Taste Pathways Research Articles

Neural circuits for taste. Excitation, inhibition, and synaptic plasticity in the rostral gustatory zone of the nucleus of the solitary tract.

The rostral nucleus of the solitary tract (rNST) plays a key role in modulating, organizing and distributing the sensory information arriving at the central nervous system from gustatory receptors. However, except for some anatomical studies of rNST synapses, the neural circuits responsible for this first stage in synaptic processing of taste information are largely unknown. Over the past few years we have used an in vitro brain slice preparation of the rNST to study synaptic processing, and it has become apparent that the rNST is a very complex neural relay. Synaptic potentials recorded in rNST neurons resulting from stimulation of afferent taste fibers are a composite of excitatory and inhibitory post synaptic potentials. Pure excitatory postsynaptic potentials (EPSP) can be isolated by using gamma-aminobutyric acid type A (GABAA) receptor blockers to eliminate the inhibitory postsynaptic potentials (IPSP). Application of glutamate ionotropic receptor blockers effectively eliminates all postsynaptic activity, indicating that glutamate is the transmitter at the first central synapse in the taste pathway. Stimulation of the afferent taste fibers originating from the anterior (chorda tympani) and posterior (glossopharyngeal) tongue results in a postsynaptic potential that is a complex sum of the two individual potentials. Thus, rNST neurons receive convergent synaptic input from the anterior and posterior tongue. The IPSP component of the synaptic potentials in rNST results from stimulation of interneurons. If these IPSPs are initiated by tetanic stimulation they undergo both short-term and long-term changes. Short-term changes result in the development of biphasic depolarizing IPSPs, and long-term changes result in potentiation of the IPSPs that can last over an hr in some neurons. This remarkable synaptic plasticity may be involved in the mechanism of learned taste behaviors. Synaptic transmission in rNST consists of excitation combined with inhibition. The inhibition does not simply depress excitation but probably serves many roles such as shaping and limiting excitation, coordinating the timing of synaptic events and participating in synaptic plasticity. Knowledge of these synaptic mechanisms is essential to understanding how the rNST processes taste information.

Neurochemical Changes after Imbalanced Diets Suggest a Brain Circuit Mediating Anorectic Responses to Amino Acid Deficiency in Rats , ,

Amino acid–imbalanced (IMB) diets induce an acute amino acid deficiency and hypophagic responses in most animals. The neural circuits underlying these responses are unknown. To ascertain potential neural circuits involved in the recognition of IMB, we measured the concentrations of norepinephrine, dopamine, serotonin, their metabolites and 20 amino acids in 14 rat brain areas in three studies. Rats were prefed a basal diet with L-amino acids as the protein source for at least 1 wk. For the experiments, either threonine or isoleucine IMB diet was offered for 2.5 or 3.5 h. Brains were taken before (using a mildly IMB diet) or after (using moderately or severely IMB diet) food intake was significantly (P < 0.05) depressed. Brain areas were dissected and analyzed for monoamines, metabolites and amino acids. Only in the anterior piriform cortex (APC), a brain area that may contain the amino acid chemosensor, was the limiting amino acid lower in IMB groups than in controls across all of the experiments. Before the onset of the anorectic response to the IMB diets, monoaminergic activity was affected in areas that have recognized monosynaptic connections with the APC. We propose a circuit for the neural responses in the initial recognition of acute amino acid deprivation that begins with activation of the APC and includes areas in the hindbrain and hypothalamus. After a significant hypophagic response, serotonergic indicators were altered in areas of the taste pathway and the limbic system. These results suggest that different circuits mediate the initial recognition and secondary conditioned responses to IMB diets.

Synaptic interactions due to convergent input from gustatory afferent fibers in the rostral nucleus of the solitary tract.

1. Whole cell recordings from neurons in the rostral, gustatory nucleus of the solitary tract (rNST) were made using the "blind" patch-clamp technique in horizontal brain stem slices of rats. 2. Postsynaptic potentials (PSP) were elicited in 71 rNST neurons by electrical stimulation of the solitary tract (ST). To investigate PSPs evoked by convergent input from the chorda tympani and glossopharyngeal nerves, the ST was stimulated at levels where these two nerves terminate. These are referred to as rostral (rST) and intermediate (iST) ST, respectively. 3. When the rST was stimulated 72% of the PSPs were depolarizing, and 28% were hyperpolarizing (n = 64). Stimulation of the intermediate ST resulted in 75% depolarizing and 25% hyperpolarizing PSPs (n = 56). 4. Application of gamma-aminobuturic acid-A (GABAA) and glutamate receptor blockers revealed that all PSPs recorded in the present study were a composite of summed excitatory and inhibitory PSPs. Application of the GABAA receptor blocker bicuculline, by eliminating the hyperpolarizing component of a PSP, revealed the excitatory postsynaptic potential (EPSP) component of the potential. Bicuculline also increased the amplitude and prolonged the decay time of the depolarizing potentials once the hyperpolarizing potential component had been eliminated. These pure EPSP revealed by GABAA receptor blockade reversed at approximately 0 mV. 5. Application of glutamate ionotropic receptor blockers effectively eliminated the initiation of the synaptic responses evoked by ST stimulation. If the stimulus strength was increased, an inhibitory postsynaptic potential (IPSP) was elicited, presumably by direct activation of interneurons close to the stimulating electrode. These IPSPs had a mean reversal potential of -88 mV. 6. When synaptic responses were initiated by stimulation of the projection areas of both the chorda tympani and glossopharyngeal nerves, all neurons tested (n = 49) responded to stimulation of both sites on the ST. The resulting synaptic potential was a sum of the two individual synaptic potentials. 7. If stimulation of the rostral and intermediate sites both elicited depolarizing potentials, the potential resulting from stimulation of both sites was the arithmetical sum of the two individual PSPs. The EPSPs summed even if the time between stimulation of the rostral and intermediate sites was separated by < or = 100 ms. 8. Inhibitory PSPs evoked by simultaneous stimulation of the rostral and intermediate ST also summed. The summation was not linear and saturated at a mean level of -66 mV. 9. When the PSP at one stimulation site was excitatory but inhibitory at the other site, the PSP wave form resulting from dual stimulation was a complex mixture of the two individual potentials. The inhibitory potential was capable of blocking action potentials resulting from the excitatory PSP. 10. These results indicate that synaptic responses in rNST are complex mixtures of excitatory and inhibitory potentials. The synaptic potentials result from excitatory afferent input mediated by glutamate and inhibitory input derived from interneurons. Inhibitory transmission is mediated primarily at GABAA synapses. Stimulation of the rNST afferent input derived from the chorda tympani and glossopharyngeal nerves reveals complex convergent input. The complexity of these synaptic interactions indicates that considerable processing of gustatory information occurs at the first central synapse in the taste pathway.

Introduction: Neurotransmitters and neuromodulators in gustation

A complex chain of neural events is initiated when a chemical is placed on the tongue that eventually leads to a taste sensation. Beginning with transduction and a receptor potential, the sensory information has to be transformed into action potentials that ascend from the periphery through the central nervous system. This process involves many synaptic junctions where the message is transformed and distributed to a number of different brain areas. While transduction has been a major focus of recent investigations in taste, synaptic processing, and the neurotransmitters and neuromodulators used by the taste system have received less attention. Obviously, an understanding of the role of neurotransmitters and neuromodulators in the gustatory system is very important if progress is to be made in comprehending how taste information is processed by the central nervous system. Thus, it is timely to examine the current knowledge of the role of neurotransmitters and neuromodulators in gustation. A symposium at the Association for Chemoreception Sciences Annual Meeting (AChemS XVII) recently focused on this topic to address the role of neurotransmitters and neuromodulators at the level of the taste bud and at the first central relay in the taste pathway—the nucleus of the solitary tract (NST). The first report of this symposium is a comprehensive review by Steve Roper of the neurotransmitters and neuromodulators at the level of the taste receptors. Once transduction has taken place at the taste cell receptor membrane the information has to pass through a synapse between the taste cells and the primary afferent taste fibers. Suprisingly little is known about this essential event or about the identity of the neurotransmirter/s involved. As detailed in this report several investigators have attempted to study synaptic transmission at the taste bud. Histochemical, immunocytochemical and physiological techniques have been used in a number of different species with many conflicting results. Often an investigator produces a flurry of papers and then apparently abandons the effort. A few years later another investigator tries again, probably because new methods become available, to re-investigate the question of synaptic transmission at the level of the taste bud. Roper describes recent work from his laboratory indicating the significant role played by serotonin as a neuromodulator in the taste bud, but also reports that the identity of the neurotransmitter released at the taste cellprimary afferent fiber synapse is still unknown. In the last three reports information is presented on neurotransmiters and neuromodulators at the level of the NST. Synaptic events at the first relay in the central taste pathway are also little understood, as are the neurotransmiters

Neural mechanisms of taste aversion learning

When ingestion of a taste stimulus is paired with internal malaise the animal remembers the taste and rejects its ingestion thereafter. A model of neural substrate for this conditioned taste aversion (CTA) is presented on the basis of our recent experiments. To establish the CTA in rats, 0.01 M saccharin was paired with i.p. injection of 0.15 M LiCl. Behavioral lesion experiments by means of ibotenic acid showed that the parabrachial nucleus (PBN), medial thalamus (MT), and lateral part of the amygdala (LPA) were crucial to establish the CTA. c- fos immunoreactivity studies showed that ingestion of saccharin induced a remarkable activation of the central lateral (cl) subnucleus of the PBN in normal rats, however, rats with the CTA to saccharin showed c- fos neurons in the ventral lateral (vl) subnucleus of the PBN. These findings suggest that the recipient zone for saccharin taste switches from the cl to vl subnuclei after acquisition of a CTA. The taste pathway from the vl subnucleus to the LPA through the MT may be a neural substrate for taste aversion learning. The switching may result from a plastic change involving long-term potentiation and depression due to a convergence of the taste input of saccharin and general visceral information of LiCl injection.

The sense of taste: neurobiology, aging, and medication effects.

The sense of taste is an oral chemical sense in mammals that is involved in the choice of foods. Initial transduction of taste stimuli occurs in taste buds, which are distributed in four discrete fields in the oral cavity. Medications can affect the taste buds and ion channels in taste-bud cell membranes involved in stimulus transduction. The sense of taste gradually declines with aging, with bitter taste most affected. Neural circuits that mediate taste in primates include cranial nerves VII, IX, and X, the solitary nucleus in the brain stem, the ventroposteromedial nucleus of the thalamus, and the insular-opercular cortex. The central taste pathways process taste information about sweet, salty, sour, and bitter stimuli serially and in parallel. Medications associated with "metallic" dysgeusia and taste losses affect the taste system via unknown mechanisms.

Intramedullary connections of the rostral nucleus of the solitary tract in the hamster

The rostral nucleus of the solitary tract (NST) figures prominently in the gustatory system, giving rise to ascending taste pathways that are well documented. Less is known of the local connections of the rostral NST with sites in the medulla. This study defines the intramedullary connections of the rostral NST in the hamster. Small iontophoretic injections of horseradish peroxidase (HRP), confined to the rostral NST, resulted in Golgi-like filling of axons that exited the NST or that interconnected cytoarchitectonic subdivisions within the NST complex. The NST efferent axons terminated sparsely in the trigeminal, facial and hypoglossal motor nuclei, but axons and endings were heavily distributed in the parvicellular reticular formation ventral to the NST. HRP injections centered in this part of the reticular formation resulted in heavy projections to the orofacial motor nuclei. Intranuclear connections, labelled after NST injections, linked NST subdivisions that receive primary afferent taste inputs to suvdivisions involved in (1) projections to the preoromotor reticular formation, (2) projections to swallowing motor neurons, (3) activation of preganglionic parasympathetic neurons, and (4) general viscerosensation. In general, the connections defined in the present study provide anatomical details about the substrate for gustatory-motor and gustatory-visceral interactions.

The Shaker and shaking-B genes specify elements in the processing of gustatory information in Drosophila melanogaster

Mutations that affect the physiological properties of neurones or the development of neuronal circuits are likely to have profound effects on sensory and motor pathways. We have examined the effects of mutations in two loci--the Shaker complex and shaking-B-on the taste pathway of Drosophila melanogaster Meigen at the behavioural and electrophysiological levels. The Shaker locus encodes a variety of A-type potassium channels that are likely to be tissue- and stage-specific. Flies containing the different Shaker alleles examined exhibit a variety of defects in their gustatory responses to sucrose, NaCl and KCl. The firing patterns of the labellar chemosensory neurones in response to these stimuli are normal. This suggests that the channels encoded by the Shaker locus are probably not involved in taste transduction, but affect central gustatory circuits. The shaking-B locus affects neuronal connectivity, though its molecular nature is unknown. Mutants at this locus show increased thresholds for detection of sucrose and fructose and lack the attraction response to 0.1 mol l-1 sodium chloride that is exhibited by the wild-type fly.

The relative advantages of sparse versus distributed encoding for associative neuronal networks in the brain

In some neuronal networks in the brain which are thought to operate as associative memories, a sparse coding of information can enhance the storage capacity. The extent to which this statement is valid in general is discussed here, by considering some simple formal models of associative memory which include different neurobiological constraints. In nets of linear neurons, trained with either a Hebbian (purely incremental) or a Stanton and Sejnowski learning rule, sparse coding increases the number of independent associations that can be stored. When neurons are nonlinear, for a diversity of learning rules, sparse coding may result in an increase in the number of patterns that can be discriminated. The analysis is then used to help interpret recent evidence on the encoding of information in the taste and visual systems, as obtained from recordings in primates. Following the taste pathway, it is found that the breadth of tuning of individual neurons becomes progressively finer, consistent with the idea that sparser representations become advantageous as the taste information is eventually associated with that coming from other sensory modalities. In the visual system, considering a population of neurons in the temporal cortex that respond preferentially to faces, it is argued that their breadth of tuning represents a compromise between fully distributed encoding, and a grandmother cell type of encoding, which would result in a given neuron responding only to an individual face.

Convergence in mammalian nucleus of solitary tract during development and functional differentiation of salt taste circuits

To determine the type and extent of neural rearrangements that are made during functional differentiation of circuits for salt taste processing, we determined receptive field size and salt response characteristics of second-order taste cells in 3 age groups of sheep. Neurophysiological recordings were made from single cells in the nucleus of the solitary tract (NST) in fetal, perinatal, and postnatal sheep. Responses to NH4Cl, NaCl, and KCl were measured, and location and number of fungiform papillae in the receptive field were determined by stimulating individual papillae with anodal electrical current. The data are compared with previous, parallel measures from chorda tympani nerve afferent taste fibers to permit conclusions about convergence or divergence onto second-order cells. Receptive field size of second-order taste neurons increases during development, in contrast to the decrease in field size observed previously for chorda tympani nerve fibers during the same period. Furthermore, receptive fields of second-order cells are significantly larger than those of first-order fibers at perinatal and lamb ages, but not fetal. Thus, there is convergence of first-order taste afferents onto brain-stem neurons, and the convergence increases remarkably between fetal and perinatal periods. Associated with the increase in convergence are increased salt response frequencies relative to afferent fibers for NaCl in perinatal animals and lambs, and for KCl in lambs. The increase in frequencies occurs before NST neurons are functionally mature, as indicated by the rapid response adaptation of many cells in young animals. Convergence in NST during development apparently functions to maximize gain for processing neural responses to NaCl. In the periphery, response frequencies to NaCl are very low in fetuses, and increase progressively during development. In the NST, NaCl response frequencies are high even in fetuses, and remain high. The process of convergence onto second-order cells is accomplished with maintenance of order in afferent projections because receptive fields of NST neurons are composed of fungiform papillae that are clustered together, not dispersed over the tongue. Our quantification of taste receptive field size at 2 neural levels provides strong evidence for increasing convergence in the NST during development. Altering patterns of afferent neural input and geometry of second-order neurons may have a role in establishing convergence. The convergence has an apparently special function: to increase gain for NaCl taste sensation. Therefore, neural rearrangements during differentiation of salt taste pathways result in specific functional outcomes.

The relative advantages of sparse versus distributed encoding for associative neuronal networks in the brain

In some neuronal networks in the brain which are thought to operate as associative memories, a sparse coding of information can enhance the storage capacity. The extent to which this statement is valid in general is discussed here, by considering some simple formal models of associative memory which include different neurobiological constraints. In nets of linear neurons, trained with either a Hebbian (purely incremental) or a Stanton and Sejnowski learning rule, sparse coding increases the number of independent associations that can be stored. When neurons are nonlinear, for a diversity of learning rules, sparse coding may result in an increase in the number of patterns that can be discriminated. The analysis is then used to help interpret recent evidence on the encoding of information in the taste and visual systems, as obtained from recordings in primates. Following the taste pathway, it is found that the breadth of tuning of individual neurons becomes progressively finer, consistent with the idea that...

Calbindin D-28k and parvalbumin in the rat nervous system

This paper describes the distribution of structures stained with mono- and polyclonal antibodies to the calcium-binding proteins calbindin D-28k and parvalbumin in the nervous system of adult rats. As a general characterization it can be stated that calbindin antibodies mainly label cells with thin, unmyelinated axons projecting in a diffuse manner. On the other hand, parvalbumin mostly occurs in cells with thick, myelinated axons and restricted, focused projection fields. The distinctive staining with antibodies against these two proteins can be observed throughout the nervous system. Calbindin D-28k is primarily associated with long-axon neurons (Golgi type I cells) exemplified by thalamic projection neurons, strionigral neurons, nucleus basalis Meynert neurons, cerebellar Purkinje cells, large spinal-, retinal-, cochlear- and vestibular ganglion cells. Calbindin D-28k occurs in all major pathways of the limbic system with the exception of the fornix. Calbindin D-28k is, however, also found in some short-axon cells (Golgi type II), represented by spinal cord interneurons in layer II and interneurons of the cerebral cortex. It is also detectable in some ependymal cells and abundantly occurs in vegatative centres of the hypothalamus. The “paracrine core” of the nervous system and its adjunct (1985, Nieuwenhuys, Ghemoarchitecture of the Brain. Springer, Berlin) is very rich in calbindin D-28k. The distribution of calbindin D-28k-positive neurons is very similar to that of the dihydroperydine subtype of calcium channels. Most of the cells containing calbindin D-28k are vulnerable to neurodegenerative processes. Parvalbumin-immunoreactive neurons have a different, and mostly complementary distribution compared with those which react with calbindin D-28k antisera, but in a few cases (Purkinje cells of the cerebellum, spinal ganglion neurons), both calcium-binding proteins co-exist in the same neuron. Many parvalbumin-immunoreactive cells in the central nervous system are interneurons (Golgi type II) and, to a lesser extent, long-axon cells (Golgi type I), whereas conditions are vice versa in the peripheral nervous system. Intrinsic parvalbuminic neurons are prominent in the cerebral cortex, hippocampus, cerebellar cortex and spinal cord. Long-axon parvalbumin-immunoreactive neurons are, for example, the Purkinje cells, neurons of the thalamic reticular nucleus, globus pallidus, substantia nigra (pars reticulata) and a subpopulation among large spinal-, retinal-, cochlear- and vestibular ganglion cells. Parvalbumin is rich in cranial nerve nuclei related to eye movements. In addition to nervous elements, parvalbumin immunoreactivity occurs in a few ependymal cells and in some pillar cells of the organ of Corti. In contrast to calbindin D-28k, parvalbumin is virtually absent from vegetative centres and pathways. Parvalbumin co-exists with GABA in cortical neurons and is, in general, preferentially associated with “fast-firing” neurons. Whole functional systems are revealed by either parvalbumin or calbindin D-28k immunohistochemistry. Parvalbumin, for example, occurs in the whole chain of neurons of the epicritic sensibility in the somatosensory system. Calbindin D-28k, on the other hand, occurs in the whole taste pathway. With antibodies against calbindin D-28k and parvalbumin it is possible to reveal at least two as yet undescribed brain nuclei: one in the hypothalamus (parvalbumin 1), the second in the medulla (calbindin D-28k 1). Calbindin D-28k and parvalbumin are evenly distributed within the various domains of the same neuron, but in some instances their levels vary between soma and cell processes. The immunostaining with calbindin D-28k and parvalbumin antisera show gradations in intensity among neurons belonging to different populations, a phenomenon compatible with the presence of varying concentrations of these proteins. Calbindin D-28k and parvalbumin are excellent new neuroanatomical markers which can be utilized to selectively visualize certain neurons and pathways in the central nervous system and peripheral nervous system.

Neural substrates for reflex salivation induced by taste, mechanical, and thermal stimulation of the oral region in decerebrate rats.

In order to investigate the neural mechanisms of reflex salivary secretion, experiments were carried out on anesthetized, decerebrate rats from which the volumes of submandibular salivary secretion and the efferent discharges in the preganglionic parasymapathetic fibers innervating the submandibular gland were recorded. Salivary secretion was induced by either infusing a taste solution, or an aliquot of hot water (45-55 degrees C) into the oral cavity, or by pinching the frontal parts of the oral region with a pair of forceps. The reflex salivation induced by noxious thermal and mechanical stimuli was markedly reduced by lesioning either the caudal (VC), or the interpolar (VI) trigeminal sensory nuclei. Taste-elicited salivary secretion was significantly reduced by lesioning the nucleus of the tractus solitarius (NTS). Of 43 preganglionic parasympathetic fibers sampled, 27 responded to both noxious mechanical and thermal stimulation of the oral region, and to electrical stimulation of the VC. Ten fibers responded only to taste stimulation and to electrical stimulation of the NTS. The remaining 6 fibers responded to both taste and noxious thermal stimulation of the oral region. These fibers responded well to NTS stimulation, but gave only a slight response to VC stimulation. These results suggest that two distinct neural pathways exist which mediate reflex salivation in the lower brain stem of the rat, i.e., the taste pathway via the NTS and the nociceptive pathway via the trigeminal sensory nuclei.

Structure/activity relationships in the arginine taste pathway of the channel catfish

To characterize the molecular/structural requirements for activation or antagonism of the arginine taste pathways in catfish, Ictalurus punctatus, structure/activity studies were performed using integrated multi-unit responses and cross-adaptation. Of all the guanidinium-containing compounds tested, only L-arginine, L-α-amino-β-guanidino propionic acid (L-AGPA) and L-arginine methyl and ethyl esters were strong stimuli

Information processing in the taste pathways and its behavioural significance

Information processing in the taste pathways and its behavioural significance

Evoked taste thresholds in a normal population and the application of electrogustometry to trigeminal nerve disease.

No standardised method for taste threshold measurement is available and therefore comparison between clinical studies is difficult. An electrogustometer was evaluated in normal subjects. No sex difference in taste threshold was noted; however, there was a significant elevation in detection threshold with age and smoking. Electrogustometric values both in patients before and after surgery for trigeminal neuralgia and in patients with trigeminal sensory neuropathy were determined. Many patients with trigeminal nerve disorders had abnormal electrogustometric detection thresholds suggesting that there is possibly an accessory taste pathway through the trigeminal nerve, although in some individuals the site of lesion may be in the brain stem. Electrogustometry is a convenient method for clinically assessing taste.

Selective localization of calcium-binding protein in human brainstem, cerebellum and spinal cord

The 28,000-Da vitamin D-dependent calcium-binding protein, CaBP, which is induced by one hormonally active form of vitamin D 3, 1,25-dihydroxyvitamin D 3, was localized by immunocytochemistry in the human brainstem, cerebellum and cervical segment of the spinal cord. Positive structures (neurons and their processes) were restricted to some well-defined motor and sensory pathways. In motor regions, the highest density of immunoreactive sites was found in the Purkinje cell layer of the cerebellar cortex, and CaBP-positive neurons were also found in the reticular formation and the inferior olivary nucleus. In sensory pathways, positive neurons were mainly localized in structures associated with protopathic thermalgesia (pain and temperature), as well as in the solitary nucleus and parabrachial nucleus of the taste pathway.

Lateral hypothalamic sites eliciting eating affect medullary taste neurons in rats

A hypothetical functional relationship between the feeding area of the lateral hypothalamus and the medullary and pontine taste areas was assessed in rats. Multiple electrodes were implanted in the lateral hypothalamus and the animals which exhibited eating upon electrical stimulation were selected. Then under pentobarbital anesthesia, the hypothalamus was electrically stimulated while unit activity of medullary and pontine taste areas was recorded. Electrical stimulation of the hypothalamus at sites which elicited eating affected 22% of gustatory neurons, 13% of proprioceptive neurons sensitive to oropharyngeal stimuli but only 0.2% of the remaining nongustatory neurons. Electrical stimulation at nonfeeding sites affected 3% of the gustatory neurons. The electrical stimulation of the hypothalamus had the same inhibitory or excitatory effect on a given gustatory neuron as the sapid stimulation of the tongue did. These results are discussed in terms of the taste pathways and their possible role in electrically elicited feeding.

Neuropeptides are present in projection neurones at all levels in visceral and taste pathways: from periphery to sensory cortex

Using combined immuno-staining and retrograde tracing techniques many of the ascending visceral and taste pathways within the rat central nervous system have been shown to be composed of a variety of neuropeptide and catecholamine synthesizing enzyme containing neurones. The pathway we examined extended from the periphery to sensory cortex and included: the nodose ganglion (periphery) →solitary nucleus (medulla)→parabrachial nucleus (pons)→ventral posterior medial nucleus (thalamus)→visceral and taste sensory areas (cortex). In the solitary nucleus of the medulla many neuronal cell bodies could be shown to be both immuno-positive for one of 6 neuropeptides including avian pancreatic peptide (APP), cholecystokinin (CCK), enkephalin (ENK), neurotensin (NT), somatostatin (SOM) and substance P (SP) or the catecholamine synthesizing enzyme tyrosine hydroxylase (TOH) and to have a projection to the parabrachial nucleus of the pons. In the parabrachial nucleus of the pons many neuronal cell bodies could be shown to be immuno-positive for one of 5 neuropeptides (CCK, ENK, NT, SOM, SP) and have a projection to the ventral posterior medial nucleus of the thalamus. In the ventral posterior medial nucleus of the thalamus several neuronal cell bodies were shown to be immuno-positive for one of 3 neuropeptides (CCK, ENK, SOM) and project to the visceral and taste sensory cortex. This is the first report of neuropeptides being present in the projection neurones of any sensory system in the central nervous system and for the first time describes an entire set of putative neurotransmitters which extends from the periphery to the sensory cortex. From previous studies it also appears that in all cases examined the relevant receptors are present in these visceral and taste relay nuclei in order for the neuropeptide or catecholamine to produce an effect upon release. Comparisons between rat and other animals suggest that a similar organization of these visceral and taste pathways may also be present in other mammals including man. Functionally these neuropeptides containing projection neurones appear to be primarily involved in relaying visceral information rather than taste information. In this capacity activation of these neurones may produce such visceral sensations as malaise, well being, hunger, satiety or thirst.

Taste pathways to hypothalamus and amygdala.

The projections of a third order gustatory relay in the dorsal pons of rats have been traced using tritiated proline autoradiography and antidromic activation of pontine neurons from electrodes in the thalamus and amygdala. Labelled axons collect in the central tegmental tract and ascend to the thalamic taste area in the medial extension of the ventrobasal complex. The majority of the fibers remain ipsilateral, but a few cross in the rostral pons and midbrain. The largest crossing occurs at the level of the thalamic termination. Many fascicles of fibers continue rostrally by passing beneath the thalamic taste area, piercing the medial lemniscus, and spreading out along the dorsomedial corner of the internal capsule (IC). The terminal field at this level caps IC from the subthalamic nucleus down into the far-lateral hypothalamus. Labelled axons grandually penetrate through the internal capsule, and ramify throughout the underlying substantia innominata. This terminal zone extends laterally into the rostral end of the central nucleus of the amygdala, which is densely labelled to its caudal exremity. At the caudal end of the amygdala labelled fibers are visible in one component of the stria terminalis. These fibers can be followed over the dorsal thalamus into a smaller, but equally dense terminal area in the dorsolateral bed nucleus of the stria terminalis. The electrophysiological data demonstrate that pontine gustatory units can be antidromically activated by electrodes located in or near the central nucleus of the amygdala. Since many of the same units can also be driven from the thalamic taste area, at least some of the axons traced autoradiographically probably convey gustatory information to the hypothalamus and amygdala.