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 Seventh Edition

ROBERT J. BROOKER
University of Minnesota
viii PREFACE
aware of the Topic at hand. Second, the question is evaluated
with regard to the Information that is available to the student.
Finally, the student is guided through one or more Problem-
Solving Strategies to tackle the question.
∙∙ Genes → Traits: Because genetics is such a broad discipline,
ranging from the molecular level to populations, many
instructors have told us that it is a challenge for students to see
both “the forest and the trees.” It is commonly mentioned that
students often have trouble connecting the concepts they have
learned in molecular genetics with the traits that occur at the
level of a whole organism (e.g., What does transcription have
to do with blue eyes?). To try to make this connection more
meaningful, certain figure legends in each chapter, designated
Genes → Traits, remind students that molecular and cellular
phenomena ultimately lead to the traits that are observed in
each species (see Figure 14.8).
∙∙ Learning Outcomes: Each section of every chapter begins
with a set of learning outcomes. These outcomes help students
understand what they should be able to do once they have
mastered the material in that section.
∙∙ Formative Assessment: When students are expected to learn
textbook material on their own, it is imperative that they are
regularly given formative assessment so they can gauge
whether they are mastering the material. Formative assessment
is a major feature of this textbook and is bolstered by
Connect—a state-of-the art digital assignment and assessment
platform. In Genetics: Analysis & Principles, seventh edition,
formative assessment is provided in multiple ways.
1. As mentioned, a feature called Genetic TIPS is aimed at
helping students refine their problem-solving skills.
2. Each section of every chapter ends with multiple-choice
Comprehension Questions. Also, compared with previous
editions, many chapters in the seventh edition are divided
into more sections that are shorter in length. Formative
assessment at the end of each section allows students to
evaluate their mastery of the material before moving on to
the next section. The answers are provided in Appendix B
at the end of the book.
3. Most figures have Concept Check questions so that
students can determine if they understand the key points in
the figure. The answers are provided in Appendix B at the
end of the book.
4. Extensive end-of chapter questions continue to provide
students with feedback regarding their mastery of the
material. The answers to even-numbered questions are
provided in Appendix B.
5. The textbook material is supported by digital learning
tools found in Connect. Instructors can assign questions
and activities in Connect, and students also have access
to our valuable adaptive study tool, SmartBook 2.0.
With this tool, students are repeatedly given questions
regarding the textbook material, and depending on their
answers, they may advance in their reading, or they are
given specific advice on what textbook material to go
back and review.
Overall, the pedagogy of Genetics: Analysis & Principles,
seventh edition, has been designed to foster student learning. In-
stead of being a collection of facts and figures, Genetics: Analysis
& Principles, seventh edition, by Rob Brooker, is intended to be
an engaging and motivating textbook in which formative assess-
ment allows students to move ahead and learn the material in a
productive way. We welcome your feedback so that we can make
future editions even better!
MAJOR CONTENT CHANGES
IN RECENT EDITIONS
∙∙ Coverage of the topic of epigenetics was expanded to a whole
chapter in the sixth edition, which is Chapter 16. In the seventh
edition, this chapter includes a new section that focuses on the
structure and function of heterochromatin.
∙∙ A chapter on non-coding RNAs, Chapter 17, was added in the
sixth edition. This long-overdue chapter was included in
response to a remarkable explosion in our appreciation for the
roles of non-coding RNAs in many aspects of molecular
biology. Although two new chapters were added in recent
editions, the overall page length of the seventh edition is not
longer than the fifth edition.
∙∙ Discussion of the role of the CRISPR-Cas system in providing
prokaryotes with a genome defense mechanism has been
added in Chapter 17, and its use by researchers to edit genes is
described in Chapter 21.
∙∙ Due to the expanding material in the area of medical
genetics and cancer biology, the chapter on medical genetics
and cancer has been split into two chapters in the seventh
edition.
∙∙ The material that was covered in Chapter 20 of the sixth
edition has been placed in other chapters. Homologous
recombination is now in Chapter 19 and transposition is in
Chapter 10.
Examples of Specific Content Changes to
Individual Chapters in the Seventh Edition
∙∙ Chapter 2, Mendelian Inheritance: The discussion of the chi
square test has been revised to better explain the connection to
the null hypothesis.
∙∙ Chapter 4, Extensions of Mendelian Inheritance: Ever wonder
why some breeds of dogs, such as collies, have white
undersides with darker fur on their backs? A new subsection
explains how this phenomenon is related to events during
development that involve the migration and proliferation of
melanocytes (see Figure 4.16).
∙∙ Chapter 6, Genetic and Linkage Mapping in Eukaryotes: The
section on genetic mapping has been revised to emphasize that
the pattern of allele linkage is deduced from the true-breeding
P generation and that F2 recombinants are produced by a
crossover.

∙∙ Chapter 7, Genetic Transfer and Mapping in Bacteria: The
section on bacterial transduction has been simplified by
eliminating the discussion of cotransduction, which is not
commonly used.
∙∙ Chapter 9, Molecular Structure of DNA and RNA: Based on
reviewer feedback, the topic of triplex DNA was removed from
this chapter.
∙∙ Chapter 10, Molecular Structure of Chromosomes and
Transposable Elements: Due to their effects on chromosome
structure, transposable elements (discussed in Chapter 20 in the
sixth edition) are now covered in this chapter (see Section 10.5).
New and revised figures regarding eukaryotic chromatin
structure were added (see Figures 10.19, 10.22, and 10.23).
∙∙ Chapter 11, DNA Replication: A new model depicting the
molecular structure of DNA polymerase has been added (see
Figure 11.8b).
∙∙ Chapter 12, Gene Transcription and RNA Modification: New
information describing how a component of the spliceosome
acts as a ribozyme has been added.
∙∙ Chapter 14, Gene Regulation in Bacteria: The discussion of
the trp operon has been simplified.
∙∙ Chapter 15, Gene Regulation in Eukaryotes I: Transcriptional
and Translational Regulation: The topic of insulators has been
moved to Chapter 16. They are now called “barriers” to
heterochromatin formation. Discussion of the potential role of
nucleosome-free regions in preventing heterochromatin
spreading has been added.
∙∙ Chapter 16, Gene Regulation in Eukaryotes II: Epigenetics:
A new section entitled “Heterochromatin: Formation, Structure,
Formation, and Maintenance” has been added. This section has
five new figures (see Figures 16.4 through 16.8). The section
entitled “Role of Epigenetics in Cancer” was moved to
Chapter 25.
∙∙ Chapter 17, Non-coding RNAs: New figures illustrate non-
coding RNAs that act as a guide and a decoy (see Figures 17.2
and 17.3). Figure 17.10 was revised to distinguish between
pRITS and pRISC, which silence the transcription and
translation of transposable elements, respectively.
∙∙ Chapter 18, Genetics of Viruses: The information on the origin
of HIV and the occurrence of HIV infection worldwide and in
the United States has been updated.
∙∙ Chapter 19, Gene Mutation, DNA Repair, and Recombination:
Figure 19.18, which concerns nucleotide excision repair, has
been revised to include the function of UvrD. The section on
homologous recombination (in Chapter 20 in the sixth edition)
was moved to this chapter.
∙∙ Chapter 20, Molecular Technologies: This chapter was Chapter
21 in the sixth edition. The topic of real-time PCR has been
divided into smaller subsections for clarity and ease of reading.
Section 20.4, formerly called “Gene Mutagenesis” is now
called “Gene Editing.”
PREFACE ix
∙∙ Chapter 21, Biotechnology: This chapter was Chapter 22
in the sixth edition. The section entitled “Human Gene
Therapy” was moved to the chapter on medical genetics
(now Chapter 24).
∙∙ Chapter 22, Genomics I: Analysis of DNA: This chapter was
Chapter 23 in the sixth edition. The section on physical
mapping has been simplified.
∙∙ Chapter 23, Genomics II: Functional Genomics, Proteomics,
and Bioinformatics: This chapter was Chapter 24 in the sixth
edition.
∙∙ Chapter 24, Medical Genetics: This chapter was Chapter 25
in the sixth edition. The topic of genome-wide association
(GWA) studies was added (see Figure 24.8). The section
“Human Gene Therapy” was moved to this chapter. The
material on cancer was moved to Chapter 25, which solely
focuses on cancer.
∙∙ Chapter 25, Genetic Basis of Cancer: This topic is now
featured in its own chapter, which includes four separate
sections. The last section focuses on the role of epigenetics
in cancer.
∙∙ Chapter 26, Developmental Genetics: The information on Hox
genes in development and on the role of the SRY gene in
human sex determination has been updated.
∙∙ Chapter 27, Population Genetics: Discussion of the topic of
inbreeding has been expanded.
∙∙ Chapter 28, Complex and Quantitative Traits: The
identification of QTLs is now covered in a separate
section.
∙∙ Chapter 29, Evolutionary Genetics: The cladistics method for
constructing a phylogenetic tree is compared with the UPGMA
method.
Suggestions Welcome!
It seems very appropriate to use the word evolution to describe the
continued development of this textbook. I welcome any and all
comments. The refinement of any science textbook requires input
from instructors and their students. These include comments re-
garding writing, illustrations, supplements, factual content, and
topics that may need greater or less emphasis. You are invited to
contact me at:
Dr. Rob Brooker
Dept. of Genetics, Cell Biology, and Development
Dept. of Biology Teaching and Learning
University of Minnesota
6-160 Jackson Hall
321 Church St.
Minneapolis, MN 55455
brook005@umn.edu
612-624-3053
 ®

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xii PREFACE

overseeing this project. I would like to thank other people at

ACKNOWLEDGMENTS McGraw-Hill who have played key roles in producing the actual
book and the supplements that go along with it. In particular,
The production of a textbook is truly a collaborative effort, and I
Jessica Portz (Senior Content Project Manager) has done a superb
am deeply indebted to many people. All seven editions of this
job of managing the components that need to be assembled to
textbook went through multiple rounds of rigorous revision that
produce a book. I would also like to thank Lori Hancock (Content
involved the input of faculty, students, editors, and educational
Licensing Specialist), who acted as an interface between me and
and media specialists. Their collective contributions are reflected
the photo company. In addition, my gratitude goes to David Hash
in the final outcome.
(Designer), who provided much input into the internal design of
Deborah Brooker (Freelance Developmental Editor) me-
the book and created an awesome cover. Finally, I would like to
ticulously read the new material, analyzed every figure, and of-
thank Kelly Brown (Executive Marketing Manager), whose major
fered extensive feedback. Her attention to detail in this edition
efforts begin when the seventh edition comes out!
and previous editions has profoundly contributed to the accuracy
I would also like to extend my thanks to everyone at
and clarity of this textbook. I would also like to thank Jane
MPS Limited who worked with great care in the paging of the
Hoover (Freelance Copy Editor) for understanding the material
book, making sure that the figures and relevant text are as close to
and working extremely hard to improve the text’s clarity. Her ef-
each other as possible. Likewise, the people at Photo Affairs, Inc.
forts are truly appreciated. She is at the top of the list for copy
have done a great job of locating many of the photographs that
editors.
have been used in the seventh edition.
I would also like to acknowledge the many people at
Finally, I want to thank the many scientists who reviewed
McGraw-Hill Education whose skills and insights are amazing.
the chapters of this textbook. Their broad insights and construc-
My highest praise goes to Elizabeth Sievers (Senior Product De-
tive suggestions were an overriding factor that shaped its final
veloper), who carefully checks all aspects of textbook develop-
content and organization. I am truly grateful for their time and
ment and makes sure that all of the pieces of the puzzle are in
compassion.
place. I am also grateful to Andrew Urban (Portfolio Manager) for

REVIEWERS Deborah Duricka, University of Alaska Willian James MacKay, Edinboro

Amy L. Abdulovic-Cui, Augusta
University
Bruce Bejcek, Western Michigan University
Dena Berg, Tarrant County College NW
Chi-hua Chiu, Kent State University
Norman W. Clippinger, Metropolitan State
University of Denver
Brian J. Dingmann, University of
Minnesota Crookston
Anchorage
Kurt J. Elliott, Northwest Vista College
Rajendra Goswami, Harold Washington
College
Vanessa Hörmann, Broward College
Vladimir Jurukovski, Suffolk County
Community College
Galyna Kufryk, Grand Canyon University
Dr. Hsiu-Ping Liu, Metropolitan State
University of Denver
University
Helen Piontkivska, Kent State University
Anthony Schroeder, University of
Minnesota Crookston
Sabeeha Y. Shariff, College of DuPage
Michael A. Thornton, Florida A&M
University
Nina N. Thumser, Edinboro University
Muatasem Ubeidat, Southwestern
Oklahoma State University
 PA RT I INTRODUCTION

CHAPTER OUTLINE
1.1 The Molecular Expression
of Genes
1.2 The Relationship Between
Genes and Traits
1.3 Fields of Genetics
1.4 The Science of Genetics

1
CC ( for “carbon copy”
or “copy cat”), the first
cloned pet. In 2002, the
cat shown here was
produced by reproductive
cloning, a procedure
described in Chapter 21.
©FEMA/Texas A&M
University/Getty Images

OVERVIEW OF GENETICS
Hardly a week goes by without a major news story announcing a
genetic breakthrough. The increasing pace of genetic discoveries
has become staggering. The Human Genome Project is a case in
point. This project began in the United States in 1990, when the
National Institutes of Health and the Department of Energy joined
forces with international partners to decipher the massive amount of
information contained in our genome—the DNA found within all
of our chromosomes (Figure 1.1). Remarkably, in only a decade,
the researchers working on this project determined the DNA
sequence (the order of the bases A, T, G, and C) of over 90% of the
human genome. The completed sequence, published in 2003, has an
accuracy greater than 99.99%; less than 1 mistake was made in
every 10,000 base pairs!
In 2008, a more massive undertaking, called the 1000 Ge-
nomes Project, was launched to establish a detailed understand-
ing of human genetic variation. In this international project,
researchers set out to determine the DNA sequence of at least
1000 anonymous participants from around the globe. In 2015,
the sequencing of over 2500 genomes was described in the jour-
nal Nature.
Studying the human genome allows us to explore funda-
mental details about ourselves at the molecular level. The results
of human genome projects have shed considerable light on basic
questions, like how many genes we have, how genes direct the
activities of living cells, how species evolve, how single cells
develop into complex tissues, and how defective genes cause
disease. Furthermore, understanding our genome may lend itself
to improvements in modern medicine by leading to better diag-
noses of diseases and allowing the development of new treat-
ments for them.
The quest to unravel the mysteries within our genes has
involved the invention of many new technologies. For example,
researchers have developed genetic techniques to produce medi-
cines, such as human insulin, that would otherwise be difficult or
impossible to make. Human insulin is synthesized in strains of
Escherichia coli bacteria that have been genetically altered by the
addition of genes that encode the polypeptides that form this hor-
mone. Grown in laboratories, these bacteria make large amounts of
human insulin. As discussed in Chapter 21, the insulin is purified
and administered to many people with insulin-dependent diabetes.

1
2 C H A P T E R 1 :: OVERVIEW OF GENETICS

Chromosomes
DNA, the molecule of life
Cell
The adult human body
is composed of trillions
of cells.

Most human cells contain

the following:
Gene
• 46 human chromosomes,
found in 23 pairs

C G

T A
T A
• 2 meters of DNA

G
C G

T A

T A
• Approximately 22,000

T A
genes coding for

A T

C G
proteins that perform

A T
most life functions
DNA
• Approximately 3 billion
DNA base pairs per set mRNA
of chromosomes,
containing the bases A,
T, G, and C

Amino acid

Protein (composed of amino acids)

FI G U RE 1.1 The human genome. The human genome is a complete set of human chromosomes. People have two sets of chromosomes—one
set from each parent—which are found in the cell nucleus. The Human Genome Project revealed that each set of chromosomes is composed of a DNA
sequence that is approximately 3 billion base pairs long. As discussed later, most genes are first transcribed into mRNA and then the mRNA is used to
make proteins. Estimates suggest that each set of chromosomes contains about 22,000 different protein-encoding genes. This figure emphasizes the
DNA found in the cell nucleus. Humans also have a small amount of DNA in their mitochondria, which has also been sequenced.
CONCEPT CHECK: How might a better understanding of our genes be used in the field of medicine?

New genetic technologies are often met with skepticism and some-
times even with disdain. An example is mammalian cloning. In
1997, Ian Wilmut and his colleagues created clones of sheep,
using mammary cells from an adult animal (Figure 1.2). More
recently, such cloning has been achieved in several mammalian
species, including cows, mice, goats, pigs, and cats. In 2002, the
first pet was cloned, a cat named CC (for “carbon copy” or “copy
cat”; see the chapter-opening photo). The cloning of mammals
provides the potential for many practical applications. With regard
to livestock, cloning would enable farmers to use cells from their
best individuals to create genetically homogeneous herds. This
could be advantageous in terms of agricultural yield, although
such a genetically homogeneous herd may be more susceptible to
certain diseases. However, people have become greatly concerned
about the possibility of human cloning. This prospect has raised
serious ethical questions. Within the past few years, legislation F I G URE 1 . 2 The cloning of a mammal. The lamb in the front
that involves bans on human cloning has been introduced. is Dolly, the first mammal to be cloned. She was cloned from the cells of
a Finn Dorset (a white-faced sheep). The sheep in the back is Dolly’s
Finally, genetic technologies provide the means to modify
surrogate mother, a Blackface ewe. A description of how Dolly was
the traits of animals and plants in ways that would have been
produced is presented in Chapter 21.
unimaginable just a few decades ago. Figure 1.3a shows a strik- ©R. Scott Horner KRT/Newscom
ing example in which scientists introduced a gene from jellyfish
CONCEPT CHECK: What ethical issues may be associated with human cloning?
into mice. Certain species of jellyfish emit a “green glow”

(a) GFP expressed in mice
GFP
(b) GFP expressed in the gonads of a male mosquito
FI GURE 1.3 The introduction of a jellyfish gene into
laboratory mice and mosquitoes. (a) A gene that naturally occurs in
jellyfish encodes a protein called green fluorescent protein (GFP). The
GFP gene was cloned and introduced into mice. When these mice are
exposed to UV light, GFP emits a bright green color. These mice glow
green, just like the jellyfish! (b) The GFP gene was introduced next to a
gene sequence that causes the expression of GFP only in the gonads of
male mosquitoes. The resulting green glow allows researchers to identify
and sort males from females.
(a): ©Eye of Science/Science Source; (b): Photo taken by Flaminia Catteruccia, Jason Benton
and Andrea Crisanti, and assembled by www.luciariccidesign.com
CONCEPT CHECK: Why is it useful to sort male mosquitoes from females?
produced by a bioluminescent protein called green fluorescent
protein (GFP) encoded by a gene in the jellyfish genome. When
exposed to blue or ultraviolet (UV) light, the protein emits a strik-
ing green-colored light. Scientists were able to clone the GFP
gene from a sample of jellyfish cells and then introduce this gene
into laboratory mice. The green fluorescent protein is made
throughout the cells of their bodies. As a result, their skin, eyes,
and organs give off an eerie green glow when exposed to UV
light. Only their fur does not glow.
The expression of green fluorescent protein allows research-
ers to identify particular proteins in cells or specific body parts.
1.1 The Molecular Expression of Genes 3
For example, Andrea Crisanti and colleagues have altered mosqui-
toes to express GFP only in the gonads of males (Figure 1.3b).
This enables the researchers to distinguish males from females and
sort mosquitoes by sex. Why is this useful? Researchers can pro-
duce a population of mosquitoes and then sterilize the males. The
ability to distinguish males from females makes it possible to
release the sterile males without the risk of releasing additional
females. The release of sterile males may be an effective means of
controlling mosquito populations because females mate only once
before they die. Mating with a sterile male prevents a female from
producing offspring. In 2008, Osamu Shimomura, Martin Chalfie,
and Roger Tsien received the Nobel Prize in chemistry for the
discovery and development of GFP, which has become a widely
used tool in biology.
Overall, as we move forward in the twenty-first century, the
excitement level in the field of genetics is high, perhaps higher
than it has ever been. Nevertheless, new genetic knowledge and
technologies will create many ethical and societal challenges. In
this chapter, we begin with an overview of genetics and then
explore the various fields of genetics and their experimental
approaches.
1.1 THE MOLECULAR
EXPRESSION OF GENES
Learning Outcomes:
1. Describe the biochemical composition of cells.
2. Explain how proteins are largely responsible for cell struc-
ture and function.
3. Outline how DNA stores the information to make proteins.
Genetics is the branch of biology that deals with heredity and
variation. It stands as the unifying discipline in biology by
allowing us to understand how life can exist at all levels of com-
plexity, ranging from the molecular to the population level.
Genetic variation is the root of the natural diversity that we
observe among members of the same species and among differ-
ent species.
Genetics is centered on the study of genes. A gene is clas-
sically defined as a unit of heredity. At the molecular level, a
gene is a segment of DNA that contains the information to pro-
duce a functional product. The functional product of most genes
is a polypeptide, which is a linear sequence of amino acids that
folds into one of the units that constitute proteins. In addition,
genes are commonly described according to the way they affect
traits, which are the characteristics of an organism. In humans,
for example, we observe traits such as eye color, hair texture,
and height. The ongoing theme of this textbook is the relation-
ship between genes and traits. As an organism grows and devel-
ops, its collection of genes provides a blueprint that determines
its traits.
In this section, we examine the general features of life,
beginning with the molecular level and ending with popula-
tions of organisms. As will become apparent, genetics is the
4 C H A P T E R 1 :: OVERVIEW OF GENETICS
Plant cell

common thread that explains the existence of life and its conti-
nuity from generation to generation. For most students, this
chapter should serve as an overview of topics they have learned
in other introductory courses such as general biology. Even so,
it is usually helpful to see the “big picture” of genetics before
delving into the finer details that are covered in Chapters 2
through 29.
Nucleus

Living Cells Are Composed of Biochemicals

To fully understand the relationship between genes and traits, we
need to begin with an examination of the composition of living
organisms. Every cell is constructed from intricately organized
chemical substances. Small organic molecules such as glucose and
amino acids are produced by the linkage of atoms via chemical
bonds. The chemical properties of organic molecules are essential
for cell vitality in two key ways. First, the breaking of chemical
bonds during the degradation of small molecules provides energy Chromosome
to drive cellular processes. A second important function of these
small organic molecules is their role as the building blocks for the Proteins
synthesis of larger molecules. Four important categories of larger
molecules are nucleic acids (i.e., DNA and RNA), proteins,
carbohydrates, and lipids. Three of these—nucleic acids, pro-
teins, and carbohydrates—form macromolecules that are com-
posed of many repeating units of smaller building blocks. RNA,
proteins, and some carbohydrates are made from hundreds or even DNA
thousands of repeating building blocks. DNA is the largest macro-
molecule found in living cells. A single DNA molecule can be
composed of a linear sequence of hundreds of millions of building
blocks called nucleotides!
The formation of cellular structures relies on the interac-
tions of molecules and macromolecules. For example, nucleo-
tides are connected together to make DNA, which is a constituent
of chromosomes (Figure 1.4). In addition, DNA is associated
with many proteins that provide organization to the structure of
chromosomes. Within a eukaryotic cell, the chromosomes are
contained in a compartment called the cell nucleus. The nucleus Nucleotides
is bounded by a double membrane composed of lipids and pro-
teins that shields the chromosomes from the rest of the cell. The
NH2
organization of chromosomes within a cell nucleus protects the Cytosine H Guanine
chromosomes from mechanical damage and provides a single N
O
compartment for genetic activities such as gene transcription. As O– O N H H
N
N

a general theme, the formation of large cellular structures arises O P O CH2

O O–
H
O– H2 N N
from interactions among different molecules and macromole- H
H H
H O P O CH2
N
O
cules. These cellular structures, in turn, are organized to make a OH H
O–
H H
complete living cell. H H

OH H

Each Cell Contains Many Different
Proteins That Determine Cell Structure
and Function
To a great extent, the characteristics of a cell depend on the types
of proteins that it makes. The entire collection of proteins that a cell
makes at a given time is called its proteome. The range of func-
tions among different types of proteins is truly remarkable. Some
FIGURE 1.4 Molecular organization of a living cell. Cellular
structures are constructed from smaller building blocks. In this example,
DNA is formed from the linkage of nucleotides, producing a very long
macromolecule. The DNA associates with proteins to form a chromosome.
The chromosomes are located within a membrane-bound organelle called
the nucleus, which, along with many other different types of organelles, is
found within a complete cell.
(inset) ©Biophoto Associates/Science Source

proteins help determine the shape and structure of a given cell. CONCEPT CHECK: Is DNA a small molecule, a macromolecule, or an organelle?
 1.1 The Molecular Expression of Genes 5

For example, the protein known as tubulin assembles into large

structures known as microtubules, which provide the cell with
internal structure and organization. Other proteins are inserted into
cell membranes and aid in the transport of ions and small mole-
cules across the membrane. Enzymes, which accelerate chemical
reactions, are a particularly important category of proteins. Some
enzymes play a role in the breakdown of molecules or macromol-
ecules into smaller units. Known as catabolic enzymes, these are
important in the utilization of energy. Alternatively, anabolic
enzymes and accessory proteins function in the synthesis of
molecules and macromolecules throughout the cell. The construc-
tion of a cell greatly depends on its proteins that are involved in
anabolism because these are required to synthesize all cellular
macromolecules.
Molecular biologists have come to realize that the functions
of proteins underlie the cellular characteristics of every organism.
At the molecular level, proteins can be viewed as the active partic-
ipants in the enterprise of life.

F I G URE 1 . 5 A micrograph of the 46 chromosomes found in a

DNA Stores the Information for Protein Synthesis
The genetic material of living organisms is composed of a sub-
stance called deoxyribonucleic acid, abbreviated DNA. The
DNA stores the information needed for the synthesis of all cellu-
lar proteins. In other words, the main function of the genetic blue-
print is to code for the production of proteins in the correct cell,
at the proper time, and in suitable amounts. This is an extremely
complicated task because living cells make thousands of different
proteins. Genetic analyses have shown that a typical bacterium
can make a few thousand different proteins, and estimates for the
numbers produced by complex eukaryotic species range in the
tens of thousands.
DNA’s ability to store information is based on its structure.
DNA is composed of a linear sequence of nucleotides. Each nucle-
otide contains one of four nitrogen-containing bases: adenine (A),
thymine (T), guanine (G), or cytosine (C). The linear order of these
bases along a DNA molecule contains information similar to the
way that groups of letters of the alphabet represent words. For exam-
ple, the “meaning” of the sequence of bases ATGGGCCTTAGC
differs from that of the sequence TTTAAGCTTGCC. DNA
sequences within most genes contain the information to direct the
order of amino acids within polypeptides according to the genetic
code. In the code, a three-base sequence, called a codon, specifies
one particular amino acid among the 20 possible choices. One
or more polypeptides form a functional protein. In this way, the
DNA can store the information to specify the proteins made by
an organism.
DNA Sequence Amino Acid Sequence
ATG GGC CTT AGC Methionine Glycine Leucine Serine
TTT AAG CTT GCC Phenylalanine Lysine Leucine Alanine
In living cells, the DNA is found within large structures known as
chromosomes. Figure 1.5 is a micrograph of the 46 chromosomes
contained in a cell from a human male; this type of image is known
cell from a human male.
©Kateryna Kon/Shutterstock
CONCEPT CHECK: Which types of macromolecules are found in chromosomes?
as a karyotype. The DNA of an average human chromosome is an
extraordinarily long, linear, double-stranded structure that contains
well over a hundred million nucleotides. Along the immense length
of a chromosome, the genetic information is parceled into func-
tional units known as genes. An average-sized human chromosome
is expected to contain about 1000 different protein-encoding genes.
The Information in DNA Is Accessed During
the Process of Gene Expression
To synthesize its proteins, a cell must be able to access the infor-
mation that is stored within its DNA. The process of using a gene
sequence to affect the characteristics of cells and organisms is
referred to as gene expression. At the molecular level, the infor-
mation within genes is accessed in a stepwise process (Figure 1.6).
In the first step, known as transcription, the DNA sequence
within a gene is copied into a nucleotide sequence of ribonucleic
acid (RNA). Protein-encoding genes (also called structural
genes) carry the information for the amino acid sequence of a
polypeptide. When a protein-encoding gene is transcribed, the
first product is an RNA molecule known as messenger RNA
(mRNA). During polypeptide synthesis—a process called
translation—the sequence of nucleotides within the mRNA
determines the sequence of amino acids in a polypeptide. One or
more polypeptides then fold and assemble into a functional pro-
tein. The synthesis of functional proteins largely determines an
organism’s traits. As discussed further in Chapter 12 (look ahead
to Figure 12.1), the pathway of gene expression from DNA to
RNA to protein is called the central dogma of genetics (also
called the central dogma of molecular biology). It forms a corner-
stone of our understanding of genetics at the molecular level.
6 C H A P T E R 1 :: OVERVIEW OF GENETICS

DNA 4. The direct result of the process of transcription is the synthesis of

a. DNA.
Gene
b. RNA.
Transcription
c. a polypeptide.
d. all of the above.

RNA (messenger RNA)

1.2 THE RELATIONSHIP

Translation BETWEEN GENES AND TRAITS
Learning Outcomes:
1. Outline how the expression of genes leads to an organism’s
Protein traits.
(sequence of
amino acids)
2. Define genetic variation.
3. Discuss the relationship between genes and traits.
4. Describe how genes are transmitted in sexually reproducing
species.
5. Explain the process of evolution.
Functioning of proteins within living
cells influences an organism’s traits. A trait is any characteristic that an organism displays. In genetics,
we often focus our attention on morphological traits, those that
FI GURE 1.6 Gene expression at the molecular
affect the appearance, form, and structure of an organism. The
level. The expression of a gene is a multistep process.
During transcription, one of the DNA strands is used as a color of a flower and the height of a pea plant are morphological
template to make an RNA strand. During translation, the traits. Geneticists frequently study these types of traits because they
RNA strand is used to specify the sequence of amino acids are easy to evaluate. For example, an experimenter can simply look
within a polypeptide. One or more polypeptides form a protein that at a plant and tell if it has red or white flowers. However, not all
functions within the cell, thereby influencing an organism’s traits. traits are morphological. Physiological traits affect the ability of
CONCEPT CHECK: Where is the information to make a polypeptide stored? an organism to function. For example, the rate at which a bacterium
metabolizes a sugar such as lactose is a physiological trait. Like
morphological traits, physiological traits are controlled, in part, by
the expression of genes. Behavioral traits affect the ways an organ-
ism responds to its environment. An example is the mating calls of
bird species. In animals, the nervous system plays a key role in
governing such traits. In this section, we will examine the relation-
1.1 COMPREHENSION QUESTIONS ship between the expression of genes and an organism’s traits.
1. Which of the following is not a constituent of a cell’s proteome?
a. An enzyme The Molecular Expression of Genes Leads
b. A cytoskeletal protein to an Organism’s Traits
c. A transport protein in the plasma membrane A complicated, yet very exciting, aspect of genetics is that the
d. An mRNA field’s observations and theories span four levels of biological
2. A gene is a segment of DNA that contains the information to organization: molecules, cells, organisms, and populations. This
produce a functional product. The functional product of most broad scope can make it difficult to appreciate the relationship
genes is between genes and traits. To understand this connection, we need
a. DNA. to relate the following phenomena:
b. mRNA. 1. Genes are expressed at the molecular level. In other words,
c. a polypeptide. gene transcription and translation lead to the production of
d. none of the above. a particular protein, which is a molecular process.
2. Proteins often function at the cellular level. The function
3. The function of the genetic code is to
of a protein within a cell affects the structure and workings
a. promote transcription.
of that cell.
b. specify the amino acids within a polypeptide. 3. An organism’s traits are determined by the characteristics
c. alter the sequence of DNA. of its cells. We do not have microscopic vision, yet when
d. do none of the above. we view morphological traits, we are really observing the
 1.2 The Relationship Between Genes and Traits 7

properties of an individual’s cells. For example, a red

flower has its color because its cells make a red pigment.
The trait of red flower color is an observation at the
organism level. Yet the trait is rooted in the molecular Pigmentation gene Pigmentation gene
(dark allele) (light allele)
characteristics of the organism’s cells. found in a dark butterfly found in a light butterfly
4. A species is a group of organisms that maintains a
distinctive set of attributes in nature. The occurrence of a Transcription and translation
trait within a species is an observation at the population
level. Along with learning how a trait occurs, we also want
to understand why a trait becomes prevalent in a particular
species. In many cases, researchers discover that a trait
predominates within a population because it promotes the
reproductive success of the members of the population. Highly functional Poorly functional
pigment-synthesizing enzyme pigment-synthesizing enzyme
This leads to the evolution of beneficial traits.
(a) Molecular level
To illustrate the four levels of genetics with an example, Figure 1.7
considers the trait of pigmentation in a species of butterflies.
Some members of this species are dark colored and others are very
Pigment
light. Let’s consider how we can explain this trait at the molecular, molecule
cellular, organism, and population levels.
At the molecular level, we need to understand the nature of the Wing cells
gene or genes that govern this trait. As shown in Figure 1.7a, a gene,
which we will call the pigmentation gene, is responsible for the
amount of pigment produced. The pigmentation gene exists in two Lots of pigment made Little pigment made
different versions. Alternative versions of a specific gene are called (b) Cellular level
alleles. In this example, one allele confers a dark pigmentation and
the other causes a light pigmentation. Each of these alleles encodes a
protein that functions as a pigment-synthesizing enzyme. However,
the DNA sequences of the two alleles differ slightly from each other.
This difference in the DNA sequences leads to a variation in the struc-
ture and function of the respective pigment-synthesizing enzymes.
At the cellular level (Figure 1.7b), the functional differences
between the two pigment-synthesizing enzymes affect the amount Dark butterfly Light butterfly
of pigment produced. The allele causing dark pigmentation, which
is shown on the left, encodes an enzyme that functions very well. (c) Organism level
Therefore, when this gene is expressed in the cells of the wings, a
large amount of pigment is made. By comparison, the allele caus-
ing light pigmentation encodes an enzyme that functions poorly.
Therefore, when this allele is the only pigmentation gene
expressed, little pigment is made.
At the organism level (Figure 1.7c), the amount of pigment
in the wing cells governs the color of the wings. If the pigment-
synthesizing enzymes produce high amounts of pigment, the
wings are dark colored. If the enzymes produce little pigment, the
wings are light.
Finally, at the population level (Figure 1.7d), geneticists
would like to know why a species of butterfly would contain some
members with dark wings and other members with light wings.
Dark butterflies usually live Light butterflies usually live
One possible explanation is differential predation. The butterflies in forested areas. in unforested areas.
with dark wings might better avoid being eaten by birds if they
happened to live within a dimly lit forest. The dark wings would (d) Population level
help to camouflage the butterfly if it were perched on a dark sur-
face such as a tree trunk. In contrast, light-colored wings would be F I G URE 1 . 7 The relationship between genes and traits at the
an advantage if the butterfly inhabited a brightly lit meadow. Un- (a) molecular, (b) cellular, (c) organism, and (d) population levels.
der these conditions, a bird might be less likely to notice a light- CONCEPT CHECK: Which butterfly has a more active pigment-synthesizing
colored butterfly that was perched on a sunlit surface. A population enzyme, the dark- or light-colored one?
8 C H A P T E R 1 :: OVERVIEW OF GENETICS

geneticist might study this species of butterfly and find that the
dark-colored members usually live in forested areas and the light-
colored members reside in unforested areas.

Inherited Differences in Traits Are

Due to Genetic Variation
In Figure 1.7, we considered how gene expression leads to varia-
tion in a trait of organisms, using the example of dark- versus F I G URE 1 . 8 Two dyeing poison frogs (Dendrobates tinctorius)
light-colored wings in a species of butterflies. Variation in traits are examples of different morphs within a single species.
among members of the same species is very common. For example, (Left): ©Natalia Kuzmina/Shutterstock; (Right): ©Valt Ahyppo/Shutterstock
some people have brown hair and others have blond hair; some CONCEPT CHECK: Why do these two frogs look so different?
petunias have white flowers and others have purple flowers. These
are examples of genetic variation. This term refers to the differ-
ences in inherited traits among individuals within a population. Traits Are Governed by Genes
In large populations that occupy a wide geographic range, and by the Environment
genetic variation can be quite striking. Morphological differences
In our discussion thus far, we have considered the role that genes
have often led geneticists to misidentify two members of the same
play in determining an organism’s traits. Another critical factor is
species as belonging to separate species. As an example, Figure 1.8
the environment—the surroundings in which an organism exists.
shows two dyeing poison frogs that are members of the same species,
A variety of factors in an organism’s environment profoundly
Dendrobates tinctorius. They display dramatic differences in their
affects its morphological and physiological features. For example,
markings. Such contrasting forms within a single species are termed
a person’s diet greatly influences many traits, such as height,
morphs. You can easily imagine how someone might mistakenly
weight, and even intelligence. Likewise, the amount of sunlight a
conclude that these frogs are not members of the same species.
plant receives affects its growth rate and the color of its flowers.
Changes in the nucleotide sequence of DNA underlie the genetic
An interesting example of the interplay between genes and
variation that we see among individuals. Throughout this textbook,
the environment involves the human genetic disease phenylke-
we will routinely examine how variation in the genetic material results
tonuria (PKU). Humans have a gene that encodes an enzyme
in changes in an organism’s traits. At the molecular level, genetic
known as phenylalanine hydroxylase. Most people have two func-
variation can be attributed to different types of modifications.
tional copies of this gene. People with one or two functional cop-
1. Small or large differences can occur within gene sequences. ies of the gene can eat foods containing the amino acid
When such changes initially occur, they are called gene phenylalanine and metabolize it properly.
mutations. Mutations result in genetic variation in which a A rare variation in the gene that encodes phenylalanine hydrox-
gene is found in two or more alleles, as previously de- ylase results in a nonfunctional version of this enzyme. Individuals
scribed in Figure 1.7. In many cases, gene mutations alter
the expression or function of a protein that a gene encodes.
2. Major alterations can also occur in the structure of a chro-
mosome. A large segment of a chromosome can be lost, re-
arranged, or reattached to another chromosome.
3. Variation may also occur in the total number of chromo-
somes. In some cases, an organism may inherit one too
many or one too few chromosomes. In other cases, it may
inherit an extra set of chromosomes.
Variations of sequences within genes are a common source of
genetic variation among members of the same species. In humans,
familiar examples of sequence variation involve genes for eye
color, hair texture, and skin pigmentation. Chromosome variation—
a change in chromosome structure or number (or both)—is also
found, but this type of change is often detrimental. Many human
(a) (b)
genetic disorders are the result of chromosomal alterations. The
most common example is Down syndrome, which is due to the F I G URE 1 . 9 Examples of chromosome variation. (a) A person
presence of an extra chromosome (Figure 1.9a). By comparison, with Down syndrome. She has 47 chromosomes rather than the common
chromosome variation in plants is common and often results in number of 46, because she has an extra copy of chromosome 21. (b) A
plants with superior characteristics, such as increased resistance to wheat plant. Cultivated wheat has six sets of chromosomes.
disease. Plant breeders have frequently exploited this observation. (a): ©Stockbyte/Alamy; (b): ©Pixtal/age fotostock
Cultivated varieties of wheat, for example, have many more chro- CONCEPT CHECK: Do these examples constitute variation in chromosome
mosomes than the wild species does (Figure 1.9b). structure or variation in chromosome number?
 1.2 The Relationship Between Genes and Traits 9

with two copies of this rare, inactive allele cannot metabolize phenyla-
lanine properly. When given a standard diet containing phenylalanine,
individuals with this disorder are unable to break down this amino
acid. Phenylalanine accumulates and is converted into phenylketones,
which are detected in the urine. Individuals with PKU can manifest a
variety of detrimental traits, including mental impairment, underdevel-
oped teeth, and foul-smelling urine. Fortunately, through routine new-
born screening in the United States, PKU is now diagnosed early. Part
of the treatment is a diet that restricts phenylalanine, which is present
in high-protein foods such as eggs, meat, and dairy products. Restrict-
ing phenylalanine allows the affected child to develop normally. PKU
provides a dramatic example of how the environment and an individu-
al’s genes can interact to influence the traits of the organism.
During Reproduction, Genes Are Passed
from Parent to Offspring
Now that we have considered how genes and the environment govern
the outcome of traits, we can turn to the issue of inheritance. How are
traits passed from parents to offspring? The foundation for our under-
standing of inheritance came from Gregor Mendel’s study of pea
plants in the nineteenth century. His work revealed that the genetic
determinants that govern traits, which we now call genes, are passed
from parent to offspring as discrete units. We can predict the outcome
of many genetic crosses based on Mendel’s laws of inheritance.
The inheritance patterns identified by Mendel can be
explained by the existence of chromosomes and their behavior
during cell division. Like Mendel’s pea plants, sexually reproduc-
ing species are commonly diploid. This means that their cells
contain two copies of each chromosome, one from each parent.
The two copies are called homologs of each other. Because genes
are located within chromosomes, diploid organisms have two cop-
ies of most genes. Humans, for example, have 46 chromosomes,
which are found in homologous pairs (Figure 1.10a). With the
exception of the sex chromosomes (X and Y), each homologous
pair contains the same kinds of genes. For example, both copies of
human chromosome 12 carry the gene that encodes phenylalanine
hydroxylase, the enzyme that is nonfunctional in people with
PKU. Therefore, each individual has two copies of this gene,
which may or may not be identical alleles.
Most cells of the human body that are not directly involved
in sexual reproduction contain 46 chromosomes. These cells are
called somatic cells. In contrast, the gametes—sperm and egg
cells—contain half that number (23) and are termed haploid
(Figure 1.10b). The union of gametes during fertilization restores
the diploid number of chromosomes. The primary advantage of
sexual reproduction is that it enhances genetic variation. For exam-
ple, a tall person with blue eyes and a short person with brown eyes
may have short offspring with blue eyes or tall offspring with brown
eyes. Therefore, sexual reproduction can result in new combina-
tions of two or more traits that differ from those of either parent.
The Genetic Composition of a Species Evolves
from Generation to Generation
As we have just seen, sexual reproduction has the potential to enhance
genetic variation. This can be an advantage for a population of indi-
viduals as they struggle to survive and compete within their natural
environment. The term biological evolution, or simply, evolution,

1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8

9 10 11 12 13 14 15 16 9 10 11 12 13 14 15 16

17 18 19 20 21 22 XX 17 18 19 20 21 22 X

(a) Chromosomal composition found (b) Chromosomal composition found in

in human somatic cells of females a human gamete (23 chromosomes)
(46 chromosomes)

FI GURE 1.10 The complement of human chromosomes in somatic cells and gametes. (a) A schematic drawing of the 46 chromosomes of a
human. With the exception of the sex chromosomes, these are always found in homologous pairs in somatic cells, such as skin or nerve cells. (b) The
chromosomal composition of a gamete, which contains only 23 chromosomes, one from each pair. This gamete contains an X chromosome. Half of the
gametes from human males contain a Y chromosome instead of an X chromosome.
CONCEPT CHECK: The leaf cells of a corn plant contain 20 chromosomes each. How many chromosomes are found in a gamete made by a corn plant?
10 C H A P T E R 1 :: OVERVIEW OF GENETICS

refers to the process of changes in the genetic makeup of a population
from one generation to the next.
As suggested by Charles Darwin in the nineteenth century, the
members of a species are in competition with one another for essen-
tial resources. Random genetic changes (i.e., mutations) occasion-
ally occur within an individual’s genes, and sometimes these changes
lead to a modification of traits that promote reproductive success.
For example, over the course of many generations, random gene
mutations have lengthened the snout and extended the tongue of the
anteater, enabling it to probe into the ground and feed on ants. When
a mutation creates a new allele that is beneficial, the allele may
become prevalent in future generations because the individuals car-
rying the allele are more likely to reproduce and pass the beneficial
allele to their offspring. This process is known as natural selection.
In this way, a species becomes better adapted to its environment.
Over a long period of time, the accumulation of many genetic
changes may lead to rather striking modifications in a species’ char-
acteristics. As an example, Figure 1.11 depicts the evolution of the
modern-day horse. Over time, a variety of morphological changes
occurred, including an increase in size, fewer toes, and modified
jaw structure. The changes can be attributed to natural selection.
Over North America, where much of horse evolution occurred,

Equus
0 Hippidium and other genera

Nannippus
Stylohipparion
5 Pliohippus

Hipparion Neohipparion

10

Sinohippus Megahippus
Calippus

Archaeohippus
20
Merychippus
Millions of years ago (mya)

Anchitherium Hypohippus

Parahippus

Miohippus

Mesohippus
40

Paleotherium Epihippus

Propalaeotherium

Pachynolophus Orohippus

55
Hyracotherium

FI G U RE 1.1 1 The evolutionary changes that led to the modern horse genus, Equus. Three important morphological changes that occurred
were larger size, fewer toes, and a shift toward a jaw structure suited for grazing.
CONCEPT CHECK: According to the theory of evolution, why have these changes occurred in horse populations over the course of many generations?
 1.3 Fields of Genetics 11

large areas of dense forests were replaced with grasslands. The 4. Evolutionary change caused by natural selection results in spe-
increase in size and changes in foot structure enabled horses to cies with
escape predators more easily and travel greater distances in search a. greater complexity.
of food. Natural selection favored the changes seen in horses’ teeth, b. less complexity.
because such changes allowed them to eat grasses and other types c. greater reproductive success in their environment.
of vegetation that are tougher and require more chewing.
d. the ability to survive longer.

1.2 COMPREHENSION QUESTIONS

1.3 FIELDS OF GENETICS
1. At which of the following levels can gene expression be observed?
a. Molecular and cellular levels Learning Outcome:
b. Organism level 1. Compare and contrast the three major fields of genetics:
c. Population level transmission, molecular, and population genetics.
d. All of the above
2. Variation in the traits of organisms may be attributable to Genetics is a broad discipline encompassing molecular, cellular,
a. gene mutations. organism, and population biology. Many scientists who are inter-
b. alterations in chromosome structure. ested in genetics have been trained in supporting disciplines such
c. variation in chromosome number. as biochemistry, biophysics, cell biology, mathematics, microbiol-
d. all of the above. ogy, population biology, ecology, agriculture, and medicine.
Experimentally, geneticists often focus their efforts on model
3. A human skin cell has 46 chromosomes. A human sperm cell has
organisms—organisms studied by many different researchers so
a. 23.
that they can compare their results and determine scientific prin-
b. 46. ciples that apply more broadly to other species. Figure 1.12 shows
c. 92. some examples of model organisms: Escherichia coli (a bacte-
d. None of the above is the number of chromosomes in a rium), Saccharomyces cerevisiae (a yeast), Drosophila melano-
sperm cell. gaster (fruit fly), Caenorhabditis elegans (a nematode worm),

0.3 μm 7 μm

(a) Escherichia coli (b) Saccharomyces cerevisiae (c) Drosophila melanogaster

156 μm

(d) Caenorhabditis elegans (e) Mus musculus (f) Arabidopsis thaliana

FI GURE 1.12 Examples of model organisms studied by geneticists. (a) Escherichia coli (a bacterium), (b) Saccharomyces cerevisiae
(a yeast), (c) Drosophila melanogaster (fruit fly), (d) Caenorhabditis elegans (a nematode worm), (e) Mus musculus (mouse), and (f) Arabidopsis
thaliana (a flowering plant).
(a): Source: Peggy S. Hayes & Elizabeth H. White, M.S./CDC; (b): ©Science Photo Library/Alamy; (c): ©Janeff/Getty Images; (d): ©Sinclair Stammers/Science Source; (e): ©G.K. & Vikki Hart/
Getty Images; (f): ©WILDLIFE GmbH/Alamy

CONCEPT CHECK: Can you think of another example of a model organism?

12 C H A P T E R 1 :: OVERVIEW OF GENETICS
Mus musculus (mouse), and Arabidopsis thaliana (a flowering
plant). Model organisms offer experimental advantages over other
species. For example, E. coli is a very simple organism that can be
easily grown in the laboratory. By limiting their work to a few
model organisms, researchers can more easily unravel the genetic
mechanisms that govern the traits of a given species. Furthermore,
the genes found in model organisms often function in a similar
way to those found in humans.
The study of genetics has been traditionally divided into
three areas—transmission, molecular, and population genetics—
although there is some overlap of these three fields. In this section,
we compare these three fields of genetics.
Transmission Genetics Explores the Inheritance
Patterns of Traits as They Are Passed from Parents
to Offspring
A scientist working in the field of transmission genetics examines
the relationship between the transmission of genes from parent to
offspring and the outcome of the offspring’s traits. For example,
how can two brown-eyed parents produce a blue-eyed child? Or
why do tall parents tend to produce tall children, but not always?
Our modern understanding of transmission genetics began with
the studies of Gregor Mendel. His work provided the conceptual
framework for transmission genetics. In particular, he originated
the idea that factors, which we now call genes, are passed as dis-
crete units from parents to offspring via sperm and egg cells. Since
Mendel’s pioneering studies in the 1860s, our knowledge of
genetic transmission has skyrocketed. Many patterns of genetic
transmission are more complex than the simple Mendelian pat-
terns that are described in Chapter 2. The additional complexities
of transmission genetics are examined in Chapters 3 through 8.
Experimentally, the fundamental technique used by a trans-
mission geneticist is the genetic cross—the breeding of two
selected individuals and then analyzing their offspring in an
attempt to understand how traits are passed from parents to off-
spring. In the case of experimental organisms, the researcher
chooses two parents with particular traits and then categorizes the
offspring according to the traits they possess. In many cases, this
analysis is quantitative in nature. For example, an experimenter
may cross two tall pea plants and obtain 100 offspring that fall into
two categories: 75 tall and 25 dwarf. As we will see in Chapter 2,
the ratio of tall to dwarf offspring provides important information
concerning the inheritance pattern of the height trait.
Molecular Genetics Focuses on a Biochemical
Understanding of the Hereditary Material
The goal of molecular genetics, as the name of the field implies, is
to understand how the genetic material works at the molecular
level. In other words, molecular geneticists want to understand the
molecular features of DNA and how these features underlie the
expression of genes. The experiments of molecular geneticists are
usually conducted within the confines of a laboratory. Their efforts
frequently progress to a detailed analysis of DNA, RNA, and pro-
teins, using a variety of techniques that are described throughout
Parts III, IV, and V of this textbook.
Molecular geneticists often study mutant genes that have
abnormal function. This is called a genetic approach to the study
of a research question. In many cases, researchers analyze the
effects of a gene mutation that eliminates the function of a gene.
This is called a loss-of-function mutation, and the resulting ver-
sion of the gene is called a loss-of-function allele. Studying the
effect of such a mutation often reveals the role of the functional,
nonmutant gene. For example, let’s suppose that a particular plant
species produces purple flowers. If a loss-of-function mutation
within a given gene causes a plant of that species to produce white
flowers, you might suspect that the role of the functional gene
involves the production of purple pigmentation.
Studies within molecular genetics interface with other disci-
plines such as biochemistry, biophysics, and cell biology. In addi-
tion, advances within molecular genetics have shed considerable
light on the areas of transmission and population genetics. Our
quest to understand molecular genetics has spawned a variety of
modern molecular technologies and computer-based approaches.
Furthermore, discoveries within molecular genetics have had wide-
spread applications in agriculture, medicine, and biotechnology.
Population Genetics Is Concerned with Genetic
Variation and Its Role in Evolution
The foundations of population genetics arose during the first few
decades of the twentieth century. Although many scientists of this era
did not accept the findings of Mendel or Darwin, the theories of
population genetics provided a compelling way to connect the two
viewpoints. Mendel’s work and that of many succeeding geneticists
gave insight into the nature of genes and how they are transmitted
from parents to offspring. The work of Darwin provided a natural
explanation for the variation in characteristics observed among the
members of a species. To relate these two phenomena, population
geneticists have developed mathematical theories to explain the prev-
alence of certain alleles within populations of individuals. This work
helps us understand how processes such as natural selection have
resulted in the prevalence of individuals that carry particular alleles.
Population geneticists are particularly interested in genetic
variation and how that variation is related to an organism’s envi-
ronment. In this field, the frequencies of alleles within a popula-
tion are of central importance.
1.3 COMPREHENSION QUESTIONS
1. Which of the following is not a model organism?
a. Mus musculus (laboratory mouse)
b. Escherichia coli (a bacterium)
c. Saccharomyces cerevisiae (a yeast)
d. Sciurus carolinensis (gray squirrel)
2. A person studying the rate of transcription of a particular gene is
working in the field of
a. molecular genetics.
b. transmission genetics.
c. population genetics.
d. None of the above is correct.

1.4 THE SCIENCE OF GENETICS
Learning Outcomes:
1. Describe what makes genetics an experimental science.
2. Outline different strategies for solving problems in genetics.
Science is a way of knowing about our natural world. The science
of genetics allows us to understand how the expression of our
genes produces the traits that we possess. In this section, we will
consider how scientists attempt to answer questions via experi-
mentation. We will also consider general approaches for solving
problems.
Genetics Is an Experimental Science
Researchers typically follow two general types of scientific
approaches: hypothesis testing and discovery-based science. In
hypothesis testing, also called the scientific method, scientists
follow a series of steps to reach verifiable conclusions about the
world. Although scientists arrive at their theories in different
ways, the scientific method provides a way to validate (or invali-
date) a particular hypothesis. Alternatively, research may also
involve the collection of data without a preconceived hypothesis.
For example, researchers might analyze the genes found in cancer
cells to identify those that have become mutant. In this case, the
scientists may not have a hypothesis about which particular genes
may be involved. The collection and analysis of data without the
need for a preconceived hypothesis is called discovery-based
science or, simply, discovery science.
In traditional science textbooks, the emphasis often lies on
the product of science. That is, many textbooks are aimed primar-
ily at teaching the student about the observations scientists have
made and the hypotheses they have proposed to explain those ob-
servations. Along the way, the student is provided with many bits
and pieces of experimental techniques and data. Likewise, this
textbook also provides you with many observations and hypothe-
ses. However, it attempts to go one step further. Most chapters
contain one or two figures presenting experiments that have been
“dissected” into five individual components to help you to under-
stand the entire scientific process:
1. Background information is provided so that you can appre-
ciate observations that were known prior to conducting the
experiment.
2. Most experiments involve the testing of a hypothesis via
the scientific method. In those cases, the figure presenting
the experiment states the hypothesis the scientists were try-
ing to test. In other words, what scientific question were
the researchers trying to answer?
3. Next, the figure follows the experimental steps the scien-
tists took to test the hypothesis. The figure presents two
parallel illustrations labeled “Experimental Level” and
“Conceptual Level.” The experimental level helps you to
understand the techniques that were used. The conceptual
level helps you to understand what is actually happening at
each step in the procedure.
1.4 The Science of Genetics 13
4. The raw data from the experiment are then presented.
5. Last, an interpretation of the data is offered within the text.
The rationale behind this approach is that it enables you to see the
experimental process from beginning to end. As you read through
the chapters, the experiments will help you to see the relationship
between science and scientific theories.
As a student of genetics, you will be given the opportunity
to involve your mind in the experimental process. As you are read-
ing an experiment, you may find yourself thinking about different
approaches and alternative hypotheses. Different people can view
the same data and arrive at very different conclusions. As you
progress through the experiments in this book, you will enjoy
genetics far more if you try to develop your skills at formulating
hypotheses, designing experiments, and interpreting data. Also,
some of the questions in the problem sets are aimed at refining
these skills.
Finally, it is worthwhile to point out that science is a social
discipline. As you develop your skills at scrutinizing experiments,
it is fun to discuss your ideas with other people, including fellow
students and faculty members. Keep in mind that you do not need
to “know all the answers” before you enter into a scientific discus-
sion. Instead, it is more rewarding to view science as an ongoing
and never-ending dialogue.
Genetic TIPS Will Help You to Improve Your
Problem-Solving Skills
As your progress through this textbook, your learning will involve
two general goals:
∙∙ You will gather foundational knowledge. In other words,
you will be able to describe core concepts in genetics. For
example, you will be able to explain how DNA replication
occurs and describe the proteins that are involved in this
process.
∙∙ You will develop problem-solving skills that allow you to
apply that foundational knowledge in different ways. For
example, you will learn how to use statistics to determine
if a genetic hypothesis is consistent with experimental
data.
The combination of foundational knowledge and problem-solving
skills will enable you not only to understand genetics, but also to
apply your knowledge in different situations. To help you develop
these skills, Chapters 2 through 29 contain solved problems named
Genetic TIPS, which stands for Topic, Information, and Problem-
solving Strategy. These solved problems follow a consistent pattern.
GENETIC TIPS THE QUESTION: All of the Genetic TIPS
begin with a question. As an example, let’s consider the following
question:
    The coding strand of DNA in a segment of a gene is as follows:
ATG GGC CTT AGC. This strand carries the information to make a
region of a polypeptide with the amino acid sequence, methionine-
glycine-leucine-serine. What would be the consequences if a mutation
changed the second cytosine (C) in this sequence to an adenine (A)?
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with every tide! This became the universal supply for us and for the
Natives all round the Harbour and for miles inland. Hundreds of
Natives from all parts of Tanna flocked to examine this greatest
wonder they had ever seen—rain rising up out of the earth. I built it
round with a kind of stone brought in my boat from the other side of
the bay; and for many years it was the only fresh water supply for the
Natives all around. Some years later a native Chief sank a well about
a mile nearer the entrance to the Harbour at his own village, and
built it round with the bricks that I had purchased for house-
building; these he grabbed and thus appropriated! Many a vessel,
calling at the Harbour, was glad to get her casks refilled at my well,
and all were apparently more friendly because of it; but the Sinking
of this Well produced no such revolution as on Aniwa,—to be
hereafter related.
For fully three months, all our available time, with all the native
help which I could hire, was spent in erecting a building to serve for
Church and School. It was fifty feet long, by twenty-one feet six
inches broad. The studs were three feet apart, and all fixed by tenon
and mortise into upper and lower wall plates. The beautiful roof of
iron, wood, and sugar-cane leaf, was supported by three massive
pillars of wood, sunk deeply into the ground. The roof extended
about three feet over the wall plates, both to form a verandah and to
carry the rain-drop free beyond the walls. It was made of sugar-cane
leaf and cocoa-nut leaves all around. The floor was laid with white
coral, broken small, and covered with cocoa-nut leaf mats, such as
those on which the Natives sat. Indeed, it was as comfortable a
House of Prayer as any man need wish for in the tropics, though
having only open spaces for doors and windows! I bought the heavy
wood for it on Aneityum—price, fifty pairs of native trousers; and
these again were the gift of my Bible Class in Glasgow, all cut and
sewed by their own hands. I gave also one hundred and thirty yards
of cloth, along with other things, for other needful wood.
My Tannese people at first opposed the erection of a Church. They
did not wish Jehovah to secure a House on their island. On the
opening day, only five men, three women, and three children were
present, besides our Aneityumese Teachers. But after the morning
service, on that day, I visited ten villages, and had worship in each.
The people were generally shy and unfriendly. They said that we
were the cause of the prevailing sickness and fever. They had no idea
of any sickness or death being natural, but believed that all such
events were caused by some one nahaking, i.e., bewitching them.
Hence their incessant feuds; and many were murdered in blind
revenge.
As we were preparing a foundation for the Church, a huge and
singular-looking round stone was dug up, at sight of which the
Tannese stood aghast. The eldest Chief said,—
“Missi, that stone was either brought there by Karapanamun (the
Evil Spirit), or hid there by our great Chief who is dead. That is the
Stone God to which our forefathers offered human sacrifices; these
holes held the blood of the victim till drunk up by the Spirit. The
Spirit of that stone eats up men and women and drinks their blood,
as our fathers taught us. We are in greatest fear!”
A Sacred Man claimed possession, and was exceedingly desirous to
carry it off; but I managed to keep it, and did everything in my power
to show them the absurdity of these foolish notions. Idolatry had not,
indeed, yet fallen throughout Tanna, but one cruel idol, at least, had
to give way for the erection of God’s House on that benighted land.
An ever-memorable event was the printing of my first book in
Tannese. Thomas Binnie, Esq., Glasgow, gave me a printing-press
and a font of type. Printing was one of the things I had never tried,
but having now prepared a booklet in Tannese, I got my press into
order, and began fingering the type. But book-printing turned out to
be for me a much more difficult affair than house-building had been.
Yet by dogged perseverance I succeeded at last. My biggest difficulty
was how to arrange the pages properly! After many failures, I folded
a piece of paper into the number of leaves wanted, cut the corners,
folding them back, and numbering as they would be when correctly
placed in the book; then folding all back without cutting up the sheet,
I found now by these numbers how to arrange the pages in the frame
or case for printing, as indicated on each side. And do you think me
foolish, when I confess that I shouted in an ecstasy of joy when the
first sheet came from the press all correct? It was about one o’clock
in the morning. I was the only white man then on the island, and all
the Natives had been fast asleep for hours! Yet I literally pitched my
hat into the air, and danced like a schoolboy round and round that
printing-press; till I began to think, Am I losing my reason? Would it
not be liker a Missionary to be upon my knees, adoring God for this
first portion of His blessed Word ever printed in this new language?
Friend, bear with me, and believe me, that was as true worship as
ever was David’s dancing before the Ark of his God! Nor think that I
did not, over that first sheet of God’s Word ever printed in the
Tannese tongue, go upon my knees too, and then, and every day
since, plead with the mighty Lord to carry the light and joy of His
own Holy Bible into every dark heart and benighted home on Tanna!
But the Tannese had a superstitious dread of books, and especially of
God’s Book. I afterwards heard that Dr. Turner had printed a small
primer in Tannese, translated by the help of the Samoan Teachers;
but this I never saw till near the close of my work on Tanna. Dr.
Geddie sent me a copy, but it was more Samoan than Tannese,
especially in its spelling, and I could make little or nothing of it.
Shortly after this, I was greatly refreshed by the visit of an
American whaler, the Camden Packet, under Captain Allan. He, his
chief officer, and many of his double company of seamen, were
decided Christians—a great contrast to most of the Traders that had
called at Port Resolution. The Captain cordially invited me on board
to preach and conduct a religious service. That evening I enjoyed
exceedingly—wells in the desert! The Captain introduced me, saying,
—
“This is my ship’s company. My first officer and most of my men
are real Christians, trying to love and serve Jesus Christ. We have
been three years out on this voyage, and are very happy with each
other. You would never hear or see worse on board of this vessel than
you see now. And God has given us gratifying success.”
He afterwards told me that he had a very valuable cargo of sperm
oil on board, the vessel being nearly filled up with it. He was eager to
leave supplies, or do something for me, but I needed nothing that he
could give. His mate, on examining my boat, found a hole in her, and
several planks split and bulged in, as I had gone down on a reef with
her when out on Mission work, and narrowly escaped drowning.
Next morning, the Captain, of his own accord, set his carpenter to
repair the boat, and left it as good as new. Not one farthing of
recompense would any of them take from me; their own Christian
love rewarded them, in the circumstances. I had been longing for a
chance to send it to Sydney for repairs, and felt deeply thankful for
such unexpected and generous aid. The Captain would not admit
that the delay was any loss to him,—his boats spending the day in
purchasing cocoa-nuts and provisions from the Natives for his own
ship. Oh, how the Christlike spirit knits together all true followers of
Christ! What other earthly or human tie could have so bound that
stranger to me? In the heart of Christ we met as brothers.
Dangers again darkened round me. One day, while toiling away at
my house, the war Chief, his brother, and a large party of armed men
surrounded the plot where I was working. They all had muskets,
besides their own native weapons. They watched me for some time in
silence, and then every man levelled a musket straight at my head.
Escape was impossible. Speech would only have increased my
danger. My eyesight came and went for a few moments. I prayed to
my Lord Jesus, either Himself to protect me, or to take me home to
His Glory. I tried to keep working on at my task, as if no one was
near me. In that moment, as never before, the words came to me,
—“Whatsoever ye shall ask in My name, I will do it;” and I knew that
I was safe. Retiring a little from their first position, no word having
been spoken, they took up the same attitude somewhat farther off,
and seemed to be urging one another to fire the first shot. But my
dear Lord restrained them once again, and they withdrew, leaving
me with a new cause for trusting Him with all that concerned me for
Time and Eternity. Perils seemed, however, to enclose me on every
hand, and my life was frequently attempted. I had to move about
more cautiously than ever, some days scarcely daring to appear
outside my Mission premises. For I have ever most firmly believed,
and do believe, that only when we use every lawful and possible
means for the preservation of our life, which is God’s second greatest
gift to man (His Son being the first), can we expect God to protect us,
or have we the right to plead His precious promises.
The vessel of one calling himself Prince de Jean Beuve, a French
refugee, who had become a naturalized American, visited Port
Resolution. He said, he had to escape from his own country for
political offences. His large and beautiful ship was fitted up and
armed like a Man-of-war. She was manned chiefly by slaves, whom
he ruled with an iron hand. What a contrast to Captain Allan’s
whaler! Yet he also was very sympathetic and kind to me. Having
heard rumour of my trials and dangers, he came on shore, as soon as
his ship cast anchor, with a body of armed men. He was effusively
polite, with all a Frenchman’s gush and gesticulation, and offered to
do anything possible for me. He would take me to Aneityum or
Sydney or wherever I wished. The ship was his own; he was sailing
chiefly for pleasure, and he had called at our Islands to see if
sufficient trade could be opened up to justify his laying on a line of
steamers to call here in their transit. He urged me, I believe
sincerely, to give him the pleasure of taking me and my belongings to
some place of safety. But I was restrained from leaving, through the
fear that I would never be permitted to return, and that Christ’s work
would suffer. In the still burning hope of being able to lead the
Tannese to love and serve Jesus, I declined with much gratitude his
genuine kindness. He looked truly sorry to leave me in the
circumstances wherein I was placed. After two hours on shore, he
returned to his ship towards evening.
Knowing that the Tannese were threatening to burn my former
house, which I wished to remove to higher ground and add to the
room I now occupied on the hill, I took advantage of the presence of
the Prince’s vessel, and set my Aneityumese Teachers and some
friendly Natives to prepare for the task; but unfortunately, I forgot to
send word to the Frenchman regarding my plans and aims. We
removed the sugar-cane leaf thatch from the roof of the house, and
began burning it on cleared ground, so that I might be able to save
the heavy wood which could not be replaced on Tanna. Our French
friend, on seeing the flames rising up furiously, at once loaded his
heavy guns, and prepared his men for action. Under great
excitement, he came ashore with a large number of armed men,
leaving the rest on board ready at a given signal to protect them with
shot and shell. Leaving one half of those brought on shore to guard
the boats, he came running towards my house, followed by the other
half, wet with perspiration, and crying,—
“Fer are dey? fer are dey? De scoundrels! I vill do for dem, and
protect you. I sall punish dem, de scoundrels!”
He was so excited, he could scarcely compose himself to hear my
explanations, which, when understood, he laughed at heartily. He
again urged me to leave in his vessel; he could not bear me to lead
such a life amongst savages. I explained to him my reasons for not
leaving the island, but these he seemed unable to understand. He put
his men through drill on shore, and left them under officers, ready
for action at a moment’s warning, saying they would all be the better
for a day on shore. He wished to take pot luck with me at our Mission
House of one room for all purposes! My humble dinner and tea must
have been anything but a treat for him, but he seemed to relish the
deliverance for once from all the conventionalisms of the world.
Before he left, he sent of his own accord for all the Chiefs within
reach, and warned them that if they hurt me or took my life, he
would return with his Man-of-war and punish them, by killing
themselves and firing their villages; and that a British Man-of-war
would also come and set their island on fire. They promised all
possible good conduct, being undoubtedly put into great terror. The
kind-hearted Frenchman left, with profuse expressions of admiration
for my courage and of pity for my lot. No doubt he thought me a
foolish dreamer of dreams.
A miserable contrast befell us in the bad impression produced by
the conduct of one of Captain T——’s vessels in the Sydney sandal-
wood trade. Whale-boats had been sent out with Mr. Copeland and
myself from Glasgow, as part of the necessary equipment of every
Missionary on these Islands. Mine being rather large and heavy, I
had sold it to one of T——’s captains; but the other had also been left
to my care. After having used my boat for about twelve months—the
best boat in that trade only being expected to last two years—the
Captain called on Mr. Copeland, and got a note from him to me
regarding the sale of his boat too. He declared, when calling on me,
that Mr. Copeland had authorized him to get his boat from me in
exchange for mine, which he had now been using for a year. I asked
for the letter, and found it to be authority for me to sell his boat for
cash only and at the same price as mine. Captain V—— then raged at
me and stormed, declaring that he would return my old boat, and
take the other in defiance of me. Swearing dreadfully, he made for
his ship, and returned with a large party of men whom he had picked
up amongst the Islands. Collecting also a company of Tannese, and
offering them tobacco, he broke down the fence, burst into the boat-
house, and began to draw out the boat. Here I reached the spot, and
sternly opposed them. He swore and foamed at me, and before the
natives knocked and pulled me about, even kicking at me, though I
evaded his blows. Standing by, I said in Tannese,—
 “You are helping that man to steal my boat; he is stealing it as you
see.”
On hearing this, the Tannese ran away, and his own party alone
could not do it. In great wrath, he went off again to his vessel, and
brought on shore as much tobacco as could be held in a large
handkerchief tied by the four corners; but even for that, our own
Natives refused to help him. He offered it then to a crowd of Inland
savages, gathered at the head of the bay, who, regardless of my
remonstrances, launched the boat, he raging at and all but striking
me. Instead of returning, however, the other boat to the house, he
merely set it adrift from his vessel, and it was carried on to the reef,
where it remained fast, and was knocked about by the waves. After
his vessel left, I, with much difficulty, got it off and brought it to the
boat-house. Imagine, when such was their tyrannical treatment of a
Missionary and a British fellow-subject, how they would act towards
these poor native Islanders.
By the earliest opportunity, I wrote all the facts of the case to his
employer, Captain T—— of Sydney, but got not even a reply, while
Captain V—— continued in their trade, a scourge to these Islands,
and a dishonour to his country and to humanity. Unfriendly Tannese
now said,—
“When a white man from his own country can so pull and knock
the Missionary about and steal his boat and chain without being
punished for it, we also may do as we please!”
I hesitate not to record my conviction that that man’s conduct had
a very bad effect, emboldening them in acts of dishonesty and in
attempts upon my life till the Mission Station was ultimately broken
up. After I had to escape from Tanna, with bare life in my hand, one
of the same Captain’s vessels called at Port Resolution and gave the
Natives about three pounds weight of useless tobacco, purchasable at
Sydney for less than one shilling per pound, to allow them to take
away my boat, with oars, sails, mast, and all other belongings. They
also purchased all the plunder from my house. Both boats were so
large and so strongly built, that by adding a plank or two they turned
them into small-decked schooners, admirably suited for the sandal-
wood traffic round the shores, while larger vessels lay at safe
anchorage to receive what they collected. Once, when Dr. Inglis and I
met in Sydney, we called on Captain T—— and stated the whole case,
asking reasonable payment at least for the boats. He admitted that
the boats had been taken and were in his service, and agreed to pay
us for the boats if we would repay the large sum invested therein by
his Captains. Calling one of his clerks, he instructed him to trace in
the office record how much had been paid to the Tannese for the
Missionary’s boat.
The young man innocently returned the reply, “Three pounds of
tobacco.”
In anger, he said, “I understood that a larger value had been
given!”
The clerk assured him, “That is the only record.”
Captain T——, after discussing the worth of the boat as being about
£80, agreed to give us £60, but in writing out the cheque, threw
down the pen and shouted, “I’ll see you —— first!”
Offering £50, to which we agreed, he again resiled, and declared
he would not give a penny above £30.
We appealed to him to regard this as a debt of honour, and to
cease haggling over the price, as he well knew how we had been
wronged in the matter.
Finally we left him declaring, “I am building similar boats just now
at £25 apiece; I will send you one of them, and you may either take
that or want!”
We left, glad to get away on any terms from such a character; and,
though next year he did send one of his promised boats for me to
Aneityum, yet the conduct of his degraded servants engaged in the
sandal-wood trade had a great share in the guilt of breaking up and
ruining our Mission. Thousands upon thousands were made by it
yearly, so long as it lasted; but it was a trade steeped in human blood
and indescribable vice, nor could God’s blessing rest on them and
their ill-gotten gains. Oh, how often did we pray at that time to be
delivered from the hands of unreasonable and wicked men! Sandal-
wood traders murdered many of the Islanders when robbing them of
their wood, and the Islanders murdered many of them and their
servants in revenge. White men, engaged in the trade, also shot dead
and murdered each other in vicious and drunken quarrels, and not a
few put end to their own lives. I have scarcely known one of them
who did not come to ruin and poverty; the money that came even to
the shipowners was a conspicuous curse. Fools there made a mock at
sin, thinking that no one cared for these poor savages, but their sin
did find them out, and God made good in their experience His own
irrepealable law, “The wages of sin is death.”
Ships, highly insured, were said to be sent into our Island trade to
be deliberately wrecked. One Sabbath evening, towards dark, the
notorious Captain H——, in command of a large ship, allowed her to
drift ashore and be wrecked without any apparent effort to save her.
Next morning, the whole company were wading about in the water
and pretending to have lost everything! The Captain, put in prison
when he returned to Sydney for running away with another man’s
wife and property, imposed on Mr. Copeland and myself, getting all
the biscuits, flour, and blankets we could spare for his destitute and
shipwrecked company. We discovered afterwards that she was lying
on a beautiful bank of sand, only a few yards from the shore, and that
everything contained in her could be easily rescued without danger
to life or limb! What we parted with was almost necessary for our life
and health; of course he gave us an order on Captain T—— for
everything, but not one farthing was ever repaid. At first he made a
pretence of paying the Natives for food received; but afterwards, an
armed band went inland night by night and robbed and plundered
whatever came to hand. The Natives, seeing the food of their children
ruthlessly stolen, were shot down without mercy when they dared to
interfere; and the life of every white man was marked for speedy
revenge. Glad were we when a vessel called, and carried away these
white heathen Savages.
The same Captain T—— also began the shocking Kanaka labour-
traffic to the Colonies, after the sandal-wood trade was exhausted,
which has since destroyed so many thousands of the Natives in what
was nothing less than Colonial slavery, and has largely depopulated
the Islands either directly or indirectly. And yet he wrote, and
published in Sydney, a pamphlet declaring that he and his sandal-
wooders and Kanaka labour collectors had done more to civilize the
Islanders than all our Mission efforts combined. Civilize them,
indeed! By spreading disease and vice, misery and death amongst
them, even at the best; at the worst, slaving many of them till they
perished at their toils, shooting down others under one or other
guilty pretence, and positively sweeping thousands into an untimely
grave. A common cry on their lips was,—
“Let them perish and let the white men occupy these Isles.”
It was such conduct as this, that made the Islanders suspect all
foreigners and hate the white man and seek revenge in robbery and
murder. One Trader, for instance, a sandal-wooder and collector of
Kanakas, living at Port Resolution, abominably ill-used a party of
Natives. They determined in revenge to plunder his store. The cellar
was underneath his house, and he himself slept above the trap-door
by which alone it could be entered. Night and day he was guarded by
armed men, Natives of adjoining islands, and all approaches to his
premises were watched by savage dogs that gave timely warning. He
felt himself secure. But the Tannese actually constructed a tunnel
underground from the bush, through which they rolled away
tobacco, ammunition, etc., and nearly emptied his cellar! My heart
bled to see men so capable and clever thus brutally abused and
demoralized and swept away. By the Gospel, and the civilization
which it brings, they were capable of learning anything and being
trained to a useful and even noble manhood. But all influence that
ever I witnessed from these Traders was degrading, and dead against
the work of our Missions.
The Chief, Nowar Noukamara, usually known as Nowar, was my
best and most-to-be-trusted friend He was one of the nine or ten who
were most favourable to the Mission work, attending the Worship
pretty regularly, conducting it also in their own houses and villages,
and making generally a somewhat unstable profession of
Christianity. One or more of them often accompanied me on
Sabbath, when going to conduct the Worship at inland villages, and
sometimes they protected me from personal injury. This Nowar
influenced the Harbour Chiefs and their people for eight or ten miles
around to get up a great feast in favour of the Worship of Jehovah.
All were personally and specially invited, and it was the largest
Assembly of any kind that I ever witnessed on the Islands.
When all was ready, Nowar sent a party of Chiefs to escort me and
my Aneityumese Teachers to the feast. Fourteen Chiefs, in turn,
made speeches to the assembled multitude; the drift of all being, that
war and fighting be given up on Tanna,—that no more people be
killed by nahak, for witchcraft and sorcery were lies,—that Sacred
Men no longer profess to make wind and rain, famine and plenty,
disease and death,—that the dark heathen talk of Tanna should
cease, that all here present should adopt the Worship of Jehovah as
taught to them by the Missionary and the Aneityumese,—and that all
the banished Tribes should be invited to their own lands to live in
peace! These strange speeches did not draw forth a single opposing
voice. Doubtless these men were in earnest, and had there been one
master mind to rule and mould them, their regeneration had
dawned. Though for the moment a feeling of friendliness prevailed,
the Tannese were unstable as water and easily swayed one way or the
other. They are born talkers, and can and will speechify on all
occasions, but most of it means nothing, bears no fruit.
After these speeches, a scene followed which gradually assumed
shape as an idolatrous ceremonial and greatly horrified me. It was in
connection with the immense quantity of food that had been
prepared for the feast, especially pigs and fowls. A great heap had
been piled up for each Tribe represented, and a handsome portion
also set apart for the Missionary and his Teachers. The ceremony was
this, as nearly as I could follow it. One hundred or so of the leading
men marched into the large cleared space in the centre of the
assembled multitudes, and stood there facing each other in equal
lines, with a man at either end closing up the passage between. At the
middle they stood eight or ten feet apart, gradually nearing till they
almost met at either end. Amid tremendous silence for a few
moments all stood hushed; then every man kneeled on his right
knee, extended his right hand, and bent forward till his face nearly
touched the ground. Thereon the man at the one end began
muttering something, his voice rising ever louder as he rose to his
feet, when it ended in a fearful yell as he stood erect. Next the two
long lines of men, all in a body, went through the same ceremonial,
rising gradually to their feet, with mutterings deepening into a howl,
and heightening into a yell as they stood erect. Finally, the man at
the other end went through the same hideous forms. All this was
thrice deliberately repeated, each time with growing frenzy. And
then, all standing on their feet, they united as with one voice in what
sounded like music running mad up and down the scale, closing with
a long, deep-toned, hollow howl as of souls in pain. With smiles of
joy, the men then all shook hands with each other. Nowar and
another Chief briefly spoke, and the food was then divided and
exchanged, a principal man of each Tribe standing by to receive and
watch his portion.
At this stage, Nowar and Nerwangi, as leaders, addressed the
Teachers and the Missionary to this effect:—
“This feast is held to move all the Chiefs and People here to give up
fighting, to become friends, and to worship your Jehovah God. We
wish you to remain, and to teach us all good conduct. As an evidence
of our sincerity, and of our love, we have prepared this pile of food
for you.”
In reply, I addressed the whole multitude, saying how pleased I
was with their speeches and with the resolutions and promises which
they all had made. I further urged them to stick fast by these, and
that grand fruits would arise to their island, to themselves and to
their children.
Having finished a brief address, I then walked forward to the very
middle of the circle, and laid down before them a bundle of stripes of
red calico and pieces of white calico, a number of fish-hooks, knives,
etc. etc., requesting the two Chiefs to divide my offering of goodwill
among the Tribes assembled, and also the pile of food presented to
us, as a token of my love and friendship to them all.
Their insisting upon me taking their present of food, laid upon me
an unpleasant and dangerous necessity of explaining my refusal. I
again thanked them very warmly, and explained that, as they had in
my presence given away all their food to an Idol God and asked his
blessing on it as a sacrifice, even to Karapanamun, the great Evil
Spirit, my people and I durst not and could not eat of it, for that
would be to have fellowship with their Idols and to dishonour
Jehovah God. Christians could acknowledge only the one true and
living God, and ask His blessing on their food, and offer it and
themselves in thanksgiving unto Him, but unto no cruel or evil
Spirit. Yet I explained to them how much I thanked them, and how I
loved them just as much as if we had eaten all their gifts, and how it
would please us to see them all, along with my own gifts, divided
amongst their Tribes.
Not without some doubt, and under considerable trial, did I take
this apparently unfriendly attitude. But I feared to seem even to
approve of any act of devil-worship, or to confirm them in it, being
there to discourage all such scenes, and to lead them to acknowledge
only the true God. I felt as if guilty and as if the hat were rising from
my head, when I heard them imprecating and appeasing their God,
without being able to show them the God of Love and the better way
into His presence through Jesus Christ. My opportunity to do so
arose over the refusal of the food offered unto Idols, and I told them
of the claims of Jehovah, the jealous God, who would not share His
worship with any other. But all the time I felt this qualm,—that it
were better to eat food with men who acknowledged some God and
asked his blessing than with those white Heathens at home, who
asked the blessing of no God, nor thanked Him, in this worse than
the dog which licks the hand that feeds it! Nowar and Nerwangi
explained in great orations what I meant, and how I wished all to be
divided amongst the assembled Tribes to show my love. With this, all
seemed highly satisfied.
Heathen dances were now entered upon, their paint and feathers
and ornaments adding to the wildness of the scene. The men seemed
to dance in an inside ring, and the women in an outside ring, at a
considerable distance from each other. Music was supplied by
singing and clapping of hands. The order was perfect, and the figures
highly intricate. But I have never been able to associate dancing with
things lovely and of good report! After the dancing, all retired to the
bush, and a kind of sham fight followed on the public cleared ground.
A host of painted savages rushed in and took possession with songs
and shoutings. From the bush, on the opposite side, the chanting of
women was heard in the distance, louder and louder as they
approached. Snatching from a burning fire flaming sticks, they
rushed on the men with these, beating them and throwing burning
pieces of wood among them, till with deafening yells amongst
themselves and amidst shouts of laughter from the crowd, they drove
them from the space, and danced thereon and sang a song of victory.
The dancing and fighting, the naked painted figures, and the
constant yells and shoutings gave one a weird sensation, and
suggested strange ideas of Hell broken loose.
The final scene approached, when the men assisted their women to
fill all the allotted food into baskets, to be carried home and eaten
there; for the different Tribes do not sit down together and eat
together as we would do; their coming together is for the purpose of
exchanging and dividing the food presented. And now they broke
into friendly confusion, and freely walked about mingling with each
other; and a kind of savage rehearsal of Jonathan and David took
place. They stripped themselves of their fantastic dresses, their
handsomely woven and twisted grass skirts, leaf skirts, grass and leaf
aprons; they gave away or exchanged all these, and their ornaments
and bows and arrows, besides their less romantic calico and print
dresses more recently acquired. The effusion and ceremonial of the
gifts and exchanges seemed to betoken a loving people; and so they
were for the feast—but that laid not aside a single deadly feud, and
streams of blood and cries of hate would soon efface all traces of this
day.
I had now six Stations, opened up and ministered to by
Aneityumese Teachers, at the leading villages along the coast, and
forming links in a chain towards the other Mission Establishment on
Tanna. And there were villages prepared to receive as many more.
These Teachers had all been cannibals once, yet, with one exception,
they proved themselves to the best of my judgment to be a band of
faithful and devoted followers of Christ. Their names were Abraham,
Kowari, Nomuri, Nerwa, Lazarus, and Eoufati. I visited them
periodically and frequently, encouraging and guiding them, as well
as trying to interest the villagers in their teaching and work. But,
whenever war broke out they had all to return to the Mission House,
and sleep there for safety by night, visiting their Stations, if
practicable, by the light of day. My poor dear Teachers, too, had to
bear persecutions for Jesu’s sake, as the following incident will
sorrowfully prove.
A native woman, with some murderous purpose in her heart,
pretended great friendship to the excellent wife of one of my fellow-
labourers. She was specially effusive in bringing to her dishes of food
from time to time. Having thus gained confidence, she caught a little
black fish of those parts, known to be deadly poisonous, and baked it
up in a mess for the unsuspecting Teacher’s wife. On returning, she
boasted of what she had done, and thereon a friendly neighbour
rushed off to warn the other, but arrived just to learn that the fatal
meal had been taken. Beyond all reach of human skill, this unknown
martyr for Christ died soon after in great agony, and doubtless
received her Master’s reward.
 In helping to open up new Stations, those dear native Teachers
often bore the greatest hardships and indignities with a noble self-
denial and positively wonderful patience. Nothing known to men
under Heaven could have produced their new character and
disposition, except only the grace of God in Christ Jesus. Though still
marred by many of the faults of Heathenism, they were at the roots
of their being literally new creatures, trying, according to their best
light, to live for and to please their new Master, Jesus Christ. This
shone out very conspicuously in these two apostolic souls, Abraham
and Kowari, as leaders among all the devoted band.
Let me recall another occasion, on which I prevented a war. Early
one morning, the savage yells of warring Tribes woke me from sleep.
They had broken into a quarrel about a woman, and were fiercely
engaged with their clubs. According to my custom, I rushed in
amongst them, and, not without much difficulty, was blessed in
separating them before deadly wounds had been given or received.
On this occasion, the Chiefs of both Tribes, being very friendly to me,
drove their people back from each other at my earnest appeals.
Sitting down at length within earshot, they had it out in a wild
scolding match, a contest of lung and tongue. Meanwhile I rested on
a canoe midway betwixt them, in the hope of averting a renewal of
hostilities. By-and-by an old Sacred Man, a Chief called Sapa, with
some touch of savage comedy in his breast, volunteered an episode
which restored good humour to the scene. Leaping up, he came
dancing and singing towards me, and there, to the amusement of all,
re-enacted the quarrel, and mimicked rather cleverly my attempt at
separating the combatants. Smashing at the canoe with his club, he
yelled and knocked down imaginary enemies; then, rushing first at
one party and then at the other, he represented me as appealing and
gesticulating and pushing them afar from each other, till he became
quite exhausted. Thereon he came and planted himself in great glee
beside me, and looked around as if to say,—“You must laugh, for I
have played.” At this very juncture, a loud cry of “Sail O!” broke upon
our ears, and all parties leapt to their feet, and prepared for a new
sensation; for in those climes, everything—war itself—is a smaller
interest than a vessel from the Great Unknown World sailing into
your Harbour.
 Not many days thereafter, a very horrible transaction occurred.
Before daybreak, I heard shot after shot quickly discharged in the
Harbour. One of my Teachers came running, and cried,—
“Missi, six or seven men have been shot dead this morning for a
great feast. It is to reconcile Tribes that have been at war, and to
allow a banished Tribe to return in peace.”
I learned that the leading men had in council agreed upon this
sacrifice, but the name of each victim was kept a secret till the last
moment. The torture of suspense and uncertainty seemed to be
borne by all as part of their appointed lot, nor did they prepare as if
suspecting any dread assault. Before daylight, the Sacred Men
allocated a murderer to the door of each house where a victim slept.
A signal shot was fired; all rushed to their doors, and the doomed
ones were shot and clubbed to death as they attempted to escape.
Their bodies were then borne to a sacred tree, and hung up there by
the hands for a time, as an offering to the Gods. Being taken down,
they were carried ceremoniously and laid out on the shore near my
house, placed under a special guard.
Information had reached me that my Teachers and I were also
destined victims for this same feast, and sure enough we espied a
band of armed men, the killers, despatched towards our premises.
Instantaneously I had the Teachers and their wives and myself
securely locked into the Mission House; and, cut off from all human
hope, we set ourselves to pray to our dear Lord Jesus, either Himself
to protect us or to take us to His glory. All through that morning and
forenoon we heard them tramp-tramping round our house,
whispering to each other, and hovering near window and door. They
knew that there were a double-barrelled fowling-piece and a revolver
on the premises, though they never had seen me use them, and that
may, under God, have held them back in dread. But such a thought
did not enter our souls even in that awful time. I had gone to save,
and not to destroy. It would be easier for me at any time to die than
to kill one of them. Our safety lay in our appeal to that blessed Lord
who had placed us there, and to whom all power had been given in
Heaven and on Earth. He that was with us was more than all that
could be against us. This is strength; this is peace:—to feel, in
entering on every day, that all its duties and trials have been
committed to the Lord Jesus,—that, come what may, He will use us
for His own glory and our real good!
All through that dreadful morning, and far into the afternoon, we
thus abode together, feeling conscious that we were united to this
dear Lord Jesus, and we had sweet communion with Him,
meditating on the wonders of His person and the hopes and glories
of His kingdom. Oh, that all my readers may learn something of this
in their own experience of the Lord! I can wish them nothing more
precious. Towards sundown, constrained by the Invisible One, they
withdrew from our Mission House, and left us once more in peace.
They bore away the slain to be cooked, and distributed amongst the
Tribes, and eaten in their feast of reconciliation; a covenant sealed in
blood, and soon, alas, to be buried in blood again! For many days
thereafter, we had to take unusual care, and not unduly expose
ourselves to danger; for dark characters were seen prowling about in
the bush near at hand, and we knew that our life was the prize. We
took what care we could, and God the Lord did the rest, or rather He
did all—for His wisdom guided us, and His power baffled them.
Shortly thereafter, war was again declared by the Inland people
attacking our Harbour people. It was an old quarrel; and the war was
renewed and continued, long after the cause thereof had passed
away. Going amongst them every day, I did my utmost to stop
hostilities, setting the evils of war before them, and pleading with the
leading men to renounce it. Thereon arose a characteristic incident
of Island and Heathen life. One day I held a Service in the village
where morning after morning their Tribes assembled, and declared
that if they would believe in and follow the Jehovah God, He would
deliver them from all their enemies and lead them into a happy life.
There were present three Sacred Men, Chiefs, of whom the whole
population lived in terror,—brothers or cousins, heroes of traditional
feats, professors of sorcery, and claiming the power of life and death,
health and sickness, rain and drought, according to their will. On
hearing me, these three stood up and declared they did not believe in
Jehovah, nor did they need His help, for they had the power to kill
my life by Nahak (i.e., sorcery or witchcraft), if only they could get
possession of any piece of the fruit or food that I had eaten. This was
an essential condition of their black art; hence the peel of a banana
or an orange, and every broken scrap of food, is gathered up by the
Natives, lest it should fall into the hands of the Sacred Men, and be
used for Nahak. This superstition was the cause of most of the
bloodshed and terror upon Tanna; and being thus challenged, I
asked God’s help, and determined to strike a blow against it. A
woman was standing near with a bunch of native fruit in her hand,
like our plums, called quonquore. I asked her to be pleased to give
me some; and she, holding out a bunch, said,—
“Take freely what you will!”
Calling the attention of all the Assembly to what I was doing, I took
three fruits from the bunch, and taking a bite out of each, I gave
them one after another to the three Sacred Men, and deliberately
said in the hearing of all,—
“You have seen me eat of this fruit, you have seen me give the
remainder to your Sacred Men; they have said they can kill me by
Nahak, but I challenge them to do it if they can, without arrow or
spear, club or musket, for I deny that they have any power against
me or against any one by their Sorcery.”
The challenge was accepted; the Natives looked terror-struck at
the position in which I was placed! The ceremony of Nahak was
usually performed in secret,—the Tannese fleeing in dread, as
Europeans would from the touch of the plague; but I lingered and
eagerly watched their ritual. As the three Chiefs arose, and drew near
to one of the Sacred Trees, to begin their ceremonial, the Natives fled
in terror, crying,—
“Missi, away! Alas, Missi!”
But I held on at my post of observation. Amidst wavings and
incantations, they rolled up the pieces of the fruit from which I had
eaten, in certain leaves of this Sacred Tree into a shape like a waxen
candle; then they kindled a sacred fire near the root, and continued
their mutterings, gradually burning a little more and a little more of
the candle-shaped things, wheeling them round their heads, blowing
upon them with their breaths, waving them in the air, and glancing
wildly at me as if expecting my sudden destruction. Wondering
whether after all they did not believe their own lie, for they seemed to
be in dead earnest, I, more eager than ever to break the chains of
such vile superstition, urged them again and again, crying,—
 “Be quick! Stir up your Gods to help you! I am not killed yet; I am
perfectly well!”
At last they stood up and said,—
“We must delay till we have called all our Sacred Men. We will kill
Missi before his next Sabbath comes round. Let all watch, for he will
soon die and that without fail.”
I replied, “Very good! I challenge all your Priests to unite and kill
me by Sorcery or Nahak. If on Sabbath next I come again to your
village in health, you will all admit that your Gods have no power
over me, and that I am protected by the true and living Jehovah
God!”
For every day throughout the remainder of that week, the Conchs
were sounded, and over that side of the island all their Sacred Men
were at work trying to kill me by their arts. Now and again
messengers arrived from every quarter of the island, inquiring
anxiously after my health, and wondering if I was not feeling sick,
and great excitement prevailed amongst the poor deluded idolaters.
Sabbath dawned upon me peacefully, and I went to that village in
more than my usual health and strength. Large numbers assembled,
and when I appeared they looked at each other in terror, as if it could
not really be I, myself, still spared and well. Entering into the public
ground, I saluted them to this effect,—
“My love to you all, my friends! I have come again to talk to you
about the Jehovah God and His Worship.”
The three Sacred Men, on being asked, admitted that they had
tried to kill me by Nahak, but had failed; and on being questioned,
why they had failed, they gave the acute and subtle reply, that I also
was myself a Sacred Man, and that my God being the stronger had
protected me from their Gods. Addressing the multitude, I answered
thus,—
“Yea, truly; my Jehovah God is stronger than your Gods. He
protected me, and helped me; for He is the only living and true God,
the only God that can hear or answer any prayer from the children of
men. Your Gods cannot hear prayers, but my God can and will hear
and answer you, if you will give heart and life to Him, and love and
serve Him only. This is my God, and He is also your friend if you will
hear and follow His voice.”
 Having said this, I sat down on the trunk of a fallen tree, and
addressed them,—
“Come and sit down all around me, and I will talk to you about the
love and mercy of my God, and teach you how to worship and please
Him.”
Two of the Sacred Men then sat down, and all the people gathered
round and seated themselves very quietly. I tried to present to them
ideas of sin, and of salvation through Jesus Christ, as revealed to us
in the Holy Scriptures.
The third Sacred Man, the highest in rank, a man of great stature
and uncommon strength, had meantime gone off for his warrior’s
spear, and returned brandishing it in the air and poising it at me. I
said to the people,—
“Of course he can kill me with his spear, but he undertook to kill
me by Nahak or Sorcery, and promised not to use against me any
weapons of war; and if you let him kill me now, you will kill your
friend, one who lives among you and only tries to do you good, as you
all know so well. I know that if you kill me thus, my God will be angry
and will punish you.”
Thereon I seated myself calmly in the midst of the crowd, while he
leaped about in rage, scolding his brothers and all who were present
for listening to me. The other Sacred Men, however, took my side,
and, as many of the people also were friendly to me and stood closely
packed around me, he did not throw his spear. To allay the tumult
and obviate further bloodshed, I offered to leave with my Teachers at
once, and, in doing so, I ardently pled with them to live at peace.
Though we got safely home, that old Sacred Man seemed still to
hunger after my blood. For weeks thereafter, go where I would, he
would suddenly appear on the path behind me, poising in his right
hand that same Goliath spear. God only kept it from being thrown,
and I, using every lawful precaution, had all the same to attend to my
work, as if no enemy were there, leaving all other results in the hands
of Jesus. This whole incident did, doubtless, shake the prejudices of
many as to Sorcery; but few even of converted Natives ever get
entirely clear of the dread of Nahak.
If not truly converted, the two Priests were fast friends of mine
from that day, as also another leading man in the same district. They