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Causality in Time Series - ClopiNet

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Causal analysis of fMRIthe SNR of BOLD fMRI <strong>in</strong> general is very good <strong>in</strong> comparison to electrophysiologicaltechniques like EEG and MEG.Although the balloon model and its variations have played an important role <strong>in</strong>describ<strong>in</strong>g the transient features of the fMRI response and <strong>in</strong>ferr<strong>in</strong>g neuronal activity,simplified ways of represent<strong>in</strong>g the BOLD signal responses are very often used. Mostprom<strong>in</strong>ent among these is a l<strong>in</strong>ear f<strong>in</strong>ite impulse response (FIR) convolution with asuitable kernel. The most used s<strong>in</strong>gle convolution kernel characteriz<strong>in</strong>g the ‘canonical’hemodynamic reponse is formed by a superposition of two gamma functions (Glover,1999), the first characteriz<strong>in</strong>g the <strong>in</strong>itial signal <strong>in</strong>crease, the second the later negativeundershoot (Figure 2B, solid l<strong>in</strong>e):h(t) = m 1 t τ 1e (−l 1t) − cm 2 t τ 2e (−l 2t)m i = max (︁ t τ ie (−l it) )︁ (6)With times-to-peak <strong>in</strong> seconds τ 1 = 6, τ 2 = 16, scale parameters l i (typically equal to 1)and a relative amplitude of undershoot to peak of c = 1/6.Often, the canonical two-gamma HRF kernel is augmented with one or two additionalorthogonalized convolution kernels: a temporal derivative and a dispersionderivative. Together the convolution kernels form a flexible basis function expansionof possible HRF shapes, with the temporal derivative of the canonical account<strong>in</strong>g forvariation <strong>in</strong> the response delay and the dispersion derivative account<strong>in</strong>g for variations <strong>in</strong>temporal response width (Henson et al., 2002; Liao et al., 2002). Thus, the l<strong>in</strong>ear basisfunction representation is a more abstract characterization of the HRF (i.e. further awayfrom the physiology) that still captures the possible variations <strong>in</strong> responses.It is an <strong>in</strong>terest<strong>in</strong>g property of hemodynamic processes that, although they are characterizedby a large overcompensat<strong>in</strong>g reaction to neuronal activity, their effects arehighly local. The locality of the hemodynamic reponse to neuronal activity limits theactual spatial resolution of fMRI. The path blood <strong>in</strong>flow <strong>in</strong> the bra<strong>in</strong> is from large arteriesthrough arterioles <strong>in</strong>to capillaries where exchange with neuronal tissue takes placeat a microscopic level. Blood outflow takes place via venules <strong>in</strong>to the larger ve<strong>in</strong>s. Thema<strong>in</strong> regulators of blood flow are the arterioles that are surrounded by smooth muscle,although arteries and capillaries are also thought to be <strong>in</strong>volved <strong>in</strong> blood flow regulation(Attwell et al., 2010). Different hemodynamic parameters have different spatialresolutions. While CBV and CBF changes <strong>in</strong> all compartments but mostly venules,oxygenation changes mostly <strong>in</strong> the venules and ve<strong>in</strong>s. Thus, the achievable spatial resolutionwith fMRI is limited by its specificity to the smaller arterioles and venules andmicroscopic capillaries supply<strong>in</strong>g the tissue, rather than the larger supply<strong>in</strong>g arteriesdra<strong>in</strong><strong>in</strong>g ve<strong>in</strong>s. The larger vessels have a larger doma<strong>in</strong> of supply or extraction and, as aconsequence, their signal is blurred and mislocalized with respect to active tissue. Here,physiology and physics <strong>in</strong>teract <strong>in</strong> an important way. It can be shown theoretically – bythe effects of thermal motion of sp<strong>in</strong> diffusion over time and the distance of the sp<strong>in</strong>sto deoxy-Hb – that the orig<strong>in</strong> of the BOLD signal <strong>in</strong> SE sequences at high ma<strong>in</strong> fieldstrengths (larger than 3T) is much more specific to the microscopic vasculature than tothe larger arteries and ve<strong>in</strong>s. This does not hold for GRE sequences or SE sequences81

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