Distinct neural processes support post-success and post-error slowing in the stop signal task.

Executive control requires behavioral adaptation to environmental contingencies. In the stop signal task (SST), participants exhibit slower go trial reaction time (RT) following a stop trial, whether or not they successfully interrupt the motor response. In previous fMRI studies, we demonstrated activation of the right-hemispheric ventrolateral prefrontal cortex, in the area of inferior frontal gyrus, pars opercularis (IFGpo) and anterior insula (AI), during post-error slowing (PES). However, in similar analyses we were not able to identify regional activities during post-success slowing (PSS). Here, we revisited this issue in a larger sample of participants (n=100) each performing the SST for 40 min during fMRI. We replicated IFGpo/AI activation to PES (p≤0.05, FWE corrected). Further, PSS engages decreased activation in a number of cortical regions including the left inferior frontal cortex (IFC; p≤0.05, FWE corrected). We employed Granger causality mapping to identify areas that provide inputs each to the right IFGpo/AI and left IFC, and computed single-trial amplitude (STA) of stop trials of these input regions as well as the STA of post-stop trials of the right IFGpo/AI and left IFC. The STAs of the right inferior precentral sulcus and supplementary motor area (SMA) and right IFGpo/AI were positively correlated and the STAs of the left SMA and left IFC were positively correlated (slope>0, p's≤0.01, one-sample t test), linking regional responses during stop success and error trials to those during PSS and PES. These findings suggest distinct neural mechanisms to support PSS and PES.

Primary somatosensory cortex necessary for the perception of weight from other people's action: A continuous theta-burst TMS experiment.

The presence of a network of areas in the parietal and premotor cortices, which are active both during action execution and observation, suggests that we might understand the actions of other people by activating those motor programs for making similar actions. Although neurophysiological and imaging studies show an involvement of the somatosensory cortex (SI) during action observation and execution, it is unclear whether SI is essential for understanding the somatosensory aspects of observed actions. To address this issue, we used off-line transcranial magnetic continuous theta-burst stimulation (cTBS) just before a weight judgment task. Participants observed the right hand of an actor lifting a box and estimated its relative weight. In counterbalanced sessions, we delivered sham and active cTBS over the hand region of the left SI and, to test anatomical specificity, over the left motor cortex (M1) and the left superior parietal lobule (SPL). Active cTBS over SI, but not over M1 or SPL, impaired task performance relative to sham cTBS. Moreover, active cTBS delivered over SI just before participants were asked to evaluate the weight of a bouncing ball did not alter performance compared to sham cTBS. These findings indicate that SI is critical for extracting somatosensory features (heavy/light) from observed action kinematics and suggest a prominent role of SI in action understanding.

Primary somatosensory contribution to action observation brain activity-combining fMRI and cTBS.

Traditionally the mirror neuron system (MNS) only includes premotor and posterior parietal cortices. However, somatosensory cortices, BA1/2 in particular, are also activated during action execution and observation. Here, we examine whether BA1/2 and the parietofrontal MNS integrate information by using functional magnetic resonance imaging (fMRI)-guided continuous theta-burst stimulation (cTBS) to perturb BA1/2. Measuring brain activity using fMRI while participants are under the influence of cTBS shows local cTBS effects in BA1/2 varied, with some participants showing decreases and others increases in the BOLD response to viewing actions vs control stimuli. We show how measuring cTBS effects using fMRI can harness this variance using a whole-brain regression. This analysis identifies brain regions exchanging action-specific information with BA1/2 by mapping voxels away from the coil with cTBS-induced, action-observation-specific BOLD contrast changes that mirror those under the coil. This reveals BA1/2 exchanges action-specific information with premotor, posterior parietal and temporal nodes of the MNS during action observation. Although anatomical connections between BA1/2 and these regions are well known, this is the first demonstration that these connections carry action-specific signals during observation and hence, that BA1/2 plays a causal role in the human MNS.

Is Brain Activity during Action Observation Modulated by the Perceived Fairness of the Actor?

Perceiving other people's actions triggers activity in premotor and parietal areas, brain areas also involved in executing and sensing our own actions. Paralleling this phenomenon, observing emotional states (including pain) in others is associated with activity in the same brain areas as activated when experiencing similar emotions directly. This emotion perception associated activity has been shown to be affected by the perceived fairness of the actor, and in-group membership more generally. Here, we examine whether action observation associated brain activity is also affected by the perceived social fairness of the actors. Perceived fairness was manipulated using an alternating iterated Prisoner's Dilemma game between the participant and two confederates, one of whom played fairly and the other unfairly. During fMRI scanning the participants watched movies of the confederates performing object-directed hand actions, and then performed hand actions themselves. Mass-univariate analysis showed that observing the actions triggered robust activation in regions associated with action execution, but failed to identify a strong modulation of this activation based on perceived fairness. Multivariate pattern analysis, however, identified clusters potentially carrying information about the perceived fairness of the actor in the middle temporal gyrus, left postcentral gyrus, right inferior parietal lobule, right middle cingulate cortex, right angular gyrus, and right superioroccipital gyrus. Despite being identified by a whole-brain searchlight analysis (and so without anatomical restriction), these clusters fall into areas frequently associated with action observation. We conclude that brain activity during action observation may be modulated by perceived fairness, but such modulation is subtle; robust activity is associated with observing the actions of both fair and unfair individuals.

cTBS delivered to the left somatosensory cortex changes its functional connectivity during rest.

The primary somatosensory cortex (SI) plays a critical role in somatosensation as well as in action performance and social cognition. Although the SI has been a major target of experimental and clinical research using non-invasive transcranial magnetic stimulation (TMS), to date information on the effect of TMS over the SI on its resting-state functional connectivity is very scant. Here, we explored whether continuous theta burst stimulation (cTBS), a repetitive TMS protocol, administered over the SI can change the functional connectivity of the brain at rest, as measured using resting-state functional magnetic resonance imaging (rs-fMRI). In a randomized order on two different days we administered active TMS or sham TMS over the left SI. TMS was delivered off-line before scanning by means of cTBS. The target area was selected previously and individually for each subject as the part of the SI activated both when the participant executes and observes actions. Three analytical approaches, both theory driven (partial correlations and seed based whole brain regression) and more data driven (Independent Component Analysis), indicated a reduction in functional connectivity between the stimulated part of the SI and several brain regions functionally associated with the SI including the dorsal premotor cortex, the cerebellum, basal ganglia, and anterior cingulate cortex. These findings highlight the impact of cTBS delivered over the SI on its functional connectivity at rest. Our data may have implications for experimental and therapeutic applications of cTBS over the SI.

Weight dependent modulation of motor resonance induced by weight estimation during observation of partially occluded lifting actions.

Seeing others performing an action induces the observers' motor cortex to "resonate" with the observed action. Transcranial magnetic stimulation (TMS) studies suggest that such motor resonance reflects the encoding of various motor features of the observed action, including the apparent motor effort. However, it is unclear whether such encoding requires direct observation or whether force requirements can be inferred when the moving body part is partially occluded. To address this issue, we presented participants with videos of a right hand lifting a box of three different weights and asked them to estimate its weight. During each trial we delivered one transcranial magnetic stimulation (TMS) pulse over the left primary motor cortex of the observer and recorded the motor evoked potentials (MEPs) from three muscles of the right hand (first dorsal interosseous, FDI, abductor digiti minimi, ADM, and brachioradialis, BR). Importantly, because the hand shown in the videos was hidden behind a screen, only the contractions in the actor's BR muscle under the bare skin were observable during the entire videos, while the contractions in the actor's FDI and ADM muscles were hidden during the grasp and actual lift. The amplitudes of the MEPs recorded from the BR (observable) and FDI (hidden) muscle increased with the weight of the box. These findings indicate that the modulation of motor excitability induced by action observation extends to the cortical representation of muscles with contractions that could not be observed. Thus, motor resonance appears to reflect force requirements of observed lifting actions even when the moving body part is occluded from view.

The impact of certain methodological choices on multivariate analysis of fMRI data with support vector machines.

Multivoxel pattern analysis of functional magnetic resonance imaging (fMRI) data is continuing to increase in popularity. Like all fMRI analyses, these analyses require extensive data processing and methodological choices, but the impact of these decisions on the final results is not always known. This study explores the impact of four methodological choices on analysis outcomes and introduces the technique of partitioning on random runs for characterizing temporal dependencies and evaluating partitioning methods. The analyses were performed on two fMRI data sets, which were repeatedly analyzed with support vector machines, varying the method of temporal compression, smoothing, voxel-wise detrending, and partitioning into training and testing sets. Smoothing sometimes slightly increased classification accuracy. Partitioning other than on the runs increased classification accuracy, and the random runs technique allowed us to attribute this improvement to the increased amount of training data, rather than to bias. The impact of the temporal compression and detrending methods varied so strongly with data set that general recommendations could not be drawn. These interactions suggest that, rather than searching for a universally superior set of methodological choices, researchers must carefully consider each choice in the context of each experiment.

Individual differences in the attentional blink. The important role of irrelevant information.

A well-established phenomenon in the study of attention is the attentional blink (AB): a deficit in reporting the second of two targets when it occurs 200-500 ms after the first. Although the effect has been shown to be robust in a wide variety of task conditions, we recently reported that some individuals show little or no AB, and presented psychophysiological evidence that target processing differs in nonblinkers (who do not show an AB) and blinkers (who do show an AB). Here we present evidence that the level of distractor processing and subsequent interference with target identification processes also differs between the two groups. In one task, two masked targets were centrally presented at varying temporal intervals, with or without additional distractors. In a second task, the masked targets were presented eccentrically, with or without the presence of a central sequential stream of the task-irrelevant distractors. In both cases, the presence of distractors led to an increased AB magnitude in blinkers, whereas performance for nonblinkers remained relatively unaffected. The results thus support the hypothesis that nonblinkers are more efficient in ignoring irrelevant information than blinkers.