the _Passeres_[29] or _Raptores_, requiring special notice, I shall proceed at once to a consideration of the _Cursores_, the best examples of which are the ostrich, emu, cassowary, and apteryx.
[28] The woodpeckers climb by the aid of the stiff feathers of their tails; the legs and tail forming a firm basis of support.
[29] In this order there are certain birds--the sparrows and thrushes, for example--which advance by a series of vigorous leaps; the leaps being of an intermitting character.
The ostrich is remarkable for the great length and development of its legs as compared with its wings (fig. 24). In this respect it is among birds what the kangaroo is among mammals. The ostrich attains an altitude of from six to eight feet, and is the largest living bird known. Its great height is due to its attenuated neck and legs. The latter are very powerful structures, and greatly resemble in their general conformation the posterior extremities of a thoroughbred horse or one of the larger deer--compare with fig. 4, p. 21. They are expressly made for speed. Thus the bones of the leg and foot are inclined very obliquely towards each other, the femur being inclined very obliquely to the ilium. As a consequence the angles made by the several bones of the legs are comparatively small; smaller in fact than in either the horse or deer.
The feet of the ostrich, like those of the horse and deer, are reduced to a minimum as regards size; so that they occasion very little friction in the act of walking and running. The foot is composed of two jointed toes,[30] which spread out when the weight of the body comes upon them, in such a manner as enables the bird to seize and let go the ground with equal facility. The advantage of such an arrangement in rapid locomotion cannot be over-estimated. The elasticity and flexibility of the foot contribute greatly to the rapidity of movement for which this celebrated bird is famous. The limb of the ostrich, with its large bones placed very obliquely to form a system of powerful levers, is the very embodiment of speed. The foot is quite worthy of the limb, it being in some respects the most admirable structure of its kind in existence. The foot of the ostrich differs considerably from that of all other birds, those of its own family excepted. Thus the under portion of the foot is flat, and specially adapted for acting on plane surfaces, particularly solids.[31] The extremities of the toes superiorly are armed with powerful short nails, the tips of which project inferiorly to protect the toes and confer elasticity when the foot is leaving the ground. The foot, like the leg, is remarkable for its great strength. The legs of the ostrich are closely set, another feature of speed.[32] The wings of the ostrich are in a very rudimentary condition as compared with the legs.[33] All the bones are present, but they are so dwarfed that they are useless as organs of flight. The angles which the bones of the wing make with each other, are still less than the angles made by the bones of the leg. This is just what we would _a priori_ expect, as the velocity with which wings are moved greatly exceeds that with which legs are moved. The bones of the wing of the ostrich are inclined towards each other at nearly right angles. The wings of the ostrich, although useless as flying organs, form important auxiliaries in running. When the ostrich careers along the plain, he spreads out his wings in such a manner that they act as balancers, and so enable him to maintain his equilibrium (fig. 25). The wings, because of the angle of inclination which their under surfaces make with the horizon, and the great speed at which the ostrich travels, act like kites, and so elevate and carry forward by a mechanical adaptation a certain proportion of the mass of the bird already in motion. The elevating and propelling power of even diminutive inclined planes is very considerable, when carried along at a high speed in a horizontal direction. The wings, in addition to their elevating and propelling power, contribute by their short, rapid, swinging movements, to continuity of motion in the legs. No bird with large wings can run well. The albatross, for example, walks with difficulty, and the same may be said of the vulture and eagle. What, therefore, appears a defect in the ostrich, is a positive advantage when its habits and mode of locomotion are taken into account.
[30] The toes in the emu amount to three.
[31] Feet designed for swimming, grasping trees, or securing prey, do not operate to advantage on a flat surface. The awkward waddle of the swan, parrot, and eagle when on the ground affords illustrations.
[32] In draught horses the legs are much wider apart than in racers; the legs of the deer being less widely set than those of the racer.
[33] In the apteryx the wings are so very small that the bird is commonly spoken of as the “wingless bird.”
Professional runners in many cases at matches reduce the length of their anterior extremities by flexing their arms and carrying them on a level with their chest (fig. 28, p. 62). It would seem that in rapid running there is not time for the arms to oscillate naturally, and that under these circumstances the arms, if allowed to swing about, retard rather than increase the speed. The centre of gravity is well forward in the ostrich, and is regulated by the movements of the head and neck, and the obliquity of the body and legs. In running the neck is stretched, the body inclined forward, and the legs moved alternately and with great rapidity. When the right leg is flexed and elevated, it swings forward pendulum-fashion, and describes a curve whose convexity is directed towards the right side. When the left leg is flexed and elevated, it swings forward and describes a curve whose convexity is directed towards the left side. The curves made by the right and left legs form when united a waved line (_vide_ figs. 18, 19, and 20, pp. 37, 39, and 41). When the right leg is flexed, elevated, and advanced, it rotates upon the iliac portion of the trunk of the bird, the trunk being supported for the time being by the left leg, which is extended, and in contact with the ground. When the left leg is flexed, elevated, and advanced, it in like manner rotates upon the trunk, supported in this instance by the extended right leg. The leg which is on the ground for the time being supplies the necessary lever, the ground the fulcrum. When the right leg is flexed and elevated, it rotates upon the iliac portion of the trunk in a forward direction, the right foot describing the arc of a circle. When the right leg and foot are extended and fixed on the ground, the trunk rotates upon the right foot in a forward direction to form the arc of a circle, which is the converse of that formed by the right foot. If the arcs alternately supplied by the right foot and trunk are placed in opposition, a more or less perfect circle is produced, and thus it is that the locomotion of animals is approximated to the wheel in mechanics. Similar remarks are to be made of the left foot and trunk. The alternate rolling of the trunk on the extremities, and the extremities on the trunk, utilizes or works up the inertia of the moving mass, and powerfully contributes to continuity and steadiness of action in the moving parts. By advancing the head, neck, and anterior parts of the body, the ostrich inaugurates the rolling movement of the trunk, which is perpetuated by the rolling movements of the legs. The trunk and legs of the ostrich are active and passive by turns. The movements of the trunk and limbs are definitely co-ordinated. But for this reciprocation the action of the several parts implicated would neither be so rapid, certain, nor continuous. The speed of the ostrich exceeds that of every other land animal, a circumstance due to its long, powerful legs and great stride. It can outstrip without difficulty the fleetest horses, and is only captured by being simultaneously assailed from various points, or run down by a succession of hunters on fresh steeds. If the speed of the ostrich, which only measures six or eight feet, is so transcending, what shall we say of the speed of the extinct _Æpyornis maximus_ and _Dinornis giganteus_, which are supposed to have measured from sixteen to eighteen feet in height? Incredible as it may appear, the ostrich, with its feet reduced to a minimum as regards size, and peculiarly organized for walking and running on solids, can also swim. Mr. Darwin, that most careful of all observers, informs us that ostriches take to the water readily, and not only ford rapid rivers, but also cross from island to island. They swim leisurely, with neck extended, and the greater part of the body submerged.
_Locomotion in Man._--The speed attained by man, although considerable, is not remarkable. It depends on a variety of circumstances, such as the height, age, sex, and muscular energy of the individual, the nature of the surface passed over, and the resistance to forward motion due to the presence of air, whether still or moving. A reference to the human skeleton, particularly its inferior extremities, will explain why the speed should be moderate.
On comparing the inferior extremities of man with the legs of birds, or the posterior extremities of quadrupeds, say the horse or deer, we find that the bones composing them are not so obliquely placed with reference to each other, neither are the angles formed by any two bones so acute. Further, we observe that in birds and quadrupeds the tarsal and metatarsal bones are so modified that there is an actual increase in the number of the angles themselves. In the extremities of birds and quadrupeds there are four angles, which may be increased or diminished in the operations of locomotion. Thus, in the quadruped and bird (fig. 4, p. 21, and fig. 24, p. 47), the femur forms with the ilium one angle (_a_); the tibia and fibula with the femur a second angle (_b_); the cannon or tarso-metatarsal bone with the tibia and fibula a third angle (_c_); and the bones of the foot with the cannon or tarso-metatarsal bone a fourth angle (_d_). In man three angles only are found, marked respectively _a_, _b_, and _c_ (figs. 26 and 27, pp. 55 and 59). The fourth angle (_d_ of figs. 4 and 24) is absent. The absence of the fourth angle is due to the fact that in man the tarsal and metatarsal bones are shortened and crushed together; whereas in the quadruped and bird they are elongated and separated.
As the speed of a limb increases in proportion to the number and acuteness of the angles formed by its several bones, it is not difficult to understand why man should not be so swift as the majority of quadrupeds. The increase in the number of angles increases the power which an animal has of shortening and elongating its extremities, and the levers which the extremities form. To increase the length of a lever is to increase its power at one end, and the distance through which it moves at the other; hence the faculty of bounding or leaping possessed in such perfection by many quadrupeds.[34] If the wing be considered as a lever, a small degree of motion at its root produces an extensive sweep at its tip. It is thus that the wing is enabled to work up and utilize the thin medium of the air as a buoying medium.
[34] “The posterior extremities in both the lion and tiger are longer, and the bones inclined more obliquely to each other than the anterior, giving them greater power and elasticity in springing.”
Another drawback to great speed in man is his erect position. Part of the power which should move the limbs is dedicated to supporting the trunk. For the same reason the bones of the legs, instead of being obliquely inclined to each other, as in the quadruped and bird, are arranged in a nearly vertical spiral line. This arrangement increases the angle formed by any two bones, and, as a consequence, decreases the speed of the limbs, as explained. A similar disposition of the bones is found in the anterior extremities of the elephant, where the superincumbent weight is great, and the speed, comparatively speaking, not remarkable. The bones of the human leg are beautifully adapted to sustain the weight of the body and neutralize shock.[35] Thus the femur or thigh bone is furnished at its upper extremity with a ball-and-socket joint which unites it to the cup-shaped depression (acetabulum) in the ilium (hip bone). It is supplied with a neck which carries the body or shaft of the bone in an oblique direction from the ilium, the shaft being arched forward and twisted upon itself to form an elongated cylindrical screw. The lower extremity of the femur is furnished with spiral articular surfaces accurately adapted to the upper extremities of the bones of the leg, viz. the tibia and fibula, and to the patella. The bones of the leg (tibia and fibula) are spirally arranged, the screw in this instance being split up. At the ankle the bones of the leg are applied to those of the foot by spiral articular surfaces analogous to those found at the knee-joint. The weight of the trunk is thus thrown on the foot, not in straight lines, but in a series of curves. The foot itself is wonderfully adapted to receive the pressure from above. It consists of a series of small bones (the tarsal, metatarsal, and phalangeal bones), arranged in the form of a double arch; the one arch extending from the heel towards the toes, the other arch across the foot. The foot is so contrived that it is at once firm, elastic, and moveable,--qualities which enable it to sustain pressure from above, and exert pressure from beneath. In walking, the heel first reaches and first leaves the ground. When the heel is elevated the weight of the body falls more and more on the centre of the foot and toes, the latter spreading out[36] as in birds, to seize the ground and lever the trunk forward. It is in this movement that the wonderful mechanism of the foot is displayed to most advantage, the multiplicity of joints in the foot all yielding a little to confer that elasticity of step which is so agreeable to behold, and which is one of the characteristics of youth. The foot may be said to roll over the ground in a direction from behind forwards. I have stated that the angles formed by the bones of the human leg are larger than those formed by the bones of the leg of the quadruped and bird. This is especially true of the angle formed by the femur with the ilium, which, because of the upward direction given to the crest of the ilium in man, is so great that it virtually ceases to be an angle.
[35] “The pelvis receives the whole weight of the trunk and superposed organs, and transmits it to the heads of the femurs.”
[36] The spreading action of the toes is seen to perfection in children. It is more or less destroyed in adults from a faulty principle in boot and shoemaking, the soles being invariably too narrow.
The bones of the superior extremities in man merit attention from the fact that in walking and running they oscillate in opposite directions, and alternate and keep time with the legs, which oscillate in a similar manner. The arms are articulated at the shoulders by ball-and-socket joints to cup-shaped depressions (glenoid cavities) closely resembling those found at the hip-joints. The bone of the arm (humerus) is carried away from the shoulder by a short neck, as in the thigh-bone (femur). Like the thigh-bone it is twisted upon itself and forms a screw. The inferior extremity of the arm bone is furnished with spiral articular surfaces resembling those found at the knee. The spiral articular surfaces of the arm bone are adapted to similar surfaces existing on the superior extremities of the bones of the forearm, to wit, the radius and ulna. These bones, like the bones of the leg, are spirally disposed with reference to each other, and form a screw consisting of two parts. The bones of the forearm are united to those of the wrist (carpal) and hand (metacarpal and phalangeal) by articular surfaces displaying a greater or less degree of spirality. From this it follows that the superior extremities of man greatly resemble his inferior ones; a fact of considerable importance, as it accounts for the part taken by the superior extremities in locomotion. In man the arms do not touch the ground as in the brutes, but they do not on this account cease to be useful as instruments of progression. If a man walks with a stick in each hand the movements of his extremities exactly resemble those of a quadruped.
The bones of the human extremities (superior and inferior) are seen to advantage in fig. 26; and I particularly direct the attention of the reader to the ball-and-socket or universal joints by which the arms are articulated to the shoulders (_x_, _x´_), and the legs to the pelvis (_a_, _a´_), as a knowledge of these is necessary to a comprehension of the oscillating or pendulum movements of the limbs now to be described. The screw configuration of the limbs is well depicted in the left arm (_x_) of the present figure. Compare with the wing of the bird, fig. 6, and with the anterior extremity of the elephant, fig. 7, p. 28. But for the ball-and-socket joints, and the spiral nature of the bones and articular surfaces of the extremities, the undulating, sinuous, and more or less continuous movements observable in walking and running, and the twisting, lashing, flail-like movements necessary to swimming and flying, would be impossible.