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

Auditory Nerve Frequency Tuning Measured with Forward-Masked Compound Action Potentials

Frequency selectivity is a fundamental cochlear property. Recent studies using otoacoustic emissions and psychophysical forward masking suggest that frequency selectivity is sharper in human than in common laboratory species. This has been disputed based on reports using compound action potentials (CAPs), which reflect activity in the auditory nerve and can be measured in humans. Comparative data of CAPs, obtained with a variety of simultaneous masking protocols, have been interpreted to indicate similarity of frequency tuning across mammals and even birds. Unfortunately, there are several issues with the available CAP measurements which hamper a straightforward comparison across species. We investigate sharpness of CAP tuning in cat and chinchilla using a forward masking notched-noise paradigm—which is less confounded by cochlear nonlinearities than simultaneous masking paradigms and similar to what was used in the psychophysical study reporting sharper tuning in humans. Our parametric study, using different probe frequencies and notch widths, shows relationships consistent with those of auditory nerve fibers (ANFs). The sharpness of tuning, quantified by Q(10) factors, is negatively correlated with probe level and increases with probe frequency, but the Q(10) values are generally lower than the average trend for ANFs. Like the single fiber data, tuning for CAPs is sharper in cat than in chinchilla, but the two species are similar in the dependence of tuning on probe frequency and in the relationship between tuning in ANFs and CAP. Growth-of-maskability functions show slopes <1 indicating that with increasing probe level the probe is more susceptible to cochlear compression than the masker. The results support the use of forward-masked CAPs as an alternative measure to estimate ANF tuning and to compare frequency tuning across species

Central auditory masking by an illusory tone

Many natural sounds fluctuate over time. The detectability of sounds in a sequence can be reduced by prior stimulation in a process known as forward masking. Forward masking is thought to reflect neural adaptation or neural persistence in the auditory nervous system, but it has been unclear where in the auditory pathway this processing occurs. To address this issue, the present study used a "Huggins pitch" stimulus, the perceptual effects of which depend on central auditory processing. Huggins pitch is an illusory tonal sensation produced when the same noise is presented to the two ears except for a narrow frequency band that is different (decorrelated) between the ears. The pitch sensation depends on the combination of the inputs to the two ears, a process that first occurs at the level of the superior olivary complex in the brainstem. Here it is shown that a Huggins pitch stimulus produces more forward masking in the frequency region of the decorrelation than a noise stimulus identical to the Huggins-pitch stimulus except with perfect correlation between the ears. This stimulus has a peripheral neural representation that is identical to that of the Huggins-pitch stimulus. The results show that processing in, or central to, the superior olivary complex can contribute to forward masking in human listeners

Threshold and Beyond: Modeling The Intensity Dependence of Auditory Responses

In many studies of auditory-evoked responses to low-intensity sounds, the response amplitude appears to increase roughly linearly with the sound level in decibels (dB), corresponding to a logarithmic intensity dependence. But the auditory system is assumed to be linear in the low-intensity limit. The goal of this study was to resolve the seeming contradiction. Based on assumptions about the rate-intensity functions of single auditory-nerve fibers and the pattern of cochlear excitation caused by a tone, a model for the gross response of the population of auditory nerve fibers was developed. In accordance with signal detection theory, the model denies the existence of a threshold. This implies that regarding the detection of a significant stimulus-related effect, a reduction in sound intensity can always be compensated for by increasing the measurement time, at least in theory. The model suggests that the gross response is proportional to intensity when the latter is low (range I), and a linear function of sound level at higher intensities (range III). For intensities in between, it is concluded that noisy experimental data may provide seemingly irrefutable evidence of a linear dependence on sound pressure (range II). In view of the small response amplitudes that are to be expected for intensity range I, direct observation of the predicted proportionality with intensity will generally be a challenging task for an experimenter. Although the model was developed for the auditory nerve, the basic conclusions are probably valid for higher levels of the auditory system, too, and might help to improve models for loudness at threshold

Chronic Reduction of Endocochlear Potential Reduces Auditory Nerve Activity: Further Confirmation of an Animal Model of Metabolic Presbyacusis

Gerbils aged in quiet show a decline of the endocochlear potential (EP) and elevated auditory nerve compound action potential (CAP) thresholds. However, establishing a direct relationship between an age-related reduction in the EP and changes in the activities of primary auditory neurons is difficult owing to the complexity of age-related histological changes in the cochlea. To address this issue, we developed a young gerbil model of “metabolic” presbyacusis that uses an osmotic pump to deliver furosemide into the round window niche for 7 days, resulting in a chronically reduced EP. In this model, the only major histopathologic changes were restricted to the hook region of the cochlea and consisted of loss of strial intermediate cells and massive edema in the lateral wall. The morphological and physiological evidence suggests that the cochlea can adapt to furosemide application over time. The morphology of spiral ganglion cells and hair cells appeared normal throughout the cochlea. CAP responses and EP values in this model are similar to those of quiet-aged ears. The spontaneous activity of single auditory fibers (n = 188) was assessed in 15 young gerbils treated with furosemide for 7 days. The percentage of recorded low-spontaneous rate (SR) fibers at characteristic frequencies (CFs) ≥ 6 kHz was significantly lower in furosemide-treated than in control ears. Recovery function tests of CAP responses after prior stimulation also showed a decline in activity of the low-SR population with CFs ≥ 6 kHz in the treated cochleas. A similar loss in the activity of low-SR fiber has been previously shown in quiet-aged gerbils. These results suggest that dysfunction of the cochlear lateral wall and subsequent chronic reduction in the EP can directly affect the activity patterns of primary auditory neurons in a manner similar to that seen in aged gerbils