Spatial Organization of Receptive Fields of V1 Neurons of Alert Monkeys: Comparison With Responses to Gratings

2002 ◽  
Vol 88 (5) ◽  
pp. 2557-2574 ◽  
Author(s):  
Igor Kagan ◽  
Moshe Gur ◽  
D. Max Snodderly
Keyword(s):  
Eye Movements ◽  
Spatial Organization ◽  
Receptive Fields ◽  
Window Size ◽  
Group 14 ◽  
Simple Cells ◽  
Complex Cells ◽  
V1 Neurons ◽  
Fixational Eye Movements ◽  
Spatial Frequencies

We studied the spatial organization of receptive fields and the responses to gratings of neurons in parafoveal V1 of alert monkeys. Activating regions (ARs) of 228 cells were mapped with increment and decrement bars while compensating for fixational eye movements. For cells with two or more ARs, the overlap between ARs responsive to increments (INC) and ARs responsive to decrements (DEC) was characterized by a quantitative overlap index (OI). The distribution of overlap indices was bimodal. The larger group (78% of cells) was composed of complex cells with strongly overlapping ARs (OI ≥ 0.5). The smaller group (14%) was composed of simple cells with minimal spatial overlap of ARs (OI ≤ 0.3). Simple cells were preferentially located in layers dominated by the magnocellular pathway. A third group of neurons, the monocontrast cells (8%), responded only to one sign of contrast and had more sustained responses to flashed stimuli than other cells. One hundred fourteen neurons were also studied with drifting sinusoidal gratings of various spatial frequencies and window widths. For complex cells, the relative modulation (RM, the ratio of the 1st harmonic to the mean firing rate), ranged from 0.6 ± 0.4 to 1.1 ± 0.5 (mean ± SD), depending on the stimulus conditions and the mode of correction for eye movements. RM was not correlated with the degree of overlap of ARs, indicating that the spatial organization of receptive fields cannot reliably be predicted from RM values. In fact, a subset of complex cells had RM > 1, the traditional criterion for identifying simple cells. However, unlike simple cells, even those complex cells with high RM could exhibit diverse nonlinear responses when the spatial frequency or window size was changed. Furthermore, the responses of complex cells to counterphase gratings were predominantly nonlinear even harmonics. These results show that RM is not a robust test of linearity. Our results indicate that complex cells are the most frequently encountered neurons in primate V1, and their behavior needs to receive more emphasis in models of visual function.

Cone Inputs to Simple and Complex Cells in V1 of Awake Macaque

2007 ◽  
Vol 97 (4) ◽  
pp. 3070-3081 ◽  
Author(s):  
Gregory D. Horwitz ◽  
E. J. Chichilnisky ◽  
Thomas D. Albright
Keyword(s):  
Linear Models ◽  
Noise Analysis ◽  
Receptive Fields ◽  
Light Responses ◽  
Simple Cells ◽  
Complex Cells ◽  
V1 Neurons ◽  
Spike Triggered Averaging ◽  
Simple And Complex Cells ◽  
Simple And Complex Cell

Rules by which V1 neurons combine signals originating in the cone photoreceptors are poorly understood. We measured cone inputs to V1 neurons in awake, fixating monkeys with white-noise analysis techniques that reveal properties of light responses not revealed by purely linear models used in previous studies. Simple cells were studied by spike-triggered averaging that is robust to static nonlinearities in spike generation. This analysis revealed, among heterogeneously tuned neurons, two relatively discrete categories: one with opponent L- and M-cone weights and another with nonopponent cone weights. Complex cells were studied by spike-triggered covariance, which identifies features in the stimulus sequence that trigger spikes in neurons with receptive fields containing multiple linear subunits that combine nonlinearly. All complex cells responded to nonopponent stimulus modulations. Although some complex cells responded to cone-opponent stimulus modulations too, none exhibited the pure opponent sensitivity observed in many simple cells. These results extend the findings on distinctions between simple and complex cell chromatic tuning observed in previous studies in anesthetized monkeys.

Internal Spatial Organization of Receptive Fields of Complex Cells in the Early Visual Cortex

2007 ◽  
Vol 98 (3) ◽  
pp. 1194-1212 ◽  
Author(s):  
Kota S. Sasaki ◽  
Izumi Ohzawa
Keyword(s):  
Visual Cortex ◽  
Minimal Model ◽  
Spatial Organization ◽  
Receptive Fields ◽  
Simple Cell ◽  
Complex Cell ◽  
Linear Filters ◽  
Simple Cells ◽  
Complex Cells ◽  
Early Visual Cortex

The receptive fields of complex cells in the early visual cortex are economically modeled by combining outputs of a quadrature pair of linear filters. For actual complex cells, such a minimal model may be insufficient because many more simple cells are thought to make up a complex cell receptive field. To examine the minimalist model physiologically, we analyzed spatial relationships between the internal structure (subunits) and the overall receptive fields of individual complex cells by a two-stimulus interaction technique. The receptive fields of complex cells are more circular and only slightly larger than their subunits in size. In addition, complex cell subunits occupy spatial extents similar to those of simple cell receptive fields. Therefore in these respects, the minimalist schema is a fair approximation to actual complex cells. However, there are violations against the minimal model. Simple cell receptive fields have significantly fewer subregions than complex cell subunits and, in general, simple cell receptive fields are elongated more horizontally than vertically. This bias is absent in complex cell subunits and receptive fields. Thus simple cells cannot be equated to individual complex cell subunits and spatial pooling of simple cells may occur anisotropically to constitute a complex cell subunit. Moreover, when linear filters for complex cell subunits are examined separately for bright and dark responses, there are significant imbalances and position displacements between them. This suggests that actual complex cell receptive fields are constructed by a richer combination of linear filters than proposed by the minimalist model.

scholarly journals Coding of chromatic spatial contrast by macaque V1 neurons

2021 ◽  
Author(s):  
Abhishek De ◽  
Gregory D. Horwitz
Keyword(s):  
Color Image ◽  
Receptive Fields ◽  
Luminance Contrast ◽  
Chromatic Contrast ◽  
Simple Cells ◽  
Complex Cells ◽  
V1 Neurons ◽  
Full Color ◽  
Common Principle ◽  
Neural Selectivity

ABSTRACTColor perception relies on spatial comparisons of chromatic signals, but how the brain performs these comparisons is poorly understood. Here, we show that many V1 neurons compare signals across their receptive fields (RF) using a common principle. We estimated the spatial-chromatic RFs of each neuron, and then measured neural selectivity to customized colored edges that were sampled using a novel closed-loop technique. We found that many double-opponent (DO) cells, which have spatially and chromatically opponent RFs, responded to chromatic contrast as a weighted sum, akin to how simple cells respond to luminance contrast. Other neurons integrated chromatic signals non-linearly, confirming that linear signal integration is not an obligate property of V1 neurons. The functional similarity of DO and simple cells suggests a common underlying neural circuitry, promotes the construction of image-computable models for full-color image representation, and sheds new light on V1 complex cells.

scholarly journals Beyond Fixation: detailed characterization of neural selectivity in free-viewing primates

2021 ◽  
Author(s):  
Jacob L Yates ◽  
Shanna H Coop ◽  
Gabriel H Sarch ◽  
Ruei-Jr Wu ◽  
Daniel A Butts ◽  
...  
Keyword(s):  
Eye Movements ◽  
Receptive Fields ◽  
Natural Behavior ◽  
Visual Neurons ◽  
Trained Subjects ◽  
V1 Neurons ◽  
Fixational Eye Movements ◽  
Neural Selectivity

Virtually all vision studies use a fixation point to stabilize gaze, rendering stimuli on video screens fixed to retinal coordinates. This approach requires trained subjects, is limited by the accuracy of fixational eye movements, and ignores the role of eye movements in shaping visual input. To overcome these limitations, we developed a suite of hardware and software tools to study vision during natural behavior in untrained subjects. We show this approach recovers receptive fields and tuning properties of visual neurons from multiple cortical areas of marmoset monkeys. Combined with high-precision eye-tracking, it achieves sufficient resolution to recover the receptive fields of foveal V1 neurons. These findings demonstrate the power of this approach for characterizing neural response while simultaneously studying the dynamics of natural behavior.

Eye Position Compensation Improves Estimates of Response Magnitude and Receptive Field Geometry in Alert Monkeys

2007 ◽  
Vol 97 (5) ◽  
pp. 3439-3448 ◽  
Author(s):  
Yamei Tang ◽  
Alan Saul ◽  
Moshe Gur ◽  
Stephanie Goei ◽  
Elsie Wong ◽  
...  
Keyword(s):  
Eye Movements ◽  
Receptive Field ◽  
Receptive Fields ◽  
Eye Position ◽  
Stimulus Position ◽  
Fixational Eye Movements ◽  
Field Geometry ◽  
Receptive Field Subregions ◽  
Movement Compensation ◽  
Definition Of

Studies of visual function in behaving subjects require that stimuli be positioned reliably on the retina in the presence of eye movements. Fixational eye movements scatter stimuli about the retina, inflating estimates of receptive field dimensions, reducing estimates of peak responses, and blurring maps of receptive field subregions. Scleral search coils are frequently used to measure eye position, but their utility for correcting the effects of fixational eye movements on receptive field maps has been questioned. Using eye coils sutured to the sclera and preamplifiers configured to minimize cable artifacts, we reexamined this issue in two rhesus monkeys. During repeated fixation trials, the eye position signal was used to adjust the stimulus position, compensating for eye movements and correcting the stimulus position to place it at the desired location on the retina. Estimates of response magnitudes and receptive field characteristics in V1 and in LGN were obtained in both compensated and uncompensated conditions. Receptive fields were narrower, with steeper borders, and response amplitudes were higher when eye movement compensation was used. In sum, compensating for eye movements facilitated more precise definition of the receptive field. We also monitored horizontal vergence over long sequences of fixation trials and found the variability to be low, as expected for this precise behavior. Our results imply that eye coil signals can be highly accurate and useful for optimizing visual physiology when rigorous precautions are observed.

Receptive-field characteristics of neurons in cat striate cortex: Changes with visual field eccentricity

1976 ◽  
Vol 39 (3) ◽  
pp. 512-533 ◽  
Author(s):  
J. R. Wilson ◽  
S. M. Sherman
Keyword(s):  
Visual Field ◽  
Receptive Field ◽  
Striate Cortex ◽  
Ocular Dominance ◽  
Receptive Fields ◽  
Direction Selectivity ◽  
Orientation Selectivity ◽  
Simple Cells ◽  
Complex Cells ◽  
Cat Striate Cortex

1. Receptive-field properties of 214 neurons from cat striate cortex were studied with particular emphasis on: a) classification, b) field size, c) orientation selectivity, d) direction selectivity, e) speed selectivity, and f) ocular dominance. We studied receptive fields located throughtout the visual field, including the monocular segment, to determine how receptivefield properties changed with eccentricity in the visual field.2. We classified 98 cells as "simple," 80 as "complex," 21 as "hypercomplex," and 15 in other categories. The proportion of complex cells relative to simple cells increased monotonically with receptive-field eccenticity.3. Direction selectivity and preferred orientation did not measurably change with eccentricity. Through most of the binocular segment, this was also true for ocular dominance; however, at the edge of the binocular segment, there were more fields dominated by the contralateral eye.4. Cells had larger receptive fields, less orientation selectivity, and higher preferred speeds with increasing eccentricity. However, these changes were considerably more pronounced for complex than for simple cells.5. These data suggest that simple and complex cells analyze different aspects of a visual stimulus, and we provide a hypothesis which suggests that simple cells analyze input typically from one (or a few) geniculate neurons, while complex cells receive input from a larger region of geniculate neurons. On average, this region is invariant with eccentricity and, due to a changing magnification factor, complex fields increase in size with eccentricity much more than do simple cells. For complex cells, computations of this geniculate region transformed to cortical space provide a cortical extent equal to the spread of pyramidal cell basal dendrites.

Binocular Combination of Contrast Signals by Striate Cortical Neurons in the Monkey

1997 ◽  
Vol 78 (1) ◽  
pp. 366-382 ◽  
Author(s):  
Earl L. Smith ◽  
Yuzo Chino ◽  
Jinren Ni ◽  
Han Cheng
Keyword(s):  
Cortical Neurons ◽  
Striate Cortex ◽  
Receptive Fields ◽  
Spatial Summation ◽  
Simple Cells ◽  
Complex Cells ◽  
Threshold Response ◽  
Binocular Interactions ◽  
Specific Complex ◽  
Left And Right

Smith, Earl L., III, Yuzo Chino, Jinren Ni, and Han Cheng. Binocular combination of contrast signals by striate cortical neurons in the monkey. J. Neurophysiol. 78: 366–382, 1997. With the use of microelectrode recording techniques, we investigated how the contrast signals from the two eyes are combined in individual cortical neurons in the striate cortex of anesthetized and paralyzed macaque monkeys. For a given neuron, the optimal spatial frequency, orientation, and direction of drift for sine wave grating stimuli were determined for each eye. The cell's disparity tuning characteristics were determined by measuring responses as a function of the relative interocular spatial phase of dichoptic stimuli that consisted of the optimal monocular gratings. Binocular contrast summation was then investigated by measuring contrast response functions for optimal dichoptic grating pairs that had left- to right-eye interocular contrast ratios that varied from 0.1 to 10. The goal was to determine the left- and right-eye contrast components required to produce a criterion threshold response. For all functional classes of cortical neurons and for both cooperative and antagonistic binocular interactions, there was a linear relationship between the left- and right-eye contrast components required to produce a threshold response. Thus, for example for cooperative binocular interactions, a reduction in contrast to one eye was counterbalanced by an equivalent increase in contrast to the other eye. These results showed that in simple cells and phase-specific complex cells, the contrast signals from the two eyes were linearly combined at the subunit level before nonlinear rectification. In non-phase-specific complex cells, the linear binocular convergence of contrast signals could have taken place either before or after the rectification process, but before spike generation. In addition, for simple cells, vector analysis of spatial summation showed that the inputs from the two eyes were also combined in a linear manner before nonlinear spike-generating mechanisms. Thus simple cells showed linear spatial summation not only within and between subregions in a given receptive field, but also between the left- and right-eye receptive fields. Overall, the results show that the effectiveness of a stimulus in producing a response reflects interocular differences in the relative balance of inputs to a given cell, however, the eye of origin of a light-evoked signal has no specific consequence.

scholarly journals Spatial and Temporal Features of Synaptic to Discharge Receptive Field Transformation in Cat Area 17

2010 ◽  
Vol 103 (2) ◽  
pp. 677-697 ◽  
Author(s):  
Lionel G. Nowak ◽  
Maria V. Sanchez-Vives ◽  
David A. McCormick
Keyword(s):  
Spatial Frequency ◽  
Response Latency ◽  
Frequency Tuning ◽  
Receptive Fields ◽  
Width Ratio ◽  
Area 17 ◽  
Simple Cells ◽  
Complex Cells ◽  
Temporal Features ◽  
Simple And Complex Cells

The aim of the present study was to characterize the spatial and temporal features of synaptic and discharge receptive fields (RFs), and to quantify their relationships, in cat area 17. For this purpose, neurons were recorded intracellularly while high-frequency flashing bars were used to generate RFs maps for synaptic and spiking responses. Comparison of the maps shows that some features of the discharge RFs depended strongly on those of the synaptic RFs, whereas others were less dependent. Spiking RF duration depended poorly and spiking RF amplitude depended moderately on those of the underlying synaptic RFs. At the other extreme, the optimal spatial frequency and phase of the discharge RFs in simple cells were almost entirely inherited from those of the synaptic RFs. Subfield width, in both simple and complex cells, was less for spiking responses compared with synaptic responses, but synaptic to discharge width ratio was relatively variable from cell to cell. When considering the whole RF of simple cells, additional variability in width ratio resulted from the presence of additional synaptic subfields that remained subthreshold. Due to these additional, subthreshold subfields, spatial frequency tuning predicted from synaptic RFs appears sharper than that predicted from spiking RFs. Excitatory subfield overlap in spiking RFs was well predicted by subfield overlap at the synaptic level. When examined in different regions of the RF, latencies appeared to be quite variable, but this variability showed negligible dependence on distance from the RF center. Nevertheless, spiking response latency faithfully reflected synaptic response latency.

scholarly journals Substructure of Direction-Selective Receptive Fields in Macaque V1

2003 ◽  
Vol 89 (5) ◽  
pp. 2743-2759 ◽  
Author(s):  
Margaret S. Livingstone ◽  
Bevil R. Conway
Keyword(s):  
Receptive Fields ◽  
Direction Selectivity ◽  
Positive Interactions ◽  
Null Direction ◽  
Simple Cells ◽  
Complex Cells ◽  
Preferred Direction ◽  
Orientation Axis ◽  
Interaction Regions ◽  
Simple And Complex Cells

We used two-dimensional (2-D) sparse noise to map simultaneous and sequential two-spot interactions in simple and complex direction-selective cells in macaque V1. Sequential-interaction maps for both simple and complex cells showed preferred-direction facilitation and null-direction suppression for same-contrast stimulus sequences and the reverse for inverting-contrast sequences, although the magnitudes of the interactions were weaker for the simple cells. Contrast-sign selectivity in complex cells indicates that direction-selective interactions in these cells must occur in antecedent simple cells or in simple-cell-like dendritic compartments. Our maps suggest that direction selectivity, and on andoff segregation perpendicular to the orientation axis, can occur prior to receptive-field elongation along the orientation axis. 2-D interaction maps for some complex cells showed elongated alternating facilitatory and suppressive interactions as predicted if their inputs were orientation-selective simple cells. The negative interactions, however, were less elongated than the positive interactions, and there was an inflection at the origin in the positive interactions, so the interactions were chevron-shaped rather than band-like. Other complex cells showed only two round interaction regions, one negative and one positive. Several explanations for the map shapes are considered, including the possibility that directional interactions are generated directly from unoriented inputs.

A model of the dynamics of retinal activity during natural visual fixation

2007 ◽  
Vol 24 (2) ◽  
pp. 217-230 ◽  
Author(s):  
GAËLLE DESBORDES ◽  
MICHELE RUCCI
Keyword(s):  
Eye Movements ◽  
Visual Field ◽  
Ganglion Cells ◽  
Receptive Fields ◽  
Visual Fixation ◽  
Natural Scenes ◽  
M Cells ◽  
Retinal Activity ◽  
Cell Responses ◽  
Fixational Eye Movements

During visual fixation, small eye movements keep the retinal image continuously in motion. It is known that neurons in the visual system are sensitive to the spatiotemporal modulations of luminance resulting from this motion. In this study, we examined the influence of fixational eye movements on the statistics of neural activity in the macaque's retina during the brief intersaccadic periods of natural visual fixation. The responses of parvocellular (P) and magnocellular (M) ganglion cells in different regions of the visual field were modeled while their receptive fields scanned natural images following recorded traces of eye movements. Immediately after the onset of fixation, wide ensembles of coactive ganglion cells extended over several degrees of visual angle, both in the central and peripheral regions of the visual field. Following this initial pattern of activity, the covariance between the responses of pairs of P and M cells and the correlation between the responses of pairs of M cells dropped drastically during the course of fixation. Cell responses were completely uncorrelated by the end of a typical 300-ms fixation. This dynamic spatial decorrelation of retinal activity is a robust phenomenon independent of the specifics of the model. We show that it originates from the interaction of three factors: the statistics of natural scenes, the small amplitudes of fixational eye movements, and the temporal sensitivities of ganglion cells. These results support the hypothesis that fixational eye movements, by shaping the statistics of retinal activity, are an integral component of early visual representations.

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