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About the Authors
James Stewart received the M.S. degree from Troy Day received the M.S. degree in biology
Stanford University and the Ph.D. from the University from the University of British Columbia and the Ph.D.
of Toronto. After two years as a postdoctoral fellow in mathematics from Queen’s University. His first
at the University of London, he became Professor of academic position was at the University of Toronto,
Mathematics at McMaster University. His research before being recruited back to Queen’s University as
has been in harmonic analysis and functional analy- a Canada Research Chair in Mathematical Biology.
sis. Stewart’s books include a series of high-school He is currently Professor of Mathematics and Sta-
textbooks as well as a best-selling series of calculus tistics and Professor of Biology. His research group
textbooks published by Cengage Learning. He is also works in areas ranging from applied mathematics
coauthor, with Lothar Redlin and Saleem Watson, of a to experimental biology. Day is also coauthor of the
series of college algebra and precalculus textbooks. widely used book A Biologist’s Guide to Mathematical
Translations of his books include those into Spanish, Modeling, published by Princeton University Press in
Portuguese, French, Italian, Korean, Chinese, Greek, 2007.
Indonesian, and Japanese.
A talented violinist, Stewart was concertmaster of
the McMaster Symphony Orchestra for many years
and played professionally in the Hamilton Philhar-
monic Orchestra. He has given more than 20 talks
worldwide on Mathematics and Music.
Stewart was named a Fellow of the Fields Institute
in 2002 and was awarded an honorary D.Sc. in 2003
by McMaster University. The library of the Fields
Institute is named after him. The James Stewart
Mathematics Centre was opened in October, 2003, at
McMaster University.
Copyright 2016 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content may be suppressed from the eBook and/or eChapter(s).
Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it.
Contents
Preface xv
To the Student xxv
Calculators, Computers, and Other Graphing Devices xxvi
Diagnostic Tests xxviii
Prologue: Mathematics and Biology xxxiii
Case Studies in Mathematical Modeling xli
Case Study 1 Kill Curves and Antibiotic Effectiveness xlii
Case Study 2 Hosts, Parasites, and Time-Travel xlvi
1 Functions and Sequences 1

1.1 Four Ways to Represent a Function 2
■ Representations of Functions ■ Piecewise Defined Functions ■ Symmetry
■ Periodic Functions ■ Increasing and Decreasing Functions
1.2 A Catalog of Essential Functions 17

■ Linear Models ■ Polynomials ■ Power Functions ■ Rational Functions
■ Algebraic Functions ■ Trigonometric Functions ■ Exponential Functions
■ Logarithmic Functions
1.3 New Functions from Old Functions 31

■ Transformations of Functions ■ Combinations of Functions
Project The Biomechanics of Human Movement 40
1.4 Exponential Functions 41

■ The Growth of Malarial Parasites ■ Exponential Functions ■ Exponential Growth
■ HIV Density and Exponential Decay ■ The Number e
1.5 Logarithms; Semilog and Log-Log Plots 52

Inverse Functions ■ Logarithmic Functions
■ ■ Natural Logarithms
Graph and Growth of the Natural Logarithm
■ ■ Semilog Plots ■ Log-Log Plots
Project The Coding Function of DNA 69
1.6 Sequences and Difference Equations 70

Recursive Sequences: Difference Equations
■
Discrete-Time Models in the Life Sciences

■
Project Drug Resistance in Malaria 78
Review 80
Case Study 1a Kill Curves and Antibiotic Effectiveness 84
vii
viii CONTENTS
2 Limits 89
2.1 Limits of Sequences 90
■The Long-Term Behavior of a Sequence ■ Definition of a Limit ■ Limit Laws
■ Geometric Sequences ■ Recursion for Medication ■ Geometric Series
■ The Logistic Sequence in the Long Run
Project Modeling the Dynamics of Viral Infections 101
2.2 Limits of Functions at Infinity 102

■ The Monod Growth Function ■ Definition of a Limit at Infinity
■ Limits Involving Exponential Functions ■ Infinite Limits at Infinity
2.3 Limits of Functions at Finite Numbers 111

■ Velocity Is a Limit ■ Limits: Numerical and Graphical Methods
■ One-Sided Limits ■ Infinite Limits
2.4 Limits: Algebraic Methods 125

■ The Limit Laws ■ Additional Properties of Limits
■ Limits of Trigonometric Functions
2.5 Continuity 137

■ Definition of a Continuous Function ■ Which Functions Are Continuous?
■ Approximating Discontinuous Functions by Continuous Ones
Review 149
Case Study 2a Hosts, Parasites, and Time-Travel 151
3 Derivatives 155
3.1 Derivatives and Rates of Change 156
■ Measuring the Rate of Increase of Blood Alcohol Concentration ■ Tangent Lines
■ Derivatives ■ Rates of Change
3.2 The Derivative as a Function 168

■Graphing a Derivative from a Function’s Graph ■ Finding a Derivative from a
Function’s Formula ■ Differentiability ■ Higher Derivatives ■ What a Derivative
Tells Us about a Function
3.3 Basic Differentiation Formulas 181

■ Power Functions ■ New Derivatives from Old ■ Exponential Functions
■ Sine and Cosine Functions
3.4 The Product and Quotient Rules 194

■ The Product Rule ■ The Quotient Rule ■ Trigonometric Functions
3.5 The Chain Rule 202

■ Combining the Chain Rule with Other Rules ■ Exponential Functions with
Arbitrary Bases ■ Longer Chains ■ Implicit Differentiation ■ Related Rates

■ How To Prove the Chain Rule
Contents ix
3.6 Exponential Growth and Decay 215

■ Population Growth ■ Radioactive Decay ■ Newton’s Law of Cooling
Project: Controlling Red Blood Cell Loss During Surgery 222
3.7 Derivatives of the Logarithmic and Inverse Tangent Functions 222

■ Differentiating Logarithmic Functions ■ Logarithmic Differentiation
■ The Number e as a Limit ■ Differentiating the Inverse Tangent Function
3.8 Linear Approximations and Taylor Polynomials 230

■ Tangent Line Approximations ■ Newton’s Method ■ Taylor Polynomials
Project: Harvesting Renewable Resources 239
Review 240
Case Study 1b Kill Curves and Antibiotic Effectiveness 245
4 Applications of Derivatives 249

4.1 Maximum and Minimum Values 250
■Absolute and Local Extreme Values ■ Fermat’s Theorem
■ The Closed Interval Method
Project: The Calculus of Rainbows 259
4.2 How Derivatives Affect the Shape of a Graph 261

■ The Mean Value Theorem ■ Increasing and Decreasing Functions ■ Concavity
■ Graphing with Technology
4.3 L’Hospital’s Rule: Comparing Rates of Growth 274

Indeterminate Quotients ■ Which Functions Grow Fastest?
■
Indeterminate Products ■ Indeterminate Differences

■
Project: Mutation-Selection Balance in Genetic Diseases 284
4.4 Optimization Problems 285

Project: Flapping and Gliding 297
Project: The Tragedy of the Commons: An Introduction to Game Theory 298
4.5 Recursions: Equilibria and Stability 299

■ Equilibria ■ Cobwebbing ■ Stability Criterion
4.6 Antiderivatives 306

Review 312
5 Integrals 315
5.1 Areas, Distances, and Pathogenesis 316
■ The Area Problem ■ The Distance Problem ■ Pathogenesis
5.2 The Definite Integral 329

■ Calculating Integrals ■ The Midpoint Rule ■ Properties of the Definite Integral
x CONTENTS
5.3 The Fundamental Theorem of Calculus 342

Evaluating Definite Integrals ■ Indefinite Integrals ■ The Net Change Theorem
■
The Fundamental Theorem ■ Differentiation and Integration as Inverse Processes

■
Project: The Outbreak Size of an Infectious Disease 354
5.4 The Substitution Rule 354

■ Substitution in Indefinite Integrals ■ Substitution in Definite Integrals ■ Symmetry
5.5 Integration by Parts 362

■ Indefinite Integrals ■ Definite Integrals
5.6 Partial Fractions 368

5.7 Integration Using Tables and Computer Algebra Systems 371
■ Tables of Integrals ■ Computer Algebra Systems
■ Can We Integrate All Continuous Functions?
5.8 Improper Integrals 376

Review 381
Case Study 1c Kill Curves and Antibiotic Effectiveness 385
6 Applications of Integrals 387

6.1 Areas Between Curves 388
Cerebral Blood Flow
■
Project: Disease Progression and Immunity 394

Project: The Gini Index 395
6.2 Average Values 397

6.3 Further Applications to Biology 400
■ Survival and Renewal ■ Blood Flow ■ Cardiac Output
6.4 Volumes 405

Review 412
Case Study 1d Kill Curves and Antibiotic Effectiveness 414
Case Study 2b Hosts, Parasites, and Time-Travel 416
7 Differential Equations 419

7.1 Modeling with Differential Equations 420
■Models of Population Growth
■ Classifying Differential Equations
Project: Chaotic Blowflies and the Dynamics of Populations 430
Contents xi
7.2 Phase Plots, Equilibria, and Stability 431

Phase Plots ■ Equilibria and Stability
■
A Mathematical Derivation of the Local Stability Criterion

■
Project: Catastrophic Population Collapse:

An Introduction to Bifurcation Theory 438
7.3 Direction Fields and Euler’s Method 440

■ Direction Fields ■ Euler’s Method
7.4 Separable Equations 449

Project: Why Does Urea Concentration Rebound after Dialysis? 458
7.5 Systems of Differential Equations 459

Parametric Curves ■ Systems of Two Autonomous Differential Equations
■
Project: The Flight Path of Hunting Raptors 467
7.6 Phase Plane Analysis 468

■ Equilibria ■ Qualitative Dynamics in the Phase Plane
Project: Determining the Critical Vaccination Coverage 479
Review 480
Case Study 2c Hosts, Parasites, and Time-Travel 484
8 Vectors and Matrix Models 487

8.1 Coordinate Systems 488
■ Three-Dimensional Space ■ Higher-Dimensional Space
8.2 Vectors 496

■ Combining Vectors ■ Components
8.3 The Dot Product 505

Projections
■
Project: Microarray Analysis of Genome Expression 513

Project: Vaccine Escape 514
8.4 Matrix Algebra 514

■ Matrix Notation ■ Matrix Addition and Scalar Multiplication ■ Matrix Multiplication
8.5 Matrices and the Dynamics of Vectors 520

■ Systems of Difference Equations: Matrix Models ■ Leslie Matrices ■ Summary
8.6 The Inverse and Determinant of a Matrix 528

The Inverse of a Matrix ■ The Determinant of a Matrix
■
Solving Systems of Linear Equations

■
Project: Cubic Splines 536
8.7 Eigenvectors and Eigenvalues 537

■ Characterizing How Matrix Multiplication Changes Vectors
■ Eigenvectors and Eigenvalues
xii CONTENTS
8.8 Iterated Matrix Models 547

Solving Matrix Models ■ Solutions with Complex Eigenvalues
■
Perron-Frobenius Theory
■
Project: The Emergence of Geometric Order in Proliferating Cells 559
Review 560
9 Multivariable Calculus 565

9.1 Functions of Several Variables 566
■ Functions of Two Variables ■ Graphs ■ Level Curves
■ Functions of Three Variables ■ Limits and Continuity
9.2 Partial Derivatives 585

■ Interpretations of Partial Derivatives ■ Functions of More Than Two Variables
■ Higher Derivatives ■ Partial Differential Equations
9.3 Tangent Planes and Linear Approximations 596

■ Tangent Planes ■ Linear Approximations
Project: The Speedo LZR Racer 603
9.4 The Chain Rule 604

■ Implicit Differentiation
9.5 Directional Derivatives and the Gradient Vector 610

■ Directional Derivatives ■ The Gradient Vector
■ Maximizing the Directional Derivative
9.6 Maximum and Minimum Values 619

■ Absolute Maximum and Minimum Values
Review 628
10 Systems of Linear Differential Equations 631

10.1 Qualitative Analysis of Linear Systems 632
■ Terminology ■ Saddles ■ Nodes ■ Spirals
10.2 Solving Systems of Linear Differential Equations 640

■ The General Solution ■ Nullclines versus Eigenvectors ■ Saddles
■ Nodes ■ Spirals ■ Long-Term Behavior
10.3 Applications 652

Metapopulations ■ Natural Killer Cells and Immunity ■ Gene Regulation
■
Transport of Environmental Pollutants

■
Project: Pharmacokinetics of Antimicrobial Dosing 664
10.4 Systems of Nonlinear Differential Equations 665

■ Linear and Nonlinear Differential Equations ■ Local Stability Analyses
■ Linearization ■ Examples
Review 676
Case Study 2d: Hosts, Parasites, and Time-Travel 679
Contents xiii
11 Descriptive Statistics 683

11.1 Numerical Descriptions of Data 684
■ Types of Variables ■ Categorical Data ■ Numerical Data: Measures
of Central Tendency ■ Numerical Data: Measures of Spread
■ Numerical Data: The Five-Number Summary ■ Outliers
11.2 Graphical Descriptions of Data 693

■ Displaying Categorical Data ■ Displaying Numerical Data: Histograms
■ Interpreting Area in Histograms ■ The Normal Curve
11.3 Relationships between Variables 703

■ Two Categorical Variables ■ Categorical and Numerical Variables
■ Two Numerical Variables
11.4 Populations, Samples, and Inference 713

■ Populations and Samples ■ Properties of Samples ■ Types of Data ■ Causation
Project: The Birth Weight Paradox 720
Review 722
12 Probability 727
12.1 Principles of Counting 728
■ Permutations ■ Combinations
12.2 What Is Probability? 737

■Experiments, Trials, Outcomes, and Events ■ Probability When Outcomes
Are Equally Likely ■ Probability in General
12.3 Conditional Probability 751

Conditional Probability ■ The Multiplication Rule and Independence
■
The Law of Total Probability ■ Bayes’ Rule

■
Project: Testing for Rare Diseases 766
12.4 Discrete Random Variables 767

■ Describing Discrete Random Variables ■ Mean and Variance of Discrete
Random Variables ■ Bernoulli Random Variables ■ Binomial Random Variables

Project: DNA Supercoiling 783
Project: The Probability of an Avian Influenza Pandemic in Humans 784
12.5 Continuous Random Variables 786

■Describing Continuous Random Variables ■ Mean and Variance of Continuous
Random Variables ■ Exponential Random Variables ■ Normal Random Variables
Review 799
xiv CONTENTS
13 Inferential Statistics 803

13.1 The Sampling Distribution 804
■ Sums of Random Variables ■ The Sampling Distribution of the Mean
■ The Sampling Distribution of the Standard Deviation
13.2 Confidence Intervals 812

■ Interval Estimates ■ Student’s t-Distribution
13.3 Hypothesis Testing 821

■ The Null and Alternative Hypotheses ■ The t-Statistic ■ The P-Value ■ Summary
13.4 Contingency Table Analysis 829

■ Hypothesis Testing with Contingency Tables ■ The Chi-Squared Test Statistic
■ The Hypothesis Test • Summary
Review 835
Appendixes 839
A Intervals, Inequalities, and Absolute Values 840
B Coordinate Geometry 845
C Trigonometry 855
D Precise Definitions of Limits 864
E A Few Proofs 870
F Sigma Notation 874
G Complex Numbers 880
H Statistical Tables 888
Glossary of Biological Terms 891
Answers to Odd-Numbered Exercises 893
biological Index 947
Index 957
Preface
In recent years more and more colleges and universities have been introducing calculus
courses specifically for students in the life sciences. This reflects a growing recognition
that mathematics has become an indispensable part of any comprehensive training in the
biological sciences.
Our chief goal in writing this textbook is to show students how calculus relates to
biology. We motivate and illustrate the topics of calculus with examples drawn from
many areas of biology, including genetics, biomechanics, medicine, pharmacology,
physiology, ecology, epidemiology, and evolution, to name a few. We have paid par-
ticular attention to ensuring that all applications of the mathematics are genuine, and we
provide references to the primary biological literature for many of these so that students
and instructors can explore the applications in greater depth.
We strive for a style that maintains rigor without being overly formal. Although our
focus is on the interface between mathematics and the life sciences, the logical structure
of the book is motivated by the mathematical material. Students will come away from a
course based on this book with a sound knowledge of mathematics and an understanding
of the importance of mathematical arguments. Equally important, they will also come
away with a clear understanding of how these mathematical concepts and techniques are
central in the life sciences, just as they are in physics, chemistry, and engineering.
The book begins with a prologue entitled Mathematics and Biology detailing how the
applications of mathematics to biology have proliferated over the past several decades
and giving a preview of some of the ways in which calculus provides insight into biologi-
cal phenomena.
Alternate Versions
There are two versions of this textbook. The first is entitled Biocalculus: Calculus for
the Life Sciences; it focuses on calculus and some elements of linear algebra that are
important in the life sciences. This is the second version, entitled Biocalculus: Calculus,
Probability, and Statistics for the Life Sciences; it contains all of the content of the first
version as well as three additional chapters titled Descriptive Statistics, Probability, and
Inferential Statistics (see Content on page xviii).
Features
■ Real-World Data
We think it’s important for students to see and work with real-world data in both numeri-
cal and graphical form. Accordingly, we have used data concerning biological phenom-
ena to introduce, motivate, and illustrate the concepts of calculus. Many of the examples
and exercises deal with functions defined by such numerical data or graphs. See, for
example, Figure 1.1.1 (electrocardiogram), Figure 1.1.23 (malarial fever), Exercise
1.1.26 (blood alcohol concentration), Table 2 in Section 1.4 (HIV density), Table 3 in
Section 1.5 (species richness in bat caves), Example 3.1.7 (growth of malarial parasites),
xv
xvi preface
Exercise 3.1.42 (salmon swimming speed), Exercises 4.1.7–8 (influenza pandemic),

Exercise 4.2.10 (HIV prevalence), Figure 5.1.17 (measles pathogenesis), Exercise 5.1.11
(SARS incidence), Figure 6.1.8 and Example 6.1.4 (cerebral blood flow), Table 1 and
Figure 1 in Section 7.1 (yeast population), Figure 8.1.14 (antigenic cartography), Exer-
cises 9.1.7, 9.2.48, and Examples 9.5.5 and 9.6.6 (snake reversals and stripes). And, of
course, Chapters 11 and 13 are focused entirely on the analysis of biological data.
■ Graded Exercise Sets

Each exercise set is carefully graded, progressing from basic conceptual exercises and
skill-development problems to more challenging problems involving applications and
proofs.
■ Conceptual Exercises
One of the goals of calculus instruction is conceptual understanding, and the most impor-
tant way to foster conceptual understanding is through the problems that we assign.
To that end we have devised various types of problems. Some exercise sets begin with
requests to explain the meanings of the basic concepts of the section. (See, for instance,
the first few exercises in Sections 2.3, 2.5, 3.3, 4.1, and 8.2.) Similarly, all the review
sections begin with a Concept Check and a True-False Quiz. Other exercises test concep-
tual understanding through graphs or tables (see Exercises 3.1.11, 5.2.41–43, 7.1.9–11,
9.1.1–2, and 9.1.26–32).
Another type of exercise uses verbal description to test conceptual understanding (see
Exercises 2.5.12, 3.2.50, 4.3.47, and 5.8.29).
■ Projects
One way of involving students and making them active learners is to have them work
(perhaps in groups) on extended projects that give a feeling of substantial accomplish-
ment when completed. We have provided 24 projects in Biocalculus: Calculus for the
Life Sciences and an additional four in Biocalculus: Calculus, Probability, and Statistics
for the Life Sciences. Drug Resistance in Malaria (page 78), for example, asks students
to construct a recursion for the frequency of the gene that causes resistance to an anti-
malarial drug. The project Flapping and Gliding (page 297) asks how birds can mini-
mize power and energy by flapping their wings versus gliding. In The Tragedy of the
Commons: An Introduction to Game Theory (page 298), two companies are exploiting
the same fish population and students determine optimal fishing efforts. The project Dis-
ease Progression and Immunity (page 394) is a nice application of areas between curves.
DNA Supercoiling (page 783) uses ideas from probability theory to predict how DNA
is coiled and compacted into cells. We think that, even when projects are not assigned,
students might well be intrigued by them when they come across them between sections.
■ Case Studies
We also provide two case studies: (1) Kill Curves and Antibiotic Effectiveness and
(2) Hosts, Parasites, and Time-Travel. These are extended real-world applications from
the primary literature that are more involved than the projects and that tie together mul-
tiple mathematical ideas throughout the book. An introduction to each case study is pro-
vided at the beginning of the book (page xli), and then each case study recurs in various
chapters as the student learns additional mathematical techniques. The case studies can
be used at the beginning of a course as motivation for learning the mathematics, and they
can then be returned to throughout the course as they recur in the textbook. Alternatively,
preface xvii
a case study may be assigned at the end of a course so students can work through all com-
ponents of the case study in its entirety once all of the mathematical ideas are in place.
Case studies might also be assigned to students as term projects. Additional case studies
will be posted on the website www.stewartcalculus.com as they become available.
■ Biology Background
Although we give the biological background for each of the applications throughout the
textbook, it is sometimes useful to have additional information about how the biological
phenomenon was translated into the language of mathematics. In order to maintain
a clear and logical flow of the mathematical ideas in the text, we have therefore included
such information, along with animations, further references, and downloadable data on
the website www.stewartcalculus.com. Applications for which such additional informa-
tion is available are marked with the icon BB in the text.
■ Technology
The availability of technology makes it more important to clearly understand the con-
cepts that underlie the images on the screen. But, when properly used, graphing calcula-
tors and computers are powerful tools for discovering and understanding those concepts.
(See the section Calculators, Computers, and Other Graphing Devices on page xxvi for
a discussion of these and other computing devices.) These textbooks can be used either
with or without technology and we use two special symbols to indicate clearly when a
particular type of machine is required. The icon ; indicates an exercise that definitely
requires the use of such technology, but that is not to say that it can’t be used on the other
exercises as well. The symbol CAS is reserved for problems in which the full resources
of a computer algebra system (like Maple, Mathematica, or the TI-89/92) are required.
But technology doesn’t make pencil and paper obsolete. Hand calculation and sketches
are often preferable to technology for illustrating and reinforcing some concepts. Both
instructors and students need to develop the ability to decide where the hand or the
machine is appropriate.
■ Tools for Enriching Calculus (TEC)

TEC is a companion to the text and is intended to enrich and complement its contents. (It
is now accessible in Enhanced WebAssign and CengageBrain.com. Selected Visuals and
Modules are available at www.stewartcalculus.com.) Developed in collaboration with
Harvey Keynes, Dan Clegg, and Hubert Hohn, TEC uses a discovery and exploratory
approach. In sections of the book where technology is particularly appropriate, marginal
icons TEC direct students to TEC Visuals and Modules that provide a laboratory environ-
ment in which they can explore the topic in different ways and at different levels. Visuals
are animations of figures in text; Modules are more elaborate activities and include
exercises. Instructors can choose to become involved at several different levels, ranging
from simply encouraging students to use the Visuals and Modules for independent explo-
ration, to assigning specific exercises from those included with each Module, to creating
additional exercises, labs, and projects that make use of the Visuals and Modules.
■ Enhanced WebAssign
Technology is having an impact on the way homework is assigned to students, particu-
larly in large classes. The use of online homework is growing and its appeal depends on
ease of use, grading precision, and reliability. We have been working with the calculus
xviii preface
community and WebAssign to develop a robust online homework system. Up to 50% of

the exercises in each section are assignable as online homework, including free response,
multiple choice, and multi-part formats.
The system also includes Active Examples, in which students are guided in step-by-
step tutorials through text examples, with links to the textbook and to video solutions.
The system features a customizable YouBook, a Show My Work feature, Just in Time
review of precalculus prerequisites, an Assignment Editor, and an Answer Evaluator that
accepts mathematically equivalent answers and allows for homework grading in much
the same way that an instructor grades.
■ Website
The site www.stewartcalculus.com includes the following.
■ Algebra Review
■ Lies My Calculator and Computer Told Me
■ History of Mathematics, with links to the better historical websites
■ Additional Topics (complete with exercise sets): Approximate Integration: The
Trapezoidal Rule and Simpson’s Rule, First-Order Linear Differential Equations,
Second-Order Linear Differential Equations, Double Integrals, Infinite Series, and
Fourier Series
■ Archived Problems (drill exercises and their solutions)
■ Challenge Problems
■ Links, for particular topics, to outside Web resources
■ Selected Tools for Enriching Calculus (TEC) Modules and Visuals
■ Case Studies
■ Biology Background material, denoted by the icon BB in the text
■ Data sets
Content
Diagnostic Tests The books begin with four diagnostic tests, in Basic Algebra, Ana-
lytic Geometry, Functions, and Trigonometry.
Prologue This is an essay entitled Mathematics and Biology. It details how the appli-
cations of mathematics to biology have proliferated over the past several decades and
highlights some of the applications that will appear throughout the book.
Case Studies The case studies are introduced here so that they can be used as moti-
vation for learning the mathematics. Each case study then recurs at the ends of various
chapters throughout the book.
1 Functions and Sequences The first three sections are a review of functions from
precalculus, but in the context of biological applications. Sections 1.4 and 1.5 review
exponential and logarithmic functions; the latter section includes semilog and log-
log plots because of their importance in the life sciences. The final section introduces
sequences at a much earlier stage than in most calculus books. Emphasis is placed on
recursive sequences, that is, difference equations, allowing us to discuss discrete-time
models in the biological sciences.
preface xix
2 Limits We begin with limits of sequences as a follow-up to their introduction in

Section 1.6. We feel that the basic idea of a limit is best understood in the context of
sequences. Then it makes sense to follow with the limit of a function at infinity, which
we present in the setting of the Monod growth function. Then we consider limits of
functions at finite numbers, first geometrically and numerically, then algebraically. (The
precise definition is given in Appendix D.) Continuity is illustrated by population har-
vesting and collapse.
3 Derivatives Derivatives are introduced in the context of rate of change of blood
alcohol concentration and tangent lines. All the basic functions, including the exponen-
tial and logarithmic functions, are differentiated here. When derivatives are computed in
applied settings, students are asked to explain their meanings.
4 Applications of Derivatives The basic facts concerning extreme values and shapes
of curves are deduced using the Mean Value Theorem as the starting point. In the sec-
tion on l’Hospital’s Rule we use it to compare rates of growth of functions. Among the
applications of optimization, we investigate foraging by bumblebees and aquatic birds.
The Stability Criterion for Recursive Sequences is justified intuitively and a proof based
on the Mean Value Theorem is given in Appendix E.
5 Integrals The definite integral is motivated by the area problem, the distance prob-
lem, and the measles pathogenesis problem. (The area under the pathogenesis curve up
to the time symptoms occur is equal to the total amount of infection needed to develop
symptoms.) Emphasis is placed on explaining the meanings of integrals in various con-
texts and on estimating their values from graphs and tables. There is no separate chapter
on techniques of integration, but substitution and parts are covered here, as well as the
simplest cases of partial fractions.
6 Applications of Integrals The Kety-Schmidt method for measuring cerebral blood
flow is presented as an application of areas between curves. Other applications include
the average value of a fish population, blood flow in arteries, the cardiac output of the
heart, and the volume of a liver.
7 Differential Equations Modeling is the theme that unifies this introductory treat-
ment of differential equations. The chapter begins by constructing a model for yeast pop-
ulation size as a way to motivate the formulation of differential equations. We then show
how phase plots allow us to gain considerable qualitative information about the behavior
of differential equations; phase plots also provide a simple introduction to bifurcation
theory. Examples range from cancer progression to individual growth, to ecology, to
anesthesiology. Direction fields and Euler’s method are then studied before separable
equations are solved explicitly, so that qualitative, numerical, and analytical approaches
are given equal consideration. The final two sections of this chapter explore systems of
two differential equations. This brief introduction is given here because it allows students
to see some applications of systems of differential equations without requiring any addi-
tional mathematical preparation. A more complete treatment is then given in Chapter 10.
8 Vectors and Matrix Models We start by introducing higher-dimensional coordi-
nate systems and their applications in the life sciences including antigenic cartography
and genome expression profiles. Vectors are then introduced, along with the dot product,
and these are shown to provide insight ranging from influenza epidemiology, to cardiol-
ogy, to vaccine escape, to the discovery of new biological compounds. They also provide
some of the tools necessary for the treatment of multivariable calculus in Chapter 9.
The remainder of this chapter is then devoted to the application of further ideas from
linear algebra to biology. A brief introduction to matrix algebra is followed by a section
where these ideas are used to model many different biological phenomena with the aid
of matrix diagrams. The final three sections are devoted to the mathematical analysis of
xx preface
such models. This includes a treatment of eigenvalues and eigenvectors, which will also
be needed as preparation for Chapter 10, and a treatment of the long-term behavior of
matrix models using Perron-Frobenius Theory.
9 Multivariable Calculus Partial derivatives are introduced by looking at a specific
column in a table of values of the heat index (perceived air temperature) as a function of
the actual temperature and the relative humidity. Applications include body mass index,
infectious disease control, lizard energy expenditure, and removal of urea from the blood
in dialysis. If there isn’t time to cover the entire chapter, then it would make sense to
cover just sections 9.1 and 9.2 (preceded by 8.1) and perhaps 9.6. But if Section 9.5 is
covered, then Sections 8.2 and 8.3 are prerequisites.
10 Systems of Linear Differential Equations Again modeling is the theme that uni-
fies this chapter. Systems of linear differential equations enjoy very wide application in
the life sciences and they also form the basis for the study of systems of nonlinear dif-
ferential equations. To aid in visualization we focus on two-dimensional systems, and we
begin with a qualitative exploration of the different sorts of behaviors that are possible
in the context of population dynamics and radioimmunotherapy. The general solution to
two-dimensional systems is then derived with the use of eigenvalues and eigenvectors.
The third section then illustrates these results with four extended applications involving
metapopulations, the immune system, gene regulation, and the transport of environmen-
tal pollutants. The chapter ends with a section that shows how the ideas from systems
of linear differential equations can be used to understand local stability properties of
equilibria in systems of nonlinear differential equations. To cover this chapter students
will first need sections 8.1–8.4 and 8.6–8.7.
11 Descriptive Statistics Statistical analyses are central in most areas of biology. The
basic ideas of descriptive statistics are presented here, including types of variables, mea-
sures of central tendency and spread, and graphical descriptions of data. Single variables
are treated first, followed by an examination of the descriptive statistics for relationships
between variables, including the calculus behind the least-square fit for scatter plots. A
brief introduction to inferential statistics and its relationship to descriptive statistics is
also given, including a discussion of causation in statistical analyses.
12 Probability Probability theory represents an important area of mathematics in the
life sciences and it also forms the foundation for the study of inferential statistics. Basic
principles of counting and their application are introduced first, and these are then used
to motivate an intuitive definition of probability. This definition is then generalized to
the axiomatic definition of probability in an accessible way that highlights the meanings
of the axioms in a biological context. Conditional probability is then introduced with
important applications to disease testing, handedness, color blindness, genetic disorders,
and gender. The final two sections introduce discrete and continuous random variables
and illustrate how these arise naturally in many biological contexts, from disease out-
breaks to DNA supercoiling. They also demonstrate how the concepts of differentiation
and integration are central components of probability theory.
13 Inferential Statistics The final chapter addresses the important issue of how one
takes information from a data set and uses it to make inferences about the population
from which it was collected. We do not provide an exhaustive treatment of inferential
statistics, but instead present some of its core ideas and how they relate to calculus. Sam-
pling distributions are explained, along with confidence intervals and the logic behind
hypothesis testing. The chapter concludes with a simplified treatment of the central ideas
behind contingency table analysis.
preface xxi
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xxii preface
Linear Algebra for Calculus

by Konrad J. Heuvers, William P. Francis, John H. Kuisti,
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preface xxiii
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Acknowledgments
We are grateful to the following reviewers and class testers for sharing their knowledge
and judgment with us. We have learned something from each of them.
Reviewers
Anthony Barcellos, American River Eli Goldwyn, University of California–
College Davis
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in Philadelphia Mark Harbison, Sacramento City College
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"Never mind," broke in Kitty hastily. "It's an old frock, and—oh, Bob,
do be quick and see it's not hurt."
Bob had set the box right side up by this time, and was trying to
open the lid; but it had become jammed, and it took him a few
minutes to prise it open with his pocket-knife.
Tim now saw that the front of the box had been knocked out and a
piece of wire netting put in its place, and that he had not been able to
see from the top of the wall. He began to feel anxiety as well as
curiosity to ascertain what the box contained. Was it something living
—some animal? It might be a pet of some kind, and that would
account for Kitty's saucer of bread and milk.
Bob had succeeded in opening the box now, and he and his sister
were bending over it, their fair, curly heads close together. A strange
quietude seemed to have fallen upon them, and when a minute later
they stood upright and looked at each other, Tim observed that the
colour had fled from both young faces, and that the tears were rolling
down Kitty's cheeks. At length Bob spoke in a voice which sounded
rather husky.
"I say, don't cry like that, old girl!" he said kindly. "I'll go in and ask
father to come out and look at it." And he rushed off into the house.
Meanwhile Kitty stooped over the box again, and took therefrom
something white, which she cuddled in her arms and wept over in
bitterest grief.
The onlooker at the window watched her in consternation, a choking

sensation in his throat. Although he had not yet fully realized the
cause of her trouble, he sympathized with her, for he owned a very
warm heart, and the sight of the little girl's tears touched him
immeasurably. What had he done to cause this grief?
He was soon to know, for in a very short while Bob returned,

followed by his parents, and the two servants, who all congregated
around Kitty, and presently Tim heard Mr. Glanville say, "No dog did
it, so Snip is guiltless, at any rate, although of course he might have
overturned the box, but I don't think he could have done that. It must
have been instantaneous death for the poor thing. You can be sure it
did not suffer at all, Kitty, and that thought is very comforting, so dry
your eyes, there's a good girl. Doubtless, when the box was turned
over the little creature was thrown against the side, and received a
blow on the head. It takes but a small knock to kill a rabbit, especially
a young one like this."
Tim knew what he had done at last, and he was both shocked and
frightened. He was anything but a cruel boy, and he was exceedingly
fond of all animals, and now that he realized that the white, fluffy
object poor Kitty held so tenderly in her arms was a little baby rabbit,
which he in his wicked, revengeful temper had killed, he felt like a
murderer. His first impulse was to push back the lace curtain, and
shout out to the group in the next garden that he was responsible for
the rabbit's death; but a minute's reflection made him change his
mind, and determine to keep his secret. In an agony of contrition, he
watched Bob get a spade and dig a hole under the big apple tree
which grew at the bottom of the garden, whilst one of the servants
fetched a shoebox, into which Kitty placed the rabbit, and then
followed the funeral. Afterwards Bob made a little mound over the
grave, and planted a forget-me-not root upon it. And he told Kitty he
would try to get her another rabbit very soon.
Mr. and Mrs. Glanville and the servants had returned to the house,
but the sister and brother lingered in the garden. They were
conversing in low tones, so Tim could not hear what they were
saying; but his guilty conscience suggested to him that they might be
discussing by what means the rabbit's hutch had been overturned,
and perhaps arriving at the truth.
Suddenly Kitty looked up and saw the curtains in Mr. Shuttleworth's

dining-room window move, and it flashed upon her that Tim was
behind them.
"That boy next door—he is watching us again!" she whispered

excitedly to her brother. "Oh, Bob, I wonder if he knows who killed
my dear, dear little rabbit. You don't think he could have thrown over
the hutch to spite us, do you?" she suggested with a condemning
glance in Tim's direction.
Bob shook his head—he thought the idea most unlikely; and a few
minutes later he and Kitty went indoors, little dreaming how uneasy
Tim was feeling, for he had known they had guessed he was at the
window, and Kitty's incriminating glances had not been lost upon
him.
Poor Tim! He was utterly miserable for the remainder of the day, and
so dispirited and dejected did he look that even Mr. Shuttleworth
noticed it, and asked him if he was ill.
Ill? No, he was not ill, he declared; and oh, how he longed to
unburden his heart to his uncle! But he shrank from doing so. And,
saying he was tired, remarked he thought he would go to bed early,
which he accordingly did.
The clocks in the house struck many times before he was able to get
any sleep, so tormented was he by his guilty conscience and the fear
that the sharp-looking children next door were beginning to have
suspicions respecting him. And when at last slumber overcame him,
and he fell into a troubled doze, he was confronted in his dream by a
vision of Kitty Glanville, her blue eyes full of angry tears, whilst in a
voice shrill with accusation, she cried, "Ah, I've found you out now! It
is you who killed my rabbit."
CHAPTER III.
KITTY INTERVIEWS TIM.
"I'VE spoken to Tom Hatch about getting you another rabbit," Bob
informed his sister on the afternoon subsequent to the one on which
they had found her late pet dead. "And he says he believes his
brother will be able to let you have one, for his doe had young a few
weeks back, and he wants to get rid of them all before leaving the
place."
The Hatch boys were schoolfellows of Bob's, who were about to

leave the place with their parents to take up their abode in a
neighbouring town. Kitty had purchased the rabbit, which had come
to such an untimely end, from Tom for a shilling, and was very
desirous to replace it.
"Will he be willing to sell me another for the same price?" she

inquired practically.
"Oh, yes," was the response. "You may be sure of that."
"And will it be a white Angora like the other?"
"I suppose so; he didn't say; you needn't have it if you don't fancy it,
you know. I think we had better move the hutch closer to the house,
near to the back door."
"Yes, then the servants will be able to keep their eyes on it, and see
it is not interfered with. Oh, Bob, I cannot help thinking that the boy
next door may know how the hutch got overturned yesterday, for he's
always watching us! At any rate, I shall ask him; there can be no
harm in doing that."
"I should not have anything to say to him, if I were you," advised
Bob; "but please yourself, of course."
Thus it came about that whilst Tim, who was feeling much bored with
his own company and was very dispirited, was doing a little
gardening by way of passing the time after tea that evening, he
heard himself addressed by Kitty's now familiar voice:
"Hi, you boy—I don't know your name—I want to speak to you."
The hoe with which he was working dropped from Tim's hands, so
startled was he, and the expression of his face was one of alarm as
he looked around hastily; but he could not see the little girl, though
he ran his eyes from end to end of the partition wall.
"Here I am," she said, with a merry laugh as she observed his
bewilderment. "Why, you seem quite scared," she continued. "Don't
you see me? I'm in the apple tree."
There she was sure enough, perched high on a branch of the big
apple tree at the bottom of her own garden, from which position she
could overlook Mr. Shuttleworth's domain.
"Oh," exclaimed Tim, "now I see you. I couldn't think where your
voice came from."
A smile, which was rather embarrassed, though certainly not

unfriendly, flickered over his plain, freckled countenance as he
spoke.
"I want to speak to you for a minute about something important," said
Kitty.
"Oh!" He wondered uneasily what "something important" might be.
"You remember meeting us—my brother and me—in the town

yesterday morning?"
"Yes," assented Tim. "So you did see me. I was positive of it at the
time, though you kept your head turned aside."
"That was because I was eating sweets, and there was a big
caramel in my mouth," she said hastily, looking somewhat abashed.
"Not that I should have spoken to you, anyway," she went on
truthfully. "You couldn't have expected either of us to do that after—
but never mind that now! What I want to know is, where did you go
afterwards?"
"Where did I go afterwards? I—I—why do you ask? What does it

matter to you?"
What a rude boy he was, Kitty thought. She flushed with annoyance;
but she was so anxious to ascertain if he could throw any light on the
matter which weighed upon her mind that she answered pacifically:
"It doesn't matter to me, except that I thought, if you came straight
home, you might have noticed if there was any one prowling about
our garden. The fact is," speaking in a confidential tone, "I had a
dear little rabbit in a big box against the wall, and some one upset
the box and killed the rabbit. Perhaps you know that?"
"I—I—yes," Tim admitted; "I saw you all in the garden when—when
you found it dead, and—and—I was sorry—"
He paused in confusion, whilst Kitty regarded him more favourably,

for he really did look sorry, quite distressed, in fact.
"It was a sweet little creature," she said with a sigh, "and it was so
sad to find it killed. I daresay you thought me silly and babyish to cry,
but really I couldn't help it. I had only bought it the night before; I
gave a shilling for it. That wasn't what made me cry, though; it was
because it was such a dear, so soft and as white as snow." She
paused and blinked away a tear, then proceeded more briskly: "Well,
what I want to know is, did you see any one interfering with the rabbit
hutch?"
"No," Tim answered, so hesitatingly, that Kitty shot a glance full of

suspicion at him. "Perhaps some strange dog got into your garden,"
he suggested, feeling himself to be very mean-spirited as he spoke,
"or perhaps the wind—"
"Oh, it couldn't have been the wind," she interrupted impatiently,
"although there was a strong breeze blowing, but not strong enough
to upset a heavy box like that. Bob thinks a big dog must have
pushed the hutch over in trying to get it open, but I don't know what
to think, except that you know something about it," she declared with
a ring of decision in her tone.
Tim was so taken aback at this sudden and direct charge that he had
no answer ready. The colour rushed to his face in a flood of crimson,
then, receding, left him quite pale.
"What do you mean?" he gasped at length, assuming anger to hide

his dismay. "How dare you say that I know something about it?"
"You said you'd pay us out because Snip had spoilt your garden—we
didn't know he'd done it, so it wasn't our fault—and I thought you
might have killed my rabbit out of spite."
"I never knew your rabbit was dead till I saw it in your arms,"
declared Tim solemnly. "I hadn't the faintest idea there was a rabbit
in the box, I didn't know what was there."
"But did you overturn the box?" persisted Kitty.
For a moment Tim hesitated. He still craved for the friendship of Kitty
and her brother, and he thought if he acknowledged his guilt they
would never have anything to do with him, so, though he was usually
truthful, on this occasion he gave way to the temptation of the
moment, and answered:
"No, certainly not."
He did not look at Kitty as he spoke; and when several minutes had
elapsed without her having addressed him again, he plucked up
courage to glance furtively towards the apple tree, he found the little
girl had gone. He did not know that he had lied in vain, or guess that
Kitty, who was very keen of discernment, believed he held the key of
the mystery which surrounded her rabbit's death.
"If that boy next door didn't throw over the box himself, he knows
who did it," Kitty declared to her brother after she had given him an
account of her interview with Tim.
"Perhaps he was only pretending to know, just to make himself seem

important," suggested Bob. "Some fellows are like that. What good
have you done by speaking to him, Kitty?"
"None at all," she was bound to admit; "and I wish I had let him be.
He seemed sorry my rabbit was dead; but he's a very odd boy, he
hardly had a word to say for himself."
"He had plenty of words yesterday when he was in a passion," Bob

rejoined with a laugh. "Cook hears that he is Mr. Shuttleworth's
nephew, and that he's going to stay here all the summer," he
proceeded to explain. "He will have rather a dull time, I should say,
for Mr. Shuttleworth doesn't appear to take much notice of him, does
he?"
"No, indeed," agreed Kitty. "If he was a nicer boy, he might be

friendly with us; but he's a dreadful temper and I believe he tells
lies."
"To whom are you giving such a bad character, Kitty?" asked Mrs.
Glanville, coming into the room where her children were talking at
that moment, holding an open letter in her hand.
"The boy next door, mother," the little girl answered promptly.
"What do you know against him, my dear?"
Kitty told all she knew—of the exhibition of temper Tim had given
them on the previous day, and of her suspicion against him, which
she was obliged to admit her brother did not share.
"I don't think you ought to jump to the conclusion that the boy has
told you an untruth, Kitty," Mrs. Glanville said gravely. "If you have a
doubt—well, give him the benefit of the doubt, my dear. Strangely
enough I was coming to speak to you about the boy next door. I find
Mr. Shuttleworth is his uncle, and that he has come to pay him a long
visit because he was very ill a month or so ago, and the doctor has
advised his not returning to school for the summer term. His home is
in Dublin—"
"That accounts for his accent then," Bob broke in. "I knew he was a
Paddy the minute I heard him speak."
"It would be wonderful if he had no touch of brogue, considering he

has lived all his life in Ireland," Mrs. Glanville remarked smilingly.
"How do you know that, mother?" inquired Kitty. "Have you been
talking to Mr. Shuttleworth?"
"No; but I've had a letter from Mr. Shuttleworth's sister-in-law, telling
me her eldest boy—Tim, she calls him, short for Timothy, I suppose
—is staying with his uncle here, and asking me to be kind to him."
"How extraordinary!" exclaimed Kitty, whilst her brother gave a soft

whistle of surprise.
"Not so extraordinary as you think, my dear, seeing that Tim's mother

was once a school-friend of mine. She was an Irish girl, and I knew
she had married some one called Shuttleworth, but it never occurred
to me that her husband might be in any way connected with our
neighbour. Tim's mother does not know we are living next door to her
brother-in-law; she simply addressed her letter to this town hoping
that I should get it, and as we are the only Glanvilles in the place, it
came direct."
"And now, what do you mean to do, mother?" Kitty asked eagerly, as
Mrs. Glanville paused and glanced through her letter once more.
"I shall call next door to see my old friend's son to-morrow," was the
response, "and most probably bring him back with me to tea. You
must not be prejudiced against him, children; if he is anything like his
mother in disposition, you will be sure to get on with him, and there
will be no difficulty in your making friends with him, for she was one
of the kindest-hearted girls I ever knew. Don't you think, when he
perched himself on the ladder and stared at you over the wall, he
might have been wishful of making your acquaintance? That is my
opinion. Had I been in your place, Bob, I fancy I should have spoken
to him; he must be about your age—a trifle younger perhaps, but not
much. You must try to be friends with him—a stranger in a strange
land. I daresay he has been home-sick, poor child!"
"What a nuisance!" exclaimed Bob, as soon as his mother had left

the room. "You will see that boy will spoil our holidays—or what is left
of them. We shall have to be civil to him—which he certainly has not
been to us!"
CHAPTER IV.
MAKING AMENDS.
WHEN Mrs. Glanville called at her neighbour's house on the

following afternoon, she found that Tim was not at home; but she
saw Mr. Shuttleworth and explained her mission to him. He had not
the faintest idea where his nephew had gone, he informed her, and
greatly regretted his absence. "For I should like him to be friendly
with your young people," he said cordially.
"I thought perhaps I should have persuaded him to return with me,
and make their acquaintance now," Mrs. Glanville replied. "But, since
he is not here and you do not know what time he will be back, will
you let him come to tea with us to-morrow? Please let him come
early so as to spend a long afternoon with us."
"Certainly," Mr. Shuttleworth agreed readily. "You are very kind and I
have much pleasure in accepting your invitation for Tim." And thus it
was settled.
Meanwhile Tim had gone on a secret errand. Since his interview with
Kitty, he had been possessed with a strong desire to make good the
bad turn he had done her, as far as lay in his power, by giving her
another rabbit. With that idea he had asked the butcher boy, at the
back door earlier in the day, if he knew any one who had rabbits for
sale, and the butcher boy had told him of a shop, kept by an old man
named Jacob Dottin, in a back street of the town, where all sorts of
animals and birds might be purchased. So whilst Mrs. Glanville was
interviewing his uncle, Tim was making for the abode of Mr. Dottin.
An obliging policeman had shown the little boy the way to go; and, at
length, after traversing several narrow, dirty streets, he found himself
before the shop he wanted.
Tom stood for a few minutes gazing into the window at rows of cages
containing birds of all sorts and descriptions, and, so engrossed was
he in watching them that he did not notice the shop door open, and
he started when a voice at his side addressed him:—
"Would you like to come and have a look at my little family, young
gentleman."
Tim turned quickly and looked at the speaker—a spare, bent-

shouldered old man, with grizzled hair which was so thick, as was
his beard, that, with a cap drawn far over his forehead, but little of his
face was visible except a pair of sharp, black eyes and a hook nose.
Tim smiled involuntarily, for he thought the old man was like an
animal himself, an animal that was a mixture of a baboon and a
parrot with his hairy face, and hook nose, and claw-like fingers,
which he twisted together as he blinked and smiled in what he
evidently intended to be an amiable manner.
"Mr. Tottin?" said Tim, inquiringly.
"Aye—Jacob Dottin, young gentleman, at your service," was the

response.
"I wanted to see you, so I will come in," and Tim followed the old
man into the shop.
His first impulse was to retrace his footsteps, for the air in the shop
was close and unpleasant, and he felt he could not breathe there; he
did not retreat, however, as, on glancing around him, he was
fascinated by the sight of a big monkey asleep in a corner with a
small, terrier puppy cuddled in its arms, and various other animals
such as guinea-pigs, rabbits, mice and rats, ranged in hutches
around the walls. Several parrots screamed in cages suspended
from the ceiling, and a raven croaked on a shelf over the door. Tim
grew accustomed to the babel of sounds in a few minutes, and did
not so much notice the offensive atmosphere, and, as the old man
desired him politely to look around, he did so at his leisure, finally
drawing up before a hutch in which were several young rabbits with
beautifully thick, white hair.
"Ah, those are worth looking at," remarked Mr. Dottin; "pure-bred
Angoras, they are. Maybe you're a rabbit fancier?" he questioned,
regarding his visitor shrewdly.
"No," Tim answered, "but I want to buy a rabbit for a—a—some one I
know. What would be the price of one of those, now?"
"Five shillings," was the unhesitating response.
"Five shillings!" echoed Tim, his face clouding over. "Oh dear, so
much as that? Then it's out of the question my buying one to-day,
thank you."
He turned towards the door as he spoke, but the old man stopped
his exit by saying hastily:
"Wait a minute, sir; don't be in such a hurry; perhaps we may be able
to come to terms."
"I'm afraid not," Tim replied regretfully. "For I haven't five shillings in
the world; that is the truth."
"Well, well, you're frank, and I like you for it. They're beautiful
creatures and pure-bred, as I said just now," Mr. Dottin observed
thoughtfully, surveying the rabbits and then Tim with his head on one
side.
"But I'd wish to oblige you and make a new customer, who'd no
doubt recommend me to his friends. Would three shillings be nearer
your prize? Well, then," as the little boy shook his head, "say half-a-
crown? Half-a-crown for a pure-bred Angora rabbit, why, it's absurd;
nevertheless, you shall have that little one for half-a-crown. What do
you say?"
Tim reflected. He was a shrewd boy, and it struck him that Mr. Dottin
had lowered his price very quickly. He also remembered that Kitty
had said she had given a shilling for the rabbit he had killed, so,
though half-a-crown was the exact amount of money he had in his
pocket, he determined to try to get what he wanted for less.
"I'll give you eighteen-pence for that little rabbit, it's the smallest of
the lot, I see," he said. "I don't believe it's worth more."
"Not worth more. Eighteen-pence for a pure-bred Angora! Why, you

can't know what you're talking about, young gentleman!" Mr. Dottin
cried, in shrill accents of protestation.
"Oh, yes I do," Tim returned, confidently. "I price the rabbit at
eighteen-pence."
"And I price it at half-a-crown," the old man retorted. "Stop, stop," he

proceeded, as Tim again moved toward the door. "You are too
impetuous, sir, too impetuous by half. I'll tell you how we'll settle the
matter, we'll split the difference."
"Split the difference?" echoed Tim doubtfully. "I don't understand.
You mean—"
"That you shall have the rabbit for two shillings—ready money, of
course. You price the rabbit at eighteen-pence, I price it at half-a-
crown. Eighteen-pence from a half-a-crown leaves a shilling—split
that shilling and the price of the rabbit is two shillings. See? There
now, that's fair, isn't it?"
"Yes, I think it is," Tim answered, smiling. "I'll give you two shillings
for the rabbit."
"You've a business head on your shoulders. I perceive," observed

Mr. Dottin, as he proceeded to open the hutch, "and I like you the
better for it. Have you anything to put the rabbit in?"
"No; but I can easily carry it inside my coat, it will be quiet there,
won't it?"
"Quiet enough, but mind not to squeeze it—these white rabbits are
delicate creatures. Well, I'm pleased to have done business with you
and I shall hope to see you again. You're a stranger in the town, I
take it? Ah, I thought as much; if you'd lived long in the place you'd
have found me out before. Any time you find yourself this way come
in and have a look round, I shall be glad to see you, and bring your
friends."
"Thank you," Tim replied; "I have no friends in the town, though I
may have some later on. Perhaps I may call in again on another
occasion as you've been so kind as to ask me."
The little boy paid his money, and unfastening a couple of buttons of
his double-breasted coat, he put the rabbit inside, where it appeared
quite comfortable and happy. Then he said good-bye to Mr. Dottin,
who parted from him most affably, and set out for home. As he went,
he turned over in his mind ways and means of conveying the rabbit
to Kitty. He wanted the little girl to have it without knowing, in the first
place, whom it came from. And, later on, he intended to tell her that
he was the donor. After which, he assured himself sanguinely, she
and her brother would gladly—and gratefully—allow him to be their
friend.
Tim planned everything very carefully. That night, he meant to get

over the partition wall, by means of the ladder, and put the rabbit in
the hutch, which he had perceived had been placed near the back
door of the house. No one would see him do it after dark; and to-
morrow morning Kitty and Bob would discover the rabbit; then, on
the first favourable opportunity, he would confess that he had bought
the rabbit to replace their dead pet. How surprised they would be
and how kind they would consider him, whilst he would have salved
his conscience to a very great extent.
It was nearly tea-time when Tim arrived at home, so he ran straight

upstairs and put the rabbit into one of the drawers in the set in his
bedroom, leaving the drawer a little open to admit the air. Then, after
washing his hands and brushing his hair, he went downstairs and
joined his uncle, who immediately told him of Mrs. Glanville's visit
and the purport of it.
"Mrs. Glanville knew mother years ago, and she has asked me to tea
to-morrow!" he exclaimed delightedly, his eyes sparkling with
excitement, after he had listened to Mr. Shuttleworth's tale. "Oh,
Uncle John, how jolly of her!"
"But I thought you didn't think much of the next door children,"
remarked Mr. Shuttleworth, with an amused smile. "If I remember
rightly you said you considered them stuck-up."
"That was because they wouldn't have anything to do with me; but
now it will be all right. We shall soon be good friends, you will see."
"I hope so," Mr. Shuttleworth replied. "You are to go early to-morrow
afternoon, remember."
"Oh, I will not forget," Tim returned. "No fear of that!"

After tea, he waited impatiently for darkness. The evenings were
light until seven o'clock now, consequently it was nearly eight before
he ventured to fetch the rabbit from the drawer and steal out of doors
with it. In the tool-house he found a basket with a cover, into which
he placed the little animal with some bits of bread he had smuggled
from the tea-table, remembering it would require food during the
night. And then, he procured the ladder, and a minute later, basket in
hand, he was sitting astride the partition wall. It was the work of
another minute only to transfer the light ladder from one side of the
wall to the other, and to descend in the Glanville's garden.
For a few seconds Tim stood quite still listening intently. No one was
about; the servants were in the kitchen, for he could see the
reflections of two figures on the kitchen blind, and the rest of the
household, he guessed, would be in the front part of the house. Very
slowly and cautiously the little boy felt his way to the spot where he
knew the rabbit hutch to be, and, having found it, he removed the
rabbit and its supper from the basket to the hutch in safety, and
closed down the lid. His errand thus accomplished, he was startled
to hear Snip begin to bark furiously close inside the back door. Not a
second was now to be lost, he told himself; and, with a wildly beating
heart, he was making a hasty retreat towards the ladder when, to his
dismay, the back door was flung suddenly open, and out flew Snip
with the series of angry "yaps" to which he never failed to give
utterance when in a hot pursuit of a cat.
Tim's dash for the ladder was favoured by success, and he had
reached it and placed his right foot on the first rung of it before the
little dog could scent him and discover his whereabouts in the
darkness. But if the boy's movements were quick, master Snip's
were quicker, and, springing at the intruder, he caught him by one of
the legs of his trousers, fastening his teeth into it with so firm a grip
that Tim knew no amount of kicking and shaking would induce him to
drop off.
CHAPTER V.
A SPOILT PLAN.
ALTHOUGH startled, Tim was not frightened by Snip's attack upon

him. The servants did not follow the dog to discover the cause of his
excitement, no doubt taking it for granted that he was hunting a cat;
and, relieved upon that point, Tim began to hope he might succeed
in disabusing Snip's mind of the idea that he was there for an evil
purpose, so he spoke to him in a conciliatory tone:
"Hulloa, Snip, old man! What's the matter with you, eh? Why, you
know me right enough, don't you?"
Snip immediately recognised the voice as that of the boy next door,
and his firm grip relaxed. Tim stooped and patted him gently,
whereupon he gave a little wriggle of pleased surprise and dropped
his hold of Tim altogether, realizing he had made a mistake and
feeling rather foolish; and after that he stood quietly by whilst Tim
mounted the ladder, pulled it up after him, and disappeared on the
other side of the wall.
Tim did not linger in the garden; but, having put away the ladder, he
entered the house, satisfied that he had carried out his plan as he
had intended, and congratulating himself on the way in which he had
conciliated Snip.
Meanwhile Snip, who was very curious even for a dog, was
exceedingly puzzled by the behaviour of the boy next door, and was
taking a stroll round to see what he had been doing. Apparently all
was as usual; but, on returning to the back door, Snip's sharp nose
made him suspect that there was a rabbit in the hutch, and having
satisfied himself that he was not mistaken, he gave one sharp,
imperative bark, which the servants knew meant that he desired to
call their attention to something.
"What's the matter, Snip?" asked cook, opening the back door.
Snip barked again, more imperatively; and cook came out, a lighted
candle in her hand.
"Why, some one's left a rabbit here!" she exclaimed, greatly

astonished. "I wonder if Miss Kitty and Master Bob were expecting it.
I suppose I'd best call them out."
She accordingly did so, and a short while later the sister and brother
arrived upon the scene, the former full of excitement.
"Who brought it, cook?" she inquired, as Bob took the rabbit out of
the hutch and they examined it by the light of the candle. "Oh, isn't it
a love?" she cried, her face aglow with delight.
"I don't know who brought it, miss," cook answered. "We didn't hear
any one, although Mary and I were both in the kitchen; but Snip
barked to be let out and, as we fancied he heard a cat about, we
opened the back door and he rushed out in a fury. I suppose there
must have been somebody here."
"Most likely," agreed Bob. "And that somebody was frightened away."
"But who could it have been, Master Bob?" asked Mary, the
housemaid, who had come out too.
"I don't know, I'm sure," replied Bob. "I expect it was an errand-boy,
or some one like that, sent by Tom Hatch. I saw Tom this morning,
and he said his brother had given him a couple of his young rabbits,
and you were to have one of them, Kitty; he told me he didn't want to
be paid for it."
"Then it's a present, Bob?" asked Kitty.

"Yes. Tom said he would bring it if he had time—you know the
Hatches are leaving B-to-morrow. I suppose he hadn't time, so he
sent it. Yes, here's the basket it came in." He picked up the basket
which Tim had dropped and forgotten, and examined it. "Snip must
have given some one a fine scare," he proceeded, chuckling with
amusement. "I say, Kitty, it's a jolly little rabbit, isn't it?"
"It's a beauty!" declared Kitty. "I shall keep it combed, and its coat will
soon be lovely. How very kind of Tom Hatch to give it to me! I shan't
be able to thank him, shall I?"
"Oh, don't let that weigh upon your mind. I thanked him for you, so
that's all right," Bob responded carelessly. "I didn't say anything
about it to you before because I thought I'd wait and see if Tom really
meant to give you the rabbit," he proceeded to explain. "The fact is,
he's such a chap for making promises he never carries out. He's
been as good as his word in this case, though. I wonder whom he
sent with the rabbit."
"I am afraid whoever it was must have been dreadfully frightened by

Snip," said cook regretfully. "I'm sure we wouldn't have let him out if
we guessed any one had been here. I'll hang up the basket in the
scullery, for I suppose it will be fetched by someone to-morrow."
Bob replaced the rabbit in the hutch, remarking upon the pieces of
bread which Kitty collected and soaked in milk for her new pet.
The little girl's heart was full of gratitude towards Tom Hatch, who
had, as she naturally believed, from all her brother had said, made
her this most welcome gift.
"I shall call the dear little thing 'Fluffy,'" she confided to her brother. "I
don't think I shall show it to the boy next door, as I feel so certain he
had something to do with the death of the other."
Bob merely shrugged his shoulders on hearing this, and made no

reply, for he and his sister could not agree upon the point. Kitty was
absurdly suspicious, he told himself.
The following afternoon when Tim Shuttleworth—his countenance
shining with the recent application of soap and water—rang the front
door bell of the Glanvilles' house, his heart beat fast with excitement.
He was shown into the dining-room, where the whole family were
assembled, and received a cordial welcome from Mr. and Mrs.
Glanville. The latter, in particular, greeted him very kindly, and kissed
him for his mother's sake, she said—a remark which made him feel
a little choky for a minute, as he was deeply attached to his mother,
and he had greatly missed her since he had been away from her—
and then she introduced him to Kitty and Bob, who shook hands with
him and looked as though they had never seen him in their lives
before. At first, Tim felt a trifle embarrassed; but, as Mrs. Glanville
talked to him and asked him questions, he soon lost all traces of self-
consciousness, and found himself chatting quite easily of his own
people—his mother and father, and his two brothers and three
sisters, all younger than himself.
Then Snip scraped at the door, and, on being admitted, evinced

much delight at the sight of the visitor.
"You are fond of animals," remarked Mrs. Glanville, as Snip promptly

accepted an invitation to sit on Tim's knees, where he settled himself
comfortably with a flattered expression on his face. "I suppose you
keep pets at home, do you not?"
"No," Tim answered, "we have no room for them. Ours is a small
house, and we have no garden. Mother had a canary once, but a
neighbour's cat came in one day when no one was about and killed
it."
"How dreadful!" exclaimed Kitty with a shudder. "What a horrid cat it

must have been! I should hate a cat that killed birds!"
"Should you hate Snip if he killed a cat?" asked Bob, laughing.
"I don't think I should ever like him again," Kitty answered seriously;
"but Snip wouldn't do it."

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1 Functions and Sequences 1

1.2 A Catalog of Essential Functions 17

1.3 New Functions from Old Functions 31

Project The Biomechanics of Human Movement 40

1.4 Exponential Functions 41

1.5 Logarithms; Semilog and Log-Log Plots 52

1.6 Sequences and Difference Equations 70

Discrete-Time Models in the Life Sciences

Project Drug Resistance in Malaria 78

■ The Logistic Sequence in the Long Run

Project Modeling the Dynamics of Viral Infections 101

2.2 Limits of Functions at Infinity 102

2.3 Limits of Functions at Finite Numbers 111

2.4 Limits: Algebraic Methods 125

2.5 Continuity 137

3.2 The Derivative as a Function 168

3.3 Basic Differentiation Formulas 181

3.4 The Product and Quotient Rules 194

3.5 The Chain Rule 202

Arbitrary Bases ■ Longer Chains ■ Implicit Differentiation ■ Related Rates

3.6 Exponential Growth and Decay 215

Project: Controlling Red Blood Cell Loss During Surgery 222

3.7 Derivatives of the Logarithmic and Inverse Tangent Functions 222

3.8 Linear Approximations and Taylor Polynomials 230

4 Applications of Derivatives 249

Project: The Calculus of Rainbows 259

4.2 How Derivatives Affect the Shape of a Graph 261

4.3 L’Hospital’s Rule: Comparing Rates of Growth 274

Indeterminate Products ■ Indeterminate Differences

Project: Mutation-Selection Balance in Genetic Diseases 284

4.4 Optimization Problems 285

4.5 Recursions: Equilibria and Stability 299

4.6 Antiderivatives 306

5.2 The Definite Integral 329

5.3 The Fundamental Theorem of Calculus 342

The Fundamental Theorem ■ Differentiation and Integration as Inverse Processes

Project: The Outbreak Size of an Infectious Disease 354

5.4 The Substitution Rule 354

5.5 Integration by Parts 362

5.6 Partial Fractions 368

5.8 Improper Integrals 376

6 Applications of Integrals 387

Project: Disease Progression and Immunity 394

6.2 Average Values 397

6.4 Volumes 405

7 Differential Equations 419

Project: Chaotic Blowflies and the Dynamics of Populations 430

7.2 Phase Plots, Equilibria, and Stability 431

A Mathematical Derivation of the Local Stability Criterion

Project: Catastrophic Population Collapse:

7.3 Direction Fields and Euler’s Method 440

7.4 Separable Equations 449

7.5 Systems of Differential Equations 459

Project: The Flight Path of Hunting Raptors 467

Project: Catastrophic Population Collapse: