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Humor(Goel & Dolan, 2001; Mobbs et al., 2003; Moran et al., 2004; Watson et al., 2007)2

Wolfenstein,1955; Dixon,1980; Veatch,1998*Martin, R. A.2011207-234TSSCI*2010232271-304TSSCI*2010232241-269TSSCI

3Nicholson,1973Yorukoglu,1977Chang, H. J., Wang, C. Y., & Chen, H. C.* (2013). The analysis of elementary and high school students' natural and humorous responses patterns in coping with embarrassing situations. Humor International Journal of Humor Research. (SSCI)

4(emotional Shift) (Morreall,1987)(Kris,1952)Freud,1950(Mindess)5Martineau,1972Lorenz,1963200436113-33TSSCI*2010232271-304TSSCI6HauckThomas,1972Wakshlag,DayZillmann,19811996535-65201146212-212011.1256443-687Chan, Y. C., Chen, H. C*., & Lavallee, J. P. (in press). The impact of gelotophobia, gelotophilia and katagelasticism on creativity. Humor International Journal of Humor Research. (SSCI)200334179-198TSSCI200435393-411TSSCI20065171-93TSSCI8()(Nazareth,1988)

330 1901James Sully 79 200143(2)137-153TSSCI1995--2424026120052655-5620065171-93TSSCI2010232173-182TSSCI102 1112 SetupPunch lineincongruity theoryNerhardt : :14

Suls1972incongruity-resolution theory -Comprehension Elaboration Theory )Wyer Collins1992 )-

( 1991,1995)

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18(in press).TSSCIChan, Y. C.,Chou, T. L.,Chen, H. C., Yeh, Y. C., Lavallee, J. P., Liang, K. C., & Chang, K. E. (2013).Towards a neural circuit model of verbal humor processing: An fMRI study of the neural substrates of incongruity detection and resolution.NeuroImage.66C, 169-176. (SCI) Chan, Y. C., Chou, T. L., Chen, H. C., & Liang, K. C. (2012). Segregating humor comprehension and elaboration process of verbal jokes: An fMRI study. NeuroImage. 61(4), 899-906. (SCI)

Verbal JokeWhile a great deal of the humor that we experience on a daily basis is verbal in nature (Brownell et al., 1983), We used verbal jokes instead of films, meaning that our material was more semantic in nature.The cognitive processes underlying the resolution of verbal jokes seem to be a part of executive functions such as disambiguation, bridge inferences, and schema shifting to reestablishment of context (Bekinschtein et al., 2011).20Nonverbal jokesMost studies using functional magnetic resonance imaging (fMRI) technology to study humor processing have focused on non-verbal processing, using, for example, nonverbal visual cartoons (Azim et al., 2005; Bartolo et al., 2006; Moran et al., 2004; Samson et al., 2008, 2009; Wild et al., 2006) or humorous videos (Moran et al., 2004; Neely et al., 2012).This study joins earlier studies (Bekinschtein et al., 2011; Goel and Dolan, 2001; Watson et al., 2007) in focusing on verbal humor processing.21Non-verbal jokes (cartoon and film)22

Kohn et al., (2011)

Moran et al., (2004)Goel and Dolan, 2001 (UCL)Participants listened to riddles containing either phonological jokes or semantic jokes, as well as a set of nonhumorous control riddles.The semantic jokes induced greater activation in regions of both the left and right temporal lobes (e.g., MTG, ITG) that are involved in semantic processing of language.The Phonological jokes induced greater activation in areas of the left frontal lobe (e.g., IFG).Subjects post-scan funniness ratings of jokes were correlated with significantly greater activation of the vmPFC, an area with connections to the limbic system that plays an important role in integrating cognitive and emotion processes. 23Nature Neuroscience

23Goel and Dolan, 2001 (UCL)What do engineers use for birth control?Their personalities. (Funny)The pill. (Unfunny)Why did the golfer wear two sets of pants?He got a hole in one. (Funny)It was a very cold day. (Unfunny)

24Nature NeuroscienceGoel and Dolan, 2001 (UCL)25Nature Neuroscience

bilateral MTGLeft IFG

Right MTG, Left ITGleft inferior precentral gyrus/insula

Left vmPFCBekinschtein et al., 2011 (University of Cambridge)26The Journal of neuroscience

Jokes versus non-jokes effects while ambiguity effect was only seen in left ITG (LITG) and left anterior IFG (L-ant-IFG; shaded bars for ambiguity).Mobbs et al., 2003 (Stanford University)27Neuron

Mobbs et al., 2003 (Stanford University)28Neuron

Funny Minus Nonfunny Cartoonsextent threshold (p 0.05) corrected at the whole-brain levelMobbs et al., 2003 (Stanford University)29Neuron

Covarying with Degree of Humor Intensityextent threshold (p 0.01) corrected at the whole-brain levelSegregating the Comprehension and Elaboration Processing of Verbal Jokes: An fMRI Study

Study 1Chan, Y. C., Chou, T. L., Chen, H. C., & Liang, K. C. (2012). NeuroImage, 61(4), 899-906.Funny vs. Unfunnyvs.Cognitive componentFunny(incongruity-resolution)(amusement)Unfunny(no incongruities)(calmness)ElaborationEmotional component+Comprehension31comprehension and elaboration of humor32

Garden pathGarden path sentences refer to sentences that are grammatically correct, but which readers typically misinterpret in their first reading, leading to a semantic dead end (i.e., incongruity).Readers are then required to resolve this incongruity by rereading the sentence to construct a sustainable interpretation (i.e., resolution of incongruity) (Christensen, 2010, Uchiyama et al., 2008). 33Funny vs. Garden path vs. Unfunnyvs.vs.vs.ComprehensionElaborationUnfunny(no incongruities)(Calmness)Garden Path(incongruity-resolution)(Calmness)ElaborationComprehension34Funny(incongruity-resolution)(Amusement)ParticipantsTwenty neurologically healthy volunteers [10 males and 10 females aged 2029 years, 23.40 2.74 years (mean SD)] participated in this study.They were all right-handed as determined by the Edinburgh Handedness Inventory (Oldfield, 1971) and native Mandarin speakers.35StimuliSixty stories in Mandarin Chinese were selected, including twelve stories for each of three different conditions: the funny, unfunny, and garden path story conditions.Twenty-four filler stories were also included. Each joke structure contained two components: the setup and the punch line.3637

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Experimental paradigm

39Image analysisGiven our a priori hypothesis based on previous imaging studies, a region of interest (ROI) analysis was performed (Poldrack et al., 2008).According to the finding of previous studies of humor, laughter, and reward, we used 13 hypothesis-driven ROIs with a radius of 10 mm centered at peak voxels of brain regions.

40humor comprehension: the garden path versus unfunny conditions

41humor elaboration: the funny versus garden path conditions

42Post-scan rating: the high versus low comprehensibility conditionsThe contrast of the high versus low comprehensibility conditions produced greater activation in the left SFG (MNI coordinates: -9, 2, 73) and left anterior IFG (-45, 35, 10; Z = 2.57, p < .01 uncorrected) with the use of ROIs.

43Post-scan rating: the high versus low funniness conditionsThe contrast of the high versus low funniness conditions produced greater activation in the left vmPFC (MNI coordinates: -6, 53, -11), left amygdala (-24, -7, -23), right amygdala (24, -7, -29), left parahippocampal gyrus (-21, -13, -26), and right parahippocampal gyrus (21, -13, -23) with the use of ROIs.44Humor comprehension (IFG, SFG)The association observed here between humor comprehension and activity in the frontal regions of the bilateral IFG and the left SFG verifies that the main functions of incongruity detection, semantic selection, and semantic integration are located in this area.45Humor elaboration(cortical and subcortical regions)The vmPFC, bilateral amygdalae and bilateral parahippocampal gyri are related to the reward associated with the feeling of amusement.

46Left and right hemisperesThe present study also reveals that the activated areas related to humor processing are located in both hemispheres, and in both cortical and subcortical regions.This finding is consistent with earlier research (i.e., Bartolo et al., 2006; Bekinschtein et al., 2011; Mobbs et al., 2003).47

Dissociating the Incongruity Detection and Resolution of Verbal Jokes during Humor comprehension: An fMRI Study

Chan, Y. C., Chou, T. L., Chen, H. C., Yeh, Y. C., Lavallee, J. P., Liang, K. C., & Chang, K. E. (2013). Towards a neural circuit model of verbal humor processing: An fMRI study of the neural substrates of incongruity detection and resolution. NeuroImage. (SCI)Study 2Humor comprehension49

Unfunny, Nonsensical, Funny

51vs.()(-----)()()(-----)(-----)vs.vs.StimuliSixty-four stories in Mandarin Chinese were selected, including sixteen stories for each of three different conditions: funny, unfunny, and nonsensical.Sixteen filler stories were also included.Incongruity detection:The nonsensical versus unfunny conditions

Incongruity resolution:The funny versus nonsensical conditionsSFGIPL

Incongruity detection: further analysis of surprisingness ratingsBased on the rating scores, the contrast of the high versus low surprisingness conditions for the ROIs produced greater activation in the right MTG (58, -1, -8; BA 21, 69 voxels; Z = 3.13, p < .05 FWE corrected) and right MFG (12, -10, 66; BA 6, 38 voxels; Z = 3.21, p < .05 FWE corrected). The results are consistent with those shown in Table 1 for incongruity detection, providing further support for our account of the neural substrates associated with the detection of incongruities in the humor comprehension process.Incongruity resolution: further analysis of comprehensibility ratingsBased on the rating scores, the contrast of the high versus low comprehensibility conditions for the ROIs produced greater activation in the left SFG (MNI coordinates: -9, 47, 45; BA 8, 96 voxels; Z = 5.42, p < .05 FWE corrected) and left IPL (-60, -49, 42; BA 40, 56 voxels; Z = 4.97, p < .05 FWE corrected). The results are consistent with those shown in Table 1 for incongruity resolution, providing further support for our account of the neural substrates associated with incongruity resolution in the humor comprehension process.Conjunction analysisIn order to examine regions involved in the incongruity detection process, we performed a conjunction analysis (using MarsBaR and WFU PickAtlas in SPM8) of regions activated in both the nonsensical and funny conditions. The conjunction analysis, masked by the main effect of incongruity detection (p < .05 uncorrected, 10 voxels), was performed to highlight regions of activations associated with nonsensical and funny conditions [conjunction (nonsensical-unfunny), (funny-unfunny)]. This analysis revealed a significant activation in the right middle temporal gyrus (MNI coordinates: 57, 2, -11; BA 21, 56 voxels, p < .05 FDR corrected).MTG, MFG, SFG, and IPLThe present findings thus suggest that the neural circuit for humor comprehension may include distinct components for the detection of incongruity and the resolution of incongruity, with initial incongruity detection processed by the MTG and MFG in the right hemisphere, followed by semantic integration and resolution processed in the SFG and IPL in the left hemisphere.These results are consistent with Jung-Beemans recent discussion of bilateral processes for comprehending natural language (Jung-Beeman, 2005).

Towards a Three-stage Neural Circuit Model of Verbal Humor Processing

General discussionNeural circuit of verbal humor processing61

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Conclusion64