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The cognitive processes involved in humor comprehension were analyzed by directly comparing the time course of brain activity associated with the perception of slapstick humor and that associated with the comprehension of humor requiring theory of mind (ToM). Four different comic strips (strips containing humorous scenes that required ToM, non-ToM humorous strips, non-humorous semantically coherent strips and non-humorous semantically incoherent strips) were presented to participants, while their EEG response was recorded. Results showed that both of the humorous comic strips and the semantically incongruent strip elicited an N400 effect, suggesting similar cognitive mechanisms underlying the processing of incongruent and humorous comic strips. The results also showed that the humorous ToM strips elicited a frontal late positive (LP) response, possibly reflecting the active deployment of ToM abilities such as perspective-taking and empathy that allow for the resolution and interpretation of apparently incongruent situations. In addition, the LP response was positively correlated with ratings of perceived amusement as well as individual empathy scores, suggesting that the increased LP response to ToM humorous strips reflects the combined activation of neural mechanisms involved in the experience of amusement and ToM abilities. Overall, humor comprehension appears to demand distinct cognitive steps such as the detection of incongruent semantic components, the construction of semantic coherence, and the appreciation of humoristic elements such as maladaptive emotional reactions. Our results show that the deployment of these distinct cognitive steps is at least partially dependent on individual empathic abilities.

The superior temporal gyrus (STG) has been found to play a crucial role in the recognition of actions and facial expressions and may, therefore, be critical for the processing of humorous information. Here we investigated whether tDCS application to the STG would modulate the ability to recognize and appreciate the comic element in serious and comedic situations of misfortune. To this aim, the effects of different types of tDCS stimulation on the STG were analyzed during a task in which the participants were instructed to categorize various misfortunate situations as "comic" or "not comic". Participants underwent three different tDCS conditions: Anodal-right/Cathodal-left; Cathodal-right/Anodal-left; Sham. Images depicting people involved in accidents were grouped into three categories based on the facial expression of the victim: angry or painful (Affective); bewildered and funny (Comic); and images that did not contain the victim's face (No Face). An improvement in mean reaction times in response to both the Comic and No Face stimuli was observed following Anodal-left/Cathodal-right stimulation when compared to sham stimulation. This suggests that this stimulation type reduced the reaction times to socio-emotional complex scenes, regardless of facial expression. The Anodal-right/Cathodal-left stimulation reduced the mean reaction times for Comic stimuli only, suggesting that specifically the right STG may be involved in facial expression recognition and in the appreciation of the comic element in misfortunate situations. These results suggest a functional hemispheric asymmetry in STG response to social stimuli: the left STG might have a role in a general comprehension of social complex situations, while the right STG may be involved in the ability to recognize and integrate specific emotional aspects in a complex scene.