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Title: Learning and Memory


1
Learning and Memory
  • Mind and Brain
  • Chapter 13

2
  • You are responsible for Chapter 13 (text and
    notes) for you final as well as all other
    chapters (and notes) covered in class.

3
  • http//www.youtube.com/watch?vJliczINA__Yfeature
    related

4
Chapter Overview
  • The Nature of Learning
  • Synaptic Plasticity Long-Term Potentiation and
    Long-Term Depression
  • Perceptual Learning
  • Classical Conditioning
  • Instrumental Conditioning
  • Relational Learning

5
The Nature of Learning
  • Introduction
  • Learning refers to the process by which
    experiences change our nervous system and hence
    our behavior we refer to these changes as
    memories
  • Experiences are not stored they change the way
    we perceive, perform, think, and plan
  • They do so by physically changing the structure
    of the nervous system, altering neural circuits
    that participate in perceiving, performing,
    thinking and planning.

6
The Nature of Learning
  • Learning can take at least 4 basic forms
  • Perceptual Learning
  • Stimulus-Response Learning
  • Classical Conditioning
  • Instrumental Conditioning
  • Motor Learning
  • Relational Learning

7
The Nature of Learning
  • Perceptual Learning
  • Learning to recognize a particular stimulus
  • Primary function ability to identify and
    categorize objects and situations
  • Each sensory system is capable of perceptual
    learning
  • Accomplished by changes in the sensory
    association cortex

8
The Nature of Learning
  • Stimulus-Response Learning
  • Learning to automatically make a particular
    response in the presence of a particular stimulus
  • Involves the establishment of connections between
    circuits involved in perception and those
    involved in movement
  • Classical conditioning
  • Instrumental conditioning

9
The Nature of Learning
  • Classical Conditioning
  • Unconditional Stimulus (US) stimulus that
    produces a defensive or appetitive response.
  • Unconditional Response (UR) response to the US.
  • Conditional Stimulus (CS) stimulus, which when
    paired with the US during training, comes to
    elicit a learned response.
  • Conditional Response (CR) response to the
    presentation of the CS.

10
Figure 13.1 A Simple Neural Model of Classical
Conditioning
11
The Nature of Learning
  • Hebb Rule
  • hypothesis proposed by Donald Hebb that the
    cellular basis of learning involves strengthening
    of a synapse that is repeatedly active when the
    postsynaptic neuron fires.

12
Figure 13.1 A Simple Neural Model of Classical
Conditioning
13
The Nature of Learning
  • Instrumental Conditioning (Operant conditioning)
  • learning procedure whereby the effects of a
    particular behavior in a particular situation
    increase (reinforce) or decrease (punish) the
    probability of the behavior.
  • Reinforcing Stimulus appetitive stimulus that
    follows a particular behavior and thus makes the
    behavior become more frequent.
  • Punishing Stimulus aversive stimulus that
    follows a particular behavior and thus makes the
    behavior become less frequent.
  • CC involves automatic or species-typical
    responses, but OC involves behaviors that have to
    be learned
  • CC involves an association between 2 stimuli, OC
    involves an association between a response and a
    stimulus.
  • OC is considered more flexible because it permits
    an organism to adjust its behavior according to
    the consequences of that behavior.

14
Figure 13.2 A Simple Neural Model of Instrumental
Conditioning
15
The Nature of Learning
  • Motor Learning
  • Learning to make a new response
  • Component of stimulus-response learning

16
Figure 13.3 An Overview of Perceptual,
Stimulus-Response (S-R), and Motor Learning.
17
The Nature of Learning
  • Relational Learning
  • More complex form of learning
  • Involves learning the relationships among
    individual stimuli
  • Includes the ability to recognize objects through
    more than one sensory modality
  • Involves learning the relative location of
    objects in the environment spatial learning
  • Remembering the sequence in which events occurred
    during particular episodes episodic learning
  • Hippocampus

18
Chapter Overview
  • The Nature of Learning
  • Synaptic Plasticity Long-Term Potentiation and
    Long-Term Depression
  • Electrical stimulation of circuits within the
    hippocampus can lead to long-term synaptic
    changes that seem to be responsible for learning
  • Perceptual Learning
  • Classical Conditioning
  • Instrumental Conditioning
  • Relational Learning

19
See Figure 13.4
  • Primary input to the HF comes from the EC
  • Axons of EC neurons pass through the perforant
    path and synapse with granule cells in the DG
  • From these cells, the mossy fibers project to the
    pyramidal cells in CA3
  • From CA3, Schaffer collaterals project to CA1
    cells
  • The 3 synapses know as the trisynaptic loop

20
Induction of LTP
  • A stimulating electrode is placed in the PP and a
    recording electrode is placed in the DG
  • A single pulse of electrical stimulation is
    delivered to the PP and the resulting population
    EPSP is recorded in the DG
  • Population EPSP is the evoked potential that
    represents EPSPs of a population of neurons.
  • LTP can be induced by stimulating the PP axons
    with a burst of electrical pulses (i.e., 100)
    within a few seconds

21
LTP
The size of the first population EPSP tells us
the strength of the synaptic connections before
LTP is induced Evidence that LTP has occurred is
obtained by periodically delivering a single
pulse and then measuring the response in the DG
to see if it is bigger than the original response
22
Synaptic Plasticity LTP and LTD
  • LTP
  • Can be induced in other parts of the HF and in
    other brain regions
  • Can last for several months
  • Can be induced in slices and in living animals
  • Can follow the Hebb Rule (in slices)
  • Associative Long-Term Potentiation long-term
    potentiation in which concurrent stimulation of
    weak and strong synapses to a given neuron
    strengthens the weak ones.

23
Figure 13.6 Associative Long-Term Potentiation
24
Figure 13.7 The Role of Summation in Long-Term
Potentiation
  • Nonassociative LTP requires an additive effect
  • Series of pulses delivered at high rate will
    produce LTP
  • Same of pulses given at slow rate will not
    (LTD)
  • Rapid rate of stimulation causes EPSPs to summate
  • Rapid stimulation depolarizes the postsynaptic
    membrane more than slow stimulation

25
Figure 13.8 Long-Term Potentiation
  • Experiments have shown that synaptic
    strengthening occurs when NTS binds with
    postsynaptic receptors located in a dendrite that
    is already depolarized
  • LTP requires 2 events
  • Activation of synapses
  • Depolarization of the postsynaptic membrane

26
Synaptic Plasticity LTP and LTD
  • Role of NMDA Receptors
  • NMDA Receptor specialized ionotropic glutamate
    receptor that controls a calcium channel that is
    normally blocked by Mg2 ions.
  • Calcium ions enter the cells through channels
    controlled by NMDA receptors only when glutamate
    is present and the postsynaptic membrane is
    depolarized

27
Figure 13.9 The NMDA Receptor
28
Synaptic Plasticity LTP and LTD
  • Role of NMDA Receptors
  • AP5 2-amino-5-phosphonopentanoate, a drug that
    blocks NMDA receptors.
  • Blocks the establishment of LTP

29
Synaptic Plasticity LTP and LTD
  • Need glutamate depolarization.but how do
    dendrites become depolarized if only axons can
    produce action potential?
  • Dendritic Spike action potential that occurs in
    the dendrite of some types of pyramidal cells.
  • Threshold for activation is very high
  • Only occurs when action potential is triggered in
    the axon
  • Backwash of depolarization across cell body
    triggers dendritic spike
  • Whenever the axon of a pyramidal cell fires, all
    of its dendritic spines become depolarized for a
    brief time.

30
Synaptic Plasticity LTP and LTD
  • Simultaneous occurrence of synaptic activation
    and a dendritic spike strengthens the active
    synapse.
  • Magee and Johnston (1997) injected individual CA1
    pyramidal cells in hippocampal slices with a
    fluorescent dye that permitted them to see the
    influx of calcium
  • When individual synapses became active at the
    same time that a dendritic spike had been
    triggered, Ca hot spots occurred near the
    activated synapses
  • Size of EPSPs produced by these activated
    synapses became larger synapse became
    strengthened
  • TTX (blocks Na current) injected near dendrite
    prevented dendritic spikes, no LTP!

31
Synaptic Plasticity LTP and LTD
  • Role of NMDA receptors in Associative LTP
  • If weak synapses are active by themselves,
    nothing happens (NMDA receptors dont open)
  • However, if the activity of strong synapses
    located elsewhere on the postsynaptic cell has
    caused the cell to fire, then a dendritic spike
    will depolarize the postsynaptic membrane enough
    to eject Mg ions from the Ca channels (of NMDA
    receptor)
  • If some synapses then become active, Ca will
    enter the dendritic spines and cause the synapses
    to become strengthened

32
Synaptic Plasticity LTP and LTD
  • What is responsible for the increase in synaptic
    strength that occurs during LTP?
  • Dendrites on CA1 neurons contain 2 types of
    glutamate receptors NMDA and AMPA

33
Synaptic Plasticity LTP and LTD
  • Mechanisms of Synaptic Plasticity
  • AMPA Receptors ionotropic glutamate receptor
    that controls a sodium channel when open it
    produces EPSPs.
  • Strengthening of individual synapses is
    accomplished by the insertion of more AMPA
    receptors into the postsynaptic membrane of the
    dendritic spine
  • CaM-KII type of calcium-calmodulin kinase, an
    enzyme that must be activated by calcium may
    play a role in the establishment of LTP.
  • Nitric Oxide Synthase enzyme responsible for
    the production of nitric oxide.
  • Drugs that block this enzyme prevent the
    establishment of LTP in CA1

34
Figure 13.16 Chemistry of LTP
  • Activation of terminal button releases glutamate,
    which binds with NMDA receptors in the
    postsynaptic membrane of the dendritic spine
  • If the membrane was depolarized by a dendritic
    spike, then calcium ions enter and activate
    CAM-KII
  • CAM-KII travels to the postsynaptic density and
    causes the insertion of AMPA receptors
  • LTP also initiates changes in synaptic structure
    and production of new synapses

35
Figure 13.16 Chemistry of LTP
  • The entry of calcium also activates NO synthase
  • This produces NO which diffuses out of the
    dendritic spine and back to the terminal button
  • The NO may then trigger chemical reactions that
    increase the release of glutamate
  • Long-lasting LTP also requires the synthesis of
    new proteins and the presence of dopamine

36
Synaptic Plasticity LTP and LTD
  • Low-frequency stimulation of the synaptic inputs
    to a cell can decrease their strength
  • Long-Term Depression (LTD) also plays a role in
    learningsome synapses are strengthened and
    others weakened
  • long-term decrease in the excitability of a
    neuron to a particular synaptic input caused by
    stimulation of the terminal button while the
    postsynaptic membrane is hyperpolarized or only
    slightly depolarized.
  • Like LTP, requires activation of NMDA receptors
  • LTD involves a decrease in AMPA receptors

37
Synaptic Plasticity LTP and LTD
  • Other Forms of LTP
  • Some forms of LTP do not involve NMDA receptors
    and are not blocked by AP5 (CA3).
  • For example, mossy fiber input from dentate gyrus
    to CA3
  • Presynaptic changes only no alterations in
    structure of dendritic spines

38
Chapter Overview
  • The Nature of Learning
  • Synaptic Plasticity Long-Term Potentiation and
    Long-Term Depression
  • Perceptual Learning
  • Classical Conditioning
  • Instrumental Conditioning
  • Relational Learning

39
Figure 13.18 The Major Divisions of the Visual
Cortex of the Rhesus Monkey
  • Primary visual cortex receives information from
    the lateral geniculate nucleus of the thalamus
  • Ventral Stream pathway of information from the
    primary visual cortex to the temporal lobe, which
    is involved in object recognition (what
    pathway).
  • Dorsal Stream pathway of information from the
    primary visual cortex to the parietal lobe, which
    is involved with perception of the location of
    objects (where pathway).

40
Chapter Overview
  • The Nature of Learning
  • Synaptic Plasticity Long-Term Potentiation and
    Long-Term Depression
  • Perceptual Learning
  • Classical Conditioning
  • Instrumental Conditioning
  • Relational Learning

41
Figure 13.21 Conditioned Emotional Responses
Information about the CS US converge in the LA
so synaptic changes responsible for learning
could take place in this area Hebb rule - weak
synapses (from tone) are strengthened when US
activates neurons in the LA.LA neurons fire and
activate CN which evokes the response (Ch 11)
42
Classical Conditioning
  • Evidence for the involvement of lateral nucleus
    of the amygdala in CER
  • Changes in the LA responsible for CER learning
    involve LTPand is accomplished through
    activation of the NMDA receptor
  • LTP
  • Injection of drugs that block LTP into the
    amygdala prevents the establishment of
    conditioned emotional responses
  • CER training (tone-shock pairings) causes AMPA
    receptors to be driven into dendritic spines of
    synapses between LA neurons and axons that
    provide auditory input

43
Classical Conditioning
Rumpel et al., (2005) used a virus to insert a
gene for a fluorescent dye coupled to a subunit
of the AMPA receptor into the LA of ratslearning
caused AMPA insertion
They also inserted a gene for a dye coupled to a
defective subunit of the AMPA receptorthe
defective subunit prevented AMPA insertion and
conditioning did not take place
44
Classical Conditioning
  • Infusion of many drugs into the LA that prevent
    LTP disrupt acquisition of a CER
  • Conclusion
  • LTP in the amygdala, mediated by NMDA receptors,
    plays a critical role in the establishment of CER

45
Chapter Overview
  • The Nature of Learning
  • Synaptic Plasticity Long-Term Potentiation and
    Long-Term Depression
  • Perceptual Learning
  • Classical Conditioning
  • Instrumental Conditioning
  • Relational Learning

46
Instrumental Conditioning
  • Instrumental conditioning involves a connection
    between a particular stimulus and a particular
    response
  • 2 major pathways between sensory association
    cortex and motor association cortex
  • Direct transcortical connections
  • Involved in episodic memory (along with HIP)
  • Connections via the basal ganglia and thalamus
  • 2 pathways play different roles

47
Basal Ganglia
  • Learned behaviors become automatic and routine,
    they are transferred to the basal ganglia
  • Leaving the transcortical circuits free to learn
    new tasks

48
Figure 13.23 The Basal Ganglia and Their
Connections
  • Neostriatum (caudate nucleus putamen) receives
    sensory input from all regions of the cerebral
    cortex. Also receives information from frontal
    lobes about movement (planned or in progress).
  • Outputs are sent to GP which sends information
    back to frontal cortex to premotor cortex (where
    plans for movement are made) and motor cortex
    (where movement is executed)

49
Instrumental Conditioning
  • Basal Ganglia
  • Studies of laboratory animals have indicated
    that
  • lesions of the basal ganglia disrupt instrumental
    conditioning without affecting other forms of
    learning.
  • Fernadez-Ruiz et al., (2001) destroyed portions
    of the caudate and putamen that receive visual
    information from the ventral stream
  • Lesions did not disrupt visual perceptual
    learning
  • Impaired the monkeys ability to learn to make a
    visually guided operant response
  • Williams and Eskandar (2006) as monkeys learned
    a operant response, the rate of firing of single
    neurons in the caudate nucleus increased
  • The activity of caudate neurons is correlated
    with rate of learning
  • Blocking NMDA receptors in the basal ganglia with
    an injection of AP5 disrupts learning guided by a
    simple visual cue

50
Instrumental Conditioning - Reinforcement
  • Neural circuits involved in reinforcement
    discovered by Olds Milner, 1954.
  • Reinforcement
  • Neural Circuits Involved in Reinforcement
  • Ventral Tegmental Area (VTA) group of
    dopaminergic neurons in the ventral midbrain
    whose axons form the mesolimbic and mesocortical
    systems and are important in reinforcement.
  • Nucleus Accumbens nucleus of the basal
    forebrain near the septum receives dopamine from
    neurons of the VTA and is thought to be involved
    in reinforcement and attention.
  • See Figure 13.24

51
Figure 4.13 Dopaminergic Pathways in a Rat Brain
52
Instrumental Conditioning
  • Reinforcement
  • Microdialysis studies have revealed that release
    of DA in the NA is caused by
  • reinforcing electrical stimulation of the medial
    forebrain bundle (connects VTA to NA) or the VTA
  • administration of cocaine or amphetamine
  • presence of natural reinforcers such as water,
    food, sex partner
  • fMRI indicates activity in NA during reinforcing
    events

53
Instrumental Conditioning
  • Functions of the Reinforcement System
  • To detect reinforcing stimuli.
  • To strengthen the connections between the neurons
    that detect the discriminative stimulus (i.e.,
    sight of lever) and the neurons that produce the
    instrumental response (i.e., lever press).

54
Instrumental Conditioning
  • Reinforcement occurs when neural circuits detect
    a reinforcing stimulus and cause the activation
    of dopaminergic neurons in the VTA
  • A stimulus that serves as a reinforcer on one
    occasion may fail to do so on another
  • Reinforcement system is not automatically
    activated when particular stimuli are present
    activation also depends on the state of the animal

55
Instrumental Conditioning
  • Detecting reinforcing stimuli
  • Reinforcement system appears to be activated by
    unexpected reinforcing stimuli
  • First DA in VTA responded rapidly when the
    reinforcing stimuli was present
  • Once the animals learn the task, VTA neurons are
    activated to the learned stimuli
  • If a reinforcing stimulus does not occur when
    expected, the activity of dopaminergic neurons
    decreases
  • Berns et al., (2001) increased activity in NA
    (fMRI) when tasty drink was given unpredictably,
    no increase if predictable
  • Activity of these neurons sends a signal that
    there is something to be learned

56
Instrumental Conditioning
  • Prefrontal cortex provides input to VTA
  • PFC involved in devising strategies, making
    plans, evaluating progress toward a goal.
  • May turn on reinforcement mechanism when it
    determines that ongoing behavior is close to goal

57
Instrumental Conditioning
  • Strengthening neural connections
  • DA induces synaptic plasticity by facilitating
    associative LTP
  • NA, amygdala, prefrontal cortex

58
Chapter Overview
  • The Nature of Learning
  • Synaptic Plasticity Long-Term Potentiation and
    Long-Term Depression
  • Perceptual Learning
  • Classical Conditioning
  • Instrumental Conditioning
  • Relational Learning

59
Relational Learning
  • Includes the establishment and retrieval of
    memories of events, episodes and places

60
Relational Learning
  • Human Anterograde Amnesia
  • Anterograde Amnesia amnesia for events that
    occur after some disturbance to the brain.
  • Retrograde Amnesia amnesia for events that
    happened before some disturbance to the brain.

61
Figure 13.27 A Schematic Definition of Retrograde
Amnesia and Anterograde Amnesia
62
Relational Learning
  • Human Anterograde Amnesia
  • Korsakoffs Syndrome permanent anterograde
    amnesia caused by brain damage from chronic
    alcoholism or malnutrition.
  • Medial temporal lobe damage also produces
    anterograde amnesia (i.e., H.M.)
  • Based on extensive work with H.M., Milner
    colleagues concluded
  • The hippocampus is not the location of LT
    memories nor is it necessary for the retrieval
    of LT memories
  • The hippocampus is not the site of immediate (ST)
    memories
  • The hippocampus is important for converting ST
    memories into LT memories

63
Relational Learning
  • Human Anterograde Amnesia
  • Consolidation process by which short-term
    memories are converted to long-term memories.

Figure 13.28 A Simple Model of the Learning
Process
64
Relational Learning
  • Spared Learning Abilities
  • Not a total failure in learning ability
  • Perceptual Learning
  • Visual recognition of incomplete objects (Figure
    13.29)
  • Face Recognition
  • Stimulus-Response Learning
  • HM could learn a classically conditioned eyeblink
    response
  • Instrumental conditioning task visual
    discrimination task in which pennies were given
    for correct responses
  • Motor Learning
  • Serial reaction time task

65
Figure 13.30 The Serial Reaction Time Task
66
Relational Learning
  • So, even though amnesics can perform some tasks
    they have no memory of having ever learned
    themled to the notion that the brain has
    multiple memory systems
  • Declarative and Nondeclarative Memories
  • Declarative (explicit) Memory memory that can
    be verbally expressed.
  • Nondeclarative (implicit) Memory memory whose
    formation does not depend on the hippocampal
    formation a collective term for perceptual,
    stimulus-response, and motor memory.
  • Appear to be automatic, do not require deliberate
    attempts to memorize
  • Acquisition of specific behaviors and skills
  • Do not need to be able to describe these
    activities in order to do them

67
Table 13.1
  • Declarative Memory Tasks
  • Remembering past experiences
  • Finding ones way in new environment
  • Nondeclarative Memory Tasks
  • Learning to recognize broken drawings
  • Learning to recognize pictures and objects
  • Learning to recognize faces
  • Learning to recognize melodies
  • Classical conditioning
  • Instrumental conditioning
  • Learning sequence of button presses

68
Relational Learning
  • Anatomy of Anterograde Amnesia
  • Damage to the hippocampus or to regions of the
    brain that supply its inputs and receive its
    outputs causes anterograde amnesia
  • HIP formation CA fields, DG and SUB major
    input is from EC
  • Outputs of Hip (from CA1 and SUB) also go back to
    EC, and PC and parahip cortex
  • Perirhinal Cortex region of limbic cortex
    adjacent to the hippocampal formation that relays
    information between entorhinal cortex and other
    regions of the brain.
  • Parahippocampal cortex region of the limbic
    cortex adjacent to the hippocampal formation that
    shares the same general role as the perirhinal
    cortex.

69
Cortical Connections of the Hippocampal Formation
  • Figure 13.31

70
Figure 13.32 The Major Subcortical Connections of
the Hippocampal Formation
  • Hip also receives subcortical input via the
    fornix
  • Fornix carries dopaminergic input from VTA
  • Fornix also connects the HIP with the mammillary
    bodies (located in post. Hypo.)
  • MMB degenerate in Korsakoffs syndrome

71
Relational Learning
  • Role of the Hippocampal Formation in
    Consolidation of Declarative Memories
  • HIP not really important for STM or LTM but seems
    to be important for declarative memory formation
  • HIP receives input from sensory and motor cortex
    and from subcortical regions (BG and AMG), it
    then processes this information and sends
    projections back to these same regions and
    somehow modifies the memories being stored there
  • Hippocampus is involved in modifying memories as
    they are being formed.
  • The order in which events occurred
  • Contextual information
  • Relationships among elements

72
Rational Learning
  • Hippocampus
  • Time-limited role
  • Anterograde amnesia is usually accompanied by
    retrograde amnesia
  • The duration of the retrograde amnesia related to
    the amount of damage to the MTL
  • Damage limited to hipp retrograde amnesia
    lasting few years
  • Damage to hipp entrorhinal cortex retrograde
    amnesia 1-2 decades
  • Damage to MTL spared memories from early life
  • Gradual process controlled by hipp transforms
    memories located elsewhere
  • Before transformation is complete, hipp is
    required for retrieval of memories
  • Later, retrieval of memories can occur if hipp
    has been damaged

73
Relational Learning
  • Declarative Memory
  • Episodic and Semantic Memories
  • Episodic Memory memory of a collection of
    perceptions of events organized in time and
    identified by a particular context.
  • Specific to a particular time and place
  • Semantic Memory memory of facts and general
    information.
  • Do not include information about the context in
    which the facts were learned
  • Episodic memory must be learned all at once
  • Semantic Memory - can be acquired gradually
  • Requires the hippocampus

74
  • Semantic Dementia loss of semantic memories
    caused by progressive degeneration of the
    neocortex of the lateral temporal lobes.
  • Difficulty naming objects
  • Difficulty understanding the meaning of words

75
Relational Learning
  • Spatial Memory
  • Memory for spatial information used to move
    around ones environment and get from one place
    to another.
  • Role of Hippocampus in Spatial Memory
  • Damage to (right) HIP causes spatial memory
    impairments
  • London taxi drivers have bigger HIP than control
    subjects
  • longer an taxi driver had spent in this
    occupation, the larger the volume of rt hipp
  • Place Cells special neurons in the hippocampus
    that are directly involved in navigation in space
    (rats).
  • Virtual-reality towns
  • Spatial Strategy maze learning strategy based
    on spatial cues activation of HIP (fMRI).
  • Response Strategy maze learning strategy based
    on a series of responses (turns) activation of
    caudate nucleus.
  • People who tend to use spatial strategies
    larger hipp
  • People who tend to follow response strategies
    larger caudate

76
Relational Learning
  • Relational Learning in Laboratory Animals
  • Spatial Perception and Learning
  • Morris Water Maze

77
Morris Water Maze
78
Spatial Learning
  • Damage to the hippocampus animals swim in what
    appears to be an aimless fashion until they
    finally encounter the platform
  • Hippocampal lesions disrupt navigation in homing
    pigeons
  • Hippocampus of birds and rodents that store seeds
    in hidden caches and later retrieve them is
    larger than that of animals that dont

79
Relational Learning
  • Relational Learning in Laboratory Animals
  • Hippocampal Place Cells
  • Found in dorsal hippocampus in rats (corresponds
    to the posterior hippocampus in humans)
  • fire at a high rate when the animal is in a
    particular location, called the cells place
    field
  • fire at a low rate when the animal is in a
    different location
  • First discovered by OKeefe Dostrovsky

80
OKeefe Dostrovsky (1971)
81
Relational Learning
  • Relational Learning in Laboratory Animals
  • Role of Hippocampal Formation in Memory
    Consolidation
  • Time limited role in memory
  • Mice trained in water maze inactivate HIP with
    lidocaine 1 day later or 30 days later (Figure
    13.40)
  • Memory Reconsolidation
  • A process of consolidation of a memory that
    occurs subsequent to the original consolidation
    that can be triggered by a reminder of the
    original stimulus.

82
Phases of MemoryReconsolidation
Acquisition - the pairing of the context/cue to
the aversive stimuli
Train
Reactivate
Test
Evidence suggests that reactivation of a memory
can return it to a labile state requiring
reconsolidation via protein synthesis
83
Figure 13.41 A Schematic Description of the
Experiment by Misanin, Miller, and Lewis (1968)
84
Reconsolidation of Memories
  • Reconsolidation requires LTP
  • Injection of anisomycin (blocks protein synthesis
    and prevents memory consolidation), blocks
    reconsolidation

85
Relational Learning
  • Relational Learning in Laboratory Animals
  • Role of LTP in memory
  • When rats learn mazes, strength of population
    EPSP in CA3 increases
  • Mutations targeted at NMDA receptors in CA1
  • Prevented establishment of LTP
  • Poor spatial learning on Morris water maze
  • Hippocampal Neurogenesis
  • New neurons can be produced in the hippocampus of
    the adult brain (DG)
  • Training on relational tasks increases
    neurogenesis
  • Easier to establish LTP with these new neurons
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