Showing papers in "Frontiers in Human Neuroscience in 2012"

Self-awareness, self-regulation, and self-transcendence (S-ART): a framework for understanding the neurobiological mechanisms of mindfulness.

Abstract: Mindfulness-as a state, trait, process, type of meditation, and intervention has proven to be beneficial across a diverse group of psychological disorders as well as for general stress reduction. Yet, there remains a lack of clarity in the operationalization of this construct, and underlying mechanisms. Here, we provide an integrative theoretical framework and systems-based neurobiological model that explains the mechanisms by which mindfulness reduces biases related to self-processing and creates a sustainable healthy mind. Mindfulness is described through systematic mental training that develops meta-awareness (self-awareness), an ability to effectively modulate one's behavior (self-regulation), and a positive relationship between self and other that transcends self-focused needs and increases prosocial characteristics (self-transcendence). This framework of self-awareness, -regulation, and -transcendence (S-ART) illustrates a method for becoming aware of the conditions that cause (and remove) distortions or biases. The development of S-ART through meditation is proposed to modulate self-specifying and narrative self-networks through an integrative fronto-parietal control network. Relevant perceptual, cognitive, emotional, and behavioral neuropsychological processes are highlighted as supporting mechanisms for S-ART, including intention and motivation, attention regulation, emotion regulation, extinction and reconsolidation, prosociality, non-attachment, and decentering. The S-ART framework and neurobiological model is based on our growing understanding of the mechanisms for neurocognition, empirical literature, and through dismantling the specific meditation practices thought to cultivate mindfulness. The proposed framework will inform future research in the contemplative sciences and target specific areas for development in the treatment of psychological disorders.

Introduction to Research Topic – Binocular Rivalry: A Gateway to Studying Consciousness

Abstract: In 1593, Neapolitan polymath Giambattista della Porta publicly lamented that he was unable to improve his impressive productivity (he had published in areas as diverse as cryptography, hydraulics, pharmacology, optics, and classic fiction). Della Porta was trying to read two books simultaneously by placing both volumes side-by-side, and using each eye independently. To his great surprise, his setup allowed him to only read one book at a time. This discovery arguably marks the first written account of binocular rivalry (Wade, 2000) – a perceptual phenomenon that more than 400 years later still both serves to intrigue as well as to illuminate the limits of scientific knowledge. At first glance, binocular rivalry is an oddball. In every day vision, our eyes receive largely matching views of the world. The brain combines the two images into a cohesive scene, and concurrently, perception is stable. However, when showing two very different images (such as two different books) to each eye, the brain resolves the conflict by adopting a “diplomatic” strategy. Rather than mixing the views of the two eyes into an insensible visual percept, observers perceive a dynamically changing series of perceptual snapshots, with one eye’s view dominating for a few seconds before being replaced by its rival from the other eye. With prolonged viewing of a rivalrous stimulus, one inevitably experiences a sequence of subjective perceptual reversals, separated by random time intervals, and this process continues for as long as the sensory conflict is present.

On the relationship between the "default mode network" and the "social brain".

Abstract: The default mode network (DMN) of the brain consists of areas that are typically more active during rest than during active task performance. Recently however, this network has been shown to be activated by certain types of tasks. Social cognition, particularly higher-order tasks such as attributing mental states to others, has been suggested to activate a network of areas at least partly overlapping with the DMN. Here, we explore this claim, drawing on evidence from meta-analyses of functional MRI data and recent studies investigating the structural and functional connectivity of the social brain. In addition, we discuss recent evidence for the existence of a DMN in non-human primates. We conclude by discussing some of the implications of these observations.

Moving a Rubber Hand that Feels Like Your Own: A Dissociation of Ownership and Agency.

Abstract: During voluntary hand movement, we sense that we generate the movement and that the hand is a part of our body. These feelings of control over bodily actions, or the sense of agency, and the ownership of body parts are two fundamental aspects of the way we consciously experience our bodies. However, little is known about how these processes are functionally linked. Here, we introduce a version of the rubber hand illusion in which participants control the movements of the index finger of a model hand, which is in full view, by moving their own right index finger. We demonstrated that voluntary finger movements elicit a robust illusion of owning the rubber hand and that the senses of ownership and agency over the model hand can be dissociated. We systematically varied the relative timing of the finger movements (synchronous versus asynchronous), the mode of movement (active versus passive), and the position of the model hand (anatomically congruent versus incongruent positions). Importantly, asynchrony eliminated both ownership and agency, passive movements abolished the sense of agency but left ownership intact, and incongruent positioning of the model hand diminished ownership but did not eliminate agency. These findings provide evidence for a double dissociation of ownership and agency, suggesting that they represent distinct cognitive processes. Interestingly, we also noted that the sense of agency was stronger when the hand was perceived to be a part of the body, and only in this condition did we observe a significant correlation between the subjects’ ratings of agency and ownership. We discuss this in the context of possible differences between agency over owned body parts and agency over actions that involve interactions with external objects. In summary, the results obtained in this study using a simple moving rubber hand illusion paradigm extend previous findings on the experience of ownership and agency and shed new light on their relationship.

Neuroimaging of child abuse: a critical review

Abstract: Childhood maltreatment is a severe stressor that can lead to the development of behaviour problems and affect brain structure and function. This review summarizes the current evidence for the effects of early childhood maltreatment on behavior, cognition and the brain in adults and children. Neuropsychological studies suggest an association between child abuse and deficits in IQ, memory, executive function and emotion discrimination. Structural neuroimaging studies provide evidence for deficits in brain volume, grey and white matter of several regions, most prominently the dorsolateral and ventromedial prefrontal cortex but also hippocampus, amygdala, and corpus callosum. Diffusion tensor imaging studies show evidence for deficits in structural interregional connectivity between these areas, suggesting neural network abnormalities. Functional imaging studies support this evidence by reporting atypical activation in the same brain regions during executive function and emotion processing. There are, however, several limitations of the abuse research literature which are discussed, most prominently the lack of control for co-morbid psychiatric disorders, which make it difficult to disentangle which of the above effects are due to maltreatment, the associated psychiatric conditions or a combination or interaction between both. Overall, the better controlled studies that show a direct correlation between childhood abuse and brain measures suggest that the most prominent deficits associated with early childhood abuse are in the function and structure of lateral and ventromedial fronto-limbic brain areas and networks that mediate behavioural and affect control. Future, large scale multimodal neuroimaging studies in medication-naive subjects, however, are needed that control for psychiatric co-morbidities in order to elucidate the structural and functional brain sequelae that are associated with early environmental adversity, independently of secondary co-morbid conditions.

A sensorimotor paradigm for Bayesian model selection

Abstract: Sensorimotor control is thought to rely on predictive internal models in order to cope efficiently with uncertain environments. Recently, it has been shown that humans not only learn different internal models for different tasks, but that they also extract common structure between tasks. This raises the question of how the motor system selects between different structures or models, when each model can be associated with a range of different task-specific parameters. Here we design a sensorimotor task that requires subjects to compensate visuomotor shifts in a three-dimensional virtual reality setup, where one of the dimensions can be mapped to a model variable and the other dimension to the parameter variable. By introducing probe trials that are neutral in the parameter dimension, we can directly test for model selection. We found that model selection procedures based on Bayesian statistics provided a better explanation for subjects’ choice behavior than simple non-probabilistic heuristics. Our experimental design lends itself to the general study of model selection in a sensorimotor context as it allows to separately query model and parameter variables from subjects.

Look at this: the neural correlates of initiating and responding to bids for joint attention

Abstract: When engaging in joint attention, one person directs another person's attention to an object (Initiating Joint Attention, IJA), and the second person's attention follows (Responding to Joint Attention, RJA). As such, joint attention must occur within the context of a social interaction. This ability is critical to language and social development; yet the neural bases for this pivotal skill remain understudied. This paucity of research is likely due to the challenge in acquiring functional MRI data during a naturalistic, contingent social interaction. To examine the neural bases of both IJA and RJA we implemented a dual-video set-up that allowed for a face-to-face interaction between subject and experimenter via video during fMRI data collection. In each trial, participants either followed the experimenter's gaze to a target (RJA) or cued the experimenter to look at the target (IJA). A control condition, solo attention (SA), was included in which the subject shifted gaze to a target while the experimenter closed her eyes. Block and event-related analyses were conducted and revealed common and distinct regions for IJA and RJA. Distinct regions included the ventromedial prefrontal cortex for RJA and intraparietal sulcus and middle frontal gyrus for IJA (as compared to SA). Conjunction analyses revealed overlap in the dorsal medial prefrontal cortex (dMPFC) and right posterior superior temporal sulcus (pSTS) for IJA and RJA (as compared to SA) for the event analyses. Functional connectivity analyses during a resting baseline suggest joint attention processes recruit distinct but interacting networks, including social-cognitive, voluntary attention orienting, and visual networks. This novel experimental set-up allowed for the identification of the neural bases of joint attention during a real-time interaction and findings suggest that whether one is the initiator or responder, the dMPFC and right pSTS, are selectively recruited during periods of joint attention.

Oscillatory power decreases and long-term memory: the information via desynchronization hypothesis

Abstract: The traditional belief is that brain oscillations are important for human long-term memory, because they induce synchronized firing between cell assemblies which shapes synaptic plasticity. Therefore, most prior studies focused on the role of synchronization for episodic memory, as reflected in theta (∼5 Hz) and gamma (>40 Hz) power increases. These studies, however, neglect the role that is played by neural desynchronization, which is usually reflected in power decreases in the alpha and beta frequency band (8-30 Hz). In this paper we present a first idea, derived from information theory that gives a mechanistic explanation of how neural desynchronization aids human memory encoding and retrieval. Thereby we will review current studies investigating the role of alpha and beta power decreases during long-term memory tasks and show that alpha and beta power decreases play an important and active role for human memory. Applying mathematical models of information theory, we demonstrate that neural desynchronization is positively related to the richness of information represented in the brain, thereby enabling encoding and retrieval of long-term memories. This information via desynchronization hypothesis makes several predictions, which can be tested in future experiments.

Working-memory training in younger and older adults: training gains, transfer, and maintenance

Abstract: Working memory (WM), a key determinant of many higher-order cognitive functions, declines in old age. Current research attempts to develop process-specific WM training procedures, which may lead to general cognitive improvement. Adaptivity of the training as well as the comparison of training gains to performance changes of an active control group are key factors in evaluating the effectiveness of a specific training program. In the present study, 55 younger adults (20–30 years of age) and 45 older adults (60–70 years of age) received 5 weeks of computerized training on various spatial and verbal WM tasks. Half of the sample received adaptive training (i.e., individually adjusted task difficulty), whereas the other half-worked on the same task material but on a low task difficulty level (active controls). Performance was assessed using criterion, near-transfer, and far-transfer tasks before training, after 5 weeks of intervention, as well as after a 3-month follow-up interval. Results indicate that (a) adaptive training generally led to larger training gains than low-level practice, (b) training and transfer gains were somewhat greater for younger than for older adults in some tasks, but comparable across age groups in other tasks, (c) far-transfer was observed to a test on sustained attention and for a self-rating scale on cognitive functioning in daily life for both young and old, and (d) training gains and transfer effects were maintained across the 3-month follow-up interval across age.

Effects of mindful-attention and compassion meditation training on amygdala response to emotional stimuli in an ordinary, non-meditative state

Abstract: The amygdala has been repeatedly implicated in emotional processing of both positive and negative valence stimuli. Previous studies suggest that the amygdala response to emotional stimuli is lower when the subject is in a meditative state of mindful attention, both in beginner meditators after an eight-week meditation intervention and in expert meditators. However, the longitudinal effects of meditation training on amygdala responses have not been reported when participants are in an ordinary, non-meditative state. In this study, we investigated how eight weeks of training in meditation affects amygdala responses to emotional stimuli in subjects when in a non-meditative state. Healthy adults with no prior meditation experience took part in eight weeks of either Mindful Attention Training, Cognitively-Based Compassion Training (CBCT; a program based on Tibetan Buddhist compassion meditation practices), or an active control intervention. Before and after the intervention, participants underwent an fMRI experiment during which they were presented images with positive, negative, and neutral emotional valences from the IAPS database while remaining in an ordinary, non-meditative state. Using a region-of-interest analysis, we found a longitudinal decrease in right amygdala activation in the Mindful Attention group in response to positive images, and in response to images of all valences overall. In the CBCT group, we found a trend increase in right amygdala response to negative images, which was significantly correlated with a decrease in depression score. No effects or trends were observed in the control group. This finding suggests that the effects of meditation training on emotional processing might transfer to non-meditative states. This is consistent with the hypothesis that meditation training may induce learning that is not stimulus- or task-specific, but process-specific, and thereby may result in enduring changes in mental function.

Naturalistic FMRI mapping reveals superior temporal sulcus as the hub for the distributed brain network for social perception.

Abstract: Despite the abundant data on brain networks processing static social signals, such as pictures of faces, the neural systems supporting social perception in naturalistic conditions are still poorly understood. Here we delineated brain networks subserving social perception under naturalistic conditions in 19 healthy humans who watched, during 3-tesla functional magnetic imaging (fMRI), a set of 137 short (~16 s each, total 27 min) audiovisual movie clips depicting pre-selected social signals. Two independent raters estimated how well each clip represented eight social features (faces, human bodies, biological motion, goal-oriented actions, emotion, social interaction, pain, and speech) and six filler features (places, objects, rigid motion, people not in social interaction, non-goal-oriented action and non-human sounds) lacking social content. These ratings were used as predictors in the fMRI analysis. The posterior superior temporal sulcus (STS) responded to all social features but not to any non-social features, and the anterior STS responded to all social features except bodies and biological motion. We also found four partially segregated, extended networks for processing of specific social signals: 1) a fronto-temporal network responding to multiple social categories, 2) a fronto-parietal network preferentially activated to bodies, motion and pain, 3) a temporo-amygdalar network responding to faces, social interaction and speech, and 4) a fronto-insular network responding to pain, emotions, social interactions, and speech. Our results highlight the role of the posterior STS in processing multiple aspects of social information, as well as the feasibility and efficiency of fMRI mapping under conditions that resemble the complexity of real life.

Effects of Meditation Experience on Functional Connectivity of Distributed Brain Networks

Abstract: This study sought to examine the effect of meditation experience on brain networks underlying cognitive actions employed during contemplative practice. In a previous study, we proposed a basic model of naturalistic cognitive fluctuations that occur during the practice of focused attention meditation. This model specifies four intervals in a cognitive cycle: mind wandering (MW), awareness of MW, shifting of attention, and sustained attention. Using subjective input from experienced practitioners during meditation, we identified activity in salience network regions during awareness of MW and executive network regions during shifting and sustained attention. Brain regions associated with the default mode were active during MW. In the present study, we reasoned that repeated activation of attentional brain networks over years of practice may induce lasting functional connectivity changes within relevant circuits. To investigate this possibility, we created seeds representing the networks that were active during the four phases of the earlier study, and examined functional connectivity during the resting state in the same participants. Connectivity maps were then contrasted between participants with high vs. low meditation experience. Participants with more meditation experience exhibited increased connectivity within attentional networks, as well as between attentional regions and medial frontal regions. These neural relationships may be involved in the development of cognitive skills, such as maintaining attention and disengaging from distraction, that are often reported with meditation practice. Furthermore, because altered connectivity of brain regions in experienced meditators was observed in a non-meditative (resting) state, this may represent a transference of cognitive abilities “off the cushion” into daily life.

Combining EEG and eye tracking: identification, characterization, and correction of eye movement artifacts in electroencephalographic data

Abstract: Eye movements introduce large artifacts to electroencephalographic recordings (EEG) and thus render data analysis difficult or even impossible. Trials contaminated by eye movement and blink artifacts have to be discarded, hence in standard EEG-paradigms subjects are required to fixate on the screen. To overcome this restriction, several correction methods including regression and blind source separation have been proposed. Yet, there is no automated standard procedure established. By simultaneously recording eye movements and 64-channel-EEG during a guided eye movement paradigm, we investigate and review the properties of eye movement artifacts, including corneo-retinal dipole changes, saccadic spike potentials and eyelid artifacts, and study their interrelations during different types of eye- and eyelid movements. In concordance with earlier studies our results confirm that these artifacts arise from different independent sources and that depending on electrode site, gaze direction, and choice of reference these sources contribute differently to the measured signal. We assess the respective implications for artifact correction methods and therefore compare the performance of two prominent approaches, namely linear regression and independent component analysis (ICA). We show and discuss that due to the independence of eye artifact sources, regression-based correction methods inevitably over- or under-correct individual artifact components, while ICA is in principle suited to address such mixtures of different types of artifacts. Finally, we propose an algorithm, which uses eye tracker information to objectively identify eye-artifact related ICA-components (ICs) in an automated manner. In the data presented here, the algorithm performed very similar to human experts when those were given both, the topographies of the ICs and their respective activations in a large amount of trials. Moreover it performed more reliable and almost twice as effective than human experts when those had to base their decision on IC topographies only. Furthermore, a receiver operating characteristic (ROC) analysis demonstrated an optimal balance of false positive and false negative at an area under curve (AUC) of more than 0.99. Removing the automatically detected ICs from the data resulted in removal or substantial suppression of ocular artifacts including microsaccadic spike potentials, while the relevant neural signal remained unaffected. In conclusion the present work aims at a better understanding of individual eye movement artifacts, their interrelations and the respective implications for eye artifact correction. Additionally, the proposed ICA-procedure provides a tool for optimized detection and correction of eye movement-related artifact components.

Correlated components of ongoing EEG point to emotionally laden attention - a possible marker of engagement?

Abstract: Recent evidence from functional magnetic resonance imaging suggests that cortical hemodynamic responses coincide in different subjects experiencing a common naturalistic stimulus. Here we utilize neural responses in the electroencephalogram (EEG) evoked by multiple presentations of short film clips to index brain states marked by high levels of correlation within and across subjects. We formulate a novel signal decomposition method which extracts maximally correlated signal components from multiple EEG records. The resulting components capture correlations down to a one-second time resolution, thus revealing that peak correlations of neural activity across viewings can occur in remarkable correspondence with arousing moments of the film. Moreover, a significant reduction in neural correlation occurs upon a second viewing of the film or when the narrative is disrupted by presenting its scenes scrambled in time. We also probe oscillatory brain activity during periods of heightened correlation, and observe during such times a significant increase in the theta band for a frontal component and reductions in the alpha and beta frequency bands for parietal and occipital components. Low-resolution EEG tomography of these components suggests that the correlated neural activity is consistent with sources in the cingulate and orbitofrontal cortices. Put together, these results suggest that the observed synchrony reflects attention- and emotion-modulated cortical processing which may be decoded with high temporal resolution by extracting maximally correlated components of neural activity.

The hippocampus supports multiple cognitive processes through relational binding and comparison.

Abstract: It has been well established that the hippocampus plays a pivotal role in explicit long-term recognition memory. However, findings from amnesia, lesion and recording studies with non-human animals, eye-movement recording studies, and functional neuroimaging have recently converged upon a similar message: the functional reach of the hippocampus extends far beyond explicit recognition memory. Damage to the hippocampus affects performance on a number of cognitive tasks including recognition memory after short and long delays and visual discrimination. Additionally, with the advent of neuroimaging techniques that have fine spatial and temporal resolution, findings have emerged that show the elicitation of hippocampal responses within the first few 100 ms of stimulus/task onset. These responses occur for novel and previously viewed information during a time when perceptual processing is traditionally thought to occur, and long before overt recognition responses are made. We propose that the hippocampus is obligatorily involved in the binding of disparate elements across both space and time, and in the comparison of such relational memory representations. Furthermore, the hippocampus supports relational binding and comparison with or without conscious awareness for the relational representations that are formed, retrieved and/or compared. It is by virtue of these basic binding and comparison functions that the reach of the hippocampus extends beyond long-term recognition memory and underlies task performance in multiple cognitive domains.

Regular, brief mindfulness meditation practice improves electrophysiological markers of attentional control

Abstract: Mindfulness based meditation practices involve various attentional skills, including the ability to sustain and focus ones attention. During a simple mindful breathing practice, sustained attention is required to maintain focus on the breath while cognitive control is required to detect mind wandering. We thus hypothesized that regular, brief mindfulness training would result in improvements in the self regulation of attention and foster changes in neuronal activity related to attentional control. A longitudinal randomized control group EEG study was conducted. At baseline (T1), 40 meditation naive participants were randomized into a wait list group and a meditation group, who received three hours mindfulness meditation training. 28 participants remained in the final analysis. At T1, after 8 weeks (T2) and after 16 weeks (T3), all participants performed a computerized Stroop task (a measure of attentional control) while the 64-channel EEG was recorded. Between T1 and T3 the meditators were requested to meditate daily for ten minutes. Event-related potential (ERP) analysis highlighted two between group effects that developed over the course of the 16-week mindfulness training. An early effect at left and right posterior sites 160 – 240 ms post stimulus indicated that meditation practice improved the focusing of attentional resources. A second effect at central posterior sites 310 – 380 ms post stimulus reflects that meditation practice reduced the recruitment of resources during object recognition processes, especially for incongruent stimuli. Scalp topographies and source analyses (VARETA) indicate relevant changes in neural sources, pertaining to left medial and lateral occipitotemporal areas for the early effect and right lateral occipitotemporal and inferior temporal areas for the later effect. The results suggest that mindfulness meditation may alter the efficiency of allocating cognitive resources, leading to improved self regulation of attention.

Dynamic BOLD functional connectivity in humans and its electrophysiological correlates.

Abstract: Neural oscillations subserve many human perceptual and cognitive operations. Accordingly, brain functional connectivity is not static in time, but fluctuates dynamically following the synchronization and desynchronization of neural populations. This dynamic functional connectivity has recently been demonstrated in spontaneous fluctuations of the Blood Oxygen Level-Dependent (BOLD) signal, measured with functional Magnetic Resonance Imaging (fMRI). We analyzed temporal fluctuations in BOLD connectivity and their electrophysiological correlates, by means of long (≈50 min) joint electroencephalographic (EEG) and fMRI recordings obtained from two populations: 15 awake subjects and 13 subjects undergoing vigilance transitions. We identified positive and negative correlations between EEG spectral power (extracted from electrodes covering different scalp regions) and fMRI BOLD connectivity in a network of 90 cortical and subcortical regions (with millimeter spatial resolution). In particular, increased alpha (8–12 Hz) and beta (15–30 Hz) power were related to decreased functional connectivity, whereas gamma (30–60 Hz) power correlated positively with BOLD connectivity between specific brain regions. These patterns were altered for subjects undergoing vigilance changes, with slower oscillations being correlated with functional connectivity increases. Dynamic BOLD functional connectivity was reflected in the fluctuations of graph theoretical indices of network structure, with changes in frontal and central alpha power correlating with average path length. Our results strongly suggest that fluctuations of BOLD functional connectivity have a neurophysiological origin. Positive correlations with gamma can be interpreted as facilitating increased BOLD connectivity needed to integrate brain regions for cognitive performance. Negative correlations with alpha suggest a temporary functional weakening of local and long-range connectivity, associated with an idling state.

The two-brain approach: how can mutually interacting brains teach us something about social interaction?

Abstract: Measuring brain activity simultaneously from two people interacting is intuitively appealing if one is interested in putative neural markers of social interaction. However, given the complex nature of interactions, it has proven difficult to carry out two-person brain imaging experiments in a methodologically feasible and conceptually relevant way. Only a small number of recent studies have put this into practice, using fMRI, EEG, or NIRS. Here, we review two main two-brain methodological approaches, each with two conceptual strategies. The first group has employed two-brain fMRI recordings, studying (1) turn-based interactions on the order of seconds, or (2) pseudo-interactive scenarios, where only one person is scanned at a time, investigating the flow of information between brains. The second group of studies has recorded dual EEG/NIRS from two people interacting, in (1) face-to-face turn-based interactions, investigating functional connectivity between theory-of-mind regions of interacting partners, or in (2) continuous mutual interactions on millisecond timescales, to measure coupling between the activity in one person's brain and the activity in the other's brain. We discuss the questions these approaches have addressed, and consider scenarios when simultaneous two-brain recordings are needed. Furthermore, we suggest that (1) quantification of inter-personal neural effects via measures of emergence, and (2) multivariate decoding models that generalize source-specific features of interaction, may provide novel tools to study brains in interaction. This may allow for a better understanding of social cognition as both representation and participation.

Event-related potential studies of outcome processing and feedback-guided learning.

Abstract: In order to control behavior in an adaptive manner the brain has to learn how some situations and actions predict positive or negative outcomes. During the last decade cognitive neuroscientists have shown that the brain is able to evaluate and learn from outcomes within a few hundred milliseconds of their occurrence. This research has been primarily focused on the feedback-related negativity (FRN) and the P3, two event-related potential (ERP) components that are elicited by outcomes. The FRN is a frontally distributed negative-polarity ERP component that typically reaches its maximal amplitude 250 ms after outcome presentation and tends to be larger for negative than for positive outcomes. The FRN has been associated with activity in the anterior cingulate cortex. The P3 (~300-600 ms) is a parietally distributed positive-polarity ERP component that tends to be larger for large magnitude than for small magnitude outcomes. The neural sources of the P3 are probably distributed over different regions of the cortex. This paper examines the theories that have been proposed to explain the functional role of these two ERP components during outcome processing. Special attention is paid to extant literature addressing how these ERP components are modulated by outcome valence (negative vs. positive), outcome magnitude (large vs. small), outcome probability (unlikely vs. likely) and behavioral adjustment. The literature offers few generalizable conclusions, but is beset with a number of inconsistencies across studies. This paper discusses the potential reasons for these inconsistencies and points out some challenges that will shape the field over the next decade.

Social attention with real versus reel stimuli: toward an empirical approach to concerns about ecological validity.

Abstract: Cognitive neuroscientists often study social cognition by using simple but socially relevant stimuli, such as schematic faces or images of other people. Whilst this research is valuable, important aspects of genuine social encounters are absent from these studies, a fact that has recently drawn criticism. In the present review we argue for an empirical approach to the determination of the equivalence of different social stimuli. This approach involves the systematic comparison of different types of social stimuli ranging in their approximation to a real social interaction. In garnering support for this cognitive ethological approach, we focus on recent research in social attention that has involved stimuli ranging from simple schematic faces to real social interactions. We highlight both meaningful similarities and differences in various social attentional phenomena across these different types of social stimuli thus validating the utility of the research initiative. Furthermore, we argue that exploring these similarities and differences will provide new insights into social cognition and social neuroscience.

The interactive brain hypothesis

Abstract: Enactive approaches foreground the role of interpersonal interaction in explanations of social understanding. This motivates, in combination with a recent interest in neuroscientific studies involving actual interactions, the question of how interactive processes relate to neural mechanisms involved in social understanding. We introduce the Interactive Brain Hypothesis (IBH) in order to help map the spectrum of possible relations between social interaction and neural processes. The hypothesis states that interactive experience and skills play enabling roles in both the development and current function of social brain mechanisms, even in cases where social understanding happens in the absence of immediate interaction. We examine the plausibility of this hypothesis against developmental and neurobiological evidence and contrast it with the widespread assumption that mindreading is crucial to all social cognition. We describe the elements of social interaction that bear most directly on this hypothesis and discuss the empirical possibilities open to social neuroscience. We propose that the link between coordination dynamics and social understanding can be best grasped by studying transitions between states of coordination. These transitions form part of the self-organization of interaction processes that characterize the dynamics of social engagement. The patterns and synergies of this self-organization help explain how individuals understand each other. Various possibilities for role-taking emerge during interaction, determining a spectrum of participation. This view contrasts sharply with the observational stance that has guided research in social neuroscience until recently. We also introduce the concept of readiness to interact to describe the practices and dispositions that are summoned in situations of social significance (even if not interactive). This latter idea links interactive factors to more classical observational scenarios.

Neural synchrony within the motor system: what have we learned so far?

Abstract: Synchronization of neural activity is considered essential for information processing in the nervous system. Both local and inter-regional synchronization are omnipresent in different frequency regimes and relate to a variety of behavioral and cognitive functions. Over the years, many studies have sought to elucidate the question how alpha/mu, beta, and gamma synchronization contribute to motor control. Here, we review these studies with the purpose to delineate what they have added to our understanding of the neural control of movement. We highlight important findings regarding oscillations in primary motor cortex, synchronization between cortex and spinal cord, synchronization between cortical regions, as well as abnormal synchronization patterns in a selection of motor dysfunctions. The interpretation of synchronization patterns benefits from combining results of invasive and non-invasive recordings, different data analysis tools, and modeling work. Importantly, although synchronization is deemed to play a vital role, it is not the only mechanism for neural communication. Spike timing and rate coding act together during motor control and should therefore both be accounted for when interpreting movement-related activity.

The brain on art: intense aesthetic experience activates the default mode network

Abstract: Aesthetic responses to visual art comprise multiple types of experiences, from sensation and perception to emotion and self-reflection. Moreover, aesthetic experience is highly individual, with observers varying significantly in their responses to the same artwork. Combining fMRI and behavioral analysis of individual differences in aesthetic response, we identify two distinct patterns of neural activity exhibited by different subnetworks. Activity increased linearly with observers’ ratings (4-level scale) in sensory (occipito-temporal) regions. Activity in the striatum also varied linearly with ratings, with below-baseline activations for low-rated artworks. In contrast, a network of frontal regions showed a step-like increase only for the most moving artworks (“4” ratings) and non-differential activity for all others. This included several regions belonging to the “default mode network” previously associated with self-referential mentation. Our results suggest that aesthetic experience involves the integration of sensory and emotional reactions in a manner linked with their personal relevance.

High frequency deep brain stimulation attenuates subthalamic and cortical rhythms in Parkinson's disease

Abstract: Parkinson’s disease (PD) is marked by excessive synchronous activity in the beta (8-35 Hz) band throughout the cortico-basal ganglia network. The optimal location of high frequency deep brain stimulation (HF DBS) within the subthalamic nucleus (STN) region and the location of maximal beta hypersynchrony are currently matters of debate. Additionally, the effect of STN HF DBS on neural synchrony in functionally connected regions of motor cortex is unknown and of great interest. Scalp EEG studies demonstrated that stimulation of the STN can activate motor cortex antidromically, but the spatial specificity of this effect has not been examined. The present study examined the effect of STN HF DBS on neural synchrony within the cortico-basal ganglia network in patients with PD. We measured local field potentials dorsal to and within the STN of PD patients, and additionally in the motor cortex in a subset of these patients. We used diffusion tensor imaging (DTI) to guide the placement of subdural cortical surface electrodes over the DTI-identified origin of the hyperdirect pathway between motor cortex and the STN. The results demonstrated that local beta power was attenuated during HF DBS both dorsal to and within the STN. The degree of attenuation was monotonic with increased DBS voltages in both locations, but this voltage-dependent effect was greater in the central STN than dorsal to the STN (p < 0.05). Cortical signals over the estimated origin of the hyperdirect pathway also demonstrated attenuation of beta hypersynchrony during DBS dorsal to or within STN, whereas signals from non-specific regions of motor cortex were not attenuated. The spatially specific suppression of beta synchrony in the motor cortex support the hypothesis that DBS may treat Parkinsonism by reducing excessive synchrony in the functionally connected sensorimotor network.

The hippocampus and imagining the future: where do we stand?

Abstract: Recent neuroimaging work has demonstrated that the hippocampus is engaged when imagining the future, in some cases more than when remembering the past. It is possible that this hippocampal activation reflects recombining details into coherent scenarios and/or the encoding of these scenarios into memory for later use. However, inconsistent findings have emerged from recent studies of future simulation in patients with memory loss and hippocampal damage. Thus, it remains an open question as to whether the hippocampus is necessary for future simulation. In this review, we consider the findings from patient studies and the neuroimaging literature with respect to a new framework that highlights three component processes of simulation: accessing episodic details, recombining details, and encoding simulations. We attempt to reconcile these discrepancies between neuroimaging and patient studies by suggesting that different component processes of future simulation may be differentially affected by hippocampal damage.

The hippocampus and inferential reasoning: building memories to navigate future decisions

Abstract: A critical aspect of inferential reasoning is the ability to form relationships between items or events that were not experienced together. This review considers different perspectives on the role of the hippocampus in successful inferential reasoning during both memory encoding and retrieval. Intuitively, inference can be thought of as a logical process by which elements of individual existing memories are retrieved and recombined to answer novel questions. Such flexible retrieval is sub-served by the hippocampus and is thought to require specialized hippocampal encoding mechanisms that discretely code events such that event elements are individually accessible from memory. In addition to retrieval-based inference, recent research has also focused on hippocampal processes that support the combination of information acquired across multiple experiences during encoding. This mechanism suggests that by recalling past events during new experiences, connections can be created between newly formed and existing memories. Such hippocampally mediated memory integration would thus underlie the formation of networks of related memories that extend beyond direct experience to anticipate future judgments about the relationships between items and events. We also discuss integrative encoding in the context of emerging evidence linking the hippocampus to the formation of schemas as well as prospective theories of hippocampal function that suggest memories are actively constructed to anticipate future decisions and actions.

Social top-down response modulation (STORM): a model of the control of mimicry in social interaction

Abstract: As a distinct feature of human social interactions, spontaneous mimicry has been widely investigated in the past decade Research suggests that mimicry is a subtle and flexible social behavior which plays an important role for communication and affiliation However, fundamental questions like why and how people mimic still remain unclear In this paper, we evaluate past theories of why people mimic and the brain systems that implement mimicry in social psychology and cognitive neuroscience By reviewing recent behavioral and neuroimaging studies on the control of mimicry by social signals, we conclude that the subtlety and sophistication of mimicry in social contexts reflect a social top-down response modulation (STORM) which increases one's social advantage and this mechanism is most likely implemented by medial prefrontal cortex (mPFC) We suggest that this STORM account of mimicry is important for our understanding of social behavior and social cognition, and provides implications for future research in autism

The hippocampus and visual perception

Abstract: In this review, we will discuss the idea that the hippocampus may be involved in both memory and perception, contrary to theories that posit functional and neuroanatomical segregation of these processes. This suggestion is based on a number of recent neuropsychological and functional neuroimaging studies that have demonstrated that the hippocampus is involved in the visual discrimination of complex spatial scene stimuli. We argue that these findings cannot be explained by long-term memory or working memory processing or, in the case of patient findings, dysfunction beyond the medial temporal lobe (MTL). Instead, these studies point toward a role for the hippocampus in higher-order spatial perception. We suggest that the hippocampus processes complex conjunctions of spatial features, and that it may be more appropriate to consider the representations for which this structure is critical, rather than the cognitive processes that it mediates.

The hippocampus and the flexible use and processing of language

Abstract: Fundamental to all human languages is an unlimited expressive capacity and creative flexibility that allow speakers to rapidly generate novel and complex utterances. In turn, listeners interpret language “on-line”, incrementally integrating multiple sources of information as words unfold over time. A challenge for theories of language processing has been to understand how speakers and listeners generate, gather, integrate, and maintain representations in service of language processing. We propose that many of the processes by which we use language place high demands on and receive contributions from the hippocampal declarative memory system. The hippocampal declarative memory system is long known to support relational binding and representational flexibility. Recent findings demonstrate that these same functions are engaged during the real-time processes that support behavior in-the-moment. Such findings point to the hippocampus as a potentially key contributor to cognitive functions that require on-line integration of multiple sources of information, such as on-line language processing. Evidence supporting this view comes from findings that individuals with hippocampal amnesia show deficits in the use of language flexibly and on-line. We conclude that the relational binding and representational flexibility afforded by the hippocampal declarative memory system positions the hippocampus as a key contributor to language use and processing.

rTMS for Tinnitus.

Abstract: I applaud the authors of the article “Can temporal repetitive transcranial magnetic stimulation be enhanced by targeting affective components of tinnitus with frontal rTMS? a randomized controlled pilot trial” because many of them are pioneers in the development of protocols that use rTMS to treat chronic tinnitus. Investigators (including myself) around the world conducting similar studies of rTMS for tinnitus often follow the procedural examples of these researcher–clinicians in Germany. In this recent publication, Kreuzer et al. (2011) compared the effects of 2000 daily pulses of rTMS delivered to patients’ left temporal region with a protocol that began with 1000 pulses of right frontal rTMS followed by 1000 pulses of left temporal rTMS. In both protocols, patients exhibited improvement in measures of tinnitus severity. However, there was no significant difference in treatment outcomes between the left temporal and the combined (right frontal + left temporal) rTMS protocols. I would like to add a few comments about this study: (1) No placebo condition was included. Because the placebo effect can be significant for many tinnitus patients (Dobie, 1999), a placebo condition should be used in most clinical trials involving this population. (2) The fact that patients’ scores for the Tinnitus Questionnaire, THI, and BDI all declined from the time of screening to the time just before the first rTMS session (baseline) illustrates, I believe, the placebo effect (or, as the authors describe it, “an anticipation effect”). This decline in scores from screening to baseline is similar in magnitude to the decline in scores that occurred during rTMS treatment. (3) The authors state that they used a left temporal coil placement for rTMS (regardless of the location of patients’ tinnitus perception) in order to compare results with previous studies that used the same coil placement. However, because Frank et al. (2010) reported that left temporal rTMS was not effective for patients with right-side tinnitus, it is time for researchers to re-evaluate rTMS coil placement in experimental designs. All of us conducting rTMS studies for tinnitus realize that many procedural questions need to be addressed before this treatment option can reach its full clinical potential. We thank our colleagues in Germany for continuing to conduct studies that increase our understanding and contribute to improvements in this area of research.