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Complexity in the Brain: Emergent behavior from complex interactions within and between neurons Stan Gielen

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Page 1: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Complexity in the Brain: Emergent behavior from complex interactions within and between neurons

Stan Gielen

Page 2: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Content of presentation

• What is complex, intelligent behaviour ?– http://www.livesteaua.com/tags/robocup-videos/– http://www.robocup.nl/achievements.html

• The neural code: – Neuronal dynamics– Emergent behaviour of nonlinear interactions:

multi-stability and hysteresis• Distributed parallel processing

– The network connectivity is flexible by• Neuronal oscillations• Frequency modulation• Flexible task-dependent organisation

– What are the implicatons for control ?

Page 3: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Once upon a time in 1997 ….

IBM Deep Blue

Gary Kasparov

Page 4: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

What is simple and what is difficult ?

Page 5: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Is chess more complex than playing soccer ?

http://www.livesteaua.com/tags/robocup-videos/http://www.robocup.nl/achievements.html

Page 6: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

BrainComputing capacity : 10 petaflop

Storage capacity ~ 103 terabytes

Energy consumption ~ 30 Watt

1011 neurons

Time constant ~ 1 ms

Fujitsu K-Computer (nov 2011)Computing capacity : 10.5 petaflop*

Storage capacity ~ 10.5 terabyte

Energy consumption ~ 9.9 MW

88128 CPU-processors

Time constant ~1 ns

1 peta flop = 10^15 Floating Point Operations

Page 7: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting
Page 8: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

The neural code

Page 9: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Currentcausing a magnetic field

Dipole,causing an electric field

EEG

MEG

Page 10: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Donders Centre for Cognitive Neuroimaging

~ 100.000 times weaker than earth magnetic field

Page 11: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

You can get much further with a kind word and a gun, than with a kind word alone(Al Capone)

Page 12: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

You can get much further with a kind word and a gun, than with a kind word alone(Al Capone)

You can get much further with experiment and theory, than with experiment alone(Computational NeuroScience group, Nijmegen)

Page 13: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Hodgkin-Huxley model

Fast variables• membrane potential V• activation rate for Na+ m

Slow variables• activation rate for K+ n• inactivation rate for Na+ h

-C dV/dt = gNam3h(V-Ena)+gKn4(V-EK)+gL(V-EL) + I

dm/dt = αm(1-m)-βmm

dh/dt = αh(1-h)-βhh

dn/dt = αn(1-n)-βnnddt

ddt

Page 14: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Neuron at rest

-70 mV

IVVW 3

3

)(1 aVb

W

W

V

Page 15: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

-70 mV

IVVW 3

3

)(1 aVb

W

W

V

Page 16: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

-70 mV +40 mV

Page 17: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Neurons are nonlinear oscillators

Fast variables• membrane potential V• activation rate for Na+ m

Slow variables• activation rate for K+ n• inactivation rate for Na+ h

-C dV/dt = gNam3h(V-Ena)+gKn4(V-EK)+gL(V-EL) + I

dm/dt = αm(1-m)-βmm

dh/dt = αh(1-h)-βhh

dn/dt = αn(1-n)-βnn

Page 18: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Multi-stability and parallel processing

Page 19: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Intra-cellular signaling

External input from other cells

nucleus

Ca2+

Calcium store

Page 20: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting
Page 21: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Cells reveal non-linear behavior

• Cell mitosis• Gene-expression and protein synthesis (e.g. pancreas) • Folding/unfolding of proteins (hormone secretion)

Discrete statesOnce started: complete it

Page 22: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Various stages of growth

Page 23: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Various stable states of cells

Membrane potentialCalcium oscillations

Page 24: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Caex

GCaLClex

GCl(Ca) ATPPMCA pump

Caer

ATP

SERCA pump

IP3R

Coupling : excitable membrane and intracellular calcium oscillations

Page 25: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Stability analysis

)( )( SocCaClCaleakKm

m IIIIIdTdVC

mmdtdm

mm )1()55( mVVhmGI CaCa

mmdtdh

hh )1(

dtBCadIIIJII

dtdCa

SERCACalk

CaERIPPMCA

Calk

Cahm

cyt ][)(3,

2

SERCACalk

CaERIP

ER IIIdt

dCa

)(3

2

wwdtdw

ww )1(

BCakCaBCaTkdtBCad

offcytBon 2][][))(( txf

dtxd

Page 26: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Poincaré map

)(~ tx

)()(~)( tytxtx

)( 1tx

)( 1 Ttx

)(~ tx

The map M : CnCn with

can be written in diagonal form with eigenvalues or Floquet multipliers with

Stability requires

)())(( TtxtxM

ses 2

s

0)( s1|| s

0))(~( txfdtxd

Page 27: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Multi-stability and hysteresis

Gielen et al., Phys Rev Lett 2009

Page 28: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Multi-stability and hysteresis

Page 29: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Conclusion

• In a network of cells with the same conditions everywhere, (clusters of) cells can be in different states (multi-stability) depending on the past (hysteresis).

• Multi-stability allows that subsets of neurons in a network can be in different states (parallel processing !)

• Hysteresis ensures that, once a process is started, it is completed, even if external input changes.

Page 30: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Liley model

1

)1(1)(

)()()(~

)(~

~,~

,)()(

)()()(

ˆ)(2

maxabs

max

,

l

ll

lklk

lk

th

llll

lklklllklklk

lklklk

lk

lklklk

lk

lkrk

eqlk

keqlk

klk

lkiel

klkkrk

kk

eSrSthS

tptthSNe

tIdtd

dtd

ehh

thhth

tIththhdtdh

Liley et al., Phys Rev E 2007

Page 31: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

P4P3O2O1FzCzPz

1 s70 V

Brain rhythmsEyes closed

Alpha-rhythm (about 10 Hz)

Page 32: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Oscillatory activity in the brain at 40 – 80 Hz

Page 33: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Hardware of the brain and functional connectivity

How is the connectivity modulated to tune the brain for each particular task ?

• dynamic interactions

Page 34: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Canolty et al. Oscillatory phase coupling coordinates anatomically dispersed functional cell assemblies, PNAS 2010

Oscillatory phase coupling coordinates anatomically dispersed functional cell assemblies

Page 35: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Dynamic interactions between neurons

1. Plasticity by learning• time scale seconds, hours, days, ….

2. Spike timing dependent plasticity• Time scale milliseconds to second• bottom-up process

3. Modulation of excitability by other neurons• Time scale tenth of millisecond• Top-down process• Responsible for rhythmic oscillations

Page 36: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Human Brain Mapping 30:1791-1800 (2009)

Role of alpha-activity

Hypothesis : information is gated by inhibiting task-irrelevant regions.

Functional inhibition is reflected in oscillatory activity in the alpha band (8–13 Hz).

Page 37: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Human Brain Mapping 30:1791-1800 (2009)

Larger alpha-activity: more false alarms !

Alpha activity reflects a state of reduced perception

We can predict false alarms !

Hypothesis : information is gated by inhibiting task-irrelevant regions. Thefunctional inhibition is reflected in oscillatory activity in the alpha band (8–13 Hz).

Page 38: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Alpha-activity after rehearsal compared to that in “forgotten” condition

Inhibition is important in a word rehearsal taskHuygensgebouw

Page 39: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Alpha-activity after rehearsal compared to that in “forgotten” condition

Inhibition is important in a word rehearsal task

Page 40: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Alpha-activity after rehearsal compared to that in “forgotten” condition

Inhibition is important in a word rehearsal taskHuygensgebouw

Page 41: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Dorsal = “WHERE”

Ventral = “WHAT

Page 42: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Activity in posterior cortical areas (dorsal stream)

Identification Orientation

Selective inhibition of dorsal pathway for tasks requiring ventral stream

Page 43: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Alpha activity is a mechanism to inhibit particular brain areas

What modulates the alpha activity ?

Page 44: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Increase of Theta before a good response

Page 45: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

OOHPS !

Theta activity from frontal cortex is anti-correlated with alpha posterior

Page 46: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

alpha (10 Hz)

Theta (4-6 Hz)

Page 47: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Summary: where are we ?1. Nonlinear system theory is the key to understanding complex

intelligent behaviour.• Neurons and neuronal interactions are nonlinear systems with

complex and highly interesting emergent properties:More is different !

Page 48: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Summary: where are we ?1. Nonlinear system theory is the key to understanding complex

intelligent behaviour.• Neurons and neuronal interactions are nonlinear systems with

complex and highly interesting emergent properties:More is different !

2. We understand the basic principles of• neuronal dynamics• dynamics of neuronal interactions• simple emergent properties

Page 49: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Summary: where are we ?1. Nonlinear system theory is the key to understanding complex

intelligent behaviour.• Neurons and neuronal interactions are nonlinear systems with

complex and highly interesting emergent properties:More is different !

2. We understand the basic principles of• neuronal dynamics• dynamics of neuronal interactions• simple emergent properties

3. We see a glimpse of• the basic principles of the neuronal code (neuronal firing, neuronal

oscillations, neuronal ensembles, flexible distributed parallel processing)

• how intelligent behaviour is represented in the brain4. We have no clue as to how the network is modified for the job and how the

connections between neuronal regions are modulated to optimize the network architecture.

Page 50: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Challenges:

1.Understanding self-organization and changes in network architecture at various hierarchical levels by bottom-up and top-down processes2.Understanding the opportunities and limitations of the neural wetware (e.g. memory, decision making)

(see paper by Barabási)

What is required is a multi-disciplinary team of research groups to address complexity

Page 51: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Thank you for your attention !

Special thanks to my collaborators

Magteld Zeitler Martin Krupa Ingo Bojak

Page 52: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Functional Disconnection of Frontal Cortex and Visual Cortex in ADHD

index

middle

Page 53: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Communication by coherence (CTC) hypothesis

Page 54: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Synchronization of neuronal activity provides a label for cell assemblies

Page 55: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Canolty et al. Oscillatory phase coupling coordinates anatomically dispersed functional cell assemblies, PNAS 2010

Oscillatory phase coupling coordinates anatomically dispersed functional cell assemblies

Page 56: Complexity in the Brain: Emergent behavior from complex ... · intelligent behaviour. • Neurons and neuronal interactions are nonlinear systems with complex and highly interesting

Modulation of Gamma by Theta Rhythm

Short reaction time

Long reaction time