Learning and Memory

Learning and MemoryLearning and Memory

Learning and MemoryLearning and MemoryWhat is learning?What is learning?– Changes in our nervous system as a result of experiencesChanges in our nervous system as a result of experiences

What is memory?What is memory?– How these changes are maintained over time and how they are expressed How these changes are maintained over time and how they are expressed

(recall)(recall)

Are they different?Are they different?– Yes, but they are interconnected…Yes, but they are interconnected…

You can’t have memory if you have not learned anythingYou can’t have memory if you have not learned anythingBut, can we have learning if we have no memory?But, can we have learning if we have no memory?

– Perhaps only at birth?Perhaps only at birth?

My thoughts:My thoughts:– Learning and memory make up who we areLearning and memory make up who we are– We are who we are because of all of the experiences we have had throughout We are who we are because of all of the experiences we have had throughout

our lives that have changed us…for better or worse our lives that have changed us…for better or worse

How we study Learning & MemoryHow we study Learning & Memory

Amnesia:Amnesia: The loss of memory, incapable of The loss of memory, incapable of rememberingremembering– Retrograde AmnesiaRetrograde Amnesia

Inability to remember events prior to injury (i.e., some Inability to remember events prior to injury (i.e., some sort of damage to your brain)sort of damage to your brain)Can’t remember your pastCan’t remember your past

– Anterograde AmnesiaAnterograde AmnesiaInability to remember events after injuryInability to remember events after injuryCan’t form new memoriesCan’t form new memories

Time-line of events in one’s life

Injury

Retrograde Amnesia Anterograde Amnesia

The Case of HMThe Case of HMBilateral Medial Bilateral Medial Temporal LobectomyTemporal Lobectomy

– Removed his medial Removed his medial temporal lobes (MTL) temporal lobes (MTL) bilaterally to alleviate bilaterally to alleviate serious epilepsy (1953)serious epilepsy (1953)

HippocampusHippocampusAmygdalaAmygdalaRhinal Cortical AreasRhinal Cortical Areas

– Epilepsy improvedEpilepsy improved

– He was last patient to He was last patient to receive this treatmentreceive this treatment

Why, you might ask?Why, you might ask?

The Case of HMThe Case of HM

After surgery:After surgery:– Intellect was above averageIntellect was above average

IQ actually improved from 104 to 118IQ actually improved from 104 to 118– Normal perceptual and motor abilitiesNormal perceptual and motor abilities– Well-adjusted individualWell-adjusted individual– BUT,BUT,

He was left with very poor memory abilitiesHe was left with very poor memory abilities– Mild retrograde amnesia (~ 2 years prior to surgery)Mild retrograde amnesia (~ 2 years prior to surgery)– Profound anterograde amnesiaProfound anterograde amnesia

The Case of HMThe Case of HMBrenda Milner (1965)Brenda Milner (1965)– ““As far as we can tell, this man retained little if As far as we can tell, this man retained little if

anything of anything of eventsevents subsequent to the operation… subsequent to the operation… Ten months before I examined him, his family had Ten months before I examined him, his family had moved from their old house to a new one a few blocks moved from their old house to a new one a few blocks away on the same street. He still had not learned the away on the same street. He still had not learned the new address (though remembering the old one new address (though remembering the old one perfectly), perfectly), nor could he be trusted to find his way nor could he be trusted to find his way home alonehome alone. He . He did not know wheredid not know where objects in objects in constant use were kept, and his mother stated that he constant use were kept, and his mother stated that he would read the same magazines over and over again would read the same magazines over and over again without finding their contents familiar… without finding their contents familiar… Forgetting Forgetting occurred the instant the patient’s focus of occurred the instant the patient’s focus of attention shiftedattention shifted.”.”

The Case of HMThe Case of HMHM is the most famous patient in the history of HM is the most famous patient in the history of neuroscience research.neuroscience research.

He has served as an invaluable teacher of the He has served as an invaluable teacher of the neurobiology of memory and has been neurobiology of memory and has been extensively tested for over 50 years.extensively tested for over 50 years.

Demonstrated that the medial temporal lobe is Demonstrated that the medial temporal lobe is critical for forming new memories.critical for forming new memories.

But, what can HM do?But, what can HM do?

Many Forms of MemoryMany Forms of Memory

Short-Term Memory (seconds – minutes)

Long-Term Memory (hours – years)

Working Memory (active maintenance)

How Long the Memory Lasts

What Can HM Do?What Can HM Do?Remember a list of 6-7 digitsRemember a list of 6-7 digits– Digit-Span + 1 TestDigit-Span + 1 Test– Performance in normal rangePerformance in normal range

Tap a sequence of 5 blocksTap a sequence of 5 blocks– Block-Tapping Memory Span TestBlock-Tapping Memory Span Test– Performance in normal rangePerformance in normal range

So, HM has relatively intact short-term memorySo, HM has relatively intact short-term memory

Milner (1965): “… Milner (1965): “… Forgetting occurred the instant the Forgetting occurred the instant the patient’s focus of attention shiftedpatient’s focus of attention shifted.”.”





OK in HM

Not OK in HM?



Episodic: breakfast this morning

Semantic: the name of the 44th president

Skill Learning: skiing, riding a bike

Long-Term Memory

Priming: more likely to use a word you heard recently

Conditioning: salivating when I see a nice steak!

Declarative (explicit): things you know that you can tell others

Procedural (non-declarative, implicit): things you know that you can show by doing

OBAMA!

What Can HM Do?What Can HM Do?HM can learn new behavioral HM can learn new behavioral skillsskills

Mirror-Drawing TaskMirror-Drawing Task

HM’s performance improves HM’s performance improves with training sessionswith training sessions– Normal sensory-motor Normal sensory-motor

learninglearning

He has He has nono conscious conscious recollection of ever performing recollection of ever performing it beforeit before

What Can HM Do?What Can HM Do?HM can learn new behavioral HM can learn new behavioral skillsskills

Rotary-Pursuit TestRotary-Pursuit Test

HM’s performance improves HM’s performance improves with training sessionswith training sessions– Normal sensory-motor Normal sensory-motor

learninglearning

He has He has nono conscious conscious recollection of ever performing recollection of ever performing it beforeit before

What Can HM Do?What Can HM Do?HM can show normal HM can show normal primingpriming– More likely to use a word More likely to use a word

if you have heard it if you have heard it recentlyrecently

Repetition Priming TestRepetition Priming Test

He has He has nono conscious conscious recollection of the words recollection of the words on the original liston the original list

Examine List of Words:

PuppyMushroomPancakeSalsaSpicy

Fill in the Blanks:

__us__r__o__m

What Can HM Do?What Can HM Do?

HM can show HM can show normal recognition normal recognition memory and memory and primingpriming

Incomplete Incomplete Pictures TestPictures Test

Recognition

What Can HM Do?What Can HM Do?Recognition















What Can HM Do?What Can HM Do?Recognition Priming


















What Can HM Do?What Can HM Do?HM can learn HM can learn conditioning tasksconditioning tasks

Pavlovian ConditioningPavlovian Conditioning

HM’s performance HM’s performance improves with training improves with training sessionssessions

He has He has nono conscious conscious recollection of ever recollection of ever performing it beforeperforming it before

Prior to Eye-Blink Conditioning:

Tone (CS) ≠ Eye Blink (UR)

Training:

Tone (CS) + Air Puff (US) = Eye Blink (UR)

Conditioned State:

Present Tone (CS) alone = Eye Blink (CR)





OK in HM

Not OK in HM?

…sort of, but he can form some kinds of long-term memories…



Episodic: breakfast this morning

Semantic: the name of the 44rd president

Skill Learning: skiing, riding a bike

Long-Term Memory

Priming: more likely to use a word you heard recently

Conditioning: salivating when I see a nice steak!



Impaired OK! OK! OK! OK!

What has HM Taught US?What has HM Taught US?Supported ideas that Supported ideas that there are many different there are many different kinds of memorykinds of memory

Different brain regions Different brain regions seem to be more seem to be more important for some kinds important for some kinds of memory, but not othersof memory, but not others

Forming (consolidating) Forming (consolidating) new episodic memories is new episodic memories is dependent on MTLdependent on MTL

Multiple Memory Systems?Multiple Memory Systems?

Why do we need 2 parallel memory systems?Why do we need 2 parallel memory systems?

Although subjects with MTL amnesia can form Although subjects with MTL amnesia can form procedural (implicit) memories, they can not transfer that procedural (implicit) memories, they can not transfer that memory to a new or different context (situation)memory to a new or different context (situation)

Two parallel memory systems enables us to use the Two parallel memory systems enables us to use the information we have gathered more flexibly.information we have gathered more flexibly.

Long-Term Memory



Other Case StudiesOther Case StudiesPatient RBPatient RB– Ischemic brain injury during Ischemic brain injury during

surgerysurgery

– Suffered from MTL Suffered from MTL amnesia (similar to HM)amnesia (similar to HM)

– Killed cells in Killed cells in CA1CA1 subfield subfield of hippocampusof hippocampus

– Perhaps it is just CA1 Perhaps it is just CA1 subfield of hippocampus subfield of hippocampus that is important for forming that is important for forming new long-term episodic new long-term episodic memoriesmemories

Other Case StudiesOther Case StudiesPatient NAPatient NA– Stabbed up the nose with a Stabbed up the nose with a

miniature fencing foilminiature fencing foil

– Resulted in slight retrograde Resulted in slight retrograde amnesia which got better over amnesia which got better over timetime

– Also had anterograde episodic Also had anterograde episodic amnesia, but this was less amnesia, but this was less consistentconsistent

Past events would Past events would spontaneously enter his spontaneously enter his consciousnessconsciousness

– Damage restricted to medial Damage restricted to medial diencephalondiencephalon

Mammilary bodies of the Mammilary bodies of the hypothalamushypothalamusMediodorsal thalamic nucleusMediodorsal thalamic nucleus

Other Case StudiesOther Case StudiesKorsakoff’s SyndromeKorsakoff’s Syndrome– Observed in chronic alcoholics (vitamin deficiency)Observed in chronic alcoholics (vitamin deficiency)

– Similar memory impairments as in MTL amnesiaSimilar memory impairments as in MTL amnesiaProminent anterograde amnesia (declarative memories)Prominent anterograde amnesia (declarative memories)

– But, severe retrograde amnesia develops later on in But, severe retrograde amnesia develops later on in the course of the disorderthe course of the disorder

– Damaged areas:Damaged areas:Mammilary bodies, mediodorsal thalamus, neocortex, Mammilary bodies, mediodorsal thalamus, neocortex, hippocampus, & cerebellumhippocampus, & cerebellum

Other Case StudiesOther Case StudiesAlzheimer’s DiseaseAlzheimer’s Disease– Memory deterioration greater than what is seen in a Memory deterioration greater than what is seen in a

normal ageing populationnormal ageing populationMore general than seen with MTL amnesia & Korsakoff’s More general than seen with MTL amnesia & Korsakoff’s SyndromeSyndrome

– Progressive and Terminal: eventually loose abilities to Progressive and Terminal: eventually loose abilities to perform simple activities (eating, speaking, perform simple activities (eating, speaking, recognizing family, bladder control)recognizing family, bladder control)

– Anterograde & Retrograde amnesia for explicit Anterograde & Retrograde amnesia for explicit memorymemory

– Impaired short-term memory & implicit memory for Impaired short-term memory & implicit memory for verbal and perceptual materialverbal and perceptual material

– Sensory motor learning intactSensory motor learning intact

Other Case StudiesOther Case Studies

Alzheimer’s DiseaseAlzheimer’s Disease– Degeneration of Degeneration of

acetylcholine-acetylcholine-producing cells in producing cells in basal forebrainbasal forebrain

– Amyloid plaquesAmyloid plaques– Neurofibrillary tanglesNeurofibrillary tangles

Twisted fibers within Twisted fibers within neuronsneurons

– Widespread cortical Widespread cortical damagedamage

Another Case StudyAnother Case Study

Clive WearingClive Wearing– Herpes EncephalitisHerpes Encephalitis

– Primary damage to hippocampusPrimary damage to hippocampusSome damage to amygdala, temporal, and frontal Some damage to amygdala, temporal, and frontal lobeslobes

http://www.youtube.com/watch?v=5ObnErfTblY

http://www.youtube.com/watch?v=5ObnErfTblY

Storing MemoriesStoring MemoriesOverall, we have now seen that amnesia results from the Overall, we have now seen that amnesia results from the inability to form new memories, or store them over timeinability to form new memories, or store them over time– But, what do we mean by storing a memory?But, what do we mean by storing a memory?

Memory Storage = Memory ConsolidationMemory Storage = Memory Consolidation– Change the memory from a labile/vulnerable state to a stronger, Change the memory from a labile/vulnerable state to a stronger,

more permanent statemore permanent state

Donald Hebb (late 1940’s)Donald Hebb (late 1940’s)– Postulated that memories are held in a short-term reverberating Postulated that memories are held in a short-term reverberating

circuit until they are consolidated to last over a long period of circuit until they are consolidated to last over a long period of timetime

Post-traumatic Amnesia: Evidence Post-traumatic Amnesia: Evidence for Memory Consolidationfor Memory Consolidation

Concussion or ComaConcussion or Coma– Usually results in amnesia for events occurring just Usually results in amnesia for events occurring just

after regaining consciousnessafter regaining consciousnessFailure to convert short-term memories to long-term Failure to convert short-term memories to long-term memoriesmemories

– Permanent retrograde amnesia for events just prior to Permanent retrograde amnesia for events just prior to the injurythe injury

– Older memories are spared!Older memories are spared!Suggests that these memories have been protected by some Suggests that these memories have been protected by some mechanism and are stronger than newly formed memoriesmechanism and are stronger than newly formed memories

Memory ConsolidationMemory ConsolidationElectroconvulsive Shock (ECS)Electroconvulsive Shock (ECS)– Used to study memory Used to study memory

consolidation with the consolidation with the assumption that disrupting assumption that disrupting neural activity would erase the neural activity would erase the memories that had not been memories that had not been consolidated yetconsolidated yet

One-trial learningOne-trial learning– Thirsty rats allowed to discover Thirsty rats allowed to discover

the location of water spoutthe location of water spout– ECS applied more than 1 hour ECS applied more than 1 hour

after training did not disrupt after training did not disrupt memory for the location of memory for the location of water spoutwater spout

– Suggests that consolidation Suggests that consolidation takes place within 1 hourtakes place within 1 hour

Memory ConsolidationMemory Consolidation

Human studies show Human studies show that ECS (used to that ECS (used to treat severe treat severe depression) can depression) can disrupt memories up disrupt memories up to 3 years oldto 3 years oldWhy are memories up Why are memories up to 3 years old affected to 3 years old affected by ECS?by ECS?

Memory ConsolidationMemory ConsolidationHuman studies show that ECS (used to treat severe Human studies show that ECS (used to treat severe depression) can disrupt memories up to 3 years olddepression) can disrupt memories up to 3 years old

Suggests consolidation is a longer, ongoing process that Suggests consolidation is a longer, ongoing process that takes more time than a short-term reverberating circuit takes more time than a short-term reverberating circuit would allowwould allow

Hebb’s basic postulate is likely incorrectHebb’s basic postulate is likely incorrect

Reconsolidation:Reconsolidation: recalling a memory seems to put that recalling a memory seems to put that memory into a more labile state that can still be memory into a more labile state that can still be disrupted before it is consolidated againdisrupted before it is consolidated again

Reconsolidation Failures:Reconsolidation Failures:Misinformation EffectMisinformation Effect

Loftus, Miller, & Burns (1978)Loftus, Miller, & Burns (1978)– Subjects watched a video of a car accident. A sports car stopped at a Subjects watched a video of a car accident. A sports car stopped at a

yieldyield sign. sign.

– Subjects were then interviewed 20 min – 1 week laterSubjects were then interviewed 20 min – 1 week later

Consistent ConditionConsistent Condition: “Did another car pass the sports car while it was : “Did another car pass the sports car while it was stopped at the stopped at the yieldyield sign?” sign?”

Neutral ConditionNeutral Condition: “Did another car pass the sports car while it was : “Did another car pass the sports car while it was stopped?”stopped?”

Inconsistent ConditionInconsistent Condition: “Did another car pass the sports car while it was : “Did another car pass the sports car while it was stopped at the stopped at the stopstop sign?” sign?”

– Subjects then shown pictures of a stop sign and a yield sign. Subjects Subjects then shown pictures of a stop sign and a yield sign. Subjects had to point to the sign they saw in the video.had to point to the sign they saw in the video.

Reconsolidation Failures:Reconsolidation Failures:Misinformation EffectMisinformation Effect

Loftus, Miller, & Burns Loftus, Miller, & Burns (1978)(1978)

Consistent questions Consistent questions reinforced the correct reinforced the correct memorymemory

Inconsistent question Inconsistent question deteriorated the correct deteriorated the correct memorymemory

– Biased subjects towards Biased subjects towards reconsolidating the reconsolidating the incorrect memory!incorrect memory!

020

4060

8010

0

Consistent Info

Neutral Info

Inconsistent Info

Retention Interval

Per

cent

Cor

rect

0 20 1 2 1 min. day days week

Hippocampus and ConsolidationHippocampus and Consolidation

HM:HM:– Temporally graded retrograde amnesia and severe anterograde Temporally graded retrograde amnesia and severe anterograde

episodic amnesiaepisodic amnesia– Suggests that hippocampus and related cortical areas are Suggests that hippocampus and related cortical areas are

important for consolidating episodic memories for long-term important for consolidating episodic memories for long-term storage in other cortical areasstorage in other cortical areas

Over time, memories are stored in distributed networks of Over time, memories are stored in distributed networks of cortical areas and become less and less connected to the cortical areas and become less and less connected to the hippocampushippocampus– Engram – the change in the brain that represents a memoryEngram – the change in the brain that represents a memory

The search for the Engram: Where is the memory stored?The search for the Engram: Where is the memory stored?

Need to systematically test which brain areas are important Need to systematically test which brain areas are important for memory storagefor memory storage

Animal Models of MemoryAnimal Models of Memory

Delayed Nonmatch-Delayed Nonmatch-to-Sample Task: to-Sample Task: monkey versionmonkey version– Test of object Test of object

recognition memoryrecognition memory

Delayed Nonmatch-to-Sample TaskDelayed Nonmatch-to-Sample Task

Monkey results are Monkey results are inconclusiveinconclusive– Damaged Damaged

hippocampus, hippocampus, amygdala, and rhinal amygdala, and rhinal cortexcortex

Rodent studies only Rodent studies only damage small damage small portions of parietal portions of parietal cortex, leaving rhinal cortex, leaving rhinal cortex intactcortex intact

Animal Models of MemoryAnimal Models of MemoryDelayed Nonmatch-to-Delayed Nonmatch-to-Sample Task: Sample Task: rodent rodent versionversion– Mumby BoxMumby Box– Test of object recognition Test of object recognition

memorymemory

Object Recognition Engram?Object Recognition Engram?

Rodent studies suggest that recognition Rodent studies suggest that recognition memory for objects is dependent on rhinal memory for objects is dependent on rhinal cortex, not hippocampus or amygdalacortex, not hippocampus or amygdala

What is the hippocampus important for?What is the hippocampus important for?

Hippocampus and Spatial MemoryHippocampus and Spatial Memory

Morris Water Maze

Pool

“Hidden”Platform


Morris Water Maze

Pool

“Hidden”Platform

Hippocampal lesions impair rats’ ability to learn the location of the platform

Control Rat

HippocampalLesion Rat

Eight Arm Radial MazeEight Arm Radial MazeHungry rats will learn Hungry rats will learn which arms contain which arms contain rewardsrewards– Reference MemoryReference Memory

Rats will learn to not Rats will learn to not enter an arm more enter an arm more than once on a given than once on a given dayday– Working MemoryWorking Memory

Hippocampal lesions Hippocampal lesions produce major deficits produce major deficits on reference and on reference and working memory working memory measuresmeasures

Where should I go???


Hippocampus seems to be Hippocampus seems to be critical for spatial memory, but critical for spatial memory, but why is this?why is this?

Place Cells!Place Cells!– Cells that fire when the subject Cells that fire when the subject

occupies a particular location occupies a particular location in the environmentin the environment

Place cells seem to represent Place cells seem to represent a a cognitivecognitive component of the component of the environmentenvironment– If the cells fire in a certain If the cells fire in a certain

pattern, the rat will behave a pattern, the rat will behave a certain waycertain way


Theories of Hippocampal FunctionTheories of Hippocampal Function– Cognitive Map HypothesisCognitive Map Hypothesis

Hippocampus contains a cognitive map of allocentric space that can Hippocampus contains a cognitive map of allocentric space that can be used to navigate to locations in an environmentbe used to navigate to locations in an environment

– Configural Association TheoryConfigural Association TheoryHippocampus is critical for learning the significance of combinations Hippocampus is critical for learning the significance of combinations of stimuli (i.e., you will be learning about biopsychology in this room, of stimuli (i.e., you will be learning about biopsychology in this room, not any other topic)not any other topic)

– Spatial ArrangementsSpatial ArrangementsHippocampus is important for recognizing the arrangement of Hippocampus is important for recognizing the arrangement of objects relative to one another (i.e., the layout of your living room)objects relative to one another (i.e., the layout of your living room)

Milner (1965)Milner (1965):: “… He still had not learned the new “… He still had not learned the new address (though remembering the old one perfectly), address (though remembering the old one perfectly), nor nor could he be trusted to find his way home alonecould he be trusted to find his way home alone. He . He did not know wheredid not know where objects in constant use were objects in constant use were kept…”kept…”

Evolutionary PerspectivesEvolutionary PerspectivesHippocampus is Hippocampus is important for spatial important for spatial memory in memory in Food Caching Food Caching BirdsBirds

Larger hippocampi than Larger hippocampi than non-food caching birdsnon-food caching birds

Hippocampus grows Hippocampus grows when food caching when food caching behaviors beginbehaviors begin– Developmental changesDevelopmental changes– Seasonal changesSeasonal changes

Evolutionary PerspectivesEvolutionary PerspectivesHippocampus is important for spatial memory in Hippocampus is important for spatial memory in Non-Non-human Primateshuman Primates

Spatial view cells in monkey hippocampusSpatial view cells in monkey hippocampus– Fire if monkey is looking at a particular location in the Fire if monkey is looking at a particular location in the

environmentenvironment– Analogous to place cellsAnalogous to place cells

Hippocampal damage produces deficits on some spatial Hippocampal damage produces deficits on some spatial taskstasks

Inconsistent results likely due to the fact that monkeys Inconsistent results likely due to the fact that monkeys are not allowed to freely move around during are not allowed to freely move around during experimentsexperiments

Evolutionary PerspectivesEvolutionary PerspectivesHippocampus is Hippocampus is important for spatial important for spatial memory in memory in HumansHumans

Hippocampal activation Hippocampal activation during performance of during performance of virtual navigation tasksvirtual navigation tasks

Place-like cells recorded Place-like cells recorded in humans performing in humans performing similar virtual taskssimilar virtual tasks

Evolutionary PerspectivesEvolutionary PerspectivesHippocampus is important for spatial memory in Hippocampus is important for spatial memory in HumansHumans

London Taxi cab drivers have larger hippocampi London Taxi cab drivers have larger hippocampi than normal subjectsthan normal subjects

Milner (1965)Milner (1965)– “… “… He still had not learned the new address (though He still had not learned the new address (though

remembering the old one perfectly), remembering the old one perfectly), nor could he be nor could he be trusted to find his way home alonetrusted to find his way home alone. He . He did not did not know whereknow where objects in constant use were kept…” objects in constant use were kept…”

Hippocampus: Imagining the Future?Hippocampus: Imagining the Future?

Do you remember Clive Wearing???

The Truth About Learning and Memory:The Truth About Learning and Memory:There’s a lot more to it!There’s a lot more to it!

Mediodorsal Mediodorsal Thalamus & Basal Thalamus & Basal ForebrainForebrain– Alzheimer’s DiseaseAlzheimer’s Disease– Korsakoff’s SyndromeKorsakoff’s Syndrome– Patient NAPatient NA


Inferotemporal CortexInferotemporal Cortex– Visual patternsVisual patterns

““What” component of What” component of visual systemvisual system

– Visual Recognition MemoryVisual Recognition Memory– ProsopagnosiaProsopagnosia


AmygdalaAmygdala– Emotional memory (fear Emotional memory (fear

conditioning)conditioning)– Lesions disrupt ability to Lesions disrupt ability to

learn CS-Shock learn CS-Shock associationsassociations

– Rodent and Human data Rodent and Human data are very consistentare very consistent

– Imposter SyndromeImposter Syndrome


Prefrontal CortexPrefrontal Cortex– Temporal order of Temporal order of

events (i.e., steps events (i.e., steps involved in cooking)involved in cooking)

– Short-term and Short-term and Working MemoryWorking Memory


CerebellumCerebellum– Implicit sensorymotor Implicit sensorymotor

tasks (eyeblink tasks (eyeblink conditioning)conditioning)

– Motor Learning Motor Learning Feedback loopFeedback loop

““Efference Copy”Efference Copy”


StriatumStriatum– Part of Extrapyramidal Part of Extrapyramidal

Sensory Motor systemSensory Motor system– Important for Important for

“allowing” appropriate “allowing” appropriate movements to occurmovements to occur

– Implicit sensory motor Implicit sensory motor tasks (habit formation)tasks (habit formation)

– Affected in Parkinson’s Affected in Parkinson’s diseasedisease

Documents

Learning and Memory