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Taste responses in the parabrachial pons of ovariectomized rats.

Since the early 1970s it has been known that female rats prefer higher concentrations of sweet stimuli compared with males. Recent data have revealed that electrophysiological responses to sweet tastes recorded in the parabrachial nucleus of the pons (PbN), the second relay in the neural pathway for taste, are larger in diestrus female rats compared with those in males. Because it has been shown that ovariectomized rats have lowered preferences for saccharin compared with intact females, it is possible to predict that responses to sweet stimuli in the PbN of ovariectomized rats might be smaller than those in intact females. To investigate this hypothesis, electrophysiological responses to representatives of the 4 basic taste qualities, i.e., salty, sour, sweet and bitter, were recorded in the PbN of ovariectomized rats. Gustatory stimuli included NaCl (0.1 M), HCl (0.01 M), sucrose (0.5 M), quinineHCl (0.01 M) and Na-saccharin (0.004 M). Comparison of taste responses of 49 PbN units in ovariectomized rats with those in the PbN of intact female and male rats showed that responses to sweet stimuli were of comparable magnitude to those in female rats but elevated compared with those in male rats. However, responses of PbN units to quinine appeared to be larger in ovariectomized rats compared with those in the PbN of both intact female and male rats. These results suggest that decreased saccharin preference in ovariectomized rats may reflect a greater sensitivity to the bitter components of the taste of saccharin. Moreover, these data provide evidence that taste responses in the PbN are influenced by both the activational and organizational actions of ovarian hormones.

Animals↗

Multichannel taste sensor using electric potential changes in lipid membranes.

A taste sensor with a multichannel electrode was developed by using lipid membranes as a transducer of taste substances. The sensor can detect the taste in a similar manner to the human gustatory sensation by response patterns of electric potential to taste substances. The sensitivity, reproducibility and durability were superior to those of humans. The same taste as that elicited by some commercial aqueous drink was reproduced by making aqueous solution mixed from four kinds of basic taste substances, the concentrations of which were determined so that the electric-potential pattern of this mixed solution could agree well with the pattern by the drink. Different brands of beer were easily distinguished by the electric-potential patterns.

Biosensing Techniques↗

Application of a taste evaluation system to the monitoring of Kimchi fermentation.

As an objective method, taste evaluation with an instrument is able to supplement the subjective sensory evaluation and to be applied to the optimization of food processing. Kimchi, a Korean traditional pickle fermented with lactic acid bacteria, is expanding its consumption worldwide. The fermentation control of it has been routinely done by measuring titratable acidity and pH. In this study, an eight-channel taste evaluation system was prepared, followed by an application to the monitoring of Kimchi fermentation. Eight polymer membranes which individually responded to cationic or anionic substances were prepared by mixing electroactive materials such as tri-n-octylmethylammonium chloride, bis(2-ethylhexyl)sebacate as the plasticizer and polyvinyl chloride in the ratio of 1:66:33. Each membrane prepared was separately installed onto the sensitive area of an ion-selective electrode to produce the respective taste sensor. The eight-channel sensor array and a double junction reference electrode were connected to a 16-channel high input impedance amplifier. The amplified sensor signals were stored to a personal computer via a multi-channel A/D converter. Two sensor groups composed of the cation-selective and anion-selective polymer membrane electrodes showed characteristic concentration-dependency to various artificial taste substances. As a whole, the response potentials of the sensor array increased during the fermentation period at 4, 10 and 25 degrees C. Even the response potentials of the anion-selective taste sensors slightly increased possibly due to the protonation of anions by liberated H+ ions, thereby leading to a decrease in the anion concentration. When the signal data were interpreted by principal component analysis (PCA), the first PC at 4 degrees C explained most of the total data variance. A close correlation was found between the values of titratable acidity and the first PC, which indicated a possible applicability of the multi-channel taste sensor of this study to the process monitoring of various pickle.

Biomimetics↗

Effects of midazolam on the expression of conditioned taste aversion in rats.

In conditioned taste aversion (CTA), the animals learn to avoid a taste substance (conditioned stimulus, CS) which was previously associated with visceral distress (unconditioned stimulus, US). The present study examined the effects of administration of midazolam (MDZ), a benzodiazepine agonist, after the acquisition of CTA on the expression of CTA. After ingestion of 0.5 M sucrose (CS) was paired with an intraperitoneal (i.p.) injection of 0.15 M LiCl (US), control rats showed strong CTA to the CS. However, a systemic injection of MDZ (1.5 mg/kg, i.p.) before the retention test prevented conditioned animals from rejecting the CS, but in the subsequent retention tests where the drug was not administrated, those animals again showed strong aversions to the CS. Aversive taste reactivity patterns to the intraorally infused sucrose and 0.3 M dl-alanine in the conditioned animals were also diminished by the similar injection of MDZ, but not by a serotonergic anxiolytic agent, buspirone (2.5 or 5.0 mg/kg, i.p.). General taste sensory deficit might not be induced by MDZ because the drug injection did not impair conditioned aversions to 0.2 M NaCl and 0.01 M HCl. Infusion of MDZ into the basolateral nucleus of the amygdala (BLA) also attenuated conditioned aversions to sucrose. These results suggest that systemic or intra-BLA administrations of MDZ impair the expression of CTA selectively to sweet-tasting substances, implying that a transient MDZ-induced CTA expression deficit is due to the enhancement of palatability of CSs with preferable tastes rather than general anxiolytic or amnesic effects of MDZ.

Amygdala↗

Orbitofrontal cortex: neuronal representation of oral temperature and capsaicin in addition to taste and texture.

The primate orbitofrontal cortex is a site of convergence of information from primary taste, olfactory and somatosensory cortical areas. We describe the discovery of a population of single neurons in the macaque orbitofrontal cortex that responds to the temperature of a liquid in the mouth. The temperature stimuli consisted of water at 10 degrees C, 23 degrees C, 37 degrees C and 42 degrees C. Twenty-six of the 1149 neurons analyzed (2.3%) responded to oral temperature. The tuning profiles of the neurons to temperature showed that some of the neurons had graded responses to increasing temperature (27%), others responded to cold (10 degrees C) stimuli (27%), and others were tuned to temperature (46%). The neuronal responses were also measured to taste stimuli, viscosity stimuli (carboxymethyl-cellulose in the range 1-10,000 cP), and capsaicin (10 microM). Of 70 neurons with responses to any of these stimuli, 7.1% were unimodal temperature; 11.3% were temperature and taste-sensitive; 7.1% were temperature and viscosity-sensitive; and 11.3% were temperature, taste and viscosity sensitive. Capsaicin activated 15.7% of the population of responsive neurons tested. These results provide the first evidence of how the temperature of what is in the mouth is represented at the neuronal level in the orbitofrontal cortex and the first evidence for any primate cortical area that in some cases this information converges onto single neurons with inputs produced by other sensory properties of food, including taste and texture. The results provide a basis for understanding how particular combinations of oral temperature, taste, and texture can influence the palatability of foods.

Action Potentials↗

Effects of drugs on olfaction and taste.

The fact that so many varied medications reportedly affect taste and smell is a testament to the complexity of the gustatory and olfactory systems. The reception, transduction, propagation, and perception of a chemical tastant or odorant requires the effective operation of numerous mechanisms--all of which may be susceptible in one way or another to a prescribed medication. Just as a diuretic may block the apical ion channels on a taste bud, or an antifungal can inhibit cytochrome p450-dependent enzymes at the level of the receptors, a chemotherapeutic agent can destroy mitosis in a replicating receptor cell and a steroid can lead to candidal overgrowth on the tongue surface. Medications not only have a perceivable taste themselves at times, but they can alter the mechanisms responsible for the ultimate perception of tastes and smells--either by direct or secondary means. It should be emphasized, as noted earlier in this article, that while many medications are to blame for the impairment or distortion of the gustatory or olfactory systems, it is not uncommon that the underlying medical problem for which they are prescribed is actually the culprit. Examples include epilepsy, migraines, hypothyroidism, schizophrenia, infections, and cancer. In fact, simple partial seizures emanating from regions of the brain such as the amygdala, hippocampus, parietal operculum, and rolandic operculum can lead to the chemosensory sensations that are most commonly considered unpleasant, such as "rotten apples," "cigarette," "peculiar," or "vomitus". While removing or changing an offending medication can reverse the effects on smell or taste perception, it is important to remember that lasting impairment may occur. This is vital for a physician to recognize prior to prescribing a medication. It is also necessary to report this to patients who may be devastated by chemosensory alterations after starting a new medication (eg, pastry chef, perfumist, wine specialist, plumber). Among the "risks" in a risks/benefits discussion with a patient regarding the use of a new medication, alterations in olfaction and taste appear to play an increasingly recognized role.

Drug-Related Side Effects and Adverse Reactions↗

Further evidence for conditioned taste aversion induced by forced swimming.

A series of experiments with rats reported that aversion to a taste solution can be established by forced swimming in a water pool. Experiment 1 demonstrated that correlation of taste and swimming is a critical factor for this phenomenon, indicating associative (i.e., Pavlovian) nature of this learning. Experiment 2 showed that this learning obeys the Pavlovian law of strength, by displaying a positive relationship between the duration of water immersion in training and the taste aversion observed in subsequent testing. Experiment 3 revealed that swimming rather than being wet is the critical agent, because a water shower did not endow rats with taste aversion. Experiment 4 found that taste aversion was a positive function of water level of the pools in training (0, 12 or 32 cm). These results, taken together, suggest that energy expenditure caused by physical exercise might be involved in the development of taste aversion.

Analysis of Variance↗

Effects of inorganic constituents of saliva on taste responses of the rat chorda tympani nerve.

The effects of saliva on the taste responses of the chorda tympani nerve to the 4 standard chemical stimuli (sucrose, NaCl, HCl, and quinine hydrochloride) and water were investigated in anesthetized rats. When the tongue was adapted to pilocarpine-stimulated whole saliva (pH 8.7), the magnitude of neural response to sucrose was about 2 times that obtained when the tongue was adapted to distilled water. Under saliva-adapted conditions, the magnitude of responses to other taste stimuli was reduced by 10-30%, and the water response appeared. These changes were dependent on the concentration of electrolytes (Na+, K+, Cl-, and HCO3-) and on the pH of the saliva. When the tongue was adapted to 10-30 mM NaHCO3 (pH 8.4-8.6), taste and water responses were similar to those under saliva-adapted conditions. Single fiber analyses revealed that the enhancement of the sucrose response after adaptation to NaHCO3 was produced by an increased overall activity of sucrose-responsive fibers. The correlation coefficients of the magnitude of the taste responses between the 4 taste stimuli remained unchanged, but the water response showed a high correlation to HCl and quinine hydrochloride responses after adaptation. Possible mechanisms for the effects of saliva on taste and water responses were discussed.

Animals↗

Conditioned taste aversion: modulation by 5-HT receptor activity and corticosterone.

Two experiments were designed to elucidate the involvement of the hypothalamic-pituitary-adrenal axis and the 5-hydroxytryptamine (5-HT) system in the acquisition of lithium chloride-conditioned taste aversion. In Experiment 1, rats were administered either vehicle or 50 mg/kg nefazodone daily for 4 weeks. Rats were treated with 22 mg/kg of lithium chloride in order to produce conditioned taste aversion to a sucrose solution. Three days later, nefazodone completely blocked the lithium chloride-conditioned taste aversion. In Experiment 2, the effects of chronic corticosterone administration on lithium chloride-conditioned taste aversion were investigated. Twenty male rats received either corticosterone at a dose of (50 mg/kg) or vehicle injections over a period of 14 consecutive days. Lithium chloride-conditioned taste aversion was potentiated in rats treated with corticosterone. Additionally, corticosterone-treated animals required more trials to reach extinction. These results suggest the involvement of both the 5-HT system and the hypothalamic-pituitary-adrenal axis in lithium chloride-conditioned taste aversion.

Administration, Oral↗

Taste preference and acceptance in thirsty and rehydrated [correction of dehydrated] rats.

Experiments were designed to determine whether water deprivation affects taste preferences and/or taste acceptance. In experiment 1, both five- and two-bottle preference tests were performed in normally hydrated rats to permit the selection of five groups of rats showing the same taste preference for one of four prototypical tastes. Subsequently, in the same groups of rats, taste preferences were determined by a two-bottle test (experiment 2), and taste acceptance by a one-bottle test (experiment 3), following 12, 24, 36, and 48 h of water deprivation. After both 12 and 24 h of dehydration, during the first 10 min of the tests of experiment 2, all rats ingested greater volumes of either NaCl or sucrose solution than water, but more water than either HCl or quinine solution, and the differences were very significant (P<.0001). After 36 or 48 h of dehydration, the differences became very small and, in some cases, the P-values were at the lowest or borderline level of the significance, suggesting that dehydrated rats poorly discriminate the nature of the fluid drunk. During the 11-60 min interval, all rats preferred either sucrose or NaCl to water, but water to either HCl or quinine. Experiment 3 was performed to ascertain whether the need for fluid might overcome the palatability of solutions. All rats, dehydrated for 36 or 48 h, after 10 min of exposure, drank equal amounts of fluid, independent of its palatability. During the 11-20 and 21-60 min interval, the fluid intake of rats changed in accordance with the palatability of the solution available. In conclusion, severe thirst in rats may override the palatability of the solutions, and the thirst drive may be so strong that they do not reject fluids because body fluid balance would be severely compromised.

Animals↗

Acute anosmia in the mouse: behavioral discrimination among the four basic taste substances.

The importance of taste and smell in discrimination of tastes was examined in normal and anosmic mice. We studied the influence of olfaction on taste sensation using behavioral and electrophysiological methods in both normal animals and animals made anosmic mice by destroying their olfactory receptor cells with zinc sulfate (ZnSO(4)) solution. Electrophysiological responses from chorda tympani nerves showed that peripheral taste receptor cells transmitted taste signals normally to the central nervous system, even when the olfactory senses were abnormal. Behavioral observations showed that mice with abnormal olfaction could not differentiate tastes.

Animals↗

Estrogen-induced suppression of intake is not mediated by taste aversion in female rats.

Estrogen treatment can suppress the intake of a previously presented gustatory conditioned stimulus (CS). This finding has been interpreted as an estrogen-induced conditioned taste aversion. However, a distinction must be made between taste aversion and taste avoidance. In particular, tastes are only considered aversive if they elicit a stereotypic behavioral response, otherwise the reduction in intake is classified as an avoidance. Although aversive orofacial responses have been reported in male rats after taste-estrogen pairings, they have not been examined in ovariectomized female rats. The goal of the present investigation, then, was to use similar procedures to determine whether conditioned aversion also mediates the estrogen-induced reduction of intake in female rats. Animals were introduced to a novel 0.1% saccharin solution and immediately thereafter were given a subcutaneous injection of vehicle or estradiol benzoate (10 microg). Responses were assessed using a two-bottle preference test, a one-bottle acceptance test, and a taste reactivity (TR) test. The results confirmed previous reports of a reduced preference for saccharin after saccharin-estradiol pairing using the two-bottle test. The reduction in intake during the one-bottle test, however, was not accompanied by stereotypic aversive responses, such as gaping. Surprisingly, a similar reduction in intake also occurred when using a backward conditioning procedure in which estrogen was injected before, rather than after, CS access. Thus, the present results show that the suppressive effects of estrogen reflect an avoidance, rather than aversion and, moreover, that the reduced intake may be due to an unconditioned, rather than a conditioned, response.

Animals↗

Changes in taste and satiety in dietary-restrained women following stress.

Stress has been shown to increase food consumption in women with high levels of dietary restraint (restrainers) but decrease consumption in nonrestrainers. The present study was designed to replicate the differential eating pattern following stress and measure physiological and subjective responses to food over repeated taste presentations. Restrained (N = 16) and nonrestrained (N = 16) women were given eight taste presentations of a food followed by an ad lib taste test. Between taste trials, half of the restrainers and nonrestrainers performed a variation of the Stroop stressor while remaining subjects sat quietly. Salivation to the food cue was measured at each trial as well as ratings of food liking, hunger, fullness, and arousal. Results showed significant effects of restraint on food liking and stress condition on hunger. Restrainers increased liking ratings over taste trials whereas ratings for nonrestrainers increased and then decreased to baseline levels by the last trial (p = 0.05). Nonstressed subjects showed an increase in hunger ratings, whereas ratings for stressed subjects did not show any stable directional pattern (p = 0.04). Salivary responses decreased for all groups (p = 0.01). A significant interaction of restraint by stress for intake was found (p = 0.03); restrainers increased consumption following stress whereas nonrestrainers decreased consumption as compared with controls. The data suggest that the Stroop task can influence intake in restrainers, but the changes in intake did not directly correspond to the changes observed during the presentations of the taste cues and stressor.

Adolescent↗

Anesthesia of the chorda tympani nerve and taste phantoms.

The chorda tympani nerve innervates the anterior two-thirds and the glossopharyngeal nerve, the posterior one-third, of each side of the tongue. Previous work showed that anesthesia of one chorda tympani increased the perceived intensity of quinine applied to an area innervated by the contralateral glossopharyngeal nerve, but decreased the perceived intensity of NaCl applied to an area innervated by the ipsilateral glossopharyngeal nerve. The data presented here corroborate that earlier finding and show that if both chorda tympani nerves are anesthetized, the taste of quinine is intensified and the taste of NaCl diminished at areas innervated by the glossopharyngeal on both sides of the tongue. In about 40% of the subjects, tastes occurred in the absence of stimulation (we call these tastes phantoms). The phantoms were usually localized to the posterior tongue contralateral to the anesthesia and they were abolished when a topical anesthetic was applied to the area where they were perceived. Phantoms like these may be a source of clinical dysgeusia in patients with localized taste damage. The phantoms may result because the anesthesia releases inhibition normally occurring between the central projection areas of different taste nerves.

Adult↗

Leptin and sweet taste.

Leptin, the product of the obese (ob) gene, is a hormone primarily produced in adipose cells, and also at smaller amounts in some other peripheral organs. It regulates food intake, energy expenditure, and body weight. Leptin is thought to promote weight loss, at least in rodents, by suppressing appetite and stimulating metabolism. Mutant mice that lack either leptin or functional leptin receptors, such as ob/ob and db/db mice, are hyperphagic, massively obese, and diabetic. Central hypothalamic targets are mainly responsible for the effects of leptin on food intake and weight loss. However, there are also direct effects on peripheral tissues. Recently, the taste organ was found to be one of the peripheral targets for leptin. The hormone specifically inhibits sweet taste responses in lean mice and not in db/db mice. Thus leptin appears to act as a modulator of sweet taste, provided a functional leptin receptor is expressed by the taste cells. This chapter reviews the genetics and molecular biology of leptin and its receptors, the receptor mechanisms for sweet taste, the modulating action of leptin on taste receptor cells, and the consequences for the regulation of food intake.

Animals↗

Intragastric copper sulfate produces a more reliable conditioned taste aversion in vagotomized rats than in intact rats.

Although copper sulfate is an emetic stimulus, preliminary experiments failed to obtain a taste aversion in intact rats following intragastric administration as had been previously reported in the literature. Several experiments were therefore run to further investigate the capacity of intragastric copper sulfate to function as an unconditioned stimulus for taste aversion learning and the role of the vagus in mediating that learning. The results of the first series of experiments showed that intragastric administration of copper sulfate (5 mg/kg X 5H2O) was more effective in reliably producing a taste aversion in vagotomized rats than in sham-operated control rats. The second experiment examined the effects of area postrema lesions on the acquisition of a taste aversion produced by intragastrically administered copper sulfate in vagotomized rats. The results indicated that the taste aversion observed following treatment with intragastric copper sulfate in vagotomized rats could be prevented by lesions of the area postrema. The present results indicate that intragastric administration of copper sulfate is a more reliable unconditioned stimulus for taste aversion learning in vagotomized rats than in intact rats. It is not certain what factors might account for the discrepant results between the present experiments and previously published research.

Animals↗

A method for in-situ tight-seal recordings from single taste bud cells of mice.

In order to investigate taste transduction mechanisms, we developed a method to irrigate the receptor and basolateral membranes of mouse taste bud cells with different solutions under tight-seal recording conditions. A peeled tongue epithelium with taste bud cells was mounted on a recording platform designed to separate irrigating solutions for each membrane. The mucosal surface (receptor membrane side) of the peeled epithelium facing an inner chamber was always irrigated with deionized water or stimulating solutions, and the serosal surface (basolateral membrane side) was irrigated with a physiological saline solution. A recording electrode was placed on the basolateral membrane of a taste bud cell under an upright-microscope with a x 40-water-immersed objective. Investigated taste bud cells generated action potentials, and 1 M glucose, 200 mM NaCl, and 10 mM quinine elicited inward current. Irrigation with deionized water for more than 1 h had no effect. The resistance of the peeled tongue epithelium was 1570 +/- 343 omega cm2. These results show that the peeled tongue epithelium protects basolateral membranes from deionized water or stimulating solutions as the tongue epithelium does in situ and that this method is suitable to investigate the role of each membrane in taste transduction.

Action Potentials↗

Co-expression of calcium signaling components in vertebrate taste bud cells.

In order to investigate the molecular mechanism of calcium signaling pathways common to the vertebrate gustatory systems, we have analyzed the expression of their molecular components. We first identified a phospholipase C (PLC) beta subtype expressed in the taste buds of pond loach (Misgurnus anguillicaudatus), designated DPLCbeta2, which is closely related to mammalian PLCbeta2 shown recently to be expressed in rat taste buds. The taste bud-specific expression of PLCbeta2 in a fish species as well as rat strongly suggests that PLCbeta2 mediates the tastant-induced second messenger response in taste buds, which is common to vertebrates. Next, we examined the correlation of gene expression of the candidate components leading to PLCbeta2 activation in rat circumvallate papillae, including G proteins, G(i2) and gustducin, and a G protein-coupled receptor, TR2. As a result, it was shown that the mRNAs for PLCbeta2 and G(i2) co-exist in the same cells, and PLCbeta2- and G(i2)-positive cells include both gustducin-positive cells and TR2-positive cells. However, no correlation was found between the expressions of TR2 and gustducin as reported previously. Our results thus indicate that a taste transduction pathway comprising TR2, G(i2) and PLCbeta2 occurs in a subset of taste cells.

Animals↗