How the brain laughs. Comparative evidence from behavioral, electrophysiological and neuroimaging studies in human and monkey.

Laughter is an affective nonspeech vocalization that is not reserved to humans, but can also be observed in other mammalians, in particular monkeys and great apes. This observation makes laughter an interesting subject for brain research as it allows us to learn more about parallels and differences of human and animal communication by studying the neural underpinnings of expressive and perceptive laughter. In the first part of this review we will briefly sketch the acoustic structure of a bout of laughter and relate this to the differential anatomy of the larynx and the vocal tract in human and monkey. The subsequent part of the article introduces the present knowledge on behavioral and brain mechanisms of "laughter-like responses" and other affective vocalizations in monkeys and apes, before we describe the scant evidence on the cerebral organization of laughter provided by neuroimaging studies. Our review indicates that a densely intertwined network of auditory and (pre-) motor functions subserves perceptive and expressive aspects of human laughter. Even though there is a tendency in the present literature to suggest a rightward asymmetry of the cortical representation of laughter, there is no doubt that left cortical areas are also involved. In addition, subcortical areas, namely the amygdala, have also been identified as part of this network. Furthermore, we can conclude from our overview that research on the brain mechanisms of affective vocalizations in monkeys and great apes report the recruitment of similar cortical and subcortical areas similar to those attributed to laughter in humans. Therefore, we propose the existence of equivalent brain representations of emotional tone in human and great apes. This reasoning receives support from neuroethological models that describe laughter as a primal behavioral tool used by individuals - be they human or ape - to prompt other individuals of a peer group and to create a mirthful context for social interaction and communication.