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[(u)12.2(s)0(e)19.1(d)-247.9(to)-232.5(s)11(p)12.2(ec)16(i)20.1(c)13.4(a)0(l)18(l)0(y)-232.5(d)0(i)16.8(r)13.2(ec)16(t)-244.7(n)12.2(o)12.2(n)0(-)20.1(c)0(l)18(e)0(a)21.4(v)0(i)16.8(n)12.2(g)-242.5(C)10.2(as)19.1(9)-247.9(R)10.2(N)13.5(P)0(s)-233.6(t)0(o)-237.9(o)12.2(ff)21.1(-)]TJ
T*
[(ta)18(r)13.2(g)0(e)20.3(t)-504.8(s)11(e)0(q)20.3(u)0(e)20.2(n)0(c)20.3(e)0(s)19.1(,)-503.3(th)22.3(er)15.8(eb)20.3(y)-502.6(p)12.2(re)21.3(ve)20.3(nt)16.8(in)22.3(g)-502.6(a)0(c)21.5(t)0(i)20(v)0(e)-494.6(C)10.2(as)19.1(9)-502.6(R)10.2(NP)]TJ
0 -1.1869 TD
[(fr)21.1(om)-317.8(b)12.2(i)0(n)16.8(d)12.2(in)22.3(g)-329.2(t)0(h)22.3(e)0(s)19.1(e)-328(o)12.2(f)0(f)21.1(-)0(t)17.8(a)0(r)21.3(g)12.2(et)-318(s)11(i)0(t)20(e)0(s)19.1(.)]TJ
7.8454 0 0 6.9729 207.5527 192.3023 Tm
-.0158 Tc
(51)Tj
10.4607 0 0 10.4607 218.6645 187.6535 Tm
0 Tc
[(La)20.4(st)15.7(ly)22.3(,)-324.5(D)0(N)16.1(A)-322.6(ol)16.8(ig)22.3(o)12.2(s)]TJ
-14.9744 -1.1923 TD
[(wi)18.1(th)-189.1(ph)19.1(o)12.2(s)0(p)17.9(h)12.2(or)20.1(o)12.2(t)0(h)16.8(i)0(o)22.3(a)0(t)18(e)-203.4(li)20(n)12.2(k)12.2(a)0(g)14.8(e)13.4(s)-205.7(d)12.2(is)21.1(pl)16.8(a)13.4(y)0(e)20.3(d)-204.5(s)11(t)0(r)17.8(o)12.2(ng)-192.3(in)16.8(h)12.2(i)0(b)22.3(i)0(t)20(i)0(o)22.3(n)]TJ
T*
[(o)12.2(f)-203.5(C)10.2(a)0(s)19.1(1)0(2)19.1(a)-203.4(ac)16(ti)20(v)12.2(i)0(t)20(y)12.2(,)-205.2(i)0(n)22.3(d)12.2(ep)20.3(en)20.3(de)14.8(n)12.2(t)-201.3(of)-196.7(th)22.3(e)-203.4(n)12.2(uc)20.3(le)18(o)12.2(t)0(i)20(d)0(e)-196.5(s)11(eq)20.3(u)12.2(e)0(n)20.2(c)0(e)]TJ
0 -1.1869 TD
[(o)12.2(f)-290.3(th)16.8(e)-290.1(i)0(n)22.3(h)12.2(ib)16.8(it)20(o)12.2(r)-290.3(u)12.2(s)11(ed)20.3(.)]TJ
7.8454 0 0 6.9729 148.9322 154.9417 Tm
-.0158 Tc
(49)Tj
10.4607 0 0 10.4607 72.9637 137.8204 Tm
0 Tc
[(In)-527.3(the)-522.5(current)-524.5(study,)-521.4(we)-525.9(assessed)-526.5(whether)-524.3(SpCas9)]TJ
-1.046 -1.1923 TD
[(could)-330(be)-332(inhibited)-325.7(with)-328.2(guide-complementary)-331.4(DNA)-328.2(oli-)]TJ
0 -1.1869 TD
[(gonucleotides)-468.4(without)-461.7(any)-466.1(chemical)-471.6(modications.)-462.5(To)]TJ
0 -1.1923 TD
[(this)-439(end,)-438.3(we)-439.2(designed)-439.6(and)-439(tested)-439.2(different)-436.2(oligo-based)]TJ
T*
[(inhibitors)-412.4(complementary)-410.2(to)-411.3(the)-408.7(spacer-derived)-413.8(part)-411.7(of)]TJ
0 -1.1869 TD
[(the)-316.6(gRNA)-313.7(\(these)-316.8(oligos)-311.6(thus)-312.1(have)-317.2(the)-316.6(same)-313.8(sequence)-315.8(as)]TJ
0 -1.1923 TD
[(the)-522.5(PAM-proximal)-511.7(part)-520.1(of)-521.9(the)-517.1(protospacer\).)-521.2(We)-519.1(also)]TJ
23.998 60.7159 TD
[(investigated)-304(the)-300.3(effect)-302.6(of)-299.7(extending)-298.4(the)-300.3(inhibitors)-298.5(with)-301.1(a)]TJ
0 -1.1923 TD
[(double-stranded)-396.3(PAM)-397.6(sequence.)-396.4(We)-394.5(show)-395.4(that)-398.7(various)]TJ
T*
[(designs)-508.5(provide)-505.1(strong,)-502.7(sequence-dependent)-505.4(inhibition)]TJ
0 -1.1869 TD
[(of)-299.7(Cas9)]TJ
/F8 1 Tf
3.4306 0 TD
[(in)-302.9(vitro)]TJ
/F7 1 Tf
3.2789 0 TD
(and)Tj
/F8 1 Tf
1.7451 0 TD
[(ex)-298.3(vivo)]TJ
/F7 1 Tf
2.867 0 TD
(.)Tj
-10.2702 -1.1923 TD
[(In)-343.1(addition,)-340.4(we)-341.6(investigated)-341.9(the)-343.7(effect)-345.9(of)-343.1(oligo-based)]TJ
-1.0514 -1.1923 TD
[(Cas9)-576.3(inhibition)-571.6(on)-571.7(off-target)-574.7(sites)-578.5(in)-573.9(the)-571.3(context)-572.1(of)]TJ
0 -1.1869 TD
[(genome)-320.6(editing.)-314.7(We)-318.6(found)-311.7(that)-317.4(while)-320.2(inhibition)-311.4(reduces)]TJ
0 -1.1923 TD
[(both)-316.4(on-)-314.5(and)-319.8(off-target)-314.6(activity)-315(of)-316(Cas9,)-315.5(the)-316.6(presence)-314.8(of)]TJ
T*
[(specic)-312(oligo)-306.4(designs)-307.9(results)-309.1(in)-308.3(slightly)-302.4(increased)-310.2(speci-)]TJ
0 -1.1869 TD
[(city.)-447.6(By)-449(comparing)-442.3(the)-446.7(inhibitor)-445.1(results)-444.6(with)-447.4(a)-447.2(Cas9)]TJ
0 -1.1923 TD
[(titration,)-358.7(we)-363.3(conclude)-356(that)-360.8(the)-359.9(increased)-359(specicity)-361(is)-358.3(a)]TJ
T*
[(general)-302.2(consequence)-299.6(of)-299.7(lowering)-298.7(overall)-300.1(Cas9)-299.9(activity.)]TJ
1.0514 -1.1869 TD
[(Lastly,)-558.8(we)-558.4(studied)-558.3(which)-556.4(mechanisms)-557.7(lead)-557.9(to)-557.6(the)]TJ
-1.0514 -1.1923 TD
[(observed)-604.4(inhibition.)-598(We)-605.8(conclude)-599.8(that)-604.6(the)-603.8(effect)-600.6(of)]TJ
T*
[(the)-370.8(tested)-368.8(inhibitors)-363.6(is)-369.1(dependent)-370.8(both)-365.2(on)-365.7(the)-370.8(length)-363.4(of)]TJ
0 -1.1869 TD
-.0153 Tc
[(t)-15.3(h)12.4(e)-316.2(o)-3.1(ligo-based)-317.4(i).1(nhibitors)-318.6(and)-317.4(o)-3.1(n)-317.4(t).1(he)-321.6(presence)-316.2(of)-316.4(a)-316.2(P).2(AM-)]TJ
0 -1.1923 TD
0 Tc
[(loop.)-418.6(Their)-421(relative)-419(importance)-420.2(is)-423.3(strongly)-416.1(affected)-417.8(by)]TJ
T*
[(the)-300.3(speed)-300.7(at)-301.7(which)-301.7(Cas9)-299.9(cleaves)-297.7(the)-300.3(targeted)-303(DNA.)]TJ
/F5 1 Tf
/Cs8 cs 1 scn
0 -2.2708 TD
[(Materials)-302.5(and)-303.8(Methods)]TJ
/F10 1 Tf
0 -1.2411 TD
[(Cell)-301.8(line)-300.1(and)-298.4(cell)-296.7(culture)]TJ
/F7 1 Tf
0 0 0 rg
0 -1.1977 TD
[(The)-238.3(chronic)-232.9(myelogenous)-230.3(leukemia)-234.3(\(CML\))-231(cell)-234.9(line)-236.1(used)]TJ
0 -1.1923 TD
[(is)-249.9(a)-246.7(rediploidized)-246.2(derivative)-248.2(of)-245.5(the)-246.1(HAP1)-243.5(cell)-251.2(line,)-246.2(which)]TJ
T*
[(was)-265.6(a)-263(kind)-256.8(gift)-263.4(of)-261.8(Dr.)-259.8(Thijn)-262.8(Brummelkamp.)]TJ
7.8454 0 0 6.9729 496.9699 416.9196 Tm
(54)Tj
10.4607 0 0 10.4607 507.5148 412.2708 Tm
[(CML)-263(cells)]TJ
-18.5893 -1.1869 TD
[(were)-352.9(cultured)-352.9(in)-351.7(Iscoves)-349.9(modied)-346.5(Dulbeccos)-352.1(medium)]TJ
0 -1.1923 TD
[(\(Gi)15(bco)10.8(\),)-295(su)12.4(pp)13.6(lem)13.2(ent)14(ed)-294.1(wi)12.6(th)-297.5(10)13.6(%)-299.7(fet)15(al)-296.3(bov)15(in)11.4(e)-300.9(ser)16(um)-296.1(and)]TJ
T*
[(1%)-439.2(penicillin/strepto)5.5(mycin.)-438.5(Cells)-433(were)-439.6(grown)-435.2(at)-437.2(37)]TJ
/F11 1 Tf
21.7977 0 TD
()Tj
/F7 1 Tf
.3956 0 TD
(C)Tj
-22.1933 -1.1869 TD
[(in)-302.9(a)-300.9(humidied)-297.1(atmosphere)-300(containing)-293.8(5%)-298.3(CO)]TJ
7.8454 0 0 6.9729 511.1998 360.7369 Tm
(2)Tj
10.4607 0 0 10.4607 515.1117 362.4944 Tm
(.)Tj
/F10 1 Tf
/Cs8 cs 1 scn
-19.3155 -2.3684 TD
[(Cas9)-304(protein,)-297.7(gRNA,)-299.9(and)-298.4(inhibitors)-299.7(used)]TJ
/F9 1 Tf
17.0447 0 TD
[(ex)-302.9(vivo)]TJ
/F7 1 Tf
0 0 0 rg
-17.0447 -1.2032 TD
(Recombinant)Tj
/F8 1 Tf
5.8152 0 TD
[(S.)-464(pyogenes)]TJ
/F7 1 Tf
5.4034 0 TD
[(Cas9)-462.5(\(variant)-465.8(with)-463.7(multiple)]TJ
-11.2186 -1.1923 TD
[(nuclear)-426.8(localization)-417.6(signals\))]TJ
7.8454 0 0 6.9729 434.4377 317.3102 Tm
(55)Tj
10.4607 0 0 10.4607 446.6833 312.6613 Tm
[(and)-422.8(high-delity)-420.2(protein)]TJ
-12.774 -1.1923 TD
[(variants)-522.1(\(SpCas9-HF1)]TJ
7.8454 0 0 6.9729 409.0392 304.8377 Tm
(56)Tj
10.4607 0 0 10.4607 422.3054 300.1889 Tm
[(or)-516.5(evoCas9\))]TJ
7.8454 0 0 6.9729 475.9369 304.8377 Tm
(57)Tj
10.4607 0 0 10.4607 489.203 300.1889 Tm
[(were)-515.5(provided)]TJ
-16.8387 -1.1869 TD
-.0001 Tc
[(by)-750.6(Geijsen)-745.6(lab)-750.3(through)-744.8(Divvly)-748.2(\(https://divvly.com/)]TJ
0 -1.1923 TD
0 Tc
[(geijsenlab\).)-480.3(gRNAs)-476.1(used)-471.3(are)-480.2(synthetic)-475.2(gRNAs,)-475.4(which)]TJ
T*
[(contain)-409.6(the)-414.1(target-specic)-410.6(CRISPR)-407.5(RNA)-412.7(\(crRNA\))-408.3(and)]TJ
0 -1.1869 TD
[(the)-403.3(scaffold)-405.1(trans-activating)-400.4(CRISPR)-402.1(RNA)-401.8(\(tracrRNA\))]TJ
0 -1.1923 TD
[(\(IDT\).)-340.1(The)-341.3(crRNA)-337.2(and)-341.5(tracrRNA)-338.5(were)-336.6(dissolved)-339.9(in)-335.4(the)]TJ
T*
[(nuclease-free)-349(Duplex)-349.1(buffer)-345.1(\(IDT\))-346.3(to)-346.3(reach)-351.5(the)-343.7(concen-)]TJ
0 -1.1869 TD
[(tration)-406.5(of)-402.7(200)]TJ
/F12 1 Tf
5.9291 0 TD
(l)Tj
/F7 1 Tf
.5528 0 TD
[(M.)-404.2(Equal)-400.3(volumes)-405.1(of)-402.7(dissolved)-399.5(crRNA)]TJ
-6.4819 -1.1923 TD
[(and)-363.1(tracrRNA)-360.1(were)-358.3(mixed)-358(and)-363.1(annealed)-360.2(by)-354.9(heating)-360.8(for)]TJ
T*
[(5)-166.6(min)-410.7(at)-410.1(95)]TJ
/F11 1 Tf
4.7801 0 TD
()Tj
/F7 1 Tf
.3956 0 TD
[(C)-407.1(and)-411.9(cooling)-405.3(down)-410.5(at)-410.1(room)-411.5(temperature.)]TJ
-5.1757 -1.1869 TD
[(The)-287.1(oligo-based)-290.3(inhibitors)-282.3(used)-287.1(were)-287.9(unmodied)-284.1(single-)]TJ
0 -1.1923 TD
[(stranded)-265.2(DNA)-263.1(oligos)-262.8(synthesized)-264.2(by)-262.8(IDT)-259(\(IDT\).)-264.3(Each)-262.8(of)]TJ
T*
[(such)-243.7(oligos)-241.1(was)-243.9(dissolved)-242.3(in)-243.3(nuclease-free)-240.6(water)-245.3(to)-243.3(reach)]TJ
0 -1.1869 TD
[(the)-300.3(concentration)-302.4(of)-299.7(75)]TJ
/F12 1 Tf
9.5927 0 TD
(l)Tj
/F7 1 Tf
.5474 0 TD
(M.)Tj
/F10 1 Tf
/Cs8 cs 1 scn
-10.1401 -2.3684 TD
[(Induced)-303.1(transduction)-301.9(by)-297.8(osmocytosis)]TJ
0 -1.2411 TD
[(and)-303.9(propanebetaine)]TJ
/F7 1 Tf
0 0 0 rg
0 -1.2032 TD
[(The)-243.7(recombinant)-240(Cas9)-245.7(proteins,)-240.8(gRNAs,)-236.9(and)-243.9(oligo-based)]TJ
0 -1.1869 TD
-.015 Tc
[(i)-15(n)12.7(hibitors)-307.5(were)-305.1(simultaneously)-306.3(t).4(ransduced)-300.8(into)-306.3(CML)-306.1(cells)]TJ
0 -1.1923 TD
[(by)-300.8(using)-306.3(t).4(he)-305.1(induced)-300.8(transduction)-300.8(b)2.7(y)-306.3(t).4(he)-305.1(osmocytosis)-302(a)-1.6(nd)]TJ
/F5 1 Tf
9.9626 0 0 9.9626 62.022 754.3557 Tm
0 Tc
(572)Tj
41.45 0 TD
[(SW)-6.6(ARTJES)-308.2(ET)-299.4(AL.)]TJ
ET
endstream
endobj
66 0 obj
<>stream
/GS1 gs
BT
/F7 1 Tf
10.4607 0 0 10.4607 59.7543 698.2297 Tm
0 0 0 rg
-.0136 Tc
0 Tw
[(pr)5.1(op)4.1(an)4.1(ebe)5.3(t)1.8(aine)-341.6(\(iTO)5.3(P\))-347.2(meth)4.1(od)-342.8(as)-344(d)4.1(e)-.2(scrib)4.1(e)-.2(d)-342.8(p)4.1(rev)4.1(i)1.8(ou)4.1(sly.)]TJ
7.8454 0 0 6.9729 291.1747 702.8785 Tm
-.0158 Tc
(58)Tj
10.4607 0 0 10.4607 59.7543 685.7573 Tm
-.0146 Tc
[(One)-380.5(d)3.1(ay)-376.3(be)4.3(for)4.1(e)-380.5(the)-380.5(t).8(r)4.1(a)-1.2(nsduc)4.3(t)-14.6(i)10.9(on,)-377(CML)-376.1(cells)-377.5(we)4.2(re)-380.5(plated)]TJ
0 -1.1923 TD
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[(at)-421.7(18,000)-419.5(cells/well)-421.7(i)1(n)-419.5(t)1(he)-418.3(Ma)]TJ
12.4001 0 TD
-.0155 Tc
[(trigel-coated)-420.6(w)-2(ells)-421.8(on)-426(96-)]TJ
-12.4001 -1.1869 TD
-.0159 Tc
[(well)-472(plates,)-470.5(s)-4.9(uch)-469.8(t)-.5(h)-3.7(a)3(t)-472(o)-3.7(n)-469.8(t)-.5(he)-474(d)-3.7(a)3(y)-475.2(o)1.8(f)-468.7(t)-15.9(r)12.8(a)-2.5(ns)-4.9(ductio)-3.7(n,)-470.4(c)-29.6(e)-24.1(l)-27.6(l)-27.6(s)]TJ
0 -1.1923 TD
.0095 Tc
[(would)-406.4(r)1.1(each)-406.4(about)-403.2(7080%)-404(conuence.)-407.1(Next)-403.2(d).1(ay,)-401.7(f)1.1(or)]TJ
T*
.0101 Tc
[(each)-340.8(well)-343(of)-345.2(the)-345(96-well)-343(plate,)-341.5(50)]TJ
/F12 1 Tf
14.4053 0 TD
0 Tc
(l)Tj
/F7 1 Tf
.5582 0 TD
.3548 Tc
[(Lo)345.4(fi)348.6(T)345.5(O)341.2(Pm)354.8(i)338.3(x)345.4(t)343.1(u)345.4(r)346.4(e)]TJ
-14.9636 -1.1869 TD
.0096 Tc
[(that)-397.7(contains)-396.7(20)]TJ
/F12 1 Tf
6.9479 0 TD
0 Tc
(l)Tj
/F7 1 Tf
.5636 0 TD
.0097 Tc
[(L)-395.2(o).3(f)-394.4(t)-2(ransduction)-395.4(supplement)-392.2(\()-4.2(Opti-)]TJ
-7.5116 -1.1923 TD
.0101 Tc
[(ME)-4.6(M)-378.5(m)-.2(edia)-382.9(\(Thermo)-378.7(F)-1.5(isher)-377.7(S)-1.5(cientic\))-377.7(supplemented)]TJ
T*
.0098 Tc
[(with)-763.8(542)-167.7(mM)-763.6(NaC)9.8(l)-7.9(,)-764.5(333)-167.6(mM)-763.6(gamma-aminobutyric)]TJ
0 -1.1869 TD
.0113 Tc
[(ac)3.1(id,)-351.1(1)1.9(.67)]TJ
/F11 1 Tf
4.2869 0 TD
0 Tc
()Tj
/F7 1 Tf
.7262 0 TD
.0106 Tc
[(N2,)-351.8(1)1.2(.67)]TJ
/F11 1 Tf
3.81 0 TD
0 Tc
()Tj
/F7 1 Tf
.7262 0 TD
.0098 Tc
[(B27,)-352.6(1.67)]TJ
/F11 1 Tf
4.2652 0 TD
0 Tc
()Tj
/F7 1 Tf
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.0102 Tc
[(non-ess)10.2(e)-8.7(ntial)-353.7(a)2(mino)]TJ
-14.5408 -1.1923 TD
.0099 Tc
[(acids,)-385(3.3)-162.1(m)-.4(M)-389.5(g).5(lutamine,)-385(167)-167.6(ng/mL)-384.1(bovine)-383.1(broblast)]TJ
T*
-.028 Tc
[(g)-37.4(r)-41.9(o)-37.4(w)-36.2(t)-39.7(h)-232.5(factor-basic,)-195.3(a)-3.7(nd)-194.6(84)-167.5(ng/)]TJ
11.7714 0 TD
-.0295 Tc
[(mL)-201.3(epider)-5.4(ma)-5.2(l)-198.3(g)-1(row)-5.2(t)2.2(h)-201.5(f)0(ac)-5.2(tor)-5.4(\))0(,)]TJ
-11.7714 -1.1869 TD
-.0285 Tc
(10)Tj
/F12 1 Tf
1.111 0 TD
0 Tc
(l)Tj
/F7 1 Tf
.5257 0 TD
-.0276 Tc
[(L)-307.8(o).9(f)-312.4(C)4.3(as9)-308(p).9(rotein)-308(\(75)]TJ
/F12 1 Tf
8.5467 0 TD
0 Tc
(l)Tj
/F7 1 Tf
.5257 0 TD
-.0285 Tc
[(M\),)-309.6(7)0(.5)]TJ
/F12 1 Tf
3.0241 0 TD
0 Tc
(l)Tj
/F7 1 Tf
.5203 0 TD
-.0283 Tc
[(L)-314(o).2(f)-307.7(g).2(RNA)-312.9(\()1.2(100)]TJ
/F12 1 Tf
6.6715 0 TD
0 Tc
(l)Tj
/F7 1 Tf
.5203 0 TD
-.0264 Tc
(M\),)Tj
-21.4454 -1.1923 TD
-.0285 Tc
(10)Tj
/F12 1 Tf
1.111 0 TD
0 Tc
(l)Tj
/F7 1 Tf
.5257 0 TD
-.0283 Tc
[(L)-389.8(o).2(f)-389(o).2(ligo-based)-390(i)-2(nhibitors,)-390.7(a)1.4(nd)-390(the)-388.8(e)-4(xcess)-391.2(volume)-388.8(o).2(f)]TJ
-1.6367 -1.1923 TD
-.029 Tc
[(nuc)-4.7(le)-4.7(ase)-4.7(-fre)-4.7(e)-460(w).7(a)-4.7(t)2.7(e)-4.7(r)-460.2(to)-455.8(r)-4.9(eac)-4.7(h)-461.2(t)2.7(he)-465.4(50)]TJ
/F12 1 Tf
14.3837 0 TD
0 Tc
(l)Tj
/F7 1 Tf
.5257 0 TD
-.027 Tc
[(L)-459(t)-.7(otal)-456(volume)-458(w)2.7(as)]TJ
-14.9094 -1.1869 TD
-.0291 Tc
[(pr)-5(epar)-5(ed.)]TJ
1.0514 -1.1923 TD
0 Tc
[(Fo)22.3(r)-219.8(t)0(h)22.3(e)-225(n)12.2(o)12.2(-)0(p)20.1(r)0(o)20.1(t)0(e)18(i)0(n)-210.8(c)13.4(on)19.1(tr)23.3(ol)16.8(,)-221.5(1)12.2(0)]TJ
/F12 1 Tf
11.7009 0 TD
(l)Tj
/F7 1 Tf
.542 0 TD
[(L)-220.6(o)12.2(f)-219.8(p)12.2(ro)20.1(te)18(in)-210.8(s)11(t)0(o)22.3(r)0(a)15.8(g)12.2(e)-219.6(b)0(u)19.1(f)13.2(-)]TJ
-13.2943 -1.1923 TD
[(fe)15.8(r)-263.2(w)13.4(as)-257.3(u)12.2(s)11(ed)-256.1(in)22.3(s)11(t)0(e)18(a)0(d)-256.1(o)12.2(f)-263.2(t)0(h)22.3(e)-268.4(C)10.2(R)10.2(I)13.2(SP)20.1(R)-266.2(n)12.2(u)12.2(c)0(l)18(e)0(a)16(s)11(e)-263(p)12.2(ro)20.1(te)18(in)22.3(;)-266.4(a)13.4(nd)]TJ
0 -1.1869 TD
[(fo)20.1(r)-241.5(t)0(h)22.3(e)-241.3(n)12.2(o)0(-)20.1(g)12.2(ui)16.8(d)12.2(e)-241.3(co)20.3(n)12.2(t)0(r)17.8(o)12.2(l,)-227.7(an)-229(eq)20.3(ua)20.3(l)-239.3(v)0(o)19.1(l)0(u)22.3(m)0(e)-229.9(o)0(f)-229.2(n)12.2(uc)14.8(l)15.4(e)0(a)16(s)11(e-)21.3(fr)15.6(e)13.4(e)]TJ
0 -1.1923 TD
[(wa)16.1(te)23.5(r)-246.9(w)0(a)21.5(s)-249.1(u)12.2(s)11(ed)-234.5(to)-237.9(r)13.2(e)0(p)20.3(l)0(a)18(c)0(e)-238.7(t)0(h)22.3(e)-246.7(g)12.2(R)10.2(NA)-238.7(o)12.2(r)-246.9(o)12.2(l)0(i)20(g)12.2(o-)20.1(b)12.2(a)0(s)19.1(e)0(d)-239.9(i)0(n)22.3(h)12.2(ib)22.3(-)]TJ
T*
[(it)20(or)20.1(s)11(.)-357(Th)19.2(e)-355.1(5)12.2(0)]TJ
/F12 1 Tf
5.3763 0 TD
(l)Tj
/F7 1 Tf
.542 0 TD
[(L)-356.1(i)0(T)22.4(O)0(P)-345(m)0(i)16(x)12.2(tu)22.3(re)-347.3(wa)21.5(s)-357.5(a)0(d)20.3(d)12.2(ed)-348.3(o)12.2(n)12.2(t)0(o)-346.3(t)0(h)22.3(e)-355.1(ce)16(ll)20(s)]TJ
-5.9182 -1.1869 TD
[(im)16(m)11.4(e)13.4(di)16.8(a)13.4(t)0(e)18(l)0(y)-302.9(a)0(f)21.3(t)0(e)18(r)-317.4(th)22.3(e)-311.7(c)0(u)20.3(l)0(t)20(u)0(r)20.1(e)-317.2(m)11.4(e)0(d)20.2(i)0(u)22.3(m)-313.8(wa)21.5(s)-314.1(r)0(e)15.8(m)11.4(o)12.2(v)12.2(e)0(d)20.3(.)-313.6(Th)19.2(e)]TJ
0 -1.1923 TD
[(pl)16.8(a)13.4(t)0(e)-399.3(t)0(h)22.3(e)0(n)-397.1(w)0(a)16.1(s)-406.3(in)22.3(cu)20.3(ba)20.3(te)18(d)-405.1(i)0(n)-395.1(a)-409.3(ce)16(ll)-391.8(cu)14.8(lt)20(u)12.2(r)13.2(e)-409.3(i)0(n)22.3(c)0(u)20.2(b)0(a)20.3(t)0(o)16.8(r)-404.1(fo)20.1(r)]TJ
T*
[(45)-149(m)11.4(i)0(n)22.3(,)-237.8(a)13.4(f)0(t)17.8(e)0(r)-228(w)13.5(hi)16.8(c)13.4(h)-237.1(th)22.3(e)-235.9(i)0(T)17(O)13.5(P)-239.3(m)11.4(ix)22.3(tu)22.3(re)-228(wa)21.5(s)-238.3(g)12.2(en)20.3(tl)20(y)-237.1(r)0(e)21.3(m)11.4(ov)19.1(ed)]TJ
0 -1.1869 TD
-.0203 Tc
[(a)-20.3(n)0(d)-311.6(e)-20.3(xc)-20.3(ha)-20.3(ng)-8.1(e)-20.3(d)-303.5(f)-7.1(o)-20.3(r)-303.7(2)-8.1(5)-8.1(0)]TJ
/F12 1 Tf
9.2838 0 TD
0 Tc
(l)Tj
/F7 1 Tf
.542 0 TD
[(L)-291.1(o)12.2(f)-290.3(r)0(e)15.8(g)12.2(u)12.2(l)0(a)18(r)-290.3(cu)20.3(lt)20(u)12.2(r)0(e)-282.2(m)11.4(ed)20.3(iu)22.3(m.)]TJ
/F10 1 Tf
/Cs8 cs 1 scn
-9.8258 -2.3684 TD
[(Isolation)-303.8(of)-300.7(genomic)-296.9(DNA)-300.6(in)-297.5(96-well)-296.9(cell)-302.2(culture)]TJ
0 -1.2411 TD
(plates)Tj
/F7 1 Tf
0 0 0 rg
0 -1.2032 TD
[(Genomic)-276.9(DNA)-279.4(of)-278(each)-278.1(transduced)-277.5(cell)-278.3(sample)-280.7(was)-276.4(puri-)]TJ
0 -1.1869 TD
[(ed)-582.1(by)-582.5(direct)-583.4(in-plate)-582.8(cell)-581.8(lysis)-579.7(and)-579.9(DNA)-582.9(isolation)]TJ
0 -1.1923 TD
[(according)-364.4(to)-368(a)-365.9(previously)-363.4(published)-360.4(protocol.)]TJ
7.8454 0 0 6.9729 255.5149 328.7621 Tm
(59)Tj
10.4607 0 0 10.4607 267.1936 324.1133 Tm
(Briey,)Tj
-19.8304 -1.1923 TD
[(after)-641.2(aspirating)-640.8(culture)-641.6(media)-638.6(and)-639.5(adding)-637.6(50)]TJ
/F12 1 Tf
20.2206 0 TD
(l)Tj
/F7 1 Tf
.5528 0 TD
.6379 Tc
[(Lo)637.9(f)]TJ
-20.7734 -1.1869 TD
0 Tc
[(lysis)-260(buffer)-258.4(\(10)-168.2(mM)-258(Tris)-256.8(HCl)-264.4(pH)-256.1(7.6,)-255.9(10)-170.6(mM)-258(ethylenedi-)]TJ
0 -1.1923 TD
[(aminetetraacetic)-650.9(acid)-644.6(\(EDTA\),)-647.3(100)-163.8(mM)-648.2(NaCl,)-645.9(0.5%)]TJ
T*
[(N-lauroyl)-513.1(sarcosine)-513.1(sodium,)-510(50)]TJ
/F12 1 Tf
13.5165 0 TD
(l)Tj
/F7 1 Tf
.5528 0 TD
[(g/mL)-512.1(RNase)-512.9(A,)-511.5(and)]TJ
-14.0693 -1.1869 TD
(100)Tj
/F12 1 Tf
1.6692 0 TD
(l)Tj
/F7 1 Tf
.5474 0 TD
[(g/mL)-376.6(proteinase)-371.8(K\))-379.8(to)-373.4(each)-375.6(well,)-375.1(the)-376.2(sealed)-376.2(plate)]TJ
-2.2166 -1.1923 TD
[(was)-368.5(incubated)-367.6(at)-366.8(55)]TJ
/F11 1 Tf
8.2812 0 TD
()Tj
/F7 1 Tf
.3956 0 TD
[(C)-363.7(overnight;)-364.4(added)-364.6(100)]TJ
/F12 1 Tf
9.9396 0 TD
(l)Tj
/F7 1 Tf
.5528 0 TD
[(L)-367(o)0(f)-364.7(ice-)]TJ
-19.1692 -1.1923 TD
[(cold)-336.9(NaCl-saturated)-334.5(ethanol)-339.1(\(for)-332.8(100)-163.8(mL)-333.8(100%)-333.4(ethanol,)]TJ
0 -1.1869 TD
[(add)-525.7(1.5)-169.9(mL)-523.5(of)-527.3(5)-166.6(M)-524.1(NaCl\))-524.9(to)-530.5(each)-527.4(well;)-522.9(allowed)-527.6(the)]TJ
0 -1.1923 TD
[(plate)-293.1(to)-292.1(stand)-287.9(still)-290.3(for)-291.9(4)-166.6(h)-291.3(at)-290.9(room)-286.9(temperature)-292(to)-292.1(precip-)]TJ
T*
[(itate)-344.1(the)-338.3(DNA;)-339.8(washed)-341.5(the)-338.3(precipitated)-344(DNA)-339(with)-339.1(75%)]TJ
0 -1.1869 TD
[(ethanol)-349.9(for)-346.1(two)-343.7(times)-345.1(and)-346.9(let)-351.3(the)-343.7(plate)-347.3(air)-348.1(dry;)-342.5(DNA)-349.8(in)]TJ
0 -1.1923 TD
[(each)-305.2(well)-299.9(was)-298.1(dissolved)-296.5(in)-302.9(50)]TJ
/F12 1 Tf
12.3459 0 TD
(l)Tj
/F7 1 Tf
.5528 0 TD
[(L)-296.5(o)0(f)-299.7(Tris-EDTA)-300.1(buffer.)]TJ
/F10 1 Tf
/Cs8 cs 1 scn
-12.8987 -2.3196 TD
[(Deep-sequencing)-297(preparations)]TJ
/F7 1 Tf
0 0 0 rg
0 -1.2032 TD
[(The)-363(regions)-365.4(of)-364.7(interest)-361.8(\(Supplementary)-361.6(Table)-361.2(S1\))-365.5(were)]TJ
0 -1.1923 TD
[(polymerase)-262(chain)-263.8(reaction)-265(\(PCR\))-262.8(amplied)-264.8(\(Supplemen-)]TJ
0 -1.1869 TD
[(tary)-390(Ta)20.4(bl)16.8(e)-376.8(S)0(2)22.3(\))-377(fr)15.6(o)12.2(m)-378.8(th)22.3(e)-382.2(e)13.4(xt)16.8(r)13.2(a)0(c)16(t)0(e)23.5(d)-378(DN)16.1(A)-376.8(u)12.2(s)11(i)0(n)16.8(g)-378(Q)13.5(5)-378(hi)16.8(g)12.2(h)12.2(-)]TJ
0 -1.1923 TD
[(d)16.9(e)13.4(l)0(i)20(t)0(y)-297.5(D)13.5(NA)-298.3(p)12.2(o)12.2(l)0(y)16.8(m)11.4(er)21.3(as)19.1(e)-306.3(\()0(N)21.3(e)0(w)-298.3(E)12.4(n)12.2(g)0(l)16.8(a)13.4(nd)-295.3(B)10.2(i)0(o)22.3(l)0(a)18(b)12.2(s\))18.9(.)-308.2(O)13.4(f)-306.5(t)0(h)22.3(e)]TJ
T*
[(pr)20.1(im)21.4(er)15.8(s)-314.1(u)12.2(s)11(e)0(d)-304.9(i)0(n)-302.9(t)0(h)22.3(e)-317.2(PC)20.3(R)10.2(s)-314.1(\()13.2(S)0(u)22.3(p)0(p)19.1(l)0(e)18(m)11.4(en)20.3(ta)18(r)13.2(y)-312.9(Ta)20.4(bl)16.8(e)-311.7(S)0(3)22.3(\))0(,)-305.8(t)0(h)22.3(e)]TJ
0 -1.1869 TD
[(fo)20.1(rw)15.9(a)13.4(r)0(d)-370.2(p)12.2(ri)17.8(m)11.4(e)0(r)21.3(s)-379.2(co)20.3(n)12.2(t)0(a)18(i)0(n)22.3(e)0(d)-370(5)12.2(nt)-368(s)11(e)0(q)20.3(u)0(e)20.3(n)0(c)20.3(i)0(n)16.8(g)-378(b)12.2(a)0(r)21.3(c)0(o)20.3(d)0(e)20.3(s)0(.)-367.6(A)0(l)18.1(l)]TJ
0 -1.1923 TD
-.0231 Tc
[(5)-23.1(7)-4(6)-428.2(a)-4.2(mplications)-375.2(were)-378.2(veried)-374(u)0(sing)-379.4(20)-162.6(g/L)-373.8(a)-4.2(garose)-378.2(gel)]TJ
24.0035 60.7159 TD
[(electrophoresis.)-363.8(E).2(qual)-365.4(volumes)-364.3(of)-367.6(the)-367.4(P)3.2(CR)-365.1(products)-364.3(were)]TJ
0 -1.1923 TD
-.0236 Tc
[(pooled)-314.9(with)-314.9(samples)-310.6(f)-4.9(rom)-310.3(t)-2.7(h)4.9(e)-313.7(same)-313.7(locus,)-315.6(bu)4.9(t)-311.7(w)-4.7(ith)-309.4(unique)]TJ
T*
-.023 Tc
[(barco)5.5(d).1(es.)-179.5(T).3(hese)-177.6(36)-173.4(pools)-174.5(were)-177.6(t)3.3(hen)-178.8(puried)-178.8(\(Zymo)-173.3(R)-1.9(esearch)]TJ
0 -1.1869 TD
-.0227 Tc
[(Z4004)5.8(\))-443(a)1.6(nd)-438.6(quantied)-444(u)5.8(s)-.8(ing)-438.6(Q)1.6(ubit)-435.4(d).4(sDNA)-437.4(BR)-440.6(\(Thermo)]TJ
0 -1.1923 TD
-.0233 Tc
[(Fisher)-243.1(Scientic\).)-244.8(Equal)-246.3(a)1(moun)5.2(ts)-245.3(of)-243.1(DNA)-248.3(from)-244.9(t)3(he)-248.3(samples)]TJ
T*
-.0231 Tc
[(were)-367.4(t)-2.2(hen)-368.6(pooled)-368.6(further)-373(t)3.2(o)-374(p)5.4(rov)5.4(i)-2.2(de)-372.8(six)-368.6(pools)-369.7(w)1.2(here)-372.8(each)]TJ
0 -1.1869 TD
[(sample)-161.5(contains)-158.5(a)-161.5(uniq)5.3(u)-.1(e)-161.5(c)-4.4(ombination)-162.7(of)-161.7(sequencing)-157.3(b)-.1(arcode)]TJ
0 -1.1923 TD
-.0233 Tc
[(and)-217(a)1(mplied)-217(r).8(egion.)-217.7(These)-215.8(s)-1.4(amples)-218.2(were)-221.2(t)3(he)-4.5(n)-217(s)4(e)-4.5(n)-.2(t)-213.8(t)-2.4(o)-217(B)3.2(a)-4.4(s)-1.4(e-)]TJ
T*
-.0227 Tc
[(Clear)-177.5(B)3.8(.V.)-179.2(f)1.4(or)-177.5(quality)-173.1(co)5.8(ntrol,)-173.8(index)-178.5(P)3.6(CR,)-179.2(a)1.6(nd)-178.5(sequencing)-173.1(o).4(n)]TJ
0 -1.1869 TD
-.0232 Tc
[(the)-264.5(N)1.1(ovaSeq)-260.3(600)5.3(0)-260.3(f)-4.5(o)5.3(r)-264.7(paired-end)-260.3(150nt-long)-260.3(reads.)]TJ
1.046 -1.1923 TD
0 Tc
[(The)-650.2(raw)-648.2(sequencing)-647(results)-650.6(are)-648.2(available)-650.5(on)-653(the)]TJ
-1.046 -1.1923 TD
[(NCBI)-955.5(Sequence)-956.3(Read)-958.7(Archive)-952.5(under)-955.5(BioProject)]TJ
0 -1.1869 TD
(PRJNA796802.)Tj
/F10 1 Tf
/Cs8 cs 1 scn
0 -2.3684 TD
[(Analysis)-296.4(of)-300.7(deep-sequencing)-298.9(results)]TJ
/F7 1 Tf
0 0 0 rg
0 -1.2032 TD
[(To)-544.4(analyze)-545.9(the)-544.2(deep-sequencing)-544.5(data,)-546.3(the)-544.2(paired-end)]TJ
0 -1.1923 TD
[(reads)-462.3(were)-466.7(programmatically)-463.6(merged)-465.9(using)]TJ
/F8 1 Tf
18.6922 0 TD
(seqprep.)Tj
/F7 1 Tf
7.8454 0 0 6.9729 542.0975 503.603 Tm
(60)Tj
10.4607 0 0 10.4607 310.8472 486.4818 Tm
[(Then,)-366.3(the)-365.4(reads)-364.7(were)-369.2(ltered)-367.4(out)-366.6(that)-366.2(do)-365.7(not)-366.6(match)-367.6(the)]TJ
0 -1.1869 TD
[(expected)-338.5(pattern)-332.7(of)-337.6(starting)-341.3(with)-333.6(a)-338.8(barcode)-337.9(followed)-336.7(by)]TJ
0 -1.1923 TD
[(a)-458.1(forward)-461.7(primer)-457.5(annealing)-459.7(part,)-459.8(and)-460.7(ending)-458.7(with)-458.3(the)]TJ
T*
[(associated)-372.8(reverse)-370.6(primer)-370.8(annealing)-373(part.)-373.1(We)-372.8(then)-369.4(split)]TJ
0 -1.1869 TD
[(th)22.3(e)-349.7(r)0(e)21.3(a)0(d)20.3(s)-352.1(b)12.2(y)-350.9(th)22.3(ei)18(r)-344.5(b)0(a)20.2(r)0(c)15.8(o)12.2(d)12.2(e)0(s)-338.6(a)0(n)20.3(d)-350.9(m)11.4(a)0(p)20.3(p)0(e)20.2(d)-350.9(th)22.3(e)-349.7(r)0(e)21.3(a)0(d)20.3(s)-352.1(to)-335.4(th)16.8(e)]TJ
0 -1.1923 TD
[(h)12.2(u)12.2(m)0(a)19.4(n)-546(ge)20.3(no)19.1(m)11.4(e)-544.8(\(h)20.1(g3)19.1(8)12.2(\))-550.4(u)12.2(s)11(in)22.3(g)]TJ
/F8 1 Tf
12.7307 0 TD
[(bo)19.1(w)10.2(t)0(i)20(e)0(2)]TJ
/F7 1 Tf
3.1054 0 TD
(.)Tj
7.8454 0 0 6.9729 478.9983 428.8818 Tm
[(61)24.5(,6)27.4(2)]TJ
10.4607 0 0 10.4607 501.9589 424.2329 Tm
[(T)12.4(h)0(e)-538(r)0(e)21.3(s)11.1(ul)16.8(t-)]TJ
-18.2695 -1.1923 TD
[(in)22.3(g)-551.4(a)0(l)18(i)0(g)22.3(n)0(m)18.2(e)13.4(nt)16.8(s)-552.6(w)0(e)21.5(r)0(e)-542.4(t)0(h)16.8(e)13.4(n,)-545.3(u)12.2(s)11(in)16.8(g)]TJ
/F8 1 Tf
14.3132 0 TD
-.0119 Tc
[(sam)-11.9(t)6.2(oo)-11.9(l)4.9(s)]TJ
/F7 1 Tf
3.4686 0 TD
0 Tc
(,)Tj
7.8454 0 0 6.9729 499.3511 416.4093 Tm
-.0159 Tc
[(63)-15.9(,)11.5(6)-15.9(4)]TJ
10.4607 0 0 10.4607 522.3684 411.7605 Tm
0 Tc
[(s)11.1(o)12.2(rt)17.8(ed)20.3(,)]TJ
-20.2206 -1.1869 TD
[(in)22.3(d)12.2(e)0(x)20.3(e)0(d)20.3(,)-481.6(an)20.3(d)-475.5(s)0(p)17.9(l)0(i)20(t)-477.7(b)12.2(y)-480.9(th)22.3(e)-479.8(l)0(o)16.8(c)13.4(us)-469.9(o)12.2(f)-479.9(in)22.3(te)18(re)21.3(st)-472.1(w)13.4(h)0(e)20.2(r)0(e)-471.9(t)0(h)22.3(e)0(y)]TJ
0 -1.1923 TD
[(a)13.4(l)0(i)20(g)0(n)-371.2(t)0(o)-368(t)0(h)22.3(e)-382.2(h)12.2(u)0(m)18.2(a)0(n)-370(g)12.2(en)20.3(o)12.2(m)0(e)19.4(.)-378.7(A)-382.2(s)11(ep)20.2(ar)21.3(at)18(e)-382.2(s)11(cr)21.3(ip)16.8(t)-374.8(w)0(a)16(s)-379.2(u)12.2(s)11(ed)]TJ
0 -1.1869 TD
[(to)-498(sc)19.1(or)20.1(e)-512.3(t)0(h)16.8(e)-512.3(am)19.4(o)12.2(u)12.2(n)0(t)-503.4(o)12.2(f)-512.5(r)0(e)15.8(a)13.4(ds)-502.4(wi)18.1(th)-503.4(in)22.3(s)11(e)0(r)21.3(t)0(i)20(o)0(n)19.1(s)-514.7(o)12.2(r)-512.5(d)12.2(e)0(l)18(e)0(-)]TJ
0 -1.1923 TD
-.0131 Tc
[(t)-13.1(i)6.9(ons)-262.2(ba)-13.1(s)6(e)-13.1(d)-253(o)-.9(n)-261(t)-13.1(h)9.2(e)-259.8(c)-13.1(o)7.1(nc)-13.1(i)4.9(s)-2.1(e)-259.8(i)-13.1(di)-13.1(o)9.2(s)-2.1(yn)-13.1(c)7.1(r)-13.1(at)-13.1(i)6.9(c)-259.8(ga)-13.1(p)7.2(p)-13.1(e)7.2(d)-261(a)-13.1(l)4.9(i)-13.1(g)9.2(nme)-13.1(n)7.2(t)]TJ
T*
0 Tc
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T*
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T*
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T*
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T*
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[(t)-6.2(hese)-481(statis)-10.3(tics,)-485.9(we)-482.6(used)]TJ
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[(tides)-582.7(could)-579.3(work)-577.5(as)-582.5(guide-specic)-580(inhibitors)-580.4(of)-576.1(Cas)]TJ
T*
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[(nucleotides)-314(\(nt\))-314.4(\(Fig.)-316.9(1\).)-309.8(The)-314.2(inhibitors)-314.8(are)-312.2(complemen-)]TJ
T*
[(tary)-487.6(to)-487.2(the)-490(PAM-proximal,)-483.9(spacer-derived)-484.2(part)-487.6(of)-484(the)]TJ
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[(itself,)-300(creating)-303(a)-300.9(double-stranded)-298.8(PAM)-294.6(\(Fig.)-300.6(1D\).)]TJ
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T*
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[(itors)-404.1(to)-400.5(CML)-398.5(cells)-402.7(using)-402.9(iTOP.)]TJ
7.8454 0 0 6.9729 450.6518 192.3023 Tm
(58)Tj
10.4607 0 0 10.4607 462.6707 187.6535 Tm
[(We)-405.3(analyzed)-398.1(editing)]TJ
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T*
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T*
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(73,74)Tj
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(574)Tj
41.45 0 TD
[(SW)-6.6(ARTJES)-308.2(ET)-299.4(AL.)]TJ
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T*
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[(and)-569.1(50%\).)-564.8(The)-568.9(8nt)-567.1(inhibitor,)-563.6(however,)-570.2(unexpectedly)]TJ
T*
[(showed)-440.2(increased)-445.7(indel)-440.6(frequencies)-441.9(rather)-439.2(than)-439.8(inhibi-)]TJ
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[(scrambled)-203.8(versions)-202.6(of)-196.7(the)-202.8(inhibitors,)-200.3(which)-198.7(were)-201.2(included)]TJ
T*
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[(tions)-340.1(and)-336(lengths)-336.1(of)-337.6(the)-338.3(indels)-337.5(\(Supplementary)-334.5(Figs.)-335.3(S1)]TJ
T*
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[(To)-403.5(characterize)-405.4(the)-403.3(observed)-398.5(inhibition)-398.1(in)-405.9(more)-399.5(detail,)]TJ
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[(we)-271.2(tested)-276.7(the)-267.8(inhibitors)-271.4(in)-270.4(a)-268.4(concentration)-275.3(gradient.)-265.5(Sub-)]TJ
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[(stantial)-202.4(inhibition)-197.6(by)-203.2(the)-197.3(20nt)-197.1(inhibitor)-201.2(is)-201.1(only)-197.1(observed)-203.4(at)]TJ
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[(and)-227.6(above)-229.1(the)-224.4(equimolar)-227.7(ratio)-228.3(to)-227(the)-224.4(Cas9)-229.4(RNP)-226.4(\(Fig.)-224.7(2B\).)]TJ
T*
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[(PAM)-619.8(inhibitors,)-612.2(a)-620.7(more)-616.3(gradual)-617.7(dose)]TJ
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[(response)-323.8(is)-320.4(observed)-317.2(\(Fig.)-316.9(3A\),)-323.5(with)-317.4(low)-322(concentrations)]TJ
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[(seemingly)-354.6(providing)-343.1(low)-354.5(levels)-347.1(of)-353.9(inhibition.)-348.7(However,)]TJ
T*
[(even)-360.5(at)-361.3(the)-359.9(highest)-357.7(concentration,)-355.9(the)-359.9(8nt)]TJ
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[(PAM)-354.3(inhibi-)]TJ
6.3301 32.1437 TD
[(tors)-560.4(are)-566.9(not)-567.1(able)-563.3(to)-563.1(reduce)-566.9(indel)-565.3(frequencies)-561.1(to)-568.5(the)]TJ
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[(same)-438.4(extent)-438(as)-441.6(was)-439(achieved)-436.1(with)-442(the)-435.8(20nt)-441(inhibitors.)]TJ
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[(At)-323.4(the)-327.4(investigated)-325.7(off-target)-325.4(loci,)-327.5(we)-325.4(observed)-328(reduced)]TJ
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[(indel)-473.2(frequencies)-479.8(with)-474.5(the)-473.8(20nt)-479(and)-471.5(8nt)]TJ
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[(PAM)-478.9(inhibi-)]TJ
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[(tors,)-570.6(following)-566.3(a)-571.9(similar)-565.7(doseresponse)-571.5(as)-571.7(was)-569.1(seen)]TJ
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[(The)-352.1(8nt)]TJ
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[(PAM)-348.9(oligo)-355.1(displays)-346.7(inhibition)-349.4(both)-354.3(on-)-347.1(and)]TJ
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[(off-target,)-210.9(but)-209.4(neither)-208(is)-212(completely)-206.7(abolished.)-212.1(This)-209(raises)]TJ
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[(the)-235.3(question)-237.1(to)-232.5(what)-238.1(extent)-237.5(the)-235.3(ratio)-239.1(between)-236.9(on-)-233.2(and)-238.5(off-)]TJ
T*
[(targeting)-391.7(is)-390.8(affected)-396.1(by)-392.8(the)-392.5(inhibitors.)-390(We)-394.5(calculated)-398.6(a)]TJ
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[(specicity)-426(score)-424.4(by)-425.4(dividing)-423.6(the)-425(on-target)-426.4(indel)-424.4(frequ-)]TJ
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[(ency)-300.9(by)-306.1(the)-305.7(sum)-301.3(of)-305.1(the)-305.7(on-)-298.3(and)-303.5(off-target)-309.2(indel)-299.7(frequen-)]TJ
T*
[(cies.)-471.6(This)-469.1(specicity)-469.4(score)-467.7(signies)-473.5(the)-468.3(percentage)-469(of)]TJ
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[(indels)-261.6(that)-263.2(were)-266.2(mapped)-261(to)-265(the)-262.4(on-target)-263.8(locus,)-260.1(corrected)]TJ
0 -1.1923 TD
[(for)-497.8(the)-500.8(total)-502.5(number)-499.7(of)-500.2(reads)-500.2(mapped)-499.5(to)-503.4(each)-500.3(locus.)]TJ
T*
-.0128 Tc
[(We)-313.7(found)-309.5(t)-12.8(h)9.5(at)-311.7(the)-313.7(h)-.6(i)-12.8(g)9.5(h)4.9(e)-12.8(r)-300.5(c)-12.8(o)7.5(n)4.9(c)-12.8(e)8.6(nt)-12.8(r)10.5(a).6(t)-12.8(i)7.2(ons)-310.7(o)4.9(f)-313.9(t)-12.8(h)9.5(e)-308.3(8nt)]TJ
/F11 1 Tf
20.1881 0 TD
0 Tc
(+)Tj
/F7 1 Tf
.5365 0 TD
[(PA)23.5(M)]TJ
-20.7246 -1.1869 TD
[(i)15.4(n)12.2(h)12.2(i)0(b)22.3(i)0(t)25.5(o)12.2(r)-360.7(s)11(l)15.4(i)0(g)22.3(h)12.2(t)15.4(l)0(y)-346.3(i)0(n)22.3(c)13.4(r)13.2(e)0(a)21.5(s)11(e)-360.5(t)15.4(h)12.2(e)-360.5(s)11(p)12.2(e)13.4(c)13.4(i)0()20.1(c)13.4(it)25.5(y)-356.3(s)11(co)20.3(r)13.2(e)-360.5(\()13.2(F)15.5(ig)22.3(.)-357(3)12.2(B)10.2(\))13.2(.)]TJ
0 -1.1923 TD
[(However,)-243.8(it)-240.1(is)-244.5(important)-242(to)-243.3(realize)-243.6(that)-241.5(at)-242.1(the)-246.1(same)-243.3(inhib-)]TJ
T*
[(itor)-594(concentrations,)-599.6(on-target)-599.8(activity)-596.9(is)-596.8(substantially)]TJ
0 -1.1869 TD
[(reduced)-299.9(\(Fig.)-295.2(3B,)-302(C\).)]TJ
1.046 -1.1923 TD
[(For)-251.7(the)-251.5(20nt)-245.9(inhibitor,)-249.3(we)-254.9(observed)-246.7(increased)-250.6(specic-)]TJ
-1.046 -1.1923 TD
[(ity)-217(only)-218.8(in)-216.2(conditions)-220.3(in)-216.2(which)-220.4(barely)-218(any)-216.8(on-target)-220.5(activ-)]TJ
0 -1.1869 TD
[(ity)-222.4(is)-228.2(left)-224.3(\(Supplementary)-226.1(Fig.)-221.7(S3\).)-229.3(We)-226.5(also)-224.2(tested)-227.9(variant)]TJ
0 -1.1923 TD
[(20nt)-245.9(inhibitors)-244.3(\()48.2(off120nt)49.7()-246.9(and)-249.3()51.2(off220nt)49.7(\))-249.9(based)-246.5(on)]TJ
T*
[(the)-690.5(off-target)-694(sequences)-689.5(\(Supplementary)-692.2(Table)-691.8(S5\).)]TJ
0 -1.1869 TD
[(These)-300.5(off-target)-298.4(20nt)-300.1(inhibitors)-298.5(did)-301.5(not)-301.5(show)-297.9(substantial)]TJ
0 -1.1923 TD
[(differences)-381.2(in)-384.2(specicity)-388.1(or)-381(activity)-385.5(compared)-380.6(with)-387.8(the)]TJ
T*
[(on-target)-366.8(20nt)-370.6(inhibitors)-369(\(Supplementary)-367(Fig.)-368(S3\).)-370.2(This)]TJ
ET
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h
20.651 .963 m
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h
25.558 1.121 m
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h
22.881 1.619 m
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24.729 2.747 24.918 2.06 24.91 1.619 c
22.881 1.619 l
f
Q
BT
/F14 1 Tf
11.9999 0 0 11.9999 118.885 693.5251 Tm
13.543 Tc
(AB)Tj
0 -8.9677 TD
(CD)Tj
ET
EMC
EMC
/GS1 gs
BT
/F5 1 Tf
9.9626 0 0 9.9626 59.7543 481.7762 Tm
0 0 0 rg
0 Tc
[(FI)-7(G.)-330.6(1.)]TJ
/F10 1 Tf
3.8753 0 TD
[(Design)-302.7(of)-306.1(guide-c)-8.1(ompleme)-8.1(nta)-7(ry)-302.6(oligo-b)-8.7(ased)-300.4(inhi)-8.6(bitors.)-305.7(Sch)-8.6(ematic)-308.4(representati)-14(on)-301.2(of)-300.4(a)-300.6(C)0(a)-7.4(s)0(9)-301.4(nucleas)-9.5(e,)-298.8(gRN)-8(A)]TJ
-3.8753 -1.2974 TD
[(\(here)-309(displayed)-310.3(as)-303.9(a)-300.6(sgRNA\))-304.2(with)-307.6(and)-301.9(withou)-10.9(t)-298.3(olig)-8.2(o-based)-306.6(inhibi)-8.5(tors.)]TJ
/F5 1 Tf
29.2722 0 TD
(\(A\))Tj
/F10 1 Tf
1.5706 0 TD
[(Witho)-11.1(ut)-303.6(inhibitors)-10.5(,)-298.1(the)-304.7(RNP)-307.8(binds)-306.2(the)]TJ
-30.8428 -1.3031 TD
[(ta)-7.1(rget)-301.4(sequen)-11.5(ce)-299.2(in)-305.9(genomic)-306.6(DNA,)-305.5(forms)-309.6(an)-300.6(R-loop)-8.4(,)-298.1(and)-307.6(proceeds)-308(to)-303.6(cl)-6.9(eave)-301.1(both)-304.9(strands)-313.3(of)-300.4(the)-304.7(DNA)-303.3(\(triangl)-10.1(es\).)]TJ
/F5 1 Tf
0 -1.2974 TD
(\(B\))Tj
/F10 1 Tf
1.5251 0 TD
[(20nt)-302.5(inhibi)-8.5(tor.)]TJ
/F5 1 Tf
6.2425 0 TD
(\(C\))Tj
/F10 1 Tf
1.5251 0 TD
[(8nt)-300.4(inhibi)-8.6(tor.)]TJ
/F5 1 Tf
5.7304 0 TD
(\(D\))Tj
/F10 1 Tf
1.6161 0 TD
(8nt)Tj
/F11 1 Tf
1.3885 0 TD
(+)Tj
/F10 1 Tf
.552 0 TD
[(PAM)-305.1(inhibi)-8.6(tor.)-302.4(For)-308.3(all)-297.8(nuc)-7.8(leic)-302.2(acids,)-308.6(5)]TJ
/F11 1 Tf
15.558 0 TD
()Tj
/F10 1 Tf
.552 0 TD
[(ends)-301.3(ar)-7.1(e)-296.7(indicate)-8.9(d.)-305.1(gRNA)-7.2(,)-298.1(guide)]TJ
-34.6896 -1.3031 TD
[(RNA)-7.2(;)-298.1(PAM)-9.2(,)-298.1(proto)-11(spacer)-305.3(adjace)-8.4(nt)-303.9(motif;)-307.1(RNP,)-304.3(ribonuc)-9.7(leopr)-8.4(otein;)-305.1(sgR)-7.2(NA,)-301.4(sing)-7.6(le)-301(gRNA.)]TJ
/F5 1 Tf
0 32.5101 TD
[(MO)-8.2(DULA)-7.6(TING)-300.8(CA)-8.2(S9)-299.8(ACTIVI)-11.5(TY)-300(WITH)-309(DNA)-302.7(OLIG)-9.2(OS)-25472.4(575)]TJ
ET
endstream
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