Neurophysiological correlate of the auditory after-image ('Zwicker tone')

The 'Zwicker tone' (ZT) is an auditory after-image that can be evoked most effectively when a band-suppressed noise (relative width of gap 1/3 octave) presented for a certain period of time has been switched off. The sensation of this purely monaural phenomenon is that of a pure tone with a frequency corresponding to the center frequency of the gap and an equivalent level of 10-15 dB above auditory threshold. The sensation decays gradually; it may last as long as 10 s depending on how long the evoking noise was presented. The search for a physiological correlate has been futile so far, probably because the search was confined to more peripheral levels of the auditory system (inferior colliculus). A neuromagnetic study was performed in normal-hearing subjects in order to look for a neurophysiological correlate of the ZT in the auditory cortex. With a stimulation paradigm especially designed for this study, we have been able to isolate poststimulus activity which appears to be related to the ZT and which originates in the supratemporal auditory cortex. It is a sustained neuromagnetic activity that shows a clear-cut dipolar field distribution, and it appears that this activity has certain similarities with the tone-evoked auditory sustained response. The hypothesis is put forward that during the sensation of the ZT a process takes place in the auditory cortex which is similar to that underlying the sustained response, and which gives rise to the sensation of the ZT. In contrast to the sustained response, however, which is due to neural activity evoked by an external acoustic stimulus, the sustained activity associated with the ZT is due to a temporary absolute or relative reduction of neural activity originating from those regions in which the ZT exciting stimulus caused an adaptation. These differences in neural activity cannot be distinguished by the auditory system from a corresponding external acoustic signal. Preliminary studies in patients suffering from tonal tinnitus yielded results which exhibit a certain similarity with those obtained in the ZT experiment.     

Pseudospontaneous activity: stochastic independence of auditory nerve fibers with electrical stimulation

We describe a novel signal processing strategy for cochlear implants designed to emphasize stochastic independence across the excited neural population. The strategy is based on the observation that high rate pulse trains may produce random spike patterns in auditory nerve fibers that are statistically similar to those produced by spontaneous activity in the normal cochlea. We call this activity 'pseudospontaneous'. A supercomputer-based computational model of a population of auditory nerve fibers suggests that different average rates of pseudospontaneous activity can be created by varying the stimulus current of a fixed-amplitude, high-rate pulse train, e.g. 5000 pps. Electrically-evoked compound action potentials recorded in a human cochlear implant subject are consistent with the hypothesis that such a stimulus can desynchronize the fiber population. This desynchronization may enhance neural representation of temporal detail and dynamic range with a cochlear implant and eliminate a major difference between acoustic and electric hearing.     

Auditory afterimage: tonotopic representation in the auditory cortex

The auditory afterimage is a sensation which occurs for several seconds after the exciting acoustic signal has been switched off, and which roughly corresponds to the inverse of the spectrum of the exciting signal. In contrast to the well-known visual afterimage, the physiological mechanism generating the auditory afterimage has been questionable so far. Neuromagnetic source imaging revealed that the source of cortical neural activity which coincides with the sensation of the afterimage is located in the auditory cortex and exhibits a tonotopic organization similar to that of the sustained response which occurs during continuous presentation of an acoustic stimulus. It is concluded that the neural processes leading to the generation of the two phenomena -sustained response and auditory afterimage - are similar.     

Transfer of noncorresponding spatial associations to the auditory Simon task.

Four experiments examined transfer of noncorresponding spatial stimulus-response associations to an auditory Simon task for which stimulus location was irrelevant. Experiment 1 established that, for a horizontal auditory Simon task, transfer of spatial associations occurs after 300 trials of practice with an incompatible mapping of auditory stimuli to keypress responses. Experiments 2-4 examined transfer effects within the auditory modality when the stimuli and responses were varied along vertical and horizontal dimensions. Transfer occurred when the stimuli and responses were arrayed along the same dimension in practice and transfer but not when they were arrayed along orthogonal dimensions. These findings indicate that prior task-defined associations have less influence on the auditory Simon effect than on the visual Simon effect, possibly because of the stronger tendency for an auditory stimulus to activate its corresponding response. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Representation of spectral and temporal sound features in three cortical fields of the cat. Similarities outweigh differences

This study investigates the degree of similarity of three different auditory cortical areas with respect to the coding of periodic stimuli. Simultaneous single- and multiunit recordings in response to periodic stimuli were made from primary auditory cortex (AI), anterior auditory field (AAF), and secondary auditory cortex (AII) in the cat to addresses the following questions: is there, within each cortical area, a difference in the temporal coding of periodic click trains, amplitude-modulated (AM) noise bursts, and AM tone bursts? Is there a difference in this coding between the three cortical fields? Is the coding based on the temporal modulation transfer function (tMTF) and on the all-order interspike-interval (ISI) histogram the same? Is the perceptual distinction between rhythm and roughness for AM stimuli related to a temporal versus spatial representation of AM frequency in auditory cortex? Are interarea differences in temporal response properties related to differences in frequency tuning? The results showed that: 1) AM stimuli produce much higher best modulation frequencies (BMFs) and limiting rates than periodic click trains. 2) For periodic click trains and AM noise, the BMFs and limiting rates were not significantly different for the three areas. However, for AM tones the BMF and limiting rates were about a factor 2 lower in AAF compared with the other areas. 3) The representation of stimulus periodicity in ISIs resulted in significantly lower mean BMFs and limiting rates compared with those estimated from the tMTFs. The difference was relatively small for periodic click trains but quite large for both AM stimuli, especially in AI and AII. 4) Modulation frequencies <20 Hz were represented in the ISIs, suggesting that rhythm is coded in auditory cortex in temporal fashion. 5) In general only a modest interdependence of spectral- and temporal-response properties in AI and AII was found. The BMFs were correlated positively with characteristic frequency in AAF. The limiting rate was positively correlated with the frequency-tuning curve bandwidth in AI and AII but not in AAF. Only in AAF was a correlation between BMF and minimum latency was found. Thus whereas differences were found in the frequency-tuning curve bandwidth and minimum response latencies among the three areas, the coding of periodic stimuli in these areas was fairly similar with the exception of the very poor representation of AM tones in AII. This suggests a strong parallel processing organization in auditory cortex.     

A comparison of contextual and generalized auditory-cue fear conditioning: evidence for similar memory processes

A number of variables influence contextual, but not auditory-cue, fear conditioning. However, several of these variables (isolation, stimulus preexposure, retention interval, and age) affect generalized auditory-cue fear. More generalized fear was found when (a) rats were isolated in a novel environment than when returned to their home cages, (b) the retention interval was 3 hr rather than 24 hr, and (c) in 18-day-old compared with 25-day-old rats. Moreover, preexposure to the auditory cue eliminated the isolation effect. At a behavioral-psychological level, these variables may exert their effects by influencing the processes that construct a memory representation of the stimulus. At a neural systems level, they may influence processing carried out in the thalamo-corticoamygdaloid auditory pathway.     

Electrical stimulation of the auditory nerve. I. Correlation of physiological responses with cochlear status

The purpose of the present study was to evaluate evoked potential and single fibre responses to biphasic current pulses in animals with varying degrees of cochlear pathology, and to correlate any differences in the physiological response with status of the auditory nerve. Six cats, whose cochleae ranged from normal to a severe neural loss (< 5% spiral ganglion survival), were used. Morphology of the electrically evoked auditory brainstem response (EABR) was similar across all animals, although electrophonic responses were only observed from the normal animal. In animals with extensive neural pathology, EABR thresholds were elevated and response amplitudes throughout the dynamic range were moderately reduced. Analysis of single VIIIth nerve fibre responses were based on 207 neurons. Spontaneous discharge rates among fibres depended on hearing status, with the majority of fibres recorded from deafened animals exhibiting little or no spontaneous activity. Electrical stimulation produced a monotonic increase in discharge rate, and a systematic reduction in response latency and temporal jitter as a function of stimulus intensity for all fibres examined. Short-duration current pulses elicited a highly synchronous response (latency < 0.7 ms), with a less well synchronized response sometimes present (0.7-1.1 ms). There were, however, a number of significant differences between responses from normal and deafened cochleae. Electrophonic activity was only present in recordings from the normal animal, while mean threshold, dynamic range and latency of the direct electrical response varied with cochlear pathology. Differences in the ability of fibres to follow high stimulation rates were also observed; while neurons from the normal cochlea were capable of 100% entrainment at high rates (600-800 pulses per second (pps)), fibres recorded from deafened animals were often not capable of such entrainment at rates above 400 pps. Finally, a number of fibres in deafened animals showed evidence of 'bursting', in which responses rapidly alternated between high entrainment and periods of complete inactivity. This bursting pattern was presumably associated with degenerating auditory nerve fibres, since it was not recorded from the normal animal. The present study has shown that the pathological response of the cochlea following a sensorineural hearing loss can lead to a number of significant changes in the patterns of neural activity evoked via electrical stimulation. Knowledge of the extent of these changes have important implications for the clinical application of cochlear implants.     

NMR spectroscopy and brain diseases. Clinical applications

We studied the relationship between auditory activity in the midbrain and selective phonotaxis in females of the treefrog, Pseudacris crucifer. Gravid females were tested in two-stimulus playback tests using synthetic advertisement calls of different frequencies (2600 versus 2875 Hz; 2800 versus 3500 Hz; 2600 versus 3500 Hz). Tests were conducted with and without a background of synthesized noise, which was filtered to resemble the spectrum of a chorus of spring peepers. There were no significant preferences for calls of any frequency in the absence of background noise. With background noise, females preferred calls of 3500 Hz to those of 2600 Hz. Multi-unit recordings of neural responses to synthetic sounds were made from the torus semicircularis of the same females following the tests of phonotaxis. We measured auditory threshold at 25 frequencies (1800-4200 Hz) as well as the magnitude of the neural response when stimulus amplitude was held constant and frequency was varied. This procedure yielded isointensity response contours, which we obtained at six amplitudes in the absence of noise and at the stimulus amplitude used during the phonotaxis tests with background noise. Individual differences in audiograms and isointensity responses were poorly correlated with behavioural data except for the test of 2600 Hz versus 3500 Hz calls in noise. The shape of the neural response contours changed with stimulus amplitude and in the presence of the simulated frog chorus. At 85 dB sound pressure level (SPL), the level at which females were tested, the contours of females were quite flat. The contours were more peaked at lower SPLs as well as during the broadcast of chorus noise and white noise at an equivalent spectrum level (45-46 dB/Hz). Peaks in the isointensity response plots of most females occurred at stimulus frequencies ranging from 3200 to 3400 Hz, frequencies close to the median best excitatory frequency (BEF) of 3357 Hz but higher than the mean of the mid-frequency of the male advertisement call (3011 Hz). Addition of background noise may cause a shift in the neural response-intensity level functions. Our results highlight the well-known nonlinearity of the auditory system and the danger inherent in focusing solely on threshold measures of auditory sensitivity when studying the proximate basis of female choice. The results also show an unexpected effect of the natural and noisy acoustic environment on behaviour and responses of the auditory system. Copyright 1998 The Association for the Study of Animal Behaviour.     

Age-related alteration in processing of temporal sound features in the auditory midbrain of the CBA mouse

The perception of complex sounds, such as speech and animal vocalizations, requires the central auditory system to analyze rapid, ongoing fluctuations in sound frequency and intensity. A decline in temporal acuity has been identified as one component of age-related hearing loss. The detection of short, silent gaps is thought to reflect an important fundamental dimension of temporal resolution. In this study we compared the neural response elicited by silent gaps imbedded in noise of single neurons in the inferior colliculus (IC) of young and old CBA mice. IC neurons were classified by their temporal discharge patterns. Phasic units, which accounted for the majority of response types encountered, tended to have the shortest minimal gap thresholds (MGTs), regardless of age. We report three age-related changes in neural processing of silent gaps. First, although the shortest MGTs (1-2 msec) were observed in phasic units from both young and old animals, the number of neurons exhibiting the shortest MGTs was much lower in old mice, regardless of the presentation level. Second, in the majority of phasic units, recovery of response to the stimulus after the silent gap was of a lower magnitude and much slower in units from old mice. Finally, the neuronal map representing response latency versus best frequency was found to be altered in the old IC. These results demonstrate a central auditory system correlate for age-related decline in temporal processing at the level of the auditory midbrain.     

An anatomically constrained neural network model of fear conditioning.

Conditioning of fear reactions to an auditory conditioned stimulus (CS) paired with a footshock unconditioned stimulus/stimuli (UCS) involves CS transmission to the amygdala from the auditory thalamus, the auditory cortex, or both. This article presents a simple neural network model of this neural system. The model consists of modules of mutually inhibitory nonlinear units representing the different relevant anatomical structures of the thalamo-amygdala and thalamo-cortico-amygdala circuitry. Frequency-specific changes produced by fear conditioning were studied at the behavioral level (stimulus generalization) and the single-unit level (receptive fields). The findings mirror effects observed in conditioning studies of animals. This computational model provides an initial grounding for explorations of how emotional information and behavior are related to anatomical and physiological observations. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

High-density recording and topographic analysis of the auditory oddball event-related potential in patients with schizophrenia

BACKGROUND: Prior research has shown reductions of the N1, N2, and P300 auditory event-related potential (ERP) components in schizophrenic patients. Most studies have shown a greater P300 reduction in left versus right temporal leads in schizophrenic patients. These studies were done with sparse electrode arrays, covering restricted areas of the head, thus providing an incomplete representation of the topographic field distribution. METHODS: We used a 64-channel montage to acquire auditory oddball ERPs from 24 schizophrenic patients and 24 controls subjects. The N1, P2, N2, P300, and N2 difference (N2d) amplitudes and latencies were tested for group and laterality differences. Component topographies were mapped onto a three-dimensional head model to display the group differences. RESULTS: The schizophrenic group showed reduction of the N1 component, perhaps reflecting reduced arousal or vigilance, but no N1 topographic difference. An N2d was not apparent in the schizophrenic patients, perhaps reflecting severe disruption in neural systems of stimulus categorization. In the patients, the P300 was smaller over the left temporal lobe sites than the right. CONCLUSIONS: The increased ERP spatial sampling allowed a more complete representation of the dipolar nature of the P300, which showed field contours consistent with neural sources in the posterior superior temporal plane.     

Experience-dependent corticofugal adjustment of midbrain frequency map in bat auditory system

Recent studies of corticofugal modulation of auditory information processing indicate that cortical neurons mediate both a highly focused positive feedback to subcortical neurons "matched" in tuning to a particular acoustic parameter and a widespread lateral inhibition to "unmatched" subcortical neurons. This cortical function for the adjustment and improvement of subcortical information processing is called egocentric selection. Egocentric selection enhances the neural representation of frequently occurring signals in the central auditory system. For our present studies performed with the big brown bat (Eptesicus fuscus), we hypothesized that egocentric selection adjusts the frequency map of the inferior colliculus (IC) according to auditory experience based on associative learning. To test this hypothesis, we delivered acoustic stimuli paired with electric leg stimulation to the bat, because such paired stimuli allowed the animal to learn that the acoustic stimulus was behaviorally important and to make behavioral and neural adjustments based on the acquired importance of the acoustic stimulus. We found that acoustic stimulation alone evokes a change in the frequency map of the IC; that this change in the IC becomes greater when the acoustic stimulation is made behaviorally relevant by pairing it with electrical stimulation; that the collicular change is mediated by the corticofugal system; and that the IC itself can sustain the change evoked by the corticofugal system for some time. Our data support the hypothesis.     

Transfer of magnitude and spatial mappings to the SNARC effect for parity judgments.

Left–right keypresses to numerals are faster for pairings of small numbers to left response and large numbers to right response than for the opposite pairings. This spatial numerical association of response codes (SNARC) effect has been attributed to numbers being represented on a mental number line. We examined this issue in 3 experiments using a transfer paradigm. Participants practiced a number magnitude-judgment task or spatial stimulus–response compatibility task with parallel or orthogonal stimulus–response dimensions prior to performing a parity-judgment task. The SNARC effect was enhanced following a small–left/large–right magnitude mapping but reversed following a small–right/large–left mapping, indicating that associations between magnitude and response defined for the magnitude-judgment task were maintained for the parity-judgment task. The SNARC effect was unaffected by practice with compatible or incompatible spatial mapping for the parallel spatial task but was larger following up–right/down–left mapping than up–left/down–right mapping for the orthogonal spatial task. These results are inconsistent with the SNARC effect being due to a horizontal number line representation but consistent with a view that correspondence of stimulus and response code polarities contributes to the effect. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The time course of auditory perceptual learning: neurophysiological changes during speech-sound training

Here we report that training-associated changes in neural activity can precede behavioral learning. This finding suggests that speech-sound learning occurs at a pre-attentive level which can be measured neurophysiologically (in the absence of a behavioral response) to assess the efficacy of training. Children with biologically based perceptual learning deficits as well as people who wear cochlear implants or hearing aids undergo various forms of auditory training. The effectiveness of auditory training can be difficult to assess using behavioral methods because these populations are communicatively impaired and may have attention and/or cognitive deficits. Based on our findings, if neurophysiological changes are seen during auditory training, then the training method is effectively altering the neural representation of the speech/sounds and changes in behavior are likely to follow.     

Interaction between drug discriminative stimuli and exteroceptive, sensory signals.

Interactions between drug discriminative stimuli based on 5.6 and 10 mg/kg sodium pentobarbital (ip) and exteroceptive stimuli (visual and auditory) were studied in 27 male Sprague-Dawley rats in a T-maze. In 3 groups, visual stimuli (light vs dark) were differentially paired with drug stimuli; the 4th group discriminated combinations of tonal frequencies (1 kHz or 10 kHz) and the presence or absence of pentobarbital (10 mg/kg). In general, visual stimuli controlled choice behavior (left or right turn) to a greater extent than did the drug training stimuli, whereas the auditory stimuli exerted no apparent control over the pentobarbital stimulus in Group 4. Tests with higher doses (13.75 and 17.5 mg/kg) indicated augmented stimulus control by the drug dimension in 2 groups (Group 1, 10 mg/kg pentobarbital vs saline; Group 2, 5.6 mg/kg vs 10 mg/kg pentobarbital) but not in the 3rd group (5.6 mg/kg pentobarbital vs saline) in the "conflict" situation, in which the exteroceptive conditions signaled one response whereas the drug stimulus signaled the opposite response. Discrimination training with only one of the stimulus dimensions resulted in stimulus control in the following order: 10 mg/kg vs saline?>?5.6 mg/kg vs saline?>?1 kHz vs 10 kHz. This indicates that the auditory stimuli were of marginal significance. It is concluded drugs can compete with exteroceptive, visual stimuli for associative strength. (23 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Validity and boundary conditions of automatic response activation in the Simon task.

Three experiments were conducted to determine whether spatial stimulus-response compatibility effects are caused by automatic response activation by stimulus properties or by interference between codes during translation of stimulus into response coordinates. The main evidence against activation has been that in a Simon task with hands crossed, responses are faster at the response location ipsilateral to the stimulus though manipulated by the hand contralateral to the stimulus. The experiments were conducted with hands in standard and in crossed positions and electroencephalogram measures showed coactivation of the motor cortex induced by stimulus position primarily during standard hand positions with visual stimuli. Only in this condition did the Simon effect decay with longer response times. The visual Simon effect appeared to be due to specific mechanisms of visuomotor information transmission that are not responsible for the effects obtained with crossed hands or auditory stimuli. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Temporal-order discrimination for selected auditory and visual stimulus dimensions

Thresholds for the discrimination of temporal order were determined for selected auditory and visual stimulus dimensions in 10 normal-adult volunteers. Auditory stimuli consisted of binary pure tones varying in frequency or sound pressure level, and visual stimuli consisted of binary geometric forms varying in size, orientation, or color. We determined the effect of psychophysical method and the reliability of performance across stimulus dimensions. Using a single-track adaptive procedure, Experiment 1 showed that temporal-order thresholds (TOTs) varied with stimulus dimension, being lowest for auditory frequency, intermediate for size, orientation, and auditory level, and longest for color. Test performance improved over sessions and the profile of thresholds across stimulus dimensions had a modest reliability. Experiment 2 used a double-interleaved adaptive procedure and TOTs were similarly ordered as in Experiment 1. However, TOTs were significantly lower for initially ascending versus descending tracks. With this method, the reliability of the profile across stimulus dimensions and tracks was relatively low. In Experiment 3, psychometric functions were obtained for each of the stimulus dimensions and thresholds were defined as the interpolated 70.7% correct point. The relative ordering of TOTs was similar to those obtained in the first two experiments. Non-monotonicities were found in some of the psychometric functions, with the most prominent being for the color dimension. A cross-experiment comparison of results demonstrates that TOTs and their reliability are significantly influenced by the psychophysical method. Taken together, these results support the notion that the temporal resolution of ordered stimuli involves perceptual mechanisms specific to a given sensory modality or submodality.     

Neural encoding of single-formant stimuli in the ventral cochlear nucleus of the chinchilla

Responses of the principal unit types in the ventral cochlear nucleus of the chinchilla were studied with a single-formant stimulus set that covered fundamental frequency (f0) from 100 Hz to 200 Hz and formant center frequency (F1) from 256 to 782 Hz. Temporal coding for f0 and F1 was explored for 95 stimulus combinations of f0 (n = 5) and F1 (n = 19) in primarylike, onset and chopper unit categories. Several analyses that explored temporal coding were employed including: autocorrelation, interspike interval analysis, and synchronization to each harmonic of f0. In general, the representation of f0 is better in onset and chopper units than in primarylike units. Nearly all units in the cochlear nucleus showed a gain in phase locking to the envelope (f0) of the single-formant stimulus relative to the auditory nerve. The fundamental is represented directly in neural discharges of units in the cochlear nucleus with an interval code (also Cariani and Delgutte, 1996; Rhode, 1995). The formant is represented in the temporal domain in primarylike units, though some chopper and onset units also possess the ability to code F1 through discharge synchrony. Onset-I units, which are associated with the octopus cells, exhibited the strongest phase locking to f0 of any unit types studied. The representation of f0 and F1 in the temporal domain is weak or absent in some units. All-order-interspike interval distributions computed for populations of units show preservation of temporal coding for both f0 and F1. Results are in agreement with earlier amplitude modulation studies that showed nearly all cochlear nucleus unit types phase lock to the signal envelope better than auditory nerve fibers over a considerable range of signal amplitudes.     

Auditory hallucinations inhibit exogenous activation of auditory association cortex

Percepts unaccompanied by a veridical stimulus, such as hallucinations, provide an opportunity for mapping the neural correlates of conscious perception. Functional magnetic resonance imaging (fMRI) can reveal localized changes in blood oxygenation in response to actual as well as imagined sensory stimulation. The safe repeatability of fMRI enabled us to study a patient with schizophrenia while he was experiencing auditory hallucinations and when hallucination-free (with supporting data from a second case). Cortical activation was measured in response to periodic exogenous auditory and visual stimulations using time series regression analysis. Functional brain images were obtained in each hallucination condition both while the patient was on and off antipsychotic drugs. The response of the temporal cortex to exogenous auditory stimulation (speech) was markedly reduced when the patient was experiencing hallucinating voices addressing him, regardless of medication. Visual cortical activation (to flashing lights) remained normal over four scans. From the results of this study and previous work on visual hallucinations we conclude that hallucinations coincide with maximal activation of the sensory and association cortex, specific to the modality of the experience.