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Perceptual integration of tertiary taste mixtures.

Integration psychophysics was used to explore the taste perception of mixtures of sucrose, fructose, and citric acid. Three levels of each stimulus were varied in a 3 x 3 x 3 factorial design. Subjects rated total intensity, sweetness, and acidity of the 27 mixtures on graphic rating scales. Consistent with earlier work, the perceived total intensity of the tertiary mixtures was found to be dictated by the intensity of the (subjectively) stronger component alone (i.e., either the integrated sweetness or the acidity, whichever was the more intense). In contrast, the sweetness and acidity of the mixture were susceptible to mutual suppression: Sweetness suppressed acidity, acidity suppressed sweetness. There was, however, a difference between sucrose and fructose in their interactions with citric acid, fructose being the more susceptible to suppression. This selectivity of suppression indicates that the two sweetnesses could not have been inextricably integrated. Implications for taste coding are discussed, and the findings are reconciled in terms of two separate coding mechanisms: one for taste intensity, another for taste quality.

Adult↗

Locus coeruleus activation transforms cortical taste representations.

Noradrenergic neurons in the locus coeruleus (LC) shape sensory processing, yet how LC activity influences population taste coding remains unclear. Using optogenetic LC activation with miniscope imaging in the gustatory cortex (GC), we examined LC modulation of multiple taste attributes. Phasic LC activation strengthens correlations between neuronal responses and palatability and expands the dynamic range of stimulus representations along a palatability axis. This expansion is driven by an aversive shift in the representations of all tastants except sucrose, the most palatable stimulus. For mixture ratio and concentration, phasic activation expands and rotates attribute axes, potentially reflecting dependencies between these attributes and palatability. These transformations likely arise from multiplicative gain modulation and more flexible tuning changes. Tonic LC activation affects fewer neurons and does not expand attribute axes. Together, the findings show that LC activation reorganizes GC population geometry in a pattern-dependent manner, linking neuromodulation with feeding behavior and affective processing.

CP: neuroscience↗

Responsiveness of the cortical taste area neurons to a mixture of the four basic tastants in rats.

The taste coding mechanism in the cortical taste area was investigated by analyzing the responses of 59 neurons in the cortical taste area of the anesthetized rat to a mixture of the four basic tastants in both absence and presence of bicuculline methiodide, a specific antagonist to the GABA(A) receptors. The mixture caused response suppression more frequently than response facilitation, both in the control state and during bicuculline application. Cluster analysis revealed that only a group of the neurons with the best response to both NaCl and HCl (group NH) showed the best response to the mixture in the control state, whereas during bicuculline application, in addition to group NH, two other groups of neurons responding to sucrose, or to HCl and quinine responded vigorously to the mixture. Multidimensional scaling located the mixture outside the space of the four basic tastants facing an NaCl-HCl line in both states. GABAergic inhibition caused the group NH to represent the taste of the mixture in the control state. Thus, the mixture probably tastes salty and sour to rats. No cortical neuron was found which specifically responded to the mixture.

Animals↗

Human taste: peripheral anatomy, taste transduction, and coding.

The anatomy, physiology and psychology of taste provide a glimpse into a uniquely heterogeneous sensory world; a world that is robust in its importance to flavor, redundant in its transductive heterogeneity and complexity, requisite in that feeding and hence life usually depend upon taste input, regenerative in that taste cells constantly turn over and regrow after tissue damage, and resistant to disease, loss of neural innervation and epithelial destruction. This chapter considers our current state of knowledge in anatomy, taste bud physiology, molecular biology of bitter, sweet, sour, savory and salty tastes, afferent signaling and quality coding, human perception, and pathophysiology and senescence of taste. We highlight some of the advances made in molecular biology of taste and point out areas where further research is needed ranging from taste bud development and regeneration, to within-taste bud processing, to central/perceptual coding networks for taste. Our hope is that this chapter will provide a background for greater understanding of taste physiology, perception, disease, and future sensory research.

Brain↗

Bitter taste in single chorda tympani taste fibers from chimpanzee.

We have found earlier that chimpanzee chorda tympani taste fibers fall into groups that conform with the human taste qualities. This study focuses on bitter taste and its relation to sweet taste. Eight fibers were classified as bitter fibers according to their responses to 31 stimuli. The stimuli included the bitter compounds quinine, denatonium benzoate and caffeine. The results indicate a clear dichotomy between the bitter and sweet fibers. Sweet fibers never responded to the bitter compounds. However, in addition to their responses to the above compounds, some of the bitter fibers were stimulated by other compounds. Most prominent were responses to NaCl-amiloride mixture, KCl and xylitol. In most cases the cause could be assumed to be a bitter taste in the compound. These results suggest that the bitter and sweet tastes are conveyed in specific and separate groups of nerve fibers in the chimpanzee. Because of the closeness between chimpanzee and human, this finding has implications on the question of taste coding in human and the concept of taste qualities.

Animals↗

Taste recognition: food for thought.

The ability to identify food that is nutrient-rich and avoid toxic substances is essential for an animal's survival. Although olfaction and vision contribute to food detection, the gustatory system acts as a final checkpoint control for food acceptance or rejection behavior. Recent studies with model organisms such as mice and Drosophila have identified candidate taste receptors and examined the logic of taste coding in the periphery. Despite differences in terms of gustatory anatomy and taste-receptor families, these gustatory systems share a basic organization that is different from other sensory systems. This review will summarize our current understanding of taste recognition in mammals and Drosophila, highlighting similarities and raising several as yet unanswered questions.

Animals↗

Sweet successes.

Mapping of the chromosomal location of genes essential for sweet and bitter taste and identification of the relevant G protein-coupled receptors reveals unanticipated complexity in taste signaling pathways. The distribution of sweet and bitter receptors suggests complete cellular segregation of these taste modalities. Sweet compounds may be distinguished through differential expression of sweet receptors. Novel heterologous expression systems to test bitter and sweet modalities now provide the tools necessary for understanding taste coding.

Animals↗

Cross-enhancement of the sour taste on single human taste papillae.

The subjective intensity of one taste quality can be increased by prior exposure of the tongue to a different taste quality stimulus. This phenomenon, called cross-enhancement, may be the result of interactions among the physiological mechanisms that code taste quality. Another possible explanation is that the water solvent of the second stimulus acquires a taste after exposure of the tongue to the first stimulus. This water taste could add to the taste of the solute in the second stimulus and result in an increase of its subjective intensity. A third possibility is that taste receptors on the tongue may be sensitized by exposure to a taste stimulus. Using a small number of highly trained subjects, we have demonstrated that sucrose can enhance the intensity of an acid taste on the single papilla. Neither water taste nor sweet taste system activation played any role in the mediation of this enhancement. Through a series of experimentally derived inferential steps, we conclude that this phenomenon depends on the removal of protons from the acid receptors. In addition, we have demonstrated in the single papilla, that suppression of the acid taste when in mixture with sucrose can occur without sweet system activity. We conclude that sugars, through their capacity to bind protons, act to reduce the availability of protons to the acid receptors.

Adult↗

Difference in receptive field features of taste neurons in rat granular and dysgranular insular cortices.

Receptive fields (RFs) of 59 cortical taste neurons (35 in the granular insular area, area GI, 21 in the dysgranular insular area, area DI, and 3 in the agranular insular area, area AI) were identified in the oral cavity of the rat. The fraction of the neurons with RFs in the anterior oral cavity only was significantly larger in area GI (74.3%) than in area DI (42.9%). On the other hand, the fraction of neurons with RFs in both the anterior and posterior oral cavity was larger in area DI (42.9%) than in area GI (11.4%). On the whole, it is suggested that area GI is involved in discrimination of several taste stimuli in the oral cavity, whereas in area DI taste information originating from various regions of the oral cavity is integrated. When neurons were classified according to the best stimulus which most excited the neuron among the four basic tastes, different categories of taste neurons had RFs in different parts of the oral cavity. It is suggested that, in either taste area, different categories of taste neurons are involved in different sorts of taste coding. The majority of neurons in both areas had bilateral RFs. In area GI, neurons with RFs on single subpopulations of taste buds were significantly more numerous at the rostral region of the cortex than at the caudal region. There was no such relation between RF types and cortical localization in area DI. Otherwise, topographic representation of the oral cavity by taste neurons on the cortical surface was not obvious. RF features of taste neurons did not differ across layers in either cortical area.

Animals↗

Gustatory processing: a dynamic systems approach.

Recent gustatory studies have provided a growing body of evidence that taste processing is dynamic and distributed, and the taste system too complex to be adequately described by traditional feed-forward models of taste coding. Current research demonstrates that neuronal responses throughout the gustatory neuroaxis are broad, variable and temporally structured, as a result of the fact that the taste network is extensive and heavily interconnected, containing modulatory pathways, many of which are reciprocal. Multimodal influences (e.g. olfactory and somatosensory) and effects of internal state (e.g. attention and expectation), shown in both behavioral and neuronal responses to taste stimuli, add further complexity to neural taste responses. Future gustatory research should extend to more brain regions, incorporate more connections, and analyze behaviors and neuronal responses in both time- and state-dependent manners.

Animals↗

Differential covariation in taste responsiveness to bitter stimuli in rats.

Variation exists in the sensitivity of individual rodents and humans to different bitter tastants. An absence of uniform correlation in responsiveness to different bitter substances across individuals within a species suggests heterogeneity in the mechanisms underlying stimulus processing within this taste modality. Here, we examined taste responsiveness of individual rats to three bitter compounds (quinine hydrochloride, denatonium benzoate, and cycloheximide) in short-term lick tests to determine the magnitude of covariation among responses to these stimuli and infer commonalities in their receptor and neural mechanisms. Rats were tested with a given pair of bitter stimuli during three sessions comprising randomized trial blocks of six concentrations of each stimulus + deionized water. Psychophysical functions were generated for individual rats for respective stimulus pairs, and concentrations of each stimulus that produced equivalent lick suppression relative to water were correlated across animals. Behavioral taste responsiveness to quinine hydrochloride strongly covaried with responsiveness to denatonium benzoate (r = +0.82). Lick responsiveness to quinine was less robustly correlated with that to cycloheximide (r = +0.44), and denatonium and cycloheximide responses failed to correlate. These results imply substantial overlap in the bitter taste coding mechanisms for quinine and denatonium but some degree of independence in the mechanisms responsible for gustatory processing of cycloheximide. More generally, these data reinforce the notion that bitter taste processing is not a homogeneous event.

Animals↗

Effect of intraduodenal lipid on parabrachial gustatory coding in awake rats.

Intestinal fat differentially suppresses sham feeding of liquid diets and preferred gustatory stimuli. Although the behavioral effect is robust, no electrophysiological evidence exists to account for its neural basis. Therefore, we investigated the effect of intestinal fat on gustatory coding in the pontine parabrachial nuclei (PBN) by recording from single neurons in awake rats before, during, and after intraduodenal infusions of lipid (Intralipid; 10 ml, 5 kcal). Intraduodenal lipid did not alter the response profiles of PBN taste neurons. It did, however, produce an overall decrease in response magnitude (-16.25%; n = 43), with the largest reduction to sucrose (-30%; n = 43). The most pronounced suppression occurred in sucrose-best neurons in response to sucrose (-55%; n = 19), and this effect was largest for the sucrose-specific cells (-77%; n = 3). After lipid infusions, nonspecific neurons in both the sucrose-best and NaCl-best categories also responded less to their best stimulus (sucrose, -46%; n = 16; NaCl, -35%; n = 13). In contrast, no significant changes were found in NaCl-specific cells in response to NaCl. All effects appeared with short latency ( approximately 5 min) and were reversible within the time frame of a meal. In controls, duodenal infusions of saline did not cause any changes in taste responsiveness. These results suggest that intestinal fat has specific effects on taste coding in the PBN that may contribute to the intake suppression of palatable food observed in behavioral studies. The similar, short latency of both the behavioral and neural effects supports the hypothesis of a preabsorptive site of action.

Animals↗

The receptors and cells for mammalian taste.

The emerging picture of taste coding at the periphery is one of elegant simplicity. Contrary to what was generally believed, it is now clear that distinct cell types expressing unique receptors are tuned to detect each of the five basic tastes: sweet, sour, bitter, salty and umami. Importantly, receptor cells for each taste quality function as dedicated sensors wired to elicit stereotypic responses.

Animals↗

The sweet and the bitter of mammalian taste.

The discovery of two families of mammalian taste receptors has provided important insights into taste recognition and taste perception. Recent studies have examined the receptors and signaling pathways that mediate sweet, bitter, and amino acid taste detection in mammals. These studies demonstrate that taste cells are selectively tuned to different taste modalities and clarify the logic of taste coding in the periphery.

Animals↗

Psychophysical investigations of cetylpyridinium chloride in rats: its inherent taste and modifying effects on salt taste.

Salts are transduced by at least 2 mechanisms: (a) antagonized by amiloride and (b) antagonized by cetylpyridinium chloride (CPC). The authors report on 4 behavioral experiments in rats that characterize the orosensory properties of CPC itself as well as its effect in suppressing the intensity of NaCl and KCl taste. Experiments 1 and 2 indicated that CPC has a quinine-like taste quality. Experiments 3 and 4 demonstrated that the recognition of KCl, but not NaCl, is modestly reduced by mixture with CPC. However, control experiments call into question the mechanism of the salt suppression of CPC, because both CPC-salt and quinine-salt mixtures had similar effects. The relevance of these studies for understanding salt and bitter taste coding is discussed.

Animals↗

Insect odor and taste receptors.

Insect odor and taste receptors are highly sensitive detectors of food, mates, and oviposition sites. Following the identification of the first insect odor and taste receptors in Drosophila melanogaster, these receptors were identified in a number of other insects, including the malaria vector mosquito Anopheles gambiae; the silk moth, Bombyx mori; and the tobacco budworm, Heliothis virescens. The chemical specificities of many of the D. melanogaster receptors, as well as a few of the A. gambiae and B. mori receptors, have now been determined either by analysis of deletion mutants or by ectopic expression in in vivo or heterologous expression systems. Here we discuss recent advances in our understanding of the molecular and cellular basis of odor and taste coding in insects.

Animals↗

Single neurons in the nucleus of the solitary tract respond selectively to bitter taste stimuli.

Molecular data suggest that receptors for all bitter ligands are coexpressed in the same taste receptor cells (TRCs), whereas physiological results indicate that individual TRCs respond to only a subset of bitter stimuli. It is also unclear to what extent bitter-responsive neurons are stimulated by nonbitter stimuli. To explore these issues, single neuron responses were recorded from the rat nucleus of the solitary tract (NST) during whole mouth stimulation with a variety of bitter compounds: 10 microM cycloheximide, 7 mM propylthiouracil, 10 mM denatonium benzoate, and 3 mM quinine hydrochloride at intensities matched for behavioral effectiveness. Stimuli representing the remaining putative taste qualities were also tested. Particular emphasis was given to activating taste receptors in the foliate papillae innervated by the quinine-sensitive glossopharyngeal nerve. This method revealed a novel population of bitter-best (B-best) cells with foliate receptive fields and significant selectivity for bitter tastants. Across all neurons, multidimensional scaling depicted bitter stimuli as loosely clustered yet clearly distinct from nonbitter tastants. When neurons with posterior receptive fields were analyzed alone, bitter stimuli formed a tighter cluster. Nevertheless, responses to bitter stimuli were variable across B-best neurons, with cycloheximide the most, and quinine the least frequent optimal stimulus. These results indicate heterogeneity for the processing of ionic and nonionic bitter tastants, which is dependent on receptive field. Further, they suggest that neurons selective for bitter substances could contribute to taste coding.

Algorithms↗

Effects of the central amygdaloid nucleus lesions on the gustatory responses in the parabrachial nucleus in rats.

To access the role of the central nucleus of the amygdala (CeA) in the gustatory activity in the pontine parabrachial nucleus (PBN), the responses to four prototypical taste stimuli (NaCl, HCl, QH2SO4 and sucrose) in the PBN were observed before and after bilateral electrolytic lesion of the CeA in the urethane-anesthetized rat. Of 29 neurons, 14 were classified as NaCl-best, 9 as HCl-best, 3 as QH2SO4-best and 3 sucrose-best. After CeA lesions, the response rates to HCl and QH2SO4 were statistically higher across all PBN neurons (P<0.01). According to the best-stimulus category, the effects on the responses to HCl and QH2SO4 were similarly subjected to these modulations in NaCl-best, HCl-best and QH2SO4-best neurons. Correlation analysis indicated that the CeA lesion depressed the effect on the chemical selection between NaCl and QH2SO4. These findings suggest that the CeA plays an important role in the taste coding at the pontine level and it may be involved in mediating the feeding behavior via modulating the gustatory responses.

Amygdala↗