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,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.
!"#$:10004025(2016)07149306 犱狅犻:10.7606/j.issn.10004025.2016.07.1493
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> ?:犕犻狀犻犪狋狌狉犲犘犻狀犵(犿犘犻狀犵)^ _`abYcde< (MiniatureInvertedRepeatTransposableElements,
MITEs)fde<犜狅狌狉犻狊狋likeg(hY6ij,^ 9:,klmnopqrstuvq MITEs,^ MITEsS(h
wxyz{|}~)*
deuqijs
,
k
,犿犘犻狀犵^de<ABq。
m犿犘犻狀犵q、de ¡、¢u£¤¥,klq¦§¨©ªqAB«
¬
,
«s®¯°±B MITEsqde²³¤犿犘犻狀犵de@AB
:
de<
;犿犘犻狀犵;犘犻狀犵;犘狅狀犵;9:
CDE6$
: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
edRepeatTransposableElements(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
ClintockÊËwrs̵Íde<,ÎÏ
qÐÑm¼+%ÒÓÔÕ
,
Ö×Ø+%ABqÙÚ
µ¬
,
ÛÜÝÞpde<[ß^àáM$«NqY
6âãs
。
¥ä¨M$,klq;åwµÍ
,
Ûf,klwdeæçqèéê45%,#$,kl
Cëß50%~80%Ù¨[12],ìíÉ;îï+
#$wde<ðñ,klqèé~äê90%[3]。
òóde,
Öø
õµq,kùú
、
=êûüùNìíýþ¡
qYÿ!^M$,kl«NqY6àáâ
s
[45]。
de<"#de²³~¼;2$f`:Class
Ⅰ(ade<)¾ClassⅡ(de<)[6]。ClassⅠ¾Ⅱ
fde<%#&)^()*+deôâ,~
;)5`de<
(autonomouselements)¾-)
5`de<
(nonautonomouselements)[7]。Class
Ⅰde<ø,klDNAd.RNA,/01ad
.RNaseH ad.DNA,234i,ke¿
qdÀ
,
56fde<½ade<
(retrotrans
posons);ClassⅡde<78C9DNA:;/<°
²³[Íde
,
¼½de<
(transponsons)。
ClassⅠde
,klm{|y@A
,
*B
,
SÉ;ClassⅡf
de
<°de£ZCëC
¥@x
q{|y
[8],
&wSÉ;ClassⅡf-)5`de
<
,
&{|yìíÙDg950t[9]。
MITEs(miniatureinvertedrepeatstransposa
bleelements)½_`abYcde<,EFG`
DNAde<(ClassⅡ),^ #$wY6qsfde
<
[10],
ÖL&HClassⅡde<ÙI,MITEs~
C9)Jc³K){|yêpyLMíNO
t
[11]。MITEsµÍFîï+E$ÊËw,3
qABwÛÜP&HsQ´R#$
、
á$Mí
Ûf , k l m µ Í MITEs q S T[12]。ø F
MITEsSAø)5`qde<«NB,ÏU_F
600bp,VK&)WX*+de(Y1),de¢
uZô%[&Hde ¡
,
¥9:,klq;å
w\*µÍ
,MITEs(hð9:,kl8.6%,]
{|y^g_100000tA[13]。%# MITEs`
aabYcbc
(terminalinvertedrepeats,TIRs)
¾d¿ZYcbc
(targetsiteduplications,TSDs)
q£Z
,
5yãeSq MITEs;i犜狅狌狉犻狊狋
likeMITEs¾犛狋犪狑犪狔likeMITEs2tg(h,f
FTSDsCë3g,qTAAhTTA¿Z,B3
FTSDsß^A2g,TA[14],6$iúqjk
~ôøF2t(hqde ¡犘犐犉/犎犪狉犫犻狀犵犲狉
¾犜犮1/犿犪狉犻狀犲狉de(h¥de<<°¿Zq
kls÷mbÓV
[1516]。
犿犘犻狀犵(犿犻狀犻犪狋狌狉犲犘犻狀犵)2n&oF2003,
^9:wnopqrst¢uq MITEs[13,1718],^
^9: MITEseS(hwps~)*qç
C
¢uqde<
[19]。
ÙIF&H MITEs,
Cë犿犘犻狀犵
¥@zq{|y,r:sït,
kluvw70t犿犘犻狀犵{|,x:‘9311’yuv
w14t4z{|[13]。{Fde¢u}C
¥@
xq{|yq£ZK犿犘犻狀犵ide
。
,
|×¥&ABqÙÚ¯°
,犿犘
犻狀犵¤&H MITEsÙy«NABw}pY
Õ
,
~ù¡I¤;<¨pÇÈ
·
[2022]。
1 犿犘犻狀犵q£
犿犘犻狀犵kL)5`de<犘犻狀犵`aäUI
B
,
ÏU430bp,` auv15tg,qTIRs,
TSDsbcCëCq3g, TAAh TTA,%
#TIRs¾TSDsbc£Z5&f犜狅狌狉犻狊狋like
g(h
[13,19,23]。犘犻狀犵ÏU5314bp,q`a
;253bp¾177bpbcL犿犘犻狀犵É;bcs
Vêp99.9%¾100%,Á8li4zq
犿犘犻狀犵bc(Y2)[13]。IÐ,¥9:,kly#
qnwµÍ
,犿犘犻狀犵Lst)5`de<
犘狅狀犵q`abc70%sV,犘狅狀犵ÏU5166bp,
L犘犻狀犵13bpIbcqTIRs(Y2)[24]。L
&H MITEs(hijI,犿犘犻狀犵Ù)*+d
eqôâ
,
k
,
LäUI
})5*+d
eq犘犻狀犵¾犘狅狀犵i犿犘犻狀犵~ôqde
¡
。
þ=de<d¿ZYcbc
(TSDs),þ=de<`aabYcbc(TIRs)
Y1 DNAde<(ClassⅡ)q
TheblacktrianglesflankingeachelementrepresentTSDs,andthegraytrianglesrepresentTIRs
Fig.1 ThestructureofDNAtransposons(ClassⅡ)
4941 ! " # $ % & 36
犿犘犻狀犵ø犘犻狀犵q2t`a8Bi,}L犘狅狀犵aäUI
。þ^de<d¿ZYcbc(TSDs),
þ=`aYcbc
(TIRs)
Y2 犿犘犻狀犵、犘犻狀犵¾犘狅狀犵q
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 ~ôqde ¡犘犻狀犵
¾犘狅狀犵
,E犿犘犻狀犵de ¡q犘犻狀犵
¾犘狅狀犵p¯°qAB。;å[,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&*+$L犿狔犫d.k
(DNAbindingdomain)s÷¥,#à
oORF1*+$~ô®LDNAL¢£,¯
1de<¢u
[2526]。ORF2*+$wuv DDE
,b
,DDE,b°^É;de±Nuw²qY
6£
[27],
}ORF2uvN2、N3¾C1äUC³
,
*+$L犘犐犉/犎犪狉犫犻狀犵犲狉de(h´
ä¥
[28],
¥µ¶ORF2*+$µ·de
©ô
,
Ö«s®AB[
,ORF1¾ORF2犿犘犻狀犵
de9?w!µ·íY6qE·
,
Ö¸);¹ª
Ù4ñ«¬
[24]。
¥F犘犻狀犵h犘狅狀犵ºst^犿犘犻狀犵»:de
¡í§ªW÷¼
。
r:
‘
sït
’
,kl~n
pS½5t犘狅狀犵¾1t犘犻狀犵q{|,*Bx
:
‘9311’,klm¾¿µÍ4z犘犻狀犵qÀT。
IÐ
,Zhang¨[16]¶·ÁµÃ%ÄÁAB[,
Sã#$
、
á$¾ÅÅ,klw!犘狅狀犵
I
bc
(犘狅狀犵like),SÉ;犘狅狀犵like!uv2
tORF,}*+$ƵL犘狅狀犵q2tORFÇÈ
sÉ
[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
,
Z4z犘犻狀犵qWX¢u犿犘犻狀犵 de,
BF ~õµ犿犘犻狀犵de}deÍÎ。
ÏF¥èlÐÑÒ39:ÙIÓÔ3~µÍ
,
犿犘犻狀犵q¢uÕÎL犘狅狀犵 q=ê9Ö,Ö×
صÍL犘犻狀犵=êÙ。;å3ào,
òó犘犻狀犵*+$)Ú¢u犿犘犻狀犵 qôâ,Ö~
ôøFfaÛá<³qù
,
ÜV犘犻狀犵*+³
Ùô°ë=ê
,
N#¥µ!¶犘狅狀犵°^
犿犘犻狀犵qde ¡。
*B
,
Ý9:EG4ÞÁwqAB,ßÙ
Iq¼
。
AB[
,EG4Áw犿犘犻狀犵qdeu
)*ÑÒ~pC
,
×àSw
40t{|qáUÙÚâã,¦§¨EG4,kl
m犿犘犻狀犵 q{|yäåâííx1000t[31]。
Teramoto¨[19]EG4Kæ3~5d3¥ÙIÓÔ
;åµÍ
,
ÎçqµM«°pèa;Mlоé"
³;NêÁÐ
,
éçmnop犿犘犻狀犵qëȾ
<°
,
IÐ|×犘犻狀犵=ê9ÖqìíNî,Íï
&Hõµ犿犘犻狀犵¢uqqçwªØµÍ9,AB
ÛjÝçµMqnÝ~ôõµ犘犻狀犵q£ú
=ê
,
ä=êq犘犻狀犵¥F犿犘犻狀犵 q¢uµ·Y6
E·
。
N~ð
,犘狅狀犵~ô^犿犘犻狀犵 ²qde
¡
,
Bsde<犘犻狀犵~ô#$µÃ£÷
ÐÝ~)Ú¢u犿犘犻狀犵deqôâ,ÖªW#
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
[13,1718],
3ÛÜPµ
Íuøäù
、
¢úûü
、
ýþÿ!¤"#¾$%¨
ÄÁ ~¢u犿犘犻狀犵de[3238],BNqçCë
õµ,klÇÈq?@¾=>?@ùú
,
ÏFq
¹Ew
,
I&Ùyynop犿犘犻狀犵qde¢u,
IеÍ,klqSãùú
[39]。
犿犘犻狀犵¢uqÙIqç,Ede ¡
59417Ý ./*,¨ :9: MITEsfde<犿犘犻狀犵AB«¬
犘狅狀犵hF犘犻狀犵q=êùN^犿犘犻狀犵 µM¢uq
s
。
IÐ
,犿犘犻狀犵mɾ5’` abcq
()*+,N9ÖL&ö÷,Íìí
,
ä-
Uq+,N¥F
犿犘犻狀犵 qö÷µ·Y6E
·
[34,40]。
*B
,Tsukiyama¨[9]ABµÍ,9:È
î.,k
(犚狌狉犿1)ù~õµ犿犘犻狀犵q
de
,
/q^
,
Ð犿犘犻狀犵mɾbcq(
)*+,N9Ö×صMùú
。Yang¨[40]µÍ¶
·01úⅡ2³3(%456)ÿ!9:~7
µ犿犘犻狀犵de¢u,^ øF%456õµ9:,k
lÇÈq?@¾=>?@ùú
。Wu¨[36]¶·9
:
‘
8988’E}¡,:(犣犻狕犪狀犻犪犾犪狋犻犳狅犾犻犪)E
¡«;"#$%
,
"#Á,klwòó¿n
op ¡,klD<
,
Ö¾ßö÷q"#
3
,^
"#Áwnop犿犘犻狀犵qde¢u,k
"#9?w~ôøFÉ;D=;
,
õµ}¡>(
“
,kl?á
”(genomeshock),
7Bõµde<¢u
。
,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)
9:,klCHG¿Z()*
+,Nq©ô
,狅狊犮犿狋3犪ù¡mCHG¿Z+,N
9Öìíî
,
7Bõµ,klmSã,k=êB
á
,
«BCá,klqö÷
。
7NABwÙDµÍ
,
ÒqçqEù~ô
õµ#$,klq?@¾=>?@Ùö÷
,
«B
¦§pde,k=ê9Ö¤DNA()*
+,N
,
2Fõµ犿犘犻狀犵de¢u。犿犘犻狀犵
“
GH
”
q£K&ino,klCá?Uq
s
,
~K&i,kl«NNqáâs
。
4 犿犘犻狀犵<°¥,klq¦§
CëDNAde
,
×ÙDEù,klS_
。
*B
,犿犘犻狀犵I)
“
:;¾<°
”
¤
“
c³¾<°
”2$deûü,Íä
=µÍr:J$ EG4wàs!DâãÆKt
犿犘犻狀犵H¿Z[31],«NABào#$,kl^LØ
âSq©bµ¬
[42],
k
,犿犘犻狀犵dehM~ôò
pNOâ
。
SãAB[
,犿犘犻狀犵mb<°,kmv<¾
,k³
(2.5kbm),"#ABào,犿犘犻狀犵q
<°~ôPQ<°¿ZR,k=êìíÜV,k
Su
[4344]。
*B
,
Sã<°¿ZT,k=ê9Ö
no[
,犿犘犻狀犵×ÙDPQR,kq=êû
ü
,
ìíDs÷?UNâUÉ;,kq=ê
。
Yasuda¨[38]ÙyßIqAB¼,IÐV
µÍst/qÍï
,
WXqç
,犿犘犻狀犵
~¸ä<°¿ZRY,k=ê9Ö
,
~s
÷?UâUE$qYW
。
øZ[
,犿犘犻狀犵~ô
)Ú\üE·æçq©ô
,
âãÉ;aüE·k<
L=ê,kq
,
«B¸äR,kq=ê
9Ö
。
Ö~ABµÍ
,犿犘犻狀犵<°3)+,N
9ÖDEù
,
¦§p<°Z,k mRNAq]:©
ü
,
ø[Í¥,kqd.3îô
[45]。
5 ¯°AB犿犘犻狀犵de²³1F
´µÛ¹õÜùH^_
Ý,Horibata¨[14]C9¥è`a¾Ù`a
犿犘犻狀犵q2$f`9:bc
,
øF犿犘犻狀犵c
c
SiLjc<q²D
,
ìí
c
¢uq犿犘犻狀犵~ômz_n
k
,
òó¥犿犘犻狀犵qdeä=p
Sã;å¾AB
,
Ö\qÎyãq犿犘犻狀犵
¿Zc
B6Q¼ÔÕ
q~ôstØF犿犘犻狀犵ªÙ¼Ýðde£
¾²³
。
DNA+,Ns7¼Ýîô,k=ê
de<ö÷wµ·Y6E·
[7,4647],
É;AB
[犿犘犻狀犵uL+,Ns÷[34],Öu
ø犿犘犻狀犵mq犜狅狌狉犻狊狋likeg(hªWABd
»+,Nqîô²³
。
¥ËPÌABµÍ
,
c
<µÃ9?w
,
l¢£ H3K9µM+,N~ôõµ
É;_RNAô³de<³DNA+,N9Ö,«
BÑ8îôSãde
~uµÍc
6^(¶JÜ
犿犘犻狀犵Mí MITEsde<~}pI²³qî
ôv
?
²³qÙÚ±B^^q´µ¶·
,
×Ø
¥de
uø犿犘犻狀犵mqS
ãde<?@;<¤,k©ôAB
ÇÈ·
。
ABµÍ
,
9:ÙIJ$¤ËPÌ
ÙIJ$w犿犘犻狀犵q;Æ¿Z¤{|y!ì
íiú
[13,15],
01£Z
,Monden¨[43]wkpr
:sït¾EG4Ñ150tiú;Æq犿犘犻狀犵,
E;<marker¥¡FYl)%Á«;yã
¿Z
(QTL);å,i©÷¿xyÝîô,
6941 ! " # $ % & 36
k,
IеÍ犿犘犻狀犵E markerqAìíä
F@Âz{|bc
。犿犘犻狀犵ÏU@}(430bp),Ù
I$$wquiúÜV&;ÆìíA
[13,15],
!犿犘犻狀犵if¥z{|bcs&q
=G;<marker¸ ~qç,6¥FÉ;,k
lD<}ñ,klobDqA¡$
$B
,
As$Íq;<marker~kl。
¶·de<<°SuI#$ù¡äÇÈ
·FA$E$w
,
Çm
vSãudeˤC9d,k^_[Íde<¢uqËPÌ
、
9:¾
¨
[49],
01Sªde<¢u
,
@
}ÐÑm~ßSãù¡[\º
。
*B
犿犘犻狀犵mb<°{|,k³T[31],&<°£
hM÷ÙIFade<犜狅狊17[50],×Ù
FI,k©ôSù¡
,
,N
,犿犘犻狀犵
)zîhâU<°¿ZR,k=êq£Z
[20],
~ôde<Ã$h[\ºINq·¸
H q
。
]
,犿犘犻狀犵 E s f u v q
MITEs,¥F¯°±B犜狅狌狉犻狊狋likeg(hMíz
t MITEsS(hq,5PY6þ,~
D?@¾«NqAB¸ ^AqÄÁ¾^Çq
Õ
。
JK!F
:
[1] WESSLERSR.Transposableelementsandtheevolutionof
eukaryoticgenomes[J].犘狉狅犮犲犲犱犻狀犵狊狅犳狋犺犲犖犪狋犻狅狀犪犾犃犮犪犱犲
犿狔狅犳犛犮犻犲狀犮犲狊狅犳狋犺犲犝狀犻狋犲犱犛狋犪狋犲狊狅犳犃犿犲狉犻犮犪,2006,103
(47):1760017601.
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