Part 18

_The Bones of the Wing of the Bat--the spiral configuration of their articular surfaces._--The bones of the arm and hand are especially deserving of attention. The humerus (fig. 17, _r_, p. 36) is short and powerful, and twisted upon itself to the extent of something less than a quarter of a turn. As a consequence, the long axis of the shoulder-joint is nearly at right angles to that of the elbow-joint. Similar remarks may be made regarding the radius (the principal bone of the forearm) (_d_), and the second and third metacarpal bones with their phalanges (_e f_), all of which are greatly elongated, and give strength and rigidity to the anterior or thick margin of the wing. The articular surfaces of the bones alluded to, as well as of the other bones of the hand, are spirally disposed with reference to each other, the long axes of the joints intersecting at nearly right angles. The object of this arrangement is particularly evident when the wing of the living bat, or of one recently dead, is extended and flexed as in flight.

In the flexed state the wing is greatly reduced in size, its under surface being nearly parallel with the plane of progression. When the wing is fully extended its under surface makes a certain angle with the horizon, the wing being then in a position to give the down stroke, which is delivered _downwards_ and _forwards_, as in the insect. When extension takes place the elbow-joint is depressed and carried forwards, the wrist elevated and carried backwards, the metacarpo-phalangeal joints lowered and inclined forwards, and the distal phalangeal joints slightly raised and carried backwards. The movement of the bat’s wing in extension is consequently a spiral one, the spiral running alternately from below upwards and forwards, and from above downwards and backwards (compare with fig. 79, p. 147). As the bones of the arm, forearm, and hand rotate on their axes during the extensile act, it follows that the posterior or thin margin of the wing is rotated in a downward direction (the anterior or thick one being rotated in an opposite direction) until the wing makes an angle of something like 30° with the horizon, which, as I have already endeavoured to show, is the greatest angle made by the wing in flight. The action of the bat’s wing at the shoulder is particularly free, partly because the shoulder-joint is universal in its nature, and partly because the scapula participates in the movements of this region. The freedom of action referred to enables the bat not only to rotate and twist its wing as a whole, with a view to diminishing and increasing the angle which its under surface makes with the horizon, but to elevate and depress the wing, and move it in a forward and backward direction. The rotatory or twisting movement of the wing is an essential feature in flight, as it enables the bat (and this holds true also of the insect and bird) to balance itself with the utmost exactitude, and to change its position and centre of gravity with marvellous dexterity. The movements of the shoulder-joint are restrained within certain limits by a system of check-ligaments, and by the coracoid and acromian processes of the scapula. The wing is recovered or flexed by the action of elastic ligaments which extend between the shoulder, elbow, and wrist. Certain elastic and fibrous structures situated between the fingers and in the substance of the wing generally take part in flexion. The bat flies with great ease and for lengthened periods. Its flight is remarkable for its softness, in which respect it surpasses the owl and the other nocturnal birds. The action of the wing of the bat, and the movements of its component bones, are essentially the same as in the bird.

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.

[84] The furcula are usually united to the anterior part of the sternum by ligament; but in birds of powerful flight, where the wings are habitually extended for gliding and sailing, as in the frigate-bird, the union is osseous in its nature. “In the frigate-bird the furcula are likewise anchylosed with the coracoid bones.”--Comp. Anat. and Phys. of Vertebrates, by Prof. Owen, vol. ii. p. 66.

[85] “The os humeri, or bone of the arm, is articulated by a small rounded surface to a corresponding cavity formed between the coracoid bone and the scapula, in such a manner as to allow great freedom of motion.”--Macgillivray’s Brit. Birds, vol. i. p. 33.

“The arm is articulated to the trunk by a ball-and-socket joint, permitting all the freedom of motion necessary for flight.”--Cyc. of Anat. and Phys., vol. iii. p. 424.

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.

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).

_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 contemplated 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).

_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:--