Mutual cross-adaptation of the volatile steroid androstenone and a non-steroid perceptual analog

Self-and cross-adaptation are believed to result from stimulationof the same olfactory sensory channels. These adaptation phenomenawere studied after exposures to 5-androst-16-en-3-one (androstenone)and a synthetic perceptual analog (DMCMC), viz. a racemic mixtureof the isomers 4(R)-(4',4'-dimethyl-cyclohexyl)-2(R)-methylcyclohexanoneand 4(S)-(4',4'-dimethylcyclohexyl)-2(S)-methylcyclohexanone.In Experiment 1, six subjects very sensitive to androstenonereceived four randomized sequences of six concentrations offour ordants (androstenone, DMCMC, amyl acetate, and Galaxolide*;plus blanks) before and following adaptation to either androstenoneor DMCMC. Exposure to each odorant resulted in self-adaptation.Measures of stimulus intensity and identification thresholdrevealed reciprocal cross-adaptation between androstenone andDMCMC, but no cross-adaptation to amyl acetate or Galaxolide.The degree of cross-adaptation was asymmetric; adaptation toDMCMC resulted in more complete adaptation to androstenone thanvice versa. This asymmetry was apparently due to intensity differences;when stimuli were matched for intensity, the asymmetry disappeared(Experiment 2). These results demonstrate cross-adaptation forqualitatively similar, but not dissimilar, odors and suggestthat androstenone and its perceptual analog DMCMC share thesame sensory channels.     

Olfactory adaptation as an aspect of odor similarity

Two experiments examined self-adaptation within and cross-adaptationbetween two structurally different substances with almost identicalbitter chocolate odors, trimethyl pyrazine (TMP) and 2-propionyl-3-methylfuran (PMF). The first experiment charted psychophysical functionsfor odor intensity under steady-state self- and cross-adaptationwith adapting concentrations matched in perceived intensityat two different levels. Participants breathed an adapting concentrationcontinuously for 2 min and then rated test concentrations everyminute during momentary respites from the adapting stimulus.Both self-adaptation and cross-adaptation weakened the intensityof low test concentrations proportionally more than higher onesand thereby steepened the psychophysical functions. Adaptingstimuli at matched levels caused equivalent self-adaptation(i.e. altered the functions equivalently) and equivalent, thoughweaker, cross-adaptation (i.e. exhibited symmetry of cross-adaptation).The second experiment examined whether cross-adaptation to TMPand PMF would prove relatively specific. At a matched perceivedintensity of adapting stimulus, adaptation to TPM and PMF againproduced equal amounts of self-adaptation and of cross-adaptationto each other, though not on three other test stimuli: anethole,ethyl butyrate and 2,3 pentanedione. Both TMP and PMF showedsome generality as cross-adapting stimuli, with PMF showinga bit more than TMP. The present protocols and others discussedherein offer opportunities to consider functional criteria forthe relatedness of similar-smelling substances.     

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.     

Structure/activity relationships in the arginine taste pathway of the channel catfish

To characterize the molecular/structural requirements for activationor antagonism of the arginine taste pathways in catfish, Ictaluruspunctatus, structure/activity studies were performed using integratedmultiunit responses and cross-adaptation. Of all the guanidinium-containingcompounds tested, only L-arginine, L--amino-ß-guanidinopropionic acid (L-AGPA) and L-arginine methyl and ethyl esterswere strong stimuli. Results of functional group substitutionsand modification of the L-arginine parent molecule indicatedthat: (i) stereospecificity was observed with D-arginine beinga much less effective stimulus than L-arginine; (ii) an L-aminogroup must be present and unblocked (-chloro-guanidino-N-valericacid and N-acetyl L-arginine were weak or inactive stimuli);(iii) a free carboxylic acid group was not necessary for activity;(iv) the distance between the anomeric carbon and the guanidiniumgroup was not critical (L-AGPA, having two methylene groupsless than L-arginine was a moderately strong stimulus as wasL-canavanine) and (v) modification or substitution of the guanidinumgroup by other basic groups including amine, methyl or dimethylamineor by an isosterc (ureido) resulted in loss of stimulatory ability.In general, those stimuli and analogs that were good cross-adaptersof L-arginine stimulation were also good competitors for L-[³H]argininebinding to a partial membrane fraction (P2) from catfish tasteepithelium. On the other hand, compounds that were poor cross-adaptingstimuli were also poor binding competitors. While D-argininewas a poor stimulus, it did cross-adapt L-arginine and competedwell with L-[³H]arginine for binding to fraction P2.     

PSYCHOPHYSICAL CHARACTERIZATION OF MUSK CHEMICALS

The odor intensities of six commercial musks: celestolide, muskolactone,galaxolide, musk 36A^®, tonalid^®, and musk ambrette^*were matched by the method of equal odor intensity. Galaxolidewas arbitrarily used as the reference odorant in order to generatea scale for the determination of relative odor intensities.The odor intensities are expressed in terms of Stevens' psychophysicalpower function with the exponent value, b, ranging from 0.43to 0.73.     

Asymmetrical neural cross-adaptation of hamster chorda tympani responses to sodium and chloride salts

Cross-adaptation has occurred when exposure to an adapting chemicalstimulus (A) reduces the response to a subsequent test stimulus(B). The degree of cross-adaptation between two stimuli is thoughtto reflect the overlap of their ‘neural activation processes’.We measured self- (A—A) and reciprocal crossadaptation(A—B, B—A) of the response of the hamster chordatympani nerve with lingual presentations of stimuli elicitingequal unadapted transient responses. Adapting and test stimuliwere 0.1 M NaCl, 0.1 M NaNO₃, 0.1 M NaBr, 0.4 M Na acetate (NaAc),0.09 M LiCl and 0.4 M NH₄Cl. Nearly complete and symmetricalcross-adaptation was seen for NaCl, NaNO₃ and NaBr. Those Nasalts paired with LiCl showed strong but asymmetrical cross-adaptation.Exposure to sodium completely eliminated the response to LiClbut not vice versa, suggesting that lithium and sodium are notcompletely interchangeable taste stimuli for the hamster chordatympani. Relatively little cross-adaptation between NH₄Cl andother salts suggested relatively separate neural activationprocesses. Strongly asymmetrical cross-adaptation was foundbetween NaAc and the other sodium salts. Responses to NaCl,NaNO₃ or NaBr were eliminated after adaptation to NaAc whereasthe response to NaAc during the reciprocal cross was strong.Asymmetries are discussed in reference to sensitivities of singlenerve fibers for the chorda tympani, effects of adaptation andthe concept of anion inhibition.     

Gustatory responses of eel palatine receptors to amino acids and carboxylic acids

The gustatory receptors of the eel palate were found to be extremely sensitive to amino acids and carboxylic acids. The results obtained are as follows: (a) 11 amino acids which are among naturally occurring amino acids elicited responses in the palatine nerve, but 9 amino acids did not elicit a response even at a high concentration. The effect of D-amino acids was always much less than that of their corresponding L-isomers. There was no appreciable difference in the effectiveness of an alpha-amino acid (alpha-alanine) and beta-amino acid (beta-alanine). (b) The threshold concentrations of the most potent amino acids (arginine, glycine) were between 10(-8) and 10(-9) M. A linear relation between the magnitude of the response and log stimulus concentration held for a wide concentration range for all the amino acids examined. (c) The palatine receptors responded sensitively to various carboxylic acid solutions whose pH was adjusted to neutral. The threshold concentrations varied between 10(-4) and 10(-7) M. The magnitude of the response at 10(-2) M increased with an increase of carbon chain length. (d) The extent of cross-adaptation was examined with various combinations of amino acids. A variety of the response patterns showing complete cross-adaptation, no cross-adaptation, or synergetic interaction was observed. The synergetic interaction was also observed when one amino acid below its threshold concentration was added to the other amino acid below its threshold concentration was added to the other amino acid. No cross-adaptation was observed between amino acids and fatty acids. (e) The treatment of the palate with papain led to loss of the responses to arginine, glycine, and histidine without affecting those to proline and acetic acid. The treatment with pronase E eliminated selectively the response to proline. The possibility that the eel gustatory receptors are responsible for sensing food at a distance was discussed.     

A model of paradoxical odour mixture perception

An identification experiment on a series of 10 graded mixturesof musk and n-propanol, with constrained (approximately constant)total concentration was modelled by a psychophysical functionwhich reproduced up to two local discontinuities in the perceptionof the intensity of one component; such results can be paralleledby analogous perceptual processes in vision. The model usedis an extension of previous models on odour cross-adaptation,with additional boundary conditions added.     

From musk to body odor: Decoding olfaction through genetic variation

The olfactory system combines input from multiple receptor types to represent odor information, but there are few explicit examples relating olfactory receptor (OR) activity patterns to odor perception. To uncover these relationships, we performed genome-wide scans on odor-perception phenotypes for ten odors in 1000 Han Chinese and validated results for six of these odors in an ethnically diverse population (n = 364). In both populations, consistent with previous studies, we replicated three previously reported associations (β-ionone/OR5A1, androstenone/OR7D4, cis-3-hexen-1-ol/OR2J3 LD-band), but not for odors containing aldehydes, suggesting that olfactory phenotype/genotype studies are robust across populations. Two novel associations between an OR and odor perception contribute to our understanding of olfactory coding. First, we found a SNP in OR51B2 that associated with trans-3-methyl-2-hexenoic acid, a key component of human underarm odor. Second, we found two linked SNPs associated with the musk Galaxolide in a novel musk receptor, OR4D6, which is also the first human OR shown to drive specific anosmia to a musk compound. We noticed that SNPs detected for odor intensity were enriched with amino acid substitutions, implying functional changes of odor receptors. Furthermore, we also found that the derived alleles of the SNPs tend to be associated with reduced odor intensity, supporting the hypothesis that the primate olfactory gene repertoire has degenerated over time. This study provides information about coding for human body odor, and gives us insight into broader mechanisms of olfactory coding, such as how differential OR activation can converge on a similar percept.     

Topographical EEG maps of human responses to odors

The physiological response to seven odors (birch tar, galbanum,heliotropine, jasmine, lavender, lemon and peppermint) was assessedby EEG recordings from 19 scalp loci from 16 young adult females.Topographic maps were constructed from the amplitude spectrain four frequency bands: delta (1–4 Hz), theta (4–8Hz), alpha (8–13 Hz) and beta (13–30 Hz). Eightseconds of representative and artifact-free EEG were selectedfor FFT analysis before onset of odor delivery, and at threetimes after stimulus onset. EEG was also quantified at 30 safter stimulus termination. Subjects differed in their subjective responses to the odors,with the most consistently arousing and strong odors being galbanum,lavender, lemon and peppermint. Heliotropine was notably weak.The most pleasant odors were lemon and peppermint, while birchtar, galbanum and lavender were consistently unpleasant. EEGmap changes occurred in one or more frequency bands in eachsubject in response to one or more of the odors. EEG map changessometimes occurred even with weak odors and even when the subjectseemed unaware of the odor's presence. This was most notablewith heliotropine. Across subjects, the most consistent responses to odors werein the theta band. Analysis of variance confirmed that certainodors caused statistically significant theta increases overthe left anterior group of electrodes. Both right hemispheregroups tended to have significant theta increases. The odorsthat caused the greatest increase in theta were birch tar, jasmine,lavender and lemon. On the other hand, during blank controltrials, theta activity diminished. There was also a significantodor by epoch interaction over the right posterior set of electrodes.Several of the odors caused increased theta at one or more epochs.Lemon caused an immediate increase in theta that abated withtime. Birch tar induced a delayed response that persisted afterthe stimulus was turned off. Jasmine and lavender tended toinduce theta sooner than birch tar, but the effect did not outlastthe stimulus. Increased theta was not associated with EEG signsof drowsiness. We conclude that all odors affected the EEG in at least somesubjects, and all subjects responded to at least some odors.Widespread increase in theta occurred in most subjects duringstimulation with such odors as birch tar, jasmine, lavenderand lemon.     

Specialized receptor neurons for pheromones and host plant odors in the boll weevil, Anthonomus grandis Boh. (Coleoptera: Curculionidae)

Antennal olfactory receptor neurons in the boll weevil, Anthonomusgrandis, were investigated through single neuron recordings.Receptor neurons for both pheromone components and host plantodors were associated with type I sensilla within the sensoryband regions. Nine types of receptor neurons were identified,based on their responsiveness to the four aggregation pheromonecomponents and selected host plant odors. Three receptor neurontypes responded to either compound I, II or IV of the aggregationpheromone. Dose—response curves were similar for eachof these receptor neuron types, which differed only in theirkey compound. In each instance, I neurons responded primarilyto (+)-I, the optical isomer produced by the boll weevil whichwas found to be active in field tests. Receptor neurons forII also responded to a lesser degree to III, its aldehydic analog,at the same stimulus load. Six additional receptor neuron typesresponded to selected host plant odors: ß-caryophyllene,trans-2-hexen-l-o1 and other six carbon alcohols and aldehydes,trans-ß-ocimene, benzaldehyde, linalool, and B-bisabolol.These neurons were as responsive as, or in some cases more responsiveat the same stimulus load as receptor neurons for pheromonecomponents. Receptor neurons responsive to six-carbon alcoholsand aldehydes were generally most responsive to trans-2-hexen-l-ol.Receptor neurons for other plant odors responded principallyto only one compound among the odorants tested. However, responsesof these neurons were not uniform, suggesting possible specializationfor other unidentified key odorants. Comparisons were also madebetween single neuron and electroantennogram responses. Theresults indicate that the boll weevil, a narrowly oligaphagousinsect, detects its host plant at some distance, and utilizesinformation about a wide range of chemical structures in itsolfactory-mediated behavior.     

Facial EMG responses to odors in solitude and with an audience

Two experiments were undertaken to examine whether facial responsesto odors correlate with the hedonic odor evaluation. Experiment1 examined whether subjects (n = 20) spontaneously generatedfacial movements associated with odor evaluation when they aretested in private. To measure facial responses, EMG was recordedover six muscle regions (M. corrugator supercilii, M. procerus,M. nasalis, M. levator, M. orbicularis oculi and M. zygomaticusmajor) using surface electrodes. In experiment 2 the experimentalgroup (n = 10) smelled the odors while they were visually inspectedby the experimenter sitting in front of the test subjects. Thecontrol group (n = 10) performed the same experimental conditionas those subjects participating in experiment 1. Facial EMGover four mimetic muscle regions (M. nasalis, M. levator, M.zygomaticus major, M. orbicularis oculi) was measured whilesubjects smelled different odors. The main findings of thisstudy may be summarized as follows: (i) there was no correlationbetween valence rating and facial EMG responses; (ii) pleasantodors did Dot evoke smiles when subjects smelled the odors inprivate; (iii) in solitude, highly concentrated malodors evokedfacial EMG reactions of those mimetic muscles which are mainlyinvolved in generating a facial display of disgust; (iv) thosesubjects confronted with an audience showed stronger facialreactions over the periocular and cheek region (indicative ofa smile) during the smelling of pleasant odors than those wbosmelled these odors in private; (v) those subjects confrontedwith an audience showed stronger facial reactions over the M.nasalis region (indicative of a display of disgust) during thesmelling of malodors than those who smelled the malodors inprivate. These results were taken as evidence for a more socialcommunicative function of facial displays and strongly mitigatesthe reflexive-hedonic interpretation of facial displays to odorsas supposed by Steiner.     

Olfactory discriminating ability of lacustrine sockeye and masu salmon in various freshwaters

In order to study the olfactory discriminating ability of lacustrine sockeye salmon (Oncorhynchus nerka) and masu salmon (O. masou), the integrated olfactory nerve response to various freshwaters was recorded by electrophysiological techniques. In both species independent of sex and gonadal maturity, each freshwater caused a different olfactory response. Source and effluent waters of the culture pond at Toya Lake Station (the source and culture pond waters) evoked the minimum and maximum response magnitude, respectively. In cross-adaptation experiments, the culture pond water abolished all secondary responses to other freshwaters, and no freshwater abolished the secondary response to the culture pond water. The concentration response study revealed that the minimum concentration (threshold) to induce response to the culture pond water after adaptation to Lake Toya water was between 0.1 and 1.0%. The present study indicates that the olfactory organ of lacustrine salmonids may discriminate different intensities of various freshwater odors.     

Efficiency and the role of adaptation in klinokinesis

Klinokinesis is a behavioral mechanism in which an organism moves toward or away from a stimulus source by altering its frequency of change of direction without biasing its turns with respect to the stimulus field. Computer simulation was used to study the efficiency of, and the effect of sensory adaptation on, this behavioral strategy. In modeling an organism with perfect performance (no error in determining the intensity of the stimulus and ability to move in perfectly straight lines) efficiency was about 70% without adaptation, and declined as the rate of adaptation increased. In contrast, models with non-perfect (noisy) performance were frequently able to double or triple their reduced efficiency by adapting to the stimulus intensity. Three types of noise that degraded performance were simulated: (1) intensity noise described random fluctuations in the intensity of the stimulus that were not associated with movement of the organism in the stimulus field; (2) motor noise described random fluctuations in the direction of locomotion as the organism moved along; (3) developmental noise described random differences between individuals in a constant tendency to turn to a certain degree as they moved forward. Adaptation had similar effects with any of the three types of noise. If a particular type of noise was strong enough to degrade performance significantly, then optimal performance occurred with an adaptation rate of about 0.2 per step.     

Receptor sites for alanine,proline, and betaine in the palatal taste system of the puffer,Fugu pardalis

Summary To elucidate the relative independence of the receptor sites for amino acids and betaine in the gustatory system of fish, the neural responses from the ramus palatinus facialis innervating the anterior palate of the puffer, Fugu pardalis, were recorded.There were observed independent amino acidsensitive and betaine-sensitive fibers.Cross-adaptation to pairs of stimulant was studied. The pair stimulants were applied reciprocally, i.e. after adapting with one stimulus the second stimulus was applied and then reversed. There were observed 3 types of cross-effects for the pairs of stimulants tested: (1) a reciprocal profound cross-adaptation; (2) no distinct cross-adaptation; and (3) a reciprocal enhancement of the response between betaine and alanine or glycine. Such an enhancement between betaine and alanine occurred in the amino acid-sensitive fibers, suggesting that betaine has an enhancing effect on the amino acid receptors.The present results suggest at least 3 different groups of receptor sites for the stimulants tested: (1) alanine sites for alanine, glycine and sarcosine; (2) proline sites for proline and dimethylglycine; and (3) betaine sites for betaine and dimethylglycine.     

Copepod reaction to odor stimuli influenced by cestode infection

The cestode Schistocephalus solidus uses copepods as first andsticklebacks as second intermediate hosts. For transmission,an infected copepod has to be preyed upon by a stickleback.We used copepods of the species Macrocy albidus to test whetherinfected and uninfected copepods differ in their reaction totwo kind of simultaneously presented odors: odors of sticklebacksand odors of sticklebacks and conspecificz. By giving this choice,we attempted to force the copepods to make a trade-off betweenthe benefit of risk dilution and possible predator confusionand the costs of food competition and other disadvantages inducedby conspecifics. Within 1–8 h after last feeding, uninfectedcopepods clearly preferred the odors of conspeciflcs under thechemically simulated threat of predation. This was in contrastto the infected copepods, who tended to avoid the odor of conspecifics.When the time between experiment and last feeding varied, infectedcopepods showed an increas preference for fish water only (oravoided conspecthcs) with increasing hunger level This suggeststhat S. solidus benefits from hunger-induced behavioral changesof its copepod host by influencing its microhabitat selection.The same effect could be found in both sexes; however, it wassignificantly more pronounced in male than in female copepods.We propose several hypotheses that could explain the differencebetween the sexes in their infection-dependent microhabitatselection.     

Time Course and Action Spectrum of Vibrotactile Adaptation

In a series of experiments designed to explore the processes underlying adaptation of the sense of flutter-vibration, vibrotactile threshold was measured on the pad of the index finger, using Békésy tracking. Unadapted thresholds were first measured, for a number of frequencies (4-90 Hz) and contactor sizes (1-8 mm diameter). As expected, these measurements indicated the presence of (1) a Pacinian system possessing spatial summation and increasing in sensitivity, as frequency was raised, at the rate of 12 dB/octave; and (2) a non-Pacinian system showing little spatial summation, and with a frequency characteristic matching that of the NP I mechanism of Bolanowski et al. (1988). These baseline data of Experiment 1 guided the selection of stimulus parameters for subsequent experiments, in which threshold for a test stimulus was measured before, during, and after periods of vibrotactile adaptation.

In Experiment 2, test stimuli of 10 H_z and 50 H_z were combined factorially with 30-dB SL adapting stimuli of the same two frequencies. When the test stimulus was 10 Hz, the two adapting frequencies were equally effective in raising threshold; however, when the 50-Hz test stimulus was used, the 50-Hz adapting stimulus raised threshold by a greater amount than did the 10-Hz adapter. These results confirm on the finger the independence of adaptation in Pacinian and non-Pacinian channels, a result previously established on the thenar by other workers. For all four frequency combinations, threshold rose exponentially with a time constant of 1.5-2 min.

In Experiment 3, an action spectrum was determined, showing the adapting amplitude needed at each of a series of frequencies to raise the threshold of a 10-Hz stimulus by 10 dB; this spectrum was essentially flat from 30 to 90 Hz. The results, taken in conjunction with what is known about rapidly adapting cutaneous mechanoreceptors, imply that the effectiveness of an adapting stimulus is not determined solely by the amount of activity it generates in first-order afferents.     

Odor Perception Phenotypes: Multiple, Specific Hyperosmias to Musks

Olfactory detection thresholds for 11 structurally diverse muskodorants and one non-musk odorant were obtained from 32 subjects.Hierarchical cluster analysis produced four groups of subjects.One group (n = 12) was uniformly sensitive to all musks; another(n = 16) was uniformly insensitive. Two groups of subjects containedotherwise insensitive individuals who were exceptionally sensitiveto cyclopentadecanone and musk xylol (n = 2) and to delta9-hexadecenolactoneand tonalid (n = 2) respectively. We propose that the lattertwo groups are odor perception phenotypes (MSHM1 and MSHM2)that consist of multiple, specific hyperosmias to musk odorants.Chem. Senses 21: 411– 416, 1996. ¹Present address: Synesthetics, Inc., Montclair, NJ 07043, USA     

Time course and action spectrum of vibrotactile adaptation

In a series of experiments designed to explore the processes underlying adaptation of the sense of flutter-vibration, vibrotactile threshold was measured on the pad of the index finger, using Békésy tracking. Unadapted thresholds were first measured, for a number of frequencies (4-90 Hz) and contactor sizes (1-8 mm diameter). As expected, these measurements indicated the presence of (1) a Pacinian system possessing spatial summation and increasing in sensitivity, as frequency was raised, at the rate of 12 dB/octave; and (2) a non-Pacinian system showing little spatial summation, and with a frequency characteristic matching that of the NP I mechanism of Bolanowski et al. (1988). These baseline data of Experiment 1 guided the selection of stimulus parameters for subsequent experiments, in which threshold for a test stimulus was measured before, during, and after periods of vibrotactile adaptation. In Experiment 2, test stimuli of 10 Hz and 50 Hz were combined factorially with 30-dB SL adapting stimuli of the same two frequencies. When the test stimulus was 10 Hz, the two adapting frequencies were equally effective in raising threshold; however, when the 50-Hz test stimulus was used, the 50-Hz adapting stimulus raised threshold by a greater amount than did the 10-Hz adapter. These results confirm on the finger the independence of adaptation in Pacinian and non-Pacinian channels, a result previously established on the thenar by other workers. For all four frequency combinations, threshold rose exponentially with a time constant of 1.5-2 min. In Experiment 3, an action spectrum was determined, showing the adapting amplitude needed at each of a series of frequencies to raise the threshold of a 10-Hz stimulus by 10 dB; this spectrum was essentially flat from 30 to 90 Hz. The results, taken in conjunction with what is known about rapidly adapting cutaneous mechanoreceptors, imply that the effectiveness of an adapting stimulus is not determined solely by the amount of activity it generates in first-order afferents.     

Odor sensitivity to geosmin enantiomers

Enantiomers of the earthy smelling odorant ‘geosmin’(t-1, 10-dimethyl-t-9-decalol) were tested on 52 human subjectsfor odor threshold ratio (+)/(–) variability. Thresholdconcentrations of (+) and (–) and their (+)/(–)ratios appeared to be monomodally distributed. For most subjects,the ratio varied within a predictable narrow range comparableto that found previously for carvone enantiomers and contraryto the large variability found for -ionone enantiomers (Polaket al., 1989). The natural occurring isomer, (–) geosminhad, on average an x11 lower threshold than (+).