Increasing and decreasing interregional brain coupling increases and decreases oscillatory activity in the human brain.

The origins of oscillatory activity in the brain are currently debated, but common to many hypotheses is the notion that they reflect interactions between brain areas. Here, we examine this possibility by manipulating the strength of coupling between two human brain regions, ventral premotor cortex (PMv) and primary motor cortex (M1), and examine the impact on oscillatory activity in the motor system measurable in the electroencephalogram. We either increased or decreased the strength of coupling while holding the impact on each component area in the pathway constant. This was achieved by stimulating PMv and M1 with paired pulses of transcranial magnetic stimulation using two different patterns, only one of which increases the influence exerted by PMv over M1. While the stimulation protocols differed in their temporal patterning, they were comprised of identical numbers of pulses to M1 and PMv. We measured the impact on activity in alpha, beta, and theta bands during a motor task in which participants either made a preprepared action (Go) or withheld it (No-Go). Augmenting cortical connectivity between PMv and M1, by evoking synchronous pre- and postsynaptic activity in the PMv-M1 pathway, enhanced oscillatory beta and theta rhythms in Go and No-Go trials, respectively. Little change was observed in the alpha rhythm. By contrast, diminishing the influence of PMv over M1 decreased oscillatory beta and theta rhythms in Go and No-Go trials, respectively. This suggests that corticocortical communication frequencies in the PMv-M1 pathway can be manipulated following Hebbian spike-timing-dependent plasticity.

Affective interoceptive inference: Evidence from heart-beat evoked brain potentials.

The perception of internal bodily signals (interoception) is central to many theories of emotion and embodied cognition. According to recent theoretical views, the sensory processing of visceral signals such as one's own heartbeat is determined by top-down predictions about the expected interoceptive state of the body (interoceptive inference). In this EEG study we examined neural responses to heartbeats following expected and unexpected emotional stimuli. We used a modified stimulus repetition task in which pairs of facial expressions were presented with repeating or alternating emotional content, and we manipulated the emotional valence and the likelihood of stimulus repetition. We found that affective predictions of external socially relevant information modulated the heartbeat-evoked potential, a marker of cardiac interoception. Crucially, the HEP changes highly relied on the expected emotional content of the facial expression. Thus, expected negative faces led to a decreased HEP amplitude, whereas such an effect was not observed after an expected neutral face. These results suggest that valence-specific affective predictions, and their uniquely associated predicted bodily sensory state, can reduce or amplify cardiac interoceptive responses. In addition, the affective repetition effects were dependent on repetition probability, highlighting the influence of top-down exteroceptive predictions on interoception. Our results are in line with recent models of interoception supporting the idea that predicted bodily states influence sensory processing of salient external information.

Heartfelt Self: Cardio-Visual Integration Affects Self-Face Recognition and Interoceptive Cortical Processing.

The sense of body-ownership relies on the representation of both interoceptive and exteroceptive signals coming from one's body. However, it remains unknown how the integration of bodily signals coming from "outside" and "inside" the body is instantiated in the brain. Here, we used a modified version of the Enfacement Illusion to investigate whether the integration of visual and cardiac information can alter self-face recognition (Experiment 1) and neural responses to heartbeats (Experiment 2). We projected a pulsing shade, that was synchronous or asynchronous with the participant's heartbeat, onto a picture depicting the participant's face morphed with the face of an unfamiliar other. Results revealed that synchronous (vs. asynchronous) cardio-visual stimulation led to increased self-identification with the other's face (Experiment 1), while during stimulation, synchronicity modulated the amplitude of the Heartbeat Evoked Potential, an electrophysiological index of cortical interoceptive processing (Experiment 2). Importantly, the magnitude of the illusion-related effects was dependent on, and increased linearly, with the participants' Interoceptive Accuracy. These results provide the first direct neural evidence for the integration of interoceptive and exteroceptive signals in bodily self-awareness.

Electrophysiological correlates of self-specific prediction errors in the human brain.

Recognising one's self, vs. others, is a key component of self-awareness, crucial for social interactions. Here we investigated whether processing self-face and self-body images can be explained by the brain's prediction of sensory events, based on regularities in the given context. We measured evoked cortical responses while participants observed alternating sequences of self-face or other-face images (experiment 1) and self-body or other-body images (experiment 2), which were embedded in an identity-irrelevant task. In experiment 1, the expected sequences were violated by deviant morphed images, which contained 33%, 66% or 100% of the self-face when the other's face was expected (and vice versa). In experiment 2, the anticipated sequences were violated by deviant images of the self when the other's image was expected (and vice versa), or by two deviant images composed of pictures of the self-face attached to the other's body, or the other's face attached to the self-body. This manipulation allowed control of the prediction error associated with the self or the other's image. Deviant self-images (but not deviant images of the other) elicited a visual mismatch response (vMMR)--a cortical index of violations of regularity. This was source localised to face and body related visual, sensorimotor and limbic areas and had amplitude proportional to the amount of deviance from the self-image. We provide novel evidence that self-processing can be described by the brain's prediction error system, which accounts for self-bias in visual processing. These findings are discussed in the light of recent predictive coding models of self-processing.

The emotional homunculus: ERP evidence for independent somatosensory responses during facial emotional processing.

Current models of face perception propose that initial visual processing is followed by activation of nonvisual somatosensory areas that contributes to emotion recognition. To test whether there is a pure and independent involvement of somatosensory cortex (SCx) during face processing over and above visual responses, we directly measured participants' somatosensory-evoked activity by tactually probing (105 ms postvisual facial stimuli) the state of SCx during an emotion discrimination task while controlling for visual effects. Discrimination of emotional versus neutral expressions enhanced early somatosensory-evoked activity between 40 and 80 ms after stimulus onset, suggesting visual emotion processing in SCx. This effect was source localized within primary, secondary, and associative somatosensory cortex. Emotional face processing influenced somatosensory responses to both face (congruent body part) and finger (control site) tactile stimulation, suggesting a general process that includes nonfacial cortical representations. Gender discrimination of the same facial expressions did not modulate somatosensory-evoked activity. We provide novel evidence that SCx activation is not a byproduct of visual processing but is independently shaped by face emotion processing.

Timing along the cardiac cycle modulates neural signals of reward-based learning.

Natural fluctuations in cardiac activity modulate brain activity associated with sensory stimuli, as well as perceptual decisions about low magnitude, near-threshold stimuli. However, little is known about the relationship between fluctuations in heart activity and other internal representations. Here we investigate whether the cardiac cycle relates to learning-related internal representations - absolute and signed prediction errors. We combined machine learning techniques with electroencephalography with both simple, direct indices of task performance and computational model-derived indices of learning. Our results demonstrate that just as people are more sensitive to low magnitude, near-threshold sensory stimuli in certain cardiac phases, so are they more sensitive to low magnitude absolute prediction errors in the same cycles. However, this occurs even when the low magnitude prediction errors are associated with clearly suprathreshold sensory events. In addition, participants exhibiting stronger differences in their prediction error representations between cardiac cycles exhibited higher learning rates and greater task accuracy.

Do non-traumatic stressful life events and ageing negatively impact working memory performance and do they interact to further impair working memory performance?

Stress and normal ageing produce allostatic load, which may lead to difficulties with cognition thereby degrading quality of life. The current study's objective was to assess whether ageing and cumulative stress interact to accelerate cognitive decline. With 60 participants, Marshall et al. found that ageing and cumulative stress interact significantly to impair working memory performance in older adults, suggesting vulnerability to the cumulative effects of life events beyond 60 years old. To replicate and extend this finding, we increased the sample size by conducting 3 independent studies with 156 participants and improved the statistical methods by conducting an iterative Bayesian meta-analysis with Bayes factors. Bayes factors deliver a more comprehensive result because they provide evidence for either the null hypothesis (H0), the alternative hypothesis (H1) or for neither hypothesis due to evidence not being sufficiently sensitive. Young (18-35 yrs) and older (60-85 yrs) healthy adults were categorised as high or low stress based on their life events score derived from the Life Events Scale for Students or Social Readjustment Rating Scale, respectively. We measured accuracy and reaction time on a 2-back working memory task to provide: a) Bayes factors and b) Bayesian meta-analysis, which iteratively added each study's effect sizes to evaluate the overall strength of evidence that ageing, cumulative stress and/or the combination of the two detrimentally affect working memory performance. Using a larger sample (N = 156 vs. N = 60) and a more powerful statistical approach, we did not replicate the robust age by cumulative stress interaction effect found by Marshall et al.. The effects of ageing and cumulative stress also fell within the anecdotal range (⅓

The social readjustment rating scale: Updated and modernised.

The Social Readjustment Rating Scale, originally devised in 1967 by Holmes and Rahe, measures the impact of life events stress. At the time, the SRRS advanced its field of research by standardising the impact of stress with a set of independently derived weights called 'life change units' (LCUs) for 43 life events found to predict illness onset. The scale has been criticised for being outdated, e.g. "Mortgage over $10,000" and biased, e.g. "Wife begin or stop work". The aim of this cross-sectional survey study is to update and improve the SRRS whilst allowing backwards compatibility. We successfully updated the SRRS norms/LCUs using the ratings of 540 predominantly UK adults aged 18 to 84. Moreover, we also updated wording of 12 SRRS items and evaluated the impact of demographics, personal experience and loneliness. Using non-parametric frequentist and Bayesian statistics we found that the updated weights were higher but broadly consistent with those of the original study. Furthermore, changes to item wording did not affect raters' evaluations relative to the original thereby ensuring cross-comparability with the original SRRS. The raters were not unduly influenced by their personal experiences of events nor loneliness. The target sample was UK rather than US-based and was proportionately representative regarding age, sex and ethnicity. Moreover, the age range was broader than the original SRRS. In addition, we modernised item wording, added one optional extra item to the end of the scale to evaluate the readjustment to living alone and identified 3 potential new items proposed by raters. Backwards-compatibility is maintained.

Changing connectivity between premotor and motor cortex changes inter-areal communication in the human brain.

The ventral premotor cortex (PMv) is an important component of cortico-cortical pathways mediating prefrontal control over primary motor cortex (M1) function. Paired associative stimulation (ccPAS) is known to change PMv influence over M1 in humans, which manifests differently depending on the behavioural context. Here we show that these changes in influence are functionally linked to PMv-M1 phase synchrony changes induced by repeated paired stimulation of the two areas. PMv-to-M1 ccPAS leads to increased phase synchrony in alpha and beta bands, while reversed order M1-to-PMv ccPAS leads to decreased theta phase synchrony. These changes are visible at rest but are predictive of changes in oscillatory power in the same frequencies during movement execution and inhibition, respectively. The results unveil a link between the physiology of the motor network and the resonant frequencies mediating its interactions and provide a putative mechanism underpinning the relationship between synaptic efficacy and brain oscillations.

Increasing interhemispheric connectivity between human visual motion areas uncovers asymmetric sensitivity to horizontal motion.

Our conscious perceptual experience relies on a hierarchical process involving integration of low-level sensory encoding and higher-order sensory selection.1 This hierarchical process may scale at different levels of brain functioning, including integration of information between the hemispheres.2-5 Here, we test this hypothesis for the perception of visual motion stimuli. Across 3 experiments, we manipulated the connectivity between the left and right visual motion complexes (V5/MT+) responsible for horizontal motion perception2,3 by means of transcranial magnetic stimulation (TMS).4,5 We found that enhancing the strength of connections from the left to the right V5/MT+, by inducing spike-timing-dependent plasticity6 in this pathway, increased sensitivity to horizontal motion. These changes were present immediately and lasted at least 90 min after intervention. Notably, little perceptual changes were observed when strengthening connections from the right to the left V5/MT+. Furthermore, we found that this asymmetric modulation was mirrored by an asymmetric perceptual bias in the direction of the horizontal motion. Overall, observers were biased toward leftward relative to rightward motion direction. Crucially, following the strengthening of the connections from right to left V5/MT+, this bias could be momentarily reversed. These results suggest that the projections connecting left and right V5/MT+ in the human visual cortex are asymmetrical, subtending a hierarchical role of hemispheric specialization7-10 favoring left-to-right hemisphere processing for integrating local sensory input into coherent global motion perception.

Causal manipulation of self-other mergence in the dorsomedial prefrontal cortex.

To navigate social environments, people must simultaneously hold representations about their own and others' abilities. During self-other mergence, people estimate others' abilities not only on the basis of the others' past performance, but the estimates are also influenced by their own performance. For example, if we perform well, we overestimate the abilities of those with whom we are co-operating and underestimate competitors. Self-other mergence is associated with specific activity patterns in the dorsomedial prefrontal cortex (dmPFC). Using a combination of non-invasive brain stimulation, functional magnetic resonance imaging, and computational modeling, we show that dmPFC neurostimulation silences these neural signatures of self-other mergence in relation to estimation of others' abilities. In consequence, self-other mergence behavior increases, and our assessments of our own performance are projected increasingly onto other people. This suggests an inherent tendency to form interdependent social representations and a causal role of the dmPFC in separating self and other representations.

The somatotopy of observed emotions.

The ability to experience others' emotional states is a key component in social interactions. Uniquely among sensorimotor regions, the somatosensory cortex (SCx) plays an especially important role in human emotion understanding. While distinct emotions are experienced in specific parts of the body, it remains unknown whether the SCx exhibits somatotopic activations to different emotional expressions. In the current study, we investigated if the affective response triggered by observing others' emotional face expressions leads to differential activations in SCx. Participants performed a visual facial emotion discrimination task while we measured changes in SCx topographic EEG activity by tactually stimulating two body-parts representative of the upper and lower limbs, the finger and the toe respectively. The results of the study showed an emotion specific response in the finger SCx when observing angry as opposed to sad emotional expressions, after controlling for carry-over effects of visual evoked activity. This dissociation to observed emotions was not present in toe somatosensory responses. Our results suggest that somatotopic activations of the SCx to discrete emotions might play a crucial role in understanding others' emotions.

When you smile, the world smiles at you: ERP evidence for self-expression effects on face processing.

Current models of emotion simulation propose that intentionally posing a facial expression can change one's subjective feelings, which in turn influences the processing of visual input. However, the underlying neural mechanism whereby one's facial emotion modulates the visual cortical responses to other's facial expressions remains unknown. To understand how one's facial expression affects visual processing, we measured participants' visual evoked potentials (VEPs) during a facial emotion judgment task of positive and neutral faces. To control for the effects of facial muscles on VEPs, we asked participants to smile (adopting an expression of happiness), to purse their lips (incompatible with smiling) or to pose with a neutral face, in separate blocks. Results showed that the smiling expression modulates face-specific visual processing components (N170/vertex positive potential) to watching other facial expressions. Specifically, when making a happy expression, neutral faces are processed similarly to happy faces. When making a neutral expression or pursing the lips, however, responses to neutral and happy face are significantly different. This effect was source localized within multisensory associative areas, angular gyrus, associative visual cortex and somatosensory cortex. We provide novel evidence that one's own emotional expression acts as a top-down influence modulating low-level neural encoding during facial perception.

[Neuroarchitecture of musical emotions]. / Neuroarquitectura de la emocion musical.

The emotional response to music, or musical emotion, is a universal response that draws on diverse psychological processes implemented in a large array of neural structures and mechanisms. Studies using electroencephalography, functional magnetic resonance, lesions and individuals with extent musical training have begun to elucidate some of these mechanisms. The objective of this article is reviewing the most relevant studies that have tried to identify the neural correlates of musical emotion from the more automatic to the more complex processes, and to understand how these correlates interact in the brain. The article describes how the presentation of music perceived as emotional is associated with a rapid autonomic response in thalamic and subthalamic structures, accompanied by changes in the electrodermal and endocrine responses. It also explains how musical emotion processing activates auditory cortex, as well as a series of limbic and paralimbic structures, such as the amygdala, the anterior cingulate cortex or the hippocampus, demonstrating the relevant contribution of the limbic system to musical emotion. Further, it is detailed how musical emotion depends to a great extent on semantic and syntactic process carried out in temporal and parietofrontal areas, respectively. Some of the recent works demonstrating that musical emotion highly relies on emotional simulation are also mentioned. Finally, a summary of these studies, their limitations, and suggestions for further research on the neuroarchitecture of musical emotion are given.

The sacred and the absurd--an electrophysiological study of counterintuitive ideas (at sentence level).

Religious beliefs are both catchy and durable: they exhibit a high degree of adherence to our cognitive system, given their success of transmission and spreading throughout history. A prominent explanation for religion's cultural success comes from the "MCI hypothesis," according to which religious beliefs are both easy to recall and desirable to transmit because they are minimally counterintuitive (MCI). This hypothesis has been empirically tested at concept and narrative levels by recall measures. However, the neural correlates of MCI concepts remain poorly understood. We used the N400 component of the event-related brain potential as a measure of counterintuitiveness of violations comparing religious and non-religious sentences, both counterintuitive, when presented in isolation. Around 80% in either condition were core-knowledge violations. We found smaller N400 amplitudes for religious as compared to non-religious counterintuitive ideas, suggesting that religious ideas are less semantically anomalous. Moreover, behavioral measures revealed that religious ideas are not readily detected as unacceptable. Finally, systematic analyses of our materials, according to conceptual features proposed in cognitive models of religion, did not reveal any outstanding variable significantly contributing to these differences. Refinements of cognitive models of religion should elucidate which combination of factors renders an anomaly less counterintuitive and thus more suitable for recall and transmission.

How the emotional content of discourse affects language comprehension.

Emotion effects on cognition have often been reported. However, only few studies investigated emotional effects on subsequent language processing, and in most cases these effects were induced by non-linguistic stimuli such as films, faces, or pictures. Here, we investigated how a paragraph of positive, negative, or neutral emotional valence affects the processing of a subsequent emotionally neutral sentence, which contained either semantic, syntactic, or no violation, respectively, by means of event-related brain potentials (ERPs). Behavioral data revealed strong effects of emotion; error rates and reaction times increased significantly in sentences preceded by a positive paragraph relative to negative and neutral ones. In ERPs, the N400 to semantic violations was not affected by emotion. In the syntactic experiment, however, clear emotion effects were observed on ERPs. The left anterior negativity (LAN) to syntactic violations, which was not visible in the neutral condition, was present in the negative and positive conditions. This is interpreted as reflecting modulatory effects of prior emotions on syntactic processing, which is discussed in the light of three alternative or complementary explanations based on emotion-induced cognitive styles, working memory, and arousal models. The present effects of emotion on the LAN are especially remarkable considering that syntactic processing has often been regarded as encapsulated and autonomous.

How is sentence processing affected by external semantic and syntactic information? Evidence from event-related potentials.

BACKGROUND: A crucial question for understanding sentence comprehension is the openness of syntactic and semantic processes for other sources of information. Using event-related potentials in a dual task paradigm, we had previously found that sentence processing takes into consideration task relevant sentence-external semantic but not syntactic information. In that study, internal and external information both varied within the same linguistic domain-either semantic or syntactic. Here we investigated whether across-domain sentence-external information would impact within-sentence processing. METHODOLOGY: In one condition, adjectives within visually presented sentences of the structure [Det]-[Noun]-[Adjective]-[Verb] were semantically correct or incorrect. Simultaneously with the noun, auditory adjectives were presented that morphosyntactically matched or mismatched the visual adjectives with respect to gender. FINDINGS: As expected, semantic violations within the sentence elicited N400 and P600 components in the ERP. However, these components were not modulated by syntactic matching of the sentence-external auditory adjective. In a second condition, syntactic within-sentence correctness-variations were combined with semantic matching variations between the auditory and the visual adjective. Here, syntactic within-sentence violations elicited a LAN and a P600 that did not interact with semantic matching of the auditory adjective. However, semantic mismatching of the latter elicited a frontocentral positivity, presumably related to an increase in discourse level complexity. CONCLUSION: The current findings underscore the open versus algorithmic nature of semantic and syntactic processing, respectively, during sentence comprehension.

Encouraging expressions affect the brain and alter visual attention.

BACKGROUND: Very often, encouraging or discouraging expressions are used in competitive contexts, such as sports practice, aiming at provoking an emotional reaction on the listener and, consequently, an effect on subsequent cognition and/or performance. However, the actual efficiency of these expressions has not been tested scientifically. METHODOLOGY/PRINCIPAL FINDINGS: To fill this gap, we studied the effects of encouraging, discouraging, and neutral expressions on event-related brain electrical activity during a visual selective attention task in which targets were determined by location, shape, and color. Although the expressions preceded the attentional task, both encouraging and discouraging messages elicited a similar long-lasting brain emotional response present during the visuospatial task. In addition, encouraging expressions were able to alter the customary working pattern of the visual attention system for shape selection in the attended location, increasing the P1 and the SP modulations while simultaneously fading away the SN. CONCLUSIONS/SIGNIFICANCE: This was interpreted as an enhancement of the attentional processes for shape in the attended location after an encouraging expression. It can be stated, therefore, that encouraging expressions, as those used in sport practice, as well as in many other contexts and situations, do seem to be efficient in exerting emotional reactions and measurable effects on cognition.

Neuroarquitectura de la emoción musical / Neuroarchitecture of musical emotions

La respuesta emocional ante la música, o emoción musical, es una respuesta universal que depende de diferentes procesos psicológicos y recluta una extensa red de estructuras neuronales. Mediante el empleo de técnicas como la electroencefalografía, la resonancia magnética funcional y los estudios con población clínica e individuos con formación musical previa, se ha empezado a dilucidar estos mecanismos cerebrales. El objetivo de este artículo es hacer una revisión de los trabajos más relevantes en los que se identifican los correlatos neuronales de la emoción musical, desde los procesos más automáticos hasta los más complejos, y comprender cómo interaccionan en el cerebro. En concreto, se describe cómo la presentación de música emocional está asociada a una respuesta rápida en estructuras talámicas y subtalámicas, acompañada por cambios electrodérmicos y endocrinos. También se explica que el procesamiento de la emoción musical implica la activación de la corteza auditiva y estructuras límbicas y paralímbicas, como la amígdala, la corteza cinguladaanterior o el hipocampo, lo que demuestra la contribución del sistema límbico a la emoción musical. Asimismo, se detalla cómo la emoción musical depende de los significados semántico y sintáctico de la música, procesados en áreas temporales y parietofrontales, respectivamente. Además, se mencionan trabajos recientes que han demostrado cómo los mecanismos de simulación emocional también contribuyen a la emoción musical. Por último, se hace un resumen de estos trabajos, comentando sus limitaciones y ofreciendo alternativas para seguir avanzando en el estudio de la neuroarquitectura de la emoción musical (AU)

The emotional response to music, or musical emotion, is a universal response that draws on diverse psychological processes implemented in a large array of neural structures and mechanisms. Studies using electroencephalography, functional magnetic resonance, lesions and individuals with extent musical training have begun to elucidate some of these mechanisms. The objective of this article is reviewing the most relevant studies that have tried to identify the neural correlates of musical emotion from the more automatic to the more complex processes, and to understand how these correlates interact in the brain. The article describes how the presentation of music perceived as emotional is associated with a rapid autonomic response in thalamic and subthalamic structures, accompanied by changes in the electrodermal and endocrine responses. It also explains how musical emotion processing activates auditory cortex, as well as a series of limbic and paralimbic structures, such as the amygdala, the anterior cingulate cortex or the hippocampus, demonstrating the relevant contribution of the limbic system to musical emotion. Further, it is detailed how musical emotion depends to a great extent on semantic and syntactic process carried out in temporal and parietofrontal areas, respectively. Some of the recent works demonstrating that musical emotion highly relies on emotional simulation are also mentioned. Finally, a summary of these studies, their limitations, and suggestions for further research on the neuroarchitecture of musical emotion are given (AU)