Lewin γονίδια χ ( μέρος iii)

Meroypoqn Kol
q;"
r q
"4
't{ n
MeroypoqlKoi Mnxovlopoi
25 3u r?
OoToypoQ(o ;le tqv o6eto rov Jie Ren rot Jun Ma, U-
niversity of Cincinnati College oI Medicine rot Cincin-
nati Children's HosPital.
Metoypoqrl otouq [lPoxoPutireq
Metoyporpn orouq Europudrreq
Iuppoqn rt EneEepYooio rou RNA
Iro0eporrlro rL Evtontopdq rou mRNA
Kototrurtro RNA
Meroqpoorl
XprlotgonottitvToq T0 [evertro K66tro
KEOAAAIO 19
KEOAAAIO 20
KEOAAAIO 21
KEOAAAIO 22
KEOAAAIO 23
KEOAAAIO 24
KEOAAAIO 25
509
III

O Phantatomix/Photo Researchers. lnc
ffi * K0T0 rIV ertqrlKuvon, I UeToypoqxn Quo0tr[60 UcToKI-
veiror roro prlrog rou DNA rr r; otruoOo rou RNA enr-
prlruveror oe roreuOuvorl 5'-+3' pe npooOflr4 vourtre-
ond[olv oro 3' oxpo.
* H Ueroypoqrl oropord ror ro d[r],trrvo rou DNA [ovo-
oXrlporl(erot g6trrq q RNA notrupepdorl onoouv6e0e[
ond ro DNA erpoye(o oe Uro 0eon repponopou.
H Borqprorrl RNA l-lotrupepooq Anoretre[ror onri l-lotrtr6g
Ynopovodeg
* 0r nupqveg rtov porqprorov RNA notrugepoorirv e(vor
notrupeprl oupntroKo -400 kD pe yevxl Oopn 02pB'o.
e H rordtruol enuetrelrot on6 rtg unopovodeg p xor p'.
To Otr05v(upo r4g RNA l-lotrupepooqg Anorele(ror ono
rov lluprjvo rou Ev(rlgou ror tov llopoyovro lypo
* H poxrrlptorq RNA notrupepriorl uno6rorpeiror orov
0z0p'o nupnvo rou ev(upou nou roro),uet Tn UeToypo-
qr1 ror tqv unopovodo o nou onoue(ror privo yro r4v
evop[r;.
ffi
ffi
@*83
Meroypoefi oTouq flpoKqpudlreq
Etooytrlyrt
* H Ueroypoqn npoyporonore[ror Ue Qopo 5'npog 3' o'
dvo ergoye(o nou 6Xet qopo 3'npog 5'.
H Meroypotprl llpoyporonore(rot ge Zeuyopropo Booetov
oe gro u0uootr[60, A(euyoporou DNA
* H RNA notrupepoor; OroXopr(er toug drlo rl,drvoug rou
DNA oe pLo eqlgeprl <quootr[do> rot Xprloqionolei rov
6vo rtrdrvo og erpoye[o yro vo roreuOuver q ouvOeo4
luog oupdrlpolUolrnq otrtr1trouXrog RNA.
* To Uflroq rqg EuootrOog ervor 12-14 bp evtir rou uppt-
d(ou RNA-DNA p6oo oq rpuootrOo 8-9 bp.
H Avr(dpoon qq Meroyporp4g'EXerTpio lro6ro
" H RNA notrupepdorl npoo06veror oe pto 06011 unorr-
vr1rr1 oro DNA yro vo oXrlpor[oer 6vo rtrerord oupntro-
K0.
* H RNA notrupepooq [envo q peroyporprl oqou ovor[er
ro d(xtrtovo rou DNA yro vo oX1por(oer pro peroyporprrq
rpuootri6o.
510 KEOAAAI0 19 Meroyporpq otoug l1poroputirreg
ffi
19
r.t
I
rss
Ew tp6)'eta Keqt€vou : Richard Gourse

tl 0 nopdyovToq o Tpofiororel lg rdulqreg npoodeolq
oro DNA r4q RNA notrupepoorlg, 6ror drore n ouyy6-
vero qg yto ro yevrro DNA vo ger6verot ror, ovr[Oero,
1 ouyy6vero Trlq yro Touq unouvrlrdg v' ou[over.
l16q Bp(oxer H RNA l-iotrugepdorl rLg Atrtr1trouX(eg rov
Ynoruv116v;
* H np6odeon rrlg RNA notrupepoolq otoug unoxrvlreg
e(vor rooo ypnyopl tlore potrtrov dev nporeuor yr'
ontrr1 6roXuor1.
* H RNA notrupepoorl npoodeveror oe ruXo(eg 06oerg
oro DNA ror lg ovro),trooer pe dtrtreg otrtrntrouxieq 6otq
drou ppe0ei evog unorrv4qg.
To 0tr06v(u;ro Tpononoe[ror Koro rr1 Arepyooio Avoyvdr-
pronq Kr Anetreu0epoorlg on6 roug Ynorrvrlr6g
s OTqv Il RNA notrupepoorl npoo6everor o' rlvov unoxr-
vt1it1, dLoXropt(er roug rtr6voug rou DNA yto vo DrlpL-
oupyrloet Uro UsToypoQrKn Quootr(60 xr evotopordrver
vourtreoridro oro RNA.
* H rvop[n ono rrlv RNA notrupepoo4 dev elvor novrore
enuuxqg.
* 0 nopoyovrog o(ygo ouvrlOoq onetreu0ep6veror ono
qv RNA notrupepoor; orov 1 veoouvre0eqlevrl otruoOo
RNA onorrrloet pflrog -1 0 pooeolv.
0 fiopoyowoq Irypo Etr6yXer rrlv l-lpdodeol oro DNA
Avoyvop((ovrog IuyxerpLp6veg Atrtrqtrou1ieq rov Ynoxr-
vrlrdv
s 'EvoS unortvrlrrlg npoodropl(erot onri qv nopouo[o
ppoXeuilv ouvorvenxrirv oltrrltrouXuirv oe ouyxerprp6-
veg 06oerg.
e 0t ouvorvtTrreq otrtrqtrouXieq Tou urroKrvrlrrl ouvrl0oq
onore),ouvror on6 gro noupivq oqv oqeqpro, evo
e[0Uep5q Ue otrtrr]troux(o nopepqepl Ue rnv TATAAT
orq 05or; - -1 0 xr 6vo otrtro e[ogepdq ge gro otrtrr1trou-
XLo nopegqeprl pe r1v TTGACA oq 0eor1 --35.
* 0t Eexttptorol unoxrvlr6g ouvrlOog drorp6pouv ond rr1
ouvorvelrrl otrtrr;trouX(o oe Uto n neproorirepeg 0e-
0etq.
e H onoretreogortx6rrlro rou unortvqq pnope( eniolg
vo enlpeo(eror xor on6 enrnprioOero orotXe(0.
H Anore),eopotxdn1ro rov Ynorrvqrtlv Mnope( v' Au[o-
ver 11 vo EtrLordveror ono Merotrtro[4
e 0r uopvlnr6qD UeToMoqerq nou pertirvouv qv orroTt-
leogortxoqro rou unorrvlrrl ouvrlOtrrg pertilvouv tr1
oupg6prprooq npoq nq ouvorverx6g otrtr1trouX(eg, evtrr
or uoeflrdq, perotrtro[erq 6Xouv ro ovrf0ero onor6tre-
0u0.
* Merotrtro[erq oqv -35 otrtrntroux(o pnope( vo enrqped-
oouv rr1v opXrxl npdodeol i1g RNA notrupepoolg.
* Merotrtrd[etq onlv -1 0 otrtrr;trouX(o pnopel vo enrlpeo-
oouv Trlv np6o6eol 11 n1v ovridpoorl onodroro{qg nou
perorp6ner 6vo x),erord ofgntroro oe ovorXrti.
l-lotrtreg lleptoXeg qg RNA l-lotrupepoorlg 'EpXovror Ape-
oo o' Enogrl pe rov DNA Ynorrvrlq
q H do[rl rou o70 ol],0(er orov ouv66er0r Ue Tov nupflvo
tou ev(upou, enupenowog ortg neproXtlg yto npoode-
o1 oro DNA v' otrtrqtrenr6pooouv Ue rov unorrvlrfl.
* l-lotrtr6S neptoXdg rou 070 otrtrrltrenrdpouv Ue rov urroKr-
vrlTn.
* H unouovodo o enfor1g oul.rpotrler on]v 0rc, , -- --
rou unortvrlq"
H Anorunoorl e(vor Mrl0odog YmInS Avotruoqg yra ::
Xoporrrlprop6 rtov Atrtrrltrenrdpooetrlv RNA l-lotru pepoorl c -
Ynouvrltrl ror, [evxdrepo, D NA- l-l poreivtirv
* 0r 0uv0rvtTrK6q otrtrltrouxreq -35 ror -10 nop6Xouv ro
neprooorepo onpeio enoqnq yto rrlv RNA notrupepoorl
orov unortvrlrfl.
6 T0 qgelo enoqrlg pp(orowor rupro:q or4 pro ntreupo
rou DNA.
0r Atrtr4trenrdpooerq Mero[0 rou I-lopoyovro Iiypo rot
rou fluprlvo ir;q RNA llotrupepooqq Atrtro(ouv Koro 11
Atotpuyrl rou Ynoxrvlr1
" Mro neproXrl rou otylro rorotroppdver ro 0toutro e[ridou
rou RNA rot npener vo perouvr;0ei yto vo dreurotrtivet
rr1 ouv0eorl rou RNA.
* Avenuuxe(q evdp[erq ouv1Oog ou;rpofvouv nporoU ro
6v(u;ro oXqpor(oet 6vo npoyporrr6 oupntroro entgrl-
KUVorlq.
& 0 flopoyowoq orypo ouvrl0oq onetreu0epoverot ond
qv RNA notrugepooq g6trrg 1 veoouvre0ergevil otruol-
6o RNA vo qrdoer oe Urlxoq -l 0 nt.
H Kpuorotrtrrrl Aoprl Ynoderrvuer'Evo Movr6tro yto 11
Meror[vr1o1 rou Ev(ugou
e To DNA petouvelrot 6rop6oou evdg droutrou oqv RNA
notrupepoorl Kor npoypoTorrote( pto on6roprl orpoqn
oro evepyo r6vrpo.
* Atrtroyeq orrl 6roprlpgtrloq ouyrerprp6vov eu6trrrrov
e[oprnporov p6oo oro dv(upo etreyXouv qv e(oo6o
rov vourLeond(orv oro evepyo x6vrpo.
Mto Iropoqpdvl RNA llotrupepoon Mnope(vo iovopXioer
# Mro oTopoTrlpeq RNA notrupepoorl pnopei vo [ovopXr-
oer rn UeToypoErl xripovrog ro RNA peroyporpo yro vo
Dqproupy1oer evo rorvoupyro 3' oxpo.
H Borrqprorf RNA l-lotrugepool Teppor((er oe Atorpudg
0eoerg
€ Ynopxouv 6uo ro[erq otrtrr1lou1uirv reppolopou. Erei
veq nou ovoyvtop[(ovrot prlvo ono rrlv idro rrlv RNA
notrupepoorl Xop(q rr1 oupperoXq rovev6g rurroprrou
nopdyovro ouv40rrlg ovorp6povror a "evdoyaveC
reppouoreq" (intrinsic terminators). Atrtreq onououv
1-rLo xurroptrrl nporelvrl nou ovopo(eror po (p) xor ovo-
tp6povtot oq (TepuoTtoTtig e[opr6gevot onri qv p,.
* 0 evdoyevnq reppouopog npoUno06rer ovoyvtirpro4
ptog otrtrqtrouX(og reppouopou oro DNA nou rordrxo-
norei pro 6opn Qoupr6rog oro RNA npoi6v.
6 T0 orlpoTo yro repgolop6 pploxovroL rupltrlg gSoo oe
otrtrltrouXreg nou 6Xouv n0n UeroypoQe( on6 qv RNA
notrupepdoq xr 6rol o repporLoprig ppforeror un6 rov
6treyXo rou erpoye(ou ro/r1 rou RNA npoirivrog nou
peroypoqer n notruUepoon.
llrirg AeLroupye( o l-lopdyovrog p;
€ 0 n0p0y0vT0g p etvot pto nptrlrefvq nou npooddveror
oro veoouvre0eq:6vo RNA rot perorrve[ror xoro pqrog
tou RNA yro v' otrtrrltrentdpooer pe qv RNA notrupepo-
orl Kor v0 qv onetreuOepdloer on6 ro o0gntroro enrpl-
KUV0rlq.
#wa
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ouvey[(erut ou1v enopevr1 oelido
511

ffi H Ynepetriro:oq Ervot lqpovrx6 XoporqptorLxo fllq Me-
roYporllq
" H opvlnrr; unepetr(roo1 ou[dver qv onoretreoporuo-
qro ronourlv unortvrlr6v eneroq poqSd qv ovr(6poor1
onodroro{1q.
a H peroypotprl dqproupye( gnpooro ono ro 6v(upo Oel-
rdg unep6trtreg ror n[otrr rou opvrllr6g unep6],rxeg, or
onoleg np6neL v' otpoLpe0oUv onrl rr1 yupootl xot qv
ronoioopepool.
qlE}i H RNA l-lotrugepool rou 0oyou T7 Elvor 6vo Xplorlio
l-lporuno IuoTlUo
& H orK0y6vsr0 ro:v RNA notrupepootirv tou T7 e(vor g.rovo
notrunenr(Oro trovo v' ovoyvrop[(ouv roug unorrvrlr6g
rou qoyou ror vo enuel,ouv notrtreg ond ng dp0orrl-
proqreg tov notrupeptirv RNA notrugepootirv.
* 0r rpuorotrtrrr6q dog6q rov RNA notrupepoorlv qg oL-
roy6verog rou T7 pe ro DNA rouronororjv qv neptoXrl
npoodeolq rou DNA, oro evepy6 r6vrpo xor uno0et-
rvuouv ;lovre),o yro rri 0rorpuyrl rou unorrvqq.
ffi O Avtoytr-rvro;rdg yro rouq l-lopoyoweg liypo Mnoper vo
Pu0p(oeL qv'Evop[1
* Io E. coli eXer enrd nopdyovreg o(ypo, xo06vog ono
roug onoloug o0ei qv RNA notrugepoorl vo (errvfloer
oe pro opodo unoxrvrltrirv nou npoo6top((ovror ono
ouyrerpLp6veg otrtrntrouX(eq -35 xot -1 0.
* 0t evepyrirnroq ToJV 0rorpopertr6v nopoydvrtov otylto
pu0pi(ovror ontl Dtorpopelxou g Lr4Xovropoug.
ffi Or flopdyovreg l(ypo Mnope( vo Elvor Opyovrop6vor oe
Axotrou0ieg
* Mro oKotroue[o nopoyovrov oiypo dnproupye(ror orov
dvog nopoyovrog oiygo onoueiror yro rq peroyporprl
rou yovr6[ou nou rodrronorel rov enoplevo nopoyov-
tu,
* To npdrruo yov[dto rou qdyou SP01 peroyporpovtor ono
rnv RNA notrupepoon rou [evrorr1.
e 'Evo
0rt0 ro nptirLgo yov(6ro roDrronotel 6vov nopoyov-
ro o(ypo nou trlOei r;v RNA notrupepoorl vo peroypo-
ger ro evdtdgeoo yov(dto.
19.1 Erooyoyfi
* H Ueroypoqrl npoyltotonoteirot pe qopo 5'npog 3' o' 5vo er-
poye(o nou eXet qopo 3'npog 5'.
H pretoypo<pr1 nopdyer pto oluo(6o RNA ge raut6o4-
pry aMqAouy[a Ue rov 5vo x].6vo tou DNA, nou Uept-
x6g gopSg role(ror x<o6rxonor4rrxrig xArivog
(coding strand). Aut6g o r<Ltovog SXer <popd 5'-+3'
xor e(vor o u pnlr1papaux6q (co m ple mentary) (6nLodn
(euyopcbver) ge ro eKpoye(o (template), nou SXeL <po-
pd 3'-+5'. I-t' out6 o nop6protog pe ro RNA x).rilvog
* Auo ono ro ev0Lopeoo yov(6ro roduonorouv unopovo-
6eq evoq nopoyovro o(y;lo nou o0e( qv RNA notrupe-
poorl vo ueToYpouer ro oQrlro yov(6r0.
H Inoproyeveon Etreyxeror ono l-lopoyovreq lypo
* H onoproy6veoq 6rope( evo porrrlpro o' evo prlrptro
rurropo nou urpiororor truor1 rr dvo onopro ttou oTit-
treu0ep6verot.
I Koee [eXopror6 6ro;rSprogo npoXoper oro enopevo
oro0ro ovdnru(qg ouv06rowog 6vo rorvoupro n0p0-
yovro olypo nou erronl(er rov npolyorlpevo,
* H rnrKorvovlo Uero[u rorv Duo dropepropdrtov ouvrovi-
(er qv eniloyri rou rorotrtrr1trou Xpovou yro Trq ovTrKo-
tootooetg ttrlv nopoyovrtov oty|to.
0 Avtrepponopoq Mnopei vo Elvor'Evo Pu0prolxo Iup-
tsov
* 'Evo o6untroKo owrrep[oTropou enup6ner orrlv RNA
notruUepdol vo 6ropooer 6topdoou reppouordtv.
* 0 Qoyoq trou0Do xpnotponorc[ ouorrlporo ovTtTtpuoTr-
olrou yro vo puOpioer rooo ro npdrgo ooo Kot T0 otIiUC
perdyporpo, otrtrd ro duo ouorrlporo treuoupyouv p' ev-
retr6g drorpopenro0g plXovrol.roug.
* H rpooDeq nopoy6vrotv oro veoouvteOetg6vo RNA
ouv66sr rtq nporelveg ovTrTrpgoTroUo0 oq 0do1 iou
reppolor1 6rop6oou proq 0qtrroq RNA.
* AvnrepUortopdq rnq peroypoqrlg oulrpo[veL en[or1g roL
oro oneprivro rou rRNA.
0 Kuxtrog rou Borrqplorou Ayyetrrorprjpou RNA
* Iro porrrlpto, n UeroypoQn ror 11 ;leroqpoon npoyU0-
ronorouvtor rouroXpovo (ou(euyU6vn Ueroyporpl/Uerc-
Qpoon), ro069 ro p$ootoporLo [errvouv q perogpoor
evoq mRNA npLv otrortrr1pol0e[ 1 ouv0eofl rou.
* To poKTrlproKo mRNA e(vor ooro069 Ue Xpovo rlpr(orqc
trQov trenrtiv qq topog.
s 'Evo
BoKTnproro mRNA pnope( vo e[vot notruotorpovtro
ge notrtrrlg rordrreg neproXdg nou ovTrororxouv oe 6ro-
qoperLro yovi6ro.
nepitrnUn
ffi
ag.i*,#,*
5! L PJ't
,fiFts{
t:t &:!( l
xolefuor xtrrilvog p4-expoyeio (non-template strand),
evri: ow6g nou l,enoupye( og er<proye(o yro rrl o[v0e-
on rou RNA rq}'ekor rLctrvog exproyeio, 6no:g tpofue-
ror ornv EIKONA 19.1.
H oUv0eoq tou RNA rotoLUeror on6 ro 6v(upto R-
NA notruyep6o4 (RNA polymerase). H peroypocprl [e-
rtvd 6rov q RNA noluprepdor; npoo6e0e( orov
unoxrvqr{ (promoter), puo'er6rrcl neproXrl, orrlv opXtd
rou yovr6(ou. O unorrvqr4g nepil.oprBdver to npd:ro
(erlyog Bdoecov nou preroypdgeror oe RNA (ogerqpio
- startpoint), roOrig Kor rrq neprBdl.trouoeq Bdoerg.
An6 ourj q 06oq, r1 RNA noluprepdo4 perorrve(tor xo-
rd pqrog rou erpoye(ou, ouv05rowog RNA prdXpr vo
qrdoer oe Uro otr).4},ou1(o repporror{ (terminator),
6nou retrerc0ver ro perdyporpo. 'Erot ;uo peroypogrxri
pov66o (transcription unit) erre(verqr qn6 rov unort-
512 KEOA,A|O 19 Meroyporprl oroug l-lpor<oputirreg
Boorxeg cworeg

K),6voq un-eKuoys[o
5' TAC * CG ET,qC G GTCAATG CAICTAC CT
3' .l';':;,:j !:.: i1:t-i:ii:l i.; lt & I'jp.'i-l G
jrt +,,$*i * A
vrlrri 6oq rov repUorrmrl. To Ko(pro Xoporrqptortx6
rnq Ueroypo<pu<qq provddoq, 6noq <po(verot orqv ElK0-
NA 1S.2, e[vor 6rt onotel,e[ 6vo rprrlpro DNA nou Xpnorpo-
noe(tor og expoye(o yro rnv nopoycoyrl ev6g provo6trof
prop(ou RNA. Mro peroypogrxrl provd6o pnope( vo rto6t-
ronoe( neptoo5rcpo on6 5vq yov(6ro.
Or oMrll,oufeq nou Bp(orowor npw on6 rnv oee-
rqp(o ovogdpowor ovo6rx6g (upstream), evdr outdg
nou Bp(or<owor perd qv ogeqp(o (g5oo orq Ueroypo-
q6pevq ol,r1LouXio) rog xo0o6rx5g (downstream),
Iuyyd 4 oMrlloul(o tou DNA ypdeeror 5rot c0ore
vo <po(verot pr6vo o rd.cbvog prrl-erproyeio, o ono(og
(6no:g npoovo<p6p0r1re) 51et rrlv (6to oMrlloul(o pe ro
RNA. Or O5oerg rov Bdoerrlv optOpo0wot Kot npog rtq
6fo roteu0Uvoerg on6 qv ogerrlp[o, nou ovopd(etot
+ 1, or de oprOpro( ou[dvouv npoq rnv r<o0o6u<q <popd.
H Bdoq Trprv rrlv oqeqp(o oprOprefuot -1 Kor ot opvnrt-
r<o[ oprOgo[ oufdvouv nrlyo(vowog ovo6trd. (Kop(o Bd-
oq 6ev 61et rov opr0;r6 0).
To opXrr6 npoi6v rnq Ueroypoqrlg nepil.o1-tBdvet to
ou0ewrr6 5' drpo ror ovoprd(etqt rporoyev6g per6-
ypqeo (primary transcript). To npcoroyev4 pretdypogo
rou rRNA Kqr rou IRNA utp(orowqr pto 6epyoo(o top(-
Uovonq rord rqv ono(o otrLr1trouX(eg'rd:v drpov toug
r6Bowor on6 ev6ovourd.edoeg. To cirptpro npoi6wo trrrv
-35-10-1 +1 +10
--.+
Avo0rro KoOoDLr<o
IVleroyporprrq povodo elvor 1uo otrtr1trouXio DNA
nou peroypoqerol o' evo evrolo RNA, [extvtlvrog ono rov unort-
vrlq r<or retreulvovrog orov repportor1.
wNw{s=Ktrtirvoq erpoyaio
METATPAOH
RNA getoypotpo
ElK0liA 19.1 AeLroupyro qg RNA notrupepoorlg e[vor v' ovlypoqet 6vov r],6v0 rou d(r],orvou DNA oe RNA.
H otrtrr1trou1ro rou RNA ervot I
oupnlqpupattr4 ge rov xtrtrlvo tergoyeLo xat raur 604 p4 ye
rov <otdtronotrlrtxo rrtrovo
= 5'UACGCGGUACGGUCAAUGCAUCUACCU
onepov(rov rRNA r<or IRNA e(vot moOepd, trre 1p6vo
(or1q nop6poro Ue ro Xp6vo yevtdq tolv Borcrqp(ov. Av-
r(Oero, ro npcoroyevrl getdypogo rou mRNA 66Xowot
oXe66v dpreo4 en(Oeor1 on6 ev6ovouxledoeg r<t e[u:-
voux.).edoeg. 'Eror, o prSoog Xp6voq (onq tou Borcrqpto-
r<of mRNA e(vor pr6vo 1-3 Lemd. Iroug europutiteg, to
pretdyporpo rRNA ror IRNA utp(orowot Korepyoo(o t<ot
ro npoi6wo nou npor[mouv e(vot oroOepd, 6nog mo
Borrqpro, oLl.d to mRNA e[vqt noLti nto oroOepd on'
6,r oro Borcrrlpro. Ot rpononoqoetq Kot 11 ono66ptqoq
rov mRNA 0' ovoLu0oriv ctrlv Ev6trya 19.23.
H preroypogq e(vor ro np6to ord6to orq Srgpooq
rcov yovrd(rov xor ro Brlpo sro ono(o oorefuot nto ou;yd
p(Opuoq. Or puO;umtxo( nopdyoweg ouprd r<o0op((ouv
ov 6vo ouyxer<prpr6vo yov(6ro 0o pretoypo<pe( pSoo qg
RNA noLuprepdonq. En(oqg elSyXouv rot petoyevSore-
po mddro rnq Ueroypogqq r<ot dMo mddto qq 6r<-
qpoonq ro:v yovr6(olv.
Ario orlprowu<6 epor(proro oletrd Ue rn Ueroypo-
qq efuor:
o t-lcirS Bp[or<er q RNA no].uprepdorl unortvrlt6g mo
DNA; Aw5 eivor pr6poq ev6g yevu<6tepou epotrlpo-
rog: l-lig (eXcop((ouv or nporefveg ng ouyrerptpr6-
veg 06oerg np6odeoqg mo DNA on6 ttg dl,l,eg
oMqLouX(eq;
o flcirS oLLrllentdporiv or puOprorrr5g npoteiVeg trle
rqv RNA noluprepdo4 (r<or pero(U toug) rbme vo
evepyonorrloouv 11 v' qvome(trouv ouyrerptpt5vo
Brigoro rotd rrlv Svop(r1, r4v entprlruvorl ri tov
rcpprotropr5 qq uerqypoeriq ;
Ie our6 ro reqdtroro 0o neprypdtloupe rtq otr4Le-
nr6pdoerg qg Borcrqpror{g RNA no}.ugepdoqq Ue ro
DNA, on6 qv oplu<ri enogrl rrlq pe tov unoxtwltti, r1
Ueroypoen Kor rnv oneleu06ptoon qS on6 to DNA p6-
l,rg oloxl4pto0e( ro pretdypogo. Ito KeEdl.oto 20, 0'
ovol0oouge tg ow(motleg owr6pdoetg prero[U rov
europutoru<c0v RNA notruprepootirv Kor rov ergoye(<ov
roug. To Kerpdl,oro 26 nopouotd(et dtdtpopouq rp5-
nouq Ue toug ono(oug or pu0ptmtrSg npolteiVeq Kot ot
nopdyoweg prnope( vo po4Orloouv ti vo nopepnod!
oouv rn Bonrqpror<q RNA notruprepdorl vo getoypdtlet
6vo ouyrerprpr6vo yov(6ro. Iro Ke<pdloro 27 e[erd(ou-
9.1 Erooytoyq 513

19.2 H Meroypogri
Zeuy6polpo Bdoeorv
A(euydporou DNA
llpoylroronoreiror pe
o€ Uro (Ouoo^i6o,)
pe nrilq or [elotpror6g pu0;uorrr5g owrdpdoerg gnope(
vo ouvdud(owor oe nol,unLor6repo 6(rcruo. To Kegd-
Loro 30 ovoL0er np6o0eroug rp6noug prlOpuoqg, pero-
[ti dMtov n Xpnorporofiorl gu<pc0v RNA, r<or qvo].Uer
nrilg ourdg or oMqlenr6pdoerg prnope[ vo ouvddowor
oe peyolUrepo 6(rcuo.
roypogrx{ quoo}.[6o", mrlv ono(o ro DNA nopo6rxd
6toXrr:p(eror orouq 6Uo provo0g drc0voug rou Kor o
xi.cbvog extrroye[o Xpnorpono€(rot yro vo xqreuO0ver
q o0vOeorlrou xl.drvou rou RNA.
H oLuoGo rou RNA ouw(0erot on6 to 5, npog ro 3,
dxpo pre npoo0qr<q v6ov voux.l.eon6(ov oro 3' drpo rqg
ou(ov5;revrlg oi.uo6og. H 3'-OH opd6o tou reLeuro(ou
vourlsort6(ou nou npoor60rlre or4v ol,uo.(6o owrbpd
pr' 6vo eroepl6;revo 5'rpupoorpopx6 vourLeoo(6ro. To
etoepX6prevo voux.l,eoo[6ro Xdver ng 6Uo re].u<6g gco-
o<popu<6q oprd6eg (y r<or B), evr6 4 opd6o o Xpnoqonor-
efuot oro goorpo6eoreprr6 6eopr6 nou ro ouvdder arnv
qluo(6o. H roXUr4ro qg ouvoLx4g ow(6pooqg yro rr1
Borcrqpror<f RNA noLuprepdorl prnope( e(vor dog -40-50
vouxLeor6ro/6eurep6l,emo orouq gToC yro ro nepro-
o6repo prerdypogo, nep(nou (oog pe qv to1tirqro pe-
rdgpooqg (15 opwo[do/sec), oMd no].ri ;ur<p6repq on6
rrlv rolrir4to owrypoqqg rou DNA (-800 bp/sec).
H RNA noLugepdorl 6qpuoupye( rr1 ;reroypogrxrl
<puooL6o pr6).rg npoo6e0e( o' dvov unorrwlrn. H ElK0l{A
19.4 nopouord(er qv RNA noLu;repdoq vo perorrvefuor
r<ord prqrog rou DNA, pre q guool(6o vo peroxrve(ror
po(( qg Kor rnv o).uo6o rou RNA v' ou[dva oe pulrog.
H 6repyoo(o rou (euyopcilUoroq rov Bdoerov Kor rrlq
npoo0{r<r1g Bdoecov ;r5oo orq <puooA(6o roro.)Jeror xt
enr4pefuor on6 r4v (6ro ulv RNA noi,uprepdo4.
H 6opr1rrlg quooL(doq 96oo oro oritrrnLoro rnq Ue-
royporp4g go(veror ar4 Aemopepn dnorln qg E|K0NAI
19.5. Ko0rig q RNA noLuprepdor; perorrvefuor r<ord prj-
Koq rou DNA erpoye(ou, ro 6(r),ovo (erultyetor oro
pnpoorrv6 pSpoq qg guool,(6og (onUe(o [erutriyprorog
e H RNA notrupepoorl droXropr(er roug duo rtrdrvoug rou DNA
oe pro eqrlgepl ,,rpu00L[60" Kor xprlouon0re( rov 6vo r],tirvo
org ergoye(o yro vo roreu0uver rr1 ouvOeo4 prog oupntr4prrt-
porxrlg otrtr1trouXLoq RNA.
* To Unroq qg quootrrdoq e(vor'12-14 bp evdr rou uppr6(ou R-
NA-DNA p6oo orq quootrOo 8-9 bp.
H peroypocprl Xprlor;ronore( q ouprnL4prollolx6rqro
ro-lv Bdoecov, 6ncog xor or dMeg owr6pdoetg noLuge-
prol.rori, dqLodq 11 owrypogq xoL 4 ;rerdgpoorl. H
.,',, ': ,', . ono6(6er q yevrKr1 opxn rnq ;retoypog4g. H
o0v0eo4 tou RNA npoyporonore(ror pr6oo oe Uro (Ue-
Kodtrdg x.tirvog
Ktr6voq eKuoye(o
0r rtrtirvor rou DNA dtoXrop((ovror yro vo oX4porLoouv
Uro UeToypoQrKrl rpuootrido. To RNA ouvrlOeror pe oupntrrlpotporxo
(euyoputlro rtov pooeov pe rov evo ono roug r),(rvoug rou DNA.
514 KEOA Al0 1g MeToypoQ1 oroug l"lporoputirreg
w;.@,TWrx"=
ffifl.M-ffix ''r*er'ffi"x-ffi
EIK0NA 19.4 H geroyporpq npoyporonorefror gdoo oe Uro quoq-
tri60, orqv onoio RNA ouw(0eror pe (euydptopo rov pdoetov 1r,
6vuv on6 roug rtr6voug rou DNA oqv nprioropo [erutrryp6vrl
neproXrl. Ko0tig perouve(ror q quootrbo, ro D(xtrrr:vo DNA [ovo-
oXrlp0ri(eror n(oo u1g, exron[(ovrog ro RNA un6 popqrl povrlg
nol,uvouxl,eo16xrl g otruo6og.
To DNA onodroTdoor.o,
I
H oUvgeo4 Tou RNA
[euvd mr1 quoo],6q
Boorxdg evvoreg
J
+ 0uootri6o peroypoqftg +,

Merorw4oq rou ev(Upou
'+
05oq npdo6eolg rou RNA
EIK0NA 19.5 Kord q Ueroyporprl, I rpuootri0o Drorrlpeiror p6oo
oq porqproxrl RNA no)upepdol, rou [erutrfyer xor [ovoru)ryer
ro DNA rot ouv06rer RNA.
- unwinding point) ror ro DNA enqvefurd)veror oro n(-
oro rprlpro (o4pe(o (ovorullyprmoq - rewinding point),
To pqrog rrlg <puool,[6og e(vor -12-14 bp, evcir ro ;uixog
tou uBpt6(ou RNA-DNA pr6oo on1 rpuooLbo pr6llq 8-9
bp. Ko0rig ro 6v(u;ro petorwe(ror r<ord prrlxog rou eK-
poye(ou, [ovoq4qpor((eror ro 6[r],rovo DNA ror ro RNA
erron((eror o g eLeri 0ep4 nol,uvo u x.leorr6rnj ol,u o(6o.
Ie rdOe 6e6op5vq onypf , ro rel,euro(o 14 prBovouxi.e-
or(6to rou ou[ov6pevou RNA Bp(orowor oe oUpntror<o
pe to DNA xo/r1 ro 6v(upro.
19.3 H Avri6poon Tnq McToypoefiC'Ex€l
Tplo lrd6to
" H RNA notrupepoorl npoo06veror oe pro 0eorl unoKrvrlTrl oro
DNA yro vo oXqporloer 6vo r],eroro oupntroro.
" H RNA notrupepoorl [exrvo q Ueroyporp1 orpou ovo[[et ro di
rtrrr:vo rou DNA yro vo oXqporloer Uro UeToypoqrKrl rpuootrido.
* Koro r1v enrpfruvor1, I Ueroypoqtrl rpuootrrdo perortvelror
roro pr;roq rou DNA ror 1 otruo[do rou RNA entplruveror oe
roreuOuvorl 5'+3' pe npoo0rlrrl vourtreortdlrrrv oro 3' orpo.
* H Ueroypoql oroporo rot ro dlrtrtovo rou DNA [ovooXlporr-
(erot gotrrg 1 RNA notrupepoorl onoouvdeOel ono ro DNA er-
poyelo oe Uro eeon repportolrou.
H ow(dpoon rnq Ueroypognq drorp(veror oe rp(o
orddro nou cpo(vowor ornv i', ' : (1) 6vop[n
(initiation), orrlv ono(o ovoyvrop((eror o unoxrvrltrlg,
6q;rroupyekor Uro guoo).[6o r<or (erLvd 4 oJv0eorl
rou RNA, (2) enrp{xuvon (elongation), orrlv ono[o
11 guool(6o ;."tetoxrve(ror r<otd pr1<og rou DNA, r<or
(3) repponop6g (termination), xord tov ono(o to
RNA perdypo<po oneleu0epcbverou<or r<).e(ver q rpu-
oo).(6o.
H (6to q 5vop[r1 prnope( vo 6Lor<pr0e[oe noMd Bqpro-
to. H ovayvdtpon rou expoye[ou lexwd pe u1v npoo6e-
ENAP:H
Avoyvtirptorl rou expoye(ou: 11 RNA noLugepdol npoo0everoi
oro 6[x],rovo DNA
To DNA {erutrryerot orov unortvrlq
llotrrj ppoXe(q qtruo6eq ouw[0ewqt Kr oflelsueep(tvovror
EflIMHKYNIH: 1
H notrupepdorl ouv06ter RNA
TEPMATIIMOI:
H RNA notrugepdon Kqr ro RNA onetreu0epdrvovror
ElKOllA 19.6 H peroyporprj 6xer rp(o md0rq: To 6v(upo npoo66-
veTor oTov unouvrlr4, onodroroooer To DNA xor nopop6ver or(vr1-
ro xord qv 5vop(1: Merouve(ror rord plrog Tou eKpoye(ou Kqrd
r4v enrp4r<uvq rqr 6r(ororor orov reppouogd.
oq rnq RNA nohupepdo1g oto Sklavo DNA oe pn aA-
l4louy[a DNA nou xaleftcl. unoxlrpr1g @romote!. To
5v(upro oXrlprot((er np6ro 5vq xtreror6 o0prtroKo
(closed complex) oro ono(o ro DNA nopopSvet 6(r<Lro-
vo. 'Enero, (eruLfuer ronrd ro tprrl;ro DNA'rou unorr-
vqrrl nou neptloprBdver rn eSon Svop(r1g r4g
peroypogqg yro vo q4qpot(oer ro ovol;6r6 oripnAoxo
(open complex). O droXroptog6g rou 6u<Lcirvou DNA ro-
Ormd rov r<i.c0vo-er<proye(o 6ro0Sorpro yro (euydpcopo
Bdoecov pre eroepX6trrevo prBovoux).eor(6ro xor oriv0eoq
-rcov
npcbrrov vour.treon6rrc0v 6eoprdlv oro RNA. H qdon
ouq prnope( vo nopo'ro0e( qn6 qvenrule[g on6nerpeg
Svop[4q, orrg ono(eg ro 6v(upro qndprer BpoX6o pret6-
ypoqo, runxd rdro rulv -10 vouxleond(cov (nt), evcir e(-
vot or<6;rq npoo6e6epr6vo orov unorrvrlri. To 5v(upo
ou;yd r<dver 6ro6oXxorig 66xtroug ovenrulcirv Uerqypo-
grilv r<or lovopX((er q o[v0eoq rou RNA. H <pdo4 rqg
Svop[r1g re^.er6vet 6rov ro 6v(upro retrrd r<oropOcOoer v'
enercrefuer qv ol,uo[6o Kor vo onopor<puv0e( on6 tov
unorlqrrl.
Krivoq erpoye(o DNA
lii .: H AvTiDpooq rrlg Meroyporplq'EyerTpLo Iro0ro 515
Boorxeg eworeg

H enLp-rrp<uvoq ouv(ororor oe npoo6eurrrrl preror<fvq-
or'l rou ev(0pou trre 6tordpo(r1 tou (euyopri-rprorog rov
Bdoeov oro d(x.tro.rvo DNA, ogrlvowog erreOerpr5vo rov
r<Lr.rrvo-expoyeio yro npooOrl<q voux).eorL6(rov, Kor Ue-
tot6nroq rnq peroyporpLxqg guooLf6og nro ro0o6rxd.
Ko0rrrg perorrve[tor ro 6v(upro, o x],covog-exproye[o qg
npoorrrpLvd (etuALyprdvlg neproXrlg (euyoprrtver pre ro
veoouwe0eLpdvo RNA crro o.r;pe(o qg o0(r1oqg. Nou-
r).eori6ro npom[Oewor ol.rorono].Lr<d oro 3' dr<po rrlg
oulovdpevrlg oLuo(6og RNA, o'Xqpor((owog 6vo uBpG
5Lo RNA-DNA ;r5o'o mqv lerulryprdvrl neptoXrl. l-1(oot
on6 rrlv [erufuypr6vr1 nepropl, o r<].ovog-er<poye(o rou
DNA (euyoprrrveL pe -rov ouOeyrrrd r<),rrivo "ro[pr" rou,
yto vo lovooXripor(oer rq 6rn).r1 dLrr<o, evctt o ou(ovdpe-
vog N).rrlvog tou RNA [enpoBdtr].eL ond ro dv(upo.
H nopo6ooLorr; on6r,l.rr1 6l r1 enr;rrp<uvor1 e(vor ;rovo-
rovq 6tepyoo(o, o-rqv ono(o -ro 6v(up_ro npocu0e(ror r<otd
pqrog rou DNA pe oroOeprl rolirqro nou owrmotle(
ornv rrpooenKn rcuv voux).eorr6(olv, dXeL oMdler ro re-
Leuro[o XpdvLo. H RNA no].uprepdorl npoyporonore(
nor-ioq 11 ordpq ror oropord oe ouyKeKprpdveg oMrl-
trouX(eg. H em6nrorl rou 3' dxpou rou RNA ond ro
evepy6 r<6,wpo trrnopei vo nporol6oer "ofiroeox6prlor)"
rrlg noLupepdoqg r<oL ogo(peoq ;_reprr<rr.lv vour<LeorL-
6(ov on6 trlv cu(ov6prevq o).uo[6o rou RNA fiprv rr]v
enoverr[v4oq.
O rep ;.rottopr6g nepiloprBdvet ovoyvcirptor; oMrlLou -
Xrorv nou 6(vouv mo 5v(upio ro orlpro vo oropotrloeL
r4v nepondpot npooOqr<r1 vour<).eol6iov ornv o).uo(6o
rou RNA. H guo'o}.(6o xoroppdel<oOriq ro uBp(6Lo R-
NA-DNA onodroroooeror Kor ro DNA lovobq;rtoupye(
6vo E(xLrr:vo, 11 611 puoupy(o goo<podLeoreprotv 6eo pur-rv
dLoxdmeror rar ro orjpn).oKo Ueroypogqg 6tolrrip((etor
oro ouorolKd on6 ro ono(o onopr[(eror: RNA no].upe-
pdoq, DNA r<oi DNA ;reroypo<po. H aAA7Aouyfu rou D-
NA nou xareu9lva r ArjEn rnq psraypaErlq ovolto(erat
reppau.orq.
'l 9.4 H Borrnplorf RNA lloAuUepdon
AnoreAeirot on6 noAAdq Ynopovd6eq
". 0r nuprlveq rov Borqproxtrrv RNA notrupepoorirv oivor notru-
pepfl oupntroxo'400 kD pe yevLrrl 00prl 0208'0r.
., H rorotruorl enustrefrot ono rLg unopovodeq p ror B'.
OL r<o).rjtepo XopqKrnprotl6veg yevettrd xor BtoXqpru<d
RNA no).upepdo'eq npo6pXowor on6 Bor<rr1pro, t6(<i.rg
on6 ro Escherichia coli. Oo16oo, oL p6veq BorcrrlpLo-
r<59 RNA nolupepdoeq yro rtq ono(eg 51ouv npoo6ro-
pLore( uUnlrlS ovd).uo11g rpuoro).Lrr<6g 6op6g,
npodplowoL on6 60o Oeppr6gil.o Borcrqptor<d e[d4, ro
Thermus aquaticus xat Thermus thermophilus. Ie 6o
ro Borrrlpro, p6vo 6vog r0nog RNA noAugepdorlg ei-
vor uneU0uvoq yrq rq orivOeoq rrov rRNA, mRNA xor
IRNA, qw[0ero on6 toug europucirreg, srouq ono(oug
ro rRNA, mRNA rqt IRNA runxd preroypdtpowor on6
6rogoperu<59 RNA noluprepdoeg, ng Pol l, ll xot lll.
l-lepfuou 13,000 g6pro RNA noluprepdorlg undplouv
oro rUrropo rou E. col/, nop6rr o orprB4g opr0;rdg
norrf],er ovd],oyo Ue trq ouv0rlr<eg oU[r1oqg. Irqv
npoyllorrr6rnro, oe rd0e 6e6opr5v4 orryprrl 6ev ep-
nA6Kowqr 6).eg or RNA noLuprepdoeg orq psroypo-
<prl, 6l,eq 6prog e(vor npoodedepdveq oro DNA, e(re
et6rrd e[re ;rr1 er6rrd.
To n)r1peg 6v(u;ro 11 otro6v(upo (holoenzyme) rou
E. coli 51er propror<6 Fdpoq -460 kD. To o).odv(upro
(qzpp'oo) pnope( vo droXroptme( oe 6Uo oumotrxd,
rov nupr;vo rou ev(riprou (ozpp'to) Kor rov nopdyowo
o(ypo (notrunenr(dro o), nou epnL6reror er6rxd orqv
ovoyvcilprorl rou unoKrvrlrrl. H oUorooq rou oe unoUo-
vd6eg ouvor!((erot orrlv EIK0NA 19.7. Ot uno;rovddeg B
ror B' go(( rpoyUoronoroUv r4v r<ordl,uoq rou RNA ror
qnore^,o0v to preyol,rirepo rpulpo r4g pd(og rou ev(0-
;rou. Or oprLvo[rrdg o].].r1].ouX(eg rot ot rpro6tdmoreg
6op6q rouq e(vot ouwqpnp6veq, x6tt nou rqxuer yro nq
peydleg unopov66eg rtov RNA no).upepoociv r<or on6
rtq rperq enrxpdreeg rqg (or1g: Bor,rrlpto, oplo(o xr eu-
t<opuc0reg (B).6ne Ev6rrya 20.2). Auto uno64l,cbver 6rr
or Boorr5g t6r6rrpeg rnq Ueroypogrlg e(vor roweg prero-
[ti rtov no].upeprirv RNA no].ugepood:v 6].cov rrov opyo-
vrogd;v. Or B xor 9' Uo(( oXq;ror((ouv ro evepy6 r5wpo
tou ev(6prou, rov r<ripro 6(oulo 6ropr5oou rou ono(ou
6rSpXeror ro DNA xo-rd tov r<0xi.o rqg preroypogrlg, ro
6ewepe6owo 6(ou).o 6ro;r6oou rou ono[ou ro prBo-
vouxi.eo{6ro rou unoorpriporog ero6plowor oro 6v(u-
[ov[6to llpoiriv Aenoupy(eq
rpoB B unopovddo
(r51 kD)
rpoA 2 o unogovd6eq
ftA Iuvopgotrdyqorl Tou ev(ugou
;; ^; i'r ql7 ovoyvopron rou unoKrvrlnl
xoOeg(o) npooddver rdnotouq
1|i'-onoqrdq
l- Kurotrurtxd x6wpo
glrpoC p' unopovddo
(1 55 kD)
o unopovo6o
(1 8-70 kD)
Etdtxorrpo rou unorrvrlrrl
or unopovd6o
(r 0 kD)
'Ev(ugo rou
E.coli =460 kD
0t RNA notrupepooeg rov euporqpiov eXouv ne.-
re tunoug unopovo0ov: oL o, p, p' ror o eXouv potrtrov oroOepc
U6ye0oq oro dlorpopetro erDrl porrrlp(ov, evtir 11 o norrltrer ne -
prooorepo.
rpoD
rpoZ
516 KEOAAAI0 19 Meroyporprl oroug llpoxopurireq
Booxdq 6woreg

uo orrlv nope(o rouq npoq ro evepY6 KSwpo Kot ro 6[-
ou^o e[66ou 6ropr6oou tou ono(ou to veoouweOetp6vo
RNA eyrorotre(net ro 5v(upro. Ie oup<pcov(o pre ro p6Lo
ourrilv ttov unopovddcov oe 6l.eq ow5q ng Lettoupy(eg,
prerotrtr6(erq oro yov(5to rpoB xot rpoC nou xtodronot-
oriv lg unoprovd6eg B ror B' en4ped(ouv 6tro ro ord6to
rqg pretoypogqq.
To 6tprepdg nou oXrlproil(etot on6 ng 6Uo q unopo-
v66eq Xp4orpre0er toq u<p(cr:pro Yto rrl ouvopprotr6yrlo4
rou KewprKo6 ev(0pou. H C-retur{ neplolli (C'termi-
nal domain - CTD) rov o unopovd6tov en(oqq 6pXetot
dpreoo o' enog( ;re to DNA rou unoKtvrlrrl rt Stot oup-
BdMer orrlv ovoyvcilptorl rou unortvqr{ (B}.5ne nopo-
rdrco). EntnL6ov, ot unoprovd6eg o Kot o e(vot ot rUpteg
enrgdveteg qq RNA nol.uprepdorlg yt' oAtrrlLenrdpdoetq
pe nopdyoweq nou pu0pr((ouv qv 6vop[q rrlq Uero-
ypoqnq. H unoprovddo to en(or1g no(et p6lo orrl ou-
voppol6yrlorl rou ev(6pou, 6nog en(or1q Kot oe
ouyr<erpq.rdveg pu0ptorrxSg ettoupy(eg.
H unopovdEo o e(vot rotd r0pto tr6yo uneU0uvrl yo
rrlv ovoyvdrpton rou unortvrltrl. H xpuorotrtrtr<{ 6opf
rou nup4vo rou Bor<rqptororj ev(rlprou 6eiXveL 6tt 51et
5vo oXrlgo oo 6oyr<dvo xoBouptorl, Ue rn Uto 6oyrdvo
vo oXq;roil(etor tcup(cog on6 rq B uno;rovd6o Kot rnv
dtrl.q rup(rog on5 q B' unoprovd6o, 6nog go(verot ot4v
Ell(0NA'1S.8. O npoteUorv Dfoutroq yto to DNA Bp(oretot
mqv enrgdveto eno<pqg rov unogovd6tov B t<ot B', ot
ono(eg oroOeponoto0v tou g 6toltoptoprdvou g, provoti g
r<).cirvouq orrl Ueroypo<prxrl quool(6o, 6nog rpo(vetot
orrlv EIK0NA 19.9.
H 05or1 rotd).uor1g Bp(oxerot orq Bdorlrqq q4totrrqg
nou oXrlptot((etot on6 rq (ooY6vto" roJV 0 rot B'.
'Evo
H ovodrrl onogl rou nuprlvo qg RNA notrupepo-
o1g, onou qo[verot ro oX11.to ,,doYrovdtv ro$ouptou, Tou ev(u-
Uou. 0r 0 (ruovo) rot p' (poQ unopovo0eq i1q RNA notrupepoonq
eXouv evov 6[ou],0 yto ro DNA erpoyeto. H o I tpolverot npootvl, q
o ll ruprvrl rol 1 o xornvl. tlpooopltoopdvo ono. K. M. Geszvain
ror R. Landick (ed. N P Higgins). The Bacterial Chromosome,
American Society for Microbiology, 2004
, t, H 6opr1 rou nuprlvo rrlg RNA no),upepoolg ono ro
porrlpro Thermus aquaticus, Ue U g unopovoDo pntre rot q $'
npootvq. Aopq ono: Protein Data Bank 1HOM L. Minakhin. et ai,,
Proc. Natl. Acad. Sci. US/ 98 (2001):892-897.
on6 to 6rjo roroluttrd t6wo Mg2* nou e(voL crnopofurl-
ro yrcl ro prllovLopr6 rrlg rord).uoqg e(vot ogrXro npoo-
6edepr6vo oto evepy6 Kevrpo rou ev([pou (B].6ne
Evoulta 19.18). To dMo L6v Mg2- tptovet crro evepyd
x6wpo oe oUpnLot<o pe ro etoepx6prevo tpt<prooqopLxd
vour.treori6to (NTP). Onoq npoovotp6pOqr<e, o nupqvog
rou ev(rjpou rov tuKopuorov dlet qv (6to pootr<q 6o-
prrl pe ro Borcr4ptor<6 dv(upo (Erx6vo 20.2). En(o1g ne-
pr51er r<onoteg entnp6o0eteg unoprovo6eg KClt
nepLLoprBdver oMrl).ou1(eg nou 6ev undpXouv oro Bo-
rcrrlpLor<6 6v(upo. Or r<ripteg 6Lotpop6g ovdi.-ieoo mo Bo-
r<rrlproxd Kor ro euKopuortKo dv(upo Bpior<ovtoL
oXe66v onor).etorLro mqv nept<pdpeto rou ev(0;rou,
uoKpro ono ro evepyo Ktvrpo.
'l
9.5 To OAo6vCuUo
AnoreAeiror on6 Tov
Tov nopdyovTo o
Tnq RNA l-loAuuepdonq
Iluprivo Tou Ev(tlpou Kot
a H Borrlptorl RNA nolupepooq unodtotpe[toL orov 02BB'o
nuprqvo rou ev(upou nou roro),0et rl1 UeroYpotp1 KoL Tl'lv uno-
povodo o nou onouoltot ltovo yto rr1v 6vop[q.
* 0 nopoyovro-Q o rpononote[ rtq tbtorlreg npoodeorlq oro D-
NA qg RNA notrupepoorlg, ersL 6ore q ouyy6veto Tlq rpoq
ro yevlr6 DNA vo ptet6verot xot, ovr[0ero, 1 ouyyeveto qg
npoq Touq unouvrlrdg v' ou[ovel.
O nuprlvog rou ev(tlptou 61et yevtr<q ouyy6veto Yto ro
D NA, xu p(r.og ).6ytr-l rllercrpoorotLxrrtv oMqenL6pdoe-
oiv 1.rero[[ tqg nporelvng, nou e(voL Bootr<r1, Kot rou
DNA nou e(vot 6(tvo. Otov npooddverot mo DNA
ror' our6 rov rp6no, to DNA nopopevet oe 6(r<trovq
lfr.li To Otrodv(uUo qq RNA tlotruUepoonq Anoretreiror ono rov l-luprlvo rou Ev(upou roirov I-lopoyovro o 5'17
Boorxeg eworeg

/ i r::,1
, .T -"'
I
llopoyovrog o
0 nupqvog rou ev(ugou npoodeveTot odtorplrog
oe roOe DNA. 0 nopoVovrog o perdlvet rr1 ouyy6veto yto npoode-
oq ove[opqrq ono rqv otrtr1trouXio ror npoo6[6et eldtroqro yto
roug unorrvlreg,
Uopqn. O nup1vaq tou ev(l1tou 6yeLqv xav6trya va
ouv06teL RNA an6 6va DNA expayeb, oMd 5e pnopel
v' av ayva p [(er uno xwryt 6q.
H Uopqri rou ev(ripou nou e(vqr uneri0uvq yro rnv
6vop[q rnq Ueroyporprjg on6 rouq unoKrvqrdq ovoprd-
(eror ol.odv(upo (o2BB'roo) (EIK0llA 19.10). Arorp6per on6
rov nup{vo rou sv(rlprou enetdri nepdXer dvov nopd-
yowo o. O napayowag o$pa 6yt 1t6vo efaoEah((et 6tt
q Baxtqpnxi RNA nolupepdorl evepyonoe[ rn pera-
ypaqi an6 optopdveg 0ioerq, aMa en[or1q peuitvet xat
qv np1odeory oe pr1 edxdq a&rltouy[eq. Xop(g nopd-
yowo o, 11 oro0epd ouyydverog yro rrlv np6o6eor1 tou
nupfvo rou ev(Uprou oro DNA per6verot rord -104 rpo-
p5q r<or o 1p6voq npr(onq rou ougnl,6rou e(vot <1
6ewep6Lenro. Aw(0ero, ro otro6v(upro npoo66vetot
fiouq unoKrvrlrSg noLrj nro orptXrd, pre oroOepd o'uyy6-
verog xotd pr6oo 6po -1000 gop6q UeyoUropn Kor
Xp6vo ngr(onq 6o9 r<or oprer6q cirpeg. 'Etor o nopd-
yowoq o ouorcrorrKd onomo0eponore( q pq etStrrl
np6odeoq rou unorwrltj.
H roX0rrlro pe rnv ono(o ro oLo6v(u;ro npoo66ve-
ror oe 6rorpoperu<59 oMrlLouX(eg unorrvqrc0v nopou-
ord(er peydl,eg 6ror<updvoetg ror yr'our6 onore)re[
orlprowrr<4 nopd;rerpo arov ro0optop6 rnq "oy0oq rcu
unoxwt1r4", drllodTl rqg onotetreoponx6rqrog ev6g (e-
Xoptoro0 unorwrlq oq npoq qv 6vop[4 rnq Usroypo-
qnq. H ouXy6rrlro rqg Svop(qg nox(trer on6 -1lsec yo
ro yov6ro rou rRNA un6 rSovrx6g ouv0{reg 6o9 < 1/30
min yro rdnoroug dLLoug unoxrqt6g. O nopdyowog o
ouv40cog onetreuOepc0verot 6rov 11 otruo(6o rou RNA
gtdoet oe ;rr1<og -10 nt, oEtivowog"rov nupfvo tou
ev(Uprou une0Ouvo yro rrlv entpr{ruvot1.
0 o onomo0eponore( q gr1 el0rKn npdo000n
I 9.6 I'ltilg Bpiorer n RNA lloAuuepdon rtq
AAAnAouxiaq Tov Ynomvnrdrv;
a H npoodeo! rrlq RNA notrupepoolg oTOUS ulloKtvrlTeg eivc.
rooo ypnyopn tiore gotrlov 6ev nporeuot yt ontrl OtoXuo1
," H RNA notrupepool npooDevetot oe ruXo(eg Odoetg oro DN"
xor trq ovrotrtrooer pe otrtreg otrtrr1trouXteg 6oq orou Bpe0s.
evog unorrvlrrlg.
H RNA nol.uprepdoq np6ner vo Bp(oret unortvqt6g ;.re-
oo oro nLo(oto tou yovt6uol-rorog. I-1cog [eXorp((ouv c.
unorrvrlrdg on6 trg 4 x 106 bp nou onopt((ouv rc
un6Lorno pSpoq tou yovtdrcbporoq rou E. coli', N
nopouotd(er ontrd prow6Lo yro to ntrrg q RNA
nouprepdorl 0o pnoporioe vo Bp(oret rtq oMqlor--
1(eg rorv unorrvqrcov prSoo oe 6eq rrg oMq).ou1[e:
trg ono(eg pnope( vo npoonel.doer. To otrodv(upo r,':
RNA noLupepdonq ewon[(et ro Xptopr6ooLio Ue rL-
1o(o 6rdXuoq ror npoo66veror pr1 erdtrd sro opvllr(c
cpoprtop6vo DNA. Ie our6-ro pow6lo, to oo5v(ut-:
6r(ororor noti yprlyopo. H 6tdXuorl 05tet 6vo ovrr-rrc-
ro 6pto yro rr1 oroOepd qq roXUtrltog otiv6eorlq r-:
rd(r1g rou <108 M'1 sec'l yro or6Xo 75 bp. l-ldwoc -
npoyprortxr; moOepd npotrl0rlttrrlq toXUtqroq 1fo'-
ward rate constant) yro rdnoroug unortvlt5c, in vtt':
rpo(vetot vo e(voL >108 M'1 sec1, oro 6pro dLdxuo1c *
xor ndvo qn6 our6, H 6nuroupy(o rot rotompocc-
f-tog oerpdg oupnL6rcov 5og 6rou (r<ord tUXrl) rl RN:
no).u;repdorl ouvowqoet 6vov unortvqrq xot 6rlptouc-
y4oer 5vo ovotXr6 orjpnlor<o u<ov6 vq rpttd[et RNA E:
4rov oXerrrd opVn 6ro6Lroo(q. 'Etot, o Xp6vog no,
onoLre[roL yto tuXo(ouq r<rix].oug dto6olrr<qq oJvdi-
onq Kor 6rdmooqg oe Xolop6g 06oerq np6o6eoqg e .-
vot noAU peydoq ror 6e prnope[ vo epprrlveUoet nc.
Bp(or<er r; RNA nouprepdoq rov unoKrvnrq tr1g. Iuvo
noq, n RNA no).u;repdoq npSner vo XprlotUonotei ci.-
o prSoo yLo v' ovo(qrrloer lg O5oeLg np6o6eo4q rqc
H Eu<6vo 19.11 6e(yyer 6rt 11 5to6u<oo(o nt0ov6v en.-
rolUverot enerdrl o opltr6g m6Xog qg RNA notru;,a-
pdonq e(vor oL6xLnpo ro yovt6(<r-r1.to, 6Xt pt6vo g.:
ouyrerptpdvq o).r1Lou1[o unor<tvqq. Au[dvowoq r:
pdye0oq rou or6Xou, 11 moOepd rqg toX6tqrog 6toX--
orlq oro DNA ou[dver ow(orotlo xoL 6ev e[vot ntrec,
neproprmu<1. l-169 pretor<tve(rot ro 6v(upro on6 gLo t--
Xo(o 06o4 np6o6eorlq mo DNA o' 5vov unoKtvfr-
I[prgorvo pe opretd 6e6o;r6vo, toutrdXtorov tpe(g 5Lc-
<popertrSg dtepyoo(eg 6topoptptivouv l'lv toltitqr:
ovo(qtrloqg unortvqtq on6 tqv RNA noLuprepdor
I-1purrov, ro 5v(upo prnope( vo petortvrl0e( oe pto 1.tovc-
6tdoroq tulo(o 6to6poptrl r<otd prlxog rou DNA (,.oI,-
o0r1or1 - sliding"). AeUrepov, enet6rl ro Xpropi6otr.rl.tc
6Xet 6or6oLridrl muXcor4 6otrrr1 oro Borrqptox6 nupr-
voer66g, ;16l.19 ouv6e0e[ oe pro oMq].ouX[o rou Xpo[o-
I
ffiy r-"*W
L"Ah!**
'%ffi92
I
To revrpLxri dv(uUo npooddveror oe onoLodrlnore DNA
To olo6v(uUo npooDdverot mov unoKtvlrn
518 KEOAAAI0 19 MeToypoQrl orouq flporopudrreg
Boorxdg eworeg

EIK0NA 19.11 llporew6pevor Urlxovrouoi yro ro ntirg n RNA notruuepoon Bpioret 6vov unortvqnl. (o) otr(o0non (p) dLorplportxrl perotpo-
pd(y)ev6oneproXrxrlouvdeoqxordrdotoolnovonnOnon.llpooopUoog6voon6:C,Bustamante,etal.,J. Biol.Chem.274(1999):166665-
1 66668.
orilgorog, to 6v(upro Bp(orerot nMov nto xowd oe dtr-
trsg O5oetg, yeyov6g nou petcilvet to 1p6vo nou onqtte[-
ror yrq 6r6mooq rt enovonp6o6eorl oe pto 6IIn 06on
("dtotUrluortrrj getocpopd - intersegment transfer" i
(ovonndnon - hopping"). Tp(rov, evril e[vot ouv6ede;r6-
vo prq erdrd oe p[o 06o11, to 6v(upro gnope( v' otr].d[et
O5oetq qro DNA pr6Xpt vo Bpe0ei o unoxtvqtfg (.dpreor1
prerogopd - direct transfer").
I 9.7 To OAodv(upo TpononoteiTot Kord Tn
Arepyoofo Avoyvtipronq Kt An€Aeu8dptrlonq
on6 Touq Ynortvnrdq
* '0T0v
n RNA notrugepool npoo66verot o' 6vov unortvrlrrl, dto-
Xop((er iouq rtrtirvouq rou DNA yto vo dlptoupyrloet gto pero-
vpoqrrrt rpuootrido rt evootgor6vet vourtreorldto oto RNA.
& H evopfi ono qv RNA notruuepdoq 6ev e(vot novrore enttu-
xts.
* 0 nopoYovroq 0tYU0 ouvflOoq oneLeu0eptlveroL ono rrlv R-
NA notrupepoorl otov 1 veoouvre0etp6vrl otruotdo RNA ono-
rqoer pflrog -10 pooetrlv.
Mnoporipe rdlpo vo neptypdtpoupe ro orddto rnq Ue-
roypogrlg oriprgovo pe ttg olLrllent6pdoerg preto[ti
ttov 6rorpoperrrrov ptoptpcov tqg RNA nolu;repdoqg
Kor rou DNA er<proye(ou. H ow[6pooq 6vop[qg prnope(
vo neprypoge( pe Bdoq ttg nopopttitpouq nou ouvoQ(-
(owor orrlv :
o H ow(6pooq oloev([prou-unoKtvrlrrl [ertvd pe to
oXrlprolopr6 ev6q xretoro0 6Lpepoiq oupnA6xou,
6nog <po[vetot offlv Etr6vo 19.12(o). "K].etor6" orl-
po(ver dtt to DNA nopop5vet 6ixtrtovo. O oXqgotL-
opr6g rou xletoro0 6tprepoUg oupntr5xou e(vot qwt-
orperrr6g. 'Etot, ouv4Oroq neprypd<petot on6 pto
oro0epd tooppon[og (Ke). YndpXet preydLo e0pog
orrg nprdq qg mo0epdg tooppono(og Yto ro cDCq-
prottopr6 tou xl,etoroU oupntr6rou.
o To xtreror6 orjpnloro petorp6netot oe ovotyr6 o01t-
rrloxo pe
"oflodtdroqn, ;rtog ptr<priq neptopiq DNA
pr6oo orrlv oMqLou(o 5nou npoo6dvetot ro 6v(u-
pro, 6nog qo(vetot orqv Etr6vo 19.12(9). l-to roug
neproo6repouq unoKtvrlr6g, 11 perorponrl on6 to
ovorXr6 oro xtretcn6 oriprnloro e(vor ptq owtorpe-
nrj ror owri q ow(6poor1 prnope[ vo neprypoqe(
on6 11 oro0epd npool0rlrxrlg toX0r4roq (forward
rate constant) (kr). Oo16oo, pepxo( unoxwqrdg (n'1'
ol unortqt6g tou rRNA), 6e drlprtoupyo6v oroOepd
ovorXrd o0ptntroro rot our6 e(vqt to xLet6( yto 11
priOprori roug. O nopdyowoq o no(et orlprowtx6
p6Lo orqv owidpooq ri€nq GMne Ev6r4ra 19.19).
Or preroBdoetg nou ou;tBo(vouv on6 tqv 6vop[4 ;t5-
Xpr rnv entprlruvorl en(o4g ouvodeUowot on6 orl-
;rowrr6g oMoy6g orq 6ogrl Kot rn oOmoorl tou
ougnl6rou,
Ot oMoy6g oro oXrl;ro qq RNA notru;repdorlq ouvo-
6efouv ug t<tv4ttx5g ;retoBdoetg nou neptypdgnKov
nporlyouprdvtog, ro0rilg r<ot q ;rerdBoon oro o6pntroro
enrpulruvoqg (nopouotd(erot firlv EIK0NA 19.13). Ito
xtretor6 oUpnloro, to ol,o6v(upro qg RNA noLupepd-
oqg roLrimer nep(nou 55 bp- DNA, errew6pevo on6 --
55 5og -+ 1. To 6[x].tovo DNA npoo65vetot rup(tog xot6
priKoq rqq puog ntreupdg tou oloev(ripou, 6nou SpXetot
o' enoqrl ;re nq C-refur6q neptoX6g rov o uflopovd6tov
roOdrq Kor Ue rtq neptoldg 2 xat 4 rnq o unoUovd6og
(BL6ne nopor6to). Kord q getdBooq oro ovotXr6 o6pr-
nlor<o, q drop6prpoorl t6oo qg RNA noluprepdoqg 6o'o
Kqr rou DNA oMd(et. Ot nto dpoonxSq oMoy6g orrl 6o-
Uri rou oupnl"6rou qo(vowot crqv Ex6vo 19.12: (o) ptto
xdprUn rou DNA -90 potp6v, nou enrpdnet orov xtrtivo
"i1l ? To 0tr06v(upo Tpononore[ror Koro rr1 Arepyooro AvoyvtirpLolq rr Anetreu06ptlorig ond rouQ YnoxLvnte c 519
Boorxfg eworeg

eKpoye(o vq npooeyy[oer ro evepy6 x6wpo rou ev(6-
Uou' (B) dvorypro tov x.trcirvrov rou DNA rou unoxwrltl
pero[6 --11 rqt +3 ovoEopxd pre qv ogerrlp(o rnq Ue-
roypocprlg r<or (y) r<tre(orpo rov (ooyovrcbv> tou ev(0prou
ciore vo neprruxl,cioer to rpui;ro rou unoKrwlr4 nou Bp(-
oKeror r<o0odxd on6 rrlv orperlp(o preroypocprlg. Iuve-
ncilg, ro oqge(o enorpriq rou ulroKrvrluj mo ovorXr6
oUpnl,oro erre(vowor on5 --55 oe -+20.
To en6prevo Brigo e(vot vo evooUorco0oUv rq 5[o
npdrro voux.treot(drq Kor vo oXqpronore( rivog <ptrlorpo-
6eorepx6g 6eopr6q prero[ri roug. Aur6 drlproupye( 5vo
rproroy6q o0pnAoxo (ternary complex) nou nepr6ler
RNA, DNA xor ro rlv(upro. Iroug neproo6repouq unoKr-
vqr6g, qXqpror(eror puo oLuo(6q RNA nou SXer pr4r<og
opr<er6g Bdoerg, Xrr:p(g getor<(vqoq rou ev(rjprou xotd
prrlrog rou erpoye(ou.
M6Ltg npoore0e( puo Bdoq, un6pXer rdnoro nt0ov6-
rrlro ro 6v(upo v' onel,euOeprioer trlv oluo[6o rou RNA,
rorolqyowog oe npoi6wq overtruxo6q (errptonx{g)
520 KEOMA|019 Metoyporprl mouq flpoxoputirreg
EIK0NA 19.12 H RNA notrupepoorl nepvoer on6 op-
rerd orddLo fiprv rnv enqrlruvorl. 'Evo r],erotri 6q:e-
prig orjpntroro gerorp6neror oe pto ovoLXrrl poptp4 u
6neno o' 6vo rproroy6g oupntroro. l-lpoooplroop6vo
on6 S. P. Haugen, W. Ross ror R. L. Gourse, Nat. Rev.
Microbiol. 6 (2008): 507-51 I
dvop{4q (abortive initiation). Merd rrlv one-
Leu05prooq rou ovenrruXorig npoi6woq, ro
6v(upro [ovopf(et rrl orlvOeor1 RNA oq 06orl
* 1. EnqvoloprBov6prevor r0xi,or ovenruXd:v
ev6p[eov 611 prro u pyo rlv o].ryovo u r].eor(6 ro
nou ouvrlOtog 51ouv pqxog tr-r6vo Liyeg Bdoetg
(nop6n prnope( vo rptdoouv oe grir<og ro 20
nt), npw ro 6v(upro npoyprorxd roro<pdper vo
6rog0yer on6 rov unorrqrl.
Or otrtr4l.enrdpdoerg Ue rrlv RNA noAupre-
pdoq te).u<d e[otre(rpowor r<ord q 6rorpuyq
rou unoKrvrlq. M6Xprg 6rou 4 oluo[6o rou
RNA ensKro0e( oe 15-20 nt, ro 5v(upro yevxd
61et nepdoer on' 6treq lg pretoBdoerq nou Xo-
porcrqp(ouv 6vo orjprnLor<o enrprlruvoqg. Or
d0o nro govepSg pretoBdoerg e(vor 4 qneLeu-
05poor1 rou nop6yowo o [ono6(6eror mrlv
Ex6vo 19.12(y)l Kr o oxqporropr6g ev6g ou;r-
nL6t<ou nou rorol,o;rBdver pr6vo -35 bp DNA,
ow( ralv -70 bp nou e[vor Xoporrrlptmu<6q
uov oupnl5rov rov unortvqrrriv. MoLov6rr 4
oneLeuO6pcoorl rou o ouvqOoq ougBo(ver xo-
rd q 6roquyrl rou unorwrlrl, our6 6ev e(vot
npoononoripevo yro tq prerdBooq oqv en[-
prrlruvoq. Ie optopr6,veg neprmdloerg o o SXer
rouronor4Oe( oe oU prnLor<o enq-rr1ruvor1g, oi.-
)d q oUvdeo{ rou Ue ro 5v(upro (ocog owrro-
tomp((er enovonp6o6eorl orov nup{vo rou
ev(0;rou r<otd 14 gdo4 rrlg enqlrp<uvor1g.
I 9.8 O l-lopdyovroq o EA6yx€r rnv np6o6€on
oTo DNA Avoyvrrrpi(ovToq luyrerprpdveq
AAAnAouxieq Tov Ynorrvnr6v
* 'EvoS
unorLvrprlg npoo6ropl(eror ono rrlv nopouo[o ppoxerotv
ouvorvelrtlv otrtrrltrouXrtilv oe ouyrerprpeveg 0eoerg.
* 0r ouvorvelx5g otrtr1trouXreq rou unorrvlrrl ouvlOtog onore-
Lo0vror ond ;-uo noup[v1 orrlv oqerrlp(0, 6vo e(opepeq pe otr-
trritrouXro nopepqepfl pe qv TATAAT oq 0601 --10 rL 6vo
otrtro e(opepeg pe gto otrtr1trouxio nopepQept Ue qv TTGA-
CA oq 0eo1 --35
* 0t qexoproTo( unouvlr6g ouvrl0rog dtotpepouv ono q ouvol-
velrq otrtr1trouX(o oe pro fi neprooorepeq 0eoerg.
* H onoretreopolr6rrlro rou unorrvqrrl gnoper enloqg v0 cnl-
peo(eror rot ono enrnpooOero orotXe[0.
Avodtrd
DNA
I-lepro1r1 3 tou o
Tlsproxn 2 Tou o
lleptoXrl 1.1 rou o
E(opepdg -1 0
Ktrtivog 91-expoye[o
Ko0o0rrd
DNAfiepLoXr1 3,2 rou o
Ktrrirvog
ergoye[o
Boorx6g eworeg

.aciexet 10v 0 Kot
-5: --3: -30 -20 -10 1 +10 +20 +30
-- ,., ...:;iJ:.i'ttti:,: oxluoT((tTot oTtq 10 Bdoetq,
.' -- :-:- .,cKorxov€rrtq0T10Q6q0116ro-356(l)qr0-55
-5: -40 -30 -20 -10 1 +10 +20 +30
. .:.:. :..':. r:.,::t, oxnuoll(tT0t ortq 15-20
-5a -40 -30 -20 -10 '1 +10 +20 +30
H RNA notruUepool opxtKo epXetot o' enoqrl pe
-,- ,- :ro -55 eolq +20.'Orov dtfororot o o, o nuplvog
, .--,, oloretrtrerot oro -30. MotrLq ro ev(upo perortvloel
- -, . - --: ','rvLro o0pntroro enLglruvolg.
--. - -:,).qAouXio DNA nou l.ettoupyio qg e(vot v' ovo-
-:.leror an6 npare'tvtq, o unoKtvnrnq 6tog6pet
--: .: cMq).ouX(eg tcov ono(ov o p6og e(vot vo trle-
-: . : loowoL. Ot nLrlpo<pop(eg yro tr1 Lettoupy(o tou
,*:...rlrrl nopdXowor oneu0e(oq on6 rlv oMqLou-
-- ::.l DNA: n 6oUn tou eivot to or1trro. Aut6 e(voL
. : ., .oorxd nopddetypro ptog cis-evepyoU 05o4q,
- --: op[m1Ks ortq Eu<6veg 2.16 rot 2,17. Aw(Oeto,
- .,:co(dpreveg neptoXdg ononorjv to v64ptd toug
,-,: lroorou q nLrlpo<pop(o prerorpep0e( orrl uopan
. r-l.ou dMou vour<Leivtxoti oE6oS n npottelvqq.
-,oq rpdnog vo oledtdoouge Svov unor<tvr1t1 0o
--:. Jro ouyr<erpttrt6vq oMrlLoul(o DNA v' ovoyvop((e-
-: :lro r1v RNA noLuprepdoq. Kd0e unortvqtrlg 0'
--::iorjwov on6 tr1 ouyxercptp6vq otrLr1i.ou1(o, r1
--,:.c1rorov, 0o rrlv nepildprBove. Iro Borcrqptor<6 yo-
i.-;-io, ro etrolroro prrl<og nou 0o prnopotioe vo no-
: rr(01 dvo enopxdg orlpro e(vor 12 bp. (OnoLo6rlnote
:::4r.rrepq oMqtrouX(o e(vot nt0ov6 vo oXqprottore[ op-
-..:,q rpop6g -onLcog xord t[p1- n 6rot vo npor<o].5oet
-.-5q orlporo. To e).d1Loro prrl<oq nou onottekot yto
-: . cbrxrl ovoyvcoptorl ou(dvet pre to pr5yeOoq rou Yovt-
- ,.roroq - 6vo np6B).npo oro europu<ortrd yovtdttrt-
..::c), H oMrltrouX(o tov 12 bp de Xpetd(erot vo e(vot
ouveX6prevrl. Av 6voq ouyrerptp6vog opt0p6q Bdoeolv
6toXrop(er 6rio oro0ep6q BpoXftepeq otrLfltouX[eq, to
ouvotrrr6 pulr<oq touq 0o prnoporioe vq e(vot )try6tepo
oTro 12 bp, enetdrj q (6to 11 an6orao4 StoXtoptogori
onoretre( 6vo tprrlgo rou otiporoq (6orol rot ov q evdt6-
peor'1 oMrlLouX(o e(vot doXerrl). It4v npoyprortr6vlto,
11 RNA noLuprepdoq ovoyvop[(u tLg DNA oMq].ouX(eg
rov unoKtvntriv oe preyd).o Fq0U6 ptSoo "dpeoqg ovd-
Yvoonq" ouyrerptpSvtov Bdoeov oro DNA on6 ouyre-
r<pr;r5vo opuvo(6o tou oloev(tiptou. Ot ewunootordg
6torpop6 g tcr4ioq trr:v 6totp6ptlv Bor,crl ptoxtirv uno Ktvn-
rdlv ono6(6owot rqrd noLf or4 6totpopenxrl onotetr'e-
o1-rottr6rqro ovdyvoo4g tcrlv otrtrrllouXtrirv toug on6
ng optvo(tx6g oLl.qlou(eq rov unopovddtov o Kot o.
Or on6netpeq yto rnv ovoyvtirptorl tov tdtorltov
rou DNA nou e(vot onopofurlreg Yto rrlv np6odeoq qg
RNA noluprepdonq (er<(vqoov ouyt<pivowoq ttg oMt'1-
),ouX(eg 6torpopettxrilv unortvrltc0v. Kd0e onopofurlrl
voux.treot6rrrl oltrr1Lou1(o 0o 6npene vo undplet oe
5oug roug unorwqr6g. Mto t6toto oMrll'oufo Xopo-
rcqp((eror ouw4p4p6vq (conserved). Mto ouwrlprlpS-
vr1 oM4l,ouX(o 6ev efvot ovoyromxd ouwqpqpSvq oe
rd0e (eXoprorrl 05o'r1qq, 6r1}'odri enmpSnerot xdnoto
norril.opropg(o. l-kiq ovoltioupe ;rto oMrl).ouX(o DNA
yro vo npoodtop(ooupe qv 6Xet ouwqprlOe( opr<etd
rilore v' owrnpooonefet 5vo ovoyvtr:pfotpro onpo;
Auvqnr6g Odoetg ovoyvrirptorlg tou DNA ;rnopef vo
npoo6roprorotiv ovotpopu<d pe pto eft6ovu<eup5vr1 oL-
Lql,ou{o 11 ono(o owrnpoooTle0et rq Bdoq nou Bp(oxe-
ror nro ouy3r6 oe xdOe 0dor1. Mto ouvolvenx{ oAtrqtrou-
;1fo (consensus sequence) xo0opi(etot on6 t4v eu0u-
ypdpptorl 6trtov ttov yvomc0v nopo6etyprdtov 6tot
rilore vo preytoronotrl0e( q oproloy(o rouq. l-to vo y[vet
ono6erc4 puo oM4trou{o oq ouvotvertr4, rdOe ouYKe-
rp4l6vr; Fdon qq np6net vo entxpote( or4v ow(orotXq
06o11 r<ot to neptoo6repo npoYpqrtxd nopode(yproro
np6ner vo 6ro<p5pouv pr6vo rotd pto 4 6Uo owtroro-
mdoetq on6 rq ouvowenrrl oM4trou1(o.
'Evo ewunootox6 Xoporrqptonr6 trrlv unorwrlrcbv
oro E. colie(vot 11 iMeulr1 et<terupivrlq ouwrTp1oqq aA-
lqAouyuhv oe 6).11 rrlv 6rcrooq ttov 75 bp nou oler(etor
Ue rqv RNA noluprep6oq. 0o16oo, rdnoto BpoXSo tprl-
proto pr6oo orov unoKtvrlrl 6Xouv ouwrlpqOe( rot e(vot
(onr<r1g oqproo(og yto t4 l,ettoupy(o tou. H ouw1p1ory
p6vo noAA Fpaxeuiv ouva:euxtisv aM7ouyuitv e(vatru-
nx6 yapaxr4ptwtx6 tuv pu1lttottxtitv 06'oeuv (6nuq ot
unoxlrySg) t6oo ota npoKapu@rtKd 6oo xot ora euKa-
puanxd yovtdubpara.
Apr<er6g oMrll.o u1(eq tov Bor,mq p rorcilv uno rtvltriv
ouverogdpouv ornv ovoyvdrprori toug on6 to oLodv(u-
Uo rnq RNA nol,u;repdonq. Ario oLl'r1i.ou1(eg 6 bp,
yvoord og ororleio -10 G10 element) rot orotlefo'35
(-35 element) (6norg rot to prlxog rnq "6toqrnUotxtlg"
oMrltrouX(og nou nopeprBdMetqt ;reto[rj rouq - (spac-
er>> sequence) e(vor ouvqOtog oqpowu<6tepeg on6 ou-
169 ro0out6g ttq oll4l,ou1(eg ovoyvrirptoqg. Ot
oll,rll,ouX(eg rov unoKtvrlrciv nou Bp(orovtot mqv
o<peqp(o r<or oxp$rig 5[nl.o t1g, ot otrtr4]'ou{eq Ydpo)
on6 ro ororXe(o -10 [yvotmd oq "6teupup6vo orotXefo -
:i.* Ol-lopoyovroqItyUoEtreyXeLUvtlpoobeorloroDNAAvoyvopi(ovrogluyrexprpeveqAtrtrltrouXieqrovYno<1,'- - 521

'l0D (extended -10 element) mqv ovodKn nl,eupd ror
"6rqxoprorig" (d iscriminator) mrlv xo0o6u<r1 nLeupdl
r<or or 10-20 bp og6ocog qvodrKd rou morXe(ou -35
[yvoor6g oq uororxe(o UP, (UP element)], en(oqg o],-
Lqlenrdpotiv rord erdu<6 rp6no Ue rrlv RNA noLu;repd-
orl Kor ouveroqSpouv orrlv onorel,eoporrr6rlro rou
unorlrprl.
. :rouq neptoo6repouq unoKrvqr6g ovoyvrop((eror
pro neproXrl 6 bp nep(nou 10 bp qvo6trd rnq oqe-
qp(oq (r1 npoygoru<r1 on6orooq on5 r1v oqeqp(o
6rog6per eLogpd on6 unoxwqrrl oe unorwqrrl). Au-
r1 q e[opepqg o].].q].ou1(o ouvq0rog ovoprd(eror
ororXe[o -10, r<ow( Pribnow (Pribnow box) ri oMuirg
xourf TATA (TATA box). H ouvowenrrl oMqLouX(o
TATAAT ouvor.l;((eror orn UopQn
Tao Ags T+s Aoo A.5o Tso
or 6e(rcreg 64Lrirvouv rnv eKorofiro(o ovo),oyfo
ulg oulv6tep4g Bdorlg, nou norr0.et on6 45"/"
6tog 96%. (Mro 05or;Xcop(g ep<povr1 npo{gqoq yro
xdnoro Bdorl 0o 6rlLcov6rov ge ro oriprBoAo N). H
ouXv6qto epr<pdvroqg owrmorle( mr1 o4proo(o
our<ilv r<ov (euyc0v Bdoeov yto rrlv np6o6eor1 qg
RNA noluprepdonS. 'Etor, ro opXrx6 noLri ourrrn-
pqpSvo TA ror ro rellr6 oXe66v on5l,uro ouwn-
pqp5vo T orqv oLL4LouX[o -10 oulvd e(vor
ro(ptog orlproo(og yro rnv ovoyvriprorl rou unoKr-
vqrrl. I{pepo yvrop((oupe 5rr ro ororXe[o -10
flpoyUoronore( er6u<69 yro rrlv oLL4Loul(o enoE6g
mrg neprolSq 2.3 Kor 2.4 rov nopdyowo o (BMne
nopordro). 0o16oo, ouq q neprofi rou unoKrvn-
ri e(vot 6ir<).tovq oro x.l,eriyr6 oriprnLoxo ror ;rov6-
r<Lovr1 oro ovorXr6: ouvenc(lg, or oM4Lenr6p6oerq
UeroEU tou morXe(ou -10 rcot qg RNA notruUepd-
o4g e(vor noLUnl.oxeg ror oMd(ouv oe drogoper-
xd o166ro rnq 6repyoo(og 6vop(r1g rnq
peroypogrlg.
. To ouwnprlp6vo e[opepdg -35 bp ovodrrd rrlg
oqerlpfog ovoprd(eror orotleio -35 C35 element).
H ouvolerrrrl oMqLouX(o e(vor TTGACA. Ie nro tre-
nrotrrep{ Uopen, n ouwnpnprivrl o},r1}.ou{o 61er
os eqnq
Ta+ Gze Aos Cs+ Aqs
Ot Bdoerg oe our6 ro morle(o ollrl).enrdporjv
oneuOeiog Ue rnv neproX4 4.2 tou nopdyowo o
(BL6ne nopordtro) ;re nop6proto rp6no 16oo oro
ovorXr6 6oo ror oro xLeror6 oriprnLoro.
r H on6moon ;rero[ri rtov O5oeov -35 ror -10 e(vor
16-18 bp oro -90% rov unoKrvrlrciv. Ing un6lor-
neg neprmdroerg e(vot '15 bp ro lly6repo xor 20 bp
ro neproo6repo. Molov6tt 4 (6n 4 aMrlLouy[a mo
peya)thepo pipog u1q napepBaM6pevrlq nepoyfig
e[vot oyenxd ao4pawq, 4 on6otao4 elvat xa[pn
enetdr1, t6yu r7q elLxoet6o0q E1oryq rcu DNA, xa1o-
pGet 6XL 1t6vo tov xardM4io 6nyapop6 rav 6Ao
neprcydtv r4q BNA ndtupepdo1q nou a)rLr1ient-
6poiv, aMa en[oqq Katro ye@petpm6 npooavarcit-
o1t6 tuv 50o 06,oeav.
o H oqetqp(o ouvrlOog (>90%) e(vor truo noup(vr1, ou-
522 KE0MAI019 Meroyporprtmouq llpoxoputirreg
yyd 4 r<ewprrq Pdon qq oMr;),ouX(oq CAT. Oordc:
q ouwrlpqol ourrlg rrlg rprn).6rog 6ev e(vor tdoo lr-
yd).r1 urore vo Oecope[tot oq unoxpeorxd oqpro.
o Opropdvo (e[yq Bdoeov ornv neproX( pero(U r:
o<peqp(oq Kor rou morXe(ou -10 ovoyvrrrp((or,.::
on6 qv nepropl 1.2 rou nopdyowo o (B).dne .:.
por<drco). l-to nopddetypo, Uro oMr;).en(6poorl o : .
r<r1 yro t4v o).).4LouX(o prero[U ev6g roro].oi:r:-
youov[vr1g orov x.,rrtvo prrl-expoye(o 6tjo O5oetq .:.
0o6u<d rou orotle(ou -10 e(vor e(orpelr<d of lrc,l .
rq yro tov r<o0optopr6 rrlg oro0ep6rqroq r3-
ovotXro0 oupnl6r<ou. 'Etor, oL 6ro<pop6g orqv o).1.--
Loul(o rou unoKrvrlrrl oe O6oetg nou 6ev e(vor n:, -
ouwrlpqpdveg prnope( vo eu0Uvowor yro rnv ere::-
yevq roXU rov 6Lo<p6prr-lv unorrvqrov.
o Or pdoerq oro 6eupupr6vo orotXe(o -'10 npooeyyil: ,.
ror on6 qv neptoXll 3.0 rou nopdyowo o (B).dne ::
poxdtrrr). H oM.r;LouX(o TGN oro ovo6rx6 dr<pc .:-
ororXe(ou -10 ouprpret6Xet oe oMq).ent6pdoerg L-.:.
repo orlUowu<6,9 yro qv 6vop[q rnq peroypcc*:
6rov 11 oMr;),ou1[o rou ororle[ou -35 orov unoKt,,-:*
6e pord(er noLU pe 14 ouvorvelrcl. Aur6 ovo6err,,,:
norg ororleroOerotjwor or oMrlLoul[eq rou uno(.,-.
14: prrpr; oprot6rlto Us rrl ouvorvetLrrl oMqAc-,.:
oro 6vo ororXe(o ;inope( v' owroroOpuore( ond ue,:.
).r1 oprot6qro o'6vo dMo orotle(o.
o H neptoxn -20 bp ovo6u<d rou ororle(ou -35 cr-:.-
Lenr6pd pre ro CTD rotv 6Uo o unopovddcov, OL i .
nrcuoerg ourcbv rov ol.Lq).en16pdoerr-rv c:-.
evepy6tqro rou unoKrvrlrrl prnope( vo e(vor op(,::
oqprowrr<69, ou(dvowog rn Ueroypoerl rou).:, .
orov rord p-uo rd(r1 prey60oug yro no).ri er<gpoi:*i.
vouq unoKrvnt6g, n.X. urlv yovr6(rov tou rRNA 'O :.
or ouyKeKprpdveg oMrlLouX(eg protd(ouv noAu p.
ouvorverrrrl, 11 neptoXrl ourl ovo<p6peror c,Dq c.: .
1e(o UP (UP element).
H 6o9( ev6g unoxtvrlrq, pre ro enrpen6pevo eupcl
norril.otrropg(oq (on6xlrorlq) on6 ro B6trrroro, nop.--
oro(eror mqv
I 9.9 H AnoreAeopoTrK6TnTo Tov YnoKlvnTd,v
Mnopel v'Au{dvel 6 vo EAAordveror on6
MerdAAoIn
+ 0r uopvrllx6qD UsToMoqerq nou per6vouv qv onoretreopc. -
rriqro rou unorrvrlrl ouv1Oog pertirvouv q ouppopqrr:c-
npoq rtq ouvorvenr6g otrtrqtrouXieg, ev6 or ,,0ertr6g,. perc;.
tro(erq 6Xouv ro ovri0ero onor6treopo.
* MerotrtroEeLq oqv -35 otrtr1trouXro pnope[ vo enrlpeooo-,
rrlv opXrrl npoodeoq qg RNA notrugepooqg.
* Merotrtrolerq orqv -10 otrtrr1trouX(o pnope( vo enqpeooc-,
qv np6odeorl 11 rrlv ovldpool onodroro[lq rou UrToTpsr:
evo rLeroro ougntroro oe ovorXro.
Boorxeg evvoreg

l-leproxn 3.0 tleproXrl 2,3-2.4 fleptoXq 1 .2
TOU O TOU O IOU O
IroLXeio UP -35 Ext -10 Uls +l
-t n0txs[0 Tou DNA Kor T0 ouoToTrKo qq 6oUnq Trlq RNA notru;repoolg nou ouppereXouv oTnv ovoyvtirptorl rou unortvlrrl
----,-, 1: tlpoooppool.t6voono:S.P.Haugen,W.RossrotR.L.Gourse, Nat.Rev.Microbiol.6(2008):507-519.
- ,- --::.c getoMd[eotv pnope( vo npooq6pouv
- -: - -: :: ryerlKo |re q trettoupy(o rou unoKtvrl-
- -. .:'-i.cEelq orouq unoxtvrltdg enqpe6(ouv to
- -.: : :-: eKepooqg tou(tov) yovtd(ou(tr.rv) nou
-, : -, ,*c[g v' oMd(ouv outd xo0outd ro yovt-
' -,: -::.l.ro. OL nepLoo6tepeg prerotrd[etg ovo-
-: -:, :. ;e Borcrqpto tog oloydg tqg oMq).ou-
- : - - - : l-','ouv oe onotreto rl Uey6).q ;re(tooq qq
. --: ::j-: :tDV yettovtrrirv yovt6(ov. E(vot yvoor5g
- , :: , nrxeq peroM6[erg (down mutations)' AMo-
- - - : :r-'.-iou1(og euO0vowot Kot Yl'o 0erlx6g pe-
-:--: iarc (up mutations) ottg ono(eg undpXet
-. -: : ". -3-ioYpoQ( on6 tov unor<tvqrq.
-- l- :. ,,8ettrd,g,, doo xot ot "opvr1ttr6g" pretoMd-
.. - :: --. .:L oe oXeoq pre q ouvry1to1tiv4 onotetre-
- -- .- -:: evdg ouyrer<ptp5vou unoKtvt'1tq nou
- - - - I : .:.rL.tret notrU. 'Etot, prto oMoyr; nou ovoYvo-
. -.-- -: ::i.1rtr<q petdLl.o[r1 o' 6vov unor<tvr;tr| pno-
. : - -- , lrcUovrr-loe( not6 o' 6vov 6Mo unor<tvqtrl (o
- - - : r. :. cto rorooroorl 0o pnopo[oe vo e(vot o16-
:-r:: cnortleoprottr6g on6 tq petol'LoyprSvrl
-,: -- -:- :cLJrou unor<tvr1q). Ae6o;r6vo on6 in vivo
- . :-: - : .r.;q ovoyvcop((ouv 11 ouvoLtr<r1 rote[Ouvot1
- - r . :.-: -rou npor<ole[ 11 pretdMo[q.
- -. ::,,.oEeiq rou ou(dvouv ttlv opot6tqro rov
- . - " " - <ot -35 Ue rtq ouvotvettx6g oMrlLoulieg 11
. :- - : - , :- pretolti toug on6oroorl rdto on6 17 bp
- - : - : :- lc. ouv rlv evepy6rrlto tou unortvrltrl. l-lo-
. -. -: : *iroMo[etg nou getovouv ttlv ogor6tqro
- - -:l-::.i nLeupog Ue rrl ouvotvettr<r1 oMrltrouX(o
- - :::::::icuv rr1 preto(0 toug on6orooq (on6 to
. - -: :-, '7 bp) o6rlyoriv oe petcol-ttivrl evepy6qto
-' ,-.n, Or opvqttr6q getotrtrd(etq ouvq0og
- .: -:*,c',/Tot ortq TIto ouwrlprlpr5veg O5oetg tou
. - ," -:- :riBeBottrrvowoq rnv t6to(tepr1 oqpoo(o ou-
-
- . ::ceolv Yto rnv onorel'eopottr6qto rou uno-
l;coo, undplouv onopodtrdg elotp6oetg oe
- --: -: :ouq rov6veq.
-: r,opddetYpo, 6vog unor<tvqtrlq ge ouvotvettxdg
- , ..'.cr1[eg oe 6].o to oroLXe[o nou neptypdqrlKov nq-
:: -:.:(erot mqv Etr6vo 19.14. Aev undpXouv 6ptolq
-:-: r. rpuotxo[ unortvqt6g mo yovt6(trl1-to tou E. coli
. : -:, ,rro( unoxLvqt6g nou totptd(ouv on6Luto 5le 11
- - r .::.<rl oMlLoux(o oe 6).eg ttg 06oetg srnv npoY-
-:- . ::-:l e(vot oo0evdorepot on6 unortvqtdg pe tou-
- l.:, evo ).d0og to(ptoopro oro ouvotvettrd
e(opepri -10 r<ot -35. Aw6 ou;rBo(ver enet6rl npoo56-
vowor 16oo orptXr6 orqv RNA noLuprepdo4 rilore ornv
npoyl.tonx6tnro nopeuno6((ouv tr1 6ro<puyf rou uTloKl'-
VNTN,
l-ro vo ro0optoroUv ot on6l'uteg entmriloetg rov Ue-
roMd[e<ov rov uTloKtvnrcilv, pretpdtot in vitro q ouyy6-
verq rnq RNA nouprepdorlg Yto rov dypto rot ro
pretotrl.oyprdvo t6no rtov uTloKtvntriv' H etepoy6veto
tr1q roxfunroq Ue rnv ono(o q RNA nol'uprepdorl npoo-
66verot in vitro oroug 6totpopetxorig unorwrlrdg o1e{-
(etor r<oLd ;re q ouly6rrlro rnq pretoyporprlg 6tov r'
ow(orotlo yov(6to errppd(owot in vivo. Enercrefuowog
ourl qv ovdl.uorl, ;rnopei vo npoodtoptore[ to orddto
oro ono(o prto pret6Mo[r1 enrlped(er r1v ono6onx6tqto
ev6g unoxrvntn. ALI'd(et q ouyy6veto rou unorwqtrl
yro np6o6eoq rqg RNA nol'uprepdorlg; Arpqvet pev ro
6v(u;ro trqv5 vo npoode0e( oM6 ov(rqvo vo (erwrloet
rrl Uerqypoeri; Atogoponoe( trlv enrppoq ev6q enxou-
pu<ori nopdyowo;
Metpdrwoq ttg t<tvtlttt<69 oro0epdq Yto ro ox!1gort-
opr6 ev5g x.tretoto0 oupntr6rou Kot rnq gerotponrig rou
oe ovotXr5 orjptnl.ot<o, prnopo6ge vo dtorp(voupe to
6rio ord6to rrlg ow(dpooqg 6vop(r1g:
. Ot opvnrtK5g geroMd(etq orrlv ol.i.r1}'ouX(o -35 ou-
vr'1Ooq get6vouv r1v toX0rlto oxnUortopoti tou
xtretorori ouptntr6rou, oLl'd 6ev ovoqr6Mouv rn Ue-
rotponf rou oe ovotXr6 oUPnl'oxo
. Ot opvnrtKdg pretoL),d(elq orrlv o).trr1l,ouX(o -10
prnopei vo petc0oouv e(re tov op1tr6 oXqpottopr6
ev6g xtretmori ou;rntr6r<ou e[te r1 petotpon{ tou
orrlv ovotXrrl Uopqi efue xot ro 5Uo.
H ouvowenrrl oMr1tr'ouX(o orq 06o11 -10 qnotelefuqt
onorletonrd on6 (e0yq Bdoeu:v A-T, Stopr6prpoon TIou
9on06 orov opXtr<6 6roXoptopr6 tou DNA oe provo0g
xLrivoug. H Lty6tepq ev6pyeto nou onorefuor q 6totd-
po[r1 tov (euydrv A-T oe ox6on Ue ro (eUyq G-C orlpo(-
ver 611 dvo ttrrripo pe (e0yq A-T onotte( prtrp6teprl
noo6rqto evSpyetoq yo to 6toXcoptqr6 tov xi.rirv<ov' Ot
otrr1Lou{eq op5oo4 y0po rot r<o0odtrd on6 tqv o<pe-
tqp(o en(oqg enqped(ouv qv 5vop[r1' Ax6p4, n np6rn
neproX{ nou preroypdqerot (on6 -+1 Udxpt -+20) enq-
pedEfl rrlv toXriqro Ue rnv ono(o q RNA noLuprep6oq
onogot<p0verot on6 Tov unoKtvnrrl r<ot yt'our6 enr6pd
orqv roXti rou unoKtvnr4. 'Etot, 4 ouvotrrr4 toXtig tou
unorwqtq 6e ;rnope(ndwore vo npoBtre<p0ei on6 ttg ou-
IroLXe[o UP
'i:':.t g 4noat^eoporLrorrlro rolv Ynoxrvlrdlv Mnopei v'Au[overot 1 vo EtrtroroveTot 0[o l,ra :):::- 523

vorvenKdq oMqAouXieg rou, ox6pr4 r<r ov tr4gOoriv un6_
qrlro enrnp5o0ero ororXe(o nou ovoyvrop((er 4 RNA no_
Aupepdo4 elc6g on6 trg oMqi.ouX(eg -10 ror -35.
. Molov6rr 4 opor6rqto Ue rrl ouvorveru<r1 oMrlAou_
{o e(vor lprlorpro epyo},e(o yro rrlv ovoyvcilprorl unoxr-
vrlrrirv pr6vo Ue Bdon qv oMql,ouX(o rou DNA, 6nog
en[o4g xoL nop6rr ot (runrKo(> unoxtvqrdg nepdlouv
eudtdr<pneg oMrltrouX(eg -35 r<or -10, noMoi unoKrvrl-
tdg 6e 6ro05rouv eurprvrl morlefo -10 ror/ri -35. Ie
notrilig on6 ow6g rtg neprmc0oerq, o unoruvrlrlg 6ev
ovoyvrop((etor on6 g6v4 rrlv RNA no).uprepdor; xor
qnorre( pto enrKouprKri npcoreivrl (,evepyonor4riq, -
activator, BL6ne Kegdioro 26) nou unepBo(ver rrlv ove_
ndprero tov evdoyevcirv oMql,enrbpdoeov pero[ti RNA
notrupepdo4q Kor unoKrvrlrl. Eivot oqpowrr6 en(or1g
vo_rov[oou1,re 6rr "g6trrrm4 evepy6qroD 6e orlpo,v
.
";r5yrorr1 evepy6rqro,,. llol,Lof unorrvqt6g dlouv e(e_
).t10e[ pre oMqtrouXfeg nou on6louv no],rj on6 to vo e[-
vor ouvqrverrr6g oxptBrig enerdrl 6ev efvor ro Bdtrnoro
yto ro rUmopo vo grrd;yer 16oo nol,[ npoi6v nou KoJ_
6monorekot on6 ro RNA pietdypoeo rouq.
l9.lO l-loAA6q t-teproxdq Tnq RNA
l-loAupepdonq'EpxovTor Apeoo oe
Enogfi U€ Tov DNA ynorrvnrfi
* H doHl rou 070 otrtro(et orov ouvdeeror ge rov nuplvo rou ev_
(u;rou, enup6n0vr0q oTrq neptoXeg yto npoodeoq oro DNA v,
otrtrltrenrdpooouv Ur Tov unorrvr;r1
er llotrtreq neproXeg rou 070 oitrrltrenrdpouv pe rov unorLvrlrl.
s H unopovodo o en(orig ouppotrtret orr;v ovoyvtirprol Tou uro_
rrvrlq.
Onog npoovorpdp0rlKe orqv Evoulra 79.g, opr<erdg
neproX6g orrlv unopovddo tou nopdyowo o Kot rrl
CTD r;g unogovddog o Splowor o, enogrl pe ro DNA
rou unoKrvrlr4. H rowono(qor1 ptog oerpdg drorpoperr_
t<c6v ouvorverLxov otrl.rlLoulLrrtv or ono(eg ctvoyvop(-
(owot on6 o).o6v(upo trie dro<popelxoUg nopdyoweg
o (dntog qo(veror o-Trlv ' , ., ,.) uno6r;i.rr;vrr o-i. :
[6ro9 o nopdyowog o npdner vo dpleror o, enoef *.
ro DNA. En[or1g unodrl].rrrvet 6l ot drogoperLxo( nop:-
yoweq o npdneL vo npoo6dvowor Ue nop6pioLo rpo::
mov nuprlvo rou ev(rjprou, dror c6me oL enLgoveLs:
ovoyvirtprorlg yro ro DNA ftov 6rorpoperrxrr:v nopoyo,,-
roJV vo rono0erotiwoL ire nop6;roro rp6no otore ,,,n
npoyporonorotiv ng xpforpreg enogdg Ue rov unoKrvr_
q r<owd ortg oMrlLoul(eg -35 xor -10.
Enrnp6o0ereg ev6e([erg drL o nopdyowog o(ypro ec
Xeror oe dpreorl eno<p4 Ue rov unor<rvr;r4 16oo orrl _35
6oo roL o-cr1 -10 ouvorvelrq oMrlLouX(o nporlLOov on:
unoro-roordoerq orrov nopdyowo o nou KoroordMo",
peroALd[erg orrq ouvorvelr<6g oMq].ou1(eg" Otov 1.ric
perd).).o[r1 oe Liro ouyxercprpr6,vrl 0dorl rou unoKrvr.lr-
nopepno6((er r;v ovoyv<rrpro4 on6 rqv RNA noLuprepc-
on Kor puo owroro0prorLrq ;rerdMo{4 orov nopdyowc
o enrpdner orrlv noi.upepdorl vo lprloLponorqoer ro f.lr-
toMoypr6vo unoxrvqq, q nro nr0ovrl eEnynol eivor or.
to olerrxd (e[yog Bdoeorv rou DNA dpleror o, enoer
pe ro optvolti nou sxer owrKqtooro0e(,
I uyr<p(oerg tlv oMq).ouyurtv opxertrtv Boxrq pto r<u_r,,
nopoydwov o un6.6er(ov ouwqpqpr6veg neproXdg o-rc
o70 Tou E.coli .,,, , ,, ,.', nou oMqenr6poriv dpreoc
pe unorrvqtdg" Aur6g or ev6e((eLg rexprqprot0qxov onc
rqv avoyvriprol Uroq xpuoroMLr<qq dopnq rou oLor,,-
([pou qg RNA no].upepdonq oe oUpnl.or<o pr, dvo r<.r.c-
ol-lo rou unor<rvqtr1. To Borcrqpro Thermus aquaticus xc.
Thermus thermophilus derlvouv notg or neprol6q npoc_
6eo4g oro DNA rou nopdyowo o mulrr.lvowor or ovr_
{dprqreg neproX6g orrg nptrrreivLr<dg neproleq 1 2
2,3-2.4, 3,0 r<oL 4.1 -4.2.
H Eu<dvo 19.'16 nopouord(er ro rprrlproro rou nopc-
yowo o nou exouv dgeoo pd),o mrlv ovoyvcilprqn roL
unorrvqrrl. Auq q eLrdvo de(pet q 6op4 -rou xrlpLcu
nopdyowo o oro n).o(oro rou oLoev(0prou. Atio Bpolec
rp4proro qg neproXqg 2 rr evo rprrlpo qg neproplq .1
(2.3, 2.4 KoL 4.2) eplowor o' enog4 ;re Bdoerg oro oror-
Xe(o -10 r<or -35, owroro(1org. H neproX4 1.2 rou nopo-
yovro o Spleror o' enorpr'1 pe rqv nsproX4 rol
unoxrv4rq r<o0odLnd on6 ro ororXe[o -10, evot rl neproxn
3.0 dpXerot Ue qv nepropl rou unoKrvrir4 prd).rg ovo6L-
rd rou oroLle(ou -10. Ko0etrr(o ond outdg trg neproldq
oX4porf(er Bpoldo rp4poro o-ti].u<og orqv npc,-rteivn
Ynogovd6o/
yov0ro
M6ye0og
(# aa)
Kord npoo6yyrorl
# tov unorw4rdrv
Aliflou1(o rou urtoKvlrn
nou ovoyvropi(eror
ILypo 70 (rpoD) 613 1000
ILypo 54 (rpoN) 477 5
IryiLo S (rpoS) 330 100
Irypo 32 (rpoH) 284 30
ILypo F (rpoF) 239 4A
I(ygo E (rpoE) 202 20
TTGACA-16 bp 6o9 1B bp-TATAAT
CTGGNA-6 bp-TTGCA
TTGACA-I6 bp eog 1B bp-TATAAT
CCCTTGAA-13 erog 15 bp-
CCCGATNT
CTAAA-1 5 bp-GCCGATAA
GAA-1 6 bp-YCTGA
Iiypo Fecl (fecl) 173 1-Z ?
Elx0NA 19.15 0t nopdyoweg o rou E coli ovoyvcop((ouv unorrvrlr6g pe drorpopelrrlg ouvorvelreg otrtrr1trouxreq.
524 KE0AAAI0 19 Meroyporprl oroug ilpoxopudrreg
Boorreg eworeg

H 6opq rou nopoyovro o oro yevrrorepo ntroioro rou o),oev(upou: otr).qtrenrdpooetq -1 0 rot -35 0 nopoYovrog o enerrel-
vrT6r Kgr or neptoXrlg rou ouvfleovror p' euetrrrroug ouvdereg. To oXrlpo 6XeL npoooplioorel ono: D. G. Vassylev, elal., Nature 417 (2002)'.
712-719. AoUrt on6 Protein Data Bank 1lW7 D, G Vassylev, elal.,Nature 417 2002).712-719
Mro r<puotoMu<q 6o;11 tou oLoev(0prou oe oripnlot<o 1.t'
5vo xldopro tou unoxtvrlni, oe ouv6uoopr6 pre nsrpdpto-
ro Ue unoKrvqt6g oro ono[o ot r<].ri:vot tou DNA roro-
oreudmqxov ri:me vo nept5Xouv oto(ptomo (eUyr1
(,,etepo6(xLrovo)), 66erqov 6rr o o70 6pXetot Kotd r<rjpro
L6yo o' enogq ;re Bdoerg orov xtrcbvo-prr1 expoyeio tou
ororXe(ou -10, oro 6reupup6vo orotXe(o -10 t<ot orrlv ile-
ptoXrl tou 6tolcoptorrl xot ouve{(et vo 6rotqpe( ow5g
rg eno<p6g t<ot getd to (etritrtypro rou DNA outtlg qg
neproXqq. Aur6 entBeBouilvet 6tt o nopdyowog o efuot
or1trrowx6g orrlv ow(6pooq onodtdtoEns FnEnS)
H Xprio'n o-ehroetdriv pro{Brov e(vot t<owr1 oe npo-
retveg nou ovoyvop((ouv oMql.ou{eg 6(r}'orvou DNA
(BI6ne Ev5qto 28.6). Apwo[5o nou on5louv pero{U
roug 3-4 06oetg Bpiot<owot ornv (6to nLeupd qg o-51't-
Koq Kor yt'out6 e(vot oe 05or1 vo 6p0ouv o' enoq4 pe
yenovrrd (e0yq Bdoeov. H ElKOl'lA't3'17 de(Xvet 6n r'
oprwo[5o rord prlxoq ptog n].eupdg tqg neptoplg 2.4
trlq o-6lrro9 5pXowot o' enogrl pe rg Bdoerg ortg 06-
oerg -12 5og -10 rnq -10 otrtrr1).ou{og tou unoxtvrprl.
H neptoXrl 2.3 potd(et pre npo:teiVeg nou npoo66-
vouv 1.rov6rtrrovo vouxleivtxd o(5o rt epnl,5retoL or4v
ow(6pooq rn[nq. Ot neptoXSg 2.1 ru 2.2 (ro nto ouv-
flpnudvo rpqgo tou nopdyowo o) egnl'6rowot oqv
otrLqtren(6poon Ue rov nuprlvo tou ev(ripou. Eu<d(erot
611 6tror or nopdyoweg o npoo66vowot onq (6eg ne-
ptoX6g rou nuprlvo trlg RNA no).uprepdoqg, on6re "ov-
royrr:v((owotD yto neptoptopr6vo Udpoq tou nup{vo
rnq.
I-1op6l o nopdyowog o 6XeL neptoldg nou ovoYvo-
pi(ouv ouyr<enpt;tdveg Bdoetg oro DNA tou unortv4trl,
q N-teLrr<q neptoXrl tou eerJOepou nopdyowo o (ne-
proXrl 1.1), evepyolwoq Log ouroovoorotrttrrl neptoXtl,
onorpUnret qv noptoXrl np6odeoqg oro DNA. M6vo
6rov q 6Lopr6p<ptr:or1 rou rpononotq0e( on6 11 orjvdeorl
rou ue rov nuprlvo tou ev([p-rou, prnope( o nopdyowog
o vo npoo6e0e( erdLr<d ortg oMqLouX(eg tou unortvqtrl
H oduvopr(o tou etrerjOepou nopdyowo o
v' ovoyvolp[oer oMqtroul(eg tou unortvqt4 dleL orlpo-
o(o: ov npoodevdrov oe unoKtvrlr6g u.tg e).e00epq uno-
provo6o 0o pnopoUoe v' ovome[].et rqv 6vop[q tqg
06on -13-12-11*10-9-8-7
ElK0$iA 19.17 T' oprvo[60 ot2.4 o-5trtro tou o70 6pXovrot o'
enoqn Ue ouyrsrprpdveg pooetq orov rodxonoulrtrd rtrtivo qg
-1 0 otrtrqtrouXiog rou unortvrlrfl.
19.tr0 [lo),treg l-leptoX6q qg RNA llotruUepoonq'EpXovrot Apeoo oe Enoqrl pe rov DNA YnoxLulnl 525

neproxeg flp6o0eonq oro DNA
tlH N-rdwr1 neptoXrl
npoo6Sver nepLoX6g
npdobeorlg DNA
mov ele00epo o
To DNA erronl(et qv
N-re)x1 neproXrl rirov
oXqgor[(eror ro
ouurroKo pre ro DNA
EIK0NA 19.18 To N-retrxd orpo rou o epno6(er Trq neprox6q
rp6o6s0nq DNA vo ouvoeeofv Ue To oTdxo Touq. Mdtrrq oxrlUoTr-
ore( 6v0 ovorXro otjuntroKo, ro N-rs,xd orpo ourlpeuor 20 A po-
rpud ror or 6Uo neproX6q rpdodeonq Tou DNA 0r6xouv 11 1uo ond
qv dtrr1 roro -15 A.
Ueroypoqnq on6 ro ol,o6v(ugo. H EK6vq 19,18 ono6(-
der g4qporrd qv oMoyri dropr6pgooqg rou nopdyov-
ro o rotd ro oxnporropr6 rou ovorXroU ouprnL6rou.
'Orov o nopdyowog o npoo66verqr orov nupr]vo
qg noLuprepdonq, n N-rellrq neproXrl ourlpe(ror -20 A
por<pud on6 rtg neproX5g np6o6eorlg rou DNA ror or
neproX6g np6o6eo1g rou DNA on5louv preto(ti roug
-15 A, nr0ovr6g rilore v' ononrloouv nro enlgrp<uoprdvrl
6topr6pgcoorl, rotd).Lr1l,q yro rnv enog4 pre ro DNA.
MeroLLd(erq e(re oqv oLL4LouX(o -10 efue mrlv -35
ovoor6tr.ouv rrlv np6o6eor1 oro DNA puog xotroBro;r5-
vrlg erdoXqq rou o70 Xcop(g rqv N-rel,u<ri neproXrj, yeyo-
v6g nou uno64l,6ver 6tr o o70 6pXeror rour6lpovo o'
eno<p4 Kor Ue trq 6rio d.)l.r1lou1(eg. Aur6 rorprd(er pre
rg nlqpotpop(eg on6 rqv rpuoroLllxrl 6oprl rou olo-
ev([pou (Etr6vo 19.16), orrlv ono(o e[vor oorpdg 6n o
nopdyowog o 61er gdMov enr;rrlxuoprdvrl Uopen, eKrer-
v5prevog oe -68 A rov 6Uo orpoec0v rou DNA.
MoLov5t 11 neproXq 1.1 rou nopdyowo o 6ev SXet
ovoLuOe[ pre r<puoro].troyporp(o, Brorpuorx6g prerp4oerg
qq ew6nrorlq rnq oro oLo5v(upro Kor oro ovotXr6 oUpr-
nloro unodql.<irvouv 6r oro eAe00epo olo6v(upo 11 N-
retur<{ neproX( (neproXri 1.1) Bp(oretor orov rewpr6
6(ou).o DNA rou ev(0prou, puproriprevq ouoroorrrd q 06-
orl nou 0o xoroldBer o unoKrvnr{q 6rov q4qprorrme(
ro oUpnloro rnq Ueroyporp{q (ElX0ilA 19.19}. Orov ro
oLo6v(upro oXqproiloer 6vo ovorXr6 oripntroxo oro DNA,
q N-reLtrrl nepropi rou nopdyowo o eKroTr((eror on6
rov r6pro 6(oulo. Iuvendlg, q ew6nroq qg oe q46or1
Ue rnv un6trotnr1 nporelvr1 e(vor notr0 euprerdB],4rr1. A].-
trd(er 6rov o nopdyowog o npoodeOe( oro rewptr6 6v-
(upo ror (ovd 6rov to oLoSv(upro npoo6e0e( oro DNA.
H 5fur<o rou DNA np6na vo trretorwq0e( nepfnou xo'rd
16 A on6 t4v opXxrl qg 05or1 nporeql5vou vu ero6),0er
mov xripro 6[ou].o tou DNA rr 6nemo vo perorrvrl0e(
(ovd 6ore vo enmpdr.fer oT o DNA vo ero6l0er oro 6(ou-
l,o t<otd ro oxrlporropr6 rou ovorXro6 ou;rnL6rou. H El-
(01{A 19.20 nopouord(er owrl q perorfur1or1, B}.5nowog
526 KE0A Al019 Meroypogrt mouq l-lporoputlreq
EIKONA 19.19 0 nopdyovtog o 6Xet gro enqrlxuop6vrl 0opr1 nou
exre(veror rord p1rog qg enrqoverog rtrlv unogov66rov Tou ttu-
prlvo 6rov oXrl;iouore( ro otro6v(ugo.
oe eyxdporo roprrl rotd pr4rog rou dlovo t4g 6).u<og
rou DNA.
l-loLot6repo enu<pcnoiloe 11 dnogq 6rr o nopdyowog
o efuor q povo6rrq unogovd6o rrlq RNA nolupepdorlg
nou npoo66veror orrlv nepropi rou unorwrlrq, I(pepo
yvrop(oupe 5rt or C-refurdq neptoyiq rov 6Uo unopov6-
6rov o pmope( entorlq vo no((ouv orlprowx6 p6lo orqv
enogf pre ro DNA rou unorwrlrq ;r5oco rqg np6o6eor;g
rouq oro ororle(o UP (BL6ne Ev6rqra 79.8). Enerdq or
oCTd ouv66owor XoLopd Ue rnv un6l,orn4 RNA nolupre-
pdon (FMne Eu<6vo 19.14), ro 5v(upro prnope( vo rprdoer
oe opxerd onopoKpuol,l5veg neproX6g evril e(vor ox6pr1
npoo6e6epr6vo qro ororye(q -10 Kot -35. 'Etor or oCTDs
nop6Xouv rwqr5q neprol6g yr' oM$.en(6ppooq pe psro-
ypogu<orig nopdyoweq npoodedepr5vouq oe 6r6gopeq
onoordoerg ovodxd rqq oger4pioq rnq Ueroyporp{q 616-
gopov unorwrpcilv (BMne lGgdloro 26,7o Onep6vto).
EIK0NA 19.20 To DNA opXtrd 6pXeror o' enorprl pe rov nopoyov-
ro o (po0 ror rov nuprlvo rou ev(Upou (yxpr). Merouve[rqr nro
po0rd p6oo orov nuprlvo rou ev(Upou yro v0 npoyporonouloer
enoq6g pe rrlv otrtr4tr0uX(o -10. M6)rg onetreuOeptoOe[ o o, ro etj-
poq Tou nepdoporog nou nept6Xer DNA ou(dver. Avor0ntooq pe
r4v d6eLo rov Macmillan Publishers Ltd: Nature, D. G. Vassylev, et
al., uol 417, pp. 712-719, copyright 2002. 0toroyporpfo pe qv
odero rou Shigeyuki Yokoyama, The University of Tokyo.
N-TetrrKn neproxn
U-

19.1 1 H Anorrjntoon Eivor Mi6o6oq YrlnAfq
AvdAuonq Vro To Xoporrnprop6 rurv
AAAnAenr6pdoeov RNA lloAupepdonq-
Y no xw nr i't xaq I €,v rx6repo, DNA-I'I p an ei' v thv
* 0r ouvorvenreq otrtrntrouXreq -35 rot -1 0 nopSXouv T0 nepro-
oorepo orlge[o enoqrlq yro qv RNA notrugepoorl 0T0v uIIoKt-
vnTrl.
= To orluelo enoqnq 9p(orovror rupfog oq pto ntreupo rou DNA.
H txov6r4ro qq RNA nouprepdo4q (ornv npoyport-
xdrqro, r<d0e nporeivqg) v' ovoyvorp((er to DNA pno-
ce[ vo XqpoKrnprore( pe onorUnooq (footprinting).
Vro oMqloul(o DNA npoo6e6epr5vrl oe Uto nptotelVrl
r6meror peputitq on5 pro evdovoux.tredoq nou enrG
0erqt oe (eltrrptoroIg grrrogodreoteptroUg 6eopotig
ldoo mo voux.treivrr6 o[6. Yn6 r<otdtr].r1].eg ouvOr1-
<eg, rdoe (e1oprm6g grj:ogodremeptr6g 6eopr6q
.rnope( vo ondoer oe rdnoro prdpto DNA. Ot O5oetg
:ou ndnrowor ovoyvtr;p((owot Xpqotpronoui:wog D-
A o4pooprSvo orov 5vo x).rivo, p6vo oro
:vo dr<po. [lp6retror yro rrlv (6to oppl nou
iraoppd(eror ornv oMqLoriXqoq rou DNA:
- peprr<r1 ndr,Uq ev6g telrrd oqpoopr6vou
*op(ou oe Uro eunoOrl 06o11 dqpuoupye( 5vo
doopro povo6u<oU prqxouq.
H 6e(Xvet 6tt petd rrlv Korep-
.'ooio pre voux.tredoq to onoop6vo r<i.doproro
)NA pnope( vo 6toltoproroUv ovdloyo pe to
*r'lKoq rouq pie rlLer<rpog6pnon oe nr1rr1,
(o0e x).oopro nou 6Lotr;pe( 6vo orll.rooprdvo
:r<po nopdyet pto po6revepVn (rr:vn. H 06ot1
:1q (rrrvqg owtororle( mov opt0pr6 torv Bd-
trocrJv oro xi.dopo. To Bpoxurepo r^doporo
*erorrvo0wot roXUrepo, 6ror q on6moorl
:16 ro orlproop5vo drpo unoloy[(etor on6
:c xdrro pr5pog qg nnKqq,
Iro e).eU0epo DNA, r<d0e eunoOnq 6e-
:-rog ondet roul.dltorov oe rdnoto p6pto. H
=L<dvo 19.20 6e(Xver 61 6rov to DNA Bpi
l(rror or oripntror<o Ue Uro npore[vq, ot 05-
leiq nou roLUnrovror on6 tqv npotelvrl
.cccpuldooowor ond qv n6Qr1, 'Etot, 6tov
5.ro owLdpdoerg rp6(ouv nopdl,).q).o -dvo
H onorunool ovoyvtrtpi(eL rtg 0e-
-.r-q onou npood6vovrot npoTelvsq oro DNA ono
-,' npooroo(o nou nop6Xouv rlvovrL qg 6qgtoup-
, .rc evropcirv.
DNA prdprupog 1op(g ropr(o np<oreTwl ru 6vo nerpogqn-
16 prfupro pre pr6pro DNA npoo6e6e;rdvo mqv nporreivr;
0o 64;uoupyrl0e( dvo XopoKrnprorrr6 npdruno. 'Oruv
gro npoodedepdvrl nptoretvq ovoore(Aer rqv np6oBoo4
qg vouri,edonq oro DNA, or 6eoiro( orryv npoo6e6e;r6-
vr1 oMqAoulfu 6e ondve oro rretpapaut<6 1t$pa Ket ctu-
t6 to t1t4pa rqq ftnKrrlq 6ev nepdya o4paopiva xid-
oparu DNA.
Iro prdptupo, onder r<dOe 6eog6g dqpnoupytitwog
pro rl,(poro (ovrirv, orrlv ono[o x60e Bdoq owtnpooo-
neJerot on6 gro (c0vq. Irqv ex6vo, undpXouv 31 Bd-
oerq. Iro npoororeuUdvo xtrdopro, 6eopro( 6e ondve
orrlv neproXri 6nou e[vot npoo6e6etrr6w1 11 npcoteivq,
on6re 6e dr1;uoupyoriwor (c0veg nou owtnpoooneUouv
rl6oproro tcov ow[morlrov preye0c0v. H onouo(o turv
(ovrrlv 9-18 orrlv eu<6vo rou'ronoe( ptto 06oq np6o'6e-
o4g q ono(o roltjmet rrlv neproXtj nou Bp[oretot 9-18
Bdoerg on6 ro or1;roopdvo drpo tou DNA. Iuyrpfuov-
roq rrl Lop(6o tou ;rdprupo Kor rnv netpogotu<r1 op(-
6o pe pro ovt[6poor1 o].].q].orixnong nou rpdlet
nopdtrtrr1tro, prnope( vo "6tqpoore6, dpeoo q owforotXq
oMrlloul[o, ovoyvrop((owog 6ror tr1 vou t<].eottdu<rj o],-
Lql,oufo qg 06o119 np6o6eo4g.
'Oncog npoovo<p6p0r1r<e (BL6ne Ex6vo 19.13), q R-
NA nol.uprepdo4 npoo6Sverot srrlv neptoXti rou unoKt-
.AJffinffihnffih
I*stu@htr@
+ /V/Va/1.
+ /-,/,/VV.
}a
b
+ .ra/./vv1.
*ry+ /L
/ HAEKTPOOOPHIH
DNA olgoogdvo oro / Ilto gorPtd on6
dxpo rou evrjg rtrtivou m ffi- ro olpoopdvo
ry ffi drpo
1ETpAMATTKH ffi ffi 1HKTH
NHKTH ffi ffi MAPWPA:
ffi To DNAnou 6ev
To orlpntroro RNA rduUepdonq-
unoKrvnrn ddXerot peprxq en(Oeoq
on6 qv DNAdoq I
0r (tiveg nou
onouord(ouv
ovoyvrop(ouv
06oerg
npdoDeonq
Anopovrrlorl rou DNA
xor onoDrorolrl oe
lrovoug rtr6voug
ffi eivoL npoodedepdvo
ffi sqynougepoorl
ffi eXer (tiveq oe Oeoerq
ffi fiou owrororxouv oro
:::: onoorpo Kooe deol]ou
llro rovro oro
orll-roouevo oxpo
' ; :: H Anorunorq e[vor M50o6og Ytpllrlg Avoluqg yro Xoporqpto;.ro Atrtrqtrentdpooeov RNA llotruuepoolq-Ynoxtvrlq xct Dl',]i-': *..., * , 527
Boorxeg evvorcg

v4q on6 -55 Scog +20. To oqpre(o oro ono[o q RNA no-
Lupepdorl.6pXeror oe dpeorl enoqrj Ue rov unoxl4rrl
rquronororiwor Ue rponono(qor1 rrlg reyyu<rjg rnq ono-
t0n<oor1 g, 1p4 orprono uirwog owr6 poonl pro no u rpono-
notoUv ouyrerptpr6veg Bdoerg oro orjprnl,or<o RNA
noLuprepdo4g-unor<rvqri. Me ouq qv npooriyyron, ro
ne(po;ro ;rnope( vo ercrel,eore( ge 6tio rp6noug:
o To DNA pnope( vo rpononotrlOef nprv npoodeOe( 4
RNA nol,upepdorl. Ie ourrl qv nep(nrooq, ov 11
rponono(qo4 egnod(oer qv np6odeoq r4g RNA no-
).upepdoqg, rouronomfuor trlo Bdo4 6nou q enoErj
efuor onopofurlq.
o To o0unLor<o RNA notru;repdoqg-DNA pnope( vo
rpononor4Oe(, on6te ro np6runo rov npoororeu-
pr6vrov (alvcbv ouyr<p(verot ge ro ovr(morxo rou
eLerlOepou DNA r<ot rou 1-trl rpononorqpr6vou oup-
nl6xou. Kdnoreg (6veg e(ogov((owor, rouronor6v-
rog 6tor 06cerg ottg ono(eg ro 6v(upro rlXer
npoororeUoer rov unoKtvrlfi on6 tponono(r1or1. iL-
Leq (6veg 6Xouv ou[r1pr6vq 6woor1, towonorciwog
O5oerg orrg ono(eg to DNA np6na vo 6roqpr10e( oe
1.rro orepeo6op4 mqv ono(o e(vor neproo6repo
erre0erp6vo orov nopdyowo n5Qqg.
Aurdg ot otri,oy5q or4v euoroOqo[o onord].utfov r1
yeolprerp(o rou oupnL6rou, 6ncog ouvotp((eror orqv El-
K0l,lA 19.22 yro 5vo runx6 unorrv4rri. Or neprol5g oro -
35 ror oro -10 nepr6Xouv to neproo6tepo on5 -ro
o4pe(o enorpfg yro ro 6v(upo. MSoo oe our6g trg ne-
proXdg, or (6teg oprd6eg O5oeov re[vouv'16oo v' ono-
rp5nouv rrlv np5o6eon qv 61ouv nporlyoug6vrog
tpononot4Oe( 6oo xor ov eprrpov((ouv ou[qpdvrl ri prer-
cr:pdvq euoro04o[o orrlv rpoTronoirlorl trrerd rqv np6o-
6eoq. To orlpefo enogqg 6e oupn(nrouv n],ripog pre
O5oerg prero,trtrd(eov: ouprBo(vouv ndwcog orrlv (6ro ne-
proproprdv4 neproX4.
A[r(er vo orlpreuoOe( 6rt or (6reg 06oeLq oe 6rorpopen-
roUg unoxlqr5g nopdXouv noMd on6 ro orlpre(o eno-
rpr'19, 6oro Kr qv rl Bdoq nou Bpioreror oe Uro ouyKe-
xprpr6vr1 06oq 6rog5per. Aur6 unodrlAdrvet 6n undpler
rivog r<otv6g ;rr1Xovtotrr6g yro rnv np6o6eo4 qg RNA
noLuprepdoqg, nop6rr 11 ow(6pooq 6ev e(optdror on6
r1v nopouo(o ouyrerprprSvrov Bdoeov oe optopr6vo on5
ro orlpre[o enoqrjg. Aw6 ro ;row5].o e[4ye( yror( xd-
noro on6 ro o4pe(o eno<pr1g 6ev e(vor 06oerg peroMd-
[errlv. Enrn].6ov, 6e Bp(or<erot r<dOe prerd].],o(r1 oe
oqpe(o enorp{g: or preroMd(erg ;rnope( vo enrlped(ouv
q yettovu<q neproXrl 1o:p(g mqv npoygolr6rnro vq
Splowor o' eno<pri pre ro 5v(upro.
E[vor r6ro[repo orlpowx6 6n ro nepdproro nou oKo-
LouOo0v q orporrlyrxrl rng npo-rpononoqorlg rouro-
nororiv Odoetg or4v (6to nepropl Trou npoorore[eror
on6 ro 5v(upro Svowt enqK6Lou0qg rponono(qor1g. Au-
rd to 60o nepdgoro perporiv 6ro<poperu<d npdyproro.
H npo-ponono(r1oq ovoyvop((er dleg ttg O5oerg nou
npdner v' ovoyvcop(oer ro 6v(upro yro vo npoodeOe( oro
DNA. To netpdgoro npooroo(og ovoyvrop((ouv 6Leg rg
O5oerq nou npdygort Splowor o' eno<pq oro 6rprep5g
otipnloro. Ot npooroteu;r5veg 06oerg nepil.oprBdvouv
6leg ug 06oerg ovoyvc0proqq r<or peprrdg enrnpdoOereg
05oerg, yeyov6g nou uno6rlLtiver 6rr ro 5v(upro npcbto
ovoyvcop[(et puo oprd6o Bdoecov onopo(r4tov yro rnv
"npooye(oon) Tou Kr Snemo enercrefuet ro oqpe(o eno-
grlg rou o' enrnp6oOereg Bdoetg.
H neproXrlrou DNA nou e(vor (eruLrypdvq oro 6rpe-
p6g oUpn).oxo prnope( vo rouronorlOe( d;reoo pe Bdon
rrq Xqpu<5q oMoy6g orq 6ro0eo4-r6tr1rd qg. 'Orov 6ro-
lcoprororiv or x.trri:vor rou DNA, or o(euydpcoreq Bdoerg
yfuowor eud.troreg oe owtdpoorrlpro nou 6e prnopotiv
vq rrq npooeyy(oouv or4 6rntr4 5Lu<o. T6roto neLpd;roro
nepiloprBdvouv 06oerg pero[ri -9 xoL +3 orrlv opXu<r] ov-
ildpooq rri[nq. I-r'qur6 11 neproXrl nou [eru]lyeror rord
rqv 6vop[4 neprAopBdver ro 6elr6 dxpo rrlg oMq].ou-
X(oq -10 ru ercre(veror pr6Xpt pr6).rg prerd r4v oEerlp(o.
Av r<orrd[oupre rpro6tdmoro ro orlpre(o enocprlq
qvodrKd rqg oLLr'1louX(oq -10 dronrmrirvoupe 6tr Bpi-
oKowor 6Lo orq puo nLeupd rou DNA. Aur6 <pofveror
oro xdrcrr Srdypogpo qg Eu<6vog 19.22, oto ono(o ro
EIKSNA 19.U2 Mto ntreupo rou unoxtvlrrl nepL6Xer
to orlpe(o enoqlg yro ro RNA.
Atrtrr1trouX(o -35
t
ALtrnlouxio -1 0
J+ t,I, ,1. l.
I lt^,t+Iitl, I K06rKoIornrtK{iq
t Ktrtivoq
lxxxxllu.IffiKtrdlvog erpoyeio
To neprooorepo orlge(o enoqlq pploxovror oq gto ntreupo rou DNA
Tpononouloerq nou egno0[(ouv r1v npdodeorl qg RNA notrugepoorlg
06oerg rinou 1 RNA notrugepoorl npoororeuet ond qv rpononoiqorl
l4ero),tro(erg nou xoropyouv 11 perdlvouv qv evepyriqro rou un0Krvlrr1
J
+
0
ti
ililil1
528 KEOAAAI0 19 Meroyporprl oroug l-lpoxoputilreg

: ^Joro enoglq onpercovowor oe pro drnArq dtrrro nou
j 1.,,'rror ond q prLo nLeupd. To neproodtepo Bp(or<ov-
:trr orov r<ovo prq-er<pioye(o. Aur6g oL Bdoetg (or,rg ovo-
.,ropi(owor r<otd tov opXrr6 olqpotropr6 evdg xLer-
:.or-i 6Lpepo[g oupnLdr<ou. Aw6 0o enSrpene orqv R-
', - .rouprepooq vo n).rloLdoer to DNA on6 14 pro ntreu-
- r (or v' ovcryviop(oer ourr; rqv dQrl tou DNA.
(o0rig [errvd ro (erritrLypro tou DNA, ;rnopef v' ovo-
.*pLoro[v r<or dtr].eg O5oetg nou opXrr<d Bp(orowot
--',, oMrl nl.eupd tou DNA.
| 9.1 2 Or AAAnAenr6pdoerq Mero(U rou
llopdyovro o Kor rou llupfivo Tnq RNA
IloAupepdonq AA deouv KoTd Tn Aroguyfi
TOU YnOKIVNTfi
n ','.0 n00t0Xl rou o rorotroppovet ro dtoutro e(o6ou rou RNA
.r. lrprnrr vo perouvl0el ylo vo dteurotruver q o[v0eo1 rou
t.i
. - ,.nTUXe[q evop[erg ouvq0tog ouppo[vouv nporou ro ev(upo
,,i-porlorr evo npoygortro o0gntroro enLprlruvolg.
" I .opoyovroq o ouvnOoq onetreu0epdrveror ono qv RNA
:,,tpepoo1 potrtq q veoouvre0elpdvq otruo(60 RNA tptooet
:: plxog -'10 nt.
- rA noLupepdoq np6neL vo ouprBrBdoer rrg ovdy-
..: !'ro dvop[q pe rtg ovdyxeq yro enrprnruvon. l-1pco-
-: , o 6(ouLoq Sroguyqg rou RNA ornv
--:iuolx6rqto e(vot r<oreArlprpr6vog on6 rprlpo tou
-r:ryowo o, ro ouv66rq (linker) nou ouv6Set rtq ne-
- r ,.3Q 3 r<or 4. 'Eror, 11 6to<puyr1 rou unoKtvrlrl np5net
: .oorloprBdver ovodLo-rdro(q rou nopdyowo o,
.- cri(owdq rov on6 ro d(outro e[66ou tou RNA
- -t vo prnope( vo ouvexrore( 4 otlv0eoq tou RNA.
' : - .3oov, q 6vop[r1 onone[ orevq np6o6eor1 p6vo oe
- -. <a<prpdveg oMq).ouX(eg (unor<tvr1r5g), evcb q enr-
--,-;ol onotre( np6o6eor; oe dreg Ttq oMrlLouXieg
- - - :o ouvowrloer to 5v(upo Kord tq preroypogtl. H
6e(Xvet nurg enil.Uetot out6 to 6il.qprpro
-.r* rlq owtorpemrlq np6o6eor1 tou nopdyowo o
- :, rupqvo rou ev(Upou. Oncog npoovogSp0lKe, ro
. --ic ouvrl0og u<p(ororor r<rirLoug oventtuXtov
- ::ieov xord q 6Lepyoo[o r4g 6to<puy4g on6 tov
--:..'rlll. Ev6oto oupBo(vouv outo( ot oventruXe(g
. -- ' :., ro dv(upro 6e perorrvekor ndv<o oro expoye(o,
: -' : -oMov rpoBd ro npdlto ).tyo vour<i,eor(6ro tou
. :::5Lr<oU DNA, npoBdl).owog ouroUg roug provo6g
. -- ,o,Jq novco orqv enrgdverd rou, oe pto dtepyoo(o
-:rrl orq <paoqoq tou DNA - DNA scrunching". Ztn
r- ,.
'.aro.
p' 6vov prrlXovtopr6 nou 6ev SXer onooorp4vr-
, . .r.r1pcog, to 6v(upo dro<peUyer on6 ouq q <pdor1
--, r','enrruxov r<rjr<.trrrlv r<ot nepvd orrl rpdorl Tnq ent-
_ -'- ,
rplyopo
I f novo
ww@w
",:1lk!'> / --*ru I
i U rpnvopo *
: ^m.:@E
i ,*oypnyopo I
i ^d!%'
:*-Ef*eB
! n*rypnyopo I
i#@r....... I
To revrprxri 6v(ugo
repgorl(er rr
oneheu0ep6veror
EIK0NA 19.23 0 nopdyowoq o ror o nuprlvoq rou ev(ilgou ovo-
ruxtrtirvovror oe dtorpopenro oqpeio qg Ueruypoqnq.
Molov6rt q oneLeu0Spoorl rou nopdyowo o on6
to o0pmtroro 6ev e(vor onopokrlrrl yto q 6roguyi on6
tov unorwqr], 11 drdoroorl tou nopdyowo o on6 tov
nuprlvo rou ev(titrrou ouvrlOoq ouprBo(ver rout5lpovo 11
Lryo perd rq 6roquyrl. O nopdyowog o e(vor oe nep(o-
oero or o16o4 pre qv RNA nolupepdoq, ouvencig 11
oneLeuO6pcoon rou on6 ro oLo6v(u;ro 6e y(verot
onL6g yro vo ro xdver 6ro06otpro yrq Xp{on o' enrnp6-
o0ero owlypoqo rou otroev(riprou. Irqv npoy;rorr6q-
ro, ot nopdyoweq o owoyrov((owol yro neproproptdvo
owfuporpo rou nupfvo qg RNA noLuprepdo4q, yeyo-
v6g nou 6(ver q 6uvot6rqro v' oMd[er ro npSruno t4g
peroyporp4g (BI6ne Ev1trTra 1 9.1 9).
O nuprlvoq tou ev(6;rou (<,xewpx6 tiv(upto", core
enzyme) cro rprprep6g oriprntroxo (nepd.o;tBdvet DNA,
veoouweOery6vo RNA r<or RNA noLuprepdor;) ouorool-
rd e(vor "x.l.et6ofr5vog"
pr6Xpr vo o).oxlr1po0e( 11 enrprl-
KUVon. 'On<oq 0o 6oripre, outl 11 e[6].t(q npor0met ev
;rdper on6 rov rp6no Ue rov ono(o ro 6v(upro neprKu-
xl6ver to DNA ru ev p6per on6'rqv of[non qg ouyyd-
verdg rou yro ro orl;rntroro Ue ro veoouwe0ettrr6vo
RNA.
KevTprKo 6v(uUo
ono0lreupevo
oro DNA
0 nopdyovroq o
ouv06eroL pe ro
revrprxd 6v(upo
To otrodv(upo
perorrve[ror
npog Touq
unorLvlr6g
To revrptxrl ev(u;ro
ouv06ret RNA
3.rtr,rltrenrdpooergMero[0rou[lopoyovrooKorToutluprlvorrlqRNAtlotruUepoonqAtrtro(ouvKoroqAroquyl:0i-'.].,.,-.- 529
Booxfg eworeg
a

To gdppor<o prgopnu<(v4 (UdLoq rrlg orroy6verog
rou owrBrorrroU prgol.rur<(vq) epno6((er q preroypo<pq
on6 q Bor,m4pror<r1 RNA noluprepdoq. E(vor ro rUpto
owrBtoru<6 evow(ov qq gupror(coonq. H rpucroMrr{
6opr1 qg RNA noluprepdonq oe o0gnl,or<o Ue rn pt-
<poprnx[vr1 e[4ye( q 6pdor1 rou cpopprdrou: n prqognr-
r[v4 npoo6Sveror o' 5vo Ori],or<o rqg B unopovddog,
>12 A prorpud on6 ro evepy6 r6wpo, oe ;uo 05o4
6nou nopoxrr:trrJer q 6to6popuj rou enrgnKuv6gevou
RNA. Epno6((owoq r1v endrrooq qg oLuo(6og tou
RNA ndpo on6 2-3 vouxl,eor(6to, 6tor5nter rrl Uero-
Ypoen.
19..I3 H KpuoroAArrrl Aopri Yno6errv0er 'Evo
MovrdAo yro Tn Merorivnon Tou Ev(0pou
e To DNA perorrve[rol dropeoou evog droutrou orrlv RNA notru-
gepooq ror npoygoronore( pto onoropq orpotprl oro evepy6
r6vrpo.
x Atrtroydq oq dropoptptrlorl ouyxerprpevov eudtrrrrtov e[oprr1-
porov p6oo oro ev(upo etreyXouv qv e[oodo rrov vourtrrol-
d[ov oro evepyo xevrpo.
Me q droLeUrovoq r1g xpuoro).).rx4g 6ogrjg rov ev-
(tipuov on6 Borcrr;pro r<or (tipre oe oUpnloro pre NTP
ror/n to DNA, onorcr4oope noMdg nnpogop(eq oxe-
nrd ;.re rq 6o;14 r<or 14 l.enoupy(o rrlg RNA nolupe-
pdonq rord rrlv enrprrlruvoq. H Boxrqpror<r1 RNA
noLugepdorl 6Xer 6romdoerg -90 x 95 x 160 A, evo-l or
s0A
RNA noLupepdoeg rov opXo(rov Kot rov europu<orrirv
e(vor eloEpcbq preyol0repeq, rup(rog ).6yo np6o0e-
rov rgqgdrcov o1.uvo(6cov r<o/r1 unopovd6ov rou ev-
ton((owor orqv neptg6pero rou ev(ri;rou. MoLorowo,
ot nuprlveg rov ev(U;uov prorpd(owor 61r pr6vo r<owr1
6opr{, mrlv ono(o undpler 6voq "6[outroq" eripoug
-25 A nou meyd(er ro DNA, oMd r<or r<ow6 pr;Xovt-
qr6 yto npooOrjrq voux.l.eou6[ov.
'Evo gowriLo owo( rou 6roUl.ou orq Bor,irrlprorcg
RNA nolupepdo4 ono6(6eror orrlv EIK0HA 19.24. H oG
Loro ouyr<pore( -17 bp DNA. Ie ouv6uoopr6 pre to -13
nt DNA nou Bp(orowor orrlv neproXq rou evepyof x6v-
rpou rou ev(0pou, owtorotloriv ornv npoororeup6v4
neproXrl tru'p<oug -30-35 nt rou noporqpe(ror oe on(>
tunc0oerg tou oupnL6r<ou enrprlruvoqg. H o0Aoro
enevdUeror pre Oeru<d gopr(o, Xdprl oro ono(o oM4Ae-
nrdpd pe rtg n^erupopvrlnr6g qrr:orpoprxdg opd6eg
tou DNA. To rorolurrx6 r6wpo (catalytic site) oX4pro-
r((etor on6 pro o1rotrrr1 ovdgeoo mrg 6rio ;reydleg
unopovd6eg rt otXl.roLor((er oro (oqy6vro" rou to D
NA t<oOcbq ero6pleror mqv RNA noi,uprepdo4. H RNA
noLupepdo4 neprBdLLer ro DNA Kor oro evepy6 rdv-
rpo rnq nepdXer 5vo 6vo rorolurx6 r6v Mg2+. To D|.lA
ouyrporefuor orq 05oq tou on6 rov r<o0odrrd ouvde-
rrjpo (clamp), evoMorcrtrq ovol.roo[o yro 6vo on6 to
ooy6vro. H EIK0HA 19.25 onerrov((er q orporp{ 90o nou
ovoyr<d(eror vo npqyporonorfoer ro DNA orrlv e(oo6o
rou evepyo0 r<5wpou elon(og ev6g 6rnlovoU npcoreivr-
ro0 re[Xoug. To pr{r<og tou uBprdu<o[ RNA neprop[G-
ror on5 6vo 6).Lo npo:reivu<5 eprn66ro, yvom6 og
rdlu;r;ro (lid). To vourctreor(dro ntoreUerot 6u ero6p
Xowor oro evepy6 r<5wpo on6 xdtco, prSoo rou 6eure-
poyevorig 6torlLou (mqv RNA nol,uprepdoq rnq E0pnS
ovoprd(eror "n6poS", pore). H rpuool(6o nepiloprB6va
8-9 bp uBpr6(ou preroyporptxrl DNA-RNA. To xdLuppro
Movr6tro nou 6e[Xver q Dogrl qg RNA no-
truUepoolq 6top6oou rou rup(ou 6roul,ou. 0r uno;-rovodec
rordrronorouvror Xpopolro coq e[lq: B po(, 0, ruovo, o
npootvo, oll, r[rprvo, o, 16rrrvo. Ipoooplroogrlvo onc
K.M. Geszvain rorR. Landick (ed. N. P. Higgins). The bacte-
rial Chromosome. American Society for Microbiology. 2004
FcTp,,l
530 KEOAAAIO 19 Meroyporprl oroug llpoxopudrreg
Boorxeg ewoleg

Avo6rxrl DNA
'Evo
nporeivtro re(Xoq e[ovoyro(et To DNA v0
:,poqr1 oro evtpYo K6vTpo. Nouxtreor[dto pnopel vo
oro evepy6 rSvrpo 0toireoou evog nopou oqv npto-
:.-ir rLq BdoeLg DNA rot RNA oro 5vo drpo tou
: :- tpl6ne Etr6vo '19,25) r<ot n Fdon tou DNA
l:.cxeroL mo dMo dt<po rou uBpt6(ou ttvd(etot
llpoor(0erot rowo0pu Bdoq xot ondve 0eogo(
'R11i-:
lrt;ii,l
lltr;111.
To 6v(uuo gerortvefiot rot ovodrlptoupye( touq 6eoU00q
il ;, r. ll
ll:t,i,
,
-:- - "-..... --,
;:.r.i;il+1li.it:; ',
Noudeot(6tq
KtuIon tou
ev(Upou
ir I
!1,_!
2To DNA
Ero6pXoTot
oro ooydvto

: .: e[o yto vo enttpSqet to (euydptol-to Ue ro et-
: -o'vo NTP.
^N
r A . ,-^-^i -'(E* h -^avia rr,lr Ftwr,lnt-
:1..q ro DNA unoore(qEn, n rpox(q tcov [elcopt-
, - . ,.i.!vcDv U6oo crro 5v(upo dev neptop((erot nlSov
:-: :- 5uoKoUqr(o 'rnq 6tnI(q 5trxog, enrp6nowog oto
- ,: . : rpoYuoronotrloet rq orpoqn 90o oro evepy6 r6v-
-: : I:.c1q, ouprBo(vet ptto peyoLr1 oMoyrl dtopt6ptpoorlq
-- l : ro dv(upto nou nepAoptBdvet to "ouvderrlpo" o
EIK0NA 19.26 H Ueroxivnon rrlq notrupepdorlg evdg vourhei-
vrKorj oE6oq rpoUn0e6Tet Oporion Kot 0v06rlUtoupY(0 6eop6v
pe vourtreotldto oe oT00ep6q e6oetq oXettrd Ue Tn Oogn rou
ev(rlgou. To vourleoridto oe our6g rtg 06oetg d0d(ouv Kdee
Qopd rou To 6v(uUo UeToKtvs(Tot KoTd Uto pdoq ndvto oT0 eK'
uoYe(0.
vouKleivtKov o[6ov nou Bp(or<owot prto 06oq nto rdrco
Kord ilnKoq tou t<.trtilvou. To ;reoo[o n].o(oto 6e(1vet 6tt
ow6 np6net vo onpo(vfl 6tt pr6tr'tg npooreOef n pdon
Kot nprv ro 6v(uUo petorwqoe[ oe oxSon pe to vour]'ei-
vrx6 o(ti, ot enogdq nou 61ouv 6r1prtoupy40e( pe er6x6g
06oetq np6net vo ondoouv 6tot cilore vo Unope( vo Eo-
vodrlpuoupyl0oriv pe rtq Bdoetg nou Kqto^,ogBdvouv
ng O5oetq our6g prerd q ;retor<[vqor1'
Optop6veg rpuoroLLtt<69 6opdq rrlg RNA nolupe'
p6onq npooq6pouv oqpowtx6g ev6e([erq yto to nc0q
ro 6v(upro 6rotqpe( rtq enog5g pe ro un6orpopd tou
ro0drg ondet Kot [ovo6qproupye( 6eoprofg qrn 6tepYo-
ofq rou r<[x].ou npoo0rixqq vour],eott6(ov rt evcir uq(-
orsrqt petot6ntorl. 'Evo eudNrro t;r(trro nou rol'efuqt
0qtud nupo66qonq (trigger loop) qo(verot 6tt 6ev efuot
mulo;t6vo Tlptv rl'lv npoo0(xr1 tov vourl,eort6(tov, dr-
d muXrivetot pr6l,tq to NTP etodLOet oro evepy6 x6v-
rpo. 'Orov olorlqpo0e( o oXrl;ronopr6g tou deoproU rot
r1 pretor6ntoq tou ev(rigou orrlv en6prevq 06o11, 11 Or1Ltd
nupo66tr1or1g (erutr'iyerot rot n6Lt, 6roq-r1 yto tov en6-
prevo r0xl.o, 'Etot, 1.tto Sopu<r1 oM.oyrl orrl OqLtd nupo-
66rr1oqg ouwov((et Tr1v orotrouo(o tcrrv oupB6wov rqq
rot6).uor1q.
:: cXqprot((et 6vo on6 to ooy6vto tou ev(fpou nou
-ouv ro xo0odtr6 DNA orn 05or1 tou.
" : :nd to 6tl.qpr;roto roOe notruprepdonq voux.e'i-
:ieoq e[vot 6tt to 6v(upo np6net gev vo 6qpt-
-:r,, oerxr5q eno<pdg pe to un6otpopo Kqt Ue ro
: . lMo en(or1g xot vo 6tor6r.l.tet out6g ttg eno<p5g
, : :.Q [ovooltlpot(oet oe rdOe r<0ro npooO4r<qg
-'r::16(ou. Ag ovoloytmo[pre trlv rotdorooq nou
. s3Qerot oTnv Mro notrupepdorl
*::onote( pto oerpd et6trrov enorpov pre ttq Bd-
: :,.l Bp(orowot oe ouyxerptp6veq O5oetq. l-tq no-
:...*c, q enogrl "1" dtlutouPYe(tot pe q Bdorl mo
:: :ou ou(ov6pevou x.l.<ovou, evco 11 enocptl "2" pt
- ::]r orov rLdlvo er<proye[o nou e(vot ougn].qptl1'to-
- . :: -. qv en6pevq B6oq nou 0o npomeOe(' Entoq-- . :: -. qv en6pevrl B6oq nou 0o npomeOe(' Entoq-
-: ..:lr, ocn6oo, 6rt ot pdoetq nou rotol'oprBdvouvl. el, wv
.: :rq 06oetg arouq r<Ltovoug ttov vout<LeivLxriv
- i: --, oMd(ouv r<dOe <popd nou npoor[0etot 6vo vou-
-: rovo.r ror to xdttrl nl,o(oto r1g etr<6vog 6e(lyouv
-. l.c rordoroorl: prto Bdo'r1 np6rettot vo npomeOe(
, :. :r(ov6;tevo xtrtrlvo. H dtotpopd e(vot 6tt mo rdttr;
- r l.c o oufov6pevog x}'tovog SXet ener<roOe( rord pto
: --- -l yeo-lpretp[o Kot rov 6rio ouprnL6rolv e(vot oxpt-
:-: - i6Lo, qMd oro xdto nLo(oto ot enorp6g "1" Kot
: :ooyUoronoro6wor pe Bdoetq tolv oLuo(6<ov tcov
l* il H Kpuorotrtrrrl Aoprl Ynooltrdrver'Evo Movr6tro yto q Meroxtvlon lou Et'[-*:- 531

19.1 4 Mlo lropornp6vn RNA l-loAupepdon
Mnopei vo lovopxioel
r Mr0 or0U0Tlp6vq RNA notrugepool pnopei vo {ovopXioer ng
pet0ypogrl xdpovrog ro RNA peroyporpo yLo vo Drlgroupyloer
6vo roLvorjpro 3' oxpo.
H RNA noAutrrepdoq npdner vo e[vor oe 05or1 vo Xerpr-
ore( rotomdosrq orrq ono(eg n Ueroypogq ovoor6).-
Leror. I-ro nopdderypro, qur6 prnope( vo ouprBe( 6rov
ro DNA SXer unoorei BIdBn. 'Evo np6runo oriorqpro
yro r6roeg neprordoerg nporUnrer ge 6ror<onri qg
enrpr{ruvoqg in vitro e(opc0wog 6vq on6 ro onopo(-
rqro np66popro vouxleor(6ro, lepropr6g nou enmpd-
ner rn gOopd rou dxpou rou RNA. 0o16oo, to [6ro
onorSLeopo 61er xou<dOe ouprBdv nou nporoLef ].d-
0og euOuypdppton rou 3' dr<pou rou RNA Fre ro evep-
y6 r<6wpo: r<dr lperd(eror yro vo lovoronoOerfoet ro
3'-OH rou veoouweOer;rdvou RNA oro evepy6 r<6wpo
6ror c6ore vo prnope( vo 5e10e[ en[Oeor1 on6 ro en6pre-
vo NTP Kor vo oxrlpor(oer rpologo6eoreprx6 6eog6. H
enoveu0uypdprgtorl ennuyldvetor pe ronq rou RNA
to dxpo d:qre vo rono0erqOe( orq ocoarrl 06oq yro
npoo0qr<r1 enrnp6oOerrr;v Bdoeolv.
Molov6rr 11 11 xonrl e(vot Spyo rqg (6ro9 rrlg RNA
noLupepdorlg, 6eye(peror nolri on6 enrroupxo0g no-
pdyoweg, noL0 6to6e6opr5voug Kor oro rp(o BroLoyrxd
Boofi.ero. Iro E coll undpXouv 6Uo'r6rotor nopdyov-
req, or GreA xor GreB. H RNA nol,upepdoq ll rov euKo-
purorrriv Xpno$onote( yro rov [6ro or<on5 rov TFllS. O
TFIIS 51er pu<pri opor6rrlro oq npoq rnv oLL4l,ou1(o
ror rq 6opr4 Ue rouq nopdyoweq Gre, otrld npoo66ve-
ror oro 6ro rprl;ro rou ev(rjprou, ro 6eurepoyevrl 6(ou-
).o (n6po) qg RNA no).uprepdo4g.
Or nopdyoweg Gre/TFllS enrp6nouv or4v noLupe-
pdoq vo x6rler prepu<d prBovouxLeot(6ro on6 ro 3'rcLt-
rd drpo rou RNA npoi6wog, ennp6nowog ritor mo
roro.l.urtr6 rdwpo qg RNA nol,upepdoqq vo eiloveu-
Ouypogrore( pre ro 3'-OH. Ko0Svog on6 roug nopdyov-
reg erodyer gto orev4 nporeiVrrrl neproXrl [orov TFIIS
e[vor gto r<op66],o geu6opyripou (zinc ribbon), oro Bo-
rrqproxd riv(upro 6vo onerpoer66g one(popro (coiled-
coll)l Bo0rd prdoo or4v RNA noi,u;repdor1, qrdvowog
noL6 rowd oro rorolurrr6 r<6wpo. A(o 6(wo oUrvo-
[5o or4v ropugrlrou nopdyowo nl,qord(ouv fio npo-
-reUov Korqluttx6 r6v ;royv4o(ou rou evepyori xdwpou
ennp6nowog rrlv e[oo6o ev6g 6ew6pou r6woq Uoyvq-
o(ou ror gerorp6nowog ftor q 06o4 rordluoqg oe pr-
Bovourtredoq.
Iuvomxd, 4 enrpqxuv6prev4 RNA noLuprepdoq riXer
rqv xov6trlto vo [eru],(yer Kor vo [ovoru].(yer ro DNA,
vo ouyKpors( roug 6toXroproprdvoug r<).c0voug tou DNA
ro0oig ror ro RNA (npoi6v), vo rorol,fer rqv npoo0rlxrl
prBovouxi.eon6(rov orqv ou[ov6gevq otruo[6o RNA, vo
enmrlpe( r4v np6o6o out4g qg ow(6poo4g Kor -Ue rrl
532 KE0A Al0 19 Metoyporprl oroug l-lpoxoputirreg
ouv6popr4 ev6q r1 60o enrxouprxcbv nopoy6wov- vo
enilUer npoBLriproro nou dqproupyo0wot on6 qv rorq
Ltycov nt on6 ro npoiSv RNA rot u1v enovdvop[n rnq
otivOeorlq.
19..l5 H BorrnploKn RNA t-toAupepdon
Teppori(er o€ AtoKprrdq @6oerq
* YnopXouv d[o ro[eLq otrtrr1trouXuirv reppologori. Exelveq nou
ovoyvopf(ovror povo ono qv [6ro rrlv RNA notrupepoorl Xo-
p[g rr1 oupperoXq rovevoq rurroprrorj nopoyovro, ouvrl0otc
ovotptlpovroL oq "evdoyevelg rep;louoreg,, Atrtreq 0n0rT0u,
pro rurroplrrl nporelvr1 nou ovopo(eror po (p) xor ovorpepov-
Tot trq <<TepUoTroreg e[oprc0pevor ono rrlv p".
* 0 evdoyevrlg repponopdg npouno06reL ovoyvtrlprorl ;rrog otr-
trqtrou1rog repportopou oro DNA nou roduonorer pto dogl
qouprerog oro RNA npoiov.
* To o1poro yro reppolopo pplorovror rupirrrg p6oo oe aAAq-
Aouyieg nou eyouv 1764 peroypagei ono qv RNA notruUepo-
o1 rr dror o repporropog pp[oreror un6 rov 5treyyo iou
erpoye(ou ro/r1 rou RNA npoidvtog nou geroypotper I notru-
uepoolt.
M6Lrg 11 RNA no).uprepdorl [er<rvrloer rrl pretoypoqq
t<ord pujr<og rou erpoye(ou, ouv06rer RNA, pr51pL vo
ouvowrloet prto o).).qLouX(o repporrorq (terminator) .
Ie out6 to orlpe(o, ro 5v(upo oro;lord vo npoo0drer
vour<).eor(6to orov ou{ov6pevo r}.rrlvo RNA, one}.eu-
Oepcbver ro oloxr1pcopr5,vo npoi6v xor 6rfororor ond
to erproye(o tou DNA. O reppouopr6g npouno0dreL
6rL SLor or 6eogo( udpoy6vou Trou ouyKporoJv ro
uBp(6ro RNA-DNA np5ner vo ondoouv ror getd vo [o-
vooxrlUorrore( ro 6(x).rrlvo DNA.
Meprrdg eop6q e[vor 6Uor<o].o vo npoo6rop(ooupe
q 05o4 tepprorLoprorj yro 6vo RNA nou dlet ouwe0e( oe
(orwov6 rlirropo, ener6q ro 3' dr<po rou prop(ou Unoper
vo 6toonoore( on6 pro 3' efovouxi.edorl r1 vo rone[ ond
pLo ev6ovouxedor1, lcop(g v' ogrloer rov6vo (Xvoq qq
05or1q dnou npoyprolr<d reppdttoe 11 RNA no).u;iepdorl:
out6 e(vor rdro(tepo rorvd oroug eur<opurrtreg. l-r'ourd
ro ).6yo, ot O5oeLg -reppolop.roU oulvd loporcrrlp((owoL
xo).0repo in vitro. C)016oo, 11 rrov6rr;ro rou ev(Uprou vo
reppot((er in vitro enrlped(eroL noLU ond nopog5rpoug
6norg q rowrxrl roltig xot q 0eppror<poo(o mqv ono(o
npoyporonorefuor r; ow(6poor1. 'Eror, o tepporLop6g oe
prro opLoprdvq 06,o1 in vitro 6ev onoreLe( on66er(q drr 11
o6v0eoq 0o reppolore( orrlv (6ro 06o11 Kor oro x0ropo
(in vivo). 0o16oo, 6rov ro (6ro 3' dr<po ovrXveUeror 16oo
in vivo ooo xor pe roOopd ouororLrd in vitro, our6 yevr-
rd Oeorpeftor utg Lcrluprl Svdetlq 6tr npdrenor yro rnv
ouOewrxrl 05or; tepprorLoproU,
Ot i.li,,..::. ., " Ko[ r'jir-:,:... i'. 'rrr oUVoqJ((oUvro 60o
r<rjpro lopor<rrlptmrr<d rcov evdoyevdrv repporror6v
Boomeq eworeg
Boorxeg eworeq

Ixe6ov otreq ot onopo(rrlreq
lrliltrouXieq yro tov repgotlopo
Ppiorovror otr1v fleptoxn
nou usroYpdQnKe
H rpoupr5ro oTo RNA
Unopei vo onotTtiTot
0t otrtrtltrouxieg DNA nou onotTolJvTot Yto Tov
-.:-lrLolto
$plorovrot ovootro Uq otrtrntrouxtog rou repportorrl,
, ,,-lorrouoq uroq QoupKer0q oro RNA unope( v0 elvot 0n0p0i
--tt
ntri nsic terminators). l-l ptrttov, ot ev6oYeve(q repUort-
: .q, or ono(ot e(vot ove[dprlrot on6 rov entrouptr6
nop6yovro po (rho factor), Yto vo oXqpot(oouv trl
1.-repotoyq doprrl rou RNA nou preroypd<petot onot-
:JV llo goupx6ro (hairpin) n).oUoto oe G+C. 'Ero4 11
-.,aypagn eoptatat an6 rc RNA npoi6v xat 5ev xa)o-
.*#.w
E*G
ffiiG
b#&
L&th
U}4F
$r$4
"6RqeAg. h*
wtrg6lE
h&r&
ilfl-E
ffi
ffi neproxrl iloriotq oe G-C oro gtoxo
ffiq
[Jels
il&T TMovoxtrovn
oend U
EIKONA 19.28 0r evdoyeve(g repponor6g nepiloppdvouv notr[v-
dpogeq neptox6g nou oXl;roti(ouv goupx6req Kot notKfiouv 0s
g4rog on6 7 etrrq 20 bp. H 6opr1 Utoxou-Ontrtdq nepiloUFdvet Uto
neproxn ntroUoto oe G-C nou oxo)rou0eltot on6 pto ostpd K0T0-
tro(nov U.
p(etu anLriq pe lleyyo n1q a&rlLouy{ag tou DNA xa-
td q peraypacpfl. 1o 6e0repo XopoKrnptsrr6 efuot pro
oetpd xotdtrotnov U (T oro DNA) nou orol'ou0o0v to
pr(o1o qg qoupr<6tog oMd nporlyo0wot qg npqYpm-
rrlq 05oqg repUortopoti. Ito yovtd(topto 'cou E. coli
undpXouv -1100 otrtrr1LouX(eg nou totptd(ouv pre owd
ro r<pttripto, yeyov6q nou unodqcilvet 6tt ndvo on6 to
gto6 pretdypqqo rou xurrdpou tepprot(owot o' ev6o-
yeve[g reppottm6g. Ot repportor6g nou efopr6wol
qn6 rov p (rho-dependent terminators) xoOop(owot
on6 t4v ovdyr<r1 npooOrixqg tou nop6yowo p in vitro.
En(oqg, pretoMd(etq 6e[lyouv 6rt o nopdyowog epnl'd-
Kerol fiov reppronop6 in vivo.
Ot tepgoror6g nox[trouv no]li og npoq rnv onotetre-
opcntr6rrpo. Og per6ypogo nopopiooqg (readthrough
transcripts) ovog6powot to pretdypogo nou 6e oropo-
roUv orovtepgonon(. (O 6poq nopoB(ooq lpqoqlonoef
rot ornv Evot7ta 25.14ytovo neptypdtpet rrlv xotooroNi
rodu<ov(cov tepporropori on6 dvo ptBooropdtto). En(oqq,
o rcpponop6g prnope( v' onotpone( on6 et6u<orig entKou-
pu<oUq nopdyoweg nou oMrltrenrdpo0v pre to RNA ro/r1
rqv RNA noluprepdorl, rotdoroorl Yvoorn oq qwlrep'
ponopr5g (antitermination). 'Eror, 6nog rot orqv nep(-
rrroorl qq 6vop[r1q r1 qg entptir<uvonq, o tepporrop6g
prnope( vo pu0ptore( oq Unxovtotl6g yro tov 6treyXo rqg
6rgpoorlg rrov yovt6(tov, 6noq 0o 6ori;re oro Ke<pdtroto
27, Zrpaqyxiq tav @dyuv.
YndpXouv Kqt dMot nopoM4Lrqro( preto[U 6vop[qg
Kor repportopro0. Kot or 6Uo 6epyoo(eg qnomo0v 0rt6-
orpro deoprdlv u6poy6vou (opXu<r1 ui[r1 tou DNA r<otd
qv 6vop[r1 rot 6tdorooq rou uBpt6(ou RNA-DNA orov
tepgottotrl6) r<ot pnope( vq XprlotUonoto6v entnp6o0e-
reg nproreiveq (nopdyoweq o, evepYonotqt6g, roro-
orol.e(g rot nopdyoweq p) nou otrq}'ent6po(v pe rov
nuprjvo tou ev(0;tou. Qor6oo, ev<ir 11 6vop(q Boo(etot
pr6vo orrlv oMr[ren(6poor1 prero[U qq RNA notru;repd-
onq Kor tou dtxtrtivou DNA, o reppottopr6g nepilopBd-
ver Kot ovoyvtilptorl onpdtov oro ;retdypotpo on6 qv
RNA nol.ugep6or1.
Ot orlpretor<6g pretoMd(etg nou trretcbvouv rllv onore-
treopour6qro rou tepgottogoti ouvrlOoq ouprBofuouv
p6oo oro p(oxo rqq rpoupr5rog ri ornv nLo0otq oe U
oMrllouX(o, 6v6et(4 Yto rrl onUoo(o outriv ttr:v oMq-
trouXtc0v oro prlxovtopt6 tou reppottopro0' To uBp(6to
RNA-DNA 6Xet preydtrrl ouvetogopd ortg duvdpretq nou
ouyrpototiv to otigntroro entpfruvoqg' Euvendrg, to
ondotpto tou uBpr6(ou 0o prnopoUoe v' qnooro0eponot-
qoet to o(prnLot<o en4.uiruvorlg, o6rlydrwog oe repUo-
rrotrt6. Atrtrntrenrdpdoetq rrlq <pouprdtog pe rrlv RNA
noluprepdoq ri 6uv6pretq nou oot<o0wot on6 ro crxnpo-
nopr6 qq goupr6tog ro0tirq ro RNA (enpoBdMer on6
ro 6(qutro e(6dou tou RNA, prnoporjv np6oxotpo vo eu-
Ouypoprpr(oouv pre trd0og rp6no to 3' dt<po tou RNA pe
ro evepy6 x5wpo tou ev(Uprou. Aurl q I'ov0oopr5u1 eu-
Ouypdprpuoq, oe ouv6uoo1.t6 pe to oouvti0toro oo0ev6g
uBp(6to RNA-DNA to ono(o oXrlpro{(etot on6 to (eUyr1
Fdoecov rU-dA RNA-DNA nou npox0mouv on6 ro 4tt1-
Uo Ue ro rqtdlotnq U, onosroOeponote[ to otipntroxo
enq.rlruvorlg.
19.15 H Bonrlptorrl RNA l-lotrupepdon Tepuqr(et oe Aroxptt6q 06oerq 5:B
lGUu
UA
GG
AA
I
H RNA notrupepdoq ror ro RNA onetreu0ep6vovrot
..'}..I. , [ }'^

Lewin γονίδια χ ( μέρος iii)

Lewin γονίδια χ ( μέρος iii)

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Lewin γονίδια χ ( μέρος iii)