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The discovery of two families of mammalian taste receptors has provided important insights into taste recognition and taste perception. Recent studies have examined the receptors and signaling pathways that mediate sweet, bitter, and amino acid taste detection in mammals. These studies demonstrate that taste cells are selectively tuned to different taste modalities and clarify the logic of taste coding in the periphery.
BACKGROUND: Besides anamnestic data and taste test, the clarification of epithelial taste disorders makes macroscopic evaluation of the surface of the tongue necessary. METHODS: In this article the contact endoscopy is evaluated for examination of the tongue epithelium and for quantification of the fungiforme papillae. These results are related to the results of the spatial taste test (1. chemical taste test with sodium chloride in 2 different suprathreshold concentrations and sucrose in also 2 different suprathreshold concentrations; 2. electrogustometry with a bipolar electrode). PATIENTS: In a prospective study we performed contact endoscopy in 16 healthy volunteers from ages 7 to 68 years. The subjects were divided into 2 age groups (< or = 45 years and > 45 years) and 2 papillae density groups. In 34 patients with either taste disorders of unknown cause or in patients with macroscopic changes at the tongue surface, the spatial and whole mouth taste tests and contact endoscopy were used to clarify the cause of the complaints. RESULTS: Density and shape of the fungiform papillae as well as the shape of the subepithelial vessels of the papillae vary among the investigated human subjects in the different age groups. No correlation between intensity ratings, papillae density und age was found after stimulation with 40 % sucrose at the tongue tip in the younger age group. Stimulation of the tongue edge with 15 % sodium chloride, however, showed a relationship between the two different density groups and the different age groups. CONCLUSIONS: The contact endoscopy allows the evaluation of the epithelium in the oral cavity. It is a nondestructive method and is additionally easy to handle, but the evaluation of the observed epithelia needs interdisciplinary co-operation with pathologists. No relevant informations are supplied in diagnostics of taste disorders.
Taste detection and recognition thresholds were monitored in 10 patients for up to 1 yr after allogeneic bone marrow transplantation (BMT). As well as a control group of 12 healthy volunteers, taste acuity was tested in 10 patients, who had undergone BMT 2-5 yr previously. Immediately after transplantation, there was significant hypogeusia of all four taste modalities compared to registrations one week before the aplastic phase and also compared to the healthy control group. Although some normalization of taste thresholds was registered 3-6 months after transplantation, most subjects still experienced dysgeusia. Of the four taste modalities, the most frequently recorded change was a raised threshold for salt. In about 80% of the patients taste acuity had recovered to the control values one year after transplantation. The group tested 2-5 yr after BMT had normal values for taste acuity.
Recent studies showing that single taste bud cells express multiple bitter taste receptors have reignited a long-standing controversy over whether single gustatory receptor cells respond selectively or broadly to tastants. We examined calcium responses of rat taste receptor cells in situ to a panel of bitter compounds to determine whether individual cells distinguish between bitter stimuli. Most bitter-responsive taste cells were activated by only one out of five compounds tested. In taste cells that responded to multiple stimuli, there were no significant associations between any two stimuli. Bitter sensation does not appear to occur through the activation of a homogeneous population of broadly tuned bitter-sensitive taste cells. Instead, different bitter stimuli may activate different subpopulations of bitter-sensitive taste cells.
1. The responses of 126 neurons in primary gustatory cortices of two rhesus monkeys were recorded during sapid stimulation of the tongue with 18 taste stimuli. Ten of these stimuli were dissolved in distilled water (DW): 1.0 M sucrose (Suc), 0.1 M and 0.03 M sodium chloride (NaCl), 0.003 M hydrochloric acid (HCl), 0.001 M quinine hydrochloride (QHCl), 0.03 M monosodium glutamate (MSG), 0.03 M polycose, 0.3 M glycine, 0.1 M proline, and 0.1 M malic acid. Seven other stimuli were dissolved in 0.03 M MSG; the last stimulus was a mixture of 1.0 M Suc and 0.03 M NaCl. 2. The average spontaneous rate (2.2 +/- 0.2 spikes/s, mean +/- SE) and response to DW (2.5 +/- 0.2) of these 126 neurons was low but within the range previously reported for neurons in primate taste cortex. Suc was the most effective stimulus for 24.1% of the neurons tested followed by NaCl (15.7%), QHCl (14.8%), HCl (11.1%), MSG (10.2%), and other miscellaneous unitary gustatory stimuli (8.3%). Binary taste mixtures were the most effective stimuli for 15.7% of the sample. The net responses (corrected for DW, in spikes/s) for Suc-best (3.3), NaCl-best (4.3), HCl-best (3.4), QHCl-best (2.3), and MSG-best (4.1) were sluggish, but comparable with that reported previously. 3. The response breadth of the 82 neurons that responded best to either Suc, NaCl, HCl, or QHCl measured with the entropy coefficient indicated a moderate response breadth for these neurons (mean = 0.79; range = 0.30-0.98). According to the response criteria adopted in this experiment (water response +/- 1.96 SD), however, 81 of these 82 neurons (98.1%) responded to only one or two of the four basic taste stimuli. The disparity between the entropy- and criterion-based measures of response derive from the nature of the two statistics. Adjustments that would make the entropy statistic less inclusive and the definition of a response according to statistical criteria less exclusive would increase their concordance. 4. Three multivariate statistics (cluster, principal axis factor, and multidimensional analysis) were used to analyze the data. Cluster analysis enabled us to divide the 82 taste neurons into groups on the basis of response similarity. Each of the four largest groups was dominated by neurons that responded best to one of the four basic taste stimuli: Suc, NaCl, QHCl, and HCl (ranked in descending order); the fifth largest cluster contained neurons that responded best to MSG. Principal axis factor analysis demonstrated that 80.8% of the total variance could be accounted for by three factors. Neurons responding best to Suc, NaCl, and QHCl each were closely associated with one of those three factors, but the loadings of the HCl-best neurons were evenly distributed across all three factors. The communality coefficient of these three factors was > 80% for the Suc-, NaCl-, HCl-, and QHCl-best neurons; the MSG-best neurons, by comparison, had very few high loadings on any of these three factors and a correspondingly low communality coefficient of 40.4%, a difference that was statistically significant from the other four groups. Thus the three factors related to Suc-, NaCl-, HCl-, and QHCl-best neurons are not relevant to MSG-best neurons. We used multidimensional analysis to arrange the neurons that responded best to Suc, NaCl, HCl, QHCl, and MSG into five loosely arranged and partially overlapping clusters. A multidimensional space based on stimulus similarity showed that MSG was as different from the four basic taste stimuli as they were from one another. 5. Mixture suppression, a common observation in human psychophysical experiments, was examined at the neurophysiological level by including binary tastants in the stimulus battery. The average response of 19 Suc-best neurons to 1.0 M Suc (4.1 spikes/s) decreased to near 0 when the solvent was changed from DW to either 0.03 M MSG or 0.03 M NaCl. Similar decrements were observed in NaCl- and MSG-best neurons tested with Suc/NaCl mixtures.
BACKGROUND: The sweet taste inhibitor lactisole acts on the human sweet taste receptor heteromer TAS1R2-TAS1R3 but not on its rodent counterpart. Recently, it was shown that the lactisole sensitivity of the human sweet taste receptor involves the part of TAS1R3 encompassing the seven transmembrane regions but not the huge N-terminal domain. Using mutational analysis we investigated which amino acid residues distinguish lactisole insensitive rat from sensitive human T1R3 receptors. RESULTS: The functional analysis of specific receptor mutants in HEK293T cells revealed that the exchange of valine 738 in the fifth transmembrane domain of rTas1r3 by an alanine is sufficient to confer lactisole sensitivity to the rat sweet taste receptor. The sensitivity of this receptor mutant is approximately 2 fold lower than the sensitivity of the human sweet taste receptor. Additional substitution of lysine 735 by phenylalanine in rTas1r3 results in a rat sweet taste receptor that is as sensitive to lactisole as its human counterpart. The exchange of valine 738 to alanine was accompanied by a approximately 50% reduction in receptor efficacy. This effect was seen with all six different sweet compounds examined. CONCLUSION: The lactisole insensitivity of rat sweet taste receptor is caused by only two amino acids in transmembrane region five, which is critical for the interaction of lactisole with the sweet taste receptor. The observation that the mutant receptor simultaneously displays a generally reduced sensitivity towards all agonists suggests that the lactisole insensitivity of the rodent receptor might be more likely caused by the inaccessibility of the lactisole binding site rather then by its direct disruption.
A 38-year-old man developed dysesthesia, diplopia, and an unsteady gait following an upper respiratory infection. IgG anti-GQ1b antibody was detected in his serum and he was diagnosed as Fisher syndrome. The patient also complained of loss of taste sensation, and it resolved along with improvement of other neurological manifestations. In Guillain-Barré syndrome, cranial nerve involvement is very common, though taste disturbance is a rare complaint. Impairment of taste has been reported in association with severe facial nerve involvement, but taste disturbance developed without facial nerve palsy in the present case and taste sensation was diminished in the area of all four nerves involved in taste sensation. These findings suggest that the impaired taste sensation in the present patient was not a complication of facial nerve palsy as in previous cases, but rather due to taste sensory specific involvement.
In various environments where primates are presently observed, as well as in forests and savannas which have been inhabited by australopithecines and early hominids, there are (or there have been presumably) categories of substances eliciting taste signals associated with stereotyped responses. Such is the case for various soluble sugars of fruits and nectars, attracting consumers, and for several plant compounds in which bitter or strongly astringent properties have a repulsive effect. The occurrence of such classes of tasty substances among natural products appears to be related to the evolutionary trends that shaped primate sensory perception (for detecting either beneficent or potentially noxious substances) in the context of a long history of coevolution between animals and plants. Here, we present original psychophysical data on humans (412 individuals aged 17-59 years) as an analogy with which to test recent evidence from electrophysiology in nonhuman primates (Hellekant et al. [1997] J. Neurophysiol. 77:978-993; Danilova et al. [1998] Ann. N.Y. Acad. Sci. 855:160-164) that taste fibers can be grouped into clusters of "best-responding fibers" with two more specific clusters, one for sugars and one for quinine and tannins. The collinearity found between human taste responses (recognition thresholds) for fructose and sucrose, as well as for quinine and tannins, is presented and discussed as another evidence of the two-direction evolutionary trend determining taste sensitivity. Salt perception appears to be totally independent of these trends. Accordingly, the appreciation of a salty taste seems to be a recent culturally learned response, and not a primary taste perception. The very existence of primary tastes is discussed in the context of evolutionary trends, past and present.
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 carbohydrates 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.
Effects of cytochalasin D on actin filaments in cells encircling taste pores were examined to clarify the functional role of actin filaments in the maintenance of taste pores in rat fungiform papillae, using a confocal laser microscope and a scanning electron microscope. Fluorescence in the taste pore cells was detected as a ring shape produced by actin staining with rhodamine-phalloidin. Treatment of fungiform papillae with cytochalasin D diminished the positive reactions in the taste pore cells and increased the inner diameter of the ring reactions. However, deformation of the taste pores in fungiform papillae was not detected under a scanning electron microscope after treatment with cytochalasin D. These findings suggest that the organization of actin filaments encircling the taste pores contributes to regulation of the taste pore's size in rat fungiform papillae.
Underground water in volcanic areas contains vanadium when the basalt layer exists among igneous rocks. The concentration of vanadium in drinking water sometimes exceeds 0.8 microM in these areas, however, the physiological effects of vanadium, especially non-toxic effects, at concentrations lower than 1 microM are unknown. In the present experiments, we examined the effect of pentavalent vanadium and tetravalent vanadium at 0.8 and 8.0 microM concentrations on the recognition threshold to taste substances in healthy college students. Pentavalent vanadium, ammonium vanadate, lowered the sweet taste threshold to glucose at 0.8 and 8.0 microM as well. Tetravalent vanadium, vanadium sulfate, did not alter the threshold to glucose either at 8.0 microM or at 0.8 microM. Ammonium vanadate also decreased the sweet taste threshold to L-proline at 8.0 microM. Ammonium vanadate did not influence the sour taste threshold to hydrogen chloride. Neither ammonium sulfate nor ammonium bicarbonate altered the sweet taste threshold to glucose. Therefore, the effect of ammonium vanadate on the sweet taste threshold is attained by vanadium but not by ammonium. It was concluded that pentavalent vanadium at 0.8 microM intensifies the sweet taste sense to glucose rather specifically. We have first shown the physiological effect of vanadium at the concentration of the underground water level.
Taste and smell functions were measured in 18 unselected patients with untreated primary hypothyroidism, and in 15 of the 18 patients after treatment with thyroid hormones. Before treatment, 9 of the 18 patients (50 per cent) were aware of some alteration in their sense of taste, and 7 of the 18 patients (39 per cent) were aware of some alteration in their sense of smell. Distoritions of tase (dysgeusia) and smell (dysosmia) were frequent complaints among the untreated patients; dysgeusia was observed by 7 patients (39 per cent) and dysosmia by 3 patients (17 per cent). Median detection and recognition thresholds for four taste stimuli salt (sodium chloride), sweet (sucrose), sour (hydrochloric acid) and bitter (urea), and for two smell stimuli (pyridine and nitrobenzene), were determined in each patient before and after treatment with thyroid hormones. Before treatment, decreased taste acuity (hypogeusia) for at least one stimulus was observed in 14 of the patients (83 per cent); the most common abnormalities were in the detection and recognition of bitter stimuli. Median detection thresholds for both smell stimuli were also markedly elevated (hyposmia) before therapy. Treatment with throid hormones largely reversed both the taste and smell defects. In one patient, taste and smell abnormalities were completely corrected after 16 days of treatment with thyroxine. This study indicates that taste and smell defects are common clinical abnormalities in primary hypothyroidism, and suggests that these defects may contribute to the anorexia and lack of interest in eating which are frequently observed.
To determine whether functional characteristics of the taste system change during development, electrophysiological taste responses were recorded from neurons in the solitary complex (nucleus and tractus solitarius) in the medulla of fetal, newborn and adult sheep. Taste stimuli included NH4Cl, KCl, NaCl, LiCl, citric acid, and HCl, applied to the anterior tongue. Fetal neurons at all ages (84-137 days of gestation) responded to stimulation of the tongue with NH4Cl and KCl, but responses to NaCl and LiCl were only obtained in older fetuses (after 114 days of gestation), lambs and adults. Responses to citric aicd were obtained at all ages; however, HCl responses were only infrequently obtained in young fetuses. Other developmental changes included a progressive decrease in latency of the responses to NH4Cl, KCl, citric acid and HCl, and an increase in the duration of the neural response discharge as a function of gestational age. Since taste buds do not acquire the structural characteristics of the adult until the last third of gestation (approximately 100-147 days), these functional changes in taste response characteristics take place concurrently with structural development. Mammalian fetuses swallow amniotic fluid in utero, and, therefore, the fetal taste system is stimulated during structural and functional development. Thus, there is an opportunity for fetal gustatory experience to influence the developing taste system.
The examples of "synergism" reported in the taste literature fit the common definition of the term but may merely reflect normal additivity in the taste system. That is, the perceived intensities of these "synergistic" mixtures exceed the sums of the perceived intensities of the unmixed taste components (addition of perceived intensities), but do not exceed the perceived intensities predicted by adding along the expandng psychophysical functions which describe the unmixed taste components (stimulus addition). True synergism requires that the perceived intensity of a mixture be greater than that predicted by both types of additivity. Subjects used magnitude estimation to judge the perceived intensity of unmixed monosodium glutamate and disodium 5'-guanylate and mixtures of these two taste substances. The mixtures showed synergism according to both crtieria. The robustness of this phenomenon is striking; for all subjects the taste mixtures showed true synergism. True synergism in taste is an extremely rare phenomenon.
Taste receptor cells are primary sensory receptors utilized by the nervous system to detect the presence of gustatory stimuli in the oral cavity. These cells are particularly heterogeneous and may be divided into various subtypes based on morphological, histochemical, or physiological criteria. One example is the heterogeneous expression of neuropeptides, such as cholecystokinin and vasoactive intestinal polypeptide. These peptides are hypothesized to participate in the transduction processes. To pursue examination of this hypothesis, this study explored the relationship of peptide expression with two important and mostly non-overlapping transductive elements--the taste-specific G protein gustducin, involved in bitter and sweet transduction cascades, and the seven transmembrane taste receptor T1R2, hypothesized to respond to sweet compounds. Double labeling experiments were performed on taste buds of the posterior rat tongue combining immunocytochemistry for peptide expression and in situ hybridization experiments for either gustducin or T1R2 expression. Additionally, vasoactive intestinal peptide (VIP)-expression in posterior taste receptor cells was confirmed using the technique of RT-PCR. More than half (56%) of the CCK-expressing taste receptor cells co-expressed alpha-gustducin mRNA whereas far fewer (15%) co-expressed T1R2 mRNA. A majority of VIP-expressing taste receptor cells co-expressed alpha-gustducin mRNA (60%) whereas only 19% of these cells co-expressed T1R2 mRNA. More remarkable was the observation that these two peptides displayed almost identical expression patterns with these signal transduction molecules, suggesting that peptides are not randomly expressed with relation to signal transduction molecules. This observation supports the hypothesis that peptides may play roles in transduction. Further physiological exploration will be required to elucidate the nature of these roles.
There is overlap between brain regions involved in taste and pain perception, and cortical injuries may lead to increases as well as decreases in sensitivity to taste. Recently it was shown that chronic back pain (CBP) is associated with a specific pattern of brain atrophy. Since CBP is characterized by increased sensitivity to pain, we reasoned that the sense of taste might also be enhanced in CBP. Detection and recognition thresholds were established for a sour taste and ratings of both suprathreshold taste intensity and pleasantness-unpleasantness perception were collected for sweet, sour, salty and bitter stimuli in 11 CBP patients and 11 matched control subjects. As a control, ratings were also collected for visual assessment of degree of grayness. There was no difference between CBP and control subjects for visual grayness rating. On the other hand, CBP patients in comparison to control subjects rated gustatory stimuli as significantly more intense but no more or less pleasant and showed a trend towards a lower detection threshold (i.e. increased sensitivity). The selectivity of the taste disturbance suggests interaction between pain and taste at specific brain sites and provides further evidence that CBP involves specific brain abnormalities.
Elements expressing the molecular mechanisms of gustatory transduction have been described in several organs in the digestive and respiratory apparatuses. These taste cell-related elements are isolated cells, which are not grouped in buds, and they have been interpreted as chemoreceptors. Their presence in epithelia of endodermal origin suggests the existence of a diffuse chemosensory system (DCS) sharing common signaling mechanisms with the "classic" taste organs. The elements of this taste cell-related DCS display a site-related morphologic polymorphism, and in the past they have been indicated with various names (e.g., brush, tuft, caveolated, fibrillo-vesicular or solitary chemosensory cells). It may be that the taste cell-related DCS is like an iceberg: the taste buds are probably only the most visible portion, with most of the iceberg more caudally located in the form of solitary chemosensory cells or chemosensory clusters. Comparative anatomical studies in lower vertebrates suggest that this 'submerged' portion may represent the most phylogenetically ancient component of the system, which is probably involved in defensive or digestive mechanisms. In the taste buds, the presence of several cell subtypes and of a wide range of molecular mechanisms permits precise food analysis. The larger, 'submerged' portion of the iceberg is composed of a polymorphic population of isolated elements or cell clusters in which the molecular cascade of cell signaling needs to be explored in detail. The little data we have strongly suggests a close relationship with taste cells. Morphological and biochemical considerations suggest that the DCS is a potential new drug target. Modulation of the respiratory and digestive apparatuses through substances, which act on the molecular receptors of this chemoreceptive system, could be a new frontier in drug discovery.
In the gustatory system, the mechanism of coding-the process of relaying the identity and intensity of the stimulus to the central nervous system-begins with the taste receptor cells. Ironically, although these are the first cells of the gustatory system to contact stimuli, they were the last from which neurophysiological recordings were obtained. How taste receptor cells decipher stimulus identity remains the subject of active research; its origins began with a series of intracellular studies. Prior to the first intracellular recording, it was unknown if taste receptor cells would be specialists, responding to only a single class of taste stimulus, or generalists, responding to multiple stimuli. The first reports established several major aspects of these cells' physiology. Taste receptor cells have varying response profiles to basic stimuli; they have obvious conductance changes during stimulation; they have low resting potentials. It became evident that multiple transduction schemes must underlie these responses, although the identity of these transduction schemes remained elusive. Additionally, these early recordings missed a major phenomenon-the presence of regenerative electrical events (i.e., the action potential) was not observed due to the low input resistance that accompanied this technique. Although intracellular recordings are essentially no longer used to study taste receptor cells, replaced by the superior method of patch-clamp recording, these early articles provided key insights into the then unknown electrical responses of taste receptor cells.