In a cancellation task, a participant is required to search for and cross out ("cancel") targets, which are usually embedded among distractor stimuli. The number of cancelled targets and their location can be used to diagnose the neglect syndrome after stroke. In addition, the organization of search provides a potentially useful way to measure executive control over multitarget search. Although many useful cancellation measures have been introduced, most fail to make their way into research studies and clinical practice due to the practical difficulty of acquiring such parameters from traditional pen-and-paper measures. Here we present new, open-source software that is freely available to all. It allows researchers and clinicians to flexibly administer computerized cancellation tasks using stimuli of their choice, and to directly analyze the data in a convenient manner. The automated analysis suite provides output that includes almost all of the currently existing measures, as well as several new ones introduced here. All tasks can be performed using either a computer mouse or a touchscreen as an input device, and an online version of the task runtime is available for tablet devices. A summary of the results is produced in a single A4-sized PDF document, including high quality data visualizations. For research purposes, batch analysis of large datasets is possible. In sum, CancellationTools allows users to employ a flexible, computerized cancellation task, which provides extensive benefits and ease of use.
Our knowledge of the anatomical organization of the human brain in health and disease draws heavily on the study of patients with focal brain lesions. Historically the first method of mapping brain function, it is still potentially the most powerful, establishing the necessity of any putative neural substrate for a given function or deficit. Great inferential power, however, carries a crucial vulnerability: without stronger alternatives any consistent error cannot be easily detected. A hitherto unexamined source of such error is the structure of the high-dimensional distribution of patterns of focal damage, especially in ischaemic injury-the commonest aetiology in lesion-deficit studies-where the anatomy is naturally shaped by the architecture of the vascular tree. This distribution is so complex that analysis of lesion data sets of conventional size cannot illuminate its structure, leaving us in the dark about the presence or absence of such error. To examine this crucial question we assembled the largest known set of focal brain lesions (n = 581), derived from unselected patients with acute ischaemic injury (mean age = 62.3 years, standard deviation = 17.8, male:female ratio = 0.547), visualized with diffusion-weighted magnetic resonance imaging, and processed with validated automated lesion segmentation routines. High-dimensional analysis of this data revealed a hidden bias within the multivariate patterns of damage that will consistently distort lesion-deficit maps, displacing inferred critical regions from their true locations, in a manner opaque to replication. Quantifying the size of this mislocalization demonstrates that past lesion-deficit relationships estimated with conventional inferential methodology are likely to be significantly displaced, by a magnitude dependent on the unknown underlying lesion-deficit relationship itself. Past studies therefore cannot be retrospectively corrected, except by new knowledge that would render them redundant. Positively, we show that novel machine learning techniques employing high-dimensional inference can nonetheless accurately converge on the true locus. We conclude that current inferences about human brain function and deficits based on lesion mapping must be re-evaluated with methodology that adequately captures the high-dimensional structure of lesion data.
Individuals with mutation in the lysosomal enzyme glucocerebrosidase (GBA) gene are at significantly high risk of developing Parkinson's disease with cognitive deficit. We examined whether visual short-term memory impairments, long associated with patients with Parkinson's disease, are also present in GBA-positive individuals-both with and without Parkinson's disease. Precision of visual working memory was measured using a serial order task in which participants observed four bars, each of a different colour and orientation, presented sequentially at screen centre. Afterwards, they were asked to adjust a coloured probe bar's orientation to match the orientation of the bar of the same colour in the sequence. An additional attentional 'filtering' condition tested patients' ability to selectively encode one of the four bars while ignoring the others. A sensorimotor task using the same stimuli controlled for perceptual and motor factors. There was a significant deficit in memory precision in GBA-positive individuals-with or without Parkinson's disease-as well as GBA-negative patients with Parkinson's disease, compared to healthy controls. Worst recall was observed in GBA-positive cases with Parkinson's disease. Although all groups were impaired in visual short-term memory, there was a double dissociation between sources of error associated with GBA mutation and Parkinson's disease. The deficit observed in GBA-positive individuals, regardless of whether they had Parkinson's disease, was explained by a systematic increase in interference from features of other items in memory: misbinding errors. In contrast, impairments in patients with Parkinson's disease, regardless of GBA status, was explained by increased random responses. Individuals who were GBA-positive and also had Parkinson's disease suffered from both types of error, demonstrating the worst performance. These findings provide evidence for dissociable signature deficits within the domain of visual short-term memory associated with GBA mutation and with Parkinson's disease. Identification of the specific pattern of cognitive impairment in GBA mutation versus Parkinson's disease is potentially important as it might help to identify individuals at risk of developing Parkinson's disease.
Although the pre-supplementary motor area (pre-SMA) is one of the most frequently reported areas of activation in functional imaging studies, the role of this brain region in cognition is still a matter of intense debate. Here we present a patient with a focal lesion of caudal pre-SMA who displays a selective deficit in updating a response plan to switch actions, but shows no impairment when required to withhold a response - stopping.
Delayed adjustment tasks have recently been developed to examine working memory (WM) precision, that is, the resolution with which items maintained in memory are recalled. However, despite their emerging use in experimental studies of healthy people, evaluation of patient populations is sparse. We first investigated the validity of adjustment tasks, comparing precision with classical span measures of memory across the lifespan in 114 people. Second, we asked whether precision measures can potentially provide a more sensitive measure of WM than traditional span measures. Specifically, we tested this hypothesis examining WM in a group with early, untreated Parkinson's disease (PD) and its modulation by subsequent treatment on dopaminergic medication. Span measures correlated with precision across the lifespan: in children, young, and elderly participants. However, they failed to detect changes in WM in PD patients, either pre- or post-treatment initiation. By contrast, recall precision was sensitive enough to pick up such changes. PD patients pre-medication were significantly impaired compared to controls, but improved significantly after 3 months of being established on dopaminergic medication. These findings suggest that precision methods might provide a sensitive means to investigate WM and its modulation by interventions in clinical populations.
Working memory (WM) is a core cognitive process fundamental to human behavior, yet the mechanisms underlying it remain highly controversial. Here we provide a new framework for understanding retrieval of information from WM, conceptualizing it as a decision based on the quality of internal evidence. Recent findings have demonstrated that precision of WM decreases with memory load. If WM retrieval uses a decision process that depends on memory quality, systematic changes in response time distribution should occur as a function of WM precision. We asked participants to view sample arrays and, after a delay, report the direction of change in location or orientation of a probe. As WM precision deteriorated with increasing memory load, retrieval time increased systematically. Crucially, the shape of reaction time distributions was consistent with a linear accumulator decision process. Varying either task relevance of items or maintenance duration influenced memory precision, with corresponding shifts in retrieval time. These results provide strong support for a decision-making account of WM retrieval based on noisy storage of items. Furthermore, they show that encoding, maintenance, and retrieval in WM need not be considered as separate processes, but may instead be conceptually unified as operations on the same noise-limited, neural representation.
The role of the thalamus in high-level cognition-attention, working memory (WM), rule-based learning, and decision making-remains poorly understood, especially in comparison to that of cortical frontoparietal networks [1-3]. Studies of visual thalamus have revealed important roles for pulvinar and lateral geniculate nucleus in visuospatial perception and attention [4-10] and for mediodorsal thalamus in oculomotor control . Ventrolateral thalamus contains subdivisions devoted to action control as part of a circuit involving the basal ganglia [12, 13] and motor, premotor, and prefrontal cortices , whereas anterior thalamus forms a memory network in connection with the hippocampus . This connectivity profile suggests that ventrolateral and anterior thalamus may represent a nexus between mnemonic and control functions, such as action or attentional selection. Here, we characterize the role of thalamus in the interplay between memory and visual attention. We show that ventrolateral lesions impair the influence of WM representations on attentional deployment. A subsequent fMRI study in healthy volunteers demonstrates involvement of ventrolateral and, notably, anterior thalamus in biasing attention through WM contents. To further characterize the memory types used by the thalamus to bias attention, we performed a second fMRI study that involved learning of stimulus-stimulus associations and their retrieval from long-term memory to optimize attention in search. Responses in ventrolateral and anterior thalamic nuclei tracked learning of the predictiveness of these abstract associations and their use in directing attention. These findings demonstrate a key role for human thalamus in higher-level cognition, notably, in mnemonic biasing of attention.
Working memory is widely considered to be limited in capacity, holding a fixed, small number of items, such as Miller's 'magical number' seven or Cowan's four. It has recently been proposed that working memory might better be conceptualized as a limited resource that is distributed flexibly among all items to be maintained in memory. According to this view, the quality rather than the quantity of working memory representations determines performance. Here we consider behavioral and emerging neural evidence for this proposal.
Apathy is a common but poorly understood condition with a wide societal impact observed in several brain disorders as well as, to some extent, in the normal population. Hence the need for better characterization of the underlying mechanisms. The processes by which individuals decide to attribute physical effort to obtain rewards might be particularly relevant to relate to apathy traits. Here, we designed two paradigms to assess individual differences in physical effort production and effort-based decision-making and their relation to apathy in healthy people. Apathy scores were measured using a modified version of the Lille Apathy Rating Scale, suitable for use in a non-clinical population. In the first study, apathy scores were correlated with the degree to which stake (reward on offer) and difficulty level impacts on physical effort production. Individuals with relatively high apathy traits showed an increased modulation of effort while more motivated individuals generally exerted greater force across different levels of stake. To clarify the underlying mechanisms for this behavior, we designed a second task that allows independent titration of stake and effort levels for which subjects are willing to engage in an effortful response to obtain a reward. Our results suggest that apathy traits in the normal population are related to the way reward subjectively affects the estimation of effort costs, and more particularly manifest as decreased willingness to exert effort when rewards are small, or below threshold. The tasks we introduce here may provide useful tools to further investigate apathy in clinical populations.
Complex manual tasks-everything from buttoning up a shirt to playing the piano-fundamentally involve two components: (1) generating specific patterns of muscle activity (here, termed "synergies"); and (2) stringing these into purposeful sequences. Although transcranial direct current stimulation (tDCS) of the primary motor cortex (M1) has been found to increase the learning of motor sequences, it is unknown whether it can similarly facilitate motor synergy learning. Here, we determined the effects of tDCS on the learning of motor synergies using a novel hand configuration task that required the production of difficult muscular activation patterns. Bihemispheric tDCS was applied to M1 of healthy, right-handed human participants during 4 d of repetitive left-hand configuration training in a double-blind design. tDCS augmented synergy learning, leading subsequently to faster and more synchronized execution. This effect persisted for at least 4 weeks after training. Qualitatively similar tDCS-associated improvements occurred during training of finger sequences in a separate subject cohort. We additionally determined whether tDCS only improved the acquisition of motor memories for specific synergies/sequences or whether it also facilitated more general parts of the motor representations, which could be transferred to novel movements. Critically, we observed that tDCS effects generalized to untrained hand configurations and untrained finger sequences (i.e., were nonspecific), as well as to the untrained hand (i.e., were effector-independent). Hence, bihemispheric tDCS may be a promising adjunct to neurorehabilitative training regimes, in which broad transfer to everyday tasks is highly desirable.
Emerging evidence suggests that items held in working memory (WM) might not all be in the same representational state. One item might be privileged over others, making it more accessible and thereby recalled with greater precision. Here, using transcranial magnetic stimulation (TMS), we provide causal evidence in human participants that items in WM are differentially susceptible to disruptive TMS, depending on their state, determined either by task relevance or serial position. Across two experiments, we applied TMS to area MT+ during the WM retention of two motion directions. In Experiment 1, we used an "incidental cue" to bring one of the two targets into a privileged state. In Experiment 2, we presented the targets sequentially so that the last item was in a privileged state by virtue of recency. In both experiments, recall precision of motion direction was differentially affected by TMS, depending on the state of the memory target at the time of disruption. Privileged items were recalled with less precision, whereas nonprivileged items were recalled with higher precision. Thus, only the privileged item was susceptible to disruptive TMS over MT+. By contrast, precision of the nonprivileged item improved either directly because of facilitation by TMS or indirectly through reduced interference from the privileged item. Our results provide a unique line of evidence, as revealed by TMS over a posterior sensory brain region, for at least two different states of item representation in WM.
Working memory and attention are intimately connected. However, understanding the relationship between the two is challenging. Currently, there is an important controversy about whether objects in working memory are maintained automatically or require resources that are also deployed for visual or auditory attention. Here we investigated the effects of loading attention resources on precision of visual working memory, specifically on correct maintenance of feature-bound objects, using a dual-task paradigm. Participants were presented with a memory array and were asked to remember either direction of motion of random dot kinematograms of different colour, or orientation of coloured bars. During the maintenance period, they performed a secondary visual or auditory task, with varying levels of load. Following a retention period, they adjusted a coloured probe to match either the motion direction or orientation of stimuli with the same colour in the memory array. This allowed us to examine the effects of an attention-demanding task performed during maintenance on precision of recall on the concurrent working memory task. Systematic increase in attention load during maintenance resulted in a significant decrease in overall working memory performance. Changes in overall performance were specifically accompanied by an increase in feature misbinding errors: erroneous reporting of nontarget motion or orientation. Thus in trials where attention resources were taxed, participants were more likely to respond with nontarget values rather than simply making random responses. Our findings suggest that resources used during attention-demanding visual or auditory tasks also contribute to maintaining feature-bound representations in visual working memory-but not necessarily other aspects of working memory.
Working memory declines with normal aging, but the nature of this impairment is debated. Studies based on detecting changes to arrays of visual objects have identified two possible components to age-related decline: a reduction in the number of items that can be stored, or a deficit in maintaining the associations (bindings) between individual object features. However, some investigations have reported intact binding with aging, and specific deficits arising only in Alzheimers disease. Here, using a recently developed continuous measure of recall fidelity, we tested the precision with which adults of different ages could reproduce from memory the orientation and color of a probed array item. The results reveal a further component of cognitive decline: an age-related decrease in the resolution with which visual information can be maintained in working memory. This increase in recall variability with age was strongest under conditions of greater memory load. Moreover, analysis of the distribution of errors revealed that older participants were more likely to incorrectly report one of the unprobed items in memory, consistent with an age-related increase in misbinding. These results indicate a systematic decline with age in working memory resources that can be recruited to store visual information. The paradigm presented here provides a sensitive index of both memory resolution and feature binding, with the potential for assessing their modulation by interventions. The findings have implications for understanding the mechanisms underpinning working memory deficits in both health and disease.
Trait sensation-seeking, defined as a need for varied, complex, and intense sensations, represents a relatively underexplored hedonic drive in human behavioral neuroscience research. It is related to increased risk for a range of behaviors including substance use, gambling, and risky sexual practice. Individual differences in self-reported sensation-seeking have been linked to brain dopamine function, particularly at D2-like receptors, but so far no causal evidence exists for a role of dopamine in sensation-seeking behavior in humans. Here, we investigated the effects of the selective D2/D3 agonist cabergoline on performance of a probabilistic risky choice task in healthy humans using a sensitive within-subject, placebo-controlled design. Cabergoline significantly influenced the way participants combined different explicit signals regarding probability and loss when choosing between response options associated with uncertain outcomes. Importantly, these effects were strongly dependent on baseline sensation-seeking score. Overall, cabergoline increased sensitivity of choice to information about probability of winning; while decreasing discrimination according to magnitude of potential losses associated with different options. The largest effects of the drug were observed in participants with lower sensation-seeking scores. These findings provide evidence that risk-taking behavior in humans can be directly manipulated by a dopaminergic drug, but that the effectiveness of such a manipulation depends on baseline differences in sensation-seeking trait. This emphasizes the importance of considering individual differences when investigating manipulation of risky decision-making, and may have relevance for the development of pharmacotherapies for disorders involving excessive risk-taking in humans, such as pathological gambling.
Some prominent studies have claimed that the medial temporal lobe is not involved in retention of information over brief intervals of just a few seconds. However, in the last decade several investigations have reported that patients with medial temporal lobe damage exhibit an abnormally large number of errors when required to remember visual information over brief intervals. But the nature of the deficit and the type of error associated with medial temporal lobe lesions remains to be fully established. Voltage-gated potassium channel complex antibody-associated limbic encephalitis has recently been recognized as a form of treatable autoimmune encephalitis, frequently associated with imaging changes in the medial temporal lobe. Here, we tested a group of these patients using two newly developed visual short-term memory tasks with a sensitive, continuous measure of report. These tests enabled us to study the nature of reporting errors, rather than only their frequency. On both paradigms, voltage-gated potassium channel complex antibody patients exhibited larger errors specifically when several items had to be remembered, but not for a single item. Crucially, their errors were strongly associated with an increased tendency to report the property of the wrong item stored in memory, rather than simple degradation of memory precision. Thus, memory for isolated aspects of items was normal, but patients were impaired at binding together the different properties belonging to an item, e.g. spatial location and object identity, or colour and orientation. This occurred regardless of whether objects were shown simultaneously or sequentially. Binding errors support the view that the medial temporal lobe is involved in linking together different types of information, potentially represented in different parts of the brain, regardless of memory duration. Our novel behavioural measures also have the potential to assist in monitoring response to treatment in patients with memory disorders, such as those with voltage-gated potassium channel complex antibody limbic encephalitis.
In Parkinsons disease the degree of motor impairment can be classified with respect to tremor dominant and akinetic rigid features. While tremor dominance and akinetic rigidity might represent two ends of a continuum rather than discrete entities, it would be important to have non-invasive markers of any biological differences between them in vivo, to assess disease trajectories and response to treatment, as well as providing insights into the underlying mechanisms contributing to heterogeneity within the Parkinsons disease population.
Impulse control disorders (ICDs) and apathy are recognized as two important neuropsychiatric syndromes associated with Parkinsons disease (PD), but as yet we understand very little about the cognitive mechanisms underlying them. Here, we review emerging findings, from both human and animal studies, that suggest that impulsivity and apathy are opposite extremes of a dopamine-dependent spectrum of motivated decision making. We first argue that there is strong support for a hypodopaminergic state in PD patients with apathy, as well as for an association between dopamine therapy and development of ICDs. However, there is little evidence for a clear dose-response relationship, and great heterogeneity of findings. We argue that dopaminergic state on its own is an insufficient explanation, and suggest instead that there is now substantial evidence that both apathy and impulsivity are in fact multi-dimensional syndromes, with separate, dissociable mechanisms underlying their surface manifestations. Some of these mechanisms might be dopamine-dependent. According to this view, individuals diagnosed as impulsive or apathetic may have very different mechanisms underlying their clinical states. We propose that impulsivity and apathy can arise from dissociable deficits in option generation, option selection, action initiation or inhibition and learning. Review of the behavioural and neurobiological evidence leads us to a new conceptual framework that might help understand the variety of functional deficits seen in PD.
Sustained attention is an essential brain function that enables a subject to maintain attention level over the time of a task. In previous work, the right inferior parietal lobe (IPL) has been reported as one of the main brain regions related to sustained attention, however, the right lateralization of vigilance/sustained attention is unclear because information about the network for sustained attention is traditionally provided by neglect patients who typically have right brain damage. Here, we investigated sustained attention by applying a virtual lesion technique, transcranial magnetic stimulation (TMS), over the left and right superior parietal lobe (SPL) and IPL. We used two different types of visual sustained attention tasks: spatial (location based) and non-spatial (feature based). When the participants performed the spatial task, repetitive TMS (rTMS) over either the right or left IPL induced a significant decrement of sustained attention causing a progressive increment of errors and response time. In contrast, participants performance was not changed by rTMS on the non-spatial task. Also, omission errors (true negative) gradually increased with time on right and left IPL rTMS conditions, while commission errors (false positive) were relatively stable. These findings suggest that the maintenance of attention, especially in tasks regarding spatial location, is not uniquely lateralized to the right IPL, but may also involve participation of the left IPL.
Patients with alien hand syndrome (AHS) experience making apparently deliberate and purposeful movements with their hand against their will. However, the mechanisms contributing to these involuntary actions remain poorly understood. Here, we describe two experimental investigations in a patient with corticobasal syndrome (CBS) with alien hand behaviour in her right hand. First, we show that responses with the alien hand are made significantly more quickly to images of objects which afford an action with that hand compared to objects which afford an action with the unaffected hand. This finding suggests that involuntary grasping behaviours in AHS might be due to exaggerated, automatic motor activation evoked by objects which afford actions with that limb. Second, using a backwards masked priming task, we found normal automatic inhibition of primed responses in the patients unaffected hand, but importantly there was no evidence of such suppression in the alien limb. Taken together, these findings suggest that grasping behaviours in AHS may result from exaggerated object affordance effects, which might potentially arise from disrupted inhibition of automatically evoked responses.
What is the purpose of attention? One avenue of research has led to the proposal that attention might be crucial for gathering information about the environment, while other lines of study have demonstrated how attention may play a role in guiding behavior to rewarded options. Many experiments that study attention require participants to make a decision based on information acquired discretely at one point in time. In real-world situations, however, we are usually not presented with information about which option to select in such a manner. Rather we must initially search for information, weighing up reward values of options before we commit to a decision. Here, we propose that attention plays a role in both foraging for information and foraging for value. When foraging for information, attention is guided toward the unknown. When foraging for reward, attention is guided toward high reward values, allowing decision-making to proceed by accept-or-reject decisions on the currently attended option. According to this account, attention can be regarded as a low-cost alternative to moving around and physically interacting with the environment-"teleforaging"-before a decision is made to interact physically with the world. To track the timecourse of attention, we asked participants to seek out and acquire information about two gambles by directing their gaze, before choosing one of them. Participants often made multiple refixations on items before making a decision. Their eye movements revealed that early in the trial, attention was guided toward information, i.e., toward locations that reduced uncertainty about value. In contrast, late in the trial, attention was guided by expected value of the options. At the end of the decision period, participants were generally attending to the item they eventually chose. We suggest that attentional foraging shifts from an uncertainty-driven to a reward-driven mode during the evolution of a decision, permitting decisions to be made by an engage-or-search strategy.
Self-regulation of brain activity in humans based on real-time feedback of functional magnetic resonance imaging (fMRI) signal is emerging as a potentially powerful, new technique. Here, we assessed whether patients with Parkinsons disease (PD) are able to alter local brain activity to improve motor function. Five patients learned to increase activity in the supplementary motor complex over two fMRI sessions using motor imagery. They attained as much activation in this target brain region as during a localizer procedure with overt movements. Concomitantly, they showed an improvement in motor speed (finger tapping) and clinical ratings of motor symptoms (37% improvement of the motor scale of the Unified Parkinsons Disease Rating Scale). Activation during neurofeedback was also observed in other cortical motor areas and the basal ganglia, including the subthalamic nucleus and globus pallidus, which are connected to the supplementary motor area (SMA) and crucial nodes in the pathophysiology of PD. A PD control group of five patients, matched for clinical severity and medication, underwent the same procedure but did not receive feedback about their SMA activity. This group attained no control of SMA activation and showed no motor improvement. These findings demonstrate that self-modulation of cortico-subcortical motor circuits can be achieved by PD patients through neurofeedback and may result in clinical benefits that are not attainable by motor imagery alone.
In recent years, diffusion-weighted magnetic resonance imaging (DW-MRI) has been increasingly used to explore the relationship between white matter structure and cognitive function. This technique uses the passive diffusion of water molecules to infer properties of the surrounding tissue. DW-MRI has been extensively employed to investigate how individual differences in behavior are related to variability in white matter microstructure on a range of different cognitive tasks and also to examine the effect experiential learning might have on brain structural connectivity. Using diffusion tensor tractography, large white matter pathways have been traced in vivo and used to explore patterns of white matter projections between different brain regions. Recent findings suggest that diffusion-weighted imaging might even be used to measure functional differences in water diffusion during task performance. This review describes some research highlights in diffusion-weighted imaging and how this technique can be employed to further our understanding of cognitive function.
Impulsivity is often characterized by rapid decisions under risk, but most current tests of decision-making do not impose time pressures on participants choices. Here we introduce a new traffic lights test which requires people to choose whether to program a risky, early eye movement before a traffic light turns green (earning them high rewards or a penalty) or wait for the green light before responding to obtain a small reward instead. Young participants demonstrated bimodal responses: an early, high-risk and a later, low-risk set of choices. By contrast, elderly people invariably waited for the green light and showed little risk-taking. Performance could be modeled as a race between two rise-to-threshold decision processes, one triggered by the green light and the other initiated before it. The test provides a useful measure of rapid decision-making under risk, with the potential to reveal how this process alters with aging or in patient groups.
Viewing objects can result in automatic, partial activation of motor plans associated with them-"object affordance". Here, we recorded grip force simultaneously from both hands in an object affordance task to investigate the effects of conflict between coactivated responses. Participants classified pictures of objects by squeezing force transducers with their left or right hand. Responses were faster on trials where the object afforded an action with the same hand that was required to make the response (congruent trials) compared to the opposite hand (incongruent trials). In addition, conflict between coactivated responses was reduced if it was experienced on the preceding trial, just like Gratton adaptation effects reported in "conflict" tasks (e.g., Eriksen flanker). This finding suggests that object affordance demonstrates conflict effects similar to those shown in other stimulus-response mapping tasks and thus could be integrated into the wider conceptual framework on overlearnt stimulus-response associations. Corrected erroneous responses occurred more frequently when there was conflict between the afforded response and the response required by the task, providing direct evidence that viewing an object activates motor plans appropriate for interacting with that object. Recording continuous grip force, as here, provides a sensitive way to measure coactivated responses in affordance tasks.
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.
Recent neurophysiological and imaging studies have investigated how neural representations underlying working memory (WM) are dynamically updated for objects presented sequentially. Although such studies implicate information encoded in oscillatory activity across distributed brain networks, interpretation of findings depends crucially on the underlying conceptual model of how memory resources are distributed. Here, we quantify the fidelity of human memory for sequences of colored stimuli of different orientation. The precision with which each orientation was recalled declined with increases in total memory load, but also depended on when in the sequence it appeared. When one item was prioritized, its recall was enhanced, but with corresponding decrements in precision for other objects. Comparison with the same number of items presented simultaneously revealed an additional performance cost for sequential display that could not be explained by temporal decay. Memory precision was lower for sequential compared with simultaneous presentation, even when each item in the sequence was presented at a different location. Importantly, stochastic modeling established this cost for sequential display was due to misbinding object features (color and orientation). These results support the view that WM resources can be dynamically and flexibly updated as new items have to be stored, but redistribution of resources with the addition of new items is associated with misbinding object features, providing important constraints and a framework for interpreting neural data.
Neurons in medial frontal cortex have been found to distinguish between whether an animal or its partner is responding on a turn-taking task, but are they really the basis of a social learning mechanism?
Impulsivity is a multifaceted personality construct associated with numerous psychiatric disorders. Recent research has characterized four facets of impulsivity: "urgency" (the tendency to act rashly especially in the context of distress or cravings); "lack of premeditation" (not envisaging the consequences of actions); "lack of perseverance" (not staying focused on a task); and "sensation seeking" (engaging in exciting activities). Urgency is particularly associated with clinical populations and problematic disinhibited behavior.
Recent studies investigating working memory for location, color, and orientation support a dynamic resource model. We examined whether this might also apply to motion, using random dot kinematograms (RDKs) presented sequentially or simultaneously. Mean precision for motion direction declined as sequence length increased, with precision being lower for earlier RDKs. Two alternative models of working memory were compared specifically to distinguish between the contributions of different sources of error that corrupt memory (W. Zhang & S. J. Luck, 2008 vs. P. M. Bays, R. F. G. Catalao, & M. Husain, 2009). The latter provided a significantly better fit for the data, revealing that decrease in memory precision for earlier items is explained by an increase in interference from other items in a sequence rather than random guessing or a temporal decay of information. Misbinding feature attributes is an important source of error in working memory. Precision of memory for motion direction decreased when two RDKs were presented simultaneously as transparent surfaces, compared to sequential RDKs. However, precision was enhanced when one motion surface was prioritized, demonstrating that selective attention can improve recall precision. These results are consistent with a resource model that can be used as a general conceptual framework for understanding working memory across a range of visual features.
The process of encoding a visual scene into working memory has previously been studied using binary measures of recall. Here, we examine the temporal evolution of memory resolution, based on observers ability to reproduce the orientations of objects presented in brief, masked displays. Recall precision was accurately described by the interaction of two independent constraints: an encoding limit that determines the maximum rate at which information can be transferred into memory and a separate storage limit that determines the maximum fidelity with which information can be maintained. Recall variability decreased incrementally with time, consistent with a parallel encoding process in which visual information from multiple objects accumulates simultaneously in working memory. No evidence was observed for a limit on the number of items stored. Cuing one display item with a brief flash led to rapid development of a recall advantage for that item. This advantage was short-lived if the cue was simply a salient visual event but was maintained if it indicated an object of particular relevance to the task. These cuing effects were observed even for items that had already been encoded into memory, indicating that limited memory resources can be rapidly reallocated to prioritize salient or goal-relevant information.
We report a case of a 45-year-old man presenting with asomatognosia, or loss of body part ownership, when he experienced difficulty acknowledging that his arm was his own. His symptoms might easily have been considered to be of psychiatric origin. Instead they turned out to be due to highly focal stroke secondary to carotid dissection, an important and often missed cause of stroke in younger patients.
Although many functional imaging studies have reported frontal activity associated with "cognitive control" tasks, little is understood about factors underlying individual differences in performance. Here we compared the behavior and brain structure of healthy controls with fighter pilots, an expert group trained to make precision choices at speed in the presence of conflicting cues. Two different behavioral paradigms--Eriksen Flanker and change of plan tasks--were used to assess the influence of distractors and the ability to update ongoing action plans. Fighter pilots demonstrated superior cognitive control as indexed by accuracy and postconflict adaptation on the Flanker task, but also showed increased sensitivity to irrelevant, distracting choices. By contrast, when pilots were examined on their ability to inhibit a current action plan in favor of an alternative response, their performance was no better than the control group. Diffusion weighted imaging revealed differences in white matter radial diffusivity between pilots and controls not only in the right dorsomedial frontal region but also in the right parietal lobe. Moreover, analysis of individual differences in reaction time costs for conflict trials on the Flanker task demonstrated significant correlations with radial diffusivity at these locations, but in different directions. Postconflict adaptation effects, however, were confined to the dorsomedial frontal locus. The findings demonstrate that in humans expert cognitive control may surprisingly be mediated by enhanced response gain to both relevant and irrelevant stimuli, and is accompanied by structural alterations in the white matter of the frontal and parietal lobe.
An influential conception of visual working memory is of a small number of discrete memory "slots", each storing an integrated representation of a single visual object, including all its component features. When a scene contains more objects than there are slots, visual attention controls which objects gain access to memory. A key prediction of such a model is that the absolute error in recalling multiple features of the same object will be correlated, because features belonging to an attended object are all stored, bound together. Here, we tested participants ability to reproduce from memory both the color and orientation of an object indicated by a location cue. We observed strong independence of errors between feature dimensions even for large memory arrays (6 items), inconsistent with an upper limit on the number of objects held in memory. Examining the pattern of responses in each dimension revealed a gaussian distribution of error centered on the target value that increased in width under higher memory loads. For large arrays, a subset of responses were not centered on the target but instead predominantly corresponded to mistakenly reproducing one of the other features held in memory. These misreporting responses again occurred independently in each feature dimension, consistent with misbinding due to errors in maintaining the binding information that assigns features to objects. The results support a shared-resource model of working memory, in which increasing memory load incrementally degrades storage of visual information, reducing the fidelity with which both object features and feature bindings are maintained.
Individuals with posterior cortical atrophy (PCA) report a host of unusual and poorly explained visual disturbances. This preliminary report describes a single patient (CRO), and documents and investigates abnormally prolonged colour afterimages (concurrent and prolonged perception of colours complimentary to the colour of an observed stimulus), perceived motion of static stimuli, and better reading of small than large letters. We also evaluate CROs visual and vestibular functions in an effort to understand the origin of her experience of room tilt illusion, a disturbing phenomenon not previously observed in individuals with cortical degenerative disease. These visual symptoms are set in the context of a 4-year longitudinal neuropsychological and neuroimaging investigation of CROs visual and other cognitive skills. We hypothesise that prolonged colour after-images are attributable to relative sparing of V1 inhibitory interneurons; perceived motion of static stimuli reflects weak magnocellular function; better reading of small than large letters indicates a reduced effective field of vision; and room tilt illusion effects are caused by disordered integration of visual and vestibular information. This study contributes to the growing characterisation of PCA whose atypical early visual symptoms are often heterogeneous and frequently under-recognised.
This series of articles for rehabilitation in practice aims to cover a knowledge element of the rehabilitation medicine curriculum. Nevertheless they are intended to be of interest to a multidisciplinary audience. The competency addressed in this article is The trainee consistently demonstrates a knowledge of the pathophysiology of various specific impairments including cognitive dysfunction including perception and management approaches for specific impairments including cognitive dysfunction including perception. The article focuses on hemispatial neglect as a common and difficult to manage problem in clinical practice.
The authors report here the case of a patient with severe deficits in arousal and sustained attention, associated with hemispatial neglect. These impairments were secondary to acute disseminated encephalomyelitis, with bilateral involvement of the medial nuclei and pulvinar of the thalamus. Treatment with the noradrenergic agonist guanfacine, previously used for attention deficits in attention deficit/hyperactivity disorder and stroke, was associated with a significant amelioration of both the spatial and sustained attention impairments in neglect. Guanfacine may prove to be a useful tool in the treatment of disorders of attention associated with neurological conditions.
Subliminal visual stimuli affect motor planning, but the size of such effects differs greatly between individuals. Here, we investigated whether such variation may be related to neurochemical differences between people. Cortical responsiveness is expected to be lower under the influence of more of the main inhibitory neurotransmitter, GABA. Thus, we hypothesized that, if an individual has more GABA in the supplementary motor area (SMA)--a region previously associated with automatic motor control--this would result in smaller subliminal effects. We measured the reversed masked prime--or negative compatibility--effect, and found that it correlated strongly with GABA concentration, measured with magnetic resonance spectroscopy. This occurred specifically in the SMA region, and not in other regions from which spectroscopy measurements were taken. We replicated these results in an independent cohort: more GABA in the SMA region is reliably associated with smaller effect size. These findings suggest that, across individuals, the responsiveness of subconscious motor mechanisms is related to GABA concentration in the SMA.
Attempts to improve cognitive function in patients with brain disorders have become the focus of intensive research efforts. A recent emerging trend is the use of so-called cognitive enhancers by healthy individuals. Here, we consider some of the effects - positive and negative - that current drugs have in neurological conditions and healthy people. We conclude that, to date, experimental and clinical studies have demonstrated relatively modest overall effects, most probably because of substantial variability in response both across and within individuals. We discuss biological factors that might account for such variability and highlight the need to improve testing methods and to extend our understanding of how drugs modulate specific cognitive processes at the systems or network level.
In the human brain, cognitive-control processes are generally considered distinct from the unconscious mechanisms elicited by subliminal priming. Here, we show that cognitive control engaged in situations of response conflict interacts with the negative (inhibitory) phase of subliminal priming. Thus, cognitive control may surprisingly share common processes with nonconscious brain mechanisms. In contrast, our findings reveal that subliminal inhibition does not, however, interact with control adaptation--the supposed modulation of current control settings by previous experience of conflict. Therefore, although influential models have grouped immediate cognitive control and control adaptation together as products of the same conflict detection and control network, their relationship to subliminal inhibition separates them. Overall, these results suggest that the important distinction lies not between cognitive or top-down processes on the one hand and nonconscious priming mechanisms on the other hand but between responsive (poststimulus) mechanisms that deal with sensorimotor activation after it has occurred and preparatory (prestimulus) mechanisms that are modulated before stimulus arrival.
Where we look is determined both by our current intentions and by the tendency of visually salient items to "catch our eye." After damage to parietal cortex, the normal process of directing attention is often profoundly impaired. Here, we tracked parietal patients eye movements during visual search to separately map impairments in goal-directed orienting to targets versus stimulus-driven gaze shifts to salient but task-irrelevant probes. Deficits in these two distinct types of attentional selection are shown to be identical in both magnitude and spatial distribution, consistent with damage to a "priority map" that integrates goal- and stimulus-related signals to select visual targets. When goal-relevant and visually salient items compete for attention, the outcome depends on a biased competition in which the priority of contralesional targets is undervalued. On the basis of these findings, we further demonstrate that parietal patients spatial bias (neglect) in goal-directed visual exploration can be corrected and even reversed by systematically manipulating the spatial distribution of stimulus salience in the visual array.
Constructional apraxia refers to the inability of patients to copy accurately drawings or three-dimensional constructions. It is a common disorder after right parietal stroke, often persisting after initial problems such as visuospatial neglect have resolved. However, there has been very little experimental investigation regarding mechanisms that might contribute to the syndrome. Here, we examined whether a key deficit might be failure to integrate visual information correctly from one fixation to the next. Specifically, we tested whether this deficit might concern remapping of spatial locations across saccades. Right-hemisphere stroke patients with constructional apraxia were compared to patients without constructional problems and neurologically healthy controls. Participants judged whether a pattern shifted position (spatial task) or changed in pattern (non-spatial task) across two saccades, compared to a control condition with an equivalent delay but without intervening eye movements. Patients with constructional apraxia were found to be significantly impaired in position judgements with intervening saccades, particularly when the first saccade of the sequence was to the right. The importance of these remapping deficits in constructional apraxia was confirmed through a highly significant correlation between saccade task performance and constructional impairment on standard neuropsychological tasks. A second study revealed that even single saccades to the right can impair constructional apraxia patients perception of location shifts. These data are consistent with the view that rightward eye movements result in loss of remembered spatial information from previous fixations, presumably due to constructional apraxia patients damage to the right-hemisphere regions involved in remapping locations across saccades. These findings provide the first evidence for a deficit in remapping visual information across saccades underlying right-hemisphere constructional apraxia.
It is widely accepted that regions within the dorsal medial frontal cortex are involved in the control of voluntary action. However, recent evidence suggests that a subset of these regions may also be important for unconscious and involuntary motor processes. Indeed, Sumner et al. (Neuron 54:697-711, 2007) showed that two patients with micro-lesions of the supplementary motor area (SMA) and supplementary eye field (SEF) demonstrated an absence of unconscious inhibition as evoked by masked-prime stimuli, while pre-SMA damage had no such effect. Here, we employ fMRI and a similar masked-prime task to test whether SMA and pre-SMA are similarly dissociated in healthy volunteers. Reaction times (RT) revealed that responses to compatible trials were slower than those to incompatible trials (negative compatibility effect, NCE), indicating automatic inhibition in every participant. BOLD signals in the SMA were modulated by prime compatibility, showing greater signal for compatible trials, but there was no change in pre-SMA. There was also no modulation in the hand motor cortex (HMC). These findings imply that the SMA is involved in automatic suppression of manual motor plans.
Motor sequence learning on the serial reaction time task involves the integration of response-, stimulus-, and effector-based information. Human primary motor cortex (M1) and the inferior parietal lobule (IPL) have been identified with supporting the learning of effector-dependent and -independent information, respectively. Current neurocognitive data are, however, exclusively based on learning complex sequence information via perceptual-motor responses. Here, we investigated the effects of continuous theta-burst transcranial magnetic stimulation (cTBS)-induced disruption of M1 and the angular gyrus (AG) of the IPL on learning a probabilistic sequence via sequential perceptual-motor responses (experiment 1) or covert orienting of visuospatial attention (experiment 2). Functional effects on manual sequence learning were evident during 75% of training trials in the cTBS M1 condition, whereas cTBS over the AG resulted in interference confined to a midpoint during the training phase. Posttraining direct (declarative) tests of sequence knowledge revealed that cTBS over M1 modulated the availability of newly acquired sequence knowledge, whereby sequence knowledge was implicit in the cTBS M1 condition but was available to conscious awareness in the cTBS AG and control conditions. In contrast, perceptual sequence learning was abolished in the perceptual cTBS AG condition, whereas learning was intact and available to conscious awareness in the cTBS M1 and control conditions. These results show that the right AG had a critical role in perceptual sequence learning, whereas M1 had a causal role in developing experience-dependent functional attributes relevant to conscious knowledge on manual but not perceptual sequence learning.
Extinction is a common consequence of unilateral brain injury: contralesional events can be perceived in isolation, yet are missed when presented concurrently with competing events on the ipsilesional side. This can arise crossmodally, where a contralateral touch is extinguished by an ipsilateral visual event. Recent studies showed that repositioning the hands in visible space, or making visual events more distant, can modulate such crossmodal extinction. Here, in a detailed single-case study, we implemented a novel spatial manipulation when assessing crossmodal extinction. This was designed not only to hold somatosensory inputs and hand/arm-posture constant, but also to hold (retinotopic) visual inputs constant, yet while still changing the spatial relationship of tactile and visual events in the external world. Our right hemisphere patient extinguished left-hand touches due to visual stimulation of the right visual field (RVF) when tested in the usual default posture with eyes/head directed straight ahead. But when her eyes/head were turned to the far left (and any visual events shifted along with this), such that the identical RVF retinal stimulation now fell at the same external location as the left-hand touch, crossmodal extinction was eliminated. Since only proprioceptive postural cues could signal this changed spatial relationship for the critical condition, our results show for the first time that such postural cues alone are sufficient to modulate crossmodal extinction. Identical somatosensory and retinal inputs can lead to severe crossmodal extinction, or none, depending on current posture.
The mechanisms underlying visual working memory have recently become controversial. One account proposes a small number of memory "slots," each capable of storing a single visual object with fixed precision. A contrary view holds that working memory is a shared resource, with no upper limit on the number of items stored; instead, the more items that are held in memory, the less precisely each can be recalled. Recent findings from a color report task have been taken as crucial new evidence in favor of the slot model. However, while this task has previously been thought of as a simple test of memory for color, here we show that performance also critically depends on memory for location. When errors in memory are considered for both color and location, performance on this task is in fact well explained by the resource model. These results demonstrate that visual working memory consists of a common resource distributed dynamically across the visual scene, with no need to invoke an upper limit on the number of objects represented.
A new study mapping the functional effects of brain lesions has revealed a surprising map of human intelligence, stimulating a re-evaluation of data from purely correlative methods such as functional magnetic resonance imaging.
Cowan & Rouder suggest that a modification to the four-slot model of visual working memory fits the available data better than our distributed resource model. However their comparisons of statistical fit are biased in favour of the slot model. Here we compare the predictions of the two models and present further evidence against the division of visual memory into slots.
Major advances during the past 50 years highlight the immense potential for restoration of function after neural injury, even in the damaged adult human brain. Yet, the translation of these advances into clinically useful treatments is painstakingly slow.
Optic ataxia (OA) is generally thought of as a disorder of visually guided reaching movements that cannot be explained by any simple deficit in visual or motor processing. In this paper we offer a new perspective on optic ataxia; we argue that the popular characterisation of this disorder is misleading and is unrepresentative of the pattern of reaching errors typically observed in OA patients. We begin our paper by reviewing recent neurophysiological, neuropsychological, and functional brain imaging studies that have led to the proposal that the medial parietal cortex in the vicinity of the parietal-occipital junction (POJ) - the key anatomical site associated with OA - represents reaching movements in eye-centred coordinates, and that this ability is impaired in optic ataxia. Our perspective stresses the importance of the POJ and superior parietal regions of the human PPC for representing reaching movements in both extrinsic (eye-centred) and intrinsic (postural) coordinates, and proposes that it is the ability to simultaneously represent multiple spatial locations that must be directly compared with one another that is impaired in non-foveal OA patients. In support of this idea we review recent fMRI and behavioural studies conducted by our group that have investigated the anatomical correlates of posturally guided movements, and the movements guided by postural cues in patients presenting with optic ataxia.
Recent models of human posterior parietal cortex (PPC) have variously emphasized its role in spatial perception, visuomotor control or directing attention. However, neuroimaging and lesion studies also suggest that the right PPC might play a special role in maintaining an alert state. Previously, assessments of right-hemisphere patients with hemispatial neglect have revealed significant overall deficits on vigilance tasks, but to date there has been no demonstration of a deterioration of performance over time--a vigilance decrement--considered by some to be a key index of a deficit in maintaining attention. Moreover, sustained attention deficits in neglect have not specifically been related to PPC lesions, and it remains unclear whether they interact with spatial impairments in this syndrome. Here we examined the ability of right-hemisphere patients with neglect to maintain attention, comparing them to stroke controls and healthy individuals. We found evidence of an overall deficit in sustaining attention associated with PPC lesions, even for a simple detection task with stimuli presented centrally. In a second experiment, we demonstrated a vigilance decrement in neglect patients specifically only when they were required to maintain attention to spatial locations, but not verbal material. Lesioned voxels in the right PPC spanning a region between the intraparietal sulcus and inferior parietal lobe were significantly associated with this deficit. Finally, we compared performance on a task that required attention to be maintained either to visual patterns or spatial locations, matched for task difficulty. Again, we found a vigilance decrement but only when attention had to be maintained on spatial information. We conclude that sustaining attention to spatial locations is a critical function of the human right PPC which needs to be incorporated into models of normal parietal function as well as those of the clinical syndrome of hemispatial neglect.
Simultanagnosia (resulting from occipito-parietal damage) is a profound visual deficit, which impairs the ability to perceive multiple items in a visual display, while preserving the ability to recognise single objects. Here we demonstrate in a patient presenting with Balints syndrome that this deficit may result from an extreme form of competition between objects which makes it difficult for attention to be disengaged from an object once it has been selected.
Current models of the visual pathways have difficulty incorporating the human inferior parietal lobe (IPL) into dorsal or ventral streams. Some recent proposals have attempted to integrate aspects of IPL function that were not hitherto dealt with well, such as differences between the left and right hemisphere and the role of the right IPL in responding to salient environmental events. However, we argue that these models also fail to capture adequately some important findings regarding the functions of the IPL. Here we critically appraise existing proposals regarding the functional architecture of the visual system, with special emphasis on the role of this region, particularly in the right hemisphere. We review evidence that shows the right IPL plays an important role in two different, but broadly complementary, aspects of attention: maintaining attentive control on current task goals as well as responding to salient new information or alerting stimuli in the environment. In our view, findings from functional imaging, electrophysiological and lesion studies are all consistent with the view that this region is part of a system that allows flexible reconfiguration of behaviour between these two alternative modes of operation. Damage to the right IPL leads to deficits in both maintaining attention and also responding to salient events, impairments that contribute to hemineglect, the classical syndrome that follows lesions of this region.
Making robust inferences about the functional neuroanatomy of the brain is critically dependent on experimental techniques that examine the consequences of focal loss of brain function. Unfortunately, the use of the most comprehensive such technique-lesion-function mapping-is complicated by the need for time-consuming and subjective manual delineation of the lesions, greatly limiting the practicability of the approach. Here we exploit a recently-described general measure of statistical anomaly, zeta, to devise a fully-automated, high-dimensional algorithm for identifying the parameters of lesions within a brain image given a reference set of normal brain images. We proceed to evaluate such an algorithm in the context of diffusion-weighted imaging of the commonest type of lesion used in neuroanatomical research: ischaemic damage. Summary performance metrics exceed those previously published for diffusion-weighted imaging and approach the current gold standard-manual segmentation-sufficiently closely for fully-automated lesion-mapping studies to become a possibility. We apply the new method to 435 unselected images of patients with ischaemic stroke to derive a probabilistic map of the pattern of damage in lesions involving the occipital lobe, demonstrating the variation of anatomical resolvability of occipital areas so as to guide future lesion-function studies of the region.
Attention modulates the availability of sensory information to conscious perception. In particular, there is evidence of pathological, spatial constriction of the effective field of vision in patients with right hemisphere damage when a central task exhausts available attentional capacity. In the current study we first examined whether this constriction might be modulated across both space and time in right hemisphere stroke patients without neglect. Then we tested healthy elderly people to determine whether non-pathological ageing also leads to spatiotemporal impairments of vision under conditions of high attention load.
Recent studies have demonstrated that memory performance can be enhanced by a cue which indicates the item most likely to be subsequently probed, even when that cue is delivered seconds after a stimulus array is extinguished. Although such retro-cuing has attracted considerable interest, the mechanisms underlying it remain unclear. Here, we tested the hypothesis that retro-cues might protect an item from degradation over time. We employed two techniques that previously have not been deployed in retro-cuing tasks. First, we used a sensitive, continuous scale for reporting the orientation of a memorized item, rather than binary measures (change or no change) typically used in previous studies. Second, to investigate the stability of memory across time, we also systematically varied the duration between the retro-cue and report. Although accuracy of reporting uncued objects rapidly declined over short intervals, retro-cued items were significantly more stable, showing negligible decline in accuracy across time and protection from forgetting. Retro-cuing an objects color was just as advantageous as spatial retro-cues. These findings demonstrate that during maintenance, even when items are no longer visible, attention resources can be selectively redeployed to protect the accuracy with which a cued item can be recalled over time, but with a corresponding cost in recall for uncued items.
Although we frequently take advantage of memory for objects locations in everyday life, understanding how an objects identity is bound correctly to its location remains unclear. Here we examine how information about object identity, location and crucially object-location associations are differentially susceptible to forgetting, over variable retention intervals and memory load. In our task, participants relocated objects to their remembered locations using a touchscreen. When participants mislocalized objects, their reports were clustered around the locations of other objects in the array, rather than occurring randomly. These swap errors could not be attributed to simple failure to remember either the identity or location of the objects, but rather appeared to arise from failure to bind object identity and location in memory. Moreover, such binding failures significantly contributed to decline in localization performance over retention time. We conclude that when objects are forgotten they do not disappear completely from memory, but rather it is the links between identity and location that are prone to be broken over time.
Active exploration of the visual world depends on sequential shifts of gaze that bring prioritized regions of a scene into central vision. The efficiency of this system is commonly attributed to a mechanism of "inhibition of return" (IOR) that discourages re-examination of previously-visited locations. Such a process is fundamental to computational models of attentional selection and paralleled by neurophysiological observations of inhibition of target-related activity in visuomotor areas. However, studies examining eye movements in naturalistic visual scenes appear to contradict the hypothesis that IOR promotes exploration. Instead, these reports reveal a surprisingly strong tendency to shift gaze back to the previously fixated location, suggesting that refixations might even be facilitated under natural conditions. Here we resolve this apparent contradiction, based on a probabilistic analysis of gaze patterns recorded during both free-viewing and search of naturalistic scenes. By simulating saccadic selection based on instantaneous influences alone, we show that the observed frequency of return saccades is in fact substantially less than predicted for a memoryless system, demonstrating that refixation is actively inhibited under natural viewing conditions. Furthermore, these observations reveal that gaze history significantly influences the way in which natural scenes are explored, contrary to accounts that suggest visual search has no memory.
Hemispatial neglect following right-hemisphere stroke is a common and disabling disorder, for which there is currently no effective pharmacological treatment. Dopamine agonists have been shown to play a role in selective attention and working memory, two core cognitive components of neglect. Here, we investigated whether the dopamine agonist rotigotine would have a beneficial effect on hemispatial neglect in stroke patients. A double-blind, randomized, placebo-controlled ABA design was used, in which each patient was assessed for 20 testing sessions, in three phases: pretreatment (Phase A1), on transdermal rotigotine for 7-11 days (Phase B) and post-treatment (Phase A2), with the exact duration of each phase randomized within limits. Outcome measures included performance on cancellation (visual search), line bisection, visual working memory, selective attention and sustained attention tasks, as well as measures of motor control. Sixteen right-hemisphere stroke patients were recruited, all of whom completed the trial. Performance on the Mesulam shape cancellation task improved significantly while on rotigotine, with the number of targets found on the left side increasing by 12.8% (P?=?0.012) on treatment and spatial bias reducing by 8.1% (P?=?0.016). This improvement in visual search was associated with an enhancement in selective attention but not on our measures of working memory or sustained attention. The positive effect of rotigotine on visual search was not associated with the degree of preservation of prefrontal cortex and occurred even in patients with significant prefrontal involvement. Rotigotine was not associated with any significant improvement in motor performance. This proof-of-concept study suggests a beneficial role of dopaminergic modulation on visual search and selective attention in patients with hemispatial neglect following stroke.
Apathy is a complex, behavioural disorder associated with reduced spontaneous initiation of actions. Although present in mild forms in some healthy people, it is a pathological state in conditions such as Alzheimers and Parkinsons disease where it can have profoundly devastating effects. Understanding the mechanisms underlying apathy is therefore of urgent concern but this has proven difficult because widespread brain changes in neurodegenerative diseases make interpretation difficult and there is no good animal model. Here we present a very rare case with profound apathy following bilateral, focal lesions of the basal ganglia, with globus pallidus regions that connect with orbitofrontal (OFC) and ventromedial prefrontal cortex (VMPFC) particularly affected. Using two measures of oculomotor decision-making we show that apathy in this individual was associated with reward insensitivity. However, reward sensitivity could be established partially with levodopa and more effectively with a dopamine receptor agonist. Concomitantly, there was an improvement in the patients clinical state, with reduced apathy, greater motivation and increased social interactions. These findings provide a model system to study a key neuropsychiatric disorder. They demonstrate that reward insensitivity associated with basal ganglia dysfunction might be an important component of apathy that can be reversed by dopaminergic modulation.
Visual working memory (VWM) is the facility to hold in mind visual information for brief periods of time. Developmental studies have suggested an increase during childhood in the maximum number of complete items that can simultaneously be stored in VWM. Here, we exploit a recent theoretical and empirical innovation to investigate instead the precision with which items are stored in VWM, where precision is a continuous measure reflecting VWM resolution. Ninety boys aged 7 to 13 years completed one-item and three-item VWM tasks in which stimuli were coloured bars varying in orientation. On each trial, participants used a rotating dial to reproduce the probed stimulus from memory. Results show linear age-related improvement in recall precision for both one-item and three-item VWM tasks. However, even the youngest age group stored a significant amount of information about all three items on the difficult 3-item VWM task. Importantly, the development of VWM precision was not accounted for by development on a sensorimotor control task. Whereas storage of a single complete item was previously thought to be well within the capacity limitations of the current age range, these results suggest protracted development during childhood and early adolescence in the resolution with which single and multiple items are stored in VWM. Probabilistic modelling of response distribution data suggests that improvement in VWM performance is attributable to a specific decrease in variability of stored feature representations, rather than to a decrease in misbinding or random noise. As such, we highlight a novel, potentially developmentally plausible mechanism that may underlie developmental improvement in VWM performance, independent of any alterations in the maximum number of complete items which can be stored.
Effective behavioral therapies exist for patients with brain injury. The main issue is one of access. Can the internet be used as a resource so that suitable patients can build up enough practice to improve? We tested this hypothesis using a web-based application for patients with a right-sided hemianopia causing slow text reading. We studied 33 patients aged 26-81 years who fitted the entry criteria and accessed the therapy website between May 2010 and December 2011, in a longitudinal cohort study. The therapy consisted of reading animated, laterally scrolling text whose content and form was selected by the patients. Reading speeds on static text (main outcome) were assessed after every 5-h period of practice had been accrued. Statistical analysis was carried out using a repeated measures ANOVA. Read-Right therapy produced significant improvements in text reading speeds at all time points with a clear dose effect: 10 % at 5 h, 20 % at 10 h, 39 % at 15 h and 46 % at 20 h. Sub-analyses demonstrated that this was unlikely to be due to either multiple exposure to the testing materials (familiarity) or to the simple passage of time. This is the first example of a clinically proven therapy being delivered effectively to stroke patients over the internet. As therapists time is more limited than patients capacity to improve, carefully designed, web-based resources like Read-Right represent a realistic way of delivering a sufficient therapy dose to patients so they can obtain clinically meaningful improvements.
Impulsivity is a feature of many brain disorders. Although often defined as the predisposition to act with an inadequate degree of deliberation, forethought, or control, it has proven difficult to measure. This may in part be due to the fact that it is a multifaceted construct, with impulsive decisions potentially arising as a result of a number of underlying mechanisms. Indeed, a "functional" degree of impulsivity may even promote effective behavior in healthy participants in a way that can be advantageous under certain circumstances. Although many tasks have been developed to study impulsivity, few examine decisions made rapidly, for time-sensitive rewards. In the current study we examine behavior in 59 adults on a manual "Traffic Light" task which requires participants to take risks under time pressure, if they are to maximize reward. We show that behavioral variables that index rapid anticipatory responding in this paradigm are correlated with one, specific self-report measure of impulsivity: "lack of premeditation" on the UPPS Impulsive Behavior Scale. Participants who scored more highly on this subscale performed better on the task. Moreover, anticipatory behavior reduced significantly with age (18-79?years), an effect that continued to be upheld after correction for potential age differences in the ability to judge the timing of responses. Based on these findings, we argue that the Traffic Light task provides a parametric method to study one aspect of impulsivity in health and disease: namely, rapid decision-making in pursuit of risky, time-sensitive rewards.
The aim of this study was to test the effect and specificity of a novel, compensatory eye movement training therapy designed to improve visual search performance in patients with homonymous visual field defects.
Although executive control and automatic behavior have often been considered separate and distinct processes, there is strong emerging and convergent evidence that they may in fact be intricately interlinked. In this review, we draw together evidence showing that visual stimuli cause automatic and unconscious motor activation, and how this in turn has implications for executive control. We discuss object affordances, alien limb syndrome, the visual grasp reflex, subliminal priming, and subliminal triggering of attentional orienting. Consideration of these findings suggests automatic motor activation might form an intrinsic part of all behavior, rather than being categorically different from voluntary actions.
Reports have conflicted about the possible special role of location in visual working memory (WM). One important question is: Do we maintain the locations of objects in WM even when they are irrelevant to the task at hand? Here we used a continuous response scale to study the types of reporting errors that participants make when objects are presented at the same or at different locations in space. When several objects successively shared the same location, participants exhibited a higher tendency to report features of the wrong object in memory; that is, they responded with features that belonged to objects retained in memory but not probed at retrieval. On the other hand, a similar effect was not observed when objects shared a nonspatial feature, such as color. Furthermore, the effect of location on reporting errors was present even when its manipulation was orthogonal to the task at hand. These findings are consistent with the view that binding together different nonspatial features of an object in memory might be mediated through an objects location. Hence, spatial location may have a privileged role in WM. The relevance of these findings to conceptual models, as well as to neural accounts of visual WM, is discussed.
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