Geoffrey M. Boynton

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.

Global effects of feature-based attention in human visual cortex.

The content of visual experience depends on how selective attention is distributed in the visual field. We used functional magnetic resonance imaging (fMRI) in humans to test whether feature-based attention can globally influence visual cortical responses to stimuli outside the attended location. Attention to a stimulus feature (color or direction of motion) increased the response of cortical visual areas to a spatially distant, ignored stimulus that shared the same feature.

Adaptation: from single cells to BOLD signals

Functional magnetic resonance imaging adaptation (fMRIa) is an increasingly popular method that aims to provide insight into the functional properties of subpopulations of neurons within an imaging voxel. The technique relies on the assumption that neural adaptation reduces activity when two successive stimuli activate the same subpopulation but not when they stimulate different subpopulations. Here, we assess the validity of fMRIa by comparing single-cell recordings with functional imaging of orientation, motion and face processing. We find that fMRIa provides novel insight into neural representations in the human brain. However, network responses in general and adaptation in particular are more complex than is often assumed, and an unequivocal interpretation of fMRIa results can be achieved only with great care.

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 observers 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.

Feature-Based Attentional Modulations in the Absence of Direct Visual Stimulation

When faced with a crowded visual scene, observers must selectively attend to behaviorally relevant objects to avoid sensory overload. Often this selection process is guided by prior knowledge of a target-defining feature (e.g., the color red when looking for an apple), which enhances the firing rate of visual neurons that are selective for the attended feature. Here, we used functional magnetic resonance imaging and a pattern classification algorithm to predict the attentional state of human observers as they monitored a visual feature (one of two directions of motion). We find that feature-specific attention effects spread across the visual field-even to regions of the scene that do not contain a stimulus. This spread of feature-based attention to empty regions of space may facilitate the perception of behaviorally relevant stimuli by increasing sensitivity to attended features at all locations in the visual field.

Individual Differences among Grapheme-Color Synesthetes: Brain-Behavior Correlations

Grapheme-color synesthetes experience specific colors associated with specific number or letter characters. To determine the neural locus of this condition, we compared behavioral and fMRI responses in six grapheme-color synesthetes to control subjects. In our behavioral experiments, we found that a subjects synesthetic experience can aid in texture segregation (experiment 1) and reduce the effects of crowding (experiment 2). For synesthetes, graphemes produced larger fMRI responses in color-selective area human V4 than for control subjects (experiment 3). Importantly, we found a correlation within subjects between the behavioral and fMRI results; subjects with better performance on the behavioral experiments showed larger fMRI responses in early retinotopic visual areas (V1, V2, V3, and hV4). These results suggest that grapheme-color synesthesia is the result of cross-activation between grapheme-selective and color-selective brain areas. The correlation between the behavioral and fMRI results suggests that grapheme-color synesthetes may constitute a heterogeneous group.

Cortical magnification within human primary visual cortex correlates with acuity thresholds

We measured linear cortical magnification factors in V1 with fMRI, and we measured visual acuity (Vernier and grating) in the same observers. The cortical representation of both Vernier and grating acuity thresholds in V1 was found to be roughly constant across all eccentricities. We also found a within-observer correlation between cortical magnification and Vernier acuity, further supporting claims that Vernier acuity is limited by cortical magnification in V1.

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.

Efficient Design of Event-Related fMRI Experiments Using M-Sequences

Rapid event-related fMRI (erfMRI) allows estimation of the shape of hemodynamic responses (HDR) associated with transient brain activation evoked by various sensory, motor, and cognitive events. Choosing a sequence of events that maximizes efficiency of estimating the HDR is essential for conducting event-related brain imaging experiments, since increasing efficiency is essentially equivalent to reducing scanning time or increasing the strength of the principal magnetic field. The efficiency of an erfMRI design depends critically on the temporal arrangement of the sequence of events and the noise in the fMRI signal. We introduce to erfMRI a simple method for generating efficient event sequences based on maximum-length shift register sequences, or m-sequences. We show that under the assumption of white uncorrelated MRI noise, efficiency of erfMRI experimental designs that employ m-sequences exceeds efficiency of the best randomly generated sequences. This is true for single and multiple event type experiments, which allow either parallel events (overlapping events design) or designs in which only one event occurs at a time (nonoverlapping events design). HDR estimation efficiency afforded by m-sequences grows with the number of event types, and is greatest when event sequences are relatively short, albeit within commonly used scan times (i.e., 63-255 total events per scan). The improvement in efficiency, however, comes at a cost of constraints imposed by m-sequence generation rules, such as predetermined sequence lengths; for nonoverlapping events design m-sequence-based designs are not available for all possible numbers of event types. Nevertheless, designs that are available with m-sequences cover a large subset of commonly used erfMRI experimental designs. Under conditions of characteristic time-correlated fMRI noise, randomly generated sequences may yield efficiencies that exceed those afforded by m-sequences for single event-type designs, since in this case one can generate random sequences that partially decorrelate MRI noise by chance. Our simulations suggest that for designs of realistic sequence lengths that use more than one event type, m-sequence based designs tend to outperform random designs, thus making the knowledge of noise inessential. Finally, within an r-th order m-sequence (generated by a shift register of length r) all possible combinations of subsequences of length r occur, and thus these subsequences are exactly counterbalanced. This property is essential for minimizing effects of psychological and neuronal adaptation and expectation.

Global feature-based attention for motion and color

We used a divided attention psychophysical task to test the hypothesis that visual attention to a stimulus feature(1) facilitates the processing of other stimuli sharing the same feature. Performance on a dual-task was significantly better when human observers divided attention across two spatially separate stimuli sharing a common feature (same direction of motion or same color) compared to opposing features. This attentional effect was dependent upon the presence of competing stimuli. These results are consistent with a spatially global feature-based mechanism of attention that increases the response of cortical neurons tuned to an attended feature throughout the visual field.