The molecular biology of taste transduction

Taste cells respond to a wide variety of chemical stimuli: certain ions are perceived as salty (Na⁺) or sour (H⁺); other small molecules are perceived as sweet (sugars) and bitter (alkaloids). Taste has evolutionary value allowing animals to respond positively (to sweet carhohydrates and salty NaCl) or aversively (to bitter poisons and corrosive acids). Recently, some of the proteins involved in taste transduction have been cloned. Several different G proteins have been identified and cloned from taste tissue: gustducin is a taste cell specific G protein closely related to the transducins. Work is under way to clone additional components of the taste transduction pathways. The combination of electrophysiology, biochemistry and molecular biology is being used to characterize taste receptor cells and their sensory transduction mechanisms.     

Molecular mechanisms underlying the reception and transmission of sour taste information

Taste enables organisms to determine the properties of ingested substances by conveying information regarding the five basic taste modalities: sweet, salty, sour, bitter, and umami. The sweet, salty, and umami taste modalities convey the carbohydrate, electrolyte, and glutamate content of food, indicating its desirability and stimulating appetitive responses. The sour and bitter modalities convey the acidity of food and the presence of potential toxins, respectively, stimulating aversive responses to such tastes. In recent years, the receptors mediating sweet, bitter, and umami tastes have been identified as members of the T1R and T2R G-protein-coupled receptor families; however, the molecular mechanisms underlying sour taste detection have yet to be clearly elucidated. This review covers the molecular mechanisms proposed to mediate the detection and transmission of sour stimuli, focusing on polycystic kidney disease 1-like 3 (Pkd1l3), Pkd2l1, and carbonic anhydrase 4 (Car4).     

Responses of Primate Taste Cortex Neurons to the Astringent Tastant Tannic Acid

In order to advance knowledge of the neural control of feeding,we investigated the cortical representation of the taste oftannic acid, which produces the taste of astringency. It isa dietary component of biological importance particularly toarboreal primates. Recordings were made from 74 taste responsiveneurons in the orbitofrontal cortex. Single neurons were foundthat were tuned to respond to 0.001 M tannic acid, and representeda subpopulation of neurons that was distinct from neurons responsiveto the tastes of glucose (sweet), NaCl (salty), HCI (sour),quinine (bitter) and monosodium glutamate (umami). In addition,across the population of 74 neurons, tannic acid was as wellrepresented as the tastes of NaCI, HCI quinine or monosodiumglutamate. Multidimensional scaling analysis of the neuronalresponses to the tastants indicates that tannic acid lies outsidethe boundaries of the four conventional taste qualities (sweet,sour, bitter and salty). Taken together these data indicatethat the astringent taste of tannic acid should be consideredas a distinct taste quality, which receives a separate representationfrom sweet, salt, bitter and sour in the primate cortical tasteareas. Chem. Senses 21: 135–145, 1996.     

棉铃虫幼虫对人类呈味物质的取食反应

利用叶碟法在室内测定了棉铃虫对人类酸、甜、苦、咸4种基本呈味物质和麻、辣味2种植物提取物的取食反应。正交试验结果表明,棉铃虫幼虫对用甜味、苦味和辣味物质(蔗糖、奎宁和辣椒提取物)处理过的烟叶取食选择率较高,对这3种呈味物质表现出有较好的适应性;而幼虫对咸味、酸味和麻味物质(氯化钠、柠檬酸和花椒提取物)处理过的烟叶取食量较少,这3种呈味物质表现出较强的拒食活性。在选择性条件下,幼虫的取食量与花椒提取物剂量显著相关;而在非选择性条件下,幼虫的取食量与氯化钠剂量显著相关。     

Genetic variation in taste and its influence on food selection

Abstract Taste perception plays a key role in determining individual food preferences and dietary habits. Individual differences in bitter, sweet, umami, sour, or salty taste perception may influence dietary habits, affecting nutritional status and nutrition-related chronic disease risk. In addition to these traditional taste modalities there is growing evidence that "fat taste" may represent a sixth modality. Several taste receptors have been identified within taste cell membranes on the surface of the tongue, and they include the T2R family of bitter taste receptors, the T1R receptors associated with sweet and umami taste perception, the ion channels PKD1L3 and PKD2L1 linked to sour taste, and the integral membrane protein CD36, which is a putative "fat taste" receptor. Additionally, epithelial sodium channels and a vanilloid receptor, TRPV1, may account for salty taste perception. Common polymorphisms in genes involved in taste perception may account for some of the interindividual differences in food preferences and dietary habits within and between populations. This variability could affect food choices and dietary habits, which may influence nutritional and health status and the risk of chronic disease. This review will summarize the present state of knowledge of the genetic variation in taste, and how such variation might influence food intake behaviors.     

Autonomic nervous system responses associated with primary tastes

The hedonic dimension of the taste sensation plays a crucial role in the control of many taste-mediated responses related to food ingestion or rejection. The purpose of this study was to evaluate the emotional reactivity associated with each primary taste (sweet, salty, sour and bitter) through analysis of the variations of autonomic nervous system (ANS) parameters. Thirty-four healthy non-smoker volunteer subjects (17 males and 17 females, mean age = 28 years) participated in the experiment. Taste stimuli were solutions of 0.3 M sucrose (sweet), 0.15 M NaCl (salty), 0.02 M citric acid (sour) and 0.00015 M quinine sulfate (bitter). Evian mineral water was used as the diluent and control (neutral taste). Throughout the test, five ANS parameters (skin potential and skin resistance, skin blood flow and skin temperature, and instantaneous heart rate) were simultaneously and continuously recorded. Results of the ANOVA evidenced a significant effect of primary taste on skin resistance amplitude (P: < 0.001) and duration (P: < 0.0001), skin temperature amplitude (P: < 0.001), skin blood flow amplitude (vasoconstriction) (P: < 0.0001) and instantaneous heart rate increase (P: < 0.0001). Skin resistance and cardiac responses were the most relevant ANS parameters to distinguish among the taste solutions. The four primary tastes could be associated with significantly different ANS responses in relation to their hedonic valence: the pleasantly connoted and innate-accepted sweet taste induced the weakest ANS responses whereas the unpleasant connoted tastes (salty, sour and bitter) induced stronger ANS responses, the innate-rejected bitter taste inducing the strongest ones. Such a neurovegetative characterization of each primary taste could provide references for the hedonic analysis of the more complex gustative sensation attached to foods.     

INFLUENCE OF COLOR ON TASTE THRESHOLDS

Increasing molar concentrations of sweet, sour, bitter and saltywere evaluated in colorless and colored (red, green, yellow)water solutions by 28 untrained students. Green color statisticallyincreased sweet taste threshold sensitivity while yellow colordecreased taste sensitivity. Red color did not affect the tastesensitivity of sweet. In the case of sour, both yellow and greencolors decreased sensitivity with red having no affect. Redcolor decreased bitter taste sensitivity with yellow and greencolor having no effect. No significant differences due to coloraffected salty taste sensitivity. Thus, psychological colorassociation can alter reports of certain basic taste sensations. ^*Scientific Series Paper Number 1764 of the Colorado State UniversityExperiment Station.     

Taste development from health education among schoolchildren: a two-year intervention study

It is necessary to develop a system of nutritional education which can be understood among schoolchildren who have not yet received a basic education. In the present study, we conducted an educational program for lower-grade schoolchildren, which contained dish selection, an agricultural experience, a cooking experience, and a lecture on digestive absorption. We evaluated the effect of this program on development by measuring taste sensitivity regarding sweet, sour, salty and bitter tastes. For the baseline period, there was no significant difference between the intervention school and the control school in each variable. At follow-up periods, both the intervention and the control schools showed an increasing sense of taste. In the intervention school, development of sensitivity to the sweet, the sour, and the bitter taste was significant. In the control school, development of sensitivity to the sweet and the bitter taste was significant. The increases in the sense of the sour and the bitter tastes and the sum of the four tastes for the intervention subjects were significantly larger than comparable values for the control subjects. These results suggest that the development of taste sensitivity is affected by nutritional education for lower-grade elementary schoolchildren.     

TASTE INTENSITIES OF OIL-IN-WATER EMULSIONS WITH VARYING FAT CONTENT

The objective of this study was to determine the effect fat has on the intensity of sweet, salty, sour, bitter and umami tastes in oil-in-water emulsions. The first experiment used two levels of fat (9% and 17% in oil-in-water emulsions) and two intensities of each taste (high and low). We compared the taste intensities of these emulsions to the intensities of oil-free samples with equal total volume, and to oil free samples of the same aqueous taste compound concentrations. Because of potential confusion between taste intensity and viscosity, we repeated the experiment, having panelists rate both thickness and taste intensity. Diluting with oil, compared to diluting with water, decreased bitterness, but increased the intensity of salty, sweet, sour and umami tastes. When compared to samples with equal aqueous taste compound concentrations, fat suppressed bitterness, but had no effect on the other tastes.     

The good taste of peptides

The taste of peptides is seldom one of the most relevant issues when one considers the many important biological functions of this class of molecules. However, peptides generally do have a taste, covering essentially the entire range of established taste modalities: sweet, bitter, umami, sour and salty. The last two modalities cannot be attributed to peptides as such because they are due to the presence of charged terminals and/or charged side chains, thus reflecting only the zwitterionic nature of these compounds and/or the nature of some side chains but not the electronic and/or conformational features of a specific peptide. The other three tastes, that is, sweet, umami and bitter, are represented by different families of peptides. This review describes the main peptides with a sweet, umami or bitter taste and their relationship with food acceptance or rejection. Particular emphasis will be given to the sweet taste modality, owing to the practical and scientific relevance of aspartame, the well‐known sweetener, and to the theoretical importance of sweet proteins, the most potent peptide sweet molecules. Copyright © 2011 European Peptide Society and John Wiley & Sons, Ltd.     

Activation of Polycystic Kidney Disease-2-like 1 (PKD2L1)-PKD1L3 Complex by Acid in Mouse Taste Cells

Five basic tastes (bitter, sweet, umami, salty, and sour) are detected in the four taste areas where taste buds reside. Although molecular mechanisms for detecting bitter, sweet, and umami have been well clarified, those for sour and salty remain poorly understood. Several channels including acid-sensing ion channels have been proposed as candidate sour receptors, but they do not encompass all sour-sensing abilities in vivo. We recently reported a novel candidate for sour sensing, the polycystic kidney disease-2-like 1 (PKD2L1)-PKD1L3 channel complex. This channel is not a traditional ligand-gated channel and is gated open only after removal of an acid stimulus, called an off response. Here we show that off responses upon acid stimulus are clearly observed in native taste cells from circumvallate, but not fungiform papillae, of glutamate decarboxylase 67-green fluorescent protein (GAD67-GFP) knock-in mice, from which Type III taste cells can be visualized, using Ca²⁺ imaging and patch clamp methods. Off responses were detected in most cells where PKD2L1 immunoreactivity was observed. Interestingly, the pH threshold for acid-evoked intracellular Ca²⁺ increase was around 5.0, a value much higher than that observed in HEK293 cells expressing the PKD2L1-PKD1L3 complex. Thus, PKD2L1-PKD1L3-mediated acid-evoked off responses occurred both in HEK293 cells and in native taste cells, suggesting the involvement of the PKD2L1-PKD1L3 complex in acid sensing in vivo.     

Coding of sweet,bitter, and umami tastes: different receptor cells sharing similar signaling pathways

Mammals can taste a wide repertoire of chemosensory stimuli. Two unrelated families of receptors (T1Rs and T2Rs) mediate responses to sweet, amino acids, and bitter compounds. Here, we demonstrate that knockouts of TRPM5, a taste TRP ion channel, or PLCbeta2, a phospholipase C selectively expressed in taste tissue, abolish sweet, amino acid, and bitter taste reception, but do not impact sour or salty tastes. Therefore, despite relying on different receptors, sweet, amino acid, and bitter transduction converge on common signaling molecules. Using PLCbeta2 taste-blind animals, we then examined a fundamental question in taste perception: how taste modalities are encoded at the cellular level. Mice engineered to rescue PLCbeta2 function exclusively in bitter-receptor expressing cells respond normally to bitter tastants but do not taste sweet or amino acid stimuli. Thus, bitter is encoded independently of sweet and amino acids, and taste receptor cells are not broadly tuned across these modalities.     

A serotonin-sensitive sensor for investigation of taste cell-to-cell communication

Taste receptor cells are the taste sensation elements for sour, salty, sweet, bitter and umami sensations. It was demonstrated that there are cell-to-cell communications between type II (sour) and type III (sweet, bitter and umami) taste cells. Serotonin (5-HT) is released from type III cells, which is the only type of taste cells that has synaptic process with sensory afferent fibers. Then, taste information is transmitted via fibers to the brain. During this process, 5-HT plays important roles in taste information transmission. In order to explore a sensor to detect 5-HT released from taste cell or taste cell networks, we develop a 5-HT sensitive sensor based on LAPS chip. This sensor performs with a detection limit of 3.3 × 10(-13)M and a sensitivity of 19.1 mV per concentration decade. Upon the stimuli of sour and mix (bitter, sweet and umami) tastants, 5-HT released from taste cells could be detected flexibly, benefit from the addressability of LAPS chip. The experimental results show that the local concentration of 5-HT is around several nM, which is consistent with those from other methods. In addition, immunofluorescent imaging technique is utilized to confirm the functional existence of both type II and III cells in a cluster of isolated taste cells. Different types of taste cells are labeled with corresponding specific antibody. This 5-HT sensitive LAPS chip provides a potential and promising way to detect 5-HT and to investigate the taste coding and information communication mechanisms.     

Some Basic Psychophysics of Calcium Salt Solutions

Detection thresholds and the taste qualities of suprathresholdconcentrations of calcium salt solutions were assessed. Averagetaste detection thresholds for calcium chloride (CaCl₂), lactate(CaLa), hydroxide, phosphate and gluconate ranged between 8and 50 mM, with no reliable differences among the various salts.Between-subject variability ranged over four orders of magnitudeand reliability coefficients for repeated detection thresholdtests of CaCl₂ averaged r = 0.52. In an odor detection test,subjects could reliably discriminate 100 but not 1 mM CaCl₂and CaLa from water. The taste of suprathreshold concentrations(1–100 mM) of CaCl₂ and CaLa was considered unpleasant.At 1 mM, CaCl₂ solution was rated as 35% bitter, 32% sour, 29%sweet and 4% salty. At higher concentrations the sweet componentdiminished and the salty component increased, so that 100 mMCaCl₂ was rated as 44% bitter, 20% sour, 1% sweet and 35% salty.CaLa solutions were considered to be significantly less bitterand marginally more sour than equimolar CaCl₂ solutions. Thus,the taste of calcium varied with both the form and concentrationof salt tested, but included both sour and bitter components.Saltiness was identified only in high (     

Identification and characterization of human taste receptor genes belonging to the TAS2R family

The sense of taste is a chemosensory system responsible for basic food appraisal. Humans distinguish between five primary tastes: bitter, sweet, sour, salty and umami. The molecular events in the perception of bitter taste are believed to start with the binding of specific water-soluble molecules to G-protein-coupled receptors encoded by the TAS2R/T2R family of taste receptor genes. TAS2R receptors are expressed at the surface of taste receptor cells and are coupled to G proteins and second messenger pathways. We have identified, cloned and characterized 11 new bitter taste receptor genes and four new pseudogenes that belong to the human TAS2R family. Their encoded proteins have between 298 and 333 amino acids and share between 23 and 86% identity with other human TAS2R proteins. Screening of a mono-chromosomal somatic cell hybrid panel to assign the identified bitter taste receptor genes to human chromosomes demonstrated that they are located in chromosomes 7 and 12. Including the 15 sequences identified, the human TAS2R family is composed of 28 full-length genes and 16 pseudogenes. Phylogenetic analyses suggest a classification of the TAS2R genes in five groups that may reflect a specialization in the detection of specific types of bitter chemicals.     

灵长类苦味受体基因研究进展

在鲜味、甜味、苦味、咸味和酸味5种味觉形式中,苦味能避免动物摄入有毒有害物质,在动物的生存中发挥着特别重要的作用。苦味味觉的产生依赖于苦味物质与苦味受体的相互作用。苦味受体由苦味受体基因Tas2rs编码,此类基因在不同物种中数量变化较大以适应不同的需求。目前的研究在灵长类中鉴别出了若干苦味受体的配体,并发现有的苦味受体基因所经受的选择压在类群之间、基因之间甚至同一基因不同功能区之间都存在着变化。本文从苦味受体作用的多样性特点,受体与配体的对应关系、受体基因进化模式与食性之间的关系、苦味受体基因的适应性进化方面对灵长类苦味受体基因进行了综述,以期为苦味受体基因在灵长类中的深入研究提供参考。     

Proteomic analysis of human whole and parotid salivas following stimulation by different tastes

Whole and parotid salivas, collected after stimulation with tastants, were analyzed by 2D electrophoresis and mass spectrometry. In whole saliva, the number of proteins affected by taste stimulation increased in the order sweet < umami < bitter < acid. Annexin A1 and calgranulin A, involved in inflammation, were over-represented after umami, bitter, and sour stimulations. Their low abundance or absence in parotid saliva after bitter stimulation suggested that they originated from other oral glands or tissues.     

Reduction of saltiness and bitterness after a chlorhexidine rinse

Chronic rinsing with chlorhexidine, an oral-antiseptic, has been shown to decrease the saltiness of NaCl and the bitterness of quinine. The effect of acute chlorhexidine on taste has not been investigated. The purpose of the present study was to examine the effect of acute chlorhexidine rinses on taste intensity and quality of 11 stimuli representing sweet, salt, sour, bitter and savory. All stimuli were first matched for overall intensity so the effects of chlorhexidine would be directly comparable across compounds. As a control treatment, the bitter taste of chlorhexidine digluconate (0.12%) was matched in intensity to quinine HCl, which was found to cross-adapt the bitterness of chlorhexidine. Subjects participated in four experimental conditions: a pre-test, a quinine treatment, a chlorhexidine treatment, and a post-test condition, while rating total taste intensity and taste qualities in separate test sessions. Relative to the quinine treatment, chlorhexidine was found to decrease the salty taste of NaCl, KCl and NH4Cl, and not to significantly affect the tastes of sucrose, monosodium glutamate (MSG), citric acid, HCl and the taste of water. The bitter taste of urea, sucrose octa-acetate and quinine were suppressed after chlorhexidine rinses relative to water rinses, but were only marginally suppressed relative to quinine rinses. Potential mechanisms are discussed.     

Temporal properties of the human taste system.

An attempt to apply linear systems methods to the temporal properties of the human taste system is described. The percentage modulation in concentration required to detect a fluctuation in intensity was measured as a function of frequency of modulation. Both a sine-wave and a square-wave input were used. The sensitivity of the taste system at its maximum, and also at the low frequencies, was greater than expected from previous work based on classical methodology. The sensitivity of the taste system at its maximum, and also at the low frequencies, was greater then expected from previous work base on classical methodology. The sensitivity of the taste system to the various qualities was, in decreasing order, salty, sweet, sour, and bitter. The taste system in insensitive to frequencies above about 5 Hz. Except for bitter, representatives of the various taste qualities yielded similar functions within qualities.