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----------------------The Plant Journal (2011) 66, 6679 doi: 10.1111/j.136-313!.2011.0"03.#

T$% P&'T %*+%: ' %*&T*' % * TT% '4 5T*

5ro horone to e8ondar etaoli: the eer;en8e

o< etaoli8 ;ene 8luter in=lant

$oi ee hu, %>a e;el?

and 'nne *ourn@

4e=artent o< +etaoli8 Aiolo;, John nne entre, olne &ane, orBi8h " 7$, C

e8ei>ed 2D *8toer 2010E re>ied 21 4e8eer 2010E a88e=ted 10 Januar 2011.@5or 8orre=onden8e (erit o< arid;e, 4oBnin; treet, arid;e A2 3%$, C.

++'

ene 8luter er;en8e. %#a=le in8lude

the aalian $o# and -;loin lo8i, Bhi8h are reuired

el, and 8luter o< leu8ine-ri8h re=eat ;ene

ao8iated Bith dieae reitan8e in =lant (*ourn and

5ield, 2009).

5un8tionall related ;ene are 8oonl di=ered

throu;hout the ;enoe in euHarote, and 8an e

8o-ordinatel re;ulated in tran. ntil re8entl it Ba

thou;ht that ;ene order in euHarote Ba eentiall

randoE hoBe>er, it i e8oin; in8reain;l 8lear that thi

i not the 8ae ($urt et al., 200"E *ourn and 5ield, 2009),

and that there are nuerou e#a=le o< 8luter o


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67 $oi ee hu et al. The eer;en8e o< eta:oli8 ;ene 8luter in =lant 67

P 2011 The

The Plant JournalP 2011 Ala8HBell Pulihin; &td, The Plant Journal, (2011), 66, 66

P 2011 The


5i;ure 1. The e e8ondar etaoli8 ;ene 8luter that ha>e een re=orted ena8in 8luter ena tri;oa (di=loid oat), re=e8ti>elE the oila8tone and =hto8aane 8luter


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6D $oi ee hu et al. The eer;en8e o< eta:oli8 ;ene 8luter in =lant 6D

P 2011 The


P 2011 The


5i;ure 2. The %8heri8hia 8oli la8 o=eron. The la8 o=eron 8onit o< three tru8tural ;ene that are reuired e


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ariou h=othee ha>e een =ut olution o< o=eron (e.;. Pri8e et al., 2006E 5ondi

et al., 2009). o-tran8ri=tion o< the ;ene Bithin o=eron

a our loBer ;ene e#=reion noie and toi8hioetri8

e#=reion o< the 8o;nate ;ene =rodu8t (o8ha, 200DE

Co>aS 8 et al., 2009). The ot hi;hl 8oner>ed

o=eron tend to 8ode


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5i;ure 3. ene 8luter ie, John nne entre, orBi8h, CE ia;e (a) and (8) Bere Hindl =ro>ided 4a>id $o=Bood, aed on

ine linear 8hroooe (&in et al., 1993).

'lthou;h e8ondar eta- oli8 ;ene 8luter areo88aionall ariale NarO re;ion 8loe to the end o< the

8hroooe, Bherea ;ene al under 8ertain en>iron-

ental 8ondition, and 8an theree ;ene 8luter (*ourn and 5ield, 2009E *ourn,

2010a). n =arti8ular, the relate to the e#=loitation o< neB

en>ironent or the ana;eent o< intera8tion Bith other

or;ani. %#a=le in8lude 8luter iiae) that =erit the

ue o< neB nitro;en or 8aron our8e, the eat 4'& and

'& 8luter, re=e8ti>el ($ittin;er et al., 200"E on; and

ol


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reuired e iunit (intera8tion Bith

=atho;en in the en>ironent) ($orton et al., 200").

uerou e#a=le o< ;ene 8luter ha>e een di8o>ered

in ation a8ro diel, it i =oile that the =riar et-

aoli8 ;ene and their i;nature ;ene 8ounter=art e>ol>ed

e ori;inated inde=endentl in the di8ot

and ono8ot linea;e du=li8ation o< 88loartenol n-

thae ;ene (5ield and *ourn, 200D). 8loartenol n-

thae i an * that i reuired


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5i;ure ". The=reued e>olution o< the Ni;natureO ;ene e ori;inated inde=endentl the re8ruitent o< 88loartenol

nthae () ;ene ed to ha>e een re8ruited alonate-inde=endent =athBa, and taHe =la8e in the

=latid (%ienrei8h et al., 199DE &i8htenthaler, 1999E oher,

1999).

The 8o=oition o< the 8luter

n addition to thee i;nature ;ene the 8luter alo 8ontain

;ene


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5i;ure . on>erion o< the =rodu8t o< the i;nature enIe to =athBa end =rodu8t i ediated tailorin; enIe. 5or the thalianol =athBa a olution o< oe 8o=onent o


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5i;ure 6. %elo=ent. (a) ad3 utant o< '>ena tri;oa (di=loid oat)

a88uulate a late a>ena8in =athBa interediate that i in8o=letel ;l8olated, leadin; to tunted root and a root hair deere dBared inde=endentl in ;rae and eudi8ot (8hulleh-

ner et al., 200D).

ene 8luter


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LaeA, global acvator

~7.5 Mb shown, schemac and not to scale

MHC histone familyMHC immunerelated

MHC olefactory receptor

5i;ure 7. $i;her le>el re;ulation o< ;ene 8luter. The 4'& 8luter i reuired ol>e the


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e#=loitation o< dru; and utation that aer e-ered in the a8tino8ete tre=to8e 8oeli8olor (+8'rthur

and Ai, 2006, 2007). %>iden8e i eer;in; ena8in 8luter, oth o< Bhi8h are de>elo=-

entall re;ulated, to i=li8ate 8hroatin reodellin; in

the re;ulation o< =lant e8ondar etaoli8 8luter. The

thalianol 8luter ha arHed hitone $3 line 27 ($3C27)

triethlation, u;;eti>e o< 8oordinated re;ulation at the

8hroatin le>el (5ield and *ourn, 200D) (5i;ure 7). n oat,

hi;h-reolution 4' e re>ealed that e#=reion o< the

a>ena8in 8luter i ao8iated Bith 8hroatin de8ondena-

tion (e;el et al., 2009).

$'T '% T$% %*&T*' * *5 P&'T

%*4' +%T'A*& %% &T%V

$oriIontal ;ene trane een

throu;h horiIontal ;ene traniiae i alo

liHel to ha>e arien ;enoi8 rearran;eent (on; and

olent a ha>e in>ol>ed ;ene

du=li8ation ed in their ori;inal ;enoi8 lo8ation. n


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5i;ure D. hroooal lo8ation o< the =lant ;ene 8luter. (a) The =oition o< the thalianol, =hto8aane, oila8tone and 2,"-dihdro#-7-etho#-1,"-

enIo#aIin-3-one (4+A*') 8luter, a deterined ena tri;oa (di=loid oat)

8hroooe uin; the ;ene a e;el).

et al., 2006), a =ro8e that Be ha>e =re>ioul tered

N;enoe deiden8e

that run 8ounter to thi ar;uent. 5or e#a=le, the

=reued =ro;enitor o< the ;ene iden8e ed 88loartenol nthae and otui


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8onditional e#=reion o< ;ene throu;h uteloeri8 ;ene

ilen8in; e8hani. elo8ation o< ;ene into utelo-

eri8 re;ion a thereer;en8e

e8aue o< lo8al ele>ated ;enoe dnai8 (AroBn et al.,

2010). The aiIe 4+A*' 8luter i lo8ated 8loe to the end

o< 8hroooe " (5i;ure Da). $oBe>er the 'raido=ithaliana thalianol 8luter and the tBo ri8e 8luter are not

in uteloeri8 re;ion. The oat ;enoe ha not een

euen8ed,ut the a>ena8in ;ene 8luter a= to the end oel re;ula-

tion o< iontheti8 ;ene 8luter. at. he. Aiol. , "62"6".

AroBn, .'., +urra, '.. and ertre=en, C.J. (2010) a=id e#=anion and

er;en8e o< uteloeri8 ;ene art, .J. and er8H-

ei8hart, 4. (200D) 'n e#teni>e (8o)-e#=reion anali tool eriolution o< o=eron

and etaoli8 =athBa. e. +i8roiol. 160, 0212.

5raer, J., oueau, +., henHer, ., 5erraiuolo, +.'., $aahiIaHi, .,

Alan8hette, +. and 4otie, J. (2009) hroatin 8onerin; natural

=rodu8t ed analti8al ethod. n +ethod in %nIol-

o; ($o=Bood, 4.'., ed.) "9. eB orH: '8adei8 Pre, ==. 991.

ierl, '. and 5re, +. (2001) %>olution o< enIo#aIinone ionthei and

indole =rodu8tion in aiIe. Planta, 213, "93"9D.


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ro, J., ho, .C., &eIhne>a, &., 5alH, J., Cru=inHa, C., hinoIaHi, C., eHi,

+., $errann, .. and +eurer, J. (2006) ' =lant lo8u eential itain C1) ionthei ori;inated . enet. , DD9D99.

$urt, &.4., Pal, . and &er8her, +.J. (200") The e>olutionar dnai8 o. enet. , 299310.

Ja8o, 5. and +onod, J. (1961) eneti8 re;ulator e8hani in the n-

thei o< =rotein. J. +ol. Aiol. 3, 31D36.

Jan;a, .., ollado-ide, J. and Aau, +. (200D) Tran8ri=tional re;ulation

8ontrain the or;aniIation o< ;ene on euHaroti8 8hroooe. Pro8.

atl '8ad. 8i. ', 10, 17611766.

Jon8IH, ., 8hidt, $., *terrieder, '. et al. (200D) %lu8idation o< the e 'A

tran=orter 8oniden8e a;aint the ela, .4., AoH, J.., raer, .'., ortan, J.., Ci,

$.., ieran, .. and Celler, .P. (2007) Tran8ri=tional re;ulation oerit in '=er;illu


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$etero8hroati8 arH are ao8iated Bith the re=reion o< e8ondar

etaoli 8luter in '=er;illu nidulan. +ol. +i8roiol. 76, 1376

13D6.

o8ha, %.P. (200D) The or;aniIation o< the a8terial ;enoe. 'nnu. e>.

enet. "2, 211233.

oher, +. (1999) The di8o>er o< a e>alonate-inde=endent =athBa ol>eent o< tran=oon-liHe eleent in =eni-

8illin ;ene 8luter re;ulation. 5un;al enet. Aiol. "7, "23"32.

hiura, C., *Hada, '., *Hada, C. et al. (2007) dentied in deto#i

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