The contribution of the right supra-marginal gyrus to sequence learning in eye movements.

We investigated the role of the human right Supra-Marginal Gyrus (SMG) in the generation of learned eye movement sequences. Using MRI-guided transcranial magnetic stimulation (TMS) we disrupted neural activity in the SMG whilst human observers performed saccadic eye movements to multiple presentations of either predictable or random target sequences. For the predictable sequences we observed shorter saccadic latencies from the second presentation of the sequence. However, these anticipatory improvements in performance were significantly reduced when TMS was delivered to the right SMG during the inter-trial retention periods. No deficits were induced when TMS was delivered concurrently with the onset of the target visual stimuli. For the random version of the task, neither delivery of TMS to the SMG during the inter-trial period nor during the presentation of the target visual stimuli produced any deficit in performance that was significantly different from the no-TMS or control conditions. These findings demonstrate that neural activity within the right SMG is causally linked to the ability to perform short latency predictive saccades resulting from sequence learning. We conclude that neural activity in rSMG constitutes an instruction set with spatial and temporal directives that are retained and subsequently released for predictive motor planning and responses.

Multiple spatial frequency channels in human visual perceptual memory.

Current models of short-term visual perceptual memory invoke mechanisms that are closely allied to low-level perceptual discrimination mechanisms. The purpose of this study was to investigate the extent to which human visual perceptual memory for spatial frequency is based upon multiple, spatially tuned channels similar to those found in the earliest stages of visual processing. To this end we measured how performance on a delayed spatial frequency discrimination paradigm was affected by the introduction of interfering or 'memory masking' stimuli of variable spatial frequency during the delay period. Masking stimuli were shown to induce shifts in the points of subjective equality (PSE) when their spatial frequencies were within a bandwidth of 1.2 octaves of the reference spatial frequency. When mask spatial frequencies differed by more than this value, there was no change in the PSE from baseline levels. This selective pattern of masking was observed for different spatial frequencies and demonstrates the existence of multiple, spatially tuned mechanisms in visual perceptual memory. Memory masking effects were also found to occur for horizontal separations of up to 6 deg between the masking and test stimuli and lacked any orientation selectivity. These findings add further support to the view that low-level sensory processing mechanisms form the basis for the retention of spatial frequency information in perceptual memory. However, the broad range of transfer of memory masking effects across spatial location and other dimensions indicates more long range, long duration interactions between spatial frequency channels that are likely to rely contributions from neural processes located in higher visual areas.

Naming versus matching and the stability of unique hues.

It is known that there is a distortion of hue and saturation in the peripheral visual field. In a previous study, when an asymmetric matching paradigm was used, four hues in the blue, red, yellow and green regions of colour space were unchanged and these were referred to as peripherally invariant (Parry et al., J Opt Soc Am A, 23, 2006, 1586). Three of these invariant hues were similar to unique blue, red and yellow. However, for most observers there was a marked difference between unique and invariant green. To investigate this apparent paradox, we have measured unique hues using a range of eccentricities and colourimetric purities. An asymmetric matching and a 4-AFC paradigm were used to establish peripherally invariant and unique hues, respectively. In the asymmetric matching task the observer matched a peripheral spot with a para-foveal spot, for 24 different hues at 18° eccentricity. In the 4-AFC paradigm, 41 hues were presented 20 times at three purities (0.5, 0.75 and 1.0) and three eccentricities (18°, 10° and 1°). The observer had to name the hues as red, blue, green or yellow. Unique hues were found to be constant with eccentricity and purity. The unique green, established with 4-AFC, was found to differ from the invariant green, determined using the matching task. However, red, blue and yellow invariant hues correspond well with unique hues. The data suggest that different mechanisms mediate the matching of green compared with the identification of unique hues. This is similar to the difference between detection and discrimination of spectral stimuli: the detection process is dominated by the cone opponent mechanisms and is most sensitive, whereas more central processes, serving unique hues, influence discrimination.

A behavioural investigation of human visual short term memory for colour.

We examined visual short term memory (VSTM) for colour using a delayed-match-to-sample paradigm. In these experiments we measured the effects of increasing inter-stimulus interval (ISI), varying between 0 and 10 s, on the ability of five colour normal human observers to make colour matches between a reference and subsequently presented test stimuli. The coloured stimuli used were defined by different chromatic axes on the isoluminant plane of DKL colour space. In preliminary experiments we used a hue scaling procedure to identify a total of 12 colour stimuli which served as reference hues in the colour memory experiments: four stimuli were exemplars of red, green, blue and yellow colour appearance categories, four were located between these categories and a further four were located on the cardinal axes that isolated the activity of the cone-opponent mechanisms. Our results demonstrate that there is a reduction in the ability of observers to make accurate colour matches with increasing ISIs and that this reduced performance was similar for all colour stimuli. However, the shifts in hue that were measured between the reference and matched test stimuli were significantly greater for the cardinal stimuli compared to those measured for the stimuli defined by the hue scaling procedure. This deterioration in the retention of hue in VSTM for stimuli that isolate cone-opponent mechanisms may be a reflection of the reorganisation of colour processing that occurs in the cortex where colour appearance mechanisms become more prominent.

The contribution of human cortical area V3A to the perception of chromatic motion: a transcranial magnetic stimulation study.

Area V3A was identified in five human subjects on both a functional and retinotopic basis using functional magnetic resonance imaging techniques. V3A, along with other visual areas responsive to motion, was then targeted for disruption by repetitive transcranial magnetic stimulation (rTMS) whilst the participants performed a delayed speed matching task. The stimuli used for this task included chromatic, isoluminant motion stimuli that activated either the L-M or S-(L+M) cone-opponent mechanisms, in addition to moving stimuli that contained only luminance contrast (L+M). The speed matching task was performed for chromatic and luminance stimuli that moved at slow (2 degrees/s) or faster (8 degrees/s) speeds. The application of rTMS to area V3A produced a perceived slowing of all chromatic and luminance stimuli at both slow and fast speeds. Similar deficits were found when rTMS was applied to V5/MT+. No deficits in performance were found when areas V3B and V3d were targeted by rTMS. These results provide evidence of a causal link between neural activity in human area V3A and the perception of chromatic isoluminant motion. They establish area V3A, alongside V5/MT+, as a key area in a cortical network that underpins the analysis of not only luminance but also chromatically-defined motion.

Disruptions to human speed perception induced by motion adaptation and transcranial magnetic stimulation.

To investigate the underlying nature of the effects of transcranial magnetic stimulation (TMS) on speed perception, we applied repetitive TMS (rTMS) to human V5/MT+ following adaptation to either fast- (20 deg/s) or slow (4 deg/s)-moving grating stimuli. The adapting stimuli induced changes in the perceived speed of a standard reference stimulus moving at 10 deg/s. In the absence of rTMS, adaptation to the slower stimulus led to an increase in perceived speed of the reference, whilst adaptation to the faster stimulus produced a reduction in perceived speed. These induced changes in speed perception can be modelled by a ratio-taking operation of the outputs of two temporally tuned mechanisms that decay exponentially over time. When rTMS was applied to V5/MT+ following adaptation, the perceived speed of the reference stimulus was reduced, irrespective of whether adaptation had been to the faster- or slower-moving stimulus. The fact that rTMS after adaptation always reduces perceived speed, independent of which temporal mechanism has undergone adaptation, suggests that rTMS does not selectively facilitate activity of adapted neurons but instead leads to suppression of neural function. The results highlight the fact that potentially different effects are generated by TMS on adapted neuronal populations depending upon whether or not they are responding to visual stimuli.

Nasal-temporal differences in cone-opponency in the near peripheral retina.

The purpose of this study is to establish whether nasal-temporal differences in cone photoreceptor distributions are linked to differences in colour matching performance in the two hemi-fields. Perceived shifts in chromaticity were measured using an asymmetric matching paradigm. They were expressed in terms of hue rotations and relative saturation changes and also in terms of activation levels of L-M or S-(L+M) cone-opponent channels. Up to 19 degrees eccentricity there was little difference in chromaticity shifts between nasal and temporal retina for either channel. For matches beyond 19 degrees L-M activation is significantly lower in the nasal field and the S-(L+M) channel was equally activated in both fields. The data are consistent with the asymmetric distribution of L- and M-cones in the nasal and temporal retinae.

The perception of speed based on L-M and S-(L+M) cone opponent processing.

We have measured perceived speed and speed discrimination thresholds for stimuli that selectively activate the L-M, S-(L+M) cone opponent and L+M (luminance) post-receptoral pathways. For low speeds and low contrasts speed discrimination thresholds for L-M and S-(L+M) are similar but are higher and have a greater dependency upon contrast than those for luminance motion. These differences between chromatic and luminance speed perception can be eliminated when stimuli are equated with respect to their individual motion detection thresholds (MDTs). For fast moving gratings speed perception based upon L-M, S-(L+M) and L+M signals is similar in terms of threshold performance and contrast dependency. These results are consistent with the view that there are separate mechanisms for the analysis of chromatic and luminance motion, the relative contributions of which may change as a function of stimulus contrast and speed. The similarity in performance for S-(L+M) and L+M chromatic stimuli across a range of stimulus parameters suggests that signals derived from the two cone opponent pathways can be used equally well. Our results argue against the idea that speed perception is compromised when it is based upon information derived from the S-(L+M) cone opponent pathway.

Reaction time measures of adaptation to chromatic contrast.

Simple reaction times (RTs) were measured to brief temporally blurred (total onset 570 ms) Gaussian isoluminant chromatic patches (s.d. 0.5 degrees) whose chromaticities lay along the cardinal chromatic axes (0 degrees, 90 degrees, 180 degrees, and 270 degrees in MBDKL color space). Bipolar adapting stimuli were employed (0 degrees versus 180 degrees or 90 degrees versus 270 degrees). These were larger Gaussian blobs (s.d. 1 degree), modulating sinusoidally between the two hues at 1 Hz. Throughout, the background was illuminant "C" (x = 0.31, y = 0.316, L = 12.5). In a single run, a series of 64 or 32 stimuli were presented without adaptation, followed by 64 or 32 stimuli each of which was preceded by 3 s of adaptation, either along the same or the orthogonal chromatic axis. Finally, 192 or 128 RTs were recorded to measure the time course of recovery from adaptation. Both adapting and test stimuli were presented at fixed supra-threshold contrasts. The effect of adaptation was seen as a lengthening of the RT, which occurred in the first few seconds of the adaptation period. After cessation of adaptation, there was a similarly rapid shortening of RT, although full recovery took 60-90 s. Adaptation gain functions suggested that the S-(L + M) system was less prone to adaptation than L-M.

Speed selectivity in visual short term memory for motion.

In this study we employed a 'memory masking' paradigm to determine which stimulus attributes are important in the storage of information about the speed of moving grating stimuli in visual short term memory (VSTM). Delayed speed discrimination thresholds were measured in the presence of masking stimuli which varied in terms of their spatial and temporal frequency content. Memory masking results demonstrate that it is genuinely the speed of the stimulus, as opposed to temporal or spatial frequency content, that is crucial in the retention of information about motion in visual short term memory. The property of speed selectivity exhibited by VSTM mirrors that reported for neurons in area V5/MT, a brain area crucial for the processing of visual motion in primate brain. This link between area V5/MT and VSTM for motion is consistent with current views which suggest that there is a close association between the neural mechanisms involved in the analysis of sensory information and those involved in its retention in short term memory.

Misperceptions of speed for chromatic and luminance grating stimuli.

Errors in the perception of speed of moving visual stimuli can occur when presented stimuli are of unequal contrast and when they appear alongside additional modifier stimuli that move at different speeds. We have examined these misperceptions for chromatic and luminance grating stimuli in order to assess to what extent these different kinds of motion cue might be utilised in the analysis of speed of moving objects. We show that the dependence on contrast of speed matching for luminance and chromatic stimuli is similar over a range of stimulus speeds greater than 4 deg/s. Differences between the contrast dependencies of speed perception for chromatic and luminance stimuli are only evident at slow speeds (< 4 deg/s) and low contrasts. The presence of modifier stimuli can directly influence the perceived speed at both high and low velocities and contrasts. This influence was found to be independent of the modifiers' chromaticity and was greatest when the modifiers were adjacent to and presented simultaneously with the test and reference stimuli. However, the modifiers were still able to induce measurable changes in perceived speed for increased separations over space and time. Taken together these results indicate that whilst differences do exist in the contrast dependencies of speed perception for chromatic and luminance stimuli, they are evident only for a narrow range of stimulus parameters (i.e. low speed and low contrast). There appears to be ample scope for interactions between chromatic and luminance contrast in speed perception where there is the capacity to pool this information over a relatively broad spatio-temporal extent.

The cortical topography of visual evoked potentials elicited by chromatic and luminance motion.

When motion-onset VEPs are elicited by moving luminance patterns, the motion specific component of the response, N2, is more prominent at electrode sites that overlay the lateral occipito-parietal cortex close to area V5/MT, than over the primary visual cortex. Functional segregation suggests that colour and motion processing should take place along different ventral occipito-temporal and lateral occipito-parietal pathways, respectively. Hence, a different topographical distribution might be expected for the motion-onset VEPs elicited by chromatic and luminance motion stimuli. We recorded motion-onset VEPs to moving luminance or isoluminant chromatic sinusoidal grating stimuli from five electrodes sites located at Oz, and at four locations (T1-T4) lateral to Oz, at intervals of 5% of the head circumference. Responses were recorded from 6 subjects over a range of speeds and contrasts. The results showed that the N2 component was maximal at similar lateral electrode locations (T2) for both luminance-defined and chromatically-defined motion. The earlier P1 component was of greatest magnitude at the occipital pole (Oz) and decreased with more lateral electrode placement and again this was the same for colour and luminance responses. These similarities suggest a common origin for VEPs elicited by colour and luminance defined motion.

Cone opponency in the near peripheral retina.

Changes of color perception in the peripheral field are measured using an asymmetric simultaneous matching paradigm. The data confirm previous observations in that saturation changes can be neutralized if the test target is increased in size. However, this compensation does not apply to hue shifts. We show that some hues remain unchanged with eccentricity whereas others exhibit substantial changes. Here the color shifts are plotted in terms of a second-stage cone opponent model. The data suggest that the S-L+M channel is more robust to increasing eccentricity than the L-M channel. Observations are interpreted in terms of the known underlying morphological and physiological differences in these channels.

The segregation and integration of colour in motion processing revealed by motion after-effects.

Analysis of the colour and motion of objects is widely believed to take place within segregated processing pathways in the primate visual system. However, it is apparent that this segregation cannot remain absolute and that there must be some capacity for integration across these sub-modalities. In this study, we have assessed the extent to which colour constitutes a separable entity in human motion processing by measuring the chromatic selectivity of two kinds of after-effect resulting from motion adaptation. First, the traditional motion after-effect, where prolonged inspection of a unidirectional moving stimulus results in illusory motion in the opposite direction, was found to exhibit a high degree of chromatic selectivity. The second type of after-effect, in which motion adaptation induces misperceptions in the spatial position of stationary objects, was completely insensitive to chromatic composition. This dissociation between the chromatic selectivities of these after-effects shows that chromatic inputs remain segregated at early stages of motion analysis, while at higher levels of cortical processing there is integration across chromatic, as well as achromatic inputs, to produce a unified perceptual output.

Effect of foveal tritanopia on reaction times to chromatic stimuli.

To investigate the effect of foveal inhomogeneities on sensitivity to chromatic stimuli, we measured simple reaction times (RTs) and detection thresholds to temporally and spatially blurred isoluminant stimuli at retinal eccentricities from 0 deg to 8 deg. Three color-normal subjects participated. Contrast gain was derived from the slope of the RT versus contrast function. With a Gaussian spatial distribution (S.D. = 0.5 deg) and modulation between white (CIE x, y, L = 0.31, 0.316, 12.5 cd x m(-2)) and blue (MBDKL 90 deg), gain was maximal at about 2-deg eccentricity and declined by approximately 1 log unit towards the center and the periphery. The red (0 deg) and green (180 deg) cardinal axes showed maximum gain in the center, whilst the yellow (270 deg) data were intermediate. Although the spatial extent of the Gaussian spot was much larger than the S-cone free zone, we wished to determine whether foveal tritanopia was responsible for the marked drop in sensitivity to the 90-deg stimulus. To align the color vector along a tritan line, we used a smaller disk (0.3 deg) with a blurred edge and measured detection threshold, rotating the vector until minimum central sensitivity was obtained. Other workers have used transient tritanopia or minimally distinct border to similar effect. By repeating this at different locations in color space, a group of vectors were obtained. These converged near to the S-cone co-punctal point, evidence that they lay along tritan confusion lines. These threshold findings were then confirmed using the RT-derived contrast gain function. The tritan vectors were less pronounced as stimulus size increased. With the vector optimized to produce foveal tritanopia, the RT gain versus eccentricity functions for the 90-deg and 270-deg stimuli both fell markedly in the center and periphery, and sensitivity peaked at about 3-deg eccentricity. There are some similarities between these findings and the underlying photoreceptor distributions. As a result, there is a greater difference in gain between red-green and blue-yellow systems in the center than in the near periphery. We conclude that the RT versus contrast function is a sensitive index of foveal opponency.

Chromatic adaptation, perceived location, and color tuning properties.

We have studied the influence of chromatic adaptation upon the perceived visual position of a test stimulus using a Vernier alignment task. Maximum and minimum offsets in spatial position are generated when the adapting and test stimuli lie on the same and orthogonal axes in MBDKL color space, respectively. When the test stimuli lie on intermediate color axes, the measured positional shifts decrease as a function of the angular separation in color space (phi) from the adapting stimulus. At low stimulus contrasts, these shifts follow a sinusoidal function of phi and exhibit broad chromatic tuning and can be accounted for by a model in which the centroid is extracted from the linear combination of after-image, formed by the adapting stimulus, and the test stimulus. Such linear, broadband behavior is consistent with the response properties of chromatic neurons in the precortical visual pathway. At high contrast, and when adaptation gets closer to the S/(L+M) axis, the tuning functions become narrower and require sinusoids raised to increasingly higher exponents in order to describe the data. This narrowing of chromatic tuning is consistent with the tuning properties of chromatic neurons in the striate cortex, and implies the operation of a nonlinear mechanism in the combination of cone outputs.

The influence of stimulus chromaticity on the isoluminant motion-onset VEP.

Motion-onset visual evoked potentials (VEPs) were elicited by low spatial frequency chromatic isoluminant gratings presented in a central 7 degrees circular field. The chromatic composition of the stimuli was varied so as to modulate along different axes in colour space. For slow speeds (<5 degrees/s) changing the chromatic axis induced large response differences between the S- and L/M-cone VEPs. At faster speeds (5-12 degrees/s) the effects were not as marked. A dichotomy between the slow and fast responses was also shown to exist in terms of their contrast dependencies, the former exhibiting a stronger dependency on contrast than the latter. These findings suggest that neural substrates with chromatic sensitivity are involved in the generation of S- and L/M-cone mediated motion-onset VEPs at low velocities. At higher velocities, responses are generated by different mechanisms that possess little or no chromatic sensitivity.

Visual evoked potentials elicited by chromatic motion onset.

Visually Evoked Potentials (VEPs) were recorded in response to the onset of chromatic and luminance motion gratings of 1 cpd and luminance 40 cd m(-2) subtending a 7 degrees field. At slow speeds (< or =2 cycles s(-1)) the motion onset response exhibits a clear amplitude minimum at isoluminance. Over the Michelson contrast range tested (0.05-0.75) the chromatic response at 2 cycles s(-1) possesses a linear response function compared to the saturating function of the luminance response and the contrast dependency of the former is a factor of 5-6 times greater than for the latter. These differences are suggestive of different neural substrates for the chromatic and luminance motion VEPs at slow speeds. At 10 cycles s(-1) the chromatic motion onset VEP exhibits no amplitude minimum at isoluminance and becomes more like its luminance counterpart in terms of its saturating contrast response function. Furthermore, the contrast dependency of the chromatic and luminance responses differs by only a factor of 1.6 at this faster rate. These findings are consistent with the idea of separate motion mechanisms that operate at fast and slow speeds, the latter having separate channels for colour and luminance motion.

Red--green and blue--yellow mechanisms are matched in sensitivity for temporal and spatial modulation.

The spatial and temporal properties of human colour vision are examined using isoluminant, red--green and blue--yellow tritanopic gratings. Chromatic sensitivity is found to be low-pass as a function of both spatial and temporal frequency along all the chromatic axes investigated, including the tritanopic confusion lines employed to examine the properties of the S-cone driven mechanism. Comparison of sensitivity to on-off and contrast reversing stimuli indicates that transient mechanisms contribute to the detection of red--green patterns but that the detection of S-cone specific patterns is governed by sustained mechanisms. By compensating for transient contributions to red--green sensitivity, it is shown that sensitivity of chromatic mechanisms dominated by L- and M-cone input are closely matched to those with S-cone input.

Motion adaptation in chromatic motion-onset visual evoked potentials.

The aim of this study was to investigate the influence of motion adaptation on visual evoked potentials (VEPs) elicited by the onset of isoluminant chromatic motion. VEPs were recorded from the occipital cortex of human subjects using a sinusoidal grating stimulus of one cycle per degree which moved at either a velocity of 2 or 10 degrees/s and subtended a field of 7 degrees with a mean luminance of 30 cdm(-2). In the first experiment the effects of adaptation were investigated via the manipulation of the stimulus duty cycle which was varied between 11-90%. The results showed a significant (p < 0.001) reduction in the N2-P2 amplitude of the chromatic response. In contrast, P1-N2 amplitude was not significantly affected by motion adaptation. Subsequent experiments demonstrated that the chromatic motion onset VEP was attenuated not only following adaptation to isoluminant chromatic motion, but also to luminance motion as well. These results indicate that the chromatic motion onset VEP is just as susceptible as its luminance counterpart to motion after effects (MAEs) and as a result it is highly likely that it is a motion specific response. Furthermore, the fact that the VEP shows that there are cross-adaptation effects between motion defined by change in colour and by change in luminance, suggests that the two types of motion stimuli have inputs into a common motion mechanism.