The alimentary canal has two coats of muscles: a longitudinal and a transverse or annular. Both coexist in most of its parts. Internal to these coats there exists in the anterior and posterior parts of the canal a chitinous layer, which in the stomach is replaced by a remarkable epithelium, the cells of which are renewed, new ones growing while the old are still in activity. We figure a portion of this structure after Miall and Denny, and may remark that Oudemans[[42]] has verified the correctness of their representation. The layers below represent the longitudinal and transverse muscles.
Fig. 70.—Epithelium of stomach of Cockroach (after Miall and Denny): the lower parts indicate the transverse and longitudinal muscular layers.
In addition to the various diverticula we have mentioned, there are two important sets of organs connected with the alimentary canal, viz. the salivary glands and the Malpighian tubes.
The salivary glands are present in many Insects, but are absent in others. They are situate in the anterior portion of the body, and are very variable in their development, being sometimes very extensive, in other cases inconspicuous. They consist either of simple tubes lined with cells, or of branched tubes, or of tubes dilated laterally into little acini or groups of bags, the arrangement then somewhat resembling that of a bunch of grapes. There are sometimes large sacs or reservoirs connected with the efferent tubes proceeding from the secreting portions of the glands. The salivary glands ultimately discharge into the mouth, so that the fluid secreted by them has to be swallowed in the same manner as the food, not improbably along with it. The silk so copiously produced by some larvae comes from very long tubes similar in form and situation to the simple tubes of the salivary glands.
The Malpighian tubules are present in most Insects, though they are considered on good authority to be absent in many Collembola and in some Thysanura. They are placed near the posterior part of the body, usually opening into the alimentary canal just at the junction of the stomach and the intestine, at a spot called the pylorus. They vary excessively in length and in number,[[43]] being sometimes only two, while in other cases there may be a hundred or even more of them. In some cases they are budded off from the hind-gut of the embryo when this is still very small; in other cases they appear later; frequently their number is greater in the adult than it is in the young. In Gryllotalpa there is one tube or duct with a considerable number of finer tubes at the end of it. There is no muscular layer in the Malpighian tubes, they being lined with cells which leave a free canal in the centre. The tubes are now thought, on considerable evidence, to be organs for the excretion of uric acid or urates, but it is not known how they are emptied. Marchal has stated[[44]] that he has seen the Malpighian tubes, on extraction from the body, undergo worm-like movements; he suggests that their contents may be expelled by similar movements when they are in the body.
The functions of the different portions of the alimentary canal, and the extent to which the ingested food is acted on by their mechanical structures or their products is very obscure, and different opinions prevail on important points. It would appear that the saliva exercises a preparatory action on the food, and that the absorption of the nutritive matter into the body cavity takes place chiefly from the true stomach, while the Malpighian tubes perform an excretory function. Beyond these elementary, though but vaguely ascertained facts, little is known, though Plateau's[[45]] and Jousset's researches on the digestion of Insects throw some light on the subject.
Respiratory Organs.
The respiration of Insects is carried on by means of a system of vessels for the conveyance of air to all parts of the body; this system is most remarkably developed and elaborate, and contrasts strongly with the mechanism for the circulation of the blood, which is as much reduced as the air system is highly developed, as well as with the arrangement that exists in the Vertebrates. There are in Insects no lungs, but air is carried to every part of the body directly by means of tracheae. These tracheae connect with the spiracles—the orifices at the sides of the body we have already mentioned when describing the external structures—and the air thus finds its way into the most remote recesses of the Insect's body. The tracheae are all intimately connected. Large tubes connect the spiracles longitudinally, others pass from side to side of the body, and a set of tracheae for the lower part of the body is connected with another set on the upper surface by means of several descending tubes. From these main channels smaller branches extend in all directions, forking and giving off twigs, so that all the organs inside the body can be supplied with air in the most liberal manner. On opening a freshly deceased Insect the abundance of the tracheae is one of the peculiarities that most attracts the attention; and as these tubes have a peculiar white glistening appearance, they are recognised without difficulty. In Insects of active flight, possibly in some that are more passive, though never in larvae, there are air-sacs, of more than one kind, connected with the tracheae, and these are sufficiently capacious to have a considerable effect in diminishing the specific gravity of the Insect. The most usual situation for these sacs is the basal portion of the abdominal cavity, on the great lateral tracheal conduits. In speaking of the external structure we have remarked that the stigmata, or spiracles, by which the air is admitted are very various in their size and in the manner in which they open and close. Some spiracles have no power of opening; while others are provided with a muscular and valvular apparatus for the purpose of opening and closing effectually.
The structure of the tracheae is remarkable: they are elastic and consist of an outer cellular, and an inner chitinous layer; this latter is strengthened by a peculiar spiral fibre, which gives to the tubes, when examined with the microscope, a transversely, closely striated appearance. Packard considers[[46]] that in some tracheae this fibre is not really spiral, but consists of a large number of closely placed rings. Such a condition has not, however, been recorded by any other observer. The spiral fibre is absent in the fine capillary twigs of the tracheal system, as well as from the expanded sacs. The mode of termination of the capillary branches is not clear. Some have supposed that the finest twigs anastomose with others; on the other hand it has been said that they terminate by penetrating cells, or that they simply come to an end with either open or closed extremities. Wistinghausen[[47]] states that in the silk-glands the tracheal twigs anastomose, and he is of opinion that the fine terminal portions contain fluid. However this may be, it is certain that all the organs are abundantly supplied with a capillary tracheal network, or arboreal ramification, and that in some cases the tubes enter the substance of tissues. Near their terminations they are said to be 1⁄30 to 1⁄60 millimetre in diameter.