SummaryThere is a resurgence of interest in the Drosophila midgut on account of its potential value in understanding the structure, development and function of digestive organs and related epithelia. The recent identification of regenerative or stem cells in the adult gut of Drosophila has opened up new avenues for understanding development and turnover of cells in insect and mammalian gastrointestinal tracts. Conversely, the physiology of the Drosophila gut is less well understood as it is a difficult epithelial preparation to study under controlled conditions. Recent progress in microperfusion of individual segments of the Drosophila midgut, in both larval and adult forms, has enabled ultrastructural and electrophysiological study and preliminary characterization of cellular transport processes in the epithelium. As larvae are more active feeders, the transport rates are higher than in adults. The larval midgut has at least three segments: an anterior neutral zone, a short and narrow acid-secreting middle segment and a long and wider posterior segment (which is the best studied) that secretes base (probably

Deciphering the genome of the fruitfly, Drosophila melanogaster, has revealed 39 genes coding for putative odorant-binding proteins (OBPs), more than are known at present for any other insect species. Using specific antibodies, the expression mosaic of five such OBPs (OS-E, OS-F, LUSH, PBPRP2, PBPRP5) on the antenna and maxillary palp has been mapped in the electron microscope. It was found that (1) OBP expression does correlate with morphological sensillum types and subtypes, (2) several OBPs may be co-localized in the same sensillum, and (3) OBP localization is not restricted to olfactory sensilla. The expression of PBPRP2 in antennal epidermis sheds some light on the possible evolution of OBPs.

Sensilla lining the inner walls of the sacculus on the third antennal segment of Drosophila melanogaster were studied by light and transmission electron microscopy. The sacculus consists of three chambers: I, II and III. Inside each chamber morphologically distinct groups of sensilla having inflexible sockets were observed. Chamber I contains no-pore sensilla basiconica (np-SB). The lumen of all np-SB are innervated by two neurons, both resembling hygroreceptors. However, a few np-SB contain one additional neuron, presumed to be thermoreceptive. Chamber II houses no-pore sensilla coeloconica (np-SC). All np-SC are innervated by three neurons. The outer dendritic segments of two of these neurons fit tightly to the wall of the lumen and resemble hygroreceptor neurons. A third, more electron-dense sensory neuron, terminates at the base of the sensillum and resembles a thermoreceptor cell. Chamber III of the sacculus is divided into ventral and dorsal compartments, each housing morphologically distinct grooved sensilla (GS). The ventral compartment contains thick GS1, and the dorsal compartment has slender sensilla GS2. Ultrastructurally, both GS1 and GS2 are double-walled sensilla with a longitudinal slit-channel system and are innervated by two neurons. The dendritic outer segment of one of the two neurons innervates the lumen of the GS and branches. On morphological criteria, we infer this neuron to be olfactory. The other sensory neuron is probably thermoreceptive. Thus, the sacculus in Drosophila has sensilla that are predominantly involved in hygroreception, thermoreception, and olfaction. We have traced the sensory projections of the neurons innervating the sacculus sensilla of chamber III using cobaltous lysine or ethanolic cobalt (II) chloride. The fibres project to the antennal lobes, and at least four glomeruli (VM3, DA3 and DL2-3) are projection areas of sensory neurons from these sensilla. Glomerulus DL2 is a common target for the afferent fibres of the surface sensilla coeloconica and GS, whereas the VM3, DA3 and DL3 glomeruli receive sensory fibres only from the GS.

In Drosophila, as in most insects, gustation is mediated by sensory hairs located on the external and internal parts of the proboscis and on the legs and wings. We describe in detail the organization and ultrastructure of the gustatory sensilla on the labellum and legs and the distribution of PBPRP2, a putative odorant-binding protein, in the gustatory organs of Drosophila. The labellum carries two kinds of sensilla: taste bristles and taste pegs. The former have the typical morphology of gustatory sensilla and can be further subdivided into three morphological subtypes, each with a stereotyped distribution and innervation. Taste pegs have a unique morphology and are innervated by two receptor cells: one mechanoreceptor and the other a putative chemoreceptor cell. PBPRP2 is abundantly expressed in all adult gustatory organs on labellum, legs, and wings and in the internal taste organs on the proboscis. In contrast to olfactory organs, where PBPRP2 is expressed in the epidermis, this protein is absent from the epidermis of labial palps and legs. In the taste bristles of the labellum and legs, PBPRP2 is localized in the crescent-shaped lumen of the sensilla, and not in the lumen where the dendrites of the gustatory neurons are found, making a function in stimulus transport unlikely in these sensilla. In contrast, PBPRP2 in peg sensilla is expressed in the inner sensillum-lymph cavity and is in contact with the dendrites. Thus, PBPRP2 could be involved as a carrier for hydrophobic ligands, e.g., bitter tastants, in these sensilla.

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