Heil P
Department of Psychology, Monash University, Clayton, Victoria, Australia.
J Neurophysiol. 1997 May;77(5):2616-41. doi: 10.1152/jn.1997.77.5.2616.
Sound onsets are salient and behaviorally relevant, and most auditory neurons discharge spikes locked to such transients. The acoustic parameters of sound onsets that shape such onset responses are unknown. In this paper is analyzed the timing of spikes of single neurons in the primary auditory cortex of barbiturate-anesthetized cats to the onsets of tone bursts. By parametric variation of sound pressure level, rise time, and rise function (linear or cosine-squared), the time courses of peak pressure, rate of change of peak pressure, and acceleration of peak pressure during the tones' onsets were systematically varied. For cosine-squared rise function tones of a given frequency and laterality, any neuron's mean first-spike latency was an invariant and inverse function of the maximum acceleration of peak pressure occurring at tone onset. For linear rise function tones, latency was an invariant and inverse function of the rate of change of peak pressure. Thus latency is independent of rise time or sound pressure level per se. Latency-acceleration functions, obtained with cosine-squared rise function tones under different stimulus conditions (frequency, laterality) from any given neuron and across the neuronal pool, were of strikingly similar shape. The same was true for latency-rate of change of peak pressure functions obtained with linear rise function tones. Latency-acceleration/rate of change of peak pressure functions could differ in their extent and in their position within the coordinate system. The positional differences reflect neuronal differences in minimum latency Lmin and in a sensitivity S to acceleration and rate of change of peak pressure (transient sensitivity), a hitherto unrecognized neuronal property that is distinctly different from firing threshold. Estimates of Lmin and S, which were derived by fitting a simple function to the neuronal latency-acceleration/rate of change of peak pressure functions, were independent of rise function. On average, Lmin decreased with increasing characteristic frequency (CF), but varied widely for neurons with the same CF. S varied with CF in a fashion similar to the cat's audiogram and, for a given neuron, varied with frequency. SD of first-spike latency was roughly proportional to the slope of the functions relating latency to acceleration/rate of change of peak pressure. Thus SD increased exponentially, rather than linearly, with mean latency, and did so at about twice the rate for linear than for cosine-squared rise function tones. The proportionality coefficients were quite similar across the neuronal pool and similar for both rise functions. Minimum SD increased nonlinearly with increasing Lmin. These findings suggest a peripheral origin of S and a peripheral establishment of latency-acceleration/rate of change of peak pressure functions. Because of the striking similarity in the shapes of such functions across the neuronal pool, sound onsets will produce orderly and predictable spatiotemporal patterns of first-spike timing, which could be used to instantaneously track rapid transients and to represent transient features by partly scale-invariant temporal codes.
声音起始段很显著且与行为相关,大多数听觉神经元会发放与这些瞬态相关的脉冲。塑造此类起始反应的声音起始段的声学参数尚不清楚。本文分析了巴比妥麻醉猫的初级听觉皮层中单个神经元对短纯音起始段的脉冲发放时间。通过系统地改变声压级、上升时间和上升函数(线性或余弦平方)的参数,短纯音起始段期间的峰值压力时间进程、峰值压力变化率和峰值压力加速度都发生了变化。对于给定频率和方位的余弦平方上升函数短纯音,任何神经元的平均首次脉冲潜伏期是起始段峰值压力最大加速度的不变且反比函数。对于线性上升函数短纯音,潜伏期是峰值压力变化率的不变且反比函数。因此,潜伏期本身与上升时间或声压级无关。在不同刺激条件(频率、方位)下,从任何给定神经元以及整个神经元群中,用余弦平方上升函数短纯音获得的潜伏期 - 加速度函数形状惊人地相似。用线性上升函数短纯音获得的潜伏期 - 峰值压力变化率函数也是如此。潜伏期 - 加速度/峰值压力变化率函数在其范围和坐标系中的位置可能不同。位置差异反映了神经元在最小潜伏期Lmin以及对峰值压力加速度和变化率的敏感性S(瞬态敏感性)方面的差异,这是一种迄今未被认识到的神经元特性,与发放阈值明显不同。通过将一个简单函数拟合到神经元潜伏期 - 加速度/峰值压力变化率函数来推导Lmin和S的估计值,该估计值与上升函数无关。平均而言,Lmin随特征频率(CF)增加而降低,但相同CF的神经元变化范围很大。S随CF的变化方式类似于猫的听力图,并且对于给定神经元,随频率变化。首次脉冲潜伏期的标准差大致与将潜伏期与峰值压力加速度/变化率相关联的函数斜率成正比。因此,标准差随平均潜伏期呈指数增加,而不是线性增加,并且对于线性上升函数短纯音,其增加速率约为余弦平方上升函数短纯音的两倍。比例系数在整个神经元群中非常相似,并且两种上升函数也相似。最小标准差随Lmin增加而非线性增加。这些发现表明S起源于外周,并且潜伏期 - 加速度/峰值压力变化率函数在外周建立。由于整个神经元群中此类函数形状的惊人相似性,声音起始段将产生有序且可预测的首次脉冲发放时间的时空模式,这可用于即时跟踪快速瞬态,并通过部分尺度不变的时间编码来表示瞬态特征。