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Biological Mechanisms of Learning
interaction between CPCA Hebbian learning together
with the network property of
inhibitory competition
as described in the previous chapter, and results in dis-
tributed representations of statistically informative prin-
cipal
features
of the input.
The biological basis of learning is thought to be
long-
term potentiation (LTP)
and
long-term depression
(LTD)
. These mechanisms are
associative
or
Hebbian
in nature, meaning that they depend on both presynap-
tic and postsynaptic activation. The associative nature
of LTP/D can be understood in terms of two require-
ments for opening the
NMDA receptor
: presynaptic
neural activity (i.e., the secretion of the excitatory neu-
rotransmitter glutamate) and postsynaptic activity (i.e.,
asufficiently excited or depolarized membrane poten-
tial to unblock the magnesium ions from the channel).
The NMDA channel allows
calcium ions
(Ca
++
)toen-
ter the synapse, which triggers a complex sequence of
chemical events that ultimately results in the modifica-
tion of synaptic efficacy (weight). The available data
suggests that when both presynaptic and postsynaptic
neurons are strongly active, the weight increases (LTP)
due to a relatively high concentration of calcium, but
weaker activity results in weight decrease (LTD) due to
an elevated but lower concentration of calcium.
4.11
Further Reading
The last few chapters of Hertz et al. (1991) on compet-
itive learning provide a clear and more mathematically
detailed introduction to unsupervised/self-organizing
learning.
Hinton and Sejnowski (1999) is a collection of influ-
ential papers on
Unsupervised learning
(model learn-
ing).
Much of Kohonen's pioneering work in unsupervised
learning is covered in Kohonen (1984), though the
treatment is somewhat mathematically oriented and can
be difficult to understand.
The paper by Linsker (1988) is probably the
most comprehensible by this influential self-organizing
learning researcher.
Probably the most influential biologically oriented
application of unsupervised model learning has been
in understanding the development of ocular dominance
columns, as pioneered by Miller et al. (1989).
The journal
Neural Computation
and the
NIPS
con-
ference proceedings (
Advances in Neural Information
Processing
) always have a large number of high-quality
articles on computational and biological approaches to
learning.
Hebbian Model Learning
Model learning is difficult because we get a large
amount of low quality information from our senses.
The use of appropriate a priori
biases
about what the
world is like is important to supplement and organize
our experiences. A bias favoring simple or
parsimo-
nious
models is particularly useful. The objective of
representing
correlations
is appropriate because these
reflect reliable, stable features of the world. A par-
simonious representation of such correlations involves
extracting the
principal components
(features, dimen-
sions) of these correlations. A simple form of Heb-
bian learning will perform this
principal components
analysis (PCA)
, but it must be modified to be fully
useful. Most importantly, it must be
conditionalized
(
CPCA
), so that individual units represent the principal
components of only a
subset
of all input patterns. The
basic CPCA algorithm can be augmented with
renor-
malization
and
contrast enhancement
, which improve
the dynamic range of the weights and the selectivity
of the units to the strongest correlations in the input.
Self-organizing
learning can be accomplished by the
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