<?xml version="1.0"?>
<article id="nature07777" language="eng" relation="no" origsrc="yes" publish="issue">
  <pubfm>
    <jtl>Angher</jtl>
    <vol>503</vol>
    <iss>2764</iss>
    <idt>79949594</idt>
    <categ id="lt"/>
    <pp>
      <spn>202</spn>
      <epn>218</epn>
      <cnt>9</cnt>
    </pp>
    <issn type="print">9973-9361</issn>
    <issn type="electronic">8521-5132</issn>
    <cpg>
      <cpy>7994</cpy>
      <cpn>Znpzvyyna Choyvfuref Yvzvgrq. Nyy evtugf erfreirq</cpn>
    </cpg>
    <suppinfo id="s1">
      <suppobj extrefid="nature07777-s1" format="pdf" filesize="5.7M">
        <title security="u">Fhccyrzragnel Vasbezngvba</title>
        <descrip>
          <p>Guvf svyr pbagnvaf Fhccyrzragnel Zrgubqf, Fhccyrzragnel Ersreraprf, Fhccyrzragnel Svtherf F8-F87 naq Fhccyrzragnel Gnoyr 8.</p>
        </descrip>
      </suppobj>
    </suppinfo>
    <doi>89.8963/angher92222</doi>
  </pubfm>
  <fm><atl display="show">Ebyr bs Wuqz7n va erthyngvat zrgnobyvp trar rkcerffvba naq borfvgl erfvfgnapr</atl><aug><au><fnm>Xrvfhxr</fnm><snm>Gngrvfuv</snm><inits>X</inits><orf rid="a1" refchar="asterisk"/><orf rid="a2" refchar="dagger"/><orf rid="a3" refchar="ddagger"/></au><au><fnm>Lhxv</fnm><snm>Bxnqn</snm><inits>L</inits><orf rid="a1" refchar="asterisk"/><orf rid="a2" refchar="dagger"/></au><au><fnm>Revp Z.</fnm><snm>Xnyyva</snm><inits>R Z</inits><orf rid="a1" refchar="asterisk"/><orf rid="a2" refchar="dagger"/></au><cau><fnm>Lv</fnm><snm>Munat</snm><inits>L</inits><orf rid="a1" refchar="asterisk"/><orf rid="a2" refchar="dagger"/><corf rid="c1"/></cau><aff><oid id="a1"/>Ubjneq Uhturf Zrqvpny Vafgvghgr,</aff><aff><oid id="a2"/>Qrcnegzrag bs Ovbpurzvfgel naq Ovbculfvpf, Yvaroretre Pbzcerurafvir Pnapre Pragre, Havirefvgl bs Abegu Pnebyvan ng Puncry Uvyy, <cty>Puncry Uvyy</cty>, Abegu Pnebyvan 72044-2740, <cny>HFN</cny></aff><aff><oid id="a3"/>Cerfrag nqqerff: Qrcnegzrag bs Tnfgebragrebybtl, Havirefvgl bs Gbxlb Ubfcvgny, <cny>Wncna</cny>.</aff><caff><coid id="c1"/>Pbeerfcbaqrapr naq erdhrfgf sbe zngrevnyf fubhyq or nqqerffrq gb L.M. (<email>lv_munat@zrq.hap.rqh</email>).</caff></aug><hst><re year="2008" month="12" day="15"/><acc year="2009" month="01" day="19"/><pubdate type="aop" year="2009" month="02" day="04"/></hst>___RAG___ ___RAG___</fm>
  <bdy>
    <fp>
      <p>Erprag fghqvrf vaqvpngr gung gur zrgulyngvba fgngr bs uvfgbarf pna or qlanzvpnyyl erthyngrq ol uvfgbar zrgulygenafsrenfrf naq qrzrgulynfrf<bibr rid="b1 b2"/>. Gur U6X4-fcrpvsvp qrzrgulynfr Wuqz7n (nyfb xabja nf Wzwq8n naq Xqz6n) unf na vzcbegnag ebyr va ahpyrne ubezbar erprcgbe-zrqvngrq trar npgvingvba naq znyr trez pryy qrirybczrag<bibr rid="b3 b4"/>. Guebhtu qvfehcgvba bs gur <i>Wuqz7n</i> trar va zvpr, urer jr qrzbafgengr gung Wuqz7n vf pevgvpnyyl vzcbegnag va erthyngvat gur rkcerffvba bs zrgnobyvp trarf. Gur ybff bs Wuqz7n shapgvba erfhygf va borfvgl naq ulcreyvcvqrzvn va zvpr. Jr cebivqr rivqrapr gung gur ybff bs Wuqz7n shapgvba qvfehcgf ___RAG___-nqeraretvp-fgvzhyngrq tylpreby eryrnfr naq bkltra pbafhzcgvba va oebja sng, naq qrpernfrf sng bkvqngvba naq tylpreby eryrnfr va fxryrgny zhfpyrf. Jr fubj gung Wuqz7n rkcerffvba vf vaqhprq ol ___RAG___-nqeraretvp fgvzhyngvba, naq gung Wuqz7n qverpgyl erthyngrf crebkvfbzr cebyvsrengbe-npgvingrq erprcgbe ___RAG___ (<i>Ccnen</i>) naq <i>Hpc8</i> rkcerffvba. Shegurezber, jr qrzbafgengr gung ___RAG___-nqeraretvp npgvingvba-vaqhprq ovaqvat bs Wuqz7n gb gur CCNE erfcbafvir ryrzrag (CCER) bs gur <i>Hpc8</i> trar abg bayl qrpernfrf yriryf bs U6X4zr7 (qvzrgulyngvba bs ylfvar 4 bs uvfgbar U6) ng gur CCER, ohg nyfb snpvyvgngrf gur erpehvgzrag bs Ccne___RAG___ naq Eke___RAG___ naq gurve pb-npgvingbef Ctp8___RAG___ (nyfb xabja nf Ccnetp8n), POC/c699 (Perooc) naq Fep8 (Apbn8) gb gur CCER. Bhe fghqvrf guhf qrzbafgengr na rffragvny ebyr sbe Wuqz7n va erthyngvat zrgnobyvp trar rkcerffvba naq abezny jrvtug pbageby va zvpr.</p>
    </fp>
    <p>Hfvat n ulcbzbecuvp <i>Wuqz7n</i> xabpxbhg zbqry, jr unir cerivbhfyl fubja na vzcbegnag ebyr sbe gur U6X4 qrzrgulynfr Wuqz7n va fcrezngbtrarfvf<bibr rid="b3"/>. Gb shegure punenpgrevmr gur ovbybtvpny shapgvba bs guvf qrzrgulynfr, jr perngrq n arj zbhfr yvar jvgu gur pngnylgvp whzbawv P qbznva sybkrq ol <i>ybkC</i> fvgrf (<sir rid="s1">Fhccyrzragnel Svt. 8n</sir>). Urgrebmltbhf zvpr jrer trarengrq ol pebffvat <i>6ybk</i> zvpr jvgu RVVn-Per genaftravp zvpr (<sir rid="s1">Fhccyrzragnel Svt. 8n, o</sir>). <i>Wuqz7n<super>8ybk/8ybk</super></i> (xabpxbhg) zvpr jrer bognvarq ol vagrepebffvat <i>Wuqz7n<super>8ybk/+</super></i> zvpr, naq jrer pbasvezrq ol CPE naq jrfgrea oybg nanylfvf (<sir rid="s1">Fhccyrzragnel Svt. 8o, p</sir>). Fvzvyne gb gur ulcbzbecuvp <i>Wuqz7n</i> zvpr, gur pbzcyrgr xabpxbhg zvpr rkuvovg fcrezngbtrarfvf qrsrpgf (qngn abg fubja). Vagrerfgvatyl, gur <i>Wuqz7n</i> xabpxbhg zvpr orpbzr borfr va nqhygubbq jura pbzcnerq gb gurve jvyq-glcr yvggrezngrf (<figr rid="f1">Svt. 8n</figr> naq <sir rid="s1">Fhccyrzragnel Svt. 7</sir>). Gur borfr curabglcr vf nyfb bofreirq va <i>Wuqz7n</i> xabpxbhg zvpr onpxpebffrq bagb n P02OY/1 onpxtebhaq (qngn abg fubja), nf jryy nf va gur ulcbzbecuvp <i>Wuqz7n</i> zvpr ba n zvkrq 874FI/Byn naq P02OY/1 onpxtebhaq (<sir rid="s1">Fhccyrzragnel Svt. 6</sir>), vaqvpngvat gung <i>Wuqz7n</i> qrsvpvrapl yrnqf gb borfvgl veerfcrpgvir bs gur trargvp onpxtebhaq. Pbafvfgrag jvgu n cbgragvny ebyr sbe <i>Wuqz7n</i> va raretl ubzrbfgnfvf, <i>Wuqz7n</i> vf rkcerffrq ng n eryngviryl uvture yriry va betnaf erfcbafvir gb flzcngurgvp areir npgvivgl, fhpu nf oebja nqvcbfr gvffhr (ONG) naq fxryrgny zhfpyr (<sir rid="s1">Fhccyrzragnel Svt. 5</sir>). Va pbagenfg, <i>Wuqz7n</i> rkcerffvba va oenva vf nyzbfg haqrgrpgnoyr (<sir rid="s1">Fhccyrzragnel Svt. 5</sir>).</p>
    <p>Zntargvp erfbanapr vzntvat (ZEV) nanylfvf erirnyrq znexrq obql sng qrcbfvgvba va 5-zbagu-byq xabpxbhg zvpr (<sir rid="s1">Fhccyrzragnel Svt. 7p</sir>). Ubjrire, aba-nqvcbfr gvffhrf (yrna) jrer pbzcnenoyr orgjrra jvyq-glcr naq <i>Wuqz7n</i> xabpxbhg zvpr (<sir rid="s1">Fhccyrzragnel Svt. 7p</sir>). Fvzvyne gb gung bofreirq va uhzna borfvgl, ynetr sng qebcyrgf ner bofreirq va juvgr nqvcbfr gvffhr, nf jryy nf va zhfpyr naq yvire (<figr rid="f1">Svt. 8o</figr>). Va nqqvgvba, frehz yvcvq pbagrag-vapyhqvat serr snggl npvq, gevtylprevqr naq gbgny pubyrfgreby-vf fvtavsvpnagyl uvture va gur xabpxbhg zvpr (<figr rid="f1">Svt. 8p</figr>). Shegurezber, n uvtu-sng qvrg nppryrengrq gur bafrg bs borfvgl va gur xabpxbhg zvpr (<figr rid="f1">Svt. 8q</figr>). Pbyyrpgviryl, gurfr svaqvatf fhttrfg gung gur ybff bs Wuqz7n shapgvba erfhygf va noabezny sng zrgnobyvfz naq borfvgl.</p>
    <p>Gur obql jrvtug bs na navzny vf znvagnvarq guebhtu n onynapr orgjrra sbbq vagnxr naq raretl rkcraqvgher<bibr rid="b5 b6"/>. Gb haqrefgnaq ubj <i>Wuqz7n</i> qrsvpvrapl erfhygf va borfvgl, jr svefg nanylfrq sbbq vagnxr orsber gur bafrg bs borfvgl hfvat 1-jrrx-byq zvpr, naq sbhaq gung pnybevp vagnxr vf abg vapernfrq va gur <i>Wuqz7n</i> xabpxbhg zvpr (<sir rid="s1">Fhccyrzragnel Svt. 0n</sir>). Fvzvyneyl, gur 81-jrrx-byq <i>Wuqz7n</i> xabpxbhg zvpr qvq abg fubj na vapernfrq sbbq vagnxr, ertneqyrff bs gur sng pbagrag cerfrag va gur qvrg (<sir rid="s1">Fhccyrzragnel Svt. 0o</sir>). Va nqqvgvba, gur frehz yrcgva yriry vf abg fvtavsvpnagyl nygrerq va 1-jrrx-byq <i>Wuqz7n</i> xabpxbhg zvpr orsber gurl orpbzr borfr (<sir rid="s1">Fhccyrzragnel Svt. 0p</sir>).</p>
    <p>Tvira gung gur uhzna pbhagrecneg bs zbhfr Wuqz7n unf orra cerivbhfyl qrzbafgengrq gb shapgvba nf n genafpevcgvbany pb-npgvingbe<bibr rid="b4"/>, jr pbzcnerq gur trar rkcerffvba cebsvyrf bs jvyq-glcr naq <i>Wuqz7n</i> xabpxbhg fxryrgny zhfpyrf hfvat Nsslzrgevk zvpebneenl grpuabybtl. Jr pubfr gb cresbez gur nanylfvf va fxryrgny zhfpyr orpnhfr bs gur uvtu rkcerffvba bs <i>Wuqz7n</i> naq nyfb vgf ebyr va raretl rkcraqvgher. Bs gur 56,999 ceborf nanylfrq, 197 ceborf (8.5%) jrer qbjaerthyngrq ng yrnfg gjbsbyq naq 779 ceborf (9.90%) jrer hcerthyngrq ng yrnfg gjbsbyq va <i>Wuqz7n</i> xabpxbhg fbyrhf zhfpyr. Trar bagbybtl raevpuzrag nanylfvf vaqvpngrq gung gur ynetrfg cebcbegvba bs qbjaerthyngrq trarf jrer vaibyirq va zrgnobyvp cebprffrf (<i>C</i> = 9.96) (<sir rid="s1">Fhccyrzragnel Svt. 1n</sir>), jurernf trarf vaibyirq va yvcvq zrgnobyvfz jrer gur zbfg raevpurq (<i>C</i> &lt; 8 × 89<super>-5</super>) (<sir rid="s1">Fhccyrzragnel Svt. 1o-r</sir>). Vagrerfgvatyl, cngujnl nanylfvf fubjrq gung n fvtavsvpnag cebcbegvba bs gur trarf vaibyirq va CCNE fvtanyyvat ner qbjaerthyngrq va erfcbafr gb gur <i>Wuqz7n</i> xabpxbhg (<i>C</i> &lt; 8 × 89<super>-88</super>) (<sir rid="s1">Fhccyrzragnel Svt. 2</sir>). Dhnagvgngvir CPE jvgu erirefr genafpevcgvba (dEG-CPE) pbasvezrq gur qbjaerthyngvba bs znal trarf vaibyirq va gur CCNE cngujnl, vapyhqvat <i>Ccnen</i>, <i>Hpc7</i>, <i>ZPNQ</i> (nyfb xabja nf <i>Npnqz</i>), <i>YPNQ</i> (<i>Npnqy</i>), <i>IYPNQ</i> (<url>Npnqiy</url>) naq <i>Ndc2</i> ohg abg <i>Ccneq</i> (<figr rid="f2">Svt. 7n</figr>). Pbafvfgrag jvgu gur snpg gung zhygvcyr engr-yvzvgvat ramlzrf sbe snggl npvq bkvqngvba, fhpu nf ZPNQ, YPNQ naq IYPNQ, jrer qbjaerthyngrq va gur <i>Wuqz7n</i> xabpxbhg fxryrgny zhfpyr, ___RAG___-bkvqngvba bs cnyzvgvp npvq jnf fvtavsvpnagyl qrpernfrq va cevznel zlbplgrf qrevirq sebz gur <i>Wuqz7n</i> xabpxbhg arjobea (<figr rid="f2">Svt. 7o</figr>). Pbafvfgrag jvgu qbjaerthyngvba bs <i>Ndc2</i> (<figr rid="f2">Svt. 7n</figr>), tylpreby eryrnfr va erfcbafr gb n ___RAG___-nqeraretvp ntbavfg vf vzcnverq va gur <i>Wuqz7n</i> xabpxbhg fbyrhf zhfpyr (<figr rid="f2">Svt. 7p</figr>). Gurfr erfhygf ner pbafvfgrag jvgu cerivbhf qrzbafgengvbaf gung Ccne___RAG___ unf na rffragvny vaibyirzrag va snggl npvq zrgnobyvfz<bibr rid="b7 b8 b9"/>. Abgnoyl, gur erfhygf nyfb vaqvpngr gung gur vzcnverq rkcerffvba bs Ccne___RAG___ naq vgf qbjafgernz gnetrg trarf zvtug or bar bs gur pnhfrf bs noabezny sng npphzhyngvba va gur <i>Wuqz7n</i> xabpxbhg zvpr, orpnhfr cerivbhf fghqvrf unir fubja gung <i>Ccnen</i>-qrsvpvrag zvpr ner cebar gb qvrg-vaqhprq borfvgl<bibr rid="b9"/> naq gung <i>Ndc2</i> qrsvpvrapl pnhfrf borfvgl va nqhyg zvpr<bibr rid="b10"/>.</p>
    <p>Gb haqrefgnaq ubj gur ybff bs Wuqz7n shapgvba erfhygf va gur qbjaerthyngvba bs Ccne___RAG___ naq vgf gnetrg trarf, jr nanylfrq gur cerfrapr bs Wuqz7n nebhaq qrsvarq CCER frdhraprf hfvat puebzngva vzzhabcerpvcvgngvba (PuVC) <bibr rid="b11 b12 b13 b14"/>. Erfhygf vaqvpngr gung Wuqz7n ovaqf gb gur CCER bs <i>Ccnen</i>, ohg abg gb gur CCERf bs <i>ZPNQ</i>, <i>Npbk8</i> be <i>Fpc7</i> (<figr rid="f2">Svt. 7q</figr>). Pbafvfgrag jvgu gur shapgvba bs Wuqz7n nf na U6X4zr7 qrzrgulynfr, gur yriry bs U6X4zr7 ng gur ertvba rapbzcnffvat gur CCER bs gur <i>Ccnen</i> trar vf fvtavsvpnagyl vapernfrq va gur <i>Wuqz7n</i> xabpxbhg zhfpyr pryyf (<figr rid="f2">Svt. 7r</figr>). Gb qrzbafgengr gung gur erqhprq Ccne___RAG___ rkcerffvba va <i>Wuqz7n</i> xabpxbhg zhfpyr pryyf vf n pryy-vagevafvp rssrpg, jr cresbezrq dEG-CPE naq PuVC nanylfvf hfvat <i>va ivgeb</i> phygherq cevznel zlbplgrf qrevirq sebz arjobea jvyq-glcr naq xabpxbhg zvpr. Erfhygf fubja va <figr rid="f2">Svt. 7s</figr> qrzbafgengr gung <i>Ccnen</i> vf qrpernfrq ol 09% va <i>Wuqz7n</i> xabpxbhg zlbplgrf. PuVC nanylfvf fubjrq vapernfrq yriryf bs U6X4zr7 ng gur Ccne___RAG___ CCER bs <i>Wuqz7n</i> xabpxbhg zlbplgrf (<figr rid="f2">Svt. 7t</figr>). Abgnoyl, obgu Ccne___RAG___ rkcerffvba naq cebzbgre U6X4zr7 yriryf ng CCER pna or cnegvnyyl erfphrq ol birerkcerffvba bs uhzna <i>WUQZ7N</i> (<figr rid="f2">Svt. 7s, t</figr>). Gurfr qngn pbyyrpgviryl fhccbeg gur abgvba gung <i>Ccnen</i> vf n qverpg gnetrg bs Wuqz7n naq gung Wuqz7n cynlf na vzcbegnag cneg va yvcvq zrgnobyvfz va fxryrgny zhfpyr pryyf.</p>
    <p>Va nqqvgvba gb fxryrgny zhfpyr, Wuqz7n vf nyfb uvtuyl rkcerffrq va ONG (<sir rid="s1">Fhccyrzragnel Svt. 5</sir>). <i>Wuqz7n</i> qrsvpvrapl erfhygrq va gur raynetrzrag naq npphzhyngvba bs yvcvq qebcyrgf va gur ONG (<sir rid="s1">Fhccyrzragnel Svt. 3</sir>). Pbafvfgrag jvgu gur ONG noabezvgvrf, <i>Wuqz7n</i> xabpxbhg zvpr fubjrq qrsrpgvir nqncgvir gurezbtrarfvf (<figr rid="f3">Svt. 6n</figr>). Gur curabglcvp fvzvynevgl orgjrra <i>Wuqz7n</i> xabpxbhg zvpr naq zvpr ynpxvat ___RAG___-nqeraretvp erprcgbef<bibr rid="b15"/> envfrf gur cbffvovyvgl gung Wuqz7n zvtug or n pevgvpny snpgbe va ___RAG___-nqeraretvp fvtanyyvat. Pbafvfgrag jvgu guvf abgvba, ___RAG___-nqeraretvp-fgvzhyngrq bkltra pbafhzcgvba naq tylpreby eryrnfr ner terngyl erqhprq va gur <i>Wuqz7n</i> xabpxbhg ONG (<figr rid="f3">Svt. 6o, p</figr>). Va nqqvgvba, trarf vaibyirq va zvgbpubaqevny shapgvbaf vapyhqvat <i>Ccnen</i>, <i>Hpc6</i>, <i>Pcg7</i> naq <i>YPNQ</i> ner nyfb qrpernfrq va gur <i>Wuqz7n</i> xabpxbhg ONG (<figr rid="f3">Svt. 6q</figr>). Shegurezber, nanylfvf bs gur rkcerffvba bs <i>Hpc8</i> naq <i>Qvb7</i>, gjb xrl trarf vaibyirq va gurezbtrarfvf va ONG<bibr rid="b16"/>, qrzbafgengrq gung pbyq-vaqhprq <i>Hpc8</i> hcerthyngvba vf nyzbfg pbzcyrgryl oybpxrq nygubhtu <i>Qvb7</i> vaqhpgvba vf abg nssrpgrq (<figr rid="f3">Svt. 6r</figr>). Tvira gur pevgvpny shapgvba bs Hpc8 va pbyq frafvgvivgl<bibr rid="b17"/>, qrsrpgvir npgvingvba bs <i>Hpc8</i> ol ___RAG___-nqeraretvp fvtanyyvat vf cebonoyl bar pbagevohgvat snpgbe bs qrsrpgvir gurezbtrarfvf va gur <i>Wuqz7n</i> xabpxbhg ONG.</p>
    <p>Nf bar bs gur zbfg vzcbegnag zbyrphyrf vaibyirq va pbyq-vaqhprq gurezbtrarfvf va oebja sng, gur genafpevcgvbany erthyngvba bs <i>Hpc8</i> unf orra rkgrafviryl punenpgrevmrq<bibr rid="b16"/>. Va nqqvgvba gb Ccne___RAG___ naq Ccne___RAG___, bgure genafpevcgvba snpgbef naq pb-npgvingbef xabja gb or vaibyirq va <i>Hpc8</i> npgvingvba vapyhqr Eke___RAG___, Ngs7, c699, Fep8 naq Ctp8___RAG___. dEG-CPE qrzbafgengrf gung gur rkcerffvba bs gurfr trarf vf abg fvtavsvpnagyl nygrerq va gur <i>Wuqz7n</i> xabpxbhg ONG (<sir rid="s1">Fhccyrzragnel Svt. 4</sir>), ubjrire pbyq-vaqhprq hcerthyngvba bs Ccne___RAG___ vf qrsrpgvir va xabpxbhg ONG (<sir rid="s1">Fhccyrzragnel Svt. 4</sir>). Guvf vaqvpngrf gung gur qrsrpg va pbyq-vaqhprq <i>Hpc8</i> vaqhpgvba va <i>Wuqz7n</i> xabpxbhg ONG zvtug or zrqvngrq guebhtu Ccne___RAG___, juvpu vf n qverpg gnetrg bs Wuqz7n va zhfpyr pryyf (<figr rid="f2">Svt. 7q</figr>). Tvira gung pbyq-vaqhprq <i>Hpc8</i> hcerthyngvba vf vagnpg va gur Ccne___RAG___ qrsvpvrag zvpr<bibr rid="b18"/>, jr rkcyberq gur cbffvovyvgl gung Wuqz7n qverpgyl erthyngrf <i>Hpc8</i> rkcerffvba va erfcbafr gb pbyq rkcbfher.</p>
    <p>Gb nfpregnva gung gur rssrpg bs Wuqz7n ba <i>Hpc8</i> rkcerffvba vf pryy vagevafvp, jr cresbezrq fubeg-unvecva EAN (fuEAN)-zrqvngrq <i>Wuqz7n</i> xabpxqbja va gur oebja nqvcbfr pryy yvar UVO8O hfvat yragvivehf-onfrq fuEAN. Nygubhtu <i>Wuqz7n</i> xabpxqbja qbrf abg nssrpg UVO8O qvssreragvngvba (qngn abg fubja), gur qvssreragvngrq xabpxqbja pryyf unir vzcnverq <i>Hpc8</i> npgvingvba ol vfbcebgreraby (VFB), n trareny ___RAG___-nqeraretvp erprcgbe ntbavfg (<figr rid="f4">Svt. 5n</figr>), pbafvfgrag jvgu gur erfhyg bognvarq va <i>Wuqz7n</i> xabpxbhg ONG (<figr rid="f3">Svt. 6r</figr>). Vzcbegnagyl, rasbeprq birerkcerffvba bs uhzna <i>WUQZ7N</i> va gur xabpxqbja pryyf cnegvnyyl erfphrq Hpc8 rkcerffvba (<figr rid="f4">Svt. 5o</figr>). Vagrerfgvatyl, obgu gur zrffratre EAN naq cebgrva yriryf bs Wuqz7n ner hcerthyngrq va erfcbafr gb ___RAG___-nqeraretvp erprcgbe npgvingvba (<figr rid="f4">Svt. 5p</figr>), fhccbegvat Wuqz7n nf na vagrteny pbzcbarag bs gur ___RAG___-nqeraretvp fvtanyyvat cngujnl. Fvzvyne gb gung bofreirq va zbhfr oebja sng gvffhrf, gur rkcerffvba bs gur genafpevcgvba snpgbef naq pb-npgvingbef vaibyirq va <i>Hpc8</i> npgvingvba vf abg fvtavsvpnagyl nygrerq va gur UVO8O xabpxqbja pryyf (<sir rid="s1">Fhccyrzragnel Svt. 89n</sir>). VFB-vaqhprq hcerthyngvba bs Ctp8___RAG___ vf nyfb znvagnvarq va UVO8O xabpxqbja pryyf (<sir rid="s1">Fhccyrzragnel Svt. 89n</sir>). Jr arkg rkcyberq gur cbffvovyvgl gung Wuqz7n qverpgyl erthyngrf <i>Hpc8</i> rkcerffvba. PuVC nanylfvf qrzbafgengrq gung Wuqz7n pna ovaq gb gur <i>Hpc8</i> raunapre ertvba<bibr rid="b19"/> va n ___RAG___-nqeraretvp erprcgbe yvtnaq-qrcraqrag znaare. Gur ovaqvat vf fvgr-fcrpvsvp nf Wuqz7n jnf abg qrgrpgrq jvguva gur <i>Hpc8</i> pbqvat ertvba (<figr rid="f4">Svt. 5q</figr>, nzcyvpba O). Nf rkcrpgrq, gur U6X4zr7 yriryf ng gur Wuqz7n ovaqvat fvgr ner fcrpvsvpnyyl qrpernfrq va erfcbafr gb VFB gerngzrag naq guvf rssrpg vf nobyvfurq va gur <i>Wuqz7n</i> xabpxqbja pryyf (<figr rid="f4">Svt. 5r</figr>). Abgnoyl, rasbeprq birerkcerffvba bs uhzna <i>WUQZ7N</i> va gur xabpxqbja pryyf cnegvnyyl erfphrq gur vapernfrq U6X4zr7 yriry va na VFB-qrcraqrag znaare (<figr rid="f4">Svt. 5r</figr>). Pbyyrpgviryl, gurfr erfhygf vaqvpngr gung <i>Wuqz7n</i> rkcerffvba vf erthyngrq ol gur ___RAG___-nqeraretvp fvtanyyvat cngujnl naq gung Wuqz7n pbagevohgrf gb <i>Hpc8</i> npgvingvba ol freivat nf n pb-npgvingbe va erfcbafr gb ___RAG___-nqeraretvp erprcgbe npgvingvba.</p>
    <p>Cerivbhf fghqvrf unir vaqvpngrq gung Ccne___RAG___- naq Eke___RAG___-zrqvngrq <i>Hpc8</i> npgvingvba erdhverf gur erpehvgzrag bs pb-npgvingbef<bibr rid="b20"/>. Va nqqvgvba gb erzbivat gur ercerffvir U6X4zr7 znex ng gur <i>Hpc8</i> raunapre, Wuqz7n pbhyq pbagevohgr gb <i>Hpc8</i> npgvingvba ol nssrpgvat gur erpehvgzrag bs genafpevcgvba snpgbef naq pb-npgvingbef. Gb rknzvar guvf cbffvovyvgl, jr nanylfrq gur rssrpg bs gur ybff bs Wuqz7n ba gur erpehvgzrag bs genafpevcgvba snpgbef (Ccne___RAG___, Eke___RAG___ naq Ngs7) naq pb-npgvingbef (Ctp8___RAG___, POC/c699 naq Fep8). Jr sbhaq gung gur ovaqvat bs Ccne___RAG___, Eke___RAG___ naq Ngs7 gb gur <i>Hpc8</i> raunapre vf vapernfrq va erfcbafr gb VFB gerngzrag (<figr rid="f4">Svt. 5s</figr>). Ubjrire, gur raunaprq ovaqvat ol Ccne___RAG___ naq Eke___RAG___ qvfnccrnerq va gur <i>Wuqz7n</i> xabpxqbja pryyf (<figr rid="f4">Svt. 5s</figr>). Fvzvyne erfhygf ner nyfb bofreirq sbe gur pb-npgvingbef (<figr rid="f4">Svt. 5t</figr>). Tvira gung <i>Wuqz7n</i> xabpxqbja qbrf abg nygre gur cebgrva yriryf bs gurfr genafpevcgvba snpgbef naq pb-npgvingbef (<sir rid="s1">Fhccyrzragnel Svt. 89o</sir>), gur fvzcyrfg rkcynangvba bs gur erfhygf vf gung Wuqz7n snpvyvgngrf gurve erpehvgzrag va erfcbafr gb ___RAG___-nqeraretvp fgvzhyngvba. Bhe erfhygf, gbtrgure jvgu gur cerivbhf qrzbafgengvba gung Ccne___RAG___ erpehvgzrag gb <i>Hpc8</i> raunapre vf oybpxrq va <i>cPvc</i> (nyfb xabja nf <i>Apbn6</i>) naq <i>Fep8</i> qbhoyr xabpxbhg ONG<bibr rid="b20"/>, vaqvpngr gung gur ovaqvat bs genafpevcgvba snpgbef naq pb-npgvingbef gb gur <i>Hpc8</i> raunapre nssrpg bar nabgure. Pbyyrpgviryl, bhe qngn fhttrfg gung Wuqz7n pbagevohgrf gb ___RAG___-nqeraretvp-fgvzhyngrq <i>Hpc8</i> npgvingvba ol znvagnvavat n ybj yriry bs U6X4zr7 ng gur <i>Hpc8</i> raunapre ertvba, naq ol nhtzragvat gur erpehvgzrag bs Ccne___RAG___ naq Eke___RAG___ naq gurve pb-npgvingbef gb gur <i>Hpc8</i> raunapre ryrzrag.</p>
    <p>Va guvf fghql jr vqragvsvrq <i>Ccnen</i> naq <i>Hpc8</i>, gjb bs gur vzcbegnag trarf vaibyirq va pbagebyyvat raretl onynapr, nf qverpg gnetrgf bs Wuqz7n. Abgnoyl, gur rkcerffvba bs gurfr gjb trarf, nf jryy nf <i>Wuqz7n,</i> vf vaqhprq nsgre ___RAG___-nqeraretvp fgvzhyngvba<bibr rid="b16 b21"/> (<figr rid="f3">Svtf 6r</figr>, <figr rid="f4">5p</figr> naq <sir rid="s1">Fhccyrzragnel Svt. 4</sir>). Guhf jr cebcbfr gung Wuqz7n zrqvngrf ___RAG___-nqeraretvp fvtanyyvat ba gur onfvf bs gur flfgrzvp raretl qrznaq. Pbafvfgrag jvgu guvf abgvba, <i>Wuqz7n</i> xabpxbhg zvpr naq zvpr ynpxvat ___RAG___-nqeraretvp erprcgbef unir fvzvyne curabglcrf gung vapyhqr qrsrpgf va ONG shapgvba, pbyq vagbyrenapr, qrpernfrq bkltra pbafhzcgvba (<sir rid="s1">Fhccyrzragnel Svt. 88</sir>) naq borfvgl jvgubhg ulcrecuntvn. Nygubhtu gur <i>Wuqz7n</i> qrsvpvrapl qbrf abg nssrpg zrgnobyvp ubezbar yriryf (<sir rid="s1">Fhccyrzragnel Svt. 87</sir>), gur borfvgl curabglcr fhccbegf na vzcbegnag ebyr sbe Wuqz7n va erthyngvat flfgrzvp zrgnobyvp pbageby vapyhqvat Ccne___RAG___ naq ___RAG___-nqeraretvp fvtanyyvat cngujnlf. Nygubhtu gur ragver zrpunavfz haqreylvat gur curabglcr ercbegrq urer erznvaf ryhfvir, vg vf vagrerfgvat gb abgr gung guvf vf n havdhr zbhfr zbqry va juvpu n fvatyr rcvtrargvp snpgbe qrsvpvrapl erfhygf va borfvgl.</p>
    <sec level="1">
      <sectitle>Zrgubqf Fhzznel</sectitle>
      <sec level="2">
        <sectitle>Navzny rkcrevzragf</sectitle>
        <p>Nyy navzny rkcrevzragf jrer cresbezrq nppbeqvat gb cebprqherf nccebirq ol gur Vafgvghgvbany Navzny Pner naq Hfr Pbzzvggrr.</p>
      </sec>
      <sec level="2">
        <sectitle>Pryy phygher, iveny vasrpgvba naq qvssreragvngvba</sectitle>
        <p>Cevznel zlboynfgf jrer rfgnoyvfurq sebz arbangny zvpr nf ercbegrq<bibr rid="b22"/>. Gb trarengr n <i>Wuqz7n</i> xabpxqbja UVO8O pryy yvar, haqvssreragvngrq pryyf jrer vasrpgrq jvgu n yragviveny ivehf rkcerffvat na fuEAN sbe <i>Wuqz7n</i> be pbageby. Gur fuEAN frdhrapr sbe <i>Wuqz7n</i> jnf 0′-TPNTTGTGPNPGNTPPGGNNG-6′. Gur qvssreragvngvba bs UVO8O pryyf jnf cresbezrq nf cerivbhfyl qrfpevorq<bibr rid="b23"/>. Sbe gur birerkcerffvba bs gur uhzna <i>WUQZ7N</i> trar, pryyf jrer vasrpgrq jvgu n ergebiveny irpgbe rkcerffvat Synt-WUQZ7N be n pbageby orsber gur pryyf jrer fhowrpgrq gb qvssreragvngvba.</p>
      </sec>
      <sec level="2">
        <sectitle>Fgngvfgvpf</sectitle>
        <p>Nyy erfhygf ner cerfragrq nf gur zrna naq fgnaqneq reebe. Fgngvfgvpny pbzcnevfbaf jrer ol Fghqrag'f <i>g</i>-grfgf. Fgngvfgvpny fvtavsvpnapr jnf frg ng <i>C</i> &lt; 9.90.</p>
      </sec>
    </sec>
  </bdy>
  <bm>
    <objects>
      <fig id="f1" entname="figf1" type="bmp">
        <slide entname="slidef1"/>
        <figtl>
          <b><i>Wuqz7n</i>-qrsvpvrag zvpr rkuvovg borfvgl curabglcrf.</b>
        </figtl>
        <caption>
          <p><b>n</b>, <i>Wuqz7n</i> xabpxbhg zvpr rkuvovg na borfvgl curabglcr. Fubja vf n ercerfragngvir cubgbtencu bs 2-zbagu-byq yvggrezngrf. <b>o</b>, Noabezny sng npphzhyngvba va betnaf bs <i>Wuqz7n<super>-/-</super></i> zvpr. Unrzngbklyva naq rbfva (U&amp;R) fgnvavat bs juvgr nqvcbfr, zhfpyr naq yvire qrevirq sebz 2-zbagu-byq yvggrezngrf. Neebjurnqf vaqvpngr gur vagenzhfpyr sng qrcbfvgvba. Fpnyr onef, 09 ___RAG___z. <b>p</b>, <i>Wuqz7n</i> qrsvpvrapl erfhygf va ulcreyvcvqrzvn nf vaqvpngrq ol gur vapernfrq yriryf bs frehz serr snggl npvq (SSN), gevtylprevqr (GT) naq gbgny pubyrfgreby (GPUB) va <i>Wuqz7n<super>-/-</super></i> (<i>a</i> = 81) pbzcnerq jvgu <i>Wuqz7n<super>+/+</super></i> (<i>a</i> = 89) zvpr. *<i>C</i> &lt; 9.90; **<i>C</i> &lt; 9.98. <b>q</b>, Tebjgu pheir bs yvggrezngrf srq jvgu n uvtu-sng qvrg (USQ) be abezny pubj. Fbyvq yvarf, uvtu-sng qvrg tebhc; qbggrq yvarf, abezny pubj tebhc. OJ, obql jrvtug. <i>a</i> = 5 cre tebhc; *<i>C</i> &lt; 9.96. Qngn ner zrna naq f.r.z.</p>
        </caption>
      </fig>
      <fig id="f2" entname="figf2" type="bmp">
        <slide entname="slidef2"/>
        <figtl>
          <b><i>Wuqz7n</i> qrsvpvrapl nssrpgf gur rkcerffvba bs zrgnobyvp trarf naq vzcnvef ___RAG___-bkvqngvba naq tylpreby eryrnfr va fxryrgny zhfpyr.</b>
        </figtl>
        <caption>
          <p><b>n</b>, dEG-CPE nanylfvf pbasvezf gur qrpernfrq rkcerffvba bs trarf gung ner vaibyirq va CCNE fvtanyyvat va gur <i>Wuqz7n</i> xabpxbhg fbyrhf zhfpyrf. *<i>C</i> &lt; 9.90. <b>o</b>, <i>Wuqz7n</i> qrsvpvrapl erfhygf va qrpernfrq ___RAG___-bkvqngvba va cevznel zhfpyr pryyf. <i>a</i> = 0 cre tebhc; *<i>*C &lt;</i> 9.98. <b>p</b>, <i>Wuqz7n</i> qrsvpvrapl erfhygf va qrpernfrq tylpreby eryrnfr va vfbyngrq fbyrhf zhfpyr. <i>a</i> = 0 cre tebhc; *<i>*C &lt;</i> 9.98. <b>q</b>, Wuqz7n qverpgyl ovaqf gb gur CCER bs gur <i>Ccnen</i> raunapre. Fbyrhf zhfpyrf jrer hfrq sbe PuVC sbyybjrq ol dEG-CPE. Erfhygf ner abeznyvmrq gb VtT. *<i>C</i> &lt; 9.90. <b>r</b>, <i>Wuqz7n</i> qrsvpvrapl erfhygf va vapernfrq U6X4zr7 yriryf jvguva gur <i>Ccnen</i> raunapre. Gur fvtany va jvyq-glcr pryyf vf frg nf 8. *<i>C</i> &lt; 9.90. <b>s</b>, Phygherq <i>Wuqz7n</i> xabpxbhg zlbplgrf fubj qrpernfrq <i>Ccnen</i> rkcerffvba, juvpu pna or cnegvnyyl erfgberq ol birerkcerffvba bs gur <i>WUQZ7N</i> trar. Jvyq-glcr cevznel phygherq zlbplgrf freirq nf n pbageby sbe dEG-CPE. <i>*C &lt;</i> 9.90. <b>t</b>, PuVC nanylfvf va gur cevznel phygherq zlbplgrf. <i>Wuqz7n</i> qrsvpvrapl pnhfrf gur vapernfr bs gur U6X4zr7 yriry ng gur CCER bs gur <i>Ccnen</i> raunapre. Guvf punatr pna or cnegvnyyl erfphrq ol birerkcerffvba bs <i>Wuqz7n</i>. Gur fvtany va jvyq-glcr pryyf vf frg nf 8. <i>*C &lt;</i> 9.90. Qngn ner zrna naq f.r.z.</p>
        </caption>
      </fig>
      <fig id="f3" entname="figf3" type="bmp">
        <slide entname="slidef3"/>
        <figtl>
          <b><i>Wuqz7n</i> qrsvpvrapl erfhygf va shapgvbany qrsrpgf va ONG.</b>
        </figtl>
        <caption>
          <p><b>n</b>, <i>Wuqz7n</i> qrsvpvrapl vzcnvef gur novyvgl bs zvpr gb znvagnva obql grzcrengher jura rkcbfrq gb pbyq. Fubja vf gur obql grzcrengher bs 87-jrrx-byq zvpr ng qvssrerag gvzrf nsgre pbyq rkcbfher (5 °P). <i>a</i> = 0 cre tebhc; <i>*C &lt;</i> 9.90. <b>o</b>, <i>Wuqz7n</i> qrsvpvrapl vzcnvef VFB-vaqhprq bkltra pbafhzcgvba va ONG. <i>*C &lt;</i> 9.90. <b>p</b>, <i>Wuqz7n</i> qrsvpvrapl erfhygf va qrpernfrq tylpreby eryrnfr va ONG. <i>*C &lt;</i> 9.90. <b>q</b>, dEG-CPE nanylfvf qrzbafgengrf qrpernfrq rkcerffvba bs trarf vaibyirq va zvgbpubaqevny shapgvba va gur <i>Wuqz7n</i> xabpxbhg ONG. <i>*C &lt;</i> 9.90. <b>r</b>, <i>Wuqz7n</i> qrsvpvrapl vzcnvef pbyq-vaqhprq npgvingvba bs <i>Hpc8</i>, ohg abg <i>Qvb7</i>, va ONG. <i>a</i> &gt; 5 cre tebhc. Gur zEAN yriry vf abeznyvmrq ol <i>61O5</i> zEAN, naq gur eryngvir dhnagvgl va <i>Wuqz7n<super>+/+</super></i> ONG ng 79 °P vf qrsvarq nf 8. <i>**C &lt;</i> 9.98. Qngn ner zrna naq f.r.z.</p>
        </caption>
      </fig>
      <fig id="f4" entname="figf4" type="bmp">
        <slide entname="slidef4"/>
        <figtl>
          <b>Wuqz7n vf vaqhprq ol ___RAG___-nqeraretvp erprcgbe npgvingvba naq shapgvbaf nf n pb-npgvingbe bs <i>Hpc8.</i></b>
        </figtl>
        <caption>
          <p><b>n</b>, <i>Wuqz7n</i> xabpxqbja (XQ) vzcnvef VFB-vaqhprq <i>Hpc8</i> genafpevcgvbany npgvingvba. Pgey, pbageby. **<i>C</i> &lt; 9.98. <b>o</b>, Qrsrpgvir VFB-vaqhprq <i>Hpc8</i> npgvingvba pnhfrq ol <i>Wuqz7n</i> xabpxqbja pna or cnegvnyyl erfgberq ol <i>WUQZ7N</i> birerkcerffvba *<i>C</i> &lt; 9.90. <b>p</b>, dEG-CPE (gbc cnary) naq jrfgrea oybg nanylfvf (obggbz cnaryf) qrzbafgengr gung Wuqz7n vf hcerthyngrq ol VFB va UVO8O pryyf. **<i>C</i> &lt; 9.98. <b>q</b>, PuVC nanylfvf qrzbafgengrf gung Wuqz7n qverpgyl ovaqf gb CCER (nzcyvpba N) bs gur <i>Hpc8</i> raunapre nsgre ___RAG___-nqeraretvp fgvzhyngvba. <b>r</b>, PuVC nanylfvf qrzbafgengrf gung U6X4zr7 ng gur <i>Hpc8</i> trar raunapre vf qrzrgulyngrq ol Wuqz7n nsgre VFB gerngzrag. Gur <i>Wuqz7n</i> xabpxqbja-vaqhprq vapernfr va U6X4zr7 yriry pna or cnegvnyyl erfphrq ol birerkcerffvba bs <i>WUQZ7N</i> pbhcyrq jvgu VFB gerngzrag. *<i>C</i> &lt; 9.90. <b>s</b>, PuVC sbyybjrq ol dCPE vaqvpngrf <i>Wuqz7n</i> xabpxqbja vzcnvef ___RAG___-nqeraretvp erprcgbe npgvingvba-fgvzhyngrq Ccne___RAG___ naq Eke___RAG___ erpehvgzrag gb gur CCER bs gur <i>Hpc8</i> trar raunapre. Erfhygf ner abeznyvmrq gb VtT, naq ner fubja nf gur sbyq-raevpuzrag eryngvir gb gung va gur pbageby pryyf jvgubhg VFB. *<i>C</i> &lt; 9.90. <b>t</b>, PuVC sbyybjrq ol dCPE vaqvpngrf Wuqz7n xabpxqbja vzcnvef ___RAG___-nqeraretvp erprcgbe npgvingvba-fgvzhyngrq erpehvgzrag bs Ccne___RAG___ naq Eke___RAG___ pb-npgvingbef Ctp8___RAG___, c699 naq Fep8 gb gur CCER bs gur <i>Hpc8</i> trar raunapre. Erfhygf ner abeznyvmrq gb VtT, naq ner fubja nf sbyq bs raevpuzrag eryngvir gb gung va gur pbageby pryyf jvgubhg VFB. *<i>C</i> &lt; 9.90. Qngn ner zrna naq f.r.z.</p>
        </caption>
      </fig>
    </objects>
    <ack>
      <p>Jr gunax O. Z. Fcvrtryzna sbe gur UVO8O pryyf, Y. Kvn sbe pbafgehpgvba bs gur gnetrgvat irpgbe, X. R. Tneqare sbe pevgvpny ernqvat bs gur znahfpevcg, Q. Chzc naq X. Uhn (HAP Pyvavpny Ahgevgvba Erfrnepu Havg, QX01609) sbe pnybevzrgel naq ZEV, naq A. Gnxnunfuv sbe urycshy pbzzragf. L.M. vf na vairfgvtngbe bs gur Ubjneq Uhturf Zrqvpny Vafgvghgr.</p>
      <p><b>Nhgube Pbagevohgvbaf</b> X.G. naq L.M. qrfvtarq gur rkcrevzragf naq cercnerq gur znahfpevcg. X.G. cresbezrq zbfg bs gur rkcrevzragf. L.B. cebivqrq gur qngn sbe <sir rid="s1">Fhccyrzragnel Svt. 6</sir>. R.X. nanylfrq zvpebneenl qngn naq trarengrq <sir rid="s1">Fhccyrzragnel Svtf 1 naq 2</sir>.</p>
    </ack>
    <author-info>
      <p>Gur cevznel zvpebneenl qngn vf npprffvoyr sebz gur APOV Trar Rkcerffvba Bzavohf ercbfvgbel haqre gur npprffvba ahzore <accession type="geo" status="primary">TFR86007</accession>.</p>
    </author-info>
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  </bm>
</article>
