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.     

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.     

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.     

The light sensitivity of the human visual system depends on the direction of view

The near-stable North-South orientation of the natural geomagnetic field provides an ideal basis for navigation. Sailors have used it since ancient times, animals for much longer. Various mechanisms have developed for this purpose. Experiments have pointed to a connection between orientation in the geomagnetic field and light perception. Such observations are supported by theoretical considerations. The underlying interaction should also modulate the light sensitivity of the visual system. Recently we demonstrated the effect of an oscillating field. Here we report the existence of a weak influence of the static field on visual sensitivity in man. By comparison with control experiments, if the directions of view line and field vector coincide the perception threshold of a light stimulus is slightly but significantly increased. This significance is lost if the view line deviates by 10 degrees from the field direction.     

Neural coding of 3D features of objects for hand action in the parietal cortex of the monkey.

In our previous studies of hand manipulation task-related neurons, we found many neurons of the parietal association cortex which responded to the sight of three-dimensional (3D) objects. Most of the task-related neurons in the AIP area (the lateral bank of the anterior intraparietal sulcus) were visually responsive and half of them responded to objects for manipulation. Most of these neurons were selective for the 3D features of the objects. More recently, we have found binocular visual neurons in the lateral bank of the caudal intraparietal sulcus (c-IPS area) that preferentially respond to a luminous bar or place at a particular orientation in space. We studied the responses of axis-orientation selective (AOS) neurons and surface-orientation selective (SOS) neurons in this area with stimuli presented on a 3D computer graphics display. The AOS neurons showed a stronger response to elongated stimuli and showed tuning to the orientation of the longitudinal axis. Many of them preferred a tilted stimulus in depth and appeared to be sensitive to orientation disparity and/or width disparity. The SOS neurons showed a stronger response to a flat than to an elongated stimulus and showed tuning to the 3D orientation of the surface. Their responses increased with the width or length of the stimulus. A considerable number of SOS neurons responded to a square in a random dot stereogram and were tuned to orientation in depth, suggesting their sensitivity to the gradient of disparity. We also found several SOS neurons that responded to a square with tilted or slanted contours, suggesting their sensitivity to orientation disparity and/or width disparity. Area c-IPS is likely to send visual signals of the 3D features of an object to area AIP for the visual guidance of hand actions.     

Functional characteristics of visual cortical neurons during local photic stimulation of their receptive fields in cats

A group of functional characteristics of 103 neurons in visual cortical area 17 was investigated in acute experiments on curarized, light-adapted cats during a change in various parameters of the local photic stimuli. The average threshold sensitivity of the neuron population was 32 dB (0.052 nit), the sharpness of orientation tuning was 37°, the critical summation time was 57 msec, and the reactivity recovery time 190 msec. Photic sensitivity was lower during light adaptation than during dark adaptation, orientation selectivity of the neurons was increased, temporal summation was lengthened, and the time required by the neuron to recovery from after-inhibition was shortened. Several properties of the cortical neurons depended on the accentricity of their receptive fields: Cells with centrally localized receptive fields on average had lower thresholds and shorter summation time and they recovered their reactivity more quickly; their activity was of a higher frequency and they more often generated short phasic discharges than neurons with receptive fields in the peripheral part of the visual field. The mechanisms responsible for changes in the properties of neurons in the central and peripheral visual channels during dark and light adaptation are discussed. The presence of several inhibitory subsystems in the cortex regulating unit activity in the primary visual projection area is postulated.     

Coding of stimulus movement parameters in cat visual system

The principal component analysis of matrices composed of spike numbers generated by visual neurons of cats in response to motion of simple and complex stimuli revealed vector encoding. Responses of detectors of moving dot direction and detectors of oblique line orientation are encoded independently in V1 and V2 cortices by excitation of two cardinal neurons. Each pair of these neurons generates sine and cosine functions. Responses of detectors in the association cortex selective to specific orientation of moving stripes depend on the activity of four cardinal neurons which sum up the excitation incoming from the direction and orientation channels.     

End-stopping and the aperture problem: two-dimensional motion signals in macaque V1

Our perception of fine visual detail relies on small receptive fields at early stages of visual processing. However, small receptive fields tend to confound the orientation and velocity of moving edges, leading to ambiguous or inaccurate motion measurements (the aperture problem). Thus, it is often assumed that neurons in primary visual cortex (V1) carry only ambiguous motion information. Here we show that a subpopulation of V1 neurons is capable of signaling motion direction in a manner that is independent of contour orientation. Specifically, end-stopped V1 neurons obtain accurate motion measurements by responding only to the endpoints of long contours, a strategy which renders them largely immune to the aperture problem. Furthermore, the time course of end-stopping is similar to the time course of motion integration by MT neurons. These results suggest that cortical neurons might represent object motion by responding selectively to two-dimensional discontinuities in the visual scene.     

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.     

The tuning of striate neurons to cross-like figures during local blockade of intracortical inhibition]

Many neurons in the cat primary visual cortex reveal sensitivity to cruciform and corner figures as well as to local orientation discontinuities. We investigated this sensitivity in 85 V1 neurons before and after the local blockade of GABA(A)ergic inhibition by microiontophoretic application of bicuculline and observed two opposite effects: a decrease or disappearance of sensitivity to crosses in about half of the neurons and its increase or appearance in about one third of the units. The results indicate substantial and drastically different contribution of intracortical inhibition to sensitivity to line-crossings in the visual cortical neurons. Some possible mechanisms of this difference are discussed.     

Selective sensitivity of the cat striate neurons to cruciform and angular figures of various orientations

Selectivity and invariability of tuning were studied in 51 neurons of the primary visual cortex (area 17); cruciform and angular figures (CF and AF, respectively) of different configurations and orientations were presented in their receptive fields. Twenty-three neurons, or 45% of the studied cells, demonstrated selective sensitivity to these figures. Their responses considerably (2.38±0.36 times, on average) increased, as compared with those evoked by presentation of a single bar of preferred orientation. In the examined group, 2 cells demonstrated sensitivity both to the CF and AF. A wide range of detector properties related to the CF and AF analysis was found in the analyzed neuronal population. Detectors of configuration of these figures are described. Selective sensitivity to the angle between branches of these figures was observed in 17 neurons, and responses of 2 neurons among them showed invariability to orientation of these figures. Four cells were selective for orientation and were insensitive to configuration, and 4 other cells showed no specific sensitivity to either of these properties, but were sensitive to the appearance of a CF itself in their receptive field (these cells were regarded as invariant detectors of crossing nodes). Data inconsistent with the hierarchic principle of detection of the above properties are presented. Possible mechanisms and functional significance of selective sensitivity of striate neurons to the CF and AF are discussed.Neirofiziologiya/Neurophysiology, Vol. 27, No. 5/6, pp. 403–412, September–December, 1995.     

Coding the presence of visual objects in a recurrent neural network of visual cortex

Before we can recognize a visual object, our visual system has to segregate it from its background. This requires a fast mechanism for establishing the presence and location of objects independently of their identity. Recently, border-ownership neurons were recorded in monkey visual cortex which might be involved in this task [Zhou, H., Friedmann, H., von der Heydt, R., 2000. Coding of border ownership in monkey visual cortex. J. Neurosci. 20 (17), 6594-6611]. In order to explain the basic mechanisms required for fast coding of object presence, we have developed a neural network model of visual cortex consisting of three stages. Feed-forward and lateral connections support coding of Gestalt properties, including similarity, good continuation, and convexity. Neurons of the highest area respond to the presence of an object and encode its position, invariant of its form. Feedback connections to the lowest area facilitate orientation detectors activated by contours belonging to potential objects, and thus generate the experimentally observed border-ownership property. This feedback control acts fast and significantly improves the figure-ground segregation required for the consecutive task of object recognition.     

Functional characteristics of visual cortical neurons in squirrels

Depending on the organization of their receptive fields and character of their responses to shaped visual stimuli the following main groups of visual cortical neurons were distinguished in the squirrelSciurus vulgaris: nonselective for direction of movement and orientation of stimuli (14%); selective for direction of movement (30%) and selective for line orientation (49%); 7% of neurons were not classified. Cells selective for direction of movement and some nonselective cells exhibited specific sensitivity to high speeds of stimulus movement (optimal velocities of the order of hundreds of degrees per second). Neurons selective for line orientation differed in the degree of overlapping of their on- and off-zones; they could include analogs of simple and complex neurons.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Moscow. Translated from Neirofiziologiya, Vol. 13, No. 2, pp. 125–231, March–April, 1981.     

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.     

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.