Sweetness intensity enhancement by pulsatile stimulation: effects of magnitude and quality of taste contrast

Upon stimulation with continuously alternating (pulsatile) taste concentrations, humans report higher average taste intensities than for continuous stimulation with the same average tastant concentration. We investigated the effect of the magnitude of concentration changes (concentration contrast) and the effect of taste quality changes (quality contrast) between alternating tastants on sweet taste enhancement. The perceived sweetness intensity increased with the magnitude of the sucrose concentration contrast: The pulsatile stimulus with the highest concentration difference (average sucrose concentration: 60 g/L) was rated as the sweetest in spite of the fact that the gross sucrose concentrations were identical over stimuli. Moreover, this stimulus was rated equally sweet as a continuous reference of 70 g/L sucrose. On alternation of sucrose with the qualitatively different citric acid, sweet taste enhancement remained at the level observed for alternation with water at citric acid concentration levels up to 3 times its detection threshold. Alternation of a sucrose solution with a citric acid solution at 9 times its threshold concentration, resulted in an attenuation of the pulsation-induced enhancement effect. Upon alternation of citric acid pulses at concentrations around the threshold with water intervals only, no taste enhancement was observed compared with continuous citric acid stimuli of the same net concentration. We propose that the magnitude of pulsation-induced taste enhancement is determined by the absolute rather than relative change of tastant concentration. This explains why 1) pulsation-induced sweet taste enhancement is determined by the magnitude of the sucrose pulse-interval contrast and 2) the alteration of citric acid with water does not enhance taste intensity at detection threshold level.     

Effects of cannabinoids (marijuana) on taste intensity and hedonic ratings and salivary flow of adults

Cannabinoids purportedly improve taste responsiveness and enhancethe sensory appeal of foods. These properties and a commonlycited oral drying effect were evaluated in a series of studieswith ‘light’ marijuana users. The first was a double-blind,placebo-controlled, acute oral dosing trial, involving an ageand gender stratified sample of 57 adults. An influence of routeof drug delivery was explored in another 11 individuals whowere administered a single dose orally, sublingually and viacigarette. To explore effects following chronic administration,six additional individuals were dosed twice per day for 3 daysorally and by rectal suppository. Taste intensity and hedonicresponses for sweet, sour, salty and bitter food stimuli weremonitored at baseline, 2, 4 and 6 hours post-dosing in the acutestudies, and daily in the chronic study. Stimulated saliva sampleswere collected at these same times. Salivary flow rate was significantlynegatively correlated with plasma drug levels, and reported‘high’ 2 and 4 h post-dosing. No effects of thedrug were observed on taste responses. Self-reported shiftsin taste responsiveness and hedonics may be related to alterationsof memory and cognition, rather than gustatory function.     

A molecular mechanism for high salt taste in Drosophila

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Activity in salt taste fibers: peripheral mechanism for mediating changes in salt intake

In order to study the role of peripheral taste sensitivity inmediating increases in salt intake of the rat, the effects ofsodium deprivation and adrenalectomy on chorda tympani nerveresponses to taste stimulation were determined. Sodium deprivationresulted in a reduction in whole nerve responsivity to suprathresholdNaCl concentrations requiring a 10-fold increase in concentrationto elicit the same neural signal of control preparations. Saltintake of sodium deprived rats was predicted by adjusting datain a 10-min intake test from control rats for the reduced neuralsignal and lower salivary sodium levels of sodium deprived rats.The whole nerve responses to LiCl and KCl, as well as to NaCl,were reduced after sodium deprivation and adrenalectomy. Themultifiber response of the chorda tympani is comprised of theindividual responses of NaCl sensitive N-best fibers and HCl/NaClsensitive H-best fibers. After sodium deprivation N-best fibers'responses to suprathreshold concentrations of NaCl were reduced;H-best fibers' responses were not affected by sodium deprivation.Future studies will determine the effect of KCl and other saltson responses of N-best and H-best fibers. Applying Beidler'sbiophysical model to the single fiber data suggests that sodiumdeprivation influences receptor mechanisms for NaCl of N-bestfibers and not H-best fibers. Because repeated NaCl stimulationresulted in increased chorda tympani responsivity to NaCl, wesuggest that sodium deprivation may alter the salt receptorsimply by disuse. Altered receptor sensitivity may be an adaptivemechanism to influence salt consumption by a shift in suprathresholdNaCl intensity.     

Contrast and range effects for category, magnitude and labeled magnitude scales in judgements of sweetness intensity

The labeled magnitude scale (LMS) is a verbally anchored quasi-logarithmically spaced response scale with properties similar to magnitude estimation. Three experiments examined whether the LMS showed context effects similar to those found with magnitude estimation and category scales. Two versions of the LMS were used, one anchored at the high end to the strongest imaginable sweetness and the other to the strongest imaginable oral sensation. In a simple contrast experiment, subjects judged the sweetness of a 10% sucrose fruit beverage in the context of a less sweet (5%) beverage or a more sweet (20%) beverage. Consistent with previous literature, the sweetness was judged more intense in the low context and less intense in the high context, for all scaling methods. In a second experiment, this effect persisted (although was smaller) when the contextual item preceded the to-be-rated item, a so-called 'reversed-pair' design. Once again, the effect was highly significant for all scaling methods. In a third experiment, a range effect was examined using wide and narrow ranges of concentration. Psychophysical functions were flatter in a wide context and steeper in a narrow context, consistent with previous observations on range-mapping bias. This result was obtained for all scales. In three common contextual effects, the labeled magnitude scale behaved similarly to other scaling procedures. Its application to comparisons across individuals may be limited if those individuals have different experiential contexts within which they make their judgements.     

Effects of amiloride on salt taste in humans

Amiloride reduces several responses of the taste system to NaCl.These effects are crucial support for the ion transport theoryof salt taste. Here, three experiments tested for the inhibitoryeffect of amiloride on salt taste in normal, whole-mouth tastingin humans. There was no evidence that amiloride reduces thesaltiness of NaCl when used as a pretreatment, mixed into solutionwith NaCl, or both. In a fourth experiment, pretreating thetongue with amiloride-soaked filter paper and presenting NaClon filter paper within the treated area also had no effect onsaltiness. In addition, amiloride, a bitter compound, did notsuppress the taste of sucrose, and caffeine did not enhancethe taste of NaCl. These results suggest the ion transport theoryof NaCl taste needs to be modified or extended to account fornormal whole-mouth tasting of NaCl in humans. ¹ Present address: 1605 Harbor Court Tower, 10 East Lee St.,Baltimore, MD 21202, USA     

Contributions of Smell and taste to overall intensity

The integration of olfaction and gustation in producing thesensation of flavor was studied using an almond extract stimuluspresented with the Two-Module Delivery System. Almond in themouth was found to influence the scaling of smell. The datasugest that this background effect is due to the diffusion ofairborne molecules from the oral cavity to the olfactory receptorsarea. At least under some stimulation conditons, almond vaporin the nose was shown to afect the scaling of taste. When scalingoverall intensity, subjects apparently added together the sensationsof smell and taste to produce the sensation of overall intensity.However, for all combinations of odorants and tastants, theestimate of overall intensity was less than the sum of the estimateof the smell and the estimate of the taste. The data furthersuggest that when estimating the overall intensity, the magnitudeassigned to the olfactory or gustatory sensations was less thanthat which have been assigned when estimating just smell ortaste.     

Effect of Maillard reacted peptides on human salt taste and the amiloride-insensitive salt taste receptor (TRPV1t)

Maillard reacted peptides (MRPs) were synthesized by conjugating a peptide fraction (1000-5000 Da) purified from soy protein hydrolyzate with galacturonic acid, glucosamine, xylose, fructose, or glucose. The effect of MRPs was investigated on human salt taste and on the chorda tympani (CT) taste nerve responses to NaCl in Sprague-Dawley rats, wild-type, and transient receptor potential vanilloid 1 (TRPV1) knockout mice. MRPs produced a biphasic effect on human salt taste perception and on the CT responses in rats and wild-type mice in the presence of NaCl + benzamil (Bz, a blocker of epithelial Na+ channels), enhancing the NaCl response at low concentrations and suppressing it at high concentrations. The effectiveness of MRPs as salt taste enhancers varied with the conjugated sugar moiety: galacturonic acid = glucosamine > xylose > fructose > glucose. The concentrations at which MRPs enhanced human salt taste were significantly lower than the concentrations of MRPs that produced increase in the NaCl CT response. Elevated temperature, resiniferatoxin, capsaicin, and ethanol produced additive effects on the NaCl CT responses in the presence of MRPs. Elevated temperature and ethanol also enhanced human salt taste perception. N-(3-methoxyphenyl)-4-chlorocinnamid (a blocker of TRPV1t) inhibited the Bz-insensitive NaCl CT responses in the absence and presence of MRPs. TRPV1 knockout mice demonstrated no Bz-insensitive NaCl CT response in the absence or presence of MRPs. The results suggest that MRPs modulate human salt taste and the NaCl + Bz CT responses by interacting with TRPV1t.     

Tactile interaction with taste localization: influence of gustatory quality and intensity

Taste is always accompanied by tactile stimulation, but little is known about how touch interacts with taste. One exception is evidence that taste can be "referred" to nearby tactile stimulation. It was recently found (Lim J, and Green BG. 2007. The psychophysical relationship between bitter taste and burning sensation: evidence of qualitative similarity. Chem Senses. 32:31-39) that spatial discrimination of taste was poorer for bitterness than for other tastes when the perceived intensities were matched. We hypothesized that this difference may have been caused by greater referral of bitterness by touch. The present study tested this hypothesis by comparing localization of quinine sulfate and sucrose under conditions that minimized and maximized the opportunity for referral. In both conditions, stimulation was produced by 5 cotton swabs spaced 1 cm apart and arranged in an arc to enable simultaneous contact with the front edge of the tongue. Only one swab contained the taste stimulus, whereas the rest were saturated with deionized water. In both conditions, the swabs were stroked up-and-down against the tongue 5 times. Subjects were asked to identify which swab contained the taste stimulus 1) 5 s after the fifth stroke (touch-removed condition) and 2) immediately at the end of the fifth stroke, with the swabs still in contact with the tongue (touch-maintained condition). Ratings of taste intensity were obtained to assess the possible effect of perceived intensity on spatial localization. Taste localization was surprisingly accurate, especially for sucrose, with errors of localization in the range of 1 cm or less. For both stimuli, localization tended to be poorer when the tactile stimulus was present while subjects made their judgments, but the difference between conditions was significant only for the lower concentration of quinine. The results are discussed in terms of both the surprisingly good spatial acuity of taste and the possibility of having a close perceptual relationship between touch and bitter taste.     

The taste of sodium chloride: masking and adaptation

Human subjects estimated the saltiness of sodium chloride anda mixture of sodium chloride and sucrose. The adapting effectsof sucrose, sodium chloride and a mixture of both substanceswere investigated. The stimuli were delivered by a flow systemto pre-defined tongue areas. After adaptation to NaCl and tothe mixture the subjective intensity of NaCl-saltiness decreased58% and 69% respectively. Also a strong masking effect of saltinesswas observed: 59.1% reduction. No indications of crossadaptationwere found. The results were interpreted in favour of the independencyof adaptation and masking.     

A model for the stimulation of taste receptor cells by salt.

A taste cell mucosal surface is regarded as a planar region containing bound anionic sites and openings to ionic channels. It is assumed that the bulk aqueous properties of the exterior phase are not continuous with the surface but terminate at a plane near the surface. The region between the (Stern) plane and the membrane is regarded as having a lower dielectric constant than bulk water. This fact admits the possibility of ion pair formation between fixed sites and mobile cations. Mobile ion pairs entering the region may also bind to a fixed anionic site. Thus, it is assumed that mobile cations and ion pairs are potential determining species at the surface. Binding cations neutralizes surface charges, whereas binding mobile ion pairs does not. This competition accounts for the observed anion effect on stimulation of tast receptors by sodium salts. The potential profile is constructed by superimposing the phase boundary potentials with an ionic diffusion potential across the membrane. The model accounts for the anion effect on receptor potential, pH effects, the reversal of polarity when cells are treated with FeCl3, and the so-called "water reponse," depolarization of the taste cell upon dilution of the stimulant solution below a critical lower limit. The proposed model does not require both bound cationic and anionic receptors, and further suggests that limited access to a Stern-like region continuous with membrane channels may generally serve to control transport of ions.     

Dissociating pleasantness and intensity with quinine sulfate/sucrose mixtures in taste

Independent experimental manipulation of subjective intensity and hedonic tone is required if one wants to study their separate effects on brain activity and behavior. This is problematic because hedonic tone and subjective intensity are related, leading to a pleasantness change each time the stimulus intensity is altered. In the present study, a solution to this problem was explored by combining a pleasant-tasting substance (sucrose) and a bad-tasting substance (quinine sulfate) into a number of different isointense mixtures. Here we show that subjective intensity as well as pleasantness can be accurately predicted, particularly in midrange, only if one corrects for mixture suppression.     

The allen correction for porphyrin estimation in spectrofluorimetry

• 1.1. The Alien correction as used in spectrophotometry is modified to be used as a general method for spectrofluorimetric estimation of porphyrins.

     

Running to the taste of salt in mineralocorticoid-treated rats

In the present series of studies, the running speed of rats in a runway was used to measure appetitive motivation for different concentrations of salt solutions. We found that when rats were thirsty, they were not interested in running for concentrated salt solutions; when they were rendered salt hungry by mineralocorticoid treatment in addition to the thirst, or even without thirst, they ran vigorously for salty tasting solutions, as high as 24% NaCl. The running speed is correlated with the degree of the mineralocorticoid treatment and the salt concentration. Moreover, drinking 5 cc of a 3% NaCl solution before the runway test decreases the running speed of the mineralocorticoid-treated rat. The experiments are discussed in the context of the effects of hormonal events on the movitation to ingest salt solutions.     

Probabilistic segmentation and intensity estimation for microarray images

We describe a probabilistic approach to simultaneous image segmentation and intensity estimation for complementary DNA microarray experiments. The approach overcomes several limitations of existing methods. In particular, it (a) uses a flexible Markov random field approach to segmentation that allows for a wider range of spot shapes than existing methods, including relatively common 'doughnut-shaped' spots; (b) models the image directly as background plus hybridization intensity, and estimates the two quantities simultaneously, avoiding the common logical error that estimates of foreground may be less than those of the corresponding background if the two are estimated separately; and (c) uses a probabilistic modeling approach to simultaneously perform segmentation and intensity estimation, and to compute spot quality measures. We describe two approaches to parameter estimation: a fast algorithm, based on the expectation-maximization and the iterated conditional modes algorithms, and a fully Bayesian framework. These approaches produce comparable results, and both appear to offer some advantages over other methods. We use an HIV experiment to compare our approach to two commercial software products: Spot and Arrayvision.