Digestion in the Tsetse-Fly: A Study of Structure and Function
Authors: Wigglesworth V. B. 
Published: June 2009

The anatomy, histology and digestive enzymes of the mid-intestine of the tsetse-fly have been investigated, and an attempt has been made to determine the functions of the various parts and to observe the changes to which they are subject during the digestion of blood.Histologically the mid-gut of Glossina consists of three regions:(i) An anterior segment of small, pale-staining, irregularly columnar cells, which comprises about half the total length of the mid-gut. The zone of giantcells containing bacteroids, which is very limited in extent, lies at about the middle of this region.(ii) A middle segment of large, deeply staining cells, heaped together in the resting state, which is separated abruptly from the anterior segment.(iii) A posterior segment, arising by gradual transition from the middle segment, composed of regular columnar cells.After a meal the blood is concentrated by the removal of fluid in the anterior segment but it shows no other change in this region. The giant-cells are greatly flattened but they do not regularly discharge the bacteroids which they contain and there is no evidence that these organisms play any part in the digestion of blood. Their possible function has been discussed.During digestion the cells in the middle segment contain globules of secretion, and vacuolated buds of cytoplasm are set free and disintegrate in the lumen. The blood shows an abrupt change on reaching this region; it turns black where it is in contact with the epithelium and amorphous masses of altered blood pigment are deposited.In the posterior segment, the epithelial cells become greatly vacuolated later in digestion and are probably concerned chiefly in absorption.The distribution of digestive enzymes agrees with these histological observations. The salivary glands and proventriculus contain no digestive enzymes, and the anterior and posterior segments of the mid-gut also are practically inactive. But the middle segment produces a very active tryptase which agrees in its pH-activity curve and other properties with the tryptase of the cockroach. A peptidase also is present but, except for a very weak amylase, enzymes acting upon carbohydrates are absent. The contents of the mid-gut are always slightly acid (about pH 6·5) and the tryptase present is well adapted to work at this reaction.These findings have been contrasted with those in a non-blood-sucking muscid (Calliphora). Here the salivary glands secrete an active amylase and the mid-gut is rich in amylase, invertase and maltase, whereas the proteolytic enzymes are extremely weak.Some observations have been made upon the tracheal supply to the walls of the gut. The epithelial cells of the middle segment have been shown to contain a very rich supply of intracellular tracheoles. These are usually difficult to make out in the resting cells but after a large meal the surface of the cells is ruptured and blood pigment enters the tracheoles and may extend to the sub-epithelial tracheoles and tracheae or even to quite large tracheal trunks. As the epithelial cells are flattened by the pressure of the meal, this pigment is set free in the lumen in the form of dark rods of haematin, which often bear a superficial resemblance to bacteria. The pigment in the deeper tubes appears to be slowly absorbed later. Intracellular tracheoles similar to these are present also in the mid-gut of Calliphora.The proventriculus in Glossina is a complex and has always been a puzzling structure. It has been shown that it acts as a sphincter between the fore-gut and mid-gut and that it is responsible for the production of the peritrophic membrane. This membrane, which is composed of chitin but contains a small quantity of protein, is secreted in the form of a fluid by the ring of large epithelial cells at the base of the proventriculus. The fluid is pressed and condensed to form a uniform membrane by being drawn through the cleft between the wall of the proventriculus and the funnel-shaped invagination of the fore-gut.The function of the peritrophic membrane has been discussed and it has been shown that it is freely permeable to digestive enzymes and to haemoglobin.