CHAPTER V. Transformations,--Outward and Inward

TRANSFORMATIONS,--OUTWARD AND INWARD

We are now in a position to study in some detail the transformation of those insects whose life-story corresponds more or less closely with that of the butterfly, sketched in the opening pages of this little book. In the case of some of the insects reviewed in the last three chapters, the may-flies and cicads for example, a marked difference between the larva and the imago has been noticed; in others, as the coccids, we find a resting instar before the winged condition is assumed, suggesting the pupal stage in the butterfly's life-story.

The various insect orders whose members exhibit no marked divergence between larva and imago (the Orthoptera for example) are often said to undergo no transformation, to be 'Ametabola.' Those with life-stories such as the dragon-flies' are said to undergo partial transformation, and are termed 'Hemimetabola.' Moths, caddis-flies, beetles, two-winged flies, saw-flies, ants, wasps, bees, and the great majority of insects, having the same type of life-story as the butterfly, are said to undergo complete transformation and are classed as 'Metabola' or 'Holometabola.' Wherein lies the fundamental difference between these Holometabola on the one hand and the Hemimetabola and Ametabola on the other? It is not that the larva differs from the imago or that there is a passive stage in the life-history; these conditions are observable among insects with a 'partial' transformation as we have seen, though the resting instar that simulates the butterfly pupa is certainly exceptional. It has been pointed out by Sharp (1899) that the most important indication of the difference between the two modes of development is furnished by the position of the wing-rudiments. In all Ametabola and Hemimetabola these are visible externally long before the penultimate instar has been reached; in the Holometabola they are not seen until the pupal stage.

Attention has already been drawn to the contrast in outward form between a butterfly and its caterpillar. As in the case of dragon-fly or may-fly, the larval period is essentially a time for feeding and growth, and during this period the larval cuticle is cast four or five, in some species even seven or eight times. After each moult some changes in detail may be observable, for example in the proportions of the body-segments or their outgrowths, in the colour or the closeness of the hairy or spiny armature. But in all main features the caterpillar retains throughout its life the characteristic form in which it left the egg. From the tiny, newly-hatched larva to the full-fed caterpillar, possibly several inches in length, there is all along the same crawling, somewhat worm-like body, destitute of any outward trace of wings. When however the last larval cuticle has split open lengthwise along the back, and has been worked off by vigorous wriggling motions of the insect, the pupa thus revealed shows the wing-rudiments conspicuous at the sides of the body, and lying neatly alongside these are to be seen the forms of feelers, legs, and maxillae of the imago prefigured in the cuticle of the pupa (fig. 1 e). The pupa thus resembles the imago much more closely than it resembles the larva; even in the proportions of the body a relative shortening is to be noticed, and the imago of any insect with complete transformation is reduced in length as compared with the full-fed larva. Now these wings and other structures characteristic of the imago, appear in the pupa which is revealed by the shedding of the last larval cuticle. From these facts we infer that the wing-rudiments must be present in the larva, hidden beneath the cuticle; and until the last larval instar, not beneath the cuticle only, but growing in such-wise that they are hidden by the epidermis. For if they were growing outwardly the new cuticle would be formed over them, so that they would be apparent after the next moult. But it is clear that only in the pupa, forming beneath the cuticle of the last larval instar, can they grow outwards.

Anatomical study of the caterpillar at various stages verifies the conclusions just drawn from superficial observation. A hundred and fifty years ago P. Lyonet in his monumental work (1762) on the caterpillar of the Goat Moth (Cossus) detected, in the second and third thoracic segments, four little white masses buried in the fat-body, and, while doubtful as to their real meaning, he suggested that their number and position might well give rise to the suspicion that they were rudiments of the wings of the moth. But it was a century later that A. Weismann in his classical studies (1864) on the development of common flies, showed the presence in the maggot of definite rudiments of wings, and other organs of the adult--rudiments to which he gave the name of imaginal discs. We will recur later to these transformations of the Diptera. For the present, we pursue our survey of changes in the life-history of the Lepidoptera and can take to guide us the excellent researches of J. Gonin (1894).

Careful study of the imaginal discs of the wings in a caterpillar (fig. 10) made by examining microscopically sections cut through them, shows that the epidermis is pushed in to form a little pouch (C, p) and that into this grows the actual wing-rudiment. Consequently the whitish disk which seems to lie within the body-wall of the larva, is really a double fold of the epidermis, the outer fold forming the pouch, the inner the actual wing-bud. Into the cavity of the latter pass branches from the air-tube system. In its earliest stage, the wing-bud is simply an ingrowing mass of cells (fig. 10 A) which subsequently becomes an inpushed pouch (B). Until the last stage of larval life the wing-bud remains hidden in its pouch, and no cuticle is formed over it. When the pupal stage draws near the bud grows out of its sheath, and projecting from the general surface of the epidermis becomes covered with cuticle to be revealed, as we have seen, after the last larval moult, as the pupal wing. Thus all through the life of the humble, crawling caterpillar, 'it doth not yet appear what it shall be,' but there are being prepared, hidden and unseen, the wondrous organs of flight, which in due time will equip the insect for the glorious aerial existence that awaits it.

As mentioned above, this hidden growth of the wing-rudiments, in butterflies, beetles, flies, bees, and the great majority of the winged insects, has been emphasised by Sharp (1899) as a character contrasting markedly with the outward and visible growth of the wing-rudiments in such insects as cockroaches, bugs, and dragon-flies. The divergence between the two modes of development is certainly very striking, and a conceivable method of transition from the one to the other is not easy to explain. Sharp has expressed the divergence by the terms Endopterygota, applied to all the orders of insects with hidden wing-rudiments (the 'Metabola' or 'Holometabola' of most classifications) and Exopterygota, including all those insects whose wing-rudiments are visible throughout growth ('Hemimetabola' and 'Ametabola'). Those curious lowly insects, belonging to the two orders of the Collembola and Thysanura, none of whose members ever develop wings at all, form a third sub-class, the Apterygota (see Classificatory Table, p. 122).

Not the wings only, but other structures of the imago, varying in extent in different orders, are formed from the imaginal discs. For example, de Reaumur and G. Newport (1839) found that if the thoracic leg of a late-stage caterpillar were cut off, the corresponding leg of the resulting butterfly would still be developed, although in a truncated condition. Gonin has shown that in the Cabbage White butterfly (Pieris brassicae) the legs of the imago are represented, through the greater part of larval life, only by small groups of cells situated within the bases of the larval legs. After the third moult these imaginal discs grow rapidly and the proximal portion of each, destined to develop into the thigh and shin of the butterfly's leg, sinks into a depression at the side of the thorax, while the tip of the shin and the five-segmented foot project into the cavity of the larval leg. Hence we understand that the amputation of the latter by the old naturalists truncated only and did not destroy the imaginal limb. In the blow-fly maggot, Weismann, B.T. Lowne (1890) and J. Van Rees (1888) have shown that the imaginal discs of the legs (fig. 11--1, 2, 3) grow out from deep dermal inpushings. Simple at first, these outgrowths by partial splitting, become differentiated into thigh and shin.

Similarly the feelers and jaws of the butterfly are developed from imaginal discs, and this fact explains how it comes to pass that they differ so widely from the corresponding structures in the caterpillar. The larval feelers (fig. 3 At) are short and stumpy, those of the butterfly long and many-jointed. The maxilla of the larva (fig. 3 Mx) consists of a base carrying two short jointed processes; in the butterfly a certain portion of the maxilla, the hood or galea, is modified into a long, flexible grooved process, capable of forming with its fellow the trunk through which the insect sucks its liquid food (fig. 2). Nothing but some such provision as that of the imaginal discs could render possible the wonderful replacement of the caterpillar's jaws, biting solid food, into those of the butterfly sipping nectar from flowers.

A curious segmental displacement of the imaginal discs with regard to the larva is noticeable in some Diptera. In the larva of the harlequin-midge (Chironomus) as described by Miall and Hammond (1900) the brain is situated in the thorax, and the imaginal discs for the head, eyes, and feelers of the adult lie in close association with it, though they arise from inpushings of the larval head. These rudiments do not appear until the last larval stage has been reached. In the gnats Culex and Corethra, on the other hand, the imaginal discs for the head-appendages retain their normal position within the larval head, and appear in an early stage of larval life. Among the flies of the bluebottle group (Muscidae) the brain (fig. 11 B) is situated, as in Chironomus, in the thoracic region of the legless maggot, which is the larva of an insect of this family, and the imaginal discs for eyes and feelers (fig. 11 e, f) lie just in front of it. Here, the imaginal buds of the legs (fig. 11--1, 2, 3) and wings (fig. 11 W, w) are deeply inpushed, retaining their connection with the skin only by means of a thread of cells. As the larva is legless and headless its outer form is not affected by the discs and it is not surprising to learn that they appear early. It has indeed been suggested that the pharyngeal region of the larva, in connection with which the imaginal head-discs are developed, should be regarded, though it lies in the thorax, as an inpushed anterior section of the larval head. In any case this region is pushed out during the formation of the pupa within the final larval cuticle, so that the imaginal head with its contained brain, its compound eyes, and its complex feelers, takes its rightful place at the front end of the insect.

The mention of the brain suggests a few brief remarks on the changes in the internal organs during insect transformation. There are no imaginal discs for the nervous system; the brain, nerve-cords and ganglia of the butterfly or bluebottle are the direct outcome of those of the caterpillar or maggot. More than seventy years ago, Newport (1839) traced the rapid but continuous changes, which, during the early pupal period, convert the elongate nerve-cord of the caterpillar with its relatively far-separated ganglia into the shortened, condensed nerve-cord of the Tortoise-shell butterfly (Vanessa urticae) with several of the ganglia coalesced. In many Diptera, on the other hand, the nervous system of the larva is more concentrated than that of the imago.

The tubular heart also of a winged insect is the directly modified survival of the larval heart.

Similarly the reproductive organs undergo a gradual, continuous development throughout an insect's life-story. Their rudiments appear in the embryo, often at a very early stage; they are recognisable in the larva, and the matured structures in the imago are the result of their slow process of growth, the details of which must be reckoned beyond the scope of this book. For a full summary of the subject the reader is referred to L.F. Henneguy's work (1904) containing references to much important modern literature, which cannot be mentioned here.

On the other hand, the digestive system of insects that undergo a metamorphosis, passes through a profound crisis of dissolution and rebuilding. This is not surprising when we remember that there is often a great difference between larva and imago in the nature of the food. The digestive canal of a caterpillar runs a fairly straight course through the body and consists of a gullet, stomach (mid-gut), intestine, and rectum; it is adapted for the digestion of solid food. In the butterfly there is one outgrowth of the gullet in the head--a pharyngeal sac adapted for sucking liquids; and another outgrowth at the hinder end of the gullet (which is much longer than in the larva)--a crop or food-reservoir lying in the abdomen. The intestine of the butterfly also is longer than that of the larva, being coiled or twisted. Towards the end of the last larval stage, the cells of the inner coat (epithelium) lining the stomach begin to undergo degeneration, small replacing cells appearing between their bases and later giving rise to the more delicate epithelium that lines the mid-gut of the imago. The larval cells are shed into the cavity of the stomach and become completely broken down. J. Anglas (1902), describing these microscopic changes in the transformations of wasps and bees, has shown that the tiny replacing cells can be recognised in sections through the digestive canal of a very young larva; they may be regarded as representing imaginal buds of the adult gastric epithelium. In the transformations of two-winged flies of the bluebottle group, A. Kowalevsky (1887) has shown that these replacing cells are aggregated in little masses scattered at different points along the stomach and thus corresponding rather closely to the imaginal discs of the legs and wings.

The gullet, crop, and gizzard of an insect, which lie in front of the stomach, are lined by cells derived from the outer skin (ectoderm) which is pushed in to form what is called the 'fore-gut.' Similarly the intestine and rectum, behind the stomach, are lined with ectodermal cells which arise from the inpushed 'hind-gut.' The larval fore- and hind-guts are broken down at the end of larval life and their lining is replaced by fresh tissue derived from two imaginal bands which surround the cavity of the digestive tube, one at the hinder end of the fore-gut, and the other at the front end of the hind-gut. The larval salivary glands in connection with the gullet are also broken down, and fresh glands are formed for the imago.

A large part of the substance of an insect larva consists of muscular tissue, surrounding the digestive tube, and forming the great muscles that move the various parts of the body, and of fat, surrounding the organs and serving as a store of food-material. Very many of the muscle-fibres and the fat-cells also become disintegrated during the late larval and pupal stages, and the corresponding tissues of the adult are new formations derived from special groups of imaginal cells, though some muscles may persist from the larva to the adult. Similarly the complex air-tube or tracheal system of the larva is broken down and a fresh set of tubes is developed, adapted to the altered body-form of pupa and imago.

The destruction of larval tissue and the development of replacing organs from special groups of cells, derived of course from the embryo, and carrying on the continuity of cell-lineage to the adult, are among the most remarkable facts connected with the life-story of insects. The process of tissue-destruction is known as 'histolysis'; the rebuilding process is called 'histogenesis.' Considerable difference of opinion has existed as to factors causing histolysis, and for a summary of the conflicting or complementary theories, the reader is referred to the work of L.F. Henneguy (1904, pp. 677-684). In the histolysis of the two-winged flies, wandering amoeboid cells--like the white corpuscles or leucocytes of vertebrate blood--have been observed destroying the larval tissues that need to be broken down, as they destroy invading micro-organisms in the body. But students of the internal changes that accompany transformation in insects of other orders have often been unable to observe such devouring activity of these 'phagocytes,' and attribute the dissolution of the larval tissues to internal chemical changes. The fact that in all insect transformation a part, and in many a large part, of the larval organs pass over to the pupa and imago, suggests that only those structures whose work is done are broken down through the action of internally formed destructive substances, and one function of the phagocytes is to act as scavengers by devouring what has become effete and useless.