1

Spiking neural networks,

an introduction

Jilles Vreeken

Adaptive Intelligence Laboratory, Intelligent Systems Group,
Institute for Information and Computing Sciences, Utrecht University
Correspondence e-mail address: jvreeken@cs.uu.nl


Biological neurons use short and sudden increases in voltage to send information. These signals are
more commonly known as action potentials, spikes or pulses. Recent neurological research has
shown that neurons encode information in the timing of single spikes, and not only just in their average
firing frequency. This paper gives an introduction to spiking neural networks, some biological
background, and will present two models of spiking neurons that employ pulse coding. Networks of
spiking neurons are more powerful than their non-spiking predecessors as they can encode temporal
information in their signals, but therefore do also need different and biologically more plausible rules
for synaptic plasticity.


You constantly receive sensory input from your environ-
ment. You process this information, recognizing food or dan-
ger, and take appropriate actions. Not only you; anything
that interacts with its environment needs to do so. Mimicking
such a seemingly simple mechanism in a robot proofs to be
insanely difficult. Nature must laugh at our feeble attempts;
animals perform this behaviour with apparent ease.
The reason for this mind-boggling performance lies in
their neural structure or brain. Millions and millions of
neurons are interconnected with each other and cooperate to
efficiently process incoming signals and decide on actions. A
typical neuron sends its signals out to over 10.000 other neu-
rons, making it clear to even to inexpert reader that the signal
flow is rather complicated. To put it mildly: we do not under-
stand the brain that well yet. In fact, we do not even com-
pletely understand the functioning of a single neuron. The
chemical activity of the synapse already proves to be infi-
nitely more complex than firstly assumed.
However, the rough concept of how neurons work is un-
derstood: neurons send out short pulses of electrical energy
as signals, if they have received enough of these themselves.
This basically simple mechanism has been moulded into a
mathematical model for computer use. Artificial as these
computerised neurons are, we refer to them as networks of
artificial neurons, or artificial neural networks. We will
sketch a short history of these now; the biological back-
ground of the real neuron will be drawn in the next chapter.

Generations of artificial neurons
Artificial neural networks are already becoming a fairly old
technique within computer science; the first ideas and mod-
els are over fifty years old. The first generation of artificial
neural networks consisted of McCulloch-Pitts threshold neu-
rons [15], a conceptually very simple model: a neuron sends
a binary high signal if the sum of its weighted incoming
signals rises above a threshold value. Even though these
neurons can only give digital output, they have been success-
fully applied in powerful artificial neural networks like
multi-layer perceptrons and Hopfield nets. For example, any
function with Boolean output can be computed by a multi-
layer perceptron with a single hidden layer; these networks
are called universal for digital computations.
Neurons of the second generation do not use a step- or
threshold function to compute their output signals, but a
continuous activation function, making them suitable for
analog in- and output. Commonly used examples of activa-
tion functions are the sigmoid and hyperbolic tangent. Typi-
cal examples of neural networks consisting of neurons of
these types are feed-forward and recurrent neural networks.
These are more powerful than their first generation predeces-
sors: when equipped with a threshold function at the output
layer of the network they are universal for digital computa-
tions, and do so with fewer neurons than a network of the
first generation [14]. In addition they can approximate any
analog function arbitrarily well, making these networks uni-
versal for analog computations.
Neuron models of the first two generations do not employ
individual pulses, but their output signals typically lie be-
tween 0 and 1. These signals can be seen as normalized firing
rates (frequencies) of the neuron within a certain period of
time. This is a so-called rate coding, where a higher rate of
firing correlates with a higher output signal. Rate coding
implies an averaging mechanism, as real spikes work binary:
spike, or no spike, there is no intermediate. Due to such an
averaging window mechanism the output value of a neuron
can be calculated in iteration. After such a cycle for each neu-
ron the answer of the network to the input values is known.
Real neurons have a base firing-rate (an intermediate fre-
quency of pulsing) and continuous activation functions can
model these intermediate output frequencies. Hence, neu-
rons of the second generation are more biologically realistic
and powerful than neurons of the first generation [3].

2
Adjusting the synaptic weights can alter the information
flow through a neural network; the strengths of incoming
signals of a neuron are altered, so most likely the output
signal will also change in strength. This is a simplified basis
for learning, also known as synaptic plasticity. Using a con-
tinuous activation function allows us to apply a gradient-de-
scent learning algorithm like backward propagation [12].
This is a very commonly used and powerful supervised
learning algorithm for training a network to give the desired
output for a certain input vector.
The third generation of neural networks once again raises
the level of biological realism by using individual spikes.
This allows incorporating spatial-temporal information in
communication and computation, like real neurons do [7]. So
instead of using rate coding these neurons use pulse coding;
mechanisms where neurons receive and do send out individ-
ual pulses, allowing multiplexing of information as fre-
quency and amplitude of sound [9]. Recent discoveries in the
field of neurology have shown that neurons in the cortex
perform analog computations at incredible speed. Thorpe et
al [24] demonstrated that humans analyse and classify visual
input (i.e. facial recognition) in under 100ms. It takes at least
10 synaptic steps from the retina to the temporal lobe; this
leaves about 10ms of processing-time per neuron. Such a
time-window is much too little to allow an averaging mecha-
nism like rate coding [9,24]. This does not mean that rate
coding is not used, though when speed is an issue pulse cod-
ing schemes are favoured [24]. Before going into more detail
about artificial spiking neurons, we will treat some more on
the biological background of real neurons.

Biological background
Maass [19] correctly points out that its a bad idea to pour
water over your computer. It would most likely stop
functioning, defining a sharp contrast with the absolute need
for water all organisms in nature have. The neurons (see fig.
1a) in our brain find themselves surrounded by an artificial
ocean of salty extra-cellular fluid. Our wetware, as we call
our brain and parts of it, bears a lot of resemblance with the
wetware of creatures that still live in the ocean. Squid have
neurons up to 1.000 times larger than we do, making them
much easier to examine. Despite the huge difference in size
their functioning is alike; the equations Hodgkin and Huxley
derived for squid neuron dynamics can also be used to
describe the neurons in our brain. These similarities are
extremely helpful in the research of how neurons, and the
brain in general, function.
Computers communicate with bits; neurons use spikes. In-
coming signals alter the voltage of the neuron and when this
reaches above a threshold-value the neuron sends out an
action potential itself. Such an action potential is a short
(1ms) and sudden increase in voltage that is created in the
cell body or soma. Due to their form and nature (see fig. 1b)
we refer to them as spikes or pulses. The spike traverses
down the axon of the neuron, axons being signal carriers that
grow quite long before they start to branch: we have up to 4
kilometres of them in every cubic millimetre of our cortex.
Bodies of Ranvier amplify the spike over the course of the
axon, and at most branching points the spike is duplicated,
so that minimal information loss occurs [4,10,15,19].
Spikes cannot just cross the gap between one neuron and
the other. They have to be handled by the most complicated
part of the neuron: the synapse [18,19], formed by the end of
the axon, a synaptic gap and the first part of the dendrite.
The synapse was first thought to only just transfer a signal
from axon to the dendrite; it has proved to be a very compli-
cated signal pre-processor and is crucial in learning and
adaptation. When a spike arrives at the axonal (presynaptic)
side of the synapse it is likely that some vesicles fuse with the
cell membrane and release their neurotransmitter content
into the extra-cellular fluid that fills the synaptic gap. Neuro-
transmitter molecules have to reach a matching receptor on
the postsynaptic side of the gap to open an ion-gate on the
neuron. Such a postsynaptic potential (see fig. 1c) can either
be positive and called excitatory (EPSP) or negative and
called inhibitory (IPSP). Once these charged particles enter
the neuron they initiate a cascade that traverses the dendritic
tree down to the trigger zone of the soma, altering the mem-
brane potential. A single neuron receives potentials from
roughly 10.000 synapses [19]. When the sum of these poten-
tials reaches a threshold value the neuron sends out a spike
down the axon. After which the neuron enters a short mo-
ment (10ms) of rest, the refractory period, in which it cannot
send out a spike again.
Contrary to spikes, which are all very much alike, postsy-
naptic potentials differ in size. This is caused by the differ-


Figure 1. (a) Schematic drawing of a neuron. (b) Incoming
postsynaptic potentials alter the membrane voltage so it crosses
threshold value ; the neuron spikes and goes into a refractory state.
(c) Typical forms of excitatory and inhibitory postsynaptic potentials
over time. [8]


Figure 2. A 4 second recording of the neural activity recording from
30 neurons of the visual cortex of a monkey. Each vertical bar
indicates a spike. The human brain can recognize a face within 150ms
[24], which correlates to less than 3mm in this diagram; dramatic
changes in firing frequency occur in this time span, neurons have to
rely on information carried by solitary spikes. [13]


3
ences in amounts and types of neurotransmitters released
and the resulting number of ion-channels activated, in short,
the synaptic efficacy [8,10,19]. The long- and short-term his-
tory of the synapse and outside influences shape the role of a
synapse as a pre-processor. The history of the synapse influ-
ences its characteristics and capabilities in the form of i.e.
chance of vesicle deployment, regeneration and the amount
of receptors. Neuro-hormones in the extra-cellular fluid can
influence both the pre- and postsynaptic terminals temporar-
ily by i.e. enhancing vesicle regeneration or blocking
transmitters from activating ion-gate receptors. These are all
examples of synaptic plasticity: influences on the effect of an
incoming presynaptic spike on the postsynaptic neuron,
which forms the basis of most models of learning and devel-
opment of neural networks.
Now we know a bit more of what is going on under the
hood of a neuron, we can take another look at figure 2. We
see that individual neurons send out erratic sequences of
spikes, or spike-trains, which alter dramatically in frequency
over a short period of time. Neurons have to use spatial and
temporal information of incoming spike patterns to encode
their message to other neurons [9,15,16]. Besides that figure 2
faintly resembles a musical score there are more parallels: in
order to recognize a piece of music you have to hear more
than just a single note, the melody and notes played by other
musicians are even more important [19].

Spike coding
There are many different schemes for the use of spike timing
information in neural computation. Because of the nature of
this paper well only cover two models here: the spike re-
sponse and the integrate-and-fire model. Both are instances
of the general threshold-fire model. The integrate-and-fire
model, which is very commonly used in networks of spiking
neurons, will be covered after the conceptually more simple
and general spike-response model. This model is simple to
understand and implement. However, as it approximates the
very detailed Hodgkin-Huxley model very well it captures
generic properties of neural activity [8,9].
As weve seen in the previous chapter action potentials are
all very similar. We can therefore forget about form and char-
acterise these by their firing times ti
(f)
. The lower index i indi-
cates the neuron, the upper index f the number of the spike.
We can then describe the spike-train of a neuron as
} ,..., {
) ( ) 1 ( n
i
t t F =

(1.1)
The variable u
i
is used to refer to the membrane potential,
or internal state, of a neuron i. If a neurons membrane po-
tential crosses threshold value from below, it generates a
spike. We add the time of this event to F
i
, defining this set as
} 0 | { > = = (t) u (t) u t F
i i i


(1.2)
When a neuron generates an action potential, the mem-
brane potential suddenly increases, followed by a long last-
ing negative after-potential (see fig. 1b). This sharp rise above
the threshold value makes it is absolutely impossible for the
neuron to generate another spike and is named absolute
refractoriness. The period of relative refractoriness, which we
call the negative spike after-potential (SAP), making it less
likely that the neuron fires again. We can model this absolute
and negative refractoriness with kernel :
s) KH(s)H(
) )H(s
t
s
( n
(s)
abs
abs
abs

=
exp
0

(1.3)

>
=
0 if 0
0 if 1
s
s
(s)

(1.4)
The duration of the absolute refractoriness is set by
abs
,
during which large constant K ensures that the membrane
potential

is vastly above the threshold value. Constant n
0

scales the duration of the negative after-potential. Having a
description of what happens to a neuron when it fires itself
we need one for the effect of incoming postsynaptic poten-
tials.
) H(s
s s
(s)
ij
s
ij
m
ij
ij

(

|
|
.
|

\
|

|
|
.
|

\
|
=

exp exp

(1.5)
In equation 1.5
ij
defines the transmission delay (axons
and dendrites are fast, synapses relatively slow) and 0<s<m
are time constants defining the duration of the effect of the
postsynaptic potential. Kernel by default describes the ef-
fect of an excitatory postsynaptic potential; by using the
negative value, we can model an IPSP from an inhibitory
synapse (see fig. 1c). We use variable wij to model the synap-
tic efficacy or weight; with which we also can model inhibi-
tory connections by using values lower than zero. It should
be noted that real synapses are either excitatory or inhibitory;
we know of no synapses changing effect during lifetime.
Neurons of the second generation work in an iterative,
clock-based manner of digital computers, but can deal with
analog input values; we can quite easily feed input neurons
with digitised values from a dataset or a robot-sensor. Due to
their iterative nature these networks are not very well suited
for temporal tasks; they do not use time in their computation,
whereas spiking neural networks do. However, such values
cannot just be fed into a spiking neuron, in some way well
either have to convert this information into spikes, or have to
employ a method to alter the membrane-potential directly.
A general approach to achieve the latter is to use an extra
function h
ext
to describe the effect of an external influence on
the membrane potential. These functions usually are too
task-specific to be covered in this paper, so this leaves us
with


+ =
i j
(f)
j
j F t
(f)
j ij ij
ext
) t (t w (t) h h(t)

(1.6)
For non-hardware solutions it might proof handier to convert
analog signals into spikes that can be fed to the network di-
rectly. An often-used solution is to apply a Poisson-process
for spike generation by the sensor neuron; a higher input
signal correlates with a higher chance for a spike. Such a
spike will then be processed and affect the membrane-poten-
tial of neurons normally. The current excitation of a neuron
is described by
h(t) ) t (t n (t) u
i
(f)
i
F t
(f)
i i i
+ =

(1.7)
where the refractory state, effects of incoming postsynaptic
potentials and eventual external events are combined. To-
gether with equation 1.2 this forms the spike-response
model, a powerful though easy to implement model for
working with spiking neural networks.

Short-term memory neurons
Analysis of neural networks has always been difficult and is
even more so in a spatial-temporal domain as with networks
of spiking neurons. An often-used simplification of the spike-
response model only takes the refractory effects of the last
pulse sent into account. Mathematically speaking, by replac-
ing equation 1.7 with
h(t) ) t (t n (t) u
i i i
+ =
)

(1.8)
we are already finished. Forgetting about the effects of earlier
refractory periods is not a capital crime; for normal operation

4
the model is still quite realistic, while analysis is been made
easier. Due to the bad memory of this model these are
called short term memory neurons.

Integrate-and-fire neurons
The most widely used and best-known model of threshold-
fire neurons, and spiking neurons in general, is the integrate-
and-fire neuron [5,6]. This model is based on, and most easily
explained by, principles of electronics. Figure 3 shows sche-
matic drawings of both a real and an integrate-and-fire neu-
ron. A spike travels down the axon and is transformed by a
low-pass filter, which converts the short pulse into a current
pulse I(t-t
j
(f)
) that charges the integrate-and-fire circuit. The
resulting increase in voltage there can be seen as postsynap-
tic potential (t-t
j
(f)
). Once the voltage over the capacitor goes
above threshold value the neuron sends out a pulse itself.
Mathematically we write
RI(t) u(t)
m
u

m
+ =

(1.9)
to describe the effects on membrane potential u over time,
with
m
being the membrane time constant in which voltage
leaks away. As with the spike-response model the neuron
fires once u crosses threshold and a short pulse is gener-
ated. To force a refractory period after firing we set u to K<0
for a period of
abs
.


=
i j
(f)
j
j F t
(f)
j ij i
) t (t c (t) I

(1.10)
The input current I for neuron i will often be 0, as incom-
ing pulses have a finite short length. Once a spike arrives, it
is multiplied by synaptic efficacy factor c
ij forming the post-
synaptic potential that charges the capacitor. This model is
computationally simple and can easily be implemented in
hardware. It is closely linked to the more general spike-re-
sponse model and can be used like it by rewriting it into the
correct kernels and [8].

Spiking neurons in hardware
Very Large-Scale Integration (VLSI) technology integrates
many powerful features into a small microchip like a micro-
processor. Such systems can use data representations of ei-
ther binary (digital VLSI) or continuous (analog VLSI) volt-
ages. Progress in digital technology has been tremendous,
providing us with ever faster, more precise and smaller
equipment. In digital systems an energy-hungry
synchronisation clock makes it certain that parts are ready
for action. Analog systems consume much less power and
space on silicon than digital systems (in many orders of
magnitude) and are easily interfaced with the analog real
world. However, their design is hard, due to noise computa-
tion is fundamentally (slightly) inaccurate and sufficiently
reliable non-volatile analog memory does not (yet) exist
[20,4].
Noise is the influence of random effects that affect every-
thing in the real world that operates in normal (so, above the
absolute zero) working environment temperatures. For digi-
tal systems this is not much of a problem, as extra precision
can be acquired by using more bits for more precise data
encoding. In analog systems such a simple counter-measure
is not at hand, there are no practical ways of eliminating
noise; at normal temperatures noise has to be accepted as a
fact of life. Our brain is a perfect example of an analog sys-
tem that operates quite well with noise, like neural networks
do in general.
In fact, performance of neural networks increases with
noise present [11]. Spiking neuron models can easily be
equipped with noise-models like noisy threshold, reset or
integration. The interested reader can find more de-tails on
the modelling of noise in spiking neurons in Gerstners
excellent review on neuron models [8].
Hybrid systems can provide a possibly perfect solution,
operating with reliable digital communication and memory
while using fast, reliable and cheap analog computation and
interfacing. In such a solution, neurons can send short digi-
tal pulses, much like weve seen before in the integrate-and-
fire model. This model can be implemented in VLSI systems
quite well [2]. VLSI systems usually work parallel, a very
welcome fact for simulation of neural systems, which are
inherently massively parallel. A significant speed gain can be
acquired by using a continuous hardware solution; by defini-
tion digital simulation will have to recalculate each time-slice
iteratively [20]. Though computer simulations have an
advantage in adaptability, scaling a network up to more neu-
rons (1000+) often means leaving the domain of real-time
simulation. VLSI systems can be specifically designed to be
able to link up, easily forming a scalable set-up that consists
of many parts operating like they are one big system
[2,11,20].

Synaptic plasticity
We saw that synapses are very complex signal pre-processors
that they play an important role in development, memory
and learning of neural structures. Synaptic plasticity is a
form of change of the pre-processing, which is a preferred
word for learning as it better describes what is at hand: long-
or short-term change in synaptic efficacy [1,4,24].
Hebbian plasticity is a local form of long-term potentiation
(LTP) and depression (LTD) of synapses and is based on the
correlation of firing activity between pre- and postsynaptic
neurons. This is usually, and easily, implemented with rate
coding; similar neuron activity means a strong correlation.
As we are using pulse-coding schemes, we have to think
about how to define correlations in neural activity using
single spikes. Pure Hebbian plasticity acts locally at each
individual synapse, making it both very powerful and diffi-
cult to control; it is a positive-feedback process that can
destabilize postsynaptic firing rates by endlessly strengthen-
ing effective and weakening ineffective synapses. If possible
one has to avoid such behaviour, most desirably by a biologi-
cally plausible local rule.
Spike-timing dependent synaptic plasticity (STDP) is a
form of competitive Hebbian learning that uses the exact
spike timing information [1,23]. Experiments in neuroscience
have shown that long-term strengthening occurs if presynap-
tic action potentials arrive within 50ms before of a
postsynaptic spike and a weakening if it arrives late. Due to
this mechanism STDP can lead to stable distributions of LTP
and LDP, making postsynaptic neurons sensitive to the tim-


Fig. 3. Schematic drawing of the integrate-and-fire neuron. On the
left side, the low-pass filter that transforms a spike to a current pulse
I(t) that charges the capacitor. On the right, the schematic version of
the soma, which generates a spike when voltage u over the capacitor
crosses threshold [10].

5
ing of incoming action potentials. This sensitivity leads to
competition among the presynaptic neurons, resulting in
shorter latencies, spike synchronization and faster informa-
tion propagation through the network [1,23].
Hebbian plasticity is a form of unsupervised learning,
which is useful for clustering input data but less appropriate
when a desired outcome for the network is known in ad-
vance. Back-propagation [21] is a widely known and often
used supervised learning algorithm. Due to the very complex
spatial-temporal dynamics and continuous operation it can-
not be directly applied to spiking neural networks, adapta-
tions [12] exist in which individual spikes and their timing
are taken into account.

Discussion
Neural structures are very well suited for complex informa-
tion processing. Animals show an incredible ease in coping
with dynamic environments, raising interest for the use of
artificial neural networks in tasks that deal with real-world
interactions. Over the years, three generations of artificial
neural networks have been proposed, each generation
biologically more realistic and computationally more power-
ful. Spiking neural networks use the element of time in com-
municating by sending out individual pulses. Spiking neu-
rons can therefore multiplex information into a single stream
of signals, like the frequency and amplitude of sound in the
auditory system [9].
We have covered the very general and realistic spike-re-
sponse model as well as the more common integrate-and-fire
model for using pulse coding in neurons. Both models are
powerful, realistic and easy to implement in both computer
simulation and hardware VLSI systems. Standard neural net-
work training algorithms use rate coding and cannot be di-
rectly used satisfactory for spiking neural networks. Spike-
timing dependent synaptic plasticity uses exact spike timing
to optimise information-flow through the network, as well as
impose competition between neurons in the process of unsu-
pervised Hebbian learning.
Pulse coding is computationally powerful [15,16,17] and
very promising for tasks in which temporal information
needs to be processed. We conclude with the remark that this
is the case for virtually any task or application that deals with
in- or output from the real world.






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