外膝状体神经元的亮度反应与感受野ON-OFF反应的关系

外膝体是视觉信息进入新皮层的主要通路,其编码亮度信息的神经机制还不清楚.我们采用随机呈现的连续快速变化(50 Hz)的均匀亮度刺激,显著地提高了猫外膝体神经元对均匀亮度的反应强度,通过反相关算法抽提出神经元的亮度反应函数.约81%的神经元的亮度反应函数为单调性上升或下降,有19%的神经元亮度反应函数为V型.通过分析这些神经元对亮度上升和下降的反应强度与感受野ON和OFF反应强度的关系,表明83%的神经元对亮度的反应模式是由其感受野ON-OFF反应的相对强度决定的,其余17%则与其感受野ON-OFF区的兴奋和抑制的变化相关.这些结果揭示了外膝体神经元编码亮度变化的机制.     

扣带皮层前部对丘脑腹侧基底核触觉神经元ON-OFF反应的易化性影响

实验在41只氨基甲酸乙酯(25%)麻醉的SD雄性大鼠上进行. 采用细胞外微电极记录的方法, 首先观察了大鼠丘脑腹侧基底核(ventrobasal thalamus nucleus, VB) ON-OFF神经元的触觉反应特性, 并对不同的ON-OFF反应进行了分型, 分析了ON-OFF反应不同型式与刺激参数之间的关系. 在此基础上, 进一步考察了刺激扣带皮层前部(anterior cingulate cortex, ACC)对ON-OFF神经元触觉反应的影响. 实验发现, ACC的刺激对触觉ON-OFF神经元的OFF反应有易化效应, 表现在降低OFF反应的阈值, 从而改变了反应构型, 或者增加原OFF反应的放电频数. 在关于ON-OFF反应神经元感受野的研究中发现, 扣带皮层前部的兴奋可以使ON-OFF反应神经元感受野边缘的ON反应改变为ON-OFF反应, 从而使其对感受野内刺激物运动的时空信息的编码更为明确, 发现了边缘系统能够对触觉上传信息进行调制.     

刺激大鼠杏仁外侧核对皮层A Ⅰ区ON-OFF神经元反应及调谐曲线的影响

在30只氨基甲酸乙酯麻醉的SD大鼠上记录神经元单位放电,观察短纯音诱发的皮层A Ⅰ区神经元ON-OFF反应的特性及电刺激杏仁外侧核(lateral amygdaloid nucleus,LA)对ON-OFF反应以及调谐曲线的影响.实验证实,A Ⅰ区神经元ON-OFF反应的模式与纯音刺激的强度、频率及作用时程有关;刺激LA可以抑制ON-OFF反应的放电频数,使反应的阈值升高,或使反应放电构型发生变化;此外,刺激LA能使ON-OFF神经元的调谐曲线变窄,Q10数值增大.研究结果不仅表明ON-OFF神经元能对纯音刺激的时程、强度和频率等多种信息进行编码,而且还证明杏仁外侧核可以在皮层水平参与听觉信息的调制,削弱或衰减某些听觉信息,导致整个调谐曲线上移变窄,从而提高A Ⅰ区ON-OFF神经元的频率选择性能,有利于检测外界嘈杂环境中特定的听觉信息.     

猫外膝体神经元兴奋和抑制过程的相互作用

用神经脉冲自动计数的方法,定量地研究了猫外膝体神经元兴奋与抑制过程在时间和空间上的相互作用。1.时间上的相互作用:在亮度变化瞬间,神经细胞的兴奋或抑制过程增强,表现为放电频率暂时地明显增加或减少。这一作用持续的时间(适应时间),瞬变细胞约0.1秒,持续性小感受野细胞约4～8秒,持续性大感受野细胞约40～80秒。2.空间上的相互作用:改变被照射视网膜面积的大小,同时计算神经元的平均放电频率,可以确定外周抑制区的存在、范围和强度。外周抑制可以同时作用于给-撤细胞的给光反应和撤光反应,也可以选择地只抑制其中某一种反应。纯撤光细胞没有外周抑制区。具有大感受野的外膝体神经元,其感受野的大小与背景光的强度有关。在高亮度情况下,感受野变小,有利于改善分辨率;在低亮度下,感受野扩大,有利于提高灵敏度。     

猫外膝体神经元兴奋和抑制过程的相互作用

用神经脉冲自动计数的方法,定量地研究了猫外膝体神经元兴奋与抑制过程在时间和空间上的相互作用。1.时间上的相互作用:在亮度变化瞬间,神经细胞的兴奋或抑制过程增强,表现为放电频率暂时地明显增加或减少。这一作用持续的时间(适应时间),瞬变细胞约0.1秒,持续性小感受野细胞约4～8秒,持续性大感受野细胞约40～80秒。2.空间上的相互作用:改变被照射视网膜面积的大小,同时计算神经元的平均放电频率,可以确定外周抑制区的存在、范围和强度。外周抑制可以同时作用于给-撤细胞的给光反应和撤光反应,也可以选择地只抑制其中某一种反应。纯撤光细胞没有外周抑制区。具有大感受野的外膝体神经元,其感受野的大小与背景光的强度有关。在高亮度情况下,感受野变小,有利于改善分辨率;在低亮度下,感受野扩大,有利于提高灵敏度。     

刺激家兔杏仁基底核对外膝体“ON”神经元感受野反应特性的影响

利用细胞外记录,在乌拉坦麻醉和三碘委按酚麻痹的３７只新西兰兔上,考察刺激杏仁基底核对外膝体神经元感受野反应特性的影响。实验观察到,在１１７个具有“ＯＮ”反应外膝体神经元中,４７个单位在电刺激杏仁基底核时不同程度地受到抑制或易化性影响,即细胞反应强度和／或感受野范围均有改变。产生这些影响的潜伏期为７－４１ｍｓ,最大时程约２８ｍｓ。撤除电刺激３ｍｉｎ后,感受野反应特性即告恢复。单独刺激杏仁核不能引起外     

特定方位的光栅识别学习不改变猫外膝体背核神经元的方位敏感性

视觉信号识别训练可改变视觉通路神经元的可塑性, 其神经机制尚不清楚。已有少数研究显示, 动物(猴) 长时间进行特定方位的光栅识别学习后, 视皮层部分神经元对视觉刺激的反应表现出与学习任务相关的敏感性变化。这种敏感性变化是否亦存在于皮层下结构尚无报道。本实验训练两只成年猫分别进行水平和垂直方位的条形静止正弦光栅的识别以获得食物奖赏, 两只猫的行为识别能力逐渐提高, 4 个多月后识别的正确率达85%以上, 用与训练方位垂直的正弦光栅检测发现, 识别正确率明显下降。细胞外记录外膝体背核(Dorsal lat eral geniculate nucleus, dLGN) 神经元对不同方位正弦光栅刺激的反应显示, 与正常猫相比, 训练猫外膝体细胞的最优方位并未向着训练方位发生明显改变, 对于感受野位于中央区15度视角以内的细胞来说, 其方位选择性强度以及在训练方位的发放强度与正常猫无明显差异。以上结果表明, 猫对特定方位的光栅识别学习不改变外膝体神经元的方位敏感性, 其行为上方位识别特异性的提高可能与视皮层细胞的方位编码可塑性有关。     

猫外膝体神经元对光栅刺激方位倾向性的一种分布特性

我们检测了猫外膝体(LGN)神经元对闪烁方波光栅的反应与光栅方位的关系。对一定对比度和空间频率的光栅,26个记录到的神经元的反应均随方位不同而变化,其最大反应和最小反应的比值平均为3.0±0.3(S.E.)。神经元的最优方位(即引起最大反应的光栅方位)随其感受野中心在视网膜上的位置而异,具有平行于其各自感受野中心与视网膜中央区(area cen-sralis)的连接线的倾向。     

综述: 初级视皮层的亮度信息加工

亮度(luminance)是最基本的视觉信息.与其他视觉特征相比,由于视神经元对亮度刺激的反应较弱,并且许多神经元对均匀亮度无反应,对亮度信息编码的神经机制知之甚少.初级视皮层部分神经元对亮度的反应要慢于对比度反应,被认为是由边界对比度诱导的亮度知觉(brightness)的神经基础.我们的研究表明,初级视皮层许多神经元的亮度反应要快于对比度反应,并且这些神经元偏好低的空间频率、高的时间频率和高的运动速度,提示皮层下具有低空间频率和高运动速度通路的信息输入对产生初级视皮层神经元的亮度反应有贡献.已经知道初级视皮层神经元对空间频率反应的时间过程是从低空间频率到高空间频率,我们发现的早期亮度反应是对极低空间频率的反应,与这一时间过程是一致的,是这一从粗到细的视觉信息加工过程的第一步,揭示了处理最早的粗的视觉信息的神经基础.另外,初级视皮层含有偏好亮度下降和高运动速度的神经元,这群神经元的活动有助于在光照差的环境中检测高速运动的低亮度物体.     

初级视皮层的亮度信息加工

亮度(luminance)是最基本的视觉信息.与其他视觉特征相比,由于视神经元对亮度刺激的反应较弱,并且许多神经元对均匀亮度无反应,对亮度信息编码的神经机制知之甚少.初级视皮层部分神经元对亮度的反应要慢于对比度反应,被认为是由边界对比度诱导的亮度知觉(brightness)的神经基础.我们的研究表明,初级视皮层许多神经元的亮度反应要快于对比度反应,并且这些神经元偏好低的空间频率、高的时间频率和高的运动速度,提示皮层下具有低空间频率和高运动速度通路的信息输入对产生初级视皮层神经元的亮度反应有贡献.已经知道初级视皮层神经元对空间频率反应的时间过程是从低空间频率到高空间频率,我们发现的早期亮度反应是对极低空间频率的反应,与这一时间过程是一致的,是这一从粗到细的视觉信息加工过程的第一步,揭示了处理最早的粗的视觉信息的神经基础.另外,初级视皮层含有偏好亮度下降和高运动速度的神经元,这群神经元的活动有助于在光照差的环境中检测高速运动的低亮度物体.     

Predictive Feedback Can Account for Biphasic Responses in the Lateral Geniculate Nucleus

Biphasic neural response properties, where the optimal stimulus for driving a neural response changes from one stimulus pattern to the opposite stimulus pattern over short periods of time, have been described in several visual areas, including lateral geniculate nucleus (LGN), primary visual cortex (V1), and middle temporal area (MT). We describe a hierarchical model of predictive coding and simulations that capture these temporal variations in neuronal response properties. We focus on the LGN-V1 circuit and find that after training on natural images the model exhibits the brain's LGN-V1 connectivity structure, in which the structure of V1 receptive fields is linked to the spatial alignment and properties of center-surround cells in the LGN. In addition, the spatio-temporal response profile of LGN model neurons is biphasic in structure, resembling the biphasic response structure of neurons in cat LGN. Moreover, the model displays a specific pattern of influence of feedback, where LGN receptive fields that are aligned over a simple cell receptive field zone of the same polarity decrease their responses while neurons of opposite polarity increase their responses with feedback. This phase-reversed pattern of influence was recently observed in neurophysiology. These results corroborate the idea that predictive feedback is a general coding strategy in the brain.     

Responses of Neurons of the Extrastriate Area 21b of the Cat Brain Cortex to Changes in Orientation of Moving Visual Stimuli

We studied the responses of neurons of the extrastriate cortical area 21b of the cat to changes in orientation of the movements of visual stimuli within the receptive field (RF) of the neuron under study. Our experiments demonstrated that 24 of 108 cells (22%) responded differentially to a certain extent to orientation of the movements of visual stimuli. As a whole, neurons of the area 21b did not demonstrate fine tuning on the optimum angle of orientation. In many cases, neuronal responses to different orientations of the movement of visual stimulus depended significantly on specific parameters of this stimulus (its shape, dimensions, and contrast). Some directionally sensitive neurons responded to a change in orientation of the movement of visual stimuli by modification of the index of directionality. We also studied spatial organization of the RF of neurons with the presentation of stationary visual stimuli. Comparison of the neuronal responses to a change in orientation of the movements of stimuli and to presentation of stationary stimuli showed that the correlation between the orientation sensitivity of the neuron under study and the stationary functional organization of its RF was insignificant. We hypothesize that inhibitory processes and subthreshold influences from a space surrounding the RF play a special role in the formation of the neuronal responses generated in the associative visual cortical regions to visual stimulation.     

Responses of neurons of the extrastriate cortex of the cat brain to moving stimuli of different forms

We examined responses of neurons of the field 21b of the cat brain cortex to presentation of moving visual stimuli of different forms. Characteristics of the responses of about 54% of the studied neurons showed that in these cases configurations of the contours of moving stimuli were to a certain extent discriminated. Most neurons selectively reacting to changes in the form of the stimulus were dark-sensitive units (they generated optimum responses to presentation of dark visual stimuli on the light background). Detailed examination of the spatial infrastructure of receptive fields (RFs) of the neurons and comparison of this structure with the selectivity of neuronal responses showed that there is no significant correlation between static organization of the RF and responses of the neuron to the movements of stimuli of different forms. We hypothesize that the dynamic infrastructure of the RF and the combined activity of functional groups of neurons, whose RFs spatially overlap the RF of the neuron under study, play a definite role in the mechanisms responsible for neuronal discrimination of the form of the visual stimulus. Neirofiziologiya/Neurophysiology, Vol. 38, No. 1, pp. 61–71, January–February, 2006.     

A linear model for symmetric receptive fields: implications for classification tests with flashed and moving images

The purpose of this study was to explore the effects of spatial and temporal properties on the expected responses of visual neurons that have linear receptive fields (RFs), particularly those having a mirror symmetric distribution of spatial subregions. Receptive fields that are symmetric in at least one spatial dimension occur in neurons of the retina, the lateral geniculate nucleus (LGN), and the visual cortex of mammals. Responses to flashing bars, moving bars, and moving edges were studied for different configurations of an analog RF model in which spatial and temporal aspects were varied independently. Responses of the model at intermediate stimulus speeds were found to agree with responses in the literature for X and Y units of the LGN and often for simple units of the visual cortex. In particular, having separated regions of response to light and dark edges, an identifying property of simple cells, was found to be a linear consequence of RF regions responding inversely to stimuli of opposite polarity. Model differences from responses of cortical complex units show that a linear model cannot mimic their responses, and imply that complex units employ major nonlinearities in coding image polarity (light vs dark), which signifies a nonlinearity in coding intensity. Because sudden flux changes inherent in flashing bars test mainly temporal RF properties, and slowly moving edges test mainly spatial properties, these two tests form a useful minimal set with which to describe and classify RFs. The usefulness of this set derives both from its sensitivity to spatial and temporal variables, and from the correlation between the linearity of a cell's processing of stimulus intensity and its RF classification.     

Receptive Fields of Visually Sensitive Neurons of the Extrastriate Associative Area 21b of the Cat Cerebral Cortex

Organization of the receptive fields (RFs) of neurons of the extrastriate associative region 21b of the cerebral cortex was studied in cats. Most neurons under study (63%) were “monocular,” while 37% of the cells were “binocular” units. Among 178 neurons examined in detail, heterogeneous RF functional organization was typical of about 76% of the units; point-to-point testing of the entire RF area by stationary stimuli resulted in the generation of various types of responses (on, off, or on-off). The rest of the neurons (24%) generated homogeneous responses. The dimension, form, and functional organization of RFs of the neurons under study depended to a certain extent on the parameters of visual stimuli used for the measurements. Examination of the influence of the visual space, which surrounded the RF, on responses of the neurons evoked by stimulation of the RF per se showed that darkening of the visual space adjacent to the RF inhibited neuronal responses to moving stimuli; in some cases the responses were totally suppressed. Analysis of spatial overlapping of the RF sequentially recorded in the course of each insertion of the electrode showed that the density of distribution of the overlapping RF areas of neighboring neurons with the RF of the examined neuron is irregular, and that the RF is of a mosaic nature. We hypothesize that the visual space surrounding the RF plays a significant role in the formation of responses of visually sensitive neurons to presentation of moving stimuli. Neirofiziologiya/Neurophysiology, Vol. 37, No. 3, pp. 223–234, May–June, 2005.     

视觉图形诱发的瞳孔反应

以图形变化刺激进行瞳孔反应研究,实验表明：（１）空间平均亮度守恒的光栅或棋盘格图形翻转能激发起瞳孔反应,为瞬态收缩波形,与ｐｕｐｉｌｌａｒｙ ｅｓｃａｐｅ相似;（２）光栅或棋盘格的空间频率的变化也能引起瞳孔反应,且反应幅度随空间频率差别增大而变大;（３）从均匀亮背景变化到棋盘格图形或者从棋盘格图形变化到黑暗背景,虽然不存在任何局部亮度增强,皆能引起瞳孔反应。实验结果明确证明了人的瞳孔反应系统除接收     

Modulation of V1 Spike Response by Temporal Interval of Spatiotemporal Stimulus Sequence

The spike activity of single neurons of the primary visual cortex (V1) becomes more selective and reliable in response to wide-field natural scenes compared to smaller stimuli confined to the classical receptive field (RF). However, it is largely unknown what aspects of natural scenes increase the selectivity of V1 neurons. One hypothesis is that modulation by surround interaction is highly sensitive to small changes in spatiotemporal aspects of RF surround. Such a fine-tuned modulation would enable single neurons to hold information about spatiotemporal sequences of oriented stimuli, which extends the role of V1 neurons as a simple spatiotemporal filter confined to the RF. In the current study, we examined the hypothesis in the V1 of awake behaving monkeys, by testing whether the spike response of single V1 neurons is modulated by temporal interval of spatiotemporal stimulus sequence encompassing inside and outside the RF. We used two identical Gabor stimuli that were sequentially presented with a variable stimulus onset asynchrony (SOA): the preceding one (S1) outside the RF and the following one (S2) in the RF. This stimulus configuration enabled us to examine the spatiotemporal selectivity of response modulation from a focal surround region. Although S1 alone did not evoke spike responses, visual response to S2 was modulated for SOA in the range of tens of milliseconds. These results suggest that V1 neurons participate in processing spatiotemporal sequences of oriented stimuli extending outside the RF.     

Functional mechanisms shaping lateral geniculate responses to artificial and natural stimuli

Functional models of the early visual system should predict responses not only to simple artificial stimuli but also to sequences of complex natural scenes. An ideal testbed for such models is the lateral geniculate nucleus (LGN). Mechanisms shaping LGN responses include the linear receptive field and two fast adaptation processes, sensitive to luminance and contrast. We propose a compact functional model for these mechanisms that operates on sequences of arbitrary images. With the same parameters that fit the firing rate responses to simple stimuli, it predicts the bulk of the firing rate responses to complex stimuli, including natural scenes. Further improvements could result by adding a spiking mechanism, possibly one capable of bursts, but not by adding mechanisms of slow adaptation. We conclude that up to the LGN the responses to natural scenes can be largely explained through insights gained with simple artificial stimuli.