Centrifugal inputs modulate taste aversion learning associated parabrachial neuronal activities

Our previous studies have demonstrated that gustatory neurons in the parabrachial nucleus (PBN) show altered responses after the acquisition of conditioned taste aversion (CTA) to NaCl. The present study was conducted 1) to examine centrifugal influences on the altered gustatory activity of CTA-trained rats, and 2) to evaluate the role of amiloride-sensitive (ASN) and -insensitive NaCl (AIN) best units in coding the taste of NaCl. Animals were separated into 2 groups: a CTA group that had acquired taste aversion to 0.1 M NaCl and a control group that underwent pseudoconditioning before the recording experiment. Single-neuron activity, in 2 separate series of experiments, was extracellularly recorded in anesthetized rats. In the stimulation studies, the effects of electrical stimulation of the gustatory cortex (GC) or the central nucleus of amygdala (CeA) were examined on firing of PBN taste units. CeA stimulation produced excitatory effect in significantly more neurons in the CTA group (n = 8) than in the control group (n = 1). Furthermore, ASN-best units in the CTA group showed larger responses to NaCl than similar units in the control group. In the decerebration experiment, there was no statistical difference among the taste responses between the 2 groups in any best-stimulus category. These results suggest that CTA conditioning uses an effective central amygdaloid input to modulate activity of gustatory neurons in the PBN. Data also substantiate that amiloride-sensitive components of NaCl-best neurons play a critical role in the recognition of distinctive taste of NaCl.     

Pontine gustatory activity is altered by electrical stimulation in the central nucleus of the amygdala

Visceral signals and experience modulate the responses of brain stem neurons to gustatory stimuli. Both behavioral and anatomical evidence suggests that this modulation may involve descending input from the forebrain. The present study investigates the centrifugal control of gustatory neural activity in the parabrachial nucleus (PBN). Extracellular responses were recorded from 51 single PBN neurons during application of sucrose, NaCl, NaCl mixed with amiloride, citric acid, and QHCl with or without concurrent electrical stimulation in the ipsilateral central nucleus of the amygdala (CeA). Based on the sapid stimulus that evoked the greatest discharge, 3 neurons were classified as sucrose-best, 32 as NaCl-best, and 16 as citric acid-best. In most of the neurons sampled, response rates to an effective stimulus were either inhibited or unchanged during electrical stimulation of the CeA. Stimulation in the CeA was without effect in two sucrose-best neurons, nine NaCl-best neurons, and one citric acid-best neuron. Suppression was evident in 1 sucrose-best neuron, 18 NaCl-best neurons, and 15 citric acid-best neurons. In NaCl-best neurons inhibited by CeA stimulation, the magnitude of the effect was similar for spontaneous activity and responses to the five taste stimuli. Nonetheless, the inhibitory modulation of gustatory sensitivity increased the relative effectiveness of NaCl resulting in narrower chemical selectivity. For citric acid-best neurons, the magnitude of inhibition produced by CeA activation increased with an increase in stimulus effectiveness. The responses to citric acid were inhibited significantly more than the responses to all other stimuli with the exception of NaCl mixed with amiloride. The overall effect was to change these CA-best neurons to CA/NaCl-best neurons. In a smaller subset of NaCl-best neurons (n = 5), CeA stimulation augmented the responsiveness to NaCl but was without effect on the other stimuli or on baseline activity. It appears that electrical stimulation in the CeA modulates response intensity, as well as the type of gustatory information that is transmitted in a subset of NaCl-best neurons. These findings provide an additional link between the amygdala and the PBN in the control of NaCl intake, modulating the response and the chemical selectivity of an amiloride-sensitive Na+ detecting input pathway.     

Development of taste responses in the rat parabrachial nucleus

Extracellular responses from neurons in the parabrachial nuclei (PBN) were studied in rats 4 days old to adulthood during chemical stimulation of the tongue with monochloride salts, citric and hydrochloric acids, sucrose, sodium saccharin, and quinine hydrochloride. Multiunit taste responses were recorded in rats at 4-7 days of age and single-unit responses were recorded from 121 neurons in four other age groups of 14-20 days, 25-35 days, 50-60 days, and adults. PBN neurons in rats 4-7 days old consistently responded to 0.1 M solutions of NH4Cl and NaCl, to 0.5 M solutions of NH4Cl, NaCl, and KCl, and to 1.0 M sucrose, 0.1 M sodium saccharin, 0.1 M citric acid, and 0.1 N HCl. They often did not respond, however, to 0.1 M KCl and 0.01 M quinine hydrochloride. Single PBN neurons in rats 14 days old and older characteristically responded to all stimuli, which consisted of 0.1 and 0.5 M salts, acids, sucrose, sodium saccharin, and quinine hydrochloride. Thus no developmental differences occurred in the number of stimuli to which neurons responded after rats were 14 days old. With the exception of responses to hydrochloric acid, there were significant increases in response frequencies to all stimuli after 14 days of age. Average response frequencies to NH4Cl and citric acid increased after 20 days of age and those to NaCl, LiCl, KCl, sucrose, sodium saccharin, and quinine hydrochloride increased after 35 days of age. Average response frequencies for hydrochloric acid did not alter after 14 days of age. The proportion of single PBN neurons that responded maximally to specific monochloride salts did not change during development. Most single neurons in all age groups responded equally well to NH4Cl, NaCl, and LiCl. No PBN neuron responded maximally to KCl. Developmental differences in response frequencies of third-order gustatory neurons in the PBN generally reflect developmental response changes in first-order neurons of the chorda tympani nerve and second-order neurons of the solitary nucleus. However, unique developmental changes are evident in the PBN. Thus the ontogenetic changes that occur in PBN responses likely relate to modifications of lower-order peripheral and central nervous system afferents and peripheral receptor sensitivities.     

Activity in the hypothalamus, amygdala, and cortex generates bilateral and convergent modulation of pontine gustatory neurons

Evidence suggests that centrifugal modulation of brain stem gustatory cells might play a role in the elaboration of complex taste-guided behaviors like conditioned taste aversion and sodium appetite. We previously showed that activity in one forebrain area, the central nucleus of the amygdala (CeA), increased the chemical selectivity of taste cells in the parabrachial nucleus (PBN). The present study investigates how activity in 2 other similarly interconnected forebrain sites, the lateral hypothalamus (LH) and gustatory cortex (GC), might influence PBN gustatory processing in rats. The potential convergence of descending inputs from these sites, as well as the CeA, was also evaluated. After anesthesia (35 mg/kg Nembutal ip), 70 PBN gustatory neurons were tested before, during, and after electrical stimulation of these forebrain sites, while responding to 0.3 M sucrose, 0.1 M NaCl, 0.01 M citric acid, and 0.003 M QHCl. Although each forebrain site modulated taste-evoked responses, more PBN neurons were influenced by stimulation of the GC (67%) and CeA (73%) than of the LH (48%). Activation of cortex (71%) and amygdala (85%) most often produced inhibition, whereas inhibition and excitation occurred equally often during hypothalamic stimulation. Of the neurons tested for convergence (n = 60), 88% were influenced by > or =1 of the 3 sites. Twenty were modulated by stimulation at all 3 sites and another 17 by 2 of the 3 sites. The net effect of centrifugal modulation was to sharpen the across-stimulus response profiles of PBN cells, particular with regard to the NaCl- and citric acid-best cells.     

Neural representation of salts in the rat solitary nucleus: brain stem correlates of taste discrimination

One mechanism of salt taste transduction by gustatory receptor cells involves the influx of cations through epithelial sodium channels that can be blocked by oral application of amiloride. A second mechanism is less clearly defined but seems to depend on electroneutral diffusion of the salt through the tight junctions between receptor cells; this paracellular pathway is insensitive to amiloride. Because the first mechanism is more sensitive to sodium salts and the second to nonsodium salts, these peripheral events could underlie the ability of rats to discriminate sodium from nonsodium salts on the basis of taste. Behavioral experiments indicate that amiloride, at concentrations that are tasteless to rats, impairs a rat's ability to discriminate NaCl from KCl and may do so by making both salts taste like KCl. In the present study, we examined the neural representation of NaCl and KCl (0.05-0.2 M), and mixtures of these salts with amiloride (0, 3, and 30 microM), to explore the neural correlates of this behavioral result. NaCl and KCl were represented by distinct patterns of activity in the nucleus of the solitary tract. Amiloride, in a concentration-dependent manner, changed the pattern for NaCl to one more characteristic of KCl, primarily by reducing activity in neurons responding best to NaCl and sucrose. The effect of amiloride concentration on the response to 0.1 M NaCl in NaCl-best neurons was virtually identical to its effect on behavioral discrimination performance. Modeling the effects of blocking the amiloride-insensitive pathway also resulted in highly similar patterns of activity for NaCl and KCl. These results suggest that activity in both the amiloride-sensitive and -insensitive pathways is required for the behavioral discrimination between NaCl and KCl. In the context of published behavioral data, the present results suggest that amiloride-sensitive activity alone is not sufficient to impart a unique signal for the taste of sodium salts.     

Gustatory responses of neurons in the nucleus of the solitary tract of behaving rats.

1. The activity of 117 single neurons was recorded in the rostral nucleus of the solitary tract (NST) and tested with each of four standard chemical stimuli [sucrose, NaCl, citric acid, and quinine HCl (QHCl)] and distilled water in awake, behaving rats. In 101 of these neurons, at least one sapid stimulus elicited a significant taste response. The mean spontaneous rate of the taste neurons was 4.1 +/- 5.8 (SD) spike/s. The mean response magnitudes were as follows: sucrose, 10.6 +/- 11.7; NaCl, 8.6 +/- 14.6; citric acid, 6.2 +/- 7.8; and QHCl, 2.4 +/- 6.6 spikes/s. 2. On the basis of their largest response, 42 taste neurons were classified as sucrose-best, 25 as NaCl-best, 30 as citric acid-best, and 4 as QHCl-best. The mean spontaneous rates for these categories were 4.9 +/- 6.2 for sucrose-best cells, 5.8 +/- 7.4 for NaCl-best, 1.6 +/- 2.0 for citric acid-best, and 5.8 +/- 6.0 spikes/s for QHCl-best. The spontaneous rate of the citric acid-best neurons was significantly lower than that of the other categories. 3. At the standard concentrations, 45 taste cells (44.6%) responded significantly to only one of the gustatory stimuli. Of the 30 acid-best neurons, 23 (76.7%) responded only to citric acid. For sucrose-best cells, specific sensitivity was less common (18/42, 42.9%), and for NaCl-best neurons, it was relatively uncommon (3/25, 12%). One of the 4 QHCl-best neurons was specific. In a concentration series, more than one-half of the 19 specific neurons tested responded to only one chemical at any strength. 4. The mean entropy for the excitatory responses of all gustatory neurons was 0.60. Citric acid-best cells showed the least breadth of responsiveness (0.49), sucrose-best cells were somewhat broader (0.56), but NaCl-best and QHCl-best cells were considerably less selective (0.77 and 0.79, respectively). Inhibition was observed infrequently and never reached the criterion for significance. 5. In the hierarchical cluster analysis, the four largest clusters segregated neurons primarily by best-stimulus category. The major exception to this was a group of sucrose-best neurons that also responded to NaCl and were grouped with the NaCl-best neurons. In a two-dimensional space, the specific taste neurons, those that responded to only one of the four standard sapid stimuli, remained in well-separated groups. These specific groups, however, were joined in a ring-like formation by other neurons that responded to more than one of the sapid stimuli.(ABSTRACT TRUNCATED AT 400 WORDS)     

Altered NaCl taste responses precede increased NaCl ingestion during Na(+) deprivation

It has been suggested that Na(+) deficiency alters the sensitivity of taste receptors, thereby rendering NaCl solutions more palatable or preferred and more likely to be ingested. Increased ingestion of concentrated NaCl solutions by rats during dietary Na(+) deprivation occurs only after approximately 8-10 days. To determine whether changes in gustatory responses mediate the deprivation-induced NaCl ingestion (salt appetite), we evaluated taste responses to a range of NaCl concentrations before, during, and after dietary Na(+) deprivation. Rats were trained to lick rapidly in short-duration (10 s) tests by mixing NaCl solutions in a dilute sucrose solution. This method elicited consistent, interpretable rates of licking, even of normally avoided NaCl concentrations, without the necessity of depriving the rats of water. The licking rate increased after dietary Na(+) deprivation of only 2 days, increased further after 5 days of Na(+) deprivation and, after 10 days, was not different from that after 2 days. These results suggest that a change in the response to NaCl taste, as evidenced by increased rates of licking during short-access tests, occurred after 2 days of dietary Na(+) deprivation. In contrast, a significant increase in the 24-h ingestion of a concentrated NaCl solution occurred only after approximately 1 week of maintenance on Na(+)-deficient chow. Thus, it is unlikely that a delayed change in the response to NaCl taste to more palatable or preferred underlies the delayed increase in 24-h NaCl intake during dietary Na(+) deprivation.     

Parabrachial nuclei damage in infant rats produces residual deficits in gustatory preferences/aversions and sodium appetite

Ten-day-old rats sustained bilateral electrolytic lesions of the parabrachial nuclei in the pons (PBN). Growth measures and tests of sensorimotor, feeding and drinking behaviors, sodium appetite, and gustatory capacities were made between age 1 and 150 days. PBN rats displayed a transient period of attentuated suckling, as evidenced by body weight loss. When tested soon after weaning, PBN rats were hyperdipsic in response to cellular dehydration and during food deprivation. This effect, however, was temporary. When tested as adults, PBN rats were hypodipsic in response to extracellular fluid volume depletion, they displayed alterations in sodium appetite, showed “exaggerated” preferences and aversions to saccharin and NaCl solutions, and they displayed attentuated quinine aversions. These results are generally similar to the behaviors of rats sustaining more central gustatory pathway lesions as adults. The functional significance of the PBN in the developing rat for preference/aversion and sodium appetite behaviors are discussed.     

Responses from parabrachial gustatory neurons in behaving rats

1. The responses of a total of 70 single neurons were recorded from the parabrachial nuclei (PBN) in awake rats. In 59 neurons, sapid stimuli (0.5 ml) elicited significant taste responses. Of these 59 neurons, 10 also had significant responses to water. The mean spontaneous rate of the taste neurons was 13.4 +/- 6.9 (SD) spikes/s. Of the remaining 11 neurons, 9 responded significantly only to water; 2 had no significant responses to the standard fluid stimuli. 2. Based on the magnitude of their response to our four standard stimuli, the taste neurons were classified as follows: 42 NaCl-best, 14 sucrose-best, 2 citric acid-best, and 1 QHCl-best. Of these, 25 responded only to one of four sapid stimuli; 20 of these specific cells responded only to NaCl. All the remaining 34 neurons responded to two or more of the four sapid stimuli, with NaCl and sucrose responsiveness dominant. For the 59 taste neurons, the mean entropy for the absolute value of the responses was 0.68; for the excitatory activity alone, it was 0.58. 3. The mean responses to NaCl and sucrose concentration series increased monotonically. Except at the lowest concentration, responses to citric acid also increased monotonically, but with a lower slope. Mean responses to QHCl, however, remained stable or even decreased with increasing concentration. Thus the power functions for the NaCl and sucrose intensity-response series were higher than those of citric acid and QHCl. 4. A hierarchical cluster analysis of 59 parabrachial neurons suggested four different categories: NaCl-best, sucrose-best, citric acid-best, and QHCl-best. These categories were less evident in the two-dimensional space produced by multidimensional analysis, because the positions of NaCl- and sucrose-best neurons formed a continuum in which neural response profiles change successively from sucrose-specific to NaCl-specific. 5. The results were consistent with previous anatomic and neurophysiological data suggesting convergence in the medulla of sensory input from receptors in the nasoincisor ducts (NID) and on the anterior tongue (AT). Taste buds in the NID respond preferentially to sucrose, whereas those on the AT respond more to NaCl.(ABSTRACT TRUNCATED AT 400 WORDS)     

Amiloride sensitivity of the chorda tympani response to sodium chloride in sodium-depleted Wistar rats.

Peripheral gustatory mechanisms that may contribute to the expression of sodium (Na) appetite have been a focus of interest for many years. Because amiloride-sensitive Na transport is involved in the generation of neural signals in response to NaCl stimulation, the present study assessed whether changes in amiloride sensitivity of the neural response to NaCl accompany the induction of a Na appetite in the rat. Na deprivation was achieved by acute depletion with the diuretic furosemide. The magnitude of the whole-nerve chorda tympani response to 0.5 M NaCl was reduced in Na-depleted, compared with Na-replete, rats, which provides qualified support for previous reports that the induction of a Na appetite is associated with reduced neural responses to NaCl. However, changes in sensitivity to the specific Na channel blocker amiloride hydrochloride as a result of Na depletion were not evident. These findings suggest that the behavioral and neural changes that occur after Na depletion are not based on changes in amiloride sensitivity in the taste bud.     

Neural representation of the taste of NaCl and KCl in gustatory neurons of the hamster solitary nucleus.

NaCl and KCl are monovalent salts that can be discriminated behaviorally by hamsters on the basis of their tastes. We examined the effects of the passive Na+ channel blocker amiloride on responses to both of these salts in 34 taste-responsive neurons of the nucleus of the solitary tract (NST) in the hamster. The effects of amiloride were assessed with two different, commonly employed stimulus protocols. Additionally, concentration-response functions for each salt were measured in 37 neurons. Cells were characterized by their best response to (in M) 0. 03 NaCl, 0.1 sucrose, 0.003 HCl, 0.001 quinine hydrochloride, and 0. 1 KCl. In neurons classified as NaCl-best, amiloride reversibly blocked responses to both NaCl and KCl. In neurons classified as HCl-best, amiloride had no effect on either stimulus. In sucrose-best neurons, amiloride blocked the response to NaCl but not KCl. These results support the hypothesis that both salts are transduced by at least two different receptor mechanisms. In the NST, information arising from these different inputs is maintained in discrete populations of neurons. In addition to differences in amiloride sensitivity, the cell types also differed in their responses to the salts across concentration. At midrange salt concentrations, NaCl-best neurons were far more responsive to NaCl than KCl, whereas HCl- and sucrose-best neurons responded equivalently to the two salts at all concentrations. Because NaCl- and HCl-best cells cannot by themselves distinguish NaCl from KCl, it is the relative activity across these cell types that comprises the code for taste discrimination.     

Synchronized discharge of taste neurons recorded simultaneously in rat parabrachial nucleus

1. Cross-correlation analysis was made in the taste-sensitive neuron pairs recorded simultaneously from the parabrachial nucleus (PBN) of rats. Three indexes were adopted to evaluate the activities of the taste neurons; namely, 1) spike response density (RD value), which is the net spike density to stimulation with the four basic tastes; 2) the frequency of correlated discharges (FC value in spikes per second), which was determined by measuring the area of the peak appearing in the cross-correlogram (CC) during application of the test fluids; and 3) the weight of correlated discharge (WC = FC/RD), which shows the relative importance of correlated discharges in the taste signals delivered by a component neuron of a given pair. 2. In 11 of 23 pairs, the CCs exhibited peaks during stimulation with tastants. These 11 pairs, which were recorded in the pontine taste area, were composed of 18 NaCl-best (most sensitive to NaCl) and 4 HCl-best neurons. In eight pairs, the best-stimulus of both of the component neurons was NaCl, and it was HCl in one pair (homo-type pairs). The remaining two pairs were composed of an NaCl-best and an HCl-best neuron (hetero-type pairs). 3. In eight homo-type pairs (7 NaCl-best pairs and 1 HCl-best pair), each pair exhibited the maximal FC value during stimulation with the best-stimulus of the component neurons (2.3 less than or equal to maximal FC less than or equal to 26.6 Hz). In the remaining three pairs, the maximal FC values were low (0.8-1.9 Hz).(ABSTRACT TRUNCATED AT 250 WORDS)     

Perinatal exposure to a high NaCl diet increases the NaCl intake of adult rats

To determine the effect of differences in perinatal NaCl exposure on NaCl intake, adult Sprague-Dawley female rats were maintained on diets containing either 0.12, 1.0, or 3% NaCl throughout pregnancy and lactation. The offspring were continued on the these same diets to 30 days postpartum. Thereafter, all offspring were fed the same basal diet containing 1% NaCl. At 90 days of age, the adult offspring were placed in metabolism cages for 7 days and fed 1% NaCl chow for days 1-2, and 0% NaCl chow for days 3-7. On days 6-7, the animals were free to consume both water and 0.3 M NaCl. When dietary NaCl was available, adult rats exposed perinatally to the high NaCl diet excreted significantly more sodium on days 1-2 and 6-7 than did the rats exposed to either the mid or low NaCl diets. There were no differences in sodium excretion during sodium deprivation on days 3-5. The 0.3 M NaCl intake of the high NaCl-exposed rats was also significantly greater than the intake of the mid and low NaCl-exposed rats. In another group of adult rats, exposed perinatally to either a low or high NaCl diet, the spontaneous 24-hr intake of water and 0.3 M NaCl was measured after repeated episodes of acute sodium depletion. Sodium depletion was induced by 48 hr of dietary sodium deprivation combined with a single subcutaneous injection of 5 mg furosemide. Acute sodium depletion was found to augment existing differences in NaCl intake between low and high NaCl-exposed rats.(ABSTRACT TRUNCATED AT 250 WORDS)     

Responses to taste stimulation in the ventroposteromedial nucleus of the thalamus in rats

Extracellular action potentials were recorded from 73 neurons in the parvicellular division of the ventroposteromedial (VPMpc) nucleus of the thalamus of anesthetized Wistar rats during gustatory, thermal, and tactile stimulation of the whole oral cavity. The stimulus array consisted of 16 room-temperature (23 degrees C) sapid stimuli, distilled water at three temperatures (0, 23, and 37 degrees C), and 0.1 M NaCl at three temperatures (0, 23, and 37 degrees C). Among all 151 neurons isolated in VPMpc, 9% responded exclusively to taste, 33% to taste and temperature, none to taste and touch, but 6% to all three modalities. Discharge rates evoked by the basic tastants were 13.8 +/- 1.6 (SD) spikes/s for 0.1 M NaCl, 9.3 +/- 1.4 spikes/s for 0.01 M HCl, 5.1 +/- 0.9 spikes/s for 0.5 M sucrose, and 4.3 +/- 0.6 spikes/s for 0.01 M quinine HCl. Water evoked mean responses at 0, 23, and 37 degrees C of 9.9 +/- 1.5, 0.6 +/- 0.4, and 1.3 +/- 0.9 spikes/s, respectively. The mean firing rate evoked by 37 and 0 degrees C NaCl was 15.0 +/- 2.4 and 17.0 +/- 2.8 spikes/s, respectively. The exponent of the NaCl concentration-response power function was 0.39. Thalamic taste cells were broadly tuned. The mean breadth-of-tuning coefficient for these 73 gustatory cells was 0.79 +/- 0.02. Two cells responded predominantly with inhibition, which accounted for the majority of inhibitory responses. The taste neurons were statistically divisible into three groups: sodium-oriented (n = 40), acid-oriented (n = 12), and sugar-oriented (n = 17). Four additional bitter-oriented neurons were not closely enough related to be defined as a group and were considered outliers. The sodium-oriented group could be divided into three statistically distinct subgroups, differing in the specificity of their responses to NaCl. With respect to polymodal sensitivity, spontaneous rate, evoked response rates, signal-to-noise ratio, proportions of cells responding best to basic tastants, taste neuron groups, taste spaces, and temporal responses, VPMpc neurons have characteristics that are intermediate between those of parabrachial and cortical gustatory neurons.     

Effects of dietary NaCl deprivation during early development on behavioral and neurophysiological taste responses

In order to determine whether the gustatory system can be modified by restricting dietary NaCl during early development, neurophysiological taste responses were recorded in rats at various times after deprivation, and behavioral taste preferences were measured in adults. Rats deprived of dietary NaCl from the third day of gestation to 12 days postnatally and then placed on a NaCl-replete diet had chorda tympani nerve responses similar to those of nondeprived rats when recordings were made at 28 days of age and older; however, preferences for NaCl solutions over water were significantly less than those of controls when tested at adulthood. NaCl deprivation in pups from the third day of gestation to approximately 35 days postnatally resulted in altered chorda tympani nerve responses to NaCl but not to other stimuli such as NH4Cl and KCl. Therefore, restriction of dietary NaCl at a period in the rat's development when peripheral and central taste responses are changing results in short-term alterations in peripheral neural responses and in long-term changes in preference behaviors.     

Development of taste responses in rat nucleus of solitary tract

Extracellular responses from neurons in the nucleus of the solitary tract (NST) were studied in rats aged 5 days to adulthood during chemical stimulation of the tongue with monochloride salts, citric and hydrochloric acids, sucrose, sodium saccharin, and quinine hydrochloride. Multiunit taste responses were recorded in rats at 5-7 days of age and single-unit responses were recorded from 111 neurons in four other age groups of 14-20 days, 25-35 days, 50-60 days, and adult. NST neurons in rats aged 5-7 days consistently responded to relatively high concentrations (0.5 M) of NH4Cl and KCl and to citric and hydrochloric acid. However, they often did not respond to 0.5 M NaCl or to 0.1 M NH4Cl. Single NST neurons in rats aged 14 days and older characteristically responded to all 0.1 and 0.5 M salts and to both acids. At least 75% of neurons also responded to sucrose and sodium saccharin, and 46% responded to all of these stimuli and quinine hydrochloride. After 14 days, no developmental changes occurred in the number of stimuli to which neurons responded. There were substantial developmental alterations in the response magnitudes to some chemical stimuli. Average response frequencies increased after 35 days of age for 0.1 and 0.5 M NaCl, LiCl, KCl, and for sucrose and sodium saccharin. Response frequencies for NH4Cl, citric and hydrochloric acid, and quinine hydrochloride, however, did not change throughout development. The proportion of single NST neurons that responded maximally to specific monochloride salts did not change during development. Most single neurons in all age groups responded equally well to NH4Cl, NaCl, and LiCl. No NST neuron responded maximally to KCl. There were also no developmental differences in response latencies in rats aged 14 days and older. Response frequencies of second-order NST neurons generally reflect changes in responses from the primary afferent, chorda tympani fibers, throughout development; however, the increases in salt response frequencies from NST neurons occur comparatively later in development. Furthermore, at all ages, the taste responses to monochloride salts include higher response frequencies and a general loss in response specificity in NST compared to chorda tympani neurons.(ABSTRACT TRUNCATED AT 400 WORDS)     

Gustatory neural coding in the monkey cortex: stimulus intensity

1. We analyzed the activity of single neurons in gustatory cortex of alert cynomolgus monkeys in response to a range of stimulus intensities. Chemicals were deionized water, fruit juice, and several concentrations of the four prototypical taste stimuli: 10(-3)-1.0 M glucose, 10(-3)-1.0 M NaCl, 10(-4)-3 x 10(-2) M HCl, and 10(-5)-3 x 10(-3) M quinine HCl. 2. Taste-evoked responses could be recorded from a cortical gustatory area that measured 2.5 mm in its anteroposterior extent, 6.0 mm dorsoventrally, and 3.0 mm mediolaterally. Taste-responsive cells constituted 62 (3.7%) of the 1,661 neurons tested. Nongustatory cells gave responses associated with mouth movement (10.1%), somatosensory stimulation (2.2%), and approach or anticipation (0.9%). 3. Intensity-response functions were determined across 62 gustatory neurons. Neural thresholds for each stimulus quality conformed well to human psychophysical thresholds. Mean discharge rate was a direct function of stimulus concentration for glucose, NaCl, and quinine HCl. The most effective of the basic stimuli was glucose. 4. Power function exponents were calculated from the responses of neural subgroups most responsive to each basic stimulus. Those for glucose, NaCl, and quinine were within the range of psychophysically derived values. Thus the perceived intensity of each basic quality is presumably based on the activity of the appropriate neural subgroup rather than on the mean activity of all taste cells. 5. The mean breadth-of-tuning (entropy) coefficient for 62 gustatory neurons was 0.65 (range, 0.00-0.98). 6. There was no clear evidence of chemotopic organization in the gustatory cortex. 7. An analysis of taste quality indicated that sweet stimuli evoked patterns of activity that were clearly distinct from those of the nonsweet chemicals. Among the latter group, NaCl was differentiable from HCl and quinine HCl, whose patterns were closely related. 8. The response characteristics of cortical taste cells imply gustatory thresholds and intensity-response functions for the nonhuman primate that conform well to those reported in psychophysical studies of humans, reinforcing the value of this neural model for human taste intensity perception.     

Taste responses in the parabrachial pons of decerebrate rats

1. Behavioral studies have shown that chronic decerebrate rats retain the capacity to react appropriately to gustatory stimuli (12), but do not form taste-illness associations (13). Little is known, however, about the effects of decerebration on the processing of gustatory information. The present experiment was designed to investigate this issue in the parabrachial nucleus of the pons (PbN). 2. Rats were decerebrated at the supracollicular level under ketamine and ether anesthesia and were prepared for electrical recording in the PbN. Thereafter, animals were maintained under Flaxedil, and wound edges were frequently treated with lidocaine. Heart rate, core temperature, and CO2 were monitored throughout each experiment. Control subjects were treated identically, except that they were not decerebrated. 3. Sapid solutions of NaCl (0.1 M), HCl (0.01 M), sucrose (0.5 M), saccharin sodium (0.004 M), and quinine HCl (.01 M) were used as taste stimuli. After a 10-s base line, each stimulus was bathed over the tongue for 10 s followed by a 10-s wait and a 20-s rinse of distilled water. The intertrial interval was at least 2 min. 4. Gustatory responses from 32 parabrachial units in 13 decerebrate rats were recorded. These were compared with responses in 31 units from the PbN of 16 intact rats. 5. Analysis of response profiles of PbN units in decerebrate rats showed that these units produced smaller responses to NaCl and HCl and larger responses to saccharin sodium compared with units in intact rats. 6. Despite changes in response magnitude, the temporal patterns of response (phasic-tonic relationships) were not different in PbN units in decerebrate rats compared with controls. Differences in the length of responses were, however, apparent. Responses to saccharin sodium were longer, response to NaCl, HCl and sucrose were shorter, and responses to quinine HCl were unchanged. 7. Results of a multidimensional scaling analysis of the response profiles across units showed that "taste spaces" for decerebrate and intact rats were similar. Units in each group were meaningfully placed near stimuli that evoked the best response in a given unit. Units that did not respond well to any stimulus were placed close together regardless of their best stimulus in both taste spaces. 8. Responses to the termination of the taste stimulus (OFF-responses) were observed in PbN units in the decerebrate rat but not in units from the intact rat. Twenty-one OFF-responses were recorded in 14 units; 6 of these occurred in the absence of a response to the stimulus.(ABSTRACT TRUNCATED AT 400 WORDS)     

NaCl味觉刺激对钠缺乏大鼠臂旁核c-fos表达的影响

目的:探讨臂旁核(PBN)在钠平衡行为调节中的作用。方法:成年雄性SD大鼠20只,分为对照组、糖精溶液口腔灌流组(Sac组)、NaCl溶液口腔灌流组(NaCl组)、注射furosemide后NaCl溶液口腔灌流组(Furo-NaCl组),观察大鼠对味刺激的摄取反应和各组PBN各亚核内c-fos表达水平差异。结果:正常大鼠对糖精溶液显示嗜好性反应,对NaCl溶液显示厌恶性反应,但钠缺乏大鼠对NaCl也表现出嗜好性摄取反应。Sac组PBN各亚核内c-fos表达水平均高于对照组;NaCl刺激增加内侧亚核(ms)内的c-fos表达,但减少外部外侧亚核(els)内的c-fos表达;Furo- NaCl组els和中央外侧亚核(cls)内c-fos表达显著高于其他3组,ms内的c-fos表达低于NaCl组,但仍高于对照组和Sac组。结论:els和cls在钠平衡的味觉嗜好调节中发挥重要作用。     

Central gustatory lesions: II. Effects on sodium appetite, taste aversion learning, and feeding behaviors.

Intake and taste reactivity tests were used to determine the effects of bilateral lesions of the gustatory portions of the nucleus of the solitary tract (NST), the parabrachial nucleus (PBN), and the ventral posteromedial nucleus of the thalamus (VPMpc) on several complex ingestive behaviors. In the 1st experiment, lesions of the PBN and the NST blocked, and VPMpc lesions impaired, the behavioral expression of salt appetite. In the 2nd experiment, alanine was paired with injections of LiCl. Control rats as well as rats with NST and VPMpc lesions acquired the taste aversion, but rats with PBN lesions did not. In the 3rd experiment, all animals increased their food intake after injections of 2 U/kg insulin and 250 mg/kg 2-deoxy-D-glucose, and their food intake was suppressed after nutritive stomach loads.