Science Magazine 5817 2007-03-09
Science Magazine 5817 2007-03-09
Science Magazine 5817 2007-03-09
for your PC
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Usingla unique bl~end of thermostabte emzymes and a set of 12 :standardi.zed premixes, optimiizati,on ot any IPCR reaction is quick, 9 SSy and r,eprodl!Jciil~:).~e,. The Fa~ISaferM peR System Amlp~ifies"Up'to 85% GC~richtemplates .' ,Any templates lip to 200kb .' Multliph~x peiR, consiistenUy
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Arnj:liitl'Calitm ,at alii 8(1-85% GC-~ld1, 1egjgll of J If!8 hJJlJIIIiI '(rngf]e:,X gene. IF'GR MIS: pmwmed US[1lIJlhl fails.atll'" peR Syslu!l!.la!lcsA-L a.ow '1'tI1I2InnPlm~~oo prnwet5 r 8almn.Q 'ri'OOI IisinQ '~e ~21ia1lSaIe PCR II?reM~. rill, iiI101~l!l'~rwe~hI: lII'IiIJ'ker. (ll}1:lmal~m1,P!il~ 'Ibn was OOtalned wi:lb FaliSaf~ PCR PreMii!! J.
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use on any r,salll,-tim,e :PGRlnstnrnent, .' Low eli Values: high sel1ls~iUvity wlirth de·~eN·.:::t~onf as o 'mew as 10 coples .
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Bringing protein analysis to life with Etten DIIGIE ond Amershorn IE:CL
WI"J>(~n cernes to life sciences, GE Heolthcare is setting the it
stendord, lens of thousands of scienusts worldwide rely on our pro ducts 0 n d proven e>::perti se ln protei n o na lysis. and de'tecti on
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By contlnuolly developing technology thot con turn your scientific ideas into reaHty, V!Je'F@ bnngmg science to Ills' end he[pmg to
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We call it Protein Analysis Re-lrncqined, Discover the full power of Etten DIGEand the Amershorr: ECLrl'l family of Western blotting systems ot
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9 MARCHI
2007
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www,5dencexpress.org PILANT .sCIENOE A G i~iI"~till'n-'{llIIp,!ed Rel:llplor 115 <llfllils.D1'11 Membrane fior the PIl<llllt ~.jo[mol1e- AhSdiSic;,Add X. Liu;, Y. YUe', It t~ Y. Ni(!, w- U W.-H" WIt t. Alto f\ mlwl'b'r'iil n~-Dol:llldl rec~ItI'DO rlOl il~(isill: ~c,id i~, entlliect r' id IPlam!t R!~cep'OO~' ~ith IEl,i,~!Fr.amilly ~g
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PHI¥SICS Floorn"tll~mp~fatQjro QUaJl1iI:urn H~~11 ffect, i!'l'Graphll!n'~ 1379 E MATI~:FUA!.S SCII!:NCtE
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The ~U~llIItll," HOIIleH~(!. ulwal!~ ~5W'!1i~r i(I d!?g rees keIVii~. O'tOUfS Olit l r'Q@1Il'I t~mpl:lr<!ture '!'Iilhin single91r~JlMI1!~ ~eel'.i, 1111 ~hi(!hD:h~ dlillr1J~' (,jmiil2Fs lbem!f@ ~,5 l'IlI3,~si~ r,~Mi!.rkS!k iJ<lrlid~s.
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~ESEARCIHI RillClE A
PHYSICS
PHYSICS QU~liItum tHalL Ene~t 'i!il Po~~r Ch[i~e Hlu~roS:UU(jt:ldr;f!S It rSUklllOki e[ sl,
All lmitri iii Sij;( lli!l~:nd·Gen'~ ~ed Elle[~r('Jl'lfc G~ss 'j,l~it~ Unlid~ rEe l1olil,al [lomaim in ~lii,drl?rdl('Jped (lIpr:at!~ Y.i(oksoko el at
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CHEMISTRY
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CHEMISTRY 1393 Mill ~ti~u"di,onI3Il EI'IIC(lcl~dl ~.alrtide~fo~ lIfIigh-lihrouglhplUtl!!,iiO<fiIn,oteclU~~ Aina i~$fuJ, D.C. Pf,~ibGlJ, M. ToJl'~r,.,f! 5. fkJyJe Via mtho9r~p-lfIy,a micJ\ljlf1I11lT[liic.d~~",ice sy.nl!li,~£iz~,pani~les can wi~hldMlif~a ble c;olll~and delll,ctioiii Il!age 1'Its: t~ genera1;e mulliJiU I!I1:llonat f>CleenS 'f,or ,gle~e'tics 01 (herl!llli~trf.
AHA iQ'5P H !:RilG SCIENCE
II'fiI~~r~e Rela'lilDns Behveel'l Afrnounll5 I)IW Polhillion A~r 139.6 iii rid Orog ra phi c IPreciipit.aliol'1l D. R(]5e.nJeid eot oil~. 1"\I.e<18Uf@fflmt~,[IW'r 5ev.~l'al. (jj'@-(:ad~~, cen~r,a~ rna $hiow that <Ii r , in (BfI polMiOl1 has d fa matlcally red IiN:M prec lp,ftalqOI1l from Mctiltfi ng ai r maliSes,1111 lrH.iimu. hilly
9 MARCHI 2007
Science
~:IEPIQRlS.CONTINUED.
IMMUNOLOGY
CONTENTS
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EVOLU'lI'ION
Ail.!topila gy-Oepl!~dl'~liitVilral Recognlit]ClIli by 1393 lfI~as;~cyt:(l~dDl!,rldriUcCells I-i. K. lee ~t at I\nl ul1ie~pedleG r\.lncti~1I .0{ au~o[)h~fJ~' kell~l~r5e!U'.,tJlg~li@fI} ls w
!!Inij~ IRNAfrom imrf,ecijln!1J",jn,JSII!5
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IE,co~og h:allSpeda lion in S'lluithAtI.a,Il,th: II:sl.i!nd F1i1(lh~j; P. G. liIyan. Po Bloomer. C i, Mo,~'Qm;;r, L].
1420
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BlOCH EMISTR'f S~rtJdll,ua~ Il'iH.igJt'lt n~o the lr',pl"!sglyc!!)syl~tiQn Stl1!P ~ 1402 of !Baoeteria L CeU·¥lIIa ~Il EliDj;J'lflth~$i$ It. t. t,fJLio~rirl9, ,l, H. ,d,1!(o ...rw, D. Utrj~N. C j. S~'I'Yl'uldk(J Thi!'t:.¥:ral ~truOurl1 a nl,e,n~~1e'CiSl\fIHill, ror bacterial ,eell !wll Oil ~ynE Itf'sis wiU all()~ ~l1m:~UrE·ba5M (:Ie'~~glf'l nNia rlti i¥lllcrobi<l,1 of agerns.
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1111URlliII1IS, ~ I'li!"W 1'IiIt!'IiIlIIJrlI 'fofm@G in ti;)t!' l'Ue~rul@ 01 an odori:s h (on.501~dMl!dl'a5l1@r ~hel1i tli~ odor is IJsed1 '[[I iMIltt': n.ctJ lSI aaivi,ty illi ~h~h ippocam ptis during ~rulb'itqLcJ:fllilSlee,p.
Alilap,hase O,ilSIR Bef(lr~ (omp'I.ii!It;~VN.A.Rep,iicaU@1i! 1411 witf1 Intact Checik Illo'int FtesI!lQ'I1ISie'.S;
).l()rI1!5·,Rosefl et. a L Ulli@$oc"!]dif~. ~~~t~ mdllly used organism f1011O~U{~d~ sllld~~~ dM~, rlet h'a¥E'!'lI medIi'!! ni~ to prevenll c~u. dwi5iun ~rDNA rn,pHci!itia n ijsitilllrUlompiete.
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to WaUdflgl with .01 Sa !ama ndler il.'obot. 1416 Df~ven by <I Spinal (oro Mod.el A. J. ii~ft ;t Crespi (1 f/:yakQ -M'. CaDeigrJen A simple cirtlJit Ulilt ~tmtf\Ol55Wlilil'lmi~1 in a robot~t :5aliJ;~~OInd'er n ca
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• Separate5
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• Offers 'cuslomizable pH g!1a}d'i:en~stQ further enhanoe
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Ilt.l!i'aly.~to Be Uw Wirlgmaln 11'1 :Soullil Ame~kaUi l1Iirtl~ ti1ealpila m~~e atwa)"$ g~1:l: a the !lIir~ 1001his, lxud'dy be.ll'fils (rom l.agg~ng iillotl9J. [)I(!dlg~ng a W!i.!Fm~f!g Sl.!'ll@'t Bal'lIIl'Iiil'lg (K5 M~ ~1"!,iTIlIved Cli.'lllIole lot IJ~ 9rN!nl'iol)l§e Iptntcel1Jjal rroml ~he.1IIUl1ospheil~, SdrntH~r!@tll S@lC!lrl::lh lEariV .AJf!iJ erlcans J'O[f E_:I\pooit:ion wi,~ [o(!lk fcrF tro!!:lIS 01 LIO"9I.dr(!MIOOclMllllill i:nll~s in GuM CeasL
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EU ROil' [; I,tatli~,n R!€'seatr:(~ Sl1ake"tI Pi Pe'~din 9' 5. Big!)i" Ilalialf'l eilf"(h~~ awaHl1Iew legilSlation Iprolfll~sin.g I'i(>j\l t(lIOSili(!alS, ~ f uncling,and!~m~dLlr'~, US·; h. Grali1'ts.,.g:Q~
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PERSIPciH]'liVIE: nllX at FClca.l AtIi'u;::5l0rls-5Lippa~'~ Olltlih, Mectri.llnlic.aL 'Gallg~, m Siglililll !)cepri~ Y. MIOny Wh1i' dCII arious r0(.11 adhesion !lmilei~"s un dlugo diUerenL dfi'9rn~s v o,f oorrelalecl relJiOoglrade 1I1l0!oleme,nl with a(I~111fi liIn-UliFllis! RIEVIEW: The Expill1di n51'ROlle 1m II1AM· B:i!seil S:ign a ~itng Pathwaysi n ~mmll.lllH~Cel.1lli C. .L Abmm alld c:. ,A. Low-fit Ac:~v~t~(m (l1'ilimmullle(ll'~1sIlly lIfIiln)' diilrerent r;E!'{e~tl)r~ooperrld~ QfIIITAM·lN!md '5ij~!'IaUn{l,
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With Pilusiof} Hig'll·fid8Iity' 0 NA PI:! [ylUlel'~~, lihore PS00 fl!ll~ ~D'cClm~lQmi~' any >ilcS1J@ct C!r ~OLJfftAper('OOIDarw:ie. A slll]eri!lr r:hoiu! 'r®r '~"':J:nillg, ~is ~E!Ol,mhirlanl polvmerase has an ~rror ~tl[! 5D'~DWItlcwer thalil r8Q DNA PolymE!{BSEI. rna',:i 1'11]' i l tihe IIl11Sl00cW\lit13 lll(l ITTIll&lable pl}lyrnern se ·aNCi i latJ!e_ Phusi 1:111 mllA.l't)lvrneraS!!:! is; sup,plieai in a ~ta!ri ,[I~~ormal~:S, v...rll~ CI. elY ,Ell' t [}L S tar I mOflJl ita t 1011 for incr-ea5erl speci~ciLy_
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~e~om lJ~ae.r~o Is and pl'E<i:~itatT(m rrOil'! cef1iI!l'al Clilin.a" (!fUl! oUhe 1II1llist polluted .amllsoftill~ wo~kI. WOii1~'ly PQUlJtecliIir proiliided O:i'!~.yhalh~ iJIl! ~(hC)r~ f;\1I)~~' :pre(ip~l~t!Q~~s did dea n!!~r, a
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Cytokine Center
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The l(lcomoHOIl oi It! esa l('lmall1(ier [pr(lvi des an OiPP orlu !I!~tyto ((llll1le(l res.ealrch (lin \l'erteb~.al~e $win~· ming '~'!J(h as j n l~e lallillp reyho rese~ rth entetreped locomotion. l~i!5peeirte,~ (p, 1"'116; ~e the at lII,ew~ slmy by Fll!nl1li:~i)d€vellop a 'Ihem:'e-'bhriill.lTloae'llo slll,ow hlJw iii lall1lp'r,~·m,je~y;t~lllI Glim be '~;I;I~I1t1ed @~iOlail~sa~"nH!nd~r l~)(O!f!:OliQJiI. ile m'DrD@'~ W T ~~pl(!f~~s U~rliSt~ii(mrrllm trfllj\~lqll1gto tlw ~landli ng l;'I,I<lll.:'es.o'~ Ibod:li' IJmldlll;aticn~, the <llJl(lmatlic 5!IYit{lll hOIl1l OI1l'~ rnede of romlll1 otiOI1 to the OI!iIler, ~he wilrdinatioH 11l@[Vffl'i!IfII.imb.~a~d bod.y d'!l!lTiog lI'Iiallkingl, andithe oon:t~ol, Qr speed and dir@(~lon. ~(I va lida l~lhe mode l, ttl e! QlI,I'th(lr~bu i It a sala Ill~nder· tik~ r[ltml(lI pal:J~e' of ~mdl,]c; ng! (and switc:hi ng be:t~~'een)~~\li ntmt~~g. SetlPell'ln~,e cu a~li I'Ig, and wa llHl'ig g,ai(:s_
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enterd awrnOile new eX(litUn!j E!r~ Glr reSE!'awcih. [.~:llrnSOFlH! instam:e's iHJW,eller, me rE!:s:o~uli(ln ,[If 'the: :ligjlu rniereseepe is the limiti WI.g raclor i WI a I'!!SW1eri g (I IJI!' sele J1IUHc rl questions, :111') ~l'e:sec~ses.lhle high e r resQ11J~~'f) (If the' electron miclf~Hi~O'P e lis e S~H!~li:al., ICllm bining bOlliillOght :;:'I,nd IilC.jr,omJ m~CIf05(;tl)IPY is myluEi:ld or irUe rest By ~ erfQ:rliflillig el S(iI' c,aJie-dllCo'l'iI"€ ~aJti\1~' IUglu IEI'~Clr!!)irl i CI'IQscolPy~C1EMl' l M 'e\:<periI1l1~~~ om: h as th~ ,~d\fii!.maJg,~ Qf ,Iiite ce:U ilma 9illigi Cltil Ii! conroe 111lillie rose (lP'fi:.. nd a~ fV!l'>Ellr!1ish aVie h igih ra sol u11lon results lram lrn e trans mis si a III e~e~lrran mic rose (1)13'(lr~hes:ame ce-It lh€llLeicii!, 'EM IUS was slle~!:Hc:aUv d'El'l€llol)ed 10 bl€llllse,dl h1 such ,e_)(!)e:rrm,e~~s,ill conj l!Iir1c.tion wilt1 ElM PAtel2. It :pro'l1i des a hi91l~ me re~~ L!~OI1i 1:1I3't.'!.!iH!,E!i"II111 li~ ht a ncllel,@,C:lron mic rosc.ope, e Cliliolwi~ ~ exc elllellL IllreS'E!llVa1!i].(lI1'I of U'llil' I,II~ra structu rre 'C~o)'SIE! til e fl all,lri31llstate. i31111E!SSiIl!ntia'lrere qllisile! rOlf to p eh1!C1FOn mic rosc,opy. It ,8Ji lows us lJO dec idle 1m p an the el'!:111t nto m ent of inru(!!n'HUII nod,study that pallic!IIlla r E!\flent ,all c high res oll,ltio!i1l.~
have! 101r:PiO'IUI VII! rfl:allille.•. M01iiX IPilall'lek I!rustitime
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to,r M,g 1'!i!~1D10'1r l~e!1I Bi 1l!,I'9SlValrldll\jel[ile~ies" L'EI~c EM PAC12 &. RTS High Pressure Frs e2 er, a
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i !~f~ct tou ~ di:;~a:; 1:;;.1nthe ,2 hu c~~t1:.ii~ it wi]! bl;' a ~~ii\of gl(1b .. l \~!'n'ililmg •. 1:'i001~rty•. and infectious I;lnd c'~rQIrni:c iseases, Li]"i;: e:\~CI:<!Il~'y in the UIDIi~~d S~tl~i;l::; now twice ~h[d of d is
~:h.e] 9th century, il1n,d envircamernal ~l.eahllil-hcall.hit':1f food. cleaner water, bener places to live (the "built en\"iromlu:n~;")-h"':-J been dl.egrea!;e.'H; contributol'" (~a~~ el:JlvirolfJmen~al ~.C'~I~I.h. rJdmsiS ~ ~sl"c~:rmwry clla]~{mgc~'~~ ~ Envilio:nmu::nnd heuhh ilm dl.e ~9dl cenuuy wmi practiced by p1iaysi.ci<J11.1S~l~d sciemists, lliWJLim:~onamly. OJ1I~ob)' business people. engineers. ]!,l\:vyer5. architects, politicians, and I
n~!I!Il)'oth~r:;~lU!ri'iid~;h·~~h~~mdscienee. Thil;;primary Hlclis fO'f healthImprovemenewere i:rm.l]-as.lrlJcmu ;EI nt1! S3m1,Ul<U re ion. fiQ~: xam pi e.. i l \v;lsFred,er e uc:k La.w Ol.m.~s~:ed.he mau belm ~Ind t IWrbml ~~ndsc'ip",[;; ~ike Nillw York City ~ (:l;lrlm;\I f'm'k. llNho headed the SOl'Ii ~ill!y ! (;,~mm~~~~(ln dl!l.I.'ing Lht" Ch'.!1 War that ~:8i\'ed tJIOlb£a.l1dls 0.1' lives,
..t~he U niVlilr!lity QI (ilF1fllmiii!1 at lil!fkel~~ ;and I®rm ~r dii FlrdDtf 'Oi~ nihe(ent~ [5 for m,sea5!l' (ol'iltrcl. a no Plre!o'en!i~m iNill,ijona~Cenler riO! 1~¥iroJ1lmental. Heoa,lth,
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To (eL@i:lrat!! lhil! 200th ,aDHlliv@~$aryQlf ChaJrl@s IOoiln1.liins birth." p.tJir or IBriU Ih.a".:e gottel1 togelitE!'r to bui~d 11wod,]IDj'g rep,H'ca of &lMS, Bliulg'lll,the S~iIP' Illlal OOI'e< DaJr'l'!Jin
a rrOlJ i1dl the
The I1IM:5, 8eogtr Pmf~(lWa'le5, IOlllnded liJy David ~m·l·pm.iWlls, a Welsh ~.i1IrliTler,. ii~d Bifili::ihr ulvfrQrllm~ntoi!ll, ~~.rileF ,Pe'~'F McG,falh, intef!:d!l th~ $hlp a~ ii1 !!aJilillg dasseoem and ialoor,atory. IdenUciJll to Dhe'miglinal. oln the (lui,side. the $6.4 mil.lioIDI, plri'v<lt.ety funded ship will b.te hilrnished wUh 21st c.elltury na'lligation •. ~ar,ely. ,and
(o'm rliIu,n~'a l1011l~6~ l!i pment, For it~ ri r~l 'Iloy-a '1Je, 'the retP~ka, c:r,eJ;'l'~ by 30' scientists a nd s.aillQ~. will S ~11.iUOL.llnd Ithe ~'Mmld~o~lowing lh,e path o:f the originiliL Smole in ~:hE11.S31.-3:6 voyag:~ lhall ilupir!:d DarwjiiQ's. O~ rile ()~fgit~ Q/.'5{)(!(ies.•tJ.ffter thlll s:cieOli~l!! "1m be ~'ble ~o use lite' Inew B~gie ~or f'E!'S@ilf'(:I1. primarily on clilll1i1.alt~ (hlng~. !SaysJ\'cGra~hr. OM' project he is wfl~ir:l'~fi!1!g~~ 'lising lI'~,e .BMgle.'!> ~og~to co m palre the di IlIl i3i~.eor ltl e lL830s 'M'I'iU'n ~h!i!llc:lif today. Bmadcasl'~ of @'.xp~rurn~nl!i.l'!~ wen as 1,e,tU.!Ft:'SwiU be a'y'ailaltil~ t,o I,ails and d:u-sroomsarour!d lhE! w~rld Via allll<ls.·yeHJiflbuilt \I\Ifebsite. The l[;)rQ1OC~i51td1ended to p~~. some ~ai.~e INlck i!'Ulo 5.{ief!ce" .QJm;faurr~<I'J, ore~lJl1g m Illeo;ple liIJlhe rr ~1d1,says ~G raUII: "We nfHN:I to use IProps lik~ tille Boogle to g;~l tlW'ir p'lIl~e~ to. 'q ui(~eli!,"' (QilS!r'uctJiQJi! is L'O be gil!1 e-a I'ly .fII\i!\Il.t.yealdf!1 MiUolid HilVef1l, Wates, OOl!gn'l. Th@ gafri~ 'is. [~n1it@d to @.xp~rl.SwhQ ar,e orr Pr<l.MED, the dli;5~asE!'lIl{Il'Iitori n9 nelwor~ of 'the IfI~iemaU()fli!l S:o.del,y r(lr Inle(~imJs mS~,M'!';;. ~~rn!1~ pe If:Sp,~c:tirve. ",elY M~ovalive our it's arnd 11!:t:.cnt,irng," :S<lrYJi FlroMED EdHClr larwfern,r:e C. Madon c'~IHaf~ard M~kal Sd'iQol iiJ1IE!O;StOfi~ I,i\I~O' n@jm teen roll aloo!J.!l :1.00 p1.a.yeF$ (S;~~ fh,l,I1~did.lo.n.JUio'i'lla_edlil}. ~IIi1i\!'ers"ilyof ~owa .. phy:5ic~an Philip POI~I~Ei!l!1'I sa'y5lh~ Iowa E~e(ttolilk Markels., wl1id11 beg:anil11968 by predkU illig ~iheouteemes o'f we5idenHtll e~e;cliDnli, ~~.arted selling Irlu r Lllmes to r the slate ol ~OIO'll<l 2: yEl1!rs ·~go•. P~~Q~,~nys, d!es,Pil:e< ~hil!lillY IP~Qr· ~
mern bers its.--wililllrers (alll g:aln ol1~YilbolJi! 'SlOO'-I.he pwj~d. di!ICiCEid to d@a~ ill ~Ol~ilJf!i1b@C:"!J~@ "'lhe~'s a~e!il~ 'th1lll Pe!(!P~etal~ less exllielfle P05~'tijOmJ!\~if real !ll"JQ!ley 1i5, i.!'IJ,!-o![ved.
!!)ff~~n il1lilue!ilizailefore Jlll.y? HeCllllh agel1a.es Ciln"t. a~r :!i~uh 'Quli!sHQn~.But they Imary gain .11 fI ii.ich be~l,e,rsense or the bfrd fru Il'lem! ee thanks
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.~ ~ h~r@ I~l@'nu ifiiiUspf'@iJdi ,al!l~ ~\lIH!Ihe~1 rl~WQ~ld .. ;..', HB:! ilh Org<lni.Mliol'l ,vi U ril~~e it~poi!nderllic __ i I/Ii'i!ini; rlg Il@v!!1 befOl1ll '1 ]rI..!~y. 1f11 ~ pri(@ p;ilf shs r~ . o rnnge:;; ILiP tCJd!J;\I'i!iln~ ciI!!pendi 1119 th~ 'rna rkel. 1 en
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'Weibs Q~UJlIlem.nm~(.~~d IlJQl~i!fQ:i ilFld gu'~s :-1 'keep cells mnning., lQ,explo,re these networks. cvisillBloGRID rro!'ri Iil1ICileolaf blolo!Jjisl Mh:hael~yer~. of [mE' Samurel Lll! FI@'n'~,~laJ R!!i@',1!!J'(lh r nsti ~L!'t~ 1n Townt!:l,.
(alfllld;'!, ilrnd
Moh1!~(uLe,s ,M,e,e'" -- -
the Bay,Q,f Ni.1lple-s..When j'~empte-d girl 79 C E., it caus!(![i!ho!lllS.:l nds of d~alrh~. PIsl mi~lr efl.lplio fl, lod~ \~ouild be fair W>Or5e; Mort~l~a f! a
lOQmHlve~ ildiU·rnul.lion people now liVE! in tme '''Red lOne,"
il
csno. Resear(~'E!'r!i headed by A:l!QlIsto Ner'j ilIl ~he N~ticmlJl I n'~Wlli~e Q~ G<eQphysks ;;!Ind \l()kilnology il'll PiSo1l, I~:a~y. hlIv@ 1iI01.!iI wmpl~~ed
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(oIh'l\'lgUe:!i,. St~wed lhrerre,aJr~aU ~lflO\'i.rlg;enelil( a If!~ IJrot~il1 im~l!ra(~101il!;flfl blJldd ing ,Y'E1E1 !>l. al.0Il9 will;h palrtitll11 ~ fou !1rll!l!1ilau. iii~ilI'IalO~leS, afil~ [rryit '~lies--miOre than 161iClOQ ,mocfa.lim1l5 in, <IlL Curalors gll€ii1ll1 d:ata IiI',tDm tm.eli~rBra~lIreand II pd,a:te~l1~ oola~di~mIIlQlilthJy. Each el'ltry maps the >l['flisOJils",r ~ j)arliuliH proteqlll 'If g~llleilnd ~,l!mma,riz:e~ l~e e~errilmel'ltal evideiil<oe 1m e.al!:h aSi5(1di;lli'~n. Abl)llle, the ilFl~erac:;tij·QfI neaw0'rk for the I1l11zyme C.dc 14. wh~th ~on!IFQ~S the e",ftl rifOIlll ml~t0-5,i5,,, >;;>0 wll'1'W.lh,~tJiQr;gJ id.il:I flI
what. ~hey :Sill! ar~ the! mest de'la~hM .~Dr.imul.Th(ions yet or just ho\!,Ithl ng:oI'I'IlgIII unfold. Above is an ~ rnag:e rnmll ~ !il1Mi~m·scdte eru fltiol1 smnario ilfWo~ving "partial collap~" of the vo~(a,rlk c:o~uflm.ll~e' t.wo glray douds r(!'p:re-sent py,ro(lal~.
ticd'ensity 'wnents-a'll(li.anclle'5 of hot goilS.eS" a41, and mck-JOD ~e<(!olld~ ';I'~te,rthe on~l of
the (o1lap.s;~.n,r: simlllaliolls anll de$cl'bad ]Ifl the 27 hbrIJary Geop~Y5ico{ Reseo«i) ,(ell,Ef5.
SC;II~Nelc. a L 31 S v
9 MAIRCH ,2007
1345
You could
Vies" il1'van bappen to )!Io m IFecSearrch~Ihenext Eppendorf . Neur"obio 1:Q.gy oo'i,IIld be~ol,J r$! This a.ranuiill' fl'lge;1!rrch and Scknce Pirizefor
be nex
pn2'@ irre{.,ogll'[email protected] accompl~sil'm@,nls
in nei!.lfiob~oJ,ogY'researoh based on methods ofmo~ecu~r and cell bio~agy." 1I1'i1e inne r ,al1ldfil'lallists, are seleot,ed by w a committ:oo of irndependelntsc~etntist:s, dhairoo by thle cdiirtoif-iilll-ChieM crr Science. Paslwinn,ers
includ,e
or age or y'OulI1g:er:
eppendatlfi
&Scie:nce
f!'FI:IZEi: F"OR NEtlIROBIOLJ1GY
" WIUD
'
BY VUDHIJI1
:BHATl'ACHARjIH
Movers
Pit:RUt RJSIIIN G. h,'aI'!Ghi.'iit~ of U!t'i a '$ P'(lra i rid;l U11iV'i.;:!'~.id~~d 'I PUCP} ,~<ISbecome tTh,e "J·c~.3eology dilrec~.or for tb~~n~tjt~H(l N;;!dl~)il,JI d~Clj,llh.~n~-in dJe'ct the mlti~ll'1';\i h~~~d ~.I'>CI.UH::ologisl overseer ef one 01' the world's mosrimpenaar and .~rchaeol ~)g~,'l,;,pani mgllii es, The:¥"ie-lnJ ined researcher, whowill a1
C a tQ~iii:ade] Peril
h'll!p ~,li 1P1JCP post S'll.C\)~ed3 Y~lri C'1Sl roo s G he'L1::~be'~o.biI,~s a grm ....ng 'Cil(j re of L arl n1,A meriean O!IJrch~l.::fll~ to i oglisti) who ;U'~ g<linin£l intemational prominence, Ahhough P~nr\,li~!l!~ h~vc m,fld~~~~~!'!'ti,al c()mribmi~)~s ro A!~d(l:f~!l studies since the d<lYs Q:If Julio C. Tetlo ( ~S80~1.1J47~. Ihe field ,~3:S typic;lilly been dmll i.H!lwd by beuc:r-fund!r:!cl~ he u.e'r-cq~1~'PI~ed. nd bettera trained ::<cieflll]st:ll from North Ame.rica .JlI1d E1lIWpe:. N·o\v. MI)'S ar.el19.eolog[!;t.lILlLi~ Cm.tiJlo ofPllC~ '''[her~'s !I ~~.ewge~eration ef ~re~eIH1cher.s"..;1:10' can \"lOm as ,Ie.~dersor equelparmersv .... irhpeople from Fmm;:(:, Sr~i u, ~Uld t~c: U S," Th(!im;::Nas~ i~ IO(:~I] t:':'x]1Cn I'H,;~ is!~. ~y~.isl~~,di~~ to "fI'1;!J~ and b!i.::'~t!;;r w!Jrk," One ~N~lmpll;i:Ghe{'~r~ ~1mly of a 23{Ki~YC'iQr--old Oh!;e:li\,'l:UOJ;Y in aertheru coastal P'el"l~, thtlt OlPP~!lD"0d .vn Science las~ '\'eek (2: MO!!reh. p, 1206]1.
sereed on the feeu l~yo f~h~ School or PlIIblk HeilUh at.lhe !Jl flIlV~ rs.ity 'of Nlowlh
Carolina, OI~pel
\!jiorl< onissses sucha s a(:cess~a,clrinb rag vlIai<er and ~nidel1iCY of di,n](all tnals. "there nlLIS,~be' a roil!i~Xl'ri1ln:di Ilimryb~sllf<@r way [~o (olldll,l et lrialls~, parfku la rly where '011,1 r compln~ngl P.O\lJIlUCilil'l come in," GHI~f'lg$ s:ays. TijH~s(II1Qol hOlpesro foster the use of"bu~in ess p r .ii(tk.~)~o, thillt~.hi2 d~ mlve-ri €iS,[m ad ill by t.~e ~dbs:11((I11li !be dis$~minatedl, ~ IR.inrler iJdds. "'There ere too moi:lflY 9 00<1 d~s(o,veriie-s ln lJublic. heo1lLthit1ut s;n Qr:1i 'shielve5." FUS·IIOIN HEAD,. !Raymond' FOJl.ck ls tlH~ ne~v hedi[l ,of the- Office ,e]'f FI!J~ioFil Elf1le~gy S(i~r'Ice at the U.S. 'De'p'artJTN~'!ll of Eller~1V (DOE). Fom:.I<,who e'dlrilil1!iF tl1ii~Y~ilr oeocdirrH!' (:hi·~f s:ciientist. of the IU5. ~:J'Iojec:lofWlce'ror the I ntern anen all Thill r mo n Lil dea r IExpel'l menla 1 R:~li:lQr, 1,I'IIm be .re~PQI15i l.e fQ r everseei ng b DOE's p<Thrticip'!il'lion in the<$12 bHliol'1~non. Hie (!(Nne5 to the PQS;i'-\lIl'Ciar!lL S,l!1Ce Iormer head Anllf1leDilVi,~s Irellrll1d in April ;;t~06from 'IJ~epmY)l(:s, f.1!(u[ty or ~me University of W]scon 5il11, Mad]sQfL
n aliona l wa iI'IUtilt!
s.chMI~o, ~timLlild~tlt jn tHI¥,~tiijnl 0 n key lcca l ~l'Id 9 leba l. IIIea ~t~l haU~nges.,. :11'1 re-c:og !lltion of the gW:, c the sdroel has Ibeen, retllame.c:llhe D~r1ni,and J,g,an Gming~ 5[hoQ,1 ,of mobal PuMk HeilllU,. ~ . ThO' SiCtlOOl'S.dean, Ba.rbam RfmQr, says ,~ thehJ,!~dswilllbe !~ed 5_t:illrt n 'endo~m~'rll ~(), a
'i
IMCI( 'N PLAY. WhHe S~U:~fir~g tp dliauJI1QsJe T:;mHlIjit. K.allillf (ablJllII',e, left), OJ paHHlibiijli)E]f'S,~ oiIi·~ Virginia lPolyl~i1 nih: l:ri:stH.uu!! a nd ~ml~ U nliVoersilyI,ll ,Black$bu F[I. \~'Olldered wl~e~herlihe[e was a wwy to r~dU',ce the: envirol'nrn~flta,l,ifllP1.ld of herr rese<:ilr(ih, Wi~h lh~ he1p of an;hilecl mr· Leagul2' Man Lutz (~bover ~ight); she's come u~ \vlt.h [Ill an:l,~\~r~ the' ";odd 's fn rst fully "'O~lalp~b'le', !liIoblile [[QJbo.r~to:ry. Th~ two-story f.iiolily C;]I'I ho,ld IiilbQm~(ny e~lJiIP:rl1le~t 'on the b>olh:m1f[ijou,.i!ldJl!di!1~1
,111
re'h~D:eta'lrIr,
and four r,llse2!wGI1!1l!"fs onlhll! s~a:o!'lld I'loow. Ill's m,i!d~ of fi!)~rglaS$ md!: and s~d~~ga~.iJnliiinumr p~!'Ie:[$lhtil tl1ld, in~~, ~l;;'(i~, The !~b
'illmliller
in Tan,z"aniii,
SC;II~Nelc.
vo L 31 S
9 MAIRCH ,2007
1347
NUCLEAIFI WEAPONS
Liiverm'01r,e Lab D:iIPS In,t'o:the Past '10 Win 'We,apon:s Desi'gn Conl'e:s,t
\;llhat's lhe trick ~t) ~l;"Dnl1ing a camperitien to des,ign :1 nuel ear hn~d~ \vh(llin you
C.Ul'l
d.eMilo:l.do!l
O~1ireemputer] r.tu;dlL::~" d~ri~!:! ~.!HQ~~il't'l:red [ u~i~g(:<)'~'Venli.o~RI ~xplns1 \I~S al Livermares S~te 3,00 r<licu~,il.y. That \\~lS both "very ~li ".ry.~ng·'i<~the teasn Md ~1:we~l~'liy lIISeli~1 j)J,\O'\rillg tbe dleSU!?iIl '''credibi lily;' S<lYS ill'l. S
bluwi
I'ull~ IW~ db
pb:ysi,c.;h>l:S
y .H
,,\I'4;i,U
j)'U!1 ~
'1>HJU!ld
Lawrence
are ben Q r" ~: haa t hei r :dvi:d, says D' .l\glf;)stiml.a r~\ft~'!~ ofthe rub ts rup'l~nui(_]n as boeiJligl'I~.e mO:IN,;: Ol~serv<uke or llill.e lVl"Olabs, c
Bali'll!: Grill i~:.llirlP~E!!.tin' world':!: h:limh 1'I10Sl powerfiUl :~i1lpem:l<impu~~r. gree '"offQln~.lu::~ear tests ga\~e it the edge, I new I~-b.)mb. 'fhl.;: d,1UI~ Ile~ped ~rm'~n~ liverlll'1O~ wii11 ~ b~ml~(le-.sig n ~oltil,pe'M ti@l'Il. n~u,,~~.pidc31~vklcltcCW1~Y nothc c'J1.ough ielllge.which piue,cl Liverto i<.eep the pr"O'Jr.lllll 1n0''I''i11g fO:J:'w<l:nil, .h.mVevt'iL. ~::O!'!nl;ll" Lhlif;'ri!~~ar~ di~(1l1!r B·fl!fC~ Taner S<.'L1'::'; mom .agatiOClst'l:reh rival Los Ali}.mos Nmio!lO;'ti by scourlngthesrebivss uf t:~~ rnu~hhl !Ie, daesn'l, Ih,intk DOE 11~<IIs ~OIne IUp with <I ye~: Labor-nOl'), in New Me1>!.i 'W3:!l IiO de5>igUl. a ee, ~000 u:~(leu:griluumlml.cle'lr~~tslh:;lt the IJl1i'ted weapon th~lt \',voll~.dn"t require nuclear testStales lsas peI'Jom-.:d. But l!hey leaned dH]f:relllt speeific melhod, to measure 'iI\~l~~:~.eJ," des;ugn the IlIiI,I;;"!I;:tN the \."rii~d;gl~ gfbeing Q(;:r~il'j;,j,bk: ··wi~·~,. in,g..apractice the US. efl'~I,;'~i'i/dyruJr$~t'Qg,'~ 1~;Fl$ fi"\..Jftllhe J:;'u~. Livetmore phy~ici:i<t and in .~ 992, A1Ud IDOE. ol~ciclals say tlm~ soundRRVV lco:!~~il' .H2111k O'Brren ~s l]~il a. Vi'~]101l Oli,llll'C([lli.ring nue IcaD: t.c&ting" as it advenlses, uess of:l iV~:lr,mJ,ml;e desi gil was 5Upei:UOI' '1:0 a '5 dle:)i£!!] tested fcur rimes ~1:Jm the I!}SO:s bur Some ,~awm'I~r!i i1:1 'C'O~,1gJ1-e;,5 are wOJ:uiedn.£ (l:eemed to be lOo"overs.nz-ed. and he<I,VY" b."lL'ie whether R.RW is. even r.1it::ede(t g,[V\cn recent IlnQre n,'tlve I pro posal ftoml Los A Iamos. "Th~~W(i~.about :sun in g o~Tw ith the mO&1 d~iri~~,! Cnhj 'War turned mlt~n he J[lg,~rad fiffildi!'!~ I)~Jt tl~ p:l~u.on~ inside t;I~~\lI~I.1J:lm:itoi [he tum Ctll1l .1.,:H!.l [anger l~l,;1Il expected ,[Seienct!, ccaservarive appreach," says Natjcnal basis: for the :1J1,ewomb. ln eentrast, Los b 8: D<;c~mber 2ll{J:6.p,. ~526l'they 'l]K~) \,\o'urry Nuclear S!<lCUrh)i Ag",ncy (NNSA) interim Ai<"!l'!Ilt\<1offici.d GI-'1:1 M,!f<l ,[!.1;,'kn(l'I;\'l~g.~~ !:~;AA head Thamas D' Agm:;t na, v.,'~m i aunonueed hiis U~"d111'j;deS!i;gJIl 'ViiIS IIDI. in the "Hlm iqtlc S\ve:~ about its impaet 011 erm.1S 10 (;mb ~~Hde3t, prerhe out(lO]l~!):' lasrweck, ~p(n_s oi.·]pI'L 'iflU~ [~dfic] .... d\:.s.:i.gJI1SL1ull \l;~r,C in lil.~rnti(lll., "[he pmgl'$1n !w£ ~. '1mllb,"C-i[.-tll~~~ 1:IiI.e Rd ii,lble Rtl,lli<lc:el;m::IDiI <J:meOiw q RR:Wl W drne s;\OOkpile.:" ,dhmlah m1JIli)" m.Iin,C'Ie<ID' te~1S and you~s.o ~ong '1:~<1I<1Jt:ter:'~,,~,)I~ -a lq)res.elilt<l.'~ ive P:CL~VilVC]fiSky ,~DLIN).~h~lir. of a I<~y I~k)u~t;: pnlg~lln \1J;!1!; ~~ up,b;r Cf!11gr.~.;to ifi;~0mit;:c ~W{l4J.;:lli>i:~:rn:rm~ II:> d~igll" Vi,tillrn basjc desIg .. ill h1l'lTIiI11:1~eCalilbrnia ~\Indillg pmtell. He 's dlli;);!l~ene<il, to C.H~,~md~ng the <1giIlS, u.s. swclp:ile. The l"\'I,'O labswere h~ cmnpeti~'!g to ~pl,,~~ <i. ,\ ... !11~~uJtll!;Qt~Ul~'f~T!!iiy ~~mll\,V~~~ O!~ to PGrftlr.~~~. 28JlOO ;.;;im~- for 'RR\:VIiJi:~~:~J.;;*,s ~G~S amere ··(1!\)h~Nilf' some :iiit:; <l~llP mis!iii I~l'! in U.S. :i~!lJmiilru~1~. mnl; yf S lated l;lx.plosi~Jn!j fthe deviee on t:t~eIeh 's e..... o ~Jh:II~~.tioiil purJ:l{l'.5l;.\. of~t,.\'i time oo.mbs are ~llQ[lre[han 30 ye.:1I":'i; O:~dand sei~OO"~~I:3I~'!JI)s s.ulX!'reompu!el,·" These rum. :mg~ A panel of .ml.c~:ear \'\ 'C<lpollsexperl's con" el]j~:u51.sCaa 't deronaie them [(I see hIJ\,v l,\i~1~ g.e~tA.-'rl hew eeuaia 1iiI,1~JlI!II.I:~'!JCluriL~g defi::lils or vened by theAmerican t\s50ci ..riou foa: the t~1ey',rc 11~f:1dup, AhhDUg!~ lim: fllC~LS is, rdlfln.ij." des~g~. dUl1mgcs\mu~dR~foct.~lS pcrfhnmmce Ad'v~utCemcnl of Seienee '~wh~,chpublishes tiry •. R~{ b; 1l~.s('1 ntcnded Il'O cn.~ne s;;"rer. I.T!Clre ~ &i(!Jlc(;,~ hm\;caned for "imlependem fit:'fiew \li.'iJ:mL s'ubjecreci to Irlldi<l!t~{ln. sh,;;~kinS.mKl high set: L.I[e, di.Ul.d gIlieenerWeapons, IBm. cr:itoc.,s say tlhe l:empe:~·:JHI,re.s.111I1 e:!llr~,y 2006, Ihelc:'ams ~:ed!~,] " to IIno !]ito[Jf he next ~lluge:sof Ime RRW s t pmgram.liCl:>AIa1lH.'II>. hm: f~hftd}1 '~L;;il given rn;l.\,tr.-.Iln, \"~hich i.1!lu~d('(]i5l mo:['(; tI!liU~ $7.25 mil· I,;\xchal1g.<;d w~~n ;(1111"<; ufl'lda~de~crH~~da~ ··pl~.{lale-1xIDk~ized'· dollsi.el'!> Or,d~ls.s~ir~oo, d,na lion by 20 ~2. is ~.ui.sdirecled .,nd COUnle:w~ ~1~'1l <lIS a ,cQm;olnl~OiI~ ~lJ.iz:e, lLive~,~M]OIl'eS [~Ie OtB;rie:lms~ws h1;!l'~··n()UiIji~(r' II<I\'~\,\,t)~.• ~nd 1'0 pmJU!(1~\;~. ··niB: could $(;'r~il;;~O "nC~l~~!~ ~le 01'1 U,eir \:\~'ip<"~n, ~m!"'oll"b for I~oorr~vi~'\I'.\,~~I! kll nm,otiv~ne~:.lllU uliI:.'t! I:he next s,t,eps.• wh,i.ch very p,rolifemtion\w are' tl::lI.ing to pl'eve~,1C' ~l~,lIDj,i,e]dled so.~~,e:~ml"ilt,o\'lelnenls,. (Smne ,of los A~a~~o!;'s, R.1W fc,fllures might be:i~h1:c,., hu;l~tdcfu~~~scal(: ~ Semim [)uQIlI1J;:F(!hL."t~in] A). ~!mlgill\eer1ng l;!udueli ~1I1tl~l!J'gmtt:::d il1llto~lme C'.. ~dbmi<l design.) Ihu the illl£, resctlrchel's llO ~~.Q\.,~~Ii1.e d~igl!ll rent;;ts to To look 1:0 the fi.lhqre" bo~~, tl:'<Lnmsslaned : wit11lhi;,': p~SL Bolstered w~th el1~J,:uu:;e]"i!lg UWli1norol~!lm.~.., O· Brien, ;;t k!.;;y !no.m~ll OlI"]ifctlme ofs:h~lke. It'<iu.k, Ilid. l"" S~s; ~ ~m Good,,>,un. ··We·'\.'e taken l:ilcfiIF8't ~~Cp5. hut ~ Cl"et">'S :suppl,ued by S."ndi<l N.u:icnll~l~, ,[,aOOr.ilifo· came h,.s~:A.~.gIU5t,\"rben a n~loc~U'pofrhe h.'s.~ lur.uggro;;ut"..jE!l1 ,KIN:liIISCIiti S ri~s i!l1!Albuq~loe~ll~e., N~,'v1\,1,e:>< itXh 't~lfi;")'~'g;it!'l \:\'~ap(ln s~o.,,'(;,j"lim Gxoo~IC,liIl ;;t!;lOOJ.;;l:nefUwidl
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ing I::llU.H5 <nul dml ,g~~I~:-;w:itl b e in d!l:ll!l:laQ,HI as. 51(.': ien~i,:!~5here embark 011 an ambi ~ious
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8Qth van A.!rd,~ and Owen-Smith me Univ'ef~ily of Pn:'wri<l,N COIIH\A~r\'.t~ tion Ecolo,gy R.es~ ...rch Unit. argues dun members of the Elc:phal]~ Science Rou~ld
of
lhll
week
h::y
research 'progrll1lil~ announeed last ~'!S part of the gcvermnere's draf] pom~
one r~mlcdy ~s to reducethe Iliu~uhel" of water~ng ho.les. and reraeve fences to create
Table, whkh
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Several such Traasfrcrrner
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~UIVC~ been
ph,lnH pcpularicrr. Tlte :policy's II10!>1 con= II'oVlCr:s'i<llpl'ill' i~io:~]wm.I~J,!)1low U rIm,n,bed clllning-\~,ihit:h is l1Iot ~~i ig doue in!J!I1j' r A Ir~ei'ln ,eollnlwy-if at he r a:ppl'oe.c.i<c;l;l'hi,L :PIl!esell'I~liiglhepiJ~lli,q;y Addo EleJ?~mill at N II 'I ional Pu r k, E 1.1 i rtl !l~");len v f\ rfairs
Cunserva-
l(;nHfk censensus, ln Dl;ce~h~L Th(; 2. I = IJ'IIembe Ii p~mel" wliil,ucb included P:lrik Service e~ep,&1~nH experts, submitted a
t~:~
Seh a ~k\'lryk :said ~~ plan, suggested by a .,;C'iegH,ifi.c old", isorypane]. "'\'iU hopl£lfu Uy reduce the s,dentific uncerralnty" <Ihmu fbe OlrdJlil,nds' numbers and movements whi le managers de'al with "immed iate ~h:2lII:~ngcs'"s~ch ~I$ !l'Dli~li~mlizin!l: t~(;: darn\',Ul
M~fli:~n\:rManhifl~$
rIlllnltuyc,Jr research
age'IIH,~Ycause in parks
<mtl reserves. ]n
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PM!:.;, Inr
thil?
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to
about 1i ,5000 sin,ee:cuIHllg was stopped in 2 I £)94..Nnm,bl,iir~ arealso reporredto be on rhe rise i~ Bt'Ylsw,an~a~md Zimb,;1'b,ve.
ThascOmiI'<IS,'ts ;':lh.nply with the Irend ~n \1l'\e51 .1)11(1 ceru~I.<l~, frica, where ivolrY peachA
ing has deeimated some elephampopula» ] ions. 13a.~ed O'Jilthe an~.ollnt<; of ivo~y sei zed .~ndDNA :;iOUf'Ce lmcld~g. <I r~s~.~n;ll group esrimared .i.1llI1 the Proceedings l~f'liw
j'\hl~i.r)n(lJ Acutfem.v of St'f(:'rJ(·i!.~ {PNAS)~QSl week H'IOl~ .2J.orUl .f\Jric'l'lIil e~.eplmI'lH'; were kiUc\(11 ~h~iu for IfUS! ~e~r alene, "Overalf , the tdepti'U'fi,1 pcpulatjons
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~11llii~tb!ii.1JC'rl,;:'ISir.l~ iHlll$ihrrp,; nv~' or ~ix ellf:jJiham r.Hlgg;: ~1'UI.ii~··in ;;;[It:I~her!ll ~~ric';i!. A :l:myslilt: Ie.,,!:!aurhor of the PNAS study, S..muel K. Will5~i;r.direcrcr of '~he('.~mer
far Conservation Biu~.ogy ill the University (lrw~~sh:ingl:orl.Seattle, Bm declin~,]Jb'g Imp~1.~:It 01.1 are <~preble rim, i,~l '~'Im otlm ell 30=pl u s 9s ·e
in seuthern Aflii,r.;:~~ I J ~p~g.(lp.l~u th~lt C.?iUS for Ih~ cloev~I{'ipri~~~t Ihrough I:~,le efforts of rhe iadepeudem of predictive mcdels o:n. c.~eph,mt managegO<1!.Is., The mi~i~r~y hOiS P~ac~ P'irks. Found,lti()!l. [u ~Idd.i~ifl~l,~s m~mt ~lnlgfig (l!f~J;:v p,~rl of ... Ioilg-ten~.l meg11p<lrb research i! Uecated ,Jboul $,700.000 lO start the work praJec;:t. van A~)fde's gruap has outflued n ndw ill seck <Id{llii.Olm fundi ng later, l al
n~n~\:~la tes,"
SII'.UCI:po,iicm.l:1ghas,I~'~~llj,led i'vory poaching m SOUl~, Afitic~'~ w.un~ p'if~~, m~my b'lu ~ ecollog:US1Sblame IHlOr managemeat-> :i iu elud i:~.g H~e dr iHi ng 0 f l1loU'et~] an
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ch<lngins
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last resert ifothcl" opl..iJom;.-bel ieves l hat the new 5(1 uth Afri.c an i~1 cl m;,hng relocat ~OI'I. ra nge )111;1 [1 i,Plll<t[iJtl~. and contr,;u:epli.m.-dun·1 '!,\fo:rn. But l'Iejitt~er reseerehprcgram shoub.! :focus mlo~tly on "adaptive ruanagemem," nr'lUrli'tori f1!~ ehe o wen-Su'ili Ih nor V<ln Aard~ .. b(l~hof who~~ .J;t.'ill,ernl]y !>upmmr~ l~,lell:lJlilli:s!er:s plan, Ihinl,ks ecolegieal ~lliTIlpm:tOrspecir.c actions ..such fh,~l t:: Un~!,gis now \\'l)(!T"~r:Ii!1;!di!:'l! Klfllls,er 1P1lrn .. m~r,~mnvh'!Jl: ~em::es Of wareri ng holes. Alld rraekiag needs roceuuaue fOIr' "seveeal and both question its l,ong-tel:iI;TIl effectiveat yen rs,' S~yi; Nm:1~lml 0 \''lMn~Sn~:il~~ii1, a Imrgc~ ness, "Thc.rc is :lmO poi~t i~ ~~,II,i:D1,gIKI1,Igc 1","' 1lffi,<l~IiIHnOl,1 ecologist who clIin:C1'5Ihe Centre s~:ys van Aarde, "We know now thatjhe for African E.C'oh~:gy the Ul'llivcrnhy (if at t. 7'~~ar c·"d1i~.gpro;,>rom ~l~re.whid~ ~movcd rJ,l.e1iJ;;,r.u~,w.al:erl:l[I;an,u il~. Jolulnnesbur.g!. ·'.~I~ost m:<Jfl.y 17.000 r::le'p~'<IIlIs.did :I~(.n reduce I~J.e~r ~f!1:)Jpa"".i'l [j,le ICHm:g.nJ.U'I:· ~ROBERT .I!)OENU) cl.eplwu~~,~lVIo\"~mG!nswdtGS so h"IW
!I
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91 d.ep'hIMH:i iu seven so mil ern A fr iCow coumries with OPS tracking collars. He
The euvircnmearrninister
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SCII~Nelc.va L315
9 MA!RCH2007
The ·~!tE'r,cof'lfSils.ts two,vislt.!~e, phialS/as, eacih wiMla o~ dlifter,'em pam sim,Thistwo-plThase,mme'( prot~ction enSllIf1eS u~ti[mate abso!!p~iio:n of aerosols Gand br:omoloo~.dea 6., (nJ~matQhing aU 'o(mvlI)f11ion'<lll·fiitlllfS. Rely on ~t, For more inFom'llatiTan glO ·~o 'iNWW.e,perndlill'f.,comldua,ntiler
eppe',ndorlf
In' f,ouch wif\b life
Yen:llI'lOCIiIdi!i.1rillutI:U': www.e,ppel1ldo"' •. o~9[dwJl:I!I' e AftjUCMten S~PP~r1:~ T41i'18lMI,OOG1 691 Ep,pendc:u1' AG " Ge~ • +49 tliOI·500 m -0 ,~IE~OOriIINOrIIh Amemi~ IIiICL 8OO~M~H30510
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,METEO,ROILOGY
E ~ NHio'~ h~jh1CI~
Three htl!rric<'l!t~s hmJiippe:d lll{;'n.~~~, HQrid~ I~!efp~;,roiled in ~ht::e~~ei"i1[ffirit:~i]Adiilntk. ;is dm,il'lg the 2.004 season, Four hurric.mes,. .~l ]1,13.)1 ~:k"lJVe OOlle l<IIs~ year "!ilhen it pUl i,m!, :SUI'" OJ ind,lL(~n,g K;nr~llI,~. f"<'I\"<'I.~>d ~l~ G~i~fC'~l~.stprise uppr:,.m~~~ b~i!lning in All.lg~l~!, in hmJ Bl~t inJ{Kl5, Now me;t~orologic3~, !;fgJI1~'!j!i,'CreU:~,<Lnthe' west, I~!.'.'arthe Cm.'.i'b,bean and the United immq,i'j :~n iti(l!l;!lelli~l,gyet .mo~~er hyperaetlve Still'e::;,. dust hOl:.'ii b~enl,~e dumiil ..~m external tlm;ilricl1'lID,e ,....e;.SOUll,. the: eigh~h,in I 0 years. 13m ~Ilfh.e:l:lce. they ~bll.nd. "'We ',Iie not dftrl;y~ng the ~()r'l;i'Cili ~I:S\J\l~re Iflr 01T th~ mark" 'The 3006 El NiilO had:a1l1l illi1!p<!lcr:' sfl}I's Lau, bliI1~"n,na)'t~e season was normal, and no liI:lllr,~~ C;i1n~~came ;ll!!l.y\vh(~;~ ~,~.~r[he IJmli~d St.nes oethe Dmi[Jbeall. NO'w [WO el imatol o!:l,lil:S are ts sllgge;o.;tin~ 'h,1l dust Mown m.:w5S t heAt'I!I:~ti{:' fro:l1tl the Saharawas ~'iVQI"11 tiu,e busloofureC'ds~s. The in dust t>~I;:~~iM,~O b~i!ve s.uppf'~s;i;ed
Sl~Iffil,ICli yil),
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Mer:ck R:~$ean:hLal.roralQri~. The Pl1(1~,ey~rom the Slanl~!2Y ~m:nlUi~e.'!mlnded by ~Il!;!'fdilTllily of ThoodtlrE! and Vad" Sta!nley. wm help 18~C!ad res.t!'dir{h~rs apply ''Ihe mOiSt advaII1iced ,g'~i1omie lQo'l~~ to Uilf biolo~y or me li1iall illn.i!'SS, sa:v::; lrni:lFVan:l PrOVQ5t Steven nlyman.
-AN!'J!RIE.W lA'W!!.I!!Iit,
em North Ad:mJ!;lc ~ml C~rib!b~'i1n by bluckln,g some ~1~ergizing 51.1n!]ghL they say. '''I If 1m lhey'IL'e on luI;; to something," s.aysmum:i.cune
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~"V~;H.Iirions :~QQkedpifQpi IIou ~ :Wtlr !:lJliO!:herb~uslcl)' ~:urrica:lrlC season 1.:II1p<nticulilr. therewas no ~i.~J~f o
E~ N:ii'io.\?!.~lOse P,1tciJicw;mnl.og cam! reach om h~ rhe All:;;Lfllicm~d;)lter atmasphericc lreul ,1fumll to. :mppn::ss hurricanes there. BlIt [hI'I!Ie~ m."rhd by lFt'!!ru~a~H~n>l.~!!1 nU!ill~maUy heavy surge d[l:SI.IJeg:nn blowing ot:lf Nm:iIl. A~ll:i{.'~1 !:l~ldil1JtO the wesrernArlan lk at tbe I. J ~LtU;: beglinnunsofthe ofJici•.d l,n.nic..-Inese!}S'Onl. Tv,'O
or
\~~k~ID.!r;;r. ~'1u·ra-c~\~'[l;.T.."; Df!:~~ \~~st~rb1 ~1~' AH3Illic began to 000]] oomp"1Jied ,\~~~~,temper~i~n,;':;i!l~:~~A;:~',~!tlu." ~~1,~(1~~, c~m".l(llog:i.~lS '\lVil] iam L~UIof NA SA.~ i Godd111'd SpiII.(.'eFlighi[ Cellle'r i.1m Greenbeb.
we have neglected an cql!<'I.lIy ilHlXJ:r.ttInl: 1~~C!:c!f .. irnnt ;;\mere ir1J'lpooii1:am f,telO:r_'" Many ~llllrric<lilll.eresesrehers are inrrigued lJl!J\ (:.mil"h.1l!!'>."TI1~ 'l~a~tlu:.ll\ how,\;,: an imrigl!i!'lg hYj}m~~.esis," ~ys Chr.~s.toflllC:r Landsea of" the' N~ti.m~ 1 H tltrri'(.".a~c Cen~~r :~n111 i.fn~1i, FI.Q~id*l, IJU[ "~:here8~cvr much evidence lhm. there is a dii\.~t. cause ~ll'1d t;:lleu ,gtling (in here," A~,dir dustwere illIwolved, it. weuld have been urore
A iMASt\. prQboe IUladillg lQ f!lQ.!loand a bnvpeall Spa ee A.g;en(.y ([$A) 5pac.e(~rt on its way lit! a I!:Ofllltt.wilUram liip In cllmin~]J'M;:tks insn ~fIlJ<5U ai efr(lrt to ob5!er'll'e ] ~piter. E5.#\':5 R!otieitta. ~~ulflched in 2004. and Wrr€n~ly 'iilll the n ~ghtborl'modl o:r Mars, liIIiI.~. exan~ ine U~erin.g ~f @I~m~c.ilny (h(.irg@,d~ntcles around l~~ gas 'gna I'll. pla,nellh 211. lII1Iay'stern hOin vokan k ~rl!pti;(ll1ils.'Ilnl ruts· oen ~IJ.lI.tk!'arn!tl1ii~eo, NASA's m New Hori:Zoll1lslIfIi~iQn {b~lllw)'~ped pasl. ~liJpiter ~ai$l\veelk aH'er le.w1ng Earth il'l]lal'luarl' 2,006. ,A~tlile pfob~ !l!SI!1!!;tilE1 pl,anillfs 'gr.a,vity to 5,Ung~hOl it~ \!,lay tQ Ilh,H;O,l~'e (lJ1booIFdl ins.tw" ml'rlt~ <I r;eIIJllOI1HmUfi 9 'th~ ]lJIpil~.r ~ys lem. Ihe rough ly~ nlulLa neOIJ5, obS€Nalio!il5 rrolfll~il)eh\IO' pr(!~e'S 'I::Oli.l p rQYi(j~ a lun iqlfe 5e'l ~d of data on Ihl' planet, ~We rnu~d n'l. pasl> Iii P' IIhi5 'opporlu fiity 10 sll!oy JI.]~il!I"s mettrofol.ogv,
c~~!l)plk"Nd ~I,~an !'>imrl~C{)!nli.~. ~y~ J~$[m a D'UI~i()nohh~ N;:~li{J\~al ce<!nlc ,!:~d AtmO',~O gr..phic lund MCieOf'o.logncO'II t..or,.llOry irl b
Mia:~!rTi. The du.~( OOllleS
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Mm:yl.md "!Ill! Kyt~·1\"yon,gKim of the U~liv,ern~ty.of Ma[fy.~ml(1. Ba.itim!Ol"c {'mL~].t.y. in B'J~lillnore 'll'd,lu!,;in du: 27 feb'JilQ<lfy iS~lIe of ~ £o,~'!:hlEll~;l.1e arr:ii'itl:l.1 oft:he lh~lck dllSI. o:Ind tille
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~ ~he~U:if[!r.,'e.l~J;'y Thill ogo~i~;\!I'em o~ to s.ly. M nigg.er ~ shin to\!,,'<IIrd ,~ess;n~VOf'db.~e .colldi~ t~o:Il~i':or i"ol'lmltiml ~J,1!1ld i:~]1\t,:~1!l!if'iclldoll of t1~.e so~orm~. ~n the\~'e.s~:ern Admni'C. t~,U;1y3rgm: ..As
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rl FIgs". all.~~(lrQl€. ~tenntes.. ,and fnl.a ~Ilelo:spheref " 5<lYS,if'IJelt~ Horizom principc.tinvesligaw:r s. A~ In Stem or Soulnwest. ~~eaKh In~tirute' j n Bol.! ld~J" (o~or€!do. The .ioint effort 11 IJg~f~ Wl?1l
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im:er f'latiof!(!I~.(Iop~,r<llljonilli Si!l"me ( Kiene£!: Stl!!m !tJk!!s. ooer as; NASA"s Ki~,ul!!'
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~NllmiRIE'W ILAWL!ER
W'll'!/ll\tsdencem ~g.org
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.... ru.,iiII_fldll. With ,ius~a [~larnlg~in ~ignall.~lreng'~J'~ h ~ tlM$ n)bo\:. m(l~@ldalfter iii sal(lIt!~lltdiM.can $MliItm
to 'sh~n~.ua'flll (Ill to !.a!ld., and: w~l~ oilWilPj. sw i~11, Once t he nell ral Ilieh\'o:~'k!' for 1110\'s i]lg,l~gs wercln place, IHdc iu,hHt.iono;Li neur~~l kc~~lry\o\la.s c !"~~Iuin,"'dljsp~~n~i"idhi~ colleagues report oump<tge 1'1,]6. "This is. !,;I~<ldy 1111i'i..;enel1~ fmj~lIl1! of ~ b:iolo£;y and robotics to test neurologieal .md e~'Q~lq.li,QIU~ry hypnrheses," S"Y~ Fg:a~,lk
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,I
<i
Pe IlGll,SYvan ia. "'TITh is pflp(':r wDil11 I high-profile example ofhow IiOb01S C:f!!1l be used as surregates nl!i U'l!i~~g;:l!'ld fbst>:illsysl.f:rns." Sa h.ll~la :IM.l!tus arca I (H :~ ke th,\! rh's l i hH1d-b.~~edt~trO!Lpods, These ~ulll'll~11].~ [IS: 1b :!Ow iIn ~lll ~l!rir1le1'It:I1!I~ Ih':I' as prillli;~,hive fl sh •. tJilc such .IS lampreys, andthey waddle, with I.egs. pl.'ly~d out, W{C' ~]H~flto.r~!lnd, l~leiu s ~~n(,~ie~u relatives, The ;j~~lt'l.m<i!ld~f "repreVeT);! tyin
senrstherefore
the evolurjonary rransirion frern swimnml~n~ to\Va:~king:' 5<:1)'3 ljspeert, ~~'h.o COIll~ bined lon,::es wil~l ~NSERM r!eurobiologi:'ll J ealli-M.llri e C :!belguen () nhe U 0) iversi~y of BordC:!lux.Fr~!1JCC. and I.WO ~r~dual~' stud~mns tn de~-erill ine how the !br,) iIli c:m.nrels U~~s:!plllphib:i~" 1';. mO\~$I.h'l'HS OO!il~p~!'~dtJl ~11,e .lInmprey 's, They fceused 01'1 two neh\'Orks of aerve cells. called central pattern generators .. located al,oll£" [:1m.:: spine. 'W.fuu:"l1. :Iiii.ct\~k is ill.
EVOLUTIOINI
is !I. triathlete. She ,..... ks, She c rawl s, She sw im s, One of \fcry 11 rew :mbots C<llliilb]e ofmulriple modes, of
!UCO~H)liOI1il.,
physicis;
thls salamanderfikc
m@c~jl'lt'
it In<ly have been ['I!::I.·.niv'(l'1~1('<1sy for ~~~y .,n~n(HII!' rn '~<I:kr,;- their r:~n'lsteps on land. From a aeurelogical
l:l~tSdenlilonS'tn:<lled I:h;lt
of Teciln,(llln,gyini iIL.ulsfullne:. Thanks IJ) SMmJ wm Ira , he and his.celjeagues have ~~~ullivnh'!l ml;;relyd~<l~g.i!lg the !'tre~:!gthi;'lf ~ the brain signal dl:ivillg im;:ornlolioll can
Hel iv.;ned. its iI'Id:iJV id aalaervec eHs alterIJ)f~leberwees fiu'i:~£ <lind bt:nn~, quiet causin g dlydlln~c~m~usc lee mn r.;~c[k~n~, Il,(J[ih the: la:lmlpn:~y ami. the salamander have one net~lii(lrl Hi d rive dl'~Ih!J dy nfilmu,; !I itt ure. For l
swimming,
this. network
sends
WJVe!'S
of
perspcceivc,
walks
0.1'
U,S.
SC'~ENCE IPOUCY
NSF is [lot ;,1.8 we]oo:m:III,ing as it ShOllld be to such res-e<ln:'Iin.:' To erase tlm;tpl;\'~ep~conl.iili'l.e report s(~!i1,~SitS f1:~W. d~Si~ittC[,mm.d ~~'p8111Ue~(mliii.dia~ ·'3.
~Flm;Ihil' US. Na(ultJi.Iiii.al deneeFmM:uliu,[CJImJ S U~SF) :s,ot~e[JOOoo cOJl2!erV<.l.tll \ff~,!A uew d mJi report from ,illS [)ve:rsiglll !Jody ealls on lbe'
~edel"O;lJl ro~~reh,
ag!;t'l-C~~I(l, emooo b
id,e:lmtDfy "pnrenrta Ily lr!l.lm-sl·onn.~,~tiV'e"" rew;t'f1Ch. il><Iy:il Bement. ~Iddi~gthatanether pr,ognltn \'.... ·ol!JlldWlt al.~.a.lready everburde·~{;~d ff. S!.'! NSF's peer-review system :ms wi.de~IY seenasthe go~d:nandOlrd fQj;" ~~'Ie;C'~in~ some scientists don't even oo~her hJ3pply fOil ~f{!l1b;. .For Dd~a5i,ll~ ~lJ1t in;:ro::;;S tbe sel.• n e:III,I:ifi c ~p~'l11beca use of \~,'tHI t l:rrn.e d, ~;Ift j,
:mli._gb-qual i'ry rc,sieal:'e~pro:p,o:l>~ll;' Blil bCllm:d memoors ~.Iy Ih,ey are wnrried till.d.
rica-wide
l1ol-ready-f~!r-priirnil'u::;-lim.El
l:~~!U"'might
<I;
sep ..fi'tle
"'ll:aul,brlll;;ltion~1 research ,i!)h:i~liv~" is irsc-I If <'I bit O'V!:tr the: ~Ilp. fhe $6b i]~iga,llhulTI,~ (I'~li O!1l is a~fe~dycl!o:iug,,ewr;y '~1Jling h c~m to
til ink~'~!:l: f&~~. iJ~~dts..y...bo~rd ~n.~lllb.t;ir ~ i~. f 1t<lll1dalt <lpl<llntphys,io~Q,W5!t .'1:1. ~ the Ug.liv!.1::iSily of M.is~.(Yulri. Col ulnlbi~,. ''1\'111(1 ~:r~ ~l g<:tes. nte LIiI'1Iillllillc'lfli,~"'r-e~f~e\\'rn,C;a1 i gljri~ulcr" ;;:
.Dmlsj<ls ~l']~JUSt yel
spit nut"
. •.
I I
I-
o~le.\"'hich
(,IOI1l11fOls
~heI~mh.....
.111 100]1" Cabelguenand ~~~s elleagues c d1i~coveTed the region of l~.e ~:<II<l[J]~mde'r's 1¥IJ]db~3i:~~. fin~g ll~at s ignall> ~fl these hV:O
of mm •. es. d Us.U!l!.ga .rc.m.c:nc contn:)]. ljspeert and JlI.iS gmd uate s tu den~ J'de ssand 1'0 C respi Sii.:I'll si.gmds of V~Uiy.~Uilgstrength 10 SoJ~~.1~J,nll'1hf .
A~ [Lim. abelguen'sexperimems, C less i!1iih;~~ISCslgnalsc ..used tile' rehot N) watk. WHh ~lropS.~i' sigimi'iis., th~ ~~'g~!.>p~d~!j)'. [hat w.utliillse stl1Ol'Igest s.ign~~ls,.the legs :slopped t moving and :)~fl/~lfUmdr(l began dil~ering."Thi:s c lese corte-
.on'
c€'!UIi~li
p:Mtc:rUil gl.;;l11!l.;;:nH·of~.\;llhli;:!~lh~
researcbers
gell~I}1 srirrurlated
lhl:is part of
:->p·ondence
fClssI~ systems."
-Frank Fish, West Chester UI1IWersity
~he brain dtlctr~c'IUy, dley caased lilt: Iil~lbs
move 3~ if waU':'UI'llg.A.s they grati.ual]y increased t~.H:;_~llp]ied CllU'el1lt,lI1e:u:L':a1 activ10
researchers [I1JayfuJ:~,,,e <t.ccur:;I[ei.y rec rea l~d some of lime aC~lHIIucural 'co 11rrol me chan is~lms iSala-
dti ring oil I)l!IbUc: (,OITIf!'!ent p!;:riQd lla st yea r, SQ!'I~e blol1lledkat sde-nti)ts ~ay Hllltnm.l<ltiom are neeessa ny to keep' pat.ent a.ppl.k:atiol1s u,p to dale,
manders
usc,"
~21)~
fI[mosl E!VE!'Y
OJle!
i.t}' in
~~~C
Hmi.H.ily
!'1i t,
biomeehsnist <U.V;Is:sm College .un :r~)u.!:Jhkee'pli:uc. NewYoik. The "est! Its, L(lmL~ .• md o~her!" say .. ~llg:;ei:a tl1'l.l ~ar]j' !.Inim!!~!; idmi'~mi€ed toumw~nl comd pleldy lj.eWll1e~u:'11 rmlh'''''.IYS ~oe.... p.I!1.1d their ~oomnQtor repeztcire, Sorne r\~:;;~"f\~h~fs tll!:!i! k ~hh, s.i~~p I~
re$iO~ndimlglv J'leg;<lllive." Wt011e'lawyer 1Y!<lIdt P~rdlHi!' of S.1~rm LlP in D.a'l~a5, Texas, 00 th~
popular P.tI~.enHr-O blv9 lla6l sum mer. 1M!) Director J~Of![Iludas disag ~~s. "·W@'r:e b e<11ll'gvery t hOlJlghUtil. Cll00Ii1t Ilhi;s .... he wid Serenee. "'P~ople dem'l. want to di a~g,(\l. Hll ~Vs ~ h!ll eJ:pect·oo thll Dai;;Mtlllh when he proIJQo:se~tile Umil'S. He Ih~~fil't decided whefiLo ;~lIe·aI nf!'ill d!tc;i~~onon the rules, AnO'th~; IDuda) il'l~tflltilre.
·aI
moving <'IIil.d
Gi:I~inh:mn,.Uon
concluded lh~IGbe I~n~bs'eerural p1UN;:rlJl .!.;ene~~t,tl:f interfered with the otla,er neural !1e'lwurk 's ahi lilY to ii:e'l up thL;; S-tv.(lWS_ This i nterferenee produced the slower body
bending necessary faX'walking. Only
,\~hle'U~1
o iffe.m'\o1111 e W
,S
flO
COlhlPrleite
was ::;11 m
network of
When it comes to improvillg U.S. 11'1 110 'IIaH(nlI, rolfli]~ ~~i5lflt~:rs a ~.e·thun kiPilglIr~g ~\l11.'l\lVN:S. orhers s afY~hat SIII1 [!JU is belter.lrh[~ week, Seni3ile Maiori[y iLI~'a:dE!rHarliY Reid (ll)-iN'W led ill biparHs.a 11 (CI.a,lition of [email protected]!tors '1I11doJsing the I\m~rlea (OMPUES, A(lt, a sp:rg,'I'I.I1i g bill[ d rawJl~rn:I'Il n the- rl1mmlilll!ndalion~ o~ a 2005 Natiollitll Ac:ade':f!1Iies reporlr:!l1l ~'ow til) suengthen the U.S. s.c:ientilk enterprise (S'df!lJtZ:f!', 21 Odober
-EILIZAIB'EiI'IIiI PEINNI.511
2005. p. 423:).1'lle
<II COlllllnl\l~
in ~lep
\v~lththe
rising
Sh~11T6 f lOp·f~lted o
llilOW
nlJ!~ i!l throe, H~ '!!:~ys; ]~~ .a!:!roo~ wid1!d~~ bO<lird 011 l~,l(: .import';l.~ce of Illhlld~IBl~as mUChp(H~M£Ol!lb n]ii$fDrmOl!!~iv~ reoo!lre~ <IIspo~~ib~.e but nol wi ttl ilt So cOJ:l:I::~u:si 011 that NSF :Jlho1Jl~(\illaunch II. SICPl!lriM,(! ru:nlw~.ill.'ltive .• .. Eveu"Y di Jie,cto[;It1e has 51J ec.i.a~lpJ'ogrnllli~ s
ftlr Frolil:t~!.'l!:'· r,~s~~n:::~:' e notes" "'f\:IlG ~v~]"y b ]JKlgrnnl olfficcrr is ,~ookiliS tk!:r e1l\,cilillS [:ImpCliS!l I~ fu l:'I:d," ~p
~la:;:riisC:ll, by 4',% sinll.1e1 2'O[lQ, WDI'tD,l ~ .Ii: fll\~l(h i~l M<:!!I MQrui~g~l:g., ~lq.i:,,\'proiui,~ i grn~m. Be:lmm~m S3yt>. VI'Ol],]d ,1C!'2IIwtev:efl I.~l!is lime to 5tlek out~he IYlIl~ ofre::''e<lllxIiIIH'OpOS.I~Sd]lU~he :report ill c .. l~iilg fur. S.~v~nllllHl~rof!l(,:!1li:n~rs $~yl~~y \'\.~tdd llike ~~~,e :fe'\~iew pr-o'cess to rdace more Illined
s.1,t!l\!,jing thiil~. Lheprojecl if> fe'.1Is:uh.le. ··Hm.i:~ iswhlilll peor,l.e;.!lre cij·.fl'ifott~ ahl~ with.W~ ~~'!.'l~d:;nl1l~l~Jing , difT~r,~ ...t~" ::>i:1YS bo'Oud cha.ir Steve~1Bee'ri 11I(g,.TIlllI e l'eil~o:1rt
ne~:il as mIlL ltd
~an
aoo
ewnph~sis Q!lrh~ i!1v~~Uga(or's id~1)$ ;u~d u~<ldk.record ancil ]e:s.'S nu 1~:IIel:iI'l1lilit,IDl~.D"Y dal<l
O~~
,:;oUlls,u'lli!H.
he says.i!'l.
!:!
he;i1'!.iY
.'1
m'HM:I.gclwe~t
~@gislaticlI1 )tamis
C!
by (il)n~res~.
W'\I'!/ll\tsdencem ~g.org
'li'I~ilfl!klli1!11! lilli"
B~~ :sdelte~' h:ol~l>~h~lrurn\t~d I)~ in C~i~l{:1., ~~.
wl~~]"e[(IP ptiIHkit!~s
<Ibolll.t
OHti!'l\li.lai(
'.'["1;';'0
!,!o~t~]l~~c
bombs mJ d UI1,¢ Mm'''; th~ dt!vdo~iI'!i:le:lil!~ ~lfthe ~Wlmlic ;)!rld ,ilrydrn.goen bo:r.l:lbs<lllJld~ll~ecc!ulI,11t1·y's,rulrslS3t~:I,I wte iin the '~9608 l\':3.deUj Uketo l'H~fJ'POU .."
:rank
O:I1II~;e outskirts
of S~:l3JlS!hili.p"155e]mger,~
me,gapmjec~:s
can glimpSJ2 whal!rmks lilkBmJ gi:u'll S;U~\'eli"~~UE tilas onthe huriznu Tlsis :.Hj,.OOO"'S<lJU~1reolJIIleteil:" ~pim'l MrM.emf\;':is n~(l ilal'lgi1ai Sym:hnltm!'! S Radi ..!t'io~l, ,I?;n:i~,i'ty,(SSRF), :schoo~I,I,edtoecme c:di.~ ilJlOfl9.
omd~1 ~!'().f1he.m:d [0 fi ve of 1:2 P:lFoj ec ts: l he seve ,II[ eth ers .ne ~p~d!",d to rc,'e0h~tlPl'l~l'!{~[lah.,1r y~!r; !~i~
I~. N D:R,C ha~~wl1li en Wmpo'r~lenl,s ilI:rgue'~hal C~]in'lneeds to inv·esl ~.nl1lilcgl'ltp[lojeCl:!i: 10 'l100S.'t its rapid otS(!eJlin as a re~;ll'd1 pO~"<eii. With I.~r~~f<lt:lil~ ities, ·,'\W ean pl.·Qud~y sland] up il~ fhe i~,~crnarienrrl ~cief'llil1c COlhllill~llt~ly andno
Irt{l]y
Can.
3IJii.
expert
IOI'igelf
en fomign.
oqUi.Pil!l.lW~.t
10
do
!Ii!'!
!1iiIiI,lJ1lY ,~1(~r-
fad~"i~}',\,'!I'ht'CIlw~ll g\'nl>irn~ p(J\"'4;·fr~l.",-~!y.s : WI' stmlyI.lJig t~~esm.u::lm'Cs: molec::u~c:.'landl Hdv<~I!I'!CedllIJUIW:ri!lls. " v 1]1sOO!'! be j(ll~"d by an~oth€:r Ile3vyweigj,n ch~mp. Lu~tmDl~th., the Chinese hClldt~lyofS(:;iem:e:s ('CAS) 8~gnoo Jilil ~Igreeil~en[wi.~1l1. GlIa.ngdollig Peavince in sou~llil~rt~ hi11i~ [ill bu ild lh(l $25[l m ill~[]!'l C
or
olTi~;:i~1 'th~ m
.is to ~mii Id ..
h dem()i~$nt'eS hO\v a I~rge ~~cililY C'.:I.1iI mke rom in (,hina:su~,:poll"'t ,II the :Iin.igh~stlevel 'Olr gll~fel.'[Jme.mt~D'~Ilg.:hirose! f decreed Uttll it l'Joe bU]~I: he'll" from :iru~:ern<l~iol1:d <'ldv~sen; and
co:Hab orarorscan dmasti rl"lJ'pon~n[ Iy" I he work of indicem:o'UlS:phY!'>i,cisl:S I:r.l.il'led in ~uclea!i \Wf!po!mS:<lmJ ~rtiel.c ri~y~icj; \\~uJ
China Sp<l~~<Itlliolli Neutron Sour-oe 't I(,SNSll, Tij'~mksi!l Po:lJrl to a pl~ctg.~ to rai1O~It&D ~1)en,dil1l11 from 1.3% '0 r gros;!:; domestic
O!Ii'reill~:u3iasli,c e
about
preduct
(;SCient:'~! .•
IlIiIlalll<1:[l,oo, to~m,',1iiI Delli!;'!; dream of <'1111 e.. penx ~iv!;:'pro~o~,<Ii.cc!:'lc'.mto:rilllo11. more modest
NDRo(' QIl1~ici;d~ decliaed to pmvide~·Ili~:"q}r" ~u.atiOI1 O:Ui!<II1iY of the prnjC'ClS., C'~a~m.il1gthat )"ea~1'>aud 1111'i;~W clllm;oepl:5 <Ire fil!jl bel;xuning c-ertain d~ta~~~H'Lre secrets. Ami ~nos t sci~t~]te reOlJluly. "fhe eeneral govermnen,lt's N<nio:lI1l.~1 enlis;~:s declsied 1.0discuss.~heuniti:ulvii::S ani D ev stopme Ilt .ul'ild Reform Cfn1~lmJissl ~1!'1 the n.::cnrcll. ~xpl"~.~sin~ COnCI;\:Vlllh,flll'~eilr {NDRC).. the 3.ge:n,cy responsible ['or all l'Cm<l:ms c:ou]d jeopardize nm:(]ing for proj~ sets no[ 'y1;!( fi~!l~.it~d. But rr(lllj~pubHshed !'naj or &ll];~~inv!;;5~nn.l;;mt~.is b.nikmill i',g 't[li;; construerion ofa. dozen major faeiliues-> reperts aad i.l:Ii~cr\l'i.e'VJS ,,;..itll~ two dozen s.cie.IDl.~ to the' tune of $750 mUUon-durullm.,g l~~C ~ists aud ('I rnil:,d9Js, 's,{,'i.t;IJ{,It; has p'icc~d ~og:ether a picture ofChiu!l's OJm:b~,I:ioll~IEhg I h1ll, S-ye.u plan, whidtruns dm:msh 20 ill0. {SS,R.F W!l~ ~au'1J(:,h~din IThiAl' !~l:\evit'iij:,~ 5~)'('-!u' Sd~miJ;,'e !~nd!l t~hlc. p",IJ55). ~ ~~
NEWSFOCIJS
h~lp.;;d Chhl,f~develap h,'ld!lic'~!capaclry,
"Co.IiIIMnu:::tion ora large fl1iC~~ityneeds :Imnnry nevel parts and teebnolegies," says; ]rlEP Director Chea Hesheug. "'M[I~leriny rile pIiO~CJ;S grcirtly iiInpl:rovCS ~ltd igenous ski ~]s."
Project tead
II
Or~ilililati{ll'l
Ellterronjn, CA:S
A'!Ii~llioo RernlDre~,nl>il1g
5~s!Mi1
In!!iliitu.w
'Q<~
A.rh:tr 1e'.!frIin~hO\!,' ~omake aecelerater rmrlcs. [H E P l'ii~'!.I' CXI~tlr~l> ::iIOl1U:: 'CO!HPfll~I~I£ m ~'';~I'~Il. Smi[h KOMI. Wl!dUt~ Unil.0d St~.~~. H~ljJyedby its success, I3EI?C has auraered fril;l:l1d;:;in h~gh pl.I!C~::;;h;:; VU'I vi:-;gt1:Jf log is u Who ·~W1.10of Chi[m!; poUti.ca~ ~.hc. TI~m.has
Eleijillg l!J~efSi~¥ 01 S<ien,"e' @lfId1r~dh1!IQ!g.giy tlelei IHlg:h "Ioagn eHe FifiildllLab. CliiS~ IHl!.ll~'lhOf!g Ulli'-'tJsrt.~ (If Scien~ .iJlfld i;ecjhH'!i!l)!~y.Wuh¥!
MIA
S~<lir~
'Cil
25
2008
ke;pl the fundo!<lHowing. In 20(m .:~~.~. .•!1 eenn giQvr,;:rmn{:;Jill apprmred SE~ million to r-eJlovont:
I~'n.c facili[J'" 1Ii1be ~upgnllJ!.l:~ca~,~~d BEPC l[~ls
scheduled [01" ,comple~ iou ne.,,'H year, lts PQ1r.!.'CJ $l~nuld~ncr~m;.,;:OO·fold. ~n~.bli!!l.g~\!~r·fin~r ] rneasu re:mne'.r,(;~s '~Ir t111L! .~ep~on rLK'lS!S., [lIowmI.lhe· mad Il ~EP plml!i to l')ui Id lhe $2biUiOJ~ :1Bc~j.n£ i Tau·Crnnn r.iCI:OI'y to e'.">lendi the,IC(-elernlof enc:rgy ~LPto 3. GJ1:V far (,_'iipe'rrimenl~ 011 bOlJ~1 ~h.e1;m.iI. lepton ,Hidl dli;]l1Iiill~llmrk., And the ins:rilute is p'lrlidpa,.i~lg ill the h~IL'fn;uio:nl<l1Li~C'~w
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Wind TUii'll'iel
WA.
··fbnowin~\.vh.a·[ ethers do" H) "oC]D1~ing up ~~!ith ·\\,h.1.Ito do:' N everrheless, s,eve'lii1I~:L,fl b science projocls<I]l~jea!L' to be "rue-teo" FaoCir~i~· .~ ;~nc~~!d CSNS. '1/\11 ichb~~n~~ iawd 'l~~.r cueb.ii~ni~to a p~ Qfbi IllilHl.1 ba~I's:' :-K\)!'f> .Zh'1\~, ies, iill,g le @ tour [:Im ]000 of [SIS. a pulsed neurrcn EIII.clmprolt1iUD, all shear 2:0 to 30 ucutrOJloS o!lrthe c sou rce ;It' RUIh:e~tl!il.-d..AppteronLabm11.t{lfY tdr£e>t g:~~.ler.!llif]Jg an intense ];ml:se~u:l' probillg nemr OxfoJ,\(t Ueited K[l'Igcl.OiIlil.l, m icro st ru c III res of S~. pe:n:o 1'1 due tors ... ltor instance, Oii p~)I.ehiil~t Phasel CD[l,~m!..lC[~flll is AC'cfirding 10 J K. Z~l~.o. ,1 senior scien~]st;U Oak Ridge Narieual Laboratory in eJi:iJecl:ed to Iflike 6 :ye:ml. Tennessee and Ol. e~l~l1!~ld~:;lm 'SNS. tM In Re~~rkahiy. 'CSNSwiU h~ llUiH ft~r from W2IS .impn::~ssed by !.he nmhjidiscip~i.llNU.'Y the Imd~.I.iOlI)131science stronghnlds of Be~ i.Dlg .reKC~.I'eh III ISIS and ~Ib:kodCA.S's lnstirute and SI~~llg.h~i[){'jnggl~!!Uli. ~I 'C.ily i.UD. the prosof .f'hysi'L"sror'l fe,1sibi lli ~y study of'J si mi !orr perons Pearl RiverDelta in scurhern C'h~ml. offered ~i~ii; md and $63 ntiUion reward h 11:1C~~H~ Chi ilij" A gIi"OI.J1P' rif neu IrOI~ selenil1 ~i.s.ts en Iisted Zhao and othel, ovesseas inlrastl·'ll.r::~ure costs ,"]f's a reasonable deal ff)f Dt:mggi,J1~n, WIWfl;': =-t.r(~~g ~~flnomic experts to h~lpdr~lnl.:S.IS-i.!1~fl!irod plan .s .. CAS .~ppfOvl;.ld the cone epu!l.'I.l. d~~ig~1 in, gn.!wil~"md w~~lk research c:!pm;il)' result in ~ligh.l}' ull~}'II<lm:edl d,evr;:]op!lnenl." sa.)!:!; .:WOi5 !1:1m.d nded a prepamlory fru leanm, led by W1I;lU of BrooikluJ1vefl Nat~omll Lab~mro[)r in, CAS"::; Uarlg, "MiI.I1y s,d,emifk 't1.~e.111S ]ifl,ay j'ie Upton., New York. i:u~d~;!~I:.r. to stan on :'Ill be anmC'!0d tOI:~M::dty." tl en,gilno::ering de.siglliD1.1]2(J{H'i To ensure reli;]b.~Ahht'lllgh .Zh,1O \\'Ollde:1rS hen!,' rmmy~n~ur .~ ity. tllne ~emm.l wi~1 <ldOpl m.U.llI:e t.ecl.ll'lolo,gy sci.elll:is~:s willJi f'lInlil~ies \..~i:Jm reIOC".u.e to Dong= .~\II/~.\;;:Ui(.eve~·po&$.ibll;:. ~I~IS Fl! Shin ht~ •.,~~~ ~~ l;U~Hl. hi!;; i=- C(lnnd~in lhO)iluV1l~versiUrl;ls~n § d!ePlll)l ,It. IlU~P: The pha:re I d.e.mgn Cil~I~ for iI. GU<ll'Igdo[lg :andH.ong K.ong wi.U pack the .:ll Ih'!t\'~lr ac~l~r.n,~)r)D1d 1R!llIkl~)'C~irl,~ sYl'I,dno· ; Itl,c!Hl'}I \'II'it~. Y.1U!'!!i:~~~..f!irdhtffi. ~I $I;I}'s,~n~(j!i ~ ,~ lron tD s]~eed.UI' protons l~) ~,6 Ge V mu~,del iver ~~i.enth'lo;;rom Be.ijingwiHlrairilhem:>:opbyti.::s . f but!'lI;;kllml\'lodgc.~ tha~ "~xiPCf'ts S!I:~ltby.IHEP ~mulliii::tJ be~mpt1\"~!.' of .120 ku~:rn\~~~I.S. :\,'1]ich c i can be dOlllble~ .~nd qmldl:1llp.~ed ]UiI: 'Iutu;~"e O'I,loliI,eent .. ilnly <Ire :!m.ot ellilOlJg~,'" So CSNS wi II
.",
tion en Adv'lItced Neutron S(:m(1;li;l~.dIe fin>l time the evem will be held .ill CM1l~"
('h~l'Ie~~ ..S[II:tJ\1i10men;.:i. I :l!iIu:.,m;u\'~lile. ~:~uld :l>OOiJrn hear a decisinn on dl.e .nnmln:li~ NIJIRC
wi II :spe!'l.dl OJ1~rl~~ '\'\.itlrl!d~~1;:af',4C$l I~!!e i~l si m.d telescope, the fi'i.le"iIi~Lr:lJd:red·lneler A:pe.nu.re
Spherie"nele~"t:upe
firsl proposed by C~m~,:lm.3.·:!iNSllio~ill AstrunOllif1ic~11Observa UTld~s (NAO lUll 19Q4. would ~H(i\'" <'I.:i:tromnnern ttl peer f'lr~hef than
ever
~~:rl!ol
NAO c;ho~ ~ Hm~s~(JJ:a~hl~;t d~rr~~siQn l 1m!ooU.th\.\1'C5·r Gu~i~{}~~ P.Ul)'!.'ifH.:e .u::;tll;l.sil~ 10 sm.p.emlilre<ic]\! ~ dns~.rn!'If~e of 400n tn:i<lnIll,£; gul~r pam.els. t~ki!l;ag ;Rlhf.~flI~]ge ofd1eIDI<li,m.a11
lOpograpl~y :to lreduce com;.tillJClioUD. cost.s. ]~ T had been ~.~.he Wiltll in,g ~o ~l!O~~' he AS t 't S (I ~I.al"e' K.~IOllil]le ter An.,,:,,!ty •. tn'll :iUIll.le i,o:II1I:I], rl,l;U fluho (Isl~'lomy proje"Cl U'I<1U would ~!l<lb]~ <I:l>lronmners 10 Obsel'~''e Ihe' formal ion .of the
:1
such
as :t\Jng.~le'UiI.liO knock
opefll ~ts. doors 10 ~o:re.i,g~ r~SiCJr>Cbcr!;., For s~"ne[IF£. DOIm,ggmlll i ill Apr~1 hO:r>l Ihe ~xtlli M~e'li!llg (l:f~l~ ~nl'0nl'-'!liq@~J C'oJI1olbf]iI1't-
'wm
'COIll-
e-~;~hn;lt~s
SC;II~Nelc. va L 31 S
9 MAIRCH .2007
NEWSFOCIUS
be'1!Ulh~~~'~,[I tlme as needed," -~;;IysF~.dar! di.rec:~od
l)hys,Lci'l.l Zh'~.I~g X,i[JIYi. who Sym:1lW~!I1On Radiation lab (NSlll ) in Hefeibefore l1l[wil1g W Sh;r~gha i."But il!Jl Olal1l1'l •.wC' ~l.,~leto Ima Id ~l~ fuci I ity as one big 'rrnjecl because allthe flliiLdilll£: e(jilii!~~$ ~itlgll~ehunl\:." iE~~.
IJ\tC Naliona~ [11addil
U:un,ivlershy
in~. iU~\1I'fiue;',~ndc: an
~rs~1
J~1.eilily S
ope.~t
budget
illg buclg\f'l. 3t
occasions,
o.Uile-lcuththe
l"OllS1u,CL10n
thl;l !~P(;;f(!iillg
thnt. FASTwi.U cost :lit ~eil£t.5;86 million ~nd ~'lke 6 )'tlou's or more ~o buikl. .111eemparison, lhc gestatien [hue oh~l,e H,igJh M,1Jgl"ie~i!: fie,1d Facil it)1 has been muc hi $'~lorit;':r. Scren~Sl1> fin;tpMflos~d'l~'ncbi~ ~ie'Unc¢ p~uj"ct:ifli .uUIU~l!l2:0()4,<'I!!lidf!.)l;."!;;]v""d N .c>R.C apPI.'mm] 'e<'lr~:i,e~'~'lhiis ye::iIJr.nt: lacHity will be hosted in h\iO chies ~o 'If!ke <ld'ii<lmuge efrhe exhidng k:d~ll~tmll C<IJJflb:i:Mly of each city:- says, Kll<I!iI,gGurtngli. dil'iXtor orCAS'\, Hefe;i liign M~l£nelic Field tub ij,l A~]:I~ui Pli(l'vilu:e, H(lF(li wll] bL;; hoU'lu; '~,!J l5~ till 40-wsl~ ~'I'<lle'N:()ole{~ and supen::a:l~dwl,cljn::, steady~ fie~J magnets, priD'lHlrM.y for m;u~r]~~sels
ence research, saeh as p:ll'obinglhe quantum ~,4 m effect,<lud for ,magnet ie resonance ..
w~U cast :1;25 rhimi,on and is expected tetake .3 }'C!l!1' to Dom.pl.C'IC. P.r'Oh"'Cl :s;~.iel1.ib;ts~{llv,~
alsu pruro~~d.r!ll
Meridian
is determined before consrrnction begiin~:' 5<lYSC AS"~ ,li.mg. Thi,~ ~,m~,l.l:;,trillgl'i reseaechers when unforeseen problems arise, r1.1r~'Ii~II'l~ipth~,\.11enNSRL in the: I 9911lSen(.'01,II'Mj. aered trouble with beam srabi lity, which ~l\:V;;1n:~d~p~rim~i.'us. ir.~o~~'"LUngbud.gl)'! \'~'" insuW'ic:iielm:to C.'IJ,"ny om a fix, NSRL.:S sy~u;hro= t troll source ~\I<I£ stabi.1 ized mn2;004. o.un.ly~t:n:C.1i" NDiRe ... prtl\'&! $15mil.liml tba:'!fml, p llpgr.'lde. ZIMng "!m'Inllts 1.0 a~)pljl I'CSSO~l:>lci21m@:d ~rmTi m'ln.lgiing lin: Hefei sy:Jiid~il.'O'tI'Ori to the consrruetiun oluthc: Shcl'mgh'fli ooUlrl;C.Hc has :;;ugges!~d t~l NDR(' that ;;i~I!l1~ !'!1UlH:ybe set
aside so users carl bid to des,ign lin.ei:libealll~
.c;u,'IFCI,e fmg:mllil,n. 1:0 1:~mll::; muiomll ar~Iy.'ii ~Jf greund-based lfn.oni ~;(~rs'o ~nh,Hwr;; SI)iUJe envulml~1,m~'~1t cil).~ell""'ltuon wori.dlwi.de. Ol~er b~ SC'iI;IU,'~ ill i~iaH\rt's. are ~~s~\\'C[ I
lines OIm.1 c:>\pedm~:~~~~~ stations. However, he snys NIDR'C p:refcned to stlck with ~1S;
usual [Ipproo!c]:u W <l!UlN'::;J[in,g mOllie)'., .Zh.mg sd~ I hOIJIil:s.lhal S SRI,:":" mElIl~lgeJmlen t ''''i I~,be fli'lor(: nc;:x~itll~~it has; akL;~.dy agreed 10 lin him d,e5i~ll h.is own beandine for eesearch o~ ~()n.dJens~d ~~1.lIU~i!"phy!>.iesand s)lrl(.':hr~)I'mn rmJi<Ii'lio:~, im<l.gunl~with rundillg
defi n'oo ,T<lke ~~~e "timared $1.25 million '~}J:'(" ~ t~'i[tl science ~e~ll~b, I~ilhy," 1I1C' AI! ild;~hlof the SARS mnlm:<lk in2(J03. :rurter Rw Zihe:s ~M~patQinghu~. Univ~rsilY ~~B~aJi!!Ig.cl';;!.C~ed
1~IrQlIf:I,1,
r~eM
Insdune
diLCeCWl
lx.'C(JIl,~,e
in 20M. But ~hep:llojeca derailedwhe:1IIl Rao left B;~ijing la!'i~}l'('.$ur to b~(]m~ preside!u (11'N~i1k,1i Ulli.ve:rsily in Tianjin, CAS Vice President
Thmnk:~ngwiid(!j1J'
\V~dl il'~C'~I~'roj I,;C'I:~ l>1'1:i)~~ti"iJgl ike bamb eo I s:JiaoOIS <lifter 01, spril:lg rain, biS science is no
In~.~~'[' fh~~()l~ i~1;.\G: ~~fCAS:~p.:~~}'!i1.'I;· rro'.. .. 1Kd· ences divi:s;ion. Oth~i;L1ew el'U:~~ie5 ~lNmfiil;lbbil .s,lIch as ~lg.rh::~l,:IIUl'\e. bilology, geoi.ngY.!Ind
re~,1Lme ~.~tli!lmg. One oftillC firnl.p:rojeeu:;NUlRC' finalized, in 2005" i.!l'the' Merid iml S:p.~ce We.nhe]' MOII~~tol"ill].!i:rlrojecL ]I :111;1::> 01\vi.de :weach, fromlh ndam(l]ilt;;~ a~Hophrys ics-~aud~& I ~!
Chi,Ollll!~u ~l~~ l"Ji,vTI;:IPOO<oo b~~i dl~ f~iI iry ,H Idi~g S.h~llgha.i:S.Z~~ngj i~ngl ]ig tu- TechPark, nesr SSRf.lu mlk(:m;h!a.mL!ag~ofl~!~c .~~.c\'!,1 s.Y¢lcl~ro~ tron source, 8m lImn)' bi:o~,~;i,,,ts ~<Iive~id dl;Jt" protein st,:i~Mje<e h. man; I.h~n ,ci1"sUillogrn~~hy <lInd lIml tline ne\.\f centc.r :$~lOlll!d:1,101: 'be so far foom ~'X~sting: iol~}~k'=ll i!1c... tiiML;;:i, b l'h~ ~J(l.iitic.. 1tussle is he~d1ng up. S(lIIIL"\:l;:~ say thin Qr:l.el'act.iof<l: is lobbying r~r the iadlity ~o 'be bui~l do\),t,~nolwml. m:~lf("ASs Shra.g;bi
I:lms[ tllJte-s li Biologk:i1J~. Scielm~<" ... Z~lmnrut!lllg D o ,j S [O() n"f wgo for e.1\.Pt:Jil~l!enb i r you M:wld)! prot:ei.llu\ ~n li\i:~n&!:, 'I;;ells." says one ['Cs!(!.rud.l>:'f,. O~hGrs.are l!J,l'ing. 101 Cflf~'('l off liunds far ~~i!~1illg labs i:llI.l3eij illIg. They ;u'glte l.lial'll ir too ]lIludl
BmmrlM would ..oo USIilI1i dnrJ,Ol have the Ct1'l,Ipkof:mi:llion dollars ff"Qil"thiskine! ofi:lmst~l1I~ ment work, I:HEP's Xian :Dingdmng. one of Chjna 's tWClre~]N)sts.y.m.c1~ fI experts .. says roue !btn I.;~ti!i!g m~~f.(yii w.~rs~I:I1~dPJI!w lin C'!11'!S,IIl,IC'" lion and 11KIIl~f'!eme:mt is popular i~ rUI,-eign $yIDlcl1rolr~nl abs, b~!~"uUJ' $:ysl~m d~if,;~mlnl ~llm~' it Ol'gm~iZl!LtimlS ~""am.to (iOfll~ro~. ;El1.llhe
rt~{}urecs th.\:y'vegoC'
Xian hopes dl.tt (]li IIiW~ '''lX'L~~'1It~on1lim~:ron S f Call dlei ben~r.Accnrdin,g ttl F~r, ('SNS has involved users IWUL1I.lthe beg~:ll1nmi'L1lg and !1f'!.(]dil1..'1,1 d~;;;~~~~;; (;'Irl'!,:ri!lr~U frO!ll I~rg~ ~!~~~I! m~~[ ing~. ·'Weultlo learned lessons <iixmllil.t: opeWo'It1llgbudl~1 :Hrom pn::v~ous j:lJI\ojeclS,:' says
so~~rc\;:
::llIU:! Im~illMjUutCe
emme;~t.
G'l~'~:migdolligPr(w.im:e ~!lr~iThtl:vas h
p~Obi,Ulg 110\111o~m fI~res ;andl.comna~ ~.nas;s s ~jecliOtis :~!!:II1U!,ml;l;; E,whts, upper ,,~~mo:;I"' pbere-to mon itO'li"i ng lle'3:11"- SirIll ~1',H::e to E en,-,I,!~ safe ::mtd~it~ o:pcr,~t~o~ .. R1!Lll by CAS":; Cem,ter. for Spacr:;: SCii:ence ,lind Aip'pHi,;d Re~furch,lh~ proj.,;\c~.\',rm equip
,Ij g:ulOund,':based obsel'vali()n s;~3tiom;.wit'hi
nHl!'!i,Oyis
W<'Li>~e inevit.,b~,e. is :~ ndced.rC:::'~!'\:!1 rC~lers 1~C"1k.1l0w]c{]l~e 1hat. Chimt's IlmG'il"S sy8~em fbr megaprojects has .<)
6
0,.;1
htp1.l1l'Or SOlU~l
]!l::;t!:~inlcmi1i.H)
r~CON
""
l!i
9 MARCH 2007
:5CII~.Nelc. W'l!Vw.sdenooml1l.g.orgl
NEWSFOCIJS
R:ETHOVIRUS MIEETIING
II
81iliSWillNA
G~QiilfOO,,"
Sifir4iir~ ~<Irly \~mmhlg.lJn~[~d no he~lU;ln[+"["h~ study. which compared abrupt weaning ill <4 mo.l1Iit.11so beeasi-fecd ing for an averilgc of t I() mum hs, found jU'l equal nnm oor .[1 flnV ~nf~lirms:. and dleamhs by 2: ye~rs ofi:\gc. tl~.e n~n!lt p~"CJmising new.\l .11: IJ:lelJ'l.~.ee~ i~l£: c~ncfn:1U' repo"~Of~fHrg~ ~'n1caeylr~ab; ufr'!.w A IDS drugs, ~m,~tIltll" ~'riJli"li'V~l[~ twit), ~"thi~ yea I'; th3t hmn: :ml.o\'e·It,,1Irb~l~. of the d:mgs.. One rnhc:gmvir. nmd;eb)! MI;:;~k & C~l.in Rull\!,·.W. NeiW Jersey. C'r.~pple:~Hrv's ilnl~~se eillZ:yIne. In studies ul'niil.,jd.y }:i'lJO pil!lili;lm5~'l.rhu had viruses dm~ were .~~£M'y r>e5.~:SHrlrn:t m exils~i~.l£ d rugs" abm~1 60% I~lil.dundetectabl C levels of 111V ~.ntheir blood after.24 wee;b; 'o~· ~"~~~meill w with mJu;).grn\'1rfl[ld~ln···upHm~~d" c(]d.;(~!ilof other dnJgs. I n the controll. ~.:roup,. whieh reeeivedjust the Oipli.~uLredreghn~ns, 'l.'ina:ll load'sbecmlll.e mlcle~OC'lt11J~.e in roughiy .lO'K~ The ~Q~lld drug ..n'0fI'IJ',.Iiroe.mrgel~ hlU1fll8in imr~mll1.e cells It'o.Ilher than lhe v.urns.. II [V egll~rs cells by ~u~ld~i~.1!l the oC04 f\,"CII;:~ptor OJ~(: i:o ~nld
rrv
.mJlH'lK
OCEJI.'1
~(lU11H.Jl~R1(A
2~
mrs, Cn~5 or CX.r:1R.4. M ara v irec, 'IiI~ade by IPnze'r' iIII NewY()J.'!k Cil)'.
Ihall:1 ]V-i iii reeled \vome:IlI, use Ibrmub .. ()~.~Iy \~~lell it is. ''<'I:(X'epT'db:le, fea!l~ ble .. ~n:Ord.d)]e. ~"i~i\iilftin~hlc. ~11:e.") Ne:l!ITly and ~\~.·o':I,hird.s. nfected mothers ill !Jeil S.Wtl:nltl ofi !l.l1W use fonmLll;.t \I!.+!i,,~lln Ii>,pr(I\'!~d!ioic]free by 'the: '!!! governnn:lll. Brut ~hal ~)oiUcy3ppe'i!U:s l~ have ; h~~d3lr.;LglcdtlwIlSld~·.Rc:l>C3rehc!"1l !>tt~.YI~lg au
~f(IS~ recommeuds •.
<lire IXlrpin~MC'k. tip. The m~!iild~un~ltic wlk 0".1:1 alt1;:n~~U\~~S to bre;<L~t~~~d:iJlg. focus€:l(1 0:1.1 B ms.v.."!L:l.1a.where the g:ovcnunenl advises aUIlIV-umlfec~ool "".\Omen to ~u;r:inf;r~tI1;~:rmul.., to avoid trn~.lsniliui"g ih.e vims, ,~r,~c XV{)I'ld. lth Org,ani.za~kllli, 'by COl ... Hea
c(i~kl:l:gyes s~ur:li~d l'Uilorl i~ :; ,20 ~ of L53 B()t:i!1l'l."I~lnchildsen '1i l'S; ho:;pit~liz.cd far d~!u·ullcn" fii UI Of tilll!Se', 33 cljed. and .aU 'IE :Ii bUll (m~ \M(\f~ lil~mi,nb-lloo. :i!ll: nle~nV~i fected !.101tll5 of n the motber and theillf.\mt. W'1:i!<mll. as$'{_loCi~te,J i1Nith;my ol'a~.e de,alilins.SI)]ol s~Ulm;p~ shewed widet;pre,ad infection 'I,'I.'i crypto5pur.~d:i'lnm.l. Esdr~;riC'M(,l th edt. and S3lloo.~dl(!l. ·'Ifs.a r.UIl1I~ii~@:5'lorny:·Sii'Ly:s rtt_[i,..~trficktn ]-[fI\~nC(!n\'';;idj,o ol"l!llu;:Uni\l'(:rsi~y
reee prors, rna r<'IJv:u me dm\le vi:mllevd s drn\!n to !I.1nJ~b;;:clablei!ll 4S.5% of l.~l.CI.n::<lil,ed roup vcr:sus .24 .•6% ~r:l, a oo:w;tm:ll g group~~atr«"eh'ed op'rill:1.ized :ltlegil~].le·lJqj aloue; David Cooper o['du:: Ull;ail.ve[s~ty of New SouLl~\:va !es h11 Sydll.ey. A ustra 1i1Ji.\l;.Iho has
.~ e~~e?i<JI .cr~~II.y.o~L~~re<~k , of d Il"ir~~C\aI~ d~~~~~.~m'i.t ii5 ~ye;l'Ir repo rted I:h<ll infllmt ~~ofITh~iI ~~,~~unl'~l~'\d ~l. . tn .~ bn:.~54..feed ing.incrl:"L~d ~ a l~ d's risk of dei:!,llh hi~ ~ t~wmd i ~nht'.~ di :fiet1,::i~ I :in times, ilike~y boci:!U~ 11 ~ severe mood cont.H1Ill" 1lJ.,u,edlhe W;J tel' m,ed 10 fl m-Ire III > :r.- m1JLll:llI,ii!I, ,_,L;; 1,;1 _",;;!.,
b:.
orK:w'aZrd:lli"N~ua] in SQiulhA:fril!~l, Creek's '['I;;<IIn has advised Botswana IU ~e\1'a]:mne ~lS nlliven>al fOIli1l1!1Ii;t pill ic~ srtrcssil1g th!u Il,~ of vir-al. u~al,~~uu& fitUlflJbrea_"triSl!.: s ion
tested botil~.drugs. s:::~ys f their potency and i ~(l~~y hold up, (:I~nki.u·l$ !'h~lUld reevaluate
when W gt,ut !:reatlmi.u:nl. CWII:rell~l:eoomm.end:Jtions cnCQ~lmgc dcla.yijng[~!n1'll~fn IJlmitil~rieus ~mn,1;UiL1e d'Jm<l.s:e Of.:ClIIiS. i'ml~u't lO redute
feedi IlIg
t11l11::>t
f~,g:f~irithel'i.1
Hm -
But
Ji;lIl'i¢.' M~.I.~tyrL;:•.!M~
§ g
12 M
ch~~dren n~ dli'U,~·he.l. u~ .~ fro[!Ii'l. 21 tile yettr before. M:Cdical cpfd(:J'lflioh gi\5~;Tn:cYC~ek Qf,t.~e .S" ~O(:DUt'er~1br ~.il~se ~
:s
owlJ:lrici'~!1"g)rni;':Ccolo.gi Sf ~[ :Bari:li.gwmllifllhllo:sp.ita I ill1 Sowe'l~l. Soulb AI1riC.I. U'lul~on~ddl!u reseru.iChcf:> ~1;ru:J po]iJl.}r.mtl;"~rs s:ilO:lJld !:1m :~ose sight of ~Im.e real gO<i!I: w make tHnh :lOOding aplll'O,tche:s S;<1.fi::r. Sev~ e:u:id ~lud~~ ~lnd'ef\"".lY 1!:~ a myri<ld or !>1:ljlI!Ie~e's to do. jm;l. (illat dUS.i'!"i.bl;i1~ing !l;:h~.QI.ri~"'.ion S!ol.u!io.~J;
dn:lgs in ~.Ulot~l!i:n; 1Ii1lli .... <L~~d
too:j~(;~lh't~" mnh~tmd\:"(lilf~s 11~$ B :p00~lJ~'~~ sui:' fe:r from~he pll'n~(mged i~.n mune sl illuul<ll ion t1'b1J[ H IV ~rigg~n.;.whid.l Ctl(~p~r~y!> is lik~~y I'i:$pgns~bllilb the ~ncreu..'ie~number ul' d~I~:l:i: fI.'Ollrl. heart, ~ iver; and k~dney disease .... If YOLI
:-;~;(]Jt
pCQple
f;<I'~
ier.
ytlU
11'1.1 igiu
tmw;: a sign]]
~r-
~.]l~
Utlh (i(l n I~r@I1ICfl'OIl Re~lr'(!Io'IrU!!lUlS·OPllonu msuc ~,",i:I ;~ rllfecH:I!IflS,25~2.a Fetl.ruary, los:ll1il9ete~, Collihllmia.,
I;CClOO. b~i~ Ino[C ~l~ss!\1dy. B1,l~<l Z!1l1"1bia!'l sluely pr-esli:ilated ,It 'tille mee~'LI.g nnql1ldll~.n ·one soollll ngly IogiC<l~\..~y m m~:k~ bre~:s~.-~~'edhlg
impact 0.1111 all ofl~sc:·lmc s:m.y:;, rOlf t!l.e V'<l.Sln~"tiO[~l}1 ,ofHIV-jnfccoc'ti ptlClI):i.ewho live in poon:omnr:i.es.,hm'!,lc\-er.Lt Irll~jl b~ many Y~;!.I'$ b.efbrl<: mo!!lf~wilf(J(~and ndwglOmir ofler.my rei i.ef. Ma:nh.i'lll :Eg~'r of tile 1JJniv~rs]lY ur Bi1;;I1'[il1;; in Sw i~t'Kr~~und,I~Hi..· I;:~d h~agMt'!l;repo.l"ted dmt 59 d~.'lfe·renl Jlmg n.'eg~mens .11"¢H\ia.Habl~ t·o rnoSil AIDS paticlli~ in Nordl. Amli:Tic<I. but only three <Ire l}ll'ic<lIHy <lv~li~.aible'i~lAsk~ a~d l\rricfl, SO~\'C (lll~pmbCi'lIll
I.em, <IlIlOlher
5tnJlICe~,
I
urruedeut
tfl
bumans, TI:i,egello:m.e of GOJ'{I'tlufax f)oMjY!(,b·.a. !for cx.3Jmplc. is 18 t U[!llCS tllilesize of mtrs.. A ~(l~l!liOI:l sons emer<dro in t~le 1910N~
h~w~ bi.gg~'rge;;'lO!n~S
d'!an
or
EVOUJTlON
so-ealled junk DNA ..In a(id[Gioll to preteincod imlggienes,.~e:lftome~ l"'OIlI[!1 ill ~I:re!che!' ,C) r DNA th~~encede RNA~1~,oICllt'li.l les or 3f¢ jW;[ w~[ig.~ 'of old g~n!;;$. Mai~}1g!!ni:lm~~ i~ltud.. ing our awa .• u,"e dominated by vjrusl ike ~~]lI!~n~~:;. DNA !;",II~d ~l(lb~I!~ of ~~rw,'lm~~h,ll ,""Ull l.llaH\;cnew copies of themselves th.u gel im~m;.'ldiuuew spots ill 'the same g~IUJme. Th.e human ~lI;lnonlt: is 98.5i% nom::o<.iing DNA.. Comp(lrimJ,~~I~t:::g(;li1ioln~ onu ving J';p!.):ces, i sc:i:e.I]j~istshave tound Ilmt geL1.0\Im,les ~rul,ex,lXIr:uJ and :;,li!J!:~f'!k qluk~kly. with !m.lobi]~d~nl'!l~lnS spl1eadilng .~ikre;1ge'.~.aIl1,~,ic lague'. The ronan p gcUl,OIllC. forexit.lll1l,J.::Ilc. U".i~:l,.lcd size ov(:,r l~.C iun,
hellP~lllg sdentist's.
SOp edes
t:E!,j3j!iIE!
,aJpallrtw:hytl1~siZie;$
Imliiid, Cill~yiiU'ligerro:rJ;c~n(~I!.~h"e(:ell.~1:0 !i:lJlipout Iotrge cifl,'t~u:Jhof li.om::O<iirlg DNA by accident, !>hrjnking their genemes iI~ tlrepD':f~cSc". li:l te~t \~~~d:iI;:r ~..,t:~~f~,~ selection pi.Ol.y~
.1
1:;::rtm.rUJ,\lUWUS H!'\". it rums Ol!L~<ld .11pre~ty sm311~ genome. A team of Amerlean aUldl Brirish~ie~,l~bl'S eM~mllresili.n ~I(om<1ined Il, r>ei<uve~y puny 1.9 bill ion base 'Pa~rs of DNA.. i a~~uk:over ha if I~,(: :SUI'_(; uf'!JlIr. o\\,'fl !,~1l0IllL3. The !S\:iem~~lS\J)j,'hocame lip wli,l:h this esti!'!J:!l!;1li!,.~long\',\lj,lh ~IU!nmM!l: filf U'!l: ~OO!l!l~." af .30 ether dfiinosmm species-s-had ~,o"lllc:ielU DNA to gtlld)r. .1: 1:t'.V,'lfl:ea:' iL~ll, beesme e::o:tiililCl
,(~5mimofl years
goncl
n~lead;
Eong
ill'l~p(h::'ton tracing the evcletion of bird gf.!nal'bles. "13 irds arc d unos<l'u:1i"$;hey·.-c' the :~mH t vestige," says; O'rg<J[l. Sciemisrs have long ncted Iha~ birds have ~rI1I'l.U genO[lrnl!es. cernpared h1 ~'~ptH.(:s.. thL;irdo~~S'I: ivi n,g~l;ni\'C1O. [ but i.l.,';"'.ISunclear maow .lllrn.d\\i.~eH that cm:i!ln!1;l eeeurred, OrJll:~m's~Hudy sl~ggl.;':~'tsth~t 111i.$ d illostlurm:toe~t()rs of b.iooJ> lll<ld evolved S!Im~OI.11 geililOJ,ues lo~g before bi rds tonk~Q the sky. "I, thin!.; !~lS vcry c;,;.cidn~:·' saysT. RYilll Gregory, "Ill expert 011
£;el:lo~,m: size at the Unn.vell'" sity of Gljelp~l in Canada "[f:s thekind ofp~lper we've ~l;,~d~dl(]{i!<1 l{l:~g tinl'l~:"
(iiillltl g:e:l!l,OIill~'5,il!ll
:S1'Wmlgole ill de~enniHung tlre size ora speciet;' r g(,llm(lm~•.&ciemisl:S h~~'!.ie~Omp;ilred wide ~I!lg~ tl of species, sC'd'rcl~imlg oorrol13tlimlS between lihr geJrnome sizeand ~JlhefHili~", Ill'[u:t:mt~]~l!bellid~lpIi ve ,.findil~g lil1ese (OITe~:Hta~:S ])mao; been dilfi(lull l~0'!.1!ro.'\i(N~. because rc:lruiwly few .g:~nomcs I13d been mea'MJOOdLllll.lil :u;eClell1l:ly.and m"my of I:~O);~ m~~~sur~~~IB'!~~S DUlIn b~\\I!"tlng.. turned Gel1l!Olne sizes are ~:sy to mi,~jLldl~e ..even ~~'ilh
moo:eCillgellQnn~' seque:unc.ilng
they disco'i,\llred a re'\i'e~:UUlg C>Qf~ rel<llti.on:: Big ge;l,lmm:slJ;."<!u! to berul.md ~llIag,lp·, mal s wi~;~, ig bone cel~.s. By Qom~ril1l~ 1~,1Jeize b s of Cli:11~in (fIil!~~1IS'-RLr fh~sil1ofifl[1i1Ifi.~ fIn ivingflnlI· m'l!li:s,~:heseientisrsgct S~ll~sli,calllysound.est~. ffi':Uil'iS I:ilf rhe !,ij~ ofthe d[no$;i!ur g~!I.~!:l~l;~ The fi nd ~ngs. lJubhls~led by /Va wro lhi is week, ~re IflQr~ tlmn just a curlosiry, Chnd~ OD:'~lln. a ~!'II.·\f.tl1d U:l1iversity ]m!ool'ltoloSi~~: ami dl~ II;~d ;';~ilhor uflli1l..:: 11t;\7!,I p~~r. Sf~ ehe eSlim~le5 slil.ed IIlle\V~ig,llilon" big puzsle: Wh,y do th~g,L;ln9!~l~~" vi~,g sp~d~sI;.'tJfi!H;: :~n ani :;;~!C'~
'U Maggl;t~:iu~g rdrilg4;l uf S~l>I:::;.V'dII:y,~ng nillm'!;l tiUHiI,
merho d;o;,,'liVhen sc 1,1;';]1.( ists sequence fI genome, I:h,ey gener.d~)' breuk it up into rdt;::,C~ t1mc!ml~ng~the'r like .1 puzzle. NO:Ulcod~.llg UNA ii' l(.lad(':dJ!;,v~~h r~p~,uil'lg,
sequences, wl11ich ~'1i'II.'! dli.Il'ILl':uh hire&<;..~mhk:pll"l;lPC'~~J~ Things aee improving .. i!>fiy ~ G rcgor}i'. N ~w leehllil
f~aglHenl:S
to
~oW~Y'
(f(!atl!!l~s:
iPi\':CiS~r!I:J"~'{~t'lure!mJ;':!1i~-rnli
instance,
11)ISgsnome
is
15'0
li~le:i!!'m(l~I:erlhmrn
bled hu~gfb;h geuome Is ]1 times bigger" R,eceE:~.iy. SOilll,e ]m-ge:-sc<lle eomparisoas of 1e'II,lOm,(! sizes h:3'!,'e suggested liKH, I'I<1Hual scl~clh)!'II m;)y f~'",or bi.g g~no!n",s inf;Om!3' s,per::~es ,mdi smailgeoomes in alhers.., :Bm SO'Uil~eikf;!'p~ !U''!I~:U1;'i th,Mt ,~~~n{lmeite m~\y n('i~ & ies s be <'Id.<I:pli,ve <lit oaJL Now. w.utUn. the ~dwe~~,l of wh~t o.~1Em~Hk~"wC<lU "'dlin~ogICllomi.('&;." Sid~
p~~p,e' ..
NI.OlI!mahil gr,as5StO;lfIper (A:ldi~Q ~di!!''''li15): :Jl6.Sbil~Q:i'I lip 'Figie;~~i,al'li!!al!i!de!i'(Ambylitomo tigllinllm): n billiolll b;p
~h~$Iil:i'~~.d~s are i
s~orod
i,1l1
~0l,:V
being
online dat!lb1.lse!>,
!li
eilli SiS C<lDilI be,sinliO ~eilse $ome <U1S,\iers by mn (iddifl,g c~li[1dSFedl~lnl:hc emcrghlgpi.cl!.lr¢
of g;e'l1<Jr1J1ecvo,llmiQDiI,.
!1n~l:hDNA '~$ f,ound .~. .~s. .1 cen of th!! dmue~tiC'Cbv.'.I. a flu·more' highlly devdo~'ti, anirllla!.'· A.s res'enrcilers !li,,,ed ~Irlno:~ ,~~IlQmcs.the p'<l,lOldio:x £J-e,.\~dl.oep;er. ,s'llme
;.1$
:rmikhlg ptl~iM~
]!ii!y$-~(,tl4;l ~.
p:ro tn~oilns
iL:Ompiiml,sons.. 'Gri:£,OIIy nl<lillilt(lum; <I dail.'thtl_sil;: nr.l~1i,rna~ :g~u)me S:~l~S,at ihe Unive'r- ~ :.ity oFGuclptli (gel":l.oo)Ji!£~uze. ~ Ci(lil~l). Kew 'Garde:lm.s bio~,Q- "" l!i "" git\,Ll>m~nfl~~01]~ thr p](llfntS ~
9 MARCH 2007
VOL 315
:5CII~.Nelc. W'liVw,sdenooml1l.g.ofgl
WSFOCIJS
a ~~d ,(lI.gM
(\liI' W' \~'.
II
hmnepil!l~ ,00:Ll1r;1iI) ... mel b1oIogis;~,~n the iL'>'Ionian University of Life SCit11CCS nrn. ;;1. d1maDa:'!C [Qr tung~.(ww\'\'.zhi,ee/:rmll,gul-g,enome~iz:!.:!!~. TQgt.~~.lc'r. ~e dahl1:mseSCQni!ai~ Il'I iorl1latuon o.n, ~ mOIre thanl O.Ol]fD S',IJec ies, One On~l(: :rir$'L 'l~orr~:~1;]t~on!>selentlses
~.h.e ize ofcells, s Il-cmllped ~IP iln a study
Or!
bp
Emu
Cr.ow
'0' my a
rliiiremtot1.f
3,2, biU(oo lip
7Jm~jMIj:'al.l~
L9b1LliDm
!i!:i~i!OJj¥dr!i'S
1.5:8;IHIJ[~ I~p,
• _
~ ~]re foWld in rel~~-d ~)rUiiC~!))[Ul!:> thmiiY'-c:e]:;eI~~rhell'l ,un the body~ The sc.tea~dSls, sli1ggest~lma~ ,~ td-1!!!11 T. n~gilf;f1lf.~ ]'[L1!Ov\,.,J huo its curreru IX()Q Iqgil..<llilm iche, iis gelliome f:xplil.1.ded :l<lJ:pid~,y.Jlhe '~ prctozoanitscl f b{:1C ~I!;HC h i~g;c!"a~aresult. ~ ~'I'hi,ch mtll!de i.l more e~;rcct~ve n chasi ng <md ~
Oi!
11I1.~Y ,oOlldllldE! !bOIl tgd~V'~ b~rd~ge~ Ih~k sma'll genom es hom their cHru~s.tl.lilr~IIIIG:e5to rs, .
~.l.appot.::ldili:> IlW~SI,l~m~'I1!ts O!llO a~, IilVG!kltion~ h
<!Ir)'ree.llulcG:l'reb!ltioilll !,!!,1;1LS good enouyh thar l th~y oOluid lISi~ iht: !'l,i~c rlfgl s~d.Q's' C1, t~'CH:yt~:l> s :~ (':n~l,ill1ng, it'> baeL'l'!i.(1IP~'YCh!l1~~lg cell size mEliy bea;lel~t ether kil.llcis ttl O1A.:c;m<neiy predi!ct its genrnci:l:ue size, Tihe sciillg eatists t'~].elm.<l,ddcdto the tree branches fer ,~ of species in ol~.cir\:I,1ays.ill some £)I'oups of 3 I species Qfd ill,'lICiil'i,H!iIf:S limd used the size ef <lIllinm~:>~$p~cie~',\'~.th high m~H]b?htC f<lleS ':! I.cnd to have small gel.lO~lCS., forexample, t h:~~ro:st.eocylC's to est imf~I:Cl:l:l.c :5uze 0 r tl~.e ir they .~ whereas :i!J1ecie:s with ::l~O\~'metabol ~:iiom::;~<lv1: genome!:>. from that iufonnaricn, ~ big ones, One possible explanarion is ~hal inferred how d~e: size of dinosaargenomes I~~:umillg.~mnlml~~ iv~ rise [0 &tm:flU blood eil'1 ls, g ll~;;td'~"'ll~\ll.;;dove r ti 1m:1;;_ ·~which have ;lhugh :Sl'Lr~ll~to.:\(ollJme rario .u~d Theiranalysis suggesrs t11;n the ceannon 1.1, ,~ l.:a1i!l~an"'r0rt oxyg00:fn. r,lst~r aero".....[h~~.!'IfI~~m· O1tIJ.~~lnrof d~.n~Ji\;lli1!r:;,. ~m~LI] rOI!JlJ-ftlO1\;orj rep.~ bmua.es. [if a VJ.1l1irm-Ib.~oodedaniD'l~al~~.eedSI.olll;~e t~~le .i:i;'I.'e-d .00llll 230 ~lu~,lion ye,Hlrs ago. had l~~;mt 3 ,~ ~i~m of O){y~c:lm to the lilS mcn:llIOOIil;~~1 slrUI~1 •.~ (;l rclatiilvt:'~)lIh.ig, yenorm~~about th~ Sl:11~£ &l!z(:&<; .l!!' genome II,nu.!;lhlga.ve ~l ,m ,evolmi:OImryedge. OJ n!IU i.~'IJ:or·s. That t'OmM.on ancestor ,~,I\~e~~e to ~1iiCr-id nuti1;)!1" '~iIJlC'h..;:o of Ji.t~t~WIn>_ 011(: of b those br.mcTmes.the Or:llll ith i&:h:iian!S.i~achLded:big ~ The: 'Ms·s~l[ Im~ii'd 'Ill Ctm~,;iSi~'~~wit h ~hi~, hyp(ll~M;;l~. h i['(ls h~l\~ ~wrilb.nfi;;1S $m;b ;;t." st,,;:gosmqifl> {.~d TJ1t;CJP/oll<t. d n,'lI!!ch8mal~e.r g~~lG!n:~.I~~lnl~]{}~ oftheir ,r~f'· Their g~~{J(~~H;::S id net ch,rlilg~ much, ··Tl~.et;;~
d lnosaurs; inc! ~l.di!l!1l.fc~uh~'r$.ra ridl !i!JI;W,lth •. and neMiulll behavior, "Yi;UI duu'r decide you N!Jli n~ to fly aJ1d 'be warm-bleeded ~iktl
3
b:~I'{iI, Ol,ndllhen 1n1lk.-e:<III Ilrn.ese~~~a1lge:s:'
:!!a:i"S
Orga.nt. "T.h.q·re all 5111<1:111 CLu'il,u~athre i:hillg;; lh<J~ go \\~W, ""',J)' buck, ~md~1meycmn~k>gether ~o pood.ucl;,'; 1.~lis end l!arm," Ah,~,ough G regolry and others pind::;;e O:rg:l.ll·s paper, some scientists HI1€: not as j,mpmfl-%"!(.t ··h~;;t cm,~p;:~'e:r"bml"rl.~' mu ~r~.,
b1y confi.dentin the
OHlQOIliilC:' S<I.YS Mic'irlllld
II-'
"' ....
tyq;gd:a oflndi,JIf!I;J! Uniwr:o;i~y. BI~l~lming~()!l . who questions whether natura] selection is re~~m~lsiblc druvi.ng g('non~.C'se dUTcru.lu filr e
sizesto fill1l.eH:une m,etaboliisllIi!. "There'S <I CO(fr(::I~Uo~ [)n:h~tWfi. hut I dOl1'! know ohuny d:irect demOIlS'l'f'<'II:ian, of' C:l'USi:\~ i'ty."
ta U~.liIi:rll relatives. Btlit ,ut birdscvelved s,I1n<"llle'r guys hWllc a lypic<lii r.::pd,i.illJlJ"Slz~J ~:I1lO:I1~e." ~ .~ gem:uru:!)lur' their hi~h metabo.lli:iln1.~~.e '1m:;$- S~-l)i!iO~. ~Iion Ila.tum'lm.yari:sc:s, ~vhenl did l~:1'I!t ,Un,r~nkagc s But illltoIJ~.CiI.·brn[lch of d hln5!mr.s,-b~pcda I ' precMo~ km.. ~,s~1merl()p()ds-e-io~"ed sig~ w.~~, ~_ ~Hke pbl!1.1e'! o'rg:l'm re~d ized (h'll di mI,Q$,1IU[ I'm;· .~ .s i.~::;~:niighlhok:l the ~11l,';\\'e.r. Il]if~.c.mlly jiln:! IleO" yen.anlile'S. Thelupoclls \!,l'O~JId
G reg"\J(ry c~J.mJl,,~-d.~s. ~""\;:!.~~fm~!(:t~b[jlis!\!~ th(ll" l can aC~QUIllt: ~iJ:rI,he M~l<lll genmne::;. of .!lI~imaI~ such as dhlOSilI.lkI'S and birds, i~ \mlm"l exp.l<!Iin ..U ~hep;;;~U-eHiS il>ci ~m~~l;:.', fi fld ,P~'ln~ fbr s,. cxampk., i~Jlvre n ::;;.~11llar co rrelal.i 011 be{\\:ccn i c ~i.e <lind ye~nOl]M£: s.ize •.. ,e~"'<lJl1Ip]e. b!JI t l
en
~or
tltley
Som~ d~!Il!)$ ilU r fossi is .a!\l 'So \:\N;;U 11_ Ii'~ 1..1 Ui~~i):~i;;ly give rise [0 birds •. "T11i!' bIo\xl~our ,= .sef\1ed t~lill;~ !!l~11 ~,1."ve'the cmrit:~esliKm. once' ohhe !,!".uel" the :ide'~.lil::H S1111l1111genomcs they held ~h~~r bm'!~ c~lls (kliLu\:V!~IT~S o$~e(i(y~e·g). e()411\'{1'I!\t~wu~1:I nli,~I'Il." S<lys.O~J,~m. '~ But ~o OimiC ilm.,d eVf:1" establ i~~,cd~ I i~,llk.bet.v.reen Org~.rt gU~B that [lten~"Qds evolved to :2 ~nlJlm: slz~ ~UldOSitCGt,'}'tc !i1¥e. ··Tha~.\'WLS,OlJ!r have h~.gh,~r metaboHc fates tha~ athcl: anioll ::;elec~ .~fitsl ~~ep:'s<li)1SOir.g:mm" They ~"ll:uilned ooUil,es J ~rlios<ltn h.~,L as a,~:-'lIh. ~.la;~um~, ~, fmm 2() sp~fe:i of biNS" reptHes. m;;l!:nm;;L~s. d,nn n'lvorod sm~I,IGr ~;eI1C1mCf> Olnd ~mo;l~hlr c:dls. Other p<lleoll~ologi5l;s have ,allw ~:b~Ia:IJd andamphiibi.,llS, W:iilh co~I~.£!lI'e::i <It HOlP!tdJd ,~ ~tld d~~U~~v~rs~~yflf R~~ding .. U.K .. he cvid€,!lC:~ f'l;)f bird bjo~.[~g)'i!l hip~d~d
:I:j
i
g
bec.m;se ~enQim:le size cl1:anges the' 'Wi;'l)' LI~l'l:~rcen::;: (:'\pll!r~:;;m:'l~ili~~ ()rm,m~poniluids. "AIIlIY orte fe<lulI:e i5Jrl'[ .re.tlly gQ(ing~D '!.JO\'elr it" Gr~m:y ~"Iy..;,'''Yem. hllVJeto :Iookfrom the' bot~om tip mmd the top OO,!/\flliill every C<I5e:"
-(ARIL ZIMMER
si zes
me, 'wilttM?
IJUttli:tll;~d ned'wrinJ}
t. ~fi allci!he
~nii!1lg
·of
SC;II~Nelc. va L 31 S
9 MAIRCH ,2007
bnl~.111 lis
~'SJX'ind
QrllyilP
however, On:le.llie'\.\1 muJ contm\ler.l:i;l~ i~ypoll~e- ofplaoe cells fires. Sub~~.~enl :smudies fOlmd sis s;~;ggesI:S'~~<1l mem,o.uy and other cegnirive I~'M: sequences of plm~~ce~l:trilrin£ that ·rn;cuD' ben~nls;!:Ire m~rdy side efWccts or the 1[IIH~ ~",[I n'!,[explores ;) m;{w I~fi\'~m!ilme[n are face of your b[~liln. funcrion of ske:p: dialil~g down ,synapse:!; dUI! re:p'.lt'I,ycdILl.cnext li.nm,eIh.cn:;;lt dozes, ill> if lhe What all lL'ai~ !Hmr;diH;:~ivi ly accomh,O!vt;:g~lU~-.Il tl>V!;;r~dl€ld by d!J)ftim~ <u::'livi ty, mlima I~m'u;1;\'S~l~ S!:t;;rli." dmoJeWlg~k'\;'lp. Arid some skeptics aren't ·conv,ufI(;ed that 1luIn31IJlS Ull<liy do :s;om,el1'n~nl.g similar, ]n .. plishes, il'anythi:ll,g~ is 11 my:stery.i!lnd p.un of lh~ c\,~YII.I!ll"gcr' PI,]zzle of wllY we sl.c~~:I• One . lil!cC'l"Il~~~.anyLhing to do \.\:.~thMClli'lJfln' m al I. liOO.:!, a rcseareh team ]~d by Pl\crl~ M~qH(:lOf , i.de~i~h'll has S~li ed fa \'01" i~1 rece n'I years n GU"~I.l.111 the uaeertainries, researehers :my l.he U l'IIivers.ily .of l iilege. Be~g~~Lm. used p.nll:it.rtJ.!Jii.!;,1Jijlis$lr.ifiiJ l.iOll~.og,'mI.lhy .'PET) t.O iUO:!l1iiE.~hat dWll":b"geer~~1 i!liis~~,g~~~ll' sl~~~~ lh~ .. th:~ qU(;:>l 1.0 unde:r::muliJ the si!i;.le~~h~~ is br~in just beginning. But tll.ey'reall'ead;y findin.g ot:'<L in lrep~.'IYS,~~", iences fliol:m, rhe day toO per i'tor bra in activ uty in :n.]CII.1 pl <ly ins 11 vi rtualliilr~~gt:I~~I} !lll;m~llryof what h~.ppl,;:~ll,;l(l !b~d~'lti!l g d~t~~ ~lj;put \';h~1!1 ~.Pl?¢~~~ hen tbL;: .h w re.~lily g~l1mle i!lii \';~h~h~lm~yl(r"'r~~d tu n~v~Suppou<rl: for this notion con1:!>:;) fm:nifl ~~\~Lri}l:lly we're olT]n Ih~ Land QfNod. g;.lle throu£Jh .~ \.~I1,'m~1town IhM:t~~,d~.y :;CC'UlL~ a ~:'rom,he shoot ~'em-up v~(.l:e'(lgame Du ke t o f'e )l:P~'1:imemswi lilm rodents <melpe c:rple. Nuke m). T;~,e:same n;>,g~{1 >0 r ~ hi ppcus 11m!!;!: inclmlgflg~. :i1e\/t.~:l'i!.ldyii1d~is issue Sl!gg:es:~~ L~t !lI1Iil! iSI~!l't!l!llt agaillil i ng that bO()isti:~.g such meraory-related T11,C :furnl c1oipocrimellltaJ evidence: I.~~at the 'I}I"!I in c<llmllH~s dun revved up \\.II~.cn the )lUbjClC~5; activiryin file ,~I.eei),~ng rain C<!J! imprnve b exploredthe irulO.lI elwirorulbe'~:~.d!i.obecame n:P~ilYS,recent eJ"];te\I,ieIl:CfSd!Jri~ sleep ceme me:1I],~ory performance illmhumans, fil'mn expell.; mcm:s 'with rats begun :i,n tile :~ 9'90s ;ICI,'U\'C\"I,hellthe men :lSI ~ilpped lillll:O s~m\r=wmre SO'l'n.~,(;; resssrehers r:>Uf;pG'>Cl ilh,fll r~p~"lyin:g, by nel!wSC'i~~nisl Brw::ll McNiatlghl'<Ul of lh~ sleepIh~~ igM:.n ~ u ssll,~pl'(~g(lis eflen con~s the recent past during sleep is more' than 3. LJ~i \:er:s.ity of Ari7JJml. Tm.::ml~,and 11 :gr"<Lr~l~ si dered the deepest: Skl1;,v~w.lli\fe sleepers aee m~mij;lr)l 'Lid, This rev~¢\"t m,~yO!b;ogive I:h~ a~e smdem Mm!h~wWils.t)~. McNmighu,'l!'i hi;~rd~(I :rtll!~~. D~urh~~ ::-:~etlp t,f~il;.'!.~l~cffi1this S brai.11I1 a chaaee 10 cau::JI1I, i.:lm~por[al'llt ~:!iifarl1i:1l'1~· and W.ullson recordedthe ·e~el.:trica~: <lc~.ivil)' of eueep ba:~ogr"<I,m {E EG)I raees S~~Q~l!''wmrcs;; .0 f ~ e:leoC[r,[c;:.nJ;H::t1vity Ihl'iHU1il;~]Ollt the braiu that ~ tkm h~l~~S""C'd firstt ime a roand, "l'll1cr,~:1;. ~~~Lrons,caUed "'pI:,1.~ Ii,lells"'illthe .r,,'I.1lilllm~ i:lu: more an:dm,ore ev~.dellic:e a'Ccn~ililg Ihmwhm: campus, These neurons, have an ;)Ili n~,tyfor peakabout once 3 se,cond.rET SC<H'IIS .~'n.ippoca~up{ll ~. w~'l:\e seeing d.uri!m.,!!, ~.~~~~p ~.Oi _~ is US~ a particular lccatlons, 100 that as ."1 riM runs revC'a~ed ll1![U the more ~fil~Il[se st lren,gtllc n ing 0 f memet ies, '" :S<'l:ys Rober ~ <lIiOllnJ,ils, em::]m,m-e, ,J given p·~3.oe cell fires <Icli.\'ilya volui1H:e:1f had d1urli.n:g slow-wave I!! Sti.ck_:gQ:~d.a ne lil_~sd(lnt ifi,l .:'It Harvard M~d:· €'~!cl~ lhWl~the mde~l p<1SS.~'l dll:,o~~~hthfl~ c~U':s \'Iloop. l h~ better ~H~pei!' l"onll~d Ihf;): ~H~" diu)' ~ I
,I
9 MARCH 2007
VOL 315
:5CII~.Nelc. W'liVw.sdenooml1l.g.orgl
NEWSFOCIJS
;;vh~n he sprhu~d lh!'O'"i;~gl~. "~nual t'l)\\,m ~!) the f~nd certain objects as qlUic:k~y as possible, M~quel and colleagues reported in the 2~"'! lll(ilbeit ']n04 issw::ofM::,unm. O M~lqlUC't's, l:llil.d~lngssl~O\:\~e:dnioe corre:la~ u liml bt'ftv.~e~ Illf\If<.!~. Lct~viy (~ur.~n~~ieeJ;)and ~ su b&~qll~IU. lnC'mory pcrl."o:nna nee, says J:;liU B[wn, .. ~1 n~urosc;:h::n[j~q .at the U!'!iv(;;~ity of
Ui,bec:llin
Ge]".I]1I,my. The namral next step,
II
hypn~.h~sis.
the hippoc<unpus al1d~a~er-pe:~~aps, bllel ma tter e f hom s or d~y s=-rransferred lor long-term M(lrdgc tn ~h~l:\f;rebr<l~.(<lrle.~. r e ~CQ{}QrIC:CSevera] lines of evidence su.pport this seerraric, <lIllC!lIfnS '~iliae,IH~he ebservation t hfil m.~myp (:03~lc \\'110 sul]":(;,' a!li1ln~.~ilafi:l'lowl i!l\g: d<imag~w!he hiprp(lc~!m!H~S c~li!'1 l'>.'rill IL"'eC'31.1 events andf,u::IS learued prior liOtheir
likely enabled
!IS
have a nJl!>.hi of .i.ns:.~gbl that solve ~he PHlb lem more q ~Lickl.y liner .3. ~ igllt of .s~.l;'eP '~lliI2ln (Iller ;:)iI sanli;h.u tlme oh\·'lke'11JI:m:~s:s"
NJ
l~,Wifl1l!lei
ll'!~t
HJljt~for BiQI(lgk~11
~(lnll~t1~.Qm!· \'I.11il~
Bot~l say~..w~]st~l seewherher <lflifi·d.,Uy boostung memmy-:rellatcd neural ..aclh"ity dur:. ing *,Iee.'pc(l'u~din(lpTtWe ~lllemo~;y pi::r:funn~ <11100.01'1 pOIge 1426. BO{r,~lS team describes an $th~lmlptIf) do _f uSllhaL nerese;u.iClhJers had! ~'olul'I~eers pby :I video vcrskmtlnh~Cflrd yi!1:!1;~ M~mory"also k!l(n,i\rl~ ~s Cmu:{'a:ulliatli,mll)~ in whech they had to leam and remcnll]f!'r the loesrl ens of Il3 i:rs or (,9:rrlS. Dr.mril1~llle :iO.1Jne i~]l"~ illl." srl.)ujJ 000 ctI[d~·. EaC'~]m~l'Ched I'~ir nu1hed ililii th~ sereen fo:l" a few seconds. one lit <lU.ii,m],1.e, with .1.U the o~ll~r c,l:udSk'I-C~f1lg dm\,Ilii .. AliCl' ~h~¥oll,llllii.t~t'N
lfui.eysleep, .1~.lli.s roup had volumeers play a g card !:,;<lm1;li~ll\m~,(;h .utool,npt~d to pr~dilt;.;! ~he'y whe ther it wm~1d "ra iI1.1.'" based CUll cards the researchers hlild S:hown~'I~.c~itl, ~me was The r.ugged so tl:l:u. Ute eard lJ.f.ilh dll.e diamonds, fur h~-nn~e~, w<vsWoUO\\.·~d SU1!1ny \1\.'11.~~~~(:r s lily 8IOU.4~f the time .. E3!l:b c3Jl:1 hold us 0\'1'1'1rule. o
~nt h~p;_dr~,.the
researchers tested the subjects' recall by nl~l1I~ illlg Cil1ne:Or~1m.t:30(,,,005 roo~upul\ld<'!~kin.g~ihem W fiLlil.d hs mntch.l1lile researchers then used
and Ji found ~haLlnw.::h like hippocampal ,t'CR s, 11!.:::lJrO]lS the \1~~,~lal ~ll CQI.'l¢l~..t~f~l~ lwepl!ay f:ilr~.:Il1,gsequences l.luring slew-wave &(;% ~('If..~~bj~c,;:n;. 'd r~~~il'!i'G-d~Qod~l!n\:hi I~ :;l~pdm!!H,II~h \VhD !~~~:rJ.;tiv~lY dm:1ng lht.:r:r!~s' " d ay~ilrn'i;l ~1.1,111.\l !1! nrli U],g. J\ilm~;n.r1M. the seienslell:p'irl.g. Sllbjifi.'i~I:s who gut rhe rose odor e:ither while awake 01' whUe il:D IREMsleep, QUI~hi::' dsa:s fbundlfui<lil the replay in the visual ClJ.I1iCX o~~tlr~tl!I.llJ. !mw..::d :Jro,o meIllor)' bOOM!nllJrdid s Ih.,ppemi ~r1 lacktli:t\P \\'ilh repl~y in the ~]ippgp[1!;!Senli~l.g the od.or Ilhlring .slm""~lw~wc sleep C'alllpUS. ··IIt',; tile I1l"st lh~1IC\'!I'e' soc s.eqmmccs .fl3 hdp :subjelo1s '!.vho h;J,drn'~oc"'en ,ej"'IJ'OSl~.~orose bot.lliL .ill die hiIPlj)Oc.~mpm;. .md lhe WI,eooCo:n,e;", .~ dur.iing the Imi l1Iill1J:!, ;es.'i~OI1. "~Ir:~. fiffi~ "'~!ill.dfY ~mdl~he if ~1C!o!JTdilm;I.Knilli nl lime:" s.~y;sM<lqLLC L s the
place ~~U !'Cf!lly de:t:lliliHl=:'I!1!ltc th~~Um!;t;: C<l~l ,~nfht~nc!,)' b mi;;:I1I],orywUltl1.1li'U:lmU lh,ll eXI~li.ci~ll)l m.:tuv:lIH::' s
sleep, dlf' ree<irchen>.~.;]\~e ~omt' afthelll:ot p~llr Ofr{lt':e~~e~m~dak Th~y'd:pre\'itlu:s:~y ,~iv(,11 some of the ~mbjec~siii whi!fotl'lose d:urin~ ~he:urin.alimllmining SC!>SIOIl \viuh i:~1f!'cards, ru.l:5onirli,g th.ll lh~odm \l'iQ~~1dem;~\."ltelll~:em" r ories of the uilLII1.ing S(I':!lSJtOll iun. th~~e :5l,11bjects \1\!i~~iULH w"tiLkil.g: ~l.eml" ]udeed, fl~nc~iomd Ill'~~ net ie resona rice iIll a gi ng (I}'/IIRJ) SC~.~.l in, s de-(';p~l'Igs~llbjecl:l: revealed that the ,od~)raetiWlted! ([le Ilippocmn~.pusiln dlOSi: who had e~~ rienced i~ pr~~liousI)'. even l~lUl!g~ the E I~'G .sll(}Wtld 1110diS!mplio:I1~" in lheslIbjecu;':slmmll:lt'!r. Ah1rn.OlJgJ~ tht;:y did~l'treniu:;·m'be. ll'!I[lclling reses i.lll.the.~r!:i,l!~et),. ,~ubjell"ts ,,,,1u) g~~the flr<l.~"'U'I~ the ~r.!l'im~~ rem~mb~r,ed, th~ Imultdu.;(i 1~9ir~ b~ut!'r ~lil.eau:)!!.1 lfi11)'. genung, 9'1% cortect ,t:Olllllp'1.red~D
A MmJy by Wi Isun [~~.dpo.s~d(J(;H1r.;f~ r~lkllii;.t Da.oy UI1I, J:i,publiis.hied il~ the Jallu;1 ry issue of Nmun: Neul'u,,,t'i~>fu.'I:slII!wons I.hi.sitk,:!t WUiSOll
lM rhe vol Wl~~ef!' d~d~nT kIDL(:I\,V1'", rules '[;Ve!'1 [ existed. As expt'cled they did IlQ bener thsn ·c~~.m:c!u.first. B~11~flt.'ir 200 prodi.t't.ions. their ::;ucoess :ratesil1ilpmlVe:d. When the subjecl:!i
came b.."iCk12 hO~1.rs~a~r til:j! .~~~I1+ t~~~y~~d t() ii'mllp:roved even mo:r>e-tnd. thase \!;'ho(l s.lepl
i~Wg"!i:)V~dabout "there
'Si~g:ro\\I
\l;~~fl
I.e.,..niin,g
Then':: s. atf)lOiBlCreao;;lug ~vid\.':lltie ~h~ltdl.ffer= ~l'It s,t,Jges {If :s.~eep~l!leil1~,'Ol,'Cd :il:l 00.1]sol idal~ "
:i:ng differel1l1 kumls ofnu::luory: says .[V[3U:!i1.eW
during !)~eep.;'
.scut>lnIIHI!e
m,,~
<J,
pn)ces:s lleurOM:i.ell&ILS;~5'c,111
U)
d~i~
M.o~'emanm@m:1ll11J Be:Y{H~d :;:i iI'n~p Iyrfl:r:~ ~ i11 ~ m~~nur ies. dH,~ f)! br<lin ~mlay!be ~nflin~ t~uou~h 'recent e:o:pel:.iem;cs. d:uri.ng :sleep to i.f,l!t,:~]t:ifyi!Julcsabout ~a~];_"'ii: <L~1H.1I elfec~ (lr o~heilrIIJse11111p;:lHe;m~. S.OIlfl,e res~areh~]'l> S~ISJ'~C[.o.~,e ofl:~e flrnt h[ril::;,of t hi s p hellO m'CIlJOI1I !Ct'Ln1'C from ~1 200"'1 N~mre j:)>1perby B.om 's grOtlr," They r~·p'0~TGdll];ilJ~i"~SC;II~Nelc. a L 31S v
\~h!lkcr. f! nl;;uro$!l::iQ:misll.'lill~flnl:r.rd MiI,;d~c.id School. Sp<ni'111Il'1£mmies.. lilke lh~ formed b~~ pt,~yinsMii;nl()1)' orb)' mIVi~l~rI,g.:Lhmulilh~ m~c.e 011" \',~rmal tIl~~'n,seem ~o be:c0Q1Si.)1ud.1ued during S.~OW-\l!J"d.V~ ~Ie.t;:p!' TIl.{: ::m~1C apll~ru.';.; to be tme !for dedamtive Ilii,~lemQries" which
~IWO~\1'e re'll~!l'll~rill~ li~~I~"""":bl'LlI10lmJ~s.\'\,fIil'li,~yothe:L'klinds, ofllm::'n~ol:-Y, Some ::itudiesw;tLve
o;Ii
9 MAIRCH ,2007
1361
.e -,rl..ne'le'
anage ..
me
bib'i~raphi{ software that S'tream ~nes research. wrim ng and pub~~hing. Reference' MallaQef has served OOfjDate. 'govemmenl¥1d ocaiemc resetrchef) ~o~dfflde for O\~f 20 ye,ars. And rtO:# l!el'5iool1 deliveG ~ew way:, to~re flooview~oor re:ferel'l(jero!locti:Jns: rust ~ ~ tolhe We-b..CoIlabffill'IE ~l.ilh((Il~glJes O'!i'ef a ool'lMrl:" Li1 tofull te-xt ~ fires, These are JllstSOIl'OO or the po'i';'elful fe.at1Jrei ~t. aI!lrait }U.i , R!eferenteManager i~1f1)Uf oommand and cootl'iOlcenter for all tIi1ngs re:~erenee relat.oo.
•Au:..~ss nll."N and upJal@dcon:t8fll files a[ WilIM .R!'fman.m •Share travelil1g libraries,¥ntn colleag,ues • Cormecr to daten ·s.ualizaOOnlools. Put inrlO'JatioFl into actioo. Ordei or upgFade' lO~.
Avaifab~' 'for\lVf ru:nljl6 in a singte-user and ni:!lWOrn edition. Phom~:801l n.2~ 1227 • 100-4\38--5502:6 • info@lsJr'@'SeEl[{hsoft(OJIl
'THOMSON
....
C! C/Jpf1fll'Xo!l_,,*~I!.o."iiI"
,I;I~'
I'~
L"t_,.,~Wt_
dl~":~IIT'1riO'
NEWSFOCIJS
$~TO!1g.('l~l,ll1.~omll 'Cl;1iPl~p(lrtl~f1t LA'!i!l~,r"rid-~y0d
II
moveurem
d~.ff~c~lhseq aence ()~!1~! ke'ybo~nl. during s'ta,g.t: .2 and REiVl :sleep {sec dimgn':lIm, pcl 36~, l W hry I:his dii,,;·iS ion ,on<Joor exii,~~ us <Ipulzle. b~ll '\~,r~I~kC'r ethers !>POCCUI:;ll(~ t~,'K: i ncr~ ,Ut~~ lh~lt d I;l!uphy$i{llogicll~ Md Ill.;;U!l(l,eh~mieO!Ii'1mJj~ it:'l~~ assnciatedwith di:Wl"cllrent slee'po stagesmay
~~l\inr ,celt.lh'1kili1m. gfn®._U41~
also bi,gge:r.[aki.ng upprecious space.•t'u'I,d f:imdly: II)O much [TP rmljl S<iltlLl~.'a~e :SYmlilipseS. 1e:<lJviffilghem uffila!Jle tD gel :lilly s(['QI1!_!;er hcu t w the bt,Jj Ii! needsto learn ,~(nm~lm!ill:1gnew. S:tee·p reSlo:res hmm~cOSl;l!Isi$ by ml(;hcliblg down symlplu:(; !-l1il'l:.'11~lhs. Tommi.<Irrgll:es. ll'~ ,01
far ~nt)1"C'In!ITlpmWnl service
~l(1dC;i~l SOl}'!!.•
boostin
in~m(jor}lp~~lofm8lne;J.,',. he
l~li!1Jn pnwi.dimi.g ~
!>.yIH\~"~e~. Pft1pOi'lellh" as
uf lh~
~l~ri'H'i:i)'-
pIOl$tidW,
Some rescaochcrs poim out. hmven:r,ljl!~u: Ihq: Hu;:n~~l!r-eQI1J\,,'hit~h ~h::~]J ~1ug;es lebl~~ W
COIl1,~
~,5% impttwenle~~t '!i",!: seein some thii:1g...;:' Toneni ~. "I.dl] iuk :il does :!lometlljl.lg much more :~'ij,nlll~~m!l;nt.d tdittl neusom lf~du.: price 1:0 \Ve IJay for IPbSl]city:" l10'!<\' mig~t sleepreser
Iished In d~\;)15J~m~Hl[Y ./owwrrl
PIJJ'~:io.log)'. They reponed li13t
fe:phl)f scenaeie fuI<1iv,egenemliy assumed, ".~ t~i!lk thi.'W QQ~ld not j~!u.tindl:lp.eIH,f~I1!~J}" .m;:~
n~um!>ci~!~li!!>l the U!1l~\I'CffiilY at ufOdifl:l[,Uli'l. los. Angeles.. S~lickgo!l(j,and other pmpommLs of." s!eca~-.nrlem(}.ry~£nk~c:.b.owh:!"'l:gc t~l"llihC'y f::ut'e l:1IllIIII)' miru!l'!tll v~ i~\lI!s ab()l1~[he roll! of sleep i;n ~n,el1lory t~onSiol:idalion. "T!lc're arc rIiliJ~i\~e qlles.~mi s tel'mJinill£ aool~t ow e%lie'mI:h sive it is, Im,v imp(jr~llll. i~is CXflct~y whi.cJ~
.1!i
~radict iml&,,'"lr ~ere is illflcon,~j5>te,UD,CY here, and .smueone has te be v.'~con!ll:· says, J erome Sie'J)';:~.
Tononi's proposal. S<lY~SI.. ckgQ]d: "lr's an ~ synaptic stre:Uil.~IIl!~'! One due. 'imHui ·~ay~,. el,egalili~. ~lypolhesi;s that doesn't hfl'vc 1lI lo[ or c1Qmes~":om.1 Stl1:tdJy by Swiss [e::;.elll,d,~e'l:s· ubp data b~.mti~ldit,"
~f
.~~"......,.......a..~.
eause enee-per-seeend
AIl1(lili.;;r \,dd~-O~!Il i~.W1J1.;lis ',\lhe~hI;:r tht;re's, a ~inkbetv.;een dlremnh'lg :'lJru! memery-related !iJ;:uF'.lI",oovjty dm!iJ.lg sleep, The kind ofdirec[ replot}' of recent experience ~mgges,led by \V~is.)11's. 'wuri<:. fUiI CX!U'1~ph::•. dassn '1 K~~. to be the stu ff of dreams, s~ ickgol d's grmlpllnas. fOlU~d th~l (lll ~y] % to, 2% (If t;:ph.ndI;;S from dIL-.e;nll.~;:; reflect events !'110mI~~eJcnevicms d~,)I. U' dre'ams don't directlyreffect the tne'rno\liy~
"Vllc'r~ ~n~.O!nj)lU~. ~'Dd;:' i1I.ys 'iVaU{er. S
ind uces <I lype of );}I'f1(1tl'Jci: ~~·\!!nt2IJk~~1Ji.mg e a ~.I lo flg -te nl~ "d, depll':~ton ,{LTD}, Thnol1Ji ~hiunlQ; if\! [!:aO ceiacidence
tliHU ihe coo rd .i [I,.nedi neural fir~ i1ll.g durin!;! :f;~OW~ ·~..we sleep iimppens, .tt
iil!';():l'as;btl;lf!-'
the sleep-
'0 iesjn
Like
S~egd [I~~OqooMu.om;;
;J~y cnnm:iClinn .n 'll.!'" b~~'W,!;':~~l~!~~p ,mdm~mory •. Siegell, SCkys, 1.I.c favo:U'£ the idea tjJal sh.'!ep Cl\.'olved to hell~ iI!~liml;li~.~, COIll'le:r'¥'e! ('1Ilcrgy for t1lileiremire b{ldies<lnd to pilir;wem ·tln:nn frolil.~, b~ililg !l1:.'t~,\I~!lllim'£Swl~~ th~i'1"l:::t:e$sI,ikdy 10
rind food and ~mmre ~i It;:eiy~o !be eaten. To b~ ~t~re. ~~l.flr}/i~~ i:ia~ u~ly f1;J1mJ.om n(l~
[~,O!'l of
:sl~p,
~,;(m~lt!:'Ub S(i:c~golcl.b~1~h~e'\..~-
defQ{,A: lh<lt it: i:s one fLulcliorn, of s~eep,iXl: J~CflS.~~11 Itn'ln1:nilills. ~s, wo gre<L1 to l~mre. Once sleep ~vol'!Jt."d. 1m s",'l!y.l, e\lollJtkm ngurod (1Ut i~m~' to
I
f
or
1363
FORUM
I PERSPECTIVES
LIETT'EIRS
a t:{lmbin.~tion 0 f rhe TWO m;.i n M<Jy.:!glyp~ hloeks): U~lJ;yda IlfM ran.. do~d}l"blJnc;;h·' glyphs as a/1m the C!l:st:*~lbl.ock ['IJ.r, (I )]. 4) M'lI'l!)' of the so-ca ned ;1)/'11 s repl~ ate deeorar ive mot ifs h ie IIN iiHEIR RIESIEAR(H AR'l'KU ~EA~lIIE5,[1" WRITING liN AMElRIC/f 1(15 fmma: Qilrl iI wide .range: oflm:gely ll,I'l~J'nMeIUinced (i!.c.. I~]eir Iom~helmiSept, .2(W6 .. p, I.() I 0). M.I. del Carmen Roclri.guez M..rl~:lIu:tzet til Ii e i~yi!ii.nOI.pr.pv~.l!ii. :l!ii~H· it Ix IU"Ovc:d I sl~'lf~~I-se.al~ !~:nif3(!:l'S. NOIl1.!' eal!ii sugget-::t:tha[!~.c im,er.bed '~CIli$ca:i~1~leek" ls I.he filt"llt.di:!lcoverjl.of b lo:hhes~ Illotifs ill tbe.iir origirm! cO[l~eil(,l:has been ideillined as 01.farm O~If'nec ''''!'iliIiS, J\.lthulIgtJ. \"'e 'lgtee w1.l.11 authers dm1 ~I pas:-;~~ nfwd tiog •. t.lH:ii:·;;]'!'!'1jP~G. "ghtph" If, 1 241.38l5.2:~s. f~]i,lfl d ~!:> p,l!"~u hh,~ the is h~(l rllJ,;9it til..;: 01mIX d~d wm'i~[~. ll::tv-t,,: trong f(:SCfVBltjOI:!S coacernwe s headdress <l:ssemIb13~'eon a Imnm,belt of the ee Its (nos. 116. :~117.I I g. 11'Q) reproduced in OJmec .rJr{ ~Ji':1 ,u;:.h~,u~J(,·."(;If.:n (2' J'; ·'glyph'· .. ir~g lh hi, flrl g f,l,ct ~~!1dittiif!d~1;'ld motHs, ~l' :Belli!£: fm.uiI.d by persons un,known in <I pile ofbu'~ldo z;~r debili.s fr 28/58 is found ()[I1 another c~h (un, Il4) .. and '''glyph''", 4 is iIllsc:1iibed 'O:~Hl s;[ol1erigll rc 1110. 47)1 agai un,n the same cam log, i does not censtiune rel~f.lbleru'ov~ml:~ce. :n The block does not fh ~tnyhlOw~;lc:alcgory of MeooRI1iTI.erl.cm~ 5)WhO!!!t'l.vecan [lulli}' describeas l~e "ceetie" glyph ~#iI2Jf501 ] n1jl~rihed tin ifact i.l ~s !l(U .! stela, celt, sc W1lpw[\l! .. OJjewel The he.!u·, fits illlClknown cmcgory of M(lSo~:mclfic~1l g:lJl'Ph~nill ~ogclbc.r\C\li~h .~and Olmee did 110~: build ii.m1. s~oU'l.e:.there¥.bn~. It CiilIlilOl be .1!1l ;nc~u~ Ibe centext onhe d Dficovery. SlITN1g~.y sngg,ests :J pract iC;11 joke ..
t\,H~t~II'~1 in~>cripl~.Ol~,. ladeed, thereare IU.flil}~~llindreds ofsilYlil,~Jr S4;:F-
peutine blocks. knuwn m l,IVenl;<lliilihll: were usedas basal ()!.nm<Lme.U1I;~ o.n earthen pl!)!l~ibrmsand !Iii! the bt.uril,;ld ·raV~imI~L;;!ln,. utnot .~ ll;i~,g~iIt. LS:.an h Fra!l~hco Stat~ I.ilI'1Mmily. San Francism .•CA94:lIJ2. USA. E.·rna,in:~blillihn~@~r,~u.edLi. ,rMid!d~e]'~~ness.e(1 Staw Uni~~rsilY, Murrree~:bQm, TN!37 HZ. liSA E·:[q'I~1In kel~er@ t one oflilm.elleh~s e:lrngmving or l.'eliefc1Jrv~J1g, The <LlUi1101"l!' mll~il1Jgs 'rnlli!!l.oou abnetthe blcek being used [or practice ."lm].re~)ea~edly erased lI'el~nmO~5 (~::;lIltilng, in a concave :-mrfac e) are farfetched. ~, J, MaIM, oMi!'StltTllnMffill ml.lin!l' :S'~h!'m.l': Pt(}f!D.g,Q~~ ~ alld Hi~POf,J'~' fOrJl AAdtm J J Knowlm Me Stoflmerk: ~m, AffiM((l1'l Ci'l!il1mlia.l'u (PriIKtIOI'lI(jl~f~, FI~l, PlimCtIOO. ~I, 1"2). writing sjfl:lu::m!'i ,n;e: ''I,'rifil~m'i'hie C!ilSl~a]1!1hu:lk. b ,. 'E.. p, 8~n~l!i!I,E,9. ~'liI fue:rlibil. I OImtc An iQj hKiem'~i~ (N~lijooalIGa~te')' 1Ioit. IlIf ei~her v-erticaUy or linealrty r or Wi!l!hiD1g1OO..trllC. 199'lt
IR.eslJ!I GlIl1i115le
WE THA.NK IB,RUHN'S AND I<HIKJE.R Ihra rlieu h\'t~Ill£, s,e'vern~..issues th~t Ot hers: h~\I~ r,;\i sed con-c~rll~ ing the Cascajal b~.ock'J'm1dl~o:w provhJing till the uppprw~ity t~1 1f~),1
them 10 rest,
.111 The
p.~IJVCI1!i1I1Ce
'see·~~~tl O!I(.Mib OII<!~. rex rappeared on ~~. an s :Ill!d e. flrl b. on a seu l:plllIX:under Cli<!c:e:P[J.!1l~~ <IUy~lrn,o rolug1l. scn1l.:i ny (.I}. The lie rote. 'lj:!,-e \vould netbe ~u!fJ:ri~'t,lIIO~~ili.l"iIl ~I.l~t('ilh~rp~ ... viou:slly cl:isCQvC:'.rede.x,<lmpIes '';'C,lI1t unnoriced, J) The s,igll~:;:ormp~tl.1Xli.o;eflill,1 sequences: t I~ey do IUJi "rando ml)' 'bunch,' ., '~!'i n!llm~ B l'Ind Kelker assert, 1',~\te pauerus inllhc Last'3ja:l1
I:t"."'~·
f':i,l«':~ll10 3'~.a1i"C"<!, ofa sqmll.'c nITlctc!1J"S lhi ~, wl :1 knowa .[lrch<!e(llogic.~i. site in Vi;r;u:nlz., Mexi,cQ., :;Vi<lny other bona f~de examples of
~ew
btwt still
'$
..J
anelem \'\,Iraling
Il~\'1;:
even
~~SJ~
secure find
'~ M~'llt.l,,,m~ricii'l'lbnhmio:l!n, Scripi <I~d E~ypf~ Rloseu~ Stone .. Such objectS> will CQ:t1tir,ullI.C' to .1lI.. t apr c~r :we.~m.n I~ly. \1!le '!II Ih.-.vc dOlilc this lO the e1'itent of Cl'~~r bil ..ty ai :~ wit h file' O:USiC'lj<l1 Mock (mel M:rll.d by our ron~
c<lrehll sl1~dy ~i;n e'Vlu el:lce of
e'x.:n:Ln,ple of
:~n the:
4) The bleek coetains s~g.~n.s found on obj0Ct~wit1~p!:o~r~nanc~ nd otlle:r$tl~~HI~ck .. it. SUI1m,eof t~.lf'lkntf'r have been kll,l(lWIJII,since I~.;:~ 9:th '(';~rlUlr1'HI-!.;.- ID~ "Humbo Id[ '~h" I (1)" One p:rev,i:ons.ly lInknovl,ln sign (glyph 191 ~~llpi;;~un~d>~~O!rs.n~r~he d~ "I SCO\l'~ ry !Jf the blm;l iaa s-6clIre<lm::h.leologk-:'11 COIlleN.t <It C!I.utoOi O]rralitCl. Chu::lp~&'M,c,~i~o {j). AU
k.1~D'I\'l!iI
wllid
.1I&sI;1~$~
rehiue to p!iC-Cx~!il.illg i~onQ.!~: rophyor ('Od! ned tl)rmboli,sm: ne,v :s.i,glm:s<lppe(u wlhien W<l'lifamed by sc.rihill needs, Any hif:'roglyphic
hi er(lglyphi!i;
sYSi:I:I1IiIIS
iniheworM
1365
lEITERS
sys~eiln lh~l·d~vi:<l~~d jhHnkic'l;1i~ ion!l:ogl"olp.hy \",-'{Iu~de uct tl!'illy unique but, indeed, M i[i!~ b exp~,i.c?}n,k' phenemenon. :5 Ji [:~ the •.~l:1 sect gJ}'pif' w 'i~~. r<l.ct i,e'll p joke. the joke sterwas an O]n1:lcoC. T~c mot iI' is :l>hO\:vnlh!\;'~-dim~liIsi;()!'Iall>, in the dimim!1.J1tuve MO.nullne,n:t 43 .,~ S<lIl, Lorenao, diseevttr~d ~lnd~}:;c;lVii!i~d by oi1~ efus (CQI!~ ii:! Hl66 ~U1Jdpublished by Coe ,uld Diehl iin, ~9'fmt 4). We ~l:ject d'ae O;~Llth{Jf's .':!pedr.c criticisms and lhe:~1'h:n.pUdi. claim [h(u the: b.iock :~~iI
II]mderUllbrge~:y.liO'!'>reve'r. we <lJJPiIr-e>cu:ne the 3S4!ai7. USA" ~hparlJIWm ol.Al'!tn roptllo gr. Il roWl'i Unj;. V@lf$ity, llioYidemm" ril!1 02912, USA. ~D~arlm~n~ 01tin'hro1 ~log~". Uni;;efSIl!l' of Callfll)fnia Jllilillul de, Rt~er$~de. 0\ 9,2'S;21, IJM,
IIiI@f;~mm:;~
l~,
;1,,!i,,
:5" .I~JI~
I\;in.!!lie\
~.rl!:I~~
!jJ~;!}m:~j
B.lUJl'C)'O,. E. Ml!jia. Ms~ ~l'iIifili,~I~ri'oe CIlIli!lr.! 'I H. d G!l\rt~<I!il1 (iijo. G1!~!'T!;lI". 20061 ~. M. £)., (m, R:. A.lDftllil., tll1hHalM'~/r~()JffiN:; vm. OJ~~'!',cf r~~ Pfli:~' ,I!,m~irnl TKo l'jl',BO~.
~~I!lir!e1;
<Il1Lho:rs-eonclusien
l1ile'~,t lotl~.e ooast
all
ClUI
the 1IIlimIJOl:1t,am::e
facrthai such challcages arethe essence of scientific enquiry and that they evemnnlly
~eillid truth. \lVe' still affirm :110 lin:M. rhe Ca~<Ijt'L1
an l~1~ weknow
bleckis
l~,e t'lldeg! ~x~lmp.le of~lj,rr~li[l~inI,he Nc\~' Worldnud mnongli1e' most inlpo:rulllI. I'intls. ever made ]n MIt:M)~mer.~c<1.
MJ\. D,ElIL (AIRMEN, 1t;~:[)R:rGUEZ MJi\RfrNEZ} PONClANO oRril'CEI!!P.lWS.~ MIC:HAIHIl), ((lil&} IRmCH:ARDA.. 'm~HIl,~ S'fEiIPHEN [I', IiiI:OUSrolN..~ KAIRIk A. llAU BE,,~ .AU1RI~D G) '!)~lGADOI CA.LIDi.R;OirJ.IJ
C10ast
OF THE SU'iNI~ICANICE
autlmrs also do not IlJJ:clltion I,h:lt ahhongh some m~lfgilln.c..~received sediments, f! nGI ImOl~of 562 klill:! of coastal rn<lrsites. nalum:ll
levee I'.idges.~md b~urii!'r [:ii:~~lIldswa£convened io open ,VlIiler dmung the l'l:wiThumieatiies I( I, 2"
i~~ml.w d@1 an"5lillJlW NadaJ]a~. d@ .AmI mpeta g i~ 'lilistmifl, e l!lei!'lj,1O .~lJ~riJ?,z N!ifl'!~'1'i) ~.2~;·iUl, Ve~acf~, Me:d(iII" 2IMli~llIl~ ,d~ Milli:pc1o-gia !leo la Unhi\EI~id'ad 'ilEIJIl(]flJZilUlil,.
Smree', Nilr.w ,",21~en, CT 06-:511, USA. ~iDeiP"v!!Jf!e!1~ at IImJllfm)pOwflY'. l!I!I1i~E1r~iwW Alllbal1llll, lu!icalo~a, At
~Jj!6 lliooon
Turner et nl.. v.,su~~)' refer lo tlre sedimentd~p()sited 'by du: t~MG! ~~urricflnc~ as 11(1\1;;.~dimerHs. in~1[) the ~~HU:;,~" f"\"'1~~hmd i.~g '~ITiY...d "frcun aft~hufi~," R('::;'l~Sp~~lSiu" se d:i menta I iO:~l from H U:I"I" ica nes Karri 11<1 and lI:"'lnspo~;I: of sedimenrs from oflshmrc ~ fm d Ril"'," Repons~ 20 Of.:L 2 U06,.p .•449'). .!nd ddw-pla in sources by storms are <~mo~lg~ Ahhongh this 'lm.€!W stud y adds to tl1,c;: odly b liliu,:!'m.ech!l:un,~smls hy which sedlme]1lts oll'igj~ of knowledge l1ont'~ming the role oftr{''1pi'~ mil]), t-iupp,lied byri'lrcr's tire dlis~ibllt~~d ~ cal stormsin s.edlimen:1 n::di'snibulion. the ac russ the M ississ i'PI)i m.HI ether dehJs ~.
I)h~~i;;ilhU1itl~d:u:l!:I!ierlmtiorl! dm·j.UiIgHurricanes Rila ,~md KI!~l'1l11a, It. E.. HLlli':mif:r ~n1\d ee..llJ..ll1mors ecaeluded Hm~ "riverine sources br1n~I'\C"I[ iv~.iy (wi vil'l~. am.QUrdii ofillorgilni~
from Sdence illS soon as it is publlshed, Sngn up fm,enlf,e";alert S,~rvi(e5 andlVQu, csn Ikl1O'1'1!' when the lilil'€stiswe of Science or Science ExpreScs has oee{1 pested,
p~n.!!s,ethe t~test ~alMeof ,COrtilenl$ for Science or Sciellce'S Signall Tral'ls.d.LliC~rUrikirnow[edglB' EniVTr()n· rnsnt,and r~d stlnllmarlif:.s of [he tOl!ma~'s re:sea rch, news
:Sl'If.1.ETIOC
We6i1!j't;aMe'
'C1Fcootent5,
cents nt, or Edito:rs' Choice (jolumn, a~l~rom your e"m~i:l ifllOOX. To start rr,e~ejv[ngecmBlill.updaltes, go to:
f:ditQf$"IO!l!o!~ H'~Iii1'1hts·otUIII~·
r~05nt lil~rili~j,j'@
Imttp:/ fV!l'lW!l.scl@ri('@;JiI'lig.orgl@mi!
,COI'llt!:nt
.lETTERS
i(3-6j. Th~~:!samples, m(l\i;~[),1~;a~~~l'Q !llOll!ry k ~10me I ers i '1'111<111 We: re no l ana I y zed in d, a mi!n[!~r thatwould ~H~If~1bI,-, [I[) deterthem ml[)le the source, ie .. offshore or inshore
\!ri~uterbOUIQ]1!;].l;. b~~wr~cl" islands, ar pretRcl'er.eIrl(t:. and N\QI~e:5
1. JlreLlmil'iili'l J5~~lIN!l: or I~(!I Vittllilnd II.c:lm~. aJliI!r Hwrii'Y!~t::i!lrirnOl o:i!lt! M~'!'!'e~~lrell:iIl(l!! 111 t.liie 1,Ji.). ~dg9i~i!I.~r~on:~ f/Jg'l\. 005 2
II
of an uaeonfieed
e;l!jsling marshes, The (lQm~~ril>Ol~ ofth~ K(dtll(lJU b;:;:!liit:nl~ of ~.lll1f'fic.mes '\Vil~.l the bene f.i H., o:W tile Caernaevuu Fr-esJIlwater Diversion structure
U11(p,:!NJmI;niJjJC.iIJl~~.91l"'.IW!~a~~r05_00 i.ltilli'il. 2:. ~. A, IB.arr;JiJ:., !}. S. G«i~ .s~/lI'. Qpm-fil~ Rep. 1!1,1l·U'1l4.2000'~'OO ~J. ~. E. lImu,pj,. ~n,~ GriM/! 81llin~ and M\11I1IIiru:, VM..J~ ~il'l.il~~Y of 'hi" GrJ.1/ oj .M!!.I;l'..--O (Nj~·(~mb'JNJ.n Sto. A. l t,!. ~(li!'!'\,. r. G. :SteN.~ ~i!I~.(KIi~wlTIl!lei~ ~e"\I '!'aft, '
]~75).
chf~nnel [IUHj rhrolllgh crevasses l:1m, (;(ln~ (l~~ s true lied levees, b u~ not <II the river mouth. They take us. to task for In il1i~nli::d.ng
by Turner
Cl
,rd, \\';)1::5
OUl~ Qf C~lll~e;'I:;t
l'igItCJ;il~j~js·
strucrurewas
des igned iat he IIIid - ~. &0s 9 1{I~on\'>i;ly filitlsh w.,lter «(;lot sediments), to manage s<lUnirrie:s. and [0 benefit oysters.
Whhol!l~ IrIi{!jor 5O~dimcl'l~ CQnlrib~ll~loIlS by ~heriver, ''R!di~el.ns. ~di:sn·.ulmled by stcsms
4. [I.. 1Ii~.lmJllrl!! :1.1,22;! t:jj9tl9~. ~ :I. L.~. Wrigll1\, c, A. ~ilru;oll(,I,.~lWlTfill1 :IS', ·4')4 m9'S,~. ill. ~.l'.'l,COleman. fl. It 1~1l:, G. W. ~~!:.1. (WSl(1r Rrt.
l~,ssa (19~'IIj,
'7. flf:OC,~pOl't'QI'lh
Uui!.lrwrh'!Jll!r(~~tgr9l'l~~n~!r~.pdf)
(~Q6).
arc l1lB domi !KI!'!~~dhll.l,;m !\()M.n;:~f<:frW~~1001m.Gs. Comraryto tl11e eonelusiuns ofTumer ('/ at.. tWO recent ~~i<lmiric panels havl;; l'UI~;estecl th.ll more ~n::s~ive diversion of I,he MiS:l>b.si.I~'Pi Rlverwaier mh:~ ~c(Hrnlcmi>is needed W build .m1.1:l1il<iifitailii ~<lllclif! omlslall Louisiaoa (7. t~)_
VilIRGI N nA ~ijIRK!EiIiT,.1 C, Gi. G'IitOAiI"~~ [I E.NIS.iE RJE EI[I~
Illl!(.~~tte, U\ n~1j9. UI$..tI.2.l~d~oli'l Sdhoo\ of 'Geopc!elf!C!l$, UI~illl!f~i1;y Te(!!~~1 UI1~~II!f 'S~~UOI~ of CflOo. A~im. rx
g. rI:i1il)i'I~1 ~o!i~th (j)i.ii"l~gll DJirl.,.J~ ~oo1s~'~' l\I'J!'JIi' MJ'lPl A'IJitfMUil'lf £lIl'mHO'liIi il'J ('oru;rill' lo.~rgf~~1'1 (J!1aii;lDnilljk~em.l~. ~r~J, ~~~i!il!iioJ1.l.'Ic;,'(ID61,
1lp-l]~'""l[HL
Res.ponSIf!'
BLIIR:KElil HAL ,A.CKMDWILHli(]E TtHA:T OFFshore sediments provide eoasral marshes with iliilor~aliiljJt; ~1l:dimc:~.t1O and ~Iuuour qUlllmific(\t~QI'l ofdn::s:e sediments "addis 1:0 the body of k now h::dJge ooJm,{;erning~:hc role of tropical storms in sedimeta redisrribud nn," They clearly realize ltlmet we made ('om~l<tiiSl(llllliof l~e ql!ll<]lIllhy of hurricanederived inergauic :sed.iI11lCI:lIS \\rw'~h the
the eontributions of river dn\'er$:~i(Jn to III a.:~.Wlt~ln ingwe Itt~ nd ~Q~~B .• wh ic h b; 11u i approach lhey r,)Vor. <l.iliuiWor i.gnOli'i:lm.s: wetI.arul loss c~msedJb)' hurricanes. T~~ey alse meutiouthar P;VO t~t~I;l!il~' panels endorse r.~vcr diversions .. We made ul!llr CO:~1I.1.p'!lri.:S!Jfl5 about sediment de! ivcry because iris part Qn.mHdi~lga sediment bl~dglC[ for this Cfl.l;l<;;;L There am Imblished estimates iJ;\'~ti.. <}b]e Jllld v:U.iOLJlS ~ !1lO'lnrl1,l:~mcl'u drl!l;!um~!nsth~n bighi~.g[llt ~d~·· ment :iint1~oduCli()lls vi;!. divi:Ji"',!;iom; as <Ii positive alnihuu~ UII.\""(~ did nut examine The I'igni.fic:,uu ~·mrric'LI1e·iIldm;li;-d :1Ii.lIOlfll.h-m-open "Wilereonversions, whlch may ~m be perm
;In.elli l
p). be cause
~i;(1.I:t .
\'\."IS
7871l2~.2S4
search (lbj~c[ivc.. We' o tTellfed, <I hypo t hesis, not <I. cone hll s. ion. 1hal these :n.c se<Bw'nn.e; nrs C <I UiI:U: w from ihe sll!lUoW w.lter ZOJ!l.e immedi~l~dy of~"'sholl'c' of the depositlen site .. The Chenier NOlin in Ib!;!: western half (If The state reeeived abom ;5.0% OfpO!![~HLr.r.i-
lambda .OG-4
Hiqh-speed
wavelength switcher
And vE!'rs;atiiitle~~'fhe Lambda DG~4offers rleall·t~me video' and dU<311wd'v!I;'l'englth ratio jlmo;l:giing with Ulr!lif.ormspatiilli iIIIuminat.~on and integl'al r!I.el!J!'Uatl dens,ity fil~.tJer ilng •
iNOi1!J1 ~SCID
IMil~,
ClllslOIi1
Xenoglrafl
Studl:es
We. Oilier 8:Xl.oellent: avai~abmry of nirU'~·lmrrunodefioient !rodent mod\l~Hs. ur Precllinioolt O SSlNioes grOLlip also !provides xenogf8!ft and otiler oncoioQY":based servlces.
._,Up to 4 interference 'filters (5 ava'iJabie ·on CHi-5;· - 1 m5~Cfilter to filter swi't-chi11'g - P'f:f;!·-i)iig'ned 175 W ,xenon
light 50Uf:Ce • Prog{:r'Jmm~b}e .~ttefl't,uatiQ:n'
Felalt!l.llfes:
..Adaptable ro .MQ.U
micr'05'Copes
.~
Eilllmpe~ ~nfa@elllw~rI.cOIIIiI iMW•. CRilVUi.ODMI
'S~t"".(lR~!iiI' .. 1
CHIAlRlES RIVER
~."lirdl Mod~,s and StNvitJE$
SUTTIER
EMA.UL,
IINSTR'UME':NT
I FAX;.: .4J~~~"ae3.0~"::ll
LABORAlI':OR~~
II!Nf'\CH~S.lJT1II'ER'.COMI U 'llVWW.SUT'lliER.COIM
SC;II~Nelc.
vo L 31 S
9 MAIRCH .2007
136'7
LETTERS
cane
sed~.mlrn.elUeposition. d sedimem
has no
hl.n'gc
'"lS <l
b~}':$ ~,(I
ae l
S;WD"<I~"C site •
ne wl y d epos ited sed in%etl rs oIL( the eeeanie shoreline Welte mostl;y sands, Burlk~u {;i (:II. ,offer no ~~t~rn~,tjvc:hYj'lot:~csis for t~le ne'~'ly deposited C~l.e:ljliier PI<'Iin ~dhn~nt 5;lll~~!,JC~S.Surl,jlyrh~ orher sn%
is P<lf!,'uaUj'
We d id no '., ,;uHI do not. 11K. kea recomI1lCnd,ll]UOI1J I!lOOl,ll buildlng "iv(!!:I" dlversions.
There are m<JII1Y factors influencing
such
management ehelces, inclm!ing S!ciC]1li Fi:e .:an<l~,yseiS and sociopofirical aspects, We trus t l:I~.~llrepulali:llc se i,enOepIlIUII,f:']:S and ~lil$iShtflll !ldmiw~~'tnuClrs mi~bt\I\·i!).h. t(l 1·~ce.UVC.IiI.Ol CXe]I~ldc. uew~uformalio:~ll. Fo:r example.we O1gree~'I.'il~, Burkeu e/ (II. ~h'll
~;,ii1I:~~,:u!y ontrol <lit the Cae,m'U;'VOIl, diverc sion it> f~ ~,h::"ig~bjl,,'i~~iv{': o nW1HU~(I c;;~h'IP~~~ oyster yields, Would <I scieece panel not cons id~:r !lif;.'i',\,' in form.rt i.QfI ifldk'al i!'l:g I be dilfl~.i:lm~:ihedw:u~h addmdonall 1:liCshwmcli"
f~.,
for
Bu~ldullg ~ marc mbus:t Slcdimcrll hudget [he-sill coastal wetlands can be part of
Lou i~hull" 5t.a,te unt\<e,r~.IW, ilatoo Rwg;e. LA 70'S/~), UStl .. 1'Worlh ~fi~·•. wrfll1ah EI<I~' N~!10i1a,1 ~ E~h,i~rli\~ R:6earch R>e'5E!tliie.
fll:i5t Offim Sox 1~~0, (l!mJg~lO~,
R~ri~VOOCi~
5C .294,<l,2, USA.
1L
'U,i
IJ1ill:p',/~m\i'fi,~~OK~anrr~.m'lll!~ljt(:aoJ!nrn~r\!Ont
Your ca;rleer is too importan~ to I~eav'eto chan!!:e. So to Hlild the right job or get career adviee, turn t@l theexperts, A~
[.0
crm .~~ Op£tl,JJIt: Rril" 2M'." ::I!2'~ utGIM}, 3. R. E. T!lHler;, BrulNie~ Cr)!1!~ts :!:~:5flOO61. 29',
2.
Gi~nt;li'Io~.Ii~ITii).
J. A.B.!!rr~$,u,s.
se i,ene!:!:. \n~itwW'w.•Scienc;eCaw'e\e
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COMPUTERS
who~.e~!ltory
\~,\'lek·striip.
p¢r0ep~~OJ!i!~ftller re.mlimlg a ,l!uideibo nk ~md <l Fin.10f,dl. giO l!'liu:k 10 allJi;:,'l.'!l~ seme of ~he orlginel SQ~trecs.. ~ fear ym.lnlgcr readers .U'lm.ighl !1ew~r ~~v,~ a Whole Erl,I#" ('cutdng. h &6\~n m<L;rno~ g.e~:W]nl:1iI, the fuss is about just fr'UlnI;l fl;'adinS Tli1m~r: .. di.;;~·ri1:'Ti~'i!l1> ;;lind Ithi,;;b~li]k'$ rUizzy 1-eJ:nudm::t iio.lll.!>flulO j)'OlIges, Ted Nelsm:l S o ompUlt'r Uhti}~~(lm McrdihrJ;:;'i! di ;.;mi~d ( !). by TUlrm::1" in a :;ingle sentence, ,di:e\v the 1;:X:HJlm", ~CI:'CU~,D,lro·rc:ybel\C~dull.'C ("f)mlml.oclion e\re~m1. ore m expli.dl:ly d'm,1Ji jtlholc earI'll, S'teNe:~:l Levy's llc:ldtf:i':!if Cl) is also a must. B,ffind's mVIl short
Im!.;jt~~l~\l"l;" tl~l\VO'rkin whi.ch a~.mru[ anybody (:,m r~II,L p~~Mi<,~"H!fdl~'l;f",~mm ; PMty mach ,.111)'tldng.
e'll,lf:
pr.t:tty
d.' <I~l~,nIm:ky
=. did:il ''Vorldw!!.Ii::
n~y
gO. ·llhe
<1.1OJiI.e sli ee dm:mgh lhii s marvel eus sl;a~)', Umilik¢ lll~mty ol:htT his~t'lri~s ~h1!l fg(.':u:o:~lilrhe
technical
ffinIllO'mliOrS~~~]e
CiQu:ld IlfI~ been domiuni11ll~ by a I~w I\Xlr~1m.'a'tiOIlS,., spoon-feeding junk-lbcd media to the masses, jus;I.li~ t~k;vis;i:on_ 0.1" balkanized cmmnulJJi.(,,;J:tio~lLs providers ,oo~Lldh;we s:oodlt1d ~~l1fS"."ith d~"C!f;:pli'!.f€: chflrgjng sel'M;;m.;"~ O'lOO ~"1ifled li(lchini(',11 ir1lmJli...~lioo.jll!i\·1~l\e l~~pll(a'e~. . ,C
Berners-Lees,
t[>g~
omD~:alw:ay.h
Mi nskys---<thi" <ll..CC01IUlIl:focuses Oml, ,Ike)' p.I<'I.}'ill' ,'\i~~.QSCrole w~s ~n~kh~glh~ CQLlmcreu~~[JlIrecyberclJllture OOlm.ectU10Ili: Sle,\'<UI BrmJld. j3,lllml i.;. C'omriblJI[ UOIm.WfIlS,verarDi f1J,~ em ahi!O p~en()me:~.olrn,: tftl.emizilllg <]bmn iI:: '~p~Ll;lrizins u: cih~~r>ll(;,:;lding. for it: aad (l~gfm~ting. mnwnlt-
'1:lIII,d ililS'p~r]llg'·Sp.u::ewar: F.l'Im)tLc l.ifennd Sym'oolic [)e,al:li~, nmmg tlhe COJllpUlif!U' B nm!i;" A
il1~..,a oopnwnding.rum~roll:>
fbit
ulmetl tl.'C~Ul~pDkc:li~ ,oonJSC.~n:l>LJI:!> v'l~lles of llm~ cO'n1~nuirl[!ies, ]ft> 1111.001 to ~ ~x:~lctl.y \vl'iiat he did. b~I.~: everybody knew hi llill •. <Imllhillt sure helped. TlltH!gh~h~bm~k~i1:;9~ .h'lls d personal de~:<Ii]jsof Bm~ul:s Ii:Fe.. tlD5 nUL <l bi.(lgm]Jhy. 111{,3ther. h :fQ(;nses, 0111 ,evelllt;:;; thm S'lvir~around him" It traces his illli,,-oI\lenit.'1U wi,tiill 1960:; ,COlllmlU!bC$. ~f1id ~oill(leprul art ClOllmH!!1IJllities, Tl~,e JVfuJ!(!!:.al'ill Cm(~/(JgOIl1d l1~i!U we ~mt>p.eiledjn ~'ls,!!1I.ch 'M} open net\w:rlk was ~ Min:sc I~, Em il\~l>~)'t mil ace i~~lu. m(l~i!'l:~s wt\l~d~s .~.ldml of primliHvlil ·111n'~~ I~.'t rel],,-"C~allhe ,,-'Oil1The l~e h'llk~ti COiil'l:ii'llilini~y [h~!l b'uiIt tad!!Y'$' c10py \~r~b'j(Iii~~~~ I
digutall inFrastnlclure
::1eI: f o
~~'" p~ti~fl{:l~in 'thl;i cybl:!'~uh ure, d~!I;,C{IIJ!.~~~rc ulture, 0 r bolh., But when you "re .11: <lcly to understand :~~(:i\\' thm ell!thu;ji.;;')'im~O!t u s 10 where '!,Ne~JIie ~.od'1Y,. re.,d Turner, To get you sl~rl;od. r~~~c!lve ~-'OUI with Br.nld'g illtroduC'~ion W ~he ··ftmali.c lire"
pieee
(3):
!lOt. C{llfIljllttC'f.'S ;m!
eOluir!g to
dl:Id so around
(I
certam
~non 'IIaI1uc!':,of eth conununities and led 10 b IJ~nld.The WELL, anearly Irn.l,e:-;::mge board
~~l ariQ)g.. perscnal ex pr'e;ssi.o~. a Ild :~nn,{:watrion, These were cere values ohheearly digi~<L1 piuneer~ (the ha eke rs) •. embodi.ed in what we pnmclly rhe •.h<!~:kt'lr ~~fuli~." Today. \\~J;! l<lkie lll.e digil;)~. r~'\!ol,tU io III fo:!" gra1m.~cd ::1:11,(11 l'iidd(nn .ap~~1'C-
or M~JIKLce?'· ghQol
of liberal crluelsm
1'D IC.yIhEm: u Un r~
Stewart Brand, l~le
p<Utidp<lI1:~S:,
c.~n
AJ IIli.allgful. be misled
into lhi~]king
in\i(;riI:t'd
~.h~\VE II
wd.a)' :~'fflr-
that
U~ll,wilh the mmarnic faUlt;.!sic.s Qr tlllc rllI"Cf!l!.ll~cr~ ,'he seleace j]9chas of No.roc'.r'i \Vucrn.cr, WlIJITI.::I1McOulkldl, J. C.
bur sLlrp:dshn.gly
ill,
r~.Lkk~hi(."r.
The'
lrcm~
Jnhn
V(ln
NCl!ll~lillm
and
'ifJlIlMV'JIF
lBu1in.
O'I.I!,\C.~
cli.1lle to
ealled \Wb 2.0. il \1.':iIIs0(.'rminlya stE'pl:o'Wl'Jrd t:l1esephellOmen'J. BnmdJ's Ghih0.l BtL~,i~l~~:> N~twOlik, a 'C:O 11 suh:ung corm;pal]l¥.
i.1t: h.cl.l~th1:0,[1'1!;l(k!army
:z
the
~'liJed make ,emmICl.'Claltlimll and to ,ybeTCU~ll![cl idc,~ ,m~c~si:ble to' hig~rk:\'el (:orporar[epJ!~m:ml.Cil':l"
iSlhlt
ef ifll~,ltIell~\::>: The YOl1ilhfi.l:l, fcirvo·r J!i.I:i,d :Wi.rmdb- st'IMi!i.hm!~n!:al1i.ulJism of •.he :r:Ii'C".flk5 1;\~~Q cli:~;!:l.:tI cmta p~lIlCr SC~'C'Ji'U;:C~ I:I.IJ, >!!'''W!li:;:!'!i!l~!y~I:l!i!gh')~!l~d research p:ro· grdi'UI frmn the very top ~rth;~' ICh::ttlllt'1,;
:[]c-pllr~;n:u:nt; ,Ill IJIlil.~X pm::wd liI.larke'I.·'
I
i
=
Tur.mI.C'l' fOl'~m::rjOU1:'fllfllliS'l now ~fliite DepOrl= fa ~'~l ofCml1!UUlliC(lliotl. SIl.'111fO!'dUnivG:.r&~ry) tfiie<llly took~e time to swei;1tmhe (jet.ilk The,we 0l!'1<l nl]!ri<ld tOf n.lSCi~l(;ni~gliu~~ ~is~or~{:fd ,a det3ils thal he dJM~ til' that will sUlI:'I~.l'ise'itmd
nanking J.tlqvcm~llt b}' the 1'lunUlf~ctmcrs o:hnmH c;]k'Lliiil[il'l,g IlIlm:::lil ir",;:l'i:and an iii· repr~ssiMe l'irlidnight Ilh.CImIllC:tlOIii. ](110'"111
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e!'lI~~ht.J,;i1lI;':V"'l1. ~hl;: key plflYi;:n; ~:~ Ih~ I.h,rlil~fl ",'\.'m.o pick lip thi:s hook He doeS;Iii'~, 11\,\"<'1.)15 get 1 tb.ung. .. right, thrn,Jgh. ~.\\''00 -t. q~~i.bb.l: 'iI\~lhV',1:~i. . 011:1 im.l.c:om.lCieslton.~ my J'.le~on<il kliiaw.ledge but o'n I)' $l1Y[hat it i;hou I~!l'tct~~oll~Y lmQk b ),011 I:e,nl 011 dl~5 :SWJ.bject. Saa,m::t:Ume:s TumcT:S <liCQOlunt hillS tI'I31 niO~-wl11iIm;g~but-'nOI-<luite_;I:~u:~ SC;II~Nelc.va L 31 S
::1..
!lI,r..;e~, ,U~~f,tmcl'
1[,
COiiliFUlertil}.;'~ Ca!'l!lllu/.li!~t
(;QtI1pU!~ N~~ill'JMq(_lrjtJ~: ~
tT.lt ~e~, 'ChiraSO. 11:97;(1). 2.S;. ~ H~~: Hel'!},~~riiJe C();I!1pr,IW tIi'~~ri~1J ma~.ed~y, tGardm (~'I': ~'I'. 1~4)'. :l_S_lIirand, R(flJi~9'Sj~'. 1 f!)~mberl W:J;, ~p. S'!)~~,
"e.f,rJfJ~ril1'/ol!(fll C~mp~!w~i:1em'-Alj~fif,ljNily&l'pfJl1
9 MAIRCH ,2007
I
lEN DOC R IINIO lO GY
Se;q;l1IardamlOUllm::d. that he had "JeJu vemuccl" hi msel f'wl thunjocl['O]lS of 1~s;~jcU].~lr rraees (he :sde~I~:fk discourse t~£Il underex tm(~s 01" dog:) andgu iaea pigs, nml~;med :S~einach.·s model Tillis episode marks the s;lbghtly of liIIllCl~O]il 11I~y opposed lila! ,l\I]dlIs1 Se·eret sex g~m.uis, suel.l ~IS !Dorothy elll1Jb<lJ'I'lI:f>silligoaaneucemem e of Qunl1lt@)s5i~'Ulcl3ll1f me modern el1dlocri.n~,ology. a begmPrice 21ml CJr'~ Nklo.re:S Sex, G.lands, and ] 932 anncuncemem uf .1] nil1l; lhm l':ilif1ic~~~ l'\J:ldllCri~W:~ll~i~t H (I rm on e-s, Hl501950 Hans USse,L' i:~,,~957£~J~I l:eg3ruooi teedback loop b~I.,...Oen the ' as "c<l~,mnirOlL$..'"IEmbarras."lfl~. pi·tuiHH'Ji .Irld tht: g'l1~ndS,'llld li'Y t:hMdfik S:rm[jIJ()pta' Bcrll~UlI:dZo]1de:k's .~934 b~~IIJ~e' BI,'own-S~iqll,.HIl~1died 9111 UqlPv.ersity a,1Chqcago dis¢t",'~ry of ~i!"(lgil;~njill th~ U~94, O~l~.y.1 f~w YG~iI"~f:"r~r his P rB 5:>, Ch icag{].l:006. 366 lamous se,lf-ell!:peril,ne:mt His, [eju~uine ofa stali~.iO:I1I. j1p. $45, (2H_511 ISB N '9];1l()2261~,g.634, .By thi: ~94il)s. '",n,lm; ri\~mllin~ h~d nm~e!~ f~~SI!lC(:lI,;.."z;;.. lulas claimed. e.,.Ianmitcms, be-
II
.•Ilt[lgon~ srn pelfSi~~ed WI'!~~c difdiypnu; Dic.;e f ! (I cliniea] praetice Q]' '3~ues. 'fake sex ;I!IfI.tfl~l)~ Ilc di.ni.c. [Ill.]~e~. '9~0s, aud 19505. eudecrinologJr~ts. studied the~hui()11 be~:li\'t'leni1!1)mH.1~e:;; nism !O:Le:..... np~e.In lheI910:.l,. Ihere~lQ.W!1ed n Vie~nu:scphys.iol.og~5a Eugen St~'illma.ch·s ex~nd 'ca~c\:r" lan;:~sl cancer. fGlu exsmple, \VJS perimems led him '~;obelieve th,;] I. male and !~.mlshl, W be caused by estregeas, If 11;0llfenb<I~:esex ~I<mds were in d keel, OIlP'Osir.ion to. m{llml.escould cause ii, Cit-midi they <l~so cure if? ~lil~ ;moth~r. thli!>.",S;l~lbli~hil)Jg d~!;;1.111ImJ.~ptlf l $t!l;)!;) red by ~mrir~'~~I~ t h~r~poei:,j~ ri;i~~ij~h ie Sex.UOII <l.1111.<II,goa.1US!I.n, The idea, thfll: male and '(guided by di:lm~,ca,1res~IJls. rather thau biofemotl e ~ri;;lilimny~p-jl'\~:!i,~I:~i;;en:iIIi~]y~~I;h~llill il;~it vrp~\r:ysi(l~.lJgi-c;dl·ind]ng~). di nic~.a[l recommended s ~ndmgenlLc a ud o;rmll~:-· m13~.0d with t~LilInlD:-JiII.gender mlQlIl1ns,. SengooPlOi
pofta[!~
o Ib~~l.(:uiwrol I
C(l~lC!f;i:Pt
of sexual
estrogenic rreauncnts, s.CliIgnopta :>~lOWS, il~,o~v Im,1:' reascni 1.£ he11 inei ~hese ;adlli1l!iu.edIy S'UlCres.....l1lil.endocrine Er~(mcnts of bre&'i.l cancer was d r,~\I'elm,by an outl.m]d\ed 'concept1Lmra. th':ll
ass,UI.n.n,edl:~.a.tI:I'n.eepposaehcrmone acted .l!IIlt;l~tff:lliM iC<liIYl)l1tlile- body. Sengoopta·,S e'.'q~,~.or.~l of the relariens ion ''1m(Ji~gt~le i~~I:!i~ectillS fon;e~ uf scieaee, cHnlie. ~nd c;: ulture deepeas 0I1I', unde.rstmuHng
of~iI;: ~:h~v~d(jp~.l~1mt QI·:m~)d~m 'I,.~~~{krt;;:rin()I()gy. lese<1,n;;IiI. on se.x, giam~its was shaped! bjr! h.e d i~ illid<Il!~~'lhe~!PGlBt.k: dPt~mi~~.n,()!ld ,H~interest h1 t1ln.epl:uysiologiil::<I~.englneeri IlJg of Ih.e
cause the OO!U10J1nlP[ ~~ml idi01.!~~ r he aroused near1Jy kUIed the b~ldc.]iug science. )
NUl !>~Jpri:"'[I1,g~y,tbis aspect of cmJocr.h.mlog i~ put 01:1 dle frOOi I'''lges. 0 f dmJ': llIJk,:~~1 annals, II does. however. open '. h~IHI",,];; Sengoopta :s.1n1lgte:rriy new hi:!>wry of £f'XlI<I] endec'ri~ol(I~W. Th~ M.f1.'11 S"en:! QUiflte.I~,wmC'~· ~f t Liji!. Examining ~I ceaurry's scientific <md ~.~QP11lufl~clmn~lon 'with e'I'lNlil:loCrirlegl~[i],ds irncl thci r Cr:ttcis on the 'body" SC'Hgool'la. (il bistorhr!1, ,~:flnG-dici~l~ and scie~c€' ,;It B,lq:lkbe.ck CoUcge. UrtI"u\fers,ity of London 1 conclndes th,)t in the l'flc;llt W'l~ .-~~~ ubOtLII sex _S~:.: \Vo;lS, wen "u the center 0:1' life. V]l:il]~.~y.and energy. and for.~time, gl~~nd~s!,jil;ll'1h;'d~n th~ keys h{lld te ~.~, "li!>foct!~ o:~ se:x allows Sengeopra to fol'~ov.r :t~uuhipk': hL~1.aric2l~:~Ialiorl~ll.ill$.. ~Ir'ti:td~ldythose between eli ill.lic:r;J1J] S and ba sic s:~~i~ijH~ltI;!Ind bI;:EW~!i:$i! bim~1e;:dleii1:~!ind the ,viderpor.m.I<l1' cu ~ture. The::sere:llat&onship.s Vi,1'eI'C dym,3'nm,ic. hll, the'
CII sc:~e]iilCe \\1',JS never
hM'1!,m body ..
th~' rd~Ltio~hipbenl'l;).C!l
moderu, cO"~:l;We.~Jng recent ~~igh1y :Jj)Ulb[iei ;t:t;d deba l'I::~,ib C:ilit the c:oh!t1e.c ~icn betwee:~, horrnene replacement dlel1llp)" f!Jrme:[Iop,lm:od ,\iQmen .m~ breast
ciltl1'C~r.The use 01' hermenes t.o oomi:l(lt
tl'n(' .ag i[l1l. preeess d~dU1lo.t di~ with Bmwn~SeqJ[[.,u,jL It \I,!iil$ Ilopwl,lar,h;.!J!d by S;h;:hlf~dll JO )''G'[~rs I@t~r; ~L1]d I!rlday ;lUll!.• ;I g~~,£! hormones. t'I£,<L iu c apture the publi(:!.; <iu~mkl!'!, Se.~,g,O()Pi'~i..~ke~the science ofl~e pf~S:I:~C'ri.QU~[Y :s.mi m~ ill> mVIiIIU!.r.n~,s,
r~fl,1.~inS 1.0 e'v,<II.mtle
~he mod~1 ~fl ov~rJ;implifi0d: prf!V~U C!l~fOOe[_ h~$ ~~~o profu~lltdly afr'=lC'wd 'J:I.~dl)owells ~md eSll1og_ens VJ.'ere· fOIlII,nl~n both byiits [el:liUted~y 'il\flilJllg. :S1D'all,ge, or ODSC L.ure ~C\'i.Ipc.),.t'l,;:rrIS Ih{l Jf~mh("m!(,flH'lr tlHhc n~~vly \"'OJll~!l ~rld ffiVll, As :S~"~O(1pw.-,ptl~ pUl!> i~c. p.;:!Sl. S(!m~'~im!;;s ··fals~" scientific th(lor~~s ~. "II,~mSl ilIlulhorili~ lrnad.. by the end of the lhir~ l)lIl"ifi.'ed honnQi.~::t1 C~I.rm::'tl>. T~ll.'{l1IgJbom. the' 'pmdutie the' r:ighl tllemments. SOIl1.letiil~le.~ t~.ey 9 li~:s. come ·~.tlrt::g'ird d~e: S.~H,,~!d$ !he' SUitl ;"$ imlspi ro .-~~.'~'i>!fI:~hlradil ifMl lhf!t $,~~lpl,l'ge:rntir~ ~ M.;i~,ee·ri~g Oli!iP~Cl Sidemif~c med!ci~l~ of sh'lped re-,seun:h '(Ill t~lle :lie:xg~'llids and its. inSt1lt!.U~lem~ illU the ,em.lcerto gf Se11'1. essenli,ll di~ipli'Ui.es. rn rh~ Mf}:.'fItS(!Cr,'l fluillfe8SCI.rc,·(' ' ~ d ink:a~ ic,1th:ms.. lIppl but not sulrk:k~&in i:h~m!>f1!lv~", endmM I:h~ (~f ~/ft.,eIliJo~ri.l1o.log y cnmerg.cs .1!> pal,t of ~ ~. to L body \v.ilh virij~i'ty or lWI1IiIinin~.I\y" Oil1,1eT imtru· proc~s.. ofm dYIJMlmic KI;,H ionship alllolli,g c~ll- ;S !n~nl;$ had to work i,~iliHII~lO~yith~'fuiG]1J~ w Il,i!~ L 'c~[ni,,,';lI.I,.~uld fie h):]l,tifCc :rorc(lsr:!1~~l.G'rl~~f! ~ lie re'tlewe-.r ~ at ~ihe' Imtitll!e 01 ~he HJ~l'Dry Mte-d~dml!', of under the b<l~on of a COIll,d~q.cliOr: the' e\I~~wnat'i,c <Jill l1icIS\1!~dit'ill.!;lto be di~c:oveil,';ed.. ~ !oI!fi~ ~>op~lfi£ UIiliIYeiF1iilyd!o>o~ ot Merdil:in~. Ilammo~!e'. S MD212JllS-2:16?, U$.I'I.E~m<iil: stder!@ilnfl'li~u pilWI,itary gland.'· 1O-:rH6I>Ciernce,lll]91J~Hl e;
!lologhl;~ saw
1920!li.t~leg{l~d~~l (Jfth~ =-~N,,g~mHI~, <lg~ m,-lil\~ b.lbor..u:ory ~.~'Q:rk as done by 'o]l~nici.amcS-.ull.d w driven by dil'lic.rl~ 'lgell1l,d9s.l[I rhe 1930 s, the clini!ci:PIn·s hcgclnony wBined <]50 Illu: 5n1r of basic bioch(l~uis~:)"y1"(\&c,t!"d~ l"OS~_ SC!~guopta SI.lOWS, •. h O'!jM-ever. Ihal; the d, in iti :Ul. g;l:irUil.ed
ju~g~its v~iij~ b)l2l;>~-~\;n!1I,lry ~!~ndOl'lJl(kAt f~r:sl &b'I~l:e. t~Jis migh:~ cl~OIppoi:m,l anyone butll:u:stor:ialls . .A n.Cli 311•. ,\'hy :;h{}~~dwe r~adI aoout i'ide:nli fie
theories thiU have been proven wmn.gl
QUlle:
.'jiU'l'ilip~},.~lilodel-,1!1 scienceis
IDill],u-
9 MARCH 2007
VOL. 31 5
SCII!:.NCtc
www.,s-ciencema9'.o~g
ClllIMATIE.
b ~ sU~JlIum,id.
required redue ~kl!'! !>ch!;idu~~:; Ail r\,'I,I'l:.' '.lre GV'.~[to ~l~~'e~e,di!l <'ipr.i~~.g .. , C the I:miklup oflhe a~]l~!l6phe~~s:'(,OJ (."(l(l~te'n,t. ,lddilli;oH<l1 element would be neces\'I.'e must mOlt.:" f tlit-c~r~~ changein (hit:: :l..~ry,The S<'Ip (see f~gm:e. rigjHl and allowed emis''Vtl)1 we v,i:~wthe pmbl~m" Most pO:~~lci.e-s~:lun berweenactual II~ilIv~b-ccn discusS1xl~m;:liUdD~gc'~p-al~,~unde slum. \'\iOu'kll hfli\1e 1:0 be Iml.d~ up I:rilher by pWI.r.cliuase ,o:reol a]~:oc:.ned sys.l:eil,llS and the Kyoto lreHly. haveneared the rl!'(~blenm, e.."1.chH;;,ive~,y hlii. u:~rms; <0 r i!lCrClmle!'n:tal to poorer narions or by burial of roductirln!'in cn2~mi~i~~I1!', l1~~.11{}1;.~e\l~r. CO2 Ct'lph!!'l;;'iI from rhe annesT p.h;ere. SH'!Huninu:. the rise in Co., wi~,~aot sl:abi~iu'311nosphe'D",ic CO~ levels; lhey only filow ~hen'lle (~I·i$lcre·~~~.I~:s[e.ml <lC[Uto \,,;olllid !require P1!rt1'cip;nJio:L~of mp: lIIf1y st{I(' the U,'~lCr(~ilIsC'. '!JI,,'C lllms'~, evC'lop the d ~.d!yi:ndWl£I.II"iHlz.ung mltiom; sHell. as ClOllCifolpl 0 f wh.u.migi'lloo c.~!Ied,l"~ ,n'bQn C~~rI<~ and India Under Ihe pie l'l'ie:·Cnlitml~.y. foreach-J gig<tlo:ms (GI) offb~ concept. 't~:1f:re 'would be ani iueentiVl;: fQri'~.I;'nl h~jei n RJ~theY\,V4;l1lld ~i~ ~~ror~!lbl~rn~d\ the 11.im.{*,p~~~ru cnn's, COl tent rises <lOO~I,t IpPI,n: im;;lud~,ng deforesiahavea cOl~s~der<lbljy 1,00l~er pel'iod
iW:l0
l\D~~
20JiJ'
,040
~(}5iO
2060
chase of the eneJgy required to accomplish I:~.C t~l)lp[urc;;n1dl~r~i1I,fthe CD:."Tof Sllmric iE'ul b l ;_(~II1I.~o~p1rnere0 increase, lh~mI. lili sl1.xe.s lire size 1 ~o!\.~i,1rue] to gene'r.lt~ this ~nJe~y is 000 hiJJ;h. IJ1~n Ihjs Slro:lJ.l!:'l,\\,I'HI~dbe in~p~lc'th;:~~I. of the carbon pic .: ~1:101"exa mp~e'.• this ~iIii\~H The ~.,,~esl ohhe costs asscciatedwith \""lilr~ [ll btl doubl,t.;; dl>e rrei!1dl~S~d~~~ ('02 1II.I1OOLnrnlt •• 560 Ilpm),. then t~.le S:ize of the pie ~ie :a~r-C'al)lme willi, be tillose asseciated 'wilhl~e' \vouJd he 711] Cit of earbon [i.e .. 4 x l560release ·ofthe CO~iom the CO;lptuiil;' m'I~~rU.:li:1 l:lllO)].. \\'ere the I ilm:il to be set <1.1450ppm, the .lnd with~11e recycl i:lJ1lg of allY ebem icals used, ~ize of ihe pie- ,,!,rmlld 'be lln~y,21'1.0 Gt, beimplemented, and faces, several ~echnQIO'cl:- A~ ~gJiimn hydro",icle ..• m obvious choice, Once Ihe size of pie :~~::; bec.~n,eSI<lMhihed. ~Ic,d c'h<!l~lenges, [f the COL carried byt~e ~~~ b(l,lds. l)(un,to CO'2 too tCrI!llcUotJ:dy. a bene. r ef~ch ~nh~ 'world ~flinm il1ns 'Ii\ 'fluld he (l11(]c~"II.Ci.¥.t Sl~.funs used m drivl;lwind turblnes wereto 'be op~ion \'i'O~111d. be O,'l ~~~~J;::ri;rd [~unw\ll1Idbe a,bl~ cap~lIIred ~~.ell enan energy-equivalenrbasis, a slice, ~Il an i.de<!l~ world" dUl' size of these ~o pick ~upC'O~ btl! would release it more slices \~'{lu:l~dbe ~uS£"d (.1'1 cpriruk~li.o!'l. llim (his llm);;pl'lly-,"k~~1i!1~e;iIiISi(l!l.j<j af the CO~ c~~ph.mife r~i.u.my.R~gartl]ess of'lvltrf~l m~.neril,1i~s to he d i case, the wU.Ii"Id's:rich countries would.get O/hlly devices would be: oml;! I % of the s\veep of used i.~.is llhso]tne.~ ,e!!se~ni:~llh;.n research on 1,l.b,cmi;,Hl'.4 afliLe pie. :~r[he~:Lmit. agreed lilI~xm cnprure Mel ~e(llleMn~lit"in b~ CO'lfDlI;i1 (Hll ~~) t.11ii.eUlibim::·s H ~,~n other WOilitlLs. in :.I sense, t were Silt) ppm, '~~!n.eil,l r:idl m'u.iom· share the ;llir streams C;J'fwy WO times mon:C02 rh,m de~,~Tm,~ne w~lelher the ,ene~ cu sf:.;c.m be broughl. do\l.~l an!lc>Of:'::pta~J!e It~v{~l t.o, C!lI~nUl~ 'muhl be abolll .1501 A.s these COUIlJII'ic.<> Idnel~!.! nc~. Ot. c u)~rel:her cuntt~l,n]y COI'ISlil me ~,OOL~~Gl of fo~ () In add iiion 'to .1]lo\vi n~ 'the gap betweeu ~~rom (,'O,,~ g<lsini:c~.lion pl<lmLS ~,hOlll.ld'I~£i.O be a,ehl~lalld p~rlni~si.b.i~, ctn~:!:s:i()n~ h) be si~ C:;ll1b[inp~r y~alt". if [[li;;Y ~o:~tjl:'lu(:'l(~ dl~,:; ful i.ri,~.pk:mcllt~d .. Iall the present p o~itical c],ima ~e..• my .,;[email protected];'!. th~if nllutmt;l!H 'Ii\'Quld. b~ o(}¥ilsllm~d iJ1l fi~h~~1.;;!ir~:x~!1J,c~iun hm; ntht;: r O1,Hn.(."i:h~I~tJ.-, just. 2: 5 yea rs.lIFaced w~l:l;l lhis ~,imit. em::hi 0 r lures., (~,) [t. cO~lld be done at !!,ite~ Y.u l'f1mu I.empt t9 achi:eve .m agr:ee:nm,em 0111eil:~,er the' ~h!i;:,';e nw kln5 \;\''Ou~clbIf;Rm;.,ed IO n~pid Iy po:p~.I~uli,O'llc~mer~ .md cI"):51;.'l bJ' d\e !'iite~ of ::;i~of;] Cilfbgn],>je ur it!'! ulku'::<I:tion .!W11:llll,g the rk:h reduce hs eiI,uissiQliliS(see ligu:re. iI'bcrIlc~,Poor cn, stol:<'lg;t::. (iii) Om~e the rise illl, cn, hnd \:vodd's mllinns,wouk'l be diftilcullo HO\?!'CVCli, ~l<:lJion;s wtmlcl: be ,a.ble ~o !iidl poniuEls; of their beell s~eil1,niiled. he ,CO~ cOlne~u of the m;gie5:1' we 'ld\' ..tm.cebey(lndlthink~J1g on~y ]11 ~ ,of,conscrvatuon am~ ill'tCriliiU,C sources p~Jf! ii;1:~ce~o til,e rilch cOi.1ll1tde:s :Jnd sliU ntave ~ lll1l1.0 spher,e COli hI be d~wn hack down to' 1(;!:rl1iili.'i e:m)ttgb ]eft 10 pCnli'JJi[ ~hc~~ [0 indl'M[~i~i:i!:(;l:" alcvd m t~.vh ieh th~ C'~~r.lll:); C!lpS \..re~ Mll~ k~ ~nd bc~i!l ~(l~]njnlk HlrnlS of m. C'!l~oon pic. w~ in H'tlhissceli111111'io \,ven~to he i:l1I~p.~e:lll1eml;~ed. bilizeJ .. ,(iii) ]I '\'I.f()uid 'pmvide:;I m~dUI11~~I~lll h<lV'e lmo ch,m.ce to S'loJ) the I.+s:e iml,<'Itm()lSo .. 1 will Ihlii~horllyisSUi!.f of (,;ef~,p~n~li on ph~ricCO~. n!1di[M~h Iy un:l~ Ut~d'UWt"~"1Inbi~lkm uf b)1 "Iihi ~l~ kl.oJl~ p~~:st ,COl: e'lllis:sio:~!\ by l'icher Il<Ili,o:u~s im::lFeillsed e ffi.ciency in ,en:lerID" lise. 'illllplemell~ &ld~mIilOO!l andl No~1)S .~ nit ion 0 f Iton-ii'o&,>i I flill.~~, energy :[;(lUI."C(!:tl. ,uli.<iJl, ~ol,llld~e l1:t!goli<lted, q i, i<. $,. lMklll~t H.·j. ilOdl:, Po Grlttil~, im Pt«~ding.~ ofrhtt Whi.le lhere is 110 .CIUe:5lon l~,nl CO~ C<1.pi '.Q capture un:02 produced imiemIl gm:!ific3lion ... 24111 lmtl'lffllMool ({}fl:/wmtf ,~1'1Cal'll !j.riNlIl'JiM & Frm h~~r[\o11f'I1~.C' i!llalfl.{JIJ;ph~re.~,doaJblc, theco~t is i .~ p~.[IlliS 'i;\i'ou](ili be capilhlc of mecl.ing the s'YI'le.m~. ~, S;jI!;!!;I:~~d.. ~I (C~.lT~~hml;l~~~.1'\';50Ntiof1, aeallWat9.I~L lW11), lIP, :e95-:996. $!i II uulkncl'\"!'ll!. C'apt~n:e: \!''Q~I,1d, be ~~=ft)iiid'ible iJ ~ 2- ~ ~rnaJl~~. (al1Iid, G. !iI~!, ii!.rndIK, :5. l<!ciro~rb '!l~CJiJ5· it C'~u~d the pri.cc or fos1>U fl,ld '~Il~rgy to ':~ lihif';J!IIlthllr i~, Ihe b<lmont"Doh~lY Earlh Qb~fl'!'G<t~1iYof "" ~11!!1'1~. at ~n;crlL\ase by ]0 ~J() JG%., Bo,:\'Ife\.'€'f. <ll<'lr~~ FnlC0: C(lhJm~iil tlm,i'li'er~['~.y, IPiliH~dIlS, HV 109M-SO(!(), USA. if E-Dl1111D: bro!l{ik@r@ldll'IM01liJl1Ililifllii1ld'u tl.Orl Oflhc::'opermung costwoukllbe Ihr~1la.e IJUIr= to which we
:~I~~O~"" 1~.C1!('Oz CO~UClll
(it u fC~fb(lll p~r y~m'. lFm'thel', lbis rQUIr~(H)ne :I<III;iowill 0:I.1ly c:hflngc slow~I}1in tin: comil1g decades, Hence. ~ ''I.'e se! Ol,desi FJbl.t: upper I iffill~l 01'1,!jheeXl~I'II~. r
~iO!i!•. \1!..1;:PlItlW (;n'ilit about ~
or the
sions lh:1llll:idl mllioJ.1s. ],he:-sOOJ.l~J' (ll1IdbUified. such an i1g,mem,elllW<lS ~Ul~hli~O ferce, the bet.ter~]m.e situation would be for these uar ious, Umi I Db/isis done, the size om~c careen pie will ,c,uD1.timu::' W shrililikil' a r.clteof 70 to SOOl perrd~~mj~. B~C<lllli\e CO2 sales would ~~n'~ l1nl)' as a t.emllOliilu)' S~OI)g:"I~',C;<I'~11H1e .oO~I1i,'OliiI,1 the of .'lm~~pliei'i6 \\,'Oillld·b-t: r'i;wess,ny. c6'J c"p~~ite fli"m:tll' the a~'Ii1.ulsphere:is reasiMe. bu! 11<:yet tn ..s
IH.~p'Q~iIiIE!~ca~ s(:el1alri~ :Ellrruise bY' ~ich liI\atfon!i~fth,e:ir 15 (H:illt wedg,e of tbe !Cat~I!i®1l pi'e".1\s "me passes, the ~)lae1:~01 rlo~~i~·ruel tll.lrniln.g over tlh ~ (I~ nishin91P~'lIDliS6ib~~ cmi,s~ron Umilll'liU Ii~~ly mi growl. re1li,uliring'an im:m<lrSI! in lhe arneenr 01 CO2 W, be cOIptmed
ror
SC;II~Nelc. a L 31S v
9 MAIRCH ,2007
1371
PHYSICS
Allilimpm'll,eli imag]n:g techniqlle reve ...~s,UHlit ,!ilenr~!Ils hn !l ma~erii31l calli !ltJH~r griidl.or:k tike Gusilli i! dly" 'Vl1i:!i milY help rle1ie,indl!r~ Ii.Irltlerst~!1d~l'Ie' mechalli'S.m by which (11Hrents now \vittllll!J l re~~r;ti3irll~ein slI.!'lermrn,d seters,
ItI:6td~,!dl;,
<'Ifi!::\!(~hli:k~n
I);;\~
QWU1~ln1l1mechanics rules in, d1is micrescopic \m:rktm:ld researchers assumed lll}iU Ihe smear~
jl1l,£ ..Ud ave:r.lging I efTect:s of the quantum
mu~ioll1s wUlurd reuder this b~h~li'!;'il'u ~,,(:~~d~ 'IliSly ~imrle. ~n S't~ld),'ili1g hi:y,h-tmvasitroll iemp~I'<ltm'"e (I:~) SlIp>t':':li'>t::OIUIHd~V i~y i:~, oppclr c oxides, ~.n'\1I.'ev.l:!lil:; physicisl:S fmmd i11!SK'!ld~1~aB:
!Ili~ cIClC'U'On ::>y1!:tell~5 ~1lIthese ~nHtC'l"i;als were
~x;~-mt~!!)l1!fl:~~yt,:(lmplil;"x,. o.m! p~gil,.~ 1380 ~)fdl~ s issue. KQhs;<1,ka ef ~r.l f ),presem .1:111 exper,im~ffil,~ ~o:~~. b!1ii1)klh~~1J~1~.$t~dJyin,g th~ ~1~;\~troUll !iYl"~'jll on the surface o~:'c::oppcel' ox ide Sllpe~1CO:1n.dLLb~Lli' by means of 5c."lI'miny fu:m~eling Sp.6'"~fOOOI)PY; By cleved)! er>;plo.ili llig~lle e Ir~l1SI(lf the declo i.l"O.[i! lnrcractiens, tll":;Y rII1m.l:!lgt:to probe t!~c 'e~,eclf-O:IIImime di:rectly. TU,1ey d iseover a wtl:rld ofa~n~zil,griehness .. J,htlp(,j bli' [h~ quantum mo Iiens ot ~he elecnon s forlill iug complex
spli[lia~ :p;2IHern.'>.
A main dTt;l;:~gfslitul\g i~'[e!'m;l1,{lilS i~the dm;s[ml,\vol'ld is 'uidlk.nlO\.l;!1}~o3l~ybodly l'iv~ ilmgin ,I mCilm~'l(),~~.I']1l Whenlhe density .Jr;e<l; of cars becemea ~('IO~lig~l d~.:tr.ingrli!1:sh hour, tile u"Ilffic CO I'IW!;. H) a S;~~l!mds;liI, Th.~ S;i'lI'!1!? ph~ ... I
ml:!:UeilOIli slumld I:J.GClU with electrons ,un ~oHds. blul [he ""\:irdbl(1s...,, f ,qt~"'llum phy.~k"s, o interferes, E~ecmrons shcmh:l eNie{;[I~:eoOl]!i~mall (;~t!:i!ml~m:tJlnotinf~:\ ~ml!he~ are $() v i~d!t;;mil!'! .• l"OIwe!lui:tlmll! metals and slJlpertiOm]~Lc:to;n.;~1iaOill Il'Dc c:Aeels of'tlincintc.~r'JelioUil~9r,(! \\c'l'ls,itedllw:!)1. hll~~1'1 roeg;Jrd, the electron !\.ystenls Ibund :illl is
TraUic: IpattE!Ms. Scannilllg nlil'lllu~iings:pe(.t.nm:oIlY 'UiD dilrec:llyp~c)b(1lliil! ~palijal. d~!illiitlillJl'tion:\1 ir~he<e~ed:r,on o lIiI()tion~ ~:f'lI.h~ooPiP~~ 'CJ}:irie~d¥E!rs1:11 hig 11·ftSiU peii'(lOn,rh~(lor. Tile pauerll1is. [am be'I;1iewed as at qlual1llU a mana· l(lgl al :Sl<IJh)mj"go, lr~lfi(. ~~:U-ll!rg'}lli!l:.jl1!g illito (In:~-diim~"$io!l~! hi:ghw~s II"'heril'H\~ elcctrolill,rolHr.c nD'i'II~ r,ei[alti!U!!lyh~el:f, s~pllri!ted by sewfl"ly jalimnedi areas. wG:llians 01 (OIlP~r.~ nd a)!'Ijg!HIi'ltom'5 atreshown belew tnl' imm:ge or elemcm flow.
Kohs;Elk<li etal. claim to ~~mvcs.ccollllP:~ished. HtJ\lli dc, Koh~11.k0\ ~·t(Jt.mOlU1<lgehi r:~b~ these electron SInWt ures? n1!~Y use .l<C<I'Irn'lIirillg hig'~.Tt S'I,.I1~¢rcO.UII.d.I;i~1)on; ~ll(·{'I~Pli()~il'ii, :rhl~ ~U·.,; wmt!tli~,g NI'~t~tlP)', a T~~.1l'iiiqw;: I.h'f[t m~9te the :s~rona pclh::111 rials exerted an, the e leesures the <n LJiml:UlIU mec 1.1:anic.II,'JlI'o'bJabi I ity or ydd~n!1: nr~UQ.~o';ling an ~1C;j:el!'o!~ ~ $pl;ldfi(: !,'it [OOllS by tlllC crystal. lattice of (;OPpcr. oxide I:m:aliolill <It 11given energy. In stro.Mgly :Unl;em.cl~ pl~~Ili~:i"Ih,~qlli!l_mmn motions are hindered w ,~ degme thilt rhe eleenen trd,l:nt'! :n~Jghleven get ing eleemm systems like those oftliu:: hig11-r~ con\lp.~ete~y jamrued. Copper o~",i.dt:s in I:heir (3). There <Ire imLdi:c;.l.Ii(lns.lhfl~~mley occur in :ompeJco:lldw';~{)f$. this wmb"b:i~ily, i::; VC<J)i herd prislinu: Slial.Care d'n.Lj1~ hlSU~:lIWI"5 i:1ndl in orde~r gtlod supercenduerers as I~n:ctmlt:1ng p:fl!ttc:r:n~.,> to inl:tr::rprel, KohSo.'1h et aI, 11~~.vcmmdln "''flY f :.n'OlJl,m:l ttlis dilfjcl.!hy. based on <L clever blJlt ~ to tumthem inro (s.upcrl~ol:1uJil~~IOP>. lfii~s (3, 4,. ,,\....ile in some cup.r~l~es 1:lley :JCfWl<IUy one oom!t to ~ standstill .. likely due l!oil.lllP'Crr[x::~ si.[[~1l1cidea of Anld~J·s.nn and O~g (6) th~l ~ ~"O I,'emove eJectro:ll,~ b~ chemiesl cl.opiJ~g (tlil<!lt L;; x ploi~s the 1;)'f1i;ct:;. ohl~ S'Wofig ~I,,'!l;rmrl·~h:~- Q ~h.'m=-fn nht.;crystal ]:l~~iec t~Ollvin~, I.:;lfl;~lsimi an is.ml~caddition ofimpmHy I,):I~m~m$·l' . ~ The electron motions i:l1I, these dopoed il1311"~o mad\";,'OL"k on oIilLrhigllv.~lys.l"hese :fl"nil.: ueu ilm:e"'''1.1.mm;.1'olIndersl.nlld. dte esseuceef ~. l"UpmHol:S ean bs Vi~t'l'~~l~ qm!llt i;.til,."'t! $'t(lr,""Mm!(.I~ SlfUpl'l!-i. ureclsarly ebservedjn bt)'[h aeurrenlh~ idJi;l,L ene jlQSt n~'(Idt' fflj)~\""!yt'Xll~·~!~~t'~: To ~. __ ' seaueriag 0) '.IIldi rasoaaat X-l"ay-scallel'il1g Int'\l'ge from ~:~.e<!mp o.ll, ~~.rn'ee:VI'-1.jI r \II;,'.ulh S[OP- ~ go, Ira Ific ,. Stop-.ailld-go traffic in our 'illlnr!d tends. 10 d~'VdQiP C'ompk:x ro!Hecl.i~ 1)lIiJt¢m!l, !md,~.~otml"fu!creqt~i.re:~ patuen:ce, whereas it is 21 '!l ~x.pcr,~m<C'ni;l;P), b\lt ll1es~~.:o;iP'~ri.l:Ji1e~ts .only pick up aVel"O\ge properties oflh,e $~Tipe~" relief to "'U)Ol an exir s:i!glIl,. fhe 5<!m,e dl:eCl ; Rlcs~~rdn:;rn\iI.'Ould I]kc to ".iew 't]~~mil1!1i'a1 OCCtI!.'!i !i~Ih~ Sl!op-;lnd~~o·~ fcc~oonsyst~~l; his ~ i1be '~IiI~or ls~n ~he Depa rtm~mt or Phv~'ks, Uni\l'erii~ty sp.ace {m; opposed to' the f~c:u'~)n)c:al ~pac,e nm.ch e<l!:>:Lem:tn [~llIJiO\'eC.h:CI:I1C:mS,th<al1l,to.uld ~ or ltid~fiI" Leidep2~33. Ihe INe'therla!'!d:~. E-iTI1:aU: (}f di ff~c lioUQ). .:I!ld this is e:»t~ct~y \V~oYf them" To ootflin lhe!)ver.;l!~] plI"Obflll:liIiI]' ofeidu~r {) i:ill1@IB,ngmtz,:lfiDe'nutl'liv.l'li
<1.11di
ro:m.elhj rig sil1:~:i.I!:~1i h:appf:'llcS~vil;I~. eleet~~u:: fb~lIldll~<ll elec~TOllilis mov,~n~11[1Q[In,cI illl the eQPper exide planes h;ndm ~rni.jl,g.ot; dl~'!f!i~~dv,~::>~ onei dim.ensianal "h~gTm\""I)"S··~~~;let:e niltYl1'emthet Lhey 1i~1t;,~ly. s~rrut~!tr]~ by i~t.;uhltillg d(l!mtins (.2'). <I ItCI inrecent years e~periffil,1.e'[I~<1~ Sl.IIpJKJU 'I~'3S licumd for I.~.e Cl!::d:stcm:;e ofsuc·h :st:rl,pe l:l~tI.>eI.~lmlS
~ro~l ~.1~.1 ca~t~Uhll~t'Il:-;" rl1y;~h: iSIS
9 MARCH 2007
VOL. 31 5
SCII!:.NCIC
www.,sdencema9.o~9
IPERSPECTIVES
remov~ng ot i:1I!dJd ing an elecuon. one has to glassy panern
(J).
II
CUfJTcm
for a
iIlustrntillmg~liDe sensitivn.ly of
A~ ~\)hn\v~ i~U~~ngijw. the br;l.ghl!lmis art where Ih~ electrons move aroundrelarively freely.al1d the dlll'k.lI"e,gJtonsare W.liIcrc·lhee:l~Cl"'-. nUll tr<'llft·:~; jammed, The i~mlge 11;,t~atemicis scale n;,'S!o:bif)ll.~U:d it is elcsr I:h~t Ilhe lfIIon.ilc;: el:ec~ru~:shH\le a pJ:'ielbre~loe for oxygen ove.r copper. Zeomiag nut to Ihe n~lUometel" scale, (ln~ d~~!rly disC4;;~'[JS the SltriP€'$,. the "rivers uf charge" st::p'll'3tIJd by inSild,ni:lmg regions, wh.ih::' 0111 '111i ew~ j1myerfidd of\,je\v t~&e H!I1m. inl.o 01
mJ,~) IiIIlld lU!"a ~'~llalDiwIyc~~I],,~~d Vlilj~ctinlgL;:k."'llip ~:rn[l$l over <I sizable voltage f"ollligeata :givi;;'1l lue~l!jun,.and by Ulkh~~ther<uio (lihl~~~ q~~rui!i" ~it'ies one obt<llil1:> a Illna~, Ih.e dec ~ml~ .of tll.'l1rfUc..
m~ld'n.fir]Qlr,egoing euin l1'n.cse hlla~f'Dh~Ul~SO)lp~~m:at liI!·i.rst glanee, ~mllhL;: llinill'ion dolhrr qne!l.l:iOirii is \\~1(!'~~er thesesubfM.iGS.rew$1 ~rnylhi!~§fd)(lmlh~ (1rj"gi!1Jof~u(l~rcQm;lnc:tuvi~y .n i:lig,lm Tc' Tlris is s!:iU as [Im.uch:1 ~~y.<lW'ry~!S il\io'".u; 21 Y~~~i,gif._J\i"I~l!;lnit W<l-~ dis-
.m ill'Llsi{m.~n by omnbin~ng the presem tech" ntque w~lh the energy·resolved ~Pt."CtroNC{Jpic i.nWor,mu:tioJl.:~l should be posSJ.ib~.e tngeneraje ha:m l·~'Vid.(:'I1~C' ~ n:OI1ility Of't!lC~ pai;n;, r~:
Ri@f'!Ilirtl1l(!I!S;
covered ,t/. i~~, It i~believed th!!!;:, as in normal sl.Ipe.room:lllCl('u"S. the e~cGtroll:S b:it:l'!l,dl I)~ i ts in ~h~n 8!ib:;;!;1CI!!e,lllllyII [I&'\I:,go Bose con.Jem"lli()~,l 'U
the SU~roOlldHctivily ..~~mJlLgil'[\1I'~11 be thfll ~he:IU1<Iges, obl'1.~f1e:d by KohsakH .:1 al .. are ~]bom electron pai rs omninl;! to a ~)arli ;11 ~tand~~i in!''ll'i1!d (If rhe (l1~t:ln1l'iI$ th""m~1 '!.",s. II, Whe~l .~SA.ill'i;!',I!,rill}, my trained eyes at tile illlil~'1ges I Sllf;; II~se pain>, e\ 'e1}'l",1n:re. This mighl wold~be
dlilll: 'C<:IU":ies
]. 't: ~hl.i'ib .Nrl~'.,ScNMte :li:l.!I. 1:38C11(6007}; ~bli!hed p ,gnLillle(I l~etlrua'1f ~(10t,lUO.'llU6ISctena'113 8!!M~, 2, J. Za.:JI\II!I1', &Jniil~ o, t'hyt, ~ 8 ~!o,1.~91l19a,9j\ :3•• \t.A. ~eli~l'Ie.1g1.,Itf'r. ~.O'd. Ph:~. '5 ...nOil GliOO~~. oil. 1. Z~Iilen. o!i!·~1trfE'lI~!~.ina (lOO6t ~" 1?,.AbbinnDme.~!Il'I..i!tcllllrePhY1. ~1,:]s·5 I(aoo~. . ,. R w. J!!t!dmoJl, IN. ~.IOJ1!i1, f. Ph~ '~em..~ 67', 1 f2006~. ,,~p!irllli fNhlR'dn I1lgli1~.~:ijpi!\I\oo"di!Jrlh<it:l" Na!~ Phy.._ z, '][33 HoIjilEh lOO,~t I. ~pl!'d~1R_r,j: Hi9M~rnJl4!tilUjR'Su PI!KQMudivil!j 1I',urn!l Sd.ef.!fe J14, 10; 2-])0\19 t:!OO6). 20.
d~..scrn,lcryOl!ldeHnical development. of penid,~lin I. ushered i~l the modern j_ll1i~ubi!ol~eera iii1d St111'ntlhHOO th.e diS;;::;O\I'f:I")I oftlle,mtibiotLcs iI], cur-
'Th~'
~U
years
P [IIJPlII1iMJ [ 5 jl,i,(l
l'E!)I\on~ pe:l!l'ic:iUims" B<3{.tertl!l U,!ie! 01 pep:~idlog ly(.ai11 Il.aVeo~ pr:Ml!LLM1,em· to Sllll'!l€s11r()1liI [)SmotijOlr~ss~S:i'lfith~is or til~s lay,ell" precee ds inISf:lf,~,roilltile ps, First,llii~idIIl~ ~Yl'llhesT~~d cdl. il'llhe It ils~l1e!l~r.j!n~ll!lf~d ttl lhll' 'ol.l!isldie.
resistance w
aflcr lilL(:ilr dus-cm't'ry. pellf:cillim: and !:Chuled fmt lbi()~ks (CiJ)I~eclive:~}1 ealled ~~act<lli1,ls) rema ill dlinl:m~~y us~fliLNI;;"'ie:n~~iot,;'s;s, 1.1l~f,,,,ma!'k~ able ;lobi:i~ly et'bacteria to develop
p,·l actam
and ()!~r
.ullliibi.oli:s meOlns~hOl.I. [here is ~II L'JOnli~'II,I(;illlicC1!dl for !.1~\.\i'inllibio[ic t.m:sels. m.~.dne~,v 'J]J:timitrahi,.d a~~!m'ts. 11 ~g~ 1.402 oU~is i~1,Ic. 0 L(lV~fing"'l(~Ll)!1;;pprl'~~ (1)'\,ud ~
~ln:1uctm'e01:'<1 bitllmciiomI bacl>e~H]1
Jilr
biirlilil(Ii:j)naltil"illiIs;91.y.:.ois,y1a"i(Hran~· peptidase. The ~iructlilr~5h!l)utllil iI~d de~!opmlln~. (!if rM:!w anliloiolic5 M~get· ii'i9ti1~ It-afl'i~I~~oos,ylati.(1f'i d()m.;ilil1.
~ll~Ul~bn!!IITL\l p;AA~in. dm~pru\'i.tf~:-iun·~'t sites 11(11 bfl:<!kcl:Qwn I[ fur ~ltml~ple. ) Io:llib:lli: pi!ni~itlll of U OJ,lly fau [).1.~lrt~ms, ut alsofi'ol: 1lit.",\:~1 31Uibi:o!I.(;S, b lUn.C cell w~r~ll during cell d.ivisionl [i1:erel'are
I?elliidJ~~nand o~hef ~ ·~W:::UI L~.~. llllll·ibiol: ics ti1~et" several bacterial enz.y,une:s, collecti\lely termed peuidllin~bindin.~ pm~eim. PBPs). PH Pi> are 'lI1IecesS<l~-y the g rnwth anClI1~3i.n:~ for ~an'l"e t,nh~ pepLid'QoJ~YC<i!lll~y.;r.\:\/I'lic!ll ftIJIViI,~ 1];".1"1of the baeterial ·c:e~,~ alll :~nd I~J!Olec:Jtl> w th:e cdJllrom o.!miiLol.tC stresll. Inhib.ition Orpel'ti®~ giYC,Hl b]os)l1l1~he5is ;Jl:id of its cmltlfolled
I~
.~~'Il:
Syl~Il<lIlSI'a:3S.es: assist i~,gl.nll aug ~.I. C.-ItQ Iy.le the fOflllHl.lkm of peptide km.d~~ between po'Iiy1:uelJ" inh ibilS (;ellgro\'\.'~:~l,Because .~ ~epeptidogly~ ~tl·;iUldl~. tlumhy m8kin.~ dIu:: 'wall more r~.!i!i,d, calli polymer is, lIbiqu~~('ms al1:d.e~...er:u.iilil tebaeThese l<~sks<IX perlanned by hifllnct~mml terial life. its ~lssemM.y .~nd m,Iilliltot:mniLce mit: ~UZY~leslh;ll conHlilil g)yc('l$yll:l1r!lsf~n1~ ~nd 1:~.JnJsJJept~dJse emains: the lilluer;:;]l~' sellsilli;ve'; d t~1[gets l:Orm~"'lY~]ntub~olilcs (2}. The P~Il~jd(lg~YC'f!~l censlsts nf a oockl'loOlJllil 1.0 ~bLC'lams. clh,uin of l:epe3.t~ng t\\'O-smgall' nnli ts (l",II~l:ed P!'fvinu:!i. $.I:l~di~ lfl"the ~·i['J.,~~ep~ orp~pl.iNAG ~~mlNAM)I ;u~d ap~~ll,,~p~plide dl11in Ihios:lic;luhcs.ig lul'l"CtCrlll.h:d to ~iOi(;l~S. bmll'ld ~o cadiil NAM (:see l!l.e ifi,glufle). llu~ 01:1 t1~e trnllspep,ti d;,;;se st,i,g;e of m.semb~y. ~lmll'le'v'Cr, th.e~llh ibilioll of u:a:l1.spep.t id3..;e NACl-NAM-·pcnt'lf.K:lPtidc m.1i"i1! ~·~Ji~,k:d ~.ipid]1) is s}1IiLtll;aesi,~oo ilmdle oe:lil <I1,lJ.Ie'thered to ~1,le\.'e'u .<Iclivily by peDiliciUinsis only halh~,e story. !l1cmbx,mc by tI, lipid! Uilker. Lipid U 1:sIhel1 ~IIUCI tbegb"Co.s,yllram;:femse"ctUv~,ty of the In:m:lsfeJlied :h)J,ll~ilil.e hl::>idle olil'~e cell to the ill fil!m.,;~kmf!1 enz.ym~ ]s ~n ... :(~~lem nl ~e~llxl1' X flll;~ii~d~.wh~Trc nl(;mbrm~ll;':-fussoci,f~wd glyoo· IIm.ede\'el()pl.m::~u. ornew ilntibiotUcs (3.4).
on!I:o [h~ pdJlyml;'l;, T.ru~.I;p1.jp,tida:s~
I.:lli.):gI~'r::[Lrii.
SC;II~Nelc.va L 31 S
9 MAIRCH ,2007
1373
PERSPECTIVES
Th~ 'bif1;il!lilCtinnril ~m~y:11l~J;1'l> i!1ld~td~PBP'.~'b Esciterichit[ c-o# iQlIldP BPi from Lovering,c:l
"nle~l~~~'-diin~~nsi(lgl,<l~ srrucmre
(If
PBP2
rl10m
tl1ci.r impor. t,~m::e~o b~lcl~ru~1pllysiolagy;tncl dliLlY disclQ\i~ cry, tlhey have ,[Csis.tcd de'Wtih,;d s~1Jcl:y,y.lil'llinly I
because lhefle ~"lr~ membrane ~J:rtut'ins <Iredif .. ricu~1 lO IH:JJ~fy.~~s.r~y. anti erysw'l~.i".1;: Lnv~rj i\g 1:1 ~l. 'h<l\!~not\!' evereume [h~-~e ahstm::les and ~,·epoil,1 the: :r.;~";. lhree-dinlelml,~ s~ ~i(lnill :titI\Pi;;l1~N QfPBP2, A7;jthe ",\~t~l(jr$ ojrll p out,t'his stmc~,ll!reisreg~:dled as the hall}' gran,1
(.;n
Im.er,ero:re In()v:~des <U15U'ers 10 long~:snU'ii.ding (W.csli.o]ls ,o:Fbacrcri ~IIb~odlC'ID].isl.ry. physio~~ ~nhilJiwt Hla~ is mostlyusedin ,anim,,~ heOlld,l ogy. i!][!l.d 'lrnt~bjmiC' aC'liVli,ty.Wh,d'sll1flXI:! (4). jViOCIlOIl'l)lci[l h. one or~ \'C'r)1 feW!l~litub~.~ \VC:lIre clC';I~y n~n(~)~ .orne\\! 1lllililliottcs w otnoi..".'i du!itovered~:lim:s Ulr th.n tm:gell:he y;~yoo~ evereome the j]l.l)hte~nof anribimic resisranee d syh rn~}l r~'~l~' reaet ifl[! in ~.I~ I.idugJ'yt:~U1 m'iid th~ ("IUL::,!;.CIl'\}C of ItCW infcCIJi.ol~s, i~m>('S
IJhy.sioICl~is>~. As plb':(lieood the structure is d,uvidlCd inlo t\\'O $e,lXU-Jte. lobes conaeeeed by a linker.,J"he C-~e:im~ml] IT'J!1!$pe:pl'h:l'1S~d;[li!Ii!~ain ~$ po .. !;iHgn~-d O~ [he Q~~~J!;,id~'lite eel l.where h '~0li1J of ,~'{e:CIt.~ijve~nl.eu<IC~ ion~ W[LbJ auumber Q,f"~U'liil,il'O find ilS sub~nItc (the ]:lCIlMlpepl.i.'Ltc dl<"1.in:il.J. ro:r a Th is is 'I~ \,!,Iilere t~1;l ~lyclropl1 ilit: ~-Iat::l'lm ru::iu.'l.mel .~c!':OtUli~illg i~:-;~igDl[ ffinity for so !!illilih(ioIIeS call1l~llil.crnN 'with the ,enzynm,e and the enzymes. The lipid tail of the 3Jnl.ib.uo~.~t' block its lilllct~ml. ,hm, cnlllrnst 'tlhe N-\~rmilli1j!J,1 ~.i kely directs t~e\li,,..Hhet1Jd of I.heallll~,bi.o~ic U) the membrane for imera!:l;iolll with the target tmnsglyQosylase domain imern'cts with the ~u~!"rib.mf!!1;i \';hof;:!1:.~~lfllirl'idll~lbti1r.Jt~i,"" ~o(,;o;m:d, g~,ycllSyltW'."!f!sfern.~4;i..ll!:~ nrill>i!llf~ti(ln U!rl'~~ inThe S~IIFUt'~mre orlhi S OOJ,nOl in does notre-semble hihimr ~n~in.ee.rlzyme active SiH: <II~so ans ...ers o~~erk ~O'!i,\~n g~~!~iy~tr.m5fuoo~".S. R;nh~f. it is, the imp(lll1,am 'lil~e$tiu!1ol·'I,,\·h~dH~r lipid II is relsredto bm;:~criop.bflge i lysozyme. which, the donor OJ the acceptor in ,glycosyl bond eleaves the pCf)lidlogl.ycnn glyoosy~ balld~lm form."Ition" "[1\14:;structure ic5o]!lIJl oonsl;Slcm II,iI!Ve~ereaetian of sym:l:u~sis(5), ..VULt! the laner,
bkl~y!Uhe:>l~ Th6 new~~b~~ri,i!uju!~spft"lvidt; ,t} 'IIi). A !'rmj~f !;;h,.~l~!lge the de\!d.{)pm~nl of jill sl<lirtillg pain! to engineer lill,is "mlibiotVc: fil,r new .'Inli.bi.O'~ic agents :~sthe <I.bility ~o icl.enl.ii}' use in l!r~iJdng~lqin~,m if]f~'t,;tkm.The gIYI."()~)I~- n~\I!i tm:~~;o; <ll1id t~~ cPt.."'<iI!i"'~ly i;,'vlffitil1ltQe '~if;_) e';oq:)~lbit transferase ICilcl:uml is iElpl1omig:ung<llldbiotlc ~heC'>oll~eClionflll.\OVCn enes, ~fui,e S~mC~lll'e of o I;'~~l". Llh i.nQel1.o!L~,~ydnI (lOO' lil!il1Le~as pote~n w fBP2 provides .m {)prorl~~nily ~o :we\'i!i<i~; .~n "ol.d" targe~ wirhnew ilClol::..Thisis a br~lk'IS· v;;m(.."Omycin, .u £~yc'ollep!~.de "'Dli~ib.ioli.c thar also mrg(:cs peptido~~yc.mii~ymnthesbL IfuimuMfui. ill 11n!~'I!dIJ1:3>t has been b0rclIli: of I.~.a b~los.Y,Im~ The structare of the il:O:l1iilP~,exreveals ~h'lt means to fllJly exptorepep!idog~yca[l lh~ glycosyh~Ln~f~r;l:s;e j1l'w closes dn'W!l on ~~I:;~i~ !n'[)d~,.!'1! with S'tiPi~[Gg~~"in d!iug, d~sc{1Ii.'Gry. rhe SlUScc1r "warhead" >llf[h~ ~1imiLa.iotic.lbrnmlill:;
R!~r~rl'l1lC~, :1. A. t, ILc\!oi!iflmg, L ID~ (!SilO., ~. 'llm, N. v...~J)'d\'lI!lilfl:a. ) SaM(f 3:.1.51 MOl f2007t .2• .(. T" 1liIilt~h"AIn.lib;"!'c:r;A!rl1o~ Or;~~ ~~JleM(!' (ASM. I?r~~. W3:!ii1ingtclII. 2003)" II)C., 3. J. HaUid~~. Mi;:~!i!~:f, ![ MAili.dOo!1'. ~ 1I:~!!rat!l[!!lI,. \11. MeiJ!~i\!j;i!nl~ B~ P'hllf_L '1, 9):1 t~illOM, ,~. ~. 10s!a5t!, ;5,. WiI'I~~f. 0piI;!. (frem. 8M. '9. ¢~9 ClI/f.
5.
~,OO)},
IP.II)~iltl,~J MtI1.B;~~~:j!6.,:n!l,1
~I"
ClcL,'l LHOLQGY
t~<ll mitos;is'jl\,1i'1Iy~lbll()\.vs
uring the normal Qt':.11 cycle,. DNA rer'lic<Ltlo.ru is comp~e·t6dbertlte t~eceH divides. ,(m~[osb; l,~ow tl1~eell en."~lrt',S
called
DNAreplic-.niam
ill
~nd
the 5~1,lJjecl of
<I
~'SImly by Torres-Rosell
~'nIE~ugUlil:l<~}~ D'~!g,,;: l ~)f~hi=-ii~~!~ (I). ~4~ I:l'IVestig.ahl'l's have :~ong k:nmAfl] that cellu lar
c~.mlrol~ C'tll!i.ldcl1~ikpoi ms ~Lt:tas ij,m:veilbl!illCl: nU'lcha:nls,mm; m guar.'tllil:et::' I:h,,][ ~.,d'D.cvt:'nl dtl,r~ ing the: reU cyde is oomplct(;dheriQrel~.c 1lC:::~1 mle beg:~ns U-n. But dle tillldi~lgs()i1·To.lr,res~ Ro~1I {;'J (II $~l!m~_"[ ~l'U I:~ds;is; ~Ujt al~,~~ys~l~ C<I:se, 31lld pose ~h..e her~li(.<tlptl1j.sib:il~ Ihal the ity cd! !T!;):!J,I b~ bUml to \"'h'Cd~~~ i~ .real.ly dom.)' is
the mi t~tic ~):und~,e. DNA.. &~m<lge checkp'Oilli~s de~ec~br-oken cil;mmOSO~lles.Wh.en 3cl.iva'led ~ith1;r !1;~~d"''IVirublQcks mitu~i:-;,,!li.~nt'ing lime for errors to be eerreeted. Thus, these c~1eckpO!in ts ~Jl :i.l~r~ lh~ll mi't1J,~i~ to! kespl aee
ml~Y \\illil.cll (ll~womQoolfii]ii;!i <lfC' h~I~Cl ilUil.dPJ)()P~ erly a£''>Clllb~.ed Orill lfui.c $p:~mlilc..:"
repl.icailiortl Ibrk. of the manly tll131 lire formed ( 6). :11l.~ighl IIIis!'" Il:lk.cn~y ~len;i;f;~hl~c i.~111L11~ cell ,cycle. liP to 1~\: lime o:fmim:~i.s [DNA lrei)llicOl~liJjll oCC'illrs.d~lr.~nS S, phase, but l3tel:'ep]_ic.'1t~.on IlJIlught occur at tiil.c
1
end ofthe
!tIre'}],
SlillCh
a. "[)iNA «:'plic<ltiOl.l--(;01lfiipleL~on ch~t'kpoim'" ,is,l;.iOIl:-5ide'Jfl::d urli'1.L!~ bectl~Q:-;~it \¥OuM. dl::le(;1 '''Illorm.,1 stnlctUl'oC p~~lmt al t!iLe \vrt:1ngtim~" iml(!C(L -r:orrcs~Ro5Clll et ell" (I iscm'~
2, ph(l~";: d"ll
normal rcp]±c~~i(ln fovk~ at [h~ wrong Um.;-_ ooordirtirucs Ih~C<}!mpI(~ljml of DNA ~'cpli~Jtjon \.\Juth t he onset of m i~m;i·s. Tllm.ewel~4;1U:J\~IUl Despite~J,l.e prli::Sel1ce ofll1e forb. celllsprooeed DNA~pli~tiotH:~~ckpoifU tril;::lgCfS '-Ilr.~h~yin i!i!~o,n~imsls...wilh CUll"~qH'I;!lUi.rd c~wflmC):;;f)m~ mi ~o!l:is llP[)11 del~ct~"g d"II1!'l.<:\~ed or St1l n.ed 1:GJ~~k.at,LOnli1!oc"DNA l'egUom,. \'i,.'here rcp.ti.<-"<l:~ ( lioll ]S I.a:ki~'ig pllilct') (5) (see lhe figuwe). I-~o~\'· ~veJ. ~.t l:Jta.y b~ qll~liC ~I di fl"ert,'iJlt ta,k lor tlte cell 1 ~Q deteHliI!ine ifllii,QIDmaln::pl ic.nio:lU, lQrkti h,iv'e
CO:I,llP leted~he.ir~l:l:sk8
ered
<Q.
~],lJUlm'llI;ye'dS~;cellmrll:
mis~gr~~.lJi~n.
c~lude
\v(fli1~d
p~dicl
be[<Il1~e
or the
www ..s-ciencema9'.o~g
IPERSPECTIVES
Their test '!WI),;'I1Irmde p![)6sffib~e by virtue Qfs]X.-'Ci<1if!::i~tlUres ~lf NA repl ication D
m05!:mu: XU.
II
al tlia.is locus, l[ach of Dfue 3SS genes ill the rDNf\~octu; ~u!il,tamm, rep:ilij.cat UO!il lodi;. ba:rrijcr a ~in:l,I(~ ''I.lli!Ch billdsa :~'lI.mlf'Un ,r.':!I~led Fob I ~7).
This $.il~ nf~!lI;",~e~inu is :~Qco:lru~d dnWllSI~]!U of the 11 ighJbt tnM1SJCribed 35S !'ON A gene <l~d I1':l)JforC~:!!ipoli!:ri t)1~~I'\~pbc~.,~io!:iJ ri\ movei ft'i 3mdooplk~llitnt ]~ thb 1'iIlt'icld. ac..;[:! e~1Iexits , me:~,t.;thm is, ~eplic'lti,tm forks r~plic<'l~ Qnly G I ~ gtt'l\lI.¥lh r'l~<l5e ofi~e cell cyd~,. ,~l the comi:l'Q, din::'ClUOIl l1!1'J5~);tTIll1cSier:ipli,0'1:I bm 11:01.ll d'n.c i IUilS 10 a specific limtHl: ~;or replicat~QfIi iUll S. ~h.e Opp05,~l;e dir:e:c~im~, IJhaM: .Ul.! for Ihe begiuJ:lil,l£, (Jif mitosis. ByI8~g~ng chr'OI'I'105tl'lmle XU with a. gvcen Evidem::c i~,budd,ing811d .I1.ssion yellSI:!; :undliI,' ~~(ne,Sl'e~t [ulei n,.~:h.e <.IlLlthors ohser\'edl!l~a~ p cates du:exi!'>tenoo ora tinr:li~.l£mec~,l.Hlis'llil th,il' ~he rDNA locus missegreg ... es in cells [ha~. S.I1I:I.<.l5 nd .'l11~ntelrll{;li n~~ protein, R3dl60, have r is independent of DNA repl icat iou, M m:lllI.l a ~)ill;.!'i!l ropliC;)J,~irm fQlmf' .~~.drepllc<ll~~ln-.;;~~'\(]:y~~~tcells th.u C.fnl~l(llini~i;u~ DNA replicaexpress mutanr formsef plrn~~j!l~ fbI uorl
conaern sterns I,n:ulill the usc onlle l'ep~lcali()Jl l'orll:bt~rmier ;i;]oc (lom~li:nirlig lecusendereie to ..... the rDNA regien, Because of ~.1\tCbarrier site, th~ 1fn1,i1:mlbc;:'r.a~l.J S1.~lJCn.lro ol'!'norm~] furksilllhL;: IwDNA fe!;io:n ,UI'I ~,uc6 IH'lm.mts,:lllhe lime of ~,iltlS~r:;. b: a~~o tlndl;aT. St"eo!ld. t~~B ()fMJ~,('6 U~L;l IIUnl<lntsto gencr<lte life repl i~<Itiolill is also of C\(ll11C'tlw.Wl~y '[l~ r~gili n ~pl~c:u~~ late ln i$ smro ]]liJl)tHJus.i~mlknml\,fl1. ami '~he[eis the :I!!i~ggillg possibility dU:1la DNA r,e:plic~lioUil-CfJxalp~:etion du:o;;:kpoint n:q:u ire~ Smd: mul SJIiI1\OO pmtclm,. R~c::m evud~1f!0C .in rhe nSS,10n yeast Sdlb()sacdu!~'Jmr.ry()t~S lJombe ::\uggc51:SI1mar
beeausethete ~ no DNA I:Clplic<!liOl~-CQi[I.~pies lion (:h0~'kp[)!~t lf'there is .~.oDNA ['ep]i!b'<Lli.o:~-cOlup1:er~('m cl~(;~cIt1?'Oin~. n~i~~ll eell ~n!lkL:; ~~irly ,~](¥~,~I a S1J1I:'!C. wuthom beil,1~ oel1<liiJl. thm mitosis [oUov.'slhe e01!ii.1.pl.cti.orl ef DNA r~:plkgljliu~'!Ol~l: id~3is th,U the oeU relies on innate rilling of mimsis
~I
~0iI,fIl1lflii!d
~'l~~
.!rtllt.,~ ,([J!i!dt{iOiJII
H!pI,i!tat,~ rilliik
~!l[~~q
-+
~mLi.~
[)iliA
I
G.l.
- - -- -'-_._.;:
- - - - - - - - - --)!o'
1140 rog~f!':!slon p
~1l1 milri~~
,"
\ •.• =. _ ~ _
~"I"
~1101'~~1!!i ,dmMkjiojn!
o rdel",. o!ro'e t ,@rci,e.li" (TOil'" I~en)flh~' d M~iol1 'Ycle aU.Q,w1i repUcaUoll of DNA befo re its ~gregati®fI (,en ~nw daILlg~1ile; cells. (11'£1111. right) The Se<iluemUI or (fli'enl5 is en$IUl~edIh~ d1>t~(ikp(liln,lS~hat sefil~~ IDN.Ain:te!mriity arnd' $tan~lI ~ r ~~lIIla9r&ll r'¢~ic~ti(),n $tr!!l(~u Iie~. {BOIUJORiI') ()I'(IN rnlay also IJ,e. elllSi1lrM by ~ D>rt4#i IrepUtiiltil)n,~ ·Th¢ cDlliiPle'tnoii ,t;l1ll'dkpoili!t (dashe~ Une). m>1liI}' c:om,1en:~I<l~e at the :ifDNA locus 1.'1').Those
ci''IItedc~,leckpoiLn
F;;o.<;sel
f:1
SigIilHlia:J@.
al do show
pf'Ol]d:elll far DNA dalll!Elge al1drep~.i,calion cltL;'C'kpgint~, BUllheg.,,;:nen'll!ily ()1·'~;I~l;.'ilrC(mdusl:gn~mllls~ be rempl;ired with their O'I:rml)'siii'i{If the' 100000li 9Jlildlmiln:~lIllS. dmt were needed to ask the qlle:stion. TJlcrc' isalse ~i~flm I suggcsrt.i ng LhiSi!a DNArep~iL(;,nioil-C{):.1111p,~et'ion l"heckpo~1li1 does
e.:dsl: (./,n, \fan: B.f'ah<llll et
lion 'e~ it G l' but do ,1iI.{Yt subl\e(~~Lellldy g:epHc::al:e ... I)NA~Uld still t1U~r rnito\S[s Ill>iril'1i."¢m a :r'e~ normal (en cyde i( 14 . .I 5). "f~.eceli~. thus r'f'I~,e:s en Ibe'~ im iu~g orONA ~'Pllk,lHmii ~nd milx]['.;!s IU ge:t ~'~job cl<~I~e. ~.nJrnif; is HrneCii~~. hm.lhe w cell c!oCSlffi'lknm'lllU~y nat hurl CIlOllgh I.or i.l to have emlvtld .1 meehanism that, <I!'~he present \\!f1l'k II rgue~, in :if9,cl dees not {!1(J~t
Rdl~Il!()fl'5
,(II .. constructeda
phase. CeU cycle k~net'ul;.~~i~~dic01ted I~I,U ~hi::; d.elayed entr}'imolnmiw:Ms .. B(II:h Tt~ITes-Roiseil
(!.~
l:l
"fhe d(!lxtl{~ ove~:~ [H'4J.\ :J!iep:lie,~t;ion-eQJ,nIs dli.s [he~'lKt \.\I'l)rd un I~e:if;;X~$tenc;e Il pk;ti.(1iL~ checkpoint of \"'i~1l'OIniml~., .nma umU DNA. IlCpl ~~21I.i IH:u.u:Jmlplel iO:lm, c hOC~!l()il1il?' O we~olved!. one ,C.lUll specu hue on the cons;eProb<lbly not T\~fOdel~]ils "utmut the elegouu. qlU~:Iit:eS of !tiOI ~!1Eu\ri['igsuch .. oo:t1ltrol, There is sy.'>1em ~lS!l~, TOl~OO,i;-R(MC~,i~I (fl. Inay ca~l~e the Cat.c nl.·~1~!Imlllillu1[l:!1 by fmgi~.~ sll~s.., ;!;pec'if:i,c ("(l!1!,~~rn ~,lmulllw ~~n~r,fl.~hy ofl~dr~()!lC'lI,ll- regitl,ns i:~G~uomu;;,()Ili11;;iS~'~'t f4;,l'J.~k..~L1ii mld I;;Lt~ s ion s. F.~r:st [t :LS lmcl.e,n: wlmt 1iii.·_',cti<:UlI of lhe tII,fiepl,une to bn:<lk in mitm.is 03). 'Pe~~,a])s DNA-rep~ iC'.uk]f~fbV'ks.~h;u p~~i~imo!'!,limsi~, Ir..-.,i Ie si~~s boc"ak b0C;;~UsedlSy (lom:rd~~oo['Una I re.semble "'~lOXl1lllI<lI"JreiPlii:atioll StlucuII:es.. llfh[s foik:s hlu,: ~11i G2, Cd.ls e.i1,~er ~lliltDt!>fs i:1nyw3y
w'I,IWiI.sde noem ago org
J.]:~n5·~m~ll e'!' ,~1" S(~,el;lS'lloliJU (:l;Ol!ln • t H. Hi'!1l1fe!l,T. A. Wel~ei!1l"Saen(1l 246..629 0. 9.8?). M, 6, t(I51i11l, J, Elii!m:~;I' N(J~~ 4:!lZ,~ll6 ~200>!t !D. ],. lew.. J:I.~, 8v.rt~e,APIW. ~. IGetlfl. ~7. as '1 ~2003~. :S" 11). llr:dfl.l4ri, M. iliDi!an'i, lapin. Cdl i51JQl. 17, S 6a ~:20a!l)" 6. M. K. FlII!ilIi!~ri!fI!~ el ai, SOIWCce' 2:914. lIS. '(20011. '.!t. bf1l,'b,erl, A. M.. ,C:'arl'. 8f«hil'l7if 87., 59'1 (~i~), 8. J. rtfrey,~me~.a N{1I. CrlltJi(J{, 7. ~n ~2001.5)" ,rill., 9. A, lG!iMlI,'T!. Hilfllno. G'fn~£Ie\!'. '19, 1:lM'12(105;I. 11.0. n. i!IIfril!M, 11.'lkllii~ml~OIIT( 1'!1~liIiID. i l{ontl, I'i, ~n[It~'!ililW~\ Mill. til!tl. 00l1. :26, 3o\1l(:![UJ6~ n. Ii:. AIIII,ilal2ii!ou, .,tllmmirot lilt J. Q'(tll!Jn,i!I~ ]. M• .Mmr!f, Mol Cgl. 8W1. 2',9~e7 ~.:i!OO6ll 1!L" A.!_ Blraban1. 'C ()_Ilu~h~a~ IL Ctl~rOOliei!UI 'I\t L f(lI1>!J!I1~!I, IB,J. !l1r,~~ .,tIlat Cdl7 705~200ln. U- M, f;, Mil.. Ii. I'll. C.a.1~~~,T_ w; fjllo~ ...(ylqSlEW~':" GeMme li'es. 1000.'2:~200,]L :M,. ].1'1, 1I'oyn, A. L.lohn~D!'I,l. H.l!!i!ll~~t~!'I, fie'!!. aWL MoL iI:5. 5,U2 ~U9s,), :115,. 11'. J. ~~. fl r.!L. C'f.:rJ:l'4, 3 n U99!ll),
:I., 2. 3. .:I.
Cit".
"'~~I
SC;II~Nelc. a L 31 S v
9 MAIRCH
,2007
1375
PERSPECT~V:ES
IMMUNOLOGY
Delectkm oil: replicating \lim~ il1v(lh,r~5.i3I (,~llLJIlf!,r p ro:J(~SS that In(l~ma Hy iliIa~f'!tail'l:l
bal.al'lIr:!!i behwie!n IJf!Ddll!cbllrl .Illndl·~:unli~\I!!r 0,1 ll:eUIUI[alf S[fV r:tur~:l>.
<I.
~rrU~~miUl~mil;l OO~poil~tl
(].~:"IiI.ii:iUillde,'jigned~n ..
Iimul p<!dmgt:n
Inet':~!:IIl]s.rnc~I'illilitsease
l1IilY-''IIif: bein.£ mpidily unraveled, 0111 ])3ge ] :3 98 of tIl,i:sisswu!, Lee e/ {J'I. IJ~ cover an ilf!l.lrig,l~i~gr1!~~Vijmt;m;.kllll\n"'ir.;'!~ s~,:l~si!'!g d Ih,~l i:llwl)]vesi!lu~opl'nagy, the 1):J:ioc:es.~.by \"'~1icJin. s celm n.orlli1i1l~tyd ispase elfi.'lte !)r.;<lIlelleH amI prowim; to ffiilinLa:iIJ1. hOIHDO!lmS~I!> (n Tbreugh i~lIrophagy, llh!(1; !O!pot;rnti.o~} of tWO classes 0 r viral-:sell,S,ung receptors o\'eih.l:p and are :1lI.ol
.,r
- -- - - -- - --- -- ---
--------- -----------
j;i0i1 hloo.i<ilY
Ol.'.g<:mli.sms senseuawanted virus lhlffillgh b<lsk' !>1.lr.lt1;lg:i~'Sth'll ~r.dyQ!'I d~1i~c'~h)Jg f~)f~lf"dregjl'. Sh[Uif;;'d b:}· a!~;
'Uilittde<iJI'ed cells, i;nlvol ves ~m~t receptors inl.llllc (.)'ilOpJ!~'~Jrill hm recognize RNA l<1lliclpmoo.bly t
DN!A, from\~n~ses (2)" \~rh~n iilCllv.ll'ed these rol~rh)fS pron~tl~!..! tll~ exp~S$ion of m!U1lY genes. illdlld:ull.g lYIJe I. illlterfe~{.oJ1is,(.IF.N-W~i), (,~tlk ines th ..tare criti.i;<il f():j" vi r:fl I rosis';;;Jim:e. T',~.e othe.r ~f"'.llegy is II."eslr,ic:ted to fe\!!.!C::rcd: ~:ii. fno~lt liilu'tiJbly of I.he ]mm~li'ie$y~~..,m. U relies QiFI .1 sl~b:let 'of receptors of thii! Toll-l ike lY'Cepl.or l rt R I r:al~l:it)'.iVi!cl udiun.grt 1t1. 1 LR7. n.RS. and l"LR 9. These rec.. "epliQl'$C<11l access. 11~ endQ"flnmaU co~mll)f~l'tnle'IliI'[L'ild ~UiIS(lllt~IL:l1c acids. Or\~mses.~h;J~ ~l<li\iebeen eu.1WiII:rt:d by the h.OSlcell. thll!ULgh tiliD:ep.rocess of ~ndocytMis,~ 2'). The emlo)lomal sy~wm can ~ftt,'CTh"(.3:ly 00 00Ilsid.e.red 3. topologi!l."f111 cmHirllllHl:ion Orl~llC extraeellu klll~Jl'<1,ct:.~ n fhi,~ sense, TL Rs. presL
l-
Au~Q;pha.g:Gs.ome De~e(tinIQi 1Iiral inifel1:t'iD:Iil.Ih ro.l!Jglh ~~g(,!f:lOIJS PJ!IIlilWlllY, RNIiliwiUlin inlte-rIiIalJil&l virnons is detf('teci an
I'D)"
TolHike ,roo~llllQr 7 in endesemes after dig:es~iQn 01 the '!o'i~<!l en'ielo;p~ and capsid ~roteilil::; ib1J hOSlt (eil efl:il.ymes. Inlil~~ le.ilflO1gel1()US Ilalhwa~ illlItop'llag¥ performs. an 11l1atogffil:li fllll'LclioFl to' ~mdm;)iilo~l5. lJ;aFl$~~rrinlg vi~a! RN'A fmmlih.e C)'itolpiaSim ttl, irilracelll!.dar C0i1'1llp,animenls (lJntahlii19 TotHi~ ll!i3Snm<1t"yLOI~d d.en,di'r:j,lic cell~s, Sipe!l:i<lll~.lfldells c flf the iili~.ml~mtt': !'>y.~~~m. ~Ik:hsmlnll;;' ~.FN~a ~eCepl\!)r '7.
~() recognize
elm·al;.,~nl;Jj.la!i
i!lt:~~ ~U's:
mjlil.eu. This means that, nn~i.lkelh.erecOgilmilli!Oll "':f.'~~m th;,!:~rei ies (JH .ci't·oP la5liil1k.~ roc~r~~}r~. l'lme endmo~.n~]TLR system can detect virus
i~.d~~nde!~I,~y 'llf,lt'lmd cellular infe>etionan "e."'O:!:l~nmi5·;recogllJition P,n~\'''ray (~e!w.e figure], IIIll.d:GClCll aelhlat~(lIlofemk,somIDIi TUb by nO[li[f;feCl~lou:S i;Ulr<l:(;ti~Lted viruses :h'~\S been
pUld.llctio:n lht'ln~re<lHb1en] of the same c:e]h: with i~;;U;:liv;rt",d nmm;;plkat~ng viru ... s, Ye'!.. e the response depend!' en fUR? in hn~' \vi~h I~:rcvious obSlel"\';u ions. lhm Il!lmm, ~yl.i()id d(!n~ dritic cells ndy heav~l:y on lhisiJJ<rr~i,cular ~C"l~~or to n;~p~mdlei RNA V~i'il,I~e.'t s l>uc~m. A TLR7like~y [ecGgliiizes "iml RNA iliep]~:C:<'IIlilil>S ill~ ful.eC'y~Qiplasm,m~h.erl:I~",n the' viral geamOlmll::
Auwph~g;y ~Hrn\is.a cell to enlgJLdf'pru:ts of observed(Jl'\7ilh.u Le~el' (II. de!n{):~l.1>~r~!l~i~. own (:~('llp[a$.tn to I"Gcyc!~ cm~li'm]k prodil~ ~lm.3tlimis exo~encms p3thw.aylism.ol~he' oll~.y uets an:UIsupp~y anabolic ueeds. Th:ils en!illl~f~ m.echi<lllism l~y 'i'I\~.ich eells C'Ull lUSC TLR:s ~tl ~u~mg~n(;nHe$vesicles {lliIJ[opht'l,~o~m~(;;s) arnnems ~tecr vur,ll presi(;,m:~"The ,mthot::; iIlol)l;ld l.h~~l that mature imo degl'<!I!CIi<lliveCOiml~.p for :m:IIll~'t' viruses, sueh as \'esicu~ar s:romat:ilis (alltO:lyw~lnes) 3).lh~ :p.mce~!>is ~1b1;cQSi.nlg virus: or Sendai virus .. ~mLfect,uQIlI o~~ mliri,~e as iill iq,'!!lPOfilmllil p:&i.yer i iii muhilJle filoets of,eeH
I~
com pi f:% ~:MlliC') class U molecules J). Add i'~ il131fil ~y_ .~utfiPh;).~y cnn 11C1 as an iml~"Ul<e effector pa~]m7lY fur ptl~:lli.ogenel.i:~null~lt.nonby p",nnin~·ng clllgjl~~n.l'lt(Jf~ of kL('~l;JfiO!!~:i.'ll\~(fl~ III t od'qefwi~.I:perNi~t withjn 1~IIa.tlgQ.~orllie~ the or {~Ylol'lm;l~ll ofinJcrl~cl cells (3). Hn\veven·;. the ~1ii"'QhiCn1emOrmlilUplm.1..SY im.lhe :inlli'l] st"J:1s:ing {~:fpal:l\i.tJgem;h~dHot '~!rn f~pOfl!~d prcv.[lJu=>JY, ;um Ihat is what Lee {f( al. setout m deLenni ne, The mJt~.o.rs ~nalyred <Luwplm.gy i1lpb.~mi\~ cywid de'~diilk'~~dl:slilk1;l!lirom a tr.Hl..sge!~ic mClfI.SC~n whic~,lhc all[Op'~~£Ooolne-1li~oc:ial:Cd pw~~in LC3< was fused io green fluorescear protela. Tell UJ 15% of the dendrltie cells dus~ p~~'C;;'d,1plmCUIW n~lO!X'~oom p""lu~mindiCfi[i\r~ of lal'ge"'sc;d.e amephagoseme fm:m<Llion. fu;r(l&l!tr~~u virus did !lOT Olher the overal I IfG'-' que.nc,·y nn~1I0~l1t c::e1m s,~iI.lth.Olllgh i~ poss~:· is ble·thm. ~}al:l!5e..<;nthe f~"qll~ncy ofau~apl\i~gy i WCJilliM be seen I]mQrlSH~e S~l~ ~mi.lIIl1l.'i1ber f eells o
p~.1Yl!:iol(Jgy.
1mlHuI101biDlogi~'1.~'btl!rmOJ)', (i!in~~ ~~rch
UK, IJOIl.dOilll LYn~(]<ll1I'!i ~ields Lilooranulfs, IInil M Ulllc~~n'5Inlil Jilidd~, LM,dcn W(VI 3lPN,m:" E·It!1~;l; (;a;e:~illtD@(am]~~.ory. uk R@jeardhi innitute.
devd.QllJlmeilt. <lnd g.elresce'~e . ./'\.mophagy a.lso regu.hu(:S im[ill~l!dly by pl.'oW.di.mI,g cyl\Jp.~aSli~~.k: mo~~.iml ucan 00 pJil)c€C'li~d fllr prc.'i'~cll~;lliiU':.lI!t [h.. ~Il"ssooj,ui:on \?!.'ithmajor hi!;tocompmihi]lhy VOL. 31 5
th~ i.l1chldin~progliiu·~.~nm(l t:'ell dea·~I~. that ~lC:UtLaUyg{;1 .ill~~cted}.To.~u;icln;:~i; ~hncliQnalimporl<lm;e (If l~i!) obselrv;uion .. the author.;;; ~lalyzed IlIie drOOl of .illhib~lifllg elM,s IU plm:sph.flddY]i'!ltlSUlol 3-ki~](l:Sl:;S;e·~yrnes th.u are (;~i~~C<1~m:ex.eollllllg a~ltophilID" Tlmey f SCII!:.NCIC www,.sdencema9'.o~g
9 MARCH 2007
IPERSPECTIVES
alse checked the' functionilJ (:,IPfiCity of~)I<Lsmru" cy~oid clenddtk: (dls~lhu lack Alg5. <1 necesS<lf)' compo.m:nrr of~he mIl:op11.agic .machim::;ry. r'l both eases, (~I.iIlli.!mti1l1iga~Uophflgy ab!llM.flWd eytekine [-esporl~e~ ~o imfectim,1 with vesculas i;l\lm~lilL~\I irus, Alt1!lg tlll~S3!U~ Ihti.;l. !llk'C~,~~l an AlgS~el1ci,em illmmll'le systeml f<~ii,i.ed to mo~mn ~.rJpid in~~rl~'fOill j'ii;-;spult~~~dC'~dGf.i1 on phlSn1J<lIc;f!oid d~nl\llrilic <;~II~~ he~.ilmht~d w 'wi.llm, vieus, "fiitlll,S;, <luwpilagy bylh'IIS inmnl".~lc' cell type isesseutia I for d elu~'eri1J:,g t:"ylitl~,1as1'I:111l" vl:rni II utelecules, \\,~licl'n, simIMm·:ty survey those srm'le ~'I."'(_I regi(;l~ where pa!hqgJ.::1'i!S em1JlJI]iJ·' are tcredHewever, just as the divisicu of labor ]Ql !nasI cell types, atlloph,jIJgy inlC1rU:~U;;~ hl b~IWOO!l MHC d~s.~1 and class Il is :~]Ol IieSjmnse 110 cellular stresses sucl~.as !l<t;l:iNIniull 'lbw,I!ill.~e. the c)'~oplasm:tc <JimJ endesomal or~r!I~ctifl!l. [n(:luc~h.ln of autt1phflgy ~.po~ ilnmt~ i!l1nl!ml!l~rc(:(lpto:n>m_~.}i be ~~i!1g ov~rinfection. either <'II'S ,,. bO:ii1 l1espO,lrt~e or ;<IS lappil~g \\'OiMs. AlI~(Jph~g_v calli :pmvid:e endea ri;l!rhog~im-drivi.'f! rmoo~" (5)1,.M~y ~hiJ!~ ~dkl1.'r g,~MI!fllL~"~lig"il~ lor MHC (:h~$[I pre~!1i!r.ilat:kli:'i 01;1b. Qther~~1.,n pla:iirhii~cyIQid ~k:g'JdJr~ljceel b to I J) '~luLas lee ",I' al. sho\v. \''.~nli Rl\v\':1· Iilr make use arT:U~.:s for dl;:~ecl.. ng pathogens u oocogfil,itiom 'by ~ "fUR • .com eil"Se~y. Cm'>S~J!lfC.S~ in the ':y'I:op!a::-;;ml. H{Jweye~;. it also raises entatian p<.Q:~~w<Jiy:sa~low ~,il(' class l moleM 1>\ pmb!cm .. TL R7 ;1"C'~f)Qn.dsto slre~c~,,~l'i of culm, mprcsem emgc'l'ftons al~~iMel'L';. much like polylQ rielyl ic acid fouml in both host (seU) and cyl:opl<lSrnic reeeptors C"dDl. sometimes med.ion:e ~,~iF.lI, RNA (6). Discriminat ion bel ween Ihe l'\'\I'O reSllOl'I!,es tosln.lmlll.i 3ppl~ed lomhe owside of <I is ~~lll~mbly li);;~Mld~mth~ f'L('g thm viral .RNA ~.J~I 1"1). ~rh;ip~ tile m"'CIl<ll1Ji,,tll;S lmd~ruying (7, is 'pm~:et:ted \1I,'.ilhi:Jm :pro~ei aceous ('ilp.~ il n id. pathegen S!e~$oi 111g, y the' .illlla~e ;I:Ild 3.d!i:JJplive b inlm~iIiJ.'":,,)'slt"ln~ ~ru lI10t,'ill tli (r~rum <lf~~.rd I. , \"'~.('\re'-'~ r RNA. !i~ :;el 1!lJnprot!!l~ed 1Jndsuseeprible [0 dt!g,rad"'~.iQn b:y exuaceltular RNA~e;!>
II
place; Tb~l;S.the i~~e of alJl~crph,,"1!Y TLR slgill1l ~aU!l!,gIn,My extend be}'tl~ld H,g;ll1cldel i\ie~y and .im::~udJeiuu';)ccU(JI:m: receprer Irnffid:.illg.
~he,reby .d,~ow,ung l:ecogni.tiO:IiI of some RNA viruses byTLR7 (f). Oneunexpeeied Ob~~~""'iUitl'!l~nhe study t eeuid 3 (feet thiscenelusiou TL Rlam:!. TLR '9 ....iyul!.di IDlgin ~~(.l!\u;ell typ€$!<:.iicit:> prodacrion
OfC)1ICddlle!l
su('11. &~ .imcr.le~,k~n~t240 qll-r2: p iFN-p:. The db;tin0ti\~t:~le:ll~of pl<lSln1q[oid clell1ld 1fi,IU!C cell slies (}OIl rt~y il1l beilii~ p4U).b~u
l'ijC!t
d.el~"efScytlgpimmllit:
RNA directly
tQ
TlR-
di:;cri~~i'l~~d{)n
comp<"Ixtment"~iheJe TIL R7' and 'fLR9 signalm lISt be aooo:nmplu~hec.l by add ililOllOlmeass. l ing is I;l~Hlpl~diJ) FN ·q~N;"p~:;;~i{J'fI!,t4)- S~II:1 Umil ml'l ...tbe T.LR sy~l;l!U~~ml . f.:ytnt'.ulk: pris ing~y.the iUI,U1m,Orsi nd Ith,lt p~mil],KL~~)~l)]cl itt'CCpt(lft) were t'~Ullgilin!t l{l~rl:brm ~!n[llog:UlJl;'i f d'el'ldt'lll.ic ,ccHs~clill~g AIg5 produce Il~ ~121~. I.hnc[ ions b:u in d:istinct regiof1s or the C~~ I, [fi'or bm IlOtlfN·o:.. in ]ie~puJ'l\;e to r;;:~",og,emousTLR i!} ii.nnmmilog;i~"'lsbmllgJn up 011l;,t diet: orud!p'~\'1i! agoni;f;[S_ Atg5 may oo.!wchrnv be illNo]v'cd i~n .h.lununily-I:!mt is" ~mWl~1,.ufU:: c.sllo.~~~~b¥ :~!rl~~~ r plmi:yres-[h[$'~.lS iIJ11en:SI.ilmg p~J"dnelj'\ wi.tl:! soni~ 11.R.g to the '<Ippropri,t~1!: ~QI1I,1Jl:Irf1ments. where sii,gm,1 imrg:lbl" IFN"ct l'.lmdm::lion C<l1l~<lke' the dlil.:llO~D\lrn,lY between Mile elass I <u~dclass
ril. Miru5hima., Ji. I..... ,a$ilk'i, Sr~Gl' ~!IS,:m8t,GtJ'7Ji, :2. If•.~'W1lii, S,A!cifa.IWJ; !mm!i!fjQ1.,', HI j:.l~l. 3. V.Derf1:i(, CUff; Oprn,.lrmmuool. :11. ~7!i ~OOt 4. lie Hooda J!'loJ•• 1II1:liWt!4.~<l,lOO~ (200I~ .. s, ]f,I!'!IH!i!'f!ii!l" xiem'I1.;!,'l2:, :illS (~~~'. 6.S.5.,Dltbotd iN DJ.. ttl" J, IlIJ\AWoo1..::16., 3:1156(~006}, "7. H. t::il1D 1:'1or" Na/jJR: ·H:1.• rol(.;l\OO6'~. 8" L GILllii!'i ~ ~N(m. Al(d. Set USA ]l)1ll,84,5' ~2(106).
]I..
APPUED
PHYSIIICS
MfcrQe~dnmks
.aJr~blJitt 'hom rnalt~ria~s ,,,hose e~e(tmn:5il1iterad s~rongjtv .anti (an 'l;1xmrbit PfQperties es iii grQI1I p tlil~til re not. seen ~n ailily 'Ol'lleIhIllrtkile,
il~1
emergent slilte' G'J,Ued the' q~IJi\lJ:nmn:HaU. ,cl'ftlCl. Thesppearance of~lr'n.c~ t \.~,'fll 1:C31~JfCS ~,:~~nl exide ~.2' a[Jm~le~~:(Jirrn,e ourabill,il), to ('llUlIt1i.'ol ill materlals. MQr-~()vC'r, l~,iilIe'il'cl of OOml,twt)l ] $110W$.<Idet~rerp'llhl:O '!illsi'lmgs~LCli1. r1ll<:t~e~~<~ls i~l 3pplk:'ldollS as \'OIlIiied as, advanced co~m~pulillg ..ud ,d~-lm il;JllI:rgypr~;ldli.H;:linrt IEmerg,efll phemmlclulI'I2' •. 'illl condensed
lh"Ul~J 1;.~'-n:iJ:lIlgit lLllid~LOSIOlJd :~n l~U~S gf ~il'al'~ hoe
1['De\V Ih,ne~ia~sj)()r
liiIl~11It~~I1,~u~b;·th~t f
be'~t,eil:'into focns" E1'Im.ergem p.h!ii:'nQrne~<'I•. iln, \'I.r~k'l'ImL.ew prypenies .~rJ~·frum dl'!;linlemcdons of 1rn'lll)f p~lrlk:h::'s ilJ1l . ." l(;omplt.'X s:ys~.cm., g~neL'me specIal ~nl:coc"St In the expc~:in.~cnts, f o Ts~ik<lz.akJiel Clf. repom:d on page 1.388 of this i&"'lle; ,{ J), 111~ !)l.Qt[mn;; ~hc)\1;' l~f]il;:\I' nov~ dc\~c~ eml be rlili~.de from 3 dOlls!s of m;,ner,ia~!>~the tnmSh~OIH~l~~1 t)~id\;.;..,._tI~l·oft~n ~~~O!i~>i~!IiLc~ $ of emer~e:n:t p~:H:n{)meml \\,'hel:l illl bun\: ~:Clnn, nJC dJ~vioe.~~nHcl.c by T ~ul<3Z~ki (Jill!. exMbh bi~ di~IItg;!e C<I'Ii'riJi!i"mobility. ne:e.h:d lOt ~ligh~
pcrf(,mnmwe :t;'iivit.dl:~ng.
iI'S 'we~lI. :usan cx,otk: :
ple iareractions bet ween 1),lin> of panieles. E.xmnph:s ofsniChplc~o.]1m,eil:la llIrc nUlgnclic exe iI~~l s fn Inw-di men sien ..~ material s, i(lT1 llUlXreUllduet ivil~ l!fQ '''~,C!)v)lel~'Cll!':!nii"111I;agnel~.'lnd fh.l~l io I'mII )' ch.ugedp<uti I eles in ,I 1'i,,!r(H~imensiomd electron gas. A lU'3d~liOll<i:l1 !Ut!~ ~{l t'IHl\;lrl'>i~ndt~g th'i;;li5<~ k~mt1l1f I,)'m\:l:igent slates is to C~iJi~e t~u::;m,illl neVI!' mm>eriills. ' [11~hi5 w'~y.,IJII1~ emil stllldiy :;t~lt!:."'\lith diI11\:~!l::nl-
1Ihe .il1lI~hllri~,rwith Bell Ldb~ A1GllM-ll!lluf1It, 600 MDUI~!:ain !I~enue. Murray Hill, NJ 07974. IUJ'51l E"F11a~l: alpr@
Ilil!'iI-il1Ib5,'oom
O:~dm!lt@;ri'i!i~ MIiIi! lnl~~ p'rQ~@wti~,~.(1L,~mS'h~m~li( (iiagra m of wide I~ycr S:1J!"\lI(II!.!t~ l~ill Tsywl:>:!,ki ef ol IlIsBd ttl OOSAi'Plie"ti1@ q 1I!<lIJ'!/llilIll HaU efha{t (Right) MkliOg raph ,of 1il e .a d.ua I. d'~l!,i'kes UI§ed. 'I'lIh@Je f'll is 'nil Q< 9 roilf~h temper:clti lie, ,PIs;F1glede1;'ic:ewilih dkneflsio;M 22 I) ~~ x 60 pm i,slilig Illig tilted. [Adaptt'd rrom U}~ m
SC;II~Nelc,va L 31 S
9 MAIRCH ,2007
1371
PERSPECT~V:ES
characteristics and either tesl; theoretical de~rip(io!\:~ ofsu~h S(~~J;:~, (:JfIire~lize~mvr~ly uew stares of mauer ..Searches fur e,mi,n(l~'.b"eln. phenomena are .,lfl~nperf(lr~~ed in blll'k or (hiJ:l~fil[Jl fill,alelrijl~!\'. either e f}1MOJI line or
IIl;llJlOIllC!LCiIi "f.iC'a~cill size. The cQII'I1me.,c,~,,1 development of <I new lu~u!~~i~1 :i(~~bi,'Lin~mcrg~Im,[bchfivioritu ~, ¢ c p~'rti,cll]!'lr device <lpplicati,on Pl'eSflIII:S.1differ~ ent selt of,eh.dlenges" J-\,ltbDlJgh Ihlis res.earClrn, ~s !1iI~m;~all be "'~lered to transport holes as well c
to
HIt'UOS
or fUI1I.GOII
as ~lecmms,
l1(l~ h
~ight;",eN:LI:~lligdiode (I
To c-refiteh igh:-Ilerror]fl.~o nee devices, hewever, ~JliN neccssmy to el,imi 1l~L1e l.lefuct'!i. Ihm tijl~
Canll"i~p
n.
I1!l;cn'iyl C(J!llStani~S: th~ electron k ch ....§1ii C' O,'Jjrld r 1PI<I:nlck's,'COil~st(!llli~ . The e,,,,u'CmeprecisiQiI1l (w h
] purl in Ual m il~ k~l1) (] II [he; ~KpeJtUlm.C;:~M.a1 resis,tanu.ce pbv:,ms \",.~sexp;I<'I~'Iled theoreti('?}J~}I by Laugh] iltl U 6) ,;1m;!since 199.0 has been the i~t-enm!~m'mI.sI.unlawc;1 (lfre,,,i~l .. nee, "['.he ql~I~Elflt~11U ~n ~1.f~!I2l been o'bE'1''l.lc~d H h~... ol'lly illl 01.few m<lter.i~lIls;;lII.u! never bd:l:re iB 3i d!evi.oe made 1i,'01n oxide mal.e'r:i<llL~. In <I re~aled de~,;eI'op.:L'n~IDIt. N(}\'l)!ioel(/v ~t di. n:p~nl~)rlp<l,g~ .1379 of this: issue II i) 111:1o~~c'rv<llio[;Q or lill.c e "~~I.mwm H,dl ~'IWedill: rmll.lll'e"ml1er.mm::: :in one of the :si~~P~~IpL"IS:Siblc d~v!c:(:s,-a Sl.1,lg~e~ al:mn-lfullick sheet [) f ,\:IC'lmlell'~fllcm;oo~'lI, CBl~ 10d graphene.Ue'roe 11iD.e ebservanon ,(I r <'11'1 etteer
1~i1!lt ad h
,ellcetIDnilS
~lldlllllUS
reduce
1Il1~' switching
speed. ],[I<}dd:itiOl~.the presence of sueh defecl:!li cau~lm p:~Wl~T [1\01; ~um~~t.im'1lofd~~kat~ (~ncr~ !;elllll,states, The absence ,of clmm-ge ll'".1,ppi~.g is measured by l11.en13te'ri<1~"s IlilllObil it y. 'U seful .,im~ <llI;':IlMw~ng iubi liwmld cvn.lp<l~ibli~Hy Im1biUtY\'<1Iw:,-; Q1;J;;Uf n\t\,'( ;,:Jjn~I'\J~l~.tely\,\iidlil rei V d with ,exisling materiala, ~nd<lt, dru.v,ullg dOVl':f11 rnl'l~e,.fN:ll1 I Cl~;l:l: -I rl (:iInCOITlrpLllt-er isplay,'i;~ to :l(J\JO crrt V-I :...-1 {iu ~iHcll'JI used i~l ~hq:OO:5t 'OJ ;1Il,~.'lJi~l.r'I1;I;~lr,in,g • .'i'lu;himi\,~ligo~tim~5 ean also IC~(~~Q qu:a~hll1liycly nco\\' I1;liltlcrJ'a~:s. IUiCIiOpl\:)ce_'iSor:sl ~o more dum 1,0i!'GIU:!V-I S-II (illl:~~.ehighen..q,~lality gallium ar~nk~c ImtFor ~dlS;l"lJIlOC"as pan Qnh~pro1iln'lln 10 s:ynlhc~ size ever-purer :5.eIH iconductor :S'tl!'UCUl res ~n erorslmotu£eS cocled down to ~,IOO mK). Reeeat th~~ 970~. Cho ~.ml/\:r[~l~!r d~vdt)p~ the ad ~r.ri!lC~in the g ~Yw~h '(If ",i !~gl e-crysw] .[!!ld (1'1), technique of m.oIWLII<IU'redrnl epit.'l.7!:.Y. \~i,1midn, OIidel.m nhnsol' ZI'IO demonstr.ned 1ll0bi]lil~le:; il~ hiShly ordered atomic hl).JQfS1!fe ~J'WV!'li oJind ~1I1~,'"c-e$$(lf2J)f:r~I,m 5(1'K ~12). [h~!$aren~ng dl~ hrr[io~~m~whe~ to t hel.:mder.I)I~.ll1g subst I'lUlll. dO!JIrw hi;]n-ql.l<di~y o..x ide <.!tl\im, Molecular 1:tC1I.~ (~pita:..y 'Wa~lfill1Jmcrn:fi!rned 10 iFtlr Iii II~ igh-'lXI.dbnuaIm,ee qU::llltuLu \\'e.~~it. is create '~;be I.wo~diUi.~elifsi.otl!oI~, s~mc[ures lea.dimiS e;<:;5eml:li<lllh'il [he defect density in the :im:erto thlil Irn(!:tiomd q'JiI.~1:tJl,l~l H~JI elI·Cd. a (:;Q.f!!l~ fsee ~g i01m b(tlow. Sbrp i nte rfaces h~ld already been achieved illm a dn,fr~r>e1H p3i~: of p.!etdy mte1'ipeded emerg-ent ~mu:eof'e!ec~rOl1ls. d il:;c(lvl:red by TSH ieu'J, (j: ),. 0.,,,, ide:'Hl'i'llerials. sl:ronti ~111~, tit3111ate/]l,mduU'mmtl The iienlilfcond~!c~jl,gbt;;h~vi(;lrJ! ~dJ;,\r~fil,g: lh!J~are ('/3). and o~dnmions, in ,t:~0c~dca,1 I ~ modem micme[ectllro:l!n~cs is fO'Lm.d.uD1, comresistance as a ~hncdon or:IJil.~OII,gneluc fj.e]d ( N). r(l~[lJiU:sor s.:i~icon or cl!Jmbint~l']~JgTh (lfm;lre::ri",~ s mHrn<d~.y auU'ibl!wd to low-defect metals, from colllnm;, HI. and V oftlinc peri.odi.cllfl.b~e, \f\Jcr,cebsereed, Tsukazaki t'l el. were able to SI.IcliD(1$ g1lmWlJl'il ~ud ~rs~'~j.c. E~ul.,;rgiCnt pheac:hi~w slmllarly ~O\V defectlevels i!i! ~hi:!!: II~m.nem<L.lrnoul'ever. oceur i~:l~. wide \',m:iel:y of \\o'cn1.:to form ZJ[lOfM.~n ~~O qJ lI<Lnl;!i:I '!ic\o't1~ nil Is m()bilili~sg:~a:Il;lr d~~~, (l~~f !f!i,~~~~a~s ~uchasc.trbPll':~l~d fl~~miC". w~lh IO;:.\I'-te'mp~r~U!re 50!JOcm2V-! s-!. . in~ennel;i'I]:~ics.'lnd trmro:llituOiNnet<.ll. 0:,,,, ides.Jt i fu; ~he ni'\id~s, I:lotvever.tlull p~t,.-m[' dl~ rich~"';1 l'h~ !f!!!t.,rii!L~9;1t! '00 iOO~ ij~~~t'1·~ft_lKtfl~O~ U ~~rh:~ {If eltl;-er~'nYtst~.'~~s. ;~ncludililg mel.~l~ lt~ru:i! sJ!w" Id ,,15cOr.1Cg I~~"ne'~h,e lr,ml ~fil:lr (l·f y Wil:~,cl(l~tmll ]1l~as!>Cs s(\':~11 hllflClred ·~inle-stht e.h;:el!l"OO;S fh1!mlll:hc buU{ hno the i.l:Iil.erface .• blue m<ts>s. !:lP;~1l :L~~;Jgnl.eti~;m l'l'ith 7~i:{l-po:im; T:i.uk<l2dil-d (it ai. n~~~d that wln:n-;I!:; lht: ZnO C:1~lmpy. a~d S:l~r~reondllclhd~y ~H n;:oc):rt1i 1il.)"ii:'l"SilaveiS~£ilbl~ 1l1:,Il,si~!C' l>lrai~,which~lld~~ high ~~:i'i!]rGmhlr,*,-M(.ioori~ls pll)lsicisJ$lhi<Ilv~ ch~rg~ S('gr'L;;g~l iOil i.hn.n~ghlhe pit'z(ie'10clrie drc<lmed of cre:~'Ltingun.ew cl:m;ses. of de\r.ices. dlect. the uffitd:erlying ~1~g;,(ZIlI"",p 1''Lyers ~l~.\'\e (~~Ll h.U"!1e55 ~w;h lin'1;l'lfJ~~~l~~t(I~~~ {rfl'!ll1]I~(lr"rH. n",gl igibllOistr.dn. nus. O'I!'l1>lT eh.ny"" dcn:-;i~y i::;; LI1~cd. \!II jllh tile'end ofMoo.re-!j la~r in s~.glu as. ~ndm:ed~1.cc~Msnatli,O<Illy lron[fuie Zn(] ~.'egion . ~Fdins.i5'roB il1l:i1liC'ropI!lJce:.;sO'~ OiIflpro;_ldl hl'l,.... - ~rll{ltile imevf~lCtl.lUle .m.hQn;~~urlher de"''I;li. Il.l.U1olne~eL'-:iC'lle d inlelm:sio'.llS. e\"elIil ~~~e.\1i I iC'O'm1 U:J~<1 <t ",ynlhesis ~oc1.Hli'luel~Ht .ll]rn1l'-ecll~ht'1n to illid~,t~t.y:~tx]:kiI,gl~~It!;mmivc !liM'I~iial:s ij:~~d v~i.ry Ute tll~~k:ncs~ of the, Iilyc~r..., h~~1C~ th!.;;;r ~ I> ou:'Chilet::Imes ;fi:J:r ,oompLft ~ng beyon.d 2:020 ( /'), om~ld study MI]lP:~es'\firdl diiTereil.ll dmrgc' denTt->\llqp'..;;lki~., cd, des~ri~ (l ~!ni~tllld~t~~~ar ~;!ii~s th~ t'''\:I-di!llePl!- .. io~l] i~~J;rr~~\~~1J:io!'i. in ~,1.e1iel:ostn']iCture ;r{):rll,~ledat. tile ilJ&ert~ce of lay~ These elfQits r-eprCS;:'Dil~<1hi gil degree 0 f ern of ,dnc oKi.dc (Z~.Qrl !I.nd a .mi<'l~!ild ('OlVlI~ Iluu(\':r.i!)Is cOIllE.roll. 21ml as iI resu~t, T:sul,;lIZald et p(lu~.ld,1\11g"Zn~~O(see ithe I]guooh These two(,) ,al.\'I/'f;!:re .ublc: ~o oooeryequmlli7.)~d v;ilu~~~f ~~111 eil:hd ~ Si1i1i'llC d~m::r,ent v;flhLUS of th~ ::>cm i~ lhe i laU 1.'e !Srill~CC 'lrt:hucd [0 tllcvobgJ.! .... (;ol1dtLc~ilg !;!,np,. the eJIa,e;t:'~yneli::declw exc ire dei!,felQped lmmwt:~dy",cJos:s a sillIiI.l:ple i.n an ce~t'lCtm:n~Il~oits co~]d:~.ction ~Ix!, Th(: d~r~ !'espc:msc to~!)ngf.lmh n!l:~ t: I;ltl'rellt 110\\1')~~tC.l'nl~ l [il:!:e:lilIl .g!:1.p vOllhu~s [uq:-1lH ~lfIfltL;;mhd !;'l!l!;l'rrg.y per.;lJuoos ri.e-"" .. ab~(lhn~ Zl;ro,. This; !I"'l-ci.ln~d q1l911il.rum rl<li~,1eflt:ct lis, !he' Ilmlai~1.es>rIlh of the r (qlu'1Lr'ltmnl 'weli~ lh<lt oollilines, elecuxms 1.0 3. ~w{)-dim~,m:;;it'lm_.1 [resi(l~, rmper~!!I'nd lypicf.nY'ln~y!JCt'L!fS in lhe hig,h~s(puri tytv.'O-·dimi:l1siollll I met3ls. Tllll:i>, ,I hOI11ZnO ;already f~m.ds~~lall:lY ~f)PI iC<ldans fi11uk l;'};,p~ulmelU. perf6;r.m~(;;'d in l~).80by illdmlilllllg pi~'7A~ll;!!l;;tr:ic t:icMJl{Ir~ (~~).• phospln)J;s iiI di:spla)f~l:!h <!lm.d lrnIls,pm:em conduct~ vonKJ~ltzillg fl.! at. i,ll1. highanmbility :<,~~,icQI1 iog.ll IfnlS'~ 10). f\$, en C'Xflmptc (1rth~versa Lci:JIHy b~terflS!.r\ictul:Cs [ 15). ~~O'I$ llowbcel1J n;;pc~~ed of ,ZI1IO. some of~'~e <mthors oll'the report by iin tIll enu;ire~.yiil.ew I1l<1ten:iah:i C~355. \~;ekno",. Tsu~01aki ~l til. I13Jll fl:l'(lm~ElvShO\\lll dliilt ZIllO lhflt the I-I ;:III r0_~~S'~Il(lC 111 the qlUil.~,~l\lIn Ho:l!I I
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cortdensed-msner pi~y~h;s-'ltre physicsof devices, and the phyS~.iCS of ~luhi functioual '!UlllQ .... dililCU1h n1agneto-. and dl:il~to-~ :sens..~n!il ~nd .r.:on~ml!--,uld the liml)IHc.u ion:; uanseend I:~~ (l:b,vi[)~$ $(:',i~!lilti 1':tt;:~lu!";;!J:;:[iomi of l~!~n\'eril,g: an el<!Joti~ S:1.<'Iteof matter in a :lIiIe\x~ :1rn1<Uelii .. 1. Th~ pr~~'\;;m""Mp~ri!'rl!i!'m1~ sug.g:eli[5 U~,H \"'1;;
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9 MARCH 2007
VOL. 31 5
SCII!:.NCIC
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G, l<ind~ iff 01" ~ f (~w~ ,61 ? U t'!Ooo~. 5- K. s, ~~~t!H erfl', •.~~~ ~lt ]9] ROQ5h 6. t: Zti1Q_rI~ 'IN. Tim. Ii!. l Slarmer; f'. Kim.. Natr.rff 43:8. 't. 201 W)(kSl 7. 1It1is ~rli: W/1I!!. '~IIPIiJOIled by 'Eirn~in!!Jl!fiD!l~nd E'h.l'!iwa'L SderuclfS ~fli~ilrm CQ~ftd,rutiJ. tl!S:FIri.MR~Hi2:7:liSiJ. wr~.a9 r:1et feu Rml-(T" 20041- ~OI&:!l'OO, 1.:1.'5. De~!Olell~ or f!tu~rm~ ~DOa '~I[)E4iJIO.NI!l(~,~lJl] ,anel O£-FGOO "O!SE~A61'~l, '!hif Rcyal Sti<tii,eiy. ILeYer~Cllmli lf~51, ~igr'DW~! (~ra!irnn, andW, M. ~~~ l'oi.inmOlIiIifl. The e~iI1iL1~Ii'IISi.w.11! pa Flia'lly Ii'l!wlclmLlm 1i~I~t ~IiDI'i~~ If"-JhM~ne"li~ field L.a!l ~dl:01Y bupP!l:rted bit N15IF W!!IlerO!ti~ a~H'iel'!le'nl no. !:liWII" oo;IJ4:m:, ~yt~~ ~i:iJt~[if f1ilooa, ;mdl byt~~ 1000 ~i11d 11liIl'~~Im!!ii ~IB~~[I;'11 ~'i4i1kl ~~;mlt Lil,b"'r~OO!Iy
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mkiiogralp4'lOI~'OI1~"O~ t~e ~i~e$ u~ed'i!i!rn~m~~lI~e!'l'lelilts,1be sea!! ~iSgi~llJby the HaU oo(~ width ~1'12~Iflt D@vic@'raMcalt1lWl pmiLetluJe':'l ,velie des:crihed'irl (5), (I,~ ff).-y ~ed) and P;o;t fullue)! ;lisa r'll~liI:l'ir,m~r 'i}1!,OOvoltages; (VJ ilna m~U]liie~(: of 2'9 T, field Pesitive v~llles OIr Y~lind!!l(e ,~le~J'QJI5,ilnllil negaltive VII! li!!f~ oJr
6 Movem!iet aOO6; i!~cep'ed n )~~.!.IOIr)' 2001 'Pti~IJ~llied 001111 :I!S f.etJiWi)' ~oOO'; ~ 10"lKl!!li!i!:i~~~,.lH'i.rol 11~'~l!!!!:!hil' l!lf!!lffi1~~Q!l1 wM!:R ~~lmllh~~ ~per.
M13 9Pl UK. ~'D~~rlllOOlII~ IliI'f!r-;~i[S. (olwn!:iiil Ulliwm· 0'1 ~il1t. NI~lIl'lI'Q)lI:. m'l!CtO'2:7. USJi.. ilN\:i~ional High fl&,glfltti(
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efleets d~!c!()the ~lur!!"u;.rins '"ll1!i~1ixderm::nl~. the hmll~~.utier h~ight., 'l11d: y.a~i1'luon, (torn ck"lC's 11tl'!i!ic IlCICi'tlgi.';lIficil), ;In:- CMtaHrl~'d iJ!i J( 1', :;) {~~ ~~,pfi1.1Ii:'Iilil!l'. ol]:~iltc loxl l ~F(Ii' ilH. sim'ipJ~ Ui~(;'[!I~IJie
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Riem01ilng e ~edmns Ilrl(ml~he (;Uo.2 ,p'LEme cd (llIp[.i3itesaitt:e rs th,e ele'clmfllk ,cQ)ne lalliorn fllIni (il!ntlly' to pmd:u(;e hig hl't.emper,<IIil! re su pe rt:oindudivily. Ali{50r1la~E1d! lll'itl'l Ul'e!'lled~i3illig{Hj;.arE!5pe(tlra ~·w.eligM translers hom the higl'N!nel"'~Y s~.t!t@',!>of tlil,eilll:liu'l,Htlf' to Il(lflll @n@rgi@:s.. In tileory. ~@S!! sho!!!hil be Icll~OCli!b~l'! as (I'!limbtJili!(f!()e l:u?twl?!l'n'Dth!! b.mn@~]IIII~ Ir.ilt@ 1,0If'e~@df(llflill.l@iCtiol1l, "'!lid' e-xtril!cliIlJl=~ tUll'!ln@!U~a5:trJ1l m@l1fy. W@' 1rU:fiod'u(@'oOOmi(,~n~s~~liJtiQ!fI~lm!l.~lijnQ-asyrlfufL@lryiriIi'.il ~ inlg" n~di 'i!!] \I'in.uiJ~ly fcl~ot~(al p,l1ef!l)ml~g i,n lW~ ~i'!Jhti¥ h61€'"d~lle~1 (!li!pral~5t (~~~Il:8Naa12C!JO;~iC.b:i1!~ld Bii2Sr!?J)Y'Q..z(go.,~,(lI~O~~ .. li!it~fI~ spo::!litla~ i.I'~l'i~itlL',llns~il1ltUilh1!tiilil!) aSyiflllm.e't~v ()·i:!C~U ~rim~rily ~n 'the p'~i;ln1ilr oXY'!J,e,nsites; their $Pcatjia1lan allge'lII'1 elil ~f,orrlilS a (,U-Q-(III' 11:10 1'iI(j-(I~nte re.d d&~jro IiII~ p~U~iI'liI withoillt 10 1ii!;)I-r l'iI!1!If or,delr Iblilt w~1h ,!lo!J~\~ d~ lInidireuiona a I. e l,ec:trQlilk ®mliil~m;, d~~pe,rs.e,d tl1irQ)uglillol!l,t (Go: tlil,e (11-0-(11 lIisM Iilil:,e). The- emergin'g I~klure ~S liil,enef a. llar1.iiail hDI~eII(lci!lll~iZ:ati®n withiin .arnrintr~lniSic eleetrenie glal~'S' ewlviing, <lit h~gh,er ~o~e ,d,em,ilies, inte mrllptete de!I(l( .. lll~:atiel'l ,~nldl biig,he'st1eIM pe riJturre$'llperoon dllll(thri ly.
Eq. .~ ~htl\.~ II\!![ ~1imii! ~ i'IImp'piin~ of ~U': di f· Ili!rcnl:i.'li1 tlJlfmc:iill1':; roadllctll~lC~dJ/d~" Jll' (i". y~cMs'''''(1.£ = It!'), Ilewever, Im'1I1e ~I.rollgly rolTCb~ccl ,clee'tl\onic stmc5\. ia a m:::;hlJly hokGOI'ca CUpLctll~ .~hC' si~ !Jalt~tln is nUtCh~nlire complex. ~'n!hl!'(_Il;" (41'., lhi~ 'e(lrreh.tio~'I.~ cause ~h~~ lio Z( 11 m Ihc~~\'>t;;I<Lg~L;'!lS.~~),-or-[':l'~~C.'l d ~hl'L· 'r!lpi~)':.!:(Ik~;s, lV(E = ~ ~o lh,~1 Wdkid of !'im~!.!~'F1'(E = I,d'" to become .iI~)'nml!cliit· hy
or
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'C~JlhCilY
Ihe (uprntc C!lIO,l: planes derives I~ J) nom l1l(}f~!lD..xygcn 2r mm cr'!lppe.r 3~ orilit:."ll!\ r F~g ~ ('Oll!l!o.Jl1ih .. A)..
or
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R~"c~IlI]Y. il h,l1': b(J(.~!lp:ropo:,.;("iII~"ulthese dQpiIl,g-ind~!iCed oonch~~ dml,ges m igh~ 'be inIDI O~'D."'CL,,"l<JliJ.k: i rcCI~ya!>l ~1J1I d .U~ mI1l1c,~r or ckl;Rfon ttulll'I:H:li!!lM, (:LUrll;nls wil.1l bims \'nll!l~~ t4. :'n cI~iX.u'm'l ~x:~mC'ltrHIII negative :smllplc fuia."bcil1g !
sU·OI1igly lin'Mt'1i over dO:I((JI] l[ljt1C'lio.UI lit jXlsi-
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prooess, \71'hic-h gt.~'irlcml:CShig:l'a~-lt~liI1Ipei!lllmc :sIIl1pctr.Oirn[ll.ICI~viIY, Il1:K'11 removcs elcelrom~ iml'l1l, the CuO~ 'P~ml.e. milti:n,t new h.ole·like d'cctronic :>~fIi[C:S \:vi~l IlfCdomin~ill.lttl~ o..xYb'C'ill2p c1m~ctcr I( 2), This is a rn.dically dini::llC.l1u pJ.'OC\.'SS~hlln lu:ill;:~ dOJ~il1g a oon\,ICnlla~l;ll .scl11tcO:lfld~!iCwrbl'Cm&:. \"'~'l'C1Il an declroJl lis ~;umm"CXi linnl <1 'OOJ,wlJloo
iw~!.!lrr~Of.~lc $W1!.Cll, \'\,l~th \I"hkh il \;\'\1IS crord""o;J arc ~Ibo ahcrcd nlradaHII'.:lHt'llly. NUlll~rictd ~11NIc:~iulg of 111Is p;tlJiCeSll. j J ~ indicale5. Ihat\",1~etl n holes IJoCJ' cell m'Culll rod!!iCec~ Ihe cQt1:c~alio:n llrail
c1hm:I,1gl:s f,!,cn~!lll1,l; ~lm;I",~··wl;ig~u. 1:r.:I1l!S~b!'l;; imm both I~'I,I~ ,IUd. ciI!Jny hi,!,;IN,;ll"'~Y bmldl.'i
tive sample bias, SuC'h ~~Teots,siHnuM be deteetahlc !;\'ith .'L Sl'lllllrl h~! liAm'MCli i,g: l1rl~'eroSCo;pc CiTM}. The STM' t~smnpJ~ utr1lldihlg, cum:1H
Dj;gh,tCiIl by
!kl(" , i:;; hlJ-p~fniC hr}W~Jl!;l rme <lind! :r~ srn.tis'l~e$i ~)IU~O~'i~~i'CrgysC'tlk,,··<:: ~~ <: V. As fiI ~C::;lof SHeil iderLo;;" W~ show .im Fig.. I C
Ihe
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or
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/1 ) IFi !!I., ill" (Iij Rereva rrt e~ed:r'Gnrlic ofb!i~L$ ,@Uhe <~,d)~ plan:~ (iI,I3d or~it!ls .tire slilll:Il.'1nin l[)ri3ililgE! I'la a
o::q!'g,en 2:p Ilrb1'U!~ilI~e slhi,ow~ ~fl billie. A s:ili!gl~ pl~~ ,"ume (If r'OlJlf (IJ al®m!S~is ~hm.",n wilhitilm the d1asliied $q,LJare Ibo:l':, an~ a s]nlgl@ CIN'J-Cl! unit ~.:;\:fltM~ the ,(jash 00 'O\l\lIt I~B~S..:lh@nTOItlil::
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Co~lnel~ Uiin.eJ1U~, 1'lIil.Wi.. filiY 1485J. USA. ~O!liPar:iJiiIlent 01 M!l,anced Malllri~1s Sderlce.\ U'nk~.ml)i !!If i~k:ro, K!I~hil'i'<!, QlIIibii 217-365,1. J~pan.li:1~~fII~mel!tde ,2ItJl" (an>ilda,
of PltpiQl.
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Wa~o. :~aITalilili 351Hll9!1., J!iI!pan'l. SWnstHlJtf: fi1mital. R.Hea!'(1i" ~~O' UI1!I~~~sil!l'. Ujl, K)\O~~ 601.-OQU, Japan.
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RESEARCH AR1ICLE
mpidly wsh i~C~;!iSillg x.. as PIl.1Iic!~,] (.~ J}, :Ifl:~ab!Xi!~I~~t[\:h ~rrQ;'!:\. arc (,iIJt,,~lllb~~ ":Ill,'lt!"]!YI1.~{)!vl?d i~~hn!qllll~ {6, hy and di~llini:ilhC's
n,
the Tt\ flrop~l~ l4. J), also Wcmir~i ~lC~~t i1~om~.scglle Pl\~ Qftlilc ,~p:~I~g-i;m,!I~~O(.>d cerrehl!~onch~n~'l_
Etl!i(l~f!l)iilk"'dt!iS,~~~ gil.i!!S~·~~~t~
hQle-do:p~d IliJlprafbil's.
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~mr!.ir,;;I~.!y!:h;;ul. 1l;:t!!ll,,,1;I;';!:m!!brn:l~~.1"g1l1(,,~.i~~I~i!f;..; d
:;!Iillpt;,~lrs<l'i ;l ~lpt.,o~':ho!~ d!;.'Ill;~i·ryp~r C~iOl1·'~.J. it) 3%, The :.;~r:rt1~}ndiJC;:1 iv ity ~i£i,iJ~llly docs nooi aH]pc,u' lnlllU I~ - :5 to 11m, TIl~ref{lrc, ~uuluh~t Inl\,,,.~tC~.l~Ol~I,I~'t;; $Iu!", int~~rVt;fl(:5; i~ ~L~W!"~ i~ Iy
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dk,a,tw beneath nn~ dashoo ~in~, m: Iemi C) E:!i8lFl1lplies Qr~he ·~ng spatia:l V'alnattnm. f n Ihe-UJri 11e:~03 !l~rnel;ry
Wiig. 2. (AJ Scl1~malic phas>e,di:a!Jlrallil ICif h(i~dope~ 'cup~ates:.l~e regkiiii$ ICifal1l:if~lroma.~IfI~I:iit:in,5u~to~ iiJ-\\'I;'3'1[e S!lIp'E!fUlllalJ)(ler (d50, .mdl !l~d~!ap 'OlGI' 1Phll'i'!li)m@ft!I)!\Jgy ar~ indk<!ted, Th!! e~!!(tronk ~·dl!sler Iglas5'~ I~E:CG)extl:!nm over.i!I regjm~ in-
mhoU!!l~lof:;!ls O)I!IlC l~ll(Ui~t: "cl U!;l~L:'It~~~I~" ~E('(I) 2n. At lilignltli dop'ing:.,. d'le En:.1 ~.ii!!\coe;t iSi \.viil.lil diimil11shi nl~ inllClt:;; II)' w ilht lit!! ),,1:i\~I'i~t~l!."'fI illlS :>iiifi,c'it:ondliCE ivill.y. bIi r:l1.illIhl!~ dl S;[Ip~ar ::;on~ewhcrelfle.mr Ii -~ 5%. A~Lh(iu~h. Ihe ECG l>t3le ~.hibil" 110 k~R.'M,I:~1 -rnll~e ~nrl~ sr:in or ~hJl~.c order, SCkI.1lC cloctronicom.-eli' (If !im'krlOwn :;'I',.n~,,~ lorm is mlways dctoc[Cd at n m
slii![(; (8 ~l!jl.iJf\',t'l
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rl(iUi1!d jn Na"CCOC an<l D1H3ii2212~ BIK.a~!!Si!i!xperimefltJ'l rUJmmlilo'ltiolilikee-;jiiS m:l@gIF1!too clIld 1I.i3i~poiSltllw biases !::omti3Jlilt, lihe la.rge-ll'oirialiom are- seen ~nl!legal~~ bias and di~edly I'dect Slll~ti<l[Vgri~t~Q!ill)lr~lle· TI!; (~llPi!flel~;!!!i1r:tiOil SFi1~. 200 pA. at 6100 rnV}. As carl 00· !jJe@·IIl. 1~1,e[U(lne·:dri!diJIIlli~ stwlIlg Lv r~on.ldl ~'IIlIr llil.i@C:li~n and \l'1~fi@~ slN!liatlly. llhesoE1 phsnemena a re 'quile· simi ~ r ~n the hvo, malt@rials, maiildl B~ (oo!>iant-' top~9rill phkima:ge-s. I~f N\a-OCOCand [ly-Bi22:U
'CllJrreflt
~e
bi<u. (mi\r)i
d:l~!-lile (14 ]tJ) hcnee !h~ "cluster" des:ign"tlinn, And. lJil"("~i~ltiJ!.;"11i~[!'flm ~lllil>BeG iit:;I~~ liml hl.~h~t~n~ll~rmm'C :'i~Ip;ilroondW!ct it)' eme fgl:lS ~v with holt' ~~I~ing,(22)1 il !!\ crinic'.lll(l d!!!!iI.'lll~~m: ~"lhOlilt it is, ]H]·W it is ~j.,mcr'M()d lli}' hol;c ci01ling. ,:3)]0 110\\' ilL;\.lti~W,~ ~Ilm Olud ()pc"X!6il~ \\Iilh Ihl/;: sW!poc['Ctmdut"lin!:l S,!UltC'_ .Mut"h lISiikniO\VIIi~oollll.l_hc ,t"UPr'.UI;!EC'G l'I!lDll;!.
fJlOilll
Fiig.3.
·inl 112"!Ilm rneW'<s, of ~e~. Ilmagklg oondilions ~r,@ (A} 50 ptA. ;i31t 600 mV and (B) 50 IflA".atl:50 m\l, Th@(lr:Q.f10@ ~(I~SIn (A) <I nd ~B)ind'ictlte
!'i~m:!mcm!\. shnwinglha! the ,uckJCrllilo.cd hole dens ily "ppll~a~h~~ C:h~~l!cod doped \"I~lH,;·~ the ~}' only\\'ilcil n ~ Hl% (15). Mlu:m-S~Jn l'()tiTI.]toll IUC'Il.S'!!.reIll1.i!llbl Illl' .IJ) findg~asS}' dYH<!JllllOCS. !IfIins tnl~ C.xh!b:~l~ng some UrI_ulown rarm. of l!«¥.:l," li;~1 or'dcf ill the mn :i'ic<d~: til...: <>llll~ill (:!llIT!ll£llltm or Ihe HTi "ci~~tL,.\t'· S;ml1i~a~I)!, mlltlli,aC,11\;)1~;n\,. ... fOlll.I:Il:ir ~mLld ies show Uti:~ :;;u[m;~m'll:iilll 11'~'t:-:'iPill. ~irnmflgllcl.i:\mll" NlIckm" mUlgllctklquadll'upo!e
or
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re:lpl\ctiwly, i'lliidi lihe (lJ~Q i:irl.l1d direclio~~ aIe ~nL')j,lff'I as PIMFS (!f of1!hi(l!g()!fqa1 blac ~ Qlrro~';IS,. (C and Il:I)R DI1:aps.
take·fiI at 150 mV [L~ .• R(F, 1S.!) !lI!\i') 1[,; +11501 1Il'IVl/{{r; - :1l5tO ~Vninl tine
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kMiR
Cda'rk) m~!'IiS
ell rrrgc h(!~ct'ogcj}C'ilY f~D the tum scoJ.c: Ihe holedellsityoomlPcmcfll oflhc ECG "dUSlC':" [i'myd(.'r ~elll1l1O\l1di IImction reveals rebtcd 1Of..'llJI lattice (bl:o:Lili(m:J 1(2 n and ~pl.ic-all SIJlCtll'Osoopy" Ihc ruJOll18~OH:> i1f1j~'_ropic ~o:ndbliGI:l \~hyilInd ck:clit'()j1J.. phmmn rourl~ngs {lh fll1iil:l ~~, :nlcutrolN'iiCattenng mea.'iUrC'lFl,.:n1.<; il1dic-J~l:l (Iilm d'lC hO:Il:s are d!llslCi.\w ill nm-sized n;:g[olls, will1. semc IOnlll orl.nll,gnetw :sll{lfI~I'lI1gc {If{.!er U.~~, ~ummliny. Ih~o~}:I.'!ll: In EC I :5n.'lLIC its ll!CrV'al'iiv~. ·c,;:hibit~t:1,g ,lolHJt1l i~ole lool~.icG;.~on :<1 ~!"IIf:w:il;hrl\u~ IOIl,g-I!',mg.r; ol.l1t.'f l~~l ill mh~ 1](1111~1hd\;~i$" iSl,IPP<lrhi 1iitl!1iI(; Ul1lkm)\lI'll tOl!illl of d"-!ClDOn.k: .doJml,iul$, A IUJlg-s.!,mding flJ:Obk:n1 .h.... t; been wh.cthcu- rhesc ·dl~ls are an iUlirillsic dC'J,rR."I. Q~' h(l~~.,doped (JLla~ ~~OC!.rolilt' ilruClllre or are· : c;;:u:,insic ('ICrl1ilfl'S uiggC':l.'Cdi by randem dopm~l ;:rnd{or i:nln1rhy d~:i!.aat!cil'. )i!j'u~'1:me r lhe ClJpmie ECG ,..lillie iuc:l'udC'. for (I c..v.!Lll1plc:.O:i:s of electronic lml'li:ilalitQna~ iin. I
[UTld
gl1a.~
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dop:ml di'8on.!cr~nduc·ed
rnm]om d~clr-o:n ic g.l1!SS ~}()~, n~·~)1 $piJnIOJ!lll;'Ou.~ (3 j ~ (II" Clo";ln~<II. induced (30. }l ~£h~\i8 I1f sc]r:..o~'I,HL~i.r;<;d;!i;"(:!I!"on( i~ 11!1Il,{I-clDl1igi~~. !!lid (IV II Ibil"tiiJU!c 1I,,~~"c!t{1!bic: liqllllid t:O:Y~~I.~ of :such lii!IJiockIHil1!i li~ JU). Bul 11
~I I
RESEARCH ARTICLE
d il,'f;cl[~"S~ s.m:'!l ideas has 1l00t been l~o.'<Sih!J:c \~!i~'h pro~ll~~, in ],i~I1tly d9jJ\...,g C~~llrnJ~C:ii" f (If e;>;<In~rk:.. >1 f>1;;l:iulrnrd dlAiU ' im;Lg~. ;1~tlrolilgh \\'C~! 'b«;IW!5:C nci~her t~c ~a~·llp..aec eleeuenic slrue~\In:of Uli: IKG :;;'m~" nOr!!il<ilt nf ~n ~I].d~\'idlll\,!! d~boo.!*, 11m ,[I dir<;'ii!I~~ l;)ri:heU.::ms ('''IN
1b~ ~kh;. .n~1iflgcl; (I kl:,lt;:II'y~~'l lID, ...uitaMI;: iI11llllg~IVgaL"Chrliqlll.(_-:.; !';xis:k;d.,
'.~ I~I~kr:'t"![Jl!IIld
or
Th~
I~~\"r
Ujlllll':;~ i:!Ilg
I[)es,ign QffA~tudii>es ilil (~uIIN~DI'[o!CUO:Z(!ii! ,:;IIilid i!lr~s:~~DY'O\zta~6(U,~Oia*,a.-' ST~\I·b.~'>{;-J il'I'Qa ;)I.g,ifilg .~ligh.t ,,~ppc<ir 1U~,1ti:'ipmrJ.ri;Jr~too~ iOHd~ diC!>ll 5i,J~~~ .,....,Wi(<;S" BLHt dUd!' ilililg;nl:ig is rrdi.lgha
:;ls.ym!~l~~1!)'U'1t;;"':-'~I~twn~'" pmi~'~!~'~ a
~hC'wpm{1l~'~OLI02, IlI~n~rn;,1iybc~n
clil'l';u)'''s!~tI..'
,m ·'e.~~wor·
~'1ot:B.ldli,; :mll,ltiml to plf(j1J.k,-m:-.with :;.j,mtd:rrd dNl! r !m~!g!ng because Eqs . .2 ..m,d 3 h;lv~ .0Il erucial
WI;
(J.J) or'lh,u~ i'l~J;rrl;i!X'!l~ 1J(1"k;~'U1 1[.,,\"0 ll!lmaC'~il1g ~r.~~~tl!ri~,,~ tlwrl'ugHi 111C Jpi.(_'! orbii!!'!!> 1tdj 1Ltcl'il. 10 a dopmm N ~~+ ionrn!lye~~lsc
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~~,'C sec inum~d!m,,!y rrom E¢I,I I~m li~1.:l un· k~~m"'n dfcc~s ill tep.;:} .are ..u ('~un(:\.':k:d Olul IJy 1.lm di\jjz,;u!il!ll PIAA'"IJ.'ii!i, 'Jh~~. Z(F. r)[l!Ild 1?P';, f,') not ollly cmllrnin ilID!poO[i1.~llil phy~caJ
illrorrn:n.iml
exprc:;-sible in temrs ofnnc'llsumb:J:c qHllm~l~es only. \I\1c .~mvc 'COflfi[l1neti l.hm,IJle unkmrwll, ~a.cI!(}I'S f(F. :) JI,'C ~f'l(kx:d C"l:1:IC:C~'OOQUi iJ~ Eq . .4
, see
£l1!jllfIoCJ.I1'!m:lg
To address Ihl~ matcriel-speeillc theorctical eoncerns ,J4. JS). we h:wc d,es..~gll£\d a sequence of idc]r~l,cal TA-iul<lllling experinmC-llts ill~\\Jo rod iCllHy dHTcrcn:l ,cuprmcs:
Filg.4. {A Ollila D'" III mllp1i of IfilJal·C(CH:am:ll)y·IB1ll!2:12. rlespe~thl',ely (takeon atn$O mV fmm lIreas~11 Ute blue [1:iI0'.x~$ Qlr Fig. 3, <: and 11ll}.il'lu~ fietds. of l;Ol:eW (A,) 5 ..0 nm Iby 5.3 f1mand '~Ii!.) 5.0 f1m by are 5.0 nlilil. n,~ bllL.le! bl»:lls ln (!!I.) i'H1~ (D) ]ndic;ate ,alrCI,U of f:igi, 4, IB arrd C, a~dl fig. ~, ElIliId F. respectlrve!ly ...UI: nd E) Higherr·re~ol!utnlln R map '~';Iiitlhlne!qUlii'i!'[I~li!f1td,omanlil$ hmn Na-cmc and [ly. , 81!22]2, r~5I,PeCliv.ely(bIDe l~uEls ,o~ ~ii!!l~ ,i!1,A al'ldl [)). Th oocatiol'ls (If the (III ,aiiomsare s:hil)'i,'1l'!l as ~~a(k crosses. ((alld IF) (o~s;taln't:<!!!rr~W!t:wlJ~glrapllic ciffiol!,g@:i 5]li1IIult"!l.~I(lUi~~Y tak$l with f~g+ 4.• B <llfldl ~ r:eSiP~[~ij'i!'ely•. Imi3igliln!1l untDiUOlll'S ilue ~O 50 pA. at M}O mV .a,J'ld (f~ 5>[)pA .alt 150m". Ihe iIl'I ~Ir't«lr$ S~rJltIi aOOmk i()CClt1(1I1! r;, !.iIS~dJ ,als(lin Fig. 4. IE! ,glnd IE. llhef~(lld~ (!If \ii.~y (If tlte'Si~~malges are ~holWl1Iill Fig. 3., Po and IB,. as oran9.!i! bOii(lls. iFhiig• 5. d(A) Wr:I]~50 re~~e to l· e o ,ilIDI. IW m'lIll'laI:l),1l'!IIIiI@(IlIO;Z ~[(!newhere ea(h ,dlt~'V :SIfledtnu'!'! <1,1tI1:~~mral(~ ,of Fig .. 4,. (and F" : ,a nd51hOllWll nln~n!1l, S!B, is mea·
~!romgl'f U ndeljdr)l~cd C~" I.SSN<lll12('lI 02C~2! ,(N,I'CCOC: cri~iclll [C'!llIl'11C'r;;l~!..Ire Ie ... 1'~ K)
(D),'.,B i22 ~2~ ~ As indliclII,,"'d ~dK~llIl;Jl.i~lIlllyi'l'I A g. 2. B <lind r:,lht"y l~fU\'C compk:ldy dilT~relil cl'ySlat. lo.gmphk SirmC~lIre, chcmkfll cO~"~lil.lle~u.s..,fLlild dO;Piillt species and sites h~ Ihc tcf:in~nmtol!, I~u)\!;;p.\ .y]ng OOHV1.J(,'ll the CuO::! pl!ml~ ilnd I.h~ ~ stM. lip. N;}~('C'OcI~,m~ :sing~c nLO::! lay~r II mld
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A,
if
-
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o~
:$ur"€d. Spoctra ~r@ mfH!s,l!rOO along l'quhlal.ent lines laliel.ed I. 2, 3. end 4 'iJil ba~h domai!illi lof FiJg.,4, EI end IE, ana Fig, SA (8) Difi',erenlial~UfIne~ing (On·
dliH::lant~~pe{ra ta~i;!nl;;I:l>01'19 p:8ll<1llel ~il'll!:; li1 r(I<I.:I~hi eqUliiv· ,al(oe;nt dolmain~ ]n Na·CCO( and
I,
I I
,,
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1
-
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1i.l~1'I l!nrnrid~nIiGjt jll!!1Qlikm w:ndlaiofli5 (200 pA,. 200 m\(lt Nl!fIiIDm (l'l'o 4) 'OOfi'€sp!i):nd 1'(1 trali~(tl)'rie~ wh~~ lhes:e~q,l.I~n~l!~ 'o~' ~~{',(tra \:toere UI kenl. L(lc~tion~ of th~ tr'.:ljecUJ'ri!l!s, reL~tweOO (he dClt!lai'I'ls..aue's,h!!J~m ~'lwe~!l F~g. 4B (0 II IIId 4 IE. by anOWli.
1382
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SICII~NCI!
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RESEARCH AR1ICLE
capped b}' ailcr:rc~!
P)'~~i~.212.h,~'l C'~~O~ bil;l)'(J'!', ~1;J.o'ii~whi~h ~ ir:!{lihl3iO !ll~~ SJO ~,IY~'fI\\\'ho~ ~'lit c~Us uu~d~QJ;o t~li; il1cmtnllCil ~u,ruR!~ ,C'r}'l'i¥i.1 :~U,p~irrllodlJl~:~li[lo!l~ non!':~oochlGlTIc:!:l'ic oxyg~1m ,onp<lnl OJ!to~u~ <I~ IOCOl~~~d inb.'r:.;~.~t.~'llIy !1L;<,!!t ~hc ElLO
,I
and D'j, !:leN Ihe R the Cf)'SI,;I!mm:: ~t~'JoCiITI~Qdl.!l'!Jion • .figmt' 2. C and ~illc blue loa.x es of fig,;;:. D, liP;: i,n<!g\~s R(r~. f" = t;'{iI' mV)~~t;i!_~\I~d ~n;Lp~,,~ r~,<!!~cdm pu~ a C\IQ<ll'l~;;l, .m"~hliJl:'llhe ~111!~~ id~J~~~e~,1 f~!'d", (l:r'J;,~\,9!- .r-~g.~,A .,t~~~~ ~O:!ll.. i!] .!,1Xi~o;;" ~,n~i~~~. P<'-~n;. ofd\h~ lin~ .13:, Th.\l ~,~pt.,".{;'Li\·I,;:ly+ These R nltlps i1n.;: !.1lll1rlccl~ silHi~ar {r!Jpn.:~t;mi~lg Il!.gh 'fA) i~l lli!ap~ illdi~..ru,xl by l? in tl,~X~U!'G mdti.bi'~.rat iil~~'r Sf1'1uiial tk'I[~i Il> IIElIIl 'lh~ arrows are pDL"'Ci'5dy 4a.o aparii mt~~ocP~"'L'Ul :inK! ~hcir l~lat1.:'d surfaee [O.l:mg!'ilr~);';i: th~ ~l;;~"0l1 is ~·.at ~hc perimeter of <II sin,glt<: Glnnmin. ln F ~g, 4, j\ ,nut'll o:t' Ihi;ir (;'Jllt1:;.:o;;~ ~~~"H rOO:!"l fC'lllIll1.;"S amI! D. ~~lul.'ip:t~, p,u.II!'d. "lao-wid!;; dor:m~i!1~ rUIl
9"
layer. TIi~n;lhrc,."'H ..~scrt [h1U TA- hrn>'!,gil'l.!!l ph(.':i:iOiil.'CIlU [hill H re ide!]1 irn I~I] t]l(:'5C n'o'o miuucr.i.. J s lli1.O'li III be 3!iiCR'ibt~,1.0 I.h,cir ,onli)' conD· ~noi1, larne'llCfi:::l~C -Ihe ~Illii.in~ie cl;~clroili;e smi:e~ d
.~ of dtc' C~O,\liJ'i~al1e. i.Ire Aiool1lllic"r,@sQLutiol1 'mll,j lti'agill1lg. ~n F'~g, :2, B and C. \~' sho., s:~nd<iI1d ,r:;LI'AWsllCCIIi3 :me3.a ,~urod under :ld(nlOC,al jnclion oondili(ll~tl O!:~ mndom !OC,M iouls on the sl!ufm;t"£ 111~i'~"H:'CO'' 0tJ1d D}'·Bi.n 12~; mlll~~ (,~nd''!'tOl \,\'CIiC "~w!~!li.;'d .. a :)It 4.2 KJ. \1\liuh,in E < 1O{l! 11f!~·V.,I]I~)I beth C'xhibillhc C'XI)l't,':~cd V~slm'~ dllell' t,':!\;'"n<C'!'oo ~m {£ = tn 16. ft U!W';'i.pctwdly. m hi~l!ci' cnl.:'r;W(lti,. ~h(l :';!~TU.,~ i~~'ul)l]~ :;'Imaia ~ \'mill ion ... ~Ulhe [1iJ !~~ t .fI,d,iJIg ~~}' !l!trn~uy u r ilpoI.:C \1!(;l!"C !}b:;;~[\',\,;'d i~l! Ita b{]IU~ um:m::1i ~lI~S~1i'Ul~'C C m! be seen V i\l.udly m the .I!e:ll-.l:nlnd p(:rimCh .. ~r Fi!!',. 2:. Brund C
t)~(;:~lI!ri!lg
w~tlli~
1."Ileh
1./q,
or
A~ lUOl:ni;c seale .. we rhcn 'sec an clcc~'ronic snncrure C{lll."'~~ting of a~I~!lg.h clcmCl~~z;. i~o d strucmre, in To 'i,!"'P~oro]~lC ~~ .. ~u:1i mT1,1allll~~~li:!li~S. or ~h~,~C' [rib,~~~1d [I diMl:rdcrcd I~!\.hio!l a!io.!1g.bo~h ('m"J() p;hcllomcn:!,wc' USlOOan a~o:mk: .l"C'so.~uli{ln H'R d·ircctiolls.With in this I'IIL::!Irix. arc eml'll..'ddcrl4ull'mIUlI'!' spatia II~}I ]m<lging 1?(.f. I..~ Eq. ,..Jib, r wide tlll OCI iroc~i()m~1 regimlS or ''dom;l!i~ls:' 'FllCSC 1'i!\."C·OllISC !.hC'}'m:tt: Ill'ft(xilic ~lIO]~g the F~Jil'llre 3, A m~d B. :iiho~'" ,ypic>!il ~(III]11J".gI'l~')h~c domajns. ilnmgcs. or~hl,.;'OI('IOInld BiO~'I)'cl'l\. ohr:."L:illC'cI by Io:!~g axi)'i, aPJ~e;lr 1$1be ()rdC'~oo- RC'if'l!C'iL1.in~ 4f1U" wid~ do:r'Ili.!~m .of lhis t.Ypc :II.IL' ZI,h,\I'ay~ Urrl~d(;,f~\,'~W;. un cl)I'nJil.~nk lIl:tI.,~hi.~h vutun:r,. of d~ltt'litmal. (;):I!Llld:un~ nloll!;!; one or ,[]lhcrClI..O Nn·C( 'OC ;)IHd By-Bi211 ~. rt:,il;peCE i\,,,I),, niL\' dil'OOtiAI't 11~~1l'i" the mn scnlc~ tllie c:lt~coroillic :n it!<righIL';SllfC,1J,1O\I'IS in Fi~, 3A inc!k'3t~ t~le locHSIirILK'IHOC ofl]~('S\!: Iighl.~)1 hO\I:~-(klpt'lielllpr.fu!c:;, I iQ,ns .0 f Clm~)I1i1s, lhm mire d ireel~Yflb{J,vc I.he eu .'IL10rn1lSnn Nn·CeOC 'WhCl\.ilS thos.c in Fig. JB Il:rea.ks bOI.h e" s,ym.mctU)' fLltd lrnn:sJfloim~.'IiJ ilildicatclhc Bi a~om~~tluu 1lireobov,e the !L'u S\}Ilmn:n:y oftb.c id~,~ $qlmre crysla~ lattice. J:n Fig, 4. A ;!Ind D. \\'C: 5iI0\\' hi,ghc-raliOlns ill IDy-Bi2::! I !, The dark cl,'Q£HI'Wpt.iJ ,uc'" giom, ill 'Fiig, 3A lttl! the mni,'<sinS n m.OllJS and. iu [lCOO!ltJllioll s~udies of ~'qllliv:llc:tn.dom'lins II,rom. f'~ 58, ,a'Ire di~~m;..::d l3i flll:milS alo~~gnnruo;hnaof' Na,.C(,,[)C iLifld .Dy-l3i2:2 I 2. rcspccllvc~y (at d'LC
1')1,
procedures nonualise [~Ie lmegrmed dOd!' OiU1!h~ pesitive sklc}, The: eorre,~pom:ljl1£\'arimiolls ill Ti\. iI~djcaW' the: cxlstence of inIClto;:C' atom ir"'.'>C:'d~C varimiom, in c'lI!CtlXll:lk
{because eur
or
dli:ib.ul.UI'ig vi !'IilIal ~y idcnl:ic~11 i uuC'mliii. :>.tn.ICllir,~ illl ~Lh mli~cr.iillJs.. \~Ie- next C-X!imin~~. in, :ritl. 4, B Ilind E. ~u: intennlli ~~1.ruClure0 nile dmn~,ii1i!l; {a!tth.e boxes of d~;;~I:tICC, The moS! obv[:o~s:l!IlJ In'll~bl)' n~O'.i>t Fi~. 3, .1-\ ,muJ B. and Fi,g. "I, ,A mad [)) wid11 ill1l~rt2lnI ab~rv<l~imlin Fi;, 3, C and D. f s 8, loss iCicm.i.fi1;;.nion of :alma tc si~C$ 11\.1111 :;hnul'.Lthe 'Iopo:grnph~'~ Fig, 4, Cmld F.I)(.~I,':ICrtthely). (I r ~d~~lmm:l,niol1a~ and 'C)fJl-ro.,m,tiol1<11 lC'.~I ~1~'{'Il~S Wi;! sl;"Cimlltf.'{li'!l,~Jy [11lo.1l the Pl'ilNlj' ~Pil!ll,,1. i!'l\1!JI',i,mocC' Ithc~I""~i,d "lIi'''L~g,;,:mcn'_~ [lof clecin [11m Lt' ~fUlC!UJ1C' ,;;r~ the' lIan ~("dl,; tht"$;¢ CITI.1(i!ii '\mrIDtiO!l.';; ill thi!' nlOlgl.-l ,,~nt'COll(:~!larr~t(ld !lO~ {lit bC~jl~ \'.in.i!II~I~~Y i!1d;i!~l.iJlgi!lli~.1[L1.d{,un N;J~ TOe 'lhJi.l('~ 5:u'~fi, bUl rather ou ~le' 0 ~~h;: \'" ij[hin ~..u;h and Dy~Bi22l2.h is atse evidcnt Fv~. 3. ('U-'Od u l!om~. Here! the (j@JniJ!~n'l>!l,)'1nml.:!H), 011.'.;.1:> ih ,,~t'IJ~g~,'cl'l~!:.d i~~C' a I :>naf1 ing .n ~hc..rnl\h1 C mld D. ~hii!lIh,(.~ ill~~~'Jn,yl h'UClW!fC S ~h~\'lC IH'b!.:;~d L A long lhir- <b:.i~... aro a li~l~ of U,XY Mi.:n "dOll111~11:;::' a" \'Ii'L,;'11 ~.II': Q\, ~.r..l.nn1~ul'!'h in 'i'\<h~'Ch ,'~~ sil(..">, 1L;!lcl~ \'\.'V~l~!] a hf.!.:ri-.!lnll~1 C'u-O-CI,ii blJVld dlC)' omJ t;llnhr;:ddcd, r~1<11U!!1S rl!llrfht~r l;iI('llf(lt$ of :lind [In (~~h:ubil.ing, higi~ R" nu~'vertical Iline dl'ClJfm:li(; COlfup1c..>:il}' 111Lh~ ~tQ;rn,ic ~C8iIt.:; .. CU,-O·CtLl b®n!rll·ce,!il1!elrre<dl el.ectmlil~:(. gl~as:ll !abe.red::!: Is. the llne ohocnical CU:-JO"·CllOOUids: Inng-.rHl1:J3IC :~piil~[11 ell'det of lilt)' kind, N,:vc'i:'il:h.~.I~:!:,::iijLOOOfIii.:Jillio:tl mml,~!\il;:'; .. :ihu\!f.:,;,nht'it th~y do :lUi\ c :il~orl-rnllg~ - 4fin >: .;kIt! pcriQdl1e:cmreh.llion.'>, w&aerc (~I!I i~ Ilt~ t'u·O-Cu
~ro.~n
.or
with (iisp!i!"rlilil 41'1o'"wid:e dOlilllilli DIS. TO. \' isu~li_ze d'LCSe: SPf~lial ~k'llllCJ1~Smore clearly •.wi:'mkc the Lill) IlIldl'm~R of Filg, :;. C ~Ltldn ,~Fig.3, E ~md
Ih,cSCOXygOl1
dincrenl
sites exhibit law.R. Tbus, R is very For the h.Qr~ZOI1HiI Cu,;Q·Cu bonds
~ end l~leVCrl.ic<l1 ('uA)-t'u
li){lJld~ a~ont .liil1c 2, even thOL\gh IheM!' bond s share a Cu m:Dln tm the' lDl"'lrner of the ssme I1J;qIJl;lU.~. ·n~ !WM W.~r!~C<!I lim: .of ClU ~1:O~t.. aW:Jiy ]rolill (ile: ;[1L.xi:sil" .i~bekd J. and Hille <I , ~I'Im~Cnli5:lhc:line ,pr Q(li.ygcn Sttes IlIm. i~lalll Kl the right of n.xi:s I. Th~ :iiOqwm:;c or hor~"'onw']
I
y,,,O,,C'u bond~~ ,Oll[)n~ linc 4· cxlrnib'j,hl a 1I11~· tOCl'.Iml'yffJ'w ft Th,,'S~ p.tIIU.c[11S ,t'xn!ihul. lim~rmi.
Filg.
,,_
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;]1:u:DI!II.~lllC' n iC'a I .Ixis I rtlt'~1J1Uirl~ w ohm.the vd"mh: t!U,llMUI'i L~predsdy ol;(~owick \lIie U~IIld.dle.~~ lunimxilll.~ drJill,KuinsnnriJll R nnl'l,p 1IIInool1ml)' disper:\odwil.h cqlm~ IProoohilit), of orh::JlWi,t:ioll aiong Ihcl'>\'O Cu..{) axes (I:jg. 3, E and F] and wh'lll v~I1IL1<1l1y idenlic:::d snru'C UUI:e in. ooth n1iili!terr..Is.. .t\ 110tCWOro],IY obser\'adon :hon;; is Lhal d1<: 0, shes wilhin (\l~..(·'ll hom"". even nhotlgh symm:cq.ry
~S~! randi!J~ly m$triiJ1.dro el.eCilrel'iK: variations oUhe (iltO·Gu Ibond slate \lliith e~llal prob .. il~lily of ,orr;elit:a~ion ~kln9 '~!h!!tJ.\II:l' ClIl"O.axe~. File eu·l) bom! dir@ctions iI,r!:! ~hm'llmas pairs I(II~ (lrtllog,onal black oil nr-o~~. lih~
uyskll1ogl1'Iph iicall>' cq;u iV.Jlcllt.mt:'i!t;l ,elco.;::U1OJlrrcally ineq,l.liv;:dcnt SlalC$ {F~gs. ,J: ruKi 4}, In,gcncml. Ihe ~ura~. ila1'JIJ1;,Tcn1C'Jl~Sof Ih~ ·y·~a.1 oond
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RESEARCH ARTICLE
cn~rgy value at w!lkJ~ tl:ldl' h"C}'1i:;m~r!Cl> (IiC'C"1H", To d,Hily the 1[H;;''''I~Q!l)l, ~~!,,~~'rc the O>J.n,g to C~I (l!"hi~,I~Si~~ C~O~ rhmc, nf the ~.ftdJ' sr';:!i;:~IfI[llI~l ~n~a!\tLmd C!~ till!;: SiLlrr~~~ll of .r~g-.iI,. C :lIldP. w~ ~htliw II :.;dWllr!;m,til;:' ill Pug, SA, The dHd r sji(;cl ra .u"(; !nc'Ii.~In."d ;dOl~ cq~li\ '~!~!"Ill!nc;s. !.!bited 1,:2.), and '" i~! dOI'll,lill& of fi£ .•l, B boll) <!I:l'idEo .md Fig. S:A.lli~ Illili~i!oiif!o;;tm la In iJ 1dnn,g
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lq'iCl'icnt:i! an ~:l\"i.ti~l,gQllp.oJ:lt!H.Ul)'10 deteemine !he il1ll,."Imli dcC'~nmk' :iit!1!!Cl~!'C ef' a 1;;1!Pr.J!.tc ·~Ifi~~i.(' {Fi~. 4J flJ.!r~~i;'lmlfliro, ~~~,ni!fo!,l; tlr dit;l o
~m~iisit;nl \.... idl,lht; "ch~I!o;'r"ph~llOI~.W· t ! J. '::)'-2S~,.[he 4(1(1.'\I~dc d'L't:ut.'!lilic dO!n,jl!~n.<; ~d~Ofqlrr.~~.'Jll c",cdk'!~1 !;~!"Id1idaile's,'to. be ~hc ~bkIU~IPUS ··(:hl~[L,.'!"8.·' of'!hg; !ECG ~!.1~!e. F:illillJ~y.out ~~I'I1pl~1~I!jp .¢:>\hil'l~1'~~W b;:ltU(1'U~
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lint~rlilbc~ed 1 ,~lJibd4 ~ItC l1ik.cn ~fl ~ ~{-qIllL,"iioc tW,'oi': fil.lnfillli'OKyg~':Jil s.i!Q;w~~1'i IOllll"empty s.ir,,""";i; .iul !tii0lWQenl. \\Iho;;:i\.'a. l.ho.'>C.aIDi~~ lines 2 and 1 are !ak.Ci11 II s~~,lleH~ or ri\'~p!lallar at cower sites wilil four lI,,'\;.ygc~l ShiCS in be~w,eell. In oo1Ji~
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~mJ1i,gJ<I~~rilil!!!il!. .. ic ,!lid Wll1i'~~~1~110 cupr.lta, \¥n~l:td pmvijl; ;J ~jll~HI~ihl~ nlcl COI:i:si..~kn~ ~'ltl~1!1. .. llaLi:O~ll "0:[wllil'loIllg •[email protected]' ~P!lnlCtl~J'gI,,"«.d~~t1d-w ~~e !iilllpo=roO:lidi!lclivi~}'[[0 ni.g~'uly hnl~·doj'k;d. 1 ... C'Llp-m[c,., ll1is. eeex ii>l:; :;I'il:'i:~hlJUy \... it~l Ih~ Irl'tJ.j]d· . states !Ire i'im. J!.!lcct(ldm. low dOpb~l!!.. Ttltln.,;:J'ii'l~ fisymi'l1ct[)' InC'r~ mcmeI1~:o; .lli,re rentcrud IHumelin!!f ~~inmelry paU.cn!$ ,d~sCi'i~d. .. hell!!. A Il'rio:l'i.~he TA. Wnliil!:ll, repre:;ci11.... vllrim,dc:s.ii:nC'd K'i yi;{!M .kC:jI inFonn1i1t r.o,n on ~IOO' c-.orro·
co."!. \'1/ hli~ in ,C'O'!ch -k"lo"widc '-.klmai¥l. ~~~. IlDijs(!lI~$iQiill aJliiliBl ((lndl!ls:iQ!i)s. B,,;;c-'oI\ISC \\'()' 1~l1d ;nuid~y ;dl;lll;q~11 ~,hcrmmr;ml .~nN,.·CCO(" ... a:lldDy-B ill·~.2 l>;lIlll,pl!l.<> orL1tdil-~~l,y d1inbi:\:J1l p.~'~yS;cil.l, chcriliiC'J.l,~uli! dOPii~l~l ::>WC'~IilIIit', U\ill1.Criil~~ specili;c cxpllaalaticlITIs. lJ4. 35) ror Ihe 11\ ,~nCCI. ,all b!: FLlk)d 01it In~t>Cad, we oolls~idC'I I hc..",c rhel1o:rncml t\) be- Lntl"insk c]cclro~k clmm'C.J;clI:i~~i~ ill\.: Cu02. rhiIlH:, fi.l['b.h..:ililmk; I,~I~ 'l:trQCDS; ,or rcpoii1~d h.ere CmllIDII. ll>!.:gm~rnoo by inldividllml d~1POlnt ll~om.,> OOC<lIIlI:SC (~Icre is m1ly .. :single dop-ant <bWnllor evcry-~.o C~-O"('lIIbonds and. in any I:1!.SC. 'they oc,cur at ,quite di fl:errenl, iocat uons in ~hc;lurlil celli> or Na"1 '0(' and Dy~Bi~::! 12,
a c:I!c(l!-on~c. Imsn(ll )Ie! been ijndt,.~1iC'I!dI!r!~ 1;::,,1 · !h~ :oiJullcrc(tnducl.i ~IIt}' cQn~~,~I$ dekx:: ~i;lod ~y i b~,IK-d lor the 0:!J'!!l1\l!.Oi uf oh.trgc-dL;'.)Jl\iily vO"tll'l• d·'W'i\'~'L.lre,C'uron p~~i¥joi, HO\v ~lIei~ dii.~~:1rO'lw er,;" r ,llI'ilfw clt)~]i!!g, ;Jl'ldi htlw 111..: flll,n, I timts ~[:n.·rTt,:~~I~ts'Ln!d .~I:S:\l OOllwin q 1i.lI,IIUllll'n ~bell);;;i:),1 O(tt;_" i,,~ :ina)' IJ~~ li....... ion (25) "!;;'<SOCf'H!:'d wid] h~lliI.·s~~ !ol(<! H h~!cl'li:r-cnc-c e IIbC'~!'l'13_'1l\, ~n ~Lddil,ioll!. ~h~ relmiun l!cti,vil)l e~lL,.'L"'gL,.'~ :fl,'om between d'te higB~"l~\1lI!~'l,g}':a HI.mP$ and dlC 10\\>- !L,.'nl,l~cr-a!ll~ro!ilu~}Cr-CQnd !.hi~:;nu.\l \'II~.J~im:roa:;.cd ,clopillg.~nl'~ ~I]to b~ C!~L,,~rgy l.Iifdl' In!i!~l;;' '~:i.ll(l~mpk~d}' 'Ulld~'rntot!~l i For t;xan~Il~~.FoLi~r ~ro~~~~mrni\; d,(id l' n~I:IiiIgl,l:;. d~'~ml inml of :U 1£ -e 50 me\( re\"'l':n!1 ",c~I\"'CkC'rLlo[IIRI" ,cl!lhU' ~e:re~e:ri!Q~$,a N,Q!tll'~' ifld ll'tfld~l~rLl~O:ifls wil.~!pcakS. m if ~,(OJ/4) lind lJ/il.ln :l" ~. Z~m~ G.¥!" )iU'!!Wty, I, W. AClen.. Phfl'. Rtlv. ,~l" 5:5. (6). TIllS imp~l'CSIWQ. at Ilr,cseJ1l. inailsti:l'Ig,lhisJ'I. 4111 (l'9M}, a ltiil.c.pos.sibi:lit~es :aMnJlt the rc]miobl5:. between R 2. r.. i. (l'Iffi flrll:, ~.t'i~, l.t'n. 66, l04 ~l9'lJ. 'I" M. II. I, M~nmi!:, it [st6, 6. kSa~~.PIl~~~,. 8 nup=> nnd dlfdV ,rn~ps: I{ill~hc pl'ljJsiC<llI crni,t)' 0:1' 46, ~'9ll6 (l99n" the l'Ionrn-cclltcu'C\l domJin~ Ill;~yill:pI)C'lU' dilTer,~" P,W" ~dtr:s«1, t.I, 1'1: 'Oil!» J. ~ Clem. SoUtf~', 6 cnl,~Y in tlilC two C'xrcr~l~wll!al ~u;mlitic~, (sec SU]l>1\ {~!)IJ~~. w~lh .. clt.'t.r K'dlll(~io~ oJ dJldJ' tOW<lOOE = 0, IlOr! UV\gCI!1Jhn~ lCXI I): :lml.ktr~ .~i) !he loc.alifXd !I. M. lii.limdl!!rna. R. ~n!11lffln, N. TrlloMl~ f.·C. 2ih~ln~. Fl'I,III: 1\ll1l. ~(. 9~w 1301001 (JOIMo}" dckl!c.llizcd lovt.i-cncl:gy Thcsc~ypcs ors~11 shapes. and tile di:ffC!l"'C'Jn de<:trol1ic domain ....L'lffec~. 6, t Iia~agiltrll !.I[, ~L.1M'~ru.rc ~;'!I(I. ]:001 ~20[J!:!), state-so po,>;;s]bly \I~,R ~<UiI.tl'ing ]nK-rJ:G'II,,'ncc (36). C'11Cfgy r.mgcs in srunlPk~ Wi~l di ITc.rcn~ Tc:. n~lay ,. K. IIlI!Elmv 6 Dr", PhYf,Ill'll'. tN/.~4, 197,Q@S (2.Q05),. But ~ndcpC'tH_!cm or Wilich Cx.pbmllio~J ror indi.c.<!.1C njlC'1r K'IJJt!1111 IOdre :illQ)cJ'C1l'Indl!lc!i\r~IY, !3. y, J, !.Jeo,~ril~l ol. J. ~(}Ir;tJt{,) (oMcq.: 4~ 2m1i' U9aal. '·c::hl.!"Cik(.'!hoalfdi'· dNd] " unodllhllliol'l~ ho!d:lt, th(,."'!I.C 1~~!ilg·r.gJIII~1l: ~le:~tlr~:n~(ml!;l(!I:Il~. Rctumin,!',l 'it D. It. 1f!a1rl1A:ii1~iit I!'t uL, P'll~fWf. B ~I!, !!5i:l! ,(US 00" ~o ll:'~~ hll:g~~ st'IIr;;, ..,llig~, R .[rm,? ~hrn.\fll1inF'i!i!:_ (l IlleW 1iA-nrr!)t!I.!ililJ!.~ l()(;Ill'I iqlll'Cii ,~D(: \""1I;:~1 dt)ril1~d, 10, Cl'I. I\iJgm~\r t~'at I'l'l~ ~ .i.l!.'I!t. 80, ~80113i~'l'!l913). U. C. f!~~~u1Ql fr.' I!f,. ~ 1M; ill ~'6. 064150:1 ~:tlOO2i. ,~~)::rJ:'i:;; II 25.m!lli'II:! Irul[~M of vut,"\v" No IOllJg.t~1.I.1~\\' 3VOOC! Ih~ s;y:sli:\lli"-l,tiC' ennIS liuln ('(;.:::,i.l1 dlldt' 12. A. '1'., ~sadt:1 6 til" i!t.i~, oIIt;v. .i!il 6~014S 24 ~:lOOZ~. oruC[ Gilirn be (k:I.t~'lod (C'I~1l fiLiI 10 50 mn: ~lig,. DI'iiillgiu!g. n.cCi.."'-" l1ighcr.cllcQl.Y sCflk'S~hmiFCvi. B. $, !iilnm~. G. J!!.lod'l. G. (on(,}j;, A. 0, Hi~lier, It I)e RtlUil 53). Bliltlhc fourier lr.lillSlorm ofstl.eh l?: n1IflpS 01l,'lly. <1I,l1d l1'1:li1y lTC' em] on~iglufl!!llf~cw ph}'5iCl'l.l My,>. lI'e~~(. 93, 207001 ~OI!), inio:rnuuiml (1 5~. I~ :imctto .F~g.6) .iI'C\'Cll~s rlirlhc r slu~r1L'>Cs: The 111- ] .. H~r.e.. c. I!'. 5IKh~[. (.t l'IIilllng. :A SrJjl~ :ll5" 3:3.9 m02~, HCCiiluse t&le TA imfl,gG ohva)'s (;m~t3~11 4(10' rrcdQmi~n3nt rell,,,,"s occ~lr lit wa'l"C' '\lCcwrs fi ... 115- 1"1.'1 Gt'~ril, I\!I. ]. (u~ M. ,fIi:i!:k~r, I! Biii:litt1~Ft. \\ ide 'Unid:i l'II.'Clionnll pa.u.CI'iI15 • .fltl o'hVUO!lS q I:IC51(3l~,O~ilnd 1O'.3/4l (il1l unilS ()b'~1itlfJO" WllC~slh,c ~J1. fM'I:. ,uti. 96,. 017002 (21DOO), peaks RI q ~ 'rl/4Jh '-'lid UJ.l./4) 11131. w(}uJII iJ.c [Iron is. whcd~cr t~cy arc :SCSHll;,'1IUS oft~'tC cilargd 16. ]" H, ChI), f. Borsa, '0. (.~OI:tn~oo. I!l. R, T'Df~f!':I!(!fj, C'xpcclCd frivially rrou~~.3 -4fll!! ;>i 4(,1'0 m{)dut~uiO:iiI spin-orucred •'san;pc;s" (17 ··41). Il is iJhtgucd rrom ~~ !i'ev. .tI~ 6, 31'19' ~l!!J'9'2L :It:;!\ F. (. Chil~ II!!' ~J.,j,~. R~: ~i!l1,.!!'1,2;lZ;! ~19'I1;!J <'IIJ'C w~kcr; We I~ndtha,t ~hc:ilc dmrnC'lcri.~ljc TA, m~u.tron SC<lIIc:rin,g tht'lll 4flQ"wide Il.iilll d iifIC"Cti.om I 113.MAil. tuHtn li!t 17l.•. ~fW., ,t~t/. 8~.,~O~ ~U'!ii~~, UmXl\IJIJtim10 al 'i - {3t4.0)3I1d tOJ/4~'Ot..:;~n· nOI l(mg.-I';lIl,g.C ell moyc 0I1k:l' forms ]hc 'b"", is of sl,mic 19" P. Me Sin9sf, .f!.. W_ IiIunt,K Im~F, Pnrs~I'l't!¥. l,e1t HfJl b«auSIC mL\j,lIlgbctwccn I ~14,,0) and II. ,O}. bl!~, :>1J'ipC's H1" ·IS}. ('on$lstCI11 \~,'jlh thispl[cwrc. 0il'6~ ~2o!m, p,rhnruii ~}' hcc<lIJSC dH~ 1l1!IIi'ilima in R typlc-alll.y I~S'OIt"II~ X"rny .~aqu~ringin .La.I;8i~Barull:f l~Oi! 2,0, R. mer; S, ~eo;y. A, ~iIOigel, A. ~ftl'!,.Mys. iI1'll'• .8 ,~,
,l..V!,h~bi~s. nUl;:'~II!,i.l:itl~. bul h)w-cu~-rg)' :1~";~Dur"\'-;:,; ,~iuni~"rIO~hOt1c lill,!.;' I: I lne ) sh(lwsh~ghl.:l' TA: .1mdline 4 shQ~vl' ~lC' h ighcsl. TA ,,\'i.h wc;d .. k~w~ ~'\!'Icr-gy fl.'<lU"H"l,;'S-,These ~~;J;t~.cp!.""of dlld i' S~lJt.;'(:1!t<I ~'.:J]ibil!JnimJr :.y~lllli!"'~lry "bout ~he Idolml;lin'~ \.'l~i:II.ic~1 ,a,i$. All spt..~tIn show II PIt'!I1f1L1Pl~-d ~~millre tlt B - +WO nmV i;.... r~ m:~lmr,d fefrturrc i~J~ ~1C"'dl"E ~ HlOm~V btu ~n~sk.oo.lJy Ihc ~pidl rise due 1.0 1.~nlncHnlga.")'lll1neIrY. The lm~'.''illC'.f.'gy I:e."m.lrcs., whcti~~r or ~'l1~" (lines I .,md 2) or shoulders ,(Ihle:!> J J~[J 4), ~l~l'cnJ' .~ ±IO> meV .Iit for N~("CO JlId ... , 20 mc,vror D!l(~Bi12.12T
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:;;udl J.lS AI OJ!ald("l~. nm.:;,;,imlUTI! ~'l!C:lldi!lg ,1U~lill~ 1110 \v:inqg.~ik~b;~lW: ~n ~ l!."'I~'i!ldOO The und lUI~k;!i.~ (krt1rn1,~in!l. "'PJ'!m~ht.,.'>'i '90'" in btllhlUidli ~'iIJ rb1.(...., SWl~h 1~~~l'!cr'~D li~~r:.:,iin~\'(;r·in(;!f(,.'<Hsil1!il un !I!~.!:ll t_k!-J;:It'C.\1 ,l:;Irpli!!."iIici!y .<JJld fk.."X.ib~1 w~\re Aft~ true p!'l!."IUC ,.;lfilin ·of -I(~" lit)' a high i.:k'l""ily (;fr l'1l!~"<!Fbands I." fO~TlliCd tFig, 3B)o, 1i1l!!.':BMG~ 'lfC~'aW!::'iI~'obll."li ~~~'(i BM'Gs in. u jR!~x''l'CC!Clk'd i lor h,~w~','ol\',~"d inll:! 'P~~yc~~I·likJ..:l :pwu.'l1'1l' WiDh l:iqllDid .n :hi.!lJlh.iI!lWK"r.ll~llJt-"'~I n s!lK'.U!lb~,~di\. $~n.' ill_g,:1('l "gj[j,iulbolJ,ndnri~!:-;.'· As. shnwu i'L~ F~!!l. 2A. Ihe 1l('I'\,~'SIDl,'SSof S1 ~n1(lf:~~(l ,[!!'diBks ,,\'!:fi::' also tbund, hlll.UQllC of Gflerca&cswiil.iI inere....:-;in'lg suain ,~n~rucss s mhli!i1ln~.Jink-d ~IttlilghdcroJ']mu~i!ln to e::tLIS(,! speeO\CJNillt'!OI. I{[lYwc\l~r, iitc' D-~upidinen.'fu.~ of now il1m~11 rrn~lLiiC. n(! SMG::; seem m dclQrm iI~ I) stress ('!\I!ten stlranl:l. is ::>~O%} is not due 1.0 tlile microscopy ITEM). and :Ilig~l~rcsollution T.EM 01 RTEMl. Elm,tk:modtrlJ:i or Ihe- BM'(js 'WI;!i'I(!' SIl:aidl.1ll9fflening.Wc eondueted com~~i"Csslioll rllo:railored by me'.1IJIS or an lI~l,rm;J('flliiC;method tCSI$, lm S2 ;l]ru.! ul1itom:lcd .[t di lTerent l'Il'i~~k l$.-0-1). The llamr~c:swi!h ,J ,~1!lIgcaspect rauo or sn::linrs, J\sshm';'ll in Fig. :2 D. !hc ~1~p'1C'S' h;lI'J\cl, ",S:~:I!:x1o-'1.s,-' 2:~. were cW Otlll of [Ire as-cast 1~mm I)nlils, lor like Sh:II:lC'im:pliCl': th::llJ fric'lion occur:-; a~ the CO!~[;;!!tl ~urfW;;~OOl\,\'l."CI~ d'le' !.'i<'H~p.~ ,md ~hc ~! iaxiul COOnl)res."mm.~c~a:o;. n u.mgstcn carbide platens (lKI. The ni'tction \'\lo~:ld Aramil)' or qururt:cm my Z'I,'CuNiAl BMJ]s was obtained with ~hc (iaJHIJ;O~ilions shewn in increase Ibw s!ress. ,c;spccimJly inlhc law Sf!1lgc of ~l green a.!W H~' f,il@" l A .l\OO1I:s,tiC measureU: dcfomml!(m because of thc:la!l!.C iacrease of tmllj;.\'l)T:'i:al 'iro<!. of th~~[X;'tJm~~!l:;mil [hllS n,u; ~lC1l1~S~how~h~t some (;o:]llf/!l'lsil[iOl1S. such as decl(."1lscd :l'L'iiptlCI. II":11Iio (J 9), iI1!~ hlllrdll.el\l'i of tile lrililll:!l.ClIls:r"'l i~1t12AII~ Z~'M, IlCu~5,.'f:'lN~ 11l.~~!UI~ ~ hctlv;iliy di:lomKid, nalkci!S onl)l ~ligh~ly MUL'ulJ;:'r and LJ:tl:lCu:13oJNitl.SAIIIO ijlmlhcled as S.~" S2, an'!:! than Ilm~.of Ill.'; ~1S-Cfi$[s1lmple. liilliggcSl in!!l 1i11~1 SJ.. re:S~"':Cl~\I'e~.Yt (~"'hibil.llargt\r Vtl,luc;';,or' v rc~mh:e 1.0 Ihm or Olht'r 131\i'iGs in ~nl~:$COl1lli only IjJ~lngilNC' slmi:n.hardelli:ng Ot'CilI~. GOIiSIru.· Sll 90 120'1501 eo P'OlliliODD mll~e (rab~e I,. Imowever. litcjlr (tcnsilie,s cnl w~lh prev[)ous. W\Ofk lJ~h?2'J\lh~' dcforl'l'nal.ioll Ir!I!E! SIrn;!!'I (%) (I r the: .BM:Gs; also c;",hibils Slm.in~l.c ooJiCUling arc very slmih~!T (Tob.k: 11)"Figure ~B simo\lI'S thm ~heseHMOs Oln be (:<lstjuw cl.lIkdy eilassy !'Ods amI ~mlltcd Ik!iW bellm/kill's. Un ~ik.c ill crys;tldlinc metals, d~sloc[ltiollwllih d i1l1'l1C'tCi':S Up to .:5 UlIml wilhmn the o~ Il'LCiltal~d de: rOl1!natiofJ is, 11m. ilvlliiilabk: In fi'l'viiGs, scn'mionl or ony O1I;pp.fLre~ncrys.1.'.\1IJiuc ,Bragg. The Imm<'llt:i,on I1ml C'\'d I:l~.ionof s,hear bands, are ~ks" Clear gilIlSS, tn.nl1sc~Lonis IlItd sh:rup Cl)'stll~~hemOlin P,ifOC>C'S:S\."S :<lCiCO~1 thai nl,for' 1,h.erl:JisticI iZiltion C'\"cms <lJ:1i: ~,,\lcd En the DSC~i1lOC'S it)' of B~II(j~, C'Spt::\ci;(!iI)!,m h}\\ICml~el<l,run;;s. O",i:g, I 1 col1,lil1lnill~ dlC gl",,,:,,>' H;)JHl!I'\:' ofillc CTCClOCl!ll:l!Hi~Qpc ,SEM,1 Ol1z;;~'V,~" BMGs. 'fhe tllCJ!l1l;l~ 1~;:r;mnlCI!t.'fSare 01100 Sl!n~~ Sc;tllning dQPl~"on Ihe .spcl;'.imen f>urlln;1; aa:cgh'l."ll in fig. 3, IIl;;llrJ;":~-d in T~b~cL Multiple I;{lmpf~.l5:;:i(m. I~~;;; \'!l·'I.;'I:l:)p'1.TJ.Qm1Il~x! .figll:~'C 3,A slmw:'i I WQ !n~l)tl~ant 1~·<ltWlf\;'l> hhc,~r n bund" O1ItU11!.;" p~Mi(:: slr .. ill Qf :;:!,7'y.... Firl'i~. tl:l.,; .1L!~'!Imnu'DClnp~I1It~II'¢ un COmIJ!l.lIS:i.l.itj,.IfI:O; 10' 'S I
ddQml!1l.io:n behm,\'iol' fill The ,1!,;"'00J1I1i(}m}~ Jclb.m!~:lbillhy is ,;]Scribed ·~o[he Imll1oge~loo~::: and OOnClIln':lll ul!il!cI~ti(IJfl lli~dl.woO!l,utiml of h~gll~Cfl~1ty ~heolr bi.U10t'l Ihmllght~llH Ihr<: ·S<lIT!I~~,whiQh <l!\': (O!~~ IPo!lyd ofh,lr-d .",,~gi(ln.<: ~\I!'WLn~cl~ld:«,rt r~gjol1i'. 'lEI)' -.m~ BMGs W~re rfCP'lh;d b~' ,~~!t m~~!in.!:lmh~ 'PlU1i.!' c]!C~f!e-nl~lI!mfc~' a !llui nloo AI' atmGspllt.':I.'c OlUld iJl sill!! S'i,ICI!,VJl1 aJ!,St"!lg i'l a Cl!~n[}ld.TIl(.; <1I~:rnlrfll1(DWI$n~~u~ of I.hc· m1-ca~1 m.n'o~'$ wa,..; :!IM;:('!~!Iim;d lJy ...·r,IY diJfl1lctioH "XRBl. diffi1:reUlial :;;e~uuDl ing e,lhJrD mL!1.ITy I DSn with a ~I.c:iling r.:LI{~or20Klun iUi, 1I1l1.bsm h,su(m ch:~I.ronl
rool1r~ lii::llll)C;f'3t!J.llfC.
2B} w.i.thcilIt nilC!~lfil1~. ,,;J l;an1.1!ler,~rhe nakes UI;! I is, InmlC' Ihan ;; III In ,md ooutJ be r!;',;~lirLllad~cl 10 ia:rgCl' );i~. rlfif'i~yiuSiliS~1 l1{1w!!h:inW of (iIe BMOi>. MOf'OO\ er, Ill!.! BMGs ceuld be Ix'~n imo dl;,'i'irod :Jmpc)1 tPig. 10 s~mili1r III Ik,xih1c !ncI:.~i;t
'I1mu~."I~ Ihrm~d in di I~i;!ren!i!\:'4;l~QI1S IJ\ en at d mh~i!lilii:!1 ~'"1.i'I.g."~ or U~l(lp~t'i~i~ d~!:OrH'l,: .. ig)ll.lti!i:ii n,.":iu~i:;. fton~ !lIw. illh;;rn~~iml of dw s!I.'1(larb"tL1d:;; lhll~ ;l~kr~urrol,mdl~ngstn.",,:-:; liL~ld~miid ~ril1f'! !II~ i!1Cn'"lS~"d Toc,1I sLrel'it,l, ~V~~ (J3l 1.0 onmpCal:s.[te ~hc sh(l;lr ~n[~dil1dllL;«i !\(l.rku!illg UI) .. :lPn..'n_lllltt~lids wouM 1,;~1ITy 1~liJt]i(.V pl::isuiC' slr.iin. a\ioiclii ng IIj~ OOC~ilf("J1t~a.f c[~ra.'ilIDphiC' liJltC'. SL-COitd. dt~ snU:"ill' balld::;,fliIJplL'f1r wa,,,..y, The jXi'iookny ofth~ wa\'~ h i1bmJlllte 3 I1iln;w~lkh i~ ool1s1~tci1t w.rtl~ the s,~ olflhchrucl remtm1;" in Ille
nc
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tla:;.'iC~. In addiliof], "villi£:likc shear band.'i. were ~bl'lllcd!lICin,g nbc ,'rim1!!1)1 ~~~C<lr ands (b~:tr.;,k. b
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I1ILand @hilstn, conilI! Ilt:s (E, ,til? shl?!t!w m:Cld'uh!l.iS G, 'th~ bllilk rIIodiu'llls K, <! nd v) ~~a~l!Ir~d by afl l!Ittr,asQ~k I'I'Ill!tho,dl(i(,rrlihe Sl.lp~~ lrl~aS'tkBMGs, ~nd a fe;PII'I~'€'(IM'~~ve I3MG Uf'l~!ij;~'~ :ZrC,uNiAIIBMGr) 111 lh'e< ~!I1~tlil IEUM.G-rorming ((Il11lll;loS'itl'Clnr l',arl1geil'ldica'led ill fi'li. lA tln~ soown. 1,h~ range or wIwes ~or v o~ lili,e ZrCd'iA.l BMG~ with poo, Of ~.]I!I1IHedl j)~.alitii~ity hll~he (O;mp®,sii~iO"'1 range
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53, r,~sp«,ti\lielly, (B:) ((lmp.r~SiOO ~L1IIf.es!(l'f S,l" 52; al1dl S,],. () 52 bent ]l1l0 diU,e!'rEnt shapes.. shOM.rii1n9 eil:QepiITIllClil dt11'~o-rm," Dl Ut:y. I~ID) $:2 d,e-o (I(lfmed '~I;! '\I'~r~©Y:s, fll(lm~i~l, :Strt!~fi1$. A bl~r~1 mil,pe Cii'Hl be ~Jl\\ern', wl1k~imp!lies thi'lt hidf:on Ibel~~~nl th~ sarn~le and the p,t~t~nl~ocnlrredl.
1386
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IDIl~)jC<lny ~r"h{lfl1lqgQllO\1'Jlll1!!lru
profllS~ amll;ol~CWlfl1l,':r!I-llhe;If~.lal.ld rrnnm!~O~ 'l~'l}, ~~!lthPl,~~h !hlil p]~~tf.t; ,'l\Inlijl~I" !~<lili~o. Th~ ~~I~ R~:,l'ilnM~ ... ~l~i of ~}'::l[{LL'i, \\ih!,):n;: pliik"liic sic.ti~1 is loealiZl,1cl: ~ll slip ir}' of lh!,; g1,1L'1L<;(:'~ i!h their ~~n.I~!U!t'. TE!I.~~ w iL'i"'¢~lig>il~Oil!i! We're P'(rlOnn{;d io OC~iI] lh~iF mI'ii.c:i['Oj;,[tllletl,lmJ, rc'~l:lmC;:o;... figlij'L:: 01",\. :l\llOW:;. lh~ !g.ng,hl-I'icklilinlige Dr '~je a:;,.-C2il>l S2. The itlo:S! nl.(.'IJj}bllc,ei~l!I_mctC'risik j:. illlHt tit\: BMG ill CtiiW~ p~ed or isolated ~iar.k znnes, rnn!l,i~1~ ffi~(,}I1~2. iD' :$ ~Im ill size. Ih,u, are :>1J1"II:(!!un.dccli ,ool1timmu:l; by brig hi, m~C's. whiCh are abmn O.s 1,0 '~ ~ml iu ~'\,I dLh ~ volume liTIct L\!!11 of dlC' bir.igln ~(lJles·is est ~m!.'t1XldW be -I ~;(," The s~r(u!g 05nlll';~l ind i~OlJlI.:~ lhm Ihebri,gill 1;.01101 are thi nJ1(V lh.~~!~h~ d.ut O!1i(;"i'i..ilfIJ).~yill1~ flhignet ~liJUlinjjllt,~,co~~l[h~ 'lC'lri,ghl ..;om,~ ~mdcr Ihe- ig!tl be,ulli. which OO',nll!:' fr-Ol~l d~\.!: ,d iflc-wJlc!(l ~n m;'li,\l)1-IO-'lllic:km.~$ can-
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~l~af rnu1d&. "'m~ '~!L;'!lpq ~l~~ i;'."~tr;:I~1g d~l;nm. mili.ly.ir5 bl.1~d ~m ~1~ di~,in~li ..'I; :;,tn.i(,'!JlI~ ol'lh~
lo c(H'R"hllc
.illborh bright. and dnrk :;!O:il~'S~ut':huJo rins;;.. oonlTInuing the gla..~"'}'m.l\lrr;;in oo!h zones,
N n.~!O!o~"S of pli1<l1~ !lcpaF.it wh~)c:I~ O\,."l;:U~ did
M!
~l~ur as
:sm"n
u~u~lIy on lh~
Cl'i,.:~~j~tl;ho\t: lilly
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r the OI.;'CtII'fCI'aCC (j rphnsc ~ep<iilibliUI~(3) (ra.~. Sol). The demi 1t.'<I ~.rnClure uf the brilllhl, mH:.I milk :!ioilll l\.."Siolls,~n Il M(l~ and cvolv e into :;i!c<J;, Zf!l1.C$ was slso exml1~I1c,d ~ mc;ms or:~ IRTIS~. bml(ns upon l'Qadi nl:!" COr.IScqllI:!Ht ~)<.. lum.lcro~ s (Fi,g. 48). lIhe lt~l:nilOll' 1I.lrn'£l.i'ke 1~;:U.lerml w ~tilQlLt .ne :JlCi1lr-b'.uld nuclei att! fOmloo cmlcMI'rcl~t.~)1 ill dl!.': regi.ti'Hs. H(~\:veN'l'q:, ~h!.':)1he<lr~b.jJ!!ld l1L'lJllilg..'l~ an,' !CryslaUine In!1g:~ which aJ:\.'! typ:uc.<"ll fcaturt!{1, £1r ~!<l"":.C.till 00 ~~~,clc~y (~hi'I,!'r..~ctLI",,;:;'!!lum.cml.id !io,l1wo!JM 00 im,I~l)d1,'d ti:!e ha~'d ro~on!s'ni:ll b}1 In;Jn{li:u!d~Il:!;I1:itu~ '[01,;'-"1}; nn I~lf; 'I~-!,.;l~l S11,11X~r Ill(!lMi(: in~~wn~"C .d!~nl U1.(:1,11it}!Jo;lgll!.il1g dirot:'uimlNwlcl. BMG:.1 ~hQW ]h.u ,I ]';W,.!;C' nlK:n!<uion or ~oml~!l. :l!.';;sil1~~· ~nlC"'U"'~l~Wlda!ltiph(C'O!!~ L'OI1., n re,-.ll:!it~n~ in Ihe liI."Qt!j!iJ.>d. Wi· !~nl,;'l!rm~ idc>:Jnicod d~pth (50n nm, ~\I.. vel ike O'!I~I'C'~!RUll(:~of ~hC:8h!.':l"!.I"liNand:;; ,md b (iO~1~!ldingl(l an i~~~I~nl:i)Jlio~~ of -4 "'!IH~ ~h~ tb:nrnl(l.tiau of ,,~'ij!~g!lnd:s, f~w irn(lU'JlI..cl~'mll ~i;<:~ U'l."il pffidll{;oo by dl~ ~ml!'ocdbl]~ily or!l;II;llbri~h~ Ix-lwl){lI1r UW S~K<.:JU; lJOJlll(!S m10 Iwro If!;:giQn,~would ;t:Qm,.~. hcut) ~d,C'S iL,~,~" OC~!,I:rw W.ig. 4A). Sjimrilur (t'il;l. SJ ~ whi~h ~mliC'''ll(,\~ !;it~l!gth 'ilri.J1!.io~~ in cmnp\1mmH~ i.h.\: soil'irt"lliflg il1d'lI(:',,-~d b~, their !llkr:osntuCl.u~i; are O!QSl"Wlc'd S I ;Jnd S3 f!"~,. tilt' ~a.~s~ ~'11~ 5Irucl'UI<'I1 slw:liL,.'S Ol)llrl1rnD the rer
(I
~allabk 0" re·mT:,~n.gC:li~~.nl ~W.~i~d ;!O;~~~llIl1' ~i1:t1;,';!oj~ ~~!~\lC.~ Ihe nlic1l:MnL1l1~nl~ [or IItlC' :;;ht~r~E~nd~. TIlt :ibilii.y 3 regi.orn to uml,,;~o ~, ~h~'..>;jrr tm.u~l>~Oml1t~ton. depend.. on d'lt: It'tC.III~l'irlicilt~i'ilClMi'C ~l'(v~The ~ihCi:lr Ira~'l.~jhlml'ltl~im~ Z(i~ occ!1!Ir 111:'Cc~'Cr~li!l~~Yin. [
!ttn.I.clun;;i~ undi;:r
!lUI
or
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ChCI11~Cful .O;HldtjO:lnpos~lif!oi'l3Jlhoi'il0g.Cih,:iIY and sil'l_gl,e gla'>:S~'1.11I1ureC!lr ~K' BNKis.. .1ioW'Cv.£:r. tlilc rre:fcr,e~u:ilr~llhiilllii1g i~dIe br.igh~ zones jru.!uca~.~, dUA t~ll! bright zoacs are !'<oncr t&~;m Ihe Wi:rk. zones or th:l~ Ihere a1)C mueh .nlOrc 1l1C'ml:l~!ly mt,,!~b1lc nlonlic-~~,e open VOllVilIDi cxiSli~lg in [h~ m~~rW!lc:;;, (:4.15), A !'O~m~Jr i'1lru.cwml R:,~· .. uee ~llL";;also been found in Pd-bascd IBMGs (.?6). \iVc sU@g,;."5.~ :lniC'rostl1!..lCl~Irc.co'.nlposcd or hanl a ["C.giOll!S sufltHl!!ldcd b}1 81"111 rcgto,ru;.(wi~hIO!rg.cr V,I]\tI."5 or v], iu jhe ~\I~f pb...;;,ic IlMCJ!ii,. '1111; :;.1 ii.!ilIU i~~C'nt"I~L' nlC ffhu::~D(:m nfttik~ :!ioli l\.';!;lin.lls i:ll wO~lrd ;]U~~lIIDI1EV SIQ;l)~Y "'I~ihGI!lII. Mlb:i:l:tII,mim.l chrmgc.....In ~)¥emll dlL."ri.';;ily l)6), The rchni,,,'Cly
~bI1i11~lto (,'fl. The cOlUi!>'lc!~e,Y1Jc1WILICUi LI'K:\'!9\1~ ui pcrio-a:id Iy of she;,'lIr hands ~n.1P~g,3Amu:1 ~131 of hml! rega'DfL~ in Pig, II 'SuJ'IJOrts the IISSLlilllllti.Oil.
!3ccJlI$c~hc son :region:s c.mnlo:t CJ!lTy ~nore' plJ:t'iI.i;c ~m<lill. IU11hcr pl;t;<::llc d:cforU1;u~ ion h[l."!o 11R1(lccr.! in hmrlrcgim~s, SO·~h"H more ~~I:l!ll'1 is ic-i~y ~iH :!It;!!i!l,,bl~. These ftlc~OIN i~,:!ht~ Ihe- Bi'vIGs ~'VilJjl ~pcr plkl::lilidty. lIRTEJ~: o.h~f."\'3Ijoa~ do
IKlt re\'CllI nmlOCrys!iiJ!.b, veum ]u;:t[vi~y dcliol1l1cc:I c ~b:kl.~" Thi!'i im.lic.rb IT~m d'tC Sl!!pcr p]a~li(~ly is ~ot '~~\.In rmr;ll~hC' ;5:rlfC.~illdl!.Dttd mI!lOCI.}ISl<ll~· li:t-'.'luicm f~1J Ili'l'lllld in mli!l;'r' BMGt; (29): ill~IC1i1cl, il i~ .lil1irirtliiC' r:mp<:Dty- AIU~·.liLli!lgoHllL' BM(~)); an
1ll'tI.l1$iliion
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mined 1:1;0 till ililu'&'i!.)l'ilk me~clOd, ~~.[u1'l~ndicruor of the ~~j£honccntrmiol1l of SLId'! open vo:iltnncs, ~n c the!' ,g;!:iliSSCS.1I:I1e (jIucstiolil 0 g~ves rise ~o Ihis, d ~~~iflcliveSlmCll!I3~ 1ic1:lU!l'C <l clmitlcngi Ilg subis: jcm ~nr~lC' rlillme,
r'''''~lat
mUy dk-:erefL..es dte fllilS~tci~y.llCC3li~C tlrlC fLnr-lcll~· iil£ C1Ltl i'CilmVC' ~hcm1illmy till1st!iblcflloi'llj.c.sc~.Ie !.1f-,..;::n hlrn.es~hm exist :in lite soil iiCg,ion.~. vo docreasing lhe rrnclDon orllle son .rcglOhlS, ;and redm:il~g v l3~~",J I). alii tire o~her han.eL. he inuodll~do:n of~, h(gh fl'OPIll.i:m,jQll of puccxustnlflg
IFi~..t.
shOl!i!i!1gd~rk
or
(~i'1'rinlnl
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BM~:s,VhI!'
me'
Piig_ 3. l~@ SEMi 1111tlg,e'.s;orf ~h :!ililll''f,acl! of 'alle 5~ :~p@[im~fII def~riilll~d'oo~rlQe 'p!astiC:st!1'3 ifll:li -2.70/(1 {AJ' 'iillfld~15,9'It(o(B), U!SP@il:tiv:!!I.y.ih@ snMr b.a nd~, in, (AI app€'~r ~1,!a'!i~.aJi!~ tliie ~~iIIgti1l ~I the \!Iav~ lis umpla r(!bll! to !hat orr Uu~ dark regions, 111 fig. 4, (All arid (l~n. Black ilIrrrnlli!! ~l'Idkal'l1'\lI'ilil[llj~e Sne[lf !b'~_r!(i~, NlI!(/&i, :sh~"'r"!il~od !lIu;c;m. TI1i~in~l ~hQ\lfS wingiikl!!5Jheaw 00 nd5: tI ~on9 1~1! pr,e,maimE! WilVY
,Cllilies; (ilI~m\~).
REPORTS
,~h\Z,;wIbml~S ~,"'n drnmmktdly inerease plil£i.tlChji rnem or ~I])CI" 1)I~tic BI\.IIO", as h V.!!lll"li'x.'r(nmwmce of!3MCis {J2')', These g\,.'%\!lt~ ~l1t1l1n d~ill <! MI1I.IL;tll!!ro! m~.lcri~l" in Q~ her 'li:;nm'Vn Of tI!!!~h~;I'''\ln "tftl:!!;Mill 1~~tll!1';l ~",,~h~ ~~Itl~ Qf llil~ ,~~q]l;)r BMO-I'g<nniI1g {IUny~ ru~do~n an area of[~c~~ pkt.jli~i'h}.~f iiJrl~BMG~ mld !hi; 11Om~~(;nmJ~IS o1'l.md1 fi.Jnlt;i11,~lI,~11ia~ ~ppllk"d il~lPOruulCC_ ,md :JI:lld O~}].lclUID;.;.nrtmlrl~~ii!Onof n~~uh irl.e ~ht:lIr mlfidi... ~hlTOlughou:~ 111:(;'samples is c!'uci,d rar the Rer;erei!lce~ .uull, INotes: i'!l'llim.Ji\, ~1'Il~Il~of i"'1a,~!.~cilY BMGK.. in 11. I, S!:h'iSt!. tL w. J~t.t!UI., ..~f,r,~~ 3fil, 0'll'1' ~:i!OO:3)r. ni~ UflUSI!II,;;L1 pl ... "ULl,;,ilt)i~ml 5~I(;"d~i.ghL o~ tI.O~'IC 2. lit S. ~~rnilr:"H. V. 'Swjg;fn'lMwel1, s, SlJiffilil, Alta t.~am;
li>l'il,it~ M lhe I:kfGllrlllllliOn anW BMCh. stich fiJ;, uh~ d~ nlllt~iel> ~Ia."~icd!!lurnlllalkm (5~ We itiiCfL.~llm;.-d 'llr~e c\'Cil~~ !Km or sh.em·~baltd ~"Paeilfl.~ d winl:i il1.c~asin.:l 19. M. ,f.!. O!1ilr'al<lmbi'*s.. S,M. iGo~. t, ll'II.afl~SDO~, l· G. WiJliiffi~,. 'i\t ~~'11<, I, o!il!lr~":>6.. !l(l, 3:375
Rl)OSl
Inr.~mMl.tif1 :10.. 1911 (M02), 21. V. 1(;, Hvrll<!g~!. l, ]i~, Y. u, !('l. fii!f;fl~:5h, J. Maler, Ref. :I,}, 144112002), ,22. 1i'!I" J" 1!'!'ri!D,1n.. 1!Ii.5il11'i1l.lN, I), tlli~ Mbll'f. rf'Qn~ ~:il. 1M2
fllllclillti,.;:nl111
fi'!I.t'llit\:
or
or
3- Ilc L G~~; Sd'M~ 261', :1'1<1" (U'.'>II ~. It. 'Q'. l'iiimgl Y. u K. T. Ralrrlil!in,. ], U. t C. IfiiJ!OOg~l
~5:a..5143 {200:ll.
~1('IIn'i
mIl
\,>,:idl E via a po\';\~r-I<lw ,r!'.:~11lI.km I d A!1'-~) whoJiC' /,1 i~a rol1~"lnl, and !J - ~.45, \\ h ich is d itfcqx":Ill, Ir'O~l the \"~ll~ m~m~ncd: by d!:;'cleoM~n,g Ih~ ~ml!ll~'ja."fI~'CIr'daii) (hc:igtiujdlii<lmc~l..;'r) 11;&"; ~o nl~l~ 1,0 SlIUib,!l'I I he glil..,;:'H:bmling !lbil i~)'of BM:G~ jblo'mi~~g illloy.~ dlC WpC'l:ll~,ls!ic~~y nhc of BMG~~ .ii<; V('~ ~'n. ..tiHl In 1!11~i, oomp{l~ilion. .i MilloI' devi,~iCi'Jl." in COI'lk'IIl «I1l!!tl-mic %) ea!l! iO'W!botarllnaUy clmll~rc t]ll: plm;;~ieil.}' of Ihl: BMGs,. 111l!' r,r:.:;ulio;indiealcUJml, C"'l'li.'I in [I~ct\~pm'loo
un.
~.m"
BMG "'~'Sl~t." ~'C]rnordinmily plrn51l.C BMGs Ilm~i be ootain~cdI !by d~e:iPlMopri:li.tc cholicc' of 'their
COnlP!G."il:iml wnhlhc' IilSC or lite ft)iiltSl:JJ1"SU;nto ,<;lmK'gy, ThC' f]~'SC'nl ilw'CSIi,g<l1.h:mhas iOcusoo on Zr=hm.,'1.ldBM(t... We c.x~c~ th;;n ~hc !1!lm~y wo~dd provid~ lIs~Hd f,!J.:!i.dC."lill~~S, tbr !he d'cvelop-
B 64.18020'1 ~200U, HI, Diml'du'l:. (. lHoodwlIrci. 1ft lillSar. IW. O. ilI,hk:, Sa'Mff 31l. UB8 ,121006). 6. ]1, ~~ E\!'.{l\\.. l"hft. Iilev. £,m 911, .:ro.~s.o {:.!OO5.}, 1 1. L F.• :1;Mng,. HI. 2ha'i1l!i,. x. F. Pan, j. [)~ ~ Ed;~ A'li~l. ~ .~iil. iI~,~~n {200S~, s. It ~'i. s, ,iije, £. il'.ue-orge. fII'Ir.;. Rcw: tell:. 96, lO:SalO~ £;:1!006'. ~I. J. Sc!~ W,.llOO_~ ~. RftI: ~ 93, ~1l55ct612004~, III IE. ~.~Il~, Mtlil't.;!., 1 f2'OOJ}, 1l1L. Il ]~, L!!Wali'ldooi;i', W. H. wang. :Il L (,ft~rr, Milat ,~_ l~Ii~a:.$. 17 l,HKI51 ::1!2. Q. ~~ng, H. 1M!iI,. E. M1I, 1LXu. ,S{l(p.ra Mctli'l'. ~~~.~41 (2'006). Jl3~ X. ]. 'Guo ,4, G. HiI:IOemC!it., $, J. Foro. G" .i. 'limirle!, ~PJj'f. U!t :e8"lll9tlS, (z,o~l ll-4. \'1. jij. waFil~, I. AppJ.. PrItt ~'}, O'.lJ~6 1(200~)' :liS. !, ~IJ!.G, (:;. !!a~l\aOO~n, W. L ~~I\n~cn Mia Mam. S:l!.
iIIJ_yf, ~, 1[1.
S.
23. ].. £t1U!1'I. tlr' iii, tnr.t.r'ml!hTlI1t:!lli1,8:l',6 ~2!OOIiJ. 24. C. HilgIl~ Iii. MIl~ E, s..nrnidll:i~, F. F~u,pe~ 'IN" UJftri. ~,..., ~e... ,1j: 6<0. 9212 ~l!999)'. .25. O. Silh, It 1:1. [lamikatdl. P. A. KumM. P. A.SU!rn@, !l !'I. ~~~~ 1.· ~.r,e'f; 11'.5;, 19:, Ol'OiH j;2QOO)·. ;16- ll&t1I~~ gt 1#,. Mft Rel'- Wl- 9~,. '14:s.5~n,
(200~).
ROO:U,
27. It So ,!!;r.gDn, Arta ~Ilm.2:7.47 {U791. .213, It F. 1~I!Fnl'lgo, S, (. I~~d~. 11'. A. Kum~r • .It .M. IF~ Il'ItJtlm~llJ{/icI 1l, lOB (~04), 29. M. W.(f1'e!1. ,I!i.. !r!t!u~. W.Z~~~ l Si!!Wl.. i~PIl~" R'i!!!': ~"'. "1l,~4~)0l ~<tOO6Jo. ~O'. L fit Yhiig.; VI. Ii, W~n.y.R, l~ W~ng. AppI. {!1l~•. ~j. 77, 11.14'1 (2:_J', 3L P. IlM,a1~ '1iI" llamamllrl!l', Acr!! l.lal<"t. ss•.U6r
f}OO5)"
B. 0, W\lrIg. Ii. T,!In, V. M, AiM:! Mi!N'N. iS~, 2:911!/I!~20OS}. 3>'1- l1~amri~l, ~~JI!!lii1i ~~fJ\Qm!he i\!!$f ~d Ori~~ ~gr~nlll~ir S&2 ]D1'i:Jil' ,lilld :til!!! ilJhii'ii!':!@ ~d!!i'iI}o' o!. ~~ii!Iri'~.
e
.~¢29 '!200l).
:l!Jl... ¥. t:""\:lmUfil,, ,A, Ihil!l~'; t Ma:!UIII!II'O, An./, 1'iIy;, H'rt. ':l'lL. 7lI'9 (1::9911],- IF_ 5;paepen,. ,1Iw t.iO!~; ~5. 'H17 n.91n 11& f. E1-Agulqr,. l. $. I'!:imlf;, A. H. Slocum" J. K. VQ~lYI, Rt!1(. SCI. Inrill'tlm. 16.. 1D7~lnQ8 GlOOS],
lter~reno~
P'Y 1~5), Eal']~' ~~!.S \,'I'C:I,iC (DIht"ill()d in Ih~ n1alCrl:d:'1),SliC!IlS< or SiISiO~ or G~AslAKiM\.,. 116, il HQiwcv(;r:" 'illerdi:!iOD\,~'iI.y ,,)rlh~; .fl'lI!:~i(lIllill
QI-m: (8" 9), the fI:'lcWI~. Im~ been ~'NIl"n~kdW a
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lic[ld-e'nbcl u·..,.n,~ilOlnr.;;
~I!I:!lijtuler.o~Materials ~a!W~h.Jcho~u U'nliloersity, iendi1i S 9'80'8577, !iI~~. ~P,RlE51()r. ]0Ipi111 ~5(ien~e allHi Vei:M~.ogy Agi!;rnty. Kaw.iI!JIdn13;~2';[!Oll. ~lIpari,J[RA.m Sermri,ooridlK!Dr ~~pMro_mies: ProJect J!lp1!!1 Sl;:ijenoe .aooT~cIli!lilolClg!1f 1!,~~!fKY. ~S:~!'!d"&9M{lOn }'<!Il'!'n. ~~OO~I:w:r'r~ r.!¥!OO~li'Q]1I~$ and '5pil1lJCIiI~3,R5,eardlllll'!ltihilre !ZIf Eliearic.alComnklnXa, tfoo. iGholl:Ul UJill\ie\r:>il)i. Se!'ld!ai 980-i8iS ]~p;v!. ~SCRESl, 'OII!I'!I'I:~i!fIOI! alild T~mnc~ l\genq', £G<iwa;guu:hi332,10012,
)iI~I'I,
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til'(; .. pin[~ .. ialgnl,w'Uh (I rZno, p;IIn:bi.Cl,IIiuriy in '~~'I,US of i!l> novel cNCilclflic pmp;;rla"-;,,. hili :1~dI1.0 ~hc ~... "{x:nl rea:ll,lll1\11l, hOIl'l'O::-;truC~LlmiligJ.lt~l'I'ili.uill~ dio<l!:s tl~ Slttd!es ofd1.e ifllrnllsi,.; POOll~''ill0C:s of ZnO h",w Y~'l:dcd p;;:cil:Kl rOf ~le prq~Mnlml hi&:hM,llmlily t:p:~Iu.}'\:11> ~I1INin!!;j tligll rnobility mId excib.'l1i:c ~mnjllcS{lcllt.:'C·witlil hng]1 Cijlloltli.LlIln d:i· dene)' (3,.01). I('crlaiul aSfK.'CIS of l.wo-dimen.~I:Of:lJIck"~mn gm;; (::mEGj b~h,l',,'ior in s~lniconlclltcWr helcro!<!.fllUCH!fClll. h,we I:wJen ~t!ild~ed by oh\'\.Cll'Vlli£ the (11U.u1tunl I i,;;J.1Icm.. ~QHE) "Cl 1~ ,vu.m~ iz:cd m ...g· I'Ic~Ol!<L~s.port ao;nmp.anicd by Sh!JIbr!il-;('Iv de
vmil:!), of ulh~r l'I'iI!ileri:a.l1 SyM.cI'I'I1',. sudl as HI,1'I~lITidc:;tUn 31lbd!lJ'3phc!lC (J!).. n.c ob1ii~i'V.mklll, of Sd U os.cfilhnio.l'l requi n;:s; cond idem::; suchas ()~l' > ~:md I~ :> /")31; when:: t~ ~s the 'Cyc~ol.ron ~roqLIt:nc)' equal to eBlm· (where I? is the diUlJ.,£C ()nlhc electnm, lJ is 111:.1£11C!1c: ndd ..,ru1.dm* ls Ihc cl~ffill e~rcclivc lnilfiS). '1:' is dll: rnmcr rcJlin.:atocm lim!l:. h is Plrnnc:k's CClnIslmn divided b~' 21O,A:~ is ~~lZmrnm~~ oonst3nt, mid f5 aboo~.!lIItt tf."l.n:pcr::IH.!I1C', A.luhmlgh ;:','C\'crnl C'J)imld,!!J, (lJddci~ClCI"(lSU:II!CI1!.1-es haw sm~s I],ed t~IC'<C wlldU iOI1.~ 1[I'J~, dlC rQlJEh~, nO'~l)CC!~(lh:;,:c<J:vcd: in,!li!o@ m,~acri,d~. 1.1 ~ow(,~\'t.;'~. QI-I E-f~kc~1;;llCW<l!"l .. ~c,,;n in ,I
rill
1;!I~11,IL~i-]1)cry:in<l.~ ,~}rhuUkIlrMoil,OI'"
UJI.11
II] otlr:>llL'dy.
(000 I ).ori,,;ulk,d:.ZnO/Mg)j]j~
or
<11,
or
IK'h.wiiilru.clmc:;; 'm~ grown by It!;s,,;r ll'Iok"C~daL" beam cpi;mNY wii(h lite l~ of a scm&<:ol~iCIlLcto.rI~r ~le.U~rlg S),:>l!Ciffi1(1), 11lc 1'I11~Zf,lI_iO b.~lCr IJi:C1.S ;j]:> ;j] pot-em ial hrm1.Cl' ~or!l1;C :!DEG in the
LUO I~r. {J.;~}, '~Vc UN";;-d Ii tt.'1I1ipr.:ru~lIre gm.dh,:ut tm;tliloJ i.lmt a~Jowoo us.to grrrw the J:i hm: 0\'<:1' ~ wide range OfICIl,lp;:m~lII::e5 on ,"' !>i~~gic :iuhsln1i~e I~J).ULe 1~111ee' smmples (~lmpl~ A. B. :md C~· d~u$scd here WCL'!! o:,r ::l'l!ch ~ligh ,qu[diW Ihal we w~,'C'.!hle tQ lodk at ~~e cfreclls or gmwlh
adj"l<Xlli.
I,I.;nlll~LIIlC !crIl;\" i1.1l
the
I Til' of Ihe 2XlO I<l}'C'I~ aIl~d Mg OlmbOillii"T ]1I)'C'" (l'ahtc I ~~~~ U5~In,
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13813
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REPORTS
of a IHm with eu .\" = OdS t:t.:J.fficr, whereas ~~<lm~~.~~'" (' \\'cr~ selected ~ro~n :B 1)I~~d ,~ow~r"<J.n,g i!~t~'n!Jl~ im1,;-..TlI !{:gio~s, ru,;"'f1~ti\~~.~y. ~!'!1IflOII1lJr fli~n~ ,.v~th ,11:1 ;~- = 0_2 b!1rri~r_ Fmrn .he ~m1i."".~dd 1.~!i~1 ,'m,:'llfIc:icntRu, catlJh:rdt:Ih."iiii(.\';.u = I/(R'1~~) c
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and 1]lL'lb,jii'!ics ~~- R.,.'Pu <Ii't I K were cl.'<I!I· 1;!.11~d~o'b;;.~ (]i,M~x~OI~ 10 ),1 l<i OIl ~m-:I: ,~~l'iJ
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i~ldqK~~Ki,.;!u n.f l,;:-m'illoi)j'~111tl:'1.: below I K. bLdwith increa""il1!:l ~cI1~I'l~raJ~JlIre .•I.~ i1flcre~~ed wh~lc jJI dceU'e1~s,.:d, e\'~11[ll!}ll!)'rcach.
Valli~:;i
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corurol d~~ fi-~~~,LITi(;~I;knlt;,uLy~ t~w;intl!'iu!sil;: dOllqr ~UllCC111irMioJl il~cn:~~~;; w iill illil,'N;m,illg T~(3 J. To ""1 \"~~Hg;.nC' !:.heclcpencleP1cc' on .'1(. it ~:ii, ~lCCCs".",~ry~o {."Cl.!l1!il idL.!f :Sp(lll!'t~I]O("l~~ .n'ld picZ>L'lo ~ IceLric 1P(i.Q;H.riz~~~ cITi:dJ;1 :1 ~on~ ih~ pc~ar iQI! (0(101 ) ofit'O'li3,'luOn or \i!'urtzih: ZltO P 6,17~. Thii.:;S;~po.~~lfjl~L~Oi,b iJidiJl'l~ SiLlt,~8eC chr~~~ li~1 itldii'!i'idLi9~ layers. ~s\.llt~n~ in accumulatlon Of
T"
(fif,;- .~A.). f~'Om highdiifrn~li9Il[U~.aI}'~~~ Wg. S],). th~ ~ij~?lQd~~i.ri,(; ~nb::1'\nil1b;hc'S iMI !;i'll",tn~,in~\d Mg.,lni_Jr0 layers, ~ hereas UP'o',I,l:ro 'pic:r:o... d~~~:ri!C'lJlollll:iz~I.iCln ~hould arise :il'l 2:00 la)l~~rs frrll'll~.cl'i~ill~ :.I[l;IQllV 4"). Thu~" the ~owI rol:ill'iZ<11I.[}:I] ddilt~~ b~' ij~
'il...
~al!Illllc'S,W<l5 identified rub-Clhe O-WX:~ (!). the d~ icm 'n<f ~,p.um"P1Cf>!\lSpog.l!lr~?~~ was tLP~ reci ion
nrd
ru~~QhUl~ml ;torny
aliso lis,liCd in
dCI)leDiOIil of :fiib'C ~lclC'trons, !II. the hc~!!roil'I.~;'I~tces (18). Becalllse the ~row~h dirocliml ofo~r
G = [P5plO]> j ,Pi~.t·l~
;JlP",,(xl
IPii"'~,II,'j
P~(.\·l
1iitlllil l<e1., Growl:!li an d ,eledron1lc: para 1mIl'i.e ~s Qlir :~f10~M9xZIil!lJ..P he~er(l~tru;du res. 'E~edroril demUies we~e Qbt~ined incle"e!'dellt~h(N'!iI the t~~\lIo:fnelld (!(Dd~idell~RHIPIlld the k~i!\I~and Mgh;-,fiekl H~l~ , ~~fi'Pfl:!i 5~,and 5,.,.. Rlspect1lvl1!,ly (5<~ :~ig. ,2C).
IE:t~(mrol'l! iIiIlIiliVI5ity t1.01:2 ~m~O!~ i'llhlbillity I.! {()nn~ ~";1:)
where P~T'~1() i:: !).lm!nlm.n~o!IJ:-O ro:~:lrit:alion In !h~ M~Zll!-.tl h')'~I"; and Pp.!~\")1 is picz(~~ eleetrle lm~iflri:i::alioll ~n Ibeslrnill,CtI ZnO layer on lllus~u~."ill1oo NI;.Z~~I_JrO. NOlc lh:ltro$.i]i\'C'
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Wi.! c!llel.1:1I~~l!d cr ;I5 . .(u IlmCQiU<r!or x ,Fig. .1B). Wh~Ii._'I(·11;; h~g&K~rI~rlnl.. OJ ,PL1'1I1fwltilOil ilidti:<:(1d: [ pOS'i,U;'llC' ch:ll'gcs all!] (bh,l~ shading) are fml'.md: :itlhc' ht!lcroillltcrr.1c~s, iHlcl Iheir tlcm.ity ill", C1'C3!1.CS ;IS x II1C1)Ca:;cs' There is 2,00[1 3\!!i"OOllC Ill, Li)ctW'C'CB C'xll'Crimentiilil II (SO~ id ei rc:lcs) and csl.in~~~oo iDle, Jcs;p.itc~]~c Mnblguhy in P;1,...!(.1{), :lL rnJlJJlc of d1Cnllc()J.'Ctic:a~'\I8h~i!Srel)1'CSClnitiJl! all orlJ~c sllilldoo rogiOllS.,~Uld nile lack (I,f eenslderm it'll] 0.1' chrn~'C cmnpell siniO'JI willI free C~I~~~tons. l.isim,g die atMCi!l'lcnlio!1cci 'pol<lri~toc:m ChilXg.C." O)!Jld b'lInd orlkcl (!9'1, \\11'.;' Qln (:Q~j!I.fljICIIhc p;i? tL;'nll:i (ul d i;)~I:;:um !1(,,',(ur 'tlbl,; hO'L'C'J1[J(i m,-TIi!Cl::.$ tFig.
EF' 614
Ea 70
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!E1
dEc
1C)l UK: SiLlbOOlld ~Il(:r~[cs ~tiJt1;ilocd u~in!!!; <11. me iri.:lal!!l.u~'lr.-pm(;luifil tliPjllroxijm~nU~)Ji am [hUlai me lisl.cdiDn the up'pcr panel. The l~u~cD]Y scp,1nltiluns 1Xtwt.'C~1Ihcc\\:o ~01!'\'C'St subhand r.cvcls f.U-egMttcr
limn
the Fcrmi
CltCfg.~. wli.ichsumJ)~.sIS,
!c~llrcmtutt: mll!!.c or~h~ ,e;o:.perintcm., described below, ~flilI'licr OCCl:lfl~ltiicUl in I.he SC'Cfir.ld Stlh'bitm:l
'~"" "
'!II
I W
e:·
t
,E'c.......
~OO nm
. II
is I~cglligibl!c, We measured the m mgndmmnlspon properlies Wig. 2:, A and :13»lIIsJnSfl sl.m(]"rn lack-ln teochlliqm;o: w idl a.c cJ...dwJ~·on 0 U ~lA, 19 1~1i!:~. ,At IO"'\' field. ,d~I'tlc :'iO!:rnpolr.'l'l e.",hibin~t1 ncg:lIij", C
nlag!l~l(l!'!;;',i~!mwc. ~1;1iK:l'iUfm~hly . because ot\vl".·;Jk!y IQCOlI~i~dC;,JIlTi(;n, A1>o,,'1; -:2 T. d~'...u: p"" (Jolo;ci~l;lI.i{lflJ; ltl,~[ \'\'lI,"PI;:~'IC[irJ~ icill II h' ~LI'lJM;;'I.~);_\J. llU1cl 'Dhcir al1mpl.i III des il1cn;:fL~(;d Will:1 iIIC[(,I!1;illlg ./J. A Ihhollil~h tlre zcro-seaistauec :>!31.>C \\'~!> ~lb::K:'nn bcca LIse Q:t"I~lC' na 1]."': S(;;[l!tcrilig. [lMI::. each ~nir~!ill~IlII'IiI, p~ l;:o.ul1£:i{k~J w~ih ih(." q~u~lIui:lJ;;,\d, of
P.(jI' p~Hicall~' "'qml~lo trl(w)}.\vhcoc v i~ the lmidimj fiUill~ hide;.;;.. Tbcsc 'OOti~rvtli~OIlI> C01'Illirmcd th.;: existence o~'i]~c'Q~ IEin om:'[l.Il1(pI'c:;
-----(m'
.~.'.
1!Ii.""..,"
[~OOI]l]]
•
="
Mg~_;!n, p _
0.1
Fiig. 1. {A) S(iheol'lia!tk
0.2
0.3
I!V
l1elli!lreiStn.!ic;tllir'~grown 0111 :5CA!lMgIO~, :Sl!i~b:ShjUES. Dellelndin!m. ,on!lh~sngln elr ~p~!;!(x)l - IP~~)III, an iUi(llImU~tI~ill>n la~r r,ellr~'Bnt.l!!~ ill)' Ibro~en llnas i'5 Jonrl'l.ecleii~er at th'~~lIlr:~a:Oe! l(luii"l~l'Ie hi!terf .. '(IB)(ak~lalt!·~ (gray al!1d b~!Seshaded rerJ1liOM) Qlnicl ;uc~. U!'I ~a,~!!i r~ ('SIOUdd rcles:) {j as a h.!PlIct~an (If Mg Wifll:Cfll x 'ilil~he b.1! ~nl~F. "h!d~h~i~,oorfg~~flml Pil(x}tell] P ,8il1idl~otid ,orange (Uli'l.l'e [,IM~(dli!~l\li't!'re [ak:llIlatt!1li by 1151'ng tliiiemeti~al va.lues ltsletl lli!. t.a11l~~Sm ,ali1ldl5,2. ~C) Ilflotel'l'l'I'al di,'l!.gra ii!iI iilear t~e 11€'~f1o~l'It'i.l'~faces" (iilktilkiOOd enea-,g'f~ar~m€'~~rs~ue ltS:ledil'l the upper IPafll~l, where .fFisthe lFenfJllf,ene"QY. foalnd E'lare til:efirst ~nd SC((Ind sllIblland cl'aelFgie~ \'I'ith IN~~PIKtto the botto!Jm of~heo iUllilldlul.:tl01Fl hand ,'n tlll'e1: ~~IIlU!]., Ir'i;?lI,pe(~.ijlllel.y. allld ,['!J£c' is the ,cQmjllil~liiQ!lIb@~,c! onset Th 'Q~tl~ Q~ th~' I)otenli~t profile (s:otiC! n!1~ ,!l!!n~~Q~l'iQFI of f" (brQ~>m ~iIiUlS) i~ th~ mi:dd Ie palll!!L cll>rrii!!Sp'Dnd to 'tihO:51! of [he llIPIle!1r p\illne!~ rre,pre'5enrtin9 ~ m.pWIiA, E, iI nd C,
~Irthe
.ZrlOJIMg\o:Zlll-,{fO
and al:I'cnl,lcd flirecl deb:: U1li1iil1i1lkm or dtc 21DEG densi~y {j\j1;O." N{)~enlml i~s,nnr'l~ A.~hc odd s!.1H!s, 5.\lch as v :3 and S.~13d mm:h wider illal:l. plJa!cm.!s amI I,luger <!Implil,~!,-b, of' PM' minima n::lalive~o tllO~C 01' lite even !)t."lt1;.'l\ ~Fig. 2A~ , '1J!1"'~qLIL.!!ll!.ly. ~IH:;;iI,"\<~~~ tt~.lol1U;:h ,IL~\' = 4, w~n.: M ..
11~1n;~)' l11I~~r'i;'ab,lL': C in
r@spectiwly.
REPORTS
wen prc,'),"'C'l'\'e~ elcvmOO~eli1il~rnIUfl:.s {::;Ie'C .rig, .M 21\ O1m! IPp~ Il"n~1of f.~g, 2D'~ ~ :-;"<u~d,~f(1;1I!<dYl"-~' ,,\!(,~~~~~ror.!~1I;'d lJSiifl~r1t:~~ ~i~gml~1~(Fig_lC). wbere ilil~ij~-s, cl.kxtr¢un a in Px.; m1diClii dF"'J/dJJ :Il1'C p,IOlk"t! :1:<: iii. i'UlTu::t:iun 1/8. Taking 1M",.· ;)I~:II:;lhigh~ficM .~Ipjles (51:..mK! Su) •. WI; .il1d~p"'nd.o11Iy "vOll~ui!.lIH,id Un) .. ~e:SL/h <1IJld ('5;&11/11. rT;,~P«'1 ~"rClly, ~~!1d COrl1pfUrnL;i dte~~ vahJ~ \v uuh fI ~Tflb:~c:: I). wlt'i,~IC~, ... )'t.11i g\,,~1cnuny C'o:rrespo.ndl.\ hllhc .:m.EG d~ll~i'Ly CO:l.'ie~bllting ev ~h~ QUE. The t1Ib'l!lii.i~10!i1di"tli \!!IILi'~ \!,,'i,.'i),,; ~ystC<J1t9.li~J.lI)' smallc-r Ihml i,!~ji 'g%lo .S! l%,
I. as C\.~tl]pp<)f(I;ntli}, indi>Cal~ Ilg!ha~ 2[) c91lli ll1:m>Cl11 be" ]'ICIfIClldiCl~ ~<lr!J 'QmJ)Q~CTlI {lJ cos comes w~.;;ik~"wi~h!n~J\Q4iI$in~. lo ~~lVc1'\ligmc deuced 'by IllC ,dm<l at (~- ;~Q"l (llil'c'.f.lc!l)ck's in n caHi~ d i!~li;:!:t~i(l!l<U! w~ t\InhJ[':f sw~!$;I ~Iwfj i~y, 2C), .it- ~ll;;;!~";IL;!}1 ith .. ~lD dw.~~~(~r nhc w o O:ri!Jul~;l,tinll dqll;nd~ll~~ of l:h~ s,J~I(dll'ct"I.llIr~1;iI~~r ';~L;'nrml g1I~. In Ihi~I"I;Elari:!, IH~\\'l;;·vcr.w~~ li.lulf!d, th(,': t'lt:I'Il.figq!l rnlion slJu(!!,,,,n itl Lh~inl$L~t F~g. 2: [l, of ill:l"(J.trI~i:;i~rurc:,~~uht'o·I:Ii::h",c"~11"~::!D c..c'I;lrm:h::d th.m'l S.~n.rp~t'A c:lIh ib!wd w~ak esei I~tioJ1, CW!1 \\'id', ~iln d!<lgrnn).;'1.<'Im! COlmer dimcJlsin'n<'!lil\}' C'~'iI· B p;L",~lIcl o 'h~il~'!;;rt;,"lC~ 1~1l= 9Wl. ,uid 5~lInlil!,lr ~ dl.ln!~llb}' ,~~gll~~lr dl,:'P~nd"'nci~ nr~h~ nsSdH c'iliati:mj~ (~ bdC!~",). l:h;:h11\. ier ~Ii'a~ (liH~liIIK-dfor &."Ifi'illr.; B ~ rl~.S6~. h'i Um:ilig G'..,tzibli:.,.J:i~\d the: plit".tit'iiCc of Slirn I nscOI~lr.;l~l. ~amrlc C• which 'wal> Ccxfh.'ctcd In have eil !liJtkM1.~... re C'X111l;ct(-d the value of m" li.'~'j<I~~ ih~ more lilteOjll~ li,cd Ire..: cml'icfl.!;, she!w~dI"".1Ltl ish· di!pCiIldcr!ec: of i.l1it: Sd! losclll-m ion in~ ·MdU9~iob will~ 0 1lippro~ehrin!! 90~. The t~n~lj)ei'ilhJl~ lact thm the OSCl [fI'~lpcriloo~ clcpc'tu:!ed oalllc !11,~t amplitude ..The ~~ulti~~!l Dintl~ ~)10l.for saa111~Ie (' is sho\\tl ill Ihe If] ~~I.of F i£. 3. We s,l.ilburLd~d
Or
n~-
or
2.51iF'1 --~--~---9'~5
45 ffiK
B
~~~~_.-:....
~
Ihe
,ClJicl in !!ir)'
m<l,~1~.C'IlOIIfI:'$b!.,~lO!·
lrorn the
I<JI'W
1K
Ie;
Sample A
Ip~(i:l') p~OWp,I!,!;IO'1 tH). Thl.~slope of a 1~!l(,;'.!I!' I!'r'lo ~h~~ <Ill !1 - J',9 Tg1i'.'~ m * = 0.32 + '0,03 m{l ~~10l 6"'.'1~~·f(lmo~~.nile ,1:h;~(;'k:c;illJon 11.\i.'I~~). ~illnihtr ~1I~d \/<11ues Wot;!'l)l;,;'(l"'IOlI~lled lor lJF.lng.i.ng frem 2.6 to
§'
.t"
4.5 T. Th~ m'" v~llu~is :>O!b1I\JWh'Ui heavicr limn ~h~ blu~1k ]jlt;!l<hW!] l1la."'" Ol2:~mo ~ ~~li!l1I.,n",d b~
Cy.c'IDtmn f~'::o.'C!J]an{,'cQ' gi\ il1g ri~ to Ih~' p~ (22). sibi I~I)' a rmU5~ tlll:l1!1ll1ocunu:nt \!\f~lh 21Jp.!:lJ :1irnllae
cerrcctien
'2 ~
'1
or
~~~~~~~~~o
-b
mhc SdU osdllmtiio:rls. The S••/Sl, ranes were fo~nd ~obe ~3 lor 'S~mph~s ~ml B. 3.11d~2 lor. A !'<a[n~~iJ,!.': These I'Ii!SLl~~S ,~uggcst Ih::u ml iDl~C'n1..i1:1 C'. ekel ric lid,d 1111ing !min rhe :ls.ynn1l.CU'i( Ir~= s ru,gul<!r well f;Wlh"mn1.~lUy removed !lpill (k;gcn1",['3Cj~,CVell~J1 zcm ' ~lOrn'llhujm!S
11il3gm;:lir:: field III I his case. I1lc nn to or the slopes d iroc~I}I gives Ihc cmncr
in two spj !1~I']lil slubbOi.JDdls,,~,. and
8m
[D
.301
1F1
~~F"""~~~T9IF~~._F"""~9
25, I.
I I I
Sample:
....
,1:1 1.1
1..0i"'"~~~_~
A. ·this 'lI.I1<Jly;",!s ~iH1ol .. <C'.ilTrI(;r d!!'n:;;i~y U!I ,on~ rofmd h"..k\~!fl!.i 1:tI!¥~ Ii); li!,u~ in the othct b:lI1d i'Il. low fudd:l; and ~Oitll carrier dem,.ilY Il:!D = Ii.. + U l IL=6 x W em- (2S. 26). As !Orimmp:~cc.,we OOHii ned dear C\' ldenec of [he 2:[) .nm.IHI:'C, arid. hCllcC' \\'C c~pcc!cd nml 1rmWoOtild be equal IOn. but ln fa{;t \'\IC rOll;lld th"l/~:2.[lwas albo!lu' hllllrof,ll . Ta£,l!'lherw:ilh tilt 31l'1l1'lJJIi. imalel.)' HllC<lr ,'S ropes. tlliis. rirrulingimpllic's dlm IJll~ miiS.'>Cd IP:u ex.lR::iiUB
0.9
V11----i----+--ii--""'I
occurred ill <I s.ystcnmi.ic: ~:·ashiml. In th is, sil:mllion, I he values or r'l~ and ~~IS;(_ are Itollonger
rc·.lliab~c.BecalQSC of ~11C ~~rol!1ger p{l~~riza~tol1J
~h.::ldsin
~<l!npll~,
B and
spJh~irlg\,,'O~dd hoe C'x.pcc:!cd ...~md..r!lY i~ dimCl!~l !O n,.1>{~h"C',.'11 bnd'UII ..~ ..~~;:;I~,,;JI, Rc!~t:~vl;ly 1~}W(;r~~ 'Indi high~r \Y 1[1,~y ~'lrmh\.·T rt,\d.1II~1; [he
llIb.lil)"I,(} .u.i~cLiminm.c t~uz landi.l!LI 1C'\;'i,:k In ~a:C'I, even :;:imcl'I 'wiih v -c I (I tendedto di~;}PPC~LL"
ill
IlL~OVC
'r.
rn~u~r
~<lrg~ 8~~in
0.:2:
sfllmpk: 1\.. which has I:he ~ligh~t ~I.1111 the d lseusslon, VI.''; ignoR:d thcposslb iII~ly I,) r
b<l;.'!C4JLUs(: lhi,!~1: WiIS 110
Y<lllc:y
Piig. 2.• (Ao) I~!ll!'lgitll,.ldin~~. fes~stMty p;;oo Hal[ resnstirvU;')I f'l:")\, a:nd dHrle'renl(ial[ Ha~t r~jjsl~'I,1ity dp~d8 \i@'rs~s S' liti~ai~ur,~dI all 4~SmK far !sample .A. 1~!lit~IJ'~~ t~,~IhiJn2:O!ll!taJlt~di. milir~ ar,~ Ul@ landall! on ~.e\i'el'~;iU1ngfa ctors defined illS v :: hf(P:7"'~}' Ol:} :P~ il mUDp~.y~e;r!>us .8' Ime'a:sIIJre~i!lt ]1 II( for :!Ia'llfllpl.'~ B (to;p} .!:Ifld I( (hoMom). (C) Stanl~a~dI fan d~a,grams e)!:~r"(tt:d fr(Jlm Ithe d~ta sh.(l\mifl (A) ~~jd '(IB~. '~ymb(l'l:s, SL andl 5!!t re$l~r to~hc ~OI~ tl mid :hig n,l1iel.d1slopes, .re:spocli'lle~lJ'· COl(l,FS ar:e (:OIil~i$lel1l .. !lith " th.(l:S~1;I~~dlou P.!jJ!'U:n (A) a IiId ((1)•.'OiP~n :s::iilli! bo,b iI're r'l: e¥iii~l;Ia'tedri!"@m dalt~1Qr sample C iiit2 K a,n d il'l I[]= 30~' S!h~l{;fl lP1 {ID}. N'li!It~ tihi!!l e!ld'l ~rl'reme!ill Ilr l~'ci'~ Qlr edrelflllr:l! ~iS· :set.t·o (I, s. (IP) Pill ~ I!!~ r d,etpend~n!'::e,of liulnnaUjjedi lTrnagnetJJ:r.esi:sttlll'lC!1! measmed lIt 2: K [orr :5alR1ip~E! (~op) ilna sample ( A
In~
evidence :indireci. baad structu 1\1.:.. \:l;/hC'i~H::ror not the spin~pl,il~i~~g. scc.~JdQ ~~likely in our s amp~cs. it iii: an i~~II~.estillg problClln 10 iJc s~lILdicd ill rullJlI'C
0:, M~Zfll
......P
illl;saU
C''>:.lllCri
mcnilS. D~pi~c ti~~~. OJl!Ctlques~io:n ..~h~l'lllSel''''!"lml (If !he v = 1: ~ .. Ic ~5~i:;:!1markoib]e.p~lf! k:ll~Nty when
mNSlilr11!·m.~Hlit.
(cmfigmallfilil.
VoOl
c.m.1:1?~In:;d w it~~~os~tl!I L;~II,ml~ A 1.~G~l!.~ G1aN ... . N ht,~~~ru!\!~rtlclurt."&, It i~ tUinici!JI It In oh1;;ai~ ;JI:1l l,r.afJ "!lIi.ic l(i~...t~tl.hlSiul ml~ CI11!,-l iti IlitridC5 l'Ui\'iiii.~
" MARCf--12007
31 5
SICII~NCI!
·www.sr.ien.-emag.org
REPORTS
IFig•.. . NQ!~li~d oog~ 3
IilltQres.isli\n1~'I!\ml.lli5 1).8 r~Jrled ai~ valiottJ:s Lem.~ peRllUi1!'~ ~or ~mp!!! C
1;0'. M. A. K!l.m.! .. H. Kumiil. ]. M. ·\t0ll HDIf(. N, P'I!l. ~~C~IiI~r. Jlj:lpt ~,,~ ~""J,r_ .MD, ~~1 f!.~~n u. K. ~ It.101.C!er!l'l~·~~, SriMriO' ~;~i!l,666 '(ioo~,. 12. Ii.. 'OIilIOlTll't. ii, 't'. lilwalng. ~tlJ~<I,2:7. II!H i(2'Ot)l), U. M. ~<lk~ et ~i.,5tJIifJ Sttl~ "grmflU!l!. ::!.!!I~'. ~51
rl~pen~MC!e o~ Ih:e· I.o!:i:ai" rilhlilil~<: ilmptihlde Oif j},.;., peak at 3,86 11(Iiunlii\raled b<y !'!~,;)'.
0 .005
1
~1. J!!IjJ1. ~flt 72. 2466 (U900. ~!~~I ~X~~ ()rI1i[1l~111i. I.. Ii!!. IKdbil!ka~,. <jdid ~r,~ (Mm~rl.,il:~, l[)'~ ~1~'800, 17. ~,,~erpf1agnm. , 'IN, ~~'M:r.lik. A!P.P1, FNJ'li, £tott; 1a. 2:45, H
u- ~~ ~1J~rl;ing
fl97n
ff aL.
18. 0" ArIl:bad:l~r NO',.}. ~. PJJj8. 8:1. 334 ~1!_)\ l~. 4. iQh'~Q f,t qJ, .$pl. PfJy>..'f'l'~7S, $1;&1) ~19\l9)" i!()l F. Sl!e11t1, Ph)1_ Kl1!Ii a :"I. 4.8511 ~:lc91i!~" .21. It.), 5I'.1d~FJ:,f'tjys. .~~ :n.o,iH7 {1'9Sro, 22. K.:I. IBII1:IQIII it a.:. PIIff. Rfff. '~ll 28. 1,6117 U~? 2.1. .2~. 11. AD!lD, .Pi_ S, fiIWl~r. ~. 511~,rn, .MDi!!, Phys.. .li~~:H' ~'" b9.62~, ~4. S, I)'. 5:iim!.a,. ~ ~. i!\! 2:]', 259·0 '(1l9831. ~~_ H_ t, S,onner ~[ ~L l'hys_ :«~. t~t.~\l. ~:!,6 ,],Si3I, .26, ~. 1". ~is@'Mlt~!, Iii. t, Si,(jntl@ll;\I.N!Ii'~~~1iulfl:~, (!" i(, G!lIlI~i!ld,llit W.iegm·aM, n.~ Rf'r. tflt. 53:. 257'9 (l9t~~ . .2'. W~ iIlia nk !j, f. Cl'iKhilim. tK. Ue~D,., anCi '[ Ful:unura
d~o!i1~.~'!J!I'IIiKI"~el!l !lit .iII Japi!fI SOdely for !hI! Pmjj)a~iJii ~~ S:cieiflce f~~l\i:lh 1~lli)'Wlhl~, ~iUJ; wtlli!!lle
ror
All,/.iIlI-.J'>.I
bmke
U 8). A p~ di[:CCliDn
.rei\ilL~ts if! pilinlg tip of dmr.g!O!s .irtduoo:l by both po~.miz.'li~ ThlJS. (lur reslhll.~ hUp'~}'I~IC1:1(Iollis. dl~18, poss.lbili~:1 orrca~izing dre rmcli!0J1.d. QUE i'n~]~c:,~rescnl PQ1~1]" hctcro:,!]1!,!Cl~I['C ir the' carrier ~nQh~lil)' elm lhl.~~rnpfiiove~l]111 ;'Iddi'ti.on. gj \ll;'n~ chemical ,C'.ompal ibuHly Wi~lOO11<1illether classes of o.N ldes, the: q 1LII~IJ"Jzed I ~<lnstare ~n;.Iybe combinoo \Yllh .. brood "mge 01" phy.:iiic<ll,nol'lC'flk~ i!l! OJ" eomplex ox idC'~C'lC1'Q!'; 1!'I.IC'Hm::.
:l!. lIt N:omUl~.1N at, Sdt!ij~ 1I0tl, i2M ~(lI'Il), Z. A.. 1c!iiJ'koY.!iki~I' t!Il., .~m"M'~w, A, ~.~ 1&2'00s;~• ~_ !!i" T~ul:Mat~ fl.. IO!i1tI!lIlllO, 1;'1" ~iW~~;IH. App'_ ~ ,~~ 00. [~21A;16 '(i!~6~_ 4. S. r, OlDlfliliu tlr!i/,. ,r.. AfJP. ,'PJj~ 99, 0931)00 ROO6)'. 5. K "'~ t1tiwn!ll, IG, ~~tI, ,f!\. I~!l~~; ~ .JIiI~I1.Lm .,,51 4~~ (19&0, 6. ]lW~~'5.. ~~ $~ :S~"JW_.~A, '~3i~U,9l'8l 7. 10. C~T~~ A. C. '~d! A:PPf, ~. t:e'lt n. ~~s,e (USl}' 8. 111(, 'fl!lIi\ It 1L>'LCiimr, Pi. C. !O!:!1'!;ml 1nJys. ~" .l(![t. 48, 15~S19(ll'9811 .. ~'. I!I. :vi'illet~ fl ,~1•• Ptl,ys. ~ ~fl[; 5,11. 1776n:![lm.
lIeren~~u:e~O!nliliN@l@~
CIt~iJv.'!' :Sl:,II!,mHIic1Iie!ll!I1Ircihgrant 1'!G50Ilfl!'! 'rromllil@ Mifll~u)' of ;,dIKalill!P., 'Cl;!lt"'r.~. :Sp:im;. Si:ieJ1(e, .~m:l 1I'~nmo!I!!j'i' oll~iiia~;:li~d:h~Fn~~fijffil;\t!l~ily I (.a~~rMIw. PI&!II<lIfI GI:t~~ 1~s;tilli!le Jor Jilaleriiiii:!;, iRt,!t,arch, l:J!~an, :SIIl!P'POITt"!rlgl Cn'Utille Mat~i!lO!I.
~.iI:~~!'!I;!g,argr~!lLrl:gn.e!1~rU!1l111:~.743Ml(1 "'l:iI!ri~ls.aDiI MI!'Ililod~
'Fig!;. S1. t>m s.6 l;!bl!;:~ 5>1.and 52 .Rt!rerM~~. 11'u~IJ~li1ed ~ 25 ]ll rn~iI!'~ 200:1'; oollil'l 10,U261$[i~~~~.H;HJ1!~O Im~tJldl:lni5 fnfQ[m:J~~OO '1'f!1I!:r] "fling U'!i~ ~per.
:w r4"",gmD~
lJ]~
~.~~Ied
K. !.. W(!l'!if.g~~' P~Wiq'i;~ K-Y. KWI'JI!l.l. X., Un,l ",i, Ji.;lJ~.l U. SI'JILa!'iki,~' Fl, !t J" tr~w~nt,l G. L. !:'I~n.eb~ iT ~, S:'tQlbQ\I/' 'T,~. R:~h~l1'l1i!IZ
'We 'rm,md ·[h.at !I IlUuaquil1Jame' diHl!I:'!:l1!$ .a11nrr.g.a ">traligM ~.iml,e~OfIOs."i[l [,lali, hi91Il1l!~symrnetric (IlI(UJ) :5urfa(e .. It ciln also IN~VIHSiM.y .~Uil(h ene Q,r t~\!O (02; motemLe'!> <:IS ",(argo'" OIlIldada~a ~'tmole(llIle ea erier," ther.eby hi! "s~ornninJgl the diUm;i'l!\!il beh a vloJ oUhe {:O~IIIl otJE!l!:ulesholll!lioobiIJpiic to lir~,e.u. Oi1IH.i~:y llUlmdionall thoory (alll:ll.l~i'lt:iemj imiiclIb!d ']!iubs~rate~edillll;ed ;a~trril(n!lnJef -0.'12:
'i!'ledrolfl \taU ~~V). Sil:tlinniill9 hll1ni~l~ rnJi:~ros:(c·~y re'1fW~!!dlil1d~lIielLiill. steps ,of till!! mole!CI.il.u IiIQ1 lcomp[ex. 0111its: difim:iofl ~altrnway,. ~'I!i~hinJueaSie-!i oif ~O.03: al'ldi -{t02 eVilll tne d~~hn,io:n barrri~r !lIpon aU[lcllll'ffUant or fn~lkst. iimcil£~mnd (0<1 mol~~~u'le., res:p~di\ll!ly .. and ITIitc'lrlOlS (IU. In. 17:01 any renli:alliol1 0:[ hl,;.'. d. C.Vcl. OP'1.1.1.Cnl 0. f ~~lQ:leCUIII'c.:.: ~.13.t,. W.l1~n.~ acloo.ihcd unto a sU!r~a(;c. ·eJ\,I!n:II:nt dyrmrmc W1m,k,-{ulilf-'SCa I~c'f1iIac.hhn.~I)'. control led tr.lf:lSDXl'rl 01: d~rronlo{;' plUpo.;rtM.,':>n·:;t,'l~Ming 1l1O.l>J.J r"rn~6" rnolL'ClIl~ i~ k~J .~inl'lcliorl:JI.li[J". II, IlilU~h uflmLc«~~opiC' o~jccb; or mflchlll£S.lla'O anracted as Irili1:ipnrlmiQll orll;meru~oric o~jl~b>ici iJilc collo;id'crabk: iUi~tij:Gn {1 .;}. Ell:.3f11pk-sill'C:[:[ldlc j,;~y j;JliltJiW•se ofbmU:lY nnachincs. (e.g,. Cflnl'l'cyor mtoU)i. t ,,). 111InOC313 allclnmm:Yw~l!.:ci:S (5 71, yar~ b.;;::iI.,>.nf1tKks}. {'.cmbml of :>urJ"acclmj .... on by p iOllis. kiu~ds, f ·ch:'(;triic.u:1 !>!.:vitdc and !cads (~, 9}. m~ans or adsol'l'!lt;."\1 m,o;l:eclIlc.o; 1tlJ}' ;ldditk!lla~:ly o G ITcrum'l'!!il O1\!Ct1ll1lC'S ror ~le 'OlJ!thlli:l3li:on or MII'Ib.,~ prnre~lliCS ~.ch .'!J1; growll';Cllila~"}I .. rem;li~:,i.ly' lPi~(E! H~U, UIIN~I!ilfmr (~mOOlia." Rw~r~d~. (1\ 9:2'521, ';!Ialrsi!':,. ,md IUic~ jnnltuoriC<llkm. U:SA. ~:DE.'~rlrm.I!IIl,t of IP!rrsi13, UIIL'I'~r:siI!l' '01 CelllJ'al Ftoridil, )\~~1~L;;i1 ~l:fuCL;~. ·dirr~~i{mi:;: ~~lC'!:a~I}' the N OJl~p'ldol H 3:t'iSi.6. wsA. 8U·tl1t~ p:rmJ~~WiEh Ih(:~OWl"Sl mL;~~ric bntri'l;iI'~ ·Vo \~hom oo~~e~p(!lfnl1fen~e illlIOuI_dI e addlre~. b IE',rnait
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niil5iionl is, random illl'lmWli'C om! i:rolrop1c surlrtocs unlessthe surtace hself l'i 31'1'ismrOj)ic U 2. !J I~ step edges H -n. an i'Kllffll)ic :SUI'IR<:l!S (lJ). and surface lCmpla.res /(i 17j em :r:e;Ulcl d ii'rt:lS,Lr.1l 10 specific ImtlnVllys or ru"Cm.... The use sueh surtacc I(;m.u~ is, hcwo'ever. a vei)' ill He.... e al'!Jnoa.ch for glJl i.cllnng ib\1 :lur.f~c~rnllspor~. l~rc'Vlm~:i\siltdics have SIlO'WIlI 111:11.he iSlJ~110j}ic dil1~lsion of ml$oul:l<ltes on t S\lrlaces call. be turned h~1fI Ih~ca_a: g~lidJcd and nmJ:k~rl by~m~a!~"f hlro!JlOl:'I~im or ci!her ~wn o :'1i,'lt]uq,~miall~' ull(wh.m !hi'ol S~Ib.:'i!'~~~~li:(.'lk<;n-; 1(,6) O:~~h'],~L't)ll~ '\11'11",,:;1:,'" l5. H} ~Itl'nIDle ';:i.d!'i~lr~lll.'!. 0111 ~fLI
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;n;~.Hz'b"tl, 21.~d~mi.gh CMccrlctl mot~on ofri.dsorooJte:: ha'> been observed ill. unrelated ~Y"'1cnls tNt HI). ~:ere l;.\'C' :i11(DW l til al an~.In·aq!JllO'llC (t\Q}~lOlcctq les (Fig. I A) i can lr<lI'L~I'Kl'1Iiom:: or two CO2 mG~eculesin a hear ,lItel ~l! idcclr<lSih ion iJcro~~iJin i~ll'OJ~ ie 'ul i :~ $urlh~. I) Thi~ !iiwdy u~{jd 'I;!J~~or!l-buihV' ..ri..bk'"
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ool! Ies;[PI~;Ir .<!is "[iq1l'1:,'l;<;ioJlS til Sl'M im<lg~:. !he}, <1rCml)bil~ in the ~111..c l~mllCl'atli!w !1'm~c (]intlL~ i~l 111] i~o~!W~'}k I~...;i~~~trig. ~). IlOS!!~ 004 onto d~~ s~:mp~~ !~r~ltlgfu! ~ :~~"4,..>d!i; :;;m~ ~h,,:;y '!t:Jl:~(l., M(l~ aha,!!! ! 0 :;.:]mpa(;;;:WI..;'11;: ·pIC·palJl"4;! 10 OUI su!"fit~ \I!I'ilhtilhtakl~Y~:m'lJi,ugc;_" AQ ~cl. of (."O.h w(; lilld lii.i:a'l;hl!IiC'lll of 0(',0;1 ~llI]o1e~ilIl,~ to oOl]f~nlll tb~ ,~n~~!:s :rcP9rrJ.,.'lBI~t:rc. each side fir f\.Q by I<U"!oom dim.lsio!l or ei~h~r I\Q m"! C'lll i ] ! I is~~nill~~'"d by SlM .1iS an ~~inng,~~(;d Ilmt!1.l!S.inn !hOll i~ ~.Iig!lcd ",.~i!fh (he st"lil;'"C;~"S o.r·~1I.e'r~~ .!'ll'IIl.i("lldo.1lin.~~.I..~y!.ll~ STM ~ir .. ~r'I U'k~ ~..\illll!U'~L ~L1IIlipkK. AQ J~'iai~"1Li~i~ itl; ~i~~~r~ui1l8r~t~~~ higHi.• \,)'ll~I1I~l!) diilt.."\CtilJln"Til"l'k~-I.. p:;.c I dHlll,:-.iob1 and i~;;;doljo!¥1ll'tl 5h~IPC~ illiagi~g re.\ ~";;16rnp.id fll:lJ ~rietljl liuaC.t'I:i.dli 11m,· ~rcti~B:i.c li.i1C'lIl1' I'!ow'" ....er; ils ec.nDer l'iil:!;lb:, ~itl'l!:i.goo as lil>lS Ilrusim~ 9.l tl.11~prrnIUDL.'" I(~wIt" 20 IK.. COl ,Iutll· as, Il'1iidk~. A. S!."liLje~~re demdun~m lind 1\.... of ~i'tFL.cI~menl. one of two ,rn~ of molccu!es rro<m AQ i.~~hQwli1 in fil::. 2..A to (' ~lo~. U!lloacll~~~
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Fiig. I.. tA, i!md I) The dirimlion or :A;Q (llw~u!j;iOO'!i,w~il'ei) and GO~ <~ru::lenti1tiom;" faU mO~&lII'l>E!1iIffil'lWE!l1 two STMima:ges i(rIJoUage' III == -1.5 V, mnrent. J == 11m pA, '~BmJ!!@1ra!ure T == ,26 II!:"alr~1 A == 95 :0; BO AJ taken ,2; l1~tI rs I'Ipi!rLilihe difh.!riion palliw<lfS ('!Wlte li.n:es) were obtained h'om ~ll!tl?1iifIl8dia1)1' l'ml'lges:..
ultc AQ-CIO;l complex ~tcr:" across lilt ,'lll'ril1;"(l:: F~illltcmiyllre !\Q mOl!;!!}, j,$ l:CXnnd to be ;jla~~d l~y ...lW wil~~ r~'p!.. ~o~he ~ul1slr.'lXC l1iglli".(:t SY.lmillcH:y di't'OC!i.OIl" l\ ~lllellcc of tln-ec :SlC-p.S
of<iLll AQ CruryUl2 I\IJ()(,O~I~lOlc:cuk'i! is SilOWIt in Fig.. 2. C 1.0 FThe c~,h:;ul'lt K.1Ill'i' i~~dicr~ti: IlmriWJ1!.d trd Ii»' 0lI 8i,lliml I:x:nh lbl'iMl~m:l\Ct1 CO2 11~~.~ctlllk~ ,nl'd 0,11,,'\1 <1IJt.. C"d co 1'\:0 ~ Fig ..3 ~ \vhich i:;;ul] agrc('Ii~C!ll "LCIl.' wit1i~ spt."'ClIt}$OU1:lic fi,·h.ll:1:>urcn~.cnl" on is.olm.ad
Fiig. 2. (Ao to 'CJ' 11i)@!adllil1:entand rll'aUa;{hrmmt {II a (OO! I'lmlecllIle(depre.Miolm, roo) on tim!! le'fi ~ilfEl Oi[ an AQ (pmM'u:Siolil, blue-\\lhffili1!')' (C tD' In DiUusl::m ofAQ \vijtll 'l\'I,I'(I' (0:2. mlo'[@cJlles, bV m,-e'aI'!S ariildijvi::l!.iI,aL S~~~ ,(!fits ~~~i1 gt~!iliiS(~ Flig. :3). ln a~limaglB, v= -35 iiliiV"I '" &0'pA,r =50 K" (and A = 20 x 40 it Note :th~ ed'ge Of:.ill ~l!llst.etrof AO l1lIol.ecu~s 'in the' botwm Il>efi,5l'I'Villi gl :n a marker, and Ihe Cu.{U ..:U s'ida~estartE: OOCi&i,tJiOOIS,
spcci.cs PI), A.l11c9ku~mliQlt." used (he \lASP rode (11) w.idl the ~~i1Iernl~~ed"£rrJdi(;i1~ fL"PpJ'ox~ im1lllioll (13) rOO'" dlt' cl'!:.Clml1ge.-oolTC~3Iiofi funcliollil~ find tbc' Illrurne-\\llVC ILo;;ctldop!~jeutim.1me~h[!(l ,t~4) \""It~lu~u:a~.tl rseml'opolcntiah, (25),. Our calcu:IiMim:ns, used .'1 iSli,pcoccll ·001"5, x 6 SUb5ir.lltC rooms" and l'ICSL1hso'blai~lcd Willl tWO and t~In;:C !iuhslI'm~e layer:;:;; \~I:e slmlilar. 11]cmlr.l.Ol..irw Interac1ioll between CO:;! and AQ W"'t,vJ wrn.s ,c-allC1.I!h.llcC:l oon1p.;)IJ~ng Ihe 1*1 encrgii.'S nt'ooppcr by
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F~!J. 3.lMorle~ of Hue diHwlQIl 'D'f an ItO' Mu\ccule wi!Jll OFle C02 atta:dud~ resu:llS ror rem C!nildiMO C02 !iflQt~liItes a ~ vrtry sJmilalr. NJ dir~li$es· aklli!g]t~e S'::l'b~!;aile lhigliii·~ymmi€tlY dir'edi~1'I l¥ ~e~ulialll.Y IiIilQvl!ll!l Iilqlth~F qn~ «;1>' liJ 'or~h~ (II~~r ~IO' Olr ~~ Q,~ien !lilc!I»Ir@w ~!1~rs.b~), Sl"I.lp;s, la @~dl ~!;I ar~ im/ersion'SYlllilm~tJ;k an~ ollitlm Otl:rnrirn direct sliIuesMoo solihi1ll1lilie mole(jul.l!1ipp~af"s to ril~p IHllw~ ,clol1lfi!!lmaHcllii:'ii ,anuli 3 (f~£I. 2). 2:
fr'ig!. ~,. Anlileml~m plo!. ,cd [the- diffl.l!llol'l of AQ I'I'II)lewh~$ ~\iilI!h~f'(I, M~, ail~~w~' O~ iIn(ll€'(l,jl~rtS ( aJt:lil.cn,oo. The r~'ttaJnt d:tftmiolFl barri~~,an~ indtCQI!ed.The ~r !la'rs;af~ ~fl'laU.er tha:iil I.he s:ym.cw~s.,~rll1il tl1~s~$itll~t ~nr,qFof N(ih Q~ th~ Ib~nrien; i£ b!.'&o~\I' meV. TiU! p:ref.adiIlf5 !1re LO\'i U;!~lz 10
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[M[u 1[t:ii'f:UIDI'C:tiil'Dln,B EII,coded I Partleles for IHiglh ..,Thr'o'Ug!hp,U'1 [8ieme [IIec ul e An a [Ilysiii:s
IiUglh·t~ rIQl!Ighrpl,lt :SCri'!rl:i'itnl,"i 'f('ilf ~ue'tk a !'I a tY5:i~, C(l!mbj[i'I~t(!lria I chem'i~tj[y" a md dbillk.al di,a9i'1~$'tij!l:!!: be.ne'fit5 homr [mlllt:i~te..'df'lg, wh'kh .all~ws. 'rOl~ jj~ ~~mLllt.ai"leli!JI:!:aS'~ayof .~Mra[l an~iililytes but n 11'jt!~(e$~tatf!S, Iii, E!1"![cod]ng sciile,mero'r a mrl!ilec:u lair iide:ntm~atnon. CI!I:rrl!!l'1It a pprna[che~r.,u m uLtil1tex.ed <llnallysi:s im'Olve (omplll(at~~1 or ,explmsivt' pmlLes!>t's;flll:r !llli1mdlift9. ~limdk)f1al:izirl{l~. ,or dilC!~ding ,aJdiw stlbslrat,es (p,arlides ,(lrmrr.,u::es) tll'ld often yi,~ld a 'r.II~ry tlmit:erm m.llll'lberr of a,flalyl~',l;pl!!dric: ,~o!les~ We preS:1!1I11 [l'lu!thod based! 011'1 (olmtilill.!ulLJs·low ~it.ho!llwi3Jp:hy'th!1l'~CIl<liTiIb3ne,s: i! pllrt'id~SjliI'Ithe~is .aln[dll'rundilil(l .[I[f1[rrii PMD@'if1l(Q,rpor.[l,t:irl!ir"l irl~~ .[I[ sin'gle pmlLe~!> ttl !!leilil,!!'rat~ ml!l't~ifl!l'n{tkma[ IP.anide.~ Dell rii 1'11£1 Ilv.eor <I milllion[ lnriq!Hl rn!l,l's:. By [usi mlgi !I'lIe1h parlide1i, we ,d'~miOl'I'fitra[te a FIiIIJItt.ipiex.ed, ~;il1,gleflru:(Hes,"eDil'GI!det:ecliom Vir DNA ,o~iglomer~ \¥ith encodie<d jlillr[iide' ~iibli'iH~e!'i that '~8JfI be scarmed r.ll"idty ilfl 11 HOI'J4i"uoughi mkro~llllidk {hiJ[l1rneL fllr~he'rmOl·'~., \'Ie dlelfl[o:rJst~1Im '~"i'ill high 'spedHd11l' the ~m@' I'JIIlILtipl.el«!iliI dei@'oCtiolfl llIiSiDl'~ ~lI'Idi!,!'idllla!t m'UllHlpl'obe l!lart~de!i.
:;;JppliQIIJ1ro!l'" p:"(jtl!lri!1,1il ul~m."i11i,gh-d:''',IlSil}' :mJ'il~ys.is. ~Il 'OO:Jll:prU'lOOCl!l!, ~1~I1lS~'o!I~ ,111'1);'_" bene~! (mm
soIUlionkin~~:i(;i;,L;<l'<t;' '
Im!l bybJueh ,!!CltOlyp.iulg
~UI1'k1'
or .,:v.._..y modilIi\clniIHl,
Illsoo 3my:>hovc
piJanml' marnys. rely Mric:tly on po.-;ilum~;~1 cncodin~. ~I);<,;t}c!lrs~!] ,Uf'dYl' hblN~' used II gre;,ll~~lImbt;;'r of cn~ocxlln!ll :;d~~'i!t1(~. Ihm (,<m be clOl....~vnoo <IS "ll!lX''''' Irom.ClJlc UI m. grnpilk .•11 U 1 Nn. e:lec~rol1iic I j' 7 In fl1' rlhy~ic-a~(1~~. J J), SiJlJti,>,C,trmm'uric: .,l'lrcooing Lli;.'l'I.~ "'f"'1,\cir.c w,wo(,... 1:~11~llL;,;; (If l.i,li;ill or~,ldia1i(m lintl\'~~nIl nUO~lphrml,_'lS 7)\ ctUllH!{Jpht.'!I:'i.,ii\(NhIfJh{I!1.on1:i(,':~lnH;;tun_'!; {4 H',. and Rflli~,uU.I<lg,:o; (Ie. U)j ~l .idcrlli(v 11
~1}cci,\."'S..r-~~J;(m;'~OCI!ll)C-";IlCooctlunicrobi::Jd .. [,4'-7, C1.l1i1 ~
he. a~nh)' to q;~~~ljrY.. nn.lI.II'il}~.· 1'~~cill::1.. c)'tok ~nC'll.or ]l!.i"I!rlc add ~qllc~~C\:'S :In l'lali:a~~cl ~)sii!lg <l :;;ingl,c sarnrlc <lHows m;:~ s'c-archers and c'~i~lk;i<llJS to ,)blil~'Il 'liliSh"(ki~si.ty
~llieiPdflUl!I1~nt 0'1 CIilelPical. Eno,ml':r!leElrlng, 1'lIa~-s;amU~!l'tI5. ~rtI· sm~~e of 'il'e~hll!oI.og~, (a!l1Mdge. MA 0,139'. [UiSA. '!B~cMEMS llescllJ~(~ Q!flI!.I!t JllId (ir!rut(!I' 101' EfIIgrnl!)~ri[1lIIj in Ml!dKilil~. Mi!i~I<I:d;II ..I!)!1!UJi, G~nEf;3I. liOSiPi~.. l il:lild filalll<lJ-d ~dk!!1 :k~t BO$~, MA Oil:U.4, uSA, "'1'0 \~hol'll oo~~e~p(!lfnl1fen~e iIIl1OuI_dI 'be addlre5.'Sie(iJ.
IPoo:flI!@l1lIil:J~d'LJ
IE',rnait
mr'lidly rlr:oo~cdl by Ili:-iJl1<gOJ'lvcnlioflfl:1 c cyl.Omc!i"1' lor on U ~r"oplj,c arrn.y.; (l4, I. inltml~,tion '\' it~ mina~m31 &.'\8il)' Lim.c,. S1l.l11J11e mrIk.ifl@. I!~cm a poPl~lar r:llarl~)ml rar l'IT!.I~,I~I)lbivl,')l~llnc. mN oCO&I. Sl!Ich nmllipk'.'\Cd rul;1l~y~i~i:i! ing. I :~o"'",c\!cr.~hcn;'arc' ~\'(rnl di~fL-d'!'<ln~<lg'C::> 1" 0 uSJngmtihiple I1mll'C:lICl;nl sigvmb >lS me;('"Ils 01" ;]CCQIli1\p.aI'lICQ l'l}' ~vcml ch,,~'lcngc:;;,ill!:;;~~!dingIIlWlmrooding. illduditlg, I il lhelimi,tccl barcode!': Iccul!ar ellcoding m'ld dte 1'tC'C(1 10 rel.:''Lirl assay scu'k.~i Invi!y, ~?-"'Cifidty,.,mCln!f\l!I!dllcjlbil~ly 'with Ihe m:hic\,.,lble (1},p'icaU.);, -11(11) because o.f spcctrJ'I LISe: of oOOl'II~;l~ mi.... u!re:3. TllCIlC are two bJ100i:! ovcrllillp, I(l,i~ lhe lack of p(lNbil~ly 1:'01' olill'k}' c]m,~'So.~·lccl:l4n{)logk~lI~d Ibr mJJI~ifiJ~c"'~1g: P~'!" !low cytomclcJcs. (ii~)added COS I widl c<lch Uuore:>.ClI! nl L;:~!cin~.r ~nd d~~¢C!Of Illced~>d. ,I'~d n.1t,u!rdY5 a .1) ;l!ld sU~prc!!!si{l1i1(I~alilick.~lilfol;"'d ~ (iv ~ ~~L;~I i..1.~.irl~k."fr(;n;m;;c of e.~~oora_g nuni "ITII¥-~ (.;I .1!).bonh elf ~Ii'~!ich~iav~"lljJ~ic.~!iolln;:s(':t.mt;~ \'!I' i[h"Ii~~~)'t~-dCHXI.im] Ouu~\_';s,oC!~t~" z\,~"'Cifi.~ )JI.d\!!Ll1I~;I~b';S. Pl.l·.IIW:iim'~jll\ :.ilL;;1l ~lh'lDNA Fat 'I]rh::"t: rc!~",olilii,. ~,illiglc.11 umi\I,:l'1cot.."i'iCC: lli.cll:i!}d~ 31]cl ~LciUfi i'iiricroalTa:,Yll. :.I£C b~";ld ~ui't1.'tI!(;'r
now
REPORTS
illO\~'d ~h~IKe bQt~ i'1f~l rnci~,ymK! \"ClIlsh u\ it}, of m~~}ft~ d~wti(ll~, \~!~1l!8~ fl· :'linllp~c dQ.Hl,1\ling ~A;;~l~lC !n J~I]1,'fnll:; r;I!i!,~k:"'.1pt ~~,!,~~'lr ! '~\~f ;1. ~udhoJl I(~ I ~(I~ tr!g. lq,PtImc;:~c.s \\I(':n1; 1;iI~. si!l~1.cc;lto 00 "R;mcl'i 11Irn1ig~"''C 1'3I1C.~. k:m'I~ their { i!;,'ill!,llh. wi ~h.diggn~llO'lll ind~t[l"{I!1l ·~halwere used 10 • Th~ I:;hcll1li~~tri:L,'s ~~'l~d Ihbr~Ci~l~ ~llIclia p1iJP"~ 'to ~doni~' >OOtlJi.: PO!\lil~gll!i"~~~he d:iL.:: fC<l!d''di[o;iCiPB'' '~i[:k~'t~i~a:Uic;: ,~- ~]lio~trft;si~lJ rr\qlllurn;~ .. dd il i[H~111 lk"f'iilP;: Ihl; ll!!lt:Dd~ 01i.CI1:I::i[~riJ (f'ii!} If: J. The I.hit, rn'lplili~ '~~iL;~ilii;h'iti~ 1.00 ,('JL~.nj u~[J;:: 1:J,ir.llti,Q~O~li~c:;. hjimg ~hapc IIt1;: ~>ilIl'll:dJ.~ l.~lpt\ !lligua llli!i..'1l11 lor 'Iv the ~tBrf~.cc. ;H:id, il~ lh.c ease (if s,lril~dl rod... s.eaml~I1!fl. irl !I fl\:rlw-l!1uQlIIgh dev:iJ::e..l]tc:>JJ[ri1~ ~"-'1I'i. Ilr.llio:nl of \~m'itlu;". cltcmi::·.In:c::...en the partid{'$ each ~11'1!..'<!al palh::m i1,cedl~ tn 'be ~n~rn.k'ti He one baAe~n!.II!! ~im~ Typicm.II)!, IJilc~~at~mt>otl a:II'O\\'5 doeod ing ,li.ndm~t ,uclcctiou tDOc acl~i.;:\-cd ~hC'~ 1.,m1i:c:1C'1' ml·ly l'IC di;;;lin£ll~IH.ld if Ihe by us~n~ Iii sil.l.l:.ie fluoropl1.-o:rc. can ~]\ICiP!Sccnce of IlI1c tilyci .~!gll<lll i~Stl Ilk ~'11li)1 TO. dcmm~Slf'itile the m'$:1I1~in.>,of 1}':11tL'CI~ synthesls, wre &C1ccH~~dyllm1'iCloo mOJlOn~Cr !l.tre:U'l:l)1. il i~l, nO'lil'-'f sr-aphflCa~~wt~INJ IN' ~nicrocal'l"i.cr A encoding ~l1. I:hc seloetivo phouob~ca eh~n~ Wilh a 111!flIii."Jflhorc ,lItd 'll'100 .<1 '\,'1I1~'of dm!mei (:(ldi.~il]1,O nU~:IITt,~t,:ntoc ...L& (/6).. In Ihi",~.li".~~~oo. dl;.'Sign..... !lltlil'O'm.i;; lo 1~~'Cr...~ oo.uin~ u ~ingk\ pJtIb¢ mll~:IP'lnidi; ~ynnh";Joi~~\l <an),d dlJl"t!dil~I:l "It;' lijal1.C~'~~O:rl. rll!,ll~ti,p~J.:: pmhc ~gi(l!'I.,'" ;,Il~d: l]mb(~·n;~iol1 cQn"lJImiv~:g_!. I~mking il an IlmHke~~ ,~.uldli,dm~ror gnidienn; Wv.:;:. tE to G~ M~!I~i!~mhe !}lII~:tli;I'cl1 i!i!!h .. ~lIDllghpLlt a:n!i!b'si~. A n~g,~dlOd Ih,~1~hmi~ trIg.· I F:k. mad~ wi!111 nJ!~ llVC' of" ch<!!ml):L~ wid! ~xi~tthm
~Q:iiIC SlM.p:h iC<lII,ci;hnique~~o ~l};'llgl!I!:y ~mbod t:lS):P'Ct1<dC;\ microen m'j,}f:)1. mI· O!1l~*!C~! lNrpl;'m;!~ rdy !D!~~h~P'lt~r!1~~!.lil of op!iqll d~Il1!!~~I~~ OJl ". lInk:mt;:nrr.i~-r; ?l!1111'~ ~x;itrn!)k~iirlchIElll s~:rip.,;tl [lid.. (/1. JJ't ridg.c-d P~!'1;:~~ ~U~. ,Iud dOl4~(~~~emr.>d palr!icl~ U 4, J 5).
A ,,~~ fe;:nIiJ~· m~r !lIlClihod is jhe dim::l inrorpGmtiml 0fl'lJ)o1lC8 into IiIw cncodcdD<l!'!iC!~~. '1l1i.~i~ oconll~~ .. i~ll;-d by ~i!np:~ ;Igl;lirl::l ~.~}'!>ft'\,~
~llOi;!iti~-cllJiomokx:~
AH ..!f ppIYI~lerilm:i:mJ, '
or
I~ il~to
th~~ ItlOn!lOlIl;;'!"~ml~liO:Il,
or
an; om":iili:.;mly OOl~p~cdto~1C; polymer' n~r\"'·mk lll~~. p~ll is ~W!iC<1!.b!C' r¢foo~h (ll:i:goI1Ul;;k;"Olidl; and rmh;ill pH~ t.M-32~ \lI;t' d~nl'irn~le II1!nlhl~ ~~DJt btirSl::. ofUV 1lI~ d ttl :'Y!l1lht~,i:H: ~rObb"'OOI~~I~~~!0d, ... ~ifli~l~ areirno1: dru.iinCi'lllll IIi Ihe IlUi'Ie'!itJi'lality of iilt\Ofj}Dmh::d ,L'}.II~~ti.~i..id.;;.'(]lid~Pre\l~rnis~y,. we ihe~"S
or
6"'lhL:C~1
:Ul(:
a!Ol!.llt".,;\11lcc~1\~L;'a!~ O'I.lI,O~i('[h~r u."L,.~ m!d,vo fk'\! ~!.;'!~.~ ey nii;ltiory mg,-,'l J '1-19<). lflhil> ;lppw!ilch ~~.'1..11)' \ W'iJ~,'\,:J'riJl 1ll;e~iIl11;~.CI.LI rows; .hbr !i!"~rl)l' !J l:t~imi[~-d ,
:;'(""\'\oT<1il
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tbr
,I d~!1.'Ct~ij!]!l~L!'
barcodes
scheme frorln 31li.l1~1!C: tlUrullHlc!ition (,I:l:iJQrcs~ eeneel, bUl the :~YI1.11~C'S1.11" rlny appreci:iibk' iL~ ~nmdJICr (1hous[l;m~d,.. or mi ~.loo.ns.}1or these decrate), mnagnelic mH1Ol)arlk.l~:;:;lre lllco:epomkd ~Tonlc mh:;fOC:h~p-Il;I$OOCaLTk:r.l. l'I1IaYr1'O\'C· 1.0 be in a g'r<!dicl'll. il maybe pQ.'<Sihk:m rm{]JJ;OC a. ("x~'lClrasivc "mci !\l4Jlw.'f~~C' :md S'C'\olC'rJloiber ~l~~hCid~dC\.'\.':~o;pcrlthrmu!lip~~~~'clm~(l!~ysi~lm\,"~ [(;nnpC'nlwlw '\·;)lri~Himlak!,~g I)~·~ide,~\\ hen st~lllullutoo i~ ~'CiIiUlil1igml1lg11c'1~.C fI.clcI (1'9}. illl b~'CI1tl10Wll:£hly re\l iewed elsewhere (25. 261.
Weullltro(h.llJC'Ca technique d~m OVCl'OOilm.'S ~mrly ihese m,.dti~~lcxil1g !iInit[!j~!o!1S. B:}' C~~ pk~~ljll:!il blllmi'I1<1r (lows dl<II'!l;:lt.~risJ.ic of mitltnnllidi~-ll, Wi; dt:nl{mSI~'a'[~ the :lilil iW 10 gCJlLll'{lW
p:;lrilicie C(1'.li1prui'lD.!i ofs~"\'l.1;n~1 Ulrg!.;'~ FUrn.li~r.. mere, probe grmLlk,tlii; (Fig .. ~G 1.~n('JdL:1J.y $untpJJy 31.l'uw~ng di.~:lL'>10nlofthe p.ro'be ::l:Cru~ S:tn::il11i1lS if! 3. IOiilg channel, rule useful for brortd,e:rrling flrl':: th:::I1:Clion runge or~llmmlyll:\";~IC~l ~s~l1g fixed a dcl"ec~iQll sc"~l.",il.i\' {'when !'IlC SigJ13~ CJ!I1 :s.mu~ly
~qur:iI'to; dCl!:)~~io-n 2. A ro C). \~'c·~'nl!h~~ ,~Fig. ~~oo !l!it'(!c b<urhCli. O'rlil<~f!idc:;;: OI1i.: ol·\.\'hich ,~.,'mJ !~'t1 ... ~~vtti~ 1b"~\;~Ill~cir Itbp~'O:JigiUll1(:.I(~).id~ 20 1Pmbt; I W-t\.'fA GCA GAT CAG CAG CCA GA.,)'ll :Ilalmh.::r with pmoo· 2: C,'·CAC "fA1i'fiCG CAli GTI eTC AT~J'~\and ::i.UIIDOO. w.iith Illl) probe, EO SCL'r"C HS a cmllool. l'~LI'g~S \!i'C['C "li.Oit'S{"£ll~ly labeled ollig.o.nuJ:clooticies Willi (om;picm.emlll"l!' roqucm::cs to Ih~l\vO Ilmhl...""S. w~ mi;o::oo ti1~ IWIkles. ruld.um;Lltmtcd Ihem Ibr .~ mill .>11 .l'OOlll 0 fClJlllCmlure in mimlw~~ils conl<!ini~lg ~ilher 1:lI"ACI
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<lu'IHlyre CltCOOil~!!lllndf...'i[rg(1[ CfipHl:rc Wig. I} [ni n Iyp:ili:"al CXp1!rl't1h.""'l1t, we likw,cd~wlI) monomer slUC9ms ~one: ~.OOd0d with ~ ll]il~"II't djfC anI£! Ibc ather with ;fln acrylal.c=l11nodifkd probe) ~1Ldjace~a~y down a, micl:ollu ~dicChrulllld and used a \;,ffi"~3.io.n (o~linllolll~'-.no"l\'Ii~hogrn!lh'y(27) of 1.0 polyf:m!rizc l);lniol~ ~willh jJo,.um burstS or ~lluuviQlel. fUV)I~£~111.;J;Cl'OS.l> tihe slI'Cm'l'ns ~JB) UllQ\lh! Sll. ,lin dl'i:s manner. pMilich:s wil~~ II 11~1\}fe-seem. gmphicailly CHl;oocd rcgl(l.ll and 01pl"OLlCIQ;:Ww I~,giollem be s.yl!lthe.~iz,cd un ...si'lfIgle Mell, E;L1;:hfl'Hilid~ i:ii au ~,\;,ln"I~d1IwO"dhm:lb~i(l~ml 1(2DJI -~11<,Ipc'tFi.g..IB) .,'hose morp:.ho[0£l.Y i:-; dl,'" ~.;:nl1l:im;;dhy a ph,{~lmli"~'I.'i'kth{lt is il1M.,;'i:t~tdinto 'iht,; ~T!i;4d'fi!:Op ~)U,r~~LLm:1 the 111:i.cml'ioop:: OllIad wl'iuS(i (If !.:llcm:i!S:tr'y u:; dCk'nuinoo, b.y 'tthc ClOJlt'Call ol'nhe
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n:o~'il1ig IUOllOHil.Cr :i'tii~ill1~. Til...: cr.~1ti:;;;Hni~cd p11~,y,uncrpmtides lh,;:n, no'l" ,C]'ownd1,c ,hafllincl [Wiihoul sticking d,tl;;w oxygen il1hilJ ~iol1 ncar ih~ dll11'l1:"I.;;I:;urtla(j~li pm, \\lh'Cil!; dn,.;y ~utc oo~~ Icci~d: ill. i!t n_'sc!,',toili.TIlt: r~~lI1j:ch.':.'i c.mb..:rin~oo of CKcessn~IDK'Imer and 1iIK.".i I u.'>Cd l.or biological <1I~YS. We ~df""Ol)'l,c~hylctIC glyoo~~ !pEG) {well kno~..n Ol!:'l <! bi~~fI!m f"Il1Yll1crl OlS ther<!rlid.c th!.fl'lda.ion 10 d~111imni;lhe ~ml'~ !!block~ 10 :<.~rtlu;~ aOJ;'f p.~J:]br:'lXil1j,ug"lio~ <Illd ~I=- 'llr.1~1."P'Ilt~ eiil. 11I"iIalqi~~lo ,111110\" .mUi~l1iis.~~O:ITi ITIltQl1I:,~tt111 or 8oigu'ia.1ltorn 1:~."HhpmI1.tcJCl~iOl~" n~(: pm,I~Tl:i~
co
IFig. :ill.. {:A) S(h~matk d~(l~r~:rn of clClt~Dd~etl Ililf'tide ~1{iiiI~n~ls~1'I!i)~~in:g po~.)im~rizadon aO'o~ t!li\(l C!ld.~~I!:~t lami!lii!r 'sn:f'li'amSW ~kll! si!l9~~P.['(I~i!_·if!{!U~~h~llrli':;l:!li'iU ~ar~d~s [shol!m tn (9)], «() .liliagraJmllllhllU( n~p.re~e~tEl'ti:!l1"l parlider~i!Itur,e1) for ellmding .and <lInalyle d~~:e~~ol1, Encodil:l.g ~d1.eme iJlf sh~~\ln g~I"(J\~~iI1~ gertera~iM Oif221J' t:tM8,S,76) ulilliqu~ I;ll:lms. ([ill' [J!irt€r~nti1al ]i'lt~ff.li'n~~(€c!!):ntrgst (101'0 1ml1l'QI!!'Ilf paltides generared by lIl:sing~he5()nem'~ 5~(I\mifl (A). (E to 6), O;i'erla,jl of nll.UJrt)(CI1i!!:e a iIi!(1 D~C ftilllage:5 o,f sili!9Jte"'Pir1l)!b~ ~~), mutUpiiObe (IF, b~tt!()nrd, i1m,d p.fQbeilrad~e;]1i ~G,l.:::,ft), ucoded IPilftides. ShQ!\i1l1 @l$(! lis, @ :Sl:hJ!lPllliltil: If(!prE!'Stlilitali(!lf1 'gf !llli!ttfp!Qb~ Pi! rt!d~-s (F, ~t;)p) ,~rl~ @ IPwt ·gf !l1Ll()re-£~emti~!.eIil:s~t.y OJilong th~ ,!:jen~e<r ~ill!e of OJI'gradil'nt pa rtide (G. ~'h'!)" S~ai[e hars m.!lJfcate 1100 ~Lmr 1r1'l ~EI)~ (IF), and (GJarnl~O pilm 1n (lH.
1
1394
VOL
31 5
SICII~NCI!
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REPORTS
d~'i;,">C1ion \"":5,inclll;;ai~d ~'C!K:~, which \\'<IS!1:1f,11I:l;! 'by Im~bol:-w~i(IJl Il!l!(lre.t:~
Pfl~lt;lUl'KOO
~icB~ OOgl;~~llli~p,."'lih :iil,l,~~ t;;l1~>iS~"''l','t'c liI.MC in dill.i,t'il;: mld hybridize ,~v(,'lF<l1 "",m ~ll(nih!ii pHl'i DeIC' h4:;x1y (].'l')" In 1L~i£'&~iIJ~tutt::c. illl:" pmtic-b
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par-
illmll~ unifmmiw (~.'!:) \)j,li.d~ high ~pt.i'l;Hich.)" 1.0 the Qli,gpr.n~ ~;~hlhilijng nl!O~"~nce m1~}'wben ~Iw i.l)I~'li rf~~rl't (f~:g.:Jq, "'[:no fitlnh~~ d~!liIlOlh r"'[\1" t~\,": ~,V<t,)<t 0" ,1tLf mul..... t~)k,dng :.d~L'Ill~, ~ fl!cTfot1ttt:1;!uh.::: ~n~l!;l SI..~.
qlfCflCC
detection
w..,~
or 1-gI1~ch:s
F l.h~~hitoil~ll!l!bf,'1';\\'~ \,'l;;;rt;: .. b~ ~1)simtll~t.,'tI'il~~1L~ly liL"'~Y Ibr uill/;:~'!NO ' rn'h~~ S~l1!J!~'lI'IL-';:~ 'lwhlrlmJ D1~!¥.;"'~ tt.'Il~trol) on ~I "i:rlgl~rlirlock_ A.!:l,!ri.Il"lht.: ~Ih"~ly ""ia~ hi;g~~~)' ~ci~c ,Wig. 2F~ ;,uu;l '!r'C~' uniform
1X1\\oiL;'1;;ti~hc:.
rrom
'GCAGAT CA.G CAG ern 'GI\-3', er 02', §''{AC 'TAI 'G,CGICAG GIl ere 1\'1'=.31 er uno prolile [1!I1!~t1lw ,oelll!Jr'ol (()] as :\!h e~~111 :S(hW!~t'Kol!my if] (AJ' cC!!l~ {E~. ShlWlll- a!N~ rep;r~ ~entalive UlJlme5!1:eme imagi£!S r'OFl5<fi'ig~~oOO (8) d mi ml!lltiiprll be {D)
IDjd (!:iilillt:r 5!ltiIJk';S. ool,lkl be I,I~~-dUll' c~tC'.i1!l\it::t[<C't ~~'nlliiplc1I:in_g p;.h~lili~. ~ Ii' uirlei' lO j'!rov~ ~UlTlhiJ::; m:e;tI'~oo of mLi~l~· ~1~ mm~ysls i.~~mcl.1:c8~ loli' hi~ll,,:tin:otl!lhpul ~pf'liCJ.urol1ii;, \\'C developed a ~in~'PlcsChe~'le io :;e·m~ 11"lJ:'lich:osh~ a flow-lJnuu~h device (.Fi£. 3i). MLVI~ip:l"ilhc 1):!.l'I.icicl' l!,;~d ~Il ~I!!!h>'blidilalion 1:';<1'~x'fi n'ttln't jW!S! dlJ.tlcrilxlCi (.Fig. i.. D'lnfl \VCfl(l [kl\'~I.'<I ~~!tI\I1:h a Ilmicmnuidit' ehrumc1 ud ul).~.",~~ 011, <In lillv~nit;;'tI nIilO:fIi:~('nee mil;I1~~~lf!(: (2il'~ PMicl~'l \\cre ,!ligll!il~ l!!>illg Uow=~OCL!~~~lg ould1~ro.!; eled dO\\\n a cl!<'I'l!1t·~ oni~' 5-ligI~~ly~-cr ..
dmn
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down ihc' c1'llllUK~5 wi~ht1lill 'iai:(.'.n m =:I (k-sigllilICd: dctcctiio.1fI regjun in~i~~ ~11ffiJ111~1 Oil eXfl.QSltl'l! \\Ii~h ofl t ~25 S as Gile'Iliuticlcs jl<1Ssctlll:c flCk! or \'~
'(us~ug I), 2U" o'bjeclhc~ ~X1lJ8C SQ1]UCI'lCl:':> were bier :lIla~yzcd 10 :(ktcmlioo!hc p<l!f!'!id~ code :md
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(~TtO\l.i.c' S1).
~:lI;J:llion~ ~oillitainin:g Ino 1:a1l!J~ts. ta.l!let 10"J![y, t~~~ '!Jel,2 ooly, '01" ooliill<lrgeh ~(Oam:! mJ. Sca!le b.;il'$, 1.00
A rcpr'C'O(laf~;lli\f(:)P'I~i.c lc imilgc 1s)lhow!l (Fig. 3B.} 'W.it~1 oID.c--sponrung :b]!ct~sutyp~OLsa~mlg o I1lc fWlC I"lr:li,ck ''iLt';!It:.iing .!!rubeS." Tile eedc a~OJrlg each lane !I;.1IJI! be 1.1otc~minod, by an:;:d)'zUlgI1JC !'il!lilap dips ,mel P~'I!L;,.all~ iu~hc iu~l~ilhiily pJUl"i. By
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a pQ",ilivc laIfJJ,~'u (to1.ccl!iorrbas Ihe conlml ;}\I"(,i'!lgc ili~nL"ity ~JlliS l~afitt SUH.l'ILlnrd irninl1'" uhr cfleh dc\ pMlid~Wc \VCIlC :l'ilhi:c [0 tlCOlII'n1CJy id.cl1Xi~.Y~he ~C:ilCC ofoolh oH!,~.mu,l.dCO'!kle uq;cts 3.1kr only
a shon I {l=mil1i :~ncuL"ujGIl.
rQ(lIlsing m~crorhi!~di!!:d!evio@o lI~d to idigtn and rel1d pa tides aftew hybridil,Oitionl '!l'xpe;rime'~t5i. IFiilrtTI~!~el'S ,Hoe directed dowm <I f1anm'l 'ch~IFlli!e l ,al'ldare iD'!a'ged by ~~liii'~fliL!ortSi:enoo miC:r'o~
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is luimilrily
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lid.:: size. 11'L(; panicles .:symhC$iZl..~ for this ::i'ltd]' are n;ro!lve~' t~.lI'g.t (X)rnllrul."'IdwhJ:l (hQS~ in 01111;1:"
H()w_;~I,lrolllgh mC'lhods. mC<l;,~ur.ing YU ~un i~l wid~h.-30 um in.lh r:cknc;~ and ll'lO l~ 170 ~I II I
d~l«tiOImI
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~hrol!lg'h dmC!~ as; s!h1l1U1iIilli W ..The ]1ln:ageIO'li1lI, [,'lp!lU red b~' I!!:s.ilflga micr,oscilJpeIiIl@UlliJ~ed !!:illll'Lef~ 'MI~ ilni ,expo~'lIrre of 1I:1L2~ SCi5~'l'e pa ritJ1de f!OIwdllll til 'vel«lty ,CIIf ~12,ao 1lif1t'1i5 th~ou§lh Ute
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,cha;r1li!el 5(:11;11$iO~ nUiOfes'cern inlten~illi' 'W@!I1! ti3!lrorli aJO'os~[i1e rive laliie~ o:ftlte lPail'ilijd~ I:() ~I, tlih~ au:fll'
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(01 <I nd l:m.w~th p.a~. the :iilOO ~ &!C' lid,~ 1111lhi~(I'Filmt(!liorl~ '~h~ t pi~el ,Qod!E1isread from riight to ~ehan:d ttI~to bo'ttl'lI'i1" \lfh~~e I, (lI,an:dx rEp~s:ental ho~!!, no hole" ali'id an aligiiiment~ m>a~kE!~, rt'.5pe(:tiwlll'. ParitJ1de ~ho~~llis '90~m !by ;270 pan. A.U., arbitralry un~~.
\'oll,llm,::. rn.lo\ll\lw~r. ~hc gll;:,n[.!XIr!ick'"I(I"1I~m:L'tk:~(,; n;prnG\I:I;; ibi li~}' "''''' hm ..~ d~'!.ll(IPl~!~·'l~CtI;l (2N~ wi~t ~1ilbJ-d 0IU1111,l!;.'h~{)wt.:r Iii.'itlundmu.:y dl,m i:. l)rpDLo:d, in nf.IW.,;:hr,o'LIg~ :;YSI!';ms..,lflilproving ,eme~ ICy. By uing cOI1SJCf'vMivc eSli:mrlfl:[cs. we :1~'liI1.dthilt !':'i!J'r~"Vs~cm should be cillpilbh:: oj'provl.d'~fIIg mpid. I'l~gh~cnsit), ,m.;lllys us 'willh a m;jm.;lls~alb~csemTr~'()volmnc: ,t28), despitethe ::i'!..",·miulgly ~f~C: prnrlidc~j,...,.e. In <JiddTtim to be~ng vccy t"'"1Irooliciblc'.wc h:wc' a:lw shown II13l OI1!lI' :Sys!e~n is ve.r.y sem;lti\!C'. With 3U=lnh~ inclJlb<Jlions,we call dctiocl DNA o'H~rnnCiS co!n~Grn.'1llh' a~ 500 <It!Olm~les wid1.m.l!l l'1111!in .... ': idin aicl~':d.~igmill an1Jl'~ik:~~io:n(2~), lhis I IL;:iLd~u,~ '~L' bdic~ [ll,U! our !'1y~lJ;.'In \,viU be .n ~C;;L"'!
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liA. ~ L'oIiIM. C" Sewher; E. I"n.~A1i~tly ~ 4125 112003). 1!.~ lIt !llr,a\ilt;~!i!!iI~ ~ ~£.•. ~; :lll. IE. l~ MDrali ,N lJl..}' .. Am',
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DeiII: Jec/ma1. 199 (i!:'O~))_ 115, 2. L ~lll,.Y. M~llit~ Q. 1H\:Ii~n', liL ''1\lg'j')iy'll, AM! GlI,Il'IIfi, MIter; ,~ l6·9' j200:n. ~«. 't'll:;!, 1078'7
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Re~e(l'1!Iii(:es .Uilltll N!OI~e:5 gliGer~~ !IWf#~ ~,a5 ~~OO;1t ,Z,. S. 1". IFQIliOl' Il!' al. NMum 36'1, S'5S fl9'93]r.
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s ,G. ~Ele<!lIn" S, t, Sl:!hrel!oe!f, SC!'~ae"!!69. l.7,~(11 t2Qil!ID. 4, ~ j. FrUI.Im, It L. ,Mcillace. 11'. L ~itl:1, L F.IKi~k~!, lit ~1lIii11~I1I~r." Clfn.C"tm.,4'~. 11.149 r:l.997~'. S;, ~]. ~!liby't'f~~.J".4m. CI!em. 5Q(. :J!,2;::.21:!J3 {,2:00I0:1. &. ",Xu tI (11,. M.IdtiCAl;~ ~',. ~1. ,~a {,gOO}. I. lit. Han" X. Gaa, j, L; SUo S. I'fji~, Mar. iIlrJt£clrf'IDf.. l'jl, 6H 8., .'!:.., \'I" hlii~o ~l£It,
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c.. IF'lnegiboo. J. ,CoIJlifn, t A. "'aU~!l1. P. 5. I[n~re-, N~r, ll!Qtr.r: 5. 3:6'5, QOO61 219, Matml'ltl. ntd 1ifi~lho!k ~'re- ~ai~m~ as ~~poiilnnQ malEna'l on S{lfm:1!' Onlim~, 29. R. '~J9~ ,f'( gr", rfEE fl~ns. M~51i'!.~, 3745 f199i~ l .~O\. A.~. Itou-'blm ltili. Bl{Jt«~~ 3:11. 1008 i::lOO3], si. A.. V,,\I'a~ri~~ll'".~ rGlr., 8i~tKh~;[rj~e'f ~9.2 f~~99l ,no n, F,. ~l. f;l~"~n ~t!!J.. 1!Ir,rdti( ,1lf!;M$1I1t/.. n, 61l'~
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IIDI'verse R,ellatiiions Betw'e,e!n A.mounts 'Oil .Air' Po~luti'oDiand 10rlogl ria Pih i,e: IP rec:ipiitat:i 10 111
1
1, 20 kill east r Xi'un .., Ch i~l:l tOOL! I~OC':llio!! ill. F i~. l} poov ides a complere rceord ,of foulL IrlC9StlrcrncIUs per dll.Y of all m~lcorolo£ie:l:1
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Oi!1niel [email protected].:h ]ll'IiI [ltd,.::!! Xi JIg l{1il1.:~ ,Zililrllillll¥1i!I Ya()I,~ Xii~ollilollilg XU',2: Xirlg ¥arl!iJl.:~ (11il1!l!oi'!ilill~m Diu:2 _. P,]rHIC:U~ilIte ak poUlllti®1l ful,al$been slIIggested as the (a'LlSe Qf the I"ecel'lny ,()b~e:rved decreasing U~l"!ds, {If 10 t~~5% Qliltne ~aUiiO bl';lllll'!{llI;:'Il!. MH.y ~1[1d ~Ip\'ii.f!d IgW~aJnd p"idml;atf'Q!1!" dOW,fhl,fil1!d I)f I!.uballl ~!i1dl~dl!l$~r!llt r!l1~eaS, ~,ql,lal'l~ij~ijoo th~ dle~~,nden(~ Qlr lhi~ 1",;;Ilti~1 ~he !I),roglr~pnici W Qf p~edp,it~ti()n~ifnhan~arlii~liilt f~(,t:or (I!iil ~h~ am.OILlmf~of ae(I(j5JD'ls i';(lilI1Ipo$~d' mosH.y Q~ polklti(ln in UU~ free tropo!splrier,e,. bi!l:sed 101'1 me.a~r'E!im~,nts at. Mt. IIHLla lIi,ear Xi'aln\in (ill Dlb'a i Chnin(ll. nil:) Ihi~lly predpJtllition ~ill1 be d'e(rea~eld by 30~®, 50~/o d~H~ng hazy (®lldill011l~ \\!iilfulViisi~ility®f le'~ [hal1r S klLome'lers at t.he m®Ull'Il!li Iiltop, Th:is tren.d' SOO\'lSthe ,role ~If d~r p oUlui[iicmirn fhe ~OS:!ior sigl1~li(,mt wlIter U~:W'lilr(eSin hiUy iHetlIS, wh~cih~s:a liI'IlI~,or prQ!ble;min C~imllam;:i I'I1;H1Yother illr'eiil!i,lDi[
clements. ,e:ric'ef;l~ [o',r OIllC' measuremenrper !Jl~y of IllfCC:i.pil;m h:m dIC,plh., The 'i~ibility (V IS I LlJelhre IQRQwm;,n..'JCorued as. I (I classes, ;lIld !IH! values ;lIICT I QSO were '[!;"'I im,lIcd d irec'tl)' ill ki~nm~t~'fl\"wi!h respect ~o 1,IIU~!lI;H'k~,of
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nrdcrto ro(lm:cl]~c eBcct oJ relativc hWl.midi'~y{RH} on '!;i~jbihl.y. RU V;!!irlu;::-; grc,>I!~cr~han40 .auld less d~an 99%\'''C'JiC (OI~A n.;rl.1,;'(1 In the cqIU!V:[II.h.'lill ,,'i,~ihi~:il)'in dry condil.im:ls (iJt., R~il<40%)" \l\lb(;'iJ] [IUri <40%" the ,,~sibillhJl did :nm. need to 'be corrcetesl for
r61f1li1
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th~ Wo'r~d.
arge CYUlQCIfI1m~nm~s Qf SIiJII.mliCl'Oll pan..i.c.,.. l!liltt;!!ir p7JlllU!lkUlIlOl\"C been dDC!.m1'C1lncd Wi SUllp~PI\ccii"hmiou"tbrmiJlg ~~ by se!'ving a:-; clklucl.·dmp oond!;lI!~,~~io~1 n~aclci ~h,~t uhrm dou'~t;; with "~~lnn {.kop;> Il¥t~ 01X\~ !ilmll'(.'IJ· In <l'l.l;Ukl'iCl.; iuto r~~illdJ:{lPl' nd W !li;.u;:i'lK: a i:1'l1O icl~h~ldTI:lln~I~~}J:£ (J -41. 1111si~ IlIiIillli re:;,!C!.i i k,~ prt"C~pitafL{m {lJ'l tho; ~roll]ltl wl](.'ilnhe
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11n:s.m~Ul!i! of Eill1h :5ct~mf~, T~ll: ~r~ Uni\ol>fsrty oj' !~~II.i~t~1l1iI. elU5<lLtMD., IISlfa~L '~Me'lecl1ll.ogI('.al In5l~IIJiU1 ~.f J :Shaaff!:til P,oo'l'ln~e, X~··!)I'!'7'![0'(H5, 'China, lKey !.aoonaI0l~1
~[lr Cloud Ph~;(s: .;md W~~!1I11f ,MOO~iK~!JjOillo of (IIln~ MllleDrotogbll!i§';'IOCl.altioo, Chilllm l'!c.;id~rny 6~ M~I.i!Oro" logl~~ S(~()e:;. Be:ij!~ [email protected], (hifl<l .. ·Vo \~hom oo~~e~p(!l[nl1k:n~e illlIOuI.dIbe addlre5.'Sie(iJ, IE',rnait
dani~Lr!lllsjmfeld'@'~~"1!Idl
(lm.e when fon:tld d~\Wl acmss thct'ildgc line. Thcrctbrc:. dccrca,_<i~llg ~ljellcls in,~hC' gmgraphh;: meaCnllitl!liCCmcl1t lhctOlr of !hc prccipil3tkm m(l~, mmr~cd dccrc.usJng trend fol' I'hc ~~'~IQjc wh ieh is ~I:!c r.nT(n tJ.clw~l !h~ hi~~ tQuo ~I,!e i:Hl!r'C!m.!!Hp!:ll'10d. Thi.~ I~nd rcllcets !hclrr<:~1CI, ~I~)wind ph~im;;pre~ill~I.<i1I!l;In. wc~ s!.!~gc:';l(d i~~CiI'[;',!8ing "i f l)!'l'[hu!.~o:U! ill ('~ inll ('9. WI d~llring lhi,ul 'ulnu;. ('rm!;,~M(;I'II.w]Lh ~ho;;: hypolh~~) .f(,: fI(..'tl tih~ tl:cnd of iUbl;r\;"L"I il:lg ;Ii r p<l'llu~ir}]l diL.lI:iilljg_dUI~. po:::ri(J!;~ (h Thi~,l'I~'lKYI:I1l:;si!\ W~ll:; o:sis 1.~I:ilt l ir ~mllrulion SI!lPI)!',"'£iC~ Orll.~lf~~;phie ; Ro ~howl\ :5I;JlislicOllly Si!!,\lliHi"'l1J~flo:rl.cd by the dOCILJII.'i'''llcd r(:;dllcliomi oj: pri;cipi~an.i.ou, 10 to .J5% .in the jJllSI hllllrcC.l1hD~~ in ililuChr 0:1:' emu dC'C~,r~inIS lrcnd" of 14~ndl 17%.. with. l'(:sp(Xt l.oDhc low-,I:~\,'cl fa iing,auges {I r Il:tllfi}' in. the fIi~(;Ilmlfllifil ~:i;UTlg~S d'le \Vcstcnil Ull.il<:d St.lles. dowll\!l illd of ull~jor 'LlrOOI!l and !nldm;:- ~l1d X~'aH {$ce locmio1'ls iii! :r-ig. I}, I'('Sp(>Clively F~g.3 ,. triol ~ug\!C'tI~ f1} I ,~OWCWf, b~c[~IlIi:ii!.': u~ack {5 of o r hi,stQrk: .. ~ records of ~I,mbiefll Fu::rosols. it The c @JIJ~ ror iihc J.'I::>.~""i3'lioitJl f vitOihi:l,ii~' o \\';1S I~OI 110S...ibllc to (1]11.mifltul,ll.1iwl)' . .,ella!.c i],I1.~ .. d Ito l:reltds C3lll ile-I.\!5l.edbr m ig u~{)rillg the lime racWr :<tFJu:JteslinC'j Ito diwclly. a~ ,I ;H1Ul~mlsof ;t~r pOHHI.i(l11, ~o n,o. We LliSca a. set G r n1eJ::J,turemen[,~ 13,ken ~~ncc' 1'9'5"1 (JIll 0), ~'Inc~k'lll of the ds;ilbmW disl:mo:. This was mOL!HtOlinlop in cenlral. Chin,,,,~o qll!."'Hti(y ~'he done I'm Ih!clJm::mTCc~cd vi~ibiluly ~.F~g. 4), imp,~'Cl l/! r ,acrQ~1 !c:i11.,ding on 'prcdpi 1<l1\ion in kcm,l!s(;' INn'}, ,of '~hc i"'~ipiuOln~on ,,,,,,c'nt~ .m: ~h,I't ,;n:tJ;:a. !;xclu.d:Cd wi~h the ~"Ojl'()tI:tcd \'~~.ibili'lj'. Wr.; hYPfJLh~,~;tJ; I.hal (I.:dlllc(\d ..i8ibilit}'~luJk~~~Jl A. lIl'h,:III".'O!"oi.o'gical ('Ih.<;oCrvOl.I,[}ry ~'L 'the top
V I SIV~S IT dry} = 1J.26 f 0.42&-5110g( HJO :rU I). whereRl I is irrl pereent ami Dl~c~l)gison the basis 01'10. If there was fog or preeiphatlon at <III 'dsfbility o'bscrvOll.io.UII t~me5 in ~ (_lay. the v~sibililY ~h31 .:Itl)! was excluded fr!!lnl ~hc . corrected ,'~si~ility setlcs, ~l':llc'dllily Iil\,'e~£:c , values arc presentcd ill Fig •. 1. TIle \'islbBily ;ltlil!C' I1.H1lmt~inwll. shown In: .f,~g. 2, haC! a
or
nr
or
or
ML i-mml (J2"':n'N.
mhc rlit~~~"cr
oJ rilo[("
3em$O~)'! mtd
c],C!~id
REPORTS
«('rN) lho.~ reduce the lPn:,'!i;ipi~<llif.ln over Ml. ! lua, U' this hYPQthqj;i~ i" i!l,~m~t .a11d\'i);;ilb!~iW ~'r.J<~ ~Nhi~!g ~~ the
C(lIlQ,ens.ul,ClI1 mldc]
M.1INI
If!n,.~ill"iwiiorl !l1~,rnln~:>. ~L iP~-dI,I~~~ v i!o! ibi I~lY :dlim~l,cl: ~~, n1~illly!I~5~1:l:1 of g)r'~~1,i,,:tR.[J}-,IIliTld nl0f(; 1~~'(;ip~ !~iQua.ul!1h",!ocal!o!; Q r v!",jb~lilY ... Clb!;lc:r..t.. ~iO~~N I( iML Hu;,!!). '[hi iii r~I(~I]OI~C~OIl ;!i,ilO~illd !.:'~I g; !J.ni ~:s;~m:: U iflliu~! uf ,d:~'Creli!s(jd ,'i.~i it)' w i~h im~l't:zlwd Pl'ot.;,'C ion n,!c.it bil i[piuu ih!.:nnmlial'la:il'k. wi~h respect 1.0 l&le reference l"fCeipi~8li(hrl iii! 1,I~myiln,.whiehi::. il.he elo:.~sl 8mJ bcst·col'rellltcd min g.,aLl~,!.!i.O L\.'II. ~:ll a, I
result, The
33AliNI
!r'ilg. :1.fopogrnphn:( lIIlilp !II the line,,!. lhe height rfFl~E1'l!elilt11'~!m[I~~Cmili, o,f
~~!lU:j¥
(OJlIDUrl],
~ao '(green),
o'l'cr Mt HwJ,1l is :J~mosl dOLlb~,~ thou nf H'uOB.}!.in ijn rd,ji(!~"!('li)' gO¢lc1 \'~8ib!Hty Q.r -20 km. 'n!OBlt rnt~o d(;c!;~JL'\¢.'l and lh!.;",mtol,lul ·of p:r~~il'!a'<Jti:Orl ~H MI. HU!I
ilI'I~I]roac:hot.~ lhm or Uuayirnm
a:\i~~I',jJ!s,C' I)rccipi~.uioll
\'iZ'ibiliillic!l of :lIlhl unci below. TI~~ c:lTcci of I.JI.~CW<l....!~g Ro \~fi[h fil;'dI!JCI.~d ibil~il.y dUl~~to !,mh.~ll1,;~ld \'~...
in!;: OV!lrpO'W!l,~OiI hi.; ~~·P{"C1.00,
;l.erollo~~ofld
Cppos,il~ bias
of
'-=----
\!~sibili~y g\;(;r .ML UUill:fll times of enhanced pn.·dpi~9i.i.OIl and RI t TM.~ OOSJ.!I'\ '.1011 Sup3 IJXYU1S ~h!!' suggC£tilOil IhfLl the' association between the ~rend:s of dCCIl."t s i~lg: \ isihUil.y and Ro is cilII.lscd 'blylhe hnpacl. of I~(! ('11~mm:ed aernsols, \\Iilh tume cm""'.illg"lre~~dI uf :>tI!ll(lrc.-;...ion !n Ihe Pl'IX i·riil..!t~¢I!l'i)VC'W M'L !~u:! . ~hmis gI:Ci:li~Ci' i~lan thaI sI'Lo"l,n by the plains .1Th~n gauge. which is m~mifestc,d as :l deO(:1iC'Jjl,l~lg 'Ire~d
tar
10
Ro.
Ct
19m
'~9M
~910
1'980 Veali"
'~900
2tOO
......
199C1
,iiMIOml
Fiig.Z. [rei'id~ 00 lA)~h!! (liJ&tl~d vis:ibiJuty .l! f'ld (B) tne !lO!Te(;too \li,'SilliLl\y' at Mt Hua d~rirt9 tli~ wl1~w '()Me:lIVatiool perkld,. Ead'l lPo~f1t w,!!~es,ents tnE!!lIVt'llOaged vi~ibil~t)' 'ror 1. ,day. iJi'iheljnel'lr 1~l!fIds ilroEla'kl!l'l.t),t:e!lil ~ep.nately forille periClds b~f!lJeM5:n ami alrter ('Vm5l1 19'&0 ~m'!i!nll1e ll~e,rv.a/~®fI rn~thMCI!o~y \lI<l'~ (h~f1g,ed. Se,e table, Sill. A ~Il~ B" [or dl'!till.
1 ,
I]~c ClU~'C:;; iIII F~ 4, R~) de(n;'.(Ih'i(.oD fmnn ..l'ITtoc'il 2 ~ p!I.,'riocjis; of gOfH!I ",.I!'i.n ibili~j'~o u~cnrly I Whl~1Il vi,~ibilily WIIS less •.mn I ~ km, Th~~~lkulllll1i~d average lIo lor v.llSibiliEy iutenlflls ".'!l:>fI1bolltl ,65 jn periods o:i"gooel, '\'~s,ilbiliIY l>20klln~mdl decreased lorlC3rly 1.2 ~llr \~is,~hml,~ less ulum N km, Th is mremgc or
MUiUil '"" IHI!J~)IIi'I ~ Ro ~l. 11100iY1n'I
AocO!dill!;!:
n_
)I' .. ~,.
...:IL~~
~~
-')""
-!Ii"'
1400
12'.211:5,.'O.DC!J5i:!,191
'MOO
~-~
HIJ.~i!ilAnli'l'nml!!lJ.Ro200l4JR01970;;: 11.31"
2: 1JJ
,!5~.::l1ll.1:l16 IPl'!(UI:H;'2
I••
...... E4!
'1.6 .:llI. .
lei
'11JS~ 11.411
1.. 4 '1.2
:.200
1900
----
1960
~'910
Yealll'
1900'
- G.' 2QOO
'.'
1
,260.
11
1971 1975
~980
'V,e'all"
2006
<1),1
Fiig.3. Ro ~
T~(ld:i of
'£I~Wfli1l:S
@flrn!!ii!I. pftic\~itat!~n
@Iif1;Ol!!nts
!I,l~Hm~~ bVlh~ regrt~lol'1 line at t~~ refie! Qr lh~ '~mll :s~Pies nd lWi'!l thlii e beginning ol~h!! !iE1'rie5~ 'ThEt ~e!lrre:wi()f1 e~PJlatiitmliIw tIiIe fmall lines are gjwn aMv!l!' tiUl! top of t&'lilh piot~~e rnlb~!l!' 52, A.uul 8, rlor daJ~.
REPORTS
IlIn:d H'LJ1l<yin1l!'S a fu nt:liiolil of the vti~mlY gt~~t.Hu'a for 1980' 'M'I 210M. Tille f@C ,i3!lilcl b'laci!: Lflrle:~~r!l
~!
"n'Ot;!
E.1rt~I\~
'~o!,
20% of the clot.:! (-1()j)IOI~ TlOfl/o !()f the data. ;ilroliln,d~'he rurrent P,()~i'U:). R!o.~ich is 'ilil~ dilStilIl!C.e' btlWOOrl th~ ~f es ",kmg titJ~ wg:a,n ~m~mic (ird~nat.e, is Ire@(ed f~lr
~ '~Ja, ~
f
:1:::
II:
Uc
.'C
~:
~ ~
.~~.
011 --:2:
2. 1)_ ~m~i!ld, G~~, ~, IE'tt. 2;6,. 3l!D$ !~'l9?~. 3. O. !!(J.~,rrf,etd,5{ienr,eo 2;67 ,.11'9:1 {~OOO}" ~_ I'I.t O. Alil!!neae ei aL. S~"" 30~. 'IlHll2@0<l1, 'ii. A. G:1v1!~lO. RO!f.nidd, J. ~. MilMNlI(lj. 43,!!ffi:e. {.2,ooo1!~" 6. o. !l:Q~!!i!f!~eI~, GiVilli, J. ~.~!i!~NliOt ~S. (l,l}]: A. {.2006}, ? r, t. JirOl~, W. fl. (gtton\ J. ~. ~~ 45,. 2:36
(2006).
n~5'it
2 5, and 42 klTil)~s
~E!r1lJU!d
by the
~inge black d~des ,lIiltn the' Villlue's onthe Irtgt!t Q~dil'l,atf·. ]~e5-Uipp:re~ S~Q;III effect :KIt!! rates a~. .. viflllllUty Id~slame 0<1-8 ~. :$,e@ t'llb~ 53, A 'lIlIldl
........... _--
_.J-
~--'
.~
B. O. A. ,(M1mlll.. M. IE. 501a~, ,I). ,,",YOOl'.'. Immltli1'r Morlif, 17,14 (ooo.s.),. 9- Q. JiI'l~~<l1I.'r/, ,L~O!II, ..urn()$,fii~rl.. 34 .. ,OO~ ~O'. ~ 1M,Z1lIl!!~ I,
a,ror
n.
d("CR~~ ~n R.o 81mlll 30fif, is; censisrcnt with the t111diulg:o 01" ihe case SJ.L1dy reported m tilDe ]tocky MOHlltaiiil-tDp obscrv:ll.ory. where D. sory" redloc Ikm hl the SnlD\V rJl£ \'\,'lIS rdmted 10 aerosols ~Iml caused ~.:;m;(l,~lc'reloud 'OI'O!P'$ .mbd redt!J(:ecl cfl~cic!lcy oJ r~hci,r ri Iuilng on S~IO/W"
or
rJiw,. ~. r, Wa!1g, (lim. ft,l~~'. ~~jJ)', 201 tWOS} (iii Oii~e:~). ft. fj, l!i:il)'t 10. H. lJDWil.nltllll. '5:• .i!i. Colin, W. O. ]. I!l1ii\iml Gr:rJ'p'/ryJ. ~ l~. )0.. 15
O. !!Olenr~hl, ], J1;tpl.,II/lL'~1,
(.:lllOO~"
~a 'c,co:!'-
~;!- I!., ~~
~~, :~:2l}a
The SCII1S,i1h i~~r Ihe prec ~I)ililliml from 0 o:nJJ:l,rnphk cJ:oudsw the delli nnC'~mllimp;ld of tmlhnio:n .HCU'(lsoL~ is not (111(H"cl~d. This. alsn
~ncan~dllin :-.,'1.,'(!ding Ih~ :si.llIlC elouds wjlh aero-
U. (.Q[Ilmnt~ 011'~d5nn\!t
l1<!!k'l)S I). "Ihese observadous sli!ou til m ~ IU'oxy to CCN cencemraueus in the free b.'QpG:ip.lllcre ~11C \,'islhi!lit)' at dlC wp or MI. l]ln:l.m. 2fJ(r[]1 Rl
<!
COIl'
preciip.itation
rnn
Fgr'~~ ~ I:nen~ i!!11 'C:~r:fI<!1!i:, (lijjfi~~ RI!!!~r~h Ct.mmme~. M:nioMl ~iid COOiiIol, Ratiirlt:rVi': F~ ,aj C!'lJ1':llfj~ aJ'~ f~f19 Ca-~r ,ol'Jd Mdi~ {IInW1\!'.Nrlif'~ '(N~qiIi!PiI~ #!lad~fI1i~ !'111!~, 1II'~l:JirJg[o!il, [)(,~:ij. 1.4. 1Ilt!~ 1m'~Il!l'llJtedlll)lbe ~iltki!li!~NalU.filt Silie~ee FO!,Hi~!l'il~of Cft1ip~ 19t<!!1~ 1I!(I575'OOiIl, the (hin~!!! "~ililt~try1 )(j~no~ .aDd fedin!illOOO' (~J~ma 0
~5~.
t.oo
rtS'~m
:;JI~m\'c ;.1(;'<1 ~C\..,C'I ~s; ,dircCI.ly (Q:IT>C'i;nod \'\/'ulhlhC' di.'t.n;;l:"i'in~ 1~l1d of Ro 01:1 .CL cl i~l1m(~!(jgiC::I'Uy tQH1trlli:tyillg Ihe: L'l;,;:';PO~!~L:or~t(;cipiI3[OCHrn 10 II'Il(;'IIl~ng.lil~ seale. The (Ih:;;~rr"cd sup;p~sitJlJI or 8(:ru....n,1;}I'!nouuns, 1JddlC'k~s 11r1~ ('CllOJl1i1l11L.".nclflllioll SiliJp:paningl p<.!C,(;ipil.lii.Ti(mm Mt IIUlli. l $ only nll1l~ ::liud~ ob-
cnhilnx:e tile rroc.i(lilatkl'J~. \.'ll,l)t_>cillly unrler (:011di~ions wh\.'fC Hll.:WJ!CI' ~~ l'C'llb~'y IromiJ!ir rolhnirm aliC' Ijw grcmC-$1. l]lis]1;1;;) ,llre<lld)' been :\110\\'1]1 An '6:!G l1~~ C11~ in: l:rimd U,n
200S0J8.3)1)99").lm@ k«!~flJlbpm ltiT~l:Iof Shillaru:il P~no~.,. OJ !if~I~e fflele!)r!ll!'!!9i!;~l!;!!li.i!L! d' S!lll~!l(!j 8 p~no~. W~ ~ir·e- r~t~1\{ll,g~l:i~I','cie~~~~[(',,1 g !1'!!!:(ffiil~lOO (fJrlIt.r el ~~!1l!:~ IPf<D-m.o!:li!<r pr'D!lid'i~£I~U d~[l! irllillis 'f~~rth, CIll'I~:i:lE! Matelrriial.
i!'/mlJlfill:lO
ill China" but giwu the extensive occuerence of lhis Ilru:no .nnelfltlll. elsewhere in Illc \vmM. it is Iikely (l~aAthis is II m:UiCir eeuse ~o.r slIJ)Jlresslng 'IOOCti lllmi,a-ll m I,he nmOUnllllbUl p
scr.vmicm
i
or'~hc U.S, N3I.iollm.1 R~(;aliCh CQ~i'l'cill t.l-n ~lmit the: eeueept of rfLtiimive lortirn£l should be expsnded lO~neu;cs, of IImll'il(jllltilvc lorei ~Dg.sueh as, those caused by~ I~e,efi"ccis (I r aernsols Oll
1I!o"M'f;5dMCem.:l!l.Il~gilcOiillenlt.rLillJ3151"5;B:l
Tlilltn S:1ta ~~ SOO ]"~!:i~elJegJi'flm .?3 MO'o'ett!~eor 2G~ aWl~'ed .~ ]aJ1!!~IrI" 2l)01 10.U2{o!'ocilffic~"lB194!9
()1I11,kll~'lic0n13inil'lg d s DNA g,CilUln1oeS. such as herpes s~mplc% viruses U~SVI and l11.tLrine cytolncg;a:iovimg,. IlfCllCcogaliZ1!.:d by '~lR9 lJ. 6). W~IClI)I.'<lS, til{)sccml~<linin]l, ~ing IC~SII'lI!Jidod ll'I'.N!A l:ssItNA) g~1l0m!i;'S, stllch as hltllucn~,1 I'iIle:liIll11g Kyu I~@e, ", }~r1r1if'!lr M. Lu n,d/'tt Ballaji RaRiI,al1i1U~~II1I. 1II.", 1 \'i'llu~ ;lind \'c.~k'lJhlr~tOIll,Hid,~ \'~rU:;j 'I VSV J, <l.f(; Notlonl Mnilllli$hil1l1:a,2~a Alkir«! Iw.a!i~k:i~· dl,~t(,1C:[(jd.by Ttl" ('. 91, ,;Vliu;11 C;ol.p~:D"jmlL.'l'lL1Im.), 1 {rgl:~~id,DO the ~l1doM'iTllC Dt),lipoliiL,ul'IlJl ... P~aSIllli3!(.ytold' ,dl!lildrii~iit:(eU~ l~pD(~ detecl '!;Iir\l,51~ iin Hu~ilcidiWiedl ndoliome.~ Illy m8'am oIhU-like ~gl!ms, plllD",i,licd (h;,D:NA <lind ssRNA G<lin r@ce-ptQ'F~ mRs). Y'e!'t. IJI[I( re'!lpo:nse!S to (:ertai III fiil'l~:le·stra ndi!d ffii,f\JA{S'~RNA) 'lo!1wllIsesC([ILn ill:n~y' aH.Ei'r Lri~ger r1.R~dcreUldmt cxrli.'C:;",h:n~ 1.1:1." '~y~)C.1 ~""I!a' 1l'1F,alfI!rE!(tio~. We pr~:ll?rIi~1~~d@n(11! ll@r@~hCllith@ ] 'reco9Jl1qij~!l (Iir :S\!I(h\l'inls.~.lbyTlR7 reql!!ir"'@s irncrlCfflf'lS. OFN:s) (7 .. ~•. M.I. MoreQ\'icr. 1I1~ 'lransp(lr[ IJIr (.yt!llS(l~]C:ll'ir.al [email protected]:atij~n ]1fll'Ei'rflili!'dliat:E!s ]!UO Ute lysosome by ~h@ proc@:;~ ,cf auto~nagll. lr.l·~ioici (,VV) imid iated ~: SV ((}:" II ,. .1.2, 1 I:n adldiith~li!'!,ll!! ~cpho.agywasfl) lUi!~ to b~ reql!ir~d for t~,~Pf()l(fuctio.r! '(ilf f!F!~~f~M()n"r~ pD(s. n€S~ ~ll1d tu::ai.~illa;ci~!,rated (7) or tOr1.lukJ.(;hyJea lr!/ ~eslillts sup/port iii key 'r~le 10lr~U'oophagyil:ll m~diiiitifi!~ sslRINA virlil:5d'~~.c:tioli!(Bodii'l'lerf~r(ii'l~[~ mixed {!J)innl1llC'ltL[I virlls Ciltl .induce I.FN-u $'ecfl~~iorl Il)i' pm:s. ~Ifld ~'l!lgge'5I!.l~alt (ytosDlk re'pljical'fiJ) i'lin~F.!!rmediat!~.(If li'k~st!s s;en.le a~ ~~:thIl{1:en fC',spom,;s compal'ubk: to ~u.~ir Hv·e comltcr~
s·ig 11'Qll.llr@5.
\rl ruses wi Illlolll IJdl1i£ direc! Iy ~il[Cel.ed <111,(1 achie\!Clhis tllI:mJl.gh lhellise Tnll-like I.'Cccpiors ~TLRs)iln th(; cm:losomcs l·fI. Vi,rn'l,
or
rl~llgl1!iled by nil'!:/',
of U~:v-~
[",hc~ nl1. !\C'v~ml d!~llllel ll"lh\\'1~YS. Tlu;: d;Is.'lic~~1\iimln;oL."'(}g'fii~ion P,t!I.!hWOlY.UII!>J~-d h'y l11o~l Gdl Iyp(-;:... iJii\'oh;~ ~ylfJplaslilic P\lil •.
oeetH'S inlde,~cndcully
or flls,ion 0'1"~.\I::f1!:~kalion.
bu!!~ t\~qlJ!in!;s. aU;;:",;hmenl .1II1d cu~do,C'y~oi.'Jis tern l:cco'gnilkm mo~c:c'l!le~ to dC'I£'cl vir-II [oi;ign<l.I.ur~s~.lH;h .. s doulbl,,,,~stmrldc.o RNA iii ilO\11I' ~!lgth.ll i ~Ii'lOlclion by Jij \ ~ vi n~)l. iSlll1l (ollRNA) ~l!'\~'W!rt.j; .a~!dS'-Iripho~ph..~(; roqllli~d {14~. l'h(;"'~1ob~,"J,'~ujml.';h,n ~ li:;d IJo RNA Il. _~). [11 co~~rn:n, hl: rlSJml~l~y~oid lh~ idea Llmll!l~ :rLR·:uawr~ij'llc:d n.:co~niuiml of ~ v~IiIJSC;!; in pDC:. f.lcCUDowil.\'!til,ll. direct ~nt'b::dcD1(1 iti~ct:Jm~ (poe);;) :lIJ:ieI.hOl!lghl to dlb'ucet ..
(.n
1398
VOL
31 5
SICII~NCI!
·www.sr.ien.-emag.org
REPORTS
~iml mn! lh<!il Illlc pwSCllce of Y~rnl :genomic ~n~c~~i~~~id.s;~"".~!hi I h~ Cndo.80!1iI"lr~y~![I)mmi'll 111 ~oHi!p,,:n.n~~n~s i:'O:sum~!'C!l~ ~n "~!!''''nt'Q "!l~i\'irr!! n ~tg'rln evem CO~lplod!(j! ...im~ re(l~i~!kln"
SV" bolill 1l9t ir~n~'M;::nZ>l (7), requires
;I
recog-
\'''~~ fC;,JL~o:r~I,;d1~h .. t ~f v ~1f.l;L]VI,1'I,~i ~;rn.!.i Q~l inkfl~l~dilll!.1,,~ \\llCT~~~II~ "'!i;:ti;)i:;:!~d in rOC:;, ]li:Jlihw,I}'~ (.f 0'1 l-I.~IQ~"'\,~\"~[,. [his 1)<Ir.ld~.gl~~. GO~S vsv !I]~dsv inrlj~lion. l.il)(:nfl,lsliM rI O[~X p]ni!~. \\II~y U V irfmtl~.. !iur~ r~mdh,::rs lo!l~wi.llg
~\!s'Cd w~thlmc green f!lll!rlirC~(C'1ll prot~ijn (GFPhwhi~h .1lII(l\\'cd us ~(ji "'ism!~i;,!.~ >l~W~ IIlh\lJi,;oMlm~ fOf!llll1!.iorl ~II rt."II! li~ncin !i",~ ~11!,~ p_?), Dl!Ilril~~ 1I~~oph~g(l~;I~.l!J fonlll1li.O'IJ.
L(_' J ~uld~rg()~~iI:tLi't.hi.uOIi fr!:..1II1~ihc!.':yin~plie . ~Qn'l1'!n u mombra!lc>;[!ssocialiC;'d fonn, which i~ sco~l,;'d ..:s l;ytorll'l~!'Il~C P~!lC'I;1 bj' n>Ull!-
P'\1r! ~c!J! i'~RW\ viruses, such as re~r·~"',!p!)! 'H~d c-nlu)' of viru~s m\15l preeede recogni.·· lar gJi"IC}'li~,1 ",inls (U!), ,~~dvsv U II. .illc~~p.l'I.M.1; 6rlll. To c.'w!i~im;: 1.ltiIi~ po~:;;ibi~il.y. \"'~ u~d a of indlJc~ri!!lIfN.dj S'Ce~limirJ'Ollib Ij;D[,~,.bi VSv :~'IUI,;;'Inl. wil~ ~I sil1gl!it,l!t~~~C! 1lt~id 'Ilic:>t; C.1~~es..Ui~liIilCcI1i1nl~i>ln u r [iL;'t\O.!:tllUlimi of dck~:ioll hi'l!;~hiiOiJ'i~lles I ~ iucl~c ili~Ir1111.M J i )oslINA vi~~',t; b'ypDD:,Il"e!:I\[ij,~IiIi~ unelear i~nld ptoldll (VS\IAr.,115 t ~ !'9). In wilild-llYl~e (\:\rr~
:'U:S!;l~J,I~.. I.hul iI SC P'~tiiH~~,lI1.\ i'b'il ~L)I b<c {'I<~rnl~ il£, to ens;ure i"C>!;;'Ol:ltt1 ilion '!,jf such vi ru,sc:-
a:~'
in I',I)C~, Tu C'.xl~lore'the i:mm«:cliliia r mOCI'l{lll i:"lu:: of )1s,RNA vi~<ll ,fj!;,,'C'o,gni~ion I}DCs. '~"Hl' ill ,,"i'I,;ml im.'(j the roqu~.rem~tH ofd~ ff1JI\l'!l1~:'~<l~~~:;' h~ \' iml in It'Cl ion. ~!lOOllt I'di'll ~(;I 1~1Csiuen L'l1W~ \\'1[11 :HSV h~;::Oig~:lili01il(,6. 11. I:?). r·oblUi'i~ i FN"n 1]1.\ocIuc~io.1'I by plllfil'iodrU('::;; I un in ruSpO'Vl~ 1:0 I,h~' s:;;RNA viruses. S:ClfRi<!u virus ,(SV),. OlrH;I VSV. IlrloXgl;.liF~.I!'~e ...iru:> ]!]rr;c~in]~ I(fig" I). Thus, U v~ imld i:lJ~I,_~ SV ;l nd VSV i:llduccd l'!Ylllpfllr.lll~\ldy Jow~r IC\ di.>o~ ~:PN~rJ p'mdllelio:1'I in! pDCs relative to ~i\lC: viruts~s. O~ig. I A I. Thi s d itfcreucc i.n If N·a levels C:Oli~,d nt'll be: aeeounicd for by inereased virion pmclm::~ioil with live viruses, because ~DC.;:railci:l w I)110ducc a. ~i8l1ilfic:'lm mmllx:r of' irions dll ~ Il.,g I~ hOlJI"" ofim:;!JIba~ion (llg, s: I). hi eeatrast W SV and VSV. ho!h UVa lrradieted and live iufluenza virus iX1L1ic]es induced ,oo'[email protected]~ h:~!C~;S If'N-a (Fig, lA).ID1 ad.oi!iO'n. !iCco,gni~ionof all nftlresc ,~;i;RNt\ "iruses dC~Cindcd cnti reb on Tlllt"l i1m pDC..; W~g- lEI). Thll'~, I~Dblll~1 !FN~(l. pmd~l(:~ ~ion in! pDCfi nWtcr s~imllll:lDio:u!\\!:iil.hVS\fnnld
or
~~hoo~ ~nd F4I(i!l~tyof l'I.II~dicill!il.flOtyQ. la~m- ~lI!llHo:m. O'.rn~llIt!l!di ft@5e~~n~ for '!)d~m:oo <I,~d T,ad'!llolcqy, l~p;ln '!~dl\n{e'<I.nd redhno'WglI' ~lI(;V. Kawag;udli. ]!Gpan,
ll)eiPar:limenj Olilm I1MJMbiofog;y, V,a~e lmw~fs!ll!i' S<:lllool ~ M.~ dkifll!. New H~lI'ell" ('1', USA. lOepO!r'liI~nt Gf Qlll Physio~~, 'ol(~ Mied~eal.0100 Dl!int<l Unllif.sily,-,Gradl'!ale
'·Th~ fPresenl
aOdI1e~'!C [t,eIP4,r~rnenl o! Iml'l1lunol'Gg~t Il.liml'ole<rs1'~ o~ Wa;hi ngtoru, .~@~Uil@, WI!., ITo Yllillonl (OI're~poll!d~lilre~h:OLlI.d be ilddrl'!~!ie& llti ! i'ffilS<!ki@1~'udu
ro.
I"OPll~,mioll I~'Y a<;sl!t<shlg OJ C,oHlbinmion of !11i:'lrp~·!Q.~ogy ,~ncl imn'llml[lll!.loT1l;!1cC'm::c I)"~.. ru Ilctiomd L\ii pr'O~cin \~~r{l'U bl>'i);;';'~)l.Ict~~~~o !~'.·.!1S,i'\UJtopl!,Il!QN(li'L11~~,~'l pUC's ,.~\tc(,'ju,uaild ~~'MI~l i!l h:i!!lh~r ~'L,w~h: di fl PN·t~ Myn!~!l,:'s,i~. I):;~l¢d by ~IHIIIl1C[;<fr ung' 011 e - .PDe A.-f" eells lndeed, VSV8M51 induced signi ficanlly dl splaY:lllg cywwHcP~HI.ct:m.'!' GfP ~MWln h ighcf how·c.il'. oflFN·n Ilmlll dijd \VI VS;V. at !(F'lg. 1). lIK'~C i!n<!Jy~c:> !'C~c,[l!~cdIhm ~tl~{)~u~ ~h~ ~.un~mll,dhp]i>ci~y,o:f i.M~~lio;~a,tMOn (F.i!!l." 'lJlh~~~n_~Ollill,,! rQm~ inn OCCiU~~ (:O!l~t;:l~l i\ cly in p[)C\. <lnd thaI ~nfm~tiO:ll\Iil11 VSV did Ilol. \ I nil] Ibe: (l1b~f!L:'~" or produ~li ..,~ virion n:: Sl,:l!~I. hl :l signi.lica.un ~l1cre:ls~ 11] th~ rCI'(:!C'HOQ~Y:r1th~si:>(~ig.. I), These rcsuhs SUll!:IOlI1cd age ('if p:I)('i> com:!ill]img 8.!liliIJP.~a~.g.Qsonnle.£; the hypol.hesis lhm vi ral l'CIJli,c2tlion n mermed latcs ge.i~crnl.ed a:ftcli" v irn I ~.·ii.t"iQ,nl 30d '(~3ble s I ). delivery of fl~c '!.tiroli ~lm:,1:e:ol)ro~cin particles We: next exantiacd the roCllLilX:rni.CI1I. 3Jutoph agy in hHUUC v j ra 1 1ie:!l:O!1l ith:m by to 'the' cylOso:1 Im:w~,diei] major s!~m.!illllS 170r I'D (',~, ·U;o!:h is ell [].wc' Mi:um ~mcd bone IFN-·n prodl!l.CI~(HJ by rDC~, 'LI1~Onl V'SV rmH'OO'W Ii:cMs wilh VS,\l inlhc I~"I!,>.'Cru;c inroc~ion, r~ W1li.~tmcle.~r~l{l>w c¥w~flli.~ vir:1I rcpH· diJlb'!;;~Hdo~~, of ph~m1<lc{DIQgj,c,d.i!lhihU~ol'.., of mJtOJlhagy. 3-f111Cthyl,l(lclli liae (3-M A) It2/~) C:JLt~OIli :inllcUlID!OO hues. rn ~ghl be: !)C(;ogillilZcd by TUl:. 7 in the ~ySOSQI]]iC of pDe!'>. A \\ cM·- mldWortUlal]uiil] (15)' A~.doses ofH\~A (110 '10 mM ~~lJld,\Morrmlil!m:L!] ij 10 to 100 !1M) ~hal known Ilalhwtt)' b~ whitll cylrnJOiic umncf'j<l~$ O!IliCho~~I~ to '~I¥."ci lc.dl:y unhih it ilutoph;lID' ~ 'If'!: i!1lfOd~~ed to the ]ysflsomc invnlves c jl.Iill.D(lpIHU~)' (111- To di,;h.:rmililcwh~~lllcr '~I!II'. 1(2'4. 151 IPN·a pr(ndutluon \!'I'm:ii lDnrnjjl,k'~c1.y <lIlbo[li~lH;dil'il rl,_~sponM; l('I VSV (Fi,g, ]i)" tnpl101!J.J! is Illh'ollvcd in [he mIHO'~IH1n Ol FLinthc.r. h~ purifi.b'l! pDC~, low,(;n' do::;.es,of cyc()sol Ie v iriI~rcpl icat DOl1lim';nltit'dia.le:;; to I.1:U::::;,Cjil~hi.biw:LS; Slm :r'Gs,tilhc(] jln :i(;~.ecl.ive the llysooomc Ihr i:'Ur7"ll.lcd~mcd dCtOC~~OJl of vimscs in j:1I[)Cs, we ~n.rsl ,e..xamjln~d 1111..': pfln~fLl ilJhilbit~on of resl)ml~s to VSV~ b~l. m:1I illnuclJ.:c3. without .a lliC>C~Bllg dl Slilrvival c rorm~lion of 3Uloplmgic vc:skks ill pDes 1(I]g.. 33 fLnd SlJp'P;Q11 ~n~ online m;fLtcria~te.xt). a ~lcr Vif.li,S s~inti lati,Q!il,. I.~ iSD.OI'ic.:llll}l'. 3,lU!0<,)~mgosom,~s h!lvC bC'l.."11 {!etcctcdhy elecl,:II'On Olle pmciltil<ll compHaHioll in the lIIS~ of Wi:mmannitl :iis, th~'l.l it '~l also ~l1hilbil cmlomrrCmSCQpr~1(EM). ~:owc\' cr. i~is dim,clIh to l d iSI i11,9 Ll ish oml qti fIIIli i~3tc Ime aLlro~)I13gic CYWisisl16). '~o rll~c OlQ~[llli::> possib:iJi;l,y. we examined the cxllent o,r infectlml 'by VSV. vacuoles!Mu ('lIller' SlliLICtllrcS by 11~nl'pho~owhIch depends on. 1:l1l,o.ncyl.iOO it' , ,~t \'3.J;yin,g :lilY 1lo:icly oosed ol~EM. ThC~CfON. W~ u~cd dOfl~;;; .1O"MA and '~Vo:l;1_m;lImin. lh;: lnlll'b~ lr •.mi:lgcllic mice c,"'P,I''C's:;,ing I,be- L(,j Ill'olcin
M j'!rm~in M('iil;kslt<~ilS.po.rt oil" mRNJ\, ,ffi~ril1,' Ih~ ~Hldcu:> 10 the cytosoll thl!)' preve I~I h1£. ~tllnS 11!lI.ion of host g.c I~'-"S.. ineludlil~~ I.FN-u. We reasened tlml. if fU!i.lol1 and virnl J".~l)II(,'iu ~)1'linhc$is were OJ preooquis.iw lor ...iml ~uc'~alilif.in. !he vs.v nu~t,lm 1!i'!,r;ki!1:g
l~IC
VSV.
pm.
1'(:'~~~!1CC' ~TIli.~~o_'>(;'ol)Yl23l. T~c LC.3·GFP ap~~i.on la!i(1 uo cmb::t o,m [he ~jolosneil~. 11lnctim't'> ~)DC~ (fig.. s.:n 1lh or obtain ~Mi~ll>l.icd,l':.;i~niliicarll qUHlhl.ilkaliQI'iI
'h'uni>g.L"i1C
of .mI0l)h3~solfil.cs. we um."tI mLlltfslloctrn:l. il~lliigin£ flow cy~ol1tlelry IMlfn. which <11lo\"..s dHl.raCH!I'iz,1Iiml or .~inglc eells wil.hin a
:s
~or or
or
B
..;L...
.1!.Ji:v4l
[!II UV4JS'atled
J~
i
i
'~5'OO 125~
-; ~OIlKl
700
.~SiC1
,~ ~oo
,
VSV ,
SV
O~--
DiUere-otijall'1e(ogllitiCl!I'I ~r siSlFtNA Vi~UI~S: by IPf)C~. (A t~ 'C}WT (Ilf {IE!} 'llR,~f-!iJIIouIS~ b~;HiI~ 1i11.lllrlFQI?I ~[lCs {Z 'K :!lJI~ W~J~' :s:ort.ed by nIJ:Oiie~(emi"e·3(tiv[l~ed (eU $In~ln:g (FA(S,)al'ld S'lii'iHdat.ed ,rQiU ]1,8 hours with ~:tA~'a!'ld or ~V"~f!a(li'l!.u~dl v'S\! (MOl = lOt S,V (2:,8; )( :l!.O~
Fiig.l.
om ~we
(h~(kei'l e,rnbr!f!) ,11~sJm:l1, Qr il"!l~h!l~u~!'IIMiRa(MO:II", 5); or K} V)V~S,:l or 'IIS'V (blJith ,al MQ'I = 10). IFN"(! ~~,~ts iI\I>e'!'i~ ,g~t,e(lIIld inl 'Il!ltl!r~ $ijp~rli!Q1tall'lt.o; ~iiii.ry,me'-lil'lbd imml!l:nn~orbel'lt assay. ,As:tefi~k, P <:: (105. !by Th~ dla~~ "IFE reiP!1~~M'ln,!jf@ .©Ir ~h!'E@ ~mi~r e~p~rimeJilts..
REPORTS
it{JII~ ;l11~he r.m.~,~ effective in St~rJH\~,Ii~,iIlJil, rFN~f~ prod'~'4;:!tl{l1l did no;~ 1I!,1~b"'l.cr~dO\';y!o",i;;; OJ' .~~IIID.~-,q!J!~ ~'1,.1)lj. ~IWO~\ ~d i!~ IIlil vsv i~!:r~~ um! i!lrI)C~ or~'1!!CI!1·1P.DC~. as clr;iJLl;Vll~~I~~ by lh~ VSV G",[jFr!,. .."pru_~i{!,!l ~H!:!:_3," Th,,~.,,~~~hs "II hCHl;tJI()pOaCllic cells l,~k;edlhe ATG genc cytl,l:~lne Sc(:fCIi9H (lfl(lldn(ll be ,>lOcoli!ll~(drn. (Ug, S4~). StIC~ln~'C h1l;g, no Q~;r'!.'im!~ ~~c1Qc~::l 'by thG: diIT«~n~<c ~.!'I.!hcmlc 01' i:vlrJ:;!~li:~n ~.fijg. ~n
~'l~I!.-",·i\i'''.!
or
f,li!"'9w1h ...
tlI~
rQi;m~~lihuh;d
~'h,;mainp~.i~l.iil;: ~"O!:l1p;;!1:n:~m,;n~l'l.,i!llcltu~~in!j:rOC:;,
in the :;pl~~"iIl. ly!~lPh t~O(~";s• md Imm~ m!lrn:nN • (.Fig, 4f\~, indicating that ATG5 ~~ not r:~quki."d for 'Ihl; dc"vc;kl:p~lL;~ll (I r pEl. 1>, ill
"'~\'fj.
a.ffi(,"CtilliJg v im I
~11lry
To
~N,~DIl,iDllC'
.1L", 1Giph·
4!D) or ~.~nliv,,]of pOC's U]~, S(~" ~n~lr)Jl~"'t'l.!.. US.V~lor Cl](i st~m~I,I;.ti~1!11,o:f ATGr-pDC!l 1C.~1iJ,~.i,{,,J in sinnih:n M.'Crd.ion of IL-~2 un.d decreased Jc\·ds of I fN -a !I~~g, ). Th<C'SC' 55 oaW i!ndic~INXa th.~~, .!;lhhCl!Llg,lf! iUIl,~l~ reeognlilunn or ~-ISV·~ ~~~X! 'iG ~,e~ti~ili~ ~lihlcl q
inH'aibit!:dVSV ~-cogni[~Oti i~ pDCs, To d:t~~1Jii!hdl'de~lto.III;!OIli'lIi!c Lh1!ro~c of 3,Ulfil)hiig~ in v im~ re(10~nil~'!.:)Il. 'we liIi:>crI ~
gcn.oClic~P'p'rruldl
3~
ill iiljIlII~e vli.rn~. ~:I."CogniliOl~, IDC~, plllril1(:t1 p from tlhc' l\'f\GS'+/+ er l\TG5~ chim~.i'ii~ were
ill fectedw
Ily cxmnihlint
,,,,,Des; iwllU!OO
in A,["G5. a gchle es:sC'Iltj,;11 fortllll~ol~h,,£os(line rorm,lIjo'hl pn Ilccam,c ATGS-'- mice a re incapable of SUI,\' ival l\I~j'tmd ~ d,~>,u]" We ~171. mature ,\TGs.-;L rDC's W~_!'I;; oiYwijJ!II;'d frorn ICllwlly ~11I,ldi;I~~'d \,VT mie~r("Cglli.~liwllL.:-d 'Wilit ~i~,i()j" <J.l'ld sl:lh..'~J~ cells Jl:om ~h~ II.T(i5-+- uecaates. Upon fuH ruc:omwill!ui.on \''''ith ATG5+ c~~ns (ji,g. S4M.
seereti;on o:f ~~oth IFN·o: ~ndiil1llerle!:l ~ iI1~~ 2 U:L-12 ~was dimi nf s hed ~ lht:~l~,~I1CC n of ATG$ {.fig. 4, 13, and C'J. The di~nh~bhcd
'~hi:: 1:r-~·f.I.pi"od!J.~l~.ml '111~~bscnet:;o:l:' ATG5. Alilhoij~h f&~.c mi.'eJlan~smb:, unknown, the seteetl \Ie dcrcif:~~n U~\I' b~l f1:01 Il-tl. -a, ~rocl~<CI.ion by the ATGS-'- 1).1)('$ mi£hl, ~:l:~oct dC'rC1C~h. e $011 ins of I be TU~ dowa~Iream j1.lgnal]ns cOflilpOnl!lll'> to lh~ !ysojmtlrn~I!L:r j,:;l aJil.cred ~Ill
:>'Om~~. FUl1lhcr.
~o i.n'i!\l;.l:;1lg,,~~CI~!~ 'r(ll,C' or ~ll~t~II1l;J!<!~yin 'lnti \,ijm~ :re·,'ip~}n.~Fi.'S I pDles ~Il bJ f \I~ \"t),. Al'G:lI+ • Oll]d A'fGs--l- dtinnl(.'lJ'ic mUIJ::'~ were 1nFC'C!iI..'d W~lh VSV. Sysll.'rnic lnfceuon w~llh VS V i:-; kll(l\v'I'! lO Wrgcl ~d~!liC' pDC~
'!IHd
in ~h~
~rum ;:a,lar;DUl1d 12 i!t!~~rs Uf~. Sil~c(; It-Ii and ulhcr it! j] ::I:!rm:rli:unry cylok h1Jl"$arc scereied by IJ.o.ll-POCS :ill re"'PQI1Sl~' to VSV illrccl~olj. wemeasercd SIC:rUI'l1l :IFN-o lewis. lnfcetien of l\IGS-J- dll lmeric I'll lee resulted hl s~gnJil'i~ CIInl~y rodMCC'd IFN-o: responses comlXlooci w~lh the Al'G5"1~ chuncras (Fig. <IE).. ('01!ecl.i'!;'dy. OL!f d31:~in,,:iicatcd 11m! illiIIOph:!liW eceurs.con sl.inH ivcJyinpDCi'1 .mdi~ Pl.,\! ui roOd ['o:r 1X.'C1O:gll:il~m~of VSV, but mol 1151\1-1. i~~-
['c;clilJ.ll ill 1i)I)CS. III a(](lli~ioll.OiUlopllt"lg.y is I'CGlllircdl ror I)D 'producljon of type: [ !FN:l1 in \'i I m[lndin vi \10. AlIIIIOph!IJ;Y i:.; :JI'1l ~m,cicnl, pmhway lor hoIiIlt'll\!'1~a.l.ir:[tlnmO\',~r of ~Oll!il.~ \.'(,'1:1 ~ I]U';:lC;CWIU~fir. i
and f(Jt:ll'LlIrkm ;H.cClUI~s~lir.!'!i duriUJ.!:'t slarvtJ:IIIOlt Recent s,I.~t1h;.\.lldic3~C(1 the hnpori nm\l!.:"!'c tt illiJ~(}ph3goSQl1nes ::lIS 3111 flll.limicmbial o
cun1lpo:tiC'I'ill. ..
and dcstliO)'
~·'_II
I· -
$(a1le bars, 10 !JIm. !;acliil lVIO'l'i:s,01I ~(lm,pl)s,it~ ,~r arreded b;y these il1_Mbit~r~ ait!:l!ilJ" dOSie'S e.~.· ill nll nedl" whre.3i5 IIFN·~ lpmr:ll!l QllJl'I \Y,ili-5 in ~ibi~I1e lC!inefE~ti!lll~.(r'Q~ralphiS of lihe sa'iilII~ (;!lllb, Th~s,e
'e!'xileriim~mts h:i!!ie' beem u:ijH!tltedi lhree 'liilil1'~
if'iig_ ,2:. lr.o:rmatnQII1I or alJllopm<llgo'jol'lfles in p,D(s, P'D(A·I~(]nlc+' (e~15 ohtalirned hom LO-GFF traI15~~rlk rnlo~ w@re ~Ji!.aitli~clri(lr lh~ presCf'!tE IIliI pUlm::Iil;Ie- G;Fpl" o'lLJto~h1l905Q1iIiI~' by ,Mlillf'C ~ ~,g!;!lf:S'aJl't~r if!c~b,pt!gf! \1!'ilh "'IS,'ll !1i't l~~ ifldk~t~ c MO~SliH m:ediiil comtretThe ILlpper fouir cells ]1:1 the media e:o(I:I'~~O r,eprest!lIlIt (yt.o!5olic GFptt' l ILelils, \lIhereas the rest i!)lff the ,C!eLIs r.epre'selii t pD{s ~'l'itlh GflPpumta. llhl! eelumns re'pr€':l.Emt i:!,rigmlIOJietd (BF)imaginl!J ~gray)r and PDCA·,l (o,r ,;1111 ge}, 1~(J·(3 FP (,g li'eel1~, DIRAQ 5 InliJc:te~r {Niue. !',ed), and GIFP/nud~.:Jlr {eveday) sta~n5"
I,~ ~:g
I
-'I--:::"-±:-:~---j
c
=~E~
a."",'''''-d.~~d
G'f,p____"
Inhibit:(lf$ ot a.yt~phglgy bt()ck VS;V Ir,!!JDrgnirtilon. IE!O\ne manaw (:ells WlHes!l~liiilli!a~d with -V5IHifGF~ {MOl", l[l} lomE! presel1.(Je I)ff~hE! ]rldiic.1! ted (fO:5I!?'S ,~f h)hilbit~ws of a.uoophaOif. WorrtmiHIn rJin, .i3lnd 3-11M, ,iHlOO{AI lnf,ed.l0IrJ (GF:P)
IFigl.~.
w@.~ .1!ss~@d. Nc·rth~r pi[iCs; (tupper Ing'hl l'ILI;ldr'II~t)nor ]l'IIri:!~tio'n 0'[ iIloni"p~ill(s (LO'Illt'r rig !It. ''il:l'adr<llnU \lla:5
cyiosolie llaillog.c:lls .. Certain vinl!.'>infecl.ions. sach as USV~ I. are known to tri:y!~cr~lJt(J)phflg)/ (2.\')1 lor the pllrJl'!'O'SC'S vi m I d.cg_IiHI:m.lion (_'9~" whereas !polIovirus, and r:hinovin~s usurp .m I<Jphi1lgosomal mach h1CI)' lor\! irn I s}\nIhC'S~~ pm,oL!l: st~Jdyrc'!,i<Ca~s liDe h~.lpOitancc ()f a.",wphagy in: ineate I«QYlliHon of "fro:! P'~lth!Or,l-'.'H~ a!UIIFN"f'~pmdlL'Clio!l ~!l~lDC's. The ~.!!ld~n!ll'iof~ll~ !·«..1l0~ bU!!lh!il!il:hlthe diff{;.~~u:!iul m(;-qtl]ll,1mC~'I. 1\;~;o!:\nlit]u!:1.n<1' divc'rl'ili.;· for cl,~~ (l,r 'IIir\il:;,(~ in pDe:>" A 'broad i>p(1C~rUnl uf vi11.I~(;S a uiJd ~hc ir ~ IlJect·i:CIIli cycles, ij-'OIH the: "~ fnOfl"'f~~"{)Cilflted g~1 a(Hlfa ic uuclcic ae ids in the endosomcs to the ~plic,adQrl (I r ,·lrn~ gC~lomcs in. the cy~Cl,sol. ill!:: detected by pD('s.Fill1illilly.
or
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bt'lt ccrloiM CIWOD~ic v;iiral ialf(;"!Ciions, m~ghl. bcue fit nl.'ll11n in~COrrQr<il,I:ill'lg :IgC1ilb toprom.tDtc ruutorlmg~j j~'ol" bOlh \'~r:Ji1 ,ek-:Srn,(t[lti;ol1 :md in Il.ate n:cogn h ion.
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REPORTS
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iII'!·anQiI¥ (8M) ~f ~he r@(.omS!lH~t€d (hi:f1iJ~rk Imk~~¥a,~!):ss.e:s&@dby Illo~v (ylOiillHflirY" S:e(iretron 'IlW (B) IFIN"fJ.and {C) Il-!l2 ~rQITlI 2S,,000 ~ACS:'50:r~ed I!lIDCs ~rom the im:Hc:ated ehlrnera \ll,[li5 measured fo~lm~ln:g "iISV In'r~H:liOnil {IMil.1I l!O}. f;adl dol reprll'~rtl:'; ~n 1rld1~vidlU'~I, m.QIUSi~, (11),) Tile ~M.e,rIlo,{ vir (!~ lnr~(,th)lfli!l Ulit Iwn~ liIl,anm\l' (~Ih ,oiltal'm,Nli IIr,emthe re~p'octive (himen'ic IiJIl~1!i\la!l: .as~~ed aUel i infec~ior1l with VW-(l/GFP' ,ilIt. MOl ~ 5. {E) Seroli1i! I FIN~-Ci:ie~pO!ii'Sf!S\lIere ilTieastilred i ffiO'm AlGS, d~i,merk m~(~' ;ah~~ ~r!'redf:(!111with 5 x l~ilaqli!e4i!l:rm,~n9 li!f!ltS (If p VS;Vil1ltra~n:o:tJ!~~.y alt. tlrae lindicailBti tilil1le pO]l1ils. Ea(~ Id,ot r~pr~ents: ,aliliil'ld'iviclllil'l mOl!ls€.Astl'r[~k" P' < (11.05. Th~ data a~€ repr~S~fltiltive (li~:'Ji,ve [{In and (0]1 or Ulr,~@[(At W;l" and (IE)] s'!milCllr 'il':qJ@ri!'!'l~!lits.
F'iig_ ~,.A~t,opl'!lagr-iS r~qlUlir~d r~r \fS'lJ i'\e~(lgll'li't'u(i[!. I~il,l~r. lil,d 'Sp'~ee~ c~lhhlm a A iPG S..,r- x ATGS 01< - pups ~\I@~'il' Uisl~dl t~, r'€wnstillJll'@' ['~hgHy ]trc! c'iiiilt:ed (57 B L6 !II1IiN!',(A) ]i1:e p,re:)effii'C)e Il'f pUC'!; 'in HH~~p~,en (511), lymph Iilod,e~ (liN), and OOI'!1E!
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&~ !11M. 19:8,513 '(20:011. 7,. 's, $, 100~b~~"T, I~i:~~ 'iL H~l'I1.m~ s. Abira, (, Il~~ ~ :s:aus:l, .5({,~f! :~O!, 1529 I"mo,t~. Ill, .L M, lL.uoo et Qi., Pf«..NaIJ.. A('Il'.d. Sr;i., ,U.S.A. a.O:li, 'Si~98
9.. ~"Hell lilt 01.•. SCiI!'M~ ~.o], '1!S26, (20011,~" lO. I!'.;. Yil.WdijjN 01" J. ImmElrlllL ji"14,, ~,i29 (2:j)6&) .. 1,1., A. titug ill' ~.W~ lDl:JI.3:, l<!iH '(21l1!J4). 12'. lit. l. ~kI'<liilM,. G" v. A.lm. 5ml\l'tl\. 1. tm.nrunm. 49; ~91.
19. II). F. £I"jdl il\t uil, (mi!(111 C'f,J'j' 4" ~,~ (OOO~). 20.. (, ~~n t;~. oer W., A'I'Q1, Cdl 6., :ll~3 tz@OOl, 2~lk 1M.Mulll!hlrn.:l, Y. (J1il!i!im~ f. ~Ojli.iroo~ rtf/I Slt.~(t
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U997). 26. lIt Ar~!Iii" M. T. JOOoR!IOI!, .1. A. Sw~m. J. ,Cdl g,w, 135, U4? (!!.'~!l6)_ 21. 1\ Kwrn~ ,i!'/ of" HDrrJl"~ .Ij,J:<l, 1.03;1 '(~OO!li~. 2!. Z. blLllIay f! I:l., ,~ ~tL ~rlr.!. Sa. SA '1!I.:l.!l'O
NIIH.j.MJl. 'I'II!~ Sl1PPDllIed I:lII 'tll~ training g~1lI1 PlIO:;r,iJfl!i rIm NUt, IU:.t was ~Ilwnd ~:fll!~ ~jnn.1J!yd S;dfll(1l ,,!HIl~~'~' !If KliIeO!.;II..!. ~n !n... iig~t~!n ~ pa~hO¥!,rnMi!; ~r [f1~~liDi!l5 DiSI!~Si!' ,,q,'fIa~ hllm Iii!!· 84Jfro!J~~U WeII.(lGin~' FlIl1dl.
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REPORTS
S:tru cllur,a I I:,nsi,ght. inlo the Tr'B,ns,gly,cosylu,tionS'tap ,oil Baeterial Cle'II~W;a B io:synt:hesis; II
PetpdcloglYGli!:J1 ,glyoos:yL~,~i'!sr~ ~a~s. {GTs} (aI:a1y~~the polym:eri;:~tio:n soop I'Jf celllbwall biosynlihesis., ar!! a ild alrl~ hig;hiy ooflselVloo a~mS$ aU b~~I.e,ri~. l(Jl'IgJ ron s.id~iied th~ ~h(J~y '!J1raU" (If aflrt,'bi~~k researth, ~h€y irepre5-eli!.t all e~en~~allil,nd ,~siiy a((e~ible ,d,r"'LgMrg;elt ~or an~lhio%k-~es.i5tant baICtflria., ll!i'u:ludin:g m!t'llhldl~ilil-,eflist"li1It .'Staphylococcus tJ'UteiJ£. 'We oolve li1ftemllinedl t~e 2.8, an!Jj5UOIlI1 slJlllcbil re af a hlihmctlortal ,reU-'l:IaUl(Irc)51s.-Lnln~]ng ,e-n;ryI1fl:E!',induding lts tra n511eptiiciase a lui GT domi:l~liIs, both IlJI1Ili'gand ed il nd mrll:l'lie.x.ec:l \lIiitJ'h ·the sillootrate ollill.og moeli1l,ol'l1~dn" lIne pelpHdogl!f\"il n G'fs illdo,pl a [:old di~tilill;t; rlrtmT~'~no~~ of ather GT das.5'es." Ihe S;lrllJot~J giwinsightil1ltl Nl"5 clfiitik~l rllilture~ 01 the 'C;lI~ ~ytic mecnl'lllri1l'FTI a1flid keyil1l~ro1l(tiCim. reqll!lrMror illllzyrrm itIhffibithm.
iIli!~iiIi!br~l!ile·b~·'ll~d, IlCb;!~~~rU'.d.'~i~"'\'<1!II .. i;;" bui~l .~.~,gll}'OO~~~ ... nmn~ ',-,,-a;';L; (I TI .U'!d 1r..11i,"P!.'llludm;~riff} I.:I~ylJ1~"l\. \'\Ill id'l l!ll!!,Clh~~rpltldut;1i,l croli~~ .. lin'kc't! pc"Pl[d~gl)'(llnrncsh ~1~1 gi\ es IhL;m-ICIl.'lfiuJ:!:l hs SBmiglh. and shripe. CtIZ)m{$:ln: e..xce~t{~il. drujil tl'ltgc.'l'> brealilsCl [hey 3It! ,a;..~entill~. 9(;" ~re
liJ,i) mo!Cn~mycin
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fJrWlflfmuff!(!r!~r I~ (71 [2gp~ ""''q~Il,;ntc id~~lli!y ~or 'til~~2.9~·io.(i, r.)n;:~oc)l1.. wid~ ... 'rnL!il 7 ~i1t\mo ~*Iru1; (~'\I;~uijOJl (RMSD) of :W A lor
OJ Ei ~hr.;r.m~t IlfJt both. 'Ill' tlte Gl s/[P tat;;llY!icclo~n~.i~~ ;II'C ('I1,Ip«imprnS:;IMc \1"]!~'fI 'I?l'lm~Irillg DJie hm C~)%lil~ ~{)nTii!>" .uradrC,:;Ilirig dml
J.
version or fi!3P2. uhe Or.ll~r lJiullIIicliomd en~m(' idCiiu.itl~ in Ihe p;alh~CiiI Stdl:Jlly!m_'()fT.1.t, mm:'u;~i. ccss1blC' rrorl'~ '~inl!perirJOllim. 3n::~ ha\11I! 110 eqllh'l)~ ill~<iilPocnzyme SU'UCIUn!' {2.9 A reseluden], 3nd a COIl~I)~;';. of Ihe 1!1K{.)'1:1lC \!ililh lllC' alIlihioti~ Il!m nil nmmml'liiall, c:dlrll!;"TP 1.'11Z}'l1'CS [also kll.Q\"'~1 nlOCno:m~Gil1 i2.R A)i. "[he gW\vlh (!If both c1:y5(l{li as IlCnicilliu~l~llndill,g protcil1~ {PBP$H:lre!he'~rgc~ roml$.lCq!iil~a:OO~hc dC~~~~l! bul)' Icl!inle~h!i'I:I1T1!i~lC" or~l~ ~bcr.."!n1 .glOUp or <!!ltiooClcriflL"" ·to \1!.rhiChfl dij!\~lIrn~I~.}'~!igh ,d~gu~ or It');G!ifu,'1[t!ochas d~\~~-la o;.;icic (lit Bolh s!nl:(:um:.'i" Im!,'~ Ii!v(lfnblc ~CI\.,'{11~ oped ovor tllIe I},1jSl '\,\~O l decades of clinical use, The dlCm[cshy (tab!c SI). w.uth R",~111~ v:.dl!C'S of It).1 .-4,6:% [or 1I1:C'O!poolZ}inae {!JiK~ 23.1/2!':,.0% pcpttdoglycan Gf pl.'OI..cin fuJllti~~j. GJ~I (W\,.",I;I,
c.ril)I.org}.has !JO~[ruC'tu.r;~rtfI!t~l:ui\'Qoi H) d,~te ;:lmo i... ~u...(ni\lc (,OlIIdilblb: I'hr Ih~ stnICl\Dre-l~l~d dnl.g ~i~ll of new :mlib.."ICkrithl! COIMpcundis. GT~~atld TI? (;IRyrl1'L"'S 1l'l1UJI'be r~nt as n1llm]oliU'iCliomd of bi Ill!nt.tiomnl pr.nrei;ll~,~.ll.\·,.,il]~ ~bc more pr,evll~cnt bifLJI1~I:ifm3~ 'Cllfl.yil1t'CSpo&~" ingan N.;tcUiininml {jT.'l~ oolllrl;ill anda e~r-c:tmiiml TP dm:rmiu. SCjh'tmlt.'tI by a sm::tll Hl1k~r region, The G'f~'loomains arc :dw'l)'s 11JC'U.lbretJ1Houn.d .. am;] they po.~yn\\''lrizc a lil,id n pell.lapliPlid:c slIIl15tmJre imp ilt~lil1kl>d l,,"acetylml1:r;m~ic m::kL
n~~J.!
Hfl';JIt l1~m'lan~ b<1~I(ri.~ ir~r'.:1l;:litm.~. . ~.ICl!·L;. WIi,l.~pi,1nt two c[)'1;I1:,l11 SEI'iI,I(:tl!l.nL'ioi o.i· a soluIJrc tru Hil'~\r.:d
IlleR: it. ~OJ'Ilt: .Ila: ibii,lilY ~ii1Ou nirIIilE~in'Dk!er I:'e.giml. The G1'jll doit'!Iin has h;,,'Oputtd.~VCOIllI~.y~j~ g;liillilJtl lc ~id i\.~~li;c. .. 1EI14mlicl E'I"11 (~)1•.Rlild Iheir ittnI10t1.1t1l1!C has been dCIIl'LO.Illf;I.m.lc.d in. nUlimtiollmJ ~1mlks on the E.~,'hol;!rldl#l [~~ti PBP ~b CII~ll1ile {9" The {!I'l,',erJH:fold (i rille lillid U~def1Cl1dent GT:H d'om(ll~~l show)l nosimiT.1fily h10 cilltCf '~I!~ClT·A 01' G T·B (bids dml h!ll"'C kIf;J1L"1~CI\'~d in ,III i)!1;,'t~in!il GT 'L;'~)lIli!() d~~s. :;,uucwr~~, The lI!:m~lUlr~ i",p.r.~~~.lUII'·U].y (..(.hcli,c;lI and is comj.'lQsoo of h!'!O scg~!lenl\1: tim: glolll~"'tr "head" rcgirll] ~illl~Hl,,;ti !ll.!"l W duo: lil:~k(l[ (kl!1mill "ll'l!il :;.H. ~HUlJ~L;r')<i~\{'~8l~Onl ~.,_ n~ltin tl~i~dOli~r w [he Jl!lL.'l]nIm~i11ll lFig. .~),The
or
gJohu~~r
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If~ri[y 10 bfLClC.iQpl~ag(! .~~ysoz.')Il'I'ne qlL) IDAILI Uf~) gives a Z score of 3.5.12% SflqI.ICflC(! id:e~l~ily fori ~'leQ$,1ll-26(l re~ ion. with an, [{MS[) 4.2 A for Cnl. Given IIUUlhc l'l.\'('1 l'flzynl~s :le! rill :i!,i~ndbti~loolf.a1Cf,:.,i1 i:-; rn,"siblc' i nll<!l~h~ GT~! modu~,(;' wW' a~-qllinxll by ~hch<lC'~
il IlliiS
or
,ill
N~OCl.y:l,gII1l~ooalnillePO~J''11f1ern I(Nt\M-N lliG hi, The IP domalm, are soli\''Cnl.c,~poscd ami calaI).;m, Ihi: cro:>.,,"link:ill,g 0:1:Ihe peptIde s\ibsl.ilu~ns on !:'he:;' acetyl: f,;ml1llS of Nil. M.lnhib~bJrs (If nhc (JTSII~'l:H~l i{Ul ~n:;J.}" be Ic:ftS s~llhjt.'1;:lli(l' t~IC ,!cvc~~ (!~mcnt of li!;,'i.'ii~I'U!Q~ !h,hUnTP'inl hibi~UK. whic'h elm be n~lldL;:r~-d ind'Hb::li:w by,~I·t1,."r'llinll uf nI:,,~ p'l.;·~Jai.dt' :;,ub~tih.l:Ciln, -no'~~Lbly vOlalCornyciu~ • . irl nx;"i:o;tf.ll1lt slmilil:' (2. J), There Ii>ORI1.y om~ '71'CH~ c:h<lfa'Ct(jj1 o;:diJ Itn ibulor 01:' the OT ~I re.:1cti(m.~hc Slrqnom;n\f:;';o; flal.urn I p;I'OOlllCI.lllOCIl()myc in. whi:,h is not cfliC'('tivc h~hU1!!llilllS be'>llllsc Or[~oor l'l.iJso.rpl.iO:I.1 prup!:'1'i~ei> {4), DQipJt(l iis. ~xltCi.l.fii\i'~ !lise as II grow~lprol1motcr. <in ~tliI'I1.. 1 Feed, ill) pkliSunid.bome resisi:Jllcc ~.('I 1'I11;~Il.omycill I~.f"" tleera ootccleel so raJ" ,~5 UindCII'S!tJndil~g )" of Ihe
D~li1llilil!,mt o~ gi~hil!ilIlis!ly ~ml MAIi.e,ul;!r 8f~, aoo
FDr Ihe' nK~~DO'!l~lCill"bol!lld limn. l1iu:PB'[P'2 \'i.·~flrc~urem\~!.'a]!l· <B biloh:IT ff,:Jd (d i:I]]l.illsi:ons 55 by 60 by II {I A~. w~111, fiTs I the :lnd liP drllllain.c'i s(pm:rmlLidi by .11 shm1. ~-dClh !iJ'Ikl'l' regiol'l (Pig. 'l.l ll1Cl~lW I1I(,DI!Ised. ~runino aeids 59 Il) J 16~ OJnhJ~dthc N·~crttllln,lI 'h'll:l1.~m1'!:mhlllltc .scr'l~el1ce and some oi' Ihe ].Q\\'~ (lo:mp1C:ll:ily C~I.CIi'Ib1Ii:n~Dfllil. 'Ekxtroll di;.'iJ'II.'>ily is, viSible foJ' I'csid'lIcs 67 to CfJ::!. cx.cq.~t I~r a ril:w s.hOrl.Io-ops in t~le Gfr ~I Llomain tli:1 7 tQ I 31) Hlld 10150to ~63 in inK!' J,poeI1ZYI11!! "m113 7 H), ~44 in
teriophase al .fI[)O.il1t lang 3£0 ill :lIS cvolutiau, ~)i~c low sequence idcl1lj!y.. dtcrc is, good o"'cl~ap of I~C :SI!COD1(_~1J)I ;iJJlIICll]1lC b'11l\\'~n the
'b~ [ljrnt pllw~i\ic ,gJIUltllnm,1,C GT S<I ,QHIll,liifl m'\.~rb}'sIlmi. or 1L ~E114 \V~~hEI9, It:5.;pcclivt;1!}';fi~, 2.1\). The: hdk'JIl ~(.roloID' is pc;m'!l)d l(ICI.\VCCIl I.be I}'So<::yll.le and tiT ~~ rol~ds.wit~ lhe· s.oltirbr.e Il3-rich
1\\r(iI C!lzymCli,. "lIld
ttrl'<!l.l)'li(;r~itjllcCl:rlhc
1l00l'Win I o:f AL repln&..'ti b,' tI~,.; r1'OClin:bmH(;' assoeiaicd (rr:!!i~ jaw sllbdomain trcsidues 1:2~!O 1, 82). The jaw :is coml'oolo)d u rlhl.w 0: hcl~()C's. wit~l~hl! 01ltmno.s1 hdii'l. cOl~~aining tl~e ~'CCO:llld ptlmtl:'vct&raIYLicl\::s,ldne. E 17 i.. pock!el I0J11Kd A
IPi !!I., I. o.reril l1shrd:lilfe of ~BP2,.n,;~ nlodular i1~lij,ge !llf lTI~ GT !i1!. ;jj!lid mlP Ifotrl!5, ts<hO'>\ITI~ ~'IIith itI'Ie p~~d~(t.eld ~p~siti~1'! of UI@'tfo'[email protected],1Hl!ll ~1E!§I1Mrepreselll~E!'!d by a vl"ftiGil illlut! ~~ta!l!i!~.
lIIhea"tjye..~ille re~~Mlils
,nn
ri®l~he
'"!head!;
l~gjOfil
sho.wn (I) r'~ Sp:h~F@), whe GT~~l,dQrmllilllis pro· IPosed ~~, b~ at Ileast lP~rtlyS!ulJmE!'rged ln~ne bila~rrilll m:eerro .a;o::e.!)S
the Up~g Irl~!JI!l!SitfC!rt~. whidli~ '(Olll~tlililt Wilh
IMembrane
'"'Jaw· lD"eglon
235>0 H~L~1fI Sdeooe~ M31l, wnc~r", C<lood~. "'1'0 \~hom oo~~e~p(!lfnl1k:n~ei!llIO~_dIbe add,re5.'Sie(il.IE·,rnait
1~~l:[email protected]!!Cllc!m,IJil!Jt:;"Q
oibservalions,
that ~at:aLli'tl~ ;!c!1l!1iily ill nd (ry:S<t:aUizationJ were depen.dE!'n·~ (lIn 'dii e ~res:el1ice ,of det~~£I'~nl$.
14.02
VOL
31 5
SICII~NCI!
·www.sr.ien.-emag.org
REPORTS
b)' residues
GI~~'lOC CQJls~r""m{ln
cl,oo~ to t~l~ ,1ignikocil" rt:!gi{lu~ shows se~n othl;!'" (;T:;~ C1117}'I!~ll- .m'\! ~!l~y be :~!~]POrl!,u~~.rilrCO!~':lcli,!~ t~~c ~!ilO (lfmh~ ~rnlb"'11IDlnbl11ll'; :\~~n\.'ll~+ A .:;:h;:I:\ b:t\'.,1.~'f~ 'hh~
(KtH);(fJSbG nwtill.s, 0),· lllC TP cm:1lyllc S)9R residue I~ -1tl A flom ~lCf1'lliln.a~...e ~;;tt;!!y!ic t f,~io~~c 14 of th~ (iT:;~~om~in. lh~ 'r.T~I !l'
Inl'dl
~)q,":'il;~:;~Jl fivl;_~ ~gallllh[Ll:;i
~(;'M"'ii~CI\.>a.o;;
mflot~r II
~()1l11li,';the imllCnll;Q~I,
heli1'i:
Qf~hc j;ll\'\f
)1l~hdQfll"1n t~~<iitdividcl',
~w l\\'O' p(l(:k-
globuhr :ludj!l\,{ ~~lx!o]l]ililj."~"i]llll1cd:;;nbiip«kC'lwilih ;'1 larglilf ptl(;kct en '~hl;' opposite ~,i:dc ormhco ~lpk\C~d.G. The ,e:lcll s.~lpL;fi~:npo~~,\'i with d~¢ ~ub~Irnk binding ~i~~~ ~llld n;b~(,\d ~.)!Sl]i.!:jlmile i~ II fold::>lF~,. 2 A), n~ 30Ii\,e :oilc or the TP Iclo.i'l'lr'i.in(ioli:!,[ldifh. l:he eheraeteristie SXXK. (Sf~r~,XfNl(_'l\'lIi1lL'i
ih":~llOllOrliirM:lim~al (I) {Iig. S~). Motil!s I ;Ind III COil· ~ain du;:GTs~ ~'!~~bl!:(: l~lUi.;JIIHii; .. cid~ ,(131 <!Ild £ ~4 En I, R:.'>p<;cIQvlity" \'Vi~hnillCif Co,~n!llt~ - ~ .A 4
G'f~dl'~r~7.:}'!l1ICiI.
inbo~~:1
1,;1:s.md ~.!ii p!iDbllh~y QJ!\'olvoo in :oi\I~lmt~ro~i. t!;tiOU1_ ('br~~~lI"l; ~H!rmS~ by t!rnl;, ~OC;I~ionof !~L .. t\~iJL:rmil~all\.';Sid~it·:> M in 7), 'dill,' hy(lfO;ll!hobio•. ,;;t~d ~hc h}'!!:fu:inphil~.'~ n.,,'ttuil; gf !he TP ,IO:lllll~l'Iti.';Id ~o a modd .!'br m~1'1!!bnni~oti~I'Ila~(I]'i iFig, t '"
r...
m~llIiltl1y~ii1~OO[Lnd.
i>inll.~tWlre (.fig.
31
~or311 ~mlips cO!hcr tllOlH d'LCC~ lipkllillut tFit. JC,.. !\'h:~cill!'}~l,lycinbil~ds in the cl:cl'lbc~.wccn Ihc head and jMiM :~~im~:s of Ihe (JT,~I fold in an.
Cl::~cndi)d cOllfom1illti:cUl [Il!ila~rolxlb~)p mimics !hal (If Ih~ gmwinkl ~~IW~N:imin MJb!i~II1.I~~ {Ii:" I~R1!!bllCl. R.illg~ BI(1i F {.Fi~. JB1 ~'nod~'l~ in 11 ch,m,ir on:nlhl1t~1Lli~.HlI. fO!llll .. l",~·i~~!..ldp]~Ul(~, wilh id~L\.\ ril~g B :rO(:;u~ld. ro~"I,~\dihi<! ~illlk~rrc,gioll . •mdring D 11WjLX:li!]1l nll~ into ~H; bulk W~\"l\l1'" C[lI!c"k.iOh;'Un will~ 'lh:i}; :1'D~I~]. mrJe'I1Q!'I'ljlcin derivmiw!, .~ftekjng, ri:I'Ig D 11C~::li:n rnldbimi<: ~Icli\'ily ~ It lJ I, Rimg A c...'(i.,~ Ihe ch,:lI:lncl 01] the sid~ .fmmlteSI. fi-ounmitc m~~nbrnnc: 8ndi'll!!l,}1 be posIiH'Qm~d: ~il~li,l1ud~to II peptide S!llbstit!llell~,hill Ii NAM
!':Ub",UIiMC. he T l'iulg
l1.C<lre~1
dlC'
'!llll ...
heEl N
(,'MU.~ytic 1\..0",,""
IFilg. 2. G1I"s:t.do!l!l!.:lifl oomp1!lfi~Ofl with l.ysozVffll~and Wr!f,~rm.atioi'laAch~~!!J~ bNWi!lE!n~h~ aIP(;II~~Zy,rl'!~ ,aJnd ligand'@dl kmns. fAl· S~nu:t:~l!Nl'=bl!!~dl i!Ug nlFllent or the 'G,iI' d'oDT1O'lirl (lJb.J!e:)wliUlllhat of i.t (grNn). ~ 1iIe.~il::esHill arild 'frl3 (if U nave m~Jnt'e'rp.!lrtsin alll re!ate1d (he-, and V-lypoE:' ~~ot.)'I1l.e' fdds (2;0). ine ptltauv.e H14 catIIIlytk n'5,d~u;~f ~he 'Gill'~:] domain at1.fJliIs~\!e~lwith that (If ;.\l '~lbothinl s~kk o rep.,e-sel'lta!Mlorn) at the Ie\m~ Ql I.l, ~eib; 1~i1. (lmet) Or~hog:onaL view \lli~h lcQIl!!i ,omiUedi. The lyooz.li\llile $lrm:b.m~rml1il depo5ililil:rl 1[l9U tg,~s ",lilo~~n,lI'IIitl1 tile poly-NAG tigandl in slirk 'Fo.mm.The ii9~re ~i~hli!!Jl1ts ooththe Mrikii'!.g sln1Harily ~j'l $Eioondi!iY $~rlll(tlJre' ~~o(aU~~d: i:JJ'ii.y w lh~ Gr~l head :~ubd~l'lIIfllilfdilll!l tll@ ,~el1:Sil?lNalio:nl!)f ilrJac:biiHiit@, den bEtween. tiill! two !!'n1yme t.ypoes:. l1he rogiolil~ beili:l~\!' d~h hol~ fI(I 'SmH:rural$lim~L;rilY ali!d ;:lIFE IP~e$'l,lunab1~ the dmerefl't be~a~~ tClf the !lleoo~(lr lh~ '61m5:~. d()lllilrirl t~1 00 to(illiim~~t ti1@ :m@lfIiIbr;:!JiI@ and :to ibillld! i! IliiJjdk ~1:!'Ils<tral.@'. (8,) (~.rdQrm;\!ti~:nilll :ehan!llE!in: the j!llwr.'Ubdolfl<1il'l upon binding rrn,oerno.l'Ilyc1n hpr'e>.5enled ]rrls<tficJ;::ro,rm)'. The '1'Il:()E!'rnomyt:1n· bo l!Pild 'rOffi'li! is bluJil',aJlld 'lh~ ~l\liIDenl'Ymlli!' is~~I,(J:\1,r,
Inocnmnyci~
in ,m ork:mmL'[)n!hall[lOtenti,dl~ diroct .. ,hie .Iil]id W'lJq,IPtm\'al'd '!h~ Ilwmlbmn<l. The il1~ratti{ms 'OCI.\VCCIII ti:lCprol,c111 and illlil±hitor arc extensive '(Fig. 3At ,crmsb;ilC1Iu \'\J]dll1lOCnomycin~ ilUlCi ~:]mcd COml'M)Wm_t<iposscssillg some or dlC It)\v\.~. mr.'t.liUIig inJhihi~rn)' onnCC!1:tr"'llm! v,rtI~c,..;; l.iJl.o\"~n~ <.!mibioui:.;;s(12)- The h(."I(~ sLlbd(ml<~ill tr.!'J il\ hl'M,,:liy idenlli:C!l~b('il\!I'"''!.'':11 ,ttfID;[·I1~n~..:: 31l1d Ihe mn.loclioln!"cil]~oolJnld GTs:1. $lnletui't.".S" \.... lIrl1Cfn'
B
04i
E
·,."....._\...-OO:NH;
[I
01'1
D
Filg. 3:. IOeMill of m!oonQ!ll1ll{(:i1i1l bili1ld]lIlg. (.1\)1 111iI~e-ra(lionsof lII1oe'JlolTI)'(:ln wii!lh lihed,ol'll!l),r Sil bS.tHii'OOs.ite' t(ifG:f~l' w~th
HO
,
r
IFU
0 0
,e
$~l~tiNJ s<ld~ (;n~iIil5 :mMt!, The pifl05phillric add 'd~f!~ler!Jrn~p' b~Il)osilio:nooili! ' ~ r,egi~:n ,~:I fg~1P()~lw~ (my!!)@. i! ~dl '~ile' h :~lIg:awr!lngs '~(i11Tlll an eoc~t:,n(h3d ~iHII@hil th~ ildive-~ite den. Ring 0 projec~ ll,ulin[u~o~lIltJiolil, illild' Inng .f!. is ~otal!:ed close,),!. tD Hll' ~ir'lker regijon. ~B) Ci'll~rnical Suutll1!re or lIlI~enQmydll1. Flln;:!i!nly, an R1, '9n~~pis II!sedin piau: ICliffhe ~~ Imoenoc:in{l& npild un"i. () Etel:JtroiFIdensnty o~ the moelilO'myd'n lllIooocllJlte. The majll i~orf .0;1, :t!rOolrCromla.t. c:on.t:omoo .at 10, 'i'roT!fl~h~~ffiI~!l mooeilalt ;,tB, A re~lliItnOIii. FQ'r r~~~~m'e" the ~I.!IMti~ calQlWtk f,!ffidll~ U1ll4 al'!d £:1.71 are :shOiivn ~Ifi~tickr\(iflT1l.
REPORTS
ch;!i~~ ooc~r in the j;;)\li' :2B,\'fhc :Sl.llC!~.y i;;O:n~r<'~ P! ~$.H!:.l.~, 'Kt~:4. ~~ jUry f\;';'iiidll~.~ ~u1d~I~~ b~i,\"~y qOl:llL1"\'q] 1\ 1~l T 15U, ~l~ E ~79 n,,~d\,JG\". :!I.II;: ~ocml.-d in i:h~hiJl!:,!,~~orrl lx:iw~L,"1fIi dlt..~,[,,"'.0 ~~bdonmrn .. '1!1(.1lImy 00 imJXlll!!nl i~ ~lIrlO s ahili:~'y of '~hc GTcH dnnl;;ual lQ 'I.In-d~Q !'iu~h .. l,;(lnrpFr!'iauion~i dRlng~. B~~..oo ('1i,,~nioJI ~~rilil~~IJ'~ il 01'1 ~i~;:'; bC't~D'i rrop.c~(;d Uild rilfli~'1 (" and E or m'ioelrlol1r~J'cin would. b~mltl :1111 S3lfleW"J), the as a N Al\ll-NAG db:aechll"kll:: Or~1(," 5il1b~~lIIilc. I h ~heconmph:::.;: orGT:i!iw:i1h~'~nmnydll •.lhe mil':: OX,~~O.eJ1S ,ll~d N-:LC:etyl ~'OlIJ)S of ril1~ C and E :lInCi<lH Ih~c 111.'1· Ilhlk~C$, are :if1ooooin :~"'Cm11!!~11\">'ilh <I li ~Iei~r glycml dmill. ;\.!'I'lM'!gl ng Ihe~L" li~lk..g~, tDfnh:c ~\'iI1S dlr1i~l S!!.!ll$.l!011~ W fini ~n~e ~1[1~CQf~hc~~lh~bitrn' pk!c!"!s!1:!L;' !txillc!Hlg iI!,'{)~·1end uf 111'1.(' dmilll lflllitJ;Ie.-.~ fmll11 Ih~ l.'OIl!OlIJilir: nmchilu:,'I)'. idc!lnii:a~w uJml on!1.\: h'fi{)!r.)'~IlC Ihm~br ,~~I' ~'fllt>'l11if.'$.
'OOnro,~I]g~i(ll};il_~ s~bdrJ!!l(l1l1in {rg~,
I;]!~
Tile ,dtfiJ;'u.nocs bC'lWOOl the moermmyc:in wing Ffphoo,'nhol'ac acid "i\.~Wl'!('l.!i mo,jei~C'S <1nd
~11'!; '~~~(~ S NAM. l~yf{JI·}ho5PJg<l!~~C):> mO~lit;:., ~n~li.i bt;t 1'l,:,<;po:rhiMc tb" f.iT 31 .i~l~i ... bHitm by ~ilL'1~"'I1Cllin~.ul!r, 11'l'V!..';SiigJJliIJimi~ hypodl~i~, To t \VC ~naddcd n I'UU(h!lC!l.\·Cc{l1l11ri~1>i: t'I.fbo!lr~ t~N ~i p~d][
;u:!d gm~\I~llgcJIOII~n t;,!JmI:rnh.~r,¥H !hCIl1.1[K,,'!Imllli}'til1.
I1IlQ(f.c~else :'iiil.lKII~r;;:;. NAM p~pli de SL!bs.tit1!c~~ the ...n ~hm IW ~[l,,~ic ;4;,h!o:'"~QC(;\!f \'vilh !l~~ cl active-she
l~h~11I
;I
l:ioll.md ~~[l,Ie~I,I:r;e. \V~i~~11crl:ipid H i~ !he dOHmr ur mx:~pIOi in tI'iC POiYI1lilCI'.Wil],r)1'i Jil.;'liCli,oJ'i hillS, bcci:i. ,~:;w~~~iiw~)' ddnilecl ,~jJ ~ Fto,i'I'Il Ciillh,,>Lmil1ts,
IH~d
impo:>~'l:i by dt,-" :>~rIiieIHm orlhc 1liPOCilz)H11!e· mOOllm~~yci~l;bou nd rorltDt~. wecan new a~lmo~! dcliniliely oo.ncilldcliml.lipiid~:~i!\ rllC I.lCceptor, :lrnd the a;ro~wh~gdl<1i-n is the donor.wilh
Slq~P(}1l
:lsl!Immmil;.'Od llr l:VcIz.c1 UJ)!.· Modelil'lg rhe !ilrrnw:il1~ ch,~inIdonnr) ~U~l'i~.rt~ mo~h~ !11t}~!m· Q !.tl)rt.~n~l!nir(,Hlirc. Nl\:GgJ'tlUp .~~ fl llil;:lo~"d 1;:1\, cr Icing E. and .he <l(t.join~ NAM SL!~<!!, !'illIp;1:I'n:g il11~'Il1!<c.>1wi!}1 ri:l,"~ f. The- rr~{lot.'nrIH'}'{: in-bu .. sed
~:rom, num('.t'(fI.!."
C'.xlliCr]mellltli
~Olll!l\.""('$,
each ril~g f 9.lIbsti'[lII!e!l~ of m~nnln}'c;~11 cmreluting \'\Iith !;h~ 42; ~~L~~~lC:rll. ol'Ni\M (e.g,. ~1C (" Ji OCUNU:l ~llj of cinl!;l fwith Ih~ C5 CI·I·21).I:1 lll,mup u r ]\JAM), I](">l'ti~i~ or itliii!i :.nnggel'ing •. Ihe ·1 p'hO;S;~jJhiilyl !J.IlOup!;of ~hehi: illl~tlhiliLI.i" alilU, lit!! !>~h.:>H'II.c poll1! in :..illIli~!Ii' ~:'L!.,>;i!in or:i.cnt!llion~, ~IftiJC3.llle~ o:~ 0 fm.oL"'Jlo-myc~in ismu£hly equiv(' I aiel,1 re the modeled rin,!_l. ,o.'\)!lliet~ of NAM ~ru,; S\lsgcstt.'<! ear~icr ~1l'l J 11'll' 111i:-: l»si~iol1.-<lhc :Ill: roll ~m, hyl 011 Col of the htll~bi~CIl' .. above the l'llilal.C 01" NAM .. 0\'\.'11'~l~ ri~~g x)'g.c!~.The mt~h)!!gm~p o li~';$ ~nW(!""l 11].(- o~tjr C~. sill,,'; (1!f ~!Ui(r(:u~'hitic d :.u'l!:iI1;k~~nd R;lgitm(~llri~doy Ih~ lEI ~4 lIIld Ih~ inc~ming4~:~ I end ·of (l'IC'>!.cccplOr, In eur ~!l1!C!il.!!'l'iS.. ~."l'UT!]'I!c:Sp::jOO lO moo~1 du~li~]id lh'-CToi.C
IF~g.4,. IIfl~i!)pOO:i!dmedl~·
(-I.)
(+.2)
IB
'il!!'IYiud. W ·s<helT!'.atiiCrw
s~del R~dlJ~ I~U~ ~(Il5 'to de,puowlltite ti1'~ a!u;:ept~r 4~("!f! ~IFl~llP" whkh '~Olil~
avoidlNl ~, prE!VMt (oofiUi~ion \'Iii!.h the mcencmlyciin liillg ~i!libelin.!lI). 1111lIlri~ 11Il® ~I!!~, ~ip~(lil 115 the' a(~'~Jnl(lr(Ii.gint ~idl:l'~ <In.d l]w grUlving glly-cen d1llillil is tffilf donor' 'aB'ft
~.lJid IIII pll'lymlllFiz.Won. FQ,f darity, 1't:1I.. and R2 gmup!i ,a rE ~~ 'inl pla~ 'o~~he OAc <3! ~ d N H'A,c:g raJ'!I ps, I r~· ~JI'I'!dwel.V. !'Dr '~®liTIpaiathl'e, fgl!! FP1i~~. the f~,1]~tl'~' ~.~ Lym,~ s9,g,~r rulb!iil@~ are ~~beled ki pa~the5e'l ,(the trad.iliclIl!larl [email protected]~ !!!~tn.gs!llbf.i~:e'S <l to f lla!5lbeetil
a~ffi'nl~d Illr'E!Cm50r,sinto a ~li1!·ijnkif!!:! product Irw~iclLI!1 E17.iLl'IlIarYas(;ttlhi~ J!!II'Ot'eSlS, by direct. protonarliiml of .he p.h~pharo:e-suga r oomn:1er by , ·~""'1'''''ln ,'..~ IPJiI' 0 !IJ 0""", !~, ""', ~ .s'~!I!1'!B!. 9 i!~!I;! ~ .\p .. , ...... .... '~r'e!1!pti1rQi!I~n illOO~tlilll1 mUll
ill diva~nt ,cation, (I) Spatial rl~;Jliret!ntation oUIhe ~ipid: II ~i:)wm:e~til),n mo~l. ]:he ~~afl,e fli!terf:alC/i! 1!h(l~li!tal bladk H~). lran:smOOlbr~me Ireg~rJtwrtx.:il. blue rl(l~n~~~t ,;3!OOmri&sing lPo'lypeptlidie {<ted blU!! Ikl!!l aJl'esJimwn {or !!:ffsct 1fIhe :~>o1i9'in ~tliuotm~ is. unmoo~fiedl Irom the nFlCl e-ii1lal,mfcl~·lxnmd: co'Tlflplex,.. \wlil ~e g~O\lliilllg gl~nO(_Min dmil~,r (~f~ side)
mow!m\FlI
stnLKIi!.W! ~il pmiimt. this ~ibllliorourrili1'1!1'.ih~stronger IJl'DiSi:liiw ,c::hiu'ge 011!he Iil!~t. side: 'of the acl"we sit~. r~l.3i~Weit(I! ~aif i!)li'!~he righ~S<idt [s~ ()1 -!'fIl1Iy as~lst fur
modeled
~htr~!'11
(Il!j1eY
lrani5klC'ilted inrlli!l1 dire('Don dffiot~cI byli1e ye11G'1\l' arrow. AIlIf s!lIgar cham ~fnger lhalll fallllr $lJ~j Ulni~ (d~l!In~~ ~ ~nWfi1ingl'lJ,Mo s~a r 1!l1ij~~.Q1rtliIea;C!c:ep'trid W~j'ld proJect: i)~Uirllm the GT~! dO:i'!'!ai!i1311d Iller!! are no S'!,e(J(: oo!'IFier~~~ ~~r 1
;nr1lJ,oo~'N!'Idi~ Iocared 'ior,~I»SitklolttJ, brim!!;1o1ih P!l'lI'(Ip:1liJsphale~ fiii! I)tiI~S-\Jb~lmt:e b mod'el. (10) D'etlib or the Il¥drophobiic pl.atfom m !!hE! GT~1~o!.d.The Vi't'lY 11; i!I~~I'J!(ii't\atety 90" fmm {(), ~vith rl~idut's.~ho~'iiiii!il'il~tnck f<.!~m. (Jirtletii, h1'tI~pI1o~cpIMblm gra~j.El.l4 and IE17iL.
pmi;itiiofll" {CJDetail of .adive,.s1te ~oc~!Ji imdl deft Residues IEU~ .[In~Et7 Jl.'Ull 'liffiimm illls.pa.~~-:frill1J~ 'foon"lli~ eteormtill~r lPo~f\tia.'Is bed, n~!l!gatN,e;bwe. positiive)ilmlkal:e a [onlll!!'lVOO regi~lmor pl1l5itive (iII!ngeacrQS~ th-e mid~le of the pcI~et. Th:i5 region Ibl mls the pfl®1<phmi( acid dier;t:er 'gnmp ®f moenOO'l}'Cin In 00 r
1404
VOL 31 5
SICII~NCI! .www.sr.ien.-emag.org
REPORTS
~~ ;!,'Cl;eptor .adjlttcn~ to Ihe I~mpo~(~ dener sile, \""~~l thc"-"OI"I end, ;!'!,'"il <l11!e Ib'r ~,nWr;:'i.ction ,,,,iUl
b{'J(hE~ 14
~[ltiitio~ of
<B~~ mKI
'C~
Or!!~L;' g~)110f~\i~<J!r,
~illlb:;;~i.tt::s. H.
lipid U i.~ c;fDrIlf!Hl"dhk Ih!li +.2 af t~!..::sub ..... irnt~ u~m It V J)!,
'0
Thii'.
or
si!'l'~i~"'ro t
We r(opo~ tlil[u~ E ~~4· i.~ Bmu,'11,t'db~~~ diin;.!;:'clyilb.. ..I.!'iI~Ca proh;)!l rootll the .iIi-OlJgro~pof th~ ~ipj.cl.U ~.oC'CptOi':. The de. porolmUlltlc] Ibn'"~ Qr .E11.4 l'I1il~ be swbi~itctl hy ~he atljacci1i1 .R.249resid'uc,. ~u;ctlycm'l~\'cd ss p.1III cd'~nQti r V: 111(.'l)uoWnl alt'>lr.ILctOCltl step I)ro~ m,blji OCClhllS m'~~ro.lniiIJnli)p,~Ilth Ihe c~OI. rol"hiUc: ...
~nigm~i(ll1 (If the dClI10r C I wwmcl lIw m:o.1'tor
,I
d11: ~ ~A-):iI1I,l"xll~rod~ICI. The lipjd"PYITIJlhoorihme ~c<l!vin~ gmt!1l' r ~lt: ~~l1or is then rh.>e tQ g it111~ ,,\\.[11(11' ~d be ~}'~I~~~ in liri~~ I.! ~"iy!¥~h\i"'Si.<;, '\~ rmf~Cl!>\,': "'mt 1J1!IIm"T~,u~{Jn 9f tti!g ']J~wb'~'o:mu,;d rruduci. ~() illC dOrl(lir :$iE~:i(S.\le.."i,:>ilL1d by .~~higi~t'r . al1ini~)' fur I:h~ p)'mplm:>W1I!J.1~r; nmiC'ly ill (he donor ~iulc lh;lI~in !he n'l;;t:cplof sik:. with ~:I~ oon·
(l!
l(ll
'ii, M. Tenr~11: et aL, IIlOt Mi:.,~cL 314,350 Il",9l'. t, Hgjllll'. (_ ·S.... IIiIij'~r, lV~·d~p(Aciw /iI~. !!:7'. ~.;~
(i~·99).
H. P.Wil!l[eI. NvL •. nor,rI!l~f.dJ\ll;';l 43i,. S85 u'9®7). 12. ]. Hi!UMli!.~. D, Ml;f(f!!It!'l~" ML!idWn,. ~, !!iJ.j~O!t!'!~f1'!. M~[jti!ffllllll1!,; 8J«ilm:. P~. 11, '9:)7 {2,OQ,6).
w.
c.
ili'fL' ..
KctVt."(H I'iol;i:tv~dy c;lm'!!,l.1d K15$, KHlJ. lli.fi7. 3 nd K'~68~id ues loC'al.cd near dl~ clmmr r,y nlp.lbOt>p.h~I.ci<liiml dl~ llcti,'':: :':i~eI Fig. "10. ni~ moocl i:sa~!Iin rerl1nni~irll. o.hhc: IYl'1m-ymc aetive site.• where the t~. lind ~2 SlIIlt s hes ~1Ult H1irl1ch the modeled GTil'1 aCl.'\.l~tor '~IJ£lli~.'SJlOS~S$ ~ the lowl.'.t>l. ~,!b.~~r.W:::I~l~ni~y (he ::ub5;~I.cS. oJ ,dl
,or
tw:«freJWit,lf 43. :~;'l3"'S ~oool'!i. ~~r,~t!!1fai. f. .l!I'o!ii(1gri'(l~ l!:S:7. :i!U5(2005l 19. M. fl!rf~1t M.. ~(i~n·DrlSlt'ClIn~.,.), ,!lat'1triel.138, Z!l211
~i;L O.
~4_ Q, ~~~~rr ,!t/ ~", T~r,'IW~n 5:3, 11$15, u'9?n. 15. A. It tEung. IiI. ,~ 11)I]~!1, ]I. f. HOmg. i.1i. III Bi!I'g1ulli!, 8ic;!cfrli'miJrj)' 40, 5665 16. 8. Scllwar!z et (i'f.,~I1M1'i~ 41 '125,52 j2002t 1.7. o. S.. 8~fJiI!'m. II. (Jjj@A, Nt. lit Iitt~~1), S. llhltl!1r"
{loon
2,11 C. ,1.
Q006~" i'~,
now 1)J'O\'itic<! b.litl)o [or 4,-Cm ~!I) ~F~g:,4, A <!ltdB~, I n ~lC mOf:n[l!~n),'~ addrol'1i11ng rl!!nlw.:r quc::[io~); ... bOUl tl~~ medmcin~'lln~pl~x •. 111..:C(I!U:i>(;t"!'cdE i 71 re~id~K1 lie..... Oi"U~lor[his i!l'!]ullmun fm~ibiOr i;)Uf:;)'nl~ OIndror do~~~r ~o lill.(l glyc(·ric aeld rm.llili~!y ~'1;mmh~ th{! d~'$ug;I'! f !l~W'h!~lUbm;~~·ri"'ls. Wf!n.. ;I~~O o nil' (l~~'1'l'" Ihe door fool' umh.;'I¥!<lfl(hg: !'i~Il!C;tlm::' and r'hfl)11111illC'~u~r~i!bk<l!~'C n~!~ ~lllO'''~l<I!C in l~· rl!!m;~jiun rol~niml:iih~l'I.s in o~l!.;', ciini:caUy ill"flOr~ our !1ub.s.~ml!C' m(ld,6~). which i!1oo!nb;i!l.n~~111" itlll ~,lIn1bJ11~h~ or lipid~~I!il'J!![:'(iTs, pH ac~iv.~ty pramc:;. nflhc E. ~'On PUP'I b I..'U!Zyl~\i' I( 16) e~I:.~,"\ ~OIlJh;: dm")~ CH~ \..'h~lhct En~rl't'!~PIl.~ ares 1.11l: iiug,:liT"plu'X''.phmc ~ink1J.!Wto :lss;;is,t o(':~m~~ RererffllCf$ andl INU'UlJS 1 (. G$!l"m, ]. ~ (:hL!)llI:ll, ilJl'u,a!lro1. Mat .Bio.I. .lieii'; 6Z, ysis.Funll(~n'U1lom. liIl.liil:1ms. ofaJlis; rcs.u;d:I:I,~ i.iIi 1,079 '11iI\l91" II. l'oli il?BP~breUllin $.mm;, It-sid1l11:lactivily, :2. T. D. BU!j!iJ eI "1.,, .8irldremi~rJf 30,~08 Ih!i!'!i'Ji}. whereas IIKiSC of our predlcred Bml1.nod base, 3. c, 1,'I'II~1l;1I,~.:t ~:t 300 f19~'3l',
l~\O Rtl'llClml!S
These
U9ge~,
r...
Ad)t,~ru:!~UgM $oo~~~.f~!! ah:tlllil: cCOI'cliimOll!I!fI:l!M f~!Jtl!Jre ri!d:oH af the !l,PDffillIIDB- .ilimd rnoM~m.~dlll ~Oli'l'lpil!'~ lmt'lb~m dejlaSiltd .al. ~!h~ Pm:l~ei'l1 i!:!m Ba~k. l'ii1ill1.~C>oo'~IQlllllimbe~:~201~U~ndi .2otV. ~'e!i~i\itty, figultli wer~ J1fl:iPiIreQ ~n'D~IIP':I'Md. Cffi'!:tI.P)'!'!JlItorgl. I\ILCJ.S~ i~ H_F1I Oi IfIwgMs Med~lli'I~ijlJi~ {1itH.M1~ irnt!lfM~~~~. '~mBt!r, ~ c,,:IiI~dlilll 1~!>I!IWli.e ~~I~h 6f ~al't;lil. ~.(I511U:Ki~n~~~, !Iild a Midlael5lfiii.ltl <
I1i)Ij'j~M10i111~~ He\l.llhl ~l'ird'i
~~~liIroMI!llt~lW. ~
Ell~,
~I\g~!
A.U. i!; a MSFHIl: olInd oCiJIIii. h!II~.1i1:t~: ~It wM!umd'ed !J,~ mil! .al!d! "'H~ ope1aii~SJfl!Ild$ .ili!\!I ~!l!rri!¥~ll1lch!«:' rlin!k rr(ft]. 'IDE!! (aJl'i~1 Ii'l)IOOaimon ~ !ilnD'l':u.iDIi om!! M5~H!ii"
ate the $~Ib~~rte, then we proPOSC'! hm it he~lIS.to a comd imm: lire r)lmpl'KIsl~halc gIllli!!p of Ihe donor. cilJ!~r diroC'lI},ar \~n <l di\ <ll~!l[ ~lOClHd cation, llw
\!m;nab:l:cpH Op:tl1110 and d i\,[Ilcn~ c;]!~ionroquil'!!meats orl~lcGT:'l' rJJ11 II}' or !!~lZ}'mcs.(f7 19) mmy roi'i~!lt fw,m ..<!!.:r)fi~,g l~oc<ll c~]\rirl(lJ~rnCilll'" f o ~hl,;'E 171 !'CSidll(:, Th~ SN1·!lUkc !iC'<l!(;I'~OIlU(lr;111'S
fu~"lW(i!.'1IId(mOnllld .ao;;cplm~ C;<lLI~;lIg invcr:iiioll
<U~
~.timii:;fH.
5.ilip'po.ifllii!!i1
.i!5 .. 1]1·74 (WOO. 6. ffliIq~i!!ls ,iI~d P1~Ll~5 ,iIl~ ,iliwililble ii:I~~I.R),I)Q!1ing nia1iE!rn~1 (iii Sdf.jJ~ O~IIDil' ,. (. CmMrl!1ilI!o"Mlim:~ I!f !!fl., J. M~I.: 8lot 355, ~8"; WlIilQ.
I. ~['aiJle-letifr ibbrf'l'iali~l~ b.r IIIe ~rni~!) il!I:Mj resiliue.t. •we: i!~ fOl[olM; j!o, J1!.g: C. c,~; ,I), i\l!fI,: ~. G'!w. f" Ph~ Go GIro H, Iil:is~ ~ ["Ie,; t::, ~J~ I.,. lUlU; M" Mi!'~ IfoI,Asm~ IP,.PrO~ Q. Glm: R~ A~: S;, :5eor; T;lhr, 'If. ~:l; lW.T~p; aliloY, [~_
rale or hii~ollc 1 [] 31:gC.(;lQmte sc[ll~ ,<;(It'lS If! adclrt."S,,> 1"\fO q Mll!S,lnOIl\.'i.. First ~s t~!cre C\.' idcnc(! lOr ~IlCI1l1lrntll~crill~i()ll_,acrem:JCIlt IB
!'if.K:lCi ilIel!mlO'!'er
:Ire thc~' <Kk!i!~ol'l.ull ll:u:."ohi1lni~nk"for hi~to,~!c I~Fl!nvC'r? To n~J;;'a'iiurc'tumm,'~rr.-II<.;'S, w~ l!I:,:od Y~1<!."l I.,;.myirng cOlrns'iWltivt;iy 1.,.\.\prl.~~ Myo-l.<Ljill(.;d, hi~m:rliJ H3. us; \\'\i,;'11 ,aiS:in indl!lcill'la.C' fi~~g~'~~gg~;d UJ (5) (fi!il. SI)_ .FlJlg-U30 Wuili inrltll"t:d rill 0.1run:'S~c1 ccns [or '\ImIryiug 3m.oUrrD~S [~me. dlffinlUllin \'\'3S. cro~s·linkcd 3m:! digcstcti 10 mI.m]'tOnlu.ck~Cffimnes (J 1). find Myc- aM FI!ilglo laggm~ 11l'it~)Jl(s\V~1C iUDmnlln.opn..'C~pimlCtI SCi!l!;:i;. 11!I~'~~m\~!d! Socolld.
or
i]~.;: dYi'I!IUnic bdli1vior of Ii> :kiey 10 umfcl'Si.all.d:i!t.lglilc _ diver:s.i;ty .or bi{llogrca~ oo,1cs of chroma.ill. Nudeooom~.. arc C'vicwtl :11 man;y YC:I::1I. pro-111Olel1' duril1,g gellc ;!ctiv.<!!~ion ~i ·1I <l!nd ~()e . fl;1.l,tci)S(m~e:;
.C
I,.
Illal".lII.ci.crh:ing
is lin !~ hOi1l1l0:iOg). PJ~gL: of RNA po!ym.cr13.3 aSI: U I~Poll n.)~~II[s ill c:dc~ion or ~IIJi.cko~o:m.es Iro'~ll S~Il'te (N). but l1C1t a~1.(0), ending i"fStGl1!s. In contl<~'<t.sl.u,lk::> ill P/~l~'amIJ~ IN!l)n.'lJittll!lrIJ slIg.gl!!l1l~h"'l 1B is no;~ reU'l..o~'.d durIng f~~ I~:
~mn:ilcr~pli(lnIf',. FUr!~lC'm1QI'!ll,L"O~J!II't.'9.IIL' {
o
}'~~~'i'!.
If.a.ruHy ~ l\ifI~ and Sd;eiiil~e~, Cemer rtll' S:P!I:eID5 l81olQIDi. 1'iI~I'i'.. rd UlVwe~i.~ (a.mlli~ii!lge, Mi\ 0213:8. USA ~SClrg~. •. o~ COlflllu1el5demre and Einglm~enin!l.Jhe· IH~llm;:w'Jniversity. ler~!em 9l!'90~,II$f<!eL ~Depaflf!'lm! 01 MeteclQJ !3r;:f1~~i(s and! Buot~dmOlog~. !fi!i(iuU'I Q~ ,~i dne. 1h~ fiI~brew Uni~Ersity, JfllI!i,~tt.rn 9111.2.01 Iw.alll. ~ Dli'parll1"l@mlt of IlMol~G~ (helfii~tIV a!illll Mole[ll!~r P~<lrmaoolQ,
!il1!ro'¥~ U... ve~~, i OU15, USA.
2!l{i
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!!~.
iM
ro,u.~mbl~d in Inn!.'!.lnp(ln!\.';Il're.s'.1'im~ {5). [~ Dro,!!fJfMriltj·,. ,It"ni\'~ 1.r..n,w.rijJl.l:m). k~,Ld)110 ['1Plal;;o~ !lk·lm~nrhiiS.tmut U3 by the V!lti<lMI i\iofanti Eil1J..3 {6., \.~~it'reSH in buddi.n.~ ~i'L~L (W~'b!lj~~oJ'ily1113
r l"ln~C'rj~)1i[]l~
~n
i~'l i!1clc~~li~!ll
di~agl\,,'C!'i1.c;:ll:l
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Illricl(lCU ... •
"T~N authoJ5 (om,tliiliuted i!qua[L'~10 Li~s WDr~. ~parimenl '0. Bie<lh<!lmi~ll)' and Mo.le@l~t fJ~~ffi1l~C~_g.g(y.:ljlll~~f$.1~!I'gf ~~(ih!ml~1~ M(:t1i~~l , ScRliool, WOOlE!SI~r., !WI Ol&lS, 1J15:A. $lo l'ihom '[(If]r~~nden.::e silnw.'il boe adld~E'~" E"f1I1i1li~:, CM:Ml!r.Riioo~:UfM5~ll'Ied:.12dlll
tPres.eiliit <KIdrilM;
REPORTS
Fii!). :11", 'I'inf!~ (OOF$fli·~r i"I~s!:e1l!l! tliimill\re(inyea15l
t4'l "3
tl.!J'i!i!)~fr~:r
23
adj~oont nlJ~l@omm!!'Sin 'G,m-ii:!nN5~ed yeast eul~Wies:, fi!ag i1li1d Myc \'I,I'l'~ imlmun(lpreci,pit~ledl a:t \\clriol.lll, time points. ,If~e,r r~g~H3 iliI,dulcthHi! b' ,a~s.)j a~d1 Rag!M.yc F@tlo~ (y ,a!(i;s)~ver~m~~red
by 1iT110'~~~. m~ A 'oomr
pulalimllalnno!!h,( redu:ces HmE!' oollrse ~.(!Ita to .. 5ing'le t.llJnwer p.<!lkI!lleF
'rl.ll~
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terr'l ,(lteqllernyof
his-
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s.entetl
the leHmO$t
S,NT1
~~I:l1ffl@n m~~~'lI~@~ii!fI~S!jm!.!liJJ~!!l
dill:<!r [email protected]<llle thO'll. ouu liIiIode~. (;dpllllr,es~he IIrnl1!jDnity {d Fil!}. 2:, IR~:t.i!t~orl oolW€@PlI lli$lOO.@' IIIiI Od~lfk.aili{JI1lili and Hl3 IllJIfIilO'!l,el', 1I11:1deoS'Or!IleO,(rn:lui1lns} V€r"lLl~ iii Illn:o-' t!illr!On.s'lr'oli\l:S~. Nucleo5'lJlmes, .n-e ,oM!1lrl;!dl by tlllmOlllleir rate 'QreM~'.MQd~'~tcatiofl and H0;1 ~els (12, 18}.are shown ill'l ye~lQW4!o·b'l1ilE! MleaJt lma~.~m:ei'l~ :~~Imllrep.re5il1!nits. l '~flrkhl'l'l~.r!t iii£! bQ"H.'OfIii panel shO\~ '~el'iloif11k ro[a~ell~ (l.?): 5 '~nd ,3' "i',S;S, ~~fe~ llO II!!ll!deo$(lm:€:!: Sllrf'(l~fldiflg nIl! trarn.crjption"i s~rt sil@; Pf(l--oIIIl ot.,e;r ~ndiutes othe-r IIIpstn;~'1lirn p,robe-s. Pro'leitn..(ooilng seqlu~m:eos ,ilJre Se;f)i3!raledinto' $', mit;,ldle,. and onl
~~:di! tl~11
m~sroll:~ tumnowr
~ru mi(l;,lIurin{! Gl arJie:st. ,~tC) [)l~Iri1lUlia,nDr tlllmmrer ~iilte~lor IIIlJdeoonme~in [email protected]@dI.~R a~nlll@dI~mDwr iFilit.@'S ,{lfl!' [~~~rw(J@(i as ~Il (B). (11)')s:aI!I'1~~ g;efJlomic:s1reldij, Wil~hliI11!cfitlOMl'.lTl!!$, I'M m'lor c:oded by hi mowr ffile.
Wi! ~OJn;paii\'.d high.~lmikllli rL.amO"'(!l'ilil~csto
1III1JI::::r::J~Elcmllllrm ••••••••
w,oo-lg-
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.',
IlJj'I"'~nr' rl'r~
II
!Pn....·i:Ouslly :lneasLjlL.'tI remure.o; of these m.l~I:Cl}. . semes (/1, /7, .1.''1) (Fig, 2. nnd !fig. N~Jr;k:o~lncs O\'Cf' l)mICin"!Dodi~lgt1:gton~ were oo.ll:lC:::l, \'Iihcn.ons, l]m~~lI"!lC'r nu,'ClOOscml£li, W('!!'C ,gCllcmlly hm. ('~CSillO!ldin.g~y. h(1! nlld('OsoJ'n.~ ~nl:l dep11ctcd (If the h iSlionc :~nod~.I:ocalioll."lhm nre "~lcrool},l)ically'" dC'pk~tcd s'l!fI'Ol!!lwing jhe [r.ln· s:::ripIto!1 $liiml site f['SS} U ,2) and \\'L1't!' eo:!l'o,'l!lNCly tnn~chod :tOr UlC'hi~mn!: I~111\ \,,~rin~!I.I~tz i (16 . .i 8),
sn
II
These
rat
l\\to tr1l","'I!\Orl~~,
.3'" O~El'.r iii ama.taticN'lS de'lll'iile illll~ OOIilQlniiolisly re pticatincg $'eq~'enc~, {ARS:st !RNA g(e'f!e'S, ,~f1~ Null (aiiW
ether 'iniulirgenk I'Iegian).
::lJ:lja(;cm. 11illCi.l'1(Jl>Olim(;S with {Uri Ilpp.roprilltely I1ilooiifkxlnluclcop~a.'i;~nic pool ooull:l bcp'.mll~lll~ Iy ill::l'!llmlSibk for promolcu pauems 0 r h is [om: unlCxliilieat i!:D:Jl. Second, C.l'ilsurc histone modil:K:31.iom' due 10 rn(pid tumnver watl~d result illl a
or
:s~e,Jtdy~sl;iltiCpknlre of srereetyped
promoter
rmiC'ly. t\m~lilicd DNAs were COll1PClilivcly ~'lybrkliize(lto a, .20 base pair (b,p'} ~S(),I~tio:n Illi\:,I:wl:my 'Qvcl,Jng 4% or ~he gen:on1C OJ ll, )'icl~:tllg P'Llg,lMyc m~in:J aI; cachiime IlQin., for eaeh neclcosome g:UI {H1r ,"'~~,<I!1' (fig. ~ \Wt, ~1!C'n At
l;'~lim"Dcd the W!or~o\'~r "II~ (UII~iIn~b(;r H3 T~rIC~ilil~';~luiax;'d t(}lllRJ¢tuc-uiu!I (If.f.il;Ig:...IH Rir.l~lo~um;:l:Il. ~'ll;rll:-; per l,m;~l of '~il;m:) llf 4,111C':h ml.;'IJ~~,. (30 [t) 4:5 IrninJ limi:L'l; uwr ability Ie, m!~J~ll:rC the IIUc!I.'i)I\UU1,.;' u.;;.illg .. i1iinlJJlc ;1n.LI:~)'ti~a1nu:xld th~l ~Til::;, CJi.pLiriim:J:U.fi I ,dfll~,.".ith a ~11ilil~:I Ul1~bcr 01' ~<t11C:>or,hc' li.mk"£l, ILLIc:k.,,(~i>QJncs, w~Diclii rt{x:,u· eJlc n
or
tha't I :~3I'Cpll~~~lli:m, r.JIl.c8 were \~Ic~lkly amlcorreImcd Willl ,~13OOClll:t3I1,c)' {jig. ::i6;. These ~~\!hs, ;u:c coll:iii:5l~'1]a, widl~hOM:' ex~~t'Clcd 1'13 I)C'J)~<lWrlleLlI thlnl ,;I I1tlCpoo~ ofl ~1I, and 'tiCmQlr~~I .. ~tc~h'l! \VCC.;,lIH .!1;"'OO~l'\Of ~~nig,u"n· tl',l1liVc~ I!.llllU\'Cr ro.n~rro~ll t~~nC-~101.mw .:x.pcri·
chronllmlILl dl!3~docs 11m ca:pi~lii.rt: mUliSicm SliUCS, potent:iaHy hidh~£ any m.. lbcr of i nJOnll'~~~VC m
or
h~S'l:o:nc mooincmion. C\'ClllS. ,Analysis or median 1'C1)ia.ocmenl J,111eS ror; variolll:il gellomic Iffi;:i conl1im1cO thatt 11W"I:IlOSl: .r.Jpid nunnver o..iC~l~ 0'1101' IJJ{u:n(l!~c~.'R, A. "n~c;l, IN ~;n,..l~U!u~k:,:(l~m:r .RNA~nl,.~, fr~~, J"A and .13:, .;;m~l, ~!g. S~), JMo.~l. m~~N:I)Ii,iL1!1;-d, given !h~, Yllillmie: d 1.11. J ~:cpJ!<l;t:;L'UlIl:TU m,('" Dl'fJ..mphiln!!jl.n;,,-.; (7;. 19),
Wilt; ~'IO
~10\li\";, :113 Iti,nm'o'C( in codd,ll!!; regions, I\:.~!lili\'c n "[llutcltlfltovCl' ;Iij}lc~:l.lQ\\1CV- \1miat iml or UiI'lIO\'c:r ;mt~>:; mong cedi IJgn:gr:on~ l'lInllOv,,;r rases, m:; expecwd! gji ...englobal 1113 dation ~>Ji(!Ialli'Cl ub ... it)i. d~l~QS"ti(l<nd:urirlg Yl1mnk rcpllkal,iOlI (,f\ilg., .2 er, (liver a wide range ofpat<mwters, 'lite rnl]o migilli. correlate with po~Ylnc:msc ilC~.~c~ FOL al~~h.. lacool' brOt""\.Ic~~eslimalecli ;r;ncs :i$ ;ro'btl'st."'I:1'tc reS'Ll ~Iing C\l<unpiJc, h Iislol1e~l1In~~)\'Ci (J1\tCU'I~IC and liS. S3). W'e 'IImlyzcd tk!lmm't.'II'rn!e.>; tn G1a.m.. >stcd ceU$·across du~eIl~ire yeast f:,reJlomc' l!sin,g I:'atc,cstimak'S~]an GncW I,woomers (ilfma~nilliidc indudblc gcne PUS; i51lKlre" l<ll~k~ill.:lip.h."1fllttol' ~iJ.l'l:i cclls,ll:ian in 1111~yalChr'On frlcd cells ~Fi,g. J.C '"Glmm::rci3~ miCJ1rnlfl':I},5\""ilh ,-lM"bp ~~glllJlion (depending 011 mc-a:5L~rcmli:lll~ phltrOI;m)oolwcal mdoo\so!1~ 111<11:Olre~Y[l!Ir'n O\'I;f .mod i~ol wld Il.g. S9) .. ,\i!i.r~ tl~~!l:rOremcarsl1ln."d. [:DOl ][ en! i( M, !7) ~Ilg. S4~ a:nclobbirl\;-d a high ro!Nh~lim~ "COll.r (m~ wli~n;Jj~m;"''1!~.l~U~l J,llc .~ lil<~!cr Ih.1U [itf: ric~1fl:liOlil 0lCrU...~ !h~ . I.'!~L~~1,; ~aN~ :;;vmlll"~. fi~dlilllg bt'w~'1/.'n !.ut~"Jo t'1l;1!11 !~~~ ~i"tino~1 1li1.~a,'il1J11I;~1~,"]'U Iw>o' ~ p.1~~~lh!lil~ lf~g, S5~ \;!i~ ..100 Ii'i!'i~:lilc"idred ~i,<ho.&;~ ti:nw !va~!,J.~n'ilyof our '''tt~rinli~i1l(.F.~" ~C md ~h"lpn~YI1i1~~m ~!irid:l~lli:J1' uvi.;"!" g'""iIWl':~",hubilt::d ,good I;:Om::I!1~lnOll r~ = R54. P -< 6 !oi m-171 ~~'ionic h:i~tl:mc Ot'£'Upllli.cy Ii. J ~0 r.!i,di~l,g F~~,J:J3).
(14, .15) ,(Fig. tEl II 10DCSI.~h,:'l'aMdity 0 fOLli' :r,,.:.'>til~lS,, ..'c'repc"!itoo D'hc ~;;;\~rilill\,'nlin lIfI~YHd~rolli~d y~st (ligS2ll. Q;bt;,"1·'!;,'~j)g "",;!!~:I.-t"XWuc,liU.c:d ~lH ,",:{)l\Sb,i~lll~y lat>h:f
p:JI:mIlil~leffi
mll:lil~e Flllg-[l3~10rc .ilLlly pmlC:~1l eau be dQ1CClCd by Wc.~cm b1ot. We thcil:,'rme MlulioJl ;lg~ilnst Jliteml imc~rclml(!11 ofllltm.CfV'CL !:<lIeS, b"E!l~SC ptM"<lIli,ctlo.'lr dlUiC{:;s~C',g" FkL~ =:1B dC.g,IflI.
p,robcs,
fihr ml~)'llI%
or lW11,O'I'Ci';
Dcspltclilc
,n
14.06·
VOL
31 5
SICII~NCI!
.www.sr.ien.~mag.org
REPORTS
fiig.3. 500\71 h(~ttJfl,e' fep!ail:OOefil ()\!!~ prol.i!mJ.· '~Clding !ijerJ!l'S. (Al' ,til'miilrl hllH\Illwr rate5 f,llr genomic
,~n!i!QI~:ti1(i!f!lS' (f~
M1iI
aOlI,:!;
wh(l~zg~(iI1fl~
dl;i~id,(B); IPr~
~~~1 dk~FiIJ:lil~ans of h~r'iserltM!dI Peg~OOiSClDmparw \ViiWil the E!lliliroedata set. Jil axis (lOSJaritihlll1llc SGl,~')
lurffilOw~ fa/II!!. Yaxis; ~h!l)\ll1 fira:clioliil nlp!llD'bes WiiWilill eech ralle m~. ('0 Ftl5.'I. 'codiFig regi~m and MiID(]ated l1!!Kleow:me5, (01.01 ,rociI.ed a(cortl'lngl kI turnover r.;lI[t"5 'i"roml hi.gll-re$oluiti'Onl miHnawray ,~Pt?rlmJ~B on l!n~(hr'llnt~d yeast oultMre: (UlP), <uld G:l-a riI'e~~d '~I~wre$ '~lOOti~, ~[I')Sr.;<llller Ipl~1 ,of
S!1:0M
~sflicol'lNA
~
,
(I
I-Ii
1-
._
'M'I '(1..1
Gl!lnomic al'l;J"~oIBliicn
Turnover
il'ilW
O,~r--------------...,
!f.
.
,ii,
'i'
+.,J>h ........
PQI H enr'~ment ~mi:cl~CD$) l1Cr!JareHremI'm~ ~F~g.3m. Ui's iN RNA 1)[llyn~I.'!l.·a.w p;l~!llC' C'\i'ic~~ in;g m.u;klPMll!l1~",Ul ,\'.Om!) C':"'!'O.1I<,;dlilollgh m,m}, h~£~nl)'Iroj~'C:l'~bcd @CnC'S (ftPi.37lJ. ror C'.x3nJl,lcl wi{hhislonc
tl!llHOVCr rates,
pfll)'lll.c!:;:h<;''1;,': ~')ticl~on rep:r(:oStIlQ.
Q.~m;;.i~~>;Jl~ wi!h
,A
CIlilibii!lo,\\'
Ilm~L!C :[!ndII~~ IU)iul~~ .1 ~d;Lu:iij,bl~Ilil'Sll. S'I,cr :I'h'.fcooinlg n(,,;!:!;iollE!i:snlme [lmID'\ er, I.ht:y ,~re 1il1li1:i'kcJ:y 1.0 aeccam ~;;)r ~hc hull\: of i~ iSlrlJ1C I'Cplocemenl (F~~. 3Al P'ltclm01ClS, of h.ol.codJnrrl.~ l\l~I](DWI!llh
ill!;!: h~:;;n(m(;
the converse is HOl. \~ I!I"C mJljmx.'111. cold In oodfn£1 qiOllS It\g.. F~.I IJ), Morcover, 1\.'I'l'lacemr..1In rates ,m premoters ~~'C~I:.mnHkc'lilo(i;c '
l'C'giOil5i,
t~mlw be hot.
btlt
m cooing L'Cguons. poorly ,oofl'~.ImoowltI1po~y~ Illl.T3ScabundrutCoe" ehher ..,t i~lCrJlOI'I'n(I'DC'r.' or moe-I:" mite cooin.s rcgto:1fI O'ig. SB)), makhl8, i~ ll1llil\;cly II1:u promoter fturHOVer is oolcl>' a l'Cst~l.Io(}rpo~y-
'hou~~:i.umcl ",I'
TATA
4. !l1Il.!!p~dltl!mo1i'er ,,~ Pr1HflOt:e.rS is"s:secii'lt~,ti Willi!! 'f'I1I!!!~Upl@ parlJialIy ~lI!!dollPIJJng f~~l!Ii'@'~. 00 m.oters \w re teJJted lor siJ~nHi(~IIlH~ '~Flriehad (p<lO") anillotatio as.. C~JUr;Uer r.allJllmlWlS !diag ~,(lt pmJrune~a:'l rQ\\I'5. anlrlota,tions (table, 56 ,lind rig. :511) as (n~uml11!l, BLa,crc bars ]l'Idkate ~ositive ,Q1i1lnl!!)'tta'l'iQfilS;r(l~ g givenl Irm)liliIot@r. {B ~o ID) Q!.iertap b@t~~INl ho:~ prOrrrleters ,~mdl pail'S ,of en rkn>ecl ,ilIl'Il'Ilo1:all0IilS:. P va lae ShiOws~igl'l~Thl(i!lIlI,,,e 'of oved .. P' betJ.'IIeen pili rs olr .aJl'Il'IIo'talio ~S. 'g~'oIIel\1'I l~,e 'eKt,el'lt ,C!f Ib'ek ~vedap ~\ii~hh~t PNifl'lOL'e'r:s (hyp'er~oometfk dl,S~Tibl.!i'~i():nt:5AGA~d~mii'lart.ed 'genes alr~ 'e-m!i(h~ for 'liAfA -<Q~U!lifl~l!1g IP:rtlflfloliers (8) C! 1iId1~re-m:Qd~~"te,ly (aw;::I,!M;ed wi th :lH;lQl! nil!! 'g'eFlliis (0, \'ihe rMS promote r~ w~th RtllpJl soit·e5a re nOll Iiill'lricill ed L!i psb'e'ilI rill el glen~ el'ih]D.ili 1'19 high IPoo~ II~ ~@\i~ls:Q!'! eur e~~~nm@n~ (D~.
Filg.
I~ot
III'S
The IU:mc'S.l ptonHOIcer:o Jm~IUid~ dtGISC c[ulJying bJn.:!il1!;; $il~ ror a, :>tul,,'>'C.t of Irun'>c;l;lIPliol1l WaelOl'S :;uc~ as Rapl. IlcbL OC~l.:ik lind Adr~). those !I11}Slrcm1Jlof gCflCS, iieg~lblcd by ,hool'l1adfliIm)(Ii.l~~LtilJ!'ooompk~xCt'l !,,;:.g,.S~n,f,,(Ti:rpl. M~t1Ltl.lor. SAGA. Swi,tSnf.f11l'1d S:iir).3I1d lbo::;~ upstream of g.C't;1e5,1ISSociMOO wilh nuclear pore
oo.mptJi~lCiHS
c'S~
Nup:!). {.~IiJIS~Cl'iUlS i1O,t rl.'O .. m~ers based on enriched aJ!n~oliI11011S yielded ~rnie.rel1clcn! dLJ_5tCl'S Wig. ,!lAm~d I.ig. S; II}, ~\!Ch ~I:'i a group {~fhot 'pmm:G1\;:~, ;l'~ ... m;:i"l.lJd w~l:h nuck:;;u, ipotoe C'OiJ!lPO~
rk:illl\, (2l)'1. These 5t,,,~r;t!c dH:;t~rn~ugg~$!!I~l
mhC'
nri!lnly
cnr.idulil~I]L~idCI'iL~ficdlr[jh:llli3Hy
REPORTS
!.'lUy !,1I,,~r:iJ<lPllll,g rneeha~i:.ms Ih:rmridpr1(lm~~cr ttmlO'''Cr W~,I;!i. 4. :S t\l. N:tlc~l!lnl~t~ple,.p;lrl
[)t SOP1!l~ ~'lf~j:m~~!1~S; -.;t!;g~"1't
d",wh)110'~~1,.""
o. h ,(mid. CalSI[!N',. by cm5'1~ Qr r.roInO~C'f ,dlfomminln .. rks, tlwtl ffiI!1S~rJ pt~.Qll;l~ ~i~init""I~(Il1 (li;~III~ D:r!!j' .~r! the o('l\ll~i~!.. i!\!!1: pn;,.~11,l1;;~ 1)1:' :,In
3';;:I.i\'~II.ill·lil :Slimuhl:ii.
15. ~mK
liI_b~~~,h~ IlIiI¢h~u~~:~n~lor ~~Ip~d !t1nlO\'\.""f t,~,g" rnpid h:i~l\ci.irI!.:" R."P.ll1.c~nIil";:1J1 at SWif~!l~.:'n;-gu buiL-d IHO!lIKltiJ~ rmwwdl be". COn~UCJlOe orSwi :nf .1.~lklJ~.}. whc{~..!'i 'l:~h:t,;r cli~;id:uUCI1I!1; .Uj~ I!;;~
iItiJ}{]rl:.1n,.
[§.
H. 113.
iII~1'l11!lli.t:illg
ojj:t liLlL::k.:iiI'rOR';,I.;<!);
i1Tlg.~Itz ~• tit NA getle:.. ~~li.ctCill"jX)i"C asseei r~~iM. , and Rat) ~ flntl Reb I sites) ;IIif'C agsocjal~d w.i.tlil lcHlull(,1.:u1:fS Ihat block. hctoi:loocill'Ontmlilii ~1~renJi~~ in )"Ca~1 11 1<1). Iilow 00 l1oO!.mdrn·iC's blbck I.uewl :-;p're,~d~ll~. "25) or chrom~ni~~ stntes? SU~"'Cll!lCd !l1(.'tI'!li.n:~-;::In." i!1c~ud!(!long :llJ;'1Pi'> be~WL,;\'~!l lluck'O."C'lnl~S, or 1UC'n.li·~I,.:"d .,!t~lyl~ ltiml C(llllfx'tL.l \'\I~~'1 slF¥n;,'fIdi~t'I~d~!.."i~~1\y.I!H![iGrrl t?6,. 27h The ·L'llflidI13 rep:I"CC'I1I~m 31 l'lQmld<ll')'~<l... <>;ocim~>d rugion~ ~l!ogg... ,'l:U~1 O1,itC'mmiw;i hyp('llIWl5,i:s: ttim~ ~
COnSI1~!U !(\.I~I>LCl!n!(·!1lof HlICklO!Wn1!;','l SL,.~"''Cl'ro,
1".aJlionstmUc:s of h istane n~lIC[b;; c,m!11dbe clln· kJ.1,undoo .~ di~l,Ilion \villi hiS~GUlli;~:;myin~ ~hc c iiv(,~I;JI;!ilCItior.1ihC;i'l~nli k .... of tl1~ .~mL,:'h~h.l:ule 'Cn". rool, l1iimkilig lIY,iitLini'ii..: at gA.".i:1di.: ~~~ldiC;ii k.,;y I.{.I d"~ciil.Iil~!".i'I'g~nl'ill:'lj"I~t'li\,,t: role::. ~.lf hi:;tn~lc l1'hl1r!-:s in m,'lfl~()riif!~iol,o[1 cM~rol. Flall9~:I~.rn~icl
tuml1m,~r·ocel:lrs ruI.chmI11lll.tin boundaries
[£I.,'C 3to;u
n.
~lJl
2:1.
IUJDlWe'Jr.lte-:; c.ao tile 'iiil'i\~d <it Ibe.tUfl~er:l'ily til C~lil!)'1l1<1,~Il'!!! cr~t GenQrn~~r'~[ an 5. (~I'l'~~ h!ll1l~f(.ompbic"(SJhwji..liLl1ItiJ"flOn!' 8. tGuim!l11!I!ll:e .fl.I'lI., Plo.S B'ML :~, 1"3M j2Jl11]:5J1. O. t(. ~QlJll; Ji\l17l., (df :J!2Z, 51.7 ~<l(f!OO). ft. M.. Ral~n~ l't ~Jt. (d] U:::i, 2:33 12:OO~~. K. .A11ImliId. s, Hle!'!i~_dl,. {I'I!x; N~ Awl!, S,f. U,S.,oll 9'. (oSl!!wWl 4), :lI64111 ~200'~), ].. M. C:i§Dlliri J!!'.r· ~L. (.t!IJ 11], 421 f.i!OO~). 0, Oon~~,c, It MiIIl1~, I. llirnl".,. Ii:. T. lWIlilk.a~. ·GeJ'lt!':!i
~; n, 6-'8 U:9?OO .
.22. K.1~liIil, G. Jilrib, C. ~jrn, G.V.!lA IilroJi!~ 'Ill. K. Lrillmrnll, Cdj lO;9. :151 'CilOO2)' H- M_ 'I}_ ll'IIlri1egh.im~ .M. Wu, 1lI-I!)_IIl<!dl1an1, Ce/ll:l!.!l, l;!S 2~. Q. 'i'u fl.cl .• N"~dfk Acids llifs. ::11. 12'JIl· ~OOa:~,. 25, L n. lI!_uS(h~. tk L lKii~ffI1~i.i!r, J', lljfi~, 5.iocmm, f~" 'lSI f2j)O]~. :2 6. :It ii, J!, :R. &roa(~1 (w. OprJ!. Ge!1~; Dw-i!:.!., 199
(2~11."Ve IIOOIHJ:';!c IiIliitt CrnS1.1re· of hb.tonc maeks ~or assooi<1JtcdI~mtei~\;~i yrnpid tumover delinl1b il<;. lite slj'rend 01" ctllmnk"l,tin )iiJ.alcs. \ll,rc runrtilcr S:1~I.'(ul~.nJ.: lhe wid~SJ'Ire.itd I~i~~o:nl!lilln¥wcr at Ih,u
{2oo,n
A:imt.i,~".
r:rn~l~~I.,;'~l.tQ!ilJlI~hol!l!l
!l'~a"'~~ltome (lOlu!d ~r'!J.~, .in ;1 ~~I!.~¢. to '\:",:"1'1<111(1"lh~ .gt·nom~ b)' !~re~'!.;'111in:g CIWOJlliiJ~in .~lmCl' or adjoccnl gCI1.Cl1. riuun "llt'JC!iUg!;,'1.ch o~hl.'r.
'r~li: cmillp;UC'I. Ehdme~(I!~ anlll iN@1m. ~1I.. Ill. ~. 8r:m!1i!'iin. C. t, UIl., IE. IL H'ilm~:r:.
~.:r.Y. H.
("r'~V(! ,1 I~U~(l[:!lnji !ip1'('.mdi.ngC'hmn~ati!i" dorn;lI~n bi.:fOI1: il spfi.""'"llclc\;; ~LI'l}' 1~l!lhcr j 11~. S ll). To, i:I.~;"o'L~ligmc I.hl!! rllllc: of Irnlzl (whose: ro.i!cln
IiXlunL'lary run.c~iofil is poor].)' IlIl1d:crs~oodJr in his-
~. O. hr~~1~in"
wile repla(~mellt •. we ~m:aS!la.~d FIkl&"":~ inco:r~ 13 p'ornljoll. hl iu: I !JJ. ~m'uml'" t1rnli Dlg !!1(]b::lIIly slowed ~1.3 inro~lX!mti:o:n but bv ~ocus-spo::ifk: C'1'li'CI!>Ml FLml~~rC'.x~-cri!ll\.'illl."willlX'! 1l(.'q~~i'!lfXi ( w wr~'1~lg~~ (i;llS 1"\:'I~rli1."'n~hip ,Illd 10 iC'VahIOlr1C!IW .role of llll)idllll'IJJJvct" ~n cill'Ollmltin. iJ.oomdarks. We have measured lE3 1t.'U]Ia.ccmCJfn1';l1C'S ~hro~§h('ll!l~hc ycolst genornc, 11 ndin,g Ihm 111W!~ dooomllL;S O\"~r oQ{li!~g n:g inns; lIre Nlllm.ccdl t~~
!1ighlmll~ri
5, 1!:62 (2001!~. 2- lH, I;l!!I!g<ir, l- 'Grie5iemtred:, .L~ :5lr~:g~rn.. It D- t:i~lm~1'lJ,. Mol. (tit It 1:537w:JO~). 3~ C. ~. ~, 1(, 5bi~1I~, '8. !':i!01 B. I), Slra~L I. I)'. lij~
t, 1Jfu,!.
:5.
~Inefb:~f. GPlla.rml
Chl'u. (I. 'fwl l~ S"j'jrllI'n~I@r, It B~ Gt-nr!'lle: ]l!lit~, J. 11.15 W)(lJ:). as, Y. Milo ~ o'f" ScN:!r:lIllt 315., '1403 mMln. 2.9. W~ ilia n!J!: K. AhImoad. N. rf<llli~~,~. GillQ!b" NI. MalbUt, 4. .iO}i!f!~1,Jjcl~".it ~1i!~lermi!~·, ,It Il'iilrg-aUt, i!~d t W.apjnS!Ii f(lr o;jti;eo!ll ftt!i!dan~of Ur~ jjj~jjUl~pi. W~ ~ih~J1k'. t::_i1 1 Ibrlti@ !J~n~ll~ !'jlft ,a'I !hi! lRS'I'tl ~fl'~1rI. O.,!.i!:. I~ ~~r!~ nm vaf! b~~,C~rI!er Aw~m;l1ii1l ei~ml~i~iII~ ~i!ffl~!:! 1~~iJIIll!! IIljjmiti~h~ Wjj~~OI!IU! It~it~.1iih1~ mtllM ·WM~1ltIJK!fljedlil1 ,;ralmlo t O.],R., :5-.18 itlll! .... t.I.F,r1OO1' .ili!~ ~iio!i!a,L !~~Iiivile 'Dr 6me~.d 'I'Iedi~~l ·to N.F. !i'oll'l !hI! mildll Sdefl(!~ FoIIi d~ign,. O-J,R, de~~medl~h~ ~rilii~ !ill{l r.tF.IlI. c~rmi!l·fhemr UIJ~ SeB, d!!i1f;l1i'l!!d.. .il!ld M ..(. {'o!rrltd wt,. flDllll d\ro!N!~~!mlmJI1(1lN'ecl~ita~n. T.iI:~ NLr". ~ 'O'JIR iJl~dl~e ~I!!. o..l.R. ~~dl ~.IF, ~~Iii
St:iIMIC!! frog~atn; anlll
ROOU.
from,
!lli~ 1i!~Il1~n
flr:nijB~
Ii'Dt ,~t!I_ ]6,. !J(II!Il {z,oM~, 4. lit 'rMIiI~~, .~f.l'. ~. r~j··1'1 1599 {2:00~I. W 5. IJ. j, 5thIi~m.tr, !P. ~~b!1ir., W. !'!Of.!, ~f,qL Ceil 27'
n,
('NXlo).
6. ,1(. AIiImad\ '5:. I-imil«lff,Nkli. C'f1l 9, 1191 (.100j!~. f. ':t'. Mil!)' ], G.lileli!iI;:OU, 5.. lie'mi!:ofl. rifJt, Gf'JIIe.t 3:7. 1000 (2rOO~). It HI. Ii. ~iiIl'i~n, K5.lrllltib MOl.. (ttll .. 15:1~!: lOtH .24,
U1e liIa~r .
nccms
{20011. 9. A. t:ili!rrjilhlin, ]'. O. S!!o~~trl!ip. tiM80 t 2~, 42)q3 vm~}. 11.0. (. Thiriet~ .. 1. !ililYt:~, (J,en~ Oelif. ·:11.9., ,677 (W05). 11. 11- Iling~r, JllK. Tl'~er. €~. e@ 5\ 1700 120(0), 1!2_ C. L Lfu n oL •. Pll!l:l': BiD!, ~,!~Zll Uoo~, 11]. Go C. Yoom .~i0,1•.,. ScfuJJre 30~. 626, 12005)1 1!A. t.l!a~~lijals .iima !I1~iITI~~5 ai\!\l,ilabl~ O!~. ~~pl!ining .allii ma1!~l11~~on .Sdl'.JiIC't·Onlfflf!·.
F~linr"f)i :t!OIljl·
iincliilicc{I to
prodl!Jt\:
biolill.:ru.IJggcdi ~B_3
rnr
~:smp~li1t~~:f'I~
~t I't(lme~li:c:giei'l:~S: byd:;-11~~~~~ry
f:bom strepiili\ idlin pt.lll~d(!iwn~say rutd. milnliCti!lll.~flbdt\d with Cy3f1;n~d C'y5 d~.
tllpMI.
[urad,
'lo l'Iuil:lea~Ii~-4tYlperse~tive sites, ilarl'I(I.ilii91 ~~"l f·Or Il,,,,lyoomihi aliiid'tni~I1",l'3x mmup pMl~n~,alil!d sil'es .of IiIllIdmsnl'l1le !!:Iepietnol'll. OI,UW rt'S!litls sug:gt's~. the ,t'X!1.s~8'!Il."e al 'l."",nl;inuol!I~ p'.rocoE!~~~hat di~rru~B al1f 1i11!1~~mSlilil1le:s mal nl<li ns am1!~sib;Wily of ci;s~regll~atmy and 'lIre<rn'91nl:s~
over Ihe Cillli~ 3ft chrom,(l)l()I1I,c mnn ~,e\"CliIkdll1ill. Ihe -35i(Jckb bilhOltlX complex (BX-{) regton dlSllluys the lowC'~1 B,3jlS il'mioof .<!!ny reg,tOIl I oomp<u<!hk: ,..iw O'l'! JR. and .he A~lcn mlped:i!l
or
hro~la!ill c,m bo!: dtill:ereni.iJLtecl b\~ I.he , rep~k?l,io.n-ilUl~pelld~ ..n;l. n.ipl.ac. c.m~.nl. or onc h!sh:me Y"flmll~',1~h .mmlllcf l J I. for C''''<lmp~oC'. i~I{lI'nc I B.J is '~Cpt)silr;:d ~hroUoghO~1 h nil; '~I~ cyocll::. I'tp],Ldll!;l1 Unh«ti:. d!cpo~irlJi;.-d d~nr.b'ig [c;plican~ln (2' 4). lJnlik.c rcp~i~3lk!H~ CGuplll.KI :i:;~.I.~I'i'!My r! 13,\\'h.i.tll uec'l,Il}; in ga;p~ o
l~lLcJ,ooso:n1'CS
or 113.3 ~~prc-ccded
~Basi( sdrlflclIl'5 DiYisilm. Ifrf!d Hutdlli~QIIII Qmm Rn@<lHh (en1Jef., F.1iM~ A.~,~n!Je ~ouJi(, SM~I~E',W"1 '9'8:1109.
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USA.~r)l«ui1lr am Geltll!'lar Blol\)g~ ~~r~rn, un~er>ll)i I)~ Wa~h[1Jj9lO'" Sll.. nJ~, ~YA '911:1!:95, U)J!, •.. HOI'l'afd H'lIgh~s
M.~~(.allnMiun.i!, fred t!:ukihi:lIISom (amcli!r .Rf~l!"r(!ii Ceflll!r. ~'alte. wfl, ~:iQI!'i., uSA, '1'0, wllOfl'l coneSll'Olrloonu' ~OOOldl be ,ad'oilressedl, e·tI1a.1i:
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REPORTS
homcg~k gem:: 'mnpJ~ (i\NT.P"'O also ,d:f5P~·<lYS 1'0\",[!3.~,H;1: ratio {f\r;;- IA~ Lo'w !·l13/1 tJ ~ar.o,'l :;J~. !1Iw hO~~ll,l'~!'~~:Jl~ dtJ!I;i!\;"'J11,,In; ~ <'Iltrj1.J,~I,1thk to lllf~W:I~I!l hi,~m~(,':n,:r~a~11..~rn!,l!lli.. ·
WQ \!1Ilu..'>IlI~!!y
~md nltli. to ItDWI~I;Lt:t!i,!OI>Onilcoor"ul},:m~Y, 'b~'Ci~I!i~": U33 !C'\,<fb. allhe B;x~(' ..re j~lr OO~QW !llOO~ml the
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~ll~!!!.h'tly.. bcn\l¢ Ihi; 1~1t,'di1U~ oVI.-nln (.fi!!}. lB~_ Even Ille h~l'I::-:i'ocIiTalflfILi.cC31mllIDi>O!'ilc" (6) il:iehIH;;k" .. ~nly ~hot1~: ,(~~.Oo.kb) s;t[e!ehil~ ..hill me as .1."'plclecl. ~111~ 3.3 Ib the nx-(: ~Fi:!;l.I C~ 1 A dt'iliiC61JJ1 'iI lew of ihe BX-C {uil ~re&tan
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{R~" :!f\~I, Nmnh~y, the ~ICll hiW1CSt pC<lb ODl'~l1d to tile funCI'Ol'llla~ lb{'ll!iI1dmf~ of the seven prmim.,]-to~di~!'i~. ci:-;..~"§u~mo!'y(j(F I'nain~ d'!l!lJ\ reg~I!:,Il:~ aJl(/,A tk.I~'l iuh4l a!l1d thL.~ ~l .d/u{··JJ lit~k~ to j(lMnhO:llt~mic ,gl/,~11J;-li~i!)I~~i;;~~'\,'I)' NUril ;ml:i."l~nr.[0 'IX'Ili1,iC[~orn tiac at~('imcn 17). i .on."pjCl!oW!~.p;;.'aksofW133,,!!i.o C'Orre~'KlI1dw duo: h;~d[I\'l~YOOIl~b f!"N1I01~~ (,;'1Linn;.'ut (PRE! m~d 'to pRlilno~~"INilbiuldlC' ~Jbd-.Bg~·n~,. w w~'1ic~ krID"~~l. i~ [r~Il'I} :)C'[i".'!I:: il1 S1 eetls l8 .. '9~ nll''[l;:fo:rc. eaeh of Ihe m.a'>l. .r,ml1lilrn(;I~1U3.3pc!::iks in t~i~ p region Colf~pollds to II pre\'iou."ly dcfllJed c.i.HeglJlmory dCme~ll.Our filldif1!)? ~n:' ~ikel'yto be W"lI~rn~. ..... IJI. pC<lb:
lI='iig. :11. fh'e 11[1,1(·(s !~~f;l'Isiil!i{:ly d!epll,e-t)oo li'l I!H~,~:.(A~ '-nL~~!{~ klg",ra~l.~' pr©fill: of dllr,@m!)~me ,~mn i 3R. Ori~M~ti(ln is pJQx:iIlTl'i1Il (01 On bCiSC"!P;:'i]F s(.:iILe) 'tQ (jijSloll (;28 Mb).1Ihe IloC.:iJU,ol'l ,(:if ll1'~ two MI,omI;Olia; gM~ ,dl!l~~rs are ]odi(ai~ed wUh h~iri:m!ntal b1'us. (113)IU.3 (red)! ar,)d H3 (bhile~'l~rQmes;ror th~ WH and na!'llki,rI!!I rl!l'~ri!J!1s. (() S:"m~as '~B~lo~ ,~hr,O!mli!slom@ 4. Th~ ~!J(,,,U;OIIlI~IOfl' ,ilflOOliOOdi lr.ll1!i1J ~S(lIfl:5, '~Ibrrowrl~~x.es~illld '~'8'JUl!i (lb!lIClkbQXIe'S) air!! shown OOrl'~ifl~ eaeh pa rI el, 0111 !!li1Ilii!'~ 5'4"0' 3' all:lO~ the ItiDil,eand 3:'·4(1·S;' be~OIO'll., li?w[iILe'5 ,aIT!!di5p1a:~d ill'> mO'liing <llv,er,ages, \llitill ,[I l·klJ (A) er a 2.541,10, ,~lB,and o mrldo~"I';illi lOCNbp flll~ervaJls"
causein blu:ki~alB}'Cai>t, rrnll1oler.; '-I!rI(1 'bmmd.lonies llIre .,doo sites 01" .imcrlsc h~~ollc' rep~[!;OCL'mml (Hh A d,..m~ris! tiC ICatlln.: of OO~~~t.Jllrn...l1:rics b mmll PilE." '11 Ihe 181)(,(' is !i~~ Iht,.')i sp:a!!dlJOXY~ .ri[1ol1.ude:JSc ~.(DNm,c]) hypmcl~~he sites ill :JL ',Mel}, of ocll~1t'I!cs" im:h.Hilng S2 cc:]ls Un. To OC~tciI.'l.~:lin'C'inc' h is~onc l"Cpl,t~"CI'I1C1!!~ pa1.lCfHS ill Ih~ vicinil)' (lr 11!Y~"'fl'lel1i!'i'li""(,: ~i~~,-", li~od we
Ihe ~ntirtt:BX.C !I! 2f11·bp, ~~oDlITion (I]g,. Si)"
2 •. (allf'lSI~~ IilI:lUS i111$t.olf'le! ~epialcl'!~ @'n'~~~ dsr1egl1lLi!~JJlry' dOIlili!i n tbolurlda rlles i!rid DNaslll~ lliIyper:!ien~':t.l~'!l\~ sil.es, \vutiinl Uli~ IS*·c" (,A) flltapan!lil ffili~tone p'r,omes. (lor tlille g'bdolrn~nall a nd rI~l'Iltin[J f@~f(lrils @flhie mt--C; f ncluding fOl!rwe-thIllCl,[lpellil PIR~-bolil,ndarij~$ [mag:ent;a 'b~!Ke$ tu, lJ. 25n.The ISicat~ Qt~p ~l1dka,I~~gle'ilQim~( lQ~iUi~f!(In lit with ,~!!m~, amll IPR~:.IJ~Llmdtarie!'S irndk:atedl 0111 '~h~ lill~ i:i~low'. ThE'li,ve Abd·.B ~romot'e'r!) ar~ n'i,arlked~vith ~'Drizonjil anQw:s. 1913.3 ~F@!l) .ilfld Hl3 (btuei') L~~~ ratio [profi les are display.ed as]!,·kb moving <I,wrages. E!~ldl'td.a~e$ belweeli! .adji3i0e'.nt di5~egu~S1k1ry dcmains ilr!! lndiMl;ed wY~n Ibra![kets, aJbove' tllile lCDgl" ratio plots" (8) IH3.3jH31 leg r.al»:s, '~tmll!'j1r:l) .ar~ sho,\'ff'I h!r 'l~~ Plli[s (lI1'a8lenta) !Hid boundarries (bllle) tQr wh~(h ONa~~hypers~m~t~w sibils h~~e
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S~~·";.tutpp:;gi9n 'oo~.~m! to hightcv~lf. of cQmh ~'(Il!P (fru) pro~eTns. \~ hich have been c!]l:;;(ln\'P;L,<;'~ CJ011i~iPlCl!!mlS p;i:<J!b o :~-!~, ~T!"1ImW in an 52 ,ceU l.~nc ;IiI high ~1t'!tiQn ('~). J U3j Wig, ~EI ~lM S I). 'I !:U enrichment ;,!!.1)~nd<L'f!~ (Fi~. ~!:!) !i!1!:ltcJnwly 1;;'O~~1)(l11.d i)g If ~1!C PRllX""~ tli!;;!~~l:~~p~;;; n!J!c~~om~ .. !~ 1-;,. r~V:p'~I~ i'1~ ~i'~ I~no~ ~tmh'l,I~l~Jc ~o hi~~:r OO~~P<lm;:y, lK't,n~I~~ "'~~1~i:dI31 id'(;Ili~U 11:11; kqlO~li'l~ql~c;:kH.'il!,,'-h}1~1'Cln~ilijw s ik~\Vi[~- ci~ii~A.Q;:~, b~lldill~. th~u we \'\lo~k1 C1'\I~'1l::l !! m>l~i:l;:il10S(l1~1~ !t'~ iic,:,'j.I~'IJh:!O '\'<L;'1i"l; ob~ip!cd tlilr I ]1].)1,11.3 pml~,k~ ill lite [(~ifm. t 1/-13). NWid~{l-;;!.: J)'p!.."~~ir1r;;~.~viiycQJ!l\.~-porldb'I~QC & bot\\i"'1i;."I~ ~~k:; ~}'cGbillldil]g idC!l~ifi~ sites of r·d<l.ivdy O1(lC\,:':1lioibl~)Nj\, [ •uw pc"b uf [13,]:, 111d~"<i.\'"hen \",'!I,) co:mp~Ire(1 Higs. S2 'In~S3) . No.1 .. ~RE~ i~!JicBX~(",In; WOlun<Jtno be SQ .Im!~hci,· 'p[ln~[;:Jl!lUif;l,j)l'Ic~<\1lo illl;lIk;.l of his1133 pmfjl~~"",ith Ih{ll~ for [~~h1U!Oi;.'f-QI!.7~~C s.i.l;'~ ofr~G bim:hng; fot ~i'i;~n1lip!t. 111~U~IR.'T F(j'b-,; '~CltiC .fC:pimC¢f.lbCllt ~ug_w..~r 16iIM c{mli~1~.ujlI,l8 dil>(Elj and :rn~tduo:r-£:.'-"'~"Ci!l'itt. IPS{") r~G :iillit'ilmio:t P(;Ji1-Si:s. d~li\d~~bly bo'IJ~~cl b.y EZ 'M I'S,C rupLUOl'i (I rrrqlli,dlZ:l~Oq:ik"" Cl!:~'i!Uljc:>,ei:..ocglU Irll,ary ~~~It_", ro~ntdl haJ~ l (} ~~l.ti. of PcG~i Iltliiig We ~.n !(,~,.Tli1e raet that ilI.U PeG :.it~ ilii.!pcalk~ oJ hi:;,.. DNA !:c:illli\!c[o surrol,lli'u.linS regiom;., ill lbc fIMOn,~ii1la:l1regPOUl 11i't' also lo~1 p~"!iks of ~pla(:(,lmenl.. not \'~.v.c~. :>liIW-c:>l.. :PRE.l»iii'idtl~l' elci~1enll!\ in t~1I!..: BX-C lIi1lL'i 113.3 ~Fi!. 3A and l3!b1c SI), Ukewisc aI. the 'IO-~'i.'i: Ilull, h.ili>tfllte rep~~"'Olltcnt at PRE;,; mud IXlliln.darie~ t\.NT,P..c., IIH Ll 1~<1k... f r~G binding in (tiC o ('Inherre~ifmsm\! bindii1l~ sites for Ilnl~tiip~e .Po,ly" i~ C'Cill~ i~lIIlj\, eaed i~ldC'lOCnd'cnt o:U'the- C'xl~,U!::s~t{l11i or d11.: homcolflC ~C'~no> ~hal thC')" re~IiJ!'lmC'•. For. C'(a~nplJc. ,U~t-EJ is c:-!l~rcs.~d <II. highlc\'cls in S2 ~"(Ilb. ...1<1 di~rl!aJ'l'> [h~ typical ! [13 S' l)C;1k ~hran ,tClive' ~~!~. ~F~!:, 2J\ J'. \,VhL;TlI.\,.'Lj Uh\';;LlIld ti~lI,i·A <In\! .~!L.'.l.r.ly il~eli\I(~ lN, '9), Y~'l ~hL;"PRE.<!
each
"'1"
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or
mil
aad LO!llllnd,u:i~ r~ulmin,g all llu-e~ g~l1Cl'l ,m.: :.[11;'$ or oon~~i~lIOIJ!~ ilil'lll'llC' re,pl,~CCH1.l.·Ht 0'\'1.:. a !r~.v ~(:k:!ilwl,I:m;L
\1!.h;. ,l!MJ c~;lrf:!iI1~-dhi~.rnll.t rcpl~,c~I~~rH. <w~r::lg~'t! ever the 17~ gC~Ii;,'rnt;wDdc.;.EZ+PSC pc."Lks Clu:t£idc or the: BX~' and ANTP-C (Iabl:!.: 'S II lll":ld ebscrved ill1 :m.J peril: (;Cm.cl'C'id.over Ih~ PeG Illrt:XdmUi1~ 'tFilg, 3,c' and ~:i:g.~)" Thcrefbre, Ihe :Slrol1~ msseelatlen ~ctwe~ul rw IlN)tein, bind illS !lnd histone re,placcrncm, is wn Iimilca
f[)
honlcol.~C
!~lk
i!> h~.f,lh!Jrhun Ihm lon~l~ I BX~' ~~~d J.\N1P:,.{'.pl'C;~mlf:lhl}' because GIller PotG-~jjdrl!!'. IB,)
~iws Jitl1C ~1t\1. Slll)cxinl!lmcd ow::r :S!Jd~deep U3J! 'I/,lIkys 'I fii,g, SS),
The (lo!OClJll~Uion (If ft'G·hindinlg ~i1).~und 10(:t1!1 p~,l&!i of n3_3~iII,~':\[~ .hilt 'th..: Ill'OO~"":.i Ilmi. dii'Ol;upllS Ii:llck'o:\£nn~~<~ ~~..c!II~.}1 ili]l~ijUI1~ Ihe m ru;cl'.,'ii~:ilJ,ili't)1 Ci5-.ncg,LllllImry DNA to PeG pooof teias, I r SQ" Ihcll~lhcrc should bea lower fuv,;!ruge ooollpmcy or aueleosomcs over shes or IPet] prolein. bi~ldin.g limn over Iheir stl:rrmmililng re~mlS (8. J·n. Tetesr rhis po~ib,ilil~.\"'C' hybridi:GO:! nltcl'co.-.rofllOl~ Di'>lA and Iragmcntcd gcnOlnic
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Fr'iig. 3. SiUe5 ef PdS ~fn,din91 mrre'!lJXln:d ml.o(.al p~~ o'FhiisNmeIrepla!l:!!!mI1Hiltal'tri r,edm:eirl nudes~!lme H,3.] k!9·rtattCJ IP~l)me U)' EZ i!lli!d PS:C prorh~ (8]'at (A) th~ BX-( ~rom, Fig. 2:#1)and '(IB)~heo A!NlrP·C. Loc:atiom ,(If prolli1in.e~t EZ end IPse Pf!i!lks are rna~ed \lIiilil 1;'entkal dG~tecl ~il1~1!. mbijta~yKatil'lg W(l~ lI$ed 'telradlll"le vI!sl)Ial (Om!p~rt5(lml betJ,~en IH33Ml3 log rail1:s$ a rnd Ill"iear lEi A end IPSt pr~f~i1e~>i!!\ctllJ!a~ '~.1!~. ·aJ,~s,h'olo'!'!f! ]111"90 55. (I IU.l ~rfat1o J!)rof'ii~s<tign'llid aro!i!lfld E+!PS( a peaiks for~h~ BX·C and AWJiP.-< regkln~(4otted lil!1~~aad f'(Pf the I"mImmaer of Ih~ gIllilQililiUil (m1.id I~ili~), :shO""If)~1 i)~~~1 aVfHi!~m: Il5iftlgl g !HJO{iiP'\JMdo~'\I, (Di)Sam~ g~ (0, e)OO~t sh{ri.'flngl tll~i'llo\tfi'i!lj ~r,age~ m rlOr Il!lI&i(lS.'lOmaligeli(!olfnnc IDNA. IJJg-ratio pmti~;
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ilWld
Filg.4. Bindin95b~ for tn;G, IprOlte~f!s, a midi po~sedl prometers iillf'el ",)~c;i(l,t:ed with ICOIll'SP~C:uou$ hi slone n~ ptacemen:t ,<ilnldl ~edu:ooo !11;,u::k'~·some'O(WPr~!i!(Y, til) A\I'I!;!f,alg~ HI3:.3JiH.3 <lInd 1l'IllJdeo~orne ~({up.an(y ~09-.r,~lio p;~oi~:~~!iigl'led at 390 Z~t!t!· a ~~ndlirilgl iii~s. (IB~'A'!Ii~fag@ H3.3 (md) i!!'QdI tl!3, (blue) ~.ogl raH:lDs at lIInindl)l(ed heat :shl'Jck g~iI!~~ [J'Ot~dI g~gyijjFIiIl!S gro h~st~nll: pr:Qmles (or aU (llnl1llt~l~dl gene~ 0'i1 .3iR; linmm 11!1 de(rea5,1I1gintelllsily trnrn tJhe1: mm~ ,~clt~-e('~QP~O%)t~lthe ~.~iSt a<:tlJve< (boU.olil1l 20%) gen'~ ~el:s. VerysimiLa r tII]L 3 pr,(lIme~ Wt'f-e ,L')b![lriltmil'1l t"'~~ dlir,· rle;I'JeFlt ~~pe.~UIi!eli!.lS SlLGt '(rig,
11.
1,
[)~Aw<liS c~e""rly dep:l!:tccl Oil a .... m_gc " Pig. 3D e table s, )1urnilm" m ~h>C'dcplh::lk'lll seen f{1l: !:IC'ti\'c g\;'l~PWH10:!l;.''f8 '~1)r.g. Sl ~ t essen!i;lll), !he ~Iru~' A;.'l'H,dU5,'1Irl,.'].~ oh"li n!~ir.l'!IVi~h d ij!.
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PRES!luJ boundar.iL......are IKltmd by lh~ ir~G protein.,>. Zi.... 9m!. G/\.GA ffi:lcto:r ,'.ie I(Gi\f ~ (l5}. T ('I h;ost Ihcptl:iiif<ilhilil.y th~lI. h~,>~o-m: It:pla.ccH1.Cn1 i~Cl1hanccd and IJlock:il..wmiC' occuIfKll'IIcyilS redoccd where ZC'S4!1; prolcin jll,L~relil!nIblly bind:s.. We' aHglloo 39U Za;.te~bindin.s. sties idcllfif'lotl by hlgh-oosolllliO'Il chmrn;li~in inul11l1l,,"" plii;c.~,Pil!I .ion (Chlr~ I;:omhin~d wi~h!iling mi'Cm~ • O!II11i)'~ Oi~P'-Chip pmm[ri~l U6'l ,IIlJ !lv>t,;it,:lgQd If.I;£ 11~lio~,of rn 13.3.1IB find 11L1ch':>lJMJ.:i:illi: OCnli-
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.www.sr.ien.-emag.org
REPORTS
f';>"!Itc}" \"!c 900CI"I'ccl " prosnincllt m",,,jmunl ~~!~nc ~''P!OJ;CC!OOl~ anda ~h..rp !1fiIluimUlH(lfml!~ C""V-'i('llIl~ O;;:C!:!l~~.Y ,~t~'d P\'~r '~h,;,: 'pt;l!~rn
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!IN!~-hiI1~~il'l,g ~I:t-I;! c;:h.rrlall1l:j.1.1; ..hin~~.ll:!il prG~~ I~ imoh·~d il] ~<m,,:n.1J:~ll::ni.oll di~l)hl}r :;hori I\,~idclt~c l~ltlC~, ami DNA. P 9i. !Ind. smn.l~IUilO~i~ ~rm~:o;cri:I~lipH l1!ctoR' S!!U\"" ,clYIm~nic ~"M;\~1avirnat ~h\:ir r~n~liG~'~! b~rfiding !ililc~ (lO. 111. A 1l1P!;ld lot Ithi~. pt~~11 bus been p~t>"ld •.~n\1"{1~viP1M I.• ~ ~clllimiJ\~ GyC les of "lueJt~ .. ~ d~mlr~im] by 3. Ba.'.1.hIltl!l·IX:]III~d SWIJSNIF c.l1:roi~'milli"'i\.,'ijnOO~~cr 8i1.J tmn.'>'Ct'iptioi1 fOCElI.lr bhi;ldii1.!il ,~1/), bii1.d· bl1i~ ()W' ,PeG and u;.;:G l)lfolcin~ is ~1'>O dYi1illnic (21., 1.:H" .!IIIl~ \,'C' 1J'Iropo.~llml a simiiw: eycle of nucleescme di"rul~tim~and ractor 'bi ndhlg mkcs place at bmiu~Jl:niC!l!!nd ~R.E.'i.N!Jd~mc ,dll>" n~I]I:ii;~ny S'lJV[/SNF .000m:!Ckling OOH1JlJex,~ wmlkJ b oct"~iiirn);d!y (lviCl !1l.lld('(l'>O!He:. l..?4) .mcl !:rJJ1:;,icfJ(~Y e:;,;:~~ DNA \,~hLd~ ~~tllddb(;~"O~!'tC ilV".Iil,lblc III OIhcr di m,l!~ii.lJ;,: locltml\, il1.dW!cling IPtiO l11'01""in. s, The cemieued toc,;aj ']~'"l'I..'HOC 01' nl!:cb~ :;tIll'!ClUI]md~km\, ,~((:lcldDl.lr;lQ~[ in ,mon.h(lf cyck! of ~t'I'!Iluddiillg. 1l\!Cl'l:'i\f:ml~ (~I)k~ni:nll. 1!lI:d~,><; hypctsL'I'l'iuti'l.'hy. :md hi.'>IruIll rcpl~rol'Jj:JU III the !;]I£':" Thlilo l1~odc~ coold aOOQ1,JInil.r(l['" 1111(; di\'CIi'Siny ofm.,G
~. 8. E. S!:h.w'lIu. t:. Ahmad.. ,tifi'lf'r.!!IfIlI: 19, 9104 (:tOOS:~• .41. (_ Wir~iJIJe r; O. lIleil~ 1)\ S[hu~E!r. ~ ~,J:~'.
!\IiI!g§L.?'Sn:. IIi<ll nucleeseme di5.iI!.lp.~ i!l: ~'i!l;:ml fi.;<'I!.w~ of IlrXGpmhil1 DNAa inn bil:!din~~il.GS. H~lJ ~l]vidi~i1tTit~t~I\;G. !ll1d IrXCi
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pruEeilj,-~jlwiljg ~uK.";s,l'L~ill" Im.il'i a ~p.~tc3Lit)I~~ i:j'ltlerh'~'ldeil~ I\.'I]Jlaet'tIK":ii11 l)i'O!."l~,OOe2I1l~~c essenI i~lIy iJknuJ.(%i1f\m~.~:; v...-:r.:c ,obli1iu~oo far:~ 1J.3~n;. whidl~ 1<Ild.ki;line N.tcI11~1nm] mll II mtd dcc.~ ItOI m....'iC~nbJk: d1.U'i~~!J. I'C,HC<lllm~ m£. Stj l, like HI!!-?end M"h~. ileal, :.hook ~cnc pill"" ~lCitCI'S me fIl'Onl!ill~ll( :>i!!O:l1 of GAF b~nclll';' na-
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:it.7611t200~.. S.IIL ]"rlid:l ~rJl •• Cel! '116; 1l!3 {2004), f..L. S!U~d u.!, 'Wi /Ji"t. ""'. 8\2:10 ~'OO4l'. ft. tt MMda\ I~.~~ tJiNllDpmfnt' U~., :lilll3 l:1!lliI)jl). Y. It SdtY,\il..1Iti!: u,!.. MU. ,('iemt<t:lB, 700 (iltOO6). er A, 6f.~~i"!l, (I.. M.. T~rm;;:r, ~ IE:.~ 11!;h!', ~:. ~~~d'9, NtlJrJrt 412, ,6'H. I'll!. f. 11)[00., T.. ~!lIilO" 11'.1. tlim,S. II!.:lfdt'!Jll'lih 0, ~. lii,mdcl ~. linied!lia!'l. Sr~JJ,e 315. 1WJ5 f2007~'. F...~iIf(11i tf ~~,l'lr:.fd~Ir:llritif ,1m;. Z2', 3:138 U99'.:!}. 4. ~ ~'Sainp !If qi, M"t C~t2:4, n. (iilOO·6.t $,. ~rg~ I!I ol. Vl;!ve.top.~r U~', 779 R:ooot B.. ~iI~P. j. M~II~r, 6~~ fJt!;j; OliO, 2.o'l'1I. (2006).. H. W.• ad;, C. Ii. fi~llief, Dw., D,n. :232~ '~3 ~2(05)'. ~ lit .M!!!~ i![ ~L. RloS r~~f. BML l, n30 j2.1lO6;}. I. A. ltib~l!iltdll fr.· 01., Mol .. ("~n. Bioi. 2~. 6148
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c·!¢t!81.l llyp<:I"$C'n~lti\'ilj' ... nd Itld.ured nll!~~~IH~ CiOOIJI>W1Cy U n l1~·.<Il '<lh:o!.1:.pr>tJ;tcilll ~~~iij !;lI1.'1)!.\'O H ~!'L.~ OQJ.ls:l.iu.lti'If"l.):I)' "poi~t;Id.'! for I,up:id IiI1dlJe~io'!l. but do not pmdllloodcl.Cl:'l1blc fnRNA~ in mh:c 1In~H'dl!eM ~'mm~,'Wc a~i~n~'>d !~S.fli'O genes H~ ~hL.~ir ' ~".\,!ui" nd "~>L;ri~I.;'!(.I mJ. al'll3 !U Ilrof1I~($. 5 a
F (If Mmr1urioon •. \W fl\lcrugl.!{iI sinilill.:ldy :il1~gl1L;,j IB.J ::md raJ p:r,o£i~l~filii :ItI ::H65gcnt:s Oil 311 Wilh );;1]0'"""111 5'aiJId .3' '(;nltiS., I.Hvitioo iuto q 1lI~ll~i les ib<2ISOOon eXIJmession Ile\!e~s.. 1m3.3pml.C'm~ were
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Sl., (2600. 19. R. D. F'h<lir6 m.w Mtll: Ct!1.r. 8&1. 24, ~93 {ilXDOO. .OlDI 0.. ~i~ 1.\[ ~I:, t.Mi!lO J. ~, 198· {ZOOO). 21. It ft Nlilgakb.. D. A. Wa1i:N, It WOlI~f4. G. L. Hage~, Mal: (t'H ~4,1l6:3 ~~,L n ].. Pllah~~li.,¥l 1(. Csiill'k, A, O1tin'.!itril~,. s, HMIl;~lt S J. C~I 81,,1: :tit 0, 129'r U9~m .. 23. G. fia, R. Heilllml~!l"" D.. I, AnmdHlWin. I)Ml~p"IJ'i!M.I
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iZulilc lind GAF). ~.I:ctOO5iOJ1l!.: n:'1I1:oodcrs tBm~!I1nll anti Kiisn!1!ll,l,and~lis~OI1.C' nmethy:I~T3ns:md fig. S 1. O~ W.it~1 h ii:>tOll.erel)lacemell~, levels rClll.'lCl'> (!r~hornLxalld A.,hI) tl~n ;wc ~pcdtk Ibr p~kh~g Oil ci~h~ ,~idc or h~~ shock pmlll'OI.it..'fS. J'\fo d(l~r;II1."lC~p~i()naU~1 aclive gl.'.IlC !ll:umlO1"I,.\U!' 1131<4, a ~lNG!iCJllml dw[ i~h~ghl" cru~ched (In I D.3. lh!i;lm-;.~!l~i~~g g~·~bi.!~nic p~ \\IIl'l\!klallow L?).,hw~ ::;hock gl;la~ di ~)Iu)' p'-Ul!nV[I!;;'rn m..3 and 1]3 dips, ill abundance Ihat arc ,ru:lribuUlbk: 00 r~ Ilf.'OlcirlS IIJ~I. plll1l11Citt 0P-P{lIsutc CI)i~Cllctii:' '0I..llJJ~!. ai, OOlllUIJO!t Ci.<\-llCgll!:IiL!tm),;~il.C':il> p.ilI.fit~~ I1t!Ctco.~ntc dcpk:~on (17). O::m~:i~u1i¥,c eoncsto II iSlm!C' repl~.ccmcll[iJ.!so ;j]~iX'.a.l'"$ 10 be a ~C<!UJJ:c Rerl!rei!lce~ :and:INotes or 1]!{)is(.'Jcl IIr"{:Illl,O'l.f.:(f.~ in veI1ll."bml)~""'" bcta~sc 'J!... S, !iIIlnl~f,l!(, .A~Ii[I~d~ A!;irltr .. R~ WI ~Jo:. 8J'frlj!1, 'l!33 11L.\ is; !'Ilmngljl cnrit;hcd in t~l.\i' llpS!n.~ll:nl"'l:,.~iOl:~ ijIIltlO). of llrl~~chioki;n 101111(:rL-OC~OIf gell!;, !lc~r~ruk'Siiof 2. !!. ~~o.l 1f{eI!~krj~I,5. "'e~lkgU, lII~t'. 6en~. 37. :~090 (200Il). Wh(~lhcl 1il'iL" gelle is nctirc or hUI.CI.lI\;'C {an
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rrlor~i TQrrl!!fI-l1t,o~e~l.l·Jt m1ilCOIlinODe' lfIi.g:coLi.l~. V'ioh!~iI ,wrdon-P'redildo. 1, S<lrail farmelr.~' Adlall1l ]1i1.lrmlll,z.,11r;eLix M .. (hil1l~·l+ IllIhililPpe P'II:!ii@ro,.z Midi1!aellffisby.~ ~ilI!Imf'5 IE. t:l13~,er,,~' 1~l!lli,!'i Ar.3gdflll!~ Celll!l'lcilrcheclkpoiiIiIIs. p:r,eve Iil~ mittl!5i5 nil [h,e pr~selil ce ,of £~:a~I.ed repUGltfQE1I fo:rk£.Whe'tn,e.r ,clheckplllll1ts.also emurelhe ulm;p[etiol"l ,of OtIJ1.Arepti:cilti:(1nI !:Ie-fore rnili05~1i h. IlInkn'DIO'iiIl.Hen!'. we '!OholO'!l thii:lt:in ~l.!I'IS'~ .5"1i'K5·sJnc6i mLl~~n't;, W~g[ht .me rel~too to (iohesinlllld wlil~:hmsil'l., .rel]l~.ici!Jtnoflis dle'laY'ed~ JiIlosl slgn~'!karildy at oMural r,~pllk.atioil1-impcdlillg I@d Uke the riboS<OrlIIaillDNA :Qene
S 1)11a."l£. TIle 3],gmm:nls SLlI~l!On:in.g lhc C'X.iSi.CIIlCl!o:r I) 1l;;plk:alimH(Un:plclkm ch~ck"Q.~nI. dc~~vc from ()l:ll>t'1'\'3lif.lJ~~denlOtlSlluting ilim lu!ddin~ yC1!.'51 o;.l!JI; <lei i"'''I1: a :1;\:\'(!rfo.,boJc chctk~](!!int W"1:!cv~ c~~1; b ~n.:·IIfC<.lcdwith th~ dnu~ hyd!l~i'i.j!l.m;,·~.(UU) (2). 1](]Wllwr,.~i~c· chI,.1~t.poinl. .i\.'};;po~!~o;;: H)I ~.U i!.ii (<;LLlSI;."\l ~y L&a~at"Cu rnwdiltio:n of ~ill@:r..""I>!~.·~Ll.u;:k\d DNA 0111 rcplicoilliolll rod:~n:l'nhc.r 'D'~I:ln IImx,l~IklllCdi DNA. {J .1~[Dld:ecd, undil"!.,~ c...·icl~nc~ ~rom S\::~~ :'Iutlics )lJLJgf~l:; ~hfLt. yea!!1 cd Is I'Il~g1n lad" 1) rqplkatro:llD~\1mp~Clh)l1 cltcckpo,inl
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~f Sm(!5~Smi(6" (hr~!if!05;~rnl'l' ir!Q!ldiiSJ~nlcl"i(l!il QOCtlr:5,a:s oil (OnS,!l'qILlIl:!l(E! olr ~H(llic '~rlny 'o'i'ithll~ril1ish~d .f~p~iI::aili()rI e-spiOO'intact ,chockpo~ilt ~:pooses, IEllhf!lif!cuing ~~I)«~~ d tha t obs~rlll(t ~ptkt;lt~ 1(11'1 'r(lrk pr@gresiS~QfI~'M(u-e:; tl1~ ~l1I1Ipootr(il ~ !,dli'l(QU plifllgl b~l\1.Ii€tfil f1e.pti~adon
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mi'ot Uflld:er
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d'lCckp;l·inl"-, 'ompl~tk{m. DNA replication ~. li~ ~ni!nlic (;!lli:r)''''' ~hn.'!lghl(t"I ~1!C l>.~bjc~~d to ~hm~lgh Ihc cdll ~>.ck whC'n DNA !selmiIQ~ ~'qgul;.riCin by .. 'eh~l'i':'kl)()il1l ~n~"It.:<IIlI~CIPIit"J;Q.I(J ~ J"J I~ l"il£chm'iism'l,', m:'t l\;ffiR"d It} ~IS ture clJilry kiltu ll'iliDrx;.il; \\ 'O!!I1d be dt'ltifilielilif~ Iii
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The SmcS·Smct'l cumrle);. it;. r..::13ic.a j.(} oohcsin and coll{l\::mia and n.!lfLd~m~sin DNA It:f}a~r (9) ..Cells C';.:tprcssillg ~hc .~·m(';}-i.'>ml.fl nm~1!nl.<llldes s~~O'wed S plmsr:;: JlJd an.aI~1m.~nll}' limes sin~iliar to dl109::! of wiild4}lXl ~ cd!s Wig,! Am~d ng. :s I A". :Ind lihe ccl1!lr.1II,hl;(:k-
~rn
poi~~. kii1;;L~~~ l'hrcl53 Wa'i ",~~i\I.,~~I.;'d .dkr Ihe GUlly limA nililo!iii ... L~lidL,."rno:~pL,."DlIil'ii&.,.j.\,C 1;'{md~ti)[m~ t.m). nt'ri~ul"I\al DNA. (rONA) IIITII)I niJi Ihe
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9 MARCH' 2007
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wl'icrt:l!i1'; l1l~ IEr~'i3~C fFig.. .1. B !iuicl O. To ~_"p~{itcpo:s...s iblc telomere ~nis;';~,~n::. dc'~~I;~ ilt! ii'eG~lk:ation r6rks. we COi llJJl1it'd IIC· g.mtcd ~o (i~\li.:' pole in ,,'mc'-.;..-rilc6 intlliUUS ~ r1~. tjl\'el~ 1~~HCillilll~ IrDJ.'\J:J\froil~ \V"m~dI sltl{ -6~SI' SIB) ( I !:J). Thus, um:csolvcrl :1~~k!l£.e:;. i bCI.\Veetl mlll.,nl eells by 'Dwo·di~nC'n::;iQ~ml (2m gel elees i...rcr chml'llmtiJs seem to CIBU'iiC ,dhrmno,<;one tropheresls (sec ~~l.Ipp(lI'~i~l~ m~liu~e maleri,~1 I. IICintlisjwu:tkm in ~h~ 1'fIl!~"II]t!i, Deletion ('if' 'We detceted an increase ill licpHc;)! ien I"(uk ~hc lIC("rimbin,ulo:n ,g.:!m:: RADJ] I):llil~<lll),' :S!lll~ b:;micr (~~H3) ,~m:!i.t. rooomilimuiol], amI WI'" ~rr;,~s~~s~le smll.·f'P'} g~m\'tll! dcfecl 0 0). \JI.~,o mimuirl!~ ~~rur!ilIll1(l~in ,WI1dJ:..f) mll1.'1n~8· duri!1g s ph:(J$,-,rfl,g. 2.A <II!d 13). The n-ct;u!ll'Ur~Ja~on ao.."CT\',!;"l;i,! nlmilc'SI!'\"dl!!ctio1! i~I uond ~ j unc... n f' !i1.n,JC~LHR;"S iJ~smL"6-9 \71'<1[" not c,ulI~edby :(111 1ce~. ()!de Group, Med':lCal I*starrJn Ccunci~ {MRO a.inf:C1!~ im,"rc,I.\~~in Ql1.gin ITirill~ h."Cal~\,VT Ic\'cl~ of
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IHlIlUnt Vig. S4~ Uncxpcl;:t1.i'{lly. w~ four!d !lm~. Ilu.li!;;.\! '\,V"f ,~UI~. RIlPt:'(:j· 9 lI~lIUIO'lJ]l" c-"J:lihil~d 0111 h'LCre~~ ~rI Y~aifC rop:~iclll~llill S;Unl)(:HIil'I::S \.~i!u.:rl cells were arrested inliJilCl'~jfI]ia.'>C (Pig.2;(") I i'J). Tbus, !l:m.r..'6·9cellsare slill replica~ilig
~itiic,<I!U1~ 1)1~h{I!rter lilm:ll 'Ihal ia WT cells (.Fig. 2D). suSgc.';;li~l~ that .~mt·6-9 m~CiI's break during (Itc 'Ifl'm~i:lool, TIl IIJ!s.!h i!' 'lm:C(:d~re lmdt'reM i~l~JI(!,~ ~h~ m~l[J,u 'of tl~11lCl?l ic,~t~d n'l !flNi'\ !;l<lP~ I]K"~I.i~m,lIO)n~ dls._~~1fldtJ,iEim~. in C ~']S smaller l11'<lD~ a fcwlkiiolC'l.l."i.l.~~ <l!f'C !loOt d~.n~C'~ by [I~i:;.technique ,~U), We cen..hle dud~ lh~~ rop~it:<I!lii!JHor ~hc :r][)NA f(;lllion i8
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dCI1i101liSlrrLlil'lg
thE!'t
·lh~ a1iUI~s~o:rn!ti~,rti!d:~ql!l!~'l!!f !.O~his~. I tfre~l:ai1l'&m Depa!l!l!lmt 'Cilfl~f!SM~ci~ B~q!Je~ F<!I!(!/IU'Ide ,MediciM, UJlim~l~t ode ~Ickl. MCi~~l Rolg .2,25008 U!k!jdil,)r;Ii!lm. :ll'fe5omt ad'drBli: umdladl & IIf'r\fI3otlj.adoo. ~I;al URi· 1lE!!5i!:ana<NIlfEl!!fal SeJibra, de ca*~rt.a\ (a~e~a del. ,RG~n:o s,iill m'W~~ 30010, 'S;;iflt;ll Quz di!! T~ne:rife, Spain,
there' is a dC'II">, i~ the replkmioll of this re!:lion, To Juml'i:cr C<Dllnrm Ibis 11(!1s.<;i~]ility. \VC
~.1I1iL~h::plicJt.i:on. n
:a[
iCI1Sl
j''o~'
ebromoscme
moniWfCldthc n.:pl icoui.on or indiv idtu;mlrDN A~ ci:lnwin i~ag chmmos<lm,~'S b}' [heir cx.~~nsirm
b~ addirmeill. iE·iT!ai~
.induced to ,~nC(nporn.lic ~l1onmodC'OxYlIricli'ulc (BnJV) in their DNA duriinlg~hc previous !fer" liC,jtion_~1'!;wdi~ion.wc hkll:;k~d tt]ms ill! mm;:t,~· I1hil!'i~ m cnsmc lh~u~ ~ht')'d~d nOI. L~lu;,r mHY.!h~·r
n. Cells
XU. We cvalliHttcd whd1CT l:cpi.iC'ali,onie;. ,",0ll1ipl,e-!ed i~1 .w~/('6-9 mu~Ul~S bdbre !ic,g~;lIkm U:01i'!l!,1, 'Pl~:lsoo·Jhi.~@ gel C'1\J(:lmpborc~~ I( !)FG E). s
~'I:I(lOI~lp~!C'~cly l"C'p:llk:alcdi chromescmcs do .!lot ecsolvc Iby .PFGE and reJlnaiuin thewells, In \1IfT eel I:;;. YDuclPlilcmcJC! (Fi!;l, 3A: urnc: () mil~) :;)Ind r~II'YfJCpl iC<I~cdI[ fig. 3 A: Ii um;: >1\0 rni:!H cll;rnllm~ol1lc..... cmL.'red thi; gJ,,~I: .o\'!.\~\~r, <BIll I
\,\I'C'I'C
A,
B
OIS1l1blJlbn ~I' ~.
C Cl
l
jJ
I,~,
;S;
lan~~
:2104111 - Gitll3liM
'41.o1!'~Y
.:oa ~
l ;,
----
,
"
'70
00, i5Q, 40, DIM:
i501
;25,
'ti
.", m '110 0
...
"
~
1J~i-9
DI~'~'
30
1
l. PaSi5eige through.:S pn",'S'l!', but not tllro~!1l11.aJn~pn<lse.ilil, the absence ,~f Sm(5· SIiI'tCO'p:mdU!l:flS cilu:omoIDme n.tlllidili;jullilc!io:n. (A) (.en qde dynamics: of ~P:lmlLifilelo ngatiolll in WT C!lnd smc6-9 ICi~H~,s!h~~'!'iJi!glt~~l sm't1i-9!ill!i,!it.;!~ts ~rt !liot det.;!y!l':d 'illl (:~1l qrde pr'ot!i~es.si'ell. The~lili(:a.ge diepkts, iii w'~;Flr'~if1!lI'Itatiw I1I1k~Ggrnph IDf spind~e~ dlli~in91 tile time IC,OIJrse. ~B~Segre.gali(m olf 11 (hmmo~ol1l'l;~ ttel0:'48 J.lin~erte~iln Iflle 'lelom~re 'britt olr ~a9 rONA. UlldiM tliie 1I1dij(at@dcGJmiijt~oni~ nrl m. smdc·9',. imdllQS"I!5-J: c:elils:. ~(~ ReiPr@s~mtali",-e mKrograpltl:5, of {!!'!~l5in (8). Rled, nlJldifilij~9 r'~;n.. teJO:t'fS'.7.
Fil!;!_
.i ::.:
,Sl~
!I.II~
,,
20
(I
10
~"b
2S,~
1412
" MARCf-12007
VOL 31 5
SICII~NCI! .www.sr.ien.~mag.org
REPORTS
Chr"(lJlKl!lii}me~I'il~~lCd l'OCI1~cr the g'C~ l\}M~e r,:;pl~c~ting (Fig,. JA:iil~~c;. 2,(1to tlO min). hI ,;;rjJ~'6"'9 I;I;I!~, 1I1!11'1;)'I,mt .:!!~ d~romo~om~ XU suess, Sim!lIlJ'
levels
of
~~'lt<!$3phosrhm'Y~-
or
~h.. 1 i~~ cells '~fig. 3A). and the dc,!otion of \i\/T RAD52 !md ~HdC'cfl~el (Fig.lA~!' To :.mll.>,2"Cw~.\.'th~r Slil'('fJ<-9 '~"C1I1~ re de... fie-iell[ titi ~hc micl.iv!ll.iOTI oW k~'iOil\'u'i checlcp,~,il'u ~~I,thwa,y~,,,,,'e '[ICl':k'!!, wh~l.h.(."i'" .wiu'6-9 It~i,lI,m~!); actii'V,ul~ .R.rtd53 in i'C:5POiiJi>~ to ~~I)lli!e!lI.io:ul Fig...Z. i~h~in IRBJlill~oiOn of me!.aprnil~ in
c~l1~rr:. !Irr~~h;d m. b~cld~~ (:dl!.;. witih o:iJIC or HO i,I].dm:iioul, bod'lm.ym:::luu]lol,ll>Wl find nueleus in reSfmJ1SC to NlJ, i~~di.'C"",.lu.n,g I~at .~'llit·f,J;''9 coTls arrested ill Gz-M ','iI'i'~h" !i-ingie both M:mi:n.1l hlll~ !:hc oc:U c},,\:'k wh~l] l.hL; S ~hcl!I!ii ,of ~h~ dlcckrQin~ Il:[ol.cin Ddc2 'I~g. phu!>.;:' ch~"ekpo~!H ~~ uJl;:ti\l~~[l;:'d" I.:. Ilbluir:'lot;-d S5A). mid n~rcl.s.3i.it"li v ~ncdlFi.g. 3q. ChCCMW
S.i,liI!H1I.l"lt(~ul't:. w~jc,n cd~~ w~re '!!Xpo"","'d to
tlut'I.I'!)11 mel.btlc~ilil~i:irlalc., .~natl~il~oii. mliUliml~ poi.llI CUt aeli'!.la.l.i{ili a.~i>Ooecul'S
;H!Or~ were dCI~~co in \\IT !:1m ..,:mdi~Q cells • c;"\po~~d!.Q ~h~ fi,bomu;:lli:"il~id¢ ~d~IC~'1,,~ h~~ib. it,or ~IU (f~. V J), Bn~h WT !]ltd J"/~.,[,t'i-9
or ~hc 'DN/\,~cl.m1il<l!!lc ch~kp(lum in [~rOcTlSC 100 .~ing!e '~Q~lh~~"~l!'ilIn~ bJJc"k {DSB,);r,t th~ "
I\lMT
:m,
!OCl,l=-i~1~~II;:OO ~_"'I]~~l'io~l by
0.1' ~1~!!0
"'lldr~lll[lld~.:!Is.~~
~.'1m,lr:;\
wh~i'i OSB:;;
00.:;:-
sm~6-Y
iil:sid~ the
3rcr~llso
COi1'ii,PC!C'I1l in
'th~
mj,dd.l:e of
rl:lNA trE9ioolli is; II'Ill"I:filll' smc6~j9 mLnaJ'!;t:5, W ~. ~iGatiolll ~llirerme!l!liatE5 0\1' iii ~[liNA 'regioolin 'W] ~d
..
;1,-.-,
Ii'
11" ~,
I
'
r--
F=iI=
r'fI
t
:S<hl'illli'!lile: n!pre:s@fl~OOn.s
by 2,[) ,~~OOf'~is.
,
I~)
~['
~'
~
_
..
..
-,-'I
-,
liew @ntlyl'1ll'@'sil~ am 'IDmVI'l. MIS, .autOOQllOOIJS' N'l.illil1oorsin pa~mh~ represe:ntdifterl'fll~ s:irull::~IleS QIi!,lh.iil.~!iJ g~.
"
ffl
WI
~~9
[~l..
q~~-
~.
,1lJI ....
~11 ..
'wltw"'e!Sll rr;emrl ~111m!lt:!· pl1l3il'.eilIId arntyl!ll!d by. 211) (A). 'Qoolllnlikatl.ml of oor~poli!din.g IMDNA $i!nJ;c-
'9ft ,~l«~,pDesis as in
~Di!' IO:MA ,mmnbing ,i1IMlys.i~ of i:!Oi1II~1eIri-~i1nfllJNA repth:.a,tiaFl aitlh@ n~ml genes arrr.i!~. TWo trepr'e'rserntltiw r[lNAIiJJ~. @itjQ@r GDilIIpletew rep'lic:al.e!rll or ~l!"
t!Jr~ ~ ~ho!.'i'iJ.
"
"
(l)
~lJ
lI.m1i1l11glllllrr,epli'l3t.m ~p~ ,~re siiI~11. Gap llerilglhs <I renndiUltedin Illi Red.
boo~@'Ji1iD BrdUl.lError m~
... ,..."
--._ .&01_-,,-
-- ...
-=. ""-
........
-_
......
U~liTGiI'!il'l!.......... . I'
=:
; 1~
_. ..... .-....... .
_.....
----= - -'Ii!Il I!o
·llrdU
-"""
__
.. .... I!O
I!'
"
.1!hdU
IU."---------------.,,
IDNA
m_
iIOI
_
W;!
m
I~~~)'
W;~;eI1l~geO'l'·
KG ,«10 :1IlO ~
u!i'i~e.plk:alled I~.A
REPORTS
of "H known ccMIl!I:u cht;::ck]}oin~:_;;;, the S I~h":iie chcc'l:'l;)Qj~~I., NA·dmIH!,g.~ D d~~c"pni~ltru~~d t'~]ilw!1,; ~1«kpQim, Th~rQ' ~lUlm~tcr.w~ d(,.1C~~~dRi'ltl~J ;!I.C!iV"l!,i;:¥~~ i.~ Ipr¢. thli; SI~w5-Smc;:(~~m~1p'11,1:o.;. i~IH;JtlJWn:l of \11,1T und ,"'Mw6~9 ,~~ti tf~~, 3D). Wi! 'l";:Om:~III'~'\i il1ul~:r~[~k in tlre dINA ;I~ abo under nors :JI~y of I.bl,,~"'~ ;~ncC'hani:m~:>. \1I"r,,: 1i!1 d thai, d(:~pi~e~hcr a 1;'~ I~<ll .Ii! known C'h~kpninH'i, uU<J dill;."Ckpoi nl ~~I"'C i ~I'.unc~. F ~n![IU]". :'l#ltii~9 ~ ~ ~ll!'.n"nl$are Or;l!'IlpL;'~Cnl i:11'[ille ~,piP"K~l.cd~~f;k· ;I.re i!rit~c~~I~ ,~}i.'(.'6· ~lllutrU!l'~~.We ~pl uc;'-<"I~~on P t~i!'1'u.bcc"l!IISC' lliL,:'i,[ ~~I'i:;;i,l.i;"it)" 1,0 ch~ j;ril~d k.... dd:'iy in~J:i¢>~ e~lI:-: doe:;. :rlat nold UI!'I Ilt~HHi(', d.\.1PoIY,~i'i~riz Idru~ b~lfi~iillYl. \,...fL~ l<.'diibT~II"aM!i;; iUi~ Ivlhm o~'WtT~I~ (lig. S.SB). We 'OOn(;1ude l.h~l. ~hc ribosomal
gc\IlCmT<l)'
.u'ld
CXp["Q~]ng
th,e
nancc
under
the
gll!m;;tQ~\l
m:!!nQ!Y
li.'fltry IbcrQI'"C cflmpk.'liag l"eplicmi~J1 (FRg, 4('). The !ll!r1c~ioll of the SmcS,SullcliJ oflompkx, is i:I'~1~.pn,[lJIl~ .~bT~pi~lg wi~h ~h~rc;:p,!iC:;Bltinn If!£QMPHn~; p3rti~ul ..r!y O!] ,t;'~aU~nging ~C!~lIrlnat:.~ :;,Ut;~! ~lc:~rDNA. \V~ (J.D, not kl.l(~\'\· til,:, C'~ act fCOI8{loJl \I~'.hj' ,~m,'5--'!,md~ nml .• rs are ... m ~!<!!rd(:d ijn~p!.uC;;~I.ijn.l1_ (jill'.; rU;IS.~ibijii'~y ·~!':.~ht .. S~ic5~Sn~c:6 tum::liO!i l'iI~ighl. ·~requi.rtd rOr
~ni!.ali~ '~UDli)' IJcforel~e COlt1J)k,timl or~.DNA replkatimo in ss«..6~~ mUI.m:111~s aot eaused by i I.ai IliIl'C$ in the ,ccll:U1:If chC!.:kp.oi III I1,Hii.chinc-rae.<;;, Our d:lIs. iIIdi:c:lte~b:u rellHca~ t{)'~l01" Ihc rDNA ,I!CSlT:Oll hi.:scvcre.~y dela)1cdi in [i~e ahscnce 01' SmcS·Smc6 lunetien (figs. 2: .~nd 3tH We ~~,tl,,\d'~",'hClh~r dimi~laliug prO{:'~i!i~'$ !l!OU pO,~I[nl,el. lil~plrc'rucioM fork.. ~nthi; rDNA \'\ID:llki . 11,;duee lhl.l ,d~Ii!lYand g\l~]flrc:'lS [he ,!:,'hronW!.vol'llc XU 'r!on{l.i~jl!.!lCliCln ~]~.ll;.·nol.1!'p~.fheifDNJ\ l I:'>
C'o:rnp>o,..w
.~
1'!HI'.5.&, ijoW
, ~1'iO,rr--I':"""-I;rrl-~-l i"""H~t-Ir
i"""T"~Ii+-+!-t'f!i'....-.t--+t-i"li!'~--+-1
-ij II!
IDi
~ionu lIy repl iC;JIC1(),b;(;"Cl)I:u~~ ~Jll~ of prescnee of :l), Ijo1!~r R.FIt m~di3l!r¢l by th~ FoM protein. \IV h:i;eh is next 'tt:'lll'h~'35' I.i'faltscrirll[ion l.cffiini:illl~ iiem she Uti . .I5l I.n \:\11' QclIs. replication rDNA hl'lS to dC1l:1\~'W~ the p.~S.C~lec of ,UBs and ~lC' dLls~erill!!: ,ot 3:C! ivc migil1ls U.?}. wh klil I generates a shua! iml where large ~:lugl'l1l.C]l~S~I'\C ro:plk<lrod by OJ single righlwllrd~~mwi~g fork ~n lil~ ~h~m;C' ~ Fob 1. \\llud~I'Ilo'V.ing arc not b~o(kcd ,m the RFB :i!.lrlC': thus. rcpliem ion is acecleraied by lite 1ln!'SC'11CC (I f l ~I[O
or 1:J'!l!dL.:'~nr1l;~pi.·'U$~I~~
lII!1~di!'l.;"c·
or
l~n
lork~
Ie
1
~CI i\1'IC
forb
rFOB
chml1mQl5fln1J; .. XII !":I·Qn~ ~ disjmli;lu~;m from 63[0 34,% (Fi!!:- 4A ,i~ldfi~. S6A), ]111 ruddituOIIIll RFBs. thl[~!ligh m~U]~ lOcripl.it'i'!i rares In !'DNA gen..::s are ml$Q Hkd:y
to pose
11ChllnCI1~e
lor
1QCljyC
ibc' locus.
I[rl'tNA)
[nllct,ilvfu~i.0i11
gene trnnscriplicm II~c simultaneoss dclleliOll polynu;:ffi;"'c (fol) I SUbLllli.: /\,135 .,mel the imrodllCl.iolJ. ora Il'IIult~.oo'py phllimkiJ ,oo.nlilaining, du: .)5:5 rRN J\ cod iIfQg rcgiollrl~:i1ed to an lJtNt\ I~o~ ll p~mlnfll.Cr (Ui;. JiIleilll)C'liYalion of' clnomesema I I:R.N.A gene lrnnSc.riplion redaeed c!JuunKl$l[lIUC XU .li1olldi~liJ!nc[iol!l from 6.:1 to I~% in :~m~1Io"9(fif,;. 4]~). He: .~i~lIlJlJ:I·OL!:n;!Q~1i1i in.1J,;l~Y' .. ti~lU! (Jr I U"o.Inl;;;c'ripiE im~ "un;! .~rl i(;,lliou b.'~P'~
or
uq.:lIknliQJ1
fIIC~
Tiling,
:2
.;
-31= .~
iIIiili
r!.;.1:";; Itl\,j:\ ,~",gr~'!il"lli(lri t~i' tl.;1illr~mnn.-;m11ll': X I.~ ill smc6-9 1]~IIIII;!.nl$. wiilhvurhl!<llly \1i,1T..:l1'ici1.:uc:y I.Fig, 41ll:lml Ji~g S(13), Thus, the d!cla.y in-I •.
'1~
stJblle fo;ri.; pro,grcs. s h:m dl1iot1s11 ill..:rnNA .. A cc:llirulilf chcckpo:ini C~HI be dc:r~lledril,'i [I ~noc'haiti:sxn that halts I'he cell cycle in \>VT celiis in response to OJ, cC:1.1alilc('l'I~dit~on (e.s., a cellular insult sucb as DNA dimn:!gc). anda gene i)l. elassificdas 1'1<1110.1' ,.1 checkpoint ir nnna~~I:;. 'oFll'!is gene dn net i.lITc~l [he cell
c h::: <l!'O.\i\lT od Iii. 00. We: Itavc ~lQW!1 ~Im~ smc6-9 mUlWnl cells I:id~1lvl; c.:.,."u(:lljl li.kil'i\i\rr cdl:;. wH~~ I't~$~ct In aet'~v:J~iL'In mi{~, IjJI~in1'~t:.,
C')'
or the
<
~i1[ub:u]hyw p'ro~
IFig. 3. smcJ!l·9 ~eU~ eliiliE!:J' an~ph,glse be'flor)~ 'C!ilmlpi.~~,lI'19 ~pH(atiQIiI of d'li'irDm05tl'liI'I'eXII, d~$pite ~h~ IPIF~IIN!(e @f ~If!M~1 dhe(klP(JI]~~ reS!'po~~s, (A) SY·iflcli1ronlz@d WT .i!lnd:r;mco,:9 cII!1tli!J1~S a~all~i!~,d by nGE. ttl, visutlli~ie yeast (IuorrmslJmesan:d therr w1eplkaltis'n, PfG) were transJ<erlF@d ilnd hybridized ~1,Fitlii a p~o,oo' 'l~ 'l,i'c:lrri~'I,l:S111',omO;SQiiI1:~St,o qlJl ~Iill'ify 1~l'Illry ,of lntliilfd ~~~ d'lromIOiS~ me's.. (I ld~'i'!t1Ir'i(a~'k1l'l1 ( of ChrOrn(lSi!'Jllile'XIII entlF:fintiJIlh;~ g;el,is ShO~¥flJo r'Nl. sIDc6,9, ilnd srnc6-9 rod52t!. (E~Is.llies;ing'le' ast~ri:llk llndl1ci1Ites ~hll axi~, La he]: tli me (in miin) o1rt!~r G:a: rre~eil:5.e at 31~'.1i11(B), IFlat!~,~ p 1~,:511lll1l.arLy r I)lhoij.lhQF~~t~d! In W'I' end sm((i-9 ·cll~l5;;!~t:eJ HILI '!~~tlille:nt. A DSB li!l M~r (C) 'C!rr[lNA (0) ~fldlJ,!~!I;ls :RatiS,3, phO<:5phouyiaticmil'l 'WT .mdl 5fl1r6-'.9 {eUs, tlr1l'"'ClI\~ ~II'I (8) tiD ('OJ ~mj~(ille Rad53p spede'5. ~Plho!>p'ho~~,ti1!dlR.a.L'iI!Bp is indic.ate!:!1 {U). VOL
31 5
SICII~NCI!
.www.sr.ien.-emag.org
REPORTS
"7!;I~'fIl'1TI- i~ deU;[lll:Cll11i!I~CI OhrmTI9foO!lII<aJ ~~re;)jks"l'ld g<lp:ll, observed in Ihe Inlcgri~y ![I the whole neither ![IngQ!llg !"ork s nQI' ~Im~pli~ ~(!mm~~. We ..lso mJIiC Ih"t Ibil~~K~C r~I,)~i,t;:;!!~e~lC'!i1p!¥<l~~ eel I~ are OtiC to ~nfl:!pr~c~tQ~~I)NA. to q1~,'\",t ~1tJ;l;rIl~n."~ qf rPNA~~'ij!l,g\t,;"'"Pl cI .. s~iil;"'! ,;iI. 11~~ .'f~ilpi-9 m~lta~tcel~~ ~h:(m ,,11 im!~i!\~"!i~in Qcffliin g";'lol~~i~n;l~~n:il.. ~w;h 1J!~~!Il~ml~~~%. i;i1romo1SOI!III!lIi n,;"::IIT,IJlgit;Vlli,ints at <11 nOll··,ONA oU!,illd pp.crliij~IUy il1d~~ .11 '~cU I,;y~k ::JI:nt.'t'>t ~h~dl"P{li!ll1\.."'Ifll;iIL"C ~Fug. 4D1 rl~.,\,,~ h,I\"Ll lU{'u:> prJ), which, fJ~ il~nmlmtl1i:l.!i~linl frugiJi..: I!J~~~!!:I il~~ rDN A as ~ It1.Odt] ~~)r ~1)liOition thrnlJgI'iI ~K:i.i\,,,li~II1 fllf l~IC "pl!:.dk oln,'ek:p.uiult. or clllnp~"'I.!"!l. Ihi;:rDNA .p::gim~ might he COl]· sites, eould be a'nr:ihu~I~C! Ihl;; $cgreg<.~iCl~~ 01" the R.ui[j!.(]C:PC'nd~nl mLd";I!iOIJl~!<1SC check(lm!!1t. ~;idcn;,d us ;1 .. ~,pcc~a~~~n{l!mi(: il:lcul~,_" HnwIn In~n!:ll<l]iall ~I!b. rr..r~i!C' :il.~~'ShOlV(; hi\:-..'Ul r<uil~~!ly~piiL;';,JI.tcd:chmlll'im~(1Im!'"",. Sw,di~,,'1 wilh t:'\'t:t.wc have si~O'1!Nn dl!ll i'l iz.; under !ioro'i1!~1 z>JlmV1l1 to n::rllic~ll~ IgI:(;: ,lIld ~n k :-;clisi~i\.'1.':'10 mh~g,.rU:!§l in ~'~l1L':1gnl]liol] m'mp~,I;x. h~~\'~ :showll C'hcckpo~lil :;,lir;.ci 11ill nee, Dcc ~LLJSCfDN repA 111at lh:ilri!l~ S.phm'~t LhC'.lt:' is nDOnk Ih:rt~h.old, ~pne-atit'H~ d.::JI1IIj'~ (n-19'~ whi:d!. ~1I.~~':iI:5 Iil'i!it fiOt ,~!1ccIqloiru ~ct~vlili()i1 (211. Similarly. illl IQrk!l pr1)g~5inglhr.oLiig1i 1.~~O;:ote [~gje..i1S i~l Ime r>!..'S~nh;8 to 12:% of Ih.e y,ellsl ~~~~ornh.\ ito[·· n~ lc enti)' belore coinp~,e~kmor mINA n:pliC'l.lliml G~ do no! ~i,!l,nlii.l!Umj1\L'11ie [t~]lI~ and thm. [I;I~ llii~rK'I'tttrbed e<:M eye~e.~.'oil~oin~ lork:; bcdow 11i~]:l'res.:1li:lld level Ini::hl r!li I to 8c~ii\"lnc !)
pZYBel'llmt
101:1
!!!)
lIB:
'lill II!!
i
l
4-
i:
:50
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IF'roml ,Sw,iimmlingl to Walki,ng' willhl a, Salamander R,obol Drive!n by 8 S:piln,al Co'rd Model
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Tihe ll<lln:£ilioll'l hom aquali~. to tl1!rrl1!~U'i[l& LOOOlil1lo,ti'onVa", <II I~;e~d~~opmE1ntiil'l \ v~rffi,bra,t:e evo~Lltloll,. We pre'5emit ill ~piinat (oro mneel dlnd HsimptemE1il1tatiol1il"l an allT1,Fliili'lliom salama:m:iler rmot tna'l lCiemo:nS<tr:atfi!:s how a .[lIrilililitive l1e!IJlral cir''ClIIii~ lor S'I'IlFTImlJm!:! Gill bE! 1E!x.tendeclbcy pilY~l!nelilcaUy mOI',e< re[I!J~t limb oiScilD.ats:ry (iel'lteJS to !axpl!aiim !!hI! ahi~itf (If saJilRlo'l nd'@rs 'M! switch 1b~~wl!ii?l"Is\vimmfn~ ,jil!nd m ~kililgl.lli~ m,lD[h~ll 5l!Qg~ts, IFleuralll1ll!lmiillllismlS. Jor rnladiul€ltian of !;i~k!(itiJ.,ti~:F@ni~:n., and type of
'!JM~lhat ar@II"[email protected] f.~fan '~l!ia;pad:'S, It pr~iclt:; !hat Umb '~$(il~tCirf mfttflts ,11:1!V@' llrw~,r ]ntrin~1t: r:r~ql!lefl.d~ thalnl body o5d'll~lO~ '~e,rU~rs,~dl \lie' ~rill~efI~ bi!olJ:lg!(ai. dail:a Sl!P'Port!IIlD this, a
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fO ImHJUt'!)' nUI!(J~L~J J ).~hi:!! Lm"'l illgpropcr! iC$ of ~ M o~illmory cC.I~~er Ihl! o!'lCillmocms 'oo!wcen t1Uf)lI~ of nl{lb,1nl.lll! mn <leI i\'i~)i nd p~ri(I(Js of a re.;,t~l~ nlmi,~~i\,:'d y 11i!!~~l!!i!i: of<! ~~h;if'~ b ,(~ciU;u~,
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lamprey ilnd ~ll!fmOOLL"ly ptodlll,Ccs [LllVClillg <lICliv<ltOO, '\\ ,~~I il tonic drive (~iC~ <lI s1)rt;. pic OOminli'lJUii;:sum\Ilmxm). 1111: timb CrG. ,'\,'hel~ a~~~<lIed. fo,i.t(S dl£:wlX'i1e I"G boo ~l:,~k ~ng mode. ~ P·(!~"liCl!!.~l}''~'d ~n 'll l'. IIy!pCI!!h~ Is ~; 111e'sI]'cnglhs of i1lC COLIp:i~l&" Ifi'OJU li<I'ID!o b&;iy G.1:CillrLllm>al1e~m~rthan ~~OSC .mn b._"l(]!y to body GSCillh~CII'S and fklln bGd.}' 10, litnb C&'iIL11l.~:s. "[~~b!, al ibl.II'S (he Iirnb C'PG~o ''0\, ~mdc'''lh~ m[lnr.d!c~"
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h~illtW..~ Cn!.1ctvuing bioJ'ogiOl;:,d
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or~h(:vodJ' CI);G to I]~UCC !r.llvdliL1,g W~I'\.1."l5 ~l 'to jl1tMlllQ; ~1!1Ji!\g W;;~\IC,,'" W:I)1)(lid);.~~. 3: Limb otiCiUato~ ,L;:iun!1t'l,l al('~lb!J.,': ;it hi,~ qllll~,nti~llilli ils,. K')' ~Iil:nlit:!IliD ;SI.o~ ffie1 ~~~cil1g d I Ul
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VOL
31 5
SICII~NCI!
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REPORTS
sbow tim! O'ur n,~ medel cOIn ~l~t:mlC Ihr"o,\'1:!!d i rllrt.riPJl wi~h \'<!ri"h~!!; $~cr;l ::md hCiildillg: (J,g". "-'"I;~'t;l~ !h~l~ 1!lQI;;-Q:ru '1x"w;Jy"!br~~,i,~~~iGm'l 1IJNJ Cj'lm011~ ;;,.ldioo lit ;J'II;,I),IRumiI~cd ~11~Jli,;):(ij,illw qli~HbrtiJ\'1,,'ly mn~pjttru- lIu:: ,gaii:s. gl~I~C;r.lft,-d IIln~ tn of dle ,ro,l~ ~.;"lImmllidC'j: arid (iii ~~!J :>~m'!r' I~1L!!]:I~
F'iig. t. (Q~fig:i,ualli~Ui1 ef 'lihe (]~G model (A) ,alnrl sallam~ mldie, robot {B), Tt!.e' rob!Ort is dri~~ al(~l,hi:ll!:e~)'., hi oge l'C1inil.$ JO'F thespij ne (iJ~Ci(k
c~m::cp~ of C'lPGs crn~ ~~ to ro·h!!81, ,~-on~otiQl1 co.mr(l!1 lOr wOOl:,.;; 'I.\,'i~~I1I,lLd~irll;!en iC\l! lnted jl:)inti'<.,
lGht,T &X~'l·!r.l!TIgmbo~ 1iS<gC~jJ'JIl!;:-d ~ "PJlm;."i.
pi~~~ UI1~i~lhc fl:~ <I~lln'!;ll. li:n]L:t\. pcrrQ:llnonJl~~ll1!9~ !1Clmtion. llWro!'1nj(lll rep!~;tlJ,'4' '!h~1~~~iorll;l!
~.1lla"tdy~lm~d~ n,J.;:kilfK::n~ti~ :rt!il,l~~l1; (If ~~<Illl~' dt::rs ~Fig. I.BoJl llt'l:ruoo, ean mmllC: ib ~iD!lIrillfihz.;:IS l \l<o\ciIll~ lJ!1L1(h.~ :r~~a.d u!~lvlilltta:~l~ {l!I'~lC s:pil'!c wilh
lfJiylo tll~ ,~UIIIl>t;!\vh~kin si;Il'~IPII~ rnrnd ;l!!!rn.\',:;' d'k: :nhl:111fli.im] lx1\~1.,1b'1sVi,·iirlg Imd slallc~ SG"iPO~lfu; Ilhe mol_gr oonlmnb1i. am b;~roI(In ~!l:l dirr~lCi:
~m
by 10
at malQJ'.s.,
wh~ch
dis~~n ·the- ~hl!lmali:c \ftell.\I' Glf~n@ rol:iQltj a I'ltl rom rot,i!'!iQml~ .io~nt5 hl,r the limbs (b~ack 'C}!iliml,ern,. (I~ is The UlmITpOSlla of a !:lady 'CP~a douhle cihaJilii fiI,~iJt9 foil' iliilvQl'I'g thE!
splllll 1fIII00~Of!F--a a fI1dl (PG-4,~mill.~atm~ror driving Ule, ~illlilbIrnurm,rs, Ihe 'ClIutput~ Gl,f~'heosdlli!imrl>
~ n mb
reS'!:'ne'igI1DrD'r
COlli"
torq,L!@S (t~ ro UQ iI tl1~~!' 1{ol.ta'ge Vr) 91rvel1~he adU81l alm'gles, ~. lih!!' C!P'G lilmde1 r'eH~i1{~£Ilen il nd rig ht drives Ii n::pre.~ lilting .ae:5{e,ndi n~, sfgfil[ll£rtr,om~hf' M llFl r,egriCllillin the brain stem, lfhe \f~l.odty, ,dir~(tioll1. (llildlylPe o~'gait e~:nib~tJm by the IroOO~ (ami be ad~!JI~'l.ed by Iilllodiryffin.g the'~e t\~oslgJflat~.
!1i'I1!(),t~w
fig. :;:I,., Sl",'il'chin:gri!'Q1'Ii! '~l!'a lking t<l Sil'o'iffl"trnmg, a(~-:ulY Qlf !he (PIG mOO',\llllllhE1nthe ,dri~ S;i~J!ilil t~p',fC!9r~S5h,l',~lyh,,(re~~d, (A) XJ signals from Hle ban body (PG O:5,{~I~iilt.UfS (M~mfltms on th~ to thst (lir :rn9l. lA. IJI fiI~tsaN! ill radians (sc,a'le !Jar e:Jl tJile tol'l right)., 'lh~ ~ed lines mll~tJ;ale Ihe lralil~ili®11i 'rifllnil :Malrnd~ng 1,'i',8J~~ (\'Ai1th syndil·!i'(I!i!Y in the' lfllllnil ~yn:c~roIilY if! thetaili31n~ 1111 (!Ir!lip~~ [email protected]:fool:l~l\\!@~!l ttiil@li.'!'CI,'\\ ,~ <: t <2'0 s) to [r<iI'Ol'elirllgt 'lO'I'.aj~~ (20 So < ,t <36, 5)" {R} ·!!If sngli'lab h(llir~~'he~~ft· lij'mb (lflG O\SdUa~,()!fS, IpsU.lIOO'f-al 'r~f\e- afl~ hindllimbs ~f_i!! 'in al1liph'(j$~., {C) ~if1S<tntan;eo!!l~hQuenci,es rne<l:iure.aas B; in (ycle~.lIhe VilIriiatioJlsill Ihe il'lsl<inti:anseus heql!l'~lld~s among in;divid'ualo~ill~tars :lItliIirlies~'"" 4 s ,a,nd r", ,20$ GClrirl!~pcif'ldto bril'r~mc:elllralions andl dE!ce!eriiltiom Ibefore l"El'.!i(llllldllronizalion. (11)1) :~][;'IlIa_rili!,~~~e 'o~ the i:lrifrVl:!' appl1el:l ~@ana$dllat®rs, ih~ d
right :!1i~~are exactly iln ain'liipha~e~. nu~ IiIl!lmb.erii[;'l~ «m'e~IlO1UrtS
c
f
....
I
II!"
i1I~,!:I ulpper
hor~illoJlll<llred Ilines [ones,p.ond 1:0 Uua lo~'ll!r (~~ ~~ 1) (~= 3, d~= 5) o~dll.a~ion tlllre:'>hoM:'lf~r ~imb
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35
40
REPORTS
.'11:~ sigll:Jiis rf('lm~he Icii atll! riigill, body ~Qks jOTll~ in the 11ock. lh~ body"lhn~ OXlmlnaosc ~~mCl!~inr d!e <'Pun:; rnotofl5.. ;Il~don ~hc pl1<1$\,"" tion in l1!O.h Iherorot "nd til(! r!C<!!ISiil.lim~1!!1{1cr '(]';or th~ ~ imh Q~c;:!l~:atofl'i rafl~lL;'~imt~ ~lKl'[[Jrs., ~J~li~ ~~ k~'!g~h(2.~,. Im;:w.l'i.i~~ the o!i~
'bt,,'1\\~Q\,'f1 ~le
I,.Il!llirm t;JI:~PL-ril'n1ll;;rll.~, V5). n1iC l'iI~oBd P!iMh;I;Coi ~ abniipl, lmll~;li[jl'i bt!W,.;cll !5!'li~ lii]l1p~Y ~' V".uyirig !hc drive j f~g.] ~ IJUrD'ig \!',nllldilJ!l (ii,c."at low ~ri"I:. i Ihc 51ro~lg (i'f\tiplliul,~ rffilll' Ilin~b 10 ooqy t:J~jlhlltib'i:. ~i,)1t:C n.11i! rod)' ["F'{j to (~illlllte !It II r,0\,,"' frcquet'I.'CJ1 w [lh m, S~..J1a!XlIiI st."klldil~ VI"<I\,!!.a'> ill~hc elee~romyo~lll {Ei\llO) lIL.'Q')roi~lg...;, {5l', 1lw.::: fi:equ,,",1.cy aJlI:l ID'l,lllliIU::lcofo....;cinL1liol'l ... i'lCreil~ rroponk~ll1ln)1 wi'!l~lhc dJ~\~. i\:U f = :W ~nh~ liJnh Q!'Cilhltl;lU5 S>\~urd~~ ..<ll'l!d!h~ indllct\'> a rapid !!lai! mUl~~t~fll]. t<l
\'\I,dki!~ i\1l!ll;;'rn:; 'l~mt~ C'IOl~istc!lltwiuh tJ,~ or ~h~~1 ~"w~lu!1ib. Jt.. Q~W~~ in MLR ~turl'i-
~;d'i~~}ml
dlilL~ do
the
aLl£r~l""lm,iiOJl of ~lt; ~'I;;ld 01' , ..~!lkil.l!!. hi~lt:1r ~!n,"(ll!I~I:J!;.Y ,llid ml~p%~~,II:1c orr~
c:inmio~ls" lhi; t~I"Jlki'ig velechics ,wl\.lin!lld mq.ge from OJB In OJm Inf~,U.WI ~D II body k~IIg1!lt!tt ( ,(I. .. f 1111t:'1~li"'t; !i'tnill'i. ~mt}(ll i.: ~lowI.TI!l11In"lfl n;~!#lii, wh.lrhwnlilotr; fll \'dOC~LLL'li' 1111 r.m,gt 01'0.1 to 0.4 'chL:' ixldy k1l!!,I!l't'iI.~. The dmbrei1.~c c:a~! he (,::li:~jlal.lled by lite- ~lJ~ICrrreq,ll~i'l<ciC$ H~dI in dne moot. 11 ~c ),wjm~ning modi! th~ IX'!bot is a~so COI1.<;.b.IC'IlI, '\Ill ith dlill of <I rca Ii sa !a.!ll1\11Wcy(Fig ... ,.
or
The
'llm'c~:illl:l \\lfl!\!(!
or body
Ulld~.I.n~O:II aU(iws.
dl~ high:r-~:retll)l1~'ltcY SWiIbl111il!ll: 111i:l(f.!;_ l'r;I\l.,.d~'Ig W,H(''"l few ,"'~I'hmting, are l\llc,~d. ,~f,LJ as in [he EM(iruoorui.].~ V). lll.ei.\!~r.w~·lli~-J;l\'.... .\es Ul· ~ C~~ ill nmpl!!W1d.l''!Ild 'kq~il:Y ,~,nh!i.: dri"'~~
fi,Jrlh~ incJl_~~. lIrllii Ih~ bod)' ~ill~~ ~.db ~ht'tlr 'i,I~y oscil.i!:uiDJ1 L1tlC:Jtah;1 cLnkl f:ilA'1lP)f;C.u~!llin!g,. ( lli~ 1t1l1l:lCracaJI 1fl~'~1E aMLtx: Wln.11 d!lfliLioo ~dnlC'IJI~tic nlfm~'y.'>II!s of tl~c g."Il1 tmlilshiOiDS. \1!'lttc;111 lotu"::lll'llll! u~Il'\lclli~lg 'Wa'"VCi> in 11.£ lClDdy asis 8i~ UillJl. ot'lSi;.'f'\IOO sillll!.l~I.anOO\t,;~ywilb limb 1'110\ 'Cf'l1lCJ'IIS, t<{). l\nCllhcr :iII1pOutmt sin:lIJIa!i:i!y Wi~l I~C MlR llIumlllkl!tio~ o;,~rl rl't1l11.">;wId \'",ilh I'W(}ITijllg.>l in!~(p(ll~ O!) is b; Sll'IJ) incroL'C of ~it'qJ)'Ml;(:i(,,:; dIlnril1!g 1IJ1~ ~rnt1lSiljo:nrmlll \/l'llJkitlg to s\vinlJilling. IJil 1L1Cmodct ~hclimlh 'OSCm;_llQrn; SIDW cb\lJ'n ~1C' I'hyQ1I1IlS dumlg w;i1king.rnld once silC!ll. t'fllpiill}! ro1C~iSC flilsh.'f ,~winmmi!!g rhy~lm'l~ ,rjltC 10 I~"C h~gJj~!ZrilllrHl';;'ii(; ~m(lWI~ill'C-i!.os.of !~l.;!~. ffiti~bllllN_ lll'il-oi (':;)In,d...rl
(;xplminl "vbay ~k1l1~nlU:lcr wOIlkiug ~i'n(])IJ1('il11C: do not o~t;rbp (3., J1,
the sab:rl'Wld~~f'mootto PI'OlJlC'I hself tb~""md ~n \\'<l!~CI: The bl~r.l!~'i!'ht1l~becn!cn~, ~~i!1!1i:~ar10 ~. lik~ly results fro!n .l CMlhi ~~,~u of lh!tlt; f1Lt.:'io~ ~hn~iJ of ahc s.o:lIJtqITl'<lIW!C'~. wilh j'XIi~u=-of Illinurn~.i di"r)"~fll(."n!. mwdi ~!g Jm~n h{;,ltl:f;) I[~il(f!~.4. '1~:lllN:'[l~ mooth,~" k'l\7i,rt;(: hqu,;;ncii(';.>;. rt.w~r I A <imd B. <lm~'\'$~,The cn\'c~Ol'K.' mro:inml :1 :l~" a.c'lIm~i,Jd jo:i 11K :md .. less Ill:oBk'd body ~lml1 end d:t~}Il1!.ct:;'n:u.;·m ~,"'~ ~!Irnll:}~ i(;ump~c;"l plOmc P: tt'('lJtlil_ Nom.:!h~~(;'%. consideri ng Ihe- rela-
hi~ld~i,l]fib~rdh:, This is ~~rob.llMy h'Ulg:c joi:nl. ;;(~ilh-c gi,tlJ~e arlO '~h,~ incn;:;;lM.'d !1l"~1I flf !hchill~~hmh ,!ll00lllk: ~irrl"Cl1"~~i1I!d i~I;I¢t;'~i'it;:Ii:I:... CC!!iI~i!s'Il;ltlwilh ~hn~'~ui~d~r k~ih.:W'i~tit:-;lind :EMG l\.':Cardi:ngs,.mi, 1i!1,t!iJIIJ 11I1iou w~\'eleu~.i!1Ilh of one 1xM:11' 1!t:I~~t~1 i ii' ~~:wiilla~ I'i~d C\ooC'J:1 when lI:ic n\"'1~i~i1.c:r r oscH· o I!I:ttoIH' h mooiified w:ith the drive, The s'lvim" ~nlin~\.'ckldtiJ!3 MIl£C n\)~n tI"<l"i IIi 0'.12. nil/s (O,.O~;100.14 boo)! h.'ill,g~ll;;:'S). re~m.iv,e terms, in Ih~ rob01 :i.whm: nm~e slowl), Ih,m P. ',L'(IMOii (Ikun 1(1.4 t(! ~ 2 lh!i1d)'! k~lg~k"A1). di~~-cn(:e .. The
~1Ilb,OV(: iiI"::
or
~~5
-~..-c---·\"?-=f'_6
res
Ih~ V('kK;ily .,nd mlJe Or~l.~t,u:~~l~l:b!!lly (.V,. In addutttmto :-;;l'liilllriLic:;. ill Il(L~!!Ignll);]Dt'('IlL'" the; t g.~~I~ IlrOOlI!l:'£ldb)1 ~ Ie wbo.1. :lnX: ~~ill1Iilar tolll.O,l>(; oi"' ,ji, re;;i .~~mf~iil:I-,(b. ~.rllhc wnlkirlg ~li,1 ",Fi,g. 3). tl:le bndy makc:::;, an. S-;.<;Impc:d Sialli!lhlg w;:we wi.ih ~odks <II.~lC gil:dles. Cn'i(eIOJloC~ :I.ucrnl dis.p~,OOL;"J,''iiCrrilS cotl1f:llil'!;,xi 10 b,h~dlireel~ml of muliml O'ig. 3(;'1 Me qlila~j1nii~\cly :l>imHru- Ib:dh,,; roooi.llilli
Pig, 2 8I1:ow:, al] 'C1JIaJI1!,plic w T:th a f>imlPlc Ib~' ,iI~c't'C<.t.Wof ~111;: 'C. ~h:: model ;ca!lL'l:adily clC'd~wiLh dri\ ~.t1\!lJl}!ly ,,~1I1{1 ;{lil~~i!lU;:llL'll~1n~~:d ~M~~~ l,L"i .lik(lly 1 \ .. QCIl;I,!!p"j gll,~ [il;"Cly 1.xil<Jvi!l,~ .tni!llid) IDld modll'~t~
~p'I:h.",,\liKC:n 1t11cn~I~('N~. wnllkill'lg Imm Ct6!o ~.2 I,ril; :i'\V'im:ll1lillikl1jm1lll1,,6 to 2,~ .~ h'lll1~e I:.: sflilamandcr Pff'll/fJde&1i utI/iNn. '1111 our IltCll!cl~'aIk~ ing.u1d ~""lI'imJn~1I£I-equ::l:lc[e:o mll,gc, ~'Ii\'Cly. I ~i'mn R2 to Oil I.~ and ~ro~1l 10 1.3 I,Ill;. The 0,9 in!!rl~1Si(;r~~lCtlCi~ oflll(; 111001::1hrn,vc been ~Ut.A'.d '(ltl~~lamj, tlIDSe ofnhc: n;;;11~'!3trumllld!.,1"tO ~:il. uth~ ro, w inihc~c:tJXjL~ limm.<j; ell: the 1'O'bm n:Kllm:s, A~l.Il0U,gJil runtg,lSwilhn
stJi'ucled fmm ,di:gilbeclvid:!lo 'rij~ld~· m:~~ Qf :1.8 rna rker by IPoinlt:S (b1acik dCits) dUlring ;ji c:ornPLer~ :!llPppin:g ,cyt:fle of one
cmL...,,---.......------,r------r--;=====::::;J
,........ RQ'bo~ 14 _
- .' - SlallaiTiMMf
0.06 Il11S "" 0,.34 lilIoof loo~irfJs)" S'qullIrll'.s lmik.aJle giil"dies. A. du~ ill th~ edrf!mity lDf ,~ limb
nc
or
Ihc sa~'lll'ml.deu.wilh fJniiltrl'liIi~ dh,«)laOOlbCli~lf;; cJn~c ~lJ~hc gjrdl~s, jnot'C t'llt"i:t tn,; hind li111Dgi.rdilc is IOC'II.wc !i:JS'C'I: 110 the tail fO'l"~hcroilOil). 'TIle' a~ bl lHadu b'l~gn." re~ull1lc Ilwl Gnilc ~lllD1lm~~dcrwitlll ~wo!.,,;\C~p{~jJIllS:The ~f'~hC' ro\li;ll i!5. bC:Jl~(I\'Cf iQ.~ "",']ull);:' k:n.W_h, \\!lltO"w,l'i.!h:;,l~P(lirth~:>al!ull'L11di.'f·~ ~,~ildld~c{~ IrD1l'i,Ull~~m.ighn..lind dK: h~td of l!l'l~ l mIJou .u~111k~""i.oIC ilHlC!J]11 Jispl~lttl]kiill:.~bct'l!uIiOC il. n
]rnn(ates ~e1 ,e-s'tiFl1lated foO'!: (QI't tut: Mmhlhe grooild, Thle horizQJ1Jlal lilles :show 'lfl~' O'II'~raU dir'ec~oo ,cnrroMerd ~awL m) S,:;lrn,~ JiliiE!asl,lD'e'ment withl 10 m~~ 001 thi! r;!'iool W~ = .2J!, '~bdty :: 0.0& 111/s. :: OJ); boo.~ ~~.i'!g~s). (<:) [nve~
(oolesp:mdfngl ro ~e
'!an
la~e~ill diSlpl'3{:emenlt5 101 the sail!t!~ooelf ~i'!d t~FQbQt h dartJl jXliint:5 aM e!'1T0f bars, ~Ij)rr,~ool
mam~
0-
n
HHI
ol~all
00
le;o
40
:20
ror t~
W' Ih~ tI~gesand stI:.lnrlilr.a Wiatii1Jf'5 ,til 5 sequences ill v,tlrimls wLOO~~ d sailamanmer and 2:5~enee",ror Ilhe rnoo~ (23'). See also 1lI111\l'ies;S1! aool 52. Ell rl'eante> body 1lE!l!ii(Jlh,
1418
REPORTS
~ively :simple ,d(!Sl.gn or the mhGI. its O\iCirnIi p~r1:O:rnmncc ~"Pl\WC:"l 1111 ckm\.'"i!l~" of ~~1'C'Si!~~ '!B}, ~:!:ll"I!~h;lr';s 1~:!lKl"~ ~~h[lvi"f, ht..;:ru] hunin~e;:[va be il]dw;cd ~l,Irin~ both w,ul"'iu!~; 1lnd :;;'!.""~Il~n~in!:l ~tprl,lyi[lg; ~l~ylllilby
rllc~cal d~ii\'c':l, Clw~'lm~c1'b and rjgl~l side:- of b ~]~~lixidy '['1'_1+ Such ,I I'!I1J;:'J1;h,u[rniSJli1iti. ,.yt;e. i~ rI'iI::~lI~wi'uh 'Ihe at:ti~ii't;:p~!'t(:n)~~ !('t lCU~o:;';l'~iJml or m.,ollffin:sti~cfii'cd dHrbig~~(,Cj,~ lul11il> ini 'mh~ llwilttlni~1g lililt'i!prey. ~SC(' (23) m~,od 11mO'oi'ueSJ,J The Imluci'cll k~Hd~~0 Ih~ 1'il~,lnwili1~rour p,~" 1 die~iQm;, {i) h P:I\o;.'ilie'l:> lll1ulllim'b oscH:lal.oNi> sat!lIr.:Iill:: :n lower tre'4rucm:ics ti~:im body OjCmawfS rth)'r('t,tll($i.~ 3 J, TIle ~!liurali.(!n COli kI be d ue d ll1.ef ~o II ~'hm~ l11oclmlll~n (i.e .. ~umb ruc1JI"ncm;, <i!l1tl inlfin~iC'.lj Iy limili.ldto k~'I'oC'F 1lci.'i!lluJ.:nt h;:',s)and/of ~Ou um;'Ch(m~~~lU~~ :L'tl!~cl!lllU~imd~~~\lIron$ i.e .• !he rt ~11L,,~..;t' !'!~'L1m~!B : iJ~uld ~!llllU1lI'!.c"mi!ling 't!I~ i~ mlli:I!OF.c~mllm!l:i I,g Ih!!.l~imb o.w~lr"llOt}i, if Ihe ,~iM!d eseeeds .. Ih~2'lhold,. r(ii)Ii:)'llOlh~.'li!'i 4 ~l'r(;dict;.. di1,~~r!IDIO!I!(.'II,I~(m :-lij~mds. 00' ~i!li~b• d .ul.' ax i:d ml~d!,!'; ... s.l'mu]:d (.·xhib:~1 ~ni rr~~r);llll ~ ull~aiml o
in~iOOtl.:. hI\.! OOC01.mt .md l~rotX)~iPl,giIIml, Il~ .;!(~ 1h;(i]lJc~dC's Ihl" the same ,tri ,Ie ':.... hen body d~~~nor {h~il!hln;my iXll1X.U ~ogcth~r wilh~hg Q~.il~l;nol'~ ,,,"~;ci59~m,kd frmlll lhn'b Q,~ci~hrt~. ~"'p;1ri!!l~n'!!~'l ,-.1~ow !hit> IlP(~~Uic'ionto h@ Wu~ (.. IN ~~lIX~i!ll;:;,ni"m imrirl"!l;: W'l~ ~~r.H~ilI'~ lb.i"'~I~~~ S~~timl " or lill,': Mllh;:nal'l ~[!ll\,'l;~1~1,;""_1~0![1~~ u) WIJ in !'\Pann! Olili;:m~mN ~:OI!lWrn;wK~ ~ ~~l('mjl: m",-dt(~~ ~miiS!~l r.,~r ille g.(,I~,,~c.;[tiun of lInllll:itip.k: gllih; il~ r~(LuCi. llll :>il1li!ar ttl iltl'; l!~111;lP.l'l:y.[lk'Oylll:lllCtril"1!l.1 d :>tullu~,"'ion Qflih!1!bmin ~lcl'l1!\/iliIiICild ~. hm~.i!1g '\it!!1,-tmJb,,~ Fi!i1~n}" !hiN 'WP.1i;. .ihio l,;o~himutCl< ~proiJouic.~. ~n !oiilhuu:;t!T!dc:u:N.(.h ~Wc r:~~-d tll;1I~"",>liL1iU1!i iet ng 11li;.'f(;.I~ el,llll;;i]I~)'rio 'M:.~~~,;ulr:!~i~hl,~ nll;::LI1t~~· thc!!~ur.l~p\!!ltlil\'I;'UYN IltHI] H!lliib (:'~nk:"~ 10 body o\log}' foj' cOIJlrollilig nlk~~oct1l'iilo[iM tl.f fObt1L~ CCl'iIIT~ W IIIIImodi fy (lie w111k ill!; gr. i! -I.h~ lIDd.)' WiEhriluhtll)l(! dC'~~ oITI\.~:tI~n. iii ~J:'[t1irukur[or will l.ell,tI to ilm'iKc la'il ...~linl.~wave.~, ::'I.lttl th~re wi] ~l~!I.1lL~:> ofooOFdlinl!liDiQfI Ix:l'IM.."Ci1 timll i11Diot:· ~~il SliC'ad,Y-:,,>lfIle IOOdl1WI·iml ifl rompl:oi:x ~\·iMt· I'IIlCl1.13!;,. 'CI,~Gs,offer Ilt1 iml~~ii1£ al)J~ro,LC~l lO mems.and boo)~mtLiubition: b'lIl, no!du) s.whn· :!ilI1\llm~ tillC' prob~el~l o.n~lnc IrajeclOI)! I:!,CtleUlltt{ll1r milliiJ ~nodc. TIu:' Im~'h~1~licrlJiml ort~li:i:,Sl!ldy IIIii'" H'~bl,<ltC 'by lII::1il'~~h..:~hail cycle h:IHlv!or '(ir CIlLlI,I\::d os-ci~I~(I1'!1 to :pr~lLCI!! Ilil!;;, m{ltrn' ooHwnmld." in :tt'.ru ~11Ot~0I~ i.~!{!I ::ti1O"" h~wn k!u3~1iOd I!oooJIlOtOOul Ihe oo:ruml,tcr com be buill rrom '~Oll of I~rirnili\~~ !im.;.,P(:i·b;Ii'iCd ('{lJ!ll![\oll aUO\'I.'l'one ~. ~dlJ(''I;.~ din]l;,'1:~i~l1>L!ily o,f~he ]oconlnt~u ~'O~I~d B)!])b!('H! :;.wj~nni!u~ CVLq;iuit I!!II(~ < ~-"pl!ain l!hl.)' ~~'!:.~l!l!~i~~t~ of whilL.' n..'1111cllillil1!jl ~.li!llhly Jil(,,1\iblc If) CD1'uin1l,1o!)!l.ly ~~. !F'oII~~;~iO:I1.th~ s.~'I;!i~t'h~t~\1,,\';" 'ltiw~~~,g urd ~j11~ ~'li.:ioci\y,di m;lim~. IIndWIlC or glIit ocooru" sWl'~ng \'I!jj\'el of l~ Wlndll!bllioll~ OIll'dIhe eo(Ip;Ji!!~l]t::Ug bciWt!!I.11~Y Dilld ~i~,tr.... his \II!OIk T ingll(:'l !he cn~ iro!l:r!W~l!).'!o1 O!1H!'d. O
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.:1:
E)'.;ampt~$ orr sympaiJric sj;!i~.iati'Olij, ~in mlUJr~ afle' rare lind hotly d~b~IIl\~d1,W~ d1e.~~rib,~ tlth~' Ipar.a,u~l ~pecii'IJtj,~1'I ~,~rinche.~ en two smli'llH i'>larful~ilij, the liri~talf11 ria Cu lliihraalr~hip ~la!!l~' i,l'l th,e, S®UI~'hI!di'ln'lii( O(Jeilll"n. Nl!.s{)spiza lilllmrtin.!DsilIl'lE!'[I ,d.allS1C E'i'.Yllitlp~e of <I sirnlpl.e ,i'lJdlaptiw Iradi<lti,on, wiithr l~'i.I~ ~ies sp min eaehisla rid: .1nI sbu ndant s l'I1IaU-b1Ued d'i~tary 91lmerra l i~1 and as:c,me' La rg)e-~iHe.d spedaltist.. lihe~r m;Orph[llogl~[a~.dfilltluity d~sell)l rnatches the ar';!'alitilli:lte!spec:tmm of seed sil!lM, and gel'let."
'8''I1'id!1II,l]e! lJ99@sts~hil~. tlil ey '8'volvscil indepen de rlt~lI'DI'I e,a;cih ~!iianld. SpE'r:iillt~ol1i scom,pl~E!te QI'I ~hl! 1 S sm'i:I~~F!ri!i~amldl, where there is;" sin!!il~ habitat wlti1 :stficnllgly llimod~l. s!1l!lld s;i2ii!!3lblL!riIti~mm,ll1l1!t is im:i!lm,~I~ete m~ UHl! I.ar9l!l!risi,.lrId, \lIherE! ,[I gn!a~E!r diWT~i~ ,of h,albit.ats has 're:!;tI~tedllr1~luee lineages. .a L!Ir .~t!!ldlysU'!J91e.SLs;that (he bl!lllltinlQ!5 have ulllide;rg,olW pilroll It~leco ~o9ric,i!l s,pe(iialkll1l.
:U1.d subseqsent
CXITI.I'I~)fcs W'e
dis:J)C'ffial (11.
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't J ~i}.(),I,I.!r' the 11'.,..,1, dr:crJ:cl;,;'.~noocls 113\,'c been , dcvclopoo )1\!.Uc~bng~lml ,"p'cimion can o(lI!)r::~ur ~h!l1Il1ghn:u!!I!ml 'or ~~XWI~ f,Clecl~ru! in 1~,IP;!!f)i o.r sympauy., \Ii, il'lII p<lrlkll or cO:lnpk:te O/\,'ClilJp be~~','CCfl 1'IO];lHII<1tt0I1S 144i)'. In!liill ~~;]jio:n is drivel] by IroqUC'llCy..aCPC!llk.·,m dii;Ull.Jlllivc selec~im1l. i~~hid! illdividulll nu'll.","" is; (k~~~rrnill'cc:Iby w ~h~ OC!nl~ilit:lI] ,n.f the roPlI~alnon Ihrml!!h 'Cfmril~ pelnciuJl. Ui", .ii:;; termed OIdruPI.U\'c: :;;~X"l:!milit:'m w' ~cUl'.."S d'L{; ~rnpoJJ1llltc~ ot biolog.icfl~ iilillcruCl.~Ql.1S I(.J), 3rrthouI~h adapl:ilvc precesses :dSCJ may reinforce ~gregatiOJlinl. ~1~OPfL~rit'f~lIydcrivro 1~~lCil&"CS I( 7, 8), :lEcolO£irnll sped aI.io.u is II si~ui~f.r process, whereby reprodeeuvc isolritil:m results from
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