Title: The Effects of Hebbian Learning on the Structure and Dynamics of Chaotic Neural Networks
1The Effects of Hebbian Learning on the Structure
and Dynamics of Chaotic Neural Networks
H. Berry1, B. Cessac2, B. Delord 3, M. Quoy4, B.
Siri1
1 Alchemy, INRIA, Orsay, France 2 Institut Non
Linéaire de Nice Odyssee, INRIA,
Sophia-Antipolis , France 3 ANIM, Univ. PM
Curie, Paris , France 4 ETIS, Univ.
Cergy-Pontoise , France
2Cortical circuits display complex structure
V2
- The local micro-connectivity structure of the
cortex is massively recurrent - At several length scales, the connectivity
structure is complex and appears partly random
V1
LGN
3Neurons show complex dynamics
- This complex recurrent structures yield a dense
riche network activity even in the absence of
stimulus ongoing (spontaneous) activity - This ongoing activity
- depends on the recurrent complex structure
- is highly variable but not only noise Arieli et
al (1995) - conditions the evoked response to stimuli Arieli
et al (1996)
Membrane potential of a cortical neuron in vivo
Lampl et al (1999)
4Plasticity yields dynamic connectivity
- e.g. cortical rewiring in ferrets Sur Leamey
(2001)
5Structure / Dynamics in Neural Networks
- Network topology conditions Neuron dynamics
- Synaptic plasticity (i.e. Hebbian learning)
Neuron dynamics conditions Network topology - ? mutual coupling between structure dynamics and
neurons dynamics - A complex dynamical system and key pbl in
neuroscience - But not really understood yet in the case of
complex dynamics
1
W01
0
2
W02
W03
3
W01
0
1
neuron state
connection weight
6Outline of the talk
- Aim Structure/dynamics relationships in random
neural networks with rich dynamics and Hebbian
plasticity - A simple but generic model to allow rigorous
analysis - Structure changes at several length scales
- Effects on network dynamics
- Effects on network function
- Perspectives competition between Hebbian and
intrinsic plasticity
7Chaotic random neural networks
- Firing rate dynamics cortex-like netwk structure
- Random sparse connectivity (15) Gabbott
Somogyi (1986) - 25 inhibitory neurons Markram et al (1997)
j
i
- Initial synaptic weights Wij(0) i.i.d. random
var. set so that initial dynamics is chaotic
(rich dynamics)
8Typical evolution of dynamics complexity
Learning an external input
N 500 neurons
- Hebbian learning generically decreases the
richness of the dynamics
9Structure changes motifs
- Motifs patterns (sub-graphs) that recur more
often than at random - Motifs composition of a complex network its
imprint (Milo et al, 2002) - e.g. 3-node directed motifs
- In the cortex, some motifs are more frequent than
by chance (Sporns Kötter, 2004)
10Motifs evolution by Hebbian plasticity
Statistical enrichment
Learning epochs
- Hebbian plasticity reorganizes the motifs
- Specific motifs are reinforced densely
interconnected ones - Dynamical consequences? (current works)
- No changes at early epochs, i.e. where most
dynamical changes occur
11The small-world of the cortex
- Small-world more-than-by-chance triangle motifs
(cliquishness) AND short mean shortest path - Efficient information transfer both locally and
globally - Experimental quantification of topology at
several length scales - Neural networks grown in vitro physical and
functional - C. elegans neural network (physical)
- Cortico-cortical area physical connectivity in
Cat, Macaque Human - Human cortical functional connectivity via MEG,
EEG or fMRI - Human cortical layer-to-layer connectivity
- All display small-world structure (not scale-free
except in Eguiluz et al. (2005) Phys Rev Lett,
94018102)
12Hebbian Learning-induced SW
Clustering index
Mean shortest path
Learning epochs
Learning epochs
- Hebbian plasticity reorganizes the weights over
the connectivity network as a small-world
structure - Again no changes at early epochs, i.e. where most
dynamical changes occur
13Spectral properties
- Exponential decay of W norm
- Largest Lyapunov exponent
Hebbian learning yields reduction of dynamics
complexity through decay of W norm
L1
Learning epochs
14Pattern Sensitivity
- Quantitative measurement of how network dynamics
changes when pattern is removed - Hebbian learning leads the system close to a
bifurcation point (edge of chaos) where pattern
sensitivity is maximal
Learning epochs
15Summary
- Effects of Hebbian learning in recurrent neural
networks - drastic changes of the motifs/microcircuitry
- synaptic weights redistribute as a small world
structure - shrinks Jacobian weight matrix spectral radius
- this reduces the dynamics complexity entropy
(KS) - Functional effects
- leads the system close to the edge of chaos
bifurcation - where sensitivity to input pattern is maximal
- At long learning times constant firing frequency
no response to pattern presence
16Perspectives intrinsic plasticity
- Activity dependent modification of the
excitability (firing threshold, gain) - No change of the synaptic weights
- Homeostatic effects
Firing frequency
Input current
17HebbianIntrinsic preliminary results
- Dynamics still rich at long times
- Sensitivity to the input pattern preserved
18Perspectives Regulations by glial cells
- Collaboration with E. Ben Jacob (Tel Aviv Univ.,
Israel) and V. Volmann (UCSD Salk Institute, La
Jolla, US). - Understand the possible effects at the single
neuron and network levels of neuronal
excitability regulation by glial cells.
Field, Scientific American, 2004
19Thanks!!
- Further results / analysis in
- Siri et al. (2008) Neural Computation,
202937-2966. - Siri et al. (2007) Journal of Physiology
(Paris),101136-148. - http//www-rocq.inria.fr/hberry/
20The model RRNNs
- 2 different time scales Learning and dynamics
- Learning occurs every t dynamics steps
- Dynamics
21Generic Hebbian Learning
- Initial weights are random Wij(1) N(0, 1/N)
-
- A generic learning rule
- A Hebbian rule if
- h gt 0 for i (postsyn) j (presyn) both active
during T (LTP) - h lt 0 for i (postsyn) inactive j (presyn)
active during T (LTD) - h 0 whenever j (presyn) inactive during T
- For numerical simulations
22Bifurcations
- Consider the Jacobian matrix at x
- DFx spectrum can be contracted by
- Increased neuron saturation to 0 or 1
- The contraction of W spectrum
- Confirmed by numerical simulations
bifurcations
l 0.90
l 0.80
Hebbian learning leads the system through a
bifurcation