Reciprocal Interaction of On- and Off-Systems as a Mechanism of Sensitivity of Visual Neurons to The Orientation of the Vector of the Brightness Gradient in Cats

An experimental study has been performed on neuronal mechanisms of sensitivity of cat visual neurons (lateral geniculate body) to the value and orientation of the vector of brightness gradient in a test stimulus. With changes of the value and orientation of the brightness gradient vector, there exists an optimal (preferred) orientation of the gradient vector, at which the neuronal response is maximal. The sensitivity of neurons to the brightness gradient at shifts of the gradient vector towards the preferred orientation increases not due to an increased excitation in neuronal reactions, but due to a reduction of reciprocal (on- and off-) inhibition, affecting this neuron, of adjacent neurons in neuronal pools. The reciprocal inhibitory interaction of on- and off-systems is enhanced by inhibiting the response of the antagonistic neuron at shifts of the brightness gradient vector in the stimulus from the preferred to the non-preferred orientation. This reciprocal inhibitory interaction is clearly seen in pairs of on- and off-neurons with superposed receptive fields (RF) at their simultaneous analysis of on- and off-responses at a change of the orientation of the brightness gradient vector by 180 degrees. Dependencies of the parameters (duration and intensity of inhibitory phases in responses) of reciprocal inhibitory interaction on orientation of the brightness gradient vector in RF of neurons are determined. Dependencies of responses of the total sample of neurons, which are plotted for on- and off-neurons, to their adequate and inadequate (on- and off-) stimuli on the orientation of the brightness gradient vector are inversely proportional.     

Sensitivity of the cat lateral geniculate body neurons to orientation of the brightness gradient vector

A new property of visual neurons: their sensitivity to orientation and the vector brightness gradient, was revealed and described. Receptive fields of the lateral geniculate body neurons in the cat have preferred orientation maximum reaction (average mean of orientation sensitivity coefficient--0.55 +/- 0.20). The preferred orientation mainly has a radial or tangential trend in the visual field. Temporal characteristics of the neuronal responses were analysed. A role of inhibition processes in the orientation sensitivity is discussed.     

Segregation of object and background motion in visual area MT: effects of microstimulation on eye movements

To track a moving object, its motion must first be distinguished from that of the background. The center-surround properties of neurons in the middle temporal visual area (MT) may be important for signaling the relative motion between object and background. To test this, we microstimulated within MT and measured the effects on monkeys' eye movements to moving targets. We found that stimulation at "local motion" sites, where receptive fields possessed antagonistic surrounds, shifted pursuit in the preferred direction of the neurons, whereas stimulation at "wide-field motion" sites shifted pursuit in the opposite, or null, direction. We propose that activating wide-field sites simulated background motion, thus inducing a target motion signal in the opposite direction. Our results support the hypothesis that neuronal center-surround mechanisms contribute to the behavioral segregation of objects from the background.     

蜻蜒运动检测神经元的光谱反应

蜻蜒腹神经束上存在着自运动检测神经元和目标运动检测神经元.我们采用了两种视觉刺激条件来测试自运动检测神经元的光谱反应.当采用控制强度和波长的闪光进行测试时、它们的光谱反应曲线与绿色光感受器的光谱灵敏度曲线极其相似,峰值位于500nm处.然而采用运动的条纹进行测试时,它们的峰值却位于560nm处.当用一种颜色的运动图案作为目标放置在另一种颜色背景的前方测试时,发现存在某个目标的照明强度值能使反应下降到自发放电的水平,这表明自运动检测器无法检测这二种颜色的差别,即它们是色盲的、它主要接受来自绿色光感受器的信号.目标运动检测神经元的光谱反应特性与自运动检测神经元的不同,目标运动检测神经元在以380nm至580nm的范围中有着平坦的光谱反应曲线,有时在紫外频段出现峰有(?)前景与背景颜色不同且固定背景光的颜色与强度而改变前景的光强时,神经元的反应不会下降到自发放电水平,当背景为绿色而目标为另一个颜色.特别是兰色时,神经元反应强烈,但当背景为兰色而目标为绿色时,它们的反应相对较弱.这些结果表明目标运动检测神经元是对颜色敏感的.     

Afferent signal transformation at subcortical levels of the visual system

A comparison was made of the characteristics of the background and light-induced impulse activity of 350 neurons of the retina and laterial geniculate body (LGB) of an immobilized cat. In the LGB, in contrast to the ganglion cells of the retina, there are relatively more neurons with high-light sensitivity, low latency of response, short-initial discharge, and low summation time, as well as rapid recovery of reactivity. It was found that these changes are characteristic of cells connected with the peripheral region of the retina. In the central channel and the system of long-latency cells, according to a number of parameters, no such transformations are found, while other characteristics change in the opposite direction. In the central channel at the postgenicular level, the frequency of the background and induced impulsation decreases, while in the peripheral channel it increases considerably. The results obtained indicate differences in the organization of the synaptic inputs of LGB neurons of different groups. The intensification and time compression of the sensory transmission in the peripheral channel are explained by the reticular properties of the structures formed by the branching and overlapping fibers coming from the retina, as well as by the great effectiveness of the successive inhibition in the LGB which blocks the late group of initial discharges. These transformations of the afferentation can provide reliability and rapidity of the detection of weak, slowly increasing, and moving light signals.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 3, No. 1, pp. 13–21, January–February, 1971.     

Simulation of Neuronal Responses Defining Depth Order and Contrast Polarity at Illusory Contours in Monkey Area V2

Neurophysiological, brain imaging, and perceptual studies in animals and humans suggest that illusory (occluding) contours are represented at an early level of visual cortical processing. Comparatively little is known about the mechanisms defining the depth order and the brightness illusion associated with such contours. Baumann et al. (1997) found neurons in area V2 of the alert monkey that signaled not only illusory contours but also the figure-ground direction that human observers perceive at such contours. The majority of these neurons showed this property independent stimulus contrast; a small minority preferred a certain combination of figure-ground direction and contrast polarity at these contours. In this article, we simulate the responses of these neurons by means of a grouping mechanism that uses occlusion cues (line-ends, corners) to define figure-ground direction and contrast polarity at such contours.     

Quantification of directional and orientational selectivities of visual neurons to moving stimuli

Directional and orientational components usually coexist and are mixed in the cell's overall responses when moving optical stimuli are used to study the response characteristics of visual neurons. While these two properties were quantified with all the previous methods for data analysis, their effects could not be efficiently separated from each other, and thus the analyses were imperfect. In this paper, theoretical evidence and examples are provided to show the defects of the old methods. In order to separate the two components completely, we propose to apply optimal regression analysis with the sine-cosine function series as the fundamental variables. Based on this separation, we defined the orientational selectivity as variation of response strength with orientation and performed integration and averaging to quantify the two properties [cf. Eqs. (5) and (6)]. The present method has the advantages of completeness and accuracy, and can detect some details which would have been missed by other methods. An explanation of the intrinsic implications of the method and our comprehension of directional and orientational selectivities and preferred direction and orientation are also given. Received: 4 January 1993/Accepted in revised form: 1 July 1993     

Visual Receptive Field Properties of Neurons in the Mouse Lateral Geniculate Nucleus

The lateral geniculate nucleus (LGN) is increasingly regarded as a “smart-gating” operator for processing visual information. Therefore, characterizing the response properties of LGN neurons will enable us to better understand how neurons encode and transfer visual signals. Efforts have been devoted to study its anatomical and functional features, and recent advances have highlighted the existence in rodents of complex features such as direction/orientation selectivity. However, unlike well-researched higher-order mammals such as primates, the full array of response characteristics vis-à-vis its morphological features have remained relatively unexplored in the mouse LGN. To address the issue, we recorded from mouse LGN neurons using multisite-electrode-arrays (MEAs) and analysed their discharge patterns in relation to their location under a series of visual stimulation paradigms. Several response properties paralleled results from earlier studies in the field and these include centre-surround organization, size of receptive field, spontaneous firing rate and linearity of spatial summation. However, our results also revealed “high-pass” and “low-pass” features in the temporal frequency tuning of some cells, and greater average contrast gain than reported by earlier studies. In addition, a small proportion of cells had direction/orientation selectivity. Both “high-pass” and “low-pass” cells, as well as direction and orientation selective cells, were found only in small numbers, supporting the notion that these properties emerge in the cortex. ON- and OFF-cells showed distinct contrast sensitivity and temporal frequency tuning properties, suggesting parallel projections from the retina. Incorporating a novel histological technique, we created a 3-D LGN volume model explicitly capturing the morphological features of mouse LGN and localising individual cells into anterior/middle/posterior LGN. Based on this categorization, we show that the ON/OFF, DS/OS and linear response properties are not regionally restricted. Our study confirms earlier findings of spatial pattern selectivity in the LGN, and builds on it to demonstrate that relatively elaborate features are computed early in the visual pathway.     

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.     

Orientational and directional selectivities of visual neurons in the superior colliculus of the cat

Based on quantitative analyses of the response characteristics of visual neurons in the superior colliculus to moving optical bar stimuli, it is demonstrated for the first time that the visual neurons in superior colliculus of the cat have, to some extent, orientational selectivity. The significance of this selectivity is discussed in reference to its morphological substrate and physiological functions. In addition, both the directional and orientational selectivities in the superior colliculus are relatively weak when compared with those in the primary visual cortex, and the majority of the neurons prefer upward or downward motion in the visual field.     

Neural mechanisms of tactile motion integration in somatosensory cortex

How are local motion signals integrated to form a global motion percept? We investigate the neural mechanisms of tactile motion integration by presenting tactile gratings and plaids to the fingertips of monkeys, using the tactile analogue of a visual monitor and recording the responses evoked in somatosensory cortical neurons. The perceived directions of the gratings and plaids are measured in parallel psychophysical experiments. We identify a population of somatosensory neurons that exhibit integration properties comparable to those induced by analogous visual stimuli in area MT and find that these neural responses account for the perceived direction of the stimuli across all stimulus conditions tested. The preferred direction of the neurons and the perceived direction of the stimuli can be predicted from the weighted average of the directions of the individual stimulus features, highlighting that the somatosensory system implements a vector average mechanism to compute tactile motion direction that bears striking similarities to its visual counterpart.     

Orientation sensitive properties of visually driven neurons in extrastriate area 21a of cat cortex

Orientation sensitive properties of extrastriate area 21a neurons were investigated. Special attention was paid to the qualitative characteristics of neuron responses to the different orientations of visual stimulus motion across neuron classical receptive fields (CRF). The results of experiments have shown that a group of neurons (31%) in area 21a with specialized responses to moving visual stimuli changed their direction selective (DS) characteristics depending on the orientation of the stimulus movement. Some neurons reveal an abrupt drop of the direction sensitivity index (DI) to certain orientation (58%), and some show significant increase of DI at one of applied orientations of stimulus motion (22%). Detailed investigation of response patterns of non-directional neurons to different orientations of stimulus motion have revealed clear-cut qualitative differences, such as different regularities in the distribution of inter-peak inhibitory intervals in the response pattern in dependence of the orientation of stimulus motion. The investigation of neuron CRF stationary functional organization did not reveal correlations between RF's spatial functional organization, and that of qualitative modulations of neuron response patterns. A suggestion was put forward, that visual information central processing of orientation discrimination is a complex integrative process that includes quantitative as well as qualitative transformations of neuron activity.     

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     

视觉细胞反应中方向选择性和取向选择性成分的分离

运动的光刺激常被用于研究视觉神经细胞的方向和取向选择性.但是,在细胞的反应中,两种因素同时存在且混杂在一起; 以往的各种分析方法均因未能有效地解决这一问题而不够准确,对此我们给出了理论和实例证明.本工作引入正余弦函数系对实验数据作最优回归分析.拟合程度好于以往的方法.并且从根本上分离了反应中的方向和取向选择性成分.使对这两种性质的准确定量成为可能.本文对此方法的内在含义作出了解释,并提出了对方向、取向选择性及最优方向、取向的理解.     

Dynamic decorrelation as a unifying principle for explaining a broad range of brightness phenomena

The visual system is highly sensitive to spatial context for encoding luminance patterns. Context sensitivity inspired the proposal of many neural mechanisms for explaining the perception of luminance (brightness). Here we propose a novel computational model for estimating the brightness of many visual illusions. We hypothesize that many aspects of brightness can be explained by a dynamic filtering process that reduces the redundancy in edge representations on the one hand, while non-redundant activity is enhanced on the other. The dynamic filter is learned for each input image and implements context sensitivity. Dynamic filtering is applied to the responses of (model) complex cells in order to build a gain control map. The gain control map then acts on simple cell responses before they are used to create a brightness map via activity propagation. Our approach is successful in predicting many challenging visual illusions, including contrast effects, assimilation, and reverse contrast with the same set of model parameters.     

Double Receptive Fields of Neurons of the Lateral Geniculate Body of the Cat

We studied spatial organization of the receptive fields (RF) of neurons of the lateral geniculate body (LGB) on unanesthetized cats (pretrigeminal brainstem section). After identification of localization and borders of the RF of the neuron under study, we scanned with a sufficient resolution the entire visual field and tried to find additional space zones, whose stimulation could influence the impulse activity of the neuron. These experiments demonstrated that 24 neurons of 167 examined units (14%) could react to the presentation of visual stimuli within the visual space outside the main RF, but we were unable to determine borders of these additional zones with a sufficient accuracy. In 12 neurons (7% of the group under study), localization, dimensions, and specific features of an additional RF (RF-2) could be clearly determined. As a rule, the center of the RF-2 was localized at a distance of 20-40^O; or even farther from the center of the main RF (RF-1). To activate the neuron from the RF-2, application of greater visual stimuli was necessary (as compared with stimulation of the RF-1). Thus, two RF of one and the same neuron had dissimilar spatial organizations and qualitatively differed from each other in their stationary and dynamic characteristics. Considering our findings, we hypothesize that the RF-2 of LGB neurons can play a certain role in perception of large objects within the visual field of the animal by promoting formation of the avoidance reaction.     

老年猫视皮层细胞对刺激反应的对比敏感度下降伴随皮层内抑制作用减弱(英文)

对人类和动物的心理学研究证实,老年个体的视觉对比敏感度相对青年个体显著下降。为揭示其可能的神经机制,采用在体细胞外单细胞记录技术研究青、老年猫(Felis catus)初级视皮层(primary visual cortex,V1)细胞对不同视觉刺激对比度的调谐反应。结果显示,老年猫V1细胞对视觉刺激反应的平均对比敏感度比青年猫显著下降,这与灵长类报道的研究结果相一致,表明衰老影响视皮层细胞对视觉刺激反应的对比敏感度是灵长类和非灵长类哺乳动物中普遍存在的现象,并可能是介导老年性视觉对比敏感度下降的神经基础。另外,与青年猫相比,老年猫初级视皮层细胞对视觉刺激的反应性显著增强,信噪比下降,感受野显著增大,表明衰老导致的初级视皮层细胞对视觉刺激反应的对比敏感度下降伴随着皮层内抑制性作用减弱。     

Different properties of two groups of orientation discriminators in the cat visual cortex

The properties of 149 neurons, divided into two groups, were investigated during acute experiments on immobilized cats. These consisted of "timers" (37%) in which latency of response and time taken for reaction to peak changed in an orientation range of not more than 10 msec. The remaining 63% consisted of "scanners" [2]. "Timers" reliably differed from "scanners" in their shorter latent periods, rising time of discharge rate, duration of response, and higher rate of impulsation at all orientations of the stimulus. "Scanners" display greater orientational tuning and "scan" much more frequently throughout the orientation range. The pattern of acuity of orientational tuning is counterphasic during response in neurons of these two groups, while the distribution of their preferred orientation is complementary in nature. Both timers and scanners were found in the orientation columns of the visual cortex on most occasions, with the latter predominating. Columns consisting of only timers or scanners were met with more seldom. The significance of the differences between the properties of the two groups of neurons in the visual cortex is discussed with a view to orientational discrimination.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 18, No. 1, pp. 85–92, January–February, 1986.     

Unit activity of the reticular and relay nuclei of the cat thalamus

Relations between neurons of the reticular and specific relay nuclei of the thalamus were studied in cats immobilized with tubocurarine. Under the influence of stimulation of the reticular nucleus (RN) unit activity in the thalamic relay nuclei was found to be considerably modulated. Cases of the appearance of IPSPs (possibly of monosynaptic nature), evoked by stimulation of RN, in neurons of the ventroposterolateral nucleus (VPLN) and lateral geniculate body (LGB) are described. During simultaneous recording of unit activity in RN and VPLN or LGB by means of two electrodes interaction of several types was found: inhibition of discharges of VPLN or LGB neurons accompanied by excitation of RN neurons: alternation of excitation-inhibition in neuron pairs in RN and VPLN or RN and LGB during low-frequency afferent or cortical stimulation (in this case excitation of RN neurons is associated with inhibition of VPLN or LGB neurons), and strengthening of the discharge of VPLN or LGB neurons during excitation of RN neurons. The possibility of the existence both of direct monosynaptic inhibition of activity of VPLN or LGB relay neurons under the influence of excitation of RN neurons and of their inhibition by activation of hypothetical interneurons of the relay nuclei themselves is accepted.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 13, No. 1, pp. 24–31, January–February, 1981.