Biomedical subjects

Laurel H Carney

Publications and source records attributed to Laurel H Carney.

At least 19 recordsLinked to original sources

PubMed2006

Neural rate and timing cues for detection and discrimination of amplitude-modulated tones in the awake rabbit inferior colliculus.

Neural responses to amplitude-modulated (AM) tones in the unanesthetized rabbit inferior colliculus (IC) were studied in an effort to establish explicit relationships between physiological and psychophysical measures of temporal envelope processing. Specifically, responses to variations in modulation depth (m) at the cell's best modulation frequency, with and without modulation maskers, were quantified in terms of average rate and synchronization to the envelope over the entire perceptual dynamic range of depths. Statistically significant variations in the metrics were used to define neural AM detection and discrimination thresholds. Synchrony emerged at modulation depths comparable with psychophysical AM detection sensitivities in some neurons, whereas the lowest rate-based neural thresholds could not account for psychoacoustical thresholds. The majority of rate thresholds (85%) were -10 dB or higher (in 20 log m), and 16% of the population exhibited no systematic dependence of average rate on m. Neural thresholds for AM detection did not decrease systematically at higher SPLs (as observed psychophysically): thresholds remained constant or increased with level for most cells tested at multiple sound-pressure levels (SPLs). At depths higher than the rate-based detection threshold, some rate modulation-depth functions were sufficiently steep with respect to the across-trial variability of the rate to predict depth discrimination thresholds as low as 1 dB (comparable with the psychophysics). Synchrony, on the other hand, did not vary systematically with m in many cells at high modulation depths. A simple computational model was extended to reproduce several features of the modulation frequency and depth dependence of both transient and sustained pure-tone responders.

Acoustic Stimulation↗

PubMed2006

Temporal measures and neural strategies for detection of tones in noise based on responses in anteroventral cochlear nucleus.

To examine possible neural strategies for the detection of tones in broadband noise, single-neuron extracellular recordings were obtained from the anteroventral cochlear nucleus (AVCN) in anesthetized gerbils. Detection thresholds determined by average discharge rate and several temporal metrics were compared with previously reported psychophysical detection thresholds in cats (Costalupes 1985). Because of their limited dynamic range, the average discharge rates of single neurons failed to predict psychophysical detection thresholds for relatively high-level noise at all measured characteristic frequencies (CFs). However, temporal responses changed significantly when a tone was added to a noise, even for neurons with flat masked rate-level functions. Three specific temporal analyses were applied to neural responses to tones in noise. First, temporal reliability, a measure of discharge time consistency across stimulus repetitions, decreased with increasing tone level for most AVCN neurons at all measured CFs. Second, synchronization to the tone frequency, a measure of phase-locking to the tone, increased with tone level for low-CF neurons. Third, rapid fluctuations in the poststimulus time histograms (PSTHs) decreased with tone level for a number of neurons at all CFs. For each of the three temporal measures, some neurons had detection thresholds at or below psychophysical thresholds. A physiological model of a higher-stage auditory neuron that received simple excitatory and inhibitory inputs from AVCN neurons was able to extract the PSTH fluctuation information in a form of decreased rate with tone level.

Acoustic Stimulation↗

PubMed2006

Correction of the peripheral spatiotemporal response pattern: a potential new signal-processing strategy.

The purpose of this paper is to introduce a new signal-processing strategy, spatiotemporal pattern correction (SPC), that is based on our knowledge of the level-dependent temporal response properties of auditory nerve fibers in normal and impaired ears. In normal-hearing listeners, tuning is sharp for low-level inputs and broadens as input level increases. When peripheral filters change their shape with input level, the phase properties of the filters and the latency of the filter's response also change. However, in listeners with hearing loss, tuning is broad for both low- and high-level inputs. Thus, there is little change in the phase properties of the filters for different input levels. The SPC strategy manipulates the temporal aspects of different frequency channels of sounds in an attempt to "correct" for this abnormal spatiotemporal response pattern of the impaired ear. Quality judgments and intelligibility measures of speech processed at various SPC strengths were obtained from a small group of normal-hearing listeners and listeners with hearing loss. In general, listeners with hearing loss preferred sentences with some degree of SPC processing, whereas normal-hearing listeners preferred unprocessed sentences. Benefit from SPC on the nonsense syllable test varied greatly across phonemes and listeners. A more comprehensive study of listeners with different degrees and configurations of hearing loss is needed to determine the amount of SPC benefit.

Adult↗

PubMed2006

Determination of the potential benefit of time-frequency gain manipulation.

OBJECTIVE: The purpose of this study was to determine the maximum benefit provided by a time-frequency gain-manipulation algorithm for noise-reduction (NR) based on an ideal detector of speech energy. The amount of detected energy necessary to show benefit using this type of NR algorithm was examined, as well as the necessary speed and frequency resolution of the gain manipulation. DESIGN: NR was performed using time-frequency gain manipulation, wherein the gains of individual frequency bands depended on the absence or presence of speech energy within each band. Three different experiments were performed: (1) NR using ideal detectors, (2) NR with nonideal detectors, and (3) NR with ideal detectors and different processing speeds and frequency resolutions. All experiments were performed using the Hearing-in-Noise test (HINT). A total of 6 listeners with normal hearing and 14 listeners with hearing loss were tested. RESULTS: HINT thresholds improved for all listeners with NR based on the ideal detectors used in Experiment I. The nonideal detectors of Experiment II required detection of at least 90% of the speech energy before an improvement was seen in HINT thresholds. The results of Experiment III demonstrated that relatively high temporal resolution (<100 msec) was required by the NR algorithm to improve HINT thresholds. CONCLUSIONS: The results indicated that a single-microphone NR system based on time-frequency gain manipulation improved the HINT thresholds of listeners. However, to obtain benefit in speech intelligibility, the detectors used in such a strategy were required to detect an unrealistically high percentage of the speech energy and to perform the gain manipulations on a fast temporal basis.

Acoustic Stimulation↗

PubMed2006

Binaural detection with narrowband and wideband reproducible noise maskers. III. Monaural and diotic detection and model results.

A single-interval, yes-no, tone-in-noise detection experiment was conducted to measure the proportion of "tone present" responses to each of 25 reproducible noise-alone and tone-plus-noise waveforms under narrowband (100 Hz), wideband (2900 Hz), monotic, and diotic stimulus conditions. Proportions of "tone present" responses (estimates of the probabilities of hits and false alarms) were correlated across masker bandwidths and across monotic and diotic conditions. Two categories of models were considered; one based on stimulus energy or neural counts, and another based on temporal structure of the stimulus envelope or neural patterns. Both categories gave significant correlation between decision variables and data. A model based on a weighted combination of energy in multiple critical bands performed best, predicting up to 90% of the variance in the reproducible-noise data. However, since energy-based models are unable to successfully explain detection under a roving-level paradigm without substantial modification, it is argued that other variations of detection models must be considered for future study. Temporal models are resistant to changes in threshold under roving-level conditions, but explained at most only 67% of the variance in the reproducible-noise data.

Adult↗

PubMed2006

Cues for masked amplitude-modulation detection.

The ability of psychoacoustic models to predict listeners' performance depends on two key stages: preprocessing and the generation of a decision variable. The goal of the current study was to determine the perceptually relevant decision variables in masked amplitude-modulation detection tasks in which the modulation depth of the masker was systematically varied. Potential cues were made unreliable by roving the overall modulation depth from trial to trial or were reduced in salience by equalizing the envelope energy of the standard and target after the signal was added. Listeners' performance was significantly degraded in both paradigms compared to the baseline (fixed-level modulation masker) condition, which was similar to those used in previous studies of masking in the envelope-frequency domain. Although this observation was broadly consistent with a simple long-term envelope power-spectrum model, there were several aspects of the data that were not. For example, the steep rate of change in threshold with masker depth and the fact that an optimal amount of envelope noise could enhance performance were not predicted by decision variables calculated directly from the stimulus envelope. A physiologically based processing model suggested a realistic nonlinear mechanism that could give rise to these second-order features of the data.

Audiometry, Pure-Tone↗

PubMed2006

Predictions of formant-frequency discrimination in noise based on model auditory-nerve responses.

To better understand how the auditory system extracts speech signals in the presence of noise, discrimination thresholds for the second formant frequency were predicted with simulations of auditory-nerve responses. These predictions employed either average-rate information or combined rate and timing information, and either populations of model fibers tuned across a wide range of frequencies or a subset of fibers tuned to a restricted frequency range. In general, combined temporal and rate information for a small population of model fibers tuned near the formant frequency was most successful in replicating the trends reported in behavioral data for formant-frequency discrimination. To explore the nature of the temporal information that contributed to these results, predictions based on model auditory-nerve responses were compared to predictions based on the average rates of a population of cross-frequency coincidence detectors. These comparisons suggested that average response rate (count) of cross-frequency coincidence detectors did not effectively extract important temporal information from the auditory-nerve population response. Thus, the relative timing of action potentials across auditory-nerve fibers tuned to different frequencies was not the aspect of the temporal information that produced the trends in formant-frequency discrimination thresholds.

Acoustic Stimulation↗

PubMed2005

The spontaneous-rate histogram of the auditory nerve can be explained by only two or three spontaneous rates and long-range dependence.

Estimates of the spontaneous discharge rate (SR) of auditory-nerve (AN) fibers are often based on measurements of the average rate over a long (e.g., 30 s) interval. These measurements are important because SR is apparently correlated with other AN properties, such as threshold to acoustic stimuli, shape of rate-level function, recovery from prior stimulation, and certain anatomical characteristics. Furthermore, histograms of SR estimates from large numbers of fibers suggest that they can be divided into two (i.e., low and high) or three (i.e., low, medium, and high) SR classes. Yet, even "simple" statistical estimates, such as average rate, can behave surprisingly poorly for processes with long-range dependence (LRD), which has been found in the spontaneous activity of AN fibers. In particular, LRD greatly increases the variability of estimates of mean discharge rate. We investigated the implications of this effect of LRD for our understanding of the SRs of AN fibers. The fractional-Gaussian-noise-driven Poisson process (fGnDP) was originally developed to model the LRD action-potential trains of AN fibers. Using rate estimates computed from this model, we were able to reproduce the shape of published histograms of SR using only three fixed SR values. Moreover, by using a Poisson-equivalent integrate-and-fire (IF) model in place of the inhomogeneous Poisson process in the fGnDP model, we were able to reproduce SR histograms using only two fixed SR values. These results suggest that AN fibers may have only two or three possible values for their long-term average spontaneous discharge rates. In other words, all "high-SR" neurons may actually have the same underlying SR. Furthermore, both "low-SR" and "medium-SR" neurons may have a single "true" SR value, or these two classes may have two different "true" SR values. Furthermore, the Poisson-equivalent IF model may prove useful in other applications involving the modeling of trains of action potentials.

Action Potentials↗

PubMed2005

Control of cellular pattern formation in the vertebrate inner retina by homotypic regulation of cell-fate decisions.

The vertebrate retina is composed of cellular arrays that are nonrandom across two-dimensional space. The determinants of these nonrandom two-dimensional cellular patterns in the inner nuclear layer of the retina were investigated using empirical and computational modeling techniques. In normal and experimental models of goldfish retinal growth, the patterns of tyrosine hydroxylase- and serotonin-positive cells indicated that neither cell death nor lateral migration of differentiated cells were dominant mechanisms of cellular pattern formation. A computational model of cellular pattern formation that used a signaling mechanism arising from differentiated cells that inhibited homotypic cell-fate decisions generated accurate simulations of the empirically observed patterns in normal retina. This model also predicted the principal atypical cellular pattern characteristic, a transient cell-type-specific hyperplasia, which was empirically observed in the growing retina subsequent to selective ablation of differentiated retinal cells, either tyrosine hydroxylase positive or serotonin positive. The results support the hypothesis that inhibitory spatiotemporal regulation of homotypic cell-fate decisions is a dominant mechanistic determinant of nonrandom cellular patterns in the vertebrate retina.

5,7-Dihydroxytryptamine↗

PubMed2005

A model for interaural time difference sensitivity in the medial superior olive: interaction of excitatory and inhibitory synaptic inputs, channel dynamics, and cellular morphology.

This study reports simulations of recent physiological results from the gerbil medial superior olive (MSO) that reveal that blocking glycinergic inhibition can shift the tuning for the interaural time difference (ITD) of the cell (Brand et al., 2002). Our simulations indicate that the model proposed in the study by Brand et al. (2002) requires precisely timed, short-duration inhibition with temporal accuracy exceeding that described in the auditory system. An alternative model is proposed that incorporates two anatomic observations in the MSO: (1) the axon arises from the dendrite that receives ipsilateral inputs; and (2) inhibitory synapses are located primarily on the soma in adult animals. When the inhibitory currents are activated or blocked, the model cell successfully simulates experimentally observed shifts in the best ITD. The asymmetrical cell structure allows an imbalance between the ipsilateral and contralateral excitatory inputs and shifts the ITD curve such that the best ITD is not at zero. Fine adjustment of the best ITD is achieved by the interplay of somatic sodium currents and synaptic inhibitory currents. The shift of the best ITD in the model is limited to approximately 0.2 ms, which is behaviorally significant with respect to ITDs encountered in perceptual tasks. The model suggests a mechanism for dynamically "fine-tuning" the ITD sensitivity of MSO cells by the opponency between depolarizing sodium currents and hyperpolarizing inhibitory currents.

Animals↗

PubMed2005

Effects of inhibitory feedback in a network model of avian brain stem.

The avian auditory brain stem consists of a network of specialized nuclei, including nucleus laminaris (NL) and superior olivary nucleus (SON). NL cells show sensitivity to interaural time difference (ITD), a critical cue that underlies spatial hearing. SON cells provide inhibitory feedback to the rest of the network. Empirical data suggest that feedback inhibition from SON could increase the ITD sensitivity of NL across sound level. Using a bilateral network model, we assess the effects of SON feedback inhibition. Individual cells are specified as modified leaky-integrate-and-fire neurons with time constants and thresholds that vary with inhibitory input. Acoustic sound level is reflected in the discharge rates of the model auditory-nerve fibers, which innervate the network. Simulations show that with SON inhibitory feedback, ITD sensitivity is maintained in model NL cells over a threefold range in auditory-nerve discharge rate. In contrast, without SON feedback inhibition, ITD sensitivity is significantly reduced as input rates are increased. Feedback inhibition is most beneficial in maintaining ITD sensitivity at high-input rates (simulating high sound levels). With SON inhibition, ITD sensitivity is maintained for both interaurally balanced inputs (simulating an on-center sound source) and interaurally imbalanced inputs (simulating a lateralized source). Further, the empirically observed temporal build-up of SON inhibition and the presence of reciprocal inhibitory connections between the ipsi- and contralateral SON both improve ITD sensitivity. In sum, our network model shows that inhibitory feedback can substantially increase the sensitivity and dynamic range of ITD coding in the avian auditory brain stem.

Action Potentials↗

PubMed2005

Encoding of vowel-like sounds in the auditory nerve: model predictions of discrimination performance.

The sensitivity of listeners to changes in the center frequency of vowel-like harmonic complexes as a function of the center frequency of the complex cannot be explained by changes in the level of the stimulus [Lyzenga and Horst, J. Acoust. Soc. Am. 98, 1943-1955 (1995)]. Rather, a complex pattern of sensitivity is seen; for a spectrum with a triangular envelope, the greatest sensitivity occurs when the center frequency falls between harmonics, whereas for a spectrum with a trapezoidal envelope, greatest sensitivity occurs when the center frequency is aligned with a harmonic. In this study, the thresholds of a population model of auditory-nerve (AN) fibers were quantitatively compared to these trends in psychophysical thresholds. Single-fiber and population model responses were evaluated in terms of both average discharge rate and the combination of rate and timing information. Results indicate that phase-locked responses of AN fibers encode phase transitions associated with minima in these amplitude-modulated stimuli. The temporal response properties of a single AN fiber, tuned to a frequency slightly above the center frequency of the harmonic complex, were able to explain the trends in thresholds for both triangular- and trapezoidal-shaped spectra.

Acoustic Stimulation↗

PubMed2005

Analysis of models for the synapse between the inner hair cell and the auditory nerve.

A general mathematical approach was proposed to study phenomenological models of the inner-hair-cell and auditory-nerve (AN) synapse complex. Two models (Meddis, 1986; Westerman and Smith, 1988) were studied using this unified approach. The responses of both models to a constant-intensity stimulus were described mathematically, and the relationship between model parameters and response characteristics was investigated. The mathematical descriptions of the two models were essentially equivalent despite their structural differences. This analytical approach was used to study the effects of adaptation characteristics on model parameters and of model parameters on adaptation characteristics. The results provided insights into these models and the underlying biophysical processing. This analytical method was also used to study offset adaptation, and it was found that the offset adaptation of both models was limited by the models' structures. A modified version of the synapse model, which has the same onset adaptation but improved offset adaptation, is proposed here. This modified synapse model produces more physiologically realistic offset adaptation and also enhances the modulation gain of model AN fiber responses, consistent with AN physiology.

Acoustic Stimulation↗

PubMed2005

Response properties of an integrate-and-fire model that receives subthreshold inputs.

A computational technique is described for calculation of the interspike interval and poststimulus time histograms for the responses of an integrate-and-fire model to arbitrary inputs. The effects of the model parameters on the response statistics were studied systematically. Specifically, the probability distribution of the membrane potential was calculated as a function of time, and the mean interspike interval and PST histogram were calculated for arbitrary inputs. For stationary inputs, the regularity of the output was studied in detail for various model parameters. For nonstationary inputs, the effects of the model parameters on the output synchronization index were explored. The results show that enhanced synchronization in response to low-frequency stimuli required a large number (n > 25) of weak inputs. Irregular responses and a linear input-output rate relationship required strong (but subthreshold) inputs with a small time constant. A model cell with mixed-amplitude synaptic inputs can respond to stationary inputs irregularly and have enhanced synchronization to nonstationary inputs that are phase-locked to low-frequency inputs. Both of these response properties have been reported for some cells in the ventral cochlear nucleus in the auditory brainstem.

Action Potentials↗

PubMed2004

Cellular patterns in the inner retina of adult zebrafish: quantitative analyses and a computational model of their formation.

The mechanisms that control cellular pattern formation in the growing vertebrate central nervous system are poorly understood. In an effort to reveal mechanistic rules of cellular pattern formation in the central nervous system, quantitative spatial analysis and computational modeling techniques were applied to cellular patterns in the inner retina of the adult zebrafish. All the analyzed cell types were arrayed in nonrandom patterns tending toward regularity; specifically, they were locally anticlustered. Over relatively large spatial scales, only one cell type exhibited consistent evidence for pattern regularity, suggesting that cellular pattern formation in the inner retina is dominated by local anticlustering mechanisms. Cross-correlation analyses revealed independence between the patterns of different cell types, suggesting that cellular pattern formation may involve multiple, independent, homotypic anticlustering mechanisms. A computational model of cellular pattern formation in the growing zebrafish retina was developed, which featured an inhibitory, homotypic signaling mechanism, arising from differentiated cells, that controlled the spatial profile of cell fate decisions. By adjusting the spatial profile of this decaying-exponential signal, the model provided good estimates of all the cellular patterns that were observed in vivo, as objectively judged by quantitative spatial pattern analyses. The results support the hypothesis that cellular pattern formation in the inner retina of zebrafish is dominated by a set of anticlustering mechanisms that may control events at, or near, the spatiotemporal point of cell fate decision.

Animals↗

PubMed2004

A phenomenological model of peripheral and central neural responses to amplitude-modulated tones.

A phenomenological model with time-varying excitation and inhibition was developed to study possible neural mechanisms underlying changes in the representation of temporal envelopes along the auditory pathway. A modified version of an existing auditory-nerve model [Zhang et al., J. Acoust. Soc. Am. 109, 648-670 (2001)] was used to provide inputs to higher hypothetical processing centers. Model responses were compared directly to published physiological data at three levels: the auditory nerve, ventral cochlear nucleus, and inferior colliculus. Trends and absolute values of both average firing rate and synchrony to the modulation period were accurately predicted at each level for a wide range of stimulus modulation depths and modulation frequencies. The diversity of central physiological responses was accounted for with realistic variations of model parameters. Specifically, enhanced synchrony in the cochlear nucleus and rate-tuning to modulation frequency in the inferior colliculus were predicted by choosing appropriate relative strengths and time courses of excitatory and inhibitory inputs to postsynaptic model cells. The proposed model is fundamentally different than others that have been used to explain the representation of envelopes in the mammalian midbrain, and it provides a computational tool for testing hypothesized relationships between physiology and psychophysics.

Acoustics↗

PubMed2003

Quantifying the information in auditory-nerve responses for level discrimination.

An analytical approach for quantifying the information in auditory-nerve (AN) fiber responses for the task of level discrimination is described. A simple analytical model for ANT responses is extended to include temporal response properties, including the nonlinear-phase effects of the cochlear amplifier. Use of simple analytical models for AN discharge patterns allows quantification of the contributions of level-dependent aspects of the patterns to level discrimination. Specifically, the individual and combined contributions of the information contained in discharge rate, synchrony, and relative phase cues are explicitly examined for level discrimination of tonal stimuli. It is shown that the rate information provided by individual AN fibers is more constrained by increases in variance with increases in rate than by saturation. As noted in previous studies, there is sufficient average-rate information within a narrow-CF region to account for robust behavioral performance over a wide dynamic range; however, there is no model based on a simple limitation or use of AN information consistent with parametric variations in performance. This issue is explored in the current study through analysis of performance based on different aspects of AN patterns. For example, we show that performance predicted from use of all rate information degrades significantly as level increases above low-medium levels, inconsistent with Weber's Law. At low frequencies, synchrony information extends the range over which behavioral performance can be explained by 10-15 dB, but only at low levels. In contrast to rate and synchrony, nonlinear-phase cues are shown to provide robust information at medium and high levels in near-CF fibers for low-frequency stimuli. The level dependence of the discharge rate and phase properties of AN fibers are influenced by the compressive nonlinearity of the inner ear. Evaluating the role of the compressive nonlinearity in level coding is important for understanding neural encoding mechanisms and because of its association with the cochlear amplifier, which is a fragile aspect of the ear believed to be affected in common forms of hearing impairment.

Animals↗

PubMed2003

CS-dependent response probability in an auditory masked-detection task: considerations based on models of Pavlovian conditioning.

Experimental studies were performed using a Pavlovian-conditioned eyeblink response to measure detection of a variable-sound-level tone (T) in a fixed-sound-level masking noise (N) in rabbits. Results showed an increase in the asymptotic probability of conditioned responses (CRs) to the reinforced TN trials and a decrease in the asymptotic rate of eyeblink responses to the non-reinforced N presentations as a function of the sound level of the T. These observations are consistent with expected behaviour in an auditory masked detection task, but they are not consistent with predictions from a traditional application of the Rescorla-Wagner or Pearce models of associative learning. To implement these models, one typically considers only the actual stimuli and reinforcement on each trial. We found that by considering perceptual interactions and concepts from signal detection theory, these models could predict the CS dependence on the sound level of the T. In these alternative implementations, the animals response probabilities were used as a guide in making assumptions about the "effective stimuli".

Animals↗