13
Electroencephalography
spontaneous activity,
evoked potentials, and
bioelectric events produced by single neurons.
Spontaneous activity is measured on the scalp or on the brain and is called the electroencephalogram. The amplitude of the EEG is about 100 µV when measured on the scalp, and about 1-2 mV when measured on the surface of the brain. The bandwidth of this signal is from under 1 Hz to about 50 Hz, as demonstrated in Figure 13.1. As the phrase "spontaneous activity" implies, this activity goes on continuously in the living individual.
The number of nerve cells in the brain has been estimated to be on the order of 1011. Cortical neurons are strongly interconnected. Here the surface of a single neuron may be covered with 1,000-100,000 synapses (Nunez, 1981). The electric behavior of the neuron corresponds to the description of excitable cells introduced in the earlier chapters. The resting voltage is around -70 mV, and the peak of the action potential is positive. The amplitude of the nerve impulse is about 100 mV; it lasts about 1 ms.
Evoked potentials are those components of the EEG that arise in response to a stimulus (which may be electric, auditory, visual, etc.) Such signals are usually below the noise level and thus not readily distinguished, and one must use a train of stimuli and signal averaging to improve the signal-to-noise ratio.
Single-neuron behavior can be examined through the use of microelectrodes which impale the cells of interest. Through studies of the single cell, one hopes to build models of cell networks that will reflect actual tissue properties.
13.2 THE BRAIN AS A BIOELECTRIC GENERATOR
PRECONDITIONS:
SOURCE: Distribution of impressed current source elements i (volume source)
CONDUCTOR: Finite, inhomogeneous
The bioelectric impressed current density i associated with neuronal activation produces an electric field, which can be measured on the surface of the head or directly on the brain tissue. The electric field was described by Equation 7.10 for a finite inhomogeneous model. This equation is repeated here:
(13.01) |
While for most excitable tissue the basis for the impressed current density i is the propagating action potential, for the EEG it appears to arise from the action of a chemical transmitter on postsynaptic cortical neurons. The action causes localized depolarization - that is, an excitatory postsynaptic potential (EPSP) - or hyperpolarization - that is, an inhibitory postsynaptic potential (IPSP). The result in either case is a spatially distributed discontinuity in the function sF (i.e., soFo - siFi) which, as pointed out in Equation 8.28, evaluates a double layer source in the membranes of all cells. This will be zero for resting cells; however, when a cell is active by any of the aforementioned processes (in which case Fo - Fi = Vm varies over a cell surface), a nonzero primary source will result.
For distant field points the double layer can be summed up vectorially, yielding a net dipole for each active cell. Since neural tissue is generally composed of a very large number of small, densely packed cells, the discussion in Section 8.5 applies, leading to the identification of a continuous volume source distribution i which appears in Equations 7.6 and 7.10.
Although in principle the EEG can be found from the evaluation of Equation 7.10, the complexity of brain structure and its electrophysiological behavior have thus far precluded the evaluation of the source function i. Consequently, the quantitative study of the EEG differs from that of the ECG or EMG, in which it is possible to evaluate the source function. Under these conditions the quantitative EEG is based on a statistical treatment, whereas the clinical EEG is largely empirical..
Fig. 13.1. Frequency spectrum of normal EEG.
Fig. 13.2. The international 10-20 system seen from (A) left and (B) above the head. A = Ear lobe, C = central, Pg = nasopharyngeal, P = parietal, F = frontal, Fp = frontal polar, O = occipital.
(C) Location and nomenclature of the intermediate 10% electrodes, as standardized by the American Electroencephalographic Society. (Redrawn from Sharbrough, 1991.).
Fig. 13.3. (A) Bipolar and (B) unipolar measurements. Note that the waveform of the EEG depends on the measurement location.
Fig. 13.4. Sensitivity distribution of EEG electrodes in the spherical head model. The figure illustrates the lead field current flow lines (thin solid lines), isosensitivity lines (dotted lines) and the half-sensitivity volumes (shaded region). The sensitivity distribution is in the direction of the flow lines, and its magnitude is proportional to the density of the flow lines. The lead pair are designated by small arrows at the surface of the scalp and are separated by an angle of 180°, 120°, 60°, 40°, and 20° shown at the top of each figure.
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