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LECTURE 10

Plasticity and Learning

I. Introduction
II. Synaptic placticity rules
The basic Hebb rule
The covariance rule
BCM Rule
Non-Hebbian rules
Anti-hebbian rules
Timing-based Rules

Introduction

Hebbs postulate
When an axon of cell A is near enough to excite cell B
or repeatedly or persistently takes part in firing it, some growth process
or metabolic change takes place in one or both cells such that As
efficiency, as one of the cells firing B, is increased.

Donald O. Hebb (1949)

The theory is often summarized as "cells that fire together,


wire together".

Conditioning:
The first attempt to model conditioning in terms of
synaptic change.
Behavior ---?--- neural mechanisms

Development:
The formation and refinement of
neural circuits need synaptic
elimination.
Axonal or synaptic competition
in neuromuscular junctions and visual
system (Consumptive and interference
competition)

Long term potentiation (LTP) - Long term depression


LTD
Changes that persist for tens of minutes or longer are
generally called LTP and LTD. It lasts for hours in vitro
and days and weeks in vivo
The longest-lasting forms appear to require protein
synthesis.
First found in Hippocampus
The physiological basis of Hebbian learning
Properties and mechanisms of long-term synaptic
plasticity in the mammalian brain may relate to learning
and memory.
Inhibitory synapses can also display plasticity, but this
has been less thoroughly investigated both
experimentally and theoretically

I. Introduction
II. Synaptic placticity rules
The basic Hebb rule
The covariance rule
BCM Rule
Non-Hebbian rules
Anti-hebbian rules
Timing-based Rules

The basic Hebb rule


dwi (t )
w
axi (t ) y (t )
dt

pre i

(a 0 )

wi

learning rate

xi and y : the firing rates of the pre- and postsynaptic neurons


1. Local mechanism
2. Interactive mechanism
3. Time-dependent mechanism

post

The basic Hebb rule is unstable

pre

dw
w
xy
dt
2
dw
dw
2
w
2 w w
2w xy 2 y 0
dt
dt

w
post

1. The processes of synaptic plasticity are typically much slower than the
neural activity dynamics.
2. If, in addition, the stimuli are presented slowly enough to allow the
network to attain its steady-state activity during training,

dy
y wj f (x j )
dt
j

Assume f ( x j ) x j , we have y w j x j w x
j

Theoretically, an upper saturation constraint must be imposed to avoid


unbounded growth. But experimentally,

LTP and LTD at the Schaffer collateral inputs to the CA1


region of a rat hippocampal slice

Is it due to that the basic Hebb rule has no LTD? Lets add LTD by
introducing the covariance rule

I. Introduction
II. Synaptic placticity rules
The basic Hebb rule
The covariance rule
BCM Rule
Non-Hebbian rules
Anti-hebbian rules
Timing-based Rules

The covariance rule

dw
w
x( y y )
dt

postsynaptic threshold, e.g. y

dw
w
(x x ) y
dt
presynaptic threshold, e.g.

dw
w
(x x ) y
dt
(x x )(x x ) w
the input covariance matrix,

dw
w
x ( y y )
dt

(homosynaptic depression)

dw
w
( x x ) y
dt

(heterosynaptic depression)

By the basic Hebb rule, synapses are modified whenever correlated preand postsynaptic activity occurs. Such correlated activity can occur purely
by chance, rather than reflecting a causal relationship that should be
learned. To correct for this, the covariance rather than correlation-based
rule is often used by network models

Although the covariance rule allows LTD and reflects a causal preand postsynaptic relationship it is still unstable due to positive
feedback

The covariance rules, like the Hebb rule, are


unstable and non-competitive
dw
w
( x x ) y
dt
w

dw

pre

dw
2w w
2 w ( x x ) y
dt
dt
2( y w x ) y

Average above equation over the training period:

dw
dt

2( y 2 y y ) 0

Competition can be introduced to allowing threshold to slide as


follows

w
post

I. Introduction
II. Synaptic placticity rules
The basic Hebb rule
The covariance rule
BCM Rule
Non-Hebbian rules
Anti-hebbian rules
Timing-based Rules

BCM Rule
Bienenstock, Cooper and Munro (1982) proposed an alternative for
which there is experimental evidence where the postsynaptic threshold is
dynamic

dw
w
xy ( y y )
dt

Hebb rule

covariance rule

One example:

d y
dt

y y

Usually set:

LTP

0
LTD

BCM rule
Postsynaptic activity

This is again unstable if is fixed.


However, if the threshold is allowed to grow faster than v we
get stability.
depends on postsynaptic activity. For instance, the threshold
for LTP decreases when postsynaptic activity is low (y
)
Here competition
between synapses appears
since strengthening some
synapses results in threshold
increasing meaning that it is
harder for others to be
strengthened

LTP

0
LTD

Postsynaptic activity

Synaptic weight normalization

constant

2
w
j constant
j

It is a more direct way of enforcing competition


Idea is that postsynaptic neuron can only support a certain
amount of total synaptic weight so strengthening one leads to
weakening others
2 types: subtractive normalisation and multiplicative
normalisation

Subtractive normalisation
dw
y
w
xy n
dt
Nx

or :
dwi
y
w
xi y
dt
Nx

(n x x j ; n w w j )
j

constant

It is easy to prove that the total increase in the weights is 0.

Evidences for
BCM rule
Evidence for a sliding
threshold:
It is easier to obtain LTP
in the cortex of darkreared animals and it is
harder to induced LTD in
these cortices
The field potentials evoked in layer III by layer IV stimulation in
slices of visual cortex prepared for light-deprived and control rats 4-6
weeks old
The effects can be reserved by as little as two days of light
exposure before slice preparation

Experimental evidences for constant total


synaptic weights
Low- and high-frequency BLA stimuli (LFS, HFS) are
known to, respectively, produce homosynaptic NMDA
dependent LTD and LTP in ITC cells.
Whether LFS and HFS also produce inverse heterosynaptic
modifications is unclear.

(Royer and
Pare 2003,
Nature)

slices of the amygdala


guinea-pigs (35 weeks old)
intercalated (ITC) neurons of the amygdala:
the basolateral amygdala (BLA):
an array of closely spaced (~150 m) stimulating electrodes

Homosynaptic LTP was induced with


HFS paired to postsynaptic depolarization.
Postsynaptic depolarization was achieved
by applying short (2ms) depolarizing
current pulses (0.2 nA) timed so that BLAevoked EPSPs would occur just before or
during current-evoked spikes

(Royer and Pare 2003, Nature)

Plot of EPSP amplitude and rise


time versus stimulation site

LTD
induction
produces
heterosyna
ptic LTP
(Royer and
Pare 2003,
Nature)

Left: Difference between pre- and post-LFS response profiles


(EPSP amplitudes) for one cell (top) and average of all cells
Right:Time course of changes in response amplitude

Result is
similar
with high
frequency
stimuli

(Royer and Pare 2003, Nature),

Their results showed that the activity-dependent


enhancement or depression of particular inputs to intercalated
neurons is accompanied by inverse modifications at
heterosynaptic sites, which contributes to total synaptic weight
stabilization

The inverse homo- versus heterosynaptic plasticity seems to


be a cell- wide event, which needs an intracellular signaling
system that can render synapses aware of each other or of the
mean neuronal activity.

How do unstimulated inputs detect the stimulation frequency


at the stimulated pathway?

I. Introduction
II. Synaptic placticity rules
The basic Hebb rule
The covariance rule
BCM Rule
Non-Hebbian rules
Anti-hebbian rules
Timing-based Rules

Non-Hebbian forms of synaptic plasticity


They modify synaptic strengths solely on the basis of pre- or
postsynaptic firing, are likely to play important roles in
homeostatic, developmental, and learning processes
Homeostatic plasticity
It allows neurons to sense how active they are and to
adjust
their properties to maintain stable function
Loosely defined, a homeostatic form of plasticity is one
that
acts to stabilize the activity of a neuron or neuronal circuit
in the face of perturbations, such as changes in cell size or
in synapse number or strength, that alter excitability.
A large number of plasticity phenomena have now been
identified (e.g., synaptic scaling and homeostasis of

Synaptic scaling
A form of synaptic plasticity that adjusts the strength of all of a
neuron's excitatory synapses up or down to stabilize firing,
avoiding quiescence and hyper-excitation at the level of
individual neurons.
Current evidence suggests that neurons detect changes in their
own firing rates through a set of calcium-dependent sensors
Review paper:
Gina G. Turrigiano. The Self-Tuning Neuron: Synaptic Scaling
of Excitatory Synapses. Cell 135: 422-435, October 31, 2008

A model of multiplicative scaling through the


removal of AMPA receptors

(Turrigiano 1999, TINS)

Homeostasis of intrinsic excitability of neurons


Activity can also modify the intrinsic excitability and
response properties of neurons
Models of such intrinsic plasticity show that neurons can be
remarkably robust to external perturbations if they adjust their
conductances to maintain specified functional characteristics
Intrinsic and synaptic plasticity can interact in interesting
ways. For example, shifts in intrinsic excitability can
compensate for changes in the level of input to a neuron caused
by synaptic plasticity.

Homeostasis of intrinsic excitability of neurons

Theoretical and experimental work suggests that intracellular Ca2+


concentration might regulate the balance of inward and outward currents
generated by a neuron
(Turrigiano 1999, TINS)

Anti-Hebbian plasticity
It causes synapses to decrease (rather than increase) in
strength when there is simultaneous pre- and postsynaptic
activity.
It is believed to be the predominant form of plasticity at
synapses in mormyrid electric fish and those from parallel fibers
to Purkinje cells in the cerebellum
Anti-Hebbian modification tends to make weights decrease
without bound

dwi (t )
w
axi (t ) y (t )
dt

(a 0 )

I. Introduction
II. Synaptic placticity rules
The basic Hebb rule
The covariance rule
BCM Rule
Non-Hebbian rules
Anti-hebbian rules
Timing-based Rules

Timing-Based Rules

LTP is induced by repetitive


stimulation with positively
correlated spike times of post and
pre-synaptic neuron

LDP is induced by repetitive


stimulation with negatively
correlated spike times of post and
pre-synaptic neuron

Spike Timing Dependent Plasticity (STDP)

An intracellular
recording of a pair of
cortical pyramidal
cells in a slice
experiment

(Markram et al., 1997)

LTP and LTD of


retinotectal synapses
recorded in vivo in
Xenopus tadpoles

(Zhang et al., 1998)

Simulating the spike-timing dependence of synaptic plasticity


requires a spiking model (e.g. Integrate-and-Fire Models).
However, an approximate model can be constructed on the
basis of firing rates

dwi
w
d [ H ( ) y (t ) xi (t ) H ( ) y (t ) xi (t )]
0
dt

where

LTP

LTD

H ( ) H ( )
Note above equation is based on a Hebbian rule
The STDP rule describes an asymmetric learning rule

About H(): a function like the solid line in previous figure.

A e

H (t )

t /

t /

A e

, if t 0

, if t 0.

Sequence learning based on STDP

The Timing-Based plasticity rule is applied throughout a training


period during which the stimulus being presented moves to the right
and excites the different neurons in the network sequentially
After the training period, the neuron with sa = 0 receives strengthened
input from the sa =2 neuron and weakened input from the neuron with
sa = 2

If the same time-dependent stimulus is presented again after


training, the neuron with sa = 0 will respond earlier than it did
prior to training
The training experience causes neurons to learn a time
sequence

Another example on time sequence learning in place


fields

Place field is negatively skewed after experience


(Mehta et al. 1997; 2000)

A variety in plasticity

Different cortical regions, such as


hippocampus and visual cortex have
somewhat different forms of synaptic
plasticity.

(Abbott and Nelson 2000, Nature)

A few properties of LTP and LTD


Long-term plastic changes can be induced in about 1 s or less
(i.e. within a rather short period, similar to short-term
plasticity)
The induced change in synaptic weight typically lasts for
hours (if no further changes are induced)
The longest-lasting forms appear to require protein synthesis

Three types of training procedures


Unsupervised (or sometimes self-supervised) learning.

A network responds to a series of inputs during training


solely on the basis of its intrinsic connections and
dynamics

Supervised learning

A desired set of input-output relationships is imposed on


the network by a teacher during training.
Networks that perform particular tasks can be
constructed in this way

Reinforcement learning

It is somewhat intermediate between these cases.


The network output is not constrained by a
teacher, but evaluative feedback on network
performance is provided in the form of reward or
punishment

Homework

1. Hebb , BCM
? ?
2 . (Homeostatic plasticity)
3 .