The neurology and evolution of humor, laughter, and smiling: the false alarm theory.

Laughter (and humor) involves the gradual build-up of expectation (a model) followed by a sudden twist or anomaly that entails a change in the model--but only as long as the new model is non-threatening--so that there is a deflation of expectation. The loud explosive sound is produced, we suggest, to inform conspecifics that there has been a 'false alarm', to which they need not orient. The same logic may underlie tickling (menacing approach followed by a light non-threatening contact). Thus tickling may serve as 'play', a rehearsal for adult laughter. And lastly, when one primate encounters another, he may have always begun with a threat gesture--to bare his canines--but upon recognizing the individual as kin he may stop the grimace halfway and 'smile'. When the insular cortex is damaged, patients giggle in response to pain, presumably because they can still sense the pain ('danger') but the pain is no longer aversive ('false alarm'), thereby fulfilling the two key requirements for laughter.