Biomedical subjects

Gordon L Shulman

Publications and source records attributed to Gordon L Shulman.

17 recordsLinked to original sources

PubMed2006

Attentional selection of moving objects by a serial process.

How does the efficiency of attentional selection depend on the number of attended objects in a display? We measured the channel capacity (CC) of human observers during the attentional tracking of moving targets. The relation between CC and target number was used to estimate target-sampling rate. The sampling rate was halved when the number of targets was doubled, indicating that tracking was accomplished by a mechanism whose processing rate did not vary with target number. Systematically varying the dynamic parameters of the display provided inconclusive evidence for the idea that the time interval between successive samples of the same target increased with target number. These results are consistent with the hypothesis that the selection of multiple moving objects involves a limited capacity processor.

Attention↗

PubMed2006

Separate modulations of human V1 associated with spatial attention and task structure.

Functional magnetic resonance imaging (fMRI) was used while normal human volunteers engaged in simple detection and discrimination tasks, revealing separable modulations of early visual cortex associated with spatial attention and task structure. Both modulations occur even when there is no change in sensory stimulation. The modulation due to spatial attention is present throughout the early visual areas V1, V2, V3, and VP, and varies with the attended location. The task structure activations are strongest in V1 and are greater in regions that represent more peripheral parts of the visual field. Control experiments demonstrate that the task structure activations cannot be attributed to visual, auditory, or somatosensory processing, the motor response for the detection/discrimination judgment, or oculomotor responses such as blinks or saccades. These findings demonstrate that early visual areas are modulated by at least two types of endogenous signals, each with distinct cortical distributions.

Adult↗

PubMed2006

Visuospatial reorienting signals in the human temporo-parietal junction are independent of response selection.

This study contrasts visuospatial reorienting and response selection signals in the right temporo-parietal junction (TPJ) with functional magnetic resonance imaging. The overall goal was to investigate whether spatial orienting signals and motor signals interacted or were independent in TPJ. The right TPJ showed a greater response to targets at in-validly rather than validly cued locations, but no significant modulation from the effector used to respond. We suggest that TPJ may work as a modality-independent 'circuit breaker' for the dorsal fronto-parietal attention system, directing attention to salient events and enabling a variety of responses to those events.

Adult↗

PubMed2005

Brain signals for spatial attention predict performance in a motion discrimination task.

The reliability of visual perception is thought to reflect the quality of the sensory information. However, we show that subjects' performance can be predicted, trial-by-trial, by neural activity that precedes the onset of a sensory stimulus. Using functional MRI (fMRI), we studied how neural mechanisms that mediate spatial attention affect the accuracy of a motion discrimination judgment. The amplitude of blood oxygen level-dependent (BOLD) signals after a cue directing spatial attention predicted subjects' accuracy on 60-75% of the trials. Widespread predictive signals, which included dorsal parietal, visual extra-striate, prefrontal and sensory-motor cortex, depended on whether the cue correctly specified the stimulus location. Therefore, these signals indicate the degree of utilization of the cued information and play a role in the control of spatial attention. We conclude that variability in perceptual performance can be partly explained by the variability in endogenous, preparatory processes and that BOLD signals can be used to forecast human behavior.

Attention↗

PubMed2005

Evidence for separate perceptual reactivation and search processes during remembering.

Remembering involves the coordinated recruitment of strategic search processes and processes involved in reconstructing the content of the past experience. In the present study we used a cueing paradigm based on event-related functional magnetic resonance imaging to separate activity in the initial preparation phases of retrieval from later phases during which retrieval search ensued, and detailed auditory and visual memories were reconstructed. Results suggest a dissociation among inferior temporal (IT) and parieto-occipital (PO) processing regions in how they were influenced by preparatory cues prior to remembering, and indicate a dissociation in how they were influenced by the subsequent validity of those cues during remembering. Regions in IT cortex appeared to show search-related activity during retrieval, as well as robust modality effects, but they were not influenced by preparatory cues. These findings suggest a specific role for IT regions in reconstruction of visual details during remembering. While dorsal regions in parietal and superior occipital cortex also appeared to show search-related activity as well as robust modality effects, they were also influenced by preparatory cues during the retrieval phase, and to a lesser degree during the cue phase. These findings indicate a role in integrating perceptual reactivation and search processes during remembering.

Adaptation, Physiological↗

PubMed2005

An event-related functional magnetic resonance imaging study of voluntary and stimulus-driven orienting of attention.

Attention can be voluntarily directed to a location or automatically summoned to a location by a salient stimulus. We compared the effects of voluntary and stimulus-driven shifts of spatial attention on the blood oxygenation level-dependent signal in humans, using a method that separated preparatory activity related to the initial shift of attention from the subsequent activity caused by target presentation. Voluntary shifts produced greater preparatory activity than stimulus-driven shifts in the frontal eye field (FEF) and intraparietal sulcus, core regions of the dorsal frontoparietal attention network, demonstrating their special role in the voluntary control of attention. Stimulus-driven attentional shifts to salient color singletons recruited occipitotemporal regions, sensitive to color information and part of the dorsal network, including the FEF, suggesting a partly overlapping circuit for endogenous and exogenous orienting. The right temporoparietal junction (TPJ), a core region of the ventral frontoparietal attention network, was strongly modulated by stimulus-driven attentional shifts to behaviorally relevant stimuli, such as targets at unattended locations. However, the TPJ did not respond to salient, task-irrelevant color singletons, indicating that behavioral relevance is critical for TPJ modulation during stimulus-driven orienting. Finally, both ventral and dorsal regions were modulated during reorienting but significantly only by reorienting after voluntary shifts, suggesting the importance of a mismatch between expectation and sensory input.

Adult↗

PubMed2005

A functional MRI study of preparatory signals for spatial location and objects.

We investigated preparatory signals for spatial location and objects in normal observers using functional magnetic resonance imaging (fMRI). Activity for attention-directing cues was separated from activity for subsequent test arrays containing the target stimulus. Subjects were more accurate in discriminating a target face among distracters when they knew in advance its location (spatial directional cue), as compared to when the target could randomly appear at one of two locations (spatial neutral cue), indicating that the spatial cue was used. Spatially specific activations occurred in a region at the intersection of the ventral intraparietal sulcus and transverse occipital sulcus (vIPS-TOS), which showed significantly stronger activation for rightward- than leftward-directing cues, while other fronto-parietal areas were activated by the cue but did not show spatial specificity. In visual cortex, activity was weak or absent in retinotopic occipital regions following attention-directing cues and this activity was not spatially specific. In a separate task, subject discriminated a target outdoor scene among distracters after the presentation of spatial neutral cues. There was no significant difference in dorsal frontoparietal activity during the face versus scene discrimination task. Also, there was only weak evidence for selective preparatory activity in ventral object-selective regions, although the activation of these regions to the subsequent test array did depend upon which discrimination (face or place) was performed. We conclude first that under certain circumstances, spatial cues that produce strong behavioral effects may modulate parietal-occipital regions in a spatially specific manner without producing similar modulations in retinotopic occipital regions. Second, attentional modulations of object-selective regions in temporal-occipital cortex can occur even though preparatory object-selective modulations of those regions are absent or weak.

Analysis of Variance↗

PubMed2004

Extrastriate body area in human occipital cortex responds to the performance of motor actions.

A region in human lateral occipital cortex (the 'extrastriate body area' or EBA) has been implicated in the perception of body parts. Here we report functional magnetic resonance imaging (fMRI) evidence that the EBA is strongly modulated by limb (arm, foot) movements to a visual target stimulus, even in the absence of visual feedback from the movement. Therefore, the EBA responds not only during the perception of other people's body parts, but also during goal-directed movements of the observer's body parts. In addition, both limb movements and saccades to a detected stimulus produced stronger signals than stimulus detection without motor movements ('covert detection') in the calcarine sulcus and lingual gyrus. These motor-related modulations cannot be explained by simple visual or attentional factors related to the target stimulus, and suggest a potentially widespread influence of actions on visual cortex.

Adult↗

PubMed2003

Quantitative analysis of attention and detection signals during visual search.

Prior work has distinguished regions in the intraparietal sulcus (IPs) and frontal eye field (FEF) involved in the voluntary control of attention, from more ventral regions in the temporoparietal junction (TPJ) involved in target detection. The present results show that when subjects search for and detect a visual target stimulus among nontargets, these regions show sensory-, search-, and detection-related signals that both confirm and refine these functional distinctions. The different signals were isolated by an additive model that accounted for a large fraction of BOLD (blood oxygenation level-dependent) signal modulation over the brain. Both IPs and FEF were activated during search through nontargets, consistent with a role in maintaining attention-related signals during search. However, unlike FEF, IPs also showed stimulus-related activations, and may combine signals related to sensory and task-dependent components of salience. Although IPs-FEF showed search-related activations, the TPJ was deactivated during search. TPJ activations were confined to detection-related signals. These results provide a much stronger dissociation between the TPJ and IPs-FEF than previous work, while indicating functional differences between frontal and parietal regions that are often coactivated in studies of attention. Finally, continuous flow models of information processing predict that during search, signals from missed targets should be fed from sensory to associative regions rather than being gated by the decision criterion. Correspondingly, missed targets significantly activated parietal (e.g., right TPJ) and frontal (e.g., anterior insula, anterior cingulate) regions, although with a smaller magnitude than detected targets. Surprisingly, many cortical regions showed equivalent signals from detected targets and the completion of target-absent trials, reflecting a widespread signal unrelated to motor execution.

Attention↗

PubMed2003

Functional organization of human intraparietal and frontal cortex for attending, looking, and pointing.

We studied the functional organization of human posterior parietal and frontal cortex using functional magnetic resonance imaging (fMRI) to map preparatory signals for attending, looking, and pointing to a peripheral visual location. The human frontal eye field and two separate regions in the intraparietal sulcus were similarly recruited in all conditions, suggesting an attentional role that generalizes across response effectors. However, the preparation of a pointing movement selectively activated a different group of regions, suggesting a stronger role in motor planning. These regions were lateralized to the left hemisphere, activated by preparation of movements of either hand, and included the inferior and superior parietal lobule, precuneus, and posterior superior temporal sulcus, plus the dorsal premotor and anterior cingulate cortex anteriorly. Surface-based registration of macaque cortical areas onto the map of fMRI responses suggests a relatively good spatial correspondence between human and macaque parietal areas. In contrast, large interspecies differences were noted in the topography of frontal areas.

Animals↗

PubMed2003

Identification of cerebral networks by classification of the shape of BOLD responses.

Changes in regional blood oxygen level dependent (BOLD) signals in response to brief visual stimuli can exhibit a variety of time-courses. To demonstrate the anatomical distribution of BOLD response shapes during a match to sample task, a formal analysis of their time-courses is presented. An event-related design was used to estimate regional BOLD responses evoked by a cue word, which instructed the subject to attend to the motion or color of an upcoming target, and those evoked by a briefly presented moving target consisting of colored dots. Regional BOLD time-courses were adequately represented by the linear combination of three orthogonal waveforms. BOLD response shapes were then classified using a fuzzy clustering scheme. Three classes (sustained, phasic, and negative) best characterized cue responses. Four classes (sustained, sustained-phasic, phasic, and bi-phasic) best characterized target responses. In certain regions, the shape of the BOLD responses was modulated by the instruction to attend to the target's motion or color. A left frontal and a posterior parietal region showed sustained activity when motion was cued and transient activity when color was cued. A right thalamic and a left lateral occipital region showed sustained activity when color was cued and transient activity when motion was cued. Following the target several regions showed more sustained activity during motion than color trials. In summary, the effect of the task variable was focal following the cue and widespread following the target. We conclude that the temporal patterns of neural activity affected the shape of the BOLD signal.

Brain↗

PubMed2003

Persistence and brain circuitry.

The orbitofrontal and adjacent medial prefrontal cortex may play an important role in normal social functioning and affect modulation. Recent anatomical studies of this area of the prefrontal cortex have demonstrated a striking correspondence of fine-grained architectonic partitioning schemes in humans and nonhuman primates. This finding allows neurophysiological recording and anatomical connectivity data in animals to be considered together with functional imaging data and lesion studies in humans. In a functional MRI study, we show that individual differences in Persistence, a dimensional trait assessed with a seven-factor personality model, may be linked to specific areas in the lateral orbital and medial prefrontal cortex and the ventral striatum. These areas are part of an anatomical circuit that has been defined in nonhuman primates and has been implicated in functions related to behavioral persistence. These findings represent a fresh approach to linking normal individual differences in personality and behavior to specific neuronal structures and subsystems.

Adult↗

PubMed2002

Neural systems for visual orienting and their relationships to spatial working memory.

We investigated neural correlates of human visual orienting using event-related functional magnetic resonance imaging (fMRI). When subjects voluntarily directed attention to a peripheral location, we recorded robust and sustained signals uniquely from the intraparietal sulcus (IPs) and superior frontal cortex (near the frontal eye field, FEF). In the ventral IPs and FEF only, the blood oxygen level dependent signal was modulated by the direction of attention. The IPs and FEF also maintained the most sustained level of activation during a 7-sec delay, when subjects maintained attention at the peripheral cued location (working memory). Therefore, the IPs and FEF form a dorsal network that controls the endogenous allocation and maintenance of visuospatial attention. A separate right hemisphere network was activated by the detection of targets at unattended locations. Activation was largely independent of the target's location (visual field). This network included among other regions the right temporo-parietal junction and the inferior frontal gyrus. We propose that this cortical network is important for reorienting to sensory events.

Adolescent↗

PubMed2002

Time-related increase of oxygen utilization in continuously activated human visual cortex.

Oxygen utilization increase is fractionally much less than that seen in glucose metabolism and blood flow soon after onset of neuronal activation, however its behavior during continued activation is less certain. We evaluated the effects of 25 min of visual stimulation on CBF, CMRO(2), and OEF using [(15)O] water and [(15)O] oxygen PET. Seven healthy volunteers underwent a PET session consisting of serial [(15)O] water and [(15)O] oxygen scans at the fixation-only baseline visual state and after 1, 13, and 25 min of the continuous visual stimulation using a black-white vertical grating. CBF, CMRO(2), and OEF values were calculated for the entire brain and for regions of interest in visual cortex centered over the area of activation. After 1 min of stimulation, CMRO(2) increased only 4.7% compared to baseline and CBF increased 40.7%. However, after 25 min of stimulation the increase in CMRO(2) compared to baseline was 15.0%, having tripled from that measured at 1 min (P < 0.05). CBF did not significantly change during this time. OEF was 48.3% at baseline. It decreased to 37.1% after 1 min of visual stimulation (P < 0.01) and then returned almost to baseline values after 25 min of activation OEF (45.7%). There were no significant variations in whole-brain values during the study. We suggest that in the activated brain, the increased energy demands initially are not fully met with oxidative metabolism and must predominantly be supported by increased glycolysis. With continued activation, oxygen utilization increases reducing the need for excess glycolysis.

Adult↗

PubMed2002

Control of goal-directed and stimulus-driven attention in the brain.

We review evidence for partially segregated networks of brain areas that carry out different attentional functions. One system, which includes parts of the intraparietal cortex and superior frontal cortex, is involved in preparing and applying goal-directed (top-down) selection for stimuli and responses. This system is also modulated by the detection of stimuli. The other system, which includes the temporoparietal cortex and inferior frontal cortex, and is largely lateralized to the right hemisphere, is not involved in top-down selection. Instead, this system is specialized for the detection of behaviourally relevant stimuli, particularly when they are salient or unexpected. This ventral frontoparietal network works as a 'circuit breaker' for the dorsal system, directing attention to salient events. Both attentional systems interact during normal vision, and both are disrupted in unilateral spatial neglect.

Animals↗

PubMed2002

Two attentional processes in the parietal lobe.

We report fMRI evidence for two attentional processes in parietal cortex. Subjects matched a feature, cued by a word, to a test display of moving colored dots. Either color (red, green) or motion direction (left, right) was cued on mixed scans while only one dimension was cued on blocked scans. An event-related paradigm separated the preparatory activity generated by the cue from the subsequent activity related to the test display. One attentional process specified task information while a second process was motion selective. During the cue period, a pure effect of task specification was observed in left frontal cortex while combined effects of task specification and motion selectivity were observed in left posterior parietal cortex. The frontal task-specification signal may have been the source of the corresponding signal in parietal cortex. Effects of task specification generalized over cue dimension, indicating that the information was coded in a sufficiently abstract form to affect color and motion processing. During the subsequent test period, task-specification and motion-selective signals were again observed in left parietal cortex. Task specification did not significantly affect occipital motion-selective regions, such as MT+, however, indicating that this process did not influence the lower cortical tier of the motion processing stream. These results provide evidence for general and specialized task representations within left parietal cortex during task preparation and execution.

Analysis of Variance↗

PubMed2002

Reactivation of networks involved in preparatory states.

We report an endogenous signal that has a widespread cortical distribution and is time-locked to the termination of a sustained state of task-readiness. In three event-related functional magnetic resonance imaging (fMRI) experiments, subjects saw an arrow cue that predicted either the direction of motion or the location of a subsequent test stimulus. A reactivation of the BOLD (blood oxygenation level-dependent) signal occurred at the termination of the state of readiness in occipital regions that were transiently activated by the cue and in frontal-parietal regions that maintained an attentional set over the trial. Moreover, a delayed activation occurred in prefrontal and temporo-parietal regions that did not initially respond to the cue and that have been implicated in re-orienting attention to novel sensory events. These latter regions may have generated control signals that ended the state of readiness in regions active during the cue period. These results indicate that terminating a state of readiness produces a widely distributed cortical signal and suggest that areas involved in a preparatory state may be maintained as a network which can be modulated as a whole.

Attention↗