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Research Progress of Mitogen activated Protein Kinase Signal Transduction Pathway

植物丝裂原活化蛋白激酶级联信号转导通路研究进展



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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 MAPKm3M‚ƒ,t4„1$…1$†‡、fˆ、‰ŠMa‹Œ7#$1Ž
m‘pŽ’“HQ”

•–„—˜™š›œž MAPKŸ Oy‚ƒ、t#¡¢›y1$%£¤¥7
MAPKOy¦c9§¨©ª,«t¬­® MAPKO¯~yP|}~56y=>°±²³´µ。
LMN
:`
abc:defg
(MAPK);P|}~;1$%£¤
OPQR$
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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:Mitogenactivatedproteinkinase(MAPK)cascadesarehighlyconservedsignalingmodulesfound
inaleukaryotes,includingfungi,plantsandanimals.AMAPKcascadegeneralyconsistsofthreecompo
nents:aMAPKKK(MAPKKkinase),aMAPKK(MAPKkinase)andaMAPK,andtheyplayessential
rolesinabioticstresses,hormones,celdivisionandplantgrowthanddevelopment.Inthisarticle,weout
linedthecompositions,biologicalfunctions,inactivationofMAPKcascades,whichaimedatprovidingsome
referencesbasisfortheresearchofMAPKmediatedsignaltransductionmechanisms.
犓犲狔狑狅狉犱狊:Mitogenactivatedproteinkinase(MAPK);signaltransductionpathway;biologicalfunctions
  ¶·œ¸l›P|MN‹¹º#$y1l
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ÏА‘ÑÒӉŠP|
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[1]。
P|}~hno‰Š4„œ¸P|7‹¹
ÔՊ›Ö:yz·“H×Ø

t‹Œ‰ŠÙÚ

Û
ÜÝÞ

Ößl1sm‰Š‘pŽ’“HQ”
[2]。
œ¸àᆇÁ

#$P|}~OâM¬ã
äGH{/
:`
abc:defgŸ ×Ø
、Ca2+
´åP|}~O
、Ca2+´åyæçè(SOS)P
|Ol ABAP|}~O[3]。8p,` abc
:defg
(MAPK)Ÿ ×Øhij、k$l#$
mno1$pqrstluvwxyz{P|}~
O
[4]。MAPKŸ ×ØéêëУ¤ìí y
defg
:`
abc:defgfgfg
(MAP
KKK)、` abc:defgfg(MAPKK)l`
abc:defg
(MAPK)[5]。îïy MAPKO
è4îïyœ¸P|ðf

ñòóô

t#$p

MAPKOt4„1$…1$†‡、fˆ、‰ŠM
a‹Œ7#$1Žm‘pŽ’º“H

[4]。
1 MAPKŸ Oy‚ƒ
MAPK Ÿ   × Ø € MAPKKKMAPKK
MAPKëŸfgÈõ‚ƒ,MAPKK ÑöÂy
MAPKKK÷ø:,÷ø:ùú¬Ser/Thrû,,•
ùúü,øýɬ SXXXS/T(X Ýþÿ!ü,
ø
),
ʐ™
,MAPKK÷ø: MAPK,÷ø:ù¬
Thr" Thrû,,€#$P|%Ÿ&ä¾ÃÄ
ÁÂ
[5]。
1.1 6789:;<=>=>=>(MAPKKK)
MAPKKKhŸ Ê4y(zM,Ñ)¬h
*+MAPKŸ ×Øp,ÔÕl-.,/yM。
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,11ЃF[6]。tRafs
45p

¶defgy6789Mwx

8p£¤:
¦;Ö,<犆犜犚犾(Constitutivetripleresponse1)
t=>¯~yP|}~×Øp‚ƒìþ?

@AB
‹¹N

C6=>D~,[7],Raf/CTR1
,<犈犇犚1t#$YEF4pÇB‹¹Q”[8],
犃狋1犵73660hz·Raf/ MAPKKKƒF,â¥G
H012yIæì
[9]。MAP3Kpy MEEKâ¥
Ñ÷ø:fc

J¬ MAP4K),Kâ¥Ñ Ras"
RhoL5yMGTP6Nde6NÅO1cì[10]。
MEKKK/犃犖犘1,߆‡4S,[11],
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],AtMEKKs4‚py AN
PKls_‹Œ H2O2 yP|Ã~ž[13]。NPKld
efgä.st`abb,lbcdeM1‚fË
Ì

_#$‰ŠyMalÙڝž
[14]。ZIKs45
pydefgGH_MAPKs×Øy‹¹NZRly
þ?Q”ž
[15]。
1.2 6789:;<=>=>(MAPKK)
#$ MAPKKt MAPKŸ ×ØpyƒF-
.,M

*+¬Z

012,<‚pò&g¨hi
60Ð MAPKKK l20Ð MAPK,jk10Ð
MAPKKюñ。öÂy MAPKKK è4œ¸P
|

O10Ð MAPKKÃÄ20Ð MAPKp,þ
ô MAPKKhöÂLáP|ylmú,KhÁÂ
MAPKyMnú[5]。MAPKKho“Rßìfg,
#$ MAPKK,iyP|ýÉhSTXXGTX
XYMXPER,#$ MAPKKwxyS/TX5
S/TýÉ,Åk$fgyýɬS/TX3S/T;bµ
ü,øýÉp„¿012y MAPKKM¬4Ђ,
M½¬A、B、ClD,8pC‚lD‚y MAPKK
y@A,<îq›qN

#$ MAPKK yNer
sþô¨zÐtuy MAPKvwú,ýɬ[K/
R][K/R][K/R]x(15)[L/I]x[L/I],{xtk$
MAPKKpŽñyýÉ[15]。yœ,MAPKKzu
y{$Rßì

1.3 6789:;<=>(MAPK)
MAPKhŸ Ê4y,®zM,t|}öÂ
‚MlÁÂ{$pǓHQ”
[16],MAPK¤÷
ø:Rß~4de

IÅfc‰ŠyF4€

*
I ‚
(犕犲犱犻犮犪犵狅狊犪狋犻狏犪 Linn.)p ƒ „ Ž ñ
MAPK¥™,h­t#$pòä. MAPKю
ñ

,<‚…ý†‡ˆ‰t012p 20 Ð
MAPK[17],Š‹(犗狉狔狕犪狊犪狋犻狏犪)17Ð[18],Œ
(犘狅狆狌犾狌狊)21Ð[19]。bµ@AydeýÉy
TXY,ýyîï,â¿8M¬TEY{lTDY{,
bµÈõŽ1žÈ¿ TEY{9ŎM¬ A、B、C
‚
,TDY{É¬D‚[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。ϑ MAPKt#$†‡F4l1
Ž‘p’Ž’º“HQ”
[21],
U012y犃狋
犕犘犓3âÑ/·àᆇfc,Q:†‡K¤fc
•fg
[22];
4” MPK4yRßìK¢M“ˆ‰犃狋
犕犘犓4²”1$l…1$†‡4S[23];犘狊犕犘犓2
t012pþ?8cì»5[]•¥78–†‡
P|—UABA、H2O2 mD~[24]。IN˜#$p
™T¨š‰y D { MAPK £¤›œ,U 犗狊犅
犠犕犓1D~犘犚,žl‰žèŸy¤ 
[25];犃狋犕犘犓9¡¢tw£‰
Špþ?

P£¤¤¥ì

tABAP|Op„
cìQÁÂǦ‹ŒyQ”
[26];犃狋犕犘犓18¤
97216§           ?1@,m:#$`abc:defgŸ P|}~O=>9¨
‹¹#$‰Š©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 MAPKŸ Ê4y£¤
#$t¯Ð1°±§p²³»¶·1$l…
1$†‡
,´
µ

æl¶·mh¸ƒ#$GOyGH


¬4„¹º1sy»u<ˆ

#$Pè¼l
ÃÄðfP|y¤ 

8p
,MAPKŸ O¿è
¼yðfP|ÃĉŠp

+®1$¢Od
e½y÷ø:l¾÷ø:Dñ„Láy‹Œ
[29]。
2.1 犕犃犘犓WXY45:ZA$[*+òz/óµóô MAPK×ز_¨#
$:%P|×Ø

=>¿/y#$›ÀP|MNp

ŠŒø
(SA)、ÁÂø(JA)lÃÄ=ø(IAA)m。
犗狊犅犐犕犓1,D~
[30];犗狊犕犛犚犕犓2þ?»JAD~[31];犗狊犛犐犘犓
Åu§ymRNAM“þô•,ÆÇp«H‚ƒ/þ?

ȔàÉÊ4Ë
(CHX)、
JAlSAÌÍÎ,犗狊犛犐犘犓 y}ϊÐÑÙX[32];
犃狋犕犘犓4tSAOpÇB‹ŒQ”,ÅtJA
OpǦ‹ŒQ”
[33]。SAq.yÒÓîÔ¸
ƒ‰ŠyÕÖ

KÑfc MAPKP|O。ULiu
m
[34]
Žñä×
(犌犾狔犮犻狀犲犿犪狓)p犌犿犕犘犓4狊yØÙ
~ژۉŠyÕÖlSAq.yÜu。
2.2 犕犃犘犓WX45]5^_`YA$aC
*+¸¶·^b¢Ý†º#$y1

#$*
Þ߃¨/·YEàá

â¨:%l$Íãäàá
œ

åO^b¢D~àá™wæ*É

éꉊç
Ùè

O1#$Kéˆ

fcYE,[35]。
Suzukim[36]”Iêëž(犘犺狔狋狅狆犺狋犺狅狉犪犻狀犳犲狊狋犪狀狊)
ì1řyz·D~NÌÍVWíΎñ

47kDay MAPK ïðÅñòóÑfc。Zhang
m
[37]
ŽñÈVW蟮˜^éÎ

VWp2·
MAPKÑfc,M½h48kDyŠŒø(SA)D~
defg
(犛犐犘犓)l44kDyô]D~defg
(犠犐犘犓)。Hem[38]IŠ‹põö™T¨z·
MAPK,<犅犠犕犓1,•,h®þ?。µ,—y&g,犃犺犕犘犓3t},pþ?ÙX¨VW„øùúû
(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]。NorthernÕüˆ‰,
犌犺犕犘犓7y}ϊл^b¢D~,}:VW®,
},(犆狅犾犾犲狋狅狋狉犻犮犺狌犿狀犻犮狅狋犻犪狀犪)l^éPVYy¤ ,¾
!SA O,X
[40]。
â"
,MAPK²_#$1$†‡ÁyP|
Ã~

2.3 犕犃犘犓[\b45=cA$BCDEOY
0d
  #$ø(ABA)t·N%Ž、&‹¹l´µ、
æ

¶·m…1$†‡p’ÇzuyQ”
,ABA
„Lᆇy34Q”GHhOD~(Hy†‡
͞,
t)*ª‰Šp
,ABAï
ðÅñòófc MAPKycì,Å+ëˆ(GA),
C6 MAPKy}ϊÐ。犘狊犕犃犘犓3t-×(犘犻
狊狌犿狊犪狋犻狏狌犿)pyþ?» GA、6BAD~[41],Å}
犘狊犕犘犓2,<012p犘狊犕犘犓2l犃狋犕犘犓17
犃狋犕犘犓2z.P ABAl H2O2 I»ì[42];01
2p ABA f c 犃狋犕犘犓3 l 狆46犕犃犘犓 y c
ì
[43],
Ŋ‹p犗狊犕犃犘犓5ycì» ABA yf
c
[44];
­®犌犺犕犘犓6,<²_ABAD~y犆犃犜1
yþ?
[45];ABA â¥D~/0犣犿犕犘犓5i.þ
?
[46];犌犺犕犘犓2tVWpþ?Ú$#$1¶„
ABAyçèì[47]。
=>h#$1°ckpy“H‹¹$½

²_
#$y2Dƒ3l®˜45m‘

KD~YE
¢Èy£¤

6U

ÈVW»^b¢7•Î


Û8ÑïðD~dey÷ø:

Iń^bžÇC
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‹Œ
¹t=>»¢y
ÁÂ

=>»¢6N¾fc犆犜犚1[49]。z/9)¬
MAPKO²_¨=>P|}~,U”=>ÌÍV
W˜Û®Ñfcz·50kDäy MBPfg,Ï·
fgh犆犜犚1ÁÂyz· MAPKfg[50]。
#$1ˆt#$1lŽ‘pKÇz
“Hy‹¹Q”

Ude¡:

ablb;y߃7
«¬yÖßm

z/=>Žñ¨defgl÷øg
t1ˆP|OpyQ”

ƒ¢óô¨MAPKh
²_1ˆP|}~yz·fg

¿:i1ˆy
VWBY2‰Š”Nƒ1ˆ,J2,4<=>=ø
(2,4D)9§ÌÍ,6246kDy MBPfgÑïð
fc
[51];


”1ˆÌÍ®

z·÷ø:“‚
犃狋犕犘犓2defgycìKT¨ÙX。ϑ6
2þô MAPKKl MAPKt1ˆ¯~yP|}
~pŽ’ºzuyQ”
[52]。
0821 ! " # $ % &                   36?
2.4 犕犃犘犓Xe]5^_A$YLf
#$t1sy‘pH4„¶·…1$†‡

éê´µ

u·¶·lYZ†‡m

6U

t012
p

@l憇¤D~A¯y MAPK Ÿ ×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102kÑ@†‡D~[54];VWyzÐ
MAPKKL5犛犐犘犓犓,È#$»ô]Ë®8þ
?.Üu
[55];
ä×犌犕犓1ycìt憇ÁÑf
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
},[57];
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)y˜lbp犕犺犕犃犘犓 ymRNAŠÐ
Ñ´µlæ\D~
[58];
tælYZ†‡Á
,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}ϊÐѵ

@læD~

¾tæD~Áy}Ï
ŠÐ,u
[60]。犣犿犛犐犕犓1t012pyþ?Ù
X¨#$yIæì

¾!D~¨†‡Íž,<
犚犇29l 犘5犆犛1yþ?[61]。Š‹p 犗狊犕犃犘犓44
ycì»æ
、´
µlQ:†‡D~

jî»@†‡D
~
[62],
Å犗狊犕犃犘犓33t´µ†‡Á8}ϊÐÜ
u

Åt憇Á}ϊÐÁ1
[63]。
­®p

犌犺犕犘犓2l犌犺犕犘犓16PI»YZ†‡y¤
 
[64],
¿犌犺犕犘犓3}:VWpÙX¨},yIµìlIQ:ì
[65]。
cìQ
(ROS)h/·†‡P|OypÌP|
MN

…1$†‡~Ú ROSt#$¢›yÒÓ,
MAPKŸ OîÔâ¥ÑROSD~,K⥋
¹ROSq.。tYZ†‡Á,VWpþ?y
犣犿犕犘犓7,¶ ROS ¸ƒy]•[66]。Š‹p,MAPKKK y
DSM1OROSyKâ²_l´µ†‡žyP
|O
[67]。
È#$»ô]Î
,犃狋犕犘犓8O¯
NROS、Ca2+ldey÷ø:Q”„ROSy¢›
ÐLNj¹Q”
[68]。


òóµþô¶·MFN/#$dey
÷ø:ùú
。NorthernÕü62þô,犣犿犕犘犓17
y}ϊл/·LᆇD~

tVWpyþ
?ÙX¨#$„¶·yI»ì
[69]。Berberichm[70]
I/0pMOý¨犣犿犕犘犓4,}wVWpŽñ•
,<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?.ôˆÜu
[71]。
t¶·S€Á
,犗狊犕犃犘犓2y
mRNAÒÓ.ˆTÜu[72]。
3 MAPKOy¦c
MAPKŸ ×Øy¦clc:ïm“H,
MAPKy¦chOTXY8USer"Tyry¾÷
ø:™‹¹

*+

òõö™T¨z‘ijlk$
y÷ø:g

Žñ¹ºP$ MAPK¦cy£¤,
Ï{ghiâ¥ÑM¬3{:oRßì÷øg
(DsPTPases),¤$ MAPKtSer"Tyrùú¾÷
ø:

Vüø÷ø:g
(PTPases),ktTyrùú¾
÷ø:
;`
üø

Wüø÷ø:g
(PPases)[52]。
DsPTPasesPcìùú VXVHCXXGXSRSX
TXXXAY(L/I)M,ÏÐRßy8lPTPaseïÀ
ì

<#

XâY PTPasel DsPTPasesZ¬ï
z{
[73]。


z/=>óô¨um#$p MAPK
yfc\]defgpTyry÷ø:,ϑ=>¬
MAPK*÷ø:"TyrùúÑ÷ø:°±¨óµ,
ïÎKóô¨#$PVüø÷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1tîïy†‡Á
`îï‚fp‚ƒ/þ?
[73]。


@APTPase
yz‘,012犃狋犘犜犘1y}ϊлu憇D~l
¶·yC6
,犃狋犘犜犘1P¾÷ø:£¤,t¢œ¤
$ z · MAPK ¦ c[76]。Gupta m[77] t = >
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