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ABSTRACT: Despite a large body of research, extant findings on the functional role of left inferior frontal gyrus (LIFG) in phonological and semantic fluency are still controversial. Based on cross-study comparisons, a recent meta-analysis of neuroimaging results suggests that posterior-dorsal (Brodmann area, BA, 44) and anterior-ventral parts (BA 45) of LIFG contribute differentially to processes of phonologically and semantically cued word retrieval, respectively. In contrast, a subsequent functional magnetic resonance imaging experiment failed to validate the proposed dissociation using a within-subjects design. In particular, no evidence for a specific role of BA 45 in semantic fluency was found. Here, we resolve this apparent controversy by showing that the conflicting findings can be accounted for when considering the influence of task demands and individual ability on resulting functional magnetic resonance imaging activation patterns. By comparing phonological versus semantic fluency, higher activation was robustly observed in BA 44. For the opposite comparison, higher activation was found in dorsal BA 45; however, this was more pronounced in posterior-dorsal parts of BA 45 for low-performing subjects and was only apparent in anterior-dorsal parts of BA 45 under high demands on controlled semantic retrieval. Our results thus disclose important determinants for detecting a functional segregation of LIFG in verbal fluency that also have implications for the controversial findings in previous lesion studies. Moreover, the present parcellation of dorsal BA 45 corresponds well with anatomical evidence suggesting a subdivision into an anterior (45A) and posterior part (45B) and may therefore represent evidence for its functional significance in humans.
Journal of Neuroscience 05/2013; 33(18):7837-45. · 7.11 Impact Factor
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ABSTRACT: In line with the frontal hypothesis of aging, the ability to plan ahead undergoes substantial change during normal aging. Although impairments on the Tower of London planning task were reported earlier, associations between age-related declines and specific cognitive demands on planning have not been studied. Here we investigated the impact of search depth and goal ambiguity on planning, which impose demands on the depth and breadth of look-ahead processes, respectively. Besides an overall age-related decline in planning accuracy of 106 healthy older adults, differential search depth effects were found: Whereas planning accuracy of subjects in the early 60s was not affected by variations in search depth, between the ages of 65 and 76 years, accuracy was significantly decreased for high versus low levels of search depth. For subjects older than 76, different search depth levels did not further impact on accuracy, which was lowest overall. This nonlinear pattern may reflect differential impairments in fluid abilities and working memory capacity across various stages of older age. As no age-related effects of goal ambiguity were found, normal aging seems to be specifically sensitive to planning demands on the depth but not the breadth of anticipatory search processes. Hence, cognitive functions subserved by the prefrontal cortex experience differential development over the course of normal aging. (PsycINFO Database Record (c) 2013 APA, all rights reserved).
Developmental Psychology 04/2013; · 3.21 Impact Factor
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ABSTRACT: Prospective memory performance shows a decline in late adulthood. The present article examines the role of 3 main executive function facets (i.e., shifting, updating, and inhibition) as possible developmental mechanisms associated with these age effects. One hundred seventy-five young and 110 older adults performed a battery of cognitive tests including measures of prospective memory, shifting, updating, inhibition, working memory, and speed. Age effects were confirmed in prospective memory and also obtained in shifting, updating, and inhibition. Yet, facets of executive control differently predicted prospective memory performance: While inhibition and shifting were strong predictors of prospective memory performance and also explained age differences in prospective memory, updating was not related to prospective memory performance across adulthood. Furthermore, considering executive function measures increased the amount of explained variance in prospective remembering and reduced the influence of speed. Working memory was not revealed to serve as a significant predictor of prospective memory performance in the present study. These findings clarify the role of different facets of controlled attention on age effects in prospective memory and bear important conceptual implications: Results suggest that some but not all facets of executive functioning are important developmental mechanisms of prospective memory across adulthood beyond working memory and speed. Specifically, inhibition and shifting appear to be essential aspects of cognitive control involved in age-related prospective memory performance. (PsycINFO Database Record (c) 2012 APA, all rights reserved).
Developmental Psychology 11/2012; · 3.21 Impact Factor
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ABSTRACT: Planning of behavior relies on the integrity of the mid-dorsolateral prefrontal cortex (mid-dlPFC). Yet, only indirect evidence exists on the association of protracted maturation of dlPFC and continuing gains in planning performance post adolescence. Here, gray matter density of mid-dlPFC in young, healthy adults (18-32years) was regressed onto performance on the Tower of London planning task while accounting for moderating effects of age and sex on this interrelation. Multiple regression analysis revealed an association of planning performance and mid-dlPFC gray matter density that was especially strong in late adolescence and early twenties. As expected, for males better planning performance was linked to reduced gray matter density of mid-dlPFC, possibly due to maturational processes such as synaptic pruning. Most surprisingly, females showed an inverted, positive interrelation of planning performance and mid-dlPFC gray matter density, indicating that sexually dimorphic development of dlPFC continues during early adulthood. Age and sex are hence important moderators of the link between planning performance and gray matter density in mid-dlPFC. Consequently, the assessment of moderator effects in regression designs can significantly enhance understanding of brain-behavior relationships.
NeuroImage 08/2012; 63(3):1454-63. · 5.89 Impact Factor
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ABSTRACT: Understanding the functional neuroanatomy of planning and problem solving may substantially benefit from better insight into the chronology of the cognitive processes involved. Based on the assumption that regularities in cognitive processing are reflected in overtly observable eye-movement patterns, here we recorded eye movements while participants worked on Tower of London (TOL) problems that comprised an experimental manipulation of different task demands. Single-trial saccade-locked analyses revealed that higher demands on forming an internal problem representation were associated with an increased number of gaze alternations between start state and goal state, but did not show any effect on the durations of these inspections of the states. In contrast, higher demands on actual planning in terms of mental manipulations of working memory contents coincided with a prolonged duration of the very last inspection of the start state (i.e., immediately preceding movement execution) but did not show any effect on the number of gaze alterations. Differential task demands on internalization and planning processes during problem solving hence selectively affect different eye-movement parameters. Moreover, these findings complement previous neuroimaging data on dissociable contributions of left and right dorsolateral prefrontal cortex in problem solving with novel evidence for a corresponding dissociation in the eye-movement patterns reflecting the associated cognitive processes.
Brain and Cognition 06/2012; 80(1):160-9. · 3.17 Impact Factor
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ABSTRACT: In a companion study, eye-movement analyses in the Tower of London task (TOL) revealed independent indicators of functionally separable cognitive processes during problem solving, with processes of building up an internal representation of the problem preceding actual planning processes. These results imply that processes of internalization and planning should also be distinguishable in time and space with respect to concomitant brain activation patterns. To investigate this possibility, here we conducted analyses of fMRI data for left and right dorsolateral prefrontal cortex (dlPFC) during problem solving in the TOL task by accounting for the trial-by-trial variability of onsets and durations of the different cognitive processing stages. Comparisons between stimulus-locked and response-locked modeling approaches affirmed that activation in left dlPFC was elicited particularly during early processes of internalization, comprising the extraction of goal information and the generation of an internal problem representation, whereas activation in right dlPFC was predominantly attributable to later processes of mental transformations on this representation, that is planning proper. Thus, present data corroborate the proposal that often observed bilateral dlPFC activation patterns during complex cognitive tasks such as problem solving may reflect functionally and, to some extent, even temporally separable processes with opposing lateralizations.
Brain and Cognition 03/2012; 80(1):170-6. · 3.17 Impact Factor
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ABSTRACT: Planning impairments mark a well-documented consequence of neurodegenerative diseases such as Parkinson's disease (PD). Recently, using the Tower of London task we demonstrated that, rather than being generally impaired, PD patients selectively fail when planning requires flexible in-breadth search strategies. For a better understanding of the interindividual patterns underlying specific planning impairments, here we performed an explorative re-analysis of the original data using a latent-class model-based approach. Data-driven classification according to subjects' performance was based on a multinomial processing tree (MPT) model accommodating the impact of increased breadth versus depth of looking ahead during planning. In order to assess interindividual variability in coping with these different task demands, an extension of MPT models was used in which sample-immanent heterogeneity is accounted for by identifying different latent classes of individuals. Two latent classes were identified that differed considerably in performance for problems placing high demands on the depth of anticipatory search processes. In addition, these impairments were independent of PD diagnosis. However, latent-class mediated search depth-related deficits in planning performance were associated with poorer outcomes in dementia screenings, albeit sub-clinical. PD patients exhibited additional deficits related to the breadth of searching ahead. Taken together, results revealed dissociable impairments in specific planning processes within a single task of visuospatial problem solving. Present analyses put forward the hypothesis that cognitive sequelae of PD and sub-clinical signs of dementia may be related to differential patterns of planning impairments.
PLoS ONE 01/2012; 7(6):e38855. · 4.09 Impact Factor
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ABSTRACT: Most neuroimaging studies on planning report bilateral activations of the dorsolateral prefrontal cortex (dlPFC). Recently, these concurrent activations of left and right dlPFC have been shown to double dissociate with different cognitive demands imposed by the planning task: Higher demands on the extraction of task-relevant information led to stronger activation in left dlPFC, whereas higher demands on the integration of interdependent information into a coherent action sequence entailed stronger activation of right dlPFC. Here, we used continuous theta-burst stimulation (cTBS) to investigate the supposed causal structure-function mapping underlying this double dissociation. Two groups of healthy subjects (left-lateralized stimulation, n = 26; right-lateralized stimulation, n = 26) were tested within-subject on a variant of the Tower of London task following either real cTBS over dlPFC or sham stimulation over posterior parietal cortex. Results revealed that, irrespective of specific task demands, cTBS over left and right dlPFC was associated with a global decrease and increase, respectively, in initial planning times compared to sham stimulation. Moreover, no interaction between task demands and stimulation type (real vs. sham) and/or stimulation side (left vs. right hemisphere) were found. Together, against expectations from previous neuroimaging data, lateralized cTBS did not lead to planning-parameter specific changes in performance, but instead revealed a global asymmetric pattern of faster versus slower task processing after left versus right cTBS. This global asymmetry in the absence of any task-parameter specific impact of cTBS suggests that different levels of information processing may span colocalized, but independent axes of functional lateralization in the dlPFC. Hum Brain Mapp, 2011. © 2011 Wiley Periodicals, Inc.
Human Brain Mapping 10/2011; · 5.88 Impact Factor
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ABSTRACT: The neural mechanisms underlying spatial neglect are still disputed. Abnormal left parietal hyperactivation is proposed to lead to the rightward attentional bias, a clinical hallmark of neglect. Extinction, another deficit of visuospatial attention, is regarded as either a 'mild' form of neglect or a distinct syndrome. Although both neglect and extinction are typical syndromes of acute right hemispheric stroke, all imaging studies investigating these syndromes were conducted at least several weeks after stroke onset, in a phase when brain reorganization has already progressed. The present study aimed at comparing the activation patterns in acute stroke patients with neglect and extinction during visuospatial processing. Using functional magnetic resonance imaging, we examined the functional state of the attention system in 33 patients with a first ever stroke (53 ± 5 h after stroke onset) and age-matched healthy subjects (n = 15). All patients had embolic infarcts within the territory of the right middle cerebral artery. Patients were divided into three groups: (i) normal visuospatial processing (control patients, n = 11); (ii) patients with visual extinction but with no signs of neglect (n = 9); and (iii) patients with visual neglect (n = 13). While undergoing functional magnetic resonance imaging, patients performed a Posner-like task for visuospatial attention with detection of the targets in the left and right visual hemifields. Patients with neglect showed the expected imbalance in the left versus right parietal activation, which however, was present also in control and extinction patients, thus representing an epiphenomenon of the acute structural lesion in the right hemisphere. Compared with control patients, neglect was characterized by reduced activation in the right parietal and lateral occipital cortex, as well as in the left frontal eye field. In contrast, the activation pattern in patients with extinction differed from all other groups by an increased activation of the left prefrontal cortex. In both patients with neglect and extinction, detection of targets in the left hemifield correlated with an activation in the left prefrontal and parietal cortex. Thus at least in acute stroke, a relative hyperactivation of the left parietal cortex is not a particular characteristic of neglect. The specific signature of neglect is represented by the dysfunction of the right parietal and lateral occipital cortex. The function of the left attentional centres might provide a compensatory role after critical right hemisphere lesions and be relevant for the contralesional spatial processing.
Brain 09/2011; 134(Pt 11):3310-25. · 9.46 Impact Factor
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ABSTRACT: In clinical and experimental settings, planning ability is typically assessed using the Tower of London (ToL) or one of its variants. For enhancing the comparability across studies, a common ToL problem set was recently suggested comprising a collection of 4- to 7-move problems. Based on previous theoretical and empirical analyses of problem space and task structure, development of the problem set accounted particularly for the influence of structural problem parameters on the detection of individual differences in planning ability. To assess its adequacy as a clinical and research instrument, the present study evaluated the psychometric properties of the suggested problem set. Results showed a clear and nearly perfect linear increase of task difficulty across minimum moves. Given a broad range of item difficulty, high- and low-achieving subjects could be well discriminated. The test scores' split-half reliability (r = .72) and internal consistency (α = .69) were satisfactory. Taken together, the ToL problem set evaluated here proved to have good psychometric qualities and constitutes a conceptually sound basis for diagnostic and research purposes.
Psychological Assessment 08/2011; 24(1):46-53. · 2.99 Impact Factor
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ABSTRACT: It is well established that the mid-dorsolateral prefrontal cortex (dlPFC) plays a critical role in planning. Neuroimaging studies have yielded predominantly bilateral dlPFC activations, but the existence and nature of functionally specific contributions of left and right dlPFC have remained elusive. In recent experiments, 2 independent parameters have been identified which substantially determine planning: 1) the degree of interdependence between consecutive steps (search depth) and 2) the degree to which the configuration of the goal state renders the order of single steps either clearly evident or ambiguous (goal hierarchy). Thus, search depth affects the actual mental generation and evaluation of action sequences, whereas goal hierarchy reflects the extraction of goal information from an encountered problem. Here, both parameters were independently manipulated in an event-related functional magnetic resonance imaging study using the Tower of London task. Results revealed a double dissociation as indicated by a significant crossover interaction of hemisphere and task parameter: in left dlPFC, activations were stronger for higher demands on goal hierarchy than on search depth, whereas the reversed result emerged in right dlPFC. In conclusion, often observed bilateral patterns of dlPFC activation in complex tasks may reflect the concomitant operation of specific cognitive processes that show opposing lateralizations.
Cerebral Cortex 02/2011; 21(2):307-17. · 6.54 Impact Factor
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ABSTRACT: Cognitive, clinical, and neuroimaging studies on planning abilities most frequently implement the Tower of London task or one of its variants. Yet, cumulating evidence from a series of experiments suggests that the commonly used approximation of problem difficulty in terms of the minimum number of moves for goal attainment is too coarse a measure for the underlying cognitive operations, and in some cases may be even misleading. Rather, problem difficulty can be more specifically characterized by a set of structural task parameters such as the number and nature of optimal and suboptimal solution paths, the required search depths, the patterns of intermediate and goal moves, goal hierarchies and the associated degree of ambiguity in the sequential ordering of goal moves. First applications in developmental and patient studies have proven fruitful in targeting fundamental alterations of planning abilities in healthy and clinical conditions. In addition, recent evidence from neuroimaging shows that manipulations of problem structure relate to separate cognitive and neural processes and are accompanied by dissociable brain activation patterns. Here, we briefly review these structural problem parameters and the concepts behind. As controlling for task parameters and selecting a balanced problem set is a complex and error-prone endeavor, we further present TowerTool, a software solution that allows easy access to in-depth analysis of the problem structure of widely used planning tasks like the Tower of London, the Tower of Hanoi, and their variants. Thereby, we hope to encourage and facilitate the implementation of structurally balanced task sets in future studies on planning and to promote transfer between the cognitive, developmental, and clinical neurosciences.
Behavioural brain research 01/2011; 216(1):1-8. · 3.22 Impact Factor
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ABSTRACT: By mapping the dynamics of brain reorganization, functional magnetic resonance imaging MRI (fMRI) has allowed for significant progress in understanding cerebral plasticity phenomena after a stroke. However, cerebro-vascular diseases can affect blood oxygen level dependent (BOLD) signal. Cerebral autoregulation is a primary function of cerebral hemodynamics, which allows to maintain a relatively constant blood flow despite changes in arterial blood pressure and perfusion pressure. Cerebral autoregulation is reported to become less effective in the early phases post-stroke. This study investigated whether any impairment of cerebral hemodynamics that occurs during the acute and the subacute phases of ischemic stroke is related to changes in BOLD response. We enrolled six aphasic patients affected by acute stroke. All patients underwent a Transcranial Doppler to assess cerebral autoregulation (Mx index) and fMRI to evaluate the amplitude and the peak latency (time to peak-TTP) of BOLD response in the acute (i.e., within four days of stroke occurrence) and the subacute (i.e., between five and twelve days after stroke onset) stroke phases.
As patients advanced from the acute to subacute stroke phase, the affected hemisphere presented a BOLD TTP increase (p = 0.04) and a deterioration of cerebral autoregulation (Mx index increase, p = 0.046). A similar but not significant trend was observed also in the unaffected hemisphere. When the two hemispheres were grouped together, BOLD TTP delay was significantly related to worsening cerebral autoregulation (Mx index increase) (Spearman's rho = 0.734; p = 0.01).
The hemodynamic response function subtending BOLD signal may present a delay in peak latency that arises as patients advance from the acute to the subacute stroke phase. This delay is related to the deterioration of cerebral hemodynamics. These findings suggest that remodeling the fMRI hemodynamic response function in the different phases of stroke may optimize the detection of BOLD signal changes.
BMC Neuroscience 12/2009; 10:151. · 3.04 Impact Factor
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ABSTRACT: In the present study, we identified the most probable trajectories of point-to-point segregated connections between functional attentional centers using a combination of functional magnetic resonance imaging and a novel diffusion tensor imaging-based algorithm for pathway extraction. Cortical regions activated by a visuospatial attention task were subsequently used as seeds for probabilistic fiber tracking in 26 healthy subjects. Combining probability maps of frontal and temporoparietal regions yielded a network that consisted of dorsal and ventral connections. The dorsal connections linked temporoparietal cortex with the frontal eye field and area 44 of the inferior frontal gyrus (IFG). Traveling along superior longitudinal and arcuate fascicles, these fibers are well described in relation to spatial attention. However, the ventral connections, which traveled in the white matter between insula (INS) cortex and putamen parallel to the sylvian fissure, were not previously described for visuospatial attention. Linking temporoparietal cortex with anterior INS and area 45 of IFG, these connections may provide an anatomical substrate for crossmodal cortical integration needed for stimulus perception and response in relation to current intention. The newly anatomically described integral network for visuospatial attention might improve the understanding of spatial attention deficits after white matter lesions.
Cerebral Cortex 05/2009; 20(1):121-9. · 6.54 Impact Factor
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ABSTRACT: Identifying overtly observable indicators of cognitive processes should provide a promising basis for a more precise tracking of the associated cognitive and neural events. In the current study we used recordings of eye movements to gain deeper insight into the time course of visuospatial problem solving as measured by the Tower of London. Single-trial, saccade-locked analyses revealed that, despite the complexity of the implemented task, gaze alternations between start and goal state followed a highly regular pattern. Consistent with the buildup of an internal representation, the first two fixations were of constant duration and unaffected by experimental variations of planning demands. Instead, planning manipulations exclusively influenced the duration of the very last fixation before problem execution. Our results demonstrate that different phases of complex cognition can be identified on a single-trial level using eye movement analyses.
Psychophysiology 05/2009; 46(4):818-30. · 3.29 Impact Factor
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ABSTRACT: The ability to plan and search ahead is essential for problem solving in most situations in everyday life. To investigate the development of planning and related processes, a sample of four- and five-year-old children was examined in a variant of the Tower of London, a frequently used neuropsychological assessment tool of planning abilities. The applied problems either required searching ahead for optimal solution or were solvable by pure step-by-step forward processing. Furthermore, the ambiguity of subgoal ordering was varied. Results revealed an age-related effect of search depth: the four-year olds' planning accuracy was particularly decreased in problems demanding search ahead, while five-year olds mastered both problem types equally well. Interestingly, this interaction between age and search depth could not be accounted for by measures of working memory and inhibition. Differential effects of age were also found for subgoal ordering with respect to initial planning and movement execution times. In sum, planning abilities showed considerable development during late kindergarten age that appeared to be specifically associated with the integration and back-validation of the anticipated consequences of internally modeled actions. The present study demonstrates that a careful consideration of problem structure may greatly enhance the insights gained from the application of a routinely used assessment tool, the Tower of London. This may be especially advantageous when addressing specific subpopulations such as children or clinical samples.
Brain and Cognition 09/2008; 67(3):360-70. · 3.17 Impact Factor
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ABSTRACT: Despite the large number of behavioral and functional neuroimaging studies employing the Tower of London (ToL), the task's structural parameters and particularly their impact on planning have not been addressed so far. In this paper, we highlight the structural properties of ToL problems and provide evidence for their systematic and substantial effects on the cognitive processes involved in planning. In a problem set with three-move problems, the following structural parameters were experimentally manipulated: the ambiguity of goal hierarchy, the demand for subgoal generation, and the existence of suboptimal alternatives. Analysis of preplanning time as an indicator for the planning process revealed highly significant effects for all three parameters which seems to reflect differences in cognitive processing due to structural task properties. Therefore, we suggest that the common consideration of ToL problem difficulty solely in terms of the minimum number of moves is not sufficient. Moreover, the applied problem sets should be more carefully selected and their structural parameters explicitly noted.
Cognitive Brain Research 09/2004; 20(3):462-72. · 3.77 Impact Factor
