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Ts 100 Tz 0 Tr 22 0 0 22 51.6761 725.8835 Tm [(Single-molecule)-333(tracking)-333(of)-333(mRNA)-333(exiting)-333(from)]TJ 0 -1.091 TD [(RNA)-333(polymerase)-260(II)]TJ /T1_1 1 Tf 9 0 0 9 51.6761 680.8835 Tm [(Joanna)-278(Andrecka*)]TJ 6.5 0 0 6 129.1881 683.8835 Tm (\206)Tj 9 0 0 9 132.8022 680.8835 Tm [(,)-333(Robert)-278(Lewis*)]TJ 6.5 0 0 6 198.3061 683.8835 Tm (\206)Tj 9 0 0 9 201.9202 680.8835 Tm [(,)-333(Florian)-278(Bru)]TJ 5.278 0 Td [(\250)-166(ckner*)]TJ 6.5 0 0 6 281.4294 683.8835 Tm (\207)Tj 9 0 0 9 285.0436 680.8835 Tm [(,)-333(Elisabeth)-278(Lehmann*)]TJ 6.5 0 0 6 375.5574 683.8835 Tm (\207)Tj 9 0 0 9 379.1716 680.8835 Tm [(,)-333(Patrick)-278(Cramer*)]TJ 6.5 0 0 6 451.1912 683.8835 Tm (\207)Tj 9 0 0 9 454.8053 680.8835 Tm [(,)-333(and)-278(Jens)-278(Michaelis*)]TJ 6.5 0 0 6 543.8054 683.8835 Tm (\206\247)Tj /T1_2 1 Tf 7 0 0 7 51.6761 664.3835 Tm [(*Munich)-278(Center)-278(for)-278(Integrated)-278(Protein)-278(Science)-278(\(CiPS)]TJ 5 0 0 5 221.268 666.6335 Tm (M)Tj 7 0 0 7 225.9886 664.3835 Tm [(\))-278(and)-278(Center)-278(for)-278(Nanoscience)-278(\(CeNS\),)-278(Ludwig-Maximilians-Universita)]TJ 0.1388 Tc 31.31 0 Td [(\250t)-139(M)139(u)]TJ 0 Tc 2.194 0 Td [(\250)-166(nchen,)-278(81377)-278(Munich,)]TJ -58.406 -1.143 Td (Germany;)Tj 5 0 0 5 85.1297 658.6335 Tm (\206)Tj 7 0 0 7 87.91 656.3835 Tm [(Department)-278(of)-278(Chemistry)-278(and)-278(Biochemistry,)-278(Ludwig-Maximilians-Universita)]TJ 0.1388 Tc 34.59 0 Td [(\250t)-139(M)139(u)]TJ 0 Tc 2.194 0 Td [(\250)-166(nchen,)-278(Butenandtstrasse)-278(11,)-278(81377)-278(Munich,)-278(Germany;)]TJ -41.961 -1.143 Td (and)Tj 5 0 0 5 66.0684 650.6335 Tm (\207)Tj 7 0 0 7 68.8487 648.3835 Tm [(Gene)-278(Center)-278(Munich)-278(and)-278(Department)-278(of)-278(Chemistry)-278(and)-278(Biochemistry,)-278(Ludwig-Maximilians-Universita)]TJ 0.1388 Tc 46.536 0 Td [(\250t)-139(M)139(u)]TJ 0 Tc 2.194 0 Td [(\250)-166(nchen,)-278(Feodor-Lynen-Strasse)-278(25,)]TJ -51.184 -1.143 Td [(81377)-278(Munich,)-278(Germany)]TJ 0 -2.286 TD [(Edited)-278(by)-278(Steven)-278(M.)-278(Block,)-278(Stanford)-278(University,)-278(Stanford,)-278(CA,)-278(and)-278(approved)-278(November)-278(5,)-278(2007)-278(\(received)-278(for)-278(review)-278(April)-278(25,)-278(2007\))]TJ /T1_1 1 Tf -0.0195 Tc 8 0 0 8 51.6761 608.3835 Tm [(Single-pair)-239(\037uorescence)-239(resonance)-239(energy)-239(transfer)-239(was)-239(used)-239(to)-239(track)]TJ 0 -1.251 TD [(RNA)-250(exiting)-250(from)-250(RNA)-250(polymerase)-250(II)-250(\(Pol)-250(II\))-250(in)-250(elongation)-250(complexes.)]TJ T* [(Measuring)-347(the)-347(distance)-347(between)-347(the)-347(RNA)-347(5)]TJ /T1_3 1 Tf 0 Tc 20.199 0 Td (\001)Tj /T1_1 1 Tf -0.0195 Tc 0.661 0 Td [(end)-347(and)-347(three)-347(known)]TJ -20.86 -1.251 Td [(locations)-506(within)-506(the)-506(elongation)-506(complex)-506(allows)-506(us)-506(determine)-506(its)]TJ T* [(position)-199(by)-199(means)-199(of)-199(triangulation.)-199(RNA)-199(leaves)-199(the)-199(polymerase)-199(active)]TJ 0 -1.251 TD [(center)-316(cleft)-316(via)-316(the)-316(previously)-316(proposed)-316(exit)-316(tunnel)-316(and)-316(then)-316(disen-)]TJ T* [(gages)-225(from)-225(the)-225(enzyme)-225(surface.)-225(When)-225(the)-225(RNA)-225(reaches)-225(lengths)-225(of)-225(26)]TJ T* [(and)-378(29)-378(nt,)-378(its)-378(5)]TJ /T1_3 1 Tf 0 Tc 7.115 0 Td (\001)Tj /T1_1 1 Tf -0.0195 Tc 0.692 0 Td [(end)-378(associates)-378(with)-378(Pol)-378(II)-378(at)-378(the)-378(base)-378(of)-378(the)-378(dock)]TJ -7.807 -1.251 Td [(domain.)-272(Because)-272(the)-272(initiation)-272(factor)-272(TFIIB)-272(binds)-272(to)-272(the)-272(dock)-272(domain)]TJ T* [(and)-296(exit)-296(tunnel,)-296(exiting)-296(RNA)-296(may)-296(prevent)-296(TFIIB)-296(reassociation)-296(during)]TJ T* [(elongation.)-241(RNA)-241(further)-241(extends)-241(toward)-241(the)-241(linker)-241(connecting)-241(to)-241(the)]TJ T* [(polymerase)-504(C-terminal)-504(repeat)-504(domain)-504(\(CTD\),)-504(which)-504(binds)-504(the)-504(5)]TJ /T1_3 1 Tf 0 Tc 30.3 0 Td (\001)Tj /T1_1 1 Tf 0.314 0 Td (-)Tj -0.0195 Tc -30.614 -1.251 Td [(capping)-313(enzyme)-313(and)-313(other)-313(RNA)-313(processing)-313(factors.)]TJ /T1_2 1 Tf 0 Tc 7 0 0 7 51.6761 468.2358 Tm [(Pol)-278(II)]TJ /T1_4 1 Tf 2.708 0 Td (\001)Tj /T1_2 1 Tf 0.651 0 Td (transcription)Tj /T1_4 1 Tf 6.374 0 Td (\001)Tj /T1_2 1 Tf 0.651 0 Td (FRET)Tj /T1_4 1 Tf 2.652 0 Td (\001)Tj /T1_2 1 Tf 0.651 0 Td (triangulation)Tj /T1_4 1 Tf 6.597 0 Td (\001)Tj /T1_2 1 Tf 0.651 0 Td (\037uorescence)Tj /T1_1 1 Tf 24.5 0 0 24.5 51.6761 435.2244 Tm (P)Tj /T1_5 1 Tf 0.005 Tc 9 0 0 9 66.2979 445.2244 Tm [(ol)-477(II)-477(synthe)20(size)20(s)-477(all)-477(euk)-30(ar)-60(yotic)-477(mRNA)-477(and)-477(c)-30(omprise)20(s)-477(12)]TJ -0 -1.112 Td [(subun)-30(its,)-185(Rpb1)-185(to)-185(Rpb12.)-185(A)-30(n)-185(atomic)-185(model)-185(of)-185(Pol)-185(II)-185(has)-185(been)]TJ -1.624 -1.112 Td [(obt)-30(ained)-437(by)-437(x-ray)-437(cr)-60(yst)-30(allog)-20(raphy)-437(\(1,)-437(2\).)]TJ -0.0149 Tc 18.498 0 Td [(Additional)-437(studie)20(s)-437(of)]TJ -18.498 -1.112 Td [(Pol)-305(II\226nucleic)-305(acid)-305(c)-30(omplexe)20(s)-305(have)-305(given)-305(insights)-305(into)-305(the)-305(elonga-)]TJ 0 -1.112 TD [(tion)-321(c)-30(omplex)-321(str)-30(ucture)-321(and)-321(molecular)-321(aspects)-321(of)-321(the)-321(transcription)]TJ T* [(mechan)-30(ism)-181(\(3\2267\).)-182(Cr)-60(yst)-30(allog)-20(raphic)-182(analysis)-181(detected)-181(the)-181(position)-181(of)]TJ T* [(the)-335(nascent)-335(RNA)-335(w)-40(ithin)-335(the)-335(DNA\226RNA)-335(hybrid)-335(above)-335(the)-335(active)]TJ T* [(site)-248(\(positions)]TJ /T1_6 1 Tf 0 Tc 5.97 0 Td (\001)Tj /T1_5 1 Tf 0.2329 Tc 0.818 0 Td [(1t)248(o)]TJ /T1_6 1 Tf 0 Tc 1.822 0 Td (\002)Tj /T1_5 1 Tf -0.0148 Tc 0.818 0 Td [(8,)-248(w)-40(ith)]TJ /T1_6 1 Tf 0 Tc 3.02 0 Td (\001)Tj /T1_5 1 Tf -0.0149 Tc 0.818 0 Td [(1)-248(denoting)-248(the)-248(nucleotide)-248(addition)]TJ -13.266 -1.112 Td [(site\))-552(and)-552(for)-552(2)-552(nt)-552(upstream)-552(of)-552(the)-552(hybrid)-552(af)-30(ter)-552(the)-552(point)-552(of)]TJ 0 -1.112 TD [(DNA\226RNA)-291(strand)-291(separation)-291(\(positions)]TJ /T1_6 1 Tf 0 Tc 17.197 0 Td (\002)Tj /T1_5 1 Tf -0.0149 Tc 0.818 0 Td [(9)-291(and)]TJ /T1_6 1 Tf 0 Tc 2.611 0 Td (\002)Tj /T1_5 1 Tf -0.0149 Tc 0.818 0 Td [(10\))-291(\(3,)-291(7\).)-291(The)]TJ -21.445 -1.112 Td [(last-ordered)-255(RNA)-255(nucleotide)-255(is)-255(located)-255(at)-255(the)-255(entrance)-255(to)-255(a)-255(tunnel)]TJ T* [([called)-364(the)-364(RNA)-364(ex)-50(it)-364(channel)-364(for)-364(the)-364(bacterial)-364(RNA)-364(poly)-30(merase)]TJ T* [(\(8\)],)-327(which)-327(is)-327(for)-30(med)-327(among)-327(the)-327(poly)-30(merase)-327(wall,)-327(clamp,)-327(and)-327(lid.)]TJ T* [(This)-193(tunnel)-194(leads)-193(f)-30(rom)-193(the)-193(active)-193(center)-193(clef)-30(t)-193(to)-193(the)-193(exterior)-193(and)-193(was)]TJ T* [(proposed)-359(to)-359(ac)-30(c)-30(ommodate)-359(ex)-50(iting)-359(RNA)-359(\(1,)-359(3,)-359(6,)-359(7\).)]TJ 0.005 Tc 22.474 0 Td [(Beyond)-359(the)]TJ -22.474 -1.112 Td [(put)-30(ative)-387(ex)-50(it)-387(tunnel,)-387(t)-30(wo)-387(prominent)-387(sur)-50(face)-387(g)-20(roove)20(s)-387(on)-387(either)]TJ T* [(side)-223(of)-223(the)-223(dock)-223(domain)-223(in)-223(principle)-223(c)-30(ould)-223(further)-223(ac)-30(c)-30(ommodate)]TJ T* [(ex)-50(iting)-264(RNA)-264(\(1,)-264(9\).)-264(Groove)-264(1)-264(w)-40(inds)-264(along)-264(the)-264(base)-264(of)-264(the)-264(clamp)]TJ T* [(toward)-384(the)-384(Rpb4/7)-384(subc)-30(omplex,)-384(whereas)-384(g)-20(roove)-384(2)-384(leads)-384(along)]TJ T* [(Rpb11)-426(toward)-426(Rpb8.)-426(Recently,)-426(nascent)-426(RNA)-426(c)-30(ould)-426(be)-426(cross-)]TJ T* [(linked)-357(to)-357(Rpb7,)-357(prov)-40(iding)-357(apparent)-357(support)-357(for)-357(g)-20(roove)-357(1)-357(\(10\).)]TJ T* [(RNA)-323(beyond)-323(position)]TJ /T1_6 1 Tf 0 Tc 9.914 0 Td (\002)Tj /T1_5 1 Tf 0.005 Tc 0.838 0 Td [(10)-323(was)-323(pre)20(sent)-323(in)-323(one)-323(of)-323(the)-323(cr)-60(yst)-30(allo-)]TJ -10.752 -1.112 Td [(g)-20(raphic)-326(studie)20(s)-326(of)-326(the)-326(elongation)-326(c)-30(omplex)-326(\(3\))-326(but)-326(c)-30(ould)-326(not)-326(be)]TJ T* [(obser)-60(ved)-286(in)-286(the)-286(tunnel)-286(or)-286(in)-286(the)-286(subsequent)-286(g)-20(roove)20(s,)-286(sugge)20(sting)]TJ T* [(that)-399(its)-399(interactions,)-399(if)-399(they)-399(ex)-50(ist,)-399(are)-399(transient)-399(and)-399(cannot)-399(be)]TJ T* [(detected)-338(in)-338(medium-re)20(solution)-338(electron)-338(densit)-30(y)-338(maps.)]TJ -0.0149 Tc 1 -1.112 Td [(To)-267(study)-267(if)-267(the)-267(nascent)-267(RNA)-267(indeed)-267(ex)-50(its)-267(through)-267(the)-267(proposed)]TJ -1 -1.112 Td [(tunnel,)-339(and)-339(whether)-339(it)-339(follows)-339(a)-339(defined)-339(sur)-50(face)-339(path)-339(beyond)-339(the)]TJ T* [(tunnel,)-282(we)-282(used)-282(single-molecule)-282(f)-90(luore)20(scence)-282(ex)-30(periments.)-282(Single-)]TJ T* [(particle)-316(methods)-316(prevent)-316(the)-316(loss)-316(of)-316(infor)-30(mation)-316(because)-316(of)-316(aver-)]TJ T* [(aging)-487(that)-487(is)-487(inherent)-487(to)-487(bulk)-487(ex)-30(periments)-487(and)-487(cr)-60(yst)-30(allog)-20(raphy)]TJ 0 -1.111 TD [(\(11\22614\).)-570(Fluore)20(scence)-570(re)20(sonance)-570(energ)-40(y)-570(transfer)-570(bet)-30(ween)-570(t)-30(wo)]TJ T* [(f)-90(luorophore)20(s)-219([single-pair)-219(\(sp\)-FR)-60(ET])-219(prov)-40(ide)20(s)-219(a)-219(ver)-60(y)-219(sensitive)-219(tool)]TJ T* [(for)-182(studying)-182(dist)-30(ance)20(s)-182(and)-182(c)-30(onfor)-30(mational)-182(change)20(s)-182(w)-40(ithin)-182(biological)]TJ T* [(c)-30(omplexe)20(s)-318(in)-318(the)-318(range)-318(of)-318(10)-90(\226100)-318(\305)-318(\(15,)-318(16\).)]TJ 1 -1.111 Td [(Here)-248(we)-248(used)-248(sp-FR)-60(ET)-248(to)-248(infer)-248(the)-248(position)-248(of)-248(the)-248(5)]TJ /T1_6 1 Tf 0 Tc 21.718 0 Td (\003)Tj /T1_5 1 Tf -0.0149 Tc 0.567 0 Td [(end)-248(of)-248(the)]TJ -23.285 -1.111 Td [(nascent)-378(RNA)-378(by)-378(relative)-378(dist)-30(ance)-378(measurements)-378(to)-378(k)-30(nown)-378(posi-)]TJ T* [(tions)-463(in)-463(the)-463(Pol)-463(II)-463(elongation)-463(c)-30(omplex.)-463(In)-463(a)-463(proce)20(ss)-463(k)-30(nown)-463(as)]TJ T* [(triangulation/trilateration,)-441(we)-441(deter)-30(mined)-441(a)-441(prev)-40(iously)-441(unk)-30(nown)]TJ 29.167 60.333 Td [(location)-366(by)-366(measuring)-366(the)-366(dist)-30(ance)-366(to)-366(that)-366(position)-366(f)-30(rom)-366(at)-366(least)]TJ 0 -1.134 TD [(three)-342(k)-30(nown)-342(points)-342(in)-342(space.)-342(This)-342(proce)20(ss)-342(allowed)-342(us)-342(to)-342(map)-342(the)]TJ T* [(path)-288(of)-288(the)-288(ex)-50(iting)-288(RNA)-288(upstream)-288(of)-288(position)]TJ /T1_6 1 Tf 0 Tc 19.423 0 Td (\002)Tj /T1_5 1 Tf -0.0149 Tc 0.818 0 Td [(10.)-288(In)-288(this)-288(article,)]TJ -20.241 -1.134 Td [(implications)-225(of)-225(our)-225(work)-225(for)-225(underst)-30(anding)-225(the)-225(transcription)-225(mech-)]TJ T* [(an)-30(ism)-318(are)-318(discussed.)]TJ /T1_7 1 Tf -0.0196 Tc 0 -2.023 TD (Results)Tj 0 Tc 8 0 0 8 314.1761 536.6562 Tm [(Engineering)-249(Fluorophore-Labeled)-249(Active)-249(Pol)-249(II)-249(Elongation)-249(Complexes.)]TJ /T1_5 1 Tf -0.0149 Tc 9 0 0 9 314.1761 526.4517 Tm [(The)-309(possibilit)-30(y)-309(to)-309(assemble)-309(c)-30(omplete)-309(Pol)-309(II)-309(elongation)-309(c)-30(omplexe)20(s)]TJ 0 -1.134 TD [(f)-30(rom)-578(endogenous)-578(yeast)-578(10-subun)-30(it)-578(c)-30(ore)-578(enz)-30(y)-30(me,)-578(rec)-30(ombinant)]TJ T* [(Rpb4/7)-177(subc)-30(omplex,)-177(and)-177(synthetic)-177(nucleic)-177(acid)-177(scaf)-30(folds)-177(\(3\))-177(enabled)]TJ T* [(the)-309(preparation)-309(of)-309(defined)-309(elongation)-309(c)-30(omplexe)20(s)-309(w)-40(ith)-309(f)-90(luore)20(scent)]TJ T* [(labels)-331(at)-331(discrete)-331(positions.)-331(L)-30(abels)-331(were)-331(readily)-331(inc)-30(orporated)-331(into)]TJ T* [(the)-348(synthetic)-348(nucleic)-348(acids.)-348(To)-348(reliably)-348(predict)-348(label)-348(locations,)-348(we)]TJ T* [(used)-546(nucleic)-546(acid)-546(scaf)-30(folds)-546(that)-546(closely)-546(re)20(semble)-546(those)-546(in)-546(the)]TJ T* [(c)-30(omplete)-408(Pol)-408(II)-408(elongation)-408(c)-30(omplex)-408(str)-30(ucture)-408(\(3\).)-408(We)-408(prepared)]TJ T* [(scaf)-30(folds)-243(w)-40(ith)-243(dif)-30(ferent)-243(RNA)-243(lengths,)-243(ranging)-243(f)-30(rom)-243(17)-243(to)-243(35)-243(nt)-243(and)]TJ 0 -1.134 TD [(c)-30(omprising)-289(a)-289(donor)-289(tetramethylrhodamine)-289(\(TMR\))-289(f)-90(luorophore)-289(at)]TJ T* [(their)-176(5)]TJ /T1_6 1 Tf 0 Tc 2.611 0 Td (\003)Tj /T1_5 1 Tf -0.0149 Tc 0.494 0 Td [(ends)-176(\(RNA17,)-176(RNA)-60(20,)-176(RNA)-60(23,)-176(RNA)-60(26,)-176(RNA)-60(29,)-176(RNA32,)]TJ -3.105 -1.134 Td [(and)-518(RNA35;)-518(Fig.)-518(1)-518(and)]TJ /T1_8 1 Tf 11.225 0 Td [(Mater)-30(ials)-518(and)-518(Methods)]TJ /T1_5 1 Tf 9.73 0 Td [(\).)-518(A)-30(n)-518(ac)-30(ceptor)]TJ -20.955 -1.134 Td [(f)-90(luorophore)-224(\(A)-30(lexa)-224(647\))-224(was)-224(att)-30(ached)-224(to)-224(the)-224(DNA)-224(template)-224(strand)]TJ T* [(either)-354(at)-354(position)]TJ /T1_6 1 Tf 0 Tc 7.603 0 Td (\002)Tj /T1_5 1 Tf -0.0149 Tc 0.818 0 Td [(10)-354(\(DNA1\))-354(or)-354(position)]TJ /T1_6 1 Tf 0 Tc 10.128 0 Td (\001)Tj /T1_5 1 Tf -0.0149 Tc 0.818 0 Td [(3)-354(\(DNA)-60(2\).)-354(A)-354(third)]TJ -19.367 -1.134 Td [(ac)-30(ceptor)-276(label)-276(was)-276(c)-30(oupled)-276(to)-276(an)-276(ac)-30(ce)20(ssible)-276(c)-30(ysteine)-276(re)20(sidue)-276(in)-276(the)]TJ T* [(Rpb4/7)-345(c)-30(omplex)-345(\(Rpb7\226C150;)-345(Fig.)-345(1)-345(and)]TJ /T1_8 1 Tf 17.737 0 Td [(Mater)-30(ials)-345(and)-345(Methods)]TJ /T1_5 1 Tf 0 Tc 9.384 0 Td (\))Tj -0.0149 Tc -27.121 -1.134 Td [(af)-30(ter)-309(other)-309(c)-30(ysteine)20(s)-309(had)-309(been)-309(replaced)-309(by)-309(serine)20(s)-309(w)-40(ith)-309(the)-309(use)-309(of)]TJ T* [(site-directed)-318(mut)-30(agene)20(sis)-318(in)]TJ /T1_8 1 Tf 11.961 0 Td [(Escher)-30(ichia)-318(coli)]TJ /T1_5 1 Tf 0 Tc 6.418 0 Td (.)Tj -0.0149 Tc -17.379 -1.133 Td [(The)-401(re)20(sulting)-401(elongation)-401(c)-30(omplexe)20(s)-401(allowed)-401(us)-401(to)-401(localize)-401(the)]TJ -1 -1.133 Td [(RNA)-389(5)]TJ /T1_6 1 Tf 0 Tc 3.136 0 Td (\003)Tj /T1_5 1 Tf -0.0149 Tc 0.707 0 Td [(end)-389(at)-389(various)-389(positions)-389(and)-389(thus)-389(to)-389(map)-389(the)-389(path)-389(that)]TJ -3.844 -1.133 Td [(the)-270(g)-20(row)-40(ing)-270(RNA)-270(chain)-270(follows)-270(when)-270(ex)-50(iting)-270(the)-270(poly)-30(merase.)-270(The)]TJ 0 -1.133 TD [(c)-30(omplexe)20(s)-184(are)-184(active)-184(in)-184(RNA)-184(chain)-184(elongation)-184(in)-184(ensemble)-184(\(17\))-184(and)]TJ T* [(single-molecule)-289(ex)-30(periments)-289([see)]TJ 0 0 1 rg 14.027 0 Td [(supporting)-289(infor)-30(mation)-289(\(SI\))-289(Fig.)]TJ 0 Tc -14.027 -1.133 Td (5)Tj 0 0 0 rg -0.0149 Tc 0.485 0 Td [(].)-384(They)-384(may)-384(therefore)-384(be)-384(v)-40(iewed)-384(as)-384(st)-30(able)-384(inter)-30(mediate)20(s)-384(in)-384(the)]TJ -0.485 -1.133 Td [(early)-318(elongation)-318(proce)20(ss.)]TJ /T1_7 1 Tf 0 Tc 8 0 0 8 314.1761 261.429 Tm [(sp-FRET)-252(Analysis.)]TJ /T1_5 1 Tf -0.0149 Tc 9 0 0 9 377.0413 261.429 Tm [(Using)-224(a)-224(single-molecule)-224(f)-90(luore)20(scence)-224(microsc)-30(ope,)]TJ -6.985 -1.133 Td [(we)-422(measured)-422(sp-FR)-60(ET)-422(signals)-422(f)-30(rom)-422(indiv)-40(idual)-422(elongation)-422(c)-30(om-)]TJ T* [(plexe)20(s)-504(\()]TJ /T1_8 1 Tf 3.395 0 Td [(Mater)-30(ials)-504(and)-504(Methods)]TJ /T1_5 1 Tf 9.703 0 Td [(\).)-504(We)-504(rec)-30(orded)-504(the)-504(f)-90(luore)20(scence)]TJ -13.097 -1.133 Td [(intensitie)20(s)-309(of)-309(donor)-309(and)-309(ac)-30(ceptor)-309(molecule)20(s)-309(as)-309(a)-309(function)-309(of)-309(time.)]TJ T* [(With)-529(the)20(se)-529(trajectorie)20(s,)-529(we)-529(c)-30(omputed)-529(the)-529(re)20(spective)-529(sp-FR)-60(ET)]TJ T* [(ef)-30(ficiencie)20(s.)-399(Using)-399(the)-399(dat)-30(a)-399(f)-30(rom)-399(many)-399(molecule)20(s,)-399(we)-399(then)-399(c)-30(on-)]TJ T* [(str)-30(ucted)-206(histog)-20(rams)-206(of)-206(FR)-60(ET)-206(ef)-30(ficiencie)20(s.)-206(For)-206(each)-206(length)-206(of)-206(RNA,)]TJ T* [(we)-179(per)-50(for)-30(med)-179(three)-179(independent)-179(sets)-179(of)-179(ex)-30(periments)-179(measuring)-179(the)]TJ T* [(FR)-60(ET)-201(ef)-30(ficienc)-30(y)-201(of)-201(c)-30(omplexe)20(s)-201(w)-40(ith)-201(the)-201(donor)-201(molecule)-201(att)-30(ached)-201(to)]TJ ET 0 0 0 RG 0.25 w 10 M 0 j 0 J []0 d 314.176 164.509 m 386.176 164.509 l S BT /T1_2 1 Tf 0 Tc 6 0 0 6 314.1761 151.8835 Tm [(Author)-289(contributions:)-289(J.A.,)-289(F.B.,)-289(P.C.,)-289(and)-289(J.M.)-289(designed)-289(research;)-289(J.A.,)-289(R.L.,)-289(F.B.,)-289(E.L.,)-289(P.C.,)]TJ 0 -1.333 TD [(and)-316(J.M.)-316(performed)-316(research;)-316(F.B.,)-316(E.L.,)-316(P.C.,)-316(and)-316(J.M.)-316(contributed)-316(new)-316(reagents/analytic)]TJ T* [(tools;)-158(J.A.,)-158(R.L.,)-158(F.B.,)-158(P.C.,)-158(and)-158(J.M.)-158(analyzed)-158(data;)-158(and)-158(J.A.,)-158(R.L.,)-158(F.B.,)-158(E.L.,)-158(P.C.,)-158(and)-158(J.M.)-158(wrote)]TJ T* [(the)-278(paper.)]TJ 0 -1.833 TD [(The)-278(authors)-278(declare)-278(no)-278(con\037ict)-278(of)-278(interest.)]TJ 0 -1.833 TD [(This)-278(article)-278(is)-278(a)-278(PNAS)-278(Direct)-278(Submission.)]TJ 4.5 0 0 4.5 314.1761 96.6335 Tm (\247)Tj 6 0 0 6 316.6786 94.8835 Tm [(To)-278(whom)-278(correspondence)-278(should)-278(be)-278(addressed.)-278(E-mail:)-278(michaelis@lmu.de.)]TJ -0.417 -1.833 Td [(This)-513(article)-513(contains)-513(supporting)-513(information)-513(online)-513(at)]TJ 0 0 1 rg 26.82 0 Td (www.pnas.org/cgi/content/full/)Tj -26.82 -1.333 Td (0703815105/DC1)Tj 0 0 0 rg 7.727 0 Td (.)Tj -7.727 -1.833 Td [(\251)-278(2007)-278(by)-278(The)-278(National)-278(Academy)-278(of)-278(Sciences)-278(of)-278(the)-278(USA)]TJ 6.5 0 0 6.5 51.6761 39.8835 Tm (www.pnas.org)Tj /T1_4 1 Tf 6.833 0 Td (\002)Tj /T1_2 1 Tf 0.5 0 Td (cgi)Tj /T1_4 1 Tf 1.333 0 Td (\002)Tj /T1_2 1 Tf 0.5 0 Td (doi)Tj /T1_4 1 Tf 1.5 0 Td (\002)Tj /T1_2 1 Tf 0.5 0 Td (10.1073)Tj /T1_4 1 Tf 3.614 0 Td (\002)Tj /T1_2 1 Tf 0.5 0 Td [(pnas.0703815105)-26701(PNAS)]TJ /T1_4 1 Tf 9.75 0 0 6.5 398.8389 39.8835 Tm (\001)Tj /T1_1 1 Tf 6.5 0 0 6.5 407.0035 39.8835 Tm [(January)-333(8,)-333(2008)]TJ /T1_4 1 Tf 9.75 0 0 6.5 461.048 39.8835 Tm (\001)Tj /T1_2 1 Tf 6.5 0 0 6.5 469.2126 39.8835 Tm [(vol.)-333(105)]TJ /T1_4 1 Tf 9.75 0 0 6.5 499.0529 39.8835 Tm (\001)Tj /T1_2 1 Tf 6.5 0 0 6.5 507.2175 39.8835 Tm [(no.)-333(1)]TJ /T1_4 1 Tf 9.75 0 0 6.5 528.7439 39.8835 Tm (\001)Tj /T1_1 1 Tf 6.5 0 0 6.5 536.9084 39.8835 Tm (135\226140)Tj ET 40 w 593.676 455.384 m 593.676 521.384 l S 0 0 0 0 k BT /T1_7 1 Tf 0 6.5 -6.5 0 585.1761 471.2386 Tm (BIOPHYSICS)Tj ET q /GS1 gs 19 0 0 815.5 9 9 cm /Im0 Do Q endstream endobj 125 0 obj<> endobj 126 0 obj<>stream 0 0 0 1 k 1 i /GS0 gs BT /T1_0 1 Tf -0.0149 Tc 0 Tw 0 Ts 100 Tz 0 Tr 9 0 0 9 51.6761 355.8835 Tm [(the)-192(RNA)-192(5)]TJ /T1_1 1 Tf 0 Tc 4.455 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc 0.51 0 Td [(end)-192(and)-192(the)-192(ac)-30(ceptor)-192(to)-192(DNA1,)-192(DNA)-60(2,)-192(or)-192(Rpb7\226C150.)]TJ -4.966 -1.116 Td [(Fig.)-532(2)-532(shows)-532(as)-532(example)20(s)-532(sp-FR)-60(ET)-532(histog)-20(rams)-532(for)-532(elongation)]TJ 0 -1.115 TD [(c)-30(omplexe)20(s)-391(c)-30(ont)-30(ain)-30(ing)-391(DNA1)-391(and)-391(RNA17)-391(as)-391(well)-391(as)-391(DNA1)-391(and)]TJ T* [(RNA)-60(26,)-484(together)-484(w)-40(ith)-484(characteristic)-484(time)-484(trajectorie)20(s)-484(of)-484(donor)]TJ 0 -1.115 TD [(f)-90(luore)20(scence,)-356(ac)-30(ceptor)-356(f)-90(luore)20(scence,)-356(and)-356(c)-30(omputed)-356(sp-FR)-60(ET)-356(ef-)]TJ T* [(ficiencie)20(s.)-291(A)-30(ll)-291(other)-291(histog)-20(rams)-291(are)-291(shown)-291(in)-291(the)-291(supporting)-291(infor-)]TJ T* [(mation)-426(\()]TJ 0 0 1 rg 3.712 0 Td [(SI)-426(Figs.)-426(6)-426(and)-426(7)]TJ 0 0 0 rg 7.055 0 Td [(\).)-426(The)-426(main)-426(re)20(sults)-426(are)-426(summarized)-426(in)]TJ -10.767 -1.115 Td [(Table)-318(1.)]TJ 1 -1.115 Td [(The)-288(trajector)-60(y)-288(of)-288(FR)-60(ET)-288(ef)-30(ficienc)-30(y)]TJ /T1_2 1 Tf 0 Tc 14.67 0 Td (E)Tj /T1_0 1 Tf -0.0149 Tc 0.949 0 Td [(shown)-288(in)-288(Fig.)-288(2)]TJ /T1_2 1 Tf 0 Tc 6.457 0 Td (A)Tj /T1_0 1 Tf -0.0149 Tc 0.963 0 Td [(is)-288(rather)]TJ -24.039 -1.115 Td [(c)-30(onst)-30(ant,)-292(which)-292(is)-292(the)-292(t)-30(ypical)-292(behav)-40(ior)-292(obser)-60(ved)-292(for)-292(all)-292(elongation)]TJ T* [(c)-30(omplexe)20(s)-215(w)-40(ith)-215(RNA17,)-215(RNA)-60(20,)-215(or)-215(RNA)-60(23.)-215(In)-215(c)-30(ontrast,)-215(when)-215(the)]TJ T* [(RNA)-398(is)-398(extended)-398(f)-30(rom)-398(23)-398(to)-398(26)-398(nt,)-398(the)-398(FR)-60(ET)-398(signal)-398(no)-398(longer)]TJ T* [(remains)-215(c)-30(onst)-30(ant.)-215(A)-215(t)-30(ypical)-215(example)-215(is)-215(shown)-215(in)-215(Fig.)-215(2)]TJ /T1_2 1 Tf 0 Tc 22.049 0 Td (C)Tj /T1_0 1 Tf -0.0149 Tc 0.686 0 Td [(:)-215(the)-215(FR)-60(ET)]TJ -22.735 -1.115 Td [(ef)-30(ficienc)-30(y)-582(of)-582(the)-582(DNA1\226RNA)-60(26)-582(elongation)-582(c)-30(omplex)-582(sw)-40(itche)20(s)]TJ T* [(repeatedly)-379(bet)-30(ween)-379(t)-30(wo)-379(levels,)-379(and)-379(the)-379(c)-30(orre)20(sponding)-379(histog)-20(ram)]TJ T* [(shows)-376(t)-30(wo)-376(peaks)-376(\(Fig.)-376(2)]TJ /T1_2 1 Tf 0 Tc 10.284 0 Td (D)Tj /T1_0 1 Tf -0.0149 Tc 0.74 0 Td [(\).)-376(Such)-376(apparent)-376(f)-90(luctuations)-376(c)-30(ould)-376(be)]TJ -11.024 -1.115 Td [(caused)-172(by)-172(either)-172(dist)-30(ance)-172(variations)-172(or)-172(photo-physical)-172(change)20(s)-172(in)-172(the)]TJ 0 -1.115 TD [(donor)-366(and/or)-366(ac)-30(ceptor)-366(molecule.)-366(We)-366(checked)-366(for)-366(change)20(s)-366(in)-366(the)]TJ T* [(ac)-30(ceptor)-258(brightne)20(ss)-258(by)-258(exciting)-258(the)-258(ac)-30(ceptor)-258(directly)-258(in)-258(an)-258(alternat-)]TJ T* [(ing)-368(excit)-30(ation)-368(scheme)-368(\(18,)-368(19\).)-368(A)-368(t)-30(ypical)-368(example)-368(of)-368(a)-368(trajector)-60(y)]TJ T* [(show)-40(ing)-423(strong)-423(modulation)-423(of)-423(the)-423(FR)-60(ET)-423(ef)-30(ficienc)-30(y)-423(when)-423(being)]TJ T* [(excited)-315(w)-40(ith)-315(g)-20(reen)-315(light)-315(and)-315(c)-30(onst)-30(ant)-315(f)-90(luore)20(scence)-315(intensit)-30(y)-315(when)]TJ T* [(excited)-277(w)-40(ith)-277(red)-277(light)-277(is)-277(shown)-277(in)-277(Fig.)-277(3.)-277(We)-277(further)-30(more)-277(analyzed)]TJ T* [(the)-502(tot)-30(al)-502(intensit)-30(y)-502(of)-502(emitted)-502(f)-90(luore)20(scence)-502(and)-502(found)-502(that)-502(the)]TJ T* [(obser)-60(ved)-326(f)-90(luctuations)-326(are)-326(not)-326(caused)-326(by)-326(change)20(s)-326(of)-326(donor)-326(bright-)]TJ T* [(ne)20(ss)-380(\()]TJ 0 0 1 rg 2.475 0 Td [(SI)-380(Fig.)-380(8)]TJ 0 0 0 rg 3.715 0 Td [(\).)-380(Thus,)-380(the)-380(obser)-60(ved)-380(f)-90(luctuations)-380(of)]TJ /T1_2 1 Tf 0 Tc 16.074 0 Td (E)Tj /T1_0 1 Tf -0.0149 Tc 1.041 0 Td [(cannot)-380(be)]TJ -23.305 -1.115 Td [(attributed)-514(to)-514(photo-physical)-514(change)20(s)-514(of)-514(the)-514(donor)-514(or)-514(ac)-30(ceptor)]TJ T* [(molecule,)-246(instead)-246(our)-246(single-molecule)-246(probe)-246(reports)-246(on)-246(a)-246(real-time)]TJ T* [(reordering)-384(of)-384(the)-384(g)-20(row)-40(ing)-384(RNA)-384(chain,)-384(w)-40(ith)-384(an)-384(apparent)-384(rate)-384(of)]TJ 0 -1.115 TD [(reordering)-318(of)]TJ /T1_1 1 Tf 0 Tc 5.869 0 Td (\004)Tj /T1_0 1 Tf 0.303 Tc 0.818 0 Td (1s)Tj /T1_1 1 Tf 0 Tc 6.5 0 0 6 122.3961 67.8835 Tm (\002)Tj /T1_0 1 Tf 0.819 0 Td (1)Tj 9 0 0 9 130.8734 64.8835 Tm (.)Tj -0.0149 Tc 21.367 74.444 Td [(In)-403(a)-403(next)-403(step,)-403(we)-403(used)-403(the)-403(mean)-403(FR)-60(ET)-403(ef)-30(ficiencie)20(s)-403(\()]TJ /T1_2 1 Tf 0 Tc 23.469 0 Td (E)Tj /T1_0 1 Tf -0.0149 Tc 0.661 0 Td [(\))-403(\(ex-)]TJ -25.13 -1.111 Td [(tracted)-240(f)-30(rom)-240(the)-240(histog)-20(rams\))-240(to)-240(c)-30(ompute)-240(dist)-30(ance)20(s)-240(\()]TJ /T1_2 1 Tf 0 Tc 21.454 0 Td (d)Tj /T1_0 1 Tf -0.0149 Tc 0.513 0 Td [(\))-240(by)-240(using)-240(the)]TJ -21.967 -1.111 Td [(deter)-30(mined)-370(Fo)]TJ 5.807 0 Td [(\250)-22(rster)-370(dist)-30(ance)20(s)-370(\()]TJ /T1_2 1 Tf 7.271 0 Td [(Mater)-30(ials)-370(and)-370(Methods)]TJ /T1_0 1 Tf 9.434 0 Td [(\).)-370(Then,)-370(for)]TJ -22.513 -1.111 Td [(each)-205(length)-205(of)-205(RNA,)-205(we)-205(deter)-30(mined)-205(the)-205(dist)-30(ance)-205(of)-205(its)-205(5)]TJ /T1_1 1 Tf 0 Tc 22.847 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc 0.523 0 Td [(end)-205(to)-205(the)]TJ -23.371 -1.111 Td [(three)-296(k)-30(nown)-296(points)-296(in)-296(space)-296(\(DNA1,)-296(DNA)-60(2,)-296(and)-296(Rpb7\226C150\))-296(by)]TJ 0 -1.111 TD [(using)-383(the)-383(re)20(spective)-383(mean)-383(FR)-60(ET)-383(ef)-30(ficiencie)20(s)-383(extracted)-383(f)-30(rom)-383(the)]TJ T* [(rec)-30(orded)-503(histog)-20(rams)-503(\(Table)-503(1\).)-503(To)-503(infer)-503(the)-503(unk)-30(nown)-503(relative)]TJ T* [(position)-241(of)-241(the)-241(dye)-241(molecule,)-241(we)-241(c)-30(onstr)-30(ucted)-241(a)-241(sphere)-241(around)-241(each)]TJ T* [(of)-184(the)-184(three)-184(k)-30(nown)-184(positions)-184(w)-40(ith)-184(a)-184(radius)-184(deter)-30(mined)-184(by)-184(the)-184(FR)-60(ET)]TJ T* [(measurement.)-376(The)-376(intersection)-376(point)-376(of)-376(the)-376(three)-376(sphere)20(s)-376(deter-)]TJ 0 -1.111 TD [(mine)20(s)-230(the)-230(unk)-30(nown)-230(position.)-230(A)-230(similar)-230(procedure)-230(has)-230(been)-230(applied)]TJ T* [(prev)-40(iously)-318(to)-318(the)-318(study)-318(of)-318(helicase)-60(\226DNA)-318(c)-30(omplexe)20(s)-318(\(20\).)]TJ /T1_3 1 Tf 0 Tc 8 0 0 8 314.1761 604.8835 Tm [(Accurate)-288(Localization)-288(of)-288(the)-288(RNA)-288(5)]TJ /T1_4 1 Tf 15.592 0 Td (\001)Tj /T1_3 1 Tf 0.621 0 Td (End.)Tj /T1_0 1 Tf -0.0149 Tc 9 0 0 9 461.4579 604.8835 Tm [(To)-256(te)20(st)-256(the)-256(ac)-30(curac)-30(y)-256(of)-256(our)]TJ -16.365 -1.111 Td [(method,)-388(we)-388(c)-30(ompared)-388(dist)-30(ance)20(s)-388(c)-30(omputed)-388(f)-30(rom)-388(sp-FR)-60(ET)-388(mea-)]TJ 0 -1.111 TD [(surements)-358(to)-358(dist)-30(ance)20(s)-358(w)-40(ithin)-358(the)-358(cr)-60(yst)-30(allog)-20(raphic)-358(str)-30(ucture.)-358(The)]TJ T* [(mean)-267(FR)-60(ET)-267(ef)-30(ficiencie)20(s)-267(of)-267(a)-267(donor)-267(molecule)-267(on)-267(DNA1)-267(or)-267(DNA)-60(2)]TJ T* [(and)-309(an)-309(ac)-30(ceptor)-309(on)-309(Rpb7\226C150)-309(re)20(sulted)-309(in)-309(dist)-30(ance)20(s)-309(of)-309(61)-309(\305)-309(and)]TJ 0 -1.111 TD [(63)-177(\305,)-177(re)20(spectively)-177(\()]TJ 0 0 1 rg 7.664 0 Td [(SI)-177(Fig.)-177(9)]TJ 0 0 0 rg 3.31 0 Td [(\).)-177(The)20(se)-177(dist)-30(ance)20(s)-177(are)-177(in)-177(good)-177(ag)-20(reement)]TJ -10.974 -1.111 Td [(w)-40(ith)-311(the)-311(value)20(s)-311(of)-311(65)-311(\305)-311(\(DNA1\))-311(and)-311(64)-311(\305)-311(\(DNA)-60(2\))-311(in)-311(the)-311(cr)-60(yst)-30(al)]TJ T* [(str)-30(ucture)-318([PDB)-318(c)-30(ode)-318(1Y1W)-318(\(3\)].)]TJ 1 -1.111 Td [(It)-330(is)-330(k)-30(nown)-330(that)-330(a)-330(TMR)-330(f)-90(luorophore)-330(att)-30(ached)-330(to)-330(the)-330(5)]TJ /T1_1 1 Tf 0 Tc 23.125 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc 0.648 0 Td [(end)-330(of)]TJ -24.773 -1.111 Td [(DNA)-211(v)-40(ia)-211(a)-211(f)-90(lex)-50(ible)-211(six-carbon)-211(linker)-211(usually)-211(is)-211(st)-30(acked)-211(on)-211(top)-211(of)-211(the)]TJ T* [(last)-407(base)-407(w)-40(ithin)-407(the)-407(nucleic)-407(acid)-407(helix)-407(\(21\).)-407(A)-30(lthough)-407(in)-407(case)-407(of)]TJ T* [(RNA,)-487(the)20(se)-487(interactions)-487(might)-487(be)-487(weaker,)-487(st)-30(ack)-30(ing)-487(of)-487(the)-487(dye)]TJ T* [(molecule)-523(to)-523(the)-523(last)-523(base)-523(of)-523(the)-523(RNA)-523(still)-523(is)-523(the)-523(most)-523(likely)]TJ T* [(c)-30(onfor)-30(mation.)-378(Thus,)-378(the)-378(dye)-378(molecule)-378(is)-378(ex)-30(pected)-378(to)-378(sit)-378(in)-378(close)]TJ T* [(prox)-50(imit)-30(y)-185(to)-185(the)-185(last)-185(base)-185(and)-185(therefore)-185(can)-185(report)-185(ac)-30(curately)-185(on)-185(the)]TJ 0 -1.111 TD [(position)-217(of)-217(the)-217(RNA.)-217(A)-30(lthough)-217(an)-217(interaction)-217(bet)-30(ween)-217(the)-217(charged)]TJ 0 -1.111 TD [(sur)-50(face)-301(of)-301(the)-301(poly)-30(merase)-301(and)-301(the)-301(dye)-301(molecule)-301(att)-30(ached)-301(to)-301(its)-301(5)]TJ /T1_1 1 Tf 0 Tc 27.172 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc -27.172 -1.111 Td [(end)-493(in)-493(principle)-493(c)-30(ould)-493(inf)-90(luence)-493(the)-493(position)-493(of)-493(the)-493(RNA,)-493(no)]TJ T* [(systematic)-364(ef)-30(fect)-364(was)-364(obser)-60(ved)-364(\()]TJ 0 0 1 rg 13.684 0 Td [(SI)-364(Fig.)-364(10)]TJ 0 0 0 rg 4.168 0 Td [(\).)-364(However,)-364(we)-364(cannot)]TJ -17.852 -1.111 Td [(r)-30(ule)-324(out)-324(the)-324(possibilit)-30(y)-324(that)-324(such)-324(a)-324(direct)-324(interaction)-324(bet)-30(ween)-324(dye)]TJ T* [(molecule)-318(and)-318(protein)-318(sur)-50(face)-318(ex)-50(ists.)]TJ 1 -1.111 Td [(Moreover,)-199(we)-199(checked)-199(the)-199(deter)-30(mined)-199(position)-199(by)-199(measuring)-199(the)]TJ -1 -1.111 Td [(dist)-30(ance)-330(to)-330(yet)-330(another)-330(k)-30(nown)-330(point)-330(w)-40(ithin)-330(the)-330(str)-30(ucture.)-330(To)-330(this)]TJ T* [(end,)-441(we)-441(used)-441(another)-441(single-c)-30(ysteine)-441(mut)-30(ant)-441(of)-441(Rpb7\227namely,)]TJ T* [(Rpb7\226C94)-591(\(Fig.)-591(1\)\227and)-591(measured)-591(the)-591(dist)-30(ance)-591(bet)-30(ween)-591(the)]TJ T* [(ac)-30(ceptor)-255(att)-30(ached)-255(to)-255(this)-255(position)-255(and)-255(the)-255(donor)-255(on)-255(the)-255(RNA.)-255(The)]TJ T* [(measured)-519(dist)-30(ance)-519(then)-519(c)-30(ould)-519(be)-519(c)-30(ompared)-519(w)-40(ith)-519(the)-519(dist)-30(ance)]TJ T* [(bet)-30(ween)-449(the)-449(intersection)-449(point)-449(deter)-30(mined)-449(by)-449(our)-449(triangulation)]TJ T* [(\(using)-246(DNA1,)-246(DNA)-60(2,)-246(and)-246(Rpb7\226C150\))-246(and)-246(the)-246(position)-246(of)-246(Rpb7\226)]TJ 0 -1.111 TD [(C94)-213(f)-30(rom)-213(the)-213(x-ray)-213(str)-30(ucture.)-213(The)-213(c)-30(omparison)-213(yielded)-213(dev)-40(iations)-213(of)]TJ 0 -1.111 TD [(on)-249(average)-249(2.5)-249(\305)-249(\(never)]TJ /T1_1 1 Tf 0 Tc 10.128 0 Td (\005)Tj /T1_0 1 Tf 0.2342 Tc 0.818 0 Td [(5\305)249(;)]TJ 0 0 1 rg -0.0149 Tc 1.995 0 Td [(SI)-249(Table)-249(2)]TJ 0 0 0 rg 4.255 0 Td [(\),)-249(thus)-249(prov)-40(iding)-249(another)]TJ -17.197 -1.111 Td [(e)20(stimate)-318(of)-318(our)-318(ex)-30(periment)-30(al)-318(error.)]TJ 1 -1.111 Td [(Further)-30(more,)-225(trilateration)-225(allows)-225(for)-225(a)-225(self-c)-30(onsistent)-225(te)20(st)-225(for)-225(the)]TJ -1 -1.111 Td [(ac)-30(curac)-30(y)-435(of)-435(the)-435(sp-FR)-60(ET)-435(dist)-30(ance)-435(deter)-30(mination.)-435(If)-435(one)-435(deter-)]TJ T* [(mine)20(s)-407(four)-407(dist)-30(ance)20(s,)-407(as)-407(de)20(scribed)-407(above,)-407(one)-407(can)-407(per)-50(for)-30(m)-407(four)]TJ T* [(independent)-325(three-dist)-30(ance)-325(trilaterations)-325(and)-325(c)-30(ompare)-325(the)-325(re)20(sult-)]TJ T* [(ing)-299(four)-299(intersection)-299(points.)-299(A)-30(n)-299(example)-299(of)-299(this)-299(procedure)-299(for)-299(the)]TJ T* [(case)-295(of)-295(RNA)-60(29)-295(is)-295(shown)-295(in)]TJ 0 0 1 rg 11.699 0 Td [(SI)-295(Fig.)-295(11)]TJ 0 0 0 rg 4.032 0 Td [(.)-295(The)-295(four)-295(positions)-295(that)-295(are)]TJ -15.73 -1.111 Td [(deter)-30(mined)-220(by)-220(the)-220(four)-220(trilaterations)-220(are)-220(all)-220(w)-40(ithin)]TJ /T1_1 1 Tf 0 Tc 20.812 0 Td (\004)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(3)-220(\305)-220(f)-30(rom)-220(each)]TJ -21.63 -1.111 Td [(other,)-393(thus)-393(prov)-40(iding)-393(another)-393(indication)-393(for)-393(the)-393(ac)-30(curac)-30(y)-393(of)-393(our)]TJ T* (method.)Tj /T1_3 1 Tf 0 Tc 8 0 0 8 314.1761 184.8835 Tm [(RNA)-505(Traverses)-505(the)-505(Exit)-505(Tunnel)-505(and)-505(Disengages)-505(from)-505(the)-505(Surface.)]TJ /T1_0 1 Tf -0.0149 Tc 9 0 0 9 314.1761 174.8835 Tm [(Trilateration)-294(analysis)-294(for)-294(RNA17,)-294(RNA)-60(20,)-294(and)-294(RNA)-60(23)-294(yields)-294(the)]TJ 0 -1.111 TD [(c)-30(orre)20(sponding)-464(positions)-464(of)-464(the)-464(5)]TJ /T1_1 1 Tf 0 Tc 14.136 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc 0.782 0 Td [(end)-464(\(Fig.)-464(4\).)-464(To)-464(ac)-30(c)-30(ount)-464(for)]TJ -14.918 -1.111 Td [(ex)-30(periment)-30(al)-275(errors,)-275(the)-275(positions)-275(are)-275(repre)20(sented)-275(by)-275(sphere)20(s)-275(w)-40(ith)]TJ T* [(a)-402(radius)-402(o)]TJ 0.3876 Tc 4.352 0 Td [(f5\305\()403(s)402(e)403(e)]TJ /T1_2 1 Tf -0.0149 Tc 4.856 0 Td [(Mater)-30(ials)-402(and)-402(Methods)]TJ /T1_0 1 Tf 9.499 0 Td [(\).)-402(The)-402(5)]TJ /T1_1 1 Tf 0 Tc 3.549 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc 0.721 0 Td [(end)-402(of)-402(the)]TJ -22.976 -1.111 Td [(RNA17)-340(is)-340(located)-340(near)-340(the)-340(end)-340(of)-340(the)-340(proposed)-340(ex)-50(it)-340(tunnel)-340(\(Fig.)]TJ 0 Tc 0 -1.111 TD (4)Tj /T1_2 1 Tf 0.575 0 Td (A)Tj /T1_0 1 Tf -0.0149 Tc 0.675 0 Td [(\).)-286(Modeling)-286(shows)-286(that)-286(this)-286(position)-286(of)-286(RNA17)-286(5)]TJ /T1_1 1 Tf 0 Tc 20.331 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc 0.604 0 Td [(end)-286(can)-286(only)]TJ -22.185 -1.111 Td [(be)-476(ex)-30(plained)-476(if)-476(the)-476(RNA)-476(extends)-476(f)-30(rom)-476(the)-476(cr)-60(yst)-30(allog)-20(raphically)]TJ T* [(obser)-60(ved)-215(position)]TJ /T1_1 1 Tf 0 Tc 7.523 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(10)-215(through)-215(the)-215(prev)-40(iously)-215(proposed)-215(ex)-50(it)-215(tunnel)]TJ -8.341 -1.111 Td [(beneath)-278(the)-278(poly)-30(merase)-278(lid)-278(\(Fig.)-278(4\).)-278(Thus,)-278(the)20(se)-278(dat)-30(a)-278(c)-30(onfir)-30(m)-278(that)]TJ T* [(the)-230(RNA)-230(leave)20(s)-230(the)-230(Pol)-230(II)-230(c)-30(ore)-230(through)-230(the)-230(ex)-50(it)-230(tunnel)-230(as)-230(had)-230(been)]TJ T* [(shown)-354(prev)-40(iously)-354(for)-354(the)-354(bacterial)-354(RNA)-354(poly)-30(merase)-354(\(8,)-354(22\).)-354(The)]TJ T* [(dist)-30(ance)-259(bet)-30(ween)-259(the)-259(last)-259(nucleotide)-259(v)-40(isible)-259(in)-259(the)-259(cr)-60(yst)-30(al)-259(str)-30(ucture)]TJ ET q 244.8000031 0 0 225.9600067 52.9261017 514.8834991 cm /Im0 Do Q 0 0 0 1 k BT /T1_5 1 Tf 0 Tc 7 0 0 7 51.6761 500.8835 Tm [(Fig.)-280(1.)]TJ /T1_6 1 Tf 4.062 0 Td [(Pol)-280(II)-280(structure)-280(and)-280(labeling.)-280(\()]TJ /T1_7 1 Tf 13.738 0 Td (A)Tj /T1_6 1 Tf 0.722 0 Td [(\))-280(Schematic)-280(showing)-280(the)-280(oligonucle-)]TJ -18.522 -1.286 Td [(otides)-269(used)-269(in)-269(the)-269(single-molecule)-269(experiments)-269(for)-269(the)-269(formation)-269(of)-269(elonga-)]TJ 0 -1.286 TD [(tion)-159(complexes.)-159(Filled)-159(and)-159(open)-159(circles)-159(denote)-159(nucleotides)-159(whose)-159(positions)-159(are)]TJ T* [(known)-379(and)-379(unknown,)-379(respectively,)-379(from)-379(crystallographic)-379(studies)-379(\(3\).)-379(High-)]TJ T* [(lighted)-195(are)-195(the)-195(labeling)-195(positions)-195(on)-195(the)-195(template)-195(DNA)-195(and)-195(RNA.)-195(\()]TJ /T1_7 1 Tf 30.092 0 Td (B)Tj /T1_6 1 Tf 0.611 0 Td [(\))-195(Top)-195(view)]TJ -30.703 -1.286 Td [(of)-336(the)-336(complete)-336(Pol)-336(II)-336(elongation)-336(complex)-336(structure)-336(\(3\),)-336(indicating)-336(labeling)]TJ 0 -1.286 TD [(sites.)-164(The)-164(core)-164(polymerase)-164(is)-164(shown)-164(in)-164(surface)-164(representation)-164(\(in)-164(gray\),)-164(and)-164(the)]TJ T* [(backbone)-376(of)-376(the)-376(template)-376(DNA)-376(\(blue\),)-376(nontemplate)-376(DNA)-376(\(cyan\),)-376(and)-376(RNA)]TJ T* [(\(red\))-173(and)-173(the)-173(heterodimer)-173(Rpb4/7)-173(\(Rpb4)-173(in)-173(red)-173(and)-173(Rpb7)-173(in)-173(blue\))-173(are)-173(displayed)]TJ T* [(with)-205(cartoon)-205(diagrams.)-205(The)-205(positions)-205(that)-205(were)-205(used)-205(to)-205(attach)-205(dye)-205(molecules,)]TJ T* [(that)-178(is,)-178(Rpb7,)-178(residues)-178(Rpb7\226C94,)-178(and)-178(Rpb7\226C150,)-178(as)-178(well)-178(as)-178(the)-178(template)-178(DNA)]TJ T* [(at)-371(position)]TJ /T1_1 1 Tf 5.466 0 Td (\002)Tj /T1_6 1 Tf 0.833 0 Td [(10)-371(\(DNA1\))-371(and)]TJ /T1_1 1 Tf 7.392 0 Td (\001)Tj /T1_6 1 Tf 0.833 0 Td [(3)-371(\(DNA2\),)-371(are)-371(shown)-371(in)-371(green.)-371(At)]TJ /T1_7 1 Tf 16.379 0 Td (Right)Tj /T1_6 1 Tf 2.871 0 Td [(is)-371(a)]TJ -33.774 -1.286 Td [(close-up)-478(view)-478(of)-478(the)-478(labeling)-478(region)-478(highlighting)-478(the)-478(distances)-478(between)]TJ T* [(Rpb7\226C150)-278(and)-278(DNA1)-278(as)-278(well)-278(as)-278(Rpb7\226C150)-278(and)-278(DNA2.)]TJ /T1_5 1 Tf 6.5 0 0 6.5 51.6761 39.8835 Tm (136)Tj /T1_8 1 Tf 9.75 0 0 6.5 68.5185 39.8835 Tm (\001)Tj /T1_6 1 Tf 6.5 0 0 6.5 76.6831 39.8835 Tm (www.pnas.org)Tj /T1_8 1 Tf 6.833 0 Td (\002)Tj /T1_6 1 Tf 0.5 0 Td (cgi)Tj /T1_8 1 Tf 1.333 0 Td (\002)Tj /T1_6 1 Tf 0.5 0 Td (doi)Tj /T1_8 1 Tf 1.5 0 Td (\002)Tj /T1_6 1 Tf 0.5 0 Td (10.1073)Tj /T1_8 1 Tf 3.614 0 Td (\002)Tj /T1_6 1 Tf 0.5 0 Td (pnas.0703815105)Tj 52.294 0 Td (Andrecka)Tj /T1_7 1 Tf 4.723 0 Td [(et)-278(al)]TJ /T1_6 1 Tf 2.057 0 Td (.)Tj ET q /GS1 gs 19 0 0 815.5 9 9 cm /Im1 Do Q endstream endobj 127 0 obj<> endobj 128 0 obj<>stream 0 0 0 0.2 k 1 i /GS0 gs 573.676 521.384 38.324 -66 re f 0 0 0 1 k BT /T1_0 1 Tf -0.0149 Tc 0 Tw 0 Ts 100 Tz 0 Tr 9 0 0 9 51.6761 445.8835 Tm [(at)-514(position)]TJ /T1_1 1 Tf 0 Tc 5.139 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(10)-514(and)-514(the)-514(obser)-60(ved)-514(position)-514(at)]TJ /T1_1 1 Tf 0 Tc 14.8 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(17)-514(is)]TJ /T1_1 1 Tf 0 Tc 2.624 0 Td (\004)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(28)-514(\305,)]TJ -25.017 -1.135 Td [(indicating)-187(that)-188(the)-187(RNA)-187(is)-187(in)-187(a)-187(st)-30(acked)-187(c)-30(onfor)-30(mation)-187(w)-40(ithin)-187(the)-187(ex)-50(it)]TJ 0 -1.135 TD [(tunnel)-501(as)-501(was)-501(obser)-60(ved)-501(recently)-501(in)-501(the)-501(cr)-60(yst)-30(al)-501(str)-30(ucture)-501(of)-501(an)]TJ T* [(elongation)-324(c)-30(omplex)-324(of)-324(a)-324(bacterial)-324(RNA)-324(poly)-30(merase)-324(\(22\).)-324(Beyond)]TJ T* [(the)-218(tunnel,)-218(for)-218(the)-218(lengths)-218(of)-218(17\22623)-218(nt,)-218(the)-218(RNA)-218(is)-218(c)-30(ompact)-218(and)-218(the)]TJ 0 -1.135 TD [(dist)-30(ance)-192(bet)-30(ween)-192(RNA17)-192(and)-192(RNA)-60(23)-192(is)-192(only)]TJ /T1_1 1 Tf 0 Tc 18.695 0 Td (\004)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(11)-192(\305.)-192(In)-192(this)-192(region,)]TJ -19.514 -1.135 Td [(the)-407(RNA)-407(end)-407(is)-407(too)-407(far)-407(f)-30(rom)-407(the)-407(poly)-30(merase)-407(sur)-50(face)-407(for)-407(direct)]TJ 0 -1.134 TD [(interactions)-318(\(Fig.)-318(4)]TJ /T1_2 1 Tf 0 Tc 7.915 0 Td (B)Tj /T1_0 1 Tf -0.0149 Tc 0.638 0 Td (\).)Tj /T1_3 1 Tf 0 Tc 8 0 0 8 314.1761 445.8835 Tm [(Extending)-279(RNA)-279(Associates)-279(with)-279(Pol)-279(II)-279(Before)-279(It)-279(Becomes)-279(Flexible.)]TJ /T1_0 1 Tf -0.0149 Tc 9 0 0 9 548.058 445.8835 Tm (For)Tj -0.013 Tc -25.987 -1.135 Td [(elongation)-593(c)-30(omplexe)20(s)-593(c)-30(ont)-30(ain)-30(ing)-593(RNA)-60(26,)-593(the)-593(single-molecule)]TJ -0.0149 Tc 0 -1.135 TD [(FR)-60(ET)-373(measurements)-373(re)20(sulted)-373(in)-373(histog)-20(rams)-373(that)-373(c)-30(ould)-373(be)-373(fitted)]TJ T* [(w)-40(ith)-241(a)-241(double)-241(Gaussian)-241(fit)-241(\(Fig.)-241(2)]TJ /T1_2 1 Tf 0 Tc 13.702 0 Td (D)Tj /T1_0 1 Tf -0.0149 Tc 0.74 0 Td [(\),)-241(thus)-241(yielding)-241(t)-30(wo)-241(value)20(s)-241(for)]TJ /T1_2 1 Tf 0 Tc 12.3 0 Td (d)Tj /T1_0 1 Tf 0.513 0 Td (.)Tj -0.0149 Tc -27.255 -1.135 Td [(Therefore,)-426(trilateration)-426(doe)20(s)-426(not)-426(yield)-426(a)-426(single)-426(position)-426(but)-426(t)-30(wo)]TJ 0 -1.135 TD [(positions)-318(bet)-30(ween)-318(which)-318(the)-318(RNA)-318(end)-318(is)-318(f)-90(luctuating)-318(\(Fig.)-318(4\).)]TJ 1 -1.135 Td [(A)-30(lthough)-467(for)-467(short)-467(lengths,)-467(the)-467(RNA)-467(lacks)-467(c)-30(ont)-30(act)-467(w)-40(ith)-467(the)]TJ -1 -1.134 Td [(sur)-50(face)-245(of)-245(the)-245(poly)-30(merase,)-245(at)-245(a)-245(length)-245(of)-245(26)-245(nt,)-245(the)-245(RNA)-245(associate)20(s)]TJ ET q 384 0 0 189.9600067 114.6761017 550.8834991 cm /Im0 Do Q BT /T1_4 1 Tf 0 Tc 7 0 0 7 51.6761 536.8835 Tm [(Fig.)-272(2.)]TJ /T1_5 1 Tf 4.046 0 Td [(sp-FRET)-272(time)-272(traces)-272(and)-272(histograms.)-272(Exemplary)-272(time)-272(traces)-272(and)-272(complete)-272(histograms)-272(for)-272(the)-272(FRET)-272(pairs)-272(DNA1\226RNA17)-272(\()]TJ /T1_6 1 Tf 54.924 0 Td (A)Tj /T1_5 1 Tf 0.994 0 Td (and)Tj /T1_6 1 Tf 2.05 0 Td (B)Tj /T1_5 1 Tf 0.611 0 Td [(\))-272(and)-272(DNA1\226RNA26)-272(\()]TJ /T1_6 1 Tf 9.65 0 Td (C)Tj /T1_5 1 Tf -72.275 -1.286 Td (and)Tj /T1_6 1 Tf 2.027 0 Td (D)Tj /T1_5 1 Tf 0.722 0 Td [(\).)-249(The)-249(time)-249(traces)-249(show)-249(the)-249(\037uorescence)-249(intensities)-249(of)-249(the)-249(donor)-249(\(green\))-249(and)-249(the)-249(acceptor)-249(\(red\))-249(\(thin)-249(lines)-249(correspond)-249(to)-249(the)-249(actual)-249(signal)-249(and)-249(thick)-249(li)]TJ 68.573 0 Td (nes)Tj -71.322 -1.286 Td [(correspond)-258(to)-258(a)-258(10-point)-258(sliding)-258(average\))-258(as)-258(well)-258(as)-258(the)-258(computed)-258(FRET)-258(signal)-258(\(blue\).)-258(\()]TJ /T1_6 1 Tf 39.837 0 Td (A)Tj /T1_5 1 Tf 0.722 0 Td [(\))-258(The)-258(trajectory)-258(of)-258(the)-258(RNA17\226DNA1)-258(pair)-258(shows)-258(constant)-258(\037uorescence)]TJ -40.559 -1.286 Td [(intensities)-261(for)-261(the)-261(donor)-261(and)-261(acceptor)-261(molecules)-261(until)-261(bleaching)-261(of)-261(the)-261(acceptor)-261(after)]TJ /T1_1 1 Tf 39.838 0 Td (\004)Tj /T1_5 1 Tf 0.833 0 Td [(12)-261(s.)-261(At)-261(this)-261(point,)-261(the)-261(donor)-261(intensity)-261(increases)-261(and)-261(remains)-261(constant)]TJ -40.671 -1.286 Td [(until)-203(the)-203(donor)-203(itself)-203(photo-bleaches)-203(after)]TJ /T1_1 1 Tf 19.499 0 Td (\004)Tj /T1_5 1 Tf 0.833 0 Td [(22)-203(s.)-203(The)-203(computed)-203(FRET)-203(ef\036ciency)-203(is)-203(constant,)-203(except)-203(for)-203(small)-203(\037uctuations)-203(attributable)-203(to)-203(photon-counting)-203(noise.)]TJ -20.332 -1.286 Td (\()Tj /T1_6 1 Tf 0.333 0 Td (B)Tj /T1_5 1 Tf 0.611 0 Td [(\))-227(The)-227(histogram)-227(of)-227(337)-227(such)-227(sp-FRET)-227(trajectories)-227(shows)-227(a)-227(single)-227(peak,)-227(which)-227(can)-227(be)-227(\036tted)-227(with)-227(a)-227(Gaussian)-227(distribution)-227(that)-227(is)-227(centered)-227(at)]TJ /T1_6 1 Tf 62.467 0 Td (E)Tj /T1_1 1 Tf 0.783 0 Td (\006)Tj /T1_5 1 Tf 1.06 0 Td [(0.92.)-227(In)-227(contrast,)]TJ -65.254 -1.286 Td [(the)-331(FRET)-331(ef\036ciency)-331(in)]TJ /T1_6 1 Tf 10.436 0 Td (C)Tj /T1_5 1 Tf 0.942 0 Td [(for)-331(the)-331(DNA1\226RNA26)-331(sample)-331(is)-331(not)-331(constant)-331(but)-331(varies)-331(between)]TJ /T1_6 1 Tf 30.59 0 Td (E)Tj /T1_1 1 Tf 0.887 0 Td (\004)Tj /T1_5 1 Tf 1.164 0 Td [(0.85)-331(and)]TJ /T1_6 1 Tf 4.386 0 Td (E)Tj /T1_1 1 Tf 0.887 0 Td (\004)Tj /T1_5 1 Tf 1.164 0 Td [(0.6.)-331(Accordingly)-331(the)-331(histogram)-331(in)]TJ /T1_6 1 Tf 15.989 0 Td (D)Tj /T1_5 1 Tf 1.053 0 Td [(of)-331(224)-331(such)]TJ -67.497 -1.286 Td [(trajectories)-278(can)-278(be)-278(\036tted)-278(with)-278(two)-278(Gaussian)-278(\036ts)-278(centered)-278(at)]TJ /T1_6 1 Tf 27.952 0 Td (E)Tj /T1_1 1 Tf 0.834 0 Td (\006)Tj /T1_5 1 Tf 1.111 0 Td [(0.58)-278(and)]TJ /T1_6 1 Tf 4.28 0 Td (E)Tj /T1_1 1 Tf 0.834 0 Td (\006)Tj /T1_5 1 Tf 1.111 0 Td (0.85.)Tj /T1_4 1 Tf 8 0 0 8 90.6761 346.8835 Tm [(Table)-333(1.)-333(Overview)-333(of)-333(the)-333(results)-333(of)-333(the)-333(sp-FRET)-333(measurements)]TJ ET 0 0 0 1 K 0.5 w 10 M 0 j 0 J []0 d 90.679 102.133 m 522.67 102.133 l 90.679 327.634 m 522.67 327.634 l S BT /T1_5 1 Tf 7.5 0 0 7.5 90.679 332.8835 Tm [(Donor)-278(\(TMR\))-2955(Acceptor)-278(\(Alexa)-278(647\))-3136(FRET,)-278(%)-3780(Width)]TJ /T1_6 1 Tf 35.792 0 Td (R)Tj /T1_5 1 Tf 5.5 0 0 5 363.699 331.8835 Tm (0)Tj 7.5 0 0 7.5 366.7573 332.8835 Tm [(,)-278(\305*)-2944(Distance,)-278(\305)-2943(No.)-278(of)-278(molecules)]TJ -36.81 -2.067 Td [(RNA17)-9259(DNA1)-7527(92.1)-5168(0.24)-4926(57)-5870(38)-8055(337)]TJ 0 -1.333 TD [(RNA17)-9259(DNA2)-7527(55.8)-5168(0.19)-4926(57)-5870(55)-8055(336)]TJ T* [(RNA17)-8036(Rpb7\226C150)-6304(69.1)-5168(0.28)-4926(59)-5870(52)-8055(323)]TJ 0 -1.333 TD [(RNA17)-8314(Rpb7\226C94)-6583(68.7)-5168(0.38)-4926(60)-5870(52)-8610(91)]TJ T* [(RNA20)-9259(DNA1)-7527(87.0)-5168(0.21)-4926(60)-5870(43)-8055(227)]TJ T* [(RNA20)-9259(DNA2)-7527(49.6)-5168(0.19)-4926(60)-5870(60)-8055(412)]TJ T* [(RNA20)-8036(Rpb7\226C150)-6304(67.9)-5168(0.23)-4926(62)-5870(55)-8055(226)]TJ T* [(RNA20)-8314(Rpb7\226C94)-6583(63.8)-5168(0.2)-5482(63)-5870(57)-8610(76)]TJ T* [(RNA23)-9259(DNA1)-7527(81.4)-5168(0.19)-4926(59)-5870(46)-8055(356)]TJ T* [(RNA23)-9259(DNA2)-7527(43.3)-5168(0.14)-4926(59)-5870(62)-8055(361)]TJ T* [(RNA23)-8036(Rpb7\226C150)-6304(69.1)-5168(0.3)-5482(62)-5870(54)-8055(230)]TJ 0 -1.333 TD [(RNA23)-8314(Rpb7\226C94)-6583(67.4)-5168(0.23)-4926(62)-5870(55)-8055(105)]TJ T* (RNA26)Tj 5.5 0 0 5 114.4315 199.6335 Tm (\206)Tj 7.5 0 0 7.5 183.8778 197.3835 Tm [(DNA1)-6416(57.8/84.6)-2944(0.21/0.23)-3814(53)-5175(50/40)-7359(224)]TJ -12.427 -1.333 Td (RNA26)Tj 5.5 0 0 5 114.4315 189.6335 Tm (\206)Tj 7.5 0 0 7.5 183.8778 187.3835 Tm [(DNA2)-6416(42.3/58.8)-2944(0.19/0.35)-3814(53)-5175(55/50)-7359(272)]TJ -12.427 -1.333 Td (RNA26)Tj 5.5 0 0 5 114.4315 179.6335 Tm (\206)Tj 7.5 0 0 7.5 174.7045 177.3835 Tm [(Rpb7\226C150)-5193(55.9/85.5)-2944(0.25/0.29)-3814(55)-5175(53/41)-7915(96)]TJ -11.203 -1.333 Td (RNA26)Tj 5.5 0 0 5 114.4315 169.6335 Tm (\206)Tj 7.5 0 0 7.5 176.7869 167.3835 Tm [(Rpb7\226C94)-5471(50.0/79.3)-3221(0.28/0.2)-4092(55)-5175(55/44)-7915(92)]TJ -11.481 -1.333 Td [(RNA29)-9259(DNA1)-7527(67.2)-5168(0.18)-4926(62)-5870(55)-8055(305)]TJ 0 -1.333 TD [(RNA29)-9259(DNA2)-7527(42.7)-5168(0.18)-4926(62)-5870(65)-8055(374)]TJ T* [(RNA29)-8036(Rpb7\226C150)-6304(59.4)-5168(0.19)-4926(65)-5870(61)-8055(148)]TJ T* [(RNA29)-8314(Rpb7\226C94)-6583(51.2)-5168(0.17)-4926(66)-5870(65)-8610(96)]TJ 0 -1.333 TD [(DNA1)-8481(Rpb7\226C150)-6304(56.2)-5168(0.24)-4926(63)-5870(61)-8055(281)]TJ T* [(DNA2)-8481(Rpb7\226C150)-6304(50.6)-5168(0.36)-4926(63)-5870(63)-8610(50)]TJ 7 0 0 7 99.6761 91.8835 Tm [(The)-308(displayed)-308(values)-308(are)-308(the)-308(results)-308(of)-308(\036ts)-308(of)-308(single)-308(Gaussian)-308(distributions)-308(\(double)-308(Gaussian)-308(distributions)-308(where)-308(indicated\))-308(to)-308(the)]TJ -1.286 -1.286 Td [(experimentally)-278(recorded)-278(histograms)-278(of)-278(sp-FRET)-278(ef\036ciencies,)-278(with)-278(the)-278(width)-278(given)-278(by)-278(the)-278(standard)-278(deviation.)]TJ 0 -1.286 TD (*Accuracy)Tj /T1_1 1 Tf 4.944 0 Td (\007)Tj /T1_5 1 Tf 0.278 Tc 1.111 0 Td [(2\305)278(.)]TJ 0 Tc 5 0 0 5 90.6761 67.1335 Tm (\206)Tj 7 0 0 7 93.4565 64.8835 Tm [(Two)-278(peaks)-278(found.)]TJ 6.5 0 0 6.5 51.6761 39.8835 Tm (Andrecka)Tj /T1_6 1 Tf 4.75 0 Td [(et)-305(al)]TJ /T1_5 1 Tf 2.084 0 Td (.)Tj 45.416 0 Td (PNAS)Tj /T1_7 1 Tf 9.75 0 0 6.5 413.5518 39.8835 Tm (\001)Tj /T1_4 1 Tf 6.5 0 0 6.5 421.7163 39.8835 Tm [(January)-305(8,)-305(2008)]TJ /T1_7 1 Tf 9.75 0 0 6.5 475.4029 39.8835 Tm (\001)Tj /T1_5 1 Tf 6.5 0 0 6.5 483.5675 39.8835 Tm [(vol.)-305(105)]TJ /T1_7 1 Tf 9.75 0 0 6.5 513.2288 39.8835 Tm (\001)Tj /T1_5 1 Tf 6.5 0 0 6.5 521.3933 39.8835 Tm [(no.)-305(1)]TJ /T1_7 1 Tf 9.75 0 0 6.5 542.7408 39.8835 Tm (\001)Tj /T1_4 1 Tf 6.5 0 0 6.5 550.9054 39.8835 Tm (137)Tj /T1_3 1 Tf 0 6.5 -6.5 0 585.1761 471.2386 Tm (BIOPHYSICS)Tj ET q /GS1 gs 19 0 0 815.5 9 9 cm /Im1 Do Q endstream endobj 129 0 obj<> endobj 130 0 obj<>stream 0 0 0 1 k 1 i /GS0 gs BT /T1_0 1 Tf -0.0149 Tc 0 Tw 0 Ts 100 Tz 0 Tr 9 0 0 9 51.6761 469.8835 Tm [(w)-40(ith)-197(the)-197(sur)-50(face)-197(of)-197(the)-197(poly)-30(merase)-197(in)-197(the)-197(region)-197(of)-197(the)-197(dock)-197(domain)]TJ 0 -1.091 TD [(\(Fig.)-487(4\).)-487(Further)-30(more,)-487(the)-487(c)-30(onfor)-30(mation)-487(of)-487(the)-487(RNA)-487(bet)-30(ween)]TJ T* (positions)Tj /T1_1 1 Tf 0 Tc 4.059 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(23)-370(and)]TJ /T1_1 1 Tf 0 Tc 3.254 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(29)-370(is)-370(more)-370(extended,)-370(because)-370(the)-370(5)]TJ /T1_1 1 Tf 0 Tc 15.095 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc 0.688 0 Td [(end)-370(of)]TJ -24.732 -1.091 Td [(RNA)-60(26)-360(is)-360(located)-360(15)-360(\305)-360(or)-360(20)-360(\305)-360(f)-30(rom)-360(the)-360(end)-360(of)-360(RNA)-60(23)-360(and)-360(it)]TJ T* [(advance)20(s)-195(another)-195(17)-195(\305)-195(or)-195(21)-195(\305)-195(bet)-30(ween)-195(the)-195(positions)]TJ /T1_1 1 Tf 0 Tc 21.743 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(26)-195(and)]TJ /T1_1 1 Tf 0 Tc 2.904 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td (29.)Tj -25.283 -1.091 Td [(For)-251(the)-251(even)-251(longer)-251(RNA32)-251(and)-251(RNA35,)-251(the)-251(histog)-20(rams)-251(can)-251(no)]TJ -1 -1.091 Td [(longer)-191(be)-191(fitted)-191(w)-40(ith)-191(a)-191(single)-191(or)-191(double)-191(Gaussian)-191(fit)-191(\()]TJ 0 0 1 rg 21.217 0 Td [(SI)-191(Fig.)-191(12)]TJ 0 0 0 rg 3.824 0 Td [(\).)-191(The)]TJ 0 Tc -25.041 -1.091 Td (5)Tj /T1_1 1 Tf 0.485 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc 0.595 0 Td [(end)-277(is)-277(f)-90(lex)-50(ible)-277(and)-277(no)-277(longer)-277(oc)-30(cupie)20(s)-277(well)-277(defined)-277(locations)-277(on)]TJ -1.08 -1.091 Td [(the)-318(Pol)-318(II)-318(sur)-50(face.)]TJ /T1_2 1 Tf -0.0196 Tc 0 -1.98 TD (Discussion)Tj /T1_0 1 Tf -0.0149 Tc 0 -1.313 TD [(We)-230(have)-230(used)-230(single-particle)-230(f)-90(luore)20(scence)-230(spectrosc)-30(opy)-230(to)-230(map)-230(the)]TJ 0 -1.091 TD [(path)-420(of)-420(mRNA)-420(ex)-50(iting)-420(f)-30(rom)-420(Pol)-420(II.)-420(RNA)-420(ex)-50(its)-420(f)-30(rom)-420(the)-420(active)]TJ 29.167 42.556 Td [(center)-217(clef)-30(t)-217(by)-217(traversing)-217(the)-217(prev)-40(iously)-217(proposed)-217(ex)-50(it)-217(tunnel.)-217(RNA)]TJ 0 -1.12 TD [(then)-420(c)-30(ontinue)20(s)-420(toward)-420(the)-420(dock)-420(domain)-420(w)-40(ithout)-420(apparent)-420(poly-)]TJ T* [(merase)-271(interactions.)-271(Once)-271(the)-271(RNA)-270(has)-271(reached)-271(a)-271(length)-271(of)-271(26)-271(nt,)]TJ T* [(it)-418(make)20(s)-418(direct)-418(c)-30(ont)-30(act)-418(w)-40(ith)-418(the)-418(dock)-418(domain,)-418(follow)-40(ing)-418(a)-418(path)]TJ T* [(bet)-30(ween)-339(the)-339(t)-30(wo)-339(g)-20(roove)20(s)-339(prev)-40(iously)-339(proposed)-339(to)-339(possibly)-339(ac)-30(c)-30(om-)]TJ T* [(modate)-344(RNA.)-344(RNA)-344(further)-344(extends)-344(toward)-344(the)-344(beginn)-30(ing)-344(of)-344(the)]TJ T* [(linker)-318(that)-318(c)-30(onnects)-318(to)-318(the)-318(CTD.)]TJ 1 -1.12 Td [(The)-522(position)-522(of)-522(the)-522(5)]TJ /T1_1 1 Tf 0 Tc 9.719 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc 0.841 0 Td [(end)-522(of)-522(RNA17)-522(deter)-30(mined)-522(here,)-522(in)]TJ -11.56 -1.12 Td [(c)-30(omparison)-235(w)-40(ith)-235(the)-235(RNA)-235(position)-235(at)-235(register)]TJ /T1_1 1 Tf 0 Tc 19.111 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(10)-235(obser)-60(ved)-235(in)-235(the)]TJ -19.93 -1.12 Td [(cr)-60(yst)-30(al)-307(str)-30(ucture,)-307(shows)-307(that)-307(the)-307(RNA)-307(traverse)20(s)-307(the)-307(proposed)-307(ex)-50(it)]TJ T* [(tunnel.)-249(It)-249(has)-249(long)-249(been)-249(k)-30(nown)-249(that)-249(single-stranded)-249(RNA)-249(interacts)]TJ T* [(w)-40(ith)-181(poly)-30(merase)-181(around)-181(positions)]TJ /T1_1 1 Tf 0 Tc 13.95 0 Td (\002)Tj /T1_0 1 Tf 0.1657 Tc 0.818 0 Td [(8t)181(o)]TJ /T1_1 1 Tf 0 Tc 1.687 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(15)-181(and)-181(that)-181(this)-181(upstream)]TJ -17.274 -1.12 Td [(RNA-binding)-264(site)-264(c)-30(ontribute)20(s)-264(to)-264(elongation)-264(c)-30(omplex)-264(st)-30(abilit)-30(y)-264(\(23\).)]TJ 0 -1.12 TD [(Our)-211(dat)-30(a)-211(prov)-40(ide)-211(further)-211(ev)-40(idence)-211(that)-211(the)-211(region)-211(in)-211(the)-211(ex)-50(it)-211(tunnel)]TJ T* [(acts)-318(as)-318(the)-318(prev)-40(iously)-318(proposed)-318(RNA)-318(binding)-318(site)-318(\(23\).)]TJ 1 -1.12 Td [(As)-507(the)-507(RNA)-507(g)-20(rows)-507(longer,)-507(the)-507(single-particle)-507(dat)-30(a)-507(reveal)-507(a)]TJ -1 -1.12 Td [(change)-305(f)-30(rom)-305(a)-305(c)-30(ompacted)-305(RNA)-305(str)-30(ucture)-305(\(RNA17\226RNA)-60(23\))-305(to)-305(a)]TJ 0 -1.12 TD [(more)-226(extended)-226(str)-30(ucture)-226(\(RNA)-60(26)-90(\226RNA)-60(29\).)-226(The)-226(RNA)-226(apparently)]TJ T* [(lacks)-378(direct)-378(c)-30(ont)-30(acts)-378(w)-40(ith)-378(the)-378(poly)-30(merase)-378(beyond)-378(the)-378(tunnel)-378(but)]TJ T* [(reassociate)20(s)-304(w)-40(ith)-304(the)-304(enz)-30(y)-30(me)-304(sur)-50(face)-304(at)-304(register)]TJ /T1_1 1 Tf 0 Tc 20.263 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(26)-304(and)-304(remains)]TJ -21.081 -1.12 Td [(sur)-50(face-associated)-263(at)-263(register)]TJ /T1_1 1 Tf 0 Tc 12.239 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(29.)-263(Reassociation)-263(of)-263(the)-263(RNA)-263(may)]TJ -13.057 -1.12 Td [(ac)-30(c)-30(ount)-213(for)-213(sequence-independent)-213(transcriptional)-213(pausing)-213(that)-213(had)]TJ 0 -1.12 TD [(been)-258(obser)-60(ved)-258(prev)-40(iously)-258(at)-258(around)-258(register)]TJ /T1_1 1 Tf 0 Tc 18.43 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(25)-258(\(24\).)-258(Poly)-30(merase)]TJ -19.248 -1.12 Td [(association)-361(of)-361(the)-361(RNA)-361(around)-361(registers)]TJ /T1_1 1 Tf 0 Tc 17.58 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(26)-361(and)]TJ /T1_1 1 Tf 0 Tc 3.235 0 Td (\002)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(29)-361(also)-361(may)]TJ -22.451 -1.12 Td [(ac)-30(c)-30(ount)-237(for)-237(the)-237(prevention)-237(of)-237(transcript)-237(slippage)-237(at)-237(the)20(se)-237(transcript)]TJ 0 -1.12 TD [(lengths)-276(\(25\).)-276(Prev)-40(ious)-276(biochemical)-276(studie)20(s)-276(also)-276(have)-276(shown)-276(indica-)]TJ T* [(tions)-421(for)-421(an)-421(interaction)-421(bet)-30(ween)-421(poly)-30(merase)-421(and)-421(RNA)-421(at)-421(the)20(se)]TJ T* [(transcript)-312(lengths)-312(\(26\).)-312(When)-312(the)-312(RNA)-312(reache)20(s)-312(lengths)-312(of)-312(32)-312(and)]TJ T* [(35)-295(nt,)-295(its)-295(5)]TJ /T1_1 1 Tf 0 Tc 4.358 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc 0.613 0 Td [(end)-295(bec)-30(ome)20(s)-295(increasingly)-295(f)-90(lex)-50(ible,)-295(and)-295(more)-295(transient)]TJ -4.971 -1.12 Td [(interactions)-211(w)-40(ith)-211(the)-211(poly)-30(merase)-211(sur)-50(face)-211(are)-211(detected)-211(in)-211(our)-211(FR)-60(ET)]TJ 0 -1.12 TD [(dat)-30(a.)-275(This)-275(finding)-275(is)-275(c)-30(onsistent)-275(w)-40(ith)-275(an)-275(ac)-30(ce)20(ssibilit)-30(y)-275(of)-275(the)-275(RNA)-275(5)]TJ /T1_1 1 Tf 0 Tc 27.173 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc -27.173 -1.12 Td [(end)-271(to)-271(the)-271(5)]TJ /T1_1 1 Tf 0 Tc 5.008 0 Td (\003)Tj /T1_0 1 Tf -0.0149 Tc 0.318 0 Td [(-capping)-271(enz)-30(y)-30(me,)-271(which)-271(modifie)20(s)-271(the)-271(RNA)-271(end)-271(when)]TJ -5.326 -1.12 Td [(a)-318(length)-318(of)]TJ /T1_1 1 Tf 0 Tc 4.862 0 Td (\004)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(30)-318(nt)-318(is)-318(reached)-318(\(27\).)]TJ -4.68 -1.12 Td [(Our)-172(obser)-60(vations)-172(have)-172(further)-172(implications)-172(for)-172(underst)-30(anding)-172(the)]TJ -1 -1.12 Td [(transition)-251(f)-30(rom)-251(transcription)-251(in)-30(itiation)-251(to)-251(elongation.)-251(In)-251(an)-251(in)-30(itially)]TJ T* [(transcribing)-516(c)-30(omplex,)-516(g)-20(row)-40(ing)-516(RNA)-516(sterically)-516(clashe)20(s)-516(w)-40(ith)-516(the)]TJ T* [(in)-30(itiation)-471(factor)-471(TFIIB)-471(finger)-471(domain)-471(\(28\).)-471(Beyond)-471(the)-471(tunnel,)]TJ 0 -1.119 TD [(RNA)-409(also)-409(clashe)20(s)-409(w)-40(ith)-409(the)-409(TFIIB)-409(ribbon)-409(domain)-409(\(29\),)-409(which)-409(is)]TJ T* [(located)-366(on)-366(the)-366(Pol)-366(II)-366(dock)-366(for)-30(ming)-366(c)-30(ont)-30(acts)-366(w)-40(ith)-366(Rpb1)-366(bet)-30(ween)]TJ ET q 194.3999939 0 0 193.3200073 78.4261017 547.8834991 cm /Im0 Do Q 0 0 0 1 k BT /T1_3 1 Tf 0 Tc 7 0 0 7 51.6761 533.8835 Tm [(Fig.)-397(3.)]TJ /T1_4 1 Tf 4.295 0 Td [(Alternating)-397(laser)-397(excitation.)-397(The)-397(graph)-397(shows)-397(an)-397(exemplary)-397(time)]TJ -4.295 -1.286 Td [(trace)-258(of)-258(the)-258(DNA1\226RNA26)-258(sample.)-258(The)-258(excitation)-258(laser)-258(was)-258(alternated)-258(frame)]TJ 0 -1.286 TD [(by)-364(frame)-364(between)-364(633)-364(nm)-364(\(direct\226acceptor)-364(excitation,)]TJ /T1_5 1 Tf 26.109 0 Td (Upper)Tj /T1_4 1 Tf 2.889 0 Td [(\))-364(and)-364(532)-364(nm)]TJ -28.998 -1.286 Td [(\(FRET)-270(excitation,)]TJ /T1_5 1 Tf 7.987 0 Td (Lower)Tj /T1_4 1 Tf 2.889 0 Td [(\))-270(\(34\).)-270(Actual)-270(data)-270(and)-270(a)-270(10-point)-270(sliding)-270(average)-270(are)]TJ -10.876 -1.286 Td [(shown)-278(as)-278(thin)-278(and)-278(thick)-278(lines,)-278(respectively.)]TJ ET q 453.6000061 0 0 224.1600037 79.6761017 105.8834991 cm /Im1 Do Q BT /T1_3 1 Tf 7 0 0 7 51.6761 91.8835 Tm [(Fig.)-231(4.)]TJ /T1_4 1 Tf 3.963 0 Td [(The)-231(position)-231(of)-231(the)-231(nascent)-231(RNA)-231(within)-231(the)-231(elongation)-231(complex)-231(as)-231(determined)-231(from)-231(sp-FRET)-231(measurements.)-231(\()]TJ /T1_5 1 Tf 50.306 0 Td (A)Tj /T1_4 1 Tf 0.722 0 Td [(\))-231(Back)-231(view)-231(of)-231(the)-231(elongation)-231(complex.)]TJ -54.991 -1.286 Td [(The)-216(position)-216(of)-216(the)-216(dye)-216(molecule)-216(attached)-216(to)-216(the)-216(5)]TJ /T1_1 1 Tf 23.115 0 Td (\003)Tj /T1_4 1 Tf 0.549 0 Td [(end)-216(is)-216(illustrated)-216(by)-216(a)-216(single)-216(sphere)-216(with)-216(a)-216(radius)-216(o)]TJ 22.942 0 Td [(f)-216(5)-216(\305)-216(for)-216(RNA17)-216(\(orange\),)-216(RNA20)-216(\(red\),)-216(RNA23)-216(\(blue\),)-216(and)]TJ -46.606 -1.286 Td [(RNA29)-168(\(green\))-168(and)-168(by)-168(a)-168(pair)-168(of)-168(spheres)-168(for)-168(RNA26)-168(\(purple\),)-168(indicating)-168(the)-168(presence)-168(of)-168(two)-168(states.)-168(\()]TJ /T1_5 1 Tf 44.196 0 Td (B)Tj /T1_4 1 Tf 0.611 0 Td [(\))-168(Cut-away)-168(view)-168(revealing)-168(the)-168(paths)-168(of)-168(the)-168(RNA)-168(on)-168(the)-168(interior)]TJ -44.807 -1.286 Td [(and)-278(exterior.)-278(This)-278(\036gure)-278(was)-278(prepared)-278(using)-278(UCSF)-278(Chimera)-278(\(46\).)]TJ /T1_3 1 Tf 6.5 0 0 6.5 51.6761 39.8835 Tm (138)Tj /T1_6 1 Tf 9.75 0 0 6.5 68.5185 39.8835 Tm (\001)Tj /T1_4 1 Tf 6.5 0 0 6.5 76.6831 39.8835 Tm (www.pnas.org)Tj /T1_6 1 Tf 6.833 0 Td (\002)Tj /T1_4 1 Tf 0.5 0 Td (cgi)Tj /T1_6 1 Tf 1.333 0 Td (\002)Tj /T1_4 1 Tf 0.5 0 Td (doi)Tj /T1_6 1 Tf 1.5 0 Td (\002)Tj /T1_4 1 Tf 0.5 0 Td (10.1073)Tj /T1_6 1 Tf 3.614 0 Td (\002)Tj /T1_4 1 Tf 0.5 0 Td (pnas.0703815105)Tj 52.294 0 Td (Andrecka)Tj /T1_5 1 Tf 4.723 0 Td [(et)-278(al)]TJ /T1_4 1 Tf 2.057 0 Td (.)Tj ET q /GS1 gs 19 0 0 815.5 9 9 cm /Im2 Do Q endstream endobj 131 0 obj<> endobj 132 0 obj<>stream 0 0 0 0.2 k 1 i /GS0 gs 573.676 521.384 38.324 -66 re f 0 0 0 1 k BT /T1_0 1 Tf -0.0149 Tc 0 Tw 0 Ts 100 Tz 0 Tr 9 0 0 9 51.6761 734.8835 Tm [(re)20(sidue)20(s)-176(409)-176(and)-176(419)-176(\(28\).)-176(Both)-176(of)-176(the)-176(obser)-60(ved)-176(positions)-176(of)-176(RNA)-60(26)]TJ 0 -1.125 TD [(are)-178(exactly)-178(in)-178(that)-178(area,)-178(in)-178(close)-178(prox)-50(imit)-30(y)-178(to)-178(the)-178(side)-178(chains)-178(of)-178(serine)]TJ T* [(S409)-201(and)-201(argin)-30(ine)-201(R416,)-201(re)20(spectively.)-201(Moreover,)-201(the)-201(sur)-50(face)-201(charge)]TJ T* [(distribution)-214(of)-214(the)-214(poly)-30(merase)-214(in)-214(the)-214(region)-214(of)-214(the)-214(dock)-214(domain)-214(has)]TJ T* [(both)-564(positive)-564(and)-564(negative)-564(patche)20(s,)-564(prov)-40(iding)-564(several)-564(possible)]TJ T* [(electrost)-30(atic)-239(interaction)-239(site)20(s)-239(\()]TJ 0 0 1 rg 12.365 0 Td [(SI)-239(Fig.)-239(10)]TJ 0 0 0 rg 3.919 0 Td [(\).)-239(Thus,)-239(g)-20(row)-40(ing)-239(RNA)-239(may)]TJ -16.284 -1.125 Td [(c)-30(ontribute)-241(to)-241(TFIIB)-241(release)-241(during)-241(the)-241(transition)-241(f)-30(rom)-241(in)-30(itiation)-241(to)]TJ T* [(elongation,)-397(although)-397(strain)-397(that)-397(ac)-30(cumulate)20(s)-397(during)-397(in)-30(itial)-397(DNA)]TJ T* [(scr)-30(unching)-491(and)-491(subsequent)-491(bubble)-491(c)-30(ollapse)-491(is)-491(apparently)-491(more)]TJ T* [(import)-30(ant)-318(\(30,)-318(31\).)-318(Because)-318(RNA)-318(extends)-318(over)-318(the)-318(dock)-318(domain,)]TJ T* [(and)-222(not)-222(v)-40(ia)-222(the)-222(f)-90(lank)-30(ing)-222(g)-20(roove)20(s,)-222(it)-222(can)-222(prevent)-222(reassociation)-222(of)-222(the)]TJ T* [(TFIIB)-229(ribbon)-229(w)-40(ith)-229(Pol)-229(II)-229(during)-229(mRNA)-229(elongation.)-229(We)-229(c)-30(onfir)-30(med)]TJ T* [(this)-281(c)-30(ompetition)-281(in)-281(additional)-281(ex)-30(periments)-281(where)-281(we)-281(added)-281(exce)20(ss)]TJ T* [(amount)-350(of)-350(TFIIB)-350(\(see)]TJ 0 0 1 rg /T1_1 1 Tf 9.777 0 Td [(SI)-350(Te)-30(x)-30(t)]TJ 0 0 0 rg /T1_0 1 Tf 2.981 0 Td [(\).)-350(Even)-350(a)-350(100-fold)-350(exce)20(ss)-350(of)-350(TFIIB)]TJ -12.758 -1.125 Td [(leads)-419(to)-419(the)-419(displacement)-419(of)-419(only)-419(a)-419(small)-419(f)-30(raction)-419(of)-419(the)-419(RNA,)]TJ T* [(whereas)-302(in)-302(the)-302(vast)-302(majorit)-30(y)-302(of)]TJ /T1_2 1 Tf 0 Tc 13.2 0 Td (\004)Tj /T1_0 1 Tf -0.0149 Tc 0.818 0 Td [(80%)-302(of)-302(the)-302(c)-30(omplexe)20(s)-302(the)-302(RNA)]TJ -14.018 -1.125 Td [(remains)-536(associated)-536(w)-40(ith)-536(the)-536(dock)-536(domain)-536(\()]TJ 0 0 1 rg 19.358 0 Td [(SI)-536(Fig.)-536(13)]TJ 0 0 0 rg 4.512 0 Td [(\).)-536(The)20(se)]TJ -23.87 -1.125 Td [(ex)-30(periments)-278(clearly)-278(reveal)-278(the)-278(potential)-278(of)-278(our)-278(method)-278(as)-278(it)-278(allows)]TJ T* [(for)-427(a)-427(direct)-427(obser)-60(vation)-427(of)-427(the)-427(pre)20(sence)-427(of)-427(transcription)-427(factors)]TJ T* [(w)-40(ithin)-251(active)-251(elongation)-251(c)-30(omplexe)20(s)-251(and)-251(their)-251(ef)-30(fect)-251(on)-251(the)-251(location)]TJ T* [(and)-318(c)-30(onfor)-30(mation)-318(of)-318(the)-318(nascent)-318(RNA.)]TJ 1 -1.125 Td [(A)-480(more)-480(general)-480(implication)-480(of)-480(this)-480(work)-480(is)-480(that)-480(the)-480(applied)]TJ -1 -1.125 Td [(single-molecule)-395(trilateration)-395(techn)-30(ique)-395(based)-395(on)-395(multiple)-395(FR)-60(ET)]TJ T* [(measurements)-307(\(20\))-307(prov)-40(ide)20(s)-307(an)-307(ac)-30(curate)-307(tool)-307(for)-307(deter)-30(min)-30(ing)-307(the)]TJ T* [(positions)-281(of)-281(f)-90(lex)-50(ible)-281(domains)-281(in)-281(large)-281(multiprotein)-281(c)-30(omplexe)20(s.)-281(For)]TJ T* [(smaller)-590(proteins,)-590(a)-590(related)-590(single-molecule)-590(approach\227namely,)]TJ T* [(mechan)-30(ical)-396(triangulation\227recently)-396(has)-396(been)-396(e)20(st)-30(ablished)-396(\(32\).)-396(In)]TJ T* [(particular,)-295(we)-295(show)-295(that)-295(single-molecule)-295(f)-90(luore)20(scence)-295(analysis)-295(can)]TJ T* [(reveal)-298(str)-30(uctural)-298(feature)20(s)-298(of)-298(a)-298(euk)-30(ar)-60(yotic)-298(multisubun)-30(it)-298(RNA)-298(poly-)]TJ T* [(merase)-193(that)-193(had)-193(thus)-193(far)-193(e)20(scaped)-193(detection)-193(by)-193(x-ray)-193(cr)-60(yst)-30(allog)-20(raphic)]TJ T* [(means)-443(because)-443(of)-443(inherent)-443(f)-90(lex)-50(ibilit)-30(y)-443(and/or)-443(because)-443(of)-444(limited)]TJ T* [(dif)-30(f)-30(raction)-318(dat)-30(a)-318(re)20(solution.)]TJ 1 -1.125 Td [(The)-291(power)-291(of)-291(single-molecule)-291(f)-90(luore)20(scence)-291(techn)-30(ique)20(s)-291(for)-291(tran-)]TJ -1 -1.125 Td [(scription)-356(studie)20(s)-356(had)-356(so)-356(far)-356(been)-356(demonstrated)-356(only)-356(for)-356(bacterial)]TJ T* [(RNA)-279(poly)-30(merase)-279(\(31,)-279(33\22635\).)-279(On)-279(the)-279(other)-279(hand,)-279(single-molecule)]TJ T* [(force)-373(spectrosc)-30(opy)-373(has)-373(been)-373(used)-373(to)-373(study)-373(the)-373(dynamics)-373(of)-373(tran-)]TJ T* [(scription)-379(by)-379(bacterial)-379(and)-379(v)-40(iral)-379(RNA)-379(poly)-30(merase)20(s.)-379(The)20(se)-379(ex)-30(peri-)]TJ T* [(ments)-292(c)-30(ould)-292(elucidate)-292(k)-30(inetic)-292(aspects)-292(\(36\),)-292(transcriptional)-292(pausing)]TJ T* [(and)-295(backtrack)-30(ing)-295(\(37,)-295(38\),)-295(and)-295(det)-30(ails)-295(of)-295(the)-295(translocation)-295(mecha-)]TJ T* [(n)-30(ism)-278(\(39)-90(\226)-90(42\).)-278(The)-278(ex)-30(periments)-278(de)20(scribed)-278(in)-278(this)-278(article,)-278(in)-278(c)-30(ombi-)]TJ T* [(nation)-362(w)-40(ith)-362(recently)-362(published)-362(mechan)-30(ical)-362(studie)20(s)-362(of)-362(Pol)-362(II)-362(tran-)]TJ T* [(scription)-367(elongation)-367(\(43\),)-367(prov)-40(ide)-367(a)-367(foundation)-367(for)-367(future)-367(single-)]TJ T* [(molecule)-295(ex)-30(periments)-295(on)-295(euk)-30(ar)-60(yotic)-295(RNA)-295(poly)-30(merase)20(s.)-295(The)-295(time)]TJ T* [(re)20(solution)-326(of)-326(single-molecule)-326(f)-90(luore)20(scence)-326(ex)-30(periments)-326(allows)-326(for)]TJ T* [(an)-388(inve)20(stigation)-388(of)-388(str)-30(uctural)-388(change)20(s)-388(in)-388(real)-388(time,)-388(re)20(sulting)-388(in)-388(a)]TJ 0 -1.124 TD [(dynamic)-376(v)-40(iew)-376(of)-376(c)-30(omplex)-376(biological)-376(proce)20(sse)20(s.)-376(Such)-376(ex)-30(periments)]TJ T* [(c)-30(ould)-256(prov)-40(ide)-256(valuable)-256(insights)-256(into)-256(unclear)-256(mechan)-30(istic)-256(aspects)-256(of)]TJ T* [(euk)-30(ar)-60(yotic)-318(transcription)-318(and)-318(eventually)-318(also)-318(its)-318(regulation.)]TJ /T1_3 1 Tf -0.0196 Tc 0 -2.013 TD [(Materials)-313(and)-313(Methods)]TJ 0 Tc 7 0 0 7 51.6761 229.8153 Tm [(Single-Cysteine)-182(Rpb4/7)-182(Variants.)]TJ /T1_4 1 Tf -0.0195 Tc 14.515 0 Td [(The)-182(yeast)-182(heterodimer)-182(of)-182(Rpb4/7)-182(was)-182(expressed)]TJ -14.515 -1.302 Td [(in)-238(a)-238(bacterial)-238(expression)-238(system)-238(and)-238(added)-238(to)-238(the)-238(10-subunit)-238(Pol)-238(II\226)-120(core\226nucleic)]TJ 0 -1.302 TD [(acid)-291(scaffold)-291(complex)]TJ /T1_5 1 Tf 9.948 0 Td [(in)-291(vitro)]TJ /T1_4 1 Tf 3.502 0 Td [(to)-291(obtain)-291(the)-291(complete)-291(polymerase)-291(II)-291(elongation)]TJ -13.45 -1.302 Td [(complex)-279(\(3\).)-279(The)-279(comparatively)-279(small)-279(Rpb4/7)-279(heterodimer)-279(with)-279(only)-279(\036ve)-279(native)]TJ 0 -1.302 TD [(cysteines)-184(is)-184(well)-184(suited)-184(for)-184(mutagenesis,)-184(and)-184(therefore)-184(the)-184(attachment)-184(of)-184(a)-184(single)]TJ 0 -1.302 TD [(dye)-143(molecule)-143(within)-143(an)-143(elongation)-143(complex)-143(at)-143(a)-143(well)-143(de\036ned)-143(and)-143(known)-143(location)]TJ 0 -1.302 TD [(becomes)-171(feasible.)-171(To)-171(this)-171(end,)-171(we)-171(performed)-171(site-directed)-171(mutagenesis)-171(replacing)]TJ T* [(native)-441(cysteines)-441(with)-441(serines)-441(to)-441(produce)-441(two)-441(mutants)-441(with)-441(only)-441(one)-441(single-)]TJ T* [(cysteine)-392(residue)-392(at)-392(locations)-392(C94)-392(and)-392(C150,)-392(respectively.)-392(The)-392(constructs)-392(were)]TJ T* [(expressed)-259(and)-259(puri\036ed)-259(as)-258(described)-259(previously)-258(\(3\).)]TJ 1.286 -1.302 Td [(Dye)-326(labeling)-326(of)-326(the)-326(single-cysteine)-326(mutants)-326(was)-326(conducted)-326(with)-326(an)-326(8)-326(to)-326(10)]TJ -1.286 -1.302 Td [(times)-313(molar)-313(excess)-313(of)-313(Alexa)-313(647-C2-Maleimide)-313(\(Molecular)-313(Probes\))-313(in)-313(DTT)-313(free-)]TJ T* [(assembly)-277(buffer)-277(\(50)-277(mM)-277(Hepes,)-277(pH)-277(7.5/40)-277(mM)-277(\(NH)]TJ 0 Tc 5 0 0 5 210.6017 119.4517 Tm (4)Tj 7 0 0 7 213.2855 120.4517 Tm (\))Tj 5 0 0 5 215.4802 119.4517 Tm (2)Tj -0.0195 Tc 7 0 0 7 218.1639 120.4517 Tm (SO)Tj 0 Tc 5 0 0 5 226.8373 119.4517 Tm (4)Tj -0.0195 Tc 7 0 0 7 229.5211 120.4517 Tm (/10)Tj /T1_6 1 Tf 0 Tc 7 0 1.0881 7 240.7825 120.4517 Tm (\001)Tj /T1_4 1 Tf -0.0195 Tc 7 0 0 7 245.3152 120.4517 Tm [(M)-277(ZnCl)]TJ 0 Tc 5 0 0 5 267.5741 119.4517 Tm (2)Tj -0.0195 Tc 7 0 0 7 270.2579 120.4517 Tm [(/5%)-277(glyc-)]TJ -31.226 -1.302 Td [(erol\))-314(at)-314(37\260C)-314(for)-314(1)-314(h.)-314(Labeled)-314(protein)-314(was)-314(puri\036ed)-314(by)-314(using)-314(G-50)-314(spin-columns)]TJ 0 -1.302 TD [(\(Amersham)-412(Biosciences\))-412(with)-412(assembly)-412(buffer)-412(\(50)-412(mM)-412(Hepes,)-412(pH)-412(7.5/40)-412(mM)]TJ T* (\(NH)Tj 0 Tc 5 0 0 5 63.7062 92.1108 Tm (4)Tj 7 0 0 7 66.39 93.1108 Tm (\))Tj 5 0 0 5 68.5847 92.1108 Tm (2)Tj -0.0195 Tc 7 0 0 7 71.2684 93.1108 Tm (SO)Tj 0 Tc 5 0 0 5 79.9419 92.1108 Tm (4)Tj -0.0195 Tc 7 0 0 7 82.6256 93.1108 Tm (/10)Tj /T1_6 1 Tf 0 Tc 7 0 1.0881 7 93.7563 93.1108 Tm (\001)Tj /T1_4 1 Tf -0.0195 Tc 7 0 0 7 98.2891 93.1108 Tm [(M)-259(ZnCl)]TJ 0 Tc 5 0 0 5 120.4173 92.1108 Tm (2)Tj -0.0195 Tc 7 0 0 7 123.101 93.1108 Tm [(/5%)-259(glycerol/10)-259(mM)-259(DTT\).)]TJ /T1_3 1 Tf 0 Tc -10.204 -2.731 Td [(Preparation)-200(of)-200(Pol)-200(II)-200(Elongation)-200(Complexes.)]TJ /T1_4 1 Tf -0.0195 Tc 19.411 0 Td [(Synthetic)-200(nucleic)-200(acid)-200(scaffolds)-200(were)]TJ -19.411 -1.302 Td [(constructed)-450(from)-450(RNA)-450(and)-450(DNA)-450(oligomers)-450(\(Biomers\))-450(by)-450(using)-450(partially)-450(mis-)]TJ 37.5 95.857 Td [(matched)-150(DNA)-150(template)-150(and)-150(nontemplate)-150(strands.)-150(The)-150(sequences)-150(of)-150(the)-150(template)]TJ 0 -1.292 TD [(and)-232(nontemplate)-232(strands)-232(as)-232(well)-232(as)-232(the)-232(RNA)-232(sequence)-232(\(35-nt)-232(RNA\))-232(are)-232(shown)-232(in)]TJ T* [(Fig.)-400(1)]TJ /T1_5 1 Tf 0 Tc 2.645 0 Td (A)Tj /T1_4 1 Tf -0.0195 Tc 0.703 0 Td [(.)-400(One)-400(should)-400(note)-400(that)-400(no)-400(backtracking)-400(has)-400(been)-400(observed)-400(for)-400(these)]TJ -3.348 -1.292 Td [(elongation)-396(complexes)-396(in)-396(multiple)-396(crystallographic)-396(and)-396(biochemical)-396(studies)-396(in)]TJ T* [(contrast)-448(to)-448(studies)-448(with)-448(native)-448(promoter)-448(sequences,)-448(presumably)-448(because)-448(of)]TJ T* [(energetic)-258(penalties)-258(caused)-259(by)-259(the)-259(mismatches)-258(of)-258(the)-259(oligonucleotides.)]TJ 1.286 -1.292 Td [(Seven)-292(different)-292(RNA)-292(molecules)-292(with)-292(lengths)-292(from)-292(17)-292(to)-292(35)-292(nt)-292(and)-292(sequence)]TJ 0 Tc -1.286 -1.292 Td (5)Tj /T1_2 1 Tf 0.545 0 Td (\003)Tj /T1_4 1 Tf -0.0109 Tc 0.322 0 Td (-AACAUUACACGAAUAUAUAUGCAUAAAGACCAGGC-3)Tj /T1_2 1 Tf 0 Tc 25.299 0 Td (\003)Tj /T1_4 1 Tf -0.0109 Tc 0.811 0 Td [(were)-489(investigated)]TJ ET 0 0 0 RG 0.301 w 10 M 0 j 0 J []0 d 322.494 671.415 m 327.469 671.415 l 336.642 671.415 m 341.616 671.415 l 351.565 671.415 m 356.54 671.415 l 365.713 671.415 m 369.912 671.415 l 379.861 671.415 m 384.835 671.415 l 394.784 671.415 m 399.759 671.415 l 409.707 671.415 m 414.682 671.415 l S BT /T1_4 1 Tf -0.0195 Tc 7 0 0 7 314.1761 663.5199 Tm [(\(the)-167(underlined)-167(letters)-167(correspond)-167(to)-167(the)-167(seven)-167(different)-167(5)]TJ /T1_2 1 Tf 0 Tc 25.406 0 Td (\003)Tj /T1_4 1 Tf -0.0195 Tc 0.481 0 Td [(ends)-167(of)-167(a)-167(17-nt,)-167(20-nt,)]TJ -25.886 -1.292 Td [(23-nt,)-275(26-nt,)-275(29-nt,)-275(32-nt,)-275(and)-275(35-nt)-275(strand\).)-275(Each)-275(5)]TJ /T1_2 1 Tf 0 Tc 22.719 0 Td (\003)Tj /T1_4 1 Tf -0.0195 Tc 0.589 0 Td [(end)-275(had)-275(a)-275(6-TMR)-275(molecule)]TJ -23.308 -1.292 Td [(attached)-155(via)-155(a)-155(C6-amino)-155(linker.)-155(The)-155(nontemplate)-155(DNA)-155(strand)-155(had)-155(Biotin)-155(attached)]TJ 0 -1.292 TD [(at)-285(the)-285(5)]TJ /T1_2 1 Tf 0 Tc 3.511 0 Td (\003)Tj /T1_4 1 Tf -0.0195 Tc 0.599 0 Td [(end)-285(via)-285(a)-285(C6-amino)-285(linker.)-285(The)-285(biotin)-285(was)-285(used)-285(in)-285(the)-285(single-molecule)]TJ -4.11 -1.292 Td [(experiments)-278(for)-278(immobilization)-278(of)-278(the)-278(elongation)-278(complex.)-278(The)-278(template)-278(DNA)]TJ 0 -1.292 TD [(strand)-217(contained)-217(an)-217(internal)-217(C6-amino)-217(linker)-217(attached)-217(to)-217(a)-217(thymidine)-217(residue)-217(at)]TJ T* [(either)-221(position)]TJ /T1_2 1 Tf 0 Tc 6.726 0 Td (\002)Tj /T1_4 1 Tf -0.0195 Tc 0.814 0 Td [(10)-221(\(DNA1\))-221(or)-221(position)]TJ /T1_2 1 Tf 0 Tc 9.811 0 Td (\001)Tj /T1_4 1 Tf -0.0195 Tc 0.814 0 Td [(3)-221(\(DNA2\).)-221(Alexa647)-221(\(Molecular)-221(Probes\))]TJ -18.164 -1.292 Td [(then)-213(was)-213(covalently)-213(attached)-213(to)-213(these)-213(positions)-213(by)-213(using)-213(an)]TJ /T1_5 1 Tf 0 Tc 26.387 0 Td (N)Tj /T1_4 1 Tf -0.0195 Tc 0.703 0 Td (-hydroxysuccinimi-)Tj -27.09 -1.292 Td [(dylester)-283(derivative)-283(following)-283(the)-283(manufacturer\222s)-283(suggestions.)-283(Labeled)-283(DNA)-283(oli-)]TJ 0 -1.292 TD [(gomers)-259(were)-259(puri\036ed)-259(by)-259(HPLC.)]TJ 1.286 -1.292 Td [(In)-169(addition)-169(to)-169(the)-169(FRET)-169(measurements)-169(of)-169(complexes)-169(with)-169(labeled)-169(RNA,)-169(control)]TJ -1.286 -1.292 Td [(experiments)-165(were)-165(performed)-165(to)-165(measure)-165(the)-165(distance)-165(between)-165(donor)-165(dye)-165(on)-165(the)]TJ T* [(DNA)-377(and)-377(acceptor)-377(on)-377(Rpb7.)-377(To)-377(this)-377(end,)-377(the)-377(template)-377(strand)-377(was)-377(labeled)-377(at)]TJ T* (position)Tj /T1_2 1 Tf 0 Tc 3.798 0 Td (\002)Tj /T1_4 1 Tf -0.0195 Tc 0.814 0 Td [(10)-176(\(DNA1\))-176(or)-176(at)-176(position)]TJ /T1_2 1 Tf 0 Tc 10.712 0 Td (\001)Tj /T1_4 1 Tf -0.0195 Tc 0.814 0 Td [(3)-176(\(DNA2\))-176(with)-176(5-TMR)-176(isothiocyanate)-176(\(TRITC;)]TJ -16.137 -1.292 Td [(Molecular)-310(Probes\))-310(followed)-310(by)-310(HPLC)-310(puri\036cation.)-310(Furthermore,)-310(the)-310(Alexa-647-)]TJ 0 -1.292 TD [(labeled)-326(Rpb7\226C150,)-326(biotin-labeled)-326(nontemplate)-326(DNA,)-326(and)-326(a)-326(nonlabeled)-326(17-nt)]TJ T* [(RNA)-200(primer)-200(were)-200(used)-200(in)-200(the)-200(preparation)-200(of)-200(elongation)-200(complexes)-200(for)-200(these)-200(test)]TJ T* (experiments.)Tj 1.286 -1.292 Td [(DNA\226RNA)-237(scaffolds)-237(were)-237(annealed)-237(by)-237(using)-237(equimolar)-237(amounts)-237(of)-237(template)]TJ -1.286 -1.292 Td [(DNA,)-215(nontemplate)-215(DNA,)-215(and)-215(RNA)-215(in)-215(TE)-215(buffer)-215(\(10)-215(mM)-215(Tris)]TJ /T1_7 1 Tf 0 Tc 25.912 0 Td (\002)Tj /T1_4 1 Tf -0.0195 Tc 0.203 0 Td [(HCl/1)-215(mM)-215(EDTA\))-215(at)-215(a)]TJ -26.115 -1.292 Td [(\036nal)-204(concentration)-204(of)-204(1)]TJ /T1_6 1 Tf 0 Tc 7 0 1.0881 7 388.342 482.6108 Tm (\001)Tj /T1_4 1 Tf -0.0195 Tc 7 0 0 7 392.8748 482.6108 Tm [(M.)-204(The)-204(mixture)-204(was)-204(heated)-204(to)-204(95\260C)-204(for)-204(5)-204(min,)-204(followed)]TJ -11.243 -1.292 Td [(by)-209(fast)-209(cooling)-209(to)-209(55\260C)-209(and)-209(then)-209(slow)-209(cooling)-209(to)-209(4\260C)-209(in)-209(1)-209(h.)-209(The)-209(Pol)-209(II\226DNA\226RNA\226)]TJ 0 -1.292 TD [(Rpb4/7)-253(complex)-253(was)-253(assembled)-253(as)-253(described)-253(in)-253(ref.)-253(3.)-253(The)-253(complex)-253(was)-253(puri\036ed)]TJ 0 -1.292 TD [(using)-144(Microcon)-144(YM100)-144(centrifugal)-144(\036lter)-144(units)-144(\(Millipore\))-144(against)-144(assembly)-144(buffer.)]TJ /T1_3 1 Tf 0 Tc 0 -2.721 TD [(Preparation)-588(of)-588(Sample)-588(Chamber)-588(for)-588(sp-FRET)-588(Measurements.)]TJ /T1_4 1 Tf -0.0195 Tc 29.27 0 Td [(Quartz)-588(slides)]TJ -29.27 -1.292 Td [(\(Finkenbeiner\))-373(and)-373(coverslips)-373(\(Marienfeld\))-373(were)-373(thoroughly)-373(cleaned)-373(with)-373(the)]TJ 0 -1.292 TD [(detergent)-211(Hellmanex)-211(II)-211(\(Hellma\))-211(and)-211(dried)-211(in)-211(nitrogen)-211(\037ow.)-211(Remaining)-211(particles)]TJ 0 -1.292 TD [(were)-402(oxidized)-402(with)-402(a)-402(butane-gas)-402(torch.)-402(Afterward)-402(the)-402(quartz)-402(slides)-402(were)-402(si-)]TJ T* [(lanized)-245(by)-245(using)-245(2%)-245(\(vol/vol\))-245(\(3-aminopropyl\)-triethoxysilane)-245(\(Sigma-Aldrich\))-245(in)]TJ T* [(acetone,)-328(followed)-328(by)-328(desiccation,)-328(rinsing)-328(with)-328(water,)-328(and)-328(drying.)-328(Hereafter,)-328(a)]TJ T* [(solution)-360(of)-360(mPEG-succinimidyl)-360(propionate)-360([15%)-360(\(wt/vol\),)]TJ /T1_5 1 Tf 0 Tc 26.13 0 Td (M)Tj /T1_4 1 Tf 5 0 0 5 503.5571 381.1562 Tm (r)Tj -0.0195 Tc 7 0 0 7 507.9228 382.1563 Tm [(5,000)-360(Da,)-360(Nektar)]TJ -27.678 -1.292 Td [(Therapeutics])-254(and)-254(biotinylated)-254(PEG\226)]TJ /T1_5 1 Tf 0 Tc 16.198 0 Td (N)Tj /T1_4 1 Tf -0.0195 Tc 0.703 0 Td [(-hydroxysuccinimide)-254(ester)-254(\(PEG-NHS\))-254([1%)]TJ -16.901 -1.292 Td (\(wt/vol\),)Tj /T1_5 1 Tf 0 Tc 3.846 0 Td (M)Tj /T1_4 1 Tf 5 0 0 5 347.5703 363.0653 Tm (r)Tj -0.0195 Tc 7 0 0 7 350.7371 364.0653 Tm [(3,400)-188(Da,)-188(Nektar)-188(Therapeutics])-188(in)-188(carbonate\226)-120(bicarbonate)-188(buffer)-188(\(pH)]TJ -5.223 -1.292 Td [(9.4\))-167(was)-167(applied.)-167(Subsequently)-167(the)-167(PEGylated)-167(slides)-167(were)-167(cleaned)-167(with)-167(water)-167(and)]TJ 0 -1.292 TD (dried.)Tj 1.286 -1.292 Td [(The)-151(slides)-151(then)-151(were)-151(assembled)-151(together)-151(with)-151(a)-151(precut)-151(sealing)-151(\036lm)-151(\(Nesco\036lm\))]TJ -1.286 -1.292 Td [(and)-256(a)-256(cleaned)-256(coverslip)-256(and)-256(heated)-256(to)-256(150\260C)-256(for)-256(1)-256(min)-256(to)-256(allow)-256(for)-256(the)-256(thermo-)]TJ 0 -1.292 TD [(plastic)-361(\036lm)-361(to)-361(seal)-361(a)-361(channel)-361(and)-361(produce)-361(a)-361(micro\037uidic)-361(chamber.)-361(Two)-361(holes)]TJ T* [(drilled)-368(into)-368(the)-368(quartz)-368(slides)-368(were)-368(used)-368(to)-368(insert)-368(and)-368(remove)-368(\037uids)-368(from)-368(the)]TJ T* (chamber.)Tj 1.286 -1.292 Td [(Pol)-304(II)-304(elongation)-304(complexes)-304(were)-304(attached)-304(to)-304(the)-304(biotin\226PEG)-304(surface)-304(of)-304(the)]TJ -1.286 -1.291 Td [(micro\037uidic)-265(chamber)-265(via)-265(neutravidin/biotin)-265(attachment.)-265(To)-265(this)-265(end,)-265(the)-265(cham-)]TJ 0 -1.291 TD [(ber)-279(was)-279(incubated)-279(with)-279(0.5)-279(mg/ml)-279(neutravidin)-279(\(Molecular)-279(Probes\))-279(in)-279(PBS.)-279(After)]TJ T* [(exchanging)-214(the)-214(buffer)-214(with)-214(assembly)-214(buffer,)-214(the)-214(preassembled)-214(elongation)-214(com-)]TJ T* [(plexes)-271(were)-271(loaded.)-271(Afterward)-271(unbound)-271(complexes)-271(were)-271(removed)-271(by)-271(washing)]TJ T* [(extensively)-259(with)-259(assembly)-259(buffer.)]TJ /T1_3 1 Tf 0 Tc 0 -2.719 TD [(Experimental)-451(Setup)-451(for)-451(sp-FRET.)]TJ /T1_4 1 Tf -0.0195 Tc 15.363 0 Td [(sp-FRET)-451(experiments)-451(were)-451(performed)-451(on)-451(a)]TJ -15.363 -1.291 Td [(homebuilt)-144(prism-based)-144(total)-144(internal)-144(re\037ection)-144(\037uorescence)-144(microscope)-144(\(TIRFM\).)]TJ 0 -1.291 TD [(In)-451(this)-451(apparatus,)-451(the)-451(sample)-451(is)-451(excited)-451(by)-451(using)-451(both)-451(a)-451(frequency-doubled)]TJ T* [(Nd-YAG)-261(laser)-261(\(Crystalaser\))-261(at)-261(532)-261(nm)-261(for)-261(the)-261(excitation)-261(of)-261(donor)-261(molecules)-261(and)]TJ T* [(FRET)-195(pairs)-195(and)-195(a)-195(He-Ne)-195(laser)-195(\(Uniphase\))-195(at)-195(633)-195(nm)-195(for)-195(the)-195(direct)-195(excitation)-195(of)-195(the)]TJ T* [(acceptor.)-150(The)-150(light)-150(from)-150(the)-150(two)-150(lasers)-150(is)-150(combined)-150(spatially)-150(with)-150(a)-150(dichroic)-150(mirror)]TJ T* [(\(Chroma)-284(Z532RDC\).)-284(During)-284(the)-284(measurement,)-284(the)-284(excitation)-284(can)-284(be)-284(alternated)]TJ T* [(between)-234(the)-234(two)-234(laser)-234(sources)-234(by)-234(using)-234(two)-234(software-controlled)-234(shutters)-234(\(Unib-)]TJ T* [(litz)-211(LS6\).)-211(Alternating)-211(between)-211(FRET)-211(and)-211(direct-acceptor)-211(excitation)-211(allows)-211(for)-211(the)]TJ T* [(discrimination)-140(of)-140(dynamics)-140(in)-140(FRET)-140(ef\036ciencies)-140(caused)-140(by)-140(conformational)-140(changes)]TJ T* [(against)-171(changes)-171(in)-171(FRET)-171(ef\036ciencies)-171(caused)-171(by)-171(\037uctuations)-171(in)-171(acceptor)-171(brightness)]TJ 0 -1.291 TD [(\(18\))-277(\(see)]TJ 0 0 1 rg /T1_5 1 Tf 4.01 0 Td [(SI)-277(Text)]TJ 0 0 0 rg /T1_4 1 Tf 2.939 0 Td (\).)Tj -5.663 -1.291 Td [(The)-272(excitation)-272(light)-272(is)-272(coupled)-272(into)-272(the)-272(sample)-272(chamber)-272(by)-272(means)-272(of)-272(a)-272(small)]TJ -1.286 -1.291 Td [(prism.)-280(For)-280(total)-280(internal)-280(re\037ection)-280(\(TIR\),)-280(the)-280(incident)-280(angle)-280(is)-280(adjusted)-280(to)]TJ /T1_2 1 Tf 0 Tc 32.824 0 Td (\004)Tj /T1_4 1 Tf -0.0195 Tc 0.814 0 Td (74\260.)Tj -33.637 -1.291 Td [(The)-342(interior)-342(surface)-342(of)-342(the)-342(\037ow)-342(chamber)-342(sits)-342(at)-342(the)-342(focus)-342(of)-342(a)-342(lens)-342(with)-342(focal)]TJ T* (length)Tj /T1_5 1 Tf 0 Tc 3.249 0 Td (f)Tj /T1_2 1 Tf 0.679 0 Td (\006)Tj /T1_4 1 Tf -0.0195 Tc 1.123 0 Td [(75)-310(mm,)-310(yielding)-310(an)-310(excitation)-310(area)-310(of)]TJ /T1_2 1 Tf 0 Tc 17.291 0 Td (\004)Tj /T1_4 1 Tf -0.0195 Tc 0.814 0 Td (70)Tj /T1_2 1 Tf 0 Tc 1.383 0 Td (\b)Tj /T1_4 1 Tf -0.0195 Tc 1.123 0 Td (35)Tj /T1_6 1 Tf 0 Tc 7 0 1.0881 7 503.4868 91.9858 Tm (\001)Tj /T1_4 1 Tf 7 0 0 7 508.0195 91.9858 Tm (m)Tj 5 0 0 5 514.1063 94.2358 Tm (2)Tj -0.0195 Tc 7 0 0 7 518.9576 91.9858 Tm [(with)-310(a)-310(power)]TJ -29.254 -1.291 Td [(density)-243(of)-243(1.5)]TJ /T1_6 1 Tf 0 Tc 7 0 1.0881 7 356.0176 82.9517 Tm (\001)Tj /T1_4 1 Tf -0.0195 Tc 7 0 0 7 360.5503 82.9517 Tm (W/)Tj /T1_6 1 Tf 0 Tc 7 0 1.0881 7 369.2237 82.9517 Tm (\001)Tj /T1_4 1 Tf 7 0 0 7 373.7564 82.9517 Tm (m)Tj 5 0 0 5 379.8432 85.2017 Tm (2)Tj -0.0195 Tc 7 0 0 7 382.5269 82.9517 Tm [(.)-243(Fluorescence)-243(intensity)-243(is)-243(collected)-243(via)-243(a)-243(water-immersion)]TJ -9.764 -1.291 Td [(objective)-324(\(Plan)-324(Apo)]TJ /T1_2 1 Tf 0 Tc 9.142 0 Td (\b)Tj /T1_4 1 Tf -0.0195 Tc 0.813 0 Td [(60,)-324(N.A.)-324(1.2;)-324(Nikon\))-324(and)-324(directed)-324(to)-324(an)-324(intensi\036ed)-324(back-)]TJ -9.955 -1.291 Td [(illuminated)-282(electron-multiplying)-282(charge-coupled)-282(device)-282(\(EMCCD\))-282(camera)-282(\(iXon)]TJ 0 Tc 6.5 0 0 6.5 51.6761 39.8835 Tm (Andrecka)Tj /T1_5 1 Tf 4.75 0 Td [(et)-305(al)]TJ /T1_4 1 Tf 2.084 0 Td (.)Tj 45.416 0 Td (PNAS)Tj /T1_8 1 Tf 9.75 0 0 6.5 413.5518 39.8835 Tm (\001)Tj /T1_9 1 Tf 6.5 0 0 6.5 421.7163 39.8835 Tm [(January)-305(8,)-305(2008)]TJ /T1_8 1 Tf 9.75 0 0 6.5 475.4029 39.8835 Tm (\001)Tj /T1_4 1 Tf 6.5 0 0 6.5 483.5675 39.8835 Tm [(vol.)-305(105)]TJ /T1_8 1 Tf 9.75 0 0 6.5 513.2288 39.8835 Tm (\001)Tj /T1_4 1 Tf 6.5 0 0 6.5 521.3933 39.8835 Tm [(no.)-305(1)]TJ /T1_8 1 Tf 9.75 0 0 6.5 542.7408 39.8835 Tm (\001)Tj /T1_9 1 Tf 6.5 0 0 6.5 550.9054 39.8835 Tm (139)Tj /T1_3 1 Tf 0 6.5 -6.5 0 585.1761 471.2386 Tm (BIOPHYSICS)Tj ET q /GS1 gs 19 0 0 815.5 9 9 cm /Im0 Do Q endstream endobj 133 0 obj<> endobj 134 0 obj<>stream 0 0 0 1 k 1 i /GS0 gs BT /T1_0 1 Tf -0.0195 Tc 0 Tw 0 Ts 100 Tz 0 Tr 7 0 0 7 51.6761 735.8835 Tm [(DV887DCS-BV;)-301(Andor\).)-301(To)-301(detect)-301(both)-301(donor)-301(and)-301(acceptor)-301(\037uorescence)-301(simul-)]TJ 0 -1.289 TD [(taneously)-177(with)-177(the)-177(same)-177(camera,)-177(one)-177(half)-177(of)-177(the)-177(CCD)-177(chip)-177(is)-177(physically)-177(blocked)-177(by)]TJ T* [(using)-239(a)-239(slit)-239(in)-239(the)-239(image)-239(plane)-239(of)-239(the)-239(detection)-239(path.)-239(Two)-239(channels)-239(of)-239(detection)]TJ 0 -1.287 TD [(then)-234(are)-234(introduced)-234(by)-234(splitting)-234(and)-234(spatially)-234(offsetting)-234(the)-234(donor)-234(and)-234(acceptor)]TJ 0 -1.287 TD [(\037uorescence)-233(via)-233(a)-233(dichroic)-233(beam)-233(splitter)-233(\(Chroma)-233(645DCXR\).)-233(Emission)-233(\036lters)-233(are)]TJ T* [(placed)-232(in)-232(the)-232(two)-232(detection)-232(paths)-232(centered)-232(at)-232(580)-232(nm)-232(\(Omega)-232(Optical)-232(3RD550)-120(\226)]TJ T* [(610\))-256(and)-256(710)-256(nm)-256(\(Chroma)-256(HQ710/100M\))-256(to)-256(isolate)-256(the)-256(\037uorescence)-256(light)-256(of)-256(the)]TJ T* [(donor)-356(and)-356(acceptor)-356(molecules,)-356(respectively.)-356(All)-356(measurements)-356(were)-356(recorded)]TJ T* [(with)-259(an)-259(exposure)-259(time)-258(of)-258(100)-259(ms)-258(per)-258(frame)-259(for)-259(a)-259(30-s)-258(duration.)]TJ /T1_1 1 Tf 0 Tc 0 -2.716 TD [(Data)-478(Analysis.)]TJ /T1_0 1 Tf -0.0195 Tc 7.013 0 Td [(The)-478(acquired)-478(data)-478(were)-478(analyzed)-478(by)-478(using)-478(custom)-478(software)]TJ -7.013 -1.287 Td [(written)-203(in)-203(MATLAB.)-203(We)-203(used)-203(a)-203(fully)-203(automated)-203(routine)-203(to)-203(\036nd)-203(FRET)-203(pairs)-203(and)-203(to)]TJ 0 -1.287 TD [(calculate)-152(and)-152(subtract)-152(the)-152(local)-152(background.)-152(For)-152(the)-152(calculation)-152(of)-152(FRET)-152(ef\036ciency)]TJ T* [(of)-258(the)-259(individual)-258(FRET)-259(pairs,)-258(we)-259(used)-259(the)-258(following)-259(formula)-259(\(44\):)]TJ /T1_2 1 Tf 0 Tc 9 0 0 9 147.2727 592.5085 Tm (E)Tj /T1_3 1 Tf 0.918 0 Td (\002)Tj /T1_2 1 Tf 1.075 0.812 Td (I)Tj 6.3977 0 0 5.7614 168.6693 598.2216 Tm (A)Tj /T1_3 1 Tf 9 0 0 9 175.2087 599.8125 Tm (\003)Tj 9 0 1.3977 9 184.8815 599.8125 Tm (\004)Tj /T1_4 1 Tf 9 0 0 9 190.5592 599.8125 Tm (\002)Tj /T1_2 1 Tf 0.242 0 Td (I)Tj 6.3977 0 0 5.7614 196.2 598.2216 Tm (D)Tj ET 0 0 0 1 K 0.25 w 10 M 0 j 0 J []0 d 165.205 595.759 m 201.148 595.759 l S BT /T1_2 1 Tf 9 0 0 9 165.4517 586.8352 Tm (I)Tj 6.3977 0 0 5.7614 168.9155 585.2443 Tm (A)Tj /T1_3 1 Tf 9 0 0 9 175.4549 586.8352 Tm (\005)Tj 9 0 1.3977 9 185.1278 586.8352 Tm (\006)Tj /T1_4 1 Tf 9 0 0 9 190.3105 586.8352 Tm (\002)Tj /T1_2 1 Tf 0.242 0 Td (I)Tj 6.3977 0 0 5.7614 195.9513 585.2443 Tm (D)Tj /T1_5 1 Tf 9 0 0 9 201.1565 592.5085 Tm (,)Tj /T1_6 1 Tf 0.0199 Tc 9.471 0 Td ([1])Tj /T1_0 1 Tf 0 Tc 7 0 0 7 51.6761 559.7812 Tm (where)Tj /T1_3 1 Tf 9 0 1.3977 9 150.8295 534.5654 Tm (\006)Tj 9 0 0 9 158.5947 534.5654 Tm (\002)Tj /T1_2 1 Tf 1.186 0.812 Td (I)Tj 6.3977 0 0 5.7614 173.3166 540.2784 Tm (A)Tj /T1_3 1 Tf 9 0 0 9 179.9384 541.8693 Tm (\003)Tj /T1_2 1 Tf 1.14 0 Td (I)Tj /T1_7 1 Tf 0.45 0 Td (\003)Tj /T1_2 1 Tf 6.3977 0 0 5.7614 198.0066 540.2784 Tm (A)Tj ET 168.852 537.815 m 203.037 537.815 l S BT /T1_2 1 Tf 9 0 0 9 168.8523 528.8921 Tm (I)Tj /T1_7 1 Tf 0.45 0 Td (\003)Tj /T1_2 1 Tf 6.3977 0 0 5.7614 176.6624 527.3011 Tm (D)Tj /T1_3 1 Tf 9 0 0 9 183.7001 528.8921 Tm (\003)Tj /T1_2 1 Tf 1.14 0 Td (I)Tj 6.3977 0 0 5.7614 198.0072 527.3011 Tm (D)Tj /T1_0 1 Tf 7 0 0 7 51.6761 511.8494 Tm (and)Tj /T1_3 1 Tf 9 0 1.3977 9 158.7443 486.6335 Tm (\004)Tj 9 0 0 9 167.0045 486.6335 Tm (\002)Tj /T1_2 1 Tf 1.163 0.812 Td (I)Tj /T1_7 1 Tf 0.45 0 Td (\003)Tj /T1_2 1 Tf 6.3977 0 0 5.7614 185.28 492.3466 Tm (A)Tj ET 177.261 489.884 m 190.105 489.884 l S BT /T1_2 1 Tf 9 0 0 9 177.2614 480.9602 Tm (I)Tj /T1_7 1 Tf 0.45 0 Td (\003)Tj /T1_2 1 Tf 6.3977 0 0 5.7614 185.0715 479.3693 Tm (D)Tj /T1_5 1 Tf 9 0 0 9 192.112 486.6335 Tm (.)Tj /T1_8 1 Tf 7 0 0 7 51.6761 463.9176 Tm (I)Tj 5 0 0 5 53.4858 462.9176 Tm (A)Tj /T1_0 1 Tf -0.0195 Tc 7 0 0 7 59.9514 463.9176 Tm (and)Tj /T1_8 1 Tf 0 Tc 2.141 0 Td (I)Tj 5 0 0 5 76.7499 462.9176 Tm (D)Tj /T1_0 1 Tf -0.0195 Tc 7 0 0 7 83.2154 463.9176 Tm [(are)-422(background-corrected)-422(intensities)-422(from)-422(the)-422(acceptor)-422(and)-422(donor)]TJ -4.506 -1.287 Td [(channels)-295(and)]TJ /T1_8 1 Tf 0 Tc 6.211 0 Td (I)Tj 5 0 0 5 96.9609 453.9062 Tm (A)Tj /T1_0 1 Tf 7 0 0 7 100.4747 454.9062 Tm (/)Tj /T1_8 1 Tf 0.258 0 Td (I)Tj 5 0 0 5 104.0941 453.9062 Tm (D)Tj /T1_0 1 Tf -0.0195 Tc 7 0 0 7 109.6761 454.9062 Tm (and)Tj /T1_8 1 Tf 0 Tc 2.015 0 Td (I)Tj /T1_7 1 Tf 0.259 0 Td (\003)Tj /T1_8 1 Tf 5 0 0 5 127.7858 453.9062 Tm (A)Tj /T1_0 1 Tf 7 0 0 7 131.2996 454.9062 Tm (/)Tj /T1_8 1 Tf 0.258 0 Td (I)Tj /T1_7 1 Tf 0.258 0 Td (\003)Tj /T1_8 1 Tf 5 0 0 5 137.1136 453.9062 Tm (D)Tj /T1_0 1 Tf -0.0195 Tc 7 0 0 7 142.6957 454.9062 Tm [(are)-295(the)-295(intensities)-295(before)-295(and)-295(after)-295(acceptor)-295(pho-)]TJ -13.003 -1.287 Td [(tobleaching,)-174(respectively.)]TJ /T1_3 1 Tf 0 Tc 7 0 1.0881 7 131.6177 445.8949 Tm (\004)Tj /T1_0 1 Tf -0.0195 Tc 7 0 0 7 136.98 445.8949 Tm (and)Tj /T1_3 1 Tf 0 Tc 7 0 1.0881 7 150.2387 445.8949 Tm (\006)Tj /T1_0 1 Tf -0.0195 Tc 7 0 0 7 155.216 445.8949 Tm [(are)-174(experimental)-174(correction)-174(factors:)]TJ /T1_3 1 Tf 0 Tc 7 0 1.0881 7 266.4633 445.8949 Tm (\004)Tj /T1_0 1 Tf -0.0195 Tc 7 0 0 7 271.8257 445.8949 Tm (accounts)Tj -31.45 -1.287 Td [(for)-195(the)-195(leakage)-195(of)-195(the)-195(donor)-195(emission)-195(into)-195(the)-195(acceptor)-195(channel,)-195(and)]TJ /T1_3 1 Tf 0 Tc 7 0 1.0881 7 264.5088 436.8835 Tm (\006)Tj /T1_0 1 Tf -0.0195 Tc 7 0 0 7 269.631 436.8835 Tm [(is)-195(a)-195(factor)]TJ -31.136 -1.287 Td [(that)-205(includes)-205(the)-205(quantum)-205(yields)-205(of)-205(the)-205(\037uorophores)-205(and)-205(the)-205(detection)-205(ef\036cien-)]TJ 0 -1.287 TD [(cies)-222(of)-222(the)-222(two)-222(channels.)-222(We)-222(determined)-222(the)-222(correction)-222(factors)-222(for)-222(all)-222(FRET)-222(pairs)]TJ 0 -1.287 TD [(individually)-469(by)-469(time-averaging)-469(the)-469(intensities)]TJ /T1_8 1 Tf 0 Tc 21.422 0 Td (I)Tj /T1_0 1 Tf -0.0195 Tc 0.728 0 Td (and)Tj /T1_8 1 Tf 0 Tc 2.189 0 Td (I)Tj /T1_7 1 Tf 0.258 0 Td (\003)Tj /T1_0 1 Tf -0.0195 Tc 0.314 0 Td [(.)-469(FRET)-469(pairs)-469(where)-469(no)]TJ -24.91 -1.287 Td [(acceptor)-254(bleaching)-254(was)-254(observed)-254(were)-254(discarded)-254(from)-254(the)-254(analysis,)-254(because)-254(for)]TJ 0 -1.287 TD [(these)-184(FRET)-184(pairs)]TJ /T1_3 1 Tf 0 Tc 7 0 1.0881 7 102.2576 391.8267 Tm (\006)Tj /T1_0 1 Tf -0.0195 Tc 7 0 0 7 107.3003 391.8267 Tm [(could)-184(not)-184(be)-184(determined.)-184(Direct)-184(excitation)-184(of)-184(the)-184(acceptor)-184(was)]TJ -7.946 -1.287 Td [(so)-303(low)-303(\()]TJ /T1_7 1 Tf 0 Tc 3.544 0 Td (\t)Tj /T1_0 1 Tf -0.0195 Tc 0.813 0 Td [(5%\))-303(that)-303(it)-303(did)-303(not)-303(lead)-303(to)-303(detectable)-303(changes)-303(in)-303(the)-303(histograms)-303(\()]TJ /T1_7 1 Tf 0 Tc 29.809 0 Td (\n)Tj /T1_8 1 Tf 0.648 0 Td (E)Tj /T1_7 1 Tf -34.813 -1.287 Td (\t)Tj /T1_0 1 Tf -0.0195 Tc 0.814 0 Td [(1%\))-429(and)-429(therefore)-429(was)-429(disregarded)-429(in)-429(the)-429(analysis.)-429(FRET)-429(ef\036ciencies)-429(were)]TJ -0.814 -1.287 Td [(calculated)-323(for)-323(every)-323(time)-323(point)-323(after)-323(\036ltering)-323(the)-323(original)-323(data)-323(sets)-323(by)-323(using)-323(a)]TJ T* [(10-point)-352(sliding-average)-352(\036lter.)-352(The)-352(histograms)-352(of)-352(these)-352(FRET)-352(values)-352(represent)]TJ 37.5 54.3 Td [(transfer)-215(ef\036ciency)-215(distributions)-215(and)-215(describe)-215(dynamics)-215(of)-215(nascent)-215(RNA)-215(products.)]TJ 0 -1.284 TD [(Although)-376(most)-376(of)-376(the)-376(observed)-376(histograms)-376(show)-376(a)-376(fairly)-376(narrow)-376(distribution)]TJ T* (\(width)Tj /T1_7 1 Tf 0 Tc 3.117 0 Td (\004)Tj /T1_0 1 Tf -0.0195 Tc 0.813 0 Td [(0.2\),)-179(there)-179(are)-179(some)-179(examples)-179(of)-179(broadened)-179(histograms,)-179(which)-179(could)-179(be)]TJ -3.93 -1.284 Td [(caused,)-258(for)-258(example,)-259(by)-259(a)-259(lower)-259(signal)-258(or)-258(an)-259(increased)-259(mobility)-258(of)-258(the)-259(RNA.)]TJ 0 Tc 1.286 -1.284 Td [(From)-397(the)-397(FRET)-397(ef\036ciencies,)-397(we)-397(computed)-397(distances)-397(using)-397(the)-397(following)]TJ -1.286 -1.284 Td (equation:)Tj /T1_2 1 Tf 9 0 0 9 404.3494 661.2046 Tm (d)Tj /T1_3 1 Tf 0.77 0 Td (\002)Tj /T1_2 1 Tf 1.075 0 Td (R)Tj /T1_5 1 Tf 6.3977 0 0 5.7614 427.1679 659.6136 Tm (0)Tj /T1_9 1 Tf 22 0 0 25.196 430.4946 660.3153 Tm (\003)Tj /T1_5 1 Tf 9 0 0 9 436.6088 668.5085 Tm (1)Tj ET 435.813 664.455 m 442.074 664.455 l S BT /T1_2 1 Tf 9 0 0 9 435.8125 655.5312 Tm (E)Tj /T1_3 1 Tf 0.918 0.63 Td (\003)Tj /T1_5 1 Tf 1.075 0 Td (1)Tj /T1_9 1 Tf 22 0 0 25.196 458.424 660.3153 Tm (\004)Tj /T1_5 1 Tf 6.3977 0 0 5.7614 463.7484 676.6307 Tm (1)Tj ET 463.744 674.168 m 467.071 674.168 l S BT /T1_5 1 Tf 6.3977 0 0 5.7614 463.7443 668.8239 Tm (6)Tj 9 0 0 9 469.071 661.2046 Tm (,)Tj /T1_6 1 Tf 0.0199 Tc 8.868 0 Td ([2])Tj /T1_0 1 Tf -0.0195 Tc 7 0 0 7 314.1761 638.4687 Tm (where)Tj /T1_8 1 Tf 0 Tc 3.103 0 Td (R)Tj /T1_0 1 Tf 5 0 0 5 340.0373 637.4687 Tm (0)Tj -0.0195 Tc 7 0 0 7 344.508 638.4687 Tm [(is)-255(the)-255(Fo)]TJ 3.283 0 Td [(\250)-166(rster)-255(radius.)-255(We)-255(used)-255(a)-255(standard)-255(procedure)-255(\(45\))-255(to)-255(determine)]TJ /T1_8 1 Tf 0 Tc -7.616 -1.284 Td (R)Tj /T1_0 1 Tf 5 0 0 5 318.3169 628.4801 Tm (0)Tj -0.0195 Tc 7 0 0 7 322.5063 629.4801 Tm [(experimentally)-215(by)-215(measuring)-215(the)-215(donor)-215(quantum)-215(yield)-215(\(using)-215(rhodamine)-215(101)]TJ -1.19 -1.284 Td [(as)-224(a)-224(standard\))-224(and)-224(donor)-224(emission)-224(and)-224(acceptor)-224(absorption)-224(spectra)-224(\(to)-224(compute)]TJ 0 -1.284 TD [(the)-211(overlap)-211(integral\))-211(and)-211(assuming)-211(a)-211(free)-211(diffusion)-211(of)-211(the)-211(dye)-211(molecule)-211(\()]TJ 0 0 1 rg /T1_8 1 Tf 31.907 0 Td [(SI)-211(Text)]TJ 0 0 0 rg /T1_0 1 Tf 2.873 0 Td (\).)Tj -33.494 -1.284 Td [(Different)-165(effects)-165(such)-165(as)-165(dye-linker)-165(structure)-165(and)-165(dynamics,)-165(uncertainties)-165(in)]TJ /T1_8 1 Tf 0 Tc 32.83 0 Td (R)Tj /T1_0 1 Tf 5 0 0 5 557.1242 601.5142 Tm (0)Tj 7 0 0 7 559.808 602.5142 Tm (,)Tj -0.0195 Tc -35.09 -1.284 Td [(and)-382(dynamics)-382(of)-382(the)-382(RNA)-382(itself)-382(all)-382(contribute)-382(to)-382(the)-382(experimental)-382(error.)-382(The)]TJ T* [(precise)-279(error)-279(of)-279(the)-279(measurement)-279(therefore)-279(can)-279(only)-279(be)-279(estimated.)-279(The)-279(experi-)]TJ T* [(mental)-245(tests)-245(performed)-245(in)-245(this)-245(work)-245(\(comparison)-245(of)-245(experimentally)-245(determined)]TJ T* [(distances)-174(to)-174(distances)-174(known)-174(from)-174(the)-174(crystal)-174(structure)-174(and)-174(comparison)-174(of)-174(fourth)]TJ 0 -1.284 TD [(measured)-403(distance)-403(to)-403(expected)-403(distance\))-403(yielded)-403(errors)]TJ /T1_7 1 Tf 0 Tc 25.783 0 Td (\t)Tj /T1_0 1 Tf -0.0195 Tc 0.813 0 Td [(5)-403(\305.)-403(Therefore,)-403(we)]TJ -26.597 -1.284 Td [(assume)-259(an)-259(experimental)-258(error)-258(of)]TJ /T1_7 1 Tf 0 Tc 14.827 0 Td (\004)Tj /T1_0 1 Tf -0.0195 Tc 0.814 0 Td [(5)-259(\305)-259(throughout)-258(this)-259(work.)]TJ /T1_1 1 Tf 0 Tc -15.64 -2.713 Td (Trilateration.)Tj /T1_0 1 Tf -0.0195 Tc 6.079 0 Td [(For)-187(each)-187(RNA)-187(length,)-187(we)-187(computed)-187(three)-187(distances)-187(using)-187(the)-187(three)]TJ -6.079 -1.284 Td [(recorded)-270(histograms.)-270(To)-270(infer)-270(from)-270(these)-270(three)-270(distances)-270(the)-270(unknown)-270(relative)]TJ T* [(position)-190(of)-190(the)-190(dye)-190(molecule,)-190(we)-190(mathematically)-190(construct)-190(a)-190(sphere)-190(around)-190(each)]TJ T* [(of)-140(the)-140(three)-140(known)-140(positions)-140(with)-140(a)-140(radius)-140(determined)-140(by)-140(the)-140(FRET)-140(measurement.)]TJ T* [(For)-275(the)-275(attachment)-275(points)-275(on)-275(DNA1)-275(and)-275(DNA2,)-275(we)-275(used)-275(the)-275(position)-275(of)-275(the)-275(C6)]TJ 0 -1.284 TD [(atom,)-252(and)-252(in)-252(the)-252(case)-252(of)-252(the)-252(labeling)-252(sites)-252(on)-252(Rpb7,)-252(we)-252(used)-252(the)-252(position)-252(of)-252(the)]TJ T* [(respective)-248(sulfur)-248(atom)-248(as)-248(the)-248(center)-248(of)-248(the)-248(sphere.)-248(The)-248(intersection)-248(point)-248(of)-248(the)]TJ T* [(three)-352(spheres)-352(determines)-352(the)-352(unknown)-352(position)-352(\(20\).)-352(It)-352(should)-352(be)-352(noted)-352(that)]TJ T* [(mathematically)-180(there)-180(might)-180(be)-180(as)-180(many)-180(as)-180(two)-180(intersection)-180(points;)-180(however,)-180(one)]TJ T* [(of)-218(these)-218(points)-218(can)-218(be)-218(ruled)-218(out)-218(because)-218(of)-218(distance)-218(constraints,)-218(that)-218(is,)-218(the)-218(RNA)]TJ T* [(simply)-370(is)-370(not)-370(long)-370(enough)-370(to)-370(reach)-370(the)-370(second)-370(point.)-370(Alternatively,)-370(the)-370(right)]TJ T* [(intersection)-259(point)-259(can)-259(be)-259(determined)-259(by)-258(measuring)-259(a)-259(fourth)-259(distance.)]TJ /T1_10 1 Tf 0 Tc 0 -2.713 TD (ACKNOWLEDGMENTS.)Tj /T1_0 1 Tf -0.0195 Tc 10.783 0 Td [(We)-170(thank)-170(Marta)-170(Blamowska)-170(for)-170(help)-170(in)-170(determining)-170(the)]TJ -10.783 -1.141 Td (Fo)Tj 0.647 0 Td [(\250)-166(rster)-400(radii,)-400(Peter)-400(Schwaderer)-400(for)-400(help)-400(with)-400(the)-400(trilateration)-400(procedure,)-400(Jo-)]TJ -0.647 -1.141 Td [(hannes)-360(Gierlich)-360(and)-360(Thomas)-360(Carell)-360(for)-360(the)-360(HPLC)-360(puri\036cation)-360(of)-360(labeled)-360(DNA,)]TJ 0 -1.141 TD [(Anass)-328(Jawhari)-328(for)-328(the)-328(expression)-328(and)-328(puri\036cation)-328(of)-328(TFIIB,)-328(Hazen)-328(Babcock)-328(for)]TJ 0 -1.143 TD [(helpful)-219(discussions)-219(regarding)-219(the)-219(data)-219(analysis,)-219(and)-219(Christoph)-219(Bra)]TJ 28.506 0 Td [(\250)-139(uchle)-219(and)-219(Don)]TJ -28.506 -1.143 Td [(Lamb)-416(for)-416(fruitful)-416(discussions.)-416(The)-416(work)-416(was)-416(supported)-416(by)-416(the)-416(Deutsche)-416(For-)]TJ 0 -1.143 TD [(schungsgemeinschaft)-458(\(SFB)-458(646\),)-458(the)-458(Nanosystems)-458(Initiative)-458(Munich,)-458(and)-458(the)]TJ T* [(Center)-258(for)-258(Nanoscience)-259(\(CeNS\))-258(at)-258(Ludwig-Maximilans-Universita)]TJ 0.1193 Tc 28.141 0 Td [(\250t)-120(M)139(u)]TJ -0.0195 Tc 2.117 0 Td [(\250)-166(nchen.)]TJ 0 Tc 6 0 0 6 55.6733 331.6733 Tm [(1.)-500(Cramer)-278(P,)-278(Bushnell)-278(DA,)-278(Kornberg)-278(RD)-278(\(2001\))]TJ /T1_8 1 Tf 21.839 0 Td (Science)Tj /T1_0 1 Tf 3.667 0 Td (292:1863\2261876.)Tj -25.507 -1.333 Td [(2.)-500(Armache)-243(KJ,)-243(Mitterweger)-243(S,)-243(Meinhart)-243(A,)-243(Cramer)-243(P)-243(\(2005\))]TJ /T1_8 1 Tf 27.86 0 Td [(J)-243(Biol)-243(Chem)]TJ /T1_0 1 Tf 5.563 0 Td (280:7131\2267134.)Tj -33.423 -1.333 Td [(3.)-500(Kettenberger)-284(H,)-284(Armache)-284(KJ,)-284(Cramer)-284(P)-284(\(2004\))]TJ /T1_8 1 Tf 22.937 0 Td [(Mol)-284(Cell)]TJ /T1_0 1 Tf 4.123 0 Td (16:955\226965.)Tj -27.061 -1.333 Td [(4.)-500(Westover)-278(KD,)-278(Bushnell)-278(DA,)-278(Kornberg)-278(RD)-278(\(2004\))]TJ /T1_8 1 Tf 23.673 0 Td (Cell)Tj /T1_0 1 Tf 2.001 0 Td [(119:481\226)-120(489.)]TJ -25.674 -1.333 Td [(5.)-500(Armache)-293(KJ,)-293(Kettenberger)-293(H,)-293(Cramer)-293(P)-293(\(2005\))]TJ /T1_8 1 Tf 23 0 Td [(Curr)-293(Opin)-293(Struct)-293(Biol)]TJ /T1_0 1 Tf 9.949 0 Td (15:197\226203.)Tj -32.949 -1.333 Td [(6.)-500(Gnatt)-278(AL,)-278(Cramer)-278(P,)-278(Fu)-278(J,)-278(Bushnell)-278(DA,)-278(Kornberg)-278(RD)-278(\(2001\))]TJ /T1_8 1 Tf 28.897 0 Td (Science)Tj /T1_0 1 Tf 3.667 0 Td [(292:1876)-120(\2261882.)]TJ -32.564 -1.333 Td [(7.)-500(Westover)-278(KD,)-278(Bushnell)-278(DA,)-278(Kornberg)-278(RD)-278(\(2004\))]TJ /T1_8 1 Tf 23.673 0 Td (Science)Tj /T1_0 1 Tf 3.667 0 Td [(303:1014)-120(\2261016.)]TJ -27.34 -1.333 Td [(8.)-500(Korzheva)-157(N,)-157(Mustaev)-157(A,)-157(Kozlov)-157(M,)-157(Malhotra)-157(A,)-157(Nikiforov)-157(V,)-157(Goldfarb)-157(A,)-157(Darst)-157(SA)-157(\(2000\))]TJ /T1_8 1 Tf 1.334 -1.333 Td (Science)Tj /T1_0 1 Tf 3.667 0 Td [(289:619)-120(\226)-120(625.)]TJ -5.001 -1.333 Td [(9.)-500(Cramer)-426(P,)-426(Bushnell)-426(DA,)-426(Fu)-426(J,)-426(Gnatt)-426(AL,)-426(Maier-Davis)-426(B,)-426(Thompson)-426(NE,)-426(Burgess)-426(RR,)]TJ 1.334 -1.333 Td [(Edwards)-278(AM,)-278(David)-278(PR,)-278(Kornberg)-278(RD)-278(\(2000\))]TJ /T1_8 1 Tf 20.616 0 Td (Science)Tj /T1_0 1 Tf 3.667 0 Td [(288:640)-120(\226)-120(649.)]TJ -26.172 -1.333 Td [(10.)-500(Ujvari)-290(A,)-290(Luse)-290(DS)-290(\(2006\))]TJ /T1_8 1 Tf 13.23 0 Td [(Nat)-290(Struct)-290(Mol)-290(Biol)]TJ /T1_0 1 Tf 9.16 0 Td [(13:49)-120(\22654.)]TJ -22.39 -1.333 Td [(11.)-500(Sosa)-282(H,)-282(Peterman)-282(EJ,)-282(Moerner)-282(WE,)-282(Goldstein)-282(LS)-282(\(2001\))]TJ /T1_8 1 Tf 27.437 0 Td [(Nat)-282(Struct)-282(Biol)]TJ /T1_0 1 Tf 7.014 0 Td [(8:540)-120(\226544.)]TJ -0.0194 Tc -34.451 -1.333 Td [(1)-19(2)-19(.)-519(Yildiz)-263(A,)-263(Forkey)-263(JN,)-263(McKinney)-263(SA,)-263(Ha)-263(T,)-263(Goldman)-263(YE,)-263(Selvin)-263(PR)-263(\(2003\))]TJ /T1_8 1 Tf 33.122 0 Td (Science)Tj /T1_0 1 Tf 3.517 0 Td (300:2061\226)Tj -34.749 -1.333 Td (2065.)Tj 0 Tc -1.89 -1.333 Td [(13.)-500(Zhuang)-298(X,)-298(Bartley)-298(LE,)-298(Babcock)-298(HP,)-298(Russell)-298(R,)-298(Ha)-298(T,)-298(Herschlag)-298(D,)-298(Chu)-298(S)-298(\(2000\))]TJ /T1_8 1 Tf 37.761 0 Td (Science)Tj /T1_0 1 Tf -35.871 -1.333 Td [(288:2048)-120(\2262051.)]TJ -1.89 -1.333 Td [(14.)-500(Schmidt)-197(T,)-197(Schutz)-197(GJ,)-197(Baumgartner)-197(W,)-197(Gruber)-197(HJ,)-197(Schindler)-197(H)-197(\(1996\))]TJ /T1_8 1 Tf 33.061 0 Td [(Proc)-197(Natl)-197(Acad)-197(Sci)]TJ -31.171 -1.333 Td (USA)Tj /T1_0 1 Tf 2.222 0 Td [(93:2926)-120(\2262929.)]TJ -4.112 -1.333 Td [(15.)-500(Ha)-279(T,)-279(Enderle)-279(T,)-279(Ogletree)-279(DF,)-279(Chemla)-279(DS,)-279(Selvin)-279(PR,)-279(Weiss)-279(S)-279(\(1996\))]TJ /T1_8 1 Tf 32.81 0 Td [(Proc)-279(Natl)-279(Acad)-279(Sci)]TJ -30.92 -1.333 Td (USA)Tj /T1_0 1 Tf 2.222 0 Td [(93:6264)-120(\226)-120(6268.)]TJ -4.112 -1.333 Td [(16.)-500(Weiss)-278(S)-278(\(1999\))]TJ /T1_8 1 Tf 8.726 0 Td (Science)Tj /T1_0 1 Tf 3.667 0 Td [(283:1676)-120(\2261683.)]TJ -12.393 -1.333 Td [(17.)-500(Brueckner)-278(F,)-278(Hennecke)-278(U,)-278(Carell)-278(T,)-278(Cramer)-278(P)-278(\(2007\))]TJ /T1_8 1 Tf 25.843 0 Td (Science)Tj /T1_0 1 Tf 3.667 0 Td [(315:859)-120(\226)-120(862.)]TJ 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