Synthesis of tastes other than the 'primaries': implications for neural coding theories and the concept of 'suppression'

It has been demonstrated that mixtures of ‘primary’tastes may be perceived as singular rather than as multiple(Erickson and Covey, 1980). This finding may be accounted forin two different and testable ways. First, suppression: oneof the tastes in the mixture may be suppressed by the other(i.e., in a mixture of A and B, A suppresses the taste of B);the prediction which follows from this hypothesis is that asolution of A (the suppressing component) should be indistinguishablefrom the AB mixture. Secondly, synthesis: the singularity ofthe mixture may arise from the formation of a new taste, andin this case both components should be distinguishable fromtheir mixture. The present data indicate that the latter istrue in some cases. Since the taste of these mixtures must beother than the primary tastes used to form them, these findingsquestion the presumption that only four ‘primaries’are adequate to describe the range of tastes.     

An assessment of binary mixture interactions for nine sweeteners

Binary mixtures of acesulfame K, aspartame, sodium cyclamate,fructose, glucose, stevioside, sodium saccharin, sucrose andxylitol were assessed using factorial mixture designs. A simpleadditive model was used to generate predictions for the sweetnessof the mixtures and these predicted responses were comparedto the observed sweetness ratings of the mixtures. It was foundthat the mixtures tended to exhibit superadditivity at low concentrations,additivity at intermediate concentrations and subadditivityat high concentrations. Synergistic and suppressive effectsin the mixtures were evaluated by comparing mixture responsesto the sweetness ratings of ‘self-mixtures’. Self-mixturedata were generated by treating a mixture of a substance withitself as if it were a mixture of two different substances.Synergism was defined as a mixture response that was greaterthan the sweetness of the component self-mixtures, and suppressionwas defined as a mixture response that was less than the sweetnessof the component self-mixtures. Of the 31 binary mixtures studied,18 showed synergism, two showed suppression and 11 did not differsignificantly from their components. It is hypothesized thatmultiple sweetness receptors or release from bitter suppressionmay account for the synergistic effects.     

Inhibitory receptor binding events among the components of complex mixtures contribute to mixture suppression in responses of olfactory receptor neurons of spiny lobsters

Responses of olfactory receptor neurons of spiny lobsters Panulirus argus to two-component mixtures can be shaped by inhibitory events such as odor-activated hyperpolarizations and inhibition of odor-receptor binding (Daniel et al. 1996). In the current study, we extend this analysis to complex mixtures by examining responses of spiny lobster olfactory receptor neurons to mixtures containing up to seven odorants, consisting of adenosine-5′-monophosphate, ammonium, betaine, l-cysteine, l-glutamate, dl-succinate, and taurine. The response to a mixture was often less than the response to its most excitatory component. The effect of adding an excitatory odorant to a mixture depended on olfactory receptor neuron type, composition of the mixture, and which compound was added. In some cases the added excitatory compound had no effect or even decreased the mixture's response intensity, thus demonstrating nonlinear contributions of the components. Response intensities predicted by a noncompetitive model, which is most representative of these olfactory receptor neurons, were improved when the model included a term for empirical measurements of inhibitory binding interactions, suggesting that inhibitory binding interactions are one mechanism contributing to mixture suppression. This model's predictions were accurate for binary mixtures but not for larger mixtures, suggesting that additional inhibitory mechanisms are needed to account for mixture interactions in complex mixtures. Accepted: 24 July 1998     

Interactions in caffeine-sucrose and coffee-sucrose mixtures: evidence of taste and flavor suppression

Taste–taste and flavor–taste interactions (suppression)in caffeine–sucrose and coffee–sucrose mixtureswere determined. Similar interactions for both types of mixturesshowed an extended hypoadditivity effect for overall taste orflavor intensity (percentage of suppression about 30–40%).Futhermore, mutual suppression among the components has beendetermined. Firstly, the physical intensity of the suppressivecomponent controls the amount of suppression of the other component.Thus, the suppression of bitterness and coffee flavor qualitiesincrease when sucrose levels increase, and similarly, the suppressionof sweetness increases when caffeine or coffee levels rise.Secondly, the magnitude of suppression depends upon the qualityof the suppressive component. Comparisons of the reciprocalactions were made at similar subjective intensities of the mixture'sconstituents in isolation. The results showed that, at similarperceived intensities, caffeine bitterness or coffee flavorwere suppressed by sucrose but sweetness was not affected bycaffeine or coffee.     

The role of energy substrates in the regulation of force-frequency relationship in the rat myocardium: the influence of ambiocor

The effect of ambiocor (15 mg/100 ml), which contains natural substrates of energy metabolism, on the contractility of papillary muscle of the right ventricle of rat heart was studied at stimulation frequencies from 0.1 to 3.0 Hz at a temperature of 30 ± 1°C (n = 7). The effect was recorded 20 min after the addition of the preparation. It was demonstrated that ambiocor causes a significant (about 70%), independent of stimulation frequency, suppression of the amplitude of isometric contractions (negative inotropic effect), which is coupled with an increase in the relative value of the rest potentiation effect (a qualitative index of calcium content in SR). The influence of the mixture leads to significant alterations in the time parameters of the “contraction-relaxation” cycle: an increase in the duration of latent period; and a decrease in the time to peak tension and half-relaxation time. The effect of the mixture is partially reversible. During washing off the preparation with the control solution, the qualitative indicators of the contractile activity of papillary muscle are substantially improved in comparison with the initial ones. The character of alterations allows one to assume that the effect of ambiocor in the papillary muscle of the rat heart is realized partly through the suppression of the activity of sarcolemmal calcium channels.     

Increased Productivity of a Cover Crop Mixture Is Not Associated with Enhanced Agroecosystem Services

Cover crops provide a variety of important agroecological services within cropping systems. Typically these crops are grown as monocultures or simple graminoid-legume bicultures; however, ecological theory and empirical evidence suggest that agroecosystem services could be enhanced by growing cover crops in species-rich mixtures. We examined cover crop productivity, weed suppression, stability, and carryover effects to a subsequent cash crop in an experiment involving a five-species annual cover crop mixture and the component species grown as monocultures in SE New Hampshire, USA in 2011 and 2012. The mean land equivalent ratio (LER) for the mixture exceeded 1.0 in both years, indicating that the mixture over-yielded relative to the monocultures. Despite the apparent over-yielding in the mixture, we observed no enhancement in weed suppression, biomass stability, or productivity of a subsequent oat (Avena sativa L.) cash crop when compared to the best monoculture component crop. These data are some of the first to include application of the LER to an analysis of a cover crop mixture and contribute to the growing literature on the agroecological effects of cover crop diversity in cropping systems.     

Electro-olfactogram and multiunit olfactory receptor responses to binary and trinary mixtures of amino acids in the channel catfish, Ictalurus punctatus

In vivo electrophysiological recordings from populations of olfactory receptor neurons in the channel catfish, Ictalurus punctatus, clearly showed that responses to binary and trinary mixtures of amino acids were predictable with knowledge obtained from previous cross-adaptation studies of the relative independence of the respective binding sites of the component stimuli. All component stimuli, from which equal aliquots were drawn to form the mixtures, were adjusted in concentration to provide for approximately equal response magnitudes. The magnitude of the response to a mixture whose component amino acids showed significant cross-reactivity was equivalent to the response to any single component used to form that mixture. A mixture whose component amino acids showed minimal cross-adaptation produced a significantly larger relative response than a mixture whose components exhibited considerable cross-reactivity. This larger response approached the sum of the responses to the individual component amino acids tested at the resulting concentrations in the mixture, even though olfactory receptor dose-response functions for amino acids in this species are characterized by extreme sensory compression (i.e., successive concentration increments produce progressively smaller physiological responses). Thus, the present study indicates that the response to sensory stimulation of olfactory receptor sites is more enhanced by the activation of different receptor site types than by stimulus interaction at a single site type.     

Taste Responses to Binary Mixtures of Amino Acids in the sea catfish, Arius felis

In vivo electrophysiological recordings in the sea catfish,Arius felis, showed that the magnitude of the integrated facialtaste responses to binary mixtures of amino acids was predictablewith knowledge obtained from previous cross-adaptation studiesof the relative independence of the respective binding sitesof the component stimuli. Each component from which equal aliquotswere drawn to form the mixtures was adjusted in concentrationto provide for approximately equal reponse magnitudes. The magnitudeof the taste responses to binary mixtures whose component aminoacids showed minimal cross-adaptation was significantly greaterthan that to binary mixtures whose components exhibited considerablecross-reactivity. There was no evidence for mixture suppression.The relative magnitude of the taste responses in the sea catfishto stimulus mixtures is similar to that previously reportedfor olfactory receptor responses in the freshwater channel catfishand chorda tympani taste responses in the hamster. Chem. Senses21: 45–53, 1996.     

Associative growth studies in three-strain mixtures of lactic streptococci

A recently developed differential agar medium was used to study associative growth patterns in 17 different heterologous, three-strain mixtures of Streptococcus lactis, S. cremoris, and S. diacetilactis grown in milk. Mixtures were made by combining equal volumes of 18-hr milk cultures of the three species. Relative populations of component species were followed through three successive transfers in milk after the initial mixed propagation. Direct evidence for strain dominance and compatibility was obtained. A procedure also was developed to estimate the extent of suppression of S. lactis and S. diacetilactis in a mixture containing a dominant S. cremoris strain. The technique described could be successfully applied in quality-control work in the dairy-starter manufacturing industry.     

Theoretical note: tests of synergy in sweetener mixtures

Some methods for examining the additivity of sweeteners, and their synergy in mixtures depend upon setting component concentrations on the basis of sweetness equivalence, usually to a sucrose reference. These methods may under- or over-predict the sweetness of a mixture, leading to spurious claims of synergy or mixture suppression. This paper points out one problem with one such popular method, in that the method can lead to a conclusion that a substance would synergize with itself. To the extent that self-synergy is an illogical conclusion for a mixture comparison, such a method should be avoided in tests of synergy. The sweetness equivalence approach is contrasted with a simpler approach based on concentrations that does not lead to conclusions of self- synergy.      

Intensity and hedonic judgments of taste mixtures: an information integration analysis

Anderson's (1981) information integration approach was usedto examine taste mixture integration for intensity and hedonicjudgments of sucrose/sodium chloride and sucrose/citnc acidsolutions. In Experiment I, total intensity and hedonic ratingswere made for factorial combinations of sucrose and sodium chlorideor citric acid. The total intensity judgments produced an integrationpattern characterized by extreme subadditivity at high soluteconcentrations. (Subadditivity refers to the tendency for totalmixture intensity to be rated as less than the sum of the unmixedcomponent intensities.) The intensity judgment integration patternswere essentially identical for the two mixture types. However,the pattern of integration for the sucrose/sodium chloride andsucrose/citric acid mixtures differed for the hedonic ratings.Sucrose tended to eliminate the unpleasantness associated withincreasing concentrations of citric acid, while it only modulatedthis trend for sodium chloride. In Experiment II, subjects ratedthe individual sweet, salty and sour components of the mixturesto determine whether mixture suppression of the component tastescould account for the subadditivity of the total intensity judgmentsand/or the pattern of results for the hedonic ratings. It wasfound that sucrose suppressed the sour component of the sucrose/citricacid mixtures more than the salty component of the sucrose/sodiumchloride solutions This difference in component suppressionseemed to account for the hedonic integration patterns of thetwo mixture types which suggests that mixture suppression isan important factor to consider when predicting the pleasantnessof simple taste mixtures.     

Decomposition of spruce litter needles of different quality by Setulipes androsaceus and Thysanophora penicillioides

Different components of functional biodiversity, such as functional type richness and composition, have been reported to affect the decomposition of litter mixtures. In spite of the numerous reports of these effects, mechanisms underlying patterns of decomposition in litter mixtures are still unclear. We analyzed whether mixture decomposition was affected by: (a) the number of species in the mixture (mixture richness); and (b) the mixture’s functional composition (% of fast- vs. slow-decomposing species included in the mixture). We then tested if variation between observed and expected values of decomposition in mixtures was associated to: (c) the initial litter characteristics of the component species (initial nitrogen, lignin, cellulose and hemicellulose content of litters); and (d) the chemical heterogeneity of the mixtures (variation in the same chemical compounds between the components in each mixture). When up to 5 species representing different functional types were included, both species richness and functional composition showed statistically significant non-additive, and in general positive, effects on litter mixture decomposition. The positive effect of mixture richness on decomposition did not disappear, but was much less marked, when considering mixture with slow-decomposing species only. Although the main driver of decomposition in a mixture is still the average decomposability of the component species (itself largely determined by litter quality), the species interactions in a mixture add a consistent source of variability that is worth considering when predicting the decomposability of a given mixture. We showed that a greater positive difference between observed decomposition rates and that expected from its component species alone was found in mixtures with higher mean nitrogen content and a higher heterogeneity in non-labile compounds. Our results offer quantitative proof that litter chemical heterogeneity, as well as the mean quality of the mixture, can affect the decomposability in litter mixtures.     

Queen pheromones affecting the production of queen-like secretion in workers

The honeybee queen pheromones promote both worker sterility and worker-like pheromone composition; in their absence workers become fertile and express the queen pheromones. Which of the queen pheromones regulate worker pheromone expression and how, is still elusive. Here we investigated how two queen pheromones, the mandibular and Dufour’s, singly or combined, affect worker ovarian activation and occurrence of queen-like Dufour’s esters. Although queen mandibular pheromone (QMP) alone, or combined with Dufour’s secretion, inhibited to some extent worker reproduction, neither was as effective as the queen. The effect of the queen pheromones on worker pheromone expression was limited to workers with developed ovaries. Here too, QMP and Dufour’s combined had the greatest inhibitory effect. In contrast, treatment with Dufour’s alone resulted in augmentation of esters in the workers. This is another demonstration that a pheromone emitted by one individual affects the rates of its production in another individual. Ester production was tightly coupled to ovarian development. However fertile workers from queenright or QMP-treated colonies had significantly higher amounts of esters in their Dufour’s gland than untreated queenless colonies. The fact that the queen or QMP exert greater suppression on signal production than on ovary activation, suggests disparate regulatory pathways, and presents a challenging ultimate as well as proximate questions.     

The mutagenic and antimutagenic effects of Ecballium elaterium fruit juice in human peripheral lymphocytes

The aim of this study was to investigate the mutagenic and antimutagenic effects of Ecballium elaterium (EE) fruit juice, which has an anti-inflammatory effect, using in vitro human peripheral lymphocytes. To investigate the mutagenic effects of the EE fruit juice, human peripheral lymphocytes were treated with three doses (18, 36, and 72 μl/l) of fruit juice alone for 24 and 48 h. For investigating the antimutagenic effects of the EE fruit juice, the human lymphocytes were also treated with the mixture of the fruit juice and 0.25 μg/ml MMC. The EE fruit juice induced the percentage of total CA when used alone (especially the percentage of structural CA than the percentage of the numerical CA) and synergically induced the percentage of total CA when used as a mixture with MMC. The EE fruit juice did not affect the SCE frequency for 24 and 48 h treatment time. In contrast, EE and MMC as a mixture sinergically induced the SCE frequency at the highest concentration for 48 h treatment time only. EE alone did not decrease the RI while it decreased the MI in a dose-dependent manner. EE and MMC as a mixture have a higher cytotoxic effect than the cytotoxic effects of EE alone. As a result, it can be concluded that EE had no antimutagenic effect while EE had a mutagenic and a cytotoxic effect in human peripheral lymphocytes. This article was submitted by the authors in English.     

Asymmetric suppression of components in binary aldehyde mixtures: behavioral studies in the laboratory rat

The aim of the present study was to assess component interaction in the perception of the 2 aldehydes butanal and heptanal when presented in binary mixtures to rats. A further aim was to develop a behavioral paradigm for testing suppression of components in mixtures using rodent subjects. Thirsty rats were initially trained to discriminate between the 2 aldehydes butanal and heptanal in an olfactometer using a go/no-go discrimination task. This involved rats learning to place their noses in a sniff port where odors were presented and to lick a tube for water reward when one of the aldehydes was presented (S+) while withholding licking at the tube to the other, unrewarded, aldehyde (S-). A mixture condition was then introduced into the task, whereby a proportion of trials involved presentation of a combination of the 2 aldehydes as an additional unrewarded condition. Rats readily learned to withhold licking on trials when the mixture was presented. The concentration of the nonrewarded (S-) aldehyde in the mixture was then systematically decreased, whereas the concentration of the S+ component was held constant. This eventually caused the S+ component in the mixture to suppress detection of the S-, as shown by an increasing number of lick responses (false alarms) on trials when the mixture was presented. These suppressing effects occurred well above the detection threshold for the S- aldehyde presented alone. Results showed asymmetric suppression in the mixture condition such that butanal suppressed detection of heptanal at much lower concentrations than vice versa. A second experiment showed that when both butanal and heptanal were present in a binary mixture at the same concentration (10(-6) volume %), then rats responded to the mixture as if only butanal was present. These findings are discussed in terms of butanal having higher mobility and being able to compete more effectively than heptanal for occupation of shared receptor sites.     

Behavioural discrimination of oilseed rape volatiles by the honeybee Apis mellifera L.

This study investigated the ability of the honeybee to discriminatebetween six compounds previously identified as oilseed rapefloral volatiles: linalool, 2-phenylethanol, methyl salicylate,benzyl alcohol, (E)-2-hexenal and 1-octen-3-ol. These componentswere tested individually or in a synthetic mixture for theirability to elicit the conditioned proboscis extension response.Three experiments were done: conditioning to the mixture atone concentration (1.0, 0.1 or 0.01 µg per component)and testing to the individual components either at the conditioningor at the other concentrations; conditioning to individual componentsand testing to the mixture (1.0 µg); conditioning andtesting to the individual components (1.0 µg). The resultsfrom the proboscis extension assay were then compared to thoseobtained from free-flying bees in a flight room. From the conditionedproboscis extension assay, a conditioning threshold level wasfound (0.1 µg for the mixture studied) below which a reliableconditioning could not be achieved. Recognition thresholds wereobserved: bees responded to concentrations ten-fold higher orlower than that used for conditioning. Responses to lower concentrationswere weaker, whilst responses were increased at concentrationshigher than the conditioning one. A hierarchy within the compoundstested was found, with linalool, 2-phenylethanol and methylsalicylate cueing mixture recognition more effectively thanthe other components. The ranking order of the six componentswas similar in both the conditioned proboscis extension andthe conditioned foraging behaviour in flight room. When conditionedand tested to the individual components, bees discriminateda learned odour from a number of others. However, the specificitylevel for the recognition of the learned odour varied accordingto the component, the most clearly discriminated being the compoundswhich were used by the bee in mixture recognition.     

Effects of decorin on the expression of α-smooth muscle actin in a human myofibroblast cell line

Myofibroblasts are metabolically and morphologically distinctive fibroblasts expressing α-smooth muscle actin (α-SMA), and their activation plays a key role in development of the fibrotic response. In an activated state, myofibroblasts cease to proliferate and start to synthesize large amounts of extracellular component proteins. The expression of α-SMA correlates with the activation of myofibroblasts. Decorin, a member of the small leucine-rich proteoglycan gene family, has been implicated in the negative control of cell proliferation primarily by upregulating the expression of p21, a potent inhibitor of cyclin-dependent kinase. In order to examine the effect of decorin on myofibroblast cell growth, we rendered a human lung myofibroblast cell line, MRC-5, quiescent by either cell–cell contact or serum starvation, and examined the relationship between decorin and α-SMA expression in these cells. The expression of decorin in cells made quiescent by serum starvation was lower than that in cells made quiescent by cell–cell contact. In contrast, the expression of α-SMA in cells made quiescent by cell–cell contact was lower than that in cells made quiescent by serum starvation. Furthermore, forced expression of decorin was accompanied by a suppression of α-SMA expression, whereas knocking down of decorin expression by RNA interference increased the expression of α-SMA.     

Improved stem cell MR detectability in animal models by modification of the inhalation gas

In vivo monitoring of cells labeled with paramagnetic iron oxide particles by magnetic resonance imaging (MRI) is complicated by intrinsic contrast of blood vessels. Distinction between T2* effects caused by blood vessels from those caused by labeled cells was so far only possible after carefully following the location of hypointense regions through subsequent slices of T2*-weighted 3-D MRI datasets, a procedure that is time consuming and not always reliable in the case of smaller blood vessels. Here, we demonstrate that the modification of the inhalation gas mixture from the routinely used composition 35% O2 and 65% N2O to a mixture containing 95% O2 and 5% CO2 results in a contrast suppression of most small blood vessels and reduces the intrinsic T2* effect of large vessels dramatically in an animal model. This change in protocol of physiological conditions was well tolerated by all studied animals, even over prolonged experimental times. The changed inhalation gas mixture thus provides a more reliable identification method for small clusters of iron oxide labeled cells in vivo.     

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.     

Effect of oral imperatorin on memory in mice

The aim of this study was to explore the effect of the acetylcholinesterase inhibiting mixture of extracts of Angelica archangelica fruit and Geranium sylvaticum on memory. Furthermore the effect of the main compound, the furanocoumarin imperatorin, which has been shown to affect several neurotransmitters, was studied. Passive avoidance was measured by step-down latency and step-through latency of 10 months old mice receiving 0.79 mg/kg of imperatorin daily, pure or as part of the extracts, for 14 days or longer. Step-down latency was significantly higher in both groups receiving imperatorin than in the control group. In contrast, no difference was found between treatment groups regarding step-through latency. The results indicate that the imperatorin is the main active component of the extract mixture.