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The neural code for taste in the nucleus of the solitary tract of the rat: effects of adaptation.

Adaptation of the tongue to NaCl, HCl, quinine or sucrose was used as a tool to study the stability and organization of response profiles in the nucleus of the solitary tract (NTS). Taste responses in the NTS were recorded in anesthetized rats before and after adaptation of the tongue to NaCl, HCl, sucrose or quinine. Results showed that the magnitude of response to test stimuli following adaptation was a function of the context, i.e., adaptation condition, in which the stimuli were presented. Over half of all taste responses were either attenuated or enhanced following the adaptation procedure: NaCl adaptation produced the most widespread, non-stimulus-selective cross-adaptation and sucrose adaptation produced the least frequent cross-adaptation and the most frequent enhancement of taste responses. Adaptation to quinine cross-adapted to sucrose and adaptation to HCl cross-adapted to quinine in over half of the units tested. The adaptation procedure sometimes unmasked taste responses where none were present beforehand and sometimes altered taste responses to test stimuli even though the adapting stimulus did not itself produce a response. These effects demonstrated a form of context-dependency of taste responsiveness in the NTS and further suggest a broad potentiality in the sensitivity of NTS units across taste stimuli. Across unit patterns of response remained distinct from each other under all adaptation conditions. Discriminability of these patterns may provide a neurophysiological basis for residual psychophysical abilities following adaptation.

Action Potentials↗

Expression of Mash1 in basal cells of rat circumvallate taste buds is dependent upon gustatory innervation.

Mash1, a mammalian homologue of the Drosophila achaete-scute proneural gene complex, plays an essential role in differentiation of subsets of peripheral neurons. In this study, using RT-PCR and in situ RT-PCR, we investigated if Mash1 gene expression occurs in rat taste buds. Further, we examined dynamics of Mash1 expression in the process of degeneration and regeneration in denervated rat taste buds. In rat tongue epithelium, Mash1 gene expression is confined to circumvallate, foliate, and fungiform papilla epithelia that include taste buds. In taste buds, Mash1-expressing cells are round cells in the basal compartment. In contrast, the mature taste bud cells do not express the Mash1 gene. Denervation and regeneration experiments show that the expression of Mash1 requires gustatory innervation. We conclude that Mash1 is expressed in cells of the taste bud lineage, and that the expression of Mash1 in rat taste buds is dependent upon gustatory innervation.

Animals↗

Neuronal cell types and taste quality coding.

Over the past 25 years, there have been two opposing views of how taste information is represented in the activity of gustatory neurons. One view, the across-fiber pattern (AFP) theory, postulates that taste quality is represented by the pattern of activity across the afferent population. Stimuli with similar tastes produce similar patterns of activity. The other view is that activity in a few distinct neuron types codes taste quality in a "labeled-line" fashion. Neurons responding best to sucrose, for example, would represent "sweetness," and those responding best to NaCl would code "saltiness." Some of these neuron types appear to have a biological significance, such as the NaCl-best cells, which receive input about sodium stimuli exclusively from an amiloride-sensitive epithelial ion channel. However, the relatively broad tuning of these neurons makes it unlikely that they are capable of unambiguously coding information about taste quality. Rather, these neuron types play a critical role in establishing unique AFPs that distinguish among taste stimuli. The relative activity across these cell types represent taste quality, much like the patterns of activity across broadly tuned photoreceptors code information about stimulus wavelength.

Animals↗

Gender influence on emotional responses to primary tastes.

Several studies have examined the gender differences in the area of emotion, but few data are available in the field of gustation. Yet, such differences are observed in clinical manifestations of nutritional disorders, such as a greater incidence of carbohydrate cravings in women. The aim of this study was consequently to examine the influence of gender on emotional responses to taste, by comparing autonomic responses and basic emotions associated with primary tastes between men and women. Thirty-four subjects (17 males, 17 females, mean age=28) participated in the experiment. Taste stimuli were solutions of 0.3 M sucrose (sweet), 0.15 M NaCl (salty), 0.02 M citric acid (sour) and 0.00015 M quinine sulfate (bitter). Evian mineral water served as the diluent and control (neutral taste). Five autonomic parameters [skin potential (SP) and resistance (SR), skin blood flow (SBF) and temperature (ST), instantaneous heart rate (IHR)] were simultaneously and continuously recorded when subjects tasted the five solutions. The patterns of autonomic responses, obtained for each primary taste and each subject, were transcribed into one of the six basic emotions defined by Ekman (happiness, surprise, sadness, fear, anger and disgust), according to a previously described method for odorants. Results evidenced that the mean variations of each autonomic parameter did not significantly differ between men and women, even if skin resistance and cardiac responses were stronger in men than in women. Concerning the basic emotions associated with each primary taste, a similar distribution for men and women was obtained for sweet, bitter and control solutions, whereas more differences were observed for salt (P=.02) and sour solutions.

Adult↗

Anion size of sodium salts and simple taste reaction times.

Simple taste reaction times (RT) and taste intensities were measured in adult humans for 100-mM aqueous solutions of sodium chloride, acetate, glutamate, ascorbate, and gluconate flowed over the anterodorsal tongue with a closed liquid delivery system. Results from 12 subjects showed a significant increase in RT with molecular weight of the tastant, and a correlation of 0.941 between RT and the square roots of anionic weights. A multiple regression analysis controlling for perceived taste intensity indicated that RT increased linearly with the square root of the anionic weight. These findings support a model that includes both the permeability of ions through the tight junctions between the taste receptor cells of fungiform papillae taste buds and the effects of ions at apical portions of the receptor cells. They also suggest that gustatory transduction of sodium salts in humans normally involves intercellular spaces of taste buds as part of the functional sensory structures, in addition to individual taste receptor cells.

Adolescent↗

Channels as taste receptors in vertebrates.

Taste reception is fundamental for proper selection of food and beverages. Chemicals detected as taste stimuli by vertebrates include a large variety of substances, ranging from inorganic ions (e.g., Na(+), H(+)) to more complex molecules (e.g., sucrose, amino acids, alkaloids). Specialized epithelial cells, called taste receptor cells (TRCs), express specific membrane proteins that function as receptors for taste stimuli. Classical view of the early events in chemical detection was based on the assumption that taste substances bind to membrane receptors in TRCs without permeating the tissue. Although this model is still valid for some chemicals, such as sucrose, it does not hold for small ions, such as Na(+), that actually diffuse inside the taste tissue through ion channels. Electrophysiological, pharmacological, biochemical, and molecular biological studies have provided evidence that indeed TRCs use ion channels to reveal the presence of certain substances in foodstuff. In this review, we focus on the functional and molecular properties of ion channels that serve as receptors in taste transduction.

Animals↗

Ketamine blocks a taste recognition memory in fetal rats.

Decisions about novelty/familiarity are critical in determining whether or not information should be attended to, and possibly encoded, for long-term storage. We have reported that fetal and neonatal rats exhibit an increase in orofacial movements (e.g., perseverative mouthing and mouth movements, and licks) upon tasting saccharin (SAC), if it was experienced previously. E19 rat fetuses can acquire this taste recognition memory and retain it for at least 5 days (P3). In the current study, we sought to evaluate the role of N-methyl-D-aspartate (NMDA) receptors in establishing a taste recognition memory. Pregnant Sprague-Dawley rats received ketamine (NMDA receptor antagonist) (doses: 0, 50, or 100 mg/kg, i.p.). One-half hour later, we performed a reversible spinal block on each pregnant dam, and E19 fetuses received an oral injection of 10 microl, 0.3% SAC or water (control) while in utero. The uterus was replaced and the pups were later born via a normal vaginal delivery. On P3, all pups experienced oral lavage of 10 microl, 0.3% SAC, and motor responses were recorded. As expected, non-drugged control neonates tasting familiar SAC exhibited significantly more perseverative mouth movements, as well as total mouth movements and licks, than did pups tasting novel SAC. However, this taste recognition memory response was not observed in rats exposed to ketamine in utero. The data suggest that early non-associative taste memories may be disrupted by NMDA receptor blockade.

Animals↗

Area postrema lesions in rats enhance the magnitude of body rotation-induced conditioned taste aversions.

The role of the area postrema, a circumventricular organ located on the dorsal medulla in the fourth ventricle, in body rotation-induced conditioned taste aversions was examined in male hooded rats. One group of rats received lesions of the area postrema, another group was given sham lesions, and two other groups received no surgery. Three groups of rats, those with area postrema lesions, those with sham lesions, and one unoperated group, were given pairings of 1-hr access to a 0.1% sodium saccharin solution with 30 min of body rotation at 70 rpm (on a schedule of 15-sec on--5-sec off). A fourth group of unoperated rats was given pairings of access to the saccharin solution with 30 min of sham rotation. The three rotated groups all developed a taste aversion to the saccharin solution whereas the sham-rotated group did not. The group with lesions of the area postrema exhibited a greater aversion to the saccharin taste than the other two rotated groups. In a second phase of the experiment all rats were given pairing of 1-hr access to chocolate milk with injections of scopolamine methyl nitrate (1 mg/kg). The area postrema-lesioned group failed to exhibit a taste aversion to the chocolate milk whereas the other groups all showed very strong aversions. It was concluded that area postrema lesions abolish taste aversions induced by blood-borne toxins such as scopolamine methyl nitrate, but enhance taste aversions induced by motion sickness. Some speculations concerning the enhanced taste aversions induced by motion sickness in area postrema lesioned rats, are provided.

Animals↗

Deficits in conditioned heart rate and taste aversion in area postrema-lesioned rats.

Previous studies have shown that area postrema (AP) lesions cause deficits in conditioned taste aversion in the rat. They also lead to chronically lowered heart rate which can be reversed by the animals' increased appetite for and ingestion of hypertonic saline. Although not previously examined in conditioned taste aversion, changes in autonomic nervous system activity as reflected in heart rate may be an important aspect of conditioning. The present study investigated the effects of AP lesions on heart rate conditioned responses (CRs) and unconditioned responses (UCRs). Two groups of AP lesioned and sham-operated rats, one that did and one that did not drink saline solution to raise heart rate, were studied. Both LiCl and scopolamine, which have opposite effects on heart rate, were the unconditioned stimulus (UCS) agents in two separate studies. In intact rats, LiCl-mediated conditioned taste aversion was associated with decreased conditioned stimulus (CS) intake and decreased heart rate Both effects were blunted by AP lesions, although all rats displayed heart rate UCRs to LiCl. The AP rats that drank saline behaved like intact rats exhibiting both a conditioned taste aversion and conditioned heart rate responses to the CS. Although CS intake decreased, no heart rate CRs developed with scopolamine. Scopolamine-mediated conditioned taste aversion was attenuated in both saline and non-saline drinking AP-lesioned groups. Thus, when conditioned taste aversion was associated with heart rate CRs, the AP lesion-induced deficit was counteracted by saline ingestion. Conversely, when there were no heart rate CRs, conditioned taste aversion was disrupted by the lesion regardless of saline ingestion.

Animals↗

Role of the medial and lateral parabrachial nucleus in acquisition and retention of conditioned taste aversion in rats.

When ingestion of a taste stimulus is paired with internal malaise, the animal remembers the taste and rejects its ingestion thereafter. This learning is referred to as conditioned taste aversion (CTA). To establish CTA in adult male Wistar rats, 0.1% saccharin and an i.p. injection of 0.15 M LiCl were used as the conditioned and unconditioned stimuli, respectively. To elucidate the functional role of the medial part of the parabrachial nucleus (PBmed) which receives taste information and the lateral part (PBlat) which receives general visceral information, confined electrolytic lesions were made to either of these regions. Rats with bilateral lesions of the PBlat impaired the acquisition of CTA, but those lesions made after the acquisition of CTA had no effect on the retention of this learning. The bilateral lesions of the PBmed abolished the acquisition and retention of CTA. The PBlat-lesioned rats showed normal taste preference behavior, but PBmed-lesioned rats showed impaired sensibility to taste stimuli. These results suggest that both the PBlat and PBmed are essential for the acquisition of taste aversion learning, but the PBlat is not necessary for retrieval of CTA.

Animals↗

Immunocytochemical localization of serotonin and serotonin transporter (SET) in taste buds of rat.

We used an immunocytochemical approach to study the localization of serotonin and its termination system, serotonin transporter (SET), in the taste buds of rats using specific antibodies against serotonin and SET. Under confocal laser scanning microscopy, both serotonin and SET immunoreactivity were detected in the taste buds of rat vallate papillae. Serotonin immunoreactivity was seen in the spindle-shaped cells with apical processes that seemed to be light (Type II) taste cells. SET-immunoreactivity was mainly localized in the periphery or interfaces between the taste cells. Double staining studies revealed that all serotonin-containing taste cells were immunoreactive for SET, while a subclass of SET-positive cells showed serotonin immunoreactivity. These data support the hypothesis that serotonin plays a transmitter role in taste receptor cells and suggest that the serotonin-induced sensation of taste is terminated by serotonin uptake through serotonin transporter.

Animals↗

Profiling taste-motivated segments.

Early adopters of unfamiliar, but nutritious foods can do so because of a combination of taste-motivations and health-motivations. Yet because taste can provide an enduring motivation for dietary change, profiling the taste-motivated segment of a particular food might prove useful in identifying and stimulating adoption among similar predisposed segments. This manuscript describes a basic qualitative and quantitative procedure--in the context of soy consumption--that can be used to begin profiling taste-motivated segments of a particular food. A survey of 606 North Americans indicates that when contrasted to health-motivated consumers of soy, taste-motivated consumers were more likely to claim they are opinion-leaders who live with (or who are) great cooks, and they were more likely to exhibit other behaviors associated with food appreciation, such as dining out and wine consumption. In light of these findings, instead of encouraging people to eat soy solely for health reasons, a more productive method may be to target those who are more likely to prefer it for taste-motivated reasons. This same method has potential for more effectively promoting the consumption of fruits and vegetables or the consumption of genetically enhanced foods among predisposed taste-motivated segments.

Attitude to Health↗

The taste of ethanol in a primate model. II. Glossopharyngeal nerve response in Macaca mulatta.

The glossopharyngeal nerve (NG) mediates taste from the posterior part of the tongue. Here, we studied the effects of ethanol on the tongue in recordings from both the whole NG and individual taste fibers of the rhesus monkey, Macaca mulatta. The results show that the nerve activity increased at 0.7 M ethanol, reaching half maximum at around 4 M alcohol. Previously, we identified three types of taste fibers in the rhesus monkey NG: S fibers predominantly responding to sweeteners, Q fibers responding to bitter, such as quinine hydrochloride (QHCl), and M fibers responding best to monosodium glutamate, NaCl and acids [Hellekant, G., Danilova, V., & Ninomiya, Y. (1997). Primate sense of taste: behavioral and single chorda tympani and glossopharyngeal nerve fiber recordings in the rhesus monkey, Macaca mulatta. J Neurophysiol 77, 978-993]. Here, this fiber classification was used to elucidate the oral effects of ethanol and ethanol mixtures with NaCl, sucrose, citric acid and QHCl. One and three molar concentrations of ethanol stimulated all fiber types. Mixtures of ethanol with QHCl elicited a smaller response in Q fibers than did QHCl alone. In S fibers, mixtures of ethanol with sucrose gave a larger response than did sucrose alone. The variability of M fibers was too large to allow a conclusion about the effect of ethanol. These results suggest that ethanol suppresses the taste of QHCl. Similarly, the taste of sucrose might be enhanced by adding ethanol to sucrose. These effects and conclusions corroborate an earlier ethanol study of the chorda tympani (CT) nerve [Hellekant, G., Danilova, V., Roberts, T., & Ninomiya, Y. (1997). The taste of ethanol in a primate model: I. Chorda tympani nerve response in Macaca mulatta. Alcohol 14, 473-484].

Analgesics, Non-Narcotic↗

Taste reactivity in high-alcohol-sensitive and low-alcohol-sensitive rats.

High, low, and control alcohol-sensitive (HAS, LAS, CAS, respectively) rats were tested for their perception of the taste of alcohol using the taste reactivity test. Reactivity tests with a single concentration of sucrose and quinine were also done. After initial taste reactivity, all rats were tested for alcohol consumption in a standard two-bottle test (water in the second bottle). Postconsumption taste reactivity tests completed the experiment. Results indicated that HAS, LAS, and CAS rats did not differ significantly in their taste reactivity response to a range of alcohol concentrations (5-40%), nor did they differ significantly in response to sucrose or quinine. Reactivity responses were similar for each group before and after the consumption tests. Despite the lack of line differences in taste reactivity, HAS and LAS rats consumed significantly less alcohol than the CAS rats during the two-bottle access tests. The present results are in contrast to research done with rats selectively bred for alcohol consumption (Alcohol Preferring and Nonpreferring rats, High Alcohol Drinking and Low Alcohol Drinking rats), which exhibit clear line differences in patterns of reactivity changes following alcohol access. The selection phenotype of alcohol sensitivity appears to be independent of rats' behavioral response to the taste of alcohol.

Alcohol Drinking↗

The taste of ethanol in a primate model: I. Chorda tympani nerve response in Macaca mulatta.

The chorda tympani nerve (CT) mediates taste from the anterior part of the tongue. Here we studied the effects of ethanol on the tongue in recordings from both the whole CT nerve and individual taste fibers of the rhesus monkey, M. mulatta. The response to ethanol consisted of a phasic and a tonic part. At the lowest concentration tested (0.3 M) ethanol gave a response in some animals and at 0.7 M in all animals. A sigmoidal function described best the relationship between nerve response and ethanol concentrations. Hierarchial cluster analysis with 26 nonalcoholic sweet, sour, salty, and bitter stimuli had earlier identified four types of taste fibers each responding predominantly to stimuli within one of the four human taste qualities. Here were found that ethanol stimulated all sweet-best fibers and at high concentration some salt-best fibers, but never any acid-best and bitter-best fibers. This may explain the sweet taste attributed to low ethanol concentration by humans. Further, in mixtures it suppressed the responses in acid-best and bitter-best taste fibers. This may partly explain the effects of ethanol on sour and bitter taste in alcoholic beverages.

Animals↗

Shh and Ptc are associated with taste bud maintenance in the adult mouse.

In mammals, taste receptor cells are organized into taste buds on tongue. Taste buds are trophically maintained by taste neurons and under continuous renewal, even in adults. We found that the receptor for Sonic hedgehog (Shh), Patched1 (Ptc), was expressed around taste buds where cells were proliferating, and that Shh was expressed within basal cells of taste buds. Denervation caused the loss of Shh and Ptc expression before the degeneration of taste buds.

Animals↗

An in vitro model for the study of taste papillae morphogenesis using branchial arch explants.

It is generally accepted that innervation is required for the maintenance of taste papillae and taste buds, but it is not entirely clear what role, if any, innervation plays in papillae and taste bud formation. Events in taste papillae formation and differentiation take place almost entirely in utero and, therefore, the study of the role of innervation in these events requires a suitable in vitro model. In the past, investigators have made use of various culture techniques to study mammalian taste papillae development in vitro and the role of innervation in this process with varying success. All of these models examined papillae development in isolated tongue or tongue fragments and have lacked the ability to manipulate the innervation of developing taste papillae in these explants. We have established a protocol for an in vitro model of taste papillae morphogenesis using branchial arch explants and roller tube culture methodology. Our results demonstrate that this model supports the morphogenesis of the circumvallate papilla with an integrated nerve. In addition, the use of branchial arch explants allows the inclusion or exclusion of geniculate and petrosal ganglia to examine directly the effects of the presence or absence of innervation on papillae formation and maintenance.

Animals↗

The genetics of tasting in mice. VII. Glycine revisited, and the chromosomal location of Sac and Soa.

Previous work which appeared to show that some strains of mice taste glycine solutions as bitter has been found to be in error. The bitterness came from copper glycinate which formed in the brass drinking spouts. Taste testing with copper glycinate shows that the genetical data identifying the gene Glb are still valid. The close linkage of Glb and Rua has been confirmed. Most strains of mice prefer glycine solution to water, presumably because the glycine tastes sweet. The degree of preference for glycine is correlated with the degree of preference for other sweet substances such as saccharin or acesulfame. The gene dpa appears not to be involved. The sweetness tasting gene Sac has been mapped to chromosome 4 at 8.1 +/- 3.4 cM distal to Nppa (formerly Pnd). The bitterness tasting gene Soa is very closely linked to Prp on chromosome 6 (no recombinants among 67 backcross progeny). It is suggested that the sweetness and bitterness tasting genes have descended from a common ancestral tasting gene which existed before the tetraploidization of the genome which took place in early vertebrate evolution.

Animals↗