Right hemisphere has the last laugh: neural dynamics of joke appreciation Ksenija Marinkovic & Sharelle Baldwin & Maureen G. Courtney & Thomas Witzel & Anders M. Dale & Eric Halgren Published online: 17 December 2010 # The Author(s) 2010. This article is published with open access at Springerlink.com Abstract Understanding a joke relies on semantic, mnemon- ic, inferential, and emotional contributions from multiple brain areas. Anatomically constrained magnetoencephalogra- phy (aMEG) combining high-density whole-head MEG with anatomical magnetic resonance imaging allowed us to estimate where the humor-specific brain activations occur and to understand their temporal sequence. Punch lines provided either funny, not funny (semantically congruent), or nonsensical (incongruent) replies to joke questions. Healthy subjects rated them as being funny or not funny. As expected, incongruous endings evoke the largest N400m in left-dominant temporo-prefrontal areas, due to integration difficulty. In contrast, funny punch lines evoke the smallest N400m during this initial lexical–semantic stage, consistent with their primed “surface congruity” with the setup question. In line with its sensitivity to ambiguity, the anteromedial prefrontal cortex may contribute to the subse- quent “second take” processing, which, for jokes, presum- ably reflects detection of a clever “twist” contained in the funny punch lines. Joke-selective activity simultaneously emerges in the right prefrontal cortex, which may lead an extended bilateral temporo-frontal network in establishing the distant unexpected creative coherence between the punch line and the setup. This progression from an initially promising but misleading integration from left frontotempo- ral associations, to medial prefrontal ambiguity evaluation and right prefrontal reprocessing, may reflect the essential tension and resolution underlying humor. Keywords Humor . MEG . Language . N400 . P600 . Prefrontal cortex Jokes often take the form of a setup question, followed after a short delay by a punch line: for instance, “What do you get when you cross a parrot with a centipede?” A brief suspenseful pause allows the audience to imagine the issue of this unlikely match. The punch line “a walkie-talkie” is surprising but also fits in a clever and unexpected way. Prominent models commonly postulate that the punch line is appreciated as funny in two stages of integration: In the first stage, the punch line is perceived as incongruous with the setup, and in the second, further consideration establishes a deeper coherence with the preceding context (Brownell, Michel, Powelson, & Gardner, 1983; Ramachandran, 1998; Suls, 1972; Veatch, 1998; Wyer & Collins, 1992). Thus, amusement may result when a “twist” in the meaning is successfully incorporated into the preceding context—that is, when the initial incongruity is resolved by reinterpreting or “frame shifting” (Coulson, 2001). Despite its important role in modulating social dynamics (Black, 1984; Kane, Suls, & Tedeschi, 1977), there have K. Marinkovic (*) : A. M. Dale : E. Halgren Department of Radiology, University of California San Diego, 9500 Gilman Dr., 0841, La Jolla, CA 92093-0841, USA e-mail: xenia@ucsd.edu S. Baldwin Aspen Institute for Behavioral Assessment, Syracuse, UT, USA M. G. Courtney Center for Memory and Brain, Boston University, Boston, MA, USA T. Witzel Harvard–MIT Health Sciences and Technology, Boston, MA, USA A. M. Dale : E. Halgren Department of Neurosciences, University of California San Diego, La Jolla, CA, USA Cogn Affect Behav Neurosci (2011) 11:113–130 DOI 10.3758/s13415-010-0017-7