Behaviour and the Brain 2:

Cognitive Neuroscience
Johannes Fahrenfort
[email protected]
Medical Faculty, room B563

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Neuroplasticity and memory
• Do large structural changes take place after
birth, and if yes, how?
• How much is genetically hardwired?
• How much is driven by experience?
• How to implement dynamic changes
(memory)?
• What are (some of) the mechanisms and
functions of memory?
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The brain as a tabula rasa
(= empty slate)?

• Empiricists (such as David Hume):

the mind at birth is a “tabula rasa”
• Nature (genes) vs nurture (experience)
debate
• Changes after birth? How does the brain
balance stability with change?

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 Neuroplasticity:
Structural changes

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Deafferentiation

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Remapping after amputation

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 Cortical reorganization
in ated brains
Auditory
processing in
visual areas!

V1

Tactile
processing in
visual areas!
fl
Cortical reorganization

https://doi.org/10.7554/eLife.82747
 Neuroplasticity
• Brain can perform long lasting
reorganization
• But also fast reorganization:
V1 starts responding to non-visual
stimulation such as touch within
2 days of visual deprivation

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 Jugglers

VBM of area V5 (motion)

New neurons?
Increased cell size? Glia? 10
 Clinical applications
• Lazy eye: covering good eye
• Constraint therapy: constraining normally
functioning limb/hand
GOOD eye

GOOD
hand
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 Clinical applications
• Lazy eye: covering good eye
• Constraint therapy: constraining normally
functioning limb/hand
GOOD eye

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How? Neuromodulation
• At a macro level, plasticity is driven by task relevance
• Attentional processes involve acetylcholine release,
enabling plasticity (change) in the brain

Acetylcholine release

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The brain uses the available tissue!
Tadpole

Like V1
Ocular dominance
Squishing Expanding
columns!

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 Important principle!
• A given neuron in a given location is not
dedicated to any given function
• What it does, is determined by its inputs
and outputs, not by its location!

Ferret, rewiring of
visual input to
auditory cortex

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 The sensitive period
• Most (but not all!) rewiring happens at an early age
• Pruning: the brain starts with a 1015 connections at
3 years of age and ends with about 1014
connections in adult brain:

This means that between

50% and 90% of the
connections are lost!

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 The sensitive period
• Most (but not all!) rewiring happens at an early age
• Pruning: the brain starts with a 1015 connections at
3 years of age and ends with about 1014
connections in adult brain:
• Clearly visible
in language
development: This means that between
50% and 90% of the
connections are lost!

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 How much is determined by genetics?
• Roger Sperry
(1960s) cut the
optic nerve of an
adult newt
(salamander)
• Rotated eye ball
• Axonal landing
pattern is
genetic /
independent
from experience!
How much is determined by genetics?
• Humans have only 25000 protein coding genes
• How does it build an approximately 100 billion
(100.0000.0000.0000) neuron system from
these genes? Interaction with the environment!

(a) Embodied perception

(b) Passive perception
 Rat neurons with and
without sensory deprivation

A. Normal
B. Enriched
C. Enriched C
B
D. Deprived
E. Deprived
F. Deprived A
D E F
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Cortical columns arise from experience!
Tadpole

Like V1
Ocular dominance
Squishing Expanding
columns!

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 Pruning
• Most wiring is caused by neural pruning
50% of the neurons die off:
‣ Necrosis: uncontrolled die-off
‣ Apoptosis: controlled ‘sculpting’

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 Neurotrophins

• Rita Levi-Montalcini (1941) 1968 Nobel Prize

• Nerve growth factor chemicals secreted by

the target neurons that allow neurons to
develop, survive and function
• In addition: synaptotoxic chemicals eliminate
synapses

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 Speed of plasticity
• Fast, unmasking:
existing connections are disinhibited, as in the
example of the blindfolded person’s V1
• Slow, axonal growth: new connections form

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neurotrophins
 Memory:
Dynamic changes

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 Memory vs plasticity
• Plasticity involves making
structural changes to a network
• This seems inef cient and in exible
for many types of memory
• So how then…?

26
fi
fl
Perceptron: memory stored in the strength
of the connections (weights) of a network

-- - -
+ ++ ++

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 Changing of connection strengths
Hebb’s rule

Long term
potentiation (LTP)

Long term
depression (LTD)

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Changing of connection strengths

NMDA receptor only activates when

presynaptic and postsynaptic cell re
together, causing synaptic strengthening
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fi
 The Hop eld network (1982)
Attractor states Content
Recurrent addressable
connections memory

Paper: ’Neural networks and physical systems with

emergent collective computational abilities’
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fi
 Content addressable memory
Not just binding, but one can
also imagine how a full
memory can be retrieved
from a coalition by only
activating the features ‘green’
and ‘square’. This would
activates the entire ensemble,
i.e. the memory of the green
square moving to the right.
Attractor state
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 Ensemble coding
1. Objects Memories may be represented by coalitions of
neurons representing the individual features
components of an object a memory.
2. Individual ‘nodes’ may participate in different ensembles,
forming memories

Not entirely clear how ensembles are formed aside from

weights, needs a ‘code’ like rate coding or synchrony
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 Explicit memory:
Medial Temporal Lobe (MTL)
Hippocampus
• Acts as a copying
mechanism
• place cells: ensembles
maximally active for
particular locations,
independent of head
position

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London cab drivers
new neurons!
>
VBM of posterior
hippocampus

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 Neurogenesis?
• Generating new neurons in hippocampus:
plasticity after all?
• Cab drivers (but also remember jugglers)
• It could be that these either store memory or merely
provide scaffolding to form new synaptic connections
• Mechanistic role in memory not yet known

Immature growing cells

hippocampus
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 Explicit memory:
Medial Temporal Lobe (MTL)
Hippocampus
• place cells: maximally active
for particular locations

Enthorhinal cortex
• grid cells: map location in
the environment

Nobel prize in 2014

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Grid cells

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 So memory is synaptic
strengthening, ensemble coding and/
or neurogenesis in MTL…?

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 Working memory

Skeletal musculature

hippocampus

Sensory regions

Frontoparietal network
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 Memory:
A massive lumping error

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 What is the phenomenon
under investigation?
• Lumping errors describing two or more mechanisms
in the brain as a single cognitive function
‣ “Memory” may not be one thing
‣ “Consciousness” may not be one thing
‣ “Attention” may not be one thing
• Splitting errors describing a single mechanism in the
brain as multiple cognitive functions
‣ “Attention” may be constituted by the same mechanism
as “Imagery” as “Working memory” etc

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 General features of
memory
• Embodied, intricately intertwined with
nearly all human functions
• Both accurate (high storage) and inaccurate
(constructive)
• The past is only useful if it aids in predicting
the future / what’s out there: imagery

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 Example:
Perception and memory

Perception cannot be achieved

without memory!
Perception is a prediction (inference)
about input based on past experience

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The indigenous Indian visiting Paramaribo
and ‘modernity’ for the rst time

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fi
 ????

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 One only ‘sees’ things that can
connect with existing knowledge

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 One only ‘sees’ things that can
connect with existing knowledge

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 Memory for action
contributes to recognition

to h
e e m w i t
’ t s m
s n
e ro on b l e
o
D a p pti
v e r ce
h a p e
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 Memory for action
contributes to recognition

a y
th w i o n
p a n i t
s a l co g
o r ed
D odi
b
Em
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 General features of
memory
• Embodied, intricately intertwined with
nearly all human functions
• Both accurate (high storage) and inaccurate
(constructive)
• The past is only useful if it aids in predicting
the future / what’s out there: imagery

50
Memory can both be precise
and imprecise

• Memory can be high capacity and

precise
• But can also be highly ‘constructive’,
involving confabulation in which
memories are ‘ lled-in’

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fi
 Feats of memory
The average person
can recognize about
5,000 faces, although
some super-
recognizers may be
able to identify
10,000 or more…

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Feats of memory
Although hard to
quantify, memory for
places is similarly
large. It is often very
easy to recognize a
known place based
on minimal cues.

(Next to Wertheim park)

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Feats of memory
People with superior
memory often make
use of the same
systems that are used
Retrosplenial cortex
for spatial navigation.

Some theories say

that all memory
systems have evolved
Hippocampus from spatial memory.

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Stephen Wiltshire
Many movies, also see this one: https://www.youtube.com/watch?v=aEevjAr1geE

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Memory can both be precise
and imprecise

• Memory can be high capacity and

precise
• But can also be highly ‘constructive’,
involving confabulation in which
memories are ‘ lled-in’

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fi
Memory like a camera?

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 False memories
The constructive nature of memory

Elisabeth Loftus

https://www.ted.com/talks/
elizabeth_loftus_how_reliable_is_your_memory
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 Why is memory both precise
and imprecise?
• Precise; sometimes exact information is
required:
‣ Who is that person?
‣ Where is my food?
• Imprecise; be effective, prevent overload of
irrelevant information
‣ No need to remember what I ate two weeks ago
‣ Have a exible/generative system that explores
hypothetical scenarios and predict the future
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fl
Why is memory both precise
and imprecise?
• Remembering the past is mostly useful to
be able to predict the future!
• Where can I hide? Where can I get food?
Who will help me? etc etc
• Prospective memory: rembering the future
“mental time travel” (Tulving, 1985)
• Imagination: generating unknown scenarios!

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 General features of
memory
• Embodied, intricately intertwined with
nearly all human functions
• Both accurate (high storage) and inaccurate
(constructive)
• The past is only useful if it aids in predicting
the future / what’s out there: imagery

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Networks involved in memory
and in imagining the future

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Mental imagery / simulation

Place cells

Grid
cells

PPA
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Working memory vs
mental imagery

64 https://doi.org/10.1016/j.cub.2013.05.065
 Working memory vs
mental imagery

V1
V2
V3

Splitting error!
65 https://doi.org/10.1016/j.cub.2013.05.065
 General features
• Embodied, intricately intertwined with
nearly all human functions
• Both accurate (high storage) and inaccurate
(constructive)
• The past is only useful if it aids in predicting
the future / what’s out there: imagery

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Questions

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