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Biomechanics, Muscle Fibers, and How to
Interface Experimental Apparatus to a Computer
Masataka Kawai

Biomechanics, Muscle
Fibers, and How to
Interface Experimental
Apparatus to a Computer
Masataka Kawai
Department of Anatomy and Cell Biology
The University of Iowa
Iowa City, Iowa
USA

ISBN 978-3-319-72034-0 ISBN 978-3-319-72036-4 (eBook)

https://doi.org/10.1007/978-3-319-72036-4

Library of Congress Control Number: 2018932165

© Springer International Publishing AG 2018

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of
the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recita-
tion, broadcasting, reproduction on microfilms or in any other physical way, and transmission or infor-
mation storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar
methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publica-
tion does not imply, even in the absence of a specific statement, that such names are exempt from the
relevant protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this
book are believed to be true and accurate at the date of publication. Neither the publisher nor the
authors or the editors give a warranty, express or implied, with respect to the material contained herein
or for any errors or omissions that may have been made. The publisher remains neutral with regard to
jurisdictional claims in published maps and institutional affiliations.

Printed on acid-free paper

This Springer imprint is published by Springer Nature

The registered company is Springer International Publishing AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
I could not tell how delighted and excited
I am about this new book by Masataka
Kawai! It is just what we have needed for
years, and it promises, at long last, to teach
the world about the complexities and
wonders of muscle — and how to deal with
it’s mechanical properties intelligently.
I also firmly believe that nobody is better
qualified to write on these topics and has
shown, through diligence and hard work,
the expertise that Kawai has. There is so
much to learn!
Marion J. Siegman, Ph.D.
Professor and Chair
Department of Molecular Physiology and
Biophysics
Thomas Jefferson University
Philadelphia, PA, U.S.A.
Preface – About the Author and STEM
Disciplines

The subject that I have been practicing longest is electronics. I was 5 or 6 when
my father, Shunji Kawai , taught me how to make a radio. In those days,
the radio consisted of 4–5 vacuum tubes (each about 15 cm tall with one used as a
rectifier) with lit filaments, and many additional parts. Radios were very expensive
(costing 11,000 Yen or $30 then, which translates to $1,700 in today’s US cur-
rency) and my father could not afford one (his monthly pay as a school teacher
was about 7,500 Yen), so he went to Nagoya City to learn how to build a radio. It
was fascinating to watch him make holes in an aluminum chassis with a hand-
powered drill, and attach parts with screws and nuts. He cut wires, put in insulat-
ing tubes, and soldered components together according to schematics. He taught
me how to perform all of these steps. Between us, we made ∼50 radios, and sold
them to villagers at a reduced price, which made us very popular. I did not know
how the radio works, but later made up my own theory because my science tea-
cher asked; subsequently, I confirmed my theory. In the meantime, my mother,
Yoshié , competed with my father as to what to teach me, and she taught me
how to knit. I ended up making a muffler and a sweater. Endless repetition of
(almost) the same step gave me patience and endurance when I later wired numer-
ous connections in computer circuits.
These experiences gave me a tremendous advantage over the years. The tall
vacuum tubes were gradually replaced with groove tubes (GT), miniature tubes,
transistors, low-cost integrated circuits (ICs), intermediate-level ICs (computer
chips), and large-scale ICs. At each step in this progression, elements became smal-
ler, less expensive, and more reliable. At each stage, I had the opportunity to wire
them to make functional units. When I was a college junior, our math professor,
Akihiro Nozaki , taught us how to write a computer program using an
assembler language. The mainframe was a decimal-based computer that used a
rotating disk for its immediate memory and therefore was very slow, but I thought
it was cool. In graduate school, my colleague Ron Taylor taught me how to write a
Fortran program, with which I modified a primitive word processing program and
used it for my PhD thesis with automated referencing, figure and table numberings.
My thesis advisor, Professor Erwin (Tack) Kuntz, encouraged me to use operational

vii
viii Preface – About the Author and STEM Disciplines

amplifiers, which are the basis of analog computers and essential for carrying out
biophysical measurements (Sect. 6.2.6). During my postdoctoral training, my senior
colleague, Professor Phil Brandt, obtained a 16-bit NOVA computer, and I was
able to interface my experiments (‘sinusoidal analysis’) to this computer. This
included D/A (digital-to-analog) conversion for experimental control, A/D conver-
sion for data acquisition, an interrupt service program, programming in both assem-
bler and compiler (Fortran) languages to control external devices and perform
experiments, and instant display of the results (Chap. 6). Luckily, the Fortran sys-
tem has survived, and I still use programs I developed more than 40 years ago.
My fascination with biology started when I entered elementary school. I raised
silkworm larvae, fed them mulberry leaves, and watched them grow. They made
cocoons, metamorphosed to adult moths, mated, and laid eggs. I watched these
eggs hatch and grow into larvae again. I was certain by then that in life there are
two sexes, and that their genes must be mixed to improve genes. I also caught but-
terfly larvae, fed them until they formed cocoons, and watched them metamor-
phose into butterflies. This was more satisfactory than chasing butterflies and
catching them, because they escaped or got damaged when caught. When I was
14, my parents sent me to Tokyo to live with my aunt’s family and to attend mid-
dle and high schools there. Apparently, my parents felt that chasing butterfly lar-
vae was not an adequate education for me. In high school, my biology teacher,
Mr. Minoru Okuyama , taught us developmental biology, and it was
exciting to learn how an animal is formed starting from an egg. It was interesting
to know that development of the individual roughly recapitulates the evolution of
the species. Later in college, I took biochemistry, and was fascinated to see the
Watson and Crick DNA structure. My mentor, Professor Kazutomo Imahori
, said that it was the “Schrödinger equation” in biology. I rushed to
make a molecular model of DNA using wooden balls and sticks, encouraged by
Professor Koujiro Iso . This was one of those “wow!” moments.
In the first year of high school, I became interested in differentiation and integra-
tion, and taught myself these subjects a couple of years ahead of the curriculum.
I derived the Taylor expansion series (Eq. 5.7), and a fundamental equivalence in
algebra (Euler’s formula): eiθ = cos θ + i sin θ (Eq. 5.6). Consequently, I was able
to follow the two-body problem in classical mechanics, and able to realize how a
planet moves around the sun (and wobbling of the sun), based on Newton’s law of
universal gravitation (force is inversely proportional to the square of the distance
between the two bodies) and acceleration (Eq. 1.7) by solving differential
equations set up based on these principles. My motivation came from a few hints
(dreams) our math teacher (Mr. Ichiro Takahashi ) and physics teacher
(Mr. Teizo Sasaki ) gave us during regular class hours, and from a
book of math formulas that my cousin, Mr. Toshisuke Kuniéda , had.
He was my roommate for about 12 months and taught me how to use the log table
for multiplication and power calculations. Consequently, I purchased a 7-digit log
table book, used an abacus to add (and multiply) the numbers, and calculated the
movements of planets. To get approximate answers, I used a slide rule. No hand-
held calculators were available in those days.
Preface – About the Author and STEM Disciplines ix

To open up a new field and perform new research, it is essential to develop a

new instrument. When I entered my profession, I was lucky that I possessed many
of the skills needed. My strategy and principle in developing an instrument for
biological research is to let experimenters focus only on biology. While doing
experiments, one must not worry about electronics, mathematics, computer opera-
tions, or any other technical details. That is how I have been designing my auto-
mated experimental systems.
One thing my father did not teach me was how to make money, although he had
a keen interest in it and dreamed of receiving a million Yen from me as a gift; it
was unfortunate that this never happened. American children are generally better
off in this respect, because they learn this skill from an early age. However, many
suffer from inadequate training in the STEM (science, technology, engineering, and
math) disciplines, which in the long run are more profitable than quick money-
making methods. Hopefully my book will stimulate the young generation to recog-
nize the importance of these disciplines. Although these may expose the human
race to immense danger at times, they will save it from extinction in the long run.
***
Dr. Masataka Kawai (spelled in Japanese) is a native of Japan, and
received his B.A. from The University of Tokyo (Pure and Applied Sciences
), Ph.D. from Princeton University (Biology; Princeton,
NJ, USA), attended Physiology Course at Marine Biological Laboratory (Woods
Hole, MA, USA), and postdoctoral training at Columbia University (Muscle
biology; New York City, USA), where he became a junior faculty member. He is
now a professor of Anatomy and Cell Biology at The University of Iowa, College
of Medicine, Iowa City, IA, USA. He has taught histology to medical and dental
students for 35 years; he also taught principles of scholarly integrity to graduate
and postdoctoral students. He has recently learned gross anatomy and teaches this
subject. His hobbies include playing the piano, singing Japanese and Western
lyrics, gardening, sky gazing, and travels.
Symbols and Abbreviations Used

Symbols Definition
A Magnitude (amplitude) of exponential process A (Eqs. 3.6, 3.10)
A(t) Concentration of cross-bridges in the attached state A
A1 Steady-state concentration of A
AC Cross-sectional area (Fig. 1.1)
AT Total concentration (of cross-bridges) (Eqs. 2.2, 2.9)
ATPase The ATP hydrolysis rate (Eq. 4.3)
ΔAHP Change in hydrophobic surface area (Eq. 4.11)
ΔAI Change in ionic surface area (Eq. 4.12)
α ≡ 2πa Apparent (observed) rate constant of exponential process A (Eq. 3.6,
3.10, 4.15, Fig. 4.13)
α Rate constant of detachment (A → B), Schemes 1, 2 (Figs. 2.1, 2.2)
α′ Rate constant of attachment (B → A), Schemes 1, 2 (Figs. 2.1, 2.2
B Magnitude (amplitude) of exponential process B (Eqs. 3.6, 3.10)
B(t) Concentration of cross-bridges in the detached state B
B1 Steady-state concentration of B
β ≡ 2πb Apparent rate constant of exponential process B (Eqs. 3.6, 3.10, 4.2)
β Rate constant of attachment (B → C), Scheme 2 (Fig. 2.2)
β′ Rate constant of detachment (C → B), Scheme 2 (Fig. 2.2)
C Magnitude (amplitude) of exponential process C (Eqs. 3.6, 3.10)
C Capacitance (electricity, Sec 6.1)
C(t) Concentration of attached cross-bridges in C (Eq. 2.12)
C1 Steady-state concentration of C (Eq. 2.12)
ΔCP Heat capacity change (Eqs. 4.8, 4.9)
CPU Central processing unit (Sect. 6.4.1)
D MgADP concentration (Eq. 4.1)
εr Dielectric constant of water, εr = 78.55 at 25 °C (Eq. 4.14)
ε(t) Step function (Sec. 3.9, Table 5.1)
F Force, Tension (Sect. 1.1)
F(t) Force time course (Eq. 3.10, Figs. 3.1, 4.9, 4.14)

xi
xii Symbols and Abbreviations Used

F1 First harmonic amplitude of sinusoidal force oscillation (Eqs. 3.5,

5.39, 5.40)
ΔG° Gibbs standard free energy change (Eq. 4.7)
γ ≡ 2πc Apparent rate constant of exponential process C (Eqs. 3.6, 3.10, 4.1)
γ Rate constant of C → A transition, Scheme 2 (Fig. 2.2)
γ′ Rate constant of A → C transition, Scheme 2 (Fig. 2.2)
η Viscous modulus, viscosity (Eq. 1.6)
ΔH° Helmholtz’s standard free energy
pffiffiffiffiffiffiffi
ffi (enthalpy) change (Eqs. 4.7, 4.8)
i imaginary number, i = − 1 (Sect. 5.2, Eq. 5.1)
I Identity matrix (math, Eq. 5.47)
I Current (electricity, Eq. 6.1)
Im Imaginary part (of a complex number, Eq. 5.1)
IS Ionic strength (Eq. 4.13)
K Matrix consisting of rate constants (Eq. 2.29)
K0 Association constant of MgADP to the myosin head (Eq. 4.1,
Fig. 4.7)
K1 Association constant of MgATP to the myosin head (Eq. 4.1,
Fig. 4.7)
k2 Rate constant of the cross-bridge detachment step 2 (Eq. 4.1,
Fig. 4.7)
k−2 Reversal rate constant of the step 2 (Eq. 4.1, Fig. 4.7)
K2 Equilibrium constant of the step 2, K2 ≡ k2/k−2
k4 Rate constant of the force generation step 4 (Eq. 4.2, Fig. 4.7)
k−4 Reversal rate constant of the step 4 (Eq. 4.2, Fig. 4.7)
K4 Equilibrium constant of the step 4, K4 ≡ k4/k−4
K5 Association constant of Pi to the myosin head (Eq. 4.2, Fig. 4.7)
k6 Rate constant of the ADP isomerization step 6 (Eq. 4.3, Fig. 4.7)
Kα Equilibrium constant of A ↔ B transition. Kα ≡ α/α′ (Scheme 2)
Kβ Equilibrium constant of B ↔ C transition. Kβ ≡ β/β′ (Scheme 2)
L Length, strain (Sect. 1.2)
L1 Amplitude of sinusoidal length oscillation (Eq. 3.5)
L1 Size of step length change (Eqs. 3.9, 3.10)
lD Debye length (Eq. 4.13)
λ Apparent rate constant, λ ≡ α + α′ (Eq. 2.3, Scheme 1)
λ2, λ3 Apparent rate constants (Eqs. 2.13, 2.14, 2.15)
Λ Diagonal matrix consisting of eigenvalues of matrix K (Eq. 2.37)
m Mass (Eq. 1.7)
ν Frequency (Eq. 3.2)
ω Angular frequency, ω = 2πν, where ν is frequency (Eq. 3.2)
P Phosphate concentration (Eq. 4.2)
Pi Phosphate
2πa ≡ α Rate constant of process A (phase 4) (Eqs. 3.6, 3.10)
2πb ≡ β Rate constant of process B (phase 3) (Eqs. 3.6, 3.10)
2πc ≡ γ Rate constant of process C (phase 2) (Eqs. 3.6, 3.10)
q A constant needed to calculate Debye length (Eq. 4.14)
Symbols and Abbreviations Used xiii

R Resistance (Fig. 6.1a, Eq. 6.1)

Re Real part (of a complex number, Eq. 5.1)
ρ0 Density of solvent (Eq. 4.14)
S Substrate (MgATP) concentration (Eq. 4.1)
ΔS° Standard entropy change (Eqs. 4.7, 4.9)
σ Stiffness, Elastic modulus (Eq. 1.5)
σ12 σ12 ≡ K1SK2/[1 + K1S(1 + K2)] (Eqs. 2.16, 4.2)
t Time
τ Time constant, τ ≡ 1/λ (Eqs. 2.7, 6.4)
T Absolute temperature (T ≡ 273.15K + θ), where θ is
Celsius temperature (Eq. 4.7)
Tcyc, T Duration of one cycle of sinusoidal oscillation, Tcyc ≡ 1/ν
(Eqs. 3.3, 5.25)
V Velocity (viscoelasticity, Eq. 1.6)
V Voltage (electrical potential, electricity, Eq. 6.1)
W Work (Eqs. 1.14, 1.16, 6.5)
X Probability of cross-bridges (column vector) (Eqs. 2.17–2.22, 2.28)
Y, Y(ω) Complex modulus, Viscoelasticity (Eqs. 3.5, 5.40)
Y∞, Y(∞) Complex modulus extrapolated to ω → ∞ (Eq. 3.7)
Z Impedance (Eq. 6.1)
* Complex conjugate (math, Eq. 5.2)
* Complement (digital electronics Sec. 6.3.3, Fig. 6.5c and d)
∴ Therefore
Important Constants in Biomechanics
cal = 4.1855 J
e = 2.71828182846
ep = 1.6022 × 10−19 C
ε0 = 8.854 × 10−12 C2J−1m−1
kB = 1.38066 × 10−23 JK−1
LA= 6.022 × 1023 mol−1
ln 10 ≡ loge10 = 2.302585
π = 3.14159265358979
R ≡ kBLA = 8.3144 Jmol−1K−1
T0 = 273.15 K
Conversion of work (J) to the heat (15°C cal)
Base of natural log
Proton charge
Permittivity of vacuum
Boltzmann’s constant
Avogadro number
Natural log of 10
Circumference ratio
Gas constant
Absolute temperature of H2O at freezing
point (1 atm)
xv
Contents

1 Basic Elements of Viscoelasticity. . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

1.1 Force . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2 Length and Velocity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.3 Elastic Modulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.4 Viscous Modulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.5 Mass . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.6 Viscoelasticity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.7 Combination of Viscoelastic Elements . . . . . . . . . . . . . . . . . . . . 4
1.8 Exponential Lead . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
1.9 Work and Work Absorption . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
1.10 Oscillatory Work (Work Generator, Exponential Delay) . . . . . . . 6
1.11 Plot of the Complex Modulus in the Complex Plane . . . . . . . . . . 6
1.12 Exercise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Reference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
2 Reaction Processes (Chemical Kinetics) and Their
Application to Muscle Biology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.1 Chemical Reactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.2 Two-State Model, Rate Constants, and the First-Order
Reaction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2.3 Three-State Model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2.4 Second Order Reaction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
2.5 Application of Scheme 3 to Elementary Steps
of the Cross-Bridge Cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
2.6 Rate Constant and the Activation Energy . . . . . . . . . . . . . . . . . . 16
2.7 More General Case: Multi-State Model with
Branch Pathways . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
2.8 Exercise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

xvii
xviii Contents

3 How to Characterize Chemical Reactions Occurring

in Muscle Fibers? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
3.1 Perturbation Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
3.2 Caged ATP and Caged Phosphate (Pi) Experiments . . . . . . . . . . 24
3.3 Step Length Change Experiments . . . . . . . . . . . . . . . . . . . . . . . . 24
3.4 Sinusoidal Length Change Experiments . . . . . . . . . . . . . . . . . . . 25
3.5 Step Force Change Experiments . . . . . . . . . . . . . . . . . . . . . . . . . 26
3.6 Other Perturbation Analysis Methods . . . . . . . . . . . . . . . . . . . . . 26
3.7 Effect of Series Compliance . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
3.8 Further Details of the Sinusoidal Analysis Method . . . . . . . . . . . 27
3.9 Correlation Between the Sinusoidal Analysis
and the Step Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
3.10 Exercise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
4 Structure and Function of Muscle Cells . . . . . . . . . . . . . . . . . . . . . . 33
4.1 Classifications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
4.2 Skeletal Muscle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
4.3 Molecular Makeup . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
4.4 Functional Characterization of Muscle Fibers . . . . . . . . . . . . . . . 36
4.5 Regulation of Contraction at the Sarcomere Level . . . . . . . . . . . . 37
4.6 Regulation of Contraction at the Cellular and
Organ Levels in Skeletal Muscles . . . . . . . . . . . . . . . . . . . . . . . . 38
4.7 Skeletal Muscle Cell’s Generation, Regeneration,
and Exercise Hypertrophy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
4.8 Cardiac Muscle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
4.9 Smooth Muscle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
4.10 Cross-Bridge Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
4.11 Elementary Step That Generates Force . . . . . . . . . . . . . . . . . . . . 45
4.12 Ionic Interaction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
4.13 Hydrophobic Interaction and the Temperature Effect . . . . . . . . . . 50
4.14 Ionic Strength Effect. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
4.15 kTR Measurements and Series Elasticity . . . . . . . . . . . . . . . . . . . 54
4.16 Thin Filament Extraction and Its Reconstitution . . . . . . . . . . . . . 56
4.17 Exercise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
5 Mathematics Needed to Solve Problems of Contraction . . . . . . . . . . 65
5.1 Equal Sign (=, ≡) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
5.2 Complex Numbers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
5.2.1 Polar Coordinates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
5.2.2 Multiplication of Complex Numbers with Unit
Length in Polar Coordinate . . . . . . . . . . . . . . . . . . . . . . . 68
5.3 Sinewave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
5.4 First Order Differential Equation. . . . . . . . . . . . . . . . . . . . . . . . . 69
5.5 Fourier Transform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Contents xix

5.6 Discrete Fourier Transform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

5.7 Laplace Transform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
5.8 Matrix Operations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
5.9 Exercise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
6 Computer Interfacing of Experimental Apparatus. . . . . . . . . . . . . . 77
6.1 Basic Elements of Electricity . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
6.2 Analog Circuits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
6.2.1 Voltage (V), Current (I), and Impedance (Z) . . . . . . . . . . . 78
6.2.2 Series Arrangements . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
6.2.3 Parallel Arrangements . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
6.2.4 Power Calculation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
6.2.5 Difference Amplifier . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
6.2.6 Operational Amplifier and a Simple Application. . . . . . . . 80
6.2.7 Examples of Operational Amplifier Circuits . . . . . . . . . . . 81
6.2.7.1 Weighted Sum . . . . . . . . . . . . . . . . . . . . . . . . . . 81
6.2.7.2 Second Order Low-Pass Filter . . . . . . . . . . . . . . 82
6.2.7.3 Bridge Amplifier . . . . . . . . . . . . . . . . . . . . . . . . 83
6.2.7.4 Power Supply. . . . . . . . . . . . . . . . . . . . . . . . . . . 83
6.2.8 Limitations of Operational Amplifiers . . . . . . . . . . . . . . . 84
6.3 Digital Circuits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
6.3.1 Binary, Decimal, and Hexadecimal Numbers . . . . . . . . . . 84
6.3.2 Integers and Floating Point Numbers . . . . . . . . . . . . . . . . 85
6.3.3 Boolean Algebra and Elements of Digital Circuits . . . . . . 85
6.3.4 Digital to Analog Conversion (DAC) . . . . . . . . . . . . . . . . 87
6.3.5 Analog to Digital Conversion (ADC) . . . . . . . . . . . . . . . . 87
6.4 Programing Computers to Perform Experiments
and Initial Analysis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
6.4.1 Hardware. Sharing a Data Line for CPU I/O. . . . . . . . . . . 88
6.4.2 Software, CPU Input and Output . . . . . . . . . . . . . . . . . . . 89
6.4.3 Interfacing Compiler and Assembler Programs . . . . . . . . . 89
6.4.4 Clock Circuit and Interrupt Request . . . . . . . . . . . . . . . . . 90
6.4.5 Interrupt Service Program . . . . . . . . . . . . . . . . . . . . . . . . 90
6.4.6 Experimental Apparatus. . . . . . . . . . . . . . . . . . . . . . . . . . 90
6.4.7 Overall Assembly of the Experimental System . . . . . . . . . 92
6.4.8 How to Make an Executable Program
From Source Codes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
6.5 Exercise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Appendix 1. Sample Input Program from a PCH Device NCH. . . . . . . . 97
Appendix 2. Sample Output Program to a PCH Device NCH . . . . . . . . . 98
Appendix 3. Sample Fortran Program to Interface
with PCHin and PCHout Programs. . . . . . . . . . . . . . . . . . . 99
xx Contents

Appendix 4. Programs to Set Up and Remove Interrupt Service . . . . . . 101

Appendix 5. An Example of Interrupt Service Program . . . . . . . . . . . . . 102
Appendix 6. Program to Accumulate (Signal Average) Length
and Force Time Course Data . . . . . . . . . . . . . . . . . . . . . . . . 104
Appendix 7. Hints and Answers to Exercise Questions . . . . . . . . . . . . . . 105
Units and Their Conversions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
Chapter 1
Basic Elements of Viscoelasticity

Abstract This chapter defines basic terminology associated with force measure-
ments, and includes length, force, elasticity, viscosity, viscoelasticity, mass, work
absorption, work generation, transfer function, complex modulus, exponential
advance, exponential delay, and how to plot the complex modulus data.

Keywords Length · force · elasticity · viscosity · viscoelasticity · work absorption ·

work generation · transfer function · complex modulus · exponential advance ·
exponential delay · Nyquist plot

1.1 Force

Normalized force (F) and normalized force change (ΔF)

F = ðforceÞ=AC ; ΔF = ðforce changeÞ=AC (1.1)

where AC is the cross-sectional area to which the force is applied (Fig. 1.1).
Sometimes “specific” is used instead of “normalized”. ΔF is called tension, stress,
or pressure. They do not depend on the physical size (AC) of the preparation. In the
following discussion, a word “normalized” is frequently abbreviated for simplicity.
Series arrangement
Force measured at any point in series is the same if there is no mass in between
(Fig. 1.2a):

F = F1 = F2 (1.2)

Parallel arrangement
Two forces (F1, F2) applied in parallel (in the same direction) are additive
(Fig. 1.2b):

F = F1 + F2 (1.3)

Force in the opposite direction has the opposite sign.

© Springer International Publishing AG 2018 1

M. Kawai, Biomechanics, Muscle Fibers, and How to Interface Experimental
Apparatus to a Computer, https://doi.org/10.1007/978-3-319-72036-4_1
2 1 Basic Elements of Viscoelasticity

Fig. 1.1 Length (L) and

cross-sectional area (AC) of a
preparation AC
L

Fig. 1.2 Series a and parallel a

b arrangements. Y is viscoe- F1 F2
lasticity without mass
Y F1 = F2

b
F1

F2 F1 + F2

1.2 Length and Velocity

Normalized length change (ΔL), and normalized velocity (V)

ðlength changeÞ ðvelocityÞ

ΔL = ; V= (1.4)
L L

where L is the length of the material (Fig. 1.1). ΔL is called “strain.” It does not
depend on the physical size (L) of the preparation.

1.3 Elastic Modulus

If an elastic material (preparation) is stretched by ΔL, and extra force develops by

ΔF, then the (Young’s) elastic modulus (σ) of this material is:

ΔF 1
σ≡ ; and c≡ (1.5)
ΔL σ

Equation 1.5 is Hooke’s law, and c is compliance. A spring is a typical elastic

element. Hence its shape symbolizes elasticity (Fig. 1.3c). If the stress for ΔL
and strain for ΔF are used, the elastic modulus does not depend on the physical
size (AC, L) of the preparation. If force change is used for ΔF, and length change
is used for Δx, then σ is called stiffness. Thus, the stiffness depends on the
physical size.
1.6 Viscoelasticity 3

Fig. 1.3 Elements of a b c d e

viscoelasticity. a Wall
(no possibility of movement).
b Work generator.
c Elasticity. d Viscosity.
e Mass

1.4 Viscous Modulus

If a viscous material is stretched, force generated (ΔF) is proportional to the velo-

city (V) of the stretch, where t is time (usually expressed in seconds or s):

dL ΔF
ΔF ≡ ηV ≡ η ; or η= (1.6)
dt V

where η is called the viscous coefficient or viscosity. Viscosity is analogous to the

friction on a piston, and is depicted as such (Fig. 1.3d). If the normalized velocity
and force changes are used, the viscous modulus does not depend on the physical
size (AC, L) of the preparation.

1.5 Mass

Force experienced (ΔF) by a mass is proportionate to its acceleration:

d2 L
ΔF ≡ m (1.7)
dt2

Equation 1.7 defines the mass (m), which is depicted in Fig. 1.3e. The mass
can be usually ignored on the discussion of viscoelasticity of muscle fibers, but it
becomes significant in wave propagation. Also, Eq. 1.7 becomes important when
a planet movement around the sun is discussed, where there is no possibility of
viscosity.

1.6 Viscoelasticity

A material with both elastic and viscous properties is called a viscoelastic material.
The viscoelasticity (Y) can be measured by applying sinusoidal length changes:

ΔLðtÞ = L1 expðiωtÞ = L1 ½ cosðωtÞ + i sinðωtÞ (1.8)

4 1 Basic Elements of Viscoelasticity
pffiffiffiffiffiffiffiffi
where i ≡ − 1 and indicates “imaginary number” (Sect. 5.2). If you are uncom-
fortable with the imaginary numbers, you can just take the real part of Eq. 1.8 and
think that you live in the real world. Viscoelasticity can be characterized by mea-
suring the force response to the length change:

ΔFðtÞ = F1 ðωÞexpðiωtÞ = F1 ðωÞ½ cosðωtÞ + i sinðωtÞ (1.9)

where L1 is the amplitude of the length change, ω ≡ 2πν is angular frequency, and
ν is the frequency of the length oscillation. ΔFðtÞ is called “force time course” or
“force transient.” The transfer function from the length change to the force change is:

ΔF F1 ðωÞ
YðωÞ ≡ = (1.10)
ΔL L1

In viscoelasticity, Y(ω) is called the complex modulus, which is a function of

frequency (ω or ν) and has both a real part and an imaginary part. The complex
modulus of an elastic element (Eq. 1.5) is σ (elastic modulus), and that of a
viscous element (Eq. 1.6) is ηωi (viscous modulus).

1.7 Combination of Viscoelastic Elements

If two viscoelastic elements (Y1, Y2) are arranged in parallel, the combined viscoe-
lasticity is (Fig. 1.4a):

Y = Y1 + Y2 (1.11)

If these elements are arranged in series, the combined viscoelasticity is (Fig. 1.4b):

Y1 Y2
Y= (1.12)
Y1 + Y2

Equations 1.11 and 1.12 can be proven by applying a displacement (ΔL) and
measuring the force change (ΔF) for each element and for both by incorporating
the principle of Fig. 1.2.
Fig. 1.4 a Parallel a
arrangement. b Serial Y1
arrangement

Y2

b
Y1 Y2
1.9 Work and Work Absorption 5

1.8 Exponential Lead

If an elastic element (σ) and a viscous element (ηωi) are combined in series
(Fig. 1.5c), then the combined viscoelasticity is, from Eq. 1.12:

σ × ηωi σωi ω0 i + ω
YðωÞ = = = σω 2 (1.13)
σ + ηωi ω0 + ωi ω0 + ω2

where ω0 ≡ σ=η. This combination is called an exponential lead (also called expo-
nential advance, or phase advance). Here, 0° ≤ arg(Y) < 90° (Re(Y) > 0, Im(Y) > 0)
(see Eq. 5.13 for arg function; Sect. 5.2 for Re and Im functions), hence the plot
falls on the first quadrant (I) (Fig. 1.5c). In Eq. 1.13, ω0 is called the (apparent) rate
constant, νc ≡ ω0 =2π is called the characteristic frequency, τ ≡ 1=ω0 is called the
time constant, and σ is called the magnitude (amplitude) of the exponential lead.

1.9 Work and Work Absorption

If a weight is lifted by ΔL, the work performed is defined by ΔW = FΔL, where

F is the gravitational force applied to the weight. This equation is rewritten as
dW = FdL for a infinitesimally small length change dL.
Let us assume that for the sinusoidal length change LðθÞ = L0 + L1 cos θ (where
θ ≡ ωt), tension change FðθÞ = F0 + F1 cosðθ + ϕÞ is observed. The work (ΔW)
performed by the forcing apparatus (length driver) per cycle is calculated as fol-
lows. Because dL = −L1 sin θdθ,
Ð Ðπ
ΔW ≡ 1cycle FdL = − L1 −π fF0 + F1 cosðθ + φÞg sin θdθ
Ðπ (1.14)
= F1 L1 sin φ −π sin θdθ = πF1 L1 sin φ = πL1 ImðY ðωÞÞ
2 2

Note that

F1 sin φ = L1 ImðY ðωÞÞ (1.15)

In calculating work, complex arithmetic cannot be used because work is a

nonlinear concept. Equations 1.14 demonstrates that the work absorption (work
performed by the length driver) is proportionate to the viscous modulus. In the
previous example (exponential lead, Eq. 1.13), because Im(Y(ω)) > 0, work
performed by the length driver is positive in all frequencies, i.e., absorbed by
the muscle preparation.
6 1 Basic Elements of Viscoelasticity

1.10 Oscillatory Work (Work Generator, Exponential Delay)

The complex modulus of an element that generates work (Fig. 1.3b) can be
described as follows:

σω0 σωi ω0 − ωi
YðωÞ = =σ− = σω0 2 (1.16)
ω0 + ωi ω0 + ωi ω0 + ω2

Because Im(Y(ω)) < 0, the work absorption (Eq. 1.15) becomes negative, i.e.,
work is produced by the muscle fiber on the forcing apparatus (length driver).
This element is symbolized as in Fig. 1.3b with the complex modulus of Y(ω) in
Eq. 1.16 and its Nyquist plot in Fig. 1.5d. For ω = 0, Y(0) → σ, which is actually
replaced by process A (Eq. 3.6) in the case of active muscle fibers.

1.11 Plot of the Complex Modulus in the Complex Plane

The complex modulus Y(ν) can be represented on the complex plane (Cartesian
coordinate, Sec. 5.2) with the real axis in the abscissa (called elastic modulus,
Re(Y)), and the imaginary axis in the ordinate (called viscous modulus, Im(Y)).
Here, the frequency (ω or ν) becomes the intervening parameter. In Fig. 1.5,
Eqs. 1.5, 1.6, 1.13, and 1.16 are plotted on the complex plane (c.f., Machin,
1964). This type of plot has been called a “Nyquist plot” in muscle mechanics
since the 1960’s. Figure 1.5a shows general elasticity, Fig. 1.5b general viscos-
ity, Fig. 1.5c an exponential lead (or phase advance), and Fig. 1.5d work gen-
erator and an exponential delay (or phase delay). One should be able to verify
that all frequency points of Fig. 1.5c fall on a semicircle, starting from the
origin (ω = 0) and ending at σ on the abscissa as ω → ∞ (σ is the diameter,
which can be proven by calculating |Y–σ/2|2 from Eq. 1.13. The dynamic modu-
lus is defined as |Y( f )|, and the phase shift as arg[Y( f )], where arg function is
described in Eq. 5.13.

a b c d
ω=∞ ω=∞ ω=0
ω = ω0
σ

ω=
0 ∞ ω=0 σ
ω = ω0
σ ω=0 ω=∞

Fig. 1.5 Nyquist plots of elements. a Elasticity. b Viscosity. c Exponential lead (advance).
d Exponential delay (work generator). See also Machin (1964)
Reference 7

1.12 Exercise

1.1. Based on Eqs. 1.5 and 1.10, verify that the complex modulus of an elastic
element is σ.
1.2. Based on Eqs. 1.6, 1.8, and 1.9, verify that the complex modulus of a viscous
element is ηωi.
1.3. Verify Eqs. 1.11 and 1.12.
1.4. Verify that the plot of Eq. 1.13 on the complex plane (Fig. 1.5c) is a semicircle
in the 1st quadrant with its center on the abscissa.
1.5. Verify that the plot of Eq. 1.16 on the complex plane (Fig. 1.5d) is a semicircle
in the 4th quadrant with its center on the abscissa.

Reference

Machin KE (1964) Feedback theory and its application to biological systems. Symp Soc Exp
Biol 18:421–445
Chapter 2
Reaction Processes (Chemical Kinetics) and
Their Application to Muscle Biology

Abstract This chapter describes basics of cross-bridge models, and how to charac-
terize muscle fiber (and myofibril) preparations in terms of elementary steps of the
cross-bridge cycle. Models with two states, three states, and multi states are exa-
mined. Mathematical derivation to relate observed rate constants to the fundamental
rate constants of the elementary steps are presented. The temperature effect of the
rate constants, their activation energy, and reaction coordinates are discussed.

Keywords Two state model · three state model · multi state model · cross-bridge ·
cross-bridge cycle · muscle fiber · myofibril · elementary steps · rate constant ·
equilibrium constant · activation energy · reaction coordinate · Arrhenius
equation

2.1 Chemical Reactions

Many cross-bridge models have been proposed to account for isometric tension
and its transients, but most of them are difficult to follow owing to their complex-
ity and the numerical approximations that have been used. Quite often, too many
states are employed and too many parameters are used to fit scanty data, lessening
the significance of the model. The purpose of a model is to account for experimen-
tal results, and to predict results for future experiments. In this chapter, I will focus
on simple cases and account for fundamental experimental results. These cases
are limited to the isometric, or near-isometric state of contraction, which I believe
is the best way to characterize the elementary steps of the cross-bridge cycle.
If a shortening of a half sarcomere happens exceeding that of the cross-bridges’
step size (stroke distance), then multiple cross-bridge cycles must follow. In this
condition, transients are limited by the slowest step of the cycle, hence fast steps
cannot be resolved. The symbols used in the text are summarized in the beginning
of this book.

© Springer International Publishing AG 2018 9

M. Kawai, Biomechanics, Muscle Fibers, and How to Interface Experimental
Apparatus to a Computer, https://doi.org/10.1007/978-3-319-72036-4_2
10 2 Reaction Processes (Chemical Kinetics) and Their Application to Muscle Biology

2.2 Two-State Model, Rate Constants, and the First-Order

Reaction

This is the simplest case, hence easiest to understand yet still of value. It is exten-
sively used in classical muscle physiology and mechanics literature (Huxley,
1957; Thorson & White, 1969; Huxley & Simmons, 1971; Abbott, 1972; Brenner,
1988). When a reactant (A) changes to a product (B) with the forward rate con-
stant α and the reversal rate constant α′, this process can be written as in Scheme 1
(Fig. 2.1). For applications to muscle mechanics, state A is recognized as the
attached state with force, and state B as the detached state without force. The rate
of the forward reaction is αA, and the rate of the reversal reaction is α′B, where
italic letters A and B represent concentrations of respective species A and B.
Although the rate and the rate constant are two different quantities, the “rate con-
stant” is often abbreviated as the “rate,” and I will follow this convention when
their difference is apparent. When discussing molecular species, I will use bold
letters such as A and B. Because the forward rate is a loss of A, and the reverse
rate is a gain of A, the rate of the increase of A is described in Eq. 2.1:

dA
= −αA + α′B (2.1)
dt

Fig. 2.1 Scheme 1 (two state a

model) A B
a’
Attached Detached

The reaction in Scheme 1 is called the first-order reaction, because one species
is involved as the reactant (A in Scheme 1). Equation 2.1 is called “master equa-
tion” of the reaction process of Scheme 1. Given that A and B are two forms of
the same molecular species,

A + B = AT (2.2)

where AT is the total concentration and is conserved (i.e., it does not change with
time). For A, B, and AT, “probability” is often used instead of “concentration,” in
which case AT = 1. An elimination of B from Eqs. 2.1 and 2.2 results in:

dA
+ λA = α′AT ; where λ ≡ a + a′ (2.3)
dt

Note that ≡ indicates “definition.” Equation 2.3 can be solved (Sect. 5.4) to
result in:
Aðt Þ = A0 expð−λt Þ + A1 (2.4)
2.3 Three-State Model 11

and

Bðt Þ = − A0 expð−λt Þ + B1 (2.5)

where

α′ α
A1 ≡ AT ; B1 ≡ AT ; (2.6)
λ λ

and A0 = integration constant. Thus, the first-order reaction of Scheme 1 results in

a time course that has one exponential process with the rate constant λ, which is
the sum of the forward rate α and the reverse rate α′ (Eq. 2.3). Because of the
summation, the faster reaction (the larger one of α and α′) determines (or “limits”)
the character of rate λ (consider the case such as α >> α′, then λ ≈ α). λ is called the
“apparent” rate constant, indicating that it is the rate constant deduced from experi-
ments by fitting the time course data to Eq. 2.4; this quantity is also called “observed”
rate constant. In contrast, α and α′ are called the “fundamental” rate constants
(Gutfreund, 1995) of the elementary steps. From experiments one can also deduce A0
and A1, where A0 is determined by the initial condition (degree of perturbation), and
is called the magnitude (or amplitude) of the exponential process.
The average lifetime <t> of an exponential process exp(−λt) is calculated as:
Ð∞
texpð−λtÞdt 1
<t> = Ð0∞ = ≡τ (2.7)
0 expð−λtÞdt
λ

where τ (≡1/λ) is called the time constant.

With time passes (t → ∞, indicating that t is much larger than τ: t >> τ),
Scheme 1 reaches an equilibrium: dA(t)/dt = 0, and A = A1 and B = B1 result.
A1 and B1 are the equilibrium concentrations of species A and B, respectively.
Here the equilibrium constant Kα is defined as:

B1 α
Kα ≡ = (2.8)
A1 α′

This equation is called “mass action law”; the second part of Eq. 2.8 is derived
from Eq. 2.6.

2.3 Three-State Model

To explain delayed tension explicitly, it is necessary to introduce the three-state

model. The three-state model has been frequently used among muscle physiolo-
gists, e.g., (Julian et al., 1974; Thorson & White, 1983; Murase et al., 1986).
Scheme 2 in Fig. 2.2 represents the most general form. This is a cyclic scheme with
12 2 Reaction Processes (Chemical Kinetics) and Their Application to Muscle Biology

Fig. 2.2 Scheme 2 (three- Step 3 (slow)

state model, redrawn from
g
Kawai and Halvorson, 2007)
Attached C A Attached
g’
b’ a’
Step 2 b a Step 1

(phase 3, 2pb) (phase 2, 2pc)

B Detached

the forward reaction consisting of the outer clockwise cycle: A → B → C → A, for

which the respective rates are α, β, and γ. The reversal reaction consists of the inner
counter-clockwise cycle: A → C → B → A, with the respective rates γ′, β′, and α′.
In Scheme 2, we assume that A and C are attached states, which generate and/or
support force, and that B is a detached state, which does not support force. Because
the analysis of Scheme 2 is very complex, we will discuss a simple case here. The
complete analysis of Scheme 2 can be found in (Kawai & Halvorson, 2007).
Now let us consider a simple case (α + α′ ≫ β + β′ ≫ γ + γ′). In the following
discussion, these rate constants are referred to as fast, medium, and slow, respec-
tively. If we focus on a fast time scale comparable to 1/(α + α′) (e.g., ∼1 ms in
rabbit psoas or frog semitendinosus fibers), then steps 2 (B ↔ C) and 3 (C ↔ A) are
too slow to be observed. Therefore, only step 1 can be observed, and the problem
reduces to Scheme 1 (Fig. 2.1), with the apparent rate λ2 = α + α′. Thus, the appar-
ent rate of a fast step (λ2) is the sum of forward and reversal rates of step 1, and it is
not influenced by the rates or equilibrium constants of slower steps 2 and 3.

Fig. 2.3 Scheme 3 Step 1 Step 2

Kα b
A B C
b’

If we focus on a medium time scale, comparable to 1/(β + β′) (e.g., ∼10 ms),
then step 1 is very fast and looks like an equilibrium, whereas step 3 is too slow
to be observed. Therefore, Scheme 2 actually looks like Scheme 3 (Fig. 2.3),
where Kα ≡ α/α′ = B/A. In this case

dðA + BÞ
= − βB + β′C; and A + B + C = AT ; (2.9)
dt

That is, A and B change as a group. By eliminating B and C from above 3

equations, we obtain

dA AT
+ λ3 A = β′ (2.10)
dt 1 + Kα
2.4 Second Order Reaction 13

where

Kα
λ3 ≡ β + β′ (2.11)
1 + Kα

By solving Eq. 2.10 (Sect. 5.4), we arrive at:

Aðt Þ = A0 expð−λ3 t Þ + A1 ;
Bðt Þ = A0 Kα expð−λ3 t Þ + B1 ; (2.12)
C ðt Þ = − A0 ð1 + Kα Þexpð−λ3 t Þ + C1

where A1 ≡ AT / [1 + Kα (1 + Kβ)], B1 = Kα A1 and C1 = KαKβ A1 are the steady-

state concentrations, and Kβ ≡ β/β′ is the equilibrium constant (Kawai &
Halvorson, 2007). Equation 2.12 shows that Scheme 3 has the apparent rate λ3
defined by Eq. 2.11. Thus, the apparent rate of a medium-speed step 2 is a linear
combination (weighted sum) of the fundamental rates β and β′ (Eq. 2.11). It must
be evident here that λ3 is influenced by the equilibrium constant of the fast step 1
(Kα), but λ3 is not influenced by the slow step 3.
In total, the three-state model has 2 exponential processes with 2 apparent rates,
λ2 and λ3; generally, an N-state model has (N-1) exponential processes. Each
apparent rate is, approximately, a linear combination (sum) of the forward and
reversal rates of the same step, and A(t), B(t) and C(t) take the form of exponential
functions. In perturbation analysis, the rates of fast steps 1 and 2 can be observed
(Eqs. 2.4 and 2.12), whereas the rate of the slowest step 3 cannot be observed.
However, because of the presence of in-series compliance, the forward rate con-
stant of step 3 can be deduced (Kawai & Halvorson, 2007; Wang & Kawai, 2013)
and as described in Sect. 4.15. Step 3 is important for calculation of the turnover
rate (Sect. 4.10, Eq. 4.3).

2.4 Second Order Reaction

If an enzyme (A) binds to a substrate (S), then the reaction is A + S → B. This is

called the second-order reaction, because two species are involved as reactants.
Under our experimental conditions, in which there is an infinite supply of S and
its concentration remains constant, the analysis of this situation is exactly the
same as for the Scheme 3 (Fig. 2.3), except that A is replaced with A + S, and Kα
with KαS, where S = [MgATP] (abbreviated as ATP). Note that […] means the
concentration. Equations 2.1 through 2.8 can be used with these replacements,
and from Eq. 2.3 the apparent rate constant becomes: λ = αS + α′. In this case,
the equations that govern the reaction are the same as Eqs. 2.1 and 2.2, with a
modification of α → αS, where α is the second-order rate constant of this reaction.
Kα = α/α′ is now called the association (or binding) constant of MgATP to the
myosin head, and KαS is the corresponding equilibrium constant.
14 2 Reaction Processes (Chemical Kinetics) and Their Application to Muscle Biology

2.5 Application of Scheme 3 to Elementary Steps

of the Cross-Bridge Cycle

The binding of the small ligand to an enzyme (such as the myosin head) is diffu-
sion limited and very fast, which can be better approximated by an equilibrium,
such as shown in step 1 of Fig. 2.4. After the binding, the enzyme (AM) under-
goes a conformational change (step 2), which is slower and its rates can be experi-
mentally observed. This case can be analyzed using Scheme 3, with Kα ≡ K1S,
k2 ≡ β, and k−2 ≡ β′ to result (from Eq. 2.11) in the apparent rate λ3:

K1 S
λ3 ≡ k2 + k−2 (2.13)
1 + K1 S

Fig. 2.4 Scheme 4. A = Step 1 Step 2

actin, M = myosin, and S = k2
K1
ATP AM + S AMS A + MS
k–2

In the case of cross-bridges, the conformational change (step 2) includes loos-

ening of the AM interaction and subsequent detachment of myosin (M) from actin
(A). This formalism was used to explain the ATP effect (Kawai & Halvorson,
1989). When ADP (D) is present, it binds to the substrate site (reversal of step 0
in Fig. 2.5) and prevents ATP binding (step 1) to the site. This is called competi-
tive inhibition by ADP. In this case Scheme 4 is modified to result in Scheme 5.
The apparent rate λ3 of Scheme 5 is:

K1 S
λ3 ≡ k2 + k−2 (2.14)
1 + K0 D + K1 S

where D = [MgADP]. Figure 2.5 and consequent Eq. 2.14 have been used to
explain the ATP and ADP effects (Kawai & Halvorson, 1989). From Eq. 2.8, the
equilibrium constant of step 2 is K2 = k2/k−2.

Fig. 2.5 Scheme 5. Step 0 Step 1 Step 2

Competitive inhibition by D. D K1 k2
K0 is defined as the associa- AMD AM AMS A + MS
tion constant when the reac- K0 k–2
S
tion proceeds to the left,
hence written below the
arrow

The Scheme 3 (Fig. 2.3) was also used to characterize the Pi release step,
with replacements of C → AMDP, B → AM*DP, A → AM*D, Kα ≡ K5P, k4 ≡ β′,
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If we sight(14) naught but seas at dawn?”
“Why, you shall say, at break of day,(15)
Sail on! Sail on! Sail on and on!”

They sailed.(16) They sailed.(17) Then spake(18) the mate:

“This mad sea(19) shows his teeth to-night,
He curls his lip, he lies in wait
With lifted teeth,(20) as if to bite!
Brave Adm’r’l,(21) say but one good word;
What shall we(22) do when hope is gone?”
The words leapt like a leaping sword:(23)
“Sail on! Sail on! Sail on! and on!”

Then pale and worn,(24) he paced his deck

And peered through darkness. Ah, that night(25)
Of all dark nights! And then a(26) speck—
A light!(27) A light? A light! A light!
It grew,(28) a starlit flag unfurled!
It grew to be Time’s burst of dawn.(29)
He gained a world,(30) he gave that world
Its grandest less’n: “On! Sail on!”

A Suggestive Outline for the Study of a Selection

I. Mastery of Main Theme

The first step in the study of any selection is to gain an idea of it as
a whole. This can best be done by reading the selection in its
entirety. If there should be strange words, let them pass for the time
being. Thus we grasp the predominant mood and significant setting
or situation.

II. Progressive Analysis

Read the selection, silently, a second time. The aim now is to
make a mental note of the several parts which make up the whole.
This demands close concentration, in order that we may unify
matters and prevent abrupt transitions. We are to break up the whole
into parts, and each part represents a thought group.
1. Punctuation makes the meaning clear, and the clear meaning
determines the various groups. Example: “It came, rushing in
torrents like an avalanche of rock.” We do not pause after “came,”
although it is so punctuated. Question: Do you find like instances in
the selection under consideration? Where?
2. The length and frequency of the pause which sets off the
groups is dependent upon the context and upon the listeners. If the
context is serious, or if the listeners are uneducated, there will of
necessity be many groups. And obversely, if the context is not
serious or difficult, or if the audience is educated, there will be fewer
and longer groups.
Question: What is the situation in the present selection?
3. In the study of the chief word in the group we must remember
that its real meaning depends upon its relation to the other words in
the same group. For instance, the word “fire” does not mean the
same thing at all times. The meaning of this word depends upon its
kinship with other members of the same group. When we say, “The
house is on fire,” this word “fire” means an altogether different thing
than when we say, “There is a fire in the stove this morning.” Let us
take care that we do not isolate words, but that we get their
associative meanings.
Questions: What are the important words in the various groups?
What is the real meaning of each? Why? Give five synonyms of
each.

III. Reference to Experience

We are now prepared to call upon our storehouse of past
experiences in order that we may identify ourselves more closely
with the author’s meaning. We are to react upon what we read. The
more vividly we can bring what we read from the page into our own
actual experience, the more deeply are we impressed with its
meaning. We translate the unseen, the unfelt and unbelieved by
likening it to what is already seen, felt or believed. If experience is
lacking, we draw upon our imagination.

1. If we are reading a description, we will see this scene in

terms of a past like experience.
2. If we are reading a narration, we will feel it in terms of a
past like experience.
3. If we are reading something we have not believed, we will
accept it in terms of what we have already believed.

Question: What experiences does this selection call upon from

me? What purposes do they serve?

IV. Classification
There are three divisions into which all selections may be put. A
selection may be written to make something Clear; it may be for the
purpose of inspiring, or elevating one’s thoughts and feelings—to
make Impressive; it may be for the purpose of enforcing some great
truth—to make Belief. This classification is based upon the author’s
purpose.
Questions: Where the author’s purpose is to make Clear some
obscure point or idea.

1. What significant words are used?

2. Is there any obscurity? Why?
3. What illustrations or comparisons are made?
4. Think earnestly of an experience which will aid you to see
clearly the author’s purpose.

Where the author’s purpose is Impressiveness

 1. Is the emotion aroused pleasurable?
2. Have you had an experience which resembles what is
referred to?
3. What mood is predominant? Is it:

Impassioned, grave, sad,

Triumphant, exalted, solemn,
Humorous, satirical, pathetic,
Inspiring, enheartening, discouraging?

4. What are the minor moods? Supply your own descriptive

mood if none of the following are adequate:

Fanciful, enthusiastic, cheerful,

Dreamy, sentimental, witty,
Pensive, tender, serene, quiet,—or suggestive of
Awe, loneliness,
Admiration, suspense, joy, anger,
Fear, rage, sympathy, grief, sorrow, surprise, anxiety.

Where the author’s purpose is Belief. The author does more than
make us see, or feel.

1. What actual experience have you had that resembles the

thing the author would have you believe?
2. Do you accept as truth what you have read?
3. What particular thought carries the most conviction?
4. Do you think others should believe what the author says?
5. Is it clear and impressive, and do you believe it?
 V. Setting
This has to do with time, place, objects, sounds, movement, or
anything that gives local color to the selection.
Questions:

1. Is it modern or old?
2. Where is the scene laid?
3. Are descriptions given in detail or mere suggestion?
4. Is dialect used?
5. Will personation aid in rendering the selection?
6. Does the power and beauty of the selection lie in narration,
description, or in character drawing?
7. Name some definite things, sounds or objects described,
that give color or atmosphere.
8. Is the movement:

Slow, swift, light, heavy,

Tripping, graceful, spirited,
Powerful, easy, varied?

VI. Vocalization
Let our guide be as Shakespeare has so well put it:
Let your own discretion be your tutor. Suit the action to the word;
the word to the action; with this special observance—that you
o’erstep not the modesty of nature.

1. Read the selection as ordinary conversation.

2. Now read again as enlarged conversation, or, as it were,
for the ears of many.
 3. Ask yourself the following questions:

(1) Do I make proper use of Pitch?

(2) Do I make proper use of Pause?
(3) Do I make proper use of Inflection?
(4) Do I make proper use of Tone Color?
(5) Do I make proper use of Stress?
(6) Do I make proper use of Movement?

4. At all times let us remember that our purpose is not to give

a pleasing performance, but faithfully to interpret the
author’s meaning.

A Condensed Outline for the Study of a Selection for

Oral Presentation

I. Intelligent Impression

A. General Preparation
Read silently the entire selection. The purpose is to gain an
impression of the selection as a whole.

1. What was the author’s purpose in writing this selection?

2. What specific intent did he have:

a. To make something clear?

b. To make something impressive?
c. To establish a truth?
d. To stimulate to righteous action?

3. Consult the dictionary for the meaning of strange words.

 4. Look up the historical references.

B. Special Preparation
Read the selection silently a second time. The aim is to make a
mental note of the respective importance of the several parts which
make up the whole.

1. What is the definite idea, or definite picture, or definite

feeling the author would have us get?
2. In what part of the selection is the author’s aim most
forcibly presented?
3. What is the relative value of the thought-groups?

II. Intelligible Expression

A. General Preparation
Before rendering a selection orally it must be given a setting. This
has to do with time, place, objects, sounds, movements, or anything
that tends to give local color.

1. Is the selection colloquial or dramatic?

2. Is dialect used?
3. Will personation be necessary?
4. Give an original word-picture of the characters and
situation.
5. To what reference to experience does it make?
6. What is the predominant mood?

B. Special Preparation
 Read the selection aloud for the first time. In doing this, ask
yourself:

1. Am I reading with correct thought-groups?

2. Do I make proper use of the pause? (Remember the length
and frequency of the pause depends upon the nature of
the subject and the audience.)
3. Am I enunciating clearly?
4. Is my voice melodious? That is, do I make proper use of
pitch and inflection?
5. Am I conscious of the change and interchange of moods?
6. Do I make proper use of stress and movement?
7. Do my tones fit the color-words?
8. Am I faithfully and adequately interpreting the author’s
meaning?
 PROSE SELECTIONS
Humorous
Pathetic
Dramatic
Dialect

THE JOY OF READING

Who can estimate the joy, comfort and inspiration reading has
afforded to the human race, how many weary hours it has solaced,
how many distracted minds it has quieted, how many harassed souls
it has soothed into forgetfulness? Who has not felt the thrill of
discovery when he has found a new author, a new poet who
peculiarly affected his mind, his soul, his risibilities, his ambitions, his
life? I shall never forget when I found Charles Warren Stoddard’s
“Apostrophe to a Skylark.” It was buried in one of his books and few
seemed ever to have read it. There was joy incalculable in putting it
side by side with Shelley’s classic “Ode” and comparing the two
conceptions. Thousands of souls have been inspired by reading to
higher, nobler, more worthy endeavor. So, like Sancho Panza, we
bless God and thank Him for the man who invented reading.
—George Wharton James.
 HUMOROUS SELECTIONS

NATHAN FOSTER
By Paul L. Dunbar
Nathan Foster and his lifelong friend and neighbor, Silas
Bollender, sat together side by side upon the line-fence that
separated their respective domains. They were both whittling away
industriously, and there had been a long silence between them.
Nathan broke it, saying:
“’Pears to me like I’ve had oncommon good luck this year.”
“Wall, you have had good luck, there ain’t no denyin’ that. It ’pears
as though you’ve been ee-specially blest.”
“An’ I know I ain’t done nothin’ to deserve it.”
“No, o’ course not. Don’t take no credit to yourself, Nathan. We
don’t none of us deserve our blessings, however we may feel about
our crosses; we kin be purty shore o’ that.”
“Now, look, my pertater vines was like little trees, an’ nary a bug
on ’em.”
“An’ you had as good a crop of corn as I’ve ever seen raised in
this part of Montgomery county.”
“Yes, an’ I sold it, too, jest before that big drop in the price.”
“After givin’ away all yer turnips you could, you had to feed ’em to
the hogs.”
“My fruit trees jest had to be propped up, an’ I’ve got enough
perserves in my cellar to last two or three winters, even takin’ into
consideration the drain o’ church socials an’ o’ charity.”
“Yore chickens are fat and sassy, not a sign o’ pip on ’em.”
“Look at them cows in the fur pastur. Did yer ever see anything to
beat ’em fer sleekness?”
 “Wall, look at the pasture itself; it’s most enough to make human
beings envy the critters. You didn’t have a drop of rain on yer while
you was gettin’ in yer hay, did yer?”
“Not a drop.”
“An’ I had a whole lot ruined jest as I was about to rick it.”
“Silas, sich luck as I’m a-havin’ is achilly skeery; it don’t seem
right.”
“No, it don’t seem right for a religious man like you, Nathan. Ef you
was a hard an’ graspin’ Sinner, it ’ud be jest makin’ you top-heavy
so’s yore fall ’ud be the greater.”
“I don’t know but what that’s it, anyhow. Mebbe I’m a-gettin’ puffed
up over my goods without exactly knowin’ it.”
“Mebbe so, mebbe so. Them kind o’ feelin’s is mighty sneakin’
comin’ on a body. O’ course I ain’t seen no signs of it in you; but it
’pears to me you’ll have to mortify yore flesh yit to keep from being
purse-proud.”
“Mortify the flesh?”
“O’ course, you can’t put peas in yore shoes er get any of yer
frien’s to lash you, so you’ll have to find some other way of mortifyin’
yer flesh. Wall, fer my part, I don’t need to look fur none, fur I never
had too many blessin’s in my life, less’n you’d want to put the
children under that head.”
Silas shut his jack-knife with a snap and, laughing, slid down on
his side of the fence. In serious silence Nathan Foster watched him
go stumping up the path toward the house.
“Silas seems to take everything so light in this world; I wonder how
he can do it.”
With Nathan, now, it was just the other way. Throughout his eight
and forty years he had taken every fact of life with ponderous
seriousness. Entirely devoid of humor, he was a firm believer in
signs, omens, tokens, and judgments. He was a religious man, and
his wealth frightened and oppressed him. He gave to his church and
gave freely.
As usual, he had taken his friend’s bantering words in hard
earnest and was turning them over in his mind.
The next morning when Nathan and Silas met to compare notes,
Nathan began:
“I have been thinking over what you said last night, Silas, about
me mortifyin’ my flesh, and it seems to me like a good idee. I
wrasselled in prayer last night, and it was shown to me that it wa’n’t
no more’n right fur me to make some kind o’ sacrifice fur the mercies
that’s been bestowed upon me.”
“Wall, I don’t know, Nathan; burnt-offerings are a little out now.”
“I don’t mean nothin’ like that; I mean some sacrifice of myself,
some—”
His sentence was broken in upon by a shrill voice that called from
Silas Bollender’s kitchen door:
“Si, you’d better be a-gittin’ about yore work instid o’ standin’ over
there a-gassin’ all the mornin’. I’m shore I don’t have no time to
stand around.”
“All right, Mollie; speakin’ of mortifyin’ the flesh an’ makin’ a
sacrifice of yoreself, Nathan, why don’t you git married?”
Nathan started.
“Then you’d be shore to accomplish both. Fur pure mortification of
the flesh, I don’t know of nothin’ more thoroughgoin’ er effectiver
than a wife. Also she is a vexation to a man’s sperit. You raaly ought
to git married, Nathan.”
“Do you think so?”
“It looks to me that that ’ud be about as good a sacrifice as you
could make; an’ then it’s such a lastin’ one.”
“I don’t believe you realize what you air a-sayin’, Silas. It’s a
mighty desprit step that you’re advisin’ me to take.”
 Again Mrs. Bollender’s voice broke in:
“Si, air you goin’ to git anything done this mornin’, er air you goin’
to stand there an’ hold up that fence fur the rest of the day?”
“Nathan, kin you stand here an’ listen to a voice an’ a speech like
that an’ then ask me if I realize the despritness of marriage?”
“It’s desprit, but who’d you advise me to marry,—Silas, that is, if I
made up my mind to marry,—an’ I don’t jest see any other way.”
“Oh, I ain’t pickin’ out wives fur anybody, but it seems to me that
you might be doin’ a good turn by marryin’ the Widder Young. The
Lord ’ud have two special reasons fur blessin’ you then; fur you’d be
mortifyin’ yore flesh an’ at the same time a-helpin’ the widder an’ her
orphans.”
“That’s so.” He couldn’t admit to Silas that he had been thinking
hard of the Widow Young even before he had of mortifying his flesh
with a wife.
Once decided, it did not take him long to put his plans into
execution. But he called Silas over to the fence that evening after he
had dressed to pay a visit to the widow.
“Wall, Silas, I’ve determined to take the step you advised.”
“Humph, you made your mind up in a hurry, Nathan.”
“I don’t know as it’s any use a-waiting; ef a thing’s to be done, I
think it ought to be done and got through with. What I want particular
to know now is, whether it wouldn’t be best to tell Lizzie—I mean the
widder—that I want her as a means of mortification.”
“Wall, no, Nathan, I don’t know as I would do that jest yit; I don’t
believe it would be best.”
“But if she don’t know, wouldn’t it be obtainin’ her under false
pretenses if she said yes?”
“Not exactaly the way I look at it, fur you’ve got more motives fur
marryin’ than one.”
“What! Explain yoreself, Silas; explain yoreself.”
 “I mean you want to do her good as well as subdue your own
sperit.”
“Oh, yes, that’s so.”
“Now, no woman wants to know at first that she’s a vexation to a
man’s sperit. It sounds scriptual, but it don’t sound nooptial. Now
look at me an’ Mis’ Bollender. I never told her until we’d been
married more’n six months; but she didn’t believe it then, an’ she
won’t believe it till this day.”
“Wall, I’ll agree not to tell her right away, but if she consents, I
must tell her a week or so after we are married. It’ll ease my
conscience. Ef I could tell her now, it ’ud be a heap easier in gittin’
round the question. I don’t know jest how to do it without.”
“Oh, you won’t have no trouble in makin’ her understand.
Matrimony’s a subject that women air mighty keen on. They can see
if a man’s a-poppin’ the question ef he only half tries. You’ll git
through all right.”
Somewhat strengthened, Nathan left his friend and sought the
widow’s home. He found her stitching away merrily under the light of
a coal-oil lamp with a red shade.
“La, Nathan, who’d a’ expected to see you up here? You’ve got to
be such a home body that no one don’t look to see you out of yore
own field and garden.”
“I jest thought I’d drop in.”
“Wall, it’s precious kind of you, I’m shore. I was a-feelin’ kind o’
lonesome. The children go to bed with the chickens.”
“I jest thought I’d drop in.”
“Wall, it does remind me of old times to see you jest droppin’ in,
informal like, this way. My, how time does fly!”
“Widder, I’ve been thinkin’ a good deal lately; I’ve been greatly
prospered in my day; in fact, my cup runneth over.”
“You have been prospered, Nathan.”
 “Seems ’s ef—seems ’s ef I ought to sheer it with somebody, don’t
it?”
“Wall, Nathan, I don’t know nobody that’s more generous in givin’
to the pore than you air.”
“I don’t mean in jest exactly that way. I mean, widder—you’re the
morti—I mean the salvation of my soul. Could you—would you—er
do you think you’d keer to sheer my blessin’s with me an’ add
another one to ’em?”
The Widow Young looked at him in astonishment; then the tears
filled her eyes as she asked, “Nathan, do you mean it?”
“I wouldn’t a-spent so much trouble on a joke, widder.”
“No, it don’t seem that you would, Nathan. Well, it’s mighty
sudden, mighty sudden, but I can’t say no.”
“Fur these an’ many other blessin’s make us truly thankful, O
Lord,” said Nathan devoutly. And he sat another hour with the widow
making plans for the early marriage, on which he insisted.
The widow had been settled in Nathan’s home over a month
before he had ever thought of telling her of the real motive of his
marriage, and every day from the time it occurred to him it grew
harder for him to do it.
One night when he had been particularly troubled he sought his
friend and counselor with a clouded brow. They sat together in their
accustomed place on the fence.
“I’m bothered, Silas.”
“What’s the matter?”
“Why, there’s several things. First off, I ain’t never told the widder
that she was a mortification, an’ next she ain’t. I look around at that
old house o’ mine that ain’t been a home since mother used to scour
the hearth, an’ it makes me feel like singin’ fer joy. An’ I hear them
children playin’ round me—they’re the beatenest children; that
youngest one called me daddy yistiddy—well, I see ’em playin’ round
and my eyes air opened, an’ I see that the widder’s jest another
blessin’ added to the rest. It looks to me like I had tried to beat the
Almighty.”
“Wall, now, Nathan, I don’t know that you’ve got any cause to feel
bothered. You’ve done yore duty. If you’ve tried to mortify yore flesh
an’ it refused to mortify, why, that’s all you could do, an’ I believe the
Lord’ll take the will fer the deed an’ credit you accordin’ly.”
“Mebbe so, Silas, mebbe so.”
—Copyright by Dodd, Mead & Co., New York, and used by
arrangement.

DOING A WOMAN’S WORK

By McKillip-Stanwood
“Breakfast ready yet?” asked Jack Telfer, as he set two pails of
foaming milk on the bench and turned to wash his hands.
“Almost,” replied his wife. “But, say, Jack, won’t you fix the calf pen
while you’re waiting? It won’t take but a minute. The calves got out
twice yesterday and tramped all over the flower beds and garden. I
had an awful time getting them in. I tried to fix it, but I don’t think I did
a good job.”
“I can’t stop now. I guess it’s all right. If they get out, why chase
them in; you have nothing else to do, and I’ll fix it up right when I get
time. I want my breakfast now. I can’t fool around here till noon. I’ve
got to cultivate the peaches to-day.”
“I’ve nothing else to do,” repeated his wife, as she dished up the
tempting breakfast. “Well, I like that, Jack Telfer. I wish to goodness I
hadn’t any more to do than you have.”
“Why, what under the sun have you to do? You have only Toodles
and me to look after and this little house to keep. I could do all the
work you do with one hand tied behind me and then find time to
throw at the birds. You see, I know what I am talking about, for I can
cook and do housework as well as any woman.”
 “You’ve never displayed any talent in that direction since I’ve
known you. It’s like pulling teeth to get you to do a chore around the
house. Not that I want a man to do housework, for I don’t; that’s a
woman’s business. But when she has every step to take and a
dozen things to do at once, a little help occasionally comes mighty
handy.”
“Well, the reason I don’t help around here is because there isn’t
much to do. The work you have is a snap, my girl, and a mighty soft
one, too. Why, my mother had nine children and did all her own work
and cooked for harvest hands and threshers and used to help the
neighbors out if they got in a pinch.”
“Well, my dear husband, I do not doubt but your mother was a
very smart woman. She must have been to have raised so promising
a son. But women are not all alike, my dear.”
“Now, your work is a sort of paper-flower work compared with what
I have to do. It would be a picnic for me to stay in the house, wash
dishes, play with the baby and do such things.”
“All right, suppose you have a picnic to-day. I can drive the
cultivator just as well as you and you can cook and keep house a
great deal better than I; at least you think you can. I’ll hitch up and
cultivate the peaches and you can tie one hand behind you and do
the work to-day and see how much time you have to throw at the
birds. What do you say?”
“Say,” laughed Mr. Telfer as he pushed back from the table. “Why, I
say I’m willing, but if you don’t get enough riding in the hot sun—”
“The hot sun,” interrupted his wife, “is no worse than the hot stove
I cook over. Will you do it?”
“You bet I’ll do it, but you must tell me what’s to be done so you
can’t throw it up to me for ever after that the reason I got through so
soon was because I didn’t do half the work.”
“First,” said Mrs. Telfer, “there’s the milk to skim and the calves to
feed and the churning to do. Skim the milk on the north shelf in the
cellar; the dishes to wash, and don’t forget to scald the churn and the
milk things. Then you can iron; the clothes are all dampened down in
the basket. You need not iron any but the plain things, I’ll do the
others. Pit the cherries I picked last night and make a pie for dinner.
And, oh, yes, you will have to kill a chicken and dress it, for you
know you said last night you wanted chicken and dumplings for
dinner to-day, and now is your chance.
“Stew some prunes for supper to-night, make the bed, sweep and
dust and get the vegetables ready for dinner. Oh, I guess you know
about what there is to do. I must be off now, for it is nearly 6 o’clock.”
And she was gone.
“Well, it’s early yet; guess I’ll smoke and read the Rural World
awhile. There’s an article on hogs I wanted to read; it seems nice to
have time to do what you please.”
After he had read a long time he at last knocked the ashes into his
hand and stretched lazily.
He went down cellar and skimmed the milk, then he fed the
calves, laughed at the way his wife had tried to fix the calf pen, went
in and took off the table cloth and piled the dishes and empty milk
things on the table.
“Guess I’ll wash up before I churn. No, I won’t, either. I’ll churn
first; then I’ll clean up all at once. Oh, I’ve got a head on me. I ought
to have been a woman.”
He brought from the cellar a large new pan of thick cream and set
it on the table, then he went to scald the churn, but the fire was out
and the dish water Mrs. Telfer had put on before breakfast was
nearly cold.
“Blame it all, I’ve got to go to the barn for peach pits; not one in the
basket. But I’ll kill the chicken while I’m out there and save an extra
trip. If Jennie would only use some management about her work
she’d have plenty of time.”
The large pit basket was soon filled, but the chicken was another
proposition. Every time he selected one to catch it seemed to know it
was a marked bird and would shy off to the edge of the flock. At last
he had to run one down, and he wrung its neck with a great deal of
satisfaction. As he entered the house the clock struck nine.
 “Wheu! Where has the morning gone? I must get a move on me.
Guess I’ll make the pie first so it can bake while the water is
heating.”
He prepared the cherries. Then he made the pie; made it as well
as a woman could. He had pushed the dishes back on the cluttered
table to make room for his bread-board, and just as he had the crust
nicely stamped down around the edge of his pie, with a fork, a
tousled head of yellow curls appeared in the doorway, one chubby
hand holding up a long, white nighty, the other rubbing a sleepy eye.
There was surprise on the baby face at the sight of his father.
Papa meant fun for Toodles, and, running to him, he put up his little
arms, saying, “Papa, high me; high Toodles, papa; high Toodles.”
And his father, dusting the flour from his hands, tossed the baby to
the ceiling again and again while the little fellow screamed with
delight.
In the midst of this jolly frolic the clock announced that it was the
tenth hour of the day.
“Hear that, young man?” said the father. “That means that we must
cut out this racket and get down to business. Your paternal ancestor
is chief cook and general manager to-day and has several little
chores to do yet. We will get Toodles’ breakfast first, then wash and
dress him afterwards so that he won’t get mussed up when he eats.
“Mamma don’t do that way, but we can give mamma a few
pointers on keeping a baby clean, can’t we, Toodles?”
And, putting the child in his high-chair, Mr. Telfer pinned a tea
towel around the little neck for a bib, took a bowl and went to the
cellar for some new milk.
While Toodles was eating breakfast his father washed the prunes
and put them on to stew, set the pie in the oven and started to build
the fire, but he was interrupted by an emphatic voice saying, “Papa,
down; papa, down.”
“All right, young man, I’ll attend to your case directly,” said Jack,
touching a match to the kindling. “Guess I’ll wash and dress you and
have you off my hands.”
 And, taking a wash-pan of tepid water, with soap, comb, rag, towel
and Toodles, he went into the sitting-room where it was cool and
pleasant. The baby’s clean clothes were lying upon a chair, where
his mamma had placed them the night before. Then what a time they
had. Toodles would catch the wash rag in his teeth and papa would
shake it and growl till the little mouth would have to let loose to
scream with the agonizing fun.
Then came the tangled curls, and it took a wonderful story about a
doggie that would say “Bow, wow,” and a little horsie that Toodles
could ride and a chicky that went “Peep, peep, peep,” and several
other mental concoctions to keep the baby quiet until the ringlets
were in order.
When the clean coaties were on and two little arms hugged papa
tight, Jack Telfer thought, “Jennie calls this work.”
The clock pounded out eleven strokes.
“Blast that clock; what’s got into it,” thought the man, putting the
child down and hurrying to the kitchen. “I’ve been busy every minute
this morning, and here it is 11 o’clock and not a thing done yet.”
He found the fire had burned out; he had forgotten to put the
peach pits on the kindling when he had stopped to fuss with Toodles.
“Well, I guess I’ll make it all right by noon,” he soliloquized. “This is
a hurry-up order, but I’ll be on time or eat my hat.”
He looked at his pie; it was nearly half baked. He built a roaring
fire, packed the stove with peach pits, pulled the prunes to the front
where they would cook quicker, and was debating in his mind which
he should scald first, the churn or the chicken, when something
rushed by the door.
“Drat those calves; they’re out again.”
Snatching his hat, he hurried after them. It was a merry chase for
the calves if not for Mr. Telfer. They were willing to go in any direction
but the right one, and by the time he got them corralled Jack was
hot, tired and cross.