Lewin Benjamin (ed.) Genes IX

nation at relatively

high frequencies (compared

to strains that lack integrated

F factors);

such

strains are described

as Hfr

(for

high

frequency

recombination).

Each

position

of

integration

for the F factor

gives

rise

to a different

Hfr strain,

with

a characteristic

pattern

of transferring bac-

terial markers

to a recipient

chromosome.

Contact between

conjugating

bacteria is

usually

broken before transfer

of DNA is

com-

plete.

As a result,

the

probability

that a region

of the bacterial

chromosome

will be transferred

depends upon its distance

from

oriT. Bacterial

genes

located

close to the

site of F integration

(in

the direction of transfer)

enter recipient

bac-

teria first,

and are therefore

found at

greater

frequencies than those

that are located

farther

away and

enter later. This

gives

rise to a

gradi-

ent of transfer frequencies

around

the chro-

mosome, declining

from the

position

of F

integration. Marker

positions

on the donor

chro-

mosome can

be

assayed

in terms

of the time at

which transfer occurs; this

gave

rise to the

stan-

dard description

of

the

E. coli chromosome

as a

map divided into

100 minutes.

The map refers

to transfer times from a

particular

Hfr

strain;

the starting

point

for

the

gradient

of transfer is

different for each Hfr

strain because it is

deter-

mined by the

site where the F factor

has inte-

grated

into

the bacterial

genome.

DONOR

BACTERIUM

F FACTOR

griT

fra region

RECIPIENT

BACTERIUM

F factor is

nicked

at orif

I

T

I

\f

...'.'l

: :.-.- :._::..',i'

|

5, end

leads single strand

:

----

|

intoreciPient

Singie strands are converted

to double

strands in both bacteria

Sl{i;*itil 1 *.i.i Transfer of chromosomal

DNA occurs

when an inte-

grated

F factor is njcked at onJi

Transfer

of DNA starts

with a short

sequence of F DNA and contjnues

until

prevented

by loss of con-

tact between the bacteria.

survival

of

the

parasite

and causes

the

plant

cell

to

grow

as a tumor.

Agrobacteria are

required

to induce tumor

formation, but the

tumor

cells do

not require the

continued

presence

of bacteria.

As with

animal

tumors,

the

plant

cells

have been

transformed

into a state

in which new

mechanisms

govern

growth

and differentiation.

Tlansformation

is

caused by the

expression

within

the

plant

cell

of

genetic

information

transferred

from the

bacterium.

The tumor-inducing

principle of. Agrobac-

terium resides in the

fi

plasmid,

which

is

per-

petuated

as an independent

replicon

within

the

bacterium.

The

plasmid

carries

genes

involved

in various bacterial

and

plant

cell activities,

including those

required

to

generate

the

trans-

formed state, and

a set of

genes concerned

with

synthesis

or utilization

of

opines

(novel

deriv-

atives of arginine).

16.8 The

BacteriaI

Ti Ptasmid

Causes Crown

Ga[[

Disease

in P[ants

The Bacterial

Ti Plasmid

Causes Crown

Ga[[ Disease

in PLants

o

Infection with

the bacterium A. tumefociens

can

transform

ptant

cetls into tumors.

o

The infectious agent is

a

ptasmid

carried

by the

ba cte ri u m.

o

The

plasmid

atso carries

genes

for

synthesizing

and metabotizing

opines

(arginine

derivatives)

that are used

by the tumor cett.

Most events in which DNA

is rearranged

or

amplified occur within

a

genome,

but

the inter-

action between bacteria and

certain

plants

involves the transfer

of

DNA

from the bacter-

ial

genome

to the

plant genome.

Crown

gall

disease, shown in

trI*i.JR{

1$.'i3,

can be induced

in most dicotyledonous

plants

by the

soil bac-

Ierium Agr o b acterium tumefaciens.

The bacterium

is

a

parasite

that effects a

genetic

change in the

eukaryotic host cell, with

consequences lor both

parasite

and host: It improves

conditions for

401

ir:;.;i;iiL

i r.1

.j An

Agrobacteium

carrying a

Ti

plasmid

of

the nopatine

type

induces

a teratoma,

jn

whjch

djfferen-

t'iated structures

devetop. Photo

courtesy of the estate of

Jeff Schett.

Used with

oermission of the

Max P[anck Insti-

tute for Ptant

Breeding Research,

Cologne.

i-ii:l-:{i

r i=-.:.i;

Ti

ptasmids

carry

genes

involved in

both

p[ant

and

bacterial

functions.

Ti

plasmids (and

thus tlire Agrobacteria

in

which

they reside)

can be

divided into four

groups,

according

to the types

of opine that are

made:

.

Nopaline

plasmids

carry

genes

for

syn-

thesizing nopaline

in

tumors and for

utilizing it in

bacteria.

Nopaline tumors

can

differentiate into

shoots

with abnor-

mal

structures. They

have

been called

teratomas

by analogy

with certain

mammalian

tumors

that retain the

abil-

ity

to differentiate

into

early embryonic

structures.

.

Octopine

plasmids

are similar

to

nopa-

line

plasmids,

but the relevant

opine is

different.

Octopine

tumors are

usually

CHAPTER

1.6 Extrachromosomal

Reoticons

undifferentiated, however,

and do

not

form

teratoma shoots.

.

Agropine

plasmids

carry

genes

for

agropine metabolism; the tumors

do not

differentiate, and they develop

poorly

and die early.

.

Ri

plasmids

can

induce

hairy root dis-

ease on some

plants

and crown

gall

on

others. They have agropine

type

genes,

and may

have

segments derived from

both nopaline and octopine

plasmids.

The

types of

genes

carried by a Ti

plasmid

are summarized in

fl'i*lifit t*.i'i.

Genes utilized

in

the bacterium code

for

plasmid

replication

and incompatibility, transfer

between bacteria,

sensitivity to

phages,

and synthesis

of other

compounds, some of which are

toxic to other

soil bacteria. Genes used

in

the

plant

cell code

for

transfer of

DNA into

the

plant,

induction

of

the transformed state, and shoot

and root

induction.

The specificity of the opine

genes

depends

on the type of

plasmid.

Genes needed for

opine

synthesis are linked to

genes

whose

products

catabolize the same opine; thus each strain of

Agrobacterium

causes crown

gall

tumor

cells to

synthesize opines that are useful for

survival of

the

parasite.

The

opines can be used

as the sole

carbon and/or nitrogen

source for the induc-

ing Agrobacterium strain The

principle

is that

the transformed

plant

cell synthesizes those

opines that the bacterium can use.

T-DNA

Carries Genes

Required

for Infection

.

Part

of the

DNA

of the

Ti

plasmid

is

transferred to

the olant ce[[ nucleus.

r

The vrr

genes

of the

Ti

ptasmid

are

located outside

the

transferred

region

and are

required

for

the

transfer

process.

o

The vir

genes

are induced by

phenotic

compounds

reteased

by

plants

in response

to

wounding.

o

The membrane

protein

VirA is autophosphorytated

on

histidine

when it binds an inducer.

r

VirA activates VirG

by transferring the

phosphate

group

to it.

r

The VirA-VirG is one of

several bacterial two

component systems that

use a

phosphohistidine

reray.

The interaction

between Agrobacterium

and

a

plant

cell is illustrated

in

E{#liSf;

i*"1h.

The

bac-

terium does not

enter the

plant

cell.

but rather

transfers

part

of the Ti

plasmid

to the

plant

Locus Function

Ti

Plasmid

vtr

snt

rot

nos

noc

ocs

occ

DNA

transfer into

plant

all

shoot

induction

all

root

induction

all

nopaline

synthesis

nopaline

catabolism

nopaline

octopine

synthesis

octopine

catabolism

octopine

tra

bacterial transfer

genes

all

lnc incompatibility

genes

all

oriV

origin for replication

all

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