NEUROPHYSIOLOGY AND HUMAN TASTE SENSATIONS

Taste sensations are of primary importance in food flavor. Any attempt to synthesize chemically the flavor of a natural food involves mainly taste active compounds. Many distinct taste sensations can be identified as associated with food compounds. Thirteen different taste sensations are discussed herein. These different taste sensations are differentiated on the basis of stimulus chemistry and peripheral nerve conveying the taste information. Neurophysiological examination of the peripheral nerves involved in taste reveals that the sensory neurons can, in any species, be subdivided into distinct neural groups. These different neural groups respond to distinct classes of chemicals and often display different neurophysiological characteristics. Altogether in four different species, nine functional neural taste groups can be distinguished. In many cases, these neural groups can be taken as analogs for the neural groups assumed to underly human taste sensations. Distinct human taste sensations can be considered to arise from the excitation or inhibition of different neural groups. For certain human taste sensations there are no animal neural analog groups; and for certain neural groups there are no analog human sensations.     

The Effect of Adaptation on the Taste Threshold Observed with a Semiautomatic Gustometer

A special "gustometer" has been built, which permits continuous measurement of the threshold of taste sensations. Taste sensations show strong adaptation even at the threshold level. The adaptation was much the same for the four primary tastes, but there were large individual differences. Of theoretical interest is the finding that adaptation to a taste can occur even with subliminal stimuli.     

TASTE ACUITY IN THE ELDERLY: THE IMPACT OF THRESHOLD, AGE, GENDER, MEDICATION, HEALTH AND DENTAL PROBLEMS

This study was designed to assess taste threshold and investigate the effect of age, dental problems, health disorders/diseases and multiple medications on the distortion of taste and to determine if the loss of taste increases with age. One hundred seven elderly from rural congregate feeding sites and senior centers in Northern Alabama volunteered to participate in this study. Water solutions of varying concentrations for sweet, salty, sour, and bitter were prepared to determine recognition thresholds and test the effect of other factors on the ability to taste. A questionnaire was administered to each elderly volunteer to assess overall health. Results indicated that there was no significant age difference for the basic taste sensations, except for sweetness. Gender, dental problems, health disorders, diseases and multiple uses of medications did not significantly affect taste threshold. The basic was a decrease in taste threshold independent of any extraneous factors.     

UNDERSTANDING MOUTHFEEL ATTRIBUTES: A MULTIDIMENSIONAL SCALING APPROACH

Multidimensional scaling (MDS) was used to study qualitative relationships among mouthfeel attributes encountered in oral healthcare products. Similarity estimates were obtained from a rapid sorting task and from pairwise similarity ratings. Configurations were interpreted as suggesting four groupings of oral sensations: numbing, astringency, pain and taste. The pain-associated sensations were further differentiated into thermally related sensations and chemically related sensations in some configurations. Two-dimensional solutions from the sorting task and from group-averaged similarity ratings were similar. Individual differences scaling solutions, however, showed unacceptably high stress in two dimensions, suggesting additional nuances in meaning to individual panelists that were not captured by group-averaged data or by sorting data.     

Preference for dietary fat: From detection to disease

Recent advances in the field of taste physiology have clarified the role of different basic taste modalities and their implications in health and disease and proposed emphatically that there might be a distinct cue for oro-sensory detection of dietary long-chain fatty acids (LCFAs). Hence, fat taste can be categorized as a taste modality. During mastication, LCFAs activate tongue lipid sensors like CD36 and GPR120 triggering identical signaling pathways as the basic taste qualities do; however, the physico-chemical perception of fat is not as distinct as sweet or bitter or other taste sensations. The question arises whether “fat taste” is a basic or “alimentary” taste. There is compelling evidence that fat-rich dietary intervention modulates fat taste perception where an increase or a decrease in lipid contents in the diet results, respectively, in downregulation or upregulation of fat taste sensitivity. Evidently, a decrease in oro-sensory detection of LCFAs leads to high fat intake and, consequently, to obesity. In this article, we discuss recent relevant advances made in the field of fat taste physiology with regard to dietary fat preference and lipid sensors that can be the target of anti-obesity strategies.     

Taste suppression following lingual capsaicin pre-treatment in humans

The effect of oral capsaicin on taste sensations in humans was reinvestigated with attention to methodological issues raised in previous studies, including the mode of presentation and temperature of the tastant stimulus, as well as the sensitizing and desensitizing properties of capsaicin. One-half of the dorsal anterior tongue was pre-treated with capsaicin, followed by bilateral tastant application (sucrose, NaCl, quinine, monosodium glutamate and citric acid). Subjects indicated on which side the taste intensity was greater in a two-alternative, forced-choice procedure and also rated taste intensity independently on each side of the tongue. Each of the five tastants was tested sequentially, with reapplication of capsaicin between trials in order to maintain a constant level of burn. Four experiments were conducted: (i) a high concentration (33 p.p.m.) (109 microM) capsaicin effect on taste intensity elicited by high tastant concentrations; (ii) a high concentration capsaicin effect on taste intensity elicited by low tastant concentrations; (iii) a low concentration (1.5 p.p.m.) (4.9 microM) capsaicin effect on taste intensity elicited by low tastant concentrations; and (iv) validation of the method for localizing taste by pre-treating one side of the tongue with Gymnema sylvestre, followed by bilateral application of sucrose. In the first experiment, a significant proportion of the subjects chose the non-treated side in the two-alternative, forced-choice procedure and assigned significantly higher ratings to that side for sucrose-induced sweetness, quinine-induced bitterness and glutamate-induced umami sensations. Salty and sour sensations were not different between sides. A 15 min break was imposed in order to allow the capsaicin burn to disappear and desensitization to set in, followed by reapplication of the tastant test solutions. There were no bilateral differences in the intensity of the sensations elicited by any of the five tastants. Similar results were obtained in experiments 2 and 3. In the fourth experiment, all 15 subjects tested chose the side not treated with Gymnema sylvestre as having a stronger sweet taste and assigned significantly higher ratings to that side, thereby validating the method for taste localization. These results indicate that oral capsaicin reduces certain but not all taste sensations and are discussed in terms of possible physiological and cognitive interactions.     

Selective removal of a target stimulus localized by taste in humans

Recent studies have shown that people can localize a punctate gustatory stimulus on the lingual epithelium in the absence of discriminative tactile cues. The present studies examined the human ability to localize taste sensations on the tongue and to use this information to remove selectively a target stimulus (a flavored, 1 cm(3) gelatin cube) from the mouth when presented with non-target distractors that vary in number and taste. Findings indicate that humans are capable of localizing and removing either an aversive or an appetitive gustatory target from a field of tactile distractors via taste sensations alone, although this ability diminishes as the number of distractors increases (implicating serial searches, rather than parallel). In addition, humans can localize and selectively remove a target taste in the presence of distractors of another distinct taste quality. Under these conditions performance is either unaffected or reduced, which indicates that contrast with the distinct taste of the distractors does not enhance performance. Humans also are capable of removing a nearly tasteless cube from a field of flavored distractors, but this is clearly a more difficult task, suggesting that 'tactile capture' of taste occurs for the tasteless target cube and interferes with the localization of taste. Finally, perceived suprathreshold stimulus intensity did not seem to be related to the ability to localize and remove a target stimulus via taste sensations and failed to account for variations in performance across individuals.     

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.     

Multiple sensitivity of single taste cells of the frog tongue to four basic taste stimuli

The electrical response of the taste cells of the frog fungiform papillae to four fundamental taste solutions (NaCl, acetic acid, quinine-HCl and sucrose) was studied by using the intracellular recording technique. The average value of resting membrane potential was 22.5 mV, inside negative. Each of the four taste solutions applied to the tongue produced a slow depolarizing potential, the receptor potential, on which no spike potential was superimposed. The amplitude of the receptor potentials increased linearly as a function of the logarithm of the concentration of the stimulus. Amplitudes of depolarizations to a given taste stimulation varied from one cell to another even within a single taste bud. Most of the cells responded to more than two of the four basic taste solutions. Sensitivity patterns in terms of the number of effective solutions and the relative effectiveness of different kinds of solutions were variable among cells. Statistical analysis suggests that at the receptor membranes of the taste cells, the sensitivities for the four basic stimuli are independent and random.     

味觉刺激引起大鼠臂旁核神经元抑制性反应

本研究以轻度麻醉的大鼠为对象,应用细胞外微电极记录技术,观察并分析了脑桥臂旁核抑制性味觉神经元的自发活动及其对NaCl、HCl、盐酸奎宁（quinine HCl,QHCl））和蔗糖等四种基本味觉刺激的反应。共分析了18个具有自发活动的抑制性味觉神经元,自发放电频率分布在0．2～5．5Hz之间,平均放电频率（2．15±0．31）Hz。18个神经元中,1个神经元对单一味觉刺激呈抑制性反应,其余17个神经元对两种或两种以上的基本味觉刺激发生抑制性反应,且抑制具有潜伏期短、持续时间较长等特征。抑制持续时间5～80S,部分神经元表现为后抑制效应。根据神经元对四种基本味觉刺激呈抑制性反应的程度,将其分为NaCl优势神经元（n=8）,HCl优势神经元（n=3）,QHCl优势神经元n=3）和蔗糖优势神经元n=4）。其中NaCl优势神经元的反应谐宽最高（0．945）。这些神经元对欣快或厌恶刺激的区别能力较低。结果提示,在脑桥臂旁核存在对味觉刺激起抑制性反应的神经元,这些味觉神经元可能在味觉的调制及对欣快和厌恶刺激的编码中发挥重要的作用。     

Behavioral discrimination between glutamate and the four basic taste substances in mice

1. Behavioural studies using the conditioned taste aversion (CTA) paradigm in mice showed that aversion conditioned to monosodium L-glutamate (MSG), which elicits a unique taste in humans, did not strongly generalize to any of the four basic taste stimuli, suggesting that mice could behaviourally discriminate between MSG and the four basic taste stimuli. 2. Denervation of bilateral glossopharyngeal nerve significantly increased behavioural similarities (the strength of generalization in the CTA paradigm) between MSG and sodium salts. This was not the case after destruction of the bilateral chorda tympani nerve. 3. These results suggest that taste information of glossopharyngeal nerve plays a more important role in the behavioural discrimination between MSG and the four basic tastes than does that of the chorda tympani nerve.     

The psychophysical relationship between bitter taste and burning sensation: evidence of qualitative similarity

Although it has long been studied as a pure sensory irritant, the ability of capsaicin to evoke, mask, and desensitize bitter taste suggests that burning sensations and bitter taste might be closely related perceptually. The current study investigated the psychophysical relationship between bitterness and burning using 2 different approaches. In Experiment 1, spatial discrimination of 4 taste stimuli was measured in the presence or absence of capsaicin. The subjects' task was to report which of 3 swabs, spaced 1 cm apart and presented to the tongue tip, contained a taste stimulus when 1) water was presented on the other 2 swabs or 2) when 10 muM capsaicin was presented on all 3 swabs. The presence of capsaicin did not change performance on the 3 alternative forced-choice (3-AFC) task for sweet, sour, and salty stimuli, while the localization error for 1.8 mM quinine sulfate (QSO(4)) increased significantly. In Experiment 2, the perceptual similarity/dissimilarity of taste stimuli and capsaicin was measured directly using pairs of stimuli applied to opposite sides of the tongue tip on swabs separated by 2 cm. Multidimensional scaling analyses showed that capsaicin fell nearer to QSO(4) than to any other taste stimulus. Cluster analysis corroborated this finding: capsaicin was closely linked with QSO(4) and the capsaicin-QSO(4) group was separated from the other taste stimuli. The latter result indicated that bitterness was more similar to burning than to the other tastes. These findings imply that despite being mediated by different sensory modalities, bitterness and burn are qualitatively similar. We speculate that this similarity reflects a common function of these 2 sensations as sensory signals of potentially harmful stimuli.     

Psychophysical evidence that oral astringency is a tactile sensation

Aluminum potassium sulphate [AIK(SO₄)₂ 12H₂O—‘alum’]was tested for its ability to elicit oral astringency independentlyof its ability to elicit taste. First, the astringency producedby alum (10 g/l; 21.1 mM) on the non-gustatory surfaces betweenthe gum and the upper lip was measured. Subjects reported thatalum elicited sensations of astringency when the upper lip wasmoved laterally against the gum. Second, subjects dipped thetongue into two solutions—alum or a mixture solution thatapproximated the taste of alum—and attempted to determinewhich solution was more astringent. A slight tendency was foundfor subjects to identify the mixture as more astringent thanthe alum. However, the same subjects reported the alum to bemore astringent than the mixture when the same solutions wereapplied under the upper lip. These two experiments support thehypothesis that tactile stimulation is important for producingastringency, whereas taste stimulation of the anterior tongueis not. Third, after the application of alum, lubricating rinses(water, sucrose, Salivart®, corn oil, and the subjects'own saliva) were compared for their ability to decrease astringency.The lubricants reliably decreased original astringency, butto varying degrees depending upon their lubricating properties.All three experiments suggest that tactile sensations causedby increased friction (decreases,in salivary lubrication) betweenoral membranes are the primary basis of astringent sensations.     

The role of sodium and potassium transport pathways in taste transduction: an overview

Recent studies have shown that taste sensations are mediatedby a multiplicity of transduction mechanisms. The taste of saltis produced in part by the entry of Na⁺ through channels inthe apical taste cell membrane. Na⁺ transport also mediatessweet perception in some species. The taste of KCI requiresentry of K⁺ through apical potassium channels. The productionof second messengers such as cAMP by taste stimuli or tastemodifiers can depolarize taste cells by inducing an enzymaticcascade that alters K⁺ permeability.     

Wine tannins,saliva proteins and membrane lipids

Polyphenols have been part of human culture for about 6000 years. However, their mode of action in relation to wine tasting while eating is only beginning to be understood. This review, using analytical techniques and physicochemical concepts, attempts to summarize current knowledge and present an integrated view of the complex relationship between tannins, salivary proteins, lipids in food and in oral membranes. The action of tannins on taste sensations and astringency depends on their colloidal state. Although taste sensations are most likely due to interactions with taste receptors, astringency results from strong binding to proline-rich salivary proteins that otherwise lubricate the palate. Tannins disorder non-keratinized mucosa in mouth, possibly perturbing taste receptor function. The 10–15% ethanol present in wines potentiates this action. Cholesterol present in large quantities in keratinized mucosa prevents any disordering action on these oral membranes. Polyphenols bind strongly to the lipid droplets of fatty foods, a situation that reduces the astringency perceived when drinking a tannic wine, the so-called “camembert effect”. Based on binding constants mainly measured by NMR, a comprehensive thermodynamic model of the interrelation between polyphenols, salivary proteins, lipids and taste receptors is presented.     

TASTE PERCEPTION OF UMAMI-RICH DISHES IN ITALIAN CULINARY TRADITION

The aim of this research project was to investigate umami taste properties of recipes based on Italian culinary tradition and prepared with umami-rich ingredients, focusing on the impact of the preparation and ingredient combination. Gustative profiles were prepared for a traditional Italian dish, tender beef bouillon, usually consumed with pasta stuffed with beef. Four different samples were designed by changing the ingredients (with or without integrating Parmigiano cheese) and the preparation (cooking time) of the recipe. Panelists were rigorously trained for umami taste evaluation using monosodium L-glutamate (MSG) aqueous solutions and were then asked to evaluate umami sensation in tender beef bouillon with or without added Parmigiano Reggiano . A majority of the panelists were able to distinguish correctly umami sensations induced by MSG. The level of umami enhancement induced by Parmigiano Reggiano was clearly perceived by the panelists, and this enhancement positively affected also other basic tastes; whereas the cooking time had no clear effect on the gustative perception.

PRACTICAL APPLICATIONS

In western countries, people do not know much about umami. This taste is defined as the "savoriness" of the glutamate. Umami-taste substances are present in several foods, but whereas the taste of monosodium glutamate and 5' nucleotides can be without difficulty identified in water solutions, which are usually employed for panel training, the ability to identify this primary taste decreases enormously in more complex matrices like food. This study describes a procedure for screening and training sensory panels and could serve as a guide in teaching panelists to recognize and quantify the umami taste in a multistimuli matrix like a food recipe or product.
It also provides a practical application in a recipe in which the umami taste is modulated by the culinary preparation and ingredients.     

'Thermal taste' predicts higher responsiveness to chemical taste and flavor

Individual differences in taste perception have been explained in part by variations in peripheral innervation associated with the genetic ability to taste the bitter substances PTC and PROP. In the present study we report evidence of another source of individual differences that is independent of taste stimulus, taste quality, or gustatory nerve. Individuals who perceived taste from thermal stimulation alone (thermal taste) gave significantly higher taste ratings to chemical stimuli--often by a factor of >2:1--than did individuals who perceived no taste from thermal stimulation. This was true for all taste stimuli tested (sucrose, saccharin, sodium chloride, citric acid, quinine sulfate, MSG and PROP), for all three gustatory areas of the mouth (anterior tongue, posterior tongue and soft palate) and for whole-mouth stimulation. Moreover, the same individuals reported stronger sensations from the olfactory stimulus vanillin, particularly when it was sensed retronasally. The generality of the thermal-taster advantage and its extension to an olfactory stimulus suggests that it arises from individual differences in CNS processes that are involved in perception of both taste and flavor.     

Contributions of Smell and taste to overall intensity

The integration of olfaction and gustation in producing thesensation of flavor was studied using an almond extract stimuluspresented with the Two-Module Delivery System. Almond in themouth was found to influence the scaling of smell. The datasugest that this background effect is due to the diffusion ofairborne molecules from the oral cavity to the olfactory receptorsarea. At least under some stimulation conditons, almond vaporin the nose was shown to afect the scaling of taste. When scalingoverall intensity, subjects apparently added together the sensationsof smell and taste to produce the sensation of overall intensity.However, for all combinations of odorants and tastants, theestimate of overall intensity was less than the sum of the estimateof the smell and the estimate of the taste. The data furthersuggest that when estimating the overall intensity, the magnitudeassigned to the olfactory or gustatory sensations was less thanthat which have been assigned when estimating just smell ortaste.     

Changes in Taste Perception Following Mental or Physical Stress

Taste perception depends not only on the chemical and physicalproperties of tastants, but may also depend on the physiologicaland psychological conditions of those who do the tasting. Inthis study, the effects of mood state on taste sensitive wasevaluated in humans who were exposed to conditions of mentalor physical fatigue and tension. Taste responses to quininesulfate (bitter), citric acid (sour) and sucrose (sweet) weretested. The intensity of the taste sensations were recordedby a computerized time-intensity (TI) on-line system. Subjectsperformed mental tasks by personal computer or physical tasksby ergometer for 10–40 min. Before and after these sessions,the duration of the after-taste and the intensity of the sensationof taste were recorded by the TI system, and in addition, psychologicalmood states were evaluated with POMS (Profile of Mood State).TI evaluation showed that after the mental tasks, the perceivedduration of bitter, sour and sweet taste sensations was shortenedrelative to the control. Total amount of bitterness, sournessand sweetness was also significantly reduced. Furthermore, themaximum intensity of bitterness was significantly reduced. Therewere no significant differences in bitterness and sweetnesssensations following physical tasks. However, relative to beforethe physical task, the duration of the after-taste of sournesswas significantly shortened by the physical task. After thephysical task, the buffering capacity of saliva was significantlyincreased. Thus mental and physical tasks alter taste perceptionin different ways; the mechanisms underlying these changes remainto be determined. Chem. Senses 21: 195–200, 1996.