Learned aversions and taste qualities in hamsters

Interralations among taste perceptions in gloden hamsters (Mesocricetusauratus) were examined using generalizaions of learned tasteaversions. If stimulus A is avoided given a taste aversion hasbeen established to stimulus B, and vice versa, A and B ‘cross-generalize’.Stimuli within five groups cross-generalized. The groups ofcompounds were (i) sweeteners (fructose, saccharin, sucrose);(ii) sodium salts (NaCl, NaNO₃, Na₂SO₄): (iii) non-sodium salts(KCl, MgSO₄ NH₄Cl) plus quinine HCl; (iv) acids (acetic, hydrochloric,citric); and (v) urea. Only two pairs of stimuli from differentgroups cross-generalized (HCl—NH₄Cl. quinine HCl—urea).Neural patterns of response recorded form chorda tympam nervefibers in hamsters suggest that taste receptors on the anteriortongue distinguish among three groups of taste stimuli: sweeteners,sodium salts, and a group including non-sodium salts, acids,quinine HCl and urea. Neurons innervating other taste fieldsare likely to provide the information that hamsters use to discriminateamong the tastes of non-sodium-salt and non-sweetener stimuli.     

Decline of IXth nerve taste responses following nerve transection

Summated impulse discharges to taste solutions were recordedfrom intact and transected IXth nerves in the Mongolian gerbil(Meriones unguiculatus). Five taste stimuli were used: 0.3 MNH₄Cl, 0.3 M NaCl, 0.01 M HCl, 0.01 M quinine hydrochloride,and 0.5 M sucrose. 0.3 M NH₄Cl was the most effective stimulus.Taste responses from intact nerves were stable for more than10 hours. Following IXth nerve transection, the peak summatedresponse to 0.3 M NH₄Cl declined by 50% in a mean of 119 min.(Some animals failed to show this taste response decline inthe winter months.) The transected IXth nerve's spontaneousactivity and responses to other taste solutions also typicallydeclined. The continued presence of normal compound action potentialsindicated that the transection-induced decline in taste responsesdid not result from a failure of impulse propagation mechanismsin the nerve trunk. The results are consistent with the propositionthat transection interferes with axonal transport of materialsvital to the short-term maintenance of taste responses.     

The sweet-bitter taste of alcohol: aversion generalization to various sweet-quinine mixtures in the rat

Rats were trained to avoid a 5% alcohol solution and then testedwith either sweet + quinine hydrochloride solutions (Experiment1) or sweet + hydrochloric acid solutions (Experiment 2). Thesweet stimuli used were sucrose, glucose, fructose and saccharin.Significant aversion generalization was found only in Experiment1 where trained rats generalized to all four test stimuli, thussuggesting that alcohol has a sweet taste (in combination withbitter) not specific to one sweetener. No significant aversiongeneralization was noted in Experiment 2 when sweet + hydrochloricacid solutions were tested. In Experiment 3, rats were trainedto avoid 6% alcohol and tested with sucrose + quinine hydrochloridemixtures with varying concentrations of each component. In general,rats showed generalization of the alcohol aversions across thevarious concentrations of sucrose and quinine hydrochloridetested.     

Parabrachial unit activities after the acquisition of conditioned taste aversion to a non-preferred HCl solution in rats

Abstract In a behavioral experiment, rats reliably acquired a taste aversion to non-preferred 0.01 M HCl that had been previously paired with intraperitoneal injection of 0.15 M LiCl. These rats showed aversions to other acidic solutions such as malic acid and tartaric acid. In a neurophysiological experiment, the neuronal activities of the parabrachial nucleus (PBN) were recorded after the acquisition of conditioned taste aversion (CTA) to 0.01 M HCl in urethane-anesthetized rats. Neuronal responses to the conditioned stimulus (CS) did not change on the whole but decreased in the dorsal region to the brachium conjunctivum. The proportion of HCl-best to NaCl-best units was lower in the CTA group than in controls. The spontaneous firing rate was lower in the CTA group than in controls. Correlation coefficients between the HCl CS and normally preferred tastes (sucrose and NaCl) were more negative and those between HCl and quinine were more positive in the CTA group than in the controls. These results may be explained by the notion that gustatory responses of PBN neurons are concerned with alterations in taste hedonics after the acquisition of conditioned taste aversions.     

Generalization of specific taste aversions to alcohol in the rat

Two experiments were conducted to examine the taste qualitiesof alcohol in the rat. In Experiment 1, rats were trained toavoid sucrose, quinine or a sucrose + quinine mixture. In Experiment2, rats were trained to avoid sucrose, hydrochloric acid ora sucrose + hydrochloric acid mixture. In both experiments ratswere then tested for generalization to 3, 6 and 9% (v/v) alcoholsolutions. Following the alcohol tests rats were tested withthe taste solutions used during training. Results showed thatrats trained to avoid sucrose (Experiments 1 and 2), quinineor the sucrose + quinine mixture generalized that aversion tothe 6% alcohol solution. No generalization was found to the3% alcohol solution. No generalization was displayed by ratstrained to avoid either the hydrochloric acid solution or thesucrose + hydrochloric acid solution. Following the alcoholtests, all trained rats exhibited strong aversions to the solutionthey were trained to avoid. In addition, rats trained to avoida single solution generalized that aversion to the mixture andrats trained to avoid a mixture generalized to the single components.     

The significance of apical K+ channels in mudpuppy feeding behavior

Behavioral experiments were conducted to determine the effectsof known potassium channel blockes, minnow extract (a naturalfood substance), and otber taste stimuli on feeding behaviorin the mudpuppy (Necturus maculosus). Groups of 10–15mudpuppies were presented with one or more of the followingchemicals dissolved in gelatin cubes: H₂O (control), NaCl (1.0M; 0.2 M), crude minnow extract (0.1 g/ml), CaCl₂ (1.0 M), citricacid (0.1 M; pH2, 0.01 M; pH 3), quinine HCl (0.1 M), tetraemylammoninmchloride (TEA; 0.1 M), KCl (1.0 M) and an amino acid mixturecontaining L-phenylalanine, L-arginine, L-histidine, L-valine,L-glutamic acid and L-lysinc at concentrations of 0.02 M and0.04 M each. Crude minnow extract was centrifuged and separatedinto components consisting of the following fractions: >500D, >1000 D and >14 000 D, each of which was presentedto mudpuppies in cubes. Results were expressed as the percentageof animals rejecting the cube within 5 min of taking the cubeinto the oral cavity. Statistical analysis using the Fisherexact test indicated that cubes containing chemicals known toblock (CaCl₂, TEA, citric acid and quinine) or permeate (KCl)apical K⁺ channels in mudpuppy taste cells were more aversivethan control cubes, and aversion was concentration-dependentIn contrast, cubes containing minnow extract and componentsof minnow extract < 14 000 D were significantly preferredover control cubes. Cubes containing a mixture of minnow extractand an aversive chemical were significantly less aversive thancubes containing only an aversive chemical. These findings suggestthat activation of the apical K⁺ conductance in mudpuppy tastecells triggers an aversive response, but the aversion can beameliorated by low molecular weight compounds present in minnowextract.     

Taste responses of the gerbil IXth nerve

Summated taste responses to 12 taste solutions were recordedfrom the IXth (glossopharyngeal) nerve of 38 Mongolian gerbils(Meriones unguiculatus). 0.3 M NH₄Cl was the most effectivestimulant. The relative magnitude of the peak summated responsewas a positively accelerated function of log molar concentration.Absolute thresholds were determined for three chemicals: 0.002M NaCl, 0.0003 M HCl, and 0.002 M sucrose. The relative magnitudesof the responses to quinine, NH₄Cl, and KCl were greater forthe IXth nerve than for the chorda tympani nerve, whereas NaClwas more effective for the chorda tympani. A similar patternis seen in the rat. Acetic and citric acid may bind to commonreceptor sites. NH₄Cl, KCl, and HCl may also have receptor sitesin common.     

Amiloride does not block taste transduction in the mudpuppy, Necturus maculosus

Activity of the glossopharyngeal nerve was recorded with bipolarsilver wire electrodes while taste stimuli were applied to thelingual surface in anesthetized mudpuppies. Taste stimuli wereinjected into a continuous stream of distilled water which wasrunning over the tongue, KCl, CaCl₂ and LiCl₂ at 0.4 M elicitedbrisk responses, as did HCl at 0.2 M and quinine at 6 x 10^–4M. Sucrose, glucose and saccharin did not elicit responses.Twenty amino acids were surveyed for their ability to evokea response at 0.04 M: 1-arginine, 1-valine, 1-phenylalanine,1-tryptophan, 1-tyrosine, 1-glutamic acid, 1-lysine and histidinealways evoked responses, whereas other amino acids either didnot evoke responses or only occasionally evoked responses. Thesupernatants from solutions of minced worms and minnows andPurina Trout Chow were effective taste stimuli. Pre-adaptingthe tongue to Ringer's solution by running a continuous streamof Ringer's solution over it eliminated responses to quinineand decreased responses to NaCl. Pre-adapting the tongue to10^–4 to 10^–3 M amiloride, a potent sodium channelblocker, did not alter the responses to NaCl, LiCl, or othertaste stimuli.     

The taste of alcohol for rats as revealed by aversion generalization tests

In six experiments, naive rats were trained to avoid alcoholby pairing its presentation with lithium chloride-induced illness.Rats were then tested for aversion generalization by presentingvarious test solutions. Rats trained to avoid either 3, 6 or9% (v/v) alcohol generalized the aversion to a sucrose + quininehydrochloride solution. The five remaining binary combinationsof sucrose, sodium chloride, hydrochloric acid and quinine hydrochloridefailed to produce significant generalization in trained rats(Experiment 1). In further experiments, rats trained to avoid6% alcohol showed significant aversion generalization to a varietyof sucrose + other ‘bitter’ solutions (Experiment2) and sucrose + acid solutions (Experiment 3). Varying theconcentration of hydrochloric acid in a sucrose + acid mixtureproduced small but uniform degrees of aversion generalization(Experiment 4). Rats trained to avoid 6% alcohol did not generalizethe aversion to sucrose alone, regardless of concentration (Experiment5). Finally, in Experiment 6, rats trained to avoid 6% alcoholsuppressed consumption over a range of alcohol concentrations.These results confirm that, for rats, the taste of alcohol hasa complex set of characteristics; sweet taste in combinationwith other tastes appears to be the most similar as it is tothese solutions that rats with alcohol aversions show the mostgeneralized avoidance.     

Behavioural and electrophysiological experiments on the sense of taste in Saguinus midas tamarin (Callitrichidae) (author's transl)]

The activity in the gustatory nerve from the anterior part of the tongue, the chorda tympani proper nerve, has been recorded during stimulation of the tongue of a New World monkey, Saguinus midas tamarin. A series of 0.3 M sugars and 0.07 M NaCl, 0.07 M sucrose, 0.004 M acetic acid and 0.00005 M quinine hydrochloride were used as taste stimuli. The concentrations of the last four stimuli were the same as those earlier found to be the lowest concentrations at which this monkey in behavioural experiments discriminates between them and water. The records showed that these threshold concentrations all elicited a neural response. Further, the neural activities during stimulation with the series of 0.3 M sugars were recorded. Using the amplitudes of the responses as a measure, the order between them was found to be fructose greater than glucose greater than lactose greater than arabinose greater than sucrose = galactose greater than raffinose. This order was discussed and related with the order found in behavioural experiments. With both methods, fructose seemed to be the strongest stimulus, but then the order among the sugars differed.     

Bitter taste intensity: the effect of tastant and thiourea taster status

Thiourea tasters and nontasters did not significantly differin their ratings of intensity of caffeine or quinine HCl, butsignificant group x concentration interactions were noted fordenatonium benzoate, sucrose octaacetate and urea.     

Neural responses and aversion to bitter stimuli in rats

Gustatory responses of rats to quinine, nicotine, caffeine,MgCl₂ and bitter peptides such as glycyl-L-phenylalanine (Gly-Phe)and glycyl-L-isoleucine (Gly-lle), were studied by recordingintegrated responses to bitter stimuli from the chorda tympaniand IXth nerve as well as by measuring fluid intakes with atwo-bottle choice method. In addition, the effect of L-glutamyl-L-glutamicacid (Glu-Glu), which has an ability of masking bitterness inhumans, on the IXth nerve response was examined. Quinine, nicotine and caffeine elicited aversive behavior atconcentrations similar to those eliciting the IXth nerve responses,while MgCl₂ induced aversion above 0.1 M although it producedneural responses at much lower concentrations in both nerves.No aversion was produced by the bitter peptides at 30 mM, althougha small response was elicited by Gly-Phe in the chorda tympaniand by Gly-Ile in the IXth nerve. Responses in the IXth nerveto mixtures of Glu-Glu with bitter substances was smaller thanthe sum of those for single solution alone, indicating depressionof nerve responses to bitter stimuli by Glu-Glu.     

Differences in the taste quality of maltose and sucrose in rats: issues involving the generalization of conditioned taste aversions

The present study employed a conditioned taste aversion generalizationparadigm to test the hypothesis that maltose produces tastesensations in the rat which are qualitatively distinguishablefrom sucrose. Since stimulus generalization can occur in boththe quality and intensity domains, an intrachemical (acrossconcentration) generalization gradient was established to aidin the interpretation of the interchemical (across molecules)generalization gradient. Moreover, since the commonly used intaketest is vulnerable to nontaste post-ingestional influences,the present study measured immediate responses to 100 µlstimulus samples, thus increasing our confidence that the behaviorwas under orosensory control. In Experiment 1, naive water deprivedrats were trained in a specially designed gustometer to maintaindrinking-spout contact for intermittent water reinforcement.Following this, rats in the experimental group were given threeexposures to 0.1 M sucrose on separate days, with the firsttwo exposures immediately preceding an injection of LiCl. Acontrol group was treated identically but received distilledwater instead of sucrose. Rats were then tested in the gustometerfor their avoidance of three equimolar concentrations of sucroseand maltose. Rats received ten trials of each stimulus quasi-randomlypresented in two sessions. Results indicated that all sucroseconcentrations were avoided (in experimental group only), butonly the 0.3 M concentration of maltose was avoided. The lowestsucrose concentration was significantly less avoided than thehigher concentrations. Intensity generalization gradients aresuch that intensities weaker than the conditioned stimulus (CS)produce just as much or less of a conditioned response (CR)and intensities stronger than the CS produce just as much ora greater CR than that elicited by the CS itself. Therefore,based on the results of Experimental, it was predicted thatif 0.1 M maltose served as the CS, the order of avoidance shouldbe: 0.3 M sucrose 0.1 M sucrose 0.03 M sucrose 0.3 M maltose 0.1 M maltose 0.03 M maltose, if it were true that maltoseand sucrose produce identical sensations that differ only inintensity. Experiment 2 explicitly tested this prediction usingthe same procedure as Experiment 1 except that 0.1 M maltoseserved as the CS. The observed order of avoidance was 0.3 Mmaltose > 0.1 M maltose > 0.03 M maltose = 0.3 M sucrose= 0.1 M sucrose = 0.03 M sucrose. In both experiments the intrachemicalgeneralization gradient broadened and the interchemical generalizationgradient steepened upon retesting. In conclusion, qualitativedifferences between maltose and sucrose explain the outcomesof these experiments better than differences in the relativeintensity of these sugars at isomolar concentrations.     

On the relationship between sweet taste and seasonal body weight changes in a primate (Microcebus murinus)

The relationship between obesity and taste, especially sweettaste, has been and is of interest. From this point of viewof a small primate, the lesser mouse lemur (Microcebus murinus),is of particular interest. It goes through a yearly cycle ofphysiological changes, one of which is an extreme variationin body weight of up to 100%. This occurs concomitantly withsignificant changes of the animal's liking for sucrose; measuredby two-bottle preference tests, the threshold for sucrose changesfrom 28–45 in lean to 77–105 mM in obese animals.It is possible that a change in peripheral taste sensitivitymight be the cause for these preference changes. To test thispossibility we studied the ability of M.murinus to taste sucrosewith electrophysiological and conditioned taste aversion techniques.The electrophysiological recordings were obtained from the chordatympani proper nerve in two heavy and three lean animals. Wedid not record any difference between the two groups in theirneural response to a series of sucrose concentrations. Conditionedtaste aversion experiments with 200 mM sucrose as conditioningstimulus and 50 and 200 mM sucrose as test stimuli gave similarresults. No difference was found between three heavy and fourlean animals; both groups rejected the sucrose concentrations.The results support the notion that the seasonal variationsin preference threshold to sucrose were unrelated to the abilityof M.murinus to taste sucrose.     

Tamoxifen produces conditioned taste avoidance in male rats: An analysis of microstructural licking patterns and taste reactivity

Estrogen receptor activation has been shown to reduce body weight and produce a conditioned reduction in food intake in male rats that is putatively mediated by estradiol's suggested aversive effects. Evidence has shown that the selective estrogen receptor modulator tamoxifen used in the prevention and treatment of breast cancer may also produce changes in food intake and body weight, which are known to impact cancer development and survival. The purpose of the present study was to examine whether tamoxifen produces a conditioned reduction in intake similar to estradiol by producing a conditioned aversion. A one bottle lickometer test was used to examine conditioned changes in sucrose drinking, while the taste reactivity test was used to measure rejection reactions, which serve to index aversion in rats. A backward conditioning procedure that consisted of 3 conditioning days and one vehicle test day was used to examine conditioned changes in 0.3 M sucrose intake and taste reactivity. Our results show that tamoxifen produced a conditioned reduction in sucrose drinking in a one bottle fluid intake test that was similar to the effects produced by estradiol (positive control); however, no active rejection reactions were produced by either tamoxifen (1 and 10 mg/kg) or estradiol. The present results suggest that tamoxifen, at the doses used in the present study, acts as an estrogen receptor agonist to regulate food intake and that the conditioned reduction in intake produced by tamoxifen and estradiol reflects conditioned taste avoidance rather than conditioned taste aversion.     

Extra-lingual chemoreceptors in the chicken (Gallus domesticus)

The oral aversion behaviour of the chicken (head shaking, beakwiping and tongue/beak movements) was measured following oralstimulation with 0.1 M quinine hydrochloride, 40% sucrose, 3M sodium chloride, 5 M acetic acid and pure methyl anthranilate.Section of the lingual and laryngolingual nerves did not affectthe oral aversion behaviour and therefore demonstrates the presenceof functional extra-lingual chemoreceptors. The results arediscussed in relation to previous anatomical findings.     

On the sense of taste in two Malagasy Primates (Microcebus murinus and Eulemur mongoz)

The relationship between phylogeny and taste is of growing interest.In this study we present recordings from the chorda tympaniproper (CT) nerve of two lemuriforme primates, the lesser mouselemur (Microcebus murinus) and the mongoose lemur (Eulemur mongoz),to an array of taste stimuli which included the sweeteners acesulfame-K,alitame, aspartame, D-glucose, dulcin, monellin, neohesperidindihydrochalcone (NHDHC), saccharin, sodium superaspartame, stevioside,sucralose (TGS), sucrose, suosan, thaumatin and xylitol, aswell as the non–sweet stimuli NaC1, citric acid, tanninand quinine hydrochloride. In M.murinus the effects of the tastemodifiers gymnemic acid and miraculin on the CT response wererecorded. Conditioned taste aversion (CTA) experiments in M.murinusand two-bottle preference (TBP) tests in E.mongoz were alsoconducted. We found that all of the above tastants except thaumatinelicited a CT response in both species. The CTA technique showedthat M.murinus generalized from sucrose to monellin but notto thaumatin. The intake of aspartame, ranging in concentrationfrom 0.1 to 30 mM was measured in E.mongoz with TBP tests. Atno concentration did we see a preference, but there was a significantrejection of 10 and 30 mM aspartame (P     

Different responsiveness of the chorda tympani and glossopharyngeal nerves to L-lysine in mice

Differential taste responsiveness and functional role of thetwo taste nerves, the chorda tympani (CT) and die glossopharyngeal(GL), were studied in mice by examining neural and behavioralresponses to an essential amino acid, L-lysine (Lys). Relativeresponses to Lys were larger in the GL than in the CT nerve.The neural threshold for the Lys response was about 2.5 logunits lower in the GL (about 1.0 µM) than in the CT nerve(about 300 µM). An analysis of concentration-responserelationships suggests a possibility that there are two differentreceptors (high and low affinity types) for Lys showing differentdissociation constants. The posterior tongue region possessesboth types, while the anterior region possesses only the lowaffinity type. Behavioral aversion threshold for Lys in intact mice, measuredby use of a single bottle test, was about 1.0 µM. Thisthreshold was the same as its neural threshold in the GL nerve.Animals whose bilateral GL nerves were sectioned showed a higheraversion threshold (about 300 µM) which was the same asthe neural threshold in the CT nerve. An aversion conditionedto Lys significantly generalized to L-arginine in the intactand CT-denervated mice, and L-arginine and L-histidine in theGL-denervated mice, but the generalization pattern across varioustaste stimuli including the four basic taste stimuli (NaCl,HCl, quinine HCl and sucrose) did not prominently differ amongthe intact, the GL-denervated and CT-denervated mice. These results suggest that taste sensitivity to Lys is higherin the GL than in the CT nerve, but taste quality informationfor Lys conveyed by two taste nerves is not largely different.     

Topographic distribution of taste responsiveness in the hamster medulla

The purpose of the present investigation was to map the multiunitresponsiveness of the gustatory portion of the nucleus of thesolitary tract (NTS) in the hamster, elicited by chemical stimulationof oral taste receptors. Neural responsiveness to four stimuli(0.1 M sucrose, 0.03 M NaCl, 0.003 M HCl, 0.001 M QHCl) deliveredto either the anterior tongue or other parts of the oral cavitywas examined at 37 NTS recording sites. Gustatory responseswere shown-to depend collectively upon the stimulus, the receptivearea being stimulated, and the location of the recording sitewithin the NTS. By comparing the proportional magnitudes ofintegrated responses across recording sites, unique topographicpatterns of responsiveness were demonstrated for sucrose, NaCIand QHCl. Responses to HCl and NaCl generated similar patterns.Further, the response patterns for each stimulus differed followingstimulation of the anterior tongue or posterior oral cavity.Spatial differences in NTS responsiveness arise as a resultof differences in peripheral gustatory nerve sensitivities andprovide a possible substrate for the coding of taste quality.     

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

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