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Sensory evaluations of fat-sucrose and fat-salt mixtures: relationship to age and weight status.

Young (mean age 22.4 years) and elderly (mean age 82.3 years) subjects tasted and rated a range of liquids containing between 0.5% and 36% dairy fat by weight. The liquids also contained either sucrose (0-20%) or NaCl (0-0.584%). Within each age group both normal weight and overweight subjects participated. The fat content in the mixtures was unrelated to pleasantness ratings for elderly subjects. The concentrations of both fat and sucrose in the mixtures determined the hedonic responses of young normal weight subjects. Overweight young subjects' hedonic responses were predominantly influenced by sucrose concentration. Fat content in dairy-salt mixtures was significantly related to pleasantness ratings by young subjects, but did not influence ratings by elderly subjects. These findings suggest that the elderly can significantly reduce dietary fat intake without sacrificing perceived pleasantness.

Adult↗

Developmental changes in sugar and starch taste preferences in young rats.

Recent studies indicate that neonatal and adult rats are attracted to the taste of sugar as well as to starch-derived polysaccharides (e.g., Polycose). The present study investigated whether the relative preference for sweet and starchy tastes changes during the postweaning to adulthood period in male and female rats. This was accomplished by giving growing rats 24-hr/day one-solution (sucrose or Polycose vs. water) and two-solution (sucrose vs. Polycose) preference tests over a 9-week period (from 28 to 98 days of age). Acceptance (absolute intake) of sucrose and Polycose was also evaluated by comparing solution intakes during one-solution tests with water baseline intakes. Dilute solutions were used to minimize postingestive caloric and/or colligative effects. When tested with 2% saccharide solutions, the rats strongly preferred sucrose to Polycose with the preference increasing in magnitude from weeks 1 to 9 of testing. Sucrose acceptance also increased, relative to that of Polycose, after the third test week. When tested with 0.06 molar solutions, the rats initially consumed more Polycose (week 1) but by the fifth test week their sucrose preference and acceptance exceeded that of Polycose. These results indicate that the preference for sweet taste, relative to that for starchy taste, increases with age. Contrary to previous findings, the sweet taste preference was somewhat stronger in the male rats than in the female rats.

Aging↗

Variations in human taste bud density and taste intensity perception.

Some variations in human taste sensitivity may be due to different numbers of taste buds among subjects. Taste pores were counted on the tongue tips of 16 people with videomicroscopy, and the subjects were divided into two groups (N = 8) by the rank order of their taste bud densities. The "higher" density group averaged 374 +/- 134 taste pores/cm2, while the "lower" density group averaged 135 +/- 43 tp/cm2. The higher density group had an average fungiform papilla density which was 1.8 times greater than the lower density group and an average of 1.5 times more taste pores/papilla. The subjects also rated the intensity for 4 suprathreshold concentrations of 5 taste stimuli placed on the same region of the tongue where taste pores were counted. The group with higher taste bud densities gave significantly higher average intensity ratings for sucrose (196%), NaCl (135%) and PROP (142%), but not for citric acid (118%) and quinine HCl (110%) than the lower density group. Thus, the subjects with higher fungiform taste bud densities also reported some tastes as more intense than subjects with fewer fungiform taste buds.

Arousal↗

Peripheral taste responses in genetically hypertensive rats.

In two-bottle preference-aversion tests, the spontaneously hypertensive rat (SHR) tolerates higher concentrations of NaCl than the normotensive Wistar-Kyoto (WKY). In contrast, the inbred Dahl salt-sensitive (S/JR) and inbred Dahl salt-resistant (R/JR) rat show similar preferences for NaCl. In order to determine if taste receptor function was also altered between the hypertensive rat and its normotensive control, we recorded electrophysiological taste responses from the chorda tympani (CT) nerve in SHR, WKY, S/JR and R/JR rats. Responses to a concentration series (0.05 M to 0.5 M) of NaCl, NaAcetate, KCl, NH4Cl and CaCl2 were recorded before and after lingual application of amiloride hydrochloride, an epithelial sodium transport blocker. When expressed relative to the 0.5 M NH4Cl response, responses to the majority of stimuli were equivalent between the SHR and WKY. By comparison, relative responses to NaCl were greater in the R/JR than S/JR; however, the magnitude of amiloride suppression was equivalent between these two strains. Relative responses to the majority of the remaining salts did not differ between the S/JR and R/JR. These results suggest that taste receptor function may be equivalent between the hypertensive rat and its normotensive control.

Animals↗

Quality-specific differences in rat taste detection performance as a function of stimulus volume.

Taste detection performance for representatives of the four taste qualities as a function of stimulus volume (5 x 10(-4) to 1 x 1(-1) ml) was examined in rats using high-precision gustometry, computer-controlled operant procedures, nonparametric signal detection measures of sensitivity and responsivity, and blind control procedures. The overall sensitivity index was positively related to stimulus volume (rs = .60), with optimal detection performance attained with a 5 x 10(-3) ml stimulus volume for salty tastants and a 1 x 10(-2) ml stimulus volume for the other taste qualities. The overall responsivity index was inversely related to stimulus volume (rs = -.47), especially for sour and bitter tastants. These results are consistent with prior observations and demonstrate that operant methods using small tastant samples produce sensitive estimates of the rat's taste detection performance and response bias.

Animals↗

Time-quality tracking of monosodium glutamate, sodium saccharin, and a citric acid-saccharin mixture.

The temporal patterns of taste-quality descriptors evoked by 1000-ms duration stimulus liquids flowed through a closed delivery system over the anterodorsal tongue tip region were indicated using touch-typing on a computer keyboard. Single keys corresponded to the taste words of a 23 item code. A computer monitor displayed for subjects the keys pressed and when they were pressed, starting at stimulus delivery. For 2 mM sodium saccharin (NaSac), 75% of the responses were "sweet," 6.5% "sugar"; for NaSac in 10 mM citric acid (ArtLem), 43% "sour," 20% "citrus," and 11% "sugar"; for 214 mM monosodium glutamate (MSG), 28% "salty," 14% "sour," and 10% 1st "soapy," then "no taste," and finally "bitter." Distilled water received "no taste" on all trials. Response durations were 657 ms for ArtLem, 594 ms for NaSac, 577 ms for MSG. MSG yielded multiple quality responses on 25.5% of the trials; ArtLem, 9%; and NaSac, 1%. These results are compared with temporal patterns for taste intensity and with unrestricted verbal descriptions of the solutions.

Adolescent↗

Canine taste nerve responses to umami substances.

The taste responses to the "umami substances" such as monosodium glutamate (MSG), guanosine 5'-monophosphate (GMP) and inosine 5'-monophosphate (IMP) were recorded from the canine chorda tympani nerve. A large synergism was observed between MSG and the nucleotides in most mongrel dogs (type A dog). The extent of the synergism between MSG and the nucleotides was much larger than that observed in any other animal examined except for humans. No synergism was observed between the nucleotide (GMP) and stimuli other than MSG, such as NaCl, HCl, sucrose, quinine, and glycine. It was concluded that the dog is a suitable experimental animal for studies on the responses to umami substances. In order to differentiate umami and salt components in the responses to umami substances, effects of amiloride on the responses were examined. Amiloride inhibited the response to MSG, but did not inhibit the response to GMP alone or those induced by synergism between GMP and MSG. The present results favor a conclusion that GMP acts as an agonist and MSG acts as a modulator for the umami receptor in the dog. The synergism can be explained by an allosteric model where the umami receptor is assumed to have two binding sites, one for GMP and another for MSG.

Amiloride↗

Structure/activity relationships in the L-alanine taste receptor system of the channel catfish, Ictalurus punctatus.

In order to understand the molecular determinants of amino acid taste receptor binding and activation, structure/activity studies were performed using analogs of L-alanine in a competitive ligand binding assay and a taste neurophysiological preparation. The presence of both the amine and carboxylic acid in a charged and unhindered form is a primary requisite for a ligand to both bind and activate L-alanine receptors. Although a number of carboxylic acid derivatives are moderately good stimuli, their neural activity derives from action at receptors different from L-alanine receptors. Of the molecular parameters examined, chirality and molecular volume of the side chain are the most important factors in determining the binding and stimulatory efficacy of L-amino acid taste stimuli. Electronegativity of the side chain did not correlate with receptor site binding. Heterologous ligand-induced enhancement of the binding of L-[3H]alanine by a purified taste membrane preparation is described. Neurophysiological experiments support the hypothesis that this phenomenon may be a basis of peripheral sensory interactions.

Alanine↗

Transduction mechanisms for the taste of amino acids.

Amino acids are important taste stimuli for a variety of animals. One animal model, the channel catfish, I. punctatus, possesses sensitive taste receptor systems for several amino acids. Neurophysiological and biochemical receptor binding studies suggest the presence of at least three receptor pathways: one is a relatively nonspecific site(s) responsive to short-chain neutral amino acids such as L-alanine (L-ALA); another is responsive to the basic amino acid L-arginine (L-ARG); still another is a low affinity site for L-proline (L-PRO). Several possible transduction pathways are available in the taste system of this animal model for these amino acids. One of these, formation of inositol trisphosphate (IP3) and cyclic AMP (cAMP), is mediated by GTP-binding regulatory proteins, while another involves ion channels directly activated by stimuli. L-ALA is a potent stimulus to cAMP and IP3 accumulation, while L-ARG at low concentrations is without effect. On the other hand, L-ARG and L-PRO, but not L-ALA, are able to activate stimulus-specific and cation-selective channels in taste epithelial membranes reconstituted in phospholipid bilayers at the tips of patch pipettes. Preliminary studies using mouse taste tissue demonstrate that monosodium-L-glutamate (MSG) did not enhance production of IP3 or cAMP. However, in reconstitution experiments using taste epithelium of mouse, conductance changes due to MSG are observed. The specificity of this channel(s) and its uniqueness have yet to be determined.

Amino Acids↗

Are umami taste receptor sites structurally related to glutamate CNS receptor sites?

Umami tasting substances, MSG (monosodium glutamate), HG (glutamic acid), LGDE (1-glutamic acid diethyl ester), DLHCA (dl-homocysteic acid), DLAAA (dl-aminoadipic acid) and 5'GMP, were tested on the hamster and the human. Ten mM MSG was routinely used in the hamster as it elicited strong chorda tympani responses. Similar response amplitudes were found for MSG, HG, LGDE, DLAAA 10 mM, DLHCA 8 mM and sucrose 100 mM. A 5 microM concentration of 5'GMP eventually was an efficient stimulus on a few preparations. Such a low concentration is very seldom efficient as a taste stimulus in rodents, indicating a higher specificity of receptor mechanisms than what is usually found for sweet taste, for example. The synergy between MSG and 5'GMP was found in the hamster CT only for concentrations lower than those of the literature, i.e., a mixture of 12 microM 5'GMP and 2.5 mM MSG showed a reinforcement of 50% in response amplitude equivalent to a 100% increase in concentration. We take this as an evidence of an umami component in the hamster CT response to glutamate; in accordance with literature data, we could not find reinforcement for higher concentrations which were in fact near saturation. Responses to MSG, HG, LGDE, DLAAA and DLHCA, among 38 other organic stimuli, were studied in 42 hamster chorda tympani. Responses to HG, LGDE and 5'GMP, among chemoreception of these compounds used as umami tasting stimuli.(ABSTRACT TRUNCATED AT 250 WORDS)

Amino Acids↗

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↗

Interaction of MSG taste with nutrition: perspectives in consummatory behavior and digestion.

Studies in the taste system of mammals indicate that monosodium glutamate (MSG) produces a unique taste sensation termed umami. As a derivative of glutamic acid, MSG is a naturally occurring nutrient in many foods; its commercial use to improve food palatability for humans is well documented. Behavioral studies with experimental animals have revealed that preference for MSG in solutions and selection of MSG-flavored diets can be explained by sensory means with no appreciable effects on preference by postingestive consequences. However, preference for umami-flavored fluids is reduced by feeding rats low-protein diets or low quality protein. MSG-flavored diets, initially unpreferred, were subsequently highly selected. However, the adulteration of diets by MSG either did not or only slightly increased food intake. In light of the significant contribution of MSG to diet taste, apparently as a signal coupled to dietary proteins, physiological studies have been initiated to reveal its role as a stimulator of the cephalic phase of pancreatic exocrine secretion. Modified sham-feeding experiments with dogs have shown that oral stimulation by MSG produced significant stimulation of both pancreatic flow and protein output in conscious dogs.

Amino Acids↗