1 2 The role of the medial temporal limbic system in processing emotions 3 in voice and music 4 Sascha Fru ¨ hholz Q1 a,b,1, *, Wiebke Trost a,b,1 , Didier Grandjean a,b 5 a Neuroscience of Emotion and Affective Dynamics Lab, Department of Psychology, University of Geneva, Geneva, Switzerland 6 b Swiss Center for Affective Sciences, University of Geneva, Geneva, Switzerland 7 8 9 10 Contents 1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 000 2. Which emotions are expressed and perceived in voices and music and how? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 000 3. The auditory system and the temporal limbic system (TLS) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 000 4. Vocal and musical emotions activate the TLS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 000 4.1. TLS activity in response to positive and negative vocal emotions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 000 4.1.1. Amygdala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 000 4.1.2. Hippocampus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 000 4.2. TLS activity in response to musical emotions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 000 5. Affective arousal, emotional intensity and multimodal processing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 000 5.1. TLS sensitivity to arousing and intense vocal emotions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 000 5.2. TLS sensitivity to arousing musical emotions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 000 Progress in Neurobiology xxx (2014) xxx–xxx A R T I C L E I N F O Article history: Received 5 June 2014 Received in revised form 16 September 2014 Accepted 29 September 2014 Available online xxx Keywords: Amygdala Hippocampus Emotion Music Voice Q2 Neuroimaging Neural network A B S T R A C T Subcortical brain structures of the limbic system, such as the amygdala, are thought to decode the emotional value of sensory information. Recent neuroimaging studies, as well as lesion studies in patients, have shown that the amygdala is sensitive to emotions in voice and music. Similarly, the hippocampus, another part of the temporal limbic system (TLS), is responsive to vocal and musical emotions, but its specific roles in emotional processing from music and especially from voices have been largely neglected. Here we review recent research on vocal and musical emotions, and outline commonalities and differences in the neural processing of emotions in the TLS in terms of emotional valence, emotional intensity and arousal, as well as in terms of acoustic and structural features of voices and music. We summarize the findings in a neural framework including several subcortical and cortical functional pathways between the auditory system and the TLS. This framework proposes that some vocal expressions might already receive a fast emotional evaluation via a subcortical pathway to the amygdala, whereas cortical pathways to the TLS are thought to be equally used for vocal and musical emotions. While the amygdala might be specifically involved in a coarse decoding of the emotional value of voices and music, the hippocampus might process more complex vocal and musical emotions, and might have an important role especially for the decoding of musical emotions by providing memory-based and contextual associations. ß 2014 Published by Elsevier Ltd. Abbreviations: ABR, auditory brainstem repsonse; AC/ac, auditory cortex; acb, accessory basal nucleus of the amygdala; AMY/amy, amygdala; b, basal nucleus of the amygdala; bg, basal ganglia; BOLD, blood oxygenation level dependent; c, central nucleus of the amygdala; ca, cornu armonis; CA1, cornu ammonis, subregion 1; CA3, cornu ammonis, subregion 3; cbll, cerebellum; CN, cochlear nucleus; CN/cn, cochlear nucleus; DG/dg, dentate gyrus; ERC, entorhinal cortex; erc, entorhinal cortex; F0, fundamental frequency; fMRI, functional magnetic resonance imaging; h, healthy controls; HC/hc, hippocampus; HG, Heschls gyrus; IC/ic, inferior colliculus; l, lateral nucleus of the amygdala; m, medial nucleus of the amygdala; MGB/mgb, medial geniculate body; MNI, Montreal Neurological institute; NAc, nucleus accumbens; p, patients; pc, patient controls; PET, positron emission tomography; PHC/phc, parahippocampal cortex; PPo, planum polare; PRC/prc, perirhinal cortex; PTe, planum temporale; SOC/soc, superior olivary complex; STC/stc, superior temporal cortex; STG, superior temporal gyrus; STS, superior temporal sulcus; sub, subiculum; TLS, temporal limbic system. * Corresponding author at: Swiss Center for Affective Sciences, University of Geneva, 9 Chemin des Mines, PO Box 60, 1211 Geneva 20, Switzerland. Tel.: +41 22 379 09 15. E-mail address: sascha.fruehholz@unige.ch (S. Fru ¨ hholz). 1 These authors are contributed equally to this review paper. G Model PRONEU 1350 1–17 Please cite this article in press as: Fru ¨ hholz, S., et al., The role of the medial temporal limbic system in processing emotions in voice and music. Prog. Neurobiol. (2014), http://dx.doi.org/10.1016/j.pneurobio.2014.09.003 Contents lists available at ScienceDirect Progress in Neurobiology jo u rn al ho m epag e: ww w.els evier .c om /lo cat e/pn eu ro b io http://dx.doi.org/10.1016/j.pneurobio.2014.09.003 0301-0082/ß 2014 Published by Elsevier Ltd.