Pascal Molenberghs

Objectives: Researchers have begun delivering mindfulness and aerobic exercise training concurrently on the premise that a combination intervention will yield salutary outcomes over and above each intervention alone. An estimate of the effect of combination training on chronic psychosocial stress in a nonclinical population has not been established. The objective of this trial was therefore to assess the preliminary efficacy of combination training for reducing chronic psychosocial stress in a healthy sample, and to establish assurance of protocol feasibility in preparation of a definitive RCT. Methods: 24 participants were nonrandomized into a single-arm trial and subjected to 16 weeks of concurrent mindfulness psychoeducation and aerobic exercise training. Within-group changes were assessed for chronic psychosocial stress, mindfulness, emotion regulation, and cardiorespiratory fitness. Feasibility criteria were also collected and evaluated. Primary analyses are based on 17 partici...

In this extended commentary, we critically evaluate a sample of empirical studies conducted in organizational neuroscience with respect to present-day best practices (i.e., using parameter estimation based on effect size and confidence intervals, and with consideration of the implications of inadequate research and reporting practices), with the primary goal of developing new recommendations to strengthen post-publication peer review. Our commentary yields several research themes we propose scholars must engage with in future critical evaluations of the literature, particularly with respect to seminal works in the field. This includes addressing the implications of a lack of transparency in reporting; interpretation of findings through extreme reverse inference; drawing conclusions by eyeballing effect size differences in lieu of statistical testing; and the use of analyses that preclude generalizability beyond the sample, among others. Because we endorse the view that organizationa...

The role of the orbitofrontal cortex (OFC) in moral decision-making is well established. However, OFC activity is highly context dependent. It is affected by the extent to which choices are morally justified and whom they concern. In the current study, we specifically focus on contextual factors and investigate the differential role of the OFC during justified and unjustified violence towards ingroup versus outgroup members. Muslims were chosen as the outgroup, as they are currently stereotypically seen as an outgroup and a potential threat by some Non-Muslims. Importantly, we also introduce a context where participants are the actual agents responsible for doing harm. During fMRI scanning, Non-Muslim participants had to decide to either shoot a Non-Muslim (i.e., ingroup member) or Muslim (outgroup member) depending on whether they believed the target was holding a gun or an object. Neuroimaging results showed increased activation in the lateral OFC (lOFC) in the three contrasts tha...

Morality is an important social construct necessary for understanding what is right and wrong. Neuroimaging studies investigating morality have used a wide variety of paradigms and implicated many different brain areas. Yet, it remains unclear whether differences amongst morality tasks are the cause for such heterogeneous findings. Therefore, in the present study, a series of activation likelihood estimation (ALE) meta-analyses were conducted on 123 datasets (inclusive of 1963 participants) to address this question. The ALE meta-analyses revealed a series of common brain areas associated with all moral tasks, including medial prefrontal cortex, lateral orbitofrontal cortex, amygdala, temporoparietal junction, and precuneus. However, individual and contrast analyses also revealed unique networks associated with each moral modality, suggesting that different moral tasks recruit specialised brain regions.

It is not uncommon for people to openly admit to pirating information from the internet despite the known legal consequences. Those same people are often less inclined to steal the same physical item from a shop. This raises the question, why do people have fewer reservations with stealing intangible items compared to tangible? Using questionnaires and fMRI we provide evidence across three studies as to the differences between tangible and intangible theft. In a questionnaire (Study 1), participants revealed that across different conditions they were more willing to steal intangible compared to tangible goods. Study 2a used fMRI to reveal that a network involved in imagining objects was more active when participants were representing intangible versus tangible objects, suggesting people have greater difficulty representing intangible items. Study 2b used fMRI to show that when stealing tangible objects versus intangible, participants had increased activation in left lateral orbitofr...

From an evolutionary perspective, one should be more sensitive when outgroup members attack the ingroup but less so when ingroup or outgroup members fight among themselves. Indeed, previous behavioral and neuroimaging research demonstrated that people show greater sensitivity for the suffering of ingroup compared with outgroup members. However, the question still remains whether this is always the case regardless of who is the agent causing the harm. To examine the role of agency and group membership in perception of harm, 48 participants were scanned while viewing ingroup or outgroup perpetrators intentionally harming ingroup or outgroup members. Behavioral results showed greater moral sensitivity for ingroup versus outgroup victims, but only when the perpetrator was from the outgroup. In support of this finding, fMRI data showed greater activity in left orbitofrontal cortex (OFC) for ingroup victims when they were harmed by outgroup individuals. In addition, effective connectivity...

Effective leaders are believed to inspire followers by providing inclusive visions of the future that followers can identify with. In the present study, we examined the neural mechanisms underlying this process, testing key hypotheses derived from transformational and social identity approaches to leadership. While undergoing functional MRI, supporters from the two major Australian political parties (Liberal vs. Labor) were presented with inspirational collective-oriented and noninspirational personal-oriented statements made by in-group and out-group leaders. Imaging data revealed that inspirational (rather than noninspirational) statements from in-group leaders were associated with increased activation in the bilateral rostral inferior parietal lobule, pars opercularis, and posterior midcingulate cortex: brain areas that are typically implicated in controlling semantic information processing. In contrast, for out-group leaders, greater activation in these areas was associated with...

Despite moral prohibitions on hurting other humans, some social contexts allow for harmful actions such as the killing of others. One example is warfare, where killing enemy soldiers is seen as morally justified. Yet, the neural underpinnings distinguishing between justified and unjustified killing are largely unknown. To improve understanding of the neural processes involved in justified and unjustified killing, participants had to imagine being the perpetrator whilst watching "first-person perspective" animated videos where they shot enemy soldiers ('justified violence') and innocent civilians ('unjustified violence'). When participants imagined themselves shooting civilians compared to soldiers, greater activation was found in the lateral orbitofrontal cortex (OFC). Regression analysis revealed that the more guilt participants felt about shooting civilians, the greater the response in the lateral OFC. Effective connectivity analyses further revealed an i...

To reduce the escalation of intergroup conflict, it is important that we understand the processes related to the detection of group-based threat and reconciliation. In the present study, we investigated the neural mechanisms of such processes using functional Magnetic Resonance
Imaging (fMRI). Functional neuroimaging techniques may shed light on quick, automatic responses to stimuli that happen outside of conscious awareness and are thus increasingly difficult to quantify relying only on participants’ self-reported experiences. They may further provide invaluable insight into physiological processes occurring in situations of sensitive nature, whereby participants–deliberately or not–may withhold their honest responses due to social desirability. Non-Muslim Western Caucasian participants watched short video clips of stereotypical Middle-Eastern Muslim males threatening their ingroup, offering reconciliation to the ingroup, or making a neutral statement. Threatening statements led to increased activation in the amygdala, insula,
supramarginal gyrus, and temporal lobe. Reconciliation efforts led to increased activation in the prefrontal cortex, anterior cingulate gyrus, and caudate. The results suggest that threat detection is a relatively automatic process while evaluating and responding to reconciliation offers requires more cognitive efforts. The implications of these findings and future research directions are discussed.

Although it is now widely recognised that social cognitive difficulties are often evident following
stroke, important questions remain about the nature and magnitude of these difficulties, as well as
the factors that determine the magnitude of this impairment. A meta-analysis of 58 datasets
involving 2567 participants (937 with stroke, 1630 non-clinical controls) was therefore conducted.
The results indicated that three of the four core domains of social cognitive function were
significantly disrupted in people with stroke. Specifically, while the effect size for affective
empathy failed to attain significance (r = -.33), moderate to large deficits were identified for theory
of mind (r = -.44), social perception (r = -.55), and social behaviour (r = -.53). These deficits were
robust across both left and right lateralized lesions, across social cognitive assessments that differed
in their broader cognitive demands, as well as in tasks that varied in their modality of presentation.
These data are discussed in the context of broader neuropsychological models of social cognitive
function.

Despite the apparent sociability of human kind, immoral behaviour is ever present in society. The term ‘immoral behaviour’ represents a complex array of conduct, ranging from insensitivity to topics of conversation through to violent assault and murder. To better understand the neuroscience of immoral behaviour, this review investigates two clinical populations that commonly present with changes in moral behaviour – behavioural-variant frontotemporal dementia and acquired brain injuries. Based on evidence from these groups, it is argued that rather than a single underlying cause,
immoral behaviour can result from three distinct types of cognitive failure: (1) problems understanding others; (2) difficulties controlling behaviour; or (3) deficits in the capacity to make appropriate emotional contributions. Each of these failures is associated with damage to
different brain regions. A more nuanced approach to the neuroscience of immoral behaviour has important implications for our understanding of immoral behaviour in a wide range of clinical groups, as well as human society more broadly.

Morality is an important social construct necessary for understanding what is right and wrong. Neuroimaging studies investigating morality have used a wide variety of paradigms and implicated many different brain areas. Yet, it remains unclear whether differences amongst morality tasks are the cause for such heterogeneous findings. Therefore, in the present study, a series of activation likelihood estimation (ALE) meta-analyses were conducted on 123 datasets (inclusive of 1963 participants) to address this question. The ALE meta-analyses revealed a series of common brain areas associated with all moral tasks, including medial prefrontal cortex, lateral orbitofrontal cortex, amygdala, temporoparietal junction, and precuneus. However, individual and contrast analyses also revealed unique networks associated with each moral modality, suggesting that different moral tasks recruit specialised brain regions.

Theory of mind (ToM) is an important skill that refers broadly to the capacity to understand the mental states of others. A large number of neuroimaging studies have focused on identifying the functional brain regions involved in ToM, but many important questions remain with respect to the neural networks implicated in specific types of ToM tasks. In the present study, we conducted a series of activation likelihood estimation (ALE) meta-analyses on 144 datasets (involving 3150 participants) to address these questions. The ALE results revealed common regions shared across all ToM tasks and broader task parameters, but also some important dissociations. In terms of commonalities, consistent activation was identified in the medial prefrontal cortex and bilateral temporoparietal junction. On the other hand, ALE contrast analyses on our dataset, as well as meta-analytic connectivity modelling (MACM) analyses on the BrainMap database, indicated that different types of ToM tasks reliably elicit activity in unique brain areas. Our findings provide the most accurate picture to date of the neural networks that underpin ToM function.

People often find it hard to disagree with others, but how this disposition varies across individuals or how it is influenced by social factors like other people's level of expertise remains little understood. Using functional magnetic resonance imaging (fMRI), we found that activity across a network of brain areas [comprising posterior medial frontal cortex (pMFC), anterior insula (AI), inferior frontal gyrus (IFG), lateral orbitofrontal cortex, and angular gyrus] was modulated by individual differences in the frequency with which participants actively disagreed with statements made by others. Specifically, participants who disagreed less frequently exhibited greater brain activation in these areas when they actually disagreed. Given the role of this network in cognitive dissonance, our results suggest that some participants had more trouble disagreeing due to a heightened cognitive dissonance response. Contrary to expectation, the level of expertise (high or low) had no effect on behavior or brain activity.

Racism and in-group favoritism is prevalent in our society and has been studied in Social Psychology for a long time. Recently it has become possible to investigate the neural mechanisms that underlie these in-group biases, and hence this review will give an overview of recent developments on the topic. Rather than relying on a single brain region or network, it seems that subtle changes in neural activation across the brain, depending on the modalities involved, underlie how we divide the world into ‘us’ versus ‘them’.
These insights have important implications for our understanding of how in-group biases develop and could potentially lead to new insights on how to reduce them.

Mirror neurons in macaque area F5 fire when an animal performs an action, such as a mouth or limb movement, and also when the animal passively observes an identical or similar action performed by another individual. Brain-imaging studies in humans conducted over the last 20 years have repeatedly attempted to reveal analogous brain regions with mirror properties in humans, with broad and often speculative claims about their functional significance across a range of cognitive domains, from language
to social cognition. Despite such concerted efforts, the likely neural substrates of these mirror regions have remained controversial, and indeed the very existence of a distinct subcategory of human neurons with mirroring properties has been questioned. Here we used activation likelihood estimation (ALE), to provide a quantitative index of the consistency of patterns of fMRI activity measured in human studies of action observation and action execution. From an initial sample of more than 300 published works, data from 125 papers met our strict inclusion and exclusion criteria. The analysis revealed 14 separate clusters in
which activation has been consistently attributed to brain regions with mirror properties, encompassing 9 different Brodmann areas. These clusters were located in areas purported to show mirroring properties in the macaque, such as the inferior parietal lobule, inferior frontal gyrus and the adjacent ventral premotor cortex, but surprisingly also in regions such as the primary visual cortex, cerebellum and parts of the limbic system. Our findings suggest a core network of human brain regions that possess mirror
properties associated with action observation and execution, with additional areas recruited during tasks that engage non-motor functions, such as auditory, somatosensory and affective components.

The critical lesion site responsible for the syndrome of unilatera lspatial neglect has been debated for more than a decade. Here we performed an activation likelihood estimation (ALE) to provide for the first time an objective quantitative index of the consistency of lesion sites across anatomical group studies of spatial neglect. The analysis revealed several distinct regions in which damage hasc onsistently been associated with spatial neglect symptoms. Lesioned clusters were located in several cortical and subcortical regions of the right hemisphere, including the middle and superior temporal gyrus, inferior parietal lobule, intraparietal sulcus, precuneus, middle occipital gyrus, caudate nucleus, and posterior insula, as well as in the white matter pathway corresponding to the posterior part of the superior longitudinal fasciculus. Further analyses suggested that separate lesion sites are associated with impairments in different behavioral tests, such as line bisection and target cancellation. Similarly, specific subcomponents of the heterogeneous neglects yndrome, such as extinction and allocentric and personal neglect, are associated with distinct lesion sites. Future progress in delineating the neuropathological correlates of spatial neglect will depend upon the development of more refined measures of perceptual and cognitive functions than those currently available in the clinical setting.

It has been suggested that the mirror neuron system provides an important neural substrate for humans’ ability to imitate. Mirror neurons have been found during single-cell recordings in monkeys in area F5 and PF. It is believed that the human equivalent of this mirror system in humans is the pars opercularis of the inferior frontal gyrus (area 44) and the rostral part of the inferior parietal lobule. This article critically reviews published fMRI studies that examined the role of frontal and parietal brain regions in imitation. A meta-analysis using activation likelihood estimation (ALE) revealed that the superior parietal lobule,
inferior parietal lobule, and the dorsal premotor cortex but not the inferior frontal gyrus, are all commonly involved in imitation. An additional meta-analysis using a label-based review confirmed that in the frontal lobe, the premotor cortex rather than the inferior frontal gyrus is consistently active in studies investigating imitation. In the parietal region the superior and inferior parietal lobules are equally activated during imitation. Our results suggest that parietal and frontal regions which extend beyond the classical mirror neuron network are crucial for imitation.

Group identification can lead to a biased view of the world in favor of “in-group” members. Studying the brain processes that underlie such in-group biases is important for a wider understanding of the potential influence of social factors on basic perceptual processes. In this study, we used functional magnetic resonance imaging (fMRI) to investigate how people perceive the actions of in-group and out-group members, and how their biased view in favor of own team members manifests itself in the brain. We divided participants into two teams and had them judge the relative speeds of hand actions performed by an in-group and an out-group member in a competitive situation. Participants judged hand actions performed by in-group members as being faster than those of out-group members, even when the two actions were performed at physically identical speeds. In an additional fMRI experiment, we showed that, contrary to common belief, such skewed impressions arise from a subtle bias in perception and associated brain activity rather than decision-making processes, and that this bias develops rapidly and involuntarily as a consequence of group affiliation. Our findings suggest that the neural mechanisms that underlie human perception are shaped by social context. Hum Brain Mapp 34:2055–2068, 2013. © 2012 Wiley Periodicals, Inc.

The Sustained Attention to Response task is a classical neuropsychological test that has been used by many
centres to characterize the attentional deficits in traumatic brain injury, ADHD, autism and other disorders. During the SART a random series of digits 1–9 is presented repeatedly and subjects have to respond to each digit (go trial) except the digit ‘3’ (no-go trial). Using voxel-based lesion symptommapping (VLSM) in
a consecutive series of 44 ischemic unifocal non-lacunar hemispheric stroke patients we determined the neuroanatomy of 4 SART parameters: commission and omission error rate, reaction time variability and post-error slowing. Lesions of the right inferior frontal gyrus significantly increased commission error rate. Lesions of the middle third of the right inferior frontal sulcus (IFS) reduced post-error slowing, a measure of how well subjects can utilize errors to adjust cognitive resource allocation. Omissions and reaction time variability had less localising value in our sample. To conclude, commission errors and post-error slowing in the SART mainly probe right inferior frontal integrity.

The parietal regions implicated in spatially selective attention differ between patient lesion studies and functional imaging of the intact brain. We aimed to resolve this discordance. In a voxel-based lesion-symptom mapping study in 20 ischemic stroke patients, we applied the same cognitive subtraction approach as in 23 healthy volunteers who underwent functional magnetic resonance imaging (fMRI) using identical tasks and stimuli. An instructive central cue directed attention to one visual quadrant. After a brief delay, a grating appeared in that quadrant together with an irrelevant grating in an uncued quadrant. Subjects had to discriminate the orientation of the grating in the cued quadrant. Patients with a right inferior parietal lesion were significantly more impaired during contralesional versus ipsilesional orienting when stimuli were bilateral and symmetrical than when stimuli occupied diagonally opposite quadrants or two quadrants within the same hemifield. In one area, the lesion-volume map overlapped with the activity map obtained in healthy volunteers: the lower bank of the middle third of the right intraparietal sulcus (IPS). In an additional 37 healthy fMRI subjects, we disentangled the effects of
symmetry, bilaterality, and spatial configuration between stimuli on activity in the volume of overlap. Only the axis of configuration
between stimuli had a significant effect, with highest activity when the configuration axis was horizontal. This constitutes converging
evidence from patients and cognitively intact subjects that the lower bank of the middle third of the right IPS critically contributes to attentive selection between competing stimuli in a spatially anisotropic manner.

The role of the mirror system in action understanding has been widely debated. Some authors have suggested that the mirror system plays an important role in action understanding (Rizzolatti and Sinigaglia, 2010), whereas others have claimed that direct evidence to support this view is lacking (Hickok, 2009). If mirror
neurons have an active role in action understanding rather than passive visuomotor transformation during action observation, they should respond differently to the observation of actions depending on the intentions of the observer. In this fMRI study, twenty participants observed identical actions under different instruction
contexts. The task was either to understand the actions, identify the physical features of the actions, or passively observe the actions. A multi-voxel pattern analysis revealed unique patterns of activation in ventral premotor cortex and inferior parietal lobule across the different contexts. The results suggest that ventral
premotor and inferior parietal areas respond differently to observed actions depending on the mindset of the observer. This is consistent with the view that these regions do not merely process observed actions passively, but play an active role in action understanding.

Right-sided parietal lesions lead to lateralized attentional deficits which are most prominent with bilateral stimulation. We determined how an irrelevant stimulus in the unattended hemifield alters attentional responses in parietal cortex during unilateral orienting. A trial consisted of a central spatial cue, a delay
and a test phase during which a grating was presented at 9
deg eccentricity. Subjects had to discriminate the orientation of the grating. The unattended hemifield was either empty or contained a second, irrelevant grating. We carried out a series of functional MRI (fMRI) studies in 35 healthy volunteers (13 men and 22
women, aged between 19 and 30 years) as well as a behavioural and structural lesion mapping study in 17 righthemispheric
lesion patients, 11 of whom had neglect. In the patients with but not in those without neglect, the addition of a distractor in the unattended hemifield significantly impaired performance if attention was directed contralesionally but not if it was directed ipsilesionally. In the healthy volunteers, we discerned two functionally distinct areas along the posterior–anterior axis of the intraparietal sulcus (IPS). The posterior, descending IPS segment in both hemispheres showed attentional enhancement of responses during contralateral attentional orienting and was unaffected by the presence of an irrelevant stimulus in the ignored hemifield. In contrast, the right-sided horizontal IPS segment showed a strong attentional response when subjects oriented to a stimulus in the relevant hemifield and an irrelevant stimulus was simultaneously present in the ignored hemifield, compared with unilateral stimulation. This effect was independent of the direction of attention. The symmetrical left-sided horizontal IPS segment showed the highest responses under the same circumstances, in combination with a contralateral bias during unilateral stimulation conditions. None of the six patients without neglect had a lesion of the horizontal IPS segment. In four of the 11 neglect patients, the lesion overlapped with the horizontal IPS activity cluster and lay in close proximity to it in another four. The remaining three patients had a lesion at a distance from the parietal cortex. Our findings reconcile the role of the IPS in endogenous attentional control with the clinically significant interaction between direction of attention and bilateral stimulation in right parietal lesion patients. Functional imaging in neglect patients will be necessary to assess IPS function in those cases where the structural lesion spares the middle IPS segment.

Damage to the parietal lobe can induce a condition known as spatial neglect, characterized by a lack of awareness of the
personal and/or extrapersonal space opposite the damaged brain region. Spatial neglect is commonly assessed clinically
using either the line bisection or the target cancellation task. However, it is unclear whether poor performance on each of
these two tasks is associated with the same or different lesion locations. To date, methodological limitations and differences
have prevented a definitive link between task performance and lesion location to be made. Here we report findings from a
voxel-based lesion symptom mapping (VLSM) analysis of an unbiased selection of 44 patients with a recent unifocal stroke.
Patients performed both the line bisection and target cancellation task. For each of the two tasks a continuous score was
incorporated into the VLSM analysis. Both tasks correlated highly with each other (r = .76) and VLSM analyses indicated that
the angular gyrus was the critical lesion site for both tasks. The results suggest that both tasks probe the same underlying
cortical deficits and although the cancellation task was more sensitive than the line bisection task, both can be used in a
clinical setting to test for spatial neglect.

It has been suggested that in humans the mirror neuron system provides a neural substrate for imitation behaviour, but the relative contributions of different brain regions to the imitation of manual actions is still a matter of debate. To investigate the role of the mirror neuron system in imitation we used fMRI to examine patterns of neural activity under four different conditions: passive observation of a pantomimed action (e.g., hammering a nail); (2) imitation of an observed action; (3) execution of an action in response to a word cue; and (4) self-selected execution of an action. A network of cortical areas, including the left supramarginal gyrus, left superior parietal lobule, left dorsal premotor area and bilateral superior temporal sulcus (STS), was significantly active across all four conditions. Crucially, within this network the STS bilaterally was the only region in which activity was significantly greater for action imitation than for the passive observation and execution conditions. We suggest that the role of the STS in imitation is not merely to passively register observed biological motion, but rather to actively represent visuomotor correspondences between one's own actions and the actions of others. Hum Brain Mapp, 2010. © 2010 Wiley-Liss, Inc.

According to a longstanding view, inferior as opposed to superior parietal cortex critically contributes to the spatial attentional deficits encountered following unilateral parietal ischemic lesions. We review the evidence on which this view is based and contrast it with more recent structural lesion evidence concerning the critical role of the intraparietal sulcus in spatial attention deficits. In a classical spatial cueing paradigm, focal lesions of the posterior and of the middle segment of the intraparietal sulcus
give rise to a pathological invalidity effect that is indistinguishable from that seen after classical inferior parietal lesions. When a competing distracter is added to a target stimulus, the deleterious consequences of focal IPS lesions are again very similar to those classically observed following inferior parietal lesions. The deficit could not be accounted for by functional effects at a distance affecting inferior parietal cortex. These single-case lesion data establish the critical role of the posterior and the middle IPS segment in spatially selective attention and are in line with a vast amount of functional imaging evidence in the intact brain pointing to the prominent role of the intraparietal sulcus in spatial attention, along with inferior parietal cortex under specific circumstances. Functional imaging has also provided hints about
the differences in functional contribution between inferior and superior parietal cortex. These hypotheses await further confirmation based on lesion evidence.