Effects of chorda tympani nerve anesthesia on taste responses in the NST

Human clinical and psychophysical observations suggest that the taste system is able to compensate for losses in peripheral nerve input, since patients do not commonly report decrements in whole mouth taste following chorda tympani nerve damage or anesthesia. Indeed, neurophysiological data from the rat nucleus of the solitary tract (NST) suggests that a release of inhibition (disinhibition) may occur centrally following chorda tympani nerve anesthesia. Our purpose was to study this possibility further. We recorded from 59 multi- and single- unit taste-responsive sites in the rat NST before, during and after recovery from chorda tympani nerve anesthesia. During anesthesia, average anterior tongue responses were eliminated but no compensatory increases in palatal or posterior tongue responses were observed. However, six individual sites displayed increased taste responsiveness during anesthesia. The average increase was 32.9%. Therefore, disinhibition of taste responses was observed, but infrequently and to a small degree in the NST At a subset of sites, chorda tympani-mediated responses decreased while greater superficial petrosal-mediated responses remained the same during anesthesia. Since this effect was accompanied by a decrease in spontaneous activity, we propose that taste compensation may result in part by a change in signal-to-noise ratio at a subset of sites.      

Selective procaine inhibition of rat chorda tympani responses to electric taste stimulation

1. The lingual treatment of 1% procaine for 10 min selectively suppressed responses of the rat chorda tympani nerve to anodal current applied to the tongue with NaCl in the bathing medium to about 50% of control but the drug produced no significant suppression in responses to chemical taste stimuli. 2. The magnitude of suppression of response to anodal current varied with concentration of procaine and kind of bathing medium for the current stimulation (larger in the order of NaCl greater than KCl greater than CaCl2 greater than HCl). 3. Such ion specificity in procaine suppression suggests that responses of the chorda tympani nerve to anodal current are provoked through the taste cell (not direct action on the taste nerve), and that the receptor mechanisms for anodal current are at least partly different from that for chemical taste stimuli.     

Taste responses of human chorda tympani nerve

Records from humans of summated action potential dischargesof the chorda tympani nerve were examined. The magnitudes ofneural and psychophysical responses were well related only whenthe comparison was made within a given taste quality. The responseto a mixture of 0 02 M citric acid and 0.5 M sucrose was lessthan the sum of the separate responses to the mixture components.Citric acid failed to cross-adapt the response to sucrose, implyingthe receptor sites for sucrose are independent of citric acid.The human chorda tympani nerve shows vigorous responses to mechanicalstimulation and cooling of the tongue that are maintained aftertreatment of the tongue with a water extract of the herb Gymnemasylvestre. Gymnema extract selectively suppressed the responseto all sweeteners tested (sucrose, fructose, saccharin and cyclamate)and also suppressed by – 50% the water-after-citric-acidresponse which has a predominantly sweet taste. Gymnema suppressedby 0 – 10% the water-rinse response following NaCl. fructoseand sucrose that have a predominantly bitter-sour taste. Water-rinseresponses were present even when mechanical and thermal stimulationof the tongue were minimized. The human chorda tympani nerveappears to have positive water-rinse taste responses. Theseare solute-specific off-responses that are probably mediatedby receptor sites independent of those responsible for the on-responseto the given solute.     

Neuropeptides modulate rat chorda tympani responses

The neuropeptide leptin has been shown to selectively modulate rat chorda tympani (CT) responses to sweet tastants. To explore whether other neuropeptides can modulate such responses, rat whole nerve CT responses to NaCl, HCl, quinine HCl, and sucrose were measured while administering cholecystokinin-8 (CCK-8), substance P(4-11) (SP(4-11)), or calcitonin gene-related peptide (CGRP). To avoid possible confounding effects on CT responses that take long times to develop, such as those that arise from intraperitoneal injections, we investigated the effects of the above peptides injected into the ipsilateral lingual artery (LA) on CT nerve responses during the initial seconds after a tastant was placed on the tongue. We found that CT responses to NaCl and HCl were increased by CCK-8 and decreased by CGRP. SP(4-11) had no noticeable effect. Peptide-induced CT responses to quinine HCl or sucrose were too small to accurately detect. These data suggest that at short latencies, after local infusion via the LA, neuropeptides can alter CT responses in a peptide-specific manner.     

Developmental changes in sugar responses of the chorda tympani nerve in preweanling rats

To clarify developmental changes in the gustatory system of the rat, integrated taste responses from the chorda tympani (CT) nerve were recorded and analyzed at different postnatal ages. The response magnitude was calculated relative to the response to the standard, 0.1 M NH4Cl. Even at 1 week of age, the CT responded well to all tested 0.1 M chloride salts (NH4Cl, NaCl, LiCl, KCl, RbCl and CsCl). The responses to 0.1 M NaCl and LiCl increased with increasing age of the rat while response magnitudes to KCl, RbCl and CsCl did not change up to 8 weeks. At 1 week, the integrated response pattern was quite similar to that in adult rats for NaCl, HCl and quinine hydrochloride (QHCl). The concentration-response functions for NaCl, HCl, QHCl and sucrose at 2 weeks were essentially the same as those at 8 weeks. These results suggest that taste buds in the 2-week-old rat are functionally mature for the detection of the four basic taste stimuli. The relative magnitude of the responses to the various sugars was smaller at 1 week compared to the adult rat and reached a maximum at weeks 3-4, then decreased gradually with age. Among the six sugars, sucrose was the most effective followed by lactose. From weeks 1-4, the magnitude of the integrated taste response to fructose was smaller than that to lactose except at 3 weeks of age. Maltose, galactose and glucose were less potent stimuli than the other sugars tested. The response magnitude to lactose at 4 weeks had decreased compared to that for the other sugars. Taste responses to the sugars in preweanling and adult rats were not cross-adapted by the individual sugars. These results suggest that after 1 week of age during postnatal development in the rat, taste information from the CT rapidly increases in its importance for feeding behavior.     

Anion modulation of taste responses in sodium-sensitive neurons of the hamster chorda tympani nerve

Beidler's work in the 1950s showed that anions can strongly influence gustatory responses to sodium salts. We have demonstrated "anion inhibition" in the hamster by showing that the chorda tympani nerve responds more strongly to NaCl than to Na acetate over a wide range of concentrations. Iontophoretic presentation of Cl- and acetate to the anterior tongue elicited no response in the chorda tympani, suggesting that these anions are not directly stimulatory. Drugs (0.01, 1.0, and 100 microM anthracene-9-carboxylate, diphenylamine-2-carboxylate, 4- acetamido-4'-isothiocyanatostilbene-2,2'-disulfonate, and furosemide) that interfere with movements of Cl- across epithelial cells were ineffective in altering chorda tympani responses to 0.03 M of either NaCl or Na acetate. Anion inhibition related to movements of anions across epithelial membranes therefore seems unlikely. The chorda tympani contains a population of nerve fibers highly selective for Na+ (N fibers) and another population sensitive to Na+ as well as other salts and acids (H fibers). We found that N fibers respond similarly to NaCl and Na acetate, with spiking activity increasing with increasing stimulus concentration (0.01-1.0 M). H fibers, however, respond more strongly to NaCl than to Na acetate. Furthermore, H fibers increase spiking with increases in NaCl concentration, but generally decrease their responses to increasing concentrations of Na acetate. It appears that anion inhibition applies to taste cells innervated by H fibers but not by N fibers. Taste cells innervated by N fibers use an apical Na+ channel, whereas those innervated by H fibers may use a paracellularly mediated, basolateral site of excitation.     

The organization of taste sensibilities in hamster chorda tympani nerve fibers

Electrophysiological measurements of nerve impulse frequencies were used to explore the organization of taste sensibilities in single fibers of the hamster chorda tympani nerve. Moderately intense taste solutions that are either very similar or easily discriminated were applied to the anterior lingual surface. 40 response profiles or 13 stimulus activation patterns were considered variables and examined with multivariate statistical techniques. Three kinds of response profiles were seen in fibers that varied in their overall sensitivity to taste solutions. One profile (S) showed selectivity for sweeteners, a second (N) showed selectivity for sodium salts, and a third (H) showed sensitivity to salts, acids, and other compounds. Hierarchical cluster analysis indicated that profiles fell into discrete classes. Responses to many pairs of effective stimuli were covariant across profiles within a class, but some acidic stimuli had more idiosyncratic effects. Factor analysis of profiles identified two common factors, accounting for 77% of the variance. A unipolar factor was identified with the N profile, and a bipolar factor was identified with the S profile and its opposite, the H profile. Three stimulus activation patterns were elicited by taste solutions that varied in intensity of effect. Hierarchical cluster analysis indicated that the patterns fell into discrete classes. Factor analysis of patterns identified three common unipolar factors accounting for 82% of the variance. Eight stimuli (MgSO4, NH4Cl, KCl, citric acid, acetic acid, urea, quinine HCl, HCl) selectively activated fibers with H profiles, three stimuli (fructose, Na saccharin, sucrose) selectively activated fibers with S profiles, and two stimuli (NaNO3, NaCl) activated fibers with N profiles more strongly than fibers with H profiles. Stimuli that evoke different patterns taste distinct to hamsters. Stimuli that evoke the same pattern taste more similar. It was concluded that the hundreds of peripheral taste neurons that innervate the anterior tongue play one of three functional roles, providing information about one of three features that are shared by different chemical solutions.     

Effect of chorda tympani nerve transection on salt taste perception in mice

Effects of gustatory nerve transection on salt taste have been studied extensively in rats and hamsters but have not been well explored in the mouse. We examined the effects of chorda tympani (CT) nerve transection on NaCl taste preferences and thresholds in outbred CD-1 mice using a high-throughput phenotyping method developed in our laboratory. To measure taste thresholds, mice were conditioned by oral self-administration of LiCl or NaCl and then presented with NaCl concentration series in 2-bottle preference tests. LiCl-conditioned and control NaCl-exposed mice were given bilateral transections of the CT nerve (LiCl-CTX, NaCl-CTX) or were left intact as controls (LiCl-CNT, NaCl-CNT). After recovery from surgery, mice received a concentration series of NaCl (0-300 mM) in 48-h 2-bottle tests. CT transection increased NaCl taste thresholds in LiCl-conditioned mice and eliminated avoidance of concentrated NaCl in control NaCl-exposed mice. This demonstrates that in mice, the CT nerve is important for detection and recognition of NaCl taste and is necessary for the normal avoidance of high concentrations of NaCl. The results of this experiment also show that the method of high-throughput phenotyping of salt taste thresholds is suitable for detecting changes in the taste periphery in mouse genetic studies.     

Responses of the rat chorda tympani nerve to glutamate-sucrose mixtures

Monosodium glutamate (MSG) has a multifaceted, unusual taste to humans. Rats and other rodents also detect a complex taste to MSG. Responses of the chorda tympani nerve (CT) to glutamate applied to the front of the tongue were recorded in 13 anesthetized rats. Whole-nerve responses to 30 mM, 100 mM and 300 mM MSG mixed with 300 mM sucrose were recorded before and after adding 30 micro M amiloride to the rinse and stimulus solutions. Responses of CT single fibers were also recorded. Predictions from models of whole-nerve responses to binary mixtures were compared to the observed data. Results indicated that MSG-elicited CT responses have multiple sources, even in an amiloride-inhibited environment in rats. Those sources include responses of sucrose-sensitive CT neural units, which may provide the substrate for a sucrose-glutamate perceptual similarity, and responses of sucrose-insensitive CT neural units, which may respond synergistically to MSG-sucrose mixtures.     

The K+-H+ exchanger, nigericin, modulates taste cell pH and chorda tympani taste nerve responses to acidic stimuli

The relationship between acidic pH, taste cell pH(i), and chorda tympani (CT) nerve responses was investigated before and after incorporating the K(+)-H(+) exchanger, nigericin, in the apical membrane of taste cells. CT responses were recorded in anesthetized rats in vivo, and changes in pH(i) were monitored in polarized fungiform taste cells in vitro. Under control conditions, stimulating the tongue with 0.15 M potassium phosphate (KP) or 0.15 M sodium phosphate (NaP) buffers of pHs between 8.0 and 4.6, KP or NaP buffers did not elicit a CT response. Post-nigericin (500 × 10(-6) M), KP buffers, but not NaP buffers, induced CT responses at pHs ≤ 6.6. The effect of nigericin was reversed by the topical lingual application of carbonyl cyanide 3-chloro-phenylhydrazone, a protonophore. Post-nigericin (150 × 10(-6) M), KP buffers induced a greater decrease in taste cell pH(i) relative to NaP buffers and to NaP and KP buffers under control conditions. A decrease in pH(i) to about 6.9 induced by KP buffers was sufficient to elicit a CT response. The results suggest that facilitating apical H(+) entry via nigericin decreases taste cell pH(i) and demonstrates directly a strong correlation between pH(i) and the magnitude of the CT response.     

Sugar best single chorda tympani nerve fiber responses to various sugar stimuli in rat and hamster

1. Sugar best single chorda tympani nerve fiber of rat and hamster were tested with six sugars. 2. Fibers were selected for this experiment, only if they responded to 1.0 M sucrose or 1.0 M maltose and they responded poorly to 0.1 M NaCl. 3. In rat, some single fibers gave larger responses to maltose than to sucrose, while in hamster nearly all nerve fibers responded best to sucrose. 4. The order of effectiveness of sugars was maltose greater than fructose greater than or equal to lactose greater than sucrose greater than glucose greater than galactose in rat and sucrose greater than fructose greater than or equal to glucose greater than or equal to galactose greater than maltose greater than lactose in hamster.     

Acid-induced responses in hamster chorda tympani and intracellular pH tracking by taste receptor cells

HCl- and NaCl-induced hamster chorda tympani nerve responseswere recorded during voltage clamp of the lingual receptive field. Voltage perturbations did not influence responses to HCl. In contrast, responses to NaCl were decreased by submucosal-positive and increased by submucosal-negative voltage clamp. Responses to HCl were insensitive to the Na⁺ channel blockers,amiloride and benzamil, and to methylisobutylamiloride (MIA), anNa⁺/H⁺exchange blocker. Responses to NaCl were unaffected by MIA but weresuppressed by benzamil. Microfluorometric and imaging techniques wereused to monitor the relationship between external pH(pH_o) and the intracellular pH(pH_i) of fungiform papilla tastereceptor cells (TRCs) following2',7'-bis(2-carboxyethyl)-5(6)-carboxyfluorescein loading.TRC pH_i responded rapidly andmonotonically to changes in pH_o.This response was unaffected byNa⁺ removal or the presence ofamiloride, benzamil, or MIA. The neural records and the data fromisolated TRCs suggest that the principal transduction pathway for acidtaste in hamster is similar to that in rat. This may involve themonitoring of changes in TRC pH_i mediated through amiloride-insensitiveH⁺ transport across TRC membranes.This is an example of cell monitoring of environmental pH through pHtracking, i.e., a linear change inpH_i in response to a change inpH_o, as has been proposed for carotid bodies. In taste, the H⁺transport sites may be concentrated on the basolateral membranes ofTRCs and, therefore, are responsive to an attenuatedH⁺ concentration from diffusion ofacids across the tight junctions.

     

Modulation of rat chorda tympani NaCl responses and intracellular Na+ activity in polarized taste receptor cells by pH

Mixture interactions between sour and salt taste modalities were investigated in rats by direct measurement of intracellular pH (pH(i)) and Na(+) activity ([Na(+)](i)) in polarized fungiform taste receptor cells (TRCs) and by chorda tympani (CT) nerve recordings. Stimulating the lingual surface with NaCl solutions adjusted to pHs ranging between 2.0 and 10.3 increased the magnitude of NaCl CT responses linearly with increasing external pH (pH(o)). At pH 7.0, the epithelial sodium channel (ENaC) blocker, benzamil, decreased NaCl CT responses and inhibited further changes in CT responses induced by varying pH(o) to 2.0 or 10.3. At constant pH(o), buffering NaCl solutions with potassium acetate/acetic acid (KA/AA) or HCO(3)(-)/CO(2) inhibited NaCl CT responses relative to CT responses obtained with NaCl solutions buffered with HEPES. The carbonic anhydrase blockers, MK-507 and MK-417, attenuated the inhibition of NaCl CT responses in HCO(3)(-)/CO(2) buffer, suggesting a regulatory role for pH(i). In polarized TRCs step changes in apical pH(o) from 10.3 to 2.0 induced a linear decrease in pH(i) that remained within the physiological range (slope = 0.035; r(2) = 0.98). At constant pH(o), perfusing the apical membrane with Ringer's solutions buffered with KA/AA or HCO(3)(-)/CO(2) decreased resting TRC pH(i), and MK-507 or MK-417 attenuated the decrease in pH(i) in TRCs perfused with HCO(3)(-)/CO(2) buffer. In parallel experiments, TRC [Na(+)](i) decreased with (a) a decrease in apical pH, (b) exposing the apical membrane to amiloride or benzamil, (c) removal of apical Na(+), and (d) acid loading the cells with NH(4)Cl or sodium acetate at constant pH(o). Diethylpyrocarbonate and Zn(2+), modification reagents for histidine residues in proteins, attenuated the CO(2)-induced inhibition of NaCl CT responses and the pH(i)-induced inhibition of apical Na(+) influx in TRCs. We conclude that TRC pH(i) regulates Na(+)-influx through amiloride-sensitive apical ENaCs and hence modulates NaCl CT responses in acid/salt mixtures.     

咸味觉厌恶模型大鼠鼓索神经对味觉刺激的电生理反应特性

目的:探索大鼠咸味觉厌恶建立后外周鼓索神经(CT)对咸味觉及其他味觉刺激的电生理反应特性的改变。方法:将14只SD成年雄性大鼠分为咸味觉厌恶模型组(CTA)和对照组(n=7/group)。实验第1日给予大鼠30min的0.1mol/LNaCl饮食,随后CTA组和对照组大鼠分别腹腔注射2ml0.15mol/LLiCl和同等量生理盐水。在第2、3和4日,测量两组大鼠每天30min内对NaCl和蒸馏水饮用量。于第4日行为学测试后,分别记录CTA组大鼠和对照组大鼠CT对口内给予系列浓度NaCl溶液、0.3mol/LNaCl与0.1mmol/L阿米洛利(一种舌上皮钠通道阻断剂)混合液和其他四种基本味觉刺激溶液的电生理反应。结果:与对照组相比,CTA组大鼠CT对系列浓度NaCl和其他4种基本味觉刺激的电生理反应特性没有发生明显变化(P>0.05);舌上皮钠通道阻断剂阿米洛利强烈抑制CTA大鼠对NaCl的反应(P<0.01)。结论:条件性咸味觉厌恶模型大鼠CT对各种味觉刺激的电生理反应特性没有发生明显改变。     

Inhibitory effect of the extract from Zizyphus jujuba leaves on sweet taste responses of the chorda tympani in the rat and hamster

1. One of the fractions obtained from the extract of Zizyphus jujuba leaves suppressed the response of the chorda tympani to sucrose, both in the rat and hamster. 2. In the rat and man, suppressive effect was found to be significant in responses to various sugars and artificial sweeteners but not in some sweet amino acids.     

Sensitivities of single nerve fibers in the hamster chorda tympani to mixtures of taste stimuli

Responses of three groups of neural fibers from the chorda tympani of the hamster to binary mixtures of taste stimuli applied to the tongue were analyzed. The groups displayed different sensitivities to six chemicals at concentrations that had approximately equal effects on the whole nerve. Sucrose-best fibers responded strongly only to sucrose and D-phenylalanine. NaCl-best and HCl-best fibers, responded to four electrolytes: equally to CaCl2 and nearly equally to HCl, but the former responded more to NaCl, and the latter responded more to NH4Cl. The groups of fibers dealt differently with binary mixtures. Sucrose- best fibers responded to a mixture of sucrose and D-phenylalanine as if one of the chemicals had been appropriately increased in concentration, but they responded to a mixture of either one and an electrolyte as if the concentration of sucrose or D-phenylalanine had been reduced. NaCl- best fibers responded to a mixture as if it were a "mixture" of two appropriate concentrations of one chemical, or somewhat less. But, responses of HCl-best fibers to mixtures were greater than that, approaching a sum of responses to components. These results explain effects on the whole nerve, suggest that the sensitivity of a mammalian taste receptor to one chemical can be affected by a second, which may or may not be a stimulus for that receptor, and suggest that some effects of taste mixtures in humans may be the result of peripheral processes.     

Expression of amiloride-sensitive epithelial sodium channels in mouse taste cells after chorda tympani nerve crush

Our previous electrophysiological study demonstrated that amiloride-sensitive (AS) and -insensitive (AI) components of NaCl responses recovered differentially after the mouse chorda tympani (CT) was crushed. AI responses reappeared earlier (at 3 weeks after the nerve crush) than did AS ones (at 4 weeks). This and other results suggested that two salt-responsive systems were differentially and independently reformed after nerve crush. To investigate the molecular mechanisms of formation of the salt responsive systems, we examined expression patterns of three subunits (alpha, beta and gamma) of the amiloride-sensitive epithelial Na(+) channel (ENaC) in mouse taste cells after CT nerve crush by using in situ hybridization (ISH) analysis. The results showed that all three ENaC subunits, as well as alpha-gustducin, a marker of differentiated taste cells, were expressed in a subset of taste bud cells from an early stage (1-2 weeks) after nerve crush, although these taste buds were smaller and fewer in number than for control mice. At 3 weeks, the mean number of each ENaC subunit and alpha-gustducin mRNA-positive cells per taste bud reached the control level. Also, the size of taste buds became similar to those of the control mice at this time. Our previous electrophysiological study demonstrated that at 2 weeks no significant response of the nerve to chemical stimuli was observed. Thus ENaC subunits appear to be expressed prior to the reappearance of AI and AS neural responses after CT nerve crush. These results support the view that differentiation of taste cells into AS or AI cells is initiated prior to synapse formation.     

Changes in number and morphology of fungiform taste buds in rat after transection of the chorda tympani or chordalingual nerve

In normal rats there is one taste bud on the apical surfaceof each fungiform papilla. These taste buds are innervated bythe chorda tympani proper nerve (CT). According to general consensus,after cutting the nerve the taste buds should disappear. Inthis study, performed on 24 rats divided in six groups, theCT nerve on the left side (singly denervated) and the combinedchorda-lingual (CT-L) nerve on the other side (doubly denervatedwere permanently interrupted. The animals were sacrificed after5, 10, 20, 35,60 and 100 days and their tongues serially sectionedfor light microscope examiation. Some papillae were examinedunder an electron microscope. The papillae were categorizedinto three groups: papillae with a normal looking taste bud,with an abnormal looking taste bud and without a taste bud.The results showed a substantial number of papillae with a normallooking taste bud present at all time intervals in all animals.More specifically, on the singly denervated side the proportionof normal looking taste buds stayed below 10% until day 60,when it increased to 15% and to 23% on day 100. The proportionof abnormal looking taste buds decreased from above 92% by day5 to 49% on day 100. The percentage of fungiform papillae withoutsigns of a taste bud was relatively low on the singly denervatedside at times (1, 5, 16, 29, 34 and 28%). On the doubly denervatedside fewer than than 4% normal looking taste buds were foundthroughout the time period. The proportion of abnormal lookingtaste buds decreased from {small tilde} 96% by day 5 to 35%on day 100. A significantly higher proportion of papillae withno taste bud was observed on this side from day 10 onwards.(1, 29, 32, 52, 60 and 63%). The reasons for the differencein tast bud number between the doubly and singly denervatedsides are unknown, but it is possible that collaterals fromother (non-gustatory) nerves have an ability, although limited,to induce and maintain fungiform taste buds. In other words,the capacity to induce taste bud formation is not limited exclusivelyto gustatory nerves.     

Functional status of the regenerated chorda tympani nerve as assessed in a salt taste discrimination task

We tested whether the recovered ability of rats to discriminate NaCl from KCl after chorda tympani nerve transection (CTX) is causally linked to nerve regeneration or some other compensatory process. Rats were presurgically trained in an operant NaCl vs. KCl discrimination task. Rats with regenerated nerves, histologically confirmed by anterior tongue taste pore counts and tested 62 days after CTX (CTX-62R; n = 5), performed as well as those tested 62 days after sham surgery (Sham-62; n = 5), but both of these groups initially performed slightly worse than animals tested 7 days after sham surgery (Sham-7; n = 4). Performance of rats tested either 7 (CTX-7P; n = 5) or 62 (CTX-62P; n = 4) days after CTX in which nerve regeneration was prevented was severely disrupted. Adulteration of the stimuli with amiloride, an epithelial sodium channel blocker, impaired discrimination performance in a similar dose-dependent manner in the Sham-7 (n = 2), Sham-62 (n = 5), and CTX-62R (n = 5) groups, suggesting that the functional status of the amiloride-sensitive transduction pathway returns to normal in rats with regenerated chorda tympani nerves. Performance of CTX rats without regenerated nerves (CTX-7P, n = 2; CTX-62P, n = 4) was further degraded by amiloride treatment, suggesting that taste receptors innervated by other nerves are sensitive to amiloride. In conclusion, nerve regeneration is an essential component underlying full recovery of salt discrimination function after CTX.