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Georg M Klump

Publications and source records attributed to Georg M Klump.

16 recordsLinked to original sources

Temporal integration in the gerbil: the effects of age, hearing loss and temporally unmodulated and modulated speech-like masker noises.

We characterized temporal integration for 2k Hz pure tones with durations between 10 and 1000 ms in young, normal hearing old and old gerbils with a small hearing loss. Thresholds determined in silence increased for durations below 300 ms and were on average more than 10 dB higher for the 10 ms signal than asymptotic thresholds for the long signals. The amount of temporal integration tended to be less in gerbils with hearing loss. Threshold determination was repeated in the same individuals in the presence of speech-like unmodulated and modulated masking noises. Threshold shift due to the maskers was inversely related to the threshold in silence resulting in a reduced inter-individual variability of thresholds in both masking conditions. Thresholds differed systematically between both masker types in a duration dependent fashion. For long signal durations (300 and 1000 ms) thresholds were on average 2dB lower and for the 10 ms signal 1.9 dB higher in the presence of the modulated masker. These differences in threshold obtained with the two maskers were significant. One hypothesis is that long signals can be detected in the troughs of the modulated masker, while peaks interfere with the detection of short signals.

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Forward masking in gerbils: the effect of age.

We investigated forward masking in 21 gerbils as a function of age (5-47 months) using 400ms maskers at 40dB SPL and a 20ms, 2.85kHz probe presented 2.5ms after the masker. Elevated thresholds for the unmasked probe were only observed in animals older than 3 years. Unmasked thresholds showed no significant age-dependent hearing loss in animals below 3 years of age. In these animals without peripheral hearing loss, we found a significant age-dependent increase of masker-induced threshold shift. A regression analysis revealed that threshold shift increased from 23dB in 1 year old gerbils to 37dB in 3 year old animals. Increased forward masking in these animals with no sign of peripheral hearing loss points to a central processing deficit.

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Asymmetry of masking in the European starling: behavioural auditory thresholds.

Psychophysical studies of simultaneous masking with human observers exhibit an asymmetry in the amount of masking that depends on the relative bandwidths of signals and maskers. For noise bands up to the bandwidth of one auditory filter, masked auditory thresholds are considerably lower when the bandwidth of the signal exceeds that of the masker compared to the reversed condition. We investigate asymmetry of masking in an animal model, that will allow to study the mechanisms associated with the asymmetry of masking effect. European starlings (Sturnus vulgaris) were trained in a Go/NoGo paradigm to report the detection of a 500 ms noise signal centred in a 700 ms noise masker. Signals and maskers with centre frequencies of 2 kHz had bandwidths of 4 Hz or 256 Hz. Thresholds for detecting the 256 Hz wide-band signal in a 4 Hz narrow-band masker were considerably lower compared to detecting the 4 Hz narrow-band signal in a 256 Hz wide masker and compared to all other conditions. The asymmetry of masking in starlings was on average 15 and 17 dB for 40 and 70 dB SPL overall masker level, respectively. Our animal model thus proved perceptual abilities similar to human subjects. The results are discussed with respect to the importance of both intensity and temporal cues for signal detection.

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A quantitative analysis of psychometric functions for different auditory tasks in gerbils.

The psychometric function relates the probability of a correct response to the variation of a physical stimulus parameter. In many perceptual tasks one point on this function is defined by a more or less arbitrary threshold criterion and threshold is used to study the effects of various treatments or age. Besides threshold, the shape of the psychometric function provides additional information. The variability of internal (neural) noise and the sensorineural transduction function will affect the shape of the psychometric function and may, therefore, reveal important features in the processing of stimulus characteristics. Here we analyze the effect of age on psychometric functions from gerbils: (A) for the detection of a tone or noise pulse in silence which is generally regarded as a measure of cochlear function and (B) for a gap detection task, investigating aspects of temporal processing that involve the ascending auditory pathway. Our data show that the slope of the psychometric function for the detection of tone and noise pulses in silence is independent of age and threshold. In contrast, the steepness of the psychometric function is decreased in gerbils with impaired temporal resolution. We discuss these observations in the context of physiological data from young and old animals.

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Resolution in azimuth sound localization in the Mongolian gerbil (Meriones unguiculatus).

Minimum resolvable angles (MRAs) for sound localization in azimuth in the gerbil were determined in a behavioral study using tones, 300-Hz bands of noise centered at frequencies between 500 Hz and 8 kHz and broad-band noise of on average 60 dB SPL overall level. Using the method of constant stimuli, seven gerbils were trained in a two-alternative-forced-choice procedure to indicate if sounds were presented to them from the left or from the right by choosing the left or right arm of a Y-shaped cage. The MRA is the minimum angle between two loudspeaker locations that the gerbils discriminated. Animals were either stimulated from the front (N=4) or from the back (N=3). The MRA for broad-band noise randomly varying in level by +/- 6 dB was 23 degrees and 45 degrees for gerbils stimulated from the front or back, respectively. Generally a gerbil's MRA for tones declined up to 2 kHz reaching 20 degrees and 31 degrees for gerbils stimulated from the front or back, respectively, and the MRA was generally increased above this frequency. Results for narrow-band noise stimuli were similar. Results are discussed with respect to the available interaural cues and physiological mechanisms of sound localization in the gerbil.

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Operant methods for mouse psychoacoustics.

Tone detection thresholds for a 10-kHz tone in NMRI mice were determined in psychoacoustic experiments using both a constant-stimuli procedure and a two-down/one-up adaptive-tracking procedure in the same subjects and applying identical threshold criteria (70.7% response probability). Constant-stimuli thresholds were on average 24 dB lower than adaptive-tracking thresholds, and there was a trend indicating that constant-stimuli thresholds were less variable than adaptive-tracking thresholds. Furthermore, in the constant-stimuli procedure the number of trials constituting the psychometric function could be reduced from 100 to 50 trials without a large loss of accuracy of threshold determination. In the constant-stimuli procedure, the threshold value was affected by the threshold criteria. The lowest and least variable constant-stimuli thresholds were obtained by applying signal detection theory and a criterion of d' = 1. Thus, the constant-stimuli procedure in combination with signal detection theory appears to be better suited than the adaptive-tracking procedure to determine auditory sensory thresholds.

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Auditory stream segregation in the songbird forebrain: effects of time intervals on responses to interleaved tone sequences.

For both humans and other animals, the abilities to integrate separate sound elements over time into coherent perceptual representations, or 'auditory streams', and to segregate these auditory streams from other interleaved sounds are critical for hearing and vocal communication. In humans and European starlings (Sturnus vulgaris) the ability to perceptually segregate a simple interleaved tone sequence comprised of two alternating tones differing in frequency (ABA-ABA-ABA-...) into separate auditory streams of A and B tones is promoted at larger frequency separations (DeltaF) between the A and B tones. In humans, segregating A and B tones into different streams also appears to be promoted at shorter interstimulus intervals (ISI) between tones within a stream (e.g., between consecutive A tones). Here, we used the ABA experimental paradigm to investigate the influence of different time intervals between A and B tones in repeated ABA triplets on neural responses in the starling forebrain. The main finding from the study is that a DeltaF-dependent effect of ISI had a large influence on the relative responses to A and B tones. Responses to B tones were suppressed, relative to A-tone responses, when the A and B tones were more similar in frequency (smaller DeltaFs) and occurred at shorter ISIs. We attribute these suppressive effects to physiological forward masking and suggest that forward masking functions as a mechanism for segregating neural responses to interleaved tones in tonotopic space. We discuss the relevance of our physiological data with respect to previous electrophysiological studies of auditory stream segregation in mammals and previous perceptual studies in humans.

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Duration discrimination in the mouse (Mus musculus).

Detection thresholds for an increment in duration of a 10-kHz pure tone were determined in the NMRI mouse using a Go/NoGo-procedure and the method of constant stimuli. Thresholds for reference durations of 50, 100 and 200 ms were obtained presenting the signals at a fixed level or at a level varying by +/-3 dB. Thresholds were determined using signal-detection theory ( d'=1.0 or d'=1.8) and the criterion of 50% correct responses. For a fixed level, the average Weber fraction Delta T/ T (criterion of d'=1.8) significantly decreased from 1.18 or 1.23 at reference durations of 50 or 100 ms, respectively, to 0.97 at a reference duration of 200 ms. Thresholds were on average reduced by 46.8 or 55.4% for the threshold criteria d'=1 or 50% correct responses, respectively. There was no effect of randomizing the level on the discrimination threshold. Duration discrimination in the NMRI mouse does not follow Weber's law. The results are consistent with a mechanism summing up neural impulses over the duration of the stimulus. The psychoacoustic data are compared with results obtained by Brand et al. (J Acoust Soc Am 51:1291-1223, 2000) on the representation of acoustic signal duration in the mouse inferior colliculus.

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Primitive auditory stream segregation: a neurophysiological study in the songbird forebrain.

Auditory stream segregation refers to the perceptual grouping of sounds, to form coherent representations of objects in the acoustic scene, and is a fundamental aspect of hearing and speech perception. The perceptual segregation of simple interleaved tone sequences has been studied in humans and European starlings (Sturnus vulgaris) using sequences of 2 alternating tones differing in frequency (ABA-ABA-ABA-...). The segregation of A and B tones into separate auditory streams is believed to be promoted by preattentive auditory processes that increase the separation of excitation patterns along a tonotopic gradient. We tested the hypothesis that frequency selectivity and forward masking operate as 2 preattentive processes in sequential stream segregation by recording neural responses in the auditory forebrain of awake starlings to repeated ABA- sequences in which we varied the frequency separation (DeltaF) between the A and B tones and the tone repetition time (TRT). The A tones were presented at the neurons' characteristic frequency (CF), and B tones differed from the CF over a one-octave range. Larger DeltaF values and shorter TRTs promote the perceptual segregation of alternating tone sequences in humans and also resulted in larger differences in neural responses to alternating CF (A) and non-CF (B) tones. Our results are consistent with the hypothesis that preattentive auditory processes, such as frequency selectivity and forward masking, contribute to the perceptual segregation of sequential acoustic events having different frequencies into separate auditory streams, but also suggest that additional processes may be required to account for all known perceptual effects related to sequential auditory stream segregation.

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Age-dependent changes of gap detection in the Mongolian gerbil (Meriones unguiculatus).

Gap detection using broadband noise was characterized in a group of young gerbils from the breeding colony of the University of Regensburg (RB gerbils), old RB gerbils, and old gerbils from the breeding colony of the University of South Carolina (SC gerbils). Data from old RB and old SC gerbils were not significantly different and were subsequently combined for a comparison with data from the group of young RB gerbils. Level dependence of gap-detection thresholds in young and old domesticated gerbils resembled the typical mammalian pattern of level dependence. Gap-detection thresholds of old gerbils were significantly elevated at 30 dB SL and 50 dB SPL as compared with young gerbils. Compared with young gerbils tested at 30 dB SL and 50 dB SPL, the distribution of gap-detection thresholds in old gerbils was broader with a spread to higher gap-detection thresholds. Some old animals retained excellent temporal resolution, while some showed impaired gap detection. The gap-detection data collected in young and old gerbils resemble previously published data from humans of different age and confirm that gerbils are a useful model to study age-dependent changes in temporal processing.

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Boosting GABA improves impaired auditory temporal resolution in the gerbil.

Elderly humans often not only experience peripheral hearing loss but also suffer from more central deficits in temporal auditory processing affecting speech perception. Impaired auditory temporal resolution has also been observed in old rodents. Other studies have demonstrated a reduction of GABAergic function in the auditory pathway of old animals. Here we test the hypothesis that deficits in the GABAergic system affect central auditory processing. Our data suggests that pharmacological augmentation of the GABAergic system ameliorates impaired temporal auditory processing in the gerbil and might be a strategy for the treatment of at least some forms of central hearing loss in humans.

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Within- and across-channel processing in auditory masking: a physiological study in the songbird forebrain.

Synchronous envelope fluctuations in different frequency ranges of an acoustic background enhance the detection of signals in background noise. This effect, termed comodulation masking release (CMR), is attributed to both processing within one frequency channel of the auditory system and comparisons across separate frequency channels. Here we present data on CMR from a study in field L2 of the auditory forebrain of the European starling (Sturnus vulgaris) using two 25-Hz-wide bands of masking noise that provide the opportunity to distinguish between within-channel and across-channel effects. Acoustically evoked responses were recorded from unrestrained birds via radio telemetry. The signal was a 800 msec pure tone presented at the most sensitive frequency of the units in a previously determined frequency-tuning curve (FTC). One band of masking noise was centered on the signal frequency while the flanking band of noise was presented either within the limits of the excitatory FTC (i.e., within the same frequency channel as the on-frequency masker) or in the suppression area of the FTC (i.e., in a separate channel). For flanking bands inside the excitatory FTC, signal detection thresholds based on the rate code were lower in noise maskers with identical envelope fluctuations (comodulated) than in maskers with uncorrelated envelopes resulting in a neural CMR of approximately 4-7 dB. For flanking bands inside the suppression areas, the neural CMR was reduced. Although the average neural CMR was below the behaviorally determined CMR, a subsample of between 11 and 26% of the recording sites resembled the behavioral performance.

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Gap detection in Mongolian gerbils (Meriones unguiculatus).

Hearing thresholds for white-noise stimuli and temporal gap-detection thresholds in six Mongolian gerbils (Meriones unguiculatus) were determined in a GO/NOGO procedure using the method of constant stimuli. Gerbils were offspring of animals collected in the field and only bred in captivity for four generations or less. Hearing thresholds for 800 ms bursts of white noise ranged between -0.8 and 6.3 dB SPL. The median minimum-detectable gap centered in an 800 ms burst of white noise presented at 50 dB SPL was 2.1 ms. At levels of 40 dB SPL and above, gap-detection thresholds were independent of the sound-pressure level of the stimulus. At 30 dB SPL and below, the minimum-detectable gap increased with decreasing sound-pressure level. Near 5 dB sensation level, gap-detection thresholds ranged from 13.6 to 29.4 ms. The median threshold for the detection of gaps inserted 5 ms after the onset of an 800 ms burst of white noise of 50 dB SPL was 2.3 ms. The individuals' detection thresholds varied between 2.3 and 3.6 ms for stimuli in which the relative sound-pressure levels of the noise before and after the gap differed by up to 20 dB. The data found in the Mongolian gerbil match results from other mammal and bird species.

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Temporal modulation transfer functions in the barn owl (Tyto alba).

Barn owls (Tyto alba) have evolved several specializations in their auditory system to achieve the high sensory acuity required for prey capture, including superior processing of interaural time differences and phase coding in the auditory periphery. Here, we tested whether barn owls are capable of high temporal resolution that may be a prerequisite for the accuracy in binaural processing. Temporal resolution was measured psychoacoustically and demonstrated in temporal modulation transfer functions. Four barn owls were trained in an operant task with food reward to detect sinusoidal amplitude modulations within an 800-ms gated white-noise burst or 800-ms periods of modulation in continuous white noise (spectrum levels of -5 dB and 15 dB SPL). Within the range of tested amplitude modulation frequencies from 5 Hz to 1280 Hz, barn owls' detection thresholds were lowest at 10-20 Hz. This sensitivity corresponds to an intensity-difference limen of between 0.9 dB and 1.4 dB. For all conditions, temporal modulation transfer functions showed band-pass characteristics with a high-frequency cutoff in the range of 37 Hz to 92 Hz, corresponding to minimum integration times of 4.3 ms and 1.7 ms, respectively. In summary, these data indicate a temporal resolution in the owl's auditory system that is good, but not unusual, compared to other vertebrates.

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An estimate of the auditory-filter bandwidth in the Mongolian gerbil.

Critical ratios (CRs) and the critical band (CB) were determined in six Mongolian gerbils (Meriones unguiculatus) using a GO/NOGO procedure and the method of constant stimuli. The test-tone frequencies were 0.5, 1, 2, 4 and 8 kHz in the CR measurements and 2 kHz in the CB measurement. Critical ratios were independent of the level of the white-noise masker. The lowest CR with a mean of 24.8 dB was found at 1 kHz for maskers with spectrum levels of 20 and 40 dB SPL. The CR increased on average by 2.1 dB per octave between the frequencies 1 and 4 kHz. The CR bandwidth at 2 kHz and at a masker spectrum level of 40 dB SPL was 417 Hz. Using the band-narrowing procedure, we determined a mean CB of 216 Hz at a test-tone frequency of 2 kHz. The results are discussed in comparison to psychophysical data from other rodent species and humans. Finally, the relation of CR and CB bandwidths to the bandwidth of tuning curves of gerbil auditory-nerve fibers and the gerbil's cochlear frequency map are explored.

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Behavioral and evoked-potential thresholds in young and old Mongolian gerbils (Meriones unguiculatus).

Age-dependent hearing loss has been well documented in gerbils exceeding 2 years of age using physiological methods (e.g. [Mills et al. (1990) Hear. Res. 46, 201-210]). We determined behavioral thresholds for broad-band noise and pure-tone pulses in gerbils as a function of age. Contrary to expectations based on previously published physiological data, we found no significant (broad-band noise and 10 kHz) or only a very small hearing loss (7 dB at 2 kHz) in 30-36-month-old animals. In animals over 3 years of age we observed an increased spread of thresholds and threshold shifts exceeding 20 dB in some individuals. Behavioral thresholds of old gerbils from two breeding colonies (University of Regensburg and Medical University of South Carolina) were similar. Data from individual animals where thresholds were determined physiologically and behaviorally indicate that results from auditory brainstem response measurements show no shift at 18 months while subsequent measurements at 28-29 months revealed age-dependent threshold shifts of 10-15 dB. In contrast, thresholds determined by behavioral methods in these same individuals at 31-33 months of age remained stable.

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