Professional Documents
Culture Documents
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.
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)
I. Introduction
II. Synaptic placticity rules
The basic Hebb rule
The covariance rule
BCM Rule
Non-Hebbian rules
Anti-hebbian rules
Timing-based Rules
pre i
(a 0 )
wi
learning rate
post
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
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
dw
w
x( y y )
dt
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
dw
pre
dw
2w w
2 w ( x x ) y
dt
dt
2( y w x ) y
dw
dt
2( y 2 y y ) 0
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
LTP
0
LTD
Postsynaptic activity
constant
2
w
j constant
j
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
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
(Royer and
Pare 2003,
Nature)
LTD
induction
produces
heterosyna
ptic LTP
(Royer and
Pare 2003,
Nature)
Result is
similar
with high
frequency
stimuli
I. Introduction
II. Synaptic placticity rules
The basic Hebb rule
The covariance rule
BCM Rule
Non-Hebbian rules
Anti-hebbian rules
Timing-based Rules
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
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
An intracellular
recording of a pair of
cortical pyramidal
cells in a slice
experiment
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
A e
H (t )
t /
t /
A e
, if t 0
, if t 0.
A variety in plasticity
Supervised learning
Reinforcement learning
Homework
1. Hebb , BCM
? ?
2 . (Homeostatic plasticity)
3 .