Lewin Benjamin (ed.) Genes IX

Loss of

plasmid

leaves killer and

antidote

Cell cycle of one

plasmid

t:

ir':!

I

Cell

grows

and

Y

plasmid

replicates

I

Cell divides

Each

cell

has copy

o{ same

plasmid

Cell

cycle of

incompatible

plasmids

Cell

grows

but

r olasmids do

not

I

replicate because

two origins are

already

Present

T

J

Cell

divides

V

lncompatible

plasmids

have

been

drslributed

to ditferenl

ceils

li:..r.iiri

: .;.::

"

Ptasmids may

ensure that bacteria can-

not live without them by synthesizing

a [ong-tived

kitter

and a short-lived antidote.

P[asmid Incom

patibi

Lity

Is Determined

by

the Replicon

.

Plasmids in

a singte compatibility

group

have

origins that

are regulated

by a common control

syste

m.

The

phenomenon

of

plasmid

incompatibility

is related to the regulation

of

plasmid

copy

number and segregation. A

compatibility

group

is defined as a

set of

plasmids

whose

members are unable to coexist in the same

bacterial cell.

The reason for

their

incompat-

ibility is that they cannot be distinguished

from one another at some stage that is essen-

tial

for

plasmid

maintenance. DNA replica-

tion and segregation are stages at which this

may apply.

The negative control model for

plasmid

incompatibility follows the idea

that

copy num-

ber control

is achieved

by synthesizing

a repres-

;:iirijiii: r.'

i.:

Two

ptasmjds

are

incompatib[e

(they

betong

to the same

compatibility

group)

if

their

origins

cannot

be distinguished

at the

stage

of

initiation. The same

modeI could

appty to segregation.

sor that measures

the

concentration

of origins.

(Formally,

this is the

same

as the titration

model

for regulating

replication

of the bacterial

chromosome.)

The

introduction

of a

new origin

in the

form of a second

plasmid

of the

same compat-

ibility

group

mimics

the

result of

replication

of

the resident

plasmid; two origins

now are

present.

Thus any

further

replication

is

pre-

vented until after

the

two

plasmids have been

segregated to

different

cells

to create

the cor-

rect

prereplication copy

number,

as illustrated

in

i:.111.11!ii-,

I

:.

i

:.

A

similar

effect

would

be

produced if the

system for segregating

the

products to daugh-

ter cells

could not

distinguish

between

two

plas-

mids.

For example,

if two

plasmids have the

same cis-acting

partition sites,

competition

between them

would

ensure

that

they would

be segregated

to different

cells,

and therefore

could not survive

in

the same

line.

The

presence of a

member

of one

compat-

ibility

group

does

not directly

affect

the survival

of a

plasmid

belonging

to

a different

group.

Only

one replicon

of a

given

compatibility

group (of

a single-copy

plasmid)

can

be

maintained

in the

bacterium,

but

it does

not

interact

with

repli-

cons

of other

compatibility

groups.

17.10

Ptasmid

Incompatibitity

Is Determined

by the

Replicon

@

The

CoLEI Compatibility

System Is

ControL[ed

by

an RNA Regulator

.

Reptication

of

ColEl.

requires

transcription

to

pass

through the

origin, where the

transcript

is

cteaved

by RNAase H

to

generate

a

primer

end.

r

The regutator

RNA I is

a short

antisense RNA that

pairs

with

the transcript

and

prevents

the cteavage

that

generates

the

priming

end.

o

The Rom

protein

enhances

pairing

between

RNA

I

and

the transcriot.

The

best characterized

copy number

and incom-

patibility

system is

that

of

the

plasmid

ColEl,

a

multicopy

plasmid

that is

maintained

at a steady

Ievel

of

-20

copies

per

E. coli cell. The

system for

maintaining

the copy number

depends

on the

mechanism

for initiating

replication

at the ColEl

origin, as illustrated

in

ilil*;:l=li:

I l;.ii*.

Replication

starts

with

the transcription

of

an RNA

that initiates 555

bp upstream

of the

origin. Transcription

continues

through the ori-

gin.

The enzyme

RNAase

H

(whose

name

reflects

its specificity

for a

substrate of RNA

f:gi#i.Jfii:

:t'i'.1i}

Repl"ication

of Co[E1 DNA

is initiated

by

cteaving

the

primer

RNA to

generate

a

3'-0H end. The

primer

forms

a

persistent

hybrid

in the

origin region.

BacteriaI

Reptication Is

Connected

to

the Ce[[ Cycte

hybridized

with

DNA)

cleaves the transcript

at

the origin.

This

generates

a l'-OH

end that is

used as the

"primer"

at which DNA synthesis

is

initiated

(the

use of

primers

is discussed

in more

detail

in

Section 18.8, Priming Is

Required to

Start DNA Synthesis). The

primer

RNA forms

a

persistent

hybrid

with the DNA. Pairing

between

the RNA and DNA

occurs

just

upstream

of the

origin

(around position

-20)

and also farther

upstream

(around

position

-265).

TWo regulatory

systems exert their

effects

on the RNA

primer.

One involves

synthesis

of

an

RNA

complementary to the

primer;

the

other

involves a

protein

coded by a nearby

locus.

The regulatory

species

RNA

I is

a molecule

of

-108

bases and is coded by the

opposite

strand

from

that specifying

primer

RNA.

The relation-

ship between the

primer

RNA and RNA

I is illus-

trated in l3Gtifril 1.:.:$.

The RNA

I molecule is

initiated within

the

primer

region

and termi-

nates

close to the site

where the

primer

RNA

initiates.

Thus RNA I is

complementary

to the

5'-terminal

region

of the

primer

RNA.

Base

pairing

between

the two RNAs

controls

the

availability

of the

primer

RNA

to initiate

a cycle

of replication.

An RNA molecule

such as RNA

I that func-

tions

by virtue of its complementarity

with

another RNA

coded in the

same region is

called

a

countertranscript. This

type of mechanism,

of course, is another

example

of the use of

anti-

sense RNA

(see

Section 13.7,

Small RNA

Mol-

ecules Can Regulate

Translation).

Mutations

that reduce

or eliminate incom-

patibility

between

plasmids

can be obtained

by

selecting

plasmids

of the same

group

for their

ability

to coexist. Incompatibility

mutations

in

ColEl map in the region

of

overlap between

RNA I and

primer

RNA. This

region is

repre-

sented in

two different RNAs,

so

either or both

might

be involved in

the effect.

When RNA I is

added to a

system for repli-

cating ColEl

DNA invitro, it

inhibirs

the for-

mation

of active

primer

RNA. The

presence

of

RNA I, however,

does not inhibit

the

initiati<,rn

F3#f-jgqfl

1?,J$ The

sequence of RNA I is

comp[ementary

to the

5'

region

of

primer

RNA.

422

CHAPTER 17

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