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Genetics and
Molecular Biology
DNA Structure
19
and Function
After you have finished reading this chapter, you should be able to:
Describe the basic structure of a chromosome, the DNA molecule, and
the nucleotide subunits.
List the four nitrogenous bases and explain how they are paired.
Discuss the process of DNA replication and explain how errors in
replication may cause mutations.
Introduction
Deoxyribonucleic acid, or DNA, has been called the molecule of life. Sci-
entists have known of DNA’s existence for about 100 years, but they have
come to understand it only since the early 1950s. In the relatively short
time since then, scientists have learned that DNA contains the informa-
tion on which all life depends.
During the lifetimes of our parents and our grandparents, scientists
discovered the secret of what was quietly, invisibly making life possible on
this planet for the past 3.5 billion years. DNA is the master molecule that
controls all of life. Indeed, life as we know it would not exist without
DNA. You will learn about it in this chapter.
403
404 Genetics and Molecular Biology
fur, size of ears, number of legs or wings, height, and countless other
characteristics. (See Figure 19-1.) By the early 1920s, scientists knew that
chromosomes were made up of two substances, DNA and proteins. The
question asked from 1923 on was: Is the genetic material—the sub-
stance that determined the traits in all living things—the DNA or the
protein?
DNA, actually nucleic acid, had first been identified in 1868 by a Swiss
physician, Johann Friedrich Miescher. Nucleic acid was thought to be a
simple molecule that contained only four different parts. On the other
hand, many different kinds of proteins were known to exist. Proteins are
made up of an almost infinite number of combinations of the 20 differ-
ent amino acids. Because of the great number of different proteins, many
scientists thought proteins were the genetic material—the source of the
great diversity of traits—found in living things.
The search for the substance that was responsible for determining
traits in living things began. Some interesting experiments, mostly using
bacteria and viruses, were performed. One set of experiments by a British
researcher, Frederick Griffith, found that a substance from dead pneu-
monia bacteria could change or transform harmless pneumonia bacteria
into pneumonia-causing ones. He called the substance a “transforming
factor.” (See Figure 19-2a.) Oswald Avery, at Rockefeller University in New
York City, conducted very careful chemical analyses on the transforming
factor. In 1944, Avery concluded that DNA was the transforming factor.
At that time, the importance of Avery’s discovery remained unrecognized.
(See Figure 19-2b.)
It took one more set of crucial experiments to demonstrate that DNA
was the substance that determined which traits were inherited. In 1952,
Chapter 19 / DNA Structure and Function 405
Capsule
Live pneumococcus
(1)
Died of pneumonia
Live pneumococcus
Figure 19-2a Griffith’s
experiment. A substance
(2) from dead, coated,
pneumonia-causing
Live pneumococcus No pneumonia bacteria could transform
Encapsulated live, uncoated, harmless
dead pneumococcus
pneumonia bacteria into
(3) coated, harmful,
pneumonia-causing
Died of pneumonia bacteria.
Alfred Hershey and Martha Chase, working at the Cold Spring Harbor
Laboratories on Long Island in New York, used bacteria and the viruses
that infect bacteria in a now-famous experiment. Hershey and Chase
LIVING ENVIRONMENT BIOLOGY, 2e/fig. 19-2a s/s (rev. 10/22/03)
knew that the viruses they used reproduced by injecting viral genetic
material into bacterial cells; the rest of the virus—consisting of its protein
coat—remained attached to the outside of the bacteria. The viruses then
Pneumococci Pneumococci
with capsules without capsules
(from “smooth” (from “rough” colony)
colony)
Cytoplasm and
DNA of dead Live pneumococci
“smooth”
pneumococci
After mixing
Transformed
live pneumococci
with capsules
Virus
Bacterium
(a) (b)
Figure 19-3a Viral DNA is injected Figure 19-3b The bacterium bursts and
into a bacterium, where it will direct releases the newly made viruses. The
the production of new viruses. The new viruses contain the labeled DNA,
LIVING ENVIRONMENT BIOLOGY, 2e/fig. 19-3 s/s
virus’s protein coat remains outside the not the labeled protein.
bacterium.
used the material in the infected bacterial cells to make more viruses. (See
Figures 19-3a and 19-3b.)
Hershey and Chase prepared two kinds of viruses. One kind of virus
had a special radioactive marker on its DNA. The other kind of virus had
a different radioactive marker on its proteins. Hershey and Chase infected
bacterial cells with the two kinds of marked viruses. Later they examined
the inside of the bacterial cells to see what molecules—the protein or the
DNA—had entered the bacteria from the viruses as the genetic material.
The answer was clear. It was the specially marked viral DNA. Hershey and
Chase concluded, correctly, that the genetic material in the viruses was
DNA. Except for a few unusual viruses, we now know that genetic instruc-
tions are carried in DNA in all living organisms on Earth.
◆ The genetic material must be strong and stable so it does not easily fall
apart, causing perhaps harmful changes to its store of information.
Scientists are always looking for interesting projects to work on. In 1953,
James Watson, a young researcher from the United States, went to Cam-
bridge University in England. He joined Francis Crick, a physicist, because
both of them were interested in learning more about DNA. Few other peo-
ple were interested in studying DNA at that time. Many researchers sus-
pected that trying to decipher the structure of DNA would turn out to be
very dull work. And they thought that it would not be at all clear how
DNA functions as the genetic material.
To learn more about the structure of DNA, Watson and Crick did not
work the way most scientists do. They did not perform a long series of
experiments in a laboratory. Instead, they used data already known about
DNA molecules and made models of molecules with structures to fit the
data. (See Figure 19-4 on page 408.)
It was known that DNA was made up of smaller nucleotide subunits.
Each nucleotide consists of a five-carbon sugar called deoxyribose, a phos-
phate, and a nitrogenous base. A chemist at Columbia University, Erwin
Chargaff, had discovered that the four types of nitrogenous bases in DNA
actually formed two pairs. The amounts of adenine (A) and thymine (T)
were always the same (A = T). The amounts of guanine (G) and cytosine
(C) were always the same (G = C). Another critical piece of information
that Watson and Crick had was from pictures taken when X rays were
shot through crystals of DNA. Called X-ray diffraction patterns, these pic-
tures had been taken by Rosalind Franklin and Maurice Wilkins. It was
determined from their pictures that the DNA molecule had a spiral, or
Figure 19-4
James Watson
(left) and Francis
Crick shown in
1953 with their
model of part of a
DNA molecule.
helical, pattern—a pattern that looks like a spiral staircase or the threads
of a wood screw. (See Figure 19-5.)
With this information, based largely on the work of other scientists,
Watson and Crick used tin and wire to build a large model. Their model
of DNA, in the shape of a double helix, rather than being dull, was about
the most exciting discovery in the history of biology.
The easiest way to understand the double helix structure of DNA is to
picture a ladder that has been twisted. The two sides of the ladder are par-
allel to each other. The steps of the ladder link the two sides to each other.
P Bond
P
D A T
D
P P
D T A
D
P P
D G C
D
P P
Figure 19-6 The structure
D C G D of DNA.
In this model, the sides of the ladder are the sugar-phosphate back-
bone of the DNA molecule. Alternating deoxyribose sugar and phosphate
LIVING ENVIRONMENT BIOLOGY, 2e/fig. 19-6 s/s
molecules make up this backbone. Stretching between the two sides, from
a sugar on one side to a sugar on the other, are pairs of molecules that con-
tain nitrogen. These are the nitrogenous bases. One sugar, one phosphate,
and one nitrogenous base make up a nucleotide. The nucleotides on one
side of the molecule are bonded together to make one side, or strand, of
the molecule. Other nucleotides are bonded together to make up the
other side of the DNA molecule. (See Figure 19-6.)
Finally, how are the two sides of the molecule joined? The answer to
this involves the nitrogenous base pairs. The Watson-Crick model showed
that the only possible way all the parts could fit was for each large adenine
base to be matched opposite a smaller thymine base. Similarly, a larger
guanine had to be opposite a smaller cytosine. These nitrogenous bases
were joined by fairly weak hydrogen bonds between them. It is now pos-
sible to realize the importance of Chargaff’s discovery, A = T and G = C.
So a molecule of DNA consists of two strands, opposite each other,
connected by matching nitrogenous base pairs. If we look at one strand,
we can describe it in terms of the order or sequence of nucleotides. A DNA
molecule may be very long. Because the nucleotides are in a long line,
the order of the nucleotides is called a linear sequence. Imagine walking
along a single strand of DNA. The bases in the nucleotides may occur in
any order. The linear sequence on a short molecule of DNA might be
A-T-T-G-A-C-C-G. Now imagine walking along the opposite strand start-
ing at the same place. Opposite the A in the first strand is a T. Because we
know the sequence of nucleotides in the first strand, we automatically
know the sequence of nucleotides in the other strand. In this example,
beginning with the T, it must be T-A-A-C-T-G-G-C. This is the key to how
410 Genetics and Molecular Biology
Kangaroo
Rabbit
Pig
Donkey Figure 19-8 Evolutionary relationships for DNA
Horse
closeness. The more similar their nucleotide sequences,
Dog
Monkey the more recently the two organisms evolved from a
Human common ancestor.
The DNA match was made possible because Arizona now has a database
that contains a DNA profile of every prison inmate. In fact, every state in
America now has such a database; and a national system, the National DNA
Index System (NDIS), was started in 1998. By 2002, the one-millionth DNA
profile had been entered into the computerized system. DNA evidence
collected from any crime scene can now be quickly compared to that of
any one of the million convicted offenders in the NDIS database. The
system is quickly growing and the technology of DNA testing is rapidly
improving. For example, a portable DNA testing kit is under development
in Britain. It will be smaller than a suitcase and will be linked to the national
DNA database of that country. It is expected that the crime scene evidence
will be put in a solution and then placed inside the mobile unit. Silicon chip
technology in the testing kit will then extract a DNA profile that will be sent
to the national database via a laptop computer. The results may be
returned in under an hour to the detective’s palm-held computer. Saliva on
discarded cigarette butts at crime scenes has already been used successfully
to provide DNA profiles of suspects.
It is hoped that someday, thanks to this kind of technology, there will be no
more wrongful convictions such as that of Mr. Krone, and more positive
identifications of those who do deserve the jail time.
are made of proteins and DNA. Chromosomes are packages of DNA that
seem to be held together by proteins. Why do organisms need packages
of DNA?
Consider that DNA molecules are very, very long. A typical cell in the
human body is much smaller than the period at the end of this sentence,
and yet that single cell contains more than 2 meters of DNA. In addition,
for DNA to do its job correctly, it cannot get tangled like a long piece of
string thrown carelessly into a drawer. Chromosomes help maintain the
DNA in the proper shape, untangled and ready for use. In a chromosome,
the long double-helix DNA molecules get wound around protein mole-
cules to form bundles. These bundles get looped together, and the loops,
in turn, get coiled and folded together. This all works well to squeeze DNA
into a very tight space and yet keep it well organized in order to do its job.
Old Old
A T
T A
A T
G
G C
G C
T A
C G
T
A T
G C
C G
A T
T
T A
G C
A T
C G
C G C G
C G C G
New New
G G
T A T A
A T A T
T A T A
Figure 19-9 During DNA replication, the T A T A
G G
double helix unwinds, the DNA strands
T A T A
separate, and new complementary DNA
A T A T
strands form opposite each of the original
T A T A
strands. Old New New Old
of the cell cycle. What we already know about its structure is enough to
explain how DNA replicates. (See Figure 19-9.)
LIVING ENVIRONMENT BIOLOGY, 2e/fig. 19-9 s/s
To make a copy, you need an original, sometimes called a template.
Because DNA is a double helix, it has templates built into it. To begin the
process, the double helix unwinds. As with all metabolic activities,
enzymes are needed for this process. Once the double-stranded molecule
is untwisted, it begins to unzip, just like a zipper. Recall that the nitrogen
base pairs A-T and G-C are connected by weak hydrogen bonds. However,
at the correct moment, through the activity of an enzyme, these bonds
begin to break apart. As the hydrogen bonds break, each strand of the
DNA molecule becomes separate. Many free nucleotides float around in
the cell. Specific enzymes match up these free nucleotides with the exist-
ing nucleotides in each DNA strand. Wherever a T is located on a strand,
an A pairs to it; wherever a C is located, a G joins up, and so on. One by
one, new nucleotides are joined together to make a new strand opposite
Chapter 19 / DNA Structure and Function 415
G C G C
T A T A
A T A T
A T A T
G C G C
C G C G
a. b.
DNA polymerase
Free nucleotides
C
G
in cytoplasm
G
C
A
A A
T
T
T
T
A
A
C
A T
G
G
G C
G
C
c.
G C G C
T A T A
A T A T
A T A T
Figure 19-10 Through the
G C G C
process of DNA replication,
C G C G identical double-stranded
d. DNA molecules are formed.
In life, nothing is always perfect. That is true about DNA replication, too.
The enzymes responsible for directing the correct pairing of nucleotides
during DNA replication occasionally make mistakes. A nitrogenous base
416 Genetics and Molecular Biology
may be left out. Or the wrong base may be matched up. Sometimes an
extra one is added. These mistakes produce errors in the linear sequence
in one strand of the DNA molecule. Such an error is called a mutation.
From what we know about the replication process, once an error
occurs in a DNA strand, it may be copied again and again. The mutation
in the genetic material in one cell can easily be passed on to future cells.
Are these mutations good or bad? It seems a strange question to ask,
because we assume that mistakes are always bad. However, what is obvi-
ous is not always true. A mutation is simply a change. Many changes in
the genetic material are harmful. In fact, many of these changes make it
impossible for the future cells, or even the entire organism, to continue
living. Other mutations cause the organism to change in an unnoticeable
fashion. Rather than harming the organism, the mutation seems to pro-
duce no effect. Sometimes the mutation gives the organism a sudden new
advantage that other similar organisms lack.
Let’s consider a simple example. Imagine that all grasshoppers in a
green field were brown in color. Birds could easily see the grasshoppers
and eat them. Then a mutation occurred in the DNA of one grasshopper.
When that grasshopper reproduced—passing on its genetic mutation—
the offspring with the mutation were green, not brown. Is this a good
mutation or a bad one? Being a green grasshopper in a green field is good
if it makes it harder for birds to see and eat you. This color change (muta-
tion) to green would provide a survival advantage over the more easily
seen brown grasshoppers. Natural selection would make it more likely
that the green grasshoppers would survive. The species would evolve in
terms of body color.
Not only can mutations in DNA be good, but they are actually an
important source for the genetic variations necessary for natural selec-
tion to occur. Much of the evolution of life on Earth has depended on
the chance occurrence of these mutations.
INTRODUCTION
In 1953, Francis Crick and James Watson discovered the structure of the
DNA molecule. This discovery enabled other researchers to begin to deter-
mine how molecules of DNA function. Like many other polymers, a
strand of DNA is made of many individual connected subunits. A mole-
cule of DNA consists of two parallel strands. The subunits on one strand
are connected to the subunits on the other strand. It is the connections
between the two strands, which can be broken and reformed, that enable
DNA to make a copy of, or replicate, itself. In this laboratory investigation,
you will use paper “models” of DNA subunits and portions of DNA
strands to learn how this remarkable chemical is constructed and can be
copied.
MATERIALS
Diagrams for steps 1–4 (from Teacher’s Manual), scissors, blank paper, tape
PROCEDURE
Step 1. Cut out the six nucleotides in the handout. Assemble them into
a double strand of DNA that consists of three pairs of nucleotides
that are joined together. (Hint: The phosphates attached to the
sugars point up on one strand and down on the other strand. Bio-
chemists say the two strands are “antiparallel.”) Tape the pieces
into place on a sheet of paper.
Step 2. Cut out the four nucleotides from the second handout. Match
them up correctly according to rules for nitrogenous-base pair-
ing. Remember that the hydrogen bonds (dotted lines) coming
from one base must match up with the hydrogen bonds in its
partner. Tape the pieces into place on a sheet of paper.
Step 3. Refer to the diagram for step 3. Use the rules that govern the pair-
ing of bases. Write the complementary base on the unlettered
strand that will bond with the base shown on the lettered strand.
INTERPRETIVE QUESTIONS
1. From the results in step 2, determine the two reasons why each base
can pair with only one of the three types of nitrogenous bases.
2. From the results of all four steps, write a paragraph that describes how
DNA is a well-organized molecule. Explain why this organization
makes it possible for DNA to be the genetic material for all organisms.
■■ CHAPTER 19 REVIEW
VOCABULARY
The following list contains all of the boldfaced terms in this chapter. Define
each of these terms in your own words.
DNA of virus
Not radioactive
Bacterium
Radioactive
16. Identify the scientists who performed the experiment shown in the
diagram.
LIVING ENVIRONMENT BIOLOGY, 2e/fig. 19-Q16 s/s
17. What were the results of this experiment? What did the scientists
conclude from these results?
18. What four characteristics must the substance that serves as the
genetic material have?
19. Explain why mutations can have positive or negative effects.
20. Create a time line that shows important events in our
understanding of DNA.
422 Genetics and Molecular Biology