The nature and duration of adaptation following long-term odor exposure

Any individual living or working in an odorous environment can experience changes in odor perception, some of which are long lasting. Often, these individuals report a significant reduction in the perception of an odor following long-term exposure to that odor (adaptation). Yet, most experimental analyses of olfactory adaptation use brief odorant exposures which may not typify real-world experiences. Using a procedure combining long-term odor exposure in a naturalistic setting with psychophysical tests in the laboratory, we present evidence to show that reduced odor intensity following long-term exposure is accompanied by odorant-specific shifts in threshold. Subjects were exposed continuously to one of two odorants while in their home for a period of 2 weeks. Exposure produced an odorant-specific reduction in sensitivity and perceived intensity compared with preexposure baselines: Detection thresholds for the adapting odorant were elevated following exposure and perceived intensity ratings for weak concentrations were reduced. For most individuals, reduced sensitivity to the test odorant was still evident up to 2 weeks following the last exposure. The persistence of the change, as evidenced by the duration of recovery from adaptation, distinguishes this phenomenon from the adaptation seen following shorter exposures and highlights the need for the study of exposure durations that are more similar to real-world exposures.     

A comparison of olfactory adaptation among seven odorants and their relationship with several physicochemical properties

A study was made comparing olfactory sensitivity before and after 100 sec of adaptation for stimuli representing several odorant classes with varying physicochemical properties. Measures taken were absolute detection thresholds and magnitude estimates of suprathreshold stimuli. Significant rank-order correlation coefficients (<.90) were found between nonadapted and adapted absolute detection thresholds, vapor pressure, and percent adaptation.     

Olfactory adaptation and recovery in man as measured by two psychophysical techniques

Olfactory adaptation and recovery was investigated in man, using two psychophysical procedures: modified category scaling and threshold detection. Both procedures yielded similar qualitative information regarding loss and recovery of olfactory sensitivity as a function of time and concentration of adapting stimuli. However, quantitative differences were observed that could be partially attributed to artifacts inherent in each procedure. Often more than 50% adaptation (and recovery) occurred within the first 2 min with either test procedure. In all experiments the rate of adaptation and recovery was greater at the higher of two adapting concentrations (10 × and 20 × the detection threshold I_t). Recovery occurred more rapidly than adaptation. The usefulness of both techniques is discussed in terms of the overall problem of characterizing the olfactory adaptation and recovery process in man.     

Paradoxical olfactory function in combat Veterans: The role of PTSD and odor factors

Stress- and trauma-related disorders, including posttraumatic stress disorder (PTSD), are characterized by an increased sensitivity to threat cues. Given that threat detection is a critical function of olfaction and that combat trauma is commonly associated with burning odors, we sought a better understanding of general olfactory function as well as response to specific trauma-related (i.e. burning) odors in combat-related PTSD. Trauma-exposed combat Veterans with (n = 22) and without (n = 25) PTSD were assessed for general and specific odor sensitivities using a variety of tools. Both groups had similar general odor detection thresholds. However, the combat Veterans with PTSD, compared to combat Veterans with comparable trauma exposure but without PTSD, had increased ratings of odor intensity, negative valence, and odor-triggered PTSD symptoms, along with a blunted heart rate in response to burning rubber odor. These findings are discussed within the context of healthy versus pathological changes in olfactory processing that occur over time after psychological trauma.     

Changes in absolute detection threshold and in subjective intensity of suprathreshold stimuli during olfactory adaptation and recovery

Olfactory adaptation and recovery to methyl isobutyl ketone at a concentration 10 times the absolute detection threshold (I_{t o}) was intensively studied in two human Ss. A combined psychophysical procedure was used that allowed comparisons of changes in threshold (I_t) with changes in the subjective intensity of suprathreshold stimuli. Information was also obtained on the effect of the adapting stimulus on the psychophysical power function for this odorant. A threshold detection procedure was used to estimate changes in I_t; an unstructured magnitude-estimation procedure was used to monitor changes in the subjective intensity of suprathreshold stimuli and the psychophysical power function. The data provided additional information on the behavioral course of olfactory adaptation and recovery and suggested that this combined method can be used profitably for further investigations of this kind. Complementary to the work of Cain and Engen (1969), the results suggested an increase in the exponent of the power function with increasing adaptation.     

The role of tone duration in dichotic temporal order judgment

Over the past two decades, a number of studies have been based on the dichotic temporal order judgment (TOJ) paradigm, to compare auditory temporal processing in various subpopulations to that of young, normal controls. The reported estimates of dichotic TOJ thresholds, expressed as the interstimulus intervals (ISIs) for 75 % accuracy among the controls, have varied. In the present study, we examined the influence of tone duration, within the 10- to 40-msec range, on dichotic TOJ accuracy and threshold. The results indicated that increases in either ISI or tone duration increased dichotic TOJ accuracy similarly, implying that changes in tone duration may affect dichotic TOJs simply by adding to the delay between onset of the tone at the lead ear and onset of the tone at the lag ear. The dichotic TOJ thresholds from three published studies that had used tones as stimuli and the dichotic thresholds from the present study all fell within 0.5 standard deviations from the theoretical line of a 10-msec reduction in threshold for every 10-msec increase in tone duration. When the dichotic TOJ threshold data from the present study and from the published studies were converted to stimulus onset asynchronies (SOAs) and plotted as a function of tone duration, they fell very close to or on a zero-slope line, indicating that when ISI and duration are summed to yield an SOA, dichotic TOJ thresholds are invariant to tone duration within the range of 10–40 msec.     

Invariance of odor strength with sniff vigor: an olfactory analogue to size constancy

Previous evidence has shown that detection threshold in humans and olfactory neural discharge rate in animal preparations both depend on flow rate of odorous vapor. But no data have been reported that show the effects of flow rate in humans on perceived odor strength at suprathreshold intensities. Subjects learned to inspire at two flow rates, one twice as great as the other, by adjusting (on a cathode ray tube) the transduced trace of a sniff-produced pressure change to match either of two target contours. They then made magnitude estimations of odor strength, while producing either weak or strong sniffs, for odorants presented over a wide range of concentrations via a specially designed sniff-bottle system. The odorant, diluted in diethyl phthalate, was n-butanol in two experiments and n-amyl acetate in two others. Subject-controlled flow rate had no effect on odor strength for either odorant. There was an apparent contradiction between these data and those on neural discharge rate that may, however, be resolved by adopting an odor constancy model: When sniff intensity varies during the olfactory exploration of an odor source, information about the rate at which odorant molecules are established at the receptor site is combined with information about sniff vigor so that the resulting percept is of invariant odor strength.     

Synaptic adaptation and odor-background segmentation

Habituation is a form of non-associative memory that plays an important role in filtering stable or redundant inputs. The present study examines the contribution of habituation and cortical adaptation to odor-background segmentation. Segmentation of target odorants from background odorants is a fundamental computational requirement for the olfactory system. Recent electrophysiological data have shown that odor specific adaptation in piriform cortex neurons, mediated at least partially by synaptic adaptation between the olfactory bulb outputs and piriform cortex pyramidal cells, may provide an ideal mechanism for odor-background segmentation. This rapid synaptic adaptation acts as a filter to enhance cortical responsiveness to changing stimuli, while reducing responsiveness to static, potentially background stimuli. Using previously developed computational models of the olfactory system, we here show how synaptic adaptation at the olfactory bulb input to the piriform cortex, as demonstrated electrophysiologically, creates odor specific adaptation. We show how this known feature of olfactory cortical processing can contribute to adaptation to a background odor and to odor-background segmentation. We then show in a behavioral experiment that the odor-background segmentation is perceptually important and functions at the same time-scale as the synaptic adaptation observed between the olfactory bulb and cortex.     

Very Early Decline in Recognition Memory for Odors in Alzheimer’s Disease

When in the progression of Alzheimer’s disease (AD) a decline in detection sensitivity and recognition memory for odors begins, and whether a fast progression in dementia is related to a fast loss in odor sensitivity, was determined by studying 78 patients with probable AD, categorized as very mild, mild, or moderate in degree of dementia, and 78 controls. Taste and vision were studied for comparison. Whereas the earliest decline in odor sensitivity and visual memory was found in mild patients, a decline in odor memory was found in the very mild patients. Similar taste thresholds in the four groups suggest that the poor odor detectability in AD was not related to task demands in threshold testing. A fast progression in dementia was also found to be related to a fast loss in odor sensitivity. The findings imply that memory-based olfactory tests may contribute to early diagnosis of AD.     

Tests of human olfactory function: Principal components analysis suggests that most measure a common source of variance

It is not known whether nominally different olfactory tests actually measure dissimilar perceptual attributes. In this study, we administered nine olfactory tests, including tests of odor identification, discrimination, detection, memory, and suprathreshold intensity and pleasantness perception, to 97 healthy subjects. Aprincipal components analysis performed on the intercorrelation matrix revealed four meaningful components. The first was comprised of strong primary loadings from most of the olfactory test measures, whereas the second was comprised of primary loadings from intensity ratings given to a set of suprathreshold odorant concentrations. The third and fourth components had primary loadings that reflected, respectively, mean suprathreshold pleasantness ratings and a response bias measure derived from a yes/no odor identification signal detection task. In an effort to adjust for potential confounding influences of age, gender, smoking, and years of schooling on the component structure, a matrix of residuals from a multiple regression analysis, which included these variables, was also analyzed. A similar component pattern emerged. Overall, these findings suggest, in healthy subjects spanning a wide age range, that (1) a number of nominally distinct tests of olfactory function are measuring a common source of variance, and (2) some suprathreshold odor intensity and pleasantness rating tests may be measuring sources of variance different from this common source.     

Temporal and spatial characteristics of attention to facilitate manual and eye-movement responses

Previous investigation found that the speed of saccadic eye movements is enhanced when a temporal interval (gap) is introduced between the disappearance of a foveal fixation mark and the appearance of a peripheral target (the gap paradigm). Attention was shown to be involved in the gap paradigm. Here, we investigated relevant temporal and spatial characteristics of attention, manipulating central fixation marks and peripheral targets. Results from three experiments indicate that (i) the speed of manual and eye-movement detection is accelerated when a fixation mark changes abruptly (in less than 100 ms) before its termination in the gap paradigm; (ii) the speed is further accelerated when a peripheral target location is pre-cued; (iii) sufficient time for fixation (1000 ms) is necessary for the facilitation. These results suggest that fast and transient attention at the fixation spot facilitates attentional disengagement process that urges a spatial-orienting mechanism. Sustained attention is required in the engagement process during the fixation.     

Subliminal smells can guide social preferences

It is widely accepted that unconscious processes can modulate judgments and behavior, but do such influences affect one's daily interactions with other people? Given that olfactory information has relatively direct access to cortical and subcortical emotional circuits, we tested whether the affective content of subliminal odors alters social preferences. Participants rated the likeability of neutral faces after smelling pleasant, neutral, or unpleasant odors delivered below detection thresholds. Odor affect significantly shifted likeability ratings only for those participants lacking conscious awareness of the smells, as verified by chance-level trial-by-trial performance on an odor-detection task. Across participants, the magnitude of this priming effect decreased as sensitivity for odor detection increased. In contrast, heart rate responses tracked odor valence independently of odor awareness. These results indicate that social preferences are subject to influences from odors that escape awareness, whereas the availability of conscious odor information may disrupt such effects.     

Auditory discrimination of anisochrony: influence of the tempo and musical backgrounds of listeners

This study explored the influence of several factors, physical and human, on anisochrony's thresholds measured with an adaptive two alternative forced choice paradigm. The effect of the number and duration of sounds on anisochrony discrimination was tested in the first experiment as well as potential interactions between each of these factors and tempo. In the second experiment, the tempo or the inter onset interval (IOI) was varied systematically between 80 and 1000 ms. The results showed that just noticeable differences increase linearly and proportionally with IOI in accordance with Weber's law except for quickest tempo (IOI of 80 ms). The third experiment investigated the role of musical training on anisochrony thresholds obtained for different IOI. It focused on differential effects of musical experiences by comparing non-musicians, instrumentalists, and percussionists thresholds. The results of the present study replicated the findings of previous experiments regarding the adequacy of Weber's law for slow rhythm and provided evidence for its departure for fast tempos. Moreover, thresholds from percussionists seem distinguishable from the ones of other listeners by their highest sensitivity to temporal shifts suggesting therefore the necessity to control the nature of musical experiences. The results are discussed according to current models of time perception.     

Olfactory detection thresholds in man under conditions of rest and exercise

Detection thresholds for seilen odorants were measured under conditions of rest and exercise. The test panel comprised six males with previous experience in odor detection. Exercise was maintained constant by using a modified bicycle positioned within the context of an air-dilution olfactometer. The effects of exercise were idiosyncratic for Ss: odor sensitivity was increased in some Ss, decreased in some, and not changed in others. Results of these experiments are discussed in terms of intra-subject variance within sessions and from session to session. In addition, two test techniques—one a fixed series and the other a tracking procedure—were evaluated for rapid estimation of an odor threshold. The tracking procedure facilitated prompt detection of changes in sensitivity for individual Ss.     

Acetylcholine and olfactory perceptual learning

Olfactory perceptual learning is a relatively long-term, learned increase in perceptual acuity, and has been described in both humans and animals. Data from recent electrophysiological studies have indicated that olfactory perceptual learning may be correlated with changes in odorant receptive fields of neurons in the olfactory bulb and piriform cortex. These changes include enhanced representation of the molecular features of familiar odors by mitral cells in the olfactory bulb, and synthetic coding of multiple coincident odorant features into odor objects by cortical neurons. In this paper, data are reviewed that show the critical role of acetylcholine (Ach) in olfactory system function and plasticity, and cholinergic modulation of olfactory perceptual learning at both the behavioral and cortical level.     

Anatomy of a flash. 1. Two-peak masking and a temporal filling-in

A modified paradigm of Crawford masking was used to link masking to brightness fluctuation, as distinct from flash brightness. Thresholds were measured for a 10 ms incremental pulse (the 'probe') presented before, during, or after a 500 ms pulse (the 'flash'). Both pulses were spatially coextensive with the background field, thus the criterion for probe detection was purely temporal. The flash occurred either in the tested eye, the opposite eye, or in both eyes. In all conditions, masking was strongly bimodal: thresholds peaked near flash onset and flash offset. The flash was perceived as a unitary event. Bimodal masking is attributed to cortical on-and off-effects, as (i) dichoptic masking was strong and (ii) the same incremental probe was masked by either incremental or decremental flashes. Strikingly, monocular probe thresholds were about equally elevated by binocular as by monocular flashes, although the binocular flashes were brighter. Therefore, some monocular features can be preserved in the larger net binocular response. A general conclusion is that masking depends on the same transient neural responses that bring about a brightness fluctuation, whereas the appearance of the flash as a single event, a unitary change of brightness, depends on a different mechanism, perhaps a sustained response that performs a temporal filling-in.     

Nasal trigeminal chemosensitivity across the adult life span

Individuals can determine the side of the nose that receives an odorant during unilateral presentation (lateralize) if endings of the trigeminal nerve are stimulated. By using psychophysical methods, olfactory detection and trigeminal lateralization thresholds for 1-butanol were obtained from 142 individuals ranging in age from 20 to 89 year. Sensitivity in both chemosensory pathways declined with advancing age, especially in people older than 60 years.     

Effects of Visual Error Timing on Smooth Pursuit Gain Adaptation in Humans

Smooth pursuit (SP) is one of the precise oculomotor behaviors when tracking a moving object. Adaptation of SP is based on a visual-error driven motor learning process associated with predictable changes in the visual environment. Proper timing of a sensory signal is an important factor for adaptation of fine motor control. In this study, we investigated whether visual error timing affects SP gain adaptation. An adaptive change in SP gain is produced experimentally by repeated trials of a step-ramp tracking with 2 different velocities (double-velocity paradigm). The authors used the double-velocity paradigm where target speed changes 400 or 800 ms after the target onset. The results show that SP gain changed in a certain time window following adaptation. The authors suggest that SP adaptation shown in this study is associated with timing control mechanisms.     

Odorization of a novel object can influence infant's exploratory behavior in unexpected ways

Although much is known about the development of object exploration during infancy, it remains to be understood whether and how olfaction can influence infants' interactions with novel objects. To address these issues, sixteen infants aged 7-15 months were videotaped during two consecutive 5-min free play sessions with a scented or an unscented version of visually similar objects. Results indicate that adding an odor to a novel object influenced the infants' behavior: the infants exhibited more and longer manipulations and mouthing of the unscented object than of the scented object. The differential responsiveness to the scented, relative to the unscented, object was noted after a 2-min delay following test onset, suggesting that in the present conditions infants do not immediately detect or react to the added odor. It may be concluded that infants do detect an odorant added on a novel object, show odor-based discrimination of visually similar objects, and express withdrawal of the scented, relative to the unscented, object. The implications of these findings for understanding how infants use their senses, namely their olfactory sense, in early exploratory behavior are discussed.     

The effects of thematic relations on picture naming abilities across the lifespan

A picture–word interference paradigm tracked patterns of activation during picture naming in 87 individuals (age range 17–80 years old). Distractor words were presented at stimulus onset asynchronies (SOAs) of ?200, ?100, and 0 ms bearing a has a-, location, or no relationship to the picture. Analyses of group naming reaction times revealed significant facilitation effects for both semantic relation types for all age groups. Analyses of temporal patterns of activation revealed significant effects primarily at SOAs of ?200 and ?100 ms. These findings provide evidence that both thematic relations are particularly salient in how semantic knowledge is organized, and that the patterns of effects from these semantic relations remain the same as one ages.