We have seen upon a former occasion the great variety of movements that insects can perform, and of the external organs with which they perform them[801]: but we are now to consider the internal apparatus, by the immediate action of which they take place—their system of muscles. When we reflect upon the wonderful velocity, their size considered, with which many insects move, and the unparalleled degree of muscular force that many exert[802], we feel no small degree of curiosity to know something of that part of their internal structure that produces these almost incredible effects. I shall in the present letter endeavour in some degree to gratify that curiosity, and give you an account of the muscles of these little animals,—first considering them in general; and then, as far as my information goes, adverting to those in particular that move the different parts and organs of an insect's body.
I. The muscles of insects may be considered in general as to their Origin; Substance and Parts; Shape; Colour; Kinds; Attachment; and Motions.
i. Origin. The origin of the muscular fibre in the higher animals is from the blood, the globules of which, by their coagulation in a series, appear to form it[803]; and in insects it is derived from the same universal source of nutrition and accretion, but not till it has been concreted into the adipose tissue or epiploon before noticed[804]. In the pupa of the cabbage-butterfly, Herold observed that this substance first assumed a fine flocky appearance and a blue-green colour, and that from it so changed were produced tender bundles of muscular fibres, extending in various directions, the epiploon itself decreasing in proportion as they were formed[805].
ii. Substance and Parts. The muscular fibre in vertebrate animals appears to consist of globules arranged in a series, and of no larger diameter than those of the blood,—the mean diameter of which in the human subject, when measured under the microscope by a micrometer, is found to be about 1⁄5000th part of an inch[806]. When Cuvier published his immortal work in 1805, the powers of any magnifier then constructed were not sufficient to enable this great physiologist to arrive at the simple fibre[807]; but Mr. Bauer, by the use of improved glasses, amongst other discoveries that will immortalize his name, was the first to detect, under the directions of Sir E. Home, the ultimate thread of which the muscular bundles are composed[808]. Chemists distinguish the substance of which we are speaking, by the name of fibrine. By the abundance of azote or nitrogen that enters into its composition, it possesses a character of animalization more marked than any other animal substance; and its elements are so approximated in the blood, that the slightest stagnation causes them to coagulate: and the muscles are without doubt, in the living subject, the only organs that can separate this matter from the mass of blood, and appropriate it to themselves[809]. The primary bundles of muscles are formed of the simple fibres, and the secondary are the result of an aggregation of the primary. The smaller bundles are not always exactly parallel to each other, but must in many cases diverge more or less, to produce those variations in shape observable in the muscles themselves: there are intervals therefore between the bundles, which in some animals are filled by a cellular substance[810]. Probably much of this statement will apply in most instances to the muscles of insects, but we may conclude that the globules that form them are infinitely smaller[811]. Lyonet has given some interesting observations with regard to those of the caterpillar of the Cossus: he describes them as of a soft transparent substance, capable of great extension, covered and filled by silver tubes of the bronchiæ, penetrated by the nerves, and containing oily particles. Each muscle was enveloped in membrane, and was composed of many parallel bands, consisting of bundles of fibres enveloped likewise in separate membranes. The fibres themselves, (but it is doubtful whether he arrived at the ultimate term of muscular fibre,) in a favourable light and under a good magnifier, appeared to be twisted spirally[812]. In spiders the muscles seemed to him to consist of two substances, the one soft and the other hard, the last forming a kind of stiff twisted filament[813]. A muscle thus composed of different bundles of fibres may be stated as to its parts, in insects, to consist of base, middle, and apex: the base is that part by which they are fixed to any given point of the internal surface of the crust, or of one of its processes, which serves as their fulcrum; the apex is that part by which they are fixed, either mediately or immediately, to the organ to be moved; and the middle is the remainder of the muscle. We usually discover in them no inflation of the middle corresponding with the belly of the muscles in vertebrate animals; they occasionally, however, terminate in a tendon, as those of the thighs and legs; but these tendons are of a different nature from the fibrous ones of warm-blooded animals; for they are hard, elastic, and without apparent fibres: the fleshy ones of the muscle envelope them, and are inserted in their surface[814].
iii. Shape. The muscles of insects are usually linear, with parallel sides; some are cylindrical, as those of the wings of the Libellulina[815]; and others, as those that move the legs in the caterpillar of the Cossus, are triangular[816]. In the suctorious mandibles of the grub of a common water-beetle[817] they are penniform, or shaped like a feather; and some in the Cossus are forked[818]. Under this head I may also observe, that the muscles are sometimes extremely slender threads, crossing each other, and often curiously interwoven in various directions, so as to resemble lace or fine gauze, as may be seen in the alimentary canal of some caterpillars[819]; sometimes also they surround part of this organ, like a series of minute rings[820].
iv. Colour. The most usual colour of the muscles of insects is white: those for flight however, according to Chabrier, differ from the rest, by being of a deeper and reddish colour[821]; and I have observed likewise that those in the head of the stag-beetle, when dried at least, are red, and look something like the flesh of warm-blooded animals.
v. Kinds and Denomination. In general, muscles may be regarded as divided into primary and secondary—the primary being the muscles by which the principal movements of any organs are effected, and the secondary their auxiliaries which are the cause of subordinate movements[822]. Every muscle almost has its antagonist, the action of which is in an opposite direction; so that when it is equal, the organ to which they are attached remains without motion; but when that of one preponderates, a movement in proportion takes place[823]. The principal antagonist muscles that may be found in insects are the following. 1. Levator muscles that raise an organ, and Depressors that depress it. 2. Flexors that bend an organ, and Extensors that unbend or extend it. 3. Abductors that draw an organ back, and Adductors that draw it forwards. 4. Constrictors that contract an opening, and Laxators that relax it. 5. Supinators that turn the underside of an organ upwards, and Pronators that return it to its natural situation. Some of these muscles in insects, like some of their articulations and their spinal chord[824], seem to exercise a double function,—thus the levators and depressors of the wings are constrictors and laxators of the trunk[825]. At first it may seem that insects, not having the power of turning up the hand, cannot have the Supinator and Pronator muscles; but some muscle of this kind must be in the Gryllotalpa, and in those that have a versatile head[826].
v. Attachment and Insertion. The attachment and insertion of the muscles in insects in general is to the interior of the crust, or to some of its internal processes as a fulcrum, and to the organ to be moved. In some cases, however, the muscles act upon the organ by the intervention of other bodies. Thus, those that move the wings are often attached to little bones, as Chabrier calls them[827], which are connected with the base of the wings by ligaments. In the Dynastidæ and other Lamellicorns, and the Libellulina, &c., a remarkable provision is made for giving a vast increment of force to the muscles of the wings, by means of caps or cupules surmounted by a tendon, which receive their extremity; the tendon terminating in a fine point attached to the wing, and thus more muscles are brought to bear upon it[828]. Chabrier seems to think that, in some cases, the back that intervenes between each pair of wings is the medium by which the muscles act upon it[829].
vi. Motions. Irritability is the universal distinction of the muscular fibre,—when put in action by the will or involuntarily, it causes it to contract or become shorter; and the intermediate agents of the will and other causes are the nerves, which, as galvanic experiments seem in some degree to prove, are the conductors of an invisible fluid or power which immediately causes that action. If a nerve is divided, the muscles to which it renders obey it no longer, evidently proving that the nerves cause muscular irritability[830]. How this contraction is immediately effected,—whether the fibre, as some suppose, undergoes any crispation, or becomes zigzag[831], or whether there is any sudden change in their chemical composition that rapidly and strongly augments their cohesion, as Cuvier hints[832], cannot be clearly ascertained, unless a Bauer could submit the living fibre to his glasses. All that we know certainly on the subject is, that muscles alternately contract and relax at the bidding of the will or involuntarily, and so occasion all the movements of animal bodies.