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Appendix B

Application of Maxwell's
Equations in Bioelectromagnetism


The behavior of time-varying and static electric and magnetic fields are governed by Maxwell's equations formulated by James Clerk Maxwell (1865; 1873). These equations simply summarize the mathematical consequences of the classical experiments of Faraday, Ampere, Coulomb, Maxwell, and others.
Maxwell's equations can be found in general texts on electromagnetic theory. However, they are essentially applicable to electromagnetic fields in free space (i.e., radiation fields). Where conducting and/or magnetic media are involved, then, although the equations continue to be valid, current sources can arise in other ways than specified under free space conditions. These modifications must be introduced through a consideration of the particular nature of current sources appropriate for the problem at hand.
Our goal here, after introducing Maxwell's equations in the form valid for free space conditions, is to specialize them so that they correctly describe conditions that arise in bioelectromagnetism. Following this, our goal is to simplify the equations where possible, based on practical electrophysiological considerations.


SOURCES and FIELDS: Time-varying , ρ, ,
CONDUCTOR: Infinite, homogeneous free space σ = 0, = 0, ε = ε0

Maxwell's equations are usually written in differential (and vector) form for free space conditions as follows, where for simplicity a harmonic time variation is assumed:


Equation B.1 is a statement of Faraday's law that a time-varying magnetic field induces an electric field .
Equation B.2 is a statement of Ampere's law that the line integral of magnetic field around a closed loop equals the total current through the loop. The current is described as a displacement current jωε0 plus source currents arising from the actual convection of charge in a vacuum.
Equation B.3 arises from Coulomb's law and relates the electric displacement to the sources that generate it, namely the charge density ρ.
Equation B.4 is a statement of the conservation of charge, namely that its outflow from any closed region (evaluated from ) can arise only if the charge contained is depleted.
Equation B.5 recognizes that no magnetic charges exist, and hence the magnetic induction , must be solenoidal.


SOURCES and FIELDS: Static or quasistatic emf, i, ,
CONDUCTOR: Finite, inhomogeneous sσ = σ(x,y,z), = 0, ε = ε0

Our interest lies in describing electric and magnetic fields within and outside electrophysiological preparations. Electrophysiological preparations are isolated regions (lying in air) that involve excitable tissue surrounded by a conducting medium (volume conductor). The conductivity σ of the volume conductor, in general, is a function of position [σ(x,y,z)]; that is, it is assumed to be inhomogeneous. Its magnetic permeq/eability is normally assumed to be that of free space (0), and, except for a membrane region the dielectric permittivity also has the free space value (ε0).
If we consider for the moment a static condition, then we find that Equation B.1 requires that = 0. This means that must be conservative, a condition that is appropriate for electric fields arising from static charges in free space (i.e., electrostatics). But in our conducting medium, currents can flow only if there are nonconservative sources present. So we must assume the existence of electromotive forces emf. Thus for conducting media, Equation B.1 must be modified to the form of Equation B.6.
By the same reasoning, we must also recognize the presence of impressed (applied) current fields, which we designate i; these must be included on the right side of Equation B.7, which corresponds to Equation B.2 as applied to conducting media. Such sources may be essentially time-invariant as with an electrochemical battery that supplies an essentially steady current flow to a volume conductor. They may also be quasistatic, as exemplified by activated (excitable) tissue; in this case, time-varying nonconservative current sources result which, in turn, drive currents throughout the surrounding volume conductor.
In a conducting medium there cannot be a convection current such as was envisaged by the parameter i in Equation B.2, and it is therefore omitted from Equation B.7. The convection current is meant to describe the flow of charges in a vacuum such as occurs in high-power amplifier tubes. (For the same reason, Equation B.4 is not valid in conducting media.) In the consideration of applied magnetic fields, one can treat the applied current flowing in a physical coil by idealizing it as a free-space current, and hence accounting for it with the i on the right side of Equation B.2. Since this current is essentially solenoidal, there is no associated charge density. In this formalism the means whereby i is established need not be considered explicitly.
Because of the electric conductivity σ of the volume conductor we need to include in the right side of Equation B.7 the conduction current σ, in addition to the existing displacement current jωε.
Another modification comes from the recognition that a volume charge density ρ cannot exist within a conducting medium (though surface charges can accumulate at the interface between regions of different conductivity - essentially equivalent to the charges that lie on the plates of a capacitor). Therefore, Equation B.3 is not applicable in conducting media.
With these considerations, Maxwell's equations may now be rewritten for finite conducting media as


In this set of equations, we obtain Equation B.8 by taking the divergence of both sides of Equation B.7 and noting that the divergence of the curl of any vector function is identically zero 0.


SOURCES and FIELDS: Quasistatic (ω < 1000 Hz) emf, i, ,
CONDUCTOR: Limited finite (r < 1 m) inhomogeneous resistive (ωε/σ < 0.15) = 0, ε = ε0

Physiological preparations of electrophysiological interest have several characteristics on which can be based certain simplifications of the general Maxwell's equations. We have already mentioned that we expect the permittivity ε and permeq/eability in the volume conductor to be those of free space (ε0, 0). Three other conditions will be introduced here.

B.4.1 Frequency Limit

The power density spectra of signals of biological origin have been measured. These have been found to vary depending on the nature of the source (e.g., nerve, muscle, etc.). The highest frequencies are seen in electrocardiography. Here the bandwidth for clinical instruments normally lies under 100 Hz, though the very highest quality requires an upper frequency of 200-500 Hz. In research it is usually assumed to be under 1000 Hz, and we shall consider this the nominal upper frequency limit. Barr and Spach (1977) have shown that for intramural cardiac potentials frequencies as high as 10 kHz may need to be included for faithful signal reproduction. When one considers that the action pulse rise time is on the order of 1 ms, then signals due to such sources ought to have little energy beyond 1 kHz. Relative to the entire frequency spectrum to which Maxwell's equations have been applied, this is indeed a low-frequency range. The resulting simplifications are described in the next section.

B.4.2 Size Limitation

Except for the very special case where one is studying, say, the ECG of a whale, the size of the volume conductor can be expected to lie within a sphere of radius of 1 m. Such a sphere would accommodate almost all intact human bodies, and certainly typical in vitro preparations under study in the laboratory. A consequence, to be discussed in the next section, is that the "retarded" potentials of general interest do not arise.

B.4.3 Volume Conductor Impedance

The volume conductor normally contains several discrete elements such as nerve, muscle, connective tissue, vascular tissue, skin, and other organs. For many cases, the conducting properties can be described by a conductivity σ(x,y,z) obtained by averaging over a small but multicellular region. Since such a macroscopic region contains lipid cellular membranes the permittivity may depart from its free-space value. The values of both σ and ε entering Equations B.7 and B.8 will depend on the particular tissue characteristics and on frequency. By making macroscopic measurements, Schwan and Kay (1957) determined that ωε/σ for the frequency range 10 Hz < f < 1000 Hz is under 0.15. But in many cases it is possible to treat all membranes specifically. In this case it is the remaining intracellular and interstitial space that constitutes the volume conductor; and, since the lipids are absent, the medium will behave resistively over the entire frequency spectrum of interest. In either case it is reasonable to ignore the displacement current jωε0 within the volume conductor in Equations B.7 and B.8. (One should always, of course, include the capacitive membrane current when considering components of the total membrane current.) Consequently, these equations can be simplified to Equations B.10 and B.11, respectively.
Thus Maxwell's equations for physiological applications have the form:



SOURCE: Quasistatic i (ω < 1000 Hz)
CONDUCTOR: Limited finite (r < 1 m) region outside the sources inhomogeneous resistive (ωε/σ < 0.15), = 0, ε = ε0

In this section we derive from Maxwell's equations the equations for magnetic vector potential and electric scalar potential Φ in physiological applications, Equations B.19 and B.21, respectively.
Since the divergence of is identically zero (Equation B.9), the magnetic field may be derived from the curl of an arbitrary vector field , which is called the magnetic vector potential. This fulfills the requirement stated in Equation B.9 because the divergence of the curl of any vector field is necessarily zero. Consequently,


Since = μ0, we can substitute Equation B.12 into Equation B.6. We consider only the volume conductor region external to the membranes where the emfs are zero (note that the emfs are explicitly included within the membrane in the form of Nernst potential batteries), and we consequently obtain


Now, when the curl of a vector field is zero, that vector field can be derived as the (negative) gradient of an arbitrary scalar potential field (which we designate with the symbol Φ and which denotes the electric scalar potential). This assignment is valid because the curl of the gradient of any scalar field equals zero. Thus Equation B.13 further simplifies to


According to the Helmholtz theorem, a vector field is uniquely specified by both its divergence and curl (Plonsey and Collin, 1961). Since only the curl of the vector field has been specified so far (in Equation B.12), we may now choose


This particular choice eliminates Φ from the differential equation for (Equation B.17). That is, it has the desirable effect of uncoupling the magnetic vector potential from the electric scalar potential Φ. Such a consideration was originally suggested by Lorentz when dealing with the free-space form of Maxwell's equations. Lorentz introduced an equation similar to Equation B.15 known as the Lorentz condition, which is that


We have modified this expression since we have eliminated in Equations B.10 and B.11 the displacement current jωε in favor of a conduction current σ. This amounts to replacing jωε by σ in the classical Lorentz condition (Equation B.16), resulting in Equation B.15. The Lorentz condition can also be shown to have another important property, namely that it ensures the satisfaction of the continuity condition.
Now, if we substitute Equations B.12, B.14, and B.15 into Equation B.10, keeping in mind that = 0, and if we use the vector identity that


we obtain


Just as emfs were eliminated by confining attention to the region external to the excitable cell membranes, so too could one eliminate the nonconservative current i in Equation B.10. In this case all equations describe conditions in the passive extracellular and intracellular spaces; the effect of sources within the membranes then enters solely through boundary conditions at and across the membranes. On the other hand, it is useful to retain i as a distributed source function in Equation B.10. While it is actually confined to cell membranes ensuring the aforementioned boundary conditions, it may be simplified (averaged) and regarded as an equivalent source that is uniformly distributed throughout the "source volume." For field points outside the source region which are at a distance that is large compared to cellular dimensions (over which averaging of i occurs) the generated field approaches the correct value.
Equation B.18 is known as the vector Helmholtz equation, for which solutions in integral form are well known in classical electricity and magnetism (Plonsey and Collin, 1961). Adapting such a solution to our specific equation gives



Note that r is the radial distance from a source element dV(x,y,z) (unprimed coordinates) to the field point P(x',y',z') (primed coordinates), and is thus a function of both the unprimed and primed coordinates.
To evaluate an upper bound to the magnitude of kr in the exponential terms in Equation B.19 we choose:


Since e-.04 = .96, these exponential terms can be ignored and we get a simplification for Equation B.19, giving the magnetic vector potential under electrophysiological conditions:


The electric scalar potential Φ may be found from by using Equation B.15 with Equation B.20. In doing so, we note that Equation B.20 involves an integration over the source coordinates (x,y,z) while Equation B.15 involves operations at the field coordinates (x',y',z'). Consequently, we get


where ' operates only on the field coordinates, which is why i is not affected. Since '(1/r) = –(1/r), we finally get for the electric scalar potential:


Equation B.22 is identical to static field expressions for the electric field, where i is interpreted as a volume dipole density source function. This equation corresponds exactly to Equation 7-5. Although a staticlike equation applies, i is actually time-varying, and consequently, so must Φ be time-varying synchronously. We call this situation a quasistatic one.
When the source arises electrically (including that due to cellular excitation), a magnetic field is necessarily set up by the resulting current flow. The latter gives rise to a vector potential , which in turn contributes to the resulting electric field through the term jω in Equation B.14. However, under the conditions specified, |ω| is negligible compared to the term |Φ| as discussed in Plonsey and Heppner (1967). Under these conditions we are left with the scalar potential term alone, and Equation B.14 simplifies to


which also corresponds to a static formulation. It should be kept in mind that Equation B.23 is not exact, but only a good approximation. It corresponds to the quasistatic condition where the electric field resembles that arising under static conditions. Under truly static conditions the electric and magnetic fields are completely independent. Under quasistatic conditions, while the fields satisfy static equations, a low frequency time variation may be superimposed (justified by the low frequency conditions discussed earlier), in which case the magnetic field effects, although extant, can normally be ignored.
Note that in this case, where the sources are exclusively bioelectric and the simplification of Equation B.23 is valid, Equation B.11 leads to Equation 7.2 ( = i – σΦ).


B.6.1 Stimulation with Electric Field

SOURCE: Steady-state electric field
CONDUCTOR: Uniform fiber in volume conductor

The above comments notwithstanding, we are also interested in a situation where excitable tissue is stimulated solely with an applied magnetic field. In this case the vector potential is large and cannot be ignored. In fact, to ignore under these circumstances is to drop the underlying forcing function, which would leave an absurd result of no field, either electric or magnetic.
We have shown in Chapter 3 that for a single uniform fiber under steady-state conditions a homogeneous partial differential equation (Equation 3.46) arises:


where   Vm = transmembrane potential
 λ = space constant, characteristic of the physical and electric properties of the fiber
 x = coordinate along the direction of the fiber

For a point source at the origin we have also essentially shown, in Chapter 3, that the solution to Equation B.24 is (Equation 3.49)


where V'(x) = deviation of the membrane voltage from the resting voltage.

In this equation


where   Vm(0) = transmembrane potential at the origin
 I0 = applied intracellular point current
 ri = intracellular axial resistance per unit length

We remark, here, that for a more general applied scalar potential field, Φe, Equation B.24 becomes


One can recognize in this equation that the second derivative of the applied potential field along the fiber is the forcing function (in fact, it has been called the "activating function"), whereas the dependent variable, Vm, is the membrane response to the stimulation. Using Equation B.23, one can write Equation B.24 as


where is the applied electric field.

B.6.2 Stimulation with Magnetic Field

SOURCE: Time-varying magnetic field
CONDUCTOR: Uniform fiber in volume conductor

Electric stimulation may be produced by applying a time-varying magnetic field to the tissue. As given in Equation B.12, this magnetic field is defined as the curl of a vector potential . Now the stimulus is introduced solely through a magnetic field that induces an electric field . Equation B.27 is still completely valid except that the applied field is found from Equation B.14, namely where = –jω.
The determination of the vector field from a physical coil is found, basically, from Equation B.20 (which corresponds to Equation 12.33). This relationship has also been worked out and published for many other coil configurations.
We also note that since the differential equations B.24, B.27, and B.28 are linear, and the solution given in Equation B.25 is essentially the response to a (spatial) unit impulse at the origin (set I0 = δ(x)), then linear systems theory describes the solution to Equation B.27, (or B.28), as


where denotes convolution. (The added factor of rm is required in order to convert the right side of Equation B.29 into a current density.) The convolution operation can be performed by taking the inverse Fourier transform of the product of the Fourier transform of V' and the Fourier transform of the second derivative of δe. Such operations are readily carried out using the fast Fourier transform (FFT).


SOURCES and FIELDS: Quasistatic (ω < 1000 Hz) i, ,
CONDUCTOR: Limited finite (r < 1 m) region outside the sources inhomogeneous resistive (ωε/ε < 0.15) = 0, ε = ε0

We finally collect the Maxwell's equations in their simplest form. These equations are valid under quasistatic electrophysiological conditions outside the region of bioelectric sources:



Barr RC, Spach MS (1977): Sampling rates required for digital recording of intracellular and extracellular cardiac potentials. Circulation 55: 40-8.

Maxwell J (1865): A dynamical theory of the electromagnetic field. Phil. Trans. R. Soc. (Lond.) 155: 459-512.

Maxwell J (1873): Treatise on Electricity and Magnetism, Vol. 2, Oxford. (Reprint by Dover, New York, 1954.)

Plonsey R, Collin R (1961): Principles and Applications of Electromagnetic Fields, 554 pp. McGraw-Hill, New York.

Plonsey R, Heppner DB (1967): Considerations of quasistationarity in electrophysiological systems. Bull. Math. Biophys. 29:(4) 657-64.

Schwan HP, Kay CF (1957): The conductivity of living tissue. Ann. N.Y. Acad. Sci. 65: 1007.


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