视网膜神经节细胞感受野的一种新模型 II.神经节细胞方位选择性中心周边相互作用机制

在前文建立的二维视网膜神经节细胞含大周边感受野模型基础上 ,结合生理实验结果模拟了神经节细胞的方位选择性特性。文中采用椭圆感受野的观点解释了方位选择性的成因。并通过中心区以外区域对中心区方位选择性的复杂调制组合 ,展示了感受野不同亚单元对方位选择性的影响作用 ;指出方位选择性的成因是感受野椭圆亚单元的存在;感受野复杂的方位选择性是由于中心和周边在不同刺激条件下竞争的不同结果造成的;同时指出对椭圆感受野 ,倍频反应也会有相应的方位选择性。     

视网膜神经节细胞感受野的一种新模型：Ⅱ.神经节细胞方位选 …

在前文建立的二维视网膜神经节细胞含大周迷感受野模型基础上,结合生理实验模拟了神经节细胞的方位选择性特性。文中采用椭圆感受野的观点解释了方位选择性的成因。并通过中心区以外区域对中心区方位选择怀的复杂调制组合,展示了感受野不同亚地方位选择性的影响作用;指出方位选择性的成因是感受野椭圆亚单元的存在,感受野复杂的方位选择性是由于中心和周边在不同刺激条件下竞争的不同结果造成的;同时指出对椭圆感受野,倍频反应     

抑制性整合野细胞的对比度响应函数研究

采用细胞外记录的方法,在单独刺激经典感受野(classical receptive field,CRF)或同时刺激CRF和感受野外区域(extra-receptive field,ERF)的情况下,测量了猫初级视觉皮层细胞的对比度响应函数。当刺激所用的中心和外周运动光栅的参数一致时,与仅刺激CRF相比,强的ERF抑制使对比度响应函数动态区增加,响应增益和对比度增益降低。当中心和外周光栅的方位相差90度时,与方位参数一致的情况相比,大部分细胞的ERF抑制减弱,对比度响应函数的动态区减小,对比度增益和响应增益增加;少数细胞的ERF对CRF的作用从抑制变为易化,其对比度响应函数的动态区与只刺激CRF相比还要小,而对比度增益和响应增益还要大。揭示了初级视觉皮层细胞的抑制型整合野在CRF和ERF图像的方位及对比度差异检测中的作用机制。     

简单细胞方位选择性感受野组织形成的神经网络模型

为了阐明视皮层简单细胞方位选择性感受野形成的动态组织过程, 试图构建一个由侧膝体神经元和视皮层简单细胞组成的, 且遵从Hebbian学习规则的神经网络模型. 通过该模型来考察简单细胞对自然图像刺激特征的编码过程和神经表达. 结果表明, 感受野的结构正反映了简单细胞的最优方位选择性, 它也是由非监督学习过程决定并自组织涌现的. 这还说明简单细胞的方位选择性是在层间细胞的相互作用基础上动态自组织的结果.     

猫纹状皮层神经元整合野结构的对称性及空间总合特性

用正弦调制的移动光栅刺激传统感受野和整合野,测量了猫纹状皮层神经元整合野各亚区的范围、抑制程度和亚区间的空间总合特性。结果表明：⑴整合野的两个侧区（以及两个端区）之间具有相同的作用性质（抑制或易化）。⑵大多数整合野的两个侧区（以及两个端区）的范围相等或大致相等。⑶抑制型整合野的两个侧区（以及两个端区）间对细胞反应的抑制程度显著相关。⑷两个侧区（以及两个端区）之间的空间总合具有非线性特性,侧区间总合与端区间总合的非线性程度相同。以上结果提示,整合野的两个共轭对称的亚区可能是由同一个与感受野同心重叠的大区域构成。     

猫外膝体神经元时间频率调谐特性的研究

用示波器产生亮度受正弦波调制的小光点刺激清醒猫外膝体神经元的感受野,以不同调制频率下神经元放电的平均频率为指标,分析了126个细胞感受野中心的时间频率调谐特性,主要结果如下。(1)大多数(93.7%)细胞呈调制-兴奋型反应,即刺激光的时间调制在一定频率范围内使放电频率增加;少数(6.3%)细胞呈调制-抑制型反应,即在一定频率范围内,时间调制使放电减少。(2)根据调谐曲线的形状和通带宽度,调制-兴奋型反应包括带通滤波器和低通滤波器两种类型,其110个调谐曲线的峰值分布接近正态曲线,多数细胞对7Hz 的调谐最敏感。调制一抑制型反应包括带除滤波器和低除滤波器两类。(3)调制-兴奋型曲线的通带旁边较當出现抑制性侧带,调制-抑制型曲线出现兴奋性侧带。(4)感受野中心区与外周区的时间频率调谐曲线的带宽和形状有所不同。     

猫视皮层17,18区神经元对错觉轮廓的反应

研究了轻度麻醉下猫视皮层17, 18区细胞对错觉轮廓刺激的反应特性, 比较了对错觉轮廓有明显反应的细胞对真实轮廓和错觉轮廓刺激的感受野特性的异同. 共记录了猫视皮层17, 18区200个方位/方向选择性细胞, 其中有42%的细胞是错觉轮廓反应细胞. 将这些细胞对真实轮廓和错觉轮廓的反应进行比较, 尽管错觉轮廓反应细胞对移动光棒和错觉轮廓光棒的方位/方向调制曲线十分相似, 但对移动错觉棒和移动光棒的反应模式(潜伏期和反应时程)不同. 对由光栅组成的错觉轮廓而言, 细胞的反应大小与组成光栅的相位无关, 并且细胞对组成错觉轮廓光栅的最优空间频率比对普通移动光栅的最优空间频率要高得多, 说明细胞确实是对轮廓本身反应, 而不是对组成轮廓的光栅的末端反应. 某些速度调制类型的细胞对移动错觉棒反应的最优速度比对移动光棒的最优速度要低得多. 进一步验证了猫视皮层17, 18区部分细胞能对错觉轮廓反应, 并且观察到这些细胞对错觉轮廓和真实轮廓有不同的感受野反应特性, 提示视觉系统对两种刺激图形的检测机制可能存在着差异.     

猫纹状皮层神经元整合野的形态和范围

用正弦调制的移动光栅测量了１２０个猫纹状皮层神经元在长度和宽度方向上的空间整合特性。结果表明：（１）大多数细胞的传统感受野具有长宽相近的结构,然而它整合野多数是宽而短或者窄而长的长条形。（２）整合野大小为传统感受野的２－７倍,平均为３．７倍。简单细胞与复杂细胞整合野的大小没有显著差别。（３）随着感受野的视网膜偏心度的增加,整合野大小有逐渐增大的趋势。（４）感受野靠近视网膜垂直中线的细胞,其整合野可     

猫18区简单细胞感受野的时空结构

通过测量感受野内不同空间位置的时间传递函数,研究了猫18区简单细胞感受野的时空结构.不同位置在时间特性上的差异主要表现在绝对相移上.绝对相移与位置的关系有两类:一类随位置变化绝对相移呈现连续性改变;另一类随位置变化绝对相移发生180度跳变.将频域特性进行反Fourier变换可得到感受野在时空域的表达.连续变化型的时间特性和空问特性不能分离,其时空结构倾斜.跳变型的时空特性可以分离,在时空平面内不具有倾斜结构.根据感受野内不同位置的绝对相移可以预测神经元的最佳运动方向和最佳空间频率.     

视皮层细胞方位选择性研究的新进展

高等哺乳动物视皮层细胞方位选择性的形成机制一直是视觉研究的热点。近来许多新的实验事实和假说,使长期占统治地位的会聚学说面临挑战。不同假说争论的问题在于：皮层下的输入和皮层内作用二者谁在方位造反性的形成过程中起讯作用？皮层内对方位选择性强度的放大作用主要是兴奋性的还是抑制的？本文对有关这一研究的新进展作了评述,试图使人们更全面地认识视觉方位选择性这一经典感受野特性。     

The responses to illusory contours of neurons in cortex areas 17 and 18 of the cats

Responses to illusory contours (ICs) were sampled from neurons in cortical areas 17 and 18 of the anesthetized cats. For ICs sensitive cells, the differences of receptive field properties were compared when ICs and real contour stimuli were applied. Two hundred orientation or direction selective cells were studied. We find that about 42 percent of these cells were the ICs sensitive cells. Although their orientation or direction tuning curves to ICs bar and real bars were similar, the response modes (especially latency and time course) were different. The cells' responses to ICs were independent of the spatial phases of sinusoidal gratings, which composed the ICs. The cells' optimal spatial frequency to composing gratings the ICs was much higher than the one to moving gratings. Therefore, these cells really responded to the ICs rather than the line ends of composing gratings. For some kinds of velocity-tuning cells, the optimal velocity to moving ICs bar was much lower than the optimal velocity to moving     

The Detection of Orientation Continuity and Discontinuity by Cat V1 Neurons

The orientation tuning properties of the non-classical receptive field (nCRF or “surround”) relative to that of the classical receptive field (CRF or “center”) were tested for 119 neurons in the cat primary visual cortex (V1). The stimuli were concentric sinusoidal gratings generated on a computer screen with the center grating presented at an optimal orientation to stimulate the CRF and the surround grating with variable orientations stimulating the nCRF. Based on the presence or absence of surround suppression, measured by the suppression index at the optimal orientation of the cells, we subdivided the neurons into two categories: surround-suppressive (SS) cells and surround-non-suppressive (SN) cells. When stimulated with an optimally oriented grating centered at CRF, the SS cells showed increasing surround suppression when the stimulus grating was expanded beyond the boundary of the CRF, whereas for the SN cells, expanding the stimulus grating beyond the CRF caused no suppression of the center response. For the SS cells, strength of surround suppression was dependent on the relative orientation between CRF and nCRF: an iso-orientation grating over center and surround at the optimal orientation evoked strongest suppression and a surround grating orthogonal to the optimal center grating evoked the weakest or no suppression. By contrast, the SN cells showed slightly increased responses to an iso-orientation stimulus and weak suppression to orthogonal surround gratings. This iso-/orthogonal orientation selectivity between center and surround was analyzed in 22 SN and 97 SS cells, and for the two types of cells, the different center-surround orientation selectivity was dependent on the suppressive strength of the cells. We conclude that SN cells are suitable to detect orientation continuity or similarity between CRF and nCRF, whereas the SS cells are adapted to the detection of discontinuity or differences in orientation between CRF and nCRF.     

Asymmetric temporal properties in the receptive field of retinal transient amacrine cells

The speed of signal conduction is a factor determining the temporal properties of individual neurons and neuronal networks. We observed very different conduction velocities within the receptive field of fast-type On-Off transient amacrine cells in carp retina cells, which are tightly coupled to each other via gap junctions. The fastest speeds were found in the dorsal area of the receptive fields, on average five times faster than those detected within the ventral area. The asymmetry was similar in the On- and Off-part of the responses, thus being independent of the pathway, pointing to the existence of a functional mechanism within the recorded cells themselves. Nonetheless, the spatial decay of the graded-voltage photoresponse within the receptive field was found to be symmetrical, with the amplitude center of the receptive field being displaced to the faster side from the minimum-latency location. A sample of the orientation of varicosity-laden polyaxons in neurobiotin-injected cells supported the model, revealing that approximately 75% of these processes were directed dorsally from the origin cells. Based on these results, we modeled the velocity asymmetry and the displacement of amplitude center by adding a contribution of an asymmetric polyaxonal inhibition to the network. Due to the asymmetry in the conduction velocity, the time delay of a light response is proposed to depend on the origin of the photostimulus movement, a potentially important mechanism underlying direction selectivity within the inner retina.     

The responses to illusory contours of neurons in cortex areas 17 and 18 of the cats

Responses to illusory contours (ICs) were sampled from neurons in cortical areas 17 and 18 of the anesthetized cats. For ICs sensitive cells, the differences of receptive field properties were compared when ICs and real contour stimuli were applied. Two hundred orientation or direction selective cells were studied. We find that about 42 percent of these cells were the ICs sensitive cells. Although their orientation or direction tuning curves to ICs bar and real bars were similar, the response modes (especially latency and time course) were different. The cells’ responses to ICs were independent of the spatial phases of sinusoidal gratings, which composed the ICs. The cells’ optimal spatial frequency to composing gratings the ICs was much higher than the one to moving gratings. Therefore, these cells really responded to the ICs rather than the line ends of composing gratings. For some kinds of velocity-tuning cells, the optimal velocity to moving ICs bar was much lower than the optimal velocity to moving bars. The present results demonstrate that some cells in areas 17 and 18 of cats have the ability to respond to ICs and have different response properties of the receptive fields to ICs and luminance boundaries via different neural mechanisms.     

A multi-stage model for fundamental functional properties in primary visual cortex

Many neurons in mammalian primary visual cortex have properties such as sharp tuning for contour orientation, strong selectivity for motion direction, and insensitivity to stimulus polarity, that are not shared with their sub-cortical counterparts. Successful models have been developed for a number of these properties but in one case, direction selectivity, there is no consensus about underlying mechanisms. We here define a model that accounts for many of the empirical observations concerning direction selectivity. The model describes a single column of cat primary visual cortex and comprises a series of processing stages. Each neuron in the first cortical stage receives input from a small number of on-centre and off-centre relay cells in the lateral geniculate nucleus. Consistent with recent physiological evidence, the off-centre inputs to cortex precede the on-centre inputs by a small (～4 ms) interval, and it is this difference that confers direction selectivity on model neurons. We show that the resulting model successfully matches the following empirical data: the proportion of cells that are direction selective; tilted spatiotemporal receptive fields; phase advance in the response to a stationary contrast-reversing grating stepped across the receptive field. The model also accounts for several other fundamental properties. Receptive fields have elongated subregions, orientation selectivity is strong, and the distribution of orientation tuning bandwidth across neurons is similar to that seen in the laboratory. Finally, neurons in the first stage have properties corresponding to simple cells, and more complex-like cells emerge in later stages. The results therefore show that a simple feed-forward model can account for a number of the fundamental properties of primary visual cortex.     

The synaptic mechanism of direction selectivity in distal processes of starburst amacrine cells

Patch-clamp recordings revealed that distal processes of starburst amacrine cells (SACs) received largely excitatory synaptic input from the receptive field center and nearly purely inhibitory inputs from the surround during both stationary and moving light stimulations. The direct surround inhibition was mediated mainly by reciprocal GABA(A) synapses between opposing SACs, which provided leading and prolonged inhibition during centripetal stimulus motion. Simultaneous Ca(2+) imaging and current-clamp recording during apparent-motion stimulation further demonstrated the contributions of both centrifugal excitation and GABA(A/C)-receptor-mediated centripetal inhibition to the direction-selective Ca(2+) responses in SAC distal processes. Thus, by placing GABA release sites in electrotonically semi-isolated distal processes and endowing these sites with reciprocal GABA(A) synapses, SACs use a radial-symmetric center-surround receptive field structure to build a polar-asymmetric circuitry. This circuitry may integrate at least three levels of interactions--center excitation, surround inhibition, and reciprocal inhibitions that amplify the center--surround antagonism-to generate robust direction selectivity in the distal processes.     

猫后内侧上雪氏区视神经元对运动棒反应的取向和方向特征

猫后内侧上雪区（ｐｏｓｔｅｒｏｍｅｄｉａｌｌａｔｅｒａｌｓｕｐｒａｓｙｌｖｉａｎａｒｅａ,ＰＭＬＳ）的绝大多数神经元（１７１／２００）对运动棒的取向调谐,６２％（１２４／２００）细胞的取向调谐宽度（半高波宽）小于９０°：按方向选择性和取向选择性可分辨出几类特征明显的细胞类型：１、强取向和强方向选择性细胞;２、强取向调谐的双向选择细胞;３、弱取向调谐的强方向选择细胞;４、无取向无方向选择性细胞;以及５、特征不明显的或中间类型细胞。它们与最近光学记录揭示的鹰猴中颞叶视区（ｍｉｄｄｌｅｔｅｍｐｏｒａｌｖｉｓｕａｌａｒｅａ,ＭＴ）的组织有很好的吻合。     

Orientation-selective neurons in the squirrel visual cortex

The organization of receptive fields of neurons sensitive to orientation of visual stimuli was investigated in the squirrel visual cortex. Neurons with mutually inhibitory on- and off-areas of the receptive field, with partially and completely overlapping excitatory and inhibitory mechanisms, were distinguished. Neurons of the second group are most typical. They exhibit orientation selectivity within the excitatory area of the receptive field because, if the stimulus widens in the zero direction, perpendicular to the preferred direction, lateral inhibition is much stronger than if it widens in the preferred direction. Additional inhibitory areas (outside the excitatory area) potentiate this inhibition and increase selectivity. It is suggested that there is no strict separation of simple (with separate excitatory and inhibitory mechanisms in the receptive field) and complex (with overlapping of these mechanisms) neurons in the squirrel visual cortex.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 11, No. 6, pp. 540–549, November–December, 1979.     

Prediction of orientation selectivity from receptive field architecture in simple cells of cat visual cortex

From the intracellularly recorded responses to small, rapidly flashed spots, we have quantitatively mapped the receptive fields of simple cells in the cat visual cortex. We then applied these maps to a feedforward model of orientation selectivity. Both the preferred orientation and the width of orientation tuning of the responses to oriented stimuli were well predicted by the model. Where tested, the tuning curve was well predicted at different spatial frequencies. The model was also successful in predicting certain features of the spatial frequency selectivity of the cells. It did not successfully predict the amplitude of the responses to drifting gratings. Our results show that the spatial organization of the receptive field can account for a large fraction of the orientation selectivity of simple cells.