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OH

OH

Base

O

O

O—

O—

O—

O—

—O—P--—O—P—O—P—O—H

2

C

5'

1'

4'

2'

3'

— —

O

— —

O

— —

Nucleoside

Nucleoside monophosphate

Nucleoside diphosphate

Nucleoside triphosphate

Nucleotides

A

B

C

Base

Nucleotide

(Base+sugar)

Nucleotide

(Base+sugar+phosphate)

Nucleic Acid

Purines

Adenine

Guanine

Cytosine

Thymine

Uracil

Pyrimidines

Adenosine
Deoxyadenosine

Guanosine
Deoxyguanosine

Cytidine
Deoxycytidine

Thymidine
Deoxythymidine

Uridine
Deoxyuridine

AMP
dAMP

GMP
dGMP

CMP
dCMP

TMP
dTMP

UMP
dUMP

ADP
dADP

GDP
dGDP

CDP
dCDP

TDP
dTDP

UDP
dUDP

ATP
dATP

GTP
dGTP

CTP
dCTP

TTP
dTTP

UTP
dUTP

RNA
DNA

RNA
DNA

RNA
DNA

RNA
DNA

RNA
DNA

Monophosphate

Diphosphate

Triphosphate

D

FIGURE 2-1. Biochemistry of DNA. (A)

Structure of the biochemical components of DNA and RNA (purines, pyrimidines,

sugars, and phosphate). (B) Diagram depicting the chemical structure of the various components of DNA. (C) Diagram of
a DNA polynucleotide chain. The biochemical components (purines, pyrimidines, sugar, and phosphate) form a polynu-
cleotide chain through a 3

,5-phosphodiester bond. (D) Nomenclature of nucleosides and nucleotides in RNA and DNA.

AMP 

 adenylate or adenosine 5-monophosphate dAMP  2-deoxyadenosine 5-monophosphate ADP  adenosine

5

-diphosphate dADP  2-deoxyadenosine 5-diphosphate ATP  adenosine 5-triphosphate dATP  2-deoxyadeno-

sine 5

-triphosphate GMP  guanylate or guanosine 5-monophosphate dGMP  2-deoxyguanosine 5-monophosphate

GDP 

 guanosine 5-diphosphate dGDP  2-deoxyguanosine 5-diphosphate GTP  guanosine 5-triphosphate dGTP 

2

-deoxyguanosine 5-triphosphate CMP  cytidylate or cytidine 5-monophosphate dCMP  2-deoxycytidine 5-

monophosphate CDP 

 cytidine 5-diphosphate dCDP  2-deoxycytidine 5-diphosphate CTP  cytidine 5-triphosphate

dCTP 

 2-deoxycytidine 5-triphosphate TMP  thymidylate or thymidine 5-monophosphate dTMP  2-deoxythymi-

dine 5

-monophosphate TDP  thymidine 5-diphosphate dTDP  2-deoxythymidine 5-diphosphate TTP- thymidine 5-

triphosphate dTTP 

 2-deoxythymidine 5-triphosphate UMP  uridylate or uridine 5-monophosphate dUMP  2-

deoxyuridine 5

-monophosphate UDP  uridine 5-diphosphate dUDP  2-deoxyuridine 5-diphosphate UTP  uridine

5

-triphosphate dUTP  2-deoxyuridine 5-triphosphate.

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16

BRS Genetics

1

1

2

B

A

C

μ

m

8,800-fold

Double helix DNA

30-nm fiber

μ

m

μ

m

FIGURE 2-2. DNA packaging. (A)

Diagram depicting the various levels of packaging of double helix DNA found within a

metaphase chromosome. Double helix DNA is wound around a histone octamer of H2A, H2B, H3, and H4 to form a nucle-
osome. Nucleosomes are pulled together by histone H1 to form a 30 nm diameter fiber. The 30 nm fiber exists either as
extended chromatin or as secondary loops within a condensed metaphase chromosome. (B) Electron micrograph of DNA
isolated and subjected to treatments that unfold DNA to a nucleosome. The “beads on a string” appearance is the basic
unit of chromatin packaging called a nucleosome. The globular structure (“bead”) (arrow 1) is a histone octamer. The lin-
ear structure (“string”) (arrow 2) is DNA. (C) Compaction of DNA in a chromosome. The double helix DNA of a chromo-
some is shown unraveled and stretched out measuring 88,000 um in length. During metaphase of mitosis, chromatin can
become highly compacted. For example, human chromosome 1 contains about 260,000,000 bp. The distance between
each base pair is 0.34  nm. So that, the physical length of the DNA comprising chromosome 1 is 88,000,000 nm or 88,000 um
(260,000,000 X 0.34nm 

 88,000,000 nm). During metaphase, all the chromosomes condense such that the physical length

of chromosome 1 is about 10 um. Consequently, the 88,000 um of DNA comprising chromosome 1 is reduced to 10 um,
resulting in an 8,800-fold compaction.

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1.

In humans, the female is functionally

hemizygous due to X chromosome inactiva-
tion. The inactivated chromosome is thus
composed of which of the following?

(A)

satellite 1 DNA

(B)

beta-satellite DNA

(C)

facultative heterochromatin

(D)

constitutive heterochromatin

(E)

euchromatin

2.

The levels of DNA packaging are depicted

in which of the following sequences?

(A)

alpha satellite DNA, heterochromatin,
centromere, euchromatin, metaphase
chromosome

(B)

constitutive heterochromatin, euchro-
matin, facultative heterochromatin,
metaphase chromosome

(C)

purines, pyrimidines, phosphates, nucle-
osome, 30 nm chromatin fiber,
metaphase chromosome

(D)

double helix DNA, nucleosome, 30 nm
chromatin fiber, extended chromatin,
metaphase chromosome

(E)

double helix DNA, histones, nucleo-
somes, extended chromatin, metaphase
chromosome

3.

The nitrogenous bases that make up the

nucleotides of DNA are listed in which one of
the following?

(A)

deoxyribose and ribose

(B)

deoxyribose, ribose, and phosphate

(C)

adenine, thymine, cytosine, uracil

(D)

adenine, thymine, cytosine, guanine

4.

Which one of the following is a major

component of centromeric DNA?

(A)

the Barr body

(B)

the XY body

(C)

alpha satellite DNA

(D)

Z-DNA 

5.

Which one of the following is the way

bases are paired in a double helix of DNA?

(A)

A-T, G-C

(B)

A-U, G-C

(C)

A-G, C-T

(D)

A-C, G-T

17

Review Test

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18

1. The answer is (C).

Both copies of the X chromosome in females are active only for a short

time early in development. 

2. The answer is (E).

Double helix DNA is coiled around histones that are organized into

nucleosomes, which form the extended chromatin that compacts into a metaphase chro-
mosome. 

3. The answer is (D).

A DNA nucleotide consists of one of the nitrogenous bases adenine,

thymine, cytosine or guanine, the sugar deoxyribose, and a phosphate group. Uracil and the
sugar ribose are components of RNA nucleotides. 

4. The answer is (C).

The 171 base pair repeat unit of alpha satellite DNA makes up much of

the centromeric DNA. 

5. The answer is (A).

In DNA, adenine (A) pairs with thymine (T) and guanine (G) pairs with

cytosine (C). In RNA, adenine pairs with uracil (U).

Answers and Explanations

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c h a p t e r

3

Chromosome Replication

19

I. GENERAL FEATURES

A.

Chromosome replication occurs during 

S phase

of the cell cycle and involves both DNA syn-

thesis and histone synthesis to form chromatin. 

B.

The timing of replication is related to the chromatin structure. An 

inactive gene

packaged as 

het-

erochromatin

is replicated 

late in S phase

(e.g., in a female mammalian cell, the inactive X chro-

mosome called the 

Barr body

is packaged as heterochromatin and is replicated late in S phase).

C.

An 

active gene

packaged as 

euchromatin 

is replicated early in S phase (e.g., in the pancreatic

beta cell, the insulin gene will be replicated early in S phase. However, in other cell types (e.g.,
hepatocytes) where the insulin gene is inactive, it will be replicated late in S phase. 

D.

DNA polymerases absolutely require the 

3

-OH end

of a based paired primer strand as a sub-

strate for strand extension. Therefore, a 

RNA primer

(synthesized by a 

primase

) is required to

provide the free 3

-OH group needed to start DNA synthesis. 

E.

DNA polymerases copy a DNA template in the 

3

S

S  

5

 direction,

which produces a new DNA

strand in the 

5

S

S

3

 direction

.

F. Deoxyribonucleoside 5

-triphosphates  (dATP, dTTP, dGTP, dCTP)

pair with the corresponding

base (A-T, G-C) on the template strand and form a 

3

,5-phosphodiester bond

with the 

3

-OH

group on the deoxyribose sugar, 

which releases a 

pyrophosphate

.

G.

Replication is described as 

semiconservative

which means that a molecule of double helix

DNA contains one intact parental DNA strand and one newly synthesized DNA strand.

II. THE REPLICATION PROCESS 

(Figure 3-1)

A.

The process starts when 

topoisomerase

nicks (or breaks) a single strand of DNA, which

causes DNA unwinding. 

B.

Chromosome replication begins at specific nucleotide sequences located throughout the
chromosome called 

replication origins

.

Eukaryotic DNA contains 

multiple replication origins

to ensure rapid DNA synthesis. Normally, the S phase of the mammalian cell cycle is 

8 hours

.

C. DNA helicase

recognizes the replication origin and opens up the double helix at that site,

forming a 

replication bubble 

with a 

replication fork

at each end. The stability of the replication

fork is maintained by 

single-stranded binding proteins

.

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