..

2002

1.
. ................................................ 1
1.1 ...................................................................................... 2
lac ....................................................................................................................................................................... 2
1.2 ? .............................................................................................................. 3
.......................................................................................................... 3
1.3 - ........................................................... 4
1.4 ............................................................................................................................................. 6
1.5 ....................................................................................................................................... 7
. ................................................................................................................................................................. 8
1.6 . .......................................................................... 9
2. .......................................................................... 10
...................................................................................................................................... 13
bgl .............................................................................................................. 14
3. - - .................................. 14

- . . ............................................... 16
rpoS ................................................................................................................................................................................. 16
rpoE ................................................................................................................................................................................ 17
4. , ............................................ 19
....................................................................................................................................... 19
- .................................................................................................................................... 21
5. ......................................................................................................... 24
6. ............................................................................................... 26
................................................................................................................................ 27
.................................................................................. 29
. ........................................................................................................................ 30
HPt- ................................................................................................................................................................. 30
...................................................................................................................................................... 30
...................................................................................................................................................... 30
...................................................................................................... 30
..................................................................................................................................... 30
. .............................................................................................................................................. 31
.............................................................................................................................................. 31
. ........................................................................................................................ 31
.................................................................................................................... 32
...................................................................................................................... 32
....................................................................................................................................... 33
. ................................................................................................................................... 33
................................................................................................................. 34
..................................................................................................................................... 34
7. ................................................................................................................ 34
7.1 ....................................... 34
.................................................................................................. 35
7.2 ? ................................................................ 36
7.3 .................................................................................................................. 37

- II -

 ....................................................................................................................................... 37
? ...................................... 38
...................... 39
. ....................................................................................... 39
8. .................................................................................................... 41
9. . ..................................................... 43
: ............................ 43
. .................................................................................................................... 43
. ..................................................................................... 44
. ................................................................................................................. 44
SoxRS ............................................................................................................................................................. 44
OxyR ............................................................................................................................................................... 45
. ...................................................................................................................................... 45
FNR .................................................................................................................................... 45
ArcB ................................................................................................................................................................................ 46
10. .................................... 47
CAULOBACTER CRESCENTUS. ........................................................................ 59
11. BACILLUS SUBTILIS. ........................................................................ 62
........................................................................................................................................ 62
? ....................................................................................... 63
...................................................... 64
12. . QUORUM SENSING..................................... 66
. ............................................................................................................................................................ 67
Erwinia ............................................................................................... 68
- . - (
). ................................................................................................................................................................. 69
13. ..................................................................................................... 69
. ................................................................................................................. 71
(GSP). .................................................................................................... 72
Sec . ............................................................................................................................................................... 72
GSP ................................................................................................................. 74
GSP ............................................................................................................................................................ 75
III ............................................................................................................................... 76
1.1. III . ......................................................... 76
1.2. III . ............................................................................................. 77
1.3. , III ................. 78
. ....................................................................................................................................... 79
IV ............................................................................................................................... 80
14. .......................................................................................................... 82
15. SOS ..................................................................................................................... 83
SOS- ........................................................................................................................................................... 84
16. ........................................................................ 84
, ................................................................ 86
...................................................................................................................................................................... 86
III .................................................................................................................................................................... 86

- III -

 ............................................................................................................................... 86
II ..................................................................................................................................................................... 87
. ............................................................................................ 87
- ...................................................................................................................... 87

. ............................................................................................................... 88

- IV -

1. .

, , ,
, , , .
, ,
, ,
, , -
.
,
.
, ,
, .
, . , , - .

, . -
, . , , ,
. ,
, .
,

, .
,
,
.., .
, ,
, -
.

( , ),
,
.
, ( ),
( ).
- (, , ).
, ( , ,
, .,
- ).
.
: ( ):
I- ( ) - cell cycle;
II :
) (+ )
) ( )
III :
) (rpoS)
) (cold shock)
IV : -
; ()

-1-

A priori ,

. ,
(, ,
). .
1.1
,
Escherichia coli.
1961 . "". , ,
( , .. ,
), ,
.
,
. , (
) ,
.
. :
1. - , -
. - TTGACA - 17 bp - TATAAT.
( , - )
.
2. Operator - . - 20 bp.
.
, , , , ,
. ()
.
. (, )
, ,
. ""
,
.
3. . , .
, ,
.
4. . .
, -,
, . -
, .
- .
, , -,
, .
-
(-) .
. . .,
:
( )
( )
lac
lacZ -
lacY -

-2-

lacA -
-
- 1000 .
- - lacI, - -.
. lacI .
, .
-
. (, )
.
, .

. , lac-
. lac-
CAP (Catabolite Activator Protein, CRP) ().
. CAP
, .
CAP-cAMP , .
, , ,
( .. ) .
, , CAP-cAMP,
.

1.2 ?

, ,
- .
.
-
- , "" .
, . ( ,
). -
, ( ) ""
.
1. . . (). ,
, , .
.
2. . - .
-,
.., . ,
N- "", ,
. .., ,
- ,
. - ,
.
. -
,

-3-

 , .
. , ,

.

( ).
.
?
-
- ,
. :
- ,
;
(
)
.
E. coli , ,
, ( ). cI .
.
, -
.
- ( , )
.
Lac

10
4000 JH Miller'a
Gal

LacI, - ( LacI)
Gal
HU .
?

1.3 -
,
-. 10 35 2 4 -. , .
- - -.
, , 10 ,
2 - -.
,
-,
.
- ,
, (
).
.
-.
-;
, .

-4-

 ,
.
RNAP
(a)
(. 1.1.)
a NTD
b/ b'
.
70

- -
a CTD
- (CTD)
-35
-10

.
35 . ,
(b)

a NTD
.
b/ b'
CTD. 70
s
AA
a CTD
,
. N-
-35
-10

-

, CTD

(c)
, .
a CTD
a NTD
b/ b'
, ,
, -.
s 70
AA
Region
4
,
61 ( lac ),
-35
-10
71, -81, -91.
CTD ?
(d)
, a NTD
b/ b'
, ,
s 70
AA

Region 4
a CTD
. ?
-35
-10
CTD -
(cI) ,
cI-
. 1.1.

-
cI cI.
cI
CTD, ,
, .
, 70- -
, 4. cI .
, 35 PRM.

( 70 ).
, -

-. ,
( ) -.
N- - ,
35 . DnaA PR
' -.
, -.

-5-

 , 35 , CTD
.
CTD . ,
, FNR ( ).
- -,
CTD. , FNR
-. -
, 35. ,
N- - -.
CTD.
, .
MerR Tn501.
, 10 35 .
, -,
10 35 . SoxR ( ,
). ,
.
. .
, , ( ) .
.
4 :
( ).
, ( );

, -. MalT
malK : - MalT
-.
35 , upstream, CTD.
.
Promoter occlusion
1.4
ara E. coli.
lac . lac
-,
LacI , ,
. ara ( araC)
, .
6 (araC).
, araBAD, araC. araC
araBAD, 147
. , araE araF ,
araCBAD, .. . ,
, ..
araC. . ,
() , , ,
, .
, araC
, . ,
, () ,

-6-

 . araC Crep Cind.

C araC araB. ,
, , araC
araBAD.

,
- . Crep
araC, Crep
. , araC
, .
()
, araBAD.
araC ,
, , araBAD.
.., lac .
Crep araC , LacI
lac . , , .
-
. , ,
, - Cind. Cind
, , , , . Cind
- , - ,
-, .
? , ara
. .
(, -72...50 lac -50...-23 gal ),
-. .
ara -107...-78.
- . , , .
. . ,

- . ,
gal lac -, ara
. ,
. -
,
, - , ,
-35 . ,
,
. ,
, , .
1.5

(lac ara), ,
(). ,
. , ,
-
. , -
,
() . .

-7-

 5 , ,
. (1 .190).
,
- .
, -.
trp trpR.
-TrpR . (
- TrpR vs trpR).
70 . ( 1000-
lac ).
- -, , -, ,
.
.

, .
, - . , ,
, .
, .
, ,
,
. .
- , ""
, .
. ,
, 162 .. trpE .
.
- GC , 8
U ( , ). ~90%
( 600-
).

RBS, ,
13 . ? 9 10
. ,
, .
, ,
. 1 2, 3
4. , . 1 2,
2 3. 4 ,
.
,
,
. ,
, ,
. -
1 2, , , 2,
2-3,
. -
-,
2-3. ,
. ,
, 1-2
2-3 , , 3-4.

-8-

 - , , ,
. - thr
thr ile, a ilv - ile, leu val.
, ,
, - - .
1.6 .
,
.
.
, - .
. ,
, .
, .
, , , ,
. , , ,
, ,
, .
- .
: (), .
"", ,
. - pN pQ.
.
. , N cro
"" "" PL PR.
N cro tL1 tR1. pN
- tL1 tR1
.
. , Rho,
, -,
. , ,
, C G,
. Rho -
- NusG.
pN ,
, .
, -
, - . ,
pN , - ,
. nutL nutR ( . N utilisation)
.
, : " ?". pN nut ,
- ,
. , 17 ..
( ). , boxB,
pN .
, boxA,
.
pN , , , BoxB
, -,
. ,
.

-9-

 , pN, ,
, NusA NusB (N utilisation substance).
NusA -, , , NusB ,
boxA. NusB
. , NusG, - ,
Rho -.

2.
: (
, ) - ,
. ,
, .
,
.
, ,
.
.
,
, , , , ,
. , , Treponema pallidum,
, ,
, .
:

(.)

+
(.)

-->

+
(.)

-P
(.)

,
- ().
- -14.7 / (, ),
- -3 /. ,
. , -,
, , -, ,
( ). ,
- ,
( , ).
,
.
, , ,
.

- 10 -

 - I (EI), HPr (histidine protein)

II (EII). EII - ( EIIA, EIIB, EIIC
EIID), , . I
HPr - 64-85 9-10 kDa,

,

. ,
II
,
(

),

EI, HPr, EIIA EIIB.


E. coli

.2.1.
(.2.1.()).

EI
,
His-189. EI HPr His-15. P-HPr ( ) EIIA, (His-90),
Cys-241 EIIBC,
-6-.
? , .

EIIC. C II -
, , - . (
EIIB) .
EIIB ,
. IIB Cys
(
).

.
,
.
, , -
, . ,
.
, ,
.
,
"" .
. , , pts
, , ,
,
.

- 11 -

pts , : ptsH ( HPr), ptsI

(EI), crr (EIIAGlc).
-.
, -
. - (P0 P1) ptsH,
(P2) - crr ( ptsI). P0 P1
. P0
-, P1 - .
, , ,
ptsH. P2 crr.
EIIAGlc
HPr EI. pts EIICBGlc (c P0 ).
, pts - EIICBGlc (?)
- ,
,
.
, EI HPr,
. Enterobacteriaceae , , , , , N-, , , . ,
( ) - ,
, , , , , .
?
, -
, . IIAGlc . IIAGlc
P-HPr IICBGlc
- . IIAGlc P-IIAGlc . IIAGlc
, , ,
, .
.
. , IIAGlc
.
, .
, IIAGlc ,
- ?
, - (EI HPr)
, ""
(.2.1.(b)). ""
, IIAGlc HPr ,
IIAGlc. -
- IIAGlc,
- , -
P-IIAGlc.
EIIAGlc ?
EIIAGlc - . , ,
, - , , .
EIIAGlc , .
EIIAGlc
, , .
- , PEIIAGlc -
(.2.2.).

- 12 -

E. coli
.
,

-
B. subtilis (.
2.3.). ,

HPr.
, HPr B. subtilis
.2.2. Escherichia coli
His-15 I Ser-46 ( , --)
- HPr- ( ptsK). Ser-46
. : , CcpA (catabolite control protein) , , , HPr
.
CcpA

,
cre (catabolite responsive
element). CcpA
LacI- .
cre
,
xyl, gut ( ),
hut ( ), lev (
) .
CcpA cre

,

Ser-46 HPr.

CcpA HPr-S46-P.
.2.3. Bacillus subtilis

-
. LevR, LicT LacT,
, - -.
( BglG E.coli),
(PRD).
HPr EII. HPr-H15-P
PRD , .
HPr Ser-46 HPr- .
( HPr-),
HPr , PRD .

- 13 -

bgl .
bgl E. coli () : bglG,
(), bglF, - bglB,
, -.
bglG bglF. BglF
- . BglF
, BglG, ,
BglG.
bgl ,
. BglG
, . BglG (
) . BglG
. - BglF
BglG; BglF BglG,
.

.
sac genes of B. subtilis
3. - -

- - ('2)
() ,
(. 3.1).
- (
10-20).

-,
,

-,

(. 3.1). ,
,
( , ,
..). -
, .

-
,
, , .
( 38). , -,
rpo (RNA polymerase),
-
( RpoS).
- E. coli
70 kDa 70 RpoD.

-,

,
54 kDa, 54
RpoN.
, ,

- 14 -

.3.1.

 -.
- 70. - ,
, , ,
( ,
, ). 54
,
.
3.1. - Escherichia coli

RpoD
RpoS
RpoH
RpoF
RpoE

613
330
284
239
202

FecI
RpoN

173
477

~1000

~100

~40

~
~5

~5

~15

- E. coli 70 kDa 70 RpoD.

-, ,
54 kDa, 54 RpoN.
, , (. 3.2). -
70. -
, ,
,

).
54

,

.

-,
,
,
-.
RpoD RpoS
( - )

, ,
.3.2. -
,
-.
,
. -

- 15 -

 () - .
- ,
, .

- . .
-,
.
- SPO1 B. subtilis.
. .
-
(55 B. subtilis).
, 28, 33 34. 28, gp28, -.
-, - .
33 34, gp33 gp34, gp28, ,
. , -
.
rpoS
E. coli - s, rpoS. -
70. , , , - in
vivo. S s ,
. -
, , s ..
. , ,
.
, pH,
.
,
, (
- ) (
).
E. coli
s, rpoS .
-
- 70 ( rpoD). , , , . s
30% 70,
s ( 100) s- .
, , .
s
,
. s ,
s . ,
,
, ,
.
, ,
, lac-. s
- s-
.
s ,
. - .

- 16 -

 , s , rpoS.
.
- Hfq (HF-I).
, rpoS .
s ,
. ,
, , pH,
, - s . s
ClpXP RssB. s
, 70. RssB
- ClpXP .
RssB s /, -
.
-, s? ,
.
ftsQAZ .
KatE, KatG Dps,
.
, , , Es E70,
. ,
, Es , E70
, in vitro
. ( Es 35 )
s- ?
H-NS, Lrp, cAMP-CRP, IHF Fis.
.
s-
. ,
, . s
, 70
.
rpoE
, s, -,
- 70. - 70
, "" 70.
( )
- E. coli, E.
ECF (extracytoplasmic factors) -. -

(, ). E , ()
, , , E (
o -, 32), -
.
E ,
. -, , ,
, -
. RseA.
E RseA. RseA
RseB. ,
, RseB ,
RseA. RseA E,

- 17 -

 RseA E, -
(. 3.3). :
fkpA, (
);
degP - ,
;
rpoH - -, ;
rpoE rseABC;
.

E.
E

RseA
RseB,

.
E

. , P.
aeruginosa E (AlgU)
alg,

.
, E. coli,
-
.3.3. RpoE.

, MucA. AlgU
, , ,
.
RseA MucA , - .
- -,
-. E , -
, 28 ( ) F ( B. subtilis).
-
, Salmonella typhimutium.
( ) 28.
28
- FlgM.
, FlgM , 28. 28
,
- , .
, FlgM, FlgM
, - .
:
1. D. Missiakas and S. Raina. The extracytoplasmic function sigma factors: role and regulation. Mol Microbiol
1998 28(6):1059-1066

- 18 -

4. ,

, (
- E.
coli).
..
,
(DnaK, GroEL)
- (Lon, ClpAP).
(
)
.

-
(
) .
. ,

.

-:


.
,
.

(
, ),

.
,
,
.
Hsp70 Hsp60
( heat shock protein + .
)
. Hsp60
E. coli GroEL,
GroES,

55 (. 4.1.).
GroES-GroEL

26S

. 4.1. GroELS

. 4.2. GroE

- 19 -

,
.
E. coli Hsp70 DnaK,

DnaJ GrpE (. 4.3).
E. coli
, SecB, ClpB IbpAB,
,
Hsp70
Hsp60 .

?

,
,
(. 4.4).

,
(. 4.5).
, ,

DnaK-DnaJ-GrpE,

.
,
, ""
. 4.3. DnaK
,
.
, GroES-GroEL.
,
.
.
. , SecB
.
, ,
. ,
DnaK 4-5 , ;
.
36 , .
.
GroES-GroEL
DnaK - , . ,
- ClpAP, ClpXP, HslVU -
, - - . ,
, . HflB (FtsH), ,
. , ,
, . (
, .. - .)
- , .
, ClpAP
, .
, ,

. :

- 20 -

 ,

. , -
,
.

-.

, 5
.
""

.
"" .

- P-
(, ,
-) A- (
).
, ssrA,
-,
, ,
10 ,
.
. 4.4.
, - 11
,
( Tsr,
- HflB ClpXP ). ,
"", , ,
.
-

(26S ),
,
, , (
).

, ,
.

,

() .

. 4.5.
(

). ,

- 21 -

 , .
, ,
, , .
,
5-10 .
E.coli 4 -
.
ClpAP ClpXP.

, ,

(. 4.6). ClpA ClpX

. clpX clpP
, clpA
.
.
ClpYQ/HslUV
HflB(FtsH)
. 4.6. - :
Zn

-
,

,
RpoH, SecY ( ). - ,
E. coli. RpoH- .
Lon.
N ,

SulA,

RcsA .

,
Lon

E. coli. ,
,
.

.

.
,

,

Lon.

.

32
(. 4.7). ,

E
.
, ,

. 4.7. Escherichia
coli

- 22 -

.. - E rpoH, 32,
(, 50C). RpoH,
,
, ,
.
E. coli, 30 42, 32 (
) , .

32
32

rpoH. 32
DnaK-DnaJ-GrpE, - ,
32.
, , HSP
( , , ,
32). DnaK-DnaJ ,
. 32 , ,
- HflB (FtsH). (HslVU,
ClpAP Lon), ,
32. DnaK-DnaJ
, . , ,
32 . -, 32
.
32 -, .
32 ,
. 5' rpoH ,
. ,
, -
.
rpoH , E,
DnaA cAMP-CRP. E. coli
, .
, (32 - ).
DegP (HtrA) (- ) FkpA,
. ,
. , E ,
RseA RseB. RseA -
E , E .
RseB RseA E.
, RseB , RseA,
E. ,
- ( ""),
RseA, E.
.
E. coli,
. B. subtilis
.
( CIRCE/HrcA) DnaKJ-GrpE
GroEL-GroES, HrcA,
dnaK. -
CIRCE. , ,
DnaKJ-GrpE 32 E. coli,
GroEL-GroES. HrcA,
HrcA

- 23 -

 groE dnaK. , GroE,

DnaK .
HrcA CIRCE- ,
-, , ,
.
o (50-100) , B,
. - -
( ).
, .
:
1. Susan Gottesman. Proteases and their targets in Escherichia coli. Annu. Rev. Genet. 1996. 30:465506
2. Susan Gottesman. Regulation by proteolysis: developmental switches. Current Opinion in Microbiology
1999, 2:142147
5.

: 30C
, ..

. , ,

: 10C
,
,

:
,

.
,
:
,

,

: CspA

(
, ,

).

(
), CspA, CsdA, Rbf .
.

CspA CspE

,

,

.

. 5.1.

,

- 24 -

 (cold shock proteins, Csp).

Csp E. coli . ,
37C ,
- , , ,
(<10) .
CspA ( ), CspB, CspG, CsdA,
RbfA, NusA PNP. CspA, CspB, CspG CspI, , CsdA , ""
(), RbfA - , NusA
, PNP - .
RecA, IF-2, H-NS GyrA.
10% , ,
CspA, E. coli. 8 ,
CspA - CspB - CspI, .
CspA, CspB, CspG CspI. ,
.
5' (5' UTR). ,
cspA 11 , "cold box".
- "", cspA.
CspA. - -
,
cspA. , CspA,
CspE CspC, CspE.
(
, , Csp . () Nus .) , CspA
, ,
. , 5' UTR
cspA 37,
. O cspA
( 12 20 )
CspA.
, . (
)
CspA . , cspA , 16S ,
14 .. , "downstream box"
(DB). cspB, cspG, csdA rbfA. ,
15 . DB
16S . , DB
, ,
. ,
RbfA CsdA. , -

, . ,
RbfA CsdA,
.
CspA . CspD
,
RpoS. CspC CspE 37. ,
,
.

- 25 -

 CspA.
-, 5 -.
.
7 ,
.
.
-
. CspA .
:
1. S. Phadtare, J. Alsina and M. Inouye. Cold-shock response and cold-shock proteins. Current Opinion in
Microbiology 1999, 2:175180
2. W. Bae, B. Xia, M. Inouye and K. Severinov. Escherichia coli CspA-family RNA chaperones are
transcription antiterminators. PNAS 2000 97(14):77847789
3. S. Phadtare and M. Inouye. Sequence-selective interactions with RNA by CspB, CspC and CspE, members
of the CspA family of Escherichia coli. Molecular Microbiology (1999) 33(5), 1004-1014
6.
(, , )

.
. -,
,
. :
( ),
;
,
;
- ( ,
).
.
?
.
-, .
, . , PhoQ -
Salmonella typhimurium, - , ,
,
PhoQ.
. Aer, ,
, FAD.
.
, . , Cpx-
E. coli, - ,
. EnvZ E. coli , ,
.
.
- (P)
(C), -

- 26 -

 . TM1
PTM2LC, L -
(EnvZ)
(MCP ).
.
, . , ,
,
, .
( )
.

80- ""
, .
, .
,
.
,
.

(),

H-P
H ATP

ADP
(),

(ATP)

D
D
D-P
.

Pi
(
).

,
. 6.1.

- , , H
.
, D

, :
1. :
-His + -His~ +
2. :
-His~ + -Asp -His + -Asp~
3. :
-Asp~ + H2O -Asp + PI

- 27 -

-
(1).
(2). ,
(3).
( ) ""
Ser/Thr/Tyr , ,
.
G , - (1)
. ,
=> ,
. N~P
. I
II , ,
" " .
. ,
, ..
, .
,
. , ,
.
,
.
.
,
. ,
,
, () ,
.


. ,

, . , Ser/Thr/Tyr .
. , ,
0 Mycoplasma genitalium 2.5% Synechocystis sp.
:
Mycoplasma genitalium 0
Haemophilus influenzae 9
Helicobacter pylori 11
Thermotoga maritima -19
Escherichia coli 62
Bacillus subtilis - 70
Synechocystis sp. 80
--------------Methanococcus jannaschii 0
Methanobacterium thermoautotrophicum - 24
--------------Caenorhabdis elegans 0
Saccharomyces cerevisiae 4 ( )
,
, Dictyostelium discoideum 11 . ,
. , ,

- 28 -

 , (5 30 E. coli),
( ). , ,
,
- .

""
. , ,
""
. , .
- ,
. ,
.
, ,
- Asp
. ,
. ""
.
,



N
H
N D F G
C
,

N
H
N D F G
C
(. 6.2).


N
H
N D F G
C
. (a)

40-200 kDa.
OmpR

5-6

D13 D57 K109

.

FixJ
D13
D57
K109

D13 D57 K109

NtrC

: .
/
CheB/SprE

""
, D13 D57 K109
, EnvZ E. coli, D13 D57 K109
CheY

, (b)
.
EnvZ

. 6.2. (a)
-,
(b).

.

. .
(
4-8) ( CheA NtrB).
(
).
, , ,
.
"" .
"", ""
. ArcB,
. ArcB N-
, , , , ,

- 29 -

 , HPt- ( His-containing phosphotransfer

)
.
,
/-
.
~350
. , ,
.
HPt-
,
. 120 ,
. HPt- ,
, .
, HPt-
,
.

( )
.
,
.
.
,
, .
PAS-.
"" , - ,
.
, .
,
. , , , , .. PAS-
(~100 ) ( N-) , ,
. ,
PAS- .
, , NtrB,
.
(
).


- .
,
. 40-180 .
- coiled coil ,
.

.


,
"" .
.
,

- 30 -

 .
, .
- ,
. ,
.
:
.
- (25 32 E. coli). -
, OmpR, NarL NtrC ( 14, 7
4 ). - .
, , , CheB, .
.
CheY, FliM,
.
.
. ,
Mg2+


. -
. ,
,
, .
OmpR, ,
, , ompC ompF,
. OmpR F C
, . -
" ", ,
, , , ,
.
NarL, , ,
, . NarL
"NarL " HTH .
,
NtrC , 54
-.
- HTH . NtrC
.
.
.
,
.

.
, .
- ( -) .
, -
, .
.

()
.
. .
.

- 31 -

 - . ,
OmpR
, ; NtrC
, .
. ,
.

.
,
, (
NarL) ( CheB,
). N- C-
.

, ,
.
.
( )
, , , ,
.

,
. , ,
, , ,
, .

.
-. ,
, HPt- , ,
( ).
,
( ).

.
.
,
. ,
His Asp, EnvZ-OmpR,
OmpF OmpC. ,
,
His -> Asp.
, CheA , CheB CheY.
/- ,
, NarX NarQ, , NarL NarP.


. (
).
, Asp -> His His -> Asp. ,
:
1. :
-His1 + -His1~ +

- 32 -

2. 1- :
-His1~ + 1-Asp -His1 + -Asp1~
3. 2- :
-Asp1~ + HPt-His2 -Asp1 + HPt-His2~
4. 3- :
HPt-His2~ + -Asp2 HPt-His2 + -Asp2~
5. :
-Asp2~ + H2O -Asp2 + PI
,
.
,
.
B. subtilis ( )
His-Asp-His-Asp .
Spo0F. Asp Spo0F HPt Spo0B , ,
Asp Spo0.

. ArcB His-
( HPt) ArcA.
.

. B. subtilis
.
,
(PhoR/PhoP), (ResE/ResD) (KinA-B/Spo0A).
-PhoP ResD .
, , , ,
-Spo0A ResD PhoP.

, - ( )
( )
,
.
, .
"" EnvZ-OmpR
ompF ompC.
, , , , "
".
, .
.
.
, :
-.
. ,
.

.
. ,
CheA,
CheW, , (
). - NtrB

- 33 -

 PII, NtrB
, , , .
. ,
KdpD - , ( )
.

, ,
(
).
, .
.
(RapA, RapB, RapE),
Spo0F, , Spo0A.
CheZ -CheY .
,
.

.
, . , VirA A. tumefaciens
Asp-
.
,
.
.
, .
, , .
- , .
:
1. A.M. Stock, V.L. Robinson and P.N. Goudreau. Two-component signal transduction. Annu. Rev. Biochem.
2000. 69:183-215
7.
7.1
E. coli .
, , .
"",
. "" ,
,
.
-
,
(. 7.1).
.
.
,
.
10 .
. , .
MS ,
, P L .
III ,

- 34 -

, ,

.

.
,

-

MS (.
7.2.).

,
(. 7.3.).

MS
, C. (FliG,
FliM FliN),

. ,
MS , ,
.
,

MS
.
,
MotB, ,
MotA
-,

. ,
Mot
-
,

.


FliG () MotA ()

40 , ,

,
. ,

.
.
flhDC , ,

"".
, , , ,
DnaA, H-NS,
flhDC ,
,

.

-
- (RpoD).

. 7.1. E. coli

MotB

S (FliF)

M (FliF)

MotA

FliG
FliN
FliM

. 7.2.

L, P, S, M,
C (
)

- 35 -

,
FlgM FliA. FliA

- 28 F,

, ,
, FlgM
-
,

FliA.
(
) ,
FlgM
FliA,


.

L P

MS

MotA/B
+
+
+ H Na

(a)

45 nm

+
7.2
+
H+ Na +

?

+
+

+
+

(
(b)

)

(,
).

. 7.3.

, - ,
. -
0.1% ,
.
, -
. ,
,
, -
.
. 10-20
.
, ""
, .

, .

"", .
( ),
. , -
""
.

- 36 -

(a)

(b)

. 7.4. E.coli

,
(. 7.4).
, ,
,
, . Rhodobacter sphaeroides
, Rhizobium meliloti
.
"" ,
.
, .
. -,
,
, , , "" 0.2
. -,
, ""
, .. , -
.
7.3
: ,
.

,
(methyl-accepting chemotaxis proteins, MCPs).
, EnvZ, ,
.
- CheA, MCP CheA.
-
, . MCP . 550 , .
. 4 MCP E. coli, (Tsr), (Tar),
, (Trg) (Tap). Tap,

- 37 -

Tcp. ,
,

,
. , MCP
pH,
.

,

,

,


.
(Aer :
MCP ,
, FAD
NifL)

4 5 ,
.

?

:

( ).

.

,

CheA
.

,
CheA.
:

,

() .
1
, ,
"", ..

. 7.5. Tar

. 7.6.

- 38 -

 ,
, .
(. 7.6).


(. 7.7):
CheA -
CheY -
CheW - "" CheA
CheZ - , CheY~
CheA-CheY

""
O 2 /e- ?
MCP
.
R
, CheY

Aer

,
?
, B
. CheA
,
W W
CheW,
A A
.

P
,
Che~ CheY. CheY~
P Y
Y

FliM
,
Z

. CheZ
CheY~,

CheY.

C-

CheY~
,

,

,

,
CheA.
CheY~ ,

.
. 7.7.
,

.
,
. .
.
,
CheR CheB.
, . ,

- 39 -


.
, ,
.
CheR - , MCP
, - S-.
CheB CheA~, CheB~ ,
MCP.
MCP, CheR,
CheB, MCP 0.5-1
.
CheA,
CheB~ , , CheY~, CheB~
CheZ.
CheA. , CheY~ CheB~
, ,
CheB~.
,
.

CheA, CheY, CheY, , ,
.
,
CheA, -CheY (
CheZ),
.
, , .
, ,
CheY, . , CheA "",
CheY~ CheB~,
( CheB~ CheZ),
.
CheY, CheB ,

. " "
.
,
"" .
, . ,
"" (
) (
). ,
CheA ,
.
:
1. M.D. Manson, J.P. Armitage, J.A. Hoch, R.M. Macnab. Bacterial locomotion and signal transduction. J.
Bacteriol 1998. 180:1009-1022

- 40 -

8.

- (, )
.
.
,
,
. , ,
.

.
. , , :
NH3 + + ATP > + ADP + Pi,
, , 2-
:
+ 2- + NADPH > 2 + NADP+.
2. :
(GlnA). , 12
55 . ,
.
( ).
- ,
GlnA . (GlnE),
" ",
.
,
. ,
, .

ATP ADP
,

gltBD.
NtrB
H
NtrB
H-P

Lrp
PII
(leucine-responsive
regulatory PII-UMP UT/UR
protein),

.
NtrC

Enterobacteriaceae.

(.
8.1)

:
glnD,

/
(UTase/UR),

NtrC

D
D

D-P

NtrC

Pi

NTR

. 8.1.
NtrB NtrC

- 41 -

D
D

glnB, PII,
ntrB, ,
ntrC, - .
NtrB ( glnL) -, ,
, . ,
NtrB, PII, ,
- .
NtrB , ,
NtrC. N- NtrB , PII, C-
, His-139.
, NtrB NtrC,
PII. ,

NtrB.
NtrC ( glnG) - ( NtrB) 55 .
54- .
N- "", ,
NtrB (Asp-54). Asp-54
. NtrC
, , NtrC
. NtrC N- ,
N- - (EN). ,
EN.
Asp-54. NtrC C- ,
"--" (
). NtrC - ,
100 .. ,
EN.
NtrC, . NtrC ,
.
NtrC - glnA ntrBC; ,
(glnHPQ), (argT) (hisJQMP); ,
(nasFEDCBA);
nifLA K. pneumoniae nac K. aerogenes.
(GlnD). PII
(GlnB), .
2- ( ),
. 2-
. , PII (GlnB)
2-.
PII (GlnB). , , 2- .
PII NtrB NtrB NtrC.
NtrB 2- . 2 PII, .
, ,
, PII. PII
NtrB. NtrB, ,
, .
NtrC RpoN- glnALG (
).

- 42 -

 NtrC-P, RpoN . nac,

, 70- .
:
1. M. Buck, M. A. Gallegos, D. J. Studholme, Y. Guo, And J. D. Gralla. The Bacterial Enhancer-Dependent
54 (N ) Transcription Factor. J. Bacteriol. 2000, p. 41294136 Vol. 182, No. 15
2. D.J. Studholme, M. Buck. The biology of enhancer-dependent transcriptional regulation inbacteria: insights
from genome sequences. FEMS Microbiol. Lett. 186 (2000) 1-9
9. .
.
,
, .
OxyR SoxR ,
, . ,

"".

ArcB FNR.
, , .
,
.
:
,
(O2-), H2O2 - HO,
, (.9.1). ,

O2
PQ

(HOCl) Ndh II
(NO).
(HOONO) (RSNO).
O2

PQ SiRase
_ SH

_ SH
Acn[4Fe-4S] 2+
2
,

.
H2 O2 + O2

Fe 2+
.
_S
Acn[3Fe-4S] 1+
_S

H2 O

HO
.

, O2- H2O2

.9.1.


.

. E. coli O2- H2O2

. NADH II
. - ,
, - , ,
, .

- 43 -

 E. coli ,
O2- 10-10 M. ,
. H2O2
. ,
, ,
. , ,
( ). ,
NADPH , NO ,
O2-, NO, HOCl H2O2, HOONO RSNO
.
,
- [4Fe-4S] .
- Fe-S
. ,
, , H2O2 .
H2O2 , ,
. , H2O2 Fe2+
HO (),
, , H2O2.
( )
.

,
.
, - SoxRS OxyR.
SoxRS
SoxRS 10 ,
(SodA), IV (Nfo)
(FumC) (AcnA).
. - ,
[4Fe-4S] (, - ) [3Fe-4S]
. , SoxRS -6-
(Zwf), ,
Fur, , , .
. - SoxR
- soxS, SoxS , . SoxR (.
9.2) - , [2Fe-2S] .
[2Fe-2S]+ [2Fe-2S]2+ SoxR.

SoxR, . ,
-.

SoxR

soxS,

SoxR

. ,
.9.2. SoxR
-

- 44 -

 , SoxR 10 35
, . ,
19 .. , soxS
- SoxR,
, SoxR - -
.
soxS soxR ,
, .
OxyR
, OxyR, H2O2 30
( OxyR 9 ), ( I,
KatG), (AhpCF), (GrxA),
(GorA), Mn- Fur-. OxyR
- Dps (DNA-binding protein from starved cells) OxyS
.
.
oxyR 34 kDa , LysR-
. , OxyR
, H2O2, .
. OxyR
, , oxyS,
, .
OxyR - ,
, - (. 9.3). H2O2 OxyR,
OxyR.
, .
(, , ) OxyR 1
(: , NADH
),
,
,
OxyR, ,
OxyR
.9.3. OxyR
,
, .. . .
.
- FNR (fumarate and nitrate reduction) ArcA (aerobic respiratory control)
,
. FNR
- Fe-S , , ArcA
, ArcB.
FNR ArcB ,
.
, .
.
FNR
FNR , 120 E. coli.
, .
,
, , ,

- 45 -

 . FNR
, , DMSO,
. FNR
NADH
.
, FNR . M
- ,
- -. FNR
- ,
(Cys-X3-Cys-X2-Cys-X5-Cys),
- .
[4Fe-4S]2+ FNR ( )
[2Fe-2S]2+ , FNR,
(. 9.4). , [2Fe-2S]2+
[4Fe-4S]2+ .
[2Fe-2S]2+, ,
. FNR ,

[4Fe-4S]2+ .

2+
[4Fe-4S]
[2Fe-2S]2+

.9.4. FNR
FNR
,
.
E. coli.
1-5
( ),
. FNR
.
ArcB

(5-10 ).
( -
). , ArcB,
.
ArcB ArcA (.
ArcB
9.5). ArcA
P
P
P
P
30 ,
Asp
His
Asp
His
ArcA
(
P
, ..),
ATP
ADP

Asp
OmpR
.
P
P

ArcA
Asp
SixA
Asp
CheY
, ArcA

.
.9.5.
ArcB
ArcA/ArcB
.

- 46 -

: , ,
, HPt -. ArcA
ArcB , HPt .
,
, "" HPt .

AerA , . - Azotobacter, - Tsr - .

AerA CheY, CheA CheW, , AerA , .
,
SoxRS OxyR.
.

. , RpoS ,
, , , . ,
RpoS, ,
. RpoS ,
katE, xthA sodC. OxyR- katG, gorA dps
RpoS.
FNR ArcAB sodA, sodC, acnA
fumC. , SoxS, MarA Rob,
SoxRS. , , Fur,
SoxRS, OxyR.
:
1. G. Storz, J.A. Imlay. Oxidative stress. Current Opinion in Microbiology 1999, 2:188194
2. C.E. Bauer, S. Elsen, T.H. Bird. Mechanisms for redox control of gene expression. Annu. Rev. Microbiol.
1999. 53:495-523
SoxRS, FNR, OxyR, ArcB
3. G. Sawers. The aerobic/anaerobic interface. Current Opinion in Microbiology 1999, 2:181187
. FNR ArcB.
10.

10.1. Escherichia coli

() ,
( )
. FtsZ
.

MinCD , , MinE,
, .
,
, , , .
,
, - ( B. subtilis)
Caulobacter crescentus (
). C. crescentus (CtrA),

- 47 -

, , FtsZ, , ,
.
, ,
,
. , ,
, - .
, , - ,

. , Escherichia coli, ,
, , ,
, .
,
. - , ,
,
,
. - ,
, .
, , ,
, .
, ,
,
. ,
, ,
, .
,

.

10.1.1. .
o ,
, N- (GlcNAc) N- (MurNAc), 1,4 (. 10.1). 29
. .
L-Ala-D-Glu-DAP-D-Ala-D-Ala (DAP - meso- )
MurNAc L-.
(DAP E. coli)
. - , D-
D- DAP
, .
DAP,
. , ,
5 % . D-
.
90 .
, E.coli, MurNAc
.
.
, ,
.
.
"" . ,
, ,
- ,

- 48 -

NAM

NAG

NAM

NAG

Muramidase

Muramidase
CH 3

OH
CH 2
O

HC
CO

CH 2

CO

OH

HC

HO

CO

NH

CH 2

CO

OH

NH 2

CH 3
OH

D-Ala

CH 2

D-Ala

CO

CH 2
O

NH

CH 2

CO

OH

O
HC

CH 3

CO

NH
O

HO

CH 3

L-Ala
D-Glu

OH

HO
O

CH 3

CH 3
O

NH
O

HC

Endopeptidase

NH

O
CH 3

CH 3

m- DAP
Carboxypeptidase

CH 3

L-Ala
D-Glu

NH

CH 2

HO
O

CH 3

OH

NH

CH 3
O

NH

CH 2

CO

OH

CH 3

NH 2

m- DAP
D-Ala

.10.1.
,
, .

10.1.2. .
",
", ,
, .
, .
E. coli . ,
,
.
. , ,
. - ,
, ,
.
, "" ,
.
"" , , ,
, . ,
,
. DAP
- . - ,
(. 10.2).

- PBP3 PBP2 ,
, .
, , , ,

- 49 -

(a)

(b)
LT

Docking str and

EP(AM)
Ala-Glu-A2pm
Ala-Glu-A2 pm-Ala

TP

Ala-A2pm-Glu-Ala

Stress-bear ing
la ye r

A2pm-Glu-Ala

Ala
Ala
- A2pm
Glu
Ala

TP/TG

Ala
Ala
- A2 pm
Glu
Ala
Ala
Glu
A2 pm
Ala

Ala-A2 pm-Glu-Ala

Murein tr iplet
Ala
Glu
A2 pm
Ala
Ala-Glu-A 2 pm-Ala

TG

.10.2.
.
,
.

10.1.3. .
.
, , , ,
, " " ,
,
, . ,
, .
,
fts (filamentous temperature sensitive). ,

,
- ftsZ (

- Chlamydia
trachomatis ftsZ). ftsZ

, M. genitalium,



( )
.
, .
.10.3.
-

,

- 50 -

Eubacteria. .


, ,
.
ftsZ - , ,
,
. ftsZ, mra 2.5 ,
,
5,000 20,000 . FtsZ
, .
FtsZ ,
. N-
- , C-
, C-
. FtsZ
in vitro -, ,
. , FtsZ
, , 20 . FtsZ
(), in vivo.
, , FtsZ,
, .

. , FtsZ
ftsA, ftsQ ftsI , , ftsW ftsK , ,

. FtsZ , rodAsui
ftsZ26.
. , FtsZ "
" (, ),
. ,
FtsZ-GFP in vivo, Z- ; ,
, . FtsZ-GFP
. 67 C- FtsZ
Z-, ,
, , ( ),
.
FtsZ Eubacteria Archaea, ,
( C. trachomatis);
, Arabidopsis thaliana, ,
.
Z-, ,
, ,
, FtsZ
, , , ,
, .
FtsZ
.
.
ftsA, ftsZ, 46 kDa ,
150 . FtsA FtsZ (1:100)
. FtsA
,
. -

- 51 -

 - , (HSP70),
, ,
. , , ,
, , ,
ftsA.
GFP , FtsA
FtsZ - ,
FtsZ, ,
FtsZ ( ). FtsA, ,
, FtsZ.
, FtsZ FtsA, .
- , FtsA-GFP,
, ( , FtsZ-GFP ), FtsZ
. , FtsA-GFP ,
FtsA,
.
ftsQ ftsA. - 31 kDa
N- -.
. FtsQ , , 50 100 .
, ,
rodA, FtsQ FtsZ.
FtsQ
. -
, ftsQ1ts
. FtsQ FtsZ FtsA,
FtsL FtsI. , FtsQ
,
FtsL FtsI, .
ftsW murD murG mra. ftsW
,
. RodA
. , FtsW PBP3,
. ftsW
FtsZ, 50 %
, FtsW, 50 %.
ftsN , 88.5 , 36 kDa
N-, N- .
50 .
,
MalG MalE,
, .
,
. ftsA12 ,
ftsI23 ftsQ1 , ftsW ftsK44
. FtsN (
), . ,
, FtsN FtsQ,
FtsI FtsL ' ' .
ftsL (mraR) mra. - 13.6 kDa o
e ,
. 30 40 ,
. ,

- 52 -

 , . FtsL
. , ,

. , ,
, FtsL
. FtsZ, FtsA FtsQ, FtsI.
, FtsQ, FtsA FtsZ FtsL.
zipA ,
, FtsZ. .

. ,
ZipA ,
FtsZ,
. , ZipA::GFP
91 % , .
FtsZ , . ZipA - 36
kDa, 50 kDa SDS-PAGE. N-
, - .
103
, C-
. , , ,
FtsZ. ZipA
FtsZ ZipA,
, FtsZ in vitro ZipA. Hale de Boer
ZipA FtsZ ( 1-10 %
FtsZ). , , , , Z ZipA,
. ZipA FtsZ-, FtsA- FtsI . ,
, FtsA ZipA.
ftsI , ftsQAZ,
, PBP3 . , , 50-100 . PBP3 , .
, . PBP3 . , PBP3
- , , PBP2
RodA . FtsW
FtsW RodA; , , .
;
.
, FtsZ, FtsA, FtsQ, FtsL.
ftsK 20 SOS- (dinH)
147 kDa.
SpoIIIE . B. subtilis

, . , C-
Tra Streptomyces
Tn916. FtsK, SpoIIIE, N-
,
.

- 53 -

, N- FtsK GFP ,
N- 15 % . N-
ftsK44 .
ftsK44 - ,
.
ftsK rodA , .
FtsK - FtsK-GFP
, . ,
, FtsK , ,
.
, - ,
SpoIIIE "-" ,
.
, N- FtsK , , ,
, .

10.1.4. .
,
, .
.
,
. ,
.
, , ,
, , , (,
, , ).
, ,
, E. coli ( )
. , ,
. - FtsK
muk .
mukB
. . MukB -
.
,
, ,
.
MukB . mukB , mukE mukF.
.
MukB . N-
MukB FtsZ. FtsZ
MukB. MukB ,
FtsZ E. coli. FtsZ
, MukB "", ,
.
SeqA.
Z- MukB.
ftsK .
FtsK SpoIIIE B. subtilis Tra
Streptomyces,
. ftsK44 , ,

- 54 -

 C- , .

10.1.5. : min.
minB .
, minC, minD minE. MinC MinD ,
,
. MinE ,
. MinC
MinD, ,
, DicB. minB ( MinC MinD)
, ,
. "" " "
, . ,
,
.
MinD
. , MinD in vitro.
, , (88 A), MinE
. N- ,
MinCD,
, C-
MinE, . ,
MinE .
MinD, MinC FtsZ. , MinE
, , , ,
FtsZ, .
, Min .
, FtsZ, ;
, ,
Min , MinE MinC MinD.
, MinE, .

10.1.6. .
, ,
, . ,
, ,
- .
, , , ,
, . , ,
, Z-. ,
Z-.

, , .
. zipA
FtsZ. N- MukB FtsZ
. FtsA-FtsZ
FtsA E. coli. FtsA Rhizobium meliloti
Agrobacterium tumefaciens E. coli, FtsZ
. FtsZ FtsA
. ,
FtsW PBP3 PBP3 PBP1B.
PBP .

- 55 -

 GFP
, .
FtsZ ,
in vivo GFP.
- FtsA, FtsQ, ZipA , FtsW.
fts
, , .
FtsA Z- .
FtsQ FtsZ FtsA, FtsL FtsI. FtsI
FtsZ FtsA, , FtsL FtsQ. ,
FtsZ , PBP3, . FtsN
, FtsZ, FtsA, FtsQ FtsI
. ftsK , GFP
FtsK, ,
; FtsZ FtsA, FtsQ FtsI
.
, ,
"" : FtsZ-FtsA-FtsQ-FtsL-FtsI. FtsN
FtsI, , FtsA, FtsQ, FtsI FtsK,
. FtsK FtsA, ,
. ZipA FtsA FtsZ
. FtsW FtsZ,
Z- (. 10.4).
FtsI - , , -
, . , Z- FtsI.
, ,
, ,
. FtsI

( )
. FtsI

,

, FtsI
.

FtsI

,
.
- ,

,
, FtsZ. ,
FtsA FtsZ 1:100. FtsQ,
FtsL (?) FtsN (?)
, FtsZ.
, ""
, "",
Z-,

.10.4. Escherichia coli

- 56 -

, . ,
.
,
, , ( )
, . ,
"" .

10.1.7. .

, - . , ,
-
, SOS .
.
dcw (division and
cell wall) mra 2 E. coli (. 10.5). mur ,
, .
16 , ,
. ,
, .
- ,
.
,
.

.10.5. dcw (mra) Escherichia

coli

- 57 -

 dcw. mra
, ftsW. ,
in trans,
, . , ,
, ftsZ, , (mra) .
mra, , dcw,

. - ftsZ
. , ,
, .
ftsZ .
, ""
E. , ,
ftsZ ddlB. ,
ddlB.
, 6 ... ftsZ
.
ftsZ
(. 10.6). tac
ftsZ ,
. , ftsZ
(
).
ftsZ .
ftsZ,
. , ftsA,
, FtsZ,
ftsZ .


, , , ..
. , ,
, ddlB, .
sdiA (suppressor of division inhibition)
. ,
- ftsZ84 .
- , SdiA
ddlB ftsQAZ envA .
,

.
,


LuxR-

"quorum
sensing"
.

( luxI),

.10.6.
, N ftsZ Escherichia coli
-,

- 58 -

 .
, luxR- . ,
E. coli luxI, : SdiA?
.
SdiA- (ftsQp2).
, (30-35%)
"" , E.coli.
, (40-60%)
sdiA. ,
ftsQp2 ,
V. fischeri V. harveyi, , "" .
, sdiA luxR-
, E.coli .
rpoS , - ,
ftsQAZ . ftsQp1, ,
ftsQAZ , ddlB-ftsZ , RpoS.
stfZ - , 5' ftsZ.
,
,
, ,
, ftsZ dicF.
.
, , mra ,
ORF . ,

. FtsZ , , ( 20,000 ), FtsQ
- . ORF
, , ATG -.
, ORF
, .
-
SOS . E.coli ,
, .
, ,
, . SOS-
SulA.
sulA (sfiA) , Z-.
SulA FtsZ. ,
SulA FtsZ in vitro. FtsZ -
, SulA . SOS ,
SulA Lon, .
FtsK SOS .
, "" .
FtsK , SOS ,
- .
10.2. Caulobacter crescentus.
Caulobacter crescentus , ,
, :
G1, S G2 ( );

- 59 -

,
Replication
Chromosome segregation
(..
Flagellum
Cell division
Asymmetric
)
Motile
Sessile
Chemot actic
Replication
Predivisional

Replication bloc k
initiation
cell
,
Swarmer
St alked
cell
cell

(. 10.7).


(..
G1 ,
St alk

G1
S
G2

).
Cell cycle

Flagellum St alk
Flagellar biogenesis
ejection
biogenesis
( Development
)

.10.7. Caulobacter crescentus
,
.

( S ).
,
.
( G2), . ,
:
() . ,
,
.
CtrA,
. ,
, (. 10.8).
. CtrA
ccrM, -, C.
crescentus. G2 ,
CcrM Lon,
, . . G2.

( S
ClpXP
).

DivJ
DivK
?
CtrA
,
DivL
C. ?
CtrA~P
crescentus, E. coli,
P1 P2
Cc kA
ctrA
,

+
.
CtrA
+
+
+

ccrM
fla
ftsZ
ftsQA
. ,
Cori
CtrA-Box
CtrA ftsZ
,
ftsQA
.10.8.
.
Caulobacter crescentus

CtrA

- 60 -

 ftsZ, ftsQA.
()
.
, , ,
CtrA PCori,
(Cori).
. Cori 5 CtrA- ( CtrA),
PCori. - CtrA PCori,
, ,
(.. ,
).


CtrA. , CtrA
.
( G2 ).
, CtrA ,
.
ctrA , .
P1,
"" P2, . CtrA ,
, . ,
CtrA P1 , P2 , ,
, CtrA S .
CtrA .
CtrA , ,
. , CtrA
fliQ 20 , ccrM, fliQ, CtrA
, , ccrM
, CtrA .
, " " ,
CtrA . CtrA

. CtrA , S-,
CtrA, PCori . CtrA
- ClpXP. , E. coli,
Caulobacter "" G1
. "" "" .
CtrA . ,
( CtrA)
. CtrA
, (DivJ, DivL CckA)
. , , , .
, , DivJ CckA, ,
CtrA. ,
CckA, ,
. CckA ,
.
CckA CtrA. DivJ
() G1 S .
, , CtrA , ,
.
"", "". ""
, , ,

- 61 -

CtrA.
:
1. Magne sters and Urs Jenal. Regulatory circuits in Caulobacter. Current Opinion in Microbiology 2000,
3:171176
2. Christine Jacobs and Lucy Shapiro. Microbial asymmetric cell division: localization of cell fate determinants.
Current Opinion in Genetics & Development 1998, 8:386391 (less detail but better text overall)
3. J. Lutkenhaus. The regulation of bacterial cell division: a time and place for it. Current Opinion in
Microbiology 1998, 1:210215
4. J. A. Hoch. Initiation of bacterial development. Current Opinion in Microbiology 1998, 1:170174

11. Bacillus subtilis.

B. subtilis
, ,
. 125 , .
-
,

, .

11.1.
. . ,
.
(),
, () .
.
(
).

,
. 6-8
,

.

,

0-VII. 0
,
, I -

, II III


..
.11.1. Bacillus subtilis

,
. ,

- 62 -

 spo0 spoIII 0 III .

(., spo0A spo0B).
, (.,
spoIIAA, spoIIAB spoIIAC).
?
, ,
, .
, .

, .
, , , , .
(Spo0A). Spo0A
. Spo0A
, .
, , , , ,
. ,
.
(),
() .
, ,
.
.
,
KinA KinB,
. , Spo0F ,
, Spo0A, Spo0F Spo0B. Spo0A~
,
.
AbrB, ,
. Spo0A~ AbrB
. , Spo0A~
, spoIIA spoIIG, F ,

,
,
. ,
,

KinA.

,
KipI,

.11.1. Bacillus subtilis

KipA,

kipI.

- 63 -

 ,
.
- ,
. Spo0E,
Spo0A~. , ,
Spo0E. ,
. RapA RapB
Spo0F~. Spo0F~ Spo0A~ ,
Spo0F Spo0A.
RapA RapB ,
. RapA
ComP/ComA,
( , ). RapB ,
,
, ,
.
Spo0A ( ).
,
phrA. RapA,
( 44 ) . -
, SecA-
. 19 . , ,
, - , RapA.
Opp,
5 - 19 .
. -, ,
PhrA , ,
. -,
-.
PhrA , ,
.

,
Spo0A

,


. Spo0A~P

,
K-,
A

H.
H

(
), .

- - F
E . F
, ,
.11.2. F

,

- 64 -

 .
, SpoIIE,
SpoIIAB SpoIIAA. F,

. SpoIIAB , F
.
SpoIIAA - - ,

SpoIIAB,
SpoIIAB.F
F. SpoIIAB
,
SpoIIAA,
. ,
F SpoIIAB,
.11.3. E K
, SpoIIAA

F. , SpoIIE,
, SpoIIAA~P. SpoIIE
, .
,
, SpoIIAA~P
. 5-7 ,
SpoIIAA , ,
F (. 11.2).
F , , (. 11.3). , F,
. - ,
, -. -
spoIIG
Spo0A~P. -
, spoIIGB, , spoIIGA ,
- . ,
spoIIR, -,
F. ,

, F
. (SpoIIR
,
.)
-?
, ,
, ,
F,
.

, spoIIIG,
-, G. -
,
.11.4.
. , G,
-
- F,

,
.

- 65 -

(SpoVT )
- K (.
11.3b). - , , K
-K,
- 20 .
-K , SpoIVFA,
spoIVB, G .
-
, ,
,
( 11.4).
1. R. Losick
2. M. Perego. Kinase-phosphatase competition regulates Bacillus subtilis development. Trends in
Microbiology. 1998. 6:366-370

.

12. . Quorum sensing

,
. ,

, . ,
"
" .
. ,
,
, .
,
, .,
, .
-
(), .
, .
"Quorum sensing" ( ).
,
, , LuxI- .
. , ..
, ,
LuxR- - , . - ,
. ( 1970 .)
- Vibrio fisheri, " "
(lux).
(Pseudomonas aeruginosa, Erwinia carotovora), -
(Agrobacterium tumefaciens), (Serratia liquefaciens), (Erwinia
carotovora) (. 12.1).
Vibrio fisheri
, lux-, -

- 66 -

.12.1
, , ,
- ,
. luxI , N-(3-)-L.
. Vibrio fisheri

. , ,
1010 , ,
.

. , luxR, ,
. . -
HTH ,
, lux-. (20 ..)
, . - LuxR
. , C- -
N- , . N , -
LuxR lux- .
, LuxR- . EsaR
ExpR, Pantoea stewartii Erwinia carotovora,
. lux-.
LuxR- lux-, -35
, EsaR -10, , ,
.
.
,
.

- 67 -

.12.2.
LuxI S- (SAM) ,
- (ACP).
S- ,
- 5- (MTA).

Erwinia

Erwinia
carotovora

,
.
,
, .

,

.

N-3--L- ,
LuxI,
ExpI,
CarI HslI. expI expR luxR. expI
.12.3.
, expR -
Erwinia
.

- 68 -

 , Erwinia carotovora -
- .

, .
CarR CarI, , ExpR
ExpI. CarR ExpR , , CarI, ExpI.
Erwinia
.
- . - (
).

- ,
, , .
,
. ,
, ABC-.
, , -,
-, ( )
. Bacillus
subtilis, Staphylococcus aureus, Enterococcus faecalis
Lactobacillus sake.
:
1. C. Fuqua, M.R. Parsek and E.P. Greenberg. Regulation of gene expression by cell-to-cell
communication: acyl-homoserine lactone quorum sensing. Annu. Rev. Genet. 2001. 35:43968

13.
20% , , .

,
,
. ,
,
, ,
.
.
. ,
,
.
, . ,
,
().
, ,
, .

:
.
.

- 69 -

- .
.
.
.

.

.
:
( ,
);
""
( N-),
;
;
();
,
( ) ;
( ,
);
(
III )
, ( III IV
)


()
. ,
,
Sec- ( II ).

( 20 ). , -
, (
,
). ,
SecB ( )
SRP .
", " (signal recognition particle, SRP)
.
,
SRP. SRP ,
6 300 . 54 kDa
SPR (SRP54) ,
. N-
. SPR
, --
, SPR SPR (SR)

.

.
SPR. SRP :
Ffh - SPR54 ( 100 ) (4.5S ), SR ( SRP)

- 70 -

 - SR - FtsY.
, SRP
. , SRP E. coli
. ,
, DnaK () SecB.
- SecB
signal recognition particle (SRP),
Ffh 4.5S .
SecB SRP . SRP
.
, (
) SRP
( SR), Sec .
SRP SecA/SecB " "
. SRP
. SRP
, ,
,
,
.

.
,
,
, . ,
, ,
.
. GroE/DnaK
,
.
.
, ,
.
. ,
"" () , , ,
, . ,
. , III
, , . ,
. ,
III , .

HSP60/HSP70.
, ,
,
, .
,
( -
)
.
.
, , ..
(GSP), sec .

- 71 -


.
.
.
,

ABC

.
,

,
. (gatekeeper) .
- ,
.
, ,
,
, (),
.

1-
C-,
60 .
-,
.

.13.1. I

(GSP).
, , ,
- Sec , - ;
. II

,
,
( );
, , Xanthomonas campestris; , ,
, Pseudomonas aeruginosa; Klebsiella
oxytoca. ,
II " " (General Secretory Pathway, GSP).
.
, -
. - ,
( , ),
, GSP.
Sec .
,
. Sec :
-
-
-

(, ).
, .

- 72 -

GSP.
10
Sec
(. 13.2).

SecB.

, ,

, ,

.13.2. Sec-

,
. SecB ""
SecA .
GroEL DnaK.
,
- . ,
, SecY, SecE SecG.
, - . SecY (10 - - , , LacY). SecE
, SecD SecF - .
SecA. -
. SecA - , - -, . -
SecA SecYEG, .
SecD SecF .
( ).
.. ,
. ,
:
:
++hhhhhhhhhhhhhhhh(PG)NNANAP Oor- (LepB - )
++hhhhhhhhhhhhhLNACD Oor(LspA - )
++QRGFhhhhhhhhhhhhNNNNNN (GspO)
^
,
. ,
.

.
SecB ,
SecA, SecYEG. ,
- SecA, SecA-SecB.

SecB. - SecB SecA.

- 73 -

SecA ( ). . . ,
. in vitro
SecA .
Sec-.
SecY.

Sec- (
).
, , ,
.
- . Sec ,
. ,
, , , .
DsbA DsbC, .
.
.
.
(
), ,
DsbA DsbC. ,
DsbAC , , .
GSP
JMB659 GspEL interaction, good intro
, ,
, " ".
, (.13.3).
GSP .
(, , ) ,
, 13-15 ..., 12-15
. - pul , out ,
gsp,
.

gsp

,
()
,
gsp
.
, Gsp

.
,

,
(GspE)

.
(GspGHIJKMN)
.13.3.

N-

C-,

- 74 -

 - . GspL ,
, . GspO GspF
.
Gsp . GspO N GspGHIJ(?). ,
, , ,
IV (, P. aeruginosa).
PilD, Phe N-
. GspGHIJ N-MePe GspO, "",
. , ,
, N-
. , ,
,
GspO .
GspE , , .
ABC-,
, - (
, ), GspL (
GspE).
, GspD GspS .
GspD ,
II, III .
,
. . C-
. N- , ,
.
GspS GspD,
GspD .
, GSP. ,
GSP ,
, ,
. , GspE
GSP. , N-
GspL, GspE, ,
GspL,
GSP.
II . , -
,
. , , E. carotovora
E. chrysanthemi, ,
90%. , -
. : ,
, ,
..,
.
GSP
, ,
, .
, GSP- , ,
- . , pul Klebsiella MalT, out
- KdgR. ,
,
. ,

- 75 -

Gsp- (, ..)
.


Out E. coli
III
1.1. III .
III ,
Bordetella bronchiseptica, Burkholderia pseudomallei, Chlamydia pitsitacii, Erwinia amylovora,
Erwinia crysanthemi, Escherichia coli, Pseudomonas aeruginosa, Pseudomonas syringae,
Ralstonia solanacearum, Rhizobia spp., Salmonella typhimurium, Shigella flexneri, Xanthomonas campestri,
Yersinia.

( , ,
),
.
III ,

.
III
(. 13.4). Yersinia spp.

14
Yop (Yersinia outer
proteins) . ,
Yops, ,
,
(ysc, Yop secretion genes)

.
37C,
,
,

III .

,

Yop-,
,

,
.
III
Yersinia spp.
24 Ysc , 11

.13.4. III
(

).

- 76 -

 III 11
10. yscC,
(), Yop .
III ,
, GspD II . , 11
(Hrc-) - ,
.
( ).
,
, .
III S. typhimurium, ,
(
), .
III
,
. , , ,
, .
37C
. III sec-. ,
, II .
, N- ,
( , ,
).
,
,
.
,
, III
. , - hrp-, , ,
HrpA . hrp, 8-10 .
HrcC ( YscC ),
HrcJ, HrcR, HrcS, HrcT, HrcU, HrcV (HrpO),
HrcQ HrcN, .
1.2. III .
, III , ,
, , .
.
YopN - , "" . LcrG,
. YopN HrpJ,
, .
YopE . ,
.
YopH - ( ,
, )
YopB YopD - . , ,
, , ,
.
, III ,
HR - Avr . ,
, hrp .
III . ,
Avr- ,

- 77 -

 ( ,
) , ,
,
, ( YopB YopD ).
(
"" ).
(40-45 kDa) ( 20%)
, .
HrpN, HrpZ
HrpW.
,
.
,

( , ) . , ,
, ,

. , HrpW ,
( , , ),
.
, , , HrpZ. , HrpZ, , ,
.
, III
, Avr-
, .
Avr- ,
. , Avr-
, ,
. Avr- R-
Avr-R .
1.3. , III
,
III ,
( ), III
. , III , ,
, ,
, .
, , .
,
(%GC),
.
, "
"
.
hrp
20-25 ... 7-8 .

,
. hrp P. syringae E. amylovora,
- R. solanacearum X. campestris.

- 78 -

-.

P. syringae E. amylovora
-,
R. solanacearum X. campestris
(, , )
AraC (. 13.5)

P. syringae pv.
syringae.
, HrpR, HrpS, HrpL

,
hrp avr
. HrpR HrpS
NtrC-
54-
. HrpR HrpS

NtrC ,
,

(
, ,

hrp-,

), -

-
.

NtrC-

.13.5.
- RpoN (54),
III
, rpoN
hrp
P. syringae. RpoN , HrpL, ECF (extra cytoplasmic functions) -.
HrpL ,
, hrp avr .
HrpL GGAACCNA-N14-CCACNNA,
hrp-. E. amylovora.
.
3-
. Syc
Yersinia, Yop .
Syc
( ).
, . ,
, .

,
( ) . ,
, . ,

- 79 -

 , 5 (FlgB, C, F G, FliE).
(FlgE), . K
, (FliC),
, FlgK FlgL, , FliD.
,
, FlgK, FlgL FliD. ,
, 25-30, (
) .
, -
.
,
FlgN, FliT FliC.
, -
III .
. .
125 N- ,
, -
40 ( -)
.
? ,
N- . Yop
.
yop , ,
15-17 . , Yop
.
IV
IV
.
( ,
),
, , , ( ,
).
IV
- T- A. tumefaciens, Tra IncN pKM101
B. pertussis.
Tra CagA
Helicobacter pylori.
, IV , ,
, , .
(oriT),
. , oriT
( - )
. 5'
. 5' ->
3',
. A. tumefaciens VirD2 -
, , VirD2, -
. ,
, .
, ,
. , Tra RP4 RecA,

- 80 -

 - VirE2 (SSB ) , VirF.

, VirE2 VirE1. VirE1 VirE2 in
vivo in vitro, Syc Yersinia.
IV
Tra , , . ,
Ptl B. pertussis PT, A/B
5 , S1-S5. S1, S2-S5
1:1:2:1. Tra PT .
Sec
,
Ptl.
.

.
IV ( Tra )
, ,
, ,
. , ,
:
, . ;
;
.
II-IV . , III
IV -.
. ( )
, . , -
.

II (
PT B.
pertussis). ,
, PT B.
pertussis,
II . ,
CTX V. cholerae
II
,
II

.

,

.13.6. I-III
"" ,
IgA Neisseria gonorrhoeae

:
1. A. Economou. Following the leader: bacterial protein export through the Sec pathway. Trends in
Microbiology 1999 VOL. 7 NO. 8 p. 315-320
2. D. M. Anderson and O. Schneewind. Type III machines of Gram-negative pathogens: injecting virulence
factors into host cells and more. Current Opinion in Microbiology 1999, 2:1824

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2. S. Lory. Secretion of proteins and assembly of bacterial surface organelles: shared pathways of extracellular
protein targeting. Current Opinion in Microbiology 1998, 1:2735
( 4 )
4. L. W. Cheng and O. Schneewind. Type III machines of Gram-negative bacteria: delivering the goods. Trends
in Microbiology 214-220 Vol. 8 No. 5 May 2000
5. C. J. Hueck. Type III protein secretion systems in bacterial pathogens of animals and plants. Microbiology
And Molecular Biology Reviews, 1998, p. 379433 Vol. 62, No. 2
14.
,
( ) ,
, EF-Ts, EF-Tu EF-G, - -,
. 510% . , ,
.
,
relA. ,
- (ppGpp) (pppGpp).
.
ppGpp - , . RelA (p)ppGpp -
.
- ,
, ,
. ,
. ,
,
. , ,
. .
, , 105 ,
,
.
(p)ppGpp, , ,
. (p)ppGpp , E. coli,
,
.
,
, - . A- , .
csgA, a, C - 17 . C-, ,
- , ( csgA) . C, ,
FruA. FruA - frz,
devTRS csgA. , Che E. coli
,
. devTRS .

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15. SOS

, .
.
, , ,
,
. "SOS-"
SOS- E. coli RecA
LexA, SOS-.
RecA.
RecA LexA.
LexA.
SOS- .
SOS- . ,
, ,
, RecA.
RecA . LexA
RecA-.
LexA ,
LexA . , LexA ,
UmuD, MucA CI. RecA
(- RecA ).
SOS- . ""
LexA
, RecA.
LexA ( 5'-taCTGTatatatatACAGta-3').
"" SOS- (
30):

/
SOS-
lexA
; SOS-
recA

ruvAB

recN
(, )
uvrA
()
uvrB
()
uvrD
(SOS-)
umuDC

sulA
SOS- ,
LexA.
SOS- ,
.
,
. ,
SOS-. , UvrA ,
, ,
20-30 SOS-
. RecA (
, 10 ), RuvAB RecN ,
30 . ""
( umuDC), 30-40 .

- 83 -

 E. coli,
50 .
, ""
SOS-
( ) ( ),
, ( ) .
, umuDC,
, . , ,
SulA-
"" .
SOS-
, SOS-
. ,
SOS- . UmuDC-
, , ( ) ,
,
. (
). ,
- .
SOS- , RecA UmuD
UmuC, UmuD ( LexA)
UmuD', UmuC (UmuD'2UmuC),
(
UmuD'2UmuC - V). UmuD'2UmuC RecA, -, ,
(, ,
SOS- ).
:
1. W.H. Koch, R. Woodgate. DNA Repair in Prokaryotes and Lower Eukaryotes. DNA Damage and
Repair.vol.1: Edited by: Nickoloff and MF Hoekstra
2. G.C. Walker. The SOS Response of Escherichia coli. The Bible. 1996 p 1400-1416
17.

.
, ,
. ,
( ),
.
. ,
. ,
20 20 .
,
, ,
( ) .
, , ,
(. 17.1). 3'
. ,
. , E. coli 500 REP
(repetitive extragenic palindromes) ,
3' .
5' .

- 84 -

5'
REP

RBS

-).
,
,
3'
35-50 5'
,

,
.17.1. ,

Protected

3 end
.

.
3
. 5

, , ,
Endonuclease
. -
,
, 5
3
. , ,
, ,
Exonuclease
(., ssrA
E. coli). , ,
, 5
3
.
E. coli
- . , II
.17.2.
(, PNPase),

3'->5'.
-, .
II , (a) 53pathway
PAB
PAB
PAB
AAAA AAAA AAAA 3
. 5Gppp

Deadenylation and decapping

.

5
3
.
3' (
- )
, (b) 35pathway
PAB
PAB
PAB
. 5Gppp
AAAA AAAA AAAA 3

"
"

Deadenylation
,
3' , .
E ( E, RNAse E), 5Gppp
3

5S ,
.

.17.3.
,

- 85 -

 , 5'->3' 3'->5'.
5' () ()-
(PAB) 3' ,
. 5'->3'
, XRN1. 3'->5'
. , E. coli 5'->3'
. 5' , 5'->3' ,
, , . 5' , ,
, .
5' ,
.
,
3'->5' (
3' ) .
,

E. coli. rne. (1061
). . .
- . N-
S1.
.
.
(A/G)AUU(A/U),
"" AU
. ,
5' , .
, . ,
5'
.
,
, .
40- rne-
.
III
III ( rnc) E. coli ,
. - ,
.
, ..
.
III ,
III. rnc B.
subtilis, E. coli.
III - 25.4 .
- . rnc ( ,
III ). III
.

pnp. 77.1 .
. ,
S1. . ,
, .

- 86 -

 II
rnb. ,
3' , , ,
. 72.3 . S1 . II
,

,
.
.
E. coli . , -
RhlB (. 17.4).
. (
) ( ) PPK DnaK GroEL;
.
. N-
. , ()
(U) . C-
, RhlB, PNP . , ,
N- , C- ,
.

,

-, .

3'->5' . ,
PNP ,

.
. mtEXO,
,
II E. coli,
Saccharomyces
cerevisiae.

S.
cerevisiae,
.17.4. -

, ,
3'->5' ,
II PNP E. coli.
Rpr4p ,
.
-
- RhlB
- . RhlB DEAD-
. - -, 8 ,
(D), (E), (A) (D). -
, ,
. in vitro RhlB .
,
. , RhlB -
. RhlB ""

- 87 -

 , .
RhlB C- .
mtEXO , RhlB.

.
()- I (PAP I) E. coli pcnB (plasmid copy number),
ColEI . PAP
I I , II
, ColEI. I, 2
, 5 5' .
( I-5) 10 pcnB ( PAP I
). E. coli (A)- (PAP II).
(A)- . PAP I E. coli (A).
, E. coli
. ,
, .
() ,
. , I, "",
"" 3' . 3'
.
3' ()-
, ().
, ()- , ,
PAB (poly(A)-binding). ()
.
:
1. M. Grunberg-Manago. Messenger rna stability and its role in control of gene expression in bacteria and
phages Annu. Rev. Genet. 1999. 33:193227

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