The Magnetic Field of a Single Axon:

A Volume Conductor Model*

JAMES K. WOOSLEY, BRADLEY J. ROTH, AND JOHN P. WIKSWO, JR.

Department of Physics and Astronomy, Vanderbilt University, Nashville, Tennessee 37235

Received 2 January 1985; revised 24 April I985

ABSTRACT

A volume conductor model is used to determine the magnetic field associated with a
propagating action potential in an isolated nerve axon. Both the forward problem of
calculating the magnetic field from the transmembrane potential and the inverse problem of
calculating the transmembrane potential from the magnetic field are examined. The
dependence of the model on all parameters is investigated, and the model is used to study
both the axial current in the axon and the ionic current through the axon membrane.

INTRODUCTION
Investigation of the electromagnetic behavior of a nerve can provide
insight into the relationship between bioelectric and biomagnetic fields. In
this paper we shall study a propagating action potential in a single nerve
axon, concentrating on the relationship between the transmembrane potential
and the magnetic field. Close to the nerve the magnetic field is like that
produced by current in a long straight wire: inversely proportional to the
distance from the axon and proportional to the current inside the axon. From
the core-conductor model we know that the magnitude of this current is
equal to the axial derivative of the transmembrane potential divided by
the axons internal resistance per unit length [l]. Therefore near the nerve the
fields are related in a simple way: the magnetic field is proportional to the
derivative of the transmembrane potential. However, farther from the nerve
this proportionality no longer holds and the relationship between the mag-
netic field and the transmembrane potential becomes more complicated. The
purpose of this paper is to present a calculation of the magnetic field which is
valid at all distances from the axon.

*Address reprint requests to John P. Wikswo, Jr., Department of Physics and Astron-
omy, Vanderbilt University, Box 1807 Station B, Nashville, TN 37235.

MATHEMATICAL BIOSCIENCES 76:1-36 (1985) 1

QElsevier Science Publishing Co., Inc., 1985
52 Vanderbilt Ave., New York, NY 10017 0025-5564/85/$03.30
2 JAMES K. WOOSLEY ET AL.

Our calculation is based on the volume conductor model, first developed

by Clark and Plonsey to calculate the electric potential outside an active fiber
[2], and later extended by Swinney and Wikswo to calculate the resulting
magnetic field [3]. The present analysis has two advantages over Swinney and
Wikswos work. First, all calculations now relate the Fourier transforms of
the fields through filter functions in transform space. This speeds the
computations, enables the inverse problem to be solved immediately, and
leads to a more intuitive understanding of the relationships between the
fields. Second, we can use an arbitrary transmembrane potential in all
computations, since we use a fast Fourier transform (FFT) algorithm to
convert between the space and spatial frequency domains. There is no longer
a need to fit the transmembrane potential to an empirical curve.

THE POTENTIAL
We model the nerve axon as an infinitely long cylindrical membrane of
radius a separating the intracellular and extracellular media (Figure 1).
Inside and outside the membrane the media have homogeneous, isotropic
conductivities ai and a,, respectively. We will use cylindrical coordinates p,
8, and z to specify radial, azimuthal, and axial positions.
The forward problem consists of calculating the magnetic field from the
transmembrane potential. As a first step towards this goal, we need expres-
sions for the potentials in the two media. If we assume quasistationarity,
these potentials satisfy Laplaces equation. Using the transmembrane poten-
tial Q,(z) and the continuity of the normal component of the current
density as boundary conditions at the membrane surface, Clark and Plonsey
[2,4] have solved Laplaces equation, expressing the internal potential P,(p, z)
and the external potential a,(~, z) as

where (Yand fi are defined as

P(lklu) =1+-J--
Y(W)

with y given by

~e~,(lw~)4J(lw)
y(lklu) = u,K,( lklu)Z,( lklu) . (9
THE MAGNETIC FIELD OF A SINGLE AXON

FIG. 1. Geometry of the nerve axon.

1,) Ii, K,, and K, are modified Bessel functions, and &(k), the spatial
Fourier transform of the transmembrane potential, is defined as

$&(k) =j-~~~Jz)e+k& (6)

The Fourier transforms of Qi ( p, z) and Qe ( p, z) can similarly be defined as

&(p, k) and &_(p, k). Inspection of Equations (1) and (2) shows that

(7)

K,(W)
&( k, = a( Ikla) K,,( Ikla) cpm(k)

Thus, algebraic relationships in transform space relate the Fourier transform

of the transmembrane potential and the Fourier transforms of the internal
and external potentials. Each such function relating two Fourier transforms
will be called a filter function. For example, the function

Id Iklp)
PCIkla) &( Ikla)
is thefilter function which gives the internal potential from the transmem-
brane potential.

THE CURRENT DENSITY

We can use the relation

J=-avcP (10)
4 JAMES K. WOOSLEY ET AL.

to calculate the current density J from the potential derived by Clark and
Plonsey. To express this relation in transform space we define /, ( p, k) and
/,( p, k) to be the Fourier transforms of the internal and external current
densities, J, (p, z) and J, ( p, z), respectively. The current density is a vector
with two nonzero components in the radial and axial directions, which we
denote with the superscripts p and z. The resulting filter functions relating
the current densities to the transmembrane potential are obtained by perfor-
ming the appropriate differentiation and are listed in Table 1.
We can also investigate the current density in the membrane. If we assume
that the membrane current density and the electric field are radial, it follows
from the continuity of current that the total membrane current density J, is
equal to both the internal and external radial current densities evaluated at
the membrane. J, represents the sum of the membrane capacitive current
density J, (due to charging of the membrane capacity) and the membrane
ionic current density JR (due to the movement of ions through the membrane),
I.e.,

J,(z)=J,(z)+J,(z). (11)

To calculate J, and Js we must know U, the conduction velocity of the

action potential; K, the membrane dielectric constant; and d, the membrane
thickness. The capacitive current is the product of the time derivative of the
transmembrane potential and the membrane capacitance per unit area,
c, = ~c,,/d, where c,, is the permittivity of free space, 8.854X lo-* Fm-.
We assume a constant conduction velocity, so we can replace a time
derivative a/at by a space derivative u a/az. J, is then the product of c,u
and the axial derivative of Qm. We can find Jg by subtracting J, from J,.
The filter functions for obtaining J, and J, from the transmembrane
potential are listed in Table 1.

THE MAGNETIC FIELD

There are two ways to calculate the magnetic field from the current
density. The law of Biot and Savart, together with the fact that the current
density has zero curl, enables us to calculate the magnetic field by integrating
the current density in the axial direction over the axon membrane. Altema-
tively, Amperes law enables us to calculate the magnetic field by integrating
the current density in the radial direction over a surface perpendicular to the
axon. The law of Biot and Savart and Amperes law provide complementary
views of the magnetic field [5]. Although they offer different intuitive pictures
of the sources and processes creating the magnetic field, they yield identical
results for all physically measurable quantities.
THE MAGNETIC FIELD OF A SINGLE AXON 5

TABLE 1

Volume Conductor Model Filter Function Relationships for the Potential and Current
Internalpotential

External potential

Internal axial current density

Internal radial current dens+

External axial current density

B,(k)=-iyk+,,,(k)

Total internal current

Return current
6 JAMES K. WOOSLEY ET AL.

THELAWOFBIOTANDSAVART

The magnetic field can be calculated from the current density using the
law of Biot and Savart,

J(r')x(r-r') d3r,,
/_!,3 (l-4

The volume u can be divided into three regions: the internal, external, and
membrane volumes, denoted u, , u,, and u,,,, where now the membrane is
treated as a very thin, but finite cylindrical layer of thickness d. Using the
principle of superposition, the magnetic field can be written as

B(r) =&(r)+B,(r)+B,(r), (13)

where B,(r), B,(r), and B,(r) are the portions of the magnetic field arising
from the internal, external, and membrane current densities. The law of Biot
and Savart then becomes

+ J,(r>x(r--T)_ d3r, + J,(r)x(r-r)

/ C b-r113 / urn jr-r/l3
(14

The law of Biot and Savart can be rewritten as the sum of a volume
integral and a surface integral by using the vector identity

The volume integral vanishes because the current density has zero curl in an
ohmic medium with a homogeneous, isotropic conductivity. Therefore the
magnetic field is obtained by integrating the current density only over the
membrane surface s [3,6,7]:

(16)

A similar equation holds for the external current density, with a change in
sign due to the different direction of the outward surface normal fr.
Equation (16) is similar to a solution of Poissons equation, such as that
encountered in electrostatics for calculating the potential from a given charge
THE MAGNETIC FIELD OF A SINGLE AXON

FIG. 2. Coordinate system used for evaluating the integral of the current density
required in the law of Biot and Savart.

density. In our case, J(r)xff can be considered the source of the magnetic
field, and l/p--r] as a Greens function. An important difference between
our problem and the analogous electrostatic problem is that we are dealing
with vector instead of scalar fields. Nevertheless, there is a close analogy
between our problem and that of calculating the potential from a cylin-
drically symmetric surface charge density. Electrostatics problems in cylin-
drical coordinates are well studied [8], and the Greens function for obtaining
the potential due to a ring of charge is similar to the result we shall derive.
Figure 2 defines the coordinate system used to solve the integral in
Equation (16), with the vector r denoting the source point and the vector r
denoting the field point. The internal current density Ji can be written

Ji(r) = Jp(r)$ + J:(r)t. (17)

The cross product of J, with the outward unit normal $, evaluated at the
membrane, p = a, is

(18)
8 JAMES K. WOOSLEY ET AL.

where & is a unit vector in the fI-direction. Because of the cylindrical

symmetry, the magnetic field will be independent of the angle 8. If we choose
our field point such that B = 0, then the distance Ir- rl is (Figure 2)

k-r1 =Jp2+ a2 -2apcose+(z-z~)2. (19)

This allows us to write Equation (16), the surface integral for the magnetic
field, as

=Jrnmc(
B;(P9Z) p,a,z-z')J:(a,z')dz', (20)

where

8*.I
G(p,a,z-ZZ) =pJ adOf
0 P2+u2-2apcose~+(z-z~)2

(21)
is the Greens function that gives the magnetic field at the point (p, z) from a
cylindrically symmetric ring of internal axial current density at position z on
the membrane.
Equation (20) is a convolution integral, and it follows from the convolu-
tion theorem that C#,(p,k), x(u, k), and Y(p,a, k), the Fourier trans-
forms of B,(p, z), Jjf(a, z), and c(p, a, z - z), are related by

Thus we see the general strategy of the calculation. We take the Fourier
transform of a field using a FFT routine. We then multiply this in transform
space by the Fourier transform of the Greens function, which we have been
calling the filter function, and then take the inverse Fourier transform of
the product, again using the FFT. Because of the speed of the FFT, the
computation proceeds very quickly provided an analytic expression for the
filter function can be found. We shall now find such an expression for the
Fourier transform of the Greens function defined by Equation (21).
The Fourier transform of G( p, a, w) is

(23)

where a change in variable has been made such that

w=z-z. (24)
THE MAGNETIC FIELD OF A SINGLE AXON 9

If the unit vector 4 is rewritten in terms of the Cartesian unit vectors

gl= -firsin&+jcosfP, (25)

it can be shown [9] that the x-component of the integrand in Equation (21)
will be odd, integrating to zero, while the y-component is even. Similarly, the
complex exponential in Equation (23) can be expanded in terms of trigono-
metric functions, in which case the sine term will produce an odd integrand,
integrating to zero, while the cosine term will again be even. Using these
observations and Equations (21) and (23), we write the function 9( p, a, k) as

(26)
where the scalar function Y( p, a, k) is the y-component of the vector
function 9(p, a, k). Since 8 = 0, it follows that 4 = jl and the &component
is the only nonzero component of 9.
To evaluate the integral over w in Eq. (26) we use the fact that the inverse
distance and the modified Bessel function K,, are a Fourier transform pair:

M coskw
dw= K,(kx). (27)
/0 Jz-7
Reversing the order of integration in Equation (26) and using Equation (27),
we obtain

9(p,~,k)=~/~cose~,(Ikl p2+a2-2ap~0~e+de~. (28)

0

We can now expand the Bessel function in Equation (28) using the
relation [ 81

Ko(kJp2
+P'~ -2ppcose)=I,(k~<)K,(k~>)
+2 E cos~ez,(kP<)K,(kP>),
m-l

(29)

where p< is the smaller of p and p, and p, is the greater of the two.
Orthogonality of the trigonometric functions insures that only the m = 1 term
of the sum will survive the integration over 8. So we finally arrive at the
10 JAMES K. WOOSLEY ET AL.

desired expression

This result is valid when p > a, i.e. for the magnetic field outside the axon.
The magnetic field inside the axon is found by interchanging a and p when
they appear as arguments of the modified Bessel functions in Eq (30) and in
all equations following from it.*
From Equations (22) and (30) we have the desired filter function relation-
ship

~ii(P,k)=CLoU~~(lklU)K~(IklP)JrZ(u,k), (31)

or, in terms of the Fourier transform of the transmembrane potential

Bi( p, k) = ip,,uqk 4(lW)K(lklp) + (kj.

(32)
P( IW) m

A similar derivation can be carried out for the magnetic field due to the
external current density, the result being

.cSe(p, k) = - ip,,uu,k
4(lk!a)K,(Iklp) + ckj.
(33)
4lW) m

When we calculate the magnetic field due to the ionic current density in
the membrane, the vector identity in Equation (15) does not simplify the
derivation, because in this nonohmic region v x J is not necessarily zero.
Thus the law of Biot and Savart must be applied directly. Although the
derivation of a,,,( p, k) is different from the derivation of .%,(p, k) and
L&?~(p, k), it leads to a similar expression

@,,,(p,k)=-ip&k(l +~)r,(lkla)4(lklp)X(k), (34)

The Greens function for the analogous electrostatics problem of a uniform ring of
charge [lo] differs from the result in Eq. (30) by containing zeroth instead of first order
modified Bessel functions. This difference originates in the cos B factor introduced in Eq.
(26) by &, and can be considered a consequence of using vector instead of scalar fields.
2Bquation (30) is similar to that derived by Scott [ll] in his discussion of the magnetic
field of a nerve axon, particularly in the presence of the K, ( 1k(p) modified Bessel function.
THE MAGNETIC FIELD OF A SINGLE AXON 11

and in terms of the transform of the transmembrane potential

Ikluil~(lkla) .KCOI(
Lq#(p,k)=-ip,udk 1+$ -
( )i 8(lw)4I(lm) +l d ki
(35)

The details of the derivation are not given here because they are quite
lengthy, and because calculation shows that this magnetic field is negligible
in relation to the magnetic field due to the internal and external current
densities [9]. This is a consequence of the leading factor of the membrane
thickness d in Eq. (35), and reflects the relative volumes of the membrane
and the axon interior.3
The capacitive current J, (or displacement current LJD/at) in the mem-
brane also makes a small contribution to the magnetic field. Shadowitz [12]
shows that for an unbounded, homogeneous, isotropic, linear medium the
displacement current need not be included in the law of Biot and Savart. This
follows from the vector identity in Equation (15) and the fact that every-
where the displacement D = rE has zero curl (the surface integral is taken at
infinity and therefore vanishes). In our case, we do not have a homogeneous
medium, because of the discontinuity in the dielectric constant at the
boundaries of the axon membrane, across which the tangential component of
E, but not necessarily that of D, is continuous. Thus, if we replace J by
~?D/at in Equation (15), we find that the second term, the surface integral, is
nonzero. However, calculation shows that this contribution to the membrane
magnetic field is negligible compared to that of J8, and thus it is extremely
small compared to the total magnetic field. We neglect this contribution
henceforth, as did Swinney and Wikswo [3].
Ignoring the contribution to the magnetic field from the current density in
the membrane, and ignoring all contributions from displacement currents, we
can write our final result, the filter function for obtaining the magnetic field
from the transmembrane potential, as the sum of Eqs. (32) and (33):

(36)

3Equation (34) differs from the result of Swinney and Wikswo [3] by a factor of
(1 + d/2o)/a, due to an algebraic error in their derivation. This makes the magnetic field
due to the membrane current density several orders of magnitude larger than previously
thought. However, it is still negligible compared to the total magnetic field.
12 JAMES K. WOOSLEY ET AL.

AMPERES L.A W

An alternative derivation of this result can be based on Amperes law

B. ~0 = PO lenclosed 7 (37)

wherezenclosedis the net current threading the closed path c, and p. is the
permeability of free space, 4a x 10 _ TmA. Cylindrical symmetry allows
us to simplify the line integral to

where the path of integration is a circle of radius p centered at z. In the case

of an active nerve, Zen&& is the sum of the total current in the axon interior
Z,(z) and the return current I,,,( p, z) in the external medium between the
axon membrane and the field point radius p. Since we know the axial current
densities in the internal and external media (Table l), we can obtain Zenclosed
by calculating the current density flux through a surface bounded by our line
integral path. To do this we shall need the two Bessel function relationships

$xz,(x)dx=xz,(x) (39)

and

j-xK,(x)dx=-xK,(x). (40)

If we denote the Fourier transforms of Z,(z) and I,,,( p, z) as &(k) and

&( p, k), we can perform the integrations to find that

2aqik
3(k) = B(,k,~)Zo(,k,~),k,zl(k~)~m(k) (41)
and

2qik
,a;,,(P,k) = a(,k,o)K,(,k,a),k,{ aK,(Ikla)-pK,(lklp))~~(k).

(42)

From Equations (38), (41), and (42) we see that the magnetic field due to the
total current inside the axon [Bit,,, ( p, z)] and the magnetic field due to the
return current [ B,, (p, z)] have Fourier transforms

pouiik
aiitot(P,k)= ~~(,k,a)Zo(,k,~),k,uzl(ka)~m(k)~ (43)
THE MAGNETIC FIELD OF A SINGLE AXON 13

and

Cc&k
are,(Prk) = pa((,+)KO(],+r)]k] { a~,(lkla)-p~,(lkl~))~~m(k).

Adding these two contributions gives a filter function relating the magnetic
field and the transmembrane potential

[a~,(lkla)-p~,(lklp)l
+ 4 l&dk,( IW)
(45)

If we substitute for a(lkla) and jl(lkla) their definitions in terms of

modified Bessel functions, rearrange terms, and use a relation which can be
derived from the Wronskian [8],

we can show that the filter function defined by Equation (45), derived from
Amperes law, is identical to the filter function defined by Equation (36)
derived from the law of Biot and Savart.
The filter function relations for the magnetic field are summarized in
Table 2.
We see that we can derive the same filter function relating the transmem-
brane potential and the magnetic field from two different starting points,
Mathematically, the difference in the two approaches is that with the law of
Biot and Savart, the radial integration of the current density is implicit in
Equation (16) and only the axial integration must be performed explicitly,
while with Amperes law the axial integration of the current density is
implicit in Equation (37) and only the radial integration must be performed
explicitly. Continuity of current, the ohmic nature of the current, and
cylindrical symmetry place such restrictive constraints on the current density
that only one integration is sufficient to specify the magnetic field com-
pletely. In transform space the two views amount to a regrouping of modified
Bessel functions in the filter function for the total magnetic field.
Separating the magnetic fields into terms produced by specific currents,
such as B, and B, or Bitot and B,,,, is often helpful in obtaining an intuitive
understanding of the behavior of the magnetic field. However, these assign-
14 JAMES K. WOOSLEY ET AL

TABLE 2

Volume Conductor Model Filter Function Relationships for the Magnetic Field

Magnetic field due to the internal current densrty

Magnetic field due IOthe external current density

Magnetic field due to the current density in the membrane

Magnetic field due to the total internal current

Magnelic field due to the return current

Total magnetic field

Ether

where

a(lkla)=-[l+v(lkla)l.
/3(jkla) =l+ 1
Y(W)
THE MAGNETIC FIELD OF A SINGLE AXON

FIG. 3. A schematic representation of the relationships between the transmembrane

potential and the internal and external potentials and current densities and their resulting
magnetic fields. A double arrow means the two fields form a Fourier transform pair.

ments are always artificial in the sense that only the total magnetic field is a
physically measurable quantity. Amperes law and the law of Biot and Savart
of course yield the same result for the total magnetic field. They are
equivalent, though they provide different views as to how this magnetic field
is produced.

COMMENTS ON THE MATHEMATICS

The results of all these calculations are summarized in Figures 3 and 4.
Figure 3 traces the relationships between the transmembrane potential, the
internal and external potentials, their current densities, and their resulting
magnetic fields. The analogous relationships between the membrane current
density and its magnetic field are shown in Figure 4. Although we have been
concentrating on the relationship between the transmembrane potential and
the magnetic field, Figures 3 and 4 show that given any one field, all the
other fields can be calculated from it.
16 JAMES K. WOOSLEY ET AL.

FIG. 4. A schematic representation of the relationships between the transmembrane

potential and the membrane current density and its resulting magnetic field. A double
arrow means the two fields form a Fourier transform pair.

The theme of this mathematical development has been the use of Fourier
transforms, filter functions, and the convolution theorem to simplify the
relationships between the fields. We can provide insight into our underlying
motivation if we recast the mathematics in the language of linear algebra. We
associate all fields with vectors in the z-representation and associate all
Greens functions with matrices. Two vectors, say a,,, and B, are then related
by the matrix equation
B=G@,,,. (47)
Because our Greens function depends not on z and z separately but
only on Iz - ~1, G is symmetric, so we can find a new representation in
which it is diagonal. This new representation is what we earlier termed
transform space, and which we can also call the k-representation. In
terms of linear algebra, the operation of taking the Fourier transform is
equivalent to a unitary transformation, represented by the matrix U, connect-
ing the z-representation and the k-representation:

UB = UGU -UQ,,, , (48)

or
~=%$~ (49)
where
z?#=UB, (W
h =U@,, (51)
THE MAGNETIC FIELD OF A SINGLE AXON 17

g=VGV-. (52)

Equations (47) through (52) amount to a restatement of the convolution

theorem. The advantage of the k-representation is that 9? is diagonal

DEPENDENCE OF THE MODEL ON PARAMETERS

As a first step in interpreting the mathematical relationships we have
derived, we investigate the dependence of the filter functions on various
parameters. We treat these parameters, as well as the transmembrane poten-
tial, as being independent quantities. Any relationships between the parame-
ters act as additional constraints on the possible behavior of the nerve.
The five parameters required by the volume conductor model are:

(1) The conductivity of the external medium, a,.

(2) The conductivity of the internal medium, a,.
(3) The conduction velocity, u.
(4) The nerve radius, a.
(5) The field point radius, p.

Two additional parameters are needed to calculate the membrane capacitive

current density, the membrane ionic current density, and the magnetic field
which these produce:

(6) The thickness of the membrane, d.

(7) The dielectric constant of the membrane, K.

In general, the dependence of the filter functions on these parameters is

quite complex. However, typical nerve fibers have action potentials of limited
frequency bandwidth, radii, and conductivities such that the two relations
]/~]a << 1 and y >> 1 hold fairly well. The largest value of k which contributes
significantly to the frequency spectrum of &(k) is roughly ~/AZ, where AZ
is the spatial length of the depolarization phase of the action potential.
Therefore the first condition, lkla =x 1, implies that the transmembrane
potential changes over large distances compared to the axon radius. Scott [ll]
interprets y as the ratio of the internal to external longitudinal impedance of
the fiber. Therefore the second condition, y >> 1, implies that the external
resistance of the bath is much less than the internal resistance of the axon.
When both these conditions hold we say that we are in the DC limit. In this
limit we can deduce the qualitative behavior of the fields by setting a = - y,
/3 = 1 and looking at the small argument limit (Table 3) of all modified Bessel
functions which contain the axon radius in their argument.
Of all the parameters, the model is least sensitive to changes in the
external conductivity a,. In the DC limit only the external potential depends
18 JAMES K. WOOSLEY ET AL.

TABLE 3

Modified Bessel Functions at

Small and Large Arguments

If X -=X1.

I,(x) -1,

I&>=&
K,(x)=-[ln(S)+0.5772...],

K,(x)=;

If X&l.

on a,, being inversely proportional to it. By setting a, equal to infinity we

lose all knowledge of the external potential, but leave quantities
all other
(including the external current density) unchanged. This justifies the assump-
tion in the core-conductor model of nerve magnetic fields [l] that the
resistance of the external medium can be set equal to zero. Another way of
interpreting this behavior is to say that as long as the voltage drop outside
the membrane is small compared to the transmembrane potential, or as long
as the external resistance is much less than the internal resistance (y > l),
the axon acts like a current source, injecting current into the external bath.
The magnetic field, which is produced by currents, is unchanged by varying
the resistive load on a current source. If the external conductivity is very
small, i.e. a, -=Ko,, then the approximation y x=-1 will begin to break down,
and the dependence on ue will become more complex.
The internal conductivity a, is a more important parameter in the model.
In the DC limit, the external potential, the internal and external current
densities, and the magnetic field are all proportional to it.
The conduction velocity u enters the model through the assumption that
the action potential propagates down the axon with a constant speed. This
enables us to convert the independent variable in Q,,, from time, appropriate
for measured data, to distance, appropriate for a boundary condition to
Laplaces equation. Although only the filter functions for the membrane
current densities J, and Jg, and their contribution to the magnetic field,
THE MAGNETIC FIELD OF A SINGLE AXON 19

depend explicitly on the conduction velocity, there is an implicit dependence

on the conduction velocity in all the calculations. The conduction velocity
sets the axial length scale, compressing or stretching the action potential
along the z-axis, and thereby stretching or compressing the transform of the
measured field in transform space.
Physically there are two competing effects due to changes in the conduc-
tion velocity. As u increases, spatial gradients of the potential become
smaller, decreasing the current density and the magnetic field. However, at a
given field point p, the contribution to the magnetic field due to the return
current relative to the internal current will be less as the action potential is
spread out further along the z-axis, tending to increase the magnetic field. If
p is chosen on the same order of magnitude as AZ, the spatial length of the
depolarization phase of the action potential, then small increases in u will
tend to narrow the signal and lower its amplitude.4
In the DC limit the internal potential and current density are independent
of the axon radius a. However, the external potential depends strongly on a.
This can be understood if we consider Qc (p, z) as being produced by a pair
of electric dipoles propagating with the nerve impulse [13]. The dipole
strength is proportional to the cross-sectional area of the axon, which is in
turn proportional to the axon radius squared. So Qe( p, z) and also J, ( p, z)
are proportional to a2 in the DC limit.
The magnetic field, in the DC limit, is also proportional to the axon radius
squared. This is most easily understood from the point of view of Amperes
law. The current density inside the axon is independent of a, so the total
current in the axon, Ii, and the magnetic field due to this current, Pitot, are
proportional to the axon cross section. The return current is always some
fraction of the internal current, and this fraction does not depend strongly on
the axon radius (unless p = u, in which case the return current makes a
negligible contribution to the magnetic field). So we expect B,,, to change in
the same way as Bit,,, when the axon radius is varied. Therefore, the total
magnetic field should be proportional to u2.
In the Biot-Savart view we find that B, is proportional to u2, but the
explanation is different than that given from the point of view of Amperes
law. One factor of u arises because the axon surface, over which we integrate

4The sign of the conduction velocity can lead to some confusion. In all plots shown in
this paper both the time and axial distance axes increase towards the right and the action
potential is propagating to the left. In all our equations IA should be interpreted as a
conversion factor between t and z, a strictly positive quantity equal to the magnitude of the
action potential conduction velocity. This is a different point of view than taken in Swinney
and Wikswo [3], so one must use care in comparing their equations with ours when u
appears explicitly in an equation.
20 JAMES K. WOOSLEY ET AL.

in Equation (21), has an area proportional to the axon radius. The explana-
tion of the second factor of a is more. subtle. In the Greens function in
Equation (21) we integrate 6 azimuthally. At two points on opposite sides of
the axon, 8 points in opposite directions. These two contributions to the
integration would exactly cancel each other except for the fact that the
contribution from the side of the axon nearest the field point is slightly
greater than the contribution from the side opposite the field point. The
difference between the two contributions increases as the axon radius gets
larger, which accounts for the second factor of a. The magnetic field due to
the external current density, B,, is proportional to u4. The extra factor of u2
is present because the external current density itself is proportional to the
axon cross section. As the nerve radius gets larger, B,(p, z) increases faster
than B,( p, z), so the relative contribution of B,( p, z) becomes more im-
portant for large axons. However, B,(p, z) does not make a significant
contribution to the total magnetic field even for fibers with radii on the order
of 100 pm. Typically, Bi/B, = 2000 for an axon with radius a = 0.02 mm,
whereas Bi/B, = 200 for radius a = 0.1 mm. So from the point of view of the
law of Biot and Savart it is a very good approximation to say that the total
magnetic field is due only to the internal current density for any reasonable
sized nerve and that the magnitude of B is proportional to u2.
When calculating the capacitive or ionic current density in the membrane,
and its resulting magnetic field, the membrane thickness d and dielectric
constant K are needed. These two parameters often, but not always, appear
together as the membrane capacitance per unit area, c, = KE~/~. The
capacitive and ionic current densities depend crucially on the value of c,.
Fortunately a relationship between c,, a, a, u,, and the action potential
shape can be found, so that c, can be determined from the other parameters.
We have found that the magnitude of the magnetic field calculated from
the transmembrane potential depends strongly on the internal conductivity
and the axon radius. However, we can relate any of the fields of interest
through filter functions, and they do not all share this strong dependence on
a and u,. An example is the filter function relating the total internal current
to the magnetic field. This filter function will be derived later [Equation (54)],
and we shall find that it is nearly independent of the axon radius and the
internal conductivity.

This provides another explanation of why the factor of cos 6 appears in Equation (26)
and why in Equation (29) it is the m =l, not the M = 0, term which contributes to the
magnetic field, making our calculation of the magnetic field from a cylindrically symmetric
current distribution different from the calculation of the electrical potential from a
cylindrically symmetric charge distribution.
THE MAGNETIC FIELD OF A SINGLE AXON 21

We have yet to investigate the dependence of the fields and filter functions
on the field point radius p. Inside the axon (p < u) the internal potential and
current density are nearly independent of p, i.e., Qi and Ji are nearly
constant over the axon cross section. But external to the axon (p > a) the
dependence on p can be quite complex. We must look at either the filter
functions in transform space or the resulting calculated fields to understand
the p-dependence.

RESULTS OF THE MODEL

Further investigation of the volume conductor model requires knowledge
of the transmembrane potential a,. All our calculations are based on a
transmembrane potential from a crayfish medial giant axon. The signal was
recorded as a function of time, but can be considered as a function of space
by using the measured conduction velocity of u =16.5 msec-. The details
of how these data were obtained, along with a comparison between theory
and experiment, are given elsewhere [14]. Now our intention is to use the
transmembrane potential to investigate properties of the fields, particularly
their dependence on the field point radius.
The transmembrane potential and the magnitude of its Fourier transform
are shown in Figure 5(a) and (b). At a spatial frequency of 5 mm- (13 kI-Iz)
the magnitude of the transform has dropped by 30 dB relative to its peak.
With an axon radius of about 0.1 mm, this implies that for all frequencies of
interest the signal obeys the relation ]k]a < 0.5, and that for most of the
signal (k 1a is much smaller.
When lklp -K 1 the DC limit can be extended to include Bessel functions
containing the field point radius in their argument. This will be the case when
p is small compared to AZ, the spatial width of the depolarization phase of
the action potential. However, potentials and magnetic fields are often
measured at field point radii which are on the same order as, or larger than,
AZ. In this case the DC limit will not accurately describe the behavior of the
fields, and the exact filter functions must be studied. In this example, the
spatial width of the depolarization phase of the action potential is approxi-
mately 3 mm.
The volume conductor model has been used to investigate the behavior of
the external potential [2,4,15,16]. A description of this behavior is included
here for completeness, and to set up the formalism from which the less-studied
current density and magnetic field will be discussed. Figure 6(a) shows the
filter function for obtaining the external potential from the transmembrane
potential. This filter function is multiplied by the Fourier transform of the
transmembrane potential e,,,(k) [Figure 5(b)] to give the Fourier transform
of the external potential &.(p, k). At k = 0 the filter is zero, implying that
22 JAMES K. WOOSLEY ET AL.

(a)
Measured a,,,(Z)

100 -

Qrn (z)
mV 5o

OO I
Time, msec

(b)
I+ mk( 4

k, km
FIG. 5. (a) The measured transmembrane potential. (b) The magnitude of the Fourier
transform of the transmembrane potential. a = 16.5 msec -.

information about the membrane resting potential is not contained in

Qe( p, z). (Note that this filter function is negative, so zero is at the top of the
vertical axis.) For small k ( < 0.5 mm-), the filter has approximately the
shape of a parabola, which in the time domain implies a second derivative
relationship between Qe ( p, z) and a, ( z).~ For large enough k the filter will
turn over and approach zero exponentially, removing high frequency compo-
nents of the experimental noise as well as the signal from the calculated field.
As p grows, this low pass filtering plays an increasingly larger role in
determining the shape of the external potential.

6The filter function &.(p,k) also contains the factor &(lklp), which is changing
logarithmically in k (see Table 3) so the relationship between ae( p. z) and Q,,,(z) is only
qualitatively like a second derivative. In general, the presence of K, in a filter function
complicates the interpretation in the DC limit.
THE MAGNETIC FIELD OF A SINGLE AXON 23

(a)
Filter Function,q&(p,k) from #,,,(k)

dimensionless

1 c
I 5
ktpmm
M
Calculated @,(p,z)

Time, msec

FIG. 6. (a) The filter function for obtaining the external potential from the transmem-
brane potential. The parameters used in this calculation are a = 0.107 mm, oi = 1.7
Wm-, o, = 2.06 Wm-, p=O.8,1.0,1.2 mm. (b) The calculated external potential.
The same parameters were used as in (a), along with u = 16.5 msec-.

The filter function for obtaining the internal potential from the transmem-
brane potential is essentially equal to unity for all k of interest. This implies
that Qi( p, z) and a,(z) are nearly identical and enables the potential
measured using a microelectrode, which is actually Oi (p, z), to be treated as
the transmembrane potential am(z). The theory can be developed using
(Pi(u, z) as the measured field, as was done by P. Rosenfalck [15], and by
Andreassen and A. Rosenfalck [17]. This requires merely solving Equation
(7) for $D~,( k) and substituting the result into all the equations in Tables 1
and 2. In certain situations this may be desirable, for instance when a, is
small enough that there is a significant potential drop outside the nerve. In
our analysis of a single nerve axon there is approximately one percent error
in taking Oi( p, z) = Q,(z). Like the external potential filter function, the
filter function for +i( p, k) will fall exponentially to zero at large k. This is a
24 JAMES K. WOOSLEY ET AL.

common property of forward filter functions, due to the behavior of modified

Bessel functions at large arguments (see Table 3).
The filter functions for the current densities can be found in Table 1. By
applying the DC limit we see that the filter function for /F(p, k) is
proportional to k, the filter functions for #F( p, k) and #i( p, k) are
proportional to k2, and the filter function for fi( p, k) is proportional to k3.
This implies that the internal axial current density J,( p, z) looks like the first
derivative of the transmembrane potential, the internal and external radial
current densities will look like the second derivative, and the external axial
current density looks like the third derivative of a,,,(z).
Taking the point of view of the law of Biot and Savart, we can investigate
the individual contributions to the magnetic field from the internal and
external current densities. The filter function for obtaining the magnetic field
due to the internal axial current density from the transmembrane potential is
presented in Figure 7. By previous arguments this is essentially the filter
function for the total magnetic field. At small k it rises linearly, implying the
first derivative relationship to Q,,,(z) predicted by the core-conductor model.
For field point radii on the order of a millimeter, the extremum of the filter
occurs where there are still significant contributions from the Fourier trans-
form of the transmembrane potential [Figure 5(b)]. So the magnetic field will
only roughly resemble the derivative of Q,(z), that is, will be only roughly
proportional to J;( p, z). Much of the high frequency information will be
severely attenuated. Therefore the magnetic field will be broadened and the
two peaks of biphasic signal will become more symmetrical. Also, the
magnetic field gets smaller with increasing p, as is seen from the decreasing
height of the maximum of the filter function. The exact dependence of the
size of Bi( p, z) on p is difficult to ascertain from the filter. In fact, since the
shape of Bi changes, it is not clear how to quantitatively compare the size of
B, at different field point radii. Swinney and Wikswo [3] discuss the p-depen-
dence further. They find that the peak to peak size of the magnetic field falls
as p-l near the nerve (p -K AZ), and the rate of falloff increases smoothly
with distance until the magnetic field falls as pP 3 far from the nerve
(p % AZ).
Figure 8 contains the filter function to obtain the magnetic field due to the
external current density from the transmembrane potential, and B,( p, z)
itself. Note that B,(p,z) is two orders of magnitude smaller than B,(p, z),
verifying our earlier statement that B, (p, z) accurately represents the total
magnetic field.
We can also investigate the magnetic field from the point of view of
Amperes law. Figures 9 and 10 show the contributions to the magnetic field
from the total internal current Bitot( p, z) and return current B,,,( p, z), and
their respective filter functions. Both the internal and return currents contrib-
ute significantly to the magnetic field. The magnetic field due to the total
internal current falls as p-l with a constant shape, approximately propor-
THE MAGNETIC FIELD OF A SINGLE AXON 25

(a)
Filter Function, B, (p, k) from y&(k)

Calculated El (p,z)

Bi(psz) 0

PT
-3oo-

0 I 2
Time, msec

FIG. 7. (a) The filter function for obtaining the magnetic field due to the internal
current density from the transmembrane potential, using the law of Biot and Savart. The
parametersusedin thiscalculationare CI= O.l07mm, u, =1.73-l m-l, CJ,= 2.06 52-l m-l,
p = 0.8,1.0,1.2 mm. The dotted line is the analogous core-conductor result. (b) The
calculated magnetic field due to the internal current density. The same parameters are used
as in (a), along with u = 16.5 msec-. To a good approximation, this is the total magnetic
field.

tional to the first derivative of Q,,,(z). It essentially represents the magnetic

field predicted by the core-conductor model. The magnetic field due to the
return current represents the volume conductor models correction to the
core-conductor model prediction. For p -C AZ the core-conductor model is
valid and the return current contributes little to the total magnetic field. For
p = AZ the contribution of B,, becomes appreciable. For p > AZ the
internal and return currents are nearly equal and opposite, and contribute
almost equally to the total magnetic field. Near the axon B,, is nearly
proportional to the third derivative of a,,, (z). The quadrupolar nature of B,,,
tends to widen the total magnetic field. Since B,, is zero at both p = a and
26 JAMES K. WOOSLEY ET AL.

(a) Filter Function,B,(p, k) from &,(k)

0.6 -

pT
mV 0.3-

Calculated i3, (p,z)

me
pT -3-

0 I 2
Time, msec

FIG. 8. (a) The filter function for obtaining the magnetic field due to the external
current density from the transmembrane potential, using the law of Biot and Savart. The
parametersusedin this calculation are n = 0.107 mm, a, = 1.7 9-l m-, o, = 2.06 !X mm,
p = 0.8,1.0,1.2 mm. (b) The calculated magnetic field due to the external current density.
The same parameters are used as in (a), along with u = 16.5 msec -l.

p = co, it must have a maximum at some intermediate p. The plots in Figure

10 must be near this maximum, as they are virtually independent of the field
point radius.
The law of Biot and Savart and Amperes law provide different physical
explanations of the p-dependence of the magnetic field. From the point of
view of the law of Biot and Savart, the falloff of the magnetic field with p is

Although the magnetic fields in Figures 9 and 10 are somewhat noisy, this does not
imply that the total magnetic field calculated from Amperes law is any noisier than the
magnetic field from the law of Biot and Savart. The filter functions for the two contribu-
tions to the Amperian calculation are approximately the negative of each other at large k,
so their sum goes to zero much faster than either one individually. Thus, the noise is highly
correlated, and cancels when the magnetic fields are added.
THE MAGNETIC FIELD OF A SINGLE AXON 27

(a)
Filter Function, Bitot (p,k) from d,(k)

Calculated Bitot (p,z)

300-

Bitot (PvZ) O
PT -300 -

-600 -

Time, msec

FIG. 9. (a) The filter function for obtaining the magnetic field due to the total internal
current from the transmembrane potential, using Amperes law. The parameters used in this
calculationare a=O.l07mm, a,=1.7 Wm-, q-2.06 Wm-I, p=0.8,1.0,1.2mm.
The dotted line is the analogous core-conductor result. (b) The calculated magnetic field
due to the total internal current. The same parameters are used as in (a), along with
u=16.5 mse-.

faster than l/p because of the finite spatial extent of the action potential.
From the point of view of Amperes law, the falloff of the magnetic field
faster than l/p is attributed to cancellation of the current inside the axon by
an increasing fraction of the return current.
Intuitively, the Biot-Savart view is most helpful for understanding the
magnetic field far from the axon (p X- AZ), where the current distribution
can be modeled quite accurately as two oppositely directed current dipoles
[7]. The Amperes law view is most useful for describing the magnetic field
very close to the axon (p = AZ), where the magnetic field is approximately
proportional to the derivative of the transmembrane potential [l]. The full
power of the volume conductor model is necessary to describe the magnetic
28 JAMES K. WOOSLEY ET AL.

(a)
Filter Function, B,,, (p,k) from y&,(k)

PT
mV

b)
Calculated Bre+ (p,z)

200 -
bet (p,d
PT
0

0 I 2
Time,msec

FIG. 10. (a) The filter function for obtaining the magnetic field due to the return
current from the transmembrane potential, using Amperes law. The parameters used in this
calculationare a=0.107 mm, 0,=1.7 Q-m-l, 0,=2.06 O-m-l, p=O.8,1.0,1.2 mm.
The filter function is zero in the core-conductor model. (b) The calculated magnetic field
due to the return current. The same parameters are used as in (a), along with u = 16.5
msec-.

field at intermediate distances ( p = AZ) where neither the near nor far field
approximations are valid.
Having now achieved our goal of calculating the magnetic field from the
transmembrane potential, we can invert the process and calculate Q, from
B. We shall find that the formalism of filter functions makes this inversion
very easy in theory, but that in practice difficulties arise from the behavior of
the filter functions.

THE INVERSE PROBLEM AND WINDOWING

The inverse problem consists of calculating the transmembrane potential
from the magnetic field. Once the filter function for the forward problem is
derived relating two fields, the filter function for the inverse problem is found
THE MAGNETIC FIELD OF A SINGLE AXON 29

by merely taking its reciprocal. In the language of linear algebra, this is

equivalent to saying that the inverse of a diagonal matrix is found by merely
taking the reciprocal of each matrix element on the diagonal The ease in
obtaining the inverse solution represents another advantage of expressing all
fields as Fourier transforms in the k-representation. As was previously
pointed out, filter functions in the forward problem have the property of
going to zero exponentially at large k. Therefore the filter functions in the
inverse problem will rise exponentially to infinity. Thus any high frequency
noise in experimental data will be greatly exaggerated in the calculated field.
To obtain a usable solution to the inverse problem we must eliminate this
exponential increase of the filter function. When calculating Q,,,(z) from the
external potential, Clark, Greco, and Harman [18] use an optimal Wiener
filter for this purpose. However, their Wiener filter does not fall off with
increasing k as fast as our filter functions grow (exponentially), unless the
noise power spectrum is known exactly. Since we are using experimental data
for which we do not know the precise noise spectrum, we cannot use Wiener
filtering.
We have chosen the alternative approach of windowing the filter functions
in transform space. We use a Tukey window8 .F(jkD [19], where we choose
the k-values for the beginning and the ending of the window falloff:

:, ( I;-h)] lklxkl t53j

2 l+cos vk , k,<jkl<kz,

b WI> k,.

This window was selected over others because it has the virtue of faithfully
passing the low frequency components, which contain most of the signal.
Taking the reciprocal of the filter function defined by Equation (36), we
get the filter function for obtaining the transmembrane potential from the
measured magnetic field. Figure 11 shows the measured magnetic field and
the magnitude of its Fourier transform. A description of how this magnetic
field was measured is given elsewhere [14]. Figure 12(a) shows the filter
function and the filter function as corrected using the Tukey window (k, = 0
mm-, k, = 4 mm-). Also shown is the core-conductor model result, which
in the time domain corresponds to a simple integration. The filter function
rises to infinity at small k. Consequently any low frequency drift in the
magnetic data will be emphasized in the calculated transmembrane potential.
The filter function also goes to infinity at large k, so small details in the
magnetic signal, originating from the action potential or from unwanted

With k, = 0, the Tukey window reduces to the Hanning window.

30 JAMES K. WOOSLEY ET AL.

(a)
Measured B(p,z)

I 2
Time, msec

(b)
lB(p,k)l

FIG. 11. (a) The measured magnetic field. (b) The magnitude of the Fourier transform
of the magnetic field. u = 16.5 m set - .

noise, will also be emphasized in Q,,,(z). The Tukey window eliminates the
high frequency noise, but will not remove the effects of low frequency noise
in the data. The result of the inverse calculation is shown in Figure 12(b). A
complete comparison between theory and data is given elsewhere [14], but we
note here that while the agreement between the shapes of the action potential
waveforms in Figs. 5(a) and 12(b) is excellent, the amplitudes of the two
signals do not match because of uncertainty in our values of a and a,.
The choice of k, and k, depends on the signal-to-noise ratio of the
measured data and on the field point radius. If the data are noisy or if the
measurement is made far from the nerve, then the calculated transmembrane
potential is quite sensitive to the choice of k,, and it may be difficult to
eliminate the experimental noise without distorting the biological signal. In
our example the signal-to-noise ratio is large enough that the calculation is
fairly insensitive to the exact value of k,.
THE MAGNETIC FIELD OF A SINGLE AXON 31

(a)
Filter Function, #+,,(k) from B(p,k)

mV
pT

Calculated @,,,(z)

Time, msec

FIG. 12. (a) The filter function for obtaining the transmembrane potential from the
magnetic field. The dashed line is the filter function multiplied by a Tukey window,
k, = 0.0 mm-, k, = 4.0 mm-t. The parameters used in this calculation are n = 0.107 mm,
0, =1.7 0-l m-l, o,= 2.06 9-l m-l, p =1.4 mm. The dotted line is the analogous
core-conductor result. (b) The calculated transmembrane potential. The same parameters
are used as in (a), along with u =16.5 mse-.

Our investigation of the relationship between the transmembrane poten-

tial and the magnetic field is now complete. All of the information relating
the two fields is given in their filter function, as expressed in Equation (36) or
Equation (45). However, en route to these two filter functions we derived
relationships between many other bioelectric fields. Figures 3 and 4 demon-
strate that starting from the magnetic field we can calculate any field of
interest. Our last task will be to show in two particular cases how such
information can be extracted from the magnetic field.

APPLICATIONS OF THE MODEL

Using the filter functions derived with this model, we shall give two
examples of how the magnetic field can be used to study bioelectric phenom-
32 JAMES K. WOOSLEY ET AL.

ena. In the first example the magnetic field will be related to the axial current
in the axon. In the second example the magnetic field will be related to the
membrane ionic current density.

TOTAL INTERNAL CURRENT

By taking the filter function for the magnetic field and the filter function
for the total internal current, we can eliminate the transmembrane potential
&(k) and generate a relationship between 4(/c) and S?( p, k):

4(k) =
I-LolWo( 2s K( IO)
I+)

In the core-conductor model we assumed that the magnetic field was propor-
tional to the internal current. Equation (54) is the analogous volume conduc-
tor result. We see in Figure 13(a) that for small k, Yi(k) and .%(p, k) are
proportional, so we recover the core-conductor model prediction. In this
sense the core-conductor model represents the DC limit of the volume
conductor model. The accuracy of the core-conductor model is limited by the
requirement that lklp be small, so it is most accurate when the magnetic field
is measured as close to the nerve as possible. The dashed line in Figure 13(b)
shows the core-conductor model prediction for the internal current. This
current is proportional to the magnetic field shown in Figure 11(a).
Note that the filter function in Equation (54) rises exponentially to
infinity at large k. We therefore must use our windowing techniques devel-
oped for the transmembrane potential. The current shown in Figure 13(b)
was calculated using a Tukey filter with k, = 0 mm- and k, = 4 mm- .
Also, recall that this filter function was mentioned earlier when discussing the
dependence of the model on parameters. This filter function is almost
independent of the axon radius a, and of the internal and external conductiv-
ities a, and a,, demonstrating that the magnetic field provides a direct
measurement of the intracellular current.

MEMBRANE IONIC CURRENT DENSITY

We have already shown how the membrane ionic current density, Jg, can
be calculated from the transmembrane potential The filter function relation-
ship for such a calculation is given in Table 1. This filter function is
interesting in that both the real and imaginary parts are nonzero. All filter
functions discussed previously have been either real or pure imaginary (the
filter function for the membrane magnetic field is complex, but only because
it contains J,). Thus the phase of the filter function is a nonlinear function of
k, which implies that in the spatial domain the shape of Jg bears no simple
relationship to Q,.
THE MAGNETIC FIELD OF A SINGLE AXON 33

(a)
Filter Function, Ii (k) from B(p, k)

M
Calculated II(z)

Time, msec

FIG. 13. (a) The filter function for obtaining the total internal current from the
magnetic field. The dashed line is the filter function after multiplying by a Tukey window,
kt = 0.0 mm-, k, = 4.0 mm-t. The parameters used in this calculation are 0 = 0.107 mm,
q =1.7 Q2- m-l, 0, = 2.06 W m-l, p =1.4 mm. The dotted line is the analogous
core-conductor result. (b) The total internal current calculated from the magnetic field. The
same parameters are used as in (a), along with u = 16.5 msec-. The dashed line is the
core-conductor model prediction.

The membrane ionic current density can also be found from the magnetic
field by taking the filter functions for f,(k) and 9( p, k) from Table 1 and
eliminating +,,,(k). We find

B(lW) ucz WI 1
4(k)-pouKl;,k,p) mT+j- k bdlkla)

X (55)
g(pk)
34 JAMES K. WOOSLEY ET AL.

(a>
Filter Function, JB(k) from@(p,k)

0.4-

tLA
mm*. PT 0.2 -

0
I 5

ktknrn

J,, Je *Jr,,

Time, msec

FIG. 14. (a) The magnitude of the filter function for obtaining the membrane ionic
current density from the magnetic field. The dashed line is the filter function after
multiplying by a Tukey window with k, = 0.0 mm- and k, = 3.0 mm-. The parameters
used in this calculation are D = 0.107 mm, o, =1.7 a- m-l, eC = 2.06 Q- m-l, p =1.4
mm, u=16.5 msec-, c, = 7.1 x 10 _ 3 Frn- . (b) The membrane ionic current density
calculated from the magnetic field, using the same parameters as in (a). Also shown are the
membrane capacitive current density J, and the total membrane current density J,,,.

The magnitude of this filter function is shown in Figure 14(a). Again the
filter function rises to infinity exponentially as k gets large, and we must
apply our windowing technique to achieve a stable solution. The dashed line
in Figure 14(a) represents the filter function after being multiplied by a
Tukey window with k, = 0 mm- and k, = 3 mm-. It was this filter
function which was used in calculating Jg shown in Figure 14(b).
The initial rise of the transmembrane potential is due only to the dis-
charge of the membrane capacitance, so that Jg remains zero until the
sudden appearance of an inward ionic current when the transmembrane
THE MAGNETIC FIELD OF A SINGLE AXON 35

potential reaches threshold. We made several calculations of J8 using differ-

ent values of the membrane capacity per unit area, c,,,. For c,,, = 7.1 X 10 - 3
Fm- 2 (0.71 cF/c&) the ionic current was zero throughout the rising phase
of the action potential. Thus c,,, can be found from the shape of the magnetic
field and the values of the other parameters. Figure 14(b) shows the mem-
brane ionic current density JB as calculated from the magnetic field. Also
shown for comparison are the membrane capacitive current density J, and
the total membrane current density J,. Recall that from Equation (11)
J,+J,=J,.

CONCLUSION
All the predictions of the volume conductor model are contained in the
filter function relationships, summarized in Tables 1 and 2. We have been
able to put these relationships in their relatively simple form by using the
mathematical techniques of Fourier transforms, filter functions, and the
convolution theorem: in essence we diagonalized the matrix representation of
the Greens function. One great advantage of this representation of the
solution is that it solves both the forward and inverse problems simulta-
neously. We can better see the physical behavior of the fields by taking the
small argument, or DC, limit of the filter functions. This limiting procedure
yields the core-conductor model results. The large argument limit of the filter
functions introduces exponential behavior which complicates the inversion of
the solution.
The physical processes involved in generating the magnetic field are
presented in two complementary but equivalent views, the first based on the
law of Biot and Savart and the second on Amperes law. Both points of view
predict that the magnetic field close to the axon will be well represented by
the first derivative of the transmembrane potential, but for large field point
radii the magnetic field will be both wider and smaller than the core-conduc-
tor model result. In relating the magnetic field to the transmembrane
potential, we found that there is a strong dependence of the amplitude of the
magnetic field on the axon radius and the internal conductivity. This strong
dependence is eliminated when relating the magnetic field to the current in
the axon, showing that the magnetic field is best suited for determining the
bioelectric currents. The magnetic field can also be used to study other
bioelectric phenomena, such as the ionic current density in the membrane.

We thank Dr. John Clark, Dr. John Barn& and Leonora Wikswo for their
comments and suggestions. This work was supported in part by the Office of
Naval Research under Contract NO001 4 - 82 -K-01 07, by NIH Grant I - R01
NS 19794 -01, and by the Vanderbilt Universiry Research Council.
36 JAMES K. WOOSLEY ET AL.

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