Quantification of fungiform papillae and taste pores in living human subjects

A method developed to quantify taste buds in living human subjectsto study the relationship between taste sensitivity and tastebud distribution was used to count the taste buds in 10 humansubjects; fungiform papillae were mapped in 12 subjects. Tastebuds were identified by staining taste pores with methyleneblue, and images of the papillae and their taste pores wereobtained with videomicroscopy and an image processor. Fungiformpapillae showed a 3.3-fold range in density, from 22.1 to 73.6papillae/cm² with an average of 41.1 ± 16.8/cm² (s.d.,n = 2). There was a 14-fold range in taste pore density, from36 to 511 pores/cm² among subjects, with an average of 193 ±133/cm² (s.d., n = 10). Fungiform papillae contained from 0to 22 taste pores, with an average per subject of 3.75 ±1.4 taste pores/papilla (s.d., n = 10). We hypothesize thatsome differences in human taste sensitivity may be related tothese variations in taste bud density.     

The effects of beta-bungarotoxin on the morphogenesis of taste papillae and taste buds in the mouse

Although it has been long accepted that innervation by a tastenerve is essential for maintenance of taste buds, it is notclear what role, if any, innervation plays in the morphogenesis oftaste papillae and taste bud development. The following studywas undertaken to determine what effects lack of sensory innervationhave on the development of taste papillae and the formationof taste buds in the mouse. Timed-pregnant female mice (n =3) at gestational day 12 (gd12) were anesthetized and a 1 µlsolution (1 µg/µl) of ß-bungarotoxin (ß-BTX),a neurotoxin that disrupts sensory and motor neuron development,was injected into the amniotic cavity of two embryos per dam.Two shams were injected with PBS. Fetuses were harvested atgd18, 1 day before birth, and four ß-BTX-injected embryos,two shams and two controls were fixed in buffered paraformaldehyde.Serial sections were examined for the presence and morphologyof taste papillae and taste buds. No nerve profiles were observedin ß-BTX-injected tongues. Although circumvallate papillaewere present on ß-BTX tongues, only five fungiform papillaecould be identified. Taste buds were present on a large percentageof fungiform papillae profiles (24% and on circumvallate papillaein sham and control fetuses; in contrast, no taste buds wereassociated with taste papillae in ß-BTX fetuses. Theseresults implicate a significant role for innervation in tastepapillae and taste bud morphogenesis.     

Salt taste and salt sensitivity in black adolescents

Limited support for the postulated association between tasteand blood pressure responses to NaCl has prompted investigationsof selected subgroups of subjects more likely to display a relationship.The present study included 20 black adolescents who were diagnosedas salt sensitive (n = 10) or salt insensitive (n = 10) basedupon blood pressure responses to 14 days of dietary salt loading(10 g NaCl/day). Each was administered a battery of taste tests,including measures of detection and recognition threshold, suprathresholdintensity responses and preference tests. No significant groupdifference was observed on any taste measure. While explanationsfor the failure to note differences between these individualswith marked variabilty in systemic reactivity to NaCl are offered,the present findings question the continued study of this issue.     

Effects of therapeutic doses of amiloride and hydrochlorothiazide on taste, saliva and salt intake in normotensive adults

Tentative evidence suggests that hydrochlorothiazide (HCTZ)and amiloride may alter salivary function, taste sensitivityand dietary intake. The effects of these diuretics on salivaryflow and composition, taste attributes (i.e. recognition thresholds,perceived intensity ratings and preference) as well as sodiumintake were further examined in 73 normotensive adults in adouble-blind, placebo-controlled, cross-over study. Followingrandom assignments to one of three groups [5 mg twice per dayof amiloride (n = 24), 50 mg/day of HCTZ (n = 24) or placebo(n = 25)], subjects took appropriate placebos for 2 weeks, activedrug for 8 weeks and placebo for 3 weeks. Relative to baselinevalues, amiloride treatment resulted in a significant 39% decreasein stimulated salivary sodium levels and a 28% decline in NaClrecognition thresholds. HCTZ significantly stimulated sodiumintake as the sodium/creatinine ratio was {small tilde}50% higherfollowing 4 weeks of drug use relative to baseline values. Subjectswere subjectively unaware of these changes. Potential mechanismsand clinical implications of these findings are considered.     

Genetic Sensitivity to 6-n-Propylthiouracil (PROP) and Hedonic Responses to Bitter and Sweet Tastes

Genetically mediated sensitivity to the bitter taste of 6-n-propylthiouracil(PROP) has been associated with greater acuity for bitter andfor some sweet tastes. Thus far, few studies have explored therelationship between PROP taste sensitivity and hedonic responsesto bitter and sweet. In this study, 87 normal-weight young womenwere divided into PROP non-tasters (n = 18), regular tasters(n = 49), and supertasters (n = 20), based on their PROP detectionthresholds and the scaling of five suprathreshold solutionsof PROP and NaCl. Non-tasters had thresholds >1.8 x 10^–4mol/l PROP. Supertasters had thresholds <3.2 x 10^–5mol/l PROP and PROP/NaCl ratios >1.70. As expected, dislikeof the bitter taste of PROP was determined by its perceivedintensity, which was greater among supertasters than among regulartasters or non-tasters. Significant correlations were observedbetween PROP taste thresholds and the sum of intensity ratings(r = –0.61) and between summed intensity and summed hedonicratings (r = –0.80). PROP taste sensitivity was weaklylinked to enhanced perception of sweet taste, but did not predicthedonic responses to sucrose or to saccharin solutions. Giventhat the dislike of PROP solutions is determined by their perceivedintensity, hedonic responses to PROP solutions may provide arapid way of screening for PROP taster status. Chem. Senses22: 27–37, 1997.     

Salivary ions and neural taste responses in the hamster

Saliva is a chemically complex fluid that bathes oral surfacesand may affect early events in mammalian gustation. We measuredchorda tympani responses to taste stimuli in hamsters (Mesocricetusauratus) while their tongues were adapted to either water, artificialsaliva or natural saliva. Artificial saliva on the tongue loweredneural responses to taste stimuli that were present in the artificialsaliva and to those stimuli that cross-adated with salivarycomponents. Changing from a water-adapted tongue to one soakedwith pilocarpine-stimulated saliva from donor hamsters led tosignificantly smaller responses to NaCl. Responses to sucrose,NH₄Cl and quinine were unaffected. Chemical analysis of hamstersaliva revealed ‘normal’ mammalian levels of K⁺,Ca²⁺ and Mg²⁺, but unexpectedly low levels of Na⁺ and Cl^–.     

Partitioning of [14C]sucrose and Acid Invertase Activity in Reproductive Organs of Pepper Plants in Relation to Their Abscission Under Heat Stress

The effect of heat stress on processes related to carbohydratepartitioning was investigated in young bell pepper (Capsicumannum L. cv. Maor) plants in relation to abscission of theirreproductive organs at different stages of development. None of the reproductive organs abscised after 5 d in a normalday/night temperature regime (25/18°C). With a temperatureregime of 35°C day, 25°C night, abscission occurredin only a small portion of the flower buds and none of the flowersand fruitlets. However, when temperatures in the day and nightwere reversed (25/35°C, day/night) all the buds and someof the flowers abscised during that time period. The young fruitat the first node did not abscise under any temperature regime.The abscission rate of the flower buds was reduced under heatstress if the developing fruit at the first node had been removed. High temperature during either the light or dark periods reducedthe export of [¹⁴C]sucrose from the source leaf (fed for 48h with [¹⁴C]sucrose). Both heat stress and fruit presence reduced the relative amountof [¹⁴C]sucrose which was exported to the flower buds, flowersand roots. Likewise, these treatments reduced the concentrationof reducing sugars in the reproductive organs. Concomitantly,the heat stress and fruit presence on the first node reducedthe activity of soluble acid invertase in the flower buds andthe roots, but not in young leaves. Overall, the results show that heat stress causes alternationin sucrose distribution in the plant, but may also have specificeffects on metabolic activities related to sucrose import andutilization in flower buds and flowers which in turn may enhancetheir abscission. Bell pepper, (Capsicum annuum L. cv. Maor), abscission, acidinvertase, heat stress, reproductive organs, sink leaves. Bell pepper, (Capsicum annuum L. cv. Maor), abscission, acid invertase, heat stress, reproductive organs, sink leaves.     

Cytokeratin 14 is expressed in immature cells in rat taste buds

We analyzed the differentiation of taste bud cells, by precisely describing expression profiles of cytokeratins (CKs) 8 and 14 in relation to those of marker molecules including label of 5-bromo-2′-deoxy uridine (BrdU) injected. In rat circumvallate papillae, cell division was observed at the basal layer of the epithelium expressing CK14 and located outside taste buds. The progenitor cells began to migrate toward the apical surface and maintained CK14 expression at 1 day after BrdU injection (day 1). On the other hand, a minor population of newly divided cells was infrequently incorporated into taste buds and also maintained CK14 expression at day 1. In taste buds, the conversion of CK subtypes occurred from CK14 to cytokeratin 8 (CK8) at day 2–3, showing the differentiation from immature cells expressing CK14 into mature or maturing cells expressing CK8. Functionally matured cells such as taste receptor cells expressing inositol triphospate receptor type 3 (IP₃R3) never expressed CK14, suggesting that CK14 would be expressed only in immature cells. On the other hand, a small but distinct population of BrdU-positive cells still showed CK14 immunoreactivity in taste buds even at day 12, which might correspond to the cells that remain undifferentiated for a long period within taste buds.     

Characterization of regulatory mechanisms and states of human organic cation transporter 2

Polyspecific organic cation transporters (OCTs) have a large substrate binding pocket with different interaction domains. To determine whether OCT regulation is substrate specific, suitable fluorescent organic cations were selected by comparing their uptake in wild-type (WT) human embryonic kidney (HEK)-293 cells and in HEK-293 cells stably transfected with hOCT2. N-amidino-3,5-diamino-6-chloropyrazine-carboxamide (amiloride) and 4-[4-(dimethylamino)-styryl]-N-methylpyridinium (ASP) showed concentration-dependent uptake in hOCT2 at 37°C. After subtraction of unspecific uptake determined in WT at 37°C or in hOCT2 at 8°C saturable specific uptake of both substrates was measured. K_m values of hOCT2-mediated uptake of 95 µM amiloride and 24 µM ASP were calculated. Inhibition of amiloride and ASP uptake by several organic cations was also measured [IC₅₀ (in µM) for amiloride and ASP, respectively, tetraethylammonium (TEA) 98 and 30, cimetidine 14 and 26, and tetrapentylammonium (TPA) 7 and 2]. Amiloride and ASP uptake were significantly reduced by inhibition of Ca²⁺/CaM complex (–55 ± 5%, n = 10 and –63 ± 2%, n = 15, for amiloride and ASP, respectively) and stimulation of PKC (–54 ± 5%, n = 14, and –31 ± 6%, n = 26) and PKA (–16 ± 5%, n = 16, and –18 ± 4%, n = 40), and they were increased by inhibition of phosphatidylinositol 3-kinase (+28 ± 6%, n = 8, and +55 ± 17%, n = 16). Inhibition of Ca²⁺/CaM complex resulted in a significant decrease of V_max (160–99 photons/s) that can be explained in part by a reduction of the membrane-associated hOCT2 (–22 ± 6%, n = 9) as determined using FACScan flow cytometry. The data indicate that saturable transport by hOCT2 can be measured by the fluorescent substrates amiloride and ASP and that transport activity for both substrates is regulated similarly. Inhibition of the Ca²⁺/CaM complex causes changes in transport capacity via hOCT2 trafficking. organic cation transport; fluorescence measurement; 4-[4-(dimethylamino)-styryl]-n-methylpyridinium; amiloride     

Partitioning of [14C]sucrose and Acid Invertase Activity in Reproductive Organs of Pepper Plants in Relation to Their Abscission Under Heat Stress

The effect of heat stress on processes related to carbohydratepartitioning was investigated in young bell pepper (Capsicumannum L. cv. Maor) plants in relation to abscission of theirreproductive organs at different stages of development. None of the reproductive organs abscised after 5 d in a normalday/night temperature regime (25/18 °C). With a temperatureregime of 35 °C day, 25 °C night, abscission occurredin only a small portion of the flower buds and none of the flowersand fruitlets. However, when temperatures in the day and nightwere reversed (25/35 °C, day/night) all the buds and someof the flowers abscised during that time period. The young fruitat the first node did not abscise under any temperature regime.The abscission rate of the flower buds was reduced under heatstress if the developing fruit at the first node had been removed. High temperature during either the light or dark periods reducedthe export of [¹⁴C]sucrose from the source leaf (fed for 48h with [¹⁴C]sucrose). Both heat stress and fruit presence reduced the relative amountof [¹⁴C]sucrose which was exported to the flower buds, flowersand roots. Likewise, these treatments reduced the concentrationof reducing sugars in the reproductive organs. Concomitantly,the heat stress and fruit presence on the first node reducedthe activity of soluble acid invertase in the flower buds andthe roots, but not in young leaves. Overall, the results show that heat stress causes alternationin sucrose distribution in the plant, but may also have specificeffects on metabolic activities related to sucrose import andutilization in flower buds and flowers which in turn may enhancetheir abscission. Bell pepper, (Capsicum annuum L. cv. Maor), abscission, acid invertase, heat stress, reproductive organs, sink leaves     

An intensity/time study of the taste of amino acids

An intensity/time study of the taste of selected amino acidswas carried out. Intensity, persistence and total gustatoryresponse were assessed at five concentrations. Ten amino acidswere assessed for sweetness and eleven amino acids were assessedfor bitterness, four amino acids being assessed for both sweetnessand bitterness. Both a linear function and a power function,I = Kcⁿ (where I is taste intensity, c is concentration, K isa constant and n is the exponent of taste intensity), were fittedto the data. The accession efficiencies for taste recognitionand taste detection were found. Kinetic equations were usedto find K_m, the affinity of the receptor site for the sapidmolecule. Limited relationships between chemical structure ofthe amino acids and their temporal properties were found.     

Taste Disks are Induced in the Lingual Epithelium of Salamanders during Metamorphosis

Morphological changes of oral cavity during metamorphosis withspecial reference to the taste organ were examined in Ezo salamanders(Hynobius retardatus) and axolotls (Ambystoma mexicanum), andcompared with those in bullfrogs (Rana catesbeiana). The non-distensibletongue of salamanders changed the structure progressively duringmetamorphosis: a small area of the rostrum protruded and developedcaudally with recession of the flat area of the tongue. Theprotrusion that developed on the tongue had numerous papillae,as seen in the frog tongue. The apical region of the papillaeoccasionally had a cell mass similar to the taste disk of frogs(termed a taste disk-like cell mass). On the flat area of thetongue, the barrel-shaped taste buds of larval salamanders weretransformed into taste buds with a wider receptor area. Thebarrel-shaped taste buds decreased progressively during metamorphosis,while taste disk-like cell masses increased. Neuronal labelingwith an antibody to neuron-specific enolase and fluorescentcarbocyanine dye showed that the taste disk-like cell massesin metamorphosed salamanders were innervated by the glossopharyngealnerve (nerve IX). Nerve IX responded to taste stimulation aswell as mechanical stimulation applied to the rostral tongue.During metamorphosis the salamanders undergo transformationand rearrangement of taste organs on the tongue possibly asan adaptation to the terrestrial environment. Chem. Senses 22:535–545, 1997.     

Changes in number and morphology of fungiform taste buds in rat after transection of the chorda tympani or chordalingual nerve

In normal rats there is one taste bud on the apical surfaceof each fungiform papilla. These taste buds are innervated bythe chorda tympani proper nerve (CT). According to general consensus,after cutting the nerve the taste buds should disappear. Inthis study, performed on 24 rats divided in six groups, theCT nerve on the left side (singly denervated) and the combinedchorda-lingual (CT-L) nerve on the other side (doubly denervatedwere permanently interrupted. The animals were sacrificed after5, 10, 20, 35,60 and 100 days and their tongues serially sectionedfor light microscope examiation. Some papillae were examinedunder an electron microscope. The papillae were categorizedinto three groups: papillae with a normal looking taste bud,with an abnormal looking taste bud and without a taste bud.The results showed a substantial number of papillae with a normallooking taste bud present at all time intervals in all animals.More specifically, on the singly denervated side the proportionof normal looking taste buds stayed below 10% until day 60,when it increased to 15% and to 23% on day 100. The proportionof abnormal looking taste buds decreased from above 92% by day5 to 49% on day 100. The percentage of fungiform papillae withoutsigns of a taste bud was relatively low on the singly denervatedside at times (1, 5, 16, 29, 34 and 28%). On the doubly denervatedside fewer than than 4% normal looking taste buds were foundthroughout the time period. The proportion of abnormal lookingtaste buds decreased from {small tilde} 96% by day 5 to 35%on day 100. A significantly higher proportion of papillae withno taste bud was observed on this side from day 10 onwards.(1, 29, 32, 52, 60 and 63%). The reasons for the differencein tast bud number between the doubly and singly denervatedsides are unknown, but it is possible that collaterals fromother (non-gustatory) nerves have an ability, although limited,to induce and maintain fungiform taste buds. In other words,the capacity to induce taste bud formation is not limited exclusivelyto gustatory nerves.     

Epidermal Growth Factor Interacts with Indole-3-Acetic Acid and Promotes Coleoptile Growth

We used coleoptile sections of Avena sativa, Sorghum bicolor,and Zea mays seedlings to examine interactions between epidermalgrowth factor (EGF) and indole-3-acetic acid (IAA) that mayaffect plant growth and development. Our 24-h bioassays employedthree controls ranging in dilution from 10^–4 to 10^–8g ml^–1: (1) 50 mM potassium-phosphate buffer solution(pH=6.0), (2) bovine serum albumin, a nonspecific protein; and(3) IAA; plus two treatments: (1) mouse epidermal growth factor(EGF) ranging from 10^–6 to 10^–10gml^–1, and(2) EGF + IAA. In all three species growth in IAA, EGF, andEGF + IAA treatments showed significant increases over controls;EGF+IAA showed significant increases in growth over IAA alone.As the concentrations of IAA decreased, the EGF and IAA interactionbecame more pronounced. At the highest IAA concentrations, EGF+ IAA increased growth rates ca. 2% to 39%, whereas at lowerIAA concentrations EGF + IAA promoted growth as much as 121%,thereby lowering the normal IAA physiological set point up tothree or four orders of magnitude. Our data suggest that aninteraction between EGF and IAA may allow plants to recognizeand respond to animal biochemical messengers, resulting in changesin plant cell elongation that ultimately may alter plant growthpatterns. (Received April 27, 1994; Accepted September 5, 1994)     

Selective interactions of lectins with amino acid taste receptor sites in the channel catfish

Five lectins of varying carbohydrate specificities, Dolichosbiflorus (DBA), jacalin, Phaseolus vulgaris (PHA), Pisum sativum(PSA) and Ricinus communis (RCA I), were used to extend characterizationof the glycoprotein nature of taste plasma membranes and todifferentially affect the binding of two taste stimuli, L-alanineand L-arginine, to their respective taste receptor sites inthe cutaneous taste system of the channel catfish (Ictaluruspunctatus). The binding of the taste stimulus L-arginine toa partial membrane fraction (P_–) from taste epitheliumwas inhibited by 68 and 74% by preincubation in the presenceof the unconjugated lectins PHA and RCA I respectively. A correspondinglevel of inhibition of L-alanine binding was seen in the presenceof RCA I (76%); however, PHA had little effect upon L-alaninebinding. DBA appeared to selectively inhibit L-alanine but notL-arginine binding (60 versus 8% respectively) while jacalinmoderately inhibited the binding of both stimuli to fractionP2. PSA had little effect upon the binding of either L-alanineor L-arginine (4 and 5% inhibition respectively). Inhibitionof taste receptor binding by all lectins was time- and dose-dependent,and was fully abolished by incubation in the presence of theappropriate hapten sugar. The biotinylated lectins DBA, jacalin,PHA, RCA I and concanavalin A (Con A) were used to identifythe glycoprotein components of the chemosensory plasma membranesafter polyacrylamide gel electrophoresis. As previously shown,numerous protein components were labeled by Con A. In contrast,only a few minor protein components were labeled by PHA, DBAand RCA I. This differential labeling of the taste membranesand the differential inhibition of receptor binding by lectinssuggest that they may prove useful as tools in the isolationand purification of taste receptor proteins.     

Cytogenetic Studies in Cochlearia L. The chromosomal homogeneity within both the 2n = 12 diploids and the 2n = 14 diploids and the cytogenetic relationship between the two chromosome levels

A cytogenetic study has been carried out within the diploidCochleariae. This has demonstrated that all the 2n = 12 taxaare chromosomally homogeneous, as are those with 2n = 14. Itis therefore suggested that the 2n = 12 group be recognisedas C pyrenaica DC, with the morphologically distinct specimensfrom France and Spain at present regarded as C. aestuaria (Lloyd)Heywood being given subspecific rank in C. pyrenaica. It isalso suggested that all the 2n = 14 taxa be regarded as C groenlandicaL. Some doubt is expressed on the existence of C. scotica Druceas a species, as all the specimens of this taxon examined provedto be tetraploid with 2n = 24. Genomic analysis has shown thatthe 2n = 14 group is a primary tetrasomic derivative of the2n = 12 group.     

Solitary chemoreceptor cells of Ciliata mustela (Gadidae, Teleostei) are tuned to mucoid stimuli

Summed potentials were recorded from the dorsal recurrent facialnerve innervating the solitary chemoreceptor cells on the anteriordorsal fin (ADF), from the ventral recurrent facial nerve innervatingboth taste buds and solitary chemoreceptor cells on the pectoral(PEC) and pelvic (PEL) fins, and from the anterior dorsal finmuscles in the rockling, Ciliata mustela. There is little overlapbetween the sumulus spectra of solitary chemoreceptor cellsand taste buds. The ADF solitary cells are particularly sensitiveto body mucus (skin water) of non-congeners like Gadus, Solea,Cottus, Mugil, Zoarces, Gaidropsarus, and Encheliopus, but insensitiveto amino acids and a variety of body fluids of fish, invertebrates,and extracts of potential stimuli like algae and sand. Pectoraland pelvic fins are particularly sensitive to amino acids, bodyfluids of fish and invertebrates, but less sensitive to skinmucus of fish, probably due to the abundance of taste buds.Active sampling by undulation of the anterior dorsal fin isessential for proper functioning; it induces disadaptation ofthe receptor elements. Solitary chemoreceptor cells provide,apparently, cues to discriminate between conspecifics and non-conspecifics.It is unlikely that they are involved in pheromone detection.     

Initial Anatomical Changes Associated with Tuber Formation on Single-node Potato (Solanum tuberosum L.) Cuttings: A Re-evaluation

Cell division and cell expansion during early stages of tuberdevelopment were studied using developing axillary buds on single-leafcuttings from potato (Solanum tuberosum L.). Cuttings takenfrom plants induced to form tubers, by short day (SD) treatment,were compared with cuttings from non-induced (long day, LD)plants. In the apical zone of the buds, cell division occurredfrom the first day after cutting, in both LD and SD cuttings.The planes of these divisions were transverse, associated withelongation of the buds. At day 5, a new orientation of celldivision was observed in the subapical zone of SD cuttings only.These divisions were longitudinal, associated with radial growth.Cell expansion occurred in both SD and LD cuttings, and wasnot uniquely related to the onset of tuber formation. Copyright1999 Annals of Botany Company Solanum tuberosum L., potato, tuber formation, cell division, cell expansion.     

The Scaling of Plant Height: A Comparison Among Major Plant Clades and Anatomical Grades

The scaling plant height h (m) with respect to stem diameterd (m) was determined for a total 610 species (mosses, n = 40;pteridophytes, n = 16; dicotyledonous herbs, n = 117; palms,n = 17; gymnosperms, n = 105; dicotyledonous trees, n = 315);axial length or mass vs. d was determined for the pteridophytePsilotum nudum ; and the scaling of critical buckling heighth_crit of gymnosperm and dicotyledonous trees was calculatedbased on the record trunk d and average Young's modulus E anddensity p of 33 wood species. The scaling exponents (based onleast squares and reduced major axis regressions;     

Expression and characterization of delayed rectifying K+ channels in anterior rat taste buds

Delayed rectifying K⁺ (DRK) channels in taste cells have been implicated in the regulation of cell excitability and as potential targets for direct and indirect modulation by taste stimuli. In the present study, we have used patch-clamp recording to determine the biophysical properties and pharmacological sensitivity of DRK channels in isolated rat fungiform taste buds. Molecular biological assays at the taste bud and single-cell levels are consistent with the interpretation that taste cells express a variety of DRK channels, including members from each of the three major subfamilies: KCNA, KCNB, and KCNC. Real-time PCR assays were used to quantify expression of the nine DRK channel subtypes. While taste cells express a number of DRK channels, the electrophysiological and molecular biological assays indicate that the Shaker Kv1.5 channel (KCNA5) is the major functional DRK channel expressed in the anterior rat tongue. transduction