Huffman R F, Argeles P C, Covey E
Department of Neurobiology, Duke University Medical Center, Durham, NC 27710, USA.
Hear Res. 1998 Dec;126(1-2):181-200. doi: 10.1016/s0378-5955(98)00166-x.
Changes in amplitude are a characteristic feature of most natural sounds, including the biosonar signals used by bats for echolocation. Previous evidence suggests that the nuclei of the lateral lemniscus play an important role in processing timing information that is essential for target range determination in echolocation. Neurons that respond to unmodulated tones with a sustained discharge are found in the dorsal nucleus (DNLL), intermediate nucleus (INLL) and multipolar cell division of the ventral nucleus (VNLLm). These neurons provide a graded response over a broad dynamic range of intensities, and would be expected to provide information about the amplitude envelope of a modulated signal. Neurons that respond only at the onset of a tone make up a small proportion of cells in DNLL, INLL and VNLLm, but are the only type found in the columnar division of the ventral nucleus (VNLLc). Onset neurons in VNLLc maintain a constant latency across a wide range of stimulus frequencies and intensities, thus providing a precise marker for when a sound begins. To determine how these different functional classes of cells respond to amplitude changes, we presented sinusoidally amplitude modulated (SAM) signals monaurally to awake, restrained bats and recorded the responses of single neurons extracellularly. There were clear differences in the ability of neurons in the different cell groups to respond to SAM. In the VNLLm, INLL and DNLL, 90% of neurons responded to SAM with a synchronous discharge. Neurons in the VNLLc responded poorly or not at all to SAM signals. This finding was unexpected given the precise onset responses of VNLLc neurons to unmodulated tones and their ability to respond synchronously to sinusoidally frequency modulated (SFM) signals. Among neurons that responded synchronously to SAM, synchronization as a function of modulation rate described either a bandpass or a lowpass function, with the majority of bandpass functions in neurons that responded to unmodulated tones with a sustained discharge. The maximal modulation rates that elicited synchronous responses were similar for the different cell groups, ranging from 320 Hz in VNLLm to 230 Hz in DNLL. The range of best modulation rates was greater for SAM than for SFM; this was also true of the range of maximal modulation rates at which synchronous discharge occurred. There was little correlation between a neuron's best modulation rate or maximal modulation rate for SAM signals and those for SFM signals, suggesting that responsiveness to amplitude and frequency modulations depends on different neural processing mechanisms.
振幅变化是大多数自然声音的一个特征,包括蝙蝠用于回声定位的生物声纳信号。先前的证据表明,外侧丘系核在处理时间信息方面起着重要作用,而时间信息对于回声定位中目标距离的确定至关重要。在背侧核(DNLL)、中间核(INLL)和腹侧核的多极细胞区(VNLLm)中发现了对未调制音调以持续放电做出反应的神经元。这些神经元在很宽的强度动态范围内提供分级反应,预计它们会提供有关调制信号振幅包络的信息。仅在音调开始时做出反应的神经元在DNLL、INLL和VNLLm中占细胞的比例较小,但却是腹侧核柱状区(VNLLc)中唯一发现的类型。VNLLc中的起始神经元在很宽的刺激频率和强度范围内保持恒定的潜伏期,从而为声音开始的时间提供了一个精确的标记。为了确定这些不同功能类别的细胞如何对振幅变化做出反应,我们将正弦振幅调制(SAM)信号单耳呈现给清醒、受限的蝙蝠,并在细胞外记录单个神经元的反应。不同细胞组中的神经元对SAM做出反应的能力存在明显差异。在VNLLm、INLL和DNLL中,90%的神经元对SAM以同步放电做出反应。VNLLc中的神经元对SAM信号反应不佳或根本没有反应。考虑到VNLLc神经元对未调制音调的精确起始反应以及它们对正弦频率调制(SFM)信号同步反应的能力,这一发现出乎意料。在对SAM同步做出反应的神经元中,作为调制率函数的同步性描述了一个带通或低通函数,大多数带通函数存在于对未调制音调以持续放电做出反应的神经元中。不同细胞组引发同步反应的最大调制率相似,范围从VNLLm中的320 Hz到DNLL中的230 Hz。SAM的最佳调制率范围比SFM的更大;同步放电发生的最大调制率范围也是如此。神经元对SAM信号的最佳调制率或最大调制率与对SFM信号的最佳调制率或最大调制率之间几乎没有相关性,这表明对振幅和频率调制的反应性取决于不同的神经处理机制。