Hemispheric dominance of cortical activity evoked by focal electrogustatory stimuli.

Functional magnetic resonance imaging was used to observe cortical hemodynamic responses to electric taste stimuli applied separately to the right and left sides of the tongue tip. In 11 right-handed normal adults activation occurred primarily in the insular cortex, superior temporal lobe, inferior frontal lobe, including premotor regions, and in inferior parts of the postcentral gyrus. Unexpectedly, the location and laterality of activation were largely identical regardless of the side of the tongue stimulated. Activation in the superior insula, the presumed location of primary gustatory cortex, was predominantly, but not exclusively, in the right hemisphere, whereas central (more inferior) insular activations were more evenly bilateral. Right hemispheric dominance of activation also occurred in premotor regions (Brodmann areas 6 and 44), whereas left hemispheric dominance occurred only in the superior temporal cortex (Brodmann areas 22/42). The electric taste-evoked hemodynamic response pattern was more consistent with activation of the gustatory system than activation of somatosensory systems. The results suggest that the sites for cortical processing of electric taste information are dependent on hemispheric specialization.     

Rat gustatory neurons in the geniculate ganglion express glutamate receptor subunits

Taste receptor cells are innervated by primary gustatory neurons that relay sensory information to the central nervous system. The transmitter(s) at synapses between taste receptor cells and primary afferent fibers is (are) not yet known. By analogy with other sensory organs, glutamate might a transmitter in taste buds. We examined the presence of AMPA and NMDA receptor subunits in rat gustatory primary neurons in the ganglion that innervates the anterior tongue (geniculate ganglion). AMPA and NMDA type subunits were immunohistochemically detected with antibodies against GluR1, GluR2, GluR2/3, GluR4 and NR1 subunits. Gustatory neurons were specifically identified by retrograde tracing with fluorogold from injections made into the anterior portion of the tongue. Most gustatory neurons in the geniculate ganglion were strongly immunoreactive for GluR2/3 (68%), GluR4 (78%) or NR1 (71%). GluR1 was seen in few cells (16%). We further examined if glutamate receptors were present in the peripheral terminals of primary gustatory neurons in taste buds. Many axonal varicosities in fungiform and vallate taste buds were immunoreactive for GluR2/3 but not for NR1. We conclude that gustatory neurons express glutamate receptors and that glutamate receptors of the AMPA type are likely targeted to synapses within taste buds.     

Gustatory processing is dynamic and distributed

The process of gustatory coding consists of neural responses that provide information about the quantity and quality of food, its generalized sensation, its hedonic value, and whether it should be swallowed. Many of the models presently used to analyze gustatory signals are static in that they use the average neural firing rate as a measure of activity and are unimodal in the sense they are thought to only involve chemosensory information. We have recently elaborated upon a dynamic model of gustatory coding that involves interactions between neurons in single as well as in spatially separate, gustatory and somatosensory regions. We propose that the specifics of gustatory responses grow not only out of information ascending from taste receptor cells, but also from the cycling of information around a massively interconnected system.     

Changes in neurotrophin-3 messenger RNA expression patterns in the prenatal rat tongue suggest guidance of developing somatosensory nerves to their final targets

While brain-derived neurotrophic factor (BDNF) messenger RNA (mRNA) has been localized in the developing gustatory epithelium, little information is available about neurotrophin-3 (NT-3) mRNA expression pattern in the prenatal developing gustatory and lingual epithelium. In the present study, using in situ hybridization histochemistry, we report on NT-3 mRNA expression in the tongue of rats. At embryonic day (E) 13–17, NT-3 mRNA was expressed subepithelially in the periphery of the developing tongue, as well as among developing muscle. At E19, there was a shift in the expression of NT-3 mRNA. It was then expressed in the surface epithelium of the developing tongue in the developing filiform papillae and, in higher concentrations, in top-surface and fringe epithelium of the developing circumvallate papillae, and top- and lateral-surface epithelium of the developing fungiform papillae. NT-3 mRNA expression in areas rich in somatosensory innervation of the tongue, as well as its specific expression in defined regions compared with BDNF, and the decreased labeling noted from prenatal and early postnatal animals to adults indicate a specific role for NT-3 in the development of lingual somatosensory innervation, as well as for maintenance of this innervation.     

Taste recognition: food for thought

The ability to identify food that is nutrient-rich and avoid toxic substances is essential for an animal's survival. Although olfaction and vision contribute to food detection, the gustatory system acts as a final checkpoint control for food acceptance or rejection behavior. Recent studies with model organisms such as mice and Drosophila have identified candidate taste receptors and examined the logic of taste coding in the periphery. Despite differences in terms of gustatory anatomy and taste-receptor families, these gustatory systems share a basic organization that is different from other sensory systems. This review will summarize our current understanding of taste recognition in mammals and Drosophila, highlighting similarities and raising several as yet unanswered questions.     

Gustatory projections from the nucleus of the solitary tract to the parabrachial nuclei in the hamster.

Taste-responsive cells in the nucleus of the solitary tract (NST) either project to the parabrachial nuclei (PbN) of the pons, through which taste information is transmitted to forebrain gustatory nuclei, or give rise to axons terminating locally within the medulla. Numerous anatomical studies clearly demonstrate a substantial projection from the rostral NST, where most taste-responsive cells are found, to the PbN. In contrast, previous electrophysiological studies in the rat have shown that only a small proportion (21-45%) of taste-responsive NST cells are antidromically activated from the PbN, suggesting that less than half the cells recorded from the NST are actually involved in forebrain processing of gustatory information. In the present experiment we investigated the projections from the NST to the PbN electrophysiologically in urethane anesthetized hamsters. Responses of 101 single neurons in the rostral NST were recorded extracellularly following lingual stimulation with 32 mM NaCl, sucrose and quinine hydrochloride (QHCl) and 3.2 mM citric acid. The taste-responsive region of the PbN was identified electrophysiologically and stimulated with a concentric bipolar electrode to antidromically activate each NST cell. Of the 101 taste-responsive NST cells, 81 (80.2%) were antidromically activated from the ipsilateral PbN. The mean firing rates to taste stimulation and the spontaneous activity of these projection neurons were significantly greater than those of non-projecting cells. Every sucrose-best neuron in the sample projected to the PbN. The mean conduction velocity of the 23 QHCl-best neurons was significantly lower than that of the other 58 PbN projection neurons, suggesting that the most QHCl-responsive cells are a subset of smaller neurons. These data show that a large majority of NST cells responsive to taste stimulation of the anterior tongue project to the gustatory subdivisions of the PbN and that these cells have the most robust responses to gustatory stimulation.     

Developmental time course of peripheral cross‐modal sensory interaction of the trigeminal and gustatory systems

Few sensory modalities appear to engage in cross‐modal interactions within the peripheral nervous system, making the integrated relationship between the peripheral gustatory and trigeminal systems an ideal model for investigating cross‐sensory support. The present study examined taste system anatomy following unilateral transection of the trigeminal lingual nerve (LX) while leaving the gustatory chorda tympani intact. At 10, 25, or 65 days of age, rats underwent LX with outcomes assessed following various survival times. Fungiform papillae were classified by morphological feature using surface analysis. Taste bud volumes were calculated from histological sections of the anterior tongue. Differences in papillae morphology were evident by 2 days post‐transection of P10 rats and by 8 days post in P25 rats. When transected at P65, animals never exhibited statistically significant morphological changes. After LX at P10, fewer taste buds were present on the transected side following 16 and 24 days survival time and remaining taste buds were smaller than on the intact side. In P25 and P65 animals, taste bud volumes were reduced on the denervated side by 8 and 16 days postsurgery, respectively. By 50 days post‐transection, taste buds of P10 animals had not recovered in size; however, all observed changes in papillae morphology and taste buds subsided in P25 and P65 rats. Results indicate that LX impacts taste receptor cells and alters epithelial morphology of fungiform papillae, particularly during early development. These findings highlight dual roles for the lingual nerve in the maintenance of both gustatory and non‐gustatory tissues on the anterior tongue. © 2015 Wiley Periodicals, Inc. Develop Neurobiol 76: 626–641, 2016     

哺乳动物味觉的细胞生物学

味觉对于生命具有重要作用,在一定程度上决定了动物对食物的选择。哺乳动物味觉识别主要依赖于舌味蕾中的味细胞,味蕾由50～100个极化的神经上皮细胞聚集而成。通过对味蕾细胞的分析显示,味蕾是一种精巧的单元结构。这篇文章综述了味蕾细胞的形态、结构功能、细胞生物学活性以及味觉信息的传导。     

Gustatory imagery reveals functional connectivity from the prefrontal to insular cortices traced with magnetoencephalography

Our experience and prejudice concerning food play an important role in modulating gustatory information processing; gustatory memory stored in the central nervous system influences gustatory information arising from the peripheral nervous system. We have elucidated the mechanism of the “top-down” modulation of taste perception in humans using functional magnetic resonance imaging (fMRI) and demonstrated that gustatory imagery is mediated by the prefrontal (PFC) and insular cortices (IC). However, the temporal order of activation of these brain regions during gustatory imagery is still an open issue. To explore the source of “top-down” signals during gustatory imagery tasks, we analyzed the temporal activation patterns of activated regions in the cerebral cortex using another non-invasive brain imaging technique, magnetoencephalography (MEG). Gustatory imagery tasks were presented by words (Letter G-V) or pictures (Picture G-V) of foods/beverages, and participants were requested to recall their taste. In the Letter G-V session, 7/9 (77.8%) participants showed activation in the IC with a latency of 401.7±34.7 ms (n = 7) from the onset of word exhibition. In 5/7 (71.4%) participants who exhibited IC activation, the PFC was activated prior to the IC at a latency of 315.2±56.5 ms (n = 5), which was significantly shorter than the latency to the IC activation. In the Picture G-V session, the IC was activated in 6/9 (66.7%) participants, and only 1/9 (11.1%) participants showed activation in the PFC. There was no significant dominance between the right and left IC or PFC during gustatory imagery. These results support those from our previous fMRI study in that the Letter G-V session rather than the Picture G-V session effectively activates the PFC and IC and strengthen the hypothesis that the PFC mediates “top-down” control of retrieving gustatory information from the storage of long-term memories and in turn activates the IC.     

Chemoreception in chronically anosmiated fish: A phenomenon of compensatory development of the gustatory system

The main results of studying the phenomenon of recovery of sensitivity to food smells after a long olfactory deprivation found in fish are presented. This ability is detected in fish of different taxonomic groups (Chondrostei, Teleostei) having normally external gustatory receptors. At the behavioral level, this effect starts to manifest itself six weeks upon anosmia, and its development apparently ends during the first three to five months in a partial, rather than complete, recovery of sensitivity to food smells. A more prolonged olfactory deprivation (6–14 months) does not cause any further decrease in the threshold concentration of stimulus solutions. The response to other types of chemical signals (alarm pheromone) is not restored. The ability of fish after a prolonged olfactory deprivation to recover sensitivity to food smells is provided by compensatory processes taking place in the external gustatory system. Recovery occurs due to substances that have taste attraction for fish. Species were found in which the food search response in the norm is provided simultaneously by the olfactory and the gustatory systems. The anosmia of such fish leads only to a partial decrease in the efficiency of search behavior. The complementarity of chemosensory systems, their functional interaction, capacity for a compensatory development, and vicariation are considered sensory mechanisms providing the reliability of realization by fish of the most important behavioral patterns, of food, in particular.     

Comparative anatomy of the tongue of Pan troglodytes (Blumenbach, 1799) and other primates. II. Findings in other primates

This 2nd part of our studies shows that the papilla foliata is fully developed in Pan, Cebus, and Macaca; in Prosimians the papilla foliata is well developed in Lemur and Chirogaleus. In Galago crassicaudatus, this papilla is missing. Among 3 individuals of Microcebus, the papilla foliata was differently developed: in 2 cases, the tongue exhibited only 2 on both sides and a very low folia. Taste buds were found in the epithelium of only one side of each folium. In the 3rd case, the folia of the papilla were developed only on one side of the tongue, whereas, on the other side, a typical papilla was missing. Instead of the papilla, the tongue of the same animal exhibited a hillock-like structure; it is a gustatory hillock which exhibited many taste buds. There were 3 gustatory hillocks in all of the specimens of Tupaia glis; they are situated on both sides of the tongue.     

Neurotransmitter and Neuromodulator Activity in the Gustatory Zone of the Nucleus Tractus Solitarius

The rostral nucleus of the solitary tract (rNST) is the firstcentral relay in the gustatory pathway. While previous investigationshave provided a wealth of information on the pattern of centralterminations of gustatory afferent fibers, the morphology ofsynaptic connections of rNST neurons and responses of secondorder neurons to taste stimuli applied to the tongue, littleis known regarding the neurophysiological characteristics ofsynaptic transmission in rNST. We have used an in vitro brainslice preparation of the rNST to study the intrinsic biophysicalproperties, neuropharmacology and synaptic responses of rNSTneurons. These experiments have revealed that rNST neurons respondto the excitatory amino acid neurotransmitter glutamate, aswell as the inhibitory amino acid neurotransmitter     

OFF-RESPONSE OF GUSTATORY NERVE FOLLOWING TERMINATION OF QUININE STIMULI APPLIED TO THE FROG TONGUE

After a Ringer-adapted frog tongue was stimulated with 0.001M quinine-HCl(Q-HCl) in deionized water, the Ringer rinse ofthe tongue elicited a large gustatory nerve response. Sincethe Ringer-adapted tongue did not respond to Ringer solution,this nerve response after Q-HC1 is termed an off-response. Itwas revealed that the off-response of gustatory nerve to theRinger rinse resulted from the enhancing effect of Q-HCl adaptationupon the response to 111.2 mM NaCl component of Ringer whichwas ineffective for Ringer-adapted tongue. Weaker enhancementof the response to NaCl of Ringer was also produced by adaptingthe tongue to water. Therefore, the enhancing effect of Q-HClin deionized water is the summed effect of Q-HCl solute andwater solvent. Microelectrode study revealed that during theadaptation to Q-HCl the membrance potential of some NaCl-sensitivetaste cells was more hyperpolarized than that of Ringer-adaptedstate. The larger membrane potential maintained under Q-HCladaptation was markedly depolarized by the NaCl component ofthe Ringer. This sufficient depolarization response of the tastecells might be associated with generation of the off-responsein the gustatory nerve.     

Development of the tongue and taste disks in Pelobates fuscus

In the tadpole of Pelobates fuscus the process of tongue formation starts at the 32nd developmental stage. In more advanced stages (older than 38th) fast anterior and faucial growth of the tongue fold has been observed. This process is accompanied by the development of the gustatory organs. The dorsal surface of the tongue fold, smooth at the beginning, in older tadpoles (developmental stages 36-39th) forms protrusions in which gustatory organs of the taste disk type (TDs) develop. In the 41 st tadpole developmental stage anlages of TDs are formed by elongated cells, located more or less perpendicularly to the surface of the tongue. The diameter of the sensory area of a TD at the 45th developmental stage amounts to 94 microm, while in metamorphosed individuals it reaches 130-140 microm. At the base of a TD the presence of basal cell morphologically similar to that of Merkel cell was observed at the 42nd developmental stage of a tadpole. Fully developed afferent synaptic connections in the sensory epithelium of a TD were found starting from the 44th developmental stage. Single synaptic vesicles with an electron-dense core were observed in gustatory cells as early as at the 41 st developmental stage of the tadpole. From the observations reported here it can be inferred that in Pelobates fuscus development of both the tongue and TDs is similar to that already described in the representatives of the Rana genus.     

Seasonality constraints to livestock grazing intensity

Increasing food production is essential to meet the future food demand of a growing world population. In light of pressing sustainability challenges such as climate change and the importance of the global livestock system for food security as well as GHG emissions, finding ways to increasing food production sustainably and without increasing competition for food crops is essential. Yet, many unknowns relate to livestock grazing, in particular grazing intensity, an essential variable to assess the sustainability of livestock systems. Here, we explore ecological limits to grazing intensity (GI; i.e. the fraction of net primary production consumed by grazing animals) by analysing the role of seasonality in natural grasslands. We estimate seasonal limitations to GI by combining monthly net primary production data and a map of global livestock distribution with assumptions on the length of nonfavourable periods that can be bridged by livestock (e.g. by browsing dead standing biomass, storage systems or biomass conservation). This allows us to derive a seasonality‐limited potential GI, which we compare with the GI prevailing in 2000. We find that GI in 2000 lies below its potential on 39% of the total global natural grasslands, which has a potential for increasing biomass extraction of up to 181 MtC/yr. In contrast, on 61% of the area GI exceeds the potential, made possible by management. Mobilizing this potential could increase milk production by 5%, meat production by 4% or contribute to free up to 2.8 Mio km² of grassland area at the global scale if the numerous socio‐ecological constraints can be overcome. We discuss socio‐ecological trade‐offs, which may reduce the estimated potential considerably and require the establishment of sound monitoring systems and an improved understanding of livestock system's role in the Earth system.     

The influence of gustatory and olfactory experiences on responsiveness to reward in the honeybee

Background

Honeybees (Apis mellifera) exhibit an extraordinarily tuned division of labor that depends on age polyethism. This adjustment is generally associated with the fact that individuals of different ages display different response thresholds to given stimuli, which determine specific behaviors. For instance, the sucrose-response threshold (SRT) which largely depends on genetic factors may also be affected by the nectar sugar content. However, it remains unknown whether SRTs in workers of different ages and tasks can differ depending on gustatory and olfactory experiences.

Methodology

Groups of worker bees reared either in an artificial environment or else in a queen-right colony, were exposed to different reward conditions at different adult ages. Gustatory response scores (GRSs) and odor-memory retrieval were measured in bees that were previously exposed to changes in food characteristics.

Principal Findings

Results show that the gustatory responses of pre-foraging-aged bees are affected by changes in sucrose solution concentration and also to the presence of an odor provided it is presented as scented sucrose solution. In contrast no differences in worker responses were observed when presented with odor only in the rearing environment. Fast modulation of GRSs was observed in older bees (12–16 days of age) which are commonly involved in food processing tasks within the hive, while slower modulation times were observed in younger bees (commonly nurse bees, 6–9 days of age). This suggests that older food-processing bees have a higher plasticity when responding to fluctuations in resource information than younger hive bees. Adjustments in the number of trophallaxis events were also found when scented food circulated inside the nest, and this was positively correlated with the differences in timing observed in gustatory responsiveness and memory retention for hive bees of different age classes.

Conclusions

This work demonstrates the accessibility of chemosensory information in the honeybee colonies with respect to incoming nectar. The modulation of the sensory-response systems within the hive can have important effects on the dynamics of food transfer and information propagation.     

Responses of the sea catfish Ariopsis felis to chemical defenses from the sea hare Aplysia californica

Ink secretion of sea hares (Aplysia spp.), which is a mixture of co-released ink from the ink gland and opaline from the opaline gland, protects sea hares from predatory invertebrates through diverse mechanisms. These include both aversive or deterrent compounds and also high concentrations of amino acids that stimulate the predators' chemical senses and divert the attack through phagomimicry or sensory disruption. The aim of the present study was to examine if sea hares also defend themselves from predatory vertebrates by interacting with their chemical senses. We used sea catfish, Ariopsis felis, in behavioral and electrophysiological experiments. Behavioral tests on sea catfish show that ink is aversive: when ink is added to palatable food items (noodles with food flavoring), the noodles are no longer eaten, and when ink is added to noodles without food flavoring, the noodles are avoided more than unflavored noodles. Behavioral tests also show that opaline and the amino acid components of either opaline or ink are appetitive. Electrophysiological recordings of chemosensory neuronal activity in the olfactory epithelium and maxillary barbels show that the olfactory and gustatory systems of sea catfish are highly stimulated by ink and opaline, and that the amino acid components of ink and opaline significantly contribute to these responses. Compounds generated by the activity of escapin, an L-amino acid oxidase in the secretion, are moderately stimulatory to both olfactory and gustatory systems. Taken together, our results support the idea that sea hares are chemically defended from predatory sea catfish largely through unpalatable chemical deterrents in ink, but possibly also through amino acids stimulating olfactory and gustatory systems and thus functioning through phagomimicry or sensory disruption.     

Ca2+-modulated membrane guanylate cyclase in the testes

To date, the calcium-regulated membrane guanylate cyclase Rod Outer Segment Guanylate Cyclase type 1 (ROS-GC1) transduction system in addition to photoreceptors is known to be expressed in three other types of neuronal cells: in the pinealocytes, mitral cells of the olfactory bulb and the gustatory epithelium of tongue. Very recent studies from our laboratory show that expression of ROS-GC1 is not restricted to the neuronal cells; the male gonads and the spermatozoa also express ROS-GC1. In this presentation, the authors review the existing information on the localization and function of guanylate cyclase with special emphasis on Ca²⁺-modulated membrane guanylate cyclase, ROS-GC1, in the testes. The role of ROS-GC1 and its Ca²⁺-sensing modulators in the processes of spermatogenesis and fertilization are discussed.