Mathematics for Life Scientists

Frederick R. Adler

c
Frederick
R. Adler, 1994

Department of Mathematics and Department of Biology, University of Utah, Salt Lake City, Utah 84112

Preface
Mathematics for Life Scientists teaches calculus, probability, and statistics as a way to introduce
freshman and sophomore life science majors to the insights mathematics can provide into many
aspects of biology. Why should there be a special book for this audience? Although the importance
of quantitative skills in the life sciences is much discussed, current realities tend to conceal their vital
role. Too often, biology is the natural science of last resort for students who believe they arent
cut out for math. Most colleges and universities require little calculus for their biology majors,
and those that do require a full calculus course doubt its worth when students emerge unable to
apply even pre-calculus mathematics in new contexts. Students are left with similar doubts when
the techniques they learned for tests vanish as swiftly from the curriculum as from their memories.
Students, biology faculty, and administrators see that biology is burgeoning as a science and as a
major, apparently unhindered by pervasive mathematical illiteracy.
In fact, mathematics has played an important if under-appreciated role in biology, providing the
impetus for breakthroughs in epidemiology, genetics, statistics, physiology, and many other areas.
As a theoretical biologist who uses mathematics to make sense of complex biological systems, I see
this role expanding, not contracting. Although a great deal of biology can be done without any
mathematics, the powerful new technologies that are transforming fields of biology from genetics
and physiology to ecology are increasingly quantitative, as are many of the questions at the frontiers
of knowledge. Mathematics is the language, the technology of thought, with which these developments are created and controlled. Students who speak this language will be the leaders of the
next generation of biologists. As biology becomes more important in society, mathematical literacy
becomes as necessary for doctors, business people, lawyers, and art historians as for researchers.
My goal in this book is simple: to teach biology majors the mathematical ideas I use every day
in my own research and in collaborations with my more empirical colleagues. These ideas are not
specific techniques like differentiation, but concepts of modeling. The skills include describing
a system, translating appropriate aspects into equations, and interpreting results in terms of the
original problem. In this process, the science is central and solving the equations is in some ways
the least important step.
Because a few dynamical principles underlie a remarkable diversity of biological processes, this
book follows three themes throughout: growth, diffusion and selection. Each theme is studied
in turn with the three kinds of model that structure the course: discrete-time dynamical systems, differential equations, and stochastic processes. Techniques and insights build on each other
throughout the course. Along the way, students learn and apply the standard material of a calculus
course (differentiation, integration, and their applications).
In addition, the course introduces matrices, vectors, and some basic calculus in two dimensions,
all in a dynamical context. Most significantly, the final section of the book teaches probability
1

2
and statistics from the same perspective, using discrete-time dynamical systems and differential
equations to describe simple stochastic processes. This section shows that correct and flexible
application of statistics requires understanding the processes that generate data, and introduces
the fundamental statistical notions of likelihood, parameter estimation, and hypothesis testing.
In many ways, students go farther than in a traditional calculus (or probability) class. Time
is saved by skipping methods made obsolete by computers. Learning more concepts and fewer
techniques is definitely more challenging. As a sweetener, students are given the keys to the powerful techniques professionals use when equations cannot be solved: graphical methods (cobwebbing
and phase space), approximation (leading behavior), and computers (labs using a computer algebra/graphics program). These techniques emphasize reasoning and visualization, and show that
applied mathematics has less to do with algebraic wizardry than with the clear formulation of ideas.
Working with computers has proven to be particularly successful in this context.
What are the benefits of this approach? All instructors know that students will not remember
every technique they have learned. This course emphasizes understanding what a model is, and
recognizing what models say. To be able to recognize a differential equation, interpret the
terms, and use the solution is far more important than knowing how to find the solution. These
reasoning skills, in addition to familiarity with models in general, are what stay with the motivated
student, and are what matter most in the end.
The book is designed to mesh in a logical way with a general biology curriculum. The dynamical
themes are distilled from the material covered in standard introductory courses: genetics, cell
biology, physiology, and ecology. When instructors of these courses find themselves freed from
reviewing basic quantitative methods, they can begin to use quantitative reasoning as an integral
part of each course. Students forge the connections that make learning stick when they see ideas
from their math course pay off in biology, and vice versa, or develop the confidence to play with
the numbers with algebraic, graphical, or computer tools.
Most importantly, the course is fun to teach. Leading students through an integrated course
for a full year removes the pressure for instant instructor gratification. (All of my students could
take the derivatives of polynomials.) Instead, one can allow understanding to develop as concepts
return for the second or third time. Students find this unsettling and yearn for instant gratification
too. But with time, they accept the challenge of thinking. When they begin to apply their new
powers to their own problems, when they solve a problem on the computer without being told to,
or when they teach me something about biology in the context of a mathematical idea, delayed
gratification starts to feel like the best possible kind.

Acknowledgments
This book would never have been written without the support of a Hughes Foundation Grant to
the University of Utah which included as part of its mission an attempt to more effectively teach
mathematics to biology majors. That grant brought together a committee of faculty to guide
creation of this book and course consisting of Aaron Fogelson, David Goldenberg, Jim Keener,
Mark Lewis, David Mason, Larry Okun, Hans Othmer, Jon Seger and Ryk Ward. Each, in his
own way, added much to this work. Particular thanks to Jon Seger and Mark Lewis for discussion
and ideas. Frank Wattenberg, Lou Gross, and Simon Levin for looked over the book and delivered
much-needed advice on the whole. Alan Rogers kindly let me use his exercise style and Nelson Beebe
helped smooth over many technical problems. Thanks to my editor Gary Ostedt for agreeing to

3
support a preliminary edition, for providing bird watching opportunities, and for making me feel
important.
This draft of the book benefited greatly from the comments and complaints of the students
who survived the rocky first run of the course: Jennifer Aiman, Ty Captain Flail Corbridge,
Brett Doxey, Ambur Economou, Brad Hasna, Robert Kane, Laura Krause, Jennifer Layman, Eric
Mortensen, Scott Nord, Kevin Rapp, Chris Reilly, Mindi Robinson, Rachael Rosenfeld, Stephanie
Spindler, Mark Stevens, Sheri Williams, Richard Wood and Gentry Yost. Ranging from those
with an excess of comments to those who suffered in eloquent silence, they helped give this book
whatever value it might have as a teaching tool. The veterans of the second run offered the same
range of honest and helpful advice: Elissa Ashby, William Bleazard, Aaron Campbell, Timothy
Christensen, Christina Davenport, Elizabeth Gloyn, Marc Hammerlund, Stacey Hansen, Catherine
Hatt, Christopher Horne, Katina Lessard, Michelle Madsen, Stacy Meola, Jennifer Mercier, Wendy
Pendry, Karin Rattlingourd, Alison Schick, Helene Segal, Sara Sharpsteen, Christen Sowards,
Samuel Webb, and Luann Witt. They corrected many errors and cheerfully pointed out pedagogical
shortcomings. My teaching assistants, Stephen Proulx, Peter Spiro, Vicky Solomon, Colonel Tim
Lewis and Kristina Bogar, were always ready to stand behind me and tell me what I was doing
wrong.
In addition to her extraordinary sartorial advice and culinary support, I thank Anne Collopy
for her inspirational example of writing with clear transitions, extended metaphors and elegant
sentence structure. And for filling the work-free interstices of life with the same.

Contents
I

Introduction to Discrete Dynamical Systems

1 Biology and Dynamics

1.1 Growth: Models of malaria . . .
1.2 Maintenance: Models of neurons
1.3 Replication: Models of genetics .
1.4 Types of dynamical systems . . .

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5
6
7
8
8

2 Updating Functions: Describing Growth

2.1 A model population: bacterial growth . .
2.2 A model organism: a growing tree . . . .
2.3 Functions: terminology and graphs . . . .
2.4 Exercises . . . . . . . . . . . . . . . . . .

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11
11
13
14
18

3 Units and Dimensions

3.1 Converting between units . . . . . .
3.2 Translating between dimensions . . .
3.3 Checking: dimensions and estimation
3.4 Exercises . . . . . . . . . . . . . . .

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21
21
23
26
29

4 Linear Functions and Their Graphs

4.1 Proportional relations . . . . . . . .
4.2 The equation of a line . . . . . . . .
4.3 Finding equations and graphing lines
4.4 Exercises . . . . . . . . . . . . . . .

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31
31
33
36
39

5 Finding Solutions: Describing the Dynamics

5.1 Bacterial population growth . . . . . . . . . .
5.2 Tree height . . . . . . . . . . . . . . . . . . .
5.3 Finding solutions in more complicated cases .
5.4 Exercises . . . . . . . . . . . . . . . . . . . .

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43
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45
46
49

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6 Combining and manipulating functions

53
6.1 Adding functions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
6.2 Composition of functions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
6.3 Inverse functions: looking backwards . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
5

CONTENTS
6.4

Exercises

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

7 Solutions and Exponential Functions

7.1 Bacterial population growth in general
7.2 Laws of exponents and logs . . . . . .
7.3 Expressing results with exponentials .
7.4 Exercises . . . . . . . . . . . . . . . .

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65
65
67
70
74

8 Power Functions and Allometry

8.1 Power relations and exponential growth . .
8.2 Power relations and lines . . . . . . . . . . .
8.3 Power relations in biology: shape and flight
8.4 Exercises . . . . . . . . . . . . . . . . . . .

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77
77
80
83
85

9 Oscillations and Trigonometry

9.1 Sine and cosine: a review . . . . . . .
9.2 Describing oscillations with the cosine
9.3 More complicated shapes . . . . . . .
9.4 Exercises . . . . . . . . . . . . . . . .

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89
89
91
94
96

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10 Modeling and Cobwebbing

10.1 A model of the lungs . . . . . . . . . . . . .
10.2 The lung updating function in general . . .
10.3 Cobwebbing: a graphical solution technique
10.4 Exercises . . . . . . . . . . . . . . . . . . .

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101
. 101
. 104
. 105
. 107

11 Equilibria
11.1 Graphical approach . . . . . .
11.2 Algebraic approach . . . . . .
11.3 Algebra involving parameters
11.4 Exercises . . . . . . . . . . .

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111
111
113
115
117

12 An Example of Nonlinear Dynamics

12.1 A model of selection . . . . . . . . .
12.2 Equilibria in the general case . . . .
12.3 Stable and unstable equilibria . . . .
12.4 Exercises . . . . . . . . . . . . . . .

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121
. 121
. 125
. 127
. 129

13 Excitable Systems I: The Heart

13.1 A simple heart . . . . . . . . .
13.2 Second degree block . . . . . .
13.3 The Wenckebach phenomenon .
13.4 Exercises . . . . . . . . . . . .

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135
135
138
140
141

CONTENTS

II

Limits and Derivatives

153

14 Differential Equations
14.1 Bacterial growth measured continuously
14.2 Rates of change . . . . . . . . . . . . . .
14.3 The limit . . . . . . . . . . . . . . . . .
14.4 Exercises . . . . . . . . . . . . . . . . .

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157
157
161
167
168

15 Slopes of Curves
15.1 The tangent line . . . . . . . . .
15.2 The equation for the tangent line
15.3 Estimating slopes from data . . .
15.4 Exercises . . . . . . . . . . . . .

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173
. 173
. 178
. 180
. 181

16 Stability and the Derivative

16.1 Motivation . . . . . . . . . . .
16.2 Stability and the slope . . . . .
16.3 Estimating slopes at equilibria
16.4 Exercises . . . . . . . . . . . .

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185
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192

17 More Complex Dynamics

17.1 The logistic dynamical system . .
17.2 Qualitative dynamical systems .
17.3 Analysis of the logistic dynamical
17.4 Exercises . . . . . . . . . . . . .

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18 Limits
18.1 Limits of functions
18.2 Properties of limits
18.3 Infinite limits . . .
18.4 Exercises . . . . .

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19 Continuity
19.1 Continuous functions . . . .
19.2 Input and output tolerances
19.3 Hysteresis . . . . . . . . . .
19.4 Exercises . . . . . . . . . .

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219
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. 227

20 Building blocks for derivatives

20.1 Linear functions . . . . . . .
20.2 A quadratic function . . . . .
20.3 The sum rule . . . . . . . . .
20.4 Exercises . . . . . . . . . . .

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231
231
233
235
239

CONTENTS

21 Derivatives of Powers and Polynomials

21.1 Derivatives of basic power functions . . . . . .
21.2 Derivatives of polynomials . . . . . . . . . . . .
21.3 The power rule: negative and fractional powers
21.4 Exercises . . . . . . . . . . . . . . . . . . . . .
22 Derivatives of products and quotients
22.1 The product rule . . . . . . . . . . . .
22.2 Special cases and examples . . . . . .
22.3 The quotient rule . . . . . . . . . . . .
22.4 Exercises . . . . . . . . . . . . . . . .

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23 Derivatives of Exponential and Logarithmic Functions

23.1 The exponential function . . . . . . . . . . . . . . . . . .
23.2 The natural logarithm . . . . . . . . . . . . . . . . . . . .
23.3 Applications . . . . . . . . . . . . . . . . . . . . . . . . . .
23.4 Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . .
24 Derivatives of Trigonometric Functions
24.1 Sine and cosine . . . . . . . . . . . . . .
24.2 Other trigonometric functions . . . . . .
24.3 Applications . . . . . . . . . . . . . . . .
24.4 Exercises . . . . . . . . . . . . . . . . .
25 The
25.1
25.2
25.3
25.4

III

Chain Rule
The derivative of a composite function
Derivatives of inverse functions . . . .
Applications . . . . . . . . . . . . . . .
Exercises . . . . . . . . . . . . . . . .

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257
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. 263

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. 273

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. 284

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287
. 287
. 291
. 293
. 295

Applications of Derivatives and Dynamical Systems

26 Maximization
26.1 Minima and maxima . . . .
26.2 Maximizing food intake rate
26.3 Maximizing fish harvest . .
26.4 Exercises . . . . . . . . . .

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27 Reasoning about functions

27.1 The Intermediate Value Theorem . . . . . . . .
27.2 Maximization: The Extreme Value Theorem . .
27.3 Rolles Theorem and the Mean Value Theorem
27.4 Exercises . . . . . . . . . . . . . . . . . . . . .

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319
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335
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. 338
. 340
. 343

CONTENTS

28 Limits at Infinity
28.1 The behavior of functions at infinity
28.2 Application to absorption functions .
28.3 Limits of sequences . . . . . . . . . .
28.4 Exercises . . . . . . . . . . . . . . .

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29 Leading behavior and LHopitals Rule

29.1 Leading behavior of functions at infinity
29.2 Matched leading behaviors . . . . . . . .
29.3 LHopitals Rule . . . . . . . . . . . . .
29.4 Exercises . . . . . . . . . . . . . . . . .
30 Approximating functions
30.1 The tangent and secant lines
30.2 Quadratic approximation . .
30.3 Taylor polynomials . . . . . .
30.4 Exercises . . . . . . . . . . .
31 Newtons method
31.1 Finding equilibria . . .
31.2 Newtons method . . .
31.3 Why Newtons method
31.4 Exercises . . . . . . .

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works and when it
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32 Panting and Deep Breathing

32.1 Breathing at different rates
32.2 Deep breathing . . . . . . .
32.3 Panting . . . . . . . . . . .
32.4 Exercises . . . . . . . . . .

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397
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. 398
. 399
. 401

Differential Equations, Integrals, and Their Applications

33 Differential Equations
33.1 Differential equations: examples and terminology
33.2 Eulers method: pure-time . . . . . . . . . . . . .
33.3 Eulers method: autonomous . . . . . . . . . . .
33.4 Exercises . . . . . . . . . . . . . . . . . . . . . .
34 Basic differential equations
34.1 Newtons Law of Cooling . . . . . .
34.2 Diffusion across a membrane . . . .
34.3 A continuous time model of selection
34.4 Exercises . . . . . . . . . . . . . . .

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431
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. 438

10

CONTENTS

35 The
35.1
35.2
35.3
35.4

Antiderivative
Pure-time differential equations . . . . .
Rules for antiderivatives . . . . . . . . .
Solving polynomial differential equations
Exercises . . . . . . . . . . . . . . . . .

36 Special functions and substitution

36.1 Integrals of special functions . . . .
36.2 The chain rule and integration . .
36.3 Getting rid of excess constants . .
36.4 Exercises . . . . . . . . . . . . . .
37 Integrals and sums
37.1 Approximating integrals with sums
37.2 Approximating integrals in general
37.3 The definite integral . . . . . . . .
37.4 Exercises . . . . . . . . . . . . . .

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38 Definite and indefinite integrals

38.1 The Fundamental Theorem of Calculus . . . .
38.2 The summation property of definite integrals
38.3 General solution . . . . . . . . . . . . . . . .
38.4 Exercises . . . . . . . . . . . . . . . . . . . .
39 Applications of integrals
39.1 Integrals and areas . .
39.2 Integrals and averages
39.3 Integrals and mass . .
39.4 Exercises . . . . . . .

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40 Improper integrals
40.1 Infinite limits of integration .
40.2 Improper integrals: examples
40.3 Infinite integrands . . . . . .
40.4 Exercises . . . . . . . . . . .

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441
. 441
. 442
. 445
. 448

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453
. 453
. 454
. 457
. 461

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465
. 465
. 467
. 471
. 472

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475
. 475
. 480
. 482
. 484

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. 487
. 491
. 493
. 495

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499
. 499
. 500
. 503
. 506

Analysis of Differential Equations

39 Autonomous Differential Equations

39.1 Review of autonomous differential equations
39.2 Equilibria . . . . . . . . . . . . . . . . . . .
39.3 Display of differential equations . . . . . . .
39.4 Exercises . . . . . . . . . . . . . . . . . . .

475

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479
. 479
. 481
. 483
. 486

CONTENTS

11

40 Stable and unstable equilibria

40.1 Recognizing stable and unstable equilibria
40.2 Applications of the stability theorem . . .
40.3 A model of a disease . . . . . . . . . . . .
40.4 Exercises . . . . . . . . . . . . . . . . . .
41 Solving autonomous equations
41.1 Separation of variables . . . . . . . . .
41.2 Pure-time equations revisited . . . . .
41.3 Applications of separation of variables
41.4 Exercises . . . . . . . . . . . . . . . .
42 Two dimensional equations
42.1 Predator-prey dynamics .
42.2 Newtons law of cooling .
42.3 Eulers method . . . . . .
42.4 Exercises . . . . . . . . .
43 The
43.1
43.2
43.3
43.4

Phase-Plane
Equilibria and nullclines:
Equilibria and nullclines:
Equilibria and nullclines:
Exercises . . . . . . . .

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predator-prey equations .
selection equations . . .
Newtons law of cooling .
. . . . . . . . . . . . . . .

44 Solutions in the phase-plane

44.1 Eulers method in the phase-plane . . . .
44.2 Direction arrows: predator-prey equations
44.3 More direction arrows . . . . . . . . . . .
44.4 Exercises . . . . . . . . . . . . . . . . . .
45 The
45.1
45.2
45.3
45.4

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dynamics of a neuron
A mathematicians view of a neuron
The mathematics of sodium channels
The FitzHugh-Nagumo equations . .
Exercises . . . . . . . . . . . . . . .

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491
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. 499

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. 506

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511
. 511
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. 517

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521
. 521
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. 528
. 530

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545
. 545
. 547
. 549
. 553

Probability Theory and Descriptive Statistics

46 Probabilistic Models
46.1 Probability and statistics . .
46.2 Stochastic population growth
46.3 Markov chains . . . . . . . .
46.4 Exercises . . . . . . . . . . .

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569
. 569
. 570
. 573
. 575

12

CONTENTS

47 Stochastic models of diffusion

47.1 Stochastic diffusion . . . . . .
47.2 Stochastic diffusion . . . . . .
47.3 Stochastic diffusion . . . . . .
47.4 Exercises . . . . . . . . . . .

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577
577
579
581
584

48 Stochastic models of genetics

48.1 The genetics of inbreeding .
48.2 The dynamics of height . .
48.3 Blending inheritance . . . .
48.4 Exercises . . . . . . . . . .

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587
587
589
592
593

49 Probability Theory
49.1 Sample spaces and events . . . .
49.2 Set theory . . . . . . . . . . . . .
49.3 Assigning probabilities to events
49.4 Exercises . . . . . . . . . . . . .

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595
595
596
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603
603
606
607
609

51 Independence and Markov Chains

51.1 Independence . . . . . . . . . . . . . . . . . . .
51.2 The multiplication rule for independent events
51.3 Markov chains and conditional probability . . .
51.4 Exercises . . . . . . . . . . . . . . . . . . . . .

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611
611
612
614
616

52 Displaying Probabilities
52.1 Probability and cumulative distributions
52.2 The probability density function . . . .
52.3 The cumulative distribution function . .
52.4 Exercises . . . . . . . . . . . . . . . . .

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619
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621
626
628

53 Random Variables
53.1 Types of random variable . .
53.2 Expectation: discrete case . .
53.3 Expectation: continuous case
53.4 Exercises . . . . . . . . . . .

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631
631
635
637
639

50 Conditional Probability
50.1 Conditional probability . .
50.2 The law of total probability
50.3 Bayes theorem and the rare
50.4 Exercises . . . . . . . . . .

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54 Descriptive Statistics
643
54.1 The median . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 643
54.2 The mode . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 647
54.3 The geometric mean . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 648

CONTENTS
54.4 Exercises

13
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 653

55 Descriptive statistics for spread

55.1 Range and percentiles . . . . .
55.2 Mean absolution deviation . . .
55.3 Variance . . . . . . . . . . . . .
55.4 Exercises . . . . . . . . . . . .

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655
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. 657
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. 663

Probability Models

677

56 Joint distributions
56.1 Joint distributions . . . . . . . . . . . . . . . . .
56.2 Marginal probability distributions . . . . . . . .
56.3 Joint distributions and conditional distributions .
56.4 Exercises . . . . . . . . . . . . . . . . . . . . . .
57 Covariance and Correlation
57.1 Covariance . . . . . . . .
57.2 Correlation . . . . . . . .
57.3 Perfect correlation . . . .
57.4 Exercises . . . . . . . . .

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13 Matrices and Markov Chains

13.1 Matrices and conditional distributions
13.2 Equilibria of Markov chains . . . . . .
13.3 Generalized Markov chains . . . . . .
13.4 Exercises . . . . . . . . . . . . . . . .

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681
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. 686
. 690

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693
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. 698
. 701

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175
. 175
. 177
. 179
. 181

58 Sums and products of random variables

58.1 Expectation of a sum . . . . . . . . . . .
58.2 Expectation of a product . . . . . . . .
58.3 Variance of a sum . . . . . . . . . . . .
58.4 Exercises . . . . . . . . . . . . . . . . .

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703
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. 706
. 709
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59 The
59.1
59.2
59.3
59.4

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713
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715
718
723

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distribution
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725
725
728
730
732

Binomial Distribution
The binomial distribution defined . . .
Computing the binomial . . . . . . . .
Binomial distribution: the general case
Exercises . . . . . . . . . . . . . . . .

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60 Applications of the Binomial Distribution

60.1 Application to genetics and calculation of mode . . . .
60.2 Application to Markov chains: definition of cumulative
60.3 Applications to diffusion . . . . . . . . . . . . . . . . .
60.4 Exercises . . . . . . . . . . . . . . . . . . . . . . . . .

14

CONTENTS

61 Exponential distributions
61.1 The geometric distribution . .
61.2 The exponential distribution
61.3 The memoryless property . .
61.4 Exercises . . . . . . . . . . .

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62 The
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62.2
62.3
62.4

Poisson Distribution
The Poisson process . . . . . . .
The Poisson distribution in space
The Poisson and the binomial . .
Exercises . . . . . . . . . . . . .

63 The
63.1
63.2
63.3
63.4

Normal Distribution
The normal distribution: an example . . .
The Central Limit Theorem for Sums . .
The Central Limit Theorem for Averages
Exercises . . . . . . . . . . . . . . . . . .

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64 Applying the Normal Approximation

64.1 The standard normal distribution . . .
64.2 Normal approximation of the binomial
64.3 Normal approximation of the Poisson .
64.4 Exercises . . . . . . . . . . . . . . . .

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735
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769
. 769
. 775
. 777
. 779

Introduction to Statistical Reasoning

65 Statistics: Estimating Parameters

65.1 Estimating the binomial proportion .
65.2 Maximum likelihood . . . . . . . . .
65.3 Estimating a rate . . . . . . . . . . .
65.4 Exercises . . . . . . . . . . . . . . .
66 Confidence limits
66.1 Exact confidence limits .
66.2 Monte Carlo method . .
66.3 Likelihood, support, and
66.4 Exercises . . . . . . . .

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. . . . . . . . . .
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confidence limits
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67 Estimating the Mean

67.1 Estimating the mean . . . . .
67.2 Confidence limits . . . . . . .
67.3 Sample variance and standard
67.4 Exercises . . . . . . . . . . .

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797
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819
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. 821
. 825
. 828

CONTENTS

15

68 Hypothesis Testing
68.1 Hypothesis testing: an example . . . .
68.2 Power and confidence limits . . . . . .
68.3 Likelihood and the method of support
68.4 Exercises . . . . . . . . . . . . . . . .

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831
831
835
836
838

69 Hypothesis Testing: Normal Theory

69.1 Computing p-values with the normal approximation
69.2 The power of normal tests . . . . . . . . . . . . . . .
69.3 Likelihood and the normal distribution . . . . . . . .
69.4 Exercises . . . . . . . . . . . . . . . . . . . . . . . .

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841
841
844
847
849

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70 Comparing experiments
70.1 Unpaired normal distributions . . .
70.2 Comparing population proportions
70.3 Likelihood . . . . . . . . . . . . . .
70.4 Exercises . . . . . . . . . . . . . .

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851
. 851
. 853
. 855
. 858

71 Regression
71.1 Linear regression . . . . . . . .
71.2 Using linear regression . . . . .
71.3 The theory of linear regression
71.4 Exercises . . . . . . . . . . . .

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72 Answers

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861
861
865
867
870
883