Part 20

_Abdominal Segments and Appendages._--We have already seen that in numerous lower insects the abdomen is formed from twelve divisions placed in linear fashion. Eleven of these may perhaps be considered as true segments, but the twelfth or terminal one is different, and is called by Heymons a telson; in it is placed the anal orifice, and the mass subsequently becomes the upper and lower laminae anales. In Hemiptera this telson is absent, and the anal orifice is placed quite at the termination of the eleventh segment. Moreover, in this order the abdomen shows at first a division into only nine segments and a terminal mass, which last subsequently becomes divided into two. The appendages of the abdomen are called cerci, stylets and gonapophyses. They differ much according to the kind of insect, and in the adult according to sex. Difference of opinion as to the nature of the abdominal appendages prevails. The cerci, when present, appear in the mature insect to be attached to the tenth segment, but according to Heymons they are really appendages of the eleventh segment, their connexion with the tenth being secondary and the result of considerable changes that take place in the terminal segments. It has been disputed whether any true cerci exist in the higher insects, but they are probably represented in the Diptera and in the scorpion-flies (Mecaptera). In those insects in which a median terminal appendage exists between the two cerci this is considered to be a prolongation of the eleventh tergite. The stylets, when present, are placed on the ninth segment, and in some Thysanura exist also on the eighth segment; their development takes place later in life than that of the cerci. The gonapophyses are the projections near the extremity of the body that surround the sexual orifices, and vary extremely according to the kind of insect. They have chiefly been studied in the female, and form the sting and ovipositor, organs peculiar to this sex. They are developed on the ventral surface of the body and are six in number, one pair arising from the eighth ventral plate and two pairs from the ninth. This has been found to be the case in insects so widely different as Orthoptera and Aculeate Hymenoptera. The genital armature of the male is formed to a considerable extent by modifications of the segments themselves. The development of the armature has been little studied, and the question whether there may be present gonapophyses homologous with those of the female is open.

[Illustration: A. After Wheeler, _Journ. Morph._ vol. viii., and Folsom, _Bull. Mus. Harvard_, xxxvi.

B. After Folsom.

FIG. 18.--Embryos of Springtail (_Anuridamaritima_). Magnified. A, Head-region of germ band. B, Section through head and thorax. The neuromeres are shown in Arabic, the appendages in Roman numerals.

1, Ocular segment. 2, Antennal. 3, Trito-cerebral. 4, Mandibular. 5, Maxillular. 6, Maxillary. 7, Labial. 8, Prothoracic. 9, Mesothoracic. 10, Metathoracic.]

In the adult state no insect possesses more than six legs, and they are always attached to the thorax; in many Thysanura there are, however, processes on the abdomen that, as to their position, are similar to legs. In the embryos of many insects there are projections from the segments of the abdomen similar, to a considerable extent, to the rudimentary thoracic legs. The question whether these projections can be considered an indication of former polypody in insects has been raised. They do not long persist in the embryo, but disappear, and the area each one occupied becomes part of the sternite. In some embryos there is but a single pair of these rudiments (or vestiges) situate on the first abdominal segment, and in some cases they become invaginations of a glandular nature. Whether cerci, stylets and gonapophyses are developed from these rudiments has been much debated. It appears that it is possible to accept cerci and stylets as modifications of the temporary pseudopods, but it is more difficult to believe that this is the case with the gonapophyses, for they apparently commence their development considerably later than cerci and stylets and only after the apparently complete disappearance of the embryonic pseudopods. The fact that there are two pairs of gonapophyses on the ninth abdominal segment would be fatal to the view that they are in any way homologous with legs, were it not that there is some evidence that the division into two pairs is secondary and incomplete. But another and apparently insuperable objection may be raised--that the appendages of the ninth segment are the stylets, and that the gonapophyses cannot therefore be appendicular. The pseudopods that exist on the abdomen of numerous caterpillars may possibly arise from the embryonic pseudopods, but this also is far from being established.

_Nervous System._--The nervous system is ectodermal in origin, and is developed and segmented to a large extent in connexion with the outer part of the body, so that it affords important evidence as to the segmentation thereof. The continuous layer of cells from which the nervous system is developed undergoes a segmentation analogous with that we have described as occurring in the ventral plate; there is thus formed a pair of contiguous ganglia for each segment of the body, but there is no ganglion for the telson. The ganglia become greatly changed in position during the later life, and it is usually said that there are only ten pairs of abdominal ganglia even in the embryo. In Orthoptera, Heymons has demonstrated the existence of eleven pairs, the terminal pair becoming, however, soon united with the tenth. The nervous system of the embryonic head exhibits three ganglionic masses, anterior to the thoracic ganglionic masses; these three masses subsequently amalgamate and form the sub-oesophageal ganglion, which supplies the trophal segments. In front of the three masses that will form the sub-oesophageal ganglion the mass of cells that is to form the nervous system is very large, and projects on each side; this anterior or "brain" mass consists of three lobes (the prot-, deut-, and tritencephalon of Viallanes and others), each of which might be thought to represent a segmental ganglion. But the protocerebrum contains the ganglia of the ocular segment in addition to those of the procephalic lobes. These three divisions subsequently form the supra-oesophageal ganglion or brain proper. There are other ganglia in addition to those of the ventral chain, and Janet supposes that the ganglia of the sympathetic system indicate the existence of three anterior head-segments; the remains of the segments themselves are, in accordance with this view, to be sought in the stomodaeum. Folsom has detected in the embryo of _Anurida_ a pair of ganglia (fig. 18, 5) belonging to the maxillular (or superlingual) segment, thus establishing seven sets of cephalic ganglia, and supporting his view as to the composition of the head.

_Air-tubes._--The air-tubes, like the food-canal, are formed by invaginations of the ectoderm, which arise close to the developing appendages, the rudimentary spiracles appearing soon after the budding limbs. The pits leading from these lengthen into tubes, and undergo repeated branching as development proceeds.

_Dorsal Closure._--The germ band evidently marks the ventral aspect of the developing insect, whose body must be completed by the extension of the embryo so as to enclose the yolk dorsally. The method of this dorsal closure varies in different insects. In the Colorado beetle (_Doryphora_), whose development has been studied by W. M. Wheeler, the amnion is ruptured and turned back from covering the germ band, enclosing the yolk dorsally and becoming finally absorbed, as the ectoderm of the germ band itself spreads to form the dorsal wall. In some midges and in caddis-flies the serosa becomes ruptured and absorbed, while the germ band, still clothed with the amnion, grows around the yolk. In moths and certain saw-flies there is no rupture of the membranes; the Russian zoologists Tichomirov and Kovalevsky have described the growth of both amnion and embryonic ectoderm around the yolk, the embryo being thus completely enclosed until hatching time by both amnion and serosa. V. Graber has described a similar method of dorsal closure in the saw-fly _Hylotoma_.

[Illustration: After Heymons, _Zeit. Wiss. Zoolog._ vol. 53.

FIG. 19.--Cross sections through Abdomen of German Cockroach Embryo. A (later than fig. 16) magnified. B (still more advanced, dorsal closure complete) magnified.

ec, Ectoderm. en, Endoderm. sp, Splanchnic layer of mesoderm. y, Yolk. h, Heart. p, Pericardial septum. c, Coelom. g, Germ-cells surrounded by rudiment-cells of ovarian tubes. m, Muscle-rudiment. n, Nerve-chain. f, Fat body. s, Inpushing of ectoderm to form air-tubes. x, Secondary body-cavity.]

_Mesoderm, Coelom and Blood-System._--From the mesoderm most of the organs of the body--muscular, circulatory, reproductive--take their origin. The mass of cells undergoes segmentation corresponding with the outer segmentation of the embryo, and a pair of cavities--the coelomic pouches (fig. 16, M)--are formed in each segment. Each coelomic pouch--as traced by Heymons in his study on the development of the cockroach (_Phyllodromia_)--divides into three parts, of which the most dorsal contains the primitive germ-cells, the median disappears, and the ventral loses its boundaries as it becomes filled up with the growing fat body (fig. 19). This latter, as well as the heart and the walls of the blood spaces, arises by the modification of mesodermal cells, and the body cavity is formed by the enlargement and coalescence of the blood channels and by the splitting of the fat body. It is therefore a haemocoel, the coelom of the developed insect being represented only by the cavities of the genital glands and their ducts.

_Reproductive Organs._--In the cockroach embryo, before the segmentation of the germ-band has begun, the primitive germ-cells can be recognized at the hinder end of the mesoderm, from whose ordinary cells they can be distinguished by their larger size. At a later stage further germ-cells arise from the epithelium of the coelomic pouches from the second to the seventh abdominal segments, and become surrounded by other mesoderm cells which form the ovarian or testicular tubes and ducts (fig. 19, g). In the male of _Phyllodromia_ the rudiment of a vestigial ovary becomes separated from the developing testis, indicating perhaps an originally hermaphrodite condition. An exceedingly early differentiation of the primitive germ-cells occurs in certain Diptera. E. Metchnikoff observed (1866) in the development of the parthenogenetic eggs produced by the precocious larva of the gall-midge _Cecidomyia_ that a large "polar-cell" appeared at one extremity during the primitive cell-segmentation. This by successive divisions forms a group of four to eight cells, which subsequently pass through the blastoderm, and dividing into two groups become symmetrically arranged and surrounded by the rudiments of the ovarian tubes. E. G. Balbiani and R. Ritter (1890) have since observed a similar early origin for the germ-cells in the midge _Chironomus_ and in the _Aphidae_.

The paired oviducts and vasa deferentia are, as we have seen, mesodermal in origin. The median vagina, spermatheca and ejaculatory duct are, on the other hand, formed by ectodermal inpushings. The classical researches of J. A. Palmen (1884) on these ducts have shown that in may-flies and in female earwigs the paired mesodermal ducts open directly to the exterior, while in male earwigs there is a single mesodermal duct, due either to the coalescence of the two or to the suppression of one. In the absence of the external ectodermal ducts usual in winged insects, these two groups resemble therefore the primitive Aptera. The presence of rudiments of the genital ducts of both sexes in the embryo of either sex is interesting and suggestive. The ejaculatory duct which opens on the ninth abdominal sternum in the adult male arises in the tenth abdominal embryonic segment and subsequently moves forward.

GROWTH AND METAMORPHOSIS

[Illustration: After Marlatt, _Ent. Bull._ 4, n. s. (U.S. Dept. Agr.).

FIG. 20.--a, Bed-bug (_Cimex lectularis_, Linn.); newly hatched young from beneath; b, from above; d, egg, magnified; c, foot with claws; e, serrate spine, more highly magnified.]

[Illustration: From Mally, _Ent. Bull._ 24 (U.S. Dept. Agr.).

FIG. 21.--e, f, Owl moth (_Heliothis armigera_); a, b, egg, highly magnified; c, larva or caterpillar; d, pupa in earthen cell.]

After hatching or birth an insect undergoes a process of growth and change until the adult condition is reached. The varied details of this post-embryonic development furnish some of the most interesting facts and problems to the students of the Hexapoda. Wingless insects, such as spring-tails and lice, make their appearance in the form of miniature adults. Some winged insects--cockroaches, bugs (fig. 20) and earwigs, for example--when young closely resemble their parents, except for the absence of wings. On the other hand, we find in the vast majority of the Hexapoda a very marked difference between the perfect insect (imago) and the young animal when newly hatched and for some time after hatching. From the moth's egg comes a crawling caterpillar (fig. 21, c), from the fly's a legless maggot (fig. 25, a). Such a young insect is a _larva_--a term used by zoologists for young animals generally that are decidedly unlike their parents. It is obvious that the hatching of the young as a larva necessitates 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.

[Illustration: After Howard, _Insect Life_, vol. vii.

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.

[Illustration: Adapted from Koerschelt and Herder and Lowne.

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 quantity of the food taken is often radically different. (2) A wingless, sedentary creature is turned into a winged one with superlative powers of aerial movement. (3) An individual in which the reproductive organs and powers are functionally absent becomes one in which these structures and powers are the only reason for existence, for the great majority of insects die after a brief period of reproduction. These changes are in the higher insects so extreme that it is difficult to imagine how they could be increased. In the case of the common drone-fly, _Eristalis tenax_, the individual, from a sedentary maggot living in filth, without any relations of sex, and with only unimportant organs for the ingestion of its foul nutriment, changes to a creature of extreme alertness, with magnificent powers of flight, living on the products of the flowers it frequents, and endowed with highly complex sexual structures.

[Illustration: After Westwood, _Modern Classification_.

FIG. 24.--Campodeiform Larva of a Ground-Beetle (_Aepus marinus_). Magnified.]

[Illustration: After Howard, _Ent. Bull._ 4, n. s. (_U.S. Dept. Agr._).

FIG. 25.--Vermiform Larva (maggot) of House-fly (_Musca domestica_). Magnified. b, spiracle on prothorax; c, protruded head region; d, tail-end with functional spiracles; e, f, head region with mouth hooks protruded; g, hooks retracted; h, eggs. All magnified.]