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Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot Neuroinformatics and Theoretical Neuroscience Institute of Biology – Neurobiology Bernstein Center for Computational Neuroscience Teaching Week Computational Neuroscience @ Mind and Brain School Martin Nawrot Oct 12, 2011 Introduction to Computationa Neuroscience Neural Coding

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Page 1: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Neuroinformatics and Theoretical Neuroscience

Institute of Biology – Neurobiology

Bernstein Center for

Computational Neuroscience

Teaching Week Computational Neuroscience @ Mind and Brain School

Martin Nawrot

Oct 12, 2011

Introduction to Computationa Neuroscience

Neural Coding

Page 2: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Time table

Exercises: Computer Pool, Room 224

Page 3: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

XX

• Course layout: time table | exercises & rooms | link for download

• Braitenberg ?

• AG: who we are, what we do

• Kursangebot: modelling / data analysis / reading seminars

• Abstraction versus detail

• Single neuron models: levels of abstraction: Herz et al.

Biology | Biophysics | Bioinformatics | Computational Biology Computational Neuroscience | Computer Science | Econometrics

Engineering | Information Technology | Physics | Mathematics

Our Team

Page 4: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Outline

Introduction

Spatial and Temporal Scales of Brain Signals

Neural Coding

Page 5: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Introduction

Page 6: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Experiment

Modeling

Data Analysis

Hypothesis

Applications

Page 7: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Experiment

Modeling

Data Analysis

Hypothesis

Applications

Statistics of Animal Behavior Spike Train Statistics

Machine Learning Linear Systems Analysis

Dynamical Systems Analysis GLM

Point Process Theory Dynamical System Theory

Machine Learning Spiking Neural Networks

Electrophysiology Imaging Data

Behavioral Experiments (Insects and Primates)

Cortical Slice Recordings

Neuromorphic Harware Neuromorphic Robots

Experimental Design Model Design

Grant Proposals

Page 8: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Experiment

Modeling

Data Analysis

Hypothesis

Applications

Statistics of Animal Behavior Spike Train Statistics

Machine Learning Linear Systems Analysis

Dynamical Systems Analysis GLM

Point Process Theory Dynamical System Theory

Machine Learning Spiking Neural Networks

Electrophysiology Imaging Data

Behavioral Experiments (Insects and Primates)

Cortical Slice Recordings

Neuromorphic Harware Neuromorphic Robots

Experimental Design Model Design

Grant Proposals

Page 9: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Models in computational neuroscience …

functional model

stochastic model

connectionist model

neural network model

phenomenological model biophysical model

abstract model

mathematical model

computational model

statistical model Artificial Neural Network

Page 10: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

top

– do

wn

bo

tto

m –

up

me

ch

an

isti

c

co

gn

itiv

e

BEHAVIOR and COGNITION

MOLECULES

approaches experiments models

Page 11: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Spatial and Temporal Scales of Brain Signals

• Measurement techniques

• Neural output: spike train recordings

Page 12: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Spatial Scales

Page 13: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Resolution in Space and Time

(adapted from Matt Fellows)

PET

20μm

Vm

intracellular

Ca-imaging NIRS

Page 14: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

top

– do

wn

bo

tto

m –

up

me

ch

an

isti

c

co

gn

itiv

e

BEHAVIOR and COGNITION

MOLECULES

approaches experiments models

EEG / MEG fMRI PET NIRS

behavioral (psychophysics, language, …)

spike train recordings intracellular measurements

Ca imaging

local field potentials multi-unit-activity

Page 15: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Direct vs. Indirect Measures of Neuronal Signals

20μm

electrophysiology imaging

direct measurement of neuronal signals visualization of single neuron activity

intracellular recording from

- single neurons / dendrites

extracellular recording of

- action potentials (SUA/MUA) and

- local field potential (LFP, indirect)

electrocorticography (ECoG)

- epicortical field potentials

electroencephalography (EEG)

magnetoencephalography (MEG)

optical imaging with

voltage sensitive dyes

functional magnetic

resonance imaging (fMRI)

positron emission tomography (PET)

measurement of electric mass signals

optical imaging of

intracellular Ca activity

- in vitro / in vivo

- 2D / 3D

visualization of average activity

imaging pics from:

Stosiek03

Page 16: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Intracellular recording

© Clemens Boucsein

‘Spike Train’

Page 17: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Extracellular recording

© C

lem

ens B

oucsein

, U

niv

ers

ität F

reib

urg

INPU

T

OU

TPU

T

Page 18: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Extracellular recording

Page 19: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Extracellular recording

► metal | silicon | glas electrodes

► spike output activity

► in vitro | in vivo

culture in vitro

Page 20: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Extracellular recording

Valentino Braitenberg & Almut Schüz

Cortex: Statistics and Geometry of Neuronal Connectivity

Springer, Berlin, 1998 (Second Edition)

Cortex

Page 21: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Extracellular recording

The Mouse Cortex Cortex

Valentino Braitenberg & Almut Schüz

Cortex: Statistics and Geometry of Neuronal Connectivity

Springer, Berlin, 1998 (Second Edition)

Undersampling !

spikes field potential

Page 22: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

State of the art : 100 channels in parallel (Utah Array)

Positioning the array Pneumatic insertion recordings

Thomas Brochier, Alexa Riehle, CNRS, Marseille.

Page 23: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

State of the art : 100 channels in parallel (Utah Array)

Monkey L

RH

Thomas Brochier, Alexa Riehle, CNRS, Marseille.

Page 24: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Extracellular recording

Page 25: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Extracellular recording: spikes

Extrazelluläre Aufnahmen im somatosensorischen Kortex der Ratte. Spontanaktivität unter Anästhesie.

Data Curtsey: Clemens Boucsein und Dymphie Suchanek, Neurobiologie & Biophysik, Universität Freiburg

Page 26: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Extracellular recording: spikes

Extrazelluläre Aufnahmen im somatosensorischen Kortex der Ratte. Spontanaktivität unter Anästhesie.

Data Curtsey: Clemens Boucsein und Dymphie Suchanek, Neurobiologie & Biophysik, Universität Freiburg

Page 27: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Extracellular recording: spike train

‘Spike Train’

Extrazelluläre Aufnahmen im somatosensorischen Kortex der Ratte. Spontanaktivität unter Anästhesie.

Data Curtsey: Clemens Boucsein und Dymphie Suchanek, Neurobiologie & Biophysik, Universität Freiburg

Page 28: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Extracellular recording: spike sorting

100

0

-100

-200

-300

µV

A_D1

_-70

18

1050 1060 1070 1080 1090 1100s

Spike Templates

Principle Components

Time (s)

X G X

-71.61

200

0

-200

µV

A_D

1_-7

018

11

200

0

-200

µV

D_2

12

200

0

-200

µV

D_3

13

Events3

254 255 256 257 258 259 260s

02 01100

0

-100

-200

µV

A_D

1_-7

0

18

11

258.20 258.22 258.24s

Voltage (

µV

)

Extrazelluläre Aufnahmen von α-extrinsischen Neuronen im Bienengehirn. Antwort auf Duftreiz.

Data Curtsey: Dr. Martin Strube, Neurobiologie, Freie Universität Berlin

Page 29: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Extracellular recording: spike sorting

Estimated number of spike sorting errors :

[1] [2]

false positive rate : 13% ~10%

false negative rate: 9% ~10%

[1] Joshua, Elias, Levine, Bergman (2007) J Neurosci Meth doi: jneumeth.2007.03.012

[2] Pouzat, Delescluse, Viot, Diebolt (2004) J Neurophysiol 91

single unit activity ≠ single neuron activity

Page 30: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Neural output: spike train

10100001000100000000001010010100000100010100000000001

binary representation

(array)

discrete representation

(list)

t1 t2 tn

‘ spike train ‘

discrete time series of events

Page 31: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

• Rate Code vs. Temporal Code

Neural Coding I

Page 32: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Rate Code vs. Temporal Code

© S. Reichinnek & C. Boucsein

Rate Code

concept: information encoded in

neuronal firing rate

population code permits encoding of

precise information

redundancy in large populations permits

fast transmission

neuron acts as ‘Integrator’

Temporal Code

concept: information encoded in

temporally precise spike times and spike

patterns across neurons

fast information transmission through

coincident spiking in neuronal assemblies

encoding with few action potentials

(sparse code) explores large coding space

neuron acts as coincidence detector

Contra:

- little conclusive evidence

- single action potentials and coincident

events vanish in the variable ongoing

network activity

Page 33: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Rate code: concept and stimulus tuning

Hubel and Wiesel (1968) J Physiol 195: 215-43

T

• spike count N in tim interval T

• rate r = N/T

Page 34: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Rate code: Time-varying rate estimate

Single unit activity from primary motor cortex of the monkey during repeated reaching movement

Data Curtsey: Alexa Riehle, CNRS, Marseille

Page 35: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Rate Code vs. Temporal Code

© S. Reichinnek & C. Boucsein

Rate Code

concept: information encoded in

neuronal firing rate

population code permits encoding of

precise information

redundancy in large populations permits

fast transmission

neuron acts as ‘Integrator’

Contra:

- rate coding requires integration time (it is

slow)

- large numbers of action potentials

consume a large amount of energy

- combinatorial explosion in large coding

space (grand mother cells)

Temporal Code

concept: information encoded in

temporally precise spike times and spike

patterns across neurons

fast information transmission through

coincident spiking in neuronal assemblies

encoding with few action potentials

(sparse code) explores large coding space

neuron acts as coincidence detector

Contra:

- little conclusive evidence

- single action potentials and coincident

events vanish in the variable ongoing

network activity

Page 36: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Rate Code vs. Temporal Code

© S. Reichinnek & C. Boucsein

Rate Code

concept: information encoded in

neuronal firing rate

population code permits encoding of

precise information

redundancy in large populations permits

fast transmission

neuron acts as ‘Integrator’

Contra:

- rate coding requires integration time (it is

slow)

- large numbers of action potentials

consume a large amount of energy

- combinatorial explosion in large coding

space (grand mother cells)

Temporal Code

concept: information encoded in

temporally precise spike times and spike

patterns across neurons

fast information transmission through

coincident spiking in neuronal assemblies

encoding with few action potentials

(sparse code) explores large coding space

neuron acts as coincidence detector

Page 37: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal code: experimental evidence I

Gollisch and Meister (2008) Science 319

Page 38: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal code: experimental evidence I

Gollisch and Meister (2008) Science 319

Page 39: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal code: experimental evidence II

single unit recorded in the auditory nerve of the cat

Aertsen, Smolders & Johannesma (1979) Biol. Cyber. 32, 175-185

cross-correlogram of trials 1 + 2

Page 40: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal code: experimental evidence III

Riehle, Grün, Diesmann, Aertsen (1997) Science 278, 1950-53

significant coincident spiking in motor cortical units related to expectation

Page 41: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal code: neuron acts as coincidence detector

Inp

ut L

ine

s

Ou

tpu

t L

ine

► coincident excitatory synaptic inputs are

translated into a single spike output

input

output

Burkitt (2006) Biol. Cyber. 95

Page 42: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal code: ‚synfire chain model‘

Kumar, Rotter, Aertsen (2010) Nat Rev Neurosci

a) Hebb 1949

b) James 1890

c) Abeles 1991

d) Diesmann 1999

f) Griffith 1963

M Nawrot ▪ FU Berlin ▪ SFA in the insect olfactory pathway

Page 43: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal code: ‚synfire chain model‘

Curtsey: Sonja Grün

Page 44: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal code: ‚synfire chain model‘

Pulspaket Eingang

pulse packett:

a = number of presynaptivc spikes

σ = temporal dispersion of spikes

.

Diesmann et al. (1999) Nature 402: 529-532

Page 45: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal code: ‚synfire chain model‘

Diesmann et al. (1999) Nature 402: 529-532

Page 46: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal Code: are neurons precise enough ?

Boucsein et al. (2011) Frontiers in Neuroscience 5:32

Page 47: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal Code: are neurons precise enough ?

Nawrot et al. (2009) Frontiers in Neural Circuits 3:1

► high reliability of synaptic transmission(> 90%)

► high temporal precision across trials (jitter < 1ms)

► modest amplitude variability (CV ~ 0.25)

t

Page 48: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal Code: are neurons precise enough ?

Nawrot et al. (2009) Frontiers in Neural Circuits 3:1

► dendritic generation is temporally precise and reliable

Page 49: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Mainen & Seijnowski (1995) Science 268

► spike generation is temporally precise and reliable

Temporal Code: are neurons precise enough ?

Page 50: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Temporal Code: are neurons precise enough ?

Susanne Reichinnek (2007) Diplomarbeit

σ = 1ms

σ = 11ms

a = 40 a = 10

ou

tpu

t s

pik

e d

en

sit

y

Page 51: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Rate Code vs. Temporal Code

© S. Reichinnek & C. Boucsein

Rate Code

concept: information encoded in

neuronal firing rate

population code permits encoding of

precise information

redundancy in large populations permits

fast transmission

neuron acts as ‘Integrator’

Contra:

- rate coding requires integration time (it is

slow)

- large numbers of action potentials

consume a large amount of energy

- combinatorial explosion in large coding

space (grand mother cells)

Temporal Code

concept: information encoded in

temporally precise spike times and spike

patterns across neurons

fast information transmission through

coincident spiking in neuronal assemblies

encoding with few action potentials

(sparse code) explores large coding space

neuron acts as coincidence detector

Contra:

- little conclusive evidence

- single action potentials and coincident

events vanish in the variable ongoing

network activity

Page 52: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

• Encoding – Recoding – Decoding

Neural Coding II

Page 53: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Olfactory system of the honeybee (insect)

30µm

Krofczik, Menzel & Nawrot (2008) Frontiers Comp Neurosci 2 Galizia & Rössler (2009) Annu. Rev. Entomol. 55

T2 03

T1 22

LN

Page 54: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Olfactory pathway in the honeybee (insect)

1st order

Receptor

Neurons

~ 60.000 ORNs

2nd order

Projection

Neurons

~ 950 PNs

3rd order

Kenyon

Cells

~ 160.000 KCs

4th order

Extrinsic

Neurons

~ 400 ENs

INPUT OUTPUT

x 100 / 100

PLASTICITY

Antenna Antennal Lobe (AL) Mushroom Body (MB)

M Nawrot ▪ INRA ▪ Feb 03, 2011

Page 55: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Encoding of odors in the antennal lobe

1st order

Receptor

Neurons

~ 60.000 ORNs

2nd order

Projection

Neurons

~ 950 PNs

3rd order

Kenyon

Cells

~ 160.000 KCs

4th order

Extrinsic

Neurons

~ 400 ENs

Antenna Antennal Lobe (AL) Mushroom Body (MB)

INPUT OUTPUT

M Nawrot ▪ INRA ▪ Feb 03, 2011

Krofczik, Menzel & Nawrot (2009) Frontiers in Computational Neuroscience 2

Page 56: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Krofczik, Menzel & Nawrot (2009) Frontiers Comp Neurosci 2

► Intracellular recordings from projection

► Reliable and stereotypic rate responses

M Nawrot ▪ INRA ▪ Feb 03, 2011

Encoding of odors in the antennal lobe : rate code

Page 57: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Encoding of odors in the antennal lobe : rate code

► >50% of PNs activated by single odor (broad odor tuning)

M Nawrot ▪ INRA ▪ Feb 03, 2011

Page 58: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Encoding of odors in the antennal lobe : rate code

► >50% of PNs activated by single odor (broad odor tuning)

► odor specific binary activation pattern (combinatorial code)

M Nawrot ▪ INRA ▪ Feb 03, 2011

Page 59: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Encoding of odors in the antennal lobe : rate code

Krofczik, Menzel & Nawrot (2009) Frontiers Comp Neurosci 2

► >50% of PNs activated by single odor (broad odor tuning)

► odor specific binary activation pattern (combinatorial code)

► rapid odor encoding within tens of milliseconds

M Nawrot ▪ INRA ▪ Feb 03, 2011

Page 60: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Encoding of odors in the antennal lobe : glomerular space

Page 61: Introduction to Computationa Neuroscience Neural Coding€¦ · Introduction to Computationa Neuroscience Neural Coding . Teaching Week Computational Neuroscience | Mind and Brain

Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Recoding in the mushroom body

1st order

Receptor

Neurons

~ 40.000 ORNs

2nd order

Projection

Neurons

~ 950 PNs

3rd order

Kenyon

Cells

~ 160.000 KCs

4th order

Extrinsic

Neurons

~ 400 ENs

Antenna Antennal Lobe (AL) Mushroom Body (MB)

INPUT OUTPUT

M Nawrot ▪ INRA ▪ Feb 03, 2011

Strube-Bloss, Nawrot & Menzel (2011) J Neurosci 31

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Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Classical conditioning: experimental paradigm

Acquisition

POST PRE

5 odors 10 trials

random order

5 odors 10 trials

random order

CS US

Experiments by Martin Strube-Bloss in the lab of Randolf Menzel

20 min 3h

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Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Classical conditioning: experimental paradigm

Strube-Bloss, Nawrot & Menzel (2011) J Neurosci 31

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Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Classical conditioning : behavioral performance (group)

* Trial 5 G=7.2; p<0.01;

df=1; Trial 10 G=10.5;

p<0.01; df=1

* Trial 1 G=5.37; p<0.05;

df=1]; trial 6 [G=8.79;

p<0.05; df=1

Acquisition [N=36]

trial

1 2 3 4 5 6 7 8 9 10

% m

17

ac

tiv

ity

0

10

20

30

40

50

60

CS+

CS-

**

PostAcquisition_Test [N=17]

trial #

1 2 3 4 5 6 7 8 9 10

%m

17

ac

tiv

ity

0

20

40

60

80

100CS+

CS-

Ctr_A

Ctr_B

Ctr_C

* *

► Bees learn odor-reward association under experimental conditions

Strube-Bloss, Nawrot & Menzel (2011) J Neurosci 31

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Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Single unit recording at mushroom body output

Strube-Bloss, Nawrot & Menzel (2011) J Neurosci 31

INPUT OUTPUT x 100 / 100

PLASTICITY

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Teaching Week Computational Neuroscience | Mind and Brain | M Nawrot

Odor VALUE coding at mushroom body output

M Nawrot ▪ FU Berlin ▪ SFA in the insect olfactory pathway

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Odor VALUE coding at mushroom body output

M Nawrot ▪ FU Berlin ▪ SFA in the insect olfactory pathway

► Reward prediction after ~ 140ms

Re-Coding : Odor IDENTIY (AL) Odor VALUE (MB)

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Neural performance correlates with behavioral performance

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FIN