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On the taste of umami in chimpanzee.

Whole and single fiber chorda tympani nerve recordings were obtained in 5 chimpanzees to stimulation with MSG (monosodium phosphate) and GMP (guanosine 5'-monophosphate, disodium salt) alone and in combination. The overall chorda tympani nerve activity was recorded to 5 concentrations of MSG, ranging from 1 to 100 mM with and without 0.3 mM GMP, and to 5 concentrations of GMP, ranging from 0.1 to 10 mM, with and without 30 mM MSG. A synergistic effect was recorded between MSG and GMP in 3 out of 4 animals. The effect of stimulation with MSG and GMP alone and mixed was studied in approximately 25 single fiber recordings against a background of the stimulating effects of 11 different sweeteners, 3 acids, 3 bitter compounds and 3 different salts. The fibers showed a high taste specificity and fell into groups which corroborated with the human concepts of the taste qualities. The umami compounds elicited moderate responses which were largest in the sweet fibers. In the 6 sweet fibers that responded to the umami compounds. 0.3 mM GMP was a more effective stimulus than 10 mM MSG. In 3 of these fibers a synergistic effect was recorded to the mixture of GMP and MSG. It is interesting that the response to GMP and MSG was unaffected by gymnemic acid, although it blocked the response to the sweet compounds. Three out of 10 salt fibers responded to MSG and GMP but no synergistic effect was recorded. No specific umami fibers were recorded. However, more data must be collected before the final conclusion on the presence or absence of specific umami fibers can be drawn.

Animals↗

Neuronal responses of the nucleus tractus solitarius to oral stimulation with umami substances.

In order to investigate coding mechanisms of special taste modality (umami), responses of neurons within the nucleus tractus solitarius (NTS) to oral stimulation with monosodium glutamate, disodium 5'-inosinate (IMP) or their mixture were recorded in the conventional electrophysiological method. Results obtained were as follows: Neither MSG-best nor IMP-best neuron was recorded within the NTS as in the primary taste afferents. Some of the sucrose-best neurons, NaCl-best neurons and HCl-best neurons responded to oral stimulation with MSG or IMP. A remarkable synergistic effect was observed in all of the sucrose-best neurons and in some of the NaCl-best neurons but not in all of the HCl-best neurons, when the mixed solution of MSG and IMP was applied into the oral cavity. As to the sucrose-best neurons, potency of the synergism was positively correlated with the responsiveness to sucrose. No correlation was recognized between them in the case of NaCl-best neurons. These results suggest a view that the sucrose-best neurons and the NaCl-best neurons which show the synergism may participate in coding umami taste.

Afferent Pathways↗

Taste and olfactory modulation of feeding related neurons in behaving monkey.

Single neuron activity in the monkey lateral hypothalamus (LHA) was recorded by multibarreled electrode during a bar press feeding task. Activity of glucose-sensitive (GS) neurons decreased during bar press (BP) and reward (RW) periods. The inhibition was caused by activation of beta-adrenoceptors and opioid receptors respectively. Glucose-insensitive (GIS) neurons were excited during BP and RW, and at cue light (CL). Excitation at CL and BP was caused by activation of dopaminergic receptors. Among GS neurons, 66% responded to taste and 88% to odor. These responses were 39% and 52% in GIS neurons. GS neurons responded predominantly to two or more taste and odor stimuli while GIS neurons responded to only one stimulant. GS neurons have dense mutual connections with the prefrontal area, and GIS neurons are connected with the motor area. Gustatory and olfactory stimulation elicited responses in 67% of GS neurons and in only 21% of GIS neurons. Data suggest that GS and GIS neurons may have different functions in feeding: GS neurons process endogenous chemical information and integrated chemical sensations, and GIS neurons process external information processing, motor control and discriminative chemical sensations.

Animals↗

Parabrachial gustatory neural responses to monosodium glutamate ingested by awake rats.

A sample of 41 gustatory neurons isolated in the parabrachial nuclei of awake, behaving rats was tested with sapid solutions of 0.1 M monosodium glutamate (MSG), 0.5 mM of guanosine 5'-monophosphate (GMP), and a mixture of MSG and GMP as well as with 0.3 M sucrose, 0.1 M NaCl, 0.01 M citric acid, and 0.0001 M QHCl. Interneuronal correlation coefficients and factor analysis indicated that both the sodium cation and glutamic anion contributed to the activity elicited by MSG. Guanosine potentiated the responses to MSG, but only in neurons that also responded to sucrose. These results suggest that the gustatory contribution to the flavor denoted by the Japanese word "umami" may be mediated, in part, by neurons that also respond to chemical described by humans as sweet.

Animals↗

Responses of neurons in the primate taste cortex to glutamate.

In order to investigate the neural encoding of glutamate in the primate, recordings were made from 190 taste responsive neurons in the primary taste cortex and adjoining orbitofrontal cortex taste area in macaques. Single neurons were found that were tuned to respond best to glutamate (umami taste), just as other cells were found with best responses to glucose (sweet), sodium chloride (salty), HCl (sour), and quinine HCl (bitter). Across the population of neurons, the responsiveness to glutamate was poorly correlated with the responsiveness to NaCl, so that the representation of glutamate was clearly different from that of NaCl. Further, the representation of glutamate was shown to be approximately as different from each of the other four tastants as they are from each other, as shown by multidimensional scaling and cluster analysis. Moreover, it was found that glutamate is approximately as well represented in terms of mean evoked neural activity and the number of cells with best responses to it as the other four stimuli, glucose, NaCl, HCl and quinine. It is concluded that in primate taste cortical areas, glutamate, which produces umami taste in humans, is approximately as well represented as are the tastes produced by: glucose (sweet), NaCl (salty), HCl (sour) and quinine HCl (sour).

Animals↗

Taste preference and protein nutrition and L-amino acid homeostasis in male Sprague-Dawley rats.

Changes in taste preference were investigated in a choice paradigm using rats under various states of protein nutrition. A preference for the umami taste substances, monosodium L-glutamate (MSG) with or without 5'-ribonucleotide (5'-guanosine monophosphate), was induced when dietary protein was within the normal range, but a preference for NaCl occurred under its marginal deficiency. A preference for both NaCl and glycine was induced under severe protein malnutrition, possibly reflecting the body's negative nitrogen balance. The strength of this preference paralleled the body's requirement for protein, and thus, like the protein requirement for normal growth, declined with age. When animals with L-lysine (Lys) deficiency consumed a Lys solution and began to grow normally, their intake of taste stimulus solutions changed from preferring NaCl and glycine to preferring MSG. The regulatory mechanism of preference for L-amino acid (AA) in rats deficient of an essential AA was related to the pattern of AA in plasma and brain. Data suggest that umami taste perception plays an important role in protein metabolism, and in maintenance of AA and ammonia homeostasis within normal limits.

Amino Acids↗

Human development and umami taste.

Previously we demonstrated that in very brief, one-bottle intake tests, human infants, both well nourished and protein calorie malnourished, ingested greater amounts of soup with added MSG compared with soup alone. The potentiating effect of MSG in plain aqueous solution was tested in the current experiments. In contrast to data with sucrose and salt solution, where infants preferentially ingest these compounds compared with a water diluent, aqueous MSG solutions were rejected relative to water. The implications of this observation for understanding the mechanisms of MSG perception and pleasantness were discussed.

Child Development↗

Temporal characteristics of capsaicin sensitization and desensitization on the tongue.

Previous work has demonstrated that repeated applications of capsaicin can sensitize the tongue to subsequent exposures to capsaicin. However, if stimulation is interrupted for a period of minutes, the contrasting phenomenon of desensitization occurs. The purpose of the present study was to investigate the effect of interstimulus interval (ISI) on the development of both sensitization and desensitization to sequential capsaicin stimuli. It was determined in the first experiment that the rate of sensitization varied inversely with ISI for intervals briefer than 3.5 min; when ISI was lengthened to 5.5 min, sensitization was replaced by a trend toward desensitization. A second experiment established that increasing ISI from 9.5 min to 14.5 min tended to reduce rather than to enhance desensitization. It was therefore concluded that the sensitization process can be optimized by decreasing the time between successive stimuli (ISIs less than 3.5 min), whereas the trend toward desensitization appears to be optimal when stimuli are separated by at least 5.5 min but by less than 14.5 min.

Adult↗

Magnitude estimation of adaptation to salt using a flow chamber for stimulus delivery.

Adaptation course to NaCl was estimated using a flow chamber for stimulus delivery and changes in the perceived taste magnitude as the criterion. The adapting stimulus of different durations was applied to one side of the tongue. Magnitude estimates were made by comparing the intensity of taste at the end of each adaptation time with the perceived intensity of the same stimulus applied briefly to the unadapted side of the tongue. The course of adaptation followed a negatively accelerated decreasing function. The relationship between taste magnitude and the duration of the adapting stimulus can be approximated by a logarithmic equation and, somewhat less accurately, also by an exponential equation, which can be related to the Beidler theory of taste stimulation. Some advantages of the closed flow technique and comparison of taste magnitudes on the adapted and unadapted sides of the tongue are discussed.

Chorda Tympani Nerve↗

Capsaicin desensitization and recovery on the human tongue.

The desensitization resulting from application of 10 or 100 ppm capsaicin was investigated, using daily testing of a capsaicin series (1-1000 ppm, in log steps). The series showed a significant decrement in perceived burn following desensitization with either concentration. Perceived burn of 100 and 1000 ppm did not recover from 100 ppm desensitization in six days, and perceived burn of 1-1000 ppm did not recover from 100 ppm desensitization in six days. When single capsaicin concentrations, rather than the series, were tested at one, two, four, or six days after desensitization, 10 ppm recovered from 10 ppm desensitization in one or two days, and 100 ppm recovered from 100 ppm desensitization between two and four days. This suggests that daily testing with the capsaicin series delayed recovery from desensitization. Nontasters of 6-n-propylthiouracil rated capsaicin burn lower than did tasters. The application method of rolling capsaicin onto the tongue with a swab was found to transiently inhibit burn. Implications for ingesting capsaicin products are discussed.

Capsaicin↗

Amiloride reduces intensity responses of human fungiform papillae.

Microdrop stimulation of human fungiform papillae was used to confirm that the drug amiloride reportedly reduces the taste response to NaCl and sucrose. Amiloride caused magnitude reductions of saltiness in most subjects and of sweetness for some but not all subjects. More of the subjects showed a reduction in response to NaCl than a reduction to sucrose and there were individual differences in the strength of the reduction of magnitude.

Adult↗

Human psychophysical studies of saltiness suppression by amiloride.

Two studies were performed to assess the effects of amiloride on the saltiness of NaCl. In the first study, four highly trained subjects were asked to report the first appearance of saltiness as NaCl microdrops were rapidly applied to an increasing number of fungiform papillae along the front, dorsal surface of the tongue. For two subjects, amiloride, a sodium channel blocker, caused a large and significant increase in the number of papillae needed to report a salty taste. The other two subjects were unaffected, as measured by this procedure. In the second study, four different subjects were trained in the same procedure as in the previous study. For testing, however, they were stimulated on a fixed number of papillae, which was two times the number self-selected during training as sufficient for perception of saltiness with NaCl, and sourness with citric acid. They reported the quality of the experience as salty, sour, or neither, with and without amiloride treatment of the papillae set. Two of the four subjects had a loss of saltiness of NaCl, but not sourness of citric acid, when the papillae set was treated with amiloride.

Adult↗

Chorda tympani and lingual nerve responses to astringent compounds in rodents.

A wide variety of compounds in foods and beverages produce astringent sensations when introduced into the oral cavity. There is controversy, however, whether "astringency," with its associated puckering and drying sensations, is a fundamental taste quality or is a tactile sensation. To address this issue, electrophysiological recordings were made from the gerbil chorda tympani nerve and the rat lingual nerve. The chorda tympani nerve transmits taste information from the anterior 2/3 of the tongue, whereas the lingual nerve transmits tactile, thermal and pain sensations from the anterior 2/3 of the tongue. The astringent compounds tested were: tannic acid, tartaric acid, gallic acid, aluminum ammonium sulfate and aluminum potassium sulfate. Tannic acid, tartaric acid, and gallic acids were tested at concentrations up to 120 mM over a pH range from approximately 2 to 6. The aluminum salts were tested at concentrations up to 160 mM only at low pH's. All compounds rapidly (and at lower concentrations, reversibly) stimulate the chorda tympani nerve in a concentration-dependent manner at all pH's tested. The rapidity and reversibility of the chorda tympani responses suggest that astringent-tasting compounds interact directly with taste cells rather than indirectly by precipitating salivary proteins. At pH 6, tannic acid, tartaric acid, and gallic acid all elicit robust chorda tympani responses, implying that the ionized forms of these compounds produce taste sensations. None of these compounds stimulate lingual nerves over the same concentration and pH ranges used in the chorda tympani experiments.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Taste prestimulation increases the chorda tympani nerve response to menthol.

Electrophysiological recordings of the summated response of the chorda tympani nerve to menthol stimulation of the tongue were obtained from 15 adult Sprague-Dawley rats. The chorda tympani nerve response to menthol was of short duration, ending within 2.5 s after stimulus onset, leaving the receptors in a state of insensitivity to subsequent menthol stimulation. Rinse durations with deionized-distilled water up to 10 min failed to bring the receptors back to their original prestimulus state. Although stimulation with menthol prevented taste receptors from responding to subsequent presentations of menthol, the chorda tympani nerve would respond normally to NaCl, NH4Cl, KCl, sodium acetate, glucose, citric acid, and quinine-HCl solutions. Prior stimulation with one of these taste solutions resulted in the recovery of the menthol response. The magnitude of the recovered menthol response depended on the magnitude of the phasic response elicited by the preceding taste stimulus. A general explanation involving possible reception and transduction mechanisms was offered to account for menthol's unexpected stimulatory effects on the chorda tympani nerve.

Animals↗

The effect of bilateral sectioning of the chorda tympani and glossopharyngeal nerves on the sweet taste in the mouse.

The chorda tympani nerve (CT) and the glossopharyngeal nerve (GL) have been considered important nerves for the sense of taste. We studied the effect of bilateral sectioning of the CT and/or GL on the sweet taste in the mouse. Before and after surgery we analyzed the daily drinking patterns, using the two-bottle preference test method. The normal mouse drank the low concentration sucrose solution (0.0125 M) more than distilled water. This report showed that the mouse who was bilateral sectioned, both CT and GL or bilateral sectioned CT, rejected drinking the low concentration of sucrose solution. In contrast, the mouse who was bilateral sectioned GL drank the low concentration sucrose solution like the normal mouse did. These phenomena suggested that the fungiform papilae play an important role to detect the low concentration of sucrose (0.0125 M) as a sweet favorable taste substance.

Animals↗

Development of some early sensorimotor behaviors in sodium-restricted rats.

Rats exposed to low levels of dietary sodium throughout development exhibit reduced chorda tympani nerve taste responses to sodium stimuli during adulthood, apparently due to altered activity of some hormone(s) or growth factor(s) during early development. We were concerned that such an alternation in the activity of some humoral factor(s) could affect development globally. To test this possibility, we utilized a battery of morphological and behavioral measures in neonatal, sodium-restricted rats, expecting serious deficits to be reflected in altered onset and expression of these behaviors. As compared with control rat pups, preweanling sodium-restricted rat pups exhibited greatly diminished body weight gain and delayed acquisition of several morphological features. However, in terms of sensorimotor development, no significant differences between sodium-restricted and control rat pups were found. We interpret these results to indicate that despite significant somatic effects, sodium restriction may not influence the development of physical prowess or of early sensorimotor function in a global manner.

Animals↗

Altered gustatory development in Na(+)-restricted rats is not explained by low Na+ levels in mothers' milk.

Placing pregnant rats on a sodium-deficient diet (0.03% NaCl) very early in gestation and then weaning the offspring (sodium-restricted rats) to the same diet precludes development of amiloride-sensitive sodium taste transduction pathways in the offspring. However, normal amiloride-sensitive sodium taste responses can be restored by permitting sodium ingestion by sodium-restricted rats. The present study tested the hypothesis that the concentration of sodium in sodium-restricted mothers' milk must be abnormally low in order to preserve altered gustatory function in the offspring. Other milk electrolyte and total protein concentrations were determined as well. Milk sodium was similar between sodium-restricted and control rat mothers at 10-13 and 16-20 days postpartum, as were levels of potassium and chloride. At 10-13 days postpartum, total protein was higher in milk from sodium-restricted mothers. Sodium-restricted mothers' milk calcium concentrations were higher versus controls at 16-20 days postpartum. These results indicate that the lack of gustatory amiloride sensitivity in sodium-restricted rats cannot be attributed to deficient dietary sodium levels during the suckling period.

Amiloride↗

Central processes in gustatory adaptation.

In two experiments the adapting stimuli at the periphery of the taste system were of a constant objective intensity but, using mixture suppression and spatial summation, their perceived intensity was varied. The results have shown that, in spite of the constancy in stimulus concentration, the adaptation degree changed with the perceived intensity of the stimulus. The adaptation to the compounds when in binary mixtures proved to be significantly less effective than the adaptation to the same equimolar compounds when unmixed. Similarly, the adaptation effects of solutions of a constant concentration, when applied to a small tongue area, were significantly smaller than when applied to a larger area. This adaptation dependence upon the taste sensation intensity suggests that in the gustatory adaptation, peripheral as well as central processes take part.

Attention↗