BS3033 Block 2: Transducing the Senses. Paul Glynn (pg8@leicester.ac.uk) Sensory Transducers: Chemoceptors & Mechanoceptors. Sense Olfaction Transducer Cell Olfactory neuron Transducer Molecule(s) GPCR Taste Taste-cell GPCR or Ion-channel Temperature/Pain Ad- or C-neuron TRP-channel and ? ITCH C-neuron GPCR + TRP-channel Touch Ab-neuron + non-neural cell Piezo2-channel or ? Hearing Inner Hair Cell Mechanosensitive channel Vision Photoceptor cell GPCR (Rhodopsin) The Chemical Senses. Evolutionarily ancient: eg, C. elegans has chemical sense. Mammals have chemoreceptors: On external/internal surfaces detect H+, capsaicin, etc. (Somatosensory transducers). In specialized organs: Olfactory epithelium (in nose) detects odorants. Vomeronasal organ detects pheromones (vestigial in humans). Taste buds (on tongue) detect ingested chemicals. Chemical Senses: (i) Taste (Gustation). Gustatory neural pathway. • Three cranial nerves (cell bodies in trigeminal ganglia) convey signal from taste-buds on tongue and epiglottis, past ganglia, to synapse in medulla (gustatory nucleus). • Secondary afferents carry signal to synapse in thalamus (ventral posterior medial nucleus). • Tertiary afferents carry signal to synapse in cortex (primary gustatory). Most taste (gustation) cells are on tongue taste buds. There are 5 basic tastes: 1. Saltiness (Na+). 2. Sourness (H+). 3. Sweetness (eg, Sucrose). 4. Bitterness (eg, Quinine). 5. Umami (eg, Glutamate). So how do humans perceive such a wide variety of tastes and flavours? Neural coding of taste. * Humans have 2000-5000 taste buds each with 50-150 taste cells. * Most taste cells have rather broad tastant specificity. * Each gustatory afferent axon receives input from several taste buds. * Each secondary gustatory neuron in medulla receives input from several primary afferent axons. * Responses of a large number of broadly-tuned neurons specify properties of a particular taste. Specificities of taste cell responses to different tastants. Expose taste bud to tastants. Record Vm of cells and afferent gustatory axons. Cell 1 and axon 1: responds to NaCl only. Cell 2 and axon 2: responds to NaCl, Quinine and HCl. General finding: Taste cells respond broadly to bitter, sour and salt, but more selectively to sweet tastants. Signal transduction in taste cells. Na+ directly permeate constitutively-open Na-channel membrane depolarisation. H+ permeate Na-channels and block constitutivelyopen K-channels membrane depolarisation. Sucrose activates GPCR AC activation cAMP PKA activation phosphorylation and block of constitutively-open K-channels membrane depolarisation. In all, depolarisation opens VOCC [Ca2+]i vesicular release of Glutamate activation of afferent gustatory axon. Signal transduction in taste cells – 2. Analogous mechanisms operate in taste cells for bitter and umami tastants. Taste (Gustation): Summary. * Tongue taste cell transducers are GPCRs or ion-channels. * Tastant binding to receptor [Ca2+]i Glu release action potential in afferent axon of cranial nerve... medulla thalamus cortex (primary gustatory). * Specific taste represents response of large number of broadly-tuned neurons.