Amiloride Effects on Taste Quality: Comparison of Single and Multiple Response Category Procedures

Although there is compelling evidence that amiloride reducesthe intensity of Na⁺ and Li⁺ salts in humans, its effects onsaltiness are conflicting. Many salts elicit not only a saltytaste but also one or more side tastes (sweetness, sournessor bitterness). Some studies have shown a suppression of saltinessby amiloride; others show no effect on saltiness but a significantreduction in sourness. In the experiments demonstrating a reductionof saltiness, subjects estimated only saltiness; in those showingan amiloride effect on sourness and not saltiness, subjectsestimated all qualities on each trial. The present study examinesthe role of the psychophysical method in these conflicting results.We have investigated the effects of amiloride on taste qualityby modifying only the instructions to the subjects, keepingall other variables constant. One group of subjects (intensity-only)gave magnitude estimates of the overall intensity of a LiCIconcentration series. A second group (salty-only) was instructedto estimate the saltiness of the stimuli, and a third group(sour-only) estimated their sourness. Finally, a fourth group(profile) rated all of the taste qualities on each stimuluspresentation, using a modified taste profile method. The ratingsof all groups were made comparable by the use of 0.1 mM quinine-HCIas a modulus. When subjects used only one response category,amiloride reduced their estimates (of intensity, saltiness orsourness), but if subjects attended to all four qualities, amiloridespecifically reduced the sourness of LJCI and had no significanteffect on its saltiness. Comparison of the saltiness estimatesof the salty-only group to the sum of the salty and sour estimatesof the profile group demonstrated that subjects combined thesesensations when presented with only one response alternative.To reveal the effect of amiloride on a specific quality of asalt, the psychophysical method must allow subjects to attendto all qualities on each trial. These data and previous resultssuggest that apical Na⁺ channels on the taste receptor cellmembrane mediate the sourness but not the saltiness of Na⁺ andLi⁺ salts. Chem. Senses 22: 267–275, 1997.     

Suppression of Bitterness by Sodium: Variation Among Bitter Taste Stimuli

Taste interactions between salts (NaCl, LiCl, KCl, L-arginine:L-asparticacid, Na-acetate and Na-gluconate) and bittertasting compounds(urea, quinine HCI, magnesium sulphate, KCI, amiloride HCI andcaffeine) were investigated. In each study binary combinationsof three or four concentrations of one bitter compound withfour concentrations (0, 0.1, 0.3 and 0.5 M) of one salt wererated for bitterness and saltiness using the method of magnitudeestimation. In most cases, perceived bitterness was suppressedby salts, although the degree of suppression varied. In general,bitterness suppression was not accompanied by an equivalentreciprocal suppression of saltiness. Only MgSO₄ and amiloridehad suppressing effects on the saltiness of NaCl at the intermediateconcentrations and no bitter compound affected the saltinessat the high concentrations of NaCl. Since salt suppressed thebitterness of urea effectively, a detailed analysis of suppressionof the bitterness of urea by different salts was conducted.Those studies indicated that the key component in this effectwas the sodium or lithium ion for two reasons: first, all threesodium salts and the lithium salt had a suppressive effect onbitterness, whereas KCl did not; secondly, the effect of a salton suppression of the bitterness of urea was independent ofits perceived saltiness; that is, NaCl, Na-acetate (which isperceived as less salty than NaCl), and Na-gluconate (whichis perceived as less salty than Na-acetate) reduced bitternesscomparably. These results suggest that there is a major peripheralcomponent to the suppression of the bitterness of urea, andperhaps other bitter tasting compounds, by sodium. Chem. Senses20: 609–623, 1995.     

PERCEIVED SALTINESS OF TABLE SPREADS OF VARYING FAT COMPOSITIONS

Four types of spread with 80% fat, but varying in the origin of the fat (milk/vegetable) and in the proportion of oil versus solid fat (0–90% vegetable oil), were manufactured at NaCl concentrations of 1.0, 1.5 and 2.0%. Their saltiness and degree of melting in the mouth were rated using the time-intensity (TI) procedure. The perceived maximum saltiness of each NaCl concentration varied widely, samples with 80% milk fat + 20% vegetable oil being the most salty and those with 15% vegetable fat + 85% vegetable oil the least salty. The differences in saltiness were not directly related to the degree of melting in the mouth. The results demonstrate that extrapolating saltiness on the basis of NaCl contents over brands of products would be risky, since saltiness and NaCl contents are not necessarily correlated even in apparently similar products.     

SENSORY CHARACTERISTICS OF POTASSIUM LACTATE AND SODIUM CHLORIDE IN A MODEL SYSTEM

The objective of this study was to assess the sensory characteristics of potassium lactate (PL) in combination with salt (NaCl) in a model system at common usage levels for meat systems. Using a gelatin-based (3%) model system containing various concentrations of potassium lactate (0%, 1%, 2%, and 3%) and sodium chloride (0%, 0.5%, 1%, 1.5%, 2%, 2.5%, and 3%), a sensory panel (10) selected for salt sensitivity (0.08% threshold or less) scored the intensity of saltiness and bitterness of each treatment combination using a scale of 0 = none and 15 = intense. As salt concentration increased, saltiness increased and bitterness decreased; as PL increased, saltiness and bitterness increased. At 2% salt, the bitterness of PL was masked; however, salt concentrations below and above 2% were less effective in bitterness reduction.     

Salt as an oral irritant

The temporal characteristics of oral irritation produced byNaCl and KCl were investigated in two experiments. Subjectswere exposed to 0.4 and 0.8 M solutions of both salts at therate of once per min for 15 min. No water rinses were allowedbetween sips. The results showed that the irritation producedby NaCl increased in intensity as a function of number of exposures.In contrast, the intensity of the salt taste remained approximatelyconstant over the 15 min period. The relative contribution ofirritation to the sensation of saltiness therefore increasedover time. The same pattern of results was found for 0.8 M butnot for 0.4 M KCl. The growth in perceived intensity of saltirritation with repeated exposures may be attributable to thephenomenon of ‘sensitization’ that is a characteristicof some nociceptive afferents. Overall, the results demonstratethat salts produce perceptible sensations of irritation at concentrationstypically used in psychophysical studies of salt taste, andthat this apparently non-gustatory component of saltiness vanesdirectly with both concentration and duration of exposure.     

The Perception of Saltiness is Eliminated by NaCl Adaptation: Implications for Gustatory Transduction and Coding

The tastes of salts to humans are complex. NaCl is the mostpurely salty of all salts, but even this stimulus tastes sweetat low concentrations and somewhat sour at mid-range intensities.Other salts taste significantly sour or bitter in addition tosalty. Previous studies have shown that the saltiness of simplehalide salts is reduced by adaptation to NaCl, suggesting thata single mechanism might be responsible for the salty tasteof these stimuli. In electrophysiological studies in rodents,the response to NaCl is reduced by application to the tongueof the Na⁺- channel blocker amiloride. Organic Na⁺ salts aremore heavily dependent on this amiloride-sensitive transductioncomponent than NaCl, and are generally less salty and more sour.In order to investigate the relationship between NaCl saltinessand that evoked by other salts, we adapted the tongue to distilledH₂O and to 0.1 M NaCl and obtained direct magnitude estimatesof the taste intensity of 15 organic and inorganic Na⁺, Li⁺,K⁺ and Ca²⁺ salts, matched for total intensity. Subjects dividedthese magnitude estimates among the component taste qualities.Adaptation to NaCl abolished the taste of NaCl and LiCl, andeliminated the saltiness of all other salts. The magnitude estimatesof the bitterness and sourness of many salts increased afterNaCI adaptation. Since recent biophysical data suggest thatadaptation in taste receptors may involve whole-cell mechanisms,we propose that saltiness is reduced by NaCl adaptation becauseit originates in the subset of taste receptors responsive toNaCl. This implies that saltiness is coded within the CNS incells whose receptive fields include the NaCl-sensitive receptorcells and that the degree to which any salt tastes salty isdetermined by its ability to drive these receptors. This modelproposes, for example, that KCl has a salty component becauseit stimulates some of the same receptor cells as NaCl, eventhough the transduction mechanisms for KCl are different thanthose engaged by NaCl. Adaptation to NaCl blocks the saltinessof KCl and other salts because they stimulate NaCl-sensitivereceptor cells. Chem. Senses 20: 545–557, 1995.     

Electrophysiological studies of salty taste modification by organic acids in the labellar taste cell of the blowfly.

Using the labellar salt receptor cells of the blowfly, Phormia regina, we electrophysiologically showed that the response to NaCl and KCl aqueous solutions was enhanced and depressed by acetic, succinic and citric acids. The organic acid concentrations at which the most enhanced salt response (MESR) was obtained were found to be different: 0.05-1 mM citric acid, 0.5-2 mM succinic acid and 5-50 mM acetic acid. Moreover, the degree of the salt response was not always dependent on the pH values of the stimulating solutions. The salt response was also enhanced by HCl (pH 3.5-3.0) only when the NaCl concentration was greater than the threshold, indicating that the salty taste would be enhanced by the comparatively lower concentrations of hydrogen ions. Another explanation for the enhancement is that the salty taste may also be enhanced by undissociated molecules of the organic acids, because the MESRs were obtained at the pH values lower than the pKa(1) or pKa(2) values of these organic acids. On the other hand, the salty taste could be depressed by both the lower pH range (pH 2.5-2.0) and the dissociated organic anions from organic acid molecules with at least two carboxyl groups.     

THE EFFECT OF SALT REDUCTION ON TASTE PLEASANTNESS IN COOKED'BOLOGNA-TYPE'SAUSAGES

This study's objective was to study how much the salt (NaCl) content of cooked sausage can be reduced without violating the perceived taste pleasantness. The 34 assessors evaluated seven cooked sausages made with added salt concentrations of 1.05; 1.20; 1.35; 1.50; 1.65; 1.80 and 1.95%. A relative-to-standard scale was used for rating the saltiness and taste pleasantness. The saltiness and pleasantness intensity of different salt concentrations was rated against a reference sausage, which contained 1.5% added salt. A reference sample was also hidden among the samples. The assessors were able to rank sausages based on the saltiness into the right order. Based on taste pleasantness, there was no significant difference between the sausages made with 1.35; 1.50; 1.65; 1.80 or 1.95% added salt (p>0.05). Several differences were, however, detected among the saltiness levels. The results of this study suggest that it might be possible to reduce the salt content of cooked sausage to 1.35% added salt.     

A variational approach to behavioral and neuroelectrical laws

Variational methods play a fundamental and unifying role in several fields of physics, chemistry, engineering, economics, and biology, as they allow one to derive the behavior of a system as a consequence of an optimality principle. A possible application of these methods to a model of perception is given by considering a psychophysical law as the solution of an Euler-Lagrange equation. A general class of Lagrangians is identified by requiring the measurability of prothetic continua on interval scales. The associated Hamiltonian (the energy of the process) is tentatively connected with neurophysiological aspects. As an example of the suggested approach a particular choice of the Lagrangian, that is a sufficient condition to obtain classical psychophysical laws, while accounting for psychophysical adaptation and the stationarity of neuronal activity, is used to explore a possible relation between a behavioral law and a neuroelectrical ,response based on the Naka-Rushton model.     

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.     

LIKING OF POPCORN CONTAINING DIFFERENT LEVELS OF SALT1

Approximately 300 subjects participated in a test to determine whether the level of salt in popcorn affected the change in overall liking observed as a consequence of consuming a serving of popcorn. Subjects tasted and rated three samples of popcorn (low, medium, high salt levels), ate a serving of popcorn at one of the salt levels and then tasted and rated the three samples again. Subjects returned one or seven days later and tasted and rated the samples again. Overall liking, salt intensity and salt liking were measured. The subjects also completed a questionnaire about their liking and consumption of popcorn. Liking scores for the popcorns decreased after eating a 3-cup serving; however, sensory specific satiety for the different levels of salt was not observed. Consuming low salt popcorn increased the rated saltiness of the popcorns, whereas consuming high salt popcorn decreased the rated saltiness — probably a frequency effect. When subjects retasted the popcorns one or seven days later, all liking ratings increased except the salt liking ratings for the high salt sample. There was also a trend for the overall liking of this high salt sample to increase less than for the medium and low salt samples. Questionnaire measures of liking or consumption were not related to the changes in liking observed.     

Viscoelastic properties of an exopolysaccharide: Aeromonas gum, produced by Aeromonas nichidenii 5797

The viscoelastic properties of Aeromonas (A) gum in water were investigated by using the Rheometric Scientific ARES controlled strain rheometer. An intrinsic viscosity of 8336 ml/g was obtained according to the Fuoss-Straus equation. The effect of salt concentration on intrinsic viscosity revealed that the A gum exists as semiflexible chain. Typical shear-thinning (pseudoplastic) behavior was observed at concentrations higher than 0.52%. The zero shear viscosity (eta(0)) increased with increasing polysaccharide concentration (c) showing a gradient of approximately 1.0, 2.9 and 4.8 in different concentration domains. The critical concentrations c* and c**, at which the transitions from a dilute solution of independently moving chains to semidilute and then concentrated domains occurred, were determined roughly to be 1.2% and 3.5%. The results from dynamic experiments revealed that the A gum solution shows characteristics of polymer solutions without any evidence of gel-like character. All the results from steady and dynamic tests suggest that the A gum is a non-gelling polysaccharide. The temperature dependence of apparent viscosity was described by Arrhenius equation and the flow activation energy was estimated to be 45.2 kJ/mol, which is independent on polymer concentration.     

Flavor Characteristics of Glutathione in Raw and Cooked Foodstuffs

The flavor of glutathione (γ-l-glutamyl-l-cysteinylglycine, GSH) was examined by several sensory evaluations. The measurement of a point of subjective equality (PSE) showed that the peptide increases the flavor characteristics but did not affect the intensity of basic tastes, such as sweetness, saltiness, sourness, and umami. However, the threshold value of GSH decreased significantly in an umami solution containing 0.05% each of monosodium glutamate (MSG) and disodium inosinate (IMP). This suggests that GSH interacts with the umami substance and has a certain effect on the flavor.

GSH had a characteristic kokumi flavor, such as continuity, mouthfulness, and thickness in the umami solution as well as in a model beef extract constructed from analyzed components at a concentration of 0.02% w/v. Some foodstuffs, including meat, were found to contain GSH above its threshold value, which implicates the contribution of GSH to the flavor.

The thermal degradation study suggested that a part of GSH have changed into its disulfide, pyroglutamic acid (PCA), and cyclocysteinylglycine in cooked foodstuffs.     

Germination and reproductive biology in salty conditions of Asplenium marinum (Aspleniaceae), a European coastal fern

Although there are several species of ferns that are important components of many coastal ecosystems, the adaptations that allow these species to live in salty conditions have been studied only in two Acrostichum species. We have studied the effects of salt on germination, development of the gametophyte and sexual expression of the fern Asplenium marinum, which lives on coastal cliffs of Europe. Cultures at four different levels of salt concentration, 0%, 0.98%, 1.96% and 3.26%, were established for this purpose. Salt caused a significant delay in spore germination, and prevented it completely at high concentrations (3.26%). However, spores were able to recover germination ability after a period of seawater incubation once salinity concentrations decreased. Variable salinity conditions are a typical feature of sea cliffs where halophytes can grow. A salty culture medium also affects the development of the prothalli. Higher salinity of the substrate produced a significant decrease in gametophyte sizes, and affected the development of gametangia as well. Only males were found in a 0.98%-salt medium, and no gametangia were observed in cultures at 1.96%-salt medium. Recovered spores after one and a half month in high salinity concentration, 3.26%, were able, when transplanted to 0%-salt medium, to develop normal sexual prothalli, and this occurred in a shorter time than spores without any prior treatment. A. marinum can be considered as a halophyte fern that is able to germinate and grow in salty conditions. But it takes advantage of rainy periods that cause decrease of salt concentration, which seems to be important in particular for gametangia development and fertilization.     

Aggregation and fluorescence quenching of chlorophyll a of the light-harvesting complex II from spinach in vitro

The salt-induced aggregation of the light-harvesting complex (LHC) II isolated from spinach and its correlation with fluorescence quenching of chlorophyll a is reported. Two transitions with distinctly different properties were observed. One transition related to salt-induced fluorescence quenching takes place at low salt concentration and is dependent both on temperature and detergent concentration. This transition seems to be related to a change in the lateral microorganization of LHCII. The second transition occurs at higher salt concentration and involves aggregation. It is independent of temperature and of detergent at sub-cmc concentrations. During the latter transition the small LHCII sheets (approximately 100 nm in diameter) are stacked to form larger aggregates of approximately 3 microm diameter. Based on the comparison between the physical properties of the transition and theoretical models, direct and specific binding of cations can practically be ruled out as driving force for the aggregation. It seems that in vitro aggregation of LHCII is caused by a complex mixture of different effects such as dielectric and electrostatic properties of the solution and surface charges.     

Neohesperidin dihydrochalcone is not a taste enhancer in aqueous sucrose solutions

Neohesperidin dihydrochalcone (NHDC) is an intensive sweetener, obtained by alkaline hydrogenation of neohesperidin. In this investigation a supposed taste enhancing effect of this substance was tested. A three-step procedure was used. In the first experiment, using a pool of 31 subjects, NHDC and sucrose detection thresholds were measured. In the second experiment, psychophysical functions for both tastants were determined. Then, 15 participants closest to the group threshold who, in addition, had produced monotonic psychophysical taste functions were selected to participate in the next two experiments. In the third experiment, taste enhancement was tested. Three psychophysical sucrose functions were constructed, one with a near-threshold amount of NHDC added to each of seven sucrose concentrations, one with a near-threshold amount of sucrose added (control 1) and one without any addition (control 2). No difference was found between the NHDC-enriched sucrose function and the sucrose-enriched sucrose function. Finally, in experiment 4, differential threshold functions were constructed with either NHDC or sucrose added. Neither the overall shape of the functions nor a comparison of the points of subjective equality showed enhancement. It was concluded that weak NHDC does not enhance the taste of aqueous sucrose solutions.     

Donnan membrane equilibrium,sedimentation equilibrium,and coil expansion of DNA in salt solutions

This is a review of applications of the McMillan-Mayer-Hill virial theory and the ionic double-layer theory to dilute colloidal solutions, in particular, solutions of DNA. Interactions of highly charged colloidal rods are developed in terms of the second virial coefficients between two rods, and between one rod and one small co-ion. The relevant cluster integrals are evaluated with interaction potentials based on the Poisson-Boltzmann equation. The treatment is extended to the intrachain repulsion responsible for the statistical swelling of coiled DNA (excluded volume effect). The theory is compared with three sets of experimental data: The salt distribution in Donnan membrane equilibria of DNA-salt solutions, sedimentation equilibria of short DNA fragments at different ionic strengths, and the intrinsic viscosity of T7 DNA in NaCl solutions. In all cases the theory agrees well with the experiments. The agreement is not convincing for the sedimentation equilibrium at low ionic strength, because here the experimental DNA concentration is too high for the truncated dilute solution expansion of the DNA-salt repulsion.     

Critical change of ion conductance and self-sustained potential oscillation in artificial membrane systems

A change in conductance of an artificial membrane at a threshold concentration of environmental salt solution was interpreted in terms of a change in adsorptive reaction rate on the interface which was derived from an autocatalytic reaction model. The model also accounted for a self-sustained potential oscillation which was observed when the salt concentration of one side of the membrane was higher than the threshold mentioned above and that of the other side lower.     

Characterizing psychophysical measures of discrimination thresholds and the effects of concentration on discrimination learning in the moth Manduca sexta

What is the spatial and temporal nature of odor representations within primary olfactory networks at the threshold of an animal's ability to discriminate? Although this question is of central importance to olfactory neuroscience, it can only be answered in model systems where neural representations can be measured and discrimination thresholds between odors can be characterized. Here, we establish these thresholds for a panel of odors using a Pavlovian paradigm in the moth Manduca sexta. Moths were differentially conditioned to respond to one odor (CS+) but not another (CS-) using undiluted odorants to minimize salience-dependent learning effects. At 24 and 48 h postconditioning, moths were tested for the presence of a conditioned response (CR) with a blank, then the CS+ and CS- (pseudorandomly) across a 5-log step series of increasing concentration. Results identified discrimination thresholds and established that differential CRs to the CS+ and CS- increased with stimulus concentration. Next, 3 separate groups of moths were differentially conditioned at either one-log step below, at, or one log step above the identified discrimination threshold. At 24 and 48 h postconditioning, moths were tested sequentially with a blank, the concentration used for conditioning, and then undiluted odor. Conditioning at one log step below the discrimination threshold established a CR, indicating both stimulus detection and learning, but was insufficient to establish evidence of discrimination. Moths conditioned at the discrimination threshold were able to discriminate but only when stimulated with undiluted odors, indicating learning, but discrimination measures were hampered. When conditioned above the discrimination threshold, moths had no difficulty in discriminating. These results establish methods for psychophysical characterization of discrimination and indicate that differential conditioning at lowered concentrations biases threshold measures.