Stimuli Of Different Durations Research Articles

DYNAMICS OF THE PARIETAL-OCCIPITAL ALPHA RHYTHM ACTIVITY DURING COMPARISON OF VISUAL STIMULI DURATIONS

This research is aimed at studying the dynamics of the parietal-occipital alpha rhythm in its connection with the process of stimuli duration comparison. EEG study was conducted in which participants (n = 48) were asked to compare pairs of visual stimuli of different durations ranging from 3.2 to 6.4 s. The time-frequency analysis of the EEG was carried out in the range of 8–12 Hz. The power of alpha rhythm increases from the stimulus onset to the middle of its presentation, but then its dynamic depends on the stimulus duration: it further increases for short durations (3.2, 3.6, 4.0 s), stays the same for middle durations (4.4, 4.8, 5.2 s) and decreases for long durations (5.6, 6.0, 6.4 s). The relative decrease of alpha power for long stimuli in relation to the short ones was related to subjective perception of time. The results are discussed from the point of view of the “dual klepsydra” model: it is assumed that alpha rhythm acts as an electrophysiological correlate of the functioning of “neural accumulators” associated with the subjective passage of time.

Binaural masking level difference for pure tone signals

The binaural masking level difference (BMLD) is a psychoacoustic method to determine binaural interaction and central auditory processes. The BMLD is the difference in hearing thresholds in homophasic and antiphasic conditions. The duration, phase and frequency of the stimuli can affect the BMLD. The main aim of the study is to evaluate the BMLD for stimuli of different durations and frequencies which could also be used in future electrophysiological studies. To this end we developed a GUI to present different frequency signals of variable duration and determine the BMLD. Three different durations and five different frequencies are explored. The results of the study confirm that the hearing threshold for the antiphasic condition is lower than the hearing threshold for the homophasic condition and that differences are significant for signals of 18ms and 48ms duration. Future objective binaural processing studies will be based on 18ms and 48ms stimuli with the same frequencies as used in the current study.

SSVEP-EEG Denoising via Image Filtering Methods.

Steady-state visual evoked potential (SSVEP) is widely used in electroencephalogram (EEG) control, medical detection, cognitive neuroscience, and other fields. However, successful application requires improving the detection performance of SSVEP signal frequency characteristics. Most strategies to enhance the signal-to-noise ratio of SSVEP utilize application of a spatial filter. Here, we propose a method for image filtering denoising (IFD) of the SSVEP signal from the perspective of image analysis, as a preprocessing step for signal analysis. Arithmetic mean, geometric mean, Gaussian, and non-local means filtering methods were tested, and the experimental results show that image filtering of SSVEP cannot effectively remove the noise. Thus, we proposed a reverse solution in which the SSVEP noise signal was obtained by image filtering, and then the noise was subtracted from the original signal. Comparison of the recognition accuracy of the SSVEP signal before and after denoising was used to evaluate the denoising performance for stimuli of different duration. After IFD processing, SSVEP exhibited higher recognition accuracy, indicating the effectiveness of this proposed denoising method. Application of this method improves the detection performance of SSVEP signal frequency characteristics, combines image processing and brain signal analysis, and expands the research scope of brain signal analysis for widespread application.

Attractor-state itinerancy in neural circuits with synaptic depression

Neural populations with strong excitatory recurrent connections can support bistable states in their mean firing rates. Multiple fixed points in a network of such bistable units can be used to model memory retrieval and pattern separation. The stability of fixed points may change on a slower timescale than that of the dynamics due to short-term synaptic depression, leading to transitions between quasi-stable point attractor states in a sequence that depends on the history of stimuli. To better understand these behaviors, we study a minimal model, which characterizes multiple fixed points and transitions between them in response to stimuli with diverse time- and amplitude-dependencies. The interplay between the fast dynamics of firing rate and synaptic responses and the slower timescale of synaptic depression makes the neural activity sensitive to the amplitude and duration of square-pulse stimuli in a nontrivial, history-dependent manner. Weak cross-couplings further deform the basins of attraction for different fixed points into intricate shapes. We find that while short-term synaptic depression can reduce the total number of stable fixed points in a network, it tends to strongly increase the number of fixed points visited upon repetitions of fixed stimuli. Our analysis provides a natural explanation for the system’s rich responses to stimuli of different durations and amplitudes while demonstrating the encoding capability of bistable neural populations for dynamical features of incoming stimuli.

Thermal sensitivity in humans at the depth of thermal receptor endings beneath the skin: validation of a heat transfer model of the skin using high-temporal resolution stimuli.

The bioheat transfer equation predicts temperature distribution in living tissues such as the skin. This study aimed at psychophysically validating this model in humans. Three experiments were performed. In the first, participants were asked to judge the thermal intensity of stimuli with combinations of intensity and duration that yielded, according to the model, identical temperatures at the thermoreceptor's depth. In experiment 2, participants' thermal detection thresholds for stimuli of different durations were measured to verify whether these thresholds correspond, according to the model, to equivalent temperatures at the thermoreceptor's location. In experiment 3, an alternative forced choice method was used, in which subjects indicated which of the two consecutive thermal stimulations was more intense. The model predicted results that agreed with subjects' perceptions. Participants judged stimuli of different combinations of intensities and durations yielding identical temperature at the receptor level as having equivalent intensity. Moreover, although cold detection thresholds for stimuli of different durations differed for temperatures of the stimulating probe, stimulations using the model's parameters showed equivalence at the depth of the thermal receptors. Furthermore, stimuli with temperature/duration combinations for which the model predicts temperature equivalence at the depth of the receptors corresponded to subjective equalization. These findings indicate that heat transfer models provide good estimates of temperatures at the thermal receptors. Use of these models may facilitate comparisons among studies using different stimulation devices and may facilitate the establishment of standards involving all stimulation parameters.

Impaired Motor Learning Following a Pain Episode in Intact Rats.

Motor learning and pain are important factors influencing rehabilitation. Despite being mostly studied independently from each other, important interactions exist between them in the context of spinal cord injury, whether to the spinal cord or the body. Ongoing or recent past episodes of nociceptive activity can prevent motor learning in spinalized rats. In intact animals, it has been proposed that supraspinal activity could counter the repressive effect of nociception on motor system plasticity, but this has not yet been verified in behavioral conditions. The aim of this study was to test whether a recent episode of nociception affects subsequent motor learning in intact animals. We trained rodents to walk on a custom-made horizontal ladder. After initial training, the rats underwent a week-long rest, during which they were randomly assigned to a control group, or one out of two pain conditions. Nociceptive stimuli of different durations were induced through capsaicin or Complete Freund's Adjuvant injections and timed so that the mechanical hypersensitivity had entirely subsided by the end of the resting period. Training then resumed on a modified version of the horizontal ladder. We evaluated the animals' ability to adapt to the modified task by measuring their transit time and paw misplacements over 4 days. Our results show that prior pain episodes do affect motor learning in neurologically intact rats. Motor learning deficits also seem to be influenced by the duration of the pain episode. Rats receiving a subcutaneous injection of capsaicin displayed immediate signs of mechanical hypersensitivity, which subsided rapidly. Nonetheless, they still showed learning deficits 24 h after injection. Rats who received a Complete Freund's Adjuvant injection displayed mechanical hypersensitivity for up to 7 days during the resting period. When trained on the modified ladder task upon returning to normal sensitivity levels, these rats exhibited more prolonged motor learning deficits, extending over 3 days. Our results suggest that prior pain episodes can negatively influence motor learning, and that the duration of the impairment relates to the duration of the pain episode. Our results highlight the importance of addressing pain together with motor training after injury.

Behavioural benefits of multisensory processing in ferrets.

Enhanced detection and discrimination, along with faster reaction times, are the most typical behavioural manifestations of the brain's capacity to integrate multisensory signals arising from the same object. In this study, we examined whether multisensory behavioural gains are observable across different components of the localization response that are potentially under the command of distinct brain regions. We measured the ability of ferrets to localize unisensory (auditory or visual) and spatiotemporally coincident auditory–visual stimuli of different durations that were presented from one of seven locations spanning the frontal hemifield. During the localization task, we recorded the head movements made following stimulus presentation, as a metric for assessing the initial orienting response of the ferrets, as well as the subsequent choice of which target location to approach to receive a reward. Head‐orienting responses to auditory–visual stimuli were more accurate and faster than those made to visual but not auditory targets, suggesting that these movements were guided principally by sound alone. In contrast, approach‐to‐target localization responses were more accurate and faster to spatially congruent auditory–visual stimuli throughout the frontal hemifield than to either visual or auditory stimuli alone. Race model inequality analysis of head‐orienting reaction times and approach‐to‐target response times indicates that different processes, probability summation and neural integration, respectively, are likely to be responsible for the effects of multisensory stimulation on these two measures of localization behaviour.

Parametric trial-by-trial prediction of pain by easily available physiological measures

Pain is commonly assessed by subjective reports on rating scales. However, in many experimental and clinical settings, an additional, objective indicator of pain is desirable. In order to identify an objective, parametric signature of pain intensity that is predictive at the individual stimulus level across subjects, we recorded skin conductance and pupil diameter responses to heat pain stimuli of different durations and temperatures in 34 healthy subjects. The temporal profiles of trial-wise physiological responses were characterized by component scores obtained from principal component analysis. These component scores were then used as predictors in a linear regression analysis, resulting in accurate pain predictions for individual trials. Using the temporal information encoded in the principal component scores explained the data better than prediction by a single summary statistic (ie, maximum amplitude). These results indicate that perceived pain is best reflected by the temporal dynamics of autonomic responses. Application of the regression model to an independent data set of 20 subjects resulted in a very good prediction of the pain ratings demonstrating the generalizability of the identified temporal pattern. Utilizing the readily available temporal information from skin conductance and pupil diameter responses thus allows parametric prediction of pain in human subjects.

Persistent perceptual delay for head movement onset relative to auditory stimuli of different durations and rise times

The perception of simultaneity between auditory and vestibular information is crucially important for maintaining a coherent representation of the acoustic environment whenever the head moves. It has been recently reported, however, that despite having similar transduction latencies, vestibular stimuli are perceived significantly later than auditory stimuli when simultaneously generated. This suggests that perceptual latency of a head movement is longer than a co-occurring sound. However, these studies paired a vestibular stimulation of long duration (~1 s) and of a continuously changing temporal envelope with a brief (10–50 ms) sound pulse. In the present study, the stimuli were matched for temporal envelope duration and shape. Participants judged the temporal order of the two stimuli, the onset of an active head movement and the onset of brief (50 ms) or long (1,400 ms) sounds with a square- or raised-cosine-shaped envelope. Consistent with previous reports, head movement onset had to precede the onset of a brief sound by about 73 ms in order for the stimuli to be perceived as simultaneous. Head movements paired with long square sounds (~100 ms) were not significantly different than brief sounds. Surprisingly, head movements paired with long raised-cosine sound (~115 ms) had to be presented even earlier than brief stimuli. This additional lead time could not be accounted for by differences in the comparison stimulus characteristics (temporal envelope duration and shape). Rather, differences between sound conditions were found to be attributable to variability in the time for head movement to reach peak velocity: the head moved faster when paired with a brief sound. The persistent lead time required for vestibular stimulation provides further evidence that the perceptual latency of vestibular stimulation is greater than the other senses.

Persistent perceptual delay for head movement onset relative to auditory stimuli of different duration and rise times

The perception of simultaneity between auditory and vestibular information is crucially important for maintaining a coherent representation of the acoustic environment whenever the head moves. Yet, despite similar transduction latencies, vestibular stimuli are perceived significantly later than auditory stimuli when simultaneously generated (Barnett-Cowan and Harris, 2009, 2011). However, these studies paired a vestibular stimulation of long duration (∼1 s) and of a continuously changing temporal envelope with brief (10–50 ms) sound pulses. In the present study the stimuli were matched for temporal envelope. Participants judged the temporal order of the onset of an active head movement and of brief (50 ms) or long (1400 ms) sounds with a square or raised-cosine shaped envelope. Consistent with previous reports, head movement onset had to precede the onset of a brief sound by about 73 ms in order to be perceived as simultaneous. Head movements paired with long square sounds (∼100 ms) were not significantly different than brief sounds. Surprisingly, head movements paired with long raised-cosine sound (∼115 ms) had to be presented even earlier than brief stimuli. This additional lead time could not be accounted for by differences in the comparison stimulus characteristics (duration and temporal envelope). Rather, differences among sound conditions were found to be attributable to variability in the time for head movement to reach peak velocity: the head moved faster when paired with a brief sound. The persistent lead time required for vestibular stimulation provides further evidence that the perceptual latency of vestibular stimulation is larger compared to auditory stimuli.

Detection performance for short visual stimuli depends on the duration of co-occuring auditory stimuli

There is converging evidence that the duration of an auditory event can affect the perceived duration of a co-occurring visual event. When a brief visual stimulus is accompanied by a longer auditory stimulus, the perceived visual duration stretches. If this reflects a genuine sustain of visual stimulus perception, it should result in enhanced perception of non-temporal visual stimulus qualities. To test this hypothesis, in a temporal two-alternative forced choice task, 28 participants were asked to indicate whether a short (∼24 ms), peri-threshold, visual stimulus was presented in the first or in the second of two consecutive displays. Each display was accompanied by a sound of equal or longer duration (36, 48, 60, 72, 84, 96, 190 ms) than the visual stimulus. As a control condition, visual stimuli of different durations (matching auditory stimulus durations) were presented alone. We predicted that visual detection can improve as a function of sound duration. Moreover, if the expected cross-modal effect reflects sustained visual perception it should positively correlate with the improvement observed for genuinely longer visual stimuli. Results showed that detection sensitivity (d′) for the 24 ms visual stimulus was significantly enhanced when paired with longer auditory stimuli ranging from 60 to 96 ms duration. The visual detection performance dropped to baseline levels with 190 ms sounds. Crucially, the enhancement for auditory durations 60–96 ms significantly correlates with the d′ enhancement for visual stimuli lasting 60–96 ms in the control condition. We conclude that the duration of co-occurring auditory stimuli not only influences the perceived duration of visual stimuli but reflects a genuine sustain in visual perception.

Five choice serial reaction time performance in the HdhQ92 mouse model of Huntington's disease

Huntington's disease is an autosomal dominant genetic disorder, with motor, cognitive and psychiatric symptoms. To date there is no cure. In order to understand better this disease and to develop novel treatments, many genetically modified animal models of Huntington's disease have been created. However, to utilize these models fully, appropriate functional assays need to be developed for behavioural assessments of the mice. Various facets of attention have been reported to be affected in Huntington's disease patients, and the Hdh(Q92/Q92) mice have been shown to have deficits on operant tasks which have attentional components. In the present study, the Hdh(Q92/Q92) mouse model is assessed on a well established test of attentional function, the operant 5-choice serial reaction time task (5-CSRT), in which the mice must respond with a nose poke to light stimuli presented randomly across a 5 hole light array to receive a reward. In the present paper, the Hdh(Q92/Q92) mice exhibited deficits on the 5-CSRT when pseudorandomly presented with stimuli of different durations. However, alterations in the pacing of the task, therefore requiring an increase in sustained attention, did not affect the Hdh(Q92/Q92) mice more than their wildtype littermates. This study indicates that the Hdh(Q92/Q92) mice may have deficits in aspects of attentional function, in particular disruption in the ability to maintain attention in the visuospatial domain, suggesting that this knock-in mouse model of Huntington's disease may be a relevant model of the disease for the testing of novel therapeutic interventions.

Perceived realism of dynamic facial expressions of emotion: Optimal durations for the presentation of emotional onsets and offsets

The presentation of facial displays of emotions is an important method in emotion-recognition studies in various basic and applied settings. This study intends to make a methodological contribution and investigates the perceived realism of dynamic facial expressions for six emotions (fear, sadness, anger, happiness, disgust, and surprise). We presented dynamic displays of faces evolving from a neutral to an emotional expression (onsets) and faces evolving from an emotional expression to a neutral one (offsets). Participants rated the perceived realism of stimuli of different durations (240–3040 ms) and adjusted the duration of each sequence until they perceived it as maximally realistic. Durations perceived as most realistic are reported for each emotion, providing an important basis for the construction of dynamic facial stimuli for future research.

Non-linear population firing rates and voltage sensitive dye signals in visual areas 17 and 18 to short duration stimuli.

Visual stimuli of short duration seem to persist longer after the stimulus offset than stimuli of longer duration. This visual persistence must have a physiological explanation. In ferrets exposed to stimuli of different durations we measured the relative changes in the membrane potentials with a voltage sensitive dye and the action potentials of populations of neurons in the upper layers of areas 17 and 18. For durations less than 100 ms, the timing and amplitude of the firing and membrane potentials showed several non-linear effects. The ON response became truncated, the OFF response progressively reduced, and the timing of the OFF responses progressively delayed the shorter the stimulus duration. The offset of the stimulus elicited a sudden and strong negativity in the time derivative of the dye signal. All these non-linearities could be explained by the stimulus offset inducing a sudden inhibition in layers II–III as indicated by the strongly negative time derivative of the dye signal. Despite the non-linear behavior of the layer II–III neurons the sum of the action potentials, integrated from the peak of the ON response to the peak of the OFF response, was almost linearly related to the stimulus duration.

Source of nonlinearity of the BOLD response revealed by simultaneous fMRI and NIRS

The nonlinearity of the blood oxygenation level-dependent (BOLD) response to stimuli of different duration, particularly those of short duration, has been well studied by functional magnetic resonance imaging (fMRI). This nonlinearity is assumed to be due to neural adaptation and the nonlinearity of the response in the oxygen extraction fraction (OEF); the latter has not been examined quantitatively in humans. To evaluate how the OEF response contributes to the nonlinearity of the BOLD response to neural activity, we used simultaneous fMRI and near-infrared spectroscopy (NIRS). The responses to visual stimuli of four different durations were measured as changes in the BOLD signal and the NIRS-derived hemoglobin concentrations. The hemodynamic response nonlinearity was quantified using an impulse response function model with saturation nonlinearity scaling in the response amplitude, assuming that the unknown neural adaptation parameters varied within a physiologically feasible range. Independent of the degree of neural adaptation, the BOLD response consistently showed saturation nonlinearity similar to that of the OEF response estimated from the NIRS measures, the nonlinearity of which was greater than that of the response in the total hemoglobin concentration representing the cerebral blood volume (CBV). We also found that the contribution of the OEF response to the BOLD response was four to seven times greater than the contribution of the CBV response. Thus, we conclude that the nonlinearity of the BOLD response to neural activity originates mainly from that of the OEF response.

Sequential actions of ERK1/2 on the AP‐1 transcription factor allow temporal integration of metabolic signals in pancreatic β cells

The AP-1 transcription factor composed of fos and jun gene products mediates transcriptional responses to hormonal and metabolic stimulations of pancreatic beta cells. Here, we investigated the mechanisms that dynamically control expression of AP-1 subunit proteins. In MIN6 cells, glucose and GLP-1 raised c-FOS protein with biphasic kinetics, an initial peak being followed by a plateau that persisted as long as stimuli were maintained. ERK1/2 activation paralleled c-FOS expression. Whereas initial induction of c-FOS protein required ERK1/2-dependent activation of c-fos transcription and de novo protein synthesis, persistent accumulation of c-FOS under sustained stimulation did not. Indeed, dependent on ERK1/2 activation, c-FOS accumulated in its hyperphosphorylated form protected from degradation through the proteasome pathway. The implication of ERK1/2 in the accumulation of c-FOS protein was confirmed in rat primary beta cells, and the functional consequences of this mechanism were demonstrated with DNA-binding and reporter assays. Altogether these findings reveal a sequential regulation of AP-1 by ERK1/2, which initially increases transcription of c-fos and, if stimulation persists, stabilizes freshly synthesized c-FOS protein to efficiently activate the transcription of AP-1-regulated genes. This ERK1/2-AP-1 module can function as a temporal integrator converting metabolic stimuli of different durations into differential transcriptional outputs.

Phase and Period Responses of the Circadian System of Mice (Mus musculus) to Light Stimuli of Different Duration

To understand entrainment of circadian systems to different photoperiods in nature, it is important to know the effects of single light pulses of different durations on the free-running system. The authors studied the phase and period responses of laboratory mice (C57BL6J//OlaHsd) to single light pulses of 7 different durations (1, 3, 4, 6, 9, 12, and 18 h) given once per 11 days in otherwise constant darkness. Light-pulse duration affected both amplitude and shape of the phase response curve. Nine-hour light pulses yielded the maximal amplitude PRC. As in other systems, the circadian period slightly lengthened following delays and shortened following advances. The authors aimed to understand how different parts of the light signal contribute to the eventual phase shift. When PRCs were plotted using the onset, midpoint, and end of the pulse as a phase reference, they corresponded best with each other when using the mid-pulse. Using a simple phase-only model, the authors explored the possibility that light affects oscillator velocity strongly in the 1st hour and at reduced strength in later hours of the pulse due to photoreceptor adaptation. They fitted models based on the 1-h PRC to the data for all light pulses. The best overall correspondence between PRCs was obtained when the effect of light during all hours after the first was reduced by a factor of 0.22 relative to the 1st hour. For the predicted PRCs, the light action centered on average at 38% of the light pulse. This is close to the reference phase yielding best correspondence at 36% of the pulses. The result is thus compatible with an initial major contribution of the onset of the light pulse followed by a reduced effect of light responsible for the differences between PRCs for different duration pulses. The authors suggest that the mid-pulse is a better phase reference than lights-on to plot and compare PRCs of different light-pulse durations.

Contamination of Auditory Evoked Potential of Goldfish Carassius auratus with Microphonic Potential

In this experiment, we used sound stimuli of different durations to demonstrate that the auditory evoked potential (AEP) of fishes was contaminated with microphonic potential. We demonstrated that the AEP contained a component that had double the frequency as that of the sound stimulus. The durations of the AEPs were in proportion with the durations of the sound stimuli. “Auditory brainstem response” has been considered to be the dominant component of the AEP of fishes. However, this study suggests that some of the AEP components of fishes are derived from peripheral organs such as the saccule, lagena and utricle in the inner ear.

In vivo intracellular responses of the medial geniculate neurones to acoustic stimuli in anaesthetized guinea pigs.

In the present study, we investigated the auditory response features of the medial geniculate neurones, using in vivo intracellular recordings in anaesthetized guinea pigs. Of the 76 neurones examined, 9 showed 'off' or 'on-off' responses to an acoustic stimulus and thus were defined as 'off' or 'on-off' neurones. Among the remaining 67 neurones, 42 showed an excitatory postsynaptic potential (EPSP) to acoustic stimuli and 25 showed either a pure inhibitory postsynaptic potential (IPSP, 7 neurones), or an IPSP preceded by an EPSP (EPSP-IPSP type, 18 neurones). The EPSP responses exhibited a mean latency of 15.7 +/- 6.1 ms, which was significantly shorter than that of the IPSP responses (21.3 +/- 8.6 ms, P < 0.01). The IPSP responses also showed a significantly greater duration than the EPSP responses (208.5 +/- 128.2 ms versus 122.4 +/- 84.8 ms, P < 0.05), while there were no significant differences between the amplitudes of IPSP and EPSP (8.3 +/- 3.2 mV versus 8.7 +/- 5.3 mV). Of the 11 neurones that showed EPSP responses to acoustic stimuli and were histologically labelled, 7 were located in the lemniscal medial geniculate body (MGB) and 4 in the non-lemniscal MGB. Another 6 labelled neurones that showed IPSP responses to acoustic stimuli were located in the non-lemniscal MGB. With a membrane potential of above -72 mV, the neurones showed greater EPSP or IPSP to an acoustic stimulus when their membrane potential was depolarized. However, upon hyperpolarization to below -74 mV, the neurones shifted to low-threshold calcium spikes (LTS)/LTS bursts. In response to auditory stimuli of different durations, 'off' neurones that responded to the offset of the acoustic stimulus and were located in the non-lemniscal MGB showed different response latencies or deviations of latencies in addition to exhibiting different numbers of spikes, suggesting that the timing of the spikes could be another component utilized by thalamic neurones to encode information on the stimulus. Given that some non-lemniscal neurones are multisensory and project to the entire auditory cortex, the selective corticofugal inhibition in the non-lemniscal MGB would enable the ascending pathway to prepare the auditory cortex to receive subsequent auditory information, avoiding the interference of other sensory inputs.

Functional organization of sensory input to the olfactory bulb glomerulus analyzed by two-photon calcium imaging.

Glomeruli in the olfactory bulb are anatomically discrete modules receiving input from idiotypic olfactory sensory neurons. To examine the functional organization of sensory inputs to individual glomeruli, we loaded olfactory sensory neurons with a Ca(2+) indicator and measured odorant-evoked presynaptic Ca(2+) signals within single glomeruli by using two-photon microscopy in anaesthetized mice. Odorants evoked patterns of discrete Ca(2+) signals throughout the neuropil of a glomerulus. Across glomeruli, Ca(2+) signals occurred with equal probability in all glomerular regions. Within single glomeruli, the pattern of intraglomerular Ca(2+) signals was indistinguishable for stimuli of different duration, identity, and concentration. Moreover, the response time course of the signals was similar throughout the glomerulus. Hence, sensory inputs to individual glomeruli are spatially heterogeneous but seem to be functionally indiscriminate. These results support the view of olfactory glomeruli as functional units in representing sensory information.