Discussion 6

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Physiology of Behavior

(Twelfth Edition)

Chapter 11

Emotion

11-2

Chapter Review

Fear

Aggression

Impulse Control

Communication of Emotions

Feelings of Emotion

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Learning Objectives (1 of 5)

11.1 Describe the three components of an emotional response.

11.2 Explain what has been learned about the role of the amygdala and ventromedial prefrontal cortex in fear, fear conditioning, and extinction based on results of research with laboratory animals.

11.3 Describe the role of the amygdala and ventromedial prefrontal cortex in emotional conditioning, extinction, and emotional memory in humans.

11-4

Learning Objectives (2 of 5)

11.4 Describe what is known from research with laboratory animals about serotonin and the neural circuitry of aggression and predation.

11.5 Evaluate the roles of heredity and serotonin in human aggression.

11.6 Summarize the role of hormonal control of aggression, citing evidence from studies involving humans and other animals.

11.7 Cite examples supporting the role of the ventromedial prefrontal cortex in impulse control.

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Learning Objectives (3 of 5)

11.8 Provide evidence for a developmental factor in impulse control.

11.9 Compare brain differences involved in impulse control between those convicted of a crime or diagnosed with antisocial personality disorder and typically developing adults.

11.10 Explain the role of serotonin in impulse control regulation.

11.11 Describe the brain regions involved in emotional aspects of moral decision making and cite evidence from the research literature.

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Learning Objectives (4 of 5)

11.11 Describe the brain regions involved in emotional aspects of moral decision making and cite evidence from the research literature.

11.12 Describe evidence in support of emotional expressions as innate responses.

11.13 Summarize the brain structures involved in emotional recognition, by addressing laterality, direction of gaze, imitation, and disgust.

11.14 Review the brain structures involved in emotional expression, by addressing laterality, laughter, and humor.

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Learning Objectives (5 of 5)

11.15 Summarize the evidence for and against the James-Lange theory of emotion.

11.16 Explain the role of feedback from emotional expressions in mood and activity of the autonomic nervous system.

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Emotion (1 of 2)

There is no such thing as a neutral emotion.

Emotion

Positive or negative react ions to particular situations

Patterns of physiological changes and accompanying behaviors—or at least urges to perform test behaviors

The word emotion refers to positive or negative react ions to particular situations. There is no such thing as a neutral emotion.

Emotions consist of patterns of physiological changes and accompanying behaviors—or at least urges to perform these behaviors.

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Emotion (2 of 2)

Hormones

Secreted by the adrenal medulla

Increase blood flow to muscles

Cause nutrients stored in and made available to muscles to be converted into glucose

Hormonal responses reinforce the autonomic responses.

The hormones secreted by the adrenal medulla—epinephrine and norepinephrine—further increase blood flow to the muscles and cause nutrients stored in the muscles to be converted into glucose.

In addition, the adrenal cortex secretes steroid hormones, which also help to make glucose available to the muscles.

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Fear

Amygdala

Lateral Nucleus (LA)

Central Nucleus (CE)

Lateral nucleus (LA)

Nucleus of amygdala that receives sensory information from neocortex, thalamus, and hippocampus and sends projections to basal, accessory basal, and central nucleus of amygdala

Central nucleus (CE)

Region of amygdala that receives information from basal, lateral, and accessory basal nuclei and sends projections to wide variety of regions in brain; involved in emotional responses

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Figure 11.2 Amygdala Projections

This figure shows a much-simplified diagram of the major divisions and connections of the amygdala that play a role in emotions.

Figure 11.2 page 334

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Figure 11.3 The Outputs of the Central Nucleus of the Amygdala

Shown here we some important brain regions that receive input from the central nucleus of the amygdala and the emotional responses controlled by these regions.

Figure 11.3 page 334

(Adapted from Davis. M.. Trends in Phemwoologicel Sciences, 1992, 13, 35-41.)

11-13

Emotions as Response Patterns Research with Laboratory Animals

Fear

Conditioned emotional response

Classically conditioned fear response

Conditioned emotional response

Classically conditioned response that occurs when neutral stimulus is followed by aversive stimulus; usually includes autonomic, behavioral, and endocrine components such as changes in heart rate, freezing, and secretion of stress-related hormones

Ventromedial prefrontal cortex (vmPFC)

Region of prefrontal cortex at base of anterior frontal lobes, adjacent to midline

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Figure 11.4 Context Conditioning: An example of a conditioned emotional response

In this example, a rat experiences a shock in an enclosure with a blue floor. In the signaled condition, the shock is paired with the sound of a bell. The animal will now experience fear in the enclosure with the blue floor, demonstrated by freezing behavior. The animal is then placed in an enclosure with a purple floor. In the unsignaled condition, the rat does not experience fear, as demonstrated by standing up and engaging in exploratory behavior. In the signaled condition, the rat will experience fear and freeze when it hears the sound of the bell, even in this new purple-floor environment. The bell has become a conditioned stimulus, even in this new environment.

Figure 11.4 page 335

The most basic form of emotional learning is a conditioned emotional response, which is produced by a neutral stimulus that has been paired with an emotion-producing stimulus

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Emotions as Response Patterns

Amygdala is involved in human emotional responses

Lesions of amygdala decrease emotional responses

Lesions impair conditioned emotional responding

Lesions interfere with effects of emotions on memory

Medial prefrontal cortex is involved in extinction of conditioned emotional responses

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Fear Research with Humans (1 of 2)

Phelps et al. (2004)

Directly established conditioned emotional response in human subjects

Paired appearance of visual stimulus with electric shocks to wrist

Extinguished response by presenting squares alone, without any shocks

Concluded

Increased activity of medial prefrontal cortex correlated with extinction of conditioned response

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Fear Research with Humans (2 of 2)

Ventromedial Prefrontal Cortex (vmPFC)

Includes medial orbitofrontal cortex and subgenual anterior cingulate cortex

Plays a role in complex analyses of social situations

Many investigators believe that impulsive violence is a consequence of faulty emotional regulation.

For most of us, frustrations may elicit an urge to respond emotionally, but we usually manage to calm ourselves and suppress these urges.

As we shall see, the ventromedial prefrontal cortex plays an important role in regulating our responses to such situations.

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Figure 11.5 Extinction

In this example, rats experience fear conditioning, pairing the sound of a bell with a shock. One half of the rats experience extinction (repeatedly hearing the bell without the shock) in new enclosures with either purple or green floors, as demonstrated by standing up and engaging in exploratory behavior. When the groups of animals are tested after extinction, the animals demonstrate extinction in the same enclosure where they learned not the respond to the bell (extinction). Extinction learning is context dependent. In this example, when extinction occurs in one environment, or context (green floor), it does not generalize to a new environment (purple floor) where the rat continues to experience fear and engage in freezing behavior.

Figure 11.5 page 336

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Figure 11.6 The Ventromedial Prefrontal Cortex

Table 11.1 and Figure 11.6 page 337

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Table 11.1 Brain Regions Involved in Fear Response

Nucleus	Function
Central Nucleus of the Amygdala	Activation produces fear-related behaviors; lesioning prevents production of fear-related behaviors
Lateral Nucleus of the Amygdala	Involved in producing conditioned emotional response
Ventromedial Prefrontal Cortex (vmPFC)	Involved in extinction of conditioned emotional response

Table 11.1 and Figure 11.6 page 337

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Role of Ventromedial Prefrontal Cortex

Amygdala

Plays important role in provoking anger and violent emotional reactions

Matures early in development

Prefrontal cortex

Matures much later, during late childhood and early adulthood

Plays important role in suppressing such behavior by making people see its negative consequences

11-22

Aggression

Species typical behavior, many related to reproduction

Aggressive Behavior

Threat Behavior

Defensive Behavior

Submissive Behavior

Predation

Threat behavior

Stereotypical species-typical behavior that warns another animal that it may be attacked if it does not flee or show a submissive behavior

Defensive behavior

Species-typical behavior by which animal defends itself against threat of another animal

Submissive behavior

Stereotyped behavior shown by animal in response to threat behavior by another animal; serves to prevent an attack

Predation: member of one species attacking members of another species, usually for food

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Figure 11.7 Neural Circuitry in Defensive Behavior and Predation

The diagram shows interconnections of parts of the amygdala, hypothalamus, and periaqueductal gray matter (PAG) and their effects on defensive behaviors and predation in cats, based on the studies by Shaikh, Siegel, and their colleagues. Black arrows indicate excitation; red arrows indicate inhibition.

Figure 11.7 page 339

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Role of Serotonin

Activity of serotonergic synapses inhibits aggression

Destruction of serotonergic axons in forebrain facilitates aggressive attack

Some serotonin agonist, decreased irritability and aggressiveness, as measured by a psychological test

An overwhelming amount of evidence suggests that the activity of serotonergic synapses inhibits aggression.

In contrast, destruction of serotonergic axons in the forebrain facilitates aggressive attack, presumably by removing an inhibitory effect (Vergnes et al., 1988).

Several studies have found that serotonergic neurons play an inhibitory role in human aggression

If low levels of serotonin release contribute to aggression, perhaps drugs that act as serotonin agonists might help to reduce antisocial behavior.

In fact, a study by Coccaro and Kavoussi (1997) found that fluoxetine (Prozac), a serotonin agonist, decreased irritability and aggressiveness, as measured by a psychological test..

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Figure 11.8 Serotonin and Risk-Taking Behavior

The graph shows the percentage of young male monkeys alive or dead as a function of 5-HIAA level in the cerebrospinal fluid, measured four years previously.

Figure 11.8 page 340

(Based on data from Higley et al., 1996.)

11-26

Hormonal Control of Aggressive Behavior

Aggression in Males

In rodents, androgen secretion occurs prenatally, decreases, and increases again at puberty

Intermale aggressiveness increases at puberty

Aggression in Females

Females less aggressive than males

Aggression appears to be facilitated by testosterone

Aggression in Males

Adult males of many species fight for territory or access to females. In laboratory rodents, androgen secretion occurs prenatally, decreases, and then increases again at the time of puberty.

Intermale aggressiveness also begins around the time of puberty, which suggests that the behavior is controlled by neural circuits that are stimulated by androgens.

Indeed, many years ago, Beeman (1947) found that castration reduced aggressiveness and that injections of testosterone reinstated it.

Aggression in Females

Androgens have an organizational effect on the aggressiveness of females, and a certain amount of prenatal androgenization appears to occur naturally.

Most rodent fetuses share their mother’s uterus with brothers and sisters, arranged in a row like peas in a pod.

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Figure 11.9 Serotonin Activity and Aggression (a)

(a) Some studies have found that serotonergic neurons play an inhibitory role in human aggression.

Figure 11.9a page 342 (continues on next slide)

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Figure 11.9 Serotonin Activity and Aggression (b)

(b) Selective serotonin reuptake inhibitors (SSRIs) block the serotonin transporter (shown in green on the presynaptic cell), preventing serotonin reuptake by the presynaptic cell. Use of SSRIs results in increased serotonin available in the synapse to bind to receptors and is associated with reduced aggressive behavior.

Figure 11.9b page 342 (continues from previous slide)

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Figure 11.10 Effects of 14 Days of Estradiol and Testosterone Administration on Interfemale Aggression in Rats

Figure 11.10 and 11.12, page 343

(Based on data from van de Poll et al., 1988.)

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Figure 11.12 Organizational and Activational Effects of Testosterone on Social Aggression

Figure 11.10 and 11.12, page 343

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Figure 11.11 0M, 1M, and 2M Female Mouse Fetuses

Figure 11.11 page 343

(Adapted from vom Saal, F. S., in Hormones and Aggressive Behavior, edited by

B. B. Svare. New York: Plenum Press, 1983.)

11-32

Effects of Androgens on Human Aggressive Behavior

Prenatal Androgenization

Increases aggressive behavior in all species that have been studied, including primates

Therefore, if androgens did not affect aggressive behavior in humans, our species would be exceptional.

After puberty, androgens also begin to have activational effects

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Figure 11.13 Alcohol, Mating, and Aggressive Behavior in Monkeys

The graph shows the effect of alcohol intake on frequency of aggressive behavior of dominant and subordinate male squirrel monkeys during the mating season and the nonmating season.

Figure 11.13 page 345

(Based on data from Winslow and Miczek, 1988.)

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Impulse Control

Impulsive violence as consequence of faulty emotion regulation

Ventromedial prefrontal cortex

Medial orbitofrontal cortex

Subgenual anterior cingulate cortex

11-35

Figure 11.14 Phineas Gage’s Accident. The steel rod entered his left cheek and exited through the top of his head.

Figure 11.14 page 346

(From: Everett Collection Historical / Alamy)

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Figure 11.15 Role of the vmPFC

The participants’ task was to reach maximal proximity to either a live snake or a toy bear, by repeatedly choosing whether to bring the object closer or move it away, while undergoing fMRI brain scanning. A display of courage was accompanied by activation of a region of the vmPFC, the subgenual anterior cingulate cortex.

Figure 11.15 page 348

(Nili, U., Goldberg, H., Weizman, A., and Dudai, Y., Fear thou not: Activity of frontal and temporal circuits in moments of real-life courage, Neuron, 2010, 66(6), 949–962.)

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Brain Development and Impulse Control

Brain Development

Role of the amygdala (matures early)

Role of prefrontal cortex (matures late)

Crime

Raine et al., (1998)

Serotonin

Development

Amygdala: provoking anger and violent behavior

PFC: suppressing such behavior

See Figure 11.16 (next slide)

Crime: Raine et al., 1998 found evidence of decreased PFC activity and increased subcortical activity (including amygdala) in convicted murderers

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Figure 11.16 Development of Amygdala Over Lifespan

Figure 11.16 page 348

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Moral Decision Making

Emotional reactions guide:

Moral judgments

Decisions involving personal risk and reward

Prefrontal cortex

Thomson, 1986 (next slide)

vmPFC

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Figure 11.17 Moral Decision-Making Scenario: The Case of the Trolley

In scenario (a) a person must decide to throw a switch to save five people or one person. In scenario (b) a person must decide to push a man off a bridge to save five people.

Figure 11.17 page 350

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Table 11.2 Examples of Scenarios Involving nonmoral, Impersonal Moral, and Personal Moral Judgments from the Study by Koenigs et al. (2007)

Brownies (Nonmoral scenario) You have decided to make a batch a brownies for yourself. You open your recipe book and find a recipe for brownies. The recipe calls for a cup of chopped walnuts. You don’t like walnuts, but you do like macadamia nuts. As it happens, you have both kinds of nuts available to you. Would you substitute macadamia nuts for walnuts in order to avoid eating walnuts?

Speedboat (Impersonal moral scenario) While on vacation on a remote island, you are fisting from a seaside dock. You observe a group of tourists board a small boat and set sail for a nearby island. Soon after their departure you hear over the radio that there is a violent storm brewing, a storm that's sure to intercept them. The any way that you can ensure their safety is to warn them by borrowing a nearby speedboat. The speedboat belongs to a miserly tycoon who would not take kindly to you borrowing his properly Would you borrow the speedboat in order to warm the tourists about the storm?

Lifeboat (Personal moral scenario) You are on a cruse ship when there is a fire on board, and the ship has to be abandoned. The lifeboats are carrying many more people than they were de-signed to carry. The lifeboat you are in is sitting dangerously low in the water —a few inches lower, and i will sink. The seas start to get rough, and the boat begins to fill with water. If nothing is done it will sink before the recue boats arrive, and everyone on board will die. However, there is an injured person who will not survive in any case. If you throw that person overboard the boat will stay afloat and the remaining passengers will be saved. Would you throw this person overboard in order to save the lives of the remaining passengers?

Table 11.2 page 350

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Communication of Emotions

Facial Expression of Emotions: Innate Responses

Facial expression of emotions appear to be innate

Cross-cultural studies and studies of blind children support belief that facial expressions of emotion are innate

Many species of animals (including our own) communicate their emotions to others by means of postural changes, facial expressions, and nonverbal sounds (such as sighs, moans, and growls)

These expressions serve useful social functions: They tell other individuals how we feel and—more to the point—what we are likely to do

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Figure 11.18 Facial Expressions in a New Guinea Tribesman

The tribesman made these expressions when he heard the following stories: (a) “Your friend has come and you are happy.” (b) “Your child had died.” (c) “You are angry and about to fight.” (d) “You see a dead pig that has been lying there a long time.”

Figure 11.18 page 352

(From Ekman, P., The Face of Man: Expressions of Universal Emotions in a New Guinea Village, New York: Garland STPM Press, 1980. Reprinted with permission.)

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Figure 11.19 Perception of Emotions

The PET scans indicate brain regions activated by listening to emotions expressed by meanings of words (red) or tone of voice (green). (Tracings of brain activity from George et al., 1996.)

Figure 11.19 page 354

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Figure 11.20 Unconscious Imitation of Facial Expressions of Emotions

When patients with unilateral damage to the visual cortex saw photographs of happy or fearful faces, they smiled or frowned when the photographs were presented to their sighted or blind field, which indicates that visual information concerning emotional expressions can take place without conscious awareness.

Figure 11.20 page 354

(Based on data from Tamietto et al., 2009.)

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Communication of Emotions (1 of 2)

Neural Basis of the Communication of Emotions: Recognition

Laterality

Role of the Amygdala and Prefrontal Cortex

Affective Blindsight

As we saw in the previous section, the amygdala plays a special role in emotional responses. It plays a role in emotional recognition as well.

For example, several studies have found that lesions of the amygdala (the result of degenerative diseases or surgery for severe seizure disorders) impair people’s ability to recognize facial expressions of emotion, especially expressions of fear (Adolphs et al., 1994, 1995; Young et al., 1995; Calder et al., 1996; Adolphs et al., 1999).

In fact, some people with blindness caused by damage to the visual cortex can recognize facial expressions of emotion even though they have no conscious awareness of looking at a person’s face, a phenomenon known as affective blindsight (de Gelder et al., 1999, Anders et al., 2004)

Affective blindsight The ability of a person who cannot see objects in his or her blind field to accurately identify facial expressions of emotion while remaining unconscious of perceiving them; caused by damage to the visual cortex

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Communication of Emotions (2 of 2)

Role of the Amygdala and Prefrontal Cortex

As Figure 11.22 shows, high spatial frequencies show fine details of transitions between light and dark, and low spatial frequencies show fuzzy images

These photos primarily stimulate the parvocellular and magnocellular systems, respectively

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Figure 11.22 Involvement of Magnocellular and Parvocellular Systems in Emotional Perception

The figure shows the stimuli used by Vuilleumier et al., (2003). The more primitive magnocellular system is sensitive to low spatial frequencies (SF), and the more recently evolved parvocellular system is sensitive to high spatial frequencies.

Figure 11.22, page 355

(From Vuilleumier, P., Armony, J. L., Driver, J., and Dolan, R. J., Distinct spatial frequency sensitivities for processing faces and emotional expressions, Nature

Neuroscience, 2003, 6, 624–631.)

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Figure 11.23 Eye Fixations After Amygdala Damage

The figure shows the numbers of fixations on a person’s face made by a patient with bilateral amygdala damage compared to a control participant. Warmer colors indicate increasing numbers of fixations.

Note that the patient does not look at the other person’s eyes.

Figure 11.23, page 356

Neuroscience.)

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Neural Basis of Communication of Emotions: Recognition

Perception of Direction of Gaze

Recognition of Disgust

Perception of Direction of Gaze

Recognition of direction of another monkey’s gaze involves neurons in superior temporal sulcus

Important to know target of another’s gaze

Recognition of Disgust

Ability to recognize facial expressions of disgust is impaired by damage to insular cortex and basal ganglia

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Figure 11.25 A Gaze Direction Cell

The graph shows the responses of a single neuron in the cortex lining in the superior temporal sulcus of a monkey’s brain. The cell fired most vigorously when the monkey was presented a photograph of a face looking upward.

Figure 11.25 page 357

(Based on Perrett, D. I., Harries, M. H., Mistlin, A. J., et al. International Journal of Comparative Psychology, 1992, 4, 25–55.)

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Role of Imitation in Recognition of Emotional Expressions: The Mirror Neuron System

Adolphs et al. (2000)

Discovered possible link between somatosensation and emotional recognition

Correlated brain damage information with patients’ ability to recognize and identify facial expressions of emotions

Found facial expression of an emotion, we unconsciously imagine ourselves making that expression

They compiled computerized information about the locations of brain damage in 108 patients with localized brain lesions and correlated this information with the patients’ ability to recognize and identify facial expressions of emotions

They found that the most severe damage to this ability was caused by damage to the somatosensory cortex of the right hemisphere. (See Figure 11.26.)

They suggest that when we see a facial expression of an emotion, we unconsciously imagine ourselves making that expression

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Figure 11.26 Brain Damage and Recognition of Facial Expressions of Emotion

This computer-generated image shows performance of participants with localized brain damage on recognition of facial expressions of emotion. The colored areas outline the site of the lesions. Lesions that resulted in good performance are shown in shades of blue; those that resulted in poor performance are shown in red and yellow.

Figure 11.26 page 357

(From Adolphs, R., Damasio, H., Tranel, D., Cooper, G., and Damasio, A. R. The Journal of Neuroscience, 2000, 20, 2683–2690.)

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Figure 11.27 Disease and Disgust (1 of 2)

The figure shows pairs of photographs with high and low relation to the threat of disease used in the online survey presented on the BBC Science web site. The numbers in red or green indicate the mean ratings (range = 1–5) made by people who completed the survey.

Figure 11.27 page 359

(From Curtis, V., Aunger, R., and Rabie, T., Evidence that disgust evolved to protect from risk of disease, Biology Letters, 2004, 271, S131–S133.)

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Neural Basis of the Communication of Emotions: Expression

Volitional facial paresis

Emotional facial paresis

Volitional facial paresis

Difficulty in moving the facial muscles voluntarily; caused by damage to the face region of the primary motor cortex or its subcortical connections

Emotional facial paresis

Lack of movement of facial muscles in response to emotions in people who have no difficulty moving these muscles voluntarily; caused by damage to the insular prefrontal cortex, subcortical white matter of the frontal lobe, or parts of thalamus

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Emotions as Response Patterns (1 of 2)

Neural Basis of Communication of Emotions: Expression

Facial expressions are automatic and involuntary

It is difficult to artificially produce realistic facial expressions of emotion

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Emotions as Response Patterns (2 of 2)

Neural Basis of Communication of Emotions: Expression

Right hemisphere plays more significant role for expressing emotions

When people show emotions with their facial muscles, left side of face usually makes more intense expression

Right hemisphere plays more significant role in recognizing emotions in voice or facial expressions—especially negative emotions

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Figure 11.29 Humor and Volition of Social Norms

The graph shows the activation of the right ventromedial prefrontal cortex and the left orbitofrontal cortex, as measured by fMRI, by exposure to humorous cartoons with increasing funniness and increasing violation of social norms.

Figure 11.29 page 360

(Based on data from Goel and Dolan, 2007.)

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The James-Lange Theory

James-Lange Theory

William James and Carl independently suggested similar explanations for emotion

Theory suggests that behaviors and physiological responses are directly elicited by situations

Feelings of emotions are produced by feedback from these behaviors and responses

William James (1842–1910), an American psychologist, and Carl Lange (1834–1900), a Danish physiologist, independently suggested similar explanations for emotion, which most people refer to collectively as the James-Lange theory (James, 1884; Lange, 1887)

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Figure 11.30 The James-Lange Theory of Emotion

An event in the environment triggers behavioral, autonomic, and endocrine responses. Feedback from these responses produces feelings of emotions.

Figure 11.30 page 362

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Figure 11.31 Imitation in Infants

The photographs show happy, sad, and surprised faces posed by an adult and the responses made by an infant.

Discussion 6

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