Daniele Marzoli

Effects of Transcranial Direct Current Stimulation on episodic memory related to emotional visual stimuli.

The present study investigated emotional memory following bilateral transcranial electrical stimulation (direct current of 1 mA, for 20 minutes) over fronto-temporal cortical areas of healthy participants during the encoding of images that differed in affective arousal and valence. The main result was a significant interaction between the side of anodal stimulation and image emotional valence. Specifically, right anodal/left cathodal stimulation selectively facilitated the recall of pleasant images with respect to both unpleasant and neutral images whereas left anodal/right cathodal stimulation selectively facilitated the recall of unpleasant images with respect to both pleasant and neutral images. From a theoretical perspective, this double dissociation between the side of anodal stimulation and the advantage in the memory performance for a specific type of stimulus depending on its pleasantness supported the specific-valence hypothesis of emotional processes, which assumes a specialization of the right hemisphere in processing unpleasant stimuli and a specialization of the left hemisphere in processing pleasant stimuli. From a methodological point of view, first we found tDCS effects strictly dependent on the stimulus category, and second a pattern of results in line with an interfering and inhibitory account of anodal stimulation on memory performance. These findings need to be carefully considered in applied contexts, such as the rehabilitation of altered emotional processing or eye-witness memory, and deserve to be further investigated in order to understand their underlying mechanisms of action.

Perceptual asymmetries and handedness: a neglected link?

Healthy individuals tend to weigh in more the left than the right side of visual space in a variety of contexts, ranging from pseudoneglect to perceptual asymmetries for faces. Among the common explanations proposed for the attentional and perceptual advantages of the left visual field, a link with the prevalence of right-handedness in humans has never been suggested, although some evidence seems to converge in favor of a bias of spatial attention toward the region most likely coincident with another person’s right hand during a face-to-face interaction. Such a bias might imply an increased efficiency in monitoring both communicative and aggressive acts, the right limb being more used than the left in both types of behavior. Although attentional and perceptual asymmetries could be linked to right-handedness at the level of phylogeny because of the evolutionarily advantage of directing attention toward the region where others’ dominant hand usually operates, it is also legitimate to question whether, at the ontogenetic level, frequent exposure to right-handed individuals may foster leftward biases. These views are discussed in the light of extant literature, and a number of tests are proposed in order to assess our hypotheses.

The Relation between Self-Reported Empathy and Motor Identification with Imagined Agents

Background In a previous study, we found that when required to imagine another person performing an action, participants reported a higher correspondence between their own handedness and the hand used by the imagined person when the agent was seen from the back compared to when the agent was seen from the front. This result was explained as evidence of a greater involvement of motor areas in the back-view perspective, possibly indicating a greater proneness to put oneself in the agents shoes in such a condition. In turn, the proneness to put oneself in anothers shoes could also be considered as a cue of greater identification with the other, that is a form of empathy. If this is the case, the proportion of lateral matches vs mismatches should be different for subjects with high and low self-reported empathy. In the present study, we aimed at testing this hypothesis. Methodology/Principal Findings Participants were required to imagine a person performing a single manual action in a back view and to indicate the hand used by the imagined person during movement execution. Consistent with our hypothesis, the proportion of matching between the handedness of participants and the handedness of agents imagined was higher for participants scoring high in a self-report measure of empathy. Importantly, this relationship was specific for females. Conclusions/Significance At least for females, our data seem to corroborate the idea of a link between self-reported empathy and motor identification with imagined agents. This sex-specific result is consistent with neuroimaging studies indicating a stronger involvement of action representations during emotional and empathic processing in females than in males. In sum, our findings underline the possibility of employing behavioral research as a test-bed for theories deriving from functional studies suggesting a link between empathic processing and the activation of motor-related areas.

Imagining others’ handedness: visual and motor processes in the attribution of the dominant hand to an imagined agent

In a previous study, we found that when required to imagine another person performing an action, participants reported a higher correspondence between their own dominant hand and the hand used by the imagined person when the agent was visualized from the back compared to when the agent was visualized from the front. This suggests a greater involvement of motor representations in the back-view perspective, possibly indicating a greater proneness to put oneself in the agent’s shoes in such a condition. In order to assess whether bringing to the foreground the right or left hand of an imagined agent can foster the activation of the corresponding motor representations, we required 384 participants to imagine a person—as seen from the right or left side—performing a single manual action and to indicate the hand used by the imagined person during movement execution. The proportion of right- versus left-handed reported actions was higher in the right-view condition than in the left-view condition, suggesting that a lateral vantage point may activate the corresponding hand motor representations, which is in line with previous research indicating a link between the hemispheric specialization of one’s own body and the visual representation of others’ bodies. Moreover, in agreement with research on hand laterality judgments, the effect of vantage point was stronger for left-handers (who reported a higher proportion of right- than left-handed actions in the right-view condition and a slightly higher proportion of left- than right-handed actions in the left-view condition) than for right-handers (who reported a higher proportion of right- than left-handed actions in both view conditions), indicating that during the mental simulation of others’ actions, right-handers rely on sensorimotor processes more than left-handers, while left-handers rely on visual processes more than right-handers.

Side biases in humans (Homo sapiens): three ecological studies on hemispheric asymmetries.

Hemispheric asymmetries and side biases have been studied in humans mostly in laboratory settings, and evidence obtained in naturalistic settings is scarce. We here report the results of three studies on human ear preference observed during social interactions in noisy environments, i.e., discotheques. In the first study, a spontaneous right-ear preference was observed during linguistic exchange between interacting individuals. This lateral bias was confirmed in a quasi-experimental study in which a confederate experimenter evoked an ear-orienting response in bystanders, under the pretext of approaching them with a whispered request. In the last study, subjects showed a greater proneness to meet an experimenter’s request when it was directly addressed to the right rather than the left ear. Our findings are in agreement both with laboratory studies on hemispheric lateralization for language and approach/avoidance behavior in humans and with animal research. The present work is one of the few studies demonstrating the natural expression of hemispheric asymmetries, showing their effect in everyday human behavior.

GABA content within the ventromedial prefrontal cortex is related to trait anxiety

The ventromedial prefrontal cortex (vmPFC) plays a key role in emotion processing and regulation. vmPFC dysfunction may lead to disinhibition of amygdala causing high anxiety levels. γ-Aminobutyric acid (GABA) inter-neurons within vmPFC shape the information flow to amygdala. Thus, we hypothesize that GABA content within vmPFC could be relevant to trait anxiety. Forty-three healthy volunteers aged between 20 and 88 years were assessed for trait anxiety with the Subscale-2 of the State-Trait-Anxiety Inventory (STAI-Y2) and were studied with proton magnetic resonance spectroscopy to investigate GABA and Glx (glutamate+glutamine) contents within vmPFC. Total creatine (tCr) was used as internal reference. Partial correlations assessed the association between metabolite levels and STAI-Y2 scores, removing the effect of possible nuisance factors including age, educational level, volumes of gray matter and white matter within magnetic resonance spectroscopy voxel. We observed a positive relationship between GABA/tCr and STAI-Y2 scores. No significant relationships were found between Glx/tCr and STAI-Y2 and between tCr/water and STAI-Y2. No differences were found between males and females as regards to age, STAI-Y2, GABA/tCr, Glx/tCr, tCr/water, gray matter and white matter volumes. We suggest a close relationship between GABA content within vmPFC and trait anxiety providing new insights in the physiology of emotional brain.

Sun-induced frowning fosters aggressive feelings

We tested whether aggressiveness can be triggered by the involuntary frowning that occurs when people face the sun, due to the fact that sun-induced frowning involves the same pattern of facial muscle activation as in the expression of anger (interestingly, Charles Darwin remarked on the sunshade-like nature of frowning). In line with data showing that experimentally and unobtrusively induced facial and body displays facilitate congruent feelings, we found that participants walking against the sun without sunglasses scored higher in a self-report measure of anger and aggression compared to those walking with the sun behind and/or wearing sunglasses. We also suggest that frowning at the sun affects mood very quickly, because we did not find any effect of walking time on self-reported aggressiveness. Our results provide the first evidence of the ecological validity of the facial feedback hypothesis.

The processing of chimeric and dichotic emotional stimuli by connected and disconnected cerebral hemispheres

Hemispheric asymmetries have been widely explored in both the visual and the auditory domain, but little is known about hemispheric asymmetries in audio-visual integration. We compared the performance of a partially callosotomized patient, a total split-brain patient and a control group during the evaluation of the emotional valence of chimeric faces and dichotic syllables (an emotional syllable in one ear and white noise in the other ear) presented unimodally (only faces or only syllables) or bimodally (faces and syllables presented simultaneously). Stimuli could convey happy and sad expressions and participants were asked to evaluate the emotional content of each presentation, using a 5-point Likert scale (from very sad to very happy). In unimodal presentations, the partially callosotomized patients judgments depended on the emotional valence of the stimuli processed by the right hemisphere, whereas those of the total split-brain patient showed the opposite lateralization; in these conditions, the control group did not show asymmetries. Moreover, in bimodal presentations, results provided support for the valence hypothesis (i.e., left asymmetry for positive emotions and vice versa) in both the control group and the partially callosotomized patient, whereas the total split-brain patient showed a tendency to evaluate the emotional content of the right hemiface even when asked to focus on the acoustic modality. We conclude that partial and total hemispheric disconnections reveal opposite patterns of hemispheric asymmetry in auditory, visual and audio-visual emotion processing. These results are discussed in the light of the right-hemisphere hypothesis and the valence hypothesis.

Hearing it right: Evidence of hemispheric lateralization in auditory imagery

An advantage of the right ear (REA) in auditory processing (especially for verbal content) has been firmly established in decades of behavioral, electrophysiological and neuroimaging research. The laterality of auditory imagery, however, has received little attention, despite its potential relevance for the understanding of auditory hallucinations and related phenomena. In Experiments 1-4 we find that right-handed participants required to imagine hearing a voice or a sound unilaterally show a strong population bias to localize the self-generated auditory image at their right ear, likely the result of left-hemispheric dominance in auditory processing. In Experiments 5-8 - by means of the same paradigm - it was also ascertained that the right-ear bias for hearing imagined voices depends just on auditory attention mechanisms, as biases due to other factors (i.e., lateralized movements) were controlled. These results, suggesting a central role of the left hemisphere in auditory imagery, demonstrate that brain asymmetries can drive strong lateral biases in mental imagery.

GABA levels in the ventromedial prefrontal cortex during the viewing of appetitive and disgusting food images.

Characterizing how the brain appraises the psychological dimensions of reward is one of the central topics of neuroscience. It has become clear that dopamine neurons are implicated in the transmission of both rewarding information and aversive and alerting events through two different neuronal populations involved in encoding the motivational value and the motivational salience of stimuli, respectively. Nonetheless, there is less agreement on the role of the ventromedial prefrontal cortex (vmPFC) and the related neurotransmitter release during the processing of biologically relevant stimuli. To address this issue, we employed magnetic resonance spectroscopy (MRS), a non-invasive methodology that allows detection of some metabolites in the human brain in vivo, in order to assess the role of the vmPFC in encoding stimulus value rather than stimulus salience. Specifically, we measured gamma-aminobutyric acid (GABA) and, with control purposes, Glx levels in healthy subjects during the observation of appetitive and disgusting food images. We observed a decrease of GABA and no changes in Glx concentration in the vmPFC in both conditions. Furthermore, a comparatively smaller GABA reduction during the observation of appetitive food images than during the observation of disgusting food images was positively correlated with the scores obtained to the body image concerns sub-scale of Body Uneasiness Test (BUT). These results are consistent with the idea that the vmPFC plays a crucial role in processing both rewarding and aversive stimuli, possibly by encoding stimulus salience through glutamatergic and/or noradrenergic projections to deeper mesencephalic and limbic areas.