Auditory Research publications

Temporal variation in listeners' sensitivity to interaural time and level differences (ITD and ILD) was assessed using the temporal weighting function (TWF) paradigm [Stecker and Hafter (2002). J. Acoust. Soc. Am. 112, 1046–1057] in the context of sound-source lateralization. Brief Gabor click trains were presented over headphones with overall ITD and/or ILD ranging ±500 μs ITD and/or ±5 dB ILD across trials; values for individual clicks within each train varied by an additional ±100 μs or ±2 dB to allow TWF calculation by multiple regression. In separate conditions, TWFs were measured for (i) ITD alone, (ii) ILD alone, (iii) ITD and ILD covarying (“in agreement”), and (iv) ITD and ILD varying independently across clicks. Consistent with past studies that measured TWF for binaural discrimination, TWFs demonstrated high weight on the first click for stimuli with short interclick interval (ICI = 2 ms), but flatter weighting for longer ICI (5–10 ms). Some conditions additionally demonstrated greater weight for clicks near the offset than near the middle of the train [Stecker and Hafter (2009). J. Acoust. Soc. Am. 125, 3914–3924]. The latter result was observed only when stimuli carried ILD, and appeared more reliably for 5 ms than for 2 or 10 ms ICI.

While it is commonly held that the capacity to learn is greatest in the young, there have been few direct comparisons of the response to training across age groups. Here, adolescents (11–17 years, n = 20) and adults (≥18 years, n = 11) practiced detecting a backward-masked tone for ∼1 h/day for 10 days. Nearly every adult, but only half of the adolescents improved across sessions, and the adolescents who learned did so more slowly than adults. Nevertheless, the adolescent and adult learners showed the same generalization pattern, improving on untrained backward- but not forward- or simultaneous-masking conditions. Another subset of adolescents (n = 6) actually got worse on the trained condition. This worsening, unlike learning, generalized to an untrained forward-masking, but not backward-masking condition. Within sessions, both age groups got worse, but the worsening was greater for adolescents. These maturational changes in the response to training largely followed those previously reported for temporal-interval discrimination. Overall, the results suggest that late-maturing processes affect the response to perceptual training and that some of these processes may be shared between tasks. Further, the different developmental rates for learning and generalization, and different generalization patterns for learning and worsening imply that learning, generalization, and worsening may have different origins.

This study compared the dependence of comodulation masking release (CMR) and monaural envelope correlation perception (MECP) on the degree of envelope correlation for the same narrowband noise stimuli. Envelope correlation across noise bands was systematically varied by mixing independent bands with a base set of comodulated bands. The magnitude of CMR fell monotonically with reductions in envelope correlation, and CMR varied over a range of envelope correlations that were not discriminable from each other in the MECP paradigm. For complexes of 100-Hz-wide noise bands, discrimination thresholds in the MECP task were similar whether the standard was a comodulated set of noise bands or a completely independent set of noise bands. This was not the case for 25-Hz-wide noise bands. Although the data demonstrate that CMR and MECP exhibit different dependencies on the degree of envelope correlation, some commonality across the two phenomena was observed. Specifically, for 25-Hz-wide bands of noise, there was a robust relationship between individual listeners' sensitivity to decorrelation from an otherwise comodulated set of noise bands and the magnitude of CMR measured for those same comodulated noise bands.

The precedence effect (PE) refers to the dominance of directional information carried by a direct sound (lead) over the spatial information contained in its multiple reflections (lags) in sound localization. Although the processes underlying the PE have been largely investigated, the extent to which peripheral versus central auditory processes contribute to this perceptual phenomenon has remained unclear. The present study investigated the contribution of peripheral processing to the PE through a comparison of physiological and psychoacoustical data in the same human listeners. The psychoacoustical experiments, comprising a fusion task, an interaural time difference detection task and a lateralization task, demonstrated a time range from 1 to 4.6-5 ms, in which the PE operated (precedence window). Click-evoked otoacoustic emissions (CEOAEs) were recorded in both ears to investigate the lead-lag interactions at the level of the basilar membrane (BM) in the cochlea. The CEOAE-derived peripheral and monaural lag suppression was largest for ICIs of 1-4 ms. Auditory-evoked brainstem responses (ABRs) were used to investigate monaural and binaural lag suppression at the brainstem level. The responses to monaural stimulation reflected the peripheral lag suppression observed in the CEOAE results, while the binaural brainstem responses did not show any substantial contribution of binaural processes to monaural lag suppression. The results demonstrated that the lag suppression occurring at the BM in a time range from 1 to 4 ms, as indicated by the suppression of the lag-CEOAE, was the source of the reduction in the lag-ABRs and a possible peripheral contributor to the PE for click stimuli.

The auditory cortex is malleable by experience. Previous studies of auditory plasticity have described experience-dependent changes in response profiles of single neurons or changes in global tonotopic organization. However, experience-dependent changes in the dynamics of local neural populations have remained unexplored. In this study, we examined the influence of a dramatic yet natural experience in the life of female mice, giving birth and becoming a mother on single neurons and neuronal ensembles in the primary auditory cortex (A1). Using in vivo two-photon calcium imaging and electrophysiological recordings from layer 2/3 in A1 of mothers and age-matched virgin mice, we monitored changes in the responses to a set of artificial and natural sounds. Population dynamics underwent large changes as measured by pairwise and higher-order correlations, with noise correlations increasing as much as twofold in lactating mothers. Concomitantly, changes in response properties of single neurons were modest and selective. Remarkably, despite the large changes in correlations, information about stimulus identity remained essentially the same in the two groups. Our results demonstrate changes in the correlation structure of neuronal activity as a result of a natural life event.

The dorsal lateral geniculate nucleus (dLGN) receives visual information from the retina and transmits it to the cortex. In this study, we made extracellular recordings in the dLGN of both anesthetized and awake mice, and found that a surprisingly high proportion of cells were selective for stimulus orientation. The orientation selectivity of dLGN cells was unchanged after silencing the visual cortex pharmacologically, indicating that it is not due to cortical feedback. The orientation tuning of some dLGN cells correlated with their elongated receptive fields, while in others orientation selectivity was observed despite the fact that their receptive fields were circular, suggesting that their retinal input might already be orientation selective. Consistently, we revealed orientation/axis-selective ganglion cells in the mouse retina using multielectrode arrays in an in vitro preparation. Furthermore, the orientation tuning of dLGN cells was largely maintained at different stimulus contrasts, which could be sufficiently explained by a simple linear feedforward model. We also compared the degree of orientation selectivity in different visual structures under the same recording condition. Compared with the dLGN, orientation selectivity is greatly improved in the visual cortex, but is similar in the superior colliculus, another major retinal target. Together, our results demonstrate prominent orientation selectivity in the mouse dLGN, which may potentially contribute to visual processing in the cortex.

Illusions are effective tools for the study of the neural mechanisms underlying perception because neural responses can be correlated to the physical properties of stimuli and the subject's perceptions. The F ranssen illusion ( FI ) is an auditory spatial illusion evoked by presenting a transient, abrupt tone and a slowly rising, sustained tone of the same frequency simultaneously on opposite sides of the subject. Perception of the FI consists of hearing a single sound, the sustained tone, on the side that the transient was presented. Both subcortical and cortical mechanisms for the FI have been proposed, but, to date, there is no direct evidence for either. The data show that humans and rhesus monkeys perceive the FI similarly. Recordings were taken from single units of the inferior colliculus in the monkey while they indicated the perceived location of sound sources with their gaze. The results show that the transient component of the F ranssen stimulus, with a shorter first spike latency and higher discharge rate than the sustained tone, encodes the perception of sound location. Furthermore, the persistent erroneous perception of the sustained stimulus location is due to continued excitation of the same neurons, first activated by the transient, by the sustained stimulus without location information. These results demonstrate for the first time, on a trial‐by‐trial basis, a correlation between perception of an auditory spatial illusion and a subcortical physiological substrate.

Much of what we know regarding the effect of stimulus repetition on neuroelectric adaptation comes from studies using artificially produced pure tones or harmonic complex sounds. Little is known about the neural processes associated with the representation of everyday sounds and how these may be affected by aging. In this study, we used real life, meaningful sounds presented at various azimuth positions and found that auditory evoked responses peaking at about 100 and 180 ms after sound onset decreased in amplitude with stimulus repetition. This neural adaptation was greater in young than in older adults and was more pronounced when the same sound was repeated at the same location. Moreover, the P2 waves showed differential patterns of domain-specific adaptation when location and identity was repeated among young adults. Background noise decreased ERP amplitudes and modulated the magnitude of repetition effects on both the N1 and P2 amplitude, and the effects were comparable in young and older adults. These findings reveal an age-related difference in the neural processes associated with adaptation to meaningful sounds, which may relate to older adults' difficulty in ignoring task-irrelevant stimuli.

Objective: Chronic neural recording in behaving animals is an essential method for studies of neural circuit function. However, stable recordings from small, densely packed neurons remains challenging, particularly over time-scales relevant for learning. Approach: We describe an assembly method for a 16-channel electrode array consisting of carbon fibers (<5 µm diameter) individually insulated with Parylene-C and fire-sharpened. The diameter of the array is approximately 26 µm along the full extent of the implant. Main results: Carbon fiber arrays were tested in HVC (used as a proper name), a song motor nucleus, of singing zebra finches where individual neurons discharge with temporally precise patterns. Previous reports of activity in this population of neurons have required the use of high impedance electrodes on movable microdrives. Here, the carbon fiber electrodes provided stable multi-unit recordings over time-scales of months. Spike-sorting indicated that the multi-unit signals were dominated by one, or a small number of cells. Stable firing patterns during singing confirmed the stability of these clusters over time-scales of months. In addition, from a total of 10 surgeries, 16 projection neurons were found. This cell type is characterized by sparse stereotyped firing patterns, providing unambiguous confirmation of single cell recordings. Significance: Carbon fiber electrode bundles may provide a scalable solution for long-term neural recordings of densely packed neurons.

Sound streams were generated by randomly choosing the levels of tone pips from two different distributions, A and B. Of the 18 tone pips, the first nine were drawn from distribution A and the second nine from distribution B, or the opposite. The listeners' task was to indicate order, A-B or B-A. In two conditions the A and B distributions differed in mean (condition 1) or variance (condition 2). In contrast to an ideal observer, listeners' strategies were consistent across the two conditions. Analyses suggest that listeners relied primarily on the more intense tone pips in making their decisions.

Mutations of SLC26A4 are a common cause of human hearing loss associated with enlargement of the vestibular aqueduct. SLC26A4 encodes pendrin, an anion exchanger expressed in a variety of epithelial cells in the cochlea, the vestibular labyrinth and the endolymphatic sac. Slc26a4 (Δ/Δ) mice are devoid of pendrin and develop a severe enlargement of the membranous labyrinth, fail to acquire hearing and balance, and thereby provide a model for the human phenotype. Here, we generated a transgenic mouse line that expresses human SLC26A4 controlled by the promoter of ATP6V1B1. Crossing this transgene into the Slc26a4 (Δ/Δ) line restored protein expression of pendrin in the endolymphatic sac without inducing detectable expression in the cochlea or the vestibular sensory organs. The transgene prevented abnormal enlargement of the membranous labyrinth, restored a normal endocochlear potential, normal pH gradients between endolymph and perilymph in the cochlea, normal otoconia formation in the vestibular labyrinth and normal sensory functions of hearing and balance. Our study demonstrates that restoration of pendrin to the endolymphatic sac is sufficient to restore normal inner ear function. This finding in conjunction with our previous report that pendrin expression is required for embryonic development but not for the maintenance of hearing opens the prospect that a spatially and temporally limited therapy will restore normal hearing in human patients carrying a variety of mutations of SLC26A4.

The mechanisms by which functional maps and map plasticity contribute to cortical computation remain controversial. Recent studies have revisited the theory of neural Darwinism to interpret the learning-induced map plasticity and neuronal heterogeneity observed in the cortex. Here, we hypothesize that the Darwinian principle provides a substrate to explain the relationship between neuron heterogeneity and cortical functional maps. We demonstrate in the rat auditory cortex that the degree of response variance is closely correlated with the size of its representational area. Further, we show that the response variance within a given population is altered through training. These results suggest that larger representational areas may help to accommodate heterogeneous populations of neurons. Thus, functional maps and map plasticity are likely to play essential roles in Darwinian computation, serving as effective, but not absolutely necessary, structures to generate diverse response properties within a neural population.

Perception of complex sounds such as speech is affected by a variety of factors, including attention, expectation of reward, physiological state, and/or disorders, yet the mechanisms underlying this modulation are not well understood. Although dopamine is commonly studied for its role in reward-based learning and in disorders, multiple lines of evidence suggest that dopamine is also involved in modulating auditory processing. In this study, we examined the effects of dopamine application on neuronal response properties in the inferior colliculus (IC) of awake mice. Because the IC contains dopamine receptors and nerve terminals immunoreactive for tyrosine hydroxylase, we predicted that dopamine would modulate auditory responses in the IC. We recorded single-unit responses before, during, and after the iontophoretic application of dopamine using piggyback electrodes. We examined the effects of dopamine on firing rate, timing, and probability of bursting. We found that application of dopamine affected neural responses in a heterogeneous manner. In more than 80 % of the neurons, dopamine either increased (32 %) or decreased (50 %) firing rate, and the effects were similar on spontaneous and sound-evoked activity. Dopamine also either increased or decreased first spike latency and jitter in almost half of the neurons. In 3/28 neurons (11 %), dopamine significantly altered the probability of bursting. The heterogeneous effects of dopamine observed in the IC of awake mice were similar to effects observed in other brain areas. Our findings indicate that dopamine differentially modulates neural activity in the IC and thus may play an important role in auditory processing.

In a complex auditory scene, a "cocktail party" for example, listeners can disentangle multiple competing sequences of sounds. A recent psychophysical study in our laboratory demonstrated a robust spatial component of stream segregation showing ∼8° acuity. Here, we recorded single- and multiple-neuron responses from the primary auditory cortex of anesthetized cats while presenting interleaved sound sequences that human listeners would experience as segregated streams. Sequences of broadband sounds alternated between pairs of locations. Neurons synchronized preferentially to sounds from one or the other location, thereby segregating competing sound sequences. Neurons favoring one source location or the other tended to aggregate within the cortex, suggestive of modular organization. The spatial acuity of stream segregation was as narrow as ∼10°, markedly sharper than the broad spatial tuning for single sources that is well known in the literature. Spatial sensitivity was sharpest among neurons having high characteristic frequencies. Neural stream segregation was predicted well by a parameter-free model that incorporated single-source spatial sensitivity and a measured forward-suppression term. We found that the forward suppression was not due to post discharge adaptation in the cortex and, therefore, must have arisen in the subcortical pathway or at the level of thalamocortical synapses. A linear-classifier analysis of single-neuron responses to rhythmic stimuli like those used in our psychophysical study yielded thresholds overlapping those of human listeners. Overall, the results indicate that the ascending auditory system does the work of segregating auditory streams, bringing them to discrete modules in the cortex for selection by top-down processes.

In the brainstem, the auditory system diverges into two pathways that process different sound localization cues, interaural time differences (ITDs) and level differences (ILDs). We investigated the site where ILD is detected in the auditory system of barn owls, the posterior part of the lateral lemniscus (LLDp). This structure is equivalent to the lateral superior olive in mammals. The LLDp is unique in that it is the first place of binaural convergence in the brainstem where monaural excitatory and inhibitory inputs converge. Using binaurally uncorrelated noise and a generalized linear model, we were able to estimate the spectrotemporal tuning of excitatory and inhibitory inputs to these cells. We show that the response of LLDp neurons is highly locked to the stimulus envelope. Our data demonstrate that spectrotemporally tuned, temporally delayed inhibition enhances the reliability of envelope locking by modulating the gain of LLDp neurons' responses. The dependence of gain modulation on ILD shown here constitutes a means for space-dependent coding of stimulus identity by the initial stages of the auditory pathway.

Two experiments, both presenting diotic, harmonic tone complexes (100 Hz fundamental), were conducted to explore the envelope-related component of the frequency-following response (FFRENV), a measure of synchronous, subcortical neural activity evoked by a periodic acoustic input. Experiment 1 directly compared two common analysis methods, computing the magnitude spectrum and the phase-locking value (PLV). Bootstrapping identified which FFRENV frequency components were statistically above the noise floor for each metric and quantified the statistical power of the approaches. Across listeners and conditions, the two methods produced highly correlated results. However, PLV analysis required fewer processing stages to produce readily interpretable results. Moreover, at the fundamental frequency of the input, PLVs were farther above the metric's noise floor than spectral magnitudes. Having established the advantages of PLV analysis, the efficacy of the approach was further demonstrated by investigating how different acoustic frequencies contribute to FFRENV, analyzing responses to complex tones composed of different acoustic harmonics of 100 Hz (Experiment 2). Results show that the FFRENV response is dominated by peripheral auditory channels responding to unresolved harmonics, although low-frequency channels driven by resolved harmonics also contribute. These results demonstrate the utility of the PLV for quantifying the strength of FFRENV across conditions.

The aim of this study is to further explore the relationship between distortion-product otoacoustic emission (DPOAE) measurements and categorical loudness scaling (CLS) measurements using multiple linear regression (MLR) analysis. Recently, Thorson et al. [J. Acoust. Soc. Am. 131, 1282–1295 (2012)] obtained predictions of CLS loudness ratings from DPOAE input/output (I/O) functions using MLR analysis. The present study extends that work by (1) considering two different (and potentially improved) MLR models, one for predicting loudness rating at specified input level and the other for predicting the input level for each loudness category and (2) validating the new models' predictions using an independent set of data. Strong correlations were obtained between predicted and measured data during the validation process with overall root-mean-square errors in the range 10.43–16.78 dB for the prediction of CLS input level, supporting the view that DPOAE I/O measurements can predict CLS loudness ratings and input levels, and thus may be useful for fitting hearing aids.

The neurophonic potential is a synchronized frequency-following extracellular field potential that can be recorded in the nucleus laminaris (NL) in the brainstem of the barn owl. Putative generators of the neurophonic are the afferent axons from the nucleus magnocellularis, synapses onto NL neurons, and spikes of NL neurons. The outputs of NL, i.e., action potentials of NL neurons, are only weakly represented in the neurophonic. Instead, the inputs to NL, i.e., afferent axons and their synaptic potentials, are the predominant origin of the neurophonic (Kuokkanen PT, Wagner H, Ashida G, Carr CE, Kempter R. J Neurophysiol 104: 2274–2290, 2010). Thus in NL the monaural inputs from the two brain sides converge and create a binaural neurophonic. If these monaural inputs contribute independently to the extracellular field, the response to binaural stimulation can be predicted from the sum of the responses to ipsi- and contralateral stimulation. We found that a linear summation model explains the dependence of the responses on interaural time difference as measured experimentally with binaural stimulation. The fit between model predictions and data was excellent, even without taking into account the nonlinear responses of NL coincidence detector neurons, although their firing rate and synchrony strongly depend on the interaural time difference. These results are consistent with the view that the afferent axons and their synaptic potentials in NL are the primary origin of the neurophonic.

Neuroprosthetics using intracortical microstimulation can potentially alleviate sensory deprivation due to injury or disease. However the information bandwidth of a single microstimulation channel remains largely unanswered. This paper presents three experiments that examine the importance of Peak Power/Charge and RMS Power/Charge for detection of acoustic and electrical Sinusoidal Amplitude Modulated stimuli by the auditory system. While the peripheral auditory system is sensitive to RMS power cues for the detection of acoustic stimuli, here we provide results that suggest that the auditory cortex is sensitive to peak charge cues for electrical stimuli. Varying the modulation frequency and depth do not change this effect for detection of modulated electrical stimuli.

Unlabelled: Opto-electronic computer holographic measurements were made of the tympanic membrane (TM) in cadaveric chinchillas. Measurements with two laser wavelengths were used to compute the 3D-shape of the TM. Single laser wavelength measurements locked to eight distinct phases of a tonal stimulus were used to determine the magnitude and the relative phase of the surface displacements. These measurements were made at over 250,000 points on the TM surface. The measured motions contained spatial phase variations consistent with relatively low-order (large spatial frequency) modal motions and smaller magnitude higher-order (smaller spatial frequency) motions that appear to travel, but may also be explained by losses within the membrane. The measurement of shape and thin shell theory allowed us to separate the measured motions into those components orthogonal to the plane of the tympanic ring, and those components within the plane of the tympanic ring based on the 3D-shape. The predicted in-plane motion components are generally smaller than the out-of-plane perpendicular component of motion. Since the derivation of in-plane and out-of plane depended primarily on the membrane shape, the relative sizes of the predicted motion components did not vary with frequency. Summary: A new method for simultaneously measuring the shape and sound-induced motion of the tympanic membrane is utilized to estimate the 3D motion on the membrane surface. This article is part of a special issue entitled "MEMRO 2012".