a more or less profound transformation or metamorphosis before the perfect state is attained. Usually this transformation comes with apparent suddenness, at the penultimate stage of the insect’s life-history, when the passive pupa (fig. 21, d) is revealed, exhibiting the wings and other imaginal structures, which have been developed unseen beneath the cuticle of the larva. Hexapoda with this resting pupal stage in their life-history are said to undergo “a complete transformation,” to be metabolic, or holometabolic, whereas those insects in which the young form resembles the parent are said to be ametabolic. Such insects as dragon-flies and may-flies, whose young, though unlike the parent, develop into the adult form without a resting pupal stage are said to undergo an “incomplete transformation” or to be hemimetabolic. The absence of the pupal stage depends upon the fact that in the ametabolic and hemimetabolic Hexapoda the wing-rudiments appear as lateral outgrowths (fig. 22) of the two hinder thoracic segments and are visible externally throughout the life-history, becoming larger after each moult or casting of the cuticle. Hence, as has been pointed out by D. Sharp (1898), the marked divergence among the Hexapoda, as regards life-history, is between insects whose wings develop outside the cuticle (Exopterygota) and those whose wings develop inside the cuticle (Endopterygota), becoming visible only when the casting of the last larval cuticle reveals the pupa. Metamorphosis among the Hexapoda depends upon the universal acquisition of wings during post-embryonic development—no insect being hatched with the smallest external rudiments of those organs—and on the necessity for successive castings or “moults” (ecdyses) of the cuticle.
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| After Howard, Insect Life, vol. vii. |
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Fig. 22.—Nymph of Locust (Schistocera americana), showing wing-rudiments. |
Ecdysis.—The embryonic ectoderm of an insect consists of a layer of cells forming a continuous structure, the orifices in it—mouth, spiracles, anus and terminal portions of the genital ducts—being invaginations of the outer wall. This cellular layer is called the hypodermis; it is protected externally by a cuticle, a layer of matter it itself excretes, or in the excretion of which it plays, at any rate, an important part. The cuticle is a dead substance, and is composed in large part of chitin. The cuticle contrasts strongly in its nature with the hypodermis it protects. It is different in its details in different insects and in different stages of the life of the same insect. The “sclerites” that make up the skeleton of the insect (which skeleton, it should be remembered, is entirely external) are composed of this chitinous excretion. The growth of an insect is usually rapid, and as the cuticle does not share therein, it is from time to time cast off by moulting or ecdysis. Before a moult actually occurs the cuticle becomes separated from its connexion with the underlying hypodermis. Concomitant with this separation there is commencement of the formation of a new cuticle within the old one, so that when the latter is cast off the insect appears with a partly completed new cuticle. The new instar—or temporary form—is often very different from the old one, and this is the essential fact of metamorphosis. Metamorphosis is, from this point of view, the sum of the changes that take place under the cuticle of an insect between the ecdyses, which changes only become externally displayed when the cuticle is cast off. The hypodermis is the immediate agent in effecting the external changes.
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Adapted from Koerschelt and Herder, and Lowne. |
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Fig. 23.—Diagram showing position of imaginal buds in larva of fly. I., II., III., the three thoracic segments of the larva; 1, 2, 3, buds of the legs of the imago; h, bud of head-lobes; f, of feeler; e of eye; b, brain. |
The study of the physiology of ecdysis in its simpler forms has unfortunately been somewhat neglected, investigators having directed their attention chiefly to the cases that are most striking, such as the transformation of a maggot into a fly, or of a caterpillar into a butterfly. The changes have been found to be made up of two sets of processes: histolysis, by which the whole or part of a structure disappears: and histogenesis, or the formation of the new structure. By histolysis certain parts of the hypodermis are destroyed, while other portions of it develop into the new structures. The hypodermis is composed of parts of two different kinds, viz. (1) the larger part of the hypodermis that exists in the maggot or caterpillar and is dissolved at the metamorphosis; (2) parts that remain comparatively quiescent previously, and that grow and develop when the other parts degenerate. These centres of renovation are called imaginal disks or folds. The adult caterpillar may be described as a creature the hypodermis of which is studded with buds that expand and form the butterfly, while the parts around them degenerate. In some insects (e.g. the maggots of the blowfly, Calliphora vomitoria) the imaginal disks are to all appearance completely separated from the hypodermis, with which they are, however, really organically connected by strings or pedicels. This connexion was not at first recognized and the true nature of imaginal disks was not at first perceived, even by Weismann, to whom their discovery in Diptera is due. In other insects the imaginal disks are less completely disconnected from the superficies of the larval hypodermis, and may indeed be merely patches thereof. The number of imaginal disks in an individual is large, upwards of sixty having been discovered to take part in the formation of the outer body of a fly. With regard to the internal organs, we need only say that transformation occurs in an essentially similar manner, by means of a development from centres distributed in the various organs. The imaginal disks for the outer wall of the body, some of them, at any rate, include mesodermal rudiments (from which the muscles are developed) as well as hypodermis. The imaginal disks make their appearance (that is, have been first detected) at very different epochs in the life; their absolute origin has been but little investigated. Pratt has traced them in the sheep-tick (Melophagus) to an early stage of the embryonic life.
Histolysis and Histogenesis.—The process of destruction of the larval tissues was first studied in the forms where metamorphosis is greatest and most abrupt, viz. in the Muscid Diptera. It was found that the tissues were attacked by phagocytic cells that became enlarged and carried away fragments of the tissue; the cells were subsequently identified as leucocytes or blood-cells. Hence the opinion arose that histolysis is a process of phagocytosis. It has, however, since been found that in other kinds of insects the tissues degenerate and break down without the intervention of phagocytes. It has, moreover, been noticed that even in cases where phagocytosis exists a greater or less extent of degeneration of the tissue may be observed before phagocytosis occurs. This process can therefore only be looked on as a secondary one that hastens and perfects the destruction necessary to permit of the accompanying histogenesis. This view is confirmed by the fate of the phagocytic cells. These do not take a direct part in the formation of the new tissue, but it is believed merely yield their surplus acquisitions, becoming ordinary blood-cells or disappearing altogether. As to the nature of histogenesis, nothing more can be said than that it appears to be a phenomenon similar to embryonic growth, though limited to certain spots. Hence we are inclined to look on the imaginal disks as cellular areas that possess in a latent condition the powers of growth and development that exist in the embryo, powers that only become evident in certain special conditions of the organism. What the more essential of these conditions may be is a question on which very little light has been thrown, though it has been widely discussed.
Much consideration has been given to the nature of metamorphosis in insects, to its value to the creatures and to the mode of its origin. Insect metamorphosis may be briefly described as phenomena of development characterized by abrupt changes of appearance and of structure, occurring during the period subsequent to embryonic development and antecedent to the reproductive state. It is, in short, a peculiar mode of growth and adolescence. The differences in appearance between the caterpillar and the butterfly, striking as they are to the eye, do not sufficiently represent the phenomena of metamorphosis to the intelligence. The changes that take place involve a revolution in the being, and may be summarized under three headings: (1) The food-relations of the individual are profoundly changed, an entirely different set of mouth-organs appears and the kind and
