The invariance of directional tuning with contrast and coherence.

The responses of motion mechanisms depend not only on the direction of a stimulus, but also on its contrast, coherence and speed. We examined how contrast, coherence and directional selectivity interact by measuring directional tuning psychophysically across a wide range of coherence and contrast levels. We fit data with a simple model that estimated directional tuning bandwidth using contrast and coherence gain parameters that were based on neurophysiological estimates. This model estimated a bandwidth of approximately 90 degrees for directionally selective mechanisms. Bandwidth was invariant across a wide range of contrasts and coherences, as predicted by models of contrast normalization.

Motion opponency in visual cortex.

Perceptual studies suggest that visual motion perception is mediated by opponent mechanisms that correspond to mutually suppressive populations of neurons sensitive to motions in opposite directions. We tested for a neuronal correlate of motion opponency using functional magnetic resonance imaging (fMRI) to measure brain activity in human visual cortex. There was strong motion opponency in a secondary visual cortical area known as the human MT complex (MT+), but there was little evidence of motion opponency in primary visual cortex. To determine whether the level of opponency in human and monkey are comparable, a variant of these experiments was performed using multiunit electrophysiological recording in areas MT and MST of the macaque monkey brain. Although there was substantial variability in the degree of opponency between recording sites, the monkey and human data were qualitatively similar on average. These results provide further evidence that: (1) direction-selective signals underly human MT+ responses, (2) neuronal signals in human MT+ support visual motion perception, (3) human MT+ is homologous to macaque monkey MT and adjacent motion sensitive brain areas, and (4) that fMRI measurements are correlated with average spiking activity.

Temporal sensitivity of human luminance pattern mechanisms determined by masking with temporally modulated stimuli.

Target contrast thresholds were measured using vertical spatial Gabor targets in the presence of full field maskers of the same spatial frequency and orientation. In the first experiment both target and masker were 2 cpd. The target was modulated at a frequency of 1 or 10 Hz and the maskers varied in temporal frequency from 1 to 30 Hz and in contrast from 0.03 to 0.50. In the second experiment both target and masker had a spatial frequency of 1, 5 or 8 cpd. The target was modulated at 7.5 Hz and the same set of maskers was used as in the first experiment. The results are not consistent with a widely used model that is based on mechanisms in which excitation is summed linearly and the sum is transformed by an S-shaped nonlinear excitation-response function. A new model of human pattern vision mechanisms, which has excitatory and divisive inhibitory inputs, describes the results well. Parameters from the best fit of the new model to the results of the first experiment show that the 1 Hz and 10 Hz targets were detected by mechanisms with temporal low-pass and band-pass excitatory sensitivity, respectively. Fits to the second experiment suggest that at 1 cpd, the excitatory tuning of the detecting mechanism is band-pass. At 5 and 8 cpd, the mechanisms are excited by a broad range of temporal frequencies. Mechanism sensitivity to divisive inhibition depends on temporal frequency in the same general way as sensitivity to excitation. Mechanisms are more broadly tuned to divisive inhibition than to excitation, except when the target temporal frequency is high.

Neuronal basis of contrast discrimination.

Psychophysical contrast increment thresholds were compared with neuronal responses, inferred from functional magnetic resonance imaging (fMRI) to test the hypothesis that contrast discrimination judgements are limited by neuronal signals in early visual cortical areas. FMRI was used to measure human brain activity as a function of stimulus contrast, in each of several identifiable visual cortical areas. Contrast increment thresholds were measured for the same stimuli across a range of baseline contrasts using a temporal 2AFC paradigm. FMRI responses and psychophysical measurements were compared by assuming that: (1) fMRI responses are proportional to local average neuronal activity; (2) subjects choose the stimulus interval that evoked the greater average neuronal activity; and (3) variability in the observer's psychophysical judgements was due to additive (IID) noise. With these assumptions, FMRI responses in visual areas V1, V2d, V3d and V3A were found to be consistent with the psychophysical judgements, i.e. a contrast increment was detected when the fMRI responses in each of these brain areas increased by a criterion amount. Thus, the pooled activity of large numbers of neurons can reasonably well predict behavioral performance. The data also suggest that contrast gain in early visual cortex depends systematically on spatial frequency.

Spatial attention affects brain activity in human primary visual cortex.

Functional MRI was used to test whether instructing subjects to attend to one or another location in a visual scene would affect neural activity in human primary visual cortex. Stimuli were moving gratings restricted to a pair of peripheral, circular apertures, positioned to the right and to the left of a central fixation point. Subjects were trained to perform a motion discrimination task, attending (without moving their eyes) at any moment to one of the two stimulus apertures. Functional MRI responses were recorded while subjects were cued to alternate their attention between the two apertures. Primary visual cortex responses in each hemisphere modulated with the alternation of the cue; responses were greater when the subject attended to the stimuli in the contralateral hemifield. The attentional modulation of the brain activity was about 25% of that evoked by alternating the stimulus with a uniform field.

Color signals in human motion-selective cortex.

The neural basis for the effects of color and contrast on perceived speed was examined using functional magnetic resonance imaging (fMRI). Responses to S cone (blue-yellow) and L + M cone (luminance) patterns were measured in area V1 and in the motion area MT+. The MT+ responses were quantitatively similar to perceptual speed judgments of color patterns but not to color detection measures. We also measured cortical motion responses in individuals lacking L and M cone function (S cone monochromats). The S cone monochromats have clear motion-responsive regions in the conventional MT+ position, and their contrast-response functions there have twice the responsivity of S cone contrast-response functions in normal controls. But, their responsivity is far lower than the normals' responsivity to luminance contrast. Thus, the powerful magnocellular input to MT+ is either weak or silent during photopic vision in S cone monochromats.

Psychophysical evidence for a magnocellular pathway deficit in dyslexia.

The relationship between reading ability and psychophysical performance was examined to test the hypothesis that dyslexia is associated with a deficit in the magnocellular (M) pathway. Speed discrimination thresholds and contrast detection thresholds were measured under conditions (low mean luminance, low spatial frequency, high temporal frequency) for which psychophysical performance presumably depends on M pathway integrity. Dyslexic subjects had higher psychophysical thresholds than controls in both the speed discrimination and contrast detection tasks, but only the differences in speed thresholds were statistically significant. In addition, there was a strong correlation between individual differences in speed thresholds and reading rates. These results support the hypothesis for an M pathway abnormality in dyslexia, and suggest that motion discrimination may be a more sensitive psychophysical predictor of dyslexia than contrast sensitivity.

Functional magnetic resonance imaging of early visual pathways in dyslexia.

We measured brain activity, perceptual thresholds, and reading performance in a group of dyslexic and normal readers to test the hypothesis that dyslexia is associated with an abnormality in the magnocellular (M) pathway of the early visual system. Functional magnetic resonance imaging (fMRI) was used to measure brain activity in conditions designed to preferentially stimulate the M pathway. Speed discrimination thresholds, which measure the minimal increase in stimulus speed that is just noticeable, were acquired in a paradigm modeled after a previous study of M pathway-lesioned monkeys. Dyslexics showed reduced brain activity compared with controls both in primary visual cortex (V1) and in several extrastriate areas, including area MT and adjacent motion-sensitive areas (MT+) that are believed to receive a predominant M pathway input. There was a strong three-way correlation between brain activity, speed discrimination thresholds, and reading speed. Subjects with higher V1 and MT+ responses had lower perceptual thresholds (better performance) and were faster readers. These results support the hypothesis for an M pathway abnormality in dyslexia and imply strong relationships between the integrity of the M pathway, visual motion perception, and reading ability.

Brain activity in visual cortex predicts individual differences in reading performance.

The relationship between brain activity and reading performance was examined to test the hypothesis that dyslexia involves a deficit in a specific visual pathway known as the magnocellular (M) pathway. Functional magnetic resonance imaging was used to measure brain activity in dyslexic and control subjects in conditions designed to preferentially stimulate the M pathway. Dyslexics showed reduced activity compared with controls both in the primary visual cortex and in a secondary cortical visual area (MT+) that is believed to receive a strong M pathway input. Most importantly, significant correlations were found between individual differences in reading rate and brain activity. These results support the hypothesis for an M pathway abnormality in dyslexia and imply a strong relationship between the integrity of the M pathway and reading ability.

Linear systems analysis of functional magnetic resonance imaging in human V1.

The linear transform model of functional magnetic resonance imaging (fMRI) hypothesizes that fMRI responses are proportional to local average neural activity averaged over a period of time. This work reports results from three empirical tests that support this hypothesis. First, fMRI responses in human primary visual cortex (V1) depend separably on stimulus timing and stimulus contrast. Second, responses to long-duration stimuli can be predicted from responses to shorter duration stimuli. Third, the noise in the fMRI data is independent of stimulus contrast and temporal period. Although these tests can not prove the correctness of the linear transform model, they might have been used to reject the model. Because the linear transform model is consistent with our data, we proceeded to estimate the temporal fMRI impulse-response function and the underlying (presumably neural) contrast-response function of human V1.

Forward pattern masking and adaptation: effects of duration, interstimulus interval, contrast, and spatial and temporal frequency.

Eight experiments are described that compare pattern adaptation and forward pattern masking by examining the effects of five variables on the contrast threshold of a target presented after an adapter or masker. The target is a Gabor pattern with a center frequency of 2 c/deg and a duration of 33 msec. Thresholds are determined using an adaptive spatial forced-choice method. Principal results are as follows. (1) An adapt-refresh regime with a 2 sec refresh and a 2 sec recovery period on each trial is shown to maintain constant performance. (2) Desensitization is very rapid, reaching near maximum in < 200 msec. (3) Recovery is very rapid during the first 100-200 msec and then very slow with the rate of slow recovery decreasing as adapter/masker duration increases. (4) Threshold vs contrast functions are step-like for certain frequency pairs. (5) Sensitivity vs frequency functions derived from adapting and masking are similar in form. (6) Masker temporal frequency (0-15 Hz) has very little effect. These results are described by a theory that postulates that the target is detected by a few mechanisms that are differentially tuned to spatial frequency. The effect of both a forward masker and an adapter is to desensitize the mechanisms that respond to it. Recovery is a weighted sum of two decay processes, one fast and one slow. The theory fits the data from both paradigms well with some differences in parameters.