Tony Ro

To See or Not to See: Prestimulus α Phase Predicts Visual Awareness

We often fail to see something that at other times is readily detectable. Because the visual stimulus itself is unchanged, this variability in conscious awareness is likely related to changes in the brain. Here we show that the phase of EEG α rhythm measured over posterior brain regions can reliably predict both subsequent visual detection and stimulus-elicited cortical activation levels in a metacontrast masking paradigm. When a visual target presentation coincides with the trough of an α wave, cortical activation is suppressed as early as 100 ms after stimulus onset, and observers are less likely to detect the target. Thus, during one α cycle lasting 100 ms, the human brain goes through a rapid oscillation in excitability, which directly influences the probability that an environmental stimulus will reach conscious awareness. Moreover, ERPs to the appearance of a fixation cross before the target predict its detection, further suggesting that cortical excitability level may mediate target detection. A novel theory of cortical inhibition is proposed in which increased α power represents a “pulsed inhibition” of cortical activity that affects visual awareness.

Changing Faces: A Detection Advantage in the Flicker Paradigm

Observers seem surprisingly poor at detecting changes in images following a large transient or flicker. In this study, we compared this change blindness phenomenon between human faces and other common objects (e.g., clothes). We found that changes were detected far more rapidly and accurately in faces than in other objects. This advantage for faces, however, was found only for upright faces in multiple-object arrays, and was completely eliminated when displays showed one photograph only or when the pictures were inverted. These results suggest a special role for faces in competition for visual attention, and provide support for previous claims that human faces are processed differently than stimuli that may be of less biological significance.

The Role of Perceptual Load in Processing Distractor Faces

It has been established that successful ignoring of irrelevant distractors depends on the extent to which the current task loads attention. However, the previous load studies have typically employed neutral distractor stimuli (e.g., letters). In the experiments reported here, we examined whether the perception of irrelevant distractor faces would show the same effects. We manipulated attentional load in a relevant task of name search by varying the search set size and found that whereas congruency effects from meaningful nonface distractors were eliminated by higher search load, interference from distractor faces was entirely unaffected by search load. These results support the idea that face processing may be mandatory and generalize the load theory to the processing of meaningful and more complex nonface distractors.

Pulsed out of awareness: EEG alpha oscillations represent a pulsed-inhibition of ongoing cortical processing.

Alpha oscillations are ubiquitous in the brain, but their role in cortical processing remains a matter of debate. Recently, evidence has begun to accumulate in support of a role for alpha oscillations in attention selection and control. Here we first review evidence that 8–12 Hz oscillations in the brain have a general inhibitory role in cognitive processing, with an emphasis on their role in visual processing. Then, we summarize the evidence in support of our recent proposal that alpha represents a pulsed-inhibition of ongoing neural activity. The phase of the ongoing electroencephalography can influence evoked activity and subsequent processing, and we propose that alpha exerts its inhibitory role through alternating microstates of inhibition and excitation. Finally, we discuss evidence that this pulsed-inhibition can be entrained to rhythmic stimuli in the environment, such that preferential processing occurs for stimuli at predictable moments. The entrainment of preferential phase may provide a mechanism for temporal attention in the brain. This pulsed inhibitory account of alpha has important implications for many common cognitive phenomena, such as the attentional blink, and seems to indicate that our visual experience may at least some times be coming through in waves.

Constraint-Induced Movement Therapy During Early Stroke Rehabilitation

Background. Limited data are available about the effectiveness of early rehabilitation after stroke. Objective. This is the 1st randomized controlled trial of constraint-induced movement therapy (CIMT) in subacute stroke to investigate neurophysiologic mechanisms and long-term outcome. Methods. Within 2 weeks after stroke, 23 patients with upper extremity (UE) weakness were randomized to 2 weeks of CIMT or traditional therapy at an equal frequency of up to 3 h/day. Motor function of the affected UE was blindly assessed before treatment, after treatment, and 3 months after stroke. Transcranial magnetic stimulation (TMS) measured the cortical area evoking movement of the affected hand. Results. Long-term improvement in motor function of the affected UE did not differ significantly between patients who received CIMT versus intensive traditional therapy. All outcome comparisons showed trends favoring CIMT over intensive traditional therapy, but none was statistically significant except for improvements in the Fugl-Meyer (FM) UE motor scale immediately following treatment and in reported quality of hand function at 3 months. Improvement in UE motor function on the FM was associated with a greater number of sites on the affected cerebral hemisphere where responses of the affected hand were evoked by TMS. Conclusions. Future trials of CIMT during early stroke rehabilitation need greater statistical power, more inclusive eligibility criteria, and improved experimental control over treatment intensity. The relationship between changes in motor function and in evoked motor responses suggests that motor recovery during the 1st 3 months after stroke is associated with increased motor excitability of the affected cerebral hemisphere.

Touch, Sound and Vision in Human Superior Temporal Sulcus

Human superior temporal sulcus (STS) is thought to be a key brain area for multisensory integration. Many neuroimaging studies have reported integration of auditory and visual information in STS but less is known about the role of STS in integrating other sensory modalities. In macaque STS, the superior temporal polysensory area (STP) responds to somatosensory, auditory and visual stimulation. To determine if human STS contains a similar area, we measured brain responses to somatosensory, auditory and visual stimuli using blood-oxygen level dependent functional magnetic resonance imaging (BOLD fMRI). An area in human posterior STS, STSms (multisensory), responded to stimulation in all three modalities. STSms responded during both active and passive presentation of unisensory somatosensory stimuli and showed larger responses for more intense vs. less intense tactile stimuli, hand vs. foot, and contralateral vs. ipsilateral tactile stimulation. STSms showed responses of similar magnitude for unisensory tactile and auditory stimulation, with an enhanced response to simultaneous auditory-tactile stimulation. We conclude that STSms is important for integrating information from the somatosensory as well as the auditory and visual modalities, and could be the human homolog of macaque STP.

Feedback Contributions to Visual Awareness in Human Occipital Cortex

It has traditionally been assumed that processing within the visual system proceeds in a bottom-up, feedforward manner from retina to higher cortical areas. In addition to feedforward processing, it is now clear that there are also important contributions to sensory encoding that rely upon top-down, feedback (reentrant) projections from higher visual areas to lower ones. By utilizing transcranial magnetic stimulation (TMS) in a metacontrast masking paradigm, we addressed whether feedback processes in early visual cortex play a role in visual awareness. We show that TMS of visual cortex, when timed to produce visual suppression of an annulus serving as a metacontrast mask, induces recovery of an otherwise imperceptible disk. In addition to producing disk recovery, TMS suppression of an annulus was greater when a disk preceded it than when an annulus was presented alone. This latter result suggests that there are effects of the disk on the perceptibility of the subsequent mask that are additive and are revealed with TMS of the visual cortex. These results demonstrate spatial and temporal interactions of conscious vision in visual cortex and suggest that a prior visual stimulus can influence subsequent perception at early stages of visual encoding via feedback projections.

Human MST But Not MT Responds to Tactile Stimulation

Previous reports of tactile responses in human visual area MT/V5 have used complex stimuli, such as a brush stroking the arm. These complex moving stimuli are likely to induce imagery of visual motion, which is known to be a powerful activator of MT. The area described as “MT” in previous reports consists of at least two distinct cortical areas, MT and MST. Using functional magnetic resonance imaging, we separately localized human MT and MST and measured their response to vibrotactile stimuli unlikely to induce imagery of visual motion. Strong vibrotactile responses were observed in MST but not in MT. Vibrotactile responses in MST were approximately one-half as large as the response to visual motion and were distinct from those in another visual area previously reported to respond to tactile stimulation, the lateral occipital complex. To examine somatotopic organization, we separately stimulated the left and right hand and foot. No spatial segregation between hand and foot responses was observed in MST. The average response profile of MST was similar to that of somatosensory cortex, with a strong preference for the contralateral hand. These results offer evidence for the existence of somatosensory responses in MST, but not MT, independent of imagery of visual motion.

Localization of the human frontal eye fields and motor hand area with transcranial magnetic stimulation and magnetic resonance imaging.

We localized the neuroanatomical correlates for control of saccadic eye movements and for finger movements using a combined transcranial magnetic stimulation (TMS) and magnetic resonance imaging (MRI) approach. Two participants underwent TMS while performing an endogenous saccade task. The motor hand area was localized by TMS and the region anterior to it was mapped to identify the borders of a region where TMS produced delays in generating contralateral saccades. MRI scans were then obtained with fiducial markers placed over the motor hand area and 2 cm anterior to it, the common cortical region that produced saccadic delays in these two subjects. It was also shown that the structural anatomy of the hand area, physiologically defined by visible contractions of the contralateral hand following TMS, corresponded to the knob-like structure recently reported [18, 19]. These results demonstrate that TMS can be a precise, non-invasive tool for neuroanatomical mapping of cortical structures when combined with structural images of the brain.

Inhibition of return and the human frontal eye fields

Inhibition of return (IOR) is a bias against reorienting attention to a previously cued location. In this study, using single-pulse transcranial magnetic stimulation (TMS), we show that the human frontal eye fields (FEF) play a crucial role in the generation of IOR. When TMS was applied over the right FEF at a time interval after a visual cue but shortly before the target, IOR was modulated in the hemifield ipsilateral to the TMS such that responses to a previously cued target were no longer slower than responses to uncued targets. Control TMS over the superior parietal lobule, as well as TMS of the FEF shortly after the cue but well before the target, had no influence on IOR. We further show that the FEF is involved with visual selection as responses to targets appearing contralateral to the TMS of the FEF, but not the control site, were delayed. These results suggest that the FEF produces IOR by biasing attention and eye movements away from a previously attended location and facilitating target detection at novel locations.