Annular Stimuli Research Articles

Parafoveal and Perifoveal Accommodation Response to Defocus Changes Induced by a Tunable Lens

The accommodative response of the human eye is predominantly driven by foveal vision, but reacts also to off-foveal stimuli. Here, we report on monocular accommodation measurements using parafoveal and perifoveal annular stimuli centered around the fovea and extending up to 8° radial eccentricity for young emmetropic and myopic subjects. The stimuli were presented through a sequence of random defocus step changes induced by a pupil-conjugated tunable lens. A Hartmann–Shack wavefront sensor with an infrared beacon was used to measure real-time changes in ocular aberrations up to and including the fourth radial order across a 3 mm pupil at 20 Hz. Our findings show a significant reduction in accommodative response with increased radial eccentricity.

Despite Impaired Binocular Function, Binocular Disparity Integration Across the Visual Field Is Spared in Normal Aging and Glaucoma.

To examine how binocularly asymmetric glaucomatous visual field damage affects binocular disparity processing across the visual field. We recruited 18 patients with primary open-angle glaucoma, 16 age-matched controls, and 13 young controls. Participants underwent standard clinical assessments of binocular visual acuity, binocular contrast sensitivity, stereoacuity, and perimetry. We employed a previously validated psychophysical procedure to measure how sensitivity to binocular disparity varied across spatial frequencies and visual field sectors (i.e., with full-field stimuli spanning the central 21° of the visual field and with stimuli restricted to annular regions spanning 0°-3°, 3°-9°, or 9°-21°). We employed measurements with annular stimuli to model different possible scenarios regarding how disparity information is combined across visual field sectors. We adjudicated between potential mechanisms by comparing model predictions to the patterns observed with full-field stimuli. Perimetry confirmed that patients with glaucoma exhibited binocularly asymmetric visual field damage (P < 0.001). Across participant groups, foveal regions preferentially processed disparities at finer spatial scales, whereas periphery regions were tuned for coarser scales (P < 0.001). Disparity sensitivity also decreased from fovea to periphery (P < 0.001) and across participant groups (Ps < 0.01). Finally, similar to controls, patients with glaucoma exhibited near-optimal disparity integration, specifically at low spatial frequencies (P < 0.001). Contrary to the conventional view that glaucoma spares central vision, we find that glaucomatous damage causes a widespread loss of disparity sensitivity across both foveal and peripheral regions. Despite these losses, cortical integration mechanisms appear to be well preserved, suggesting that patients with glaucoma make the best possible use of their remaining binocular function.

Mapping spatial frequency preferences across human primary visual cortex.

Neurons in primate visual cortex (area V1) are tuned for spatial frequency, in a manner that depends on their position in the visual field. Several studies have examined this dependency using functional magnetic resonance imaging (fMRI), reporting preferred spatial frequencies (tuning curve peaks) of V1 voxels as a function of eccentricity, but their results differ by as much as two octaves, presumably owing to differences in stimuli, measurements, and analysis methodology. Here, we characterize spatial frequency tuning at a millimeter resolution within the human primary visual cortex, across stimulus orientation and visual field locations. We measured fMRI responses to a novel set of stimuli, constructed as sinusoidal gratings in log-polar coordinates, which include circular, radial, and spiral geometries. For each individual stimulus, the local spatial frequency varies inversely with eccentricity, and for any given location in the visual field, the full set of stimuli span a broad range of spatial frequencies and orientations. Over the measured range of eccentricities, the preferred spatial frequency is well-fit by a function that varies as the inverse of the eccentricity plus a small constant. We also find small but systematic effects of local stimulus orientation, defined in both absolute coordinates and relative to visual field location. Specifically, peak spatial frequency is higher for pinwheel than annular stimuli and for horizontal than vertical stimuli.

Photoreceptor-Specific Loss of Perifoveal Temporal Contrast Sensitivity in Retinitis Pigmentosa.

PurposeInherited retinal diseases affect the L-, M-, S-cones and rods in distinct ways, which calls for new methods that enable quantification of photoreceptor-specific functions. We tested the feasibility of using the silent substitution paradigm to estimate photoreceptor-driven temporal contrast sensitivity (tCS) functions in patients with retinitis pigmentosa.MethodsThe silent substitution paradigm is based on substitution of lights of different spectral composition; this offers considerable advantage over other stimulation techniques. We used a four-primary LED stimulator to create perifoveal annular stimuli (2° inner, 12° outer diameters) and used a triple silent substitution to probe photoreceptor-selective tCS. Measurements were performed in a heterogeneous cohort of 15 patients with retinitis pigmentosa and related to those in a control group of nine color-normal healthy observers. Age differences between groups were addressed with a model of age-related normal contrast sensitivity derived from measurements in 20 healthy observers aged between 23 and 83 years.ResultsThe age-related loss of tCS amounted to 0.1 dB/year in healthy subjects across all photoreceptor subtypes. In patients, tCS was decreased for every photoreceptor subtype; however, S-cone- and rod-driven sensitivities were most strongly affected. Postreceptoral mechanisms were not affected.ConclusionsThis feasibility study provides evidence that the silent substitution technique enables the estimation of photoreceptor-selective tCS functions and can serve as an accurate biomarker of photoreceptor-specific contrast sensitivity loss in patients with retinitis pigmentosa.Translational RelevanceWe aim to develop tests of visual function for clinical trials of novel therapies for inherited retinal diseases from methods that can currently be used only in vision research labs.

The spatial properties of L- and M-cone inputs to electroretinograms that reflect different types of post-receptoral processing.

We studied the spatial arrangement of L- and M-cone driven electroretinograms (ERGs) reflecting the activity of magno- and parvocellular pathways. L- and M-cone isolating sine wave stimuli were created with a four primary LED stimulator using triple silent substitution paradigms. Temporal frequencies were 8 and 12 Hz, to reflect cone opponent activity, and 30, 36 and 48 Hz to reflect luminance activity. The responses were measured for full-field stimuli and for different circular and annular stimuli. The ERG data confirm the presence of two different mechanisms at intermediate and high temporal frequencies. The responses measured at high temporal frequencies strongly depended upon spatial stimulus configuration. In the full-field conditions, the L-cone driven responses were substantially larger than the full-field M-cone driven responses and also than the L-cone driven responses with smaller stimuli. The M-cone driven responses at full-field and with 70° diameter stimuli displayed similar amplitudes. The L- and M-cone driven responses measured at 8 and 12 Hz were of similar amplitude and approximately in counter-phase. The amplitudes were constant for most stimulus configurations. The results indicate that, when the ERG reflects luminance activity, it is positively correlated with stimulus size. Beyond 35° retinal eccentricity, the retina mainly contains L-cones. Small stimuli are sufficient to obtain maximal ERGs at low temporal frequencies where the ERGs are also sensitive to cone-opponent processing.

Correlations in V1 are reduced by stimulation outside the receptive field.

The trial-to-trial response variability of nearby cortical neurons is correlated. These correlations may strongly influence population coding performance. Numerous studies have shown that correlations can be dynamically modified by attention, adaptation, learning, and potent stimulus drive. However, the mechanisms that influence correlation strength remain poorly understood. Here we test whether correlations are influenced by presenting stimuli outside the classical receptive field (RF) of visual neurons, where they recruit a normalization signal termed surround suppression. We recorded simultaneously the activity of dozens of cells using microelectrode arrays implanted in the superficial layers of V1 in anesthetized, paralyzed macaque monkeys. We presented annular stimuli that encircled--but did not impinge upon--the RFs of the recorded cells. We found that these "extra-classical" stimuli reduced correlations in the absence of stimulation of the RF, closely resembling the decorrelating effects of stimulating the RFs directly. Our results suggest that normalization signals may be an important mechanism for modulating correlations.

Localization of MEG human brain responses to retinotopic visual stimuli with contrasting source reconstruction approaches.

Magnetoencephalography (MEG) allows the physiological recording of human brain activity at high temporal resolution. However, spatial localization of the source of the MEG signal is an ill-posed problem as the signal alone cannot constrain a unique solution and additional prior assumptions must be enforced. An adequate source reconstruction method for investigating the human visual system should place the sources of early visual activity in known locations in the occipital cortex. We localized sources of retinotopic MEG signals from the human brain with contrasting reconstruction approaches (minimum norm, multiple sparse priors, and beamformer) and compared these to the visual retinotopic map obtained with fMRI in the same individuals. When reconstructing brain responses to visual stimuli that differed by angular position, we found reliable localization to the appropriate retinotopic visual field quadrant by a minimum norm approach and by beamforming. Retinotopic map eccentricity in accordance with the fMRI map could not consistently be localized using an annular stimulus with any reconstruction method, but confining eccentricity stimuli to one visual field quadrant resulted in significant improvement with the minimum norm. These results inform the application of source analysis approaches for future MEG studies of the visual system, and indicate some current limits on localization accuracy of MEG signals.

Amblyopic Deficit Beyond the Fovea: Delayed and Variable Single-Trial ERP Response Latencies, but Unaltered Amplitudes

Amblyopia was first described as a deficit of central vision. However, it has long been debated whether this dysfunction is limited to the fovea or whether extrafoveal vision is also affected, as studies concerning the latter are equivocal. The purpose of the study was to resolve this issue. We investigated the amblyopic effect on event-related potentials (ERPs) with foveal and perifoveal stimuli, either matched in size based on cortical magnification or presented as large annular stimuli. In two separate experiments we measured ERPs on amblyopic patients and control subjects using face images. Latency and amplitude of averaged ERPs and their single-trial distributions were analyzed. When the fovea was stimulated, latency and amplitude of the early averaged ERP components increased and were reduced, respectively, in the amblyopic compared with the fellow eye. Importantly, perifoveal stimulation also elicited similar amblyopic deficits, which were clearly significant in the case of using cortical magnification scaled stimuli. However, single-trial peak analysis revealed that foveal and perifoveal effects differed in nature: Peak amplitudes were reduced only in foveal stimulation, while latencies were delayed and jittered at both the fovea and perifovea. Event-related potentials obtained from fellow eyes were not significantly different from those of normal observers. Our findings revealed the existence of amblyopic deficits at the perifovea when the stimulated cortical area was matched in size to that of foveal stimulation. These deficits manifested themselves only in the temporal structure of the responses, unlike foveal deficits, which affected both component amplitude and latency.

Exploiting individual primary visual cortex geometry to boost steady state visual evoked potentials

Objective. The steady-state visual evoked potential (SSVEP) is an electroencephalographic response to flickering stimuli generated partly in primary visual area V1. The typical ‘cruciform’ geometry and retinotopic organization of V1 is such that certain neighboring visual regions project to neighboring cortical regions of opposite orientation. Here, we explored ways to exploit this organization in order to boost scalp SSVEP amplitude via oscillatory summation. Approach. We manipulated flicker-phase offsets among angular segments of a large annular stimulus in three ways, and compared the resultant SSVEP power to a conventional condition with no temporal phase offsets. (1) We divided the annulus into standard octants for all subjects, and flickered upper horizontal octants with opposite temporal phase to the lower horizontal ones, and left vertical octants opposite to the right vertical ones; (2) we individually adjusted the boundaries between the eight contiguous segments of the standard octants condition to coincide with cruciform-consistent, early-latency topographical shifts in pattern-pulse multifocal visual-evoked potentials (PPMVEP) derived for each of 32 equal-sized segments; (3) we assigned phase offsets to stimulus segments following an automatic algorithm based on the relative amplitudes of vertically- and horizontally-oriented PPMVEP components. Main results. The three flicker-phase manipulations resulted in a significant enhancement of normalized SSVEP power of (1) 202%, (2) 383%, and (3) 300%, respectively. Significance. We have thus demonstrated a means to obtain more reliable measures of visual evoked activity purely through consideration of cortical geometry. This principle stands to impact both basic and clinical research using SSVEPs.

Comparative Regional Pupillography as a Noninvasive Biosensor Screening Method for Diabetic Retinopathy

We describe infrared regional pupillometry as an objective comparative assessment of midperipheral to central retinal sensitivity and to correlate with midperipheral retinal ischemia in diabetic subjects. We tested 12 normal and 17 diabetic subjects using bilateral infrared pupillometry. The diabetic cohort included seven subjects without, five with mild, three with moderate, and two with severe non-proliferative diabetic retinopathy (NPDR). Central and annular stimuli of varying intensity were presented to one eye, and pupillary amplitude and constriction velocity were measured from both eyes. Light stimulus of increasing intensity was presented as 20 consecutive trials (stimulus duration of 300 ms with 3000 ms intervals). The ratio of central to peripheral responses (Q values) was calculated for each stimulus configuration. Average responses with respect to the stimulus strength were regressed with Gompertz sigmoid function. Control and moderate/severe NPDR cases comparison showed statistically significant differences in amplitude (Q(A)) and constriction velocity (Q(CV)) (Wilcoxon rank sum test P = 0.002, respectively). Age difference for these groups was not statistically significant (Wilcoxon rank sum test P = 0.15). The comparison of control and diabetic subjects without NPDR/mild NPDR was statistically significant for Q(A) and Q(CV) (Wilcoxon rank sum test P = 0.0002 and P = 0.001, respectively). Q(A) and Q(CV) differences were statistically significant between moderate/severe NPDR cases and subjects without or mild NPDR cases (Wilcoxon rank sum test P = 0.013). Q(A) and Q(CV) values correlated highly with the severity of diabetic retinopathy, but not with the duration of diabetes.

Visual gamma oscillations: The effects of stimulus type, visual field coverage and stimulus motion on MEG and EEG recordings

Increases in the power of neural oscillations in the gamma (>40 Hz) band are a key signature of information processing in cortical neuronal networks. However, non-invasive detection of these very small oscillations is difficult due to the presence of potential artefacts (both muscular and ocular) in the same frequency band and requires highly optimised paradigms. Numerous studies have shown that the properties of visual gamma-band responses to simple pattern stimuli are highly tuned to the stimuli parameters used. The aim of this work was to compare gamma oscillation response properties across some of the more commonly used stimulus configurations. To do this, MEG and EEG recordings were made during the presentation of eight different stimulus types in a 2 × 2 × 2 design. For the first stimulus factor, "Type", the stimulus pattern was either an annulus grating or a square wave grating. For the second stimulus factor, "Field", stimuli were presented in either four visual field quadrants simultaneously or only in the lower left quadrant. Finally, for the "Move" factor, stimuli either drifted at 1.33°s(-1) or were stationary. For MEG gamma band responses, the following main effects were observed, a) gamma-band power was increased for annular stimuli compared to square wave stimuli, b) gamma-band power was increased for full field stimuli compared to single quadrant stimuli and c) gamma-band power was larger for drifting compared to stationary stimuli and were of significantly higher frequency. For the detectors used, the signal to noise ratio was substantially higher for MEG than EEG. The advantages and disadvantages of the different types of stimulus types are discussed.

Suppressive Surrounds of Receptive Fields In Monkey Frontal Eye Field

A critical step in determining how a neuron contributes to visual processing is determining its visual receptive field (RF). While recording from neurons in frontal eye field (FEF) of awake monkeys (Macaca mulatta), we probed the visual field with small spots of light and found excitatory RFs that decreased in strength from RF center to periphery. However, presenting stimuli with different diameters centered on the RF revealed suppressive surrounds that overlapped the previously determined excitatory RF and reduced responses by 84%, on average. Consequently, in that overlap area, stimulation produced excitation or suppression, depending on the stimulus. Strong stimulation of the RF periphery with annular stimuli allowed us to quantify this effect. A modified difference of gaussians model that independently varied center and surround activation accounted for the nonlinear activity in the overlap area. Our results suggest that (1) the suppressive surrounds found in FEF are fundamentally the same as those in V1 except for the size and strength of excitatory and suppressive mechanisms, (2) methodically assaying suppressive surrounds in FEF is essential for correctly interpreting responses to large and/or peripheral stimuli and therefore understanding the effects of stimulus context, and (3) regulating the relative strength of the surround clearly changes neuronal responses and may therefore play a significant part in the neuronal changes resulting from visual attention and stimulus salience.

Topography of Cone Dark Adaptation Deficits in Age-Related Maculopathy

Despite widespread agreement that dark adaptation is abnormal in age-related maculopathy (ARM), the optimal retinal location for detection of this deficit is unclear. We quantified the diagnostic potential of cone dark adaptation as a function of retinal eccentricity and compared this with the diagnostic potential of the time to the rod-cone-break (RCB). Cone dark adaptation was monitored after an 80% cone photopigment bleach in 10 subjects with ARM and 10 age-matched controls, using four achromatic annuli (0.5, 2, 7, and 12° radius) centered on the fovea. Threshold recovery data were modeled and the time constant of cone recovery (τ), final cone threshold, and time to RCB were determined. Diagnostic potential was evaluated by constructing receiver operating characteristic curves for these parameters. Cone τ was significantly longer for the ARM group at 2, 7, and 12°. The greatest difference between groups was observed at 12° from fixation. At this location, the mean τ was 3.49 (±2.02) min and 0.64 (±0.38) min for ARM and control subjects, respectively (p = 0.002), and time to RCB was 17.68 (±5.37) min and 9.05 (±2.11) min for ARM and control subjects, respectively (p = 0.001). Correspondingly, receiver operating characteristic curves showed that the diagnostic potential of dark adaptometry is greatest for stimuli presented 12° from fixation; for cone τ, the area under the curve = 0.99 ± 0.02 and for time to RCB, area under the curve = 0.96 ± 0.04. This study has shown cone-mediated dark adaptation to be significantly impaired in ARM. Our results provide compelling evidence in support of the diagnostic potential of cone dark adaptation and the use of annular stimuli at 12°. The observation that cone τ is highly diagnostic at this eccentricity is significant clinically because this parameter may be quantified within a few minutes.

Mesopic luminance assessed with minimum motion photometry

We measured the relative contribution of rods and cones to luminance across a range of photopic, mesopic, and scotopic adaptation levels and at various retinal eccentricities. We isolated the luminance channel by setting motion-based luminance nulls (minimum motion photometry) using annular stimuli. Luminance nulls between differently colored stimuli require equality in a weighted sum of rod and cone excitations. The relative cone weight increases smoothly from the scotopic range, where rods dominate, to photopic levels, where rod influence becomes negligible. The change from rod to cone vision does not occur uniformly over the visual field. The more peripheral the stimulus location, the higher is the light level required for cones to participate strongly. The relative cone contribution can be described by a sigmoid function of intensity, with two parameters that each depend on the eccentricity and spatial frequency of the stimulus. One parameter determines the "meso-mesopic" luminance--the center of the mesopic range, at which rod and cone contributions are balanced. This increases with eccentricity, reflecting an increase in the meso-mesopic luminance from 0.04 scotopic cd/m(2) at 2° eccentricity to 0.44 scotopic cd/m(2) at 18°. The second parameter represents the slope of the log-log threshold-versus-intensity curve (TVI curve) for rod vision. This parameter inversely scales the width of the mesopic range and increases only slightly with eccentricity (from 0.73 at 2° to 0.78 for vision at 18° off-axis).

L‐ and M‐cone input to 12Hz and 30Hz flicker ERGs across the human retina

We recorded L- and M-cone isolating ERGs from human subjects using a silent substitution technique at temporal rates of 12 and 30 Hz. These frequencies isolate the activity of cone-opponent and non-opponent post-receptoral mechanisms, respectively. ERGs were obtained using a sequence of stimuli with different spatial configurations comprising; (1) circular stimuli of different sizes which increased in 10° steps up to 70°diameter, or (2) annular stimuli with a 70° outer diameter but with different sized central ablations from 10° up to 60°. L- and M-cone isolating ERGs were obtained from five colour normal subjects using a DTL fibre electrode. Fourier analysis of the ERGs was performed and we measured the amplitude of the first harmonic of the response. For 12 Hz ERGs the L:M cone response amplitude ratio (L:M(ERG)) was close to unity and remained stable irrespective of the spatial configuration of the stimulus. The maintenance of this balanced ratio points to the existence of cone selective input across the human retina for the L-M cone opponent mechanism. For 30 Hz the L:M(ERG) ratio was greater than unity but varied depending upon which region of the retina was being stimulated. This variation we consider to be a consequence of the global response properties of M-cone ERGs rather than representing a real variation in L:M cone ratios across the retina.

Nonlinear characterization of a simple process in human vision

Perceptual processes are often modeled as linear filters followed by a decisional rule. This simple model is central to the understanding of visual processing in humans. Its scope may be extended to capture a wider range of behaviors by the addition of nonlinear operators or kernels, but there is no evidence in human sensory processing that these operators are able to enhance the linear description. We focused on a simple process in human vision, the perception of brightness in a center-surround annular stimulus. We used psychophysical reverse correlation to fully characterize this process up to its second-order nonlinearity. The resulting characterization was then used to reconstruct/predict individual responses by the human observers, a process that was significantly enhanced by the addition of the nonlinear kernels. These results provide direct evidence that behavioral second-order kernels can be successfully derived using reverse correlation, and furthermore that they can be effectively exploited to simulate human vision. We show that the former result does not imply the latter by performing a second series of experiments involving orientation-defined textures, for which no measurable benefit was gained from the addition of second-order kernels.

Perceived Color of Hallucinations in the Charles Bonnet Syndrome Is Related to Residual Color Contrast Sensitivity

We sought to determine whether the change in cortical excitability secondary to deafferentation in patients with Charles Bonnet Syndrome (CBS) who hallucinate in a predominant color or combination of colors is related to an alteration in color contrast thresholds and whether the change is specific to the color of the hallucination. We prospectively categorized each patient's hallucinations using the Institute of Psychiatry Visual Hallucinations Interview. We measured color contrast thresholds with a computerized test designed to assess red-green and blue-yellow color confusion axes against a background of luminance noise. We calculated the ratio of red-green threshold to blue-yellow threshold (R-G/B-Y ratio) for each patient. Because central vision was impaired in all patients, we used a sectoral annular stimulus that projected to the retina at 12.5 degrees eccentricity. There were 10 patients with age-related macular degeneration and CBS who were hallucinating in a predominant color or combination of colors at the time of recruitment. Patients hallucinating in red, green, or a combination of red and green had R-G/B-Y ratios of less than 1.0 (n = 5). Patients hallucinating in blue, yellow, or a combination of blue and yellow had R-G/B-Y ratios of greater than 1.0 (n = 2). Patients hallucinating in purple had ratios between the red-green and blue-yellow hallucinators (n = 2). The 1 patient hallucinating in white had the lowest thresholds for red-green and blue-yellow confusion axes. Comparing the R-G/B-Y ratios for the "red/green hallucinators" and "blue/yellow hallucinators" returned a significant result with Fisher's exact test (P = 0.047, n = 7). Deafferentation and secondary cortical hyperexcitability in CBS have a correlate in psychophysical threshold. This change in sensitivity relates specifically to the hallucinated color axis rather than across all colors. This is the first published evidence for cerebral hyperexcitability leading to a decrease in color contrast thresholds.

Measurement of macular pigment optical density and distribution using the steady-state visual evoked potential

The purpose of this study was to specify isoluminance at different retinal eccentricities and to characterize macular pigment optical density (MPOD) and distribution using the steady-state visual evoked potential (VEP). Redgreen (B/G) gratings were generated within two circular stimulus fields (radius = 0.55 or 1.1 deg) and within four annular fields (maximum mean radius = 6.0 deg) on a color monitor. Temporal frequency was 15 Hz. Isoluminance was determined for each stimulus using minimum flicker photometry. Steady-state onset-offset VEPs were recorded to the same annular stimuli as the luminance ratio between adjacent chromatic components was changed from 0.25 to 0.85 in 11 automated steps (0.5 representing photometric isoluminance). Fourier analysis showed that the power of the first harmonic was minimized at the isoluminant ratio specific to each subject. Relative OD was computed by comparing the isoluminant ratio at any location with that for the most eccentric annulus. To compensate for the broadband characteristics of the monitor, OD values were corrected according to minimum flicker measurements made through known concentrations of carotenoid solution. MPOD was additionally measured using minimum motion photometry.There was high correlation between the isoluminant ratios determined by minimum flicker and VEPs for both R/G and B/G stimulation (r = 0.91, P < 0.005, slope = 1). Calibrated OD values computed from VEP estimates of B/G isoluminance correlated with those derived from minimum flicker (r = 0.96, P < 0.0005, slope = 0.85) and motion photometry (r = 0.94, P < 0.0005, slope = 0.88). OD values derived from B/G VEPs increased toward the fovea and corresponded closely with minimum flicker and minimum motion assessment of MP distribution profiles. The steady-state VEP can be used to determine isoluminance at different retinal eccentricities. MPOD and distribution can be measured by steady-state VEPs to B/G stimuli.

Response Facilitation From the “Suppressive” Receptive Field Surround of Macaque V1 Neurons

In primary visual cortex (V1), neuronal responses to optimally oriented stimuli in the receptive field (RF) center are usually suppressed by iso-oriented stimuli in the RF surround. The mechanisms and pathways giving rise to surround modulation, a possible neural correlate of perceptual figure-ground segregation, are not yet identified. We previously proposed that highly divergent and fast-conducting top-down feedback connections are the substrate for fast modulation arising from the more distant regions of the surround. We have recently implemented this idea into a recurrent network model (Schwabe et al. 2006). The purpose of this study was to test a crucial prediction of this feedback model, namely that the suppressive "far" surround of V1 neurons can be facilitatory under conditions that weakly activate neurons in the RF center. Using single-unit recordings in macaque V1, we found iso-orientation far-surround facilitation when the RF center was driven by a low-contrast stimulus and the far surround by a small annular stimulus. Suppression occurred when the center stimulus contrast or the size of the surround stimulus was increased. This suggests that center-surround interactions result from excitatory and inhibitory mechanisms of similar spatial extent, and that changes in the balance of local excitation and inhibition, induced by surround stimulation, determine whether facilitation or suppression occurs. In layer 4C, the main target of geniculocortical afferents, lacking long-range intra-cortical connections, far-surround facilitation was rare and large surround fields were absent. This strongly suggests that feedforward connections do not contribute to far-surround modulation and that the latter is generated by intra-cortical mechanisms, likely involving top-down feedback.

Macular pigment and the edge hypothesis of flicker photometry

Heterochromatic flicker photometry is commonly used to measure macular pigment optical density (MPOD) in the human retina. It has been proposed, and accepted by many, that the MPOD so measured represents the value at a retinal location corresponding to the edge of the flickering, circular stimulus. We have investigated this proposal by using a series of annular stimuli to determine the MPOD distribution in the central 1.5° of the retina for both eyes of 10 subjects. The MPOD obtained using a 1.5° circular stimulus matched the MPOD distribution at a retinal eccentricity that was always less than the stimulus radius, and averaged, for the 10 subjects, 51% of the stimulus radius. Similar results were obtained using a 1° stimulus. Thus the edge hypothesis is inconsistent with our data. We suggest that involuntary eye movements may be responsible for an apparent edge effect.