Donald I. A. MacLeod

Chromaticity diagram showing cone excitation by stimuli of equal luminance.

In a space where Cartesian coordinates represent the excitations of the three cone types involved in color vision, a plane of constant luminance provides a chromaticity diagram in which excitation of each cone type (at constant luminance) is represented by a linear scale (horizontal or vertical), and in which the center-of-gravity rule applies with weights proportional to luminance.

Spectral sensitivities of the human cones

Transient chromatic adaptation produced by an abrupt change of background color permits an easier and closer approach to cone isolation than does steady-state adaptation. Using this technique, we measured middle-wave-sensitive (M)-cone spectral sensitivities in 11 normals and 2 protanopes and long-wavelength-sensitive (L-) cone spectral sensitivities in 12 normals and 4 deuteranopes. Although there is great individual variation in the adapting intensity required for effective isolation, there is little variation in the shape of the M- and L-cone spectral-sensitivity functions across subjects. At middle and long wavelengths, our mean spectral sensitivities agree extremely well with dichromatic spectral sensitivities and with the M- and L-cone fundamentals of Smith and Pokorny [Vision Res. 15, 161 (1975)] and of Vos and Walraven [Vision Res. 11, 799 (1971)], both of which are based on the CIE (Judd-revised) 2 degrees color-matching functions (CMFs). But the agreement with the M-cone fundamentals of Estévez [Ph.D. dissertation, Amsterdam University (1979)] and of Vos et al. [Vision Res. 30, 936 (1990)], which are based on the Stiles-Burch 2 degrees CMFs, is poor. Using our spectral-sensitivity data, tritanopic color-matching data, and Stiles pi 3, we derive new sets of cone fundamentals. The consistency of the proposed fundamentals based on either the Stiles-Burch 2 degrees CMFs or the CIE 10 degrees large-field CMFs with each other, with protanopic and deuteranopic spectral sensitivities, with tritanopic color-matching data, and with short-wavelength-sensitive (S-) cone spectral-sensitivity data suggests that they are to be preferred over fundamentals based on the CIE 2 degrees CMFs.

Long-term deprivation affects visual perception and cortex

Recovery after long-term blindness was first studied in 1793, but few cases have been reported since. We combined psychophysical and neuroimaging techniques to characterize the effects of long-term visual deprivation on human cortex.

Visual adaptation and face perception

The appearance of faces can be strongly affected by the characteristics of faces viewed previously. These perceptual after-effects reflect processes of sensory adaptation that are found throughout the visual system, but which have been considered only relatively recently in the context of higher level perceptual judgements. In this review, we explore the consequences of adaptation for human face perception, and the implications of adaptation for understanding the neural-coding schemes underlying the visual representation of faces. The properties of face after-effects suggest that they, in part, reflect response changes at high and possibly face-specific levels of visual processing. Yet, the form of the after-effects and the norm-based codes that they point to show many parallels with the adaptations and functional organization that are thought to underlie the encoding of perceptual attributes like colour. The nature and basis for human colour vision have been studied extensively, and we draw on ideas and principles that have been developed to account for norms and normalization in colour vision to consider potential similarities and differences in the representation and adaptation of faces.

Equiluminance: spatial and temporal factors and the contribution of blue-sensitive cones

Equiluminance ratios for red/green, red/blue and green/blue sine-wave gratings were determined by using a minimum-motion heterochromatic matching technique that permitted reliable settings at temporal frequencies as low as 0.5 Hz. The red/green equiluminance ratio was influenced by temporal but not spatial frequency, the green/blue ratio was influenced by spatial but not temporal frequency, and the red/blue ratio was influenced by both. After bleaching of the blue-sensitive cones, there was no change in equiluminance ratios, indicating no contribution of the blue-sensitive cones to the luminance channel even at low temporal and spatial frequencies. The inhomogeneity of yellow pigmentation within the macular region was identified as the source of the spatial-frequency effect on the blue/green ratio.

Punctate sensitivity of the blue-sensitive mechanism

Abstract Thresholds were measured for a tiny, brief, violet flash on a long wavelength, B cone-isolating background in foveal locations spaced only 4 or 5′ of arc apart. Large spatial variations in B cone sensitivity were found just beyond the foveal tritanopic area even though thresholds for the same wavelength test flash hardly varied at all across these same retinal locations when the flash was detected by G cones. The relative constancy of G cone threshold suggests that these spatial variations are intrinsic to the blue-sensitive mechanism and cannot be explained by prereceptoral filtering. The spatial variations in B cone sensitivity are consistent with physiological evidence that B cones are scarce in the retina. In one observer, it was possible to discern discrete peaks in sensitivity spaced roughly 10′ of arc apart. A model is described which takes optical spread and eye movements into account to show that these peaks may represent individual B cones (or clumps of B cones).

A visual nonlinearity fed by single cones

An intensive nonlinearity in the visual system can produce distortion products, or difference frequency gratings, when observers view two high contrast, high spatial frequency interference fringes of slightly different frequency or orientation added together at the retina. These distortion products are visible even when the two fringes imaged on the retina are above the resolution limit. Our experiments take advantage of this nonlinearity to measure the spatial filtering in the visual system following the formation of the retinal image, but preceding the site of the nonlinearity. The point spread function corresponding to this spatial filter is so small that it can be entirely explained by light integration within the apertures of foveal and parafoveal cones. The small size of this point spread function implies that (1) laser interferometry avoids contrast losses inherent in the eyes optics at spatial frequencies as high as 130 c/deg, (2) retinal scatter causes negligible image degradation in the fovea and parafoveal retina, (3) eye movements have little or no effect on contrast sensitivity to the distortion product and (4) that there is no neural spatial summation in the visual system prior to the site of the nonlinearity. Distortion products could also be observed when a bright interference fringe was briefly flashed on the fovea and a test interference fringe was viewed through the resulting afterimage. Measurements of the point spread function at stages in the visual system that precede the generation of this distortion product were similar to those obtained with simultaneous presentation of the two fringes, implying that the aftereffect of light adaptation is extremely local, no larger than the dimensions of single cones.

Blue-sensitive cones do not contribute to luminance

By using violet backgrounds we selectively altered blue-cone sensitivity but found no change in flicker photometric sensitivity. This indicates that blue cones do not contribute to luminance as defined by flicker photometry.

The temporal properties of the human short-wave photoreceptors and their associated pathways

Flicker modulation sensitivity measurements made on high intensity orange steady backgrounds indicate that signals from short-wavelength sensitive cones (S-cones) have access to two pathways. At low S-cone adaptation levels the frequency response falls quickly with increasing frequency, but at higher adaptation levels it extends to much higher frequencies. At these higher S-cone adaptation levels, the following procedures can selectively expose either a process sensitive to low frequencies or one more sensitive to higher frequencies: (1) at high flicker frequencies, the S-cone signal can be nulled by a long-wavelength sensitive cone (L-cone) signal of suitable amplitude and phase, but at low frequencies a residual flicker persists; the modulation sensitivity for the residual flicker is lowpass in shape with a rapid decline in sensitivity with increasing flicker frequency; (2) sensitivity to flicker in the presence of a 17 Hz S- or L-cone mask is also lowpass with a similarly steep loss of high frequency sensitivity; yet (3) sensitivity to flicker during transient stimulation of the S-cones at 0.5 Hz is comparatively wideband (and slightly bandpass) in shape. The S-cone signal produced by the high frequency process is almost as well-maintained towards high frequencies as M- and L-cone signals. Furthermore, it is capable of participating in flicker photometric nulls with M- and L-cone signals. At low frequencies, however, when the low frequency S-cone signal is also present, satisfactory nulls can not be found. From these and phenomenological considerations, we identify the low and high frequency S-cone processes as S-cone inputs to the chromatic and luminance pathways, respectively. The phase adjustments needed to optimize flicker photometric nulls reveal that the S-cone input to the luminance pathway is actually inverted, but this is demonstrable only at relatively low frequencies: at medium or high frequencies the S-cone influence can be synergistic with that of the other cone types because of a delay in the transmission of S-cone signals.

Flicker photometric study of chromatic adaptation: selective suppression of cone inputs by colored backgrounds

Flicker photometric equivalence is both additive and transitive when the test and standard are alternated upon a relatively more intense colored background. When the balance of red versus green cone excitation from the background is unequal, the contribution of one cone type to flicker photometric spectral sensitivity may be depressed in relation to that of the other by at least 1 order of magnitude more than Webers law predicts. The resultant spectral sensitivity is determined predominantly by only one class of cone. The cone spectral sensitivities of normals are then seen to be the same as those of dichromats, although there is some individual variation. A model is developed to explain this surprising phenomenon.