|All electrical currents produce electromagnetic fields, and our body is inundated by currents of all sorts. The muscles and the heart are two well-known and strong sources of electrophysiological currents, qualified as ‘animal electricity’ by early scientists like Luigi Galvani, who were able to evidence such phenomena more than 200 years ago. The brain also sustains ionic current flows within and across cell assemblies, with neurons as the strongest generators. The architecture of the neural cell – as decomposed into dendritic branches and tree, soma and axon – conditions the paths taken by the tiny intracellular currents flowing within the cell. The relative complexity and large variety of these current pathways can be simplified by looking at the cell from some distance: indeed, these elementary currents instantaneously sum into a net primary current flow, which can be well described as a small, straight electrical dipole conducting current from a source to a sink.
Intracellular current sources are twofold in a neuron:
- axon potentials, which generate fast discharges of currents, and
- slower excitatory and inhibitory post-synaptic potentials (E/I PSPs), which create an electrical imbalance between the basal, apical dendritic tree and/or the cell soma.
Each of these two categories of current sources generates electromagnetic fields, which can be well captured by local electrophysiological recording techniques. The amount of current being generated by a single cell is however too small to be detected several centimeters away and outside the head. Detecting electrophysiological traces non invasively is conditioned to two main factors:
Fortunately, a great share of neural cells possesses a longitudinal geometry; these are the pyramidal cells in neocortical layers II/III and V. Also, neurons are grouped into assemblies of tightly interconnected cells. Therefore it is likely that PSPs be identically distributed across a given assembly, with the immediate benefit that they build-up efficiently to drive larger levels of currents, which in turn generate electromagnetic fields that are strong enough to be detected outside the head.
- that the architecture of the cell is propitious to give rise to a large net current, and
- that neighboring cells would drive their respective intracellular currents with a sufficient degree of group synchronization so that they build-up and reach levels detectable at some distance.
Illustration of the basic electrophysiological principles of MEG and EEG
Large neural cells – just like this pyramidal neuron from cortex layer V – drive ionic electrical currents. These latter are essentially impressed by the difference in electrical potentials between the basal and apical dendrites or the cell body, which is due to a blend of excitatory and inhibitory post-synaptic potentials (PSP), which are slow (>10 ms) relatively to axon potentials firing and therefore sum-up efficiently at the scale of synchronized neural ensembles. These primary currents can be modeled using an equivalent current dipole, here represented by a large black arrow. The electrical circuit of currents is closed within the entire head volume by secondary, volume currents shown with the dark plain lines. Additionally, magnetic fields are generated by the primary and secondary currents. The magnetic field lines induced by the primary currents are shown using dash lines arranged in circles about the dipole source.
Neurons in assemblies are also likely to fire volleys of action potentials with a fair degree of synchronization. However the very short duration of each action potential firing – typically a few milliseconds – makes it very unlikely that they sufficiently overlap in time to sum-up to a massive current flow. Though smaller in amplitude, PSPs sustain with typical durations – a few tens to hundreds of milliseconds – that make temporal and amplitude overlap build-up more efficiently within the cell ensemble.
Interestingly, though PSPs were thought originally to impress only rather slow fluctuations of currents, recent experimental and modeling evidence demonstrate they are capable of also generating fast spiking activity (Murakami & Okada, 2006). One might assume that these latter may be at the origins of the very high-frequency brain oscillations (that is, up to 1KHz) captured by MEG (Cimatti et al., 2007). Indeed, mechanisms of active ion channeling within dendrites would further contribute to larger amplitudes of primary currents than initially predicted (Murakami & Okada, 2006). Hence neocortical columns consisting of as few as 50,000 pyramidal cells with an individual current density of 0.2 pA.m, would induce a net current density of 10 nA.m at the assembly level. This is the typical source strength that can be detected using MEG and EEG. Other neural cell types, such as Purkinje and stellate cells are structured with less favorable morphology and/or density than pyramidal cells. It is therefore expected that their contribution to MEG/EEG surface signals is less than neocortical regions. Published models and experimental data however report regularly on the detection of cerebellar and deeper brain activity using MEG or EEG (Tesche, 1996, Jerbi et al., 2007, Attal et al., 2009).
Cellular currents are therefore the primary contributors to MEG/EEG surface signals. These current generators operate in a conductive medium and therefore impress a secondary type of currents that circulate through the head tissues (including the skull bone) and loop back to close the electrical circuit. Consequently, it is key to the methods attempting to localize the primary current sources to discriminate these latter from the contributions of secondary currents to the measurements. Modeling the electromagnetic properties of head tissues is critical in that respect. Before reviewing this important aspect of the MEG/EEG realm, we shall first discuss the basics of MEG/EEG instrumentation.
At a larger spatial scale, the mass effect of currents due to neural cells sustaining similar PSP mixtures add up locally and behave also as an current dipole (shown in red). This primary generator induces secondary currents (shown in yellow) that travel through the head tissues. They eventually reach the scalp surface where they can be detected using pairs of electrodes in EEG. Magnetic fields (in green) travel more freely within tissues and are less distorted than current flows. They can be captured using arrays of magnetometers in MEG. The distribution of blue and red colors on the scalp illustrates the continuum of magnetic and electric fields and potentials distributed at the surface of the head.