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Book - Solutions To Exercises PDF
Book - Solutions To Exercises PDF
In the following sections, solutions are given for some of the exercises.
Answer 2. 17
Answer 3. 0.686
Answer 5a. A · B = 8 − 10 + 2 = 0
Answer 5b. A · B = 0, so one knows the vectors are perpendicular. That’s because A · B =
|A||B|cos(θ). With neither |A| and |B| zero, the result can be zero only if cos(θ) = 0,
and that will occur only if the vectors are perpendicular (angle of 90 degrees).
Answer 7a. 4
Answer 7b. 0
Answer 9. 42.
Answer 11. The gradient points in the −r direction, i.e., in the “uphill” direction of ψ = 1/r.
Answer 15a. No; instead ∇(1/R) = −∇ (1/R). Note the negative sign.
1
2 SOLUTIONS FOR CHAPTER 1: VECTOR ANALYSIS
Answer 19a. With φ = 1, a constant, both the gradient and divergence of φ are zero. Therefore,
the first term on the left side of Eq. (1) and the first term on the right side become zero.
Answer 21. The key to these questions is taking advantage of the fact that surface S is a closed
surface. In (a), K · dS is the portion of the surface oriented in the direction of K. In (b),
r · K is a constant, so the result is a multiple of the integral of dS over the whole surface.
One may be able to see the general answers by picturing or computing the results for a
particular example, such as a cube.
Answer 1a. Because conductivity is the reciprocal of resistivity, the corresponding units will
be 1/(Ωcm) or Siemens/cm.
Answer 1b.
σ = 0.01 S/cm
Io
φ=
4πσr
Consequently,
−Io 1
J = −σ∇φ = ∇( )
4π r
Io
J= = 1r
4πr2
Answer 5. Take the derivative with respect to x twice to find the divergence. Then find where
the divergence is zero, which is x = 0.33 cm.
Answer 7.
Io ρ 4π50
φ= = √ = 7.1uV (microVolts)
4πr 4π 52 + 52
√
Answer 9a. The magnitude is 22 + 32 + 42 = 5.38 µA.
Answer 9b. The vector area is A = U × V /2. The division by 2 occurs because the cross
product’s magnitude is that of the parallelogram formed by the vectors, and the triangle
is half that. Performing the cross product, A = 0.5az .
Answer 11. Begin with the solution to Exercise 10. Twice differentiate with respect to x (not
r). The solution then is
Io 3(x − e)2
A(x) =
4πσ r5
.
Answer 13d. The answer is the same as that for 13c, since the same question is asked, using
different words.
Answer 15b. The maximum slope occurs at the peak of the gradient, so again at x = 0.
Answer 15c. The numerical value of the maximum slope is sech2 (x) with x = 0, i.e. 1.
Answer 17a. In equation form, the potential with resistivity 1 from source 1 can be written as
φ11 = (ρ1 /4π)(1/r1 ) and φ12 = (ρ2 /4π)(1/r1 ). The ratio φ12 /φ11 = ρ2 /ρ1 = 2. The
same argument will apply to potentials from the other source. Therefore, the ratio asked
equals 2.
Answer 17b. By definition J = −σ∇φ. With φ11 = (ρ1 /4π)(1/r1 ), the equation for J includes
no σ (or ρ) term. Therefore, the result is not a function of the resistivity, and the requested
ratio is 1.
Answer 17c. Again inspect the equation φ11 = (ρ1 /4π)(1/r1 ). Note that ∇φ has a direction
determined by r, not by ρ, since ρ is a constant insofar as the gradient operation, i.e., ρ
is not a function of x, y, z. Thus the dot product is 1, since the directions of J does not
change.
Answer 19c. At the origin, the divergence is greater than zero. This result is found by using the
definition of divergence, Eq. (1.23).
Answer 21. Hint: Ignore any interaction between the electrodes. Find the potential on an
imaginary electrode of the same size and location as a real arising from a point current
source at an electrode’s center. Make use of linearity to find results for two electrodes as
the sum of results for each one separately.
Answer 23d. There are sources, proportional to x. There is no source density at x = 0. Source
densities are negative when x < 0. Loosely speaking, these sources are associated with
currents from regions with more positive x to those with less positive x, and analogously
for negative x, but no current across the plane x = 0.
Answer 25. Hint: Since any potential field can be considered the superposition of point-source
fields, choose an arbitrarily located point source and demonstrate the validity of the
statement for this case.
BIOELECTRICITY: A QUANTITATIVE APPROACH 5
Answer 27. Hint: Think of the two sides of the tank as plates of a parallel plate capacitor, since
each one is a conducting region separated by a dielectric. Then use Gauss’s law with a
volume enclosing a square centimeter of surface at one interface, or use reference results
for capacitance with parallel plates.
Answer 29a. a = 10−3 cm. Thus ri = Ri /a2 = 102 /10−6 = 108 Ohms/cm.
Answer 31. The resistance of the total path is the resistance of the top membrane Rtop , the resis-
tance of the material inside the box, Rinside , and the resistance of the bottom membrane,
Rbottom .
Rtop = Rm /Atop . Atop , the area of the top surface, is Atop = a×L = 10×100×10−8 =
10−5 cm2 . Thus Rtop = 10000/10−5 = 109 Ohms. Rbottom = Rtop .
The resistance of the inside of the box is relatively insignificant, being Rinside = Ri ×
a/(L × a) = Ri /L = 100/(100 × 10−4 ) = 104 Ohms.
Thus the total resistance of the path Rtotal ≈ 2 × 109 , and the steady-state current is only
0.1/2 × 109 = 0.05 × 10−9 Amperes, or 0.05 nanoamperes.
Answer 33. Hint: use Gauss’s theorem. Draw an imaginary sphere just outside the inner sphere.
mv 2 3 RT
= (1)
2 2 No
Answer 8.
σ = (Di F Zi )Zi Ci (F/RT ) (2)
because E = −∇Φ and J = σE
Answer 9A. 9.456E-5 Amperes
Answer 9B. -8.78E-5 Amperes
Answer 10. 4E-7 moles/cm3
Answer 11. 1E-12 moles
Answer 12. 7.448E-6 moles/(cm2 /sec)
Answer 13. 2.98E-9 moles/sec
Answer 14. 29.56 cm/sec
Answer 15. 5.37E-5 seconds
Answer 16. 53.1 millivolts
Answer 17. -83.6 millivolts
Answer 18. -60 millivolts
Answer 19. Three ion species are not in equilibrium. (At this transmembrane potential, no
individual ion is in equilibrium.)
Answer 20. Two ion species are not in equilibrium. Cl− is in equilibrium.
Answer 21. 143.6 millivolts
Answer 22A. 24.1 Angstroms
Answer 22B. 1.15 microfarads per cm2
Answer 23. 4,000 square micrometers
Answer 24. 4.85E-5 microfarads
Answer 25. 558,036,000 Ohms
BIOELECTRICITY: A QUANTITATIVE APPROACH 7
Answer 6. N p
Answer 7. N q
Answer 8. 1
√
Answer 9. N pq
Answer 14b. Multiplying cell surface area (600 µm2 ) by the channel density (40 ch/µm), one
finds the number of K + channels to be 24,000. This number times p1 gives 3,525 channels
open.
28 Exercise [28] restatement from text: In the model cell, examine N a+ and K + currents
with Vm1 of -57 mV and Vm2 of -20 mV.
1
a. At t1 , what is IK ?
10 SOLUTIONS FOR CHAPTER 4: CHANNELS
1
b. At t1 , what is INa ?
2
c. At t2 , what is IK ?
2
d. At t2 , what is INa ?
32 Question 32, from text. In the model cell, evaluate the requested currents if Vm1 is
–55.5 mV, and Vm2 is –24.5 mV.
1
a. At t1 , what is IK ?
1
b. At t1 , what is INa ?
b
c. At tb , what is IK ?
b
d. At tb , what is INa ?
2
e. At t2 , what is IK ?
2
f. At t2 , what is INa ?
Answer 32a. 1.34E-10 Amperes
Answer 32b. -8.40E-12 Amperes
Answer 32c. 5.54E-10 Amperes
Detail of c:
1 The current desired is the product of the K + current through an open channel
times the expected number of open K + channels.
2 (Vm − EK ) = 55.5 mV, so the current through an open channel is Ichan =
γ × (Vm − EK ) = 0.555 picoamperes.
3 Because vm1
= 35.5 mV, αn is 0.2766 and βn is 0.0802, making n = 0.4515 at
t = ta (using the result of exercise 31).
4 The expected number of open channels is Nopen = Ntotal × n4 = 24000 ×
0.0416 = 997 channels.
5 Thus the current from potassium ions is 998 × 0.555 = 553 picoamperes.
Answer 32d. –9.27E-10 Amperes
Detail of 32d:
1 The current desired is the product of the N a+ current through one channel times
the expected number of open N a+ channels.
12 SOLUTIONS FOR CHAPTER 4: CHANNELS
Answer 34. No, as judged by these exercises, never the same sign. INa and IK have opposite
signs most of the time because (Vm − EK ) has a different sign from (Vm − ENa ). For
the sign to be the same, Vm would have to be outside the range EK to ENa .
Answer 2a. vm = Vm + 60
Answer 4. If
Im = IC + Iion
then
IC = Im − Iion
For IC to be zero, one must have
IC = Im − Iion = 0
Thus
Im = Iion
In an isolated patch,
Im = Is
so the condition for IC to be zero, and thus no Vm change, is
Is = Iion
Answer 13. Both set A and set B have the same value for Vm and hence IL . With the state
variables of Set A, INa is greater and IK is smaller than with the state variables of Set B.
Thus set A leads to a negative total ionic current (and thus a positive ∆Vm . In contrast, set
B leads to a positive total ionic current (since potassium dominates) and thus a negative
∆Vm .
Answer 16a. A computer procedure that computes every α and β for a given vm , including the
code needed for the special cases is:
i vm an bn am bm ah bh
0 0.000 0.06 0.13 0.22 4.00 0.07 0.05
1 5.000 0.08 0.12 0.31 3.03 0.05 0.08
2 10.000 0.10 0.11 0.43 2.30 0.04 0.12
3 15.000 0.13 0.10 0.58 1.74 0.03 0.18
4 20.000 0.16 0.10 0.77 1.32 0.03 0.27
5 25.000 0.19 0.09 1.00 1.00 0.02 0.38
6 30.000 0.23 0.09 1.27 0.76 0.02 0.50
7 35.000 0.27 0.08 1.58 0.57 0.01 0.62
8 40.000 0.32 0.08 1.93 0.43 0.01 0.73
9 45.000 0.36 0.07 2.31 0.33 0.01 0.82
10 50.000 0.41 0.07 2.72 0.25 0.01 0.88
Answer 18: Here are some suggestions to consider for writing code that will be usable for a
range of questions:
(1) Keep track of the iteration number (here called the “loop count”) as an integer. (Most
people do this instinctively.)
(2) Keep track of elapsed time as an integer. (Most program writers do not do this
instinctively.) Integer microseconds work well.
(3) Enter and record ∆t as an integer value (µsec). Whenever it is needed for computa-
tion, convert it then to a floating point form.
(4) Convert explicitly between the iteration number and the elapsed time, using ∆t.
(5) Keep track of the stimulus start/stop time as integer values.
(6) The goal of items 1 to 5 is to avoid issues of floating point round-off in keeping time,
identifying the stimulus starting time, etc.
(7) Keep track of the interval between times that output lines will be created with a
distinct integer variable. In general, do not write output for every iteration.
(8) If there is a stimulus, start the first stimulus at t = 0. Begin calculation, however,
before that time, so that a baseline can be established.
Generally the program is easiest to manage if elapsed time is converted explicitly from
the iteration number, because, in this way, the iteration number increases from zero as
expected, while elapsed time can begin with a negative value. Additionally, program
decisions about whether it is time to start or stop the stimulus, or to print output, can
be parametrized on the basis of elapsed time, rather than iteration number. Importantly,
interval ∆t can be varied, which often is desirable for testing, independently of the
specification of when the stimulus should occur or output lines should be displayed.
If there is a discrepancy between the output table as printed and your results, consider the
following aspects of how the table above is computed and verify their correspondence.
Here, line k + 1 is found from line k by:
(1) Using vm for line k to compute α and β values for line k, and ∆n, etc, for the time
of line k. Note that vm for line k = 1 is not used.
(2) Using n, m, h from line k (not line k + 1) to get currents and vdot at the time of line
k, and from these estimate vm at time k + 1. Note that newer values of n, m, h for
time k + 1 are not used.
(3) Being sure the stimulus does not extend into the period 150 to 200. That is, at time
150 the stimulus must stop.
After time 200 µsec, voltage vm does not continue to fall. Rather, it rises to a substantially
higher value in the course of generating an action potential.
dTime: 2 microseconds
StimAmplitude: 200
StimDuration: 150
Answer 22b. For duration 150, just-above-threshold stimulus at end of stimulus vm 6.46 mV.
Answer 24. The time duration until stable return within the envelope of the initial conditions is
24100 µsec.
Answer 25b. Some answers are more fun if you see for yourself. Inspect the return to baseline
carefully, perhaps by plotting vm (t).
Answer 26a. Is =50, 2nd AP at 16800 microseconds from start of first stimulus.
Answer 2a. Recall that the HH model has a linear response to transmembrane voltage changes
near the resting potential, so the membrane resistance can be found from the conductances
in the HH model. Use Table 13.5, which gives resting values of HH membrane, to find
the conductances (g values) of HH membrane at rest. Using those values,
Comparing the last two parts of the equation shows that rm = Rm /(2πa). Using the
result for Rm found in the answer to the preceding question, one has rm = Rm /(2π0.003)
or rm = 78, 248 Ωcm.
Answer 2c. The intracellular resistance per unit length characterizes the volume inside the
membrane in a way that takes into account the medium’s resistivity and the fiber’s radius.
After ri has been found, one need only make a later choice of a length L (distance along
the axis) to find resistance R of the segment. Thus resistance R will be
To make the last two terms equal, one makes ri = Ri /(πa2 ). In this fiber, the result is
ri = 5, 305, 169 ohm/cm.
Answer 2d. The connection of the intracellular resistance per length with the region inside the
membrane (in the preceding part) seems obvious, but the statement that “Extracellular
currents flow to twice the membrane radius” seems much less so. Indeed, in the absence
of a physical boundary of some kind, asserting that currents flow between radius a and
radius 2a is at best arbitrary. Even so, the statement recognizes that extracellular current
flows most intensely near the membrane, and it provides a specific basis for computing
re , even if it is only an estimate. Note that the cross-sectional area inside a is one-third
the cross-section between a and 2a, i.e., assuming 2a as a limiting radius for extracellular
current assumes that extracellular current flows through a greater cross-section, but still
within a radius of the fiber membrane. With this assumption, then using similar reasoning
to that of the previous part, one has
so that re = Re /[π((2a)2 − a2 )]. In this fiber, the result is re = 589, 463 ohm/cm.
BIOELECTRICITY: A QUANTITATIVE APPROACH 23
1 ∂Vm
Ie = [ + Ire ] (8)
(ri + re ) ∂x
1 ∂ 2 Vm
Im = ( − re ip ) (9)
(2πa)(ri + re ) ∂x2
where ip = 0 because there is no stimulus. As the question asks for Im , i.e., current per
cm2 , the Im equation requires that dimensions (e.g., of a) are in cm.
Answer 5a. Vm (x) = av tanh(x) has negative Potentials for x < 0, positive for x > 0. The
action potential is moving left, toward the more negative region.
Answer 5b. Ii (x) = −ai sech2 (x). Intracellular current is flowing left.
Answer 5c. Ie (x) = +ae sech2 (x). Extracellular current is flowing right.
Answer 5d. Im (x) = −2am sech2 (x) tanh(x). Membrane current is outward left, inward right.
Answers 5a–d. The expressions given depend on undefined constants av , ai , ae , and am , which
are proportionality constants that include such factors as the radius and the axial resis-
tances. The proportionality constants are not defined in detail here, so that this exercise
can focus on the curve shapes rather than magnitudes.
Answer 7. The difference in Question 7 (relative to Exercise 6) is that in Ex. 7 it is asked that
membrane current be found “per unit area.” Dividing the earlier result by the geometric
factors of one segment, one has
1 (v2 − 2v3 + v4 ) 2
3
Im = mA/cm (10)
(2πa∆x) (R + r)
Answer 8. The transmembrane current at crossing 0, at the left end, is the same as the intra-
cellular axial current between crossings 0 and 1. Thus I 0 = (v1 − v0 + rS)/(R + r)
milliamperes.
Answer 9. The expression must take into account the different manner for computing the deriva-
tive, and the different area of the segments at the fiber’s ends. Thus
2 (v1 − v0 + rS)
0
Im = (11)
(πa∆x) (R + r)
1
Answer 10a. Im = +1.085 mA/cm2
Answers 10A-b. Note that the computation can be simplified by substituting λ algebraically
into the equation for Im .
0
Answer 11a. Im = −6.508 mA/cm2
1
Answer 11b. Im = +6.508 mA/cm2
Answers 11a–b. These answers can be obtained by finding the solution again from scratch, or
as a variation on the answers to exercise 10, looking at the relationships of the given
voltages.
Answer 15b. The mesh ratio indicates instability. The bad outlook is tempered by the HH
membrane having fairly low Rm , and the resistance present along the extracellular as
well as intracellular axial direction, which is not taken into account in the mesh ratio
calculation.
Answer 15c. The manifestation of instability will be that, at some time step, values of Vm will
begin to oscillate with increasing amplitude, and after a few time steps will be out of the
computable range.
Answer 15d. Stability can be markedly improved by reducing ∆t to a lower value. For example,
making ∆t = 1 µsec would reduce the mesh ratio to 0.30, a change in mesh ratio that
likely would correspond to a stable calculation.
Answer 1. rheobase
Answer 2. threshold
Answer 3. chronaxie
Answer 5. a/ 1 − exp[−500/(RC)]
Answer 6. W
Answer 7. 1000U/R
Answer 8. 1000W/a
Answer 9. W
Answer 10. Begin by rearranging the Weiss–Lapicque equation so that e−T /τ is the only term
on the left. Rewrite the rearranged equation twice, once for each condition. Divide the
first equation by the second, and combine terms so that there is only one exponential term
on the left. Take the natural log, and rearrange the result to solve for τ . The result is:
I (a − i)
τ = (D − d)/ log
i (a − I)
Answer 21a. P (x) = S/ (h2 + (x − e)2 )
Answer 21b. A(x) = S(−1/r3 + 3(x − e)2 /r5 ). Note the marked difference between the
answer 21A (which naively seems to be the effect of the stimulus on the fiber), and
Answer 21B. Although A(x) is not an equation for Vm produced by the stimulus, it does
at least provide the initial direction of change for Vm .
Answer 22c. No. In stimulation, extent of change is important, but amplitude is also important,
and often more so. A significant point of interest is, however, that both source and sink
stimuli produce regions of positive A(x), as this result indicates that either polarity might
serve as a stimulus, but would stimulate to different regions of the fiber.
Answer 24. A(x) > 0 is a region where, at least initially, Vm is growing more positive, and
thus is more likely to reach threshold and initiate an action potential.
Answer 26. 4
Answer 27. The ratio L2 /L1 = 2. The answer shows that the portion of the fiber being depo-
larized by the stimulus is affected markedly by the location of the electrode relative to
the fiber.
Answer 28. The ratio L2 /L1 = 2. Note that a ratio other than 1 occurs because of a change
in the separation of the electrodes, not a change in orientation of the electrode pair. The
changed ratio indicates that more of the fiber is being affected, i.e., L2 > L1 .
Answer 39. The authors know the answer to this question, but have chosen not to include it
here, so as not to take away the reader’s enjoyment of finding the solution.
BIOELECTRICITY: A QUANTITATIVE APPROACH 29
Answer 1. At t = 10 milliseconds. On the graph, one sees the midpoint of the upstroke at about
32 mm. Dividing by the velocity (4 mm/msec), one finds that this action potential occurs
about 8 msec after the one at x = 0. Additionally, parameter t1 adds a 2 msec time delay
to the upstroke.
r . Matlab is the registered
Answer 2. See Figure 14.1. The graphs here were made with Matlab
trademark of The Mathworks Inc.
Answer 3. In an action potential the activation phase always comes first. Therefore, on the time
waveform, the activation phase (fast upstroke) is on the left (lower time). On an action
potential waveform traveling toward higher x, the activation phase is on the right (higher
x), since activation is moving to the right. Otherwise the action potentials appear largely
the same, though real action potentials may show more subtle differences.
Answer 4. See Figure 14.2. Here V̇mmax = 95.5 mV/ms. For comparison, measured transmem-
brane potentials from healthy tissue usually have V̇mmax between 100 and 200 mV/ms, as
abnormalities inevitably leads to a lower value. This (unnatural) AP also is on the slow
side.
Answer 5f. Activation crossing at x = 33.9 mm. Recovery crossing at x = 12.0 mm.
Answer 6b. With the velocity doubled, the AP waveform of Vm (x) occupies twice the distance
along x. Also, the leading edge is about twice the distance from the origin.
Answer 6d. With velocity doubled, the AP wave shape of Vm (t) remains the same, except that
the time of onset is shorter.
Figure 3. Ii (x) at time t = 10 msec. The vertical scale extends to 2.5E-4 mA.
BIOELECTRICITY: A QUANTITATIVE APPROACH 31
Answer 17. The transmembrane current is plotted in Figure 14.5. In the figure, each panel
shows the potential that arises from the lumped monopolar sources (the solid line) and the
potential that arises from the continuous sources (the dashed line). Panels A, B, and C are
for distances h of 0.1, 1.0, and 10.0 millimeters, as marked. All plots are for time t = 10
msec. When h is small, as in panel A, the approximation of the continuous by lumped
sources is poor, so lumpiness is striking. Conversely, as h increases to 1 millimeter, the
potentials as generated by the lumped sources are smoother. At 10 millimeters, the lumped
solution becomes similar to those from the continuous sources, even though distance h
still is less than the distance between source M1 and source M3 .
Answer 27. V = 7.162 × 10−5 V. This value was found using Eq. (8.71). The potentials are to
be found at points far enough from the cell that a simplification is possible, because the
solid angle is virtually a constant and so can be factored from the equation.
32 SOLUTIONS FOR CHAPTER 8: EXTRACELLULAR FIELDS
Figure 5. Extracellular potential Φe (x) from Lumped Monopolar and Continuous sources.
BIOELECTRICITY: A QUANTITATIVE APPROACH 33
Answer 1. Cardiac electrophysiology, compared to that of nerve. Correct statements are num-
bers 1, 3, and 5.
Answer 5. 10
Answer 8. 22 ± 3 ms
Answer 9. 44 ± 4 ms
Answer 15. 0.036 ± 0.005 V. Some variability depends on exactly how the edges are defined.
Answer 16. 40 ± 15 mV Making an enlarged figure is helpful. Note the calibration bar.
Answer 17. 15 ± 5 mV Making an enlarged figure is helpful. Note the calibration bar.
Answer 18. Constants in the differential equation are a = 1/(cqR) and b = u/(cq).
34 SOLUTIONS FOR CHAPTER 9: CARDIAC ELECTROPHYSIOLOGY
Answer 32. Increasing the velocity. Choose 1 and 4. Choices other than 4 are evaluated in the
preceding question, while 4 is evaluated in an earlier section.
Answer 49. Select 2 and 3. The design fails because requirement 1 is not satisfied.
Answer 50. Select 1 and 3. The design fails because requirement 2 is not satisfied.
Answer 52. –0.12 V. Note that ∆vm has a negative sign. It is the value across the excitation
wave, i.e., the value in the tissue ahead of the excitation wave minus the value in the
region the excitation wave has passed.
Answer 75a.
1 σM ∇φH · dS Hin 1
−σφ∇2 dV = − σM φH ∇ · dS Hin
v r H r H r
1 1
− σM φL ∇ · dS Lin − σL φL ∇ · dS L
L r L r
1
− σM φB ∇ · dsB (13)
B r
In Eq. (??), vector surface element dS Hin points into the heart region H, vector surface
element dS B points out of the body surface, vector surface element dS Lin points into
the lung region L, and vector surface element dS L points out of the lung region L. The
lung conductivity is designated σL , while the remaining torso conductivity is described
by σM .
In getting Eq. (??) from Eq. (13.18), several particular aspects of current flow in the torso
volume conductor were used. First, ∇2 φ is zero at every point within the volume, since
the volume has been drawn in such a way as to exclude the active tissue in the heart and to
exclude regions where σ makes a transition (i.e., the site of secondary sources). Second,
current flow across the lung boundary is continuous. Finally, the normal component of
electric field, ∇φ · n, approaches zero as a point approaches the body surface, since the
exterior of the body surface is a nonconductor (hence no current can leave the volume
conductor).
Answer 75b.
1
−σφ∇2 dV = 4πδ(r)σB φb (14)
V r
where φb and σB are the potential and conductivity at the point b, where r = 0. This
point is chosen arbitrarily and located just inside the body surface. This is the point for
which potentials will be determined.
BIOELECTRICITY: A QUANTITATIVE APPROACH 37
Answer 75c.
∇φH · dS Hin 1
4πσB φb = σM − σM φH ∇ · dS Hin
H r H r
1 1
−(σL − σM ) φL ∇ · dS L − σM ΦB ∇ · dS B (15)
L r B r
where unit normal vector nH points out of heart region H, nL points out of lung region
L, and nB points out of the body region B. Vectors arH , arL , and arB are all of unit
magnitude and their directions are associated with the radius vectors (rH , rL , rB ) from
b to the heart, lung, and body surface, respectively.
Using the equation for the solid angle in the equation above (??) gives
σM σM ∇φH · dS H
φb = − φH dΩH −
4πσB H 4πσB H r
(σL − σM ) σM
+ φL dΩL + φB dΩB (17)
4πσB L 4πσB B
This expression gives the potential at one point on the body surface in terms of several in-
tegrals. The integrals thereby can be interpreted as identifying the major factors affecting
the potential at a body surface point.
The first two integrals take into account the potentials and the gradients of potential (i.e.,
the currents) that exist on the cardiac surface. Although shown here as two separate
integrals, it is useful to note that the potentials and gradients on the cardiac surface are
not independent of each other.
The third integral takes into account the lack of uniformity in the volume conductor.
Specifically, in this derivation the integral takes into account the different conductivity
assigned to the lung volume. In problems with other inhomogeneities, additional integrals
would be present to take into account their effects. For example, the skeletal muscle is
another inhomogeneity (which also is anisotropic) that is thought to play a significant
role in determining the body surface potentials.
Finally, the fourth integral shows that the potential at a point on the body surface also
depends on the potentials at other points on the body surface. Why is this so? The body
surface defines the interface between the conducting torso and the nonconducting air,
locating an obvious discontinuity in conductivity. Accordingly, secondary sources arise
at this surface and the potential of any surface point will depend on the field generated
by all such secondary sources. In fact, the secondary source distribution will be such
that the field it generates when added to the field from all other sources (i.e., applied
sources) must satisfy the aforementioned boundary condition, namely, ∇φ · an = 0 on
38 SOLUTIONS FOR CHAPTER 9: CARDIAC ELECTROPHYSIOLOGY
B. In general, the body surface sources cause the potentials measured there to be higher
than they otherwise would be, by approximately a factor of 2 or 3.
While the integrals in (14.17) arise from field theoretic considerations, note that the
contribution from the lung and body surface contains the secondary source double layers
described by (8.64). In the sense of (14.17) the first two terms could be thought to
represent primary sources, namely, a double layer (with strength σM φH , as shown in
the first integral) and a single layer (with strength σM ∇φH · an , as shown in the second
integral). However, such sources are actually equivalent sources, and (14.17) can only be
used to calculate fields outside the heart.
The integrals in the expression above make unequal contributions to the body surface
potentials. A first approximation to the resulting body surface potential’s variation from
point to point can often be obtained after ignoring the last two integrals altogether. Fur-
ther, because of their different r dependence, the contributions from the first and second
integrals may be quite unequal.
Answer 75e. The equation is the same except that the lung integral drops out.
BIOELECTRICITY: A QUANTITATIVE APPROACH 39
Answer 24. The exercises ask for portions of computer code that handle the Frankenhauser–
Huxley (FH) membrane model. The responses below are taken from a report by Chad
R. Johnson, which includes a complete program listing. (Johnson CR. 2005. A mathe-
matical model of peripheral nerve stimulation in regional anesthesia, pp. 359–367. PhD
dissertation, Department of Biomedical Engineering, Duke University.)
function [] = iFH(vm)
% IFH Initialize FH membrane model.
% iFH initializes the parameters and gating
% variables for the FH membrane model.
%
% INPUT vm: Transmembrane potential vector.
% -- PREAMBLE ------------------------------------------
global pFH;
global CONST;
global DATA;
global UDATA;
’mbeta’, 0, ...
’halpha’, 0, ...
’hbeta’, 0, ...
’nalpha’, 0, ...
’nbeta’, 0, ...
’palpha’, 0, ...
’pbeta’, 0, ...
’gates’, 0, ... % Gating variables.
’I’, 0); % Ionic currents.
(vm(pFH.Rv)-22.0)./ ...
(1.0-exp((22.0-vm(pFH.Rv))./3.0));
pFH.mbeta = pFH.Q10.mbeta.*0.4.* ...
(13.0-vm(pFH.Rv))./ ...
(1.0-exp((vm(pFH.Rv)-13.0)./20.0));
pFH.nalpha = pFH.Q10.nalpha.*0.02.* ...
(vm(pFH.Rv)-35.0)./ ...
(1.0-exp((35.0-vm(pFH.Rv))./10.0));
pFH.nbeta = pFH.Q10.nbeta.*0.05.* ...
(10.0-vm(pFH.Rv))./ ...
(1.0-exp((vm(pFH.Rv)-10.0)./10.0));
pFH.palpha = pFH.Q10.palpha.*0.006.* ...
(vm(pFH.Rv)-40.0)./ ...
(1.0-exp((40.0-vm(pFH.Rv))./10.0));
pFH.pbeta = pFH.Q10.pbeta.*0.09.* ...
(-25.0-vm(pFH.Rv))./ ...
(1.0-exp((vm(pFH.Rv)+25.0)./20.0));
IK = pFH.Q10.PK.*pFH.PK.*(pFH.gates.n(1).ˆ2.0).* ...
(pFH.Vrest.*pFH.F./pFH.D).* ...
((pFH.Ko-pFH.Ki.*exp(pFH.Vrest./pFH.D))./ ...
(1.0-exp(pFH.Vrest./pFH.D)));
IP = pFH.Q10.PNa.*pFH.PP.*(pFH.gates.p(1).ˆ2.0).* ...
(pFH.Vrest.*pFH.F./pFH.D).* ...
((pFH.Nao-pFH.Nai.*exp(pFH.Vrest./pFH.D))./ ...
(1.0-exp(pFH.Vrest./pFH.D)));
Il = pFH.Q10.gbarl.*pFH.gbarl.*(-pFH.El);
Iion = INa+IK+IP+Il;
while (abs(Iion)>CONST.myzero)
INa = pFH.Q10.PNa.*pFH.PNa.*(pFH.gates.m(1).ˆ2.0).* ...
pFH.gates.h(1).*(pFH.Vrest.*pFH.F./pFH.D).* ...
((pFH.Nao-pFH.Nai.*exp(pFH.Vrest./pFH.D))./ ...
42 SOLUTIONS FOR CHAPTER 12: FUNCTIONAL ELECTRICAL STIMULATION
(1.0-exp(pFH.Vrest./pFH.D)));
IK = pFH.Q10.PK.*pFH.PK.*(pFH.gates.n(1).ˆ2.0).* ...
(pFH.Vrest.*pFH.F./pFH.D).* ...
((pFH.Ko-pFH.Ki.*exp(pFH.Vrest./pFH.D))./ ...
(1.0-exp(pFH.Vrest./pFH.D)));
IP = pFH.Q10.PNa.*pFH.PP.*(pFH.gates.p(1).ˆ2.0).* ...
(pFH.Vrest.*pFH.F./pFH.D).* ...
((pFH.Nao-pFH.Nai.*exp(pFH.Vrest./pFH.D))./ ...
(1.0-exp(pFH.Vrest./pFH.D)));
Il = pFH.Q10.gbarl.*pFH.gbarl.*(-pFH.El);
Iion = INa+IK+IP+Il;
Answer 24B. Code to compute the membrane currents for the current membrane state:
function [] = iFH(vm)
% IFH Initialize FH membrane model.
% iFH initializes the parameters and gating
% variables for the FH membrane model.
%
% INPUT vm: Transmembrane potential vector.
% -- PREAMBLE -------------------------------------
global pFH;
global CONST;
global DATA;
global UDATA;
pFH.Q10.hbeta = 2.9.ˆ((pFH.T-293.15)./10.0);
pFH.Q10.nalpha = 3.2.ˆ((pFH.T-293.15)./10.0);
pFH.Q10.nbeta = 2.8.ˆ((pFH.T-293.15)./10.0);
pFH.Q10.palpha = 3.0ˆ((pFH.T-293.15)./10.0);
pFH.Q10.pbeta = 3.0.ˆ((pFH.T-293.15)./10.0);
pFH.gates.h(1).*(pFH.Vrest.*pFH.F./pFH.D).* ...
((pFH.Nao-pFH.Nai.*exp(pFH.Vrest./pFH.D))./ ...
(1.0-exp(pFH.Vrest./pFH.D)));
IK = pFH.Q10.PK.*pFH.PK.*(pFH.gates.n(1).ˆ2.0).* ...
(pFH.Vrest.*pFH.F./pFH.D).* ...
((pFH.Ko-pFH.Ki.*exp(pFH.Vrest./pFH.D))./ ...
(1.0-exp(pFH.Vrest./pFH.D)));
IP = pFH.Q10.PNa.*pFH.PP.*(pFH.gates.p(1).ˆ2.0).* ...
(pFH.Vrest.*pFH.F./pFH.D).* ...
((pFH.Nao-pFH.Nai.*exp(pFH.Vrest./pFH.D))./ ...
(1.0-exp(pFH.Vrest./pFH.D)));
Il = pFH.Q10.gbarl.*pFH.gbarl.*(-pFH.El);
Iion = INa+IK+IP+Il;
while (abs(Iion)>CONST.myzero)
INa = pFH.Q10.PNa.*pFH.PNa.*(pFH.gates.m(1).ˆ2.0).* ...
pFH.gates.h(1).*(pFH.Vrest.*pFH.F./pFH.D).* ...
((pFH.Nao-pFH.Nai.*exp(pFH.Vrest./pFH.D))./ ...
(1.0-exp(pFH.Vrest./pFH.D)));
IK = pFH.Q10.PK.*pFH.PK.*(pFH.gates.n(1).ˆ2.0).* ...
(pFH.Vrest.*pFH.F./pFH.D).* ...
((pFH.Ko-pFH.Ki.*exp(pFH.Vrest./pFH.D))./ ...
(1.0-exp(pFH.Vrest./pFH.D)));
IP = pFH.Q10.PNa.*pFH.PP.*(pFH.gates.p(1).ˆ2.0).* ...
(pFH.Vrest.*pFH.F./pFH.D).* ...
((pFH.Nao-pFH.Nai.*exp(pFH.Vrest./pFH.D))./ ...
(1.0-exp(pFH.Vrest./pFH.D)));
Il = pFH.Q10.gbarl.*pFH.gbarl.*(-pFH.El);
Iion = INa+IK+IP+Il;
pFH.I.P = zeros(size(vm));
pFH.I.l = zeros(size(vm));
pFH.I.myel = zeros(size(vm));
function [] = gatesFH(vmNew,dt);
% GATESFH Update FH rate constants and gating variables.
% GATESFH(vmNew,dt) updates the membrane channel
rate
% constants using the vector of updated nerve fiber
% transmembrane potentials vmNew, then finds the gating
% variables for the next iteration given the integration
% time step dt.
% -- PREAMBLE ------------------------------------------
global pFH;