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The when and where of spatial storage in memory-guided saccades
Abstract
The memory-guided saccade paradigm is an ideal experimental model for studying spatial working memory. Both the posterior parietal cortex and frontal cortex are known to play a role in working memory; however, there is much debate about the degree of their involvement in the retention of information. We used event-related potentials and electromagnetic tomography to clarify the precise time course and location of the neural correlates of spatial working memory during a memory-guided saccade task in humans. We observed sustained activity in the inferior parietal lobe and extrastriate areas that persisted for the entire duration of the sensory- and memory-phases. This time course reveals that these regions participate in both initial sensory processing of visual cues and in the short-term maintenance of spatial location memory. Similar sustained activation was also observed in the anterior cingulate cortex, probably reflecting attentive control during the task. Differential activity between conditions was also recorded in the dorsolateral prefrontal cortex and in the frontal eye fields, but only during the initial part of the memory-phase. This finding suggests that these areas are not involved in the storage of spatial information, but rather in response selection and in transformation of spatial information into a motor coordinate framework, respectively. By exploiting techniques that provide exquisite temporal resolution and reasonably precise anatomical localization, this study provides evidence supporting the key role of inferior parietal lobe in the storage of spatial information during a working memory guided saccade.
... Both frontal and posterior parietal cortex are involved in working memory. Brignani et al. (2010) utilized event-related potentials and brain tomography localization to shed light on the exact timing and location of the neural correlates of spatial working memory while performing a memory-guided saccade and visually guided saccade task. Sustained activity observed in the inferior parietal lobe and extrastriate regions lasted for the whole duration of the memory and sensory phases. ...
... A Network of brain areas such as frontal and posterior parietal cortex are associated with working memory (Brignani et al., 2010). ...
... Our findings confirm previous work that has shown lateral intraparietal regions code the saccade target locations (Steenrod et al., 2013). This finding confirms previous work that has shown that the inferior parietal lobe contributes to short-term spatial memory maintenance also in the storage of spatial information in a WMGS task (Brignani et al., 2010). ...
Working memory (WM) can be considered as a limited-capacity system which is capable of saving information temporarily with the aim of processing. The aim of the present study was to establish whether eccentricity representation in WM could be decoded from eletroencephalography (EEG) alpha-band oscillation in parietal cortex during delay-period while performing memory-guided saccade (MGS) task. In this regard, we recorded EEG and Eye-tracking signals of 17 healthy volunteers in a variant version of MGS task. We designed the modified version of MGS task for the first time to investigate the effect of locating stimuli in two different positions, in a near (6°) eccentricity and far (12°) eccentricity on saccade error as a behavioral parameter. Another goal of study was to discern whether or not varying the stimuli loci can alter behavioral and eletroencephalographical data while performing the variant version of MGS task. Our findings demonstrate that saccade error for the near fixation condition is significantly smaller than the far from fixation condition. We observed an increase in alpha power in parietal lobe in near vs far conditions. In addition, the results indicate that the increase in alpha (8–12 Hz) power from fixation to memory was negatively correlated with saccade error. The novel approach of using simultaneous EEG/Eye-tracking recording in the modified MGS task provided both behavioral and electroencephalographic analyses for oscillatory activity during this new version of MGS task.
... Accordingly, our tasks imposed memory delays between 500 and 1,500 ms, which are within a range commonly used in goal-directed motor tasks. Memory delays in this range provide ample time for spatial working memory to decay in the quality of metrics that are stored ( Rolheiser et al. 2006) and also to exhibit a memory-related cortical recruitment ( Brignani et al. 2010). In other words, our study provided visual cueing in the time window immediately prior to saccade onset that was similar to what participants would have seen in a typical single memoryguided saccading task. ...
... Using this framework, one can easily adapt the model to the results of our current study. One supporting example suggests that it is important to consider the role of the IPL because this area has been ascribed functions related to saccade planning and maintenance of memorized spatial coordinates ( Brignani et al. 2010;Schluppeck et al. 2005). The planning-control model also accounts for a source of collateral projections to the motor system from the IPL and SPL. ...
Recent investigations have revealed the kinematics of horizontal saccades are less variable near the end of the trajectory than during the course of execution. Converging evidence indicates that oculomotor networks use online sensorimotor feedback to correct for initial trajectory errors. It is also known that oculomotor networks express saccadic corrections with decreased efficiency when responses are made toward memorized locations. The present research investigated whether repetitive motor timekeeping influences online feedback-based corrections in predictive saccades. Predictive saccades are a subclass of memory-guided saccades and are observed when one makes series of timed saccades. We hypothesized that cueing predictive saccades in a sequence would facilitate the expression of trajectory corrections. Seven participants produced a number of single unpaced, visually guided saccades, and also sequences of timed predictive saccades. Kinematic and trajectory variability were used to measure the expression of online saccadic corrections at a number of time indices in saccade trajectories. In particular, we estimated the minimum time required to implement feedback-based corrections, which was consistently 37 ms. Our observations demonstrate that motor commands in predictive memory-guided saccades can be parameterized by spatial working memory and retain the accuracy of online trajectory corrections typically associated with visually guided behavior. In contrast, untimed memory-guided saccades exhibited diminished kinematic evidence for online corrections. We conclude that motor timekeeping and sequencing contributed to efficient saccadic corrections. These results contribute to an evolving view of the interactions between motor planning and spatial working memory, as they relate to oculomotor control.
... We designed a behavioral task to maximize swap error rates and measured participants' brain activity with fMRI while they completed ≥ 200 trials across two scanning sessions. To maximize detection of mnemonic neural activity, the task involved maintenance of specific spatial locations of colored circles, which is known to recruit early visual cortex (Blacker & Courtney, 2016;Peters, Kaiser, Rahm, & Bledowski, 2015;Pratte & Tong, 2014;Sprague, Ester, & Serences, 2014;Brignani, Bortoletto, Miniussi, & Maioli, 2010;Munneke, Heslenfeld, & Theeuwes, 2010;Awh, Anllo-Vento, & Hillyard, 2000). Moreover, we ensured sufficient numbers of swap errors for fMRI analyses by using color as a selection cue and requiring participants to report the location of the cued item (Rajsic & Wilson, 2014), and by prescreening participants to select individuals with high swap error rates (similar to Cai et al., 2020). ...
Working memory is an essential component of cognition that facilitates goal-directed behavior. Famously, it is severely limited and performance suffers when memory load exceeds an individual's capacity. Modeling of visual working memory responses has identified two likely types of errors: guesses and swaps. Swap errors may arise from a misbinding between the features of different items. Alternatively, these errors could arise from memory noise in the feature dimension used for cueing a to-be-tested memory item, resulting in the wrong item being selected. Finally, it is possible that so-called "swap errors" actually reflect informed guessing, which could occur at the time of a cue, or alternatively, at the time of the response. Here, we combined behavioral response modeling and fMRI pattern analysis to test the hypothesis that swap errors involve the active maintenance of an incorrect memory item. After the encoding of six spatial locations, a retro-cue indicated which location would be tested after memory retention. On accurate trials, we could reconstruct a memory representation of the cued location in both early visual cortex and intraparietal sulcus. On swap error trials identified with mixture modeling, we were able to reconstruct a representation of the swapped location, but not of the cued location, suggesting the maintenance of the incorrect memory item before response. Moreover, participants subjectively responded with some level of confidence, rather than complete guessing, on a majority of swap error trials. Together, these results suggest that swap errors are not mere response-phase guesses, but instead result from failures of selection in working memory, contextual binding errors, or informed guesses, which produce active maintenance of incorrect memory representations.
... Studies using transcranial magnetic stimulation (TMS) over the dorsolateral prefrontal cortex (DLPFC) (Hamidi et al. 2009; Mottaghy et al. 2002;Oliveri et al. 2001;Postle et al. 2006) further confirm the role of the prefrontal cortex in working memory. A recent study using EEG examined the delay phase during a memory-guided saccade task and localized prefrontal activity only during the initial part of the delay period (Brignani et al. 2010). The extent and potential timing of prefrontal activity during the immediate transition from a visually guided motor task to a memory-guided motor task are still not well established. ...
It is well established that the prefrontal cortex is involved during memory-guided tasks whereas visually guided tasks are controlled in part by a frontal-parietal network. However, the nature of the transition from visually guided to memory-guided force control is not as well established. As such, this study examines the spatiotemporal pattern of brain activity that occurs during the transition from visually guided to memory-guided force control. We measured 128-channel scalp electroencephalography (EEG) in healthy individuals while they performed a grip force task. After visual feedback was removed, the first significant change in event-related activity occurred in the left central region by 300 ms, followed by changes in prefrontal cortex by 400 ms. Low-resolution electromagnetic tomography (LORETA) was used to localize the strongest activity to the left ventral premotor cortex and ventral prefrontal cortex. A second experiment altered visual feedback gain but did not require memory. In contrast to memory-guided force control, altering visual feedback gain did not lead to early changes in the left central and midline prefrontal regions. Decreasing the spatial amplitude of visual feedback did lead to changes in the midline central region by 300 ms, followed by changes in occipital activity by 400 ms. The findings show that subjects rely on sensorimotor memory processes involving left ventral premotor cortex and ventral prefrontal cortex after the immediate transition from visually guided to memory-guided force control.
... Here, we combined tractography with functional MRI (fMRI) to investigate structural connections between nodes in the cortical network for gaze control in humans. This system is of fundamental importance as oculomotor responses can be used to probe many aspects of cognitive control Sweeney et al. 2007) including spatial working memory (Husain et al. 2001;Brignani et al. 2010), visual search and attention (Binello et al. 1995;Mannan et al. 2005) as well as topdown control related to response initiation and suppression (Sumner et al. 2006;Anderson et al. 2008). It is now well established in nonhuman primates that a distributed network, including the frontal eye field (FEF) in the dorsolateral frontal cortex (Bruce et al. 1985;Huerta et al. 1987), supplementary eye field (SEF) located in dorsomedial frontal cortex (Russo and Bruce 2000), and parietal eye field (PEF) within the lateral intraparietal (LIP) area (Thier and Andersen 1998;Goldberg et al. 2002), is crucial for saccadic eye movement control (for a review, see Johnston and Everling 2008). ...
Functional magnetic resonance imaging (fMRI) techniques allow definition of cortical nodes that are presumed to be components of large-scale distributed brain networks involved in cognitive processes. However, very few investigations examine whether such functionally defined areas are in fact structurally connected. Here, we used combined fMRI and diffusion MRI-based tractography to define the cortical network involved in saccadic eye movement control in humans. The results of this multimodal imaging approach demonstrate white matter pathways connecting the frontal eye fields and supplementary eye fields, consistent with the known connectivity of these regions in macaque monkeys. Importantly, however, these connections appeared to be more prominent in the right hemisphere of humans. In addition, there was evidence of a dorsal frontoparietal pathway connecting the frontal eye field and the inferior parietal lobe, also right hemisphere dominant, consistent with specialization of the right hemisphere for directed attention in humans. These findings demonstrate the utility and potential of using multimodal imaging techniques to define large-scale distributed brain networks, including those that demonstrate known hemispheric asymmetries in humans.
Background:
Among the elderly, dementia is a common and disabling disorder with primary manifestations of cognitive impairments. Diagnosis and intervention in its early stages is the key to effective treatment. Nowadays, the test of cognitive function relies mainly on neuropsychological tests, such as the Mini-Mental State Examination (MMSE) and Montreal Cognitive Assessment (MoCA). However, they have noticeable shortcomings, e.g., the biases of subjective judgments from physicians and the cost of the labor of these well-trained physicians. Thus, advanced and objective methods are urgently needed to evaluate cognitive functions.
Methods:
We developed a cognitive assessment system through measuring the saccadic eye movements in three tasks. The cognitive functions were evaluated by both our system and the neuropsychological tests in 310 subjects, and the evaluating results were directly compared.
Results:
In general, most saccadic parameters correlate well with the MMSE and MoCA scores. Moreover, some subjects with high MMSE and MoCA scores have high error rates in performing these three saccadic tasks due to various errors. The primary error types vary among tasks, indicating that different tasks assess certain specific brain functions preferentially. Thus, to improve the accuracy of evaluation through saccadic tasks, we built a weighted model to combine the saccadic parameters of the three saccadic tasks, and our model showed a good diagnosis performance in detecting patients with cognitive impairment.
Conclusion:
The comprehensive analysis of saccadic parameters in multiple tasks could be a reliable, objective, and sensitive method to evaluate cognitive function and thus to help diagnose cognitive impairments.
Background and Objectives
Among the elderly, dementia is a common and disabling disorder with primary manifestations of cognitive impairments. Diagnosis and intervention in its early stages is the key to effective treatment. Practically, the test of cognitive function relies mainly on neuropsychological tests, such as the Mini-Mental State Examination (MMSE) and Montreal Cognitive Assessment (MoCA). Although these tests are widely used at the present, there are noticeable shortcomings, e.g., the biases of subjective judgments from physicians and the cost of the labor of these well-trained physicians. Thus, advanced and objective methods are urgently needed to evaluate cognitive functions. Accumulative evidence indicates that the saccades in certain tasks are highly correlated with the performance in some cognitive functions. However, only a few studies directly compared saccades with the performance in neuropsychological tests depicted by their scores. Thus, the reliability of using saccades as a behavioral biomarker to evaluate cognitive functions has rarely been explored. Methods310 subjects performed three sequential designed oculomotor tasks, pro-saccade (PS), anti-saccade (AS) and memory-guided saccade (MGS) and the saccadic parameters including error rate, saccadic reaction time and spatial error are studied.ResultsIn general, most saccadic parameters correlate well with the MMSE and MoCA scores. Moreover, some subjects with high MMSE and MoCA scores have very high error rates in performing these three tasks due to various errors in saccade control. The primary error types vary among tasks, indicating that different tasks assess certain specific brain functions preferentially. Thus, to improve the accuracy of evaluation through saccadic tasks, we built a weighted model to combine the saccadic parameters of the three saccadic tasks. The receiver operating characteristic (ROC) curve analysis shows that the discrimination between cognitive impairment patients and control subjects is better through the output of our model than the MMSE test. Conclusion
Measuring saccades in multiple tasks could be a reliable, objective and sensitive method to evaluate cognitive function and thus to help diagnosing cognitive impairments.
A remarkable capability of the brain's action representation system is that neurons with 'mirroring' properties respond both when an agent (human or monkey) executes an action and also when the agent observes a similar action performed by someone else. Curiously, however, observed actions involve numerous viewer perspectives, whereas execution of actions occur with respect to 'self' coordinate space, and the mapping between viewer perspectives is not a known property of the so-called mirror neuron system (MNS). Toward a resolution of this paradox, we demonstrate in humans striking new evidence that key areas defined as part of the MNS of the frontal lobe, directly encode viewer perspective during action observation (Exps 1, 2). What is more, this property applies only to observation of embodied actions and not when disembodied movements of objects are observed (Exp 3). These findings raise the intriguing possibility that the MNS plays a critical role in the formation of action memories.
Functional MRI was used to examine cerebral activations in 12 subjects while they performed a spatial attention task. This study applied more stringent behavioural and cognitive controls than previously used for similar experiments: (i) subjects were included only if they showed evidence of attentional shifts while performing the task in the magnet; (ii) the experimental task and baseline condition were designed to eliminate the contributions of motor output, visual fixation, inhibition of eye movements, working memory and the conditional (no-go) component of responding. Activations were seen in all three hypothesized cortical epicentres forming a network for spatial attention: the lateral premotor cortex (frontal eye fields), the posterior parietal cortex and the cingulate cortex. Subcortical activations were seen in the basal ganglia and the thalamus. Although the task required attention to be equally shifted to the left and to the right, eight of 10 subjects showed a greater area of activation in the right parietal cortex, consistent with the specialization of the right hemisphere for spatial attention. Other areas of significant activation included the posterior temporo-occipital cortex and the anterior insula. The temporo-occipital activation was within a region broadly defined as MT+ (where MT is the middle temporal area) which contains the human equivalent of area MT in the macaque monkey. This temporo-occipital area appears to constitute a major component of the functional network activated by this spatial attention task. Its activation may reflect the 'inferred' shift of the attentional focus across the visual scene.
The location and possible function of the human frontal eye-field (FEF) were evaluated by reviewing results of cerebral blood-flow (CBF) and lesion studies. A remarkable consistency was found regarding the rostro-caudal (Y: from -6 to 1 mm) and dorso-ventral (Z: from 44 to 51 mm) location of the FEF, as defined by the CBF method within a standardized stereotaxic system (the zero point for all X, Y and Z coordinates coinciding with the anterior commissure, Talairach and Tournoux [Co-planar Stereotactic Atlas of the Human Brain, Georg Thieme, Stuttgart, 1988]. In contrast, there was a marked variability along the mediolateral axis (X: from -24 to -40 mm for the left hemisphere and from 21 to 40 mm for the right hemisphere). The human FEF is thus located either in the vicinity of the precentral sulcus and/or in the depth of the caudalmost part of the superior frontal sulcus. In either case, this location challenges the commonly held view of the FEF being located in Broadmann's area 8. With regard to FEF function, the results of CBF studies failed to support a role for the FEF in the cognitive aspects of oculomotor control, such as the execution of anti-saccades. Blood-flow activation data are consistent in this respect with the results of lesion studies. It is proposed that future research on FEF function in human subjects may benefit from focusing on the visuomotor rather than the cognitive aspects of oculomotor control.
A variety of biological evidence has identified a frontal-parietal circuit underlying spatial working memory for visual stimuli. But the question remains, how do these neural regions accomplish the goal of maintaining location information on-line? We tested the hypothesis that the active rehearsal of spatial information in working memory is accomplished by means of focal shifts of spatial selective attention to memorized locations. Spatial selective attention has been shown to cause changes in the early visual processing of stimuli that appear in attended locations. Thus, the hypothesis of attention-based rehearsal predicts similar modulations of visual processing at memorized locations. We used functional magnetic resonance imaging to observe posterior visual activations during the performance of a spatial working memory task. In line with the hypothesis, spatial rehearsal led to enhanced activation in the early visual areas contralateral to the memorized locations.
Voluntary attention is often allocated according to internally maintained goals. Recent evidence indicates that the frontal eye field (FEF) participates in the deployment of spatial attention, even in the absence of saccadic eye movements. In addition, many FEF neurons maintain persistent representations of impending saccades. However, the role of persistent activity in the general maintenance of spatial information, and its relationship to spatial attention, has not been explored. We recorded the responses of single FEF neurons in monkeys trained to remember cued locations in order to detect changes in targets embedded among distracters in a task that did not involve saccades. We found that FEF neurons persistently encoded the cued location throughout the trial during the delay period, when no visual stimuli were present, and during visual discrimination. Furthermore, FEF activity reliably predicted whether monkeys would detect the target change. Population analyses revealed that FEF neurons with persistent activity were more effective at selecting the target from among distracters than neurons lacking persistent activity. These results demonstrate that FEF neurons maintain spatial information in the absence of saccade preparation and suggest that this maintenance contributes to the selection of relevant visual stimuli.
We studied the activity of single neurons in the frontal eye fields of awake macaque monkeys trained to perform several oculomotor tasks. Fifty-four percent of neurons discharged before visually guided saccades. Three different types of presaccadic activity were observed: visual, movement, and anticipatory. Visual activity occurred in response to visual stimuli whether or not the monkey made saccades. Movement activity preceded purposive saccades, even those made without visual targets. Anticipatory activity preceded even the cue to make a saccade if the monkey could reliably predict what saccade he had to make. These three different activities were found in different presaccadic cells in different proportions. Forty percent of presaccadic cells had visual activity (visual cells) but no movement activity. For about half of the visual cells the response was enhanced if the monkey made saccades to the receptive-field stimulus, but there was no discharge before similar saccades made without visual targets. Twenty percent of presaccadic neurons discharged as briskly before purposive saccades made without a visual target as they did before visually guided saccades, and had weak or absent visual responses. These cells were defined as movement cells. Movement cells discharged much less or not at all before saccades made spontaneously without a task requirement or an overt visual target. The remaining presaccadic neurons (40%) had both visual and movement activity (visuomovement cells). They discharged most briskly before visually guided eye movements, but also discharged before purposive eye movements made in darkness and responded to visual stimuli in the absence of saccades. There was a continuum of visuomovement cells, from cells in which visual activity predominated to cells in which movement activity predominated. This continuum suggests that although visual cells are quite distinct from movement cells, the division of cell types into three classes may be only a heuristic means of describing the processing flow from visual input to eye-movement output. Twenty percent of visuomovement and movement cells, but fewer than 2% of visual cells, had anticipatory activity. Only one cell had anticipatory activity as its sole response. When the saccade was delayed relative to the target onset, visual cells responded to the target appearance, movement cells discharged before the saccade, and visuomovement cells discharged in different ways during the delay, usually with some discharge following the target and an increase in rate immediately before the saccade. Presaccadic neurons of all types were actively suppressed following a saccade into their response fields.(ABSTRACT TRUNCATED AT 400 WORDS)
We studied single neurons in the frontal eye fields of awake macaque monkeys and compared their activity with the saccadic eye movements elicited by microstimulation at the sites of these neurons. Saccades could be elicited from electrical stimulation in the cortical gray matter of the frontal eye fields with currents as small as 10 microA. Low thresholds for eliciting saccades were found at the sites of cells with presaccadic activity. Presaccadic neurons classified as visuomovement or movement were most associated with low (less than 50 microA) thresholds. High thresholds (greater than 100 microA) or no elicited saccades were associated with other classes of frontal eye field neurons, including neurons responding only after saccades and presaccadic neurons, classified as purely visual. Throughout the frontal eye fields, the optimal saccade for eliciting presaccadic neural activity at a given recording site predicted both the direction and amplitude of the saccades that were evoked by microstimulation at that site. In contrast, the movement fields of postsaccadic cells were usually different from the saccades evoked by stimulation at the sites of such cells. We defined the low-threshold frontal eye fields as cortex yielding saccades with stimulation currents less than or equal to 50 microA. It lies along the posterior portion of the arcuate sulcus and is largely contained in the anterior bank of that sulcus. It is smaller than Brodmann's area 8 but corresponds with the union of Walker's cytoarchitectonic areas 8A and 45. Saccade amplitude was topographically organized across the frontal eye fields. Amplitudes of elicited saccades ranged from less than 1 degree to greater than 30 degrees. Smaller saccades were evoked from the ventrolateral portion, and larger saccades were evoked from the dorsomedial portion. Within the arcuate sulcus, evoked saccades were usually larger near the lip and smaller near the fundus. Saccade direction had no global organization across the frontal eye fields; however, saccade direction changed in systematic progressions with small advances of the microelectrode, and all contralateral saccadic directions were often represented in a single electrode penetration down the bank of the arcuate sulcus. Furthermore, the direction of change in these progressions periodically reversed, allowing particular saccade directions to be multiply represented in nearby regions of cortex.(ABSTRACT TRUNCATED AT 400 WORDS)
Previous work in nonhuman primates and in patients with frontal lobe damage has suggested that the frontal cortex plays a
critical rolein the performance of both spatial and nonspatial working memory tasks. The present study used positron emission
tomography with magnetic resonance imaging to demonstrate the existence, within the human brain, of two functionally distinct
subdivisions of the lateral frontal cortex, which may subserve different aspects of spatial working memory. Five spatial memory
tasks were used, which varied in terms of the extent to which they required different executive processes. When the task required
the organization and execution of a sequence of spatial moves retained in working memory, significant changes in blood flow
were observed in ventrolateral frontal cortex (area 41) bilaterally. By contrast, when the task required active monitoring
and manipulation of spatial information within working memory, additional activation foci were observed in mid-dorsolateral
frontal cortex (areas 46 and 9). These findings support a two-stage model of spatial working memory processing within the
lateral frontal cortex.
We have taken advantage of the temporal resolution afforded by functional magnetic resonance imaging (fMRI) to investigate the role played by medial wall areas in humans during working memory tasks. We demarcated the medial motor areas activated during simple manual movement, namely the supplementary motor area (SMA) and the cingulate motor area (CMA), and those activated during visually guided saccadic eye movements, namely the supplementary eye field (SEF). We determined the location of sustained activity over working memory delays in the medial wall in relation to these functional landmarks during both spatial and face working memory tasks. We identified two distinct areas, namely the pre-SMA and the caudal part of the anterior cingulate cortex (caudal-AC), that showed similar sustained activity during both spatial and face working memory delays. These areas were distinct from and anterior to the SMA, CMA, and SEF. Both the pre-SMA and caudal-AC activation were identified by a contrast between sustained activity during working memory delays as compared with sustained activity during control delays in which subjects were waiting for a cue to make a simple manual motor response. Thus, the present findings suggest that sustained activity during working memory delays in both the pre-SMA and caudal-AC does not reflect simple motor preparation but rather a state of preparedness for selecting a motor response based on the information held on-line.
Functional MRI was used to examine cerebral activations in 12 subjects while they performed a spatial attention task. This study applied more stringent behavioural and cognitive controls than previously used for similar experiments: (i) subjects were included only if they showed evidence of attentional shifts while performing the task in the magnet; (ii) the experimental task and baseline condition were designed to eliminate the contributions of motor output, visual fixation, inhibition of eye movements, working memory and the conditional (no-go) component of responding. Activations were seen in all three hypothesized cortical epicentres forming a network for spatial attention: the lateral premotor cortex (frontal eye fields), the posterior parietal cortex and the cingulate cortex. Subcortical activations were seen in the basal ganglia and the thalamus. Although the task required attention to be equally shifted to the left and to the right, eight of 10 subjects showed a greater area of activation in the right parietal cortex, consistent with the specialization of the right hemisphere for spatial attention. Other areas of significant activation included the posterior temporo-occipital cortex and the anterior insula. The temporo-occipital activation was within a region broadly defined as MT+ (where MT is the middle temporal area) which contains the human equivalent of area MT in the macaque monkey. This temporo-occipital area appears to constitute a major component of the functional network activated by this spatial attention task. Its activation may reflect the 'inferred' shift of the attentional focus across the visual scene.
The dorsolateral prefrontal cortex (DPFC) and the posterior parietal cortex (PPC) are anatomically and functionally interconnected, and have been implicated in working memory and the preparation for behavioral action. To substantiate those functions at the neuronal level, we designed a visuomotor task that dissociated the perceptual and executive aspects of the perception-action cycle in both space and time. In that task, the trial-initiating cue (a color) indicated with different degrees of certainty the direction of the correct manual response 12 s later. We recorded extracellular activity from 258 prefrontal and 223 parietal units in two monkeys performing the task. In the DPFC, some units (memory cells) were attuned to the color of the cue, independent of the response-direction it connoted. Their discharge tended to diminish in the course of the delay between cue and response. In contrast, few color-related units were found in PPC, and these did not show decreasing patterns of delay activity. Other units in both cortices (set cells) were attuned to response-direction and tended to accelerate their firing in anticipation of the response and in proportion to the predictability of its direction. A third group of units was related to the determinacy of the act; their firing was attuned to the certainty with which the animal could predict the correct response, whatever its direction. Cells of the three types were found closely intermingled histologically. These findings further support and define the role of DPFC in executive functions and in the temporal closure of the perception-action cycle. The findings also agree with the involvement of PPC in spatial aspects of visuomotor behavior, and add a temporal integrative dimension to that involvement. Together, the results provide physiological evidence for the role of a prefrontal-parietal network in the integration of perception with action across time.
It is controversial whether the dorsolateral prefrontal cortex is involved in the maintenance of items in working memory or in the selection of responses. We used event-related functional magnetic resonance imaging to study the performance of a spatial working memory task by humans. We distinguished the maintenance of spatial items from the selection of an item from memory to guide a response. Selection, but not maintenance, was associated with activation of prefrontal area 46 of the dorsal lateral prefrontal cortex. In contrast, maintenance was associated with activation of prefrontal area 8 and the intraparietal cortex. The results support a role for the dorsal prefrontal cortex in the selection of representations. This accounts for the fact that this area is activated both when subjects select between items on working memory tasks and when they freely select between movements on tasks of willed action.
We investigated the hypothesis that the covert focusing of spatial attention mediates the on-line maintenance of location information in spatial working memory. During the delay period of a spatial working-memory task, behaviorally irrelevant probe stimuli were flashed at both memorized and nonmemorized locations. Multichannel recordings of event-related potentials (ERPs) were used to assess visual processing of the probes at the different locations. Consistent with the hypothesis of attention-based rehearsal, early ERP components were enlarged in response to probes that appeared at memorized locations. These visual modulations were similar in latency and topography to those observed after explicit manipulations of spatial selective attention in a parallel experimental condition that employed an identical stimulus display.
Several lines of evidence suggest that planning eye movements and directing visuospatial attention share overlapping brain mechanisms. This study tested whether spatial attention can be enhanced by altering oculomotor signals within the brain. Monkeys performed a spatial attention task while neurons within the frontal eye field, an oculomotor area within prefrontal cortex, were electrically stimulated below the level at which eye movements are evoked. We found that we could improve the monkey's performance with microstimulation when, but only when, the object to be attended was positioned in the space represented by the cortical stimulation site.
The dorsolateral prefrontal cortex (DLPFC) plays a key role in working memory (WM). Yet its precise contribution (the storage, manipulation and/or utilization of information for the forthcoming response) remains to be determined. To test the hypothesis that the DLPFC is more involved in the preparation of actions than in the maintenance of information in short-term memory (STM), we undertook a functional magnetic resonance imaging investigation in normal subjects performing two delayed response tasks (matching and reproduction tasks) in a visuospatial task sequence (presentation, delay, response). In the two tasks, the presentation and delay phases were similar, but the expected response was different: in the matching task, subjects had to indicate whether a visuospatial sequence matched the sequence presented before the delay period; in the reproduction task, subjects had to reproduce the sequence and, therefore, to mentally organize their response during the delay. Using a fMRI paradigm focusing on the delay period, we observed a significant DLPFC activation when subjects were required to mentally prepare a sequential action based on the information stored in STM. When subjects had only to maintain a visuospatial stimulus in STM, no DLPFC activation was found. These results suggest that a parietal-premotor network is sufficient to store visuospatial information in STM whereas the DLPFC is involved when it is necessary to mentally prepare a forthcoming sequential action based on the information stored in STM.
The internal organization of a higher level visual area in the human parietal cortex was mapped. Functional magnetic resonance
images were acquired while the polar angle of a peripheral target for a delayed saccade was gradually changed. A region in
the superior parietal cortex showed robust retinotopic mapping of the remembered target angle. The map reversed when the direction
of rotation of the remembered targets was reversed and persisted unchanged when study participants detected rare target reappearances
while maintaining fixation, or when the eccentricity of successive remembered targets was unpredictable. This region may correspond
to the lateral intraparietal area in macaque monkeys.
Because environmental information is often suboptimal, visual perception must frequently rely on the brain's reconstruction of contours absent from retinal images. Illusory contour (IC) stimuli have been used to investigate these "filling-in" processes. Intracranial recordings and neuroimaging studies show IC sensitivity in lower-tier area V2, and to a lesser extent V1. Some interpret these data as evidence for feedforward processing of IC stimuli, beginning at lower-tier visual areas. On the basis of lesion, visual evoked potentials (VEP), and neuroimaging evidence, others contend that IC sensitivity is a later, higher-order process. Whether IC sensitivity seen in lower-tier areas indexes feedforward or feedback processing remains unresolved. In a series of experiments, we addressed the spatiotemporal dynamics of IC processing. Centrally presented IC stimuli resulted in early VEP modulation (88-100 msec) over lateral-occipital (LOC) scalp--the IC effect. The IC effect followed visual response onset by 40 msec. Scalp current density topographic mapping, source analysis, and functional magnetic resonance imaging results all localized the IC effect to bilateral LOC areas. We propose that IC sensitivity described in V2 and V1 may reflect predominantly feedback modulation from higher-tier LOC areas, where IC sensitivity first occurs. Two additional observations further support this proposal. The latency of the IC effect shifted dramatically later (approximately 120 msec) when stimuli were laterally presented, indicating that retinotopic position alters IC processing. Immediately preceding the IC effect, the VEP modulated with inducer eccentricity--the configuration effect. We interpret this to represent contributions from global stimulus parameters to scene analysis. In contrast to the IC effect, the topography of the configuration effect was restricted to central parieto-occipital scalp.
Contrary to our rich phenomenological visual experience, our visual short-term memory system can maintain representations of only three to four objects at any given moment. For over a century, the capacity of visual memory has been shown to vary substantially across individuals, ranging from 1.5 to about 5 objects. Although numerous studies have recently begun to characterize the neural substrates of visual memory processes, a neurophysiological index of storage capacity limitations has not yet been established. Here, we provide electrophysiological evidence for lateralized activity in humans that reflects the encoding and maintenance of items in visual memory. The amplitude of this activity is strongly modulated by the number of objects being held in the memory at the time, but approaches a limit asymptotically for arrays that meet or exceed storage capacity. Indeed, the precise limit is determined by each individual's memory capacity, such that the activity from low-capacity individuals reaches this plateau much sooner than that from high-capacity individuals. Consequently, this measure provides a strong neurophysiological predictor of an individual's capacity, allowing the demonstration of a direct relationship between neural activity and memory capacity.
The most compelling neural evidence for working memory is persistent neuronal activity bridging past sensory cues and their contingent future motor acts. This observation, however, does not answer what is actually being remembered or coded for by this activity. To address this fundamental issue, we imaged the human brain during maintenance of spatial locations and varied whether the memory-guided saccade was selected before or after the delay. An oculomotor delayed matching-to-sample task (match) was used to measure maintained motor intention because the direction of the forthcoming saccade was known throughout the delay. We used a nonmatching-to-sample task (nonmatch) in which the saccade was unpredictable to measure maintained spatial attention. Oculomotor areas were more active during match delays, and posterior parietal cortex and inferior frontal cortex were more active during nonmatch delays. Additionally, the fidelity of the memory was predicted by the delay-period activity of the frontal eye fields; the magnitude of delay-period activity correlated with the accuracy of the memory-guided saccade. Experimentally controlling response selection allowed us to functionally separate nodes of a network of frontal and parietal areas that usually coactivate in studies of working memory. We propose that different nodes in this network maintain different representational codes, motor and spatial. Which code is being represented by sustained neural activity is biased by when in the transformation from perception to action the response can be selected.
Posterior parietal cortex (PPC) is thought to play a critical role in decision making, sensory attention, motor intention, and/or working memory. Research on the PPC in non-human primates has focused on the lateral intraparietal area (LIP) in the intraparietal sulcus (IPS). Neurons in LIP respond after the onset of visual targets, just before saccades to those targets, and during the delay period in between. To study the function of posterior parietal cortex in humans, it will be crucial to have a routine and reliable method for localizing specific parietal areas in individual subjects. Here, we show that human PPC contains at least two topographically organized regions, which are candidates for the human homologue of LIP. We mapped the topographic organization of human PPC for delayed (memory guided) saccades using fMRI. Subjects were instructed to fixate centrally while a peripheral target was briefly presented. After a further 3-s delay, subjects made a saccade to the remembered target location followed by a saccade back to fixation and a 1-s inter-trial interval. Targets appeared at successive locations "around the clock" (same eccentricity, approximately 30 degrees angular steps), to produce a traveling wave of activity in areas that are topographically organized. PPC exhibited topographic organization for delayed saccades. We defined two areas in each hemisphere that contained topographic maps of the contra-lateral visual field. These two areas were immediately rostral to V7 as defined by standard retinotopic mapping. The two areas were separated from each other and from V7 by reversals in visual field orientation. However, we leave open the possibility that these two areas will be further subdivided in future studies. Our results demonstrate that topographic maps tile the cortex continuously from V1 well into PPC.
We used functional magnetic resonance imaging (fMRI) to investigate the role of the human posterior parietal cortex (PPC) in storing target locations for delayed double-step saccades. To do so, we exploited the laterality of a subregion of PPC that preferentially responds to the memory of a target location presented in the contralateral visual field. Using an event-related design, we tracked fMRI signal changes in this region while subjects remembered the locations of two sequentially flashed targets, presented in either the same or different visual hemifields, and then saccaded to them in sequence. After presentation of the first target, the fMRI signal was always related to the side of the visual field in which it had been presented. When the second target was added, the cortical activity depended on the respective locations of both targets but was still significantly selective for the target of the first saccade. We conclude that this region within the human posterior parietal cortex not only acts as spatial storage center by retaining target locations for subsequent saccades but is also involved in selecting the target for the first intended saccade.
An automated coordinate-based system to retrieve brain labels from the 1988 Talairach Atlas, called the Talairach Daemon (TD), was previously introduced [Lancaster et al., 1997]. In the present study, the TD system and its 3-D database of labels for the 1988 Talairach atlas were tested for labeling of functional activation foci. TD system labels were compared with author-designated labels of activation coordinates from over 250 published functional brain-mapping studies and with manual atlas-derived labels from an expert group using a subset of these activation coordinates. Automated labeling by the TD system compared well with authors' labels, with a 70% or greater label match averaged over all locations. Author-label matching improved to greater than 90% within a search range of +/-5 mm for most sites. An adaptive grey matter (GM) range-search utility was evaluated using individual activations from the M1 mouth region (30 subjects, 52 sites). It provided an 87% label match to Brodmann area labels (BA 4 & BA 6) within a search range of +/-5 mm. Using the adaptive GM range search, the TD system's overall match with authors' labels (90%) was better than that of the expert group (80%). When used in concert with authors' deeper knowledge of an experiment, the TD system provides consistent and comprehensive labels for brain activation foci. Additional suggested applications of the TD system include interactive labeling, anatomical grouping of activation foci, lesion-deficit analysis, and neuroanatomy education. (C) 2000 Wiley-Liss, Inc.
It is controversial whether the dorsolateral prefrontal cortex is involved in the maintenance of items in working memory or
in the selection of responses. We used event-related functional magnetic resonance imaging to study the performance of a spatial
working memory task by humans. We distinguished the maintenance of spatial items from the selection of an item from memory
to guide a response. Selection, but not maintenance, was associated with activation of prefrontal area 46 of the dorsal lateral
prefrontal cortex. In contrast, maintenance was associated with activation of prefrontal area 8 and the intraparietal cortex.
The results support a role for the dorsal prefrontal cortex in the selection of representations. This accounts for the fact
that this area is activated both when subjects select between items on working memory tasks and when they freely select between
movements on tasks of willed action.
The location and possible function of the human frontal eye-field (FEF) were evaluated by reviewing results of cerebral blood-flow (CBF) and lesion studies. A remarkable consistency was found regarding the rostro-caudal (Y: from −6 to 1 mm) and dorso-ventral (Z: from 44 to 51 mm) location of the FEF, as defined by the CBF method within a standardized stereotaxic system (the zero point for all X, Y and Z coordinates coinciding with the anterior commissure, Talairach and Tournoux [Co-planar Stereotactic Atlas of the Human Brain, Georg Thieme, Stuttgart, 1988]. In contrast, there was a marked variability along the mediolateral axis (X: from −24 to −40 mm for the left hemisphere and from 21 to 40 mm for the right hemisphere). The human FEF is thus located either in the vicinity of the precentral sulcus and/or in the depth of the caudalmost part of the superior frontal sulcus. In either case, this location challenges the commonly held view of the FEF being located in Brodmann's area 8. With regard to FEF function, the results of CBF studies failed to support a role for the FEF in the cognitive aspects of oculomotor control, such as the execution of anti-saccades. Blood-flow activation data are consistent in this respect with the results of lesion studies. It is proposed that future research on FEF function in human subjects may benefit from focusing on the visuomotor rather than the cognitive aspects of oculomotor control.
This note provides a statistical-graphical method for the evaluation of the statistical significance of difference potentials from a group of subjects, and for the comparison of difference potentials between two groups. A table of the lengths of statistically significant intervals for various sampling interval lengths, numbers of subjects, and autocorrelation parameters is presented.
In macaque monkeys, the posterior parietal cortex (PPC) is concerned with the integration of multimodal information for constructing a spatial representation of the external world (in relation to the macaque's body or parts thereof), and planning and executing object-centred movements. The areas within the intraparietal sulcus (IPS), in particular, serve as interfaces between the perceptive and motor systems for controlling arm and eye movements in space. We review here the latest evidence for the existence of the IPS areas AIP (anterior intraparietal area), VIP (ventral intraparietal area), MIP (medial intraparietal area), LIP (lateral intraparietal area) and CIP (caudal intraparietal area) in macaques, and discuss putative human equivalents as assessed with functional magnetic resonance imaging. The data suggest that anterior parts of the IPS comprising areas AIP and VIP are relatively well preserved across species. By contrast, posterior areas such as area LIP and CIP have been found more medially in humans, possibly reflecting differences in the evolution of the dorsal visual stream and the inferior parietal lobule. Despite interspecies differences in the precise functional anatomy of the IPS areas, the functional relevance of this sulcus for visuomotor tasks comprising target selections for arm and eye movements, object manipulation and visuospatial attention is similar in humans and macaques, as is also suggested by studies of neurological deficits (apraxia, neglect, Bálint's syndrome) resulting from lesions to this region.
This note provides a statistical-graphical method for the evaluation of the statistical significance of difference potentials from a group of subjects, and for the comparison of difference potentials between two groups. A table of the lengths of statistically significant intervals for various sampling interval lengths, numbers of subjects, and autocorrelation parameters is presented.
Accuracy of saccades toward a remembered target positions in the dark (memory-guided saccades) was studied in 11 normal subjects. The subjects were instructed to execute saccades with amplitudes of 20, 40, 60, and 80 degrees, centering on the primary position. Saccades were initially performed for 30 s with visual fixation targets. The targets were then switched off. The subjects continued saccades in the dark with the given amplitude. Most memory-guided saccades overshot the target. Saccades with an amplitude error of 12.7 +/- 7.3 degrees (mean +/- SD) were followed by corrective saccades, while no corrective saccades occurred following saccades with an error of 4.3 +/- 4.0 degrees. The accuracy of initial memory-guided saccades decreased with time. However, the amplitude of the memory-guided saccades was corrected when the error was beyond about 5 degrees. These results suggest that memory-guided saccades are not repetitions of visually guided saccades, but nonvisual error signals relate to the control of eye movements in the dark.
The need for a simply applied quantitative assessment of handedness is discussed and some previous forms reviewed. An inventory of 20 items with a set of instructions and response- and computational-conventions is proposed and the results obtained from a young adult population numbering some 1100 individuals are reported. The separate items are examined from the point of view of sex, cultural and socio-economic factors which might appertain to them and also of their inter-relationship to each other and to the measure computed from them all. Criteria derived from these considerations are then applied to eliminate 10 of the original 20 items and the results recomputed to provide frequency-distribution and cumulative frequency functions and a revised item-analysis. The difference of incidence of handedness between the sexes is discussed.
The three types of responses described in this report were all related to either saccades to remembered targets or to the targets to be remembered. All the responses were a decrease in discharge rate. The new paradigm frequently used in this study required the monkey to remember the location of a stimulus presented briefly while it was fixating; a later saccade was rewarded if it was made to the position of the no longer present stimulus. The three types of responses were revealed by the use of this paradigm; they were less obvious or undetectable in conventional paradigms in which the monkey responded to the stimulus that was still present. The first type of response was to the visual stimulus that the monkey had to use as the target for a subsequent saccade (memory-contingent visual response); a minimal response occurred if the monkey made a saccade to the stimulus while it was still present or if the monkey continued to fixate. Latencies and receptive fields for this response were similar to those for simple visual responses (17). Of 93 substantia nigra cells with some sensory-oculomotor response, 29 cells (31%) showed this type of response. The second type of response was temporally correlated with a saccade made to the point where a visual stimulus was once present (memory-contingent saccade response). Nearly half of these cells showed no significant response if the saccade was made to the stimulus while it was still present, whereas others showed a comparable response in both conditions. None of them showed a change in activity in relation to spontaneous saccades in darkness. The onset of a memory-contingent saccade response usually preceded the saccade onset by up to 280 ms (most frequently by 70 to 240 ms). The response was usually spatially selective; for most responses, contralateral saccades were associated with an exclusive or greater response compared with ipsilateral saccades. Movement fields were demonstrated for some cells. Of 128 cells tested, 41 cells (32%) showed this second type of response. The third type of response began after the briefly presented stimulus and continued until the saccade made to the stimulus position (memory-contingent sustained response). Of 95 cells tested, 15 cells (16%) showed this type of response. These cells frequently also showed a memory-contingent saccade response. These three types of substantia nigra cell activity are related to the special type of visuooculomotor behavior in which a visual input, particularly its spatial location, must be stored and then used as a target for a saccadic eye movement. One of their efferent connections, the nigrocollicular pathway, may act as a channel for the stored visual spatial information to be executed as a saccadic eye movement. Discussions of basal ganglia function generally emphasize one of three functions: sensory, motor, or cognitive. All of these three functional aspects appear to be combined in the substantia nigra pars reticulata, wich is presumably a final stage of processing in the basal ganglia. In single substantia nigra cells, however, they are combined or gated in different ways so that the sensory or motor activities of the cells are specialized for the different contexts in which behavior occurs.
Recent brain-imaging and neurophysiological data indicate that attention is neither a property of a single brain area, nor of the entire brain. While attentional effects seem mediated by a relative amplification of blood flow and electrical activity in the cortical areas processing the attended computation, the details of how this is done through enhancement of attended or suppression of unattended items, or both, appear to depend on the task and brain-area studied. The origins of these amplification effects are to be found in specialized cortical areas of the frontal and parietal lobes that have been described as the anterior and posterior attention systems. These results represent substantial progress in the effort to determine how brain activity is regulated through attention. While many philosophical and practical issues remain in developing an understanding of attentional regulation, the new tools available should provide the basis for progress.
Electroencephalographic (EEG) deflections in humans related to the performance of memory-guided saccades were studied in this work. The EEG deflections were recorded during 2 spatial oculomotor delayed response tasks in which the subject was instructed to make a saccade either to the right or to the left depending on the spatial location of the cue which had been shown in the beginning of the delay period. The EEG deflections were compared to those recorded during a control task in which the subject also made a saccade to the right or to the left after a delay but the requirement to keep spatial information actively in mind was minimized. A slow delay-related shift was recorded during all task conditions. The slow shift was positive in the most frontal and negative in the more posterior recording sites. The negative slow shift in the more posterior recording sites was larger in the memory tasks than in the control task. Since the memory and the control tasks differed mainly in their requirement to hold spatial information in mind it is suggested that the difference in the magnitude of slow shifts between the memory and the control tasks reflects neural activity related to spatial working memory. But although the oculomotor responses in all tasks were similar, the preparatory activities for the impending eye movements may not have been similar and in addition to working memory may have contributed to the observed differences in the slow shifts.
Single-unit recording studies of posterior parietal neurons have indicated a similarity of neuronal activation to that observed in the dorsolateral prefrontal cortex in relation to performance of delayed saccade tasks. A key issue addressed in the present study is whether the different classes of neuronal activity observed in these tasks are encountered more frequently in one or the other area or otherwise exhibit region-specific properties. The present study is the first to directly compare these patterns of neuronal activity by alternately recording from parietal area 7ip and prefrontal area 8a, under the identical behavioral conditions, within the same hemisphere of two monkeys performing an oculomotor delayed response task. The firing rate of 222 posterior parietal and 235 prefrontal neurons significantly changed during the cue, delay, and/or saccade periods of the task. Neuronal responses in the two areas could be distinguished only by subtle differences in their incidence and timing. Thus neurons responding to the cue appeared earliest and were more frequent among the task-related neurons within parietal cortex, whereas neurons exhibiting delay-period activity accounted for a larger proportion of task-related neurons in prefrontal cortex. Otherwise, the task-related neuronal activities were remarkably similar. Cue period activity in prefrontal and parietal cortex exhibited comparable spatial tuning and temporal duration characteristics, taking the form of phasic, tonic, or combined phasic/tonic excitation in both cortical populations. Neurons in both cortical areas exhibited sustained activity during the delay period with nearly identical spatial tuning. The various patterns of delay-period activity-tonic, increasing or decreasing, alone or in combination with greater activation during cue and/or saccade periods-likewise were distributed to both cortical areas. Finally, similarities in the two populations extended to the proportion and spatial tuning of presaccadic and postsaccadic neuronal activity occurring in relation to the memory-guided saccade. The present findings support and extend evidence for a faithful duplication of receptive field properties and virtually every other dimension of task-related activity observed when parietal and prefrontal cortex are recruited to a common task. This striking similarity attests to the principal that information shared by a prefrontal region and a sensory association area with which it is connected is domain specific and not subject to hierarchical elaboration, as is evident at earlier stages of visuospatial processing.
Event-related potentials (ERPs) were recorded as 12 subjects performed a delayed matching to sample task. We presented two bilateral abstract shapes and cued spatially which had to be memorized for a subsequent matching task: left, right or both. During memorization a posterior slow negative ERP wave developed over the hemisphere contralateral to the memorized shape. This effect was similar in high and low memory load trials while the memory figures were visible (for 1000 ms). As the figures disappeared (for 1500 ms), the effect persisted only in the low memory load conditions. We suggest that the contralateral negativity reflects a visual short-term memory process and that capacity limitation in the high memory load condition causes this process to collapse.
A sophisticated study of error-related brain potentials in patients with prefrontal lesions addresses how we monitor performance and adjust cognitive control based on task demands.
An automated coordinate-based system to retrieve brain labels from the 1988 Talairach Atlas, called the Talairach Daemon (TD), was previously introduced [Lancaster et al., 1997]. In the present study, the TD system and its 3-D database of labels for the 1988 Talairach atlas were tested for labeling of functional activation foci. TD system labels were compared with author-designated labels of activation coordinates from over 250 published functional brain-mapping studies and with manual atlas-derived labels from an expert group using a subset of these activation coordinates. Automated labeling by the TD system compared well with authors' labels, with a 70% or greater label match averaged over all locations. Author-label matching improved to greater than 90% within a search range of +/-5 mm for most sites. An adaptive grey matter (GM) range-search utility was evaluated using individual activations from the M1 mouth region (30 subjects, 52 sites). It provided an 87% label match to Brodmann area labels (BA 4 & BA 6) within a search range of +/-5 mm. Using the adaptive GM range search, the TD system's overall match with authors' labels (90%) was better than that of the expert group (80%). When used in concert with authors' deeper knowledge of an experiment, the TD system provides consistent and comprehensive labels for brain activation foci. Additional suggested applications of the TD system include interactive labeling, anatomical grouping of activation foci, lesion-deficit analysis, and neuroanatomy education.
Spatial selective attention and spatial working memory have largely been studied in isolation. Studies of spatial attention have provided clear evidence that observers can bias visual processing towards specific locations, enabling faster and better processing of information at those locations than at unattended locations. We present evidence supporting the view that this process of visual selection is a key component of rehearsal in spatial working memory. Thus, although working memory has sometimes been depicted as a storage system that emerges 'downstream' of early sensory processing, current evidence suggests that spatial rehearsal recruits top-down processes that modulate the earliest stages of visual analysis.
Healthy subjects performed saccadic eye movements in one memory (MEM) and two delay tasks (delay, DEL and modified delay, M-DEL) while we recorded scalp event-related potentials (ERPs) from 25 electrode sites. In the MEM task the subjects were instructed to retain in memory the location of a visual target for a delay of 1-6 s and then perform a remembered saccade at the go signal. In the DEL task the target remained on until movement completion and in the M-DEL task the target, that was visible during the delay period, disappeared synchronously with the go signal. A reduction in response latency and an increase in the percentage of dysmetric movements were observed for the MEM task compared to the two delay tasks. An increased ERP activity at the central-frontal electrode sites compared to the parietal sites was significant only for the MEM task early on during the delay period (500-1000 ms). During the period preceding the onset of the saccade, a parietal increase of activity was observed for all tasks. Furthermore the activity was smaller for the frontal compared to the parietal areas only for the memory task thus indicating a near reversal of the previous pattern of activity observed during the early delay period. This specific activation pattern of frontal and parietal areas, observed for the MEM task only, requires further investigation focusing on the temporal pattern of activation of large brain areas involved in working memory processing.
The brain cannot monitor or react towards the entire world at a given time. Instead, using the process of attention, it selects objects in the world for further analysis. Neuronal activity in the monkey intraparietal area has the properties appropriate for a neuronal substrate of attention: instead of all objects being represented in the parietal cortex, only salient objects are. Such objects can be salient because of their physical properties (recently flashed objects or moving objects) or because they can be made important to the animal by virtue of a task. Although lateral intraparietal area (LIP) neurons respond through the delay period of a memory-guided saccade, they also respond in an enhanced manner to distractors flashed during the delay period of a memory-guided saccade being generated to a position outside the receptive field. This activity parallels the monkey's psychophysical attentional process: attention is ordinarily pinned at the goal of a memory-guided saccade, but it shifts briefly to the locus of a task-irrelevant distractor flashed briefly during the delay period and then returns to the goal. Although neurons in LIP have been implicated as being directly involved in the generation of saccadic eye movements, their activity does not predict where, when, or if a saccade will occur. The ensemble of activity in LIP, however, does accurately describe the locus of attention.
Objectives:
We used a 3-compartment boundary element method (BEM) model from an averaged magnetic resonance image (MRI) data set (Montreal Neurological Institute) in order to provide simple access to realistically shaped volume conductor models for source reconstruction, as compared to individually derived models. The electrode positions were transformed into the model's coordinate system, and the best fit dipole results were transformed back to the original coordinate system. The localization accuracy of the new approach was tested in a comparison with simulated data and with individual BEM models of epileptic spike data from several patients.
Methods:
The standard BEM model consisted of a total of 4770 nodes, which describe the smoothed cortical envelope, the outside of the skull, and the outside of the skin. The electrode positions were transformed to the model coordinate system by using 3-5 fiducials (nasion, left and right preauricular points, vertex, and inion). The transformation consisted of an averaged scaling factor and a rigid transformation (translation and rotation). The potential values at the transformed electrode positions were calculated by linear interpolation from the stored transfer matrix of the outer BEM compartment triangle net. After source reconstruction the best fit dipole results were transformed back into the original coordinate system by applying the inverse of the first transformation matrix.
Results:
Test-dipoles at random locations and with random orientations inside of a highly refined reference BEM model were used to simulate noise-free data. Source reconstruction results using a spherical and the standardized BEM volume conductor model were compared to the known dipole positions. Spherical head models resulted in mislocation errors at the base of the brain. The standardized BEM model was applied to averaged and unaveraged epileptic spike data from 7 patients. Source reconstruction results were compared to those achieved by 3 spherical shell models and individual BEM models derived from the individual MRI data sets. Similar errors to that evident with simulations were noted with spherical head models. Standardized and individualized BEM models were comparable.
Conclusions:
This new approach to head modeling performed significantly better than a simple spherical shell approximation, especially in basal brain areas, including the temporal lobe. By using a standardized head for the BEM setup, it offered an easier and faster access to realistically shaped volume conductor models as compared to deriving specific models from individual 3-dimensional MRI data.
The posterior parietal cortex (PPC), historically believed to be a sensory structure, is now viewed as an area important for sensory-motor integration. Among its functions is the forming of intentions, that is, high-level cognitive plans for movement. There is a map of intentions within the PPC, with different subregions dedicated to the planning of eye movements, reaching movements, and grasping movements. These areas appear to be specialized for the multisensory integration and coordinate transformations required to convert sensory input to motor output. In several subregions of the PPC, these operations are facilitated by the use of a common distributed space representation that is independent of both sensory input and motor output. Attention and learning effects are also evident in the PPC. However, these effects may be general to cortex and operate in the PPC in the context of sensory-motor transformations.
An important cognitive function underlying unified, voluntary behavior is attentional control. Two frontal regions, anterior cingulate cortex (ACC) and dorsolateral prefrontal cortex (DLPFC), appear to be particularly involved in attentional control and monitoring. In this study, we investigated whether ACC is involved in monitoring the preparatory allocation of attention during task switching, or whether ACC is active only when subjects are processing target stimuli and selecting a response, via a cued-attention design. Event-related BOLD fMRI activity was examined using a cue-target paradigm in which subjects performed task switches that selectively required reallocation of attention when tasks changed. There were three cue conditions: informative switch, informative repeat, and neutral. There were four target conditions: informatively cued switch, informatively cued repeat, neutrally cued switch, and neutrally cued repeat. Significant ACC activity was observed following both informative switch and informative repeat cues, but not after neutral cues. No significant ACC activity was observed following any of the target conditions. Significant DLPFC activity was observed following all three cue conditions and following neutrally cued switch targets. Overall, our results suggest that ACC is involved in monitoring the preparatory allocation of attention for conflict at the level of activation of competing attentional sets. The results also support the role of DLPFC in holding cognitive goals in working memory and allocating attention to the appropriate processing systems to meet those goals.
Several decades of psychophysical and neurophysiological studies have established that visual signals are enhanced at the locus of attention. What remains a mystery is the mechanism that initiates biases in the strength of visual representations. Recent evidence argues that, during spatial attention, these biases reflect nascent saccadic eye movement commands. We examined the functional interaction of saccade preparation and visual coding by electrically stimulating sites within the frontal eye fields (FEF) and measuring its effect on the activity of neurons in extrastriate visual cortex. Here we show that visual responses in area V4 could be enhanced after brief stimulation of retinotopically corresponding sites within the FEF using currents below that needed to evoke saccades. The magnitude of the enhancement depended on the effectiveness of receptive field stimuli as well as on the presence of competing stimuli outside the receptive field. Stimulation of non-corresponding FEF representations could suppress V4 responses. The results suggest that the gain of visual signals is modified according to the strength of spatially corresponding eye movement commands.
Scalp electric potentials (electroencephalograms) and extracranial magnetic fields (magnetoencephalograms) are due to the primary (impressed) current density distribution that arises from neuronal postsynaptic processes. A solution to the inverse problem--the computation of images of electric neuronal activity based on extracranial measurements--would provide important information on the time-course and localization of brain function. In general, there is no unique solution to this problem. In particular, an instantaneous, distributed, discrete, linear solution capable of exact localization of point sources is of great interest, since the principles of linearity and superposition would guarantee its trustworthiness as a functional imaging method, given that brain activity occurs in the form of a finite number of distributed hot spots. Despite all previous efforts, linear solutions, at best, produced images with systematic nonzero localization errors. A solution reported here yields images of standardized current density with zero localization error. The purpose of this paper is to present the technical details of the method, allowing researchers to test, check, reproduce and validate the new method.
The dorsolateral prefrontal cortex (DLPFC) plays a crucial role in working memory. Notably, persistent activity in the DLPFC is often observed during the retention interval of delayed response tasks. The code carried by the persistent activity remains unclear, however. We critically evaluate how well recent findings from functional magnetic resonance imaging studies are compatible with current models of the role of the DLFPC in working memory. These new findings suggest that the DLPFC aids in the maintenance of information by directing attention to internal representations of sensory stimuli and motor plans that are stored in more posterior regions.
Over the past two decades significant progress has been made toward understanding the neural basis of primate decision making, the biological process that combines sensory data with stored information to select and execute behavioral responses. The most striking progress in this area has been made in studies of visual-saccadic decision making, a system that is becoming a model for understanding decision making in general. In this system, theoretical models of efficient decision making developed in the social sciences are beginning to be used to describe the computations the brain must perform when it connects sensation and action. Guided in part by these economic models, neurophysiologists have been able to describe neuronal activity recorded from the brains of awake-behaving primates during actual decision making. These recent studies have examined the neural basis of decisions, ranging from those made in predictable sensorimotor tasks to those unpredictable decisions made when animals are engaged in strategic conflict. All of these experiments seem to describe a surprisingly well-integrated set of physiological mechanisms that can account for a broad range of behavioral phenomena. This review presents many of these recent studies within the emerging neuroeconomic framework for understanding primate decision making.
Rehearsal in human spatial working memory is accomplished, in part, via covert shifts of spatial selective attention to memorized locations ("attention-based rehearsal"). We addressed two outstanding questions about attention-based rehearsal: the topography of the attention-based rehearsal effect, and the mechanism by which it operates. Using event-related fMRI and a procedure that randomized the presentation of trials with delay epochs that were either filled with a flickering checkerboard or unfilled, we localized the effect to extrastriate areas 18 and 19, and confirmed its absence in striate cortex. Delay-epoch activity in these extrastriate regions, as well as in superior parietal lobule and intraparietal sulcus, was also lateralized on unfilled trials, suggesting that attention-based rehearsal produces a baseline shift in areas representing the to-be-remembered location in space. No frontal regions (including frontal eye fields) demonstrated lateralized activity consistent with a role in attention-based rehearsal.