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A receptor scaffold mediates stimulus-response coupling in bacterial chemotaxis.

The mechanism of stimulus-response coupling in bacterial chemotaxis has emerged as a paradigm for understanding general features of intracellular signal transduction both in bacterial and eukaryotic cells. Until recently it was thought that the mechanism involved reversible stochastic interactions between dimeric receptors freely diffusing in the cytoplasmic membrane and several soluble signal transduction proteins within the cytoplasm. Recent results have shown that this view is an oversimplification. The receptors and most of the signal transduction proteins are organized together in a higher ordered structure at one pole of the bacterial cell. The scaffolding network within this structure appears to be composed of C-terminal alpha-helical extensions of the membrane chemoreceptor proteins held together in a lattice by tandem SH3-like domains. Results suggest that stimuli are detected through the perturbations they induce in scaffolding architecture.

Bacterial Outer Membrane Proteins↗

Convergent carotid sinus nerve and superior laryngeal nerve afferent inputs to neurons in the NTS.

A population of 43 neurons in the nucleus of the solitary tract (NTS) was identified in pentobarbital sodium anesthetized, paralyzed, and artificially ventilated cats that received convergent inputs from carotid sinus nerve (CSN) and superior laryngeal nerve (SLN) afferents. In 21 neurons, electrical stimulation of the CSN and SLN each evoked an excitatory postsynaptic potential (EPSP; mean onset latency +/- SE of CSN-evoked input = 7.2 +/- 0.8 ms, range 2.1-14.1 ms; of SLN-evoked input = 10.3 +/- 2.1 ms, range 2.8-46.8 ms). In 22 neurons, electrical stimulation of either the CSN or SLN evoked an EPSP/ inhibitory postsynaptic potential (IPSP) sequence (CSN-evoked input = 6.7 +/- 0.6 ms, range 2.1-12.2 ms; SLN-evoked input = 8.4 +/- 0.8 ms, range 3.0-19.4 ms). Spatial interactions (facilitation, summation, occlusion) and time-dependent inhibitory interactions were observed between the convergent inputs. Natural stimulation of specific receptors indicated that 14 cells received a convergent excitatory input from carotid sinus baroreceptors and laryngeal mechanoreceptors, 10 received a convergent excitatory input from carotid body chemoreceptors and laryngeal mechanoreceptors, and 5 received a convergent excitatory input from baroreceptors, chemoreceptors, and laryngeal mechanoreceptors. The interactions and various patterns of convergence suggest a significant integration of convergent inputs from disparate afferent sources by these neurons.

Afferent Pathways↗

An important functional role of persistent Na+ current in carotid body hypoxia transduction.

Systemic hypoxia in mammals is sensed and transduced by the carotid body into increased action potential (AP) frequency on the sinus nerve, resulting in increased ventilation. The mechanism of hypoxia transduction is not resolved, but previous work suggested that fast Na(+) channels play an important role in determining the rate and timing of APs (Donnelly, DF, Panisello JM, and Boggs D. J Physiol. 511: 301-311, 1998). We speculated that Na(+) channel activity between APs, termed persistent Na(+) current (I(NaP)), is responsible for AP generation that and riluzole and phenytoin, which inhibit this current, would impair organ function. Using whole cell patch clamp recording of intact petrosal neurons with projections to the carotid body, we demonstrated that I(NaP) is present in chemoreceptor afferent neurons and is inhibited by riluzole. Furthermore, discharge frequencies of single-unit, chemoreceptor activity, in vitro, during normoxia (Po(2) 150 Torr) and during acute hypoxia (Po(2) 90 Torr) were significantly reduced by riluzole concentrations at or above 5 microM, and by phenytoin at 100 microM, without significant affect on nerve conduction time, AP magnitude (inferred from extracellular field), and AP duration. The effect of both drugs appeared solely postsynaptic because hypoxia-induced catecholamine release in the carotid body was not altered by either drug. The respiratory response of unanesthetized, unrestrained 2-wk-old rats to acute hypoxia (12% inspired O(2) fraction), which was measured with whole body plethysmography, was significantly reduced after treatment with riluzole (2 mg/kg ip) and phenytoin (20 mg/kg ip). We conclude that I(NaP) is present in chemoreceptor afferent neurons and serves an important role in peripheral chemoreceptor function and, hence, in the ventilatory response to hypoxia.

Action Potentials↗

Cnidocyte mechanoreceptors are tuned to the movements of swimming prey by chemoreceptors.

Cnidocytes, the stinging cells of cnidarians, discharge nematocysts in response to physical contact accompanied by the stimulation of specific chemoreceptors. Cnidocytes in fishing tentacles of a sea anemone are now found to discharge nematocysts preferentially into targets vibrating at 30, 55, and 65 to 75 hertz. Moreover, in the presence of submicromolar concentrations of known chemosensitizers, such as N-acetylated sugars and mucin, these optima shift to 5, 15, 30, and 40 hertz, frequencies that correspond to the movements of swimming prey. Hence, chemoreceptors for these substances tune cnidocyte mechanoreceptors to frequencies that match the movements of the prey.

Animals↗

The fine structure of the olfactory and vomeronasal organs of a lizard (Tiliqua scincoides scincoides).

Olfactory epithelium in Tiliqua scinoides is of a loosely packed pseudostratified type. It receives secretion from the supporting cells and the underlying glands of Bowman. Its surface bears microvilli and cilia from sensory cells and microvilli from supporting cells. The vomeronasal epithelium is also pseudostratified but higher and more closely packed. Its surface carries microville from sensory and supporting cells but lacks cilia. Vascular connective tissue penetrates it almost to the epithelial surface but is always outlined by basal cell processes and a basal lamina. There are no secretory cells in or under the sensory epithelium but some cells in the epithelium of the mushroom body contain secretion granules. Sensory cells of both epithelia are bipolar neurons. The perikarya of the vomeronasal cells are more neuronal in character. Axonic processes are similar in both, dendrites are distinctive. Olfactory dendrites end in rounded rods bearing microvilli and cilia of an unusual type. Microville with filamentous cores occur on vomeronasal dendrites. There are no cilia, but 2-6 centrioles appear below the cell surface. Basal cells are structurally similar in both epithelia, but axonic processes of olfactory cells are surrounded by supporting cell processes, while vomeronasal axonic processes are surrounded by basal cells before they leave the epithelium. The presence of cilia and microville on the surface of the sensory cells is discussed in relation to the physical conditions surrounding them.

Animals↗

A tissue specific cytochrome P450 required for the structure and function of Drosophila sensory organs.

Cytochrome P450s have generally been acknowledged as broadly tuned detoxifying enzymes. However, emerging evidence argues P450s have an integral role in cell signaling and developmental processes, via their metabolism of retinoic acid, arachidonic acid, steroids, and other cellular ligands. To study the morphogenesis of Drosophila sensory organs, we examined mutants with impaired mechanosensation and discovered one, nompH, encodes the cytochrome P450 CYP303a1. We now report the characterization of nompH, a mutant defective in the function of peripheral chemo- and mechanoreceptor cells, and demonstrate CYP303a1 is essential for the development and structure of external sensory organs which mediate the reception of vital mechanosensory and chemosensory stimuli. Notably this P450 is expressed only in sensory bristles, localizing in the apical region of the socket cell. The wide diversity of the P450 family and the growing number of P450s with developmental phenotypes suggests the exquisite tissue and subcellular specificity of CYP303a1 illustrates an important aspect of P450 function; namely, a strategy to process critical developmental signals in a tissue- and cell-specific manner.

Amino Acid Sequence↗

Solitary chemosensory cells in the developing chemoreceptorial epithelium of the vallate papilla.

The solitary chemosensory cells are considered typical of aquatic vertebrates and have never been described in the oral cavity of terrestrial vertebrates. We describe elements with ultrastructural characteristics of the solitary chemosensory cell in the gustatory epithelium of rats in the first week of extrauterine life. These elements appeared isolated, located among keratinocytes, and wrapped by glial-like elements. They showed a bipolar shape with a round cell body, a thin apical process, and a thicker basal one. Nerve fibers contacted the cell body and the processes. The cells showed small dense granules mainly located near nerve contacts. Small bundles of tonofilaments were visible in the perinuclear cytoplasm. Similar elements were not found in rats after the first week of extrauterine life. The present study demonstrates in a mammal that the development of taste buds is preceded by the presence of epithelial elements with ultrastructural characteristics of the solitary chemosensory cells described in lower vertebrates. This finding suggests that the oral chemoreception in the suckling rats may be mediated by three different pathways (i.e., gustatory system, common chemical sense, and solitary chemosensory cell system).

Animals↗

The anion paradox in sodium taste reception: resolution by voltage-clamp studies.

Sodium salts are potent taste stimuli, but their effectiveness is markedly dependent on the anion, with chloride yielding the greatest response. The cellular mechanisms that mediate this phenomenon are not known. This "anion paradox" has been resolved by considering the field potential that is generated by restricted electrodiffusion of the anion through paracellular shunts between taste-bud cells. Neural responses to sodium chloride, sodium acetate, and sodium gluconate were studied while the field potential was voltage-clamped. Clamping at electronegative values eliminated the anion effect, whereas clamping at electropositive potentials exaggerated it. Thus, field potentials across the lingual epithelium modulate taste reception, indicating that the functional unit of taste reception includes the taste cell and its paracellular microenvironment.

Animals↗

O2-sensing mechanisms in excitable cells: role of plasma membrane K+ channels.

Although carotid chemosensitive glomus cells have been the most extensively studied from the vantage point of how cells sense the lack of O2, it is clear that all tissues sense O2 deprivation. In addition, all mammalian cells can trigger a cascade of events that, depending on the severity and duration of hypoxia-induced stress, can lead to permanent injury and death or to adaptation and survival. Crucial in this cascade, we believe, how the cascade is initiated, how O2 lack is detected by cells, and how these initial steps can activate further processes. In this chapter, we focus on the initial steps of O2 sensing in tissues most commonly studied, i.e. carotid glomus cells, central neurons, smooth muscle cells, and neuro-epithelial bodies of the airways. Recently it has become clear that plasma membranes of various tissues can sense the lack of O2, not only indirectly via alterations in the intracellular milieu (such as pH, Ca, ATP, etc), but also directly through an unknown mechanism that involves plasma-membrane K channels and possibly other membrane proteins. This latter mechanism is suspected to be totally independent of cytosolic changes because excised patches from plasma membranes were used in these experiments from carotid cells and neurons. There are a number of questions in this exciting area of research that pertain to the role of this plasma-membrane O2-sensing mechanism in the overall cell response, identification of all the important steps in O2 sensing, differences between O2-tolerant and O2-susceptible cells, and differences between acute and chronic cell responses to lack of O2.

Animals↗

[Protein-sensitive elements of the labellar sensillas of Musca domestica flies].

Using electrophysiological technique of registration of impulse activity in chemoreceptive cells of the labellar sensillae of the housefly, it has been demonstrated that taste hairs are not uniform in their properties. They differ from each other by the set of receptive elements which exhibit different sensitivity, range of selectivity and pattern of impulse activity. It was shown that albumen solution (10(-5) M) evokes the activity in 1--2 cells of a sensilla which are classified as water and sugar receptors. Among these receptors, protein-sensitive and protein-insensitive cells may be distinguished. Considering the inhomogeneity of sugar receptor sites, it was suggested that chemo receptive membranes in most sensitive to protein cells contain more numerous fructose receptive sites, that glucose ones.

Animals↗

Fine structure of the sensilla of Peripatopsis moseleyi (Onychophora).

Three types of sensilla occurring on the lips and on the antennae of Peripatopsis moseleyi have been investigated by scanning and transmission electron microscopy. On the lips sensory spines can be found which contain numerous cilia originating from bipolar receptor cells. They reach the tip of the spine where the cuticle is modified. The perikarya of the sensory cells, a large supporting cell with a complicated surface and a second type of receptor, form a bud-like structure and are surrounded by a layer of collagen fibrils. The second receptor cell bears apical stereocilia as well as a kinocilium which are directed towards the centre of the animal -- thus the cell appears to be turned upside down. The sensilla of the antennae are 1) sensory bristles containing two or three kinds of receptor cells, one of which bears an apical cilium and one kind of supportive cell and 2) sensory bulbs located within furrows consisting of receptor cells with branched cilia and two kinds of supportive cells which are covered by a modified thin cuticle. According to the electron microscopical findings the sensory spines on the lips are presumably chemoreceptors. The sensory bristles on the antennae can be regarded as mechanoreceptors and the sensory bulbs as chemoreceptors.

Animals↗

Intracellular recordings from salamander olfactory receptor cells.

Intracellular recordings were obtained from salamander olfactory receptor cells. The occurrence of an intracellular spike in response to the antidromic stimulation of the olfactory fibers was considered as a physiological criterion of a neuronal impalement. The mean resting potential was -56 +/- 9 mV (mean +/- S.D.; n = 70). Fifty-two cells presented a spontaneous spike activity lower than 2 impulses/s. Appropriate olfactory stimulation generally evoked a slow and graded decrease (up to 28 mV) of the intracellular potential. The input resistance of the cell decreased markedly during the response. The slow potential change induced a repetitive firing. Increasing the intensity of the olfactory stimulation increased the instantaneous frequency of firing (up to 25 s-1) and reduced the spike amplitude. The spikes presented an inflexion in the rising phase indicating a two-stage depolarization. With the strongest intensities of stimulation the impulse activity was stopped during the repolarizing phase of the cell response when the membrane potential was still appreciably depolarized.

Animals↗

Electrophysiological aspects of reflexogenic area in the chelonian, Geoclemmys reevesii.

Electrophysiological and anatomical experiments were performed to determine the baro- and chemoreceptive area of the turtle, Geoclemmys reevesii. The baro- and chemoreceptive sites are located in the aortic and pulmonary arches and the truncal region. The aortic arch is innervated by both the superior truncal nerve (aortic nerve) and a branch of the inferior truncal nerve arising from the ganglion trunci of the vagus. The pulmonary arch is innervated by a fine branch of the inferior truncal nerve, a branch containing mainly barosensitive and a few chemosensory fibers. On the peripheral cut end of these nerves, afferent electrical baro- and chemosensory activities were induced by NaCN and hyperoxia. Fluorescence microscopy revealed that the glomus cells are located in these regions.

Afferent Pathways↗