Qualitative and aqualitative intensity components of odour mixtures

The perception of the intensity of components of odorous mixturesis related to the overall mixture intensity in a complex mannerwhich has not so far been adequately modelled. Hypoadditivity,where the total perceived intensity of a mixture is less thanthe sum of the component intensities which arise when the substancesare presented in pure form at equivalent chemical concentrations,has been known for many years. An experiment on series of 56mixtures of amyl butyrate and bergamot at supra-threshold concentrationsfound that the total qualitatively unspecified intensity ofsuch mixtures can be greater than the sum of the identifiedcomponent intensities as judged in the context of the mixture.To reconcile an apparent contradiction, and to bring into onepsychophysical framework results from diverse procedures, amodel of mixed qualitative and aqualitative odour intensityis advanced, incorporating some evidence on the difference betweendetection and recognition processes. Each component in a mixturemay, in this theory, both partially mask the qualitative intensityof the other, and simultaneously convert some part of the other'sintensity into an aqualitative form.     

Odor intensity of binary mixtures of odorous compounds

This paper presents the MBO model for the perceived intensity of odour mixtures. This model is based on an equation previously reported by our team, intended to model the whole stimulus-response intensity curve of pure odorous compounds. The MBO model was applied to a set of published data, and compared to other published models. The results show a high modelling efficiency of the MBO model compared to other proposed equations, especially for binary mixtures exhibiting significant asymmetry of intensity for different ratios of the two components. Furthermore, the MBO model includes parameters specific to the respective effects of each component in the mixture, which may help to clarify the masking and synergy effects that are often sought in odour mixtures.     

Perceptual interactions between characteristic notes smelled above aqueous solutions of odorant mixtures

Twenty-two experienced panelists rated odor intensity of aqueous solutions of citral, octen-1-ol-3, and hexanal. The panel assessed unmixed components and mixtures (9 binary and 4 ternary). In sensory sessions dedicated to mixtures (n = 6), evaluation was focused on one target odor, presented at a fixed concentration. All components had lower odor intensity on mixed presentations. In many cases, information obtained from simpler systems was not extended to complex mixtures. In a mixture, the competition between odorant molecules on qualitative aspects (dominance/suppression) imbalanced components contribution, anticipated from the quantitative distribution. Hexanal appeared to be the potentially weaker odorant in paired combinations, whereas octen-1-ol-3 had a lower relative impact on ternary systems. Suppression of the odor of octen-1-ol-3 and a concomitant increase in the odor of hexanal was common to all ternary mixtures. Reciprocal inhibition of octen-1-ol-3 and citral odors through perceptual interactions was suspected. Mutual suppression is suspected to have eased the perception of hexanal intensity.     

A comparison of the detection thresholds of odour mixtures and their components

It was the purpose of the present study to investigate whetherthe perception of odour mixtures differs from the perceptionof their components with regard to (i) the reliability of detectionthreshold measurements and (ii) the threshold values themselves.Detection thresholds for a variety of unmixed odorants and 3-,6- or 12-component mixtures were determined for a total of 44subjects, tested repeatedly in five separate sessions usingsniff bottles and an ascending staircase method. Threshold determinationsfor the mixtures were found to be as reliable as for the unmixedsubstances, with intra- and interindividual variability in thresholdvalues actually decreasing with increasing stimulus complexity.The four mixtures for which a systematic comparison of the thresholdvalues of components in the mixed and unmixed state was conductedfailed to provide evidence for the hypo-additive effects ofcompromise or compensation, but rather suggested a positive,sub-threshold interaction between components and possibly evenenhancement. Finally, a significant improvement in individualthreshold scores was seen across test sessions, with experiencedpanelists showing the most stable performance.     

Sequence matters – selective adaptation in electroantennographic response to binary odour mixtures by the Colorado potato beetle

Chemically mediated behaviour of insects is often strongly affected by mixtures of odour stimuli and their temporal characteristics. Both sensory transduction and central processing of odour mixtures can give rise to several different kinds of interaction, which can influence how individual components are perceived and processed. In particular, odour mixtures have been examined in model experiments as premixed binary mixtures in comparison with pure odour stimuli. Only in few experiments, the influence of the temporal structure of odour mixtures on odour perception has been taken into account. Natural odour stimuli often have a pulsed structure and may in general be superimposed on a background of irrelevant or interfering compounds, which can fluctuate with different frequencies, depending on their source. To achieve a better representation of these natural conditions, our odour mixing experiments apply a new kind of stimulation protocol: odours were not premixed but superimposed with a specific time pattern; one odour stimulus was presented as a longer persisting background and the second stimulus was a superimposed short test signal. To gain an overview of odour interaction patterns in the Colorado potato beetle by causing adaptation of one receptor population at naturally occurring levels of concentration and time intervals, electroantennographic recordings were made on excised antennae. A matrix of 12 stimulus compounds led to 132 pairs of compounds tested, each in the role of background and test stimulus. In 64 cases, the interaction was significantly different, when the role of background and stimulus was exchanged. Interaction patterns ranging from no interference (independence) to suppression were found and assigned to four clearly distinguishable types. We discuss that the observed effects of the presentation sequence in odour mixtures may contribute to the mechanisms of olfactory pattern recognition and olfactory contrast perception by insects.     

A critical analysis of the odour unit number and its use

The "odour unit number" used in applied flavour research andknown by several different names, is neither a unit of absolutephysical concentration of an odorant, nor is it a psychophysicalmeasure for perceived odour intensity. The use of this concepthas been shown to rest upon implicit assumptions which are contradictoryin some respects to present psychophysical knowledge. A specialconsequence for the mixture threshold value of the defined relationshipbetween odour unit numbers of components and that number ofthe mixture from which they are parts was shortly discussed.Finally, a few words were devoted to a psychophysical approachin the study of interaction phenomena in odorant mixtures.     

Neonatal representation of odour objects: distinct memories of the whole and its parts

Extraction of relevant information from highly complex environments is a prerequisite to survival. Within odour mixtures, such information is contained in the odours of specific elements or in the mixture configuration perceived as a whole unique odour. For instance, an AB mixture of the element A (ethyl isobutyrate) and the element B (ethyl maltol) generates a configural AB percept in humans and apparently in another species, the rabbit. Here, we examined whether the memory of such a configuration is distinct from the memory of the individual odorants. Taking advantage of the newborn rabbit''s ability to learn odour mixtures, we combined behavioural and pharmacological tools to specifically eliminate elemental memory of A and B after conditioning to the AB mixture and evaluate consequences on configural memory of AB. The amnesic treatment suppressed responsiveness to A and B but not to AB. Two other experiments confirmed the specific perception and particular memory of the AB mixture. These data demonstrate the existence of configurations in certain odour mixtures and their representation as unique objects: after learning, animals form a configural memory of these mixtures, which coexists with, but is relatively dissociated from, memory of their elements. This capability emerges very early in life.     

Synergy and masking in odor mixtures: an electrophysiological study of orthonasal vs. retronasal perception

Perceptual interactions in a model of wine woody-fruity binary mixtures were previously reported in a psychophysical study performed through orthonasal stimulation only. However, recent studies suggested that the perception of food-like and nonfood-like odors may depend on the route of stimulation. The aim of the present study was two-fold: first to examine the neural correlates of perceptual interactions using electroencephalogram (EEG)-derived event-related potentials (ERPs) and second to test the influence of the stimulation route on quality perception. Therefore, we designed an experiment with 30 subjects to study perceptual interactions in woody-fruity mixtures and compared ortho- vs. retronasal stimulation sites on perceived odor quality and ERPs. The results revealed synergy or masking of the fruity component, depending on the woody component level. Synergy was supported by larger N1 amplitude of the ERP. Furthermore, mixtures including a medium level of the woody odor elicited a strong increase of P2 amplitude only retronasally. This study evidenced for the first time electrophysiological correlates of both perceptual synergy and masking on the early component of the ERPs and confirmed that retro- vs. orthonasal stimulation route induces different neural processes that are reflected in the late component of the ERP.     

Prediction models for the pleasantness of binary mixtures in olfaction

Whereas the rules underlying the perceived intensity of binary mixtures have been investigated, minimal efforts have been directed at elucidating the rules underlying the perceived pleasantness of such mixtures. To address this, 84 subjects ranked the pleasantness and intensity of 5 distinct binary mixtures (15 pairs, inter-stimulus interval = 4 s, inter-trial interval = 30 s, flow = 6 l/min, pulse = 2 s) constructed from different ratios (0:100%, 25:75%, 50:50%, 75:25%, and 100:0%, olfactometer-generated vapor phase). We found that in the majority of cases, the pleasantness of the mixture fell between the pleasantness values of its separated constituents and that it was strongly influenced by the relative intensities of the constituents. Based on these results, we proposed a prediction paradigm for the pleasantness of binary mixtures from the pleasantness of their separated constituents weighted by their respective perceived intensities. The uniqueness of the proposed paradigm is that it neither requires presetting an interaction constant between the mixture components nor require any factorization of the pleasantness weights. It does, nonetheless, require solid psychophysical data on the separated components at their different concentrations, and currently it can only explain the behavior of intermediate pleasantness of mixtures.     

Latency and accuracy of discriminations of odor quality between binary mixtures and their components

Subjects made timed, same-different discriminations of odor quality, with the following principal findings: (i) latency reflected accuracy, with difficult discriminations, i.e. those between 50-50 mixtures and their components, requiring more time than less difficult discriminations, i.e. those between unmixed chemicals. This finding demonstrated the face validity of latency as a measure of qualitative similarity. (ii) Latency provided better resolution among pairs of odors than did errors of discrimination. This finding demonstrated the utility of collecting response times. (iii) Latency-based similarities among odors tested previously predicted similarities among pairs not yet tested. This finding demonstrated internal/predictive validity. (iv) A signal detection model assuming a differencing strategy best described the pattern of errors. Subjects appeared to make relative judgements regarding quality. (v) Finally, latency-based similarities between mixtures and their components demonstrated additivity. This finding suggested that binary mixtures fall on straight lines connecting their components in 'odor-space'.     

Behavioral discrimination of binary mixtures and their components: effects of mixture interactions on coding of stimulus intensity and quality

There is mounting evidence that mixture interactions resultin a physiological response that is different from that predictedfrom observed responses to individual mixture components. Mixtureinteractions that act to alter the neural coding of mixtureintensity (intensity mixture interactions) or quality (patternmixture interactions) may ultimately lead to dramatic differencesbetween the perceived intensities and qualities of a mixtureand its components. These perceptions could be expressed andobserved at the behavioral level. Toward examining this question,we have tested the ability of the Florida spiny lobster (Panulirusargus) to behaviorally discriminate between three odorant compounds[adenosine 5'-monophosphate (AMP), L-glutamate (Glu), and taurine(Tau)] and their binary mixtures through the use of a differentialaversive associative conditioning paradigm. Six groups of lobsterswere used, each being conditioned to avoid one of the singlecompounds or binary mixtures. Behavioral expression of intensitymixture interactions was evident. Preconditioning response magnitudesto binary mixtures were either less than those to their components(e.g. AMP + Glu) or less than predicted from responses to theircomponents (e.g. AMP + Tau). Behavioral expression of patternmixture interactions was also observed. Relationships betweenthe quality of each binary mixture and the qualities of themixture's components were determined from the results of analysisof variance and multidimensional scaling analysis. Analysesincorporated observed responses to all stimuli and ‘predicted’responses to the binary mixtures. Lobsters easily discriminatedbetween the qualities of AMP, Glu and Tau. The quality of themixture of AMP + Glu was different from either component aswell as from the predicted value for this mixture. The mixtureof AMP + Tau was intermediate between both components and wassimilar to the predicted value. The mixture of Glu + Tau, whilemore similar to Glu than to Tau, was different from the predictedvalue, and there was some indication that the Glu was actingto suppress the response to Tau. Behavioral results for AMP+ Tau, which suggest no pattern mixture interactions betweenthese compounds, are in accordance with results of recentlyconducted binding assays which indicate independent receptorsfor these compounds (Olson et al., 1992). Results, especiallyfor AMP + Glu and Glu + Tau, are consistent with results ofour electrophysiological analysis of the effects of patternmixture interactions on coding of stimulus quality and intensityby olfactory receptor cells (Derby et al., 1991a,b). This providesfurther evidence for the effects of peripherally initiated mixtureinteractions on the coding and perception of the quality ofodorant mixtures. ¹Present address: Departments of Psychology and Biology, GeorgiaState University, University Plaza, Atlanta, Georgia 30303,USA     

Sensory integration in citric acid/sucrose mixtures

The scale values of perceived sweetness, sourness and totaltaste intensity of unmixed sucrose, unmixed citric acid andseveral citric acid/sucrose mixtures were assessed, using afunctional measurement approach in combination with a two-stimulusprocedure. The data showed that the scale values obtained werelinear with perceived taste intensity. It was demonstrated thatcitric acid suppresses the sweetness of sucrose and that, inversely,sucrose suppresses the sourness of citric acid. However, thissuppressive effect was not symmetrical in the range of concentrationsused. While the degree of sweetness suppression depended onlyon the citric acid level, the degree of sourness suppressiondepended on the sucrose as well as on the citric acid concentration.With regard to the perceived total taste intensity of citricacid/sucrose mixtures, it was shown that the sum of sweetnessand sourness approximately equals the total taste intensity.The implications of the present findings for the analytic—syntheticcontroversy and for taste interaction theories are discussed.     

Binary taste mixture interactions in prop non-tasters, medium-tasters and super-tasters

It is generally assumed that the mutual, but asymmetric, suppression of the components in binary taste mixtures is an invariant property of the human psychophysical response to such mixtures. However, taste intensities have been shown to vary as a function of individual differences in sensitivity, indexed by the perceived bitterness of 6-n-propylthiouracil (PROP). To determine if these variations in taste perception influence taste mixture interactions, groups of PROP super-, medium- and non-tasters assessed four binary taste mixtures: sweet-bitter [sucrose/quinine hydrochloride (QHCl)], sweet-sour (sucrose/citric acid), salty-bitter (NaCl/QHCl) and salty-sour (NaCl/citric acid). In each experiment, subjects received factorial combinations of four levels of each of two tastants and rated individual taste intensities and overall mixture intensity. For each taste quality, super-tasters typically gave higher ratings than either medium- or non-tasters, who tended not to differ. There were also group differences in the interactions of the mixtures' components. Super-tasters rated the overall intensity of the mixtures, most likely reflecting integration of the taste components, as greater than medium- and non-tasters, who again showed few differences. In sweet-bitter mixtures, non-tasters failed to show the suppression of sweetness intensity by the highest QHCl concentration that was evident in super- and medium-tasters. These data show that the perception of both tastes and binary taste mixture interactions varies as a function of PROP taster status, but that this may only be evident when three taster groups are clearly distinguished from one another.     

Contextual effects in the perception of quinine HCl/NaCl mixtures

Stimulus distribution, stimulus spacing and stimulus range affectmean intensity ratings of solutions of unmixed tastants. Thepresent study compares contextual effects for mixture ratingswith those for unmixed stimuli: if ratings for mixed and unmixedstimuli are differentially affected by context, the degree ofmixture suppression inferred from the responses is context-dependent.Bitterness intensity ratings for unmixed quinine and quinineHCl/NaCl mixtures were not differentially affected by shiftsin stimulus frequency distribution. The subjects' tendency tobe consistent in their responses to identical stimuli, resultsin a transfer of stimulus context of previous sessions to thenext.     

The influence of sodium salts on binary mixtures of bitter-tasting compounds

In order to study potential mixture interactions among bitter compounds, selected sodium salts were added to five compounds presented either alone or as binary bitter-compound mixtures. Each compound was tested at a concentration that elicited 'weak' perceived bitterness. The bitter compounds were mixed at these concentrations to form a subset of possible binary mixtures. For comparison, the concentration of each solitary compound was doubled to measure bitterness inhibition at the higher intensity level elicited by the mixtures. The following sodium salts were tested for bitterness inhibition: 100 mM sodium chloride (salty), 100 mM sodium gluconate (salty), 100 and 20 mM monosodium glutamate (umami), and 50 mM adenosine monophosphate disodium salt (umami). Sucrose (sweet) was also employed as a bitterness suppressor. The sodium salts differentially suppressed the bitterness of compounds and their binary combinations. Although most bitter compounds were suppressed, the bitterness of tetralone was not suppressed, nor was the bitterness of the binary mixtures that contained it. In general, the percent suppression of binary mixtures of compounds was predicted by the average percent suppression of its two components. Within the constraints of the present study, the bitterness of mixtures was suppressed by sodium salts and sucrose independently, with few bitter interactions. This is consistent with observations that the bitter taste system integrates the bitterness of multi-compound solutions linearly.     

The similarity between odors and their binary mixtures in Drosophila

How are odor mixtures perceived? We take a behavioral approach toward this question, using associative odor-recognition experiments in Drosophila. We test how strongly flies avoid a binary mixture after punishment training with one of its constituent elements and how much, in turn, flies avoid an odor element if it had been a component of a previously punished binary mixture. A distinguishing feature of our approach is that we first adjust odors for task-relevant behavioral potency, that is, for equal learnability. Doing so, we find that 1) generalization between mixture and elements is symmetrical and partial, 2) elements are equally similar to all mixtures containing it and that 3) mixtures are equally similar to both their constituent elements. As boundary conditions for the applicability of these rules, we note that first, although variations in learnability are small and remain below statistical cut-off, these variations nevertheless correlate with the elements' perceptual "weight" in the mixture; thus, even small differences in learnability between the elements have the potential to feign mixture asymmetries. Second, the more distant the elements of a mixture are to each other in terms of their physicochemical properties, the more distant the flies regard the elements from the mixture. Thus, titrating for task-relevant behavioral potency and taking into account physicochemical relatedness of odors reveals rules of mixture perception that, maybe surprisingly, appear to be fairly simple.