Cortical cholinergic activity is related to the novelty of the stimulus

A number of studies have related cholinergic activity to the mediation of learning and memory. However, the acetylcholine (ACh) participation has been recently implicated in the early stages of memory formation but not during retrieval. The aim of the present study is to evaluate ACh release in the insular cortex (IC) during presentation of different taste stimuli and during their re-exposition by means of the free-moving microdialysis technique. We evaluated the changes in ACh release when a novel taste, saccharin or quinine was presented to the rat and after several presentations of saccharin. Unilateral microdialysis was performed in the IC 1 h before and 1 h after the presentation of: (1) a familiar stimulus (water), (2) a novel taste (quinine), (3) another novel taste (saccharin), (4) a second presentation, (5) a third presentation, and (6) a fourth presentation of saccharin. The volume consumed by the animals was registered as a behavioral parameter. The ACh levels from the microdialysis fractions were analyzed by an HPLC-ED system. Biochemical results showed a significant increment in the cortical ACh release induced by a novel stimulus compared with the release observed during the presentation of a familiar stimulus. The ACh release observed after several presentations of the stimuli decreased to the same levels as those produced by the familiar taste, indicating an inverse relationship between familiarity and cortical ACh release. These results suggest that the cholinergic system plays an important role in the identification and characterization of different kinds of stimuli.     

Neurons in the posterior insular cortex are responsive to gustatory stimulation of the pharyngolarynx, baroreceptor and chemoreceptor stimulation, and tail pinch in rats

Extracellular unit responses to gustatory stimulation of the pharyngolaryngeal region, baroreceptor and chemoreceptor stimulation, and tail pinch were recorded from the insular cortex of anesthetized and paralyzed rats. Of the 32 neurons identified, 28 responded to at least one of the nine stimuli used in the present study. Of the 32 neurons, 11 showed an excitatory response to tail pinch, 13 showed an inhibitory response, and the remaining eight had no response. Of the 32 neurons, eight responded to baroreceptor stimulation by an intravenous (i.v.) injection of methoxamine hydrochloride (Mex), four were excitatory and four were inhibitory. Thirteen neurons were excited and six neurons were inhibited by an arterial chemoreceptor stimulation by an i.v. injection of sodium cyanide (NaCN). Twenty-two neurons were responsive to at least one of the gustatory stimuli (deionized water, 1.0 M NaCl, 30 mM HCl, 30 mM quinine HCl, and 1.0 M sucrose); five to 11 excitatory neurons and three to seven inhibitory neurons for each stimulus. A large number of the neurons (25/32) received converging inputs from more than one stimulus among the nine stimuli used in the present study. Most neurons (23/32) received converging inputs from different modalities (gustatory, visceral, and tail pinch). The neurons responded were located in the insular cortex between 2.0 mm anterior and 0.2 mm posterior to the anterior edge of the joining of the anterior commissure (AC); the mean location was 1.2 mm (n=28) anterior to the AC. This indicates that most of the neurons identified in the present study seem to be located in the region posterior to the taste area and anterior to the visceral area in the insular cortex. These results indicate that the insular cortex neurons distributing between the taste area and the visceral area receive convergent inputs from gustatory, baroreceptor, chemoreceptor, and nociceptive organs.     

The effect of taste stimuli on histamine release in the anterior hypothalamus of rats

We studied the effect of gustatory stimulation on hypothalamic histamine release. Administering a four-basic taste mixture significantly increased histamine release, but not in the chorda tympani-transected rats. 0.1 M NaCl significantly increased histamine release, whereas 0.5 M sucrose, 0.02 M quinine HCl and 0.01 M HCl had no effect. However, when the concentration of HCl was increased to 0.03 M, a significant increase in histamine release was seen. These results suggest that taste information via the chorda tympani activates the histaminergic system.     

GABA-mediated corticofugal inhibition of taste-responsive neurons in the nucleus of the solitary tract

The nucleus of the solitary tract (NST) receives descending connections from several forebrain targets of the gustatory system, including the insular cortex. Many taste-responsive cells in the NST are inhibited by gamma-aminobutyric acid (GABA). In the present study, we investigated the effects of cortical stimulation on the activity of gustatory neurons in the NST. Multibarrel glass micropipettes were used to record the activity of NST neurons extracellularly and to apply the GABA(A) antagonist bicuculline methiodide (BICM) into the vicinity of the cell. Taste stimuli were 0.032 M sucrose (S), 0.032 M NaCl (N), 0.00032 M citric acid (H), and 0.032 M quinine hydrochloride (Q), presented to the anterior tongue. Each of 50 NST cells was classified as S-, N-, H-, or Q-best on the basis of its response to chemical stimulation of the tongue. The ipsilateral insular cortex was stimulated both electrically (0.5 mA, 100 Hz, 0.2 ms) and chemically (10 mM DL-homocysteic acid, DLH), while the spontaneous activity of each NST cell was recorded. The baseline activity of 34% of the cells (n=17) was modulated by cortical stimulation: eight cells were inhibited and nine were excited. BICM microinjected into the NST blocked the cortical-induced inhibition but had no effect on the excitatory response. Although the excitatory effects were distributed across S-, N-, and H-best neurons, the inhibitory effects of cortical stimulation were significantly more common in N-best cells. These data suggest that corticofugal input to the NST may differentially inhibit gustatory afferent activity through GABAergic mechanisms.     

Hypothalamic histamine release by taste stimuli in freely moving rats: possible implication of palatability

Our previous study indicated that taste information via the chorda tympani (CT) activates the central histaminergic system in anesthetized rats. However, the physiological roles of taste-induced histamine release remain unknown, thus to further investigate the relationship between histamine release and gustatory information, in the present study we investigated the effect of taste stimuli infused intraorally on histamine release using in vivo microdialysis in free moving rats. Consistent with findings from our previous study, application of NaCl and HCl caused significant increases in histamine levels further supporting the suggestion that this phenomenon is attributed to the excitation of the CT. When rats were intraorally infused with quinine HCl (QHCl) solution, a significant increase in hypothalamic histamine release was observed. On the other hand, histamine release was decreased by sucrose and saccharin solutions. When rats were conditioned to acquire taste aversion to sucrose solution or saccharin solution, instead of the histamine decrease seen by the palatable solutions, the pattern of histamine release was similar to that seen by QHCl solution. From these observations, it is concluded that the histamine release by the infusion of these tastants may be explained by two mechanisms-by causing a transient increase after taste stimulation and by causing a decrease relative to the tastant's palatability.     

Comparison of c-fos-like immunoreactivity in the brainstem following intraoral and intragastric infusions of chemical solutions in rats

To examine whether the activation of brainstem neurons during ingestion is due to orosensory afferents or post-ingestive factors, neuronal activation in response to intraoral and intragastric infusions of taste stimuli was compared in the area postrema (AP), nucleus tractus solitarius (NTS) and parabrachial nucleus (PBN) by the c-fos immunohistochemical method. An aliquot (7.5 ml) of 0.5 M sucrose, 5 mM sodium saccharin, 1 mM quinine hydrochloride and distilled water was delivered into the oral cavity or the stomach in each rat, which had been deprived of water and food overnight. Water induced little c-Fos-like immunoreactivity (c-FLI), but both intraoral and intragastric infusions of sucrose, but not non-caloric saccharin, induced strong c-FLI in the AP, caudal NTS and the external lateral subnucleus of the rostral PBN, suggesting that these areas receive general visceral inputs. Other areas in the NTS and PBN may receive gustatory inputs since more dominant c-FLI was detected by intraoral rather than intragastric infusions of the stimuli. Functional segregation of neurons reflecting qualitative and hedonic aspects of sweeteners (sucrose and saccharin) and bitter-tasting substance (quinine) was suggested in the PBN, but less evident in the NTS. These results indicate that c-fos induction in brainstem neurons during ingestion reflects gustatory inputs and postingestional factors depending on the kind of food ingested.     

Responses of the hamster chorda tympani nerve to binary component taste stimuli: evidence for peripheral gustatory mixture interactions

Studies of taste mixtures suggest that stimuli which elicit different perceptual taste qualities physiologically interact in the gustatory system and thus. are not independently processed. The present study addressed the role of the peripheral gustatory system in these physiological interactions by measuring the effects of three heterogeneous taste mixtures on responses of the chorda tympani (CT) nerve in the hamster ( Mesocricetus auratus). Binary taste stimuli were presented to the anterior tongue and multi-fiber neural responses were recorded from the whole CT. Stimuli consisted of a concentration series of quinine. HCI (QHCI: 1–30 mM), sodium chloride (NaCl: 10–250 mM). sucrose (50–500 mM) and binary combinations of the three different chemicals. Each mixture produced a unique pattern of results on CT response magnitudes measured 10 s into the response. Sucrose responses were inhibited by quinine in QHCI-sucrose mixtures. Neural activity did not increase when quinine was added to 50–250 mM NaCl in QHCINaCl mixtures. However, the neural activity elicited by sucrose-NaCl mixtures was greater than the activity elicited by either component stimulus presented alone. The results demonstrate that gustatory mixture interactions are initiated at the level of the taste bud or peripheral nerve. Mechanisms for these interactions are unknown. The results are consistent with one component stimulus modifying the interaction of the other component stimulus with its respective transduction mechanism. Alternatively, peripheral inhibitory mechanisms may come into play when appetitive and aversive stimuli are simultaneously presented to the taste receptors.     

Forebrain contribution to the induction of a brainstem correlate of conditioned taste aversion: II. Insular (gustatory) cortex

The induction of c-Fos-like immunoreactivity (c-FLI) in the intermediate division of the nucleus of the solitary tract (iNTS) has been shown to be a cellular correlate of the behavioral expression of a conditioned taste aversion (CTA). To further define neuroanatomical structures and pathways that contribute to this cellular response and to CTA learning in general, electrolytic lesions of insular (gustatory) cortex (IC) were combined with immunostaining for c-FLI. Rats were given either unilateral or bilateral electrolytic lesions of insular cortex or `sham' operations. Following surgery, `paired' animals were given a single conditioning trial consisting of intraoral infusion of 5-ml 0.15% sodium–saccharin followed by injection with LiCl (0.15 M, 20 ml/kg, i.p.) while `unpaired' controls received a non-contingent saccharin–LiCl presentation. Rats with bilateral lesions showed no behavioral evidence of having acquired a CTA. Increases in c-FLI in iNTS were evident, but reduced, relative to `sham' animals. Rats with unilateral-lesions displayed a CTA by rejecting the saccharin, although increases in c-FLI on the side of the iNTS ipsilateral to the lesion were reduced relative to that seen in `sham' animals. A comparison of these results with those obtained after amygdala lesions supports the conclusion that amygdala and insular cortex are necessary, but not sufficient, for the behavioral expression of a CTA.     

The contribution of gustatory nerve input to oral motor behavior and intake-based preference. I. Effects of chorda tympani or glossopharyngeal nerve section in the rat

Two-bottle intake tests and taste reactivity (TR) tests were used to reveal whether changes in ingestive behavior would follow bilateral section of either the chorda tympani (CT) or the glossopharyngeal (GP) nerve. Rats received two-bottle intake tests to compare 24-h ingestion of water to that of NaCl, MgCl2, quinine, or sucrose. Prior to each long-term intake test, rats received a 1 min, 1 ml intraoral infusion of the same chemical stimulus. Ingestive and aversive oral motor responses elicited by these 1 ml infusions were videotaped and subsequently analyzed. GP-section did not alter quinine or sucrose preference; overall, preference of MgCl2 and NaCl was also similar to controls. In contrast, TR tests in GP-sectioned rats revealed that most quinine, MgCl2 and NaCl stimuli elicited significantly fewer aversive oral motor responses. In addition, the latency of aversive responses to these 3 chemical stimuli was increased for these rats. Intake-based preference tests failed to show any difference between rats with CT nerve section and controls. In TR tests, however, CT-sectioned rats displayed significantly fewer ingestive oral motor responses to NaCl, MgCl2, and quinine than controls. Neither sucrose intake nor sucrose-elicited TR were altered by CT or GP nerve section. This report confirms the failure of long-term intake tests to uncover behavioral deficits following the section of gustatory nerves. In contrast, the use of a different behavioral test makes clear for the first time that gustatory nerve section has dramatic consequences on ingestive behavior. The examination of taste elicited oral motor behaviors reveals a coherent and nerve specific pattern of neurological deficit following peripheral nerve section.     

Effects of insular cortex lesions on conditioned taste aversion and latent inhibition in the rat

The present study tested the hypothesis that lesions of the insular cortex of the rat retard the acquisition of conditioned taste aversions (CTAs) because of an impairment in the detection of the novelty of taste stimuli. Demonstrating the expected latent inhibition effect, nonlesioned control subjects acquired CTAs more rapidly when the conditioned stimulus (0.15% sodium saccharin) was novel rather than familiar (achieved by pre-exposure to the to-be-conditioned taste cue). However, rats with insular cortex lesions acquired taste aversions at the same slow rate regardless of whether the saccharin was novel or familiar. The pattern of behavioural deficits obtained cannot be interpreted as disruptions of taste detection or stimulus intensity, but is consistent with the view that insular cortex lesions disrupt taste neophobia, a dysfunction that consequently retards CTA acquisition because of a latent inhibition-like effect.     

Hunger Modulates the Responses to Gustatory Stimuli of Single Neurons in the Caudolateral Orbitofrontal Cortex of the Macaque Monkey

1. In order to determine whether the responsiveness of neurons in the caudolateral orbitofrontal cortex (a secondary cortical gustatory area) is influenced by hunger, the activity evoked by prototypical taste stimuli (glucose, NaCl, HCl, and quinine hydrochloride) and fruit juice was recorded in single neurons in this cortical area before, while, and after cynomolgous macaque monkeys were fed to satiety with glucose or fruit juice. 2. It was found that the responses of the neurons to the taste of the glucose decreased to zero while the monkey ate it to satiety during the course of which his behaviour turned from avid acceptance to active rejection. 3. This modulation of responsiveness of the gustatory responses of the neurons to satiety was not due to peripheral adaptation in the gustatory system or to altered efficacy of gustatory stimulation after satiety was reached, because modulation of neuronal responsiveness by satiety was not seen at earlier stages of the gustatory system, including the nucleus of the solitary tract, the frontal opercular taste cortex, and the insular taste cortex. 4. The decreases in the responsiveness of the neurons were relatively specific to the food with which the monkey had been fed to satiety. For example, in seven experiments in which the monkey was fed glucose solution, neuronal responsiveness decreased to the taste of the glucose but not to the taste of blackcurrant juice. Conversely, in two experiments in which the monkey was fed to satiety with fruit juice, the responses of the neurons decreased to fruit juice but not to glucose. 5. These and earlier findings lead to a proposed neurophysiological mechanism for sensory-specific satiety in which the information coded by single neurons in the gustatory system becomes more specific through the processing stages consisting of the nucleus of the solitary tract, the taste thalamus, and the frontal opercular and insular taste primary taste cortices, until neuronal responses become relatively specific for the food tasted in the caudolateral orbitofrontal cortex (secondary) taste area. Then sensory-specific satiety occurs because in this caudolateral orbitofrontal cortex taste area (but not earlier in the taste system) it is a property of the synapses that repeated stimulation results in a decreased neuronal response. 6. Evidence was obtained that gustatory processing involved in thirst also becomes interfaced to motivation in the caudolateral orbitofrontal cortex taste projection area, in that neuronal responses here to water were decreased to zero while water was drunk until satiety was produced.     

Opioid modulation of taste responses in the nucleus of the solitary tract

Gustatory processing within the medulla is modulated by a number of physiologic and experiential factors. Several neurotransmitters, including excitatory amino acids, GABA, and substance P, are involved in synaptic processing within the rostral portion of the nucleus of the solitary tract (NST). Endogenous opiates have been implicated in the regulation of feeding behavior and in taste palatability and gustatory responses in the parabrachial nuclei are reduced by systemic morphine. In the present experiments, extracellular recording of neuronal activity within the NST in response to taste input was combined with local microinjection of met-enkephalin (Met-ENK) and naltrexone (NLTX) to determine the effect of these agents on gustatory activity. The anterior tongue was stimulated with anodal current pulses to determine the time course of drug action (n=85 cells) and with prototypical taste stimuli (0.032 M sucrose, NaCl, and quinine hydrochloride, and 0.0032 M citric acid) to investigate the effects of these opioid compounds on taste-evoked responses (n=80 cells). Among these 165 taste-responsive neurons in the NST, the activity of 39 (23.6%) was suppressed by Met-ENK. These effects were dose-dependent and blockable by NLTX, which alone was without effect, suggesting that opiates do not maintain a tonic inhibitory influence. Immunohistochemical experiments demonstrated both micro - and delta-opioid receptors within the gustatory portion of the NST; previous studies had shown numerous fiber terminals containing Met-ENK. These data suggest that endogenous opiates play an inhibitory role in gustatory processing within the medulla.     

A search for the cortical gustatory area in the hamster

The gustatory area was searched in the cerebral cortex of the hamster by means of a combined approach using electrophysiological, behavioral, and histological experiments. The chorda tympani (CT), which innervates taste buds on the anterior part of the tongue, projected to a confined area anterior to the middle cerebral artery and just dorsal to the rhinal fissure. The trigeminal component of the lingual nerve (LN) area was located anterodorsal to the CT area, and the glossopharyngeal nerve (GN), which innervates taste buds on the posterior part of the tongue, was posterior to the CT area. The center of the CT and GN areas belonged to the dorsal part of the dysgranular insular cortex, and the LN area was within the primary somatosensory granular cortex. Bilateral symmetrical ablations of the CT and GN areas abolished the conditioned taste aversion (to sodium saccharin) that had been acquired before ablations, indicating a role of these areas in some cognitive processes of taste perception. Injections of horseradish peroxidase conjugated with wheat germ agglutinin (WGA-HRP) in the CT and GN areas, centered in the dysgranular insular cortex, revealed that this cortical region had major fiber connections with the contralateral homotypical cortical area, ipsilateral amygdala (central, lateral and basolateral nuclei), ipsilateral parvicellular part of the posteromedial ventral nucleus of the thalamus, bilateral parabrachial nucleus, contralateral nucleus of the solitary tract, raphe nuclei, and the locus ceruleus. Conversely, injections of WGA-HRP in these target areas showed anterograde and/or retrograde transport in the similar dysgranular insular cortex and additionally in the ventral part of the granular insular cortex. The present results suggest that the cortical gustatory area of the hamster is about 1.5 × 1.5 mm in size with the topographic organization between anterior and posterior parts of the tongue, and is located mainly in the dysgranular insular cortex around the middle cerebral artery.     

On a neural mechanism for cortical processing of taste quality in the rat

The responses of 31 chorda tympani fibers and 47 cortical neurons were recorded in response to 6 concentrations of NaCl, and single concentrations of sucrose, HCL, and quinine hydrochloride solutions applied to the anterior portion of the tongue in rats. The neural responses were analyzed in terms of the two hypotheses of quality coding: across-neuron response pattern and across-region response pattern notions. In a behavioral experiment, animals were given a conditioned taste aversion to one of 5 concentrations of NaCl solution by pairing it with a gastrointestinal illness caused by i.p. injection of 0.15 M LiCl. Behavioral taste profiles were constructed for each stimulus from the suppression of drinking, which indicates the extent of generalization of aversion to each of the 4 basic taste stimuli. Among the two neural analyses employed for the chorda tympani and cortical units, across-region correlation coefficients for cortical neurons that were derived from the across-region response pattern theory showed the highest correlation (r= 0.89) with the behavioral suppression rates. Across-neuron correlation coefficients in the chorda tympani fibers also showed a good correlation (r= 0.81) with the behavioral data. However, the taste profile for 1.0 M NaCl in chorda tympani fibers was very similar to that for the lower concentrations of NaCl, in spite of the suggestion from the behavioral experiment and the neural analyses of cortical responses that 1.0 M NaCl has HCl and quinine components besides the NaCl component. The present result confirmed the idea that processing of taste information in the cortex involves differences in both response magnitude across neurons and the spatial localization of those neurons.     

Cardiovascular responses to gustatory and mechanical stimulation of the nasopharynx in rats

The effects of natural (mechanical and gustatory) stimulation of the nasopharynx or electrical stimulation of the pharyngeal branch of the glossopharyngeal (PH-IXth) nerve on the changes in heart rate (HR) and arterial blood pressure (BP) were investigated in paralyzed and anesthetized rats. Afferent responses in the PH-IXth nerve were also investigated. Electrical stimulation of the PH-IXth nerve elicited a tachycardia and an increase in BP. Among the gustatory (1.0 M NaCl, 0.03 M HCl, 0.03 M QHCl, 1.0 M sucrose, H₂O, and 0.9% NaCl) and mechanical stimuli applied to the nasopharynx, 1.0 M sucrose and 0.9% NaCl were ineffective in changing HR and BP; the rest of the stimuli were strongly effective as was the case with electrical stimulation of the PH-IXth nerve. Responses were evoked in the PH-IXth nerve by nasopharyngeal stimulation with the stimuli which were effective in producing cardiovascular responses. On the other hand, 1.0 M sucrose and 0.9% NaCl, which were ineffective stimuli for cardiovascular responses, did not produce any response in the PH-IXth nerve. There was a high correlation between the magnitude of the responses in the PH-IXth nerve and those of the cardiovascular system. These results indicate that gustatory and mechanical information carried in the PH-IXth nerve innervating the nasopharynx plays an important role in cardiovascular regulation as well as the sense of taste.     

The effect of satiety on responses of gustatory neurons in the amygdala of alert cynomolgus macaques

An alert cynomolgus macaque was fed a sweet solution to satiety as the activity of a gustatory neuron in the amygdala was recorded to that solution and to four other taste stimuli. This experiment was conducted a total of 14 times in two monkeys. The responses of individual neurons to the satiety stimuli were suppressed by as little as 1%, and as much as 100% by the induction of satiety (mean suppression = 58%). Nine of the 14 cells responded to the satiety solution with excitation, and their responses were suppressed by a mean of 62% by satiety. Five neurons responded with inhibition, and their responses were suppressed by a mean of 50%. Responses to other taste stimuli, not associated with satiety, were affected to a lesser extent. The amygdala is a taste relay between the primary gustatory cortex, where satiety has no influence on responses to taste stimuli, and the lateral hypothalamic area where the effect of satiety is total. The data presented here indicate that the amygdala is a functional as well as anatomical intermediary between these two areas, and serves as a stage in the process through which sensory stimuli are imbued with motivational significance.     

Cortical responses to electrical and gustatory stimuli in the rabbit.

The distribution of surface positive cortical potentials evoked by electrical stimulation of the chorda tympani, glossopharyngeal and lingual nerves which innervate the tongue was mapped in rabbits. All projections were bilateral. Judging from the extent of the cortical response area and the amplitude and latency of the responses, the major projection of the chorda tympani was ipsilateral, whereas that of the lingual and the glossopharyngeal nerves was contralateral. Both the chorda tympani and the glossopharyngeal nerve project to a confined area in the insular cortex and the lingual nerve projects to the appropriate part of the somatotopic pattern of somatic sensory area I. Further, a single unit study was undertaken to characterize the response of units in the cerebral cortex which was induced by gustatory stimulation of the anterior tongue, Twenty-four gustatory units were found in the insular cortex and the claustrum. The gustatory units were divided into an early response type (21 units) and a late response type (3 units) based on latency measurements. Gustatory units were also classified according to discharge patterns into excitation type (21 units) and inhibition type (4 units). Eleven units responded to 1 or 2 kinds of conventional taste stimuli, and 13 units responded to more than 3 different taste stimuli. Sensitivities of cortical units to the 4 conventional taste stimuli were found to be mutually independent and randomly distributed among cortical units. The frequency of discharges increased in the excitation type units and decreased in the inhibition type units monotonically with the excitation type units and decreased in the inhibition type units monotonically with an increse of NaCl concentration exfept at the highest concentrations.     

Taste neurons in the cortex of the alert cynomolgus monkey.

The activity of single neurons in the gustatory cortex of alert cynomolgus monkeys was analyzed. Taste-evoked activity in response to the four prototypical taste stimuli was recorded from a cortical gustatory area comprising the frontal operculum and adjoining anterior insula. Spontaneous activity for 364 gustatory neurons was 3.9 +/- 4.9 (mean +/- SD) spikes/s. Mean net (gross minus spontaneous) discharge rates for all gustatory neurons were: 1.0 M glucose = 4.9 +/- 11.6, 0.3 M NaCl = 3.2 +/- 7.1, M quinine HCl = 2.6 +/- 5.8, and 0.01 M HCl = 1.7 +/- 4.6. The results from intensity-response functions imply that the perception of each basic taste quality in the nonhuman primate is based on the activity of the appropriate neural subgroup rather than on the mean activity of all taste cells. Therefore a more meaningful index of the effectiveness of a stimulus may be the discharge rate it evokes from the subset of gustatory neurons for which it is the best stimulus. Glucose was the best stimulus for 142 cells (including ties), from which it elicited a mean net response of 10.3 spikes/s; NaCl was best for 107 neurons which gave a mean 8.7 spikes/s; quinine HCl evoked 6.2 spikes/s from the 74 cells that responded best to it; HCl elicited 5.9 spikes/s from the 49 neurons for which it served as best stimulus. The response characteristics of cortical taste cells indicate heterogeneous features, and significantly different patterns from those reported in other nonchemical sensory systems.     

Gustatory cortex in the rat. I. Physiological properties and cytoarchitecture

The precise cytoarchitectural localization of taste-elicited cortical responses in the rat was studied using a combination of anatomical and physiological techniques. Multi-unit responses to tongue tactile, thermal and gustatory stimuli were recorded along 97 electrode penetrations positioned parallel to the lateral convexity of the brain and marking lesions were placed at the sites of transitions in these functional properties. Lesions made at sites that received different sensory inputs were consistently located within different cytoarchitectural subdivisions. In this manner, taste cortex in the rat was localized to the agranular insular cytoarchitectural region, in contrast to its traditional assignation to granular insular cortex. Instead, tongue temperature was found to be represented in the cortical area previously termed gustatory, i.e., in ventral granular cortex where layer IV attenuates.