Ana Alves-Pinto

A variant temporal-masking-curve method for inferring peripheral auditory compression

Recent studies have suggested that the degree of on-frequency peripheral auditory compression is similar for apical and basal cochlear sites and that compression extends to a wider range of frequencies in apical than in basal sites. These conclusions were drawn from the analysis of the slopes of temporal masking curves (TMCs) on the assumption that forward masking decays at the same rate for all probe and masker frequencies. The aim here was to verify this conclusion using a different assumption. TMCs for normal hearing listeners were measured for probe frequencies (f(P)) of 500 and 4000 Hz and for masker frequencies (f(M)) of 0.4, 0.55, and 1.0 times the probe frequency. TMCs were measured for probes of 9 and 15 dB sensation level. The assumption was that given a 6 dB increase in probe level, linear cochlear responses to the maskers should lead to a 6 dB vertical shift of the corresponding TMCs, while compressive responses should lead to bigger shifts. Results were consistent with the conclusions from earlier studies. It is argued that this supports the assumptions of the standard TMC method for inferring compression, at least in normal-hearing listeners.

Forward Masking Estimated by Signal Detection Theory Analysis of Neuronal Responses in Primary Auditory Cortex

Psychophysical forward masking is an increase in threshold of detection of a sound (probe) when it is preceded by another sound (masker). This is reminiscent of the reduction in neuronal responses to a sound following prior stimulation. Studies in the auditory nerve and cochlear nucleus using signal detection theory techniques to derive neuronal thresholds showed that in centrally projecting neurons, increases in masked thresholds were significantly smaller than the changes measured psychophysically. Larger threshold shifts have been reported in the inferior colliculus of awake marmoset. The present study investigated the magnitude of forward masking in primary auditory cortical neurons of anaesthetised guinea-pigs. Responses of cortical neurons to unmasked and forward masked tones were measured and probe detection thresholds estimated using signal detection theory methods. Threshold shifts were larger than in the auditory nerve, cochlear nucleus and inferior colliculus. The larger threshold shifts suggest that central, and probably cortical, processes contribute to forward masking. However, although methodological differences make comparisons difficult, the threshold shifts in cortical neurons were, in contrast to subcortical nuclei, actually larger than those observed psychophysically. Masking was largely attributable to a reduction in the responses to the probe, rather than either a persistence of the masker responses or an increase in the variability of probe responses.

Psychophysical estimates of level-dependent best-frequency shifts in the apical region of the human basilar membrane.

It is now undisputed that the best frequency (BF) of basal basilar-membrane (BM) sites shifts downwards as the stimulus level increases. The direction of the shift for apical sites is, by contrast, less well established. Auditory nerve studies suggest that the BF shifts in opposite directions for apical and basal BM sites with increasing stimulus level. This study attempts to determine if this is the case in humans. Psychophysical tuning curves (PTCs) were measured using forward masking for probe frequencies of 125, 250, 500, and 6000 Hz. The level of a masker tone required to just mask a fixed low-level probe tone was measured for different masker-probe time intervals. The duration of the intervals was adjusted as necessary to obtain PTCs for the widest possible range of masker levels. The BF was identified from function fits to the measured PTCs and it almost always decreased with increasing level. This result is inconsistent with most auditory-nerve observations obtained from other mammals. Several explanations are discussed, including that it may be erroneous to assume that low-frequency PTCs reflect the tuning of apical BM sites exclusively and that the inherent frequency response of the inner hair cell may account for the discrepancy.

Detection of high‐frequency spectral notches as a function of level

High-frequency spectral notches are important cues for sound localization. Our ability to detect them must depend on their representation as auditory nerve (AN) rate profiles. Because of the low threshold and the narrow dynamic range of most AN fibers, these rate profiles deteriorate at high levels. The system may compensate by using onset rate profiles whose dynamic range is wider, or by using low-spontaneous-rate fibers, whose threshold is higher. To test these hypotheses, the threshold notch depth necessary to discriminate between a flat spectrum broadband noise and a similar noise with a spectral notch centered at 8 kHz was measured at levels from 32 to 100 dB SPL. The importance of the onset rate-profile representation of the notch was estimated by varying the stimulus duration and its rise time. For a large proportion of listeners, threshold notch depth varied nonmonotonically with level, increasing for levels up to 70-80 dB SPL and decreasing thereafter. The nonmonotonic aspect of the function was independent of notch bandwidth and stimulus duration. Thresholds were independent of stimulus rise time but increased for the shorter noise bursts. Results are discussed in terms of the ability of the AN to convey spectral notch information at different levels.

SIGNAL DETECTION IN ANIMAL PSYCHOACOUSTICS: ANALYSIS AND SIMULATION OF SENSORY AND DECISION-RELATED INFLUENCES

Highlights ► Signal detection theory was applied to ferrets’ behaviour-detecting tones in noise. ► Sensory acuity was stable but the decisions depended on the set of stimuli. ► Decisions also depended on responses and rewards in previous trials. ► Computer simulations accounted for the data quantitatively. ► Behaviour reflects a stable sensory representation with a dynamic decision process.

Relating approach-to-target and detection tasks in animal psychoacoustics.

Psychophysical experiments seek to measure the limits of perception. While straightforward in humans, in animals they are time consuming. Choosing an appropriate task and interpreting measurements can be challenging. We investigated the localization of high-frequency auditory signals in noise using an “approach-to-target” task in ferrets, how task performance should be interpreted in terms of perception, and how the measurements relate to other types of tasks. To establish their general ability to localize, animals were first trained to discriminate broadband noise from 12 locations. Subsequently we tested their ability to discriminate between band-limited targets at 2 or 3 more widely spaced locations, in a continuous background noise. The ability to discriminate between 3 possible locations (−90°, 0°, 90°) of a 10-kHz pure tone decreased gradually over a wide range (>30 dB) of signal-to-noise ratios (SNRs). Location discrimination ability was better for wide band noise targets (0.5 and 2 octave). These results were consistent with localization ability limiting performance for pure tones. Discrimination of pure tones at 2 locations (−90/left, 90/right) was robust at positive SNRs, yielding psychometric functions which fell steeply at negative SNRs. Thresholds for discrimination were similar to previous tone-in-noise thresholds measured in ferrets using a yes/no task. Thus, using an approach-to-target task, sound “localization” in noise can reflect detectability or the ability to localize, depending on the stimulus configuration. Signal-detection-theory-based models were able to account for the results when discriminating between pure tones from 2- and 3-source locations.

Behavioural estimates of auditory filter widths in ferrets using notched-noise maskers

Frequency selectivity is a fundamental property of hearing which affects almost all aspects of auditory processing. Here auditory filter widths at 1, 3, 7, and 10 kHz were estimated from behavioural thresholds using the notched-noise method [Patterson, Nimmo-Smith, Weber, and Milroy, J. Acoust. Soc. Am. 72, 1788-1803 (1982)] in ferrets. The mean bandwidth was 21% of the signal frequency, excluding wider bandwidths at 1 kHz (65%). They were comparable although on average broader than equivalent measurements in other mammals (∼11%-20%), and wider than bandwidths measured from the auditory nerve in ferrets (∼18%). In non-human mammals there is considerable variation between individuals, species, and in the correspondence with auditory nerve tuning.

Decision Criterion Dynamics in Animals Performing an Auditory Detection Task

Classical signal detection theory attributes bias in perceptual decisions to a threshold criterion, against which sensory excitation is compared. The optimal criterion setting depends on the signal level, which may vary over time, and about which the subject is naïve. Consequently, the subject must optimise its threshold by responding appropriately to feedback. Here a series of experiments was conducted, and a computational model applied, to determine how the decision bias of the ferret in an auditory signal detection task tracks changes in the stimulus level. The time scales of criterion dynamics were investigated by means of a yes-no signal-in-noise detection task, in which trials were grouped into blocks that alternately contained easy- and hard-to-detect signals. The responses of the ferrets implied both long- and short-term criterion dynamics. The animals exhibited a bias in favour of responding “yes” during blocks of harder trials, and vice versa. Moreover, the outcome of each single trial had a strong influence on the decision at the next trial. We demonstrate that the single-trial and block-level changes in bias are a manifestation of the same criterion update policy by fitting a model, in which the criterion is shifted by fixed amounts according to the outcome of the previous trial and decays strongly towards a resting value. The apparent block-level stabilisation of bias arises as the probabilities of outcomes and shifts on single trials mutually interact to establish equilibrium. To gain an intuition into how stable criterion distributions arise from specific parameter sets we develop a Markov model which accounts for the dynamic effects of criterion shifts. Our approach provides a framework for investigating the dynamics of decisions at different timescales in other species (e.g., humans) and in other psychological domains (e.g., vision, memory).

Neuronal Measures of Threshold and Magnitude of Forward Masking in Primary Auditory Cortex

Psychophysical forward masking is an increase in the threshold of detection of a brief sound (probe) when preceded by another sound (masker). These effects are reminiscent of the reduction in physiological responses following prior stimulation. However, previous studies of the response of auditory nerve fibers (Relkin and Turner, 1988) found probe threshold shifts following stimulation that were considerably smaller than those found perceptually. Although such threshold shifts are larger in some units of the cochlear nucleus (Ingham et al., 2006), these are either inhibitory interneurons or project to inhibitory neurons. A better account is obtained at the level of the IC in the awake marmoset (Nelson et al., 2009).

Rate versus time representation of high‐frequency spectral notches in the peripheral auditory system: A computational model study

We investigated how high‐frequency spectral notches are represented in the auditory nerve. Our approach consisted in modeling the paradoxical result of Alves‐Pinto and Lopez‐Poveda [J. Acoust. Soc. Am. 118, 2548 (2005)] that discriminating between broadband noises with and without high‐frequency spectral notches is most difficult at 70–80 dB SPL than at lower or higher intensities. We tested two possibilities: (a) that discrimination is based on the difference between the inner‐hair‐cell excitation patterns for the two stimuli, a representation related to the auditory nerve difference rate profile, and (b) that it is based on the difference inner‐hair‐cell modulation‐rate pattern, a representation related to the difference in the temporal pattern of auditory nerve discharges. The simulations support the latter. This suggests that spectral features as high in frequency as 8 kHz may be encoded in the temporal pattern of auditory nerve responses despite the rapid decay of phase locking above 2 kHz. The simul...