Kathy T. Mullen

The contrast sensitivity of human colour vision to red‐green and blue‐yellow chromatic gratings.

A method of producing red‐green and blue‐yellow sinusoidal chromatic gratings is used which permits the correction of all chromatic aberrations. A quantitative criterion is adopted to choose the intensity match of the two colours in the stimulus: this is the intensity ratio at which contrast sensitivity for the chromatic grating differs most from the contrast sensitivity for a monochromatic luminance grating. Results show that this intensity match varies with spatial frequency and does not necessarily correspond to a luminance match between the colours. Contrast sensitivities to the chromatic gratings at the criterion intensity match are measured as a function of spatial frequency, using field sizes ranging from 2 to 23 deg. Both blue‐yellow and red‐green contrast sensitivity functions have similar low‐pass characteristics, with no low‐frequency attenuation even at low frequencies below 0.1 cycles/deg. These functions indicate that the limiting acuities based on red‐green and blue‐yellow colour discriminations are similar at 11 or 12 cycles/deg. Comparisons between contrast sensitivity functions for the chromatic and monochromatic gratings are made at the same mean luminances. Results show that, at low spatial frequencies below 0.5 cycles/deg, contrast sensitivity is greater to the chromatic gratings, consisting of two monochromatic gratings added in antiphase, than to either monochromatic grating alone. Above 0.5 cycles/deg, contrast sensitivity is greater to monochromatic than to chromatic gratings.

Human peripheral spatial resolution for achromatic and chromatic stimuli: limits imposed by optical and retinal factors.

1. The aim of this study was to determine whether optical, receptoral or higher‐order neural properties limit spatial resolution (acuity) in human vision, especially in the peripheral regions of the visual field. 2. Both achromatic and chromatic stimuli were used, and measures were taken to ensure that the resolution estimates were not contaminated by the detection of spatial sampling artifacts. Spatial contrast sensitivity functions were measured at retinal locations from 0 to 55 deg along the naso‐temporal meridian for: (i) discriminating the direction of drift of luminance‐modulated (black‐white) sinusoidal stimuli drifting at 8 Hz (achromatic task); and (ii) for detecting isoluminant red‐green sinusoidal stimuli drifting at 0.4 Hz (chromatic task). Achromatic contrast sensitivity functions were also measured along the vertical meridian for eccentricities of 8 and 40 deg. Each achromatic function was extrapolated to a contrast sensitivity of one (100% contrast) to estimate achromatic acuity. Chromatic acuities were obtained by expressing chromatic contrast in terms of cone contrasts and using the same method of extrapolation. We compared the results with recent data on human optical properties and retinal anatomy. 3. Both achromatic and chromatic acuity decline with distance from the fovea, but at a faster rate than that dictated by the known optical and/or receptoral properties of the human eye. We conclude that, for stimuli of either achromatic or chromatic contrast, peripheral spatial resolution is limited by post‐receptoral mechanisms. Also, chromatic acuity declines more steeply than luminance acuity with eccentricity suggesting that there are additional post‐receptoral limitations on colour resolution in the periphery. 4. A clear naso‐temporal asymmetry is seen in the resolution whose dependence is qualitatively, but not quantitatively, similar to the Nyquist limits imposed by the asymmetric density of human retinal ganglion cells. We discuss the possibility that in peripheral vision (beyond the optic nerve head) the spacing of ganglion cells may pose a fundamental limit on the resolution of achromatic stimuli, but not chromatic stimuli.

Tolerance to visual defocus

Low-resolution optical systems are more tolerant to defocus than are high-resolution systems. We wished to determine whether this principle applies to human vision. We used psychophysical methods to measure the effects of defocus in normal eyes under low-resolution conditions. Modulation transfer of sine-wave gratings was measured as a function of dioptric defocus at low and medium spatial frequencies. We defined the depth of focus at a given spatial frequency to be the dioptric range for which the modulation transfer exceeds 50% of its peak value. For dilated pupils, depth of focus increased from about 2.5 diopters (D) at 3.5 cycles/deg to about 17 D at 0.25 cycles/deg. From our results we predicted that tasks requiring only low spatial frequencies will be more tolerant to defocus than tasks requiring higher spatial frequencies. This prediction was confirmed in a letter-recognition experiment. The increasing tolerance to defocus at low spatial frequencies also implies that individuals with low acuity will be more tolerant to defocus than people with normal vision. We confirmed this prediction by measuring tolerance to defocus in 30 low-vision eyes.

Colour vision as a post-receptoral specialization of the central visual field

The experiments address the question whether there is evidence that the central visual field is any more specialized for colour than it is for luminance contrast detection. The decline in contrast sensitivity across the visual field for colour-only (red-green) gratings is compared to that for monochromatic luminance gratings at a range of spatial frequencies in the nasal and temporal fields. Measurements are made of the chromatic spatial summation area and the relevant parts of the chromatic temporal contrast sensitivity function at different eccentricities in order to control for their influence on the decline in contrast sensitivity. Results show that at each spatial frequency colour contrast sensitivity declines with eccentricity approximately twice as steeply as luminance contrast sensitivity. The more rapid decline in colour contrast sensitivity than luminance contrast sensitivity across the visual field reveals that chromatic mechanisms are more confined than luminance mechanisms to the central field.

Wavelength discrimination at detection threshold

The experiments that we report aim to elucidate the linkage between cone outputs and color sensation. This is investigated by measuring wavelength discrimination between stimuli at threshold levels of detection. Stimuli are large spots (0.75 deg) presented on a white background. A 2 x 2 alternate forced choice method is used to measure simultaneously the detection of different wavelengths and discrimination between them. This method reveals at least four distinguishable colors, indicating the presence of four different sets of mechanisms at threshold. These are associated with the color sensations of orange, pale yellow, green, and blue. There is also evidence for a fifth imperfectly distinguished color (violet) in the shortest wavelength region. Results show that the boundaries between the distinguishable colors have little variation in their spectral positions. This is compatible with the presence of fixed perceptual boundaries in the spectrum dividing the different types of detection mechanism. The correspondence of the spectral locations of the distinguishable colors to the cone opponent responses revealed in the spectral sensitivity function suggests that these color sensations are postreceptoral in origin, arising from different combinations of the three cone outputs.

Postreceptoral chromatic detection mechanisms revealed by noise masking in three-dimensional cone contrast space

We used a noise masking technique to test the hypothesis that detection is subserved by only two chromatic postreceptoral mechanisms (red-green and blue-yellow) and one achromatic (luminance) mechanism. The task was to detect a 1-c/deg Gaussian enveloped grating presented in a mask of static, spatially low-passed binary or Gaussian distributed noise. In the main experiment, the direction of the test stimulus (termed the signal) was constant in cone contrast space, and the direction of the noise was sampled in equally spaced directions within a plane (the noise plane) in the space. The signal was chosen to coincide with one of the three cardinal directions of three postulated mechanisms. The noise plane was selected to span two of the cardinal directions, including that chosen as the signal direction. As the noise direction was sampled around the noise plane, the signal detection threshold was found to vary in accordance with a linear cosine model, which predicted noise directions yielding maximum and minimum masking of the signal. In the direction of minimum masking (termed a null direction), the noise was found to have no masking effect on the signal. Moreover, the null was not orthogonal to the signal direction but lay instead in one of the cardinal directions. Our findings suggest that detection is mediated by only three mechanisms. In a further experiment we found little or no cross masking between each pair of cardinal directions up to the limit of our noise mask contrasts. This further supports the presence of no more than three independent postreceptoral mechanisms.

Differential distributions of red–green and blue–yellow cone opponency across the visual field

The color vision of Old World primates and humans uses two cone-opponent systems; one differences the outputs of L and M cones forming a red-green (RG) system, and the other differences S cones with a combination of L and M cones forming a blue-yellow (BY) system. In this paper, we show that in human vision these two systems have a differential distribution across the visual field. Cone contrast sensitivities for sine-wave grating stimuli (smoothly enveloped in space and time) were measured for the two color systems (RG & BY) and the achromatic (Ach) system at a range of eccentricities in the nasal field (0-25 deg). We spatially scaled our stimuli independently for each system (RG, BY, & Ach) in order to activate that system optimally at each eccentricity. This controlled for any differential variations in spatial scale with eccentricity and provided a comparison between the three systems under equivalent conditions. We find that while red-green cone opponency has a steep decline away from the fovea, the loss in blue-yellow cone opponency is more gradual, showing a similar loss to that found for achromatic vision. Thus only red-green opponency, and not blue-yellow opponency, can be considered a foveal specialization of primate vision with an overrepresentation at the fovea. In addition, statistical calculations of the level of chance cone opponency in the two systems indicate that selective S cone connections to postreceptoral neurons are essential to maintain peripheral blue-yellow sensitivity in human vision. In the red-green system, an assumption of cone selectivity is not required to account for losses in peripheral sensitivity. Overall, these results provide behavioral evidence for functionally distinct neuro-architectural origins of the two color systems in human vision, supporting recent physiological results in primates.

Absence of smooth motion perception in color vision.

We have tested the behavioral evidence for a separation of the processing of color contrast from motion in the human visual system. Two different aspects of motion perception are examined; the identification of the direction of movement of a chromatic grating and the perception of smooth motion. The results show that color vision is at no great disadvantage in the identification of direction of movement, since this can be done at color contrasts quite close to detection threshold over a wide range of spatial and temporal frequencies. However, we find that subjects can identify direction without having the genuine perception of smooth motion. Smooth motion perception is revealed to be highly impaired since it is detected only at very high color contrasts and over a narrow range of spatial temporal conditions.

Estimation of the L-, M-, and S-cone weights of the postreceptoral detection mechanisms

We have obtained two- and three-dimensional detection threshold contours in cone contrast space for sinusoidal gratings for three subjects at three spatiotemporal conditions (1 cycle/degree (c/deg), 0 Hz; 0.125 c/deg, 0 Hz; 1 c/deg, 24 Hz). These conditions were chosen to favor the response of each of the three postreceptoral mechanisms in turn. Contours were obtained from measurements in as many as 60 axes in (L, M, S) cone contrast space and were fitted by superellipses. Our technique permitted us to improve on earlier estimates of the cone weightings to the mechanisms. We found that the red–green mechanism has an input cone weighting of L−M with a 2% S-cone input; the luminance mechanism has a weighting of kL + M, where k varies between 3 and 5 at the high-temporal condition, with a 5% S-cone input in opposition to L- and M-cones; and the blue–yellow mechanism consists of S inputs in closely balanced opposition to L and M inputs. These cone weights were found to be consistent among our three subjects.

Deficient responses from the lateral geniculate nucleus in humans with amblyopia

Amblyopia or lazy eye is the most common cause of uniocular blindness in adults. It is caused by a disruption to normal visual development as a consequence of unmatched inputs from the two eyes in early life, arising from a turned eye (strabismus), unequal refractive error (anisometropia) or form deprivation (e.g. cataract). Animal models based on extracellular recordings in anesthetized animals suggest that the earliest site of the anomaly in the primate visual pathway is the primary visual cortex (corresponding to the striate cortex, cytoarchitectonic area 17 and area V1), which is where inputs from the two eyes are first combined in an excitatory fashion, whereas more distal and monocular processing structures such as the retina and lateral geniculate nucleus (LGN) are normal. Using high‐field functional magnetic resonance imaging in a group of human adults with amblyopia, we demonstrate that functional deficits are first observable at a thalamic level, that of the LGN. Our results suggest the need to re‐evaluate the current models of amblyopia that are based on the assumption of a purely cortical dysfunction, as well as the role for the LGN in visual development.