Attention and the detectability of weak taste stimuli

Subjects detected weak solutions of sucrose or citric acid under conditions in which attention was directed toward one of the tastants or the other. Detection thresholds were measured using an adaptive, forced-choice procedure, with a three-down one-up rule, which computer simulations suggest should be more reliable than the popular two-down one-up rule. The thresholds were modestly but systematically lower for attended tastants than for unattended ones. Similar results have been reported in other sense modalities, including vision (greater sensitivity to stimuli presented to attended versus unattended spatial locations) and hearing (greater sensitivity to stimuli presented at attended versus unattended sound frequencies). Taken together, the findings are consistent with a general hypothesis regarding attention in sensory systems: gains or losses in detectability occur when a central attentional mechanism (or, conceivably, a preattentive mechanism) selectively and preferentially monitors signals arising from particular subsets of peripheral neural inputs.      

Spatial summation in taste: NaCl thresholds and stimulated area on the anterior human tongue

Spatial summation has been demonstrated in several sensory modalities.In this study, spatial summation of sensitivity to NaCl wasinvestigated as a function of stimulated area. In a two-alternativeforced choice procedure, a filter paper stimulation method wasused to determine the sensitivity to, and the detection probabilitiesof, five low-intensity NaCl stimuli. Each of the stimuli waspresented 32 times in two conditions, a circular area (ø9 mm) and half that circle in four counterbalanced orientations.Only one side of the tongue was used. The results showed that(a) the summation found was partial; (b) the observed relationshipbetween threshold intensity and area obeyed Piper's Law, and(c) detection probabilities increased according to chance. Adhoc analysis of orientation yielded sensitivity differencesalong the vertical but not the horizontal axis.     

Differential spatial displacement discrimination with interfering stimuli

Differential spatial displacement discrimination thresholds were determined for a configuration of three blobs with Gaussian spatial and temporal contrast envelopes. This task is similar to the well known three-dot alignment hyperacuity task. Thresholds determined in the presence of interfering stimuli were identical to thresholds determined without these flanking stimuli. The thresholds scale linearly with stimulus size over at least two decades. We conclude that (i) the mechanisms that compute differential spatial displacement for the three-blob alignment task are not disturbed by the presence of neighbouring stimuli, even when these enter the region over which the computations are performed and (ii) at all levels of resolution similar mechanisms are used to compute differential spatial displacement.     

Concentrations of NaCl and sucrose: equality in perceived taste intensity

Results from previous studies suggest that sucrose and NaClsolutions have an equal perceived taste intensity when the molarconcentration of sucrose is 1.5–1.75 times the molar concentrationof NaCl. However, according to other studies, sucrose and NaClsolutions taste equally strong when their molar concentrationsare about equal. This issue was further pursued using the methodof constant stimuli, where subjects matched the perceived tasteintensity of NaCl solutions to five sucrose references and viceversa. The results concur with previous findings that sucroseand NaCl solutions have equal perceived taste intensities whenthe molar concentration of sucrose is 1.5–1.75 times themolar concentration of NaCl.     

Facial electromyography: responses of children to odor and taste stimuli

The study investigated the potential for facial electromyography (EMG) to be used as a clinical tool for measuring the responses of children to pleasant and unpleasant smell and taste stimuli. Responses in the zygomaticus major and levator labii muscles to 4 odorants and 4 tastants were recorded from 34 children aged 6-9 years. The results indicated that EMG activities in the 2 muscles discriminated between pleasant and unpleasant stimuli within each modality in a manner that indicated that the children perceived the hedonic qualities of the stimuli in a manner similar to that reported for adults. Importantly, there was unanimous agreement across the children as regards the differential nature of the activities exhibited. These outcomes together with the results of earlier facial expression studies suggest that facial EMG may provide an objective procedure that could be suitable for the clinical assessment of taste and smell function in newborns and young infants.     

Monosodium glutamate and sweet taste: generalization of conditioned taste aversion between glutamate and sweet stimuli in rats

Even though monosodium glutamate (MSG) is a prototypical umami substance, previous studies reported that a conditioned taste aversion (CTA) to MSG, mixed with amiloride to block the taste of sodium, generalizes to sucrose. These findings suggest that the taste of glutamate mimics the taste of sucrose and raise the question of whether glutamate has a broadly tuned sweet taste component. To test this hypothesis, CTA experiments were conducted to test for generalization between MSG and several sweet stimuli: sucrose, glucose, maltose, saccharin and SC-45647. Strong bidirectional generalization was seen between MSG mixed with amiloride and sucrose, glucose, saccharin and SC-45647. Weak generalization was seen between MSG and maltose, and sucrose and maltose. None of the CTAs generalized to NMDA. These findings support the hypothesis that the taste of MSG has broadly tuned, sweet-like characteristics, possibly due to the convergence of afferent signals for MSG, natural sugars and artificial sweeteners.     

Generalization of learned taste aversions in rabbits: similarities among gustatory stimuli

Although rabbits have been used in a number of electrophysiologicaland anatomical studies on the gustatory system, there have beenfew behavioral experiments on these animals and these have beenlimited to studies of taste preference. The similarities amonga number of gustatory stimuli were assessed in rabbits by measuringthe generalization patterns in a conditioned taste aversionexperiment. Rabbits were trained to take their daily rationof water within a 30-min session, during which the number oflicks per 10-s presentation of a drinking tube could be recorded.During one of these sessions, one of 12 stimuli (sucrose, fructose,Na-saccharin, NaCI, NaNO₃, Na₂SO₄, KC1, NH₄C1, CaCl₂, HC1, QHC1or urea) was presented, followed by i.p. injection of LiCl toproduce a conditioned taste aversion. Animals were then testedwith all of the stimuli and the amount of suppression of lickingwas used as a measure of stimulus generalization. The patternsof generalization were compared for the test and conditioningstimuli separately. Some nonreciprocities were seen betweenthe conditioning and test stimuli, which reflected the occurrenceof multiple taste qualities and the tendency for aversions togeneralize more to stronger stimuli than to weaker ones. Principalcomponents analysis of the stimulus relationships showed thatrabbits responded to these stimuli in a fashion similar to thatof other mammals, including humans. Within the principal componentssolution, there were strong similarities among the sugars, thesodium salts, the nonsodium salts and the bitter-tasting stimuli.     

A four-channel analysis of the tactile sensitivity of the fingertip: frequency selectivity,spatial summation,and temporal summation

Thresholds were measured for the detection of vibratory stimuli of variable frequency and duration applied to the index fingertip and thenar eminence through contactors of different sizes. The effects of stimulus frequency could be accounted for by the frequency characteristics of the Pacinian (P), non-Pacinian (NP) I, and NP III channels previously determined for the thenar eminence (Bolanowski et al., J Acoust Soc Am 84: 1680-1694, 1988; Gescheider et al., Somatosens Mot Res 18: 191-201, 2001). The effect of changing stimulus duration was also essentially identical for both sites, demonstrating the same amount of temporal summation in the P channel. Although the effect of changing stimulus frequency and changing stimulus duration did not differ for the two sites, the effect of varying the size of the stimulus was significantly greater for the thenar eminence than for the fingertip. The attenuated amount of spatial summation on the fingertip was interpreted as an indication that the mechanism of spatial summation consists of the operations of both neural integration and probability summation.     

A four-channel analysis of the tactile sensitivity of the fingertip: frequency selectivity,spatial summation,and temporal summation

Thresholds were measured for the detection of vibratory stimuli of variable frequency and duration applied to the index fingertip and thenar eminence through contactors of different sizes. The effects of stimulus frequency could be accounted for by the frequency characteristics of the Pacinian (P), non-Pacinian (NP) I, and NP III channels previously determined for the thenar eminence (Bolanowski et al., J Acoust Soc Am 84 : 1680-1694, 1988; Gescheider et al., Somatosens Mot Res 18: 191- 201, 2001). The effect of changing stimulus duration was also essentially identical for both sites, demonstrating the same amount of temporal summation in the P channel. Although the effect of changing stimulus frequency and changing stimulus duration did not differ for the two sites, the effect of varying the size of the stimulus was significantly greater for the thenar eminence than for the fingertip. The attenuated amount of spatial summation on the fingertip was interpreted as an indication that the mechanism of spatial summation consists of the operations of both neural integration and probability summation.     

Amiloride-sensitive NaCl taste responses are associated with genetic variation of ENaC alpha-subunit in mice

An epithelial Na(+) channel (ENaC) is expressed in taste cells and may be involved in the salt taste transduction. ENaC activity is blocked by amiloride, which in several mammalian species also inhibits taste responses to NaCl. In mice, lingual application of amiloride inhibits NaCl responses in the chorda tympani (CT) gustatory nerve much stronger in the C57BL/6 (B6) strain than in the 129P3/J (129) strain. We examined whether this strain difference is related to gene sequence variation or mRNA expression of three ENaC subunits (alpha, beta, gamma). Real-time RT-PCR and in situ hybridization detected no significant strain differences in expression of all three ENaC subunits in fungiform papillae. Sequences of the beta- and gammaENaC subunit genes were also similar in the B6 and 129 strains, but alphaENaC gene had three single nucleotide polymorphisms (SNPs). One of these SNPs resulted in a substitution of arginine in the B6 strain to tryptophan in the 129 strain (R616W) in the alphaENaC protein. To examine association of this SNP with amiloride sensitivity of CT responses to NaCl, we produced F(2) hybrids between B6 and 129 strains. Amiloride inhibited CT responses to NaCl in F(2) hybrids with B6/129 and B6/B6 alphaENaC R616W genotypes stronger than in F(2) hybrids with 129/129 genotype. This suggests that the R616W variation in the alphaENaC subunit affects amiloride sensitivity of the ENaC channel and provides evidence that ENaC is involved in amiloride-sensitive salt taste responses in mice.     

Tight junctions in taste buds: possible role in perception of intravascular gustatory stimuli

Exogenous chemicals having low taste thresholds elicit particulartastes when injected into the bloodstream. This phenomenon iscalled intravascular taste. To explore the origins of intravasculartaste we investigated the permeability properties of the paracellularpathways (tight junctions) between taste cells and between epithelialcells in canine fungiform papillae. This was achieved by showingthat the transepithelial resistance (TER), which is a measureof the paracellular pathway resistance, increases upon the additionof LaCl₃. Thin-section electron microscopy of the same epitheliaused for the TER measurements showed that lanthanum depositsare found exclusively in the extracellular spaces. In the epithelium,LaCl₃ added to either the mucosal or serosal solutions did notdiffuse past the tight junctions at the interface between thestrata cornea and granulosa. The blockage of epithelial tightjunctions by lanthanum is responsible for the increase in TER.LaCl₃ added to the serosal solution was observed throughoutthe extracellular spaces between taste cells including the extracellularspace beyond the tight junctions in the taste pore. Thus, tightjunctions of taste cells and epithelial cells differ in theirpermeability to LaCl₃. From these observations we conclude thatthe tight junctions between taste cells are more permeable tomolecules of small molecular weight than are the tight junctionsbetween epithelial cells. Therefore, small molecules that leavethe bloodstream can diffuse into the taste pore and interactwith receptors in the microvilli of taste cells resulting inintravascular taste.     

Sensory irritation and taste produced by NaCl and citric acid: effects of capsaicin desensitization

Three experiments were conducted to measure the sensory irritationproduced by two prototypical gustatory stimuli: citric acidand NaCl. The stimuli were applied to the tip of the tongueon filter paper disks. The first experiment revealed that solutionsof NaCl and citric acid that produced approximately equal tastesensations also produced similar amounts of irritation; thatthe psychophysical functions for irritation were approximatelytwice as steep as the functions for taste; and that irritationgrew over time for NaCl but not for citric acid. When viewedas a percentage of the taste sensation at 25 s, NaCl irritationaveraged 23% at the lowest concentration and 70% at the highestconcentration; citric acid irritation averaged 44% at the lowestconcentration and 98% at the highest concentration. The secondexperiment investigated whether the irritation produced by thesetwo stimuli was mediated via capsaicin-sensitive (CS) fibers.The experiment included a pre-test, an irritation treatmentwith either capsaicin (a desensitizing agent) or zingerone (anon-desensitizing agent), a 15 min rest period and a post-test.Reductions in irritation and taste occurred following treatmentwith both capsaicin and zingerone. A third experiment demonstratedthat the majority of the effect of zingerone on taste and irritationwas due to a perceptual context effect. After the context effectwas taken into account, capsaicin desensitization remained significantfor both salt taste and salt irritation at the highest concentration.A similar pattern of results for citric acid suggests that bothcitric acid and NaCl produce irritation in part via CS fibers.The results are discussed in terms of the ability of subjectsto discriminate the gustatory and chemesthetic components oforal sensations and the role of salt and acid irritation inflavor perception.     

Salivary secretion elicited by taste stimulation with umami substances in human adults

Salivary secretion from the parotid gland was measured whenumami substances (monosodium L-glutamatc, disodium 5'-inosinateand disodium 5'-guanylate) were applied to the anterior or posteriorparts of the tongue in human adults. The amount of secretioninduced by a low concentration (0.01 M) of umami substanceswas very small and similar to that elicited by 1 M sucrose.Salivation induced by a high concentration (0.1 M) of umamisubstances was about twice that induced by the low concentration,but less than that induced by 0.1 M NaCl. The volume of secretioninduced by a mixture containing the low concentration of bothmonosodium L-glutamate and disodium 5'-inosinate (or disodium5'-guanylate) approximated the arithmetic sum of secretionsinduced by the individual components, whereas a mixture of thehigh concentration of these substances elicited much less salivationthan the sum of secretions for the components. More copiousflow of parotid saliva tended to be induced from stimulatingthe posterior tongue compared to the anterior tongue. The presentresults are discussed in terms of the taste properties of umamisubstances as shown in previous psychophysical and elcctrophysiologicalstudies.     

Measures of taste discriminability for sweet, salty and umami stimuli: Japanese versus Americans

Japanese and American subjects performed triangle tests withvarious concentrations of sucrose. NaCl and MSG. Both sets ofsubjects were tested in their own countries in their own languages.The Japanese discriminated significantly better for sucroseand MSG, while for NaCl there were no significant differencesbetween the groups. To protect against variation in water puritybetween the American and Japanese laboratories. 10 mM NaCl wasused as the solvent, while judges were pre-adapted to the solventto render it tasteless. Changing from purified water to 10 mMNaCl as the solvent, alters the discriminability of the stimuli.     

A general rule for sensory cue summation: evidence from photographic, musical, phonetic and cross-modal stimuli

The Euclidean and MAX metrics have been widely used to model cue summation psychophysically and computationally. Both rules happen to be special cases of a more general Minkowski summation rule , where m = 2 and ∞, respectively. In vision research, Minkowski summation with power m = 3-4 has been shown to be a superior model of how subthreshold components sum to give an overall detection threshold. Recently, we have previously reported that Minkowski summation with power m = 2.84 accurately models summation of suprathreshold visual cues in photographs. In four suprathreshold discrimination experiments, we confirm the previous findings with new visual stimuli and extend the applicability of this rule to cue combination in auditory stimuli (musical sequences and phonetic utterances, where m = 2.95 and 2.54, respectively) and cross-modal stimuli (m = 2.56). In all cases, Minkowski summation with power m = 2.5-3 outperforms the Euclidean and MAX operator models. We propose that this reflects the summation of neuronal responses that are not entirely independent but which show some correlation in their magnitudes. Our findings are consistent with electrophysiological research that demonstrates signal correlations (r = 0.1-0.2) between sensory neurons when these are presented with natural stimuli.     

Monosodium glutamate and sweet taste: discrimination between the tastes of sweet stimuli and glutamate in rats

Generalization of a conditioned taste aversion (CTA) is based on similarities in taste qualities shared by the aversive substance and another taste substance. CTA experiments with rats have found that an aversion to a variety of sweet stimuli will cross-generalize with monosodium glutamate (MSG) when amiloride, a sodium channel blocker, is added to all solutions to reduce the taste of sodium. These findings suggest that the glutamate anion elicits a sweet taste sensation in rats. CTA experiments, however, generally do not indicate whether two substances have different taste qualities. In this study, discrimination methods in which rats focused on perceptual differences were used to determine if they could distinguish between the tastes of MSG and four sweet substances. As expected, rats readily discriminated between two natural sugars (sucrose, glucose) and two artificial sweeteners (saccharin, SC45647). Rats also easily discriminated between MSG and glucose, saccharin and, to a lesser extent, SC45647 when the taste of the sodium ion of MSG was reduced by the addition of amiloride to all solutions, or the addition of amiloride to all solutions and NaCl to each sweet stimulus to match the concentration of Na+ in the MSG solutions. In contrast, reducing the cue function of the Na+ ion significantly decreased their ability to discriminate between sucrose and MSG. These results suggest that the sweet qualities of glutamate taste is not as dominate a component of glutamate taste as CTA experiments suggest and these qualities are most closely related to the taste qualities of sucrose. The findings of this study, in conjunction with other research, suggest that sweet and umami afferent signaling may converge through a taste receptor with a high affinity for glutamate and sucrose or a downstream transduction mechanism. These data also suggest that rats do not necessarily perceive the tastes of these sweet stimuli as similar and that these sweet stimuli are detected by multiple sweet receptors.