




TWO polar bears are perched on a block of floating ice. One says to the other: “Do you know, I keep thinking it’s Thursday…”
To some, this kind of surreal humour is side-splitting. Others are baffled by it and can’t even raise a smile. Yet despite the importance of humour to human psychology, it is only the advances in brain imaging during the past decade that have enabled neuroscientists to pin down how the brain reacts when a joke tickles us. Armed with this knowledge, they are now solving the puzzle of why some jokes are funny to some people but leave others cold.
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So what is a joke, exactly? Most theories agree that one condition is essential: there must be some kind of incongruity between two elements within the joke, which can be resolved in a playful or unexpected way.
Take the following exchange from the classic British sitcom Only Fools and Horses, when an anxious “Del Boy” Trotter visits his doctor for a heart check-up. “Do you smoke, Mr Trotter?” asks the doctor. “Not right now, thank you doctor,” he responds.
The joke’s incongruity, of course, lies in the unlikely offer of a cigarette by a doctor to a patient concerned about his heart. It is only once we understand the mismatch that we get the joke. “Humour seems to be a product of humans’ ability to make rapid, intuitive judgements” about a situation, followed by “slower, deliberative assessments” which resolve incongruities, says of Duke University in Durham, North Carolina.
But which parts of the brain carry out these processes? To find out, , then at Dartmouth College in Hanover, New Hampshire, used functional MRI to scan the brains of volunteers while they watched popular TV sitcoms. The experiments revealed a distinct pattern of neural activity that occurs in response to a funny joke, with the left posterior temporal gyrus and left inferior frontal gyrus seeing the most activity. These regions are normally linked to language comprehension and the ability to adjust the focus of our attention, which would seem to correspond to the process of incongruity-resolution at the heart of a good joke ().
Further research, conducted by , then at Stanford University in California, uncovered a second spike of activity in the brain’s limbic system – associated with dopamine release and reward processing – which may explain the pleasure felt once you “get” the joke ().
Examining one particular part of the limbic system – the ventral striatum – was especially revealing, as its level of activity corresponded with the perceived funniness of a joke. “It’s the same region that is involved in many different types of reward, from drugs, to sex and our favourite music,” says Mobbs, now at the MRC Cognition and Brain Sciences Unit in Cambridge, UK. “Humour thus taps into basic rewards systems that are important to our survival.”
Yet humour is a far more complex process than primeval pleasures like sex or food. In addition to the two core processes of getting the joke and feeling good about it, jokes also activate regions of the frontal and cingulate cortex, which are linked with association formation, learning and decision-making (). The team also found heightened activity in the anterior cingulate cortex and the frontoinsular cortex – regions that are only present in humans and, in a less developed form, great apes. Indeed, the fact that these regions are involved suggests that humour is an advanced ability which may have only evolved in early humans, says Watson, who conducted the research.
“Humour is a far more complex process than primeval pleasures like sex or food”
No two brains are the same, however, and how these differences are reflected in our sense of humour is the subject of much research. Men and women, for example, seem to process jokes slightly differently. Although both sexes laugh at roughly the same number of jokes, women show greater activity in the left prefrontal cortex than men (). “This suggests a greater degree of executive processing and language-based decoding,” says Mobbs. As a result, women take significantly longer than men to decide whether they find something funny, though that doesn’t seem to spoil their enjoyment of the joke. Indeed, women show a greater response in the limbic system than men, suggesting they feel a greater sense of reward.
“Women take significantly longer than men to decide whether or not they find something funny”
Perhaps unsurprisingly, personality also appears to play a key role in humour. Mobbs has shown that people who are classed as extrovert and emotionally stable have increased activity in reward areas of the brain during exposure to funny stimuli. Neurotic people, in contrast, have less of a reward response compared with the average person (). “This suggests that personality style may be important in how we process humour,” Mobbs says.
Twisted logic
Whether our neural circuitry can explain specific preferences for certain types of humour remains an open question. To investigate, at the University of Fribourg in Switzerland used MRI to scan volunteers’ brains while they looked at 90 different non-verbal cartoons reflecting various styles of humour. As a control, the volunteers also viewed pictures that could not be interpreted in any meaningful or funny way.
Surprising results emerged from that experiment. Although you might expect the subject matter – music or politics, for example – to determine joke preference, Samson found that it is the way a joke is solved that is most important. “The logic by which the incongruity is resolved matters most, in terms of what kind of person a joke appeals to,” she says (see “What your sense of humour says about you”).
There is a serious note to this work. The researchers hope that pinning down the brain processes involved in understanding jokes could shed light on a number of medical conditions. Mobbs, for example, hopes that studying humour will provide insights into depression. “It is believed that the reward system is disrupted in depression and it would be interesting to see if this deficit extends to more complex social processes such as humour,” he says.
Samson, meanwhile, hopes it could contribute to our understanding of autism. Previous research has suggested that people with autism have difficulty understanding comedy, but her work shows that they can understand and appreciate certain types of jokes as well as anyone (see “The mechanics of a joke”). This could change the way we interact with autistic children, she says.
More than anything, the recent research confirms the fact that humour, an oft-neglected trait when considering our cognitive skills, requires a tremendous amount of brain power. “Getting a joke would seem – on the surface – to be a very trivial, intuitive process. But brain imaging is showing us that there is more going on than we might think,” says Samson.

What your sense of humour says about you
Most types of humour, including jokes and cartoons, rely on some kind of incongruity between two elements that needs a second’s thought before it can be understood. The extent to which this mismatch can be resolved differs between jokes, however.
Some have a clean punchline that ties up all the loose ends, while in “nonsense” humour the incongruity can only be partially resolved, leaving a gap in the person’s understanding. The cartoons to the right should give some idea of the difference between the two styles of joke.
For years, nonsense jokes have been considered to be more sophisticated and philosophical than classic, resolvable humour (known technically as “incongruity-resolution humour”) – consider the reputation of Monty Python’s Flying Circus compared with that of Friends, for example. “It was previously thought that nonsense humour was more complex in terms of thought process,” says psychologist Andrea Samson at the University of Fribourg, Switzerland.
Samson’s recent work suggests otherwise. When comparing MRI scans of people as they viewed both straight and nonsense humour, she found that straight humour evoked significantly more brain activity than a surreal joke in most volunteers. “Making sense out of opposed scripts and integrating this information seems to be a more complex process than simply laughing about nonsense,” she says.
The degree to which Samson’s volunteers “got” the joke was reflected in one small region of the brain called the temporoparietal junction (TPJ), with the most activity occurring when the resolvable cartoons were viewed but no activity for the unfunny control images (). The surreal cartoons fell somewhere in between.
“Although the attempt to resolve the incongruity is present with nonsense humour, this effort does not lead to a complete resolution of the incongruity and therefore to less activation of the TPJ,” says Samson. What’s more, if someone failed to get the joke, the rostral cingulate zone of the brain became more active – a region thought to pick up on errors in the way we behave and monitor conflicts.
Experience Seekers
Not everyone reacted more strongly to resolvable humour, however; those with one particular personality type found the surreal cartoons more rewarding. These people, dubbed “experience seekers”, are defined by a desire to pursue novel sensations, stimulation and experiences, whether it’s through art, travel, music or an unconventional living style.
When processing any type of funny cartoon, experience seekers showed greater activity in the TPJ, hippocampus and prefrontal areas of the brain than their fellow subjects, which might reflect their adventurous mindset, says Samson.
“The hippocampus is an area known to process novel stimuli,” she says. “It could be that humorous stimuli give experience seekers an opportunity for mental exploration of novelty, and this ‘lights up’ the hippocampus.”
Indeed, a previous study at the University of Kentucky in Lexington found that experience seekers have greater hippocampal volume, which would seem to fit with this result ().
The difference was most marked when the experience seekers viewed the surreal cartoons. Importantly, unlike the other subjects, their brains responded most strongly to the nonsense humour rather than the incongruity-resolution humour.
Samson reckons that the nonsense humour may allow the experience seeker’s inquisitive brains even more opportunity for exploration than the resolvable humour, which could explain their preference.
The mechanics of a joke
Most jokes can be divided into certain “logical mechanisms” that determine which cognitive process your mind goes through before it understands the humour.
Many cartoons, for example, rely on our understanding of other people, playing on the fact that one character doesn’t understand what the other is thinking. To get the joke you need a “theory of mind”, allowing you to understand the different state of mind of each character. Perhaps unsurprisingly, brain scans have shown that areas involved in social cognition are activated when viewing this kind of cartoon.
The degree to which we empathise with others has a profound impact on our appreciation of this kind of joke. Andrea Samson at the University of Fribourg, Switzerland, showed this when she compared the responses of people with two different personality types. It turned out that “empathisers”, who identify emotions and thoughts in others and respond appropriately, found the theory-of-mind jokes much funnier than “systemisers”, who prefer to think about things in logical, abstract terms.
At the far end of this scale are people with autism, who have an impaired ability to empathise with other people. Some previous studies had found that people with autism have trouble understanding jokes, but since these studies hadn’t considered different styles of humour, it wasn’t clear whether they were unable to understand all kinds of humour, or whether it was simply theory-of-mind style jokes that had them stumped.
Samson decided to investigate. She found that while volunteers with Asperger’s syndrome had difficulty understanding and appreciating theory-of-mind-based cartoons, they enjoyed visual puns, which do not rely on empathy, to the same extent as a control group. “Visual puns are much more abstract than theory-of-mind cartoons,” says Samson. “To understand the joke, you have to realise that one visual element refers simultaneously to two meanings.”
Some researchers had suspected that an element of empathy is needed for all kinds of humour – not just theory-of-mind jokes. But the fact that people with Asperger’s syndrome get these visual puns shows that they don’t lack an overall sense of humour, says Samson, just that they are poorly equipped to “get” a certain type of joke.