Neuroethology:

Learning & Memory 2

 Learning paradigms for two memory types

Memory

Declarative
Non-Declarative
(‚conscious‘)

Episodic Semantic
learning learning
(events) (facts)

Non-Associative Associative
Procedural
learning learning (classical Priming &
learning
(habituation/ and operant perceptual learning
(skills, habits)
sensitization) conditioning)
Pavlov’s Classical Experiment with Dogs:
1) Before Training. . . .

UnConditioned Stimulus (UCS)

produces
UnConditioned Response (UCR)

2) During Training. . .

UCS is paired with Neutral

(“Conditioned”) Stimulus

3) After Training. . .
Conditioned Stimulus (CS)
produces a
Conditioned Response (CR)
Classical Conditioning of the Gill Withdrawal Reflex:

Conditioning

Sensitization

Conditioning

Sensitization
Classical Conditioning of the Gill Withdrawal Reflex:
Classical Conditioning of the Gill Withdrawal Reflex:

Two “associative” synaptic mechanisms of long-term potentiation (LTP) involved

in the classically conditioned gill withdrawal reflex:

1) Heterosynaptic -- results from simultaneous activity in sensory neurons

and facilitating interneurons.
2) Homosynaptic -- results from simultaneous activity in sensory neurons
and motor neurons
 Sensitization
at the sensory-motor synapse

Facilitation works without

activity in the presynaptic cell.
Classical Conditioning: Heterosynaptic Mechanism (pre-synaptic)

Sensitization - presynaptic cell inactive Classical Conditioning - presynaptic cell active

associative
mechanism!
Classical Conditioning of the Gill Withdrawal Reflex:

Two “associative” synaptic mechanisms long-term potentiation (LTP) involved in

the classically conditioned gill withdrawal reflex:

1) Heterosynaptic -- results from simultaneous activity in sensory neurons

and facilitating interneurons.
2) Homosynaptic -- results from simultaneous activity in sensory neurons
and motor neurons
Classical Conditioning: Homosynaptic mechanism (postsynaptic)

• postsynaptic NMDA receptors are closed at rest

• if postsynaptic cell is activated, NMDA receptors open
• Influx of Ca2+ in motor neuron

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Classical Conditioning: Homosynaptic mechanism (postsynaptic)

• Ca+ influx triggers signal pathways in the cell

• postynaptic receptors are modified
• retrograde signaling activated (?)
• this only works if both pre- and postsynaptic
cells are active = associative mechanism

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 The importance of timing:
Hebbian learning and Spike-timing dependent plasticity (STDP)
Hebbian learning:
"Let us assume that the persistence or repetition of a reverberatory activity (or
"trace") tends to induce lasting cellular changes that add to its stability.… When an
axon of cell A is near enough to excite a cell B and repeatedly or persistently takes
part in firing it, some growth process or metabolic change takes place in one or
both cells such that A's efficiency, as one of the cells firing B, is increased."
‚Cells that fire together, wire together.‘

STDP:

Donald Hebb
STDP and classical conditioning as an example for
unsupervised learning

input output

.. ..
. .

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Summary on neural mechanisms of memory:
Both short-term and long-term forms of memory result from changes in synaptic strength
and synaptic connections.

Short-term Memory is process based.

• Strengthening of pre-existing synapses.
• Relies on action of protein kinases to modify pre-existing proteins.
• No gene transcription and no protein synthesis.
• No growth of new synapses!!

Long-term Memory is structure based.

• Strengthening of pre-existing synapses.
• Relies on action of protein kinases to modify pre-existing proteins.
• Requires gene transcription and protein synthesis
• Requires growth (or loss) of synapses!!
Learning in humans is mostly a loss of excitatory
synapses in cortex

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Operant conditioning: Teaching bees to discriminate visual
patterns via association learning

Test (without sucrose):

Training with sucrose
The bee chooses

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Training Test

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Bees can learn abstract features: example symmetry

trial #

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Example for experiments using classical conditioning:
the proboscis extension reflex

US: sucrose
UR: proboscis extension
CS: odor

sucrose

restrained bee

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Relative timing of CS and US is important

onset of US (sucrose)

time [s]

conditioning works in a small time window only

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VUMmx1: A neural correlate for US (sucrose) - a ‚reward neuron’?

VUMmx1 is activated by sucrose VUMmx1 can substitute sucrose

sucrose presentation
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Time course of classical and operant conditioning
are very similar - same underlying mechanism?

classical

operant

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 Unsupervised and supervised learning -
Classical and operant conditioning
Unsupervised: Supervised:
feed-forward feed-back

input output input output

reward
or
punishment

unsupervised learning is also good for 24

perceptual pattern learning
 From single cells to populations:
Memory engrams
(sensory) output
input engram
The combination of cells that
are active forms an engram

Different cells can take part

in several engrams

.. ..
. .

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Hippocampus plays important role in memory
formation

CA3 Cells

Hippocampus

CA1 Cells
How do we measure and test populations responses?

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Activity dependent gene expression using promotors for
immediate early genes

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Activity dependent gene expression to mark memory
engrams during fear conditioning

30
Optogenetic activation of the engram alone evokes fear
response

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 Learning paradigms for two memory types

Memory

Declarative
Non-Declarative
(‚conscious‘)

Episodic Semantic
learning learning
(events) (facts)

Non-Associative Associative
Procedural
learning learning (classical Priming &
learning
(habituation/ and operant perceptual learning
(skills, habits)
sensitization) conditioning)
 Priming -- an improvement in the ability to
detect or identify objects after recent
experience with them.

• Enduring mental representation of an object’s attributes.

• Important for learning to recognize a particular stimulus.

An example of priming. . .
Priming:
Priming:
Priming:
Priming:

What did you see first?

Learning to Detect Prey: The Search Image

an example of “priming”.
aufpassen: pdfs sollten diesen Teil nicht
enthalten (oder erst hinterher hochladen
Learning to Detect Prey: The Search Image
Priming may help to allocate limited ‚attentional‘ resources

Percent prey in environment

 Perceptual Learning -- an improvement in the
ability to discriminate simple perceptual attributes
as the result of performing the discrimination
repeatedly.

• Enduring mental representation of an object’s attributes.

• Important for learning to recognize a particular stimulus.

Perceptual Learning:
Imprinting - an instance in
which an object acquires
significance as a result of early
exposure.
Konrad Lorenz
Learning to Obtain Food: Tool Use in Birds and Primates
 Learning paradigms for two memory types

Memory

Declarative
Non-Declarative
(‚conscious‘)

Episodic Semantic
learning learning
(events) (facts)

Non-Associative Associative
Procedural
learning learning (classical Priming &
learning
(habituation/ and operant perceptual learning
(skills, habits)
sensitization) conditioning)
Procedural Learning: also called “sensorimotor learning”

The improvement that occurs in the

performance of specific motor patterns with repeated
practice of certain skills or habits.

Important for the acquisition of useful skills.

Some human examples. . .

• riding a bike
• athletic abilities (e.g., a tennis serve)
• typing
• threading a needle
• playing a musical instrument
• language!!
Procedural Learning: also called “sensorimotor learning”

The improvement that occurs in the

performance of specific motor patterns with repeated
practice of certain skills or habits.

Higher Order Learning

Sensory Behavioral
Info. Programming

Sensorimotor Learning
Reflexes
Local Learning

Sensory Motor
Inputs Outputs
Procedural
Task
Learning to Obtain Food: “Übung macht den Meister!”

With experience, prey handling times improve.

an example of skill acquisition

Learning to Obtain Food: “Übung macht den Meister!”

With experience, prey handling times improve.

an example of skill acquisition

 Learning paradigms for two memory types

Memory

Declarative
Non-Declarative
(‚conscious‘)

Episodic Semantic
learning learning
(events) (facts)

Non-Associative Associative
Procedural
learning learning (classical Priming &
learning
(habituation/ and operant perceptual learning
(skills, habits)
sensitization) conditioning)
Long-term Declarative Memory:

Hippocampus plays important role in

converting short-term memories into long-
term memories = consolidation.

CA3 Cells

Hippocampus

CA1 Cells
Long-term Declarative Memory:

Hippocampal cells exhibit LTP = “long-term potentiation”

•Produced by series of repeated

action potentials in the
presynaptic cells (CA3).

•Results in stronger EPSPs in

postsynaptic cells (CA1)

•Two forms: Early and Late

Long-term Declarative Memory:

Retrograde Signal
(NO)
Long-term Declarative Memory:

Early LTP Late LTP

Do other animals have “declarative” memories too?

Mother A Mother B

Control Stimulus

looks at looks at
Mother B speaker

Distress Call of
Mother B’s Baby