Roles of chemical mediators in the taste system

Recent advances in neural mechanisms of taste are reviewed with special reference to neuroactive substances. In the first section, taste transduction mechanisms of basic tastes are explained in two groups, whether taste stimuli directly activate ion channels in the taste cell membrane or they bind to cell surface receptors coupled to intracellular signaling pathways. In the second section, putative transmitters and modulators from taste cells to afferent nerves are summarized. The candidates include acetylcholine, catecholamines, serotonin, amino acids and peptides. Studies favor serotonin as a possible neuromodulator in the taste bud. In the third section, the role of neuroactive substances in the central gustatory pathways is introduced. Excitatory and inhibitory amino acids (e.g., glutamate and GABA) and peptides (substance P and calcitonin gene-related peptide) are proved to play roles in transmission of taste information in both the brainstem relay and cortical gustatory area. In the fourth section, conditioned taste aversion is introduced as a model to study gustatory learning and memory. Pharmacobehavioral studies to examine the effects of glutamate receptor antagonists and protein kinase C inhibitors on the formation of conditioned taste aversion show that both glutamate and protein kinase C in the amygdala and cortical gustatory area play essential roles in taste aversion learning. Recent molecular and genetic approaches to disclose biological mechanisms of gustatory learning are also introduced. In the last section, behavioral and pharmacological approaches to elucidate palatability, taste pleasure, are described. Dopamine, benzodiazepine derivatives and opioid substances may play some roles in evaluation of palatability and motivation to ingest palatable edibles.     

The afferent code for sensory quality.

"Many of the more recent findings in sensory psychology and physiology derive from the application of electrophysiology to the study of sensory processes." Experiments involving the taste sense and utilizing the recording and amplifying of nerve impulse traffic in sensory fibers "en route" to the brain are reviewed, and their general implications for the theory of afferent coding are discussed. An important principle of sensory coding is that "the same afferent fiber may convey different information depending upon the amount of activity in another parallel fiber." The relative rather than the absolute amount of activity in any one set of afferent fibers may determine the quality of sensation. "It is not only the activity in parallel fibers that is important, it is the relative amount of such parallel activity. Studies of the other senses indicate that these principles are not unique to taste." 21 refs. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Control of the phosphorylation of the astrocyte marker glial fibrillary acidic protein (GFAP) in the immature rat hippocampus by glutamate and calcium ions: possible key factor in astrocytic plasticity

The present review describes recent research on the regulation by glutamate and Ca2+ of the phosphorylation state of the intermediate filament protein of the astrocytic cytoskeleton, glial fibrillary acidic protein (GFAP), in immature hippocampal slices. The results of this research are discussed against a background of modern knowledge of the functional importance of astrocytes in the brain and of the structure and dynamic properties of intermediate filament proteins. Astrocytes are now recognized as partners with neurons in many aspects of brain function with important roles in neural plasticity. Site-specific phosphorylation of intermediate filament proteins, including GFAP, has been shown to regulate the dynamic equilibrium between the polymerized and depolymerized state of the filaments and to play a fundamental role in mitosis. Glutamate was found to increase the phosphorylation state of GFAP in hippocampal slices from rats in the post-natal age range of 12-16 days in a reaction that was dependent on external Ca2+. The lack of external Ca2+ in the absence of glutamate also increased GFAP phosphorylation to the same extent. These effects of glutamate and Ca2+ were absent in adult hippocampal slices, where the phosphorylation of GFAP was completely Ca(2+)-dependent. Studies using specific agonists of glutamate receptors showed that the glutamate response was mediated by a G protein-linked group II metabotropic glutamate receptor (mGluR). Since group II mGluRs do not act by liberating Ca2+ from internal stores, it is proposed that activation of the receptor by glutamate inhibits Ca2+ entry into the astrocytes and consequently down-regulates a Ca(2+)-dependent dephosphorylation cascade regulating the phosphorylation state of GFAP. The functional significance of these results may be related to the narrow developmental window when the glutamate response is present. In the rat brain this window corresponds to the period of massive synaptogenesis during which astrocytes are known to proliferate. Possibly, glutamate liberated from developing synapses during this period may signal an increase in the phosphorylation state of GFAP and a consequent increase in the number of mitotic astrocytes.     

Color as a factor in food choice

From birth, nature teaches us to make judgements on our environment based in large measure on color. As such, it plays a key role in food choice by influencing taste thresholds, sweetness perception, food preference, pleasantness, and acceptability. Its role is elusive and difficult to quantify, however, which at times has placed color in a secondary role to the other sensory characteristics, a position not entirely consistent with the facts. Color, in a quantitative sense, has been shown to be able to replace sugar and still maintain sweetness perception in flavored foods. It interferes with judgments of flavor intensity and identification and in so doing has been shown to dramatically influence the pleasantness and acceptability of foods. Studies in the literature have used cross-sectional population panels to study these effects, but a recent investigation of color-sensory interactions in beverages has compared the response of a college age group with the response of a panel consisting of a more mature population. Interestingly, the older group showed significant differences from the college age group in their response to the effects of color on several sensory parameters as well as showing a direct correlation between beverage consumption and color. Color is often taken for granted, but this position must be reevaluated in view of such studies and the need to create more appealing foods for different segments of our society.     

Evidence for the interaction of glutamate and NK1 receptors in the periphery

It is known that Substance P (SP) enhances glutamate- and N-methyl-D-aspartate (NMDA)-induced activity in spinal cord dorsal horn neurons and that this enhancement is important in the generation of wind-up and central sensitization. It is now known that SP and glutamate receptors are present on sensory axons in rat glabrous skin. This raises the issue as to whether SP and glutamate interact in the periphery. Using the tail skin in rats, the present study demonstrates 1) that unmyelinated axons at the dermal-epidermal junction immunostain for antibodies directed against NMDA, non-NMDA or SP (NK1) receptors; 2) that glutamate injected into the tail skin results in dose-dependent nociceptive behaviors interpreted as mechanical hyperalgesia, mechanical allodynia and thermal hyperalgesia, which are blocked following co-injection with glutamate antagonists; 3) that peripheral injection of SP potentiates glutamate-induced nociceptive behaviors in that the co-injection of SP+glutamate results in a significantly longer duration of behavioral responses compared to the responses seen following injection of either substance alone. These data provide support for the hypothesis that primary afferent neurons might well be subject to similar mechanisms that result in wind-up or central sensitization of spinal cord neurons.     

Chemosensory function and dysfunction

Taste and smell are fundamental sensory systems essential in nutrition and food selection, for the hedonic and sensory experience of food, for efficient metabolism, and, in general, for the maintenance of a good quality of life. The gustatory and olfactory systems demonstrate a diversity of transduction mechanisms, and during the last decade, considerable progress has been made toward our understanding of the basic mechanisms of taste and smell. Understanding normal chemosensory function helps clarify the molecular events that underlie taste and smell disorders. At least 2,000,000 Americans suffer from chemosensory disorders--a number that is likely to grow as the aging segment of the population increases. Smell disorders are more frequent than taste disturbances, due to the vulnerability and anatomical distinctiveness of the olfactory system, and because a decline in olfactory function is part of the normal aging process. Common gustatory and olfactory complaints are due to a number of medications, to upper respiratory infections, to nasal and paranasal sinus diseases, and to damage to peripheral nerves supplying taste and smell. Most chemosensory complaints have an identifiable cause. Although diagnosis of taste and smell disorders has improved considerably over the last two decades, treatment of these disorders is still limited to conditions with discernible and reversible causes. Future research is needed for a better understanding of chemosensory mechanisms, establishing improved diagnostic procedures, and disseminating knowledge on chemosensory disorders among practitioners and the general public.     

Direct glutamate-mediated presynaptic inhibition of sensory afferents by the postsynaptic motor neurons

An in vitro preparation of the crayfish central nervous system was used to study a negative feedback control exerted by the glutamatergic motor neurons (MNs) on to their presynaptic cholinergic sensory afferents. This negative control consists in small amplitude, slowly developing depolarizations of the primary afferents (sdPADs) strictly timed with MN bursts. They were not blocked by picrotoxin, but were sensitive to glutamate non-N-methyl-D-aspartate (NMDA) antagonists. Intracellular recordings were performed within thin branches of sensory terminals while electrical antidromic stimulation were applied to the motor nerves, or while glutamate (the MN neurotransmitter) was pressure-applied close to the recording site. Electrical motor nerve stimulations and glutamate pressure application had similar effects on to sensory terminals issued from the coxo-basipodite chordotonal organ (CBTs): like sdPADs, both stimulation-induced depolarizations were picrotoxin-resistant and were dramatically reduced by non-NMDA antagonist bath application. These results indicate that sdPADs are likely directly produced by MNs during locomotor activity. A functional scheme is proposed.     

Why Americans eat what they do: taste, nutrition, cost, convenience, and weight control concerns as influences on food consumption

OBJECTIVE: To examine the self-reported importance of taste, nutrition, cost, convenience, and weight control on personal dietary choices and whether these factors vary across demographic groups, are associated with lifestyle choices related to health (termed health lifestyle), and actually predict eating behavior. DESIGN: Data are based on responses to 2 self-administered cross-sectional surveys. The main outcomes measured were consumption of fruits and vegetables, fast foods, cheese, and breakfast cereals, which were determined on the basis of responses to questions about usual and recent consumption and a food diary. SUBJECTS/SETTING: Respondents were a national sample of 2,967 adults. Response rates were 71% to the first survey and 77% to the second survey (which was sent to people who completed the first survey). STATISTICAL ANALYSES: Univariate analyses were used to describe importance ratings, bivariate analyses (correlations and t tests) were used to examine demographic and lifestyle differences on importance measures, and multivariate analyses (general linear models) were used to predict lifestyle cluster membership and food consumption. RESULTS: Respondents reported that taste is the most important influence on their food choices, followed by cost. Demographic and health lifestyle differences were evident across all 5 importance measures. The importance of nutrition and the importance of weight control were predicted best by subject's membership in a particular health lifestyle cluster. When eating behaviors were examined, demographic measures and membership in a health lifestyle cluster predicted consumption of fruits and vegetables, fast foods, cheese, and breakfast cereal. The importance placed on taste, nutrition, cost, convenience, and weight control also predicted types of foods consumed. APPLICATIONS: Our results suggest that nutritional concerns, per sc, are of less relevance to most people than taste and cost. One implication is that nutrition education programs should attempt to design and promote nutritious diets as being tasty and inexpensive.     

Carnosine modulates zinc and copper effects on amino acid receptors and synaptic transmission

Carnosine is a dipeptide which is highly concentrated in mammalian olfactory sensory neurons along with zinc and/or copper, and glutamate. Although carnosine has been proposed as a neurotransmitter or neuromodulator, no specific function for carnosine has been identified. We used whole-cell current- and voltage-clamp recording to examine the direct effects and neuromodulatory actions of carnosine on rat olfactory bulb neurons in primary culture. Carnosine did not evoke a membrane current or affect currents evoked by glutamate, GABA or glycine. Copper and zinc inhibited NMDA and GABA receptor-mediated currents and inhibited synaptic transmission. Carnosine prevented the actions of copper and reduced the effects of zinc. These results suggest that carnosine may indirectly influence neuronal excitability by modulating the effects of zinc and copper.     

Responses to the sensory properties of fat of neurons in the primate orbitofrontal cortex

The primate orbitofrontal cortex is a site of convergence of information from primary taste, olfactory, and somatosensory cortical areas. We describe the responses of a population of single neurons in the orbitofrontal cortex that responds to fat in the mouth. The neurons respond, when fatty foods are being eaten, to pure fat such as glyceryl trioleate and also to substances with a similar texture but different chemical composition such as paraffin oil (hydrocarbon) and silicone oil [Si(CH3)2O)n]. This is evidence that the neurons respond to the oral texture of fat, sensed by the somatosensory system. Some of the population of neurons respond unimodally to the texture of fat. Other single neurons show convergence of taste inputs, and others of olfactory inputs, onto single neurons that respond to fat. For example, neurons were found that responded to the mouth feel of fat and the taste of monosodium glutamate (both found in milk), or to the mouth feel of fat and to odor. Feeding to satiety reduces the responses of these neurons to the fatty food eaten, but the neurons still respond to some other foods that have not been fed to satiety. Thus sensory-specific satiety for fat is represented in the responses of single neurons in the primate orbitofrontal cortex. Fat is an important constituent of food that affects its palatability and nutritional effects. The findings described provide evidence that the reward value (or pleasantness) of the mouth feel of fat is represented in the primate orbitofrontal cortex and that the representation is relevant to appetite.     

Molecular neurogenetics of chemotaxis and thermotaxis in the nematode Caenorhabditis elegans

Chemotaxis and thermotaxis in Caenorhabditis elegans are based on the chemical senses (smell and taste) and the thermal sense, respectively, which are important for the life of the animal. Laser ablation experiments have allowed identification of sensory neurons and some interneurons required for these senses. Many mutants that exhibit various abnormalities have been isolated and analyzed. These studies have predicted novel signaling pathways whose components include a putative odorant specific transmembrane receptor (ODR-10) and a cyclic nucleotide-gated channel (TAX-4/TAX-2) functioning in taste and thermosensation as well as in smell. The emerging picture of the mechanisms of sensory transduction in C. elegans seems to be basically similar to what is known of visual and olfactory sensory transduction in vertebrates. Thus, molecular and cellular analyses of chemotaxis and thermotaxis in C. elegans have proved useful and will continue to provide significant implications for the molecular basis of sensory systems in higher animals.     

Effects of obesity and set point on taste responsiveness and ingestion in humans.

Two experiments, with a total of 120 undergraduates, investigated the relationships among degree of obesity, nearness to set point for adipose tissue mass, and responsiveness to taste. In Exp I, Ss rated milkshakes varying in sweetness intensity. Overweight and normal weight Ss did not have significantly different detection thresholds or preference ratings. However, overweight Ss worked longer than normals to obtain their most preferred substance. Exp II varied the ease of tasting the milkshake. Preference and food intake of moderately overweight Ss were significantly more influenced by tastes that they found positive and negative than was the preference of normal weight or obese Ss. The ease of ingesting the taste substance also significantly influenced preference and food intake of the moderately obese only. Nearness to set point, operationally defined as weight stability for 2 yrs, had no significant effects. (27 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Acquired aversions as the basis for varied diets of ruminants foraging on rangelands

Ruminants eat an array of plant species that vary in nutrients and toxins. This selection makes intuitive sense, but no theories adequately explain this diversity. Some maintain it reduces the likelihood of overingesting toxins, whereas others contend it meets nutritional needs. Nevertheless, herbivores seek variety even when toxins are not a concern and nutritional needs are met. I offer another explanation for this behavior, one which encompasses the avoidance of toxins and the acquisition of nutrients. A key concept in this theory is aversion, the decrease in preference for food just eaten as a result of sensory input (a food's taste, odor, texture, i.e., its flavor) and postingestive effects (effects of nutrients and toxins on chemo-, osmo-, and mechano-receptors) unique to each food. Aversions are pronounced when foods contain toxins or high levels of rapidly digestible nutrients; they also occur when foods are deficient in specific nutrients. Aversions occur even when animals eat nutritionally adequate foods because satiety (satisfied to the full) and surfeit (filled to nauseating excess) represent points along a continuum, and there is a fine line between satiety and aversion. Thus, eating any food is likely to cause a mild aversion, and eating a food too frequently or in excess is likely to cause a strong aversion. Aversions are involuntary and are not the result of conscious decisions by an animal. Aversions yield benefits (e.g., obtain a balanced diet, reduce ingestion of toxic foods, optimize foraging and rumination times, sample foods, maintain a diverse microflora in the rumen) that are often mistaken as the cause of varied diets. In this article, I discuss the subtle ways in which aversions diminish preference and cause animals to eat a variety of foods.     

Gustation and ingestive behavior in the rat.

Studied the contribution of taste to the quantitative control of intake and body weight in 15 male Wistar rats. Gustatory deafferentation was produced by combined bilateral section of the chorda tympani and glossopharyngeal nerves and the pharyngeal branch of the vagus nerve. The procedure did not affect the condition of the mouth, impair orientation to sensory stimuli, or severely disrupt the consummatory sequence. Nevertheless, deafferented Ss showed reduced responsiveness to food and water, elevated quinine HCl aversion thresholds, and impaired feeding efficiency. Food and water intake and body weight remained below control levels for at least 40–50 days postoperatively. Food intake deficits were not observed when Ss were offered ad lib pablum, a result suggesting that increased palatability could compensate for decreased gustatory input. The order of recovery of food intake through different diet types suggests a heightened responsiveness to taste stimuli. The effects of gustatory deafferentation are compared with those seen after section of trigeminal orosensory nerves, and the differential contributions of oral taste and oral touch are discussed. (42 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Psychoanalysis, Neuroscience, and Cognitive Psychology: Some Samples From Recent Research.

As a demonstration of ways in which the burgeoning work in neuroscience and cognitive psychology have important relevance for psychoanalysis, samples are offered from the literature of the past 9 months. Reports address such topics as neural transmission at the cellular level; a revolutionary new instrument capable of recording neurons at work in the brain; the latest data on the effect of psychotropic drugs on children; the research validity of measuring emotional expression (e.g., do baby rats cry); longitudinal work on inhibited children; the importance of implicit cognitive processes; emotion, attention, and the amygdala; and recent findings about glutamate, "the workhorse of the brain," and how its deleterious effects in neurodegenerative disorders may be modulated. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Increased histidine preference during specific alteration of rhythm of environmental temperature stress in rats.

It has been reported that specific alteration of rhythm of environmental temperature (SART) stress induces various physiological changes. In this study, changes in taste preference during SART stress were investigated in rats. Rats were given free access to six amino acid solutions, saline, and water in a choice paradigm. During SART stress, daily food intake increased significantly by 50% whereas the rate of body weight gain decreased significantly to one third that observed during the prestress baseline period. In addition, consumption of histidine solution increased significantly, whereas intakes of water, monosodium glutamate, saline, glycine, arginine, lysine, and threonine were unaffected. Results suggest that a specific preference for histidine emerges during SART stress, which may be related to the stress-induced changes in the histamine turnover in the brain and peripheral tissues. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

HLA and graft-versus-host disease: a population-based study of HLA phenotypes of Jewish and Arabic bone marrow transplanted patients in Israel

Secondary epidermal solitary chemosensory cells (SCCs) are widespread among the primary aquatic vertebrates. They resemble taste bud sensory cells in fine structure and may be innervated from facial or spinal nerves. According to previous studies, SCCs may constitute a water sampling system in the contexts of predator avoidance, habitat recognition and, in some cases, finding food. By quantitative scanning (SEM) and transmission electron microscopy (TEM) in 60 specimens (57 SEM, 3 TEM) of 16 developmental stages, from pre-hatchlings to adults, we describe the ontogenetic development of SCC densities and shapes of sensory apices in the zebrafish, Danio rerio. This is put into perspective with the ontogeny of external taste buds. Just prior to hatching, 3 days after fertilization (3d AF), sensory apices of SCCs penetrate between the squamous epidermal cells, whereas taste bud pores only appear at the onset of exogenous feeding (5d AF). SCC densities increase sharply from hatching shortly after metamorphosis (25d AF) up to 6 x 10(3) per mm2 on the head and remain relatively constant in density thereafter. Conservatively estimated, there may be approximately 3.2 x 10(5) SCCs on the head and 1 x 10(6) SCCs on the entire body surfaces of a zebrafish 180d AF. SCCs are spread evenly, but are 2- to 5-fold higher in density along the head than along the body. Sensory apices are brush-like in hatchlings and early juveniles, but tend to consist of a single villus in the adults. This ontogenetic change of SCC apices parallels the evolutionary change from 'oligovillous' cells in lampreys and elasmobranchs to the 'monovillous' SCCs in the advanced actinopterygian teleosts.     

Effects of degree of obesity, age of onset, and weight loss on responsiveness to sensory and external stimuli.

Four experiments tested responsiveness to external and sensory stimuli in human females who differed in degree of overweight and in age of onset of their obesity. A total of 196 overweight Ss, ranging in age from 12–58 yrs and representing both juvenile- and adult-onset obese groups, served as Ss; in Exps I and IV, a total of 14 normal-weight Ss were used as controls. The effects of weight loss on external responsiveness were assessed by testing Ss before and after weight reduction in order to determine the role of energy deficit and deprivation in mediating heightened responsiveness to external cues. There was no significant positive correlation between degree of overweight and degree of external responsiveness. In general, the age of onset of obesity was also not a factor in the degree of externality. Weight loss did not change responses to visual and cognitive cue salience manipulatons in measures of feeding, emotionality, time perception, and short-term recall, while responsiveness to variations in taste palatability increased following weight reduction. While external cue responsiveness does not appear to result from adiposity per se or from deprivation, responsiveness to taste stimuli may reflect these parameters. The role of external and sensory cues in the development and maintenance of obesity is discussed. (42 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Expression of metabotropic glutamate receptor 1 isoforms in the substantia nigra pars compacta of the rat

Metabotropic glutamate receptors, which are linked via G-proteins to second messenger systems, have been implicated in the physiological regulation of dopaminergic neurons of the substantia nigra pars compacta as well as in neurodegeneration. Of the eight known metabotropic glutamate receptors, metabotropic glutamate receptor 1 is the most abundant subtype in the substantia nigra pars compacta. Metabotropic glutamate receptor 1 is alternatively spliced at the carboxy terminal region to yield five variants: 1a, 1b, 1c, 1d and a form recently identified in human brain, 1g. We used an antibody recognizing metabotropic glutamate receptor 1, and another recognizing the splice form la only, to study the localization of these receptors in dopaminergic neurons identified by the presence of tyrosine hydroxylase. Metabotropic glutamate receptor immunoreactivity was present within the somata, axons, and dendrites of substantia nigra pars compacta neurons. The 1a splice form specific antibody, however, did not label these cells, suggesting that they express a metabotropic glutamate receptor 1 splice form different from 1a. In situ hybridization with splice form-specific oligonucleotide probes was used to determine which of the other known metabotropic glutamate receptor 1 splice forms might be present in the substantia nigra pars compacta. Each probe produced a very distinct labelling pattern in the rat brain with the exception of the 1g specific probe which produced only background signal. Substantia nigra pars compacta neurons were most intensely labelled by the metabotropic glutamate receptor 1d splice form specific probe. Metabotropic glutamate receptor 1a was expressed weakly whereas there was no detectable 1b, c, or g signal in the substantia nigra pars compacta. These data demonstrate that metabotropic glutamate receptor 1 protein is present within the perikarya and processes of dopaminergic neurons in the substantia nigra pars compacta. The majority of this protein is not the 1a splice form, which is abundant in other brain regions, and may be the 1d isoform. Since splicing alters the carboxy terminus of the receptor, it is likely to affect the interaction of the receptor with intracellular signalling systems.     

Dualism redux in recent neuroscience: "Theory of mind" and "embodied simulation" hypotheses in light of historical debates about perception, cognition, and mind.

How do we, as humans, take in the feelings and thoughts of other people? Theory-of-Mind (ToM) and Embodied Simulation (ES) approaches hypothesize divergent neural and behavioral mechanisms underlying intersubjectivity. ToM investigators assert that humans take in the belief states and intentions of another person by holding "a theory of mind" that cognitively posits the other person's mental contents, with some experiments identifying the right temporo-parietal junction as a specific ToM brain region. ES theorists hypothesize that humans perceive the other's state of mind by simulating his/her actions, emotions, and goals in the "mirror neuron system" in the brain. A historical review suggests these understandings rely on opposing, dualist models of cognition and perception. William James's intervention on this earlier debate is informative in anticipating recent findings in low-level sensory neuroscience. Of specific interest are studies showing that intersubjectivity and low-level sensory attentional filtering are both processed in the same cortical area (the temporo-parietal junction) suggesting that the ability to entertain other minds may be related to the ability to perceive salient stimuli during attention-demanding tasks. (PsycINFO Database Record (c) 2010 APA, all rights reserved)