    ]`

RNA .)<=/=,_^iNmRNA (XW8+417

ëì ÷Þ 1,2ĤÕÛ¡ÙŠūłĽÛōŗĊ 1
113-0032 ĤƝė¯øł 1-1-1ĤÕÛÕÛƭęƴÌªĉŐÛōŗŐlX@EsąÎłÀåĕ 2
113-0032 ĤƝė¯øł 1-1-1

Hiro-oki Iwakawa1,2
How do plants avoid self-attack by unwanted RNA silencing?
Key words: Antiviral defense, PTGS, RNA silencing, siRNA
1

Institute of Molecular and Cellular Biosciences, The University of Tokyo, Bunkyo-ku, Tokyo
113-0032, Japan.
2

Department of Computational Biology and Medical Sciences, Graduate School of Frontier
Sciences, The University of Tokyo, Bunkyo-ku, Tokyo 113-0032, Japan.
DOI: 10.24480/bsj-review.8b4.00114

po!$ RNA LAtvMvIĮī* 20−30 ÐÍœõ)ç& RNA ="ŉŶň&ƞ£=4!ƛÙ=ƍtes, 0ƍütes#ċ¥9ƛÙņŀ¥þĮī#9)Įī*¡®5ņł&%Ĭ&ł³Ă=¥þ9)1&7, ÓģħƠ=đƮ9łƪþĮī$")û©4ď
¸<" 9(Ghildiyal and Zamore, 2009; Bologna and Voinnet, 2014; Castel and Martienssen, 2013;
Kobayashi and Tomari, 2015)ľ'ĨĽ5ěŲ&%#*Ź&ČBAsOƪþĮī$"*7$ŋ7:" 9ĨĽ'*ç¡Ù RNA )ĭň RNA

77'ç¡Ù RNA =8ç¡Ù RNA ÑñĮīÚÊ93, RNA LAtvMvI)ĭň$&9BAsO RNA &%*Ʊ # #  !"#        ð'ù Ƈ)¥þ=µ9$$&9 (Ding, 2010))ç¡Ù RNA ÑñĮī)ĞơŹ³Ă*yĢƥ)ĭň RNA =ç¡Ù RNA )¦Ƹ#9Ʀ Ģƥ RNA 'ÒĒ9ƙœ#9, ŭƌ) mRNA =251'Ģƥ'ÒĒ"0 "*Ļőô& RNA LAtvMvIöƉ:,ĨĽ)ƛÙċ¥:9Ĉ'Ư "0 )3, ĨĽ*ıð& RNA $Ņð& RNA=ź¡, Ņð& RNA )1= RNA LAtvMvI'öƎ27 )lE\Pk=ď " 9$ũ 7:9
RNA LAtvMvIĮīņź:" 7Å°}ŝţ, 6 5ıð& RNA$Ņð& RNA =¯¤91¡ 8!!9Ģŕ#*ĨĽ) RNA LAtvMvIĮī,6,ç¡Ù RNA ÑñĮī="Óģ)Ņð& RNA =đƮ9lE\Pk=Īſ9$$4', ŭï) mRNA =ĕĔ& lE\Pk=Ğę)ŋź=Ƌ0 "ũã $ă 

qoMDK



cFT

 QT

 ĨĽ) RNA LAtvMvIĮī* 20
ÐÍ 7 24 ÐÍœõ)ç¡Ù RNA ="öƉ:9ƞ£ľŅň&ƛÙņŀ¥þĮī#9ĮŬň&Ʋ 7*ç¡Ù RNA $ŉŶň& mRNA )¢Ę,Ũżċ¥=ų ƍüNxvLAtvMv I ƺ PTGS: post-transcriptional genesilencingƻ$ŉŶň&ƛÙ= DNA )lUs®="ċ¥9ƍ¥þƺTGS:transcriptional gene silencingƻ) 2 !)ţ
Ɗ'¡ :924 ÐÍ)ç¡Ù RNA *TGS #$ŋ7:" 9yĚ#
20−22 ÐÍ)ç¡Ù RNA * PTGS 'Ʃ< # #  !"#         

9, :7*ł¸ĉƙœ)ƚ '68 microRNA (miRNA)$ short-interfering RNA (siRNA)'Õ¤:9 (Bologna and Voinnet, 2014)

qpo 

cFT

 QT

 Ê)ƛÙ=¥þ9=4! miRNA *J^k'KxZ:ç¡Ù RNA #, RNA hrlqxR II '68d?avīƖ=ď Ʀ ¦Ƹ RNA $"ƍ:9d?avīƖ*DICER-LIKE1 (DCL1)$¿+:9 RNase III ËƟŞ'6 " 20−22 ÐÍ) miRNA Ģƥ'cuQ
MvI:, lUsÍƍŒƟŞ#9 HEN1 '68 3′ġŘlUs®:9 (Reinhart et al.,
2002; Park et al., 2002; Kurihara and Watanabe, 2004; Yu et al., 2006; Li et al., 2005)miRNA Ģƥ*
RNase H ĬZlAv=ď! Argonaute (AGO)$¿+:9Tv`Hƈ'´8Ǝ0:ü, yĚ)ƥƺFAZƥƻ
* AGO 'ĳ8, 4 yĚ)ƥƺ`VQvNnxƥƻ
* RISC7đƮ:9 (Vaucheret

et al., 2004; Iki et al., 2010; Bologna and Voinnet, 2014) "ģ RNA Tv`HƈŸ¸*RNA-induced silencing complex (RISC)$¿+:, miRNA )FAZƥ$ŉŶň&ƞ£=4! mRNA=¢ĘwŨżċ¥9ƺÈƼƻ(Bologna and Voinnet, 2014; Iwakawa and Tomari, 2015)MuA][P[* 10 ŔƷ) AGO =KxZ" 8, ' AGO1  miRNA $ RISC =úĉĭňƛÙ)
ņŀ=¥þ9qqo 

cFT

 QT

 Ʀ Ģƥ RNA 687:9ç¡Ù RNA = siRNA $¿-Ʀ Ģƥ RNA *Ĭ&ţƊ#ł¸ĉ:9, BAsO RNA )ƹİīƖ5Ÿŷ~ƨ, ²Ě»)ƍ'6 "ģĢƥ
RNA, "üƒ9 RNA ÚĄ RNA hrlqxRƺRDR: RNA-dependent RNA polymeraseƻ'6 "ģĢƥ RNA &%Đ7:9 (Ding, 2010):7)Ģƥ RNA * DCL2/3/4 '6":: 22, 24, 21 ÐÍ) siRNA 'cuQMvI:924 ÐÍ) siRNA * AGO3, 4, 6, 9
'Ť¸, ' TGS 'Ʃ<9 (Havecker et al., 2010; Zhang et al., 2016; Mi et al., 2008) yĚ, 21 Ð

 # #  !"#         Í, 22 ÐÍ)ŌsiRNA * AGO1, AGO2, AGO5 $ RISC =úĉ PTGS 'Ʃ<9$ŋ7:"

9(Mi et al., 2008; Takeda et al., 2008; Montgomery et al., 2008)RISC úĉƙœ*`VQvNnxƥ
)đƮ' AGO )¢ĘĶĄ=āŹ$9Ó* miRNA $¹Ĭ#9$ũ 7:" 9 (Iki et al.,2010)ƺÈƼƻ
qroMDKV&D_^ MuA][P['* 10 ŔƷ) AGO KxZ:" 8, yƜƦĄ4Ž37:9, ::Éğ)ĮŬ=ď " 9 (Bologna and Voinnet, 2014)ŢŦ'69ņŀƣ)î5, ņŀžæ)ğ
Ļ&%'68, Šū#ņŀ" 9 AGO )_qvOŅ&9, ŸĖ) AGO *yŠū#¹ĝ'ņŀ" 9(Bologna and Voinnet, 2014)
)6 &Ŀĵ'"ç¡Ù RNA %) AGO$Ť¸9) *ĩ3"ơŹ&Äƶ#9MpBNpB_D#*ç¡Ù RNA Ģƥ)ú)ƚ
'68ƺ~Ö)jOiVU)ğĻƻƽ!9 AGO )%7'9 =Ĵ3" 9 (Tomari etal., 2007; Kawamata and Tomari, 2010; Montgomery et al., 2008), ĨĽ'"*'ç¡Ù RNA )5′ġŘ)ÐÍ%) AGO $Ť¸9 =Ĵ3" 9ĨĽ) miRNA )Ô* 5′ġŘ U #8,
5′ġŘ U )ç¡Ù RNA $ùŤ¸9Ąƈ=ď! AGO1 $Ť¸ĮŬ=ĥ (Mi et al., 2008;
Takeda et al., 2008)0 AGO2 * 5′ġŘ A, AGO5 * 5′ġŘ C )ç¡Ù RNA $::Ť¸
9$ŋ7:" 9 (Montgomery et al., 2008; Mi et al., 2008; Takeda et al., 2008)AGO7 *ç¡Ù RNA ) 5′ġŘ)ÐÍ)1&7 miRNA Ģƥ)ƞ£5īƖ=UCVH9$# miR390 $miRNA $ľŅň'Ť¸9$ŋ7:" 9(Endo et al., 2013; Montgomery et al., 2008)

ro]`UMDK IO_^rp*);0 S\MDK zƒ6

 IO_^'BAsOćĦ9$, DCL2/4 )'68Ÿŷ~ƨ&%)Ʀ

Ģƥ

 # #  !"#        

RNA

7BAsOńģ) siRNA 8:, RISC ="BAsO RNA *¢Ę:9

(Bouchéet al., 2006; Deleris et al., 2006; Diaz-Pendon et al., 2007; Fusaro et al., 2006)
ĨĽ*ç¡Ù RNA=Ññ9Įī=ď!$#68ù«'BAsO)Ÿŷ=ƫá9$#9)~Āňû
©=č )yĢƥ RNA =Ģƥ RNA 'ÒĒ9 RDR #9MuA][P['* 6 ŔƷÚÊ9BAsO)ç¡Ù RNA Ññ'"ơŹ& RDR * RDR1 $ RDR6 #9 (Wang et al.,2010; Garcia-Ruiz et al., 2010; Diaz-Pendon et al., 2007; Donaire et al., 2008)RDR1/6 * RNA Ť¸Tv
`Hƈ#9 SGS3 5))Tv`Hƈ)¬=8" RISC ¢ĘBAsO RNA =Ģƥ RNA 'ÒĒ, )Ģƥ RNA = DCL2/4 õ¢Ę9$#ç¡Ù RNA *Ññ:9ƺÈ
2ƻ(Ding, 2010)

 # #  !"#        rqJ[mAKLB-.5<2/9=>   
 1990 ó Napoli 7şů)fUo\?'ůŞ¸ĉƛÙ#9EsKv¸ĉƛÙ=æƙ¨ņŀư, Ć'³?vYM?\v¸ĉ*ƫá:Ňů)Ű=Á9=ņź(Napoli et al., 1990)EsKv¸ĉƛÙ mRNA )űŖƣ=ƀ/"19$ƙ¨ņŀ:" & +8 , ƢłË) 50 ¡) 1 œõ0#ĸè" (Napoli et al., 1990)æÓģƛÙ)1&7ŉ¹ƞ£=4!ÊƛÙ)ņŀ0#ċ¥9KLctVMpv$º7:)ŀƆ*, BAsO RNA =đƮ9Ï¸$¹Ĭ'ç¡Ù RNA )Ññ= RNA LAtvMvI'6 "öƉ:9$¡" 9 (Brodersen and Voinnet, 2006)BAsO RNA )Ï¸$Ņ&9Ɯ¡*ƙ¨ņŀÓģƛÙ*Ê mRNA $¹Ĭ)yĢƥ RNA $ĺ#9,

üƒ96 'ĨĽ*ÓģƛÙńģ)Ņð&mRNA =ź¡ RDR6 '6 "Ģƥ®9
$# RNA LAtvMvI=öƉƺÈ 2, 3ƻyĢƥ RNA '68 RNA LAtvMvI=öƉĮī)$= S-PTGS (Sense transgene-induced post-transcriptional gene silencing)$¿-rr  hlMDK IOg(?CHMDK



cFT

 yƜ)Ê RNA * RDR6 '6 "Ģƥ RNA 'ÒĒ: trans-acting siRNA (tasiRNA)$¿+:9Êç¡Ù RNA =8$#, ŉŶĄ=4!)ÊƛÙ)ņŀ=¥þ9$
ŋ7:" 9(Bologna and Voinnet, 2014)ĞƐ'& ", ĜŎ&ĭň*ď&  RDR6 Úň'łĉ:9Êç¡Ù RNA 4ĖÔņź:"(Fei et al., 2013):7)ç¡Ù RNA *phasiRNA (phased, secondary, small interfering RNAs)$¿+:9 tasiRNA )łĉ'* RDR6, SGS3,))Kb>HTx)', ľĲ& RISCƺ22 ÐÍ) miRNA $Ť¸ AGO1 5 miR390 $Ť
¸ AGO7ƻÊ) TAS RNAƺTRANS-ACTING siRNA (TAS) precursorƻ'Ť¸9$āƳ#9$Î½:" 9 (Fei et al., 2013; Arribas-Hernádez et al., 2016; de Felippes et al., 2017)
tasiRNA/PhasiRNA ţƊ* S-PTGS Įī$" 9ĺÔ , S-PTGS ţƊ'ľĲ& RISC )Ť¸

 # #  !"#        āƳ ¼ *ĠĜ7 #&so  .)<=/=,iN(Branscheid et al., 2015)

  (XW8+417 ç¡Ù RNA )Ññ= RNA LAtvMvIĮī*à'ù«&łƪþMOWk#*9, ¥þ&Ê) mRNA 'Ń9$ƛÙņŀ:, ĨĽ)łŪ*ƫá:"0 &Ê) mRNA *ç¡Ù RNA )Ññ= RNA LAtvMvIĮī)ĭň'&7& )#;ƾĞƐ)ƃ=ŻĴ9 4:& ŸĖ)Ɓėņŵ:sp′Ze6:  n RE ¦ƒ6 ', ÓģƛÙ=æ9$, ĨĽ* RNA LAtvMvI=öƉ, ÓģƛÙ)ņŀ=ċ¥9Ÿŷ~ƨ&%)Ʀ Ģƥ RNA =8BAsO RNA $Ņ&8,ƙ¨ņŀÓģƛÙńģ) mRNA *Ê mRNA $¹Ĭ)yĢƥ RNA #93, ĨĽ*%' "ÓģƛÙńģ) mRNA =Ê mRNA7¯¤" RNA LAtvMvI=öƉ&"*&7& ŊħłĽ) mRNA * 5′ġŘ'GnVcīƖ, 3′ġŘ'hr A ƥ=ď, :7*Ũż)Ƙ 6, mRNA )Ýß®'â|" 9 (Gallie, 1991):0#)Î½#, Gn
VcīƖ5hr A ƥ=į Ņð& RNA  S-PTGS =öƉ±Ç'&9$ŏÃ:" (Luo and Chen, 2007; Gazzani et al., 2004; Parent et al., 2015)Ʀ ƨĨĽ%)6 &¡ÙĮī#hr A ƥ=į  mRNA =ľŅň' RNA LAtvMvIĮī'æ *ƃ# , ĞƐ,
RDR6 '*hr A ƞ£=į  RNA )1=Ģƥ RNA 'ÒĒ, 3′ġŘ'hr A ƥ=4!ıð
& mRNA *Ģƥ RNA 'ÒĒ& $Ů¾ķ ÍƈľŅĄ9$Ĝ7 '& (Baeg et al., 2017)&<ıð& mRNA 4! 3′ġŘ)hr A ƥ*:0#ŋ7:"  RNA)Ýß®5ŨżƘ#&, RDR6 Úň& RNA LAtvMvIĮī)ĭň'&7& 6'9,<+ “ŭï mRNA ) ID” $")û©=( " 9$Ĝ7 '& Ů¾ķ $' TAS RNA 'Ʃ"*, hr A ƥ 7Ģƥ RNA ¸ĉ=ƧØ#96 #9 # #  !"#        

(Rajeswaran et al., 2012) 7ľĲ& RISC * TAS RNA 'Ŗĩň' RDR6 =¿,Ǝ2*7
=" 9)$ũ 7:9 (Arribas-Hernádez et al., 2016; de Felippes et al., 2017)

sq Gkfb/037%" @j/037 RE ł#*ð' RNA )ÂƈŚŁMOWk­" 8Ņð& RNA =Əƕ'¡Ż9$#:7)űŖ=ƪ # 9(Inada, 2013)0ıð& mRNA 4ð'ƍ$¡Ż)LAHs=ŧ8
Ƒ$#, ­ň&òŴĿĈ= " 9¡Ż=Ƙ9Ĭ&Tv`HƈÇÙ5MOÇÙ*ĖÔź!

" 9, ıð& RNA 4Ņð& RNA 4Ğšň'*DGSrg]Ht?xR'6

9 5′3′Ě»0* 3′5′Ě».)¡Ż=µ9(Garneau et al., 2007)mRNA *ƔðGnVcīƖ$hr A ƥ'6 "DGSrg]Ht?xR'69¡Ż 7ƌ=Ü " 9, DvZrg]Ht?xR)¢Ę=µłhr A ƥ=į  5′) RNA Ęļ* 3′5′ DGS]rgHt?xR'
6 ", GnVcīƖ=į  3′) RNA Ęļ* 5′3′ DGSrg]Ht?xR'6 "::¡Ż:9 (Garneau et al., 2007)DvZ]Ht?xR'69¢Ę=& ¡Ż'"*,0hr A ƥ¡ŻƟŞ'6 " mRNA )hr A ƥŌť:, GnVcīƖXGnVavI
ƟŞ'6

"´8Ʈ

:ü, Ğšň' 5′3′ DGSrg]Ht?xR'6"¡Ż:9(Garneau et al., 2007)
 :0#ŸĖ)Ɓė# RNA )ÂƈŚŁ'Ʃ<9Tv`Hƈ5DGSrg]Ht?xR& ĨĽ'", S-PTGS Ñù:9$Î½:" 9(Moreno et al., 2013; Gazzani et al.,

2004; Branscheid et al., 2015):7)Ťĥ* RNA )ÂƈŚŁMOWk5 RNA )ƄţƊç¡Ù RNA Ññ= RNA LAtvMvI$ĎČ" 9$=ŏÃ" 9 (Tsuzuki et al.,2017)
ĞƐ'& " mRNA )¡ŻţƊùƫá:9$, ĨĽ)Ĭ&ÊƛÙ 7ç¡ÙRNA 8:Ê)ƛÙ=Ĺ8'ċ¥93, ĨĽ)łŪ'Ņð=$Î½
:(Martinez de Alba et al., 2015; Zhang et al., 2015))łŪƫá* RDR6 5 SGS3 &%) RNA
LAtvMvIƩƗÇÙ=įē9$Æÿ9$ 7, DGSrg]Ht?xR'69 RNA # #  !"#        Ƅ*ŭƌ) mRNA ç¡Ù RNA Ññ= RNA LAtvMvI)äƆ'&9$=ƪ
# 9$ŏÃ:9 (Martinez de Alba et al., 2015; Zhang et al., 2015)

sr  .)<=/=,iN  (XW8+417 #$ :0#ŻſŤĥ=0$39$İ)6 &mXsũ 7:9Ýß'ÚÊ" 9ÊmRNA *hr A ƥ=4 " 93 RDR6 '6 "Ģƥ®:& ƺÈ 3 ízƻ0, RDR6)ƤË$&8ý9hr A ƥ=į Ņð&Ê mRNA *Ĭ&±Ç'68ł"* 9, Ɣð* RNA ƄŜ'6 "Əƕ'¡Ż:" 8, RDR6 )Žƅ=:" 9)$ũ 7:9ƺÈ
3 í{ƻÓģƛÙ=ƙ¨ņŀÏ¸ř'"* RNA ƄŜ'69¡ŻƓ !  RDR6

×2hr A ƥ=į Ņð RNA űŖ93, RNA LAtvMvIöƉ:9)#
; $ũ 7:9ƺÈ 3 ·ƻ # #  !"#        '&
 RNA LAtvMvIŭï) mRNA =ĕĔ& lE\Pk$",Æ mRNA )hr A ƥ$ RNA ƄţƊ)ơŹĄ=Đ, RDR6 5 SGS3 )Ƶś=úĉ9$'68ç¡Ù RNA Ññ=öƉÏ=Ƭé9$5 (Kumakura et al., 2009), RNA ) 3′ ġŘ'ÚÊ9ùÉ&
RNA īƖ5 RNA Ť¸Tv`Hƈ&%4ŭï RNA = RDR6 Úň& RNA LAtvMvI= 7
Ü9*7=" 9¶ŬĄ49 (Baeg et al., 2017)ĨĽ*:7)Ôêň&¥þĮī=4!$'6 "*3"Ƃ )§$4&8ý9ç¡Ù RNA Ññ= RNA LAtvMvI
$

ĩ3"ù«&ħƠƪþĮī=ðĝƞ, ÊƛÙ¥þ'0#ĂŃ#" 9) 4

:& 

PdYa

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