Kv4 channel expression and kinetics in GABAergic and non-GABAergic rNST neurons.

The rostral nucleus of the solitary tract (rNST) serves as the first central relay in the gustatory system. In addition to synaptic interactions, central processing is also influenced by the ion channel composition of individual neurons. For example, voltage-gated K+ channels such as outward K+ current (IA) can modify the integrative properties of neurons. IA currents are prevalent in rNST projection cells but are also found to a lesser extent in GABAergic interneurons. However, characterization of the kinetic properties of IA, the molecular basis of these currents, as well as the consequences of IA on spiking properties of identified rNST cells is lacking. Here, we show that IA in rNST GABAergic (G+) and non-GABAergic (G-) neurons share a common molecular basis. In both cell types, there was a reduction in IA following treatment with the specific Kv4 channel blocker AmmTx3. However, the kinetics of activation and inactivation of IA in the two cell types were different with G- neurons having significantly more negative half-maximal activation and inactivation values. Likewise, under current clamp, G- cells had significantly longer delays to spike initiation in response to a depolarizing stimulus preceded by a hyperpolarizing prepulse. Computational modeling and dynamic clamp suggest that differences in the activation half-maximum may account for the differences in delay. We further observed evidence for a window current under both voltage clamp and current clamp protocols. We speculate that the location of Kv4.3 channels on dendrites, together with a window current for IA at rest, serves to regulate excitatory afferent inputs.NEW & NOTEWORTHY Here, we demonstrate that the transient outward K+ current IA occurs in both GABAergic and non-GABAergic neurons via Kv4.3 channels in the rostral (gustatory) solitary nucleus. Although found in both cell types, IA is more prevalent in non-GABAergic cells; a larger conductance at more negative potentials leads to a greater impact on spike initiation compared with GABAergic neurons. An IA window current further suggests that IA can regulate excitatory afferent input to the nucleus.

Inhibitory modulation of optogenetically identified neuron subtypes in the rostral solitary nucleus.

Inhibition is presumed to play an important role in gustatory processing in the rostral nucleus of the solitary tract (rNST). One source of inhibition, GABA, is abundant within the nucleus and comes both from local, intrasolitary sources and from outside the nucleus. In addition to the receptor-mediated effects of GABA on rNST neurons, the hyperpolarization-sensitive currents, Ih and IA, have the potential to further modulate afferent signals. To elucidate the effects of GABAergic modulation on solitary tract (ST)-evoked responses in phenotypically defined rNST neurons and to define the presence of IA and Ih in the same cells, we combined in vitro recording and optogenetics in a transgenic mouse model. This mouse expresses channelrhodopsin 2 (ChR2) in GAD65-expressing GABAergic neurons throughout the rNST. GABA positive (GABA+) neurons differed from GABA negative (GABA-) neurons in their response to membrane depolarization and ST stimulation. GABA+ neurons had lower thresholds to direct membrane depolarization compared with GABA- neurons, but GABA- neurons responded more faithfully to ST stimulation. Both IA and Ih were present in subsets of GABA+ and GABA- neurons. Interestingly, GABA+ neurons with Ih were more responsive to afferent stimulation than inhibitory neurons devoid of these currents, whereas GABA- neurons with IA were more subject to inhibitory modulation. These results suggest that the voltage-gated channels underlying IA and Ih play an important role in modulating rNST output through a circuit of feedforward inhibition.

Regulation of Rostral Nucleus of the Solitary Tract Responses to Afferent Input by A-type K+ Current.

Responses in the rostral (gustatory) nucleus of the solitary tract (rNST) are modified by synaptic interactions within the nucleus and the constitutive membrane properties of the neurons themselves. The potassium current IA is one potential source of modulation. In the caudal NST, projection neurons with IA show lower fidelity to afferent stimulation compared to cells without. We explored the role of an A-type K+ current (IA) in modulating the response to afferent stimulation and GABA-mediated inhibition in the rNST using whole cell patch clamp recording in transgenic mice that expressed channelrhodopsin (ChR2 H134R) in GABAergic neurons. The presence of IA was determined in current clamp and the response to electrical stimulation of afferent fibers in the solitary tract was assessed before and after treatment with the specific Kv4 channel blocker AmmTX3. Blocking IA significantly increased the response to afferent stimulation by 53%. Using dynamic clamp to create a synthetic IA conductance, we demonstrated a significant 14% decrease in responsiveness to afferent stimulation in cells lacking IA. Because IA reduced excitability and is hyperpolarization-sensitive, we examined whether IA contributed to the inhibition resulting from optogenetic release of GABA. Although blocking IA decreased the percent suppression induced by GABA, this effect was attributable to the increased responsiveness resulting from AmmTX3, not to a change in the absolute magnitude of suppression. We conclude that rNST responses to afferent input are regulated independently by IA and GABA.

Responsiveness of neurons in the hamster parabrachial nuclei to taste mixtures.

Responses from hamster parabrachial nuclei neurons to stimulation of the anterior tongue with sucrose, NaCl, HCl, quinine hydrochloride, and the six two-component mixtures of these stimuli were recorded. A cell's response to a mixture approached its response to the mixture's more effective component in the majority of cases, but was sometimes greater or smaller than this response. The best predictor of a neuron's response to a mixture, then, was its response to the mixture's more effective component. The single-component stimulus producing the maximum response was determined for each neuron and the response to this stimulus was compared with the responses evoked by the six mixtures. For 30% of the cells, a mixture elicited a response reliably, but only 1.1-2.1 times greater than the response to the best single-component stimulus. Thus, there were no neurons specialized to respond to these mixtures. The across-neuron patterns elicited by mixtures and the responses of best-stimulus classes to mixtures were studied for comparison with psychophysical data on taste mixtures. Mixtures were usually correlated with single-component stimuli in the mixture, but not with stimuli not in the mixture. In fact, five of the six mixtures fell directly between their components in a multidimensional scaling plot. In addition, a mixture was most effective in stimulating only those classes of neurons maximally stimulated by the mixture's components. These results correlate with psychophysical data suggesting that mixtures of taste stimuli evoke the same taste qualities as evoked by the mixture's components.

Effects of high-fat diet and gastric bypass on neurons in the caudal solitary nucleus.

Bariatric surgery is an effective treatment for obesity that involves both peripheral and central mechanisms. To elucidate central pathways by which oral and visceral signals are influenced by high-fat diet (HFD) and Roux-en-Y gastric bypass (RYGB) surgery, we recorded from neurons in the caudal visceral nucleus of the solitary tract (cNST, N=287) and rostral gustatory NST (rNST,N=106) in rats maintained on a HFD and lab chow (CHOW) or CHOW alone, and subjected to either RYGB or sham surgery. Animals on the HFD weighed significantly more than CHOW rats and RYGB reversed and then blunted weight gain regardless of diet. Using whole-cell patch clamp recording in a brainstem slice, we determined the membrane properties of cNST and rNST neurons associated with diet and surgery. We could not detect differences in rNST neurons associated with these manipulations. In cNST neurons, neither the threshold for solitary tract stimulation nor the amplitude of evoked EPSCs at threshold varied by condition; however suprathreshold EPSCs were larger in HFD compared to chow-fed animals. In addition, a transient outward current, most likely an IA current, was increased with HFD and RYGB reduced this current as well as a sustained outward current. Interestingly, hypothalamic projecting cNST neurons preferentially express IA and modulate transmission of afferent signals (Bailey, '07). Thus, diet and RYGB have multiple effects on the cellular properties of neurons in the visceral regions of NST, with potential to influence inputs to forebrain feeding circuits.

Extranuclear projections of rNST neurons expressing gustatory-elicited Fos.

Previous studies have demonstrated that gustatory stimulation evokes expression of the immediate-early gene, c-fos in the rostral division of the nucleus of the solitary tract (rNST) (Harrer and Travers [1996] Brain Res. 711:125-137; DiNardo and Travers [1997] J. Neurosci. 17:3826-3839; King et al. [1999] J. Neurosci. 19:3107-3121). The present investigation further defined the phenotype of those neurons by determining their projections, by using immunohistochemistry for the Fos protein and retrograde tracing with Fluoro-Gold. Tracer injections were made into the two major extranuclear targets of rNST, the parabrachial nucleus (PBN) and medullary reticular formation (RF). These structures are thought to play differential roles in higher-order discriminative and homeostatic (PBN) versus reflexive function (RF). After PBN injections, approximately 18% of the Fos-like immunoreactive (FLI) neurons were double-labeled; after RF injections the proportion was 9%. Because only a minority of FLI neurons appear to project to targets outside NST, this suggests that most of these cells have local, intranuclear projections. Comparable proportions of cells were double-labeled after sucrose or quinine, consistent with roles for both tastants in higher-order and reflexive function. On the other hand, regardless of stimulus, twice as many FLI neurons projected to the PBN as to the RF. This could suggest that more FLI neurons contribute to functions mediated by the ascending pathway. However, the results of a recent study prompted a different hypothesis: Because glossopharyngeal nerve section similarly devastates quinine-induced FLI and oral rejection but leaves discriminative function unimpaired, it was proposed that FLI neurons are more important in driving oral motor behavior than discrimination (King et al. [1999] J. Neurosci. 19:3107-3121). A plausible hypothesis for reconciling this apparent discrepancy is that many FLI neurons make local projections in rNST, that in turn give rise to RF connections.

Ascending and descending projections from the rostral nucleus of the solitary tract originate from separate neuronal populations.

Anterograde studies have shown that neurons within the rostral (gustatory) nucleus of the solitary tract project to the parabrachial nucleus, as well as to sites within the medulla including the reticular formation and caudal nucleus of the solitary tract. In order to determine the degree to which the same neurons contribute to both projections, injections of retrograde tracers were made simultaneously into both the parabrachial nuclei and medullary reticular formation of the rat. Only a small proportion of neurons were double labeled. Consistent with studies in hamster, labeled neurons projecting to the parabrachial nuclei in rat consisted of both stellate and elongate neurons, concentrated within the central subdivision of the rostral nucleus of the solitary tract. Injections into the medullary reticular formation also labeled both stellate and elongate neurons but these were concentrated in the ventral subdivision of the nucleus. The results of the present study demonstrate that different populations of neurons in the nucleus of the solitary tract contribute to ascending and descending pathways. This suggest a possible functional specialization within the nucleus of the solitary tract for those neurons whose output eventually reaches the forebrain compared to those neurons with local connections.

Taste receptors on the anterior tongue and nasoincisor ducts of rats contribute synergistically to behavioral responses to sucrose.

In four groups of rats, behavioral responsiveness to sucrose was tested by allowing them to lick solutions in a computer-controlled gustometer (10-s trials; 0.01-1.0 M). Rats with cautery lesions of the nasoincisor ducts (NID) behaved no differently from controls. After bilateral chorda tympani nerve (CT) section, which removes taste input from the anterior tongue (AT), rats demonstrated a marginal attenuation in their responsiveness to sucrose. Combining the two lesions, however, had the greatest effect on the concentration-response curve. By shifting the curve to the right and lowering the asymptotic licking rate, the combined lesion reduced the area under the curve by one third. The effects of the combined treatments were larger than would be predicted from the sum of either one alone. This presumably reflects the central convergence of primary afferent axons from the NID and AT. Neurophysiological data have demonstrated such convergence within the nucleus of the solitary tract.

Topographic organization of Fos-like immunoreactivity in the rostral nucleus of the solitary tract evoked by gustatory stimulation with sucrose and quinine.

Fos immunohistochemistry was used to elucidate the pattern of activation elicited by two qualitatively and hedonically distinct taste stimuli, sucrose and quinine, within the first-order gustatory relay, the rostral division of the nucleus of the solitary tract. Compared to unstimulated controls, both sucrose and quinine elicited significant increases in Fos-like immunoreactivity in the rostral central subnucleus, the region of the rostral solitary nucleus that receives the densest primary afferent input. Within the rostral central subnucleus, neurons that exhibited Fos-like immunoreactivity following quinine stimulation were concentrated medially, but neurons that exhibited Fos-like immunoreactivity following sucrose stimulation were distributed more evenly along the mediolateral axis. Despite their differential distribution, sucrose- and quinine-activated neurons also demonstrated notable intermingling. Further, the chemotopic arrangement was only partially consistent with what would be predicted if chemotopy was merely an outcome of orotopy. Our results suggest that a rough chemotopy characterizes the organization of taste responses in the nucleus of the solitary tract, and that the topographic pattern of taste afferent terminations in this nucleus is related to their chemosensitivity as well as to their peripheral spatial distribution.

Convergence of lingual and palatal gustatory neural activity in the nucleus of the solitary tract.

The responses of 54 neurons to independent sapid stimulation of 4 taste receptor subpopulations associated with: (1) anterior tongue; (2) nasoincisor ducts; (3) soft palate; and (4) foliate papillae were recorded from the nucleus of the solitary tract (NST) of the Rat. Neurons responding to stimulation of receptor subpopulations in the anterior oral cavity (anterior tongue or nasoincisor ducts) were located more rostrally in the NST than neurons responding to stimulation of receptor subpopulations in the posterior oral cavity (soft palate or foliate papillae). Half of the sampled neurons responded exclusively to stimulation of one receptor subpopulation with the remaining neurons responsive to stimulation of two or more receptor subpopulations. The most common pattern of convergence observed was between responses arising from stimulation of the taste buds on the anterior tongue and those associated with the nasoincisor ducts of the hard palate. The sensitivity of NST neurons to anterior tongue and nasoincisor duct stimulation with the 4 standard taste stimuli was determined. When stimulating the anterior tongue, the order of effectiveness was NaCl greater than HCl greater than sucrose greater than quinine hydrochloride (QHCl). When the nasoincisor ducts were tested, however, the order of stimulus effectiveness was strikingly different: sucrose was the best stimulus, followed by HCl, NaCl, and QHCl. If both the anterior tongue and nasoincisor ducts are included, stimulation of taste receptors in the anterior oral cavity of the rat produces good responses to stimuli representing 3 of the 4 classical taste qualities: sweet, salty, and sour.

Gustatory and tactile stimulation of the posterior tongue activate overlapping but distinctive regions within the nucleus of the solitary tract.

Both the gustatory and somatosensory systems provide necessary sensory input for the initiation and control of oromotor behaviors. Behavioral studies indicate that somatosensory input from the posterior tongue (PT) is important in initiating swallowing, whereas PT taste input is particularly important in gustatory rejection reflexes. However, there have been few studies of the central representation of PT gustatory or tactile responses. In the present study, electrophysiological multi-unit recording techniques were used to map the location of PT-mediated taste and tactile responses in the nucleus of the solitary tract (NST) of the rat. A stimulation technique that allows taste stimuli to be introduced directly and specifically into the papillae trenches was used to optimally activate PT taste receptors located within the circumvallate (CV) and foliate (FOL) papillae. The results demonstrated that non-PT responsive sites dominated the rostral half of the rostral division of NST (rNST), while PT-responsive sites dominated the caudal half. Some PT-responsive sites extended into the caudal NST. Both gustatory and tactile stimuli were effective at 28% of PT-responsive locations (taste-tactile sites), whereas at the remaining locations, only tactile stimulation was effective (tactile-only sites). Although these two types of PT-responsive sites exhibited some anatomical overlap, their distributions were distinctive, with taste-tactile sites restricted medially and the laterally located tactile-only sites offset caudally. On the other hand, responses arising from stimulation of the CV and FOL exhibited no anatomical organization, i.e., responses to stimulation of both papillae were coexistensive. On average, of the four tastants used (0.01 M Na saccharin, 0.3 M NaCl, 0.01 M quinine hydrochloride, 0.03 M HCl), HCl was the most effective stimulus for both the CV and FOL. The present results delimit the regions of the NST that provide a substrate for the gustatory and somatosensory limbs of PT-mediated oromotor reflexes.

Welding arc maculopathy and fluphenazine.

A 45-year-old male patient presented with a bilateral maculopathy following unprotected exposure of less than two minutes' duration to a manual metal arc welding unit. He had been receiving the drug fluphenazine for the previous 10 years for treatment of depression. We believe that the drug fluphenazine, which had accumulated in his retinal pigment epithelium, may have rendered him particularly susceptible to retinal photic damage.

Taste reactivity and Fos expression in GAD1-EGFP transgenic mice.

The central distribution of QHCl-elicited Fos-like immunoreactivity (FLI) suggests the location of a brain stem circuit that controls the oral rejection response. Although many species display an oral rejection response to bitter stimuli, the distribution of FLI associated with this response has been investigated only in rats. Fos data are minimal for the mouse, a species of increasing importance, due to its use in molecular and transgenic studies and taste-evoked oromotor responses are also only incompletely described in these rodents. We investigated these questions in FVB/NJ mice and a related transgenic strain (FVB-Tg(GadGFP)4507) that expresses green fluorescent protein in a subset of GAD1-containing neurons. QHCl, sucrose, or water delivered through intraoral cannulae yielded behavioral profiles that clearly differentiated QHCl from sucrose. Similar to rat, the number of neurons expressing FLI in the medial third of the solitary nucleus was elevated following QHCl compared with the other stimuli. In mice expressing green fluorescent protein, there was a pronounced distribution of GABAergic neurons in the ventral half of the solitary nucleus. Approximately 15% of solitary neurons expressing Fos were GABAergic, but this proportion did not differ according to stimulus.

Coding the sweet taste in the nucleus of the solitary tract: differential roles for anterior tongue and nasoincisor duct gustatory receptors in the rat.

1. A variety of chemicals that humans describe as sweet drive neurons in the nucleus of the solitary tract (NST) of the rat more vigorously when applied to the taste receptors associated with the nasoincisor ducts (NID) than when applied to taste receptors on the anterior tongue (AT). 2. The differential effects of sweet stimuli applied to the AT and NID also are evident in the set of across-neuron correlations produced by these stimuli. The psychophysical similarity among the sweet stimuli is better accounted for by responses to stimulation of the NID than by responses to stimulation of the AT (mean correlation between pairs of sweet stimuli = +0.70 for the NID, +0.44 for the AT). 3. Disaccharides or polysaccharides of glucose, i.e., maltose (0.3 M) and Polycose (0.1 M), are poor stimuli on the NID, evoking responses only 17.8 and 26.7% as great as the response elicited by sucrose (0.3 M), an optimal stimulus for this receptor subpopulation. This suggests that Polycose and maltose interact with receptor sites distinct from those with an affinity for sweet stimuli. Polycose and maltose also are ineffective stimuli on the AT, evoking responses only 11.8 and 4.9% as large as the response evoked by an optimal stimulus for this receptor subpopulation, a mixture of electrolytes (0.3 M NaCl, 0.03 M HCl, and 0.01 M quinine HCl). 4. The relative effectiveness of the sweet sugars in driving NST neurons (sucrose greater than fructose greater than glucose) correlates with their order of effectiveness in generating preference behavior in the rat.(ABSTRACT TRUNCATED AT 250 WORDS)

Organization of orosensory responses in the nucleus of the solitary tract of rat.

1. The receptive field and topographic organization of single orosensory neurons located throughout the rostral division of the nucleus of the solitary tract (rNST) was studied by determining their responsiveness to gustatory stimulation of the entire oral cavity and to gustatory and mechanical stimulation of restricted oral regions. The rNST contained roughly equal numbers of two distinct populations of orosensory neurons, one responsive exclusively to oral mechanical stimulation (M neurons), the other to gustatory stimulation (G neurons). Some G neurons also responded to oral somatosensory stimuli, but usually less vigorously than to gustatory stimuli. The distribution of these two populations of rNST neurons was topographically organized: G neurons were centered anteriorly and medially to M neurons. 2. Eight of 44 G neurons responded only when the whole oral cavity was stimulated, but the remaining 36 cells responded to circumscribed stimulation of taste buds on the anterior tongue (AT), foliate papillae of the posterior tongue, nasoincisor ducts, retromolar mucosa (RM), or soft palate (SP). Overall, AT and SP stimulation were the most effective, and RM stimulation the least effective, for activating nucleus of the solitary tract (NST) G neurons. 3. Approximately half of the G neurons for which a receptive field could be defined (N = 36) responded to stimulation of a single taste receptor subpopulation, but the remaining neurons received convergent input from two or more taste bud groups. The receptive field configurations for convergent G neurons were orderly: convergence occurred preferentially between receptor subpopulations either within the anterior oral cavity (AO) or the posterior oral cavity (PO). An AO-PO distinction also was reflected in the topographic organization of gustatory responses. The mean location of neurons responding optimally to AO gustatory stimulation was more anterior in the NST, and also tended to be more lateral and ventral than the location of neurons that responded optimally to PO stimulation. 4. Forty-four rNST M neurons responded to innocuous mechanical stimulation of restricted areas of the tongue, palate, buccal mucosa, or periodontium. Stimulation of the hard palate and circumvallate papilla were most effective, whereas periodontal stimulation was least effective for activating these cells. 5. A majority (32 of 44) of rNST M neurons responded to stimulation of more than one of the oral sites tested.(ABSTRACT TRUNCATED AT 400 WORDS)

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.

Taste bud distribution in the rat pharynx and larynx.

The present study evaluated the number and distribution of taste buds in the rat pharynx and larynx using light microscopic techniques. The average number of taste buds in this region was 141.9 (S.D. = 29.4, n = 10). Pharyngeal and laryngeal taste buds could be grouped into three subpopulations: those associated with laryngeal structures (LA), the nasopharynx (NP), or the palatopharyngeal eminence (PE). Laryngeal taste buds were most numerous (mean = 103.2, S.D. = 23.6). They were observed on the laryngeal surface of the epiglottis and extended caudally along the aryepiglottic folds, reaching peak density at the caudal extreme of the folds. Further caudally, when the larynx and rostral esophagus separated, taste bud density decreased sharply. Fewer taste buds were associated with the NP (mean = 31.9, S.D. = 13.7) or PE (mean = 6.8, S.D. = 4.0) and taste bud density in these subpopulations peaked at the anterior border of the nasopharyngeal hiatus. Taste buds in the rat larynx and pharynx account for 10% all taste buds in this species, a proportion similar to that accounted for by taste buds on the soft palate. Taste buds in this region appear to be ideally situated for protecting the airway during accidental aspiration of food or fluids. Most pharyngeal and laryngeal taste buds are unlikely to be involved in triggering normally occurring swallows, although they could be stimulated as the bolus is propelled from the oral cavity to the esophagus.

Altered taste responses in adult NST after neonatal chorda tympani denervation.

Anatomic and behavioral changes have been observed in the taste system after peripheral deafferentation, but their physiological consequences remain unknown. Interestingly, a recent behavioral study suggested that peripheral denervation could induce central plasticity. After neonatal chorda tympani (CT) transection, adult rats demonstrated a marked preference for a normally avoided salt, NH(4)Cl. In the present study, taste responses were recorded from the nucleus of the solitary tract (NST) in similarly CT-denervated rats to investigate a physiological basis for this behavioral phenomenon. We hypothesized that alterations in functional connectivity of remaining afferent nerves might underlie the behavioral change. Specifically, if NST neurons formerly activated by sodium-selective CT fibers were instead driven by more broadly tuned glossopharyngeal (GL) afferents, neural coding of salt responses would be altered. Such a change should be accompanied by a shift in orotopic representation and increased NH(4)Cl responses. This hypothesis was not supported. After CT denervation, orotopy was unaltered, NH(4)Cl responsiveness declined, and no other changes occurred that could simply explain the behavioral effects. Indeed, the most pronounced consequence of CT denervation was a 68% reduction in NaCl responses, supporting previous evidence for a critical role of this nerve in coding sodium salts. In addition, we found "reorganizational" changes similar to, albeit smaller than, those observed in other sensory systems after deafferentation. There was a trend for increased responses elicited by stimulation of receptor subpopulations innervated by the GL and greater superficial petrosal nerves. In addition, the spontaneous rate of nasoincisor duct-responsive cells increased significantly. This effect on spontaneous rate is opposite to that produced by CT anesthesia, suggesting that acute versus chronic denervation may affect central taste neurons differently. In conclusion, the taste system at the medullary level seems more resistant to large-scale plasticity than other sensory systems, but nevertheless reacts to lost afferent input. Because the most robust plastic changes have been documented at cortical levels in other sensory pathways, the substrate for the behavioral effect of neonatal CT transection may be located more centrally in the gustatory system.

Anterior and posterior oral cavity responsive neurons are differentially distributed among parabrachial subnuclei in rat.

The responses of single parabrachial nucleus (PBN) neurons were recorded extracellularly to characterize their sensitivity to stimulation of individual gustatory receptor subpopulations (G neurons, n = 75) or mechanical stimulation of defined oral regions (M neurons, n = 54) then localized to morphologically defined PBN subdivisions. Convergence from separate oral regions onto single neurons occurred frequently for both G and M neurons, but converging influences were more potent when they arose from nearby locations confined to the anterior (AO) or posterior oral cavity (PO). A greater number of G neurons responded optimally to stimulation of AO than to PO receptor subpopulations, and these AO-best G neurons had higher spontaneous and evoked response rates but were less likely to receive convergent input than PO-best G neurons. In contrast, proportions, response rates, and convergence patterns of AO- and PO-best M neurons were more comparable. The differential sensitivity of taste receptor subpopulations was reflected in PBN responses. AO stimulation with NaCl elicited larger responses than PO stimulation; the converse was true for QHCl stimulation. Within the AO, NaCl elicited a larger response when applied to the anterior tongue than to the nasoincisor duct. Hierarchical cluster analysis of chemosensitive response profiles suggested two groups of PBN G neurons. One group was composed of neurons optimally responsive to NaCl (N cluster); the other to HCl (H cluster). Most N- and H-cluster neurons were AO-best. Although they were more heterogenous, all but one of the remaining G neurons were unique in responding best or second-best to quinine and so were designated as quinine sensitive (Q+). Twice as many Q+ neurons were PO- compared with AO-best. M neurons were scattered across PBN subdivisions, but G neurons were concentrated in two pairs of subdivisions. The central medial and ventral lateral subdivisions contained both G and M neurons but were dominated by AO-best N-cluster G neurons. The distribution of G neurons in these subdivisions appeared similar to distributions in most previous studies of PBN gustatory neurons. In contrast to earlier studies, however, the external medial and external lateral-inner subdivisions also contained G neurons, intermingled with a comparable population of M neurons. Unlike cells in the central medial and ventral lateral subnuclei, nearly every neuron in the external subnuclei was PO best, and only one was an N-cluster cell. In conclusion, the present study supports a functional distinction between sensory input from the AO and PO at the pontine level, which may represent an organizing principle throughout the gustatory neuraxis. Furthermore, two morphologically distinct pontine regions containing orosensory neurons are described.