Emotion and motor preparation: A transcranial magnetic stimulation study of corticospinal motor tract excitability

In the present study, we examined whether preparing motor responses under different emotional conditions alters motor evoked potentials (MEPs) elicited by transcranial magnetic stimulation delivered to the motor cortex. Analyses revealed three findings: (1) Reaction times were expedited during exposure to unpleasant images, as compared with pleasant and neutral images; (2) force amplitude was greater during exposure to unpleasant images, as compared with pleasant and neutral images; and (3) MEPs were larger while participants viewed unpleasant images, as compared with neutral images. Hence, coupling the preparation of motor responses with the viewing of emotional images led to arousal-driven changes in corticospinal motor tract excitability, whereas movement speed and force production varied as a function of emotional valence. These findings demonstrate that the effects of emotion on the motor system manifest at varying sensitivity levels across behavioral and neurophysiological measures. Moreover, they validate the action readiness component of emotional experience by demonstrating that emotional states influence the execution of future movements but, alone, do not lead to overt movement.     

Negative emotional processing induced by spoken scenarios modulates corticospinal excitability

Emotion is assumed to facilitate the preparation of behavioral responses to environmental stimuli. In the present study, we examined whether emotional processing induced by spoken scenarios of positive and negative content, related to the self or to other people, modulates corticospinal excitability. Motor evoked potentials (MEPs) elicited by transcranial magnetic stimulation (TMS) in 20 volunteers indicated that processing of negative emotional content increased MEP amplitude, regardless of the perspective taken in the scenario. By contrast, positive emotional processing did not reliably alter MEP amplitude. These results provide the first TMS evidence that the auditory processing of emotionally negative information triggers action preparation.     

Musical groove modulates motor cortex excitability: A TMS investigation

Groove is often described as a musical quality that can induce movement in a listener. This study examines the effects of listening to groove music on corticospinal excitability. Musicians and non-musicians listened to high-groove music, low-groove music, and spectrally matched noise, while receiving single-pulse transcranial magnetic stimulation (TMS) over the primary motor cortex either on-beat or off-beat. We examined changes in the amplitude of the motor-evoked potentials (MEPs), recorded from hand and arm muscles, as an index of activity within the motor system. Musicians and non-musicians rated groove similarly. MEP results showed that high-groove music modulated corticospinal excitability, whereas no difference occurred between low-groove music and noise. More specifically, musicians’ MEPs were larger with high-groove than low-groove music, and this effect was especially pronounced for on-beat compared to off-beat pulses. These results indicate that high-groove music increasingly engages the motor system, and the temporal modulation of corticospinal excitability with the beat could stem from tight auditory–motor links in musicians. Conversely, non-musicians’ MEPs were smaller for high-groove than low-groove music, and there was no effect of on- versus off-beat pulses, potentially stemming from suppression of overt movement. In sum, high-groove music engages the motor system, and previous training modulates how listening to music with a strong groove activates the motor system.     

Catechol-O-methyltransferase Val158Met genotype affects neural correlates of aversive stimuli processing

It was previously shown that variation of the catechol-O-methyltransferase (COMT) gene modulates brain activity during the processing of stimuli with negative valence, but not for pleasant stimuli. Here, we tested whether the COMT genotype also modulates the electrophysiological correlates of emotional processing and explored whether the environmental factor of life stress influences this effect. Using the early posterior negativity (EPN) paradigm, event-related brain potentials were measured in 81 healthy individuals during the processing of pictures that evoked emotions of positive and negative valence. As was hypothesized, the COMT genotype affected the EPN amplitudes for unpleasant stimuli, but not for pleasant ones. Specifically, Met/Met carriers respond more sensitively to unpleasant stimuli, as compared with Val/Val carriers. We did not find evidence that life stress moderates the effect of the COMT genotype on emotional stimuli processing.     

Unpleasant stimuli differentially modulate inhibitory processes in an emotional Go/NoGo task: an event-related potential study

Threat stimuli typically elicit a psychophysiological response pattern supporting the organism's preparation for active defence. Differently, blood stimuli prompt a distinctive autonomic response pattern and sustained processing, which do not call for clear-cut mobilisation for action. However, the contribution of motor disposition in these response patterns remains unclear. One way to address this issue is to investigate whether threat and blood stimuli differentially affect the active suppression of an ongoing motor activity. Thirty-two undergraduates were presented with threat, mutilation, pleasant, and neutral pictures in an emotional Go/NoGo task. The amplitudes of the NoGo-N2 and NoGo-P3 components of the event-related potentials were analysed as indices of conflict monitoring and inhibition of motor response, respectively. Reaction times to Go trials were significantly faster for threat than for mutilations. The NoGo-N2 was significantly larger to threat than to mutilations, whereas the NoGo-P3amplitude did not differ between the two conditions. These findings suggest that threat stimuli facilitated the execution of a prepotent response and enhanced conflict monitoring when action must be withheld. In contrast, blood stimuli did not either promote action in the Go trials or increase conflict in the NoGo condition, suggesting a response pattern compatible with defensive immobility.     

Motor excitability during movement preparation in Tourette syndrome

Tourette syndrome (TS) is a neurodevelopmental disorder characterized by the occurrence of motor and vocal tics. TS has been linked to the impaired operation of cortical‐striatal‐thalamic‐cortical circuits that give rise to hyper‐excitability of cortical motor areas, which may be exacerbated by dysfunctional intra‐cortical inhibitory mechanisms. That said, many individuals gain control over their tics during adolescence and it has been suggested that this increased control arises as a result of the development of mechanisms that operate to suppress corticospinal excitability (CSE) ahead of volitional movements. Here we used single‐pulse transcranial magnetic stimulation (TMS) in conjunction with a manual Go/NoGo task to investigate alterations in CSE ahead of volitional movements in a group of adolescents with TS (N = 10). Our study demonstrated that CSE, as measured by TMS‐induced motor‐evoked potentials (MEPs), was significantly reduced in the TS group in the period immediately preceding a finger movement. More specifically, we show that individuals with TS, unlike their age‐matched controls, do not exhibit the predicted increase in mean MEP amplitude and decrease in MEP variability that immediately precede the execution of volitional movements in typically developing young adults. Finally, we report that the magnitude of the rise in MEP amplitude across the movement preparation period in TS is significantly negatively correlated with clinical measures of motor tic severity, suggesting that individuals with severe motor tics are least able to modulate motor cortical excitability.     

Accelerative and decelerative effects of hedonic valence and emotional arousal during visual scene processing

Perceptual processing of natural scene pictures is enhanced when the scene conveys emotional content. Such “motivated attention” to pleasant and unpleasant pictures has been shown to improve identification accuracy in non-speeded behavioural tasks. An open question is whether emotional content also modulates the speed of visual scene processing. In the present studies we show that unpleasant content reliably slowed two-choice categorization of pictures, irrespective of physical image properties, perceptual complexity, and categorization instructions. Conversely, pleasant content did not slow or even accelerated choice reactions, relative to neutral scenes. As indicated by lateralized readiness potentials, these effects occurred at cognitive processing rather than motor preparation/execution stages. Specifically, analysis of event-related potentials showed a prolongation of early scene discrimination for stimuli perceived as emotionally arousing, regardless of valence, and reflected in delayed peaks of the N1 component. In contrast, the timing of other processing steps, reflected in the P2 and late positive potential components and presumably related to post-discriminatory processes such as stimulus–response mapping, appeared to be determined by hedonic valence, with more pleasant scenes eliciting faster processing. Consistent with this model, varying arousal (low/high) within the emotional categories mediated the effects of valence on choice reaction speed. Functionally, arousal may prolong stimulus analysis in order to prevent erroneous and potentially harmful decisions. Pleasantness may act as a safety signal allowing rapid initiation of overt responses.     

Event-related potentials of emotional memory: encoding pleasant,unpleasant, and neutral pictures

Emotional events tend to be remembered better than nonemotional events. We investigated this phenomenon by measuring two event-related potential (ERP) effects: the emotion effect (more positive ERPs for pleasant or unpleasant stimuli than for neutral stimuli) and the subsequent memory effect (more positive ERPs for subsequently remembered items than for subsequently forgotten items). ERPs were measured while subjects rated the emotional content of pleasant, unpleasant, and neutral pictures. As was expected, subsequent recall was better for pleasant and unpleasant pictures than for neutral pictures. The emotion effect was sensitive to arousal in parietal electrodes and to both arousal and valence in frontocentral electrodes. The subsequent memory effect at centroparietal electrodes was greater for emotional pictures than for neutral pictures during an early epoch (400-600 msec). This result suggests that emotional information has privileged access to processing resources, possibly leading to better memory formation.     

Electromyographic activity,H-reflex modulation and corticospinal input to forearm motoneurones during active and passive rhythmic movements

Four subjects produced coordinated movements, consisting of flexion and extension of the wrist in ipsilateral (right wrist only), contralateral (left wrist only), inphase (both wrists in flexion or both in extension) and antiphase (one wrist in flexion, the other in extension) conditions. Electromyographic (EMG) activity was recorded from right wrist flexor and extensor muscles. In one session, transcranial magnetic stimuli (TMS) of the left motor cortex, around threshold intensity, evoked short-latency responses in the right wrist extensors and flexors. In another session, the median nerve at the cubital fossa was stimulated to elicit an H-reflex in the right flexor carpi radialis (rFCR). A movement cycle was divided into 8 segments. In total, 10 identical stimuli were delivered during each segment in each condition, at two movement frequencies. The magnitude of the EMG reponses to TMS was modulated markedly during movements made in the ipsilateral condition, and in both bimanual conditions. EMG activity was greater, and motor-evoked potentials (MEPs) were larger in the antiphase condition than in the inphase condition. When the amplitudes of the MEPs were normalised with respect to background EMG, no significant differences between the bimanual conditions were obtained. For H-reflexes, significant differences between the two bimanual conditions were observed, suggesting differences in levels of excitability of the Ia afferent pathway. These differences were attributed to segmental input associated with changes in muscle length arising from limb movement, and upon descending input to the spinal cord, possibly mediated by Renshaw cell inhibition. During rhythmic passive movement of the right limb, H-reflexes were inhibited and MEPs potentiated in a cyclic fashion. Passive movement of the contralateral left limb resulted in inhibition of both responses.PsycINFO classification: 2330; 2530; 2540     

Emotional facilitation of sensory processing in the visual cortex

A key function of emotion is the preparation for action. However, organization of successful behavioral strategies depends on efficient stimulus encoding. The present study tested the hypothesis that perceptual encoding in the visual cortex is modulated by the emotional significance of visual stimuli. Event-related brain potentials were measured while subjects viewed pleasant, neutral, and unpleasant pictures. Early selective encoding of pleasant and unpleasant images was associated with a posterior negativity, indicating primary sources of activation in the visual cortex. The study also replicated previous findings in that affective cues also elicited enlarged late positive potentials, indexing increased stimulus relevance at higher-order stages of stimulus processing. These results support the hypothesis that sensory encoding of affective stimuli is facilitated implicitly by natural selective attention. Thus, the affect system not only modulates motor output (i.e., favoring approach or avoidance dispositions), but already operates at an early level of sensory encoding.     

Depth of facial expression processing depends on stimulus visibility: Behavioral and electrophysiological evidence of priming effects

Participants performed a priming task during which emotional faces served as prime stimuli and emotional words served as targets. Prime-target pairs were congruent or incongruent, and two levels of prime visibility were obtained by varying the duration of the masked primes. To probe a neural signature of the impact of the masked primes, lateralized readiness potentials (LRPs) were recorded over motor cortex. In the high-visibility condition, responses to word targets were faster when the prime-target pairs were congruent than when they were incongruent, providing evidence of priming effects. In line with the behavioral results, the electrophysiological data showed that high-visibility face primes resulted in LRP differences between congruent and incongruent trials, suggesting that prime stimuli initiated motor preparation. Contrary to the above pattern, no evidence for reaction time or LRP differences was observed in the low-visibility condition, revealing that the depth of facial expression processing is dependent on stimulus visibility.     

Gaze Patterns When Looking at Emotional Pictures: Motivationally Biased Attention

Pictures of either emotionally neutral, pleasant (affectionate), or unpleasant (threat and injury) scenes were paired with nonemotional control pictures and presented for 3 s, while eye fixations were monitored. Preferential attention to the emotional picture of eah pair occurred at an early processing stage: The probability of placement of the first fixation and the proportion of viewing time during the first 500 ms were higher for both pleasant and unpleasant pictures than for neutral pictures, whereas later fixation location and time were not differentially affected. This suggests that, when emotional and nonemotional stimuli are presented simultaneously, emotional meaning captures initial overt orienting and engages attention early, as measured by gaze direction and duration. The possibility that this effect could be accounted for by perceptual differences—such as luminance and complexity—between emotional and neutral pictures was ruled out.     

Mindful attention predicts greater recovery from negative emotions,but not reduced reactivity

This study investigated the role of dispositional mindful attention in immediate reactivity to, and subsequent recovery from, laboratory-induced negative emotion. One hundred and fourteen undergraduates viewed blocks of negative pictures followed by neutral pictures. Participants’ emotional responses to negative pictures and subsequent neutral pictures were assessed via self-reported ratings. Participants’ emotional response to negative pictures was used to index level of emotional reactivity to unpleasant stimuli; emotional response to neutral pictures presented immediately after the negative pictures was used to index level of emotional recovery from pre-induced negative emotion (residual negativity). Results indicated that mindful attention was not associated with the emotional response to negative pictures, but it was associated with reduced negative emotion in response to the neutral pictures presented immediately after the negative pictures, suggesting better recovery as opposed to reduced reactivity. This effect was especially pronounced in later experimental blocks when the accumulation of negative stimuli produced greater negative emotion from which participants had to recover. The current study extends previous findings on the relationship between dispositional mindfulness and reduced negative emotion by demonstrating that mindful attention may facilitate better recovery from negative emotion, possibly through more effective disengagement from previous stimuli.     

Emotion affects action: Midcingulate cortex as a pivotal node of interaction between negative emotion and motor signals

Affective pictures drive the activity of brain networks and impact behavior. We showed previously that viewing unpleasant pictures interfered in the performance of a basic nonemotional visual detection task. In the present study, we employed functional magnetic resonance imaging to test the hypothesis that behavioral interference may result from the interaction between negatively valenced and motor-related signals in the brain. As in our previous study (Pereira et al., 2006), participants performed a simple target detection task that followed the presentation of unpleasant or neutral pictures. Our results revealed that an unpleasant emotional context modulated evoked responses in several regions engaged by the simple target detection task. In particular, the midcingulate cortex was recruited when participants performed target detection trials during the unpleasant context, and signal responses in this region closely mirrored the pattern of behavioral interference (as revealed via reaction time). Our findings suggest that the midcingulate cortex may be an important site for the interaction between negatively valenced signals and motor signals in the brain and that it may be involved in the implementation of defensive responses, such as freezing.     

Emotional state affects the initiation of forward gait

The aim of the current study was to determine the extent to which pleasant and unpleasant emotional states impact the initiation of forward gait. Participants initiated gait and walked for several steps following the presentation of low arousing pleasant, high arousing pleasant, low arousing unpleasant, high arousing unpleasant, and neutral pictures. Reaction time, displacement, and velocity of the center of pressure (COP) trajectory, and length and velocity of the first and second steps were calculated. Exposure to the highly arousing unpleasant pictures reduced reaction times compared to all other affective conditions. Compared to the low arousing unpleasant pictures, exposure to the high and low arousing pleasant pictures increased the displacement of the COP movement during the anticipatory postural adjustment phase of gait initiation. Additionally, exposure to the low arousing pleasant pictures increased the velocity of the COP movement during the anticipatory postural adjustment phase, compared to the high and low arousing unpleasant pictures. Exposure to the high and low arousing pleasant pictures increased the velocity of the first step relative to the low arousing unpleasant pictures. These findings demonstrate that highly arousing unpleasant emotional states accelerate the initial motor response, but pleasant emotional states generally facilitate the initiation of forward gait due to the approach-oriented directional salience of the movement. These findings extend the scope of the motivational direction hypothesis by demonstrating the effects of emotional reactivity on the initiation of gait.     

The persistence of attention to emotion: brain potentials during and after picture presentation

Emotional stimuli have been shown to elicit increased perceptual processing and attentional allocation. The late positive potential (LPP) is a sustained P300-like component of the event-related potential that is enhanced after the presentation of pleasant and unpleasant pictures as compared with neutral pictures. In this study, the LPP was measured using dense array electroencephalograph both before and after pleasant, neutral, and unpleasant images to examine the time course of attentional allocation toward emotional stimuli. Results from 17 participants confirmed that the LPP was larger after emotional than neutral images and that this effect persisted for 800 ms after pleasant picture offset and at least 1,000 ms after unpleasant picture offset. The persistence of increased attention after unpleasant compared to pleasant stimuli is consistent with the existence of a negativity bias. Overall, these results indicate that attentional capture of emotion continues well beyond picture presentation and that this can be measured with the LPP. Implications and directions for future research are discussed.     

Evaluative ratings and attention across the life span: emotional arousal and gender

This study was designed to investigate the evolution of emotional processing over the whole adult life span as a function of stimulus arousal and participants’ gender. To this end, self-reported affective evaluation and attentional capture prompted by pleasant and unpleasant pictures varying in arousal were measured in a large sample of participants (n?=?211) balanced by gender and equally spread across seven decades from 20 to 90 years. Results showed age differences only for affective evaluation of pleasant stimuli, with opposite patterns depending on stimulus arousal. As age increased, low-arousing pleasant cues (e.g. images of babies) were experienced as more pleasant and arousing by both males and females, whereas high-arousing stimuli (e.g. erotic images) were experienced as less pleasant only by females. In contrast, emotional pictures (both pleasant and unpleasant) were effective at capturing attention in a similar way across participants, regardless of age and gender. Taken together, these findings suggest that specific emotional cues prompt different subjective responses across different age groups, while basic mechanisms involved in attentional engagement towards both pleasant and unpleasant stimuli are preserved in healthy ageing.     

Intoxication level and emotional response

We used affective modulation of the eye-blink component of the startle reflex to examine effects of three levels of alcohol intoxication and a no-intoxication control on emotional responses to pleasant, neutral, and unpleasant pictures. Non-problematic student drinkers (n=101; 48 female) were randomly assigned to intoxication groups. Normal inhibition of startle during exposure to pleasant pictures was intact across groups. In contrast, potentiation of startle during viewing of unpleasant pictures was evident in the no- and low-intoxication groups, compared to the intermediate- and high-intoxication groups, in which it was significantly reduced. This pattern suggests that a direct and selective anxiolytic effect of alcohol can occur at higher levels of intoxication without an analogous impact on response to emotionally positive stimuli at similar levels.     

Reappraisal modulates the electrocortical response to unpleasant pictures

Cognitive strategies such as reappraisal reduce the intensity of negative experience and brain activity that is sensitive to emotional salience. The time course of reappraisal-related neural modulation remains unclear, and it is unknown whether the electrocortical response to emotional stimuli is sensitive to reappraisal. Event-related brain potentials were recorded first while participants passively viewed pleasant, unpleasant, and neutral pictures, and then during an emotion regulation block in which participants were instructed to attend to or reappraise unpleasant pictures. The late positive potential (LPP) was enhanced for pleasant and unpleasant pictures in the passive viewing block, and reappraisal resulted in a reliably reduced LPP—a protracted modulation that began 200 msec after stimulus onset. The degree of LPP modulation was positively related to reductions in the self-reported emotional intensity that followed emotion regulation instructions. These results indicate that reappraisal modulates early electrocortical activity that is related to emotional salience, and that the LPP is a useful tool for studying emotion regulation.     

Responses of the human motor system to observing actions across species: A transcranial magnetic stimulation study

Ample evidence suggests that the role of the mirror neuron system (MNS) in monkeys is to represent the meaning of actions. The MNS becomes active in monkeys during execution, observation, and auditory experience of meaningful, object-oriented actions, suggesting that these cells represent the same action based on a variety of cues. The present study sought to determine whether the human motor system, part of the putative human MNS, similarly represents and reflects the meaning of actions rather than simply the mechanics of the actions. To this end, transcranial magnetic stimulation (TMS) of primary motor cortex was used to generate motor-evoked potentials (MEPs) from muscles involved in grasping while participants viewed object-oriented grasping actions performed by either a human, an elephant, a rat, or a body-less robotic arm. The analysis of MEP amplitudes suggested that activity in primary motor cortex during action observation was greatest during observation of the grasping actions of the rat and elephant, and smallest for the human and robotic arm. Based on these data, we conclude that the human action observation system can represent actions executed by non-human animals and shows sensitivity to species-specific differences in action mechanics.