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Right hemisphere has the last laugh: neural dynamics of joke appreciation

By Ksenija Marinkovic, Sharelle Baldwin, Maureen G. Courtney, Thomas Witzel, Anders M. Dale and Eric Halgren


Understanding a joke relies on semantic, mnemonic, inferential, and emotional contributions from multiple brain areas. Anatomically constrained magnetoencephalography (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 subsequent “second take” processing, which, for jokes, presumably 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 frontotemporal associations, to medial prefrontal ambiguity evaluation and right prefrontal reprocessing, may reflect the essential tension and resolution underlying humor

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  1. (2011). (in press). Sequential temporo-fronto-temporal activation during monitoring of the auditory environment for temporal patterns.
  2. (2008). A cortical network for semantics: (De)constructing the N400. Nature Reviews.
  3. (1998). A MEG analysis of the P300 in visual discrimination tasks. Electroencephalography and Clinical Neurophysiology,
  4. (1999). A rose by any other name: long-term memory structure and sentence processing.
  5. (2007). A special role for the right hemisphere in metaphor comprehension? ERP evidence from hemifield presentation.
  6. (1992). A theory of humor elicitation.
  7. (1998). A theory of humor.
  8. (1972). A two-stage model for the appreciation of jokes and cartoons: an information-processing analysis. In
  9. (2005). Accessing world knowledge: evidence from N400 and reaction time priming.
  10. (2005). An ERP study of P600 effects elicited by semantic anomalies.
  11. (2007). An fMRI investigation of the neural correlates underlying the processing of novel metaphoric expressions.
  12. (2009). An fMRI study of processing novel metaphoric sentences.
  13. (1967). analysis of presentday American English.
  14. (1990). Appreciation of metaphoric alternative word meanings by left and right brain-damaged patients.
  15. (2002). B o o t h ,J .R . ,B u r m a n ,D .D . ,M e y e r ,J .R . ,G i t e l m a n
  16. (2007). Beyond the sentence given.
  17. (2007). Big brown dog or brown big dog? An electrophysiological study of semantic constraints on prenominal adjective order.
  18. (1994). Brain potentials elicited by garden-path sentences: evidence of the application of verb information during parsing.
  19. (2008). Brain states measured as the P3. In A. Ikeda & Y. Inoue (Eds.), Event-related potentials in patients with epilepsy (pp.
  20. (2000). Cognitive and emotional influences in anterior cingulate cortex.
  21. (1975). Comprehension and appreciation of humorous material following brain damage.
  22. (1986). Comprehension of humorous and nonhumorous materials by left andrightbrain-damagedpatients.Brain and
  23. (2001). Conflict monitoring and cognitive control.
  24. (1999). Cortical surface-based analysis: I. Segmentation and surface reconstruction.
  25. (1999). Cortical surface-based analysis: II. Inflation, flattening, and a surface-based coordinate system.
  26. (2000). Differential effects of right- and left-hemisphere damage on understanding sarcasm and metaphor.
  27. (1999). Disrupted temporal lobe connections in semantic dementia.
  28. (2002). Distinct neurophysiological patterns reflecting aspects of syntactic complexity and syntactic repair.
  29. (1998). Distinct time courses of word and context comprehension in the left temporal cortex.
  30. (1998). Distinguishing lies from jokes: theory of mind deficits and discourse interpretation in right hemisphere brain-damaged patients.
  31. (2000). Dynamic statistical parametric mapping: combining fMRI and MEG for high-resolution imaging of cortical activity.
  32. (2002). Dynamics of visual feature analysis and object-level processing in face versus letter-string perception.
  33. (2003). Electrophysiological distinctions in processing conceptual relationships within simple sentences.
  34. (2000). Electrophysiology reveals semantic memory use in language comprehension.
  35. (2000). ERP effects of listening to speech: semantic ERP effects.
  36. (2004). Event-related brain potential studies in language.
  37. (1995). Event-related potentials and language comprehension. In
  38. (1994). Event-related potentials and stimulus repetition in direct and indirect tests of memory. In
  39. (1985). Event-related potentials, lexical decision and semantic priming.
  40. (1981). From words to meaning: a semantic illusion.
  41. (2002). Functional MRI of language: new approaches to understanding the cortical organization of semantic processing.
  42. (2001). Getting it: human event-related brain response to jokes in good and poor comprehenders.
  43. (2002). Good-enough representations in language comprehension.
  44. (2002). H a l g r e n ,E . ,D h o n d ,R .P . ,C h r i s t e n s e n ,N . ,V a nP e t t e n
  45. (2004). Handedness, hemispheric asymmetries, and joke comprehension.
  46. (2005). Hemispheric asymmetries and joke comprehension.
  47. (2005). Hemispheric contributions to lexical ambiguity resolution in a discourse context: evidence from individuals with unilateral left and right hemisphere lesions.
  48. (1999). Highresolution intersubject averaging and a coordinate system for the cortical surface.
  49. (1999). How many subjects constitute a study?
  50. (1986). Human medial temporal lobe potentials evoked in memory and language tasks.
  51. (2006). Humor and smiling: cortical regions selective for cognitive, affective, and volitional components.
  52. (1977). Humor as a tool of social interaction. In
  53. (2006). Humor comprehension and appreciation: an FMRI study.
  54. (2003). Humor modulates the mesolimbic reward centers.
  55. (1999). Humour appreciation: a role of the right frontal lobe.
  56. (1993). Improved localization of cortical activity by combining EEG and MEG with MRI cortical surface reconstruction: a linear approach.
  57. (1986). Inference deficits in right brain-damaged patients.
  58. (1990). Insights from evoked potentials into the neuropsychological mechanisms of reading.
  59. (1994). Interpreting magnetic fields of the brain: minimum norm estimates.
  60. (1990). Introduction to linear models and experimental design.
  61. (2007). Is metaphor special?
  62. (2002). Is there an anatomical basis for category-specificity? Semantic memory studies in PETand fMRI.
  63. (1995). Language-related field potentials in the anterior-medial temporal lobe: I. Intracranial distribution and neural generators.
  64. (1995). Language-related field potentials in the anterior-medial temporal lobe: II. Effects of word type and semantic priming.
  65. (2007). Late positivities in unusual situations.
  66. (2007). Laterality in metaphor processing: lack of evidence from functional magnetic resonance imaging for the right hemisphere theory.
  67. (2007). Lexical ambiguity in sentence comprehension.
  68. (2004). Linguistic focus and good-enough representations: an application of the change-detection paradigm.
  69. (2007). Localization of syntactic and semantic brain responses using magnetoencephalography.
  70. (2006). Making sense of discourse: an fMRI study of causal inferencing across sentences.
  71. (2006). Metaphorical vs. literal word meanings: fMRI evidence against a selective role of the right hemisphere.
  72. (2006). Monitoring in language perception: the effect of misspellings of words in highly constrained sentences.
  73. (2002). Monte Carlo simulation studies of EEG and MEG localization accuracy.
  74. (2007). Multiple effects of sentential constraint on word processing.
  75. (2004). Neural correlates of humor detection and appreciation.
  76. (2006). Neural localization of semantic context effects in electromagnetic and hemodynamic studies.
  77. (2007). Neural mechanisms of language comprehension: challenges to syntax.
  78. (1997). Neurophysiological aspects of language processing.
  79. (2007). On sense and reference: examining the functional neuroanatomy of referential processing.
  80. (2006). Processing stages underlying word recognition in the anteroventral temporal lobe.
  81. (1994). Psycholinguistics electrified: Event-related brain potential investigations.
  82. (2007). Questions versus statements: challenging an assumption about semantic illusions.
  83. (1980). Reading senseless sentences: brain potentials reflect semantic incongruity. Science,
  84. (2001). Recovering meaning: left prefrontal cortex guides controlled semantic retrieval.
  85. (2003). Repair, revision, and complexity in syntactic analysis: an electrophysiological differentiation.
  86. (1998). Right hemisphere activation during indirect semantic priming: evidence from event-related potentials.
  87. (2005). Right hemisphere activation of jokerelated information: an event-related brain potential study.
  88. (1999). Right words and left words: electrophysiological evidence for hemispheric differences in meaning processing.
  89. (1992). Semantic dementia: progressive fluent aphasia with temporal lobe atrophy.
  90. (2001). Semantic leaps: Frame-shifting and conceptual blending in meaning construction. Cambridge:
  91. (1993). Semantic priming and stimulus degradation: Implications for the role of the N400 in language processing.
  92. (1993). Semantic processing in the right hemisphere may contribute to drawing inferences from discourse.
  93. (2001). Sequence of brain activity underlying wordstem completion.
  94. (1999). Sources of distortion in functional MRI data.
  95. (1994). Spatio-temporal stages in face and word processing: I. Depth-recorded potentials in the human occipital, temporal and parietal lobes.
  96. (2003). Spatiotemporal dynamics of modalityspecific and supramodal word processing.
  97. (2004). Spatiotemporal dynamics of word processing in the human cortex.
  98. (1992). Story processing in right-hemisphere brain-damaged patients.
  99. (2003). Structure and limited capacity in verbal working memory: a study with event-related potentials.
  100. (2011). Surprise but not coherence: sensitivity to verbal humor in righthemisphere patients.
  101. (2002). Switching languages, switching palabras (words): an electrophysiological study of code switching.
  102. (2005). Testing the limits of the semantic illusion phenomenon: ERPs reveal temporary semantic change deafness in discourse comprehension.
  103. (2009). The anterior midline field: coercion or decision making?
  104. (2000). The effects of listening comprehension of various genres of literature on response in the linguistic area: an fMRI study.
  105. (1990). The effects of right hemisphere damage on the pragmatic interpretation of conversational remarks.
  106. (2001). The functional anatomy of humor: segregating cognitive and affective components.
  107. (1998). The neurology and evolution of humor, laughter, and smiling: the false alarm theory.
  108. (2005). The p300: where in the brain is it produced and what does it tell us?
  109. (2000). The P600 as an index of syntactic integration difficulty.
  110. (2011). The potential distribution generated by surface electrodes in inhomogeneous volume conductors of arbitrary shape.
  111. (1993). The processing nature of the N400: evidence from masked priming.
  112. (2007). The role of animacy and thematic relationships in processing active English sentences: evidence from eventrelated potentials.
  113. (1981). The role of the right hemisphere in the apprehension of complex linguistic materials.
  114. (1993). The syntactic positive shift (SPS) as an ERP measure of syntactic processing.
  115. (2003). The visual word form area: expertise for reading in the fusiform gyrus.
  116. (2001). Thematic roles assigned along the garden path linger.
  117. (2002). What does the frontomedian cortex contribute to language processing: coherence or theory of mind?
  118. (2006). When heuristics clash with parsing routines: ERP evidence for conflict monitoring in sentence perception.
  119. (2006). Where in the brain is nonliteral language?