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Transcript
Neural Coding
Computing and the Brain
How Is Information Coded in Networks of
Spiking Neurons?
• Coding in spike (AP) sequences from individual neurons
• Coding in activity of a population of neurons
Dept. Computing
Science & Mathematics
Spring 2010
(Single Neuron
Computation, pg 88)
2
1
Coding in Single Neurons
• Binary coding
– Presence or absence of spikes
– Logical 1 if neuron fires an AP in a given time window; 0 otherwise
• Rate coding
g
– Average firing frequency over a given time
– Analogue (number)
• Temporal coding
– Interspike interval
– Temporal sequence of spikes
Dept. Computing
Science & Mathematics
(CS92 Fig 3.4)
Spring 2010
3
Rate Coding in Sensorimotor Systems
• Muscle stretch receptors fire in proportion to stretch (weight)
– Experimental data from Adrian, 1928
• Sensation of a stimulus is proportional to the firing rate
– Hypothesis
yp
by
y Adrian
Dept. Computing
Science & Mathematics
Spring 2010
4
2
Temporal Coding in Sensorimotor Systems
• H1 neuron in the fly visual system
• Responds to movement of objects in the world
– Angular velocity
• Movements can be reconstructed from measurements of the
interspike intervals
Dept. Computing
Science & Mathematics
Spring 2010
(from Spikes, MIT
Press, 1997)
5
Dynamic Range of Codes
• Suppose a neuron is capable of firing from 0 to 100 spikes per sec
• Neurons that receive inputs from this neuron have 200 msecs to
“decode” the signal from the neuron
– Information coded in spikes fired by neuron in a 200 msec time window
• Given this scenario, what “code” is best?
– Rate versus temporal coding
• Rate code
–
–
–
–
Rate is number of spikes divided by time interval
Roughly 0 to 20 spikes could occur in 200 msecs
Only 20 states can be distinguished
0, 5, 10, 15, 20 Hz etc
Dept. Computing
Science & Mathematics
Spring 2010
6
3
Dynamic Range (2)
• Temporal code
–
–
–
–
Measure interspike intervals with 10 msec precision
20 time bins in which spikes can be detected
20 element binary vector
Over one million (220) possible states
• Different spike patterns giving different binary vectors
• 200 msecs is often too long in the real nervous system
– House fly performs visual responses in 30 msecs
– Human can recognise visual objects in 150 msecs
– Time for only 1 or 2 spikes per neuron
Dept. Computing
Science & Mathematics
Spring 2010
7
Coding in Networks
• How are “features” of a stimulus encoded in a network of neurons?
– Eg shape, colour, size etc
• Local coding: single neuron per feature
• Scalar coding:
g feature encoded byy firing
g rate of a single
g neuron
• Vector coding: feature encoded by firing rates of a population of
neurons that have overlapping tuning curves
Dept. Computing
Science & Mathematics
Spring 2010
8
4
Temporal Binding
• How can the activity of neurons responding to different features of a
single stimulus be combined?
• Cell assembly: group of neurons that fire at the same time
• Temporal
p
binding
g
– E.g red sphere and a blue cube
• Coincidence detection
• e.g. Cell C encodes a red sphere
Dept. Computing
Science & Mathematics
Spring 2010
9
Vector Coding of Features
• Three colour system of colour encoding
[8, 4, 0]
[4, 2, 0]
[0, 4, 8]
Dept. Computing
Science & Mathematics
(CS92 Fig 4.15)
[0, 2, 4]
Spring 2010
10
5
Vector Coding of Features
• Visual stimulus orientation
encoding
• Example shows 3 neurons that
respond maximally to different
orientations of a bar of light
• How many neurons with
different tuning curves are
required to encode orientation?
(CS92 Fig 4.21)
Dept. Computing
Science & Mathematics
Spring 2010
11
Prediction of Hand Movements
• Recording from 100 neurons in
groups of 20 from different
areas of motor cortex
• Monkey moves joystick to track
onscreen cursor
• Predictions of hand movements
based on:
1. Average firing rate of each
neuron in 100msec time bins
2. 10 bins per neuron, covering 1
second
d off activity
ti it iimmediately
di t l
preceding movement
(Wessberg et al, Nature 408:361-365, 2000)
Dept. Computing
Science & Mathematics
Spring 2010
12
6
Prediction of Hand Movements (2)
• Linear and nonlinear (ANN)
models
– Inputs are neuronal firing rates
– Output
O tp t is predicted mo
movement
ement
• Initial model parameters
derived from first minute of
experimental data
• Parameters updated on each
10 minutes of data
• 60% accuracy
• Estimate 90% accuracy from
1000 neurons
(Wessberg et al, Nature 408:361-365, 2000)
Dept. Computing
Science & Mathematics
Spring 2010
13
Neural Control of Cursor Movement
• Again, monkeys use joystick to
move onscreen cursor
• Recorded between 7 and 30
motor cortex neurons
• Hand movement predictions
based on neuron firing rates
measured in 50 msec time bins
• Linear models with parameters
based on 3 minutes of data
• Once model was trained the
joystick was disconnected from
the computer…
– Performance maintained
– Sometimes monkeys stopped
making hand movements!
(Serruya et al, Nature 416:141-142, 2002)
Dept. Computing
Science & Mathematics
Spring 2010
14
7