Reading: BCP Chapter 8 www.knowabouthealth.com The Chemical Senses Most ancestral and common of the senses. Present even in the simplest single-cell organisms (e.g., bacteria). Functions: Finding food sources Judging nutritional value and safety of foods. Avoiding predators and hazardous environments. Social communication, mating (“pheromones”). Monitoring internal physiological state. www.georgeretseck.com NSC201 Basic Neurobiology 2 “Taste” vs. “Flavor” Taste refers to the sensations relayed by taste receptor cells in the oral cavity. Foods activate different, unique combinations of only 5 basic tastes. Flavor depends on both taste (gustation) and smell (olfaction), i.e. flavor is a multisensory percept. Visual, auditory (crunch) and somatosensory (texture, pain and temperature) factors also influence the percept of flavor. NSC201 Basic Neurobiology 3 Tastes Five basic tastes: Sweet (non-ionic) Sour (ionic) Salty (ionic) Bitter (non-ionic) Umami (MSG, glutamate; non-ionic). Basic tastes are innate Other complex tastes are acquired and modified by experience. NSC201 Basic Neurobiology 4 Taste and Survival A sensitive and versatile taste system has great survival value for organisms exploiting wide variety of food sources. Gustation divides the world into Nutrients: generally attractive, e.g., sweet and salty. “Anti-nutrients”: repulsive, e.g., sour and bitter. Sensitivity to bitter tastants (which number in the thousands) can be extreme (nanomolar thresholds), because such tastants in the natural world are often poisonous. NSC201 Basic Neurobiology 5 Taste Chemistry Some taste perceptions are directly related to tastant chemistry: Salts (nutrients) taste “salty”. Acids (anti-nutrients) taste “sour”. Other common taste perceptions can be elicited by a wide variety of unrelated chemicals, e.g., “sweet”: Sugars Certain proteins (e.g. thaumatin) Sugar substitutes: saccharin (benzoic sulfinide), aspartame (aspartic acid/phenylalanine di-peptide), sucralose (“Splenda”, a chlorinated sucrose). NSC201 Basic Neurobiology 6 The Tongue Taste is primarily a function of the tongue. Taste buds are grouped in three of the four accessory structures called papillae: vallate, foliate and fungiform There are subtle regional differences in sensitivity to different tastes over the lingual surface, but most of the tongue is sensitive to all tastes. NSC201 Basic Neurobiology 7 Taste Buds and Taste Cells TB’s are morphologically specialized accessory epithelial structures containing taste cells. 2000 – 5000 TB’s on human tongue. There are roughly 50 – 100 taste cells per taste bud, along with basal stem cells. Modified epithelial (“short”) sensory receptors. Differentiate from basal stem cells in the taste bud. Turnover about every 10 days. NSC201 Basic Neurobiology 8 Functional Morphology of Taste Cells Taste cells are “short” receptors Apical pole: microvilli Protrude through taste pore into mucus of oral cavity. Provide large surface area to maximize contact with dissolved tastants. Basolateral pole: Contains typical organelles of epithelial cells. Synapses onto primary gustatory afferents (first-order taste neurons), which project to brain via the central gustatory pathway. NSC201 Basic Neurobiology 9 Taste Transduction Direct transduction: Some tastants are ions that carry currents through ion channels. Indirect transduction: Other (non-ionic) tastants bind selectively to specific G proteincoupled membrane receptors. Individual taste cells may employ both types of transduction. NSC201 Basic Neurobiology 10 Direct Transduction: Salty and Sour Salty: Na+ ions permeate amiloride- Salt Sour sensitive Na+ channels, directly depolarizing membrane. Sour: H+ ions (protons) permeate amiloride-sensitive Na+ channels… … AND block K+ channels, directly depolarizing membrane. Depolarization opens Nav and Cav channels, leading to AP’s (sometimes) and transmitter release, respectively. NSC201 Basic Neurobiology 11 Indirect Transduction: Bitter/Sweet/Umami Bitter, sweet, or umami tastants transduced by G protein-coupled pathways. Binding of tastant to receptor activates G protein/PLC/IP3 cascade. IP3 elevates internal [Ca2+] IP3 causes release of Ca2+ from internal stores. Opens unique Ca2+-activated Na+ channel, depolarizing membrane, opening voltagedependent Cav channels. Ca2+ influx further raises [Ca2+]]i, triggers transmitter release, synaptic stimulation of primary gustatory afferents. NSC201 Basic Neurobiology 12 Early Processing of Tastes Receptor potential magnitude proportional to both type AND concentration of tastant. Selectivity in Taste Cells Taste cell selectivity for basic tastes varies: 90% respond to two or more tastes. Primary gustatory afferents branch many times, innervating numerous taste buds and, within each taste bud, several taste cells. Thus, the electrical activity recorded from a single sensory fiber represents the input of many taste cells. Afferents exhibit taste preferences, suggesting they receive input from taste cells with common tastant selectivity. Taste Selectivity in Gustatory Afferents NSC201 Basic Neurobiology 13 Central Pathways for Taste Gustatory afferent neurons leave the mouth as part of the 7th, 9th, and 10th cranial nerves to the solitary nucleus of the medulla. Medulla ventral posterior nucleus of the thalamus primary and secondary gustatory cortex (ipsilateral). 10th 9th 7th NSC201 Basic Neurobiology 14 Central Representation of Taste Many central gustatory neurons exhibit a strong response to a specific tastant but are also able to respond to other tastants. Imaging studies of the gustatory cortex show that different tastes (salty, sour, sweet and bitter) are represented by specific spatial patterns containing both distinct and overlapping regions. These data support both a labeled line and a population (distributed) code of taste. www.cell.com NSC201 Basic Neurobiology 15 Pathways from Primary Gustatory Cortex Forebrain: Taste/Flavor Perception Primary Gustatory Cortex • Orbitofrontal cortex (location where chemical signals first merge to form percept of flavor). Medullary motor nuclei: Feeding Behavior Hypothalamus, Amygdala: Motivational & Hedonic Value of Food • • • • • Swallowing Chewing Gagging, vomiting Salivation Respiration • Hunger • Palatability NSC201 Basic Neurobiology 16