Alimentary and Nutritive System.
The alimentary canal occupies the median longitudinal axis of the body, being situated below the dorsal vessel, and above the ventral nervous chain; it extends from the mouth to the opposite extremity of the body. It varies greatly in the different kinds of Insects, but in all its forms it is recognised as consisting essentially of three divisions: anterior, middle, and posterior. The first and last of these divisions are considered to be of quite different morphological nature from the middle part, or true stomach, and to be, as it were, invaginations of the extremities of a closed bag; it is ascertained that in the embryo these invaginations have really blind extremities (see Fig. 82, p. [151]), and only subsequently become connected with the middle part of the canal. There are even some larvae of Insects in which the posterior portion of the canal is not opened till near the close of the larval life; this is the case with many Hymenoptera, and it is probable, though not as frequently stated certain, that the occlusion marks the point of junction of the proctodaeum with the stomach. The anterior and posterior parts of the canal are formed by the ectoderm of the embryo, and in embryological and morphological language are called respectively the stomodaeum and proctodaeum; the true stomach is formed from the endoderm, and the muscular layer of the whole canal from the mesoderm.
Fig. 69.—Digestive system of Xyphidria camelus (after Dufour): a, head capsule; b, salivary glands; c, oesophagus; d, crop; e, proventriculus; f, chyle, or true stomach; g, small intestine; h, large intestine; i, Malpighian tubes; k, termination of body.
The alimentary canal is more complex anatomically than it is morphologically, and various parts are distinguished, viz. the canal and its appendicula; the former consisting of oesophagus, crop, gizzard, true stomach, and an intestine divided into two or more parts. It should be remarked that though it is probable that the morphological distinctions correspond to a great extent with the anatomical lines of demarcation, yet this has not been sufficiently ascertained: the origin of the proctodaeum in Musca is indeed a point of special difficulty, and one on which there is considerable diversity of opinion. In some Hemiptera the division of the canal into three parts is very obscure, so that it would be more correct, as Dufour says, to define it as consisting in these Insects of two main divisions—one anterior to, the other posterior to, the insertion of the Malpighian tubes.
It should be borne in mind that the alimentary canal is very different in different Insects, so that the brief general description we must confine ourselves to will not be found to apply satisfactorily to any one Insect. The oesophagus is the part behind the mouth, and is usually narrow, as it has to pass through the most important nervous centres; extremely variable in length, it dilates behind to form the crop. It may, too, have a dilatation immediately behind the mouth, and in such case a pharynx is considered to exist. The crop is broader than the oesophagus, and must be looked on as a mere dilatation of the latter, as no line of demarcation can be pointed out between the two, and the crop may be totally absent.
In some of the sucking Insects there is a lateral diverticulum, having a stalk of greater or less length, called the sucking-stomach; it is by no means certain that the function this name implies is correctly assigned to the organ.
The gizzard or proventriculus (French, gésier; German, Kaumagen) is a small body interposed in some Insects between the true stomach and the crop or oesophagus. It is frequently remarkable for the development of its chitinous lining into strong toothed or ridged processes that look as if they were well adapted for the comminution of food. The function of the proventriculus in some Insects is obscure; its structure is used by systematists in the classification of ants. The extremity of the proventriculus not infrequently projects into the cavity of the stomach.
The true stomach, or chylific ventricle (Magen or Mitteldarm of the Germans), is present in all the post-embryonic stages of the Insect's life, existing even in the imagines of those who live only for a few hours, and do not use the stomach for any alimentary purpose. It is so variable in shape and capacity that no general description of it can be given. Sometimes it is very elongate, so that it is coiled and like an intestine in shape; it very frequently bears diverticula or pouches, which are placed on the anterior part, and vary greatly in size, sometimes they are only two in number, while in other cases they are so numerous that a portion of the outside of the stomach looks as if it were covered with villi. A division of the stomach into two parts is in some cases very marked, and the posterior portion may, in certain cases, be mistaken for the intestine; but the position of the Malpighian tubes serves as a mark for the distinction of the two structures, the tubes being inserted just at the junction of the stomach with the intestine.
The intestine is very variable in length: the anterior part is the smaller, and is frequently spoken of as the colon; at the extremity of the body the gut becomes much larger, so as to form a rectum. There is occasionally a diverticulum or "caecum" connected with the rectum, and in some Insects stink-glands. In some Hemiptera there is no small intestine, the Malpighian tubes being inserted at the junction of the stomach with the rectum. The total length of the alimentary canal is extremely variable; it is necessarily at least as long as the distance between the mouth and anal orifice, but sometimes it is five or six times as long as this, and some of its parts then form coils in the abdominal cavity.