We wish to confirm, by example, the prefatory claim that matrix
algebra is a useful means of organizing (stating and solving)
multivariable problems. In our first such example we
investigate the response of a nerve fiber to a constant current
stimulus. Ideally, a nerve fiber is simply a cylinder of radius
aa and length
ll that conducts electricity both
along its length and across its lateral membrane. Though we
shall, in subsequent chapters, delve more deeply into the
biophysics, here, in our first outing, we shall stick to its
purely resistive properties. The latter are expressed via two
quantities:
-
ρ
i
ρ
i
,
the resistivity in
Ω
cm
Ω
cm
of the cytoplasm that fills the cell, and
-
ρ
m
ρ
m
,
the resistivity in
Ω
cm
2
Ω
cm
2
of the cell's lateral membrane.
Although current surely varies from point to point along the
fiber it is hoped that these variations are regular enough to be
captured by a multicompartment model. By that we mean that we
choose a number NN and divide the
fiber into NN segments each of
length
lN
l
N
.
Denoting a segment's
- Definition 1:
axial resistance
R
i
=
ρ
i
lNπa2
R
i
ρ
i
l
N
a
2
and
- Definition 2:
membrane resistance
R
m
=
ρ
m
2πalN
R
m
ρ
m
2
a
l
N
we arrive at the lumped circuit model of
Figure 1. For a fiber in
culture we may assume a constant extracellular potential,
e.g., zero. We accomplish this by connecting
and grounding the extracellular nodes, see
Figure 2.
Figure 2 also
incorporates the exogenous disturbance, a current
stimulus between ground and the left end of the fiber. Our
immediate goal is to compute the resulting currents through each
resistor and the potential at each of the nodes. Our long--range
goal is to provide a modeling methodology that can be used
across the engineering and science disciplines. As an aid to
computing the desired quantities we give them names. With
respect to Figure 3,
we label the vector of potentials
x=
x
1
x
2
x
3
x
4
x
x
1
x
2
x
3
x
4
and the vector of currents
y=
y
1
y
2
y
3
y
4
y
5
y
6
.
y
y
1
y
2
y
3
y
4
y
5
y
6
.
We have also (arbitrarily) assigned directions to the currents
as a graphical aid in the consistent application of the basic
circuit laws.
We incorporate the circuit laws in a modeling methodology that
takes the form of a Strang
Quartet:
-
(S1) Express the voltage drops via
e=-Ax
e
A
x
.
-
(S2) Express Ohm's Law via
y=Ge
y
G
e
.
-
(S3) Express Kirchhoff's Current Law via
ATy=-f
A
y
f
.
-
(S4) Combine the above into
ATGAx=f
A
G
A
x
f
.
The AA in (S1) is the
node-edge adjacency matrix -- it encodes the
network's connectivity. The GG in (S2) is the diagonal matrix
of edge conductances -- it encodes the physics of the
network. The ff in
(S3) is the vector of current sources -- it encodes the
network's stimuli. The culminating
ATGA
A
G
A
in (S4) is the symmetric matrix whose inverse,
when applied to ff,
reveals the vector of potentials, xx. In order to make these ideas
our own we must work many, many examples.
With respect to the circuit of Figure 3, in accordance
with step (S1), we express the six potential
differences (always tail minus head)
e
1
=
x
1
-
x
2
e
1
x
1
x
2
e
2
=
x
2
e
2
x
2
e
3
=
x
2
-
x
3
e
3
x
2
x
3
e
4
=
x
3
e
4
x
3
e
5
=
x
3
-
x
4
e
5
x
3
x
4
e
6
=
x
4
e
6
x
4
Such long, tedious lists cry out for matrix representation, to wit
e=-Ax
e
A
x
where
A=-11000-1000-11000-1000-11000-1
A
-1100
0-100
0-110
00-10
00-11
000-1
Step (S2), Ohm's
Law, states:
The current along an edge is equal to the potential drop
across the edge divided by the resistance of the edge.
In our case,
y
j
=
e
j
R
i
,
j=1
,
3
,
5
and
y
j
=
e
j
R
m
,
j=2
,
4
,
6
y
j
e
j
R
i
,
j
1
,
3
,
5
and
y
j
e
j
R
m
,
j
2
,
4
,
6
or, in matrix notation,
y=Ge
y
G
e
where
G=1
R
i
0000001
R
m
0000001
R
i
0000001
R
m
0000001
R
i
0000001
R
m
G
1
R
i
0000
0
0
1
R
m
0000
00
1
R
i
000
000
1
R
m
00
0000
1
R
i
0
0000
0
1
R
m
Step (S3), Kirchhoff's Current Law, states:
The sum of the currents into each node must be zero.
In our case
i
0
-
y
1
=0
i
0
y
1
0
y
1
-
y
2
-
y
3
=0
y
1
y
2
y
3
0
y
3
-
y
4
-
y
5
=0
y
3
y
4
y
5
0
y
5
-
y
6
=0
y
5
y
6
0
or, in matrix terms
By=-f
B
y
f
where
B=-1000001-1-1000001-1-1000001-1
and
f=
i
0
000
B
-1000
00
1-1-10
00
001-1
-10
0000
1-1
and
f
i
0
0
0
0
Looking back at AA:
A=-11000-1000-11000-1000-11000-1
A
-1100
0-100
0-110
00-10
00-11
000-1
we recognize in BB
the transpose of AA. Calling it such, we recall
our main steps
-
(S1)
e=-Ax
e
A
x
,
-
(S2)
y=Ge
y
G
e
, and
-
(S3)
ATy=-f
A
y
f
.
On substitution of the first two into the third we arrive, in
accordance with
(S4), at
ATGAx=f
.
A
G
A
x
f
.
(1)
This is a system of four equations for the 4 unknown potentials,
x
1
x
1
through
x
4
x
4
.
As you know, the system
Equation 1
may have either 1, 0, or infinitely many solutions, depending
on
ff and
ATGA
A
G
A
.
We shall devote (FIX ME CNXN TO CHAPTER 3 AND 4) to an
unraveling of the previous sentence. For now, we cross our fingers and
`solve' by invoking the Matlab program,
fib1.m
.
This program is a bit more ambitious than the above in that it
allows us to specify the number of compartments and that
rather than just spewing the xx
and yy values it
plots them as a function of distance along the fiber. We note
that, as expected, everything tapers off with distance from
the source and that the axial current is significantly greater
than the membrane, or leakage, current.
We have seen in the previous example how a current source may
produce a potential difference across a cell's membrane. We
note that, even in the absence of electrical stimuli, there is
always a difference in potential between the inside and outside
of a living cell. In fact, this difference is the biologist's
definition of `living.' Life is maintained by the fact that the
cell's interior is rich in potassium ions,
K
+
K
+
,
and poor in sodium ions,
Na
+
Na
+
,
while in the exterior medium it is just the opposite. These
concentration differences beget potential differences under the
guise of the Nernst potentials:
- Definition 3:
Nernst potentials
E
Na
=RTFlog
[Na]
o
[Na]
i
and
E
K
=RTFlog
[K]
o
[K]
i
E
Na
R
T
F
[Na]
o
[Na]
i
and
E
K
R
T
F
[K]
o
[K]
i
where RR is the gas constant,
TT is
temperature, and FF is the
Faraday constant.
Associated with these potentials are membrane resistances
ρ
m
,
Na
and
ρ
m
,
K
ρ
m
,
Na
and
ρ
m
,
K
that together produce the
ρ
m
ρ
m
above via
1
ρ
m
=1
ρ
m
,
Na
+1
ρ
m
,
K
1
ρ
m
1
ρ
m
,
Na
1
ρ
m
,
K
and produce the aforementioned rest potential
E
m
=
ρ
m
E
Na
ρ
m
,
Na
+
E
K
ρ
m
,
Na
E
m
ρ
m
E
Na
ρ
m
,
Na
E
K
ρ
m
,
Na
With respect to our old circuit model, each compartment
now sports a battery in series with its membrane resistance,
as shown in Figure 6.
Revisiting steps
(S1-4) we note that in (S1) the even numbered voltage
drops are now
e
2
=
x
2
-
E
m
e
2
x
2
E
m
e
4
=
x
3
-
E
m
e
4
x
3
E
m
e
6
=
x
4
-
E
m
e
6
x
4
E
m
We accommodate such things by generalizing
(S1) to:
-
(S1') Express the voltage drops as
e=b-Ax
e
b
A
x
where bb is the vector of batteries.
No changes are necessary for (S2) and (S3). The final step now reads,
- (S4') Combine
(S1'),
(S2), and
(S3) to produce
ATGAx=ATGb+f
A
G
A
x
A
G
b
f
.
Returning to Figure 6,
we note that
b=-
E
m
010101
and
ATGb=
E
m
R
m
0111
b
E
m
01
01
01
and
A
G
b
E
m
R
m
01
11
This requires only minor changes to our old code. The new
program is called fib2.m
and results of its use are indicated in the next two
figures.
-
G. Strang,. (1986). Introduction to Applied Mathematics. Wellesley-Cambridge Press.