D’YOUVILLE COLLEGE BIOLOGY 659 - INTERMEDIATE PHYSIOLOGY I CHEMICAL SPECIAL SENSES Lecture 16: Chemical senses, Taste & Smell Chemical Senses: (chapter 53) • gustatory & olfactory senses detect information that establishes food preference & safety; smell also aids in detection of other animals - linked to pathways governing primitive emotions & behaviors (especially smell) • taste – leading role in perception of flavor of foods, augmented by smell & touch (texture of food); receptors are taste buds (ppts. 1 & 2) - qualities: now appears to be five primary tastes: - sour - related to acid; specific hydrogen ion receptor has been found - salty - mainly sodium; 2 sodium receptors, 2 potassium receptors and 1 chloride receptor have been found - sweet - associated with numerous organic chemicals such as sugars, alcohols, aldehydes & certain amino acids; 2 sweet receptors, also adenosine & inosine receptors associated with sweet taste - bitter - alkaloids (such as quinine) or organic nitrogen compounds, which are often poisonous; 2 bitter receptors known - ‘umami’ - poorly understood; believed to be associated with glutamate & aspartate (beefy, aging cheese taste); specific glutamate receptor has been recognized - sensitivity: relatively low concentrations of substances can be detected, (second messenger pathway that amplifies tastant stimulus), especially bitter substances, which often are toxic Bio 659, lec. 16 - p. 2 - - structure & location of taste buds: located mainly along sides of lingual papillae (circumvallate, fungiform & foliate), projections of mucosa on dorsal surface of tongue; also found on palate, epiglottis, mucosal arches & esophagus; receptor organ resembles cluster of bananas – sustentacular cells mingled with taste cells, whose microvilli surround an apical taste pore (fig. 53 – 1 & ppt. 3) - transduction (generation of taste nerve signals): tastant interacts with protein receptor on microvillus membrane causing opening of ion channel directly (salty or sour tastants) or via a second messenger pathway (sweet or bitter tastants) - causes depolarization; receptors excite taste nerve fibers to deliver strong signal initially, followed by sensory adaptation (weaker tonic signal during continuous tastant stimulation) - pathways (fig. 53 – 2 & ppt. 4): oral taste signals (Cr. N. VII – chorda tympani fibers of facial n., carried by lingual n.), posterior tongue & pharyngeal signals (Cr. N. IX – glossopharyngeal n.) & some pharyngeal signals (Cr. N. X – vagus n.) are conveyed to solitary tract in brainstem; synapse in gustatory nucleus of solitary tract - collaterals connect with brainstem salivatory nuclei, instigating salivation reflex - second order fibers ascend to ventral basal nucleus of thalamus (in proximity with medial lemniscal fibers carrying somatosensory signals from tongue to lateral post central gyrus); third order fibers from thalamus terminate in cortex of anterior insular lobe (ppts. 5 & 6) • smell: olfactory sense informs of presence of other animals even recognition of individuals (not well developed in humans); contributes to perception of flavor & influences emotional responses - olfactory area of nasal cavity: sensory field at summit of vestibule in each nasal cavity, spreading laterally over superior and some middle concha & spreading medially, over upper median septum (about 5 sq. cm. total for both nasal cavities) (ppt. 7) Bio 659, lec. 16 - p. 3 - - olfactory sensory cells: olfactory hair cells (modified bipolar neurons), surrounded by sustentacular cells; hair processes (modified cilia) spread into matlike mesh in mucous coating (product of Bowman’s glands)(fig. 53 – 3 & ppt. 8) - qualities of smell: current research has discredited former popular theory of seven primary odors; approx. 100 genes coding for specific odorant receptors & approx. 50 specific anosmias (loss of specific odorant sensitivity) have been identified - transduction: chemicals dissolve in mucus and bind to protein receptor that triggers second messenger pathway (fig. 53 - 4 & ppt. 9); second messenger (cyclic AMP) opens sodium channels causing depolarization; resting membrane potential (-55 mv.) causes background tonic signal (slow volley of action potentials) – receptor excitation results in increased signal strength (more rapid volley of APs) - pathways: receptor axons traverse cribriform plate of ethmoid bone to synapse with mitral cells & tufted cells in olfactory bulb; synaptic fields clustered into glomeruli – possibly sensitive to specific odorants (fig. 53 –3 & ppt. 8) - some second order fibers (fig. 53 – 5 & ppts. 10 to 12) in olfactory tract enter medial olfactory area (very old system), which governs primitive reflexes such as licking the lips & salivation; some sources have thrown doubts on existence of this area - other second order fibers in olfactory tract enter lateral olfactory area, which is subdivided into a ‘less old’ and a ‘newer’ pathway - less old pathway passes through portions of limbic system (amygdaloid nuclei and hippocampus) as well as medial cortex of temporal lobe (paleocortex), bypassing thalamus; appears responsible for development of food preferences or aversions - newer pathway passes through thalamus (dorsomedial nucleus) en route to frontal cortex (inferior surface); appears to facilitate identification of odorant and perceptual analysis (like other sensory cortex areas) - centrifugal (corticofugal) fibers pass in retrograde fashion (from higher brain levels back to olfactory bulb); deliver inhibitory signals that perform lateral Bio 659, lec. 16 - p. 4 - inhibition (sharpens input signal) and screening of sensory inputs (likely the major basis of sensory adaptation); mediated by granular cells of olfactory bulb that secrete GABA