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Accepted manuscripts are peer-reviewed but have not been through the copyediting, formatting, or proofreading process. Copyright © 2017 the authors This Accepted Manuscript has not been copyedited and formatted. The final version may differ from this version. Research Articles: Behavioral/Cognitive Social Laughter Triggers Endogenous Opioid Release in Humans Sandra Manninen 1 , Lauri Tuominen 1 , Robin Dunbar 2,3 , Tomi Karjalainen 1 , Jussi Hirvonen 1 , Eveliina Arponen 1 , Riitta Hari 2,4 , Iiro P. Jääskeläinen 2 , Mikko Sams 2 and Lauri Nummenmaa 1,5 1 Turku PET Centre, 20520 Turku, Finland 2 Department of Neuroscience and Biomedical Engineering, School of Science, Aalto University, 00076 AALTO, Finland 3 Department of Experimental Psychology, University of Oxford, OX1 3UD, Oxford, UK 4 Department of Art, School of Art, Design and Architecture, Aalto University, 00076 AALTO, Finland 5 Department of Psychology, University of Turku, 20014, Turku, Finland DOI: 10.1523/JNEUROSCI.0688-16.2017 Received: 2 March 2016 Revised: 13 March 2017 Accepted: 10 April 2017 Published: 23 May 2017 Author contributions: L.N., L.T., R.I.D., J.H., R.H., I.P.J., and M.S. designed research; L.N., S.M., L.T., and T.K. analyzed data; L.N., S.M., L.T., R.I.D., T.K., J.H., R.H., I.P.J., and M.S. wrote the paper; S.M., L.T., R.I.D., and E.A. performed research. Conflict of Interest: The authors declare no conflict of interest. This research was supported by the Academy of Finland (grants #265915 and #294897 to LN, #276643 to IPJ and #218072 to RH), ERC Starting Grant #313000 to LN and ERC Advanced Grants #232946 to RH and #295663 to RD. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Correspondence to: Lauri Nummenmaa, Turku PET Centre, Kiinamyllynkatu 4-6, FI-20520 Turku, Finland, email: [email protected] Cite as: J. Neurosci ; 10.1523/JNEUROSCI.0688-16.2017 Alerts: Sign up at www.jneurosci.org/cgi/alerts to receive customized email alerts when the fully formatted version of this article is published.

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Page 1: Social Laughter Triggers Endogenous Opioid Release in Humans · ZÀ]}µ ]v }oÇPu}µ } v ~Wvl oXUíõôì v DKZ rP v rlv} l}µ u] µ ] oÇ (] ] ]v Zu v ZÀ]}µ ~D}o oXUîì ìð XZZ

Accepted manuscripts are peer-reviewed but have not been through the copyediting, formatting, or proofreadingprocess.

Copyright © 2017 the authors

This Accepted Manuscript has not been copyedited and formatted. The final version may differ from this version.

Research Articles: Behavioral/Cognitive

Social Laughter Triggers Endogenous Opioid Release in Humans

Sandra Manninen1, Lauri Tuominen1, Robin Dunbar2,3, Tomi Karjalainen1, Jussi Hirvonen1, Eveliina Arponen1,

Riitta Hari2,4, Iiro P. Jääskeläinen2, Mikko Sams2 and Lauri Nummenmaa1,5

1Turku PET Centre, 20520 Turku, Finland2Department of Neuroscience and Biomedical Engineering, School of Science, Aalto University, 00076 AALTO,Finland3Department of Experimental Psychology, University of Oxford, OX1 3UD, Oxford, UK4Department of Art, School of Art, Design and Architecture, Aalto University, 00076 AALTO, Finland5Department of Psychology, University of Turku, 20014, Turku, Finland

DOI: 10.1523/JNEUROSCI.0688-16.2017

Received: 2 March 2016

Revised: 13 March 2017

Accepted: 10 April 2017

Published: 23 May 2017

Author contributions: L.N., L.T., R.I.D., J.H., R.H., I.P.J., and M.S. designed research; L.N., S.M., L.T., andT.K. analyzed data; L.N., S.M., L.T., R.I.D., T.K., J.H., R.H., I.P.J., and M.S. wrote the paper; S.M., L.T., R.I.D.,and E.A. performed research.

Conflict of Interest: The authors declare no conflict of interest.

This research was supported by the Academy of Finland (grants #265915 and #294897 to LN, #276643 toIPJ and #218072 to RH), ERC Starting Grant #313000 to LN and ERC Advanced Grants #232946 to RH and#295663 to RD. The funders had no role in study design, data collection and analysis, decision to publish, orpreparation of the manuscript.

Correspondence to: Lauri Nummenmaa, Turku PET Centre, Kiinamyllynkatu 4-6, FI-20520 Turku, Finland,email: [email protected]

Cite as: J. Neurosci ; 10.1523/JNEUROSCI.0688-16.2017

Alerts: Sign up at www.jneurosci.org/cgi/alerts to receive customized email alerts when the fully formattedversion of this article is published.

Page 2: Social Laughter Triggers Endogenous Opioid Release in Humans · ZÀ]}µ ]v }oÇPu}µ } v ~Wvl oXUíõôì v DKZ rP v rlv} l}µ u] µ ] oÇ (] ] ]v Zu v ZÀ]}µ ~D}o oXUîì ìð XZZ
Page 3: Social Laughter Triggers Endogenous Opioid Release in Humans · ZÀ]}µ ]v }oÇPu}µ } v ~Wvl oXUíõôì v DKZ rP v rlv} l}µ u] µ ] oÇ (] ] ]v Zu v ZÀ]}µ ~D}o oXUîì ìð XZZ
Page 4: Social Laughter Triggers Endogenous Opioid Release in Humans · ZÀ]}µ ]v }oÇPu}µ } v ~Wvl oXUíõôì v DKZ rP v rlv} l}µ u] µ ] oÇ (] ] ]v Zu v ZÀ]}µ ~D}o oXUîì ìð XZZ
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