{ "items": [ "\n\n
The right inferior frontal gyrus (rIFG) and the presupplementary motor area (pre-SMA) have been identified with cognitive control-the top-down influence on other brain areas when nonroutine behavior is required. It has been argued that they \"inhibit\" habitual motor responses when environmental changes mean a different response should be made. However, whether such \"inhibition\" can be equated with inhibitory physiological interactions has been unclear, as has the areas' relationship with each other and the anatomical routes by which they influence movement execution. Paired-pulse transcranial magnetic stimulation (ppTMS) was applied over rIFG and primary motor cortex (M1) or over pre-SMA and M1 to measure their interactions, at a subsecond scale, during either inhibition and reprogramming of actions or during routine action selection. Distinct patterns of functional interaction between pre-SMA and M1 and between rIFG and M1 were found that were specific to action reprogramming trials; at a physiological level, direct influences of pre-SMA and rIFG on M1 were predominantly facilitatory and inhibitory, respectively. In a subsequent experiment, it was shown that the rIFG's inhibitory influence was dependent on pre-SMA. A third experiment showed that pre-SMA and rIFG influenced M1 at two time scales. By regressing white matter fractional anisotropy from diffusion-weighted magnetic resonance images against TMS-measured functional connectivity, it was shown that short-latency (6 ms) and longer latency (12 ms) influences were mediated by cortico-cortical and subcortical pathways, respectively, with the latter passing close to the subthalamic nucleus.
\n \n\n \n \nVentral premotor cortex (PMv) is widely accepted to exert an important influence over primary motor cortex (M1) when hand movements are made. Although study of these interactions has typically focused on their excitatory nature, given its strong connections with both ventral and opercular frontal regions, one feature of the influence of PMv over M1 may be inhibitory. Paired-pulse transcranial magnetic stimulation (ppTMS) was used to examine functional interactions between human PMv and M1 during the selection and reprogramming of a naturalistic goal-directed action. One of two cylinders was illuminated on each trial. It was then grasped and picked up. On some trials, however, subjects had to reprogram the action as the illuminated cylinder was switched off and the other illuminated simultaneously with reach initiation. At a neurophysiological level, the PMv paired-pulse effect (PPE) on M1 corticospinal activity was facilitatory after the initial target presentation and during movement initiation. When reprogramming was required, however, the PPE became strongly inhibitory. This context-dependent change from facilitation to inhibition occurred within 75 ms of the change of target. Behaviorally, PMv-M1 ppTMS disrupted reprogramming. Diffusion-weighted magnetic resonance image scans were taken of each subject. Intersubject differences in the facilitation-inhibition contrast of PMv-M1 interactions were correlated with fractional anisotropy of white-matter in ventral prefrontal, premotor, and intraparietal brain areas. These results suggest that a network of brain areas centered on PMv inhibits M1 corticospinal activity associated with undesired movements when action plans change.
\n \n\n \n \nCognitive flexibility is known to depend on the striatum. However, the striatum does not act in isolation to bias cognitive flexibility. In particular, cognitive flexibility also implicates the frontal cortex. Here we tested the hypothesis that the human frontal cortex controls cognitive flexibility by regulating striatal function via topographically specific frontostriatal connections. To this end, we exploited a repetitive transcranial magnetic stimulation (TMS) protocol over frontal cortex that is known to increase dopamine release in the striatum. This intervention was combined with functional magnetic resonance imaging to determine the functional and topographic specificity of its consequences at the whole brain level. Participants were scanned both before and after off-line TMS while performing a cognitive switching task that is known to depend on a specific striatal substructure, the putamen. Frontal stimulation perturbed task-specific functional signals in the putamen, while reducing fronto-striatal functional connectivity. There were no such effects of TMS over the medial parietal cortex. These data strengthen the hypothesis that cognitive flexibility involves topographic frontal control of striatal function.
\n \n\n \n \nIt has been suggested that the frontal operculum (fO) is a key node in a network for exerting control over cognitive processes. How it exerts this influence, however, has been unclear. Here, using the complementary approaches of functional MRI and transcranial magnetic stimulation, we have shown that the fO regulates increases and decreases of activity in multiple occipitotemporal cortical areas when task performance depended on directing attention to different classes of stimuli held in memory. Only one region, the fO, was significantly more active when subjects selectively attended to a single stimulus so that it determined task performance. The stimuli that guided task performance could belong to three categories--houses, body parts, and faces--associated with three occipitotemporal regions. On each trial, the pattern of functional correlation between the fO and the three occipitotemporal regions became either positive or negative, depending on which stimulus was to be attended and which ignored. Activation of the fO preceded both activity increases and decreases in the occipitotemporal cortex. The causal dependency of the distributed occipitotemporal pattern of activity increases and decreases on the fO was demonstrated by showing that transcranial magnetic stimulation-mediated interference of the fO diminished top-down selective attentional modulation in the occipitotemporal cortex, but it did not alter bottom-up activation of the same areas to the same stimuli when they were presented in isolation. The fO's prominence in cognitive control may stem from a role in regulating the level of activity of representations in posterior brain areas that are relevant or irrelevant, respectively, for response selection.
\n \n\n \n \nBehavioral economic studies involving limited numbers of choices have provided key insights into neural decision-making mechanisms. By contrast, animals' foraging choices arise in the context of sequences of encounters with prey or food. On each encounter, the animal chooses whether to engage or, if the environment is sufficiently rich, to search elsewhere. The cost of foraging is also critical. We demonstrate that humans can alternate between two modes of choice, comparative decision-making and foraging, depending on distinct neural mechanisms in ventromedial prefrontal cortex (vmPFC) and anterior cingulate cortex (ACC) using distinct reference frames; in ACC, choice variables are represented in invariant reference to foraging or searching for alternatives. Whereas vmPFC encodes values of specific well-defined options, ACC encodes the average value of the foraging environment and cost of foraging.
\n \n\n \n \nFunctional magnetic resonance imaging was used to measure activity in three frontal cortical areas, the lateral orbitofrontal cortex (lOFC), medial orbitofrontal cortex (mOFC)/ventromedial frontal cortex (vmPFC), and anterior cingulate cortex (ACC), when expectations about type of reward, and not just reward presence or absence, could be learned. Two groups of human subjects learned 12 stimulus-response pairings. In one group (Consistent), correct performances of a given pairing were always reinforced with a specific reward outcome, whereas in the other group (Inconsistent), correct performances were reinforced with randomly selected rewards. The mOFC/vmPFC and lOFC were not distinguished by simple differences in relative preference for positive and negative outcomes. Instead lOFC activity reflected updating of reward-related associations specific to reward type; lOFC was active whenever informative outcomes allowed updating of reward-related associations, regardless of whether the outcomes were positive or negative, and the effects were greater when consistent stimulus-outcome and response-outcome mappings were present. A psychophysiological interaction analysis demonstrated changed coupling between lOFC and brain areas for visual object representation, such as perirhinal cortex, and reward-guided learning, such as the amygdala, ventral striatum, and habenula/mediodorsal thalamus. In contrast, mOFC/vmPFC activity reflected expected values of outcomes and occurrence of positive outcomes, regardless of consistency of outcome mappings. The third frontal cortical region, the ACC, reflected the use of reward type information to guide response selection. ACC activity reflected the probability of selecting the correct response, was greater when consistent outcome mappings were present, and was related to individual differences in propensity to select the correct response.
\n \n\n \n \nDiffusion imaging can be used to estimate the routes taken by fiber pathways connecting different regions of the living brain. This approach has already supplied novel insights into in vivo human brain anatomy. For example, by detecting where connection patterns change, one can define anatomical borders between cortical regions or subcortical nuclei in the living human brain for the first time. Because diffusion tractography is a relatively new technique, however, it is important to assess its validity critically. We discuss the degree to which diffusion tractography meets the requirements of a technique to assess structural connectivity and how its results compare to those from the gold-standard tract tracing methods in nonhuman animals. We conclude that although tractography offers novel opportunities it also raises significant challenges to be addressed by further validation studies to define precisely the limitations and scope of this exciting new technique.
\n \n\n \n \nThe medial frontal cortex (MFC) has been identified with voluntary action selection. Recent evidence suggests that there are three principal ways in which the MFC is an essential part of the neural circuit for voluntary action selection. First, the MFC represents the reinforcement values of actions and is concerned with the updating of those action values. Because it is particularly concerned with the rate at which action values should be updated, it mediates the influence that the past reinforcement history has over the next choice that is made and it may determine the learning rate. The MFC's representation of action value does not just reflect the potential reward associations of an action but instead represents both the reward and effort costs that are intrinsic to the action. Second, the MFC is important when an exploratory action is generated in order to obtain more information about action values and the environment. Third, the MFC is critical when conflicting information in the immediate environment instructs more than one possible response. In such situations the MFC exerts an influence over how actions will be chosen by other motor regions of the brain.
\n \n\n \n \nTranscranial magnetic stimulation (TMS) is one of the most recent techniques to have been used in investigations of the parietal cortex but already a number of studies have employed it as a tool in investigations of attentional and sensorimotor processes in the human parietal cortices. The high temporal resolution of TMS has proved to be a particular strength of the technique and the experiments have led to hypotheses about when circumscribed regions of parietal cortex are critical for specific attentional and sensorimotor processes. A consistent theme that runs through many reports is that of a critical contribution of parietal areas when attention or movements are re-directed and representations for attention or action must be updated.
\n \n\n \n \nCategory-related brain activations have been reported in the posterior fusiform gyri when people view pictures of tools and animals, but only a single study has observed this pattern when the stimuli were words, rather than pictures. Here we replicate these category effects with words and provide evidence that distinctive patterns of activation are task specific. The results suggest that category-related activation in the posterior fusiform gyri can be driven either \"bottom-up\" by visual processing of images or \"top-down\" by word processing.
\n \n\n \n \nFunctional and structural neuroimaging of the human cingulate cortex has identified this region with emotion and social cognition and suggested that cingulate pathology may be associated with emotional and social behavioural disturbances. The importance of the cingulate cortex for emotion and social behaviour, however, has not been clear from lesion studies. Bilateral lesions in the cingulate cortex were made in three macaques and their social interactions were compared with those of controls. Subsequently, cingulate lesions were made in the three controls and their behaviour was compared before and after surgery. Cingulate lesions were associated with decreases in social interactions, time spent in proximity with other individuals, and vocalisations but an increase in manipulation of an inanimate object. The results are consistent with a cingulate role in social behaviour and emotion.
\n \n\n \n \nPET was used to study cerebral dominance for the selection of action. In one condition the subjects moved one of two fingers depending on the cue presented (choice reaction time), and in another they moved the same finger whatever the cue (simple reaction time). There was also a baseline condition in which cues were shown but no movements were made. A conjunction analysis was performed to reveal those areas which were more activated for the choice versus simple reaction time, irrespective of whether the right or left hand was used. The activations were in prefrontal, premotor and intraparietal areas, and they were all in the left hemisphere. Thus, while there were activations in the right hemisphere for the choice versus simple reaction time task when the subjects used their left (contralateral) hand, there were activations in left prefrontal, premotor and parietal areas whether the right (contralateral) or left (ipsilateral) hands were used. It is argued that the results suggest that the left hemisphere is dominant not only for speech but also for action in general.
\n \n\n \n \n