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Our Ancient Laughing Brain

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30 December 2003
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Michael Ragland
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  1. Our Ancient Laughing Brain Courtesy of Cerebrum

    The Laughing Brain

    Scientists have been studying the play behavior of young animals to identify its neural basis. Some
    are also evaluating whether and to what extent early social experiences such as play or lack of it
    influence neural structure and function. Although the neural bases of laughter are not fully
    understood, we are slowly putting together a picture of the various brain regions involved in
    producing laughter.     The ancient limbic system of the brain, including two structures involved
    with emotions - the hippocampus and the amygdala - is involved in laughter, suggesting that it is
    deep in our animal nture. But our cortex, too, appears to have a role, relating laughter and speech
    and bringing humor into the equation. A. Primitive brain (archipallium) - Self preservation,
    aggression. B. Intermediate brain (paleopallium) - Limbic system, emotions. C. Rational brain
    (neopallium) - Neocortex, intellectual tasks.   Laughing in potentially aggressive or competitive
    situations disarms our social companions. On the verge of becoming our adversaries, they pick up
    from our laughter that the situation is not threatening

    The evolving human brain acquired three major parts that developed in succession and became
    superimposed (see illustration above). The archipallium, or primitive (reptilian) brain comprises
    the structures of the brain stem; the paleopallium, or intermediate (old mammalian) brain comprises
    the structures of the limbic system. The neopallium, also known as the superior or rational (new
    mammalian) brain, includes almost the whole of the hemispheres (made of a more recent type of tissue
    called the neocortex) and some subcortical neuronal groups. These three make up the brain of the
    superior mammals, including primates and humans.

    When we examine brain areas involved with laughter, the limbic system (part of the old mammalian
    brain) seems central. It controls behaviors essential to the survival of all mammals—for example,
    defending oneself and finding food. The same structures that are found in the human limbic system
    are found in the brains of much earlier animals such as lizards, snakes, and toads. In them, the
    limbic system, linked to the sense of smell, is involved in hunting, eating prey, and defending
    territory. In humans, the limbic system is more important in motivation and emotional behaviors. Two
    limbic structures involved with emotions are the hippocampus (a seahorse-shaped structure) and the
    amygdala, a small, almond-shaped structure deep in the brain. The amygdala connects with the
    hippocampus and the medial dorsal nucleus of the thalamus, enabling the amygdala to have an
    important role in mediating and controlling mood expression and emotion-laden activities like
    friendship, love, and affection. The hypothalamus, particularly its median part, has been identified
    as a key contributor to the production of loud, uncontrollable laughter.

    If laughter is such a primitive feature of the human primate, with origins in lower mammals, it
    stands to reason that its neural substrate would be in areas of the brain we share with other
    animals, probably the same places aggressive and emotional behaviors are organized.

    Neurobiologist Provine believes that laughter emerges not from our conscious mind (in other words,
    the cortex) but from a more primitive part that he calls the "pre-cognitive brain." He says, "We're
    talking about something that's very deep in our animal nature." Panksepp, too, sees evidence that
    laughter is triggered by genetically older structures of the brain. He writes that "it is not
    exactly a cerebral activity. Then laughter must be an ancient response, part of our earliest
    evolution into mammals, and is triggered by brain regions that evolved from ancient times."

    Recently, however, scientists have discovered that the brain's cortex also plays a critical role in
    creating laughter. Studying a young epileptic woman, researchers electrically stimulated a part of
    the cortex called the supplemental motor area (SMA), and found that low-intensity current induced
    her to smile (5). A higher-intensity current transformed her smile into a contagious fit of
    laughter. The SMA, associated with the control of speech, lies just behind the pre-supplemental
    motor area, which is also found in nonhuman primates. Neurophysiologists believe that the SMA is
    part of a larger brain circuit responsible for processing emotions related to humorous things: the
    cognitive part ("getting it"😉 as well as the motor part (smiling and laughing), which involves
    muscles of the face and respiratory system.

    Based on those observations, researchers conclude that there is a link among the motor, emotional,
    and cognitive components of laughter. They propose that the SMA spot they found "is part of a
    further development in humans to accommodate the specialized functions of speech, manual dexterity
    and laughter." So, not surprisingly, there is a relationship in the brain between speech and
    laughter. After all, both are communicative activities, for which we use the same muscles of the
    face and need to control the way we breathe. Primates have highly mobile facial expressions, and we
    rely on our faces more than any other body part for communication, including speech.

    Tickling is a fascinating instance of the connection between playfulness and social bonding.
    It begins in early infancy, when parents tickle their babies to evoke laughter and
    pleasurable sensations in them. Although there are a few "unticklish" children and adults,
    tickling almost always produces laughter. Probably tickling and laughter evolved in part to
    help us relate to others.

    One remarkable feature of tickling is that we do not laugh when we tickle ourselves, only when other
    people tickle us. This implies that the brain may have a different mechanism for responding to the
    two types of tickling. Charles Darwin first explored the link between tickling and social relations,
    arguing that tickling provokes laughter through the anticipation of pleasure. If a stranger tickles
    a child without any preliminaries, catching the child by surprise, the likely result will be not
    laughter but withdrawal and displeasure. Darwin also noticed that for tickling to be effective, you
    must not know the precise point of stimulation in advance. He reasoned that this is why you cannot
    effectively tickle yourself.

    Researcher Sarah-Jayne Blakemore confirmed Darwin's propositions by investigating how the brain
    distinguishes between sensations we create for ourselves and sensations others create for us. She
    used robotic arms to tickle people and found them to be as effective as real people in provoking
    laughter. When her subjects used a joystick to control the tickling robot, however, they couldn't
    make themselves laugh.6 Her studies suggest that when you try to tickle yourself, your cerebellum
    sends to your somatosensory cortex precise information on the position of the tickling target and
    therefore what sensation to expect. Apparently some cortical mechanism then decreases or inhibits
    the tickling sensation.

    Why is the brain built this way? "It may be that there is no point laughing to your own tickle
    because it is not biologically important," says Blakemore. But tickling by your playful companions
    is indeed socially important, as we have seen.

    Humor: Evolved Ticklishness?

    Quoted message said:

    From these studies an apparent contradiction arises.


        If laughter can be triggered by  stimulating highly developed cortical areas of the brain, how
    could laughter be, at the same time, a primitive feature of both human and nonhuman brains? The
    answer may lie in our prodigious human capacity for humor, which we do not seem to share with
    other animals.

    The main argument here is that laughter induced by a joke or by watching something funny may be a
    highly evolved version of the laughter induced in our ancestors by play or tickling. Provine
    suggests that the human brain is able to enlarge the capabilities of one of its ancient, more
    primitive features by processing lower functions. in higher areas. "When we hear laughter, we are
    hearing something that is very primitive. In the more modern version, laughter induced by jokes
    involves cognitive systems, something that evolved long after speech and language."

    Chimpanzees show laughterlike behavior in response to physical contact, such as wrestling, chasing,
    or tickling, but humans laugh more in response to conversational and visual stimuli. Both Darwin
    and German evolutionist Ernst Haeckel theorized that jokes were essentially "psychological
    tickling." Haeckel believed that laughter was a kind of reflex, which relieved the irritation of
    vasomotor nerves brought on by physical or mental tickling. Ethnologist Glenn E. Weisfeld, an
    expert on humor, adds that "the subject matter of humor often pertains to ticklish situations." The
    word humor is from the Latin humere, meaning "liquid" or "fluid," and betrays the Aristotelian
    belief (continued into the Middle Ages) that all emotional states were related to the essential
    four body fluids that drive us.

    One definition of humor is "the mental faculty of discovering, expressing, or appreciating the
    ludicrous or absurdly incongruous." Visual and verbal jokes often emphasize the exaggeration,
    absurdity, eccentricity, or inconsistency of a situation, a behavior, or an outcome. This
    exaggeration becomes the core element provoking laughter: the punch line.   Rat pups emit short,
    high frequency, ultrasonic vocalizations during rough and tumble play, and when tickled. They
    "laugh" far more than olders rats.

    Another key element of humor and laughter is the release after danger and surprise. When we laugh
    our heads off at someone slipping on a banana peel or getting a pie in the face, we are laughing at
    situations that involve surprise, incongruence, potential damage to the body, or an unexpected
    outcome—all stimuli that trigger fight-or-flight in our primitive, jungle-primate selves.
    Philosopher John Morreall believes that human laughter may have its biological origins as a kind of
    shared expression of relief at the passing of danger. The relaxation of tension we feel after
    laughing may help inhibit the fight-or-flight response, making laughter a behavioral sign of trust
    in one's companions.

    This connection between humor and the release of tension makes sense in brain terms. Animal (and
    human) brains do not like novelty and change; they are hardwired to produce a certain amount of
    tension when something unexpected occurs, because any environmental change could be a threat. A
    sudden noise startles us. A playful lunge at our belly by a friend provokes instinctive recoil and a
    suitably scared expression, then laughter. If the stimulus were a gunshot, however, or a real punch
    in the gut, we would most assuredly not laugh but seek cover or defend ourselves.

    Watch people laughing at an amusement park and you see this connection between release from danger
    and laughter. A boat swooping down a steep chute into water first evokes expressions of horror and
    fear. When no danger ensues, laughter erupts. In the same vein, slapstick comedies feature a lot of
    head banging, falls, and edge-of-the-cliff situations to induce laughter. We enjoy make-believe
    aggression and danger and we laugh to relieve the tension and signal to our companions that all is
    well. Weisfeld notes that "lots of humor pertains to aggressive or competitive situations in which
    someone gets hurt." See how professional comedians or clowns use stooges who are constantly
    embarrassed, hit, and harassed.   Release from danger or threat - or the civilized substitute for it
    during the amusement park plunge down the water chute - is followed by an explosion of laughter as
    we share our feelings of relief.

    Appreciating Humor

    The brain's right hemisphere appears to be important in appreciating humor (7). In particular, the
    frontal lobe processes cognitive tasks needed for us to get to the joke, including abstract
    interpretation of aspects of conversation such as irony, affective intonation, sarcasm, and
    innuendo.8 The frontal lobe is part of a wider neural circuit involved in controlling emotion. Many
    neural pathways connect it with the "emotional" part of the brain—that is, the limbic and
    hypothalamic systems.

    Recent studies by Prathiba Shammi and Donald T. Stuss at the University of Toronto suggest an
    interesting asymmetry in the frontal lobe's function in humor. It seems that the right lobe is more
    involved than the left, but why? PET (positron-emission tomography) studies have shown that the
    right hemisphere, particularly the right frontal lobe, mediates the indirect interpretation of
    information, a process key to so many forms of humor. Appreciation of humor also often requires
    interpreting current information in the light of our past experience, and the right frontal lobe is
    critical for the retrieval of memories.

    Why Laughing Is Good Medicine

    Laughter is not only associated with release of tension induced by danger and signaling
    nonaggression but also with expressing good, positive emotions. It is a social glue that facilitates
    approach, contact, and intimacy between people and decreases stress from potential conflict. This
    could be the basis for the intuitive notion that "laughter is the best medicine."

    Serious research is showing that this notion is true. Laughter and humor decrease stress and
    anxiety, reinforce immunity, relax muscle tension, and decrease blood pressure and pain. Modern
    medicine is beginning to take advantage of these positive effects; hospitalized children who see
    clown shows have shorter hospital stays than those who do not.   Laughter in humans serves as
    social glue. It helps us break the ice, get closer to people, dampen hostility and aggression, or
    soften a refusal.

    Photo by Vietnam Travel-Pictures: Hanoi, Da Nang, Hue, Saigon

    Laughter initiates a chain of physiological reactions. First, it activates the cardiovascular
    system, so heart rate and blood pressure increase. The arteries then dilate, however, causing blood
    pressure to fall. Repeated short, strong contractions of the muscles of the thoracic wall, abdomen,
    and diaphragm increase blood flow into our internal organs. Forced respiration (the ha! ha! of
    laughter) elevates the flow of oxygen into the blood. Muscle tension decreases, and we may
    temporarily lose control of our limbs, as in the expression "weak with laughter."

    People suffering from chronic anger have a higher incidence of elevated blood pressure, increased
    cholesterol levels, and heart attacks. While anger, depression, and frustration disturb the function
    of many bodily systems, including the immune system, laughter helps the immune system to increase
    the number of type T leukocytes (T-cells) in the blood, which combat damage and infection. Some
    researchers have dubbed T-cells the "happiness cells."

    Laughter may also produce beneficial hormonal changes. Scientists speculate that laughter releases
    neurochemical transmitters called "endorphins," which reduce sensitivity to pain and boost endurance
    and pleasurable sensations.

    Laughter's Social Power

    Why does laughter have such pervasive power in our lives? Beyond its physical effects, I believe
    that the answer lies in our social nature. Laughter appears to be a basic aspect of bonding.

    We are creatures who need to build stable social structures to live well. Thus we need to enjoy
    peaceful relations with the people around us. Laughter is a kind of message we send to communicate
    this joyful disposition and a willingness to play. We rarely laugh when we are alone. We even feel
    that someone laughing alone may be crazy.

    Laughter has many subtle effects on our social companions. It breaks the ice, achieves closeness,
    bonds us, generates goodwill, and dampens hostility and aggression. Observe how we laugh when we
    want to deflate tension between strangers or need to say no to someone. We often laugh when we
    apologize.

    Laughter disarms people, creates a bridge between them, and facilitates amicable behavior. Even
    babies laugh. Since they are too young to have a sense of humor, smiling and laughing must reinforce
    their connections with their parents and others close to them. Laughter's function in social
    relations may go still deeper. Studies have shown that socially dominant individuals like bosses or
    tribal chiefs use laughter to control their subordinates.

    When the boss laughs, everyone laughs. Is laughter, then, a form of asserting power? Morreall
    speculates that in this way, bosses are "controlling the emotional climate of the group." Provine
    and his colleagues observed that women in an audience laugh more often when the speaker is a man.
    Does this suggest gender differences in how we use laughter? Or reflect men's generally more
    powerful social role?

    Age differences in laughter have also been noted. Adolescents use it more when they are playing or
    flirting; executives use it more in a professional context, to increase rapport with someone or win
    a negotiation. On the other hand, laughter may have a negative connotation, even in our own culture.
    There is a widely appreciated difference between "laughing with" and "laughing at" someone.   Using
    sarcasm and scorn toward people may cast them out of a group or block their entry. Laughter then
    becomes an instrument of social hostility.

    Humor usually means trouble - preferrably for someone else. Humor and laughter can be a way to deal
    with surprise, incongruence, and threat. As tension mounts, along with all the stimuli that trigger
    the fight-or-fight response, laughter can be a release that we share with others.

    When Laughter Is Not Funny

    Some people have fits of abnormal laughter, producing an inappropriate, unrestrained, uncontrollable
    laughter dissociated from any stimulus. To observers, the laughter often appears childish or
    violent. Clinical researchers report pathological laughter in three main conditions: psychiatric
    illnesses such as hysteria and schizophrenia; pseudobulbar palsy (a disorder affecting the muscles
    that control the tongue, throat, and parts of the face); and gelastic epilepsy (a form of temporal
    lobe epilepsy with fits of mirthless laughter).9 Scientists believe all these conditions involve
    disinhibition of impulses in the higher brain stem.

    Although the neuroanatomy of pathological laughter is not fully known, three levels of the brain are
    probably involved: the cortical level, the bulbar level, and an integrative level probably at or
    near the hypothalamus. Patients with pathological laughter almost invariably have brain damage.
    Laughter may also cause neural pathology, however, leading to or aggravating some key symptom. For
    example, in a disease called cataplexy, characterized by sudden lack of muscle tone that leads to
    falls, laughter is a potent triggering stimuli.

    Collective contagious laughter is also pathological. One of the most extreme cases occurred in 1962
    in Tanganyika. It started among young girls who erupted in fits of laughter that did not stop,
    spread to neighboring communities, and became so severe that schools were forced to close. The
    epidemic persisted for almost six months and may not have been psychological—at least not
    entirely—because victims had symptoms such as fever and headache.

    The Risks of Not Playing

    We have examined how important laughter is in our lives, how it forges social bonds, how it stems
    from our playful behavior as children. In this light, the trend in modern Western societies to
    restrict the play of children is worrying. What will be the consequences, if, indeed, play is
    important in developing personality and social behavior?

    Watching television and engaging in socially isolated, individual play with video games and
    electronic toys has increasingly displaced social interaction and group play. Fear of crime, illegal
    drugs, and automobile traffic makes many parents curtail the custom of sending children outdoors to
    play freely. Parents who arrive home late from work have little time to play with their children in
    the evenings. Schools cut into playtime to cram more classes and topics into the day; they use
    detention or suspension of playtime to punish pupils for misbehaving or for getting poor grades.
    Teaching children to compete seems to have overtaken teaching them to relax, play, and socialize.

    Where will all this lead? There are indications that these shifts may have some bearing on many of
    today's social ills: increasing violence in schools, very young children using weapons and drugs,
    and social and racial intolerance. Scientists such as Panksepp speculate that the recent astounding
    increase in the prevalence of attention deficit hyperactivity disorder may be fueled by restrictions
    on play in American schools. Deprived of exercise and time to play, normal young children become
    fidgety; their attention spans decrease.

    A Better World of Laughter

    Laughter and play are important in our human behavioral repertoire, a response both ancient and
    modern. When we laugh spontaneously, we use a capacity rooted in our most primitive biology. When we
    laugh at a subtle joke, we build on this primitive process, employing an advanced capacity of our
    brains. In both cases, our laughter has a surprisingly important role. Panksepp summarizes this with
    wisdom and beauty:

    Play and laughter not only fertilize the brain but they fertilize the human spirit. These are
    the types of systems that allow us to be joyous, sharing creatures that do the right things in
    the world—or usually do the right things. If other people are interacting with us in positive
    ways and we respect the way they feel and they respect the way we feel, if we read each other's
    minds correctly, then I think we have a better world. And play and laughter are a big part of
    that.

  2. Michael Ragland said:

    Our Ancient Laughing Brain Courtesy of Cerebrum

    The Laughing Brain

    Scientists have been studying the play behavior of young animals to identify its neural basis.
    Some are also evaluating whether and to what extent early social experiences such as play or lack
    of it influence neural structure and function. Although the neural bases of laughter are not fully
    understood, we are slowly putting together a picture of the various brain regions involved in
    producing laughter.     The ancient limbic system of the brain, including two structures involved
    with emotions - the hippocampus and the amygdala - is involved in laughter, suggesting that it is
    deep in our animal nture. But our cortex, too, appears to have a role, relating laughter and
    speech and bringing humor into the equation. A. Primitive brain (archipallium) - Self
    preservation, aggression. B. Intermediate brain (paleopallium) - Limbic system, emotions. C.
    Rational brain (neopallium) - Neocortex, intellectual tasks.


    [snip]

    This is the "triune brain" hypothesis, first enunciated by Paul Maclean between 1949 and 1970 and
    later taken up and popularized by Arthur Koestler. It was in turn based on James Papez's very
    influential 1937 paper which claimed that the limbic system is primarily concerned with emotion. He
    put forward the "Papez circuit", involving, among other structures, the cingulate cortex and the
    hippocampus.

    Maclean's theory seems to be an up-to-date version of Plato's charioteer (the horses are the
    emotions, the charioteer is the intellect who controls and commands them). It has a certain appeal
    but it doesn't really fit the facts.

    1. Maclean regarded the hippocampus as an "emotional keyboard" (a bit like the Organ of Corti in the
    inner ear, I suppose). However, we now know that the hippocampus is concerned with memory
    formation, not emotion. Damage to the hippocampus and mamillary bodies has little effect on
    emotion but a lot of effect on memory, contrary to what Maclean predicted.

    2. The distinction between phylogenetically "old" cortex and "new" cortex is no longer widely
    accepted and for this reason many anatomists prefer to avoid terms like "archicortex" and
    "neocortex" and speak of "isocortex" and "allocortex" instead. It also now appears that the
    forebrain is actually phylogenetically *older* than the midbrain and hindbrain (Christopher
    Wills, _The Runaway Brain_).

    3. Although the amygdala, which is regarded as part of the "limbic system", is related to emotion
    (especially fear), it is misleading to think of the limbic system as *the* emotional centre of
    the brain. Emotion seems to be a property of the brain as a whole.

    4. Maclean's theory had the merit of setting emotion in an evolutionary context, but it does not
    really accord with the facts as we understand them today.

    References:

    Barr L M, Kiernan J A. (1972) "The Human Nervous System" (6th ed.). JB Lipincott, Philadelphia.

    Gloor P. (1997) "The Temporal Lobe and Limbic System". Oxford University Press, Oxford.

    Ledoux J. (1998). "The Emotional Brain'. Phoenix, London.

    Damasio A R (1994). "Descartes' Error: Emotion, Reason and the Human Brain". Papermac, London and
    Basingstoke.

    Wills C. (1993) "The Runaway Brain: the evolution of human uniqueness". HarperCollins, London.

    AC.

    --
    Using Linux GNU/Debian - Windows-free zone acampbell.org.ukacampbell.org.uk (book reviews and articles)
    Email: replace "www." with "ac@"

  3. "Anthony Campbell" <[email hidden]> wrote in message
    "]news:[email hidden]...

    Quoted message said:


    3. Although the amygdala, which is regarded as part of the "limbic system", is related to emotion
    (especially fear), it is misleading to think of the limbic system as *the* emotional centre of
    the brain.

    It is much more misleading to say as you do, that it is misleading!

    The general principal reason behind anyone wanting to make your 'diluting' point the way you did is
    (put with EPT simplicity) that: When insidious CURSES reinforce our instinctive AEVASIVE
    inclinations we ordinarily tend to put up mental barriers or remove our mental self (and indirectly
    that of others) from the possibility of associations (ultimately to memories of CURSES type) that
    *if allowed to flow and push-through to energize neurally most directly correlated actention
    modules, the result would most likely be disintegrative distress or emotional overload.

    Quoted message said:

    Emotion seems to be a property of the brain as a whole.

    Fully conscious *feelings* even more so.

    P

  4. "Anthony Campbell" <[email hidden]> wrote in message
    "]news:[email hidden]...

    Quoted message said:

    On 2003-12-19, Michael Ragland <[email hidden]> wrote: It also now appears that the forebrain
    is actually phylogenetically *older* than the midbrain and hindbrain (Christopher Wills, _The
    Runaway Brain_).

    I think it only fair to tell people that this is true only as far as that it started-out as a
    processing station in service of olfaction.

    P

  5. Peter F. said:


    "Anthony Campbell" <[email hidden]> wrote in message "]news:[email hidden]...

    Quoted message said:


    3. Although the amygdala, which is regarded as part of the "limbic system", is related to emotion
    (especially fear), it is misleading to think of the limbic system as *the* emotional centre of
    the brain.

    It is much more misleading to say as you do, that it is misleading!

    The general principal reason behind anyone wanting to make your 'diluting' point the way you did
    is (put with EPT simplicity) that: When insidious CURSES reinforce our instinctive AEVASIVE
    inclinations we ordinarily tend to put up mental barriers or remove our mental self (and
    indirectly that of others) from the possibility of associations (ultimately to memories of CURSES
    type) that *if allowed to flow and push-through to energize neurally most directly correlated
    actention modules, the result would most likely be disintegrative distress or emotional overload.

    Quoted message said:

    Emotion seems to be a property of the brain as a whole.

    Fully conscious *feelings* even more so.

    I'm sorry: I have no idea what you mean by this.

    A.

    --
    Using Linux GNU/Debian - Windows-free zone acampbell.org.ukacampbell.org.uk (book reviews and articles)
    Email: replace "www." with "ac@"

  6. On 2003-12-19, Michael Ragland <[email hidden]> wrote: Our Ancient Laughing Brain Courtesy of
    Cerebrum The Laughing Brain Scientists have been studying the play behavior of young animals to
    identify its neural basis. Some are also evaluating whether and to what extent early social
    experiences such as play or lack of it influence neural structure and function. Although the neural
    bases of laughter are not fully understood, we are slowly putting together a picture of the various
    brain regions involved in producing laughter.       The ancient limbic system of the brain,
    including two structures involved with emotions - the hippocampus and the amygdala - is involved in
    laughter, suggesting that it is deep in our animal nture. But our cortex, too, appears to have a
    role, relating laughter and speech and bringing humor into the equation. A. Primitive brain
    (archipallium) - Self preservation, aggression. B. Intermediate brain (paleopallium) - Limbic
    system, emotions. C. Rational brain (neopallium) - Neocortex, intellectual tasks.   [snip] This is
    the "triune brain" hypothesis, first enunciated by Paul Maclean between 1949 and 1970 and later
    taken up and popularized by Arthur Koestler. It was in turn based on James Papez's very influential
    1937 paper which claimed that the limbic system is primarily concerned with emotion. He put forward
    the "Papez circuit", involving, among other structures, the cingulate cortex and the hippocampus.

    Maclean's theory seems to be an up-to-date version of Plato's charioteer (the horses are the
    emotions, the charioteer is the intellect who controls and commands them). It has a certain appeal
    but it doesn't really fit the facts.

    1. Maclean regarded the hippocampus as an "emotional keyboard" (a bit like the Organ of Corti in the
    inner ear, I suppose). However, we now know that the hippocampus is concerned with memory
    formation, not emotion. Damage to the hippocampus and mamillary bodies has little effect on
    emotion but a lot of effect on memory, contrary to what Maclean predicted.

    2. The distinction between phylogenetically "old" cortex and "new" cortex is no longer widely
    accepted and for this reason many anatomists prefer to avoid terms like "archicortex" and
    "neocortex" and speak of "isocortex" and "allocortex" instead. It also now appears that the
    forebrain is actually phylogenetically *older* than the midbrain and hindbrain (Christopher
    Wills, _The Runaway Brain_).

    Response: I think it is ludicrous to suggest the forebrain is phylogenetically older than the
    hindbrain. Only a crackpot would really believe this. I found the following on the internet which I
    think is in accordance with what science knows:

    During embryonic development, the brain first forms as a tube, the anterior end of which enlarges
    into three hollow swellings that form the brain, and the posterior of which develops into the spinal
    cord. Some parts of the brain have changed little during vertebrate evolutionary history.

    Parts of the brain as seen from the middle of the brain. Image from Purves et al., Life: The Science
    of Biology, 4th Edition, by Sinauer Associates (www.sinauer.com) and WH Freeman (www.whfreeman.com),
    used with permission.

    Vertebrate evolutionary trends include Increase in brain size relative to body size. Subdivision
    and increasing specialization of the forebrain, midbrain, and hindbrain. Growth in relative size
    of the forebrain, especially the cerebrum, which is associated with increasingly complex behavior
    in mammals.

    The Brain Stem and Midbrain

    The brain stem is the smallest and from an evolutionary viewpoint, the oldest and most primitive
    part of the brain. The brain stem is continuous with the spinal cord, and is composed of the parts
    of the hindbrain and midbrain. The medulla oblongata and pons control heart rate, constriction of
    blood vessels, digestion and respiration.

    The midbrain consists of connections between the hindbrain and forebrain. Mammals use this part of
    the brain only for eye reflexes.

    The Cerebellum

    The cerebellum is the third part of the hindbrain, but it is not considered part of the brain stem.
    Functions of the cerebellum include fine motor coordination and body movement, posture, and balance.
    This region of the brain is enlarged in birds and controls muscle action needed for flight.

    The Forebrain

    The forebrain consists of the diencephalon and cerebrum. The thalamus and hypothalamus are the parts
    of the diencephalon. The thalamus acts as a switching center for nerve messages. The hypothalamus is
    a major homeostatic center having both nervous and endocrine functions.

    The cerebrum, the largest part of the human brain, is divided into left and right hemispheres
    connected to each other by the corpus callosum. The hemispheres are covered by a thin layer of gray
    matter known as the cerebral cortex, the most recently evolved region of the vertebrate brain. Fish
    have no cerebral cortex, amphibians and reptiles have only rudiments of this area.

    The cortex in each hemisphere of the cerebrum is between 1 and 4 mm thick. Folds divide the cortex
    into four lobes: occipital, temporal, parietal, and frontal. No region of the brain functions alone,
    although major functions of various parts of the lobes have been determined.

    3. Although the amygdala, which is regarded as part of the "limbic system", is related to emotion
    (especially fear), it is misleading to think of the limbic system as *the* emotional centre of
    the brain. Emotion seems to be a property of the brain as a whole.

    Response: The limbic system is important when it comes to emotion. It consists of several parts. I
    would agree emotion seems to be a property of the brain as a whole but I would state there are some
    parts of the brain which are more important in this area than others.

    4. Maclean's theory had the merit of setting emotion in an evolutionary context, but it does not
    really accord with the facts as we understand them today.

    Response: I'm not familiar with Maclean's theory. The article I posted really didn't make reference
    to Maclean's theory. It stated the limbic system was involved in laughter, not totally responsible
    for it. Nor did it state the limbic system was the emotional center of the brain. It did state,
    however, that the limbic system is an ancient part of our brains and it qualified by stating,
    "Although the neural bases of laughter are not fully understood, we are slowly putting together a
    picture of the various brain regions involved in producing laughter." Besides the limbic system it
    mentioned, "But our cortex, too, appears to have a role, relating laughter and speech and bringing
    humor into the equation."

    I saw no evidence in the article I posted being analogous to Maclean's theory of Plato's charioteer
    (the horses are the emotions, the charioteer is the intellect who controls and commands them). It
    has a certain appeal but it doesn't really fit the facts. I thought the article was overall holistic
    in its look at the causes of laughter.

    I posted the article because in the past I've seen people post to sci.bio.evolution asking about the
    evolutionary signifigance of laughter and this was the first article I read which I thought had some
    really insightful points to make about laughter.

    Michael Ragland

  7. On 2003-12-22, Michael Ragland <[email hidden]> wrote:

    [snip]

    Quoted message said:

    4. Maclean's theory had the merit of setting emotion in an evolutionary context, but it does not
    really accord with the facts as we understand them today.

    Response: I'm not familiar with Maclean's theory. The article I posted really didn't make
    reference to Maclean's theory. It stated the limbic system was involved in laughter, not totally
    responsible for it. Nor did it state the limbic system was the emotional center of the brain. It
    did state, however, that the limbic system is an ancient part of our brains and it qualified by
    stating, "Although the neural bases of laughter are not fully understood, we are slowly putting
    together a picture of the various brain regions involved in producing laughter." Besides the
    limbic system it mentioned, "But our cortex, too, appears to have a role, relating laughter and
    speech and bringing humor into the equation."  

    Whether or not the writer of this article is familiar with Maclean, this is where the "triune brain"
    theory originated. My reference to Plato is my own impression; one could also cite Freud's notion of
    the superego and the id, which seems to stem from the same tradition.

    Apart from the fact that the hippocampus, one of the principal components of the "limbic system", is
    concerned with memory and not emotion as Maclean proposed, I think this whole way of looking at how
    brain function has arisen is misleading in a number of ways. It presupposes a very hierarchical view
    of the brain, derived ultimately from the anatomist Hughlings Jackson in the 19C.

    IMO Antonio Damasio is on the right lines in saying that the notion that the intellect sits on top
    of the unruly emotions and keeps them in check ("Descartes' error"😉 is wrong and leads to a mistaken
    understanding of how the brain works. Thinking is a late evolutionary development. Long before there
    were thoughts there were feelings, and we are still predominantly feeling organisms. It is
    impossible to function effectively unless our decision-making is "weighted" by our emotions. Damasio
    cites instances where patients who have lost this element were unable to make quite simple
    decisions.

    A.

    --
    Using Linux GNU/Debian - Windows-free zone acampbell.org.ukacampbell.org.uk (book reviews and articles)
    Email: replace "www." with "ac@"

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