Werblin Frank S
Department of Molecular and Cell Biology, University of California at Berkeley, Berkeley, California 94720, USA.
Vis Neurosci. 2010 Mar;27(1-2):1-8. doi: 10.1017/S0952523810000076. Epub 2010 Apr 15.
Early retinal studies categorized ganglion cell behavior as either linear or nonlinear and rectifying as represented by the familiar X- and Y-type ganglion cells in cat. Nonlinear behavior is in large part a consequence of the rectifying nonlinearities inherent in synaptic transmission. These nonlinear signals underlie many special functions in retinal processing, including motion detection, motion in motion, and local edge detection. But linear behavior is also required for some visual processing tasks. For these tasks, the inherently nonlinear signals are "linearized" by "crossover inhibition." Linearization utilizes a circuitry whereby nonlinear ON inhibition adds with nonlinear OFF excitation or ON excitation adds with OFF inhibition to generate a more linear postsynaptic voltage response. Crossover inhibition has now been measured in most bipolar, amacrine, and ganglion cells. Functionally crossover inhibition enhances edge detection, allows ganglion cells to recognize luminance-neutral patterns with their receptive fields, permits ganglion cells to distinguish contrast from luminance, and maintains a more constant conductance during the light response. In some cases, crossover extends the operating range of cone-driven OFF ganglion cells into the scotopic levels. Crossover inhibition is also found in neurons of the lateral geniculate nucleus and V1.
早期的视网膜研究将神经节细胞的行为分为线性或非线性以及整流型,就像猫中常见的X型和Y型神经节细胞所表现的那样。非线性行为在很大程度上是突触传递中固有的整流非线性的结果。这些非线性信号是视网膜处理中许多特殊功能的基础,包括运动检测、动态中的运动以及局部边缘检测。但某些视觉处理任务也需要线性行为。对于这些任务,固有的非线性信号通过“交叉抑制”被“线性化”。线性化利用一种电路,即非线性的ON抑制与非线性的OFF兴奋相加,或者ON兴奋与OFF抑制相加,以产生更线性的突触后电压响应。现在已经在大多数双极细胞、无长突细胞和神经节细胞中测量到了交叉抑制。在功能上,交叉抑制增强了边缘检测,使神经节细胞能够用其感受野识别亮度中性模式,使神经节细胞能够区分对比度和亮度,并在光反应期间保持更恒定的电导。在某些情况下,交叉抑制将锥体驱动的OFF神经节细胞的工作范围扩展到暗视水平。在外侧膝状体核和V1的神经元中也发现了交叉抑制。