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+ Stimulating Neural Activity Numerous techniques exist for activating neurons in the brain. Electrode stimulation Chemical activation through cannulae Photostimulation: stimulation with light + “Two light-sensitive proteins from unicellular organisms have been harnessed to rapidly activate or silence neurons. This optical remote control allows precise, millisecond control of neural circuits.” ChR2 Ion Channels – photosensitive proteins channel which depolarize the membrane when blue light is presented. NpHR Ion Transporter – photosensitive protein channel which hyperpolarizes the membrane when yellow light is presented. + Genetic Methods Classical and conditional knockouts (KO). Classical KOs - the function of the gene is abolished from a very early stage of development. Conditional KOs - there is either a temporal restriction (gene function is abolished at certain premeditated time windows) or a regional restriction (no gene function in certain brain regions) Transgenics Foreign gene, e.g. human APP, is inserted into the genome Knockins Specific mutations are introduced in the gene leading to a loss of activity of the proteins encoded by the targeted gene (although the gene expression per se is not voided as it is in KOs). Classical or conditional + Assessment of Species -Common Behaviors Assessment of behaviors displayed by all members of a species Open-field test – general activity Colony-intruder paradigm – aggression and defensive behavior Elevated plus maze – anxiety Social interaction Tests of sexual behavior + Traditional Conditioning Paradigms - Learning Pavlovian conditioning Pairing an unconditioned stimulus with a conditioned stimulus Pavlov’s dogs Copyright © 2006 by Allyn and Bacon + Fear Conditioning Protocol TRAINING Day 1 Novel Context (CS) + Tone (CS) + Footshock(US) CONTEXT TEST CUE TEST Day 2 Training Context (CS) Tone (CS) + Traditional Conditioning Paradigms - Learning Operant conditioning Reinforcement and punishment Self-stimulation Animal works for electrical stimulation + Seminatural Learning Paradigms Mimic situations that an animal might encounter in its natural environment Conditioned taste aversion Pairing something that makes an animal ill (emetic) with a taste Radial tests arm maze spatial abilities Copyright © 2006 by Allyn and Bacon + Seminatural Learning Paradigms Morris Rat water maze – tests spatial abilities must find hidden platform in an opaque pool Conditioned defensive burying – following a single aversive stimulus delivered from an object, rats will spray bedding at the object Antianxiety drugs decrease the amount of burying behavior + Mind and Brain Emotion Chapter 11 + Chapter Overview Emotions as Response Patterns Fear Anger, Aggression, and Impulse Control Neural control of aggressive behavior Role of 5-HT Role of vmPFC Hormonal control of aggressive behavior Communication of Emotions Feelings of Emotions + Emotions as Response Patterns An emotional response consists of 3 types of components: Behavioral Autonomic i.e., dog defending its territory might bark, growl attack Mobilization of energy; activity of sympathetic branch of ANS increase while parasympathetic activity decreases Hormonal Released from the adrenal medulla Epinephrine and NE further increase blood flow to the muscles Steroid hormones Cause nutrients stored in the muscles to be converted to glucose + Amgydala Small, almond-shaped structure in the medial temporal lobe Adjacent to hippocampus + Figure 11.1 The Amygdala Ventral striatum Dorsomedial nucleus of thalamus (projects to prefrontal cortex) Ventromedial Prefrontal cortex + Emotions as Response Patterns Fear Amygdala Lateral Nucleus (LA) – receives sensory information from neocortex, thalamus, and hippocampus and projects to basal, accessory basal, and central nucleus of the amygdala. Central Nucleus (CN) – receives information from the basal, lateral, and accessory basal nuclei and projects to many brain regions involved in emotional processes. + CN Single most important part of the brain for the expression of emotional responses provoked by aversive stimuli Threatening stimuli increase neural activity and fos expression Damage to CN reduces or abolishes a wide range of emotional behaviors and physiological responses Animals no longer show signs of fear Act more tamely when handled Stress hormones are lower Less likely to develop ulcers or other forms of stress-induced illnesses Opposite is true with stimulation of CN + Emotions as Response Patterns Some stimuli automatically activate the CN and produce fear responses (loud noises)…but learning which stimuli are dangerous is also very important The most basic form of emotional learning is CER Figure 11.3 Conditioned Responses ear Conditioned Emotional Response – classically conditioned fear response. + Classical Conditioning Physical changes responsible for CC occur in LA Neurons in LA project to CN, projects to hypothalamus, midbrain, pons and medulla Responsible for behavioral, autonomic, and hormonal components of conditioned emotional response + Extinction Repeated presentation of the CS alone (without the aversive stimuli), then the CR eventually disappears Extinction is not the same as forgetting, new learning Animal learns that the CS is no longer followed by an aversive stimulus Expression of CR is inhibited (memory for the association b/w CS and aversive stimuli is not erased) Inhibition is supplied by the medial prefrontal cortex + Research with Humans Amygdala is involved in human emotional responses. Lesions of the amygdala decrease emotional responses. Lesions interfere with effects of emotions on memory. + Lesions of the amygdala decrease emotional responses. Bechara et al., (1995) and LaBar et al., (1995) found that people with lesions of the amygdala showed impaired acquisition of a conditioned emotional response (similar to rats) Angrilli et al., (1996) found that the startle response of a man with right amygdala damage was not augmented by an unpleasant emotion In both cases the amygdala plays a role in the expression of the fear response + Medial prefrontal cortex is involved in extinction of conditioned emotional responses in humans. + Lesions interfere with effects of emotions on memory. When people encounter events that produce a strong emotional response, they are more likely to remember that event. Cahill et al., (1995) studied a patient with bilateral amygdala degeneration (patient SM) + Lesions interfere with effects of emotions on memory. Cahill et al., 1995 Told a story about a boy walking with mother on his way to visit his father at work Showed a series of slides during story During one part of the story, boy was injured in a traffic accident, and gruesome slides showed his injuries Normal subjects – remember more details from the emotionladen part of the story Patient SM – no increase in memory + Lesions interfere with effects of emotions on memory. fMRI studies confirm lesion data Cahill et al., 1996 Ss (subjects) watch both neutral and emotionally arousing films (scenes of violent crime), later asked to recall the films fMRI showed increased activity of the right amygdala when the subjects recalled the emotionally arousing films but not when they recalled the neutral ones Ss were most likely to recall the emotionally arousing films that produced the highest level of activity in the right amygdala when they were originally viewed Isenberg et al., (1999) Seeing words that denote threatening situations increases the activity of the amygdala Activation in human amygdala…read words, look at pictures Ratings of emotional intensity of facial expressions by controls and patient S.M. + Anger, Aggression, and Impulse Control Aggressive behaviors are species-typical, usually related to reproductive behavior (defending territory) or self-defense Threat behaviors are more common than actual attack Threat Behavior – stereotypical species-typical behavior warning another animal that it may be attacked; postures or gestures. Defensive Behavior – species-typical behavior an animal uses to defend itself against threat of another animal. Submissive Behavior – stereotyped behavior shown by an animal in response to threat by another animal. Predation – attack of a member of another species; does not result in same level of arousal Predator not angry with its prey…it’s simply food and must be killed + Figure 11.6 Neural Circuitry in Defensive Behavior Gregg and Siegel, 2001 •Series of studies using cats showed that stimulation of the PAG elicited attack and predation •Hypothalamus and amygdala can influence these behaviors through connections with the PAG + Anger, Aggression, and Impulse Control Activity of serotonergic (5-HT) synapses inhibits aggression. Destruction of serotonergic axons in the forebrain facilitates aggressive attack. Howell et al., 2007 5-HT activity in monkeys (examining 5HIAA in CSF) High levels of 5-HIAA in CSF – increased 5-HT activity Young male monkeys with the lowest levels of 5-HIAA showed a pattern of risk-taking behavior, including inappropriate aggression 46% of monkeys with low levels of 5-HIAA died (killed by other monkeys) Selective breeding of rats and foxes – tame animals (increased levels of 5-HT and 5-HIAA + Anger, Aggression, and Impulse Control 5-HT also play an inhibitory role in human aggression Decreased 5-HIAA in CSF is associated with aggression and other forms of antisocial behavior Fluoxetine (Prozac) is a serotonin agonist and decreases irritability and aggressiveness People with at least 1 short allele for the 5-HT transporter have higher anxiety and depression Right amygdala of people carrying the short form of the 5-HT transporter gene showed a higher rate of activity during task (looking at faces expressing fear or anger) + Anger, Aggression, and Impulse Control Impulsive violence may be consequence of faulty emotional regulation…in frustrating situations we can usually calm ourselves down…probably due to the vmPFC Ventromedial Prefrontal Cortex (vmPFC) Includes medial orbitofrontal cortex and subgenual anterior cingulate cortex. + Figure 11.9 The Location of the Ventromedial Prefrontal Cortex + Anger, Aggression, and Impulse Control vmPFC Plays a role in complex analyses of social situations. Serves as interface between brain mechanisms involved in automatic emotional responses and those involved in the control of complex behaviors Includes using our emotional reactions to guide our behavior and controlling the occurrence of emotional reactions in various social situations Moral judgments/dilemmas, decision making + Phineas Gage + Anger, Aggression, and Impulse Control Hormonal Control of Aggressive Behavior Aggression in Males In rodents, androgen secretion occurs prenatally, decreases, and increases again at puberty. Inter-male aggressiveness increases at puberty. Organizational effects – influence development of an animal’s sex organs and brain Effects are permanent Activational effects - occur later in life, after the sex organs have developed. ie. Hormones activate the production of sperm + Figure 11.13 Organizational and Activational Effects of Testosterone on Social Aggression Early exposure to androgens has an organizational effect that stimulates the development of testosterone-sensitive neural circuits that facilitate male aggression + Anger, Aggression, and Impulse Control Effects of androgens on male aggression are mediated by Medial Preoptic Area Implanting testosterone in the MPA reinstated intermale aggression in castrated male rats. + Medial Preoptic Area See Figure 10.18 See Figure 3.21 + Anger, Aggression, and Impulse Control Males attack other males, but rarely attack females Discrimination between sexes based on pheromones Intermale aggression was abolished in mice by cutting the vomeronasal nerve (input from vomeronasal organ) + Anger, Aggression, and Impulse Control Hormonal Control of Aggressive Behavior Aggression in Females Less aggressive than males. Aggression appears to be facilitated by testosterone. Most rodent fetuses share their mom’s uterus with brothers and sisters – peas in a pod A female mouse may have 0,1 or 2 brothers adjacent to her Being next to a male increases blood levels of androgens prenatally Females located between 2 males had more testosterone in their blood and, when tested as adults, showed increased aggression + Anger, Aggression, and Impulse Control Females of some primate species are more likely to engage in fights around the time of ovulation Mostly with males – likely due to increased proximity to males Increased fighting before menstruation Females tend to attack other females + Anger, Aggression, and Impulse Control - human Boys are generally more aggressive than girls Small, but significant increases in aggressiveness in female twins that shared a uterus with a male, versus another female Girls with CAH - exposed abnormally high levels of androgens during prenatal development Castrated male (heterosexual and homosexual) criminals tend to show less aggression (and sex drive) Show increased aggression Lack controls Athletes that take steroids (including testosterone) tend to be aggressive Difficult to prove – may be that more aggressive people take steroids + Chapter Overview Emotions as Response Patterns Communication of Emotions Feelings of Emotions + Basic Emotions Finite set of universal, basic emotions Darwin (evolved) Universality of facial expressions + Emotions & Facial Expressions Ekman et al., (1960s) Analyzing hundreds of films and photographs of people experiencing real emotions Complied an atlas of facial expressions Dr. Cal Lightman Emotions and Facial Expression Six primary emotions Surprise Anger Sadness Disgust Fear Happiness Naturally occurring expressions are usually variations or combinations of basic ones + Additional Facial Expressions Amusement Contempt Contentment Embarrassment Excitement Guilt Pride Relief in achievement + Universality of Facial Expression Several studies People of different cultures make similar facial expression in similar situations People can correctly identify the emotional significance of facial expressions displayed by people from different cultures + Isolated New Guinea tribe Ekman (1971)devised a list of basic emotions to test tribesmen of Papua New Guinea. He observed that members of an isolated culture could reliably identify the expressions of emotion in photographs of people from unfamiliar cultures They could also ascribe facial expressions to descriptions of situations. Ekman concluded that the expressions associated with some emotions were basic or biologically universal to all humans + Communication of Emotions Facial Expression of Emotions: Innate Responses Young blind children show similar facial expressions as normal sighted children. + Neural Basis of the Communication of Emotions: Recognition Laterality of Emotional Recognition Right hemisphere is more important for the comprehension of emotion. Bowers et al., 1991 found that patients with right hemisphere damage had difficulty producing or describing mental images of facial expressions of emotions George et al., 1996 had Ss listen to some sentences and identify their emotional content. Comprehension of emotion from word meaning increased the activity of the PFC bilaterally, the left more than the right. Comprehension of emotion from tone of voice increased the activity of only the right PFC. + Role of the Amygdala in Recognition Important for emotion recognition, especially for facial expressions (of fear). Amygdala lesions impair ability to recognize fear expression FMRI studies show large increases in amygdala activity when people view photographs of faces expression fear Affective Blindsight – ability of a person who cannot see objects in his/her blind field to accurately identify facial expressions of emotion without conscious perception of them. + Amygdala and Fearful Facial Expressions Adolphs et al, (2005) Computer software that exposed only parts of either a fearful or happy facial expression to determine what regions of the face the subjects relied on to discriminate between expressions Results Control subjects consistently relied on eyes to make decisions about expression S.M. – did not derive information from the eyes She did not even look at the eyes of any face, regardless of emotion + Facial Expressions and the Amygdala Why the specific problem with fearful expressions? Most expressions contain other cues that can be used for identification Happiness – smile Fear – increase in size of the white region (sclera) of the eyes + Facial Expression and the Amygdala Amygdala appears to be an integral part of a system that automatically directs visual attention to the eyes when encountering any facial expressions + Communication of Emotions Perception of Direction of Gaze Important to know if another’s gaze is directed toward you or not. Recognition of the direction of another monkey’s gaze involves neurons in the superior temporal sulcus (Figure 11.22). fMRI study confirmed monkey data Pelphrey et al., 2003 had people watch an animated cartoon of a face. When the direction of gaze changed, increased activity was seen in the right STS an + Communication of Emotions Role of Imitation in Recognition of Emotional Expressions Mirror neurons – neurons located in the ventral premotor cortex and inferior parietal lobule that respond when the individual makes a particular movement or sees another individual making that movement Mirror neuron system is activated when we observe facial movements in others and may provide feedback important for empathy + Disgust Anterior insula is essential to both detection and experienced of disgust Imaging data Patient with anterior insula damage Same area of the anterior insula was activated both when Ss viewed expressions of disgust in others and when they smelled unpleasant odors Additional evidence that the insula is important in disgust Understanding the emotions of others may require stimulating and thus mildly experiencing emotions ourselves