猫V1区不同外周整合细胞的经典感受野属性研究

以家猫为动物模型,采用细胞外记录的方法,测试了82个初级视皮层细胞的方位和方向调谐以及感受野大小.基于细胞的面积整合特性,区分出52个外周抑制型细胞和30个外周无抑制型细胞.所有被测细胞均存在强的方位选择性,而外周无抑制型细胞比抑制型细胞有更强的方位选择性.两类细胞的方向选择性没有显著性差异.外周抑制型细胞比外周无抑制型细胞有着更大的动作电位发放率.采用两种不同方法测量两类细胞的感受野范围,却产生了不同的结果:用最小反应区测量发现抑制型细胞的经典感受野更大,而用面积整合曲线测量时外周无抑制型细胞的感受野更大.     

Heterogeneity of neuron populations with complex receptive fields in the cat visual cortex

The selectivity of striate neurons with complex receptive fields to the orientation, direction, and velocity of movement of various stimuli was investigated in unanesthetized and uncurarized cats. On the basis of all characteristics obtained by the study of single-unit responses to a stationary flickering slit, a moving spot of light, and a moving oriented stimulus, four groups of complex neurons were distinguished. The characteristics of group I neurons indicate a mechanism of orientation selectivity in the organization of their receptive fields, group IV neurons have a mechanism of directional selectivity, and neurons of groups II and III possess both mechanisms. The existence of separate neuronal systems coding the orientation and direction of stimulus movement is suggested.V. Kapsukas State University, Vilnius. Translated from Neirofiziologiya, Vol. 11, No. 2, pp. 109–116, March–April, 1979.     

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.     

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.     

The visual cortex "on command": 2 modes of operation of neuronal mechanisms (a topical article)]

Orientation tuning (TO) of 152 single units in the 17th area of cat cerebral cortex was studied under different contrasts of the light bar flashed in RF (2.3, 10 and 100) and also under local and general anaesthesia (Sombrevin, 4 mg/kg or Nembutal, 35 mg/kg). An invariance of preferred orientation to contrast or anaesthesia was revealed in 40-50% of cells. Noninvariant cells shifted significantly their preferred orientation on 22-90 degrees. Invariant units more often have a simple RF, more sharp OT and preferred horizontal and vertical orientations. Under the low-contrast conditions or under general anaesthesia, noninvariant neurons shifted their preferred diagonal orientations. Different role of the two neuronal groups in detection of the reper (horizontal and vertical) and other orientations in a normal conditions (high-contrast, alert state) in comparison with their behavior under worse conditions (low-contrast, narcosis or sleep) is discussed.     

Rabbit visual cortical neurons with simple and complex receptive fields

Receptive fields of neurons of the rabbit visual cortex selective for stimulus orientation were investigated. These receptive fields were less well differentiated than those of the analogous neurons of the cat visual cortex (large in size and circular in shape). Two mechanisms of selectivity for stimulus orientation were observed: inhibition between on and off zones of the receptive field (sample type) and oriented lateral inhibition within the same zone of the receptive field (complex type). Lateral inhibition within the same zone of the receptive field also took place in unselective neurons; "complex" selective neurons differed from them in the orientation of this inhibition. A combination of both mechanisms was possible in the receptive field of the same neuron. It is suggested that both simple and complex receptive fields are derivatives of unselective receptive fields and that "complex" neurons are not the basis for a higher level of analysis of visual information than in "simple" neurons.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 10, No. 1, pp. 13–21, January–February, 1978.     

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.     

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.     

Relationship between Size Summation Properties,Contrast Sensitivity and Response Latency in the Dorsomedial and Middle Temporal Areas of the Primate Extrastriate Cortex

Analysis of the physiological properties of single neurons in visual cortex has demonstrated that both the extent of their receptive fields and the latency of their responses depend on stimulus contrast. Here, we explore the question of whether there are also systematic relationships between these response properties across different cells in a neuronal population. Single unit recordings were obtained from the middle temporal (MT) and dorsomedial (DM) extrastriate areas of anaesthetized marmoset monkeys. For each cell, spatial integration properties (length and width summation, as well as the presence of end- and side-inhibition within 15° of the receptive field centre) were determined using gratings of optimal direction of motion and spatial and temporal frequencies, at 60% contrast. Following this, contrast sensitivity was assessed using gratings of near-optimal length and width. In both areas, we found a relationship between spatial integration and contrast sensitivity properties: cells that summated over smaller areas of the visual field, and cells that displayed response inhibition at larger stimulus sizes, tended to show higher contrast sensitivity. In a sample of MT neurons, we found that cells showing longer latency responses also tended to summate over larger expanses of visual space in comparison with neurons that had shorter latencies. In addition, longer-latency neurons also tended to show less obvious surround inhibition. Interestingly, all of these effects were stronger and more consistent with respect to the selectivity for stimulus width and strength of side-inhibition than for length selectivity and end-inhibition. The results are partially consistent with a hierarchical model whereby more extensive receptive fields require convergence of information from larger pools of “feedforward” afferent neurons to reach near-optimal responses. They also suggest that a common gain normalization mechanism within MT and DM is involved, the spatial extent of which is more evident along the cell’s preferred axis of motion.     

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.     

Double orientation tuning of cat visual cortical neurons

Orientation tuning of 148 primary visual cortical neurons was studied in acute experiments on unanesthetized, curarized cats by analysis of their spike responses to flashes in a receptive field of a bar of light of optimal size. Orientation tuning of 88 neurons (59%) was found to be bimodal: Besides the principal preferred orientation there was a second, making an angle with the first. The second tuning maximum in some cases (64%) was exhibited only with a change in stimulus intensity or background brightness. Analysis of orientation tuning by the time-slice method, i.e., on the basis of individual cuts of the spike trace, showed double tuning to be present in 69% of cases only at certain moments after the beginning of stimulation. The results of analysis of the model showed that the double orientation tuning effect may be the result of the specific configuration of the receptive field, the use of a stimulus longer than the receptive field, the presence of a series of alternating excitatory and inhibitory zones in the receptive field, and also of end inhibitory zones on the narrow ends of the field. The unequal change in zones of the receptive fields in time explains the appearance of double orientation tuning in individual fragments of the spike trace. The functional role of double, "cross-wise" tuning in some primary visual cortical neurons and their role in the detection of the features of visual patterns are discussed.     

A neural network model on self-organizing emergence of simple-cell receptive field with orientation selectivity in visual cortex

In order to probe into the self-organizing emergence of simple cell orientation selectivity, we tried to construct a neural network model that consists of LGN neurons and simple cells in visual cortex and obeys the Hebbian learning rule. We investigated the neural coding and representation of simple cells to a natural image by means of this model. The results show that the structures of their receptive fields are determined by the preferred orientation selectivity of simple cells. However, they are also decided by the emergence of self-organization in the unsupervision learning process. This kind of orientation selectivity results from dynamic self-organization based on the interactions between LGN and cortex.     

A neural network model on self-organizing emergence of simple-cell receptive field with orientation selectivity in visual cortex

In order to probe into the self-organizing emergence of simple cell orientation selectivity, we tried to construct a neural network model that consists of LGN neurons and simple cells in visual cortex and obeys the Hebbian learning rule. We investigated the neural coding and representation of simple cells to a natural image by means of this model. The results show that the structures of their receptive fields are determined by the preferred orientation selectivity of simple cells. However, they are also decided by the emergence of self-organization in the unsupervision learning process. This kind of orientation selectivity results from dynamic self-organization based on the interactions between LGN and cortex.     

Modulation of synaptic transfer between retinal cones and horizontal cells by spatial contrast

We studied the influence of steady annular light on the kinetics and sensitivity of horizontal cell (HC) responses to modulation of the intensity of small concentric spots in the turtle retina. As shown by previous investigators, when the intensity of the annulus was equal to the mean spot intensity, spot response kinetics were the same as those for the modulation of spatially uniform light. Turning off the annulus attenuated dramatically high-frequency flicker sensitivity and enhanced somewhat low-frequency sensitivity. This phenomenon reflects a modulation of synaptic transfer between cones and second-order neurons that is mediated by cones, and it will be referred to as cone-mediated surround enhancement (CMSE). Our main results are as follows: (a) The change in test-spot response sensitivity and kinetics upon dimming a steady surrounding annulus is a consequence of the change in spatial contrast rather than change in overall light level. (b) Introduction of moderate contrast between the mean spot intensity and steady surrounding light intensity causes a marked change in spot response kinetics. (c) The dependence of spot response kinetics on surrounding light can be described by a phenomenological model in which the steady state gain and the time constant of one or two single-stage, low-pass filters increase with decreasing annular light intensity (d) The effect of surrounding light on spot responses of a given HC is not determined by change in the steady component of the membrane potential of that cell. (e) Light outside the receptive field of an HC can affect that cell's spot response kinetics. (f) In an expanding annulus experiment, the distance over which steady annular light affects spot response kinetics varies among HCs and can be quite different even between two cells with closely matched receptive field sizes. (g) The degree of CMSE is correlated with HC receptive field size. This correlation suggests that part of the enhancement mechanism is located in the HC. Taken together, our results suggest the involvement of the inner retina in CMSE.     

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.     

Reorganization of Receptive Fields of Cortical Field 21a Neurons and Formation of Responses to Moving Visual Stimuli

Using extracellular recording of spike activity from single neurons of field 21a of the cat neocortex, we examined in detail the spatial organization of receptive fields (RFs) of such cells after conditions of presentation of an immobile blinking light spot (a static RF) and moving visual stimuli (dynamic RFs). As was shown, the excitability of different RF subfields of a group of neurons possessing homogeneous on–off organization of the static RF changes significantly depended on the contrast, shape, dimension, orientation, and direction of movement of the applied mobile visual stimulus. This is manifested in changes in the number of discharge centers and shifts of their spatial localization. A hypothesis on the possible role of synchronous activation of the neurons neighboring the cell under study in the formation of an additional neuronal mechanism providing specialization of neuronal responses is proposed.     

Laminar distribution of neurons with different types of receptive fields in the rabbit visual cortex

Investigation of receptive fields of 232 primary visual cortical neurons in rabbits by the use of shaped visual stimuli showed that 21.1% are unselective for stimulus orientation, and 34.1% have simple, 16.4% complex, and 18.5% hypercomplex receptive fields, and 9.9% have other types. Neurons with different types of receptive fields also differed in spontaneous activity, selectivity for rate of stimulus movement, and acuteness of orientational selectivity. Neurons not selective to orientation were found more frequently in layer IV than in other layers, and very rarely in layer VI. Cells with simple receptive fields were numerous in all layers but predominated in layer VI. Neurons with complex receptive fields were rare in layer IV and more numerous in layers V and VI. Neurons with hypercomplex receptive fields were found frequently in layers II + III and IV, rarely in layers V and VI. Spontaneous unit activity in layer II + III was lowest on average, and highest in layer V. Acuteness or orientational selectivity of neurons with simple and complex receptive fields in layers II + III and V significantly exceeded the analogous parameter in layers IV and VI.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 17, No. 1, pp. 19–27, January–February, 1985.     

Standardization and invariance of neuronal detector tuning in an orientation column of the cat visual cortex

Variability of orientation tuning of primary visual cortical neurons in single orientation columns and the degree of its stability during changes in the level of contrast between stimulus and background were investigated in acute experiments on immobilized cats. Several types of orientation columns were found, with the following properties of orientation tuning of their neurons: relatively high, standard, and stable; varying widely from neuron to neuron; invariant regardless of the level of contrast; noninvariant; mixed (invariant-noninvariant). Properties of standardization-nonstandardization, on the one hand, and invariance-noninvariance of the neurons, on the other hand, may be combined differently in a column. Differences of orientation tuning within a column were observed most frequently in neurons of the upper and lower layers of the cortex. Possible differences in the functional role of the variance of orientation columns described and in the mechanisms of formation of the detector properties of their neurons are discussed.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 17, No. 2, pp. 175–182, March–April, 1985.     

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.     

Double orientation tuning of neurons in the primary cortex of cat at different levels of alertness

Orientation tuning (OT) of 225 cat neurons of the primary visual cortex (field 17) to the flashing of a light bar in the discharge centers of their receptive field (RFs) were investigated. It was found that 43% of the cells investigated were monomodally tuned, i.e., were primarily detecting horizontal and vertical orientations. The remaining 57% of the neurons exhibited double OT, i.e, exhibited, in addition to a main preferred orientation (PO), an additional preferred orientation (aPO) at a right or acute angle to the main orientation (the mean angle between the two OT maxima equalled 71.4±2.4°). In bimodal cells, the additional maximum of OT was comparable in magnitude to the main maximum (averaging 0.7±0.03 of the PO) in half the cases. The orientational properties of the main and additional maxima were almost indistinguishable. Under light or moderate anesthesia, approximately half the neurons with double OT became monomodal; at the same time, a small fraction of monomodal cells (12%) manifested double OT. Under anesthesia, the angle between two the preferred orientations decreased, while the ratio of amplitude characteristics remained unchanged. Monomodal neurons frequently exhibited simple RFs and OTs unaffected by anesthesia. Neurons with double OT, on the other hand, exhibited simple and complex types of RFs just as often and their OT changed under the influence of anesthesia. It is suggested that neurons with double OT can function as detectors of angles and angles of intersecting lines; such angles, together with line orientation, are important attributes of images. In contrast, monomodal neurons may provide a benchmark for a stable reference system of orientation coordinates. The interaction of the two neuronal systems mentioned may allow effective analysis of image attributes at the level of the primary visual cortex.Institute of Higher Nervous Activity and Neurophysiology, Russian Academy of Sciences Moscow. Translated from Neirofiziologiya, Vol. 24, No. 3, pp. 260–269, May–June, 1992.