The three articles on laughter describe the neural substrates of laughter, the purported social

functions of laughter, and the practical applications of laughter in pedagogical or clinical

settings. Berns (2004) emphasizes the 'hedonic' element of laughter. He reviews evidence that the

viewing of humorous cartoons is associated with the activation of mesolimbic reward systems.

He notes, however, that reward-system activation is not ubiquitous in studies of the neural

correlates of humour. Panksepp (2007) confirms the role of mesolimbic reward structures in

laughter through reference to his own research on rats. Panksepp draws a parallel between

human laughter and the 50khz ultrasonic vocalizations of rats. His analogy is based on

observations that the rats' vocalizations were observed in similar contexts to human laughter, and

seemed to serve similar functions, such as the encouragement of playful fighting and the

encouragement of social interaction. The analogy is also supported by similarities in the neural

correlates of laughter and 50khz USVs. Panksepp argues that we can learn about human laughter

by assuming that similar phenomena in rats indicate an analogous affective experience which

serves a similar purpose. Thus, we can apply our understanding of the neural processes and

social functions related to the 50khz USVs of rats to our study of human laughter. He denies that

our inability to empirically confirm the existence of affective states in animals precludes drawing

analogies between affective processes in animals and humans.

While I do not see a problem with inferring that animals have something resembling our

own affective states based on the similarities of our brains and behaviour, I think that it is

problematic to assume that we can understand those affective states by analogy to our own. Rat

affect is likely very different from our experience of emotion. Panksepp theorizes that rats

experience basic 'primary-process' emotions, and that humans experience primary-process

emotions and more complex, cognitively-moderated emotions. The problem with his
characterization is that it is unlikely that a socialized adult human ever experiences primary-

process, cognitively unmoderated emotions. Our emotions are so bound up with our cognitions

that we are incapable of imagining a non-cognitive emotion. Consequently, calling rat

vocalizations "laughter" can only lead to conceptual confusion. To take a basic, empirically-

based observation like "humans and rats both make vocalizations that share analogous neural

substrates and have a social function" and derive from that the concept of "rat laughter" is to

enter the realm of folk psychology – to sully scientific understanding with 'common sense' ideas

of mental function.

This is not to say that animal analogies of laughter can't provide insights into the neural

mechanisms and functions of human laughter. In fact, the kind of conceptual clarity required for

such a comparison is essential for the understanding of the function of laughter in humans and

the possible application of such knowledge. Further research should tease out the specific neural

substrates of different manifestations of laughter to conceptually clarify what laughter is. Does it

always cause limbic activation? When is it associated with 'social'' brain areas? Does shared

laughter differ from 'derisive' laughter (in function or in neural correlates)? All of these are

essential to understand if our understanding of laughter is to be practically employed. Penson et

al. (2005) note the difficulty of applying humour and laughter in the context of cancer care.

Laughter has so many functions that it is readily misinterpreted. The authors caution that while

humour can be effective,it has to be used delicately. For example, attempts to facilitate social

interaction must be differentiated from humour used to moderate a doctor's own discomfort.

1. Does the sexual dimorphism of humour indicate its evolution by sexual selection?

2. Is laughter to basic a process to have a specific function?

3. Can biological understandings of laughter inform its clinical application?