----- Original Message -----
From: Basil Hall
Sent: Friday, December 28, 2012 3:48 PM
Subject: Humor Research

Dino, below are messages between me an Panksepp.(Just seen your message in the humor research forum)

Cheers Basil

Jaak, I emailed you a few years ago concerning laughter. I really do believe you and Davila-Ross have it wrong. If human laughter is an exapted displacement activity, as I believe it to be, then the original laughter must have been a response to an emotional conflict.(Initially a fight or flight conflict). If you have both the time and the will to read my essay:

Laughter as an exapted displacement activity: the implications for humor theory and neuropsychology and neurophysiology in general

I think you will find it convincing, if indeed you are still open to the possibility that your "joy" approach is erroneous. My essay can be found at:

Davila-Ross refused to accept that the initial driving force behind laughter was the conflict between fear and the cognitive appreciaion that there was nothing to be feared.


Basil Hall

A small extract from my essay;

A discussion of laughter would not be complete without an examination of the laughter inducing phenomenon we call tickling. If we cannot induce laughter by tickling ourselves it is obvious that laughter is a response to being touched by someone or something else. It would be quite disruptive if every time our hands accidentally brushed our bodies we drew back in fright and we have enough body maps in the brain (both in the cortex and cerebellum) to tell us when two parts of our body are in contact, and such contacts are ignored. We can also detect the difference between touching and being touched - the difference between moving part of our bodies on a foreign object and foreign objects moving on our bodies. Anything moving on our bodies is probably alive and thus a potential danger. One only has to have an insect running about under the bed sheets to know how quickly and emotively we react to being touched in a certain way.

Not everyone is ticklish, but those who are have little conscious control over their reaction to being tickled. Laughter induced by tickling is indicative of a conflict between our emotive system, which is sending out danger signals, and our appreciation that no threat to our organism exists.

The fact that being tickled on the bottom of the feet and the ribs elicits the most vigorous response suggests that these areas were particularly vulnerable in our primate ancestors. A rapid withdrawal of the foot from an unseen, sharp object, or an animal, on the ground would minimize damage or the effect of a sting or bite. The fear reaction of all primates to snakes, and their specific warning calls that signal a snake's presence, suggests a long history of primate predation by large snakes: some writers even view the relationship between primates and snakes as being of prime importance in our evolution (Isbell, L.2006). Primates are particularly vulnerable to large, night hunting, constrictor snakes, and as constrictors target the chest area to prevent their prey breathing, a rapid reflex response to being touched in this area would have been of significant survival value to our ape ancestors.
Significantly, the bottom of the feet, for most of the time, and the whole of the body on moonless nights, cannot be seen, and so sight is useless in immediately determining the source of the contact so that the appropriate action can be taken. In fact, any cognitive interference with the reflex response could prove to be fatal.

When dealing with snake predators silently hunting in the dark a primate cannot resort to its senses of sight and hearing and it is probable that the system that immediately responds to particular types of touching functions independently of the other senses. The immediate response to tickling is a form of startle, an instantaneous escape reflex that gives rise to a secondary brain mediated fear response. This explains why, even though people can see what or who is tickling them - friend, stranger or machine - the reaction is the same. What an individual hears or sees during a bout of tickling cannot directly lead to the inhibition of the tickle response. However, the reality of the situation is registered, and the conflict between the tickle response and the cognitive grasp of the situation leads to the disinhibition of laughter. Displacement activities inhibit redundant motivations, but in the case of tickling there is a problem as every new movement of the fingers on a person's body rekindles the response and a continual cycle of motivation (fear), disinhibition (of the laughter process) and inhibition (of fear), takes place.

Hello Jaak,

First of all an extract from a paper that, does not disprove your ideas but does bring them into question. (My highlight)

The suggestion that ultrasonic 50 kHz vocalizations by rats might reflect a specific form of positive affect such as ‘‘joyful affect’’ is intriguing. On the other hand, 50 kHz vocalizations also are emitted by rats in aggressive situations when one rat invades the home territory of another (Haney & Miczek, 1993;Thomas, Takahashi, & Barfield, 1983). An aggressive situation seems potentially complex, and might involve both negative and positive processes, or even mostly negative ones. If so, one might hesitate to infer that an intruding rat is in a state of unalloyed ‘‘joyful affect.’’ In any case, the story of 50 kHz vocalization continues to unfold, and more information will be useful to evaluate its affective significance.

As I point out in my essay, it is the use of what I call cultural language (everyday, default language) that often biases the thinking of researchers and possibly sends them on the wrong track. The word "joy" is a cultural word, and its meaning immediately biases our thinking towards the positive - feeling good is positive. Yes, on a psychological level; but my contention is that on a neurological level, and in its basic state, the cause of laughter is a negative process - a displacement behavior brought about by the confliction of opposing emotional states. Why would chimps laugh when being chased, grabbed and touched in a certain manner by other chimps when being chased grabbed and touched by a different species would give rise to fear.To put their laughter down to "the joy of play" gets us no nearer to finding "evidence at the primary-process level ". Why do chimps laugh? Why do some people laugh when nervous or in disbelief when in an unbelievable but potentially dangerous situation.?

If one accepts that laughter is a displacement activity, then I believe it explains much about its evocation in many different situations and impinges on many aspects of brain function.

I have never written that the tickle induced FM 50 kHz USV's, or any other form of laughter, reflects a negative state. What I have said is that the neurological process that gives rise to laughter is negative in nature. ( a form of inhibition rather than promotion as against the undesirable rather than desirable )

Some, if not most researchers, view laughter as reflecting a positive state, rather than the laughter process producing the positive state. The fact that during laughter the "reward" system is activated does not negate a displacement theory of laughter, nor does it disprove the idea that negative processes can give rise to what can be viewed as a positive state.

In the same manner as "joy" has no place as a descriptive word in neurology, neither has the word "reward" (another cultural word). Being a cultural word "reward" biases our expectations of the nature of what goes on in various areas of the "reward" system. The fact that the "reward system" is activated during the laughter process in no way proves that that the process itself is not negative in nature. I suspect that any need or problem that is resolved, or in the case of the joke (I have already given my explanation of tickling laughter) conflict that cannot be resolved, but can be voided by displacement, results in some form of activation in the reward areas, probably mediated by an opioid.

One last point. Although I am willing to accept that the 50 kHz vocalization of rats is a form of vocal displacement, it is analogous to primate laughter not homologous.


Basil Hall

----- Original Message -----

From: "Jaak Panksepp" <>

To: "Basil Hall" <>

Sent: Saturday, February 18, 2012 5:47 PM

Subject: RE: Laughter and humor

Hello Basil,

There are many theoretical approaches to this fascinating topic, and there is surely a dark side to human laughter. . . but I expect that arises at the tertiary-process level (cognitive), and not at the primary-process level (affective) where we seek to understand this mystery. Here the data always has to trump theory. Everywhere in the brain where we evoked rat FM-chirping with deep brain stimulation, animals treated the stim as a reward:

Burgdorf, J., Wood, P.L., Kroes, R.A., Moskal, J.R., & Panksepp, J. (2007). Neurobiology of 50-kHz ultrasonic vocalizations in rats: Electrode mapping, lesion, and pharmacology studies. Behavioral Brain Research. 182(2): 274-283.

A variety of other experimental psychobiological analyzes have gone the same way (Davila-Ross has never done anything remotely like that, so our evidential base is vastly different than hers and should not be conflated). We have direct expermetal evidence for positive affect, she does not.

I am as skeptical as the next scientists, and realize how theory can be more compelling to many. I have to go with the weight of well collected evidence over any theory. Your's is well thought out and fascinating from certain philosophical and theoretical perspectives, but where is the evidence at the primary-process level? Our arguments are based on attempting to work at that level, so you would have to point me to concrete data that would help me change my mind (it always does, if there is any, for I subscribe to negations at the only way to sustain an honest understanding of nature).

I also love Schopenhauer, but all philosophers make many mistakes. Affectively relevant evidence right now suggest that the tickle induced FM 50 kHz USV's, reflect a positive rather than a negative state. Don't know of any data that negates that provisional conclusion so far. Do you?


Jaak Panksepp

Please note, I am no longer at this address, so all e-mail should be sent to:

Jaak Panksepp
Dept. of VCAPP, College of Veterinary Medicine
Washington State University
Pullman, WA 99163

Jaak, the following is to be read in rtf as I have typed in different colors and also highlighted some words. Thanks for the papers you sent me , I found them very informative. I hope what I have written below helps to clear up some misconceptions that may have arisen due to the brevity of my communications.

---Every data-based conclusion in science is open to question. But here, with a hierarchical organ like the brain, one must always consider if folks are talking past each other, by discussing different levels of neural organization.

Jaak, the bottom line is that, in the case of laughter, (and for the present I am only dealing with laughter) we agree that the result of the psychoneural processes concerned with laughter/chirping is, for want of a better word, "rewarding" in nature (as I see it, the secretion of an opioid). We differ in, as I see it, the all important question of the nature of these processes, which can be boiled down to: does laughter reflect a positive affect or is the laughter process instrumental in bringing about the positive affect?.

Indeed, "displacement" is a vague concept, just like "joy", that can be used in many different ways.

I find this comparison a little unfair in the context of my essay. The word "joy" is clearly "understood" on a cultural level and is only "vague" in the context of psychoneural theory. On the other hand the word "displacement", however ill-chosen by its original users it may be, does designate a psychoneural process. And not only this, in my the essay I acknowledge this point, give a standard definition and even update the standard definition, later distinguishing this type of displacement from the "displacement" of the "killing the messenger" type.(I have noticed that some individuals writing on the topic have failed to see the difference) It is not actually vagueness to which I object - I object to all the cultural words: joy, reward, wanting, liking and seeking, as they represent biasing presumptions on the part of the authors concerning the nature of the neurological states they are attempting to understand. I know we are all guilty of this; it is almost impossible to write in a scientific manner without using cultural terms in an attempt to elucidate our ideas. I am hot on this topic as I believe that the fathoming of the nature of laughter and humor has been handicapped by such biases - as I explain in my essay.

I am glad you included a table of rebuttals in your paper (2011) as a displacement theory predicts all the facts you have listed. Below are notes, mostly extracted from my essay, that will lead in to my critique of both sides of the arguments you have listed in your paper. Remember when reading the following that it is only my opinion and I am quite happy to be proved wrong by a hypothesis that better suits the facts and has the backing of experimental evidence.

Human displacement activities take the same form as those exhibited by other vertebrates and are associated with change, or anticipation of change, in activity, internal conflict, or motivational ambivalence (Schniter, 2001). Laughing and crying fit particularly neatly within these criteria, suggesting that laughing and crying are exaptations of antecedent vocal displacement activities. (Lorenz 1963; Russell 1996; Kozintsev and Butovskaya 1999) The motivational ambivalence and internal conflict aspects of the phenomenon are suggestive in explaining the efficacy of the particular structure of jokes in precipitating laughter.

The neural processes that produce the laughter vocalization in humans I consider to be the same as those in animals that produce similar vocalizations, but the underlying psychoneurological processes that elicit laughter I view as having changed with changes in the complexity of the hominid cognitive and emotive systems. The origins of human laughter can be traced back to the play fighting of primates in which the reduction of emotive neural activity during bouts of vocalization allowed them to practice maneuvers without harming each other.

The circumstances in which chimpanzees "laugh" are limited to play-fighting, tickling and chasing, and although the "this is an attack / this is not an attack" conflict may have laid the foundation for the joke format, it is obvious that the variety of circumstances in which humans laugh requires further explanation. The whole question appears to center around the brain's processing of conflicting cognitions and emotions, and I propose that play-fight vocalization was exapted in the hominid line to include the disinhibition of the basic laughter response during decision making, particularly in fight or flight situations. Ramachandran V. The neurology and evolution of humor, laughter, and smiling: the false alarm theory. Med Hypotheses 1998 Oct, 51:351-4

Jaak, in the extract you have already received, concerning tickling, I have pointed out how responses to stimuli, which are both frequent and fatal for a species, tend to become reflexive, thus cutting out cognitive responses that might delay an escape reaction. Two of these stimuli are being chased and being grabbed. As with tickling these stimuli are outside primate’s repertoire of positively affective, slow, direct stress relieving behaviors, such as cuddling and grooming. Being chased and being grabbed, immediately gives rise to an involuntary fear response. Human and ape youngsters, recognizing that such activities are not serious, but having less control of their emotions than adults and, during chasing, a high state of arousal, will emit loud FM laughter.

In this essay I define the neurological basis of human laughter as the disinhibition of a stereotyped behavior in response to the opposition or redundancy of the neural processes responsible for the expression of negative (fear-based) behaviors.

Table 1 (Panksepp)

Non-affective hypotheses of 50 kHz USVs emission with their rebuttal

I. 50 kHz USVs are an artifact of locomotor activity-induced thoratic compressions (Blumberg, 1992). Only 10% of 50 kHz USVs were coincident with thoratic compressions, and 50 kHz USVs could be dissociated from locomotion (Panksepp and Burgdorf, 2003).

I agree with you here.

II. 50 kHz USVs are a non-affective contact call (Schwarting et al.,2007). Flat 50 kHz calls appear to be a contact call, occurring at the highest rates during non-positive affective social interactions (Burgdorf et al., 2008).However, FM 50 kHz calls appeared to be selective for positive affective social

I disagree with both sides here. The calls may be flat, but this does not change the fact that these are 50kHz calls. Human laughter has many forms but all are still considered (other than non-Duchenne laughter) genuine laughter. The flattest type of laughter in humans is probably nervous laughter which occurs when individuals are in situations that make them somewhat anxious.

Although many people have their own particular style of laughter, the range of laughter types in humans is, in the main, dependent on the level of ambivalence and the degrees of fear and arousal a specific situation evokes. The loudest FM laughter is precipitated by, what are in reality, benign situations that for various reasons give rise to ambivalence and/or mild fear and a high level of arousal, as in the case of a child being chased, or a highly effective joke. In the case of rat explorative situations, the flat chirping can be viewed as the result of conflicting motivations with a relatively low level of the fear component.

III. 50 kHz calls are evident during aggression (Berridge, 2003). 50 kHz calls occur primarily before the onset of aggression, and the vast majority the 50-kHz calls were of the non-affective flat variety (Panksepp and Burgdorf, 2003; Burgdorf et al., 2008)

This situation mirrors the displacement behaviors exhibited by many species of male birds when they rival each other for mates or territory. For example, two male birds may preen themselves, or peck at the ground before fighting, or between bouts of fighting, for the ownership of a territory or a mate. In the case of the rats, the 50 kHz calls stop when they begin to fight as the calling is a vocal displacement behavior that temporarily puts on hold the decision to flee or fight. Once either option is taken the chirping ends.

In humans, the volume and the level of the modulation of laughter are indicative of the degree to which emotive responses are being inhibited and the extent to which the arousal, engendered by the various situations they find themselves in, is being lowered. One result of loud highly modulated laughter (often characterized as mirthful laughter) in humans is a drop in muscle tone. The specific circumstances facing the confrontational rats demands that fight or flight is put on hold until they weigh up their chances of success, giving the weaker rat time to back down and avoiding a fight that may result in injury and death( a strategy common to many male mammals). Obviously a lowering of arousal could be fatal in these circumstances, and it is probable that the flat chirps are indicative of the temporary inhibition of a physical response but the maintenance of the bodily readiness (arousal) to effectively carry it out.

IV. 50 kHz calls reflect a non-positive affective "wanting" state (Schwarting et al., 2007). 50 kHz USVs were increased in the anticipation of delivered reward, which in humans has been shown to elicit a positive affective state (Knutson et al., 2001). However, during extinction bursts of "frustrative non-reward" such appetitive behavior decreased rates of 50 kHz calls and increased rates of aversive 22 kHz calls (Burgdorf et al., 2000).

Human displacement activities take the same form as those exhibited by other vertebrates and are associated with change, or anticipation of change, in activity, internal conflict, or motivational ambivalence (Schniter, 2001).

Robbins, T and Koob, G. ( Selective disruption of displacement behaviour by lesions of the mesolimbic dopamine system. Nature Vol. 285 5 June 1980 p 409-412) and

It would be interesting to repeat Robbins and Koob’s experiments to see if the adjunct drinking was accompanied by 50hz chirping or, being a appetitive/intake type of behavior, would the presence of water have afforded the rats a displacement behavior more appropriate to the situation than chirping.?

V. Adult and infant rat ultrasonic calls reflect a state of high arousal that is not specific to positive affective states (Bell, 1974). Highly arousing aversive stimuli such as predatory odor, foot shock, and bright light, decrease rates of 50 kHz calls, whereas rewarding stimuli increase rates of 50 kHz calls (Knutson et al., 2002a,b)

Here I agree that high arousal on its own is insufficient to produce laughter/chirping.

In humans the joke mechanism creates the brain state that initiates the disinhibition of laughter - the volume and duration of which is dependent on the level of bodily arousal at the time of disinhibition. Berlyne believed that over-arousal was enough to cause laughter (Berlyne 1960), and Schachter and Wheeler found that pre-injections of epinephrine (which increases arousal) increased laughter when subjects watched a slapstick comedy film (Schachter, Wheeler 1962). Berlyne's theory was not validated by experiments carried out by Godkewitch (Martin 2007), but Schatcher's and Wheeler's experiment suggests that, although laughter may not be disinhibited purely due to high arousal, it does fuel laughter after its disinhibition.

If chirping is contagious as laughter is in humans then mass chirping in families of rats is easily explained. "Completely letting go" - emitting loud FM chirps -is only appropriate and safe when the rats are in situations where they can afford to lower stress, (lower their state of arousal) i.e, when they are all together in a danger free environment.

50HzCalls from pups when reunited with the mother or group is mirrored by a human child’s laughter in similar circumstances.

The laughter of a young child on the return of a primary caregiver after a short separation can be viewed in a similar light. The reappearance of a trusted individual makes the neural activity underlying separation anxiety redundant - the child's laughter nullifying the fear response and also inducing laughter in the caregiver, which aids in the reaffirmation and strengthening of mutual bonds.

( In this essay I define the neurological basis of human laughter as the disinhibition of a stereotyped behavior in response to the opposition or redundancy of the neural processes responsible for the expression of negative (fear-based) behaviors.)

In a way the individuals whose views are antagonistic to yours are partly right. As I view the situation, the 50 Hz calls are not positive-affective (a response to a positive emotion) but neither are they non-affective ( not a response to emotion) but anti-affective (moderating negative emotion) which is registered as a "reward"..

As you can see I am not denying any of the conclusions you have come too through your experimental results but putting forward a hypothesis concerning the processes that give to "rewarding" situations.

Neither do I disagree with your statement : I think the evidence for experiential functions, even in animals is enormous. We do tend to overestimate ourselves and under estimate animals, but I do baulk at the usage of certain terms that enthusiasts use in an effort to bridge the gap between ourselves and animals. I refer to the use of the word "moral", by some writers, when applied to animals. Again, "moral" is a cultural word, a human judgment of the behavior of human individuals, the scope of which can change with time and circumstances, and differs from one society to another. I do not think the term is particularly apt when applied to the undoubtedly altruistic responses of some non-verbal species. But that is another discussion.

Kind Regards



Jaak, What are your thoughts on this video. "Laughing" penguin


DINO, do we have to go further back in evolution to find the common ancestor of birds and mammals. Seeing that birds and mammals evolved from two different reptilian groups then we have to take the evolution of "laughter" back to the ancestors of the dinosaurs. If indeed the noises made by animals when played with have the same neurological basis as human laughter.


Jaak, I think we will have to wait for further experimental work to be done before the situation can be clarified. If it can be proved that laughter and chirping are indeed displacement behaviors ( have identical neurophysiological correlates as known displacement behaviors) then all that can be said is that some animal groups (including rats) utilize a vocal displacement behavior in certain situations to moderate fear and thus the arousal/stress engendered by the endocrine system.

I cannot think of a vertebrate group that does not exhibit displacement behaviors, even some invertebrates, such as the octopus, do so. In the end I cannot support your contention that there is an evolutionary relationship between rat vocalizations and the "joy" commonly accompanying human social play, other than the fact that rats and humans share a form of stress release through a process common to all vertebrate groups. But time will tell.

Kind regards


Just found this paper;

No Laughing Matter: Intranasal Oxytocin Administration Changes Functional Brain Connectivity during Exposure to Infant Laughter.


1] Centre for Child and Family Studies, Leiden University, Leiden, The Netherlands [2] Leiden Institute for Brain and Cognition (LIBC), Leiden University, Leiden, The Netherlands.

Riem MM, van Ijzendoorn MH, Tops M, Boksem MA, Rombouts SA, Bakermans-Kranenburg MJ.


Infant laughter is a rewarding experience. It activates neural reward circuits and promotes parental proximity and care, thus facilitating parent-infant attachment. The neuropeptide oxytocin might enhance the incentive salience of infant laughter by modulating neural circuits related to the perception of infant cues. In a randomized controlled trial with functional magnetic resonance imaging we investigated the influence of intranasally administered oxytocin on functional brain connectivity in response to infant laughter. Blood oxygenation level-dependent responses to infant laughter were measured in 22 nulliparous women who were administered oxytocin and 20 nulliparous women who were administered a placebo. Elevated oxytocin levels reduced activation in the amygdala during infant laughter and enhanced functional connectivity between the amygdala and the orbitofrontal cortex, the anterior cingulate, the hippocampus, the precuneus, the supramarginal gyri, and the middle temporal gyrus. Increased functional connectivity between the amygdala and regions involved in emotion regulation may reduce negative emotional arousal while enhancing the incentive salience of the infant laughter.

Neuropsychopharmacology advance online publication, 21 December 2011; doi:10.1038/npp.2011.313.