An Introduction to Core-conductor Theory

I often say when you can measure what you are speaking about
and express it in numbers you know something about it; but when
you cannot measure it, when you cannot express it in numbers, your
knowledge of it is of a meagre and unsatisfactory kind.
Lord Kelvin

Introduction

It would not be inaccurate to suggest that the manner in which electric current
flows along a semi-insulated conductor forms a central conceptual core in theoretical neuroscience. The passive properties of dendrites and the propagating
action potential are only explicable within the context of the cable equation.
The cable equation is a second order in space, first order in time, partial differential equation 1 . A form of this equation was probably first derived by William
Thomson (1824-1907), later Lord Kelvin, during his involvement with the laying,
design and analysis of the first trans-atlantic telegraph cables beginning in 1854.
The cable equation is based on the simple notion of transverse current leakage
between an inner and outer conductor due to an imperfect insulator as a consequence of the longitudinal flow of current within the inner conductor.

Assumptions
the cell membrane is conceived of as cylindrical boundary of finite thickness
separating the intra-cellular fluid (ICF) and the extra-cellular fluid (ECF).
the ECF and ICF are homogeneous, isotropic and Ohmic.
cylindrical symmetry is assumed for all variables subsequently defined i.e
f (r, x, , t) f (r, x, t) f (x, t) (r will obviously constant at the level of
the membrane).
an electrostatic (quasi-static) approximation will be adequate.
1

mathematicians would call this a parabolic partial differential equation, physicists would be
more inclined to call it a diffusion or heat equation

 currents in the inner and outer conductors will flow in the longitudinal (x)
direction only.
trans-membrane current flows in the radial (r) direction only

inner conductor

insulator (membrane)

a
outer conductor

Figure 1: Cross-section of the model cable illustrating the geometry of the inner
conductor, insulator and outer conductor.

Definitions
Io (x, t) = the total longitudinal current flowing in the +x direction in the
outer conductor. (A)
Ii (x, t) = the total longitudinal current flowing in the +x direction in the
inner conductor. (A)
Jm (x, t) is the membrane current density flowing from inner conductor to
outer conductor. (A m2 )
Km (x, t) is the membrane current per unit length flowing from inner conductor to outer conductor. (A m1 ) 2
Keo (x, t) is the current per unit length due to external sources applied in a
cylindrically symmetric manner to the outer conductor. (A m1 )
Kei (x, t) is the current per unit length applied to the inner conductor.
(A m1 )

2
this is the neurophysiological or physiologists convention and is often referred to as the
inward negative definition in that the trans-membrane current is defined as negative when it
flows from outer conductor to inner conductor. We will return to the issue of conventions at
the end of this chapter.

 Vi (x, t) is the potential of the inner conductor. (V )

Vo (x, t) is the potential of the outer conductor. (V )
Vm (x, t) = Vi (x, t) Vo (x, t) is the trans-insulator (i.e trans-membrane)
potential. (V )
ri is the resistance per unit length of the inner, cylindrically symmetric
conductor. ( m1 )
ro is the resistance per unit length of the outer, cylindrically symmetric
conductor. ( m1 )
a is the internal radius of the cylindrical shell. (m)

ri x

PSfrag replacements a

x
Figure 2: Geometry of the inner conductor
Based on these definitions and assumptions we are now able to represent concisely
in diagrammatic form the structure of the cable. This is illustrated in Figure 3.
Note that we have collapsed the cylindrical symmetry of the cable.

Derivation of the core-conductor equations

While it is quite easy to derive the cable equation directly without noting any
intermediate equations it is more profitable to derive a limited form of the cable
equation, called the core-conductor equations, which require no assumptions be
made about the structure of the cell membrane or insulator. The core-conductor
equations are easily derived by the use of Kirchoffs electrical circuit laws and a
spatially discrete cable.

K ei (x,t) x

K ei (x+x ,t) x
inner conductor

I i(x+x ,t)

I i(x,t)
(a)

ri x

Vi(x,t)

(b)

ri x

Vi(x+x ,t) ri x

K m(x+x ,t) x

K m(x,t) x

insulator (membrane)

I o(x,t)

I o(x+x ,t)

(c)

ro x

Vo(x,t)

ro x

(d)

Vo(x+x ,t) ro x
K eo(x+x ,t) x

K eo(x,t) x
outer conductor

x+ x

Figure 3: Schematic diagram of the model cable based on the definitions of the
proceeding sections

4.1

Kirchoff s First Law (conservation of current)

By referring to nodes (a) and (c) in Figure 3 and by noting that charge (current)
must be conserved we obtain

Ii (x, t) + Kei (x, t)x = Ii (x + x, t) + Km (x, t)x node (a)

Io (x, t) + Km (x, t)x = Io (x + x, t) + Keo (x, t)x node (c)

(1)
(2)

rearranging, dividing both equations through by x and taking the limit as

x 0 we obtain

Ii (x, t)
= Kei (x, t) Km (x, t)
x
Io (x, t)
= Km (x, t) Keo (x, t)
x
4

(3)
(4)

4.2

Kirchoff s Second Law (conservation of energy)

From Figure 3 and a simple application of Ohms Law we obtain

Vi (x, t) Vi (x + x, t) = ri xIi (x + x, t) nodes (a) and (b)

Vo (x, t) Vo (x + x, t) = ro xIo (x + x, t) nodes (c) and (d)

(5)
(6)

dividing through by x and taking the limit as x 0 we obtain

Vi (x, t)
= ri Ii (x, t)
x
Vo (x, t)
= ro Io (x, t)
x

(7)
(8)

However Vm = Vi Vo and thus by subtracting equation (8) from (7) we obtain

Vm (x, t)
= ro Io (x, t) ri Ii (x, t)
x

4.3

(9)

The core-conductor equations

The key to understanding the derivation of the core-conductor equations is in the

combination of equations (3), (4), and (9). Therefore differentiating equation (9)
with respect to x we obtain
2 Vm (x, t)
Io (x, t)
Ii (x, t)
= ro
ri
2
x
x
x

(10)

and substituting in equations (3) and (4) we obtain

2 Vm (x, t)
= (ro + ri )Km (x, t) ro Keo (x, t) ri Kei (x, t)
x2

(11)

This result is known as the core-conductor equation, and forms the skeleton
about which the full cable equation can be developed.
5

Derivation of the Cable Equation

In the derivation of the core-conductor equation no assumptions were made regarding the form of the total trans-membrane current per unit length, Km (x, t), in
that the electrical properties of the insulator (membrane) were ignored. However
early electro-physiological studies had established that, to good approximation,
the electrical properties of a cell membrane could be represented by an equivalent
circuit consisting of a capacitor and resistor in parallel. Still assuming cylindrical
symmetry then a small section of our model dendrite or axon (i.e cable) of length
x can be represented diagrammatically as in Figure 4. cm (x) is the capacitance of our small section of cable (which has a total surface area of 2ax).
Similarly rm (x) is the trans-membrane resistance of this small section of cable.
Vi (x, t)
inner conductor

Km (x, t)x

cm (x)

rm (x) = 1/
gm (x)

PSfrag replacements

outer conductor

Vo (x, t)

Figure 4: Equivalent electrical circuit for a cylindrically symmetric segment of

cell membrane. cm (x) is the total capacitance and rm (x) is the total (transmembrane) resistance of this small segment of cable.
By referring to Figure 4 the membrane current per unit length is then easily seen
to be given by the sum of a resistive and capacitive component

[Vi (x, t) Vo (x, t)] + [Vi (x, t) Vo (x, t)]/

rm (x)
t
capacitive
resistive

gm (x)
= cm (x) [Vi (x, t) Vo (x, t)] + [Vi (x, t) Vo (x, t)]
t
Vm (x, t)
= cm (x)
+ gm (x)Vm (x, t)
(12)
t

Km (x, t)x = cm (x)

As defined the membrane capacitance and membrane resistance are functions of

x. In order to take the limit as x 0 a more explicit relationship between
these quantities and x is required. Therefore assuming that cm (x) and rm (x)
are not functions of x we can define the the following relationships

cm (x) = cm x = 2aCm x
gm (x) = gm x = 2aGm x

(13)
(14)

where cm and gm are the capacitance and conductance per unit length respectively
(F m1 , S m1 ) and Cm and Gm are the capacitance and conductance per unit
area respectively (F m2 , S m2 ). By inserting these definitions into equation
(12) and dividing both sides through by x we obtain

Vm (x, t)
+ V m gm
t
Vm (x, t)
= cm
+ Vm /rm
t

Km (x, t) = cm

(15)
(16)

Be careful to note the meaning and units of rm in the last equation ! By substituting equation (16) into (11) and rearranging we obtain the following second-order
partial differential equation
Vm
rm
rm c m
=
t
ro + r i

2 Vm
+ ro Keo (x, t) + ri Kei (x, t) Vm
x2

(17)

This result is commonly referred to as the cable equation. Ignoring any external
current input (i.e Kei = Keo = 0) it is often convenient to rewrite the above
equation as

Vm
2 Vm
= 2c
Vm
t
x2

(18)

p
where c =
rm /(ro + ri ) and m = rm cm 3 . For reasons that will soon become
apparent c is called the cable space constant and m is called the membrane time
constant. Further by defining the dimensionless quantities X = x/c and T =
3

the student should note that rm cm = Rm Cm , where Rm = 1/Gm

t/m equation (18) can be written as the following parabolic partial differential
equation
2 Vm
Vm
=
Vm
T
X 2

(19)

Solutions of the partial differential equations so defined depend, in addition to

the stated electrical properties, on the initial conditions and the boundary conditions (i.e the value of Vm at either ends of the cable). Further, under certain
assumptions, equations (17-19) are able to describe the passive properties of multiply branched and tapered cables. The ability to incorporate multiply branched
cables is of particular interest in describing the passive electrical properties of
dendrites and axons. The interested reader should consult Rall (1989) for further
details.

5.1

The cable space constant

Let a constant trans-membrane current be applied at some point along the model
cable. As the membrane capacitance becomes polarised with the passage of time
the trans-membrane current density will become a function of axial distance only.
Vm
Thus for sufficiently long times t
0. This corresponds to a steady state.
For increasing distances away from this site of current injection the membrane
potential would be expected to decrease due to the continual trans-membrane
leakage of current. Thus at steady state the partial differential cable equation is
reduced to an ordinary differential equation (ODE)

2c

d 2 Vm
= Vm
dx2

(20)

As can be verified by direct substitution, a general solution to this ODE for

constant current applied at x = 0 is 4
Vm (x) = A exp[x/c ] + B exp[x/c ]

(21)

where A and B will depend on the boundary conditions. For boundary-value

4

This is easily derived by substituting exp[x] into the differential equation. The solution of
the resulting characteristic equation (which is a simple quadratic equation) defines the solution
basis set. The student should refer to any textbook on the solution of ordinary differential
equations to be reminded of the procedure.

solutions it is often convenient to use another pair of solutions to the second-order

ordinary differential equation which are (as can be verified by direct substitution)
Vm (x) = A cosh(x/c ) + B sinh(x/c )

(22)

where cosh(x) = (exp[x] + exp[x])/2 and sinh(x) = (exp[x] exp[x])/2.

The space constant is useful as a parameter for estimating how far an axon or
dendrite can propagate activity passively. Typical values for large myelinated
axons are 1 2 mm whereas for small unmyelinated axons or dendrites the
values range upwards from 30 m. We will now consider the determination of
the coefficients for the following three cases
infinite cylinder
a finite cylinder of length l with a sealed ends at x = 0 and x = l
a finite cylinder of length l with a sealed end at x = 0 and a killed end at
x = l (i.e the end at x = l is clamped to the resting membrane potential
Vm = 0)

5.1.1

infinite cylinder

For the case in which the cable extends from to + we require

V () = V () = 0
and thus for x 0 A = 0 and B = V0 in equation (21), where V0 is the membrane
potential at the site of the current passing electrode. Similarly for x 0 A = V 0
and B = 0 and thus
Vm (x) = V0 exp[|x|/c ]
for a constant current applied at x = 0.

(23)

5.1.2

finite cylinder, sealed end at x = l

The idea of the sealed end is that axial current is prevented from flowing across
a boundary. For the case now considered this boundary will be the end of the
cable at x = l. Further we will assume that a current, Iapp , is injected at x = 0
at the inner conductor and Vm has attained its steady-state value. By referring
to equation (9) it is seen that the following conditions must be satisfied

dVm
= ri Iapp
dx x=0

dVm
= 0
dx x=l

(24)

These conditions are known as Neumann conditions 5 . These boundary conditions

are used to solve for the coefficients A and B in our general solution (22) by
observing that
A
B
dVm
=
sinh(x/c ) +
cosh(x/c )
dx
c
c

(25)

Thus at x = 0 ri Iapp = B/c i.e B = ri Iapp c , whereas at x = l

0 = A sinh(l/c ) + B cosh(l/c )
which gives

A = B coth(l/c )
= ri Iapp c coth(l/c )
where coth(x) = cosh(x)/ sinh(x). Thus the solution is
Vm (x) = ri Iapp c [coth(l/c ) cosh(x/c ) sinh(x/c )]
by rearranging the above equation we get
5

More specifically Neumann conditions specify normal gradients on the boundary.

10

(26)

Vm (x) =

ri Iapp c cosh(l/c )
[cosh(l/c ) cosh(x/c ) sinh(l/c ) sinh(x/c )]
cosh(l/c ) sinh(l/c )

which can be rewritten as

Vm (x) =

Vm (0) cosh[(l x)/c ]

0 x l sealed end at x = l
cosh(l/c )

(27)

with Vm (0) = ri Iapp c coth(l/c ) and where we have used the identity
cosh(A B) = cosh(A) cosh(B) sinh(A) sinh(B).
1

0.8
sealed end
V/V(0)

0.6

0.4
infinite cable
0.2

killed end

0
0

0.2

0.4 0.6 0.8

1
1.2 1.4 1.6
x (in units of the space constant)

1.8

Figure 5: Attenuation of the membrane potential as a function of distance in a

cable with differing boundary conditions. The middle curve is a plot of equation
(23) for x 0. Distances are dimensionless and are in units of c . Note that the
attenuation of voltage with distance for the infinite cable lies in between that for
the sealed end (upper curve) and the killed end (lower curve) cables of length c .
Both finite cables have an electrotonic length, L = l/c , of unity.

11

5.1.3

finite cylinder, killed end at x = l

The idea of the killed end is that one of the boundaries is clamped to the resting
membrane potential, which in the absence of ionic batteries will be Vm = 0. Such
a boundary condition is known as a Dirichlet condition. Again we assume that a
current, Iapp , is applied to the inner conductor at x = 0 and that Vm has achieved
its steady state. Therefore our solution (equation (22) ) must satisfy the following
conditions

dVm
= ri Iapp
dx x=0
Vm (l) = 0

(28)

In a manner similar to that of the last section our solution is

Vm (x) =

Vm (0) sinh[(l x)/c ]

0 x l killed end at x = l
sinh(l/c )

(29)

with Vm (0) = ri Iapp c tanh(l/c ) and where we have made used of the identity
sinh(A B) = sinh(A) cosh(B) cosh(A) sinh(B).

5.2

The membrane time constant

Vm
= 0. The cable is then
The other extreme case of the cable equation is when x
collapsed to a single isopotential element and the cable equation then becomes
the following ordinary differential equation

dVm
= Vm
dt

(30)

which has the general solution

Vm (t) = A exp[t/m ]

(31)

where A is a constant that will depend on the initial conditions. Typical values
for m range from 5 50 ms.
12

5.3

The addition of ionic batteries

Trans-membrane current is mediated by the flux of ions, through protein pores,

between the intra-cellular and extra-cellular fluids. The flux of each type of
ion will depend on its electro-chemical gradient, which in turn depends on the
trans-membrane concentration gradients and the cell membrane potential, and
the selective permeability of the membrane to any given ionic species. Further,
the membrane permeability for any given ion may be regulated by one or more
conductances. All of these conductances may be regulated by endogenous ligands
(e.g neurotransmitters), membrane potential and the concentrations of other ions
in the extracellular and intracellular fluids. The total trans-membrane current
density will depend on the contributions of the individual ionic currents. Figure 6
displays a circuit diagram describing the electrical properties of nerve membrane
which incorporates the individual ionic currents.

intracellular
Jm (x, t)
PSfrag replacements

Gi (x, t)

Gj (x, t)

Gk (x, t)

Ei
Ji (x, t)

Ej
Jj (x, t)

Ek
Jk (x, t)

Cm
Jc (x, t)

extracellular
Figure 6: An equivalent electrical circuit of a small section of neuron membrane.
See text for definitions.
From Figure 6 it is easily ascertained that
Jm = J c +

Jk

(32)

where Jm is the total trans-membrane current flowing per unit area (A m2 ) which
Vm
is composed of a capacitive current (Jc = Cm t
) and k ionic currents (Jk ). For
each ionic conductance we have
13

Vi

2aJk (x)x
Ek = Vo
2aGk (x)x

and thus
Jk = (Vm Ek )Gk

(33)

where Ek is the equilibrium or reversal potential for the ionic species k and where
the explicit dependence of Jk and Gk on x has been removed. Ek represents the
membrane potential at which there is no net trans-membrane current for ions of
type k. Ek is determined for a given internal and external ion concentrations by
the Nernst equation

Ek =

RT Cko
ln i
F
Ck

(34)

where Cko and Cki are the external and internal ion concentrations respectively.
By noting that
Km (x, t)x = 2aJm (x, t)x
the cable equation (equation 17) can be re-written as (see also equation 12)
1 2 Vm
Vm X
=
C
+
(Vm Ek )Gk (x, t, Vm ) Jei (x, t)
m
2ari x2
t
k

(35)

where ro ri and Jei (x, t) is an internally applied (positive outwards) current

density (A m2 ). Note that the dependence of trans-membrane ionic conductances on the membrane potential, time and space is made explicit. There may
be more than one conductance associated with any given ion. The essence of
the Hodgkin-Huxley equations, which describe mathematically the form of the
action potential, is in the removal of any explicit time dependence of the ionic
conductances by the use of empirically determined auxiliary differential equations.

14

5.3.1

A note on conventions

op
By lumping all trans-membrane currents into a single term, Jion
, and assuming
spatial homogeneity equation (35) becomes

Cm

dVm
op
+ Jion
= Jei
dt

which implies that an externally applied current (outward positive or inward

negative) will depolarise the cell membrane whereas transmembrane ion currents
(outward positive) will hyperpolarise the cell membrane. The superscript, op,
op
ip
indicates the outward positive convention. By noting that Jion
= Jion
we
can rewrite the above equation so that all currents have a consistent effect of the
membrane potential i.e

Cm

dVm
ip
= Jion
+ Jei
dt

P
ip
where Jion
= k (Ek Vm )Gk . It is important to appreciate that such a convention is of no conceptual significance rather it allows all currents to be treated
consistently without making a special case for ionic currents.

References
Johnston D and Maio-sin Wu S. (1995). Foundations of Cellular Neurophysiology, MIT Press, Cambridge,MA.
Rall, W. (1989). Cable theory for dendritic neurons, in C. Koch and I.
Segev (eds), Methods in Neuronal Modeling, MIT Press, Cambridge, MA,
chapter 2, pp. 9-62.
Weiss J L (1996). Cellular Biophysics, Volume 2: Electrical Properties,
MIT Press, Cambridge, MA.

15