Recent reports suggest the ON and OFF pathways are differentially susceptible to selective vision loss in glaucoma. Thus, perimetric assessment of ON- and OFF-pathway function may serve as a useful diagnostic. However, this necessitates a developed understanding of normal ON/OFF pathway function around the visual field and as a function of input intensity. Here, using electroencephalography, we measured ON- and OFF-pathway biased contrast response functions in the upper and lower visual fields. Using the steady-state visually evoked potential paradigm, we flickered achromatic luminance probes according to a saw-tooth waveform, the fast phase of which biased responses towards the ON or OFF pathways. Neural responses from the upper and lower visual fields were simultaneously measured using frequency tagging - probes in the upper visual field modulated at 3.75 Hz, while those in the lower visual field modulated at 3 Hz. We find that responses to OFF/decrements are larger than ON/increments, especially in the lower visual field. In the lower visual field, both ON and OFF responses were well described by a sigmoidal non-linearity. In the upper visual field, the ON pathway function was very similar to that of the lower, but the OFF pathway function showed reduced saturation and more cross-subject variability. Overall, this demonstrates that the relationship between the ON and OFF pathways depends on the visual field location and contrast level, potentially reflective of natural scene statistics.

Previous studies have shown that with the use of tinted lenses (or colored filters), individuals with red–green color vision deficiency (CVD) report an improvement in their performance on certain color vision tests. In this context, this study examines the effects of a digitally generated red-colored filter and identifies the mechanism mainly responsible for the changes in red–green CVD observers’ performance on a D-15 arrangement test performed using the filter. We simulate the red filter digitally with the spectral transmittance similar to that of the X-Chrom, which is a red-tinted lens. Fourteen red–green CVD subjects are subjected to the D-15 test on a computer monitor under four filter conditions, consisting of one condition without the filter and three conditions with the filter, corresponding to the opacity of the red filter. The results show that while the simulated red filter improves the performance of deutans to arrange the caps in the D-15 test, this is not the case for protans. In addition, considerations based on the human cone-contrast model enable us to identify that the improvement in deutan observers largely results from the increase in the luminance contrast between stimuli and a background. To summarize, the red filter simulated in this study induces different changes in the red–green CVD observer luminance contrast between the protan and deutan types, with the result that the performance of deuteranopes improves while that of protanopes deteriorates.

The retina of the zebrafish (Danio rerio) provides an unusually favorable preparation for genetic and developmental studies of the retina. Although the retina has been studied extensively for two decades, the neuronal response of the inner retina is largely unknown. This report describes a prominent local field potential of the inner retina, the Proximal Negative Response (PNR). It is best evoked by small (100 μm) precisely positioned spots of light and is exceedingly sensitive to negative luminance contrast. The polarity, waveform, and other properties of the PNR suggest that it arises primarily from ON-OFF neurons of the proximal retina. The dominant response to negative contrast and its enhancement by light adaptation is believed due to a dominant presynaptic input from OFF bipolar cells. Color contrast was investigated by analyzing responses to a green bar moving on green versus red backgrounds. Over an intermediate range of irradiance, the response to green on red was larger than the response to green on green, thereby providing evidence for the encoding of color contrast. The present findings complement the classic principle of color contrast for human vision known as Kirschmann’s third law and bring to mind the view of Walls that color contrast may have been the driving force for the evolution of color vision in lower vertebrates. In sum, the PNR of zebrafish provides clear evidence for the encoding of color and luminance contrast in the inner retina. It exhibits the defining properties common to many other vertebrates, reinforcing the view that the zebrafish may further serve as a model for retinal function and that the PNR may provide a new approach for studies of development, genetics, and retinal degeneration in zebrafish.

Full-field electroretinograms were recorded from five normal human subjects using white light (mean luminance: 250 cd/m2) sine wave stimuli at different frequencies and contrasts. In agreement with previous studies, we found that the amplitude of the fundamental component displayed a dip at about 12 Hz, coinciding with a maximum in the second harmonic component, indicating frequency doubling of the responses. By including measurements at different contrasts, we were able to recognize two (sine-like and transient) response components. We found that the waveform of the transient response was relatively frequency independent. An algorithm to separate the two components was developed. The interaction between these two components can explain the frequency-doubled responses around 12 Hz. The sine-like component is more linear and prominent in the low-frequency region, whereas the transient seems to be more nonlinear and prominent in the high-frequency region.

Saccadic reaction times (SRTs) of three human subjects were analyzed. The gap paradigm was used (i.e. fixation point offset precedes target onset) to obtain high proportions of express saccades (i.e. saccades of extremely short reaction times) in the SRT distributions. In one set of experiments, the luminance of the (red) saccade target was varied from brighter to darker than the (green) background including an isoluminance condition. Express saccades were obtained in response to pure color contrast stimuli with about the same frequency and reaction time as to stimuli with both color and luminance contrast. In a second experiment, the luminance contrast of a white target on a white background was lowered below 10%. Again the number of express saccades was not reduced. Thus, in contrast to other perceptual phenomena the visual neural mechanisms underlying the generation of express saccades are not affected by isoluminance nor low contrast luminance.

Is neural binding of color and form required for perception of a unified colored object? Individual cells selectively tuned to both color and orientation are proposed to moot the binding problem. This study reveals perceptual misbinding of color, thereby revealing separate neural representations of color and form followed by a subsequent binding process. Low luminance-contrast, rivalrous chromatic gratings were presented dichoptically. Each grating had alternating chromatic and gray stripes (e.g., red/gray in the left eye, green/gray in the right eye). Observers viewed the two rivalrous, 2 cpd gratings for 1 min. The duration of exclusive visibility was measured for four percepts: left-eye stimulus, right-eye stimulus, fusion of the two colors, or a two-color grating (e.g. a red/green grating). The percept of a two-color grating (misbinding) was observed with Michelson luminance contrast in the grating up to 20%. In general, for a given level of luminance contrast either misbinding (low luminance contrast) or color mixture (high luminance contrast) was observed, but not both of them. The perceived two-color gratings show that two rivalrous chromaticities are both represented neurally when color and form are combined to give a unified percept. “Resolution” of competing chromatic signals from the two eyes is not restricted to color dominance and color mixture. The transition from misbinding to color mixture by increasing luminance contrast shows that luminance edges have an important role in correct localization of color.

Human visual evoked potentials (VEPs) were recorded for abrupt reversals of 2 cycles/deg (c/deg) square-wave gratings combining high red–green contrast with different levels of luminance contrast. Response characteristics—2nd harmonic amplitudes and peak latencies as a function of luminance contrast—were compared for four different reversal rates ranging from 6.25 Hz to 12.5 Hz. At every reversal frequency, the VEP amplitude and latency plots were nonsymmetrical with respect to isoluminance. The amplitude dropped to a minimum within a region of rapid phase change, always at a red–green luminance contrast for which the green color had the higher luminance, at about 40% or 50% Michelson luminance contrast. The rapid phase shift around this contrast suggested a sudden change in the relative impact of VEP generators with different latencies, possibly dominated by parvocellular or magnocellular input. The most prominent VEP waveform through most of the luminance contrast range, P110, is interpreted in terms of a parvo-mediated response that is attenuated with increasing reversal frequency. Contrast-dependent changes in the P110 amplitude appear to be responsible for the VEP asymmetries reported here.

Human visual evoked potentials (VEPs) were recorded for abrupt 6.25-Hz reversals of 2 c/deg square-wave gratings combining red–green contrast with different levels of luminance contrast. Response characteristics— amplitudes and peak latencies as a function of luminance contrast—were compared for four different pairs of red–green colors and an isochromatic yellow grating. For each of the red–green color pairs, the plots of VEP amplitudes and latencies were nonsymmetrical with respect to isoluminance. The amplitude dropped to a minimum within a region of rapid phase change, at a different contrast for each color pair but always at a luminance contrast for which the greener color had the higher luminance. When the contrast-response curve for each of the four red–green pairs was modeled by a simple |CL − CM| opponency of L- and M-cone contrast using a fixed CL/CM weighting ratio of about two, there was a close correspondence between the contrast giving a null in the modeled response and that giving a minimum in the VEP amplitude. So for the stimulus parameters applied here, the reversal VEP appeared to be dominated by L/M-opponent response contributions for which the signed CL/CM-cone weighting ratio was close to a value of minus two rather than to a value of minus one, which is characteristic of the psychophysical red–green detection mechanism and representative of CL/CM weighting ratios for precortical cells in the parvocellular pathway.

To investigate the influence of voltage-sensitive conductances in shaping light-evoked responses of retinal bipolar cells, whole-cell recordings were made in the slice preparation of the tiger salamander, Ambystoma tigrinum. To study contrast encoding, the retina was stimulated with 0.5-s steps of negative and positive contrasts of variable magnitude. In the main, responses recorded under voltage- and current-clamp modes were remarkably similar. In general agreement with past results in the intact retina, the contrast/response curves were relatively steep for small contrasts, thus showing high contrast gain; the dynamic range was narrow, and responses tended to saturate at relatively small contrasts. For ON and OFF cells, linear regression analysis showed that the current response accounted for 83–93% of the variance of the voltage response. Analysis of specific parameters of the contrast/response curve showed that contrast gain was marginally higher for voltage than current in three of four cases, while no significant differences were found for half-maximal contrast (C50), dynamic range, or contrast dominance. In sum, the overall similarity between current and voltage responses indicates that voltage-sensitive conductances do not play a major role in determining the shape of the bipolar cell's contrast response in the light-adapted retina. The salient characteristics of the contrast response of bipolars apparently arise between the level of the cone voltage response and the postsynaptic current of bipolar cells, via the transformation between cone voltage and transmitter release and/or via the interaction between the neurotransmitter glutamate and its postsynaptic receptors on bipolar cells.

Intracellular recordings were obtained from 73 cone-driven bipolar cells in the light-adapted retina of the tiger salamander (Ambystoma tigrinum). Responses to flashes of negative and positive contrast for centered spots and concentric annuli of optimum spatial dimensions were analyzed as a function of contrast magnitude. For both depolarizing and hyperpolarizing bipolar cells, it was found that remarkably similar responses were observed for the center and surround when comparisons were made between responses of the same response polarity and thus, responses to opposite contrast polarity. Thus, spatial information and contrast polarity appear to be rather strongly confounded in many bipolar cells. As a rule, the form of the contrast/response curves for center and surround approximated mirror images of each other. Contrast gain and C50 (the contrast required for half-maximal response) were quantitatively similar for center and surround when comparisons were made for responses of the same response polarity. The average contrast gain of the bipolar cell surround was 3–5 times higher than that measured for horizontal cells. Contrast/latency measurements and interactions between flashed spots and annuli showed that the surround response is delayed by 20–80 ms with respect to that of the receptive-field center. Cones showed no evidence for center-surround antagonism while for bipolar cells, the average strength of the surround ranged from about 50% to 155% of the center, depending on the test and response polarity. The results of experiments on the effects of APB (100 μM) on depolarizing bipolar cells suggest that the relative contribution of the feedback pathway (horizontal cell to cones) and the feedforward pathway (horizontal cell to bipolar cell) to the bipolar surround varies in a distributed manner across the bipolar cell population.

We report VEP studies which delineate interactions between chromatic and luminance contrast signals. We examined responses to sinusoidal luminance gratings undergoing 4-Hz square-wave contrast reversal, upon which standing gratings with various admixtures of luminance and chromatic contrast were alternately superimposed and withdrawn. The presence of the standing grating induced a VEP component at the fundamental frequency of the contrast-reversal grating. This VEP component appeared without any appreciable lag, and did not vary in amplitude over the 4 s during which the standing grating was present. The observed fundamental response differed from the fundamental component that would be expected from the known interaction between the luminance component of the standing grating with the modulated grating (Bodis-Wollner et al., 1972; Bobak et al., 1988), in three ways: (1) The fundamental response was not nulled for standing gratings that were isoluminant or near-isoluminant. (2) The chromatic dependence of the fundamental response implied an S-cone input to the interaction. (3) No single mechanism (driven by a linear combination of cone signals) could account quantitatively for the size of this response, particularly when the standing grating strongly modulated two cones in phase.

The impulse discharge of single ganglion cells was recorded extracellularly in superfused eyecup preparations of the tiger salamander (Ambystoma tigrinum). Contrast flashes (500 ms) were applied at the center of the receptive field while the retina was light adapted to a background field of 20 cd/m2. The incidence of cell types in a sample of 387 cells was: ON cells (4%), OFF cells (28%), and ON/OFF cells (68%). Quantitative contrast/response measurements were obtained for 83 cells. On the basis of C50, the contrast necessary to evoke a half-maximal response, ON/OFF cells fell into 3 groups: (1) Positive Dominant (26%), (2) Balanced (23%), and (3) Negative Dominant (51%). Positive Dominant cells tended to be relatively contrast insensitive. On the other hand, many Negative Dominant cells showed remarkably low C50 values and very steep contrast/response curves. Contrast gain to negative contrast averaged 8.5 impulses/s/% contrast, some four times greater than that evoked by positive contrast. In most ON/OFF cells, the latency of the first spike evoked by a negative contrast step was much shorter (40–100 ms) than that evoked by a positive contrast step of equal contrast. OFF cells typically showed higher C50 values, larger dynamic ranges, and longer latencies than those of Negative Dominant ON/OFF cells. Thus, different pathways or mechanism apparently mediate the off responses of OFF and ON/OFF cells. In sum, the light-adapted retina of the tiger salamander is strongly biased in favor of negative contrast, as shown by the remarkably high contrast sensitivity and faster response of Negative Dominant cells, the remarkably low incidence of ON cells, and the insensitivity of Positive Dominant cells. Some possible underlying influences of bipolar and amacrine cells are discussed.