全 文 :书!"#$%&
,2016,36(5):1066-1072
犃犮狋犪犅狅狋.犅狅狉犲犪犾.犗犮犮犻犱犲狀狋.犛犻狀.
!"#$:10004025(2016)0510667 犱狅犻:10.7606/j.issn.10004025.2016.05.1066
%&(
:20160225;)*&%+(:20160412
,-./
:
()*+%,-
(31200209);./01+%,-(2013M531277);234/01+%,-(1201070C);235%6789
:;,-
(11JDG121)
0123
:
<=>
(1982-),?,/0,@0ABC,DEFGH#$IJA$KLMN。Email:lnding6@126.com
4567893:;<$=>?;@ABC
!"#
1,
$%&
2
(1235% AO+%DEP,23Q2212013;2RSTU5% ;$V%P,RS210095)
D E:#$IWX$YKLZ[\].,#$^_`Aabc[defg,hi`#$[jk^b,l#$mA
nWopq
。
rstuvw#$ixyz{|
、
}~|nWj
defB.[
。
_3deYKLZF[DE, ¡¢NPR1、EDS1、MPK4£
¤¥¦§¨©ª«b.[LZ
,
h¬1[DEs®¯°
。
FGH
:
jkpq
;
de±²
;
¥¦
;
de³B
IJKL$
:Q945.8 !MNOP:A
犆狉狅狊狊狋犪犾犽犫犲狋狑犲犲狀犇犲犳犲狀犮犲犘犺狔狋狅犺狅狉犿狅狀犲狊犕犲犱犻犪狋犲犱犛犻犵狀犪犾犻狀犵犘犪狋犺狑犪狔
DINGLina1,YANGGuoxing2
(1ColegeofLifeSciences,JiangsuUniversity,Zhenjiang,Jiangsu212013,China;2ColegeofVeterinaryMedicine,Nanjing
AgriculturalUniversity,Nanjing,210095,China)
犃犫狊狋狉犪犮狋:Intheprocessofplantpathogensinteraction,aseriesofsignaltransductionanddefensesystem
areexcitedinplant,whichtriggersrapidandeffectivedefenseresponses.Afteractivationofdefensere
sponses,wilcausesynthesizingandaccumulatingofaseriesofexogenoussignalingmolecules.Theplant
hormonessalicylicacid(SA),jasmonicacid(JA)andethylene(ET)arerecognizedaskeyplayersinthe
regulationofthesignalingpathwaysinvolved.Inthisreview,therecentresearchadvancesontheinterac
tionamongthethreedifferentsignalingpathwaysaresummarized.Italsopresentedthefunctionofseveral
keyregulatoryproteinssuchasNPR1,EDS1,MPK4,whichareknowntoplayimportantroleinSAand
JA/ETsignalingcrosstalkandcoordinatingtheircomplexrelationships.Inaddition,theprospectoffu
tureworkisdiscussedinthereview.
犓犲狔狑狅狉犱狊:defenseresponse;signalmolecular;regulatoryproteins;signalingpathway
)*´.,#$µ¶·¸¹º»¼½¾W
X$
(
y¿À
、
ÁÀ
、
ÂÀ
、
Wã
)
[ÄÅ
。
#$H
AÆÇ\].ÈÉÊËabcª«[jk
MN
,
ÌÍÎÏ»§¬nWXÀ[ÄÐ
。
#$
IWX$YKLZ[\].
,
#$^_`Aabc
[defg
。
rstÑ[nWdefBÒÓ
Ôz{|
(salicylicacid,SA)、}~|(jasmonic
acid,JA)(ethylene,ET)ÕB[。SA
¶#$^_Ö×ØÙ[a_ÚÛÜ$Ý
,
#$
¼½ÞßWXÀÄÐ1àá5×âãSA。SAI
äånobæçèono[éêëìY
,
Bí
ÞßW]Y¥¦
(
yPR1,PR2PR5)[î
ï
[1]。JAET)*´.ðñÆ,¶6£#$
^_[_ÚAòó$Ý
,
ôõBßöA÷ø
$[ùê
、
úûüýþÿ!"#$ê%£&[A
()Ì
。JAET*¶+,de±²,#$-.
¼½WXÀ/01ÄÅ¡2345×»âã
[2,3]。
56WX$7Ò_AÆ[F8
,¸
!9WX
$±H:^;<=
、
>?;<=@:^;<
=
[4,5]。
:^;<=WX$AÌB:[#$¿C
-..çèDE[;<$ݯAòFG
,
y
¿ÀoHIW
(犚犪犾狊狋狅狀犻犪狊狅犾犪狀犪犮犲犪狉狌犿)!"5ß
J¦KW
(犘狅狑犱犲狉狔犿犻犾犱犲狑)LMW(犇狅狑狀狔
犿犻犾犱犲狑)[íWÀ。Yp,>?;<=WX$B?
[NÒ-..çOAÆFGDE[;<$Ý
,
yÁÀPMWÀ
(犅狅狋狉狔狋犻狊犮犻狀犲狉犲犪)QRWÀ(犃犾
狋犲狉狀犪狉犻犪犫狉犪狊狊犻犮犻犮狅犾犪)。@:^;<=WXÀ,¶S
WX$ÄT#$¡îUH:^;<[A:VW
,
4X1YmA>?;<[A:VW
,
yZ[\MW
À
(犉狌狊犪狉犻狌犿犵狉犪犿犻狀犲犪狉狌犿)、]^WÀ(犕犪犵狀犪
狆狅狉狋犺犲犵狉犻狊犲犪)"_`abÀ(犉狌狊犪狉犻狌犿狅狓狔狊狆狅
狉狌犿)£。¼½· Ü=[WX$ÄÐ/01Ä
Å[¡c
,
#$ámAde[de±²
,
ySA、JA
ET£。fgmA[J×、-ê"¡h· ,
l#$iöõB[j
pqôdeo
[6]。
¨©d
e±²âã1
,
jái:Yq[de³B
,
· [j
pq
,
¬n· [klm
。
ano
,
SApIi:#$¬n:^/@:^;<=WXÀ
[j
pq
;JAETI#$¬n?^;<=WX
À/qr01[j
pqY
[4,7]。
*4)*´
.
,
#$ssE ¡IYt4[ßWXÀY
¬n
,¨
ujávw#$XiõB[j
pq
,
#$
¸!x\óSA、JAET[zy,wzj
Y
,{[îï
,
§j
de³B|ª«[Y
Kb
,
B4i:a}[j
pq
[811]。
{~
,
#$E}[MNq¬,[wÇ
。
j
pqde³B|[¬H#$
¨MN
。
*4
,
#$¶y§¨©ª«[
YKb!".[±²MN¶·¶Î
。
R.
,
tua©pISA、JA
ET¬[¤¥¦,lg¬· de
³B[§ MNzÏ[
。
1 SA、JAETÕB[j
de
[YKLZ
1.1 犛犃Q犑犃<$R:=>;ST0U
ÞßDEîvSAJAõB[de³Bx
Ƭ
。
5å±6SAJAde³
B¶YKn[b
(
1)。¸Ì{Hy~,
SAq[i:l#$z¼½IJAj
pq
Y[klm[ÄÅ
[12]。
R.
,
:^;<=
WXÀ<CÀ
(犘狊犲狌犱狅犿狅狀犪狊狊狔狉犻狀犵犪犲)[Ã
À¡¢`SAÕB[j
pq,£NJAde³B
,
lÄÐ1[-.z¤?^;<=WXÀQ
RWÀ
(犃.犫狉犪狊狊犻犮犻犮狅犾犪)[8]。Koornneef£*Y
¥[&¦
。
§g`Uß;<7YKLZ\].
,
SA¨JAde³B©ª。LmÎßR
[A«=.ଽSAÕB[¬JAq,{
犘犇犉1.2犞犛犘2[£NLZ[13]。RobertSeila
niantz£DE®#îvSA¬JAde³B[
nLZθ̶x\£NJAÕB[¬JAZ¥¦
[¯$U[
[14]。
°¶*a©&¦±HSAJAde³B
|Ƨ LZb
。
01d²x3³´¢
`[R¬<CÀ[ºµpq ¡i:
SAJAj
de³B[15]。¶·.,WX$
Y[±²¸8
(PAMP)õB[j
pqESA
JA [§ [16]。 ¡¹ºÙ»¼[SA
JA(10~100μmol/L)¨ ½¹ºJA z̾¿
JA/ETq,{[îï,îv#$IWX$YK
b.
,
ix[»¼¬ÀÁ[®#¶Î¤
[
[1718]。
R.Þß,{ ¡¼½SA
JA[õB,Â[¬¸ÌÊËÃixÄ
LZ[,Åh
,2ix[-ù¸Ì¢Æ#$³Ç
-[È
,SAJAde³B|[de
ɨgÊË[ª«
[19]。
1.2 犛犃Q犈犜<$R:=>;ST0U
ÌSAJA,SAET[KL#$Iq
r01"WX$[qpq.*`ÍLZ
。
¨y
R¼½qr01ÎH1
(犘犻犲狉犻狊狉犪狆犪犲)Ïq1
ámAET,lèÏq1[-.¬SA[qz2,
{~znÐÑÒÓWÃ
(TCV)[ÄÐ[20]。·+
#$ÔÕ.
,ETde³Bx\SAÕB[¬
JA[£NLZ1ÖõB[j
pq[21]。
CeratoplataninRúû.¸! ¡õBSA
ETÕB[j
pq,4×JAde³B[22]。
Ør.
,ET¢`SA5Ù[¬nØrÚúWÃ
(TMV)[bæçèono.`ͤLZ[23]。
RÜ¥ÛÜpq[WRÝw^1(犺狉犾1).,
j
,{[îï"¬Ã ¿ÀÂÀWXÀ[no
¼½SA JA/ET j
de³B[§
[24]。LeonReyes£DEîv,ET̾¿5ÙSA/
76015W <=>,£:#$jkixÕB[de[¬
1 SA、JAETde³BKL[Þ
Fig.1 ModelforthecrosstalkbetweenSA,JAandETdependentsignalingpathways
NPR1[犘犚1îï,hßET!·5Ù犖犘犚1[F
8SAÕB[¬JAde³BnLZ(
1)[25]。R.,ÃàpIETde³B
³Ç{²EIN3EIL1Â5Ù犛犐犇2[SA
[ùê
,
4áâWX$Y[±²¸8¢`[º
µo
(PTI)j
pq[26]。
1.3 犑犃Q犈犜<$R:=>ST0U
DEîvJAETde³BÄ ¬n?
^;<=WXÀ/qr01[ÄÅ
,
4SAde³
Bã`ͤLZ
[27]。
R.
,JA
ET§ i:j
Y,{ 犘犚3、犘犚4
犘犚12£PR[îï[4,28]。ERF1¶JAETde
³B[ä{²
[29]。
#$-.IJA$[K
åÄAíW\].àEJAET[§ L
Z[&¦
[3031]。
Jæc±çîvR.5è
50%[,{ ¡¼½JAETõB[32]。4ß
}~|·Ü¤Ýw^
(犮狅犻1)·Ü¤Ýw^
(犲犻狀2).,QRWÀ(犃.犫狉犪狊狊犻犮犻犮狅犾犪)ÄзÌõB
犘犇犉1.2,{[îï[28,33]。Laudert£DEîv
JAET¶é~ùê[êi$,°JAET|
h·µ¶YK§ [
,¨
yJAi:[ëÖq,
{¼½ET[£N[34]。ERF1ìJAETde
³B§ ¬nWXÀÄÐ"ET¬JAõBë
ÖqnLZ[íî
(
1)[3536]。
2 SA、JAETde³B
¬[¤¥¦
2.1 犖犘犚1 NPR1¶SAde³B[ä
,{
,
¬õBSA5Ù[PR,{[îïbæ
çèono¶Î¤[
。TGA³Ç{²¶aÜÖ
ïðñoòó|ôõ®ö÷[³Ç{²
(bZIP),{øù#$#$WXÀ:;²úc.[
TGACG/as1/ocsûü4èý,Ij
Y[
PR,{[îï[37]。NPR1I TGA³Ç{²YK
LZi:qSA[PR,{,£NJAde³B
[3839]。NPR1SAIJAde³B[
¬.`ͱ²þ[LZ
(
2)[5,40]。£N
NPR15Ù[SAde³B¸!
6JA5Ù
[¬qr01[no
[41]。NPR1[ÿì¬i:
SAÕB[j
,{îï¶Î¤[[42],°·¶£
NJAq,{îïD!E[。¨ovSA IJA
|[nLZ¸Ì¶x\¿CÝ NPR1
N[
[43]。
2.2 犈犇犛1
EDS1(enhanceddiseasesusceptible1)¶:^
/@:^;<=WXÀ¢`[#$,Åno[ä
{²
,
¬Tol¦Õx1¼^(TIR)Ü=[ÿ"
|®ù#Öòó|ª(NBLRR)¥¦¢`[n
o*¶!E[
[44]。EDS1 IKL{²,ÓÔ
PAD4(phytoalexindecient4)SAG101(senes
cenceassociatedgene101),pIÛÜpq/]
úo¿C?$
(HR/PCD)SA[âã,SAY¸!
x\äp%¾¿EDS1/PAD4/SAG101îï;
EDS1/PAD £ N JA/ET d e ³ B ,{ ~
EDS1/PAD4pISA IJA/ETde³B
8601 ! " # $ % & 36&
[nLZ
(
2)[45]。EDS1(CCNBS
LRR[R¥¦ÕB[j
de³B.de»
ISA)Ì)*[46]。
2.3 犠犚犓犢RVWX
WRKY¥¦¶#$de[ß,{(+³Ç{
²
。
¨Ü³Ç{²{HÖ®ùDNA[ WRKY
®ö÷4Oý
。
f[N,60àó,|.Öï
ð[ WRKYGQKúc,f[C,ÖÜ¥CC
HC/H[-S,ú[47]。WRKY³Ç{²I#$
j
pq"#$üý
。
5ÙSA[j
pq. WRKY ³Ç{²LZ NPR1 " At
MYB44./[4849]。R.\×îï犠犚犓犢70
BíSAq,{[-êoîï,
6¬:^;<
=WXÀ¦KÀ[no
,
°¶£NJAq01,
{犘犇犉1.2[îï,2â¬?^;<=WXÀQR
WÀ[no
[5051];
Yp
,
y#£N 犠犚犓犢70[î
ï
,犘犇犉1.2³Çzy[36,¬QRWÀ[no¾
¿
[49]。
¨©®#ov WRKY7045SA5Ù[j
pq[ä{²
,
!"JA5Ù[j
pq[
Â{²
(
2)。{~,f¨Ãde³B
.±²þ[)Ì
。
*4
,
狑狉犽狔701Ýw
^.
,WRKY70[6:h·Ìvw¬SAJA[
q
,
*·Ì7ª犮狅犻1 Ýw^¬JA[q。¨o
v WRKY70¬SAJAde³B8*
°h·¶!·¸9[
,
SAJA[qF
¸ÌÆfa©I WRKY70)Ì)*[
{²
[52]。
R:A=#¡.犠犚犓犢62¼½ MeJA
SA[§ õB,°¶狀狆狉13Ýw^.õBU
Ë;6
[53]。
§gx\±ç<6çè)ÌÝw^
`U犠犚犓犢62¸!£NJA q,{犔犗犡2
犞犛犘2[îï(2)。¨©®#îvR.
犠犚犓犢62¸ÌpISAÕB[¬JAqj
pq
[£N
。
z][犠犚犓犢13*Ü¥[LZ[54]。Y
p
,
R³Ç{²犠犚犓犢33¬?^;<=W
XÀPMÀ[no¶Î¤[
,犠犚犓犢33[)ÌÝ
w^¸!
6SA [zy,i:SA ÕB[j
p
q
,
£NJAÕB[j
pq[55]。
2.4 犌犚犡480
=>?@{² GRX480*¶aàASA
JAde³BKLb[{²。¥¦p
I¥¦Ý:o[BÇX
。SAõB[ NPR1
i:GRX480,*1ITGA³Ç{²YKLZ,
q SA [ PR , {,£ N q JA [ , {
犘犇犉1.2[îï(2)[56]。Koornneef£&¦
SAJA ¬¼½BÇX[$C
6
[13]。
2.5 犕犘犓4
D±EX:Ç¥¦4(MPK4)¶YaàpISA
JAde³B¬[¤{²。
R. MPK4ÂSAde³B,ä
JAde³B[57]。犿狆犽4Ýw^.,SAzy
6
,SAq[PR,{F-êoîï,¬:^;
<=WXÀ<CGÀHIíWw
(犘狊犲狌犱
狅犿狅狀犪狊狊狔狉犻狀犵犪犲pv狋狅犿犪狋狅)[no¾¿;JAq,
{[îï¼½£N
,
¬?^;<=WXÀQRWÀ
2 SA,JAETde¬[¤¥¦[LZ¸8
Fig.2 ModelformolecularplayersactioninSA,JAandETsignalingcrosstalk
96015W <=>,£:#$jkixÕB[de[¬
[ n o 2 â
[57,58]。EDS1/PAD4 ) Ì h ì
MPK4[./,I MPK4[LZYp,i:SAd
e³B
,
£NJAde³B(2)[58]。R.,
MPK4¶ PAMP õB[j
pq[{²。
MPK4I WRKY33³Ç{²ÿ_!ªù^[é
8Æ
,
ªù^5Ù MPK4[J$ MKS1。¼
½<CÀ/¿ÀKL¥¦nMÄСBí
¬ MPK4[i:"¬ MKS1[N|Ç,X1¢Æ
MKS1 WRKY33B MPK4hOP,i:¬n?
^;<=WXÀ[Q,{[îï
,¨
.jÓÔ
(ùênÀÇù$#ïx[RPAD3[59]。
2.6 犚犛犜1
R犚犈犛犝犚犚犈犆犜犐犗犖1(犚犛犜1),{(
aàìÝS[¥¦
,
TÖ11àUS®ö
÷
。狉狊狋1Ýw^ú².Ý^"ÝVÝ[×à
W¾º
。狉狊狋1Ýw^¬Xo:^;<=ÁÀ¦K
À[¤Wo¾¿
,
¬?^;<=ÁÀPMWÀQ
RWÀ[nWo¾¿
。
I~Y¬q
,狉狊狋1Ýw^.
JAÖ×"犘犇犉1.2[îï×Y36,4SAÖ×"
犘犚1犘犚2[îïׯÙ[60]。{~,犚犛犜1¶a
àZ[pISAIJA/ETÕB[j
pqde
nLZ[,{
(
2)。
3 µ®°
rsDEîvSA、JA、ET5Ù[de³B
#$nWpq.`ͤ[LZ
,
.SA
[±HÒ#$¬WXÀ[jkpq
,
4#$
¬qr01M\]ëmA[nopqÒ¼½
JAET[。^^ß[6îv¨3d
e±²ÕB[nWdefBh·¶½Ë[
,
4
¶Æ_n"§ [YKLZ
,
h!~H,Åé
êaફ[jk^b
。
*4
,
5WXÀno¾
¿I01no¯ÙY[¡c
,
#$yx\`a
Ob[±²MN§¨©ª«[YKb
,
!$U
¬W1Öjkpq[cwEzdT[D
E
。
efrs¬#$nWpq[defB[DEO
èabc
,
°de§ /n[YK
LZ[åì"MNg<9hi[$C6
。
¨yr
sjjѦq{²ERF1ìJAETd
e³B§ ¬n?^=WXÀÄÐ[íî
,
°ô^[±²MNg<9
。
an±H
SA¬JAÆnLZ,°¶¬JA¬SAde
fB[£NLZrshkÎßDE&l
。
m
,
g¬WX$}q$yx\noixùê
de³B4l#$íW*·¶Î
。
#$nWopqMNp±ª«ßu
,
#$IW
X$[KL
、
nWde[³B"j
pq[`A].Æ_abc[ó{²,{
,
fgHß
j
de[pq
Î5[q
,
B4r#$©ªi:saàd[
,
$
U¬¬,[Àtjk
。
,
_·9D
Emäí uvdeYK[¥¦
,
°
¶h×D{²[DE®#àaí
,¨
y
MPK4"RST1,fgixKL.[¤LZg
Æwx
,
ô^)Ì!"ixdex.[L
Zyaz[DE
。
¥¦¶¯j
de
¬[,Å
,
*¶ó#${^nWo[
¤{²
。
uvZ[¨u[|bî9¶1#$n
WdeDE[}î|a
。
X_±²A$%[`
,
¸8#$R.t±~½¾Ü=[
nWÝw^
,
h,{
。
Z#$,{]
¾³,{nW#¡YtÊêhqZ½TUAm
.
,
·°HTL$
[vþaZ[
,
¡9¬ï,A«ya
Z
,
ôdù[DEÞ&[qZs®
。
YZ!M
:
[1] GRANT M,LAMBC.Systemicimmunity[J].犆狌狉狉犗狆犻狀
犘犾犪狀狋犅犻狅犾.,2006,9(4):414420.
[2] LOUISJ,BASUS,VARSANIS,犲狋犪犾.Ethylenecontributes
tomaizeinsectresistance1mediatedmaizedefenseagainstthe
phloemsapsuckingcornleafaphid[J].犘犾犪狀狋犘犺狔狊犻狅犾.,
2015,169(1):313324.
[3] HAELCONRADV,ABOUMANSOURE,DíAZRICCIJC,
犲狋犪犾.ThenovelelicitorAsEStriggersadefenseresponsea
gainst犅狅狋狉狔狋犻狊犮犻狀犲狉犲犪犻狀犃狉犪犫犻犱狅狆狊犻狊狋犺犪犾犻犪狀犪[J].犘犾犪狀狋
犛犮犻.,241:120127.
[4] GLAZEBROOKJ.Contrastingmechanismsofdefenseagainst
biotrophicandnecrotrophicpathogens[J].犃狀狀狌.犚犲狏.犘犺狔
狋狅狆犪狋犺狅犾.,2005,43:205227.
[5] SPOELSH,JOHNSONJS,DONGX.Regulationoftradeoffs
betweenplantdefensesagainstpathogenswithdifferentlifes
tyles[J].犘狉狅犮.犖犪狋犾.犃犮犪犱.犛犮犻.犝犛犃,2007,104(47):18
84218847.
[6] DEVOSM,VANOOSTENVR,VANPOECECRM,犲狋犪犾.
0701 ! " # $ % & 36&
Signalsignatureandtranscriptomechangesof犃狉犪犫犻犱狅狆狊犻狊
duringpathogenandinsectattack[J].犕狅犾.犘犾犪狀狋犕犻犮狉狅犫犲.
犐狀狋犲狉犪犮狋,2005,18(9):923937.
[7] THALERJS,HUMPHREYPT,WHITEMANNK.Evolu
tionofjasmonateandsalicylatesignalcrosstalk[J].犜狉犲狀犱狊
犘犾犪狀狋犛犮犻.,2012,17(5):260270.
[8] BARIR,JONESJD.Roleofplanthormonesinplantdefence
responses[J].犘犾犪狀狋犕狅犾.犅犻狅犾.,2009,69(4):473488.
[9] GRANT MR,JONESJD.Hormone(dis)harmonymoulds
planthealthanddisease[J].犛犮犻犲狀犮犲,2009,324(5928):750
752.
[10] DINGL,XUH,YIH,犲狋犪犾.Resistancetohemibiotrophic
F.graminearuminfectionisassociatedwithcoordinatedand
orderedexpressionofdiversedefensesignalingpathways[J].
犘犔狅犛犗狀犲,2011,6(4):e19008.
[11] YANGYX,AHAMMEDGJ,WUC,犲狋犪犾.Crosstalka
mongjasmonate,salicylateandethylenesignalingpathways
inplantdiseaseandimmuneresponses[J].犆狌狉狉犲狀狋犘狉狅狋犲犻狀
犪狀犱犘犲狆狋犻犱犲犛犮犻犲狀犮犲,2015,16(5):450461.
[12] NAIDOOR,FERREIRAL,BERGERDK,犲狋犪犾.Theiden
tificationanddifferentialexpressionof犈狌犮犪犾狔狆狋狌狊犵狉犪狀犱犻狊
pathogenesisrelatedgenesinresponsetosalicylicacidand
methyljasmonate[J].犉狉狅狀狋犘犾犪狀狋犛犮犻.,2013,4:43.
[13] KOORNNEEFA,LEONREYESA,RITSEMAT,犲狋犪犾.
Kineticsofsalicylatemediatedsuppressionofjasmonatesig
nalingrevealaroleforredoxmodulation[J].犘犾犪狀狋犘犺狔狊犻
狅犾.,2008,147(3):13581368.
[14] ROBERTSSILANIANTZA,NAVARROL,BARIR,犲狋
犪l.Pathologicalhormoneimbalances[J].犆狌狉狉.犗狆犻狀.犘犾犪狀狋
犅犻狅犾.,2007,10(4):372379.
[15] SONG GC,CHOIHK,RYU CM.Gaseous3pentanol
primesplantimmunityagainstabacterialspeckpathogen,
犘狊犲狌犱狅犿狅狀犪狊狊狔狉犻狀犵犪犲pv.狋狅犿犪狋狅viasalicylicacidandjas
monicaciddependentsignalingpathwaysin犃狉犪犫犻犱狅狆狊犻狊[J].
犉狉狅狀狋犘犾犪狀狋犛犮犻.,2015,6:821.
[16] HALIMVA,ALTMANNS,ELLINGERD,犲狋犪犾.PAMP
induceddefenseresponsesinpotatorequirebothsalicylicacid
andjasmonicacid[J].犘犾犪狀狋犑.,2009,57(2):230242.
[17] MURLAJ,KENTONP,ATZORNR,犲狋犪犾.Theoutcomes
ofconcentrationspecificinteractionsbetweensalicylateand
jasmonatesignalingincludesynergy,antagonism,andoxida
tivestressleadingtoceldeath[J].犘犾犪狀狋犘犺狔狊犻狅犾.,2006,
140(1):249262.
[18] DINGLN,YANGGX,YANGRY,犲狋犪犾.Investigatingin
teractionsofsalicylicacidandjasmonicacidsignalingpath
waysinmonocotswheat[J].犘犺狔狊犻狅犾.犕狅犾.犘犾犪狀狋犘,2016,
doi:10.1016/j.pmpp.2016.01.002.
[19] PROIETTIS,BERTINIL,TIMPERIOAM,犲狋犪犾.Crosstalk
betweensalicylicacidandjasmonateinArabidopsisinvestiga
tedbyanintegratedproteomicandtranscriptomicapproach
[J].犕狅犾.犅犻狅狊狔狊狋.,2013,9(6):11691187.
[20] DEVOSM,VANW,KOORNNEEFA,犲狋犪犾.Herbivorein
ducedresistanceagainstmicrobialpathogensinArabidopsis
[J].犘犾犪狀狋犘犺狔狊犻狅犾.,2006,142(1):352363.
[21] PAUDELJR,BEDEJC.Ethylenesignalingmodulatesherbi
voreinduceddefenseresponsesinthemodellegume犕犲犱犻犮犪
犵狅狋狉狌狀犮犪狋狌犾犪[J].犕狅犾.犘犾犪狀狋犕犻犮狉狅犫犲犐狀狋犲狉犪犮狋,2015,28
(5):569579.
[22] BACCELLII,LOMBARDIL,LUTIS,犲狋犪犾.Ceratopla
tanininducesresistanceinArabidopsisleavesthroughstoma
talperception,overexpressionofsalicylicacidandethylene
signalinggenesandcamalexinbiosynthesis[J].犘犔狅犛犗狀犲,
2014,9(6):e100959.
[23] VERBERNEMC,HOEKSTRAJ,BOLJF,犲狋犪犾.Signaling
ofsystemicacquiredresistanceintobaccodependsonethylene
perception[J].犘犾犪狀狋犑.,2003,35(1):2732.
[24] DEVADASSK,ENYEDIA,RAINAR.The犃狉犪犫犻犱狅狆狊犻狊
犺狉犾1 mutationrevealsnoveloverlappingrolesforsalicylic
acid,jasmonicacidandethylenesignalinginceldeathand
defenceagainstpathogens[J].犘犾犪狀狋犑.,2002,30(4):
467480.
[25] LEONREYESA,SPOELSH,DELANGEES,犲狋犪犾.Eth
ylenemodulatestheroleofNONEXPRESSOROFPATHO
GENESISRELATEDGENES1incrosstalkbetweensalicy
lateandjasmonatesignaling[J].犘犾犪狀狋犘犺狔狊犻狅犾,2009,149
(4):17971809.
[26] CHENH,XUEL,CHINTAMANANIS,犲狋犪犾.ETHYL
ENEINSENSITIVE3and ETHYLENE INSENSITIVE3
LIKE1 repress SALICYLIC ACID INDUCTION DEFI
CIENT2expressiontonegativelyregulateplantinnateimmu
nityin 犃狉犪犫犻犱狅狆狊犻狊[J].犘犾犪狀狋犆犲犾犾,2009,21(8):2
5272540.
[27] ALONM,MALKAO,EAKTEIMANG,犲狋犪犾.Activationof
thephenylpropanoidpathwayin犖犻犮狅狋犻犪狀犪狋犪犫犪犮狌犿improves
theperformanceofthewhitefly犅犲犿犻狊犻犪狋犪犫犪犮犻viareduced
jasmonatesignaling[J].犘犔狅犛犗狀犲,2013,8(10):e76619.
[28] PENNINCKXIA,THOMMABP,BUCHALA A,犲狋犪犾.
Concomitantactivationofjasmonateandethyleneresponse
pathwaysisrequiredforinductionofaplantdefensingenein
Arabidopsis[J].犘犾犪狀狋犆犲犾犾,1998,10(12):21032113.
[29] SHERIFS,EISHARKAWYI,PALIYATHG,犲狋犪犾.Dif
ferentialexpressionofpeachERFtranscriptionalactivatorsin
responsetosignalingmoleculesandinoculationwith犡犪狀
狋犺狅犿狅狀犪狊犮犪犿狆犲狊狋狉犻狊 pv.狆狉狌狀犻[J].犑.犘犾犪狀狋犘犺狔狊犻狅犾.,
2012,169(7):731739.
[30] VANWEESSC,VANDERENTS,PIETERSECM.Plant
immuneresponsestriggeredbybeneficialmicrobes[J].犆狌狉狉
犗狆犻狀犘犾犪狀狋犅犻狅犾.,2008,11(4):443448.
[31] PLETTJM,KHACHANEA,OUASSOUM,etal.Ethyl
eneandjasmonicacidactasnegativemodulatorsduringmu
tualisticsymbiosisbetween犔犪犮犮犪狉犻犪犫犻犮狅犾狅狉 and犘狅狆狌犾狌狊
roots[J].犖犲狑犘犺狔狋狅犾,2014,202(1):270286.
[32] SCHENKPM,KAZANK,WILSONI,犲狋犪犾.Coordinated
plantdefenseresponsesin犃狉犪犫犻犱狅狆狊犻狊revealedbymicroarray
17015W <=>,£:#$jkixÕB[de[¬
analysis[J].犘狉狅犮.犖犪狋犾.犃犮犪犱.犛犮犻.犝犛犃,2000,97(21):
1165511660.
[33] THOMMABPHJ,PENNINCKXIAMA,BROEKAERTWF,
犲狋犪犾.Thecomplexityofdiseasesignalingin犃狉犪犫犻犱狅狆狊犻狊[J].
犆狌狉狉.犗狆犻狀.犐犿犿狌狀狅.,2001,13(1):6368.
[34] LAUDERTD,WEILEREW.Aleneoxidesynthase:ama
jorcontrolpointin犃狉犪犫犻犱狅狆狊犻狊狋犺犪犾犻犪狀犪octadecanoidsigna
ling[J].犘犾犪狀狋犑.,1998,15(5):675684.
[35] LORENZOO,CHICOJM,SANCHEZSERRANOJJ,犲狋
犪犾.JASMONATEINSENSITIVE1encodesa MYCtran
scriptionfactoressentialtodiscriminatebetweendifferentjas
monateregulateddefenseresponsesinArabidopsis[J].犘犾犪狀狋
犆犲犾犾,2004,16(7):19381950.
[36] LORENZOO,PIQUERASR,SANCHEZSERRANOJJ,
犲狋犪犾.犈犜犎犢犔犈犖犈犚犈犛犘犗犖犛犈犉犃犆犜犗犚1integratessig
nalsfromethyleneandjasmonatepathwaysinplantdefense
[J].犘犾犪狀狋犆犲犾犾,2003,15(1):165178.
[37] THUROWC,SCHIERMEYERA,KRAWEZKYS,犲狋犪犾.
TobaccobZIPtranscriptionfactorTGA2.2andrelatedfactor
TGA2.1havedistinctrolesinplantdefenseresponsesand
plantdevelopment[J].犘犾犪狀狋犑.,2005,44(1):100113.
[38] DONGX.NPR1,althingsconsidered[J].犆狌狉狉犗狆犻狀犘犾犪狀狋
犅犻狅犾.,2004,7(5):547552.
[39] TADAY,SPOELSH,PAJEROWSKAMUKHTARK,犲狋
犪犾.PlantimmunityrequiresconformationalchargesofNPR1
viaSnitrosylationandthioredoxins[J].犛犮犻犲狀犮犲,2008,321
(5891):952956.
[40] CAARLSL,PIETERSECM,VANWEESSC.Howsalicyl
icacidtakestranscriptionalcontroloverjasmonicacidsigna
ling[J].犉狉狅狀狋犘犾犪狀狋犛犮犻.,2015,6:170.
[41] STOTZHU,KOCHT,BIEDERMANNA,犲狋犪犾.Evidence
forregulationofresistanceinArabidopsistoEgyptiancotton
wormbysalicylicandjasmonicacidsignalingpathways[J].
犘犾犪狀狋犪,2002,214(4):648652.
[42] KINKEMA M,FAN W,DONGX.Nuclearlocalizationof
NPR1isrequiredforactivationof犘犚geneexpression[J].
犘犾犪狀狋犆犲犾犾,2000,12(12):23392350.
[43] SPOELSH,KOORNNEEFA,CLAESSENSSM,犲狋犪犾.
NPR1 modulatescrosstalk between salicylateandjas
monatedependentdefensepathwaysthroughanovelfunction
inthecytosol[J].犘犾犪狀狋犆犲犾犾,2003,15(3):760770.
[44] HUG,DEHARTAKA,LIY,犲狋犪犾.犈犇犛1intomatoisre
quiredforresistancemediatedbyTIRclassRgenesandtherecep
torlikeRgene犞犲[J].犘犾犪狀狋犑.,2005,42(3):376391.
[45] WIERMER M,FEYSBJ,PARKERJE.Plantimmunity:
theEDS1regulatorynode[J].犆狌狉狉.犗狆犻狀.犘犾犪狀狋犅犻狅犾.,
2005,8(4):383389.
[46] VENUGOPALSC,JEONGRD,MANDALMK,犲狋犪犾.En
hanceddiseasesusceptibility1andsalicylicacidactredun
dantlytoregulateresistancegenemediatedsignaling[J].
犘犔狅犛犌犲狀犲狋,2009,5(7):e1000545.
[47] EULGEM T,RUSHTONPJ,ROBATZEKS,犲狋犪犾.The
WRKYsuperfamilyofplanttranscriptionfactors[J].犜狉犲狀犱狊
犘犾犪狀狋犛犮犻.,5(5):199206.
[48] EULGEM T,SOMSSICHIE.NetworksofWRKYtran
scriptionfactorsindefencesignaling[J].犆狌狉狉.犗狆犻狀.犘犾犪狀狋
犅犻狅犾.,2007,10(4):366371.
[49] SHIMJS,CHOIYD.DirectregulationofWRKY70byAt
MYB44inplantdefenseresponses[J].犘犾犪狀狋犛犻犵狀犪犾犅犲犺犪狏,
2013,8(6):e20783.
[50] LIJ,BRADERG,KARIOLAT,犲狋犪犾.WRKY70modulates
theselectionofsignalingpathwaysinplantdefense[J].犘犾犪狀狋
犑.,2006,46(3):477491.
[51] SHIMJS,JUNGC,LEES,犲狋犪犾.AtMYB44regulates
WRKY70expressionandmodulatesantagonisticinteraction
betweensalicylicacidandjasmonicacidsignaling[J].犘犾犪狀狋
犑.,2013,73(3):483495.
[52] RENCM,ZHU Q,GAOBD,犲狋犪犾.Transcriptionfactor
WRKY70displaysimportantbutnoindispensablerolesinjas
monateandsalicylicacidsignaling[J].犑.犐狀狋犲犵狉犘犾犪狀狋犅犻
狅犾.,2008,50(5):630637.
[53] MAOP,DUAN M,WEIC,犲狋犪犾.WRKY62transcription
factoractsdownstreamofcytosolicNPR1andnegativelyreg
ulatesjasmonateresponsivegeneexpression[J].犘犾犪狀狋犆犲犾犾
犘犺狔狊犻狅犾.,2007,48(6):833842.
[54] QIUD,XIAOJ,DINGX,犲狋犪犾.犗狊犠犚犓犢13mediatesrice
diseaseresistancebyregulatingdefenserelatedgenesinsalic
ylateandjasmonatedependentsignaling[J].犕狅犾.犘犾犪狀狋犕犻
犮狉狅犫犲犐狀狋犲狉犪犮狋,2007,20(5):492499.
[55] BIRKENBIHLRP,DIEZELC,SOMSSICHIE.犃狉犪犫犻犱狅狆
狊犻狊WRKY33isakeytranscriptionalregulatorofhormonal
andmetabolicresponsestoward犅狅狋狉狔狋犻狊犮犻狀犲狉犲犪infection[J].
犘犾犪狀狋犘犺狔狊犻狅犾.,2012,159(1):266285.
[56] NDAMUKONGI,ABDALLATAA,THUROWC,犲狋犪犾.
SAinducible犃狉犪犫犻犱狅狆狊犻狊glutaredoxininteractswithTGA
factorsandsuppressesJAresponsivePDF1.2transcription
[J].犘犾犪狀狋犑.,2007,50(1):128139.
[57] PETERSENM,BRODERSENP,NAESTEDH,犲狋犪犾.犃狉
犪犫犻犱狅狆狊犻狊mapkinase4negativelyregulatessystemicacquired
resistance[J].犆犲犾犾,2000,103(7):11111120.
[58] BRODERSENP,PETERSEN M,BJORNNIELSENH,犲狋
犪犾.犃狉犪犫犻犱狅狆狊犻狊MAPkinase4regulatessalicylicacidand
jasmonicacid/ethylenedependentresponsesviaEDS1and
PAD4[J].犘犾犪狀狋犑.,2006,47:532546.
[59] QIUJL,FIILBK,PETERSENK,犲狋犪犾.犃狉犪犫犻犱狅狆狊犻狊MAP
kinase4regulatesgeneexpressionthroughtranscriptionfac
torreleaseinthenucleus[J].犈犕犅犗犑.,2008,27(16):2
2142221.
[60] MANGHG,LALUKKA,PARSONSEP,犲狋犪犾.The犃狉犪犫犻
犱狅狆狊犻狊RESURRECTION1generegulatesanovelantagonistic
interactioninplantdefensetobiotrophsandnecrotrophs[J].
犘犾犪狀狋犘犺狔狊犻狅犾.,2009,151(1):290305.
(
!"
:
#$%
)
2701 ! " # $ % & 36&