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PubMed1992

A novel method to analyse response patterns of taste neurons by artificial neural networks.

Despite several notions on the gustatory code proposed over three decades, investigators have not yet reached a consensus. This paper describes a new approach to analyse gustatory neural activities. Three-layer neural networks were trained by the back-propagation learning algorithm, to classify the neural response patterns to four basic taste qualities. The discrimination by the trained networks on taste qualities in the response patterns of rat chorda tympani fibres (CT) and cortical taste neurons (CN) was consistent both with the correlation analysis and with behavioural experiments. By examining the connection weights of each neuron, some input neurons representing CN were 'pruned' without deteriorating the ability of the network to discriminate taste. This characteristic of the network is contrary to a previous hypothesis, that taste neurons are of equal importance in the neural coding.

Animals↗

PubMed2005

In vivo recordings from rat geniculate ganglia: taste response properties of individual greater superficial petrosal and chorda tympani neurones.

Coding of gustatory information is complex and unique among sensory systems; information is received by multiple receptor populations located throughout the oral cavity and carried to a single central relay by four separate nerves. The geniculate ganglion is the location of the somata of two of these nerves, the greater superficial petrosal (GSP) and the chorda tympani (CT). The GSP innervates taste buds on the palate and the CT innervates taste buds on the anterior tongue. To obtain requisite taste response profiles of GSP neurones, we recorded neurophysiological responses to taste stimuli of individual geniculate ganglion neurones in vivo in the rat and compared them to those from the CT. GSP neurones had a distinct pattern of responding compared to CT neurones. For example, a small subset of GSP neurones had high response frequencies to sucrose stimulation, whereas no CT neurones had high response frequencies to sucrose. In contrast, NaCl elicited high response frequencies in a small subset of CT neurones and elicited moderate response frequencies in a relatively large proportion of GSP neurones. The robust whole-nerve response to sucrose in the GSP may be attributable to relatively few, narrowly tuned neurones, whereas the response to NaCl in the GSP may relate to proportionately more, widely tuned neurones. These results demonstrate the diversity in the initial stages of sensory coding for two separate gustatory nerves involved in the ingestion or rejection of taste solutions, and may have implications for central coding of gustatory quality and concentration as well as coding of information used in controlling energy, fluid and electrolyte homeostasis.

Action Potentials↗

PubMed1993

Perceptual consequences of electrical stimulation in the gustatory system.

To investigate the role of temporal coding in the neural processing of taste, trains of electrical pulses of varying frequency were delivered to the nucleus of the solitary tract (NTS) in awake rats. The temporal patterns of these trains mimicked the temporal patterns of electrophysiological responses of single neurons to natural tastes. In Experiment 1, water-deprived rats were first trained to lick water in an experimental chamber. On training days, licking water produced a sucroselike electrical pulse train in the NTS. At the end of these sessions, experimental animals were made ill by an injection of LiCl and subsequently learned to avoid licking when LiCl was paired with NTS stimulation. In Experiment 2, rats refused to lick water when licking produced a quininelike pattern of NTS stimulation but licked enthusiastically when licking produced a pattern of NTS stimulation similar to the natural response to sucrose.

Afferent Pathways↗

PubMed1996

Gustatory neural coding in the monkey cortex: mixtures.

1. Psychophysicists have shown that the intensity and quality of a taste stimulus, as perceived by humans, is modified by including that stimulus in a mixture. Gustatory neurons in the primary taste cortex (anterior insula and frontal operculum) of the cynomolgus macaque are involved with the coding of stimulus intensity and quality, and so should reflect the impact of these stimulus interactions. 2. We recorded the activity of 48 neurons in primary taste cortex in response to the oral application of each of the four basic stimuli, their six possible dyads, the four triads, and the tetrad of all four. Stimuli were maintained at a constant intensity in all mixtures by increasing their concentrations as the number of components rose. 3. Glucose was the most effective basic stimulus, followed by quinine HCl, NaCl, and HCl. The mean response to dyads was suppressed by 50% from the sum of responses to the two unmixed components. The response to triads was 62% lower than the sum of responses to their three components, and activity evoked by the tetrad was suppressed by 74% from the sum of all four individual responses. Therefore there was nearly total suppression in the sense that the responses to the mixtures were approximately 1/2, 1/3, and 1/4 the sums of responses to two, three, and four components, respectively. 4. Neurons could be divided into four subtypes: those that responded best to each of the basic stimuli. All subtypes except HCl cells were about equally suppressed when their preferred stimulus was included in a mixture. HCl was a particularly ineffective stimulus, such that this subtype responded poorly and so was less susceptible to mixture suppression. 5. Taste quality, as indexed by correlation coefficients among profiles of activity, was quite predictable for dyads. If the mixture included HCl, the profile it generated correlated poorly (about +0.20) with that of HCl and rather well (about +0.60) with that of the other component. If HCl was not included, the mixture's profile correlated about +0.40 with that of each component. 6. The profile generated by the mixture of three stimuli was predictable only if one of the components was HCl. In that case, the triad elicited a profile midway between those of the other two components, i.e., the contribution of HCl was largely ignored. When HCl was not involved, or when all four basic stimuli were combined, the resulting profiles were poorly correlated with those of all basic stimuli. 7. The contribution made by each basic taste to human perception and to the macaque's neurophysiological response was compared for all mixtures. The contribution was often quite similar for human and macaque, but when differences occurred, they were typically due to lower activity from HCl cells in the macaque, a loss that was replaced mainly by larger responses from glucose neurons. 8. The magnitude of responses to mixtures in the macaque taste cortex matches well with expectations from human psychophysical studies. The presumed quality of the response to mixtures is also similar, except that HCl is less effective in monkeys and sugars more so.

Action Potentials↗

PubMed2003

Taste and pheromone perception in mammals and flies.

The olfactory systems of insects and mammals have analogous anatomical features and use similar molecular logic for olfactory coding. The molecular underpinnings of the chemosensory systems that detect taste and pheromone cues have only recently been characterized. Comparison of these systems in Drosophila and mouse uncovers clear differences and a few surprising similarities.

Acids↗

PubMed2006

State-dependent modulation of time-varying gustatory responses.

Sensory processing is modulated by attention, which is a function of network states. Here we show that changes in such states do more than a simple gating of stimuli: they actually re-arrange cortical coding space to emphasize emotional valences. We delivered taste stimuli to rats before and after a spontaneous state change ("disengagement") that is associated with a reduction in attention and a concurrent emergence of cortical mu rhythms. The percentage of cortical neurons that responded to tastes, and the average response across neurons, remained stable with disengagement, but the particulars of the responses changed drastically. The distinctiveness of sucrose and quinine-which represent the high and low ends of the palatability spectrum-increased, the distinctiveness of the two aversive tastes (quinine and citric acid) decreased, and the distinctiveness of sucrose and NaCl, which were almost identically palatable to start with, did not change. Overall, then, the changes appeared to be palatability-specific. Two additional findings were consistent with this conclusion: rats' palatability-related behavioral responses to the tastes changed in similar ways with disengagement and disengagement-related neural changes specifically appeared late in the response, when palatability-specific information emerges in cortical responses. These data suggest that neural state changes can change the content of neural codes.

Action Potentials↗

PubMed2018

TAS2R38 Predisposition to Bitter Taste Associated with Differential Changes in Vegetable Intake in Response to a Community-Based Dietary Intervention.

Although vegetable consumption associates with decreased risk for a variety of diseases, few Americans meet dietary recommendations for vegetable intake. TAS2R38 encodes a taste receptor that confers bitter taste sensing from chemicals found in some vegetables. Common polymorphisms in TAS2R38 lead to coding substitutions that alter receptor function and result in the loss of bitter taste perception. Our study examined whether bitter taste perception TAS2R38 diplotypes associated with vegetable consumption in participants enrolled in either an enhanced or a minimal nutrition counseling intervention. DNA was isolated from the peripheral blood cells of study participants (N = 497) and analyzed for polymorphisms. Vegetable consumption was determined using the Block Fruit and Vegetable screener. We tested for differences in the frequency of vegetable consumption between intervention and genotype groups over time using mixed effects models. Baseline vegetable consumption frequency did not associate with bitter taste diplotypes (P = 0.937), however after six months of the intervention, we observed an interaction between bitter taste diplotypes and time (P = 0.046). Participants in the enhanced intervention increased their vegetable consumption frequency (P = 0.020) and within this intervention group, the bitter non-tasters and intermediate-bitter tasters had the largest increase in vegetable consumption. In contrast, in the minimal intervention group, the bitter tasting participants reported a decrease in vegetable consumption. Bitter-non tasters and intermediate-bitter tasters increased vegetable consumption in either intervention more than those who perceive bitterness. Future precision medicine applications could consider genetic variation in bitter taste perception genes when designing dietary interventions.

Adolescent↗

PubMed2005

Gustatory coding in the precentral extension of area 3 in Japanese macaque monkeys; comparison with area G.

The precentral extension of area 3 as well as the transition between the frontal operculum and insula (area G) comprises the primary gustatory cortex in the subhuman primate, receiving projections from the thalamic taste relay. However, in contrast to the extensive studies that have been carried out on the latter area, only a few taste units in the former area have been recorded. To clarify gustatory coding in area 3, we investigated the taste response properties of neurons in area 3 compared with those in area G in alert monkeys by infiltrating into their mouths seven taste stimuli [0.3 M sucrose (S), 0.1 M NaCl (N), 0.01 N HCl (H), 0.003 M quinine-HCl (Q), 0.1 M monosodium glutamate (MSG), distilled water (W), and orange juice (OR)] and artificial saliva (SA). A larger number of HCl-best units and a smaller number of quinine-best units were found in area 3 than in area G. The onset latency and response duration were significantly shorter in area 3 than in area G. Weighted multi-dimensional scaling showed that area G divided eight stimulants into four classes, i.e. two groups (H-Q-W and S-MSG-OR), N and SA, whereas area 3 divided them into three classes (N-MSG-W-OR, S-Q, and H-SA). This suggested that tastants not separated in area G were separated in area 3, and vice versa. This indicates that both areas complement each other in the representation of taste stimuli, each contributing to taste information processing in a different manner.

Algorithms↗

PubMedDate not supplied

Issues of gustatory neural coding: where they stand today.

The basic issues of gustatory neural coding are revisited. Questions addressed and conclusions drawn are: (1) what is the physical dimension across which gustatory neurons are sensitive, and upon which taste perceptions are based? The dimension that unites the various taste qualities is not physical, but physiological: a dimension of well-being, bounded by toxins at one extreme and nutrients at the other. (2) How broadly tuned are taste cells across the dimension? There are instances of specificity, but most mammalian taste cells respond to a range of qualities. (3) Are there basic taste qualities? Sweet, salty, sour, and bitter are widely accepted as basic tastes. Umami and starch tastes are considered basic by some. (4) Is taste topographically organized? There is some degree of physical separation among neurons most responsive to different taste qualities, but this does not appear to be sufficient precision to act as a meaningful coding mechanism. (5) Are there gustatory neuron types? Neurons, separated into categories according to their response profiles, respond as members of their category to the challenges of conditioned aversions and preferences, sodium deprivation, hyperglycemia, and receptor blockade, while cells from other categories react differently. This indicates the existence of functionally distinct types of taste cells. (6) Is the quality signal coded within the activity of the single most appropriate category of neurons, or is it carried by the pattern of response across neuronal categories? Both the breadth of responsiveness and the logical ambiguity of the signal in any one category of neurons argue that the taste message is carried by a pattern of activity across gustatory neuron types.

Animals↗

PubMed1997

Transfer of information about taste from the nucleus of the solitary tract to the parabrachial nucleus of the pons.

In the study of the neural code for gustation, the relative sensitivity of a cell to a variety of taste stimuli is defined as its response profile. To study the construction of response profiles from incoming signals, electrophysiological responses to NaCl, HCI, quinine-HCl, sucrose and Na saccharin were recorded simultaneously in pairs of single cells: one in the nucleus of the solitary tract (NTS) and the other in the parabrachial nucleus of the pons (PbN), respectively the first and second synapses in the central pathway for gustation. Of 37 units recorded in the NTS and 32 in the PbN, 12 (32%) pairs showed evidence of functional connectivity. Although PbN responses were significantly larger than those in the NTS in general, no amplification of NTS activity was apparent among those units that were functionally connected. Analysis of NTS-PbN connectivity patterns suggests that PbN units receive input from NTS units with response profiles that are both similar and different from their own pattern of sensitivities. Further analysis suggests that the stimulus-selectivity of the response profile of a PbN unit may be determined by stimulus-selective input from NTS cells that show similar response profiles. However, input from NTS cells with response profiles different from their own appears to be non-stimulus-selective. Analysis of the organization of response profiles in the two structures suggests that the cells in both the NTS and PbN cannot be easily distinguished by their patterns of sensitivity to taste stimuli.

Animals↗

PubMed1997

Preference conditioning alters taste responses in the nucleus of the solitary tract of the rat.

Aversive conditioning has an impact on the neural signal for the gustatory conditioned stimulus (CS). Here, we determined whether the code is also affected by preference conditioning. We paired the taste of MgCl2 (CS+) with intragastric nutrients in some rats (MG), and citric acid (CS+) with nutrients in others (CI). A control group (Control) experienced both tastants without nutrients. Preferences (>90%) developed for each CS+. We recorded responses to 16 taste stimuli in the nucleus of the solitary tract. Responsiveness of acid-oriented neurons to MgCl2 in MG rats was lower than in Controls, and its profile was more distinct from those of acidic and bitter stimuli. Total activity to citric acid was unchanged in CI rats. However, its temporal profile showed a decreased phasic component, making citric acid temporally distinct from nonsugars. Therefore, the responses to both CS+ were modified, each in its own manner, to be more distinct from those of aversive stimuli. The effects of preference conditioning, however, were weaker than those of aversive conditioning.

Animals↗

PubMed1998

Electrophysiological responses to bitter stimuli in primate cortex.

Studies investigating fine details of gustatory coding in the domain of each basic taste quality have been completed for sweet, salt, and sour stimuli. In the present experiment, we used chemicals that humans describe as predominantly bitter. We recorded the activity of 50 taste neurons in insular cortex of two cynomolgus macaques. Stimuli were water, fruit juice, glucose, NaCl, HCl, and 16 bitter solutions. In a multidimensional taste space the 16 bitter stimuli formed a coherent cluster composed of three main subgroups: (1) QHCl, phenylalanine, theophylline, caffeine, propyl-thiouracil (PROP), and phenylthiocarbamide (PTC), all of which humans describe as rather purely bitter, (2) MgCl2, CaCl2, NH4Cl, and arginine, which humans describe as salty-bitter, and (3) urea, cysteine, and vitamin B1, which are described as sour-bitter. Vitamin B2, histidine and nicotine were in the center of the bitter cluster. Human descriptions of taste qualities conformed well to the presumed quality of each stimulus as inferred from its position in the multidimensional space (MDS), reinforcing the use of the macaque as a neural model for human gustation.

Animals↗

PubMed2005

The representation of taste quality in the mammalian nervous system.

The process by which the mammalian nervous system represents the features of a sapid stimulus that lead to a perception of taste quality has long been controversial. The labeled-line (sparse coding) view differs from the across-neuron pattern (ensemble) counterpoint in proposing that activity in a given class of neurons is necessary and sufficient to generate a specific taste perception. This article critically reviews molecular, electro-physiological, and behavioral findings that bear on the issue. In the peripheral gustatory system, the authors conclude that most qualities appear to be signaled by labeled lines; however, elements of both types of coding characterize signaling of sodium salts. Given the heterogeneity of neuronal tuning functions in the brain, the central coding mechanism is less clear. Both sparse coding and neuronal ensemble models remain viable possibilities. Furthermore, temporal patterns of discharge could contribute additional information. Ultimately, until specific classes of neurons can be selectively manipulated and perceptual consequences assessed, it will be difficult to go beyond mere correlation and conclusively discern the validity of these coding models.

Afferent Pathways↗

PubMedDate not supplied

Neuron types, receptors, behavior, and taste quality.

Neurophysiological studies on chorda tympani (CT) single fibers and behavioral studies on generalization of learned aversions in hamsters (Mesocricetus auratus) are reviewed. The work on hamsters is compared to work on other species, including the laboratory rat and several primate species, including humans. This body of data demonstrates associations between response profiles of physiologically defined specialist CT neurons and behavioral stimulus generalizations on one hand, and characteristics of putative taste receptors, on the other. Response profiles of generalist CT neurons are similarly associated with receptor characteristics, but are not associated with specific behavioral discriminations. The associations of peripheral nerve data with both receptor and behavior strongly suggest specific codes for "sucrose-like" and "NaCl-like" taste qualities. Definitive conclusions regarding "patterns" or "labeled lines" requires an understanding of mechanisms of central neural processing of the several specialist and generalist taste-afferent inputs.

Animals↗

PubMed2006

Variation in the human TAS1R taste receptor genes.

We have performed a comprehensive evaluation of single-nucleotide polymorphisms (SNPs) and haplotypes in the human TAS1R gene family, which encodes receptors for sweet and umami tastes. Complete DNA sequences of TAS1R1-, TAS1R2-, and TAS1R3-coding regions, obtained from 88 individuals of African, Asian, European, and Native American origin, revealed substantial coding and noncoding diversity: polymorphisms are common in these genes, and polymorphic sites and SNP frequencies vary widely in human populations. The genes TAS1R1 and TAS1R3, which encode proteins that act as a dimer to form the umami (glutamate) taste receptor, showed less variation than the TAS1R2 gene, which acts as a dimer with TAS1R3 to form the sweet taste receptor. The TAS1R3 gene, which encodes a subunit common to both the sweet and umami receptors, was the most conserved. Evolutionary genetic analysis indicates that these variants have come to their current frequencies under natural selection during population growth and support the view that the coding sequence variants affect receptor function. We propose that human populations likely vary little with respect to umami perception, which is controlled by one major form of the receptor that is optimized for detecting glutamate but may vary much more with respect to sweet perception.

DNA↗

PubMed2006

Facial and affective reactions to tastes and their modulation by sadness and joy.

This study examined adults' affective and facial reactions to tastes which differ in quality and valence, and the impact of sadness and joy on these reactions. Thirty-six male and female subjects participated voluntarily. Subjects each tasted 6 ml of a sweet chocolate drink, a bitter quinine solution (0.0015 M) and a bitter-sweet soft drink. Following a baseline period, either joy or sadness was induced using film clips before the same taste stimuli were presented for a second time. Subjects rated the drinks' pleasantness and intensity of taste immediately after each stimulus presentation. Facial reactions were videotaped and analysed using the Facial Action Coding System (FACS [P. Ekman, W.V. Friesen, Facial Action Coding System: Manual. Palo Alto, CA: Consulting Psychologists Press; 1978., P. Ekman, W. Friesen, J. Hager, Facial Action Coding System. Salt Lake City, Utah: Research Nexus; 2002.]). The results strongly indicated that the tastes produced specific facial reactions that bear strong similarities to the facial reactivity patterns found in human newborns. The data also suggest that some adults' facial reactions serve additional communicative functions. Emotions modulated taste ratings, but not facial reactions to tastes. In particular, ratings of the sweet stimulus were modulated in congruence with emotion quality, such that joy increased and sadness decreased the pleasantness and sweetness of the sweet stimulus. No emotion-congruent modulation was found for the pleasantness and intensity ratings of the bitter or the bitter-sweet stimulus. This 'robustness' of bitter taste ratings may reflect a biologically meaningful mechanism.

Adult↗

PubMed1995

Taste responses in neurons in the nucleus of the solitary tract that do and do not project to the parabrachial pons.

1. Mechanisms of neural coding of gustatory stimuli were studied in the nucleus of the solitary tract (NTS), the first relay in the neural pathway for gustation, in anesthetized rats. Taste-responsive NTS units were identified as "relay" or "nonrelay" based on the electrophysiological response to electrical pulses delivered to the parabrachial nucleus of the pons (PbN), the second relay in the neural pathway for gustation. Coding mechanisms in each group were analyzed separately. 2. Taste responses to sapid solutions of NaCl (0.1 M), HCl (0.01 M), quinine HCl (0.01 M), sucrose (0.5 M) and Na-saccharin (0.004 M) were recorded in single units in the NTS. After gustatory stimulation, electrophysiological responses to electrical stimulation of the taste-responsive part of the ipsilateral PbN were recorded. A 0.2-ms pulse was delivered at 75-250 microA at a rates of 1, 25, 50 and 100 pps through a bipolar stainless steel electrode. An antidromic response was defined as a time-locked spike that occurred at a fixed latency after PbN stimulation that followed high stimulation frequencies. A collision test also was performed. 3. Of 42 taste-responsive NTS units, 19 (45%) were relay units, 22 (52%) were nonrelay and 1 unit was activated orthodromically by PbN stimulation. Latencies of evoked spikes ranged from 1.75 to 4.0 ms 2.1 +/- 0.2 ms (mean +/- SE, median, 1.75 ms). 4. Examination of general response characteristics revealed few differences among relay and nonrelay units.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed2003

The receptors for mammalian sweet and umami taste.

Sweet and umami (the taste of monosodium glutamate) are the main attractive taste modalities in humans. T1Rs are candidate mammalian taste receptors that combine to assemble two heteromeric G-protein-coupled receptor complexes: T1R1+3, an umami sensor, and T1R2+3, a sweet receptor. We now report the behavioral and physiological characterization of T1R1, T1R2, and T1R3 knockout mice. We demonstrate that sweet and umami taste are strictly dependent on T1R-receptors, and show that selective elimination of T1R-subunits differentially abolishes detection and perception of these two taste modalities. To examine the basis of sweet tastant recognition and coding, we engineered animals expressing either the human T1R2-receptor (hT1R2), or a modified opioid-receptor (RASSL) in sweet cells. Expression of hT1R2 in mice generates animals with humanized sweet taste preferences, while expression of RASSL drives strong attraction to a synthetic opiate, demonstrating that sweet cells trigger dedicated behavioral outputs, but their tastant selectivity is determined by the nature of the receptors.

Animals↗