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ORIGINAL CONTRIBUTION
YVES BURNOD
Institut des Neurosciences, Paris V
Abstract--We propose in this paper a new connectionist unit that matches a biological model o[ the cortical
column. The architectural and functional characteristics" o f this unit have been designed in the simplest manner
in order to simulate human-like reasoning, and to be as similar as possible to the main known ]batures of real
intracortical networks. We use a new type o f learning rule which can easily take into account goal-oriented
combinations o f actions in behavioral programs. These learning rules are both simple and biologically plausible.
We show in this paper that such units can be used in multilayered networks to perJbrm pattern recognition, with
[eedback connections effecting an attentive gating of sensory information flow. Computer simulations were
performed to assess the ability of a multilayered network made O[ these biologically inspired units to perform
standard speech and visual recognition. Such simulations show levels o f perJbrmance equivalent to the best
currently available connectionist networks .(or typical human-like problems, with very fast learning and recog-
nition processes. Furthermore, this type o f "cortical" unit can be used in more general multilayered networks
with units controlling different types o f external processing, in order to learn programs o f actions which may
be included in the process of recognition.
15
16 L Alexandre, E Guyot, J.-P. Hal(m, and K Burnoa
tie, 1978; Hubel & Wiesel, 1977: Szentagothai, 1975) Behavioral tasks, such as pattern recognition~ can
which is a repetitive circuit perpendicular to the cor- be conceived as involving several sets of laterally
tical surface. interconnected columns working in parallel (like cor-
We thus describe a new connectionist approach, tical maps), which effect serial processing on infor-
with a processing unit which does not correspond to mation originating from the cnvir~mment.
a neuron but to a cortical column (Burnod, 1988).
The main problem is to determine which mathe-
matical operations could model in a simple way the
2.1. Architecture
basic functions of such a multicellular unit (Shaw.
1988). Two sources of inspiration are relevanl. The cortex (and thus, the column) has a six-layered
The first one is neurobiological; the model has to organization. We, like others (Ballard, 1986), have
be biologically plausible: its architectural and func- chosen to group these six layers into three major
tional characteristics have been determined in the input-output divisions. The model unit thus has three
light of neurobiological considerations (see justifi- divisions in direct correspondance with the three ma-
cations in section 2). Even if the cortex is far from jor input-output divisions of tea! cortical columns
being understood, it is possible to use general prin- (Mountcastle, 1978; Jones, 1981):
ciples to define the main types of inputs, outputs,
and interactions. I. The pyramidal neurons of upper layers (layers
The second problem is cognitive; the processing 2 and 3) are specialized in cortico-cortical con-
unit combines essential features for human-like data nections that form direct or indirect connections
processing, but with parallel and distributed repre- between any two columns (Szentagothai, 1975;
sentations. Jones, 1981); the upper division of the unit will
Consequently, the proposed unit will be inter- provide a similar set of connections. These con-
preted both in terms of neuronal activity and in telms nections will be named internal input and internal
of data processing. output (see, for example, Figure 1).
In this paper, we attempt to model this basic cor--
tical function via a minimal set of rules, which never- , The intermediate layer (layer 4) receives the
theless perform more elaborate functions than main sensory inputs, from two different sources:
individual neurons.
either directly from the thalamus which provides
A formal network is developed, inspired from bi-
information from the "external world";
ological findings on the cerebral cortex (Burnod,
1988); this unit can be used to build large multilay- or from other cortical areas involved in earlier
ered networks for general artificial intelligence (AI) stages of sensory processing: connectivity upon
tasks such as pattern recognition (Alexandre et al., this division produces a progressive integration
1989) or reasoning. ol sensory information, by a divergence-con-
vergence resulting in a progressive increase in
the size of the receptive fields (Van Essen &
2. BIOLOGICAL INSPIRATION Maunsell, 1983; Zeki & Shipp, 1988).
The processing unit is designed to be in direct cor- The intermediate division of the model will also
respondance with the "cortical column," a stereo- have an external input which includes feed-for-
typed interneuronal circuit (about 100 /2m wide) ward connections.
whose main characteristics are repeated throughout
the cortex (Mountcastle, 1978; Szentagothai, 1975). . The pyramidal neurons of the lower layers (layer
These clusters of interconnected cells are not defined 5 and 6) are more specialized to effect "actions"
by anatomical boundaries, but rather from their ho- toward the external world or controls upo n the
mogeneous activities: when the firing frequency of information it produces (Zeki & Shipp, 1988;
cortical neurons is recorded in response to specific Van Essen & Maunsell, 1983):
stimuli (Mountcastle, 1978; Hubel & Wiesel, 1977),
or during stereotyped behaviour (Evarts & Tanji, they project to subcortical structures and com-
1974), homogeneous activities reveal sets of cells ar- mand different levels of actions and behavioural
ranged in vertical clusters or columns. It is thus pos- adaptations, and
sible to delimit groups of highly interconnected cells they participate in the feedback projection to
sharing the same set of inputs and outputs (Szenta- previous cortical areas and can thus effect se-
gothai, 1975; Jones, 1981). The lateral width of this lective control of this sensory information flow.
basic unit is determined by the functional homoge-
neity of component pyramidal neurons, in general The lower division of the unit will also provide
due to the intersection between subsets of connective an external output which also includes feedback
stripes (Hubel & Wiesel, 1977). connections.
The Cortical Column 17
Within each area, a column is connected from Wiesel, 1977), or linked with ongoing behavior
both upper and lower layers to neighboring columns (in associative areas; Mountcastle, Andersen, &
of the same "primary indice" (as defined in Ballard, Motter, 1981), or resulting in command of a
1986; e.g., in area 17, the same retinotopic position), movement (in motor areas; Evarts & Tanji,
and to more distant columns sharing similar func- 1974).
tional properties, with the same "secondary indice'"
(e.g., in area 17, the same orientation; Ballard, 1986; . Finally, we include a nul activity state (E0),
Gilbert & Wiesel, 1981). which describes the effects of inhibitory pro-
The three divisions of our unit will thus respect cesses.
major outlines of the cortical connectivity between
columns. Its connections can be used to produce a
2.3. Activation Rules
multilayered network (like a multilayered percep-
tron) but with feedback pathways (like recursive net- Activation rules of the unit will model input-output
works; e.g., Hopfield nets). As columns, the units transformations performed by a column using these
are not fully interconnected; a column is connected three activity levels; they are summarized in Figure
to a limited and rather constant number of cortical 1 which details activities of the two output divisions
columns (Mountcastle, 1978; Szentagothai, 1975). (internal and external for respectively upper and
The general organization of the columns in the lower layers) depending upon activity levels of the
cortex displays cytoarchitectonical variations, such two main inputs (internal and external, respectively
as the relative thicknesses of the six layers. These to upper and intermediate layers) and the previous
variations are introduced in the unit, which has sim- state of the column. Such activation rules will match
ilar parameters; for example, increasing the width of the following three important features of cortical
layer IV (thalamic input) in receptive areas is taken physiology.
into account by changing the weights of the external
input of the unit (see Figure 1). Conditional inhibitions. Strong lateral inhibitions
are observed in the cortex when produced by fo-
2.2. Levels of Activity cal, well-patterned stimuli although there are also
direct corticocortical excitatory connections
In the cortex it is necessary to consider different (Mountcastle, 1978; Hubel & Wiesel, 1977). In a
states of activity to describe with minimal complexity cortical column, inhibitory and excitatory inter-
several important features, such as selective atten- neurons are branched in parallel on the direct
tion, anticipation, or output actions which result in connections between cortical and thalamic inputs
reentrant feedback inputs. Considering all-or-none
activity, like in Hopfield nets (Hopfield, 1982), is
insufficient in the case of the cortex. In our unit, we
Inputs , prior Outputs
will distinguish three states of activity in correspond- InternallExtemal state IntemallExtemal Comments Depends upon:
ance with three cortical functional states (see Figure E0 E0 E0,EI E0 E0 Inactivity
1): E0 E0 Motor
E0 E2 E0,E1 El E0 Associative Map
E2 E2 Sensory
1. The low level, that we will call E1 in the unit, E0 E0 Inhibition
represents moderate neuronal activities (action El E0 E0,E 1 E1 E0 Gating Learning
potential frequencies in the range of 10 spikes/ E2 E2 Triggering
sec) which are insufficient to result in output El E2 E0,E 1 E2 E2 Amplification
actions: however, such activity, mostly intra- E0 E0
E2 E0,E2 E0,E1 Inhibition
E1 E0
cortical, can reflect active processes during se- E2 E0,E2 E2 E2 E2 Reactivation
lective attention, as seen for example in parietal
areas when stimuli do not match ongoing be- FIGURE 1. Activation rules for a simple model of the comical
haviour (Mountcastle, Andersen, & Motter, column. An in-out table imposes the state of activation of
1981), and also anticipation, as seen for example the two outputs (OF,, OEJ, in the right part) when the activities
in frontal areas during waiting for a go signal of internal and external (11/,, IEi) inputs, as well as the prior
(Fuster, 1977). state of the unit, are known (left part of the table). Three
activity levels are considered: E0, inhibition (or FALSE); El,
low level (or PERHAPS); E2, high level (or TRUE). The six
. A higher level of activation, which we term E2 lines of the table display the different possible combinations,
(action potential frequencies with a greater mag- and are ordered by the intensity of the internal inputs (see
nitude, 50-100 Hz), models activities observed text for detailed description). When more than one internal
during sensorimotor interactions with the exter- input is active, each potential level of activation is evaluated;
the more numerous is selected to be the real level of acti-
nal world, for stimuli matching intrinsic cortical vation. In case of equality, the unit is set to the uncertain
filtering functions (in receptive areas; Hubel & state El. After learning, this case tends to disappear.
18 E Alexandre, l-i Guyot, .]-E ttato:a and Y Bumod
onto pyramidal neurons (Szentagothai, 1975; properties and has no simple physiological correlates
Jones, 1981); the relation between columns is at the neuronal level.
controlled in parallel by excitatory and inhibitory It is possible, however, to define learning rules
interneurons, which tend to produce nonlinear which match the following features of the cortical
effects, with a spatiotemporal selectivity depend- network, both at the global and local levels.
ing upon branching patterns and channels of in-
terneurons. The combination performed by the The global logic of the learning rules corresponds
unit (line 5 of Figure 1) will be qualitatively dif- to operant conditioning: if a strongly active col-
ferent for different levels of activity; an increasing umn participates in an action outside the cortex,
level of activity can result initially in excitation and if this external action reactivates the inputs
(for a low level) followed by inhibition (for a of this column (equivalent to a " t~cward,'' or more
high level). Consequently, a strong activity level generally, to a "goal"). this input will gain b\.
will inhibit related units with a lower activity. learning a new influence: whet~ it has a low ac-
tivation (corresponding to :l 'drive"), it sclec.-
Gating and amplification. Moderate cortical in-
tively activates columns whose actions were
puts have a gating effect on other inputs, for ex-
previously efficient in satislying the drive state,
ample, on thalamic inputs (Connors, Gutnick, &
Prince, 1982; Asanuma, Waters, & Yumina,
Local learning rules correspond mostly to acti\-
1982). Consequently, in the unit (line 3 of Figure
ity-dependent plasticity of cortico--cortical path-
1), an internal input alone will have a weak in-
ways: these pathways are mediated by glutamate
fluence (at level El), but the coactivation of two
and involve receptors with potentiating proper-
inputs will generate a strong nonlinear increase
ties (Barrionuevo & Brown, 1983). This plasticity
of activity (E2).
occurs when a strong depolarization of the cell
Selective filtering oJ" inputs, As described in the (upper-pyramidal neurons) is followed by a
cortex, the operation upon the exernal and feed- strong reactivation of its inputs (due to glutamate
forward inputs will be modelled by a spatial fil- release). Long-term changes in the efficacy of this
tering process, with a central positive region and pathway will increase the influence of moderate
a peripheral negative one. This type of filtering cortico-cortical inputs (gating properties), but
is similar to the transfer function assumed to pro- will not influence the other combinations, p a r t >
duce orientation selectivity in visual areas from ularly with higher inputs.
thalamic inputs (Hubel & Wiesel, 1977).
Consequently, we will differentiate four stages of
Combining these rules results in a prediction: cor- learning that can be easily interpreted as activity-
tical activity will spread in the cortico-cortical net- dependent changes in the different types of cor-
work (via the upper layers) for a moderate activity, tical interneurons. The prediction made by con-
with no output outside the cortex. In contrast, a sidering this unit as a model of the cortical column
strong local pattern of activation will have a strong is that different types of interneurons can modify
inhibitory effect on less-activated columns and will their transmission efficiencies ~or specific acti-
produce a precise combination of output actions (via vation patterns which depend upon the specific
lower-pyramidal neurons). Such a model gives an input-output connectivities of each type of cell.
interpretation of neuronal activities within the cortex
The learning rules of the unit (Figure 2) sum-
and their behavioral correlates: a higher level of well-
marize the overall long-term activity-dependent
patterned activity corresponds to reaching a goal,
changes of neural transmissions within and be-
whereas lower activity levels correspond to a search-
tween columns, mainly by their global effects oll
ing process (propagation of activity) or an attentive
the input-output roles. We do not take into ac-
state.
count synaptic weights from ~,euron to neuron,
but transmission coefficients between input and
2.4. Learning Rules output divisions of the units. Figure 2 (in the
Local learning rules, such as the Hebb rule or the right-hand side) shows how a moderate internal
delta rule, enable neural nets to learn global in-out input (from a unit A, "presynaptic") can produce
functions; mathematical generalization of these rules four different effects on the target unit (B, which
for multilayered networks was provided with back- is the learning unit), depending upon previous
propagation algorithms. Such algorithms have been patterns of activities. Critical features for learning
used to model the transformations performed by (left part of Figure 3) are time coupling and states
cortical areas (Zipser & Andersen, 1988). But of activity, measured by two "repetition factors"
generalization of learning rules such as the back- called P2 and P 0 : P 2 represents the probability
propagation algorithm is based upon mathematical that the learning unit B is active (in state E2)
The Cortical Column 19
complementary interpretation in terms of data pro- Oli(t), OEi(t), where j and k represent the connected
cessing. units and t stands for the time.
We propose a realization with six main features, This distinction corresponds to the upper layers
sufficient to specify a connectionist network (Ru- of the cortex for internal relations, to the lower layers
melhart & McCielland, 1986). A description and a for external outputs, and to the intermediate one for
data-processing interpretation are given for each external inputs.
characteristic.
Inputs and outputs of a processing unit i are thus the external input is either a stimulus or a feed-
divided into an internal component (for concepts) forward input from other maps. These external
and an external one (for stimuli and actions), as connections (both input and output) are orga-
shown in Figure 1; they are denoted by II~ (t), IE~ (t), nized in continuous overlapping receptive fields.
ssA
Connections
Internal Input/output intra-areal
( External output
FIGURE 3. Organization and connectivity of the processing levers (model.r.: cortical a r e a s ) ~ in the simulation. This
network tot visual ~ i has ~
arrows show feed-forward and ~ internal
sensory areas. Transfer functions of milts in the ~
in o w r ~ ~ ~ ~ by~ ~ l . ~
units in associative area, both extra- and Intlll4reel.
The Cortical Column 21
The internal neighboring connections can be re- Finally, it is necessary to define a rule to combine
garded as potential pathways between concepts several synchronous internal inputs at time t for a
where information propagates step by step. unit i.
If the event "internal output j of a unit i at time
t in state Ex" is denoted by OI ~,x(t), and the "number
3.3. Activity Levels
of event Ollx(t) f o r j = 1 t o n l + n2" is denoted
A three-valued logic is a minimal modelization of by nx, then OIi(t) = Ex if max(n0, nl, n2) = nx.
human-like processing, in order to take into account In case of equality, the unit is set to El, uncertain
anticipatory and attentive behaviours. Moreover, the state,
three values directly correspond to the three states The global external output is computed in the
of activity described in the neurobiological model same way.
(EO, E l , E2): This truth table (Figure 1) is built from the neu-
robiological data and is a model of neuronal inter-
E2 stands for the " T R U E " value: depending actions in a cortical column. Moreover, in the light
upon its localization, it can be interpreted as a of previous interpretations, the table corresponds to
"sure" stimulus, action, or concept. a logical data-processing mechanism. Only the cases
which modify the outputs of the unit are taken into
E0 stands for the " F A L S E " value: in the neu- account.
robiological model, it is often due to an inhibi-
tion. Line 5 illustrates the inhibition mechanism, that
selectively limits the number of active units. Val-
E1 stands for " P E R H A P S " : it can be interpreted idation of a hypothesis (decision) results in a
as a hypothesis. Generally, when a hypothesis is suppression of competitive hypothesis.
emitted, the system will search to validate it; tran-
sition from E1 to E2 stands for the validation of By contrast, in line 6, two different decisions,
such a hypothesis, that is a decision. when successive, are not competitive; further-
more, this temporal relation will result in learn-
The intermediate state of activity can propagate in ing.
the network; this propagation allows a limited num-
ber of connections between units. The communica- Line 3 is learning dependent, according to the
tion between distant units can be performed by coefficients P0{ and P2~ (see "learning rules").
spread of activity from one unit to the next. Prop- When a hypothesis is emitted, its influence de-
agation of state E1 is thus interpreted as propagation pends upon previous experiences as described in
of hypothesis, that is, an active search. the learning rules section. It can suppress other
possibilities (E0), emit a new hypothesis (El) or
trigger a decision (E2).
3.4. Activation Rules
The unit has to compute its outputs (internal and • Line 4 shows how a hypothesis is validated when
external) as a function of its inputs, of its prior state, it matches an external stimulus.
and of previous learning. The two following functions
have to be defined: In line 2, three different kinds of topologic maps
are distinguished in the network, each kind de-
OL(t) = f(IIi(t),IE~(t ), OE,(t - 1),P0J,P2 ', ), voted to a specific step in the information anal-
ysis. A stimulus will imply a certitude in "sensory
OE,(t) = g(IF(t), IE~(t), OE,(t - 1),P0',P2',),
maps," a hypothesis in "associative maps," but
where j =- 1. . . . nl + n2; k indexes the external cannot directly trigger an action in "motor maps."
receptive field and P~ measure previous learning, as
described below.
3.5. Learning Rules
At time t, for each unit i, a global external input,
denoted by IEi, is computed as a function of the Like activation rules, learning rules are both consis-
IE~(t). It measures the correspondance between the tent with neurobiological data (see Section 2) and
input mask of a unit i, defined a priori without learn- with a data-processing interpretation. As in stan-
ing, and its external inputs IE~; IEi is set either to dard neural nets, learning is locally computed. Figure
E2 (exact correspondance) or E0 (too much differ- 2 shows how an internal input j (from a unit Af ) can
ence). produce four different effects on the target unit A,
We then compute local outputs (OE~, OI~) for (right part of the table), depending upon the prob-
each internal input II~ modulated by IEi thanks to ability of their coactivation expressed by the repe-
the truth table shown in Figure 1. tition factors P21 and P0 I. If N(e) measures the
22 k Alexandre, F Guyot, .f.-P. Hattm. and E Burnod
number of occurrences of the event e from tO up to quency ranges from 0 to 8000 Hz. and from 10
current time, then to 50 frames allocated to the word.
p0~ = N(IU2(t).and. OI,0(t - d t ) ) Connections between initial encoding and SMI
N(II~2(t)) are unidirectional and retinotopically (or tono-
and topically) organized. In SMI, units respond to the
input, selectively oriented (Hubel & Wiesel,
p21 = N(IU2(t).and. O1,2(t - dt)) 1977) into local 3 × 3 overlapping receptive
N(II ~,2(t)) fields, as illustrated in Figure 3, Similarly, in the
auditory case. units are specialized for time and
where dt stands for a little delay.
frequency range.
The four cases may be interpreted as follow:
As illustrated in Figure 3, each unit in SMI re-
• Before learning, propagation of hypothesis is ran-
ceives its "'external" 3 x 3 inputs from a field of
dom,
SMI (feed-forward), sends back its actions to
SMI, exchanges symetrically its internal inputs
During learning, if a concept (Aj in state E0)
and outputs with its neighbors in SMII, and is
never participates to the validation of another
connected with a group of adjacent units in AM.
concept (A i, E2), the latter will gain an antici-
patory inhibitory effect (P0',I = 1).
In the associative map AM. each unit carries out
a coactivation test between two neighborhoods
But if the concept Ag is sometimes linked with
taken from SMII and SM, and has reciprocal links
the validation of the concept A/, learning will
with adjacent units in AM (internal relation).
result in a selective propagation of hypothesis A,
toward A~ (0 < P0{ < 1, 0 < P2', < 1).
The map SM contains either a set of 26 units for
visual recognition of letters (see Figure 3) or l0
Finally, if the coactivation pattern is always ob-
units for number recognition in speech process-
served, a hypothesis from Aj will directly trigger
ing; these units are reciprocally connected and
a decision of A~, even in absence of external con-
receive inputs from groups of adjacent units in
firmation (P2~ = 1).
AM.
g
UJ
N'H"
:+: :+:÷:
. '5:::::'+::
.'...
': iiiiii i
I
: !
Pr ubab i ba'~pI
FIGURE 5. Internal and external representation. This figure represents configurations in the network for recognition of isolated
words, in order to Illustrate the basis on which the matching operation is effected. The two pictures on the left show the
memorized coefficients between units of AA and two different units of SA recognizing a "2" (first picture) or s "3" (second
picture). This "mental representoUon" is obtained after learning 20 x 4 occurrences of the digits. By contrast, the two pictures
on the right ~ the Initial o n c ~ s for two spectrograms, both corresponding to the digit "3," and produced by two
different speakers (note their disparity). These occurrences: are recognized by the stored coefficients of the mental represen-
tation of "3" (second picture).
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