Deductive reasoning is fundamental to science, human culture, and the solution of problems in daily life. It starts with premises and yields a logically necessary conclusion that is not explicit in the premises. Here we investigated the neurocognitive processes underlying logical thinking with event-related functional magnetic resonance imaging. We specifically focused on three temporally separable phases: (1) the premise processing phase, (2) the premise integration phase, and (3) the validation phase in which reasoners decide whether a conclusion logically follows from the premises. We found distinct patterns of cortical activity during these phases, with initial temporo-occipital activation shifting to the prefrontal cortex and then to the parietal cortex during the reasoning process. Activity in these latter regions was specific to reasoning, as it was significantly decreased during matched working memory problems with identical premises and equal working memory load.
Attending to the location of an expected visual target can lead to anticipatory activations in spatiotopic occipital cortex, emerging before target onset. But less is known about how the brain may prepare for a distractor at a known location remote from the target. In a psychophysical experiment, we found that trial-to-trial advance knowledge about the presence of a distractor in the target-opposite hemifield significantly reduced its behavioral cost. In a subsequent functional magnetic resonance imaging experiment with similar task and stimuli, we found anticipatory activations in the occipital cortex contralateral to the expected distractor, but no additional target modulation, when participants were given advance information about a distractor's subsequent presence and location. Several attention-related control structures (frontal eye fields and superior parietal cortex) were active during attentional preparation for all trials, whereas the left superior prefrontal and right angular gyri were additionally activated when a distractor was anticipated. The right temporoparietal junction showed stronger functional coupling with occipital regions during preparation for trials with an isolated target than for trials with a distractor expected. These results show that anticipation of a visual distractor at a known location, remote from the target, can lead to (1) a reduction in the behavioral cost of that distractor, (2) preparatory modulation of the occipital cortex contralateral to the location of the expected distractor, and (3) anticipatory activation of distinct parietal and frontal brain structures. These findings indicate that specific components of preparatory visual attention may be devoted to minimizing the impact of distractors, not just to enhancements of target processing.
We used fMRI to investigate competition and on-line attentional selection between targets and distractors in opposite visual hemifields. Displays comprised a high-contrast square-wave grating, defined as target by its orientation, presented alone (unilateral) or with a similar distractor of orthogonal orientation in the opposite hemifield (bilateral displays). The target appeared unpredictably on the left or right, precluding anticipatory attention to one side. We found greater activation in target-contralateral superior occipital gyrus for unilateral than for bilateral displays, indicating suppression of the target's visual representation by distractor presence despite the competing distractor projecting to a different occipital hemisphere. Several frontal and parietal regions showed greater activation for bilateral than unilateral trials, suggesting involvement in on-line attentional selection. This was particularly pronounced for regions in bilateral intraparietal sulcus (IPS), which also showed greater functional coupling with occipital cortex specifically on bilateral trials that required selection plus some repetition-suppression effects when target side was repeated, but again only on bilateral trials requiring selection. Our results indicate that competition between visual stimuli in opposite hemifields can influence occipital cortex, and implicate IPS in resolution of this competition by selection.
Our results provide causal evidence that circuits originating in the human FEF can modulate activity in retinotopic visual cortex, in a manner that differentiates the central and peripheral visual field, with functional consequences for perception. More generally, our study illustrates how the new approach of concurrent TMS-fMRI can now reveal causal interactions between remote but interconnected areas of the human brain.
The neural processes underlying tactile decisions in the human brain remain elusive. We addressed this question in a functional magnetic resonance imaging study using a somatosensory discrimination task, requiring participants to compare the frequency of two successive tactile stimuli. Tactile stimuli per se engaged somatosensory, parietal, and frontal cortical regions. Using a statistical model that accounted for the relative difference in frequencies (i.e., Weber fraction) and discrimination accuracy (i.e., correct or incorrect), we show that trial-by-trial relative frequency difference is represented linearly by activity changes in the left dorsolateral prefrontal cortex (DLPFC), the dorsal anterior cingulate cortex, and bilateral anterior insular cortices. However, a circumscribed region within the left DLPFC showed a different response pattern expressed as activity changes that were monotonically related to relative stimulation difference only for correct but not for incorrect trials. Our findings suggest that activity in the left DLPFC encodes stimulus representations that underlie veridical tactile decisions in humans.
Conscious perception depends not only on sensory input, but also on attention [1, 2]. Recent studies in monkeys [3-6] and humans [7-12] suggest that influences of spatial attention on visual awareness may reflect top-down influences on excitability of visual cortex. Here we tested this specifically, by providing direct input into human visual cortex via cortical transcranial magnetic stimulation (TMS) to produce illusory visual percepts, called phosphenes. We found that a lower TMS intensity was needed to elicit a conscious phosphene when its apparent spatial location was attended, rather than unattended. Our results indicate that spatial attention can enhance visual-cortex excitability, and visual awareness, even when sensory signals from the eye via the thalamic pathway are bypassed.
Iconic memory and spatial attention are often considered separately, but they may have functional similarities. Here we provide functional magnetic resonance imaging evidence for some common underlying neural effects. Subjects judged three visual stimuli in one hemifield of a bilateral array comprising six stimuli. The relevant hemifield for partial report was indicated by an auditory cue, administered either before the visual array (precue, spatial attention) or shortly after the array (postcue, iconic memory). Pre- and postcues led to similar activity modulations in lateral occipital cortex contralateral to the cued side. This finding indicates that readout from iconic memory can have some neural effects similar to those of spatial attention. We also found common bilateral activation of a fronto-parietal network for postcue and precue trials. These neuroimaging data suggest that some common neural mechanisms underlie selective spatial attention and readout from iconic memory. Some differences were also found; compared with precues, postcues led to higher activity in the right middle frontal gyrus.
It has often been proposed that regions of the human parietal and/or frontal lobe may modulate activity in visual cortex, for example, during selective attention or saccade preparation. However, direct evidence for such causal claims is largely missing in human studies, and it remains unclear to what degree the putative roles of parietal and frontal regions in modulating visual cortex may differ. Here we used transcranial magnetic stimulation (TMS) and functional magnetic resonance imaging (fMRI) concurrently, to show that stimulating right human intraparietal sulcus (IPS, at a site previously implicated in attention) elicits a pattern of activity changes in visual cortex that strongly depends on current visual context. Increased intensity of IPS TMS affected the blood oxygen level-dependent (BOLD) signal in V5/MT+ only when moving stimuli were present to drive this visual region, whereas TMS-elicited BOLD signal changes were observed in areas V1-V4 only during the absence of visual input. These influences of IPS TMS upon remote visual cortex differed significantly from corresponding effects of frontal (eye field) TMS, in terms of how they related to current visual input and their spatial topography for retinotopic areas V1-V4. Our results show directly that parietal and frontal regions can indeed have distinct patterns of causal influence upon functional activity in human visual cortex.
During voluntary action, dorsal premotor cortex (PMd) may exert influences on motor regions in both hemispheres, but such interregional interactions are not well understood. We used transcranial magnetic stimulation (TMS) concurrently with event-related functional magnetic resonance imaging to study such interactions directly. We tested whether causal influences from left PMd upon contralateral (right) motor areas depend on the current state of the motor system, involving regions engaged in a current task. We applied short bursts (360 ms) of high- or low-intensity TMS to left PMd during single isometric left-hand grips or during rest. TMS to left PMd affected activity in contralateral right PMd and primary motor cortex (M1) in a state-dependent manner. During active left-hand grip, high (vs. low)-intensity TMS led to activity increases in contralateral right PMd and M1, whereas activity decreases there due to TMS were observed during no-grip rest. Analyses of condition-dependent functional coupling confirmed topographically specific stronger coupling between left PMd and right PMd (and right M1), when high-intensity TMS was applied to left PMd during left-hand grip. We conclude that left PMd can exert state-dependent interhemispheric influences on contralateral cortical motor areas relevant for a current motor task.
Reinforcing effects of reward on action are well established, but possible effects on sensory function are less well explored. Here, using functional magnetic resonance imaging, we assessed whether reward can influence somatosensory judgments and modulate activity in human somatosensory cortex. Participants discriminated electrical somatosensory stimuli on an index finger with correct performance rewarded financially at trial end, at one of four different anticipated levels. Higher rewards improved tactile performance and led to increased hemodynamic signals from ventral striatum on rewarded trials. Remarkably, primary somatosensory cortex contralateral to the judged hand was reactivated at the point of reward delivery, despite the absence of concurrent somatosensory input at that time point. This side-specific reactivation of primary somatosensory cortex increased monotonically with level of reward. Moreover, the level of reward received on a particular trial influenced somatosensory performance and neural activity on the subsequent trial, with better discrimination and enhanced hemodynamic response in contralateral primary somatosensory cortex for trials that followed higher rewards. These results indicate that rewards can influence not only classical reward-related regions, but also early somatosensory cortex when a decision is required for that modality.
