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,2016,36(6):1278-1284
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J K:` abc:defg(MAPK)hij、k$l#$mno1$pqrstluvwxyz{P|}~O
,
MAPKKK、MAPKKl MAPKm3M,t41$
1$、f、Ma7#$1
mpHQ
。
MAPK Oy、t#¡¢y1$%£¤¥7
MAPKOy¦c9§¨©ª,«t¬® MAPKO¯~yP|}~56y=>°±²³´µ。
LMN
:`
abc:defg
(MAPK);P|}~;1$%£¤
OPQR$
:Q257 !STUV:A
犚犲狊犲犪狉犮犺犘狉狅犵狉犲狊狊狅犳犕犻狋狅犵犲狀犪犮狋犻狏犪狋犲犱犘狉狅狋犲犻狀犓犻狀犪狊犲
犛犻犵狀犪犾犜狉犪狀狊犱狌犮狋犻狅狀犘犪狋犺狑犪狔
JIANGShengxiu,LIDelu
(GansuPsammophyteEngineeringTechnologyResearchCenter,MinqinDesertBotanicalGarden,Minqin,Gansu733300,Chi
na)
犃犫狊狋狉犪犮狋:Mitogenactivatedproteinkinase(MAPK)cascadesarehighlyconservedsignalingmodulesfound
inaleukaryotes,includingfungi,plantsandanimals.AMAPKcascadegeneralyconsistsofthreecompo
nents:aMAPKKK(MAPKKkinase),aMAPKK(MAPKkinase)andaMAPK,andtheyplayessential
rolesinabioticstresses,hormones,celdivisionandplantgrowthanddevelopment.Inthisarticle,weout
linedthecompositions,biologicalfunctions,inactivationofMAPKcascades,whichaimedatprovidingsome
referencesbasisfortheresearchofMAPKmediatedsignaltransductionmechanisms.
犓犲狔狑狅狉犱狊:Mitogenactivatedproteinkinase(MAPK);signaltransductionpathway;biologicalfunctions
¶·¸lP|MN¹º#$y1l
,
y»¢deO¼½P|MN¾
¿8ÀÁÂÃÄ
,
IÅÆÇzÈÉy1$:%Ê4
7deËÌyÍÎQ
,
ÏÐÑÒÓP|
}~
[1]。
P|}~hno4¸P|7¹
ÔÕÖ:yz·H×Ø
,
tÙÚ
、
Û
ÜÝÞ
、
Ößl1smpHQ
[2]。
¸àáÁ
,
#$P|}~OâM¬ã
äGH{/
:`
abc:defg ×Ø
、Ca2+
´åP|}~O
、Ca2+´åyæçè(SOS)P
|Ol ABAP|}~O[3]。8p,` abc
:defg
(MAPK) ×Øhij、k$l#$
mno1$pqrstluvwxyz{P|}~
O
[4]。MAPK ×ØéêëУ¤ìí y
defg
:`
abc:defgfgfg
(MAP
KKK)、` abc:defgfg(MAPKK)l`
abc:defg
(MAPK)[5]。îïy MAPKO
è4îïy¸P|ðf
。
ñòóô
,
t#$p
,
MAPKOt41$
1$、f、M
a7#$1mpºH
Q
[4]。
1 MAPK Oy
MAPK × Ø MAPKKKMAPKK
MAPKëfgÈõ,MAPKK ÑöÂy
MAPKKK÷ø:,÷ø:ùú¬Ser/Thrû,,
ùúü,øýɬ SXXXS/T(X Ýþÿ!ü,
ø
),
Ê
,MAPKK÷ø: MAPK,÷ø:ù¬
Thr" Thrû,,#$P|%&ä¾ÃÄ
ÁÂ
[5]。
1.1 6789:;<=>=>=>(MAPKKK)
MAPKKKh Ê4y(zM,Ñ)¬h
*+MAPK ×Øp,ÔÕl-.,/yM。
012
(犃狉犪犫犻犱狅狆狊犻狊狋犺犪犾犻犪狀犪)pòÑ3)80/
·
,
ÑM¬3Ð45:MEKKs45,21Ð;Rafs
45
,
48Ð;ZIKs45,11ÐF[6]。tRafs
45p
,
¶defgy6789Mwx
,
8p£¤:
¦;Ö,<犆犜犚犾(Constitutivetripleresponse1)
t=>¯~yP|}~×Øpìþ?
,
@AB
¹N
,
C6=>D~,
,<犈犇犚1t#$YEF4pÇB¹Q[8],
犃狋1犵73660hz·Raf/ MAPKKKF,â¥G
H012yIæì
[9]。MAP3Kpy MEEKâ¥
Ñ÷ø:fc
(
J¬ MAP4K),Kâ¥Ñ Ras"
RhoL5yMGTP6Nde6NÅO1cì[10]。
MEKKK/犃犖犘1,
UV W
(犖犻犮狅狋犻犪狀犪狋犪犫犪犮狌犿)MEKKK / , <
犖犘犓1¤ÙX},<#$
1$yI»
ì
[12]。AtMEKK1lAtMEKK4â»YZ、5[
\]l^byD~
[6],AtMEKKs4py AN
PKls_ H2O2 yP|Ã~[13]。NPKld
efgä.st`abb,lbcdeM1fË
Ì
,
_#$yMalÙÚ
[14]。ZIKs45
pydefgGH_MAPKs×Øy¹NZRly
þ?Q
[15]。
1.2 6789:;<=>=>(MAPKK)
#$ MAPKKt MAPK ×ØpyF-
.,M
,
*+¬Z
,
012,<pò&g¨hi
60Ð MAPKKK l20Ð MAPK,jk10Ð
MAPKKÑñ。öÂy MAPKKK è4¸P
|
,
O10Ð MAPKKÃÄ20Ð MAPKp,þ
ô MAPKKhöÂLáP|ylmú,KhÁÂ
MAPKyMnú[5]。MAPKKhoRßìfg,
#$ MAPKK,iyP|ýÉhSTXXGTX
XYMXPER,#$ MAPKKwxyS/TX5
S/TýÉ,Åk$fgyýɬS/TX3S/T;bµ
ü,øýÉp¿012y MAPKKM¬4Ð,
M½¬A、B、ClD,8pClDy MAPKK
y@A,<îqqN
。
#$ MAPKK yNer
sþô¨zÐtuy MAPKvwú,ýɬ[K/
R][K/R][K/R]x(15)[L/I]x[L/I],{xtk$
MAPKKpñyýÉ[15]。y,MAPKKzu
y{$Rßì
。
1.3 6789:;<=>(MAPK)
MAPKh Ê4y,®zM,t|}öÂ
MlÁÂ{$pÇHQ
[16],MAPK¤÷
ø:Rß~4de
,
IÅfcyF4
。
*
I
(犕犲犱犻犮犪犵狅狊犪狋犻狏犪 Linn.)p ñ
MAPK¥,ht#$pòä. MAPKÑ
ñ
,
,<
ýt012p 20 Ð
MAPK[17],(犗狉狔狕犪狊犪狋犻狏犪)17Ð[18],
(犘狅狆狌犾狌狊)21Ð[19]。bµ@AydeýÉy
TXY,ýyîï,â¿8M¬TEY{lTDY{,
bµÈõ1È¿ TEY{9ÅM¬ A、B、C
,TDY{ɬD[20]。µ&g,A{犃狋
犕犘犓3、犃狋犕犘犓6l 犗狊犕犃犘犓5;B{ 犃狋犕犘犓4
l犣犿犛犐犕犓1;C{犃狋犕犘犓1、犃狋犕犘犓2、犘狊犕犘犓2
l犌犺犕犘犓7。Ï MAPKt#$F4l1
pºHQ
[21],
U012y犃狋
犕犘犓3âÑ/·àáfc,Q:K¤fc
fg
[22];
4 MPK4yRßìK¢M犃狋
犕犘犓4²1$l
1$4S[23];犘狊犕犘犓2
t012pþ?8cì»5[]¥78
P|UABA、H2O2 mD~[24]。IN#$p
T¨y D { MAPK £¤,U 犗狊犅
犠犕犓1D~犘犚,
[25];犃狋犕犘犓9¡¢tw£
pþ?
,
P£¤¤¥ì
,
tABAP|Op
cìQÁÂǦyQ
[26];犃狋犕犘犓18¤
97216§ ?1@,m:#$`abc:defg P|}~O=>9¨
¹#$©ª«¬y£¤
[27];
®
(犌狅狊狊狔狆犻狌犿
spp.)犌犺犕犘犓16 yþ? » : % l 1 $ P | y
D~
[28]。
2 MAPK Ê4y£¤
#$t¯Ð1°±§p²³»¶·1$l
1$
,´
µ
、
æl¶·mh¸#$GOyGH
<
,
¬4¹º1sy»u<
,
#$Pè¼l
ÃÄðfP|y¤
,
8p
,MAPK O¿è
¼yðfP|ÃÄp
,
+®1$¢Od
e½y÷ø:l¾÷ø:DñLáy
[29]。
2.1 犕犃犘犓WXY45:ZA$[*+òz/óµóô MAPK×ز_¨#
$:%P|×Ø
,
=>¿/y#$ÀP|MNp
,
ø
(SA)、ÁÂø(JA)lÃÄ=ø(IAA)m。
犗狊犅犐犕犓1,
[30];犗狊犕犛犚犕犓2þ?»JAD~[31];犗狊犛犐犘犓
Åu§ymRNAMþô,
,
ÈàÉÊ4Ë
(CHX)、
JAlSAÌÍÎ,犗狊犛犐犘犓 y}ÏÐÑÙX[32];
犃狋犕犘犓4tSAOpÇBQ,ÅtJA
OpǦQ
[33]。SAq.yÒÓîÔ¸
yÕÖ
,
KÑfc MAPKP|O。ULiu
m
[34]
ñä×
(犌犾狔犮犻狀犲犿犪狓)p犌犿犕犘犓4狊yØÙ
~ÚÛyÕÖlSAq.yÜu。
2.2 犕犃犘犓WX45]5^_`YA$aC
*+¸¶·^b¢Ýº#$y1
,
#$*
Þߨ/·YEàá
,
â¨:%l$Íãäàá
,
åO^b¢D~àáwæ*É
,
éêç
Ùè
、
O1#$Ké
、
fcYE,
Suzukim[36]Iêë(犘犺狔狋狅狆犺狋犺狅狉犪犻狀犳犲狊狋犪狀狊)
ì1Åyz·D~NÌÍVWíîÎñ
,
47kDay MAPK ïðÅñòóÑfc。Zhang
m
[37]
ñÈVWè®^éÎ
,
VWp2·
MAPKÑfc,M½h48kDyø(SA)D~
defg
(犛犐犘犓)l44kDyô]D~defg
(犠犐犘犓)。Hem[38]IpõöT¨z·
MAPK,<犅犠犕犓1,,
(犛狆狅犱狅狆狋犲狉犪犾犻狋狌
狉犪)yI»ì[39],犌犺犕犘犓16t012pyßùþ
?ÙX¨#$yK^ì
[38]。NorthernÕü,
犌犺犕犘犓7y}Ïл^b¢D~,}:VW®,
},
!SA O,
[40]。
â"
,MAPK²_#$1$ÁyP|
Ã~
。
2.3 犕犃犘犓[\b45=cA$BCDEOY
0d
#$ø(ABA)t·N%、&¹l´µ、
æ
、
¶·m
1$pÇzuyQ
,ABA
Láy34QGHhOD~(Hy
Í,
t)*ªp
,ABAï
ðÅñòófc MAPKycì,Å+ë(GA),
C6 MAPKy}ÏÐ。犘狊犕犃犘犓3t-×(犘犻
狊狌犿狊犪狋犻狏狌犿)pyþ?» GA、6BAD~[41],Å}
犘狊犕犘犓2,<012p犘狊犕犘犓2l犃狋犕犘犓17
犃狋犕犘犓2z.P ABAl H2O2 I»ì[42];01
2p ABA f c 犃狋犕犘犓3 l 狆46犕犃犘犓 y c
ì
[43],
Åp犗狊犕犃犘犓5ycì» ABA yf
c
[44];
®犌犺犕犘犓6,<²_ABAD~y犆犃犜1
yþ?
[45];ABA â¥D~/0犣犿犕犘犓5i.þ
?
[46];犌犺犕犘犓2tVWpþ?Ú$#$1¶
ABAyçèì[47]。
=>h#$1°ckpyH¹$½
,
²_
#$y2D3l®45m
,
KD~YE
¢Èy£¤
,
6U
,
ÈVW»^b¢7Î
,
8
Û8ÑïðD~dey÷ø:
,
IÅ^bÇC
6Q
[48]。犜犚1、犈犜犚2l犈犐犖4h=>y»¢,
犆犜犚1@Az·l RafL5Íxyde;犆犜犚1t
=>P|OpQ¬B
¹t=>»¢y
ÁÂ
,
=>»¢6N¾fc犆犜犚1[49]。z/9)¬
MAPKO²_¨=>P|}~,U=>ÌÍV
WÛ®Ñfcz·50kDäy MBPfg,Ï·
fgh犆犜犚1ÁÂyz· MAPKfg[50]。
#$1t#$1lpKÇz
Hy¹Q
,
Ude¡:
、
ablb;yß7
«¬yÖßm
。
z/=>ñ¨defgl÷øg
t1P|OpyQ
,
¢óô¨MAPKh
²_1P|}~yz·fg
,
¿:i1y
VWBY2N1,J2,4<=>=ø
(2,4D)9§ÌÍ,6246kDy MBPfgÑïð
fc
[51];
#
,
1ÌÍ®
,
z·÷ø:
犃狋犕犘犓2defgycìKT¨ÙX。Ï6
2þô MAPKKl MAPKt1¯~yP|}
~pºzuyQ
[52]。
0821 ! " # $ % & 36?
2.4 犕犃犘犓Xe]5^_A$YLf
#$t1sypH4¶·
1$
,
éê´µ
、
u·¶·lYZm
。
6U
,
t012
p
,
@læ¤D~A¯y MAPK ×Ø:
犕犈犓犓1犕犓犓2犕犘犓4[53],犕犃犘犓犓4 l 犕犃犘
犓犓6Ñ@læD~,Å 犕犃犘犓犓1Ñæl´µD
~
,犕犃犘犓犓102kÑ@D~[54];VWyzÐ
MAPKKL5犛犐犘犓犓,È#$»ô]Ë®8þ
?.Üu
[55];
ä×犌犕犓1ycìtæÁÑf
c
[56];
BC
(犆狌犮狌犿犻狊狊犪狋犻狏狌狊Linn.)犆狊犖犕犃犘犓 t
},
D2
EFGuN
(犕犪犾狌狊犺狌狆犲犺犲狀狊犻狊Rehd.var.狆犻狀狔
犻犲狀狊犻狊Jiang)ylbp犕犺犕犃犘犓 ymRNAÐ
Ñ´µlæ\D~
[58];
tælYZÁ
,犜犕犓犘1
tMH
(犜狉犻狋犻犮狌犿犪犲狊狋犻狏狌犿 Linn.)pÑD~þ
?
[59];
IJN
(犛犪犾犻犮狅狉狀犻犪犫狉犪犮犺犻犪狋犪)犛犫犕犃犘犓犓
y}ÏÐѵ
、
@læD~
,
¾tæD~Áy}Ï
Ð,u
[60]。犣犿犛犐犕犓1t012pyþ?Ù
X¨#$yIæì
,
¾!D~¨Í,<
犚犇29l 犘5犆犛1yþ?[61]。p 犗狊犕犃犘犓44
ycì»æ
、´
µlQ:D~
,
jî»@D
~
[62],
Å犗狊犕犃犘犓33t´µÁ8}ÏÐÜ
u
,
ÅtæÁ}ÏÐÁ1
[63]。
®p
,
犌犺犕犘犓2l犌犺犕犘犓16PI»YZy¤
[64],
¿犌犺犕犘犓3}:VWpÙX¨},
[65]。
cìQ
(ROS)h/·P|OypÌP|
MN
,
1$~Ú ROSt#$¢yÒÓ,
MAPK OîÔâ¥ÑROSD~,Kâ¥
¹ROSq.。tYZÁ,VWpþ?y
犣犿犕犘犓7,
DSM1OROSyKâ²_l´µyP
|O
[67]。
È#$»ô]Î
,犃狋犕犘犓8O¯
NROS、Ca2+ldey÷ø:QROSy¢
ÐLǹQ
[68]。
y
,
òóµþô¶·MFN/#$dey
÷ø:ùú
。NorthernÕü62þô,犣犿犕犘犓17
y}Ïл/·LáD~
,
tVWpyþ
?ÙX¨#$¶·yI»ì
[69]。Berberichm[70]
I/0pMOý¨犣犿犕犘犓4,}wVWpñ
,<ÙX¨},<#¡¶·yI»ì
。
PQR
(犗狉犻犵犪狀狌犿狅狀犻狋犲狊L.)犗狅犕犃犘犓犓1t¶·Á
þ?.ôÜu
[71]。
t¶·SÁ
,犗狊犕犃犘犓2y
mRNAÒÓ.TÜu[72]。
3 MAPKOy¦c
MAPK ×Øy¦clc:ïmH,
MAPKy¦chOTXY8USer"Tyry¾÷
ø:¹
。
*+
,
òõöT¨zijlk$
y÷ø:g
,
ñ¹ºP$ MAPK¦cy£¤,
Ï{ghiâ¥ÑM¬3{:oRßì÷øg
(DsPTPases),¤$ MAPKtSer"Tyrùú¾÷
ø:
;
Vüø÷ø:g
(PTPases),ktTyrùú¾
÷ø:
;`
üø
/
Wüø÷ø:g
(PPases)[52]。
DsPTPasesPcìùú VXVHCXXGXSRSX
TXXXAY(L/I)M,ÏÐRßy8lPTPaseïÀ
ì
,
<#
,
XâY PTPasel DsPTPasesZ¬ï
z{
[73]。
[
,
z/=>óô¨um#$p MAPK
yfc\]defgpTyry÷ø:,Ï=>¬
MAPK*÷ø:"TyrùúÑ÷ø:°±¨óµ,
ïÎKóô¨#$PVüø÷ø:g
,
tTyrù
ú¾÷ø:¾!$ MAPK¦c[74]。012oRß
ì÷øg犃狋犇狊犘犜犘1t Tyrùú¾÷ø:,¾$
犃狋犕犘犓4¦c,~Ú犃狋犇狊犘犜犘1tîïyÁ
`îïfp/þ?
[73]。
y
,
@APTPase
yz,
¶·yC6
,犃狋犘犜犘1P¾÷ø:£¤,t¢¤
$ z · MAPK ¦ c[76]。Gupta m[77] t = >
MAPK cìyD^pñ,犃狋犕犘犓6ycì»
H2O2 ¹,t¢犃狋犘犜犘1¤$犃狋犕犘犓6¦c;
,I-×lä×pKMOTAtPTP1[78]。Ha
ringm[79]I_`a(犆犺犾犪犿狔犱狅犿狅狀犪狊)pMOý¨
z·oRßì÷øgVHPTP13,P¾÷ø:y
£¤
,
<#t¢¤$b®py犛犐犕犓 l
犕犕犓2¦c。
4 ¨ c
MAPK ×ØQ¬P|}~×Øp
HyM
,
òdd/yMAPK,põöT
,
¾8£¤e¨¨=>
。
*+
,
`
MAPK,<£¤lQbÍy=>khñfgª,
åstz/hi
,
<#
,
MAPK ×Øy/
jkål9zmno
。
GHéê¥ÁpÐjk
:
(1)#$t9:pߨzq
³Aryè4
¸ðfy56
,
¶·¶.ydefg7ïz{f
18216§ ?1@,m:#$`abc:defg P|}~O=>9¨
gyîï4{ËÌtP|ÃÄpüsQ
,
\¥tñ=>,töuH9zmvK¶·def
gËÌyÍÎÈ
;(2)9zmwô MAPK ×
ØyÁÂJ7t#¡¢¯~y1:ÝÞl1Í
;(3)wô#$p_ MAPK ×ØÍ
yP|}~MN×Ø
,
¥7MAPK ×Ø_#$
p8P|×Øyk2ÎQÈ
。
Wg!S
:
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abioticstressesinplants[J].犆狌狉狉犲狀狋犛犮犻犲狀犮犲,2005,6:1771
1780.
[2] LIUY,ZHAOHY.Acomputationalapproachforordering
signaltransductionpathwaycomponentsfromgenomicsand
proteomicsData[J].犅犕犆犅犻狅犻狀犳狅狉犿犪狋犻犮狊,2004,10:16.
[3] XIONGL,SCHUMAKERKS,ZHUJK.Celsignaling
duringcold,droughtandsaltstress[J].犘犾犪狀狋犆犲犾犾,2002,1:
165163.
[4] HWACHM,YANGXC.TheAtMKK3pathwaymediatesABA
andsaltsignalingin犃狉犪犫犻犱狅狆狊犻狊[J].犃犮狋犪犘犺狔狊犻狅犾.犘犾犪狀狋,2008,
30:277286.
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