Clarke R J, Ikeda H
Exp Brain Res. 1985;59(1):83-90. doi: 10.1007/BF00237669.
The luminance detectors in the olivary pretectal nucleus, which are likely candidates mediating the pupillary light reflex, responded to all frequencies of sinusoidally modulated light up to 12-25 Hz. At low frequencies (0.05-4.0 Hz) the luminance detectors responded with modulated firing to different stimulation rates. The modulation depth of the cell response increased with the increase in stimulation frequency up to 20 Hz, then rapidly fell. There was a delay between the peak intensity of the stimulus and the peak firing of cell response of about 30-40 ms. The amplitudes of the consensual pupil responses to the same sinusoidal stimulus, on the other hand, decreased with an increase in frequency and no discernible response was recorded above 2 Hz. The pupil responses were little affected by sympathectomy. The differences in the frequency response characteristics of luminance detectors and the pupil were attributed to the sluggish dynamic properties of the pupil muscles. This was demonstrated using an electronic model of the iris muscle which modified the responses of the luminance detectors giving output waveforms broadly resembling pupil responses to square and sinusoidally modulated lights.
橄榄顶盖前核中的亮度探测器可能是介导瞳孔光反射的候选者，它对高达12 - 25Hz的所有正弦调制光频率都有反应。在低频（0.05 - 4.0Hz）时，亮度探测器对不同的刺激速率以调制放电做出反应。细胞反应的调制深度随着刺激频率增加到20Hz而增加，然后迅速下降。刺激的峰值强度与细胞反应的峰值放电之间存在约30 - 40毫秒的延迟。另一方面，对相同正弦刺激的交感瞳孔反应幅度随频率增加而减小，在2Hz以上未记录到可辨别的反应。瞳孔反应受交感神经切除术的影响很小。亮度探测器和瞳孔的频率响应特性差异归因于瞳孔肌肉的缓慢动态特性。这通过虹膜肌肉的电子模型得到了证明，该模型改变了亮度探测器的反应，给出的输出波形大致类似于瞳孔对方形和正弦调制光的反应。
