Fig. 6.1
Linear reference TMCs obtained with and without the CWN. Symbols and dotted lines: mean experimental data; error bars illustrate ± one standard deviation. Continuous lines: computer model simulations
Figure 6.1 also suggests that the CWN shifted the mean linear reference TMC downwards by ∼1.14 dB. This would be consistent with the expected effect of the CWN activating the MOCR; it reduces cochlear gain and thereby the response to the fixed-SPL probe. The observed shift, however, was neither statistically significant nor consistent across subjects. Indeed, it was smaller than the masker-level step size (2 dB) used in our adaptive procedure. This suggests that either the CWN did not activate the MOCR at 4 kHz or that the effect of MOCR activation on the fixed-SPL 4-kHz probe was so small that it became undetectable by our measuring method. Physiological studies demonstrate that a 60-dB SPL CWN is sufficient to evoke the MOCR (Lilaonitkul and Guinan 2009). Therefore, the latter explanation seems more likely.
The Effects of the CWN on the PTCs
The CWN had no significant effect on the PTCs at 4 kHz (Fig. 6.2, bottom). At 500 Hz, by contrast, the CWN led to lower masker levels at masked threshold in some conditions (Fig. 6.2, top). This effect occurred for all masker frequencies at the longest masker-probe time interval (50 ms). For the shorter intervals (2 and 10 ms), it only occurred for masker frequencies remote from the probe frequency, thus broadening the near-threshold PTCs.
Fig. 6.2
Mean behavioural PTCs (symbols) and model PTCs (lines) at 500 Hz (top) and 4 kHz (bottom), with and without the CWN. Each panel is for a different masker-probe time interval, as indicated in the bottom-left corner of the panel. Error bars are associated to the symbols and illustrate ± one standard deviation. Asterisks in the left-most panels depict the probe levels
The present results appear inconsistent with the study of Vinay and Moore (2008) on the effect of a CWN on near-threshold PTCs. They showed that the CWN had a different effect on the PTCs depending on the probe frequency. At 500 Hz, the CWN typically shifted the low-frequency side of the PTCs upwards and increased the tuning significantly. The difference with the present results may reflect methodological differences between studies. Indeed, Vinay and Moore (2008) measured PTCs using simultaneous rather than forward masking, and their probe and CWN levels differed from those used here.
At first sight, the pattern of the present results appears inconsistent with the effects of electrical activation of the MOCR on high-frequency basilar membrane (BM) responses, which shifts only the tip of the tuning curves upwards (Cooper and Guinan 2006). The computer model simulations described below will show that this inconsistency is more apparent than real.
3 Computer Model Simulations
A computer model of forward masking with efferent control was developed and used to test the assumption that the pattern of experimental data was consistent with the hypothesis that the CWN reduces cochlear gain by activating the MOCR. The model was inspired by the BM-temporal window model (Plack et al. 2002). In the current version of the model, the dual-resonance nonlinear (DRNL) filter (Meddis et al. 2001) was replaced by a version with efferent attenuation (Ferry and Meddis 2007). The latter simulates physiological observations at the level of the BM and auditory nerve by means of a single parameter that attenuates the input signal to the nonlinear path of the standard DRNL filter. Another novelty of the current model is that it accounts for the off-frequency listening effects at 500 Hz. This was deemed necessary because no precaution was taken experimentally to minimize off-frequency listening effects on PTCs that may have occurred due to the brief duration of the probe (10 ms).
The model was implemented in the time domain and evaluated for identical stimuli and conditions as used in the experiments. Its parameters were optimized automatically as follows. First, the parameters of the standard DRNL filter (without efferent attenuation) and the temporal window were optimized simultaneously to mimic the 4-kHz PTCs and the linear reference TMCs measured without the CWN. The resulting temporal window parameters were held constant for the other test frequencies and conditions. It was assumed that the rate of recovery from forward masking is constant across probe frequencies and that it is unaffected by CWN (see Fig. 6.1). Second, the parameters of the standard DRNL filter (without efferent attenuation) were optimized at 500 Hz using the experimental PTCs measured without the CWN. Lastly, the parameter controlling the amount of efferent attenuation was optimized to maximize the fit between simulated and experimental PTCs measured with CWN. The value of this parameter was allowed to vary across probe frequencies.
Importantly, rather than fitting the model to the mean data, a different set of model parameters was obtained for each individual data set, and the individual model responses were averaged and compared to the mean experimental PTCs.
3.1 Results and Discussion
The grey continuous line in Fig. 6.1 illustrates the mean linear reference TMC resulting from optimizing the model to simultaneously fit the experimental linear reference TMCs and the 4-kHz PTCs. Its slope was slightly steeper than that of the corresponding experimental curve (0.22 vs. 0.16 dB/ms). Although the difference in slope was not statistically significant, the goodness of fit to the experimental PTCs became poorer when the temporal window parameters were independently optimized to match the slope of the linear reference TMCs. This suggests that the actual recovery from forward masking could be slightly steeper than suggested by the measured linear reference TMC. This is not unreasonable considering that the experimental curve is based on measured values and it disregards runs that called for values higher than the maximum system output (105 dB SPL). The experimental linear reference TMC might have been steeper if the masker level had been allowed to go higher.