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Research Progress of a MITEs mPing in Rice (Oryza sativa L.)

水稻MITEs类转座子mPing研究进展



全 文 :书!"#$%&
,2016,36(7):1493-1498
犃犮狋犪犅狅狋.犅狅狉犲犪犾.犗犮犮犻犱犲狀狋.犛犻狀.
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./*
(1983-),0,12,34,56789:;<=>?@%AB。Email:xiaoranwang@hotmail.com
CDEF:GHI,12,JK,5678#$;45 犕犐犜犈狊6789犿犘犻狀犵:;<=
!"#
1,2,
$ %1,& 1,()*1
(1OPQRS% MT+%%U,VW450002;2OPQRS% OPX9:M$%YZ[\],VW450002)
> ?:犕犻狀犻犪狋狌狉犲犘犻狀犵(犿犘犻狀犵)^ _`abYcde< (MiniatureInvertedRepeatTransposableElements,
MITEs)fde<犜狅狌狉犻狊狋likeg(hY6ij,^ 9:,klmnopqrstuvq MITEs,^ MITEsS(h
wxyz{|}~€)*‚ƒ„…deu†qij‡s


,犿犘犻狀犵^de<‰Š‹ŒABqŽ。
‘’“”•–—˜™šm›œŠ犿犘犻狀犵qž、deŸ ¡、¢u£†¤¥,klq¦§¨©ªqAB«
¬

­«s®¯°±B MITEsqde²³¤犿犘犻狀犵de@AB

de<
;犿犘犻狀犵;犘犻狀犵;犘狅狀犵;9:
CDE6$
:Q789 !FGHI:A
犚犲狊犲犪狉犮犺犘狉狅犵狉犲狊狊狅犳犪犕犐犜犈狊犿犘犻狀犵犻狀犚犻犮犲(犗狉狔狕犪狊犪狋犻狏犪犔.)
WANGXiaoran1,2,YANGLu1,LIZhen1,CHENXinjian1
(1ColegeofLifeSciences,HenanAgriculturalUniversity,Zhengzhou450002,China;2HenanKeyLaboratoryofRiceBiology,
HenanAgriculturalUniversity,Zhengzhou450002,China)
犃犫狊狋狉犪犮狋:犕犻狀犻犪狋狌狉犲犘犻狀犵 (犿犘犻狀犵)isaprominentmemberin犜狅狌狉犻狊狋likesuperfamilyofMiniatureInvert
edRepeatTransposableElements(MITEs).ItisthefirstactiveMITEsidentifiedinthericegenomeand
oneofthefew MITEswhichkeeplowcopynumbersandmaintainmobilizationundernaturalcondition.
Therefore,itisanirreplaceablematerialforthestudyrelatedtotransposon.Thispapercomprehensively
demonstratesresearchprogressof犿犘犻狀犵throughanalyzingitsstructure,transposasedonor,activation
characteristics,anditsinfluenceongenome,lookingforwardtolayfoundationonadvancedresearchof
transpositionmechanismofMITEsandthefurtherdevelopmentandutilizationof犿犘犻狀犵.
犓犲狔狑狅狉犱狊:transposon;犿犘犻狀犵;犘犻狀犵;犘狅狀犵;rice
  de<(transposon)»¼½­dek<(trans
posonelements),^ ,klw~)5c³¾¿Àq
sÁÂÃÄÁ

€ÅÂM$,klw;ÆÇÈ
,^
Å
ÂM$,klqY6liÉ;
。1951 ™ Mc
Clintock€ÊËwrs̵͗de<,€‰ÎÏ
qÐÑm¼+%ÒÓÔÕ

Ö×Ø+%ABqÙÚ
µ¬

ÛÜÝÞpde<[ß^àáM$«NqY
6âã‡s

€¥ä¨M$,klq;åwµÍ

Ûf,klwdeæçqèéê45%,#$,kl
Cëß­50%~80%Ù¨[12],ìí€É;îï+
#$wde<ðñ,klqèé~äê90%[3]。
òóde
Öø
ˆõµq,kžùú

=êûüùNìíýþ¡
žqYÿ!^M$,kl«NqY6àáâ
‡s
[45]。
de<"#de²³~¼;­2$f`:Class
Ⅰ(ade<)¾ClassⅡ(de<)[6]。ClassⅠ¾Ⅱ
fde<%#&)^()œ*+deŸôâ,~
;­)5`de<
(autonomouselements)¾-)
5`de<
(nonautonomouselements)[7]。Class
Ⅰde<ø,klDNAd.­RNA,/01ad
.ŸRNaseH ad.­DNA,234i,ke¿
qdÀ

56fde<½­ade<
(retrotrans
posons);ClassⅡde<78C9DNA:;/<°
²³š[Íde

¼½‡­de<
(transponsons)。
ClassⅠdec³/<°qde9?,k
ˆ€,klm{|y@A

*B

SÉ;ClassⅡf
de
<°de£ZCëC…‰¥@x
q{|y
[8],
&wSÉ;ClassⅡf-)5`de
<

&{|yìíÙDg950t[9]。
MITEs(miniatureinvertedrepeatstransposa
bleelements)½­_`abYcde<,EFG`
DNAde<(ClassⅡ),^ #$wY6qsfde
<
[10],
ÖL&HClassⅡde<ÙI,MITEs~
C9)Jc³K){|yêpyLMíNO
t
[11]。MITEsµÍFîï+E$Êˇw,3š
qABwÛ܉P€&HsQ´R#$

á$Mí
Ûf , k l m µ Í MITEs q S T[12]。ø F
MITEsSAø)5`qde<«NBš,ÏU_F
600bp,VK&)WX*+deŸ(Y1),de¢
uZô%[&HdeŸ ¡

¥9:,klq;å
w\*µÍ
,MITEs(hð9:,kl8.6%,]
{|y^g_100000t‡A[13]。%# MITEs`
aabYcbc
(terminalinvertedrepeats,TIRs)
¾d¿ZYcbc
(targetsiteduplications,TSDs)
qž£Z

5yãeSq MITEs;i犜狅狌狉犻狊狋
likeMITEs¾犛狋犪狑犪狔likeMITEs2tg(h,f
FTSDsCë­3g,qTAAhTTA¿Z,B3
FTSDsß^A­2g,TA[14],6$iúqjk
~ôøF2t(hqdeŸ ¡犘犐犉/犎犪狉犫犻狀犵犲狉
¾犜犮1/犿犪狉犻狀犲狉deŸ(h¥de<<°¿Zq
klœs÷mb†ÓV
[1516]。
犿犘犻狀犵(犿犻狀犻犪狋狌狉犲犘犻狀犵)2n&oF2003™,
^9:wnopqrst¢uq MITEs[13,1718],^
^9: MITEseS(hwps~€)*qçƒ
C…¢u‚qde<
[19]。
ÙIF&H MITEs,
Cë犿犘犻狀犵„…‰¥@zq{|y,r:sït,
kluvw70t犿犘犻狀犵{|,x:‘9311’yuv
w14t4z{|[13]。{Fde¢u}C…‰¥@
xq{|yq£ZK犿犘犻狀犵i­—deŽ

˜™š

|×¥&ABqÙÚ¯°
,犿犘
犻狀犵¤&H MITEsÙy€«N‹ŒABw}pY
Õ

~€ù¡žI¤;<€¨‹Œ‚p—ÇÈ
ƒ·
[2022]。
1 犿犘犻狀犵qž£„
犿犘犻狀犵kL)5`de<犘犻狀犵`aäUI…
B‚†

ÏU430bp,` auv15tg,qTIRs,
TSDsbcCë­C‡q3g, TAAh TTA,%
#TIRs¾TSDsbc£Z5&ˆf­犜狅狌狉犻狊狋like
g(h
[13,19,23]。犘犻狀犵ÏU­5314bp,‰qŠ`a
;‹œ253bp¾177bpbcL犿犘犻狀犵É;bcs
V†êp99.9%¾100%,ŠÁŒ8l”i4zq
犿犘犻狀犵bc(Y2)[13]。IÐ,€¥9:,kly#
qnŽwµÍ
,犿犘犻狀犵Lst)5`de<
犘狅狀犵q`abc70%sV,犘狅狀犵ÏU5166bp,
L犘犻狀犵€13bp‰IbcqTIRs(Y2)[24]。L
&H MITEs(hij‰I,犿犘犻狀犵Ù)œ*+d
eŸqôâ



L‡žäUI…})5*+d
eŸq犘犻狀犵¾犘狅狀犵‘i­犿犘犻狀犵~ôqdeŸ
 ¡

’þ“”=•de<–—d¿ZYcbc
(TSDs),˜˜™þ“”=•de<`aabYcbc(TIRs)
Y1 DNAde<(ClassⅡ)qž
TheblacktrianglesflankingeachelementrepresentTSDs,andthegraytrianglesrepresentTIRs
Fig.1 ThestructureofDNAtransposons(ClassⅡ)
4941 ! " # $ % &                   36š
犿犘犻狀犵ø犘犻狀犵q2t`aŒ8Bi,}L犘狅狀犵ŠaäUI…。’þ›œ^de<–—d¿ZYcbc(TSDs),
™þ“”=`aYcbc
(TIRs)
Y2 犿犘犻狀犵、犘犻狀犵¾犘狅狀犵qž
犿犘 犻狀犵issplicedbybothendsof犘犻狀犵,whichhasthehighhomologywiththeendsof犘狅狀犵;TSDsaretheblackarrowheads
flankingeachelement,andthegraytrianglesrepresentTIRs
Fig.2 Thestructureof犿犘犻狀犵,犘犻狀犵and犘狅狀犵
2 犿犘犻狀犵 ~ôqdeŸ ¡犘犻狀犵
¾犘狅狀犵
  ˜™š,E­犿犘犻狀犵ž€deŸ ¡q犘犻狀犵
¾犘狅狀犵‚p—¯°qAB。ž;åŸ[,犘犻狀犵
¾犘狅狀犵mÉ uv2t ORF(ORF1¾ ORF2),
ø¡FORF1*+q¢£¤,Ãbc‰¥†êp
60%,BORF2*+¢£¤,Ãbc‰¥†^äê
84%[13,24]。¦§­¨,ORF1q©ôªÙ4ñ«¬,
Ök&*+­$L犿狔犫d.kŒ
(DNAbindingdomain)€s÷‰¥†,#ˆà
oORF1*+­$~ô®LDNAL¢£”,¯
1de<¢u
[2526]。ORF2*+­$wuv DDE
,b
,DDE,b°^É;deŸ±Nu†w²qY
6£„
[27],
}ORF2uvN2、N3¾C1äUC‡³
Œ

*+­$L犘犐犉/犎犪狉犫犻狀犵犲狉deŸ(h€´
䉥†
[28],
¥µ¶•ORF2*+­$µ·deŸ
©ô

Ö«s®ABŸ[
,ORF1¾ORF2€犿犘犻狀犵
de9?w!µ·íŠY6qE·

Ö¸);¹ª
Ù4ñ«¬
[24]。
¥F犘犻狀犵h犘狅狀犵ºst^犿犘犻狀犵»:de
Ÿ ¡í§ªW÷¼

r:

sït

,kl~n
ŽpS½5t犘狅狀犵¾1t犘犻狀犵q{|,*B€x
:
‘9311’,klm¾¿œµÍ4z犘犻狀犵qÀT。

,Zhang¨[16]¶·ÁµÃ%ÄÁABŸ[,
Sã#$

á$¾ÅÅ,klw!€犘狅狀犵ž
I…bc
(犘狅狀犵like),SÉ;犘狅狀犵like!uv2
tORF,}*+­$ƵL犘狅狀犵q2tORFÇÈ

[15,25,29]。Wang¨[30]54z 犘犻狀犵、犘狅狀犵 ¤
犘犻狀犵qORF³;‹L犿犘犻狀犵ÊdNËPÌ3µ
Í

Zœ4z犘犻狀犵qžWX¢u犿犘犻狀犵 de,
B›ŠF ~õµ犿犘犻狀犵de}deÍΉ˜。
ÏF€¥è—lÐÑÒ39:ÙIÓÔ3~µÍ

犿犘犻狀犵q¢uÕÎL犘狅狀犵 q=ê9։Š,Ö×
صÍL犘犻狀犵=ꐀŠÙ。“”;å3ào,
òó犘犻狀犵*+­$)Ú¢u犿犘犻狀犵 qôâ,Ö~
ôøFfaÛá<³Œqù

ÜV犘犻狀犵*+³
ŒÙô°ë=ê

N–Ÿ#¥µ!¶•犘狅狀犵°^
犿犘犻狀犵qdeŸ ¡。
*B

˜Ý9:EG4ÞÁwqAB,߂—Ù
Iq¼

ABŸ[
,EG4Áw犿犘犻狀犵qdeu
†€)*Ñҁ‚ƒ~‚pC…

×àSw
40t{|qáUÙÚâã,¦§­¨EG4,kl
m犿犘犻狀犵 q{|yäåâííx1000t[31]。
Teramoto¨[19]€EG4Kæ3~5d3¥ÙIÓÔ
;åµÍ

ÎçqµM«°pèa;Mlоé"
³Œ;NêÁÐ

éçmnop—犿犘犻狀犵qëȾ


IÐ|×犘犻狀犵=ê9ÖqìíNî,ˆÍï
€&Hõµ犿犘犻狀犵¢uqqçwªØµÍ9,AB
ÛjÝ­€çµMqnÝ~ôõµ犘犻狀犵q£ú†


ä=êq犘犻狀犵¥F犿犘犻狀犵 q¢uµ·Y6


“N~ð
,犘狅狀犵~ô^犿犘犻狀犵 ²qde
Ÿ ¡

Bsde<犘犻狀犵~ô€#$µÃ£÷
ÐÝ~)Ú¢u犿犘犻狀犵deqôâ,ÖªWŸ#Ÿ
ñ^(犘犻狀犵¾犘狅狀犵€ÙIÐÝ[Í¥犿犘犻狀犵q
òóîô

3 õµ犿犘犻狀犵de¢uqk
犿犘犻狀犵qde¢u2nµÍF9:iöç
¤R÷qlÐÑÒ9?w
[13,1718],
ˆ3Û܉Pµ
Íuøäù

¢úûü

ýþÿ!¤"#¾$%¨
ÄÁ ~¢u犿犘犻狀犵de[3238],BN–qçCë
õµ,klÇÈq?@¾=>?@ùú

€ÏFq
¹Ew

I&Ùyynop—犿犘犻狀犵qde¢u,
Iе͗,klqSãùú
[39]。
犿犘犻狀犵¢uqÙIqçƒ,E­deŸ ¡
59417Ý              ./*,¨ :9: MITEsfde<犿犘犻狀犵AB«¬
犘狅狀犵hF犘犻狀犵q=êùN^犿犘犻狀犵 µM¢uq
€‡s


,犿犘犻狀犵mɾ5’` a–—bcq
()*+,N9ÖL&ö÷†,Íì퉊

ä-
Uq+,N¥F„… 犿犘犻狀犵 qö÷µ·Y6E
·
[34,40]。
*B
,Tsukiyama¨[9]ABµÍ,9:È
‰Šî.,k
(犚狌狉犿1)ù~õµ犿犘犻狀犵q
de

œ/q^

ˆÐ犿犘犻狀犵mɾ–—bcq(
)*+,N9Ö×صMùú
。Yang¨[40]µÍ¶
·01úžŸⅡ2³3(%456)ÿ!9:~7
µ犿犘犻狀犵de¢u,^ øF%456õµ9:,k
lÇÈq?@¾=>?@ùú
。Wu¨[36]¶·9
:

8988’E­}¡,:(犣犻狕犪狀犻犪犾犪狋犻犳狅犾犻犪)E
­ ¡«;"#$%

"#Á,klwòó¿œn
op ¡,klD<

־߂—†ö÷q"#

,^
€"#Áwnop犿犘犻狀犵qde¢u,k
ˆ"#9?w~ôøFÉ;D=;

õµ}¡>(

,kl?á
”(genomeshock),
7Bõµde<¢u

ˆ›
,Cheng¨[41]€9:
犗狊犆犕犜3犪,kù¡(狅狊犮犿狋3犪)mnopuø
犿犘犻狀犵€mqSãde<¢u。@ÓAð,犗狊犆
犕犜3犪,k)œ„…9:,klCHG¿Z()*
+,Nq©ô
,狅狊犮犿狋3犪ù¡mCHG¿Z+,N
9Öìíƒî

7Bõµ,klmSã,k=êB
á

«BCá—,klqö÷†

7N–ABwÙDµÍ

›ÒqçqEù~ô
õµ#$,klq?@¾=>?@Ùö÷†

«B
¦§pdeŸ‰Š,k=ê9Ö¤DNA()*
+,N‚

2Fõµ犿犘犻狀犵de¢u。犿犘犻狀犵

GH

q£†K&i­no,klCá?Uq€
‡s

~K&i­,kl«NNqáâ‡s

4 犿犘犻狀犵<°¥,klq¦§
CëDNAdede

×ÙDEù,klS_

*B
,犿犘犻狀犵I)

:;¾<°

¤

c³¾<°
”2$deûü,Íä
=µÍr:J$ EG4wàs!DâãÆKt
犿犘犻狀犵H¿Z[31],«NABào#$,kl^LØ
âSq©bµ¬
[42],

,犿犘犻狀犵dehM~ôò
pNO‡â

SãABŸ[
,犿犘犻狀犵mb<°,kmv<¾
˜,k³Œ
(2.5kbm),"#ABào,犿犘犻狀犵q
<°~ôPQ<°¿ZR˜,k=êìíÜV,k
Su
[4344]。
*B

Sã<°¿ZT˜,k=ê9Ö
noŸ[
,犿犘犻狀犵×ÙDPQR˜,kq=êû
ü

ìíD€s÷?UNâUÉ;,kq=ê

Yasuda¨[38]Ùy߂—‰IqAB¼,IÐV
µÍ—stœ/qÍï

€WXqçƒ
,犿犘犻狀犵
~¸ä<°¿ZR˜Y†,k=ê9Ö

~s
÷?UâUE$qYW†

øˆZ[
,犿犘犻狀犵~ô
)Ú\üE·æçq©ô

âãÉ;aüE·k<
L‰Š=ê,kq”

«B¸äR˜,kq=ê


Ö~œABµÍ
,犿犘犻狀犵<°3)+,N
9ÖDEù

¦§p<°Z,k mRNAq]:©
ü

øˆ[Í¥‰Š,kqd.3îô
[45]。
5 ¯°AB犿犘犻狀犵de²³œ1F
´µÛ¹õÜùH^_
  ˜Ý,Horibata¨[14]C9¥è`a¾Ù`a
犿犘犻狀犵q2$f`9:bcµÍ

øF犿犘犻狀犵€c9?w`a犿犘犻狀犵qbc<¼Ù`a犿犘犻狀犵qb
c
SiLjc<”q²D

ìí€
c
¢uq犿犘犻狀犵~ômz_n



òó¥犿犘犻狀犵qdeä=p
—Sã;å¾AB

Ö\q€‰Îyãq犿犘犻狀犵
¿Z€c
B6Q¼ÔÕ
q€~ôst؊F犿犘犻狀犵ªÙ¼Ýðde£
†¾²³

DNA+,Ns7¼Ý­€îô,k=ꄅ
de<ö÷†wµ·Y6E·
[7,4647],
É;ABŸ
[犿犘犻狀犵u†L+,N€s÷‰Š†[34],Öu
ø犿犘犻狀犵€mq犜狅狌狉犻狊狋likeg(hªWABšd
»•–+,Nqîô²³

¥ËPÌABµÍ

c
<µÃ9?w

l¢£ H3K9µM+,N~ôõµ
É;_RNAô³de<³ŒDNA+,N9Ö,«
BÑ8îôSãde[48]。犿犘犻狀犵qAB
~uµÍc
6^(¶•JÜ
犿犘犻狀犵Mí MITEsde<~}p‰I²³qî
ôv

²³qÙÚ±B^­—^Žq´µ¶·

×Ø
˜™š¥de
uø犿犘犻狀犵€mqS
ãde<€?@;<€¤,k©ôAB‹Œ‚
Çȃ·

ABµÍ

9:ÙIJ$¤ËPÌ
ÙIJ$w犿犘犻狀犵q;Æ¿Z¤{|y!€ì
íiú
[13,15],
01‘£Z
,Monden¨[43]wkpr
:sït¾EG4‡Ñ150tiú;Æq犿犘犻狀犵,
ˆE­;<marker¥¡FYl)%Á«;yã†
¿Z
(QTL);å,i©÷¿—xy݉Šîô,
6941 ! " # $ % &                   36š
k,
IеÍ犿犘犻狀犵E­ markerqA‚†ìíä
F@Âz{|bc
。犿犘犻狀犵ÏU@}(430bp),Ù
I$$wqu†iúÜV&;Ɛ€ìíA‚
†
[13,15],
!­犿犘犻狀犵i­f¥z{|bcs&q
=G;<marker¸ —~qç,6¥FÉ;,k
lD<€}€ñ,klob€‚DqAƒ¡$
$B„

A—s$œÍq;<marker~kl。
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