tutorial for assc9 on new developments in eeg research walter j freeman university of california at...
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Tutorial for ASSC9 on new developments in
EEG research
Walter J FreemanUniversity of California at Berkeley
http://sulcus.berkeley.edu
“A movie, with its taut stream of thematically connected images, its visual narrative integrated by the viewpoint and values of its director, is not at all a bad metaphor for the stream of consciousness itself. .... The mechanism of our ordinary knowledge is of a cinematographical kind.”
New York Review, 2004Oliver Sacks
Questions regarding cinematographic frames
If the cinematographic hypothesis is valid:
• How many screens are there?
• Where are they, and how large?
• What are the frame rates, durations, sizes?
• How do frames form and sequence?
• What is the structure of their contents?
• How can EEG be used to answer the questions?
Part 1. Temporal analysisIntroduction to basic conceptsTemporal Fourier transform Gabor and Morlet wavelets
Hilbert transform
Part 2. Spatial analysis Design of electrode arrays, 1-D, 2-D
Spatial Fourier transformSpatial patterns of phase
Spatial patterns of amplitude
OUTLINE
Introduction to basic concepts:
LinearityStationarityGaussianity
Measurement Decomposition
Brains are neither linear, stationary nor Gaussian.
EEGs are usually linear, stationary and Gaussian.
This discrepancy must be dealt with explicitly.
Basic concepts
Linearity: Superposition
Additivity1. Outputs for multiple inputs are additive.
Proportionality2. Outputs are proportional to inputs.
Testing is by paired-shockusing single-shock electrical stimuli and averaging the cortical evoked potentials.
Superposition
Biedenbach & Freeman, 1965
Accompanying properties:
1. Output frequencies are the same as the input frequencies - no harmonics.
2. Gaussian amplitude distributions of input give Gaussian output amplitude distributions.
By this test, EEG has robust near-linear domains, provided that the amplitudes of evoked potentials do not exceed the maxima of background EEG.
Gaussianity
alpha
Gaussianity demonstrated by use of amplitude histograms.
normal density function
Amplitude histograms
Stationarity:
Frequencies don’t change with time.
Major state transitions reveal nonstationarity of cortical dynamics:
• waking vs. sleeping • seizure onset vs. offset
EEGs recorded during attentional shifts and cognitive activity reveal drifts and jumps in
frequencies that reflect nonstationarity.
Stationarity
Measurement
expresses some quantity of interest in numbers.
It requires a unit of the selected dimension:
• Time - sec
• Space - cm
• Magnitude - microvolt
• Phase - radian (1 rad = 360°/2= 57.3° )
• Wave form - basis function
Measurement
Decomposition
Upon measurement, the wave form of interest is said to be “decomposed” into the matching sum of
the selected basis functions.
The simplest basis function is the digitizing step.
A string of square waves is defined by the latency, the duration, and the number of squares
we add to match an EEG at each time step.
Decomposition
When EEGs conform to linearity and stationarity, they can be decomposed for measurement with linear basis functions:
Sines,Cosines,
Exponentials….
These basis functions are the solutions to linear differential equations.
Linear, time-invariant neurodynamics!
Family of linear basis functions
Fourier Transform: The inner product
Rodrigo Quian QuirogaSloan-Swartz Center for Theoretical Neurobiology
California Institute of Technologyhttp://www.vis.caltech.edu/~rodri
Brain oscillations decomposed in a semi-log plot of power spectral density (PSD) with classic bands.
Gabor Transform - Gaussian envelope, fixed duration, selected frequencies, moving window
Effect of the window duration on display of seizure onset
Morlet Wavelets - Gaussian envelope with fixed frequencies and fixed duration of moving window
Clin. Neurophysiol. 110: 643-654 (1999)Brain Research Protocols 8: 16-24 (2001)5.5 cycles
Multiresolution decomposition - calculation at discrete time steps and frequency scales, implemented by a
recursive filter bank - faster than the FFT!
Event-related alpha responses by decomposing visual EP in an oddball paradigm
Clin. Neurophysiol. 110: 643-654 (1999)
Linear analysis is easy to implement, and it tolerates wide deviations from linearity
and stationarity. Examples:
FFT, ARMA, Discriminant analysis, SVD, Karhunen-Loéve,
PCA, Factor analysis (Bayesian), ICA,Laplacian operators,
ERP, AER …
Why do these techniques work so well?Where and why do they fail?
Tools for linear analysis
Walter J Freeman University of California at Berkeley
Decomposition by FFT and wavelets requires the assumption that frequencies are discrete.
We can relax this assumption by using a coordinate transform to re-plot the power spectral
density (PSD):
log power vs. log frequency
The log-log plot indicates the existence of continuous distributions of frequencies.
An introduction to PSD in log-log coordinates
Temporal spectra from frontal scalp
From Freeman et al. 2003
EEG awake and asleep: intracranial recording from right superior temporal gyrus in epileptic patient.
PSDt of EEG
Slope = -3.46
delta
EEG awake intracranial
Slope = -2.21
alpha
beta
Histogram of power-law exponents - asleep
Histogram of power-law exponents - awake
EEG(t) and EMG(t)
500 ms
100 uv
Human scalp recordings, right paracentral
EEG PSDt in spatial spectral bands
Frontal scalpEyes closed
Decomposition of temporal spectrum by spatial pass band
Resting Slope = -2
EMG PSDt in spatial spectral bands
Frontal scalpEyes closed
EMG power covers all parts of the spectrum: white noise
EEG with strong alpha
Occipital scalp
Eyes closed
EMG with persisting alpha
Occipital scalp
Eyes closed
Walter J Freeman University of California at Berkeley
An introduction to 1/f PSDt and PSDx
The EEG spectrum tends to 1/f for amplitude, 1/f2 for power, except in sleep and before seizure.
The EMG spectrum tends to be flat - white noise.
The parameters of EEG appear to be fractal.
EEG frequencies are not fixed; they vary over a spectral continuum.
The Hilbert transform can address that property.
Walter J Freeman University of California at Berkeley
The Hilbert transformAn introduction to the Hilbert transform
At each time step the EEG gives the real part, Re(t), of a complex number. The Hilbert transform gives
the imaginary part, Im(t).
The complex number gives the analytic amplitude:
A(t) = [ Re(t)2 + Im(t)2 ] 0.5
and the analytic phase:
P(t) = atan [ Im(t) / Re(t) ]
The HT serves to decompose an EEG signal into independent functions of amplitude and phase.
Simulated EEG - 20 Hz cosine with phase slip
Polar plot, simulated EEG
analytic amplitude:A(t) = [ Re(t)2 + Im(t)2 ] 0.5
Calculate analytic phase and differences
analytic phase: P(t) = atan [ Im(t) / Re(t) ]
Calculate analytic phase and differences
“ Instantaneous” frequency is calculated by dividing each frequency difference by the digitizing step in s to give radians/s.Division by 2 gives frequency in Hz.
Walter J Freeman University of California at Berkeley
EEG from 8x8 pial array, rabbit auditory cortex.Spacing: 0.79 mm 1st component PCA: 94%
Digitizing step: 2 ms Nyquist frequency: 250 Hz
8x8 recording in waking rabbit
Walter J Freeman University of California at Berkeley
Subtract channel means; normalize globally to unit SD; FFT 64 EEGs in 500 ms; average PSDt
PSDt in waking rabbit
theta gamma
Time series
Rabbit auditory cortex; CS+ given at 400 ms
Polar plot
Elapsed time is shown by counterclockwise rotation.
Analytic amplitude
A(t) = [ Re(t)2 + Im(t)2 ] 0.5
Analytic phase
P(t) = atan [ Im(t) / Re(t) ]; p(t) = P(t) +
P(t)
p(t)+
-
Coordinated analytic phase differences (CAPD)
Time is on left abscissa, channel order is on right
Relation of amplitude to phase dispersion
Maximal amplitude occurs when the phase stabilizes.
Evidence for stationarity on average
This result can explain the success of Fourier analysis. Most of the time the rate of change in phase
(frequency) is quite low, implying stationarity.
But, epochs of phase constancy [frames] are bracketed by phase instabilities and discontinuities: phase slip.
The successive frames have different average rates of change in phase (frequencies), but they are similar
owing to band pass filtering and low rates of change.
The EEG can appear to be stationary, but it is not; the
analytic amplitude varies with the phase discontinuities.
Interim Conclusions
Interim Conclusions
The Hilbert transform reveals ‘phase slips’ within empirical EEG beta and gamma ranges.
These discontinuities in phase may demarcate state transitions by which cinematographic frames form.
Further evidence is needed to support the cinematographic hypothesis.
That evidence is in spatial analysis of the analytic amplitude of EEGs from multielectrode arrays:
Part 2. Spatial analysis
Kohler
Kohler’s Category Error
Roger Sperry
Disproof:Roger Sperry inserted silver needles or strips of mica into the visual
cortex of cats and monkeys that distorted
the electric fields. Visual perception was undiminished.
But – EEG fields are
epiphenomenal – not
Köhler’s Gestalt fields.