【第9回】ゼロから始めるゲノム解析（R編）

by Naoto Imamachi

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θϩ͔Β࢝ΊΔήϊϜղੳʢ3ฤʣ $IBQUFS$I*1TFRBOBMZTJT /BPUP*NBNBDIJ

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ຊ೔ͷิ଍ࢿྉ ˔ ͪ͜ΒͷϦϙδτϦʹɺ$I*1TFRղੳͷ 3ͷίʔυΛ·ͱΊͨ +VQZUFS/PUFCPPLΛ༻ҙ͠·ͨ͠ɻ IUUQTHJUIVCDPN*NBNBDIJODPNQVUBUJPOBMHFOPNJDTXJUIKVQZUFSMBCS ˔ Ϋϩʔϯ͢Δ PSϒϥ΢β͔Β +VQZUFS/PUFCPPLΛӾཡ͢Δ͜ͱ΋ՄೳͰ͢ɻ ʢҎԼͷσΟϨΫτϦ௚Լʹ /PUFCPPLΛ഑ஔ͍ͯ͠·͢ɻʣ IUUQTHJUIVCDPN*NBNBDIJODPNQVUBUJPOBMHFOPNJDTXJUIKVQZUFSMBCSUSFF NBJOOPUFCPPL

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సࣸҼࢠΛհͨ͠Ҩ఻ࢠൃݱ੍ޚ 3FHVMBUPSZQSPUFJO%/"JOUFSBDUJPOT (Ref: http://www.mun.ca/biology/desmid/brian/BIOL3530/DEVO_10/devo_10.html) సࣸҼࢠʢ5'ʣ ˔ ଞͷҨ఻ࢠ܈Λ੍ޚ͢ΔҼࢠɻ ˔ ಛఆͷ%/"഑ྻʹ݁߹͢Δɻ ʢ5'#4సࣸҼࢠ݁߹αΠτʣ ˔ ੍ޚ͢ΔҨ఻ࢠͷൃݱΛ ༠ಋ PS཈੍͢ΔػೳΛ࣋ͭɻ ˔ ૊৫ಛҟతͳҨ఻ࢠൃݱʹసࣸҼࢠ ͕د༩͍ͯ͠Δɻ ʢ,BEBVLF FUBM5JTTVFTQFDJGJDNJUPUJDCPPLNBSLJOHCZ IFNBUPQPJFUJDUSBOTDSJQUJPOGBDUPS("5"$FMM ʣ ˔ ΋ͪΖΜɺసࣸҼࢠ͚ͩͰ૊৫ಛҟత ͳҨ఻ࢠൃݱ੍ޚΛઆ໌Ͱ͖ΔΘ͚Ͱ ͸ͳ͍͕ʜɻ ʢ4POBXBOF "CIJKFFU3BKFOESB FUBM6OEFSTUBOEJOH UJTTVFTQFDJGJDHFOFSFHVMBUJPO$FMMSFQPSUT ʣ

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ώετϯम০Λհͨ͠ΤϐδΣωςΟΫε੍ޚ 3FHVMBUPSZQSPUFJO%/"JOUFSBDUJPOT (Ref: https://www.abcam.com/epigenetics/histone-modifications) ώετϯम০ʢ)JTUPOF NPEJGJDBUJPOʣ ˔ ώετϯ͸ɺΫϩϚνϯͷओཁͳߏ੒ ੒෼Ͱ͋Γɺ%/"Λύοέʔδϯά ͢Δ଍৔ͱͯ͠ػೳ͢Δɻ ˔ ଟ༷ͳ຋༁ޙम০Λड͚ɺసࣸ׆ੑԽ ʹؔΘΔॏཁͳҼࢠͰ΋͋Δɻ ʢ%/"ͷޙఱతͳԽֶम০ʹΑΔ Ҩ఻ࢠൃݱ੍ޚͷ͜ͱΛɺ ΤϐδΣωςΟΫε੍ޚͱݺͿɻʣ ˔ ࠓճͷղੳʹग़ͯ͘Δώετϯͨͪ ˓ ),NF ˓ ),NF

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(Li, Bing, Michael Carey, and Jerry L. Workman. "The role of chromatin during transcription." Cell 128.4 (2007): 707-719.) ώετϯम০ͷ%/"্ͷ෼෍ ˔ ),NF ˓ 544ʢసࣸ։࢝Ґஔʣͷपลʹ 4IBSQͳϐʔΫ͕ݟΒΕΔɻ ˔ ),NF ˓ (FOFCPEZʢҨ఻ࢠྖҬʣશମ ʹ#SPBEͳϐʔΫ͕ݟΒΕΔɻ ʢͦͷଞʣ$I*1TFRͷσʔλͱ͍͑͹ɺ 3/"ϙϦϝϥʔθͷ෼෍ ˔ 1PMᶘ ˓ 544ʢసࣸ։࢝Ґஔʣͷपลʹ 4IBSQͳϐʔΫ͕ݟΒΕΔɻ ώετϯम০Λհͨ͠ΤϐδΣωςΟΫε੍ޚ 3FHVMBUPSZQSPUFJO%/"JOUFSBDUJPOT

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$I*1TFRͱ͸ʁ .FBTVSJOHQSPUFJO%/"JOUFSBDUJPOTXJUI$I*1TFR ΫϩϚνϯ໔Ӹ௜߱๏ʢ$I*1ʣ ˔ ͬ͘͟Γ͍͏ͱɺλϯύΫ࣭ʢ5' )JTUPOFNPEJGJDBUJPO 1PMᶘͳͲʣ͕ಛఆͷ%/"ྖҬʹ݁߹ ͍ͯ͠Δ͔Ͳ͏͔Λ֬ೝ͢ΔͨΊͷ࣮ݧख๏Ͱ͢ʢղੳʹ͸ R1$3ͳͲ͕࢖ΘΕΔʣɻ ΫϩϚνϯ໔Ӹ௜߱๏ º ࣍ੈ୅γʔέϯεʢ$I*1TFRʣ ˔ λϯύΫ࣭ʢ5' )JTUPOFNPEJGJDBUJPO 1PMᶘͳͲʣ͕݁߹͍ͯ͠Δ%/"ྖҬΛγʔέϯε͠ɺ ର৅ͷλϯύΫ࣭ͷ%/"݁߹ྖҬΛήϊϜϫΠυʹಉఆ͢ΔͨΊͷ࣮ݧख๏Ͱ͢ɻ (Ref: https://www.genewiz.com/Public/Resources/NGS-Platforms/)

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$I*1TFRͱ͸ʁ .FBTVSJOHQSPUFJO%/"JOUFSBDUJPOTXJUI$I*1TFR ΫϩϚνϯ໔Ӹ௜߱๏ º ࣍ੈ୅γʔέϯεʢ$I*1TFRʣ $SPTTMJOL ϗϧϜΞϧσώυΛ༻͍ͯɺλϯύΫ࣭ͱ%/"Λڞ༗݁߹ ͤ͞ΔʢDSPTTMJOLʣɻ %/"഑ྻ λϯύΫ ࣭

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$I*1TFRͱ͸ʁ .FBTVSJOHQSPUFJO%/"JOUFSBDUJPOTXJUI$I*1TFR ΫϩϚνϯ໔Ӹ௜߱๏ º ࣍ੈ୅γʔέϯεʢ$I*1TFRʣ $SPTTMJOL %/" 'SBHNFOUBUJPO %/"഑ྻ λϯύΫ ࣭ ௒Ի೾ PS߬ૉʹΑΔ EJHFTUJPOʹΑΓɺ dCQఔ౓ ͷ%/"அยʹͤΜஅ͢Δʢ%/"'SBHNFOUBUJPOʣɻ

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$I*1TFRͱ͸ʁ .FBTVSJOHQSPUFJO%/"JOUFSBDUJPOTXJUI$I*1TFR ΫϩϚνϯ໔Ӹ௜߱๏ º ࣍ੈ୅γʔέϯεʢ$I*1TFRʣ $SPTTMJOL %/" 'SBHNFOUBUJPO %/"഑ྻ λϯύΫ ࣭ ର৅ͷλϯύΫ࣭ʹର͢Δಛҟతͳ߅ମΛ༻͍ͯɺ%/"ͱ ݁߹͍ͯ͠ΔλϯύΫ࣭Λ *1͢Δʢ*NNVOPQSFDJQJUBUJPOʣɻ ߅ମ *NNVOPQSFDJQJUBUJPO ʢ*1ʣ

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$I*1TFRͱ͸ʁ .FBTVSJOHQSPUFJO%/"JOUFSBDUJPOTXJUI$I*1TFR ΫϩϚνϯ໔Ӹ௜߱๏ º ࣍ੈ୅γʔέϯεʢ$I*1TFRʣ $SPTTMJOL %/" 'SBHNFOUBUJPO DSPTTMJOLΛղআͨ͠ޙɺήϧిؾӭಈ PS࣓ؾϏʔζͳͲ ʹΑΔαΠζηϨΫγϣϯΛߦ͏ɻ IUUQTXXXCFDLNBOKQSFBHFOUTHFOPNJDDMFBOVQ BOETJ[FTFMFDUJPOTJ[FTFMFDUJPO *NNVOPQSFDJQJUBUJPO ʢ*1ʣ 4J[FTFMFDUJPO

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$I*1TFRͱ͸ʁ .FBTVSJOHQSPUFJO%/"JOUFSBDUJPOTXJUI$I*1TFR ΫϩϚνϯ໔Ӹ௜߱๏ º ࣍ੈ୅γʔέϯεʢ$I*1TFRʣ $SPTTMJOL %/" 'SBHNFOUBUJPO *NNVOPQSFDJQJUBUJPO ʢ*1ʣ 4J[FTFMFDUJPO 1$3"NQMJGJDBUJPO 1$3Ͱ໨తͷ഑ྻΛ૿෯ɻ

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$I*1TFRͱ͸ʁ .FBTVSJOHQSPUFJO%/"JOUFSBDUJPOTXJUI$I*1TFR ΫϩϚνϯ໔Ӹ௜߱๏ º ࣍ੈ୅γʔέϯεʢ$I*1TFRʣ ˔ *1ͨ͠λϯύΫ࣭ͷ݁߹αΠτͷ෼෍Λௐ΂Δ͜ͱ͕Ͱ͖·͢ɻ )BOTTFO -BST-1 FUBM5JTTVFTQFDJGJD$5$'rDPIFTJONFEJBUFEDISPNBUJOBSDIJUFDUVSFEFMJNJUTFOIBODFS JOUFSBDUJPOTBOEGVODUJPOJOWJWP/BUVSFDFMMCJPMPHZ

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ ߅ମͷಛҟੑ ˔ λʔήοτͱͳΔλϯύΫ࣭ʹର͢Δ߅ମͷಛҟੑ͕௿͍৔߹ɺ %/"഑ྻ͕ඇಛҟతʹ *1͞Ε͖ͯͯ͠·͍ɺ͜ΕΒ͕ϊΠζͱͳΔɻ ˠ࠷ѱͷ৔߹ɺσʔλΛਖ਼͘͠ղੳͰ͖ͣʹؒҧͬͨ݁࿦Λಋ͖ग़͢ةݥੑ͕͋Δɻ ˔ ಺ࡏੑλϯύΫ࣭ʢ&OEPHFOPVTQSPUFJOTʣʹର͢Δಛҟੑͷߴ͍߅ମ͕ݟ͔ͭΒͳ͍ ৔߹ɺ)"΍'-"(ͳͲͷλά෇͖ͷλϯύΫ࣭ʢ&YPHFOPVTQSPUFJOTʣΛൃݱ ͤͯ͞ *1͢Δख΋͋Δɻ

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ ߅ମͷಛҟੑ ˔ &/$0%&ϓϩδΣΫτͷ߅ମσʔλ ͔ΒɺಛҟੑΛνΣοΫࡁΈͷ߅ମ ϦετΛऔಘՄೳʢΒ͍͠ʣɻ ʢ-BOEU .BSJOPW ,VOEBKF FUBMl$I*14FR (VJEFMJOFTBOE1SBDUJDFTPGUIF&/$0%&BOE NPE&/$0%&$POTPSUJBz (FOPNF3FT r ʣ ˔ ࣮ࡍͷ߅ମσʔλͷࢀরํ๏͸ͪ͜Βɻ ʢ&/$0%&ϓϩδΣΫτͷ/(4σʔλΛ׆༻͢Δ ͍Ζ͍Ζࢼͯ͠ΈΔʣ

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ 4FRVFODFEFQUIʢγʔΫΤϯεͷਂ͞ʣ ˔ ࣍ੈ୅γʔέϯεͰಡΜͩϦʔυ਺͕গͳ͗͢ΔͱɺͦͷޙͷσʔλղੳʢQFBLDBMMʣ ͕೉͘͠ͳΓ·͢ɻ ˔ ώτͷ৔߹ɺd.ϦʔυʢdສϦʔυʣҎ্͕ඞཁͱ͞Ε͍ͯ·͢ɻ ʢ΋ͪΖΜɺࡉ๔छ΍ *1͢ΔλϯύΫ࣭ʹΑͬͯҟͳΓ·͢ʣ ʢ+VOH -VRVFUUF )P FUBMl*NQBDUPG4FRVFODJOH%FQUIJO$I*14FR&YQFSJNFOUTz /VDMFJD"DJET3FT Fʣ

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ 4FRVFODFEFQUIʢγʔΫΤϯεͷਂ͞ʣ ˔ ̎܈ؒൺֱ͍ͨ͠৔߹ɺҎԼͷϊʔϚϥΠζํ๏͕͋Δɻ ʢྫ͑͹ɺ௿ࢎૉঢ়ଶ)ZQPYJB/PSNPYJBͰ )*'$I*1TFRͷσʔλΛऔͬͨ৔߹ʣ ˓ 4DBMFOPSNBMJ[BUJPO ˓ 3PCVTUOPSNBMJ[BUJPO ˓ #BDLHSPVOEOPSNBMJ[BUJPO ˓ &YUFSOBMOPSNBMJ[BUJPO ਤ͸ $IJQDIJQͷσʔλͰ͢ɻ ʢ$I*1TFRͷ͍͍ײ͡ͷσʔλݟ͔ͭΒͣʜʣ ʢ9JB 9JBPCP FUBM*OUFHSBUJWFBOBMZTJTPG)*' CJOEJOHBOEUSBOTBDUJWBUJPOSFWFBMTJUTSPMFJO NBJOUBJOJOHIJTUPOFNFUIZMBUJPOIPNFPTUBTJT 1SPDFFEJOHTPGUIF/BUJPOBM"DBEFNZPG 4DJFODFT

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ 4FRVFODFEFQUIʢγʔΫΤϯεͷਂ͞ʣ ˔ ̎܈ؒൺֱ͍ͨ͠৔߹ɺ 0SMBOEP %BWJE" FUBM2VBOUJUBUJWF$I*1 4FROPSNBMJ[BUJPOSFWFBMTHMPCBMNPEVMBUJPOPG UIFFQJHFOPNF$FMMSFQPSUT ੺ઢͱࠇઢͷࠩΛݕग़͍ͨ͠ɻ

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ 4FRVFODFEFQUIʢγʔΫΤϯεͷਂ͞ʣ ˔ 4DBMFOPSNBMJ[BUJPO ˔ ૯Ϧʔυ਺ͰϦʔυΧ΢ϯτΛׂΔํ๏ʢ$1. 31,.ͳͲʣ ˔ *1ޮ཰͕ಉ͡Ͱ͋Δ͜ͱ͕લఏɻ ˔ ҟͳΔλϯύΫ࣭ͩͱ *1ޮ཰͕ҟͳΔͨΊɺҟͳΔλϯύΫ࣭ͷ $I*1TFR σʔλΛൺֱ͢Δ৔߹͸ෆద੾ͳϊʔϚϥΠζํ๏Ͱ͋Δɻ

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ 4FRVFODFEFQUIʢγʔΫΤϯεͷਂ͞ʣ ˔ 3PCVTUOPSNBMJ[BUJPO ˔ ̎܈ؒͰର৅ͷλϯύΫ࣭ͷେ൒ͷήϊϜ ྖҬ΁ͷ݁߹਌࿨ੑ͕มԽ͠ͳ͍͜ͱ͕ લఏɻ ʢ3/"TFRͷϊʔϚϥΠζʹࣅͯΔʣ ˔ มԽͷͳ͍ྖҬͷશϦʔυ਺Λ TDBMFGBDUPS ͱͯ͠༻͍ͯɺશମͷิਖ਼Λߦ͏ɻ ˔ πʔϧ ˓ ."OPSN ˓ ."OPSNʢ࠷৽όʔδϣϯʣ 4IBP ;IBOH :VBO 0SLJO BOE8BYNBOl."OPSN"3PCVTU .PEFMGPS2VBOUJUBUJWF$PNQBSJTPOPG$I*14FR%BUB4FUTz (FOPNF#JPM 3 5V 4IJRJ FUBM."OPSN GPSRVBOUJUBUJWFMZDPNQBSJOHHSPVQTPG$I*1 TFRTBNQMFT(FOPNF3FTFBSDI .MPH TBNQMFTBNQMFϦʔυ਺ "º MPH TBNQMFº TBNQMFϦʔυ਺

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ 4FRVFODFEFQUIʢγʔΫΤϯεͷਂ͞ʣ ˔ #BDLHSPVOEOPSNBMJ[BUJPO ˔ ήϊϜྖҬΛҎԼͷ̎छྨʹ෼ྨͰ͖ΔͱԾఆɻ ˓ όοΫάϥ΢ϯυྖҬ ˓ γάφϧྖҬʢϐʔΫྖҬλϯύΫ࣭͕݁߹͍ͯ͠ΔྖҬʣ ˔ όοΫάϥ΢ϯυྖҬͷϦʔυ਺Λ TDBMFGBDUPSͱͯ͠༻͍ͯɺ શମͷิਖ਼Λߦ͏ɻ ˔ πʔϧ ˓ /$*4 -JBOH BOE,FMF㶆l/PSNBMJ[BUJPOPG$I*14FR%BUBXJUI$POUSPMz #.$#JPJOGPSNBUJDT "VHVTU όοΫάϥ΢ϯυྖҬ όοΫάϥ΢ϯυͷϦʔυ਺

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ 4FRVFODFEFQUIʢγʔΫΤϯεͷਂ͞ʣ ˔ &YUFSOBMOPSNBMJ[BUJPO ˔ ֎དྷछ PSਓ޻తͳ TQJLFJOΫϩϚνϯ%/"ΛαϯϓϧʹࠞͥΔɻ ˔ ҰఆྔͰ͋Δ͜ͱ͕Θ͔͍ͬͯΔͦΕΒͷ%/"ྔΛج४ͱͯ͠ σʔλΛิਖ਼Ͱ͖Δɻ #POIPVSF #PVOPWB #FSOBTDPOJ FUBMl2VBOUJGZJOH$I*14FR%BUB"4QJLJOH.FUIPE1SPWJEJOHBO *OUFSOBM3FGFSFODFGPS4BNQMFUP4BNQMF/PSNBMJ[BUJPOz (FOPNF3FT r 0SMBOEP %BWJE" FUBM2VBOUJUBUJWF$I*14FROPSNBMJ[BUJPOSFWFBMTHMPCBMNPEVMBUJPOPGUIFFQJHFOPNF $FMMSFQPSUT

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ 1$3ʹΑΔෳ੡ʢ1$3EVQMJDBUJPOʣ ˔ 1$3ʹΑΔ%/"ෳ੡ͷաఔͰ ($DPOUFOUͷόΠΞε͕ੜ͡Δ͜ͱ͕͋Δɻ ˔ ͜ͷόΠΞε͕ QFBLDBMMJOHʹӨڹ͠ɺGBMTFQPTJUJWFͳϐʔΫ͕ಉఆ͞Εͯ ͠·͏Մೳੑ͕͋Δʢ($DPOUFOUͷภΓͷิਖ਼͕ඞཁʣɻ 5FOH BOE*SJ[BSSZl"DDPVOUJOHGPS(D$POUFOU#JBT3FEVDFT4ZTUFNBUJD&SSPSTBOE#BUDI&GGFDUTJO $IJQ4FR%BUBz (FOPNF3FTFBSDI ಉ͡σʔλΛऔ͍ͬͯΔ͸͕ͣɺ ($CJBTͷӨڹͰαϯϓϧؒͰ݁Ռ͕ େ͖͘ҟͳΔʂ

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ #JPMPHJDBMSFQMJDBUFT ˔ ͜Ε͸ݴΘͣ΋͕ͳɻ ˔ ࣮ݧޡࠩΛߟྀ͢Δʹ͸ɺ#JPMPHJDBMSFQMJDBUFTΛऔΔʹӽͨ͜͠ͱ͸ͳ͍ɻ ˔ ʮαϯϓϧ਺͕૿͑Δ ͓͕͔͔ۚΔʯͳͷͰɺ༧ࢉͱ૬ஊʹͳΔɻ

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ $POUSPM࣮ݧͷσβΠϯ ˔ $POUSPM࣮ݧͷछྨɻ ˓ *OQVUDPOUSPM*1લͷήϊϜ%/"Λγʔέϯεɻ ˙ $I*1ͷαϯϓϧΑΓ΋ 4FRVFODFEFQUIΛਂ͘ಡΉඞཁ͕͋Δɻ ʢ$I*1αϯϓϧͷ৔߹ɺೱॖͨ͠ήϊϜͷαϒηοτ͕γʔέϯε ͞ΕΔͷʹର͠ɺ*OQVU͸ήϊϜશମΛγʔέϯε͢Δඞཁ͕͋ΔͨΊʣ ˓ *H(DPOUSPMඇಛҟతͳ *H(߅ମͰ *1ͨ͠αϯϓϧΛγʔέϯεɻ ˙ े෼ͳ%/"Λ֬อ͢Δ͜ͱ͕೉͘͠ɺαϯϓϧௐ੔͕ࠔ೉ɻ ˓ ,OPDLPVUDPOUSPM*1͢ΔλϯύΫ࣭ΛϊοΫΞ΢τͨ͠ࡉ๔Λ༻ҙɻ ˙ &TTFOUJBMHFOFʹ͸దԠͰ͖ͳ͍ɻ ˙ ϊοΫΞ΢τͷӨڹͰ $I*1ͷϓϩϑΝΠϧࣗମ͕ݩͷ΋ͷ͔Βେ͖͘ มΘͬͯ͠·͏ڪΕ͕͋Δɻ ˒ *OQVUDPOUSPMΛ༻ҙ͓͚ͯ͠͹ɺͱΓ͋͑ͣ͸0,ɻ

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$I*1TFRͷσʔλ඼࣭ʹؔΘΔҼࢠ 'BDUPSTUIBUBGGFDU$I*1TFRFYQFSJNFOUBOEBOBMZTJTRVBMJUZ $POUSPM࣮ݧͷσβΠϯ ˔ *OQVUͱ *H(*1DPOUSPMͷ̎छྨΛ༻͍ͨ΄͏͕ਫ਼౓͸্͕Δ͸ͣɻ 9V +JOSVJ FUBM5PNPDLPSOPUBDPNQSFIFOTJWFDPNQBSJTPOPGNPDL*1BOE%/"JOQVUGPS$I*1TFR /VDMFJD"DJET3FTFBSDI HLBB *H(*1Ͱݕग़͞ΕΔ ϐʔΫΛআ֎͢Δ

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$I*1TFRσʔλͷήϊϜ΁ͷϚοϐϯά લॲཧ 1SFQSPDFTTJOH$I*1EBUB ˔ Ϛοϐϯάπʔϧ ˓ #PXUJF ˓ #8"ʢ&/$0%&ϓϩδΣΫτͰ࠾༻͞Ε͍ͯΔʣ ˔ .VMUJNBQQJOHΛڐ༰͢Δ͔Ͳ͏͔ ˓ %/"ͷෳࡶ౓ͷ௿͍Ϧϐʔτ഑ྻʢϨτϩτϥϯεϙκϯɺQTFVEPHFOFͳͲʣΛ ղੳର৅ʹ͍ͨ͠৔߹͸ڐ༰͢Δɻ ˔ %VQMJDBUFESFBETͷѻ͍ ˓ ಉ͡ήϊϜྖҬʹෳ਺ͷϦʔυ͕Ϛοϐϯά͞ΕΔ৔߹ɺ ˙ ϥΠϒϥϦதʹಉ͡%/"ϑϥάϝϯτ͕ෳ਺ଘࡏͨ͠Մೳੑ ˙ 1$3ʹΑΔෳ੡ͷՄೳੑ ͕ߟ͑ΒΕΔɻ ˓ సࣸҼࢠͰɺڱ͍%/"ྖҬʹ݁߹͠͏Δ৔߹ɺ%/"αϯϓϧ͕ೱॖ͞ΕΔͨΊɺ %VQMJDBUFESFBET͕ग़ݱ͠΍͍͢ɻ ͦ͜Ͱɺ%VQMJDBUFESFBETΛ /·Ͱڐ༰͢Δ͜ͱͰײ౓Λ্͛Δख΋͋Δɻ

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$I*1TFRσʔλͷήϊϜ΁ͷϚοϐϯά લॲཧ 1SFQSPDFTTJOH$I*1EBUB ˔ ϒϥοΫϦετͷ%/"ྖҬ ˓ DFMMMJOFɺ؞ࡉ๔ͳͲͰ͸ɺϦϑΝϨϯεήϊϜʹଘࡏ͠ͳ͍ॏෳɾૠೖͳͲͷ ήϊϜߏ଄ͷҧ͍ΛؚΜͰ͍Δέʔε͕͋Δɻ ˓ ͜ͷΑ͏ͳྖҬ͕ϨϑΝϨϯεήϊϜ্ʹҰՕॴ͔͠ଘࡏ͠ͳ͍৔߹ɺϚοϐϯά ͨ͠ࡍʹೱॖ͞ΕɺBSUJGJDBMͳϐʔΫͱͯ͠ݕग़͞ΕΔةݥੑ͕͋Δɻ ˓ ্هͷ໰୊ྖҬΛϒϥοΫϦετʹ·ͱΊͨσʔλ͕ެ։͞Ε͍ͯΔɻ ˙ &/$0%&#MBDLMJTU ˔ IUUQTXXXFODPEFQSPKFDUPSHBOOPUBUJPOT&/$43)'' "NFNJZB )BMFZ. "OTIVM,VOEBKF BOE"MBO1#PZMF5IF&/$0%&CMBDLMJTUJEFOUJGJDBUJPOPGQSPCMFNBUJD SFHJPOTPGUIFHFOPNF4DJFOUJGJDSFQPSUT ϒϥοΫϦετʹڍ͕ͬͯ ͍ΔྖҬʹϚοϐϯά͞Εͨ ϦʔυΛআ֎͢Δɻ

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$I*1TFRσʔλͷήϊϜ΁ͷϚοϐϯά લॲཧ 1SFQSPDFTTJOH$I*1EBUB ˔ &/$0%&ϓϩδΣΫτʹ͓͚Δ $I*1TFRղੳύΠϓϥΠϯʢࢀߟʣ ˓ )JTUPOF$I*1TFR%BUB4UBOEBSETBOE1SPDFTTJOH1JQFMJOF ˙ IUUQTXXXFODPEFQSPKFDUPSHDIJQTFRIJTUPOF ˓ 5SBOTDSJQUJPO'BDUPS$I*1TFR%BUB4UBOEBSETBOE1SPDFTTJOH1JQFMJOF ˙ IUUQTXXXFODPEFQSPKFDUPSHDIJQTFRUSBOTDSJQUJPO@GBDUPS ˔ ιʔείʔυ ˓ &/$0%&%$$DIJQTFRQJQFMJOF&/$0%&$I*1TFRQJQFMJOF EY8%-ʢϫʔΫϑϩʔݴޠʣͰύΠϓϥΠϯ͕૊·Ε͍ͯΔɻ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ $I*1RVBMJUZDPOUSPM ˔ $I*1TFRσʔλͷ඼࣭ΛνΣοΫ͠ɺ඼࣭ͷ௿͍σʔλΛಛఆ͢Δɻ ˔ ҎԼɺओͳ̐εςοϓ αϯϓϧؒͷ૬ؔ܎਺Λ࢖ͬͨΫϥελϦϯά ˔ #JPMPHJDBMSFQMJDBUFTಉ࢜ͷ૬͕ؔߴ͘ͳΔ͔Ͳ͏͔νΣοΫɻ ήϊϜϒϥ΢β্ͰͷσʔλՄࢹԽ ˔ ੜσʔλΛՄࢹԽͯ͠ɺϦʔυ෼෍ΛνΣοΫ͢Δɻ %/"ϑϥάϝϯτͷฏۉ௕ͷಉఆ ˔ ϐʔΫΛݕग़͢Δࡍͷύϥϝʔλͱͯ͠࢖༻͢Δɻ ($CJBTͷՄࢹԽ ˔ $I*1FOSJDINFOUͱ ฏۉ ($DPOUFOUΛϓϩοτ αϯϓϧؒͰ෼෍Λൺֱɻ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 5IFEBUB ˔ ͜͜Ͱѻ͏σʔλ ˓ సࣸҼࢠʢ5'ʣ$5$' 4.$ ;/' QPMᶘ 3/"QPMZNFSBTF ˓ )JTUPOFNPEJGJDBUJPO),NF ),NF )LBD )LNF ˓ ֤छ *OQVUαϯϓϧ

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QPMᶘ QPMᶘ $I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 4BNQMFDMVTUFSJOH αϯϓϧؒͷ૬ؔ܎਺Λ࢖ͬͨΫϥελϦϯά ˔ CQ͝ͱͷ XJOEPXʹ۠੾ͬͯɺϦʔυΧ΢ϯτ $1.Λܭࢉɻ $1.ສϦʔυ͋ͨΓͷϦʔυ਺ʢϦʔυΧ΢ϯτº ૯Ϧʔυ਺ ʣ ˔ αϯϓϧશͯͷ $1.஋ͷϖΞؒͷ૬ؔΛܭࢉ͠ՄࢹԽɻ CQ QPMᶘ $1. $1. $1. $1. $1. $1. $1. QPMᶘ $1. $1. $1. $1. $1. $1. $1. R2 = 0.99

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 4BNQMFDMVTUFSJOH αϯϓϧؒͷ૬ؔ܎਺Λ࢖ͬͨΫϥελϦϯά લఏ஌ࣝ ˔ 4.$ίώʔγϯͷαϒϢχοτɻ ˔ $5$'ίώʔγϯͷෳ߹ମͱ݁߹͢ΔసࣸҼࢠɻ ˔ ͭ·Γɺ4.$ͱ $5$'ͷ $I*1TFRͷσʔλؒʹڧ͍૬͕ؔݟΒΕΔ͸ͣɻ )BOTTFO -BST-1 FUBM5JTTVFTQFDJGJD$5$'rDPIFTJONFEJBUFEDISPNBUJOBSDIJUFDUVSFEFMJNJUTFOIBODFS JOUFSBDUJPOTBOEGVODUJPOJOWJWP/BUVSFDFMMCJPMPHZ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 4BNQMFDMVTUFSJOH αϯϓϧؒͷ૬ؔ܎਺Λ࢖ͬͨΫϥελϦϯά ˔ αϯϓϧશͯͷ $1.஋ͷϖΞؒͷ૬ؔΛܭࢉ͠ՄࢹԽɻ σʔλ͔Βݴ͑Δ͜ͱ ˔ $5$'@S $5$'@S 4.$@S ͷؒʹ͸ ڧ͍૬͕ؔݟΒΕΔʢ૝ఆ௨Γʣɻ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 4BNQMFDMVTUFSJOH αϯϓϧؒͷ૬ؔ܎਺Λ࢖ͬͨΫϥελϦϯά ˔ αϯϓϧશͯͷ $1.஋ͷϖΞؒͷ૬ؔΛܭࢉ͠ՄࢹԽɻ σʔλ͔Βݴ͑Δ͜ͱ ˔ $5$'@S $5$'@S 4.$@S ͷؒʹ͸ ڧ͍૬͕ؔݟΒΕΔʢ૝ఆ௨Γʣɻ ˔ ҰํͰɺ4.$@Sͱ *OQVUσʔλͷؒʹ ൺֱతڧ͍૬͕ؔݟΒΕΔɻ ˠ4.$@Sͷσʔλ͸ *1ʹΑΔ &OSJDINFOUͷ ௿͍ɺϊΠζͷଟ͍αϯϓϧͰ͋Δͱߟ͑ΒΕΔɻ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 7JTVBMJ[BUJPOJOUIFHFOPNFCSPXTFS ήϊϜϒϥ΢β্ͰͷσʔλՄࢹԽ ओͳήϊϜϒϥ΢β ˔ *(7σεΫτοϓΞϓϦʢΠϯετʔϧඞཁʣɻ ˔ 6$4$(FOPNF#SPXTFS8FCΞϓϦʢΠϯετʔϧෆཁʣɻ ˔ (WJ[3্ͰͷՄࢹԽ ˠࠓ೔͸ίϨΛ࢖͍·͢ʂ ϚοϐϯάσʔλʢPSήϊϜϒϥ΢βͰՄࢹԽ͢ΔͨΊͷΧόϨοδσʔλʣ ˔ #". ˔ CJH8JH ˔ #&% ˔ #FE(SBQI ͳͲͳͲɻ (WJ[ʹΑΔՄࢹԽ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 1MVTBOENJOVTTUSBOEDSPTTDPSSFMBUJPO %/"࠯ͷΫϩε૬ؔ $I*1TFRͷϚοϐϯά͞ΕͨϦʔυͷύλʔϯ ˔ Ұൠతʹɺ$I*1TFRͷϦʔυ͸ɺ຤୺͔ΒಡΜͩ৔߹͸ ࠯ɺ ຤୺͔ΒಡΜͩ৔߹͸ ࠯ʹϚοϐϯά͞ΕΔɻ *1͞Εͨ%/"഑ྻ 'PSXBSEϦʔυ ࠯ ࠯ ࠯ 3FWFSTFϦʔυ ࠯ λϯύΫ࣭ͷ݁߹ྖҬʹɺ ̎ͭͷϐʔΫʢࢁʣ͕Ͱ͖ΔΠϝʔδɻ ʢࠨͷਤ͸ތுͯ͠ॻ͍ͯ·͢ʣ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 1MVTBOENJOVTTUSBOEDSPTTDPSSFMBUJPO %/"࠯ͷΫϩε૬ؔ +BDDBSE܎਺ʢྨࣅ౓܎਺ʣ࿨ू߹Λੵू߹Ͱׂͬͨ஋ ˔ ϦʔυΛ dCQͷൣғͰγϑτͤ͞ɺ+BDDBSE܎਺ʢdʣΛܭࢉɻ 'PSXBSEϦʔυ ༝དྷͷϐʔΫ ࠯ 3FWFSTFϦʔυ ༝དྷͷϐʔΫ ࠯ খ େ +BDDBSE܎਺ ʲٕज़ղઆʳू߹ͷྨࣅ౓ +BDDBSE܎਺ɼ%JDF܎਺ɼ4JNQTPO܎਺

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 1MVTBOENJOVTTUSBOEDSPTTDPSSFMBUJPO %/"࠯ͷΫϩε૬ؔ +BDDBSE܎਺ʢྨࣅ౓܎਺ʣ࿨ू߹Λੵू߹Ͱׂͬͨ஋ ˔ ϦʔυΛ dCQͷൣғͰγϑτͤ͞ɺ+BDDBSE܎਺ʢdʣΛܭࢉɻ %/"഑ྻ λϯύΫ࣭ dCQ *1ͨ͠ %/"GSBHNFOU௕͕ ͍͍ͩͨ CQͰ͋Δ͜ͱ͕Θ͔Δɻ ˠ͜ͷ஋Λ QFBLDBMM࣌ʹ࢖͏ɻ ·ͨɺ͜ͷ஋͕ Ͱͳ͍ ୯ௐʹݮগͯ͠ ͍ͳ͍ͷͰɺ$I*1TFRσʔλ͕ಛఆͷ%/" ഑ྻΛద੾ʹ *1Ͱ͖͍ͯΔͱ൑அ͢Δɻ ˒ ޙͰɺ$POUSPMσʔλͰࢼͯ͠ΈΑ͏ɻ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ ($CJBTRVBOUJGJDBUJPO ($όΠΞεͷఆྔ ˔ 1$3ʹΑΔ%/"ෳ੡ͷաఔͰ ($DPOUFOUͷόΠΞε͕ੜ͡Δ͜ͱ͕͋Δɻ ˔ ͜͜Ͱ͸ɺαϯϓϧؒͰ ($DPOUFOUʹภΓ͕ͳ͍͔Ͳ͏͔νΣοΫ͢Δɻ ˔ CQ͝ͱͷ XJOEPXʹ۠੾ͬͯɺ($ؚ༗཰Λܭࢉ͢Δɻ σʔλ͔Βݴ͑Δ͜ͱ ˔ ($DPOUFOU͕ҟৗʹ௿͍ PSߴ͍ྖҬ͸ 1$3PSαΠζηϨΫγϣϯ͕͏·͘Ͱ͖ ͍ͯͳ͍ʢภΓ͕͋Δʣɻ ˔ ͦ΋ͦ΋ɺ͜Ε ($DPOUFOU͡Όͳ͍ʁ ʢ$Q(JTMBOEΛؚΉؔ܎Ͱɺ͙ Β͍ʹͳΔ͸ͣɻʣ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ ($CJBTRVBOUJGJDBUJPO ($όΠΞεͷఆྔ ˔ 1$3ʹΑΔ%/"ෳ੡ͷաఔͰ ($DPOUFOUͷόΠΞε͕ੜ͡Δ͜ͱ͕͋Δɻ ˔ ͜͜Ͱ͸ɺαϯϓϧؒͰ ($DPOUFOUʹภΓ͕ͳ͍͔Ͳ͏͔νΣοΫ͢Δɻ ˔ CQ͝ͱͷ XJOEPXʹ۠੾ͬͯɺ($ؚ༗཰Λܭࢉ͢Δɻ ࢲͷٙ໰ ˔ ͜Εɺ($DPOUFOU͡Όͳͯ͘ɺ ($Ԙجͷग़ݱස౓ΛݟͯΔ͚ͩ͡Όʜʁ ˔ ҎԼͷܭࢉ͔ࣜΒ LC͋ͨΓͷ ($DPOUFOUΛܭࢉ͢Δͷ͕ਖ਼͍͠͸ͣɻ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ ($CJBTRVBOUJGJDBUJPO ($όΠΞεͷఆྔ ˔ 1$3ʹΑΔ%/"ෳ੡ͷաఔͰ ($DPOUFOUͷόΠΞε͕ੜ͡Δ͜ͱ͕͋Δɻ ˔ ͜͜Ͱ͸ɺαϯϓϧؒͰ ($DPOUFOUʹภΓ͕ͳ͍͔Ͳ͏͔νΣοΫ͢Δɻ ˔ CQ͝ͱͷ XJOEPXʹ۠੾ͬͯɺ($ؚ༗཰Λܭࢉ͢Δɻ #JPMPHJDBMSFQMJDBUFTؒͰͷൺֱ ˔ ͍͍ͩͨɺ෼෍͕ࣅ௨͍ͬͯΔ͔Βɺ ($όΠΞε͕ͳ͍ͱ൑அɻ ˔ $1.͸αϯϓϧؒͰ͹Βͭ͘ͷͰߟྀ ͠ͳͯ͘ྑͦ͞͏ɻ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 4FRVFODFSFBEHFOPNJDEJTUSJCVUJPO ֤ϦʔυͷήϊϜ্Ͱͷ෼෍ ৼΓฦΓ (Li, Bing, Michael Carey, and Jerry L. Workman. "The role of chromatin during transcription." Cell 128.4 (2007): 707-719.) ώετϯम০ͷ%/"্ͷ෼෍ ˔ ),NF ˓ 544ʢసࣸ։࢝Ґஔʣͷपลʹ 4IBSQͳϐʔΫ͕ݟΒΕΔɻ ˔ ),NF ˓ (FOFCPEZʢҨ఻ࢠྖҬʣશମ ʹ#SPBEͳϐʔΫ͕ݟΒΕΔɻ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 4FRVFODFSFBEHFOPNJDEJTUSJCVUJPO ֤ϦʔυͷήϊϜ্Ͱͷ෼෍ ˔ 544 &YPO *OUSPO JOUFSHFOJDͲͷྖҬʹϦʔυ͕෼෍͍ͯ͠Δͷ͔ݟͯΈΔɻ ˔ Ϧʔυ͕ෳ਺ͷྖҬʹ·͕ͨΔ৔߹ɺҎԼͷ༏ઌॱʹΧςΰϦʔ෼͚͢Δɻ ˓ 544FYPOJOUSPOJOUFSHFOJD ˔ Ҩ఻ࢠ৘ใͱͯ͠͸ɺ&OTFNCMʢ(&/$0%&ʣͷσʔλΛ༻͍Δɻ ˓ ίϨʹ߆Δඞཁ͸ͳ͘ɺ3FG4FRͳͲ޷͖ͳ΍ͭΛ࢖͑͹͍͍ɻ Ҩ఻ࢠྖҬ

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$I*1TFRσʔλͷ඼࣭νΣοΫʢ2$ʣ 4FRVFODFSFBEHFOPNJDEJTUSJCVUJPO ֤ϦʔυͷήϊϜ্Ͱͷ෼෍ ˔ 544 &YPO *OUSPO JOUFSHFOJDͲͷྖҬʹϦʔυ͕෼෍͍ͯ͠Δͷ͔ݟͯΈΔɻ ˔ Ϧʔυ͕ෳ਺ͷྖҬʹ·͕ͨΔ৔߹ɺҎԼͷ༏ઌॱʹΧςΰϦʔ෼͚͢Δɻ ˓ 544FYPOJOUSPOJOUFSHFOJD σʔλ͔Βݴ͑Δ͜ͱ ˔ *OUFSHFOJDྖҬʹϦʔυ͕ଟ͍ɻ ˔ ͋͑ͯݴ͏ͳΒɺ ˓ $5$'&YPO*OUSPO ˓ ),NF544 ˓ ;/'͸ 544 ͷ෼෍͕΍΍ଟ͍ɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLDBMMJOH ͍͟ɺQFBLDBMM΁ʂ ˔ ͜Ε·Ͱͷ2$ͷ݁Ռ͔Βɺ$I*1TFRͷσʔλ඼࣭ʹ໰୊͸ͳ͍ͱ൑அ͞Εͨ৔߹ɺ ԼྲྀͷղੳʹਐΉ͜ͱ͕Ͱ͖Δɻ ˔ $I*1σʔλͱ $POUSPMσʔλͷΧόϨοδσʔλΛར༻ͯ͠ɺλϯύΫ࣭͕݁߹ͯ͠ ͍ΔྖҬΛݟ͚ͭΔʢQFBLDBMMʣɻ ˔ ͜͜Ͱ͸ɺOPSN3 #JPDPOEVDUPSύοέʔδΛར༻͠·͢ɻ ˔ ޙड़͢ΔɺTIBSQPSCSPBEϐʔΫͷ྆ํΛݕग़Ͱ͖ΔπʔϧͰ͢ɻ ˔ ͜ͷπʔϧ͸ɺ#JPMPHJDBMSFQMJDBUFTʹରԠ͍ͯ͠·ͤΜɻ ˒ 3ʹͩ͜ΘΒͳ͚Ε͹ɺ."$4ͳͲͷ QFBLDBMMFS͸ͨ͘͞Μ͋Γ·͢ɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 5ZQFTPG$I*1TFRFYQFSJNFOUT ˔ ର৅ͷλϯύΫ࣭ͷੑ࣭ʹΑͬͯɺ$I*1TFRσʔλ͸ओʹҎԼͷ̏छྨʹ෼ྨ͞ΕΔɻ 4IBSQϐʔΫ ˔ ਺ CQఔ౓ͷ୹͍ήϊϜྖҬ͔ΒͳΔϐʔΫɻ ˔ ओʹɺసࣸҼࢠ΍ώετϯम০ʢ),NFͳͲʣ͕֘౰͢Δɻ #SPBEϐʔΫ ˔ ਺ CQʹ౉Δ޿͍ൣғͷήϊϜྖҬ͔ΒͳΔϐʔΫɻ ˔ ओʹɺ(FOFCPEZʹ෼෍͢Δ ),NFɺϙϦίʔϜෳ߹ମʹ෼෍͢Δ ),NF ͳͲ͕֘౰͢Δɻ .JYFEϐʔΫ ˔ ্هͷϐʔΫ͕ࠞࡏ͢Δέʔεɻ ˔ 3/"1PMZNFSBTF͕ͦΕʹ֘౰͢Δɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 5ZQFTPG$I*1TFRFYQFSJNFOUT ˔ πʔϧͷબ୒ΛݟۃΊΔ࠷΋༗ޮͳखஈ͸ɺσʔλΛՄࢹԽ͢Δ͜ͱɻ ˔ ͜͜Ͱ͸ɺ3Λ࢖ͬͯɺҎԼͷ $I*1TFRαϯϓϧΛͦΕͧΕՄࢹԽͯ͠ΈΔɻ ˓ ),NF 4IBSQ ˓ ),NF #SPBE ˓ 1PM .JYFE ˔ ),NF544ʹ TIBSQͳϐʔΫ͕֬ೝͰ͖Δɻ ˔ ),NFHFOFCPEZશମʹ FOSJDI͍ͯ͠Δͷ͕ ֬ೝͰ͖Δɻ ˔ 1PM544ྖҬʹ TIBSQͳϐʔΫɺHFOFCPEZશମ ʹ΋ FOSJDI͍ͯ͠Δͷ͕֬ೝͰ͖Δ NJYFE ɻ 544

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLDBMMJOH4IBSQQFBLT ˔ 4IBSQϐʔΫͷݕग़ ˓ ),NF ˔ *OQVUͱ $I*1αϯϓϧͷ $1.Λൺֱɻ ˔ ઢΑΓ্ͷ఺͸ɺ$I*1αϯϓϧͷ΄͏ ͕ FOSJDI͍ͯ͠Δɻ ˔ ઢΑΓԼͷ఺͸ɺ*OQVUαϯϓϧͷ΄͏ ͕ FOSJDI͍ͯ͠Δɻ ͱ͍͏͜ͱΛࣔ͢ɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLDBMMJOH4IBSQQFBLT ˔ 4IBSQϐʔΫͷݕग़ ˓ ),NF ˔ OPSN3ϥΠϒϥϦΛ༻͍ͯɺϐʔΫΛಉఆ͢Δɻ ˔ ੺ɿಉఆ͞ΕͨϐʔΫɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLDBMMJOH4IBSQQFBLT ˔ 4IBSQϐʔΫͷݕग़ ˓ ),NF ˔ OPSN3ϥΠϒϥϦΛ༻͍ͯɺϐʔΫΛಉఆ ˠ(WJ[ͰՄࢹԽ͢Δɻ ˔ ॎ࣠ͷ஋͕ଗ͍ͬͯͳ͍ͷͰɺΘ͔Γ ʹ͍͕͘ɺόοΫάϥ΢ϯυͰ͋Δ *OQVUʹରͯ͠ɺ$5$'σʔλͰ͸ TIBSQͳϐʔΫཱ͕͍ͬͯΔ͜ͱ͕Θ ͔Δɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLDBMMJOH#SPBESFHJPOT ˔ #SPBEϐʔΫͷݕग़ ˓ ),NF ˔ OPSN3ϥΠϒϥϦΛ༻͍ͯɺϐʔΫΛಉఆ͢Δ ˠ(WJ[ͰՄࢹԽ͢Δɻ ˔ ਫ৭͕ಉఆ͞ΕͨϐʔΫɻ ˔ Ҩ఻ࢠྖҬʹରͯ͠ɺCSPBEͳϐʔΫ ʹͳ͍ͬͯΔ͜ͱ͕Θ͔Δɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLRVBMJUZDPOUSPM1FSDFOUBHFPGSFBETJOQFBLT ˔ ϐʔΫͷ඼࣭؅ཧʢશϦʔυͷ͏ͪϐʔΫྖҬʹ઎ΊΔׂ߹ʣ ˓ *OQVUαϯϓϧͷ TFRVFODFEFQUI͕े෼Ͱͳ͘ɺ͜ͷ৔߹ɺޡͬͯϐʔΫΛ ݕग़ͯ͠͠·͏έʔε͕͋Δɻ ˓ *OQVUͱ $I*1αϯϓϧʹ͓͚Δɺઌ΄Ͳಉఆͨ͠ϐʔΫྖҬͷ FOSJDINFOUΛ ܭࢉ͢Δɻ ˔ ϐʔΫྖҬʹؚ·ΕΔϦʔυ਺ͷׂ߹͕ *OQVUΑΓ΋ ),NFͷ΄͏͕ଟ͍ɻ ˠ$I*1αϯϓϧͷ඼࣭͸ѱ͘ͳͦ͞͏ɻ ˔ Ϧʔυ਺ͷׂ߹͸ɺ߅ମͷ࣭ʹ΋ґଘͯ͠ ͍Δɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLRVBMJUZDPOUSPM%/"NPUJGTPOQFBLT ˔ ϐʔΫͷ඼࣭؅ཧʢશϐʔΫͷ͏ͪϞνʔϑ഑ྻʹ઎ΊΔׂ߹ʣ Ϟνʔϑ഑ྻΛؚΉϐʔΫͷׂ߹Λௐ΂Δɻ Ϟνʔϑ഑ྻͷ෼෍Λௐ΂Δ ϐʔΫྖҬͷதԝʹϞνʔϑ഑ྻ͕ଘࡏ͢Δ͸ͣ ɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLRVBMJUZDPOUSPM%/"NPUJGTPOQFBLT ˔ ϐʔΫͷ඼࣭؅ཧʢશϐʔΫͷ͏ͪϞνʔϑ഑ྻʹ઎ΊΔׂ߹ʣ Ϟνʔϑ഑ྻΛؚΉϐʔΫͷׂ߹Λௐ΂Δɻ ˔ $5$'ͷϞνʔϑ഑ྻ͕ϐʔΫྖҬʹؚΉׂ߹Λܭࢉͯ͠Έ·͠ΐ͏ɻ ˔ ͜͜Ͱ͸ɺ.PUJG%#ͱݺ͹ΕΔ #JPDPOEVDUPSͷσʔλϕʔεΛར༻͠·͢ɻ ˔ .PUJG%#͸ෳ਺ͷϞνʔϑ഑ྻσʔλϕʔεΛू໿Խͨ͠΋ͷͰ͢ɻ ˔ ҎԼ͸ɺ$5$'ͷϞνʔϑ഑ྻͷ֤Ԙجͷग़ݱස౓ͷදͰ͢ɻ ˔ ্هͷϚτϦΫε ߦྻ ͸ɺసࣸҼࢠ͕ର৅ͷ഑ྻʹ݁߹͢Δ͔Ͳ͏͔ɺͦͷ֬཰Λܭࢉ͢ΔͨΊʹ࢖ΘΕ·͢ɻ ˔ ͔͜͜͠͠Ͱ஫ҙ͠ͳ͍ͱ͍͚ͳ͍͜ͱ͸ɺόοΫάϥ΢ϯυͷ֬཰Λߟྀ͍ͯ͠ͳ͍ͱ͍͏఺Ͱ͢ɻήϊϜ഑ྻ͸ ͣͭۉ౳ʹ " 5 ( $ͷ֤Ԙج͕෼෍͍ͯ͠ΔΘ͚Ͱ͸ͳ͘ɺ͋Δఔ౓ͷภΓ͕͋ΔͨΊɺόοΫάϥ΢ϯυͷ֬ ཰Λߟྀ͢Δඞཁ͕͋Γ·͢ʢࠓճ͸ແࢹͯ͠ɺͣͭͱ͓͖ͯ͠·͢ʣɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLRVBMJUZDPOUSPM%/"NPUJGTPOQFBLT ˔ ϐʔΫͷ඼࣭؅ཧʢશϐʔΫͷ͏ͪϞνʔϑ഑ྻʹ઎ΊΔׂ߹ʣ Ϟνʔϑ഑ྻΛؚΉϐʔΫͷׂ߹Λௐ΂Δɻ ˔ ઌ΄ͲͷϚτϦΫε͔ΒɺόοΫάϥ΢ϯυΛߟྀͨ͠είΞΛܭࢉ͢Δࣜ͸ҎԼ ͷͱ͓ΓͰ͢ʢ͸͍ɺখ೉͘͠ॻ͍ͯ·͕͢୯७ͳׂΓࢉͰ͢ʜʣɻ ˔ #L όοΫάϥ΢ϯυͷԘجͷग़ݱ֬཰ ˔ 1J L Ϟνʔϑ഑ྻͷ֤Ԙجͷग़ݱ֬཰ 1 2 3 4 A 0.17 0.23 0.29 0.10 C 0.42 0.28 0.30 0.32 G 0.25 0.23 0.26 0.27 T 0.16 0.27 0.15 0.31 1 2 3 4 A -0.56 -0.12 0.21 -1.32 C 0.75 0.16 0.26 0.36 G 0.00 -0.12 0.06 0.11 T -0.64 0.11 -0.74 0.31 ˔ όοΫάϥ΢ϯυ ΛԼճΔ৔߹ ϚΠφεʮʯ ˔ όοΫάϥ΢ϯυ Λ্ճΔ৔߹ ϚΠφεʮʯ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLRVBMJUZDPOUSPM%/"NPUJGTPOQFBLT ˔ ϐʔΫͷ඼࣭؅ཧʢશϐʔΫͷ͏ͪϞνʔϑ഑ྻʹ઎ΊΔׂ߹ʣ Ϟνʔϑ഑ྻΛؚΉϐʔΫͷׂ߹Λௐ΂Δɻ ˔ ϐʔΫྖҬ TFRVFODF ͷԘج഑ྻʹରͯ͠ɺ֤ԘجͷείΞΛܭࢉ͢Δɻ 1 2 3 4 A -0.56 -0.12 0.21 -1.32 C 0.75 0.16 0.26 0.36 G 0.00 -0.12 0.06 0.11 T -0.64 0.11 -0.74 0.31 1PTJUJPOTQFDJGJD TDPSJOHNBUSJY 144. 4FRVFODF " $ " 5 4FRVFODFείΞ "$"5഑ྻʹରͯ͠ɺ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLRVBMJUZDPOUSPM%/"NPUJGTPOQFBLT ˔ ϐʔΫͷ඼࣭؅ཧʢશϐʔΫͷ͏ͪϞνʔϑ഑ྻʹ઎ΊΔׂ߹ʣ Ϟνʔϑ഑ྻΛؚΉϐʔΫͷׂ߹Λௐ΂Δɻ ˔ NJONBYOPSNBMJ[BUJPOͱ͍͏ํ๏Ͱɺdͷൣғͷσʔλʹิਖ਼ɻ ʢ͸͍ɺখ೉͘͠ॻ͍ͯ·͕͢ɺ࠷খ஋Λʹɺ࠷େ஋Λʹิਖ਼ͯ͠Δʹա͗·ͤΜʣ 1 2 3 4 A -0.56 -0.12 0.21 -1.32 C 0.75 0.16 0.26 0.36 G 0.00 -0.12 0.06 0.11 T -0.64 0.11 -0.74 0.31 1PTJUJPOTQFDJGJD TDPSJOHNBUSJY 144. 4FRVFODF " $ " 5 ࠷খείΞ 5"5"഑ྻʹରͯ͠ɺ ʔ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLRVBMJUZDPOUSPM%/"NPUJGTPOQFBLT ˔ ϐʔΫͷ඼࣭؅ཧʢશϐʔΫͷ͏ͪϞνʔϑ഑ྻʹ઎ΊΔׂ߹ʣ Ϟνʔϑ഑ྻΛؚΉϐʔΫͷׂ߹Λௐ΂Δɻ ˔ NJONBYOPSNBMJ[BUJPOͱ͍͏ํ๏Ͱɺdͷൣғͷσʔλʹิਖ਼ɻ ʢ͸͍ɺখ೉͘͠ॻ͍ͯ·͕͢ɺ࠷খ஋Λʹɺ࠷େ஋Λʹิਖ਼ͯ͠Δʹա͗·ͤΜʣ ิਖ਼ͨ͠είΞ "$"5഑ྻʹରͯ͠ɺ ˔ ׂͱዞҙతͰ͸͋Δ͕ɺείΞ͕ Λ͖͍͠஋ʢΧοτΦϑ஋ʣͱ͢Δͱɺ ্هͷείΞ͸ͦΕΛԼճ͍ͬͯΔͨΊɺ"$"5഑ྻ͸Ϟνʔϑ഑ྻͰ͸ͳ͍ͱ ൑அ͞ΕΔɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLRVBMJUZDPOUSPM%/"NPUJGTPOQFBLT ˔ ϐʔΫͷ඼࣭؅ཧʢશϐʔΫͷ͏ͪϞνʔϑ഑ྻʹ઎ΊΔׂ߹ʣ Ϟνʔϑ഑ྻΛؚΉϐʔΫͷׂ߹Λௐ΂Δɻ ˔ QPTJUJPOGSFRVFODZNBUSJY 1'. Λ࢖ͬͯɺϚτϦΫε಺ͷ֤Ґஔ Ԙج ͷ ॏཁ౓ ৘ใྔ Λܭࢉ͢Δ͜ͱ͕Ͱ͖·͢ɻ ˔ ͜Ε͸ɺసࣸҼࢠͷಛఆ഑ྻʹର͢Δ݁߹͠΍͢͞ΛఆྔԽ͢Δ΋ͷͰ͢ɻ ˔ ͞Βʹɺ݁߹ϚτϦΫε ߦྻ ΛγʔΫΤϯεϩάͱͯ͠ࢹ֮తʹදݱ͢Δ͜ͱ ΋Ͱ͖·͢ɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLRVBMJUZDPOUSPM%/"NPUJGTPOQFBLT ˔ ϐʔΫͷ඼࣭؅ཧʢશϐʔΫͷ͏ͪϞνʔϑ഑ྻʹ઎ΊΔׂ߹ʣ Ϟνʔϑ഑ྻΛؚΉϐʔΫͷׂ߹Λௐ΂Δɻ ˔ ૬ରΤϯτϩϐʔ dͷൣғ ͱͯ͠ఆྔԽ͞Ε·͢ɻ ˔ ૬ରΤϯτϩϐʔͷಛ௃ ˓ ֤Ԙجͷग़ݱ֬཰͕ۉ౳ͳ৔߹ ಥग़ͯ͠ग़ݱ֬཰ͷߴ͍Ԙج͕ଘࡏ͠ͳ͍৔߹ ߴ͘ͳΔ ˓ ͋Δ̍ͭͷԘجͷग़ݱ֬཰͕ߴ͍৔߹ ಥग़ͯ͠ग़ݱ֬཰ͷߴ͍Ԙج͕ଘࡏ͢Δ৔߹ ௿͘ͳΔ ˠಛఆͷԘجͷग़ݱස౓͕ߴ͍΄Ͳɺߴ͍਺஋ʹ͍ͨ͠ʂ ˔ 1J L 1'.ͷΧϥϜ ͷ Ԙج ͷग़ݱ֬཰ɻ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FBLRVBMJUZDPOUSPM%/"NPUJGTPOQFBLT ˔ ϐʔΫͷ඼࣭؅ཧʢશϐʔΫͷ͏ͪϞνʔϑ഑ྻʹ઎ΊΔׂ߹ʣ Ϟνʔϑ഑ྻΛؚΉϐʔΫͷׂ߹Λௐ΂Δɻ ˔ JOGPSNBUJPODPOUFOUʢ*$ʣ ࠷େΤϯτϩϐʔ͔Β؍ଌ͞ΕͨྻͷΤϯτϩϐʔΛࠩ͠Ҿ͍ͨ஋ɻ ˔ *$ͷಛ௃ ˓ ͋ΔԘجͷग़ݱ֬཰͕ߴ͍৔߹ߴ͘ͳΔ ˓ ͢΂ͯͷԘج͕౳͘͠ग़ݱ͢ΔՄೳੑ͕ߴ͍৔߹௿͘ͳΔ ˠಛఆͷԘجͷग़ݱස౓͕ߴ͍΄Ͳɺߴ͍਺஋ʹͳͬͨʂ ˔ Τϯτϩϐʔͷ࠷େ஋ ˔ 1J L 1'.ͷΧϥϜ ͷ Ԙج ͷग़ݱ֬཰

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$I*1TFRσʔλ͔Βͷ QFBLDBMM 1FSDFOUBHFPGQFBLTXJUIUIFNPUJG ˔ ϐʔΫͷ඼࣭؅ཧʢશϐʔΫͷ͏ͪϞνʔϑ഑ྻʹ઎ΊΔׂ߹ʣ Ϟνʔϑ഑ྻΛؚΉϐʔΫͷׂ߹Λௐ΂Δɻ ˔ ٯੵࢉάϥϑɻ ˔ શମͰ ఔ౓ͷϐʔΫ഑ྻͰϞνʔϑ഑ྻ͕ಉఆ͞Εͨ͜ͱ͕Θ͔Δ ஋͸ߴ ͚Ε͹ߴ͍΄͏͕Α͍ ɻ ˔ ϐʔΫϥϯΫ͕ ˓ FOSJDINFOU͕࠷΋ߴ͍ϐʔΫɻ ˔ ϐʔΫϥϯΫ͕ ࠷ԼҐͷ ˓ FOSJDINFOU͕࠷΋௿͍ϐʔΫɻ ࠨͷਤΛಡΈղ͘ͱɺ ˔ ϐʔΫϥϯΫ dͷ͏ͪɺ ͕ϐʔΫΛؚΉɻ ˔ ϐʔΫϥϯΫ dʢશମʣͰΈΔͱɺ ͕ϐʔΫΛؚΉɻ ͱ͍͏͜ͱ͕Θ͔Δɻ ϐʔΫϥϯΫ͕௿͍ॱʹ Ϟνʔϑ഑ྻͷଘࡏ༗ແͷׂ߹Λੵࢉ

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$I*1TFRσʔλ͔Βͷ QFBLDBMM .PUJGMPDBMJ[BUJPO ˔ ϐʔΫͷ඼࣭؅ཧʢશϐʔΫͷ͏ͪϞνʔϑ഑ྻʹ઎ΊΔׂ߹ʣ Ϟνʔϑ഑ྻͷ෼෍Λௐ΂Δɻ ˔ $I*1TFRͷσʔλͷ඼࣭͕ྑ͚Ε͹ɺϞνʔϑ഑ྻ͸֤ϐʔΫͷ௖఺ʹہࡏ͠ ͍ͯΔͱ༧૝͞ΕΔɻ ૝ఆ௨ΓɺϐʔΫྖҬͷதԝ͕ ࠷΋Ϟνʔϑ഑ྻͷଘࡏׂ߹͕ߴ͍͜ͱ͕Θ͔Δɻ