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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 versa. 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 (cb, 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 When, however, the hand is withdrawn, even in the dead bird, the wing instantly folds up; and in doing so reduces the amount of inclination in the several surfaces referred to (cb, def of the same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.1 It is further proved by this, that the bird can, and always

1 Mr. Macgillivray and C. J. L. Krarup, a Danish author, state that the wing is elevated by a vital force, viz. by the contraction of the pectoralis minor. This muscle, according to Krarup, acts with one-eighth the intensity of the pectoralis major (the depressor of the wing). He bases his statement upon the fact that in the pigeon the pectoralis minor or elevator of the wing weighs one-eighth of an ounce, whereas the pectoralis major or depressor of the wing weighs seven-eighths of an ounce. It ought, however, to be borne in mind that the volume of a muscle does not necessarily determine the precise influence exerted by its action; for the tendon of the muscle may be made to act upon a long lever, and, under favourable conditions, for developing its powers, while that of another muscle may be made to act upon a short lever, and, consequently, under unfavourable conditions.-On the Flight of Birds, p. 30. Copenhagen, 1869.

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.1 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

1 A careful account of the musculo-elastic structures occurring in the wing of the pigeon is given by Mr. Macgillivray in his History of British Birds, pp. 37, 38.

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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,1 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.

1 "The humerus varies extremely in length, being very short in the swallow, of moderate length in the gallinaceous birds, longer in the crows, very long in the gannets, and unusually elongated in the albatross. In the golden eagle it is also seen to be of great length."--Macgillivray's British Birds, vol. i. p. 30.

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 nonoblique 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.1 If the wing did not strike

1 Prevailing Opinions as to the Direction of the Down Stroke.-Mr. Macgillivray, in his History of British Birds, published in 1837, states (p. 34) that in flexion the wing is drawn upwards, forwards, and inwards, but that during extension, when the effective stroke is given, it is made to strike outwards, downwards, and backwards. The Duke of Argyll holds a similar opinion. In speaking of the hovering of birds, he asserts that

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