Effect of auditory efferent time-constant duration on speech recognition in noise.

The human auditory efferent system may play a role in improving speech-in-noise recognition with an associated range of time constants. Computational auditory models with efferent-inspired feedback demonstrate improved speech-in-noise recognition with long efferent time constants (2000 ms). This study used a similar model plus an Automatic Speech Recognition (ASR) system to investigate the role of shorter time constants. ASR speech recognition in noise improved with efferent feedback (compared to no-efferent feedback) for both short and long efferent time constants. For some signal-to-noise ratios, speech recognition in noise improved as efferent time constants were increased from 118 to 2000 ms.

Frequency Tuning of the Efferent Effect on Cochlear Gain in Humans.

Cochlear gain reduction via efferent feedback from the medial olivocochlear bundle is frequency specific (Guinan, Curr Opin Otolaryngol Head Neck Surg 18:447-453, 2010). The present study with humans used the Fixed Duration Masking Curve psychoacoustical method (Yasin et al., J Acoust Soc Am 133:4145-4155, 2013a; Yasin et al., Basic aspects of hearing: physiology and perception, pp 39-46, 2013b; Yasin et al., J Neurosci 34:15319-15326, 2014) to estimate the frequency specificity of the efferent effect at the cochlear level. The combined duration of the masker-plus-signal stimulus was 25 ms, within the efferent onset delay of about 31-43 ms (James et al., Clin Otolaryngol 27:106-112, 2002). Masker level (4.0 or 1.8 kHz) at threshold was obtained for a 4-kHz signal in the absence or presence of an ipsilateral 60 dB SPL, 160-ms precursor (200-Hz bandwidth) centred at frequencies between 2.5 and 5.5 kHz. Efferent-mediated cochlear gain reduction was greatest for precursors with frequencies the same as, or close to that of, the signal (gain was reduced by about 20 dB), and least for precursors with frequencies well removed from that of the signal (gain remained at around 40 dB). The tuning of the efferent effect filter (tuning extending 0.5-0.7 octaves above and below the signal frequency) is within the range obtained in humans using otoacoustic emissions (Lilaonitkul and Guinan, J Assoc Res Otolaryngol 10:459-470, 2009; Zhao and Dhar, J Neurophysiol 108:25-30, 2012). The 10 dB bandwidth of the efferent-effect filter at 4000 Hz was about 1300 Hz (Q(10) of 3.1). The FDMC method can be used to provide an unbiased measure of the bandwidth of the efferent effect filter using ipsilateral efferent stimulation.

Effect of human auditory efferent feedback on cochlear gain and compression.

The mammalian auditory system includes a brainstem-mediated efferent pathway from the superior olivary complex by way of the medial olivocochlear system, which reduces the cochlear response to sound (Warr and Guinan, 1979; Liberman et al., 1996). The human medial olivocochlear response has an onset delay of between 25 and 40 ms and rise and decay constants in the region of 280 and 160 ms, respectively (Backus and Guinan, 2006). Physiological studies with nonhuman mammals indicate that onset and decay characteristics of efferent activation are dependent on the temporal and level characteristics of the auditory stimulus (Bacon and Smith, 1991; Guinan and Stankovic, 1996). This study uses a novel psychoacoustical masking technique using a precursor sound to obtain a measure of the efferent effect in humans. This technique avoids confounds currently associated with other psychoacoustical measures. Both temporal and level dependency of the efferent effect was measured, providing a comprehensive measure of the effect of human auditory efferents on cochlear gain and compression. Results indicate that a precursor (>20 dB SPL) induced efferent activation, resulting in a decrease in both maximum gain and maximum compression, with linearization of the compressive function for input sound levels between 50 and 70 dB SPL. Estimated gain decreased as precursor level increased, and increased as the silent interval between the precursor and combined masker-signal stimulus increased, consistent with a decay of the efferent effect. Human auditory efferent activation linearizes the cochlear response for mid-level sounds while reducing maximum gain.

Improved psychophysical methods to estimate peripheral gain and compression.

It is possible that previous psychophysical estimates of basilar membrane gain and compression using temporal masking curve (TMC) and additivity of forward masking (AFM) methods using long-duration maskers (>30 ms) could have been affected by activation of the medial olivocochlear reflex (MOCR) (Jennings et al. ; Plack and Arifianto ). In experiment 1, AFM and TMC methods were compared to a new fixed-duration masking curve (FDMC) method in which the combined masker and signal stimulus duration is fixed at 25 ms. Estimates of compression were found to be not significantly different for TMC, FDMC and AFM methods. Estimates of gain were similar for TMC and FDMC methods. Maximum compression was associated with a significantly lower input masker level using the FDMC compared to the TMC method. In experiment 2, the FDMC method was used to investigate the effect of efferent activation on gain and compression estimates by presenting a precursor sound prior to the combined masker-signal stimulus. Estimated gain decreased as precursor level increased, and increased as the silent interval between the precursor and combined masker-signal stimulus increased, consistent with a decay of the efferent response.

Estimating peripheral gain and compression using fixed-duration masking curves.

Estimates of human basilar membrane gain and compression obtained using temporal masking curve (TMC) and additivity of forward masking (AFM) methods with long-duration maskers or long masker-signal silent intervals may be affected by olivocochlear efferent activation, which reduces basilar membrane gain. The present study introduces a fixed-duration masking curve (FDMC) method, which involves a comparison of off- and on-frequency forward masker levels at threshold as a function of masker and signal duration, with the total masker-signal duration fixed at 25 ms to minimize efferent effects. Gain and compression estimates from the FDMC technique were compared with those from TMC (104-ms maskers) and AFM (10- and 200-ms maskers) methods. Compression estimates over an input-masker range of 40-60 dB sound pressure level were similar for the four methods. Maximum compression occurred at a lower input level for the FDMC compared to the TMC method. Estimates of gain were similar for TMC and FDMC methods. The FDMC method may provide a more reliable estimate of BM gain and compression in the absence of efferent activation and could be a useful method for estimating effects of efferent activity when used with a precursor sound (to trigger efferent activation), presented prior to the combined masker-signal stimulus.

Using visual direction in three-dimensional motion perception.

The eyes receive slightly different views of the world, and the differences between their images (binocular disparity) are used to see depth. Several authors have suggested how the brain could exploit this information for three-dimensional (3D) motion perception, but here we consider a simpler strategy. Visual direction is the angle between the direction of an object and the direction that an observer faces. Here we describe human behavioral experiments in which observers use visual direction, rather than binocular information, to estimate an object's 3D motion even though this causes them to make systematic errors. This suggests that recent models of binocular 3D motion perception may not reflect the strategies that human observers actually use.

Estimates of compression at low and high frequencies using masking additivity in normal and impaired ears.

Auditory compression was estimated at 250 and 4000 Hz by using the additivity of forward masking technique, which measures the effects on signal threshold of combining two temporally nonoverlapping forward maskers. The increase in threshold in the combined-masker condition compared to the individual-masker conditions can be used to estimate compression. The signal was a 250 or 4000 Hz tone burst and the maskers (M1 and M2) were bands of noise. Signal thresholds were measured in the presence of M1 and M2 alone and combined for a range of masker levels. The results were used to derive response functions at each frequency. The procedure was conducted with normal-hearing and hearing-impaired listeners. The results suggest that the response function in normal ears is similar at 250 and 4000 Hz with a mid level compression exponent of about 0.2. However, compression extends over a smaller range of levels at 250 Hz. The results confirm previous estimates of compression using temporal masking curves (TMCs) without assuming a linear off-frequency reference as in the TMC procedure. The impaired ears generally showed less compression. Importantly, some impaired ears showed a linear response at 250 Hz, providing a further indication that low-frequency compression originates in the cochlea.

Masking by inaudible sounds and the linearity of temporal summation.

Many natural sounds, including speech and animal vocalizations, involve rapid sequences that vary in spectrum and amplitude. Each sound within a sequence has the potential to affect the audibility of subsequent sounds in a process known as forward masking. Little is known about the neural mechanisms underlying forward masking, particularly in more realistic situations in which multiple sounds follow each other in rapid succession. A parsimonious hypothesis is that the effects of consecutive sounds combine linearly, so that the total masking effect is a simple sum of the contributions from the individual maskers. The experiment reported here tests a counterintuitive prediction of this linear-summation hypothesis, namely that a sound that itself is inaudible should, under certain circumstances, affect the audibility of subsequent sounds. The results show that, when two forward maskers are combined, the second of the two maskers can continue to produce substantial masking, even when it is completely masked by the first masker. Thus, inaudible sounds can affect the perception of subsequent sounds. A model incorporating instantaneous compression (reflecting the nonlinear response of the basilar membrane in the cochlea), followed by linear summation of the effects of the maskers, provides a good account of the data. Despite the presence of multiple sources of nonlinearity in the auditory system, masking effects by sequential sounds combine in a manner that is well captured by a time-invariant linear system.

Effect of efferent activation on binaural frequency selectivity.

Binaural notched-noise experiments indicate a reduced frequency selectivity of the binaural system compared to monaural processing. The present study investigates how auditory efferent activation (via the medial olivocochlear system) affects binaural frequency selectivity in normal-hearing listeners. Thresholds were measured for a 1-kHz signal embedded in a diotic notched-noise masker for various notch widths. The signal was either presented in phase (diotic) or in antiphase (dichotic), gated with the noise. Stimulus duration was 25 ms, in order to avoid efferent activation due to the masker or the signal. A bandpass-filtered noise precursor was presented prior to the masker and signal stimuli to activate the efferent system. The silent interval between the precursor and the masker-signal complex was 50 ms. For comparison, thresholds for detectability of the masked signal were also measured in a baseline condition without the precursor and, in addition, without the masker. On average, the results of the baseline condition indicate an effectively wider binaural filter, as expected. For both signal phases, the addition of the precursor results in effectively wider filters, which is in agreement with the hypothesis that cochlear gain is reduced due to the presence of the precursor.

Type 2 tasks in the theory of signal detectability: discrimination between correct and incorrect decisions.

It has been known for over 40 years that there are two fundamentally different kinds of detection tasks in the theory of signal detectability. The Type 1 task is to distinguish between events defined independently of the observer; the Type 2 task is to distinguish between one's own correct and incorrect decisions about those Type 1 events. For the Type 1 task, the behavior of the detector can be summarized by the traditional receiver operating characteristic (ROC) curve. This curve can be compared with a theoretical ROC curve, which can be generated from overlapping probability functions conditional on the Type 1 events on an appropriate decision axis. We show how to derive the probability functions underlying Type 2 decisions from those for the Type 1 task. ROC curves and the usual measures of performance are readily obtained from those Type 2 functions, and some relationships among various Type 1 and Type 2 performance measures are presented. We discuss the relationship between Type 1 and Type 2 confidence ratings and caution against the practice of presenting transformed Type 2 ratings as empirical Type 1 ratings.

Psychophysical evidence for auditory compression at low characteristic frequencies.

Psychophysical estimates of compression often assume that the basilar-membrane response to frequencies well below characteristic frequency (CF) is linear. Two techniques for estimating compression are described here that do not depend on this assumption at low CFs. In experiment 1, growth of forward masking was measured for both on- and off-frequency pure-tone maskers for pure-tone signals at 250, 500, and 4000 Hz. The on- and off-frequency masking functions at 250 and 500 Hz were just as shallow as the on-frequency masking function at 4000 Hz. In experiment 2, the forward masker level required to mask a fixed low-level signal was measured as a function of the masker-signal interval. The slopes of these functions did not differ between signal frequencies of 250 and 4000 Hz for the on-frequency maskers. At 250 Hz, the slope for the 150-Hz masker was almost as steep as that for the on-frequency masker, whereas at 4000 Hz the slope for the 2400-Hz masker was much shallower than that for the on-frequency masker. The results suggest that there is substantial compression, of around 0.2-0.3 dB/dB, at low CFs in the human auditory system. Furthermore, the results suggest that at low CFs compression does not vary greatly with stimulation frequency relative to CF.

Inferred basilar-membrane response functions for listeners with mild to moderate sensorineural hearing loss.

Psychophysical estimates of cochlear function suggest that normal-hearing listeners exhibit a compressive basilar-membrane (BM) response. Listeners with moderate to severe sensorineural hearing loss may exhibit a linearized BM response along with reduced gain, suggesting the loss of an active cochlear mechanism. This study investigated how the BM response changes with increasing hearing loss by comparing psychophysical measures of BM compression and gain for normal-hearing listeners with those for listeners who have mild to moderate sensorineural hearing loss. Data were collected from 16 normal-hearing listeners and 12 ears from 9 hearing-impaired listeners. The forward masker level required to mask a fixed low-level, 4000-Hz signal was measured as a function of the masker-signal interval using a masker frequency of either 2200 or 4000 Hz. These plots are known as temporal masking curves (TMCs). BM response functions derived from the TMCs showed a systematic reduction in gain with degree of hearing loss. Contrary to current thinking, however, no clear relationship was found between maximum compression and absolute threshold.