1. Consider the source-fiber geometry of Fig. 7.5 of Plonsey and Barr (shown on p.

8 of Lecture
#19), such that the extracellular potential is given by Eqn. (7.70):
I0
φe = ,
4πσ e r

where r is the distance from the source to the fiber at the axial distance z from the middle of
the fiber. h is the distance between the source and the fiber where the source is
perpendicular to the fiber, i.e., at z = 0 .
Find:
a) the extracellular potential as a function of z , i.e., φe ( z ) ,
∂ 2φe
b) the activation function ,
∂z 2
c) the positions of the peaks of the resulting regions of initial depolarization and
hyperpolarization predicted by the activation function, stating which are a peak of
depolarization and which are peaks of hyperpolarization, and

d) the positions of the boundaries between the initial depolarized and hyperpolarized regions.

a. The distance r from the electrode to the position z on the fiber is:

r = h2 + z2 .

Therefore the extracellular potential at position z on the fiber is:

I0 1
φe ( z ) = .
4πσ e h + z2
2

∂ 2φe
b. The activation function is then:
∂z 2

∂φe I z
=− 0 ⇒
∂z 4πσ e ( h + z 2 )3 2
2

∂ 2φe I 0 3z − ( h + z )
2 2 2

=
∂z 2 4πσ e ( h 2 + z 2 )5 2

I0 2z 2 − h2
= .
4πσ e ( h 2 + z 2 )5 2
 ∂ 2φe
c. The peaks of the activation function are located where its derivate equals zero:
∂z 2

∂ 3φe I 0 9 zh 2 − 6 z 3
= =0
∂z 3 4πσ e ( h 2 + z 2 )7 2

9 zh 2 − 6 z 3
⇒ =0
(h + z2 )
2 72

⇒ 9 zh 2 − 6 z 3 = 0
⇒ z 3 = 23 zh 2
⇒ z = 0, ± 3
2 h.

z = 0 is the location of the peak of the hyperpolarized region, and z = ± 3

2 h are the locations
of the peaks of the flanking depolarized regions.

d. The boundaries between the depolarized and hyperpolarized regions are located where the
activation function equals zero:

∂ 2φe I0 2 z 2 − h2
= =0
∂z 2 4πσ e ( h 2 + z 2 )5 2

2 z 2 − h2
⇒ =0
( h2 + z 2 )
52

⇒ 2z2 − h2 = 0
⇒z=± 1
2 h.
2. Consider the case of a pair of extracellular stimulating electrodes that are placed such that a
line drawn between the two electrodes is parallel to the fiber axis, with the distances between
the electrodes and the fiber shown in the figure below. Simultaneously, both electrodes start
injecting an equal positive current of I 0 .

z
0.5 cm
I0 I0
1.0 cm 1.0 cm
Find the resulting:
a) extracellular potential as a function of z , i.e., φe ( z ) ,
∂ 2φe
b) activation function ,
∂z 2
c) positions of the peaks of the consequent regions of initial depolarization and
hyperpolarization predicted by the activation function, stating which are a peak of
depolarization and which are peaks of hyperpolarization, and

d) positions of the boundaries between the initial depolarized and hyperpolarized regions.
(25 pts)

a) The distances r1 and r2 from the electrodes to the position z on the fiber are:

r1 = h 2 + ( z − d ) = ( 0.5) + ( z − 1.0 ) ; r2 = h 2 + ( z + d ) = ( 0.5) + ( z + 1.0 ) . .

2 2 2 2 2 2

Therefore the extracellular potential at position z on the fiber is:

I0 1 I0 1
φe ( z ) = + .
4πσ e ( 0.5) + ( z − 1.0 )
2 2 4πσ e ( 0.5) + ( z + 1.0)
2 2

∂ 2φe
b) The activation function is then:
∂z 2

∂φe I
=− 0
( z − 1.0) I
− 0
( z + 1.0) ⇒
∂z 4πσ e
(( 0.5) + ( z − 1.0) ) 4πσ e (( 0.5) + ( z + 1.0)2 )
2 32
2 2 32
 ⎡ ⎤
∂ 2φe I0 ⎢ 3z ( z − 1.0 ) 1 3 ( z − 1.0 ) ⎥
= − −
⎢
( ) (( 0.5) ) (( 0.5) )
52⎥
∂z 2
4πσ e ( 0.5) 2 + ( z − 1.0 ) 2 52 2
+ ( z − 1.0 )
2 32 2
+ ( z − 1.0 )
2
⎢⎣ ⎥⎦
⎡ ⎤
I0 ⎢ 3z ( z + 1.0 ) 1 3 ( z + 1.0 ) ⎥
+ − +
⎢
( ) ((0.5)
) ((0.5) + ( z + 1.0) )
52⎥
4πσ e ( 0.5) 2 + ( z + 1.0 ) 2 52 2
+ ( z + 1.0 )
2 32 2 2
⎢⎣ ⎥⎦

=
I ⎢
⎡
0
3z ( z − 1.0 )
−
(( 0.5) + ( z − 1.0) ) − 3 ( z − 1.0) 2 2 ⎤
⎥
⎢⎣ ( ) (( 0.5) + ( z − 1.0) ) (( 0.5) + ( z − 1.0) )
4πσ ⎢ ( 0.5) + ( z − 1.0 ) 2 2 52 2 2 52 2 2 52⎥
e
⎥⎦

+
I ⎢
0
⎡
3z ( z + 1.0 )
−
((0.5) + ( z + 1.0) ) + 3( z + 1.0) 2 2 ⎤
⎥
⎢⎣ ( ) ((0.5) + ( z + 1.0) ) ((0.5) + ( z + 1.0) )
4πσ ⎢ ( 0.5) + ( z + 1.0 ) 2 2 52 2 2 52 2 2 52⎥
e
⎥⎦
⎧ ⎫
I0 ⎪ 2 z 2 − 4 z + 74 2 z 2 + 4 z + 74 ⎪
= ⎨ + 52⎬
.
(
4πσ e ⎪ ( 0.5) 2 + ( z − 1.0 ) 2
) ((0.5) )
52
+ ( z + 1.0 ) ⎪
2 2

⎩ ⎭

This activation function is plotted in the figure below.

c) The peaks of the activation function ∂ 2φe ∂z 2 are located where its derivate equals zero:

⎡ ⎤
∂ 3φe I0 ⎢ 10 z 2 ( z − 1.0 ) 4z ⎥
= − +
(
∂z 3 4πσ e ⎢ ( 0.5) 2 + ( z − 1.0 ) 2
) ((0.5) )
72 52⎥
+ ( z − 1.0 )
2 2
⎢⎣ ⎥⎦
⎡ ⎤
I0 ⎢ 20 z ( z − 1.0 ) 4 ⎥
+ −
⎢
( ) (( 0.5) )
52⎥
4πσ e ( 0.5) 2 + ( z − 1.0 ) 2 7 2 2
+ ( z − 1.0 )
2
⎢⎣ ⎥⎦
4 ( z − 1.0 )
35
I
− 0
(
4πσ e ( 0.5)2 + ( z − 1.0 ) 2 7 2
)
⎡ ⎤
I0 ⎢ 10 z 2 ( z + 1.0 ) 4z ⎥
+ − +
(
4πσ e ⎢ ( 0.5) 2 + ( z + 1.0 ) 2
) (( 0.5) )
72 52⎥
+ ( z + 1.0 )
2 2

⎣⎢ ⎦⎥
⎡ ⎤
I0 ⎢ 20 z ( z + 1.0 ) 4 ⎥
+ − +
(
4πσ e ⎢ ( 0.5) 2 + ( z + 1.0 ) 2
) (( 0.5) )
72 52⎥
+ ( z + 1.0 )
2 2

⎣⎢ ⎦⎥

−
I0 ( z + 1.0 ) 35
4
= 0.
( )
4πσ e ( 0.5) + ( z + 1.0 ) 2 7 2
2
 ⎧ ⎫
∂ 3φe I 0 ⎪ ( −6 z + 18 z − 4 z + 4 ) ( −6 z − 18 z − 4 z − 4 ) ⎪
3 2 63 15 3 2 63 15

⇒ 3 = ⎨ + ⎬
∂z
⎩ (
4πσ e ⎪ ( 0.5 )2 + ( z − 1.0 )2 7 2 ( 0.5 ) + ( z + 1.0 )
2 2 72
⎪
⎭ ) ( )
=
I0 ( −6 z 3
( )
+ 18 z 2 − 634 z + 154 ) ⋅ ( 0.5 ) + ( z + 1.0 )
2 2 72

e
(
4πσ ( 0.5 ) 2
+ ( z − 1.0 ) ) ⋅ ( ( 0.5 )
+ ( z + 1.0 ) )
2 72 2 2 72

I ( −6 z − 18 z − z − ) ⋅ ( ( 0.5 ) + ( z − 1.0 ) )
3 2 63 15 2 2 72
4 4
+ 0
=0
(
4πσ ( 0.5 ) + ( z − 1.0 )
e
) ⋅ (( 0.5) + ( z + 1.0) )
2 2 7 2 2 2 7 2

⇒ ( −6 z 3 + 18 z − z + ) ⋅ ( ( 0.5 ) + ( z + 1.0 ) )
2 63 15 2 2 7 2
4 4

+ ( −6 z − 18 z − z − ) ⋅ ( ( 0.5 ) + ( z − 1.0 ) ) = 0
3 2 63 15 2 2 72
4 4

⇒ ( −6 z 3 + 18 z − z + ) ⋅ ( ( 0.5 ) + ( z + 1.0 ) )
2 63 15
2 2 2 7
4 4

− ( −6 z − 18 z − z − ) ⋅ ( ( 0.5 ) + ( z − 1.0 ) ) = 0.
3 2 63 15
2 2 2 7
4 4

There are seven finite, real solutions to this equation. z = 0 cm corresponds to a local
minimum in the middle region of depolarization. The middle region of depolarization has two
peaks at z = ±0.34 cm, shown by the dashed red lines in the figure below. z = ±1.0 cm are the
locations of the peaks of the flanking hyperpolarized regions, shown by the dashed magenta
lines in the figure below, and z = ±1.61 cm are the locations of the peaks of the outer
depolarized regions, shown by the dash black lines in the figure below.

d) The boundaries between the depolarized and hyperpolarized regions are located where the
activation function equals zero:

⎧ ⎫
∂ 2φe I0 ⎪ 2 z 2 − 4 z + 74 2 z 2 + 4 z + 74 ⎪
= ⎨ + 52⎬
=0
(
∂z 2 4πσ e ⎪ ( 0.5 )2 + ( z − 1.0 )2
) (( 0.5) )
52
+ ( z + 1.0 ) ⎪
2 2

⎩ ⎭

(
⇒ ( 2 z 2 − 4 z + 74 ) ⋅ ( 0.5 ) + ( z + 1.0 )
2
) + ( 2 z + 4 z + ) ⋅ (( 0.5) + ( z − 1.0) )
2 52 2 7
4
2 2 52
=0

⇒ ( 2 z 2 − 4 z + 74 ) ⋅ ( 0.5 )
2
( 2
+ ( z + 1.0 ) ) − ( 2 z + 4 z + ) ⋅ ( ( 0.5 ) + ( z − 1.0 ) )
2 5 2 7
4
2 2 2 5
= 0.

There are four finite, real solutions to this equation, shown by the dashed green lines in the
figure below. A middle region of depolarization (activation function > 0) is bounded by
z = ±0.67 . Two flanking hyperpolarized regions (activation function < 0) extend from
z = 0.67 to z = 1.35 and from z = −0.67 to z = −1.35 . Two outer depolarized regions
(activation function > 0) extend from z = ±1.35 cm out to z = ±∞ .
 2 2
∂ φe/∂z (normalized)

−8
−6
−4
−2
0
2
4

−5
0
z (cm)
5
3. Consider the case of a pair of extracellular stimulating electrodes that are placed on opposite
sides of a fiber such that a line drawn between the electrodes is perpendicular to the axis of
the fiber, as depicted below.
I0

rP
hP

hN
rN

−I0
One electrode, at a distance hP from the fiber, is approximated by a positive point current
source I 0 , while the other electrode, at a distance hN from the fiber, can be approximated by
an equal and opposite point current source − I 0 .
a) Find the extracellular potential as a function of z , i.e., φe ( z ) ,
∂ 2φe
b) Find the activation function .
∂z 2
c) State the conditions on the values of hP and hN such that the membrane is depolarized in
the region directly between the electrodes, i.e., around z = 0 .
d) State the conditions on the values of hP and hN such that the membrane is hyperpolarized
in the region directly between the electrodes, i.e., around z = 0 . (15 pts)

a) The distances the electrodes to the position z on the fiber are:

rP = hP2 + z 2 and rN = hN2 + z 2 .

Therefore the extracellular potential at position z on the fiber is:

I0 1 I0 1 I0 ⎡ 1 1 ⎤
φe ( z ) = − = ⎢ − ⎥.
4πσ e hP2 + z 2 4πσ e hN2 + z 2 4πσ e ⎢ hP2 + z 2 hN2 + z 2 ⎥⎦
⎣
 ∂ 2φe
b) The activation function is then:
∂z 2

⎡ ⎤
∂φe I z z
= 0 ⎢− + ⎥ ⇒
∂z 4πσ e ⎢ ( h 2 + z 2 )3 2 ( h 2 + z 2 )3 2 ⎥
⎣ P N ⎦

I 0 ⎢ 3 z − ( hP + z ) 3 z − ( hN + z ) ⎥
⎡ 2 2 ⎤
∂ 2φe
2 2 2 2

= −
∂z 2 4πσ e ⎢ ( h 2 + z 2 )5 2 ( hN2 + z 2 ) ⎥⎦
52

⎣ P

⎡ ⎤
I 0 ⎢ 2 z 2 − hP2 2 z 2 − hN2 ⎥
= − .
4πσ e ⎢ ( h 2 + z 2 )5 2 ( h 2 + z 2 )5 2 ⎥
⎣ P N ⎦

c) For the membrane to be depolarized initially in the region around z = 0 , the negative current
source must be closer to the fiber than the positive source, i.e., hN < hP .
This can be argued intuitively from the perspective that the net change in the extracellular field
at z = 0 will have the same sign as the electrode that is closer to the fiber, and depolarization of
the membrane is produced by a decrease in the extracellular potential.

This can be shown mathematically by finding under which condition the activation function is
positive around z = 0 , which produces depolarization. That is:

⎡ ⎤
∂ 2φe I 0 ⎢ 2 z 2 − hP2 2 z 2 − hN2 ⎥
= − >0
∂z 2 4πσ e ⎢ ( h 2 + z 2 )5 2 ( h 2 + z 2 )5 2 ⎥
⎣ P N ⎦ z =0
−hP2 hN2
⇒ + >0
( hP2 ) ( hN2 )
52 52

1 1
⇒− + >0
(h 2 32
) ( hN2 )
32
P

1 1
⇒ >
(h ) 2 32
N (h ) 2 32
P

⇒ (h ) 2 32
N < (h ) 2 32
P

⇒ hN < hP .

d) For the membrane to be hyperpolarized initially in the region around z = 0 , the positive current
source must be closer to the fiber than the negative source, i.e., hP < hN .
This can be shown by the converse of the intuitive and mathematical arguments given for part c.
above.
4. Consider the case of an extracellular stimulating electrode pair that is well approximated by
an idealized dipole source p that is oriented parallel to a fiber, as depicted below.
z̄

h̄ r̄

θ
p̄

The extracellular potential that is created by the idealized dipole source is:
1 p ⋅ cos θ
φe = ,
4πσ e r2
where p is the magnitude of the dipole source p , r is the magnitude of the radial-distance
vector r , θ is the angle between the vectors p and r , and h and z are the magnitudes of the
distance vectors h and z , respectively.
a) Find the extracellular potential as a function of z , i.e., φe ( z ) ,
∂ 2φe
b) Find the activation function .
∂z 2
c) For the case where the dipole source is turned on suddenly, determine the regions of the
fiber that are initially depolarized and the regions that are initially hyperpolarized, giving
the values of z that form the boundaries of each region. (15 pts)

a) The radial distance r and the cosine of the angle θ vary with axial distance z according to:
z z
r = h2 + z2 and cosθ = = ..
r h + z2
2

Therefore the extracellular potential at position z on the fiber is:

p z
φe ( z ) = .
4πσ e ( h + z 2 )3 2
2

∂ 2φe
b) The activation function is then:
∂z 2
⎡ ⎤
∂φe I 1 3z 2
= 0 ⎢ − ⎥ ⇒
∂z 4πσ e ⎢ ( h 2 + z 2 )3 2 ( h 2 + z 2 )5 2 ⎥
⎣ ⎦
 ∂ 2φe I ⎡ 3z 6z 15z 3 ⎤
= 0 ⎢− 52 − + ⎥
∂z 2
4πσ e ⎢ (h2 + z2 ) (h + z ) (h + z ) ⎦
2 2 52 2 2 72⎥
⎣

I 0 ⎢ −3z ( h + z ) − 6 z ( h + z ) + 15z ⎥
⎡ 2 2 2 2 3⎤

=
4πσ e ⎢ ( h2 + z 2 ) ⎥
72
⎣ ⎦

I 0 ⎡⎢ −3z ( h + z ) − 6 z ( h + z ) + 15z ⎤⎥
2 2 2 2 3

=
4πσ e ⎢
⎣ ( h 2
+ z )
2 72 ⎥
⎦

I 0 ⎡⎢ −9 zh 2 + 6 z 3 ⎤⎥
= .
4πσ e ⎢ ( h 2 + z 2 )7 2 ⎥
⎣ ⎦
c) The boundaries between the depolarized and hyperpolarized regions are located where the
activation function equals zero:
⎡ ⎤
∂ 2φe I 0 ⎢ −9 zh 2 + 6 z 3 ⎥
= =0
∂z 2 4πσ e ⎢ ( h 2 + z 2 )7 2 ⎥
⎣ ⎦
−9 zh + 6 z
2 3
⇒ =0
(h + z )
2 2 72

⇒ −9 zh 2 + 6 z 3 = 0
⇒ z ( −9 h 2 + 6 z 2 ) = 0
⇒ z = 0, ± 3
2 h, and ± ∞.

Note that the solutions at z = ±∞ occur because the denominator is of a higher order that the
numerator in the activation function equation above.
Between z = −∞ & z = − 3
2h and between z = 0 & z = + 3
2h the activation function is
negative, which corresponds to regions of initial hyperpolarization.
Between z = − 23 h & z = 0 and between z = + 23 h & z = +∞ the activation function is
positive, which corresponds to regions of initial depolarization.
 2

1.5

∂ φe/∂z (normalized units)

1

0.5

−0.5
2

−1
2

−1.5

−2  
5h 3 0 3 5h
− h + h
2 z 2