Divergent Responses in the Gap Prepulse Inhibition of the Acoustic Startle Reflex in Two Different Guinea Pig Colonies.

Animal models of tinnitus rely on interpretation of behavioural or reflexive tests to determine the presence of this phantom perception. A commonly used test is the gap prepulse inhibition of acoustic startle (GPIAS), which is often combined with prepulse inhibition (PPI) to ensure that reduced GPIAS suppression is not due to hearing loss caused by the acoustic trauma commonly used to trigger tinnitus development. In our laboratory GPIAS and PPI are routinely used on two colonies of outbred tri-colour guinea pigs. However, our results show that these colonies show divergent results even before any tinnitus-inducing treatment, which impacts their suitability in tinnitus models. Although colony 1 and 2 show similar results in PPI (~95% of animals showing significant suppression), only ~30% of colony 2 also shows significant suppression in GPIAS compared to ~75% of colony 1. Cochlear sensitivity measured using compound action potentials showed no significant differences between colonies. Therefore, peripheral threshold loss was excluded as a possible factor. Our results show that similar strains of laboratory animals can show highly divergent results and GPIAS testing for tinnitus will not work for every animal strain. In addition, our data support the notion that PPI and GPIAS responses may rely on different neural circuitry.

Using individual differences to test the role of temporal and place cues in coding frequency modulation.

The question of how frequency is coded in the peripheral auditory system remains unresolved. Previous research has suggested that slow rates of frequency modulation (FM) of a low carrier frequency may be coded via phase-locked temporal information in the auditory nerve, whereas FM at higher rates and/or high carrier frequencies may be coded via a rate-place (tonotopic) code. This hypothesis was tested in a cohort of 100 young normal-hearing listeners by comparing individual sensitivity to slow-rate (1-Hz) and fast-rate (20-Hz) FM at a carrier frequency of 500 Hz with independent measures of phase-locking (using dynamic interaural time difference, ITD, discrimination), level coding (using amplitude modulation, AM, detection), and frequency selectivity (using forward-masking patterns). All FM and AM thresholds were highly correlated with each other. However, no evidence was obtained for stronger correlations between measures thought to reflect phase-locking (e.g., slow-rate FM and ITD sensitivity), or between measures thought to reflect tonotopic coding (fast-rate FM and forward-masking patterns). The results suggest that either psychoacoustic performance in young normal-hearing listeners is not limited by peripheral coding, or that similar peripheral mechanisms limit both high- and low-rate FM coding.

The effects of ipsilateral, contralateral, and bilateral broadband noise on the mid-level hump in intensity discrimination.

Previous psychoacoustical and physiological studies indicate that the medial olivocochlear reflex (MOCR), a bilateral, sound-evoked reflex, may lead to improved sound intensity discrimination in background noise. The MOCR can decrease the range of basilar-membrane compression and can counteract effects of neural adaptation from background noise. However, the contribution of these processes to intensity discrimination is not well understood. This study examined the effect of ipsilateral, contralateral, and bilateral noise on the "mid-level hump." The mid-level hump refers to intensity discrimination Weber fractions (WFs) measured for short-duration, high-frequency tones which are poorer at mid levels than at lower or higher levels. The mid-level hump WFs may reflect a limitation due to basilar-membrane compression, and thus may be decreased by the MOCR. The noise was either short (50 ms) or long (150 ms), with the long noise intended to elicit the sluggish MOCR. For a tone in quiet, mid-level hump WFs improved with ipsilateral noise for most listeners, but not with contralateral noise. For a tone in ipsilateral noise, WFs improved with contralateral noise for most listeners, but only when both noises were long. These results are consistent with MOCR-induced WF improvements, possibly via decreases in effects of compression and neural adaptation.

Is it necessary to occlude the ear in bone-conduction testing at 4 kHz, in order to prevent air-borne radiation affecting the results?

OBJECTIVE: To re-evaluate the current BSA recommendation that the test ear should be occluded during the bone-conduction procedure at frequencies above 2 kHz to prevent audible air-borne radiation. DESIGN: Pure-tone audiometry was undertaken during routine hearing tests. The audiograms of fifty-two ears met the criteria for the study and were included. Bone conduction at 4 kHz was tested in three different conditions: test ear open/occluded by earplug and occluded by circumaural earphone. STUDY SAMPLE: Forty-four adults aged 41-77 years with average hearing levels from normal to severe loss. All complied fully with the test procedure. No audiogram had a significant conductive element. RESULTS: There was no significant difference in each of the three test situations. Only two audiograms showed any (5 dB) difference at 4 kHz when bone conduction was retested with the ear occluded. CONCLUSIONS: The errors that result in a false air-bone gap at 4 kHz would not appear to be due to air-borne radiation. Failure to occlude the ear canal at 4 kHz, where air-borne radiation is greatest, makes no significant difference to the audiometric results. It is therefore suggested that it is unnecessary to block the test ear during routine pure-tone bone-conduction testing to prevent audible air-borne radiation, and that this should no longer form part of normal clinical practice.

Computational modeling of Adelta-fiber-mediated nociceptive detection of electrocutaneous stimulation.

Sensitization is an example of malfunctioning of the nociceptive pathway in either the peripheral or central nervous system. Using quantitative sensory testing, one can only infer sensitization, but not determine the defective subsystem. The states of the subsystems may be characterized using computational modeling together with experimental data. Here, we develop a neurophysiologically plausible model replicating experimental observations from a psychophysical human subject study. We study the effects of single temporal stimulus parameters on detection thresholds corresponding to a 0.5 detection probability. To model peripheral activation and central processing, we adapt a stochastic drift-diffusion model and a probabilistic hazard model to our experimental setting without reaction times. We retain six lumped parameters in both models characterizing peripheral and central mechanisms. Both models have similar psychophysical functions, but the hazard model is computationally more efficient. The model-based effects of temporal stimulus parameters on detection thresholds are consistent with those from human subject data.

Age effects in discrimination of intervals within rhythmic tone sequences.

This study measured listener sensitivity to increments of a target inter-onset interval (IOI) embedded within tone sequences that featured different rhythmic patterns. The sequences consisted of six 50-ms 1000-Hz tone bursts separated by silent intervals that were adjusted to create different timing patterns. Control sequences were isochronous, with all tonal IOIs fixed at either 200 or 400 ms, while other patterns featured combinations of the two IOIs arranged to create different sequential tonal groupings. Duration difference limens in milliseconds for increments of a single sequence IOI were measured adaptively by adjusting the duration of an inter-tone silent interval. Specific target IOIs within sequences differed across discrimination conditions. Listeners included younger normal-hearing adults and groups of older adults with and without hearing loss. Discrimination performance measured for each of the older groups of listeners was observed to be equivalent, with each group exhibiting significantly poorer discrimination performance than the younger listeners in each sequence condition. Additionally, the specific influence of variable rhythmic grouping on temporal sensitivity was found to be greatest among older listeners.

Tracking of nociceptive thresholds using adaptive psychophysical methods.

Psychophysical thresholds reflect the state of the underlying nociceptive mechanisms. For example, noxious events can activate endogenous analgesic mechanisms that increase the nociceptive threshold. Therefore, tracking thresholds over time facilitates the investigation of the dynamics of these underlying mechanisms. Threshold tracking techniques should use efficient methods for stimulus selection and threshold estimation. This study compares, in simulation and in human psychophysical experiments, the performance of different combinations of adaptive stimulus selection procedures and threshold estimation methods. Monte Carlo simulations were first performed to compare the bias and precision of threshold estimates produced by three different stimulus selection procedures (simple staircase, random staircase, and minimum entropy procedure) and two estimation methods (logistic regression and Bayesian estimation). Logistic regression and Bayesian estimations resulted in similar precision only when the prior probability distributions (PDs) were chosen appropriately. The minimum entropy and simple staircase procedures achieved the highest precision, while the random staircase procedure was the least sensitive to different procedure-specific settings. Next, the simple staircase and random staircase procedures, in combination with logistic regression, were compared in a human subject study (n = 30). Electrocutaneous stimulation was used to track the nociceptive perception threshold before, during, and after a cold pressor task, which served as the conditioning stimulus. With both procedures, habituation was detected, as well as changes induced by the conditioning stimulus. However, the random staircase procedure achieved a higher precision. We recommend using the random staircase over the simple staircase procedure, in combination with logistic regression, for nonstationary threshold tracking experiments.

Bidirectional effects of aversive learning on perceptual acuity are mediated by the sensory cortex.

Although emotional learning affects sensory acuity, little is known about how these changes are facilitated in the brain. We found that auditory fear conditioning in mice elicited either an increase or a decrease in frequency discrimination acuity depending on how specific the learned response was to the conditioned tone. Using reversible pharmacological inactivation, we found that the auditory cortex mediated learning-evoked changes in acuity in both directions.

Discrimination of brief gaps marked by two stimuli: effects of sound length, repetition, and rhythmic grouping.

We examined the effects of sound marker length, marker repetition, and rhythmic grouping on auditory gap discrimination. The discrimination ofthe duration of a gap between two markers was impaired by lengthening these markers (from 150 to 262.5 ms). Discrimination was impaired by lengthening the preceding marker relative to lengthening the following marker, while the impairment was not increased when both markers were lengthened compared with when only the preceding marker was lengthened. This indicates that the level of discrimination is not decided by a simple summation of the effects of the preceding and of the following marker's length. Moreover, discrimination of a gap between a short (S) and a long (L) marker and of a gap between a long and a short marker was improved by repeating the presentation of these gaps (ie by repeating the markers alternately as SLSLSL...): both types of discrimination led to near identical performance. Finally, under the repetition condition each type of discrimination was not related to the tendency for each individual to perceive the stimulus sequences as segmented into rhythmic chunks of a short tone followed by a long tone (as [SL][SL][SL]...), or those of a long tone followed by a short tone (as S][LS][LS][L...).

Effects of sensorineural hearing loss on temporal coding of harmonic and inharmonic tone complexes in the auditory nerve.

Listeners with sensorineural hearing loss (SNHL) often show poorer thresholds for fundamental-frequency (F0) discrimination and poorer discrimination between harmonic and frequency-shifted (inharmonic) complex tones, than normal-hearing (NH) listeners-especially when these tones contain resolved or partially resolved components. It has been suggested that these perceptual deficits reflect reduced access to temporal-fine-structure (TFS) information and could be due to degraded phase locking in the auditory nerve (AN) with SNHL. In the present study, TFS and temporal-envelope (ENV) cues in single AN-fiber responses to band-pass-filtered harmonic and inharmonic complex tones were -measured in chinchillas with either normal-hearing or noise-induced SNHL. The stimuli were comparable to those used in recent psychophysical studies of F0 and harmonic/inharmonic discrimination. As in those studies, the rank of the center component was manipulated to produce -different resolvability conditions, different phase relationships (cosine and random phase) were tested, and background noise was present. Neural TFS and ENV cues were quantified using cross-correlation coefficients computed using shuffled cross correlograms between neural responses to REF (harmonic) and TEST (F0- or frequency-shifted) stimuli. In animals with SNHL, AN-fiber tuning curves showed elevated thresholds, broadened tuning, best-frequency shifts, and downward shifts in the dominant TFS response component; however, no significant degradation in the ability of AN fibers to encode TFS or ENV cues was found. Consistent with optimal-observer analyses, the results indicate that TFS and ENV cues depended only on the relevant frequency shift in Hz and thus were not degraded because phase locking remained intact. These results suggest that perceptual "TFS-processing" deficits do not simply reflect degraded phase locking at the level of the AN. To the extent that performance in F0- and harmonic/inharmonic discrimination tasks depend on TFS cues, it is likely through a more complicated (suboptimal) decoding mechanism, which may involve "spatiotemporal" (place-time) neural representations.

Across-site patterns of modulation detection: relation to speech recognition.

The aim of this study was to identify across-site patterns of modulation detection thresholds (MDTs) in subjects with cochlear implants and to determine if removal of sites with the poorest MDTs from speech processor programs would result in improved speech recognition. Five hundred millisecond trains of symmetric-biphasic pulses were modulated sinusoidally at 10 Hz and presented at a rate of 900 pps using monopolar stimulation. Subjects were asked to discriminate a modulated pulse train from an unmodulated pulse train for all electrodes in quiet and in the presence of an interleaved unmodulated masker presented on the adjacent site. Across-site patterns of masked MDTs were then used to construct two 10-channel MAPs such that one MAP consisted of sites with the best masked MDTs and the other MAP consisted of sites with the worst masked MDTs. Subjects' speech recognition skills were compared when they used these two different MAPs. Results showed that MDTs were variable across sites and were elevated in the presence of a masker by various amounts across sites. Better speech recognition was observed when the processor MAP consisted of sites with best masked MDTs, suggesting that temporal modulation sensitivity has important contributions to speech recognition with a cochlear implant.

Responses to deviants are modulated by subthreshold variability of the standard.

Auditory mechanisms automatically detect both basic features of sounds and the rules governing their presentation. In the oddball paradigm, the auditory system detects the sameness (or no-variability) rule when the same reference tone is consistently repeated. We used two oddball protocols, the classical one with a fixed reference and a modified one with a jittered reference, to determine whether the auditory system can detect subthreshold violations of sameness. We found that the response to the repeated standard was not modified by the small jitter. However, the response to the frequency oddball was smaller under the jittered protocol, indicating hypersensitivity to sameness. The sensitivity to jitter was largest when the oddball deviated by 8%, was smaller for 40%, and disappeared at 100% deviation, indicating that sensitivity to sameness is context dependent; namely, it is scaled with respect to the overall range of stimuli.

Mismatch negativity and P300 to behaviorally perceptible and imperceptible frequency and intensity contrasts.

Mismatch negativity and the P300 have been investigated as electrophysiological indices of behavioral auditory discrimination of duration. Using an oddball paradigm, responses were evoked to stimuli that had been behaviorally demonstrated to be either perceptible or imperceptible. The results indicated P300 events were present with the perceptible contrast but absent for the imperceptible contrast with all participants, while mismatch negativities were present in approximately 80 and 20% of participants to the perceptible and imperceptible contrasts, respectively. The present study extended the findings by applying the same paradigm to the discrimination of spectral and amplitude contrasts. 10 young adults with normal hearing were participants. Assuming that auditory assessment should involve representation and processing in all acoustic domains (i.e., temporal/duration, spectral/frequency, and amplitude/intensity), the effects seen with temporal contrasts were predicted to be similar for spectral and amplitude contrasts. The findings generally illustrated that the P300 was more accurate than the mismatch negativity in reflecting behavioral discrimination. Together, these studies challenge the use of mismatch negativity as an electrophysiological correlate for behavioral discrimination of auditory perceptible contrasts.

Vibrotactile stimulation of the upper leg: effects of location, stimulation method and habituation.

In this study vibrotactile stimulation of the upper leg and its usability for feedback was tested. Three experiments were performed on ten healthy subjects using pager motors. The first experiment was to test the perception of the vibration at different frequencies and at different locations of the upper leg. The second experiment tested the ability of subjects to estimate location and number of stimuli in an array. In addition it was evaluated whether simultaneous or sequential stimulation is better interpretable. Thirdly the habituation of the vibration was determined. The experiments showed that vibrotactile stimulation is well perceived and can be useful in providing feedback on the upper leg. Further experiments are needed to determine the effectiveness of vibrotactile stimulation for feedback in trans-femoral prostheses.

Non-linear laws of echoic memory and auditory change detection in humans.

BACKGROUND: The detection of any abrupt change in the environment is important to survival. Since memory of preceding sensory conditions is necessary for detecting changes, such a change-detection system relates closely to the memory system. Here we used an auditory change-related N1 subcomponent (change-N1) of event-related brain potentials to investigate cortical mechanisms underlying change detection and echoic memory. RESULTS: Change-N1 was elicited by a simple paradigm with two tones, a standard followed by a deviant, while subjects watched a silent movie. The amplitude of change-N1 elicited by a fixed sound pressure deviance (70 dB vs. 75 dB) was negatively correlated with the logarithm of the interval between the standard sound and deviant sound (1, 10, 100, or 1000 ms), while positively correlated with the logarithm of the duration of the standard sound (25, 100, 500, or 1000 ms). The amplitude of change-N1 elicited by a deviance in sound pressure, sound frequency, and sound location was correlated with the logarithm of the magnitude of physical differences between the standard and deviant sounds. CONCLUSIONS: The present findings suggest that temporal representation of echoic memory is non-linear and Weber-Fechner law holds for the automatic cortical response to sound changes within a suprathreshold range. Since the present results show that the behavior of echoic memory can be understood through change-N1, change-N1 would be a useful tool to investigate memory systems.

Modulation of auditory evoked responses to spectral and temporal changes by behavioral discrimination training.

BACKGROUND: Due to auditory experience, musicians have better auditory expertise than non-musicians. An increased neocortical activity during auditory oddball stimulation was observed in different studies for musicians and for non-musicians after discrimination training. This suggests a modification of synaptic strength among simultaneously active neurons due to the training. We used amplitude-modulated tones (AM) presented in an oddball sequence and manipulated their carrier or modulation frequencies. We investigated non-musicians in order to see if behavioral discrimination training could modify the neocortical activity generated by change detection of AM tone attributes (carrier or modulation frequency). Cortical evoked responses like N1 and mismatch negativity (MMN) triggered by sound changes were recorded by a whole head magnetoencephalographic system (MEG). We investigated (i) how the auditory cortex reacts to pitch difference (in carrier frequency) and changes in temporal features (modulation frequency) of AM tones and (ii) how discrimination training modulates the neuronal activity reflecting the transient auditory responses generated in the auditory cortex. RESULTS: The results showed that, additionally to an improvement of the behavioral discrimination performance, discrimination training of carrier frequency changes significantly modulates the MMN and N1 response amplitudes after the training. This process was accompanied by an attention switch to the deviant stimulus after the training procedure identified by the occurrence of a P3a component. In contrast, the training in discrimination of modulation frequency was not sufficient to improve the behavioral discrimination performance and to alternate the cortical response (MMN) to the modulation frequency change. The N1 amplitude, however, showed significant increase after and one week after the training. Similar to the training in carrier frequency discrimination, a long lasting involuntary attention to the deviant stimulus was observed. CONCLUSION: We found that discrimination training differentially modulates the cortical responses to pitch changes and to envelope fluctuation changes of AM tones. This suggests that discrimination between AM tones requires additional neuronal mechanisms compared to discrimination process between pure tones. After the training, the subjects demonstrated an involuntary attention switch to the deviant stimulus (represented by the P3a-component in the MEG) even though attention was not prerequisite.

Processing interaural cues in sound segregation by young and middle-aged brains.

BACKGROUND: When listening to one speaker while another conversation is occurring simultaneously, we separate the competing sounds by processing physical cues such as common onset time, intensity, frequency harmonicity, and spatial location of the sound sources. Spatial location is determined in large part by differences in arrival of a sound at one ear versus the other ear, otherwise known as interaural time difference (ITD) or interaural phase difference (IPD). There is ample anecdotal evidence that middle-aged adults experience greater difficulty listening to speech in noise, even when their audiological evaluation does not reveal abnormal results. Furthermore, it has been shown that the frequency range for IPD processing is reduced in middle-aged adults compared to young adults, even though morphological changes in the auditory evoked potential (AEP) response were only observed in older adults. PURPOSE: The purpose of the current study was to examine early aging effects (< 60 years) on IPD processing in concurrent sound segregation. RESEARCH DESIGN: We examined the change AEP evoked by detection of a mistuned and/or phase-shifted second harmonic during the last 1500 msec of a 3000 msec amplitude-modulated harmonic complex. A passive listening paradigm was used. STUDY SAMPLE: Ten young (21-35 years) and 11 middle-aged (48-57 years) adults with normal hearing were included in the study. DATA COLLECTION AND ANALYSIS: Scalp electroencephalographic activity was recorded from 63 electrodes. A temporospatial principal component analysis was conducted. Spatial factor scores of individual spatial factors were the dependent variable in separate mixed-design ANOVAs for each temporal factor of interest. Stimulus type was the within-subject independent variable, and age group was the between-subject independent variable. RESULTS: Results indicated a delay in the upward P2 slope and the P2 peak latency to a sudden phase shift in the second harmonic of a harmonic complex in middle-aged adults compared to young adults. This AEP difference increased as mistuning (as a second grouping cue) decreased and remained evident when the IPD was the only grouping cue. CONCLUSIONS: We conclude that our findings reflect neurophysiologic differences between young and middle-aged adults for IPD processing in concurrent sound segregation.

Stimulation threshold comparison of time-varying magnetic pulses with different waveforms.

PURPOSE: To clarify whether sinusoidal pulses possess lower thresholds than rectangular ones at perception threshold, a statement often made that contradicts the theory of stimulation. MATERIALS AND METHODS: The results of a nerve stimulation study with 65 volunteers and with trapezoidal and sinusoidal gradient pulses were used to apply the combination of the electric field, induced in the tissue of the human body, with the "Fundamental Law of Electrostimulation." This law claims that the waveshape of a pulse is not essential as long as the amplitude of the pulse does not decrease below rheobase (rheobase condition). RESULTS: If the rheobase condition is applied to sinusoidal waveforms and the pulse duration and amplitude is corrected accordingly, both trapezoidal and sinusoidal gradient pulses have identical threshold amplitudes as a function of pulse duration. CONCLUSION: The "Fundamental Law of Electrostimulation," including the "rheobase condition," proved to be a good basis for describing magnetic field stimulation (magnetostimulation) and that application of it to magnetostimulation is suitable as the basis for describing magnetic field stimulation with various waveforms. For nonrectangular pulses, pulse durations and pulse amplitudes must be corrected according to the "rheobase condition." The exponential Blair Equation is less suited to be applied in magnetostimulation.

Perception of stationary and moving sound following unilateral cortectomy.

The perception of motion is an essential prerequisite to responding adequately to the dynamic aspects of sensory information in the environment. The neural substrates of auditory motion processing are, at present, still a matter of debate. It has been hypothesized that motion information is, as in the visual system, processed separately from other aspects of auditory information, such as stationary location. Here we report data on auditory perception of stationary and motion stimuli from a subject with right-sided resection of the anterior temporal-lobe region including medial aspects of Heschl's gyrus, and from three subjects with unilateral (right-sided or left-sided) hemispherectomy. All these subjects had undergone cortectomy decades earlier. The subjects with hemispherectomy were completely unable to perceive auditory motion, but showed slight to moderate deficits in judging stationary location. The subject with temporal lobectomy exhibited quite similar stationary auditory deficits as found in the subjects with hemispherectomy, but was completely normal in judging auditory motion. Thus, there was a clear dissociation of the effects of unilateral temporal lobectomy and hemispherectomy on auditory motion perception. Collectively, these findings suggest that the unilateral anterior temporal-lobe region plays a significant role in the analysis of stationary, but not moving, sound. One may assume that the cortical "motion network" is distinct from the "stationary network", and is located either in the most posterior aspects of temporal lobe, or in non-temporal, most likely parietal, areas.

A method to estimate the spatial extent of activation in thalamic deep brain stimulation.

OBJECTIVE: The goal of this study was to develop, evaluate, and apply a method to quantify the unknown spatial extent of activation in deep brain stimulation (DBS) of the ventral intermedius nucleus (Vim) of the thalamus. METHODS: The amplitude-distance relationship and the threshold amplitudes to elicit clinical responses were combined to estimate the unknown amplitude-distance constant and the distance between the electrode and the border between the Vim and the ventrocaudal nucleus (Vc) of the thalamus. We tested the sensitivity of the method to errors in the input parameters, and subsequently applied the method to estimate the amplitude-distance constant from clinically-measured threshold amplitudes. RESULTS: The method enabled estimation of the amplitude-distance constant with a median squared error of 0.07-0.23V/mm2 and provided an estimate of the distance between the electrode and the Vc/Vim border with a median squared error of 0.01-0.04mm. Application of the method to clinically-measured threshold amplitudes to elicit paresthesias estimated the amplitude-distance constant to be 0.22V/mm2. CONCLUSIONS: The method enabled robust quantification of the spatial extent of activation in thalamic DBS and predicted that stimulation amplitudes of 1-3.5V would produce a mean effective radius of activation of 2.0-3.9mm. SIGNIFICANCE: Knowing the spatial extent of activation may improve methods of electrode placement and stimulation parameter selection in DBS.