^jɂFDNA̕Jn̋@\

<strong>r OV</strong>
i`w `j
emailF<a href="mailto:hiaraki@nig.ac.jp">r؍OV</a>
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̈Zr[, <strong>5</strong>, e006 (2016)@<a href="http://dx.doi.org/10.7875/leading.author.5.e006">DOI: 10.7875/leading.author.5.e006</a>
Hiroyuki Araki: <strong>Molecular mechanism of the initiation of chromosomal DNA replication in eukaryotes.</strong>

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<h2>v </h2>
@^jɂFDNÄ͓̗̕悩JnD̕Jn_ɂOrcCDNAwJ[[̐G}RAłMcm2-7փeZʑ̂ƂG1Ƀ[hDJn̉ߒזEɂ萧䂳邱ƂɂDNAƍזE͋邪Cɂ͍זE̐̌ƂȂTCNˑLi[[Dbf4ˑLi[[ɂ镡^pÑ_dvȖSDȂ킿C烊_ɂCdc45GINŜƂCMĜJn_Ɍ`CɂĊDNAwJ[[ƂĂ͂炭悤ɂȂ邱ƂJn̐ɂėvƂȂDāCDNA|[[N[gDNAJnD_ɂďڍׂ͕sCMcm2-7փeZʑ̂CMĜ̍\CāC<em>in vitro</em>ɂč\zɂCCMĜ̌`̋@\ސł悤ɂȂĂĂCJn̋@\ɂđSẻ𖾂߂ÂĂD

<h2>͂߂</h2>
@ۂ𕪎qx̋@\ƂĒm肽ƂCǂ̂悤Ɍi߂邾낤H@NbVbNɂ́C1jɊ֗^^pN݂C2j̃^pNψق킦肷邱Ƃɂω甽@\𐄎@f\zDɁC3jP^pNgݍ킹邱Ƃɂ蔽<em>in vitro</em>ɂčč\z邱Ƃł΁Cf̏ؖƓɂɏڍׂȋ@\m邱ƂłD^jɂDNA̋@\̌́CECXp<em>in vitro</em> DNAn\zC3j̒iKɂ܂œBDCECX͏hזE̕n̂̂ł͂ȂECXƎ̋@\߁CזÊȂŋN镡mɂ͕s\łD܂CAtJcKG̗otpn\z͂i񂾂C2j̒iKłƂ܂ĂDCVvȒPזE̐^jłyfƂɂẮC`wIȎ@ɂ1j2j̒iǨi݁C݁C3j̒iKɐi݂D
@ŁC^pN̍\͂̋Zp̐iW߂܂DXɂ錋͂NMRɂntɂ\͂ɂ킦CߔNCdqpPq͂̋Zp}ɐi݋ȃ^pN̍\qxŌ肳悤ɂȂĂD̍\ƁCɏqׂ͂gݍ킹邱ƂɂC^jɂ镪qxł̕@\̗́CIɂ͂ċ}ɐiłDł́Cŋ߂̌𒆐SɁC^jɂFDNA̕Jn̋@\D

<h2>1DJn_</h2>
@FDNA͐̕F̂ɎU݂̕ʂł镡Jn_邢͕Jn̈悩JnDJn_͍̐yŖ1000Cqgł͐ɂȂD̕ʂK肷̂6̃TujbgȂOrcłDOrcATPƌATPasełC2{DNAƌDyꂩqg܂ł悭ۑĂCyVEWEoGOrc͓dqɂPq͂X\͂炻̍\炩ɂĂ<a href="#R1"><sup>1-3)</sup></a>DOrc̐Jn_߂邤ŏdvȖSDoyi<em>Saccharomyces cerevisiae</em>jɂĂ͓11 bp̔złACSƌ邪Cyi<em>Schizosaccharomyces pombe</em>jɂĂOrc4ATtbNƂ΂\AT𑽂܂ޗ̈ƌ<a href="#R4"><sup>4,5)</sup></a>DCqgVEWEoGOrc͓̉zFȂDāCyłDNAzɂ蕡Jn_K肳邪CזEłDNAzނN}`̏ԂȂǂɂOrčʂ܂ƍlĂ<a href="#R6"><sup>6)</sup></a>DہCOrĉƂ傫ȃTujbgłOrc1̓qXgƂ̌ʂC̕ʂƐF̂ɂOrčlω<a href="#R7"><sup>7)</sup></a>D܂COrcƐFDNAƂ̌́CoyɂĂ͍זEċN邪CyɂĂ͍זEɂ茋̏ԂςƂ̕񍐂DɁCMނɂOrc1G1猋Mɂ͕<a href="#R5"><sup>5)</sup></a>D

<h2>2DMcm2-7փeZʑ̂̕Jn_ւ̃[h</h2>
@DNADNA|[[2{DNAɑ΂DNA邱Ƃ͂łȂDDNA̍͂߂ɂ́C^ƂȂ1{DNAуvC}[KvłD̂߁CJn_ɂ2{DNA1{DNAɂقǂȂDNA̍͊JnȂD咰ۂɂĂ͕Jn_ɕDnaAƂ̋ߖT2{DNAقǂ邪C^jɂĂOrčɂ蕡Jn_ɂ2{DNAقǂƂ񍐂͂ȂD
@^jɂĂ̓TCNˑLi[[̊̒ႢM̌G1ɂāC2{DNA1{DNAɂقǂDNAwJ[[̐G}RAƂȂMcm2-7փeZʑ̂OrčJn_Ƀ[hi<a href="#F1">}1a</a>jD̐FDNAɂOrcMcm2-7փeZʑ̂Ȃ\𕡐JnÓipre-replicative complex邢pre-replication complexFpre-RCjƂԁDJnÔ̌`ɂ́COrcMcm2-7փeZʑ̂ɂ킦Cdc6Cdt1KvłDoyɂCdt1Mcm2-7փeZʑ̂ƕ̂`ĂCCdc6OrcƂƂɂ͂炭DOrcCMcm2-7փeZʑ́CCdc6ATPaseCATP̃GlM[pMcm2-7փeZʑ̂𕡐Jn_Ƀ[hD

<a name="F1"></a><div id="fig1-caption-text" style="display: none;"><strong>}1@y̌ɂƂÂ^jɂFDNA̕Jn̋@\</strong>
iajOrčJn_ւMcm2-7փeZʑ̂̃[hD
ibjJn_ւSld3-Sld7̂Cdc45Ƃ̂̌̕D
icjJn_ɂ郍[hÔ̌`D
idjJn_ɂCMĜ̌`D
iejDNA|[[DNA|[[ÂɂDNA̐LDł́CDNA|[[Â[fBOCDNA|[[MOLƂD
PF_D
<a href="http://leading.lifesciencedb.jp/wordpress/wp-content/uploads/2016/07/Araki-5.e006-Fig.1.jpg" target="_blank">[Download]</a></div>[hs_figure id=1&amp;image=/wordpress/wp-content/uploads/2016/07/Araki-5.e006-Fig.1.png&amp;caption=fig1-caption-text]

@Mcm2-7փeZʑ̂̂ꂼ̃Tujbg͋ʂDȂ킿CN[OBtH[hуWNtBK[hCCC[AAA<sup>+</sup>t@~[ɑATPasehCʒuDJn_ɂ2qMcm2-7we6ʑ̂N[ނ悤Ƀ[h邪CdqɂPq͂Ȃǂ炨̂̂̃Tujbg̔zu񍐂Ă<a href="#R2"><sup>2)</sup></a>i<a href="#F2">}2</a>jDāCMcm2Mcm5Ƃ̂ɌԂC̕p2{DNAɃ[hƍlĂD܂CMcm2-Mcm6̑ɂԂώ@DɁC΂Mcm5Mcm7N[уWNtBK[hC̑ݍpɂ茋<a href="#R8"><sup>8)</sup></a>D

<a name="F2"></a><div id="fig2-caption-text" style="display: none;"><strong>}2@Sld3-Sld7̂̍\Mcm2-7փeZʑ̂ɑ΂zu</strong>
iajSld3-Sld7̗̂ސ\D
ibjSld3-Sld7̂2qMcm2-7փeZʑ̂ɑ΂zũfD̃fɂẮCSld3͌Mcm2-7փeZʑ̂ł͂ȂCΑMcm2-7փeZʑ̂Cdc45GINŜD
CBDFCdc45hCCPF_D
<a href="http://leading.lifesciencedb.jp/wordpress/wp-content/uploads/2016/07/Araki-5.e006-Fig.2.jpg" target="_blank">[Download]</a></div>[hs_figure id=2&amp;image=/wordpress/wp-content/uploads/2016/07/Araki-5.e006-Fig.2.png&amp;caption=fig2-caption-text]

@Mcm2-7փeZʑ̂Jn_փ[h@\́CoyɂĐ^pNpč\nɂ͂ĂDcOȂC΂Mcm2-7փeZʑ̂ǂ̂悤ɕJn_փ[h邩͂܂Sɂ͂킩ĂȂD[hꂽMcm2-7փeZʑ͕̂K΂ɂȂĂ邽߁C΂Mcm2-7փeZʑ̂ǂɃ[ĥł͂ȂƂ̐C1qώ@ȂǂɂƁC܂1qMcm2-7փeZʑ̂[hCɂ1qMcm2-7փeZʑ̂[hĂ<a href="#R9"><sup>9)</sup></a>CMcm2-7փeZʑ̂̃[hɋNĂ邩ǂ̌m@\悤ł<a href="#R10"><sup>10,11)</sup></a>DɂCȋ@\CƂɁC2qڂMcm2-7փeZʑ̂̃[hɂĂ͕sȓ_Č҂Ȃ΂ȂȂD

<h2>3DMcm2-7փeZʑ̂̂DNAwJ[[̊</h2>
@Mcm2-7փeZʑ̂DNAwJ[[SCMcm2-7փeZʑ̂ł͊ȂiɂẮC̃TujbgȂ镡̂ア̂jDDNAwJ[[̊ɂG1̌SɂĊTCNˑLi[[Dbf4ˑLi[[ɂ郊_CсCɂƂȂ̃^pN̕Jn_ւ̏WKvł邱Ƃoyp͂킩ĂDł́CMcm2-7փeZʑ̂̂DNAwJ[[̊ɂāCoy𒆐SɐD
@Dbf4ˑLi[[Cdc7G}TujbgƂCTujbgłDbf4ƕ̂`DG1̌<em>DBF4</em>`q̓]ʂCDbf4̃^pNʂ邱ƂɂDbf4ˑLi[[͊DꂽDbf4ˑLi[[Mcm2CMcm4CMcm6N[̍\ƂȂ̈ɂ镡SerThr_DƂɁCMcm6Dbf4ˑLi[[ɂ郊_Sld3-Sld7̂Cdc45Ƃ̂̕̕Jn_Ƃ̌𑣐i<a href="#R12"><sup>12)</sup></a>i<a href="#F1">}1b</a>jD
@Sld3-Sld7̂2qSld3N[̈ɂSld7ʑ̂ɂ茋ꂽłȍ\Ƃ<a href="#R13"><sup>13)</sup></a>i<a href="#F2">}2a</a>jDSld3̒ɂCdc45Ƃ̌̈悪C1qCdc451qSld3ƌ邽<a href="#R14"><sup>14)</sup></a>CSld3-Sld7̂2qCdc45ƌDזEɂSld3-Sld7̂Cdc45݂͌ɈˑĕJn_ƌ邪<a href="#R15"><sup>15,16)</sup></a>C<em>in vitro</em>ɂāCSld3-Sld7̂Cdc45ȂĂJn_ƌł邪CCdc45̕Jn_Ƃ̌ɂSld3Kvł<a href="#R12"><sup>12)</sup></a>D̂悤ɁCDbf4ˑLi[[ɂMcm6̃_ɂSld3-Sld7̂Cdc45Ƃ̂̕Mcm2-7փeZʑ̂ƌDM҂́CSld3-Sld7̂̍\炻̔zu𐄑Ăi<a href="#F2">}2b</a>jC̔zúĈɏqׂ銈^DNAwJ[[̌`ɂs悢̂ƂȂĂD
@TCNˑLi[[͏oyɂĂSld2Sld3_邱Ƃɂ蕡Jn𑣐i<a href="#R17"><sup>17)</sup></a>DSld2ɂ̓TCNˑLi[[ɂ胊_ꂤ11ӏ̕ʂ݂D̂CThr84TCNˑLi[[ɂ胊_Dpb11ƌDDpb11C[N[Ƀ^fɂȂBRCT̈C̃^fɂȂBRCT̈̓_ꂽyv`hƌ邱ƂmĂD_ꂽSld2Dpb11C[ɂ^fɂȂBRCT̈ƌDCDpb11N[ɂ^fɂȂBRCT̈̓_ꂽSld3ƌDSld3TCNˑLi[[ɂ胊_ꂤ12ӏ̕ʂCC[ɂThr600Ser622̃_Dpb11Ƃ̌Ɋ֗^Dǂ炩_Ό͉\ł邪CƂ_ƋD
@TCNˑLi[[ɂ胊_ꂽSld2Sld3Dpb11ƌ邱ƂɂCǂ̂悤ĕJnîł낤D̐̂܂ɁC^DNAwJ[[ɂĐق₷낤DMcm2-7փeZʑ̂DNAwJ[[ƂĂ̊Cɏqׂ悤ɁCMcm2-7փeZʑ̂ł͊ȂDזEɂẮCMcm2-7փeZʑ̂Cdc45GINŜCMĜƂ΂鋭łȕ̂`CCMĜɂĊDNAwJ[[ƂĂ͂炭<a href="#R18"><sup>18)</sup></a>Dɏqׂ悤ɁCCdc45Sld3-Sld7̂ƌCMcm2-7փeZʑ̂ɑ݈ˑIɃN[gDGINŜSld5CPsf1CPsf2CPsf34̃TujbgȂ镡̂ŁCM҂ɂC̐̓{ǂ݂goCichiCniiCsanƖÂꂽ̂łDSld3Cdc45̃N[gGINŜ̕Jn_ւ̃N[gɕKvłDāCM҂ɂCSld2̃TCNˑLi[[ɂ郊_GINŜ̃N[gɊ֗^Ă邱Ƃ炩ɂC̋@\ɂĂ͂ĎʁC_ꂽSld2Dpb11ƌƁCDNA|[[ÂGINŜ̂̕ɂɂ邱ƂꂽD̂̕[hÓipre-loading complexFpre-LCjƖÂC[hÔɊ܂܂Dpb11Jn_ɂă_ꂽSld3ƌ邱ƂɂCGINŜ͕Jn_ɃN[ĝƍlĂ<a href="#R19"><sup>19)</sup></a>i<a href="#F1">}1c</a>jDDNA|[[Â͍זEGINŜƈȕ̂`ĂCDNA|[[ÂGINŜDpb11ƎキݍpĎSld2ɂ艻Ă̂ł낤D
@MނɂĂDpb11ƑΉBRCT̈9TopBP1mĂCTopBP1̌^pNƂSld3ɑΉTreslin/Ticrr݂<a href="#R20"><sup>20,21)</sup></a>DAtJcKG̗otp͂ɂCTopBP1Treslin͓ƗɃN}`ƌ邪CTopBP1TreslinƂ̌̓TCNˑLi[[ɂTresliñ_ɂNCJnɕK{ł邱Ƃ킩DɁCTreslinɂMTBPCMTBPJnɕK{ł邱ƂĂ<a href="#R22"><sup>22)</sup></a>DꂪSld3ƕ̂`Sld7Ɠ͂炫Ă邩ǂ͂킩ȂDCMނɂSld2Ǝア^pNƂRecQL4mĂDRecQL4C[ɃwJ[[hCC̃hČ͑Vǂ̂ЂƂłRothmund-ThomsonǌQ̔ǂ̌ƂȂDwJ[[hC͍זȆBɂ͕K{ł͂ȂCSld2ƑN[̗͕̈K{ł<a href="#R17"><sup>17)</sup></a>DAtJcKG̗otp͂CRecQL4͕Jň㔼ɂĂ͂炢Ă悤ŁCDNA|[[̃N[gɕKvƂ̕񍐂C̉͂KvłD
@Jn_ɌMcm2-7փeZʑ́CCdc45CGINŜ͋łCMĜ`<a href="#R18"><sup>18)</sup></a>i<a href="#F1">}1c</a>jDCCMĜ̌`͂ɂ͕Jnɂ͂ȂȂD́CMcm10CMG̗l̋łȕ̂͌`邪CJn͋NȂłDMcm10͈΂CMĜɂނ2{DNAقǂ悤ɂ͂炫ƍlĂ邪i<a href="#F1">}1d</a>jCq@\͂킩ĂȂ<a href="#R23"><sup>23)</sup></a>DMcm10̋@\͕Jnɂcꂽ~Xe[łD
@łCMĜłߒ͂܂킩ȂCMcm2-7փeZʑ̂CMĜ̍\͓dqɂPq͂疾炩ɂĂ<a href="#R8"><sup>8,24,25)</sup></a>DGINŜCdc45Mcm2-7փeZʑ̂Mcm2Mcm5Ƃ̂̌Ԃ߂悤ʒui<a href="#F3">}3</a>jDɁCN[ނ悤`ꂽ΂Mcm2-7փeZʑ̂̈ʒu֌ẂCMcm2Mcm5Ƃ̂̌Ԃ͔΂̕ނĂi<a href="#F2">}2b</a>jDMcm2-7փeZʑ̂DNAɃ[hdqɂώ@ƁC2{DNAƂɂ݂DŁCCMĜɂDNAwJ[[̉͂CCMĜ1{DNÂ3f5f̕ɓĂ邱Ƃ킩i<a href="#F4">}4</a>jDȂ킿CCMĜ̓[fBO̒^DNÂ𕡐̕ɓĂ邱ƂɂȂD̂ƂCMcm2-7փeZʑ̂͂܂2{DNAOɂƂCꂽ_邢CMĜ`ꂽ_1{DNAOɂƂD̂ƂCMcm2Mcm5Ƃ̂̌Ԃ̈ʒu̕ނȂ̂́CJn_Ƀ[hꂽ΂Mcm2-7փeZʑ̂邢CMĜ2{DNA1{DNAւƏ芷̂ɂ͓s悢炩ȂD

<a name="F3"></a><div id="fig3-caption-text" style="display: none;"><strong>}3@CMĜ̍\</strong>
iaj猩ƂD
ibj֐؂ɂƂD
<a href="http://leading.lifesciencedb.jp/wordpress/wp-content/uploads/2016/07/Araki-5.e006-Fig.3.jpg" target="_blank">[Download]</a></div>[hs_figure id=3&amp;image=/wordpress/wp-content/uploads/2016/07/Araki-5.e006-Fig.3.png&amp;caption=fig3-caption-text]

<a name="F4"></a><div id="fig4-caption-text" style="display: none;"><strong>}4@tH[NɂDNA|[[̔</strong>
<a href="http://leading.lifesciencedb.jp/wordpress/wp-content/uploads/2016/07/Araki-5.e006-Fig.4.jpg" target="_blank">[Download]</a></div>[hs_figure id=4&amp;image=/wordpress/wp-content/uploads/2016/07/Araki-5.e006-Fig.4.png&amp;caption=fig4-caption-text]


<h2>4DtH[Ň`</h2>
@Jn_ɂ2{DNA1{DNAɂقǂƁCǂDNA|[[̏oԂłD^jɂFDNA̕ɂDNA|[[CDNA|[[CDNA|[[ÂKvłCDNA|[[ǂ̂悤ɕJn_ɃN[ĝڍׂ͕sł邪CDNAwJ[[CMĜ₻ɌCtf4DNA|[[Ƃ̌dvȂ͂炫邱Ƃ<a href="#R26"><sup>26)</sup></a>D
@Jn_ɂĂ܂͂炭̂DNA|[[łDDNA|[[̓vC}[RNAvC}[[Ƌłȕ̂`ĂC̃vC}[[̍RNAɂÂĐVDNADDNA|[[Ctf4iMނɂẮCAnd1jƌDCtf4DNA|[[ƌ̈ɂCMĜɊ܂܂GINŜƂDɁCCtf4̓zOʑ̂`邽߁C2qDNA|[[CMĜĕtH[NɂƂǂ߁CDNA|[[悭g悤ɂƍlĂ<a href="#R27"><sup>27)</sup></a>i<a href="#F4">}4</a>jD
@DNA|[[DNA|[[Â͂ƂDNA|[[̍DNAvC}[ƂDNAL邪CDNA|[[ÂGINŜ̃TujbgłPsf1Ƌ邽߁CGINŜCMĜƌ<a href="#R28"><sup>28)</sup></a>D܂CDNA|[[Â̓[hÔ̍\^pNƂĕJnɂ͂炫CɁCJn_ɃN[gƍlDāCDNA|[[DNA|[[ÂDNAwJ[[CMĜƓɌC[fBOуMO̍Ɋ֗^̂ƎvD
@DNA|[[DNA|[[ÂDNALۂɂ́C5̃TujbgȂNv[_[łRFCvC}[3f[ɌCDNA|[[̒^DNA̒Eh~郊ÕXCfBONvłPCNA[h<a href="#R29"><sup>29)</sup></a>i<a href="#F1">}1e</a>jDDNA|[[ǂ̂悤DNAi߂Ă邩͒c_̂킩ƂłCDNA|[[DNA|[[ÂɂrNTP荞݂₷ψّ̂Cp͂CDNA|[[MOCDNA|[[Â[fBOƂčƌ_ꂽĎ_́C܂kNI`h荞݂₷DNA|[[̕ψّ̂p܂ł̉͂ƈvDCrNTP̎荞݂̉͂ɂDNA|[[̖_͈ꌏƎvCrNTP荞DNȀCɂă[fBOƃMOƂ̂ɂ͍Cۂɂ́C[fBOуMOƂDNA|[[ɍĂƂ񍐂ȂāC̖͂܂ׂƂȂɕ荞܂Ă<a href="#R30"><sup>30)</sup></a>DDNA|[[Â[fBOCDNAwJ[[CMĜƌ悭ĂƂ̕񍐂邪CDNA|[[DNA|[[Â̖S͂肵Ȃߕ]̂킩ƂłD݂̂ƂCDNA|[[ǂ̂悤ɕJn_ɃN[ĝ͂킩ȂD
@[fBO͕tH[N̐isƓɐiݘADNA\ł邪CMO͕tH[N̐isƂ͔΂̕ɐiނߕsADNAND^jɂĂ̓MO̒kNI\[ɂK肳ĂC̓kNI\[ق1ɑ165 bpƂȂ<a href="#R31"><sup>31)</sup></a>DDNA|[[RNAvC}[p30`50kNI`hDNA邪CMOłDNA|[[DNA̍͂߂̂ŁCMO̒lƃMO1/4`1/3Ă邱ƂɂȂDCDNA|[[DNA|[[Â͂܂Ď荞񂾃kNI`hZp̃GL\kNA[[CDNA|[[͂̊ȂDāCDNA|[[̍DNȂDNA|[[DNA|[[Âɂ菜Ă̂ł͂ȂƂwED
@MO̓vC}[RNAꂽ̂DNAK[[ɂADRNȀɂ͕̃XebvlĂ<a href="#R32"><sup>32)</sup></a>DRNase HRNA-DNAnCubhRNA𕪉yfŁC̍yfvC}[RNAƍlꂪł邪CRNase H̓{kNI`hfILVkNI`ȟ̃{kNI`h邱ƂłȂDۂɂ́CDNA|[[DNẢߒɂăvC}[RNAɂԂƁCRNA𒒌^DNA͂C5f3f̃GL\kNA[[tbvGhkNA[[łFEN1RNA̕܂1{DDNA|[[ɂ͂ꂽ1{ƃwJ[[уkNA[[Dna2tؒ̕fD̔ɂ茄ԂłȂDNA|[[߂̂CDNAK[[ɂsDNAƘAD

<h2></h2>
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<h2> </h2>
<ol>
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</ol>
<div class="au-profile">
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