Defining Time in a Minimal Hippocampal CA3 Model by Matching Time-span of
Associative Synaptic Modification and Input Pattern Duration
Kurt E. Mitman
1
, Patryk A. Laurent
1
, William B Levy
2
Department of Neurosurgery
2
, College of Arts and Sciences
1
, University of Virginia
P.O. Box 800420, Charlottesville, VA 22908
Abstract – This paper quantifies the time shifting of neuronal
codes in a sparse, randomly connected neural network model of
hippocampal region CA3. As this network is trained to learn a
sequence, the neurons that encode portions of this sequence
characteristically fire earlier and earlier over the course of
training. Here we systematically investigate the effects of the N-
methyl-D-aspartate(NMDA)-governed time-span of synaptic
associativity on this shifting process and how this time-span
interacts with the duration of each successive external input.
The results show that there is an interaction between this
synaptic time-span and externally applied pattern duration such
that the early shifting effect approaches a maximum
asymptotically and that this maximum is very nearly produced
when the e-fold decay time-span of synaptic associativity is
matched to the duration of individual input patterns. The
performance of this model as a sequence prediction device varies
with the time-span selected. If too long a time-span is used,
overly strong attractors evolve and destroy the sequence
prediction ability of the network. Local context cell firing − the
learned repetitive firing of neurons that code for a specific
subsequence − also varies in duration with these two
parameters. Importantly, if the associative time-span is
matched to the longevity of each individual external pattern and
if time-shifting and local context length are normalized by this
same external pattern duration, then time-shifting and local
context length are constant across simulations with different
parameters. This constancy supports the idea that real time can
be mapped into a network of McCulloch-Pitts neurons that lack
a time scale for excitation and resetting.
I. INTRODUCTION
The sequence learning, recoding theory of hippocampal
function [1] depends heavily on a temporally asymmetric rule
governing associative synaptic modification. This temporal
asymmetry plays a critical role for a special kind of sequence
completion: it allows learning that produces timely
predictions [2]. That is, it enables the network to meet an
important requirement − use an early part of a sequence to
generate the rest of the sequence before the rest of the
sequence occurs. The asymmetric associative temporal
characteristic of synaptic modification arises from the on- and
off-rate properties of the NMDA receptor [3].
The recoding process itself has been predicted as time-
shifting to earlier firings [1,4] resulting in a somewhat loosely
overlapping code (see “rough-counting” in [1]). In this way a
sequence of neural firing resembles an incrementing, shifting
binary counter when the neurons are visualized via re-
ordering by the temporal order in which they fire. In terms of
biology, the earlier or backwards shifting of firing in the
model is similar to that recently demonstrated in vivo [5]
where it was complemented by a negative skew in the activity
of a large portion of the neurons [6]. The combination of
earlier-shifting of firing with the formation and lengthening
of place-cell type firing results in a neural code for the
present input which becomes increasingly similar to the
future patterns. This similarity is what allows timely
predictions [1].
When the model learns a nonspatial task, cell firing
resembling place cells occurs, and these neurons are called
local context neurons because they identify a particular
subsequence of a longer sequence. The average duration of
such place-cell type firings is denoted E[L], the average local
context length. When training the model to learn cognitive
tasks, an intermediate to high value of E[L] correlates with
good performance (e.g., [7,8] ). However, if E[L] becomes
excessively large, the network state can be drawn into a noisy
attractor, which could prevent the appropriate sequence
recall. This motivates us to study E[L] as a function of
parameters such as the time constant of synaptic associativity.
As an expected result, we find that E[L] increases with
increasing values of the time constant of associativity.
A central finding of this paper is a matching between the
time-span of associative modification and the duration of the
inputs (stutter length) to the network. We show that this
matching produces a compromise between the predictive
component of the neuronal codes developed during learning
and performance. Specifically increasing the LTP time-span
beyond this compromise does in fact marginally increase the
predictive component of the neuronal codes, but it also
degrades performance and the robustness of the codes. That
is, too large of a time-span can cause the formation of
performance-destroying stable attractors. The other
important observation here concerns the mapping of time
between computational cycles of the simulations and real
time.
By ratioing against input pattern longevity (stutter
length), the neuronal codes developed by training are
constant across parametric changes of the NMDA receptor
off-rate time constant that maps the model into real time.
Thus, the longevity of neuronal firings and the time shifts
measured are also mapped into real time.
II. METHODS
A. The Model
0-7803-7898-9/03/$17.00 ©2003 IEEE 1631