M/EEG source analysis Rik Henson MRC CBU, Cambridge (with thanks to Christophe Phillips, Jeremie Mattout, Gareth Barnes, Jean Daunizeau, Stefan Kiebel and Karl Friston) Overview 1. Forward Models for M/EEG 2. Variational Bayesian Dipole Estimation (ECD) 3. Empirical Bayesian Distributed Estimation 4. Multimodal integration Overview 1. Forward Models for M/EEG 2. Variational Bayesian Dipole Estimation (ECD) 3. Empirical Bayesian Distributed Estimation 4. Multimodal integration Bayesian Perspective Forward Problem m Model p(Y | , m) Likelihood Posterior Parameters p( | Y , m) p( | m) Prior Evidence p(Y | m) Inverse Problem Y Data Forward Problem: Physics Current density: j Orientation r Location Likelihood Y f ( j,r ) Quasi-static Maxwell’s Equations: Kirkoff’s law: j 0 Electrical potential E E E 0 B 0 B j Y Y B (EEG) (MEG) Forward Problem: Physics j Orientation Likelihood Y f ( j,r ) r Location f depends on: location (orientation) of sensors geometry of head conductivity of head (source space) Can have analytic or numerical form… Forward Problem: Head Models Concentric Spheres: Pros: Analytic; Fast to compute Cons: Head not spherical; Conductivity not homogeneous Boundary (or Finite) Element Models: Pros: Realistic geometry Homogeneous conductivity within boundaries Cons: Numeric; Slow Approximation Errors Other approaches (for MEG): Fit local spheres to each sensor; Single shell, spherical approx (Nolte) Forward Problem: Meshes 3 important surfaces for BEMs are those with large changes in conductivity: Scalp (skin-air boundary) Outer Skull (bone-skin boundary) Inner Skull (CSF-bone boundary) (Represented as tessellated triangular meshes) Extracting these surfaces from an MRI is difficult, eg, because CSF-bone T1-contrast is poor (use PD?)… A fourth important surface (for some solutions) is: Cortex (WM-GM boundary) Extracting this surface from an MRI is very difficult because so convoluted (though FreeSurfer)… Forward Problem: Canonical Meshes Rather than extract surfaces from individuals MRIs, why not warp Template surfaces from an MNI brain based on spatial (inverse) normalisation? Henson et al (2009), Neuroimage Recap: (Spatial Normalisation) fMRI time-series Anatomical MRI Template Smoothed Estimate Spatial Norm Motion Correct Smooth Coregister m11 m21 m31 0 m12 m22 m13 m23 m32 0 m33 0 Spatially normalised m14 m24 m34 1 Deformation Forward Problem: Canonical Meshes Rather than extract surfaces from individuals MRIs, why not warp Template surfaces from an MNI brain based on spatial (inverse) normalisation? Mattout et al (2007), Comp Int & Neuro Individual Canonical (Inverse-Normalised) Template “Canonical” (Also provides a 1-to-1 mapping across subjects, so source solutions can be written directly to MNI space, and group-inversion applied; see later) Given that surfaces are part of the forward model (m), can use the model evidence p(Y | m) to determine whether Canonical Meshes are sufficient Henson et al (2009), Neuroimage Forward Problem: ECD vs Distributed j Orientation r Location Likelihood Y f ( j,r ) For small number of Equivalent Current Dipoles(ECD) anywhere in brain: f is linear in j but non-linear in r Y f r j For (large) number of (Distributed) dipoles with fixed orientation and location: f is linear in r Y F r1 r2 rN J Overview 1. Forward Models for M/EEG 2. Variational Bayesian Dipole Estimation (ECD) 3. Empirical Bayesian Distributed Estimation 4. Multimodal integration Inverse Problem: VB-ECD Standard ECD approaches iterate location/orientation (within a brain volume) until fit to sensor data is maximised (i.e, error minimised). But: 1. Local Minima (particularly when multiple dipoles) 2. Question of how many dipoles? With a Variational Bayesian (VB) framework, priors can be put on the locations and orientations (and strengths) of dipoles (e.g, symmetry constraints) j r j Y f r j e r Y p(r , j , r , j , e | m) p(Y | r , j , e , m) p(e | m) p(r | r , m) p(r | m) p( j | j , m) p( j | m) Kiebel et al (2008), Neuroimage Inverse Problem: VB-ECD Maximising the (free-energy approximation to the) model evidence p(Y | m) offers a natural answer to question of the number of dipoles Kiebel et al (2008), Neuroimage Inverse Problem: DCM Dynamic Causal Modelling (DCM) can be seen as a source localisation (inverse) method that includes temporal constraints on the source activities David et al (2011), Journal of Neuroscience Overview 1. Forward Models for M/EEG 2. Variational Bayesian Dipole Estimation (ECD) 3. Empirical Bayesian Distributed Estimation 4. Multimodal integration Inverse Problem: Distributed Given p sources fixed in location (e.g, on a cortical mesh)… …linear Forward Model for MEG/EEG: Y = LJ + E E ~ N (0, C(e) ) Y = Data n sensors J = Sources p>>n sources L = Leadfields n sensors x p sources E = Error n sensors… …draw from Gaussian covariance C(e) (Free orientations can be simulated by having 2-3 columns in L per location) Fact that p>>n means under-determined problem (cf. GLM and ECD)… …so some form of regularisation needed, e.g,“Weighted L2-norm”… Inverse Problem: Standard L2-norm Y = LJ + E E ~ N (0, C(e) ) ( e ) 1 / 2 J arg min{C 2 (Y LJ) WJ } 2 (WT W)1 LT [L(WT W)1 LT C(e) ]1 Y “Tikhonov Solution” ||Y – LJ||2 “L-curve” method WI “Minimum Norm” W DDT = regularisation (hyperparameter) ||WJ||2 “Loreta” (D=Laplacian) W diag(LT L) 1 “Depth-Weighted” Wp diag(LTp C y1L p ) 1 “Beamformer” W Phillips et al (2002), Neuroimage Inverse Problem: Equivalent PEB Parametric Empirical Bayesian (PEB) 2-level hierarchical form: Y LJ Ε(e) E(e) ~ N (0, C(e) ) C(e) = n x n Sensor (error) covariance J 0 E( j ) E( j ) ~ N (0, C( j ) ) C(j) = p x p Source (prior) covariance Likelihood: p(Y | J) N (LJ, C(e) ) Prior: C( j ) p(J) N (0, C( j ) ) C (e ) J Posterior: p( J | Y ) p(Y | J) p(J) L Maximum A Posteriori (MAP) estimate: Jˆ C( j )LT [LC( j )LT C(e) ]1 Y Y cf Classical Tikhonov: (WT W)1 LT [L(WT W)1 LT C(e) ]1 Y C( j ) (WT W)1 Phillips et al (2005), Neuroimage Inverse Problem: Covariance Components (Priors) Specifying (co)variance components (priors/regularisation): C i Qi i C = Sensor/Source covariance Q = Covariance components λ = Hyper-parameters ( e) Empty-room: # sensors “IID” (white noise): # sensors 1. Sensor components, Qi (error): # sensors # sensors ( j) Multiple Sparse Priors (MSP): # sources # sources “IID” (min norm): # sources 2. Source components, Qi (priors/regularisation): # sources Friston et al (2008) Neuroimage Inverse Problem: HyperPriors When multiple Q’s are correlated, estimation of hyperparameters λ can be difficult (eg local maxima), and they can become negative (improper for covariances) αi ln(λi ) i exp(i ) 2) impose weak, shrinkage hyperpriors: p(α) ~ N ( η, Ω) η 4 Ω aI, a 16 Anti-Averaging Smoothness Depth-Weighting Sensor priors 1) impose positivity on hyperparameters: Prestim Baseline Source priors projected to sensors To overcome this, one can: uninformative priors are then “turned-off” (cf. “Automatic Relevance Detection”) 0 Henson et al (2007) Neuroimage Inverse Problem: HyperPriors When multiple Q’s are correlated, estimation of hyperparameters λ can be difficult (eg local maxima), and they can become negative (improper for covariances) To overcome this, one can: 1) impose positivity on hyperparameters: αi ln(λi ) i exp(i ) 2) impose weak, shrinkage hyperpriors: p(α) ~ N ( η, Ω) η 4 Ω aI, a 16 uninformative priors are then “turned-off” (cf. “Automatic Relevance Detection”) 0 Henson et al (2007) Neuroimage Inverse Problem: Full (DAG) model Source and sensor space η, Ω Q1( j ) Q(2j ) ... Q1(e) Q(2e) i( e ) i( j ) C( j ) Fixed ... C( e) Ε J Variable Data L Y Friston et al (2008) Neuroimage Inverse Problem: Estimation 1. Obtain Restricted Maximum Likelihood (ReML) estimates of the hyperparameters (λ) by maximising the variational “free energy” (F): λˆ max p(Y | λ ) max F 2. Obtain Maximum A Posteriori (MAP) estimates of parameters (sources, J): Jˆ max p (J | Y, λˆ ) max F j j 3. Maximal F approximates Bayesian (log) “model evidence” for a model, m: ˆ) ln p(Y | m) ln p(Y, J, λ | m) dJdλ F (Y, αˆ , Σ m {L, Q, η, Ω} ˆ ) tr(C1YYT ) ln | C | (αˆ - η)T Ω1 (αˆ - η) ln | ΣΩ ˆ 1 | F (Y, αˆ , Σ Accuracy Complexity (…where αˆ and Σˆ are the posterior mean and covariance of hyperparameters) Friston et al (2002) Neuroimage Inverse Problem: Multiple Sparse Priors # sources Hyperpriors allow the extreme of 100’s source priors, or MSP # sources Left patch … Right patch … Q(2)j Bilateral patches … Q(2)j+1 … Q(2)j+2 Friston et al (2008) Neuroimage Inverse Problem: Multiple Sparse Priors Hyperpriors allow the extreme of 100’s source priors, or MSP Friston et al (2008) Neuroimage Inverse Problem: PEB Summary Summary: • Automatically “regularises” in principled fashion… • …allows for multiple constraints (priors)… • …to the extent that multiple (100’s) of sparse priors possible (MSP)… • …(or multiple error components or multiple fMRI priors)… • …furnishes estimates of model evidence, so can compare constraints Overview 1. Forward Models for M/EEG 2. Variational Bayesian Dipole Estimation (ECD) 3. Empirical Bayesian Distributed Estimation 4. Multi-modal and multi-subject integration Multi-subject Integration (Group Inversion) Specifying (co)variance components (priors/regularisation): C i Qi i C = Sensor/Source covariance Q = Covariance components λ = Hyper-parameters (e) Empty-room: # sensors “IID” (white noise): # sensors 1. Sensor components, Qi (error): # sensors # sensors ( j) Multiple Sparse Priors (MSP): # sources # sources “IID” (min norm): # sources 2. Source components, Qi (priors/regularisation): # sources Friston et al (2008) Neuroimage Multi-subject Integration (Group Inversion) Specifying (co)variance components (priors/regularisation): C i Qi i C = Sensor/Source covariance Q = Covariance components λ = Hyper-parameters (e) Empty-room: # sensors # sensors “IID” (white noise): # sensors 1. Sensor components, Qi (error): # sensors ( j) # sources 2. Optimise Multiple Sparse Priors by pooling across subjects Qi # sources Litvak & Friston (2008) Neuroimage Multi-subject Integration (as before) Source and sensor space η, Ω Q1( j ) ... Q1(e) C( j ) Fixed i( j ) i( e ) J Ε ... C(1) Variable Data L Y Litvak & Friston (2008) Neuroimage Multi-subject Integration Source and sensor space η, Ω Q1( j ) ... ( e) Q11 λ1( e ) λ( j) C( j ) Ε1 J1 Fixed ... Q(21e) λ (2e ) C1(1) ... C(1) 2 Ε2 J2 Variable Data L1 Y1 Y2 L2 Litvak & Friston (2008) Neuroimage Multi-subject Integration: Leadfield Alignment Concatenate data across subjects J1 A1Y1 , , A s Ys A1L1 , , A s L s E1(1) , , E(1) s J s …having projected to an “average” leadfield matrix A i Li L : L A i Li Common source-level priors: C( j ) k( j )Q(k j ) i s.t.: Ai max arg | LLT | : tr (LLT ) n Subject-specific sensor-level priors: Ci(e) ik(e) AiQ(ke) ATi Litvak & Friston (2008) Neuroimage Multi-subject Integration: Results MMN MSP MSP (Group) Litvak & Friston (2008) Neuroimage Multi-modal Integration 1. Symmetric integration (fusion) of MEG + EEG 2. Asymmetric integration of M/EEG + fMRI 3. Full fusion of M/EEG + fMRI? Multi-modal Integration “Neural” Activity Causes (hidden): (inversion) Generative (Forward) Models: Data: fMRI Balloon Model Head Head Model Model MEG ? EEG ? (future) Daunizeau et al (2007), Neuroimage Multi-modal Integration “Neural” Activity Causes (hidden): Generative (Forward) Models: Data: Balloon Model fMRI Head Head Model Model MEG ? EEG Symmetric Integration (Fusion) ? (future) Asymmetric Integration Daunizeau et al (2007), Neuroimage Multi-modal Integration 1. Symmetric integration (fusion) of MEG + EEG 2. Asymmetric integration of M/EEG + fMRI 3. Full fusion of M/EEG + fMRI? Symmetric Integration of MEG+EEG Specifying (co)variance components (priors/regularisation): C i Qi i C = Sensor/Source covariance p(X) N (m, C) Q = Covariance components λ = Hyper-parameters ( e) Empty-room: # sensors “IID” (white noise): # sensors 1. Sensor components, Qi (error): # sensors # sensors ( j) Multiple Sparse Priors (MSP): # sources # sources “IID” (min norm): # sources 2. Source components, Qi (priors/regularisation): # sources Friston et al (2008) Neuroimage Symmetric Integration of MEG+EEG Specifying (co)variance components (priors/regularisation): C Q (e) i ( e) ji ( e) ij j Ci(e) = Sensor error covariance for ith modality Qij = jth component for ith modality λij = Hyper-parameters ( e) Q21(e) # sensors # sensors E.g, white noise for 2 modalities: Q11(e) # sensors 1. Sensor components, Qij (error): # sensors ( j) Multiple Sparse Priors (MSP): # sources # sources “IID” (min norm): # sources 2. Source components, Qi (priors/regularisation): # sources Henson et al (2009) Neuroimage Single Modality (as before) η, Ω Qi(2) ... Qi(1) i(1) i(2) C(2) ... C(1) Ε1 J Fixed Source and sensor space Variable L Data Y Henson et al (2009) Neuroimage Multiple modalities η, Ω Qi(2) ... Q1i(1) λ1(1) λ (2) C(2) ... Q(1) 2i λ (1) 2 ... C(1) 2 Ε2 Ε1 J Fixed C1(1) Source and sensor space Variable L1 Data Y1 Y2 L2 Henson et al (2009) Neuroimage Symmetric Integration of MEG+EEG • Stack data and leadfields for d modalities: Y1 L1 E1(1) (1) Y2 L2 J E2 (1) Ed Yd Ld C (e) C1( e ) 0 0 0 0 0 Cd( e ) 0 C2( e ) (note: common sources and source priors, but separate error components) • Where data / leadfields scaled to have same average / predicted variance: Yi Yi 1 mi T tr (YiYi ) Li Li 1 mi T i i tr ( L L ) mi = Number of spatial modes (e.g, ~70% of #sensors) Henson et al (2009) Neuroimage Symmetric Integration of MEG+EEG ERs from 12 subjects for 3 simultaneously-acquired Neuromag sensor-types: (Planar) Gradiometers (MEG, 204) Electrodes (EEG, 70) μV fT RMS fT/m Magnetometers (MEG, 102) Faces Scrambled ms ms ms Faces - Scrambled 150-190ms Henson et al (2009) Neuroimage Symmetric Integration of MEG+EEG +31 -51 -15 MEG mags MEG grads +19 -48 -6 Faces Scrambled Faces – Scrambled,150-190ms EEG +43 -67 -11 IID noise for each modality; common MSP for sources (fixed number of spatial+temporal modes) FUSED +44 -64 -4 Henson et al (2009) Neuroimage Symmetric Integration of MEG+EEG • Fusing magnetometers, gradiometers and EEG increased the conditional precision of the source estimates relative to inverting any one modality alone (when equating number of spatial+temporal modes) • The maximal sources recovered from fusion were a plausible combination of the ventral temporal sources recovered by MEG and the lateral temporal sources recovered by EEG • (Simulations show the relative scaling of mags and grads agrees with empty-room data) Henson et al (2009) Neuroimage Multi-modal Integration 1. Symmetric integration (fusion) of MEG + EEG 2. Asymmetric integration of M/EEG + fMRI 3. Full fusion of M/EEG + fMRI? Asymmetric Integration of M/EEG+fMRI Specifying (co)variance components (priors/regularisation): C i Qi i C = Sensor/Source covariance p(X) N (m, C) Q = Covariance components λ = Hyper-parameters (e) Empty-room: # sensors “IID” (white noise): # sensors 1. Sensor components, Qi (error): # sensors # sensors ( j) Multiple Sparse Priors (MSP): # sources # sources “IID” (min norm): # sources 2. Source components, Qi (priors/regularisation): # sources Friston et al (2008) Neuroimage Asymmetric Integration of M/EEG+fMRI Specifying (co)variance components (priors/regularisation): C i Qi i C = Sensor/Source covariance p(X) N (m, C) Q = Covariance components λ = Hyper-parameters (e) Empty-room: # sensors “IID” (white noise): # sensors 1. Sensor components, Qi (error): # sensors # sensors ( j) fMRI Priors: # sources # sources “IID” (min norm): # sources 2. Each suprathreshold fMRI cluster becomes a separate prior Qi # sources Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI Source and sensor space η, Ω Q1( j ) Q(2j ) ... Q1(e) Q(2e) i( e ) i( j ) C( j ) Fixed ... C(1) Ε J Variable Data L Y Friston et al (2008) Neuroimage Asymmetric Integration of M/EEG+fMRI η, Ω Y fMRI Q1( j ) Q(2j ) Source and sensor space ... Q1(e) Q(2e) i( e ) i( j ) C( j ) Fixed ... C(1) Ε J Variable Data L Y M / EEG Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI T1-weighted MRI Anatomical data {T,F,Z}-SPM Functional data … 1. Thresholding and connected component labelling Gray matter segmentation Cortical surface extraction … 2. Projection onto the cortical surface using the Voronoï diagram … 3D geodesic Voronoï diagram 3. Prior covariance components Qi( j ) Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI 1 2 4 5 SPM{F} for faces versus scrambled faces, 15 voxels, p<.05 FWE 3 5 clusters from SPM of fMRI data from separate group of (18) subjects in MNI space Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI Negative Free Energy (a.u.) (model evidence) Magnetometers (MEG) * * * * Gradiometers (MEG) * * * * Electrodes (EEG) * None * * Global Local (Valid) Local (Invalid) Valid+Invalid (binarised, variance priors) Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI Negative Free Energy (a.u.) (model evidence) Magnetometers (MEG) * * * * Gradiometers (MEG) * * * * Electrodes (EEG) * None * * Global Local (Valid) Local (Invalid) Valid+Invalid (binarised, variance priors) Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI Negative Free Energy (a.u.) (model evidence) Magnetometers (MEG) * * * * Gradiometers (MEG) * * * * Electrodes (EEG) * None * * Global Local (Valid) Local (Invalid) Valid+Invalid (binarised, variance priors) Henson et al (2010) Hum. Brain Map. 3.2 Fusion of MEG+fMRI (Application) Negative Free Energy (a.u.) (model evidence) Magnetometers (MEG) * * * * Gradiometers (MEG) * * * * Electrodes (EEG) * None * * Global Local (Valid) Local (Invalid) Valid+Invalid (binarised, variance priors) Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI Negative Free Energy (a.u.) (model evidence) Magnetometers (MEG) * * * * Gradiometers (MEG) * * * * Electrodes (EEG) * None * * Global Local (Valid) Local (Invalid) Valid+Invalid (binarised, variance priors) Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI IID sources and IID noise (L2 MNM) Magnetometers (MEG) Gradiometers (MEG) Electrodes (EEG) None Global Local (Valid) Local (Invalid) Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI IID sources and IID noise (L2 MNM) Magnetometers (MEG) Gradiometers (MEG) Electrodes (EEG) None Global Local (Valid) Local (Invalid) Henson et al (2010) Hum. Brain Map. 3.2 Fusion of MEG+fMRI (Application) IID sources and IID noise (L2 MNM) Magnetometers (MEG) Gradiometers (MEG) Electrodes (EEG) None Global Local (Valid) Local (Invalid) fMRI priors counteract superficial bias of L2-norm Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI IID sources and IID noise (L2 MNM) Magnetometers (MEG) Gradiometers (MEG) Electrodes (EEG) None Global Local (Valid) Local (Invalid) fMRI priors counteract superficial bias of L2-norm Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI Differential Response (Faces vs Scrambled) Right Posterior Fusiform (rPF) +26 -76 -11 Right Medial Fusiform (rMF) Right Lateral Fusiform (rLF) +32 -45 -12 +41 -43 -24 Gradiometers (MEG) (5 Local Valid Priors) L Differential Response (Faces vs Scrambled) R Differential Response (Faces vs Scrambled) Left occipital pole (lOP) -27 -93 0 Left Lateral Fusiform (lLF) -43 -47 -21 NB: Priors affect variance, not precise timecourse… Henson et al (2010) Hum. Brain Map. Asymmetric Integration of M/EEG+fMRI • Adding a single, global fMRI prior increases model evidence • Adding multiple valid priors increases model evidence further Helpful if some fMRI regions produce no MEG/EEG signal (or arise from neural activity at different times) • Adding invalid priors does not necessarily increase model evidence, particularly in conjunction with valid priors • Can counteract superficial bias of, e.g, minimum-norm • Affects variance but not not precise timecourse Henson et al (2010) Hum. Brain Map. Multi-modal Integration 1. Symmetric integration (fusion) of MEG + EEG 2. Asymmetric integration of M/EEG + fMRI 3. Full fusion of M/EEG + fMRI? Fusion of fMRI and MEG/EEG? “Neural” Activity Causes (hidden): Fusion of fMRI + MEG/EEG? Data: fMRI Balloon Model Head Head Model Model MEG ? EEG ? (future) Henson (2010) Biomag Fusion of fMRI and MEG/EEG? η, Ω Qi(2) Qi(1) ... i(2) C(2) Fixed Source and sensor space i(1) ... C(1) Ε1 J Variable Data Y L1 M / EEG Henson Et Al (2011) Frontiers Fusion of fMRI and MEG/EEG? η, Ω Qi(0) C(0) Fixed ... Qi(2) i(0) Qi(1) ... i(2) C(2) Ε0 Source and sensor space i(1) ... C(1) Ε1 J Variable L0 Data Y fMRI Y L1 M / EEG Henson Et Al (2011) Frontiers Overall Conclusions 1. SPM offers standard forward models (via FieldTrip)… (though with unique option of Canonical Meshes) 2. …but offers unique Bayesian approaches to inversion: 2.1 Variational Bayesian ECD 2.2 Dynamic Causal Modelling (DCM) 2.3 A PEB approach to Distributed inversion (eg MSP) 3. PEB framework in particular offers multi-subject and (various types of) multi-modal integration The End Forward Problem: Physics Current (nA): j Orientation r Location Likelihood Y f ( j,r ) Maxwell’s Equations: E B 0 B E t E B j t Ohm’s law: j E Continuity equation: j t Inverse Problem: Simulations Multiple constraints: Smooth sources (Qs), plus valid (Qv) or invalid (Qi) focal prior Qs Qs Qs,Qv 500 simulations Qs,Qi Qs,Qi,Qv 500 simulations Qv Qi Mattout et al (2006) Inverse Problem: Simulations Multiple constraints: Smooth sources (Qs), plus valid (Qv) or invalid (Qi) focal prior Log-Evidence Qs Bayes Factor Qs 205.2 7047 Qs,Qv 214.1 Qs,Qv,Qi 214.7 (Qs,Qi) 204.9 Qv 1.8 (1/9899) Qi Mattout et al (2006) Inverse Problem: Temporal ~ Y LJ E E ~ N (0,V C ) J ~ N (0,V ( j ) C ( j ) ) ( e) ( e) ~ C(e) = spatial error covariance over sensors V(e)= temporal error covariance over sensors C(j) = spatial error covariance over sources V(j) = temporal error covariance over sources In general, temporal correlation of signal (sources) and noise (sensors) will differ, but can project onto a temporal subspace (via S) such that: S TVe S S TVj S S TVS V typically Gaussian autocorrelations… V KK T (i j ) 2 K ( )ij exp 2 2 ~ 4m s then turns out that EM can simply operate on prewhitened data (covariance), where Y size n x t: ˆ EM ( 1 YS ( S T VS ) 1 S T Y T , Q) Nr Jˆ MYSST Friston et al (2006) Inverse Problem: Temporal Friston et al (2006) 3.2. Fusion of MEG+fMRI Gradiometers (MEG) Electrodes (EEG) Local Valid ln(λ)+32 fMRI hyperparameters ln(λ)+32 Magnetometers (MEG) Local Invalid Henson et al (2010) Multi-subject Integration: Results MMN + 3 fMRI priors MMN + 3 fMRI priors (Group) Henson et al (2011) Frontiers