Copyright & A K-C

MADS-box .
( 03.00.03 - )

- 2002

Copyright & A K-C

-
.
:

,
..
,
..

*.

,
..

..

,.

2002 .

.006.027.01

> ~<- --

'-27550, . ,

. .;
. iv - -

"

,

..

Copyright & A K-C

,
.

MADS-box (Theissen
et al 2000; Ng and Yanofsky, 2001; Theissen, 2001).
MADS-box

, /

. -
MADS-box

.

MADS-box
.
MADS-box
.
MADS-box
,
MADS-box
.
. -
MADS-box ,

MADS-box , .
.'..-
, - ^ . \
;. cvivCK.-'X-Y). !ta..-;v)',',nn
I
. . .
. i'- CI

Copyright & A K-C

:
>

MADS-box -box
MADS-box : apetalal, agamous agamons-like.
> MADS-box
( ,
),
.
>
MADS-box
MADS-box {Arabidopsis thaliana L.)
{Antirrhinum mqfus L.).
.
MADS-box
.

(Murai et a]., 1998), (Schmitz et al., 2000)

(Lolium temulentum L.) (Gocal et al., 2001).

Genbank accession numbers AFI5709-AF15712

AF332885-AF332887) (AY055112AY055115) MADS-box

, MADS-box

.

MADS-box ,
,

- -

. MADS-box
() ,

Copyright & A K-C

().
3 4 MADS-box apetalal.
, MADS-box
, , -,
.
, ,
MADS-box

, .
.

.
.
, , ,
, , ,
, 111 , 1 .
119 , 2 32
.

.
.
(Secale montanum Guss. ssp. kuprijanovii),
(Secale. cereale L.), , ( 9585
9693, ) (Triticum aestivum L.,
1, Triticum spelta ssp. asialbishicatum dorofeevii, Triticum compactum ssp.
fetissovii, Triticum sphaerococcum ssp. Rotundatum; 42790, 45369, 35633,
23769, )

(Hordeum vulgare L.), , (Zea

mays L.), 188. ..
.. (, . ) .. ,
( ).
3

Copyright & A K-C

15- ,
Saghai-Maroof (Saghai-Maroof et al., 1984).
E.coli DH5 JM109
pGEM-T ("Promega", ).
MADS-box
pBluescript SK+ ("Stratogene", ),
R.J. Schmidt . A. Ambrose
( . , -, ) A. Kleinhofs D. Kudrna
( . , , ).
-
(Maniatis et al., 1982).
"" ().
2. .
2.1. MADS-box
agamous-like apetalal .

MADS-box ( 1) -box (
3 5) MADS-box .
MADS-box -
MADS-box ,
MADS-box
- agamous-like apetalal.

MADS-box

1)

,
agamous-like, agamous squamosa!apetalal.
, 1,
1 MADSbox . , ,

Copyright & A K-C

,
MADS-box .
MADS-box .1.
MADS-box

1-region

El
AglFAplF
Mf I
Mfl * *Mfl *-

K-box

II

V. 12

-* Mrl

14 5
-AglR
- AplR
- Ki 2

2
Kf3
Aglinner
-*Aplinucr

El 180
E1-E3-1200-7000
E3-Z5-450-550
E1-E5-1700-7500

. 1. MADS-box agamous-like apetalal

. '- ; - , I ;
() 5' 3'- ();
agamous-like - AglF, AglR, Aglinner, Kf2, Kr2, Mfl, Mrl; apetalal
- AplF, AplR, Aplinner, AplF2, AplR2, KG, Kr3, MF1, Mrl.
, Mrl ,
. 1
MADS-box agamous-like .
GenBank ,

, MADS-box MADS-box
TaMADSU (GenBank accession AB007S05) Zag3 Zag5 (L46397
L46398), agamous-like.
-box Kf3
( 480
..)

MADS-box HvMADSS (AJ249144),

TaMADSU (AB007504), FDRMADS6 (AF139664)
5

Copyright & A K-C

Zapl

(L46400),

(ScMADS2)

MADS-box

(SmMADS2)

apetalal

apetalal.

GenBank accession AF159711 AF159712, .

.
500 ...

MADS-box

apetalal

TspMADS, TsphMADS, TcomMADS TaMADSl (GenBank

accession AY055112-AY055115, ).

MADS-box ScMADS2 SmMADS2,
TaMADSl I (AB007504).

2.2.

MADS-box
.

(ScMADSl,

SmMADSI,

ScMADS2, SmMADS2)

(TspMADS, TsphMADS, TcomMADS TaMADSl)

MADS-box ( !).

ScMADSl

SmMADSI

agamouslike2/agamouslike6. (ScMADS2 SmMADS2)

(TspMADS, TsphMADS, TcomMADS TaMADSl)

apetalal.

- -box :
, - -
MADS-box , , ,
, C-A-rich-G-taw
6

Copyright & A K-C

, (Riechmann et
al., 1996; Riechmann et al., 1997; Theissen, 2001).

K-box

. ,
3 4 apetalal . 3 4
agamons Zag2 (X80206) Zmml (X81199) ,
APETALA]/SQUAMOSA

(63701), (Z16421)

(AF126723-AF126728 AF126734)
' .
(ScMADSl
SmMADSl,

ScMADS2

TcomMADS,

SmMADSl)

TaMADSl)

(TspMADS,

TsphMADS,

MADS- .
MADS-box (ScMADSl

SmMADSl)

1 ,

apetala3/pistillata,

agamous

MADS-box

agamouslike2/agamouslike6

apetalal,

(.

5).

MADS-box . (ScMADSl SmMADSl)

agamouslike2/6. ,

, -
.

(Schmidt and Ambrose, 1998; Theissen et al., 2000;
Kyozuka

et

al.,

2000).

AGAMOUS-LIKE6 Zag3
Zag5

TaMADS12

,
7

Copyright & A K-C

1. MADS-box ,
(% /
)

agamous*

SmMADSl 78-91/81-88
78-93/79-87
ScMADSl
SmMADS2
ScMADS2
TspMADS
TcomMADS
TsphMADS
TaMADSl
-

MADS-box
agamouslike2/
apeialal***
agamouslikeS **
86-91/91-95
88-91/88-92
-

72-76/83-87
74-77/83-84
90-99/81-97
91-99/82-97
90-98/80-96
91-99/82-97
90-97/81-95
90-98/82-95

apetalaS/
pistUlata****
60-70/76-80
64-70/76-77
-

. ,
(GenBank accession numbers).
* agamous: (ZagI, LI8924; Zmml, X81199; Zmm2,
X&12Q0; Zag2, X802O6; Zag4. L46399; Tmzl-15a, Tmzll5a, Tmzl-120a\\Zmm4\
Zeml-Zem3, X85334-X85336, Zmovl3 Zmov23, U31521 1522; Zag
homologue, L81162) OsMADS3, L37528.
** agamousIike2/agamomlike6: (TaMADS12, 007505),
(OsMADSI, L34271; OsMADS5-OsMADS8, U78890, U78782, U78891
U78892; OsMADS24 OsMADS45, U32109 U31994; OMADS79, Y15008);
(Zag3 ZagS, L46397 L46398; Zmm3, Zmm6-ZmmS)\
{SbMADS2, U49734) {HvMADS7 HvMADS9, AJ249145 AJ249147).
*** apetalal: {TaMADSlI, AB007504); (UMADSI
UMADS2, AF035378 035379, Gocal et al., 2001); (SbMADS2,
U32110) (Zap], L46400).
**** apetala3/pistillata\ (TaMADSSl, AB007506)
(OsMADS2, OsMADS4 OsMADS16, L37526, L37527 AF077760.
( et at., 1995;
Murai et al., 1998; Theissen, 2000). ,
AGAMOUS-LIKE2,
, , OsMADS24 OsMADS45,
, (Greco et al., 1997).

Copyright & A K-C

xagS i

ScMADS2

taxnad&12 p ^
osmadsd j
agl6*
osmads7
tmm7

sdmadst
osmadsS
o*rt\ads?9
osrnads4S
os mads 24
agl2
smmads 1
scmads 1
xmin3
osmadsS
zmrn8
osmads 1
tmil-125a
z e m l ]
z e m 3 i r~
z e m 2 r"^
*ag4
t m z 1 -15a
tmzl-l20a
r a g h j
z a g l l
osmads 3 '

zmm2

SmMADS2
TsphMADS

TcomMADS
TaMADSI
TspVAOS
TaMADS11
HvMA0S5
UMADS1

FDRMADSS
OsMADSU
RAF1B

11MADS2
HvMADSS
0sMADS15
OsMADSIike
SbMADS2

ZmMADS3

zmov23 - |

Zap1

t m o v 13 i I
a>aR2~
J,tmads2
apl
Sbmads2
zmm4
tarnadsl
lemads
GS m a d s 4
osmads2
fc*mads5l i
o s m a d s 16

AP1
SOUA
HA1ADS3
FDRHMDS7
OsfMDSI

HvMADS7
OsUADSS

. 5 (). MADS MADS-box - squamosa/apetalal,

agamous-like2/6 agamous.
1.
agl2 agl6 - AGAMOUS-LIKE2 AGAMOUS-LIKE6
( GenBank accession 55551 55554}.
agamous-like2/6 .
.. 6 (). -
apetalal .
: ScMADS2 SmMADS2 (AF159711 AF159712); TspMADS,
TcomMADS, TsphMADS TahtADSl (AY055112-AY055115) TaMADSI 1
(007504); HvMADS3, HvMADSS, HvMADS7 HvMADSS (AJ249143 AJ249146); LtMADSl UMADS2 (AF035378 AF035379);
OsMADSM OsMADSIS (AF058697 AF058698), RAP1B (041020),
OsMADSl OsMADSS (L34271 U78890), OsMADS-like (AB003325)
FDRMADS6 FDRMADS7 (AF139664 AF139665); SbMADS2 (U32110);
Zap) (L46400) ZmMADS3 (AF112150); {API, Z16421)
(SQUA, X63701).
9

Copyright & A K-C

, .
OsMADSI,

apetalal-

.
, , , ,

(Prasad et al., 2001; Theissen, 2001). ,
(ScMADSl SmMADSl)
agamouslike2/6

(. 5).

, 1
Zag3 TaMADSI2 (90% ),
OsMADSI (88% ). 1,
Zag3
TaMADSl 2.
MADS~box
2.
2. MADS-box ,

1

90 % TaMADSl2

(007505)
ScMADSl

SmMADSl 90 % Zag3 ( L46397)

ZagS (L4639S)
95 % TaMADSl 1
ScMADS2

(007504)
SmMADS2

84 % Zapl (L46400)
TspMADS
97% TaMADSl 1
TcomMADS
(007504)

TsphMADS
86 % Zapl (L46400)

TaMADSl

10

Copyright & A K-C

K-box

MADS-box

apetalal/squamosa (.6) -

apetalal ,
in situ .
apetalal
-box
TaMADSlI , HvMADSS , LtMADSl
OsMADS14/RAPlB (Murai et al., 1998; Kyozuka et a)., 2000; Schmitz
et al., 2000; Gocal et al., 2001).
,
, -
apetalal. ,
MADS-box apetalal
apetalal .

APETALA1/SQUAMOSA

,
(Ng and Yanofsky, 2001; Theissen, 2001).
apetalal
: , apetalal,
, - .

,
,
. ,
.

11

Copyright & A K-C

2.3.

MADS-box

.
MADS-box

MADS-box .
,

MADS-box ,

apetalal,

agamous agamouslike, .

.

, .
I.

TaMADS12 (AB007505),
(. 7).

5'

3'
MADS-box
El

I-region

K-box

II

E2

AglF
AgF Mf 1
Mfl
Mfl .

12

13

E5
AglR
AgR
-AglR2
AgR2

AglF2
AgF2
AglRl
- AgRl
AgE2F

- - - AgR
AgR2

. 7. MADS-box agamous-like agamous

. F2 R2 - .
12

Copyright & A K-C

agamous-like
.
.

MADS-box

(ScMADS3) (SmMADS3) AF332885

AF332886,

ScMADS3 SmMADS3
MADS-box Zag3 (L46397) Zag5 94-96/95-98%,
. ScMADSl
SmMADSl (AF159710 AF159709) 1 ScMADS3
SmMADS3

( 89/90 89/86 % ),
. ,
. ,
agamous-like
3-5 3 4.
ScMADSS (AF362364).
MADS-box
TaMADSl2 (AB007505).

agamous.
MADS-box Zagl (L18924),
OsMADS3 (L37528). MADS-box
(ScMADS4) {SmMADS4)
AF332887 AF346894, .

KF1 KR1, -box MADS-box

, 190 ..,
.
13

Copyright & A K-C

:
3 5, 3
4.
Zrnml Zag2 (X81I99
80206). MADS-box
agamous Tkz-1 (AF276682).
,
(ScMADS3, SmMADS3, ScMADS4 SmMADS4)

MADS-box

(Bennetzen, 2001; Harrison et a!., 2001).
ScMADSS (AF362364)
(AF276682)
-, MADS-box
, ,
,
(Bennetzen, 2001).
, NetGene2,
,

MADS-box

ScMADSS .

agamous-like (ScMADSS)
NetGene2 ,
3 ,
5'- . 3 - 48 5' 4 branch-point-,
3 4 3"- (. 11).

14

Copyright & A K-C

<

branch-point

^ ^ ^ ^ g
AACTCAGAGGTTccattccactt

5'

!
i

ttttgagcagaCACTTGCTT

. .

ScMADSS (AF362364), NetGene2
(NetPIantGene server, http://www.cbs.dtu.dk/NetPlantGene.html)
,
AGgt TGgt,
.

5' GGCCAGGAAG GTAAGACATGT 3* Zmml ( 4- 4)

AGGAAG GTACTG ATTT

TspMADS ( 4- 4)

-box agamous ()
,
,
3
4 . 3, 4
4 (. 12).
NetGene2 branch-point-
4 4 5, ,
,
4. ,
3 3 AGgt

Copyright & A K-C

, NetGene2
Zmml.
Zmml .

13

Tkzl
1 CTACCAGCAGGAATCACCAAAGCTGCGCAACCAGATCCAGATGCTGCAAA-CAACTAACAGgtagag
66
Z.Tjnl 3693 ....ft.cGT***A^****^****A*-*"*******
3158
Zzq2 4760 A**-*"G*"*A' > ****
.......*,....._,,........... 4 8 2 s
It
Tkzl
67 ttctgatgtaagtaacttagagttcgtgcaggctaggttgtttatttgtcagactctaattgattgaa
136
ZirJitl 4213 " " ,
.>4[.>...t.at..."'..4....."......>...g....ct....
4250
E5
Tkzl 137 ttcttqtttcaqAGTGAGCTGCTGTCTGCTGAGRTTGCA-TTACATGGCAAAAAG
190
433J
ZOTI 4281 . - . . . . . - . . * . G . C * " * - - " A * * * " " * * C * * * * *
,
2ag2 5269 *.' .*.....5....G-*> *G*"'*> c*** 5321

. 12.

Ffc7(AF276682) NetGene2;
branch-point- 4 ;
.
, ,

. ,
. ,
MADS-box
(Riechmann et al., 2000)
, "" MADS-box .

.
1. MADS-box ,
MADS- - ,
- - .
2. MADS-box
() ,
().

16

Copyright & A K-C

3. 3 4 -box MADS~box
apetalal .
4. MADS-bcx
.
. .
1. ., . , . , . .,
.., MADS-box
!88 // . 1999. . 46 .
600-604,
2. .., .., .., MB., ..,
.., .. MADS-box
// . 2000. . 47. . 100107.
3. ..^ .., . , ( ..,
.. -box MADS-box //.
. 2000. . 47. . 631-633.
4. Ermishev V.Y., Karyagim A.S., Khavkin E.E., Narodltskii B.S., Zabrodina M.V.
A putative transposed -box homolog of zmml and zag2 II Maize Genetics
Cooperation Newsletter. 2000. V. 74. P. 53.

/ .

. 2 3

127550, , . , 44

Copyright & A K-C