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Custom VLSI implementations of neural networks

Early attempts
The idea of making custom analog VLSI implementations of neural networks
dates back to the late 80s - early 90s:

Analog VLSI circuits


for spiking neural networks

[Holler et al. 1989, Satyanarayana et al. 1992, Hammerstrom 1993, Vittoz 1996]

Giacomo Indiveri
Neuromorphic Cognitive Systems group
Institute of Neuroinformatics
University of Zurich and ETH Zurich

General purpose computing

Competing with Intel steamroller

Full-custom analog
implementation

Communication - bandwidth
limited

Neural network accelerator


PC-boards

Difficult to program

Current research

September 17, 2009

Technological progress

Application-specific focus

Power-dissipation/computational
power
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Silicon neuron designs

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Embedded system integration

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Silicon neural network characteristics

Many VLSI models of spiking neurons have been developed in the past, and
many are still being actively investigated:

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Most designs can be traced back to one of two types of silicon neurons,

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Above threshold (strong inversion)

Below threshold (weak inversion)

Mixed analog/digital

Fully analog

Rate-based

Spiking

Real-time

Accelerated-time

Conductance-based

Integrate-and-Fire

Large-scale, event-based
networks

Small-scale, hard-wired

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Why subthreshold neuromorphic VLSI

MOSFETs in subthreshold

Vd

Exploit the physics of silicon to reproduce the


bio-physics of neural systems.

Ids

Vg

n-FET subthreshold transfer function

Ids = I0 en Vg /UT eVs /UT eVd /UT

Vs
where
I0 denotes the nFET current-scaling parameter

n denotes the nFET subthreshold slope factor


UT the thermal voltage
Vg the gate voltage, Vs the source voltage, and Vd the drain voltage.
The current is defined to be positive if it flows from the drain to the source
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Diffusion and saturation

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Exponential voltage dependence


Subthreshold n-FET

Vg

Ids = I0 en Vg /UT eVs /UT eVd /UT

Vd
Ir

-3

-4

10
V

Vs
Ir

Qs

Ids =

Vd

Ids =

If
Qd

subthreshold

10

is equivalent to:

If
Vg

-2

10

I0 e

Ug UVs
T

If

I0 e

-5

U g Ud
T

10

T
-6

10

Ir

If Vds > 4UT the Ir term becomes negligible,


and the transistor is said to operate in the
saturation regime:

Ids (A)

Vs

above threshold

-7

10

-8

10

-9

10

-10

10

-11

10

Ids = I0 en Vg /UT Vs /UT

-12

10

0.5

1.5

2.5

3.5

4.5

Vgs (V)

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n-FETs and p-FETs

One, two, and three transistor circuits

In Complementary Metal-Oxide Semiconductor (CMOS) technology, there are


two types of MOSFETs: n-FETs and p-FETs

Vd

Ideal current source

Vs

Vg

Vg

I out

I in

Vin

Vb

Current-mirror

I out
M1

M2

Vd

Vd

Vs

Inverting amplifier

In traditional CMOS circuits, all n-FETs have the common bulk potential (Vb )
connected to Ground (Gnd), and all p-FETs have a common bulk potential
(typically) connected to the power supply rail (Vdd ).
The corresponding (complementary) equation for the p-FET is

Differential pair
I1
V1

Vin

I2

M2

Vout
Vbn

Ids = I0 e

p (Vdd Vg )/UT

G.Indiveri (NCS @ INI)

(Vdd Vs )/UT

(Vdd Vd )/UT

I2 = I0 e

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I1

V1 Vs

G.Indiveri (NCS @ INI)

V1

V2 Vs

M2

UT

Ib

Vbn

Vs

V2

M3

Vdd

Vdd

M4

M5

M1

V1

V2

x 10

e UT + e UT

V1

I2

M1

I1

M2

I1, I2 (A)

V1

e UT

V2

V1

UT

+e

V2
UT

V2

2UT

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0.2

0.1

0
V1V2 (V)

0.1

0.2

(V1 V2 )

Ib
2UT

is a tunable conductance.

0.3

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where

0.4

0
0.3

e UT + e UT
G.Indiveri (NCS @ INI)

Iout gm (V1 V2 )

0.2

e UT
V1

M3

Iout V
out

gm =

V1

I2 = Ib

Vb

In the linear region (|V1 V2 | < 200mV ):

Ib

0.6

V1

Iout = Ib tanh

Vout

I2
Vs

V2

0.8

I1 = Ib

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Vb

Iout
I1

I0

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I2

UT

Vb

UVs

M1

The transconductance amplifier

Ib = I1 + I2 = I0 e UT
e

V2

The differential-pair

I1 = I0 e

M3

Vs

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What is a synapse?

Synapses in the nervous system

In 1897 Charles Sherrington


introduced the term synapse to
describe the specialized
structure at the zone of contact
between neurons as the point
in which one neuron
communicates with another.

Eex (Na+, ...)


Glutammate

Electrical | Chemical
Excitatory | Inhibitory

Vmem

Depressing | Facilitating
Gl

Cmem

AMPA | NMDA

GABA

...

Einh (K+, Cl, ...)


2005 winner of the Science and Engineering
Visualization Challenge.
by G. Johnson, Medical Media, Boulder, CO.

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Synaptic transmission

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EPSC and EPSP


730

O. SACCHI, M. L. ROSSI, R. CANELLA, AND R. FESCE

In chemical synapses the presynaptic and


postsynaptic membranes are spearated by
extracellular space.
The arrival of a presynaptic action potential
triggers the release of neurotransmitter in the
extracellular space.
The neurotransmitters react with the
postsynaptic receptors and depolarize the cell.

FIG .

2.

Typical synaptic currents at ganglionic synapse evoked by supra-

low the linear fit. This might be partly because of inward


rectification by the ACh-evoked current (Fieber and Adams
1991; Mathie et al. 1990), but it is at least partly accounted
for by the changes in ionic gradients that are associated with
even small ionic flows (especially accumulation of K / ions
in the perineuronal space) (Belluzzi and Sacchi 1990).
In all neurons the decay of EPSC was well fit by a single
exponential function. The average time constant, t, was 7.5
ms at 075 mV (n 14) and was scarcely voltage-dependent:
it decreased exponentially with a constant of 260.4 mV from
0105 to 025 mV (Fig. 3A). EPSC amplitudes decreased in
the average (5 neurons) to 69% of the initial value when
Ca 2/ concentration in the bath was lowered from the usual
5 to 2 mM; the current decay time constants were also somewhat decreased (by 21% in the average). The voltage dependence of the decay time constant and the estimated reversal
potential did not appear to be affected by calcium concentration (Fig. 3B).

maximal preganglionic stimulation.


Superimposed EPSCs recorded in a
Superimposed
excitatory
sympathetic neuron at different membrane potentials between 010 and
NATURE OF THE POSTSYNAPTIC RECEPTOR ACTIVATED BY
0100 mV (in 10 mV steps) under 2-electrode voltage-clamp conditions.
post-synaptic
currents (EPSCs)
Excitatory
post-synaptic
AC . To
correctly model andpotential
reproduce evoked
Cell was held at 050 mV and then repeatedly stepped, at 20 s intervals, to NATURAL
membrane levels at which synaptic current was observed. Note presence synaptic currents, it was important to determine whether or
recorded
in0a
neuron
at different
(EPSP)
in response
multiple
30 mV
tracings. External
Ca concentration was 5 not
of fast I in 010/
conductances
other than thoseto
associated
to the nicotinic
mM.
receptor were activated by nerve-released ACh. In fact, ACh
membrane potentials (from Sacchi et
pre-synaptic
spikes
al.
release
by the presynaptic
nerve(from
terminalsNicholls
is thought toet
gencell was estimated for the rat sympathetic neuron in culture, erate in the ganglion a broad series of side effects, in addition
al.,
1998).
1992).
whereas
the activation-deactivation time constants for putative M- to
the central role of sustaining the fast excitatory transmisH

Chemical synaptic transmission is


characterized by specific temporal dynamics.

2/

Na

currents were 220 ms in the 060/ 030 mV voltage range (Owen


et al. 1990).

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sion. Autoreceptors were proposed to control evoked transmitter release (see Starke et al. 1989, for a review). The
evidence for a physiological role in the ganglion seems to
R E S U L G.Indiveri
TS
greatest for the muscarinic autoreceptors, which are pos(NCS @ INI)
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tulated to mediate a negative feedback loop that reduces
Synaptic current at the sympathetic ganglionic neuron
transmitter mobilization (Koelle 1961; Koketsu and Yamada

Neural network models

VLSI synapses in classical neural networks


The role of the VLSI synapse in implementations of classical neural network
models is that of a multiplier.
Multiplying synaptic circuits have been implemented using a wide range of
analog circuits, ranging from the single MOS-FETs to the Gilbert multiplier.
Iin1

I1

Iin2

I2

In classical neural network theory


signals are (tipically) continuous values that represent the neurons mean
firing rate,

Ib

neurons implement a saturating non-linearity transfer function (S) on the


inputs weighted sum,
Figure: Schematic of half of a Gilbert multiplier. This circuit multiplies Iin1 and Iin2 by Ib
if Iin1 + Iin2 = Ib .

the synapse implements a multiplication between the neurons input


signal (Xi ) and its corresponding synaptic weight (wi ).
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VLSI synapses in pulse-based neural networks

IWi

Vi =
Vmem

IWi
Cmem

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A linear integrator is a linear low-pass filter. Its impulse response should be a


decaying exponential.
With VLSI and subthreshold MOSFETS its fairly easy to implement
exponential voltage to current conversion, and linear voltage increase or
decrease over time.

Cmem

Vdd

In pulse-based neural networks the weighted contribution of a synapse can be


implemented using a single transistor.
In this case p-FETs implement excitatory synapse, and n-FETs implement
inhibitory synapses.
The synaptic weight can be set by changing the Wi bias voltage or the t
duration.

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Linear pulse integrators

Vdd

Wi

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Vg

Vdd

Id
Vg(t)

Vc

Id =

d
C dt
Vc

Id (t ) = I0 e UT

(Vdd Vg (t ))

Id(t)

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Linear charge-and-discharge integrator

Iw = I0 e
M

Csyn

I
Vw

Vsyn

I = I0 e
Msyn

Mw

Ic = C

Isyn

Iw

Vw
UT

, c ,

(Vdd V )

d
dt

UT

Isyn (t ) =

Csyn UT

Iw = I0 e UT

Csyn UT

, d ,

M
I

I
Vw

(Vdd Vsyn )

Isyn = I0 e

Mpre

Log-domain pulse integrator

Csyn

I = I0 e

Vsyn

Msyn

Mw

UT

Mpre

(t t )

I e+ ci

dt

(charge phase)

syn

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The diff-pair integrator (DPI)

d
dt
d

Vsyn

Msyn

Iin
Vthr

Mthr

Isyn

Min

I = I0 e
Ic = C

UT

(Vdd V )

d
dt

(Vdd Vsyn )

Isyn = I0 e
d
dt

d
dt

Isyn + Isyn

+e

UT

Vthr

Iout (ti +t ) =

UT

(Vdd Vsyn )

Isyn =

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(Vw Vdd )
UT

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Iout

I 1 +

dt

Iout + Iout

Ig

Iin ,

Iout
Ig

 Iin

if Iout  Ig , Iw  I

Isyn

d
dt


ti ti 1
t
t
Ig Iin 
1 e + Iout (ti )e , Iout (ti ) = Iout (ti 1 +t )e
I

Response to an arbitrary spike train (t ) = i (t ti ), with t  :




Z t

Ig Iin
t
e
e ( )d
Iout (t ) =
I
0

UT

UT

Vsyn

Iw Igain

:
Mean response to spike train of mean frequency


< Iout >=

(Bartolozzi, Indiveri, 2007)

G.Indiveri (NCS @ INI)

Vsyn

Response to t pulse at time ti :

UT

Vsyn

Iw
Mpre

UT dt

Csyn UT

, Iw0 = I0 e

UT

Iin = Iw

Mw

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DPI transfer function:


d
Iout + Iout
dt

Vw

Vsyn

Vw

G.Indiveri (NCS @ INI)

I0 Iw0

UT

Isyn Iw

Isyn + Isyn =

Isyn + Isyn =

(Vdd Vsyn )

DPI equations
Iw = I0 e

Csyn

(Vdd Vsyn )

Isyn = Isyn

(t t + )

+ di
I e

dt

dt

Isyn = I0 e

(discharge phase)


n
c +d )
c
1
c f ct (
t

d
Isyn (t ) = I0 e
, with f =
, f<
t
c + d t

UT

Isyn
Iw

(Vdd Vsyn )

syn

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(Vdd V )

Ic = C

(Vsyn Vw )

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G.Indiveri (NCS @ INI)

Ig Iin
I


t , =

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t + ISI
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DPI measured response

DPI response to spike-trains

300

200

400

Vw=300mV

Vw=440mV

350

Vw=320mV

Vw=460mV

300

Vw=340mV

EPSC (nA)

150
100

250
200
150
100

50

50
0
0

0.05

0.1
Time (s)

0.15

0
0

0.5

1.5

2
2.5
Time (s)

3.5

100
80
60
40
20
0

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Proc. Natl. Acad. Sci. USA 94 (1997)


Short-term depression

50

100
150
Input Frequency (Hz)

G.Indiveri (NCS @ INI)

200

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Short-term depression

Va

Vgain
Va

Vw

M6

M1

M7
0.35

Id

Slow recovery

C2

Vx

0.3

M5
Ir

Vd

Ir

Cd

Vd

M2

C
Vpre

Vd=0.26 V

0.25 Update

Isyn

Vx (V)

EPSC (nA)

250

Vw=420mV

Output Frequency (Hz)

450

l
e
e
f
e
h

M4

Vd=0.28 V

Fast recovery

0.2
Vd=0.3 V

0.15

Vpre

e
n

M3

0.1
0.04

(a)

P,

V =1.30V
w
V =1.33V
w
V =1.35V

120

(C. Rasche, R. Hahnloser, 2001)

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0.06

0.08

0.1
0.12
Time (s)

0.14

0.16

(b)

Figure 1: Schematic for a depressing synapse circuit and responses to a regular input spike
(M. Boegerhausen, P. Suter, and S.-C. Liu 2003)
train. (a) Depressing synapse circuit. The voltage @A determines
the synaptic conductance
D

;
C
E

while the synaptic term
or B
is exponential in the voltage, @GF . The subcircuit
DC;E . The presynaptic
5 ( , 5IH , and 5K
J , control
consisting
of transistors,
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(NCS @ INI)
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5
J
input goes to the gate terminal of
which acts like a switch. When there is a presynaptic

STDP and beyond

Spike-driven learning in VLSI I


Alternative spike-driven learning algorithm
Spike-driven weight change depends on the value of
the post-synaptic neurons membrane potential, and on
its recent spiking activity.

J. Arthur and K. Boahen.


Learning in silicon: Timing is everything.
In Y. Weiss, B. Schlkopf, and J. Platt, editors, Advances in Neural Information Processing Systems 18. MIT Press, Cambridge, MA,
2006.
A. Bofill-i Petit and A. F. Murray.
Synchrony detection and amplification by silicon neurons with STDP synapses.
IEEE Transactions on Neural Networks, 15(5):12961304, September 2004.

Fusi et al. 2000; Brader et al. 2007

E. Chicca, D. Badoni, V. Dante, M. DAndreagiovanni, G. Salina, S. Fusi, and P. Del Giudice.


A VLSI recurrent network of integrateandfire neurons connected by plastic synapses with long term memory.
IEEE Transactions on Neural Networks, 14(5):12971307, September 2003.

Recipe for efficient VLSI implementation


1

bistability: use two synaptic states;

redundancy: implement many synapses that see


the same pre- and post-synaptic activity

P. Hfliger, M. Mahowald, and L. Watts.

stochasticity & inhomogeneity: induce LTP/LTD


only in a subset of stimulated synapses.

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G. Indiveri, E. Chicca, and R. Douglas.


A VLSI array of low-power spiking neurons and bistable synapses with spiketiming dependent plasticity.
IEEE Transactions on Neural Networks, 17(1):211221, Jan 2006.
S. Mitra, G. Indiveri, and S. Fusi.
Learning to classify complex patterns using a VLSI network of spiking neurons.
In J.C. Platt, D. Koller, Y. Singer, and S. Roweis, editors, Advances in Neural Information Processing Systems 20, pages 10091016,
Cambridge (MA), 2008. MIT Press.

- Slow learning: only a fraction of the synapses memorize the pattern.


+ The theory is matched to the technology: use binary states, exploit
mismatch and introduce fault tolerance by design.
G.Indiveri (NCS @ INI)

A spike based learning neuron in analog VLSI.


In M. C. Mozer, M. I. Jordan, and T. Petsche, editors, Advances in neuralinformation processing systems, volume 9, pages 692698.
MIT Press, 1997.

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Neurons . . . in a nutshell

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Neurons of the world

A quick tutorial

Complexity

Real Neurons
Conductance based models
Integrate and fire models
Rate based models
I
I

Sigmoidal units
Linear threshold units

(adapted from B. Mel, 1994)


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G.Indiveri (NCS @ INI)

Equivalent Circuit

Spike generating mechanism


Eex (Na+, ...)

ENa

Glutammate

gNa
Vmem

Vmem
Gl

Cmem

Einh (K+, Cl, ...)

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Cmem

EK

If excitatory input currents are relatively small, the neuron behaves exactly like
a first order low-pass filter.

G.Indiveri (NCS @ INI)

Gl

gK

GABA

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Spike properties

If the membrane voltage increases above a certain threshold, a


spike-generating mechanism is activated and an action potential is initiated.
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The F-I curve


Iin=I1

Spike Frequency (F)

Refractory
Period

Refractory Period

Pulse Width

Iin=I2 > I1

Input Current (I)


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Hardware implementations of spiking neurons

Conductance-based models of spiking neurons

The first artificial neuron model was proposed in the 1943 by McCulloch and
Pitts. Hardware implementations of this model date almost back to the same
period.

Hardware implementations of spiking neurons are relatively new.

One of the most influential circuits that implements an


integrate and fire (I&F) model of a neuron was the
Axon-Hillock Circuit, proposed by Carver Mead in the late
1980s.

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Conductance based Si-Neurons

In 1991 Misha Mahowald and Rodney Douglas proposed a


conductance-based silicon neuron and showed that it had properties
remarkably similar to those of real cortical neurons.
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Conductance based Si-Neurons


Silicon neurons measurements

Sodium

Vdd
Vdd
Sodium Current

1V

Vm

+
VNa

Vthr

INaoff

+
GNaoff

[Ca]

INa

INaon

Vthr

ENa

Passive Leak

GNaon

Vm
[Ca]

Vmem

Vdd

Eleak

Vdd

Vmem

Gleak

Passive

+
VK

Potassium

Vthr

Cmem

IK

Vm

IK

GK

[Ca]

Potassium Current

EK

EK

I
50 ms
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The Axon-Hillock Circuit

The Axon-Hillock Circuit


voltage

Positive Feedback

Input current
Membrane voltage

Vmem

Vmem

Vout

Vout

Vmem

Vout

Output voltage

Vout

time

Vpw

Reset

Slope = A

Vpw
Vmem

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Capacitive Divider

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Positive Feedback

Given the change V 2, what is V 1?

C2
Q

Q = C1 V1 + C2 (V1 V2 ) = constant
C1 V1 + C2 (V1 V2 ) = 0

C2
C1 + C2

V1

Vmem

A
C1

V2

time

V2

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G.Indiveri (NCS @ INI)

Vout

Vmem

Vout

Cm

Vpw

G.Indiveri (NCS @ INI)

voltage

Cfb

V1 =

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Positive Feedback
Cfb
Vmem =
V
Cm + Cfb dd

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Axon-Hillock Circuit Dynamics


voltage

Cfb
Iin
Vmem

Gain

Vmem

Vout

How to make voltage gain

Vout

Cm
tH

tL

time

Ir
Vpw

tL =

Cfb + Cm
Iin

Vmem =

Frequency Iin
G.Indiveri (NCS @ INI)

Cfb
Iin

Vdd

tH =

Cfb + Cm
Ir Iin

Vmem =

Cfb
Ir Iin

Vdd

Pulse width 1/Ir for Ir  Iin


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Whats bad about this?


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Power Dissipation

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Conductance-based models
Integrate and Fire vs Hodgkin-Huxley

The Axon-Hillock circuit is very compact and allows for


implementations of dense arrays of silicon neurons

Traditionally there have been two main classes of neuron models:

BUT
it has a major drawback: power consumption
During the time when an inverter switches, a large amount
of current flows from Vdd to Gnd.

Integrate and fire (I-C)

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G.Indiveri (NCS @ INI)

Conductance-based (R-C)

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Conductance-based models

An ultra low-power generalized I&F circuit

Integrate and Fire vs Hodgkin-Huxley


PositiveFeedback

Adaptation

But recently proposed models bridge the gap between the two:

M11

M5
Iin

Vahp

Vrest

Vthr

M1

DPI

M7

M12

M6

Ifb

M8

Cmem

Leak
M9

M15

M19

M16

M20

Vspk

Imem
M22

Iahp

Vtau

M18

Vmem

M3

M2

Vthr_ahp

M14

Cahp
M10

M4

M17

DPI
Vtau_ahp

Vrf

M21

M13

RefractoryPeriod

(G. Indiveri, P. Livi, ISCAS 2009)

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Generalized Integrate and Fire models can account for a very large set of
DPI
neuron sub-threshold equations
d
M5
Vahp

dt

M11

Cmem

M12

+ F (umem )

M18

where F (uDPImem ) is a non-linear function of umem (t ).


Vthr

M2

Vthr_ahp

M1

M6

Vmem

M3

Cmem

Imem

Vtau

M9
M4

M15

M19

M16

M20

15

Vspk

Vthr=0.325 V
Vthr=0.350 V
Vthr=0.375 V

14
12
10

M22

Iahp

Leak

20

Ifb

M8

M7

DPI

Cahp
M10

Vrf

M17

Vtau_ahp

M21

Imem (A)

Vrest

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PositiveFeedback
in

M14

M13

10

Imem (A)

Iin

umem =

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SPICE simulations

behaviors captured by far more complicated Hodgkin-Huxley models.


Adaptation

G.Indiveri (NCS @ INI)

8
6
4

RefractoryPeriod

d
dt

G.Indiveri (NCS @ INI)

Imem + Imem
1
I

I0+1 , ,

Ig Iin
I

0
2

+ (Imem )

6
Time (ms)

10

20

40
60
Time (ms)

80

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2 + 1

+1

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Experimental results

Experimental results

Single spike

Population activity

2.8

Vthr= 0.0V
V = 0.3V

Voltage (V)

2.7

Vthr= 0.6V

2.6

Vthr= 0.9V

Neurons

thr

2.5

30

30

25

25

20

20

15

15

10

10

0.5

1.5

0.5

1.5

0.5

1
Time (s)

1.5

2.4
2.3

Neurons

2.2
2.1
2
1.9
0.01
G.Indiveri (NCS @ INI)

0.02

0.03
Time (s)

0.04

0.05

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30

30

25

25

20

20

15

15

10

10

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Spiking multi-neuron architectures

0.5

1
Time (s)

1.5

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Spikes and Address-Event Systems


1

Vmem (V)

0.8

0.6

0.4

Encode

Decode

0.2

0
0

0.05

0.1
Time (s)

0.15

0.2

1
2
3

Address Event Bus


1 23 2

Networks of I&F neurons with adaptation,


refractory period, etc.

Inputs

1
2
3

12

Outputs

Source
Chip

Synpases with realistic temporal dynamics

Destination
Chip
Address-Event
representation of
action potential

Winner-Take-All architectures
Spike-based plasticity mechanisms
Action Potential
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15
10
5

Distributed multi-layer networks

0
180

120

60

60

120

180

Rotation around y axis (degrees)

Figure 1 Tuned units in inferotemporal cortex. A monkey was trained to recognize


a three-dimensional paperclip from all viewpoints (pictured at top). The graph
shows tuning to the multiple parameters characterizing each view summarized in
terms of spike rate versus rotation angle of three neurons in anterior inferotemporal
cortex that are view-tuned for the specific paperclip. (The unit corresponding to the
green tuning curve has two peaks to a view of the object and its mirror view.) A
combination of such view-tuned neurons (Fig. 2) can provide view-invariant, object
specific tuning as found in a small fraction of the recorded neurons. Adapted from
Logothetis et al.13.

AMS 0.35 m
3.9mm 2.5mm
128
28 128
4 128
32 128 | . . . | 1 4096

Technology:
Size:
Neurons:
AER plastic synapses:
AER non-plastic synapses
Dendritic tree multiplexer:

G.Indiveri (NCS @ INI)

ICANN09 Tutorial

Summary
Vg

Ir

Ir

M11

M5
Vahp

Vrest

Vthr

M2

M1

DPI

Vthr_ahp

M7

M6

M9
M4

V4/PIT

V4

V1

V1

Figure 2 A model of visual learning. The model summarizes in quantitative terms


other models and many data about visual recognition in the ventral stream pathway
in cortex. The correspondence between the layers in the model and visual areas is
an oversimplification. Circles represent neurons and arrows represent connections
@ INI)
between them;G.Indiveri
the dots signify (NCS
other neurons
of the same type. Stages of neurons
with bell-shaped tuning (with black arrow inputs), that provide example-based
learning and generalization, are interleaved with stages that perform a max-like
operation3 (denoted by red dashed arrows), which provides invariance to position
and scale. An experimental example of the tuning postulated for the cells in the
layer labelled inferotemporal in the model is shown in Fig. 1. The model accounts
well for the quantitative data measured in view-tuned inferotemporal cortex cells10
(J. Pauls, personal communication) and for other experiments55. Superposition of
gaussian-like units provides generalization to three-dimensional rotations and
together with the soft-max stages some invariance to scale and position. IT,
infratemporal cortex, AIT, anterior IT; PIT, posterior IT; PFC, prefrontal cortex.
Adapted from M. Riesenhuber, personal communication.

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Thank you for your attention


Additional information available at:

769

http://ncs.ethz.ch/

2004 Nature Publishing Group

Vd
If
Qd

M18

M15

Cmem

Leak

IT

Ifb

M8

Imem

M19
Vspk

M16

M20

M22

Iahp

Vtau

AIT

M14

M12

Vmem

M3

Ident.

PositiveFeedback

Adaptation

Iin

Qs

Categ.

If
Vg

Vs

AER INPUT Y

PFC

NATURE | VOL 431 | 14 OCTOBER 2004 | www.nature.com/nature

Vd

Vs

types of algorithm developed within learning theory corresponds to


networks that combine the activities of units, each broadly tuned to
one of the examples (Box 1). Theory (see references in Box 1) shows
that a combination of broadly tuned neurons those that respond
to a variety of stimuli, although at sub-maximal firing rates might
generalize well by interpolating among the examples.
In visual cortex, neurons with a bell-shaped tuning are common.
Circuits in infratemporal cortex and prefrontal cortex, which combine activities of neurons in infratemporal cortex tuned to different
objects (and object parts) with weights learned from experience, may
underlie several recognition tasks, including identification and
categorization. Computer models have shown the plausibility of this
scheme for visual recognition and its quantitative consistency with
many data from physiology and psychophysics25 .
Figure 2 sketches one such quantitative model, and summarizes a
set of basic facts about cortical mechanisms of recognition established
over the last decade by several physiological studies of cortex68. Object
recognition in cortex is thought to be mediated by the ventral visual
pathway running from primary visual cortex, V1, over extrastriate
visual areas V2 and V4 to the inferotemporal cortex. Starting from
simple cells in V1, with small receptive fields that respond preferably to
oriented bars, neurons along the ventral stream show an increase in
78
receptive field size as well as in74
the/complexity
of their preferred stimuli.
At the top of the ventral stream, in the anterior inferotemporal cortex,
neurons respond optimally to complex stimuli such as faces and other
objects. The tuning of the neurons in anterior inferotemporal cortex
probably depends on visual experience919. In addition, some neurons
show specificity for a certain object view or lighting condition13,18,2022.
For example, Logothetis et al.13 trained monkeys to perform an object
recognition task with isolated views of novel three-dimensional objects
(paperclips; Fig. 1). When recording from the animals' inferotemporal
cortex, they found that the great majority of neurons selectively tuned
to the training objects were view-tuned (see Fig. 1) to one of the training
objects. About one tenth of the tuned neurons were view-invariant,
consistent with an earlier computational hypothesis23.

communication

AER OUTPUT

A minimum-size chip implementing a


reconfigurable AER neural network.
Neurons and synapses have realistic
temporal dynamics. Local circuits at
The basic problem with these models is, of course, generalization:
each synapse implement
the bi-stable
a look-up table cannot deal with new events, such as viewing a face
from the side rather than the front, and it cannot learn in the predicspike-based plasticity mechanism.
tive sense described earlier. One of the simplest and most powerful
Indiveri, Fusi, 2007

which several models have been suggested24; see also review in this
issue by Abbott and Regehr, page 796), since it depends only on
passive experience of the visual inputs. However, the weights of the
combination (see Fig. 3) depend on learning the task and require at
least some feedback (see Box 2).
Thus, generalization in the brain can emerge from the linear combination of neurons tuned to an optimal stimulus effectively
defined by multiple dimensions25,23,26. This is a powerful extension of
Analog
processing, asynchronous
digital
the older computation-through-memory
models of vision
and
motor control. The question now is whether the available evidence
supports the existence of a similar architecture underlying generalization in domains other than vision.

AER INPUT X

Spikes per seco

A spike-based learning chip

239

20

M17

DPI

Cahp
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Vtau_ahp

Vrf

M21

M13

RefractoryPeriod

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