The effects of adaptation on the perception of similar and dissimilar odors

Adaptation among several odors was studied using a multiple-alternative,forced-choice procedure where six concentrations of each odorantand three blanks were presented in a random sequence beforeand after adaptation to each of the odors. Adaptation was expressedin terms of changes in both identification threshold and perceivedintensity. In the first experiment, Galaxolide® (a syntheticisochroman musk) and 5-androst-16-en-3-one (androstenone) showedno cross-adaptation in spite of sharing, for some people, amusky note. In the second experiment, Galaxolide and Thibetolide®(a synthetic macrocyclic musk) showed significant, but asymmetric,cross-adaptation. When Galaxolide was the adapting stimulus,shifts in indentification threshold and magnitude estimatesfor Thibetolide were not significantly different from thoseobtained for Galaxolide, a result consistent with the conclusionthat cross-adaptation was as effective as self-adaptation. WhenThibetolide was the adapting stimulus, shifts in identificationthreshold and magnitude estimates for Galaxolide were significantlyless than those obtained for Thibetolide, suggesting that cross-adaptationwas less effective. This result is puzzling given the perceptualsimilarity of these two musky compounds: subjects did not distinguishbetween them during random presentation. The distinct odorsof amyl acetate (banana) and d-limonene (orange) showed no cross-adaptationin either experiment.     

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.     

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.     

Electrophysiological evidence for acidic, basic, and neutral amino acid olfactory receptor sites in the catfish

Electrophysiological experiments indicate that olfactory receptors of the channel catfish, Ictalurus punctatus, contain different receptor sites for the acidic (A), basic (B), and neutral amino acids; further, at least two partially interacting neutral sites exist, one for the hydrophilic neutral amino acids containing short side chains (SCN), and the second for the hydrophobic amino acids containing long side chains (LCN). The extent of cross-adaptation was determined by comparing the electro-olfactogram (EOG) responses to 20 "test" amino acids during continuous bathing of the olfactory mucosa with water only (control) to those during each of the eight "adapting" amino acid regimes. Both the adapting and test amino acids were adjusted in concentrations to provide approximately equal response magnitudes in the unadapted state. Under all eight adapting regimes, the test EOG responses were reduced from those obtained in the unadapted state, but substantial quantitative differences resulted, depending upon the molecular structure of the adapting stimulus. Analyses of the patterns of EOG responses to the test stimuli identified and characterized the respective "transduction processes," a term used to describe membrane events initiated by a particular subset of amino acid stimuli that are intricately linked to the origin of the olfactory receptor potential. Only when the stimulus compounds interact with different transduction processes are the stimuli assumed to bind to different membrane "sites." Four relatively independent L-alpha-amino acid transduction processes (and thus at least four binding sites) identified in this report include: (a) the A process for aspartic and glutamic acids; (b) the B process for arginine and lysine; (c) the SCN process for glycine, alanine, serine, glutamine, and possibly cysteine; (d) the LCN process for methionine, ethionine, valine, norvaline, leucine, norleucine, glutamic acid-gamma-methyl ester, histidine, phenylalanine, and also possibly cysteine. The specificities of these olfactory transduction processes in the catfish are similar to those for the biochemically determined receptor sites for amino acids in other species of fishes and to amino acid transport specificities in tissues of a variety of organisms.     

Cross-adaptation in taste measured with a filter-paper method

A filter-paper method was applied to self- and cross-adaptationin taste. The original method was modified in order to overcomethree problems: (i) unreliable replacement of the test stimulusafter removal of the adapting stimulus, (ii) incomplete removalof the adapting stimulus liquid, and (iii) migration of stimulusliquid to unadapted surrounding tissue. With NaCl, sucrose anddi-quinine sulfate >70% of complete self-adaptation was retainedafter substituting the adapting stimulus with an identical testpaper. Citric acid behaved differently. Only 45% of the completeadaptation survived the change from adapting to test stimulus.It is speculated that this exception of citric acid is due tospecific physico-chemical properties of acids combined withthe use of the filter-paper method. The experiments show thatthe filter-paper method provides a useful and reliable procedurefor the investigation of cross-adaptation in taste.     

Perception of approaching and withdrawing sound sources after exposure to broadband noise: The importance of spatial domain

The existence of space-specific differences in auditory aftereffects has been tested under short-term (5 s) exposure to broadband noise (20–20000 Hz). Adapting stimuli were emitted as constant-amplitude noise sequences. Test stimuli could be of constant and changing amplitude: increasing amplitude of noise pulses in a sequence mimicked an approaching sound source, whereas a decrease in amplitude was perceived as withdrawal. The experiments were done in an anechoic chamber. Auditory aftereffects were assessed under the following conditions: (a) adapting and test stimuli were emitted through a loudspeaker mounted at a distance of 1.1 m from the listener (i.e., subjectively near); (b) both stimuli were emitted from a distance of 4.5 m (subjectively far); (c) adapting and test stimuli were emitted from different distances. The results showed that the characteristics of perception of the imitated sound source motion were similar in proximity and remoteness, which was observed both in the control (without adaptation) and after adaptation to noise. In the absence of adaptation, the psychophysical curves were asymmetrical: the listeners reported approaching of test stimuli more often for both spatial domains. However, the overestimation of test stimuli as drawing closer was more pronounced when they were emitted from the distance of 1.1 m, i.e., from near the listener. After the adaptation to noise, the aftereffects showed spatial specificity and were observed only when adapting and test stimuli belonged to the same spatial domain. These aftereffects were similar in their pattern and strength both in proximity and remoteness: the listeners reported withdrawal of test stimuli more frequently as compared with the control. As a result of these aftereffects, the symmetry of psychometric curves was restored, and the estimation of the direction of sound source motion in the experiment became equiprobable.     

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.     

Chemosensory context effects: role of perceived similarity and neural commonality

Seven experiments investigated how stimulus context affects judgements of the magnitude of chemosensory stimuli. In each experiment, subjects gave magnitude estimates of the intensity of several concentrations of two substances, with the contextual set of concentrations varying across experimental conditions. Different experiments used different pairs of substances, which could be tastants (sucrose or NaCl) that were sipped, odorants (orange or vanillin) that were sipped (i.e. presented retronasally) or the same odorants sniffed (i.e. presented orthonasally). Varying the stimulus context affected the judgements differentially when the two substances were compositionally different (sucrose and NaCl; sucrose and orange; sucrose and vanillin) but not when they were the same (vanillin or orange presented orally and nasally). Judgements of qualitative similarity of the same pairs of substances, obtained in a separate experiment, failed to predict accurately the pattern of differential context effects. Taken together, the results suggest that differential effects of context relate only indirectly to perceptual dissimilarity per se but may primarily reflect the result of stimulus-specific adaptation-like processes.     

Neural population representation hypothesis of visual flow and its illusory after effect in the brain: psychophysics,neurophysiology and computational approaches

The neural representation of motion aftereffects induced by various visual flows (translational, rotational, motion-in-depth, and translational transparent flows) was studied under the hypothesis that the imbalances in discharge activities would occur in favor in the direction opposite to the adapting stimulation in the monkey MST cells (cells in the medial superior temporal area) which can discriminate the mode (i.e., translational, rotational, or motion-in-depth) of the given flow. In single-unit recording experiments conducted on anaesthetized monkeys, we found that the rate of spontaneous discharge and the sensitivity to a test stimulus moving in the preferred direction decreased after receiving an adapting stimulation moving in the preferred direction, whereas they increased after receiving an adapting stimulation moving in the null direction. To consistently explain the bidirectional perception of a transparent visual flow and its unidirectional motion aftereffect by the same hypothesis, we need to assume the existence of two subtypes of MST D cells which show directionally selective responses to a translational flow: component cells and integration cells. Our physiological investigation revealed that the MST D cells could be divided into two types: one responded to a transparent flow by two peaks at the instances when the direction of one of the component flow matched the preferred direction of the cell, and the other responded by a single peak at the instance when the direction of the integrated motion matched the preferred direction. In psychophysical experiments on human subjects, we found evidence for the existence of component and integration representations in the human brain. To explain the different motion perceptions, i.e., two transparent flows during presentation of the flows and a single flow in the opposite direction to the integrated flows after stopping the flow stimuli, we suggest that the pattern-discrimination system can select the motion representation that is consistent with the perception of the pattern from two motion representations. We discuss the computational aspects related to the integration of component motion fields.     

A comparison of the discriminatory ability and sensitivity of the trigeminal and olfactory systems to chemical stimuli in the tiger salamander

Summary Trigeminal receptors can respond to a wide variety of chemical stimuli, but it is unknown whether these receptors mediate discrimination between chemical stimuli matched for equal perceptual intensity. The present electrophysiological and behavioral experiments address this issue using tiger salamanders, Ambystoma tigrinum, and four compounds (amyl acetate, cyclohexanone, butanol, and d-limonene). In addition, the relative sensitivities of the trigeminaland olfactory systems to these compounds are compared. In electrophysiological cross-adaptation experiments (amyl acetate vs cyclohexanone; butanol vs d-limonene), there was complete cross adaptation such that only concentrations above the background (crossa-dapting) stimulus concentration elicited responses, suggesting that chemical stimuli may stimulate trigeminal receptors nonspecifically. In behavioral experiments (amyl acetate vs cyclohexanone; butanol vs d-limonene), only animals with intact olfactory nerves could discriminate between perceptually equivalent concentrations, that is concentrations that elicited the same level of responding. Both electrophysiologically and behaviorally, the trigeminal system exhibited higher thresholds than the olfactory system. We conclude that trigeminal chemoreceptors, at least in salamanders, are unable to discriminate between these two pairs of compounds when matched for equal perceptual intensity, and that trigeminal chemoreceptors are less sensitive than olfactory receptors.Abbreviations AA amyl acetate - CH cyclohexanone - LI d-limonene - BU butanol - EOG electro-olfactogram - ISI interstim-ulus interval - ONX olfactory nerve cut - ppm parts per million (1 l of compound in vapor phase/1l of air=1 ppm)     

Tilt aftereffects generated by bilaterally symmetrical patterns

Tilt aftereffects were generated by bilaterally symmetrical dot patterns. Both expansion and contraction effects, similar in size and magnitude to effects usually reported with luminance contours, were observed after adaptation to symmetrical patterns tilted 15 deg or 75 deg respectively from a vertically oriented test. Large effects were found when both adapting and test stimuli were symmetrical patterns while smaller effects were found when the adapting stimulus was symmetrical and test stimulus was a grating. A third experiment, which manipulated the number of dots near the axis line, confirmed the above findings; expansion and contraction effects were observed again. The results of these experiments suggest that the neural mechanism underlying the perception of luminance contours may be linked to the mechanism for the detection of symmetry.     

The Effect of Stimulus Duration on Frequency-Response Functions in the Pacinian (P) Channel

Psychophysical experiments on human observers and physiological measurements on Pacinian corpuscles (PCs) isolated from cat mesentery were performed to explain certain discrepancies in the psychophysical—physiological model (Bolanowski et al., 1988) for the sense of touch in the vibrotactle Pacinian (P) channel. The model was based on correlations among the psychophysical frequency response obtained on human glabrous skin and physiological frequency-response functions measured on two PC preparations: PC fibers innervating human glabrous skin (Johansson et al., 1982) and PCs isolated from cat mesentery. The three frequency-response functions were qualitatively similar. However, the low-frequency slope for the human PC fibers differed from the slopes for the psychophysical and cat mesentery PC functions by being 3 dB/octave less steep. This discrepancy can be explained theoretically by differences in methodology involving the effect of stimulus duration and the property of temporal summation known to exist in the P channel (i.e., a 3-dB increase in sensitivity per doubling of stimulus duration). To test this, experiments were performed using two methods of stimulation: (1) a constant stimulus duration for different test frequencies, as generally used in this laboratory; and (2) a constant number of stimulus cycles (n = 5) for each test frequency as used by Johansson et al. The method of least squares was used to calculate the low-frequency (50 to 150-Hz) slopes of individual psychophysical and physiological functions. The mean slopes that resulted from using the two methods of stimulation were consistent with the theoretical expectations.     

Representation of moving stimuli by somatosensory neurons.

The findings obtained in neurophysiological and psychophysical investigations using tactile stimuli that move at constant velocity across the skin are reviewed. For certain neurons in the postcentral gyrus of the cerebral cortex (S-I) of macaque monkeys, direction of stimulus motion is a "trigger feature" i.e., moving tactile stimuli evoke vigorous discharge activity in these neurons only if the stimuli are moved in a particular direction across the receptive field. This directional selectivity is maximal when stimulus velocity is between 5 and 50 cm/sec, and falls off rapidly at lower or higher velocities. The capacity for human subjects to correctly identify the direction of stimulus motion on the skin exhibits a similar dependence on stimulus velocity. The similar effects of velocity on neural and psychophysical measures of directional sensitivity support the idea that direction of stimulus motion on the skin can only be recognized if the moving stimulus optimally activates the group of S-I neurons for which that directions of simulus motion is the trigger feature.     

Differential covariation in taste responsiveness to bitter stimuli in rats

Variation exists in the sensitivity of individual rodents and humans to different bitter tastants. An absence of uniform correlation in responsiveness to different bitter substances across individuals within a species suggests heterogeneity in the mechanisms underlying stimulus processing within this taste modality. Here, we examined taste responsiveness of individual rats to three bitter compounds (quinine hydrochloride, denatonium benzoate, and cycloheximide) in short-term lick tests to determine the magnitude of covariation among responses to these stimuli and infer commonalities in their receptor and neural mechanisms. Rats were tested with a given pair of bitter stimuli during three sessions comprising randomized trial blocks of six concentrations of each stimulus + deionized water. Psychophysical functions were generated for individual rats for respective stimulus pairs, and concentrations of each stimulus that produced equivalent lick suppression relative to water were correlated across animals. Behavioral taste responsiveness to quinine hydrochloride strongly covaried with responsiveness to denatonium benzoate (r = +0.82). Lick responsiveness to quinine was less robustly correlated with that to cycloheximide (r = +0.44), and denatonium and cycloheximide responses failed to correlate. These results imply substantial overlap in the bitter taste coding mechanisms for quinine and denatonium but some degree of independence in the mechanisms responsible for gustatory processing of cycloheximide. More generally, these data reinforce the notion that bitter taste processing is not a homogeneous event.     

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.     

Objective and subjective indexes of auditory motion processing

The objective and subjective indexes of sound stimulus discrimination have been studied in order to get insight into individual stages of signal processing in the human brain. The experiment employed two methods: electrophysiological (mismatch negativity or MMN recording) and psychophysical (two-alternative forced choice). Two types of spatial sound stimuli simulated gradual and abrupt sound motion from the head midline. The subjective discrimination between the gradual and abrupt motions was estimated as a function of the stimulus trajectory length. MMN as an objective index of spatial discrimination has been obtained in response to the subthreshold and the suprathreshold levels of psychophysical discrimination. An increase in the angular displacement of the moving stimuli resulted in an increase in both the MMN amplitude and the subjective discrimination, although their correlation remained below the significance level. The results obtained are discussed from the point of view of preconscious perception of auditory spatial information.     

Variability in 5- and 8-year-olds' memory for duration: an interfering task in temporal bisection

The purpose of this experiment was to investigate 5- and 8-year-olds' long-term memory for stimulus duration in a bisection task. Children were trained to discriminate between a short and a long standard duration presented as visual stimulus for 2 and 8 s, respectively. They had then to decide whether an intermediate stimulus duration was more similar to the short or to the long standard in two identical testing phases separated by an interfering task lasting for 15 min (immediate test vs. deferred test). The results showed that the 5- and the 8-year-olds produced orderly psychophysical functions. However, the 8-year-olds produced psychophysical functions, which increased more abruptly with the increasing stimulus duration. Nevertheless, whatever the age of children tested, the psychophysical function curves were flatter in the deferred test than in the immediate test, and more particularly in the 5-year-olds. Furthermore, the 5-year-olds produced fewer 'long' responses in the deferred than in the immediate test. Modeling of the data suggests that the variability of the representation of the standard in long-term memory was higher in the 5- than in the 8-year-olds and that the interfering task increased this memory variability.     

The taste of creatine and creatinine

Four experiments investigated the possibility of a taster-nontasterdimorphism for creatine and creatinine, and whether perceptionof these substances was related to sensitivity to phenylthiocarbamide(PTC). Threshold measurements, suprathreshold scaling of perceivedintensity, a cross-adaptation experiment and a category scalingtask produced consistent results. No relation to PTC was observed.In contrast to previous reports, no evidence of a taster-nontastereffect for creatine or creatinine was found. Creatine was anotably ineffective taste stimulus for all subjects tested.     

Effects of physiological and pharmacological stimuli on dopamine release in the rat globus pallidus

A major aspect of understanding functions of the globus pallidus (GP) within the basal ganglia is the significance of its dopamine innervation. Here, we used in vivo-microdialysis in rats to characterize pallidal dopamine release in response to a number of physiological and pharmacological stimuli known to activate dopamine neurons. Results reveal that an aversive stimulus, i.e. handling for 20 min, significantly increased dialysate dopamine in the globus pallidus to about 130% of baseline levels. Likewise, a novel and appetitive stimulus, i.e. presentation of unfamiliar, palatable food, significantly elevated pallidal dopamine to about 150% of baseline levels both in rats which did and did not consume the food reward. These findings provide evidence that increases of dopamine (DA) efflux may largely reflect stimulus saliency implicating an involvement of pallidal dopamine signalling in control of behaviour governed by salient stimuli. Results further showed that reverse microdialysis of D-amphetamine and cocaine in augmenting concentrations of 0.1-100 microM elevated dialysate dopamine in a concentration-dependent manner suggesting a role of pallidal dopamine in mediating behavioural effects of psychostimulant drugs.     

Cross-adaptation and bitterness inhibition of L-tryptophan, L-phenylalanine and urea: further support for shared peripheral physiology

A previous study investigating individuals' bitterness sensitivities found a close association among three compounds: L-tryptophan (L-trp), L-phenylalanine (L-phe) and urea (Delwiche et al., 2001, Percept. Psychophys. 63, 761-776). In the present experiment, psychophysical cross-adaptation and bitterness inhibition experiments were performed on these three compounds to determine whether the bitterness could be differentially affected by either technique. If the two experimental approaches failed to differentiate L-trp, L-phe and urea's bitterness, then we may infer they share peripheral physiological mechanisms involved in bitter taste. All compounds were intensity matched in each of 13 subjects, so the judgments of adaptation or bitterness inhibition would be based on equal initial magnitudes and, therefore, directly comparable. In the first experiment, cross-adaptation of bitterness between the amino acids was high (>80%) and reciprocal. Urea and quinine-HCl (control) did not cross-adapt with the amino acids symmetrically. In a second experiment, the sodium salts, NaCl and Na gluconate, did not differentially inhibit the bitterness of L-trp, L-phe and urea, but the control compound, MgSO(4), was differentially affected. The bitter inhibition experiment supports the hypothesis that L-trp, L-phe and urea share peripheral bitter taste mechanisms, while the adaptation experiment revealed subtle differences between urea and the amino acids indicating that urea and the amino acids activate only partially overlapping bitter taste mechanisms.