In capitate hydropolyps, the spherical end-knobs of the short tentacles present an exceptional concentration of sensory functions in one of the evolutionarily oldest nervous systems. The tentacular spheres are the basis of sensation and discrimination of objects and of capturing of prey-objects by the discharge of nematocytes (stinging cells). Recent electrophysiological studies of the spheres revealed combined chemo/mechanosensory functioning of the nematocytes and mechanosensitivity of further types of cells. The present electron microscopical study made use of the small size of the spheres of Coryne tubulosa to characterize all cells of some spheres. Five types of ectodermal cells were found to have sensory structural features and to be separated by or enclosed in supporting cells: 1) nematocytes of the stenotele type; 2) short and 3) long ciliated concentric hair cells, which carry a cilium-stereovilli bundle, similar to the cnidocil apparatus of nematocytes; 4) cells having a recessed cilium-microvilli complex equipped with a thick cell-traversing rootlet (rootlet cells); and 5) cells having a recessed short cilium with no microvilli and only a short rootlet and containing, apically as well as basally, aggregations of dense-core vesicles (vesicle-rich cells). Types 1-4 vary the configuration of a concentric cilium-microvilli complex (variations of a concentric hair bundle) and were demonstrated or inferred to be mechanosensitive. Apical exocytotic activity, which is well known for the nematocytes (discharge of their cnidocyst), is indicated by ultrastructure for the nematocyte-resembling concentric hair cells and for the vesicle-rich cells. The tentacular spheres are considered an early paradigm of a sensory epithelium. Its synaptic structures and extensive connectivity are the subject of a subsequent paper.

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Mechanoelectric transduction and its ultrastuctural basis were studied in the cnidocil apparatus of stenotele nematocytes of marine and freshwater Hydrozoa (Capitata and Hydra) as a paradigm for invertebrate hair cells with concentric hair bundles. The nematocytes respond to selective deflection of their cnidocil with phasic-tonic receptor currents and potentials, similar to vertebrate hair cells but without directional dependence of sensitivity. Ultrastructural studies and the use of monoclonal antibodies allowed correlating the mechanoelectric transduction with structural components of the hair bundle. Two other types of depolarising current and voltage changes in nematocytes are postsynaptic, as concluded from their ionic and pharmacological characteristics. One of these types is induced by mechanical stimulation of distant nematocytes and sensory hair cells. It is graded in amplitude and duration, but different from the presynaptic receptor potential. Adequate chemical stimulation of the stenoteles strongly increases the probability of discharge of their cnidocyst, if the chemical stimulus precedes the mechanical one. Simultaneously, the probability of synaptic signalling induced by mechanical stimulation is increased, reaching nearly 100%. The chemoreception of the phospholipids used could be localized in the shaft of the cnidocil, because of the water-insolubility of the stimulant. This chemical stimulation itself does not cause a receptor potential; its action is classified as a modulatory process. Electron microscopy of serial sections of the tentacular spheres of Coryne revealed synapses that are efferent to nematocytes and hair cells besides neurite-neurite synapses, each containing 3-10 clear and/or dense-core vesicles of 70-150 nm diameter. The only candidates to explain the graded afferent signal transmission of nematocytes and hair cells are regularly occurring cell contacts associated with 1(-4) clear vesicles of 160-1100 nm diameter. Transient fusion and partial depletion of stationary vesicles are discussed as mechanisms to reconcile functional and structural data of many cnidarian synapses.

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Stick insects (Carausius morosus) develop pseudotumors in aging adults. Pseudotumor formation starts at the M2 midgut region where an accumulation of stomatogastric nerve terminals is observed. Pseudotumors arise from dying columnar cells whose basal parts form an "amorphous substance" at the basement membrane whereas the apical parts, including the nucleus, are expelled into the gut lumen. The "amorphous substance" is ensheathed by hemocytes. These nodules, which do not melanize, characterize the phenotype of the pseudotumors. With age, cell death and pseudotumor infestation increases. It is shown that the maintenance of midgut tissue homoeostasis is disturbed and becomes more serious with growing pseudotumor incidence. The increased death rate of differentiated columnar cells is no longer compensated by the proliferation of regenerative cells, i.e., intestinal stem cells, in the midgut nidi. The appearance of "holes" in the intestinal wall is evidently a causative factor of premature death. Extirpation of the hypocerebral ganglion in young adults of the stick insect (before the onset of spontaneous pseudotumor formation) provokes the apoptosis of a large number of columnar cells within 24 h and the formation of pseudotumors that are histologically identical with spontaneous ones. We conclude that the stomatogastric nervous system plays a decisive role in the regulatory mechanism maintaining midgut tissue homeostasis. The possibility of experimentally manipulating the regulatory system provides a valuable tool for the exploration of extrinsic factors involved into the feedback circuitry of tissue homeostasis. The fact that comparable pseudotumors were observed in a number of orthopteromorphan species, where they could also be induced by the interruption of the stomatogastric nervous system, indicates that its role in tissue homoeostasis may be widespread in insects and possibly represent a general principle.