The Y cell visual pathway implements a demodulating nonlinearity

Neural encoding of sensory signals involves both linear and nonlinear processes. Determining which nonlinear operations are implemented by neural systems is crucial to understanding sensory processing. Here, we ask if demodulation, the process used to decode AM radio signals, describes how Y cells in the cat LGN nonlinearly encode the visual scene. In response to visual AM signals across?a wide range of carrier frequencies, Y cells were found to transmit a demodulated signal, with the firing rate of single-units fluctuating at the envelope frequency but not the carrier frequency. A comparison of temporal frequency tuning properties between LGN Y cells and neurons in two primary cortical areas suggests that Y cells initiate a distinct pathway that carries a demodulated representation of the visual scene to cortex. The nonlinear signal processing carried out by the Y cell pathway simplifies the neural representation of complex visual features and allows high spatiotemporal frequencies to drive cortical responses.     

一种新的视觉皮层简单细胞工作机制的模型

提出了一种基于独立元分析（ICA）的视觉皮层简单细胞工作机制的模型。用Gabor函数逼近对自然图像进行ICA而获得的基函数,揭示了ICA基函数与视觉皮层简单细胞感受野反应间存在内在的关系。并对水平条纹的图像进行ICA,模拟在特殊视觉环境下生长的幼年动物的视觉皮层发育过程,证实了1970年Blakemore和Cooper在幼猫上的实验结果。从而说明ICA可以模拟动物的视觉皮层简单细胞工作过程。     

Relaxation network for Gabor image decomposition

The so-called simple cells in layer IV of feline primary visual cortex have been shown to have Gabor function spatial receptive field profiles (RFP's). Since Gabor functions are not mutually orthogonal, the decomposition of an image into Gabor function coefficients is usually performed by minimising some measure of the error between the original image and that reconstructed from the coefficients. A cortical relaxation model is proposed which performs this minimisation implicitly, and is used to examine the biological relevance and feasibility of reconstruction error minimisation.     

The uncertainty principal in vision

During research into the relationship between the bandpass width of the spatial frequency characteristic (F) and the size (D) of the receptive field of the cat striate cortex it was found that F·D=1.2. Inaccuracy in spatial signal representation is thus related by an uncertainty principle to inaccuracy of frequency representation within the system of elements involved in information processing in the cat association cortex. The magnitude of the constant shows that the distribution functions of elements are sinusoids or cosinusoids modulated by square-wave impulses rather than Gabor elements. The magnitude of the constant and the fact that an uncertainty principle operates would support the hypothesis that the receptive fields of the visual cortex constitute a quasilinear basis.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 18, No. 3, pp. 307–312, May–June, 1986.     

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)     

A ring model for spatiotemporal properties of simple cells in the visual cortex

 A neural model is proposed for the spatiotemporal properties of simple cells in the visual cortex. In the model, several cortical cells are arranged on a ring, with mutual excitatory or inhibitory connections. The cells also receive excitatory inputs either from lagged and nonlagged cells of the lateral geniculate nucleus in one setting or from nonlagged cells in the other. Computer simulation shows that the cortical cells have spatiotemporally inseparable receptive fields in the former setting and separable fields in the latter; spatial profiles at a given time in the spatiotemporal fields are described with a Gabor function whose phase parameter varies regularly from 0 to 2π with rotation along the ring; the inseparable cells have directional selectivity as observed physiologically. Received: 13 November 1995 / Accepted in revised form: 1 July 1997     

The roles of polarity and symmetry in the perceptual grouping of contour fragments

We describe two experiments that investigate the roles of polarity and symmetry in the perceptual grouping of contour fragments. Observers viewed, for one second on each presentation, arrays of oriented, spatial-frequency band-pass, elements, in which a subset of the elements was aligned along a twisting curve. In each of five conditions we measured observers' ability to detect aligned combinations of even- and odd-symmetric elements, of the same and different polarities, against a background of 'noise' elements. As with previous experiments we found that the 'path' could be reliably detected, even when the elements of the path were oriented at angles of up to +/- 60 deg relative to each other. Detection of the path was still possible when the polarity of path elements alternated. However, the probability of detection of the path was raised significantly when the path elements were all of the same polarity. Perceptual grouping of even-symmetric elements was no different to perceptual grouping of odd-symmetric elements. The results provide evidence, that in achieving integration of contour fragments, the visual system uses a process that is to some degree phase selective. We use the results to describe how the visual system may resolve natural contours when they occur against backgrounds that vary over a wide range of intensities. The data presented here have been published in conference-abstract form (Hayes et al., 1993; Field et al., 1997).     

A functional model of the wiring of the simple cells of visual cortex

A computer model of the simple cells in the mammalian visual cortex was constructed. The model cells received inputs from a great number of isopolar centre/surround cells assumed to be located in the lateral geniculate nucleus (LGN). The distribution of input to the model simple cells was either inhibitory/excitatory or inhibitory/excitatory/inhibitory. Such arrangements produced receptive fields containing four or five consecutively antagonistic subfields. Responses produced by the model cells to different types of stimuli (periodical as well as nonperiodical) were obtained and compared to responses of living cells reported from various laboratories under comparable stimulus conditions. In all the situations tested, the responses of the model cells corresponded qualitatively very well to those of living cells. It was seen that the same wiring mechanism was able to account for orientation selectivity, spatial frequency filtering, various phase relationships between stimulus and response, subfield orientational selectivity, and slight end-inhibition. Furthermore, the receptive fields of the model simple cells closely resemble Gabor functions.     

Directional inhibition: a new slant on an old question

In this issue of Neuron, Priebe and Ferster describe the direction selectivity and spatiotemporal organization of excitatory and inhibitory inputs to direction-selective simple cells in cat visual cortex. Their most surprising finding is that inhibition shows the same preferred direction as excitation.     

The visual cortical association field: A Gestalt concept or a psychophysiological entity?

The receptive field of a visual neurone is classically defined as the region of space (or retina) where a visual stimulus evokes a change in its firing activity. Intracellular recordings in cat area 17 show that the visually evoked synaptic integration field extends over a much larger area than that established on the basis of spike activity. Synaptic depolarizing (dominant excitation) responses decrease in strength for stimuli that are flashed at increasing distances away from the centre of the discharge field, while their onset latency increases. A detailed spatio-temporal analysis of these electrophysiological data shows that subthreshold synaptic responses observed in the 'silent' surround of cortical receptive fields result from the intracortical spread of activation waves carried by slowly conducting horizontal axons within primary visual cortex. They also predict that a perceptual facilitation may occur when feedforward activation produced by the motion signal in the retina travels in phase in the primary visual cortex with the visually induced spread of horizontal activation. A psychophysical correlate has been obtained in humans, showing that apparent motion produced by a sequence of co-linear Gabor patches, known to preferentially activate V1 orientation selective cells, are perceived by human observers as much faster than non co-linear sequences of the same physical speed.     

Non-linearities in cortical simple cells and the possible detection of zero crossings

A theory of early visual information processing proposed by Marr and co-workers suggests that simple cortical cells may be involved in the detection of zero crossing in the retinal output. We have tested this theory by using pairs of adjacent edges (staircases stimuli) and recording from edge-specific simple cells in cat striate cortex. The zero crossing hypothesis gives rise for such stimuli to non-obvious predictions that were generally confirmed by the experiment.     

The relationship between the Gabor elementary function and a stochastic model of the inter-spike interval distribution in the responses of visual cortex neurons

In a previously reported study (Berger et al. 1990) we analyzed distributions of interspike intervals recorded extracellularly from cat visual cortex under four stimulus conditions. Stimuli were gratings differing in orientation and spatial frequency. The probability density function of first passage time for a random walk with drift process, which is defined by its barrier height and drift coefficient, was used to characterize the generating process of axonal discharge under resting and stimulus conditions. Drift coefficient and barrier height were derived from the sample mean and standard deviation of the measured inter-spike intervals. For cells with simple receptive fields, variations in spatial frequency produced changes only in drift coefficient. Variations in barrier height were produced only by changes in orientation of the stimulus. Currently, the method used to analyze these data was implemented in a simulation which displayed the relationship between the interval distribution of impulses, the random walk which represents the time series characteristic of the spike train model and the Gabor filter function which represents the geometry of the receptive field process.     

The development of the binocular depth cells in the secondary visual cortex of the lamb.

In most respects, the response properties of cells in the secondary visual cortex of the newborn lamb were indistinguishable from those in the adult. The cells were sharply selective to orientation; the orientation preferences were the same in each eye, and they varied systematically as the electrode penetrated the cortex. The receptive-field organization did not differ noticeably from that in adults, and complex, hypercomplex, and a few simple cells were all observed. The ocular dominance distribution was similar to that in the adult. Most importantly, binocular cells were found with disparate receptive fields even in newborn, visually inexperienced animals. As in the adult, the disparities were largely horizontal, and they appeared to be arranged in columns. Many of the cells responded preferentially to a binocular stimulus at a particular disparity setting (often approximately zero), but unlike those in the adult almost all the binocular cells in the newborn lamb would also respond monocularly, and the enhancement at the optimal disparity was less than in the adult. The full development of binocular selectivity took several weeks, and was blocked by binocular deprivation. We conclude that the basic wiring of stereoscopic mechanisms is innate, but the development of mature binocular interaction may depend on an adaptive process which makes use of the visual information received during binocular stimulation.     

Visual receptive field organization

Increasingly systematic approaches to quantifying receptive fields in primary visual cortex, combined with inspired ideas about functional circuitry, non-linearities, and visual stimuli, are bringing new interest to classical problems. This includes the distinction and hierarchy between simple and complex cells, the mechanisms underlying the receptive field surround, and debates about optimal stimuli for mapping receptive fields. An important new problem arises from recent observations of stimulus-dependent spatial and temporal summation in primary visual cortex. It appears that the receptive field can no longer be considered unique, and we might have to relinquish this cherished notion as the embodiment of neuronal function in primary visual cortex.     

Low response variability in simultaneously recorded retinal, thalamic, and cortical neurons

The response of a cortical cell to a repeated stimulus can be highly variable from one trial to the next. Much lower variability has been reported of retinal cells. We recorded visual responses simultaneously from three successive stages of the cat visual system: retinal ganglion cells (RGCs), thalamic (LGN) relay cells, and simple cells in layer 4 of primary visual cortex. Spike count variability was lower than that of a Poisson process at all three stages but increased at each stage. Absolute and relative refractory periods largely accounted for the reliability at all three stages. Our results show that cortical responses can be more reliable than previously thought. The differences in reliability in retina, LGN, and cortex can be explained by (1) decreasing firing rates and (2) decreasing absolute and relative refractory periods.     

Linear models of simple cells: correspondence to real cell responses and space spanning properties

Despite their limitations, linear filter models continue to be used to simulate the receptive field properties of cortical simple cells. For theoreticians interested in large scale models of visual cortex, a family of self-similar filters represents a convenient way in which to characterise simple cells in one basic model. This paper reviews research on the suitability of such models, and goes on to advance biologically motivated reasons for adopting a particular group of models in preference to all others. In particular, the paper describes why the Gabor model, so often used in network simulations, should be dropped in favour of a Cauchy model, both on the grounds of frequency response and mutual filter orthogonality.