A binocular model for the simple cell

We authors propose a mathematical model for simple cell binocular response. It comprises two Gabor-type receptive fields (RF) having the same RF center, preferred spatial frequency, and preferred orientation. The model integrates the equally weighted signals from both eyes and performs a threshold operation. Poggio and Fischer (1977) classified binocular disparity cells in the striate cortex into four groups: tuned excitatory (TE), tuned inhibitory (TI), near, and far cells. They also found that most of the TE cells are ocularly balanced and that the other three types are usually unbalanced. This model can imitate these four types of disparity sensitivities and their ocular dominance tendency. We perform model fittings to Poggio's data using the “simulated annealing” method and discuss parameter dependence of the model's response. The model can also respond with exceptional disparity sensitivity: i.e., flat type, alternating type, and intermediate type.     

Receptive field microstructure and dendritic geometry of retinal ganglion cells

We studied the fine spatial structure of the receptive fields of retinal ganglion cells and its relationship to the dendritic geometry of these cells. Cells from which recordings had been made were microinjected with Lucifer yellow, so that responses generated at precise locations within the receptive field center could be directly compared with that cell's dendritic structure. While many cells with small receptive fields had domeshaped sensitivity profiles, the majority of large receptive fields were composed of multiple regions of high sensitivity. The density of dendritic branches at any one location did not predict the regions of high sensitivity. Instead, the interactions between a ganglion cell's dendritic tree and the local mosaic of bipolar cell axons seem to define the fine structure of the receptive field center.     

A reaction-diffusion model to capture disparity selectivity in primary visual cortex

Decades of experimental studies are available on disparity selective cells in visual cortex of macaque and cat. Recently, local disparity map for iso-orientation sites for near-vertical edge preference is reported in area 18 of cat visual cortex. No experiment is yet reported on complete disparity map in V1. Disparity map for layer IV in V1 can provide insight into how disparity selective complex cell receptive field is organized from simple cell subunits. Though substantial amounts of experimental data on disparity selective cells is available, no model on receptive field development of such cells or disparity map development exists in literature. We model disparity selectivity in layer IV of cat V1 using a reaction-diffusion two-eye paradigm. In this model, the wiring between LGN and cortical layer IV is determined by resource an LGN cell has for supporting connections to cortical cells and competition for target space in layer IV. While competing for target space, the same type of LGN cells, irrespective of whether it belongs to left-eye-specific or right-eye-specific LGN layer, cooperate with each other while trying to push off the other type. Our model captures realistic 2D disparity selective simple cell receptive fields, their response properties and disparity map along with orientation and ocular dominance maps. There is lack of correlation between ocular dominance and disparity selectivity at the cell population level. At the map level, disparity selectivity topography is not random but weakly clustered for similar preferred disparities. This is similar to the experimental result reported for macaque. The details of weakly clustered disparity selectivity map in V1 indicate two types of complex cell receptive field organization.     

Two-dimensional substructure of stereo and motion interactions in macaque visual cortex

The analysis of object motion and stereoscopic depth are important tasks that are begun at early stages of the primate visual system. Using sparse white noise, we mapped the receptive field substructure of motion and disparity interactions in neurons in V1 and MT of alert monkeys. Interactions in both regions revealed subunits similar in structure to V1 simple cells. For both motion and stereo, the scale and shape of the receptive field substructure could be predicted from conventional tuning for bars or dot-field stimuli, indicating that the small-scale interactions were repeated across the receptive fields. We also found neurons in V1 and in MT that were tuned to combinations of spatial and temporal binocular disparities, suggesting a possible neural substrate for the perceptual Pulfrich phenomenon. Our observations constrain computational and developmental models of motion-stereo integration.     

Parallel processing of binocular disparity in the cat's retinogeniculocortical pathways

In the cat, parallel streams of information processing have been traced from X-, Y- and W-type retinal ganglion cells to visual cortical areas 17 (X-, Y- and W-type), 18 (Y-type) and 19 (W-type). In the present study we have examined, in the anaesthetized and paralysed adult cat, the role played by X-, Y- and W-subsystems, projecting to areas 17 and 19, in the processing of binocular retinal disparity. The tapetal reflection technique was used to monitor residual eye movements and to provide a map, for each eye, of the retinal blood vessels which could later be compared with retinal wholemounts stained with cresyl violet to reveal the area centralis. The receptive-field disparities of cells recorded from areas 17 and 19 were compared with each other and with reference to the visual axes defined by the area centralis of each eye. Cells of area 19 (receiving W-type input) had horizontal receptive-field disparities that were significantly more divergent than those of the cells in area 17 and 17-18 'border region'. Referred to the area centralis, the mean horizontal receptive-field disparity in area 19 was -0.5 degrees (+/- 0.8 degrees). The mean horizontal receptive-field disparity of area 17 (receiving X-, Y- and W-type input) was convergent with respect to the visual axis at +2 degrees (+/- 0.5 degrees). Finally, the mean horizontal receptive-field disparity of the cells in the 17-18 border region (which receive mainly Y-type input) was even more convergent (2.6 degrees +/- 1.5 degrees) than that of area 17. Binocular interactions of cortical neurons were tested with the Risley biprism technique. Area 19 cells had maximal responses to binocular stimulation when the receptive-field disparities were either close to zero or slightly divergent. In contrast, area 17 cells tended to respond optimally to disparities that were either slightly or strongly convergent. At the level of the lateral geniculate nucleus there were significant differences between the receptive-field disparities inferred from the comparison of receptive-field positions of adjacent neurons recorded on either side of the border between the A and A1 geniculate laminae and those inferred from a similar comparison at the C1-C2 border. The mean horizontal disparities inferred from the interlaminar comparison at the A-A1 border were +2.1 degrees (+/- 0.3 degrees); those inferred from the interlaminar comparison at the C1-C2 border -0.2 (+/- 0.2 degrees) were more divergent.(ABSTRACT TRUNCATED AT 400 WORDS)     

Receptive fields of disparity-tuned simple cells in macaque V1

Binocular simple cells in primary visual cortex (V1) are the first cells along the mammalian visual pathway to receive input from both eyes. Two models of how binocular simple cells could extract disparity information have been put forward. The phase-shift model proposes that the receptive fields in the two eyes have different subunit organizations, while the position-shift model proposes that they have different overall locations. In five fixating macaque monkeys, we recorded from 30 disparity-tuned simple cells that showed selectivity to the disparity in a random dot stereogram. High-resolution maps of the left and right eye receptive fields indicated that both phase and position shifts were common. Single cells usually showed a combination of the two, and the optimum disparity was best correlated with the sum of receptive field phase and position shift.     

Small-field, binocular neurons in the superficial layers of the frog optic tectum

Binocular neurons with receptive fields about 5 degrees across were recorded just beneath the pia. Most of them responded to dark stimuli in the lower half of their receptive field and to light stimuli above. There was almost no vertical disparity between the left and right fields and the modal value of the horizontal disparity of the population of cells was 1.7 degrees. Because frogs do not verge their eyes it is possible to calculate at what distance the receptive fields through the two eyes are superimposed. This calculation suggests that the neurons are tuned to detect features in the external world about 50 cm away. This is too far for the neurons to be involved in the frog's everyday distance vision. It is more likely that they are concerned with assessing the vertical position of a horizontal surface.     

猫视皮层神经元对光栅反应最优方位在视网膜上分布特性的研究

用金属微电极记录了114个猫皮层17、18区细胞对不同方位光栅图象刺激的反应。细胞最优方位与其感受野中心在视网膜的位置间有系统性关系,即最优方位总是倾向于垂直于各感受野中心与视网膜中心区(area centralis)的连线。这一规律对在18区或17、18区全体记录到的细胞而言,有统计意义。 在17、18区内,仅对于感受野位于视网膜离心度(eccentricity)大于9°视角的细胞、具有较窄感受野(宽度小于2.5°)的细胞以及感受野处于视网膜垂直经线附近的细胞,上述规律才有统计意义,而对感受野离心度小于9°的细胞、感受野宽度大于2.5°的细胞以及感受野在倾斜经线附近的细胞,上述规律不明显。     

Two-dimensional substructure of MT receptive fields.

Neurons at progressively higher levels of the visual system have progressively larger, more complicated receptive fields, presumably constructed from simpler antecedent receptive fields. To study this hierarchical organization, we used sparse white noise to map receptive-field substructure (second order Wiener-like kernels) in an extrastriate motion processing area (MT) of alert monkeys. The maps revealed a clear substructure, on a spatial scale comparable to the receptive fields of the V1 inputs. There were both facilitatory and suppressive interactions that differed in spatial organization and time course. Directional interactions were remarkably precise over a very small spatial range, and reversed when successive stimuli reversed contrast--a neural correlate of "reverse phi" motion perception. The maps of some cells had an unexpected, curved shape, which challenges existing models for direction selectivity.     

Border ownership from intracortical interactions in visual area v2

A border between two image regions normally belongs to only one of the regions; determining which one it belongs to is essential for surface perception and figure-ground segmentation. Border ownership is signaled by a class of V2 neurons, even though its value depends on information coming from well outside their classical receptive fields. I use a model of V2 to show that this visual area is able to generate the ownership signal by itself, without requiring any top-down mechanism or external explicit labels for figures, T junctions, or corners. In the model, neurons have spatially local classical receptive fields, are tuned to orientation, and receive information (from V1) about the location and orientation of borders. Border ownership signals that model physiological observations arise through finite range, intraareal interactions. Additional effects from surface features and attention are discussed. The model licenses testable predictions.     

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.     

Stereoscopic mechanisms: binocular responses of the striate cells of cats to moving light and dark bars

New knowledge concerning the internal structure and response properties of the receptive fields of striate cells calls for a fresh appraisal of their binocular interactions in the interest of a better understanding of the neural mechanisms underlying binocular depth discrimination. Binocular position-disparity response profiles were recorded from 71 simple and B-cells in response to moving light and dark bars. Predominantly excitatory (PE) cells (N = 48) had disparity response profiles that were spatially closely similar to their respective monocular responses. In addition, the centrally located excitatory subregions were flanked on one or both sides by non-specific inhibitory regions. PE cells with a preferred stimulus orientation within 30 degrees of the vertical (N = 17) showed binocular facilitations with maximal values that were always more than twice (mean 3.3) the sum of the two monocular responses to the same stimuli and generally greater than the facilitations shown by cells with orientations more than 30 degrees from the vertical (N = 29; mean 2.2 times the sum of the respective monocular responses). The strength of the binocular facilitation depended on the stimulus contrast, the facilitation decreasing with increasing contrast. The receptive-field disparity distribution of the 31 PE cells capable of making significant horizontal disparity discriminations has standard deviations of 0.37 degrees and 0.40 degrees, respectively. Predominantly inhibitory cells (PI) (N = 23) showed two basic types of disparity response profile: symmetric (N = 17) and asymmetric (N = 6). Uncertainty regarding the precise location of the binocular fixation point in the anaesthetized and paralysed preparation made it difficult to categorize PI cells adequately.     

Structural differences of spatial-frequency filters in the cat visual cortex

Sensitivity profiles of receptive fields (RF) of the cat's visual cortex along the width and length of the field were determined. They served as criteria for determining structural differences of the earlier discovered types of spatial-frequency filters. It has been shown that RF with spatial-frequency selectivity in a narrow range of orientations close to the preferable one, have the "one-dimensional" sensitivity profiles in the same range. For RF showing spatial-frequency selectivity along the RF length and width two-dimensional sensitivity profiles along these RF dimensions were revealed. A group of fields was found whose pattern of sensitivity profiles depended on the geometrical properties of images which stipulated the functional changes o RF filtering properties. Possible role of different types of spatial-frequency filters in image processing is discussed.     

Stimulus-entrained oscillatory activity propagates as waves from area 18 to 17 in cat visual cortex

Previous studies in cat visual cortex reported that area 18 can actively drive neurons in area 17 through cortico-cortical projections. However, the dynamics of such cortico-cortical interaction remains unclear. Here we used multielectrode arrays to examine the spatiotemporal pattern of neuronal activity in cat visual cortex across the 17/18 border. We found that full-field contrast reversal gratings evoked oscillatory wave activity propagating from area 18 to 17. The wave direction was independent of the grating orientation, and could not be accounted for by the spatial distribution of receptive field latencies, suggesting that the waves are largely mediated by intrinsic connections in the cortex. Different from the evoked waves, spontaneous waves propagated along both directions across the 17/18 border. Together, our results suggest that visual stimulation may enhance the flow of information from area 18 to 17.     

Temporal pattern discrimination within the receptive field of cat retinal ganglion cells

ON-center and OFF-center receptive fields of cat retinal ganglion cells can be divided into two categories: sensitive (type N) and insensitive (type L) to three statistical temporal visual stimuli with different second order statistics but identical first order statistics (Tsukada et al. 1982). The temporal pattern sensitivity of type N response is closely related to the nonlinear stage of Y cells depending on the interaction between center and surround mechanism. The temporal pattern sensitivity of type N responses has a spatial profile within the receptive field; it is highly sensitive in the center region of the receptive field and less sensitive toward the field periphery. The temporal pattern sensitivity in the center region of the receptive field to statistical properties (irregular or regular) of a surrounding flash annulus shows modulation like a switching element: when the surrounding area is stimulated by a more regular flash stimulus with normal distribution of inter-stimulus intervals the system is sensitive (switching on) to the temporal pattern, while a change to an irregular one with an exponential distribution makes it insensitive (switching off) to the temporal pattern.     

Are receptive fields of the visual cortex detectors or spatial frequency filters?

Besides its principal maximum, the spatial frequency characteristic curve of the complex visual cortical receptive field of curarized cats also has additional maxima and also negative regions, as predicted by the theory of piecewise Fourier analysis. Comparison of responses of the complex receptive field to sinusoidal gratings completely and incompletely contained in the field and comparison of responses to sinusoidal and square-wave gratings indicate that the receptive field, as a spatial frequency filter, has linear properties. The response of the complex receptive field rises with an increase in the number of periods of the sinusoidal grating. Several periods of optimal frequency match the complex field. Receptive fields tuned to a broad band of spatial frequencies were found in neuron columns. The results confirm the view that complex receptive fields are spatial frequency filters and not detectors.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 11, No. 5, pp. 403–411, September–October, 1979.     

Development of spatiotemporal receptive fields of simple cells: I. Model formulation

A model for the development of spatiotemporal receptive fields of simple cells in the visual cortex is proposed. The model is based on the 1990 hypothesis of Saul and Humphrey that the convergence of four types of input onto a cortical cell, viz. non-lagged ON and OFF inputs and lagged ON and OFF inputs, underlies the spatial and temporal structure of the receptive fields. It therefore explains both orientation and direction selectivity of simple cells. The response properties of the four types of input are described by the product of linear spatial and temporal response functions. Extending the 1994 model of one of the authors (K.D. Miller), we describe the development of spatiotemporal receptive fields as a Hebbian learning process taking into account not only spatial but also temporal correlations between the different inputs. We derive the correlation functions that drive the development both for the period before and after eye-opening and demonstrate how the joint development of orientation and direction selectivity can be understood in the framework of correlation-based learning. Our investigation is split into two parts that are presented in two papers. In the first, the model for the response properties and for the development of direction-selective receptive fields is presented. In the second paper we present simulation results that are compared with experimental data, and also provide a first analysis of our model. Received: 18 June 1997 / Accepted: 16 September 1997     

Independent component filters of natural images compared with simple cells in primary visual cortex.

Properties of the receptive fields of simple cells in macaque cortex were compared with properties of independent component filters generated by independent component analysis (ICA) on a large set of natural images. Histograms of spatial frequency bandwidth, orientation tuning bandwidth, aspect ratio and length of the receptive fields match well. This indicates that simple cells are well tuned to the expected statistics of natural stimuli. There is no match, however, in calculated and measured distributions for the peak of the spatial frequency response: the filters produced by ICA do not vary their spatial scale as much as simple cells do, but are fixed to scales close to the finest ones allowed by the sampling lattice. Possible ways to resolve this discrepancy are discussed.     

Independent component analysis of natural image sequences yields spatio-temporal filters similar to simple cells in primary visual cortex.

Simple cells in the primary visual cortex process incoming visual information with receptive fields localized in space and time, bandpass in spatial and temporal frequency, tuned in orientation, and commonly selective for the direction of movement. It is shown that performing independent component analysis (ICA) on video sequences of natural scenes produces results with qualitatively similar spatio-temporal properties. Whereas the independent components of video resemble moving edges or bars, the independent component filters, i.e. the analogues of receptive fields, resemble moving sinusoids windowed by steady Gaussian envelopes. Contrary to earlier ICA results on static images, which gave only filters at the finest possible spatial scale, the spatio-temporal analysis yields filters at a range of spatial and temporal scales. Filters centred at low spatial frequencies are generally tuned to faster movement than those at high spatial frequencies.