Does L/M cone opponency disappear in human periphery?

We have assessed the optimal cone contrast sensitivity across eccentricity in human vision of the two cone-opponent mechanisms [L/M or red-green, and S/(L + M) or blue-yellow] and the luminance mechanism. We have used a novel stimulus, termed a 'sinring', that is a radially modulated sine-wave arc, Gaussian enveloped in both angular and radial directions. This stimulus overcomes the problem inherent in Gabor stimuli of confounding stimulus spatial frequency, size, and eccentricity and so allows contrast sensitivity to be tracked accurately into the periphery. Our results show that L/M cone opponency declines steeply across the human periphery and becomes behaviourally absent by 25-30 deg (in the nasal field). This result suggests that any L/M cone-opponent neurons found in primate peripheral retina beyond this limit are unlikely to be significant for colour contrast detection measured behaviourally.     

A ratio principle for a red/green and a yellow/blue channel?

There is strong empirical evidence that, under adaptation to another achromatic color stimulus, the lightness of an achromatic color stimulus depends on the ratio of the luminances of the two stimuli. In the present study, the suitability of this ratio principle is tested for two chromatic postreceptoral opponent channels. A Hering red/green channel and a non-Hering yellow/blue channel are specified as chromatic channels. The yellow/blue channel is defined by extrapolating the plane corresponding to unique green-white linearly to the reddish part of color space, using the plane’s surface as the channel’s equilibria. The experiment was run on an isoluminant plane, measured individually for each observer. Moving along an observer’s measured opponent axes, eight adaptation stimuli were selected for each channel and spanned the whole range of the channel’s coordinates. Red/green equilibria or yellow/blue equilibria were measured as excursions along the adaptation axes. For both presumed channels, the ratios of the equilibrium coordinates of test and adaptation stimuli were essentially constant. This supports the principle’s suitability. However, small asymmetries were found with respect to each channel’s opponent hues. The status of the proposed yellow/blue channel is discussed, as are conditions that might have favored the present findings.     

Biological components of colour preference in infancy

Adult colour preference has been summarized quantitatively in terms of weights on the two fundamental neural processes that underlie early colour encoding: the S−(L+M) (‘blue–yellow’) and L−M (‘red–green’) cone‐opponent contrast channels ( Ling, Hurlbert & Robinson, 2006 ; Hurlbert & Ling, 2007 ). Here, we investigate whether colour preference in 4–5‐month‐olds may be analysed in the same way. We recorded infants’ eye‐movements in response to pairwise presentations of eight colour stimuli varying only in hue. Infants looked longest at reddish and shortest at greenish hues. Analyses revealed that the L−M and S−(L+M) contrast between stimulus colour and background explained around half of the variation in infant preference across the hue spectrum. Unlike adult colour preference patterns, there was no evidence for sex differences in the weights on either of the cone‐opponent contrast components. The findings provide a quantitative model of infant colour preference that summarizes variation in infant preference across hues.     

Edges,colour and awareness in blindsight

It remains unclear what is being processed in blindsight in response to faces, colours, shapes, and patterns. This was investigated in two hemianopes with chromatic and achromatic stimuli with sharp or shallow luminance or chromatic contrast boundaries or temporal onsets. Performance was excellent only when stimuli had sharp spatial boundaries. When discrimination between isoluminant coloured Gaussians was good it declined to chance levels if stimulus onset was slow. The ability to discriminate between instantaneously presented colours in the hemianopic field depended on their luminance, indicating that wavelength discrimination totally independent of other stimulus qualities is absent. When presented with narrow-band colours the hemianopes detected a stimulus maximally effective for S-cones but invisible to M- and L-cones, indicating that blindsight is mediated not just by the mid-brain, which receives no S-cone input, or that the rods contribute to blindsight. The results show that only simple stimulus features are processed in blindsight.     

Binocular summation of chromatic changes as measured by visual reaction time

We determined visual reaction times to monocular and binocular changes in the luminance of isochromatic stimuli and to monocular and binocular changes in the color of isoluminant stimuli. Two isoluminant color changes were tested: chromatic variations along the red-green axis of Boynton's (1986) two-stage color vision model and chromatic variations along the yellow-blue axis of the same model. The results indicate a greater degree of binocular summation for luminance change than for color change. This result was largely independent of the motor component of reaction time.     

An oblique effect of chromatic gratings measured by color-mixture thresholds

Contrast sensitivity is lower for obliquely oriented achromatic gratings than for vertical or horizontal gratings at high spatial and low temporal frequencies. Although this response is suggestive of mediation by P-pathway cortical correlates, no clear sensory (i.e. class 1) oblique effect has been demonstrated with isoluminant chromatic stimuli. In the present experiment, a two-alternative forced-choice detection task was used to measure observers' sensitivity to spatiotemporal sinusoids varying in orientation and color contrast. A maximum-likelihood method fit ellipses to the thresholds, with the length of each ellipse taken as a measure of chromatic contrast sensitivity at isoluminance, and the width as luminance contrast threshold. A chromatic oblique effect was observed at about 3 cycles deg-1 suggesting an orientation bias within the cortical stream conveying P-cell activity.     

Color adaptation in a yellow-blue system: proof of a simple numerical relationship]

Three axes in color space are specified: a (unique) yellow - (unique) blue axis, a (unique) green - magenta axis and brightness. Based on the two chromatic axes two linear opponent colors systems are defined: a red/green-system and a yellowish/bluish-system. A numerical relation is presented to describe color adaptation for the yellowish/bluish-system under adaptation to (unique) yellow and (unique) blue: two pairs of color stimuli are equivalent with regard to the yellowish/bluish-system - consisting of a test stimulus and an adaptation stimulus, respectively - if the ratios from the yellowish/bluish-coordinates of test stimulus and adaptation stimulus are identical. A control of brightness and the red/green-system is presupposed. For several (unique) yellow and (unique) blue adaptation stimuli it is examined how a test stimulus that appears neither yellowish nor bluish changes its location on the (unique) yellow - (unique) blue axis within color space. Three observers take part in the experiment. For each observer a plane of constant brightness and the opponent colors axes are estimated experimentally. The data show that the ratios from the yellowish/bluish coordinates of test stimulus and adaptation stimulus are essentially constant. The results are compared with analogous data for the red/green-system. The findings provide evidence for the specification of the three phenomenal axes. The specification is discussed with regard to Hering's opponent colors theory and Krauskopf's three "cardinal" axes [1982, Vision Research, 22, 1123-1131].     

Movement-induced modulation of orientation-specific color aftereffects

The intensity of the McCollough effect is modified when, following exposure to the inducing chromatic stimuli, the achromatic test gratings are seen oscillating orthogonally to their orientations. Green aftereffect seen on stationary test gratings is enhanced by oscillations, while pink aftereffect present on the stationary gratings fades upon oscillation of the test stimulus. These opponent changes are tentatively accounted for in terms of an interaction between Fechner-Benham type induced color and processes that mediate the orientation-specific chromatic aftereffects.     

The L/M-opponent channel provides a distinct and time-dependent contribution towards visual recognition

The visual pathway has been successfully modelled as containing separate channels consisting of one achromatically opponent mechanism and two chromatically opponent mechanisms. However, little is known about how time affects the processing of chromatic information. Here, parametrically defined objects were generated. Reduced-colour objects were interleaved with full-colour objects and measures of recognition performance (d') were compared by the continuous serial recognition paradigm. Measures were taken at multiple delay intervals (1, 4, 7, and 10 s). When chromatic variations were removed, recognition performance was impaired, but at the 1 s and 10 s intervals only. When luminance variations were removed, no impairment resulted. When only L/M-opponent modulations were removed, a deficit in performance was produced only at the 1 s and 10 s intervals, similar to the removal of chromatic variation. When only S-opponent modulations were removed, no impairment was observed. The results suggest that the L/M-opponent pathway provides a specialised contribution to visual recognition, but that its effect is modulated by time. A three-stage process model is proposed to explain the data.     

An analogy between colour and spatial coding

The early stages of colour coding are well established in that the trichromatic receptor stage is followed by a set of opponent colour channels. One interpretation of the sequence is that opponent channels carry unrelated aspects of the colour stimulus, unlike the cone channels. The overlap of the cone channels can be removed by decorrelating their spectral-sensitivity functions, and this procedure has been found to give opponent colour channels which match those found psychophysically. Since the known spatial-frequency channels also show considerable overlap, the question arises which aspects of the spatial stimulus are captured by decorrelating the spatial-frequency channels. The results of decorrelating the spatial-frequency channels are that the first decorrelated spatial filter acts as a broad bandpass filter which has a peak sensitivity at 7.9 cycles deg-1, and that the second decorrelated spatial filter acts as an opponent spatial-frequency channel, with a minimum output at a low (4.1 cycles deg-1) spatial frequency and a maximum output at a high (15.1 cycles deg-1) spatial frequency. The characteristics of the first decorrelated filter closely resemble the properties of the foveal perceptive field which have been used to explain the Hermann grid illusion. Thus, the decorrelation analysis produces a model for the functional organisation of the channel implementation at the neural and psychophysical levels, but which directly relates to the subjective appearance of the visual stimuli.     

Constructing isoluminant stimuli for word recognition research: A precautionary study

Isoluminant stimuli are used increasingly often to investigate processes underlying visual word recognition. However, construction of isoluminant stimuli is not straightforward, and inappropriate construction may have the result of misinforming theories that relate word recognition to neurological function. To inform the use of isoluminant stimuli in studies of word recognition, the present article details two experiments in which isoluminant stimuli were constructed using physical onscreen luminance matching and heterochromatic flicker photometry (HFP) with four different stimulus types: disks, squares, rectangles, and letter strings. The findings reveal (1) substantial differences between isoluminance determined by physical onscreen luminance matching and HFP, (2) substantial differences in HFP isoluminance across stimulus types, and (3) substantial differences in HFP isoluminance across participants. These findings indicate that, in contrast to common practice in word recognition research, HFP provides a better indication of isoluminance than physical onscreen matching; but HFP stimuli should match those used in the experiment proper and should be used to assess isoluminance individually for each participant.     

Colour matching of isoluminant samples and backgrounds: a dimming effect

Sequential asymmetrical colour matching of forty Munsell samples simulated under illuminant C and one of eight test illuminants was carried out. The subjects matched the appearance of each sample under illuminant C with its appearance under the test illuminant. Samples and background (N7) were presented for 1 s under the test illuminant and were isoluminant with each other. Subjects adjusted hue, chroma, and value under illuminant C. The experiments distinguished two groups of subjects; some observers needed to reduce the luminance of the sample to make a match while others did not. This 'dimming' occurred when the matches were close to cardinal axes, especially the tritanopic confusion line. A model of luminance and cone-opponent mechanisms contributing to brightness can account for the dimming effect. Details of analysis in cone-opponent space (L - M, L + M - S, L + M) are presented in the companion paper (Stanikunas et al, 2005 Perception 34 this issue).     

An n-dimensional Weber Law and the Corresponding Fechner Law

Weber's law of 1834, DeltaS/S=c for the just noticeable difference (jnd), can be written as S+DeltaS=kS, k=1+c. It follows that the stimulus decrement required to elicit one jnd of sensation is S-DeltaS*=k(-1)S. If generalized for two stimulus dimensions and two corresponding response dimensions, Weber's law would have to state such equations for all directions of change in the plane. A two-dimensional Weber law with exactly these properties is realized by [S(x)+DeltaS(x)(straight theta), S(y)+DeltaS(y)(straight theta)]=[k(sin(straight theta))S(x), k(cos(straight theta))S(y)] which determines the stimulus coordinates for all stimuli just noticeably different from the stimulus (S(x), S(y)) in all directions 0R(2)) is [x, y]mapsto[log(k)(x), log(k)(y)]. The solution is generalized to arbitrarily many dimensions by substituting the sin and cos in the generalized Weber law by the standard coordinates of a unit vector. Copyright 2000 Academic Press.     

The binocular combination of chromatic contrast

How is chromatic contrast combined binocularly? One index of binocularity is the binocular contrast summation ratio (BCSR), which is the improvement in contrast sensitivity with binocular rather than monocular presentation. Simmons and Kingdom (1998, Vision Research 38 1063-1071) noted that BCSRs with some red-green isoluminant stimuli were suggestive of full linear summation. This suggestion was investigated further in four subjects by measuring binocular and monocular contrast thresholds for the detection of 0.5 cycle deg(1) isoluminant (red-green) and isochromatic (yellow-black) Gabor patches. These Gabor patches had either vertically or horizontally oriented carrier gratings and were either dichoptically in phase (same coloured bars in binocular correspondence) or in dichoptic anti-phase (opposite coloured bars in binocular correspondence). Full linear summation would be indicated by BCSRs of 2 for the in-phase and close to 0 for the anti-phase conditions. Mean BCSRs at isoluminance were 1.93 and 0.90, respectively, for the in-phase and anti-phase stimuli with horizontal carriers, the former being consistent with full linear summation, but the latter not. Despite these results, BCSRs obtained with isoluminant and isochromatic stimuli under similar conditions were not statistically distinguishable from each other, although there was a tendency for summation at isoluminance with in-phase stimuli to be higher and anti-phase stimuli to be lower. These data fall short of demonstrating full linear summation of chromatic contrast between the eyes under all presentation conditions, but they do indicate that there are strong binocular interactions at red-green isoluminance, which are similar to, and possibly even stronger than, those obtained with luminance stimuli.     

Directional sensitivity to a tactile point stimulus moving across the fingerpad

The ability of subjects to discriminate between directions of a point contact moving across the fingerpad was examined. Subjects were required to report, using an adaptive two-interval, two-alternative forced-choice procedure, whether in two sequential stimuli the direction of motion changed in a clockwise or counterclockwise direction. The overall mean orientation-change threshold across eight stimulus orientations was approximately 14°, with the lowest threshold for the point motion toward the wrist. This observed lower threshold in the distal-to-proximal direction is thought to be due to stretching of the skin at the tip of the fingernail, to which one may be particularly sensitive. For all orientations, thresholds were generally more uniform and higher than those reported on vibrotactile linear contactor arrays for horizontal and vertical orientations.     

Sources of stimulus control during multiple discriminations

Pigeons learned to respond to the middle-sized member of six or seven sets of three stimuli differing in size. The sets were used successively, each serving as the discrimination problem from 10 to 16 times. After attaining criterion with one set, the birds received the others as probes. The number of responses in probes was related to the similarity of the probes to the prevailing discrimination problem. The birds responded either to the probe stimulus to which responding had most recently been reinforced, or to the probe stimulus closest in size to the positive member (S+) of the prevailing discrimination problem. Responses to a middle-sized probe-set stimulus occurred when it was the probe-set member most recently correlated with reinforcement, when it was one of two stimuli closest in size to S+, and when the stimulus closest in size to S+ was a negative member of the discrimination problem. All of the behavior could be explained in terms of control by the absolute sizes of the various stimuli.     

Emergent simple discrimination in children: Preference for non-preferred stimuli

Previous research (Smeets, 1991) suggested that when given a new discrimination, children respond on the basis of physical similarity with previously discriminated stimuli. They respond to a stimulus similar to another preferred stimulus (S+ transfer) and respond away from a stimulus similar to another nonpreferred stimulus (S- transfer). When both types of transfer apply to the same stimulus, S+ transfer prevails, S+ Priority Transfer (S+PT). The present study demonstrated that S+PT also occurs when the criterion task consists of two nonpreferred stimuli. When given a choice between two previously nonpreferred stimuli, one similar and one dissimilar to other preferred stimuli, children select the first one. They do not so, however, when a nonpreferred stimulus resembling another preferred stimulus is presented with a new nonpreferred stimulus. These findings suggest that the children's preferences were not based on the physical resemblance with other (non)preferred stimuli but on the functions (S+, S-, S0) of individual stimulus components. A theoretical model is presented that accounts for all experimental data reported in the previous and present study. The model implies that discriminative responding not only results from but also determines the functional properties of individual stimulus elements.     

The opponent process theory and affective reactions

Solomon (1980) proposed an opponent process theory to account for motivational and affective dynamics. This theory asserts that the brain avoids extremes of emotional experience by countering the stimulation it receives with an opposite or opponent reaction. Opponent processes are thought to be responsible for the characteristic changes in affective experience that occur over time, and to account for the dynamics of affective response to such stimuli as skydiving and sauna bathing, which have heretofore been difficult to explain. However, the relevance of this theory for affective experiences in general (beyond physical stimuli and addictions) has yet to be demonstrated. The present paper examines opponent process theory predictions in two settings, involving affective responses to situation-scenarios and emotion-provoking slides. In each study, significant habituation to both positive and negative affective stimuli was found, as the opponent process theory would predict. Subjects also showed a reversal of affect when the stimuli were reversed from positive to negative or vice versa. However, contrary to opponent process theory predictions, there was no evidence that withdrawal responses were greater after habituation to the affective stimulus. The only instance of a significant difference in withdrawal responses was actually in a direction opposite to that which the opponent process theory predicts. All other predicted differences were not significant. The opponent process theory, therefore, was not supported in these data and appears to need revision or qualification as to its domains of application.     

Effects of relative reinforcer frequency on complex color detection

Pigeons were trained under a discrete-trials detection procedure in which one of a set of color stimuli was presented on the center key and a single response turned off the stimulus and illuminated two side keys. Single responses to one or the other side key produced occasional reinforcers depending on the value of the color stimulus. In Experiment 1, one color-stimulus set comprised 559, 564, 569, and 574 nm, and right-key pecks were occasionally reinforced following presentations of members of this set. The other stimulus set comprised 579, 584, 589, and 594 nm, and left-key pecks were occasionally reinforced following presentations of members of this set. Across seven experimental conditions, the left/(left + right) relative reinforcer frequency was varied from .1 to .9. In Experiment 2, one stimulus set contained only one member, 574 nm, and right-key responses were occasionally reinforced following its presentation. Over 12 experimental conditions, two manipulations were carried out. First, the number of stimuli comprising the other stimulus set was increased from one (579 nm) to two (579 and 584 nm) to three (579, 584, and 589 nm) and to four (579, 584, 589, and 594 nm), and left-key responses were reinforced occasionally following center-key presentations of members of this set. Second, for each stimulus combination, the left/(left + right) relative reinforcer frequency was varied from .1 to .5 to .9 across three experimental conditions. The principal finding of Experiments 1 and 2 was that reinforcers and stimuli interacted in their effects on behavior. In Experiment 3, pairs of adjacent stimuli (5 nm apart) in the range 559 to 594 nm were presented in each experimental condition, and the left/(left + right) relative reinforcer frequency was held constant at .5. The data from all three experiments were analyzed according to a detection model describing performance in multiple-stimulus two-response procedures. This model provided independent measures of stimulus discriminability, contingency discriminability, and bias. The analysis showed that (a) consistent with the color-naming function, pigeons were better able to discriminate between higher nanometer values than lower nanometer values; (b) their ability to discriminate between the stimuli was independent of the number of wavelengths comprising each stimulus set; (c) they allocated delivered reinforcers very accurately to the previously emitted response; and (d) no consistent biases emerged.     

Peak shift revisited: a test of alternative interpretations

In Experiment 1, 2 groups of human subjects were trained to respond to 1 of 2 light intensity stimuli, S2 or S4, and then were tested for generalization with a randomized series of increasing values from S1 to S11. Both groups, including the group trained to respond to dimmer value, showed peak shifts to a brighter more centrally located test stimulus. In Experiment 2, which used line angle stimuli, both the size of the difference between S+ and S- and the range of test stimuli that extended beyond S+ were varied. The larger the S(+)-S- separation and the larger the range, the greater was the peak shift obtained. In Experiment 3, training involved an S- (line angle) surrounded by 2 S+ values with testing symmetrical about the training values and covering either a narrow or a wide range. The wide range produced greater peak shifts in both directions from S-. All 3 experiments support an adaptation-level interpretation of intradimensional discrimination learning and generalization test performance in human subjects. Related work with animals suggests the presence of similar processes.