1. Psychology 250 Lecture 12 Kevin R Smith

2. Three Types of Behavior Reflexes: involuntary responses to stimuli Reflexes: involuntary responses to stimuli Instincts: stereotyped responses triggered by environmental stimuli Instincts: stereotyped responses triggered by environmental stimuli Learning: a relatively permanent change in behavior due to experience Learning: a relatively permanent change in behavior due to experience © Renee Purse/Photo Researchers, Inc.

3. Types of Learning Nonassociative learning: a change in the magnitude of response to environmental events Nonassociative learning: a change in the magnitude of response to environmental events HabituationHabituation SensitizationSensitization Associative learning: a connection between two elements or events Associative learning: a connection between two elements or events Classical conditioningClassical conditioning

4. Habituation and Sensitization Habituation Habituation The response to steady or repeated stimulus decreases over timeThe response to steady or repeated stimulus decreases over time Example: You don’t hear your air conditioner after it’s been running awhileExample: You don’t hear your air conditioner after it’s been running awhile Sensitization Sensitization The experience of one stimulus heightens the response to a subsequent stimulusThe experience of one stimulus heightens the response to a subsequent stimulus Example: People are “jumpy” following natural disasters, like earthquakesExample: People are “jumpy” following natural disasters, like earthquakes

5. Classical Conditioning and Pavlov Dogs eating habits Dogs eating habits Noticed that the dogs salivate prior to getting their foodNoticed that the dogs salivate prior to getting their food Measured the amount of salivation that occurs when the dogs are anticipating foodMeasured the amount of salivation that occurs when the dogs are anticipating food Anticipation cues were Pavlov at first Anticipation cues were Pavlov at first Changed to a toneChanged to a tone

6. Pavlovian Conditioning Before Training Before Training No salivation to a toneNo salivation to a tone Salivation to meat onlySalivation to meat only After Training After Training Salivation to a toneSalivation to a tone

7. Classical Conditioning A conditioned stimulus (CS) is an initially neutral stimulus that acquires the ability to signal important biological events A conditioned stimulus (CS) is an initially neutral stimulus that acquires the ability to signal important biological events An unconditioned stimulus (UCS) is an event that elicits a response without prior experience An unconditioned stimulus (UCS) is an event that elicits a response without prior experience A conditioned response (CR) is a learned reaction to a CS A conditioned response (CR) is a learned reaction to a CS An unconditioned response (UCR) is an unlearned reaction to a UCS An unconditioned response (UCR) is an unlearned reaction to a UCS

8. Using Aplysia Californica to Study Learning The gill is used for breathing The gill is used for breathing The gill can be covered with the mantle shelf The gill can be covered with the mantle shelf Waste and seawater are released through the siphon Waste and seawater are released through the siphon The gill-withdrawal reflex occurs when touching the siphon produces a retraction of the gill The gill-withdrawal reflex occurs when touching the siphon produces a retraction of the gill Touching the siphon repeatedly yields a decrease in the gill withdrawal reflex: Habituation

9. Eric Kandel and Habituation in Aplysia The siphon is served by 24 sensory neurons, whose cell bodies are in the abdominal ganglion The siphon is served by 24 sensory neurons, whose cell bodies are in the abdominal ganglion Siphon sensory neurons form synapses with: Siphon sensory neurons form synapses with: excitatory and inhibitory interneuronsexcitatory and inhibitory interneurons six motor neurons serving the gillsix motor neurons serving the gill

10. Possible Hypotheses for Habituation Do sensory neurons become less responsive? Do sensory neurons become less responsive? Single cell recordings from sensory neurons did not decrease with repeated touching of the siphonSingle cell recordings from sensory neurons did not decrease with repeated touching of the siphon Daniel L. Geiger/SNAP

11. More Hypotheses Does the gill muscle lose its ability to contract? Does the gill muscle lose its ability to contract? Electrical stimulation of the motor neurons produced muscle contraction, even after habituationElectrical stimulation of the motor neurons produced muscle contraction, even after habituation Do changes occur at the synapses between the sensory and motor neurons? Do changes occur at the synapses between the sensory and motor neurons?

12. Kandel’s Explanation of Habituation Repeated siphon touches reduce entry of calcium into sensory neurons, which release less neurotransmitter Repeated siphon touches reduce entry of calcium into sensory neurons, which release less neurotransmitter Smaller EPSPs occur in interneurons and motor neurons Smaller EPSPs occur in interneurons and motor neurons Motor neurons produce weaker response, leading to less gill withdrawal Motor neurons produce weaker response, leading to less gill withdrawal

13. Aplysia also show sensitization After a electrical shock to the head or tail, there is an increased gill- withdrawal response when the siphon is touched After a electrical shock to the head or tail, there is an increased gill- withdrawal response when the siphon is touched

14. Sensitization in Aplysia Shocking the tail stimulates interneurons that form synapses with sensory neurons serving the siphon Shocking the tail stimulates interneurons that form synapses with sensory neurons serving the siphon Sensory neurons release more neurotransmitter onto motor neurons, and the gill retracts strongly Sensory neurons release more neurotransmitter onto motor neurons, and the gill retracts strongly

15. Long-term Changes Normal Aplysia showed 1300 axon terminals on sensory neurons Normal Aplysia showed 1300 axon terminals on sensory neurons Aplysia experiencing sensitization had 2800 terminals Aplysia experiencing sensitization had 2800 terminals Aplysia experiencing habituation had 800 terminals Aplysia experiencing habituation had 800 terminals

16. Classical Conditioning in Aplysia Electrical shock to the tail leads to gill-withdrawal reflex Electrical shock to the tail leads to gill-withdrawal reflex Touching the mantle alone does not trigger anything Touching the mantle alone does not trigger anything Pairing the mantle with the shock leads to gill-withdrawal reflex Pairing the mantle with the shock leads to gill-withdrawal reflex After learning, touching the mantle alone leads to gill-withdrawal reflex After learning, touching the mantle alone leads to gill-withdrawal reflex

17. Classical Conditioning

18. Mechanisms for Classical Conditioning Touching the mantle leads to NT release Touching the mantle leads to NT release Amount released is too small to get a gill-withdrawal reflexAmount released is too small to get a gill-withdrawal reflex Touching the tail leads to an increase in the NT released at the gill- withdrawal reflex Touching the tail leads to an increase in the NT released at the gill- withdrawal reflex After frequent pairings After frequent pairings More NT is released when the mantle is touched by itselfMore NT is released when the mantle is touched by itself

19. Conditioned Emotional Responses and the Amygdala Typical Paradigm: Typical Paradigm: Tone (CS+) followed by shock (UCS) results in reduced feeding (CR) in ratsTone (CS+) followed by shock (UCS) results in reduced feeding (CR) in rats Recording during training shows an increased response of the amygdala to the tone Recording during training shows an increased response of the amygdala to the tone Lesions of the basolateral nucleus of the amygdala prevent the learning of emotional responses Lesions of the basolateral nucleus of the amygdala prevent the learning of emotional responses Blocking NMDA receptors in the amygdala prevents learning of conditioned emotional responses Blocking NMDA receptors in the amygdala prevents learning of conditioned emotional responses Thus: the amygdala is involved in Classical Conditioning Thus: the amygdala is involved in Classical Conditioning

20. Learning and the Cerebellum Cerebellum is involved in motor control Cerebellum is involved in motor control Also probably involved in motor learning Also probably involved in motor learning Highly complex network of neurons controls learning Highly complex network of neurons controls learning

21. Long Term Depression Long term depression Long term depression A weakening of a synapse that lasts hours or daysA weakening of a synapse that lasts hours or days Thought to be associated with learningThought to be associated with learning Found in the cerebellum and hippocampusFound in the cerebellum and hippocampus Long term potentiation Long term potentiation A long lasting strengthening of a synapseA long lasting strengthening of a synapse Thought to be associated with memoryThought to be associated with memory

22. Classical Conditioning of the Rabbit’s Nictitating Membrane A tone (CS) followed by a puff of air to the eye (UCS) leads to closure of the nictitating membrane (UCR) A tone (CS) followed by a puff of air to the eye (UCS) leads to closure of the nictitating membrane (UCR) Activity in the lateral interpositus nucleus (LIP) of the cerebellum increases in response to the tone over trials Activity in the lateral interpositus nucleus (LIP) of the cerebellum increases in response to the tone over trials

23. Human Research and the LIP Humans learn classically conditioned eyeblink responses Humans learn classically conditioned eyeblink responses PET scans show increased cerebellar activity during classical conditioning PET scans show increased cerebellar activity during classical conditioning Individuals with cerebellar damage have difficulty learning conditioned eyeblink responses Individuals with cerebellar damage have difficulty learning conditioned eyeblink responses

24. Memory Memory refers to the storage and retrieval of information Memory refers to the storage and retrieval of information No absolute boundaries between learning and memory No absolute boundaries between learning and memory Learning and memory may be viewed as a continuum Learning and memory may be viewed as a continuum

25. The Atkinson-Shiffrin Model of Memory

26. Divisions of Long-term Memory

27. Locating the Memory Trace (Engram) Karl Lashley observed the effects of lesions on rats’ maze learning Karl Lashley observed the effects of lesions on rats’ maze learning The larger the amount of cortex damaged, the more errors the rats made The larger the amount of cortex damaged, the more errors the rats made Lashley believed that the engram was distributed across the cortex Lashley believed that the engram was distributed across the cortex

28. Brain Stimulation and Memory Wilder Penfield stimulated the cortex of patients undergoing surgery for epilepsy Wilder Penfield stimulated the cortex of patients undergoing surgery for epilepsy 8% of the patients experienced specific memories when their temporal lobes were stimulated 8% of the patients experienced specific memories when their temporal lobes were stimulated Lead to the theory that memories could be localized in the brain Lead to the theory that memories could be localized in the brain Penfield’s results may have been influenced by the patients’ epilepsy Penfield’s results may have been influenced by the patients’ epilepsy

29. Patient H.M. and Memory Large areas of H.M.’s temporal lobes were surgically removed Large areas of H.M.’s temporal lobes were surgically removed H.M.’s personality and IQ were not affected H.M.’s personality and IQ were not affected H.M. experienced profound anterograde amnesia H.M. experienced profound anterograde amnesia Inability to learn new informationInability to learn new information Courtesy Dr. Suzanne Corkin, Massachusetts Institute of Technology

30. Amnesias Retrograde Retrograde Loss of memories of the pastLoss of memories of the past Anterograde Anterograde Inability to form new memoriesInability to form new memories

31. H.M.’s Memories Are Not Equally Affected Short-term memory allowed H.M. to converse Short-term memory allowed H.M. to converse H.M. retained the ability to learn procedural tasks H.M. retained the ability to learn procedural tasks H.M.’s deficits appear in explicit memory tasks H.M.’s deficits appear in explicit memory tasks

32. The Delayed Nonmatching to Sample Task Monkeys with medial temporal lobe damage do poorly on the DNMS task Monkeys with medial temporal lobe damage do poorly on the DNMS task The DNMS task requires the ability to form long-term memories The DNMS task requires the ability to form long-term memories

33. Conclusion thus far The amygdala and the cerebellum seem to play a role in forming associations (very basic learning) The amygdala and the cerebellum seem to play a role in forming associations (very basic learning) Animal studiesAnimal studies The temporal lobe seems to play some role in memories The temporal lobe seems to play some role in memories HM and MonkeysHM and Monkeys What about the Hippocampus? What about the Hippocampus?

34. The Hippocampus and Memories HM had major lesions to his hippocampus HM had major lesions to his hippocampus Anterograde anmesiaAnterograde anmesia LTP occurs in the hippocampus LTP occurs in the hippocampus LTP results when synapses become more efficient LTP results when synapses become more efficient LTP can be demonstrated indefinitely in living animals and for hours in slices of hippocampus LTP can be demonstrated indefinitely in living animals and for hours in slices of hippocampus

35. LTP Shares Characteristics with Long-term Memory Unsure of whether LTP is the mechanism for Long term memories BUT Unsure of whether LTP is the mechanism for Long term memories BUT Both LTP and long-term memories last indefinitelyBoth LTP and long-term memories last indefinitely Both LTP and long-term memories result from very brief inputBoth LTP and long-term memories result from very brief input LTP is consistent with cellular learning models proposed by Donald HebbLTP is consistent with cellular learning models proposed by Donald Hebb NMDA receptors (Glutamate) closely associated with LTP NMDA receptors (Glutamate) closely associated with LTP

36. The Hippocampus and Human Memory The right hippocampus is active during spatial memory processing and the left hippocampus is active during verbal memory processing The right hippocampus is active during spatial memory processing and the left hippocampus is active during verbal memory processing Rostral portions of the hippocampus are more active during encoding, and caudal portions are active during retrieval Rostral portions of the hippocampus are more active during encoding, and caudal portions are active during retrieval The hippocampus does not store memories, but transfers them from short to long-term storage The hippocampus does not store memories, but transfers them from short to long-term storage

37. Patient N.A. A fencing foil produced a lesion to N.A. ’ s left dorsomedial thalamus A fencing foil produced a lesion to N.A. ’ s left dorsomedial thalamus N.A. experienced profound anterograde amnesia and some retrograde amnesia N.A. experienced profound anterograde amnesia and some retrograde amnesia N.A. ’ s memory loss was similar to H. M. ’ s N.A. ’ s memory loss was similar to H. M. ’ s Courtesy L.R. Squire, University of California San Diego

38. The Diencephalon and Memory Korsakoff’s patients show anterograde and retrograde amnesia Korsakoff’s patients show anterograde and retrograde amnesia Thiamine deficiencies result from chronic alcoholismThiamine deficiencies result from chronic alcoholism Untreated thiamine deficiencies damage dorsomedial thalamus and mamillary bodiesUntreated thiamine deficiencies damage dorsomedial thalamus and mamillary bodies Monkeys with lesions in the dorsomedial thalamus have difficulties with the DNMS test Monkeys with lesions in the dorsomedial thalamus have difficulties with the DNMS test Temporal-diencephalic circuits may process long-term memories Temporal-diencephalic circuits may process long-term memories

39. Localization of Semantic Memories Semantic memories are widely distributed in the cortex Semantic memories are widely distributed in the cortex Prosopagnosia: inability to recognize common faces Prosopagnosia: inability to recognize common faces Localization of Memories Localization of Memories Encoding and retrieval may activate different areas Encoding and retrieval may activate different areas

40. Episodic Memory and the Cortex Greater right frontal and temporal activity is associated with hearing your own story rather than another person ’ s Greater right frontal and temporal activity is associated with hearing your own story rather than another person ’ s In source amnesia, a patient forgets when and where a memory was formed In source amnesia, a patient forgets when and where a memory was formed Episodic memory is superior in families where left- handedness is common Episodic memory is superior in families where left- handedness is common

41. Short-term Memory and the Cortex The prefrontal cortex may be involved with short-term memory The prefrontal cortex may be involved with short-term memory Patients with prefrontal damage do poorly on the Wisconsin card-sorting task Patients with prefrontal damage do poorly on the Wisconsin card-sorting task Object permanence tasks: Object permanence tasks: Are mastered by infants at the age of 7 or 8 months Adult monkeys with prefrontal lesions respond like immature human infants © Bob Daemmrich/Stock, Boston

42. The Basal Ganglia and Procedural Memory Lesions of the hippocampus impaired declarative memories (remember where you went last time) Lesions of the hippocampus impaired declarative memories (remember where you went last time) Lesions of the caudate nucleus (basal ganglia) impaired procedural memories Lesions of the caudate nucleus (basal ganglia) impaired procedural memories Patients with Huntington’s disease and Parkinson’s disease have procedural memory deficits, but not declarative memory deficits Patients with Huntington’s disease and Parkinson’s disease have procedural memory deficits, but not declarative memory deficits

43. Genetic Manipulation of Learning Ability Drosophila can learn to avoid odors associated with electric shock Drosophila can learn to avoid odors associated with electric shock Blocking the expression of the Protein Kinase A gene prevents learning and formation of short-term memories Blocking the expression of the Protein Kinase A gene prevents learning and formation of short-term memories Too much CREB-2 blocks long but not short-term memories Too much CREB-2 blocks long but not short-term memories Extra CREB-1 produces immediate long-term memory Extra CREB-1 produces immediate long-term memory

44. Unusual Memory Phenomena Repression Repression Flashbacks Flashbacks Flashbulb memories Flashbulb memories Stress affects the amygdala and hippocampus Stress affects the amygdala and hippocampus Cortisol may damage the hippocampus Cortisol may damage the hippocampus KHBS KHOG/AP/Wide World

45. Eye Witness Testimony Beth Loftus: False Memories Beth Loftus: False Memories Had students go home and ask younger siblings: Hey, do you remember that time when you got lost in the mall? Had students go home and ask younger siblings: Hey, do you remember that time when you got lost in the mall? The siblings had never been lost in the mall The siblings had never been lost in the mall The siblings conjured up detailed stories The siblings conjured up detailed stories Bugs Bunny at Disneyland Bugs Bunny at Disneyland

46. Memory and Normal Aging In healthy adults, most measures of cognitive ability remain stable In healthy adults, most measures of cognitive ability remain stable Memory loss may accompany changes in brain systems that utilize the neurotransmitter acetylcholine (ACh) Memory loss may accompany changes in brain systems that utilize the neurotransmitter acetylcholine (ACh) Attaining an above-average education; engaging in complex, non-routine professions; and earning high income appear to protect against cognitive decline Attaining an above-average education; engaging in complex, non-routine professions; and earning high income appear to protect against cognitive decline