The Wings of Birds.
The Bones of the Wing of the Bird—their Articular Surfaces, Movements, etc.—The humerus, or arm-bone of the wing, is supported by three of the trunk-bones, viz. the scapula or shoulder-blade, the clavicle or collar-bone, also called the furculum,[84] and the coracoid bone,—these three converging to form a point d’appui, or centre of support for the head of the humerus, which is received in facettes or depressions situated on the scapula and coracoid. In order that the wing may have an almost unlimited range of motion, and be wielded after the manner of a flail, it is articulated to the trunk by a somewhat lax universal joint, which permits vertical, horizontal, and intermediate movements.[85] The long axis of the joint is directed vertically; the joint itself somewhat backwards. It is otherwise with the elbow-joint, which is turned forwards, and has its long axis directed horizontally, from the fact that the humerus is twisted upon itself to the extent of nearly a quarter of a turn. The elbow-joint is decidedly spiral in its nature, its long axis intersecting that of the shoulder-joint at nearly right angles. The humerus articulates at the elbow with two bones, the radius and the ulna, the former of which is pushed from the humerus, while the other is drawn towards it during extension, the reverse occurring during flexion. Both bones, moreover, while those movements are taking place, revolve to a greater or less extent upon their own axes. The bones of the forearm articulate at the wrist with the carpal bones, which being spirally arranged, and placed obliquely between them and the metacarpal bones, transmit the motions to the latter in a curved direction. The long axis of the wrist-joint is, as nearly as may be, at right angles to that of the elbow-joint, and more or less parallel with that of the shoulder. The metacarpal or hand-bones, and the phalanges or finger-bones are more or less fused together, the better to support the great primary feathers, on the efficiency of which flight mainly depends. They are articulated to each other by double hinge-joints, the long axes of which are nearly at right angles to each other.
As a result of this disposition of the articular surfaces, the wing is shot out or extended and retracted or flexed in a variable plane, the bones composing the wing, particularly those of the forearm, rotating on their axes during either movement.
This secondary action, or the revolving of the component bones upon their own axes, is of the greatest importance in the movements of the wing, as it communicates to the hand and forearm, and consequently to the primary and secondary feathers which they bear, the precise angles necessary for flight; it in fact insures that the wing, and the curtain or fringe of the wing which the primary and secondary feathers form, shall be screwed into and down upon the air in extension, and unscrewed or withdrawn from it during flexion. The wing of the bird may therefore be compared to a huge gimlet or auger; the axis of the gimlet representing the bones of the wing, the flanges or spiral thread of the gimlet the primary and secondary feathers (fig. [63], p. 138, and fig. 97, p. 176).
Traces of Design in the Wing of the Bird—the arrangement of the Primary, Secondary, and Tertiary Feathers, etc.—There are few things in nature more admirably constructed than the wing of the bird, and perhaps none where design can be more readily traced. Its great strength and extreme lightness, the manner in which it closes up or folds during flexion, and opens out or expands during extension, as well as the manner in which the feathers are strung together and overlap each other in divers directions to produce at one time a solid resisting surface, and at another an interrupted and comparatively non-resisting one, present a degree of fitness to which the mind must necessarily revert with pleasure. If the feathers of the wing only are contemplated, they may be conveniently divided into three sets of three each (on both sides of the wing)—an upper or dorsal set (fig. [61], d, e, f, p. 136), a lower or ventral set (c, a, b), and one which is intermediate. This division is intended to refer the feathers to the bones of the arm, forearm, and hand, but is more or less arbitrary in its nature. The lower set or tier consists of the primary (b), secondary (a), and tertiary (c) feathers, strung together by fibrous structures in such a way that they move in an outward or inward direction, or turn upon their axes, at precisely the same instant of time,—the middle and upper sets of feathers, which overlap the primary, secondary, and tertiary ones, constituting what are called the “coverts” and “sub-coverts.” The primary or rowing feathers are the longest and strongest (b), the secondaries (a) next, and the tertiaries third (c). The tertiaries, however, are occasionally longer than the secondaries. The tertiary, secondary, and primary feathers increase in strength from within outwards, i.e. from the body towards the extremity of the wing, and so of the several sets of wing-coverts. This arrangement is necessary, because the strain on the feathers during flight increases in proportion to their distance from the trunk.
Fig. 98.
Fig. 99. Fig. 100. Fig. 101.
Figs. 98, 99, 100, and 101 show the muscles and elastic ligaments, and the arrangement of the primary and secondary feathers on the ventral aspects of the wing of the crested crane. The wing is in the extended condition.
y (fig. 98), Great pectoral muscle which depresses the wing.
a b, Voluntary muscular fibres terminating in elastic band k. This band splits up into two portions (k, m). A somewhat similar band is seen at j. These three bands are united to, and act in conjunction with, the great fibro-elastic web c, to flex the forearm on the arm. The fibro-elastic web is more or less under the influence of the voluntary muscles (a, b).
o, p, q, Musculo-fibro-elastic ligament, which envelopes the roots of the primary and secondary feathers, and forms a symmetrical network of great strength and beauty, its component parts being arranged in such a manner as to envelope the root of each individual feather. The network in question supports the feathers, and limits their peculiar valvular action. It is enlarged at figs. 99 and 101, and consists of three longitudinal bands, r s, t u, v w. Between these bands two oblique bands, g and h, run; the oblique bands occurring between every two feathers. The marginal longitudinal band (v, w) splits up into two processes, one of which curves round the root of each feather (x) in a direction from right to left (c, b, a), the other in a direction from left to right (d, e, f). These processes are also seen at m, n of fig. 100.—Original.
The manner in which the roots of the primary, secondary, and tertiary feathers are geared to each other in order to rotate in one direction in flexion, and in another and opposite direction in extension, is shown at figs. 98, 99, 100, and 101, p. 181. In flexion the feathers open up and permit the air to pass between them. In extension they flap together and render the wing as air-tight as that of either the insect or bat. The primary, secondary, and tertiary feathers have consequently a valvular action.
The Wing of the Bird not always opened up to the same extent in the Up Stroke.—The elaborate arrangements and adaptations for increasing the area of the wing, and making it impervious to air during the down stroke, and for decreasing the area and opening up the wing during the up stroke, although necessary to the flight of the heavy-bodied, short-winged birds, as the grouse, partridge, and pheasant, are by no means indispensable to the flight of the long-winged oceanic birds, unless when in the act of rising from a level surface; neither do the short-winged heavy birds require to fold and open up the wing during the up stroke to the same extent in all cases, less folding and opening up being required when the birds fly against a breeze, and when they have got fairly under weigh. All the oceanic birds, even the albatross, require to fold and flap their wings vigorously when they rise from the surface of the water. When, however, they have acquired a certain degree of momentum, and are travelling at a tolerable horizontal speed, they can in a great measure dispense with the opening up of the wing during the up stroke—nay, more, they can in many instances dispense even with flapping. This is particularly the case with the albatross, which (if a tolerably stiff breeze be blowing) can sail about for an hour at a time without once flapping its wings. In this case the wing is wielded in one piece like the insect wing, the bird simply screwing and unscrewing the pinion on and off the wind, and exercising a restraining influence—the breeze doing the principal part of the work. In the bat the wing is jointed as in the bird, and folded during the up stroke. As, however, the bat’s wing, as has been already stated, is covered by a continuous and more or less elastic membrane, it follows that it cannot be opened up to admit of the air passing through it during the up stroke. Flight in the bat is therefore secured by alternately diminishing and increasing the area of the wing during the up and down strokes—the wing rotating upon its root and along its anterior margin, and presenting a variety of kite-like surfaces, during its ascent and descent, precisely as in the bird (fig. [80], p. 149, and fig. [83], p. 158).
Fig. 102.—Shows the upward inclination of the body and the flexed condition of the wings (a b, e f; a´ b´, e´ f´) in the flight of the kingfisher. The body and wings when taken together form a kite. Compare with fig. [59], p. 126, where the wings are fully extended.
Flexion of the Wing necessary to the Flight of Birds.—Considerable diversity of opinion exists as to whether birds do or do not flex their wings in flight. The discrepancy is owing to the great difficulty experienced in analysing animal movements, particularly when, as in the case of the wings, they are consecutive and rapid. My own opinion is, that the wings are flexed in flight, but that all wings are not flexed to the same extent, and that what holds true of one wing does not necessarily hold true of another. To see the flexing of the wing properly, the observer should be either immediately above the bird or directly beneath it. If the bird be diminishing from before, behind, or from the side, the up and down strokes of the pinion distract the attention and complicate the movement to such an extent as to render the observation of little value. In watching rooks proceeding leisurely against a slight breeze, I have over and over again satisfied myself that the wings are flexed during the up stroke, the mere extension and flexion, with very little of a down stroke, in such instances sufficing for propulsion. I have also observed it in the pigeon in full flight, and likewise in the starling, sparrow, and kingfisher (fig. 102, p. 183).
It occurs principally at the wrist-joint, and gives to the wing the peculiar quiver or tremor so apparent in rapid flight, and in young birds at feeding-time. The object to be attained is manifest. By the flexing of the wing in flight, the “remiges,” or rowing feathers, are opened up or thrown out of position, and the air permitted to escape—advantage being thus taken of the peculiar action of the individual feathers and the higher degree of differentiation perceptible in the wing of the bird as compared with that of the bat and insect.
In order to corroborate the above opinion, I extended the wings of several birds as in rapid flight, and fixed them in the outspread position by lashing them to light unyielding reeds. In these experiments the shoulder and elbow-joints were left quite free—the wrist or carpal and the metacarpal joints only being bound. I took care, moreover, to interfere as little as possible with the action of the elastic ligament or alar membrane which, in ordinary circumstances, recovers or flexes the wing, the reeds being attached for the most part to the primary and secondary feathers. When the wings of a pigeon were so tied up, the bird could not rise, although it made vigorous efforts to do so. When dropped from the hand, it fell violently upon the ground, notwithstanding the strenuous exertions which it made with its pinions to save itself. When thrown into the air, it fluttered energetically in its endeavours to reach the dove-cot, which was close at hand; in every instance, however, it fell, more or less heavily, the distance attained varying with the altitude to which it was projected.
Thinking that probably the novelty of the situation and the strangeness of the appliances confused the bird, I allowed it to walk about and to rest without removing the reeds. I repeated the experiment at intervals, but with no better results. The same phenomena, I may remark, were witnessed in the sparrow; so that I think there can be no doubt that a certain degree of flexion in the wings is indispensable to the flight of all birds—the amount varying according to the length and form of the pinions, and being greatest in the short broad-winged birds, as the partridge and kingfisher, less in those whose wings are moderately long and narrow, as the gulls, and many of the oceanic birds, and least in the heavy-bodied long and narrow-winged sailing or gliding birds, the best example of which is the albatross. The degree of flexion, moreover, varies according as the bird is rising, falling, or progressing in a horizontal direction, it being greatest in the two former, and least in the latter.
It is true that in insects, unless perhaps in those which fold or close the wing during repose, no flexion of the pinion takes place in flight; but this is no argument against this mode of diminishing the wing-area during the up stroke where the joints exist; and it is more than probable that when joints are present they are added to augment the power of the wing during its active state, i.e. during flight, quite as much as to assist in arranging the pinion on the back or side of the body when the wing is passive and the animal is reposing. The flexion of the wing is most obvious when the bird is exerting itself, and may be detected in birds which skim or glide when they are rising, or when they are vigorously flapping their wings to secure the impetus necessary to the gliding movement. It is less marked at the elbow-joint than at the wrist; and it may be stated generally that, as flexion decreases, the twisting flail-like movement of the wing at the shoulder increases, and vice versâ,—the great difference between sailing birds and those which do not sail amounting to this, that in the sailing birds the wing is worked from the shoulder by being alternately rolled on and off the wind, as in insects; whereas, in birds which do not glide, the spiral movement travels along the arm as in bats, and manifests itself during flexion and extension in the bending of the joints and in the rotation of the bones of the wing on their axes. The spiral conformation of the pinions, to which allusion has been so frequently made, is best seen in the heavy-bodied birds, as the turkey, capercailzie, pheasant, and partridge; and here also the concavo-convex form of the wing is most perceptible. In the light-bodied, ample-winged birds, the amount of twisting is diminished, and, as a result, the wing is more or less flattened, as in the sea-gull (fig. 103).
Fig. 103.—Shows the twisted levers or screws formed by the wings of the gull. Compare with fig. [53], p. 107; with figs. [76], 77, and 78, p. 147, and with figs. [82] and 83, p. 158.—-Original.
Consideration of the Forces which propel the Wings of Insects.—In the thorax of insects the muscles are arranged in two principal sets in the form of a cross—i.e. there is a powerful vertical set which runs from above downwards, and a powerful antero-posterior set which runs from before backwards. There are likewise a few slender muscles which proceed in a more or less oblique direction. The antero-posterior and vertical sets of muscles are quite distinct, as are likewise the oblique muscles. Portions, however, of the vertical and oblique muscles terminate at the root of the wing in jelly-looking points which greatly resemble rudimentary tendons, so that I am inclined to believe that the vertical and oblique muscles exercise a direct influence on the movements of the wing. The shortening of the antero-posterior set of muscles (indirectly assisted by the oblique ones) elevates the dorsum of the thorax by causing its anterior extremity to approach its posterior extremity, and by causing the thorax to bulge out or expand laterally. This change in the thorax necessitates the descent of the wing. The shortening of the vertical set (aided by the oblique ones) has a precisely opposite effect, and necessitates its ascent. While the wing is ascending and descending the oblique muscles cause it to rotate on its long axis, the bipartite division of the wing at its root, the spiral configuration of the joint, and the arrangement of the elastic and other structures which connect the pinion with the body, together with the resistance it experiences from the air, conferring on it the various angles which characterize the down and up strokes. The wing may therefore be said to be depressed by the shortening of the antero-posterior set of muscles, aided by the oblique muscles, and elevated by the shortening of the vertical and oblique muscles, aided by the elastic ligaments, and the reaction of the air. If we adopt this view we have a perfect physiological explanation of the phenomenon, as we have a complete circle or cycle of motion, the antero-posterior set of muscles shortening when the vertical set of muscles are elongating, and vice versâ. This, I may add, is in conformity with all other muscular arrangements, where we have what are usually denominated extensors and flexors, pronators and supinators, abductors and adductors, etc., but which, as I have already explained (pp. [24] to 34), are simply the two halves of a circle of muscle and of motion, an arrangement for securing diametrically opposite movements in the travelling surfaces of all animals.
Chabrier’s account, which I subjoin, virtually supports this hypothesis:—
“It is generally through the intervention of the proper motions of the dorsum, which are very considerable during flight, that the wings or the elytra are moved equally and simultaneously. Thus, when it is elevated, it carries with it the internal side of the base of the wings with which it is articulated, from which ensues the depression of the external side of the wing; and when it approaches the sternal portion of the trunk, the contrary takes place. During the depression of the wings, the dorsum is curved from before backwards, or in such a manner that its anterior extremity is brought nearer to its posterior, that its middle is elevated, and its lateral portions removed further from each other. The reverse takes place in the elevation of the wings; the anterior extremity of the dorsum being removed to a greater distance from the posterior, its middle being depressed, and its sides brought nearer to each other. Thus its bending in one direction produces a diminution of its curve in the direction normally opposed to it; and by the alternations of this motion, assisted by other means, the body is alternately compressed and dilated, and the wings are raised and depressed by turns.”[86]
In the libellulæ or dragon-flies, the muscles are inserted into the roots of the wings as in the bat and bird, the only difference being that in the latter the muscles creep along the wings to their extremities.
In all the wings which I have examined, whether in the insect, bat, or bird, the wings are recovered, flexed, or drawn towards the body by the action of elastic ligaments, these structures, by their mere contraction, causing the wings, when fully extended and presenting their maximum of surface, to resume their position of rest, and plane of least resistance. The principal effort required in flight would therefore seem to be made during extension and the down stroke. The elastic ligaments are variously formed, and the amount of contraction which they undergo is in all cases accurately adapted to the size and form of the wings, and the rapidity with which they are worked—the contraction being greatest in the short-winged and heavy-bodied insects and birds, and least in the light-bodied and ample-winged ones, particularly in such as skim or glide. The mechanical action of the elastic ligaments, I need scarcely remark, insures a certain period of repose to the wings at each stroke, and this is a point of some importance, as showing that the lengthened and laborious flights of insects and birds are not without their stated intervals of rest.
Speed attained by Insects.—Many instances might be quoted of the marvellous powers of flight possessed by insects as a class. The male of the silkworm-moth (Attacus Paphia) is stated to travel more than 100 miles a day;[87] and an anonymous writer in Nicholson’s Journal[88] calculates that the common house-fly (Musca domestica), in ordinary flight, makes 600 strokes per second, and advances twenty-five feet, but that the rate of speed, if the insect be alarmed, may be increased six or seven fold, so that under certain circumstances it can outstrip the fleetest racehorse. Every one when riding on a warm summer day must have been struck with the cloud of flies which buzz about his horse’s ears even when the animal is urged to its fastest paces; and it is no uncommon thing to see a bee or a wasp endeavouring to get in at the window of a railway car in full motion. If a small insect like a fly can outstrip a racehorse, an insect as large as a horse would travel very much faster than a cannon-ball. Leeuwenhoek relates a most exciting chase which he once beheld in a menagerie about 100 feet long between a swallow and a dragon-fly (Mordella). The insect flew with incredible speed, and wheeled with such address, that the swallow, notwithstanding its utmost efforts, completely failed to overtake and capture it.[89]
Consideration of the Forces which propel the Wings of Bats and Birds.—The muscular system of birds has been so frequently and faithfully described, that I need not refer to it further than to say that there are muscles which by their action are capable of elevating and depressing the wings, and of causing them to move in a forward and backward direction, and obliquely. They can also extend or straighten and bend, or flex the wings, and cause them to rotate in the direction of their length during the down and up strokes. The muscles principally concerned in the elevation of the wings are the smaller pectoral or breast muscles (pectorales minor); those chiefly engaged in depressing the wings are the larger pectorals (pectorales major). The pectoral muscles correspond to the fleshy mass found on the breast-bone or sternum, which in flying birds is boat-shaped, and furnished with a keel. These muscles are sometimes so powerful and heavy that they outweigh all the other muscles of the body. The power of the bird is thus concentrated for the purpose of moving the wings and conferring steadiness upon the volant mass. In birds of strong flight the keel is very large, in order to afford ample attachments for the muscles delegated to move the wings. In birds which cannot fly, as the members of the ostrich family, the breast-bone or sternum has no keel.[90]
The remarks made regarding the muscles of birds, apply with very slight modifications to the muscles of bats. The muscles of bats and birds, particularly those of the wings, are geared to, and act in concert with, elastic ligaments or membranes, to be described presently.
Lax condition of the Shoulder-Joint in Bats, Birds, etc.—The great laxity of the shoulder-joint in bats and birds, readily admits of their bodies falling downwards and forwards during the up stroke. This joint, as has been already stated, admits of movement in every direction, so that the body of the bat or bird is like a compass set upon gimbals, i.e. it swings and oscillates, and is equally balanced, whatever the position of the wings. The movements of the shoulder-joint in the bird, bat, and insect are restrained within certain limits by a system of check ligaments and prominences; but in each case the range of motion is very great, the wings being permitted to swing forwards, backwards, upwards, downwards, or at any degree of obliquity. They are also permitted to rotate along their anterior margin, or to twist in the direction of their length to the extent of nearly a quarter of a turn. This great freedom of movement at the shoulder-joint enables the insect, bat, and bird to rotate and balance upon two centres—the one running in the direction of the length of the body, the other at right angles or across the body, i.e. in the direction of the length of the wings.
In the bird the head of the humerus is convex and somewhat oval (not round), the long axis of the oval being directed from above downwards, i.e. from the dorsal towards the ventral aspect of the bird. The humerus can, therefore, glide up and down in the facettes occurring on the articular ends of the coracoid and scapular bones with great facility, much in the same way that the head of the radius glides upon the distal end of the humerus. But the humerus has another motion; it moves like a hinge from before backwards, and vice versâ. The axis of the latter movement is almost at right angles to that of the former. As, however, the shoulder-joint is connected by long ligaments to the body, and can be drawn away from it to the extent of one-eighth of an inch or more, it follows that a third and twisting movement can be performed, the twisting admitting of rotation to the extent of something like a quarter of a turn. In raising and extending the wing preparatory to the downward stroke two opposite movements are required, viz. one from before backwards, and another from below upwards. As, however, the axes of these movements are at nearly right angles to each other, a spiral or twisting movement is necessary to run the one into the other—to turn the corner, in fact.
From what has been stated it will be evident that the movements of the wing, particularly at the root, are remarkably free, and very varied. A directing and restraining, as well as a propelling force, is therefore necessary.
The guiding force is to be found in the voluntary muscles which connect the wing with the body in the insect, and which in the bat and bird, in addition to connecting the wing with the body, extend along the pinion even to its tip. It is also to be found in the musculo-elastic and other ligaments, seen to advantage in the bird.
The Wing flexed and partly elevated by the Action of Elastic Ligaments—the Nature and Position of such Ligaments in the Pheasant, Snipe, Crested Crane, Swan, etc.—When the wing is drawn away from the body of the bird by the hand the posterior margin of the pinion formed by the primary, secondary, and tertiary feathers rolls down to make a variety of inclined surfaces with the horizon (c b, of fig. [63], p. 138). When, however, the hand is withdrawn, even in the dead bird, the wing instantly folds up; and in doing so reduces the amount of inclination in the several surfaces referred to (c b, d e f of the same figure). The wing is folded by the action of certain elastic ligaments, which are put upon the stretch in extension, and which recover their original form and position in flexion (fig. [98], c, p. 181). This simple experiment shows that the various inclined surfaces requisite for flight are produced by the mere acts of extension and flexion in the dead bird. It is not, however, to be inferred from this circumstance that flight can be produced without voluntary movements any more than ordinary walking. The muscles, bones, ligaments, feathers, etc., are so adjusted with reference to each other that if the wing is moved at all, it must move in the proper direction—an arrangement which enables the bird to fly without thinking, just as we can walk without thinking. There cannot, however, be a doubt that the bird has the power of controlling its wings both during the down and up strokes; for how otherwise could it steer and direct its course with such precision in obtaining its food? how fix its wings on a level with or above its body for skimming purposes? how fly in a curve? how fly with, against, or across a breeze? how project itself from a rock directly into space, or how elevate itself from a level surface by the laboured action of its wings?
The wing of the bird is elevated to a certain extent in flight by the reaction of the air upon its under surface; but it is also elevated by muscular action—by the contraction of the elastic ligaments, and by the body falling downwards and forwards in a curve.
That muscular action is necessary is proved by the fact that the pinion is supplied with distinct elevator muscles.[91] It is further proved by this, that the bird can, and always does, elevate its wings prior to flight, quite independently of the air. When the bird is fairly launched in space the elevator muscles are assisted by the tendency which the body has to fall downwards and forwards: by the reaction of the air; and by the contraction of the elastic ligaments. The air and the elastic ligaments contribute to the elevation of the wing, but both are obviously under control—they, in fact, form links in a chain of motion which at once begins and terminates in the muscular system.
That the elastic ligaments are subsidiary and to a certain extent under the control of the muscular system in the same sense that the air is, is evident from the fact that voluntary muscular fibres run into the ligaments in question at various points (a, b of fig. [98], p. 181). The ligaments and muscular fibres act in conjunction, and fold or flex the forearm on the arm. There are others which flex the hand upon the forearm. Others draw the wing towards the body.
The elastic ligaments, while occupying a similar position in the wings of all birds, are variously constructed and variously combined with voluntary muscles in the several species.
The Elastic Ligaments more highly differentiated in Wings which vibrate rapidly.—The elastic ligaments of the swan are more complicated and more liberally supplied with voluntary muscle than those of the crane, and this is no doubt owing to the fact that the wings of the swan are driven at a much higher speed than those of the crane. In the snipe the wings are made to vibrate very much more rapidly than in the swan, and, as a consequence, we find that the fibro-elastic bands are not only greatly increased, but they are also geared to a much greater number of voluntary muscles, all which seems to prove that the musculo-elastic apparatus employed for recovering or flexing the wing towards the end of the down stroke, becomes more and more highly differentiated in proportion to the rapidity with which the wing is moved.[92] The reason for this is obvious. If the wing is to be worked at a higher speed, it must, as a consequence, be more rapidly flexed and extended. The rapidity with which the wing of the bird is extended and flexed is in some instances exceedingly great; so great, in fact, that it escapes the eye of the ordinary observer. The speed with which the wing darts in and out in flexion and extension would be quite inexplicable, but for a knowledge of the fact that the different portions of the pinion form angles with each other, these angles being instantly increased or diminished by the slightest quiver of the muscular and fibro-elastic systems. If we take into account the fact that the wing of the bird is recovered or flexed by the combined action of voluntary muscles and elastic ligaments; that it is elevated to a considerable extent by voluntary muscular effort; and that it is extended and depressed entirely by muscular exertion, we shall have difficulty in avoiding the conclusion that the wing is thoroughly under the control of the muscular system, not only in flexion and extension, but also throughout the entire down and up strokes.
An arrangement in every respect analogous to that described in the bird is found in the wing of the bat, the covering or web of the wing in this instance forming the principal elastic ligament (fig. [17], p. 36).
Power of the Wing—to what owing.—The shape and power of the pinion depend upon one of three circumstances—to wit, the length of the humerus,[93] the length of the cubitus or forearm, and the length of the primary feathers. In the swallow the humerus, and in the humming-bird the cubitus, is very short, the primaries being very long; whereas in the albatross the humerus or arm-bone is long and the primaries short. When one of these conditions is fulfilled, the pinion is usually greatly elongated and scythe-like (fig. [62], p. 137)—an arrangement which enables the bird to keep on the wing for immense periods with comparatively little exertion, and to wheel, turn, and glide about with exceeding ease and grace. When the wing is truncated and rounded (fig. [96], p. 176), a form of pinion usually associated with a heavy body, as in the grouse, quail, diver, and grebe, the muscular exertion required, and the rapidity with which the wing moves are very great; those birds, from a want of facility in turning, flying either in a straight line or making large curves. They, moreover, rise with difficulty, and alight clumsily and somewhat suddenly. Their flight, however, is perfect while it lasts.
The goose, duck (fig. [107], p. 204), pigeon (fig. [106], p. 203) and crow, are intermediate both as regards the form of the wing and the rapidity with which it is moved.
The heron (fig. [60], p. 126) and humming-bird furnish extreme examples in another direction,—the heron having a large wing with a leisurely movement, the humming-bird a comparatively large wing with a greatly accelerated one.
But I need not multiply examples; suffice it to say that flight may be attained within certain limits by every size and form of wing, if the number of its oscillations be increased in proportion to the weight to be raised.
Reasons why the effective Stroke should be delivered downwards and forwards.—The wings of all birds, whatever their form, act by alternately presenting oblique and comparatively non-oblique surfaces to the air,—the mere extension of the pinion, as has been shown, causing the primary, secondary, and tertiary feathers to roll down till they make an angle of 30° or so with the horizon, in order to prepare it for giving the effective stroke, which is delivered, with great rapidity and energy, in a downward and forward direction. I repeat, “downwards and forwards;” for a careful examination of the relations of the wing in the dead bird, and a close observation of its action in the living one, supplemented by a large number of experiments with natural and artificial wings, have fully convinced me that the stroke is invariably delivered in this direction.[94] If the wing did not strike downwards and forwards, it would act at a manifest disadvantage:—
1st. Because it would present the back or convex surface of the wing to the air—a convex surface dispersing or dissipating the air, while a concave surface gathers it together or focuses it.
2d. In order to strike backwards effectually, the concavity of the wing would also require to be turned backwards; and this would involve the depression of the anterior or thick margin of the pinion, and the elevation of the posterior or thin one, during the down stroke, which never happens.
3d. The strain to which the pinion is subjected in flight would, if the wing struck backwards, fall, not on the anterior or strong margin of the pinion formed by the bones and muscles, but on the posterior or weak margin formed by the tips of the primary, secondary, and tertiary feathers—which is not in accordance with the structure of the parts.
4th. The feathers of the wing, instead of being closed, as they necessarily are, by a downward and forward movement, would be inevitably opened, and the integrity of the wing impaired by a downward and backward movement.
5th. The disposition of the articular surfaces of the wing (particularly that of the shoulder-joint) is such as to facilitate the downward and forward movement, while it in a great measure prevents the downward and backward one.
6th and lastly. If the wing did in reality strike downwards and backwards, a result the converse of that desired would most assuredly be produced, as an oblique surface which smites the air in a downward and backward direction (if left to itself) tends to depress the body bearing it. This is proved by the action upon the air of free inclined planes, arranged in the form of a screw.
The Wing acts as an Elevator, Propeller, and Sustainer, both during extension and flexion.—The wing, as has been explained, is recovered or drawn off the wind principally by the contraction of the elastic ligaments extending between the joints, so that the pinion during flexion enjoys a certain degree of repose. The time occupied in recovering is not lost so long as the wing makes an angle with the horizon and the bird is in motion, it being a matter of indifference whether the wing acts on the air, or the air on the wing, so long as the body bearing the latter is under weigh; and this is the chief reason why the albatross, which is a very heavy bird,[95] can sail about for such incredible periods without flapping the wings at all. Captain Hutton thus graphically describes the sailing of this magnificent bird:—“The flight of the albatross is truly majestic, as with outstretched motionless wings he sails over the surface of the sea—now rising high in air, now with a bold sweep, and wings inclined at an angle with the horizon, descending until the tip of the lower one all but touches the crest of the waves as he skims over them.”[96]
Birds of Flight divisible into four kinds:—
1st. Such as have heavy bodies and short wings with a rapid movement (fig. [59], p. 126).
2d. Such as have light bodies and large wings with a leisurely movement (fig. [60], p. 126; fig. [103], p. 186).
3d. Such as have heavy bodies and long narrow wings with a decidedly slow movement (fig. [105], p. 200).
4th. Such as are intermediate with regard to the size of body, the dimensions of the wing, and the energy with which it is driven (fig. [102], p. 183; fig. [106], p. 203; fig. [107], p. 204).
They may be subdivided into those which float, skim, or glide, and those which fly in a straight line and irregularly.
The pheasant, partridge (fig. [59], p. 126), grouse, and quail, furnish good examples of the heavy-bodied, short-winged birds. In these the wing is rounded and deeply concave. It is, moreover, wielded with immense velocity and power.
The heron (fig. [60], p. 126), sea-mew (fig. [103], p. 186), lapwing (fig. [63], p. 138), and owl (fig. 104), supply examples of the second class, where the wing, as compared with the body, is very ample, and where consequently it is moved more leisurely and less energetically.
Fig. 104.—The Cape Barn-Owl (Strix capensis, Smith), as seen in full flight, hunting. The under surface of the wings and body are inclined slightly upwards, and act upon the air after the manner of a kite. (Compare with fig. [59], p. 126, and fig. [102], p. 183.)—Original.
The albatross (fig. [105], p. 200) and pelican afford instances of the third class, embracing the heavy-bodied, long-winged birds.
The duck (fig. [107], p. 204), pigeon (fig. [106], p. 203), crow and thrush, are intermediate, both as regards the size of the wing and the rapidity with which it is made to oscillate. These constitute the fourth class.
The albatross (fig. [105], p. 200), swallow, eagle, and hawk, provide instances of sailing or gliding birds, where the wing is ample, elongated, and more or less pointed, and where advantage is taken of the weight of the body and the shape of the pinion to utilize the air as a supporting medium. In these the pinion acts as a long lever,[97] and is wielded with great precision and power, particularly at the shoulder.
The Flight of the Albatross compared to the Movements of a Compass set upon Gimbals.—A careful examination of the movements in skimming birds has led me to conclude that by a judicious twisting or screw-like action of the wings at the shoulder, in which the pinions are alternately advanced towards and withdrawn from the head in a manner analogous to what occurs at the loins in skating without lifting the feet, birds of this order can not only maintain the motion which they secure by a few energetic flappings, but, if necessary, actually increase it, and that without either bending the wing or beating the air.
The forward and backward screwing action of the pinion referred to, in no way interferes, I may remark, with the rotation of the wing on its long axis, the pinion being advanced and screwed down upon the wind, and retracted and unscrewed alternately. As the movements described enable the sailing bird to tilt its body from before backwards, or the converse, and from side to side or laterally, it may be represented as oscillating on one of two centres, as shown at fig. 105; the one corresponding with the long axis of the body (fig. 105, a b), the other with the long axis of the wings (c d). Between these two extremes every variety of sailing and gliding motion which is possible in the mariner’s compass when set upon gimbals may be performed; so that a skimming or sailing bird may be said to possess perfect command over itself and over the element in which it moves.
Fig. 105.
Captain Hutton makes the following remarkable statement regarding the albatross:—“I have sometimes watched narrowly one of these birds sailing and wheeling about in all directions for more than an hour, without seeing the slightest movement of the wings, and have never witnessed anything to equal the ease and grace of this bird as he sweeps past, often within a few yards, every part of his body perfectly motionless except the head and eye, which turn slowly and seem to take notice of everything.”[98]
“Tranquil its spirit seem’d and floated slow;
Even in its very motion there was rest.”[99]
As an antithesis to the apparently lifeless wings of the albatross, the ceaseless activity of those of the humming-bird may be adduced. In those delicate and exquisitely beautiful birds, the wings, according to Mr. Gould, move so rapidly when the bird is poised before an object, that it is impossible for the eye to follow each stroke, and a hazy circle of indistinctness on each side of the bird is all that is perceptible. When the humming-bird flies in a horizontal direction, it occasionally proceeds with such velocity as altogether to elude observation.
The regular and irregular in Flight.—The coot, diver, duck, and goose fly with great regularity in nearly a straight line, and with immense speed; they rarely if ever skim or glide, their wings being too small for this purpose. The woodpecker, magpie, fieldfare and sparrow, supply examples of what may be termed the “irregular” in flight. These, as is well known, fly in curves of greater or less magnitude, by giving a few vigorous strokes and then desisting, the effect of which is to project them along a series of parabolic curves. The snipe and woodcock are irregular in another respect, their flight being sudden, jerky, and from side to side.
Mode of ascending, descending, turning, etc.—All birds which do not, like the swallow and humming-birds, drop from a height, raise themselves at first by a vigorous leap, in which they incline their bodies in an upward direction, the height thus attained enabling them to extend and depress their wings without injury to the feathers. By a few sweeping strokes delivered downwards and forwards, in which the wings are made nearly to meet above and below the body, they lever themselves upwards and forwards, and in a surprisingly short time acquire that degree of momentum which greatly assists them in their future career. In rising from the ground, as may readily be seen in the crow, pigeon, and kingfisher (fig. [102], p. 183), the tail is expanded and the neck stretched out, so that the body is converted into an inclined plane, and acts mechanically as a kite. The centre of gravity and the position of the body are changed at the will of the bird by movements in the neck, feet, and tail, and by increasing or decreasing the angles which the under surface of the wings makes with the horizon. When a bird wishes to fly in a horizontal direction, it causes the under surface of its wings to make a slight forward angle with the horizon. When it wishes to ascend, the angle is increased. When it wishes to descend, it causes the under surface of the wings to make a slight backward angle with the horizon. When a bird flies up, its wings strike downwards and forwards. When it flies down, its wings strike downwards and backwards. When a sufficient altitude has been attained, the length of the downward stroke is generally curtailed, the mere extension and flexion of the wing, assisted by the weight of the body, in such instances sufficing. This is especially the case if the bird is advancing against a slight breeze, the effort required under such circumstances being nominal in amount. That little power is expended is proved by the endless gyrations of rooks and other birds; these being continued for hours together. In birds which glide or skim, it has appeared to me that the wing is recovered much more quickly, and the down stroke delivered more slowly, than in ordinary flight—in fact, that the rapidity with which the wing acts in an upward and downward direction is, in some instances, reversed; and this is what we should naturally expect when we recollect that in gliding, the wings require to be, for the most part, in the expanded condition. If this observation be correct, it follows that birds have the power of modifying the duration of the up and down strokes at pleasure. Although the wing of the bird usually strikes the air at an angle which varies from 15° to 30°, the angle may be increased to such an extent as to subvert the position of the bird. The tumbler pigeon, e.g. can, by slewing its wings forwards and suddenly throwing back its head, turn a somersault. When birds are fairly on the wing they have the air, unless when that is greatly agitated by a storm, completely under control. This arises from their greater specific gravity, and because they are possessed of independent motion. If they want to turn, they have simply to tilt their bodies laterally, as a railway carriage would be tilted in taking a curve,[100] or to increase the number of beats given by the one wing as compared with the other; or to keep the one wing extended while the other is partially flexed. The neck, feet, and tail may or may not contribute to this result. If the bird wishes to rise, it tilts its entire body (the neck and tail participating) in an upward direction (fig. [59], p. 126; fig. [102], p. 183); or it rises principally by the action of the wings and by muscular efforts, as happens in the lark. The bird can in this manner likewise retain its position in the air, as may be observed in the hawk when hovering above its prey. If the bird desires to descend, it may reverse the direction of the inclined plane formed by the body and wings, and plunge head foremost with extended pinions (fig. 106); or it may flex the wings, and so accelerate its pace; or it may raise its wings and drop parachute-fashion (fig. [55], p. 112; g, g of fig. [82], p. 158); or it may even fly in a downward direction—a few sudden strokes, a more or less abrupt curve, and a certain degree of horizontal movement being in either case necessary to break the fall previous to alighting (fig. 107, below). Birds which fish on the wing, as the osprey and gannet, precipitate themselves from incredible heights, and drop into the water with the velocity of a meteorite—the momentum which they acquire during their descent materially aiding them in their subaqueous flight. They emerge from the water and are again upon the wing before the eddies occasioned by their precipitous descent have well subsided, in some cases rising apparently without effort, and in others running along and beating the surface of the water for a brief period with their pinions and feet.
Fig. 106.—The Pigeon (Treron bicincta, Jerdon), flying downwards and turning prior to alighting. The pigeon expands its tail both in ascending and descending.—Original.
Fig. 107.—The Red-headed Pochard (Fuligula ferina, Linn.) in the act of dropping upon the water; the head and body being inclined upwards and forwards, the feet expanded, and the wings delivering vigorous short strokes in a downward and forward direction.—Original.
The Flight of Birds referable to Muscular Exertion and Weight.—The various movements involved in ascending, descending, wheeling, gliding, and progressing horizontally, are all the result of muscular power and weight, properly directed and acting upon appropriate surfaces—that apparent buoyancy in birds which we so highly esteem, arising not from superior lightness, but from their possessing that degree of solidity which enables them to subjugate the air,—weight and independent motion, i.e. motion associated with animal life, or what is equivalent thereto, being the two things indispensable in successful aërial progression. The weight in insects and birds is in great measure owing to their greatly developed muscular system, this being in that delicate state of tonicity which enables them to act through its instrumentality with marvellous dexterity and power, and to expend or reserve their energies, which they can do with the utmost exactitude, in their apparently interminable flights.
Lifting-capacity of Birds.—The muscular power in birds is usually greatly in excess, particularly in birds of prey, as, e.g. the condors, eagles, hawks, and owls. The eagles are remarkable in this respect—these having been known to carry off young deer, lambs, rabbits, hares, and, it is averred, even young children. Many of the fishing birds, as the pelicans and herons, can likewise carry considerable loads of fish;[101] and even the smaller birds, as the records of spring show, are capable of transporting comparatively large twigs for building purposes. I myself have seen an owl, which weighed a little over 10 ounces, lift 2 1/2 ounces, or a quarter of its own weight, without effort, after having fasted twenty-four hours; and a friend informs me that a short time ago a splendid osprey was shot at Littlehampton, on the coast of Sussex, with a fish 5 lbs. weight in its mouth.
There are many points in the history and economy of birds which crave our sympathy while they elicit our admiration. Their indubitable courage and miraculous powers of flight invest them with a superior dignity, and secure for their order almost a duality of existence. The swallow, tiny and inconsiderable as it may appear, can traverse 1000 miles at a single journey; and the albatross, despising compass and landmark, trusts himself boldly for weeks together to the mercy or fury of the mighty ocean. The huge condor of the Andes lifts himself by his sovereign will to a height where no sound is heard, save the airy tread of his vast pinions, and, from an unseen point, surveys in solitary grandeur the wide range of plain and pasture-land;[102] while the bald eagle, nothing daunted by the din and indescribable confusion of the queen of waterfalls, the stupendous Niagara, sits composedly on his giddy perch, until inclination or desire prompts him to plunge into or soar above the drenching mists which, shapeless and ubiquitous, perpetually rise from the hissing waters of the nether caldron.
Fig. 108.—Hawk and quarry.—After The Graphic.
AËRONAUTICS
The Vauxhall Balloon of Mr. Green.