Heil Peter, Matysiak Artur, Neubauer Heinrich
Department of Systems Physiology of Learning, Leibniz Institute for Neurobiology, Magdeburg, Germany; Center for Behavioral Brain Sciences, Magdeburg, Germany.
Special Lab of Non-invasive Brain Imaging, Leibniz Institute for Neurobiology, Magdeburg, Germany.
Hear Res. 2017 Sep;353:135-161. doi: 10.1016/j.heares.2017.06.011. Epub 2017 Jun 27.
Thresholds for detecting sounds in quiet decrease with increasing sound duration in every species studied. The neural mechanisms underlying this trade-off, often referred to as temporal integration, are not fully understood. Here, we probe the human auditory system with a large set of tone stimuli differing in duration, shape of the temporal amplitude envelope, duration of silent gaps between bursts, and frequency. Duration was varied by varying the plateau duration of plateau-burst (PB) stimuli, the duration of the onsets and offsets of onset-offset (OO) stimuli, and the number of identical bursts of multiple-burst (MB) stimuli. Absolute thresholds for a large number of ears (>230) were measured using a 3-interval-3-alternative forced choice (3I-3AFC) procedure. Thresholds decreased with increasing sound duration in a manner that depended on the temporal envelope. Most commonly, thresholds for MB stimuli were highest followed by thresholds for OO and PB stimuli of corresponding durations. Differences in the thresholds for MB and OO stimuli and in the thresholds for MB and PB stimuli, however, varied widely across ears, were negative in some ears, and were tightly correlated. We show that the variation and correlation of MB-OO and MB-PB threshold differences are linked to threshold microstructure, which affects the relative detectability of the sidebands of the MB stimuli and affects estimates of the bandwidth of auditory filters. We also found that thresholds for MB stimuli increased with increasing duration of the silent gaps between bursts. We propose a new model and show that it accurately accounts for our results and does so considerably better than a leaky-integrator-of-intensity model and a probabilistic model proposed by others. Our model is based on the assumption that sensory events are generated by a Poisson point process with a low rate in the absence of stimulation and higher, time-varying rates in the presence of stimulation. A subject in a 3I-3AFC task is assumed to choose the interval in which the greatest number of events occurred or randomly chooses among intervals which are tied for the greatest number of events. The subject is further assumed to count events over the duration of an evaluation interval that has the same timing and duration as the expected stimulus. The increase in the rate of the events caused by stimulation is proportional to the time-varying amplitude envelope of the bandpass-filtered signal raised to an exponent. We find the exponent to be about 3, consistent with our previous studies. This challenges models that are based on the assumption of the integration of a neural response that is directly proportional to the stimulus amplitude or proportional to its square (i.e., proportional to the stimulus intensity or power).
在每个被研究的物种中,安静环境下检测声音的阈值会随着声音持续时间的增加而降低。这种权衡背后的神经机制,通常被称为时间整合,目前尚未完全被理解。在这里,我们用大量时长、时间幅度包络形状、脉冲串之间静音间隙的时长以及频率各不相同的纯音刺激来探究人类听觉系统。通过改变平台 - 脉冲串(PB)刺激的平台时长、起始 - 偏移(OO)刺激的起始和偏移时长以及多脉冲串(MB)刺激中相同脉冲串的数量来改变时长。使用三区间三择一强制选择(3I - 3AFC)程序测量了大量耳朵(>230只)的绝对阈值。阈值随着声音持续时间的增加而降低,其方式取决于时间包络。最常见的情况是,MB刺激的阈值最高,其次是相应时长的OO和PB刺激的阈值。然而,MB和OO刺激的阈值差异以及MB和PB刺激的阈值差异在不同耳朵之间变化很大,在某些耳朵中为负,并且紧密相关。我们表明,MB - OO和MB - PB阈值差异的变化和相关性与阈值微观结构有关,阈值微观结构会影响MB刺激边带的相对可检测性,并影响听觉滤波器带宽的估计。我们还发现,MB刺激的阈值随着脉冲串之间静音间隙时长的增加而升高。我们提出了一个新模型,并表明它能准确解释我们的结果,而且比其他人提出的强度泄漏积分模型和概率模型表现得好得多。我们的模型基于这样的假设:在没有刺激时,感觉事件由泊松点过程以低速率产生;在有刺激时,由更高的、随时间变化的速率产生。假设一个处于3I - 3AFC任务中的受试者会选择事件发生数量最多的区间,或者在事件数量相同的区间中随机选择。进一步假设受试者在与预期刺激具有相同时间和时长的评估区间内对事件进行计数。由刺激引起的事件速率增加与带通滤波信号的随时间变化的幅度包络的指数成正比。我们发现该指数约为3,与我们之前的研究一致。这对基于神经反应积分假设的模型提出了挑战,这些假设认为神经反应与刺激幅度成正比或与刺激幅度的平方成正比(即与刺激强度或功率成正比)。