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The Physiology of the Alimentary Canal of Tyroglyphus Farinae

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The Physiology of the Alimentary Canal of Tyroglyphus Farinae 45 The Physiology of the Alimentary Canal of Tyroglyphus farinae BY T. E. HUGHES, M.A. (From the Zoology Department, Birkbeck College, University of London) HIS investigation into the activities of the gut in Tyroglyphus farinae Linn. Twas undertaken because so little is known of the physiology of the tyro- glyphid mites. The work was started in 1940 and has been continued as opportunity arose since then. It was hoped that the region of secretion of the digestive enzymes, together with the seat of carbohydrate and protein diges- tion and absorption, might be determined. Since the Malpighian tubes in the Tyroglyphidae (Acaridiae) are reduced or absent, it was hoped to find what region of the gut is responsible for the excretion of nitrogenous waste. An attempt was also made to identify the chief nitrogenous katabolites. METHODS Mites were cultured on food material stained with various indicators; by this means the pH of the gut was determined subject to protein error, and the passage of food was followed. Pig meal and bemax were found to be good culture media for this purpose. The cultures were kept in cells formed by fixing a glass ring on to a slide with gold size; they were covered by another slide held in position by a rubber band. The cells were kept in a dessicator at a relative humidity of 90 per cent. Changes in colour of the indicator could be observed by transparency through the body-wall of the living animal. Cultures were also made on rice starch, potato starch, gluten, and yeast cells to localize carbohydrate and protein digestion. Animals cultured on starch were mounted in a dilute alcoholic solution of iodine. Gluten was used as a protein, because if fed on whole grain the mites attack the germ which is rich in this substance. Fibrin and dried red-blood corpuscles were consistently refused by the animals. Some difficulty was encountered in finding a method of mounting which would render the mites clear without interfering with the colour of the indicator. The method finally adopted was to drop a live mite into glycerine on a slide; on this was placed a coverslip smeared with glycerine jelly. The slide was warmed just sufficiently to melt the jelly; this greatly helped the clearing, and on cooling the weight of the coverslip was generally found to have spread the legs without bursting the body. Mounted in this way, the animal could be examined with an oil-immersion lens. Quarterly Journal Microscopical Science, Vol. 91, part 1, March, 1950. 46 Hughes—The Physiology of Efforts were made and pursued for several months to dissect out the gut, but without success. As the body-wall of the opisthosoma is punctured, the parenchyma loses fluid and the mite collapses. It appears that mites cultured at a relative humidity of 90 per cent, have a positive internal pres- sure in this part of the body; this is indicated by their turgid and glistening appearance. Extracts were made of whole animals collected by standing a beaker of heavily infected material in a half petri dish of water. The mites leaving the culture collected on the water and could be filtered off and washed; after grinding with glass powder, various extracts were made by shaking with the extracting fluid and centrifuging off the solid debris. Serial sections were cut from paraffin, celloidin, and ester-wax blocks, and stained with Mallory, haematoxylin and eosin, mucicarmine and muci- haematein to investigate the histology of the gut. Some sections were cut after fixation with a 1 per cent, solution of osmium tetroxide to determine the distribution of fats. After fixation with Pasted and Leonard's fluid (Lisbon, 1936), sections were made and stained with Best's carmine to find the site of glycogen storage. Bauer's reaction was also successfully employed in this respect, as a more specific histochemical test for carbohydrates. Extracts were made from heavily infected cultures, and specific tests were carried out for various nitrogenous katabolites. Changes in size, opacity, and birefringence of the faecal pellet in its passage through the gut were also investigated. In the following description the term 'dense culture' is employed for cultures containing more than io4 mites per gramme in all stages of the life-cycle excluding eggs; young cultures are those containing less than io3 mites per gramme, and here the mites are hard to find. In the former, the whole medium appears to be in motion when seen under the binocular micro- scope, and ultimately such cultures come to consist of exuvia, faecal pellets, and mites only. ANATOMY AND FUNCTION The anatomy of the gut of the Tyroglyphids has been described by Michael (1901) and its homologies discussed by Lonnfors (1930); a general descrip- tion is also given by Vitzthum (1940). The nomenclature used by these and other authors is by no means uniform, and an attempt to correlate it was thought to be useful. The arachnid gut like that of all Arthropoda is divisible into three regions, a fore-gut lined with chitin, an endodermal mesenteron, and a chitin-lined hind-gut. It is characteristic of the group that the hind-gut—derived from the proctodaeum—is short. The fore-gut becomes differentiated into a buccal cavity and usually a suctorial pharynx of varying complexity, followed by an oesophagus. The mesenteron always develops a system of caeca; and in spiders, pseudoscorpions, pedipalpi, and scorpions, terminates in an enlarge- ment into which the excretory structures discharge, and which connects with the chitinous hind-gut. The Alimentary Canal of Tyroglyphus farinae 47 In the Acari the fore-gut, if a chitinous lining is to be taken as any guide, certainly gives rise to the buccal cavity and pharynx. The oesophagus is only reported as having a lining of chitin in the prostigmatic Trombidiformes (Sig Thor, 1904), the Halacaridae (Thomae, 1925), and the Analgesidae (Lonnfors, 1930). Other authors have reported the oesophagus, in the forms studied by them, as having no chitinous lining and consisting of a thin epithelium. Thus the homology of the oesophagus of the Acari with that of other arachnids, where it is demonstrably part of the fore-gut, is by no means clear. The mesenteron has been ascribed various limits, though the term, when used, has been most frequently restricted to that part which bears the caecal out- growths. The term hind-gut, or Enddarm of German authors, has also been used in a loose sense to cover the regions of the gut which are not lined with chitin, and which properly should be regarded as part of the mesenteron. It is proposed here to limit the fore-gut to the buccal cavity, pharnyx, and oesophagus, and to regard the alimentary canal between the oesophagus and the chitinous region of the rectum as being mesenteron. The term rectum is restricted to that part of the posterior end of the alimentary canal which is lined with chitin; the mesenteron then becomes divisible into an anterior stomach provided with caeca, an intermediate colon, and finally a post-colon. The gut of T. farinae is of the typical tyroglyphid type (Text-fig. 1). The mouth lies below the insertion of the chelicerae; and the buccal cavity, which is bounded above by the very short epistome and the bases of the chelicerae, and below by the fused bases of the pedipalps, leads into a pharynx. The pharyngeal wall is heavily chitinized and crescentic in cross-section with the concavity dorsal. The dorsal wall of the pharynx can be raised by means of five pairs of dilator muscles which have their origins on the internal skeleton of the gnathosoma, which here projects back from the sub-cheliceral shelf into the gnathosoma as a pair of rods. The muscles are inserted on to the roof of the pharynx near its lateral edges. Starting behind the second dilators, four constrictor muscles alternate with them, running transversely from horn to horn of the pharyngeal crescent. Food appears to be seized by the chelicerae and pushed into the buccal cavity through the mouth, the free palpal parts of the pedipalps probably assisting in this. The action of the pharyngeal muscles is to raise the roof of the pharynx by means of the dilator muscles and to allow food to be pushed back into its lumen. By successive contraction of the constrictors from before backwards, the contents of the pharynx, provided they are not too dense or solid, could be squeezed back- wards into and along the narrow oesophagus. Whilst the process is going on, the pharynx is held firm by a mid-ventral attachment to the body-wall, and lateral connective tissue attachments from its edges to the ventrolateral body-wall. Michael (1901) described paired salivary glands in the Tyroglyphidae as opening by a short common duct into the posterior end of the buccal cavity through its roof. He said that the ducts were very difficult to follow. All authors, with the exception of Lonnfors, are agreed that the Acari in general 48 Hughes—The Physiology of possess such salivary glands (Vitzthum, 1940). In T. farinae the ducts of the glands open into the posterior angles of the buccal cavity (Text-fig. 2). In T. farinae the food, whether starch grains from flour or pieces broken off whole grain or coarse meal by the chelicerae, is always relatively dry, and I mm. TEXT-FIG, I . Dorsal view of Tyroglyphus farinae to show the alimentary canal. it has to traverse a long pharynx and longer oesophagus. This penetrates the solid central nervous system, and has walls which are thin and certainly devoid of muscles.
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