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3',

5',

Linear DNA

Fork oroceeds

3',

DNA synthesis initiates at left 5' end

5

strand is disolaced

when tork reaches

end

I

Termini base

pair

to form duplex origin

$ISUi{il 1S,f

Adenovirus DNA replication is

initiated

sep-

arately

at the two ends

of the motecute and

proceeds

by

strand

disp[acement.

adenovirus

DNA.

This

suggests

the model

illus-

trated

in Ft*Llft*

1*.d;.

The complex

of

poly-

merase

and terminal

protein,

bearing the

priming

C

nucleotide, binds to

the end of the

adenovirus

DNA. The free 3'-OH end

of the C

nucleotide

is used to

prime

the elongation

reac-

tion

by the

DNA

polymerase.

This

generates

a

new strand

whose 5' end

is

covalently

linked

to the

initiating

C nucleotide.

(The

reaction

actually

involves displacement

of

protein

from

DNA

rather than binding

de novo.The 5' end of

adenovirus

DNA is bound to the

terminal

pro-

tein that

was used

in the

cycle.

The old

terminal

protein

is

displaced

by

the

new terminal

protein

for each

new repli-

cation cycle.)

Terminal

protein

binds to the

region located

between

9 and

l8 bp from the end of

the DNA.

The adjacent

region, between

positions l7 and

48,

is essential

for the binding of a

host

protein,

nuclear

factor I, which

is also required

for the

flG#ft* t*.3 The 5'terminaI

phosphate

at

each end

of

adenovirus

DNA is covalenttv

tinked

to

serine

in the 55

kD

Ad-binding

protein.

Serine g

Hrr

lpl;

,zN

rlr'"r

Polypeptide

Y

PolYPePtide

CHz

Protein-DNA

link4l

Y

$;Stlftil i*"si

Adenovirus

terminaI

protein

binds to

the 5'

end

of DNA and

provides

a

C-OH

end to

prime

synthesis

of a new DNA

strand.

initiation

reaction.

The

initiation

complex

may

therefore

form

between

positions 9 and

48, a

fixed distance

from

the

actual

end

of

the

DNA.

16.3

TerminaI

Proteins

Enab[e

Initiation

at

the

Ends

of Virat

DNAs

@

RolLing

Circles Produce

Multimers

of

a Replicon

.

A rotling

c'ircte

generates

singte-stranded

muttimers

of

the originaI sequence.

The

structures

generated

by replication

depend

on the relationship

between

the template

and

the replication

fork. The

critical

features

are

whether

the template

is circular

or linear,

and

whether

the replication

fork

is engaged

in syn-

thesizing

both

strands

of DNA

or only one.

Replication

of

only one

strand is

used to

generate

copies

of some

circular

molecules.

A

nick

opens

one strand,

and

then the free

3'-OH

end

generated

by the nick

is extended

by

the

DNA

polymerase.

The newly

synthesized

strand

Template

is circular

duplex DNA

Initiation

occurs on one

strand

,,^

,/ \\

/ \\

3',-OH

il ,5,-p

Nick

at orisin

\\ ./

V

Elongation

of

growing

strand

displaces

old strand

-

Growing

strand

Displaced

strand

After

one revolution

displaced

strand

reaches

unit length

Continued

elongation

generates

displaced

strand of multiple

unit

i:ii:t"iHI

"tj.."t

The rotting

circle

generates

a multjmeric

single-stranded

tait.

CHAPTER

16

ExtrachromosomaI

Repticons

displaces the

original

parental

strand. The

ensu-

ing

events are

depicted in i*3fi["1ft{

t$.*.

This

type of structure

is called

a rolling

cir-

cle, because

the

growing

point

can be

envis-

aged

as rolling

around the

circular

template

strand. It

could in

principle

continue

to

do so

indefinitely.

As it moves,

the replication

fork

extends

the outer

strand and

displaces

the

pre-

vious

partner.

An example

is shown

in

the elec-

tron micrograph

of

Fl$;#Ril

i$"f'.

The newly

synthesized

material

is

cova-

lently linked

to the

original

material,

and as

a

result

the displaced

strand has

the

original

unit

genome

at

its

5' end. The

original

unit

is fol-

lowed

by any number

of unit

genomes,

syn-

thesized

by continuing

revolutions

of

the

template.

Each revolution

displaces

the

mate-

rial

synthesized

in the

The rolling

circle is

put

to several

uses

in

vivo.

Some

pathways

that

are used

to replicate

DNA are

depicted in Filii.jH&

tii.l.

Cleavage

of a unit length

tail

generates

a

copy of

the original

circular replicon

in linear

form. The

linear form

may

be maintained

as a

single

strand,

or may be

converted

into

a duplex

by synthesis

of the

complementary

strand

(which

is identical

in sequence

to the

template

strand

of the original

rolling

circle).

The

rolling

circle

provides

a means

for

amplifying

the

original

(unit)

replicon.

This

mechanism

is

used

to

generate

amplified

ribo-

somal DNA

(rDNA)

inthe

Xenopzs

oocyte.

The

genes

for ribosomal

RNA

(rRNA)

are

organized

as a large

number

of contiguous

repeats

in the

genome.

A

single repeating

unit from

the

genome

is converted

into

a rolling

circle.

The

dis-

placed

tail, which

contains

many

units,

is con-

verted

into

duplex

DNA; later

it is

cleaved

from

l:.1{;iJFI

:$"*

A rotting

circle

appears

as

a circutar

mo[e-

cu[e with

a [inear

taiI by electron

microscopy.

Courtesy

of

Ross B.

Inman, Institute

of Molecutar

Virotogy,

Bock

Lab-

oratory and Department

of Biochemistry,

University

of

Wisconsin.

Madison.

Wisconsin,

USA.

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Lewin Benjamin (ed.) Genes IX

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