scholarly journals Binocular Spatial Phase Tuning Characteristics of Neurons in the Macaque Striate Cortex

1997 ◽  
Vol 78 (1) ◽  
pp. 351-365 ◽  
Author(s):  
Earl L. Smith ◽  
Yuzo M. Chino ◽  
Jinren Ni ◽  
William H. Ridder ◽  
M.L.J. Crawford
Keyword(s):  
Spatial Frequency ◽  
Striate Cortex ◽  
Orientation Tuning ◽  
Absolute Position ◽  
Simple Cells ◽  
Complex Cells ◽  
Spatial Phase ◽  
Binocular Interactions ◽  
Phase Tuning ◽  
Phase Sensitive

Smith, Earl L., III, Yuzo M. Chino, Jinren Ni, William H. Ridder III, and M.L.J. Crawford. Binocular spatial phase tuning characteristics of neurons in the macaque striate cortex. J. Neurophysiol. 78: 351–365, 1997. We employed microelectrode recording techniques to study the sensitivity of individual neurons in the striate cortex of anesthetized and paralyzed monkeys to relative interocular image disparities and to determine the effects of basic stimulus parameters on these cortical binocular interactions. The visual stimuli were drifting sine wave gratings. After the optimal stimulus orientation, spatial frequency, and direction of stimulus movement were found, the cells' disparity tuning characteristics were determined by measuring responses as a function of the relative interocular spatial phase of dichoptic grating pairs. No attempts were made to assess absolute position disparities or horizontal disparities relative to the horopter. The majority (∼70%) of simple cells were highly sensitive to interocular spatial phase disparities, particularly neurons with balanced ocular dominances. Simple cells typically demonstrated binocular facilitation at the optimal phase disparity and binocular suppression for disparities 180° away. Fewer complex cells were phase selective (∼40%); however, the range of disparity selectivity in phase-sensitive complex cells was comparable with that for simple cells. Binocular interactions in non-phase-sensitive complex cells were evidenced by binocular response amplitudes that differed from responses to monocular stimulation. The degree of disparity tuning was independent of a cell's optimal orientation or the degree of direction tuning. However, disparity-sensitive cells tended to have narrow orientation tuning functions and the degree of disparity tuning was greatest for the optimal stimulus orientations. Rotating the stimulus for one eye 90° from the optimal orientation usually eliminated binocular interactions. The effects of phase disparities on the binocular response amplitude were also greatest at the optimal spatial frequency. Thus a cell's sensitivity to absolute position disparities reflects its spatial tuning characteristics, with cells sensitive to high spatial frequencies being capable of signaling very small changes in image disparity. On the other hand, stimulus contrast had relatively little effect on a cell's disparity tuning, because response saturation occurred at the same contrast level for all relative interocular phase disparities. Thus, as with orientation tuning, a cell's optimal disparity and the degree of disparity selectivity were invariant with contrast. Overall, the results show that sensitivity to interocular spatial phase disparities is a common property of striate neurons. A cell's disparity tuning characteristics appear to largely reflect its monocular receptive field properties and the interocular balance between excitatory and inhibitory inputs. However, distinct functional classes of cortical neurons could not be discriminated on the basis of disparity sensitivity alone.

Spatial Phase and the Temporal Structure of the Response to Gratings in V1

1998 ◽  
Vol 80 (2) ◽  
pp. 554-571 ◽  
Author(s):  
Jonathan D. Victor ◽  
Keith P. Purpura
Keyword(s):  
Spatial Frequency ◽  
Spike Train ◽  
Temporal Structure ◽  
Spike Timing ◽  
Dependent Manner ◽  
Data Set ◽  
Simple Cells ◽  
Complex Cells ◽  
Spatial Phase ◽  
Contrast Information

Victor, Jonathan D. and Keith P. Purpura. Spatial phase and the temporal structure of the response to gratings in V1. J. Neurophysiol. 80: 554–571, 1998. We recorded single-unit activity of 25 units in the parafoveal representation of macaque V1 to transient appearance of sinusoidal gratings. Gratings were systematically varied in spatial phase and in one or two of the following: contrast, spatial frequency, and orientation. Individual responses were compared based on spike counts, and also according to metrics sensitive to spike timing. For each metric, the extent of stimulus-dependent clustering of individual responses was assessed via the transmitted information, H. In nearly all data sets, stimulus-dependent clustering was maximal for metrics sensitive to the temporal pattern of spikes, typically with a precision of 25–50 ms. To focus on the interaction of spatial phase with other stimulus attributes, each data set was analyzed in two ways. In the “pooled phases” approach, the phase of the stimulus was ignored in the assessment of clustering, to yield an index H pooled. In the “individual phases” approach, clustering was calculated separately for each spatial phase and then averaged across spatial phases to yield an index H indiv. H pooled expresses the extent to which a spike train represents contrast, spatial frequency, or orientation in a manner which is not confounded by spatial phase (phase-independent representation), whereas H indiv expresses the extent to which a spike train represents one of these attributes, provided spatial phase is fixed (phase-dependent representation). Here, representation means that a stimulus attribute has a reproducible and systematic influence on individual responses, not a neural mechanism for decoding this influence. During the initial 100 ms of the response, contrast was represented in a phase-dependent manner by simple cells but primarily in a phase-independent manner by complex cells. As the response evolved, simple cell responses acquired phase-independent contrast information, whereas complex cells acquired phase-dependent contrast information. Simple cells represented orientation and spatial frequency in a primarily phase-dependent manner, but also they contained some phase-independent information in their initial response segment. Complex cells showed primarily phase-independent representation of orientation but primarily phase-dependent representation of spatial frequency. Joint representation of two attributes (contrast and spatial frequency, contrast and orientation, spatial frequency and orientation) was primarily phase dependent for simple cells, and primarily phase independent for complex cells. In simple and complex cells, the variability in the number of spikes elicited on each response was substantially greater than the expectations of a Poisson process. Although some of this variation could be attributed to the dependence of the response on the spatial phase of the grating, variability was still markedly greater than Poisson when the contribution of spatial phase to response variance was removed.

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.

Plasticity of neuronal response properties in adult cat striate cortex

1998 ◽  
Vol 15 (1) ◽  
pp. 177-196 ◽  
Author(s):  
J. MCLEAN ◽  
L.A. PALMER
Keyword(s):  
Visual Cortex ◽  
Striate Cortex ◽  
Direction Selectivity ◽  
Orientation Tuning ◽  
Tuning Curves ◽  
Spatial Phase ◽  
Associative Conditioning ◽  
Phase Tuning ◽  
Adult Cat ◽  
Cat Striate Cortex

We have utilized an associative conditioning paradigm to induce changes in the receptive field (RF) properties of neurons in the adult cat striate cortex. During conditioning, the presentation of particular visual stimuli were repeatedly paired with the iontophoretic application of either GABA or glutamate to control postsynaptic firing rates. Similar paradigms have been used in kitten visual cortex to alter RF properties (Fregnac et al., 1988, 1992; Greuel et al., 1988; Shulz & Fregnac, 1992). Roughly half of the cells that were subjected to conditioning with stimuli differing in orientation were found to have orientation tuning curves that were significantly altered. In general, the modification in orientation tuning was not accompanied by a shift in preferred orientation, but rather, responsiveness to stimuli at or near the positively reinforced orientation was increased relative to controls, and responsiveness to stimuli at or near the negatively reinforced orientation was decreased relative to controls. A similar proportion of cells that were subjected to conditioning with stimuli differing in spatial phase were found to have spatial-phase tuning curves that were significantly modified. Conditioning stimuli typically differed by 90 deg in spatial phase, but modifications in spatial-phase angle were generally 30–40 deg. An interesting phenomenon we encountered was that during conditioning, cells often developed a modulated response to counterphased grating stimuli presented at the null spatial phase. We present an example of a simple cell for which the shift in preferred spatial phase measured with counterphased grating stimuli was comparable to the shift in spatial phase computed from a one-dimensional Gabor fit of the space-time RF profile. One of ten cells tested had a significant change in direction selectivity following associative conditioning. The specific and predictable modifications of RF properties induced by our associative conditioning procedure demonstrate the ability of mature visual cortical neurons to alter their integrative properties. Our results lend further support to models of synaptic plasticity where temporal correlations between presynaptic and postsynaptic activity levels control the efficiency of transmission at existing synapses, and to the idea that the mature visual cortex is, in some sense, dynamically organized.

scholarly journals Comparison Among Some Models of Orientation Selectivity

2006 ◽  
Vol 96 (1) ◽  
pp. 404-419 ◽  
Author(s):  
Andrew F. Teich ◽  
Ning Qian
Keyword(s):  
Peak Shift ◽  
Orientation Tuning ◽  
Complex Spectrum ◽  
Complex Cell ◽  
Phase Information ◽  
Simple Complex ◽  
Simple Cells ◽  
Complex Cells ◽  
Spatial Phase ◽  
Opposite Contrast

Several models exist for explaining primary visual cortex (V1) orientation tuning. The modified feedforward model (MFM) and the recurrent model (RM) are major examples. We have implemented these two models, at the same level of detail, alongside a few newer variations, and thoroughly compared their receptive-field structures. We found that antiphase inhibition in the MFM enhances both spatial phase information and orientation tuning, producing well-tuned simple cells. This remains true for a newer version of the MFM that incorporates untuned complex-cell inhibition. In contrast, when the recurrent connections in the RM are strong enough to produce typical V1 orientation tuning, they also eliminate spatial phase information, making the cells complex. Introducing phase specificity into the connections of the RM (as done in an original version of the RM) can make the cells phase sensitive, but the cells show an incorrect 90° peak shift of orientation tuning under opposite contrast signs. An inhibition-dominant version of the RM can generate well-tuned cells across the simple–complex spectrum, but it predicts that the net effect of cortical interactions is to suppress feedforward excitation across all orientations in simple cells. Finally, adding antiphase inhibition used in the MFM into the RM produces a most general model. We call this new model the modified recurrent model (MRM) and show that this model can also produce well-tuned cells throughout the simple–complex spectrum. Unlike the inhibition-dominant RM, the MRM is consistent with data from cat V1, suggesting that the net effect of cortical interactions is to boost simple cell responses at the preferred orientation. These results suggest that the MFM is well suited for explaining orientation tuning in simple cells, whereas the standard RM is for complex cells. The assignment of the RM to complex cells also avoids conflicts between the RM and the experiments of cortical inactivation (done on simple cells) and the spatial-frequency dependency of orientation tuning (found in simple cells). Because orientation-tuned V1 cells show a continuum of simple- to complex-cell behavior, the MRM provides the best description of V1 data.

Oblique Effect: A Neural Basis in the Visual Cortex

2003 ◽  
Vol 90 (1) ◽  
pp. 204-217 ◽  
Author(s):  
Baowang Li ◽  
Matthew R. Peterson ◽  
Ralph D. Freeman
Keyword(s):  
Striate Cortex ◽  
Oblique Effect ◽  
Orientation Tuning ◽  
Neural Basis ◽  
Linear Filters ◽  
Simple Cells ◽  
Tuning Curves ◽  
Complex Cells ◽  
Behavioral Studies ◽  
Spatial Frequencies

The details of oriented visual stimuli are better resolved when they are horizontal or vertical rather than oblique. This “oblique effect” has been confirmed in numerous behavioral studies in humans and to some extent in animals. However, investigations of its neural basis have produced mixed and inconclusive results, presumably due in part to limited sample sizes. We have used a database to analyze a population of 4,418 cells in the cat's striate cortex to determine possible differences as a function of orientation. We find that both the numbers of cells and the widths of orientation tuning vary as a function of preferred orientation. Specifically, more cells prefer horizontal and vertical orientations compared with oblique angles. The largest population of cells is activated by orientations close to horizontal. In addition, orientation tuning widths are most narrow for cells preferring horizontal orientations. These findings are most prominent for simple cells tuned to high spatial frequencies. Complex cells and simple cells tuned to low spatial frequencies do not exhibit these anisotropies. For a subset of simple cells from our population ( n = 104), we examined the relative contributions of linear and nonlinear mechanisms in shaping orientation tuning curves. We find that linear contributions alone do not account for the narrower tuning widths at horizontal orientations. By modeling simple cells as linear filters followed by static expansive nonlinearities, our analysis indicates that horizontally tuned cells have a greater nonlinear component than those tuned to other orientations. This suggests that intracortical mechanisms play a major role in shaping the oblique effect.

Responses of neurons in cat striate cortex to vernier offsets in reverse contrast stimuli

1995 ◽  
Vol 12 (5) ◽  
pp. 805-817 ◽  
Author(s):  
N.v. Swindale
Keyword(s):  
Striate Cortex ◽  
Tuning Curve ◽  
High Sensitivity ◽  
Orientation Tuning ◽  
Vernier Acuity ◽  
Simple Cells ◽  
Tuning Curves ◽  
Complex Cells ◽  
Simple And Complex Cells ◽  
Opposite Contrast

AbstractThis paper examines how the responses of cells in area 17 of the cat vary as a function of the vernier offset between a bright and a dark bar. The study was prompted by the finding that human vernier acuity is reduced for bars or edges of opposite contrast sign (Mather & Morgan, 1986; O'Shea & Mitchell, 1990). Both simple and complex cells showed V-shaped tuning curves for reverse contrast stimuli: i.e. response was minimum at alignment, and increased with increasing vernier offset. For vernier bars with the same contrast sign, γ-shaped tuning curves were found, as reported earlier (Swindale & Cynader, 1986). Sensitivity to offset was inversely correlated in the two paradigms. However, complex cells with high sensitivity to offsets in a normal vernier stimulus were significantly less sensitive to offsets in reverse contrast stimuli. A cell's response to a vernier stimulus in which both bars are bright can be predicted by the shape of its orientation tuning curve, if the vernier stimulus is approximated by a single bar with an orientation equal to that of a line joining the midpoints of the two component bars (Swindale & Cynader, 1986). This approximation did not hold for the reverse contrast condition: orientation tuning curves for compound barswere broad and shallow, rather than bimodal, with peaks up to 40 deg from the preferred orientation. Results from simple cells were compared with predictions made by a linear model of the receptive field. The model predicted the V-shaped tuning curves found for reverse contrast stimuli. It also predicted that absolute values of tuning slopes for vernier offsets in reverse contrast stimuli might sometimes be higher than with normal stimuli. This was observed in some simple cells. The model was unable to explain the shape of orientation tuning curves for compound bars, nor could it explain the breakdown of the equivalent orientation approximation.

scholarly journals Spatial phase sensitivity of complex cells in primary visual cortex depends on stimulus contrast

2015 ◽  
Vol 114 (6) ◽  
pp. 3326-3338 ◽  
Author(s):  
H. Meffin ◽  
M. A. Hietanen ◽  
S. L. Cloherty ◽  
M. R. Ibbotson
Keyword(s):  
Visual Cortex ◽  
Primary Visual Cortex ◽  
Receptive Fields ◽  
Spatial Summation ◽  
Phase Sensitivity ◽  
Simple Cells ◽  
Low Contrast ◽  
Complex Cells ◽  
Spatial Phase ◽  
Phase Sensitive

Neurons in primary visual cortex are classified as simple, which are phase sensitive, or complex, which are significantly less phase sensitive. Previously, we have used drifting gratings to show that the phase sensitivity of complex cells increases at low contrast and after contrast adaptation while that of simple cells remains the same at all contrasts (Cloherty SL, Ibbotson MR. J Neurophysiol 113: 434–444, 2015; Crowder NA, van Kleef J, Dreher B, Ibbotson MR. J Neurophysiol 98: 1155–1166, 2007; van Kleef JP, Cloherty SL, Ibbotson MR. J Physiol 588: 3457–3470, 2010). However, drifting gratings confound the influence of spatial and temporal summation, so here we have stimulated complex cells with gratings that are spatially stationary but continuously reverse the polarity of the contrast over time (contrast-reversing gratings). By varying the spatial phase and contrast of the gratings we aimed to establish whether the contrast-dependent phase sensitivity of complex cells results from changes in spatial or temporal processing or both. We found that most of the increase in phase sensitivity at low contrasts could be attributed to changes in the spatial phase sensitivities of complex cells. However, at low contrasts the complex cells did not develop the spatiotemporal response characteristics of simple cells, in which paired response peaks occur 180° out of phase in time and space. Complex cells that increased their spatial phase sensitivity at low contrasts were significantly overrepresented in the supragranular layers of cortex. We conclude that complex cells in supragranular layers of cat cortex have dynamic spatial summation properties and that the mechanisms underlying complex cell receptive fields differ between cortical layers.

Direction selectivity of complex cells in a comparison with simple cells

1975 ◽  
Vol 38 (6) ◽  
pp. 1524-1540 ◽  
Author(s):  
A. W. Goodwin ◽  
G. H. Henry
Keyword(s):  
Spontaneous Activity ◽  
Visual Information ◽  
Striate Cortex ◽  
Cell Types ◽  
Direction Selectivity ◽  
Complex Cell ◽  
Simple Cells ◽  
Complex Cells ◽  
Sequential Processing

Following our earlier study on direction selectivity in simple cells (5), the present findings on complex cells made it possible to compare the direction selectivity in the two types of striate cell. Common properties were found in the dimension of the smallest stimulus displacement giving a direction-selective response and in the role of inhibition in suppressing the response as the stimulus moved in the nonpreferred direction. However, the effectiveness of this inhibition varied in the two cell types since it suppressed both driven and spontaneous activity in the simple cell, but only driven firing in the complex cell. It is argued that direction selectivity must enter the response before the complex cell if the inhibition responsible for it's generation fails to influence the spontaneous activity of the cell. The consequences of this finding are considered in the terms of parallel or sequential processing of visual information in striate cortex.

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.

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