Neural Networks and Physical Systems with Emergent Collective Computational Abilities

J. J. Hopfield
PNAS 1982;79;2554-2558
doi:10.1073/pnas.79.8.2554
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Proc. NatL Acad. Sci. USA

Vol. 79, pp. 2554-2558, April 1982

Biophysics

Neural networks and physical systems with emergent collective

computational abilities
(associative memory/parallel processing/categorization/content-addressable memory/fail-soft devices)

J. J. HOPFIELD
Division of Chemistry and Biology, California Institute of Technology, Pasadena, California 91125; and Bell Laboratories, Murray Hill, New Jersey 07974

Contributed by John J. Hopfweld, January 15, 1982

calized content-addressable memory or categorizer using extensive asynchronous parallel processing.

The general content-addressable memory of a physical
system
Suppose that an item stored in memory is "H. A. Kramers &
G. H. Wannier Phys. Rev. 60, 252 (1941)." A general contentaddressable memory would be capable of retrieving this entire
memory item on the basis of sufficient partial information. The
input "& Wannier, (1941)" might suffice. An ideal memory
could deal with errors and retrieve this reference even from the
input "Vannier, (1941)". In computers, only relatively simple
forms of content-addressable memory have been made in hardware (10, 11). Sophisticated ideas like error correction in accessing information are usually introduced as software (10).
There are classes of physical systems whose spontaneous behavior can be used as a form of general (and error-correcting)
content-addressable memory. Consider the time evolution of
a physical system that can be described by a set of general coordinates. A point in state space then represents the instantaneous condition of the system. This state space may be either
continuous or discrete (as in the case of N Ising spins).
The equations of motion ofthe system describe a flow in state
space. Various classes offlow patterns are possible, but the systems of use for memory particularly include those that flow toward locally stable points from anywhere within regions around
those points. A particle with frictional damping moving in a
potential well with two minima exemplifies such a dynamics.
If the flow is not completely deterministic, the description
is more complicated. In the two-well problems above, if the
frictional force is characterized by atemperature, it must also
produce a random driving force. The limit points become small
limiting regions, and the stability becomes not absolute. But
as long as the stochastic effects are small, the essence of local
stable points remains.
Consider a physical system described by many coordinates
X1 XN, the components of a state vector X. Let the system
have locally stable limit points Xa, Xb, **. Then, if the system
is started sufficiently near any Xa, as at X = Xa + A, it will
proceed in time until X Xa. We can regard the information
stored in the system as the vectors Xa, Xb, . The starting
point X = Xa + A represents a partial knowledge of the item
Xa, and the system then generates the total information Xa.
Any physical system whose dynamics in phase space is dominated by a substantial number of locally stable states to which
it is attracted can therefore be regarded as a general contentaddressable memory. The physical system will be a potentially
useful memory if, in addition, any prescribed set of states can
readily be made the stable states of the system.
The model system
The processing devices will be called neurons. Each neuron i
has two states like those of McCullough and Pitts (12): Vi = 0

ABSTRACT Computational properties of use to biological organisms or to the construction of computers can emerge as collective properties of systems -having a large number of simple
equivalent components (or neurons). The physical meaning of content-addressable memory is described by an appropriate phase
space flow of the state of a system. A model of such a system is
given, based on aspects of neurobiology but readily adapted to integrated circuits. The collective properties of this model produce
a content-addressable memory which correctly yields an entire
memory from any subpart of sufficient size. The algorithm for the
time evolution of the state of the system is based on asynchronous
parallel processing. Additional emergent collective properties include some capacity for generalization, familiarity recognition,
categorization, error correction, and time sequence retention.
The collective properties are only weakly sensitive to details of the
modeling or the failure of individual devices.
Given the dynamical electrochemical properties of neurons and
their interconnections (synapses), we readily understand schemes
that use a few neurons to obtain elementary useful biological
behavior (1-3). Our understanding of such simple circuits in
electronics allows us to plan larger and more complex circuits
which are essential to large computers. Because evolution has
no such plan, it becomes relevant to ask whether the ability of
large collections of neurons to perform "computational" tasks
may in part be a spontaneous collective consequence of having
a large number of interacting simple neurons.
In physical systems made from a large number of simple elements, interactions among large numbers of elementary components yield collective phenomena such as the stable magnetic
orientations and domains in a magnetic system or the vortex
patterns in fluid flow. Do analogous collective phenomena in
a system of simple interacting neurons have useful "computational" correlates? For example, are the stability of memories,
the construction of categories of generalization, or time-sequential memory also emergent properties and collective in
origin? This paper examines a new modeling of this old and fundamental question (4-8) and shows that important computational properties spontaneously arise.
All modeling is based on details, and the details of neuroanatomy and neural function are both myriad and incompletely
known (9). In many physical systems, the nature of the emergent collective properties is insensitive to the details inserted
in the model (e.g., collisions are essential to generate sound
waves, but any reasonable interatomic force law will yield appropriate collisions). In the same spirit, I will seek collective
properties that are robust against change in the model details.
The model could be readily implemented by integrated circuit hardware. The conclusions suggest the design of a deloThe publication costs ofthis article were defrayed in part by page charge
payment. This article must therefore be hereby marked "advertisement" in accordance with 18 U. S. C. 1734 solely to indicate this fact.
2554

Biophysics: Hopfield

Proc. Natl. Acad. Sci. USA 79 (1982)

("not firing") and Vi = 1 ("firing at maximum rate"). When neuron i has a connection made to it from neuron j, the strength
of connection is defined as Tij. (Nonconnected neurons have Tij
0.) The instantaneous state of the system is specified by listing
the N values of Vi, so it is represented by a binary word of N
bits.
The state changes in time according to the following algorithm. For each neuron i there is a fixed threshold U,. Each
neuron i readjusts its state randomly in time but with a mean
attempt rate W, setting
Vi

1
I T.,V.
Vi0if
joi

< Ui

Thus, each neuron randomly and asynchronously evaluates

whether it is above or below threshold and readjusts accordingly. (Unless otherwise stated, we choose Ui = 0.)
Although this model has superficial similarities to the Perceptron (13, 14) the essential differences are responsible for the
new results. First, Perceptrons were modeled chiefly with
neural connections in a "forward" direction A -> B -* C -- D.
The analysis of networks with strong backward coupling
proved intractable. All our interesting results arise
as consequences of the strong back-coupling. Second, Perceptron studies usually made a random net ofneurons deal directly
with a real physical world and did not ask the questions essential
to finding the more abstract emergent computational properties. Finally, Perceptron modeling required synchronous neurons like a conventional digital computer. There is no evidence
for such global synchrony and, given the delays of nerve signal
propagation, there would be no way to use global synchrony
effectively. Chiefly computational properties which can exist
in spite of asynchrony have interesting implications in biology.
The information storage algorithm
Suppose we wish to store the set of states V8, s = 1 n. We
use the storage prescription (15, 16)

Tij=

(2V -

1)(2Vj - 1)

[2]

but with Tii = 0. From this definition

Tijjs =E (2V,- 1)

VJ(2Vj-1) Hjs.

computation (20, 21). Complex circuitry is easy to plan but more

difficult to discuss in evolutionary terms. In contrast, our model
obtains its emergent computational properties from simple
properties of many cells rather than circuitry.
The biological interpretation of the model
Most neurons are capable of generating a train of action potentials-propagating pulses of electrochemical activity-when the
average potential across their membrane is held well above its
normal resting value. The mean rate at which action potentials
are generated is a smooth function of the mean membrane potential, having the general form shown in Fig. 1.
The biological information sent to other neurons often lies
in a short-time average of the firing rate (22). When this is so,
one can neglect the details of individual action potentials and
regard Fig. 1 as a smooth input-output relationship. [Parallel
pathways carrying the same information would enhance the
ability of the system to extract a short-term average firing rate
(23, 24).]
A study of emergent collective effects and spontaneous computation must necessarily focus on the nonlinearity of the input-output relationship. The essence of computation is nonlinear logical operations. The particle interactions that produce
true collective effects in particle dynamics come from a nonlinear dependence of forces on positions of the particles. Whereas
linear associative networks have emphasized the linear central
region (14-19) of Fig. 1, we will replace the input-output relationship by the dot-dash step. Those neurons whose operation
is dominantly linear merely provide a pathway of communication between nonlinear neurons. Thus, we consider a network
of "on or off" neurons, granting that some of the interconnections may be by way of neurons operating in the linear regime.
Delays in synaptic transmission (of partially stochastic character) and in the transmission of impulses along axons and dendrites produce a delay between the input of a neuron and the
generation of an effective output. All such delays have been
modeled by a single parameter, the stochastic mean processing
time 1/W.
The input to a particular neuron arises from the current leaks
of the synapses to that neuron, which influence the cell mean
potential. The synapses are activated by arriving action potentials. The input signal to a cell i can be taken to be

[5]

I Tijvj

[3]

The mean value of the bracketed term in Eq. 3 is 0 unless s

- s', for which the mean is N/2. This pseudoorthogonality
yields

2555

where Tij represents the effectiveness of a synapse. Fig. 1 thus

/

>

TiVs (Hs') (2Vs' - 1) N/2

[4]

and is positive if VW' = 1 and negative if Vf' = 0. Except for the

noise coming from the s # s' terms, the stored state would always be stable under our processing algorithm.
Such matrices T,. have been used in theories of linear associative nets (15-19) to produce an output pattern from a paired
input stimulus, S1 -* 01. A second association S2 - 02 can be
simultaneously stored in the same network. But the confusing
simulus 0.6 Si + 0.4 S2 will produce a generally meaningless
mixed output 0.6 01 + 0.4 02 Our model, in contrast, will use
its strong nonlinearity to make choices, produce categories, and
regenerate information and, with high probability, will generate
the output 01 from such a confusing mixed stimulus.
A linear associative net must be connected in a complex way
with an external nonlinear logic processor in order to yield true

~~~~~~~~~/

I'

a)-jaz -Present Model

t
--Linear Modeling
wW
0

.'C

-0.1 /
0
Membrane Potential (Volts) or "Input"

FIG. 1. Firing rate versus membrane voltage for a typical neuron

(solid line), dropping to 0 for large negative potentials and saturating
for positive potentials. The broken lines show approximations used in
modeling.

2556

Proc. NatL Acad. Sci. USA 79 (1982)

Biophysics: Hopfield

becomes an input-output relationship for a neuron.

Little, Shaw, and Roney (8, 25, 26) have developed ideas on
the collective functioning ofneural nets based on "on/off" neurons and synchronous processing. However, in their model the
relative timing of action potential spikes was central and resulted in reverberating action potential trains. Our model and
theirs have limited formal similarity, although there may be
connections at a deeper level.
Most modeling of neural learning networks has been based
on synapses of a general type described by Hebb (27) and Eccles
(28). The essential ingredient is the modification of T; by correlations like
AT41 = [Vi(t)Vj(t)]average

[6]

where the average is some appropriate calculation over past

history. Decay in time and effects of [Vi(t)]avg or [Vj(t)]avg are also
allowed. Model networks with such synapses (16, 20, 21) can
construct the associative T., of Eq. 2. We will therefore initially
assume that such a Ty1 has been produced by previous experience (or inheritance). The Hebbian property need not reside
in single synapses; small groups of cells which produce such a
net effect would suffice.
The network of cells we describe performs an abstract calculation and, for applications, the inputs should be appropriately coded. In visual processing, for example, feature extraction should previously have been done. The present modeling
might then be related to how an entity or Gestalt is remembered
or categorized on the basis of inputs representing a collection
of its features.
Studies of the collective behaviors of the model
The model has stable limit points. Consider the special case T
= Tji, and define
E =-2

Tij VjVj

[7]

Tij Vj

[8]

ioj

AE due to AV1 is given by

AE =

-AVi
joi'

Thus, the algorithm for altering Vi causes E to be a monotonically decreasing function. State changes will continue until a
least (local) E is reached. This case is isomorphic with an Ising
model. Tij provides the role of the exchange coupling, and there
is also an external local field at each site. When T.j is symmetric
but has a random character (the spin glass) there are known to
be many (locally) stable states (29).
Monte Carlo calculations were made on systems of N = 30
and N = 100, to examine the effect of removing the T.1 = T.
restriction. Each element of T., was chosen as a random number
between -1 and 1. The neural architecture of typical cortical
regions (30, 31) and also of simple ganglia of invertebrates (32)
suggests the importance of 100-10,000 cells with intense mutual interconnections in elementary processing, so our scale of
N is slightly small.
The dynamics algorithm was initiated from randomly chosen
initial starting configurations. For N = 30 the system never
displayed an ergodic wandering through state space. Within a
time of about 4/W it settled into limiting behaviors, the commonest being a stable state. When 50 trials were examined for
a particular such random matrix, all would result in one of two
or three end states. A few stable states thus collect the flow from
most of the initial state space. A simple cycle also occurred occasionally-for example, . A -* B -- A -- B

The third behavior seen was chaotic wandering in a small

region of state space. The Hamming distance between two binary states A and B is defined as the number ofplaces in which
the digits are different. The chaotic wandering occurred within
a short Hamming distance ofone particular state. Statistics were
done on the probability pi of the occurrence of a state in a time
of wandering around this minimum, and an entropic measure
of the available states M was taken
In M=

-2 pi In pi.

[9]

A value of M = 25 was found for N = 30. Theflow in phase space

produced by this model algorithm has the properties necessary
for a physical content-addressable memory whether or not T
is symmetric.
Simulations with N = 100 were much slower and not quantitatively pursued. They showed qualitative similarity to N =
30.
Why should stable limit points or regions persist when Tij
# Tjj? If the algorithm at some time changes Vi from 0 to 1 or
vice versa, the change of the energy defined in Eq. 7 can be
split into two terms, one of which is always negative. The second
is identical if Ty1 is symmetric and is "stochastic" with mean 0
if Tij and Tji are randomly chosen. The algorithm for Tij # Tj,
therefore changes E in a fashion similar to the way E would
change in time for a symmetric Tij but with an algorithm corresponding to a finite temperature.
About 0.15 N states can be simultaneously remembered before error in recall is severe. Computer modeling of memory
storage according to Eq. 2 was carried out for N = 30 and N
= 100. n random memory states were chosen and the corresponding T.9 was generated. If a nervous system preprocessed
signals for efficient storage, the preprocessed information
would appear random (e.g., the coding sequences of DNA have
a random character). The random memory vectors thus simulate
efficiently encoded real information, as well as representing our
ignorance. The system was started at each assigned nominal
memory state, and the state was allowed to evolve until
stationary.
Typical results are shown in Fig. 2. The statistics are averages
over both the states in a given matrix and different matrices.
With n = 5, the assigned memory states are almost always stable
(and exactly recallable). For n = 15, about half of the nominally
remembered states evolved to stable states with less than 5 errors, but the rest evolved to states quite different from the starting points.
These results can be understood from an analysis of the effect
ofthe noise terms. In Eq. 3, H' is the "effective field" on neuron
i when the state of the system is s', one of the nominal memory
states. The expectation value of this sum, Eq. 4, is N/2 as
appropriate. The s # s' summation in Eq. 2 contributes no
mean, but has a rms noise of [(n - 1)N/2]'2- a. For nN large,
this noise is approximately Gaussian and the probability of an
error in a single particular bit of a particular memory will be
P=

e-x2/2a2 dx.

[10]

2 N/2

For the case n = 10, N = 100, P = 0.0091, the probability that

a state had no errors in its 100 bits should be about eC0O9' 0.40.
In the simulation of Fig. 2, the experimental number was 0.6.
The theoretical scaling of n with N at fixed P was demonstrated in the simulations going between N = 30 and N = 100.
The experimental results of half the memories being well retained at n = 0.15 N and the rest badly retained is expected to

Biophysics: Hopfield

Proc. Natl. Acad. Sci. USA 79 (1982)

The model was studied by using a "clipped" Tij, replacing T4,

in Eq. 3 by 1, the algebraic sign of Tij. The purposes were to
examine the necessity of a linear synapse supposition (by making
a highly nonlinear one) and to examine the efficiency of storage.
Only N(N/2) bits of information can possibly be stored in this
symmetric matrix. Experimentally, for N = 100, n = 9, the level
of errors was similar to that for the ordinary algorithm at n =
12. The signal-to-noise ratio can be evaluated analytically for
this clipped algorithm and is reduced by a factor of (2/r)1"2 compared with the unclipped case. For a fixed error probability, the
number of memories must be reduced by 2/ir.
With the 4 algorithm and the clipped Tij, both analysis and
modeling showed that the maximal information stored for N
= 100 occurred at about n = 13. Some errors were present, and
the Shannon information stored corresponded to about N(N/

1.0

0.5

0.5

N-

n=

100

10

N= 100
2&

0.2

8) bits.

0.2

0.2

n=1

N= 100

3 6 9

10- 20- 30- 40- >49

19

29

39

49

Nerr = Number of Errors in S tate

FIG. 2. The probability distribution of the occurrence of errors in
the location of the stable states obtained from nominally assigned
memories.

be true for all large N. The information storage at a given level

of accuracy can be increased by a factor of 2 by a judicious choice
of individual neuron thresholds. This choice is equivalent to
using variables ip = 1, Tij = 1,,u4,4j, and a threshold level
of 0.
Given some arbitrary starting state, what is the resulting final
state (or statistically, states)? To study this, evolutions from randomly chosen initial states were tabulated for N = 30 and n
= 5. From the (inessential) symmetry of the algorithm, if
(101110 ) is an assigned stable state, (010001 .) is also stable.
Therefore, the matrices had 10 nominal stable states. Approximately 85% of the trials ended in assigned memories, and 10%
ended in stable states of no obvious meaning. An ambiguous 5%
landed in stable states very near assigned memories. There was
a range of a factor of 20 of the likelihood of finding these 10
states.

The algorithm leads to memories near the starting state. For

N = 30, n = 5, partially random starting states were generated
by random modification of known memories. The probability
that the final state was that closest to the initial state was studied
as a function of the distance between the initial state and the
nearest memory state. For distance c 5, the nearest state was
reached more than 90% of the time. Beyond that distance, the
probability fell off smoothly, dropping to a level of 0.2 (2 times
random chance) for a distance of 12.
The

phase

space flow is

2557

apparently dominated by

attractors

which are the nominally assigned memories, each ofwhich dominates a substantial region around it. The flow is not entirely
deterministic, and the system responds to an ambiguous starting state by a statistical choice between the memory states it
most resembles.
Were it desired to use such a system in an Si-based contentaddressable memory, the algorithm should be used and modi-

fied to hold the known bits of information while letting the others adjust.

New memories can be continually added to Ti.. The addition

of new memories beyond the capacity overloads the system and
makes all memory states irretrievable unless there is a provision
for forgetting old memories (16, 27, 28).
The saturation of the possible size of Tij will itself cause forgetting. Let the possible values of TY be 0, 1, 2, 3, and
Tt, be freely incremented within this range. If Tij = 3, a next
increment of +1 would be ignored and a next increment of
-1 would reduce Tij to 2. When Ty, is so constructed, only the
recent memory states are retained, with a slightly increased
noise level. Memories from the distant past are no longer stable.
How far into the past are states remembered depends on the
digitizing depth of T., and 0, , 3 is an appropriate level for
N = 100. Other schemes can be used to keep too many memories from being simultaneously written, but this particular one
is attractive because it requires no delicate balances and is a
consequence of natural hardware.
Real neurons need not make synapses both of i -- j and j
i. Particular synapses are restricted to one sign of output. We
therefore asked whether Tij = Tjj is important. Simulations were
carried out with only one ij connection: if T- $0, T.i = 0. The
probability of making errors increased, but the algorithm continued to generate stable minima. A Gaussian noise description
of the error rate shows that the signal-to-noise ratio for given
n and N should be decreased by the factor 1/F2, and the simulations were consistent with such a factor. This same analysis
shows that the system generally fails in a "soft" fashion, with
signal-to-noise ratio and error rate increasing slowly as more
synapses fail.
Memories too close to each other are confused and tend to
merge. For N = 100, a pair ofrandom memories should be separated by 50 5 Hamming units. The case N = 100, n = 8,
was studied with seven random memories and the eighth made
up a Hamming distance of only 30, 20, or 10 from one of the
other seven memories. At a distance of 30, both similar memories were usually stable. At a distance of 20, the minima were
usually distinct but displaced. At a distance of 10, the minima
were often fused.
The algorithm categorizes initial states according to the similarity to memory states. With a threshold of 0, the system behaves as a forced categorizer.
The state 00000 ... is always stable. For a threshold of 0, this
stable state is much higher in energy than the stored memory
states and very seldom occurs. Adding a uniform threshold in
the algorithm is equivalent to raising the effective energy of the
stored memories compared to the 0000 state, and 0000 also
becomes a likely stable state. The 0000 state is then generated
by any initial state that does not resemble adequately closely
one of the assigned memories and represents positive recognition that the starting state is not familiar.

0 rl
,2558

Proc. Natl. Acad. Sci. USA 79 (1982)

Biophysics: Hopfield

Familiarity can be recognized by other means when the

memory is drastically overloaded. We examined the case N
= 100, n = 500, in which there is a memory overload ofa factor
of 25. None ofthe memory states assigned were stable. The initial rate of processing ofa starting state is defined as the number
of neuron state readjustments that occur in a time 1/2W. Familiar and unfamiliar states were distinguishable most of the
time at this level ofoverload on the basis ofthe initial processing
rate, which was faster for unfamiliar states. This kind of familiarity can only be read out of the system by a class of neurons
or devices abstracting average properties of the processing
group.
For the cases so far considered, the expectation value of Tij
was 0 for i # j. A set of memories can be stored with average
correlations, and Ty = Cij # 0 because there is a consistent internal correlation in the memories. If now a partial new state
X is stored
[ill
ATUj = (2Xi -1)(2Xj -1) Q~ k < N
using only k of the neurons rather than N, an attempt to reconstruct it will generate a stable point for all N neurons. The
values of Xk+l-.XN
*
that result will be determined primarily
from the sign of
--

E Cij Xj

[12]

j=1

and X is completed according to the mean correlations of the

other memories. The most effective implementation of this capacity stores a large number of correlated matrices weakly followed by a normal storage of X.
A nonsymmetric T.. can lead to the possibility that a minimum
will be only metastable and will be replaced in time by another
minimum. Additional nonsymmetric terms which could be easily generated by a minor modification of Hebb synapses

ATU = A >

(2Vs+1 - 1)(2Vj - 1)

[13]

were added to T . When A was judiciously adjusted, the system

would spend a while near V. and then leave and go to a point
near V,+,. But sequences longer than four states proved impossible to generate, and even these were not faithfully
followed.
Discussion
In the model network each "neuron" has elementary properties,
and the network has little structure. Nonetheless, collective
computational properties spontaneously arose. Memories are
retained as stable entities or Gestalts and can be correctly recalled from any reasonably sized subpart. Ambiguities are resolved on a statistical basis. Some capacity for generalization is
present, and time ordering of memories can also be encoded.
These properties follow from the nature of the flow in phase
space produced by the processing algorithm, which does not
appear to be strongly dependent on precise details of the modeling. This robustness suggests that similar effects will obtain
even when more neurobiological details are added.
Much of the architecture of regions of the brains of higher
animals must be made from a proliferation of simple local circuits with well-defined functions. The bridge between simple
circuits and the complex computational properties of higher

nervous systems may be the spontaneous emergence of new

computational capabilities from the collective behavior of large
numbers of simple processing elements.
Implementation of a similar model by using integrated circuits would lead to chips which are much less sensitive to element failure and soft-failure than are normal circuits. Such chips
would be wasteful of gates but could be made many times larger
than standard designs at a given yield. Their asynchronous parallel processing capability would provide rapid solutions to some
special classes of computational problems.
The work at California Institute of Technology was supported in part
by National Science Foundation Grant DMR-8107494. This is contribution no. 6580 from the Division of Chemistry and Chemical
Engineering.
1. Willows, A. 0. D., Dorsett, D. A. & Hoyle, G. (1973) J. Neurobiol 4, 207-237, 255-285.
2. Kristan, W. B. (1980) in Information Processing in the Nervous
System, eds. Pinsker, H. M. & Willis, W. D. (Raven, New York),
241-261.
3. Knight, B. W. (1975) Lect. Math. Life Sci. 5, 111-144.
4. Smith, D. R. & Davidson, C. H. (1962)J. Assoc. Comput. Mach.
9, 268-279.
5. Harmon, L. D. (1964) in Neural Theory and Modeling, ed. Reiss,
R. F. (Stanford Univ. Press, Stanford, CA), pp. 23-24.
6. Amari, S.-I. (1977) Bio. Cybern. 26, 175-185.
7. Amari, S.-I. & Akikazu, T. (1978) Biol Cybern. 29, 127-136.
8. Little, W. A. (1974) Math. Biosci. 19, 101-120.
9. Marr, J. (1969) J. Physiol 202, 437-470.
10. Kohonen, T. (1980) Content Addressable Memories (Springer,
New York).
11. Palm, G. (1980) Biol Cybern. 36, 19-31.
12. McCulloch, W. S. & Pitts, W. (1943) BulL Math Biophys. 5,
115-133.
13. Minsky, M. & Papert, S. (1969) Perceptrons: An Introduction to

Computational Geometry (MIT Press, Cambridge, MA).

14. Rosenblatt, F. (1962) Principles of Perceptrons (Spartan, Washington, DC).

15. Cooper, L. N. (1973) in Proceedings of the Nobel Symposium on
Collective Properties of Physical Systems, eds. Lundqvist, B. &
Lundqvist, S. (Academic, New York), 252-264.
16. Cooper, L. N., Liberman, F. & Oja, E. (1979) Biol Cybern. 33,
9-28.
17. Longuet-Higgins, J. C. (1968) Proc. Roy. Soc. London Ser. B 171,

327-334.
18. Longuet-Higgins, J. C. (1968) Nature (London) 217, 104-105.
19. Kohonen, T. (1977) Associative Memory-A System-Theoretic
Approach (Springer, New York).
20. Willwacher, G. (1976) Biol Cybern. 24, 181-198.
21. Anderson, J. A. (1977) Psych. Rev. 84, 413-451.
22. Perkel, D. H. & Bullock, T. H. (1969) Neurosci. Res. Symp.
Summ. 3, 405-527.
23. John, E. R. (1972) Science 177, 850-864.
24. Roney, K. J., Scheibel, A. B. & Shaw, G. L. (1979) Brain Res.
Rev. 1, 225-271.
25. Little, W. A. & Shaw, G. L. (1978) Math. Biosci. 39, 281-289.
26. Shaw, G. L. & Roney, K. J. (1979) Phys. Rev. Lett. 74, 146-150.
27. Hebb, D. 0. (1949) The Organization of Behavior (Wiley, New

York).

28. Eccles, J. G. (1953) The Neurophysiological Basis of Mind (Clar-

endon, Oxford).

29. Kirkpatrick, S. & Sherrington, D. (1978) Phys. Rev. 17, 4384-4403.

30. Mountcastle, V. B. (1978) in The Mindful Brain, eds. Edelman,
G. M. & Mountcastle, V. B. (MIT Press, Cambridge, MA), pp.
36-41.
31. Goldman, P. S. & Nauta, W. J. H. (1977) Brain Res. 122,

393-413.

32. Kandel, E. R. (1979) Sci. Am. 241, 61-70.