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Science
SC~E.NCIEEXflRlESS
MOLECULAR BIIOlOGY (ollrlrol ofSt,re5s-IUJependlelFll Cardiillc GrO\l.I'[n ,8ImIIIGeme- [Expression

CONTENTS

.1

by a Mkr6U4A E. va~ ROfJif et ill.

An WiiRNIiI (odledl'.~:tJilgi'l al'1ill'toolll 01 a my!),» n gNile ~nJ!:reM~5 !h!1' mall.acl.aplhl'f

PHYSICS

N,ega;tive l'I:'e'rra'cl1olilat ViiSllllle Irir-eq uendes H. J tezec, ). A ()ioJ'l,ne, H. A. Atwaiell'

10.112 6I:sd@l'!cE'.::IlB9089
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10. 1 12.e:Js.dll'rlCil' .1.'13'92,56, h~Tn r(Hlghpu t Analyses Ma 1iI1~otill! Rl~$PQIi,~eof E. coN [

tv P'~rt!.!i'b'ii'ti!J(ns, N. Ishii,e't at
lin !l1li1ililtai nnn9 lQ~bdlk

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For authors who wish or need to spO~I~oropen

aoce$s~oU"lleir pulbli~hedl research an-ticles, YOiUrare

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Science
IR:EIPiQRlS
CONTINUED ...
CELL B~OLC)o(~Y fl.llilalbRity iill'1d N\(!i~e [)e,peflden~'e in II:)UfeliE:nili!l'l~On DYl!lall1!:1li(~ G~ M.. SueJ et at A 9~netk circuit h:lllrJ.acte.r~at 1l~11 Uel:lll'ltfi~ti:lJtII~xMDU15 a ,d1i 5Jurp,msliF1'!i~~ 'l{d!ri~d re~rto'ire 01 d¥namk ~np.omSM tlna:t de~nd I~eamou~t of noise ~I'I(he (Ompunelflt biOoChemkal reaction5.
INEUROSC~EN GiE

CONTENTS

.1

PrdYSICS, A!fl AtQIIlilI'C; Sel~C!w Swflt£hi formed Ib:f m~ilCI 1696 Asymrnifli'h~( l'!-G~IOi~~f$ Of! '" GI~ (OOl~ '$,Uifra(le S K. romatsu E:t a L O",ng1f!g tl1~ ~Q~i~~n ,q~tir'l <!w .... s i[)oorpO'!O!ioo il'l'i@ i!g~rilll!"f!il[lm ~urfaoo ~iik;'h~~ih@ ~IAl'JI;tr.it;;~l oond~,tivit)' ,EInI gr Q;ff .gI~~;lII}g llQkI~~n APPLIED PHYS~C8 Mar) !fIifying S'UF,lelrl_~I1~ ill th,e Vi$ible fre!!;lltH~~i!:YRalflg,~ 1699

171.6

en

I, I. .sm(l/yanin~.w. t:-j. HUrJ'g, C C D~v-.i~

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feli!l1porat Frll'lqUElf1 ity of 5ulbthres!i'l(l~d Osd'llat~(lI1s; Scah~s \l,Iit~ El1tillrhiiliiall Grid (,oeUPield Spacing t. M'. Gl~(,{l\ll'l~ E. A. lHlt f. frlJll~~It•.M, t. H{J$~rmo

1 119

1701 Tl(lpka lt\!irnca (l'iOI',eor ~as~ D'~ ladati(l,f11 the j. W. H. W,eijers. E. ScheiujJ. S, S(;!lOl1~'(!n,J. S, 5, D\t'l~)srt
IDuring deg Iadm'ion,. Wilrmiltig ur Irop'iGll Jlfuicll r~lmi\l\~ to th!l!' A.l.lilImk Oi!!Olaninm!l1Isro (he tillil!lh'I~<1 th,e:Ulfla~'9rfl,d1QmaM~Jhlls, oenlfill. Aflrncan r.QJifilal~ GEOLOGY PI Ve~~ng~ o~ IEarth 's Oldes~ OpMo~fJle 1704 Ii ttJmes et ~I, IR:em naRiS,i)1 oceanic. cw'Stlormed at,iiI s(IYH!'<I.dirng (~nter 3 • .8; Ilmiclil ~ears, ago a re P rfi!ilrYed in Gfiet;J1J'l<Jindl. imll~yliJ1liQ th!at ~me h:mlil 01 p~atet«~Clnic5 '~a5, aopoemtijlngIhen. :O!i> ~fW~ "~ p. 165t'l ATMOS 1'19~.R I.e SCIE NCE. ~on:om·1J p[)l~te!rlll'rinatl o:n of .Ain.:5ea .M.mn!!IIl~L1m 1707 Ex('hal1ge lIindera .Mar o,r 1in)Il~(a l Cy<I.o.ne f. J(HOS~ ,D, A. M'fr'heN. D. 1M W(mg, W.). reO£lue mfiut gD.s:efijla~ioI!l5r>eVea l that 'tl'ie' 't!f~;/'j~fer <OIr ii'IiIome;nllJm b~ winds
1:0 OCN!!I
CU~fi!Jlt~·O!i'I-iflI ~

CLIMATE CHANGE (O~ F,ll~d 1 h@iFi]'IiJall Q.,.dlHy~~l()gi(a ~IEvl'Jil~,.ltij(J:n of

A$ faU IDD'>iea'!:lDut, til!J1 oS[ill<li~iol!l hE!Qjul!m:jl!'S of (ortii:<lI. ooumn~ alrrolN,BdIin dl 9!'illl1!'prM~nl ho.w nelJralactivily ma:P5 ~he rat·51l.D~tiCln. l ilil SflM!2.
NEURoscrEN ell!

IEl!I1I!ergellu:eo.~INiW:e~.Co' lor U"hioni 1'1 ,Micill '[l23 IEIiIgim1,ee.redo IExplI"E!'li:!iill 1HL!I1iIlIi! t nGtme Photopigrnel'lt. G. H. Jambs, G, A. W1:Wl1ms, H. (aMi. J, NatRI{JfJS Mice M,ginMr~cI to !f.!q)nlS'5 .tI'I~tliJ~n lDng·~a.... el!l'n!ltJ;j o~iII in ad'(jiticm 110 it~ Gillin I!'.iO (io.'lor v~iorn pigrrtent!! Ol()Qjuinl' ol 1'!11!~ alOO litf
to dis:tijiJguish colors.

srs

UCTURAl. BIOLO Civ

(rYfitalStnJc:hm~s of the ,Adjen~lat€' Senrsllor ~ro:m

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m

This Week f'n SCierKf'

~)liIlring a I'l j I11ll!l1lL1ne .reCiidion" T (el ~~divid e r .apirll'y .lind d iU,eorelilli,i:llleprodlLllrl ng iii VB riety or T a:eU lypl'~ ~hal, r,e-s,poml ~P,Il:f(lpr-i.alt~lylo t~,~ pan~cularthf@2!t M@lffilory I (@H~.aIsQ @:m@fg~ hQmth~ $Cl11iIl.€ IIlOPli~,Qlionand remain in th e b od'y 'I,IllIlil sUl~hlime as a ne~ inf,e<_tion (i!U5.on ce ag~alirl lor their a~tenlioll. i(llilallil:g e:l,ut (:p. 1687. pu btl'S~i!d c:nlT1'1~ llM.alr~h: r.e@ l~E!! P\!r$p'~:tlVI!:! by L:iUJflI!~rlali1d 511'1911} rno,\l,I sh~w ~ha't sirl£ll~ li cells I,IIlwI~:rgiO,,an ilm!ilia~aiSyrrnlil1el,ri( (,en divLS,iO:!l il'!l resp0l15'~ 10 a palillogen, produ~in~ Itwo dau grl'ter (eillsl,l'ilil~ "tterlfl~te rates. ,After f:(~rl'lijilf!gj~i'I iiniill!i ~e :sy~a,p~~ wil'il an alfltigiEH'I· pre s.e IilLing ceU (A'PO, va dow; priGtei'll~, ~ndud iog '50flGL'1' re5plllilsilJle- For s~gl'I\lllin 9 and' tlsymlllu~trk 'celhl~'{j,si~!h wer~ ~"'Qri~l1tedwjtltiilll the llc.etl::i, AJu.!~ dil!ij~l(!!FI, dalDt;lhtefs th~t ~,!ilf;:f~ I)ro~ima~ t~ the AIP'C:-1i' cell s.yna pSI:! became eH,edo:r cells, \!1111~ their dl sta ~ )i'5,I€f!> bera I'I1N~fII1Iore mem o:ry-li Ile .allld le ",b11:! to (Qrr~!il,rIbetter pratl:!(,IJOn: ,~h!:ll'!lrallllsffil ~red to mice.

Division D'edsJ arts

WhllO I~ nsplres bra~ n'w',oves 'w'hi~('eI~ studywater waves,?

'~OOeli(~lelllll!nt$ via Ii! Nfl! iI'lleJ'(eJ'El ce. D lIriillg ~h!!: filii tlJrall-;P~I'l@ra't~oUil ph~'gi!-re:sisHIPU Stre:ptoccccus: n or ~/1E!rmopMt1s '19 lytic pha geob~a in eel hom yogu rl,. B,ura D'£I 011 ~/' li't. (p. 1 709'; s~e the lliewss.lolY by Mil rx) 110'ulntlthat!hl? ~nlcgi1t'~on iJ~' vira'ls.eq ue n(l!s as new spacc.:~ Into (1RIIS·PR klt:i ind~E!d'C.OIi,r·~rs. ~mmu dyag:a irlS:~.v~rule lilt ph.agle) ina specUk. aCQui red, and iIIerH.ab ~e l!I1il nner, AddlltjoJil and d,~I,!ltjQ~ or spacers all>el$ :sensil'ivurv to Vi~!,liSes., nd CRIS[iliR-as:S(ldgl~dl '~e:nes rna)' be diredl..y ill'lYQlv,oo ill! e ~heIr~!J5italllC:e II'IU1Cihllllli'SI1li.

Reg~!,l~a!fdl.k!H.er:s ,of reI) ea IS sep.a r ate-dl by ~pg{e~s (llf ~fI'Iit~~ length (OUSPiR) are li\Iid e~.yd~s'~lriblltedh'! ~ih~ gem) m~s cf li3acl@fia andl Ard;:II:!a. ,and a r~ dl Sltil!1(lllJtll.y hyperva ~ijablE!.n~i! ~p.a(E!rs sha re' seq uemE! ~o,ool.ogy with bal:~,eriol1h a:ge i!I nd ptilsmill:l :5'~IHm(€S,8inti may !H(lvideilll'li'n Illlliity <lilai nst for,~ig n

The' Fwna[ Crunch

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Stayinqin Charqe
Ulnd@f 1I1I'~rl11l.a'l 't:ond~tjens, l~@ifill tracE!'llllJlair ,~tDIl!Uln,uation o~' ~Jduo~m~ tfLI'llil05 pbte (AlP) is Olil l~e order ~.f I mi Ui· mOl.flf.ll'/JumlllftlL!ls I!lrl,~ylllle1.5and nlgl!'~t[lry ,prol.llins ~iI!~yan this ~'!!I1vE'r:sal c.urrel'llC.yof energy ror 31'!"boli!t:. ULalbo'[ic, ilnd g~n.Eirat h(HjS~eeping iJ'fQC'~:S~:S. Crill ~f the prim~i)i
elllll'l'I1Ie!>

I study the m;a,themati(;aJl squatlonsjhat deo5crri,lie tli1e matlcn of w;a,t)e~'w~!.Ies. lD~ff€rent equatiens represent dlifferent W1Bves ~ w·aves comfliiil 0 nto a bea.cll" wav~s in iii pu:d:dll!i!" or 'l;\IiiltV!!S ~nya'~H bathUJlb. Then whenll"ve SlJinFeiCiI tme msth, I ll k@ 'I1o:thi ng beU!i!r than l105tpend the ~e:5tof ttl e day S~ r~ rig ithe w.aves, lhis~eki is wry ili'illPortoJ!Ilt The beUer

Ihall

N~g.U

lillteos ATIP' levels 1S the- AMp..a(~i\l'a[ed

pW'teilPil khH,!~e- (Ai!!'IPtQ. whl(;n ~efrl~e~ the lfl:l1!!tiveo ri!!iQ

of AlP ~o .AMP;. W~en ihis ra~ioraU!>, Alil!.PIK ~h,o~phoryl.ales PIl~tabIllH( enzYln~'~, whil:h then censume less Aff and mike I1IIOf<!~or lit T'C'\M!:Itey a,lIId' Shl<lipiWlIJ' 1l726 .•j:llJb· (p. Ushed (llilli ~e 8 Feb:rua ry; see ithe PeISIPe'<!i lie by 1111 (I ~die) hiJl\i'e' ~[ll!{edllh~ slim (In.H~ oflhl1 Hssion yl'1ast AMPiK I1mnotog dnd ~e'ln4)~~l~g ~he cem petit ~ve,bf!fldifl'!J orA]!IP' ~f1'd A~ I" Cit a 5;j ilg l~ ~ l'lIudeolide s~Le. I,'IIhere U1ea bsen (e of (olll'lle'rion s a ppears, to a I!I1pl.ify the disClril1l1na licm 1:IE~lween I.ne FIlIOflO-and lrillJilosphlte ~.'r~and~.

.we: earn 11110 d'el wat~~ waves, the beuer we ta!ll predicHhe paitllrns of -.. beach ereslen Clnd natural diS03J$:te:rs.
B.eLng a mern be r off AMS mssns I R,el to ie-am ~lboL!t ·i!lreas or Tliiiter~slil lmight net a.th@'rwise !E!t1courr!:er~ II. giv@s me 'Wli~u;able opp'LI'rtul'li~ies

to e)l!chi1li'lg,eideas wUI1 eellea!l:UIE~s other~eld5. AncHhis he~P5 in

!1fleflnd new ·aWf1O~ches to !fiV ",
0WiI1 work,

The hOrl!'fiQ:ne a b5dska(id (A,(l.Mreg u!~ ies ~ v(lri ety Qr dev~~opmenit.;l ~a~d physiQkig ka l ()roce~'~ s in ~! [~'he:rI)~a(:!ts, L:i ~ IItt t11.• (p•.ll712. p~ b us~'ed oll1llfi'!1€: 8 MaJC~; :s~~ilI,fit er~pfit'~ilf~~~ GriUiill!ild ChJri:iU~ P
mJ<lliIIdhaw 11Ow~de nlified ,[I membrane -hoend pro~~i'lilUrn,[I,trllncliam il,S ailll ABA re(f ptOI'. The Il1m~,e'in. GCFl2. hal) r~a~JlIlf'@:S or C!IG prot£: V!iI,.-!W1iI pl.ed receptor. ~hidi ~E: th I) usa rids of va,riarns in afl imaJ h 'Lells, b'L.ll~. uy few Iknown ltaIrEallits in pl:afb~.oells, '!Ie

Plant Hormone Sillgna[in,g Receptor

D~. IKii th€rine Socha is an a s<sis[arJ t professor ,of mClthem<:l'lics, <It SL Mary's «(llh~g'e' ,.Ma ry~an dI.Silets a~$;O' a marn b,~r M AAAS ..
S~'11!""ideo C lips of rhls story alld others 1:11 www.'HliiI!t.crg/stol'ies

.~ U.E1 1'1III( r~tr po~e lQomp ~eoj(IN!PO (onlrc'[s thl! e,;(('hi1lI19~or m{l~emle5 b l'~l,'!,Ieli1l'1 (ytopl.i3sm am! liu! ( the ~udelil!l, Thit$.$ upr,il!ll!lwleci..ilalr a$;~ lili!bly ~s(!lim POiS~t10~ ;:i !i:E:t of proteins termedill! ~cle ~P'OriF!s (l1.lJPS), E! ,Mel~liiik til'. (p. 172.9) dl1!!;criibe t he~lruc:lme 01 i!I mrnpl.ex, of nupS 13; i!I n d ml p4 5., whkl1are eS5.enllat (olil1lIDr'].E~~l." Qi( '~he (J;!r1irall (:llal1ne~of lh~ NP" Th ~ lWQ !Tn l!J(le,opmim form ;:it.abl.e d imer~ ~hal (1,1 riher a<~Q,ia'~e J,rII,Ql>etrallllll'S. Two c!),stal, ~·Qrllllsconl.ained four HJnr~lrrm~f~ or the tetramer.l;iini,h d]ner~d ~ ]1'1 l~ eo~ater al onset beh~~,1l d il1!1ers. WhUi5" th e5Jf: ffill,J de,opi)r;l!l~ IiJla,Yhal;\(' dYl1!'llrl]k iIllteuac iion lffi1ler'~ [ac;es ... nd5ilid~ rel!J'tlvt la elt(h GJ:therun cmr~r kI adjll!f>~'~llul'dlal1let~r Qr~he tftlllSp~rl ch ..m~e'l to thll' .~ :s,ize g:~the carglo.

Diameter Modulators
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of trn'E! Nudear

POH~

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1635

Looking for solid ground in the ever-changing landscape ofscience & technology policy and budget issue's?
loin the natlon's tOIP SA.T experts at the' 32nJd Annual AAAS Forum on Science It. Te'chnolol:gy IPoUcy :3 :4 ,M.,ay :2007 ·W,ils,lhington DC
lntamatlonal 'Tra,deCenterln the

R.Or1I,~dd IRea,garn B!uHd~nl,g

The AA.AS l'onJ m on ScierKe an d Tech nology :Polky p!',ovides for issues f~d ngthe sci enee, engi!1 eeri't1lg, end higlltf~r ed ucation nifties. Infrtiat:ed tn '1976 as~he MAS
8! seUlng

d lscusslon a nd d,e'ba~e ebout the f-ederal blll dget a nd other po,licy

(0111!ili! 1.1"

ssn (oUo,quilHlI11

wi~11aJ!Jout tOO

partie ~pa1'11:5, the Fo ~um has, emerged as tb eomalor publitc meeUng tin the 1II.S. d@\il)Md >tos6e'nce ai'ild~e( ni1lo~Ggy polley issues. II. a r'li1IUaHy' dI~;;!IWSupwa ~ds oif 500 of
'I

UM~' natio

n's, to,p' S&, r policy exp erts,

'I

G,e~a. 'ful~ alt"liElIvsi5 of the Preside ntis fed era! ~ R&I[) fu.ndin,g pwposals, Hav@' an OPPOfUj,nfrlly' t.o meet dhectly with kiey S&r ,po,H cyma'ker5"

lI!

• Lea m how the changes in CongJfess a'!I~eai:ecti,t'1lg S&.T po~itCy lssues,

'I

Regisr;r .l1111;swi II reC~ iV@"at the ~fionml. AA5 Repo'rt A XXXI'~ R'eSfE.rlfrh OFid Delle/Dip· men,r~ FY2Q{)S~ a wmpr,J;,nen· sive a:naly5,~sof the proposals, for the Flf 2008 budget" prepared by.AAAS end a group, CI! fits affilliated Sde ntl fie, engine'erilng., and higher eodtll(,at! II) III asseclation s,

N etwe uk with co I ~eagu es, including top de<cisdonmakers in sclence ,alr'i:(i 'techno,I,ogy policy from all. 5eC~OI'S.

for more eemalete deteils on

t.he p!',ogrram, hotel r,egfstration <lind on-line Ir,egist1raUOIr'i, pleli}s~ visit II:h~websH!€:

• Learn about hroader n2l1ional and lntemanonal deveklp· merits that: wm afrle'c~$trat~· gltC plarmiiWlg in unlverslfles, ind U'stri~ s.and goverr nme nt

www..a.iillas.o~rlfun.Jm.

www.aaas.org/forurn

-:,D

Catherine' 'CE:s,ar:skJ' 'is dire~~or 9J~e rat of th'il'

lEur(;lIX'<Jll! $Qulihem

Ob~IWIWI)' ..nd pr,~s'l~liil of Ih:e IlnmrlililHoi'l!ll Astrom.rnka!, U nJQIII.

AT A TIME WIHUIjIUIIE POUTKAl VI'~mN OF EUROIF'IE 1.5 5rR!LJGoG,~jING TO REoGA,~NlfOCUS ANII!) n'iLgme!~l\lm, ~lirnp~<il~ E <!:s~rOi~n:I!L~~n;are tvorki~g Lp,gl;'tlile-rmor~ [h~~u,t;.\iJ,;jf on n~w projecrs, Despile t~e fm;! thiQ~ m~dl:tl!llJl:si:l'oI1Qmy 1;1"'1:I,1;1;fgwlr{lmi'~eE~~mpe.m R'i::lfgili'Slj<lfl~~.the om1!li~ Uil,eni'slead 'cru~;ul!l~ed i.UD,the 20th oel'll1\ry. Al tb~t time, visionary ~,~,h1:dsin t[lC United, St.nos exph):it1;d hnye eccnemic thnml.-e5 ~llC(m~tnu,;'~ ~Je,u: ohs-er\ .J:I{lIr1:fi:!) that op!,;:U1leda g,leW Willldu\7I,' onto rhe di;!;f{!j,1liI nl.wrse. IEumpc;:'. treco\'~:ring from rhe \\:,\'ll'ild wars., was unable m CODllpotl:l wl1(')11 u tliael.ha ired S\;J~ ~~mIil,ched, its O'w;:ir\;\lln:II1,~ing~y OOffiPfd'1.I;.m:5ive space progrml11i hl. til~e I95lh ~ mid ]960s. But tm:k~y, the Anner.i.C'.nm, <1lldEuropeall silmuims .in <IS-tro:l,~.o\L~.ly lII10I.'e balanced are and El~!\lr~ml ~"m]l'!(1111~~I(]{]j;; lO\~!<~rd fl!ll!:r~~)j,Ii~]~enewed (]Jll~!ni~,~l. 'thllUgb dKWare \Mtl~ll the r even m'\,'l!Iliet~l'U. given the ~'C!iSl projects :Jiml the <!'l;iali lable 11und!;, anl eheiees \,,ri:~~ave to be made, of h h OEil.i;;imp-orWC1t rea,lll'll.l for rhe EU~lp~~n r~~lrg~l1:t,v is t~~ut n.~i,~hI",,'_!;.ciem~~lcm~d, j)o;li1tt.~.,l : lcaders: cre,lted the EUlro:~~3n Somhern ObSCI"WlltCIIi"y (ESOl' ~In 1.9611.0 d~V{;~[('ip [lbs,~U'va~ol'i~ in I~(.'So1ill~]~I,.,"Qli1~ini!li~hErc. SOimi~~wh~11W!;;:r,. the
F

El~mpemll Sp.u:e .I-\gency U~S,t\I e.Hlbl.l.ri<:edon <11 \!te~I~pl.lD1ned P.fIlgF'<'I!Ill f o I!':r~~~mis.s.io!lS, AnimP'l-)J!'lUIQ1~ ~ldv"IIHi~~ f'~hasei !~i~rgf,)\I~ 0 rnmental o:rganizHtli,o:ll1i!S dun they CiHll, rely Qilll a st,llble budget, year ;<'I,ner ear, is, y De.:::plte iI budget less thl~n on:;:~llml t~:mt ofLhe U.S. N3idnm'l.l Ael'Qnmnic;s and Space Adminis~Ir<bTion ,~NASA). ESA l!lu5!lalvl'Iiched sU(.>ces.:SflUI m.~~ siens with weil-chesen goals, such a:'>I~.c Hippm.'iCos satellite :'lind the [nfi:lIred Space Ol):serv;;l~ory" On !heg,ruund ESO's n~s]aip. the Vety L."II'g-cT~kscope' i.~. Chile. has ~oh~cwd rocog~i[iDIl ~5; the \\ro~jkrs most pu\\;('l"!ru:~ flpl~,cO'lII~I~"(lpl;l:. oo"'id~!~ga \\I'e-ilnh of dm~ en obje;~[s!'ai~1;h'!g p Imlf1.1. tihe solar sy;slellill ~.oDh.e tarlile$l r-e'.!ItClmes, tile' lLI'Iiverse. As a :result.. of ESO has ;nl:~u;:tedn~wnm(;;m~rstr!~~.wi~11Ct~~:;(i:qu~r!li.:t~ i~~cl"lSf!~$. l~l;; in lim,Jige[ .<rmJ~.1lJcol,iabQ{f.-ating imi!S~.itutions... ,Co:l:lsidernhle hUIlUl'mi.ell:clun.g!e and , iiiel.;;\'Otl:.iil'ig., of'~~]jJfo~tered by th~ EU~OP!li!'~ifll U~klll {EU). l1a"ii;:l'esul~d in several bil:alernl Of m.l.llt~,H-entltlltd)~nak~Jn,gs '!."ithin and beyond Europe, Theresult :lilIOb been an ~lII.~~i'lins. com~ gf Wtiltinn tbat has CI'lhaj~ood rolhcrdl~Ul! hi.ncl~u:ed, ti~acro:5s I.hoc Atla:UiIlbc.ESf\ ¥1l.'b.<;; \V(J:rkod, iu long ,~lmu;~rship 'iMLthNASl.A.lalilhcmghi lin: rd<l~iomiJ,lip W,L'l not .ahv<'.)":l>ea::>y i. On the gmm~~. EmDpc. North Am~rkfu. and l~i&\ f\l)itl fl~ 1lt)\",1ii1Jvolvoo, ln CQllstfUCril!li!:[l: ALMA. a \.~!(]rld-cli));~s ,;urn;~jiof ~c1:es!""Xlpe',';~oe):lp~ore cold UIlUVel'llf. ESO is, [e~d~1:Ugdie EmiQiJe<l.n (!:nblits.p~l.nill1g [1's"e1'lp;e¥ti,~e the i~mi. mml11ging~arge ..un:frnstru.c[liJIfe project>; m the dil~pOS111 of:sul:):n[i:~~,irn.eleli raduo 1!S~rnlunl1y. Eurnp~nl ;;k.~lVCmorll!yi:-:; 11["'' ' (.~,m~id~nirigfiL!hll'e., [StY:, c(l1'lJ~mnrgil)irn.lz.; l":'wbl~shed"m ;JlT!I.l[l.&its tiOUI.!;. ~l1i\'Cn~ry of scicnnfle opporHIJl:ilies imi.s~uoe ~lSm:m.Qi.mi]y : ami.d. lmU!lOI!iJgy in the :1:0 1.5-102:5, p ~ime frume" "_Inetli ,.. 'osm ie Vi!l~GI.1 ;"I!SA has jill$! issued ae'liJl! finr propoS1l1~flO~d'1e pR1g;r.m~,·l<. rUfl'~: SpilCC c.x1)!!nrl!tlli.I"JI'~l\)JiAAiO!u. ESOaJmid its. ~otnmu:mi.~ly ~lr1;: eonduc~.hlg &Iudies on the E~lropcl!ll Extre!Yi;Iil~Y Lu!"§ T~I~~.,a ~1(i" ~ de.~igij1i:Jrh~ world'~ h.!l'gl;-i;~ pli.C'aI!illilhu~d h,~I~;.!~:.'Op~,.whkh l;: t o willlllievolutuoniz:egmulld=hased astrol.ull.ny.~ts tm'ge.t lor cO!lm:s~ructkU]is 201.0. ELilropean ro:llrlio
~1,.,,;,!fonl;lln~~.bl;]o[}~il;o~1by EU,~lpi;l'S L{¥\,\! Frol\:l,W!~I.'!C)·· Ami}' ~;r~~~cl,are h~wib!h~\}]v~-.Ji!llf~l~ing II l.inure v,lQdd\\ideprnjocl. tll.ll:: SqUMe K.i~(}Jneter Arnuy {~O be ~ile{l in A,usurnl:ia or South Altic<lJ. The high (.'o::-ot ~m;h ile;\,'proji~~.'l!"KIth!;) ill1!h(l·rn.lliry 'Of ~d!l!irlC'~. l);;u~le fm.luWlr.I.§W 'l!.'inIOrlOln,""l~ yf

frollll,bmh sides of the Ad,,"tic~Qlbml p.utm:rs..hipswith t'i.SlroIIilO:l1.1fl'S fi'olllodn::r cnminents ofdu;l 'wolJ'1,c:~,3ndfllrl thaI is HlItl1i:r );tlrport~11 by D~e~~'ternm iOrl:Fli1 AS'l:ml'lomieal 'Un.irn~, To achie~le evelil gre,ner uniuy. the fllnd~J1!l ,<Ig.eillll.c.i:es f varinus E'~lrupem] counmes bave come o ~O§~thl!r\Ii;!ithESO and [SA inan EU·'$pm)~or.~d Mt~\'tlrl.ASTRONET, to ~!OO~iKha ro.w map for the next .2:0years Ihal wi mil enm.lilllpaSS iIE SO and! ES A pmljl,l'OIUlfiIIS aud fb.stelr cellahorations muong :EuruIle:,jJ~counlr.~e:;in other ell~deavorn.M ..jQJ;,org;mihl.~iolls stu:'hl ~sE'SA3nd ESO are 'Ib!e: 10 pool mtclliTet,ril1 cr:itie,d :Iill.u.rs~. with re.s.pect lO boll~,h~I~11fl'ml;"nd ~iil1mlci~.1 resources, o,\!e~' lau1S ~riotr~, onione, Ofe~llra:~ iI~n:mlec: L~[11m. as j:'J'ublicOOdies., they mLII1i<!. serve tl~e (loJmnuuni.ly. Tlris p:~(n.\'iJdes, po\"'t:rf~~1 impe'tus ~or exeellenee irl. !heways ilii '!,i\;hich they arenite, ltalse Oi. ensures constara upgr~wli 11g of flJlcl Htictl la rernai n c:o:~~.pciit ive, [f IEl!:I,fOPC '50glo b.:.1J~. ~e~.d a i~.c:bJcs k~""ri.flg""Sl!i:1Jl(lmy:n~l~e ; for..:foont !llf!,ifl~ili.~i~glh~~u~ky flfEuru~(m o.[g'lnil.;nijnf!!~ ~ud IHi ::;s,iOlI:1S wi II be vital dlei:r

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23 MARCH

2007

EDITED BY GIllBERT CH~N ,AlND JAKE YESTON

IDi,amo nd D,tv'r!lrsfty
1I'he cbemhtry of diai'i'l(l'i'iI~h bir<iIii!;jM I.Ip h'f.I'~ 'E,iUth'5 rna nHe-n eta hI!!, t iii ai r \vi de~v ra Im'giing l!1iitrog~11! ,C(lPiite"n5~f!d rlitIN:igjeF!!1!fld (,iJfb'i:! III isctope valllUes-has enrn p Itc:aled U IiId,ernt ami] I'Igt o,f their ,o~t1Yli~s. It is comllfilo'!'iJlf li,hoUl~ht,that: !I!llaf!y d]i'!l'I1Ionds fiormlfrom HiH! fi11Iovem'ellit of CiIII"DI)nIridil 'flui~lsii!ilto deep 1ln111lrltiero(k~ o,~a con'U'a'>t]rlg ,c:ornpo,siUon, UH~f'e'b:yindlldl1g dii:lmlllJ:ndpr!!~ dpi[ati,~~. To beUe~ (on:O;;U'i3i~,nl U,ese Uuids ,i3Jn,~ :5mHre~. Tilomilsliol et,t],t studlie.dil'l detllilllfle.i3irly 60 d'idllll1lofild~ COl1ltCililUld \vilhiilfl ol'le smaU (..::3:0 (ml)1 liliI.anUe :;ol!m,l'll~ CaH~i!dllo tine SUlI'f.:!(@ hll ol! lkillillbe.rti~e vokalllo lin South Afil'ka. SI!Hprhin~,ly, till!! li1itr~'91!!1'i~QnU!l'it:s· al1di:S!ClbJ,picvai,~e~ o~ UH!s! di~lfIfIeftidis jim! 'lhtls O'I'Ii!~ill!r'lpL! :~iJ!aftllne,d~ taifge P,Ul or thf ralnges ' Obi5ie~,drIf'OIn[lll dlii3if'!lloliild~w~rldwide. Tlll'e (alv.;;llriation$' or tn'~ do!!~a i1lI1pLy 'Ula'~ t~'~$e di!!l m,(Dml~r:~rm~d r,r®lII1li3irnetlha iii!'~. !fi'ch 'flll!id. !'Iota ml1ll11[t Q:):idh:ed n~~di !(;Q~lmo'flb' aSiSltliili1~dI. as: wide !,I'alriati~I'ICiillrl b,e'~,rodi!Jced I~y~h~ hdhll'l.ilti~!'I,C!f' IIlfilmg!iil'lii .midl carbon during '21rm'ltlh Q~' the diialililomas '(ll{<E!rirae, Such HuiiJ1> ma,y alse <Ii«:QI!.I1111 t f(lr~he v"lrillID~e 'iJ);,idaitliOin state ,of the !ili!1~m~t~ ~1i'Q!~"th, Ea rth' s most arl( ii!!!1t. (!'l!!st ~ 113I H

n~

I!IU)C H UI'U:5

ntr

Adivatin'g En Pa.ssa.lIlIt.
tll!ItCDtranspoftE!r~a re<a. ralnl~~r of had~fiall viruhmce PlIOte;f'I s U'q,aJlare Rr:$~tli'3i'i~\oca~Ed across l~~e iMer membr!1!l1~ and then i!lmt~ inw the Qut!er membrane, liful!f~5T1lQUer C.;tM:!rtTllinall dOI'll"l n a·dll'pt~ CI ~ ba rr'e1,~~ru!;t1lre ~lat s ~1I11~!~eouter IH\ll1librane ~ ilnd~r1te-s as a na~:5!~II'i'all' IO'I th e largeu N·lifirR11· nsl "pl3~~nge(' d~lIIlillin, ~\Ihkh is lramplllrted ~h.rollgh U]e barrel: s.mne p.<I '>Sl"rlger do:ma~rnllre! ~!eas~d qlitW'th~ e)jtra~elllo!la~5pa.oo by p'roleo!ly.si!l.. 11'1 n ~xt.ern;r\l'·e ~@rles ofgelfl'~tkaml a btoclu~,m· ~c::ll~)l!il~ri!1'!~.rUs. D.i!mu~er oj. sh~w ~Ilil~ pas~htl s~n£le r domaill'l or f.Sd.rf!rifliin fOJiElutDl:r ,ms,porler IE~pP is rl<l':(!ve<i if! a 11lJm,h~!,Ia~'r(il~hi)ol'lc lfI~t by ~ IlEt,nllpILa!lIll1K or illite, membrane' ]llIrQt~al~',but bJt it~~lf" ~nZYIl\il:tk hydrolysi~ '(i!~ i1I p.>ep'tide b~nd is cU5to:miuilyinmatoo by an a1:Jill~ted I]ud.eopilile.

in9 m~(ih!l)ll!isnl appealrs 'to ~ ,uUli2@Cil :tw Ii);tlh~f al!ltotr,[IrI~poorl:er~ as. \~eU a5 bV el!k~ryo<lk 'IIim~e~ dlllrilm'lJ c.1p5id 1'I1I~t.UlrClliQn. GjC EM!l'O J. 2)6, :lUlI03 B/sj.emb(J].76[J:l!.638 (Zl)on.
Mil (RQI~ ~'O W'GY

At tlh e :P~i nt '0,. ,Alta ck

Jub ereulesis kilL; 'lI\Pp'ro.:liil1lale.~'y3 mmiofl pe'op l~ eec nyeiR ifneol!)l'!Itho.g€iTnic ag ent Mycob.(l'('le.rlu.m iu~{(ulo1i5~m'!l,1Ildesalld rell~ki!tes. wilhin Ina crop:ilag,e's, ,c'onsMuctililt;l i~li)ritSl!'lt~' a n j ima c~llularvsruole thaL ~heUers it hmnirmllune '~Ulr· 'veiUafin{'E!'and aUad:. Alteri!!'f tJi. ~Ia\i'eilwe$~j~ gatedl ~orw' tuberwl&i5 blnd's, to ~lnd irw.ac!@) M.

lh ~ Sf! roil or Ill,b@ F(!J!m5is, ~,ti!nts mntai r~@dal'1ti-' bm;liE,>S.thal rec:og6l!il:ed t11e' pilill'l $ul:ll!Hlit Till:!' 11fi:tHUicil!la!l!d id!~n't~~l(atiol1 \\!hal may r'@pr@" ,Ilr ;;lent a Jiiey ~roH~iiil ii'll ~Ile .e':uly 5Lag;e·~of 1l05t (oloi'ii~alio!'l b:y hi, wbewwiosis !iIiIay lead lO the develolPtmel'll o'll'I~w thera.p],E!'s and ViJ(c·ines. - :SMH' 1',''', ~(!,!t .4c(!d, 5d U.S.A. 104" 5145 (lCG7).
(H!EI!Il,~STU

51Otlftii ntg :Siti,O r,ag'f 0 pili on s

~ ~ !Olorage' ror vehide applkaliomi, Hght 1111.121.<11. ~ hydriJde Ullfnp,oufilds sueh a~ UBH~ ... ~e all1p~.ilI.inSi ~ b@(lUSE! (If th!i r Il'ililm \'IoQ'ig h'l. pe rcimtlllg,e ilf ~ o:! l1ydr[l~erl, HiOW'ever, lil'!any .~uc:h :iii Clll:rlilpo lmd''5 ,ar<l~ '~l1eni1odyl:! sm icaUy ~ qlJl~t.e!>talbte, andsu Irequir~ e;«(f'5si'V~ ~~lillg lo IUb@!'a,~e' H.~, ne O me!iUk1i {Dfa~dre'Sl>illlg~hf~ problem I J!! lh e 'S1~.arch rOJ g p.r,adi cal nil odl!ll oir hrll'Qlwgu

p(il~nlia I h<ls,t oe~,l~. On

the SlU rfac@' Qr Ul!e' rlliicrobe .• tilillydns(ov~~ d!il!l~ r1 b~rs,

U~the di!SiS~(~lsf?~iflepf~~eal~' (ata~:i'lk triad., ~\~er~ ~heCi{!rlxt:wlat@ Qf ()1i'Q aSPClr~at~ residue

pulls
01111the

hydrMyi.

prol.M

of the ilctive-site ser-

R,re rred to' <IS pill,. that C!~e2~'o 3 nm wide ~!i1d are Inik.eiy to !be impor·
t~rgel cells. lsolat~ th,E' pil.i aJnd have Ci1arac'tef'lz.ed l~eir
tant in l:!.I'IabUln!1 the rI~itJvb!!!ln .1I.dlumi! to

r1'iembfa~~. nis li:arbo:;;~aL~ pu H!lOli! th~ am1de pfamn of im aspa ragil1~ 1~~5idueil1lh~!jra ng l'Ii!iili

in eo(Vi'f! ,allliil(l!~ ~rn,ediarv l1is.tidin@'), 8pP' also ll!ses an aSIl~ rlQlt.e, I,\Ihk:l1 happens re res~de on the 11'11'1@rst!lFra!c@' ![If thE! 13 baiU@'l ~nrl is locaJl~d muglllll haUwar i!(rm:'5 ltlll!thickne1ls or Ihe mrter

ney

M. l!ltlJo~K~~Qt;S wi~ih piili,

p.Ms<eilgE!~dli)mail~; ~li$ aCli~'&~~ Ih~ amid~ i1fUIi)g:~,n(or ~Ua(k 00 ~ nd' dB8J\I'llgeo~ lh,~ peptide 1:I~(lkbon~, yie'Id'i~g .jl15u(cilrnimlde lhal ,ool,ild be rt50l!{e!:l as a m~;(tm~ 'of <l5\fMragllmlll !lInnd[i5;o-

CClitilpo~ition l:Isun.g ImaSS, ~p~(trom~lry ~ll'Id ~mm~IIHI!:;h~mislry •.lhe pil.i .a~e as~tmb~ed hom llijw-mo~ecll!la II-wight prQl~~mI

~ubpflib;l~e5,e bind ~olll~' proleilllaminrin, which i50o1lifI' alnmd 1I111l rulpolleint(llf the eXlr<ilCietlell
lu la r mll<l,lli~with

.\iIspalf'a(Jine"[his o;IiSiP~ragil'le-.QIs'~artate' selr~cl.e;w·

un 1li1lDllU ti~'5 ue). IFulrlherrlll:lf~',

willh liBH.4l~ Y~Ii!~d MgBl. Alalpalfi el ,ot. !have lused p·~al!1e~~a'l,!'~di!!!uitv rll!nUt(mall~ecryin an i!'Miiirl to nanow the d'.;ll,mtingly l:;lrge r.;;llnge of polemt~aL "iJI1I1ptl!Ynd ~QlllilbinaU!:lm 'i\lo'flh e-;liIlWr~1n9fllil this ¥ein. Sper;ifh:aUy, they pe,rrormed ~ r(llllr;lh illllerY:fJ;;iliI;UI.aHon 1'502 knll\lIlli Ug h11lfl!!t.a ~ ,,;o'lid"i alnd us.ed the rre5uUs to ~m~ellmor,e than

is ~o IllI i:( ~og~thef 1\110 or Iil'! iJre hydride (.ompoUlld5 which, 01115 relea,sing hy~rO<gel'l, 'can '~ormasteb!!1 w-p1rmlll!(t ~ha'l dl"i!olle) the' raselk~!'l~MgH~, rlOr exal[1fQr;lI.e, ream ~

III

~
~
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g_... •

Ii:
i1i!

ar

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ED ITOR'S'CHO'ICE
3;00 1I1"H~p'aI'Led ~Q5Sible reartions among them, "l'iIliFI€'enr~c[i(liFIIs th'iM, ~~nithill g plr,omiS'~IiI~ w r,ulge< ~!1ilhe! H;)'i~s Uric! lJ~ i!1Jg a LClw~r as as up pel" bound, so .al~lei ellS ure ~~,a~ibie re,hyd r(!!' 9,~n\dHomJ'cf the m~l@[ia I~ \\leJ re' tn'~iIls ubjE!clla'd to !!l'Irnr~ (lI'Ii!PU ~alio~QtUy 1F1t.e~~ive ph®1l01il (fE!n!5nr-Qr-~,a~,@~ c1lIlCl.!~1Iiti ems. .ilIU thors nol,!!! [hal 'th~irtlppro;1d is IlfmitJ!2d y I,he ;U!;uJ,llIIp~ion b oJre;;!!(t ion !o ~ kl1l~\~lll ITIQrplliOkigy ,jl!l'!d also leave's open the ql.lesliQn Oil rnmabl.e kinetics"

cemectlens, the wIl5l1qtlencesof e:.;dusion

p r,ooomi
1'1.]

.ne

or

wen

nd,v neglali'li~-!eetil1l,g

hu,rt, ~(li ifng

nl?

INQn~~he~ess.. (o:rll!puwd e!ltha'lp~t:s 'of knQ~fl reactions prwed ~'Lr~fkjeIlH,y<lll[(ur.ate (wilhiUll ]0 lkJllnliot tO~ eNJpe'ri11iI'~1l t) to one r 11 promiSl1'19
IJrelulfilina ry sl mn 91m'll'dll'ln ism~o[ gllJI~ding

1r@~lig.~r'~nJtly, IDI"~u:fopting ~·llo!n!H~!lI.f ~~h~Sityl@OIl!Ild weoge ,el'tlt llave ~egl,lnl to eY..<!millle'what T l!lit;! ~t med iatl2 th€ se aiJ ilil[@'mI U~ a't¥pi(.al ~espol'!se$. Us.il1g 11 var~ety or e:l:poer~m!!ifI!:al (ont!!)(~s budltlS l~'~ Gllnollllicil[spilll~d-p1l!lildb inddenU ~lIiId mll\iiSU res ('5 IlIcha,5dOI1~ l~O~!i'!;It ,~)(Inw Of (oope rali (lnt~1'I a p,risoiller's dilemma ga me). th ey n!1dth~11eing (h"Hld~Iif',,~d~IS hC!1tiII1Qa high b Lilkelihood off,~ prosoclal tife'>lj'~e r.nith I1Illli'IY
~tmn.g roe OI'!~hrip5. s LIChtoil~ malnia ge, resuued 1~li in p1l rtfd pa 1ll:Ji tl!~lpilllg to pf:ck l!P' j]eooll1i (011 1I'Ver;;l,ge. {I ou l ~F lQ' spmed~ \1ers us '~he per fOFliIl· iillIII'('e': or lh oS'", labe~ed alii be ing apt te llead sell~ry Ihl~51(~es,s.thai'll p@'nd~pidood liP). As, to what f!lictor$ lJl.eciate the el(t@nt (or abs~l1Ic~) oJ PJlCD~m:ia~ be~avio~,'iOlillle 'CDf the likely cElndidaltes (liUS H 119till! othe r or having a '~~m5t!o~r elongb ing) did Il(Iil regi$l~r,. w~elrea~ e,1'Ilp~~hii{wm;~m dld" ComiJinll19 this rirnHIiI9 ~'!!ith,]l n ~.aJrli'HQne~ whim ~holo'l't'd I.hat £o(~a~ exc.lIlSIl!:lI'I acl'i'llClte1! the oural dF(ilJt~slil~od~!1g 1W~!l!. prod~c€:s. the spec· e u~ lUOI'Itha,l ,alii aner"efrNt or reiedi:ol'l is aill ~1Ti o~ioi'l€ll ~ I!Imbn@ss, or i!!ilI ~fi~ bl~iity 1!!llir!'Or 'tIl~ 'to aHl!!cUvE! lltal!ls ,t:l~ o~h'ns. 'G]( l: i¥Ts. )QI\', P!¥,h. 9i2, 56 [200]).

From rife on M,ars to life sciences

For careers in science, tum to Science

h,iwre experl meflls... - J S¥

Pliy5, (lIMI,

(lr~m. h~ '9,. 1CU039JlJ6lji92NI P (20~.m.

l13ef~f~ lt~'e ~Iealld~fla\l:afd~ or u~t,avlold (UlV) Ugllu e~poSLJI"eIi' [ull.y <lppre[ia.tf~. £1Jl1'I worshiippers ..er'~ applltoo lotfons hopil1!g toten r,euher 'thal!1t btu~. Skin t.ill1mil'Lj]reSIJl~L5, from the< prOl:lmtL(m or the ptr;Jmenlt itl1l!!I,a~'111[1, ioh ~bs,Qrbs tIIV Ii@dial~o," ~\i~ and can partiaU.y prot~(;t ullls IIQIlI'Ithe UVindUlced !DNA damage l~.Qit.(alii 'llilimate:ly (.ilWi'e ski 11 (,i! I'K!! r, Wi'th-out ll'Iel.anin,., G@II:5iU@ highlly S~S(ep:lib~ tOo s!.mUghl; sUflb urn is the b(!)d.y's re:ip0l'I:;e to lhi,s dllllilltl,ge. Cui et a,!'. sh ow thalliile lumm!;uppressm p5.3:, whl!;h hi!f1Ct~QIIiI:i .ai~~~rril! nS(JiplirDfl fi!l(~Qf ,jjfldli~

I~you want. your caree~ to s,kym[ il:e't. V15~t ScIerlt:;.eCareeil':5.,Qrg. 'We: are <committed to h'elpiliiig YOMfindl [!he

one of the ImQstinl€1ll5<ely 5.luciJiiei:l pmtei'l1$irn 1:1101ag 'f, Iliays a 'nidi:! l HII.e in UV'-indl.l!,ed melalliin prQdudiol'l. S[l,Idyimg p53~defideFll mif~e ;;I,S well as n;()rrnC'l~human srkin$~rli!"lf$. the:i <lirld th~n UV Hgh'l.adivate~ 1J53 inskin kerr.CIlin.{!g{l~~~th~outE!frn!1)SlC~lI5) artie! ti10a l p'S,3 at! ilji;Ue~ lh~ t;i ~!il~ elll::.oding pm-op1mnelano[mtifl {~OM.O.The IF'[J~~( prol!:!in i'5 ~hf:!ri Wl if! S1l!l'erat IPla(€;5 tlJ !l~f'l@rat@pl!plides,u~dl!dlin91·Cl-m~1a nocyte-sthnutatillg ilorffliolle, which ~tijtrnlJlal~~ lII'Iela.l1o(y~~ ()~ll5,

In H!lt~~~tr!~ mQ~i:l~!J'l'~:i. tm"':'Jocycl~!ls, are he~d O:rito. rrodliike ~:ores by bulky end groUl!)!> that eUe(tiV\~ly a' I:~5 slapll'er;s. Rot i1l:l;o1! Jilt s'YTllh~s.is lJ~illaUy 'eqU1~n'5 Hr$llhread1l'1gall open-ended rod thr'(l!.Jghtlle r:~clle, IfoU,('i\\l('d b:i lij)ll~ ~dditi:(m

right adivice,

l(lb.
Our

all'l Ii die li\i\E! ifig Use'Flil ~ k.l'!iJw~e-dlge is Wirl!'l'llv expertise of Scier1ce.

TOIll nd,ed 01'11 file! <lill d th e

ton g e:<pe: ri em ea of MAS i~ is: [he nlalural

adl!,lan (un g science around the wOlr~~. :5denl[!eCQJue'e,r'5,J)r;g $etec:UQil .

•.... ·8 ..
. ··1 D'~ oi'J

!
F;;

lm~aled ,1l!t the' ba 5:e. or ~Ih~~pidemli~~ Lt:lIprQ~ lI!Ul rnelaflli r:~.Int.e !'*.l$~i~lgl.y. POM( I)UQ~.eil)l~ysi$ iilt~ g;e'm~r:ates the opio,id PIl!pUd:e a-end!lrphin, wliliich to su 11L

lhi rdl mol.tclld~ '10 the r rjiN~ en d or the rod in o:rde r t@ ~Ofljn th~ se(lrlds.~oppe-r. C~i lJ e.f Qr. IlIewnt ill t~lO·lOmtpOllellit fivs.t~rR1illi I,'IIhkliliaril
elll1j,l·.:l!·villlykydopmpalle end group of the mdCllh@flhtl, mi!'CFO(lI'(

Fe-at~ ~e'svl1~tu:d:ii!" !II Theusands tO~j(lb pos"Ufigs

; th~' ~ u~h~rs_ 'iP~[U~ ~~emight (Qntrl:~I~

~ seekl'1IiI9 b!!:I'Hil~(lr

~,Car;ee r toels lirolin N,ext WEIve! ~ G rafllt information

~ Re:s'lIll1ileIOi~t,aba5.e • C.;areer FOfilJlm

i
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~

IlUlmC!l1s. -

PlK CeUU.a:, as?!

C2mm.

lijk'e rIlIol@:c!.de ~s~,'etls"

I@has

~ fSYC!ilOLOG't

.~ AJI

Ernpa,thy. BL'Q(:lk
_

.i5 IEiII',~ryday eXIJoer~elUe ((I1'lr!~ms l ~~ g ~1il,ef3il beti~f

~ thall1im1l1mSi3rEl5Jocia!. animalls; ~h~ Ine<ur<li~ alh· p ~~ays5.ubs~rvili.g ,prosi'j(iallbeh(iif~Cif:s i!ve' i! S1lIb.~ ject Qf !;Ilme~~t ~$ealrclh. and~ht: e'wlulliQnary cni-

J
=>

been ~hreaded, A Cope rea rr Cllflglelille III thal.v5 )10'1'1 01! t [room I~m p'~rf!hH@ O((,u rs Oi!ft@r 2: diJiY~ rD,r he.allillng ill 501"(, (uealing ill bulky (ydoheptadj· ell,e (;;I glp,i I1Ig! gtrou p. ltIyd Jogel'laliofl of the r<l(!Ell'l1lk p:rm:lud mh:tur'e' yi~ld~ a !iin~le~<ltu· r~t@d is,ome~ far the !Fcdl:ikJi! m~'l~wlf. The <l1lI! hors demon ~trratl1!l~'E' tech n iq ue-w~'th two

din~fefit
i;llfld

fn3.CfOCYCll'lS.,

(Onrfini1.ing !by !llJd~a~

Iflag n e'~jc re:flt:llllili:llllW :ii!)'e'ctll'Q~~QPY that lh,e f~ds

~ gJUI1I50of lh~e !behaviQrs: i'lre ho~ly d eb.ated. ,~ ilitthou gil th~rEli~ t'l!~den'I1m!h~t M!!dlll ~);.du~ion ~ (al'll elki[ red()u~l,ed e;ffoil'ls ~(1 de'lle~op sodal

ril'lgss1.JI((es<:>hdly iIlltedo d:. -IF!:OS J, I1tn. (III,rm So,.. 12'. lO.1J021Jj.aOM'I31~~

(2007).

1639'

SCIence
UIKI Ntw1tOfi~Jl/IiMIJI!!,. MY
·iWiiI~ingl)im',.IK ElfIDflilIl:

~WW.Sc;j,Ii!I"I(if:!m.1g.,QrQl

~~IT~·"!'~H~I,r
IliJirlu~WllmllOii:

Dii:!i!!r:!r~1 ~tnnectll· M:ll!mriii M. l!f.ulrb:rd

Inn [;D·jtQIl !CIl!'ViIlIllQf~

'~1:~O\Il1'o'li~RISII~R

,~~II)

!. ~M!H'

mfI!J5H~B!i!fl'I~fI'

D[PiJ""~~I]Il!~~. It. B!'l!~u HiIDlOOl,.ll;IfMOl R.1iii~~~ Km'il]::I LllllllJu

HI~Il~

:ilGOOli

2~NW.~551a fA.'( 202·~:N'5oi'i2 N~ m·:3~~'SOO, r,I!.'( :;:~Hi'1~~7

BI~tm~n llilou~t".B2~:e:IHnlll Riodl t~rJlbri!IJ~~,.IU,l!! (liZ lIlQ' 'i<44 ~Ol ~:~ 32~S.'l'!o,. FA:( .~.q. 0) t~n:t26S0:!l. (

_11

.i!:b:l,...,.[j4-1!J1AS ~l:~2:n Ci!' 2O!N26-6417,. FM .j!Q2-;8:42-106S. ~Uf!!! .adil!t~t:!~ MPS,.P.O. i1<';u; 9~P~. ~~ml'l'iJql;fl<, ~r::~09J)'6t7:a< il'f ~ ~mb~ :S~~~~ .m~;~ ~r~, ,!'(;{cm!.l.1'" ~W. ~ljJl1!J~, ()( ,OO~!i
oodN:nmi:b,
iii1TI~i1"N~iiiI,l S3U liIJ~:~Iil~H q~liOfI5
REll'otIJijR:

Sg~~~~I'!1>!i!! ~~'!!!'~ "~C~~!"I~~~I oiIc!df~j.J" ,~IH!if!~!~~IM. r!~

l,w, ~ntl!~iIlfI~:

~k!0i~~

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Ilfyou"re a yaung sci:einHst makingl inroad!s ln neurobiology resealfclil,t'l:le lIlext l~:ppe'lildorf and Science ,~ri'z:efor Neur'obiol:ogy Gould be yOl,Jr,s~ naiisamnual 'rfi's"earch p:n!€! r,eGo9nj:i1ie~, aC(;IOmpllisil'rl1!lHflts in n~llIrQbtQI'o,gy m~ealr'C:h based on method's of mo[leculaJ and cle~1 bio1ogy,'"TIlle winneI' ,8lnd finallsts are seleG'ted by a committee Qf independent. saie:nt.is.ts, chaired by the of S,cJ~,rl'ce, Past. wilrnners include cdi~or-ill1-Chier

post~doctolralscholars

and assistant prdessors.

To iae eligibh!!!, 'lOll mus,t tlEfl'35 ye3ff5, o~ ,age or youngler.

~f you'r,e select~cI as this:yeaJr's

W'if1.lrner, you w~1 ir,eceive

$:25.'OOIQ"have y:our work tpub'lisrneci irrntlhe ~re.stigiol,Js jQlum:a~ Sderu;e ,iilno be inv.[ted'tolAiSnI:I:ppe:nd arf ln iHambiJlrg, uerm<:!J'I\y. G,e'~ reclo'gll1mZ!ed~

Deadline for entrieS,7

Junle' '15, 2007

'W w-cw"eIPlpe'n(;ilorLcorfl/prize'

www.erppe·lI1id orfscfi,en ce prize ..org

eppencIaIf
PIUZE,

& Science
FIOIR NleUROBrnOl.OGY

I,

r
BY CONSTANCE

E·
HOlDEN

EDiilED

Great Materials Moments

WlrlOsays, malerials :science isn't s~:;:y? At 'Ihe internationall meeting o:llhe Mineril~s.,
Melal~ andl Mil.efials Society liiiSlllliioliith in

Orl:allldo. IFlorida. ~2001 mem:bers \10['00 0111 a list c'I'the Greatest Materials Momenl$ il1l History. 'He,re "5 tlileir lop, 10.

A Proteeme to Dlrool(, O'VI!lr

Sal.iviiI is; a 1O'I'llrld r u~, o own, lEellilingl with badertllJ, mucus, e-nzym,es.. skin Ciell!s, bloodl (EUS,. and hundreds 01 dil'ltH4Hlt prot~im;-'Ihe' produrt oil Ifill It]!'le glaJ,ds, serum lea kage. drain 9C' 'Jrom nas.al cavities. and what~\I'er you put in f'our mouth. SalIva is routin!.'ly il5ied to llE'stfor I1mIilOI'U:! levels and tilLega!ldmQ$. Su! ~o ra J the .~ III nly d iseas'f' it is used to C:!.'t,!.'{l is. AlIOS., .~ llloi3Jt may so 011 c'hi:mgl@, lh,~nb to 'lhe ~ Sali'll,a Proleol1le Pmje(t For 3 ~ears, several I!:! '~ inslli'tulions hav~ been cataloging every PII'(Io lain ~halappe'ar'5, in h~alll.hy people's, s,piL iii ~ Wi'lh 1500 proteiJlIls if! 'the del'la bcmk. sciel'l! list'S, flOW !N.cllnil. to eellect samples Irnm patients, ~ ""Hat diseas,es lhat might reveal their pFI!S' nee Bvla sallijviI, dental ulsearc:her David Wong 'of ~ [he Uillln~ve.rs.ilY CalUO:l"inia, tos .AllIgeles. told or the ilIllermlltiomd A~~!}~laliol'l hn Dellitall :~ Reseali(h lh1;s \V€e,1.;:n New Ofll€(lns, LOlUisia!ll~L l Although the AIDS511~i:llillest merely ~ (hecks ror HIV an:lioodies, scl!.'nlhls look o lO'.\l,UrJ mere ,comple:;;. tests+rnonitorinq raties or vari ous 5,l.Ibs,I<lln ill lh e sal.;va(es ,~ .., in 'Ihe rullulFe. Wun9 expects tnat testsfer oral ,~ (al'l(:~r nd S:io,gren's :syndrome, an alUloa ~ ~.:. ili'liflllf1,e di[~lea~i!!llfiat aif(lecu sali'll3 j)rodllc!;! lion, ".rill be available in ~he nest mu,p le or ~ !y~rs, Fuuih~r d'ClWlll tile rusd, he Ipredi'cl:S slI:i-

The Plerillildi, 'Tilble of Ewmenb

I ron

z
3
01

s melti 1'19
CUDO SLf.J

iran !ii Sl:Of

Glm
OJ;lti(al mh;rQ<SCQpy

5
6

U668~
(175~U (300 B.C.EJ (5000 B.C.E.) (1912)

(cn rete Crllci~ e steelmaking

Copper e tractien and !tolsting

s
9'
1,0

X-ray diftlfrilcthm
Lteogeml!r j:JrQt~~'.ii

n!l56~

IRehalbiLitaling P'ltut!DI
The lalf~t sLrike inl 'LhePl.!,ltowa~ has cOll1e h,om
tile New' Me~i(O Sl'al.e H'IlUS!! of Repres!!I'I'!aliv!l's.
LaWil"i~

Ta,x,ono,my, the'
EalrLy ¥elalrs
!By th~ lale liCOs, sdel1ltisls had ~tegorhred more lflan 4 000 '5pe des o'f an im~t'll. -orren lUdked away iiii out-or'll ri nt pu blicatio'H§,
1

'i
u, (;I

lets ~here last week defi ed the ded si Oil

!ly ~ne Interna l~I;Hlal, 11.5inm!1m i~1 UqliQ 11 (~AU)['0 redassiry P;lUlt(HlIS a ..dr.lil rl p lal'let "
lna stal.emenl approved 7'0-0. lhe Hlou~e declared flulo "a plalle'I"andi U Man::h-the day o:f the vO:!e?'-,1li5~Pllllo' Plane'! Daly" in New l\~e~i(O. It was 'on 13 MalfcIil1930 thai 24-y~ar·old Clyde fombaugn ,or LH Cruees, all amaleu r a s.lrofillo
.:I 1'1III C' 1.1

~hese e-alr!)fdewiptiOifiS ca 11be diffh::ult. rl)f loodern Ire)fanJleir~~o nl dowifl. AlrIim.tll.Base hom hu ~e U:ni\i\~r'lii'tv or GoUingen in Gemlany opens up!i11? d~ssi( ta.!l:onolfliic ~il~rall!re. ilu~l.ibrary stmes ,Dr l.ill:~ to digitized
lIer!;iOllnS, oJ more Illaill

endsts

§ fgr lung, breast.and p",ncrea lu(, 'ane'!! rs, as. ~ lO'I'eU as, diabetes and! ~'iI~1II AlzhefIfrH~r',s, disease. ~ Wong sa]'5 11 "Salhl'1;l1rY D~agnosth:
,~ RCl'adlll1il1p -tombinil:l!l both the proteome and IE va tal From anOliler projecl cataloging pi ces of .~ IR A lln,all r~lal~ 10 the pml@ill~~will be avail· ;;l alble i111a bout ,<I nl0Ii1ll1~.
w

wm be ablew

I1Hi~r,

detect. pro'le~1IImiilrkef5

nC.e"d his dfs-

c~ry

or PllIlQ,

7 00 books a ii1d papers, 501ne 'hom 815 ra r backas ~heH ~OCS. A~o:ngwi Ml a stack O'r
1

irl;)piri ng to calprid e lhiill a,ppa re filly

works by (arcdu:i

Linnaeu'5. llie 5~\I~dis;h

bOlta llIi,!>t ~'ilh a relorrned t<IXOIIIOIlilY in lite the 1101d-

endures. Tomlbau'gh
died ill 1'997, bu' his rj\lid'I:lW Pahy was pu~s;en't '[CDrlhe vol,e. wTh!!1fl!are ,Il!llople ~~ho tillke [the IAU'!; .OI(UOI'I] as an anw'onl to Ar:JJiE~ri(;lIl11 i!I'slrllllomy." 5,ays pt~nel.ary SdUli'S~ S. Al~nl SlilHfl,of the .scHlhl,'l,lE!'sl. 'Fiesear,(h Im.stilule in San Allrlonio,. 1~~s.. i~e disco\'ery or Plmo '\I'as~po<hat It wa5 heralding the Kuiper b!.'ll-{lliu~ ollhe hcnlie~t topics ill planetarysdence. ~ The Nel~1 Me~ic;Q Sellatte must vote on the meilliure[o make it ,0Ui cia I.
Goat5 kilil"'(llv'@lied

noos.

ings imlude le~'Brkrlown (Ontrntbuti[m~

S'USCIIOI IFish@r:.~Vhil ~\Iorr.:.s'IJIn~hE!Sa Ii va IProteOnll'! Pm.ie!1:'l Qit ll1e Uni'li'er!ln'ly oii '11 (alUcmnfal, San Flam:~:u.(I, s.ay:s lha~. i,!;, bio~ .~ II:hell1li5~ry i:s, I1Jmi;lv.eled, ~ there are i;llwi;llj5 'il'er)f interesling SI!.l uprises ~-as sci~i'll~SlS. have' dis(Overed, for e.xample. fromthe blood test [or

~
L:J

.~
g ...
",.

prostate caacer; So sh'e se1ys.iI's ,possible sativa

1:1

win yield informilliolil that can', be obtailil'!.'~d

~ from Ibllood

lest's"

boob frem late ~uthas the 1768 tr,eantlll <lind '!!,1Irlly l~se by the Am~ri,1Il natJllBUiI ,cernltu ri'es. ufilllrs.t]o5ie'phLls INlicG;llavslaureflti that d,esuil;ies; 'he trl!le l(lads. ~.8ufotTo help Yisi'lors Irad dOMVIi1 th~ (irs'! LRsr: or a particular ~demrliH( name, euraters have ~@gl!lfl COlli bing the Le:llS 'ror rnel1llioo s or species ami other taxonomic groups.»
\'P,WI.anim1ilb~@,org

'oWNII'.'SlCiencem~g.org

S,C~EIN'CI!Val 315 ,23 MARCH 2007

1643

" WIUD

'

BY VUDHIJI1

:BHATl'ACHARjIH

Pioneers
rnDliUtIJ]lG, Vijay P~JlJd~dn~~ n·l \va~~t te SilOP s cllilikll.'len from playullg video game s, hi fact ..

the StllnFord Ulli:vens,ily chemist th:illlik.s; they \,'hal~Th~g,euj AJzhcimci'S d isease, Lastweek, r<U1ide .~~d:StUll}' COMputer

ca~, h~lp hin~. understand

ElIlc'U'~a~llInelllt AmodclI nnnouneed i,~l'Ilby I,he

ead or dl~~ ~l(mth. S(ln}lrl;lySt~ti~n 3 (PS31 (io.n~PlJu:lr ellte~,lt3,ill:llm~ml syslielHS. 'will be ;Ible t~ ~mm:(:lliO IP~Lml~':;; F~ll.di!lg@Yk)!l'L~e ~F@H.) 1~1·l)gJ·Rm:. TII~e:~rogf.Ellmis .1 d~:s!.I,·ibll~~d eompl~l~n; IH'QiecIOI1.pW!ehl fllldil:1S '~rld misfo.~dirig, \\'m ic~~emil, ~I~md, AI z;~ei11.11ft's i$1'" to d ~l1S~. ~n'l1longothers, .1 rS.3 users ~tctiV-i'lic I:h~ f
CQlrnlrn.eclioll,. t~:l~irll1.~~iCh~ Hes-wiiil.eill idEe- iNi ,SlUlOln;:lli,eOlI]y S,}'!lch up widl h!~c!e ~~ nCl\Y{l'rk ,;md ~r~'!lch ~~u~~Ib~r.;, in p.ro Idn~ fold i[lg simu la~io:l1}l. he rns,rer pmces._<;(lr ch~:ps. the 'rS3,~\'ill cxtcnitlilhework dane on: F@H by nearly 2: :m1,lil~ T lin lion desklOpOOntilplitern ~Lnce OCl:o'ber 1000.
1

Albl:idc;,pi ',g rssaareher

DlJilli BI~~C.IUHI~i:;;

:steppi llig

P<.!nd~is hoping the projec l rnigtu even s~:(';er some; !'!u.H:ie!'lrto ~(I\.\(;_.rdCilreii:t.!!! in science by g.iving Ibe:l1~~he chance [0 W!lt"d.lthe ~n'otc']Il~fo~dJi]1lg simu~!illiollin real time 2I!mlllllllnipubll:c proteins on screea, "~.~'s <I uniqueway nl educaiepeople .liH}'lItscience Oil :lnrn[!lecuh1i1l
scale," he
S<1)fS.

of the a.~yrn1.1pic AIl!~~)olic~d b.oora~ory" whick ~'n.e fuull<if{!d 2:5 y:e.,u.s. <1~Q .tt llile UiJivCIll.il.y of Cal~f~x!11Ii~c. Ai!l.g,~I,~, ('filii!'!. (,:8. is Los credited,?i,l ithul'lc(lVednS l'I1!fumysports dbp:ing schemes. fi.e ·wi.11 ckvou~ lusrirne ·~o rese~n:h !l~ the luni-Dop.il'lg Resl::ilJeh
lnstinna, a Illm~~uont sat uplaliit ~all:, ~le

dO'I/VIIII~JS cliU'ec'lt!r

'Ill!: Look.ing

$1!J'~KeedsWi l~li~rnHi;Irris,. whQ left SIFI l~st yeil r

tQ ba'c.oml!lhe.~d of th~
Ariwn~.

I.lu! ElIfQI}G!iilflI Mo1e(ula~ Biology ILallonnolry,

1~~o~ii'!I~ the[ieal~Y ,mmi~(c.'I;i\!lr]e5 '0r dl1!!J:g busls lf~IIlI~lr'll!' ~.iil!!ld3'~ruy PIIIOIIC~ @,f1 You k:~{n.v. h'~ h~!'(! W be proud of ~ bli!l.~t.Il~~~ln:knOfbo~,e~btl:li1.e~s: one of 'Ihe
nm.G~!J; i!llP(N'Ui~t

lJ.itl;:~ \i,,~t;j'\I~li'

did.lluu

Ultll. he'lped identify [me gen~~ r@'.5pol1s]olef:or lh~ gro~'IIlh and speddU~atioll of e\fllbry(mic stllm oolls. An d .A!ilile :rVlda ren, 11ril!s~a rell as:so-,diilte~t tme, lJinli'lllersily of (.~mb,ridge, U.Jt,l!\IifliS "'" the honor 'f'O,r ih~~ ('0 nlril:illltiom;.l:O th~ Iiietd of ~ :~ reprQd lJclive t~dunolog(li', 'iirld IJdiin~ the first :::> 8 su~(essful. ilU.~mpt at grcrwa nga rna use err! bryQ t ~ in 011est tu be, ami then irn,p'l.<JIIliLirtgl it for na1uwal !Z bir~L Tll~ehll"Owin'il~Ns sham $250,000. !::
:~

Janel ROSSglnt, a :5~m cell. r'eSeClfChiew fit. the Uf'li\li~FSiity of TorO!lto~1I'1 Callao"", rec:eives~he' award fDr work

impn:n!il'lgl [hie i~ealUHJi babIes-

EM13Q: i~ $~mIoQking fo~ a~Ll:C(;eSSQr.

IEiACK ~fnoo r Acce lergloor Cent!.:! r (SlAC) Direocto:r
S~lIrl!r() rd

Sdeme Fo,ulfldiation

'!i~'~l:'; ~ibn:!I1lflili.et (If de~igll!.er ::l,h.ifOid", ii

ro PHV:SH::S.

\Viruel1 ~ flugure{il, ~~~,u one out, :1 .knfl'\!lf 'dm~ lhen:: wen: peop~le schel1lllli,~g and dc'veiopinl,1; de:;~iglllC;.r slemwds th~.[ we

oOlldd[I 'I se e or fiad,

)onatha n DorIan wW step d01j\l1"l Uris f~U aher 8 ye"::!lrS~S rne;!ld

of theradlHy, w~i:c:hhi: fl,Jn~ed by tne IllS. Depan.ment of IEl':!ergy (DOE) and Uifl1maged by' Slillnfofd' UllfV€Fsity .. The (ontrad m rLlrn~heS3:00 rolUtoifll1llb is e-xpec:t:ed 1:0 bE! put up for bid S(!!O!1. S~:a1fOlr,a plans [0 cl)rnlJ!et~, r bl!rl Da.rfailli ~\!'i:lI1t5he u rui'l.limily~o find! !!! i:le!.11 t l~ad«J'.'~'rn lhinking of :5I.AC's fUltIJ.;U!l',~· S.fBVS" he IDmfJlil has b~li!i9at. s~,]"rrmdfor 30 y~'Jnl I ariel ~ifS he's looICi!lg fOrl!'IIard to r8'luf.l1!iing to I,ne oonch to e):,p~or:e 'the ~\I~ry sf9 l'1irff.c.!Int. ~y5lerie{ in dark maltler a J'i·dcos!nolo9V. DorftJll'I ill'~iOplalM ~Olhe~p "lith ·i:i!ffo,r!5Ia brirug file Illtel'll'aliorilglllLillear C'QUjderr~(llhe Ulflile>d St!1ltt'lS, whicih me s.m!ys WQu!d s~~I:aiflIWi.S. high-

'o~HasVD<Ur

wo~ik,(f~ii!I,ng€d hGw~'O'!Jlf~@W. ab0!ilt !!!,'or~? YC'<I.i1,. a \~13y. I .re:JUy love line I{) IYrl.1P ic ill
'm{l{jl~L \:vhl;lre 200 eeumrles can :!III g.~~

cmt l~Je;ir bes.t l!I.lthl.eles aad C(lHlIpeJ~c. <l11lI.d '11~lit"OOS'~ llflIf':un <lind wemen \v.u 111.If~ beautlt~::~tl e~"I.Cili,lIb.g.. 2ilIlld ~[ showdd be: ,'c:ty .nlld 'p~1.I:~. But. DIIQWl;I:d~y.li some~~!In~s your

;~ M O\U!: IRs

lruo'pes ".111(; d iIshed ",;'helil

se-aad-so is d iny,

y01JJL

read rh at

; GO[ING HOMlE.lrnsh biologist Flr,1.!llk

~ Gannon, 59', has been tlppoint!'ld dil"~(It.or g!l'l1.~el"al of the Sdenc·eFQl.mdilliol'l of Ireland ~: (SF!). a filll~ding aglenc.:y \llith ill '€175 m~UUlOln .~ annual budget. He had ~Illloun!!:ed his reilir>e•. mM! as. dife<'l.or ,olf the: Europe.1i n MClh~clJ~.alr ~_~

''ll= D:oYQU ~M1111( elj!le r be~llIlb~~to liJ'ut w,e'lm

alii em!~ll'O' ~lrIls1
[fyfllllf

.
1:0 .~~rflU

drU:!f" out fIr ;i]U sportsJorever, you're going tiO die very u1l'i~~Wy. B\!~ y,rwi ~hl'!ll!ld be illh!l;l.~(~

obJ1ec~-i\i,e is

§ Biotogy

~. :2006, p~1665). " . .~ GilrifiOIl, who also m~adoo ,a glroUJP s:tIJdy.~ iflg gene Ulllitrol b·y tile e~luoge,n receplorr iit

Chg~riI]::l:~l'i(lnl (EMBO) ilil IHcidelb;efg,

l_ ," .•

..

German

~i1I

December (Scienc€', 15 De:ciimlbeli

~~hm dl:e' HmUy ·lruigh~~:~.d ~I"am;. witt'lre the b],I!; f1·::un.(l::>.~re cmn,pet'~ng. me clean.
.;:e~. C()JIU1fo:~,e£loM.~h

energy p:hysk'j.fQr ~ gemeratiOJl,

1645

BClw~·~n 199& and 2007 + the uunrher of $t,~i1d,~!i'd i!w~-,\lH~lor·'i!1Jilj.atil>id ,mO[ J grn!~~~ roughly douMed to about ::lOJJrOO.Such

~"'rIHl,~i~~n ~Il~ir,.".:;, .. hm,g-oolfrl,~ "'XJ(n~mi~~'qMnt~

:5<1]01, because the <ligeocy prevides mos~ researchers wii]n, '~l ~il:i~t 'I~JQnion efthe it )!<~~,,~ry covers overhead eosrs, and "\1V~ boug'l'Dl in" [0 the deubl ing" "Illlid IKJW we're gettilmg ClH." says Joan B:il~~_ge" chair uf~l~~ (:(:11bi'olug.)1 deparnnem ~l researchers

Haevard

MedlLcml Sc~rn.oot

jill

au interview

befere she leMi fied Iii! t heheariug. Bru[u:.\'C. whQ II eg.u1. l,t! :>lmiy ,C.lllc'er i['ace II eg,e i~nt\r
he!"~'~s[ll:r as d.u~gno~(!d, ~,t~la f2liI~1 w w brain IILI'lilIl!or.::J.iiIi,Ylii lhal the slu\'\bd!own is espeei .. nly ~'rw;'tJr~~i:!'nM g i",~~re.cclltlld vauecs in U['iidcr.~t'l.~djf!,g h~b.u~il; bitdtlg)' ~)fC'H~~clf.Or. '~l'hi;;; ~ noit onlylhrt:cl:i't<l lis ~'I.·Dgrr:55 but ereatesa n

2(11)8, BUOf;eT

nlj),c~:l~,(:~1 arehand se ntor ~,~V~Siigi~tOQ:S rese Two powerful charnpions ofbiol1ledicjJ,~ m]~y stem cell ~ ines created be fQi~-e I) A:uglJSl 1e;;iJ,villlthe fiel.d <l,fter rein~, unable ~o secure Nm~l~ldil:'l!.tR('Ib~1!'tSiHekm.,o, wlm ~tud~l.;;s re~~rchbl~IJ'lOO th~ ,!;Vbil~ Heuse s pr,opo~~i 2001 m~y be ~!t-:,~d r~dcraU)' nJ~d~d in to cut funding for the Natn,Citl.l.,.I ,I:~~itllles of roes.e'<Ircilil,. However, Zerhouni said, "ifs very i. n V at Johns Hop,ki ns Uuiversi ~y ~,:Im. h ~,~ Ba H,Ci"Ihh fNn·[) i:11I 2008 and ,Ur.lviN:d research dear .. , that t~lese c:~IU~i.nles.w.m not be SlI fI'iullore. MmylOlJJd said he used to spend 3'O%o:ff" cient t:o dOH~~,the :~~.~(1;hlhm \\'1; need t~, o:' d ioei!den:. to vent their 0'1;\,'0 frustrations ill it hlis time 'l[lplying 'lor gru.m s. Nll\,V. ]~e lollljlihe scnatcrs, :il·S jmuped, 1.0 (i.cl'flk, 'Senate hearing this week, Senarors Tom 'f~'rtcAdlni.rili:su'<lliof'il has decHned to Ilmd new h~uli:l<LnI!:mbl)'{Ja:Jlic stem ceU lines and ,~.titl ~ii~Ilniversiries are a h-~<udy mQbiJ:ilil~l,g to Harki n ~ D-l A land A rle 111 Specter ~R- PAJ'. '!'Ina ~'D,e,~d slII1x'omnm,iuee ~hm Im'lmdlc::sNl~i, ~Trn,e rle abour l~.e extent to \IA:l ic~. federal do liars, lobby for morel1.lllui ing, .~il,~unedi3'lfJ:y fl.er the a ~l~otl!d bfuck Ihis r~C'.fiucll, S~i!'I:fLH~ h~a[,~I,g, ,[I coo:tiittm~ ofUil ineinsti !~ll !rlJ:itoi fUi:!,dl~l:lg" ,gd lie d NIH Di ree (or E ~las Zerhcuni 011 1,9 M;lI,iCh ,;loom the :~mfl;u:;l of Zeill1'n(liUIl~"S "-;<111.Cm bom>ling li~lIldung 1'0:1" and 20 scientists. .unCiudill,l<g the fom \I;~l.o testified, released jlglt]~~y. 21~p.;;~gB repoil rhat W~~LfW~~uld he rhe fif!~ c'OIlNecmi'!,.'€= 'yt;lIr of hu~mlL~l'm.h[Y(~IDIk M1<lnl cell research cnmes as N.~~:I is stil] .s,lIr~l,ggli:lllg te apPol'lioml, its su'binfh1.lio,l'DllITY budgets for Nil I. They heanl describes 1.'IJ.':CeII.llridesin eaneer; s:p~n<llt"ord Sl $28JI billion 200'7 LHI!dlg~l.,enacted .5 weeks i:Leajul)I, ::Ind ot~lI~'r dis~atlCS,!lll'gl,.liug t~;,?tl NI:~'-I senior sciCil'tus:~sdl:scrilx:::I bleakresesreh elimare in ,"11i,chtlte ~Jeij'oelm.lg,e of funded NIH a,go. Od.'ler <1I,gt:n~ie~,<liI:'C s~nlg!tdi~,.~. too: I:he gJ·..I'lfiltsJrewell ~pe.~t mild l'allill.ellil:i!~g theelTects 1 of flat n~!ldi.mig. gJ[11iut~i~pliC'mijol:'i~, ba ...drop~d rrml':l 30'% (0 pru:oid~1H '~20m: budg'Cl pmpClt>i:~ "~r~lty 2a~Xn. And the se,~~.~torsprnlHised to press for mlud.l ,fb.l--fwllildi Ij'lj!, e'l'erytlll.:ilnig Od1CT IJ~<Jn Bul: beyon:CIlhe mlecdolies. {he' researchers mf)N~!(m~y fer bi(l!n~dkll~~~1ll:'dl i~~2:00fl:, d~~~:ut~:' ~,f~S Ju~ R~ll~;;In: d iIl;'t':tur of I\lgfund tI !~iv~~ity ..Jilm~!lijS!r'-H!(lrS ~iln~li!.;d up i".;w N{llle of tbi~ ..... unexpected; H,ltki~ ,.1:-; .md i~~mh~~l~lliun:jlur Ih~ fif;.ld~roltiofl ft\me~'i~ g h<l.rdl·ig:ltre~ O:1!l, lb~ harm fhn bwuJ,~l;ln~<.In;: Specter helped wi:l'D.NU4 111.2%h;u::I'~ase In nlll Sacieties ~('lr E.K~)en.·.imeI;u3,1 Ilicdogy in c-.ul,siug .... I.know th'~s wi.~:l1'D.Dt \1I1'O:rkut [0 be <I : o m.Hkmu~ic,1i] RHillW!<I:; said SI)~C'~r;.burhe 2001 dun ~be \Vhi(f;: Home dkln't request Belhe5da., JVi~ry~,a~~.d. Some <1.gc'llc:ues th.lu l professed Fmstrnlkm ata II'ock of data d~alhc (Science •. 23 February, p.1 062:~. J'V!orc fund research i.~lim phys[ciiil sc lcnces, lneludi]lI:!, rhe NrLl:iolfllai :Sc.ie.[l(,"eFQlII'rad~~lio:lJil. \\iOIJlId, ,mi(ghl effer his more skeptical Seuare eolsurprising was <In iillll'lla8S~,oned speech by Zer,liloulTlli abmt!: ~_~;eneed fOi~ Ifedel,TI:I,~yl1.nldfd iJlo¥.'eVl~'~ receive I,"C<l,iaerease s ILl m~jJt:'rhe .. I t leagues, "Whla~"So goi ny to hOlppelli W N~ i~ ~j,r I'\d~~ in~l'~'!"l!lt 's bWld$ClprOp()smh, (S(,f,'rn't', iOml, hunlil.m ~mbryonie $H';f!1 cell research. In Ul~ b~tcllg.clis elll by $500 nlilt~UonTt he ~''''''<!!f!!~d respoase to a question, he divel.·:gedfrom 9 fCbI'!lJlliIry. p, ;501. lO know. "'It would be very be.ipf\J~ to li:n:tn~' Iwnic'aUy. even U~(~il~h Congress ,g3i;/'tl iH)Wm~~)' research projects you .Ire 1Jlil!!d~r~Adm iT11 ~m'!'~it_H'I icy. assert in ~ Ihlil~t~~!? 'pol l[;ne:nls t~,nu !ldlll h 1>~emoeUs Cl'il1 :~rfOll'm the NUI'I smfl~,i inc['!J.'!'I!l~iIl2007.t:he ~ency is ~l!Ik:ung.1:n1.J1,how 11I1'1IIny you're '~lHnilligfl\""dy.'· ~ The ~]~I.'j)dnl·J ~MS the SCmlillC~!>, otrl(:I],iflJj~ <! W ~me i~s.kl'i,!;'IS i:lu:cmb.ryollic;: V'dri.t1t.y "do 110 l ~L'l'(I'cr ~.I)il.rlic~llar ~lr:Ii1i.111 bcci:lJu~ .it .is ~~!illl~llg 1101,O\'1; in i.t:> co~sid,enlt~,Qn QfNlfr~ 2008 ; Ihold ~c'iem,ilfi,c \\'<tt>er." He <lidded that <lily olTIl~IS!1'1l~m~,8 i,[o 1998 to .2003. \".'~,u:n it 5av~' aU~mpl '·to sidel.i!llC' N ~H ll!l a~,uS&l~~ (:If ~~~I~h its bl~d~ctdouble. 'l1lliU ~)rol~.ll:'ll.ooIm:;;~~'U11i- b"lll.dget"1 proC~S;ii Ill!)! is c1l!;p~~~ted~o If~"J::,at imporr~liI,cc i's ~,hortsig1in-ed:' \rer..;;:utie::; to eo,;pal1l{t cm.lSt:mcl: Iffie'!.,,"' :Wad,liti.cs. Ie-d:st· ~Iiiliti]tin: fall" ~ f'lWCsideu~(i(lI)rg.~\~(I3I)l:sh has wlled that .md recl"l1Jt n~w ill'l,Il'.fitig,ltOrs, S<fi)'$ R.el;,;kll[ -:JENNI~ER: (:QUZIN' S

Senators IOlfflerSymlp'8the,t'i,c Earr' to 'C'omlplainlts, 0,01 NIH's, Hseal ,Sli"de

~um(l~phil:re (lfunce:~~t1i~~y asd ill!'tiJ\il;>ny:~ she

told the seuaters. Scientists ,',]'SQ spO:kl;:of Wld!';(g.flIdu!It!!;!:s and ~l"adlL.m~e:m~dl':[lil\'l tu D'llliilTllgm:VJY from bio-

1M AT:H EIMATIIICS

For lTIil,o:re th<'l.~.a e e um ry.n~.a tbemat le ians have :liln~ggled to cOflllpn;l:I~.1;l1H1 a vast, 248~diltLensi()m:'lI entily~ b,O\?r.'11Io them only t a" Es' The::t ~~a\'~ d(l'scuiiKldit a... ··m!l.!.1:ic" !!Iml "mli raeu leu s," 1b'~.1 mn.i~ IJiOf.~t. lhey cou ld Imol !:'1.l'O'llly Uml(;l11a::;;~ml how j! i~ p~lt wg~~.ln(:f. Thli.:!i: week, :1111 inreraaticaal teara of ! g llU'uhcm~!i,cia:~ll> ..nd com puter Si!.:uelui:sts caUed ll~e AI.I'I!S .rn1'Jt:cthe.~(,lJed by leffrt:y
Adaurs of the Unhrer~i ty 0 ,. Marjll:!lnd. C g Ilege Pa rk, 1\~,11JIO~~ ed l h at .t su pernc e'O~]p~lterG!IUed S!l.~1; IUJu:> ~~H;C~.5,sfu.ily "mapped" l;;~,.hi the wnrd:ii of (:I.r~gg Zuckerman, a m~tihC'mi'l.ti;d~11i "i~.le Unh'el'~ <It

s.iQlmls. hey3100 much :1lQ,l.Q:reconunoUl •. and Ihd,r L ::;;ymm~[riesm;-e e1rl:pre~~M:d y Lie groups. b Even the si:rI]ll~e!lt Lie group. lhemta~ lions of <lIi sphere, has r:rofom~d S'cielil~

tific

OII,]lj)QrI:<lllce·,

th~ i>:~P';:(If electren OIDlllak The m!1(~ ~.nleresl= SO{J)

or t\. ~omrols

Th is

gl,i()U~:l\.c.1IIed

ing Uc group, quarks.

SUO·) or A".

describes the ~vrlilrl1t!~ries ."

ol

lVi mray Gell··l' .... blllm

n.~ck.nO!lIl1,~,ed rl'fi.e"i!i.ghlfllid it W'd"j," because ~l is ~l1Q. eight-

d i!:nenr:-;i.Qf!;JI] gO);u?Phy~i·~~.isIS'
quest fer gmm:i ~m,ii'ie{11til~oo~~es

shy. "uever before has the mm~dageof pure

nmt~,lC[lm;~ltUcs,m.d 3upel.'t:ornplll~ Ilg Plx)(iillced

such ~ prized !JnSpri~]g.'·

!:'fl'(l rs

L aJlnlc~.led :~n 2002. l:me Al~.I!lS,Pmje·ct has d~cp ~nUiL~;;J!t'hc!:trilat~ ~~~stary, The ell ancient Greeks were Iase inated by e r)'s[<Ih and j:'lolih~d~. b~~ml~ lI~~jll"riC~~ S)1I1'Hmle'ny. ln 01, e IfY"'i1~<'Ii.however, :;Jln.n 11'J.elryemes 'illl c disc.rele ebunks ami Hmlhed numbers.~nl~~e

h~],~, to ever-l tll:g.er g.WliJIpr--.led !lnul Es, is dlC' biggest, ~U)51 exmi'C.'! ~!ysy nnae tric ~J f~~le!n ;d]. lt s
mathematical richness makes it a m~gtIlc~ fur stri I:!g theorists, a!!1nlO~$
I

or

o~J;ll::rs."The're's <1. ph ilosophical question af \\.'~}' !~U,iI,l!.'~\vou!d pick ene ~~nt[,p ,~:we'r ~:n,o~1'nea:':· .ays Jollilml, B.aez.'1 IIlmllllln.e\lnm~ s ileal ph)'~h:is.tat Ihe Univers;uty oITliI.lilornb,

~id~191hl entury, c

the N{ln\;egi;:nll. 111~tlhellilli;Hi"'~IC(;") SUI]'I,~d

Riverside, "!Ell: is so awe ... ~]~d~in ll!obQdy u

'~Qu.~d quan:e~wJ.l.h this C:~l~)ice." The German mathematician Wilhel:~~ KiUm ng posited ~t~.e istence QfI~ i~1 lSS1. :II] ... e..x
V'i.st~UilIS '@.f s)'ifI1lll11etry.• l1he ei9 hu,·d ~lIlum:!!iOl'llill met 1.1 U.1Cie ~a ~here l!lrll~~l.~el!l j ntea IP~~IIH'!)s H~.e~ or l (elr~nu(lloe:u~-·a ~1.;u::eNt1~r~i~'klr!fll~li@!il about [ii~S

e ian Sophus L ~(;':. l PliolJ1!n.O'I,IJlC0d

~t!ldy:iing obj:e:cls, hh smeeth rotarienal symw metries. Such objeets are ran: in threedirrrcnsinnal $pOf~Ce~ spheres, cyhfldeti;, doughmus, ;m,nt! ones, But ]1:1 hi,ghf'D'dirnenc

paper [llitl b!"flkJ;:: Il~Ipn$-~Able Li~~!"m.ap1O drnViii ~nlQ Ibur ~n~J1i~e f3111il.ies ~Iabe!ed,!\ through Fn 1"11'1"\\ flve ·;,.·. ·'.nnio ,11 ,:o[o,,)n!o;·'· (G] •. F' E .~ ... bI! f" -"'- II . ._, '''''''''''1'· _. !.,,- !:'_. '·1'" .:1; ~. ~.

reopr~5e9t1l10n~ is :5lmed

il1l

(0)

mprll'Ssea I'orln.

~iniJl. ef~llI~:.
M;rthematicilllil· pF(Jg:rarnrmer IFdkko

s; and Es)'

HisFrench

t.'oi'reagLl.e :EiIi,t:Car!I:ul

abstract. and solIll!tulmesrtm"l'e.:ll.lineorem.s

group theorists

of

d~s.I;!r~bedE~. in .~ &94. In essence, K.il]ing and

n~mmtoget~.lfr

into wu~k]~.g, ~1.gOiiHh.~M'.

suppl ied the t1~:\:JO:I1()m'y f lie o

;;Lii1I;j~dIO

du (~Oi!J;!;.:5P~!lIt hl!S· 1.as1i:months, 1iry'i'rrlg toil fifll$h t~ ~B pr()l~(;t.

!il1Jlf~~tP~. AII(!i~ Pro1~el. ilihe I.heillt gelfiiOlm:s,

cumllu~!..:

h~r!i'i!~h~!~ll~tk'i!lh;:!iins., lhe MIWtil ~t ~Yl,.Qttu !TlUp each gnmp ':-;"irreducible n:pn:"entilrlons": l~,C: set ordii'fen:n! a-dlmeuslnual
spaces o1.1,\'l.111ciflthe gro~TP 's rol.:rt~OIl$ can
l1l~tlX:I1i1l2Ltlk:irulS" [Ildld llig UCt

"M<lIthem'otti,c:iilllS olen t1Jb: shoftculS 1bflt'''1l~ [hej' k~l('i\V \Vh~H ha~to be tiI'iL1C,b~LI th~ Cfimlputer doesn "I':' says D<Ii\~d \logall of the M.'I5S{!lChl!~ll~lm,tihute u!TGchntlIQg_y,on~ of th~ p;;micipmlt:i in the ~li[Oj~Ct."f:nkkill \"~r1l bm,;:k, and fOllnd a'UI:he d:'tiljl~S, tlhL<'II\verel1i't uitel:ighl q

i1,nd mm.lie it .11:1;1 perioct"

Du C~oux ,cillc~tIO'llCd th<lt EIJ:has; .453.{160 fruni Iies.. 'V~.nlilliid expeced <loom ~.b.illl±on.)
That meantthe '·ge'~iOme·· ofE8 wouid be a
w:b[(;l tV~dl 45.1060ITl\:n.

Em'iUer

Zuckc:mmn

~ \\'h,o\~rdS

Admuss thesis

Old\f]lSe'r).

had premed

Illma~: repeeseruanons can be divided into lite f1!mil~$" ef!t;"h,!j(iml.~fah;.d y~l m'iII:l :fu~mll1l!i!. ]n bs r.hoo.wy. it should be PQ.~il:Jl.e 10 use d,e rorlllll~ as
(0

almJ 4:53,(160

eelurnas,

or more l~l'.Ulll00 billion entries.

DuCloux

eril!'lk

~Jtil

if:r~d~~.cibh~.ll~n~·s~rH'I'~h,J!\c$f n tl

'f."\''tll)'gwmLp-i ~.dJI"ri,n£ Ell;-3:n iI fin:lle dine,

t!Ogl(llU~!'tld,d~ I~~ gll!tHil~s dH·{ll.~h ET biiJ!~he d.~dn 't~l:weabig eJ,1Qugh COfTIpnler. wtonc~, Es.

B~

pUle'r sd~!liI'~hHwith ~ ~jn wrn~ng ahe for

t no O:I~C knew :Iinmv IO!:lg the t;:QiJ~.1Pllii1lii.oJ1 mig~,t IMI'ke,.Qr even if:il was femiible:. One pel:s{J!~made it 1~'l:s.ible+Fokko diu CIO~Ll'I.WioiS <1, Be.rgl;U'I m;J!~,u:iIllllti)l;i~1II1 com~ <lnd

Theil di~1s;1:c=rS'lruck, l~ Novclubcr .21005, du 'C.~mtN\"'~S diagnosed wiliru ulllI~y,)trophic .~~u~r:illsde:rof;,ls. By M~ h~ \VOlIS b~lrjdd~1l) .,Iii.d cmnld b.~eml1.e only \l;.'ilh <I .fC)')pilltfMttlr. ~:But b~ wa" ·oom,p~e[~ly ellig.f~~~{r' Adi~~~SsaY$ ,.

'. M rudi1fl,um p.roleetl:ol'1l 891ainsl aerooo IS

'.'eIP I:h,wailfil~erfJI.P,:S, availaJtJ~le Tnten sij~es;

The tnew EppendlQrl 'ElIP' lJafiiiU,e1l' T.,I.tjl:,$.! w,ith~he'ir ID

Yn~(liln~' tWQ,~~:t.a:se ~il~er~ prov~tdie~hep-eriect5~hi,el~dI agairnst oont:a11l1 il1lEilitiolll.

Theriiltet COrlS~stsof two 'liisib:le p~a;s@s, eadl with a diff,ere'lilt potro' Silze. ThiS t'W(l!<pharSe fiUer protection ,ensures ultir!flal);! abserptlen of aerosol is D and biomol'ecules €i', oUitmatcn'ing alii ,oO:t'lv.ef1iHonaIM'e'I':s. R~y en it.

R:H' me m inJiorma:t~ont go to www.ep;pendorl.cQmldualfiller

Feati!,JII'IElS,'oii tih,ii;! ep [)lJIalfilllter ',.~.P.S., Doub~eprotection provided by thetwo ph asefilter .' Provides rnaxlmum protection for b o,th pupeUe and sample • llIHimate abscrptlon of aerosols 81nd biomolecules .' Fire!;!,rom PC R tinhibi:tor add iiHves f .' Pall'e'lit peooir:lg two phasemter technology .' SldiPPHoo Sltlerile, Epperridowf PCf! eleen ane pyrogen-free .' 1'110conrorrnHy • 8artctrJ-rel i1ltoo certiflc,i~dee a.vaili1lible
1

eppendor'f
In fo'uc,h with ,Ii"s
Your lOcal dli~~ributer.:www.~ppeildiOr1"co.l!lII<fw.Qrld.li.IffiIe +. A.pp:li~ation.SUp.p!:Irt: +4:9 ~8!(l4'666789 tEppemorf M, . Germany .• 49 40 fSSB0143 j Eppenoon Noon A~ InC. 000-645-0050

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I

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SY!.!ii!lT!d!li!.!ln~~;!ij:).p~.A h.!ridio!fl~~ e~ ~ ,nlltie 240 ~mc~so'~ lE,a's 'FlInt laUicp, with ,~ighl wlors irlldicalil'lg Jet~~ij'()I'!~hil1l.s

includes

fi ~,O( of

"IUIl it.;u:y.·' wh

represen ul~i~)ns d~£IIl ren't a ic hi ~r,e the eqlll h'11~elU o,f j~l.llk

Itchiimg 'fiolr aSp'ialce Fi.ght

'The batHe be ~.wel?nlhe Wh ite H euse ,1Ind ('on--

Iletw~n the roots.

"He "Mould lie nn :his back ~in l)iOIl~.

\\'~Jtll~.Vi~~l projQ'Ctnr pOinli!lg ~l d,~ 'ceilh~g.l \""o~.ld~~ here i!'iB ..HimQre••md he would see onth:ecei~i~

DNA iarhe lnuuan gcno.rn1Ie.Nevettheless, AdmH);. expecrsrhe \vorl. W h<ii~eapracticnl ilU~CI vel"Y soon, cSIJX"CimUy 1b!I":unu.1rn~lbc'r thenrisl>;. "The l)'Pi1ci.! I tbi mg iis 't~Ut~:I gat ,I e!l U fmL~l. a numher th~olri~'t whn S.fI'f~ 'I hrwc sueh sud Mic1\i a~~r-~~.~.la~i{li'l .•C ..n )"0\1M~ !'i'W: if it j:s lmilwryT .,. says Ad3 mI,U; , "Until now. that's been :ol \l1;:iiyp01inrl:!~ Pfilc~h~re t(l'figt~ ru eut,"
~:~;lOklUp."If

'gre-ssiol1<!ll D~moJ[r.ats over NASA's budg;elt

d,~mim!Mll d t~V(l ~learrii lim we!!'t Sen ~hl;r !1gs Sarillllra MiikuI5~.i ([J-Mm, 1,\1110chaks N'As:A'~ I rl!! rQding lPaln~l. wId N:ASA AchTll rli~lratoll"

'\:rhm

1W;I[;; typ:irlg.W~ would use Skype to talll;;, AU he wmned totalk

,;lbYUI

\'\I"dS

i.H"lhemu:~k::-i-he never

oorlffllplauned

Ihat happened~o him,' Du Cleux f~l'IaHy srresmlined Ihl1t ~tlf[\'\,I~ti,; ~!'!ough (h.U,l super,(;omput.e:r m ight be 3ble 10 do th,e ealeulation, Bm :l,e di.t! 11:0t live to see its l~ump,~etion. He died (j'm! I 0 Nmrcmlru~'r 200'6. 011 ~, J.1I1l:l,1JrY. Sage, a s!JI,per~ ,co.l1il,pute.r!It tl~e Uillrol"(;;rs,uty ofW:ac ..hi.!l!gtO~1.~~.lmp~lt"d

abous d1is ~e:[u:ib1tethi I~

\~:'it.h eheartas, it ecnld beeome <I simple there were such inl '!l~ m'•. I"d bLAyil iJmnedi.nely:" SOl),,!; Pe:ter: Samak, a number
M~lthelnal~:c~:!.Ils

at Pri neeton Ul1i\;'~ri\:ilY" .]nd physicists look ll kdy H'I ~l in~ Nl~ df!~,l~$~ for }'!~!irS[O eeme, .iierun:mm Nieolai, ill theoretical phys:i:c.is;t:ll titl! IlInav€.L'Suly of POls..d~mnil1 Gcrnmlll;l:Y •. says d'ltl ~in~erprnrlt:) of~ can be fO!JIlad,IU over hetc'm~ ie ~tn; ng, thml.'Y,. the ~~lOSl: .P'OPullar vel"~ sign ofquantum gr'lvi~y. "lf'yeu ask nile if til i s wi] I he helpfu.] tumor row, [ esn not ~:Hy
y~t:" 5'IY:;; Ni~g'~,ai. "]f! the eud.] 'thh~k rhe syrnrl:Ll,ctm:yof qjl'I~~nn[r,t1gl'OIv~ly nl:ugh;~ be real-

theorisr

Mittrlael Guiiffin l.ilat ~Lin,~rei~ :5imp~y ~(lO rnseh p'~ssu~e (i!'l NASA's. budget "IHer pn);pos~d ~o~lIlttm--.add~tng n billion liO the a9!!nt:y's S 17 ,3 biimofl :cN)O'h'e\llJ es l~~ Id rei.lri,se a bijM rt.isan eflort UuL ra~ last ye.aJ. led' ~rUer the same dalY, Home 'S(le!'l[f' end Te<:h1IiI~'lQg~ CommltU::e {h Cliilf' Re pJiE!sel1:ta ~ive ~f!,JA.SAi~h~l'!d!id (:or a~!railn1wF@d" ~ Both llawmilke'~ rl~iHH1Qll ~iel1{e ~'l'~~llI1e fobbed! 00 pay rQr rising s pace station, s hu ut~. amj '~;'(lPl()falion eests, ThE1Wh,'te HOlliS!! ls like'ly 'to oppos.E .~ bigger lbudg~t--ANlIRIEW IIAWL!ER
B'~lglnall1liBiaJ!l::k
BlIn Gomolil OJ-TN} \lIilm~d GriiUifll.~ilt

TURI'N, ITAI~Y:=G~O¥arll!'!i IB~g~ami 11~'S relumed 10 the Itil~illn sp~ .a.glen(v, !his lime is ]11)l f~uds;h~d. Like theHuman ~ed :~:Ulfl more subtle '\,,<1)' lh~~n weundersrand .~ its 'ChlfiKlw. Tlh~ 'Sj)aCi! s;(i~i!'i~itSt was s'v~iuinc yet. III dUll event. it wi,l~ be very useful m Genomt: P:~ujecl. it has produced f<lf ,aijlrlil'/;;tt;)J hQIITl19'9'i I;Q200::!. re~iQnin911lfter 109. mUlc~ data ror mui.hem<!'ti.c~,<!!!~, have •.1 ,gui.de or alias." '-I!!tJtAMACKEl",Z1E It~ly dedded to sa:veS 2,0 ilflillUQIl bV hadIng (lH to <1!'ls,~'m,i over,1miglln. Ad so. i~, o.~1ll!1 M~ckenrie~ i! wnm in5<!nlil C!'IUl, C;!llirl)m~ late its if:llrll mitllllll'llt to IbllJild 11r ada u in~lnHllumt f'or ,~ NlA5A slP~(;e(lrah goi og to Ma,~, IBigtnamf says he a.nepted the lob, 'ofr€'rl~d I.aS1W€ elk.,beyuse he "s,lra:ng~y Ihe 't:<Ib~e~bJ:E~.:BUl the AHasProjf."Cl :
b ~li@'~" ltallyi:5

the last (;luH~yiu)

ill(Jiribit

co IHrniU@d

to J@Lil!IUP1loChiJllg

its :space reseolrch proglram. Bignal1'li curThe U.S. House of Rep

re..'lCJIt

atives has

have

Sp;EM'lluxl i~ ct):ungressiOll!l.IUIl'V(!Sligal.ioll.

for go'!{eU,"llmeJllu s.c:~cm:usts who dlan:mm.hall:lli,eir basses t hfuV~ und~nni!led til€' :sd~iruific precess 0[" suppressed ,illJl{,11·m;illlioIIL Last week.
.IlJIrOlecl:ilODlI

bro adened

Ross SilYS the bill! is needed because curreat protections are '\!','Orne than ~I seless" <11 Jre£~d<lia
tor}! :;;~geml.d~s SUChYSlh~ IPOA

and do

umtiy (nal ['S. t~!! 51!Ji'!(8 S[~ E'mlCI! ~dvi~(lry" co I"Ilm~ttee,r,or the Elimp ~a In Spi!(let Aoge'f1c.y. ~Hi~ h~glfl~il!~eiF~,atliCi!r1li!1 pj'\(im~ win he~p Il.alian s PllUl! s(n ~n(!~," '5.i'!ys Milmello 000 ki of' lh~ U~;""~f:sul~ olJ RQ!TtI~, L.a ..
S,jl~l~!'!li!. ~FI~NlCI~~Cl'll!) Ii! 'IRf!W!I~

IH)t

cover scientific d isputes,

le,gg~ll4~n~par:i:.i"'"cl \'I'ti~~j;'blO\,;...;'lf m·, The P

lecl~,oll Ellhi!Lilm.c;e'lIle~nlU: Act. by~ m.ugin wide: cnt'l~lgh-J 31 [0 94-to \'II ithtlt1~Qld
p:ro m~iisecl vet!:l fH1IH PresidcUli t Ge(l~e \1i,I: Buz.;h" Bm thL;:m~<l;t;~u'~f!:!~'\;~, uphilll ~n
<no [i;J ..

ad :~nthe' Se.nale.

But ethers :Fel.;il .~1)ae biU would fUfc;e t hI,"; :s~ielluiric questjo:~M ont.Soidc their ~x~~Cr!ji;(': .. ·'·[fan ~gc'Ql,ey or Ihe Adln ~n~ stf<l,t iM d:uSOIlgrecs w,~d]IIhe fi I1d~~'~g!'; ·fi f a part ic:u ~,f~r:;G'.il.;lmi:>t, 'live shOij Id ill!}! b~ opeuing up om:- jndici, ..~.syst,em fCH;' tlm~e
,c;'Our~s 10 add,I'!;!:!;S

ReflllO"oI'lli 111g '~In'e101 1];'0,al ns;

111e (I@ ~ sde~(e ~(lnUm,1 !'lily is reIJf!,iti 11{1l"v~ ,anl., Of!ll~llli.l.a:tiQnS ill hopes UlIi3't.OI"l.>e \!'okeli\'ill. speak lcmderrro (o~gre:5i'i ilnd the White Hl:Ius-e-". The (ol'!soUf um io~ Ocean Rese.archa!l<i Ed ~'ali:on KOI1:E}-<l1U~ar·old .advocBte for o.:e.m r@~e;jjrd. ,oom::atiQn, and polity-ud tliJl! n·member joilll Oooo'mogr1l!phk Im,tijtu:lions qOi)-!!Ilanlagelr of large-S4:aie' WEs;eaNSh pro'gr.a1Til5 !!l~h OOet.1lll drilllllllg:-htld ~~Ilil!ra~e{j s lIS inl 2000 a~ Q lim~ (!if e~pa!ildiI!1Q SIiJp\port r'(lf O(.e;;UlIr,~e.aIF(h, BLlll'low, rn.a 1'11" of ~hm-e projoc~:sare be1tn~ sqtJ:ee'zed by SOilr1mlg (Iii p:rkes. 1Iih~ IIIiIt rued o:rganlzatlOfl wUlpfo'l,i'qde ~a, uflite,o \!(Ike ~m ih~ QOffiilfltliilily/ SilyS ]01 uwwd d~.a~r Nlarda ~~~Nlrt.t of tnt iWlgifll~il'ey~y AqooriUITl Re5e~n:h ~nstiUne '~ffi! Moo$ ~Elndif!,g.
(;l!Uf.ornii!_ Stev€fI Bohl@11i r'lOW :rUlflSJOt .ai'!d r~6Irerl.Admi ra~ Rici!atrdl \IlI,e-~~ &:lens IJIP (ORE. h -lAj~(Iili~lriIlD I!:IE.R:IR. A.

Th~ bilL HJl_ 9S;, t~(I"\1r15i,wid~ms i~l'\.'QI dll: ·'di&5I;:mimu"iM Qrlld~ or !His" \fhlg
leadil'll!,l':'" sciCillti,~!(;illforlllm iOIl or "K:'IDQIlS ~htlt l"Ol:Hpronmri:;e ,·th~v;IIi:d ily'" of f6'tle.f<JireSetlfCm

d~~~igro~!n.;:~~s,to b~ I.itig~!'t:~d .is f~dl,;:r~1

'~a.npl o)'l.~ perMJln:nd i :oiSUII:5.·' jOlid R:~Pll:8I!; n~ m:cive :I3iUS~Ii (R- :I[)) dl'tri.b'nlg an UD1sm::(;ess.~ ful ,~Il"o:IflO:ITU.the Iloor!o ~cmp' t~,e 5f,,;Jien('~ pm'llu.sio:~lS. Sd.elmc~ pollicy expert Roger pue.lke ofdlt: Un~\~ersil"yQfCoLor,lldo. 8cnll~ de-f. \\I'() [lclers. !~m~1 IlliteprmrisiolH" wOll~.d be
'(lHf(l,r{':ed,pn=-iH~g <'I

h defines thosc!lell0ns f~S~n ··~hL1l:$C autho:r" of ~ ity,"tlre k:.gusJ:miorIJ. \I,I~ich '!i\,{)ldd,lihlo ~,p;p.ly m ~ h~~t;m(e!> in ,,\iil:i,ch~ovemme:~n sciel]~'i!Hs,are ~ 'Pre"~m!~d fmm ru.!b]~"hung d~Hl ~!'l:Clrl.!oos pro~ Qne'Ili"~_'ovcli1lmelli~ .,dvoc;lu;:"~ S""'j o vis:tons t~l.U t"

situ<'!tiof'li

rI \~I~~ ic:~."n

~ win

S~'J,'t; 'i;\11!~Slh:;:blOii,"'("f$ il b~u..,n:'I~!l1ce

l(1Pfl;;·

redel'll I ""'n~1.1t and to m:.oid relribmiol'l. "II's; \'\'lI'}I imlP'm·til.mI:t~·' say.-; Di!,vid RosSo, :f~ z forl1m,er Food <In:dDnl,g A.dlninistrntioo (FDA) !;tam~r \"~'.OgC ilCC~~!HUiOfbS tho:'!tUl.e {Ig:~ IlC Y :i] ~ i!;llllor1:d d.al:jl, on ~iver d.,m.,~,e :i,,~,p.uit:nHi
fiii"'i!.il
illl

~11u:xlr.rec~VI:emin.er forec:l:Sl OOlli.d be ~;<Ibeled ,m ~!bl!S~ (i f m! Iho.ri ly bee ~nlse: i~ (:lln~ll iit!i;;;d

F,.lst: infoJ:lmniclun.

I
.,
i!:!

A ~.obbyllstfor t he UI1I~O~ of C ont;:~r~led Sc~en~is~s. which ::>upports th'e ljegi!'iIMiim~.

;].~"mcnts C~1U Idhold s\~lIi1i!l tJl1.e S,elll.ue.\"'·~,li(;h pl:~scn~ t~"much more ~iYS
th,l[ MI!;:h

~I(m~

,f!ppwoviil~~h~ ~t~bifl[~C Ke~ek in2l0rJ4

oe:b,1~:~I;:!1.!;li ng"

{llnd!'onlll(l!ll.

..jEllJl KiINiiI'i51(]11

1649'

N,EWS OF TI-I E WEEK

MIICIAOBII!OLOGY

INew Bacterial DI181810S'8' Aga,insl IPha,geInvaders ldemlfled

!~:NH'd~pfDbl~m 'with phuge:" foi!1Jing lh~1j]' fl'orn beilteY.ph'ige~ under 01· ·c~1n.;hIlUthreill ofin""';;~~~lll1. .~,c~~!:iiil!. 1\j~"]Mml~bll,ct1i:lfitli S.tl~lun~ B t:o:r example, are awash in a sea ~)I'viruses The D'IIIlU.:SCOteam swosk also provides k nown .1$ ool;tc~:i(}p~~~g'l::~" '''Evl;}I'Y 2 d;IY~, [l1alf i~ellrM biollJ'gic .. ~ evidence for .~I·m'H;:ti,o:l~ of ~:h,e bm::te~i'l on E,rlit~,l,ne U.led [by bi1JC~eufioso-called CR1SP:1Rsequences .. which were ph;clgcS]I," s.~ ph.fIIg(: expert Vincent H s{)h~lM: i idm~[iri~d in 2002 by Leo SC~lo~LI1> nf the of Roeke!eUeJ Uni\l\l'!II,';Si[yin Nf',V York Ult;y. N;u]olml'lll.nsti.1lUe of fl~blic Health in a C'(l~ ~ntlJt lll;;linl\l:"~~f~f(:ih~p' ha,,~ nn\'!,I BUlhn\"~!'!. Il~e N(:[hl3rkund~. allndhis colleagues, These sequences, fbrun<~.lllyknown idel'ltif~,ed'i ne\1I1 defcIlUiI'l1ll1l,ec::lKmis:ll'Il!hm hc~ps, b,~ctCI\il,1 1\toldtheir OWl]1 1.1:1, b>llR!C. d~i!> hy the dC;l;C.ri1'l'tivcname or·'c!uslert."'(ll rcgl.lOn p..ge [7'09', ,I t,e,nn~tld byPl~.ihppe loldy il'lh::l1i.p,m::t'id s[lor:t p'llundlmmicrep~"il:S" Horvadl. and .R,odo~,pbe Ba:rnu.gou or ooca.~Lsc oft[,e ~y Ihey ,31:1:' armng~d i~mIhc gelloim~e. are 'i!\'i.de:~y distributedin l~,H: Daniscc. .~D.tliiish cmnp<:LllIy that prcduces ~~nomcs ul'bol:~~, B~ciiCri~alld Alich,tca. 1Il<1~te-~a] '~IJ~l1,~r.Ci>~~d ()th~r mSilil,;:rii'!I,~Ior Ibe : t~~md-prg(;l;:~'iirig hld~.:Hry. fl;ip~lfh;h<!ll baete( AI,;·C'U~l.lpm"!lyhl,g ~h", ('Rn:U'J;'R; sequenees ria use a sysrem, ap;p.m(lIll~:I:y akinl t:n t~,u::INA <I1't:: a su itt: of perhaps fOLU tol (I cas ifi1l(~rrerel~C'1il (RNAi) !i.y$u~m on~i .. herQrg!ln.~ g I( CRISPR-ass,od~,(!d) genes, Re~(\..rehers h'flive Imide a number O['pliOpOsals.,about whflt isms, to block I1h11gcreproducti on. thus lllil~3k~ illl,g them R::i;i:i't.~mio ]IO!eclrut'll1!. tl~.e$egenes misiu do, For example, Euge'ne The \v!Drl( cnuld lrelp the food mmd Koo~.il'l andKiru Mak:m:w<I on~c U.S"N.at'[olfl.t] bio[(;;dlllul~yi~l~.hil!llrk":S ..\:\'bi:ch ~i$'; b<l!C1Cllri,ai C~IlI~rfor BjQWc~noklgy ]n~urlll~t~o!lill cll~Hl,lres.o mak,e p:~odm;ls l:>m::I~.:Ischeese :<IImJ B,elhe..<;da. Mm;Y~;<llmd, and their celleagues t Oi!!l,~ly!'~dIhe ~,(fS sequences ~~d. based un yo~~.lrt as wsll ;;'I~ p:l'ate:ills; ftlf hlUm:u~m~diCtiJl'b(!, These ind~lsjltries,.:r-ischeui So'lys. "have a ~ll1.o;s:e strucnnes, Sllgge.~ted in 2002 llinl!lJt they
Hi.JJ:r1HU'!S iiltO~1;l im! H~ pa~]hlogen:s:even the' simplest
org,na~Slll:l:li

.!In:~ot

h1.ving

fe'nd 011'
ate

Cll~:t~l;re-s eeuldbenefit and

·'urs

Im/.~siQI1 •. a"cteri"~tJ!<h as this 0 I'lil millY' .1!~;qJu~r>e kll\V dllhmsiYe1SlHI'U Bl'lce~ Imllilil'lfE1:diOIl~ bacterie[ph<l9!l', .~~ta(ih.(ldl 't~~, at n1iigllt
cllcmJi.;:!lI !m(.'\V

DNA ~~pair.liY~wt:l'~, But

nlil(]rl;.ir~cl;;fn]y,. Koon ins; fly~ •• U1U'[heridea enu::~g:ed ,<IS :M:'1 CI:3,'1gl'OU'~ClS:found lh3.~ the
I

spacer sequences wlthin ,"RISPR reg.ions

.~~S(ln i

Biol{J~y

rese mh~e those of sequence» in pJrnage mild plasmids, small exnnebmmcsomal pieces of DNA t~,m:can be rransnsiued b.i;:hV(:"C1l b~c[¢:ri,lI~species, I n <I secoad i!I.IIH'Llysis.pllb~ished by
Dil'li!'I.'1
[I:~

16 Mart~h (lf~;f~sty~@r.

M<lkarov<I.

Koon:~I1. a nd cnlle ..g ues a-

GE.O!lOGV

(j~(}Ingi~l$ have di$~o\a;rud l~ l~'!rili!il~t ll~(l\!,ill remn~l'It!i-by hi ~~ ions of YC31."S.-of P~i!WteeW!!rlUCS, ]~c large-seale l\lIovcmcntorEar~h's [ crust, The KICks, .an:::' pres-eni:l;.ld in pd!,Jin ~igThn
!i~Umlg [ilL;

Dm iI'£: fmm 13l.1n1!~ early

lion
}'ei:ln;:!IgA

adoleseence 3.·8 hi~~

dl~ $.L"'~Hoof' frc]!'!1 ~1.nd~liyi!~gil!:~s.. reek thm d

solid ified .i 11 magma cham bers th<'ll fed the dikes, and uevcr-melted 1'U1'2L]]tic !XH:'k bt:]m'V

~ncJ.;'~;l~cly SL'!.ld:i~d illld~'~lI'Oeks of south west G fee niland. <I gro up of ~eologi:s!;s n:rmrt'O ~mp1igt.~ ~j'CJi4, The;;;.$ d,,)'t', ho~'1~\1!i' ~,~ 2006. Fm:I'J:,:; h<;Lclll'lHg Mml~ed mueh ygL!~ger ntlorR.nm~ from l!il'lg!ru~l midl-o~,m rid~~ ~ SC~IPJ~fooe.'lll Ctf.US[ llun: had become s;tm:~.d~d o spn.mdsa:\,~' as h caels, 'lI~dle\','Cnurul~ydives back imodite deep :hller.~or;ln irs ,~;;qrl.y J:~l)'S. en i!ald c;an~dopb:iolit~5.bl!l '~~m' he "Vol,'; ~ dllY Efl:U'lh W!IS st U~ 50 hOI t~'1I.II':011lgl~mH that ~(10k~.:~g sea-flncr lavas lrlrn.at miglu hold for researchers ha\l;e t\'Ondl!filed ,,,11etmer ]ftle p~~If];et traces ofaru::.OC:nt m,itmbi<11bor~.g.s. Ti:leJ:l Fumes J!IK~his c;o~,le:iilgueg.came lq){)ll mi!!.h![ have been I:iddi ns :~~elf ofheat by some t~ ~ll,'r.;:qed dik~:;..These band~'d ~JZb ~ d)~ (If!I1!rely d ifferomn~~):iJib~. 8m dle tl~\\r d.~soO\'~!)' .. illdrc.u.es. ltn.ere '\?!'3!S <I, .n~.Qderl'l"d~yl~le tt'!cp hall:llrn:!l,;kof o,~I~,io~tes an,u! thus of se<l~fl(lor i tonqlcs op~ni'ri!1g:;:htyrUy aflA;;.r f~1irm~lq(J~uf ~p~H'I.U~. 1B~!illmke,i ~rtek of e<lIids. ~hey~~ E 91th," S<iy~gea~ogtist Vi fd,irim Dilek of Mfial1l1li cm,nposed elUi.r~'ly or the ~llll ~heeli; of 011(;"'" i moitt;:11 rock ~njce~cd i~lo I.~.C'crests ofmid~ UlliJ~'cr.;it}' i.~ (lxffl!I'd Ohi~. Inmlull1l,~emb,te geolggisl:S !mv'e 'i,'I.'<I~k;ed,md oce.m r.ids.e',sOI.S ~he ~le\v~lbrilled. pi;Jtes sprei1ld rlm!;IU (J\'~r is(lm~l\:\''OOOllCen~~nd'~12':kHmnetE:rfl\v:~r from the ri:d1i!.~'"1~ls~!a ~hoomd dilk-.e:~ ne<~r~:me'~,QlI id]el,]t~fie(l compnm:tns oroUphi~ sly ~,oams s,~lreld~ofbak!:I\':L i:\",.iste:<L WInd lonuJied rud kfl(}\f\i~l';1$· the: 1su;;'!s~lpracrus'wl bd~_ ~)~tt'5: d.isd ~t'Clive '"'PWO\,Ilr" I;;i:v~ e:"trud~d on i

thelsua rockshold cities to how 1.lreYOU!I~g~l:~mtN\\I'Orkcl'd blCk Wh~I~,IiI~ ~l:ug;~t h,I\~~ go~~en s~.lttoo..~I.lfac~. it '\roll' the :s;earc],llJor mi~ro.;;C'op!ie s~g!1iij)oh3.rly m~that brou~ln seo~og~:Sl: "f:<l~Jd fumes oft!h.eUnh,ren.ity of H Bl;:rgil;~!l.N~jJr\.\i~}t.ll4'id tCO IJ\;: gli~sti) ]S~i~1 ill "

IImt. "The nmj(1IfCOlllliJ'Oriellit.l ['Of,U1 ophiolite]i ,nl~~~~~r be all there," ~3~ g~o~ogi~c Kt'm to G:milie of the New Mexico lU1Sh ~ll.e M.il~ing ~ of nnd T'e~.im()~(..gyi1ii! S!X\Jf,!'O, "I'm ~~)I\~'i!lC~t., Sa lE<.trthhad sea-floor :;,pn:Oldil!1lg ulmosr .l ili IIian yelllfS earl ~e., dum p:reviO'u1>lykmlov.~. Wh.n "lbol!~:d~t:other end ohhe tectonic process? 1';oday, old sea "nor dives sliC~p~y iitHO the deep interior om. lop ofa relat h1dy coM,
rigid s]!<Ibofoo!!:lonli,cplale. ;;! process called subclueHun. BlUt S(lm~ ~~Clphys~cisH:;hfld suspecled th<!l al{l OC::C3i.Il, phHB miglu once have fe~yd~1.! lll~!nse]v~s dIi:~'f'i.;jrf;;m~Y-!}IlY.b:.Y s~nkillg )1.lr<li:ght .~nt(} a hot ll1iE1.gll11l mm;h <Ill .if ~

il were (luiek.'l.alld], I PUnleSDW illks. b\ltC.1U··I: l1m~.re:.~h'l~ :rolhe:- ~ Ibi~g like madem~~~ s~~bdlLt'[iml 'W;)S go.il'l1;l.~
)~<LlF1l, go. ~{JC~ a

on ~~ ,5MI ian

<!Idj a'celli~tothe ~

]sut! (lP~li.o]h(;g:e(Jchel1l~.caUy j]'e~~~!l1bl,.,;: a-

, "'I

'

I-

('m~JSPR spacer sequences. w~kh were 'pre,SlIlll.1,ably picked UIT'by [he bacteria duro iI,gp~ior phage in fec~ions. ~(l£el he r COIls'ti~ tutc a 'boa~l(!'rimJ I ini:llllIiJlJ~~system th~u \\lurb by a meclHI'lilism $ilL11ilmr t"(.l dmt ofRNA,:~ in h:ig:Jtcrol'~rn is.n~s"Ul e idea is [hm l~C slwC ~

PU'~)P(~S(ld. that

lll~ C.lIS P rOlei ns and

U'I keep b41:CI,~riflp~~~g~ lmc.~1! conl'rot"

Dennis ,RQrtl'I,ero. <I :~.l:embell"oflhe [).allmisco tellm nlli~e oomp<lIlY:~ bib i:lm. Madison, Wi s-.

New :St!l',aJ~elgr''tiD f~gh~: AlD,s

PFI.ETOI1UA, sOruJnH AiFlfrUCA.A.lmt\\! g)ll\!ie;rnml'lIIt pl.. !!!i liTlS~~Hut So ullh Arril:l~HIV m irlf'~li0f11 J1!ltf in Ihi!l~ i!Uld. UJ'q yadr !J1l'~eline: n umber (Ii~lnfe(led pe r50ns re(l'~'I{ing i! nli·

consi,n .. ~~[y~ .hat lh!! CR.ISPR.

~li:1.a,!;flm~~ ~HUj

.\ly$.!e;~nml,lHy h~vc~. whj{,'f i.lmclli.ol1las well, "Ill ,m:lJ~lioI1l10

m~1liChi111; pl~.tge. the splJ:!:1ers,al!':ol!:ll.,rch ehmpi.=l.1>Ili)f:d scquenees," [he .UilOt¢S. ~md thul'l< d'i~YMight help OOi'ilfni I\pml~!] bae~:er,i;dgene 'ICt.~V~,ty. \li.'1~~'h1£r~lftH)~,ih at is the ~'1:5A;l, t:h.: find' img;s;O:PI:'I:I the deer to us.ing the CRIS,PR sys~em10 Mock sp>l:>c:ific gem~ <lcli v it)' ill b"lif,,~t1;lr,lla.[usr 35 R NAi is used ill h i~~U:::f ':gan~sms .. m Ami [hCIlI~(;re i~ lPo!:U~b~ i:ty or produei li'I'M the I [I]iIO fieph;lg_e -resisraar baere rial srra in s Ibr i~dl~,~tri!ll ni~t10lii~d e,l('cumpli sl~I,'d l))! U~. b genetnc:ally engineering b.JCt:er:~OI \'II:ilhappm=
priate CRJSPR sp3t'er sequences;
$ill)''<;.

f@lrovJfa~ tARV) lnerapf by 2:011. The 5-y~r

ma,l,e'fJlI'. p:r,~eifll~;eri at <II mnrfe re m1{e laSl wee k, 5,el:s 1~f9!;ls t<l meet l~ne(inl~itlill elillS mad e by SQuth Arrkil'~ '!Ike pr6id~llIt UIII Kenilnir D (Sc:h~'tlce, 1 OeceJ11I(xof 2:006.IP, 1373), 'The 'g,a'!l~rlmnn~1lI1 iU. asl( IP':IirQ~amenlfor m1~alrl.y w 50 bil.~iiOml,<I Do:ul4iO% or r"ihkh would pay fo r l. ARV Irl'E!d~all~Dm.all1d 'WaMS bllSiiII1e:5S diol1ors to match that sum, Fuamo,i:5Venter. ~ead of the S.o~~~herl1 Af rican HIIV (lin i(ij,]! ImSSod@ly. 'e"SHtn3~e thal ".!:l~me l~Q n 03: miHio;fl ~ South 5

t:r~ m~k~ ~h~lli't RNA sequeneesthat a

can

hind to ccmplememaey sequences .inmessenger RNi\~ made by illV~~dill1g ph<lJgc:'. "[",ful,]$. would block thei r tD;llIm;~I t iOil ~nto pro ~ ..

tein s- ,UiI d mark them for dif;;!l;l1<l.d,ni en by Cu:s proteins. scme of which resemble rhnse
km"l\'If1'iIto be imvol\l,~d ~lmI, RNA~"

The Danisco group

di:rot·~:G'vid.e:nc~

f~)f

\'!,! i ttl lin:' ill .. cre:r iu m S I rttp toeoecus tJr(!rmuphihl~".\,'!I'l~,ic::~, is; w.idely wIDd 1:0 make

n~~H l~ypo~~l~$is..\1\l(]rking

liUJ:i>

now provided

Horvalh

ArFi( a!lls. l,li\Outd: bi;t on ARVs in S ye(l'r~ iir (he'

p,ialll i!i lu l~yirl~p~~,mMled. . Aooul5.'S flf!ilHClI!! SiiHHh Ah'h:::!!'!salFl'l inrre.nei:!Ily H 1:V,iUlid reu gh ly' 230,100(1 now re~eive AIR\( 'therapy •. Rollin Wooo, co,-difl'l(W' 'of t~e IDIl~mol'ld rU~1J H!V Centre in COlipe TOW1'!, (.aHs. t he pia fI ~a 9I1'ea,tadvi;ll1i(e." AIt'holJlg h Ihe go;:1I. 11I11Y bEl d'~ffifku~1 to rn<ll~h, h es ~ys. "irs beuer to set tall] el'S 000' hig ~ lh'linI~D h lIve nota rgets. "

)'Ogurt 1J!~ldcheese, lh~rese"ufl.<hen-;, found

lilt!!

however; thm

"""I.tl~~CO

has no 'pl<1!nsto'

i'~.ft.,c:~km of~:I1Jc 'b.acte:liia 'wit:l~ phflge'!cad~, to incorpera ti;ol'l ofp~,l:iils,e-re lated sp ..cer r ~~W~Il(,'~ \l;'i~li',l fuCR]SPR ~gio~ su>c~ 'br~~. ter!f! became ~i'ii.:.;t.~mtQ 1~lr,Ii;he~ i~~~t'ti(i:l~ i by rhe phage strains [hat conrri OIJ led lill:ose
sQqu~!1iC~.. BUl·'.ifyou. t~,I;'

do til13l i I~ ]i,ght of eensumer eeaeerns t'lbout: f~e use Q,fGM 0 [g'e'l.1.e~i C'~llmy~l'i.od:il fied o'tgmlisms], Nlrlicul.nrly in Europe"" Th~ f!.':~1;i~~n,;hl;;fii' r'hu!'1!u!1l\t~,'ldhl s.impl.y expose' i:l<l.cter,U!1 to various ph'lge :srmn:ltl:sand

ihe spacers eut, the

resist3nce :U5!m.t" says Horvath, who WOl'ks.<lt. DmllisollN k~b in D.mlg.e-:Silim-R,Qlm~"m. Fn1.1'!(e. l[,Iiu,eream a:lso showed that at least oue eelS ~I~~. \I\,i~ ich encodes a 'possib~~ RJ\J\A..;tj~.~c.i~.lg .nlll.de<L."Se.is l'Ieces;s;<lry :b;~he'l.lh<1~ resistanee ..

~'~.I;lnelect ~br 1~1C1!'i!;: ~hm ere resis't<n:'!t They s U5W) their knlowledge or the 'CRrSPR :spn~I;TS hdpscreerl rOll bacteri .. to thatcarry the right spacers H) confer the IXl s~st an CG' t he y W~fM ," }\~ [hI) i(1g~ w~ 1;:<)1j.~
can. however, genelicHUy e:~'l:l ineer.'t Remero
fUli!ld

T~~is~ltlo\\o'S.Fise:h\)ui

~f,ys.t1HU ~et0ri~

have

lhat Jm.:U:l!:~ro: C1Ul

dnl'~C

"a vel}' neat mechanism by which I~~ey Ire OlbIe <

'Om lor us,"

!I"Y:i>," "we

Purdue 'W;elbr:almes M':aliln 1'ls:Ulute

Pl,lrd~~UillWer'5~tv in w~m ILafayelte, Il"Idl.:laa, has di ncht!d ill dBlILloioil1 whlll~he Allfred M~fHI Fou !'!o.al!io!il ~101P'llSwiUbif' a billttl!1dol~tlr·l:!llus netvrork to silll'phe rrt! u lliv.~;rs]ty b~Qm@di((iI. iF!~lIiItio rlS to th i! mar ~t Alhd Malll1instHutl's are designed! 00 be g O\l'e rnnedlby II bOi;l rd equ1lilly s~ti t betl,Jl;iee~ lhe uni\l'er~Hy <llndlthE1 (;a'liforlii1l-bal~l!tl MitliHI FOld ndi;llil)l~,Somel,m i~~ es hil,ve b~lk:r:,d at r~;~i the proRloi5oed <:Ina mlg,emlen~, r,~a ring tI 1.QI5:5Il<r !

~J MNI,MAII!lJil

Sign.nure nxk. WrlijcillAy l.ayelioo ro!;k diS(o~erlld ill1 Gt1~f1I~nd conlirms. ptal@ tectonics 0 n early EiI~t!:!.

yea~. two

ulltrol

G~N

No rlh (i.!J(I~il'lCi unfllersi [res lurl1e-d Inlilillm, dlown 10111 il M,1I1l11le'lldOl1'i'l'I1lelll (Stie-llce. 26 MflY 2Q06, p, llZn, .Bili!hQu~h

the~r l'n~e'U.ectual property.

Last

Mark (Irowel~, te(hnokigy U.8ll1!Srer oHki,i!llat lh~ U!il~'IIersilYof Nonh (a~olif1li1, Chap~lHilt '~ays til E' dom is'iitilll '[D!lli!n to 11l!l{lolititioIllS. Oil~ Ilrolli~iO!1 g i\l'~S,priority to ,If'!,{I !lalla
(omptlniiesin Lke'nsl ng 0 r Ipmd181!>i teclii· ng jiJ·oIQgil'l~tlll!~l(j!pe(J a!, Ihe ir'is,lit~le, Pu rd UiI! Presidelll Melir t~n.l i~,hke Ij\IQ~' tells,;;!i.!s:!i othe r "They'"ri!' ~~,md w e:r.:.plaill in a:ffiiYother "",:".)',," "(mdi~ C<!!!fl' ,qtllh~[!~Ne. "I dOll'! thi~lk ifs mO% die fil'lithre,"· he s<ly::>. "'Ti'JJel.'((sjm;t (~Iloughi.'!~],bi.gui.ty~hal it may 0,," Ill,V)' not" have been ~l'Illi,rdy fnQdenrn s1:JI,bdmailon, EmI.ough ~mbi,guit}' thlat h'l~a:gcologiS[s \\Im 00 ,Iineadingo..lcll;. 1:0 thefi.cld\'!·ith 'liI.ew eyt:s lfui s Sumt~ler+ ..J({II!(HI!I!t!D IlfFlR A.

delao's ~f lfl~a~eeme ~t,allW~li F1:e:ed 16 Marc:!:!,

~ rocks ca~~,ed, on~niles. lillese' rocks <I:re b cooked 5l 1Ji.p !:mly bel'.!!i;~~l'~i!;:I~1ind ehl1ins~rc~ed ui,'er

'Ii

,r.mb<dm::tiCiIl!1. zonC:!l like 1iI.101i>ein tod!ay'S \!.'eJSterll ~ p~H::UnC" 11·Is.u~ Ill<l.Sbona fide bonillitc:f;,.[1 ~rnit1.gll1m m~Hlh"muld 11m ,'{urk. ~ A~I, '[[Ime ,lsu~lJocl~s conl!C fro .." ··'0 pr,~:uy r ~ \:vell~. est .. :bl is.Tm.ed subd ~l!CliOJ,l zone s,i'lm1I <Iii" ro i

but ~Iys e\l'~ry{lne's IjIfl IiY ih1tp,!)y with ill. IIAa IiII'I iflll~F1dsto lirn:lltce alleiilst ]0 1l1i(l;re iI1st~tules, II IlIrlJi~otype lI\![li5sel up i 1m 1998 at the: Unive r~ily Qr SOiUi~herl'lCalirmni<ll ~111 Los Angeles, i3lul (I S~Qrll!J ,,,as (f,~~d l~s! OCltlb@'~ll f~chn:ion i IJInri\fe l':Sil~ 'IllI!hra~L

~wl~,f~[ 'we h~ve tod@y,~' ,co!lciud.;:s Oi le,k.

1651

Havilnga 811,8.51, Wish You Were Here

ilrhe LalrlglB H,adili'11UIIComm:h3!!,,at ICERINI will SUil,ilS.IiJI,pialli1th::le's at Il!JnpI1Bc:emie'lllited e'IIiIIBfjgy'(llliId nUllY €Ilpen n,e'w' re'ailims of: cilliscovlel",.llltwlilll~ ~laFH::lunFlIElutr'D,pei's ,asIllN::mdimc, ilil: p,aJldic:le, .,hys:i,c:s: for YIHlI~S'1m IC(lHlIil18
N~AR GI!NI!VA.

Arehana
<II

Sl~anl~at

III

I):~ly~~ch;t:~tCERN.

Ger.\'l,g pus.hes the \'l.l;dl0 f metal. "l fyt)li gel

feel. for Ihe resonant fl1e:quenn.cy .. you can ..;t~;;;:h~ to (lsci]~;;it'~. MlyS., Su.re ~~.ough •. it" he the gi'lIll srirs,"Okay ..ok.,y, it's moving!" ~~1,10ut~ ~h.}rm'i:l. s !llll~iom.0 f dollars '\~nrt h a ofddk"te equi.pllm~f:~ltsways: even'so s~i.g~~dy across thegl'3in ofth,c cenerete, Gen:viy j'!'iIl'1 the orl~Yene here who I!' <I
bttl(:g~ddy with I1JCn'DU'S exeitcmenr.Jn few mlol'l~h:->" ('ERN researeherswiljhave 21 ~ ~

SW~lnIEIRLAND-MellKU!:'.~·!1JMhj·MI~~Sf-e:!~e:rgypi!r~ic:l(i
The celossus

~!lill;l~~u~r. [he

large

I ~ meters a~ms:s ~n~.d ~~~eighillg, ~,3.0I0 m,et~ ric tons, the enormeus disk of ma.()I'd~.e:ry dangles fro.m bundles of cables ~ike'l ~iganti.c yo~yo. Sectioned :llike an o.ral~gc "ndl'estoO~'ir.::d with !::~ec'tdcOilI c a bles, Ihe CQlltfflpt.ion CO~I~dbe mistaken for a I.lying
M~UC~~ hoiswd

I hi dron CoU i,dell" ( L HI(') he re <Ill the E lIro~ pean :~~rlich~~~~:ys,ics ~.i!i,bm'atory. CERN ..
hO\'er~,1

few (ei'lhWl1tl(er~

on ~dg,I;i,.l]!J tact, h's ]X~rloff! Solenoid lCM:S}. db1!t ,,,riU

I)

".i:mv,~the ct:mel,''Cte flcor of.~ c,~l\i'crmmsS,li1ll~ terranean hall, A]~day~workershave lowered .utdewua s~lan b,!re~,y wideeueugh ttl take ~t The ]nO-me~rj(Jlurrwy slJ,;fli.~~Ehe nerves,

h ~I,g:cbanc:lj~:s haped partlcl e dlt."lcCtor, l~~,e s<lys.

Hubert G e::~"\I'U;g,, ellig ineer au

.H CB

RN.

(;onrp;a;ctMuon
SOOIl

sn~ re b it S

r matte

u: I~I"O ~ In he

new

No'llv~ha~it 's u]m(l~t over, GIL;;r\~'ig COil!'! ~,~elm, I~Hle. "\l\hrDll to sec it move?" he asks ;1

(,\[~mpl~T~d~h~ 27·kil;t}n'il~t~"-long LHe and ~ in Nl1wmb6r. th~)f hqj:i~ bJ' rH:l~ the ~ar~esl and Il1;OS.t ,COJ.ll pie.... experi mentel devu:ce ever i built thrHillghits w,lml-up ~'lp~. Sm<lshi~lg. ~ ]Mrlicl!es at cw.crgies; seven tU·nill.C's .higher than e;

NlEWSFOCUS
.-.IIIrBr(I~d~hC)!,!l]d~$. Pr()IOtlSWlIll ooU'id~u~ till!' cet11ters el' WalTi:, ATLAS.~rKI lhr\\~ c~heJ deitettcrs. S wiI3d.:;:n,"}\~u,~the U.S . .rccogu.iz~s: lhM became ~f8.inve:Sli·lil,~ <L lot of money iin th!e !;:'1(perime~lI.ll lrere,"
No one k~,ow'S'1vh.n the L.F.~C wiliU find. B;lbttl~i:s much ~sdeaf: The U-[.C ~10:!!l> ~lroady created Oil re\,(l~tuhm :~np:mi~de physics,. UIiC ,0:1" ~ Es:r (~,Go:-ris~,n CERN, "Yhu cou~dconslder tilt; I'] iggsboson the pe riiod Oon t he end of the shtmldl.!l:nl-mtlOdc1 sentence:' he says. Bm p hy sie ists h ore [he sl;m.cll<~rd model is net the fi naI werd ~l'Iult hBiltHe wi l,~bj:ll5tOiUt ol~~cr p'~ri.id:e~ m:ad surprises (!'lee p. ~6.57)" "We h~d :S1.~ph~l.l H.a\lJiki,m1ghere, :;Iud :~~cIn~d us

"!!!!III

tbt hI;:\W!i:illl'C w :ltlre wl;~\1 nnd th~ ~Hgg~ bO!5>O~and that hewas more interested in

mini-black holes," ]~~ll1i ~tly::;,··P'I;;CiiP~1; ha".~

diffe.Jilellt ideas,"

It is dimCLllllO describe
ecsonlng
10 S;~lpf:I.'latives.

the LHe '1'\,r.it~,om

Not only will the lHe smash 'p.lrtides 011the lmi£h~::'1~I~.erg~t::s. hUl.il \!o,'i]~ ;Jlwlieed tlrelargest and lrnost CQJ,U~ p]~"1.p1l:r~id(;i(,kl~C~(l1'S0Wr bu~hfoOl;;'! collider, d~a.~~in ~~:r.each cou'~dn 1~.;jlJstsek of DVD~ ~ 25 .k~~;OInetersh iyj:l, The un:: will ton8:lIIl1l'1.e .,1 reeexl ~. 0 mi~,~gi'!WHl of power,. e no ugl~ [0 2 ~~ sustain !)"ery heusehold ~~~ canton of the G Ci!nC,\>'l'l" At ~.cost of 4.7 bill len Sw iss fil':1I11~:f; {SJ,s. biHitCm,. ifs~~me 111001 e:!\,pensivtlcoUilder ever bnih .. "file Unuted S~.JI,lC:S, ~s d'Dippirlmg in $5 31 m~ lllon. m!:!!Sdy for die:~~ClOlf'S. Numl)CJ.'s alene 'c.mlliO~. o~mveY~:~le c immenS~ly,nHhe pmj,i;:ct, liowever; S~~p into the hOJllI
~l(nlsin,g the ATlA.S detector', :iIJ'mtdo~t fund y They will pump om the geeatesrtorreut of

Mighty ATLAS and eMS will race for these brea kthroughs 'It l~ee~]e[lj:yiolll ielr.
""It's going to cnlli'il.~ down 1.0 \:\"110 is better preparedand whose detector is more com-

plete" ,Yhe:~~ tHe starts nl!l~i~g. !';):ys tl~~

Tej:uDJiiJ,erVjrdee of hnper:i'<li~. OJUe~,e Leudon. spokesperson for the 135Q~I'Iillcmlber eMS co.~hlbora~imL "They can cOrlfirli1,l [our discoveries], Il~",t1;, IIIHow{)d." "['~1C' U-IC'
COlJ Id

cli 5C(lI\ier lhep~:rtidespr'ed

icted by ~

cencept (!aU(:ld iwpersyml11'!iC:I!:)' after runnii'lg for just 0:1 Y'e~I!'.Vird~~ \'i;l)';j. Sl!t the

"Soc[iety is wi~liI1gtD' pay

at

y(lu~lffao~ ~hi~':gcauhl

10

1'3(':(' ith w

:;!11~adl~ I~C ~~ig S1:nIII

ries Iklill 'iU'I~1 hallf <Is.~on~ as a soeee r field, The

ml the

mw", ora el.Hh~dJrolbut

certain amount and no m ore, ,and ·tlfJ mrake the LH:C poss,lblle"wE!! had t'o be [inrnouat,hJl':!",ul

instead ofl~n:HoOly Spirir,

eno~!gtlig other gil:mIlG~w

p,u'licie Tit.'lckl;i'S.~igI'iH"miui~gc:~y~~al$, ~nd

fiU ~,Oln mU/io'll d1~t'l
ehonnels, Amid, it'sas prec ise !'IS i~ b.lg. says is CERN'::.P>el..:r Jenni, spekesperson fer the ~.'UIO·m~mbcr AlLA,S (lollabomtlo:Ul,. It can ; measare the curv,ung p<11h cf'a p.rnide called a ~ULUlnto. within 40 micrometers. ~}';11.rthe

in. packed with

-Lyndoln Evans. CERN

W~dlh~~f <!i humml~~LI r,

e~,reUng belQI\I!,' llmc .F:re.un,c.hi cnuneryside

lh~ previous rece rd. [he L Nt" :iihO'~~ bl,'1M cur ld ~'he'one' bi.t ofm:luclI" misJ!.~llg from Ilhys.idsts· Ih~{}r:r{I ftll'; ~~l(j\\m p~irtieles, 11¢(H~ld aI~o SlJiit OLUa slew of otil.e'f particles <md open a !~~W era of disctw~ry.,. Tha,;:lHCwiU mak$ eERN t~l:ewQxld s censer for part id:ephy sics, COf!l;idlit.inS ~ts size aad t~('Ilf!o~og.ic a 1
cOIifJI~)lexi'~y.i~e tHC' "i sthemodern Il'qui V"<I.~ l.e.IiH t he PYI.'ilIIilr1Ii~d:S s ~Iy$ :Pcu::r Umon ," F,erlll1i National Acceleraror Laboratory ,t Fcrrni]lab) i:t!l Batavia. UI~naj &. who~s werkilil,g 01'1 tlre mJd'lline. Bltillhe 1JK' is n~,Q(we than a techmdog~.c ..1m8irvel. It embodies <I, iJ:1,uru:ielf

b09l'\'i;';~11 Lake Geneva to the ~~t and [he Jm,1 .MOlLIliHliltl!> 1.0the west, ~he!l~ocle!II'SiIOr itself

leeks Iii bit like ~ g 10.11 fkd ~\Vt;.r J'ii~~. V~~i[(iF~ ro l~.eL~:es o~h.eF\,,'O\~dlyrunnel must oI:arty
~Y!ifg~l~ rl,lcks.

i~(;~~'~~i),~~~,\IC'hin~'~t,':r)'Q~!li~ .LHr~\"iJ:~. !1:i,Sl!J fl1:ed LW'O spC'c:]~li.zcd deteotoni to stake daiinb ro i~eadeIf5h~.p orher lal
!Ireas

system le aks ~mrrocatung heli,mn: workerson b~k~!'i s~il~uk~!p lh~ imlu",mi,rot:. Th~ Ute en

or

or

lies i:'llo~,g nne side of the gemly Cl!H'ing

t.mmc::LAI] endless nile orb~!i! blne ,ey!i]l ders coauected end to eud like Sllm<:~ges. These a:r:c n'l.c re'!'uM.im~~ry .!])JZIglllCH' thai I. s;llC~r the
beams around the ring, Twice as strollJg 3:!J these at FermUab''s Tevarren, they in fact h ~J~~:;:e: aeee 1101'." tnr1' earryl It!::. protons ln f\\I(l

as: well,
Cll~k,:d LHCb\lr.I.iU

A detector

srudy 'the

mtl\ierl~~m. i!~l pilnidl;lp~ysks.Fur

d~c~!d~~.

p2ll.rl.lc ~C~ eontain i.llg eleurenrary bits ef marter called bOUII)lii!~ 'I narks> <Illd rhei raati matter roj,~~ .P[hJI:S~Ic;i,s;ts . at spec illl!~hred oomdl;:i'S in t ~J;l U!il,i~i;;dSmM:
.. ·mI.d .J <I. pEl 11 h~.ve s:tiudiled lhe subl.1 e' d~:rrere mlce~ lIstht;: hOUmt11S (I(:'C'l;liY to oth(:r "'f:i~VOl:;>;" qU<lr:k. i~;lI'n,QP~::l,!iJ 1'1ndi ng hi,ln::; of Ite'i,{ p ..rt~,~ C~~S (!kif:llcC', l3 OChl,be:r 2010,6. p..24!'!,) • :E,'ell,l ifAT.LAS and eMS s.ee l~,lOll'e p'anic~ll:~ dir~c!.I)i. "Y0i:,[W;;lJ'Utfl s'r)iJ.dy how fMngs

,I &yILulm~u'~c~ \ViC:I:':H bel

~W,~ n~ted S~'Me~ ~)i:!.(;'edllhe U fi!eld. No"'. ",l> the lHe 1;;di'l~e5 rerm]lOll.b~ Tcv,ul''Oi'i ~(:Illid,et. EIJI.·opc ta.~es lIb': [[!Cad. "It is certainly IrLJU':' .~ II'a~l dl~ ·cem~rof gr.l.vily 0 rphy:s~~ has movGlfI .~ ~o CERN:' says ~;h'ln=:i!-Ake Gm;~<It:Ss(m an .. :~ ex.pC1rhVl~IHer from lund U[i),Hv~n;i[y in

OPI)o.si te di mc::l iOIllSTheb:Ii'I.1i\,!,Ii'lY oe{lIUdel'·1i~tim'gs. fO[[elll~~i.'J1lu ti[scover one thi:lmg: ,I IOllg-soug~'nllp<I'tic:le C~lied I~e H i.g~s boso I). T h~ .F1i.~~s,o,uu Id compllcre lhe oo~c<'lI~.ed!!t'l,il,uJ'll1c;1lllode'i of rhe 1\11(]\\'!1 m'l!.~t!id.es. ~ys J unord]\\'!~ fin:; •.f1 s

or

NEWSFOCIUS
co~.p"lleto

CIERN's
Sev~1I1

T<'lt3~I,y3,

d~cser:l(r~v psrrictes." -~,lYS, Nakada, "Alld lh,u··sw.h,n

you call do with flavor

Ihq: U-!C wm sma~~,1lead m~~~ld, I !l;D1Ie:~,ie5 ~ 28: times higher. letting A;t]CE pee:f cle~pe,r
into lhe~Jeetillg plasma. 5il'l.}ls CER-N·s Jfh:gelll, Sc~mik"'dlt1. "111e llil,itlll,!:!lS yo~, eanlcck .;;Jl here

p]nys.i,c:s;;" kih:nne~ersa\li."dy. a del;e;i;;Wf ~;rmed AU(,E w~U study a soup ofpi:tl1!idcs eaU-cd a qu n rk-gl UOIl 'P~'~S:l.u.\.Th e l~ll:ro'lbo~:p lasma ft~rh~'dt'l~("~nnUll universe, fUr.)di Ilhysic.is(s ~Il Bro~lkh~wl1! N~~i[in~~ L;;!bU'f.'iloii'j'in U~m.'if'i. N e'iovv'OI::k, Ilm,3ve rec re ...led, it by s~nOllshi~,lg gold nuclei wjth th,~ir R~:'~~!th" H~nvy i«111I i:;~h: Collider (RJUC). For a fe;w weeks a year,

m:liIll!:ing 13m flrsr. researchers must cnmplete the coUider. Afl:er~ decade of conslruc~jon. rhey OlIll:ie (lin schedu ~e ~o ri.l~i !i1Ii!'; YIl:.H, say.s s,~. CERN's LYliQ.do~lEva I~S. who l~ead.s t~,e clforL (R~se-.?!rehC'r$ CXpeCIlO :h]\~r tht: last
magnet ~I'IW the tunnel in m~d~Apl'iL) A

Tw·o dle:ca,olesiintlhe

il:'or t11C;:LHe h~~ boon

11

malP;lthon. PbY-iiclsrs

dre<lInl the coil~ider up more than ,20 YC:<li1"$ <Igo eve 11 as, CE RN bu i~t another maehiue. the

l~rge Elecrron-Pasltrou eoU~.der {lEP).

which nu~ from ~939 to 2flo.O, In fuet. they pbnllecl.LO reuse LBP's HnlL'1el Ollrn.dWeed lill,e UIC wil~ ~X.i$li~~.g ~c-cc~cr,monL ··\VijthoI.U I~~l.t \i,i\rnuld h,av~ '~~~imtp~'i~!).ibl(;; :~ll' uiM i h I:~.e machine on 3. ceustaut rl:lll] bud!j.'e1t." !><I)'S

volee ~tml silv~r 113ir. Evans has the phlegl1l<1Jlic demeanor of Wr;:I~hm;;'ln with a
$njfWnlU~

(}n~

'1I.r~.·tlhas d!;lil~ t

~.\'ilh

C'lt]$'!;;:;;i

IUfg~

and

yo~tcan't l(lUk ,fit 'wHh RHIC it. for 50 years," he ,"Oily:!>.

~"€:rIifyllU

nm

smilJL Oill. his com r~lte r be p ~~nsUP!; mph I;l.flcl" gro,pll of p rograss on the LHe 's mlyri~ld subsystenrs. Din each a r.DS[ifn.g 'fed "jiusl in tinrn.~·'I:~nestands our. "][~,fIS same mm~.gh::al properties," E\,'~IHl s:lys."TJm.:i,:mil£s lend 100 bounce vir it" 10 "l<'l)'l on schedute,

('ERN Oi.n;:,,;j;~rl~nt;ltl~RQooift Aymn;r, Atlh~ ' 1~lne.physid:sts in the' United Suites wese plallill'li.iIIg rhe S7-ki.~.gnl[lor;:t>e:r~lung, Super~ cmulm::tuns Supe:1F Co llider (SSC). The Ll~C

]n ii.pi'~(; of the d,I,~~I[O the fil'li$~,. t1~ep'~;,\1h

(_~ouldn'l !1ilifl,l:ch[he sse':> clt(:rgy. but II cnul~d srnash more parueles, S;IYSCERN chiefscielll~· list Jos E~lgc~l;~.ln 1993. ~~l~ US. Cn!lgr~~ ki.lled the uncnmpleted SSC.[ie<ivill~g,t~ll~riie~~d

Sta b iIlity,llnternlat:i,ool,1 I Char'a ete r

Honedl'CER.N's 'Comp'etil'iiv'e Ed'g'e
l

blocks that [ripped. lI:P ~1],!: {S('iellc~" 3 Ocmhcr 2.003. p.. 3 S:)~~n~cmd of buill:d.ing ~l ;1,111 exi~linshlb. ofl'tL'Cia]schese a [oCm01:es;iile i.1'I

sse

'i[lfuI€!'

qualiitil@,!I~~ait: li!;el,el!i!l~hE! ~,a,iIlImlak~ thE! Lll1If ~li1llthe! rra(!e '~:,ortliile next {ilre:att ;pall'~h::~.esmaiShe:r

.III

[E!ality ceuld

put it a!l,t)~p aheed

IIIIiL~tleI 980s. pll)1!:i[ci 5.\:5 11~ rn!m:::1fI,~d Olnplmll!s, !'"ora.gall'g<:llltmm panJ:de: smlffl~h.« ll:Ult 'I'KI~tld reveal '!TIile key bit gf Wl.-r~i!e~r lhlllWo.l.l]d COJ1il1Jplete 11me~.rlhillory oftlme ~IIIIO~lim particles, Tin: be~'l~!l!,ot h ,\:l,,'olmld liI5.QblOi!!'l 'lIM:l scads uF l'!iI;;"\'V ~ pajrtuel~ HIlI:d OpelllillC;;::';V \'i£(!JS ofim,u::!npa.oe. ]It ij,"1.'Ili~.d b ~·Iin.ebJ) [lJOO'~n·whiehIJ~e \\:{)rlt~ of
p3lljiLic:le pl:lysic£; woul.d 1.IilI,m for dle(.:lIIdcs. Me~un'l.'h i~.~. f~\"'re:l>~~r~h~r's. at t':~.e ~

tm.'ge Hadro~. Collider U-[C).. T'!NOdecades later, time LHC' is abom [{I dl ase the disccveries never made by ll:i<iJll other rnaehine, tIJIU! tun [UlmI,l:OllS, Supereolll,dm::t= iug S!J,p'~r Co] lid ... (S.SC )1, Ol;;sig~G'd [,t) r
reflC::.~ em:ll'g'ies l hrree umes higher

\IJlll)Wllil~i;:~n~,.l'~!s.:i!'C'O~~:~J;;~ tnf1ld~ ,a sl'ii!i1~. but expeasive design c.i:m,us,e: {lileUnited S~Wl> ~ric;:d t'l.l go it !.1Ii.o~~ tlnd ~mlgillifl~ernati on'lrup'!rll'len only bd~lt,""d Iy. T1it,~tHe mlt:\;."Oeded f'Of I:t':ilISOM ~1~lcl'Il.~yoOllC'reLe-,ntci lho.'lel~.clOfli, iCQIIn.ldgive CERN dl(:!: :~;e iIiII1~e , ~
CGlll~'letllion .for 1Iw: ~i!e);Jr ig/llillic co~]li{'l!eu:,~he g IJlroptlsed 1[-:bl()lme:te:r~·I()ll~ strlli.glilil'~sbot II:IIU~i[ilia<ilil[)l131 Liillleru.· eoi lider (lJlC),

:Europe,m lab, CER.N, aear Geneva, Sw~tzerhHtd. mused of buUdi.!lI.,g ;lI wlf1I~~r macaine 0111 the c.be~;Il"They called it t:n,c:

bmlgcL was .. !-ley reits S'IlIOC(:'!\S iIll, buii Id il:bl" ·of lite' U;lC ..the died n...~com~~h~h;N;1~he tHe. 111'1 k~pil'!:s. \\!hh tlil~ tru~ly l~liU i,ll~ 19\f30 w~lell its lu~t!,g.e~:FI~.looll'!.:lclr~.·om cre,neJlI the lsb i~ 1954. each of,(,ERN's now b ~4.6 'bi~Wifm tOo InOI"i~ l:~nlil~] $lU bUWi,on;md 20 l1l~moor nat~ons, s~.PIUln·tsllli~eI~b illl !~rQ~ z ~)Qniolil till' ~Its gross oo.I:I~.estiiCproduct"n~e i!i the US. Congres~ ~dll.ed it

A~I agree tl~.il[ CERN'!;. reck-steady

sse

]1Il3ll~.~~{k~e' l

Why did th~ sse li:llil ~mdIlle: l He SUC~ ',,-eecli'! ~~J)l\Sicis.1S '1IIn poc!Ii~l~ID IliiL'i!IIIly Sti!U11iI.b!imag

~r~~ly creatas :s:uloil.iIY b~~JUs~lh~ ~'~]'!J.~beli ~ C(1Wllll,1l~ie5et:o£jll ize that th:~5 islm,'l .oomething r 5

NEWSFOCIJS
open for
glieen ~igbi~.~ fhl lowu ng yellll': ~l.e Even I.'el:ycli)l.~ as much as lbey could,

II

OIRN .. \iij~ich g~w~the tHe (h~

Hl

srand i[~ hear c!(Xl'lductor. g ]b be $'llll'e. the lHC has. hi~ so.me pOlho~e..,
:!i.I.o.n,g w'!Iy. In .]OO:~"areview shewed that the W.L-; 1fU1i1~iifl£. behind !i.Cnedllk: and 20% O\.'ef Itmdget,. forc.~llg Illiu: ~ab ~Qscale baek other projoclS .md. relfiru1lrnt.'t! the LHe (Sd(Jri'v.?~

more correctly, theother end (l,nhe @CCG:I~f;ltor-,U the end of the umnel .. Soon workers \~.'iil put dtrwl1i their wrenches nm:l welding
torches . .j:~ndfes.ealrl:he~$ will begin Hi brri['lg rhe [DHll!::hUI~C to I~fc. "We .~['~ very exeltcd,"

researehershad

j)'l!:-hthe liurits

Ofl~C.~~

the pimject

no~og~l.Ev~m;!;~'y:S;~ ··SOCI.ct}'IS vdUing to pOly acert a i~1 mnOullit and no more. and to lli113lkclht:· LHC p~~siMe .: ~'VC h~d 10 he innoVUiVf;;:" R~$!(;;.afCht;r!' l~~we de~igi:l~dth~ strongest ~,n.agnel:S far form.l acoeEe ....It!OJ.·. 'by crammed lwo uccelerntors into nw.C" )lC~of magnets. used wires of h[gll,u:lnl)eratllre' superconductor to dlislribl!~epower, and piom:ered.a type o,rmdiatioru·hard dectroUll.~ lcs ~(lt·[heir deiecrors, "f~~e:yeven ehill rhc hqllil~d he]i.1J11l that coels the magnerstc an ('L'Ii.~ rll!~ ~jcl L9 i;a~ in to make it [! fro(;~ frl lv
flowing superfluid, which is also an our-

En:g:e~ell S.'Lys .. ·'wld ... bil worried

because

28 J~U1C .2:002.p, 2317 ,.1~12004. p:rob~cm~ emt'Ii.'li_g1o"\J with tl~,e"!)'{J:'~~Ilk'lii:iit!! th~t U'~l!!~rocl¥T the: Iigni.d hel ~mn ~o the IL~.13gne:~s. WO:I.t::ern had t{l rip out, t~pi.ir, iIItdreiiilst.d I 3: ki~~~m~ter.:; f o Hue. cre<lllug an enonnous b:ldmp ef'the 1111l<Lgnels t'lie:i(1i been ill.H;"dl~f1gmi snon as they arrived" .. ~U ~m'l£,i~:led ::!IOO'I:.se Ibr 50 [l1l3gl'l!!lS.
a 11(1 ilU~'II;:£nd .llrIl!Id l.il'k1 roam to
EV<IL1£, \MllO S("'Jlte~-ed

now w~ h{iV~ to d:C~ivcr." Se<e :yiijlJlin Swi;n~dali!dl

A ~re'<1dy.Ilhysi!c]s:~sare flo ek ing 10 CERN in mHjdDl!'~i()~L Some 75·mJ ~J·f ! ~.I dill~f~nl nati.o:l1<lI:inies have registered to' work on the
t:ii'U4;:, .!~ the

1m I (KJ(l."'lliays

tiWl.H.rM overlhe 1Itb. N(I\v. abaur 2: Y(,f!rrsbehind du~ir Origh1!1l1 sehedul e. researchers see tliu,:: gIH-or. h

exper] merit'S. suelras CD P and DO,~t Ft':D.ln~II~h, ··W~.CIl ~ was hire.:!. lstartedre 'Il\1'O':kon

LHC hn:i::st;.liI:!1i1 ~MW'L,r ffl:un u~~~~r

DO."
Stoll}!

5<l1YS Ad<LL~.1L ¥urke\vicz.

a pesrdcc at H~~:ml rl.iWflOity ill N~\.vYork. "But ] U

~ 'c,o$mop?J~h<'!lllil. m~!rIIYo:.egro!JfuJ) hO!;is sdentiSIS .... QI111 <:hlfle:r'li!.rll nil!(.onah~n~. yuu vmc up a<rdown ev.cl:)' ye<lr," ~mys.CtIEI)U'I~

W3:1ll N)

buUd the

nU'lehil'u,~

eless

to ~1\if:ilUe •

1Uiid "the U.S. ']lid J~lIp<'lln ruld reue!' ~.oo!k ~LP ,uuJ ~I\i,lImb~.y~.ean1i.Irem CERN'S: hilstQry ilLn(l

esperieaee,"

"~I.\yt>

N Obll ll)~J.; 'Ic)~'tllib e Jf!~~(l~-

p~~.C[RN1> un!l\I'W!I'~'r'i:lli£l 1j:IWI.dl~leeuld .!lC'IU.aUybe ... liilabihty iii the 'col1lp!=litiolil fb:r Ihe I.LC. A.lthollgh eEItNsl\nuii'IlI!l_s;I~"':l d~p l'flp~"d h:;t;1bly; iJt ",I!"IJ d~I~~~ 't d~mb quIckly. ;l!::;i. Dm';feI!!~.I]lcoIJmtRlsr. ~m~ .S. gl..}\1'e!'m~l~nntcan U 1'3:p.id~y ronil~lp ~IP fll.uul'ing fo:rp;rojecl:S .. ''Time' u.s, syg".icn~li$ Illflo:!'Cd~mllmn.c~ml eru1!re~c[ to tiruilJig:s more 'Iu~ck:~y:' say~ Pier Oddol1~, diu;"CrorufFI3f!ui N!li:ia]lal Acc~e!M1;j):r Ul~ I'<lu:nry in B<I[<lvl<I,III·inoi".n,n.may be impoF~ tarn FDrlh~ iLC, whiehwill prl1b~!blycost b~lli'i(1en~ III bi~~~oi'i $ U bi IIian, with the {!rmd. host picking lip' tJ.iI~rLh.e tab, r~!ly~ic]~:i!~nNo:nhrAi~l~riC<1<IrW Asill ~~::5g I1Q~e dlal,a~thou~h CEIRN is an intcn~1 iO'UD:lI:~ D'lrtmmlto.ry .. il dues not embrace <1.11 loul'lu'ies. C c<!l,Lmlly. CERNs. 20 IrTIlrel1mher mldom. ,cnjoy <I.
speci(lru statns O(U'lIil.,P'<lffi"cd m"giliJest" natlens such I!IJS §<lP,IU .nnd t~:lf: rli~edStl:'Ules, .:.:m.d Jn5UlI)' U ~Il!lt~lkl!' \'1~l~!h~"fI1iD_.e I~ty ~H\!cl~reisn~ibl~ cmI.Oiu.gh 1.0<lC;o!!}JnI~~e i!I.IDru~ygIJob31j)mjecl. l'ill')ling 1'I':m)!lIx;: :1l:J.>,)!" ClE RN \\'.1Ube pe,yi mig ,offlhe 1..]1(' unti,l 20 I I. <MOOOl.ill"ter dl<l:t. IEoe I<lb p'~~~~!O u"gm<!d~' tll!£'lUiI~hht~It1I'1~dlt~ l'O ~1.I~n
3J~.

ese ."l!::oc.lc~lil'orl.".lOOl~~oryK]1K. in Tsukli1lba , [l]rt"i::(or Genl':1IiaWRoben Aym:!!r: lhe-ljIHiHlgemen I :s;C'IS thelab bml~,ct S; yeail'$ ln advance St HLTo:g.e 3dd~;. "everyone has 01 long\'I/uy ie ~,~d '(,lwn O!I~IOi,:Vili: offi~i~ls tulOarwrn:v ~g;lil1!;:n go [0 learn how to :1m1l3kellii.e n.c a. sueeessliid .g~Qi~1 p.rojecloofore jlli1.niilip~ng (Weli each fUtJi.lllreim:O:UlI,1e,,,s~ihey d~1 iIDl.2002 wlfl.eJ] they . other U) see 'ido is tiC) hO~I. it" ful.! n:d Ihtltt t!mtl l HC\ll'!n ru fll:'!i~:g, 20% over bw~dg,eL In
oorrJ~r,ilI!>L the

2'0 l]1Iemb~'1" :~·.uiolls UU1.Sl

tQgl."!il!:

on tiny

VUlnei;iljbl~. I~ ~he Uni~,~d St1!~I¢S.Congress fmlds. [he ~ii~~O:I1i.aJ1 .Y'eaf b}' year., '1,,'hic:~, .~;11OO 1lle!'21trM;:;13.'b b!i1dgcls flUCiILHI~e :[!Iud projects
:;nlQ~ IS the sse nice .~.h,e<ll'l: IqJIl:<1Itedly. .• "VI~liIl 'bl,i Ud Ihc LH~('.c:[m.:N~ko b~TI,Ci~g

sse '1\t3S far

m.ore

I1l'ed ~fr,(:JtT! mlJvh~g com illtn,{tln$ly l'rom one C(l:~I~ ider to l~~e l!iI.c:.t. ("ERN researchers bega:UlI ... desig!'!.i~ 'fik L HC even as they buJiJlrnD,oth,tw
maehine, the Large E~eelm.~l-li\Isilr01il ol IK1!ef' C (L I3P), l!li.11~t;ilIIlIJ!1 fmn ~ 9:8.9 ~{l :2:000,. By 'l!!lsi]1Jg: LEP'~ u:l!I:melllo~l('l\lse LHC' and! ,exiM.i'lrng ~'e
~~~~e~lflrs ~or~d it. CERN f~lnchds saved IliIlinn s of SiIW iss: frmilcs .ulId b lili ~l~he LI~C

\\ridlt~~lt~n if!Coc'8~ i~llw bib'!>. ~~u~~t. COIlt:iml~ll:y '~<'IS 3100 he'lred Ll:te~3b acc mc l<ll,~luedper,';OmlL(~:1I. "TIl'e [W,G.S l iIII ['l orHmD. thi I~ ru in,~tii]g. a p.cOj~~~ike this. \\iOmi5 the I
rcOtph~workiIDlg u~.dt'ir )lDlU;" S:'I)'~. CERN'S Lyndon EVo'Ins.t~h;o leoos L~C conslru>cti.oril.. "Onlli: ofollr <Id'llal~wg~jO [il:mI. q~l!alhy (lfl~e

~(Ye[ y IprobablliVI ICEA~~ha~s ...

an advanltagle over any Dther pllao8 to host an 8ven-m ore-ill te:rnatilClna I effort.than"fjhe LHG.
-Robert Aymar. CERN

m;'limai~i!lg
pl.'OjeGt~

u::~ wbi.ch 'W3lilm.'t so Im:l~ ago.'"

bdO:I1e
lh~

$Wm

is~l~ru \l;~ ~!U~l~ olT OloothGf

lHe is Ull
311'!d n!Mi rig.

E~,'~!l

TIii:;;n 11~ii1."[I·~ s[npped e:U'~Jyjodteyiililg.. b.O\V~

1l'i.{}llN:ollisiollS. So ('ERN \:vilm ~,l<l\'e its.hands !l!.1i! umn midd~(lOn~lefl!~~'>;t d.~~~ldQ:,n [ {,'(J~tur'L'!t. dle Unltlo:d S~ruili:l) \.\'~~1h<!iv~ ~Q !;:'UIHd,· en; I~'orpartic:~:cp~.lYs£is in o:peu.tion f1l):Ii:J 2fUJ9, bpmfl. :~~lti:iirrti:lill'r.'coJillideff thm Ct im&l}' iI l~pgmdc and 'wUi soon be fmishull~g iI. large pl'Olll:lib. ;lcoe]e,['.uo:r conrrupleJ£. "j] f 'We pl.oJIIil ~o blJ~~!dI~lliIe i'~.~]m.e 2.010s. CERN is ill ,gmld ltC

*~

fl!~~ys:ieis'~s HI:lo1l1lndthe world ar~ I,!!)oldllg lm.!t.;)lIdlh.e ~.e."'.1bi~ 'lcc~l.e[.~tOIf'!rId <Ire .~irnS lO drl1\v :SOi'ne :les5'ol'ls. fronl I:he ,co:titr3:S~$fl~ f:fl~~s (J!ft~1~ LHe a:!Jld th~ r:~.f;,1" S~ly z they 'tv.i~:]]l.eed the .II..C~9 study iml. clemi I the I

{,l\I'lo\:r. EunllfH;;~nll P'l1i[ll to CERN's Sl!~C~Sg COl!~~did:u~"" :saY'sKEK'$ Min~u~ld Nn~;lkL "1iilLh the U :~c its. exp~icillj)l' i:~:~ema;~a~~~ "Ho~.~1e'l'c:'r.f we ~~i.sh10 ~mn constnt'C'D:ion and i

sse.

.~ !l~W

'S

!\i 2:~ F'ebrmJH'Y

D,~itdc~r.;51.~1;: LHe ,~h(Juldi spot l~),"lC'I'I(;q. 20n3, p. ~Il 71 ). Reseml:;ltIie-rs iU1l.

l'h.~Ullii.:~~dSH!tC:S • .I3,~')<an. and IEulio'pc 3.U

ch.ilraet~r :as big plus.s~s..··Vel'yproblUbl)t. CiER:N has <II:} adv<unlta.._g.eover a~ other plRt'e to h>o.:;t!I.vn. e\'I:!l])--IUQJ.'l.,"- i~t~rnmii(l!!Jll!L1 dTbrlt~11~rI <IIUieildly exi:its," says CBRN'S Aymar. B~lt ~;apilJm.ese :fII1lJ! A~l~Co.m plCWfiid SIS say du:y hay'll: mlV'.tIltages of thdT O\",~a,.for ex ...~~~

SO(ln .. fI!.1: Ul~fll"$l [dIl:Iys:i<:sre,!: ..t!I~scllm~om of th~ U Ie around .lO 10. then the US. ~ntl J1<1pan ~ru lb,'!! mruly ~!Ilb;(ic ~~n:dndat~" or CiJl:j~.\~\~1ether ~l~eILe gets :b1:.lli m a~~ r It de.~'[Ids 'Oll whelhcr th~ [.I-I'e disoov~rn ;;111),lhing\'\'ca.1ll1])' of f'i:iirlhc;U' ::o1udy.. =A.('.

1655

R(':$.;>;n,rche:l's:mU$T go

&~O\1!.1

b~'C~~H'£ lilt;

ILH(: is the fill:S,l col Ii.oor powcl"fnl elH:mg:~,l to <II1::$1I.·oy itself, EI.I.ch of tlre tlle's he!l .. s m

B.I!~~ii!ii Ilh, ~!li:j'm~l:nt~ CERN )al,red, bi~lianl5

by hliHdlrng~ 'lnf me:

inl ~I'lJulfH1@l drfU(!dfii)f an earlier acl~eiL~r,at:Qr.

P<'Icb 11 ~~OiL.ggerirlg362 m-e.!:!..tiou les of ~nergy. l~~.~· (."qluh,l!ll,c~lltof 90 k i I,ogrrmm:,orTN Tand en ough 1,0.nilleIt j 00 k il o~mrlll'!' of copp er, Should (1.1(: niil~chi~le.!IrCeid~:um~JI:r steer aJ l:H:!i:1!iIlin(l it~ ~)W~l i~mu!(d~, ~jf't)mils ~Oi..dd i ~h~ dlrin O!I.liiIo]e~eUilJ> Iflneter:s ~m~g,potel'll i<'lU}' [) h];ki rig the lH C o~riIf >ll:~ion I"vr mvn:th:;" ( 1"0 preve.lJJt such a ca~,am:~ly, accelerator phY!'i~~,i:!Jt::; have gone~iO eJur!:lm~~ to pnl'w'Cl Im,e'LH'C from i!s.e~.f Nior-e than 4{100 ::mpc:'Jrfast beam-Joss monltorn \1111 1[1ense protons s spr:l.y~lI1Ig Olll tlin,ebeam.Tndependently, '~!l!lH;:urrem nlorli~or~ \!;'ill h~fer losses by

or

m,eOll,SlIrUrlg t:~.e amount

of

circlI]:ui.lllg

dcwl:~<Irsubsystem, bee .. use you lose toaeh if~[ht\ Iilru'fi;om:,it~.,; he plat'erordi~Q'I,i1;t)C r w ~d\iDhe enJi,r·~detector," he !-wy~. Thm:ewho C,II1IiJ(llmol,'C: ~{ji,E!1Iu-\')peface Inll~u:;t CER.N feels a bit li'ke ,I reso:I,tlbir the Chfl:~i.~11~~ ofr,;,ecp.ing contm;:l with experd1~ nerdy set, MOlky b~tildil1i~" (b(',:sde ~ihm:g streets named for Ei,:Il1i,sleilll., :feymn']l1I.'Jn~! iments dtommnd:s of[ldlorn.eiers <Ir'I,\'<liY. That's other rf,Q]lO~1s ph)'sl.dsts.. H()n: ~md there ~ie~ a big problem forA[n~rican D'~.Y.~~c~s~s. who make up 20% of the ATLAS team and equipmeut :aw<d'~:i .u>se~.nb~.y,,In the ng even ing~. friends meet iU:llhe c'lfe~ed3 to chat 30% ohliLe eMS team. Teaddress it physi-

~old{hem 1 ''W9)rned In~\\'iw~,o. ATlAS t

because

ou~ () r th,e

charge. And bC'IIn-I~josiliml monitors ~H3r1Se wilJ;:nchebe'JffiS sln~>' fwm. l~~.eiD' proper course, These S.y::iICHl:!:l esn trigger m;l.g~LM: lh'~1 ~s can safely k~ck rh e be;! ms
ft!Hu;:hilli!,~

wm

in

~lhie r,e\.\' hundred

mi~n.)M;:cofl~;h. ~ i tf!k~s l~1nH.ke tw'U ur

l:hre\O

rcvo]m.ions. Iess Ii me than .~.~ rakes a 'wobbly beam to veer offcearseeudrely, "Foll The IL~.~I'C. ",'C'v,e tried From the start to cover ~.I:li

over
~I,l<lrl

<t

beer .. U D,nl~~h Curb,berrg

S'iy~

{D]"

Czech

cisrs Me fdyblg hl part on '1mi,gEu,oC<lP<lcil}!

DOl!Iillm ring ne[\'II'Ork C!l~[edl'lr'n.C Gridl~o trans~Irni.ld~~wW key ~.,ubs11i {~l11erl'lOUliirrie:ii. These ~ ":Jllalys~ ~mppan ,ce'l:n:efS" wi I~ s serve asgarhI;~ri.!l,g places U~albri!lS lh~ Ul(' a b~lt~~, dO!l:~r to home, says Miehael Tuts ofC'olumbi'l
I

I~e differene :po$lsibJ~ fflihlre scenarios ""'0

E RN'~ RGdi ger St::111l1l dt. E~re:rm if noth~ll.lggoc;s\\':mn,!;l. lphy~icists t\r:tIlust ll;'l!k~ I:!r~~ordi stepsju ~t to !mktl i.;)'; mH)' IklC~k :~ 1.:1protons i~,w each beam ,<I~,1(i iI] usl 0 one I nomi] Iionth of ~hem flew irI to ~. magnet, thc:::~ would ~'n.el~l enough to lc:mporalvit installed hundreds or called co]limawn. !~.n l,,,'i]~l;~!d~ i~twiM'II,~w.~wiT~fdpani~1~~ eles, "OUll of ('!Ivcry 1000 particles [ile~uJed Itn1j,l~!rdl~le t:o.M.inl'ilOl.'i]. m:n mnre l.h'UlQne ~l1011ld ,esc.J]"e 1.0 re'<Ich rhe n~I,lgnels dQ.\tnresearchershave
::Idjr.mable eonstrieriens
~Lr~al1l," !l;f~ys CERN's Ih!H we don'~: have ,HI accidem,"

Budweiser,

"'There is I!I tlli Ifcrel1l <Ill1lo.~pltere

s<I)ls

,David Si.I\'el'1ul.yr. ;1 S,\,l;'dc ("rom Oek Ridg.e NlfIli.oIJ~~ l~lmr.u.o.ry !Iii! T;;;;lllll.~~"~-"Ifyou ~o Iii) I~~dl~~re. people"
<liretllU::il1lg ;!l'WJUll.'We·wl!!xcited abCim'l~Ji~: a~

in the U.S ..:'

'\\~r~ goin,_g '!.O hliiLt~ Ihm: ]lll([W U S.. pe(j.j:l'I~ U~l~vt;r$ily. \:\"h~~ m~nlage!l.lhr; ATLAS ~llIik: 'IOOIJ~. olll.dg,el s," research pnrognun. "TI\ie:(,re places where BlUt iftlle IU-I.C is; c.~~]1.gillg,~hem!a() of y~1\1can go ~.M.d g~~the water-cooler COHv(::rp'_m:~c~:ephy:-J~:c:;. alsomarks <I~e'lp i~.lIhe sation," he :says. it field ~~evol tlli.ol~lO\wrd e'V(lr.bigge.1" I)mj ~U;, ~11fact, evenas CE RN draw 5,penp ~e:. he T Since the ~. :60'5. exp,e:ri11.1:ellt:J I co,~~.abor<ilti:()n3Grid shnuld heJp eii!,teud~~.e re<.lc.~. f pani9 o hllve !ilmwn too.i~.d.ml~ dozens. d1,enhUIb~ cle physics across the M~ob~. says 1:I!luvey dre~ls. ml!..l 11,{J\.\.'llm{Ju~~~d~ nf ~ie;nli:1!lS. That Ne:\'lim'!l!1iloft1lll; Californla I!l'as(iMA;; (lfJ~ch~[)I ogyii I1IIPaS!Id.el13 .wl],[l eha irsthe boa nl explosree expansian has led 'Some particle p~:ly~iciM~ to seek more ~migll'~W ~Ill\'im~s~rll that eversees lilt' US::. C,",IS [etH1l .: 'There othe.r nelds: lSdelll:.'e. 5 Ja:~l1Jary. p. §,6), Bm i~. :lI,ILCCOIJnHiies, t:h~t \.V{~reu't inthe field hl.~1 doe~I'I'l" filzt: those whoh.we chesenre \,!,io:~k ser iO'UIS Wt.y. i.1lHI 1'10\"" I heyare there," lie outthe LllC . "l'd :stil~I have <I :sense Silll:isfac~ says .. FOI." ,eximl p.le. phys~,dsts in. PlJ.k iSliMl. ~im~ m!li~t~f \\tl@t \1,i3s, d~~lm'~r~d~f~dl~p'w ludi;!!., ~I!~dBrmdl \vill h!l'!i'~ ,1!~(,:Iil~S h.l th~ I~O b:ig ~ role :1 Imd in it," YlLnk!.!"\,vi,cl sa.ys....l·d LI~IC da!<II ill tlln:ur [Emu: 'CQUlllm.r::s. kril~l·\1l'lhOllll (,If)!1trib~l~ed;'' Ne".erl~.ndess. 1.·e!'Se"<1 rche:r-s rc'(;C!!gn:u~e du~ Wa kii !iligtffil;e gillint The fum tOl:rerlll oflembytes I1my be ill \\~.rile iu (': 11<1'111 CI~~~ ,of ri silmg :fh:nn ::H1Ch a crowd to 3 to p(R;~donoflc:iilde:liSltup. "h i:s a vcry ,compe'~~ cm;lliUlIg. howeve.r. R~Sleali"Chen;pla:rm seaJld prou.m::; .]r(II)J[1d the rring hI Novetl(lI_'M;:!" ,nld iti'V@ ~nv~n)'l'Dmellltt:'~~y:s Rosy Nikot~ido~ begilll~ki~lg d~m liI.cxl~pring. EVC:I1l.1iJ.cnthey of CEA Sad.<lY. ,h<lllce, ,,\<110work"", O·i]. !:I.la[J1l<J1f~be ATLAS. "Eaeh iJ~y.. j.'Ol,i!. hfU\fC 10 prove thfll \';.,m stan .~l~O';.'1energy-Ie.ss TeValmllls--3l'1d \l;I~lhImv bc~lm i~~te.liis~ly. orr ytlU ~!re lh~ b~st Oind t~,n y·ou. d'~:>l::rvl.! >,m~r ·'Iumil'no..'>ily:' ''If\\'I;;: (<;In g~l up to !l ~e!l'~hof chnnce:" Nekl.arioS1 [llh~IH~kO!l.an ATLAS desilgn [~j'lllm'~lu::~l()sity ener~y] ...f1e.r the .11 111111 member fmm the M.11;\l p~,"r'iC'" [i'i~lih!lt:: far P~ysics in Munich, Gerrn.nn,y. says yO~Hlg firll:l y~;j.r.] think U~OlllwOI:dd b.:; imlir;ICl~lo~t~~" phys,i~ re:;eOln::her .. mmsl Ih~llrk ~wll.,e~~C'~.ny to says; 1Vii.chad L;;)mo.n,L "'~I,~~u::oe~e'.mlm .. " avoid. for c~... . .,mp]e. bei 11£ pugeonholed" ":fOi.~ c'i M ;Ai c: ERN. "Amllthillik i~,e ~7'i.pef~m tl t~f'S sure. ~f'Oli~'!fiIU~t ~nt stick T!)(, ~uH~h toa po:~r- w(l~~.ld be quite h<~)P)1w.ith th'1!t:·

u.s.:~

Ihe LHC run, The co:lliluder iili> designed '10

i:IIy [k.lIU ~·~:s ::iupeixonductivi.ty",lg,,~gg'er~,:~,£ "I beam durrsp, To avu ld SUC~1. "'qnC[J,ehC1!s,"

or

R!I~i~':~~' A~,:;m~!UL

"Wi th(m~: ~his :system. 1111 e L HC cOIn I1Q~ m:lm.,. 11~fl'S~f~rl;lj~sl dw: t~ch!~k'RI oCh~I]I~gc,~"
When the dnu:lfina'Uy m"r~, t'o I.HJVi.l'Crup ..es Ihe mlac·hh'a(1;. the m3i,~ ,challenge be ml:'lm,gi I]g Ihe peop~e. L~nifLont :i<<I)'S. "Tlu:re N going to be :'I IOl of peop~e st.'illlding un the control WOOl. m<1ybe IlOt n\'idd~imt~ the knobs bm ~oO:ld.~.gm!~r ycmr i>h<OlLldcr.~nld

w:m

I~i!l!t'~U

it t>IU:flddb~~~:hif;iSf.S ~;>;d~iny,~.il
"Bul whm you re<lUy

gC~!l.:'· L~'LT:lI.l!o:un ays. s

,\I'd!'!! iS f~)lUi~uys

~iui~,;ooh~l!1id dO~.!;ldmJ'fS d qlu:iet~.y n.gLu1ng OUI. how 'rG ma~e Url\v{)J,~k." "fhOJ;i; fo'l,l r wi I~. a'iJ\,: ta cope wi[~ tile CR1Wd. ~] Who could Mmne anyon~ for \'"lRIiII~~!ng hl'tJ.e l~le]'G \Vhli,lllihe tHe ushers i~l ne\V ,err~ if! ~li

~rli'de phY1riics'!
~,AIDIRIANllnIO'

e;

NEWSFOCIJS
Theparricu I arr ch~~I~~l.g~ ~sTO give IVlI)SS to p<ll"lides C'<1I1ed t~~eW and. Z bO:HHlS,.
\\']nklm i;oMilvey lhe'n;:~k nuejear Foree.

II

Ma!li1ry' r the UiC~1[l rColllglh 1I1!lon~,y til e one paln:i c:I~e~hey\r~,soug hHor deead es, fea Some \!illolWJIl~ r,~~~I'l,er n0~~,ing neW!ll!t ,ai[ll, see Suppoo:se you i1r'~,'l r~rlh;:!ephy$ici;.;tA. score of :UIli!l~OIl8 :h.u given y(l'l~several bi!ho]IJ1 Swiss fmnc.-; to build a nHlch~ne lh,<]t "\,'.ill P rn '1\H~ ihii:: a,ru, gl n S (I I~ m~ss,t h~1~.neffable i .solllilclhlil1g rbat keeps au objec[ :irl s'tc:ady metienualess shoved by <I 1ib:l'iCe.¥oll:f IJilTOpo~cl exp i@!laHon of mass r~qtD ires ~ tt!JG\',V pauiclle'. uyplic3,Hy dubbedthe ~FI,~ggs buson, rhat yeur maehsue aimsto ~~rY. \V hen. i1.flIe.r 2, deeadese fpl'ep.~.mli 0 1:11. Y{IU gC'l rc~dy to s;~,v[tl:::b yom rig. you 'W01.JI.1I 011 ~:.eilr~:M:dilig utore lh'lIli the possibili ly [~li1'~ )'m,J\'!Ie'~ ~,vrl]llg ~ml th~ par~,i:.ledoe::;~,l'l exist, rughr"! Net exactly, M,I~Y pawti~:~c ph)lsi;Ci.s'ts ~~y the] r gre~u:ej~ l~fI:ri:5 dlm [he if gr-~nd 111;;:\\/
machill1i,e!-the Large Uadl'o:lil Col" I:kJer ~U~IC~ under C'OJ.l:;'lfutlio~ .Il the European particle physics laboraWf)'.

~~ilh~."~;~il:mind,·,bogglingly Mgh ~rq~q;y ;i\t wh j,(h, gra'l,'il}f pulls as, hard as the other lurces oL~mllllff;:" The Higg5 ,done could I.':s8em.iaUy mark iI. di$srl:li's,iY.~ng C1:1m,d abe ·to ag.es·,k'mg quesr i~~llQ essence of:mril,"~h:~: dH::: If. on the other hand. t!rut:: :lHf: sees no

Ace o.rdi.~lg to I~U::: 1!l~]nd!J.rd mode l, the weak force that causes a typ,e ofr!!ldio~c:~h,'e decOl)" and the el!ec~rom'lg:~.en ie :Wolfce Ih ,n (H1W'C[:i; Ughlll:illg mild ~~pH1P (I~]ni~pHl.~'n; !I:re l\'VO f.u:el~ of the same .si,UiI,~I,e thin~. The two forces ~re:lll't precisely IL:~,h;:'I·ch'll1llge~ able: EI~cll!l;)m~.gnloltk:f~lrc'e;:; cun stretch
between thesters. v.~~iI!re<ls weak force the

d nesn 'I: even reach acruss ~he uromic IlUd&~1,S,.T~;HUrange d~rfc'rence arlses h eeauscp ho tOIHL l~'l q uanm m pa rlie Ies e that make lip an electromagnetic l'i,el,d.

new

pM~Ech;f>.lllt ~It then

tl~ev~ry r~de~ (If

h ave n.f~m ~$S. ] n con H ast, UnG'p.f! rt icl es

l~al

mn mec hanie S and even Ei nstei nJ 's special l.h~ory '01'rol~lh'ic.y mtu,1 be 'WfoUg .: oH: woLIIJd :1I1i~el!J]1 tbat everyth in,g 'V,,'e th(mgl'u we knewabout everyt.hing fal:lls {Ipalrr;' says
Gl1I311ilt

make ~.ptile' Wf'i:lk force fit':~d.lhl.c W<lnld t Z besons. are ~h,OlU &6 and dmes as massi vel.'!' [he proton ..

en

Unfn.rHLnaw.ly,

the perssiekety

stan-

dard model fans ;lpm~, if rheorists ti~~!1i!p.!y i:I.s;s~gn ~~~~t~S;e:fl:~O tbc w. Z •. and other part icle s So the masses mllst snusehow ari se !'I'om intera ctl '!)I'Q,s (l f the odl.erw ise m assle ss pa:1iticles the mse lves, I.,~.flme 1:96JO~.
fer~r l~lig,gs ..atheerist mEdioblill'~~

('ERR

near Geneva, Switzer-

U~iversi~y in the UK.,I\eillJ[zcd

i f th~ LHe dfl~sn '~U"I,;;W~j] a p~t;::~ h:O~l of new particles ~,:ml additien to the Higgs,. Itl~~ny ~hey wCJlln!l,d rather it :say see nothillg new at ..IL
That uillay seemperverse,

I.alrn.d:'_\!,'iII spol~heH i~~soo!iOn,md notl~i!lS elsa, II"~n.p3niclcrhy~i~-~ could {llimid to halt. 'they say, In f11CL

~mp,ty $,p~e~!ll.ighL he wjth O!l fie ~d.<i b~I,1 <I n elect rie fie kll. ~la l ike

rm"d

~~~''HH

but Pt~t yourself'again ill the shoes ofa I)m'ti,· etel:i11ysic ist, [n the .~ 960::; and .~ '970s.

researchers hammered

OU(

m t~'(;;Oi)j Ha rvey Ne \,\'IlIiI!IIl.

il n

pru!bI<.bi~kL~:-;,amI n~h~f noosel] s,e. So ll~e coHuder hm:i ~o cough up ~:(JlrU!l!ii11M new" fl:5(!f1rchiern :lay,1 Fit s:pils, om mllylheiiigg:i., hOhtever.lilJe Ile:w gi()~.de[liI.ge o I"d isco¥c!1'" cOI,.I]dclind ~tSSOUll as it b~gi m;_ Inhe Io:rael:li.g£s hOl:sj!lJstlhe righlll.]l<lS.<; <libom 'II90 limes tile 111:I,;I!OS f <I p:roto:Ill.-i~. 0 \il.1\ou~d [i~ Ihe $t<:ln!(l\;).!t~:m(~dd'~ Iml~~ ~tP t'mls. <Ii nidi ]e<l\'e pllJ1sici Sl5> eve~, mo.re th,ol1ough I.y ~It)']il~ied~l1'Nl bef(li'f::. ~!y:l. JOI'h~dlai1! Ell i$., <~ !heo.rist ilIl CERN."Th~::; 'Il\,'o~dd'be tht;: real .~ five-sHu d~S!'ls,ler." Iw salIs. "b~C;.'lU:lH;: 'Ihal .~ ~~~ou,ld ImI.lle<lUthere \.rouldn'! IL'fo:ed to be any :~ f),e\'\ip]tysic:> a]l the wa.)' up [0 ~h~ PI<l~d

called the standard model that. ill spite oHeavins; uur gravity and Kufferi!fl~ fll"nm other shorrcumlngs, has csplaiued ~\I~l"yth jug scenl n coil i,(l!cr .;:'X~rime:l'il~ CV~:~ since and ,~ef\ phy.si eistsw ith few cl ues lo a d~p~r [ll~l;]!iY, At~:~~ 1,;!'il,;lrgi~J; ~h~ tHe will reacilD. I:he standard :U:I.l,odd goes hayw:~Iie.

ex pe r imerue r

OIl l he

Cal i. ForuI.a lust ltute .of TcdulO~Qg)i'iu r'fUS.a1dl~lU"i, Thflt \Vi,lul'd ~hriU IiiIUUlj1 b~[ i:; =>.0

ml~,i.kely ~'l'aat ~ i would be "esseruiallyimpos:.;ib]~" ftJof ~h!'\:U-[C to s....~... ul;l,t~i~~.!!J'li'~\V.

NC'I'!'m'IDll, says. O~Ihl.ers .. ~.

spinin,g

nu~' eg.-.dv~ n

PI.l)'~id$t's ~H!\'I;:nn -~imila:!r ,gL!'~fmMI/;4;llh.n Z slllgg.~s~ Ih~~.l Hi~ ,lln1W1'>120U !.im!.;;;<; ,"~ hefty <I:j ~he pll(li)O~.l Imlkz:;i Il! Ihat dow:!.. COIIl~he tHe ",vi.111 re\fl!il~n<l'I onl!y dle Higgs but
also e'x.ot~!C new paJrltd,cs
d~2IIl '\1!''(mld plJ'.hn 10. ne'w ph}'sic,~ :ntd ,opel'll R l.1e'\'Ve:.... of dI,is'C(wcry" S.O Uu;LHC is~] gmnbh:. afld IMi1rY ar~ pu~.iing r{m~l~e more exciting long! slnr~.,

eeuld dr~!~~t~partid('~Il'~ gh~ H~~~,~ inertia .. the essence '0 f" mass. The HddwguJd c flln~]:ii~efa n.e;W P;:)!f[lde. the Higgs boson. h~fkil1!g"virl:UllUy'" 11'1he vacuum, t Nature appears to (oll(n ... 'lh:~s scheme. Us,h~~h .. tfu!om'isls prcdi!C~ed the masses or dn~\iV mull ,2:. And .n C ER.N inl 9'!jS,1. thetwo parucles w~,ighil,;dln juseasexpecredjn t.'oUisi ens e'I.U: rgetic cllml~~ lOpoop!~~],em:~, m,Hof Ullll VIle ~umt NO\\', mounds of datill. point to tile Higgs. For example, til" lifetime .1inJ ~lh(t.1" IPrul~'~~vlies ofl~,e' Z depend on, the ,ckmd of vi rtua I: p~Jriid~$ fli~ii!1Jg art)ul~J i~ n i~s s\\1'tm'!11Ji!:ig l~kl,C: <I rouen I1mn S<lIudwk:i:I. ,r~cise :S~JU,diLes the of

Q'l!IClck IL~lk@, H1i.g:!lIs a

Ell$i~~1 lh~ !m.Hl~~ famuuspanid~ ~Ol ~t llliscovered. Ih,e Higgs has, even beel1l, C!WD'Wllled [h~ '~G[idp~~nkl.e;" Me N~lbet~nuru~~l~. by hl irealli,ry. howev.f:I,·, it i1s merel)f a:m in.d l].oc :sohldOll ID atlabs;trll:S~ proDkm ]111, the stllndard lI,uodel: how 'to guve panide$, IDll::;S,

IlZlrillg the [lMliSSCS onhe\¥ iII.lld ill. p!.Irl.ic]c c:!Ued the ~lP qlliJ.ark sho\,\i,S. ot ::;irl1lil<1f Ih~:~,£. s,lIYS Gordm~, Kmmc. ~I.ll;u::ori$t ;al lh~ lJ I*~el'~ sily ()if M.~t:l~i£alll.Ann Arbor~ "These <Ire lWO c:: oml"lete.~)' inde'pel:l,de l1I,l p.ieces of evi de'n,ce l~~~tdl~ro is stl1~wthi~~ (bill \\."alks .lIldlall-:s and qua.d:s ~:i~e 3 UIUggs," Ka:~le !>!Iyil. ~'The ~...:i~:~t:!ncenhhe Hi.ggs;i~llh:e LHC r~ri,gi;; is eSSlel.lti<I~~,Y t::erialn:' Discov~rhlig dw IJ~i§gswO~jld Cfl~npl~[.;; the tltand;ud InQd,e~, B ut ~':i;,IlI,d ,(}ul.~'the i~l,g

Ii ~ggswmdd gh'e ph}~silCi:Sl~ .~it[ Ie u) go on

ill

165'

NEWSFOCIUS
~ht:i~: quest as wherher
1;0

the

anSW"e'f detrptnr ''1ueMiom;. ~~il.CitL rO;~][ forces of nature flfG'

somehow di fferelH aspects of the same ~hi!l!!!. S;IYS Aldlo Deeudeea, a thecrise at I,he' Uli1ivershy ()fLj"or! [ ~!Il PmnQI.;.. --rfyol! have just <I I:liggs tbill is eonsi stent 'W ith I::he:~mm,~ dar-d ~n()c]el. liu.e'nl you rrohab~y don't knew whal. I.{I do IliC:XI,"~e sa:rs."'\;V.h~lt 'IJlCIli'!" 'G~(jd~a!i:te' a lill!! extra idillOen:!li,oll1l:S ~ it has th e r~ght masatne ~\ll st lLi'es.eOlIF,;hers S~\yll1ey''u never race lh<'L~; e Hi!;ms and nothing alse qU~:;lkm becausethe UK' wil] d~,~f.,.~(lv~r 'IJwlo'ul~d1 the rela~1 be ~iv& ph::!1:ry cfot her t hilil~. Many expeer it to bi<l:iU mil pRrd{;les pred~:cled by a ()m~cep~. nUed a s,la r di:S,ilsrterl beleau/se supersymmetry (SUSY). which posits ~ heavier "superparjner" for every k~,U)wn 'p,.1r~ thalt 'wolulld me'allr1l there .ii'i"D-c-ul~d---n _ n __, ,...iI 1"'0-' b-e- ,,,",1--'111.· -I!l't-~· ~:h:::.~e;.I~l:1l mi;vy s.eeJJil.l uncl~!lycomplicaesd, but !.!'!!" ~ __ u ,~___~nl' SUSy solves preblems \~,'hh~n the sl3rmdll:rd n,e'w phys'i,cs.!U mod~L PO~~~$W\''''iu'cl~ deeper ~b~(lr~I.~nd ~J(lnaithan Ellis, CERN mOlYeven explaiu t~u;:my:sier~omi, dil'lrk marter \!i.ilJtoS'(;;Sru vity hoi d s ~h~ ~al <lxil;s ltl.!:j:GttHH'; "SUSy is: unique ilm,ll~ZllU~ does,I~lllhese interacts wuh normal matter, andthe least Ihi!,g~ ,,-.u~gi1l'lZlli,,,,,:~~,y:' CERN10 E~~is;;~}I~. $ massjve sup e,t'parllilfr n~],:rghtj lIst:fit Illite' bi II. Mo~teollicrdel)f. SUSV solves etechni\illilh .~II evide:rrillW suptponing ir, :sus\" this (:;lIJ problem c;;Luscd bythe I~J.ggs boson i i&~II most too beaut ifull, to be 'wrollg" 5licn],~e itself. r:hel~,lig.g:s. too .. IHLH'I be :;.hmtuied ,UUiI theerists say. ·'AU these clues c!(mld be misv II:Ul.a I panlcles, ~~,d l}'n~y CiVglll 10 send irs .I(!~di~.g:·WUcz.ck says. "hut lhlflll wou~d bea mass skyrocketing,. SUSY \\rOlUld expl<l,un re-il~ly C'nll,;'!~jeke by MO:lh(l1!' N(.\u,IrC:-OTund in whyl,lfI.e Higgs is ::IShgln as it 3ppears 10 be, IlIell,~I,y bad taste on her ]:N:llrl.·· because ,milben~in~i(,!lny the 'eff!;c~s ofrm:r~The tHe mi~h~ else r~\I'I;'~~rarwUd{;)r ncr and s:LJ1pcrp,<lrlrlerom,t!~.e H~gg!i: tHUS I.cnd pheaemerra, such ~s ~,:~l'~lerP'lrt~to el,eC'~l'O.b1i!!; toeaneel each ,mtle-f. SUSY would aI!'i(l hdp a~ld otJ~eli':;;tl,pPQ~ed,ly .im;livi:;~blc bits ef'mat~xp I~i IlJ ll,l~ erigi n of 1~,cH:ugg,~. ich !J:. ju SI. ter; tiI11)' b.llm;:k wh holes. or even new dimensioss ul~k~d ontoll'ii~ !>Uumd<lrdioodel bm ~~nl;:rgt's of SpfllCC that opn. only iU '''cry :Im C1\cr.gh~s, igh l'Iah.n'a~,lyfrom ll,le stlFlilI,ct1Llreof SU SoY. : The spare "'oom SUSy ~'flUld ~]s.o h~lp uni t'yl he four f orce s. The s to:!lHBu:rd mudd "cCU1H'!I!:> for duet: of rhem: the electro-

cotdd I::,,,,plnin. for exampl e, \'t,'llY grow it)' :~5 50 m~!c:~'n \w~k~1' t~~an the o~he'l' 1:O~e:~,. Some" Ihing like eJ",I:II:<ldimensjeas I give a very
SIH<lII

probability,"

s<lys Mi,chad

Turs,

<I

phy~id~t ;;II~Columbia Ulmii!lf;;r~ilY."IB~H the pol e b1i1a:11s so big t hm :ufs y,el.·Y exei ~.il~.g:· ii
AS!lI!~e

more exetie possihilhies is: parriele phjl~i.ci8l~ say dl;at just discovera,:IIii,sthe Hig;gs\,;'Oukl be a tru.~L!.~';!Jph, the ~Hg:gs is. flruy~hi!lg Hk.1ilthi;;o"if ri stspredict ... ve w,illlr.imld it:" 'Says Perer J~rm,u. an experimente r at C ERN ...··We shou klrl'~ be d i~fliPiP{)l if'we db." P;hys,k~.i!>ts11:$0adm]t that, regardless or : Illite imr[eUe(:~:md loment it. \\"Q~!Mcasse, f:imJirm,g. m~Jrl~ngwould create pl:oblc~1~.S.3l I least wi:Gh du,;&rt)"'enulle"!ll$~~~ut for the tHe ~id "JUSi! in.nag,illleifwe go to, the CERN Coum:i~11 ~!l1d:s:uy.'Th'U1k; you VI~ry muclt.\.~~'\,~ j~~sl :t!pelml:b~.Hi.oll!> of SW.~~!l francs, and lhe're~
~t:I~r~liitccd. !,\uti

bet No:n.e ef'rhese

moo

!'IOd~~il,g ~here:"

Ems

Si!y~.

'''I ~hiif~ they

might be [!, la·d di;:;,IP'Poimed:' 1~lowc:ver. Hndi.!lg on.ly lh~ Hig,gs ]lUIY rn<lke~ifeli1,e'ir~ly as dUJiCLI:tl fo:1" ~~.hy5iciM~ luyi:rlil.gI:Qpern~~1Iidk: .~0\1'~mmC;l:uDlS bu ild the 1:.0 next gr,eat particle smasher.jhe proposed rlllern;Jt~o:n~[ L ineor Coil ider (ILC). COSli Ilg ootween i III billion and $15 billhnt I~c 1LC wotLhl m,lp out t:~e conceptual terrain opened by the L~.~IC ,~St'if!.fln'. February 9 'po 7461_ By oCoUiiding ~rmdiv i101 blc c lcerrons ~uld positrons, the ItCwould ~,~n~r=.:t,tt;:
cleanes
':;'01

Iisions

t~~, shou id ,reveal, ..t

Ili:Iagrmclic force" the \\'U:!lik toree, <IImltJile,sltr-ong nuelear torce (hal binds particles c~nc!l quarks
il1l10 pmtons.,.E~ll[-OI1l:S,.

'C,QMM.fNil'S A/sgood a~ dl'5(;QYl!red. ~:om.e S{!.iy

o
.

d\;'li:;:li Is of ft~wp;:).!tic les ihO![\,~,'~:l l~ l obscured by the ~,nes;sy pro~oM,-onprolmll;.'ll~ll5ion 'l~ H C. L

rBulif the lLC has

(:I;l]ly

the H igg::;
A

to stmly. th~rm,~l becomes .'~ vcry hanl

,:md other part~d.cs. T.lrn.c sl,rength:s mFlhe Ilm~e im: nn~$e It,!,'i (mt!! ~l'Ier~y of,colllo.s,im,1s". ;md ifLhe I,IJmhtc'D'S.of)

t'"

is

~,tlp,ets)"IIiI.1me~t~c.hel'i,JI~ I

B1.!ilSiU!rlpli'l's~~, 5u~'eli':sy etI11 [iIl11!,.

1),peU U'le iJrll!~~ecled,

;t-l
$-'1 14'-'1L

sell both sc ient ifica Uy <Jilldi pol ]].t cnlly." says D~\lid Cina'bm ... prlrti.d(l" ~ p.l!il.y:!i ie isr-t II rn e'd ~ '!.Sit ron a,mlme :U D' Wayne Snue U~,live::rsity j,n:i: De'lmi~. Mk:hi!;'JI1I •• '~ ~hin'k )lm!'U h<'Iv~ ~I reaUy ~~IJcl, tume arguung 1:!iI~ts what
you
\'4I1U

lao be~!J'l~llIJllo be

IiItIrl'lflg?

$~. 0

Pi Ui:mlrolf."' he ~y!>.

bcgruml,IO tllg~qlll~.lly :hlllrd ,It pre~ d!S~ly the sam~!.,;n(;;r,!:lY !iiOiVle\1i~1Jlu::.re bc~,O'w the Pbnck :sc~Je.

M.of~i3!rllllS;'I11red g;ue!o13, l~,lim .;l.1lredkll0:11 Bigg.er lll:ias htti'ttl,~ lliltler 1~1i!~!>•• •

Od~(:rs.S!I!Y sucl~ £.peC~li.ilIl.ion i)O \

prom'I.I~~I'~ ,I!ld pL;;:;,s.im;is;L.ic "'''I,Vf; •. ~re :so 'Lls~d to dUSo-cus.sin,lg d:le II,U~:W

Th~tshot~ldmilke

h e~1!s.l,er to'roU

Ilu;::IiI~,ilrm,dgJI:<lLV]ly tOge'I!iI.erulli o:lrne

Ci!llmp!lJoSill-e: e~ec:'ti!'l!.Ins,
!!illC •.

lerri~o:uy th>l!( '>W;:: go-i!lg w ~i1J~~r are

lhi'lil sO'blm,et1m,e:s\'t.'t:'
kl'b,()\1!.1

gt.md Iil!IiJifil0d IlLt'i~lry. s<!y~Fnl!'~k Willcz.ek. :a Iheori~~ .1J [he M"2'lS<I.~ I.!lw~setl:sh~s.~i tc' of '-;~ChllOklg)' m in Cm.1lbr,i:d~e. S USy Ul ig~.l wn p:ru\'~.d!.,;the ~ d.uk ~t11;mer tlll11t glues l:~lCgOll.I:lx~ ies together. Physi.ci5lS benev€: ~hni d.lrkmnth .. mmH 'cml~i:;;t (If ::r SOH'1le'stabl.e pa.rtic.ie lIK~~. barely

think [hm \'o'C '\lhai w~ 'lrr~ goung tu fiindJ." ~ays Jos E:ngelell. chief,~(ie:m1li.!'il 1iI1 ClEiRN .. '··'\Vell.we don't. Oi"md I I.hillk iI wi n ~ellllUt;;1l !lIiJlOre eXiculin~ lh~n we ¢.It;Pl'~I.''That .~~'.:;Iy be'. b~~.l:~,is much is certa.in t
i:l:be,ldy: Evel,'yone·lll,opes [00:11' ~
ffiQ!:i;l

th"lnJ~].~tthe Hig,g:;.

~ ~

-AJ!)I!I:IANI (HO'

NEWSFOCIJS
.... Iil'Ol1il!e'\W'~k~r.
~OOIli (i(l!!J~ha

II

ns rid R:~rlLmila

~'l::ilil CmlG:N finn.ing ar$'enic:·(.onta fllinated wa~e1'.

and dill~l~ffir"illl-law to

An lMl~hpir
atJ!!lill:iollilml
~(I

a rse nie' s;e,;ourgc. says.A II an Sm 1,th, f~n epidemiologist atthe Un~,ven~~,ly C'iL~ of ifomia .. Berke key. ij;I:~iiiI~_rgeg~ ., lPu~ooilt~g of a r~lpl~~<.lIlh,_J~ iitil hitiilury:t h di.dll·l, have tD l~LrIIJ1Qui, thisway=-cee(uil1 iy nct jn Ilm.d~.~. whose ,g(}vefrH1IiJ~lltl;rte", qlltelJil ]ywUI.s l he Qcmn.lry 'sbu rgC(lIm jug sc i-

enee mid

r~i{.~hl!l,{}lllgyCt~ptl.ci~y. ere H

in\Ve'ljl
to

Beng;..l orici"i:~S> ~:['we~~Hd aql~l:mler",er~~lrry

taekle th€' conramlnauon, (iBaflii!l~d,esh ~e;)1I'II1ed. the l11 real a dec ade la ref.) Yel the of
g(jv~mmil,.i!'H f~ik;d toi~WB~i.g.:f!!~i,J ,l~I(;':(I~!;l~""]y il [)r pro\.'~de J ]terr];n ive W<lle r re sources to .. lfeererl area s, e rit ie ~ C:ll.•rrge, ·'.Fan: mill I) Y

y~mJ;.,.governmem olTicials accused us of

I,ying a:~ld ('~fi.ggcrllling the l'jol'ob~cm:' !;Hy~ den:i.111~ol.(Jgist Kshi~ish Sahi]. \\,1io uncovered the fir ... C~S~!'lof·.Hr$~mli~m;is while ~.[ the t

Sdnml (rfTrnpk,d Ml;;dit;~i!1e in KQlht.",

TOX~COLOGY

,A Slugglish Re's,'ponlse 10 HlulmalDlity:s Bi'g"gel:sl' Mass IP'ois,oning

1

Arseni.c;-1a(ied

w,a,t!!:!rhas sid~ened t~oll!sam!lsillilSouth Asia. A:fmer deLal)ts; a.ndfa~Sies;tads.

l'h011 sands

S~nce thee<lrly IOOOs.when [ndim]a~I'I~.o:i:"'ilies b~g<Lr! orespond more VI.~Orul!~~to Ih.e e t crisis .. sn~t,em~dmHiolla:i: govermmlents hilve pl!mpc;d lens, of mimOID of dol lil I"$. imfl MDIH~ rionsauned ,H pmvi.d ing safe water, The f~~1iIiIts ha\I~' bcc~I<!Lc~J~s~t:l[; A S.7 mil honillilimi\"e'~o iiI: wells with <I~I~ic r.llr.l:l:ioiD1!illnits r~ln~d heeauss flf lmproper !'!"Hli:JmtB~a:~ce,.
Another
s.~r<ltieg)!~d[ull~,:lmg deep \:v,e.~ls~:~3.l aqilli,:Wers-has

I'ndia ~s,addfii:HlSin,g ~1i1:e'j':olbl,em 'Mth a $,500, miUio,n safe~wa'te<r 'iflliitiative' p

,C:HAIrl!:iII[!,AIl.lTHI, I~NDIIIA-U nti I dllcm]cI-1990s. ~he bi,gg.est :!be 'ofGotvt::h,HlI, and Re,Ln~h'lljJ.Ari <lind thei r e.:...tended nun iIy was poverty. Then <'iID'I~ll,rt;:ill~id~o~l" menace 'begtll'l tfl:-ind~ Ihe AlI"i h01l1M.:~[L Chan,d<lJ~1hi.~1 dlll~~lef' of mud huts {In the Io<dgeof a ye]lo"v llIH~s~~l!rd fie]d same 6[1 kilometers 11I0l"th of Kolksta, The' fri:lI's;t :!!:i,gll.~S of lll'o~1.bi'l'wcr'C brow.~. ~pC!lL'S {Ill

bypass arsenic-tamted

pru-

dueed mixed resu Its..

of !1"nnU les
i.bl.[~C'S!ilIile0 nl!,1e~t

Ute

most dep:r.o~bl~aSj.l~

Or[~C tragedy •.

critics :·my. is 11l;1t Indi,11Il omd<l~::;' ~:\i.,.ve resisted Bengal hav'e been <~frected by this biigl.H. 1vlore: 1h'1I:I, 4{1nl'~~iOin. peopi~ here I iye i'Im<lllieOl:5 edLlC'dllng vi Ihl~er.'! aimul lhe threat, p<!ully mil I W~'~'~ ,e~l.;':v:HJcd 1~'\.'B!sof miI~turflHy w':Clllrrii:1,g (If concern fh;;LI lfulb.:; C(luhJ 10::1d to ~l1c,~ewll arsenic in the !il roundw,lte:r. AM.lhol'lti.es estimarethat 5mill~u!lin \\'~Sl B!.;l~l,g~~1 drink VI'<IJler v.,·,i~:~l arsen ie COll(;eJilln:a~ions above the
gmrcr~uncll[ sl!l'I'Dda:l"d of:50 lll'llemg,r'l;tms per li,~:er.Iu l1ei.shborj,D1g g'In1:;I<ldeS\h, more th'Ul 82: ij'ijl~II ion P~()l:il~:1 i11 ive C!lin~lifii~atoo !i~at, . 1IIlfei5l.111 is ~i!surtc:{)m;Ci'mb~lble, ~'1y.s DipmllGlf

Ch,,1\ rr![llH!i.

;U~

'~ll~!,d~Ollunem,11

seleu lis t ,U

J;;uhwp:ur Ulliv,el1>ity (lIU)1 .UIlKO.liGli:.l. "I.f pee(,le. ~l''Cl1l:llilll~ to rca:Li:,;:e the darlgcf'l> ofdrilddng
<l rsen ie-cc I'IIt." IIIinated

~heili' hands: a:l~d feet I,hat. !IS the monlfus p&s~~d. d("v(~~,op~diij[o t!:iid eO'lJllus:cl'] and lesions. h, "'<IS several yei,lfs later t~.l<l:~d:octmi15 vi !I ~t~~~th~ ~'1If(:'1rel)t.l:g!l i?'!4d tlm~ ha n~~ ark :;y.l'illptom~ of arse aic I'oi:-;~ning, TC:SlS conflrmed th:'ll water fn:lIn [he v..'ell

water, t~;e)!w~~.1<~ke t

eare ll"·h:lir

mV:fi ~'1I~ry,"

:~i;: yt>. ~

A Ildl·h.e 'Pm blfllll is \\rideln i[l~;:ln recent f.es~a.rch~l!"!'l have

Yeliits.

West Bel'l.W11 ():~.~OC:i<1ls Lckm.mvlet~ge thiOit 1:11111;: < st aie.errcd. [3:~H they
~y ~6l,! h,df·b:iUklfl"' dollar initiative now

lound hi.g,h levels 'Of

grmLrI.:.i,w,JlfCr

~he ;c\rgs,were lusingw.t$~t.cle~;lwi!h arsenic, :

Their oldest 5(l~'. 31ild his wife
\lIti

in

several
<1.:11

arsenie ot her PnJde ~fu.

were d!.lIgJlflSCd

Ili1Id !nll ~t~l~s..,~ i nelud-

~hskin cancer; a d isease ~~[Ik:oot'l.'nth chronic

exposure, GOUCh~ll~~, old his s r~1y

i:1ID,g Un

low-levelarsenic C(l\'V. goucs.

IMhElf.<Ind M<lnipulI:.

under WtiI)' re in~~~uU eighr surface-water rre.nDllllelll ,1pE,1 ntsand 360' hig~,l~cap~ci~'y,

i'mm,e;"
9 z

for th~ir m:;)~mi.::f1l. The Cl(mp~e died! <liny\\.... Alffi.·:;,:i:dof sur~ li1y.. f~ri!'l~the "'1.1~1'1~ f:Ut~. I\\'(l )i{llml,gi01!' $01'!J5 mO\h"G ~D oflin.er:I~I1IS of[ndb. "An;cni~ de.stn:lyed mJ_r s~s GC!uchm], ,~Ii"llil ,'6'Y~II:"old ,villi) ~1!'.,.~b",vjith;) :llimp because ofalS:l1Iic lesitUns.

and I(.luc'k$ to

A~dlaUJ::.!h [1~~ are IIlI e rel i:,lble stanstie s cuarseaic vict~msun

I.rtclil'l andl B:mI~gll:lde£h ..

d~p wells
long,ler

fiUi;.'(!

\vith
fil:Cil~

arsenie-removal

hies willprevide
\i!j~':;li '.<Ull:~r.'; as

In :!remedy '10

one reseereh group

ih3S cQul1It'ed at [e<L:st

the biggesl
MOIrik @i~i:!i kimler. IlJl1li1lUb<l~!:li',~ pi!~n'lS be.a'r the b I'GWI'I (,.II.IJ~t's, lflm ilIrrea hd l~mil rk :S'!l'mpt.clliIl, III <IIf:'>t'llIicol>i,5.

one .of public

'Ili ·'.I.'m li.r,ed of showin~, mv ,eal.II\Ii.ses to

; st~,'ans~r"S,~" _.,.<1kl1s ReJili,ub<lI;. ·'\~U:l~'c.mm1der~ all~se:L~:iIC{l:S,:US West In .~ sH~lld OUiJnl~~!:y~!" Bel:lgal .(l~'O!ll,,>. 'fh~

~. .,000 4

'CM~S

of

hCilhh d~S~$h~r!\ of t J~,emode r:lI1. ''''01'1 d. ··V~, w.1ilOO have b~ell

NEWSFOCIUS
dl;lhl}~$:" says [:I. N. Guha Majumd<'1;r. u g(llSin.'"Of:l1Ite rolog~ s,t 01'1 \,VeSl.tBeing<ll·s arsen ie ~asktorce, "BlJt l.hegmrenmJm:1C'I'Il.isaCli:~g 1'lQ/Ii.V;· f'n~$lra(~d~C hakraberrl took off his gloves, (h is e<nsy to mi::lt<lke the: j:lugmu:io1:j;s
:!i:c.iel'lib.~.stor <ttl aetivi st. He once scolded the f suue 'sea:lvim;;nuhelill milli sl;er for !-;moki.!IfIg '~l,~ meeting: 0111 il:~oLhcr ecesslen, he adviseda prulm]ill(ml arsenic researcherte ~;,~keI eeurse < i11 wm(:r 1~z::t.ing.1ChfLkrab urt j nl'~'lh~~d an

lOOe]1lDnsudledin al''eflswil:!a ptlwbte !;l.rolUl~dwater, "\lhil~ 73 were oolivel:ingw~ltef with ::Jirsen~.1C coneentrarions .above the pent]iss~.bl.e lil1~,il.A~lld some ~.15unirs ill lowed water
through with unaeccptablelcvels
Oril'liI~ .•

Ulrnhllhe mli:d'·i960~. much ofWe:Sl BeIlS.11 I",e:lk::don m~lU'C=lu~d ~~u:r nnr.!l1lllm~d&. ~vt:'rz::, l!l!ld op(;':n \i\/i!]~: 01."- a ri;'~Mh. (!.ilOh.Ttl ~md ·O!lhi:f
lethal ¥>~lle:L'lJ{li,i",ne diseases taok a heavy to:~L v!"ells,pipcs bored

A savior <!rrh..edin th~ form 'i;lf!Shf!~IO\lI'i:liilb~ ir.u~,od'ne g!L'iClml:dwith 3,hand ~. 96.,p .• ~·74;.. llep~I:I, vietims frol:li~. 9 and eenpmnpm lh~lOp. Vi/~hen~U:~dU10~t}gy lllr,liiflkrer, ·'1 bd 19 arsen ic pu uentssi ui rI,g in the in£~I~i~ son of'well became afll)'rd<lib~e., ~lt: Ilr$!~")w." he: S<lYS, Se've:llitee:~]~(\'ves.ince died. l
The ~j995 cenference made Chakraborti

int~m~.~io~la~ ,conl~f~ne.: ~u JU in febrtli~Jy ~. 9:5. at least inpsrt, he SOl),!>. 10 ~'I~TIlbaJriL<Ls:s 9 vffili;'i~!I::! into lil~:lion. tS{:'itmctl.11 October

~ov~rnmelll ant! prlvaee Cinzens begflln, ilm:s~~tll them by the HlOlISlrlmds. Deaths. from illi.fl,
imJr~Cl!O~liO

persona non grdt;1 to t~,lestate !il9vOOI~]lnenl: He

d~~,,"~S fell SIKi!1)br.

\!,'i!'!US

But the tube

spawned

,3

new epidem ie .A fter S<L1~ni rsl f linked bro'wl'I clll]us:el' to groundWSilccr (I1"l>C'n te i.:Uii, 1981. I~H~ ~t1e=:l1 govermneral ~]()roillitetil: ap;,~ne] to examine th,e problem !l~HIind f cn~~~ltl.::~unil;,''' :;,~~r,~~. Over thl;i r!!;':xt 5 yem~, reams lunu lite Sc:Ilf)o.~of

CH IINJ!A. N E P'.AIL
I~Ui lIlTAN

"Overall, d'ae ~H~t1y shawed ~h<ll ~2% ohh,e [units] were !lot useful:' rhe researchers i\iJH~nGd~as.~ y~<.'!rii'i Ithe' ~r'ht.(:'t QUQ#/.~· Resf!m·,dl .lolluwl ofCal'l(ula. They h],mu~d .it I{Ill ;_QI~~:k (}rn.~l.1ini\.iJl1<1nl.1l;l. includ i~.g al~i Im~ ~~~ perioo~.C'a[~y :repl<tee adsorpricamed ia, \M..,M Ikng.ill o~lici,Jls disp~~h:: Chtlk,·abm1i::' .~h~l'itysis,Al1IiI:iytl Bal1t'~jee. ch~efengineer of the lliWt~'S Puhlie H~a~th E:mJgin.ecring D~I"€'CI;Ol1!t~., d;1 i.u:LliS rhar 10% {I f t:lm,e ins talled arseni c~ rCluovOlliUIliIS ~l~~workin,g fi 11'£. :Bm he aCk;~.Ol!!.;ledges th:nmhey rue [1m bC'ill'D.g lfIa.i:~lt!l.~ncd well. Even before h;_~kl!~uboni':sSlm.$y 1:<11111;:
out. l~~.e \VC:l>ll],cngQ I gm'C'rI,lt-ue&n ilUiearly 2006 .m~.lomu;ed t1.:g<Jt it

'\:\'o~l~d no Iengcr

equip

I~Mf!d

punlp~ wIth

anicen~l;.i-!1:m(I\..,l, u!liL".

""\Ie re.<llize,c:I~:~latIbe @O\'Crmrnml

TrtrriC'a~ M:edicin~ ..the AU Indi .. lnstirure of ~~lyg1ene ami Ptlb~li'c [-I:etdlhl. ~!JJdother ]IlM:lWUOns doemm,ented.evidence [)fchm~,ic rexidl)' !'Itnong hund~d$ or'!.':i~~~glJr:s iL~, dustricts of\~feSl iBeng<ll, six AIl'hcl,U.gh S{)!n" e:xperl!f; ~~:!1}1 Olll blamed tlli.eillnesses on indns~:ri.j,\lpQnUlio~. it soon became d:eiJr Iha~~'he cu IPl.-it was <l1~~.~i,c iualluvial aqulfers, Re5e~rehcU'i!
sfud}'IIli,g the plu:~Q!~TI,c~nm-S!l:elli in many other eeumries, including Chin;;!. Vn~m~m. Chile ..find the Ul'li.~ed Srares=-now believe

'C"l!moroversee the mf!]m~m!r!~lo,:: ()if

lile.<;e units," Banerjee snys. C hOI kn,hon]'s grou p has <118.0 :lSsailed the wov~rmneln'S!!ll,,~gjl for bari~lg d\;:1eptube \"l.:US i~~di~~,cr.imLml~e~){.rlllJe 'P;;~'i:l '0yi!t'If'S. the l 1
l1;.~re~.t-n;.hfl\!L;:liJum] lI~rHat ]~~Sl 20 deep wells leO to :Il.50 meters deelP-iu~ West Be'D1Jg.al'sNorth

I NIDIA

24 r<lJrg<mm, and M~II.1ihid>lbaddismets, havegone ~,hnnha\ ing vinlil-'

B ~I R M A

'11Ly no aU:llenic iI~ the wawJr <lll~.e

~

!~.m ~,oil

~lIlV~rol11i.'~li~rnl:t;;

,lfS£lrtlk

frnrn harmlesspymes

ill;l the 31 ~lJv:uurnl. Ope,n,

we Us, even l n ~1/rJ>enie-rich airel!s, t yp h;,~.Uy h avel 0\\' arsen ie ccncearrat iens because when Ih~ '!!Wller'sl:~mdsxpoS!~d 1.0 iii" :For day,t;, ~ il fh,c;: et a I binds to :irr-OJ,1x ides.and otil.ler comm o p!)~u~ds @:~d~'I;ipit'l,tG'.~ ~Jut of th~ w~ti~r (:01m~H'I" Thi,8 doe:.:; 110\.~lrpen iU1l ~ndos.ed ,Ill
tube we~,l.

rn~ ~su~~s, n~c~ivi!m,!;lI~ss;thl;m h,,1 f f,)f th~ ..... n."{lu~t rUt Yifthl; nWl14;:Y e/';p'l~uied tlrIi iniliudYe to :sink deep lube we~,ls iJIl.:'I1l el'ilb·:rr-t to Hlp

say s be has been shim OLUt I) f gO\l\f! r nme Ilit~ spensered $lh'::~1ing:li and accused o'·lliieiUii.g a tf:'olUI;or. BlIl~lrn.eev~mlmdl an inmpt\cl. The ne~ :t ... )'@Ir.\V0St B:il.@1 CJilfk~ !~II)II.~I~d $200 IIlril· iab I'uonl}-o:~] I:he :m<I~li.omlg.(wel11l]ll1.en& :ro:n:omncf'"

ou~set to COJJOO r1UrHl ions ili.".ng i~lg fr.fHliIl 511 'ttl ~5U 1'I'1licfllgr.!llnspeii hlCl' I~e;",cemillg the' SO.JI],lk:~gaTIDn [i m it) 'within 7 yli.~r~. E\.1(;l!'annw, people SJee~mL1 blind to the risks: In inrervi ews ~'I'id'll. St:!.f/J'Ut;, ~om,c v~.I~.~g~rs Norlh 24 in r<lI"j'~mJS d ~smCilsaidli:l.ev ~flJIS'ted~hll~ the \~"Uf!li' .~ ~ 1~,Cl' drink from one uf t'~tl~(; \:\1'l;:lb;is ~a~:~. Ch:iiL1kI[':",JlxlIlI'Uiugne.;: t~<'LI: deep tWI.oo\\~II:s shmll!d '~. S1,i!lk {lid}\, i~ 1!~~'" 'vh~~ d~n,;~itoO thick di.!.Y lit hiiwrrier ~h\;~~ll the sha ~~ow. <n~n~·coll!l<uninated .,.q~t.iftnmd ~lte deepe.r aquifer being

I
~
~. :::f ~ ::;;

i
~.

TIle crisis pe:i,stLu,ued Chakmboml, to svve~.p the U~illXI Stales mui ,'ct1ll:1l to h~.s, K:olk ...m .D"OOh· ,ilJl 1.98S to fOll:mLcl tlilc Sdu{ll. of E un.vil'o:un]rn,U:'I]I<L~ t~Ldlies at JU Ove. l'Im.e ne~u S 6 ~f!rs. h~'<'I!ld his gro'"l~r·~l1~.~~'d~~ndred~~ of 111Ibe wei]s_ <1.S,""''C~<I:S ski nl. 1m Ir. In.,:il~. a:ndl urine' I gl.1mple:~.They ~1'om1:dt;h<l1 ~!rsenic ctmwli'ld!1l~~'_ don ""OilS wideslue<liJ .. lll smue aJle<lS, Ih.e gov~ Crfm;lell& du.~d~p \veUs.. B'lt nn1ck"llsdi!l'J'l'lJtl1:d ~hem;;lsni,lude ofd-.e pmblen~ al1rl~Qlmom'l;d a~~'s c fmm e~'nak"'OiboJ~iand O't·fuJerslQ ¥l;.'lmG;5s\.Vcsl BelJgal:s,plef]ui1:"lIl S,lqrn~(.-e~wat~1i' rc;mqrce> IQ~
>fII ('a1'OCf ill

IT! l{.l(lm~-l'en"l~.sol~tiu~.

lunHlin~~d \\'ateD' OO~O\lI' lilt: ul:l.uvial .. (~uif~rs,. ~lppe;;d. "Subsurli:l.l::e g.Gokl£Y ~I~ould 8uid(;: I(he A.ml f.~ 1998. West l8:engill began L'<lu ip piU'llg :snllu~y,'" he says, = 2400 1~lI[ld-pllIl1ped shaUo\!I.,t1i.!be \\.dls \~!i,t~l O~]le[!'i di S~J'.!.n:e.. .Ale.luu'Ider van Geen.'1 oil ~1.m:nlic'«lllmllil[l~ln~ IS:: S~.500 <ncisorpliol1ltD~,\'= geo~,ogi~!"U: Co~mnhu3U ~1.1ivers~,ty. is, a :stron£ ~ .., gudVU(",H!;l of dee:p rube \wll:> flS f~S,i:!.o~t-~~Jfm ~ ~ff, ~C:k4;ld WiUl $LUb~mnc(;'S such fl." ~lwlITl!ma IOf £olmiall ~ilnEkmglo"de.sh. [Ie says 'tIhat l1le!C'hHllii~ ~ iron O:1(i,d,e'g,~~<l:lfmles. n.e g,HUe .!.Iso bega.1l l'Olli 1~!lure [)n~u,~,v~lh.. n(l~u~t'~,ie l~ch~rQ.\:!. ~m~l bl!ildil~~ ,"' l>~~rr:f!;~e-'\¥.rH'er lt~mn~~rn't'phHU ill (leeper ~LqLJI.ireffi. is, ~~)blrunoC for the hllndn~i of Maldt"l, nue of the' U:QirsD-·'ltl~~eddi~trkts;, v.:fL,~II&. '··Beclll.J~ on:l.H\,,~d1 ~om;.tl11J.C'li!OIl!, ~. for Cb~krabmti.lhis 'VIIS not ~'lol;l,gh, ~:~nt~d Be~'1\'e!;~nH,lS1'9 ~]iid 2!l15 .. lw .md his,oolleag'nf;S you ,end,.]) "b:m"i Illy the :s11<d 0\'>'·wmel.·:·says ~ 1 '\!l:llQ GClCIl,,\~:lf.l~ t~ell"1i [flo;)£i. d.QC'i;Jlr'!I1.~nllt~ll I\bt~v ~ C'va!J::i.atlild ll\!$p:;:cu:Fo],l'n;;m.c;c of twar1):6QtI) onh~ sm::h f<1lh~I1e.'ll (yut of .51, ~eep !lube \~'ell:s InQllli- J!l ~,llIld·pmnpe<B ...u;l'ienic~rcmmral t1lJ1i l'.~,TIley ' U!l1l(:u\~'II'0durnf~~~€rol;J1S1 pmhI6111& I:iil), ~ 111 Its had rf!!'(;d {YV~:! 5 ~~~ i!l the A:rrn ih"t.:O:1J fegion. illl ~
::!l
1

iii.

NEWSFOCIJS
B OlIm!.gI.ad0s;l~.. A lt ho l~gh he ag.r~~S~1i1 irh Chakraherri S PllSh fOI':mrll1ce \V<lter m; alongterm. w1mial1ll,
V:!IJlll

II

Gecn s~ys.deep tube \~rdh;,

;1

reguhuty n(joni~or~d, are "bardlo beat' as

AVoID no :S,ci,elrtisl :SIlI18[p e d by Aldv1elrsity

l
l

sou,ree of ~fe wm:er mer the nCN:t I0 yca.rs.. TediDnica~: fix.e!' m,IY be dehared .. but !u'Jlboodi)l disputes the need fur better p~lblic ~1i.;"'11ren~~. VtJ<lW: ~~g~ \V>l.'$~. ~n:~1 g~]\i~nm~ I.~~e menl. d.ocidt>d liD paint tubewells wlillh potable
\1i.'H~er :~,~~ g{~r;:::Fi~ndlle~!\!\':tUl!i,Olfr;:: l~lb~t~:~h b or

IUUKA,fA.-Filf$l his, I!Il11dJe ~1,I(r.umbed. Than Iili$ fath@f. l!fl'8l1 iii is; 1!llUlit 1111 2.004" whu Karlik ~r'~'mIIiis.a\yhis IllfIolher.[) ~Iali. develop Ihe ~%!ll'IIe ~esI!l):n:s~ha~t pre~9'f!d ~he diel1!llh of ~is '(l!l~er ro1H1lily members, hE1'tnok tI 5-:hour rid~ by bus

""nd uain~'rol!l his'!i'il.l:::!ge i~ N'i!dlia dislrkt to'

r,;:olkata to s.eek (l1ll'~.ICHpallka r Charlomabm1:n, iii HIlI:s~d ing arse!'!i( rlliS~i!!lr(lher liLt Jiild"vp'u~ Lllil;Wli.r:sily: lille' em:ol!lIitf!r cnalfl'Qtd Bis\V~:s,'s life. (h\a~r.al)Oui hel!l1e'dl gel nula~i aldmirted 1Ill
;ill

nm1tJ!<Irked-rnJ:I:l:ier !l1am,pa il1;t:them red as sllg~ gif;lM~d lhl:: gl;)lh'l;;Tlllimem\ arsenic ~.,~k[force. b:)' "TIle nd'n,~in~:\tr"liolli dtOllght d~!woulI.! create ll.l1rn£cc&'it'lrYpi~l1ic,'" ~mysOl.ltlnd,Et:1ll S£n!:~upta. a !<I!l.kliorr::e me.m:ber \'.1h.O fOrnLe:liy :nmIull<lged a

QIQVeHHII'II!HIII.~HH;~ilta'L

Issts cenlinaed

UN]CEFpn1i.;~t ,:dml"dall;f~c"Ji~l!i!. <l~~iC' tb~

problem, W~.U:~~.1 JUresel1lrcil,~er:s :i,n,the m.li,d.~

QlrSll;!fI~ (osi§;" ;[l.Ifld

$~e

rr,J1l~'erwe uU a ~tifll !Jr,a~l en

.1900stook u upon themselves to pa.ia.lt ml!>i!lfe· \1ltelhi red. Chuikill<loow~'i ~YI>."()lle 1t1".;is~~~ltldlltcl !lli.e ~lIbe wenls r.mjnl>OO. ~t:l.~ and~~le~ll aml:Jl.n~d, wirith a loudspeaker tel lin IiDg vi II ,Ige rs that the water io~ I!J~m,"WIS line,'"

Ko~kat;~. Dim :stain.\If!!I~s <Ire :!lpla~.rer-ed with red Gmllndl~"r.n~r '-\1 I~ conti nUl:: to be the l,no"i I1rSI<~y. buill. Together wit:l!m.three plants <1:1eady r ll'1r<lrks from pL;;.t'lph.:: sp.iuillig J){!~ILa llop~l.I~J UU1ld:~r~ $501] mill:h:!J~iIIlW !ll~VCfunded johnl)' eommisslcned i~North 24 :P<lu~l1ms.. Seuth by the !i.l<1le <I'lrl.lI mU1i:a~<'L~ ;mrermnel'l~;s:.~:vesJt 24 P;Jrga:Ii1<1~• lIlid Malda. lh,ey \,,'~JI serve • sua elk ,r,;OlilS i~l iIli~ e f fr'l~ !'31U cond] menrs 30% of lh~ af.fuc~edpop~~1Ui(l:n.To p~ ~~)r wra.r~d i.1"! (l ~~1[.1~.I~~.h 1:~r'l.l!1g\!Jilhwmt..'ilY& Be~,!pul II>sinklng 36fJ lo:lJ~'0-diO:'li!ml~t£r deep mbc dors making tea on :lli:ltle .kelcselne s~oves, .!:Im a ¥''e~L~,eq~l.ipped \VUdl HfSelmtc~tre."unem :rlJ~ illities operon ing cests, the ~O\rel:1l.1melml:\" ..iill charge large, open olfice room, desks are covered tn pipe \\'a~er to 70% ort~.e alfected JiKlPllbtiO'~. famil ies a eouaectien feeand abou~.$. ~ <I ~'i.! l:hmo(lliIt.~ ins ofpaper, i h~ltm;1ny of the Government ,~ngin~r5,.vli ~I ::i1![)erv~~11il<iliI~~e-meruh, T]~on 's ~. ris ky :HI:ll:~egy. lndi .• n v i1~ di:m::lomte's, d!~.rks and mid-n1]lkil;lg e:l'l~p'~oy- ~arn::c:C1fthe m'Cam]~u plams, Ba.1ilc!jee says. I,agcrs ty,icaHy don't IULY I'a:r.water, \\r.~,kh eC5;J[I.l;l nowhere m be seen ..Across the hall, ('i11'u~kr<LOO1:it:iiooru~Lni::>lh<Lt fini IIiIg ~tl:c~,we:11 [) flki ,11:5.<Ldmo!"dedge ma kes il d i~11.C:lIl[ to aides s;tand £_uo,,:ro: outside t~~.eriva~e offices. wuth <In .arsellli.c-tre<lllme'l1:t p.~31~1 .1. w.aglle of gel them tiC! .\iM.irch fw.m. ,I publ ietabe ~fC1110 p is of s~J1lior ufficials. such w" cl~iid (:I~gbfil.ee§ ~lon~y. ()rm~i:d~ri~g ~lJrJi'I!t nnly a small pereem- ~I piped \"''!l~lef~tYmMl~eli{Yml"1 ~ the 3 }ie.lrs that Arniya B,merjee. the SOli ~fu.24 P.:;rrg.'lIIl;I:ilp bun has been in oper~Ige orwell~ is Hkely ,lit ion, OI]~Y 25JI(~O~lm !he 30f.UJ'lIO l~()ml;;f; of 10 beemae enntamiB·~~~~di~1; ~Yf' Ihm nltlted.. \fan (l eel~ lOO urteudedto be' covered by the plaat have ~IH~' gcverumenr's hand ling of t~u: labels tbe pb~.11 ~~!l lak,cb1 ~ .(~.oQlIu;:Clion. ~r$enit: Cri!)i.) i'll now fJ.11\\'ed. "In SU;l,UJ of O~lile\F e:fliol1ls ullider '1,\I3Y ~.nc::~ude '[~le dislllii.Hlbu:ii;[. "\;Vt,: k~{;,\M 311 ikm~~l!:~iC'mh"i]; :[;iud ~ driv~ 10 $;eni"~ lip tlH~ u~~n- bUllOijof Cht::~LP' ~:lmsu[e 1~<]taU6(10.000 pll.ivlJ[l::' ~lII.bewell:s <tilre along thill surfHc:e ment phlllit:s ri~~ll W;flt~r hm:l tn b ~ ~h~ I;'i"ii\'fuy.il ~.l"lkl;':~ !Vllnr·r,;: i!~~t(;d for !,l~lii!.i~, Nk~jtwl'i!~bfJiI'\lJk>~t B~I!g"rs; ( :;;en~e ~o d~~ign ... 'Ir.'l!l;!Dlic [u:jkl(m;.~ il:N.pecl:s, thOl:t eVl;lfybody i~.l lo:vu.g-tenn ..ohtt i'n:l!i, p:mmpf:ng !>ys.tcrn d~.at West Beng.al win ~.13ve !';.al"e d:ui~;nking 'w<IItcr b~tlv.'Cmfc~ded <I qui.c'4>. could ~ (Qrulec~~dto lI~~~lhi fix in lhl;;:im~r.~m."~,e [I 3 ~"ffi" says,. n::fC1!r.l'hlgo the t CIUlkfilhoni h;. rJ:llt ~s optilmrusde. Ami he Bill aI:5~n~.C=lreiltl1lent deep tllbe\lIre~h: i!lJJd mool~lein [he fm~l r "~u[lclersho~ mrmy ~no[epeople wi~~sul"e~-if rei .. <II,.':Se'~jc=[eJlum,..~~ its lUll Ihe need!. ari s.e.~," be "\Ji'1tre:lrn.~sus not I1i1Irtlde an ua.'g1!]mt ~Jf~Ori~)i.O.lm <I d:ii'ukt I~ menl mrm with for h~llld-p~,Hnred 'D @~I!l.rli::t; umerous.a fS~mit:·r!!'mo'll'i:!1Ull it·s.irls.lillll~ says:. If Ih~ g.ov~r~- rl;;c~mv~~l ~!IJN;ld~,11 N ~'I'eH!>. B'lIlcrjee a]so ~)llhe W'ElslImllgall ,£!olJernment no ~0lI1 ge~ work 0100 rm.em lis deu::.mli~.ed to. d,<Is:si.c an,enicos.is. l!csions 'W~;l0had never b~fnre doetofS di:lS;no!<.i;;d d~f~nd5~h~d~;H;isill!1 't(l' lad: of malil1ltenafllll~. inSI.~~ 11.<!rge affi~!:lt:- h~ard ,ib('lm I.!!f:>.eoic: i u'ea'lm.el1ll. pla:lITlts.. be o 1110l'[0 IH'I.inl m1Sa:~e hi!!: d!ise<l!le~aSI December. '''T~~i!> 1n1l1l d:~~nk <1.rgut:ti,.i:l we,,!d: be wiser U;l p\I~'Hyw~l!er fif;\llm c~jlUntal!r~h~3tC'd \:\:'~"1iler for ~ars ~n:d!ilmCllllun! to l1,J!hc w>L,!~I~ cX!pll!li~.ing th!ll doing :oO!wol~ld ml 5 I :i!ke ~;yb~T\jedl:terred vi ll~.age[srrm:n dr<1\"':~ng slmUm"l. oolll~.amimHeclaq~I~I'e.rsfind COl~~erV'e sen his ~mld to find mit '\;~"ll 'i"'~ sulfe.n!ng he z rro!1~l." Cha'krabani say~;L "'~~['~l h; liai,\pp~m]flg 'I!;! wnoor for sHf~u~~ ~uch as was:~i!!l:!il.'. th~d~i)~]".a~n ic-rrc~aq~lif~rs for futul'C llS,'C', AI5'o ull1l.cl,er[~n.c West Beng;J:1 illiti~ui.ve., he:11'e is 11gll:<tV't; ilJjL~S~iQe.:' ; 1111 \,1U1C, ~ne~ee S3YS, thC' !.'1Jvt:l.'um~.ellll <Illy .~i8 nOI\'\I'i Inp~.enI'U~IUngIW~~!il·l~rm S!~) n iO):l , i h s £ilii!,~'tU:;'C-\'\,~Hlrrmr:etHi!mH~~l[ i~~~t'nt:s. a~ b~hlrg -VUDFlIIJllit IBIliIAliIA'CIiflPiRJIEIE

Ov·en;omi ngin,ed.iia l'h~ he~dqlmu~~f", of'We:,\il Bl'l~g.-"rs Pbhlic ~:ealth Engineerh~g DireC:I£ii~'UH! is: [ocatal {In l ~~e~i."'lth flOtlf of "I dlull hugh·rist: m the ]relilll or

her pallllll. Afte'r bri'l!lging liIi:1i. mother hSllilu!, Bnswai5 :51~.reald the wlmi Ol'lim'E!llii[-~,or ]mlalu~, by Cld!ofi.~il!llg \I'~IH;;Ige,f!;whrll(h (ube ~'I'.~~l~ had le~te'd positi\le~'or ,afS'eln'i( i~l Ctl'.abal:lorti's lai~-andl clltlect(?d r,ia.tersan1IP~E!shH hmlllu ~es!.ill1g. Ill!,2~005, IIIU~~li 'g ot !>kk i1Igailfi .:I 1iI!lI warS d!agrto'>ed w~~h !>kilflc.aln,~elr"IIII:mt.rns 11 fiI1I Putat'M ~er .tJlllllii•. ~lI! t lliieo Coli,ti~r h:a:!J aJli',eady spreiild" and' 'S Ili~ di'!!'d loast O(.lJijbi!~. " "lis f.aL!, Bi~li'IIalS,wn~ ;'5 (o1iliI.p~~U~SI.a1 fITI~tHer's d~glr~ i~!le@gW\!l~hyalill h,,(a~toHege, wW.joun Ch~lk~ab flnl's llilb {IS at Ph.D. slude,ilt l'el :UllIdiy 'Q(iUiflt.ermeasu rtl~ ~o g f(Ql,Jlfhd\l!Iattl ~ c@:F!talflnifiglfo!f!, 'Si!l~ Bisw.l!l5,. whooe pilhm illlld!eet Dellr the nr... rks or liI1i ld an~ni:Qos~s,. '~~ob®dy~hmdd hllv~ to Sil!E! tfl1e~~ mOI~her sllrh~lr ll~ke I did, .. ~Y •.B.

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LIETT'EIRS
~'ll.]til:::>.oJ rm~~idti ~lbmn the poli4i.i bi I i~yof rIMS h.1.\~ng ~Ol1g-~c:nJil,l, ~legal~ ve oom.eque:n.ce:::l. or O~t! Iy• .100111.1.': orthC[ll~ ~mhors I1m/(': ~1 ~duTMS tn treat nelJ['oiPsychi.,1l1nC disorders OIl tl'H!

hasis orit!>.long-l;;l!st!!'\~
JERRY GUI()'S ,ilI,RHCH
cOUint. 1l11.~perhaps

!'ilfi:i.:C[S

r.?I. Ro~mtgen's
h<ll.111r]!ess,

~e[hil1l,ology "'!"'<lS, aliso once Lhou!lln HRIV~IR DOLPHlIt4S

I.ULt· [News
w

DOWN

FOR THE

:!Illd

x-rays

\,\i''':.~'~

used

10 check shoe

sizes

oft-he W(lek. 22 Dec. 2006. p, n~6(1) revealed 1'10 :s:~,shl~ns:s of <I baiji cl,g]l:plrnin (Llpm('s 11",~'l111{t:j"Jd~Mng!in,~Q:m c~n~;p~h~llsiv~s~r\fl,;oj'. [f. <IIsexpoected,I~,lis species iSlmlye'li:,Ii'rI,ct;., thentheloss to rorh th~ nah!f<:~~ and phySi,icoi~ ~iG11~~~ :~='g]l~lru r(liTl).~Qmld.thun mm;t realize, This ananal is oncofmlly two species o:rrivcrdol,phi~cS fl~c othc II" 'iocingthe G2Ing,.~1O ri'llcr dol phi:Iro" lrlUmi:yJtl 8Jrm~t'ri(Yl) P liml. allholl:gh bereft 0[\,]$.10[1 and 01:6:<11::1:0'1" sense, ,ne able ro m lgrate, loemle prey. :;Iml Ilndmstcs \~~j[e ,ll:IJ.vfWJ[I.,ingul")J a highly dymulIiIiic riverine ellvir"(n:Iin~:el1il: ( It llfhe bi,olQl;Y ofoot~.lspecies is poorly knewn, Hewevcr, illsig;fuit ~lnto thdr biol.(l~y eeuld be ~,1<P'I'I~tl;:d liO It;ad vo ~d\l.a~l(x:m~fltf;~n aeeusueb<ls(ld sensors, £:~)I~Jcm~orl,.md ~rwi~li{JI!il~n ,i,;)xtrt;!'!'\i;l ~lwiro!1m~ms. as,\!liI,;'lU as the dt'lvelt11Jmetlt or tedmo~Clgies to <Issi~~visioJ,l-in1J~aired. :JPeI:S{lI~];!;. lh:: io:;.:~onhi~oi'p.r'iism h~gh~~gln:Hhe[ii1e1.~Ibii'ul!!ibi~'d pri(j:riti:l:a~ign ,ofooril$ei'vl1li,oi'l bi'do!S'YprojeclS. within the sc ieatific QommLl]1,i~i". Broader ~cient[iir PQ~el1iill~ liontriblltuons need to be considered in .~ddit~i!Jrlto ge,l!1ernleCtl::;;y.::teP.1~1 health, IIl\lrhOlp.s,. other species "It rli:5~ 'i,'I.'Ou~d iiece:ilve Inore,.~;!'eilli.on ifih.e :~allilj[ifi:c<u:ioos, ohbea.· d~lJiI.li::i!l:: were bener pre.sclm::d ~o the publ,ic.
MIVC:HA!~I~D. KASS
R:e'I'~mi!11K@

~3).We knew beHeii IltW"'. LESUIE SAIHi'~JI!nJONIi::S 1lli:«la,~ IRr~f6$Q.r:" Pharma!COIQgjjt P.h,YSI,ol~~nd a :NeUfO$deftce,. $cho.o~ !)i .M@di~lifi~, Me.mb~~. Car@li!l1,a" olumbia, :~C 29208, US:A. E·in<l,il.: 1~slie. C
i~Il.i!5@mllf'!"('~du

11II'5Htulioo.. ~ RlrIrim'l' BU,n.cl\ UiI'l~'!Ier!i'ity

or

'>!J1Jl1ilI

iI!I!~e!'\enc!l$
L II(,Ma(hi~ 11)., Colh~n. (. ~nnwl~~net, .. !l p~u",r ILJ!!:I:i\f!o, ,il/tUJ'IlpllysJD,I; ][17, 4~~ '(2DliJ6.I.

am.

l. C.,M.. t,~ll'k!r\.l'IIt'i!~,

"M1niZ, M m'l!IJlel:l;; llID!lC.WJ6 a

~lttllip~~mstiIil('m!n'.!loli'lllidtl!.WM1!9iJtsi!Ei\iil~I<Q\ 3. 1. Iil:.ln~ml c. Ill. R. 11f~tf!t, 1m 91 260 rnDOO)..

IRre,siPomls,e
~\lIETHANK lONES IFOR HIER UTIER., WHICH

cffers the QPp(m~ll~i~y to d iscuss d1e :Ni~fe~y of

trnflscmnnal ~m13.gmledc:Sl~Imtk'ltkm (TMS) and the ethia; 'or[''},~~s re~earc:l~ilm ~nnmllm::i ;HliC.~ to' Oilddr~:.s (,'!)]1rltn!.m :rw!;':jltd~!X~ !dX1UI [IS applic~ ~Ioll in ljl€'~llilhys;~lbj~(1fs_R.e;.~c'$iI·chOI1I~UffilHIj] :fiubjec~::sshould ~.ltdeed.be done with unnos:

1.

)@@'

hLI~cl1lMIIiIMi:IlroD.~I.~w'f'ublI{,!t[~mlZO!I'Goort2'OO3N,RJ);:~l1DDlphliM.r.lm.

.,ll\1!uiol'lto the pW{:I;;"Clkm Of!i~~r<!fI~~ipmlTs.

TheEth hc:s of Tr,ans.c Ira 111 I i81~

M agln eti~lc:St'iinlull;81t,ilon
WHENI SCIENCE PUBLISHES R'ESEAR(H IUS,!NG ~l!:'. !thy hmlJ<lriIi .\)\.bj!.'lC't'j.. one assumes there .
was, m:inlm3.1 ri S;k3'~.d/or viml clink;:!! value,

i
=E

. Thi!'i dees not "P'PWV-IO be the

b)'1

Thetrn.-stl'll$un !hl.;;do!'SQlat~iP;'IIlrrerhlJ~l<l[ C~.!2 h~'X':S [o~e iI] j udgrn.ent!l of faiirness, <IIncli :;! itm~re~l afeinl\,;;ro~li~:q~" bullihG,)' larg.ely vall,iii d aued whm. 'W,Ii:: 1IIu,I")<ldy suspeoteJI.. iil: .,~... IExp,~dme""till.1 :mbjcCb; f\,"'l'eived r(:'pcLiii.ve ~ tmllscmlliaJ :liim~gml.el.ic:j.tim~I~"lioJm (rTMSj :t~)r ; '~ lS n~il' prod~iOG:"[;'Urppif:'iC&,,~rnl ~o o,r~cti'll.ity t~ 5 tlu::st~.rml,la~ed brain [(;e'JLon:"U.e 11:]"NJS.sel.l.or;:r~
,~

i rrom~d certex,"
i

reciproc .... H~ifiLless. y d~:m~plcinghe right pre1 b t

RepoiftS" .3 No,,~. 0'('J6. p, 829'). 2

In. Knoch

COl se

IU\rt~.le\'\'{lrl

:'md colleagues ("Dhu~nish~ng

~e~r~

.~ ,U€-d ,fIn dG.{;tric ~mu;:~sl ~nn

pf]w{;rt~I'~ ,e!lO~lghIQ

Aim,in!.e'rl'~liomd\'!'O rkshop en tile !mfely of nvtS held <'lit the NaH{;Irm~ hmitwl's Qn-h:.,lt~l ~n June ~Q96r;g,ru::luded ~hat rhe risks of ~i~,.g:~\;:1Pul~.,;:. ~~.wcJ·l~'U~e, ~Itd :.>ll1wj"C'PC'[is ~~~'ie T~I,ISI S; I Hz] are mi rililrnl<llf<n' jW[pLlhlliol1s i in huml.ult><, k~vwn ri ~k!'l !\'I1'!.g~lh)!ll h(,';fid~.ch~!O with()~I~ ~~'rt,ln p~dhj,pl'lSi.ng~tlnd~tiolW:, P!Uto, more nlvdy, seizures or psychcsis (.i). vided that <Ippropl'iOlte safety guidelines <lind Lnng-tenn OCC~l:~lchange!; and Sldl:re[po:rt:oo: ~):t!e('..aul~,cms, a:re i'ollo-wed U). :h.l OLrlll" smdiy, we syn'lipwlTlillin het!l.hhy ~ubjecl~ ~l~1.\le been l.lO~ O1ppUeod ~~O'\I'. ~-H ziiepe(il~U\'llt 1M S' 0( rTlIlt s) ill s;1.ricl~J(lh(!l:el1:;;c. 1:0 l he rcc mllu:lm£~ldcd sludl:h~t!"and rTMS eonrisues (O be used for sl:ud:~esboth fuse i1lm :lril1lS~lIld fi'i volous (jus; gll.ide~ ..nes j i which have been endorsed cltcC:k~hcl~lerfLmQ:'~ ). b)' the ]Il~cml:ll~h:m~~ oden~Di(lllfor Clinical F 1'11.(;': u~ohTNIS (Jill h1.;"1lJrhy subji;x:l:;; dees N~'!Uroj'lhy$i!IJkJg.y~ll. net meet the defiuitxm of "rrrinimal risk" ln piflledi~po.~ed pmie.~I.s,e.g,.. these on certil; CfR ~c~ i{lll "l6..~ risks.; '"l'liOl :y'~.~Il;'ir mal'! m~(.Hl·<ltions~)r:vid'l underlyimiS n~u!\lp:i;y01: \ .... t:h,,1n Illose ... e~rlCO[I,I1iII;';re:d in ,dla~~.y I ife"'). ehiatric QOl'lld.it.Lons.IJ\lcre is ,J 1;<I1.'eposgib.il~.l)r 1/IlVe knm~ that healthy s~lbj eels J(l~'~I"bik for serious adverse ~lfeC'I.~" most l'i(i'I<i~bl.Y'i ::iIeii res or psyebosi si n their "dai l:y li:re ,," se izure or the induction of l'1~yr:;:ho;tic symplJU '~li/l'~t,\1{f.: d(m't k:uu.w.:is wh;.:u the residual m:nms (3' ~.,H(J\\I~'\'¢r. th~s¢ blWVG Ul~'W QlIX'~u·:1Cd c'II'ef.:ls of~h:13 <liCl~V i~y-swampi,:lag ~s,mlliilm i of with slow rTMS illil,h~'lhhys11bjecl:s. In add i~i!i.l~l.ill(lill (]~Ir stttdic;)._~.e<lch ~·~.bj r~rl]CieC'~ eh~c[ffi"'ic;)l~ clllrrom 1m;,:.The Re"POf[ d~mO!ldi::;rlJLr~all o\(,livily~br 7 D1~,irl. Arlinl,~1 rTMS research tw·~thmrere."'p0:ii.Wlre m\ ~11i E.D50 dnJg toxicity ~LUd~,cs} sIIlo\w dl.M !Uiiyl~til,gMuJ.iiOOI te,g,. receptor ~e\'e~s~ is :~.~.odiil~.ed. rTlvlS For

1663

lEITERS
pared vo~untm-Jly a:lmd on tl~.c basis or~he p~i-

:-oinnohU li'e:~ii!qi!l!~Il~onn,nl.o~. experii The menta I ~.lj!U1.l1!lC of the .imended procedine W<I.<S. made elenr ill 1~leUUtNe,tOlnlqj .'IU pUl'ili:c.ipmns were fu ny inl:Orlll1.ed of .illY r-e,lSo!il'<lhiyrore!ee~ ;:ilil~ris~s llrdi~mlii!rts~l!1d ;)~Il [1'1("at;~ll1~;;H r
they \vQWI.Id not de'~ve

7. It p;m_l!~HiIIQ~~. H.~, ~.M.~~~ml~Il]~,]. ~L!)t'!lwel~ ,Ei. ~. !\!!ii,H~a!rqJTtO&{flHri~' MtJ--!li!1l\rff

S!im!l'/t:ii!i~ (Amllid Ne$$j. I;Ondo~'. ~OO]J-

S. M. s:.. Sli!~m. A.IP.:!milIl.fu!lliM.ln ~MrJr,oNhia: tJ'tfo!~ !fh! 1S.l'tIM~ In Rl!tMl(1i; M1<I~MrlFDI'"¥.J!.llles, IBrli, (Cidcrd lInf ....f!~ O~ord\ lIK. 2OOfr).PF!. 2101,..,2:t1.

'Mel1l1ilil'l~~(hes 'Ilersorgu n gSlemnllll, liil m. IDr; Lii~~, nl'r. l~l1!dli!r. lH~r@lIW@!I:n, 49:15:5 flHJn~cer, 6i!rman.~. ]ln~mUlf'6f (ilil1i(ol~(helliliW'),' iOO i.a'bQ!'~ikiorf Me~lcIl~e,~liPh<!\1<!!! iiiiUJii;~~IjWi"e!l it)< o~ MID rlmi!', 4,B:1~'9' MlQns'er,(ie~I'Il~It)'+
Labc!I<IllDrlul'I'I::mJ:lditi Rie're~etn{JeS; 1. iiL E:. (j~f!f!J1 t'l ai.• NtJ!ltl~444,:~30(2(f06). 2. e, M.. Lambe". (. ~. M!ll~'f. W",r,l!fe 441l, .215 f2(l11D61, ~, [l. f. HlmWiI'i, rhe l'.JodJ'li~'H of Mal'll olldE~; I:iiOIlr !i{M.rJP"fMia ~pe;d MJmtmfl:r liBa~liIif1Il fr~ [andll:1l,

Geril!l,lIn)".

th~irp.at"!it,;

h~~tiOil j i,~Thre ~~:udy. lli~i!~t,;

:tilly

dilre(;'~benefit

fr<OJn

1~.dGcl

IlJot:ilJc<lltOn of the poss!lbility ofselzm.'CS. CQlltrnry 10 the opink1!un.~;.;pre.<;St.-:-,d by Jones, \'\,'e fifmlyboe:l~eve dlitf Ihe fimJi ug."iofonr .~~Id'y

NI1:81 nde:lrtiha I, ,alndl HUDlliilln GllnOme:S

IC 01111 pa riing
THE lUCENT SEQUENON'G, OF SUBSlANI~Al
parts of N~'OlI:noolfllll.,.1D.NA. ["Seqw::::U1u;~ng and ml!'lly~ils ofNI;'i~mler1lhill g~no;U'J.~ic NA,," 1. P; D Noonan ewl.• Rel'>~IT!'hAmlt;tcJ~ ••~7 NCi~,~006. 2 p" .~]I]3t 'r was. preceded by releases of
dr;il:b of the (:.li'rlimpll'nzee ami ~:mlf]lm~. M:OI,lOtl:aleii

~)rfwidc ~lO\fl:.iilm:sigl;lT5hn'O~'JIl,Crolt: of lhc righi. cl.orSCl~ate'b111p:rrefmnt;11 eerie: x mtrrC") ill the eenaol of se 111:.c.eb1i~';iI'e:dnmoti\'es iI~ tmmd overridi~.g~ol'!:mnic l~Ul~rwQion$.SM(;hnm.J~nyfi may 11l<l'\icp:wfmnwllp'IfII&c<n ieus lor a v:.uj:e~yofueuK'rp:-.)'chimric rondidgra",. f;llf ~,\<l,mpl:e..im.::re.~'ling[~'nc :~e'lelor3ct.iV.uty in the righl DLPFC ~~l~!:r:r [(n!il0~~ [he inU)ilory l'ontm I of 'prep potent, impuhJ.ive responses and t]u.er-e~.bre·

~. if. ( £~n. iii, [riM, i:i[i;l~tlil:Qll·ruldiru.i!£.ujm!; E~~Ml.H¥iy Atm 70, J45 {~~~,
~L ~.HLlFSI.~~$(i178 .. 56·tzOOn, 6. ~'.!i)_ira. il'rSilrit.lfH~f~ of eMrm !JDJrtWfl1 ~MiJrfiJ)'i tailOOl1, 190n ..

2ooU ..

in2005 aM 200]. r~~ecHv·dy. Gr.0~!1 (II. ,t I ~ fJf e:l!.pet:·t :re.::.:oV'el:')' of the complete Neanderthal ERREN HAL APIPEAR 10 rlAVE NOil'ECHNKAIL !;~n(lml..'l l,lvi(hi!1 tlil~ !1I;;X~2: Yl;;ars•.'I.\ihkh. H: objf.'e~ 10m; i.O Q1U· ~uJ(ly Imlr!lthe.1" pm tanh a .. lsilopedt'i'i I] <lll,t}w com·p.ua sam. ·of <III. '~l\r:U'e~ ;:enenll rhi]o~Clp'hi.c~ll(j!b~,ect iO[ll:O wheleg~nelmc,." to ex9b'lrThine ~~u:r~lie basis of rUl~l,e.~ genmne anaiy.-;is. The'), would Ill'~fer t:il!U we the
nic'.!§lrd 1.0 ~m~!1Jy~vol~l~ionili.ryqr,n£5ti.(ul,~. Lambert f!~dMinlii..1J' .~!;Iggest~"d t Imt m:u:ti)7::hlg (

les,pons:e

dimi!l1itoih
SUdl

~l'\,c!:;:.&~V~

r~s.k-!'fiIk~n~

I;Iehllvin!'

i:~ li.omd differences between the s:.pecies.\1i,lith

dUlerenc:es between Neandlmth3Iar.r,dlliullnan brnhw '.\I()iU Id be of grem il'l'te:m.::!'>1. i~iov.~\'eil:.'lIthDWJ~~1uch t'{)llll)arisons <lireof s inreresr, it is ~lOi~dle; ~UI:~ic ~eoome PUl ~mhefhe t dYlll:!Lmic PJ.\i;)~:COJ:m:~:llat determinesme p~ellio~

j:liltients.\li,·i,tmr, inm;Pl~lse corlll:ml disorders,

U' go.

search for rhe molecu hnt has is of hUlml.anspecific traits by fOC'~lSIlg UI~ pa rtie ular i

en i!l'!i,<;;rvI;11li'l)l1 mi.~H.I~-:,·e.",~unp~!.i:. pruvG' useful in cllnical PQ~)~ll.l!t~on.s\\'iu:hl dl1:ll.gor nousubstance adc!i.ctiOlllS ,[e..g., plLliI.ologi.ca~gm1ilJ~

d ars~l'i of g~:!Il~~~ u lut!;;:d ttl cu~tr~lbl~t~ sp~c 'I~ I

SOI~.l~io~ogi.c.<I:~ b dil~i:'rom;e berween il~:u:~laus

'!lnd (Jtl~er spec ies..\~'e :~1dtosee rhe ad\"l!I~m.g:e l

b.lilng,~.]Il\.\fhich

impairmeuts

of decision-

~mlli~]gseem to teflect a breakdown of tit esc contrOlI processes ~4).. Preiimi:Dl<1lryfindin~s. iiii eoea i~l!,1'addi"t s 1n."I~<'l~ that h~ghmfroqU:(;lm~y rTMS '[lJ lIW~~l~ DLPFC'. \~~~ .~~ ieh d'!(l'u~lll ~IO inerease cortical e,';,:cilnb!i~i!:r in the ulr:geted ~'jrnin regiO:n.rer:Ii.ices cni'!,ling ,t5). Thee~:~.iC'<I~. p1.i.n~~.pl.es iscUSised i:rm d rel<liii.on. to TMS re~emc1m.ill h'l.l~HHl subjectswere IIliiti<lUy articulated by Green et al. (61. and subseq ~H;:!~~ diii.'CU$sirHl's ~Ild updates have <lppel'lred ( 7. m.Y!\'e beli.eve that OlU 5lwLCty fol~
!~J,,;''(ldIhcl'j.(; <If'~k~ul!;lit('dhig~~
!>Hl!ldfll'.th.

;;lind

~hus dis.agJrt~ with Jgn~:s'~ imp]~,t:lllio:~:s O1,btlu~ the, a]lpll',opriatencss or our cXlPcr.~lnent
D.All1!lHA I(N:OCH,.:~ A!LVAJI1!O P,A;S!CUAL-L.E.o

NI~.2; ERiNSiIi FEH R.J

i!~Slm~iJI!1!llor ffillllplrinl Iill!s@ardl rm,j~cDliclI:rnic~, UniN~rsiiy IIf Zurich,Bmllilli~I!p!ltr.m.!'! :l0, 8OCH:iZuriuli. ·5if1il1l~r~iliiljd. ~ynl,'ip~ U, 4). ASI·~il1t: ~lre'~fc)(,,~rjmi(llt;; ~(iMler f!r)l' No njm<la~i\'-e Bn~~ S1.1mtl:la11~, H!)l"IIlIr.a Me~r(;~~ ~ngem:mli.<c DN!A. or genee1>i:pressilJOI] llre]a~,ed to Smoo1 ~:IlId ~~h 15roil8!1i)!)0!«!i~~ M~i:lic;ai Cen~l!r. i!~:on, lipid bi!tJCh,,;:~~.~b;t~y~d. ~~l\)'i.ib~l'l~!ir!l~ ~ co~dcl b~ !WI Olnl'J. lIS;.A. cam,e(]!. (Iut

whole-geuome ("(]f[lf]parisom. The re"SOJriL such candidate ge~e ,::Jtl'<.m~J!!.:i:!'!s\ ..'ere used in the past "'11';,"" dlue te IJ1.; :1 ck of g("!i'D.olmlk: 1'!I~1~ ~~1.",,1Il11e· a s.cqUGncing l}il~~of'an ('I~I!li"!u. S~]if;l~l ~ampll;,~ il~llUdc the c<lterpil:~Hr ami the' t~dpole. 'VVhi:C:~l sh,l:re ofn~jllhipl.e ~:eIliO:llne!l. ill.lciliding human and \.~ gcntl:n1~s \vith 1·~",b~Hr.;rflyal1d ft"('lOJ, ~srec" til imp.~t!J'I;le~.h!;~s~"lnt)\i~d Ih~ (lb$I~1)cl~. d~l t~Vf':lly.but '~~~.1ch Il.l<1.ve "'ery di.~!e.wellit: roteomes i p i'Lo~kiiO\V ill ~!(lv~n'iC~\.\ihk:h gel](;S 0.1 "I~:~elf nmkung them inlu very dilfer.ent o:rgm;aisms .. ~uncli..ollilll elements have c~~.,,:rlg~du hmll<'lll i Thus. Il.~~her ~h.,np:::ri'brmi~1g~~IfI:[Ufg~edom- evulmillim.h ~herefore seems ~~.~)nl'iigmted c m parlsens afsi7.;abl.c genome!>. we stlggC:51lh.u it gltC'!')s dlat mUl'l~tiQtl:siln. lipid genes .• h) (~iIC Illi.gh~:be more ~l:seft~'to ..~dre.~!ii.~'h questiou I is Ilu: .mthors"example .• nre re~p'on!iiib.~elu:w u&r.n!;! a 51:mn<.l:~1JI"dYTKJt[le5 is-,(~r~veilJ ~ppl'tl~d]. Ifum:tion<ll di.fferences b€:l'~weel,l limman.md h N~und~iI'lln~~ bmh'iI $. ~. chf~i1Ige!l.lm, g;i.,m~ ~gt_nr i al1t~SMdlfil.l'l;l[lillt:iI:1:[!iglin.roo lI\8' ··nu lit iJ ill:.uti on" hYI)othcsi!> .. wl1.idl ho;lds l11~MI~.ekiey mWI:n!l~ l:at]ug n'll mct~looli:snl dlo mrm.tll"ib~l:l:eto ~~u~mrl1l.~ don...dm~diff~r~imti<lte\II." fltlm N!;l.md~r~h;lb; :.;p""cifk u,}hs .• ~ljjV~.oIe-gt3m)mlOi ;lpp~,!p~!ch wi~] ~1I1(jgrem llL~"~me :ij,l t~U': ~I1I.es ~ ooding for lj]ilO~ el],iej:cil;lt~y dI.elcct. itll:ll :sigl1l:llI.3S \"It:]II.~s ;El1.11h.e t~i~lS reo~l!~',ni fin met<l.ooliMlt ~up,luni.il'Ulhlr. o!l(!r gene !!:.lhat l!'l.e~m:~ol:'s' "I ipid-centrk" those re,gu 1~.·I·insI:he p·lms:p.~.(lii.pids in bm i~l <IPJJ:ltO<:'Lclil,w(JuM Mli s.~. i

or~~ .!.ppJiOflC:~ over unblased is;

I.~

BDWARD' M. I!UJBIN AN~)1AMESP. NIOONAN

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'V~can list IlUlny rc:w~rC:h.I:o·Jh.cdsid(1 accOiIm]pii:shm!l:l~r$,. A discOVfU-Y iu our chernLo;;uy labs by Ri~;J~~dSi lverman ~~dt'O Ith~ d~g L)fll,ic a, l,i(.."ell1~ed 10 Pfizer. whic:hi has proved ~ n,
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rhe research un~v~~itie:s.

V;(I:jtech Ir:Il~""I1lHIlIVI' P'iliVe~Dro:l:dI., Scott IE. Milb~, M;~rQ~I~a'iol' KUU,!lIII,. Mil~alifl D!alillda. ¥:v~$IB,<I$$iH~ Geol~,g:e l[ll•.We~ibl.elill
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r1!U[l1iI !>lril;'t1!

~nm~l m~C.~lnDc<'!ll.

objoc~.ivity. Th,i~ ~~e;;njllb.scs,.o;;ion w:~d'l(l~jet~ iV:Uly e au ~d d iSRI:rt . <IS \.\ie'U <IS dmil)~. be p~.einomI ~~~ b()in.~ ~x~mi~lI~~ T~t~~1.g Le>C:i!tiQ~ llI~iffe!i~l1Ilce:s, in liIeol,d' S;hil:pi~!O. ~hrei"ol,~ists., ~ ~Io!ch as mI1Il~1hill,lg-:ij,ltellige~rlce, tor ex- WiUia m B~al~y,ill]ued !or "'~efe!bra~ to (aUzaiion (¥r ffiootallwutiol1ls. ~
ilm~.l.piil;.1~l5~.:gg~sl~ t:hf!~ there is a t.'ID,lgible: e:n:lhy to 'e;x.• ,ui~:e, sometimes u
.1

debal:ahk pl'O'POOidm~, I The F:reJ1(hp:ii,ychokl£'U~~ A~fred Binet did mere tha!lJ any ot!mr inloijividWl~1to ~~pp~y osip tivi s m's dliC!l[rms ~o 'illiles,tlldy onnleUi,geIlctl. gmund. The!)' believed thilt ~ more nrtk'l~,~1 Bi~~t. with hL cu~le·~gucl1u~odarc' S~.tlI'!(jlll!. s and j~'St St)Ci~1erder c:otlld be l'lll:>tn~c~~d 01':1 d~vL~6'd 'the m(l~t c(ll,nprellejlS~Ve igl~l;lm,~~lloo thebasis ofmell'i~ or: to lISC' a 1ll0We hislCU"iiJ:::,dly test ro that dille. Yet. OIS Carson i n:si,gIHaeeurare rerm, "talents," To 11dIYlill~k:~, ther.P.il vl::~i,'I;}i'I :tlf el(r~,~5Sl~,:US.view .•Condmac. the tesl. ( .~ (05) '\V<lS Ileve'rr:lme~t 'HH! MeaSb.iffi 1111 M,eril Hel,\.'~t~u~. Rm!~.m, and other to g~:n~e:nHe.iJ s.~ng~:e number to Talents_ Im.ell:lgence. describe an imliv~:(]nals, melll;3i1 ~)h~Io!'li1ph~rs S.y:5.tellldili cally 11nr:1I fI eql Lbll ill V III lh e acumen, However, for reasous tl'lfl~yz()d 0 broad !llf.roy of huFre:ncll and AIli:!Hican manaptitudes, The:ur (ii5CUS~ ,that <til: not d!ear. OOI,W~il,l 1905 Republlcs,17S0-19.r:j[] sionswem impnn{i.m. for '['\;~'O and I 911 Bitten rC'lri:!\t':{i1 the ~f;;SL: b yJ.rJ!m Ci!I'J'SrlJI'I reasons" First, a]t],lOugh they by redueing the les~ ndminisdid m.m 9,lw~fS 1!l.£:u:ee on the pl1rti:l!~,~us. they tel!1i,ded~('I P rQrl·ce1,Q n U rl,iversit'l( Press. Pniru:e!'ol'l, NJ, ;l(I[lJ. ~2' pr. S39.50, £25,95. IS8 N 9780591 Olij 71 50.
tr<l~or's d isercrlen amount (a eerta in

i§lsii lUt iO'!l(t~ exigencies, l hL;,i r-rc!l~~, Ann)' (lUdikJ~ ihe Am('Jiem'am.mlluy) never ald(l:pwd and Ih,lfI,S; neverV<llid.~lred the use of .unteHi-

gence

WSb;

<lindthe presence of fully tune-

lion2l~ :>o:.,t~ng mechanisms

.~nfrnnce (l!I

1ii,llio[)J;;11 s1ih'itelif1l (If :'<i;!COnd'I[Y sC~lo(1,ls .md ani-

versities I. A"[I~e.f:LC'ans. 0111 the other hmul COlli" f'rulu~"d di n€lr~!liltd rc I;J1 m~I!Ui!.C~l':, ~!1dw~ro 1~1,e.relio:remO.re willing to cOIJ(ler limeUecmEll1 ~mJ [)rtli.Wi~a1 \r'J 1~le iim'remg.~nc!; tt'~,t, Ah:~ n!~ ... i.ffiI,~,port'1Il1; wereanxjeties over the lincl,'e<ls:ing
ilet,crogene:ily of' Alm~l"iCiLI1 sod,ely 1IIldb.~C nee(l. e.... a'e:'iM:d by .::lom~. ~() create imdt~c~~ p m'll andrrm;:ifui hkrorclrnics tu nma!nJai~l,social erder, Even sa, Carscu concludes thatAmericans l1fu\'~ n~v~'1" f~ml~yaccepted imtic~UgC!lilm (;;IS meil~n~r,~d ~~a!tlill\vlydes]g!l(ld I~~sl a."I~e by oflly c',riterioll afmerit A,tlhough Carsen tires nO[!~l~~t xplie .• ~ this m11biv,lh;iiJiLCtl. he eore n .reedy notes it 11>numn~cS1. ill~ C unen l dc'rnltc~, over the Sc'hClI'~"lic Apli.l:i.JldIil Te:.'!, T1re J\ I(msnre ojMe'j'il offer.; oj, useful s~w~,e8 !Vl:oi;~m bQakso by Sh.:ph~'1} J<1Y Ga~l~d~J) .llld z to 9 Steven .PilIlkier (21', One HiI,ighl wish th.u, ('[Ir:sm ::i h;;~d de'rml'l;xi,ie.'i:;; ~~m:m~'lmt~l '~l~~I :)Ih 'c~ru~.U!.ry~ end more to lite 2:0LI1I-t"{;pec:ially m po:JliI.ica:J1 iii:
u

of s,~lbjecli\,'ilY\"'I~

d(:fi~l~'91(:nl~~1!Ir~, a~lu'll.h'3.1l;:l1il t pheucmenea dml could be

el1JII,101ncedby variou s nreans.

im;:v~l:ab~!c:) and 1)~f1ci.fI,gglX:.l~D:cr

~mph<l.s.i~ 01.1 !'he in struL~.lean;11

outcomes (l1'ne test:" <Ib:ilUty to

in,d~lding edu~.ui.l'ln, :S~~(l'nd.

as Cm~O!lllnetes, \'I.'~O undertook

p:rocb;''I;!;bloc~!t.; tlni mlhti dual

[If

::!

ge"!,!c~.Assocg:f~~~d mo.s~ dose"ly wilh the'

on an ~,~tell~gence cantiauum], Furthel: revi~iurJ,:;', ,~ndudil1ig illtn~dm:lioil of ~I'I "intelliin~t':IMgCll('f) Itl~lC tena lllal would U'l1:pla.oe ta.l~ gence qltoti.eni,~' ·by StalfDfo:ru Uni,\.'C]'sity psyellibs by [he mid~!'9IIln, Gfli.~l.ry). C c~~.o.!o'g~s;1 Lm""~s Tcm~i2iIn inn,~.9 ~.fl. pmduceda . ]n I'l erh<Lps.~!h.e oook. ~ stmngeSA ject~,ol'I. ~est l1imaa nV,~]Olle(]] ~mell:~.!i!em:eas bio]ogic::dll)' e C~ffiorl(.l)l4u,i!l~" bo'w pffiih\lL~U'1I U"!l~:!>forlfm~,r,~ d~N.,:!:~ni~cd, ullic!ary. hL;lr~dirillry. limd r~>icd :s, the SI.lldy <'Indl,perClepl'ii,Qi.lrn,~f hlJm~n i~.ne:Ui~ Lhroughmu ;3~1 in:c:livi.d[l,~1 [lei feliml.~f:. o
C<lil1\.CKmI olflC:n>inlri,!,HI i!lg ~};plf!mH iUIl~ fOf M'/~IyFn~rlee ,md Americ~~re..lcred di Il"erernly Ittl " inti.':Uugcncc tc'sI.S, The ~~n:nch Ill~c:ed I.(;s!l'>

i~: as prim<lIt1lyphi~Qs(lop.he,]'s w Ih~ em1y inve~lagt1:ti()n.~

:sdtmtis~ Chm:ks, Murray's conl~11tioU:; as,<;er;" liolts .]bOLl~ dn: clli:!lmribu~al1i oHmell ig,ence 3). N'OlU::thdc.."i<.~ s.L'1~.olmhl 5I;\.lCllll d~scifJl~ w ]~~ ~~C$ ~~[1.0 C:lrMll~'S aQ;llHIIle~lS. [relf'V'~'Un and emg:tgi:l1g. Si!rll!nmo~s
:1< 2. ).],
@i'j.

~
~ ~ ~ ~
Q

FrelmC'i'l phi~o~op~~er A~L;l:l~~ CtMmt'. po.sit~vis.'I11sought to !I~:fp.ly the s(:ie.lmaifie rn~thQd I!D soci;d ptoblen15. ('onUe ~~ilnselfemphnsiil':ro the ~l,ecd for socifll thCQri~.s to be p.r(:.d]ct~ve., The' ~~yjel!r'e'rlsi:n Ih~ IDeillafim~ntof Iid'ucaliOOi. IFlllrlll'iI:f1i Un~I!\N~Lt~. noo Podim,1.fltWlllgliw\J~. 'Greenv~!L~, S(

i:lfI,l'PO[~<lf'ICi6 a.l'i I,be~'il 1"01.' ,e'll'enJJ re'l SOil s·: s p.I'\O lie s,~i na~ F~l.li~~.~fI]U.I o &I'I'J[I;. 'l: •.g,. d.~[QI:;i;'I!:ld

l,l9\6,B-l:B4, US/I. ~·milil: 5r[ml.J'!Imd'~son @Fumllllnl.l'!w

le;~che:rs '\li.'e'1'e (ImN:i~I ilmg to cede!~ei r.mthority ~o p5ych o~:oglsts: :intell i:~HlI,<!I1 trc~.ds.., W~enc~,l S{;hoIO!LI'l' 'beg.ml ~Q) emph4;l&i i'Jil s l~.bje,cdvhy (1i1l1OSl ~~Dtab~y.:UnlUilion:J ovel." ~igidobj'l'X:I.ivi~y:

Nm 'llbr~ 20Ul}; iiM~~ ~.~I'.~I~!'\lijt S~ei!'17. 2~12{<!OO~}. 3. 1Ft. j, liI~rl'l'l:!lBil1l" MllJn<1)':~' 8el~ C (~ Uir~ I"i\Mt INI!W't'Q!k. 19'9o!11).
~

S. fiflll@(,

Go(ild\ :r~Mi'$.I!'le~.ofM'qn 2., 1!jl96J.

(Monan" 1\Il~'I'Ilrk"

Bt".~wS1atl!' WiliRg!,

CI 8
::t

~
@

I
!;;

l!i

UrlJ,lMCie~':!J!,.'~

36 ~

B·OOKS, ET AL
PUBLIC HEAIL'fH
<l\Mtiy at oldl scientific papers, eoarr deeumeras, alld; books. "fh.e'll he app]ie~ ~lmdem stari s'!i;,~ .al1d kno\~lledge'lbol!l leal! 10 re'!l!calwhm h~lppclk~. The Imfmfu~ e ITec:~", .~e:ujw<uer pipe's of b\:-ea~~~ ~ppflrl.ml a~'b'Lmtas s(lO~;l!I.sdtie~'l I:hi:: t'ilorld (w~r b~gar:l'to· im!~[f!Hth~!'1Iii. Ob~~i'v';:),n[ d oeWI::S and scientists began <Ilt~:blUl sicking ~Ii';:-i .ll1d d(;1'Hh~) lead pip~:s ns 'imriy" ~ thl;l ~. 50s. E.aJr~y :irl'lllh<ll decade, Erasnm,lIs Fe'Il,n.e:l:. 8

of di,u,;-uend iffilf~~l mion by diggi!li,g

Plumllb Crazy

go'!(ermneni~ reporrs,

\Vlitmg"? There was even a (~leory lO bolster their dis~:!:'~.mj: th(1! d.octr.inc of p~olcclive BUi<l humll1ed years ag\Q-lJe!)ol1e h:<l<Jlled powenwhich :~.eld~h.lll o\'er time ~lm;.t lead M:~$UI~!l~~ ~h~ ud~'''P~fudtlll~ ~.1Jl~. w nf ~e .d'~"MJ .. pOl,i'l!1it-dril1 ki Ill!;l;,,,r<l~erconta nmlinllt~d. byk~ad 1)ipes developed <I preiecrive il1i~eTi,or,cO...,:~jtl]g Ihr.!· iJlhibi[ed ~ut~eql~l>'IlnIpl~lkl;ief'Iead, Th.e: ~ \'iI'.<.IH:pipeswas d'n.~· ~.l1)ln source of hm]"Jjilllll~ P~)l 1!1:iOg .Purriu r M ~l(),t iced that ~'lC]~lmp:-.i'-l il1lgesu..,j, lead, In ~. 9!)O. 85% (I!'~,,,~U.S. ci'tLe~ (p]"egn.uu:y~re]!'Jred. seizures and ,emu3) VIiIa:> concept h'<ls merit .un nmny cas,es. but cities moot cemmon i!1ll(lir~h£;i11'l Eng~~Hwl(lmIW.d~lj. made the ·oo:[)!\~e!'!iemmi::;u~keofe;,,:pt;'cting it te IIml lead piJP'i!sand so di.d most mtljor Europeall c]rle;s.. In regions :mpphed w~lb cQr.ro~h'e where the' \N<Illel."chemis~l'Y was lU""ely to diis-- <I]JJl.ly~mlV\ers!llly. The couns cmnpmmded 1~.C w~u~n;.such as Mu~aclm£clls:alld northern selvc ]~cl:!d. cp~tblishcd nul tlte ·'c~tmul~th,l·L:: prehlcm wht~n.JiIJI£iI:~cs ~idool'~'ilaI. c~:!>nDIIli.u:::rs. H d 1l0~ nml'li.dpO! I i~ies.1\[f,l[erespO:IMi.iblc ' E'ellg~<l:nd,wp tv~!~~r eenral !l~d for :!Illy Jl.)or.!!~tlm~!lJ)llkll'l th:;l:! eame h~llI.uied!> of La rnes more lead r fi'ln, .pip~imi1tanc(lby the cuies, ~·~~u,ml(ld~m &1,.:m:d~rd:~ i nllnw. Owr time, th~ !~~~\"e levels {'If Those [I1i3!>s.ive $JmOUlns h~.d mil ~si\fe cen ~tI'l\lie~!:"es: ~:~ It'~idli'idrimrtii'lg \""o:Lt~J'have d~'Cli'i1I;;,d. i E'ngi neers ·h'<l~re~e~r.ll,oo o,w to coah TlIt! G:IW' .t.~"~dipt!Di.~.a;.rr/{W. P Irol t'~.e corrosiveness of water, Wemc~ Trocsken eeacludes lind :~eil.dpipc;s aJr·egl.~lduaHy b~iI]g du:y brm~!;lJu. Hh.le:ss and ffe~ re:ph~oed. ~ml'r\le:>:ken'g tale is. mere q~ICI1iI~ d~a:ll~ to mllllons of ~opl:e ..As .• t poim ofcomj:li<1J'~~ Ih<llm. <lUli linten::.',;tfl;ng~~i~to.ric<ll root·~ nore, Lead \.V;lil(;r pir~'_;;i are ~tilll in :SOl!l. Troeskienla l1.istor.iall :lIthe use, and t1:1el"ieare other sourees .ali U ni \.<eJrsity [If Pil[,..bu!('Jh} notes that 131C~1.91:hi..cc.~.lury l;!,,'Obllil.eJll le.u;i in drinkl~gw'ltr;;r .. i~~.h!ding solder and brass plm]lbitllgI"UXIiLJI.'Cs.. If),r.~n;'I;{~ babywoukl olie:~!:'Ik>c piUs m~dc of Ie. d "lleadJ helps ~:o'91ll!a.rd )lClil!llr hl!.a~~:IiI."lile ilIl.Isroraliotl'll'm ,m ad! in tlile Nmrernber Betweeu 2000!lnd 2005. \\Vashtinga T Lell:d ton .. [)J: .. f'atled., le.Ld.'cr:iISJiin which s pl~~t~r. liO\i,MSwith U11; WOM 19'23 /IjInrim,{j'l G~!lmpMc. Me< ~~ <I :l:el'ie1;hy ~he IN<li~i~I~'a'l CO.iTlJlaIilY. In Lho1..t~a~cl~of Ilu!n.es had '~~C!\lml.;d l.e;Jd In'-ob~ellls" a w'Qm~m. coukl gild tit.,.; ~'1ln~ d~~~C)n~rd by dirYllking mgl;,'!:s.;~.of eill~ts of i1Il11'i'1m ilesitl.md dm:es, one<.ld'· ( II .;.IIld I.e'ld ]11 t~.lfi r d_ri IlkiilliH "",,.!.'ter: ill [he v;t1rst C[I.._';;.es;. :-mc ~o ~ VI'm.er, Gi ",elmthese level:::; ()r'collt.nni:~<Lli(nlI. i~ ~.s. wr~)h;; ~ 1934 bOllk (2) ~~lfll~n i~~y this I~~ omU~tmi;!i!(.[U(l!l!G:\I~I~ wro.;:'j:c~l~'Jtho!>A;l liu~!~ \"'~mkr;r [ililt in) J\!.IM~iilc:I]'LIliICE:ts.and l.mrrtl\J.~ ~~l~liI~u.::e. lik~'F~nlle~·;. ~ ~"r'~.~lyignor~ .. ~~~ in !he e.urLy W(~Olii. In !Jl<'lnywO!ys. h W,lti Troesk~ns ~C{1'mr.nil .In The 611"(11' L(!.t/,t/ r:f!(;JI1.er enl E:rrng~'mld lead w.ue:r pipes. ~rpemed LO Tmeslon sltov.'S d'IiILl Po:rr.riu'Wffi righ~. Armed iIlc.reasein famu 1'I1lonaiMDjI r!lte.~.Aml ~l.U~ I bul'l l ",filth nwr'~claro. epiclenliolol:!-ic'l.~ rnodel~.'lllcl Pipe Dis(l81£T re!'iOIiI~~e!illlca nn ily wi,!h s;t!llistk[l~ Inct~nods tllil.a.tCllIll cOI1i~.ooIWor(:on~ \Vasfuil ngton's. p:roMcm. A ·~vell-.untemlt':d: C~lHllgC .rutes by up 110 25i%. Bi! ~m.y me;iSUJI:e.. t~.lis W'!I!S ibl~ndi:ng fm::l:o['S,. he\.,indic.nes Ponin. TIl,e ~m \ ....llel' lrlel.!l!1!I~,lemn h,ld tlrn.e unde.sil:edlll(1J11Sl1~ O!~~()fthc !lm~1 ~rioU5 c!l'lli!X'm.n~ctl[al dis~$. lruWS of de.nh fmm ed<l.mq:>'.Sia\\ r;:'re tl;.\1'Q to lhr'tlill [IUC:r11!:e' im: rea~~ng Ihe\'~ter's cOFll'Osin'! ness... of ~.:.rs (lf~he fX\"l 2()J Yo(.,'<iIfl;.. YE~ f~wp~opl~ 11~(] hm~s a:;;l~~ght~!1W·Jlt~rswi~:~ grea:((if' t~~ldency Publ k;. uffichds uied (.()e'xP~,!'Ii~l ' .1 IhJ;;~:t~.oo;nl\,\illy O recog~,i7c.ed it :U lh~ l.~I1lC <IIul ~1~~)OSl o one' n to. d il;s;o.i\'c I~;<ld .. w.~lh <~mooe:rm vcr1!:iQ!lYD. ohh.e docl1i.ine Dfj:liFO~ knll\~rs abou[ .11 [lO~\I'. l'r~k~!'! tbm 'Il~tttl <.:Iremil·T'!Y ~.~)~S Tli'oe:s:kcll l~nei:lrlhsllhi.c eaWiSlroll~~CI.J1:!!:itlmg i! ~~elivepfl\"~r. W.~lIY'the eddc:~;ce' "".<'IS igi~lQI"ifX!la:Uild t~le dOC:Im"s' combilIi11tio.[L of hiswric,ll de~~l'i\.'e wo:ffi R!diiili!fC!I1!()~S wal':Jmh,~s wenl ullhecd~" ChhJ!!illICleadl c:.;:poI(focm;.:ing on Emlgl<lnd <Qml jM.3SS<1.Qlimseus} ilnd 1. fit P'rottt Br. MM J. Jl9~::L92~iS )IlL~193U. mod .... lI ~Tiid~miulogy, ~.~e 1 !Uif!.~s'\f'~'iiI'~U~J;!,I!lt~ sure p.mdm;es ~YlrmJJtmtl ... thl.!l;~ .nel1l.Q~ mriqm:. 2. H.l'am1t, ~Mer.!ilI.e(}lld~fJfly4EdrJmpsiai'fl and itscrr~C'ts; ;:Iregmt so d.rol!u~tic aSI:~C: !\l~ddr;;11 fll~IJmdMIl I¥tlll!\l'(11. Mmoodl, tL.ondon193'l~. ', dedit~,l <lss«i~ted with ~i~.olem imd o~her inl~'c:ciihl!1 ~l!wewel' is <lKie<n(!ewri~er in Wi~~iamstPOli!:. PA, USA. d,o~~sdi~!;t$C'~~~~luwr<1~d'I,,;;d ~OC~~1~'(:S !it ~bi.;; E-milil~:H:mll~r@n"sw.ilJr'9l l '

m~;un.la,1 h)lol,ill,.wldch (;311:1 be Il<uc.lnfu~. CY·CIl lUInI very ::ilfliilU I.tllI'IOUIll5. [l'ol.i·liiIt. dusr, and !\Q;i:~.ne the ~UOSt Cu.llllrlrUUl soun.::c~ of Jea.d. ]n.1 m~.l1Iry places, thelead hili. Sl~iIIC;lrJm~ rrom~le 1(!1id~d gasoline th.u was :Wormer~y wid ely used .un. Ii'!! l!:~nm()t)il~s. W.~t..,f. too. C01!1! be ;:i SO~! n;'e: ()f lead, md na-endy :5Q~m:: e it~es~mClll!gii.m\s US. in Bm~il.. France, <nHI Germ''ll)' huve n~ul pJ50blelllill8: wirh high levels of Ie-. d in Iheir a

mllEly. !iCiCI1l.liiS:~:i\know <I ~I,"e211 deal aboul lend, ChlildH::!!l'D am,l :il1lfanl:S, are
\'t!'r'y

on ..dullS ruther I!)i:lill mO\Iiie'~vu~lter3ble c~lm~ d.I"e.l!L And ,}llhmLgh lTh.C problem \~I~tS widespread, il\\';IS ~lot'lhN'<lYS .liSwd.lued with lead pipes. For Icar!l,roQol1ll~l1linate' W~llCIJ".I:~'n.eWh,:r \ needs teeermde ~:e<ld. he factors ~ttolt cause T !,M~I!;;1' to be eerrosivc ,car.! be ~ubtlc"U1d make dru\vil1g U'Lt:ctlM$i)l ~~nk m~r~ difTi~u~t
But :scuenlifi!c "rgulllcnts
Qn~)' p.lI:t]j'

dl1!1(l. [!'IJ aiddhiorn. !llOSlllf]CI.O::ffllt'l~d~-d.ll) RJol1l.IS

,'~!h]l';:r.!lJle this neurodevelop~o

~.'I\pl.~h~ th~rt!lhm: nf ~~r.IY-2mh-0I;;:1l~tl!Fyocis eties toaeeept tthe lead pipe plDb]elm.l. There
were :>~}c:ii,,1faC'~u.!ls as

p'~omi~.em Ne'\ll.' 0 rleans physicu a 1m. lr<u::~dan epidemic

<1,

'.\1'(;111

or Cfl~ ic

and COI1VilJ lsiens

~othe

The ·[irealLead
Pij~(l Uijs.a~~r

Water

Munlci:~ll ities were in deni.[lll about the dangers. Lead pipes

were an 1;:'Xpe:I1LS,lve' unprnve-

city·seltp,m.di'~]g system ofk'~Jd

W~lJiirpipes, The Brilislin physici~{nJ

bll WefolJf1, 1ia1l'sJ\:·.fl'J'l'

NOf=

dJrinkill~ \.~'~Lcr;

t~mln Porritt Inlt!. JUI even mm<! ~"'i,v.r.'epiIliiS tll1cllers~ambg .ofti'll! probli2:ll1Il. Pt!l'h!lI~~ personal Il~j; e1Jl:pe~~elice rm~y :bve piq~l.ed h:~:s i.II1I.~eirest in lead's dfcci'f;, as ~e h\~(,~ came dm~ w~th lead

MIT P're ss, Ca fn b,rpd(J·e, A, M 2000. alB IlP. $19.95. !;. i9:95. IS B N 910026220 '116,'4_

meutthat

bn.)uglu pure \l,,';,1i~r

to homes; how could thal be

"loon .,

~I

t!(j~~.

1669'

ClllMATE

CHAN G E

(arbon emi~$i() ns trading '(tnsi,I,res.n,ot (l1I'I~Y that 'I:a r9lets wI'll be met by a: s pe~ifk da te bill o:lIb~'ros,lersintemgtiolil·~l co-opelr,~ni(lf'l.QIml
,(i'~I,Ii~lQp, (l>rl'~Q.i!Jj,ll(tiv@1i, m

A
g<1 ses,

s. th~ U,'ni~:C'.d, SWL'.~,~n~W£U~l.~Vin}~, I)'.., ~~j\I"I:ilrdd im1'lt~ l~g!!.~]iu~~ln. di:S(:l!~!~)n hOI:!; shi,liedi :i'O~,H the science 1:0 the' p~lhey ~rpiions fot s I~l\viig g em i !'i"'~~ln::;of j earboa dioxide ,(CO,} ami ot:~er gree]1l~.olllse·

n:d(J1~cd lil'l~bc~· h~l'\I~s~ ~.~ O:~C' It'H;miUlil ~:;:]'iS11',~a!J;.-.ed ]!~~~;j~d ~'j:y

S·omJ.~e; 111vura

referred to asu C~~l'i.( , ~.O •. er:s-It~.v(lr gm'efn~ h ment i\ubsii.di~ I!~ )"1 If hig~. eiltm~gh [0 aJlter eonsumer beha vior, a carbon t:o\X would reduce ~Im~$~~ons )r r~Jsft'!g th~ cn"'-eli'l.'~ price b ca rbon-imensi ve ene ~f re:ll,ltive to. ":J1IIOOml,= I:ree seurees, SLJ.'lL!sk~iec~ nUl)! speed develep~,~le.mJtoh~C'i I,jc.titrsel~d lew-C tt!dl~o.hj~es" Sllt.~ :1~~l;IrJket..billloo.'Yt;tl:m~Vilh!ln ! C'Cm1O[j~~

it",. ml CO, !;ln~li~::IiQn~-

hm'\'e..,I:elsewhere to :Ulllflet oe),1I1.md fl;l( I~~mb~r), t~~e eredibil ity of baselines ina,c3,pped and uneapred :;y.,\ilem~, .• !mllhlo: li~lIl,;lig}l<lte efl!~e(IS oJ enhilllce.d bklloSic.a'11 gmwth must ~ad'drri.~.s.~--d(9-10)"
HO~'\leve'f, these pmbj.eL~.]!i>~II.e allll,~ m ~lig~~WI; f f 1), fr[ml~~\lo!tS o:!ml m.eLhodolojg:ies for do~u~.M.~ntiing nh~sizeand W1i~.idn.ly ofc::m'lXm off=sets b'<I'sedi nn l<l~ld-l:1;1tlm:".g;f;lmeg1[ p....etices a:re ava i'~ablc V2, H). roll!owi"g such a nl!!;!~~oo.ology\:\lin IliO~be ~ lri\'i~]~ c~roci se, It wiU itrlvolv~ Ctl<!i.ts t;i1ii):~ win HI]~ll:~o~ hnpiLm.g;tlJ ~~l.~ffi,~t (]!J~t.l;it:;;, Ehltl:h\O <Idvdflli:age ora mm'kcl"ba:sed s:ysrem is that it

or

leclll'lolggjfg~~~.'l reduce (JO;> emissioas ~11 I~e L~!!SI ~ll&I:.tmu;,nv~ndl ~'XP~I;l~ of the pro1'111~ ~:I;)!,vi!lig disdnct mJY<l:!lUl';;lg~:-!0'1. MUNt i.nIilPQr~ \"n~uIdb~mi~imj!;rNd, taut, 11 cap-and-rrade SYSieI1:1. c:'Ou~)led \~~il'~~1 Cuning emissions ofpo~h~t~rlin:s is <l!tllmit~ 3Jdeqtlm'~ elmforeemi;lIR.f!~Ur~,'i; tharenvironled~3fnQ:~as com.pUc<~~ed!JlS, c uning CO-; tim ismenral goals aCI.~lfl~,iy~/Il'{)uldbe ~lch.i~d 'by 3. sions. <1m]transaetien C()S,t~ COI.Il, be a -r<'l(Hor. Nevl:rl lire Iess, due Um~k:d S Ulll;;~\'V.'ls, able ro cert<limt date, Gi\'en~l1e ~lotellni<!lllure~lIhniml,g d.u]1ij,1g~s nd the mgem need 1:0meel specific a redm.:e S\l~fu:wO~ ide f'mJ] iss~lnnsahe,ld of s,oCwed emi!i\iiofl HU]~)L;,Its £i[u:::~ecrt;;l~llly if: a major (4)., U le and ~l J [li)k of the p~.'t)Jccl:edcost using oulh'.<lll&mg.e. A. fiede'm~,,c<lp-:,mdi"lmde system <J IjllHrke~=b;I:s~d eap-and-uade s)I!i.wm I j). co~tld 'b\;li!10tlrporn:t('<d irlln ~isth~g ,L:)mi~"iO:~t~ m~mimui.ttlii\ of 1~;Jdfrom ,g11!;nl~!1~amI ph asetad~ng fr.nrn1ev1orks. and lnal"ket~ SUC~]3::;:the mil 001' ozone~dep let ung chem i~:11~S 'were also Kyoto P,roliOCO]:~ intemi1ti,on<l1 market or subr<l:c.~I,i~ated.b)r!!tillus..s ions. lr.l.di~]g pnJg;l'mns. e 11~~liml.]1 ones like the Regioni!~ G~iIi:'n~mqw On:-.eu:ing emissioas by storing ~Olroolliin Gas,lnidatuv-c, soils, ~bro:l;tS. and Ol~'n.er fo·nm:;. oflrliOinass ill ~:I'litM, ,liM:ll:eis '~ljlOgli.c aoo~qt the loc,f1c the U:L~iled St<IWS has the pot:t:llIti.;II, ]I) I 0 'tu 2'OUAiof u.s., e'[JIii,is:5il(:m~,in 2025 at relat~on <L1ul~Yp'eofCO.,eJ.n~s;siO:mliS ami is sensitive Q!'I!Y to I~t: li[l~fl! hlnrJ~ll nl'CO~" h makessense. l iv~l:y ·h:l\.\t COSt ~see C'l~;)l'n belew ;imid HI bl~ S ~.~.umliernat ional OJlJlorlllni~i,e:531!>o ex,~st ] thereforc.~D desig:Jm. 111 clliMlle po]iqr ~IJa:~ tt!:~ Def~oo%l{di~,m IJQpic.~1 rn.info:n":$t-sis curof all ~)~,.;;jbl""W(ltH~~ m lim~t n!<l~ CO" emissions, T.r.ui.ing o!" emissions aemss l:I~l£e(,~Drs I'el'llly e~.im'lted to cause mO.Jre tlllar:l 7000 ~,u~~,~ of tll(: economy addresses thil> by !l1]Q'\,!,Ii~l! Umll ~netricltJi~"~ I)CIi"yt;"iHofCO~cmL'>l>iC!n!\,. nhe otl:.libout 25% of wor;tdl,:\i:i,de emJ~i~a f:l1!ililters In"JilCh.,~ cm:ool'lc)'i'fi:;e~sITmll l:m:-li~ ~q'LIi\,:'!llel1l nesses ~h~ti.l~ a'bl..- [.o~rnvoi:'r.th~i[' OW~ e-mi~~ si[}:n~ll fr.mu ~bss~I "J.~] burnin.g [(jday~i~ 20125 sions below their alleeation, Ifl:r.a:di,:lmg,were lll1.e pe:rcelll,l:<lge isestimated toO be <libont iU~flrpr~r.l!l!~di:rlH) ;:'!!~ i!lwn~~t:io:~;;tl s.y~~~n~. US, 15% '~~~Ibl~S2;J, U~ijmilg a!lillt~r!:la[i(j'f!!~1 firms and I;,'unslll.me:~~ ~'O~~ldfn~e't ~ c.,.p-muJ-lmd.~ ITmlket Cl1fl1iSf>tOm.la:rgms ilIl reduoed (:o.SI'5 to ,e:ompcnsau! nado:~.s lor s~;owin£! e lore:stm ion d b~' ~W1bs.~~llIl le$."> e,"lP'l'Jl!:1ive C~b ~l1g in.,lhr ,~",a1]l ie. dc\"eI.oph~g L'OUfl~ p \VOllld In~ng s.igll~ :r.CHl~t 3. ~ries •. fOreXptl.11s:ii!i.i(: en'iLi~i.tllu; cw", bk~ck of emi:ssions uuader
O~]

wide cap

emissicns

<In,,,1: tta~llilig of emis-

sicn ~~~ lo\V,Muces

\'VO'U

ld do~Jn(t ..;~um.e.\;l,I'hiIC'

pn)v-ides au') im::emiveIi:Jii! inn(watioo,-which

infO

~mmtranslare

inexpensive

CO., emission
,m'l)'

reduel

ions.\iVhywuu!d\.\

1;\V,rn:t 00 ~('iude

seeree of the economy Ju"Om this;camp!:n ilill'.m. IEll.3hlll~ Ollt:;:\\'idl such h'l[gl;: pOl~~t.~I.?
ReJerences andl
2. Mol" Hofletltl!jof..
]J..

N~ll!S

:1. ~ H. SotnLesifLge~,S!'ifJ),e .3a.4, 12117 UOOf,),

Srifn~Z911 .• 9S:I OOO~. C.~l,YilillJ, ,." OppMl'll.1m~. rum,. (INmg~ 810, 1!1' (~on .

onset

Ed~!AWt1i1r~ D..m!l~~ (!lm~f~ Chf1ngf! ~r,"nbllidg~ ~i'II. Pfes5, Camb IT~, 21OO6t 5. ~r~rAcM,~I~,gi~,~~~r P~~r.am ileparr
S\;iefLo!e

.t,

1'1.1, 5dw!milihutllli'.

r-ofWlgflU: An Jm~~(lJedAmUi\!?etl1 ~UIS, Nil~OIMI. ,iiIl'!Il'edJlo1o~)!' Cg~f!t'iibW~~i~~~~, DC, :il'OCl)l;

".",m.e~"II!IiIi!~g(i~d.i'J!;ORS/ii~~~p;i~~~).pt!I,

':0

in the United St,ues," iBec<lu!:ie in\l~s!mtlilt "''\'l)uid,bl;; fu~~oel.ed to

O.
I

l,lKlmlg;emenL~"~ii11i prt!= Ie
~fV

ing iroopl,l,l'e"Of~bll-ll~Otl~
providing

!\rS;~e,UI,~s; .,nd

IEl'!\I'rro!un@n~l [)~!!J'!se, 2 57 P"r~ lWenlH! '5cullh1. New '(Qflk,. Ni'i l.o0:ilO, USA. '~od'n:J·W'Wioon SdtJooI. and: Di!p.arlm~ml
Ililrin(el.on, N'J1)8544, USA.,

200'1

,2:0125:SlSl'ttiln ~5,on:om

~1'iC~lme l de'!.~J(1~i~~ K t;Qll-

nQl.n~,es (6-.Nl

(Illub lIJrIJiI'.I~~, ooriunt In ~~ .. 13, ]I,,~, ~1!imi~1) tl' (fl~ ,Ei;l5. l~jei9a~~!l7r Pll'llel (jllj Gimtlll' Ch.im~;~; Go.tld .Ptar!:tiu. GuJt!~'.frJ( laoitoil tQi!ld·U~ ,rhtm!Jf!" QN/ ~'lilf ,nlt!i~f1l1J1i!'!or ,Glioml. IEn~~rnPe~liIll S.lr.jU~[li5-, If.l!yamil(, t(a~a.g~Wi!. ,~ap.iln. lOO:3:}; Wl'i'I'j,~c·ngi!Cli,p.l!iI!'.!.or ~~lj]Hibli(/ qpqlulucl[l

(;" IP.lYloo11noo.~" $cl'llNliIlilRlIn" fIk." rro[:l".~l'l)~M:!lOr:~ 'i!M (lim~~~ ~ (J!im~IZ9rn In~~1\J.JI~ f~r IEmrir'Qm:!iI~nl.a~ IftI!:II!:l.IHtI. Bil!1i!m, p.ar.\, SlTafil, .i!Jl!mI\I. 7.. lit illfl!1i~,:5. SdmiYlllmt;lllll M. 'O~!Jeflihejl1llf~ ,I, Blaom.f'ti4L SciL'Jfi'{e :2:~a~11Y63: {;ZOOOl 13" ~,~.a~lmi,~ 1l'1~(:r1m, crra. n" 2'1IS ~2005~. '1 .~, IF: K~p~t~r!!lf (1'"fb:al\lrf 411l'9',1$7 ~'006:1. .~IJI. S, Gibl!l~d tl (1J~ G~~ fI~ ~Ir!. 32, 1.J37(:J'5~:2000h 11. t, Olill'litl!t~!)o ~lnr 5rif!nt!jic fr'rlJif{fg5' ,rJrlfkrmiflt thf Grmu(e fJl1:tJef~ ofS~~e#r~r;afj ;~ f~~1$.7 (~i(~ 1lI~5 ~m~~iWik!'.ID1l11!fiI'Ii, irK .~OO:6);WiM'f,nldl1ioIMJd!JwdW i i~iliWlflrneibiinewalltf;pd'l 12, ~idlli!liJ5 !n~j~Whi',HrJ'ffln.ri~!lf~111'1~ ~t 17M ilJ rhe t.~~rbWllL-on.o.my.' .IIM 1.0 ('«ali', M'iNl5Wr_.ooQ' Vt'Jik G.l'ffn1:!JQ~ G~ Offi:e(;j BinN M .Sl'Oli~9 Calb;:!fI rn f~5 r:rtrt/ SlJoil ~JJolf Rtt!!i<!nB flfJi%l'ofl~frorn It!llld !M

rile.

use.

i:,ff!eirt 61 "lIfboln ,n!l!mlIU, 1I;S., 91~i!tfilhQ!Ji~ gas; lffilf!!~iajwfQr ~;oG4 and 202~!l 01 Ge(lsc~en(6, fri:n(eto!! Unl~~~,~I/I'.

¥li1lh

·Aw:~iqll)r 'fo:[' corresp.oMell(e. E-rna l~; bdlilfll'!eiid'e:s @em...iro:r;une~l~at~l!!ffl?m~lOrg

us

UJ:~, ~!1d·mar'1illgi~mejjt of~~~

i",di

~milll!lM ~~ ~~lJiI~ n~!ue');~:mll]laJ~ed

E~~~lr.un!illhe illh::gr.~y of s:tlc:l~:a system will

iJl'!llllh;d.htfl!lt.

P~9~

~SG per ton DrCOl o(.gr.e;~!'!~;l = 1 ~i!lign'Onl.{t~blLe:liU,

in m<!~, .;1

5i!!!!l'Purti!19O"Ii!1 ~ ~u:rii!l roqll~rc r.igom~J;; n~.iQl~itQ:fI'· "'_',~Ci~~~!!®ig~-glc:glR.gm~!Il!;J[!Ill5tS·a,1t~tl(,.7QmCt ~ ,<lllU.diitill.1g. and res181rd~ ng.. dOlt L.;;akf~,€'(e"g., \l.ih~~

l670

__
I

_._

,_
I_

8lI,OGHEMI

ST:AV

Iha

strlUci!m,e

oHhe·

Ci!J,f@

tha'~ senses amicl respends to! cha J!lges iPi! t~~ cell's ener'9V blllliln(oE1 na,$ been solved.

Qr a ~ IlI'li.jlffliHOffl

P~e]l!

The ~!bjliry of imlaMm~ iaml AM PI{ !.{"pho~~ I pil!Ol"yla,le'downstreamt ,11~gel:pm[e:u.ns, us. in~ ~ng~p~d~lfiIlk:' prQfKlrtkmii \\';or~d\v~dr;:. ~Jte<l~~d ::.I OO-r~~ld by anaehment of p~'gA]:)phate to i~:l> 'et subunit by m;lCIllln.erpl'olei~l ,{J. A key !lIirget lor drug de\'eloPJllemt. i[llhlis area !io'~em:.'lto phUS~~~.lor)lk.te,1\ M PK is t:l:leouJew n~i~1e lfIo~m'flh{l:liph~I~e-"cl.iv,lte,d Thi:ii pPOU::i~.l pliIJu:ein kinase (AMrK) complex, which r'Cl£u~ C()lllStRn:~ly~ rhephospbareis iKwm.d Iy rapbLI~ iates oc:llh.nlnre:1l~~y balance, For example. knyr:cmO'V~d by yel anOlIil.cr cn7._:yn~e. ~hal' ')';0 AMPK iii. s"\'~~c::hed ~J1muscle dmlllJa:g e:tl::: r- AMPK .illllmediJ[ely rerums jo the inactive on else as a :l"i;)suhofincreased ~n(;rg}1d~~u;,~ml &''i.11(:, Bi!m.dilng {dAMP te n~c y~~~Ubl!!1!it ir~hihi.ts ~r.Ggg~rjll1.g r1m,et:!'bo,li,c changes such :M the t1rnis deplmsphorylannn (!,'. 8]1,pl'o\l~dill~ iii S\l;,'HWh :r:~'()ilIl, ~lmi:nlg to C{']i,1Il3111'1"11.ung [lit!!> a:lld, mechanism for CO.n':Ii\,'Crli~.1gthe kinase to its c~~roo~)~d:~ne. AM:PK activ'.uiQ,iJg, clmilmg W~lI- fU::I~"e~:~te clliJl ril1l,g ti mes of tlJi1e~y de fici r, a i~w'exerelse helps to 'IPMICl!N!lgai nst ahes:il), AM~r b.illd~ng alsoiaduees a e~lru~gc struein
,~~'6 dis~r-

ders of energy balance lm2Llare 1.'Ie'K::h~

I'fii;:it"oitY and lyp~ 2 di<,!b~t~l>

~n

hi.Ji'Ql~lijl ,r,;(llupl~ ""i~h pro!cI;:i~,-d~,g:mdirig enzyme$lO g.en.eml.e the more cmllIJP'o<LGt.3ll.1d rig~d 'Core Qhhc c'(~mr!.:x.,Thie C'(lfl;l coatained Ih!~ em ire y ::mbull~.b but oilly the C _;~es,m.i 11:<1:11

diurniiili 11 S of the

cmnpl¢::c Bm:;[el"UlI were then p:I\OgD{l~lmed W

J3 :5\!lmni.I~.whIch were knrnMall;obereq~~i red rOrIUf'milHio\l~of the: a~,y

0: and

lure th<lil: C'IUS!.!S 111fil O!IJddi~liolli.li activation: the the hlrg~l Jor~lIltid.h)bc~ic dnL~:> s~L:chas the ~oI11bi!1At~m~ the I.wa ~n~'Cts; prudtLc1ilS nf b~~!.!~!1!id~.$ .-.~'U,dlbi<!7.ol~db!.~(,U~I!)~~. (I),. O~ >W[ll1-f(dd Olelgxmi:f)i!l (~). Hl!'!iVI.we-r. f!e~t'~~r e:cnre<:~ of AMP appears je occur \1!1ith the ~Ji1.ge I.'726 Qir~lIni:s :i!Sstle, lhv.',1mI~y and Shnpir'Q (J) de:5C~,bec~t.-.1 suueuues ftlf the core relased prol:ein from ~'lUddi!~yeast ((lit even thcr\.M PK ()mnp~ex from fission yeast T~H~ thollgh lite AM:P-bimlil:lg sites are.conserved, smdyprovidcs .~niiig~liS jl1i~oh.if.ldi.ng of AMP L~rgcmdlipmidnlcmnp.I~~cs arc nflcn d~,ffi.l'lllt te ,cl,'yst,d I ize because of lliii:eircomand udencsine tripho~pl.mle 'l ATr~. h~d, w may be v~ry helplid in th!.;: (;!Iesiguefnew dnJ~s. pll~X sh~pe1.l!l:d n~~'(i bi !:ity. ThL;: :M n;).tc~ adM<I.l.nm"\~ ian AMP K regnhlleS t~~e.Jtlybal~ oplied by To~vl'lle'y,JIld Simpoim ""'l1I.'> to tre:-.,U; ih.e O!!!~C~ 'by mn!1l~hl:rinS d)~~!lgt;;s.i~ lh~ c'CUul~r

~ttt~ di'lbel:es.;llliidl,l~is

prCJIreilli. eomplex is also

1l111i\'lkJe care ccmple, x by gellD.etic mElllipu~:J~ thus It¢"~!'I~!;;ch!'!iqlt~S.but s~uit<lbl(,'; (:rysUlI~ ,,\wr(;i olJt<luned only wi[lm the protein from f.iss:i:on yeast and IIUlt \'\1]I.h~:lnHI:~~m1lliUl]!U,13. Tbis is tll'll\1numlll);'. because liU~eis !:.;tltwm,lbom the fi~ ljon J:e!l~t.('I11Zyl~C, .. !I~.I~Ol:ilg~. sequence is il~ mm;e si llIil:ii~'lIf to lh"l of theprCJIreilii. from [YUlddi:JmS ~~t;1 Ih~1l to tl~~ one ~om hu.lll:If'In~

l~ th~if snuctare. thl;i C-oormif!.ilI dom.lif!

of the
"-~e'm~iml~ dnmaim of the r3 :subuni~ ~l.fddl,g areuud .~ll'see the fig.lUJ1e). There' ls mf.:lilim<l:11
COIlt.;"lCl between 0:
~I

or

0:' SIl]:b'~II1I,it has

a eempaer

S.~J:<lpe. with

the

eoncen W~~lion ~ of dl!l: fi ue leol ides AMP ;ntd ATP. A.n inerea se ill A "tIP ~om:eJiltJ:al :tO~,-

in:t1ic.~ling au energy dr:f~cil-~witche.'i the

kina se
Oll
(1Il1.

and ybm exrcnsive cmlililct betwe.elm and..,. Th.cal1ill:ino acids .involved rnn IJ1~:~-yi~~el"'.1~ttoll t~l"!.d tabe sh~d~~ finAMPK re'huivC'S in 011.11 species, wlrie 1~SlI!;1'Ces,ts lhm the siUbullhi!lle'me lions \\1 alse be 5~mil<'lf" 1]1the structuses ~'epo.rled byTQ\~'IiI,'~ey <I][IIdSl!Hlpim. the

COlllIJS.il.l,g

hloat

lilC~~pho,i;p.hiiltc

I~

do\\'m.'!'iremn ta~e~ proteies, dU::lM.:by

\\'O,'I}iS

!ioWiH::i:li il@.

preteias hl"ro:hOied in, ATr~gelle["dlillg p~lh~ ;;iJnd siiviIC;h,il~g olr'~~.~ hl"'l.l~\f'L;:d in ATP CalhS1J:IlIlpliollL Conversely hi.ghATP' COlll.oell~
t nuitl~:s-i

N ~ anlid C -term lnal Bateman domains (domains Po <InJ ,BI fm:-e each other like mirror iIliI~~@"'''t[~'icry!O~(li~.gft)\'\t]il. in the

presence ,ofAN[P

QI"

ATP. a molW"L[i;

ndic.,,'ting '~l1!<il"JY suffic.ie;ncyall:ffilc

~:Iin~~

~c·u~~(If l"~!clli ucleotide n

jJlrevcntAMP frum s\1!li~:chi.D1g en the kinase by

L'JOn'1Jpt:linlg forb;inding £iW"

bound iua very SiD'nlhi.~alliilm.laI1uil.er. bm mdy ilil. dUI1iil~in B. Although

AMPK and itswe.latives lin

nOITlIIiI,l,;ID'IMml~iiallil

spcc.i~ n:~ eomposcd of!:! t:l:Iitl'lIJ.f~k~ S~lbl,lQ1H (,cq o;nrl~"\iO regu~,~lory s~.lmnJits (p mm.d IIThe

crystallized in I:lle presenee of Mg~·. the ATP cemplsx !;lO:!l1imetal, ll'lliI:us agrees "~'~!h fi:!fliIJ.in:lb" [b~i,t TP [)'i!ldiI1~ A rained
110

~~id1,;l(ltid~~binding ~hl\;:s ~r;ek)I;:'<!:t~d iut\\tQ "Br~t1;lm<!~l {~rn~~,iJ!l:J." yf wl~icb is :fullnoo d well

to du:l3atr,;::~~afi dmll<!:in!'<

by two repe<l.I:> of ,<I p".I.N~I.·iI;]of 31ll i~,o <I/G[ds ~oemlf;ld .1 ,tB s :il~Olif (J'J" Thelm,udeol idebfnding ~~liC ~uu;bt:~11 mmle.le.l 011 theM.,l$. of t.he stn!l~mn:: of oHler 13'1I[e1~.'D.ml donmi 11 ;]]1d S The core of~1iIe' comptex •.TtlesirlU::lur,es ®1'Tol'lI1ley an.d Sililpim, (,?) (m the l:m.·:.1~lioJ:hS :lllml:Jtioll!; «;<'1.[1i 11~ lhulml:lll, of s (r~) h~,m d~s4;1asl; lh~l ~ru knowl'l tu ~nt~rflilr;L;\ oonl~ailnllih,~(4erm'ini:!l! d:(lmi1liIilSII)~ tile- O\{~U.o\!ji}{Ind ~ $iuliii,Jl1If15 Sh(lWfl at l~~ t!Jp. @Pl~Ihll ,~rrti~ l'Siulb",lit., The B a I1td "f SliJbUlnit:si'!llteJ~ [ \'\,·:illinn1:u.deoli.de billdung (3. 4). Toofll]ey illud act main'ly viii the cellltral P' slilee<L Bil!lotiimlllll domain~ A (Ieoo edl from ShiiJP~m nr}\V :>hn\i,.t hm!'! the sl!i.'lu!'I!its: b!!le;rrU;~ in•.3'IliI.d. ~,1Q~!!' nuc.leot ides are btl~md tD. the d'ae core ofl~~C aI3"N~omp~ex fm:J:11flss1lr1l1l yea~t,
Tme ilulhQ:r i~in ~ihe'oi'l'isk!n 01 'oo~et'IJlilr Phy.ii.Q~,a,~y. (!!!III!'!1iu~ ILirl! Sci~rHil$, l!J.nive~zy ~l D\lnd~~, Ii'ltil'llilll'e
!DIm :!'IEH, t:. E;-m!lil: d.g.h;ud~@d~nd@Ii!'.iI(.IJI~ u

did net wCJjtl.i.re JVlg2~ ,(3). h 11180 hel'p.~ to

~~ pl<:1ln htmr Azo,,1P t.m ,e:mnpete ... t'!1f~c~ilw:~yfor bindi[l]!:l, wuth AlP. e\,'e'l'i tllough the 10000l:te r .i s, U'iI,or1l.na]I~Yprese'nl[;;;Jl UlO~ro:M
~idgh>ef cn~C'.emr<:!liol1s: Mn;;ot AT.P ill lhe cell, is pre'se nt as, '~h~ M.~ATp1'c('1mp~~x.\ilihere~S> t.he COIH:CIIU'I!U,UOu:ll of free' AT Pis w.1.t~dl rmvcr a~d closC:;.!i to t'l:lai of A Mr. Bat(!:n~<ll'l domDli.n ,Ao ''''fL'). lJIiii" ooC::~1.pie(ll>y nudootide. d~.pite

CSSJ! ~p'i!i\1 (652) ~i'idB {CBs3 3Jr.dCBS·~) f.g(e eii~1'i G)tJ:ler[i,~~ m~irglr i I'IH!Q@s;. with dClmai 11 B t~tJill! IrCllllt i 11 th~$ 'fiew. A s,lngki AM.P I'll Cl'k!n1lei~ Ibound inl dOFIII<lilii It with 1hf1lte9i[J~q\l'l!ly rlld'rged ~hosll'i'li3lt'(' glroup (~dCJx.yg.Nlail,Oljlls) inl.erilcf[n ~ ml~ ;;II pO$i~i1i.~:lyd1!arged $[cIe (o1l;io ,(In (654. The "ppro(l:ij m~lt(,>~os~'t~()f1!S lWf.! @Quivillel1!t ~@sitijllt (J'~ ~ide d1a~ inhuman CBS] and CI:!S:t1.a al~o if'ld[Latfd;t~' eseooliif' flot n.s. re wnser\ld inlhe yea,st enzymes.

hhil, AMP(Uld AlP

("Orti.."eIlUl1-

1671

PERSPECTIVES
d~)n5.duril1g c.ry.$u~]!.i~ti(m. Tht~was unexthtiidea th~[ d(l~'lfdm!, A ahl'"l binds: AMP in dru.gS.~il1l:l~ll aitrear 1'!1t;:!'l[ t'l f rhe epkl~ rI'llk l)eCrett. becausethe isollmed Biilierrullldom.,lins, humans. Recently, mY~<Ibonu.ory has pruof obe's.ily,md d iabetes, rmm~11.cma:lrn~m!llill:1rn lsubull] its lililflid £\\1'0 mo]~ vided evidence [or a meeban ism ofactivaRt'rellmces eeules (If AMP ur ATr (3}. This i's~ikdy m be ti g.~ () r the human enzy me by A MP ~f J ]1 :t [).G. ~~dte,ArlIW. Rev. ~~~t r~~ 4", 1850 :I, .~'IJTh~ti:m!C: d~$l:Crr1;."i!lCe between the ~]ii!~1~11a.li~ul, mvolvcs lhi.~.l.;hng AMP to these side that of ~lOtNl and yea:'<l" em:ymes, cl~ain~. The se residues are I1lH con~elfved in 2. R. TlIwn!~, L,Sil'!!lMmJ, ~~ ~l~ In~I$'(::!OQ7}" 3. J.1i'I\. $~tllur fll..j, (1111\. {l'lVlS:~.113,,274 (2~~. In the :I'H.~W strU~HII\!K,~1 posili'il'c:~y lJm~ fission or btHJldin!.li )le'~sl C'nzy~ucs. 'I. J.Adilm~ fU}t"i!l:ralemm :n, tss C2004i. Chiifg1;ld sidJ;l dl ~Jin in dbm,~~h~(~.O\~ t,;t .'+" \vhi.(:h Might ~Xrhl~r1 why llilll; ~~Iwr ls !lot B 5_~. A. H'......I~ eloi., J, BiaL a 28 ~2Q)(I131 sy mbQ I ~1:1l1i, l~H': :t.gure,' imerec ts wi lhnegOl ~ ,JiI:D~"3ted by A,MP. 6. #l, ~D~ ~~,,, CWl\, 8'iQL n, ,J)(I;I1, (,OO]). tivdy d'l<lfgi;id ph(~:_;.ph'lte(~} (lj;! AM~f or Re~Q~,'IdIVilgthese ],~inJII'ljm~liI.g ~mcelt~IiJ:~,- 7 ">" 1': iJi!\!i~, !'AIIR. 1TI~~il!\P: 1, ""~ CO~E!iiI. n G, ITIMdre, FlB:!:.it'!rJ.~n, 4211(1.99;). ATP. A mutarion llln.e equli,valel1t. side ties and anomalies \"'1.111 require structures S. M, t 5<1mli!lll>"f, 0. Gro~dif'l" B, mI, H~g~~ M.,A, cluniLliiin the hm~mn en2:ymtl("I2 V<lf~ilutl ~h<'!1 grn~,.m~,~.(I~i'Ulcompl~x~;s in the p're:i>~nJl;.'ii: 'SnoWilll'm, n, UirL1J!lg. 8~m. J. :ro,:lG421,B]2006B21l ~200!9. C,U1se5 severe heart disease also grea:lJy of AMP' or ATP; w£ell~eJ \""u~.]nO!heT merh-

or

~d~~ccs bj,liild~ 0 rAM m1lg

:r I~ !{)l~

n the ~nIlIDIilul,

enzyme, rnaratious in positively charged tl~{; st W CHI res rep erted by T uwuley ~~nd side chains (H>~~lp)'i~g $in~i~;Hposltlons i:n CBS'~ .1ml CBS2 ~upper .'+" ~ymbolsin ~:!ae: S.hapiro. The; effort \"'~II~ ver, \'I.'orth.be I'~g ure) cause simil ,l:Ii e I]ects (3). £lIJl~'Il~J:ll"l~ng whne If ir racHil~[Cs deeclepmem ef new

o(.lls h~ study domain in.teracl~on!'L es,p,eci;JUy of ttl,o~e dmU<I:]IlIS, um preseut iu

'9~ M. SUlert,r iJJ~J.I!IidI. (Iltrn. :~IIl..)2~O] W"~l·" ::il1J). 8" BU1'I\i~'l:.ri. !~~,,,Am. $ H,m 6tM.t~6,lI!QIlI~ e (~ID, U. J.~. kOI~ F.A. F!1J~~., I. t::.liu, II). G. ~~dre', ~g.Ol. 2i~J

806 (2007J.

PHNS~CS
Protoli1~[Irre liI1I,ar:ie lir q 1I<1rks a lila gWOIil'5, llUlt G thil;lTr$pln~ (f.olil't ~dd !l!p. N~'1. e~p'lliflm'ent~ ma:y

Iile'lp re'5~)ilveth]S di:5uepancy"

hm\,wer; 11~.c :>pillOn~li;l~ pilrHe~~$ L"all i!1l:ri~s ic iI;~tl<i!I'ih~i'I~~1~!:ld111il~cai ph~nil;.)itteilOlm. Thi 5

sm:~.as e:leClma s, ·pro· ~.o. ...~.1.d l.leUU"O~ll> •. bL;:[~~"'C.h.~~ p.ilfli.,.w. 'Irn~ n ~! [I ing ~ops,. '1JI.llliked<llss,~c.d reps,
.~ny~nic'l.e!'i'

spill: i5 .re:sponsibte for

!'I'HL'IIIly

r1.J1,Dl.dmnellil"'lrm~

liD e';lJl'ress thesecgluQ,F!(1! Q¥efll1 VriiI h:lgh· (lr I!'neli9Y pankle ocillis.1CJ1I.111'1 ly~e of experiment, one O! l\!IolJi!rized muon q.iiJ O!!I!d iii P olariOl:oo ~r:(lHl!'l (p) illlpmlilch ~iildh either 011 lh.e ~e"t. han d fiid~. As ~hey il'I~.eraJ(l"l:he IIl'IIIQ n e",'d, aRlges a po,la,rrized ~h atoll
t1~ this

5;pi JI.

~wry~

lP'h~$idi:;LS u $~

~~y\fliQ':U1di.;lg!"~lQs

!S!!Ii::h

erties ofrnal~~M",ind.t~di~lg l~.e 'pmt(fg:j's ~~l<!g-

netic mmMIl1,. 'ti1le dil!t::rellt p!l<~SC~~ o~:':maucr in 1000i~tempr:r..I:~nephysk1i.~he !)lUpoel,1ie=:; f neuo ~'IU.I:IIstars, and the s'tability of the known ~ll1ii~ verse. In reeent experlmems, 0) nlumber res.e'kuc:lilI gro'WI.pos ha;Voe been seeking to shed

()'). P~ir~Qr(!\.lrllil-.,;In~,nariITIl;J~:1Ir~pgrt&d~ k~)aJ~ prcd IJll:cd;l~C' pr~ds;e num i)(!r Qf~h ~S'll'pa rtid.e s

cre.ate(jl d ~penlli~ o:n tl'1~ Si,pin eli thill'gluC)'rlis, (Gl in th~

pi(llMilz~d pro:roll, Oi'hich .il1l~!i'i'~the: !\Ilili! (lllh~
gll!lOmlO

or

D(l ffioonstfu.ct!!d ..

some light on !h~ rluNdi~,g, migil!1.of ~ri!ll ami

del> b!:'~w~nt:tiDC(II)'a:lild
P'll'li,des

11m.",this, D't%i~n resolve some btrge di,'>Crep<m.~ C1<p!rd Imc~lL

such mi. Ihe proten are

ens .. which: hiwe~liS:pin of ] (~,'here spin is ~IUi(l~l;7dinlUlh!\. f P~;)!nek:~ o C(](~.!i.wm di\'id(;ld by ll:)'. Whelmwc' probe' deep inside the proton •. the ~truli1~th{)fql!flrk·'g!mon1! ,M1:d ;lmJII1"~Jmlr1

'lcl;~mUy

i;~:rernl'tion)1:is small because uf<'m;ymp'ootie

fr~cdO~liJ;' This iI_InU~~i~J i{llea iU~ns; th1U •. lildike some interacucns, suchas electrostatic f~]~I~:;,~:~~ fa~~ bet\\I'(;.;n ("ll~i'l~fk~ ,gi~O~lz.: ,Ind weekensas they geldO£e'r~:ogeth::~I". If (I quark niles, to csc<lpe. lhml,gb, the force beCQII1I1:t:S

cambiaaticus of mure basic ~illHic~ 'e!1~I~d L i~.;~U"f\.~'l1Cl~nnor Ce~~t~rI h~lv~m.ughl US tluU qj~l<lrks ~nd £~uom; (which bind d;le quarks the eontribution n--am the spin of rheq uaiks ~oge!h~.{). O!l~ I',Jnh~ d,:,~U~Il~g*, pl~'<iicist~, ~(l i!lS.kJ)~ i~ SII),dt on~y,~lb~iilJt {/-'n l"hb; 30%
OV~I"

the past 20 ~,,~~ hm. been

l'Q

l!ml~Rihmd

!'lh(ni~1I Of[(;.'llli

.1

::;l~bstml!~jalCh<!n!i:l~g;etQ

OMr

hQ\\,' th~ PIOliOJ1!s :spfll is bulk up fix:tm, hs qU3rk lJllildel.'S~mll.ding 3bolLt~:fui.eSlllIClure of the p.rotOIrn. To :i()nmi~.{llIt fl. \'~~or(ll!~ £1~Jbal PJ·O£It'l!l1l has and B'iuol'l cm:l::Hi~Ieni~"" Modds Orl~.epml{ln !,"Ct'nCii:fLI:~y prodifct l.hl1ltaloom 61JM~C1fthc ~~lioton's, p'iO'l!LlcedaiboUJt I 000 t~u;:arctk:mllhlLpers. aUld

J;pi.~ ~'!f)Mk]bl)' c~rriGiI }'~ile ifnTlill';ic ~i!'! ofiJ:$, b ~lJree qU::Im:s. with '~he n,'s~:cillTued by om:illall angu~ar m(l~ll~Clli!lli..lmI~~tis. th~ ,qmll'.kf; I~y]ng t mUlJ1nd illrns.i:de the' Ilw~on~, [Iowever. e)l.peii~ met'H:, ~~lCI< RN U~uwpea~~ O!'~,H'!i7,:adof!
CO:JI

dedi.c.m:d ~PJ[I expe~illllLe[l1S <Ire llnlde.1f

CERN.

'iMlI)'

,n

DESV: .Jel:f'eil'Solli uoor.nory,.ubdi RI-UC (R(th:!I:~~ii~tk~ Ion Co~~~d~rl !l'!!!irp~fJdifll'fIV)1 w '\,1 idmd quark:.Jn,ul. glum] ,iUl:gnl:lIr Illu!Omennum

hm: lh~ ~Ilmrks mHJgluOJ~s[:lre ahv,riYs bounrlunside lIulcicl'lr pm'l~des sm:h as the pmron~ they nre Ile'\rcr ubSr(~!Jv(;-d bJ' thcmSJClvl'R'."m;: f'ooepilWlkl~s.
:->llQtI!:!,e'r-:50 ~ rolilg.~n tilc't,1 [iii ~'O\1!.~~I;;'!'Qf;lflJY IL;':XI)(lf1!l1~!U$.

~hl;P'!ll)'(Oll

cil;)lmfb~lions '~u pro~lt)n,~ lh~ ::-;p~n.

Nm:h:m" R~~~rch~. DESY ~Deil!llM:.mes.

lind SL,\("
I~

n~

bel~avcsluke'
!n<lSi!!

<I

.sys~emof ttnee :~mnoSlsive ''con~

<!b!.:lUl

~'i~r]-

~~it~!!',;m" rtw~~rk<; q eu.ryins

Orl]~e

[13 1;\;;1chh~tl o

E~.cktntlllen":Synchoolm!m.),

Si~l1ifoj:(1

I~ue.nts<'I,ll! :~10\"1 j'ieldirl~g ,e1'!.citil1g .resu~t~ (j). Ille prul.m~ i:5 d(.':sclihod by ql!alItm~.lchro-

]::i.roton. W~,lell v,-e

~.ool,: de:e:lP'f!r

~nQd)"Il.,mics {{.leD., the dl~IY 'of '1m~rks,ancl g~.l~~$' (I·S. a bUi,md Slf~t~ ohhr~~ CiOnnm~-d The ilulhQ:r 15 ,~tlh~ Imlilute j'or Th~!J·retkilt ~rn,V.~i~~, "'val!em::e" qmrwb j 6) .. Tbt: qU;JlrJ\;sh<li\~e:tOp.illl Unw~,rsi~t In~rl!ldk~ Ai60i20 ~rt"sbfUd, t!iIJS!!ri~. ·m~it E sMi'S@millil,{~I1Il.{h liZ fllld itHerf!C( l])n~rou.gh!~e ~~'>((:~Ut~g€'l;l.lut of

in~,ide ill higll~r'ie!:g)'i.S1>ip.;fimii;;:~l'S.~hes>l;l oonsl:~tuenl qmuks diis:>olve il1llmne~f m'l~td.e:8s

'\:~Irl"(.·nl"qu~ks ~Jild~,~a (lfql~;fIrk·am~iquaik
j}3ill"S1.u,UJ gl~I{)J,l~"

T,l~~ !lipin

~~]X'ri!lfIe~ts

at CE,JrN,

DESY.,

IPERSPECTIVES
Jeilterson :l!loorntory. and SLf\C illvo:llve I]r~ il1lg hig.l~,-e!]ergy electrons Of muons i~t.oJ Il:Mrge;l
of" p:ro~.o.n1:.'! \\1'ith <I~i,gIlJOO spins, Tlh.c .incm;n~ng e leer re n lllterm: t $. w~t~ .OJ .~ .Irgel prolol'! by exchmlg~n!:l ~uImi!:l~l~llewgy phCllOUi!.1~~21.t enables

II

measurerrrents are extracted from the 111'0duet il.:l!'l of d].umpmlk les (seethe figare)
:!lllui charged P]Of:lS: W.ilrl~. brgc transverse~H.'Imer:wull1.1illlloi;"r~~zfldnrnon-nucleou celli-

sinus, The Ri~~C ~1e~S~tl"(:mCI.HS; are: extracted re:;~arel~en;.eprnbe dC;l~p t ~!lsid¢ lhl;l p!'tll{'I!1. r~om high-I;lf!€'r.gy p~r~ c k: prflduc,tiol~ in i pe hlr:iizedp:lrOton~plfOhJ9~ coW S~OI1 s, MeaThe p~.otIl:n LlI'~lbe <Ibwrood by <1~U<ll1kpolarby IIIi.'; iit1din t1~~(IPlpO~.h!S direetien to the:p~ot{~n $i.LremenU ,of the :S1;;<!pOhid7.:<l~tngl FIERMI!S (J[ERA Meas.~I.rf!'l[1:e:nl 001' Spi,:~ .. but 110~by one polOllr.ized i~lthe s.am.1:direction as ihe phfll:(l'll. This <llll(li'iiv:-i us t'o extract where H ERAi~ the H~ldrom E~~k~ronR~IfI.g infb~;lInilition abmn the~p:~nonhe QJ,lq4i~~b; when A Ill.~ge A ccel e'li'~lOl'at DE S V I' .':!ugyesl: tl~.n one cnJurols tht':: spin pelarizarion of bOI:fuI, th~ t hi sis also s[lm~,II, 100 SIl1Ia.III.CIresol \Ie the Sipi ~1 beam and the prmou lOl[get TIl.e RHlC sp]:n, puzzle. R~,d lht'L~:the J,O~VoGI,llI .• uk sp:il:ll'o:trntlribmiol':li is; ~ppmxim~IBly~nlrnte{11 by v!ll~n{.-..e experjments ilW{l:~VC li\lii!l:~:·encrk'Y ro'llflri~d p.lia~:on~protoll co.~~,isiQlI1Is •.l<Itbe:rth3D eleeuouqlmrncomr]bmioJls (91. promn :iilllcmC'~ions. New. mose precise glueu-pelarization In OlI'l.lIY7.iI'!.I; fhese COm~iO~l:;'. the boy 1m'~.iI~~lremems wm saen be "l\fr!i~f.blefrom the :W06 d:u~tl'lkCIl lit eOMIPASS m,u! Ri~nC. q~lcsl.i.Qi~l:<; nare:\;Vb~t { ll~.'lp:pensto S:P~iO the' in I,trn.depell.d:em ·nmJ;.·';I!iuremei1n~ofdae v'l~e[u;e .1I11d ~1i'lmi[imlfrOillill CUIiIi~~1t I!!ll.tr1!;s Q lt~lo~e pmoed in ~.igh'iell""~I:gy exp~I:~InCr.l~s;o eenstiment t :oe~-,[II~lkli~k u:Jl"JtI'ibutuol1:>. \"'in Sllc!~1be awilablc q~uMb; Hh(l btti~di ng: bh.l cks 'IJ I· ~h~ prol\)!1 )?' fr~n'~ COMPASS, It wjll be iJtH~n::sti!lg ~u How is the. Sj),il'l :~/2 value Qfli:te PIiOIOIli bLnih see whether these 'CERN data confirm the !Up ftymlh1;l Spi~1:O~~!1droilOi!~ .!.f!,gtLl~~f.m,um~~,· 1:1 o ERM ES resu Its. Ex pedml;lflt:Sf!l j'4;l fr"'~f'llun nun of the quarks mulg~ILOm;illsi.de'!'IVhy is L~,'I)onuory are prnbiing lite sr!lin pmperl:ies of rhe ~'il!i!i.[lsur~dquark spin comrlburloa SO the\>a]enc'e quarks in kinem<lliC'~.'\~l~re they S~lIiI.li"U c(l.mlpmfed\!'r.'ilh (I~LiI.t m!odelpredic .. ure sensitive 10 the coafiuemeru process, lions', ls the: '~niss~.ng 1;pini"a valence qua:rl 'efT!::cl ()f~lnui.bu!:;'IMe ttl the sea OfqUOl"bJ OIml l!l!lli,qlil.lm'ks'l Are tll~(l: ~~ci[ath:lIls ofl~l(: qIUl.llr.lt· ~tMMUNOlOGY .umi.qm.ltk sea pO~<li.wizedl un the o~\Mpo~i~eireed
[ion to the
11110[01'1';;

The p lannedl J"'GeV ~;[pgl'ad)e f t heri'1c~,i ity o will meke \lUul cO!Hrihutiom to our uaderslanding of.orbit;1i 3mi,gular InOmC91UII1l QCl.IliI.ri~ hutions to the ~lriUloil:s ~pi~.l. Spin rrteasuremems h~ve'~. brig hi. fUlU.re
,11Ji!dCQ"Ui!i~~!lU~to

ch~n~~g~l)l:r Ui!tdcrs1:(I:~.d.i!1g o
<Ind:l.h:l1I.d<L~

..bout the srrucrure anile preton

mental

;;'i~~~~~

.of quark dynamte:>.. ''1Vh~eh

e:x.cili:~g progl;'ii"l.ss. 11m.be e 1111 made, The next years pIflJmi,~~ be equOlUy ·~)ol,cili!\g, ~(i!
UUm;)peiln Ml!I~rn(Q,'llaliJorali!!!liI,_J. flip-i. aaas, 1~-Ua.9i}. 2:. 9;. M~~ tl' at (Silin M~M Cotli!lxlr.lliol'lll,l'h,rs. ~v. Di5>1t1.12001 U 99'.8J. :3:, ~. ~b~ el.~1. (llIlACU,4l (dllilbDratlan). P~Yf. 1'1i'1'. 1~5a:,
1!ll{103 U ~9tll
oj. A. klp.!lp~~~tI~ ~

:t J- I\'lmlfIla!1~[~.

R!etleJlern!l!2;s.

Di7s" (112007

'l'200ill_

~J"1E.F.Mil:$ Coll~liar~!irin~,.PIi,'t

.i;i~;

5,.>. I)',. B;il$~ C Ii.. ,AI~~lol.ilt). MIld, My$. ,1121.l~07 l ' f2006t 6- A,~; 1hQm~~VI, we~~e.lire :Ii!nll;'(trI't()/ ~he1lltJ{leoo ~Wi~~VCIi1. I!!~!il\ ;!iOOl}. I.~. Ill. EtaS~i .w.... i0Jt .Ph,s. 77, ·h2SH~iJ'iJ$J. 1 8. s, mlM, _.u;il",.m'QJmfiP-~)j(j~1!2all S. ,. ~ERiY..ESC»Llabllfition fA. A~~~l'!f~ia~i ".1.1. ~ fl !Woe\'; W,~'~<l,(Jll~O'05i(i!~l'.

Sl~,in(thus caneeliag some

of thIS$pi~l)'~ n.e spin ·ofl:~.cgluons t~nUbind i:hepml.on C;U1 ,>1I;,~OOellthe spin '(lflhe qU<:l!rbn~,emi'\lnld in hig h C11C1I"g)" experi nm.C'JDts,. ]'t~akillgl~]C' sp:un loOk dimiQl:~~Il!~d. This ~n~"C'l hi proponloaal t(l ~heg~.Ha~] pclarizerion .&g. 13m how ~af'~' is
oi

A:SYlmiml'try and Immune Ms,molry

.AsVIll111i[1e~ri'Lcell diV'isjiol:! olf LYIiI~phO(yt!e$ensu res that emf .i'lG:I.a.pllvehnmu11!'~ bal.i.in ced proiJ lICliolll o·~two, di (ferl! nt~ypl!!l. of T (!!!H~.

~Illis glllOI'll po!@1fiZllTj(Yi1iI? il'h~

q[l{IIlUUIn superpcsirionef

'QCD'I!fU(:lutUITlI!J!.i

{i

of

~Hl~l;.~ Cb"f.tct'r;l:lfi7_e,c.!

i.[1JfinUle mnnber by nuraui\'i~l!~ ~pil!'!

an

'l;Mh en. ene puts .1 Y...te~~cl[;! 'Clm'lrkiin IThis V(!!L'ilU,lI.U, its S]lill can beeeme dc.io~l~~.zl:)d so !h.n the tt'ltld spill~ becemesn pmpeny or Ih~ rml:o~l ratbcr thlli1 the s~m,ever {hi,;:i.ltdi~ \·idm.11qm.uks. ~J:mbed ~Iiihigh~llel;,gy e;;:pel.'i~ ment::;, ~io"" big i:s dllli:s efl"ocL'
.swUC!l!rc.

~I.oilJ'ie: of ilI.llilig,(.'11-spedll'icce lis ~B,lIld 'T cells] (h~[ rcs.P't")!md (lind dimiji]2tI(~ divcr.oe 10 p...hosn::~nn;$.. The sy::<ternC<ln,ds.o 'rec.'IU~J!rlwit lUis inr~e~ ~l!I'nd[¢\i:pr.md '~1~Orc imls .Ii)p.id.iy .!I!ld ~iffec~ively whenree1>!.polSed to .11 pil.tho,~en.'1 f~~tlli!!:~ kl~(}\:vn~-~s i~~'m~.!loi.ngi~~i ~~~!ll~Iry,

[lr ;,u:iaptiVle .i m rnJll.lle syste m is

endowed

sys1l'fIl maitrdai

mi.!; ..

.I111ikl!lob.bllc:h;I]lenge.

'T cells ap~;J1"

[oli1l;l'.I'{~

w~l~~D'1: erl(]ril'n~{ll~:-;reper-

Nhm,\lurements byl~iI: COMPASS lC!Jw· J\J!UOr:l1 :Pmton .AppI'lrnlm; for StmclLll.I'e· .md Spectroscopy) Co~I~.abon!Lljol'l <11 CERN
I'llion

i-1uwT cells di nl:re;!li'~iat1;lint~1bc1tb s~{}rt-~ iv~d

efl·ocwr'l,£U~ li~cl'Il ~OI1J];h<1lI~IlIDctlm.1S <U1:dkmg-

~hred lliIlenlilory cdl!:! dH:.t prmect U~ Co:[[ye,u~

h<'lsooen ~ 'C,c~ltrdl qLLcs,tion .~n imllfllilloiogy ~ Oill p::!geI6~n i~l thi.s is~W!e. Cli1<m:~ (tl at. 1). . {l)pJiopose tlm~ effector ~ndl memory T IL"e~.I:s !liiC si mu It~.rt{.l~ii.!$IY~ll€:rf!~~d~!:om ltl~ dJvisian ,(m~Ii()S is) .of <I T ce.I~!. ller it. responds !i(l n .

PHENIX (P'iom::c'ri:~]g, H~g~1,.Elmctrgy Ni:J1de1u· l~t(lr¥l~~iDn Expcrin~~]'D[)1 .. ~d STAR '~So:l.el1ioid TI~n::ke11'<11R.i.~I:~Cl xpel'iilllmel1l1t:;a~. e RHJC ~g.gest U~!Uth~ ghwn polariJcilrriomL ~~ much too SIll<11I 10. explain ~hl!:di ff7erence' bet'liw.:en the q~li<lfk model p!li!ductuon of-OO% t~or the qmltlk sri 1'1 cmltrib~t~;[011.,uld the Ilie,lsm~d v::Ihm of ~30%, ~ithough il. m~y :!':lill m3kennimpor~<lIiu i;lQl:llribmio:m 1:0t~.le net £.pin oftbe p:rotmn,{S,' Nj. The COMPASS ~nd RH Ie exp",ri !l1ililm~lL~ed ifl"i.lr~~l proce%e:s ~)o ~uoce.'i<s:he gtuomlpul<lrialtiOiIi1. TIme COMfAS:S I

,f1In(~ the

:-~~klp~edn e.\~tllu tiona!"i Iy '!I1C h;;mml';an~. uf a allymnu:tr.ic.aily p3rlition~ng ol'dl flllle determi, 1l,,'lIl1l&.dm~ cnslI1ring balanced pmd~~Clio1l1l of bl'l~:h cell 'types ,mrl iI.\'oidimg depletion of the T T ~~Ill~p(!'r!oire, WIiiI.e~ microbu31. 'p3thosens breach om !1Il!i,n.~o!ii<r.1'!fl'h;)!'.\i ,~tHI~h ~s f~l~1 itli!.~~~.g th~ b [~~:' go~sl~.-olnl~stin:~1 or !1:5pur<!lory u-.!cl), thli: in1n1\'1ilehnl1mm: sys~t~I'I1I:respcmd..., by Pl'{)ce&o;;~ i~g m~dpresemi~.g p ..~hogem1.cQ]111pOn~i'lb~(I T l.~elk. Mk:I.l!)hi~~. !,l~l~ge:ua:s are il,oaded onto m<~jor 11 iS~'(I(;omplu~ bi lily cOHlIp]ex 'IM tiq rI'I;o~ecu~ejj,0 [II 1he ~m:lFr<l.ce of cleli1dri tic ce,lh, ~~!ld '~hl!~ ilUemc( with .m~igcll~cep[ors an IJill" S!u:lllceo:f];li'lI:hot,"eIl..lsl,ecn:Ulc T cells.. TI1 is imltt1rm.;Uon $~.hmd~:teslhrma'ti(~r'I of ,1111 il'1te-r··

11:~lIce known <I:sl lhe i:~~mm~o,~()gi,t:OlI.1 s:yn1l1~~e 0- 111. t~mafl is ~!'I~:h!!lnIo~rdl Hugn~ Medical ~lnsU~UIlie, U {Slce the ri.g:u:~~m.ld involves, ,~di!Otri.br~.tiQl~ SlMJa Ll Institute o.r aioll1,o~e(!IiII.arMedid m~·. r.l~~·'II'(nK , U!ii~e~:I~$choo\ &f Ih!dki!'le, Ne~YQI'k, 10016,I.ISll '4'1' (If other T ~!'i:,Ml'\lrnlC'e compoliie'ntslo the H. Slnghi~ in Ih~ tI~IW\Hd HlUJgh{!~ Ih!,mj[(allInstil'~II~, gynflp~.illd liJ.d:ilillgca,n;Qc~ror !.l1o~(;cl~k!fl D4 C lJiepnlmem of t!.Wlecw.l<Ilr i(jem~otk~. iI nd (e4!~ 1l10I.o!l~" Qr CDS, adh'e~i,OJil p:1f(J1t:ci j i~ntegrim!, . OiI,mi 1'180 ~ Uni~rsi'~ of (liIiiugo,. Cllii·C,19i11,. t ~(l63'. USA_ [-Imo'iil: I cyr(]l;dn~ree{':p~on.; (3. 4h IUIlIIbiII,@~ulmli.ml1lll.It1IJI.Ild'IJ; In~iDJIh@I:JMi1:"g!Jl.llldu

1673

PERSPECTIVES
V~r.y e-fdY~!l sy~o:1p'\le'rOTn~;H'~O!1J.I'hero ls localized p:),~ymeriZfllion or the actin cytoskel~tf.~'I1'!1 andthe micretubule o!:Y;;tJf!~,xing center-the structure from which cytoske I~T.. I Ini c retn hules rad i~tre~orit;l1!i~ Rn~I,Jrd the S)'m.'lipse. Otileli' C(HJi1Irpo~n:n~s. indudin~ cytuskcletal ezrln ~iU~dl¥lo(i$:~~

It cannm )'.;;1 be W~!L3d out diD~ulille C'D8lhi and C Di810 cells

used i~ Ih~trollsf~l! e:K~rimenrs are ge'~e.l";!lted mde-

penci&1ulyr,u[ler llllall1 derived

fram the same p:mge:uai~oD' T cdl.lf.lnavj~,~1 tn lcroseopy with AmnescenUy ~.''lbe]l'd pn:l;t~i!Us cfiuld ~ddw~6;dl~ll issue 11 nd perhaps SUPPOI(;t the <lsynMuellic cell tliii'i'is,ion model. A'fo~Ii1e g~llerilliu~ of <~ memory

and the ceU s~~rr<!,ce prote:ur.l ('D,,*1. redis,~ u:ilHt~e hl the pol~ OPI)t'i~jtc [h~ t>:y~~~p~~ (5,.
Synapse ftmctiron in l:'egui:.~ti,:lilg c')'tokine' secretien ~fil.d: T cell reeeptor signaling has been illt1ens eJy swd,iied. and it had been, S;pCCU~iII,U;J tbat coupling of mltosis to tin: cytaplasm ic polarizet ion II~ is; inrlueed 1I pon ~H sYr!~p~ r~umiilUon ~ould Syn~ pliic pr,01!~in!i Iresult ill an asymmetri C eell Ptp!lideiMHlC .JI(!I: div lsion ( "). OWQ T cells remain ililleemact ~F:i!I,-l wit"lI all!!j~""!lpreSt;flli~,:;dM.. Po'lanl'l' prc.teilU dritic cells ftn several heurs, QFl{(J:; 5i;U~L:S.1~.lg th'!l co~~~ts ~ou~d .'ScJib'b11! extend. ~hi\cmSt1ilmjtosis Ill). 10 Nwml:l ChMg e~ed. I~t~d the a.~)'!n'

bur net lin ef[l>:et'or(cywwl",icJ

c Dg T cell

rc:sporue,

ens T eell

:1i"C501.lonse, IlPCqUilUCS eD4 T h~]p~f cells It J I).. th~

I"D~.e T lhelper cells in rhe of

ses wiU .d~n b~ needed '1[1 [)~~feii\i!!il~ di!g,~ht(!n. oClHlnl!! ei aL {2} propo~' lhal the daughter ce~l.adja eli!I'll (® the ~(mill ulilol.ogkill )'yn.ap.se g:il,l!!,s pI'OV~ lIl1It CO',IHponems 0 f ~:ii1.e rn~e tc) 11 n 'E!i~re(tor cykl~o.x~(T ~~liDpho(vte, iHiliiereil~U1i1t ~m~lfiCnioC~ P,ltr p:;l.l~'\iiJ'.'LY !Iii\: rt:qu ired tJO g,enerarethe se de(i'o'e[lfmm l.n~[hMfI~ pole dinf)'relillr,ate~ orl(11 illi)l'Igi-~i",~d distinct T eell I",ul~~s,S;~lb~lJlIemo,l)I'c2ll.
in neul'Oblij:!$I;S

a.'\,yl1l'r~~~~ric d,ivi~ion n..e;ds ~Ci be' addressed. Gem:t.ic 3JlJiII.ly~

metric c~~]dhr~sil:lllhypOlh~is
b~ .1)~13~.yzi~~g T cells that ",'ere

elil,gineeredte express a T cell reeepeor !'Opeeifie :Worn Listeria nw,wf:I'U'lgl?Ul:'j: il1m.l~.gefl.

The T cells were ~aooh~dw i~ <I fI.!l!lo:Fescem h

dye <In:d i'lmJeC'~.edinte mice infected with the
1:J~I~og~!tPtaUiilrmiorlnrTe~ll$wa1O ~'l!)nlitored. f!IX ".i\10 by stepwise d illlll:io:r~1 Qf~11.edye. Thi~ p~j'j,iiJl~UOO ~!}i'ling (if lind~vicl:t\d y~l seuvated: T eel Is as ~<d.~as these tlh.tl.had d~\lide!tl: Q:ll.C:C Polarl zlltl.kU':I Ofpr~]l~:hm • ie ~nul ruirotle T cells \"~<ilsvel'if~:roy ~}oc:<a~,iz1Jtn:~:mof,1 b microt~lbull!;:organ£z.~n!il eenrer !ll (he remnams Of3~1

(1m. ,~!as.I.~wboeen impl iClli~e(il1 inllC'~il1~lxllSled

oogr~:,g::llion of$~~I~~ling oomp(lI~C!ll~ to tilL;: immunolegieal sym~p~e(6'), Together; rhese fl~d~!'lg~ SlL~~st t!llll $igna~inS FruIn 'th~ ~)"n;Ip'S!i;;! result ~,~Il an <!~1mfl~dric :iftlli~Q'$is~h,Jir resembles those i:n i:n.rertc:b:mte systej[f;]:1i. Ah~,m~g1fgi~~s g~ner';I]ly assumed th'lt both effector and mCIDulU'Y T cells can be del-,iived fr(~WJ,l t:~.e snme p:r-ogenlwr T cell, rorm'i~ pmolHor such aline,lge rdaliom.hip has been ~.;;te:kill,g (ll Ch(lng~>I' cd- [I~J[ked that atter activariou ·O[ the dye-labeled T ceUs in ~~~p~ewd mice. fho~~l;e]l~ Ill"ll had divided once cOlul.c:1be separated into subIm'p~l ..tlons ex pre;ssi.l'lJgh ig h or :110\\' :u~oun i~ of CD'S (COt-lhi ,n1Ld 1(' D8~1J l.Plile sUllrn ... bly. [11is eoere spoud SI.O the d aug i'i[,c r cd Is l~~t M~ gener;l~ed IP'roxinuUy <linddisl"'UY., respec~ li.wl.y. ~~l~h~ sy~l.[lPtiiL;': f~]rgC;;c'd by the: p'1Inmt~.1 T {.1ell.md d~fldri~ic the f~gwLr!.:).ln laci. I:he CDShi eel]s had p~.enolypic milrk~ e;rs dmrilc'ter~,sl iC (l,r eUl;:ClOr T cell s. \\ihc·rem •• he CDSI\) c:eillls ,exprcs~ed memory .eeU :~~.'lfl;-.ers.\Wheli1 ese l~(;.db~Jr.lere St:P'I'· m r:ne·d all~d u~Jin:sfeJFI:ed illilo I'ccipiem mic;e' .. bO'I~ arlimO!l~pflrlul~tion~ l,lfJi;~li'll~lyc'lc~r,ed 3'1~ m:::me i~rWecdar:1: wi.th ./., ".wJI,oeymgl!u(!s" H(l·\',:~wcr.\,~.·hi;;n mice w~re i~lJe:C'I(ld30 days trallSfei(," orlll~e two I.YI~es orT 0el,~s.. 'the CD8~~ ceU:s 'Wt,:1'\! lll~lch IllOl'e ~ffecti\!e at. cl!:.~nii;WJg microbe. These rle:S~lltsre 'con· the a si:l>~~~t itil;;lCt ivated T ce lis ~m.dc:rgoin.g all w a:SYII.lmu:'~r,k division tiO ge:lile~'<l,te effe",~;or a!l!d ~.nel1lory d~.~gJhtJcr (;"(:]Is.

of N~LllilJh from tllu: apical pu~e

(Iuve;mly. ~.l'lM~ be i!'!l'lPOi~''lItt W II udentify t1:1e molocllll~u deteemjl:1lHli~\':l ()f·~ffecwr and memery cell ~~~J~i.lnd explore how I:hcy are a.syml1lcn·~c~ll~y segregated duriugthe first cell division.'I!>; we ll <I!S hmll' liley promote d~:s~,um:tsmtes 0 f
~~m!~ ~Npre$$il]l1<

One major mech<ll1'irsm forailymmetric<l~]l)'

~]nitiofling cell f'lH~d"'t~!'inilil(l:rln;ii1vo~\'i.St-l

polarized
l[l~y

transport

of RNAs, (12). and. ll.~is:

a lso be used by T cells, The fi:~.dI of hlgS

clustered T cell reeeptor, ('m~_~lmll~le- integrin LFA-I. To determine ifcylop.~mmlic 'Pohuizatiio:n. could ~iEl<!d di:;.tinct mitotic pm~en)'. i!le (0 am~.o.r£ ~lla~Y2Dd Ij~~ subeellular di1>~.,il:!Utirmll of col~lpunent:-; of ,111 evolutiotmrily l't'lll!lserved ;l:sy.nmmetric een di.vis:iolll p<Llhw~.

imnnmclegical

:::lyill<Jpl'lf:

that stil] dispt,)y~t

'h~"lng It! al.. '~!{J11Ii,g wirh Iho~ on <I!j,Yml,~],m:tric eell divi~~o.Ull th~UilHII.mllifli~.iiH1. in eplthellurn 0'3). sug:g.est that. the Par pathway ~:n<lyregulat~oo~l~\ren¢lllm[~ ~Icmcell and cd~JLI:l'Ir di I:" ~ere~~i~~i(ln in .~V<l~~ty (!If dJ;:\iel(lfil'Ql~" f<.lwli.
la~ C0I1U:eXts..
RerlB~OO(~, iI. V. ~Iil, S. S;w!!:at T, s, G.oofr~" 8, t R'OO~@, [;t, A:rniii!ied, (!JfJ', OpJ'lt.lmmuirQI" ~;El. 2:5-) ~OO1'iI"

.2.

liliIw~i,Ji:r<JoI;'pi.tI.~~i.i~1

~~Ut>

of

{JmJ;(}pMlcl

rnel'anogasJer~. anc! if] the Caenori!tJilditis ete:t!fUS ~Y:!~Jlte. t hili p<l'lh.,\'t'l)" ru.~~ i!l [lnJI!;li n C'IJffip.I~1:t rp~I'r.l. RlI'6. lIm:1;,t~k<l~ ~:mClrein ~dnase o
(' I( P';:I.r p~thWfiY)

c~n(~.~ .

LriIJ7 ~2:007J; ~wblli!h.1I!!!I ,~nlilll~ ~.~~141 ~O(l1 U'(il,!'U:;:6~rjgm,ct-nl~'iiI~l. l. A" Gt~.a~i t.r:"1..~~~'2115,2;21.U999). -=_ C, R. ~n~. ~_i!.. Iir~lbef9'. 1ft ii;u~lillr;i'lLS!;ii~l:)'.
J!L tiLllpl~~lilJ'ljl\r~J9S .• 92 '(19'ii~).

J. 'K (!1amgr'~ 11/" 5d~t!CP 3Jl5.

5. P,l(:!t'!~~\ll!l,A.'. pi!!~iil1q.ll.!i( l!ri!o!_~"r~1;,I~. S B

11~1,i',

for sC.greg,alit~g 'l~eU

nne

de~elf,ilIiIi~~t:lTn$.'O the <tpieul ceJm pole before l IluitOiiiis (tI,_ Ch~ng ut til. ~otmdlhllt lhe !'Ifyp:i~ C<l1 pilfotei 1m kirm<'lseC ~ loca~,i2)es, ,,,,'it~.l llli!Cro~ a U1bulc or~UIl~:dn,g ~C!int~'r,at th~~ po~c o!lrpm.ilt~ ~he ~~m!l'!<;!n!$of'[~1!e in~mli!!llo:~ll~li·t:!11~ynar!S~. In OIdill:~[ion. N mnb (1:'1:1'1 i nil ib~tor of Now 11.

c:. Cti1~l1l (WliC]~ 2!1~ t'WOO), ']- C :5I!Ji[I~I'I,it ~pel, l I. B,~~, !I!!_ R WI'IAndI1i .... , Immlfmr¥ .i1::1,315 (OliOO4!~" (j, ]. !!et~,~i~!!Ie-~ ,II.. ti!i ootidh, Ctrf,; 8i'oL 1!~,Rolli! J•. ~2004), 9. ~+R~~~I~, '1(. Nt,Jlim. {!<II'!'; Oprl1. (~jOflJ~ JI~. 195
6. M. IL 11)i,iis1ID,A. ~2oa·¢)"

u'U(l!Im,

fifl',

r~lhw"y 5igl'i,d~llg ~ segrecSOtted:<Jw.!)"

j'lJlOlein

..ner

from kimlSe C ~. sinl~.liilr lfiWlllcll hil:d been,

~:Il

d~~:cribed
Sf')

'l~YIDIlme~ric mito",]sin
NQa:lUHI

IJn"J·

n.

10'. F, '!'II, C T. KuD,. y, rrl•. Iii~"NetJr.rJrr 5c.!..n '('W~6). ill.•~.;'I. aed1, It. Ahmild, SdtMl! ]iW:Q,2'6~ ~Oin~. K l2.!I. fmlrrstenbl1r~• .l.IBrwd~l" C. Q. l\!OIi'.Ili.), ~ Bioi. [ ';Ol,~i' fl:\i(l8}.
T, [eth1E'l; Eo ~uill!. MJWI~

43';

2jlS t200!l1.

1)1i Ua ep ilhe~ ial c'eUs 19}.

scle

~ly(J5i~ll

\\~l k;~ CQlllribwl;':s ~fl [h~(;xciu:;.io~

IPERSPECTIVES
GIEO IPHYSIICS

II

Tile 2004· SLlrna:tra "Anda matf'! ~al'tl'l~!iJ~h !I)~cum~d at ill s!lIrpri!>ili!g ~il[ll'e and IpILa:~e:he t ~<e'~SOMl~lme,d 1i!1<jY' ~.eolprosral cl1)rlHnlmiti~$ ill the fu t!J.mt

l!al:'.g~roccur .~ -;ery i.lllfn::queuady but C.'UllI cause widespn:f~d damage ~mJ loss 01' life .• 15 \\11; sawwi tht~ite Sm~,l"n:'d,-A.:nda~~.l,ul ~~rtjhqua.keull IDc~cun]~,:u3'r .:?.:I:i04, Mo.o;l oj'~Ll~e earthquakes occur .\1: dle type of tectonic boundaries where oncrlal~ .'ii! ides ~lita ,~Clltl~ angle heneathauother (;:~pr<.H';~SS known as subduetica). Beesuse theyhappen lIlos;lly b~il1eaD~the ocefllttih;,Yoflo1olfl se~t:lmtl:
di::'.sIr~I!l.:liv(! tsunami
\I.,'IIYeS,

;!:rU~t._~l!lf. !k. '~!i\.Vit.h 0:1 lilliig[JlJill.!ide9 ..

1 ~

11,119.) er .

••~ IIJ~2 ttl O.IU !tl~t~n;~!"y~ilr: Ih~~. ;M'i~~r'Ige time be'tM'ee'l1l them :us.10Cl te I m:m years, asS\nni.ng all the' slip. is by Me;! tllJiliru:S
•. ~r :;'011l1C

slip

YCC!JI~ tll,lj;iOUg;~]::;mfl~~.el; uakes or creep. q

rheinrerval will ~~oll~er, From ;'!!iIi obsc:uvaliClmi~siandpoinlo Ihi::\. ~ol1gilmtcnml is p~OM~:liil8tk:.beC;;;~'~ inmost places, relleblereeerds l'Df~;.l~tbcluak;l;i:> dm~ baek Oi.Im.ly ill eentury, Histori~ aCCOIJIIHSiill.ld

1:l.~lll:(I;icob:!i.erv"tT(1ns

C"lll

be used

~t'l xtend e

[h~ reoord. but they lack detail.

pn)p~r[~G~·onh~diITereil1l[ ~lubdudio,Mi Z~~fI!;1'~ . .An undeil,iying premise \\1<116, that. as a Jre;Sultof geologic l'acIOJi's. some Slt'b(lU<ctu.QJ1 zones are inlr:in::<>icalilli il1calmble of g~mH·4~il.£ an ,MQ ll~lakc,lf~ruc~ !:Ms \,\i'Ol,ul.dhe .illinPQilfI!!nt m klli)!,~··. Acc:arcdi n£ U) .UI. (::.uly ~dell".lhe <I£€: and gpe<L)d, onh~ .'iiHMnClil'll.~ Fia!e~'oI'€f02 li,mpOrl~nI: [f fhe subdw;t]!'!g: p'i~~le 'liv.;r~~ ~~'O!ngk"Uy .}'O[Lrlg, (rnindli1.creJfort:, 'W<l111Il mul buoy.nul. 0.11' m(,~.ri~~l!-Illlck~y",or both. Ihto;fI i~:,o.; sh<J~ ua' It'lli!.! j CClQJ:y i.llitOI~l.~ maorle wou~dnn!.~e~l stick mm.'!;l: ttl lmt'pl.ue .\bov~it. le'l:t~iIilSH'!Digg,er quakes (l). This ~)de1lhad e:m.p~:~c<l~: :mpflorl ~'ruln the

I UO-~<If 1I!~i!itori~~1 ~!.1.fthq!i,l".k~

CUiI,

record 'lJ)" Othen;.il.~lilru' :suggest ions based

1[I.aln;~.erz:one:s.SulildllJdiQnz®lI"l:f1 ~gr!neI1lLS {bh.l9' (mve~) anll1 t~(l(lfili( plate, Iml!mlila.rt~ (brown (Iurves) with mile d drd~s shlJwvnili !oca lions 01' ~l1.aWflIl',a·rtl'l(JlJabs, 1ilI.7.'5 or 91~e-alersince l:900" O,pef'll::i~les shol,\';' incompl.ete samJp~e or rl'l~erred

or

I(!PJ~~t ~rthqy~k~s;rn:lm1700' to 1900 (6), ~me e s1.1~JI1e:(ll!~ i!r<!~Ir~~ 0'( i!.~,9 l~iIIl i!itllrlil (JlJlo1kll~ in 'the' past 100 ll,(li!rs,h3,d IJh em 1i1 tbe pre-vEcws.200 ye.il~. pl>lt:e:11~£iCh;nm.k::s were that the :I<lte~a.~ lnatlCHJI gftltt:: ::mb;Jucted plate ~n Ei~nh:il'mil~;de r1!'I;{ld,if.ic:cl. 8~U'e!):il u lhe pl<ll:e bOUII,uiary (41 'J'I,uB~1mt o earthquake occurrad in ,1 '!.i(':r~ llnllike ~yreginn. ;J.ooordi~g 1:0lfuese e;~p],ml:1li!ol1is. (6).
l.r;I eoanast to. mec.~~nic;::]llpm(:esses, :sulJ>.. ~;<IU]l(:r), The Sunmlm~Andllm;nl

Ihick sedlmenrs in the trench h~briC'al<ed ~h~

dm;ll:ml, tDf yOlllln~. hot lithosphere :h.eOil,~~ the "fhcli'i:~ ]'l:lOI'e dum 40,000 krn ofsuMuc,. m~ h~ ~~,II~ce~where long l~b:;t:orrics~11"l:: :!IN'dil~ ~:au]t zene, Bind ell.nh(IWI"~!>]'):111)' be inhibited able, ~tn.e~li mes between gr-e,lt ,em;thc~mlkes in this settiug because h:ij~~·ternpemlures t i0111 bcundaries t ~e~liu'l ~im:e)" The rupture g uf(iij)' oi1~ enl1ti~QUs; ~I;:gnnl~~t -·gOO km or' arJ~~m"ttl bfl: h~igh:~ym-gu~!iil:". in C;;!M.~di,fU, for wi~hhl I~!lrtl~pm~nm.~duetllc ,OVi;;:!:' ·lriiriUl~ i . ftll'i;lN in ~~qlgth C<'ll} pr:udw';e an M9 earth~:x;alnp]~, d ish!l:ba~4;I;~· f d~·~ () ::;(llfl ~diVl~.~rlt~ deformmkm (7). R~'g~~-si9n~ ~I;!I,ting·~~ri:h·qua ke, Se.i~l.nQYogi SI:5. fuI<LW l!ong tried 10 deterby shaiking and d.qmsu~:s 01' sand by tsunamls, quake behavicl:rl.o f<Jmlt.~zO:mle t~mp'ernlures urine \\,'hkh ::I~£m~n!s <lr(!:ifLWN ~.ik~ly dR~ll ..~ both ~1~ggestive of pll"'~ gnml ~,U"t~ulu.lke~, guvl:. :::;imil,~r~ti~I'~:5lic,l~~1.~:s rhe l:nechaglical show an ii1,veiliage'tillle betv,lcel:l c:v·el;lt:HJF600 others to break, Yel. 1:i1iJ,e;' cal·~ltql~l.iilk.e of M9' I1i1mlels (.NJ. W lowever, I:~.crnmll consudem~i:ons 2004 1lI.I1)U!rud<l.~e,gJuem ~'~~ut ..s theught 10 ye;}l[$ ,t h HO\\·tlver.. I:he actual rintes tan,ge ,,"' p~Lt Andill:11'IIl. 5lLbduil'ri!!)ll-ll(mel~lU~l in the l~.e be amon~ the .1~ls!.likely togo. \Nkn yovcr,mN frunm, 200 ~(I ~ 5001 yl!<1.rs. :reve:diung: a 'reI')' ~'<.I[ge hig,h.·l1i1Lt<lgHilude class; It!l> ave.l1I:ge lempemtme is p.mhably elose [0 ll~iiu ,[If ceatral Chi I.;:. the frt:qll~.cy ofthese massive OILW];~~. and mr!d~)tn!W~s \\il~Bnth~m~rgillbrn;llk&' to are a 1,11 subductinn segments cap~lbl:e ofpm~ Thewcrldsmajor StLbdJ~1.(,1uon )ones.CIIn 1'lC where d~e ~arge;sl known subdm;tiol1~zoWll:e 1 dt!d.[JlJg oncl dhtid~d imo [;(\v.e~.ls(\I.mem$ bfL~l;ldflml 11if1til.ln·d ~:i:rtht'!lruuk~(M95H'Iccurrei(hn I 96fl (9). lEa:lidl,lqmd;;e frt."',qlll.'!mcy cam. be esti mated ou geologic changes or earthquake lilisUJries (see The rate of eenvergeuee between plates ).. the basis ofWiI~il\:'tecU)lj~cs..A.11i .M9 e'II.·~I\i~u.:Jlkc the ~:·.i.gm·elSaC~l:>egln~:I~thl long erla ~I,g~to cm~ Slff'cctthe generation Qif·.£ rl:ilI! cl.'lnhq na~es, ac'Coulll~:S;tor 'Ibml~, 20 meters of slipen th,!! pmducem'i MI:) quake .. :Fiye ~,Jve had ~vl,9 inancther \\1'ay. Theon::tic01UY. the frequency bO'U:ldary bcH.\lCC1l tWl.lplal!eSl,\·vbic11t,:aIlVCltgc ~Iualke~.hl the pm;;t. C~!ffi.tu~.·y: .Immy otile!1l'Shave C'iJlrlh<il~',;Ik;;':10f 1:'1 given size i!lcrocfl~C'.s \Vid~ o nQ"~pPDduced (!Jm~kes gremer dmn MS in Ihe mle oh11e te]!,ltiIVle pi'ltellllOfion <t Ul)t, ~,r\'!'e recm.'dled liIiskJry,. hllhe 1:ri:1SI: decades. :st:ils~ lew observe the em,'lhqJlII ,Ike pl·nee.s=:; fo r <I fi~,l he Jihif' .~uthor h;intil1le' E)@paitl'llent!J1 E:ilrth ami Ili"n~iro" mologi:-;(s h..v~ f~}C1U,,~don ~!:yiM]g:~(leKm\d;i1:~(l d!l'!~.h Sl~u'RlsHJ relii5flfl thm ~,t:.i!IXll!:eti()~~Iit(IWI(t~ I1iml~IIt~1 $~i~nf!et R!lIiml~~f IPo'lyl),d I"slitute. 'llrov, nil'; IN'l' 12180., IL!lifl. ~-ffiiI!il::ffi[(jdr@rpK.I~d'1J tl'ais V"dll'ii;"lbi.e eil~I"~'~lqml1keehavior with ol:~lcr b with f.l.ster s,lip nnes amI. he'nee. nmre fre-

0"

1675

PERSPECTIVES
queJ;H M9~,"\IiI'H~ haw more M9 ~ th;;~!l ~h~lS~ thil.t are slo'w:hll.d:eed the number ofM9 em.1111.~ quakes that have occurred in the pi:l~S'l 100 )ie'l'irS ( five ~i~ widl~none ofdmse expe(led O!I~ raurlam if we mlm i~tn ~1(;COIi.I~H mdy the raicI'lre~li.clecli'L~ tervals belWl;'!el'l~~.l m 11J l, Th~s e ~c ~.r~1l1Ioc.:t 'Ctln(:~IU ~1>:"plf~i.~ hue seme '0 fth~ e:~~r]ier pes iIi';i~ eerrelatl (In!>·of e~~!'th.ql,;jke siz;ewill1. :!!:ll~.p .r.lte.Uov.fe'\'eii.,~l dliiJers ftmdam~tlt<~Uy frpm the ~lt:cJm~ic041 and thermal explarl<lil.ions .• ~nthat the 1000~.H;r pn',(Jict. some

b~dl. ro \il]ild~:!'S~;)lndhe pa~~~[!Il.f{'ln~ exlsrs. t i f:or po~:~cYIJ~IJPos.es. one l:e3S0.ua. we sliaolJld takc a~·""f)y fWL'nl the Sllln:Mrn~Auncl~"lmilin earth(lllOlke is that every :'>'Ibduc~ion zone ispotenliaHy~ockc{t io."lt!~tand dail1gcmU$. To lo~u:l> on Mlffie mHli;lll'ne ethers m ay be fo.l.~y ..
~toC~r ~I1c~ly :P'OI~uk~l~d land ~I~&~. d 'l~d Uh~jPatet:l t i<'!~ iUllilputl~ (if !>~ii:kiil~ iJlp]d tsunarnlis cannetbe m~I;"Slated. We learuedfhat
&:VCr,;~11l~
1

IIKi:ii~d.gl.nbO;liI...varni ~~g .8<yst(;~n~.s.\W shou ~(t by sastained educal.io:n ...embed ll'n.ciessoml

of 2004 iun. the eultural memories of ~d]

coastal ,eomm!!lJrdlie:l'.
Rererr·!Mc!eS.
;1.

c, i(ji~d~~~~e~(1I... A!!m!. R~ .f;~rrh~ilmel: Sri. 3·'1,

!ii)'!iW)![m.

g~!it -:<!rthq~wkl;!~ose a uuique hazard h) p

dh;;t<mt coasts: Title kmgnl~~tlln::' ge~.lernt,ed tsunam i \",,1\'11;:5 Ibm .~ I!<we1ed ever Vi!! st eceans '\iLlh Htde IO;;~ot<Lmpl.i~mtedue W silreacliny. The: gre-m n;"8clli OfDh~ 2004 1:'l:~liIlami (12). .nui. the expected .hmg time interval bel ween ~ld~·~'lt!CnRro(lu1oo:; (h~~~l~~$~ lessonspcn;jsl
OV!:f,1 wide tl:"i.p'o'iIliSe: of li me and spaJc~. A ~nR]ll ,1:~~Qmn {Jfklm.O\....edge i.n.lh.c I'igillt. place l can S,I\'e IliiIm1ylives, us in ~lt.'l~ory of the: H)· yC3,r~oW BI'lii!;h girl w110 had learned of ts.~luil.n:lllli in ~clloo~ uadwamed ~eU:O\,\1 suns bathers iI] Th,.il.illiid to run :b' h]gheilf gr'\1~mCl.

be :i'Il(;'lpl.lble yf h,n,,1ng an M9.\li,hel.e'is the fcm.nerlmMs. tlmnh.ey are

!'.iubdtL~liu,l~
L:~m:5~Q

2. ~. lJ'j\i!di\, H. K1IIlil1'IUU'l./. ~j'j: Rts. ;a'o1I.m~j) ~·li:9'9t 3- t !tUrt liLKi~"'m1Qri, f1i~ ftMIh. ~" lI'l~ )1'3, 2.m(i~iJ.). ~. c. It Sdli!il1..l. r.ilIll111~J. GP.op~. ~ :lia~\ :1'1']1:13

n:!m}.

5- t Rtm, J\li't~I.

mere Iy ilm;lpmooblc" The Sllm<ltrd··i\mJiaman

(;, ·~.SMii1.~.
~20CjJ7),.

rn:1l~ BI.rII, sms.rnm

GefJfihp,· l~9'.~&;1 {l9691Sac. Am. '1, 5279

elllrt~.lqU<lke sur-

7. ~,.I!I!!!'r!!!I!I1"'~.~, W'!n!J,.J GeopIi~ R~ './I" ;W;l'!,! 8. R, Ml:(~fi~;}. GMl'~ •.fIitj_ :li(ljZ, ~2!l3,9 (1997). 9. ~. tifueffiJe-:;.J, G~~IWs •.9~, 6415tl~:S9t 1!!IJ\ i'_MIlIIi1;1f,8r"J! SeJ'r.moJ, :)Qc,.AII1')I. t!9, 1l1S '11'1179>, :n. R, iIM::~f~~ l!.QL. E(g !ra1l1', AiIW S"l~~2} U:<lllM.l!IlI1_ SiuPill,) J!tiilr~UJi!il·B.cl\1!: (2;oo6i. ].2. v. Til'O\l\tl'I"~. 5cirnu ~" 20o!l:5t200Sh :li;l_nUp1l1ri~.na1.io~lg~r;aphic,comt,~~O!OO5J 'O,1Jl)1:M_n,»Q:11;E._J~lmlIli'Ill'llM!lr·~IiI~Ii).t5i!I:IR,mlm~

rrhOOmf!~lY Earfh :sd~!~list:-t by oeeurrlng i:n al1! unexpected pl..,,ol1, El.!lrl~~ gave LIS ,I ;\iaOclik
IL'emindter ofthe' i.m,1lOl:t:ml diITbrcllc:c·hct\lIJeenl improl)~~bilily unci impos!'ibih~y. Our unders;tanding d f w~.e:~ and wh e!:ll the next gre~u. earlh~l,tq.;lke will !I,l<JPPi::illiis in its inf~nlcy a~ best \Ve iKIi\.'eIli!J~had e'1l1IJ~lgh ~ to decipher ~ me Mt) ~anhtl~q:o'!lk~ be~""1\'iot, ltwill hlk~ ma.!1:y num: eenmries, or m<Ilnyrn.orc· q m:]k;es. or

~l~~ )).

pfnl~]y
~ ven

15~;V.illg the~~(J3).

'!N11.~ lewe

dJe\'e~,op tee hl1o:li().gy~

8·0TANY

A h D!"Vl1ene

pia i:'!'l d,ev,e:lop.t'Ii!E! n'~ ~!'Idl Sill ~'I,I'iv~la(l:Sthro ugh a me-mboer or a

In.1llt L':Qntr(ll$

re~Ie':ptor 'f.alllility~ho:sie ether m em bars allle pe-.rva,s.i~ in olilfiimal Ceills,

wide ~:'lnge of b iologictll~. pl.Uc:\"S~e::=:i iUiI. theanimal world. V~tl;llpbys.iQI.ogicallllnc~ t[m;R, such 3S visicu jasre, 3'IlUl oiJ,lctiou recruit G p:l1o~:eiJl-ccmpled Irece'.ptIll."S to relay ~);;t'~I1'I:i:~l si!!f:IiI:li~!'. iato cells, '[f) ~;I ~[he appr~ici priate responses. Likewise, G preteia-coupled reCeplit~r)i medi<!tere,~p~)fl~~S to endo;1olIllOliS sigml ~~ encoded by pe~~lidc:s.. eueleet~dc..., orl·~p~d~ adjust cJ.;[~grmvth !I!!md W~ ..to d 11;:l1enlialiolll. mel<l'bollism,. embryosenes,is, a!l!d dl;:\\l~~)PI'~'H;:!.l~ c~trf~ !~l hy!>ifl]ogic(;li to p d.el1il~~nds.Jhe lin.n.nllrl g'eIilD[I.1I1,c· enemies I.nOn::: tl1.~If! gno G P!:flite-i !l-C~I~~pJ-I.'ld !\;I;:·l;.ptof~. h~ c!o~:UTmilro lhi 5 pel!v~$ive.~.e:ss. p·!.nu::; 5lx:m Ilot to h::l'1'>Ii: evolved s.~lch a dcpe:ll,d.en,ee (Hl. fhese recepwrs. Thege:n:tHII.l:e oflile p~,mU A/~.d}jdr!lPil~is~/l(likllw 'CllC(ldcs fubom 25 '·c,mdid ...te" Gproteim,-.cuu,pJed receptors.-

~.· T

e.. rgJ. g:-l.k.l'il. c.~w~: ~~l~Iy _of p.·.. ,rnl..eii~s i~.-.·' ~la.. ..~~'I. , a 1:'0, nOlsurpnsmlg.~~ ~11YO,~VC <l d nn.

~vl.gle~~iuul ftlr ;~c,lI1iJid:iiloo r,I'lm G proteinoOl-tpled rceeptor hss bcenknowu ,Now !li~.C't ,at.. 01.1 IJ;J~C 1,'1r2 i~1.th~ s :~s.sue (/~. ~r~pol1t~h,n a cand idaw G po.'ol{;in-COlilp;~ed rocepmr 0 f A.,rJJbidofJ;',ds is the recepi~or for the phytohor~ moue ::Ibscisi!l::~uc~d. his is 5<1ti.slf'yilm.gnot T ~)nl¥ because H e...'<lb]i~h~s ~l~enrl'~ t functional member or this receptor li.'IIni~y in the plant world, bur it .1I~5{~ identifies t'l ltms stOnght !lfh:'r rccepter .filol.' an ~ml>.lOirt!ll!:ltplant
d~vt:r!iip!n~~rill!l~. :~OIr~'lliU)~'H:,

I1QW :::l~{l\.v t~~1il 'GCR2 is ,I plasma memIlra.lle-IQc<l~.izcd.(] pmlCln-CGupl.cd rerep~o!i I~at speciflta .• bin.tli::; to lliaulr ..By ()c~un:illlg lly ~bs,d:5lc add. but nm.w l:hcp.~.ysiol.og.i:c~J]y

in<L'l,;liiveisomer (t~:<Lllis.\.. bscisic ~d(h. 1;0C'OIlIJol ::>l>mn<[tt!Ll. c:I.O::;UI'e. seed germinatien, and

~~dliir!lg

,glllll,n!h,

Abscisi.c <lcid servol!!s

<IS il.

pl<]nt~spedf~.c

I rJ, i!ldd :i,l.i to sesen-uansnrembrene on doI1Iii!L 01. (~I nrow~ ins, l1!-co~lp~::;;dreceptor 1m ,IS .1! c~H)~o~ic do:nl~o.lun l~~~[ CI~ ~s agHani:nu:a Iliudootjd~ C'_'i1.ch!lnge· tfildor Inr hel~ri'~l.rim~rie fiITp·.,b:iintl:ing pro~e:ili!s, 'ijG p:lfOte.im;:}. Upon

1\igmll d~!rim!,gd~"''t'h,lrR1'n,~I1Ji .;;lfld ,.rld high clJ!n"1J.!!1it!1!~~Mn'of

[Sf!

i!l ~ti.fl~l~lzo;~ tu elllv:ilfOmnelltal slre,sse;; such as ,o~d.. dro~ltHh].

it hllh~ ~l!i], lh!;;:

phy~ilologic<lli.~~$pq)nse-$. iT eli:dt$.ind~!dl'llhe

.!too

bi~di~l.g to ;;!. ~ig,1Im;ll:hAl f'4;'C(~p~r.r ~l~~~~J~~.~~~ p e......chmmge QlfbO'Llil.1!C.i G Dr' :i'orGTP illl al1l3ssoci~ Ci p~l.ll~ht, r~:o;utt~ if'! ~Wphif di~(lc-itltii:>

£n~on from theG pru~ein •.ThJ;l G 1J.I.'Oi:c:in i.i:5df

do.,~~lre of~e.:1f stOIl'l<:ltRI ~m:r'CS~o.restric:t U'<Ln~ di:S5Dc.iat~'1!l~uo OC:li ami O~iVColllple.."e1i liltal i S1)~nn,[on. <I~jl!stmem of meli:lbollisl'iJ] ww.ler· tm,enw:rget t](UMIl:ll't!re{Ul1l ,emeCl'O.rs such'IS ~te dlCskC'at~fi~ !'lind t:CI~d iiC.f"I1Ipc~rowrcs, 3.!'I,d M~~~m}'Iy.1cYc]::l&C.·pmtdll k~~§L~s;.or rims", phol]pm:l;is, There is 'or!ly n~~ c.-.null!(,<l1 Go:. l!1hilhiliQIl uf$e{;d .~ciIW'ino:uit1Jn limd ~~d[ing growth.llioc.linem.~ci'l1 <'[nellelectrophys.iolog,i~ pbl:sm<l meiL1.1br;llle'-loc<l~izel:!t ~}l'Ote'im; wi~~.iI. ollie GI~ ,find t\li, 'Qo G 'If i>lI1:mll~ts eXI~re'ssed ill i OO\i!';iHnl:IfnSm~tnl bra~le ·t~Jrllll(l.l:lS ~h,u chf!ww::- t'';;!:~sM1~es:pro\'id.e evid\;;~lC~Ihr tx)IJ!l~~~rl!!.Ce~~ ~f11r.lb;d()PNi:~'. Previil;)iIJ!$ Iwuletinm)~1imdysfs:o f te.lrizes: llinis; .r,ecepto.r family. Mm·t\ovel". no~ on. ~:ul<lr..nd ~.rm,'acelhnli<l1L ,percepDiofi "i:l' llhl.c hClr'Illa:~t GI,'I.'Ole.in s;.LlI,1tllln:i~ ~mplic,ned the~,:1I' I'ilone ,~.2. ). :Recoernly.th~ ~ud~<Jr RNA-bind3 imL:\,'olveill~min phy LuhuJ'1ml:t.llfne rC':l<pO:i1 ~~. im.clud~[lg <Ibs,cJl~ic<JIc id ::iigmding (I'l). .11i ingprOlein FC A .whilch OOJ.1i ro~s' l~:mvering
.W1~ iluUIOfS are· ~I'I ~11e·IDeiPa!flJlJenl ,01 ~Ianl Scie Me, ie![hmkal IJni~iIlrsi!)' Munio'h. IliIu!I iHlXh" nger 4. :8 53S0 Frei~,ng.'~'Ife;i~>einMep~an, 'GermOlI!l)'. 'mall, erwin.gfil~@ E
~,lUIll!d'e lhVil~ (4' ).fllld~}l.(;t

Mg-chcla.tase sl~b'l.i1llit H l:m,"(lIted in 'C;I.l~O[Qpl<lSI~5}. '>\iere identlfled as {

o:.cidre;,'Ce:pltifffi. Uur,::J

!TJiU.t<lllilpl;;tnlJ'i

l;lICk illg GO! (o:rA ~J. n;:gul!(I[ tC~f!

il1troceUu~~ra~i>.;ic

ai,

ofstDulat.,1 mO\'eL~.U1ni~: is impai ~-edand gerlm i!llfH ion is lrYpiilr:sensh i,.'(; m iib~c]sIi!c ;iC id (h

IPERSPECTIVES
1:Im.ci11.entH1.C<llion bj' Liu et a/. of
<I pilysi('1IL1

II

GPAI wh:h PlDo: is de::ir<lhu~.ized when 01fP

im'emctiolli OOl"!i'"~(!:~he G pm~.dn-c1Ol~pll;id t

bin:lI~ the

(ii()~, ~Wbll!ll~. 9~- This (

i:-<

comp<Jtibl!.':

rcce'ptor GCR2 ~nd the OOt .sUbHl1:~l G R!\.~ prov'I'ullln.3U abscisic acid-depend~rul release of vides t~1e m~~~.ng I:h'lkbetween .1 G pn1\l<li n ilmJd PU)o: from the Go. :subunit abselslc acid p£:rocI)lion. The :lIUlhQ:rss.!.l{l\:\' I$'a~n '~l aaimal ~~~.s.p~~ru;p~~licl:ic~ci!ll recruits ~T1~ing(l~ial~kinase t(l li:tt:: p]~lS!U~1 i:li! pl~"l~ I;;u;:kin~ GPA I and GCR2.rus:p>l.Jl1WS th~ ~u;ty.~~e to abscils;i,Io:~ll~!CIare ~mpo.air-e<i bUI. 110t abolished, GCR I of .lMwbitiopsis .US;<I c<l:Irn.dl~d1Jle preG II:li.le[J~:bm:alle· artd the:lreby control!; !:.~nernt~on of t..-ill--(.'t)liph,d ~~!!~prnr ..~hl;,':~ig~i~ld (If\l.rhki1 i~ iihi~. ls 0t)!~~h;~A;!U\;\1'Hh[! ~rallel'~lf ~du~I.i~f.i1 1I1lknOWIl. GCR:I i.Il;~eITLc~S nrel: ~:ly wIilh GPf\ .. d 1 him"m.ume zesponse palhw,~}'~;llat'll ·P'Os.,>i.hly iiTIl\rn~v~::;~~_,e iflir,U;!;;II~I]<lr h~~I1l.l~':nl~ ro~'~p~(JJi i~ and functions l;1!nl"igoni,,1i~nl~y~" GCR2 by I)romul~,ng gl!:m%imn·.~olU, ,u.d shonenilllg the clduWlJp 1<1t:-., A.m~IY:iis of 'l!:Jl~IHI'IiI,t I.mas dlefi-· s p ti me to nawl;:ui ng (/./]. Th is; jml ican~.~ COIlla dent in both extnwfl:~h~lar and illitr~c€'lIu:blr abscisic OJl:.ud [ecepltUOIii. sites .':Ohoulclcklllly this pctillg ~JII~:en!l.ctiolIll, (jeRi for GPA·~. hus, .dth T rOi1lilL A ~tcrmltivel)'. GC'R2.;rcl1fttt'lfl p(l'it)OC':im.;ilnd PAl ~ppCi;lr!; to repE~l1i~ ;:'J. node \\Iitem eliI:' nmlCilll;o.l1Iicdl (ii C( Stllb~llriJ.S ,lIJmighl be fml(.;rim~,.· fert':[fI~ s>~n>lni[lg :I;t<'I!th\~~lYS; coJlvel'glt;:. The iJdJelJ1ltif]calion of'GCRl as. all a bseisie alll~.yl.ro1JJ~.dalll in re~3yin£ ~I'~, <I'b!;cisi.c 3ci:d si.!;;1I:. liJlocl:.§, Il!l;~tT~c l'WU lRo',~ln GCR2-jyij,;IIll6-d pllt~le[m:iii i ,!cid 'rt.~eplo:rOrins,s G ~r\Yteill!i sign(liil~. il!rI:(I $Iomi1!~a d.osUlfe plants £hare 42% i!f1d63% id~il~l.it)l3t the ~lmi:n.o the U~1~elig;!~.tor~mnr:;!~'<li1'iC~1, iJ2Ile, s;l~I£llics To m;id lllvel \",'i~l G.RC2 . .amI liM t1m'ree 'p<llneimiare plimt[ !'>igrm~~ny fOCU5.t"d 'Onlt1UC il1t:-rich liitrn'l'ituruUyrel~md,~o~~ ~~UII'l.~l~Ullim:l ~plide-. rll:'.pe'lt roCi1: IDr~ .. The A'lrJh~(I{j/l~j;,·genome p r!ilJfldif}ti~M ha!lthi!Q!~li~1il ",y!.lth:el<,"~~ --UIk# proC ~1fqgod'_''!>m(Jr~ t1~~!~ (i{]O~e~u~i us-rich ntPC~~ tein •. l ill .el al. <l~SOrO'lIncllh<lil the insem;:iili.vi.1y receptors, '!Nh,:uch are involved i~ pathogen of hJr~~!1~lli\'y of mHta!lilpl;;lJ1u~ d)f!tl<~d(:1'$ !:~~l~J~iti(l(lla d in reg~f!lhlig ~l,\!~n~!U::> ~vel.. m d mrel''C:..press nCR.2:. ms;pecdvely. to abscisie epmental processes. :1:Ullconllrnst.llte 'l:I'lkl1Ys~~o:r ~c id depeads on ~.~ 'GC(s\!bl!n~t GPA I." Th~s p redkt€'d~ven-tr~n$!.ne:!il~bruneprolei nr-;has poirals ~o ·OJ pO:lli.l:i\'e .regli~~Iru;ry role of Gc."R1 in beea n(!:gl cted, G protei~:l coup,~edreeepter e abseisic ~id siigl~l. [rnmdllc~it1I]" t:mn:did!!ws in ..:f.I"f}.bit/ops.i.'l': :~Ildude the bl:!!.rl.cy W'~.'ll hsppens t~eDl. lj,\'i:le'[1 GCR2 is ac I:i~ mi idew res iM'lnce~ocus. 0 3111,d heptJ he]ilc<11 WiltCtll by abselslc add'? III [h~'don,.,,~.ic (liilll"md.igm pn)t(lills 1~.;)Il~re~tm!;:'d ~(1~1IWLmmil~.fUl adipoof G protein si£ml'l il~'!:1 (rll. G .PA.~ lrJr' ~md _,G nectin .lml pmges.~in:~-eceptors (whid] posses s:evetl-tr;:n]S;me:mb.rdlle domains] lJ'l. 1'3)..It G~V are predieted 1.0 dissociate from I]le r'l."OejJtOi" ILIplJll l'ecep~QIf-!Jal'.l'Cis.ic~Li;;;id ~m4'\[1llC"115 ~l1rd ~opredh:::~ hm ... m~!I'iy Jifft-rE'rlil. G pltl~ I;knil,. GPA:I mmd G ~Iy bun it s t hen serve as su ~~un-cnupled :.a'eplo:rs exisr in p.1;)~ts: nd :~1(HV a Sign;:liimJg molecules Ilmi. remain tethered to they inighi: :inlJel<ll;~. The limited lU!mber o~~difrhe Il:]flSI11lf~ ili:lJc;mlmbraIDW (t:lrl:WU gil I.~pid me i.~ r~:ure!~lfuielemmcl:ic G prnte.il1 s~ib~lllits, hQW(':li.es)1 (:5(;)1:) the n.'}~!1ll. G~ h<1:$ been Hn,k0d ro eV1;;I'~.~;m~l\fide\; excellent S;}IS.~~IVlI10 fil!iJ.(:Ii(Jll{ll'l a I~)fe:.llIddla~ the spec ific.~ly~udnnteg~lltiOJ~ or , several plant hormone responsesand 3110\.\1111 to iiilllilihit 11'lt~f",:llmm inHiatifl!'l in re~pofllsl;i to [he Gprol.t:~!ill-oo~!pl~d recepmr ~ign()lil)g ar lilt;; level orGp:m~eiUII:S, lmrrnene ~lIxun. GP also lI1f!g~ltuvely[1IJ!gLdales tyD.~d~ic;;~ekl:>'ig!l<llirl!g, dl.!rin,g .gJ,;rmiml·~j~ln and emdy :iieed~ingde~,'e~opmem (fl. R!elieJr-e!!llle:s ]1.. .'(_ lnu t'lrll~,Si!'ienOl!.~15, HU (~oon; llN~d The effects ofGPA I. ]n\lohlCpro~dns t~!l:ltl anUm! 8 I'.'l1ird.200, '~:il(lo,'l'l:il~~iem,!!-.1 BS&82). bind w it. These' include pho~pho~ ipase 2'. A, i(illirtUma:nn.5 ~" Plamf§D1.i31 ,U.412.00·6t Dn.1 (PLDct). wlmkh, i~in,v(jh\~d ijimj s'H)mllt~1 3.. R, R. ~inkl!lfo1l!~I'iS. :5:,. G.llm~L!J. (. [l" Ha(l\. Plant ntl:tfl .• responses, and th!e ell pi D:l1-dm,u11·il n pa:nleiln ~IiiPj!!~,)\ (2002~. SU

lllil.c Iipids:phi ngesi nel ~phos,p.~f1te. SiPlm ~n~osine I-pho:sphu~!.': then <1ICl:5<,JS all! eX!r<lceilublf ligand [0., Gp:rot.ci.n-cmI3),.~cd reccpiers, ]n pl<'lfll.s.. ~plli.lrng(l.sirlel ~phQ~pj~~:~re lJ\mn~oues G PA ~,~d.epcndcnl sIObl1iat!l.1 elosu re (I O)~nul !1liil)' .lIso ae~\.~,~1 G Pm<!~!k'()~:ph;"d ~ r~OOIlIO:r.

or

ru~we

AtP i.i~,l I. wh ich Ii.H,m::1ons downstream n of (~PA[ i~l seed g,trmi)rimnK~fl umle;"dy ~ed~ i~g devi1:l:op]1~,em. [n addition. t:~.epl\Qtdn illn,y~

I..wid fmll!il.'lti:o~ I ~nteracts \viJh (iPAI .md

I,unks G pl'l)tdn s.igJIlaling to fI: SlUg<'IN:;eJm:l;]ng pat]l).l''',I'ay. Arn-cisic acid-5.T:imu~;l[h,;d phosp.fui.olip,.id t:lea'It:I;g,~ b}r PLDu gener'~les:the s:i~n;Jl~ ilm,g~:,n(d",c\l~!~lm~lmtidic p add ]n dl", pl;l$m3 mi;;:I1I~.bra:L~(!'. in.osplu'IIid.uc acid fHrlllCr !.'l\!!o· P mol.e.<.; illMl~usie llI.cid, sigmiJ~ng by r;'CCmiJ.tng J~ f\:m 1.<1 li;eS~lli\~ lI10ir:lI!i.lm.leresporlse llegld.llo.l' :~ {AH~ I :UI1L'er-iLC~S. w.~th diflefICnl odlu~~I'~IlI~ClS .~ such .tS lhel]:mt5cr:~pti(mrncwr A:!HBi5 to Ilieg~ ;:: nl.!'I'¢ly rog-~~ l!l~c;)bsc~sk .fldd FC:;'Pt).~,Sles). to 6!.1ard~or! h;Dm!Q~~S..
at~l~,pore '~[I a

h~a,lis (()filVO lid by

(1i(lp)

l1M.~p~rtlJr~@r

<IS,tOIl!1'

OSlin DrfglU ~liolll (Iof

tldi'(J Ii] Ulardil;eHs" Wh~VI '~!It~ PflJ'lein",aupi~~d G o:~p. tor 'GCIl!:2 I:; inilctivt, the prO:iein phosphatols,e ABU inhibits sloinutil d.o:l:uJileby attenuating a,bms~c a(fidl ~l:rJna I U ai'l·SltIu(jt~i()'n1. (BiO'UOIiliI) "ypo<sm.l)ti!: C~ndi'tijOfltSi~gluardl cells (rclali~e~() suntJ,uncii i'!!} tissucl (aH~<~ ~.kfm·a tal ~(I ~Ij)~e. This iirn'l'Q a(tivaliOH'l of l~'e$ G(R2 by.!lbs(is,i!:: "Qidi" whidl '~ng!g!lfS :spe6Ih: sign,j!I.~ i IiIQ pa~hwa~:5 that. ~eiI~to d~os~ )"lOfilata, SGln njl1l~ eLec.lmm miUlograplll~ ,ai ·stomal:a a~e<from {14~.

<!. ~,A,~:em, A. ~!"trere~!ll)';S. !t.A~r~m.~, ~ilL R.D', 1II'alu~ ;U?, 2~1l' {;rog~}. 5·. 'i': 'I. S!!.tn f! ili'J" W'Q/f!I'e ·44],,8,231 (:!OO 6). ~_ A. 1M.JQ$, S,M. As~ilItafin..£Ji!6..o'~. s; 5H (~~" ,.~. i\lilldf!<j', [, G, C~I!I'I, A. M. }omes, S. 1M. Alim'lann, plllf.lt Ph~itJ1,,:II.41. 243 t2:006:), ~. 'G. Ml~'tlr.", w. Zh<!~g, F; Oeng,. ~. Zhilo, X.W~~g.xifN,e :11:1;"2M (:lOQ6~. ? J. Zh~~.X,'I!'IIi!!'Ig,j, 8Wl (hem.. Z7'.n9~ (Z.oG4). l(l,~,(Il~.,gl ~tN~~~~~. ~H~~oo3l ~ 1]. l. ~~IIl!,hib~Q&i,A. M. ]1JllW!\,. S,M.~m~ilUi1, C-!tl~ Opl.n,Pi~ro!!l~i{lJ.1. n~ floo t .. ~]Q~. . ilemlo!.'! oJ.,J,. MaL trnl, 56, n (Mum. A l!3. hi. iii. K~ri\ H. tI\ (jooomllo.j. fJp. lll:li!:.S!SIU37 (,005). ::1.4. E. Grm. 1-1. Zil!g:lf1r, 5dll\iMl' 2:82. ::152. '(1.9 99J • r~~!i;i!1 '~~!1~ 6 MilI"(~.iJ:001; ~I
1.{hlN6I:l6!NiI~,1lll(li]'!l1L.

'Ill ~h~~'!<lsma nl>emb~lale:: this rei oC~'dit:;f!!~kl!1!of .~ Allarl blm:k:s .~lS. ""i ~y J. lIIhe iIl.~rd.cd(}nor <I{lll q8

1~~hlde~!i~iHf{llTrl.a!i!l!n ~e!l citi~gl~rniS p~r_

1671

PERSPECTIVES

A.

{j"

Mac:.lClillrmh:l,ill Cnerni~t ~\lh 0' (Q·diis(ove'rl'!d

.A'lan IG. MacDiarlmidl t'921-20'0'1~

:2 00 1'. Alan G raham 111~e:1i' phllyed an indirect role i.l,l his Nobel rriz·e-'w~~miing c(ldiscl[)\~r}l ('if 1fie'l'fullk arMaem~n~~id .i~~jill PhH~~dl;~~phi~.m d _ age 19 .• I!;]cr l"lilmS"I,rhen abeut to ~al'li.c polyme:rs. dep::ut :for 3 visit to his Il;<l~.lJve New Z~<Iblrtci. Ablu earned ooc'l'orntc:s Ifr.cnnthe !J ni\fell~i ty of Wi ~,cgn~in and Caml.1.br~.rJg4;l Un iHI;l su tle:u-ed from a blood diseuse dH'~W.,.5 expected 10 soon e~]dhris I irIC. versl ly ~nd bu~r beeame a. p:ro~essor at t~.l!C: Aim] N.lm::D.~,.rmi.d.whQ shared ~he 2m:1IJ UIl~veI'sity of Pe~lby~v']rllia. whe.re he made
.n."1 february
I

1IIl@[ilHic '~Jlg~nli[ pol}'lfTlers. hd ~ r.u'IDhnmd $E! nse (lJ rl'~Pr C!!flSl'biil iiW IC!!wa rei iindi"lfidl,l.;;lls a Iild irlu1mi!F1kind ..

Nobel Prize

ilL~ (,I~cmistry

with AJ!l1il1.Uc-ege.r

pio:[D,oel'ill.!f bl'Caiklhmlllg,hs

O\limlHidcld Sfuirnkm.. ....\. bclic\f",;1.1 lhfll \v!!lQl]ng in &.moon d~~~ni$lry ..\\ihid~ the prize g<I,vellll:im ~i s:p!l:ci<ll oblig<ltiQJ,l for eaFlll.edhil'lml the 1.971 Fre<IJ· fud\l'ln,f1;e~ng hum.tnkind.liif' jndomh~'bl~lq'li.ril Ai~!'I V(lh!!1I~liJ.erlild((I ~(i' erich Swim~~1 Kippin~i\\~1 of [he American Ollemic<lJ slone into harm's ~~1<1y and sense or:l'"es·~l1sibi.~it)l never dwindh%l. H,H\rill1g b~~1'iJclose ~udr.l~uh 2 ffiumths e a rlil;;r :il!'! S{)f.~iely. In ,21J02. he beI.~eli mi mn'i<: !~t:: d <IIl~er. China. he "!,IIiIS back on ~heroad tb~sJanuary to came prefesser of chemWi till <In ~~l~enj :sa~:~:I:.r ~hilld WBlUS, he tucked ~uDCIl~lrlm.;nt of E!:u~rgy lahm:~Ul)ry l'md ~1is h;:try alld .!raflrll~-~ 'VOn Ehr iIilJ:.tii;I:~.t(;:· thelJlni'l,oers.ily orJe~",~'8 ,U atDallas, D~::>~ ing~l.ished C'.I1a.3 iri n <I1~o1l.ebook~mo:lmi:; tilxmsHis p~'~!Q:!n!,;,~d I.O-day lr.~p llO N\i"'liv ZC'l]~H~d Sd~M.cC o;tf!:d 1~c:h!lolQ!fY ;;ill ers to f~infflrcc p.mtt:cincluded ~ l:ilre-su::;;~OIi~.lillgblood lr.J'L1:snH.sion, tion a (worcled b}' bO:II1,~b the Universil~Y efTexas at Dallas .. di5pOS!'l~ clm~.fJng,aml Il]rreekCjlllme~ec~.LIIlies .. discussions '''hilI gov~ efll!!tI!e.l!llmin isrers, a TV :i ntervl ew, mld ~. In the I970s. there \\'i!~ i!1~d ,! ~ls]U i['1grod ro Q'~Jl .1ilkely ~~Mtr:EIre\,'I::(!~~ lam i Iy, to mnch excitemens about ill \~dveandavert a porenA 1<1~ 1;,~~g,ocmll:~n NC\'I.'Ze~!I!,mtllOJ) 14 Apri.11 new~y diseovered kinds of I i~t~. <lt'lsimphe. c i 927. The rnv~llg,.t!s Qr~hc a~:lil DC1P!:l:$StOIl :t~n:tall ie IlfIJUIt."eu~~~lr dllll_rge· A~~:!ll was ,;I here i~1 guml~f~!lg-·t~rm~ln~mp~JOJInlt'l1[~Jr~is . ~~I~i~J;;'i;l,n-;:un~f~foompl~-..\it'S.an ;l~~ lm~ny Ol]n('r'w~ys.~]eere~::'it~.l.erc;msedAhJUIIto live in p{Welft~1 uri:rng his d m;ed iustitntes in New in ·\'1~.1ch Ala~IHf.lege'r!lnd i if; l;;ind!'il~rk di$~xweri:~:-:. Z~ah~liId,ell i:!il~.lfld:ill. Sou[h Ki1lreil. Bntt.il 'yOtU~, Vel:h is f(lm~~Y1is~isl~dotJiler LmJ.~p(li\r~r- 11 group t;,,<eJl,;;n~~aking i:shedlll~mi l:i!es 011> est they t:'oukl. llih is tn:adi~ b M.<I'~Y researehers were tryi n,gto make .~nslllat~ ~mdTexas •.wh~Jch, e'l1.1.phasized rim! illg sustair» ~:iq!l 01' helpi !'g '~lt~l\:r$.'\i~£c~~ hi 11,did i~~tl \ ... \ il\g org,~tlic p(l:~}'m,eP.' 1'i11.(!lnnic-but ~i~h~!r ••ble wlu~i(m5 for (h~ i~cW1ltoing I1.Q~d fTtJr ene n~. ChJl~Ili,g CtU]tc.Lereucep'n,~lter sessioas, he m:mj' fields and OOllntr~}~. ~s <In in~p{)J,"t"'i'll pan with tile'lI'OOJ:lg polymer m' l~,e \'I.'HJ"'~ OOP<Hlit of h is .icgacy,. nu;:n, M0111Labes ~liIld en-werkersat T~Tlil,p:l~ w()~lld focus on lite work. of yotmg people, He UIl]vlt:ilfsity showed metallic condn(~i'!{iJ:y Iovould listen inl~erlsdy 1:0 their presentarioas, A Iler cle~h'em;i~ milk in the eOlrl.y momin.![l 01.1 %~ bicycle, AI:Thn be gall primary $C~flo'l, d(l~1itn 1.0 4,1{ filII" lite- um!oped iUOl]:illlk: polyask pcl~Dmtin!:! q.UI('stiol'ls.and .hen l'I"{lIlltt;;nflty ;11lildsincerely wnUln.e:a:u~:Hmt <~}'Iom ,"''ark is so classes in :I. twa-reem schoolhoase ..where mer] SN)x- wi'lJ'icll can be made fffiniJ: S''IIN.. "most of Illy sehuel d1lJITIS \~'eI.'C M:!IJor.i boys. M9.cHi9l!"rn,id and [ h:1!ger beg:.'11lto {;.·oi:II~ll)QT'<li~e veliY i!ueres~.ing."1"11i~~:n.d other encourage9 t1ind;ir]~ :iul1l !i\i~(ji1fl1l.11e-'lir:J!111t1.\lu[Jmw;;I1t' ~ h OJ;] (SN)II:' ''lim) fbmld [h,n bmm in~ dctpiIDIlgpru1t!("U from such 01. ~rem :>ci:;:miHt ins~'lir~d £,~!'I-' tn ho.UII.or ofthe mnivc M<toll"i ~1Jj(1 ~1,i Ne\.w' due-cd II IO-frold. illOl'~lgc ~n eleetriea] camndm:s erou,lons. of :>l~IA..~!enls> tivity~Wh~le visiti~s l~,e Tokyo .1::rnsl.inule Ah.m ,,\;'H)\eXlllaerdllit kwi!ll,g. seulsilwve. and .Ze-,d.ln:d ~le!dtrlS'Ii;'. AI!lln c:hrooed the M'IOri I'mhl at the: !)11~nigbl ~'iud~:UiltO(1:'icbrati('m t:1:ltdi.lii,g ~he nch~'o~ogy in~975. Mm:eDial!'mid ~hO\,,~d ~~t_iv(: in ~t>pcmding 10 ilmd~,'vidu~I~.umanl N~)b~1 ['riA: ~c:~!n~i(_liti~S. ,llld ~~ Ill,"l}' o~h~~ I~Hdl;:k~ ilitlk.iiW~1 Sh gol d~~ 'C[ys.t~b; of(S N i,r' !l~~ds ll~'iIdt~o;-.,:;~ f l~u~'iI.al1kJ~d. His i!lsightz.; p i O! ml Sllri mk<l\!;.'~. shov.'Cd :~im r.ilnms of hi s sil~ 3m] disclQve:wi'e:s:. bOoth fumilmnellt31 amil ;<up~ celelmniol11s ",.allirl frienclls. p I it'll Wl:~ ~ fi't i!J;o:!i fVJ bIl;,'gi!miID!.g th-e ~\"~(md A hu~.~ltiw ~lf I;.~hemi~~tfY g~~!1 l:n.. .mi,l.gfill 0.. WI)' puiYW;:~lyl~!1~ (C H I S~aring.! v ~t;;i(~!1 ... uf \:vhen l~e s~arted. too ~'e3d i~is (hiller"s ,",he~nu5m~ Ip.oill5Ub~,y tr.lffilslbrmimg I{"i~])x fronrnl ,mimmli'ltm.' age of p(}~Y:lmle;::'.r.s;.wherepolynu:r.; become ~e%tJb(l!)llrom lh!IJ Imel RlO{f,,;;; ~.fl!1i prtHl!iI<lA .iruo .11 po:lyme.ric n'il~t'll S~i~l;lka\\f'o!I joinoo fully fumaiomd eli(l~tronic m.~wlI'i<~h;.. nereJ tlq]ie.ly len h~gb sclloo1 m he~.psupport his J~Ili1- Heeger <l:ndi :JcOiarmid ~t tile" University of M w(J~IId~'li\ie been mtne iI] st~llUU::St1ml he 1110iUlI::- :i! o i]y. ~md l.o.II"Hula.tdy found a p~rI~[il1i!lc ~s job PCllil1isylW'llni1t 1.0n~~Lkc:hiSiIw~ppcfl. t is:hf,j~lld 1Th111l}' more fu~kthroug~~.:s.ifhc~~'ld ~. I<lb boy 3·mJ.d j.ml irQri [I tIl'!'! !C~emistry de'pju!~ l1n: di SCOV'elf)f IhHt iI],-m 1:Jli~.g or-go:lnit::po~,yE l,j\redju1t.<I luule ~JQ~e'r.He b. slnvi¥(."\J. by ilmis ~ !1i'IIc;~l ofVictori.~ Uniwr&iw ,OU~g.Io;l. h \'l."aS ~l.1;r& C:(lll~d ib(l C>Qrllwrl('d il1ilo mClalllic CQi~dl.!CE w~f~"~ :>i51~r.and two brothcn>, l~isli)lIU" ,chi~- S ~ h~oo ~h~1he,e<!n1ed hL~b,teh~I(l:r':;; 1:1.nd:rnl<)sle;r~ mrs by dupiimg ,~rnil>'d Het;l.~r. M~~cD~<lrril~i~L de·gree.~ ..l~i~ first fl~.blished ~per •. in :11949. "Ulid!.sh]mk;B~\,<I. the Nobe~~P,liize ill~ C'hemisl.Ii"Y. R!eflE!iJrel!hOI2'5 and No'Les h ]mas~.u:l ,e~1(iml0usfmld;imt:Ilt,!~ .mid prnc~i\!\",jS,on lhlii cydic S;1jN~mole(:Wile.. \'\il[dl mm~h
o.

sensors, anifici<~~.muscle~. lr.n~sistors. S.upe:f~ bmt~!'i~s. ~1I1d li~L;;~(,1~1:1,~ A. is raduate s:rudent once rorn. ~l'lto A~<In's ol[Dce s.ay iug lh.n. he lill igh~ have accidenta il)f c end ~~1 Mold '1.C'e 1)'1erie. the~;eby po~enl.i'IUy cre'!uing ;I, powerful bOlllb. as Lin: .'I:ccl:yl.cnc g ••to prel~I[S(lr to K"H}:l; beccmes MgMy explcsi.vc when liqucli'icd. The fI:~ d~1,,\l'!1~l~Ul\~S c,dh,;d and the builk~i~l!Jcle ..red.
rC~pf~i.I!l;P-'1"

or

I.

ca~ i~lljf}Qrlil.mI.o~Rea] ize:d o:r p~(l'p-ose(IJ .. jjl"odilheil!:lithllr ~ilIU1~ ~ilmo]"eo;h Ini~iI.ut~. lLI'nl~;nity

1. ifh]~QiNiII@ i~ IKIIiIIAIlll m's. ~w!nb109t~liIltl'l' onillil~ rfwl

of Tf~~

~II IDa'lli!~. IX 75090.

IJt&~!M,ll'dlJ

U')J!I., E·m~~:;.r.'l.~.b.~!J~1ti 1lIl1l1n@

~'l;ti:~~nC I"de. for e'.";;lI!rnple,'lnt~~toltiiC ,l'nd (QITOs:iOI1l,·pliOtect io n gmlteria~~:s.elec lr'IJmagn:e!k: shields .. Ugbt-,~~niHi~~ devic~s. S!~)I~r eU5~ ~

Web Mle ~lJIl!beliP.riite.orQ!l'll!DbelJprilBldl~mi'IJ}'J ~uPMe:ilo!:(lIllOOni!(d'j!nmiill'~!!~o~i!l.h!JmU" W,1~OOtiernct'.nllill!U3

I
:Ii

'"'
~

l!i

IR'B':s,t,o,r'alion all 'Ihe Mii:ssi:ssippi De!I'ta::Le:S:SO'IDIS from Hurricanes K,8'lr'iilnal and R,ta, i
~lol1l"1\111.Djlll'l'r.,lIiO ID,tm~h!1 f. Boe:s"',,~ 1E:~U5J. Cla~ri1lillil,,3\G. '01.11 I, Kemp.~' 5 hlh~~y Ek I~~$k;\ Wmiil md. Mi~'~c:: ,~ K@ll!rleth, Odh)' HtI:ss.i!,!1,Maslilri,qlui.~ O@il!llSE! J. FI,e:ed,? s h L~'!Ii'a r'd5i1ablilili n.~(I (h:arles .A.5~mel!ls;t~ ~~~BmJ. S~)r'~@v@r,:~:I; !R,o iJ~r~lit. 'mw~U~y/ (h,e.:~te'r C WaltS!IJ ",1:3 Jolhllfl T. 'W,~Us. 14 Oe,l!lllfllsF, Wl1m!l!halun1~ I-iIlIln'i:l:ilIFli!5 Kilitriinal andl Rita, sihoweiti~he vU~I1~r~bilRyir ul~staJl wllllimul1itjie~ ~Iul ow human V h ~(tMli@s;~hoi!l (alu~oo (jleOOrioroi!lio~ or 1i1,e Miss.i!iiipp,i I[)e[tafic Plo11~(IM,I.)P) @'';,i!!:~rbat@di '~hliS \ill!~IfIIe;r~b'il'ity. hs MDP J(lrmE!'dl by' d~raaii1i!~~ f1t@,r,ac:t!iollls I ~ bet~~~efl nver a~~ C~NIs;t at. VCl,fii(l!Ji; t~mp~lfall alfldl spatial s(all!$., a!il~ hmg!l~mtvijty has !iooll!Ci~d tih~~~ ~Fil~fa(li(l!1l5 ,!It aU s('.i!,h~s. Res:t@ra,tlof!, !~nofl'~ (lim 'lQ IfFN'$i:albUsih[his. dYll,pllillk: ~11!t.e~aclj(l:!1,wHh >eil1llph,1'Is1$ <11'11 r~('OI'ln:tH':ti I'lgtn>e rive'f t~[ne:! d,~lMic .pLaron. SE:ien(ie rnust guide' MDP' re'5t~'rdiU(m. IO'IIhidll \lIiWlIProvi:d~in$;,ghb irnto dlelf;3 re'MJDrati~n ,eolsewher'e' andl gen:~ra~ly ~il'1ltD coasts ff<l(ing dimat!E.' ~hang:eo ilil times of [l!11Smm;:e ",ca~dw.
~l.e Mi£. Si:5.. S TPi. D.C'I.~3~.JCr.)~,ailll (!VI DP) lS .[1 25.000-knll· d)!nl~mic laindsCflpC of water, "\iI.!tl~:mds. m~.d.low u~li1nd ridges. IOflllC'di 3~~ series of oV'C'I'J~p~~ing de~!J lob~s, An llULicr.-;lI."lIWiug ofhQW .tlHHl.l~I1i'" and HurriC'MK"S Kanilllt and JUm ~rfTcctcd tile M D:P i.nI.

01' olki(!, iii, er sctIi:lncms II,,",Sll~lX'lliJcd rrom. bay..ruld ~N IlL.... n1dl,{~:re ~u~r of !VIC!! i(m or crod~c1 from o~b~, \"'("!~n\!~ is ~~'Ci ..~.~yn~p{1l.L~m i d~lf~
irrg,wil111.1,lt; f>~I;l~ U!1(3,h.l[lrriCiurw:.\~,7. H1, I!l. Htlwl

To f>~n"i\'~.the soil :iimr;L~l: ,p,r ~'(l;!:'ii~1 \11.'1;.'\!;Lf!~l{;;,m!il~t~m,\ Vl;:rii~O'I.~!y k!C1;lIP"Q~\~~lh to klQl~ ~!,\""L h;"':L~tl!~~:;i :'i cr1!k'i11 il~I!l~ M.DP,whc.TG ~ !l,Lulogic: s'IJ~icl,,:ut1~ C'tUL~{': if. fL;~kttiv~~"lI-k:vt::J ~ rise ~R.SLR) (] r abtrnu~I en }'~ar,~ ol;}ma~:ltId ~o -I ,:) HUilfyC<lr of 't;lL~Wlijc SUt I? kul I J!l,mw~h ~m'lw~mil~ ()!'"!§3I~it;: SC1!i~...! Iht' [~':.I or~}j(: verli~.1 .gfll:)\vl.~i toiii.lL:.';S, rmrri1 !'iiii~'iiCi!'iI~ &':tIU~li.."J'IIS ( I). [tdv~rin~ ~ lil)i.JI::; bt..'fl!.!Il •. (:00:<'1.9.1. \'I,1~llrilil't.h;, in :>(:y.. '-TIll \"'3,~: M~fibCr.J1:.cdhui."inl,> increase ,ftccj"(:~toi'ii ~i1~dulk Ja~il.:r •. nlitricnr.~ eld]'IiI'l'!:1.": ~am~mwl'h. b p ~~l.lw!l!er bu ~J:Ci'S salx\\" 1Cr il'lII'lllsion, row iron !'iI.'OCip[tatc::< ~~xic :-.lJmdcf. ~N. Q I, Tile depcil<i~ ton

C'!!JC'L".mo....~ .'l,dm:cnt is imrOOll~"'d di~~CI~)i lrem ~hc I'aver 0 2 ~ !n 'Ii~~~ MDI?; ~ier i\il:"nd~ ~rnw .md d1i~!lin~l U Jt C~lrsef s~dum~ll" are t!~po~itcd :u m::'lnvc: ri!,'(;r !ilntlluhs, ,auld :.ts the ddn:rLaclvIllICc,,,, Sf.IlI:! is 1r.tll.'l;POili!OO Im~ny .~ form beseh ridge::. lU.bc:r. cb;lIljlc~ .a'b:JiDlclkmlTIcnt,d:cJ11I,..l'mm~IRls, src It ... . wo~--d IOfooo erosional he&l~ml(b;.tlullched !('I ImiBI1!!:': bchi Ild, 1411iC 'b:!imcr,Wa¥~"S[llld C\Ure~11 s ~wt(lrk and It'ldislfibl..l!C'~I(,'.OLdbDld :Qn,a:.; latct[lily !t) foom Hwnking h(ulTi~'riskmds, ami Ihei;;,,!m:nds M.10VC' Iruld~,V3rd as .<;[1m::li5 ImI:lSPO:l'too in washin 'l~I~Ullctiolli
will] cl(l~I"Li;:l~cycJ~

[Fi,g.. 21

t 5). M:tny

or ki IQmCil~us ror dd,I:TL lobes dislrilbl:l~lrtc. ... fUIlC:tio-ncrl, C'Uthcr ':n!..'1IinrillM!~Y or :scaoonally. :111 til!! b::,g,il1na.,y 0:1" ElIlnJ~;Ul (okm17 ...l.OC1Il. ,aIDwl!! 17M A sJ..rek'llal l lj;mllC~' Dfk. nf diiislribLII.11), ridge;. al!~[]b~cr i-ilana$,
pared
1.0

hundreds
rm1l11~

(6}P:Uh;:c~(J

.imericlI" ,nb.si:ICD: wetlands :rrortl~ Il1Iru~J1.e

lOGS requires !oJowl~dgc' about IJaC OOnll)k,,'I; fbrees nnd sakwatcr i~luuskm(FIg.. I), rroOC>S1CS thai r{)m~cd ,mld sU:5mi ned tMlC dc~tEl for !'l~ lenn ~j l)cfo~"C hu.~mn in![p3c~, Ute' dc.~L.'l i~
('lliIcr-gcd about
i,)'lIl'ilillic

:S1.1b:il~ t Fvg_ I) II ill£ld A \1,1r!lL'Ly or prru.\.'.ilNC-i rh:m~(;t1 and Sl!l'illl inoo Ih(; dcha and im~rem,.cld il;;; o'!l'(;ru.ll~. ize ,(Thb~c I) ~ 1(·II"R.hrCJ:1,ltc scdiunc:nt~ were (kpOIl.i.tm at dvcr nm:llutl,s; and via O\'('!~bf~"knoodhl.~. cl'C"\:<!isse fornnmto!lJ. alt(~ older di.~~tiihmaJrics, ll. 31. Crevasses sealevel
were !JIsluallly sh{)!,t'iHivcd d~:tli:GIH.'LI Splo:.""Sf~bout

,(~ooo10 n}{lo

}'cm1',~gn u!,k::r

-n

«i'oo

YCi,11IS)

!!ltd

[DI1I11.oo

111 km

w~dc. as <:O"I11.~

S«I~. ~ii'!Mrsll.)'ilir NewO'~. W'ew ~.ILA701~8, U:SI\,.'~Oi.enmllg1j RiMlf Vi'el.ttund Ili5f8IDIJ Paft. ~ 00.;0' Stlll.e i.ii~i~!'$.!!)', 3$2 V!U [ioQf,jdge Sl~et,. Cbl!!mb1Js, OJ! ';32:02. ~DE,Jlltrnernler,(M\~ EFmrt!!Tirii~ EDliimem!i9. ~iiRIi' ~St.He Il.IIm~~ty, lIa!om, ~, III 7tOS'l16, 'lISA. ~Oep.;ii!lmE'l'lt !OO' G~ ~rnd' ij~~ Il.!Imiver,;;ity 01 Ni~ O.-eGm, Nm Orlea!!'!!~ LA 70 U8~ 'USA:.. ~e~ur.cez rot' t~ IFl!Jitu~. 1616 P St~ Nlilli.W~11iJlg~m,DC 20036, USAn~ool (If II!liUil~C ilrid ~she-tIJI' 5ciill1i~@s. BD~, 5502Q. !1.inMi5iyal ~ Wa~hUiJg. to!il\5~~[]]l.e,!III ~8(115., USA. 1Zeu_))jJ~Oril'~Ori~h). \ ~ omce ~ ~966:].a I\~ldln~r:" At;995'l9-tl6~.t USA. UE~gil'iee.ri'ngl Rli1liiilKIi Cffite<r, GlIora&l Sta!e I.!Il1i1i'o\friIW., f>on (omn,s, co 6'01S.a.., U5A"vlrgin1a 1ii!~'it~lg ~f Marine ~i~nll!', BID:: 134 6., Gloo~a:nl!r P&imil~ VA .:!306,:!, USA. l~£IfII1~oni,am ~mironnlimlilil. R~~Hh Cmcer, 800;, 2!1, E.'IiI!)Il'ffa~gr. MO ZiQ37" US!'!. ·Vo \~hom oor.~e~p(!l[nl1fen~e illlIOuI_dIbe add,re5.'Sie(il.IE',rnait
jclilm EI~.y®tu'I!d'1!I 'IJ:9!." ',III!S{iIJ,Hlelor Wa1el' It~~, lUIS Jiirm)I ~ Gf Em· QiDl!@15, il71J'1]@!@g~~~ Rwdl. A~D1dlii!i,llA 2'2315,. U~

D~~Q~n~ ~ntl!'l, U.s; ~y COIJi!!< i Eng&-i~. 'i,lioobt,!rg, Q MS,3911lO. u:5A ~lhe flmtir.lll!i! (il?n~l. Lmri5tiJ1lIiI slam U~ioreffiilil'; BUm. ~, lJI 70000\" USA. ~D~iilmmt of

lDewal1n'!ernt of ~e.l.lllCl1lJap1~~ am C~1Ll~ S!:ietM;e~ lOui:si~IIII' Sta'l~ U~;y. bton R~. lA 70a.o:~, lISA. ?UMoe:rS1\y~ Mar)flllJdI(eri[fl I'u (nwOnnwrtal SOEDrf. PosI. Of~e BOil: 1'75, C<IIIl'brldlje, '-10 lU.U UiSJli. ~1E.ir!gimeer I~!'i'(!l and

IFi g •. 1 •. iFhie MDP ~~as f'orimm by .81~erij l"~ (]If ove rrLappifll!j de ~ta~Ob1'5<lIStill' nive ~ 0 cIl:UJlt~d, ,~if.erll!rlt f (tliiT~lltalld abandOfl,W Iriv,~r chafllfie~, bari!ierisliHiidis. g nd e'';.t:elli~~Vfl: (1J~~lal w'flila'n~5. (urrerlUy; ,about l.·f(H.ii1hcl:~ of is dis:d"arged vital tM lOIl!ll~r Mi5Si~:5ippi ,ci~re(t.~, t® the GLJUdlFld on~,;lhiinj]i~ dli5flil<llrgoo'll'id Ih~ At:th:afa!aya River to il shaillQ~loI

cllii ~liIels" 11le-d~t~a liS(J11!1U(i(ooriz.'!1!d by

II ifll:!W d~I!t.:11 J1or:mil'!g,. i~

nQ~

l;Iaywhre

lc{ati'lll'I Q:f tine' MRG,()" The turbiid p~!LJmle shewn !:Iefore '~he pH!~Emt [Moirliirie,m tram 'W6))

The 1o~~Hof!i

@~

Il~l/ll:es ~s S.'hQWfI (I'f! the WWl!,f rr'll~r ~~ Wl~U

the right Ire5,ult:!> hOFTIti
river

o:!:s

th~
BP',

,tin!

dli~rsion.

11679

over :liJln5. As \~'::II;)n4; d'etc-ri{!mJil;!!. a banier i~~llKl m:c is li;lr~n.~. Over time, jhc bm.i~ !rn,gnl~~
ill!O srIID]~I,)"f·s!!iI~l~l.1;,. i ~l{i
.~~

~~t~j1.~~\~ l\,,,~hpl\.'~r rer~

oir ..andy ~tbml~S .arl; tor~ll;\d i~~~h{]!n;: ofih!ii i:;;~m](b.eVI,,1I:lIHuUypT!!Jd!llci[lgru ~~tbJl!IIL~d tymIiIpl(;". 'hQIll'l o:md i'l:I1l.'d !'.heI.'L'L Ih~)1I'rcx:t\1.'S con-

or...

~i~ll~ <11:110 the

L1nti~ .. a"lOlh~r

dil'.tI.ihW!lm)' dimmd ~b(QI1i!ol

iJ~iri,

c)'tk

bI;:.~LI1.~

DI!~eWmQMlt]OD1 of t~e ,M'D,P'

the C(IJH;I:ruC'~kln el' imcr'K)~mdll,lCnl~ hove· jJ~~I.!~d the hydrokl·I.':Y Ilml i'IIJ!smin.";.lhc ~~g~n t';O~. Sp!JiI b.;:Jnll:;" ~~s~ial~~\'\.'~i1! 1;:;;111 :~;;; [!I;Ir;J.\It;:~ .. !In~"'i Ilrn\ of W,r:r~f ~lfO!lI:!l1;hw~l.~nr;l~ (1/), Dt1Q':!, sJi:nlighi. !:ltHti.g;:I:fion~:11~1~Gl[i~ Sah\\';llTLTir!in1:;iori and !Ilt d!Ot~~hof fh::~h\'l.l...ll,,"i!· pl!:i!'!! cmmHl.mnil:ics. (17), QIIIL;: Ilw mG!ltu!o'!:<lbk i'l ~fl~~ M~M>i:iisippi Il~RTGlillfOlutk~ [M R.GD1.;J Il~by-300-niJ ~"'IlW dK"tIg~\dliI:inlllgt~ lhe B~.o.rrn SouiT1d Bflsili .UI'i %3. J Sl1k",'!JJ.cr blhll~;ijJ~'!i'b tlte MRGO kitiliXl. rl'lmi-

wetland surfaces. hllll.ping 100 ollloct RSLR. ;)nd

~1~1",;Ire

i!li"!llQmm~ !l;lr Ulc ,S\I;<;lai~lAAb~l,~)'

(/', ll),. ~IlIJ!nilf;;ml.c~ K.,t!ril~<1 ~l1d [Ril\;;'l\\'{J1"l; Ihe ~i1!~~·~h fij Rh. 1110..1 ptil'I\ ~'rli.lll :\torm,,~r) :\<i:!nil<:.~ ru!~~ s
lh~ Mnp since I 1193 W~i~ln,.":>~'Ci to IJmNU~ilWm1 ~",ind~c.;.'i~ .. t 111undrhli. b!1~ w~re more 1'CfI1>'11ri>.oilil~ i~oouh '1;'<1I~ lor Ih~ hUI1!'~I1I;'d:;; ·~rkillmm;t1;pl or lht' ,ro!~~ nl1i;CI.ccl. by ;)I S.tOt.~il liiU:~ of mnre ih;m 3rj11. A~. K1u~ril1alPro~~~('d m~:ro&~ :Btt'ltOll, SclLUldm~di L~lkc BOlljIi.C lis. 8 cmcgen)' j. :sternl1
I

or

n],,~"5I~C8

or

S~ncci900. about -l:900 :k.ilm~ or ·~ldllJi1tLi iul coa."tffi.ILt'ltlj~.ilInll Im,,"c- OOcn~ !ost;!!it rates as hi~111 as 100 ktnJlycar (/4. 15), \VcaJiand 10..'>$.is muc'h lowcr ol1~hc ocnlrnl cmlSi. where the Atchillfa1aya Ri'\J~r. ,I d istribull:iuy til:ll cmijcs mu::~i~lkd(Dflh~' 11Q~""{lj'!he' M~:;~i:ss~altli iver, di~.;(:lm!"J.~s iUl~(}lI R slmUc>w in.;;;JJore buiy U,(ih ti['lN, ~~ .;;r.t!hl.'; w~llul1.d ~\clgil,; k%.. ![..;,t' .¢l1' "'1;1.\'(,; (:I'm.!.~~n :lnldi~a im~rior w~IJ<ll1d'lby ,~~bl~li.'\l'b~illCC suil aecreas ~in!]l~ilf,: 00 ~p lI!l) Wilh RSlN, un M~" ~os. .. W,I~ il]i~i<B.~ly i~u/i;'!11,d, but Ol~·\~·,~~Im~'~!i· Qlih,,1l,oo. up.
w ... e l~roi\!ionh,i)i v
~CCOI~liJ

of rre:-.hr\"I~ler 'wel~ 11:liId fm·t!.·"t> .. As KmriltUl'~ I)mh CI\"'ISt;Cid B~to:n Sound, le ..'CCt1 al o:n~ the M RiGO \\,"r;!~ bre:lchCil and t>1onn
~'k.'CUlii\~:.

snit'!:!:; of

:~nClf\:lill"flO:I:iI::Uli~. U .. }'h

AhhCM,lg~1 a d~~l'l gl\li\\'''' a:nd,~~\Cily£. ~L~.autl.llffll Olill{.!O'um:: of I.h.c ddltl. lcbe ej\cl~·, the M D'~' experienced alii (wernli Ilel. !J,roWdl [orr severel !]rIOi!lS,~I1~ years aller the S!l!JI l.c'~'C1 s~lIbH:izcd. nmnm~ m::l.lvi!ic:;. dlJrillJ!~h.C' 20~h rem.urry I'CVc.rSloo this u~ml (J 5, !7, /ifJil.

Th:,l main cause of In:-1s.w:!Sthe i~bli(}ll or ~h~ river IUjDm l!he MDP (J 1, If!). n~c r~\'tlr is 110,\1taJnroSl. cmnp\I~CI!cly leveed, prcv\'1I1~hlg OVCTOlill:rlk Iloed
il'i.
ing

:iiLlrgc :1.1J11·IU:h~t1lhl\Ol~11 hC' ItGO :.lI1d~nl~}(he ~M Gu ~r~mrncoru1m~\Va.1.I::u''!;\,,,y 10 oCln~,.flMf~e~('i rhe Ilooding !OrN~w Ork·.m~. The ~"'i!htl'r.'!:wru. orCllj~, oOlulr.ril l;lJ.1.,..... md !oml,u'i{ll~ \\'UlCf~ ~OW~I\;'d ~. ~L1!'(.'1riin ILlllcli;'Q.·I>'.~ng~~"{~~ogie' W<1IIlIr¢.'{. 1~ln1:i(ulb!Jy cm.II~~n.Y d(lwnlbilQ'liting ,md inm"<1~il1g Ihr: mtl!! l]f M!~'l.s~d~nt:e t\'\lO~O dln.1C' tim~.s ,during aeuve by oil <1111::1 M~l.;';!mxh)IClir101l a~~I'. The C!CI!i$C.flJ.(;'!ieu (If rcs~r'!,'oilN i~l a1w M is:;;is~irpi basin drrr:rnu·l.iC'~lIyreduct:dchi.! $l.lIpply (I r bolii su spcnded iiud b:::d~.o:ildediments s to the: del i::l. trj). [llpu!S of sand are panlcu linly ill1portmJ'lI.for I1Imilll.~iflilng Il:lamcr isllm~: t~Ii.lS. aH barrier :ishloos .illlhr: ,Ck:~t:lic J,"I']3in 8fC detC'riol<lting l'lt.."lC:'Ju..~ (he dewram.mioll phase (I r Ihe barrier island cydiC ilm.<;. ~l(ioo~ermcd w~li IC!}l~ ~~~vC'IO~i).~~.l(:)nl plm~ has been gfi;.'.iI[~Y

rnl~lQl) III lhc mWil. CXJ)(cl~"""tI II;."!mirll't;. m~.} 3 24). In s(l\i~ll~""il,t 01Iisam~;II.commlUnu~~ ~n· 1 1P!'O.\~~I\:'d b)'t~,"(;'"C5 W()!\~Ull~I!&tcd,. .,~]d Ihe ~iJfln dCi-il!roycdk:Vtl!..'!; pn;lI!~'i;:'!iF!~,,\,~·;t~m N ... m O~c.ms and St Bl'rmlnl<!!llti Pl:lquC'lniIK'l'! p,lf~ i~h!-:'.... ~~. Ihl.:' :>Ol!tl:lilflU (... ~.. f'iood\\'alb fil!.i~cd
·\...:t~,Cl '

of ],94 ~ un h.olU..I). it SCilC'i:'mcd, I!i Sl.om~ Stlrg.~ l1mt c%croded 110'I'll on lh~ M:i:":'" ~i§il)pi. ceast and n\'!.'<lSllred ull to 6 un s(lIllIDleasl. of New Orlean s, with 'LIr 10 2111 or 3dcli~iol1 11 ..
!iL.l::;I.'1il~~dWii~(b;

<1IiiI¥ng dmim,~c

aud

.!I1,u"'~g!J:li[m c.u:l;l~S CO!'!!lI.I(;I}L.'d,

u.n

In< bk~ ~:I,t]l]llill~Lnim!ll <3f!d .1k!1]_IlCo ilIlm~~.in1g lI.mi. ofuhc 11:.$1 of New {]iC1:Cfnnli,lk~uIJl5l': much ·nh:is. men iis bCluw S.e9 b,.cl.l!le Illoo(hvm.l~fS l'elnaiumd for J or Ino:rc W(!CID> while e!l1e~-L'ney repairs. were made· and Ihe ",,'mer was pumped out. More limn I 500 poo'P~!e[lied as. a. di reel or in/a ~n::C:1 IJC'Sult or W lurricane Ktiltn ua, a In~(iSI.

t~r

1100 nf'them ill Louisiana.

K,'I[["in.[l and Riw dC'I~Q~i!.'Cd5!£I

[1Cdm::ed.
'HIL'lrrnCaml:5:

SI..-d hncnt
{II).

O\'CI.' l:!i1:gC

to em uf areas of eoasial \1'ic(IJ,mr:1S

dI~"lriblU!;lri~" or ~h~ I'i\~r han:tix'Wn~ "'[he ~o\.\rerM1Moias"i,ppiiN IJ:rcv~nl(ld rmm ,on::ckiillg fI ShQiJl~~1i COu~· LO dtc GI.I.1J \'i[~lh(; AH:hflWr.layfJ by lite Old River ['Qll[ro~SIn:I('I~lre.

,clisdmrgc is imo l1hc.(k.~pG1..!1r or I\llC'xioo ,(f$g,.I). \\l:i111 lil(: ,1;,'XOt..llinnn of fh!.:;! i'udliJl rb1.1:Y.~ of
Riv1.'r_

and

Cl,!;:V:J8Se

fonmJ;~i()1l1, &'0 111051.

.illlillllMis~i~iS"il~:~i Del~~:<1 etlallds W I hllTie,Hw;:s .;;r:~ .,u I.ugu~j';if ~pisrxlic, lo:r'{:e ill nile

,II:!

C;lfi... tt t

M D P. Tholl~n{h ~)II'~['rJfliC1l]:stn~n!\. ,I~jbc~(;d lilt': ddru Ol!ZOi ill grew oV"c:rlihe pmilil. 6CI00 1.07{100

0'\,'('1" I S,O{IO kl1l'1or l~ill'tl" Imyc~ii:n

drcrlg,c.d

~b.r.lm\' ig.11.iml. drnilmf'C.m~d lo£ging.. lnn mDsl!'" Iy lilr oulalld £;ll::> (l'cvcl~opmenl:(17),. This and l1alllle

11Ildcr smnl~ C(n:rtlil~(rrn~,.nllf~olT.g~l~r.!OC:tI by lrmr.ncmlc pmx:~i~mocm imroouiCCS. fn:;;;;..h w~ler lind mU!i".i.ellrts. dUll. ]:'C(hICC SlIIIi~1i.ly ~md. CiiliH~mec c[m,'S'l,~1 plll"XIucth'ily t:'J~ Iluu:ricmle5 <1100 dc, posit hll'p,c :m.10l!lllt,s ohc:rusllij::nded S!dimcnlS 0011
yCi'liS"

1. A. liIier<Ht:hy o~ riOrdllg'~ or pUlis]1'19 evelill'Si1linecdn{l de'ltas. [Mo~ iii«lho rn (4r)~

~he rQrmaU~1lI

.md sl.Jis·!:ainOlbil.ily IJ'r

Major' d"aI1111Ies ;1'1 riiver

Qn~ill!l~t5·

INew ,delta Ilob~roi"maifiol'l (aJ!j'~dsi(ll1iS) nilllij(lil' :5~d~m:~liIt •.

d~p~itioJlI
1I""ulis'tl)fi1 efil!1lE1ilc:emelrit. rnajo:r f S'ed:lment liiepos.i'ti.olll, em ilhll'lCemellt of creva~se rO.M1'i1I,tiolll .a n:d gll'1C)iel\'Itn

Majo r5edimt'llt depo!s~tilliii.

enh<lWLCiedlpwdlilctiol'l 'cn'h,lIiu::ed ~edillitent dI~p@sitigll~ hi~:sliumll!'!~ ~ttl"w~,rsali!'l~ty), uuJ!tfient : i!'lr;lut, enlhal1!~lIld1P~~m{l ~Ild ~ :<~condl~rry.pr"Od1ucliDI'I 'En'h.~H'i~ed s~dim,~rUd!ii'p@Siti(ln~ enhal1,oedl org:a l'IisFIJItrilllls.p!:!:rt,
~f;!J~~!' !11l:1 !ID!!~~rf<iSiU'o!!!l!'ll5PQ!l I

Blll about ~OOkm.:2 of \\fC~ Imtdls in the BifC~on SO~IIKi Basil~~}ii!1g .in Ihe $~!l'IrlnI),nll \'\Icre OOIl\'eDlJ4:d 1.0 oren W;;)ilcr (25)" 1\lth<l'LIgh S1l}1.1lC of I!hi~ !l1('<I i.. riM'!.' Inti {If rn!'lt!i.: d~.i!:;'r, :IHo,'iII. oflhc: (hu~lln.g(-d fll'I,)al is :;;hlllHowul'il;llc;l 1111111\ in~J:r~ $po..~rs.cdw'iEb~ myriad llIlar.;h CIUiliIlj]S upromcd, by H!.c SUlmlll.. Th~ di5>turua nee of l.JuOJlrrlnlto~v~ S""d~aliEY mlll'lil~cs ,>viall I()'w-de"si~y o.~"tflie soils o~lel1 OCall"$. dtulng 111~lmc.Uiles. T.~le(';:'llC.till."lrvon. ri"er di.vc'fs[{m s~:rUCtllllC i~pre~ntly bc:iJ'I,g opermoo to, the max.ium.IIH'I C,"'IClJlpo.'\'llble 10 l;.'Whancc n:llfflh reem·crJi in. Ihc musl il.c'iivHy al1C'Cloo ~L~i1. ~nil~rul, abscn·'.mio:m; hl.dk8~!:; SUOOlIU11il11m8rnll: PX{lvcry, 111:urricane .~j!,<Ii uml<le IiIJ1;dIll:ll ncar SOlib!lnc p~~" m~hcILQU!isi;tlm"Tc..."Ias bor-dcV"on ~4 St,," tenlix.:r2005. j,:i,mcr.nin,g. ,~ l'i,l{1run S!.11;gc or ~Ial '~o .;:l III (.Fig, .3), .. ntl .rollrJ;l)diU!!!l Il;ln.;; N~~wOf~ kOlIn, mrm,; til~m 200 k~ll 'i:,!M of I~u]dr~d~, Crnl~~".J. um'lm!,llI,li ci('.!;;ill CflJlm~mn~ ~Jt~ri~h \\'"(.n: d~~croyeJ. illH.l pmt.. oJ [~IC city CD r Laki:: <t111arll.'S C",~.)!,;ric.ll~d 2-1:o-j;';iI1Il-d~r 1100d in.g 'I!>:roC! oo wi ~l. m

nr

rurgc Ilrop;~:gm.ing ~p

.1

:Ship c'hMilIel.TQ

Ihe

rt4 ~mnat >iOllm'~el1lt:;

Uronta~ pas5.!I!Jje)

~~~~ ~r~ifl.\lg~,!sliil1l!I;Ilal,~d 1l!1!1'r$~ p(FQ(iuctil)nl. I,g,w n~,t tlranSp!I):rt Qr w,at:er ililldl finlst:eriarll!s

~'>I.~hc .30-:Io-50-krrn w id.c C'~lC'Jlii.:'r Plain \1I'Ctktnld:'rc-dltt~tI SLU:gC iil'lilItd. B!..'C"Ju'>C' ~J~ .r..~~L of ~n;n :;U1];'t: mtd IO\!l cr rilopLl~~lion ..ilicfi. rC\1,,'i!'L dct1i. mh_aa 10 1l!!..."'Oplc!Dst I:hdlf livl!S diroc~ly <I:,.;: 11.m,iJ~l of Ri!a';s wiltds Jilud :surge. R~la's surge dispJaood uesidcnls trotn "II LoIiI~ iilLn>l co<1Sla~ ~::ui~~u~, .. hO",'Cvcf,1l!nd dro~<c s,.lt w..tcr.~cns. 01:' kilomclcl.'S in l~lIld. k.illi.ng rKc,.;hw~tcr w~lbllds in.url;i.~ci;;!II}'
i!~u!1~h~{
1lJ:(;,IL~

{.!5}.

I~umt ll: Ric,u's. h~glu;i'it .. n

nicm~y
'Il\

tI~MnliUl'g~ W~I!l gIL":!! ::rs lhe ~lI.rg~ cora[mn'iil'ig ihe

REVIEW
c;)i:1tt:rn sicJ)c QfN\.'iW 'il1dC<lJl~ du,lng KawiJtl!" blU had ~ CI"Q,"s;1:010 '0 km, of Chenier rl;'~n '''''el.~
!!!n:t~s
wh~1'\l'lIs 1<1II(il1;I':, :Ii~I'N,~'!""~Il>kliS Lm~~¢d as i~

p,:;·Jb~~"!l;:hirl:gll~li!li!l1pqfl'!.J!<JIQ!QI~

Qt;1f!!elll<,

dep!'h of large ll,.Y'S aJ(lng!hc !..t1tl,i~!m~ Qoo:"t AII!1.9\I{lh ~w I'J;'!ativc dT-o;;,t::;. (D!' ~h[i.~IO!\'\f OPt,;:rq. \JI,1!J~~N \'~tn:i'i imiJ~.t(li<I~wei.-

cq~~H:i.bri1Jm hutd..

a!~oroq~iro

by Ii:H,'gC 1l1Iv:]gmion channels, which 111m)! m.JW e!pb(lffilli? :gJ.~e5.m~d (li~hC'relosare

~!I1\I~l~l,n;;.£;,

~;!~

both

i;....'1r'i.'~

l'1II~'1

~~1\~1~~t[~ruligh hUlge bgpml.... degrudodw(ll.h,ult1;.1;1,~dooW~ci~l! channels. Barrier i$bnd.~,

shml~!'i.. nG wl;ll~.uiJd!>(:OIn~cluO;:~Iprm ~U!gC'and . w .....cs, bWil t~i~ 1:1.1II l:a1ll!:lJL; 01' the;;;," dR·t;t:.: i~ unt

s,t[o.l~g :I~urric;m~src'nmil'l

:s:rorval to lJC:

it is c'k:;~r Ih<'!t [he iniaet b!IITil!ri~b!iod~. wethlUlcl:~,<lnd riJgL;'~ Ih ..l (l'!rnct ~hm-a(:'~C'iri;.tl;dl l}jc CQ~IM.. J lal]clsc1~~ ~)r Ltl'ui~~i;vj"~~ nhI'd~"d !>lIb:>lfi!ilia.~proecc'lit:!.n'i lQ. Nc:w Or1e11~!O arid other coos;~a:1 OOmi1iHjl'i~I~es. l11at (,~1ua!~;tl1 depend ... be -d

rlll.~~

SWI ~I.:"$

fD1m~ Y [ot,,';S,(i!hteJ,

TItle IEvoliorillllg .MDI~' R~stDI'<IItiollil Effort PI1!lln~I.,g ~he !iI;; ,ornlio!l 01' 'the OO~J1;;l.,~r..ndscagw.; ..... n;qlli!'ll.~ lilt: ~1b-;iMll ~b'I"ciau.IbILC ~WiystOliis 1.1!111 il1t!,;'"gr.llt: hl,lIl11'lll SOck1y wifi1 Ili~ !:l1u~I!II!<"U1 OL:.b1V~i\)lin'i"j~iii~ PJ- 3 7) lI.iu:l'O\;nfuwhl:i Gi;ti'kr e]:i~~j, a:g,ains.l U~:3ihil'alproct~~"". SlJeh ~~o]o!;jie!1,1Cj,1!!iii~.!!C:r.iH~, pprnaehes relyp:ria nmri:l,y 0:11 Lhll! ClleJ:ll;i.CS. oiYr 1U.lIUIfC. 'I~'iiIin ,Immilin Ci'LC'1!)1 l'Cin~ used in the d~si~l and oontrol 0:1" :kot..:j' plUOt..~~ &cau~ of the d~ H1it!Hsions of [he de],I.'.~ p:~nbl!O~n.~. . !Ir.-di,tmmd Oinginl.'it,;~n~",~p:ro;lch!.;'~ ~Ich as I~~L;~ (,,·m!~IrlI.(:tiOlll(ind '~hc pb::ing. ·of ,d"",tycd scdinK'Ill.";; m

or

wd I
3l;,;'¢

l'iUWill,itL1] ill. pre~1i::1it by ~'it{]toII IlLU:i'iL:'r:JCl'i1 m'iOd~kA~th.olii~hil !lOiS1lo.::1~~~iO\v:1ii Ihlil[Jilfi'!I··

(rom tlI~e 1004 ~fjtlirttl [lee-,m I:llUlffimi \\'I.IS, less il'l CQlffl'IIt~Ulfli'li~'S sheltered ~y iunoo1 n1al1~roves.
{1ti}.lhc CXb1.t~lac of:m l::;:iI.~l'l.~l\'1C' b".lI'l'Icr isbnd ~,>'j1!ern olf (I r IllC'
[VI i~i.":-:;!,l~l~i OO".~I did 11m. rm'!(lCI il

fmm a ~O·m i'llJ1l.~~ dLUi~~.g ..lri!H~ K Oh..;tT'\l<B~im!.c"of W"R.Cr kw~L~ irr~d~

("'die

IlIlilil Rim'$. SlJIr~

Oill

wa.'>: .mellliF'

•ned
~J\.i!,

an a:~""''fllge'....re of 4.7 alll kikl~~lL','l1.:T of w~'lhl'l\>d ~m~~

'\I! h~~ ...

plli;o '!(,'\lll! irf(;Id,

ofMDP
fr,eshwaJl:er' IMafSh

IfOillf.Jr.)Ltil:lH

An irn~X1'I1.ml g-m~I . ~... I(hIC <lW~ic': " ...

~U~liC

~·l(lml~K+' ~"',;)~t.~ ~10il.

tien 01' lJriiJ op'[i~nLlllillinix of ,'"Co-

is sll'll~lar to jJrc~i!)iIIS 'In~nicm:ies. inchJding 1.;lliJllrlcane i\mJrow in. t992. inldicminlg,

P!("}£It1:.l1l~~

l'!lic:fI1 8Ilid l>L!aI!ld3iru 'L;'ulgincerilTlg :.lpp:roacile&. Wi~.h iJilis.in mind Iwr

~~I1CtI.li a~clr,.~Il)ruil.mtllOIl

,0;101111 SMifll.~'ancm~:Ilioll of7;Q em. p'C'!' kilomcler for ~n~t wetlands mIO~l!! !~I!~ reiIII rn ~ LouisiJI!l:l ~OOSl 1(17 2'9t Em~rrJl.;llt 'C'anr.lpics of 11.1rc.... ld K

6~d"!,
~11l!S

\'\!c~lag)ds em g['Cady dim hl ish wind pctnc~iml"lhcroh}' reduci~,g the 'wir!d~~;ii, a;'N;li!~bk: !n
,gt.'1K~I".JI1.I}'''!IL1b:r.~
,~~(Jml_"~l~C \V<J,.\i\I;o';'i i.1:,1, \1I.'C~1 :~

Bame~ Evoii!rlilo-:n iii'! AOOn.OOMd IDiIllllaJ

(30" jl). If!L;' sl1!.~lh:r-

il1g cn"(;t;1 of Ihe:>c:CllllOpi:Cd rurcm;; also ~llTl'CIS lite 1(~I~hovClt w!l:ich

wave

(fc':velopm<:meflk~" pb.oe.. Sh,\'I!lQW wm.cr d~pth5i 3U.cmn'LlC

fn1cllmlfmd breakillg. wllCro."l.~loC,get."llio:n pJlIWIIVCS \' i)a,

bonem

vklcs adclii~iorw IlIkt~OIm:1drag i:md W!M: mre.nlIlmiOlll rJ2) ml~ ~.IOO lim!.!.,,; SIfU:r~ WfL\'l!: sClur ~U).I~,irai,;!

D D D ,.

AetiiS:l1

Ib'amer

are ilelilg t":~ timtill,pirunlf:d. oQL hrll,k-ilnel:11C[J nr~I.he M DI?: (.~ )l'tJ.."lC"OIlI1CClill£l.lhc ri ver jo 'the (_lC'~mic "I:lin vii:! r~vc, reumro= dueriens, 11u:~r>C11~xmi'I1.\lOr old db'lribu.larics. and ac\,'<ls:sc-sp~J)' devclopll1Cllil Vi. 37. J8}. {)vcr nbc p;)jSI~\'I.lQ decades, it has becmnr: iucrcrlsilngly dC'<JIF hila tili:.; ~ ~'j~ hm\'~ ~ll be anile 0111 a l[il:l~e III
:.o"Codc_ Usil~g dmdgm] scdimemM to create mIdi i1.'$J1OI'C ,vcthmd .. b~

rn

Sub~er:gedl ~;;Indl sihOOI

i~ C'llCIg)' Inunl \Vttvcsclrt:hcr'

pt;n1piulg them over disilill1.C~S of tens o'.f kUmll.clcrn, This is CX~ pensivc., Il!~H because dredged ~limcl1ts, Crull be I!! sed to ercaie weilends qLlicikly. lhis[C'Chllique m.m,)! be uscjllil lilrrc.~olling \:\'Otland s lJ:ml wotJ1l:cI seen be .I\:n:H or,
qu ilckl~ crcatlJlg !large 3JlC;'l!Soi"
\"'O~lkI

by bJ'C;iking or irac:rC'as~,j, d:mg

rcdLl'C'cSdesnuctive wave ,[1;'1.101] un k~'\;"1;l"ll,DIJI~i .K.u~~I.iml,"'~';;n'1.-'" ng h1d'u~!l,\d nHI-\I~l and O\iCiU{lP'pill!!1. w;llih~(j1I\1I':.!Iy~1"Ul)' mi~~'J,; f hoIufo co ~"l:rtlJ l~II'1.~len~ ,I,q;vtie.<; ,ll'ong 1I1"lC' MR{.iO t1<~,. f.;;wwClimnd~ (IT rf~: klma&; pl\)I.I."{;I~d tll~ ~oC"V""CS frut 111

wetlands

I.hal

then be

~ligh-Ci'IC'!tV :>u.g~c 'CLJmlllISa~dl wmoc:s Ihill.bro~c en the levee race.

CQlwI,,:rliCly, Ll1:lrili.~rt"aJtlt'l.'1JIk"YI.~ I~ruby t~ml:were mrr..,'t'Ii.Gpped by db(.' :lw!!'-vclociDY surge. btu. fronlcti by C'xtJ!tlsivc: ~vcl'I~lI1d~. c9C"i1po!!tl sulJ:o;l:mli.d dilm~ age ~3JI. Dcu~nd un,g on the 1Ol1.c of :R.Sl'R. '~':O<!lS1,11

:,dvc lc\,(;c:;. CO~"qIJc:tI~y.m[linihlliil,if1,g ami, wltcre sib~e. using dc~llm.ic ,~mc,e$scsw ~ilC.rea.W
aut''''

onlod>!), <ltld nms1; ~ liCpl.ilocd by

mllO'[t;

m:\i;~I-'i!fiS-

sl~!'ilaqm(1 Ilw(l'llI,l hrh ~I' ~~mm' .. ! d~IClirm~ (J9). (3} R.c~jorillg b;Ir.~ul.'Ti~~"I1c,11ii by I}un~piglg ;:;;aliluls fro~:n,of't~liu~, con~llnJcli Ill:! gro,ill~ and break'wmcr.;. I:llildltg rip~p. end lIS,ilflg I~nces and pJ!antullgs 1:0 :>~billi7';c sand dUI1I.'S (4i~. 1/ Because Mnp bm:'ri:C-r bJrrmd:. do net jt:lsti~'iigrni.c Irlui. d(;tcnarnic O\iCrllli1t1c., :reshlmlll.Q.11 wiU require

()1~a:Jl1g fl'ilail~lci1mtce'.

R...""St!.lrn~tom~ .mil

In;fLin~tc.-

\'Vcl,l1!!n,15Illminl<!i!l! ,"' I~!r "SC'Jc]l:lw,::1 e~f!\!alklTl b}i ~.ruppin.g ~edin1.Cn'lS ;;IIW;~ fn-!Tuilig o.I'gt.1:r!C-fic'h i soii~~, Thu..... (:l~~l!l{1s rh~)IU!i ilillpc~r~n~. m kh~ W 1 ~'i~~ill:l~~i:I'iil1g Cb.';IIUOmlJQL;"-.l1:t l';~'l Icy-d. i~mill,"~"~ '~o Uk: -J 'io -I-i'i~ elGvi'il.iO:I'is 't~~il. hm1i\;'l~,ti>'JC"Ute ~

f'rr..ll.egtc Inca~inn.,> f~'ol!l!ld pr"CII\.' "" sci F-5III!sl!.1ining t'de l;omi1Icmc'ill~ ~!!l the !5.U-lJctur.~1 P1Tlt,i:.\C~ ion nf kv~",,- UI~IOrw.!l;,Jh::!Y. the pli.}l.'lic;d ..nd hydro...
io~,ic nnc~gtiDy of Ih~ wCLl;mcli.... ~ou!lic<n;l of Nc"'\l!' Or.I'(;~:tis 113:. oct:.ri ~Ix.,'til~y m)liiiprol'ili~d

or 'mars'~~cs.

~hc

rn<l'l1,gn:l'VC"s.>llld SW.JlUp:s in

m~allccc;m, L'IC'j !i]stilir;:(i, Ila~'vc\'er.becau.~ i",I@;nds I'tdl.lcc \!i...\'1~s:~I~d s!or,l'nslu:ge ,md [ll1U\,~dc imptJlrt<1I1iI h;l~lit,J1:S inl'h!.':oo;l:.~i!1 l~mo.ds~~J~' ~n. ~hc 1i1~~I1'~"" mllj(lhi~i;'-<!!lifm (If sand 11I'~'pp;:;din dlr.!
I;Ip-b;I~in
1';8iL;!"...

ni~

I~l[i.l'

b'lO'Cml¢

,I SOl!llj~

[If

oo~_r.;.;: S~d~lllCJII:' ~hal wi II ;;lid in IJ.arFjc .. ~~13Iocl .."

ni~iIt1:ll:Un~llIg

REVIEW

Fr'iig_3 .• A alUn;po.lli~e fi{lUJre~h~m[ilmgl p.ath~ or Hlurical'les lKa!irinla~nr:l RiM,. :wrlge lerelts [inl llIW~ers, a:5 1E!':5!:ilffilillt@dy th,e AIIJ(IF« b floodleci., silE'~ of L~el~ faJil!l!r~:::",!I!ild w~t!al!ld loss dU'eOO the hyni(amJe. l4). Ih'''::''Wrii'l!;,hydrolo&irC'f~ 1'1'O(;cs.s<:'S by re~~lO\ling ,spoil~.['mks" bfI.CMlml1lgC<.llI1~,L s, dosing deep naVi~,!;<Mim~ chlln~.ds (:su~h as jhe MRGOl inslaiiling: locks {oj':), tmppins soo~mcnts ,~"'Jj.

llIlod181 (6Wt,a~e.u

<lIndprol~itlg~mc:raor .$hDiiCl~fI~rt(i,fllil1s'L eresion, Suchutsmral.iong<:n~:m lIy ol1li;;cI.s. a rc'.lil~iycly s mall .;Jj1liC3, blll,1 eml be ll,;mi:cIi.Q13~I)i ,,11~ljyc i ,,1001; inO(i'IlWllnC'~iol1i w ilh d I:\.. ersiens

I'I'll1Hng_cmCI1.1.f~lC rcCQ11"1C'Clt1:m of wc~ kli'idS lind 1100d IJ:laln::o.m~d :reducing :lM.din,g..s of l~mll'knts, 1"rI,}~uag/l'icllIItHrn~ lands 11m! :L'Cslll~ iIl ~DYllOXkl in the Gul j" or Mc:'\'i,k:o(47 PI). (J~OI),1I~clinwtc oe'lailllgc ilnd ,he 3\,flilabll Ll,Y and
cOO!.
del[;;'1: ,'Cs'lQf;lItion (;~Ol"

so ]Imt rlvcr \'\,'tlter Irs used

Ulmll

cf'l:~[hte~y.

.f.\!cn \1I'i~h iI,~00 lI'lQ.i 1111!li1Iwml !1i."'Wlun~ i~ !Illll:.;;l ~ 1~11~11bt:I~d !iH~1 ~h1;[VI DP i)'l ~" '\Hlrking I1mll\C (.m, 441,.tmd: I\~l(mu~ ln~~l k~ intr.m ~l;gml~xl W:Udl IIU'II,lgiltOClJI ili"td Ilnodll~ro&cctiol:1 ililfmSlfllm~IJ\;. ~;gI:Jcul~{llIn"(;. IilIrkm dc\:el()pJl1{;n~. COJTllnlli..':I:'CJfl~ad fO:'l'Cilti.OII[li,1 fishiID.i:, aad ol I and a g.<iS produ~::~Jo.nl. In hilfni, d'teSc ilC'1 i",m.cs w ill l~m"C w ;)Idapl, 10 Ilrojlxl;~ such ;[lIS di versions,lh<1l1 roc'k ton..'tllm tillc dehu W :II, mare Ilillurnl l>lcu&(!. This h; 0jI. lessonto 'bee~eamilc,jreaflrdl~ng mml dC.~rns" Coos~n'l. l:C.'>l!ol'mlion win be' more cneclavc il' it takes. imo aecount ch;:u18,Q'l: in lres~~. a~.cr $.\IPw ply. sU~)leTldod =tcdimCllt and nmmoenl, 11W1l'i1!S in ~he Miss~issirpi Ri;\'cr B<!;Sin 145, ';6). [t silol!tki ~""o!'k ~oopt;.'ml±v!!lly enhrl~to ooh~: nmm~g.~ ~"'~Ih
.ul!;l~~oTI;:~h~ 1t;-S'OllI~ ;1I1:d ~n\' iffi!Ut'il[;'n'~~ IhL;b~w.>il~. tim:ltld'il1g 1]~~r(;'~10tdlio'l1 u:f t~~ Mi:;;~

il.:ccclcrntcd sea-level rise, c hal:l,gcs in. ~11,'ot.~iprmliOI~ I1Il1~cm.", .. d chm.ng,e:li, m in the- r~~CJl~ellcy and iInCUIS it)' hll!ij'!"JC'!Ilc:ii lH ;'l4) :Iflu<llile taken i~l~"O a.ccO~Un! n d~Jrg:n' ~ ]n!: crl1~¢I;','c !~!JJr.uli.r.m :-;u·"!1.I.'gi~. U~~ 1;.'1'1~~J-

or Cll.{'1r.gy

have

iI~ljl\Oflmn

:iJnpliil:'l.lli!llls

lor

or

~!as i100vidcd lip to $50 m illiion per ycnr iill ~le UUli~cd Simes, bUI. i~ became fipJ)illI!m IhatllfU'ger~ 9:.alc 1'CS[oITLtioll clforts weuc' nccdoo 1(;'i6t .1-\ nrore ilU::l~sfl\'c eeosysiem 1'oI.."Sto;r.l[ion plan, "CO,)Sl lU5io- toward 8. Slis~.inab~c toosrm.l Lmjlisl,flIlll," \~S dcvc~Gf)Cd in~'9g.:i!, 'which im:lm'lcd ,f! dilVCI'S\: .mna!g:nlll:8l km ofprojccts (I r '\'a:riimls sJ>tcs :md pW!rpos~, :10C"3w,1 IJU{lL!gllOlIl the iJoo~ml
>'lOlIe (;:i 7),

To 111:!1hcr ~Ihu:-!hc
U.s,

COO~~ 20:50 PI~l!lI, Ih~

intCI"'iiv~ J\::;tol,ni(mlt-chniql!lQ1\ lI~t u~c [1:1.,; cm;~e,,, 001' n:ilWJ1.:. R~lh~r tlrl1un dVl'imllliJ'lg imd CO~'l~Y' ro!;:;il .twlcl:> (5'5, shoodd 00 cmpian:;:ill!J (.·W J"

·m.

J\rmy C'mpt> nf .Eug~t:lC(.'1!]i;. IUIHJcrwo~ Ihe loui:;;imm r()m>lU~ .An;OI. (,lC'A) ECU.)I~.tt;1111 R~ ... [QI<9lioJ] SILIdy '15~Yl.Tlu ..l(.'A ; Si,udiy prudut:tid "h::lnik'tl qLl[ullti,mih'c ,armly:;~ o:r Y;n~ious rc...lara-

A N~
Fermost

IIif!s,ril~~;iQllilaJl FfaJm@Wor:Ir~ uf ilte 20l1rl ,x:n.tu:ry. IJlIbiJ.ic dc-ct",ioJ'l:-i,

roe~J em nood rro~(X.tioll, mll\I~g.1Iion. on and gas c'~,lmC't ocm, Of wi idme m,ma,gemciln.. Grrnll'ing awaren~ of the dimcl1sto![lS and oon$CCJlKllll."'Cs
ucl iUI\'~I:~m:l:lt coastal Leuisiana .. i~l
G

r wof;dOl~dI 1(1;51$

lioIl IC.atLl:~<Illd of iheoOMflJl.d c:l:lr.'Ciivcf1,c::>s or ruh:cs o r various .l'c;tlhl["CS in ;];chicvj,ng c(os},stem bi.zm:'l:its,mng [liS ill 19l<llco:Sl: ~rom$$ bl lliol1lto 17 billion, 1]1~ O.!l'ic(; OrM~lUi11tg.~1~.1~n!i.rul~d B~dl· g_(!i. dliroctrudtC' t'O.tp;,,>hl prepmrc .:ll. )JC[ii.loo-hack LC /I.. :PIan thm was 'suhmiU(;.Id 1:0 C'Oll~SS in. j,mualj' 2OtJ5I~M~ :[1, twmrncmkd <lLlihor~.~,kll1; u or !h~e "!'lC\il1'-It.'fIIlli cril'ic;Jil, "'Q~y)1alrri:SloutllKm.
~Ca[ure:'>.·'·:1 science mid ~Imo]{lg.}' p.ljogrmn" :! OC!ll(ulSlll<!lijonprogr<ul1!, kncfkiaJ use or dn;d,g!.;d "'i1L!.1.,.".ti;I]~, ,,~~d 1l11rlhtr
mi(.'a['-U·!1~ll:ti':~o:r.ll~ori

has :L'f.:~L!h.cdin t1onsidl!:'r.dde re·

or

8<OOrn

illt,d,

VPfi'l~g' N.ili~",I.""\lippu in.n" ~t;~!:n oaf II

und Ilbnnil!£ ror ~l.d:lSt;,!inl ial r1ib] ij¢ i~vCt;~1:I;nl" l'o.rlit'5~or.H.lim~ oflhl..': M DIP. ThJ;;: roo!:i1d G':Ifb'inal\,Vl,."ll;md .., f'la1lu.ing" P,·,rl!tci;tioD1. arul R(''llIO:~Uliofl Act ~('\N'.PPRA.~ ~)f 1990

Iil i~'m~ ad\~OC'LCY

im~1).~i.r,l,u~iq!lc'" of

(1~Wr:

Ic.:l.l!un.. ;It U C('I!-;tof ti l'lJlt I)' ..... Slhill1 ir::!Ui. lli~A&...;;;s~anl Scellt~m"y uf the Ani,y

1682

REVIEW
Nq~.C'~ttd I~ro~mnmml.ic ,anthm~i z-.n~on Ib1' ele~lC~~'l tol,;;J,ling $U.:!ili,~riQ:JIl. which c!.I!r!"';;mb,l

tCI;~iol'l imQ ooosm~ ecosystem reS'u~r;n~(ln in a COllla~tib.lc m"H'lN'~:.

NO!IJ;:I&!",k:!'i:~. Ih~~OOS h~iml:;mu,.'l'i, h,,'l'("1; .. ~,<;O ",wlli!~p<I:<~"gl;:nr:ru \l\l<l!~f R~fiOl,lrQC"'P.~"\'dop!!,li,;'ITI'tAct or :I.OII!fIl~ otlwt S!mJ~'~I\,'_ .g i"'~n Ilm,\ ~]~.Il\,~i~L~ ~, Ilmol.'l;'; I;;mti]ln;."ht ..illsa \.~ al1d ' aMOC""l;i\,~ ,~)~.1 CQo:;;yst~"1nl!!.':;ir~"1tiojl ~I~:i!· A.NilIi itmal.~l~scmt h Cou Uc i~ i~w 0 f i,h~ I!'CV lC API<.1!l1 coneladed: "';1 Ilhou,gh the i~~cl~vir;llllrli "mat::h.",5.~hlln dtos~~ illl::latdcd ill lire :!OO5 1iC'A p,mjcl:;ltRill [he $~lII.oy ,,~~t ,u~nlifk::d~Y8ound. ~ P~~lll p.r'lJ~t.'d I,t] CO!;!i!!~tiS. Th~ ~ndudt; ~cr. I

UII:::[ot,;' llhnllM bL,: IfiQ[<; ;IJ~dl k;.~~r Soc .. k proj~~~:. H1aJU prn;;.idc a CUillP~~It'ITI~!i ... e a;ppt.oHe1'i io i!i:d~ d rl~~,iUI.!1. lan'ld ]'~ OVCii' llilieh 1:1 bjJ'~ l:iiI!tlH. MODe ii~i,po.riaill~y. 'l.iJI~ :i.ltld,~1 sh;IJIiII~l:d ~ ~(uidcd by ~ detailed nUIp< () nheCKIXICk':cl IU.llillre !iJndsc!lpc of oo,",,>~all Luulslsna Ihru, i,~,d.e\'c·kn~cd from tlLg:t\.'Cd lIl)OU SOlI 1$ ful' the region ,mlL! I]IDC' na~ion:' l.i'Q" 6r~). (·'Ol~resli. diroc~oo the CCll~ t:o d~~:dlbr" IPI1lll fot:cln.~ro (~r l;;~Mlil'.G ttl d~l!'F ~ )
I;h:;;~H!l'WI£:,;'Ilijfm. ,u!d in D0CA.1lIlb.;!"200flllhc C011lJo

lli\'~r M,,~hm il,'ore or ihe s~d~me~l l(jj}d or [1iI~ rh er I=>reuILil,~OOil~ tile 1'tC'al'!>hore m~bC til'! 0011· tli~u~cto ON1~n.lctil\'C' aad sUi,;;nailiiin1:!: dellaic j)roccsses:. FllI'lh~m'K)re. ~lC dmml~C \\'~m~uby th'C']ntW~t,lU~i<l" lessened ~!me l~rc'\'iolls :-:.ocim,1 h,~s
(ilb~\ac:lc)1
1(;1 nl!!C':i);l l1lC'lfC

st:~II~ divrnmn.....'tliie lL'il'ig·di."l~UlC~ ~lnlfl.'i:!'1mo~~of :<..~diin'~ll :\I~rn'i~ .and ~irigtii'locring or I!~ iiiil,l,I· bgai.ioil8~ aeeess an:the i'ill'Oi:~~l of'the Mi:-l:oi:>sippi

W'lrr,~ of Ihe eoast. And Ihe jJm! has (lHC of Ihe hh;ilil~ ~.h."qt:II1,'l!K~e,'l orlmpit,,1 ~yd[lnc imlli1C'~1l il~ 1lu;t\1i'Qrlo _ Th~ mlllla~I,:'l11Alll,n"pp!ml.l;hl,.~ dr..~ '!L~I~ped ~,,;:o~ and :li1,L'ii",i~~ i:h~ M[W il~ Ih~ to or PIk"WJ1i .•dil.Y r(m::~'.\i wiU Ul1d{Jllbh:.dly innuC\!!~ ~iUl:ure.tiliip1l!Bjtio~t{~ d im~l~ cl:!>u~gc im· p~L'C"'" i;:~~~"\'I'hC'~:t;, O>lI~.i.drr1' dl~ling ,I pt:;'!ioo of i'("l'iOl!lrCi: ~m:~ity. hi .l!I:dtl ~1:iG!.l, l~ (~:\ifl'C'.rnt'lioe ill d mile M.DP indi(';JI1l:sliIun flL.":>[otatiul:I ~loch .liIIi'gc on ~!ilcs jj~iIIir,e.!l 10.11£ time ~H~riOcl.. 3I1d oon\p-k~ ...

~a(1,,~

~.~kehoMCit"

C1'ig...~S,t'liItCi'l.

agg_I\.':'.<ivc

,~~lm~IIc":>

'by

Ibroj~lgrel'Cl!ca~ioll "!"'''Jl)' th:HIl !.~K~ I;."ML\1I,.("i.~!:I.~fl!J

~a~~~ of n."£jrJ!!,IDt~R. ;nld,,(nr il~ft.:lMl!'Uehll!'i,!. ;ll1d low~ri'I'l,gPl~b~ic tilicmf1Cle IOI.'~l!ih1!.!c:~i:on.~ by drodl 00.\ i~~~i{l!1J; mlll'O\~' im~!1f.~. i\~ t~qi..~i." c\'id~'lC~ limll h~'f(l lor UL;llbn:- I]!r:; hum~1I1~", (~f 2005. pi<l!1nilllM OIU-d Pi' ,1 ~TO\~'j~l:g f!;.,'i;.\[l'gnilin<!l [I~~ c1i,;lw u\,;liiliIJr.. aion a m1~~hun:i~n~li.'iWi'mtC:C1 im~. ~~,i~1d()m~nd ~.~U,il'~ of d.~"Ci!>btm,·!l'if~li:ill!:!: :1i:Jr IkU;l reslOr<uljon r'i.'\!llI:J-UI'IL;,J :liL:tuvi~ thm. are mu:d:l g~ucr ill seale ::md ies l;lQ!.Cly S~UrulLiC r'Uum ~'im rtll:' storm ~~wtC'Clio:nl and lI:iV~WlttaiJ1 ~"tnlin LeA ~:iII:liI1Ulillg. \:~t(Hli- more profound [h{iflthos(; considered bardy 3. 1 HOI1l ~e-amres \vcre eV:l~.l!Ial(.',d on the basis I.h'::C2I:dc ago .. C'{';osysl!(.'nn bcnefhs and !l~lralldal cos!:,'>. :5)0 t~m~. UU~ Gulf of Mexico Eitc~y Sccl1riity Act. imo flI"..iu DI!X('n1hcr :W(I{\ ,gil\'~ LnuiflibCmost cost-e:~roctivc' .au:r:LJ!y (I r r~lIml!S could be- si!:l'~~oo ide-~IIHicd. !kulcllts, dldnet ~peC'iJiC;llly ~1'lCh!dc Si311<l and Qid,cr (hllf ('O:lSI sw:tCl'3?% of the r~\ ·cnl.l~ Iro!1l1new];}' (1fIil'llOOoii ~ and gas.~mCls. ~hc V~I,ll:C'of ~l'Onl!l d:!lll;agc· l~clltc~Kl!1" 11m! costs Lol!,i~:i~.amlm::; ~k,,~i]~lij0l1ul,I~1 d~>d~'C'UJwdth~ w~ro onbi nn;l!lc~M OlU]"y:;, l~.~' ID'Iv('!nrni"l'~lt~. !.wa~ ['C\e-.~me.s to ooasunl. 1\."S~orn:tioll amI prowc~ ion. ~hough !Jibe [cmmcs,lliIligall iItljOOt'!C costsor yield fiI,\OOI]lll1(lndl:-d th"n I}I~ c1101mlIJI 00 'W-Il;.ll'im1i(;"Ililly blocked and 1101 n1<!inruincc.l even Ih!l'~h<llllo~,,'~

[Fcre·~er,eiFl:tlti, ,illfid N(lte~: 1. H. N. 1'11k. E. M~Fil!!lilO, Ki:lllii\ J. Wilber!, J. 'S~ rPetfd. ~,J!6 U'}~<l~, 2. it ~iI!r, ~-ph()lugf ~itd O.ll'1lri!l'tlllJl' ,Gt,IlI~jt: I'mfo.lj' ,,;·w ImVi!r Mfuf:!li'ppi VIl'JiI.ejI lJ\.s.IimIy C«tp~ ~ o~ EiIIginl!f!~'LI'[~lJf'9, ~l6,.1!'J9~J. ll 1. H l. H. Hi. Itct.l!r~. J. (CD!fJ'l~ ~:i!.]. (,(i)5 U,9'97). ~. ]. ~. ~y ]1' .• ]. ~. ~lI1ifl, l Ci!~~l)!;h,.P. "'.]'emp!.e~, C~¢ '''It;1~, ~~. U5 '(199'1). ].IW~~l!:,]l (mJlMlilfl, .~!lfuDt. CrMMDf~PlJ Sid. 2&, 111 fl9a7}. 1']. It I'L ~~~. rmr~ Gr!Bl L'OIif./tl ~~i111, tll3 '(:l269J. :i!'. 0. (;]IlMlI Ill. ~,~ [!i;;y l~., MDr. GtcL tZ!t 1.f:i,~ID.

'c.

s,

or

S. I. A• .Mell1d~'jj~!lfl'ITl, T. tJ!fiJrll'il. ~11 .1. ri!i'I'd Calth'o .. l5Pe: rll :nd~~ l'~ f(~. I!I. P. W.eil1$leim. 0. A. Kf!~U, [d~ ~tm!lfllr,f!r A~1!mic. HOfKfJl., 200m,
1!1l'-5HO.

'!.wK~

9" It 'il 1!lI!!'l1!uM, S, lit ~lII!~k~ W{lMlr~I'S'()iJ fittlIu/, 3, 'h67 (2JO(I,3~. '1;0'. It 1'1. BiU!'T\iI!I1 1"1,. J. W. Diy JJ., C l\.. Mil!er" 5dtml! .2:24,. 1I»I!3 U9'3A~,. 11. it E.. ftJm!r:,].~. Boo~II;m,.fl:. HI. ~M..I, s, ~~'prar,

t\klfl~ ',\'iih oHlcr mllh::ipm.\.'\.1 rc~'elJllC Sh'!!;m:1S, f:L..-41Cn-; and (~i~id g.as and mwig-mon 1@!~"Si[;,,),. ~his '~TIlit'd IfJirl\ ide- <lpplilYx.imlil~d}' £1 hilhon 'per m YCl~r uver 30 ycw-~ rOlf IhJ;>.lcPlLrpll'i\."£, COMSC· Tlu.-..e ,m;ilytiC;!lI1 lil'l1:it<lILiQll S I.:'n:ocl~w.~b' i:.;ol;.ilfoo
[JCllclilS In ClllT.::ln ecosystem IJ.!SCDl(sudl as

n. s. feQL~!ull R.BQ¥dl
U. n9.~B). L ~it'!c~, 1'.

Xi!l'h{~ ~"h4A~ QO~, 12. ]..~N, C:Clemam, If. I'll..Rcb~rI$.. G. 'MI. $tmne. 14, 693 (1m~.
1)111'10111,

f. (oos11l1 Kts,

.. Si:!ler, J•. Sed: ~ffoi 5:B. '~3j!

rt.~1tm:~rilm pbn fnmlWll'iIIi:cm from ,tllht;:1' rrol~nlli:il M.!~niC'bi W:Dt~l nOM Ilrrmt:clio:ll. ~wnn dIT['mg~redlle[[oI1. alma :nfl\lDM,fuli.rm,.

qur;:<rnl)'.llil\.,': :s;!rr~c IllII}' hOlvc Ihe !("lilll!lr'C~s lo

5'jlnerW!l~ or

~[ b3~ Ik'1erullC

d(:f11t llen ,ouly m scicnll.isDS

rmsui: .toa"'I;}1 Ctlos,y.ilr;;:m f1C;S.roruion on .1 _..r.1lc .. . Ifu:gcr dmll1 ~lllyolher U.S. qiorL -n]i~ ptlSl."1O II m~jor ChfUlklJ~(! 10 scicmoc1und scienoo.boscd

.'Itnd ,enlgiaeei"S ~J~J bu~ f.L'io to 8 ~:rovr.'ing scg~ ~ncnt oFdDC pllbHc i1ndpolil.i(,'3IICfl,dersh~p lhm

p lilllning
dror;:I~\!'C

to

(k"dop

lhe mo:>]

Slratt';~ic:

!lild

snnu.cgics. whik

mi~limizilTlg~h~

con·

1(5. S' G<I,gL~RI), ti. ~r·.;!![end!.. K., Wi!lk~. l'J1lllit, (j~lf ('~tt!1. A:;!t:il.. GM/, 5~t'.:rr,M~.ll, 2'9.5It19Sll. '11.6. llit(ill!(liI .• ]1. l)unDaf; J. .(jr;Ir;Ist,liIj ~.9', :9 24 tJ.. 93:t 9 '17. 1. Il'!l. ~ ]r. ft ,,>to f'ifiR1!ies 2'3.• ,1125 '(~OOOL :lil3_ ],.!l ,~atFa5, ~. E., Bt:iu~~il. IL lit Hi:ln~ley, l£I~d~ COQ'YGI LO!.iI~.: 1956,90 ~OpIlm fil{i jjJi!pilfl 9'l·(1:l(,

J~.c"1'1.$~1'l,1 '.'.l'. ~

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be~lt.'.wi~h II1Cirn.lt'!JfiOl<UioilOr ~I, jl'liloolilOi'rlrial b'1!l~ lnnO!li nu:::k:ru'!:lC~lt.'Iii'i}e. l1lC'tr.m~m,'ed ~t1.e

m[~l~

tllcn

oo;lu~i'C :ii.'q!ll~

oollli::rrintnmclemr

c~rrussion ,:ll'td f;!;,'qucm:e:;:rn'S,"ling tlile ~~.'IC'I""OOllCl ' tOdlC m~~och{lll1ilionJ I, L;t:1~I:r.the nliRiClIOI!lcli'ial ( oOJ~'mu,ll be ~lcnooiJJ or toot r!l'l:!nih~ p."')lIL1l~:;Ill \,'I':~IJ'~'!.C madc,ll: copy il;, n.'1'li~I~U UI!l~l1 .. \.!,h:01i,lll r~i~

rli!tctiooaIOOiFiii(;s of :1051 rttiu:l!L'~lzyttclri1~, ~gt{S rue· lI.sjlmJ.l~ ~ai.IJR! i11,Ill': U~iJd0~i'i (I,.2t wilh c:;:rep1it.'!l~it:mo\',,'.d IilL\ln (Jim'ru'iII~)'~s (d.'t.Oil:\.'ti in SONI IleXt). Sim ill!nrly. o!llriJl:Ib'cn:c of 11Ii1,.,.,~1IDI uq,nxIlvIclbre nwle is ill'Il)!..'ifCd. (310M IC1Iil), mil ~lL.!S ~[Cj' lU'JohuJ:on oft&1e lhllil1~ o:f {!.<."iI1C' U.llllSrcr.
!HIid IIii3t

~nrnp~ ~li~R. I~i§h O"'lN pn..~~ ii~leliicll!. 1 . g.;..'J'l.~ t'!lY.ltil1in\liuiou s, ~"+Icrc:IS n:~.C'LirJ:n whhl.O\!I' OM},! 1.J~.JI.b apflJ1 15j.T:lC llmiumiol~. pitJI.C!'Dliu,lly m
U~~it~l1lI\"'O l'l\.'luri.~!~kl!Ud
t~!ll@k'l]Il;'llt.'l,

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im,'o~\'e..,\ieo::ld:aJ'lIrLti:tm oflnilOHudG'M ~1(; cvmbilrlal.iA.'lus,I:!cr. i~1bilil.y of 1l1C'SC ("(ubilimiLl'lk> is 1tt'Ccs..~lliY I~r 3.

t~· pJt~1>
org~'I(:lr.'IJI!';r~r l:C5ijJOI'ISC [0

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\V!1CIl a. get}!!' moves Iro~n~bC' Illi,tocIwl:drkli to ~hJ,'l !U!CilCll5,i'L';; cHvimml1C'nI dlil!!l!!!~$ ntll11"rri~ ~.lJfill)'J~o!!"~!if.>(:Onlhjni~g" I~]]k~id, l!!nir<ll~m,dl)' ia~~l;'rir~'CI m~od~o!!drol MJ,,;'~mQ ~) a I'I);.TIIJib~l~i~!g. bijp.lt~m~~Y~l.lh~"'ikid mn:'ki<"14gcn~i(lI1~. lh,ls. ~11~ ("V(l.J\I~o.ll:I!)' ;uh,·,]:n!!."lJ;,l;(;'s oJ ~uoolnbjr~I~ion ma~i d riw ml1irO!1lllC~c;'ar gC'llC~ral1 .. lt·[. OOc,ru!\.l; I'C'(omhiml1!tm "HfIWN 1l:I~lI.(i>Cimllldm~ gt;11CcS10 l;~,pt.;' d('!lJlLd'fitiml b~ MWlII!..~rsn-.nchCl, [11: im:.;lcc-;,iMc .1I.CClliln.lki.tion of dCk:tC:OOlLi IlntLlmnol:IS.,or ~~'£n::;i:SC:S; ill!! ~LI.cof sf!!t'fLd of odVllIl!fltgOOll$ I1bUlffilcms {,4\1 r d~hcr of 1'~k::SCsclOCl~,VC [CI~ d.rii'l'CS mi~OllUQkfJl" gCI1Cll'ruISler., Il1DIi'C gene IirnIlSi('cr sbeuld he 01), ~'>t..'f¥ed ~n OU~crosrull£ mxa, bcouse Ihey experience

~lOrc 'elitoctivc roco.l.1lbJm~ioJlII1<l:nl sc:llflII£' or clonal

~'lJ."'<!!'

001 t~l(! pos.'iibiiHrry Ih31 !his ~.'C~ilil.L" dU.~'lo .iII ,~hynli! M:tll gcIl011£.'S,. II is l.,m'bJb~'!lkqtro-iuhmlmlCC is. 1Ci'b"Cn~k C"Oill.ctdC'~cc. Flu1l1£nvaore~ a po.'>:il,ivc' asalso 'C8scn~,111O ~~l!looes!!tlillf'lmK'tiOfilJI gcl'C ~I<lll~'tcr !'I[ICi;UliJ()ll ~et\",'eC'n O,I>\1Nand g.c1'LC Imm;lcr is t5). A pl\'i:iti",'e afWl:i~nio.n bc~w!.'li!nQ&'inhmm!:lcc (o:n.~~tcl:1l.with thi! Ob:f.cf"!,..o:i 'I~hyIOb"al~.ic dis!'rih!.ujon 'Oi"llllIUY rcean bUl tcw .;I~~cicnl uU!'l~R~1N; :Iud tr..rns-I'cr ~ullunb;:~ woWl'ld 'Jllfl')'~'idi/lSUJitlOl'I iF sc~ l1~lgand don~:lJ'!Cp]'odtlc'liOlln.:pI'CSClll ,J , 'd::nd ~l!i!'ihypr~~.lI.:1'i!';~ emf' (7), gcnoc tr.u~b In these h i\gh O.Mc-.:! gro~()S is Alll~nr~,~~\~~.y. nril\JI!J)IckUl" ~nI!1s r\'\r t'Ul!lld 100 a 11Cl!~ Pl'l)."""CSS,:J;llllidoc1 IliIlldmn ge!~tic drin, or by il'b~!c\vo:iL~ [hcir cKlinclicm. .In ;fl(brl~W: mces;>l, p :r.'\~idl;,\dy gt'V~~, \\'ilh .Qddilivc b Evid~"J'!,(X tha1. cl~~arol'l~1S1.[g..nU~~C<ifg~'illc 1I<1!~Is.~1lN, :1'1"HlN(;I11'(;IU of rtu::::k'{ir .~"La~ :Il'nung dl.ltIL~lil· cll~L'" .on liul:.'.\li\ l1(:'glmlk.~ of rnmi:llg ~~~'l~_ Ndl!llC:f 'l( IhCJi!CUI}.poiJ~C5C:i m.Ik(';S l1Iily pu1 i:(:lit:tll RlIU;:-s. or InO~iC:rl~m of ~1le:.;b.:I.\Vt'.l"l t:!Ml> :iI!l:l, vcgtiro:1~ytLllilSl1l'llitlOO t1MooJnthimm. fire :Ilssoc!ut[IJ(I; co:m.clll:rbl~ Ill.: ","'1miOllis!up bcI\Vrenl (I"'~N mnd ihc Ililllmb!..,- of'hnl1sler 1C"',"~1t.~ wilh hi,gher 0 would rLlrlhc:r:iUlr~poJ1,!)tluhypmt~sk \;\t; t~c1 dH "~>"C rompcl~i'lg hYrjoGlI!l'lL"":l!C.~ ~, ex.. Uvc:t:all,lhcs.e nUDditl.~ rU1.dpn."'diclio:ns d~unlfinsttill.e Ilmtlliulg d'LCrclmio.u::;;hip OO'\'\iCCn, imfcpcnldclH ~'l:M lhe hCll11abHhy of gem! c.:m~bnllfll;il1~s provide> a "!,wrk tor the C'~o.llII:i.on of galOfbnC mitooholl.dril'ill gellC losses ldocillU'lClltW ill 'J}I 111,\ldoclivc :tlmn.. and <!!blce.slrn~ OMN (~I!lketl. ;IS hi,glli. intc:111L-diate. rud'lill.cdJlll't:. or [bl,,\,) aeress 1')10 m,gi:m;'!!CI'lTI. gelWl [see Slq~ IRe~e~e:n:~e;~!!!!ild N~t~.~

ncr lir3f1si1.c'I"n::quires sim:ullrU'LCOIJS ,gene~ic;cl:w~lt-~,

scl!clio.l!l. Bacauf.C' ad~i\;!C

Itll1.Ctk:n

rhCiml<ll~ebi;lliJ.Cl' i~fClleLIO¢ of m~1rnl 0MN. :ljlIlw~din.g 0:1. stm!1gI;.;'r[~'$I"nf' 0(111' bl'iP01h~"~ \I\,!", fbund .<Ii \\1!!11~~g.. pt'si~~\.~ a"roci~[~icm 'beI\\'I,."O!D OMJ';'lfUld liIli;U~Ulltlbc!' of iUl1di(mill glo!l1e Il1l!Ilsfcrcvcm~ 'tSJ':!Cru'!na'll's r - (J,,2:1l3.1\'I.'(F!iBlili.>d P = 0,.002. fig, I~. oon:,i~H~ 'i\ljlh d1e ~'dil;lit'l!1l ~m g'\,'j'!l...cQ·ird1JCri~w·K"Cff~ili!,ul." liwlc~imml ,f,I.~!1!1.: ~ mlW!£R"t 1010 Upposih..[j,1 pn...r,l:imionn.~ lL{~~I:gg~ ! n mh~ aodV'dlIU'l~S ::lre d~~ ~\(J rooorrlb3n~u.Li!1l. ~4).The l""illII!f!,C' of \ w.imioH illi L1tC' n~l1i.oot (If trIullIOllC.r '·\lcn~ ( rllJdithe vahle of G~j~ within clool;.'S {fig. S]) rules
'\\I(mlki

ror

Ahh(lWl!;lh :;.cUing O!LmJI don"lrop.r(lduc~

1,,8

~iOJ] diminish: rcco:Ill,'bi~Imli(ln, they iD1C11Cas'C '~'MN' lhc ~u'Ob~b~~i~y ~hn~~~lil"m~toC~~Lr ~C'J'tI;'
IIJi,l:i~1S
ill

IBMNI .llrJi,gh
.11'fIi~e:l1II!Iedlats' (hil: '~8) (NI:: IB,2) eNI =, '90)

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cal by

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nigh ml ~Oll~J~ combinm KlI1.<:'

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6. I. ~~ M,. w. j;~!!Irron,nrt!(ff, PDp/J, 8"'1.1 lEI!(19M}. 7. G, l.'~~bt:Ams,~ Hm, 9.1, J37't1'}!57). 8. ~JI(ifU!riI ~ ~m,M~Fpr,ild'Oi101',~J fe[fiW!ltipla, Y.s.., am MIH Iliil'!1n!jl !lI'a!!ll wM,S,8.,. i!1ilI!~!!'I '!ilfa~ RO:1GM65UA! tc ~-W. ~~I!~ frg[[lI)..MI1~!~.I'!,\.IHI~m, tl t[]<eo1~ l. It'.lJfI'I1f, t PI!"~, ~. K:lne, A i!iidlt!~dZ!it c., :lierltl}'.:01 ID<:!OO!l'I'li, 'I)i~ur liI~' ~Jl' illl1PrO'!'~tbll; rIliIlI1f1llmpt

(2L')QtJ). ~- j,. t ~~~ ffl, ljlf;leh, ,i'M!t~,r; u. 31!5 ~,(1(1!111 $, Mo]" W3il1~.(, J. GoiXlnfglll, tmlrtfun 60, 6'1.~~200!il},

:5I!1p',pDlitinglhll~~'I'1~ ( Ma~~lniial.
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C"{lJmhhmtiolls' int:~:lsin,g [lIe ellbt..-th.'CI1.cs$ 001' ~-cIcl:tt'{lJn an m [ton~!c~c,!f and nllclcl!t-nlldlc<~rcpfls. n.1!.'Il.\'i. (6'1..

Fig, S1
Riefer'etIQe:5,

6 Se'!}t'l:mber llO(t6; i!m!pled lQ.n:i!~;elfl~" 1.13418·9

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g('iJ..~ lTI.IIilii!~

"1'0, wllOfl'l (onesll'Olrloon()e ~OOuld be ,ad'.JiI~5sedl, e'l'I1a1il: yllrill*i!@'llldi~rn'.HIJ

BREVIA

IF'ar...Fi,eldl'D'pliiic,al HYP,erle!n:s, M,aglni'fyingl I S:ub-,IOiffra"ction-,Liiilmi'ted Ob,jacls,

~Qlv',"~ WI!h an (:Ijltk .. :1 mK::r-.o:'lc(lJ'l'e 1.lmmcrk<ll ;;Ipt;l~~11:t (NA I = I ,41, IhllS d~mQl1Slrot~r!gmagn~ ~k\IJ!iOl!~ IImlprQjl,:'1;.I!:r.JoH ~f 11 !t~ih-din!"I,;~ion'

the h)'pcrlcn,~. The hn~ pairroll'ld !10~ be~ol'\l~dI bt('';;l!,!I~ ofl!.hc di,~ht~~i{m Ihuil VJNA = 260 Hml (F~~. I.,B ;ilitl C~, lkC'~LlL~~lIl~hypt.11'\,1i~ Ih"Cc~l 1.liicolt:t:i:ClLI studiesen :ill op:1iC~11hyper. ~~,Pfi1."}J.'l.ojJli!C: ~]m[i::!~ftl!ItiOfil a vel)' bmad :.r.~"'" 1 or ~.le . ~~:lra.cDio~. ~iu~'ii'l ?rlliglll:.;,,~h iC'. h..:i~ C1IIILi&l.'1:! by 1.llc less l"ll '~\I:!~l,,~:;ccmW8\!~!!' Icu~~ fLltti a 11iI~t;li~!LCt:ria~, ci)':t\lal :'lipcrlCI!.~ have nrurll' or wave v~cror.>. ~l !.':::;i1n DiI~a~11~1)' aruiili'ai)' r pr,oposcd the use of an 3.i1~:>olropic m~"tI~lIin"l, obj\:Cb, wilh lii.ib-,dj frnlclirn1l-~j lillit~d Pl.."1>Ci!li.ti.Oil. in I.~IC fi1r tield lhm C'~rry h~th l>.p'.!iililfil The ;rt'OOrucd ii,nmage or the 1\::I~c~"ON" shO\\ls fi'Cql.lcncy in~:onmltion • .IIi n~iI::; the resolution of ,\lith~ hypcrboHc di~~cffiion Sll~1ri thai, ordi, ('I'r~k!lllc;ns.cs 1!~Ci erder ,~hh",\'i,lLI,IC'lcllll,lh of nauy cvanescent waecs [)])eJl1agatc :fLh:mtl Ihc the Ihe 11 icmllll'CS or the objoci (Fig. 1 n~ The ue l:'ad!;)1 dil~C~iol1i{lnh:c' 11Iiyeroclltlet.'1~nilLtcri:Ji1(7' 9'1. !-iubdi m:acIL'On re:;:ohniClu of 130' ~1'I:nwa .. ac'hie\'~d ligh~ used 10 irllas;C til!! o'bjcct ~:rnjcc~illl:l'" sub, dinl::ac.ie'~~~i~nil(.~d im,~e haw Ihe flu Ucld Micro'o5.C'op~~aU)'. ~hc ~a:l'g!c )lI}<lIia~ freQ~,1K"L1CY ,(Hg,. S II, AIIID10USh ~his work de..II~ ..\'ilh dll.': would IlC'qli.t!irerottl-\'cry orll1i!:! !i.!!\,I'II1C~OC.Il1I. ~~l"".\'l~, \\"W'C~, l~rOilXlg,i~c ~hrollgh coll.t!Wlled il\lrr."leC c}'limIlL~c<l1 h)'iPCrk'l!l~~it ~hfll~ki be ~~ltill'k:: '~o i(1~1t' t)'l W'I.·A;I~hL.: '1I11ctll.niC'~1~.Y·· d,,':ili!!lJl it ~pJlcfje<l1 h)'pi:!.h:'fl~ U~n (;"a:~1 b ~ .In(!,g.~!i.ry in A l111~;II:!l.'f!(·riat~·b;a.~~cl ~u'~¢rkH~ o~bF.i [h~~1 ri;ISmOi'Jj c",,~~i~;u .U1m; diul1t;rt,.,iollS. lJl~1k(: ~'I{ r"ll:(;W o;l~1ie';..~ n~~~ .. ,rt~ilJ,il.i't)'(I,. Ahhul!IIgh ~M}!ikc :!u~('rkn8~'~ !.in- pO', ~I.WC (h!moH!'>~l7ItC'd .';:l~LNl.i i<lct ~O:II-~ ilCd resn il1i~ We di(,"lfllm1"~ rn~C' :1 m'.gn ~f)' hl,g orl leal h>,~ Cffi'\C-Opy ·~~!iiltlI!$C$ a ['i,l W .:;;c,m d1C ol*,cl.. eur ClII">"",'<i Glii.:riooic :.lla.cl<\, (lptic .. ~ .hyp.:ricn."il'll!l!gJli~ks .. :-il!I~lIiffl'K·!i{1n· elution 11' 41 ]!lmging~B~he nCo"'!' Held, Hl<llll' I'u.:rll.:lb ,oon,~i:->~iulg ~~i(k:: ·of ~\lb~"".)w(l]IL,.\nwh~~~ut~ll'~~nw the aaion linl:~'fl.ldin1i!llil~ <Iud n):rojcel~ it iH!.Onh(l f~n fl~ldI. of Ag (3.5 m~H ,Iud AI~.l {35 fU~Udl;I)QMi~l.o,don , .. l.l1LII.·-<;yl.illdrt(·,d~.1V.ily rabri~O,IlltdOll :J q!ll<lrlz This 'L!Jq;ll.'rinl;(;"i1J~. cl"'nil!llll'llOlllOl'I, !}l~e,(!!Jl<!bil;ly fflr a~(:I'd:!:1m.11m be",n pm ......thlc. II. ~"" ~ prupOJiL;,J , ~ SiubS'tr:full,':' l.Fig., ~A). Slllli-dorrr~~L'I)n.lhtilit~'iI obor :I. hypt.:"rl!l:li'> tor slllb-d!iflfrnl;:liC!I~~lnnfl~cdimlJoUll a mnjglllify~lIog Slllpl'rl·Il~~ couM be: C{_lll~ llging •. The: '1IIYPc'rh:ns, :il1Il::lgitli ikslhl~ ohjCtlL by "'1,ruC'[~'iI by WiSiltlg q'II.lndriCldun~UalnMmkuls ~:i.rn, j~Cl\ were inscribed imo,~ 50·mn·thick ehrome ~ll~(;rloeared 31[the ill· nr.tlu:forrning the sceucred cvm~~nn. W:l\CS i[l~o 1PfiC!.j13g.1Iing w~~es hl the 3nh;;mro,pk Illl!diium. ne r surface 'I ai r side], projClC'Iin:!:!. ~ligh-Dt$oll!lliOil nlillffig,e into the :tiltr. ,~ Th'" 1ll1k~olJ:'lQlpicunCHi:A m::lI'

1;ln~i~~ inUl~ in~o 111.1; r3rli~!d,h~[~ ~OIlaml ~ p<.!ril.'iK11i, ih~ lin,,: pair ObjIC1t':~\V3S in~ag~-cl""lilt-

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rlg!tilcs
1li1(;,(~UIl'l nrrd pnlp-

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~<;ib:i~ittJ."'l'
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li.lllg~l~tilLl:llwavcveaers

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{200M,.

arc pJ.'og.Il1.'S."iii 'C;~y 001111.. pressed as the \\1IVCS.

mw~ CfLIL\'!,iU':l, n;'S\llting in a 1l1:ilg'n iflc{j hna,gc at tlte O1II.CI: botindillJ.ji or .he hypcrlen....
Or!l;~ Ill: nili~~~]II!CJi~ jb;i.'
1!~!i1:

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~if1!1;: pair ~bie~t '1iii~h:1!.5i()-ii!m$'Jladn.g (rod).•'1W1:efeas a di~ffilCli:~f'I·~mmlOO]iIIl!age '(!Ibt!i!ifi~dI in the OO!ltl~t ~xpefiment(gfeei!ll'.,A.U" ~Tbl~!'oIuy lJ!!tilS, (!1l') AliI a1rbll[i'l~' ohjed "\ON'"]nm~,ed' wIth :SlJ'bdi~liri'ldilj)~ rie-!ilDludiofil. ILiine widlh oflhe ,I)bj)ett 15 :about 40 fltlli. l1~:e-lllypenilem is m:ad\i;!, oi'18 IllI'yt!:i1: (if ~tA~~!3:'

(AI Srnerna~( ,(If hype:rlem and! or~ma9lililgl Ci( ~ub"diflra{ltijoo"l~mited obj~cil':S~ (IB) tilyperl.e.rns 'ililihi!gifl9 of~inilp,dr Ob,~.llfl wiiblii ~in.e \l.!i:r;ith ,nr 35 rnm and ~padng ,O!~ 150 n'm. From [@rt'ttI right SGlJ'Iiillng ~~octl'Onllm~(m500~e hml:ge ,[It the ljne p;JU d~j@(tral:M~cat~@dl near the l~!l;Elf sid€' ,~fl!1@ hYiPIl'.IWflI5, nlllg rdfiedl hyp?.r~f!s im'ag~ sn,ml,!ing t~<lt '~he 150·n m·5pElc:e~ !linl!! pail" ohje,u. ~aml be, dear~~ r~sd~d~ andl11e F~s.\Jilting di~fr,gl(ttiiOfi·timil€dimageh!'l1 a l)~nIJF(j'leX~fimrtnl wliWhout the l'lyperlelil'5,;. (() Ib averagedl (rQS~ section of 1l1'IPer~~F1~ hnage offille
Fiig. ]." .glllli~n;g OjJlical[. nypertens'"
~Illlill:~.rli!;:a~ samilllo'llitl:ri

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:8 714,lJ5U6 (l01J6l, 12. 11m!! lIijll'iM Ilhm: It. ~M\~i'i0\l .!iffid ~ ],'Iodi f~ JlilIil'i\;i1 d~l)~i;)~ .andl~~riflg ~~iI !~j\eti~l; Irj~eutell~~~~r, hIofliliti r'IlI!IiMlloo Ihk 1I!);,-pi!Fi'lI'II!flI. llifIi~~i1k ~. su,pplli'!!'0!1 h~ I~ .Im'rj ~F~ Q:li~MJ.Jllidil~poo'~1Y ilJllM!ml)' ~r(h !!litlillgl,le m!Jll!!~ ~wgr<f!l ~r~!1I~ !I!l. :5ij>'!!3'il'!\!i· MUrul, Air FDfC~OffiCii iii S6~1'1i: Iii~'<i'di MIjJRI (~1 00, IFA95S0oG~·1-643i4),ilmdl th~ ir4S'F NIlE.CIl.~' ilWlIro
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~'II:~M[tmil9,Qrgl<illit'c:omi.flnt;'l'LiLl!'31~:el9J·]~85l[1Cl f!l'!ale'filill!; ·aOO I'.'letljlll~

IlI,;T!1.11s lJ..,i~:~t~~ 00· b:' iurd n1iClial11lst>~Fig. I A ~ I'n our ~'xperi~n~lt. the ooji:Cl, ~nlm,goo wilh

:Fig, S1 9 IitIC'l'tmD!I' 2:OO&; .:lmpl.td8, liditlillll)! .:!OO" l(1,l12!ll5tiemc~.liln611 SUO IEldmt'o'efryH .. I~ I~S:FlNancs~!t SdefilCeand IE!ilIg1~ee~ (II!nler ~NSEC), IJnulfi!ll'5'It~ 01 (;!ur~iil, IB'l1fk~e)', (A 9117~o..17o!lo. USA.

Ihe hyperk<Jl;~ wa.~a p'.Iir or 35"~Hn~\\~idclin!c:!i ~l;;l£Ocl l:5ID 11m ~p~r! trig, ~Bl Th'G lirmgl!li.
ff'icd ~n'i~~~ (J:50.rillil ~ac;.ili:l~Jt"~IU I.lIe d~'tll[ly

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Asymmellr,ic 'T Lympho,cyt:'B D iii'v'i siii on ill 'th,s' IInlf ali 0n 10il A,dap'tiivIB' IlmmuDiB ReslP"ollsas
~lol1l"1VCII jI,mg',l", Vi bafllTl R. P,a[ilIlllliv'@l/L'~ II:chiik'il!Il K~l'IjIVQ, '1 FI!!~m)!Scliltlltiilbflc:h/ :I\'l1ldlre<1!iI IlfI~;leko~el~','l Alrrl1l!ob IBilIlliJ.l!I~el!,.'1Sarah, ,A, t~ongillilortll... 1. l<f~still1le iE" Vifll.ljl •. M. ".1 p.;J l! I..Mr ~~s}Jo\lIl1!@ (lU,Ult1!'~ Nig@i, Kilne@'m~ Jo:r,~l;!lrl S. Q!ra,m,~~!j;So'illFo1l1 M,. Ibus@U?,!:i ~ 'Wolfgang Wlenml1!!Jielr/5~1e'~,~n I!.. Reillf!el(flt _ A hall,malrk I)I~ ~aI!T!maJlialf'll i,llnmUl'ilily 'i:;ile h!llil.er~gef'lleilY (If c.eur.a~:~h.at e:ltists amli)f'lg p.atil1o'gel'!" I~)lperiielw~d lylll'llpno~yte~. We ~hO'\lI '!iI1.at oil d~'!,tndJilr'lg ij·Y!il'ilpal(l'CYtein~tii~l~y res:pol!"ldif'llgl ~o .a mkrrobe T I~)lhnb11t UlIIequ at pa~titiD min ng 'of plr0~elns :l.ilIat nlle~i .. u~ ~~gllila,lnng. ~e~l f'at!espIH:;ifi~1'Ition~. ,alnd s <Ii~ynlm,e4:ri( eell d~vn~i(ln" AsYllfllm'~trk8e~re~! a t~{ln iJf d,ele~miin<ll'lts <llpllll!!an 1!0 be nlorrd~nal~ed bV ~HI)'~O Ilgled ~n:t.erac:h(llil bel\~e~fllih~ T cell ,31!"1d ~it$ anti'ge'f'l·p~eSE','!1Urn9 clel~ be'f'(lJ~ d~\tisi(llil. MdillOllhil ~ily.tl!ile ~ij'st 1:\00dalt:IJg;nt:eT cElis dri$p'~aYBd p henOl.ypk .aIld 'f'liJlildllJ 1m~llind1ic.a liors, of bf!rin!l!l rr differl!!lilltiaillyrate,d tllw,i31rd eU'el:t orand memory l]liIeag'e1ic. Tliie5tE! Ire:Slill.ts ~liIgl~Je''>1:a. fI'Iechalili~m by which dl si rnglLIl!lymphmyle (<I n apportion di1,l\e'r~e (:,e.U fal~es l'Ie~essaf)l 1m adaptivci1l ]1!111 mUl'liily"
1

f;(lIrled \ly "0\\' q'tr.l'lTIeu)' .;rlld cXllmim:d hy mkm~py.'JiIli:;; ;rppr~~~ 'I,'!] :'l!)l\;i(;!, nlmt 'I.\I~' \,V~ ~:X1l1l1~ll T«[~ 1'D~'i;:pari~1gfor i~!!! their Hi.~r d~vi:;ii[1II1,
Qpl1r~~!

Undivided (pal'euwll T eells were ~1<:!i!1cd .mlik!odi,~ a!}'lin~l p·'luhuli," and 'V<lfiol!l!s T c'L;1I stgli>J li~~ rt11~ci~~ 31id r'C'~Lcl ily id~lilliU'i(jd .:!I:' SJdiV!ll.cdllJla.:.L~ jjy I.~~il' inel'C'~I~d: C~Ui;i;:tC 'll]g. S ~I. Most OJ nlt~bIM\:; ,ror!~ai:lh:~c1!Ii ~:illj~I~ iVITO('. ii~diicmimi!1, Ihm ~hcy !'1!i>L:iX: 3Cli!'!8~'tI, btU. ~~~lilo,t~e;. ,~'CIILr; f~£.. I 1\.• ~ila' len). In morenlmi1r ( 90'"/n or iliUC&Q «Iktile eenuosemc colocaliizrJd \\'~I.h a, 1'01iI.!,i7xdp';ill.liC'!:b cOlnl.a.i ni n~ leukocyte ~\mcI10!u"a"l...;ocj'll~C1 anligcn :~j lfA-I).. CDK ,HId "D3 ~'Fig,~A." roll'kcn~. lhi.~ rX'i:~arily W.'I$ I1'tlI ,I dfll\:"(1 rtilll h 01' ~h~ ;ld(ll~l~iv'I.I-'rr.Ul~R-r pro<..~,
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l'x.'C'rI ;."cr~edw an~bl.'C'n c",!1!ibj~~'" di nlL"'C' ~<li!1r e iug ohhe ~~ne rocf,;~p'lP'Iil' S2l~~nt"(ll1tm"~ tn tn~ ~hc l'i!Oh~ri~y ,!.';,.;h~i~~"d e{J;m,j]Qllrcr:tt~kll.o\'l,iH 'to by P1u1! tl; i;p{t1~(l il'l~ll'!rnunol.o~c;)!J ;;.ymlpM.:· FiDll'H,;nifw. mile loc,~li:lmimlOf 'l"D90, a 110J1!\YI18plie 1J'.II]1Si~m::nnl:nll.nl~ rooeplor. W~ fOI,ln:id[o be dlmlis.~illl

tiSSlloi:SU£8(;"SIS n:hal ~.m ~c:ti\!.u.ed T ce'lll unclE:~, susmimxl inlerntCli~n \\1Ih :lflti,2C'n-oo;:uin.E dcadr.itic rellhl. fDC's ~ dll:dn,g 111£'~1]lCLVIJII I)rct)crrling its finn cell di,visio:n (5 In. If' !i;y~mps~~,ikc poObriuy eceers dl! ri~~gIJI~ p.(m~ 01." ~iI'C]~ il!!~ !he r~'gil.'.v~~r.lli.w cap~le~ I.YIn llm~nt;)Liv~ \\/ilh PC~, timl 11In:.;I."!I)d'I::~ h~!in::lT ~ ~h~ ~k~I(;'I[~~irl~',;n,g~~ ~n).;~nrnycc~~~l. nunl~~ml~ ~nVlgil,;"d ","Orn,,~~ ·
~l~[pnx~~

~lC C'b~l[ll selection 'tilco1)' of ~iliipth',C' iI_lllnLln illy s:u~er.;~ s d1.i1l~ roU [cm!~on n.t' .~ p single lymnrllOCY~C: ,~lDu~d ~}li[jvtdc ::mrIk~enln~ nClioll fur ~lC'~H~ elense ~clTC'Cwrcells), d

d","vc!'flIprnl'!:I!.~:rldhtmlLl(x-;ru;;i~,

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~1.'fH.xI J1Il!X:imnism is IDled to cmlfel" di;qm_r.J!tC' th'OC'li i1n1!_11~J,l cbul,llm;:r cells, This; !11(".dlltll1lli:-;nl!,
'5}'.u!1l1l~lriC (id~ di\ i~icl'Jl.mrl\'oh·~:;;~ bri411 fll;.!!i1U11IC1~l of dt.'1!I;_:nnilllllm o( oclll1u[", pcrpc~~icl!.lliJr~o [he mit(lt[.c spindk.. Ihu~ Clk,i)lrin'llt Ihe ltihl.ltNi~ illh.crrUll1J)CC ol' crill.ClI,1 rl.'IOlo::ulcs fJJbCi d:i~1lIre of da!llgh1<:1f' liI.iS l!J. j\.ozy·ml~'~iC' ,(XU cell di vfuMo.n could dlllS represent a IX)leilli~! nm:X::h~mlsm 10 en.'it'll\! d1ill al);Propr.iatc diversi.ty of eel ~rme arises from th~ cJk!lll!il~ dl~oeud\<UlIS or,[I sin:);lle lynP'JJOC)'lC' d'!.U'h~galll i~~llnm~ef.:51'101!.'lC. f T '0011 i!~II.'f<IC'!ionw~tll a!'ai,g;(.l!.1·prorendn,g iCcll...
lI::l1.o\VlI1a..

i~ (;haroc~~rizl;id by orif.'Jlu'~lioJl of

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~C!.lb.1.!lil1l cylookdijoJlNUI t'L'J.)'IHll11C'IL1;1; ~gI'Cgalio:n 01:'5lWllililin.jil and .u_I~r.:.ilt\"C proleins to\I\':lJl,lI ~I~e OrhlIC!l;cll:ul,w t1pm<LC~ (,n.Wi~h~ll~l);;!I.h~~-dI.s~WI,d.a ing to thi.; T ';:'4;U. i!I!lr.,i~lh!!:~"J'\."(.l!riU~I!ijQ~1.1;:<111 ~'il for hOlm, il~ yulm '~3,. A.hll.oUlg;h 111,1;: imm!lII~tll!Jg~

sale

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rWlJ,ty 1'I..';;!)~yOOin rlYQc of IJ'I1:I~}hnltl

,2002• ..ntI!lli'iIiia ~ep<!!1JTleru d' Mi@-witlOW .!!Il'!d liHrmt!liI01· Il>g]f, 1J1lI~1~ III t(aj)ijo~ 5.ilJll t1r:ilmd~~!J.i (.1\9'1143, IJ~ ~l:Iepal1tntm~ ef Pediil!ri~ IUI~Mr.li1y II!l Poemn~iiI, Pltlla, oolph.1a~ PA 'll9'lI;W1 U51\,~GeQ1Ief fog"M!croPhI!ll:OOi:s, Fcar;:J!!~~!(if E'ingfneering <llfIdi ~dt$l<t Sde-, Sw.i.r~ ~(y of TIi!dnD1.ot.J~,ViClm:a1 :3i1l~ Almril~" ·lhese ~1I!Ulm tll:l1lliiMed: eq/llaUII' til Ulis wM:. tTo whom cOlTes),\IOJide!IJce' sholilld be addrts.~ed IE',rnait
~~~m~r@m1~1~,rn~dLIIp~m~u

lAb~OIl ~l11ity C!!~er Re;s~1it.1il1 11~~W",e... d illep;Irtrnent O'~ ~dKfne, W~nlli~ Q~ Pen~;v~ Philildel~ PII 1,910~ USA. iIIm1!l!.mology Pr:QgJ'M1. The Wis,t1~ 11~~hr,e. PJlilade~lp;llia, f'Jj, 191{1!l, U~ ~limlDIlle ~MJ~·~1Ii1!l91atJ.t:;rdtl:Ky, l P>Mer .Pk:ClIluID eanGl'ff (mire, l[asl Millbootrme. Vidoriol

pemlieuW'1O ~he I11IlWtLcSpJH1dle, 3 m.orp~mlom' n::mn~lisoo~1 or usymmnelrie 0::111 division. "w\rc next examined if flsYl]1Imellk T cc~J d~d'l<l~asy~urnclroc cc~1 dh']siml can inhia1.C dilllCr~ "~ SiO.ll ~QuM 3~~O be dNccted in CD4"' helper '" cell it:spO!~~ses,l.dsti.rlw.I,ri'thsll'Ccilk T eel:1 rc. cnt~!lion as!'1{ldill(,~d \'\,'~t~ ~~e ;td<lNh<c inmunnc io ~rH(lr (TC~}u.'an$g.el1tc CD4 T ~clls were [il':'N'l!I_}n.'<f,.~ oj" T cc!1 ~s. labeled wilh ("FSE ~md ;lclq'p~i\'e:IY~1,msliC't'iI'CC1 S'~glr@i!1l~t~~!'1 ,~f hinl'lf!liIfl!~ w@ir.:epton: ililild ~Inoostl~all pot!:! ri~yIP~~t~'i'J!:siP1nwitOilk T (~US', intu 'wil{J·tYIW :lfCcipiclll."ilhal had tl!Ct!n In Wclcd \lVhcl~~cr i.~!hijJl~ cylQkiDlC5i~ nai\'c T OCUli, 24 h'{l~!,~ rctlli il;''' wiuhl.. IJW_WI" ~ U~.AI. 32 ~'KllIJS ;!!If""~ruhc n'~!lI!.rCf. I!JImJivi,kid, bhl"ll~ Wl;."IX;' C'X<.!_m· GII;O~l'l' d~lr.in~ I;:OUl.111;!. wil.~ D'CI'I, {11' ,I:al;'f ill~i~~ (br~hl; kl(.<lIlliil'JI.i~}H LF/\.· ~, ("D4. ulld Ul~ {lr di~Mn::j,miun IMIl!~ DC:.. h,l!.>; m bOC(;111 :L');:~ol n ,,~d (5, ('1;1, 12•. JJ). Wel.hcrl:foJc tc~ti:d the IJQ'lcmi.aJ .n::.t:l.iplur.uor il1l.'Cif~rnll.'f (.IFNyR" wh~('h ~ID:Jutill.!L;" for !l:;,yrml'i('! dh,'i~ion b.y i:;ol!l!lil\.g, T relll~ ~cl~I tOI:he ~.l1rIlYiltmO:~oMj{'j}:I,lS)'IWpSC d~I~'ilJ\kl~cl:i'l'.ual'On i\1I~hfJOO NQo.~U:m; ilPPc.:ul\.'t! ~Q 'be prq:mrin£ Iorl hdr Ii rst d h·~· oR"D4+ T <lClis ial v.UllJ'Ollj~ pflCmil.otlc bliJ.'lIS s:i!.-m ~il "1 \10 ,~ <i cXf~mil'lifl£ their ~~qboc~J'lillar rolocalizcd whhlhc .Mlor ... p<l111tiolflcclira m~ch;:mtCI.crl~~io..;: C};; vh10 {til}" Nal\'c mOllise CDs+ ruiI,d remllill,OO ~ymn~dicall)!' T ,cll~~~rnllsgC:lilk rorlhc P 14 T ceU l\.'Cql~or lotte' ("D4,+ T j;CUS in \1·j,y'o IFig. IC I. As.y~IU1i'iCiFie l!t'gcgaticm or signL'iling nmk:'Cullel': may. tlttj,~,b~ \\ eTC L'f,hclc~ \l;,'i!i'I ~rooxyl1il{)fC;~lH ~irnl;:c!me )lJIJa;~nlimidyic.':m:r ~.. "FSE~, and Jdo.pti.dy IlThllS· ill leaiult!.!or'both l11lajor T «.I~~ b:i;~s iLIUtlergOi~lg Ihe Iii lSI d il,'isionl d!JIriilg OlJilimmLlnc'~~)ol1se. Icl1iliC'\l inlo wi Id-~YIli!n::cip]el1.ts.. Th.escUl!cip.i~nts Olll rhe basis (II rlhdr s.uSgcslcd~ok: cl\JI.'i~lg had l'lCCill in reeloo 24 hO~JI'spre\l:i{lLls~.y with ill" T cdl rnigration ,mdndiv:Jllion in v~u;o 06), tm,,'Cl]oI!S 11I.;'J;:{Ullbill<lnl .U,"l.tda flJtmO(ltj)~"ll"'~ ~'II~nA;"" t c1:;..ami(:le<.i llll;: IOCtlliO!<H ion Cllr ;H~cc-slrn], I:mclcri~ '~"lf!e5'.1ing :~ sa}C'CiJlic- 'I!l~ig~gii; gpJ,3-41 ~.g~!lI'~!Q.~of ;1S!)'ll1.Il1~I:£1i.rCL;(;n di, ~s~,on,pro~l;1in pt.."\P!lidc ~."pilP'l~ (~~,.33 mm!(.!It)'(tlg~}~~'t'1i') N~. L I kiJ:l;lIs.c ("-I; (.pK:r~)and St;:rillble. in CIJS+ T AI. 32 hO~ifl'i [~~hcr 'tn~ll'I;~ fi;r. Wt~'idi\. idl.,"d donor 'f Cells. l[trlil'Sm~(-d hy lill(" brugl1k~L i(]~SE f!CllkJ rell~ jlil.-p...'iI".iTi!; fur di\r.i~O:I't !pIKa; is'1~n'iiiI'WliT:tI-

~I.!Ud~vi~iO'.u,~t li!>pu~hltl Ul;!! ~b~Yl1nll!...'rny mighL C!1Sl!C- S!:i.~bh: oril'l]i::."Ltion ~t~co:rigiJ~J'I ~nicro· ~ubtll'c o~D.l1i:linJilccnfC'[ (MTOq in ,~ ~,")'!lIilBl.'iC· rrol(iI11(1~pt'li'l;~itm ~hrollgh ~hc lirn~ or nrnito1'!!li WG'lIldu.:;Sll h. ill! lilt';"d~lplit<ilk~ cclltrm;[lllili, nmving io lite cl:it;ml ptJl!.:d:LIHin~!!I ptopJI1G"c. If ~':gI't.;"gmion of si.!ilnfLl ing OOJllpOJgl:;, \V8S also nllllinDalnOO dimrir~!.'ldlC ~m(!M~ of SliSuhlU!1lool1;13CI, then their oricnterion til a pole orlbe mriwli,e ~pindlc wo.uldlillfill lhclrmUnmrks fL".~nunetric []i\ islnn, ComSCl.1llLCnl~y" 'we 'Ci'i.pro,ltlcl tUnc roo~ibilily

or

T c,c.lmsrro'.~l in reeled reeipieets C?ig. I j\J. AWaol.ii,ghluch tess ['I'c~jjCnl Ihilltl the preluilm tcb!laslS~ n~n~£l! eel L~ Uho.~' ~I'I propIU.ISi!' lc ~lmMsh ~mplliL~clceuld he Kl'cnli~iod bylhc 11It\Sf. l'IIO!! of lwtlop~ile~}' rKing M~IUC::1, IWig. IAl', Ukc~hci r tlfCf!liHl~ic 'C:Ol!ml;"Il'!an~, th~'f,C:~niu:nie T cells eXIIlibited IlronOlII1Ccci pollmized dfisnr.f:· llilltion of lfA·]. C[)~.~mdl m fFig. ll\). As e witlia p:I'IC!Milotit bhl$t'", tile :-oynr!.l]lic 'OOHlpm!~"IS
coIC'lC<~h~d\\'i.~h one ,\ill'OCwhich con.scq~L'C!Il~~ I)' [1I..';pn..,:,,(~til~cd ml~ ofuhc two poles of nil; 11'1~. rrllic spindiC' rWig_ I A ~ The in]ili,,~ a::U di\I;:si;Ol1!ii 'I'Ia: olJ:iCr\ cd in ~hu$ e",.pl:l'inncmtl~ $~Hulig of (_1)~+ T eell l\.":.lIDl'I.'K'SWIi:IC, I.l~errclore, d1:Uffie-lC'ri:2l~~ PflirtiDioning of .siglflllilinlgprold~~s per~ hy

or

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nt

or

11687

RESEARCH ARTICLE
limn hmll(l\IOJ;!l <!!M'lica ~ I~KC.a COLHpommt an "",~"'I:;f1li,.l, ~C!~ 'PO~!I!ity ~Qmr~C"',wn""linig~,g, !1:~r_) a!l,o, r.!r~ O. S!;:ribb~'g ."" ;;ii, :sign!litlN C'Q31~ppll~~~ of 0l110i!r'I,,'l'k~ypol [Uril), 9tnVlpl,~, il1ut is 0 llJ.,:ll rt]sitiDJ1IJ~ 'oppf,l" i:t(l"~l1c I'lJlyr'H=!11 PKC"'i:t'lI1~]ni~!g oomp,&3;", 17~ ln l~remitO'!ic T i c'~ll bh!!,,!:..rKC~,~\;,<!~ ob~~n'>\;"dtD Iloc,llli~ opp(il>itC lh~ MTOC ~lI1d pc[5~I~di!1~!l Ilfiit(Ni~ ,~~~,

or

or

was

17",

r..JI'l[lg~n ...'l;,_"''Ilgci1!!14>d ~~Ihl:!;,: t D(, Qiv.~:lIio~.

related

~I~~

sustained

"ln~lwd \'\\/9 .. ~h"mgu~i\e z;;~imt~l~ I;~tcilijll~ I;:~n v i~i s,im! i~! 0., III ~hlll in,t, !t".l'l~g.t~lt~~ vj\, f T ~1I!l'!Nl;:n;:
~rm~~,fi:[!"\;~ lymp]rltJIK.~lic lLMj 1,..dd:jc;el1~ into, 11.~-

Cl,1lJ:1mct \;vith ,,'Vc ex-

00 Iml :'l1lll1"l:xJl1, 1~ s~ainoo ~

lif)oIl~C~

QQl1!Wlct d~11~:hil!.'f:nC't~~i;t,1l; p.l'p;:;lucli\'e~mml!ln~

iml;,'l'\~ls.,.. T cdl re-

c:irk'cll~ ~ il'lili<J!tc cd~ d~v:isi[]n in theaascaec of r<l~hog~!1. ll~ij~,riJl'nlI of dh ision is ,~ Ihid ,1I,l;:~1~" ..
oti iipi5C'ir.c ~oo~tii'[ioll.

(2.1). !n IXI'II! t~r.;: ~,ni 11 K\d(~IY~lIlphqp¢ulij~ ~nil;:~ ~mdthe iJ1~i.'CiJL-cl iCAM ~-dd~ciC'lli l7l~ipir;~I~. ,,'i£om~s cell di'>'i!lign gfd1~U.rrIl~r:en:\t·d T cell poiP~~I'1I!~kll~ \1i'0l." !;i!Jpptll'i!~-d (fig- 3 C, Rg. S4).
Uicl\:
\I.o!ll':.

'iJ.i~ u:'J'mri:~drit: ,PQ(jJitit'Jirn p{)1c (f~.!}.... IA ~i1d 2).

('0

1~, [lie

'pWi~,1lt~,\'e

ed I~. 'bUt iilt>l.cftd is. kilit~.tt"tl ilj,i't'$flC'il~e to' s~J:r~ mtli~cn ~n ~U~cmli roi1oiVlieil~,,dc\ u.i:d ef olh\:; 1j'1i1l1phoeytes OJ!' I, In Ihe seeond linL'XI~1Ii. II rO~ei.lJ,9l micl'oh.flaJ: ,~Iimu~us \Ul:.~ used, ~ut in rcc:iw.icnt)1 Ilili:::kunS. il1,tcI'tcn~clar adim.k'IJl m.oicckllie 1. {Ie I\!M II ancestral fI('I~'lliIJi 11!~~,WO!L'ka'~~IH-:arcd~o be C$!:ablishocl before miw,sisOlml ~~~i'it\1d umo M and !iO '!limbic 1.0 ~UPllO'J'1 lIFA-1 ,do.l~jI!(l'cnt lme!" "~lijOII~ bc~w~lO'nT cells a!1dOC~ (J9~. Thi)': !'I1l!la' !llm~ wilh dmrHClcri.-:!ie o:ri~~I<ui(,'iJl at O(lll(l.silC .~(l!k~ ofille nJ,h,{1Ii~ ~pindl~. ~ion \"''<f1;.d~~~·n b('lCOilISC soch i!lK1lfit~dh; ..~in : SIl$t'a~It@dl$y~.a,IlIS~ I~eq redlf:~,r al[>ym'iriu~Ul ~I!IIIJ.:;I".1IJ;1Liol'I'" hU\\:"~('ll shm\ln 111Ib<;~ cr'i!i(,,~J Ihr IIi '~ufillf!g 'HlII1itasis, To d~cfl!ni~l~~whctJIlLNcnh~:l;~yn~~ (lrrAmlj/ln,~""\YlmIK'lrj.c 0011 dh'is-ioH in I1Illlll!UliII," ~.l!.'~a.y ..\<\~ a:h!1~r\\,1CI iUI the intli..11 m rto~icT 101.;'11 li.ul $kin (~"t(J) and ~lt;'Cau..;e[lC:-l l,lCkiJ!g ]C'AM!
COI11IGiSt"

,Ill

dl,>I8J1 Sctib1b~je

lmnk'O~~llic rmlilti"OltiD'!l ~md d~~ Ii~ (;k:~lid IOrt~!!ll'ilin1iigel~by r :

or

how(."oct.

U ;!i,1iJI~t1u'~li~L]~O~~

of

a\~~Iii.ltt{'ilry .iul

!O!.;"&lized with the Ml"OC lit Lh~ pLJI~ltive synliptic pole of 1\t'CI1Tli,lotic lffi."n:>~!I'i~. mKi b S3). ~his. lO!.J, 1K'1'!>i:.lcdinlillrn1ilClsi:.- (f"il:!- :2 I, TIHI;.. ,au

11li.ltiLi·uie cells l'!th"lI!;""-'t1 !'rol'i'i Ihe T ~mIUlJll! lfl,iCl;y,d,lhd iiiILII~~ di)i,hiliHM.i:On of :SjliliiiJ-' lie aed polaffiity piiL'il,di~ .lind I'IIQ,,[j~p'tl\.·t'l M.\:itlalitm oOrllt~~ I'rot~il~s to h;,"lnil.Gti.c spindle :(Fi£. 1. •. 1-\ and B),. These Ilill(lin~ $u."'i1~llat cvcm'):
'~he 1)Iill!O:I1~,Cdcmllacl, bCIWOOJl T ~d~ ;[md

durin:

OC~ pJ'ocooiu~ Ihe first T rei! dh.'~sion.~Ol~hcr !h,m g~lcra! mi'ag<:!nic gugn<'Llirlg, :!1lC' 1:cqW!,ir«i'~O 1.."l!<.1Mi~ha".j' Il1I'l1!CU}'.
A$YI1l~11IU!U11c;iiillhefi1:~mce'~ff.\;I~~ det:en'l1i, 11!\l~1S in,~li!e firstt da!litg1'!iI.E!ir mil!;, To delC'l'Il1Iim:

if asymmc~ric ,\1egfl.'!g.,,~ of i{ua

'Ilyni!p~ i'c~}mtcin$

,A
,....

ir:t ,...iI

• WiICf·iyP8 (il1l~8eted) _ Rag 14 ..,r(uninf'8icted) bI taaAl'l'4;. (inrtec~oo)1

lF~g!, l\. A$ymm~~rk p;ro~rin Stgi'\eg~lt'Qlili!1! !iii!jt:Qi~ic'l' ce'll5 ~$p!l,i;r!di[lg ~1IiI i"dcUiol1. (rPJ IUlildl~aedP:I!.4 CDr TClR tr.c!-riSigenic T (ii?ltshClm :!!p:ieen!l (If il1:dkated ~d,l:Iieffiits ~~~rehal'Ve~ted3,2 h®~HS~ft~dI:11 \Iii;1lI0'~~pal~~r~ ~he ttl Tubwll'in Merge TuilululII lInIiOOg@flrtc ,~tim!!!lll (fifd,~:~fiOJ1, ~mthr gp,3l-L mOl1acj:\iiOgmes 'rQr MkI-lYP~ .lin:d ,co[(I1-1- ffil~(:e~~YJn~h.opelll!iaforr fl,(j!Jj-l- mb)awtd e~liI1lined Ily mrdri(all mtCuo5(oP¥ .lirtpr ~l:ainingrCDr Jj"ttlbuLfIl ~red) and IUWA-l" ([B" (D8, rK(~. i),r (1)190 {g~een,).l"al1lels aro rtlPre~ill,nta.~v~ of Inine er mJl)I'e mmroti( (e!I_$ lIl!~r g:r!!)i!Jp.~'alr left panets alFe pji~l'Ir"~~liclasu ,rfflif'll if'!f~(Md 'Iv:1'ld·lY1P~,rtdp.i;elil'S narg;~ b ,ulll~. DfUil:sangLe MTClC); all~ rE!m~]rli!19 panel~ are mitotic tells MUill~ oPIJ'O)ilt.l?ly Wadn{j ,MiliOCs). ("B:~Qu~ntllrk,crl:ij(ln or po!arized r,~~plQr5 'or! IfIhot-uc OD,E!""rr !JeRs : rllpresEl'rrltffiin (A~.fhe Iilll rnber af n.n~tDotkmlls examil'le~ fur 'each reCi~pl:or [pe r ~Imup (If ~\likl-tYJle1',. R:agl..j..; and /f,(Jtnl""-mioo, r'espe(tilo'e~~)\va~ 8;5,oRm.,.;: f U'A~l {Zl. 30, 9');CJI)3 (1.9. 27. U)~ (0,3 (15. 2:7'. 9). ~I" s '0,02:5<;~"I';'$ (11.001. {C) ~n;di\l'id'lldl~on~~ l"k.hm~m~-~II!~mc ClI)~ TCRlr~J;iI:rg~n~c T (1;!I,t_:s, r'~©m infeded Wi'ld·lype recijpierl'l5we;re hil,l"IIe~ted ~rQmdrtl~ Iling IlYllllp~ l1ode:si1d'ler .32 hDurs am:!1stain,ed 'wil~tlnlibodies ilg!lIil'lst I3-tL:lhu~il'l(red~ and LFA-Jl., (1)4,. or IfNI~IR {s~eefll). Pli~[lrity (If IU~,~.-l, (~, a midiIIFNylfi. m~ oDs.er""ed in (.iO~Jm(R. ~ ]5)1 ],el.% 'rn '" 9),ainr:l71%, (n '" Ill) ,ar mi)llIIlk iJ~~ts:,re~~&!iiI;Ie~y ..

16813

RESEARCH AR1ICLE
10 ~'1c.'\:\F1>li n \"~li,,'iher ~lgJIml~f1;g P!i!l!I~\Vil.y:'i, ~Clyt~ gf1'lII.'Q ~:m~dienrnilli.l'lg .si.gtm~ung p<llh,\'OlYS t~'l'ro~~gh miro"i;l:. we ~x;Jl!1Ilil!1li,,>dells c inwl~~ctN in 'o::~~.t~~~might be ;;l~~JI'fII!!lflCU"ic<!~l)' !h~l<: 11ro\ ides 1l 1~~!tM~~n~h:.mi ..m thr ,.sym~1!1I~~.going 0"t9ki~.~~.~:ii,$QrI~d.~!ld~vidod T ~~l1,llril;: 0011 divi~!on !ogI£Pl~f!j!W- im[<JIdm~~1 T d~· ~cil~ U1,lt 't1ld"h',~dy ·!\,,:'irmlde~.]4;1 \',<uri,nl;I" i.~ 1),,1.ri!~·iiorJ.c~in CDiF!+ T c~~!",\'1.~1,.~"~II,i!J)i';lIIFNJll. \1 I\'0 :;;1 il~1!,l1~ w~!'\' C!,IIi!.1I,11rt,Idin IliJ(.,d'~'i QO]l[[lIilf]ing i in i;lL;:n~'~r~oltdin~.o k'flH-:l.. }'~tmr.191f}~~jl!.":f, ','~lSity i;1i[:irillg dlCimmlJl1R;' r,t<:;pO:Ii.'ii~. ~ Jbymmetrffir ·~ivis.]OIl yi>etdling e-ffedo,r- anllll tytl[:u;:h"1.I;I::;:i~lU,o i!Uhibit :JICi:~n p~}llym..::rizjdio:i1t t {'m'l..i~kll't w~th ii._.;. l:lmxi~~lt'L1 ~oe:lLio:ri Ma:.i:; ~li mem(iry·fat~dj p!r(igel~y, COri.8i~I(""!lt wi ~h the Ul:!;l.. Sfl~ rn;rl;l~n!i<l.~ i~~heri!,~~cc n:f [FN'fR W,I!!, hcfm-c in~<lging ~14 I. 1h~ 1~'S'ul~",~i!I!lht;',~~~ pliL,-d~{;IA.:-d mk for "SYUQui~L;!r.j,t:' cell di~'i~iO:!l in ofC'}'~okinq_"jii~ inr~~~nl T ~~b aHcln,lltil1,!! ttl obs~J.",,~"d ill Ih~ PllII'lli\,i,lPfQ",·~m.. J. d~~I,J~hkt of cOrljcil)t"d ~,.\,in ~dR. ~II~ an il~ ~c~,g11Iion Sf'M;-ci r>, iu;gdlillcnml ,a1i'!lghlli.'t li[,:t:L';S.iM: ob~l'vCd !I cl i'!,Ii~~(" }' i,,:IJil'l!g I;Qnjo~I'iedIW~I] d:ulIgliit.J:r.' M;.'(:>'. :i\,~~~,:r rmi~'i l~ puklli'!r\L:' d~~,ld19liighlet. whtt:h ptrnLOLl:Jie,"Udi~rilr iUi rtlclnj~ypLe i~'1I1:k!..Th of n C lli~~ijjil!lg 0 rew: [Uli!:! ,PKq, ~V(~l led l.lllfi~'t!I U:!il cJ1ecll'l'l m~.::I:1:'I~lnQ~1 cells b,e,tW ... X,l LIte 1)I.iUljnvc p'rotdi'i inheritanee i~1 d'.JItighltc:r T ,ecHl5 l11ljL'I;.>1.c· il~~~ei'lils.the ~~'I:atoi" ~:iiJt'IIkil'i of PKC~, iPig. "'). Emil~ il1, the inU1Jmne ~POil:;1!" d~c li!fli'ld lor proximal m~e1Jl~!a1diiiJ,J;ihl~i:li. re."iX~l:i,\'d'y. The quemto ~lC'iil\'i\.o hmmliile re;:,i}Ol~sc Ifig,. lA}, IFN'l',R, m~~I~1 be S!.'ClIIClOO,by {,[»:,'" '1' cc]ll<;, tlbClin· CDfl311d P.KC{,ro~siSA~l~l)' !;eg,re~!JJ,CdM oJ)I}f.;"'t~IUlI i\, e I1mxim9~. dau~l~l!rs '(poiiSi."!.."">if'b!:. noon: ahmClm1t Cr»::)\,>'Cit'e 1'~ug.:1i"ll :>.ire. ,~'mcl~i1C1ie"!':-loo i si[C ~J.1ughle~,in accord wilillill! syn!l:~'sC'-dis;ml :;.c:~vcs:(liS .. S6]" nci~hool:ijnl:J. Ilalmnil kil~ eells, 01' DC~ 12'), In m.1ivmoo CDS+ l' eells, WN-y gm~~tlIMly., <U1dC'xlWC'liSed km lecels of CD(~l loca ~iZlnion o,fPKC~ in 1)lfCmiwtic and mi~olic l~ut Iligllcr k\cll> of 'D(,/).' 'D43. ' '1)25. ,lIld :i.;ignill~~ns. ilUlllCC~ lite kc)' ~'I,'H~$Clll)I,~On t,lctOI': hl<liS"t.~.In conllaS"L cells lun(kl~oiI1S .lClM~ ho11>44 ~Fiy:. SA. li~, S3). ~. 11mm(: con.si ...~n( T~~1111:g, S 71. \'\ il~,dlis luufcr(!nti,diy c;"'iI~re..\lscd 1!1I1OO!-lla'!i.e pmli[C'mlij{lI~. n.'Il~.nIing nen :m.t~Il.~ (17), !ill I;iO!ltm.'\l., di~wl, in cfloor~r CI..'U~ ~'j)!:!lP~U!\;'dt:o, th~l~liJU10'1)' ~illf.""!1ll";; \viij! the ~lfccl(u' ~.il1(.~g,~ drh ,,~1eel ~di\ ision, ~~i.~laJ«I unitbm il1h,,;ril:!~fl~ nilo.~ '1;'" it&! k~ ab'lJll~ll~ CDiii:), l"d ,I ~ 113.l,n h iii: lilus (tm~c~v'lM~tbt din~wn~t.·:\. ~11I!11g'hU;:J.'" of CD.~"L!lId PKq 1,1;) I;r;t.llh drlll~ilh;n:: Irig ..lAt I"lmme more QI:;IIl~.i\o;,tC'1lIw~lh the ml~'I:'a~IUC'rIlO~)' in IFN-"l z;;~mHng c01dd dil"l,,~ly tlr indiroclJy Sinii ~,lI'n;'illdIS ~J;icre o'bmi!1cd rml'l1I, c)'ook inetie inllu!.\~~'C Ii"~"<l~imod goC!1C',~, ~iol] x [he (dill;: c,""''lmim:d diocctl)' ,~':,~ vil\'O d~n \~I\I,.'ronot eulli!l()'I.g,~~2i'1: ihese cdJ~ \V(lrl.;' ~m<!llcr,k"N:"gll..nUlUf. ~]Iill .;__.,..pn.",~'dhigh f\'!\,'\'!L~of CD6;;!L bllll li!$l ,dulJ~i~!(:!f (\ills dlDv!!!g .. n i~!lr'l1lJ~~ R.."'/flfm.\"-i:, T hlll!\.1d ill t}'toch;da.\.;_~!~ B UIll·S:;). Inw~.r I\;"\!!.d~of C[l'69. COLD. CEJ!25.a&ld CII):;'I4 Un~~I!II~~1 inh~l'"jl:!1il1re NlJl'I'Ih.. ~u!q~uihibiDm of UC! ~'b"Ylllillt!~'!rit:'!ii~gre~[ion'l of CDii': be,(pig. SA. fiig. S8\. We-;liku O~f'I!~'tIoooruill(ue 11r NOIeh5igfI:[1.~il!!:!. it;; mm~ll(:r ;l!lie~r.ilI haU1natk IW(~J:I cO!!joirl!..\I twin d3.lIJ!l,~lcrt"CIL~ suggt'5:[ud ~illllC~imml dis;~J21:tiiL}'illl proJd.l]~zbl: find di,S!rll d:!uglt:thm il. ~l1iig~!t e ~IDlis,~blcto mCllS\i[J,! ,nm'l! cytob [I r fl£!o'IInllUL:lliiC cell dh'us.iun "n hl COltiolnoed I~ r~DXim[,l1 d::!uglm:1'S cxlilibhcd !1ireaw.ii e:li:,. ml(!~rlC evideeec fQrBh~s; f!l,"Ccplor dis;parity inl lwill. cells, \.\'C llIbs.Cf\!i(.-,j p:r,crcreu~li;]1 illheuiMlIl.CC Nmnb in Ihcpro:li:.inmll datlgllll.cr, based ![m :its, pn-ss;ioiil {'If the C~ICCllor ,gcncproclucL<;. I.FN-y ~l;[!jsce~l~ dru:lIghtCY pop'll~miol'lis. l:n in't1x:loo wildru1d1 Gmnzyme B ,[I:ig. 56}, w:ilcrc.l'!1 dism~ d~lugl1l~ypel\,"'C~ph::nts.~.imooal dislriblltioll or, DS 1111\1 roscgre!o;llioll 10 (~lI£h!~rs I.honiI:II~Yil.od grealcr lFA"l S'l1lhling was cvi[]cm, in 1'I!1!IIkpO;ptll~:lliOl~S CD~ 'Wig,. 4). Illh\:"ril,ilIflJcc oFNmnb by p!'I,1):l!~~llmJ t~ ex hib.loo grcr<ucr C.XPfi,\·;:S~Oil of in~crlcuki u-7 ~1;)cpitQr 0: (.~L";7ltr~) m~!tNA ,(Fig. 5C'~ a ITlnmker of d.lugln""rT cells (Fig. 3fHln contGlSl, dm~!gh~ d:auglu.cll' was C'Q1l:sil:'1l£m\\,!i~1 II::. rell{lFt(;ld loof (.'<!r~y m(;!I!I!Or}' Pl'(.'C!,I]1i:OL'l\. (28), c,d iO'4!~im~ W the irI.muuldJngi(:a~ .~}'m.!I);;'lC \' iun in ~Cf T cells II1mth:.~ ~nl~~",(~ from ht[liunOO~I:!I~'C' A r~ic~ion of tlI~eseob~el"','miolli.'J, is lIun Ihe rro~ itb.TIlion e:-lhi'bil~dI~l uni modo II dis~rib1ll1im~ (25). DjS«}i'tI~ly ill. Notch ~ign8iling could 0011dfcclOr-fmcd d::mghtccr'S. \\i,l] M1l1frna]II),' dliITc.Ii!CHl'lll."'CMll'C NCilCh of co-rec!!pl0r stainil'lg ,Wig. :.In. TIle b~moo[ll nibl!lC:!Oo initi![!JI .r:JJtc divagence, li,g,mT!ru:, 11IolVC bO:11 [Olmd '!iO' be indlllliOl."'d 'Oll ,,,~ui!'irl'.il C\~d~l~ in dre III'$!T eell dj\'~ion or nlle time. without 11l1lbsw_[l~~ r~J;l.c'.Ilcmti ve tJIJ!"'C;~ ~l~ it}!, Itil '1;,'I[l1111:<""". !1l.cmoIy~r;)!I,-'(1 d:"!I:g~U.t..Wl\ \V{I!uld irm]1\ml! roI.'S11!I:l!1!!'ICIll"}", Ihll:.i" IKI\\~ ,"-~preSCllW r':)lt]~OJW'11·Slilll\d".'[(;(1 D " {!6j. like t~l~ n~)'[ll· ~1lJ:inlml~:'Il;'ml (k'\'dopm,cntnJ IIJ!!lSlii;:'i~y LrI ~(llh m~rit· inh~tm.l;(';- :~l;'n Ihr IlFNyNt, Ilmi. orNum pl'oximnl mld di!!'ilkll d:1lIi1!hl~ pOjjXilruci0J1S, wil~h ;[U ~~ ~::.:r(11ew and ICI11IIIlJ'l;.1Ilyilfli'T(~lli!lIJ;.':. l:(m~:iNlC!ll d ~t(j" tI:nd k~~"t .1IibUrltlmltt nf synapuic:: rO".... . li'- IIIf!PC"Lrc~ SpC!tillic [0 dnulgin.'1J1T cdl:s rt:~u)[u]:ing W inlec,tiorl. btX:fllUS(~ 01]1), ~)lnlIll'ilClric: M:gNg11wilh the meo!!!inizcd :lCinuuc." of aocillIDry T ~i h. rOTh" ~pccl:h(d,)'. Howcvcr.whiy Ihc pUiftti\lC' dr... ,!]9). COns.eql~elil~y, bolll~ clrr~or flnd memo!), don lhese proitdli1s WllS d.,;;mcl1.'i) inl~lc conull dfliilighler Y;iru... ItOI ~.tved 1.0 ~JroliUi.~e !'fCCtlffiOrs shQuM be cupablc 01' proilCc~i(m joined l win SCtS orilO.in£, lrom [LCUiIA.': homco,suuic :mb.'1I!illitialiy beyond d\'C nr..'1 rli\'l~iol1 '~Fig. 3. B and q, i.'>hlffi C'IIidi!n.1 31td n::mm.ihlS 10 be e.'i.p~Q:rodL d·i\>i:si.ID1l Fig. 4), Sl!gre~m l'cm ofc~i.ti('8l: COntnIJO" ;[llgi~ns.. a1ll acute Ch3~k':llige. whcrea.\i deln)'cd dmlkn£C should be bcncr c~ntmlk:d, by Ihc n.cmmy-like p:rlJctL1.·sol'l> (1.';). To test Ihis Iln::dkFiig. 2. BipulaJr ~eg;rega· lioJ1. we sor,ed 1l1llJ!llive prr,o:;,d11111 anld dimal !II liIDn of [Q1i!~ii'!ied r~lIdflt~~ll~CI---S :lmllfll'fimili)' recrIP!cn~$ nne! Imtlstcli'ir>\,,>(I ~~tci'i;Or 'M-~iiilietrk (flit dtClH tOo a new ~t o:f 1l1l1'V(!SQ.,,{)llduyrccip~l~ diviSien durirlig mitosiS.
pe:I'SiSIi5

rn

or

or

(@11s hml ~~~'eoted w~i1dF@dpf;etr1ts IW@r'!!' hal" (03 'Dr

L)'~

Vl"EI:e~ <lIS, lf1

5l!ainoo falrr LfA-1, Pl«~,

Fi§l', lA .mel ~greeliJ), So1iDbte

Ihm we~ d~I!Ic<r-.gOO Jlnrn~clioll;ly or 31 ,50 d<'l:Ys ~j;lcr tl3ns~;;:l' w~th gp~3 I~ I1WIWq1t)gt'lt~ 10 !iI."~'$~, pro!J.;,,::!kll~ 04}. AI 30 ,days ,,~I'lC1.' ~r.lIl)1li;or. df~11 dml!1;.:hl,l.'!N w~ ind':'icd (r;!l,uodto p:1'Q\'idc

~·tubllJnrrl ~b~lIe),"lIuli (!1i1d~, (.!)'~(.Qitiwtiolil (03, \li'~th lFA-l ,and!

~r

Sc Mbbte was ~~tvtdin 94% (n= :116) and

l!OO~/~ (I]' "" 10), or (eII~ r(lS-'~cthtllly.. PKG;, was polamdi (lWIlSilE! (03 in '9~/a, {f1 .:;: (if ~l'IIls"

T t'll:~:liS Wig. SCI). Iluwc'l'!;.Y, f1roxi:lilii~ daug.h~c.l" IJrovid~~ equal ar oon"';-.r pm'l~l10n, ~hflll Iheir d~:m~dCO~ ll.ClpJ,nlli when dnal'IIi.'I".g,ed immctliztccl,Y .r1~\G·lrnllIS feU'If ig. 51)). 111C ji.liIlclkmai pm[lcl1~es of d~ 1.1rsl]\\<O dfilil!:lhtct5 \'\,'CiI'Ii:iiJUl': roll:;:i~elJi. wiih ,our Ils;:.;igl1"
In~nt ai, t.!ftl'(!~;or .tLl1dmCmO~1 -prccuffion;; ml~h~ l~ili;~ of 1~K.'llC!iypk and flltld,1011JI mmt.CI]s, [)iSi(I~s:s;io:lii. U'le !'unclion ~ii' IhcinH'lUnIDlogical $yn[!jp5c. ;)5 it h~ been SiludklO in \111m. h:~:iop!maTmxl envg:m:atic pt)). ·rh~ cmcl;ging I'I."!C~ Q§ni~ion !~m~d~ p~rent T ccU ,1Ii~dDC ILmck;ijgo pro:~(l!'!MOO. ~1on~cl ~jl 'I'~ rl;:]"li\·c]}~ 1~l.c "Jt.lO't ,'0 iULt illt .acliv;iilioIQ t5 II ~ 1i!!l::Il11j'itlld us ~o 'eQlbi:dL:."t mhc ~ibulilY of:1 fih:\,':irMllily lnnhiOwl'i IlmiC~ton

~a1i;"~hl,,ibly b;;;,u,.:r pmlxill:lfhl d[uu~ltcr

1j.1RJI,~'(,'~in:l1~huu

''''l!);rc~liJIl1d wilh

1m

RESEARCH ARTICLE
F'ilg. 3. v!sp~~ijty 'im!~ry~apdc~f!:dp~ Ia rii!!/ pro:teiili! ir!Mrii~ll!(I~ ~$ dla_u~11H.@r" (08+ T o~lls" (4) U\rltdivi:llecl P]'4!JansgE!nrlc {Ofr' T cells {110m ]l1f,~dec:i wikl-'l.ype or I!In]nfie:cted R~gl-4- r,oopJenb> \llere 1~ll!ltllred11111 me~~a wnti1!ilnin.'g cylQ~h,;:Ilcl~ill1l B ~~r 4, ~Hmm at 3:1~(o ,ane!>t I~y~{ll<iliu~:\jis" t (j~l~~ theml 'lIer!:!' $tai!"l>edr~r {[l~ (gre>l'iii'l)',. PK(~ (rlld), IHubul~ili! (blu@}, ol!~d IDNA (gr,ayslCalet G~l~s lI!'nd~r!moilil!ll q!'loltiin~i$ "'er~ident1mBd by P.MiI1ollltl(1~cI (y~l'l" ~,!~s,UJ:!f( d~ft by brig~hm~l(lldi and duail fHld~i ~fiIld1 were 5c!(lIN?d mOIf .ts,ymmeVh:. :5legr@galitm ~d s!:a~rn~ l!"lg htwOO'1'!1 'c(ll'll.~oil'!ed d~UJghtelr!celts. Merq~s ill!' CDiS ""illih PK<:~, ""er~ sli.!p!!'rilfllllll!JiS~dQfltJJ' b01!h~he lu'l:mlfllil and brightln.il~d ilillla9le;5(~om~hOllila ~eWnIUTl l~oll!llmrnS, rrE~~(~iveM. A5ymmeuk s;eg~ga~~(l!ill'l'ff (DB ln wil.dl·type! versus R,'ag:1-+- mioe O(iCllmlltiim 69 % (ll =29)~!1ir.ru5, lil% (n 3:5) ,(!If t.winl5lE!ls, ~es~!l:th,l'~ly.Th~PE!!fl:,~t aSY'i'Ii!liI'twk s~gJ'e,gi:lt~(lili! lOll PIK(~ ii'l!a~ 6,2 % (n' 3:4) ln m'ldl-:typ@ {I!l.d 1,4% m "" :29) in Rt191~- rniee, For bmh IITlOleCl.lte5,~ne

A,

EI:

1001

I
CIFSIE

c
CiFSIE;,

oj

im:id:e(lcE:

oif ,atsymml~uy

\¥,M

in ('elQ~.rnlm 'wi!Qj~lypll conn:paredi

m elc(ul~.es a syli1 I1iIlltricall¥
l

sigua~~k!l!liltl¥,Br€!l!U!!'
to R~9'l: -m~!:e
whel1e botm C[IS allid

(P ::i O.OO:U. IEllcosmai,nimlgl.experimeiilits

palrlitiQtlec,

PtzC~s.egr~ate~

to oPP'O~'~!! iltJ1£I'I1t'e',minl 100% d

aJbulndarlCE!' or sVIni'!ptk pro·l'@·in~ lrl'l the ~irst rd<l,ugh~er if cells re!lpoOlIlla:iiiing to lnrll!trio,n. CiF.SIE·lla'beled ~ra Ilsg,enic:: ([[18'" 1 Icellilrmm ]rd~cted \'IIlldHt,ype lredpi~il~ were aifli;lllyzed by flow IcyOOm-euy4:8
~'Cl!.U5

(jJ '" '9) Q,f ~viltll·lYl1e l\lIi~$.l1!'Is. ~B) IKliUerefnial

CIOB
tr.am,fEmEid n~lh (righl uppl'r ~<ll'Ieil) are ~h(!l~~I1I. Re(!~,pli(lf aiJu ndaJl:lce (:t a:!:.is),(!I~ etocuon:i~illy ggJted daJught~r C€U$ (rewlild b~ghOO~t (IPSE IP~~oI1Mis;

o1!flie;r infection, [l~vhiollil, trep'.r,~r~too by <:fSE dih.,rlioli! U' o1!l~ist and ~e(ept~r alb1,lliiida~(e '(yaocis,)'~If.a:lt tra nsr~md ceib (le,h liipper pa IiU~1) a iii!:! CDS"" ,4. AsYli'l1!mewkinhe""ta m:~ 'of ~'g/l~detei'fl'll·

dJi~p~ved Oln~riDlfKlio n
~+

~~e'rUI[,I\el1rsusl'aJit:e' i

(()

'

Comparison ,~fCDS .abll!ndarl~@ ]~ da! l!gh!:er li!olilllem1'1lIn.: dlMfll)[ln hi!lht)"

.C~US aJri~]rig

F'Ilg.

lfI~nu ]n~h!!' ~hS!t; da Lll)!hteril' llfm~holl;yti"S;. Ct~hii 1(Ii Ma:ill'edi ;alnldl a !ilally~~dl al5: 1!f;l F~9.· 311. \lIe~~ Icost;[iii~cirOif .IIF,NyR .i!lnd PIKC~, (liiiPj)~,r ro;,i,iS) (lr (DS and! Numb (lower rows},. plus Il"tub:t.l~inra IIIdl DNIl, As)jmm:elri:.c _£E!9Ire'gatio n o~ IIfN~R ][11 wild· ly~ versus ROfJl-f- miite o«(um~d ]n 77'o/fJ '(ill .""
B)' ve~lIJlS 1%(n ::; 14:) ,oHWilIl ~ls, resp ~~vl'!ly.

Tine perceililt .a1~Yf1llmft.rij.( s~regalion or Nl!lliI1lbw.[Il!> 67'% (n "" !11m, if'! wi~<f.'lYpil'and 130/0('11' '" 'lli'5}if!l Ragl-J- mrke. FOlr ~O'lh IfNIi'R ,tlr'ld Numb.~'he im:il;i:en(e Qlf ~sym!f!~uy W<.1iS ~gl'!r~f!(al'lltly (jj~00'~~ i.ri (~ltrrQm wih:Hype i:(lmp.alr~dlte R,Ciq,:Z+ mice: W :!::: O.OOJi)., In c~s.taiililinl~1i!!~pE!rimenrts whi!.i!;le both im~wCl!~es.aSfmm~llfica~l~ partflt!l(lm~d., .IIFIMyR !i!fiIdi p~C~ SE!g regate~~o O~IKIS,1tt!d~U.9h~lf.rs In 1.00010 W '5.) of wild~type t~'II]n sets, ~~nile (])8aln:d NUlnnb $e-gJl.lgalied~o the same di.a,lI!gnt'e!r'f n :I.l)~1l:i {11"" 8)

I~

b II
1:1,

'~

I(lf

\lfiild-type' h¥iir'l!iie~.

~:

for: iJI~ ~~,'nrips~1l mnely. [hiM it hdpt<: ooordillnlc <I'IsYl11bm~iric ('ell tlivit;lOi'i, .Ai. iiS .s;ll1lpbt level.. [I

the
ti.ODl

I:lsdiJ~ ,dolll.:tlu'OLlgh

tire ~venll th<ll.oUc~foctt,r

bm.'cih.mi:>m lor .J.i>ymnle4i1:ic cdl division mjl~h!. rrovkl\: J sohnion In an ~~nli.dpI\1Mem racing .:JI{h.lpth·,eiummmity, lf'an irm:nlill1c ~]Qn$C were

n 1.1\!'"'1nmy lirn;.'Ig~ ~n cells die. Ar'iC1'~solu· of in~lcclio:n. the !>IQ\\'. cyi.()kin~hivel1.1Pro-

~o~gil1 from .he .L.cli\'a~ionand luolifcrnliall or a sungle .1un~igWl·!i1POci lie 1!1I~'\'C' T cell, simulitJ~
flC<li!,ll:O;, ,~"_sigm;l'!eJ~.~.

~ill:eli<llkm(If~lCItlol)' T cells mistll.lhc~1 Irl\'(llve f;ym~l1c1ncdivi::;ilonsl(:l, nlifnlairl stalble nunlbcrs G r ~,unlila:dyl!i!hl,'!d {_L.1:lIghllcl' ~1Is. This mug!11 be

or c rfQclot

,Ikful
[I\,'c,

~tl

Ihe :$>I!n~.l~~ic ht1l·m~osl.'l!k divi~,i{lll we

T CGIIb;!I!".u"f~f!ij,;d ~o ;unigJ:l'.!1. ... N\i"C0U1d1wlJi ("pG:ifl1~!!>'C'ni-pn:~'uil:ii1~ DO. 1'i'iI.ignt

;,JIllti n~~~lOI)'

J'IiI'lp"

(Il~'<C'~"'il(;"dfh:r. Il>IYvl; SiIJl'e hi llil:: reJllt~al.

di Ir~'IT'i11d~ulIgfi'lii;l':!i; ~\,li:iuM \."~I~urt aeure dil'iiilJ'iatiOll or diCIiil'iit:'I'O~~,\ilhilJ.: tw=rvial_g

C'xlje;s ~~1' ('I ~",

11'mphopeJii£ tQiict. lipan

(!~illi!I~1.: lJi1JiJ!i.lK!r episode of I'k.. )'llilnK~1ty ill.~h~ m.cmo.ry J cell, leadi In!£ 10 Li:nnillfLlly dl m;:rel~lialoo c'lJroctor :Iiltcagc'. as ,vd~as 3. !9C'11:' re1i1C'>ving l'I1!l!mol:Y ~ineol;gc" f.iucil <I u1iOd.;:~ is censisl.cnl withthe stlYil cell like l~llQm'C$~lall]g\,c been rroJ~(ilscd ror 11l1CIU(1)' T eel r.s (]!J, 1/1 mid is DIN II!!~~we ,Ire' CUILTCI\t~y inv~'tlliw!.ing by 1..'_'\,arni!.1i ng tlru;~.iOTphnl()M¥ ;nidph~nlJlYrrc (If n'i~1iIfi.(j~)' T cd~~ ,~Did Ih~ir &.\lghl~!N ill 1111; 1\,.... ~nRJ.: to a S(':C(l~l(huy c:l1il~.k!Il!!F'. \WhJ.:~lier Ihe

nhel1

1:11,

RESEARCH AR1ICLE
Fiig.S,. In'll:!~ght(lr rilte~dllltiJn£iimmllllliil!)l.

(orr (~I15~cqiJ!~ri!1;g ,c1lsp,ari:'!ltea5

lo~l~s;\'IE!r'~ ~~all~~

fA~ CF5tE.t1Ib~k!d CDS'" iI' in f1g. 3, B ,~F!d (",. with

A,

,Aoute

alnil:~'booies~gain~: (08 and ~1lE!ofll1l! rcUIQml'l1~1 lm,olecllIl.es: CD62:L,(069, (043, (D,25. OJ (104,41, PhieDiio~wk ii1ila:rt(@W$ elector a ~d memmy (:eli5 ,Ut ,of <IllM a mdl (IDS a1tmndlall(]e (y o]lis) of eLe(hollliaJIU:;

m@~~s (If re:t!l~~JI'Hlire:1{~rs@tfa.I1Wiptiol1l=poty.rIU!~1i!' m:7=l!l;L;;...-. proll! ,enailill readiolll hom serted proximail and d~sLat I~' dijs~a:1 d~lllght:ers.~I)), $Ol1ed pmiltilillla'l. and! di~lal '~(I!JIgliI'IQI Q l~ir~ \wrE! tr<lllsilerrna nlilto new Ilill\re rli!c~'iJielmts.." IJintranderre(i mllll~ml and redpi~,nt mice :W~if1e 1':Ny ,chail~en~edl~lh l. mOiWCY!'(lgei1es: e,lther inlim~dii" ,atIi!J~(Acut!), (IT 30 days W@I~y@d~':!fi.!r ransf,~.r+ . Four days an~rirl'rE(~O~, q,l,.I~otiil~li~ b~d,Ni(ll bl;!irden:s 'iii! sph;!en:s \~me' d\el~omned.ME.'alnl ~cterial burden (If .ue(jpiellt iiIiIj(~ (It'H Ull rigi"'!;, 11 :=3, 4. 4.o!J} 15 lE!'xprc!i$'~d 3:; ill ~@r.celll:.1lg,i!' rl~13t1lv!! to Um@~li! Jlti@1'i ~r !!! !lIra n!ir@ffedl (.QntJ,(!illlI'l'ii!:~. AH '!Iir(lll~!! w@~, sn9iili~ici!lntl'l! pr(Joo(]@d. J!l.Clt~'Il@·OO ~fI'l!I'ans:l'@rr@~ WrUfQ!S (P $ 0.006). A.th'E!r de!l.<irysdc!hill~mge, distal daughlhl rs·aiiWomed sig llifh:amUy bei'lt~r pmt@c:tiiolm lillam pr,o:K!lmal da~ghter~ (OJ' ~ 0.0.3).

,gated daugln€r c:el!ls (3.'e(ond bri[l'htest (1F:~I~eak p I(l!o:ly)are sh!ll,\1«I,C~U size ~el)f<,:s~il~ 'r(l!rw~rd Ugl1t :s"catllw; gralllllJ!tarity Irllpr~wlS side: tlg~t ~U~iF+ P!l;.Io:reSi:eiUe iI:!Jre,~~i,lY ,of'5-:;,JFfface I'!~~'PW staliniiiigi is ili~e~ ]11 fig!> sa. (I} G~lt!;\!,t@1\e' 3 !lialv·zedi3,s in W~ wil!.ih ,an:tll'bod~~ to d:eteclimtr.uellul:n IIFN--:f IJI" IGr,<llnlyrrt~ s. (()~l~7RIJI rnRNA \llollS m;l1"a:sl!Ired !:IV
1

[II

:tJ

~S;YflllIl1lCI.ricd~HIg1l[CUSof n~I\,'c T ct!:lls ate aClll!i~

S« cc:i<llUzcd Sl:lbS.~IS, oihcr !har~ c'llcclor and memory cells, l~ml'ID~,er. cm~~l()t 'c:li,dmkd atlh ~I i me, ~~e Alth!}l!~~ !he pro,g,eny or a newly :!clivahxi CDt' T C'l;!11 mill,}, I[Xl C<!l't<llllc of'[~~\ iding~~~d! difa ICrl::JlIiming witbmn Jhtthcr cxpoourcw m1fljtgeJ~ 1(32. 3.1). ~l has bc!!~ Sllggcstl.'d l]l;JJ! cllidcm I D4+ T eell fC!'POl1~ nmy rotll!ire cOnl.ifll!oo
l.iiJJ~ HliD.cdte oooom~ C'XBM'J.SUIfC 'IInl.b~!l hy Ih~~ urli~i1d d~.Ilugh~~'f I(~ ~1If.i 02, 3</), ml!liill,g lil~ po&...ihil~ly 111m. 1l1il0'1'L! (ll1L~~

mighl be n,ilrorediu! Ilhe bc11flVior oU'(nher tli· \ idh'lg IYilnphocy[i,!'s and :i11 d'le h:u.cr::tcilKms of o'lCtasDmic 001001" cell~ \~~~h~h!! c~nulmr orexunc(:!U\~Ii[r Oo:n~[)OJbeln.;: of t~~eirprovisjjo:n[1 !lichc. Th!.'I'C rct'~mil1 m!!merouN :!"'~';In!~s 1:0. Ibm~:ln)' 19:5! [h~ B~W\<~cm;'C' ;u:nd~mptnl'I<!lU;C ·of rnij'llUll~~~C' re~1 dh isioain ~~)li.VC' lmmullh}! ~]jJ;clinl mMlUlfmJiall

23. ~ III. ~t~lll.cil~ ~t,~I" IIltlt, ,rmmtlJlDl, 6, 'll2:3;fj (200:)~. 24. U. MmKI'to, Pl.tt ~n~~mfli1; T. ~0I'II!r.~p,,E:. ~ W1ie1l)'. 1. 5. l ,~~~; J, I~"t 1.'i1'7. l'51S Qa~t ,2~. A, tC, And'eF~~n ~I",1". J. {.mmtrffl!i'rl. 1.74, (l9~ {.;!OO5:J'. .21\. 0. Plri~1!II .~ Cd 1:1:1.,sa5 ~004!, H. III. Mmju~~lh ~ ut, .f. C1ir:i\ {t:i~. 1I.oa, ;614 t2(}0'11 2B, $, ,M.. f.aern il't ttl" ,~tfl. mutlUJ'lt1i ~, li'iii. ,(20tU,. 29. o. T.I~~roo, 11', Iili:mOOH, s, [):, agmer. SMff' 2.9,;3, W

II'..

dcvdopIllC'~lIand ~IDI1IlI.cos~~is,:i2.xlCllSl'ofls,oflllc rmHl::\,,'!!J1ik 1Jl1I.."9::'!llccl here mi,gJlIt. Illus, C,"'IlMd [he prKriciph.: of ",,!;.ymn~Cilric cell division !o
ytJ:r:iom .~1~1~1..~rB :1L1'1d l!},mlplulCylC inrnmuui~l)'. o I Or 'fJniI\\irk illiSigh:t iri~:() hc cl~If.iUVl!..!'tiroUIl'lliMiIIECS L prr"'tlt-:ding <Liyun:ut:;Hl:t div Istrm of henmlCipo:i~ it lind 11Ctlp:l!)Siic ~erl'~cells (40. 4!).

~itcil~lme ihc diversilIT:calio:ll o:r Cifl.'i:ltOii ehoiees ~T helper cdl .~ !~TIIIJ, '1.12. ,md T~"unlin, 00,. d!ittOil ~(I",or ~tlslcad 01:.s;ilillp:l.y !:"x::tl~titlSc~'C'~r am'! mCinory I)ro~ny. Intlllils \\<1)'. tll~e ~'!pl.'ttI\lm {'If ('04··· t cdl etlktGr~lur:.'>,. wh~cb arise onl)i afler QC'IIdJ,visiDfl t151, might all be IlCp~nli.cditl ~h;: ~nhial cllolnal bmsl. Th~'I,":,al~CI':. only i~le 1llOO! ~!sen.l~1 f 111C' d ~~I,'l'IC'1W'c1'g~n~ I~.lighl Lmcil::lb"O o ~.l!filhCf :5C1b;:lt0I1, o,u 'the basis of~1~ tY11<: of' p'~'hQ~,cn CIf~OOlUlt{;;1'Cd, Indl.,'\\d..~hc dl;.'1-e!Tlliu,lIlt. ... ~]Iln ~1,l :-;t];I1;j;l-blll;;o,O in lhc ini~i;d C:DR~ T ccU di~
vi~;oJl ,IrQ' <lIM]' ~:l.J\.';S!I,IJ1,1pli;v{,i n.:g~lilal.Qn; M CDlI,~ Til ~.T:! 12 I ~nt~lg,c ehrii.("{; l L5• ..2'6., Jt$-J9), \~titlil c\'cr-i Iilipl;{I!{,illilg ii~(;1h{ldoJlog)i fOJ .uII",i ",0 ill~ag~ ifl,~ill Id Ii i!I~ ~mclfl!:l'it n~y SOClllne p.")~:siblc ~,oOOi":l:;-~l\lCt a {lclaillr.)Q IiiLtC' n~p of a sifl£lc T erell donal, bursl. d!lllrimg iii \ <lrie~.y ofinllJi..me clm~I~nb.gcti. Par I;IOW,. :lmw~.:"!'!.T. 1lIK: ro.l~ (II;' 3symlt1::irit: cc:U divis10nin divc[!;i ry ing ("04·+ T ccl~ ffllc r ..fIlillin::.: ~!iI be IIGmK~II}' c:v;aluatc.a:l ' 1\lIholllgh iurmn1aii,o:n IU'OtIl t~le clwironn!C1'I!.

ouw rotund 'L'}f fISyll1.l:llto'1.ric diviSion miglu occur, Rc~[cFo:r~i'\JIe m~L!'J[i~,oil ~i£)'IInnlCll)' daring dtc innliml donal c'qll:l!1tJ.ioc!.orll OJ'I,'" Tedl m:lgbl, i[\hlS"

2i1l8 (l;QOD, Lfli. M. ]. MIII1!~,~ :5.. 9i~MI', J. rttH~J'lfl. 1.70, 117 QOO~!}. ,1. C. j,. ~111~~ e-l ,ilIl,. I"{~;, Ni1N, AlI;'ll1d,.~;. ,v-S,Al 10~, l3Q.1I (1006). ~2. 5, M. ~e(~, fl AhmtL Hor; fmrotli'Jol. 2,~:lS fOlOOill'. H. M. l. Yal"! SH~f!, E. E. IlI!niITie~', 5, P. 5thO~ber9e;r; mIl fflImuMi,Z,'::Z) '(£OOll
3![ll j.

Re'r~reJ!l(:e~ iNo~e$ and

1. IL 8ellldliD1Jer.l. :II.. K~idJl(.. (lirr. 8!'Q.!, :L4. 1R671! ilil:!!li!Il$t 2. (, It MOOits, B, A. IFrl!!ibi!fg.. iii, ~ilI,pl'i!r, 1M. S(&II~r, A. IDIillrer, N"owre ~~,!i,a,2 U:9~at .• 3~ IL I!!, ~~!!Pi!, "'" ""~rn.1!:r.. C. 5ll!mm. ~, .M. r!l'~t l~f, >1, 701~'t200:Il'. 4. 11'.Rt~~hiltl. Il:. L.. IMnl'i.ml1~ Eo. Ing:u/lUI, It l~nfi:lmt HI, 1, fmroUIJ~ 1166,.4~13 !2i1l]ll'. S. ~tol~ j. ~~'lil', M. T. fl. tiri. G!!maim, ~df!l.ta

I" .&p. med. ,~~1j 1515<S (~OSl .~5. ].. ~. 81rd tllll:, uruOOlllO' 9, ,20~9 0\1'516).

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,t

5(79 ~OO.1l;~.

8.. T. !t ME\mpel, ~ Ill'. "'ie~ritkiOP1. IU. H'. YI!l!f! AAdriiil", ND~42:7, M<!i '(.H!OU 9. :5. H!lI£lUil~' fl ,r'JI:.Nat. ,1romJ'l.!ilrl~ 5,. lG'~:5 {~04}" 1J1). m .~ Millf.r, O. 5~rrina, t I'arile:r. ~ III (,;jl!ii~n, }. Up.. M'm:. 2\00, B'~l!' {,UIq.~J. 11.1. G. ~1iI~lihalf\lO' •• Mat hllJII,(Inol. 6, '07 ~JOOS), ll.2. S. Celli, P. IBw.i"IO, 1..Ef,p. .~ •. j!~ 1271 (;W05), :113. i!.!I. Iflt}mm~,. ill. ft)'trj!l1\ J. Etp. t.JW" lL:91, ,269~2(1O~~.

~6. M. AI~~rf.an et I!~,M:I;Jimm.ru;!c1. 3. S.~9 '!2002~. H. I\. Jl lts;m.~1 ir/ oJ, ~~ I'II'tlii'l.Al::Il'@..Hi_ USA. "8, 1I$U".l ~~jj). 313. f'.I!\'nti!i! el,g1., Pt.ar. ~l. ~ ,Sit U.s:.A,.:!.G2;,9SU (2:1l11)5). 39. L fll J!J DI•• }. &'1'0 NfM:. 2101':, ]:0.3:7 't200>~. ~'O'. S, .i, MD!IIi~!'!., ]. Kimb'l~, NaiiNe 441 1008 f2:006}. 41. l.(- ~llIg, l, E:, ,!)Ildl\, 1~rJJ, .(;e11 ~~I.U" ~1)4 ~OO~), C 42. w~ iIiI:i.nll: M. 1~1~i1ml :lild E. WiliII'fIl1lil.ll f~ nil! g~!i!~ U5!!: of ~!l!~ a:;5i~ti!~,~ will'! t!:!e ~o~r~c.J'l mi'HOSCqpe-: r;J'. nbmcill,. ~, N~flhillJp, A 1'i1i'~m, ~ P,lmn,. L ~~" $..OulJlJ, run!! J. t~ffi::I! f&i 1r!~ill!ilfI(~ <lml!!Iiil(u~K!!n.SlIJ.P,pllmd tt)! riIH gf;,m~ AI~:237o. ru(l~.3(l21. ind I!iI0'616,:!l\l <lilt! .iI'a~:II,'b!iltli~~ F,ii\l),ibi

am

z. ~~
~n

(S,l.IU; MIH Q,~!].Jj CAil.14114 ilnd A!069~O M.WJ;' "!H t~!iIill!ll g!i!!iIiS !lll:OOi)'Q66 't1.lCJ..II,[OS,5.1i1;2"$ 300 G/;IOV1170 {'!,r,IR,'l'j .1f!,HI(l75(3:Z (A~,u~, (M.:lll.'1.2 .1Il'Id (A.B,). ~M th~ IPtltJ Mil ((a[l~m 'Coot!Jlf i(enlre. Nlalioolll He-ill!! iiiM Me,jiKil R!l~IiI!l1lr! C~II!!riI • .an~ "1W~I!iLlijf!l ~!ld (o~ng~ 5,1'otR 1 'Sl!l!p~oJitingl O!l!tt!i!E! M .. teip'!,!l
~;!Ci~i:I'i!!g,Q~~L!'etillll!nb'rillLIIl13Ij1;1Ij131DCl

14. MiI!eriil:l5 ,iJrnd m~1iI5

,ilfiOi

awililbl~ ~~,~~,porling

is es~lttial in: choice or re'n Ihle. the la:::lIIi~"Ltivc 1l1i~ia~ 01',1 migrn1ir~g I;:cll, Ims gfl.crJ I1fmde .~ d:iF fJcuh w ~dcn~i ~l(; n1GS1 criH,:!! ::;.iglmk It it' ry ~"~1~~!1_gIf .I !;.:pt;iCL.d ..!~~ii1l! thl; l]mIDn~od ~nn .'T~li:d~ . witti di(: all1ig,c~,-P~~!1liJlg n11, whkh sQ::mi!igiy cootain'li,""f> lill!: :It.)'nlilticlty rlf 'f ec.~1 di,,'i~lJlh.

Iii. iI. PIMldornad'o.. Ij}~JI iAliM. it Sdin!tlB', L Iii. Gliffl'lttlJ~j'. NalrM ,1131. 52.7 (2,QQ4~. :ll6. m ,~ Ii.lJtlfrl'~~MIi!nWil~ m" ,NIlIJ'j.~nil}l 22; 7H 't2Q051 fii' '11.7. A.lwlJ~~ :5:. Ollmo, J. (l~ tkf. 119. !l!i9 j;:ll)Q:ii~, ]3. It IN., (I«lIrall1l ~,.I, Be~~, !.JrnNlIil~ 1:!i. la3 {1~9'!1)'. '!!.?, m l ~~n.ll,. A.5wliimge!'. l!j't,tWl..r 304t 6:l;? ~~',ila"l'. 20. f. let:h.1t!i', IiIlW, YDWft4~7. l {200'j. il1l.. i), PHiigDrma, lM'!r.ll)nil~ IromJl'lui!li~'IiDn. 2:2. III M,. fiJUdrt (fl al., l.r:en.Os. lmmtrtml.2Z, 5"l~ ~:2OilJilI~
jj5,

ma1!ernfll

SdenCi O~IiI1!!l'.

Mllwli!l:lilfldl Mil!iIlhJXl~ ,1':'91" SI t@ !ts

'~~I@.r·M~~ ~ I~~m~r

n.,

~OO6,;

~Q~

e, F>lilbl1J;¥~

~OO7

'Pu~li~Ii1M1I!'111~!!1l ~~(h :lUr: lOt:!.ll61~iem::e. Hln!.ll 'iii'ldll!l~ 'IM~lmlll<lm;1~'OOIWh~n i1:d$ pa;pBr. dtlii9

1[69~[

IEnlh,Blnc:ad Bondiing 01: GUild INanlop 8rtiiic:les en Oxid:izle,d TiO 2(111O)

l1
I

ing

The TiO-'!l ~.~ ii'wr"f'acc oC'9I1..~isis of i11.~:mmII~ !"Q'i,\,'tl of rh'l;!ro~~klooroi~p~~ Tj l5f"'~.J ..tolill:ii
U7i.f(l:r(!!!d-(:oo~il]11Th.-d

:!I.l1~ pmlD1,.!clal!!l, U'li:yg,i;:n

(Ot,.." ~tmi1;-;"The

~h(; Ohr~wms .u'Csi",(old conrdinated 0lli5 ,any Qlhli;l!" l~ .... i"l $!okhioJ11i;"!Jl!C !iUtillc. An Ar+ 01"11
SpU'L~crud ~D'Id "~LC\I,;n'l1!""iIl!IIi~"'~k'cl"~{ I.•~0) CI)'S~1~1. 301 i'i-lypc l>C'lm:it1Q:11I1 i. ~.ii::lot,iUlili ul>; i>tiIl1:lCt: Im.~a. Jtlin'~r or,~ \11Letlltcj:~. (5,. S'rt\il ii1~!l!lF' of iI1lc Ti:O~I.~IOl' ~.rJil.c~ .fun: (kliWill"I(ld, by elcctffi:l,k ,ell"ec!s (~.c.,. bri~~t ro"V"~ C:OJ:i~_"" !ipO~ld tti '11~ctil ttol~I'$), wh~ttc;'li."j;o!.!!OIIl.'C,tLiica1ly ~rndil'~ iQ", a(('ilw),f!I'I'!(l:u d'l.Lk ~5~ }\,croru-

Tu :IJiomsl!llucll,':l11C1Hh

bri:d!;':illlg

D a. MditU'n,e¥. J.G. W<liIlQ~ 5,. W!!ll'Id~~ J..M<!nhie.~enj it Sc;lfrI<!Jub, E. Li!9s9<!:~rd; B. HljJlfiIIl1TIIl@r ,Be:5~l1Ilfu1ad'l@Ir''' IF.
j.

n~~

Wesl:llIdi@di thi!: rJlucilE!lIt~oa1 of golldchJI$~@rS 011 i1']CJI~(UO) S1Jda('~ ]11 UU,I2@ diUe'rellit Qx.i~.i3itio'fI !i~d~@S b,y nrig'h-r@sohrliDJ'I 5c,a,rlil'l']rtQ tU!lJ'l@l:in9 lIT! i(r'Q~(o,py. ll1C'Uiln..,~ 1il,oo:,('llllO) SlIIpporltS (:l:1os:e~ !Were ~@dll'C'OO(n~ri!l9 bridgil'lgl~~ygl!f! V.1lU~IN:i,~), (]O ny~r~~e'dl'fh~vinlg Mdgi.lfIg llydr'~iI::)d grol!p~~, aJ!'Idl (lim 'o~idii~.edl{I1!11!f1Jf!g' Oc:(yg:e-IiI, aJdiatollll!s), At ~~om ~1!'''li!PIUalU!lr~.!Jokl oanodul$ter:s Ji!~d~~te ' ~'omog<eflel:N;JiSly 'ol'llihe-ter~QI(e~ ofU'l~ ~edl!lced ""i'l~ I).!(idil'~ .5!UIPPQrts.,~'lih~l'\~sM the hryldr.;iI~OO l'i;Q2(JllLO) $urra,c;!!" ,ellll$tef~riO rIiIl pre!fe~enti!il Uy ~t(he sl~p 'e'd!ges. Fl'lOlm ilil.~!lirplay \~.th d,~,f'Is~~ •

iIlSI),.

di!C'

Illan;l

:l.ll!Ot)1 ~lm,t ~'O:IU1CC~ 'th~'

bri,glll

11

h.!If'!CllioJf!al th@QfY ~al(;l!~tio,ns

.• W€ ld~{!li~i~d tMio <iifr~(lon,1:g(Jlrl~n02(ilLO) ~dh~~i(lIfl'n1Iocha,!'Jli:;m~ror

the- l1ooLlc~d and ~\Ilh:fri~~d 5Y,P:pCuts. l11~ adhesh)1\l of g@ldi dlistil'~S is stf'CU'lg~t. 0'" t,h@ o!idd~~d
:~uppo.nt~ .;ili)lcllihe' ~m ji](ij~~ti(III1'$ or thi~~'nd]i'lgr{lr {(I'~ tylk ,~pp ~katij 111 ,dijS(U~5ed. o,ns re

OXII:J!C ~i.JlII~POIllS O. J) has Slllll1ll.ialCdi C)!;~ !.cfk.~h·cresc>llreh .acti\'ilic.., {1 .'i),HlI'IU 8. gcn.eml consensus IJOW C:l! rlSt.<; on :SC\'CfI1 II asp-a",""!$;or Ilhis ,.,-ystcm. TJIC' size of Ihl!gQld pm'! ides ru[:.",'1;:m~ i~lIy<llfoC'Cl.. ~h~t:"<1J.taiy~ic ;u:livity ... ~d~he go!d clu s;t!.'~ Xml!'1l be ~rm~h::r Il!>lal :5 ~ml for j~igh~ cmaiylic JLcliYi~y to OOClH ~~ .n. The dmkc of ~lIc o.. xjc._rc :mpllOfl innl]cl~CCS the cal~iy~ic ee~idty,so ~lerc isn s! ro!li~ "S'llllll!On clf\.'(: f' iun lJIlrd ilion~o th,c "Si7-C' (:1l1'<;(" l2. -s, 9 J', and red ~~cihl~nxides are O'Id",,1]1 aM4;:'o~L~1h'ii COIlII'p;mT1I;-d

-T

he '0 !£em
(~n~!..s

or d~sp~'Sc-d goM

~11)1' 0 rdh;;AnlTlc~.i\, L:"

cmdyl:h:: :prop.-

I1lll~Piiliii'!ick~!>ml

t~11JlPC:f{liDU""'S. as s.i,own ~11111(: Ijionl,~.riltg snldi(,~ by U:lJ'lJllil U. I'H. GoOOUiiiii1l7. U'I)\ 3t:i.(1:Itll~,",,'urk

or !!)theil'SU!

23').

~!ol!!gh.s arise "'Om '(\.r \':.!C'.mcic{'l. Wig..IAI (1'4). S!1I!rti!~!ll \\'~dl n dl.'Im. ~\:'~dWl·~"¢d. TiO~l~ un ~/'"T~01! I ~. I),]1 ~urn1;Ce. \I!1'epT"~JI"fed twC! liurlil{."r [ ~~,\!'n-<ll.:l'i:rlooT~U1.(]I n I surtaces lI! nd'cr 'll~~rnh~,gh '!,':~'C'L.!lI!m l"(mdulion!' ,(24. 13}.Pi!l'i ••, \%'(1 :proclu{:~'d O!l ]!ydmt~\d.TiD'll I.~O) Ih-TiO~I'~ IGl'1 M.ld"hcc with U M3IO~~l~.~.u~llilTlg~0!.1~ o:fQh~' (lilr ~U'tl'Jn. b~ . kuing \mHiIt di~:;oci{lre in ~. \r;3c~~ci!..~trill. lUl, The C'JPI'Iin!!\[I filQl'l'nS .!onrn bmklgi~l!1\ hydroxyL~ (01 ~10r)~ln'lt~ppcat" as brigh~cf Ilffilru-

rediuoed

~{jl'lo.nruclll.ciM.c

ones (9" Hl~"~,lowc'I/iCr, 1.11,(;

rcl.'iI.uoJ]of the adhesion prnpcrlics of l1i1lnOi;I1:JOO .!',loM whl~e.1I.tlllylic ~e~h ity is still llllrrc .. solved, A number of studies (fI~fllinly C3tiillysis wOTk dose 011 hi£l~ area :>ilppol1 Illmcrialls) 6 ~.LVC ~'epmtcd that s:lilghl.~ yoxidiz.."".d gold l Alit }. ina>ddlilOCl',1!I III ~m:ta;mc sokl (Atl(!ll. is,il1ll)()r!llm1 W achieve high aJ:ili,vit)l or di:Sj:l!Cr.ilCd All cm.aly'St~ IJ, J. n-M~, I'll pilirnll~el•. ~yci·i111:!;urracc science ,slud!-t..'S (11:1:' modcl systl;:ms r;wOJ'iCd an i:UiC"J.1ll'Ot;lI'lonwhcre Ox,)igeul vacancies Imv,C' been Invoked as bcin,gt'csIUJnsib]~ ror tlilC' ~~a~ l~i~i<,:.;jJ!1 Kln<J!r~cl, :iC'~i'\'ily fir .1\\1 !;::i1.lsh,;11;, on !Jilc !ilL!PO
.~~mt U

' 2m.

\'Ve Nll:.ldlicd:

t1i1"~ fi.md..nl~~J]!tlll n:J~dl~Ulli~H Il!.~ of"

rI1C'[,-d

oxidc-:;.uppollt

adh~iol,1

by

1l1,C';nll:s

or-

~mliIJ~

I1m:ll~lingIl~iicrol\c()tly (SiM) mid den:>i~,y rLlfl.cIi,O:f1alllioory "DfT) c.IIcLlia,tioI18.\VC COmpiliI'C I he 1~~I,clC'i1~io'nl r A!IIJ cluners o on Ol1iC Sl1IPlloOr'il lTII."9.~ri>l1 llml, Imd been IlUCpafoC'd in dirfCR:1ti 0.;\ it:!~in{)Jl :jHlit~."rVc ehose rutile T~O::rll !0) 11I.~ the mood suppmi becsuseaurrercus snrdics

ex ~,,! H the AurnO-,! moo'c] 5)1~I.em •• md Al' ,riO.: Fig •. 1. SIMI ~nl!ol~@~ (!if f" (pJ', h-- '~B), and .~ (C) TiCJ,j!(1.10)'Sl!ria:CFS; be,f()~ ~: i~xpoS.\.!rc (130 x no A~. O is " good C;JJ!:Jilysl, for {Xl oxida[im~ 0!I1~O\.~, Symfim'js 1l1ldltrilIP '(\r l;!il.\,II_Iln(~e1) ~liJare},. HI ad:atD.m en '1\r5il\~ Olf!wagoN, .ami Oc!! <J~:otlI1IS in 'the T'i uallgm {drd~~. IrI_~l'S3(.1 x 30 $:'1 s~,~~e poiru d~oc~ a!' ~aM~~ .e~b f'gi~d,~(Jr','~ ~etailW p'rorl",~:'~(li'lrOCIJ)~s, ( see (25). '~D'tv' II) sm fJ"llag e5 an~r .3°10 ML AIJ e~po~w!~l RiIi [1 Mlis d'etf[nE!!ilo'l51.387 x OOrc' ~1:o.nfI~ per (mOl I nWl1di~rlpIJnm![Jlry tbOOscll!ll(ll! {MU!!' ~iIl4A!iQl}and Di!iPi!Iil'· a lIl~mm !Ji~P~i!:s. dllllil A!iIH!nOIm'l', Ulli~!lf~~ly J!.lirlitlJ~ ID!(~ ('Qr~pot'1diflg~' AiU'llU/l. II'm~ge .~~ are iso xl~O A2 [([») t{ll ~F)ll ~ 3$0 x~SO il,z ~~G)UI' ml~ '01 f~oct~ly;. ,I'll [(ID) tOo O:n '~1le OO!~~t~ of tlile .Au dustl,lrs ;;!ro~ QI~fi, by (mtooT li!ll~~ :l!..2', 32, .and! S"ZA it ,eOoC! ~m~<$ c. IClePllrnrt.. aooVl!' Ul!! t.e"'ra~e. Oir e{liol1l!5.are as im:tiGlteaill{G), ilnd 11.!l1lileL arameters Ul.!lllrnefitrl9'wn~J1l il.:s: o;m M" p ·Vo \~hom oo~~e~p(!l[nl1fen~eilllIOuI_dIbe add,re5.'Sie(iJ. IE',rnait
i

fme;®in~IIIHik

UJiMl11ting voUa;ge 'lI~ 1,2V)a~ '"

idelii~caldiirn'~hout

tlile p.1poi1!f.

1692

REPORTS
sions between the Ti ITn:v:stl1i;m~hrl:iic ~'b'$,"iQCim~x! \">'~~l the O~\lm;l;11nd~ (.U. 16),. SL);ilIIHi!, \\nC pn.Wq-~'(1 m~ o;xidiz!;'>,QT~o.~U !O), [t:J.·T~n~ W)j ~. ~lIlrjiloL;: l'Dy I~ting '0'4 dis,,,oci:;i!f~~~ Qi:o' 'iI;,l~l]cill,.""
{!4, ..?7), rc"uhi.l!g in JIll a-Ji02{1.~n) stn:,rJJ~~ dl .. t:,!;t;;~~~i~>d by pcrt1.'Cl 0hr K'lW~ mid by <I rlUllQbt.'if of 0 <ldHc1,{l~l.~(Oot)tacO'lI!cd iIi lhoe 1i
IIDughi!o 'Fig_ I C)" lllt.~(:
,;llJffIT~t(l fPDlI~

A n);r c~"msing~hc~' three ,Cmn:I\.':Il~n02l11 (1 I Al! 1:1:% ,()f .. TlIDfID!~Y(~T (MLU "'\
i'~ll~p~~.!~P~t RT~. we
Q!l~!~T"'Q~' 'iiI..! i~.;t

HO\\lcver. oll~he ~'fiOl~IIO;

~i;:!~m!Au elusters
QI~

w·

Sll.HIJb.OC, '!i,~"C:~ilin

fl~!·dC'il!e hQn!ogCill~YS!Y

Ihe i.I:l!'mO~ \ fig. !,

diffil;~nt AJ.I ~t~ISlII,.·T n!Oqlilltl~ogil,.'8c ~Il ~~c"'1;of th.G r- 1101 ~.~0 I smJiou:::c, rI~m!cro~~. ~]jj±~I~:r S.~llill.lll i\l!!. C~U:S~i;.'n; are d is~r~]lI!~~'Iclh~nogot;.!1ooll! sly on

,ID:;~:U;:1i; C1.f hmnoMI;_~rl~'l'!IS!Y :>lTi1JU A:ilI t~!u]O"~~li~tCr:s the J~.li01j nil

Th..;:

r mlG1, n-

di...lnl!~I,,\d,
U (II :;~Ir-

~i1~ ill1p]~ '~ha·tIllIO!lo!n!;;'rk:

AI..! !l~l!llOd~L"'~!;,\fll

J\.!lQ ~AUII )

.;;IndsmaU

dl~ STi\!l:~mll.!SCs.11:; ~MULJ!;\il;,idil~. di"mr~h~ll(-.duiloJ1g lJ:le" T~ lrotig~l~ <''It'Idm'e ~OL·me,don. U),P of Sf:' Ji ::lites 12;1,.26).

D.ci ;llnnllgi,

show

I..!r'i~1

~h~ terraees Wig- I. D ;JIid G", ~n CO:r!lms~.'On ~hc 'J.:1i01{! ]0) x~I.·ril~, w~ 1:"(~I!.m~~ riliir~ylfi~~~ Au
clilit-ilci'!i.lh l\[j~~,,,, fu~. ]'!t(f~i'tll1i~l~

1:.-

(li::col1uh.'

the

:>'li:p

of I:I~C'Mlb;:;lmh.;, and uc M1\a.l11 A.li Cl:lllil('ffi on th...: ~mlO~ are ~~~n IFig. l, E ,uud U,.

dinh'<l: !\'\~u;i.i.my <J1IRT on ·th~~1; s.,irr,1.c(:. lJ.1is !ili!di.!1g II;; ~n ;J~ti.,~Jit:i;:tn \lI'ilh rlt'" ,·iious RT·S,TM ~tlldh.!l.i.addllt'slill:g Ali!JiD1( i 'l (7, 17. 28. 19).wh~.r,e 5ii~l.Cri ul.g i:ll.cO:inil~g.Ail.ll ~lrl!.~ been
i:'Cf.!ru1!~.

In i.heprc-s.ct~twurk.

or

en

We'

Ii:m~d

Ii
OL

60

ga% tIL ~'iCDj110)1.1!I.

!:iI1:m. ML~~'~

N'" ,~

10)

..S1~C !II: FIT

!ii:
~,

·om

10J "" ~

1:
I~'

• a
~Je
(I

Itm:mogeu~ootl~'IIY d[~"~ri.ll11noo .n\i! ~I Au dllID.tcr:;: s ru::'!.''i on Ihe li:a'i<lCCJ; of dlC tr-TlO::( ~IOj, surface whcnl ~ co oooiedllhc s.l!bSlrn~e dCllwn lit) S7QC oofm'c C'XpGSWlfC (R.~ 2. A mtd i3).Acrortling ·~o !hesc It'SWl]U;, \\'1.: ~~I[cr IJ ml !hc i~~l!C'rac~iO'l~ .... ~1It. I\Vel!!l AUI C~U\..:t1.1l}< ,bud !:h~ TiO~ l t[ll.~utfutc~ i,\1 Jj~ll.irly\\r~.... {J(ll, On IHIC lXliliUr.UY. l""t)lh for th~ ~k I'" TiO:!( 110) :.;mf1!:iX' ~wi!h Ob~ v..eaneies I ,md ~hc (J~T!i.O:!!tII (I) 1ill!IFf.·U:C' (wi[h o.~I'ltorl1r''lI. [~lf(:
~'1'i·!,I!I'" 1l1'l.;."".!l!IJ;.\i:,1. iu FiJil.1 ijmpl~i 1~)!H!t. m~ ~h~1\1(': Iwo T!OJl UO) !'I\.I1!1~'e&.. Slmllg",r A~I.Til~ in:-

on!hl!&! two TiO]: surfaces (Fig. 2. em F). S~mrting p(lh~1.SIn lillcllC c~pcrhnC'Jlts were 1'- ~md
ul~t4.~~"1·~0)IiIlIm
g;!!I~~~i1iirIC

uhal:l011 uhe h-TIDJt I.~0) surfaee, G~ V~ll!mhil s.tmngc,. il'llCruCI.i.on.~!,!Ii ilh tlill: l'IDld tlte (J'- 110~ .~ {H surfnces, we ~'(pJorecli Ih~ l. .:l1.Ted0:1: hC3t~n!l Oil lire All elaaermorpho logics
[(1rdelll'OJt"OOt~liJr

~ Uo.2{.~10) !i!llrf."lces ~hal wecxpesed to 3% Ml AI.! a~ RT {fig. I. D and mld ffg. L f and D,.reSllCCli ve~,)'}. After subsequellt hc,Hililg of

!.Il!:!Silllll"'~eS to .aR"C
(,"II

All

sill~cring is dcrnrly
!lO~

idi:.'~l for AlI'/~'~ TiO<L{ 110 ~{Fig- 2(,), btll

I) (

Ibr
A

J\uiO'·TiOzl ~IOJ fFi.g_ 2E~. Tille diffi':rcll~ SI;JbiI i!k~ 111'1) A~I (:1Itl~OC'ifS agllJ~~MJ:ll!:;'rirlg Ol~ Ih~ ~i
Iiql!{ 11 0) :>udfi.i;CSi cOll:;;ideacd ,Jilt; even mu'KIliC ebvious \"h(,iIl~ook[rigwu. the cmr.~pfmd. lug Au hc:i~u. h~srol;l;uru1nS fFi~. 2, [) tUfld FJ. ;rnr It!fcrCIlOC, !{~'C tulsa ShD\V t~IC heig,~it~isIDgroiills
1\10'0

I Ii:

Au/o"llO.2:,( 110')
•

S1N1 ~gl1t (A)

'II!I

..
-.

•
11

F
Ii.lI 3% ML .~T1~111~·~ lei a'~!er iI'lWI klWC III: FIT

ooE3lnoo en the Au c'~P'OscdsiJrracl;.'S belbre he<ltill£, (ha~chcd red baa'S). Ar:I!.CfIhe hC;1[ing or lite AlIIJr-TiO::!( 110) llurface (i}J.ne L"I3f3 in Fig. 1:D'1 Ihc' sharp fc.:lmre .m. AlLl dU~l'i)j"hcl~hL.~.01" i.M 10.2.6 A (labeled ''0:'') is a~most l~u'll;.'Climcs less intense I.hml1bctlhrc·. :Bccausc the "n" Il(:ilk COiN:\floOlld,.. to At~~. tr;qJJ:h.~ ill (lilt vaeanclcs (see
'bdnw),~hi~ 1\"SIJ~l IlldiC"'dil.'l.'S d~~ ;100111. h">'rh-ihirds (yf the Allil ~m'tQt~il,~"C cO<lr~~~. whll oO:thcr Al..!n 'I;~u~o(.'I.l'i, 'Ihe "i!l~illg {l,r f\\I~!A,~, lJ~~'~ d t~lnJ'IL:: r.. irO:l;(l ~O) ~ld';,;J;'1; k"Ld~ m !he 1~:nm~.llLI(m: T (lrirug'!.C Alii" el'I[:oilcl!> \vilh n.M htigJm; lip ·t~) 1.0 A (.f'.ig. :m, J·IIl>CI}. N(ll.hill!(! iUI ..he S1.M drrl!l polliu.£VQ the oocyr~l(;C ora rip...'ning, mceheiIllml. lnstead, OHi" data .. are COIl:il.llSIellt with ce-

"

...
•
,~,

.,
(EI)

Ol~the il-<TiOzfnl()l) surfa:oo after 3% Ml All! e)i\p®~l!Il\e --5 7~( (350 ;.::350 j.'l, a nd ait All n~i!!!~'U hiMO!JtaiM!$Cilnespll.nd~iiii!D1~ AlII/h·TlO,2I(UO)strr·fi'il::es {b,r~'lJlb~tJiale t tE!~~~alltJre~o~ ~7~( ~h~och@d IF lid w'm .and RT UltUJ~I~iil!rs).ThIl·111&t0t1il~'i:!m~ en sl!:a~fi€dI ar~ ~r -1~ !ilflil~ @a;clil(-1000 All! rely dllii~ei!5 Irner hii5·t~~nll). A 'tnresl1otd or 1.2 A aoove the terra(e was !l:h~rl to, ell):'lillre that il'..'Iidus.i.velyAt! dlllstt'1'S i!ft (Of15id~f,wF(1r ~ill 111;stoqrgm~. () 5TM inlge ~'r the A~r-TiO:21trlO) S'lli'rg.c~ (350 x 3;50,i1!l) pr~i!lClrw by .3% Ml All! '~~Pos]~~11at IU J-o!llo\\ltla by' I1tatMl~ up 'w 6~C (11)) Au het9ht hislogr:aD1ili5 'o~"'tl'5P<Jr1dil1g~o .AW'r~;JfijOtlU.();)befi~r!:: '~~Ud1100red bar?} iilffildl ahe~(bl~ bars) l1eallUil!D1 ilflhe5am~~~. ln

Fiig. ,2:. (A) S1FMirmage

..

..

$TNI

~fr~ (A)

alescenee
:::;inh:'o:iltg

On the Q- 'TIOz(.~I 0} )\U1:".~8(i~. iil~ hdghl. Au hbloya.ms I Fig.:lF) slurwpoo:nmmccd ~h::ak:!<; ofi.dcn~k:ll :ill1.tcl1s~iics an. STM he~~lls bel.wec'rI 1.2 .md 3.4 A {ll:lbele!Jl "3") bOlb before

o'.f AUn dllstm 11l..~h1rniSt.n.

as ihe

i1~'\'diH~lg

{V), '~he pe.:ljj;'i!ldi~too by ":(1 ~~oHe:sPQf\;d~tQ A\\ U~~~dl if! 'Otl' WCoilfiCiili:s. ltle l!rrQ~'i'ifll~~ !11~t in\!k.c!t~ ~arg~ A.UJ..J, dJll~~l1!r5tlilat are· 0111 lrillSlUUcd slri,blring. i!11: nd IF) l~lke·(Oalfldi (D) bu,t r~r lfrIe AWQ·Tio'2:(110) Mlrra(]!. a In ~IIf).he t 'iilidicated !by "3" rll\rffi~pOI'l~Sto'Ai~ dll~lPts of O:rlE!' M!y;~\rhe-i~ltt. No sirl,t.ering ~\\[Ij!>Ci,~~fV@d.

~i!I"

and .d'lo; healing of Ihe s.o.unpk'. The clmrnoC~ !cl~.s!.ic ·"fl"'...k rowmd fo:nhc AWJJc·Ti02( l !I]I ..... 8W!rr.. OO i.!Il 1~~f,~I~icl"",,tlaut b1il;l1WC'!I" 'th;;ui ~lC m~r.Il~wr~ p('.:)~k ·~yp'ie<l~. at AUI on the r·:[iO~Ien r ~ :!oilllrfa'L'C.. 111~ ~ l1L.'Sl..!~ oem'i ~ C'x,pa:JI,i!1>tdliJrjl t!'it: D.

REPORTS
~xi~tc!l,ce of m<lI1}' stable A\!Ol cluster c!:)fII~g~

II~ 02l ~. (I), sur fm::e ~f!c r Ti I

(Fig, ~.. J

AlICXI)Qi)l!1!C

at

tmUi(]!!lfIonthe

t~

TiO'!llllH

surface, whereas

[~:r d<!!!ll p;ro1nl::l.Kl'IlS in I!his s~ rm~ m ....<J.llx! in the ' STM ml.'a."ll!li;n.:'!1~.
Ol~~.l1;
h;p,;;
t'J-'TIOl(

o~~ f"(;'w ~'iiUIIM~A~ ~'!~I~tI;ri;on~:1~g~t~Ql~'" !y .., ha Pig, 4,\~ 1;l0001IlF,,~~1l!~ 1;,'111,:'IIgcl~c."In)' !!I!Olit ~~":I1Jl ~[I I,,~i:;,t 0111 the I'" TiO~(~.10) ~~lIrf~I~_ Th~ ' 1~\O:r:.'lM~ A!.JI :-'1Q.I,;;:h~ ~i11,JI]d un D:F'f rnlC;:~III,Iclear d~U~Il.clcrilt~ of the .Au hci gin cl isi:rib~i~ion ti:ol1(s '!.\Iii.l.h the bliE,!h.,-r!.::;oll,J'cima STM iU113gC:i on r- "fiD'2('~ !0 I ..$ 0pPQ5~d 10 the i~wa!~m'lce mosl.undant SH'!il!.1! AWl ...elusiers, Or! r":Ti0lt! I(I~ Ihi': di~"lrili:l;l!.!~ion (j-n(l~I,~ I!.BIlKm h¢~II.in~ ~h~j;;Iir!~lIk."! on 10) Au·~hlrt;'d protrusii:l1l-" (fig. .:jDI1Iri~ ~C! bf.l"C i'(:v~~l~ !Im~ lht; Au.jj dll~l~'I'~ bilild nlm(: li-elli! A!ill in 0bo vlie.m,t~C's, ll!t;."C~L1,IOC 'rb¢ d(:lh ... i~}' u:f ~1.!l(iib1!5ly to ,tI1~ O-i'tch (i~id:C ~Li;P~XIl1. th.:1'lC' ::"fit!ll~k"iSt ~lIrornl;;iotij<' iancrefLls¢,';; l!JI(lon .Au c_\,[.. To rmio.n,:'iJizclite ob~aillctl A.i.I i::l'tL~ler dispoa~a~ 31. lh~ t:'J/i;llIC'l'I."~or dlC [t'!:itr-"i!y of d~e faiitt tribLJI~O~'i!i &Iier Alii 1i:\Ii:~urem RT. we CtiiW~ pro1[Usim~:" i1lri~i~1~ IrChln OI:J,\~lCilndc:" (/7, 1,1,:1. p'~I]:ednhc ,cl,i n:1L~io:n rAu~ ol1lhe th n::c d~ .i:1i'i-:ct1! o TIle second ,,,11IliilUeM l)Oolnlsim'l~ Founcll (.FiG:. 4. T~Oll ~, O) $il11"r."LO:S il~lcre.M by me,lnS (if DFT I E am!1 F') (u".ilgiil<itc ~roi1m ~incal' A.ulrim.c'~

~,IO)I!j~rfu~,

we

{~~,,rl;-.d

;I

or

or

(.Ii f1~'OCJI[diSl.ri.'b1)lI iOll 0 f pt'i:h~ll!IIsiol1,,,of A iii"

~ltiN-

(F.ig, 4\ J la' L).•\"'ilt~re :;;()lIl1c ofih~ ~]1Nl~ksi Au ~oflogrnphie~ 'I~i:: ,dcpic!II.:"d. C""l~IJn,l Oll!ll & 0!I11ll11c'i ~c ar,;; UI;:k':.)lirm ~il~~ for i~C'flIl1!i.!'I_g Alu. n Wig. 3D). IllI: Og, :at{)nil" art f'lmOlidlty .~iti~1rpol{u[!.5CI, intO 'n1ic AHjj e~l~(~fil. dli(~'ta.-O:r1~ m:.....i~jjl,1)!;lil of A
~~'k'.."I!"ilU~OO proh~i~lRo [0 n'll.1f1illl\iJ,i"j.J~L elillskli':;i

i...

calC:!laI<lli,on~ I u some df);;t.1ncc' of tile lX!'im defects, ;1~1Iht'li.'C:-lIrlaCl.~ c.m bie- cr:m,,,id'cro:! as l\I~j~[g Sl()~ch~~111i1.'tL"k; 11~:"T~O~(lIOn lhil. C<L~" ]n !lll'l difliu~(m 'Llam~J1'Ol!'C f(llJ~cI to .~ \'cr), low (iti .• AWl 1 lfL.'I.Idi cl il~iii:«'~ ;U I)' JDin!;i;siL11~ I)f AUI on :~ 1iO:i1(! Ul)i!1, ~'Kl~~iblC'lIot onl}' in Ihe ~~~,~ I di~clion blll <!i~Oi.!l ~lC' q Tn) di!1l.'C~iQIII (fig. .. 3/1.1. '~~,i(l!llI~ DFf C,d!/;llIl!(Hlio!k~ ~ikc\liIi:ii(l i~lldi:~lhJ fiL'C'ill}diifflll~~(Ql or :;!llll(lill Atl, 1IWI'Iic'~'S lUI "'IOlChhlI1tL'1l:tIie fi:»kl,.: SiurfaC\$ Il20, J2-34l .lPor Hit: !P-TLO;ll ~.I serfaec, we IX'~CJk-d (!J) iticunieai c(mellil~im!s. bccmJs~ Ihe dil:Tt~iClI~barrkrs are a!nUOS1 ~illChal1igoo (Fig. 3B) .. 1~IO\\~"\:e~, flibC jll.,U, monomers C1l1'1 be 11i!I~~rcd m l)Oj!'lI, d:ctbcb: [Ill (~lC IlCirrd~es Ihc r- and 0- TLO;!,( IIO}

un

(i,r

~~ri1:trivisl, butthe Ji~l.inci .~IC'~,1~I. dislrihul.iOJ1 of Ih~ All ChJ6tC'J:}:on, the ,tJb JiO~:( II (I) surtaee m!I lows
!Ib

w di :;tiJ'I~.llli~D~IW!.'Cn

A~N c:I:u."le~ cO:I'lSiSli~.~

OJ

m Au

~A 1IJ t TI1 ls a:lt,.'<l~am'lCl1~ relics

0111the

folkii\"":~n,~:

or~lI1c. t\\O • or C"V~'Jl il~C Au t

Rn

tlimcfl\ (/\ l~2) a:I,C ealeu ~atc;,l(]~('1 adhere less ~.!m:li!gl)llo rhe \~~Ilpml I.h;~n A~I~ e~U~iCfl;t . (T uhll.'l l I'. and W); DfT l,;'lku! u~kl!!." l\!iIU Au;:! CI;lIl r.gl!lla;niI;:iJI,~. ci I.~!~·,· sy.m!lrlwlti~';llIy lFh:. 4]8., O:rO!isYll1un!:tlfilCaUji ,~.F~!ll. 4C)m[m:.l!cd jo Ihe ()I;~ vue am:::y, 'I~fiI.;'C ~',dyrclulldlJoc~h~ l\\IU mo~, ..bUJ1"

or

byers (nS, S2t more ,oolnpl~,,'ilC S\If'\ Ie)' f(l( ,\l~blc,\!U" dlt,;l'toL~ '~"" h I :s Jj :s 5 ould ,~ = r' on h ~hc tliUil":'~ TiO~( 110I ~~rrm.c& or iillt~re_~l mtc· ~ll~~ TIw DFf re~!J~l~of
~lari~'d i,,!'f,Ii}I~
I. }\.. at;:bt.·11C11.~. \~ :.Hli'ii[l' een-

:>iOOl'lld .~ ]10.:1010). "~~ll1l~ghlh~ .~l!rfhC'C ",,'as

m'l1~1~td~oo c."p:~;jml;'f1!li<!ny+ ThcmOlVt ,~!all!ceeu-

,~uri;tw.;,1(1~~nsidc or Fig. 3, C and f), l'ClllllC(;"li\'cb'), hl ~11L~ cases, the 1:~lrJf1i~·rr. hvg'h~ ~re
C'j.l(mgh to p.uc'Vcntd:~IT~sio:Jlevents at RT. These

or

calculauons

an: ru~ly in line Wll11

OIilU'

STh1: ob--

si!'.!,'\',~tim!!5 ofh(}mQgclJl;>'OllSJ~1 ~li"uibl!\td ~lmJ ~All ~1,lnocTlNt~'1l' e;~!C'!lI:;;i\,\dy (m!h~ I'" ,u1(1 ~O~ 1!0) r,J.o '~ILllft1u;;i.:_~ rind sinlC'I'inl;l of Au c:l1lI~tCrl\, on 1]l.L;'

at!ached on an 0lii- 'vacancy (I), ilrndll1ool,i3iltac:h:e!rl1 A.lJl~ (0. ~D m F) Z(l~nl-ii III S'I'M iml~g,!l'S' {40 ><<110 A.'!) shewing Aul,q. duskers th<l'l ,areasumea 'm the (iii h::u'[.a,ted A~ dlcister 'L')I!'lj'di~ur,allfQili$, C(lrn~s.por'!dililgl UII1f! p:rOI1'~tDI Ules.e rm~l" S1J~m~n~ are $ho\min 'fi~. sm. (SCllt().m) M~st 'rgv·~raI;)Le:s!ru(hl!ffl~r'OU!1l11il ~n on
syrmmetriC:i1IUy

IFig. 4. {To;p} Mil:Slt. f.~l;'or.able! structure'S rO'L!lndiWi DR (.<I ~(ju!atijonls 11m AUjJ. trClIP\lled ]11 an Ol:w v(!;canq W. PtU3; s,rnrn@tri!!:i.!Uy

~Uon {)l-1riQ,,(liLOO; _~ m U Zoo m-in SuM l!rIII!1!ge:s, (40 x 4,0 Ill) Qr Au.n du~t.eJsi1!t.·
ta<hedifll
the 'II11dnity or O.;malclrlls that a~ t:el1ta'ti~y .a~~'glil'~ !.Q1~'hf? ~kula~ed Allin dli~t.ef' confi~liral~oifis.

cakiu!atiollc's1or

AWl

{Q,. ~

~Hlhand

AJU7

s: ..(D1;5
!. -~,O
w

I(UJlt B: h-TIo'lI

mfOO1] .:Ei .. ~.O;g... II:

i~r.!5
0
1

.
~'............lo'
I~ IC. ;':,
f.

'[110'1
, .~

8?

!I ..(11,5

".0,.'~

·,~.,.T10.·.',·~ [0011,] t<

11 '.

_i ,-~,0

..,

'"

i ,-,~,o~

lEi!: 0-"0,2 .

.',

~
D;Ff.b\il~dI APfs.]11 'fi:V(per Au" cruste-r) tdlhe most s~blt Alii."d\Jisblrs.llrithli. S n:::s 50~nd fi'~: 7 s-, and Q-TiO.2'~lUO ~lIriaGe1i, re~tNely. SOlil1le,,,r Ih~e liulimbers.me~I.otl!i!clinf~g. 5E ~see'also '~~" 53 to 5.7).
Om! r-, Ii-,

_ ~1.5 __ _ _' , . ~DIl11r--..,\2.0 -

T,ilbl~el.

<11.

e;

a:

ReaoJ:1on P,alli1l~A)

to

Goldl
,l!l1U:1l
AIUii1:

dU:U!ei~

f-'iI'~.O~(llO)

.h-'TiiiO~titl0)

.s-rno;!tll0) ---'0.61. ~.10 -].36~"97

o!-TiOz(1'lLO) -2.3[1
4,941
-.3,(1';1

Fr'iig. 3. Potential ene rgy p:JiOfj.~~~ for Aui i daJ'I1ler diHtlsi~iI1I(11'11 s& (AI, h· ~B). r (C),Cll1!d o·~D}no,2(:1l1O). The liIlri~irrHI at. whidl AiU!l ~5tJap~ed at 'lJhe ,polrr1t de'fe'~_s (Iii! t- iilnd 0-'I'i02(nO} alr~ fndkated {a:rrO~Vii!(, ml:l\,!!11·~ ~·~~~!!:V),· ~S-i!~dl o·liO:!(UO,), o A!lI~ di~h.ll:!i~,llonl!J '~heObr Irows, wherelll~,el1 Ii· ami i r-Tij02!(UO),AIII~. dinaJ~~ alolil[llth~fi; uoug!ls.

-:U.4 --{),it

A!\l.~
~1!J11

AiU5 AIiJ~,

-:l41 ~"57 -{1.3'1

--{l,'52.

-0..5'1 -11105 -:U6,

-1).,64

-4,041

-1,(!4

-~.~:;:!

-3,1$,7
-3.:f1~

-(I.S'

~:Il.:ta:

1694

REPORTS
~i.gurn~i('lllSon :~1iO~ 1~'0) ;]!1CI\,V~ ~lnCt!t'*]Il;']1 (lD) "t!1.~~!'\e~ I~":ilt ,~. ~.t~'l.I;~"lW via !I~ f\.. :::!t\ll~:;;, <ti11~~ 1.!1!JI~lLI'~ ;I!.~iWl\l~ ~:I~ II(XU J ,d~Iil.":l;~ion, ill 10
§'!~~ i!1~gh~ i!lit~\\,'IW Ih~ m(lN

Al!i.! ,~!.!~~. billd

As, fl,,1K)~ pn:v:iIij!JI~,~,. ah~:;;~ s1!~IH;.tLm,.,;>m;!h!l,'I'iLl only wC1lkly 'lJJ), Oni,hc Il-TIOz:tll 0) :;;nrlit!.X;W~ ~bund even snmllcr m;:TI~I;;g;iolmrolenli.d ~~~ayit\". I(APE~) tor thf,; ~]to~! l"l~lhlc oonngur.'li(Ul~ r.!Il 1;0JlI1(,1..,t OIl [~i~, ..:ri0,l( 110, Slil~tlilcc.AUI 1~nd A\I~ cilWl!!C \'I'~ oh.Wf\'cd Ih~~,d'l,l!>li¢1' \~'il1:i 111e SThl. bind tdilti~ ~1ro:i:1.gl'yin '000 ..'!)tm:leD~'i>"\lI'h~ii:~I~ (Fi.g. 4E". Arml,y"'ling Ihcbt'llirliL1.!:'l qliillJuliitil vdy ~I.~ AUl,A.i14 (J\liltel:r.1l:lIli:~),. AM::; lA~1 re.nmil.~ers.}. ill l~rec tl1i11iIIt:ibi.{!il~ (Fig,. 5AJ..wc li'liEnd. lh!lt ~:I~,(;W~l ~lIrg(! 8OCUi'i'll~~.II.efi d (n'tl joo~ i'lIJees} and i\u, j Ali SCtl~Lt~1C'1S)~jjld \'c~ w~I:y. r-'on.ill.: "',1.11C1.!( I Bl) ~I:lrfacc. ·W!! (oMnd ~..~r.i1M1inlli!iII'y bCI:W'b'!CJll the Au a!iO:lUS andthe :iiillbslll:a[C S. r:, 11 u~N'lnmJsmplHl.IiI. imlk<niili! (he fQnu.'l~i()ll r eeo 5lron£ 'oondillJ:: of all Al~r dMS1i.':i:'S con.:.i d'ciioo, i\ccnn:.HI,; 10 the DFT liesill[I~$<l(1dre~in~ .r- "ak.'11~bonds. A mmc qmmtimJir\'c irul;\'Llysi~. ",vh~rc' and TiOl~ lin), d1C CIlC'I:g)' Jb~' Au" d~)t"I~r dl:!- li.rc {i:nly cOllsidcrod the Chill'l'gC :recll~ubulion rCI'I'lend~cular ro [~\'C SIl!II~X'I!'!, sho\\ s ti~.u dlC Au ~<!.dUTIcn! rmm [he dclcebl onthe '''l'jjO~ II OJ 5~rmc~ docu\\O~ :l'lIpidly w~th dlll~'itiL;'rsi~ hu~ ilHl-11"," Illf'J~l1<Jriill' ~1;(~riL;'~~ce dlllL~~ Ill)~,~:ri~..:u.ion ~IOl~m ~11; ,I- ,[[,~O::it 10) ~W1 I rl~~L.'. where th~dI;I.InXlnOO~rpl~O~1lFi!l" 5B~I, ()n t111~ fJ..110il W~ Slll:!~l'I;(,. t/!.'Cstl!d:tcd Ihe ~m::ll1nCnl CI1CIll,Yrumal~.s h~~. This mcarts lilm. IOf ,! gvwn Au dij~I)f.~i(JH. a hi,~lI;,>fp.:r,.li~!.nClc cxtr.mrdimll:i~y ~limt;Jk: AIiJ,j dUS1.Cl' in nil!/;! ~,'jci!!jl)" of .m OfjtOli(o:rn. III this. e.."'~. clllOlJ~~ !'li.1d,!;sinlTl,u!ion i!'i O'!!lUiR"~, ~~ncrin!:!: 'IJp¢fH h~li~!ll i,.",lik(;i)' ~~ c'Jj"ili~ S!;l!iIl~~.t!llg!lIID All! 0 brmd ax~ Wig. set indio C!l~ rhc lJ.TiO~lIIO~'!l;ULfiti')i; ~z.; cOlfl]l;.m'd TC!' ~liJ :;itCfI!nfilg lIi~J!. Ihl; l\.u-O b(u~d~ c(!lltmlbulJ.. In ihe ~ iLlmnol;1 (11~ IhL: ,'- JiO~ I'~ $1Jrillre Thas, ih~ DIT 0) !I1J1.\')~liJn(;i(li of til::' j\lll C'l1.lli[C'f. The q~:!.L:iltiiillhc ~\~ts. pJ:'L~lltoo i~Tsble I e,OJ'lclJar.,lOC: I.he: SfM firdin£,S ~(W, n inl fig, 2, C 1.0 E 3ll!lll¥sis (lFi!!i. .5D) n!\'c,ll~ !'lam clecll1'O~1 dm~ h3S
I\'j-

!'n{l~gly on the o":'fi~llI0~ &\lrf~1X ~l~;~~~~~ r·ll('~t~. -~w.::ra~\. \Ii'l? ~!1~~~~1 ~ ~Ol~ d{l"'IfC!il ~h~ITlWdf;Il~L'Y dilri;;n,..w~ ni1ll~~ Inr two s.t:Jb.=.k'tl Al!Jnd1,J:>iJ.!r.:; (Pig. 5, A m~d C], For ~lllc f-nO~( 110) ~urrilce.. \\'0 chose ,111i\~l cluster K1~·1'i!'fiIL;'triC'ln" .[!~a(:hoo in ,Ill {\r, ~',;;U:;<I!l(:)'. be·

m~lc'h

in:Q":cd been trnn:J.l~clT-ed!(!I~hc surlaec OO!iWnl'~' A~IOI, 'C~l!;;;~"''f hOl'l become C<1a~'rl!lic. Thi~~ mc;ln~ ~hm th~ bm1(I.~I]g 0 r l\u~,\1ir~ lhc 0-TiG 2~ 1m) :;,url~IC;:1;.i~~)!Irli.ll!!Y 1:0!] ic, C(nliriblllim1J elf imlir b!l.1ldin;g illih~ OlSL~or AI.!~ en a-Ti02i~1WI ~~cO:!1Sisl:~nl ~d~b Ih~ I;;;JI~. OI!I,~t~ .. tomk: J\\I 5<1 jloplllhn:ion,\i Upt1'1l 1!cl~01P" liOl~or AI,I~ :and All~ 10'1'1Ihe r- :md lI'J·lfiDl(! un
'!.""~1;CNby the
~rf1LC('i>.

R""'I)ot1i\oC.~y.

A:;i

com.~irccl It!,

Alii!

,aliid

ALl" i~l 1i1.1~ ru. p.hru;c. the roktl 5dpoPli~~I.irnl g larger reduclion

ii'i

n::duc-ed by 0.24 and (1.55, r:,- for ALiJ!r~TiD2.( i llIJi sud A.lIl~I.c,,·1iOlti UN. re::;plXtiv~ly 136). The

of the .5d pOlH11.nioll~ [or l"Onaribute5 022 or

Au<!Ia-T~o.zt 1.10) C<l.~~be traced b!llck Im:inly ·~o Ihe .~inslle aWnllllll~ l\lindtl,d il'OCll.:.' to the O~ t\u
t!l()!lft~alll"'c

I hi:.>:AI!.!

m(lm

~h~1 ~-. 0,56

The ,~WTC'~(l1tL btlm;l :"tt'thgni~l:l!i;.¢If Inu C(ln~ ~i,dc!1l."dAu" clusters ( Fig. 5. A 10 D)I rclkcl tlWl ~hc re,cllltoci I'~ <met i'·TiO~ i l,n}I'mih~ <Ire

filg. S. 00 (~aI!1:e' dertsflty diffe:r-' ~f ill Al!!3 du~re:tr Qlit~~d to an ~ lo\iK:i:loc.y. Y,eilO'ill ~hE,'l'e:s
lll'rJOO ffiil,p

a!'ld diliShed~iFd6mark ~:si'OOli'lli (J!f All! awms.,. Bt!J)~ (jFl~ rndIsWJr.~JI:e;s indicall-e d~pleliDn .and addlilio n, ~Pllc~~~~ ~r)0£)15·(2,':'" peJ jJ,3 .• (0 Chilr!]ii! d'e:nsity dtiHere~u:e milp of
Au., attacliie~ 001 o·nO,~(:1l1Q). ~ I
S\lrfaC$ ani!

or

cl~"Ctm.11:;; :pct elK VflCWICY rund OH~ !i!J.>l\o cEcs.,resPLIC'~.i'IC'~y L \l.lhc.nca~ 1.1lL':f}.[i02l II 0 I silIirllmc ~s electron d(!fiCJCUll (R'!rililli2ll~y mis.."iiIl.!1\ 1\\10 ~k'IGIIDJ'IEper O(li! mom ~ As 11.l:itmcticn of ~hJ! sMr~illcc ox id,~tiQ;jl S.tfllC. the ALII,,;-·n0'2(1 10 I adhesion .is .~loml1ion(}Imil!: for II -I. .• , 2, and 3 Witll I~f! Jo.WCl"t3rlll~iQIl ~'rel1g~'l" on the h- and ,,-,.TiOi·1 ~.), N!.Jr1u.cc,..;: W~. 5E),. ~.:;;or !h~ srn:JU n All", (:Tu!\t\.'f1':, !h~ w;ll1~i(m ,<:Ilxngth incn',I,\\:!~,011 !''-TIO~ll O~" bcc.'1IJ.1SC tbe oo\'3'~enl bonds be... 1.\'I/CCIl TI i;md At!! rOlTIl. I1l0!'C readily Whl.'ill 110t ~n: the T] "tenus, a4!C ill !lw~r R~H:joxidized :5W1C. .
Ct:~!> !~iW~\'Cr. mll :ldlt{."SiOll!nrnOC!'I[llllism

(.111J(!m:mriel] (fornmn!y

:tl!il\!ing two

<l!mj IrJlJtI; C'x-

itl]\'nJiv~lI1g

c;:J~ l(mic

Au ,~(::~rs when UtC~lI r:1'1I.I;(;iSi o'i\idirod ~(.l. TiO,d ~10 11[,whit]! l!%'lds It] 11 umeh ~1ID1lif,'i:r.

dE!I~~dl s e

il1l

(Al ~B

AUjI"!>lIW!'pG:11.

ooJidklgilum
(lW

1!hfLl n Ihe: r~,TI();!llllll u

,alnd 1!iI) 1E~(lrofil ~h[lrg~ deniliity

dif'f,~nce ,~p) fbrlhe5i!!iflIl~~~em'5' as 1m! W ·~md (0, rts~ivdl~; bl.!t. ilntlil.g r. ate~ OV1HUhe iiOllnd r d~iIil!~(ii'iS< oflh~ R!!,ilIilir·,ooIL1lUlez pDsilicl1IS oii 0 ilndAu a.ooms .11 Fe '£I~en 'Mlh Irespl1ll:1to 1lhE!! f-Ii arom:5 S (1= Ql. iMler@bV~oo ~.aool H2 x All" i~ ~ lQindic1IIle the z pn.51liDrn:!ii (If ·~Ile·I:M' IW a~oms,~ililIMJol'e il:lentk:.!l~. z' Wlhlesinh two AUJI d,ill*,~' IQon!iioered. ~E) I'krt. of l~ f.alOll'laled AP~ror !h~ moo~~tabl~5i!I,aU A'U\:r dLRSI:efS (n = 1.001 4i)as.arime:ltio.!'i of
the ''liO~;(l,lO) ,~X!jdi3itilm 'S<tate" (Oii!ill)(!~ with Tgb~ 1).

[I]

g
.3.
I'SII

JI;.

~.")

i' '"
I I

[1~epre50medcxlA--riJltl,;n!111 illld ~'lllis •. we OOUhDtlila. 0 voc."Ulcics on oxide SIJIrlaccs tarc relevantm :iiI~ fO:l"nhe st3bili~1ion to f ,d rspcr.ll.xi At:!. ~10LTttcb'1II1'ldel' :re<!il 1t".:u,;Ii:ml cond:i:tiQ~1s Wil.~1 u.,>\llilllly high oxygen pr:c:ss.nres" [nos.earl, our fC'SlJliS SlJl£gCSl. 111[1'1, Cl,/,etl, under :81~ghll& Ol'I,k!l.dng. oond:i~~{)ns as u~d IlX::I'I:. lite Aill Il<JJllictcs ;.1ll;1.:: $t3bm~-d vim, Au 00 ~II 'oom!.~,~whcl'C M l'Crrc3.:n~ a mC!iill ,aI{llU l I,he ~~J~PO~1h!g (iI!'!\l:dc), fhi[ii i:l~l~'fll~~!.ionis in h~

surf.."lCC.. In Iliglll oomptllnlimml

or

IE, 0.0

lS~U) = .!. ."

\"'i!11 ~n~ n;r~m:~ (l!l)c.~~i~llic:Alii K1'l!;.'J;;:I1,'l'i Ibr 1\1.1, ~LPilfJlr1t;"d {m ~ll'lgB1t1>L~. In. ~1.~ri'LU2, ./5)"md iru.ra QN:i'UI::(14}. In! ~III (~r tlt1.:: l,mJdel s1udik.-:> Il(&lrci>~ill!j Au :SUppOrted on !iWfllll whcrc~l.~ Cl'Ltmlynicamh'iI), of the :s:",s1cms consiodc.it:'d was :ilMYWfl. very high o::.:.ygcn 'c'xfl'IJ;."llIIX:S had l'lo!:!cll Ii.l:i!.cdto prepare or. 10 ·llCi~'9~I.:" 111l~ ea)ta~ys.b;, (7. .0;, n,:, 23), ..
I~

n~ ~

IW

ID.
0(

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-:3J:I
~"O

'

, ..lIDO!

~1i02

'"ifi02

f{lfll!lI:m

o~.diz:@d

11)-11°2

<It11ad(l~timml IDlt\(:m whyw~ qll:cstkl'n~ pre\liot1l.sIYllI'Opruc-d :olructU!"C' m(!!dcL~, illclmlillg oor O~'i'FI (/7. J5}, \vherc ~ ~V<lS impJ r:cd Ihal ill, 0 \'acm~cies allm:hcd All ch.J;... tcl.':ii comsl;;!!~hC' CO,l,t,dy.,ic m:-~ivity .in "fe-a.!'" CJ~,!I~SlS (i,co,. O,l~ hi;g~ p~"S~\lre ~. "V~ l~ml'lOOe!h.r1 }\u, Pl'1!1!ic!~5di~pt;;'~'d flU !'~-d~cibk n;>iidi.......;:a.!!:t;' beH1,O't OI:tallyslli>;i Umlll I.h~ t';1,I!pport.crl 0.11~lor!!mc1ucibk: ~nt~~Ciu.:alh....c litth:i,eibl~oxI:d~~ •. 3ltCrnllOlfC c;irmhkz
l,bs..cfYiliio:i1 :i is

REPORTS
of r(l!111~~ngO"-ooh At~
1>1!11fl'011

in~jb"~c.s.. N()t~

~h~~O':viQ)~ ~yrt<!,l;'l'!I~I1fnim!i!iQn, s have !J;;;I~nidcn~iiiQr;;j !tOt OIl!,}' f'r:tr li![Il~!ia (J4. 37) h!,li ~d:<o f<l' ~"I,\~ml oi!u,;r t\1n'udblc Q1(id~"." illd~diI1.g \'1II~~
d ilillm tmdjrurl 01;: ide j38). i\ mll;ch ~~rong.erAll 0..... ide !:1uppml,
C'Xi~~~~l

;ilh~"$im~

::Ii.2'. Q.!F1Il. Ifl. SaVtibllr~, ~, ~'l'ffiIl!li-~If{p~a!1J~lIla!l, SGieme! lOll, ~;I:) (il;OQ;I). :i8; IL flJ, !!r' ~i:,J. p~, ,O'!il'iM. iii 100., 3,']00 ~:2rOOsJ'. :Il~. G. ]L !ftlolk;I'IUlg~ I!t C~fal.242., 71 (2006). Jl.S. J(, iI'.iIi.!~J!Ila~" S, Cij;f,r~ltin. A, 'Jjm1~J.Arrr. O'i!Ml', S~. :12'7. 12Jl6 ILOO!i}. ]6. ]1. G. W~~!J',II. "'ilmll1iJ, 1M¥>;;/fl'. ~erl~97" 13:~,lOf

lIl. mile Au,,1!,Il;es QfI~l!ie vali!U

~~~vlla~edl ~qgrdilflg!!l
~j:jj ~;~

e.:

fJ>.. rich Au :'(!J)11I\"111intG!ii,~\;'s Ih!b~ m~ O,-:pnoroxid¢ l'ill,p~:mt~ s!'urfu~_ For (:tlla~ytil;:11.r-pJiCiUOCllill. 'uhi~ oc-i>\uh ~!!\l:,"!$L~, .he OCClUtlI:!i:liC~'

~aIiMi~
Ciil~

I~lllure. roc I.h.i! tw TiOz( 11101' ~_rraCc. W~ RfL.ii'ild !Ilm q.,\mro~~cics allC'M1 rarllt..;: .st<lbili23li0l11 or gold 1'OOl1u~l1Ci'S and £oM lIill1!ai~. htn 'lal'Bt-'rt:!l,oki (1l1~1C\I'S, anllm be' )<tq f1kicl1tly :in.'!bm~. ''A~ ,cmc ~ll<lSl~ 111.11 a:I'p1'O<lch of .'ilUd)I~llg ,(ll.!r iheexidc ,~W!ppo!il In a l:o:!Ilgl!! of Gi,.. ~d:_ni{:J1I1 K!,m~ i:;. <!ll~pJi~ c..ble ~1 :nlllm~\at)lU$ o!h~!" nlodd ,'\Yl'1!t.~n;'l.
R.@f~ilmrn(;~s ,IIM:I NiIJ!~S 1. M, :fI!irll~, N, lfal'l'l!ld~,. 1. K\:ib~\filIilI~ s, 'Jlma.}:, Clitlli:

!i;'{lkl

m tile

AlLi-;;ilir~1

t'II

~i1IlcriiiC~ <l'i

or

g'Cij.

.t/Ili, ji~ l/.i!<Ii:. mt. 9Q, 00'6101 12m3}, 16. M, ~ (ibm, D. \!I,GiJ;ooomn •. Si::ifm'f 2.5,~(;roM), 1l~'. I!. 1>1. ~Ifliedr~k;;, HI lo~II:" J. tt ~""~C~ ~ Cilem\. lilt, .f,'rj, 4·11, 182,4 120(5). 20. I!!, 5.. wQr&. Ul !i!~1~, F. C1!!q,Yi!!l~ G. f&hii;.!!li, J, f'hy.i.. (hfm. 1iI'109, UAU Q;(I!!I5I, ~lli. 1[, W\ll\1!iI~1

~1O(6).

la"..

&.c {Au"J\ 1100 £1)01 h~plH) ,illt 'Il!ie 'tallli (!nllfl)i!l!~ allhe ~o~i!ile-;;iI .t!l~ ~ .iI·~~1li:am'lil~hil:le A:I.I.. ~,I) ditn1l'f~. ,Mill th~ Tif);I(1:L1lJ smfa;c~ in ., ~.I!l1t.'lil!l 'D:d":olI.ilIHiIli i1l<!1~. re~lw~!I'. ~elJalWt A~~ im!illl:al.~n~~ ~h1~i(lm 01 !ihI~ JiI:!" ~!!~,~r~, Rl~ ,~~~, ~ US,. H. A. iJelViUQ\ G. paa:hmi, r. [)i!lbt!q. 1'. :S.~IJ1;{!I.
/. ,~ C~lm. 8 1'09, 91011(1~:2005~. It !d~itr; 5, I~ Nt. 1.'1. EII"Ownmf3', M. M. ltiIiib. tL'l'J~ ,l[ei.(. ,15) 08·i410"~f!J' (2005,}', 3<1, H, !~~if;S, Nt, D. Bte~~~fJ', M. M. ~i~. ~,ft i!iWL', ,~ ".l, Ol~9 Oii!U;)< (~O(6)_ 35. L. M. MOIil'lla. M" I). Il:.!l'lll'lllmn. 5. Hiamtll1ilr. J. Cilem, 1'~. 1210, 7673 ~2004l. ~6_ Oi.!Ing~~ ifilllile Au ,6~ <lI'IIlI,6p IpcitlIlIOM~OM~I!:<i Kernr ~iIQr! mlJli! bi!' ,to !he' QMf 'I:Ir 0.,2 $ e- ~~~urmll~!ija"l per Au cfuiler f~ j!wjr-lIiD~tl;lO) il'1 On;!e1I(l ''l!!!lkrl!!l! wilh H~" 41:1, ~fll,e gmt:;' th;;lr,~~ Pl'Iaril~tiom ·0Dd !il;l' nei diOl rglnm i;;. 1Iwiwn, iIihI~ dUiliSjl!;~~i'l1Iti! ,rw ~~ Mc6;p jlO,p~llM~ IC.'!m~ot bt 'IJ!Iilnliie!l beGllAle t~, ClibllilJJ; c¥elt!ip ~brull"~i3ll)o ~ililti other Q"blI;tlIc~if'l!l"~ h"!ltillill~l!N"Ii'IQ ~IIM'IS.. .~,. V.). lI!ed! I!lt at "'~~, t.m, .,~,Qi3610~ ~2Q(4). 313. M. #ob~ !'!O!ij",~ D~.,'SrJ.rf; 15!00, 1~9j!' '!;!OO6l, and H.

Trot~ltOI ~u!i!IKI!1l'5 :were APE= flili {!!Ill\, + ~pponj - .illj:jj f~uppori;'l, or.b!!lIl~fj,,1 V!mj, T ~lJpptiIi'~,

_eID~

n. Ogm.

2.11. IF. BI'I«IJUl. f2000<t

A.

Ch.i::.-IIlII),,}, Phjl1.,

(hl!m..

Ii 100, S~U

22'" lB. Si!:hurn:lc~~r. 11, Plza~ M,_ Kinn~. ~. ]_ 8elirn,

115, ~oatLU9t C. Bo.llII.. I), "Ii. TharnpSi.'iD, 6!Ild ,&11, ::Il~l ~200m" 3:, f. CQ~m~)'. t E. Mace'j'". SUfi. Rfv. ~f~t. l. 73, (2:00U. .k R. ~e<f, C. l£milfe. S~llt Sbaillllnlll!diI1C1't, H,.·~,. Freund,. ,~d I'liill.l.. 3:7, 12. ~:200.U. '~,u OieOOldi. SUf/. Sa ~ 4a" $:1 {;roo.~, 6, G, R. ~mW'l!iW, S. \fw.1ld~ 1, N~lk&ll'l~r.I. . H!II'~, M ftltm, Li,rr: ,~4.ell '[1997~. 'I, M. Wld~!l,X. ~~i, h!~" Q, ijl,!I1I, 1;1, W. ,G;).g.drTlil,rt, It;:, Sden(F ~Sl. 1M f i[1J;9900" as Lt~,c, f<lm.. T. WU,S;" t, Ant:Imomd', An:!. C1lll'lIJ, 50 e, l,t;,

Cll'lfli:~511.a9. lG9 (0Il00!~" 2]~ T. S. ~, 1. s S1ie", C. T. ReM5, III], M~eIf. C B. ~li!1S, J..1m_ ~. S~'U~, ;;:00.8WOlI, ( 24. S. WHWlt II!I al. •. 'Jjj[f. Sd, $911,. ,lM (2001l), 25. Mille-liiils ,ilmd rnll1illl~~5 ,iI~ cl'illiliibl!! in ~l\P,pll1ni~~ mal!etii~~on Sd,r.nc~, O~liflil!. 26. O. Bik.(lOOoaIN 1l'1,.,N,,,r, .tM/,e1. 5,189 (2006), 2.7. W. $. (P:'!i~g. C. iii, f. r=!\ M, A. ~!i!de!ili~ UI, o;e~ Sri/. sa; o1I12-4l!l, 3H ~19'nJt2®. WliifJII M tI!lDl1fili!i'i!<I ItIt ~ ~R!in·IIfi!~~ ,oompli!l1!Ml.!j,~ dirl~ r..el 'willi llf,iMol.l5 dil!1.i! fremo~r g!"Qli!~' (J7i'.i~ li;et:lJfI';~ ~pjlIIirl!!tit t~M,~ (~ll Ali ;;,\U EiI'¥iOf.!li!!lll 011'0
ll(}ill.Ol Rlff,am of 'I'arious, dl'ilf~(l()er.Fa" ~Il~ l'!Jw.-

i~.ri!i~~ ~~eri~er!.tsifl ~Ul"lh!<: ~i<iri'Il!l !'K'1!l!1 ~ ~ tI~n f·Ti:OJ'(l~l ~~!ilat(!c orth~ Au '~~!1J~ ~ at ~igl1~~ l!!.r~I~n::!' m!:!~l!:i'af. 'un~Iit!fiHo.~aIJy/t·1iCMlll!J) rurlillC~, WEirE' !mlll,T~Iii!1!l1 t~iiI, iFiIQ',~cmllllil" 'nnM!'ilI, lull. "~IiUI'ttnl MIImfI lliIe S'liM 'l!iIlU5 1111 7I ,aM !hi!! U
IM,eseai

rI!!~m~ 'i~~iro. 39. w~ :l!CtnMtd'~·linillii'lml Siipt!i.'l(l [i\OliII Ih~ I><lmiltl MiIll~U)' of 5den(f., 'l"etm~e!k!gil'. ill!~ Im~Q~i'l!n th~IJ'1lb iMmO., lhe' HlIi'fill'!bIk!~j'(.n COiii'i!fi~, ~i'id ,lIt~ iJ~rnl~~1 Cenl~ fo; $d~l'1fi~~ (ompWlt!j, D..M. ii!db1~d~e!i ~rI rro!11J ill;Je- ~ t#.alit!l1<11 Scte~~~, rQ!!nd!~~CI!)" We!.~ nli. 11 !;'II_ GilldmM ~rlf rruiltu~ !!Iii~iu~i\lfi'!, SIliP;P0.iitifl!il1 Orn~~rI~ MatlNnal.
"'lM'l'.lCi~l'I)ilg,Q!W(QiI(l!I1ltliiU'rlJllll:ll5fS:e19ll!.6921I!1Cl

Sa.

~216. 56a~ f.HIO-=),

9. M. M •. Su.umgt tot ,Cmtl7.l,i91" 113 (2001). 10, S. It IOwlitJlJfY d Ifl,. ((I'M', !.ftlt. 95. 99 {;!00!4')" U. ], 'WiiJ;i~~, !j:., C 'G<l'iI~, }. Am. OIL'f.1!.s:ar. UI6',il61,;!

m.. ):

~elI·ifiQ2~D.O~ aQ 11ft. 29. X, T!lIag tr:' bi, (1tMI, Si!ll'. :ll21.,ll~51~ (mOS)'. 30- (."[, c~nn~:it)_ C_.. ar~l!r. I). E. Sto'!fif, SdMl.t 2:'ill!i, &n

1m Dolli diIlO!~Ilt!!, -:&M.'D or.he Au dll~11 r(!uOO iI! I!w m'p e-dges ii~~rAu '~JI!(!~Wr\l ~tQ
Wl)r~,

Mi!ie<.rful15iliOO ~..li:il1gd~
'Fi!Jl;, 51 ~~ S7
,liid~etlOe:1

ll',,',

.~ Ol;:tDl;Ier Ol(l{:i6,; ~~ep1iW ~, lF~brw~f~ 2001

l200,H"

(;j!JOO!2).

10,.i:li!&rscl~m~~in;~~2

Ml~~"'·dlo, ~Dep.arment or iliPtllllld Ma'ilfloematlia and p~r('S,. 'FO'Ilo.' 11Inlvers~• .,-u.n, MliI1\¥1'1l,
~"Qliln.,
~\lfII\iI,

rovc~'C ofSll 31fm.lshad !been de:posiwdL Th~sc adalmt'16 im::o:rporn.tc iIl~O .he d hiler rows.!O Ci'Ciucbuckled Sl1I-UC dimers dmt CfliliD adOjlt two dfITcJ'Clll con:l:iglJl.rnliom, 111.11.can be swfltchcd 1\."versibly wi! ~~~, SC3ft11Iblg Uiwm~1illl!]!; m iic;msoopc. I~STM) i.~p"WC IiIld dlal olle of Ihc.,_'\l:' c{m~'igiJmliolls malllflailliS dl"C "mgutl,i'l'c frc1o,"-clC'\:tl:'Oll ::1'i.J[C. Wnentim (SIIlI) a~om~<I.re d'ePO~itl!!rllen1l deilln 9~rn'hQn~um (Ge~ (OO]) smr<l.cl:! at Iroom 'll11m!le~aitlllfe. "ihcl'c<!S lhc {l~hcr ,t'C~h."ts Ihe Cl'}lld~c~i()n eiCoCm.ul:Wded,!Ili'mers o'rl:gin\1ltilll~1from the Sri ilt.ems are fOmledillt t~e G~·dil'il1er posiitli1l1l1.We~d~mlilie~ !rom, and ::lOtS up <l :;.~,nClin~,~w,;w'Ci.1:ii.1iC'. :' 'the diflll~r tiS ,ill het~ro~!!rumm, Sn-Ge ,dimer Ib:f ~@1j!ers'n9 its :bUlcktll1~ oriienl:atnol'l with 8i s.til,nll'lin!jll On a d(,~,Il] Gv ~OO )~ll.l:.li';lI.cc.l\\'O a'tQ~!l:-' I tl.lnU'le~;lfI£!II1Iklro:S~~J1e{SliM) at so. kelJ..iliil. Alil at~mic se~~v Sllfil.l:h w..~ Iomled fOr'~I1:~diilmem;ional ~t~'m ,I dijm,,'f wt~l mld~i'lillc tmuds of It ,Iud a !~~ct:mnb::mntll!ldion in nt~ Ge !rl:iilil1ier-m'll dirediol'l hy 'I.nln~'l'he SiJiM tD Ireveni.b~y [;Ii", '~he bm~lililgJ 1;_'(}qjl.Jg<IIIim~~"Th!..~,di~ni:;r lilr;:;; (bl,lc'k I~i.'i) fi.um Ihe ,oruE!1nl:i3tiolllof the Sn-GEld~mer iiI!Il:d to S'E!~. !Up :~t.uadin[l-\.. ve sl.a~e.5. .a :>I,IIrjlflC(; ~dl!nli.:. ~uuJ dlli:: bondng It iIJ'ld 3111 il~lldliug 1];- ~unC$l ~OCiilJ (m the IIp'lPCr "'~Id: IOVl'lC.r iii'.i!i: ol1iSiderable cil:"on. hl&~ I:l'l:CIli dcvft:1.0IiI t1.) l/-5l A. mo.llooular con ro:mmlnon challgj;; hilS i."ll.oms or tile diUim::r. g:espct'lh'cly l.tl,.. rile Gc dl.1ltcrn <1:1 ig-11 'iT! Ihe ~~,~O direC"I~.ogl, I:orminl,g. il J ~'IC swilcining cl:ec1:iJ.1l1ic C\1!ulduClilllll bQCI! propo.'K:d t~Cl\,"'iliJ: <lOOl.her lypC oflll'lFlo,",crue .:lintel' ~~I" Tille Ilc~ghborlng ~inne:rs ifl Ihe smn'C _ dlro1.lgil the 1l10v\:"iI1'tCiITI .i aWJ:llS. Atomic o s""n~c~ la 111IIQIc(;HI~1: :>\,'itch~ (6 U), I:~owcvl!r. the ~w~tche~ have bec~ rD.ill1cd l'iol"':I,""''OOl tW"{l< cl!."'C- d~~cl obs~rva~uoJ:l of di~t'ci\cm OOllducling 1'!'.}1I'1l- d1mcr rov.~ :lire lJulckJoo imt~IK' O:ppa~iiC CcOl\oIlig~_U:fUltol1.~ltlclthe t::' dc:<.:tro:n bch3VCS Ii k~ il ~'rodc.s t~!o11 ~ .;J mct<!.l lip 1Il!ld ;)I. mCl\o.llstlrr~~) w<!lY::1wi~b<!ltnom~!; ru'Q1~!tiQmI ha..~llC~1'I dimc1Jl~l w j. hy c(mlJiO~Ii!l1g tll~ i'>11;lli"1cemi.!!J;mli.o:n of atiOms or aciJi,i!\.'c. 'Oai::'climCLI~ly illl C()wd<lIJng Ch.al~ 1~1 ]J) rn:-e declrolJ <uh:mg dIe dinne-rrow ,~.IJ It ~hc fO:l'I"ual.ion and ;llln~~li!laii.on ol·m~a,tol:nic blidgc aiJomic ril. .. . iliofLS with chan~'e\ in oond~cl_h!hy ~$ ~\'3S recently :l:hiW<t11l that 'Ihe hlUd:!liIIlS Orl'Cillalim; or the Ge dimcr ,an 11-C r.cvcrnibly co~lml ~cd by tlu~i1:1, 11I100il "m~... ..11115 dut I1<1W ll;."'CIi1 sillud ied!. ·)'51 llm!ltilute f,[f ~LldI :fiI:a1.e IPlnysb. lMi'iffiity[]r ~o~ H-5 ~he condll.l!clln,g ~~Iht~~yis PCIJ)C~~dtcuhli In tile a $ul!'f"-Ce'lli",,<; vo In..gc of Ihe STM ll4)r.1iI is Ka~iiw8!'!Omdl,. I('a~hi\'!'~hi.. Chlb.~2 n..aSOl lap.a;m, ~RJK.EN dUreCll.ml ~h,Il t'<.'!.nre iloogW wi!h $C'.allIl~n.g pttlhc 0011 fbf!"111~i{)n Ch<lXlg<(;i3~ncluc!l.-d bj'ine]~lStic

A,n At10lmic S;e,eS8IW S,w'ilch 'Fo,lrm,ed lby TilledA:s!Ylmm,etrilc ,811-6e D,imers on a, G e (0011) Sll1Illrfae e I

tx_!IW(:CiI1 con~d(ICI:i.lIgm~cl lloJ1Nlilduttilllll ~mltcs of Ollc-d;ufli£msiomll ~.'~ r,t)\\I'.:i; of d~mcr.>. .011 II :;C1l1i:m roildue~i~I.!i- Gc tom, sur,i~i1eon vi'~~ich spl]iP.:\e II

.C

or

.HI;!ai!ll~ I~;U~~ at Sprlng·t ~-i·1l K~,

If-I~@ 6]9"$148, ].alj)iIfi

Tbtoori oi!o8Q,-SS<52,

JO!jl.lIr.I,

~l1i,;tlmcl~ orulic s\\'.~~'hi~\g .. bc!W(;1;'11 ~ll.dru(;'~il1£ aud l1oUlemaclUJ;'tunM ~onng~n~U~i(lIiiS i~nOI. !'l;:\'cr..ibl~. ~A: n.~1IC!tt. '~ik':lJ~r'I!"3L~Oi1 of !owitdlil1g \ IlOw

SI:;'{l!1A;:l"ll1£

gf

~~~C1;:~~~

c;nTi~rs
~]C

(mill t~~1;; Sl'~\iI~ir 0.", ZI [~u~l

'Ino lhe ;!i,lIId~DL:' iI,I:n!dli::f

,decl:tic I!i~~,d.

Cfrdt~

~iilli~r)li~I~~.

REPORTS
Hence, \'I.'l; can i,cl.ellli~}i~bc X QlfIlCI" as <l dimer ro\",sin~;:l1!{!i!1g Ihe X d~~1~"I~, mQ~va i~fig., III Allhill, s~;,ge, the ch~m~c;l!lIy I!&YlllHm'tl'ic di ~lCL~, Su-Ge ,Hmcr . ;:I:;,. a Uii,;n; li,'!!~ define ~hL;:~:il¥!l.'ri!~mg~>d, h.~ghI,1T !h!l[l i!ll"~ or ~~i;i X dil!IQlr bt,;~<I!m~,hI,; ;llIII!~ 'i..~!Ihm Qlf thl; G~ d i'!l~T.A n~r hll-;illgillllol il~1mJgii;tl Fig. 11.1::1:, nh~ Ge ~ il!llCf in the fiJ,~~II;l,-siwl~ i i~l\'l~ !'IS IU lJ WL,: :;1.1b_>i'L"ql)L;llI~~Y ~c,n~lk~ !lH ofi:he iatmgl.::d ~~~~l di 1l~I;.'r. :Ilild dud ilt'm~-d~ik(,': i.~mc dim .... us m~ tic <II VI; = -!.2 V ;Incli I = 1.£1 nA thlf'Cc lime;;.. l dim~,. wllich r~!u!l'lcd lh~ ~rl;',al~'Icl 1](.h2~ dnlifla in !n nfclc~' loO d¢tr;!l!n~n~ which ;Ilt"}~n{I r tht:: back 10 lilt c(4x:l) do'~i'i ii:1. 'Jhi:-: prooe&,,~I~o .. S~'"G!b"di~'k"!N .i1s ,L\ di~ ui1'PC'~~- ~OW(T"~~fm,'i poor [cVCri>(::cl il~iC hu e~ I i~.lg o~icntr11~{!I'iI (I r~he X ~itic!i1I. w~n'ttllc'i1,un:,"(llIj~' bl~h"\,Onatt~ dt'~D1dtli£C dil1tl::i' back 1.0 lilt, !:Iri!jlitla:il poS.i[lio~l. and itt>, ~~'lh~:>~d,~l'n(TI>.Inlhe ~illloo-:ol'l:.1A";:~I'i'ljJY,~' (.ri~,~. . imag.cheiyln reMilfic>ti.lb shewn ii'lFi~. !COn A arid f':), .hl! lirpj'Cr IltO'i1"'~ lwh(:fC ther.: Sltfltc:> the same surface, we II'oUlr.d 1:1101ihe X ,dimers loo!lli.z:c) are im!lged ,~ prolrusiol't:>" .run(l:in Illc \\/C'I~ inm~oCl hi;J:hcrll'aalll ~lc (led LI~CIiinlhc cr:I'ply-sl<lle il:l'lm~ (Fig. 2,,,B mid D),. Ihe IM\"C'1i' p( 21-!2) .aoll.min for Vii! 0.4 v: ne~r ~IIJ~);[lrollt m()rn~ 'f"\'ilcll:lilc!t" $1:lICSlocali~c) m'C seen, The ~wo ~lCilblc MIIl~r.·:tilIClml;.':S ~hcck'an Gc (fIOl ~ heiglu bee ..me the same as Ihm of~hc Ge di~m tJ climer \'V<!~ ~r~cd iriSIl",,. Urla~~ Ge dijmcr in Ill!! ~OIm~ {/!;J' WIH.1l1 I~I~ dom,~in ,~~rlbc\c!.;'~;Ir.U:1(,'ri;wd ]b.}' 'w!lt!her Ihe bluck~in~ rners for!h~~ both lH~od~ ~ml'ly-~"'!Cimllo$"''l' Fig. 1, A al~c1 and l B), \,'!'bK~·~, Illc 1 dim""r W~\~ 1l1,~g~'d .. ~ t i :'~!T!,il.~rly (l\ffi;n~~io~on~lC !If,;;ighbori.ng dhlW'Fmw~ .~~.~M sh<:'lur" Wil~ ch'in!1lcd to c'j4!x2ll. Thus, lihc !\Uf'of or in I\~!IL'I£~: a e~ "xl) !JOTlll!I.VI] Fur {l,,J:t-of.ph,~~~ I~L~~~'l~,I~c:~tru:el UUtl i;I()Cs n(j~ govclll 'ch~ ,app;.9!i"· ~Mo !h~ Gl;~ I.! in1tif inl!~h,,~ fi[1k"tl-.\:;I;:JIICin'l:)lg('! u ~d : hi ghc.1i" 111 the GI.:' dimcf in the C'H~I]tY"!'illltc' 1!!~l (lriC'lIl:llI[iO~Siil!lJld III !l{:2x2) dalml.in r~n in"~}.h'~"(,' Cnl hci!jlh~of the X <.Ii mil;.'i!": These rcsld~s givecruicat inrbf~nm~onon oric<n~l!.iom. {fl.. I'.J,. In Fig. ~A. !lI!Ol'~ or~h(.; iIII'M!gc IF~g. 2.. C ;..n'~ D ).~n the STi\i~l1b!'ierwlTi(m ill It'r: l:no.... of~hc X dinlL,;'~ ~",hibi~ [h~ 1 dillJlL.\f mw~ Inu:lkle il1w ~.hl;;c(4",2)d'01I1!10lin. L;:1>. ~h~ CO!l1p~i.lion ~]11.:l' dimCl'r. !f WI(lX din~~r X tl:Irrnc b~a,\A'o~!.iIg~~ dl;p(:~ld(',~II,;,,~;is lh,il of the U (:cpr (til: aile X ,d.~lllt.l'J-., \\'hieh ;IPJX;af~ IlighilT ~um i& :1 dl.\l~:nicully ::oy11II!l1llJlru~ ,dilllll~,.'r. s u(:'h ;JL" OJ p~I[Q' n d~[n~[,. i[ should be in'MIgtlcl :~Ili S the sasne d~IlIK':f 06, f 7l. fhu:!i..nhc U d in'!!lr is IlilOon;: SI.m1" 111(:: illImgc- limn ~h~ onhc.r GL~ diI1l1ielS3i!llcl: is; o~ manner, i[I)('S~"Cli \.'c of its buckl i~clg .o:ricmmian. ble thlUt the L dimcr, ""Ct.'\'ciCIfil lh~ Sllib&l,tillC Gc.dinfmcr !:bOJ;;ilj;tllliuHlaCJ~~ by Il:Ie d:ol't... ch\::~. -d

'When \'!!.'C dCl)oS~l1:l" slIIbnmnobyCl ~\'CI"~ agcs 11<0.4 m'(mo! a}'~ {NIL) U5H ~t' Sn "lQJn,'l C!'~~!Olh~ GI(l ~OO') "l)rr,,~, ~oo\r~n~m i~1li!1lC'f~ ~t~l~ IRH ;;I l':itll,;kltld cli~lu,,'jr X dlin~"I·~" i~\'lfhicl~ 1.11~ c;:lR,Jt:iroliic: si~lll.';S. \"""'-oc d~Ili"I1:l1lrru~~l tJ:IO';o;i;l'or ~hc Gc dimt,;~, W;M. orncl"\,~ld ..l~hc !'!J!hslmU;; (i(;·di!l~l;'.r [fID,;;i~tioJL~ nSTM Ill"'~LjU!\:.~1"'n!~1~1 ~ FlT I( /fl, J l~.uc;tL~i\~ Jillcd.loil!<lte il~11LJgI;.'S. t~ilnJiloe S bi1~:;'volm~e Vb = -0.4 V iillld, 'GILiI1i~dili..!!, Oun'('TI1 .I = lJ:II'ilA) of Ihe s,~,..JCPO"'ih ..'d «0.0$ J\ill L) Ge I Inll) surfaee Ht :-:0 IK. ate :s!w\vn ill~ ,F'ig..l. Am C jJ,"I}. !lndFil~ I, D Iii) F. :J\m\··s tine sJ!nlcluml medels ooI1'CSl)md~1t t~ lite' Ul':clml\~lIflr.:liIl"C."I ~~lIIrrolJmfcdby s(i~:id lines ~n Fig" :lA. 111,coro m:e

the ~imcrs inlbe

or

:5l1lCGe ,[(IO~)surface CilIll be rcversib~y c;.;,chlln~~"<i IPi!!!_ ll. "A~o by 31M below so K U 4).\Yhcn ~VC' :ll>l~lioo ;[1 (:e~5,i'lfe filLed~sl:a;teiflllpUl$c t~ia.'i.\'O!tilQ(.' f~ ~ -t 0.7 'V [mill 0.4 V i~~ iI'ges ~I'r.[1 SfI-depoS'itd ~h~ c{4x2} d'om;JIiv~. bll!ck~ing(lricm~l!.ir.H~ of G,~ (OOU smhu:~a(!he the d imers .irl~l.c dii n~cl' ,ITI\\' jwJ!. under (he l~p' qU~I~ed,at eo :K (Vb '"' -0.4 '1.1,1= 1,0 !1iAJ an~ ,ll~ixwal'l ~:C\ACI'S'OO. and a p(~::t2) domein elour(D')t~ (lrn~e s~~U1clln~lillg ill ~hc:dinn;'C'"fiow diroc~ion"'PlleJl'Cd.\oV'I!CIl a [ mcnd'els mrlre",pond~ \'Ire ~.C'.,m!~l n\'cr~hi~ slII'1iHcc ,)lit Itb:S OJ) V"~bf.' . .... ]flgt~ the f@(IIi~"i1g!!J~alr ptb.]) d(mrl~1il~in~hc ~ctLl1Il()di urea W,IL~cm~I!;'.d. iH~ain tAl 'The do~te~ and lillt: c(4x.2 J ,dOmUJll :1it.'1Ippb.lr(;(t\~l!lCltlhL\' drd~ 'il'ldi'.i1ll1.e thef' P'001i" bin ... \pn.llhlgcwns 0"6 V -< ~~ < +111 V, Ihe l\tlr~ doP1!S ()f'~h~ X dijiflll~f.

'Jhc c'.il1)(2) :Ind p(2:t2J' oomains

iJJl ~1C'

c~efm

fhel: sHpersmbclIUr,1;! :rem!linccll.bc Sfl11llC, Alflcr uhscr\< b~lgt~DC slirliHcl;! SchOWD~ijl fig. I A. we fJPllJikd a.PUJ]s.c'v'olmgeo If' i 0'.::1 V ror 5.QU illS. w11111 tr,pheight bed m the I)osil.ion hldicmoo the b,y the whille CIIO:5:5, in l~iS. IB. Tire (:ocdbnc.k ~oop

B:eM~erl (A) ·ami (lin, \~ iI pptioo ~ p~ilsil!vo't!alg~ (l'f +O.a: V ~'QJr 500 Ir'I'l:S~ ~llilt '~hle t~ip h!~i,g I1t h

ofl,he S ..11Vi (10:111:1'0 I w.ms Cliil at I"~- 0.4 V ilntl/t ,0 alA 10 lix iltc t~p' h~'lghi. /i, l)(lx2l dmmlhl W<llI cJ'oC.11led b}' l\.'Ycrsing Ihc h~ck'ijil'lg 'o;J1,eull1llio:n of'

~'i:l:l!l'd ~t t~,i!' pos~ili®1'1 (!), Pli!y 'IOI.lage ~ ,-+ 01.6V (3) seal'lll a~· 1.2' V ]ndiciillsd by '~hl'! while noss In (B). lE!eh'IJeen ami (0, \W s(anln.ecl~he 'im<l'ged are<lat VI:i=~lL2: V <lInd f= 1.0 'riA. l1h~ (8) bll!Jes.ph~r>I~!i im~ne S'~FlI.II:tmalIIIliDd'e~ weprlils,ell t the X dhnEirs.

fl,1iI. 2:. tAilIlnd lBi} Empty- ,Sind ml.ed~·:nate

U1dimer

'th~1J dim:~r aJnd [~C) and' '(10)) th(lo:5!

'CI~

imo1! es ~

.&n

Be

,O!flhe l~ dhif'l;Filil',

~~ ,a1f1d F) S~rtlJ)ctlJjl",a:1mDd'~ls of the U and t ,~imerrs.

+ 0.. 6 eV' 0

REPORTS
We nexr ex;mnim:d Ihc energies

~lll!IJt liiiKl;S,~ cd' !l'ie :11'* CkCTmljg~!Sc,..:!._-d ·tllml!,!i!ilh the R ~2. ~V .Ic~ s!Lmible Lhlilll;';'·!fI...,mh~ II dii'IIK'i" wil.h!.l1J~. [oCll'ctliiJJ1. 'ni.t: It<ttgc dillii:.fi.'Ii.ec ill 'abc renoC'~iojj, probalt!m.l)' Ix-~\I,~~ili the U and Sll-G~ d:bl'l\.~ wilh 9. Si~ .l1om Ifll. Ill!.: !JJiPfi.'"~'-IIKu\l1 po~ilion. This ~11.rulcs eut tl~ jJOl\:::lilb~,c L dim.::U', lFi~. 3E}f1lso IIP~I\.'tI iili Ihe c(4x2} .. ~mb!!~i~lIIjonf Si'I 310J1'i:l hno UIC StOOJtJ li1iyer. o dom!lill. These p'heilimn~~mtCJn he qUlIHllmti\'cly unI~CilOC.the U aild :l t1hnM rcre~ 10 I~i! fX'lS~liml dersteod b)' :-.trucnu<l!1 and clb::lrill1ic dH[cfences of Ihi: :!:"in :ll.om (Fi~. 1, !E mlcli F). Oml lite cleen Ge ,~onl) surfaee, a In Fn.'C- bCl,\ 001 tllc iIJ ,md L dil'l1crs (Fig. 4). On the clean 'Gc (00 I i MUf.,cc, ~hl.::r.:. h.and ~Nhibil!ng ~Icc!mn s.yl'~C'm in l41u' d ifm!r~!l'H!'1 d ire(;!!~Jl i:'i fMbl~d. by !h~ ~'!1!1ply Jt*~I~"tm'!1.", .U!!~ e;ld~ r!\;'C·dl::l1:!ro!l ;Ch"II:;;L(:'~'fis ~\~iU.lly Cf:l'll]~s,~;d of uroi~..b. of ~I~~ 'IQw('r·dilll1C!' ,1~Oll~~ d in1..,:r row ·~'1111}\,~T(;~rd~"d.,IN ml ~'!,'u:~!:li~ ~ ';.vire. bl the ..!oallJ~~· om~r ro .~n\'·~l'i~igm!.;'Ihe >eOlHlucticm of ~~~!.;' ID (./1 .. J O~lC, werds •. 11u:'p;'" dl.'CtLllHN I'unp" ..gwl.! ulong I'hc di ~1!.'f mw lhl'OlIgb rhe 1{I~\'r':!," ('k'cnmn~(; :>n.rtl.:. \\"1.;' obi'I,.;'f\·,I.'dslW'~dh]g W',!t"'~ C-olllf;((1 fuy !]~~ r~n\;l(;l!Oll .u !!he SH-G ~ d~fm.~G~~ "~rJ.ml'iid~. The L di!~J:;'r illl tl~~dinw~' f(1o\\! has ;). rniIJm(':Ill i~.1 "dlld r) ill1~lgi(£. TIll(: dlJtll' ~~g;Jt:i.l is. Sn <I1t!n~m dJ(; low.~r-.mClln pos.ilitll] ,tpi:g. 2Ft tllillA ;~!>. he Sn ;:u.mn .~~I'Iled uri 'mh~Il'"clecuon t proprnlio:ilal. to Ih~IL1{.·falclem:ify of S!!ll.t~ :n. a specifle 'L:i!e~1 IC!,ldUml ro.~I)(l.l'id& 1.0 the bl~~" path, The .It'" CI!.!cl.I1JJasporoll!l.glIliJi,lhl, along m'l:le

!:lU~1~~ ioll{1~ ca1~~I~1bliO'!lL'l 7), U Sn-(ii; {~ij~l].tJr W~~l .:11 Sn amm[u~ i:h~ t!J)llIC'r-@I,pllfq JiXNibiol~ O"'ig, 2E) W<l" ~(~~Iud be 0,06 tV more :;;il[l~JJlc i.hml ttl ~h.. ~w~~h.,1 S n <110m at~hc ~O,\I"~'!~"1~1m ~ poiil~~im~ I(fi,~, In. \Vc ..[sn COl~~lra!r.\d [h~II;:llttID' of !itL;: G~ dilticr w~d'i ;J S".• 110111 ijl'i Ih~ ~t;cmi:d kfl'L."f,

k!nd~ or'~n.(;;

,dimc~,

on

d~i,;basis

or thesetwo

n~

or {_!i."l1S!tYa

Th~!1\ the 11". electrons were reflected .at I!W l dime!". lu centrast, '~~nrty~."t;IJ~eF~g. J(')I and l
w

dJ/~U" (rig ..JD), illil~f,!~t'O Or~¥!.¢'i,II~~~ ! dilll~!"!"oW iall thli; U di 1~1~r. a~~ui~~!!1 'lJl1gJ~f IIl~ :;;m~w
Ilrlc

be pCrLmbi.,'lI;iI by those oflhe SI1 <11.0111 b~CiI~!iSC !h~ [i~ ~w~r atom ~hcm~ci1I.~y bonds \\.g~1il the s~ ~PI~f <JI~J(l!i1I, The ~,jf~li:!il!blere:nl.lc!ion(:"11
.~ ~tttrib~I~~ m the wll4k:;1rntilmrrn/JiUlg
ii{~in of
II

il~t~m~

c{]mli[~o~:1 m.. the L!1:Ii m~r. did m1mn:-\:",-u1

r:r~lCC

(,"{liU~lIigatiml,

of any M'iHd!l!lJgW;l!vc~. al. I~'~ bias

1i.II~~'~ I

voll~g.~WI/;

rio = 0.4 11;1 L2 V I. Th~~n1~,U!l~

The U mid L dimcn;,

can 00 rr.wl!rsibl}"

ex-

TIi!ft.. dime-r

;'viiI:>.

occur \\JIm~~1 lilL;: bias \,{lh,I~~ it! OJ; V < I'b ~ ~o, V I FiJi} 1.1, l!~ f(~I~L" illl Fig, 3 5l'itli\'il Ilml. 7 1 IDlI9wmie :;;wil~h ean ~ rom~~. for ] D eleentmlic (~ndUel~ml of! db: r~iuli1ei:" OO\\'. tlnlikc c-rntvelllion~1 !liim~~i.c swik:hc.~ V -5 ~hi.~mml1Iic :>Wilcti J~ .11(111.:Jccol1rlpanicd by til,.;: mass til:'olJl~!'iOrL,i,j:r~I:oms. bllllll:\C$ reversible c:on[ormal.ion cbal1~ d~mk re\'CI"iJ,I~s; of 1IIIc dhnc:r_:~ltkli~g ori""lr~mi()n ch...:miC'O'Iil), .<t;<;)'!1lQncll'ic dimcr.~11i ~h:~;W.!'I.\1L;'. we (;'al!lro~l\d tII~is:;.wil!Ch <liS. 'lll"ltomie

(;h~!1~cdi [jy STM, mid ~W1ch~~"d1<1ngr.:::dot'S no!!

'I,

or••

.n~n

~X~'lW WH.l

in movie S~U 7),.

:,~",'i1i;h"'b"ultu~u".lilwd

in t~!CiU.c'i.Ct!-; o.rf'i.g. 4

IFCe:rer,e;J'lCiIi!'~aiildN(IItes
;l

O. M.. ~!#r. (. f.\ lUll;

(l991).

!,II.

E. :Rl!~ge-. N~Me

~5-'.

600

2. O. P, ( •.Sm'ill1t, .xrenre Z6:~'. ] 7:1 (:li.'il,Sl Pfi,.: RfV- ~rl. 93tlE28:3~3: (lD0II~. 'Ii. ],.A. SIm:!OQ. it J. (dlll.a.. 5c~.('1:1~ :3:D6, 2~2 f20041. $. K. T@r~k ]'. H:aSI!g:ll~ .~'. ~'kj)'omla • .til. PoroD, lItarurt 43i3~ ~.j' ~~OO5~.
fi. C ~. CoUitr
(jj

\lQll~c!.

dimc·.r ro\~ meet the SI:! .mmn 91. lil(! L

1!11C'

di mer.

3. f.-Q .. XI~. iL. HitII~~. 1Ot!. OlM:rmiliff.

liM.. SchimmeL

Emply-glille

hnm:gcs .or

(f~g. 3A), lind dlMF (Fig. 3B) same dinl"r ro'\\' in II1\:' pax::!}

domain \\'il1l1Ihe L d~rncr llC',\q~a~ ~~cc:!rm~k fln '''Iomding WJ\ e UQ). [!."i. 'w<!!\l~lcn,glhim:roa::1cd as
~h~ bii.1~.\:Oi!.1g~ d!XjJ~a.'>\,'<I. and '\Arc Q"Iur!m.wd ~hm~hc:band dh;:pcf)l:icm a,grccs wcll~"':ilh ihat of ~llC:n. oo.nd ol1~lIc c1ei11J!~ Gc: ((10 1.1 :iilmilCC'~ 1J J'.

wlsich c~nll"Cfllcct the jI'. eleetrens, U()Iwc\',e.r. the U dI~ mer in the d i~lcr 1\(Ilov~las; II Su al.om at [he uppcw-,wJm posiiion ,(.fig. 2E~.~nd the lnwor mnr~l<; nrc .~Iwr!l}'s (ie' mOHl:-l Ihw!ll!~hOln
[h~ ~ill1~r r't'}W. 'lfln~,
thl;l :no!< c~ocl!f\n!ls '~m:l! he

(1/., 5dm<nr~ :Z8S, 39'.:1. U~~t,

scattered weakly b,y the U dim.Cl': The elecu'()i]~i!c s~atcs or ~hc Gc ;[litOIll, ho\\fC·\.'r;;:.l",shollld

t, 8, 9. 10.

(.P. Co1lier (jj i!IL. 5drMlI."~ .0l39, ~l!:rl2: 00). 0 z..I. D!!l.mau$ei' e.[al... 5'deM~.'~jl •. H!H (~OOi]J, G. K R.l'I'ID(h~.ndrllnel (1[•• !icamt: :3!()i0! loliB ~2003.~. Y. ~Uilal<;e,. l. S'Ili, J. m~~il&:i~ :stJ~!"!olii;na.,'I'. Mdjilm, lit.
~ lift 5, lOS7 (lOOS).

n.
Figl... 3. im~gi!!$ (~Il '" +0,;1' Y, t = HI nl\.) or a iliIimE!r rvm ~F1dlldillg lll~ L dlimerall'ld U()ii1Ii!~ O~}~ ~heseind.lIldJingl I.ti@ U lCiimer inl the p(2x2} d'o(.A. a,i'!~ :B.} Emptv"slatll aJ'11lil ,(filii!

Ill.

M. l~tillli~ ~ tli.•.SdfJlre 31l8;,l:00t1I 110(5). 12. MI. ril~~~, M,. t P~Iffit~,.I. lilt, )oommopguto:s, I'hy.1 .1i.'ev.Be 38, S1)4·3 n~e)"

B. K. r.II;!iat!llj~ 'y. la'hgi.IH_ KIJ~~Mf~,oil fIll1]~F. Koomirlln,. M~, Rtrt. .~ 1'2. 1.<1. Y. l'a~ig~'I'.. ~~lliI11D'1o.~. ~a~I~~ii\ F. K«OO!i. 'i!J1. 5,i\ S.5'~. 1 ~!lQ04~. 15. Mt ,1&. tlelii'H!t:i1 <'!I~, Ilri~ oomlio!!r demit\" or Ih~ ~ O!I~ iliilI Ih~ Gf ~OO:l!)·!\Urla:~1l' Ii ML • 6..25)( jJO·t!j illtcim~m~. 1J:nd~ih~~ !iD~di!iIioiii~ a 'r~ .X dinnelS .af.~ f,arm~ pl!1

:l:~n.Qa~2oo5'.

imam tni').

Th@ dmtOO ck(resm frI@ ,~mJll'tS'~~ nn!o.'lglt'5. i!ldij. (.al:@ tJue L~and U4diiTiIa

~me pi'Qrites ~t1oJ!gthe· di~J rQ~ indllldil'lg [he L. '~r UI dliUIlier hi! '~ll!i{jiliN fiT1~~.
(E)

E;:mpty~ts'!ildiJdlll

Empty'llfalil'

,d]I"d\I!

l dilifiier

lUll,climer

pooiOOn5.

Filg.4. 5chmatics OIf proiJ~galllDn the :ilt:" ei'llctroll15. a;'Oll mil {A} the L and (B~ the UI dimea:.1Ilte red alnd b1Jue: atoms; reprl!'s)~rnGe ;lind! Sn d!tolilil~,. r'E!s~ectil;re~y,.a~dl U,le y-eUQ~~'~lJIlrv!'!~tlo\ns a ~!ll:ndu(:ij()1'1 p3'1hway~'o~t~@ It" ,~l·oc~I'(J'rl.hi! l ~.~.el.ectrCl"!,~are l1e'~le(tlld .at liIle· l dim.er because tlile Sn ,gl~llmis ~I)l(atoo .oil UI~r pathW'ay~ :111'1 [I F1" ( lramt. ITfLmt. af the' l'!~'eLedr'!Jm. pa5-.S (hwll!~'h l~~ ~ ~m~~b!?'~~iSe th~ ~o~er atm1l1~ are ilJllways GI! ilIItOI'JilS '~hrlilll!Dh(!ilJl~ t~e dilliner aow.
I()f

::i!O~·::!O-llrrt :lll!~. :~6. K. T~!1I;n~1J,II::'N~kilt~uj~ ll. l~mOO..f. Koml:.t 'Sif4 Sa,. 111 ~~$$.; IP!~ri~1 W~~e. "t 1it!F'1f.;ir;r~.O!'9i<!b~ comd'fI'IlIl1Ol'lll15,17. 1'. :;JM if1l1lil!J~ or tl1l~)',: 'hm~at m. d~jjlls ill am d ~clhtioo ~(q.rp, :li~dl fI!Iolif)l!' '5.1 r:I\!rnrrmltOllin~ ~.~VldlJ'.Iil·n~~w.rtlCflilil~ wlU',lIh~ b~dJ@d Sn-GI!' d!lnu .ilftl· 3'o\aila~~ as WP,pilfri~fJl maltrf~l a\!! S6~.e' Orrllin~. lit 1I1ni! SifM ii'I1~]1I!ftn~_1!! IP~ilft!,rrmd .il1: 00 ~. U~I' !!11T.!-!!\!~~·~t!!J~mtQfldi!:i~m {~1!)I. 10~~ ~.!d w.;ii ••~~ d~pasit~ oo'to ~ d~an SIJ~1l1'11lE .IIIIRli. Ill!! .aK!rr~ge SI'I CQlmlM.l~ W3i1 ,lt~ than alli) ML. n. 1l1neeNId!! tB!~Ii!~ wt!!le' ~tQ,v:ife!!l !Jl! t.!l~ll;lc_~.{jm'lech!Iiq.~~ M!I'hI!~~ lip fl~~i'i<!dI; rum~~~~\~ b~ mU'!'!I~ ~ mllxl'ul~I~d ~.li'ell',mM f~~~ '10 lIlf,lI'kJ' ~·t 3.0 IkHt. j!IJ !i'l'e-ihank ~ 'I"~liIliIdOi i!I'Hn.I~li1glla~ rad!fl!i~~~! d~llurl~lrn, Thi~ilfo(~ wa~ ~anrnl!dl oot !:It pJliil r_dil ~llh~'ln!il~llItf! lor:r.dill5Jt.il~I?,~kJlI!l th~ IlInWt~~ 01
]'o~

:S~ppfirtin!!ll CI1I'I1r1eMa~~li'ial _.ldM[trnil!J.argl~!ii.l'clllll'tnWwI.l!':01S1S;ej,.9jl69Ql[1C.1

ll!i!!elri\jlI;~dll!,':.e1fu~~

Fig,

51

.ffi~I"rllfl~~.

M:Mi~ Sl!
.21.MC!'I'emb~2GM; ",(apled :8 IF~a'fli'~O!Di l{tT12~~i~mci!.1)184111 1

1698

REPORTS

IMI!aglni'fY'ingl Superlsas in 'thl,e' Visiiible IFrBlquencyR,811!g6!

su,ped@l!s, that: >GIrl Ibeililt~g,r,at~d ~nw, (II 'CQrrvootlorKJilr.:n~rMd 'Clpnli~l rni(fios(OPI~+ Our dl!siglfl is oo,~,d Of! ;1! IlflI!.dtit~Yl'l:r pJh(looll1k m~tall11(1t~ria'l 'C~!1si:sliF!g ,O!J .ah~~natil!l~ lay~s of '~Q:shh~ and f!eg"tff\l'~ rehil!c:t~ve ]!1d~, as orig]F!gllly !p,r~iPosed by
We dem~,f1i5t(~le a! ITl'ClgllifyillQ
!(I!f

W1Wl$<C v~l~ilY}. Thev ... d~~ .h~ PMMA r~I.,gs r {If d'!, is chosen so ~Iml n ~~l! n'!,fiz. w!hcro d~ ~lC is '... '~"!Ill ~f th~ g9J{!-V\'l!f;;:~UtllHi!' '~I'!~.rm~ of the
illt~~rlitC'C_A~iholll.~'~~le i~n['f~i,~,g
a(lt~{m of Oti!t sLi;pt.;rik~u:;:

knl:; i.. b"~lldm~ i.h~ o.ngil~1iJ1 1)IIIIHlf

icl~\'l. ils rll!.lg_nit1C<llio!~ d~,pcnd$ 011 the ~ht'! Ilwl '1~11!'I~ _~ the; !;;upotd'~n~ I.c;:ru;t 1o rmp<!i!;l"l~ in !'<I}' in
mli~l~di311 din;cliou wh~n '~I :l:= -11~1:l fig:. i It). d ( Tltft.,b\:ht'l'l'iol" \\i,l~ obi>clVc,d iu time ex,peri 1i1it'i1I. upon iinlimi~ul:tit~i'i dl.C' ~en:;,~\'i.h)...= 495 iHl' or Iru.'i\.'i" li~1t (oottoi'il~ poi'ti;(m orF'i~. JB~lor·w~l~ch j1rltl il",d;;. The nm~row b.;!<Iln \1~:i:r~l.cin du: il~l~S.C us p:1:oo!lcedi by 1\,1l*!:,uit~~C~f-i nll;!'g;iI1S(if $ lite [ocall poi ~n b>' '~he<l~~em.u,il'l(l:!. la.yc~cs or m,aI.e-1'1:1])< \\' II h pesltive and n!!g.al iw re:trm:'1 j, c i~ IdC1<.

N~!'ili!'1M(I!i!' !lli'ld IEI:!.ghei:a, We iI(hlllr'l€,d .;II lFI!:soh!lthJI'I 'Cmlihe order i;lf 70 mJaln(lm,~~rs,Ttue IJIse ~u(iila inil:Q ~Hyilllg sup erLe,ns :'l tlolll~dlnnd II1l;1im~rol,l'5 ~ ~t1.(ati(lln~ j;llima'g~if'l9' a

On Ih~ OlnL;'r Jlmld, if 5 15,.;nm ~igI1~ i~used, Ihe _ T:CfLICnlili,e:;:.lmagil1~!oo~s Wl.dl C'\'Ci('IL;'~!$ OOCClIl1~';S lU!llCO!llll~C!I!~t~d ami the 0ll~ ic'.d ~it:ld d~,!ri1:rlhlutim]~I",,,it.k,: !~ k'u-~ ["i;pru.dutl.'.); Ihe U C:WS!!.lOp), tcchniqlle (6~.<I!nd in pm1.icl!lal'on inc~'ksin~ ~IMI,\iOlI 1)C'W~lJit areL'C'lui redi rlhiC L!On ... cmroru rate or I'L·l}g~:. in mmllt~dll](IIoIzy and the !i!~!lIsm!l olll ic.~of:'llurn.II.'C !~~a",l~lOnI~Qlolla·~ ~icld disuib:\!liol,1 in tbe nomml··!~~a.unonic :~~m~" ~nic!'obio~.og1'" W' cOn!ijnu1.1'. I~a~\a.w~~r. ~ ~hO.iiP;lll.i'!I iW!1." ,(SrPs). Tile' ~f'CJpcnic~ of I~~II.~ ~wu· O!!.'l J~~cri'lx.-d till,,!,} (top pOrlion on:'jg 2St ~lowf('m~lIlion 0 r CI:lBvcmiol!uij l"IliCmiie.opjr~"~in~iIL.O'd din'l~~I1.~ic!l!!~ul ol~~·ic~ll~nOlJQ;;. .wei l'~u"'en.i~'!H I,.!I.'ICT. in I]~i:,;~.unplc'('·~h~"O~r~~li !.k".'CripDio!] of by l1:lL; d~tlhH~'[io:r~ f 1iglu \'II.8I'I~~10 a \'.~~I'Ui.:l mh~ o On WO,IySllj ~\Xrt:'i[!'!· d]!,_~m.. n.: d~scrib!,)d in d",~il in mh~ 111~:gn!iJyill:l;l ~11}l]1r1!~.~11,.~ht'! (D1l!~i~ l)ic~l,Iru !1~}I' pt"SCJ'II~-dill Fig. 11\ lt1~y Ilc.!..1dto be S~IPljlcOruL"f at" 2UOlfllli'l!. Thus, vil1~. P'IDlClus, [liN'!\. (;tl TIle wave ",~emf or dIle :Si}.Ps;i~ d(;fi~[j~J by ~IlCIilIOO hy~hc :1II'1lwpic ~ dfl\C'[i\'{; tl.1OOI.ililt1l 11m.. iiI.lt'li(\Cll~(!J;,al1dilmurlY othL"1" Sl:amplc'S 'UreUm~K18~ the expression ,~ibl~ to vfil.Olimli,Z{': Wi~l 3. regtlJillr fl~~ero.~pe. One ory presented i.iiI 1-1,.5). suggcfjrtcd wily to OVCroOll~' [hi.~ limilal,ion is Theunagl'liryillg Deliou of I:he '~\Ipcrl.I::'lls is (I) demonstmwdill ,Figs. :; and 4. Rnws OflWO or oo~"t!on 11lC' CQIlCI!~ 0 r a SL1Pcr"-'il'IS ~11. white IiI relies an lite, n!I$C OrmKltC'riaJls ormllC'L:."lJIlllIteri;]Jls Ihrcc ~1\l1 t\ dOh. wen:' plooLli:cd rte:lf the :i nner M ~llm h;lIvc IlcgiU Lverelia(tiw ~ndco;x ill thC' \'iz;,ib~~~ where £~Irm) and I!d/{ffi') arc the fl.x;qllcm:y~ I'IIlS oJ the slQ:lC'rkltlS (Fig. 3. B flnd O. These dependent dlclcctric OOI1Si,,1!IJlS of the mct..'t1and r.1X1] ~CUJC)' :mngc. NCill"~.~icldsupcrklliS iImtginll F{m.'s,of rMMA dots had O.:5·~m ~!It.'!'i{:K~iciIY in WIlS rccenl~y dCI1l1(}J\s~lilled 3\, blll~ ~le· W!::h.t!. the (I:i,z~(,'(lrk" respccti'ocly •. and (" is the speed of ~he md lal tI~rcclion. so that [)IJinSC' 1'Ilm.chi~g '!ncI\\,'Cicnl lhC' inc~Jdel]\ 1~'C'1' Ilghl tll'Jld surface II ique· is ~indtod ~)i the r<lC~,lilal, tilt: Ullillgnli flcalio:n 1~11" Above the reSOfnJIlI, rreqLtciIlcy Ill) dcsC'.I'.ibcd !Il'l ;!sInOns ool~ld be ach]cvcd (fig. 1: \ I. 1I1lOI1 J of tl~(.' l~km1!!f uperl'cns is (.'qUI:i! to l. Thus, lhili~ by the condition s
1

luical
[001

1].liI.'TOilC(lI'~:~!,""ml

~n\'OJ~ualbi,C'

Ihl' s~W!d:il;,.~· o:rvmncrio1.l~ and biolog~

,:oe{ka~1~I'iOJU,l!'.1Isof ~J:Il "o'il'ilic,ll hYIliC'rlcmi'· ~4~ "m~wl11"lCri;d C'fy.~I a 1 lens" i 5) ,mo on the reeemly developed ,dru.~mo~HlssiNlcd lIli·
.. l1d

,I

pl~rl:;.l!1i -~ullcricns C:nlm'~ be· i'l~lcgr ..I1£d 111110;(1 i (ti!n\,~l1t~ouml ~plk-..lllrnliC'ms,(:op¢ m inm~ nl1j(;('~s smaller lIulll. I~'~diffmotl(mlill1il. lilt" SPP groLlp mId phase \"d(Jci~i!.':'s :ltuy have \\f(; (ksel'ihe I:he rc~IIi:t.miHI'I of i'l1iII'Ifl.g,nify- or~N:ilk: :;,~gnl::> tfig. !B.I. The im.cill'llfll !\IlliIJiCI.llIIrc illl!:'; supcrlens (Fig, Ii\) lind di..'lltotlslr.JI,t(;iIS o r lhcllillilgL1~ fy ill!; Sllp(;',rJJ;!I~S (Fig,. 21\"), ecusists imc&~t~!DiIliii'llJO! I) regliilllr far-ncM optk.fll mi~ (I r cmlccntricril'lg5 (I rpo~J'(iiIIlmhj! In~cLhfle:ry!91~ l o crnscepc. O!llr design is based 011 ~l'IC~hco~ (p'I\,mvlA) {,h;"po.~~~t4n a g(l.i'd Ilinn :i!I,Jr!"i~C'c:_ i~or ~iE S.lP'lP'dii8plniol'l law I,bl"~h(:~old~l\IftCtml]1 <Illd gold.PMMA il'ncrJ~ocs (rig.i]3:} ~i1.IJ~eoc'Qu{,'f!. f De,pali'!rn@n~ cf IEl@~iI!rk~'.~ml CQm;puwr ~ngi'n~f!IIiIl!], cy tOlilngC marked by I.hc bo..... .r)~vIMJ\ lUis. m:;g· , 'U1li~\!'f5ity or Many~nd. C,l)1l~ lPikJt..MD2~Jl~,2. U'iIL Ilin v.:r:efmctil\'c hlcb::x III .::: U iI.'> l)CIjuil'vcd by "'1'0 whom oorm~I'I~d.!n~i1! ~u!di Iii@ <lddlri1~. [-(Tlail: piitlmml'lS Ohe gm1!p vclOoCilt.y ns oPl}ooitc~o (111c ~1l'Ot~n9;JumdLt~

iihlm~nmion\\,'ilh an C~K~!'IIlf.'l1 J<J~er.UK~ 1~~ft'C mws (!rp~'.lIMAdo~s ;in Fig. JB ~,I''''er,iiff; l!'ll!n'c~ 11Hlirui di,\'Cl'gcm p;1 [~m{M:b '~rny~, which m''C ,.

c:'kurly vLs ible in lite P~fLSiffil't)llimn!!';.; in Fig, 3D obtaiaed w~d~ <1], oollvcnlkillilil epueal I1miem.. scope. The emss-seefioual analysis of.this illbi'll:jc across r~~c plmll1!lt1 ··rn.)I'5·· W iog. J.F) imii.,cuJtcs re~o~uliol1 orm lIeast:70 mn. or -.An. ll.hc lateral s<c;p,amlion iJ.C'!\\"C'C'Jl these mys inl::llC'a soo ~)' I) ~~lJ(~'hU I] as ]hc mys reacihcc] dte omc;:r dIU 0 f {I ri ~hC' :il!llp'er~CI~s" TIlis, inlC~~ :lJI!{lf\~'CfJ visu~liza~km (:If the I ripici. 'by iCm1vem iO.llJlcnicroscopy. ~11:

Fiig.l. [AI Sdllen[ie Qf the lfI1agJfliWiIi!!D: SliIJ.lerleflll 'irm~~rratl'rl i~w a oo;rI'!lI~illio~al flil!croooope. The pl!ilII,m®lliIs genera!.ed !by ~he
ph!ase-fl1IaWilingWU:OOIlf ilttll'l'li·

A.

mI"re ~e samplE posrtioned ,oo.:!r (he [eIil~et' oil 1:h~5lLJlj(IJ~l1lrH\•. 'The ~~~.raldi~;a!!1c:e ~~m inrI· agli!'S ,,,''Cu:hpce,d oy '~Ile·~tcm:atT rig ~aye;rs or ma,teil11alli \\IIl~h positive ,!!lInd l!legai~ r~rlrall:~ve irIdex '~r'l':I\'lS '1Wi1h distlll!KP at®nu ![he

me

rradill~ The m.<l!!lnlifil'd image!! vI(liol\'ed by ~ F€g,uI..ar ~kroe 'S(Oj)e ~B}IR~l iIJrIIdlima;giFlCl!i1J
,l'! Fe

111m,

parts or tile S~fra(e pl~m.on

if)5

,---~--------~~-- 15<9 la' U l.t!, 21~ il>O ,,ilj, ~:J

<I!',g;

kat UJe,~ol,(f-PMl'.'lAC!ndl ~~"v.l!wlJ!min~fi!oos ~'S ,~ t~n(lfQJiI of h~q,~em~y+ 11'I1Utll' heq,l.!eflty range lI1l'lIrkM b!Ythe !lax, PMMA llillS: Iru!g~tiw reflradJlolleil1ldl!11:1:. 85,pe-rt:l!'iwd by Iplasm.olu, Ii'IIherea~, the! ,~,old1"\1ac IJ!lI!mi!ilti~rf'a(e l,Ooks Ink@ g, medtumr w i~h p(lS"iti1;i~ !',e'rraclirljie 'ind~x..
ViQ\fe. ¥~I;O!f

kc(GV)

11699

REPORTS

IFim. ~. AI~ fmi'lg~s (Po to ,e} ood GDl1'IIIeI1l1:i:!IDi!i optical mh!:rn~c,~pyim·

il~~S (II ~1iId1E) 'Ilr tlll!!

~~sv'l.ution lesl .s.ampLe!i compo5.ed of ~hre~ (A and 18) illncl t.IkVO~C) ril" dlall rrn~ :(If PMMt\,,~ots
(111!11 r~~d [lrf~rrololii:S) po-

of ~he iITI~g.Ulif'Yin S~[j

[p~,l'lens.

s1l1clnleti ne<1lr tme ,cerl!tllr

ill. lniclF'oscoPY im!lIgle!i ~~mted[ tnl ~lin and (El' (Qf['@1Sp(lInd to' (h@ sam-

n@ GOfl~!iO:n·

[preS S'nOllll'lilli (B) .me:! ,~'O. re:o;,penlve!ly, ne IrO\l{;\i,Qir PI!M\A ~o,tsg i'!,l\~

rise h), '~ither~lllre!!~

<Or

UlKi dii\N~F;g:e~t iPl~i'il!Qi'i ~raiy!( (m.<llrk'ei!lil b:v awIr(l~~)). 'IIhkh al"~ vnS'lbt!? ~ ]II! '~h~,a(ln'lj\~nt~l)nii!~DiP-

'tij{al 1iI1Kl'"OlWOP'(imag8S. •. ~F) Ote cros~ rection Glf the- IJptl(;a~.im<lge o1I.1lDlgl the< Ufleshmllnill (10:,. ]Iildic.a~e;;; IresCil~ti~1'I ~f .~~
lle-asl: JiQ nm 'Dr - ),,0.
OptiCill~. ~(If1!1!t,X,

!E

dislalii~~.

Z.

Fiig. 2:. ~A) AfMilll1l1ge o~ the nl'1I9lilifyUllig ~uper[eIil5 mdlt' of ,PMI\,~A dnlgl:'l on '~'he' g;Gl\d

fijllm

S!J! rraC!(!'"(I)

Qlpti,allileki

,dis iribultij@!'Ill'! t:he

i

lIIIIilgllilUy~ng :SlIlpilHl~~s iU\llIlliin::l!ted bey el'l'li~rrliall ~{l ~ ~In theflleqji,u;:i1CY ~ fa iij,ges where' Jl1:lid~,;;.1;' ~2~1. (top) :i!fld ~l~l =: -R2d~ (.b~U:om),

It

a s,imil.,'lr~lls~liQll, 'the t\VO rndi!;ll ~mvs oOrl?MMA dots ji!l(W.~Util F~ 3(~ gavc rise to t\W)p~aSlllml rnys.. which me vi:sl],~Iiz,t._x! Fig. joE. in The: COJHI)C!silC inbilgc in fIg. ~ :I" " ;~lJIpe,vms h ~n or the a~omic: iliJiUCC !IlIic!iI),'<oopy (AfM) in'k1lgc trom Fig. 3A and tl'lC' cOUl'C~o,u~cting optic,,"11 ilnage obmined b.y oollvC'lnw,n~1 OI)tll;JI ,UIlk:l)OSi;;0PY. It i:lll!5]1~1IoCfi, liJC :i!1lmging meeha,~i(5!no:f Ihclfn".,\l!.liIYi~!g <,uper,lc!~;".Ihcp;'Slii~I~C ul'ilh~'i~11{li~!r.;~y....~h[l)!~x,;Jl Ihe .;;oHeen'lrie ;,q,~,a h,:u,-nl~uin~g IUy~J::;' or nr~.r~t;Li,ls wi~h ~w:;;ilh'~ .. nd . .. nl!C.!¥J:~ u~'CUC~i,itt'~ ~v·c iucl.cx im:MI~C;; Ih~ IrJil(iHII

XliluJ Fi:gl. 4", (:Aj Conl p~nih! ,@,~ the AfMi millig,l? ~rom rig'. 3Asl.!!,p@rirnpo:li?dl Q~'to th'I;1 ,(jorrespoll'tdi~!lI 1m'ag'''' 'obl;1Milne!di by co n'!lelnt~oimall opltical m1.C!rGlllmpy" HlIII,S-uaUng the ilmi1lgfng m.echanism oJ t~ ma~mifyilil'Q ~lIlperrem;. Near l!lie ed9ie of '~I1e'S~ ~~d('iij$. ~hoese ~ar~tkln illf ~~re~ ~a~ (m:;llrkoed' Ibyam)il'\f~ is lo1:lrge ,~n(lllglhw be fe'5(llved 1!J5inl~ ,ill ()cn\l~nl]j)nal ~ ptkal ili'!iicn~5(QPli+ (B:) The- (Iritis!; till'(ti:Ofi i\)·f the j)'ptk,a,llmagll' oiilo:fig the l]ne shewil"lil'l~A) lr1dka:~e<s:thi! thna,e rays.

oflhC'lhn.;-c ['lIYS (I1mrn<;,j by .ftOOWS fill. lower d~.t). N carthe c~c of Ihe ::;lI[~cfleiil:>,he t :>~iilrn,ti~11 ls ~argc emYLlgb 10 be i\.~Qlvcd Wdlh a '"O:n1;'CJlliOI1<l~ op.ti.c .. mh;;IQ,~rc,tll!Js d ,dCmllOfI:stroning ,J 11il:l,gn:frfYlng ~~p.;:if.l'C,t"I~ in tile \ilsiMc
..,qp:ili.l\tiOll [f'C{IIiiCllcymugc'.

Thc IhCOll~~iCJI rcsol:ut'fron of sueh a miC'm~

scepe nmy IlI;:"C~~ 1l[[IfIG.rnel~rscale (J. the

~h'U", as Ihe ~1i.'HtbllO become :m im.',jlluabh! h ~ool In mooic<!~and biolQgic<l!~ im<lg.un£.• wh~re j~lf'~11dd opllc<llinm·Wn!l, of ]ndh idulll vinl':5f.!~
and DNA
nllUllol;:'lJl~, U1I;IY lII1;;'1;'Om~~l n;:.al~ty. II <

,no h

We C1(I'~""Ci: thil~ ~jiltl~lli[j~eptiea ~m,~n..<J:mat~~rio~!i ~h.;qUC'nc,lf romIgC'. such SiS. dlC ;n,,':Ccuily sugmay :Inc desi~llc,d and illljJrCfliIcnlled ~l8inlS the !o.!stcdi ('~, .1") <11'11.1demon sl mi."-,;! ~ U) "illvisi[)i I~ . tJl1 ncir~c of our mmgll i~yil\S seperlens, Bec(1iusc il,Y c]!o'a 10;." (d. c/z) muilhc ,Md'!,. "atio (FiS,·1'!'\~ are easy IF.!Eifer'!il:i'ie!~,~ad Nl~t('s ~ to \"'J I)' locaUy, 'Ihe effective .mi~tl'-.oph:lrC· 1. ]. B" ,Piem:lr~, ~YJ, fifv-. h!'1. lIl5, 31166 ti!OO(U. l"!.7I!cli\10' i~dc!o\ of~l~c!lu,d~i.~.ayer III <lh:nia~,m..}"
;!, ~.

b~ \'.<Il'iod ~!'lI:~nlUOU'!'il>'rlfCH!i hll'g~!lCg<l~~,~ to Ii :lLnO\I.:. \,"c-ry ~hil1pl~.t"'~1, q:O~'U51, :uu:~ ~tmi!!lM~ hi.~e r{]J~,ili.\~ "'''~I!IIil:'t>, Thus, l!II.i.u~i1l1 na:110r~'iOh~];1 dt:·".ie('!> l'i1ii.y bi.:. t.'r'o.:'~l~d i:1i the \.'~siIDolc ic fOIlMlrd hlil!I!!,'C' eqlUisi'~ iOI,1. a

f2OO~).
(lJlllTi5}.

Fc"~~\ H. ,I.A:~, C. :$1;11'1, ;t~~~~g\sri~~ 30>e;,5J~

~. 0. O. S,MeMLI~It~, fiLai~l[',Opt; ~pre!iJ '13.2:1,2:7

REPORTS
4, ,Z, tacdb.. L \1'. ~~jkJt)?"" E. Hil!I'im'In!:lv" 6i!47l20fl6l
'5;, ;A..

Q{}1; bjKm 14,

II.
~I.

:z. Liu, J•. M. 'S1eele,

rnl(il~(~OO6)"

H, Let" le,. Zh.a~~. Appl. f1j~

If'U;

as.

u, o. S!;~lIti~

~tlL,

S,iffK,~ 314",97t (2006);

~l:i;~diiIllD,. N, ~i\i!ll'h~Ia,. ,PJj~ ~~,. 8: 1.... 01!) I£!3 Q:OOM. 6, ~ I. 5!i!1~,mi!1ll<v, C, C. IDij.'~, J. e;UilIln, ,~. II.Z~~tl:, I¥'I:~.~, ll'.~t.94. IOI.l'~Cil. ~200~'. '1, A.~. Z~,!t$.I.1I Sml~.~m1M'i1(, Jli, A, Mllfo!du!liill Ph,YJ., Rtf,). 4(!e!, '~1 (~OlOiI.

]l ilL Pendzy, Il\ :Sdl,"~' l'l. II. Smllh, 5crf1JC'e l;!!2; 1780 tl'OO6~.:~1iI~li~t1~'rnlrn!! :;N M~~06 fW.llZ6f~QMce, "112'~7>, ll{t U. IL~ru~ S<,ietrr:.? :H~ :~;rntz(l(l6~; piil:fl~m@dlo:r!lin~ 2A Ma!!, 2006 ~'HI.l12''''cil!na!,.
U2M'9in,

JI'J~~h~d o~jiffl' 18 OOober ~OO6 n;iln2'Mti!1~{"'.

:ilt3J~:!I), U. ~p,p~edl:byMSF(I;fallll~mi.G104G4.6 ilnlil (i(J.o50SlH~

to! 'O!ectml!er 2006;: Mlci!pWd e, f,e'bruM)' ~OO7 1O,1126J:s1:iB~ce.ln:B74!6

C,oupled Thermal and Hydroll:,o,gical IEvoll'ulion 10il 'Iropieal ,Africal IO'V'I!lr fhl's' Last Ds'gll,Bciatiil,o'ln
th@ dist~'bllJti(lm of bran(II1(t(J1 ~tmgetJher rn~m'b~,~ft~J~s d;~r1il.iiedfro,m ~i~ bJQ,(t~ria~n h ~ m.alrii!li'~5~dim,~mt r~Il:~~dthgt ''JIas, n:((I\I'eredl do~ t(D '~il~((lmlg!l) River (ll,uftOIi\l" ,~i!ild the f\~$LjlllS 'e'nlal:!~~d 1,115o ~ec~l"I!Strl!ln lar!!.le~alile (Oln~in€tltat t~mp~raUllre 'CIilQlfi;gesinl tn'l,pical MriC1'll~hat ~p'ln t th~ pal:'lt ,i:!.5,OOO<y,e-.;rs, Tall:picilil Afrl1~aln tMlperatul"les ~radlu\ilnyin(f\e'ii"OO hOill1l ... ;n ~®, 25"( 'elver th:~ last degladarnolll, wh1'ch iill, ,iii larger ~\!al"lIlllln:g~halii estima~ed (or the tmpi(a~ ..A:Hiill'ltiil: OCMII1i.
W~i1!ln!1!lf!ed
Q

To !!ilill1 beuer inl':Oi.!5'1ulllfuo c~nlRll A~Ifi(1l11, lhe t~nnpcr.iituuc d~\'!!kifu'\i."i'Ie Q.\'~r tl~e tl1::..t d~lm. eia,tioil. its ~~!IIion to globAl clilnmic ehrul~es. and its clfcet on tile c{)1~lijncntal hlydro~osica:l, cycle. \\ C lI)lcdi rhe Mel,hylalioll index of Br.m<Ci1\!'d':e~r.tclhcrs ~MBTI nad ' )'dizatlf,'i(1,
I" ~i{l of u

Br.l!m:hcd ];C1r.u;:lhc~ {ClH)

b~Ii,'o.'Od on

lilr.. a'I'Ch~t1 ~1.>,!O"ml ,d,i".ll<.yl gl)'~X'ro~

,tGDGTsJ'

~~1r.l(llhc·~'$ (.1!) pa"$.i;.~IlA ;1 nmril1£ oll('Jru roc;~l\'\,r~, ill

close U) ~h(':Ct:m.go Ili\"~r CJll;tllow ~(ii~B 6511'1 ..1, Q5"353'5." 11'913.3'E. \\"~~rd!.'P'!JI oP:J(a2. un. rl,Q, I l'. U~:un!!l.hC! M 8:T and the C' ]3:"[. w~ ~\I;'cnol!.\qru(:lod mh~;1.unu'l~ :11W,UU OI.~rl'll~ ~lrl~'L1~ (MA T~'of the [ Congo R~\C'r h~ll\il! (.I2JI whi(,.:~1 couldllE comI'1flred with me S.C.l'Lurlaec h::nlll)cmlilu'C (SST! s A d~:~(~ ((IiiIiIl)'arison with sea"'.5l.1rraC~~f!Illpera'hlre estnlriii'l!e!lff1om the S!;l,i'IiI'~core reveal.e,d that 1\'!1OO1t'd1 ebtained frof1i1I.~IC same eore l.r~g, 2:). the la n d"'5:eii tempe~alime d]U~rleil'l(:e ~~a$, Uilr()iugh: the me ~mta I p re5's.'lIl'ie[ilrad~enl:..i1Iln 'imporlta nt ~ml~ehcd GDGTs lI'ig,. S I ):Jire ~bLm(j:JilJI core' ,col1l1rot 1:1'1"11 'il:flFltrall A:frkanl predp]tatnoll pattennll., Ine~n~rnl1ic I~p.~d:iiderived fll'Olll bUiC~C'iI'j8;~hal: ~Iuu\'(!'ill ~oih:;{ 11. 13, 14~.As the soil erodes, !hl~ GD(iT:;: ;!~ l1ill\'~!!lIy 't~~E!.n:\.po!1'rledrt1lthe' 'lal::JmCICIN~I1;:)i1have gnllui!mxxl ~hc soeree-waer O:nI~I1CJ.l~a. dil.ra:u·.,c.d.l:lI1ge du~jalg li..t" tw;.~ I. l , dcgl&::i;n.i(lln. !(,,.,p..~i,dl}' i~!h.c ~ropi~,ii>o !.1rnnl'lCl:-<i~ km, 1:lw )iCmlt ~!~,,;IotlJmml!C'l~"llire oo~u OC('ml. ~ndl(i\.;dI.~~!c}'h,~\",~hcen sh(}w~·!~[) 1'lC:In Olwli,lklbl(; 1(;w!lle Atrit1111 (:(Iln~i¢Il;.'n.irrl{p~y ;1~"~1· I,;Xcci!cm ~r..c;cr of til\'! r.~!C'",r :It'IIi! off,Finic n'!<1!~~ !1.o.,I!.~ \~~nwu:l~'IS'nod ..~ i. ~:i.for!hl;' . ooCCalt~ (I 3~.!For Atnca.,. <:1 COIL'If.'1'i,';!1!S is CH~CQl.- IK'l'<II1!rC di ITcrcncc of ~3.5<:Olo f.I~(' bctwccll d1C tel' i~~ ('(m8:0 dCClP-sc.1l rim ~!) 1lI.;nddlc Bay lhe J Last Glucii,ll M:~l;xim.mn (LG~'I) ru~d lhe presem elf Bi~:.lY U5~ !Ii:g. S2}. He k!rg.c c~!chmc!ll iug on pas~.ch"J1,g~s in hunniciity anci!hcj!" C:Jl""'1."l' :;)Ili1:,;)I, of"~hc C'uugo Ri\'l.~f 0.7 )( ! 0(' knll:2) ex~!l'lOO 01] bk~k."\i\!1 and l'I>nll!;:!] :ii~lJcli:e::>,IL~wl1;:H(ls day ,(2, 5, 9, lOt bluetl~i.;'-~c often [il.IPt't_'iCI~I"!J. d~I'1 wnds (mill about ,(i,QN~(l 13"'5. 11110. rrom uboWiA ~h\;' dl::U[L'l'ium crmjl~n nf pl,ml. \\'~'L"C'l 1415), h,u ['I.~lmi\'~~y loe(il ~i!,1;ll'll orm:c i!lm:Jnpk'~e IJ.:.'l:'m!::Iis._ ~.ct~lpcm.tLln:'reror(tc" tbr ~UCh~roPLCiJ~ Cmltlll1i111td T~lU:", kn~Ciwl~d.gl·(ifil Alri~liIn Utlp~ctul (l'~.lpo1::r.· 13'" [(l J3"E ~lli~!llil1fll Alri!;.'tl (f~~. 1.1\). w;lh elofiLI'C'..f1!S R:rm~in, f\C,fllu:: and IrI'CCn1l1p.iCI.C;. In C(lrI~rrl$l 00' euae ,d'b'UI~.gc o"Gr~hc kL~I.(!cgI:iC~1i1~0I1s rnt~lC'r ,'mio:n.'O bclWCt:U 300 lind 12:00 J.'n except rOt i Ille n.lflnm:: cMvimmllCI1.l. lew q~flll~hflli\'c l.cmll'C'l'~ ~i:rnltocl~ ~c:i~Jlj' for the \1ISl Impi.c311 roini"ofl'St one :Olllll:l.lI~Pfl~:t, leeaied ttl iDS CflSDCr!1 lID~lnd8ry. .1LUII'C proxies C"'Lllit ibr ih~ll!rrCstriftil Ci1I\' imUlmCIl[, area t~le Congo Bl)si~i. O:m1:ilflllouS. high reseIlmi. fJ:o';C'S f(OOv c ::WO(l m (I fit The Icmpern~lU'C and cmnttJ:lll.aotJ!.'> l(mg.{rnn dilllmc rn'ChiIII'G 001Ikmd ~l]~io,'n.~O:l1'Ig,~cmll'1.!Co.msor@u~tillcnmkllCaJ'c: t'C~& C>la mM.C'S obt5LinDcd room our mariulc roro. therea:re limited, For i~'I;.~~1:nlQC.i=K')J Il:n~ VC~cI~.lIjml pCl,<I!I~I'!!dmUJge8.ll: mLAc~a need 'cd 10 improY'Clhis, ~.o,rc. 1:\.1I)!iCSCnl a cmclmlclll:-in~c£f<lllCd terres,)~udiC'5.a widely used mc~llod tOr clwiromncnml kUlOW'k'd,b"C ffml ,crmb:lc'propor cmnpariso:Jl \\iidl Iri;1I :tcl'I1iper.:m.l!r'C SigJm3,i derived from land of ~;I\l~ of m~ri~~e~m~)cl!3aure dm 11il.'C5 , low 10 intcrH'lcdi<rI)C' clc\'a:tion, Amlij1'sis the' rt."CO'JlSUlI!ClkII!lS ru~kUld {51'. mlC cmllplicmcrl in il.hc lropii~s IjIOCl!U'>'Clhc cllCc!S ~1'iai112,e. lu 1J<:1'I'n..

or

pcliuure .a1i'C dil~icLJltlo diSlillgtlL'\h :~I'Ol1il !!bose of' ,h;;rn,g.l:s ~UI 1}ut:cipi~'1cioll. lelllll!elil11~11'C c~thnl;];[eti b,Ji,'~cCl 11 ;l:!lOlhr;,';J'Illtdmtil, smblC o;."yg.cniroto]!!C 0 (;QnttJl~ of lCi.wboJ1l.IH,-r; ,md $~licM¢g. .Re \\,klcly !l!ppl~(';ll in kU;:U,'IlI!·im; ~im!;:m.., ,IUd :;;1)i,ih,,'(~lh(,111:>.
1~(l\~'C""~I\.'i.I~:It)!IIg'1iI (~K'M; e:;l:Lrn,II,leJ; ,In; ,UPiJJ"lflru~ ate Ru h~gh..rc~~ul.km qUL'l.I~tHI.i\'c p~,k;ocl'il1mL>C ruroUl~LruCliolJ~. l , .'f}. qlill<linl~Ii.l;:ilI:ioD~ c'lill;Hlle of c:hail.gc in terms pa~tc~npe~~LII\.'S roq;L1i~ te~lI.ous <I.S.'>ti!i'IlpI:LO:IlS (1I.lOlit 1.'I1C'past 'Ch~Ll1ige:>inl

or

or

or

tRoy.!! Ne!herLuocls ~mtilUl~lor5~,

~cl:f(h,>

Depa~rne'JM

of ,M;;irin~ S~g~iJ ~hi2mistl)' alta T(I:tK~logiY, P\Dsl OI'fi,[.I! 8;o~59, 17m /\1!. [let! aurg-T~~ N~tl\ed~dt ~U~id~

,FoQ~~\!i!'l9$9.ei'!"lll~~~~..il, ~(lFGHilll~a1~h, (!lil!1ilr 'O~n M3rq,ins, l!Im,i~,~fiiilJ of Brmen, lm'bemer' 5lrm,e, :2835<9' ,S~meJi'!i.. GeiiTill~P'!¥. 'To \i!hOIlil oo!T,e~p~~l!oe sli!oWJ ·1Je atkl,r,e~.E~!fIai~ lwe1j,Ilf1@Ttij()~~1 tPI1l~Eml, a!lldl'e.w. ~nsl:ilutle f\i)r' GfO~ciel:H:;El$, lilri~li.a C iii'

Albrechl'"JlJnWer~ll)i Kiel, LU~~g·~JI'"SU"~se

K~0,G~~Il\lI:J1I~.,

lO,.2,;rn:J!.~

re(o,veredc:wiSe tolhf' Collig<l R.iv~r o~itfl~\1,I'rr~'m a waler depth ~,~'962. m. alild tliiI'ee);~lief'lt.I(lf~'he C~l'Ig)(l River drailiJ.i~lg,e' b~$il"!' {""h]w ,gI:lU1M} ~Ul' plQu~ tAl' oln ,;:! d1igi'l.ll~ ~wv;atfe!1 rn~p or Afrk~ [ph:hHehorn N.ASA J:~t, PIN~PIJ15i~:n~ilbel'aJtol), 'CallliorDna II'nstii~ll~.e 1reclhnology] ilnd (B) (lin ill map she~\1]1iI9 i1InnlJaI, o,f the MAll diis;triiHlri,on llii A.~rii:a IlilIVdi~ilIbl~ the ,~atlb<lll~ i,n i,n

IF;!!!. ll. O~en,l11~li 1trlaip.5 of Afd(a. "Iihe lIloSihio'll! ofcol1e Gl?o,B651,6.

um:J.

117CU

REPORTS
'!.'\II1I1:.

:limMe C<J.nlO'I!l .i:iiowpi!; c(lmcm {BJ']"l of rJ>!!mfr-~ ~kl1nes Imf>~hown I hat till: O;m~Ci' R,i \'<t;'I:
~~~[Ii~l dlf.\Lir!IJ(~

nh~r<'~~l 0.000 y~~~ 16), 2

')I

~illlbre.;;;t·\

(,ig;L;'~"~~~ ~!iIJ'~~ (w<t;:':r

The

11II~!ill\1cl\'tll!it3~S.of

·our

OCt1Utd IDnllt:-

O!Il .;~)' igh~clllJ1It;lF.d h h

pm(;-d ~ '~uh I~~\"violll~ c1inmlc ft'.Cou!b;,are in:5 I'd..

reso] ulioll

1~-2!(J(1 y~';U'l;.).

mlil,~ltUQ.1.IS ~l~ll,U't:,."ltd la~ ~ogmphirn] t:'.nvLT3!!\C ,~ile.,'OOlll"dl. IJrClpiad Afri~L). Ilia~J:im'Urc of

eur clilll.iJI aiil~)WI..'tI ...to u

Ih~

~:LtI

deft!l:iIJ~d ii15ight

imo

W::i1~R~1 ~,l'::\I<cIop:i'bCttlt of Ilmpic'llil Afi'f:elli'il

tcnnpcmtun:! oVl.'iI"lhe 1&'>1deglnei1l~iml. milC' 1iI'16 ~~u,11 M.A]" ~\;",'COtlill orth&: '('Ol'IYO B~1~ii1. ~vcals aill ovcrnllwan:nins shlc'C' IJw I..!GM ~-24.il(nr) ·~o
.26

Z;§, 24-

[~PIICADome

U'::.OOH ealibm1.l:d }'C1I1N I.1tcfOJ:C ptl.'Scnt h24 ·!O lfl !l:,~1 y .B.P;}j of4,Qo lU 4.5t;l(' (Fig. 2C~. T11!i~ it; k
w.I!l'i~'i,I\;nl ~.vi!]! ~,I!n

l.im::l O!]P!Lk'tl!~(,;"~.lp!Ct.l!il.lru~

I~

iii(

2,2'

~itlm ,~ fi.::w ether IUi;01!lrullli. ill lftli~'lic<~1 Jile,1 A out...d~ dlC Ccmgo ~3<l~~1~. f·ICI!lc!1rba.>l;!d ~mr i 'Ipc!31tu:re'1Jf;,,(l.ms4ftl!cli~~ .j;l~gg,C'$I. an I.!(iM ,11..1 . h1~doo!.;'l~ to "'5<;1e '~~!;;irij~OUl md~1)' 'nhm is, 4'l'

21

2S, =241
- 22,

.. 4'lC ~".OOI~il~ alt~~Burundi higi~~ands. (;5) mu:l

...4" SClCt'ook~r ~u tlk~ &OlIlht'lT:lLak~ T3illl!l)lllHlyiko i)fkS:UIlI (J 7),rurtl'i~fIiIlorc.. LG M 1.t.1J'ipt:'rullLlres lhnl. \\c:rc~3.5Il'C lower 1:1'1111'1 IJo.iliI,y 1~H1v~bo.."!.::.nl reprrJ'il:OO ~l1.'>Cd 01'1 TEXtoi~ '(lCIrn.C,thCf:" inac1oi.. of w~m::.1hcr.o.; oorls.iS!ting 86 carbon alolns) ~;d.:.e· !'l~L·.tiicc ~1i1\l)cr.U!UJ.'CIlC'(OIlSirue! ions for no:rilhclU bkc' MaI<liw:i UJl')~ t.;U]lf:;'f d~n"Jcnccsbc~wccn

20

or

5..2 :5.4.
- ::1:'1

:::c

:fi~6 :5.8

€iI.

6"0 6.2

- ~170

l{iM ,~nd P'I:cr.c:m-dOJY wmp('r~!!hl!J\'~ have been l't1l'oncd based on illIDllyscs or ~ta!lagillti~c s!~bl,c l'illwpc~ and ~Jl()t.Jlf)dwatCl· nob]!::, g3:foCS. Saeh 1.'I..mlJ)_vscs UK~i-aJJtcd d'kl!~lJ;,';!~~ures were ~S.7°C (,'{mjl::r iu ,I Solnh A If!<;<.m SpclC(lDil,'I;11m ,~Uf~' rld H N~il'J]ibinmad somh Africa ~9, J91,•.llllll u~ dffitfl '!i!{()J1C dcrh!("li fmm ~~k..s SulJlll~lnl~ rdIy mnJu:r
~oll'[ia ·ofthc Ct1Jj,%t(lrlll~ Co.ngo E3a,'iiI1c, Oun: rcOOf!::l, 5<;1 ~o {'f'C
~1ot~~t in grtltl ndwmer "~~Lli lCu1i

ill

p'

im:licmes Ihm. degl'lcial

\io1nrmillg has OC"C1l f o

sill'l1lir>ar I11aflgnitLKle Iforihc wbo.lic of Iropkill ,'Jrica. rllOmdlc' western :md eennal lowlands ~o 11 .

01
·'1 .

·:2'

f·

...
:5

101 'C:a~elJilldali"ag:e (kl' B,P.,

20

lite eastem hig~n~allds. ,,~C onset of p6Slgiacinl Wlln'l'n:illg un mr. reeord ;n ."brrLlt ~; i!::al ky I1.P; (trig .. 2('1 ~'> hI UIlC ~ ith tl'lC:Sotlllu.;rn iLCI'I1Iisl1heric c~bn:uc dcvelepmeru, w.ilh d1CE~H\)'pe;!I,1 1)1.'<Ojcc~ 001' .~'C Coring i~ An~9:I:c~k!:l (.EPICA) DOI'I'te r D'C!.!t~ ri~am .fJ;,'Qolld:iho<\,ving I,h~ Oi~~SCI f wanla:ing .. I o ;Ib(]~uu I j~S cui ky B..I?: ali) {Ffg. 2B) .md <lI!m~l 17 'lOll ~ kv B.P. in ,l i>a~~1 m~n1Jl.tc1'!! I sOnx:md ii-nm . Si;:l11til iricu t! N~i,{l1(""('"~'1I, \,!{~lI'k bL~iJd o;a b@Bc

~lP,l~S I)~ SJ1l1'l ba(lll!~lill C(),fqJ!«l;e~B ·~518·1. (F) The BI) lr~cord (Jr (.:~~ "hJIl~aM pbi!1lt waXci't;S in (,(Dr~ 'GeilllB 6S 18-1, reHeHtng hll!fI!Ilcmy (hatn~'esilli tropi ccd ,cen'~rCll frk~ ~6). (GI) The laim:l-sel.'lrempeftl,l1.!IH! A '!Jrlil~hml bel\'lEMmt eentral tr'(Dplcal Ahij·(.[J allld the tmp~c;a~..#!~l.[mth: O(fl'ilr'l based on remrn50 (C) am:1 {to,), ~l@l:kwia!1t;!l~.@n the @:me ;K@teo lrldic~t~~hil;l' Uc i!«J~lt:r.l!t~'r IfTIo:!SSi spe(tmmetllY r@,dijoc~dl!)n .date::;'e el'gved hllm mixed platrl kl.onl!!: foro1m~ niireral uls~d WeI" sta,biisiling UlIll ch rotrlo,Lo9Y of ,!!:or'e6;eeB e &~ u,·! (6). VD'., Young:~iI" liJ,ryl'ls'; ACR, Amtardic: (o~d [R,ev.!!'r~1Il

Fil!)_ .2:. Temp!~ri.l·ture (ha!r!g~s ~1,i'fl' thE: ~aSli: :25,OOOV~i.lr5 'inl ·Ir,op!(;i!~.(~Plitra[lAriri(oI! !l:O'miU;i!r~d with A.~·ric;,m Inmilidiit.y ~haillg;el:'i and .Ardkalld AlFltlln:;I;i!l; dijlilbClte S~9~1lih!. {A} The Gre:elll~[md ke Sl1eet P'mjed 2 {GI[,sP2} 31:80 reil:Qrdil!llcl~!:ative olf GireellllCilild alir-'i:E!'liIiIperature 'illll!::rualiom: Ul).SMcrW, :S:i:.Gllnd.!l.dI me'tJn Ilcean wat~r. (18) 1 hll! EllfllCA OOIiTil e! ( ·BD r~'lll"d ]trldiciil~i!le r off AiIllaJrd:iiI:1.I~r" temperiltme cinarngles: (2'Ol (C] The amrma[l MAT remrd Qlr the Congo Ba!>nll based IOtrnlhe' M[BT'ilnd,ex ,pll1ldl e[1n r,ati ill, of~h;e. ~railided 'GDGT Upi~5; !ill soil b:lu:teliia ~rI (OTie Geo[B 65U:·l. ([0) Tile SST ,changes ~~ t~@ '~II~uIOFial AUan·ti( O~~n b.i!$~dan a ~b!Ii'i:!i1~[p.a~~eoU'!~I'i!1l()~f1Il~~ry G@~B6:518"1 (6). ,n slt~ un A reeord or 'he alvei!'a:g~~!ilU pHint th~ (om'go Basiin based Oil the ([In r<IItio' (If brsnehed (i[Ii)G1

c""~Llre dalill!!;; OFllllJI1l1l1L'S (.~) 113:;. :;I~o'i!i!nLhal. ill Ihe mh:j·.lati~Lld,"'I;. htalih h"<llllil>p!'J,(,'lj\."S,. I,he onset lhl; :liI~ dq;.ll;\Cimion was i1C~, sYllchro..

or

<or

IlJOtI:8 at alL.Xltlit 1:7..2 cal ky 11,P.: 0\ sirn illar limi~lg or 17.3·~fLl ky 8.P;. wlL .. ~.lfcnud rar ih(; <O:ll.iCot or lh~lll:;~:dcglncnnJiionin nile' Sotl~.h A~IlC1Ci(:an nI\Yp:i~s or :Peru ;t~~d1301 a,. Om r Iropkall.eJnpcriVii ill UfC' recerd shows. lhal, the 'i!I.1~~g~!l1cia~ wJrm!~nlg in.tro;pica~ celliIJ:~1 liiea s~t1!ed allillc smnc Iu file'. A

afler

O!

~hOl1i. eooljng episode between

H'i .md

17 cod k~' BJt pro'ilWi~g ~tron~ cvidcllwl'Or .. gklball,1' ~Jueh!fQmll!~ Ol~n ~f!l~L;: hl!ii~ d~b:;'~uifl'!1. Wil,~i V3lu~ c:Jf 20.CSao 'Ul1 11' " De'ilpt;,;r"~In,~ 'I\~tlX: rulhl~t CMSj!'JlllD. uLiririg Ihe LGM. n~rule

1702

REPORTS
of S,1Ii'lS(;ijLIClflt dcg'I'<!Ci<ll.~;llmli~g w
'!D<I,.'1\\I§C'rI

was hlgnesi 1,' "ad 135 c;!1 ky B.P; "nd br,,'1\\IC~~ ll5 ,:,1110, 9 C<I~ k~ I:!,P'.Wilh ;;lin "\c~'!1I!l,i:;\,VOPT!I!!np,;

of -(0,1"(" ky~:II. PLlri:I'~ 111~ ilt'i(,'lI'\{t~l~iltg ~riod fn:mm t35 tu I L,j cilil ky [l:r., knlflot:m~wln.:;S :~ rlli'linoo re~i!li,\'cly !lrublC' a! ,!OOll:t 12.5" ;uK! w~n;, 11Vi!<JNy. nQ!~~d]~IOIl;;cd by U1C J\n'<J~\;,!i:c ('aM RC'~m~llu~ l:hL,~ Stllll!hL';!'n ~kll'iiJisrhtl\: 'Of' by IIj~ Vt'jllill:i~!' [l:rya:o: co!dt,PllilC'!dL:: hllbo~ uu:t~'k:i'il Ik'lil1IiJ;ph~,. Ermn I) kI :5.5 esl ky !J.P.. Ii:m;pe:rulure.s .only ~e ~]owl~ 1:1,), -0.311'(" k~~q. reaching ~nlfll,'(iI:l1flm\·.!lI~e'.~ 2:S~c. [,hilWil1l;llhc

whe~ thlC~!1l'emopi~1 ('GlYverg~m;c ~w' i( !TC'Z) i:l. lini~"" foQlI.u1h~mm9~ lr~,ut~n .. ;;!lld ~lC SPl.U~hi;nl I-!L;'~~.~i,';plw!'l; .\I>']i!I"W~ !he!lfC'Z i~il~ i~ ~~L'}rnll~Il'l· ... ~no~IXli'!~iijm~ ('I}•. i~s !"'I~lpo:,~l1IiIJ,1!\~ ~\~Ohlll~on ~:;. likdy!1i,:~~I~J;.~ H) dillUaI...; dmugl':i :in ~Qil .. hcmispheres. This cOlLplif!g to both ilIilJni"phercs. !;iUI]pnrL~nh~ ~d!l;ll~ !!I~t mhc o.!l."~~ l~iC d{,.'g~m;i~.1 ,[If \\ 1l!'11t~l1g ill 01U:,"I'('.Colt~ :ll aC!i:!lIt i1 (:~d k)! !lV. u:; g~{!tm~ ~It thar.n.::k'L .11.~~bo ltij,i\!:ihl.t:"'.P.l.!liil~EhL: lack
o r elear kl'i1iiJcm'[liHC si~n[ill! relmiicl to ~je Amt~· Ilrcltic Cold Revcr:s.al, af'l.ddlc' YOIliIfl~r Dry3l>. which were as)11~chron.ollt.. ~1~le'~!li cold event'; in thlc Soulhctn and No:rlhcrn h.emisl'hen:~. re~~livcly ~17),Ccnpclbsa!t1on Of~lCSC coolin!:: event .. by 'W,lmlCIi :li r room Ihc ow!f1'Gsi~e i1emi" S:1~hC'~'chl !hL;'~r(!Jp:iC".drealm appm-crlliy ~c~ul'l!:!d in!]!~ n~e,,'od peried Ofr:l.1~atiwly ",wbl\j '!I~num.1 MAT~v/.llr ~'I:;~u:!u.mildlmpLt;uil.}\'fitit;3.frum ~,3.jlo

!hie ~D' 1\"'C9~ and fclmg '\lei}, high. $(I~:Ipli Y<l~lIlCS ,Wig. 2:. E 00 Gh SHi!"n~r ncg.'i~\'C' or C\'~r1rO\'i·~ ,;,,'1; ..... ~~IC~ i~~di~a~iJflg \ll'1,.",k {l,!l:'I;u!ll,l or~"\'~!Il (.Il~· I,~ndj :.t~r ilJov.'. ~~rmld II['!' "\I~H~r m;nd.~Li)[l!l:ii :mdlowc;:r 1;;o:~1 p~I v~u~~;.,_ l1lut>., ihc .~lftd·-M.:n
~hL;'I1'111a~ picl]1 "PIUR"!'l't~JI xens .>!J :.l!w;n_g e ·on '~'\L;rllr.dJ\ fric,m p:r(,~il]ilj ..ion rml!J;;'l1ll,~, UK; cl i~el'(lJ~m;yoclwt,'tn ahi::' !'l.Timd~';Il n:crltrJ, ru~clthe ~!llll'ill,idity ~\Xltd:, diunlil'iJ!lliic t-.Cll"t'lI11;!li!IlI1 t l..'1I'the 1J()'creit~ iuighL tic due hl ~ verywesk Ir[n~ea ·te~~'ip'~rollire ieitll ~n [hi:. I.imc in~ tCr\~J 111,11. allowed o1hl!r ror.cin£, 1iI1~·lh.~ni:;l'I'~s, ·!O bccol1i1c l:'Ircdom~mlllt I~ has been :iil~~~ed tlml. I,h~ meridiom.:! IIUWI'ic<lI"MlbuJOpiclll s"'I),"f .sradicm in the S\lUdl,
w!lIltd.I.

or

seeend

jjiiH'I

of the

~>'(I.:V~ U10i~c.TIle
!11t:l Cl1f)iNOP

rel1!l:lil1.cd about 2SIklC, when the W'rnpernlure dccu\."ru1cd :<Iiglltl)'~;lin

2...

loloccne. 1Q1nl:ICl<tUllf\.'$ 'C-",cql~1 .ror the 1mS:1 .7 k:y 0

~j

~d~n~"'nl is simil<~f 10 ~]!I; r~~cI11~ d.1cY ,<I:ltrl~I"LI Ar (! I' Uu:: C!l!'1~O Ri \,,.;V bti5i~:I. M OI!bOllU23. 7~C lm'«'d ,1;1H '!.;"HJ1J:uo;'U' \\ ~tI~io~~ elm:l 121~
(f.ill,!.,

.s C
g

1'lX00Jl!{tl'lJc!t>(1 from

A'~lmmiC' ()Q;c"!_al cx.crl~ Ol~.!mng control .

ml'OC\lllr.d

IUl,

Our record .1,1;;0 b:~. a rnCJ~ ~~!J1LilL,,'d bl.i[lili(i.'i!l l!Iie ~'(l!1l!.irli.l'm <Uld the OOI.'.1il1 :J}I:'!~ld I;:Ql1li.p~~riS[llnf Impic ..~ cUlluincnM o t~J1l1p!:nllu!:'l,: h.w~ ~~a4;11] iunp<IC'~ 011 ,L;{.ll1tinl.mE<I:~ hy(iI.'ology change wj,d] .~16~1 c~imm.c dl~'I'dOJ:mtclll m'Ki through lhcl'flr~dl..f;(\f1 l[h~IlIt]ld~iGldiC'IlILi\iL.'S.Cm. S~.,. records. .ill:l p.lilltieuJ~lr :rmm '~hlll '1I!Op.i£fll i.h:IY Afri)!.:'ain 11Iiu!f,~11 \':lri3.lIJi~J,I.y i~ in IllH'gC p~urrt Al~llll1<i.c Ocean. SST. rc«n~IR!ctiQns bfli.",oo 0111 3ssociM(~d with SST disuibulicms il1idlC Sollnl!. alkenone pa I(!Olhct11f1omc~ryindiCII~c -1,,7OC· AI.lmu ie Occm~ in th@i~IIt!;! dll;!l'umll pilI:.'S:Sllire lower SSfs fOil" till!;! .~m;lian OCellI.]l and ~~.7°C £md~ell~, dinc~'S bct""'C!."'11 dlC .Mric,m co~ni,II\:I1:l lower SSTs 1'01" the ~uxmiCJ,~AI~anlic fJccan and the ocean ~!tIl'fllre U/il i\ large' m.'g.nin:!' lilnd=:,,:alcmpcr;umc gmdic.nl dlkl~l.J.;,r1.:k .. :i vdy d~H'ing jhe LCiM I)~~,ativc Un'J:ooC'm:!cmpc;r.. iI\I! annes (11). Thi~ aglil\,~ \~cll !;,vi'~h l1~c-25°C warm ~~n ,[!ld coo! (:m!lin~mwiU !\''.fIld~ill a m.lkCillone-ba.~ U]M-·I ]O!QCCIlC SST. mlll!.c!]ji. :Qcrn::mll)' oiT-·llI,ud illir flow. pliC\Clnil1£i moi.~Ii:lir ~llis siie (Fig. :!DI, l\..'1w!sent::ltiivc rWIJJ.c c·i!!..<;,1,cm rm'Jn nile ocean rronn. nowing ontcthe comDII.Cl1t

' n ..

I L5 coalk~ B.j]~ No~~bly .~IK.! d~fn."'t·!l()~ tfi.'1,lIac:v,d'~\O)n11i!lg in

AlnC';lll Il~'(;eiriwliO:I! ChiIll~" (6), Thi~ nlL.'Oh'li1I1~~m\'IIIL':!rk.l'i~ll~h~ l!iaL~1C' cl iR'Cli¢r,!:I0:1.... !.h~t..Jtd~1 Illcnnru gmdiclIll: A. p;:1<lt~~"cJ,r ~W3a'ln lmpic:' •.:1 00(.';l1l w~l~re,>luh 111~.mnge:r ~K':r~diCl~ml w~nd:ii ns ~""en~ orr~I!ilJld \,~linclf>.b(j!!h Il~\ ~~lll!.in!lmoii-;.] a~, ~iom nt]i\'\'ing (mm the ,p~~ ..~ O:~IB.{~th~~("i.llilt!:·
.liC'III,. BC'rom

the

:11.C!~oocu~~. IC' H

Iand-sea

[hCillttlil

!l,r:;UJit'll~~~'~)S~c

rutd ~pfiucljj~ly cxclt1,"~d:I. dtllTI:~'

mhc land-sea Iltel1b119~ P i:cnt re~nail1.cd w~ak.the In~ridi:oll all SS.T grn.dicnn increa5.Cd and L1CC1llbl'i: donu Il:llll in conuilYJ ~in8 im:cil'iUl!~iQ:n ill cC'll,trnl !mptcll~ Africa. !1K'roby (:,msing. inCTC!!s.il'!,g;ly 01)'

11&1~[ro~~lrol. \!!..hereas

during the 1.~oIClCClle. W:l~e11

~1tIglion,'l

1(6).

H\;~lCC,l~lCc(lllilii ued

CV(ll\r~Ll)!l

of"

Hupical Atla!tlic Ocean (6), TIm$" cQm(l1lJll.~ wUlh Ure (lOJ;.:'<JII'l.the rcCO!1S~n,IC:'!~'tl cnntin!:I!1I,d cl~J,lI:u(;~~ .ITUlllliMg Qf 4" [0 .:j,-r[' is (1rnbid~llll~ W
bly ~ml1~"::If_ l"hi~ tlill1b.t::Ilt1!: ll1IC~wi,\c!:lcOn:Linul~ml <lU'Id ot"Cflln it t1Cf!;.inciful \van'fliltg in l~lc Impics, is in 3~cmc:lTU. will'll dimm,e model Siuaics., which

1'1.11tdV~~lmgC' fllii.'C(Hl:s4mcling ~ O

co!ni!1I!:'Ilml AIl'i"

Imld tC1npclmlmC mod SST. in diilTcrom pmts of~hc i\~kmlic Ocean cunuXl~loolPl1!.'Cir:i;I~~mlImJloc:ms in CClnrill iUh'ctl Idluritl!!, lhi'.: P":-tl ~5.'non,'(,'illS. ) IIire:t'er,Ill'F1t!e-,jO..a.liI!! N[l~e:i.
S,it:1l«: '~II. :ill 3 {l);9'lN1. 2. I.. hiii'@tIlMo]" CIJirl.. .!:ljij'l. 1~.,:!i:l~ (t~W). 3. 1,M. Schaefer 6 01,,, SrWJl~ 3:12; lli:MII i]0I]6}' ~. F. Gam. Q.~tlt 5£1. ~; U" 189 {:roam, ~_ ~ il(Inn~iDIe, .1_C,. ll!O!!i!a'iIIll,.1 Gw~ N):;onult' ]iI'U, .;Il'ii!l (1990). 6- E. SdieluB, 'Sc S'd1Q~~~, R IR.Sd'lli~l(!.er. ~1)Jt;g 4;!~', 400;1
;l E,. 1l!;lI~,

C;~!1!c[np CWlyn.;s riDnI~~,~ marine COfl! is: Ihe- po.:o<. sibilil)1 of C(ln~I:I<!lrinlil 'nJ{; ©~IIIA!'i WUI~l ssn.
dcri\'(Xl fffilll I[he ~fm~e OO.1iC1~6) IFi!:J_J[)~; We

u:;,oo Ih:i~

.e!,lIlilPflrii1iQJi It) JJ1:1CJ:mt\lll1ue1. tiE

Ill~l]m.l

:mggcst tha;tthc 3\'Ci'fIgC cmnin.cntal deg.liieifil warmhlg in, Ihe tropiC'S was ~[bo'llt 1.5 titllCS SlUl'mger Ihilm thc dcgJacial watil11iinlgof b!lrop'" icll~ occmlS {.23. 2"n ,[his ,<!ill'npJlilioo, contincnml
W!llitnh1!l( may hrr:ve Ix;cn ~~,ffil1Se d'l\:' cont~lI1eims

cool

I1l!lOIl(!

dmhlg: gllada~lim.cs.

AI high m'ld

gmd:h~ll[©erWOii!UD oc.uurnl A frtc3 fllld~hc ,e<1l5>Li!J1Il l'K'IU{lwr:ial Atliflmll.ie 'llcetltl over dl(: pfl5l. ~:S."OOO yctll'S (a..11md~: fig., 2Gb" D!llrinlg ,g~ll:ciallimcs. the cli.il:liJOO~ ,r,(!cmd 8C!11cmlly s,lm~s \!:JI.lucs or I,<:11)0 3~C. \vhkh di.~PiX'W· Imdwl" ClLlick~y<I rlcr d~ OU1SCl of dlC dcgl;tcimli!:m .. ~1},\Tivc \"'l~HeS d('JIwli~ tl.'l 2~ I:'ellt111 ,m. 300m, ~,2. cal ky Illt

(2!Il05j,

7. M. ]l. iltnq" 1200.:1),

1. 0,

Mal'lhil!l~Oua.t .:SCi. .ilJw. 2:~, au.

:Z3, '~Ol {z,oo;1).

!h F.l',!icl!)eiiJ!®ii, Q,lIIilIt Si{.i. _.

ullid-~<'Ltil1.ld~s. rllC' IllfCSCltCe of It chan.~'Ci[J mK! olwn l~dUlocd \,¥..lI~~m~ cm'>C'l:" d!IJl1ulg ~Iaci;)jls rcSl! ~t~ [lnOUllh i!1CNa~d ~alb«lo, in CII1imnoc.<i (;'Qolh~!ll Qnh~~,lI1d~llrl;.~ •. l-iO\~\'c~" iUI 1JiI;>:p:k:;d OlU\~'as.!hi~ 'clr"-"'I 4-;; c;owU)L; cl~'Id by .;;~.~!~g.,lth',~ ... f(1l..xil:mck I[~ln:!, n;:du~;:t:d ~v;'PQ"~lij.{Jn.wh~chn;lOul:ls.rn;~.r~.'1 dC1l;n:~;~,!>{\d: Ilr~lflicOlI rilillj(m:_~~ 'lR:jJ. nH'; rodll'C(1t1 cVf1po:r~Iif}Jl :Icnd:-;1)001decl.U<:I&;d 10.::;8 OE

ooincidcm whh ~ rasl SST

lilltil~~

Tnuhc

C'Ilsti:t11

c'Iml:tonm.l AI:I,mllc (').ccan, and gmd~!;)jlly dis1!p~lc;~r... ain m ~le St;."1:Ll of I~DClloloccm:. Dm:ing g ~h!i;: ~ll oku;,f.: n~ .~hc ] 1Ii~~d·~I~.;:mpC!<lWI!C gN~· di~ul """,Ii F.B!h(.;f~"I:M~. wilh \,.,~II"'L;'~.juM~!'I~"'tl,,'r
dl!lU 10"('_

9. 1, T. t:illlorigMrui, rJ. ~, l{iU1l11,IE. '" St:IMJ~ G.I!tIp'~, kttf. ~#.t 31, llJn(h4 ROO:!l)i" 1!Q. L A. W!:iMi~ t.r'Qi., Gt,ophf3: Rtf, Ult. ~:l:,. lO[i,OJ!ll
l]"

bIJt~I~thCil,t <1m:! l~luS n:~mily~~y \C'I'llnllcr slIir[are ~,(;mpc.illllJiI\,.'$. 1\::; ;0, fCS:Im 11,.l,!Ql,ct."LtiOIll chmJlgcs inl Orrl;mH~\ IHllkaru::s from the S3IlnC l\.'COro,(ti) ~mpic-al ... \:I'r:icaal"e Ih~Hl~UI(j! rc:.'ull C"¥(!Pltlllallly iu autl with Ihe soil pH: ~];t't1 dcriV('iI lrom ill&:: iii, n.cgl'~n~~"::~0j~1W11..~~liUre ,,~;';'Ct ~lj). .A 11IIoD~ likely c,xpllmlJinOll for th.:: !.."iIl~u~nccd glndal Q;lO~~ CST :miitl of dlC 'brom::h...:d GOOT mcmbr~ulc il1l,£, (l\f the o::mlillen1a:llropics ~nay Ihn .. beml ililcrcascd Iml e"t~'1Llat{llrlC'nlpern~UI.'C gmdf~nl. rest! ~t~Il,g in OJ sln~lgI,helledi ,and cnl.d ~IOldle.)" " 'dl ci~L~lalion (1m .mld, 1.""{ll1sc'lLlenrll:y, ~m in", C[~a."II;': ill n.:la!il\'Cly 0001 iii f lhlll flkll!j,\~ iLcom

lln; inl1l1i,,;lll:(; '[~:iC Imld·$~l!L l:h~m'Hil ;gm· d lieu'll un Cl;11'l1; li,;;n1.U:~ydrultlg,J eaJl~1he cy;aluull..:d h by ron1li}ctri"~Olli wil.h CC'.ruroJI Ahr.i.c<l1l huullidily challlgcs rell:cc!oo iii I h..: I)C!lJitcrfllll'n1 OOlll:Cr~t (,~m

or

12-

taooS}" ]. 1!'!1. H.Weijert, S. 5thg~'~i!'l" ], VOl" '~~nl)~!ker,. E. C IHliprn~ J. S. Simm;F1~h~ Oaiftl!:l~, Gt«h'm COillIla.llim. JIno71. 703 l(iilOOll'). 1ru'!~;!Jt~ ~ rod! ffI~l~ ~ r~ ~lllamt~ a~ !I~~rI!ifi!;I

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ch<lIl£i:'S are '[]ss!llIMcd to ~rk:c~ dli],llg~

~oi:l pU ill pre-

dPLt,ltToill iItIClrn,~ty given dut itl ti~c ~ongtcml. wil":rc<ldg~ng pn;:lj;'.C~cs.IlCSI.!~l in dlher ~t~lJll,ger

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Ga'

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fIlI~'l!ri:l'l &/I 5dM!! !)inlim•., 1l. e, C. &i!l!l'1ilrn, .iN: 01•• E:rl.rtlr f'lai!lft; Sri, ~f.rt, 2;2i11. 1~j!' ~h 111. ~,w.H. W~ijM.~ al., If:rlJ;(.J"lM, t.~li!'m~ioJ,8, MEl {20015). 15. G. ~~1l9!l 'Itt .,;>f" :sri~~3U" ~3 {iro~)'. ~6- S,.h. It;aT.ad~ F. IK.aI!liJ.J 1&!I"p,li~.~Ar.<rtl.lll!l, U:l!il U'99.6}. 17. 11'" (i1<l1j~.C. R. kad: Sd. Pam !l:i!O ,w5 (U9~. .• ~e·, K. liiJ~rngr!!m t\I' .(It, Ql!!~t. 56- ~: ~, l3:U QQlm19. ]. HI. IE. HellIon,· A. ~ T~lml. ]. c. V~gef, (.l.twr, Re>, ,lS" 19 U9.B6). ,~lJl L Au!llJ~1ifl ~r aL, .rIIali:ifc ~2;9, 6:n ROO4)1 21. f!a)\lII'J~lmtiLanm Mflb~KIkiogIcal tfi'!1lj!jjb~ Welltd aill:!~te !'nfooma,j~ (WKI .iI,OJ; <I'I'<!iliil!le- OIllirn!
~);nrm.Ii1~~!T]~()IJI~!I11;!.!~1:\l11Hl~~~

d(,'t:JiI.'ilI~(:ji, .fl:_5p~livdy

22. A. R~5eDl-i\'..{i1i! •• r:;i1OfJ~ RH. btl; 3;1,1 t!OO~)" ft'tl1 H, 8, L otW"!!~i~~er { m" t (lim. :!.~, l~,6 QOO6), ~ :2·11.$, I, Shim M oJ•• Cum. Dji'fi. 20,. :127 t2.003~.

REPORTS
~S;, ~,

iI!I. W~lI!tI:a. B. ].MrAVar!e)',

~2(10!l1
,~, Ham~ll!!iirl,

ClYro: D.!'rI. 1", 9'2]

n\t!wl',

30. C, 1, WillmDlrt. (H9S).

j~l1..

S. Ill. Ih'be'501ll1 llilt..~', G"matocl.

:IS" 2'2:1.

S.l!Ip1PDl'tingl OIll'Ui!lle N1i1t~rriill

"'lM'I.5(,lilOO1m.i1g,aro1Ggi!CO!lllftl'lU\fIllI.lB15.f5:819J:l!7(1lJl:!tl M<!I~~I$ .;nI metfilgd~ SOM 1t~id
:F~, 51 iilnll ri2

'If•. ~!I!~fln'i-l.@' 'if~~l!Ilit b!

(lim', D~

M. :Foo~~on,S.~a~lIwml!, ~1996~.

14. ,B3,

~$.lulm'. ii. M., '~8, ()ool. fI.~ S!, 217 ~~.oo!ID), 32. We linin!! lit R, ~IlIlHidHf for ~r>O'iidlng~lIr'lIp.1~ !rain Wof Ge!!6 615,la·1 <1M ~, ~~ !-!'o,p!mn~rvr <i""l)'Ii:i!1 ~!'I~!iI
iliiln ~n~hlgll"pi!iffcrtin11l00! llqillilt dmm1lIDgra~ll.)II'fI'Ii!~ ~1l!~rHmel!y ~m!ll.!1l1!lntf!l.lIihii! m~rdl!""",,~, 'iUll~l!Iflled, br !il~E!~.~ ~f~Xi~Il~.(t;I!,mei!gl'1h~ ~''!l,1t1~ri~ Or~ilnl!::llir.tl far St:iIr!nl'llillU!~,:rldi .illrDil Dl!u~ !~

2i" 'iI., Bliim1er ~ 111£. ~~ iI.~ ii!l!. :2'4, 2693 l(n.99'~ Z!, P. (.a1!1!1lMilitil1lS. j\allioon. !. IlIooc:ard. 'nl. L (1!'11lafol. 2:1.
2.9., D., W.~1iIilKM:I. 11'.~ ~!ill4f1,. I), [. ifadm, It B. Sll~ralk:. ,

!MeretlQe-:5, ,l5l,r.tO'l'MIlhl!f20M; ~O!l!i!lIM :2.2F~r~~ir'J 20(!l] 10,1'l2il1ociB"(~.IH8B 1

~n (~@(IIl~~

C. F. Tretlin., Wortl Ail

~(jaPQiM~:il.Jl)S"2~1

(1993),

fl:CdJu~g!ii.!,(!mf!lll!;cJ'rilll,

~M!IIli &: ~R!~Sh'I91,:~ IKalffl~m!iiM.u@M~!'Illa>ch:s&

t111:uaLd ~i~I~'Il~$,.I~ ~lIr~el1

de \'11~~,:!:.!'i llilbel'~:s:tall!ldiget~ H

Ii, :'l1t~tM~dijk~ ((Ili'npl~x '1,Iifi(liIil'llihe -.3.8·biHiilln-year-Ill'!d tsua S!1lp~.;IcrUSI:a[Ib~l~ ~I$B} in ~liIthw~.5~ 'G:reenh:md ~'ro\lid!e1 ~he otdes:~ eviciel'lce of,'lllc£!iillnk ,crus~li!l({~eliol'l by ~pr~ad'ing. Ihe geOlLhiemltStl!:r1 Qi~ the di&:ie-.s, lind as!ltl d[li~e(jl ~mm!l laeas di1!rmm~tr:al'81l an f1lillr<loOi!\1!nk i~l:im d .<Ir{ .<IlfI:d n11.d·Me1ln IrndEllHi.ik@ < !il'nilil{l" al'l(l~heir (lX,yogelilisalD;Pe1i :HJggill!it. iii h~dro~h:erl1lliill 'O~El'a liI"floClr'-'lyP1i! m,etal'llmplhi~m. The' pl.llO'LllS aJl'ld dlir,res are: a,$$OdaltM with gabbroik !:lInd u'ltramafk lr(l.r;I!:s th:.atlOgetliu!!rr l'IiIiI~ Up~UI ill,phioUti:c dis"mdiatijon: the Pai~roi!ln::hei!lii! tsua ophIidlil~ ,mmpLe)t Thes.e she,~t:ed dlift~ illir:t.e'r evidiel"l()E!lor r"e1!l111il:imls ,(!I~O:(j~al'lk enllst. illifrnoed by '5.eai,,'ilom spn2'adi'n~ of t.he l~aJlil's;t ]ntld mc:1l:.:s E:.a.irlh. 00
SC'CIL"()]fIS (I r oce.II~~k: crni'.'1.that were ~lJel<ltod 'by sea-Ileer _ Sjllt!'ilding (Iml ~;illwrCll~pbccd O~IIOCfH~!inCnl~1 ma~n~ n Origiml'l~y. (~phi{llit~ \'1I'QfiC' a..... "H!!]l~ld l,(~rep]:~':'l!3~! OJ;:~1~~!1!i,c:cm:il r{lmKod,u~ rnid~lXlWl ridg{;~. but ~]is \~I,;"\'I;' 11l!l~d~nMJ.'\i r..ldic,1! Iy. Ul'1rl .~~ i.. ~~"oI~~lIg dl.'>u· u1ml Ill(: n!<l~ jOi.cilyO:r ephiolites ":LI:tJgcltl~I1Il\.-din s up:r.t ,~~bdllC'~ ~'On"'Y_onc ,cra\1~mlTlnl.'ill~S(1). Dl.;'lxw.;ling on IIl"teir cwllit 1I.;lwumm:rwnu of fo:nllmiOill ,nl'iJ ~ ~hcir ShiLlc'[l!.Imll .rrrcilitOCllJffi <'1:[Uj g['OChl"n'JiC';rr~ a I~ ~~llitics,M'i!lncro:-.t!o;c ophioli[Qi ,om be d~~'1I;;l!ioo 11110 di ~]"cllCrrn lypes, bul. lhc llilfdoriny are .W:;llCIgw caMy rc;J!!lredtoslldn.i:UC"llon. ~nironmcl~~s(I.';t A cm,np'lel.c ophiQlite cc::msiS"tsof submerine 'ooS3llnc \loiJCllIllk .ltOCks (maInly piUo\\, :lil\\[1.s )" sheeted dikes, ,<I Il~utonk ,comJ,)k-1:l.<!i~d UIrPc\Nllrult~:C

oceanic Cl1IlIstrOj'.~noo. by ~e,~"f1oor ~1'i'l'Cad~nt'~\!3:;: re'l~nedl w :Pltan'C'rO"~otc-1i ke p~me ti."Clo:nics ha.<;::lO Illr .rcnmimxl, conjoclLlr.lll because oflhc :ihilC'rlCC oroom~lli~g kiltcl'llmil": evidence' W d~l~inllrmtc bCI\\~(;11o{lr.~gin~~hmugh hori:lOll~~1 I!Hrktiolil (Ir Ihe phl!l,.~ .n di\'~1rg>L;'nI jp],IW bl'U!J!cl,lri~ !U" 11:ri\wgh v(lrlie'IJ I~ol.imj ;.Ib(WL; m!lll"ltk p.!IUIi!.lL,'S. S!lt:·tW~d. ¢nJL!1> p'nwid~ all. :ll'i:,;w~r 'tu these qliill"":OLiolv, because lilley limn by sca· n:oor spread ill!;; .rmd. ru:ecD\cli()l~ durnnlg h.tlriito.r:ua!~nQ\'\:m~m at dhlcrgl,!Jll[PIm:Lt' bm,III1u;!OII1'i(\lI.

mtd

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be mJei~~1

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It has OOC~l stl!!gc'Swd~hfrj several AlfCh!J!an grecll· slIme bcl~s. host dh'IHClwbcrcd {l-phioiil.Cs ,(~. I)'). N ml£lllC~C;",....~h,c 'Clues! ion or WilC'lhlcor ,1\l'c~acan

repert jhc t1iSllffU::1i)' of,ffi, S.hC(:iJl:d-dJikero;rnp~ej(. withi n the Palcoorehclln [SlI3 stlprncru..,ml be~l. ~IS:l)l, 11Ji,s rmd d'l(! .rus.'iocimiX! rocks, 1J}g.~thcr: ~~ ith!l~cir camposltiees, meke lip 3 ~3.,~-hilliOl1iyc.."l.t"-o:ld ~1Iliol~!C'. \,\'hich in tum 11m., :>tmtlg
Fi!!jl. :It. ('.11.) Lo,cati(i!i'l (If the ISS inl 'S/(!I ulhwe~:tl!! [In GrE!'Ei'n~dill'ld. lh~ bl.a;c:kSflU!dlre'~h®w.. the I.orati'Of!1 ,(!f thrt C!lr~1 d~t~~tel!:liF! (S}. (B)

I~

IS8 and (!dij~QUU gln~i$S!!'S. 9!'1~tl bo(ati~1rI (If th~ ~Wea detariledl inl (C). (() S:impiil'lied '!tie(!l.ogic~l !iIlap
ef the we~tem !ilrm o1f~he IIS8. sh!Ol~vi g ~,(i(ati~l!,!iS ('1 to3) ,(jf thEt n l!O()%shil!~~E!d"d.i iw com p t!!'x (3: ~Bltitlll{h~ ~,5.05 .BS;(lN~ lel'lglitud~ 50.]IL661"W)i gra{iio'g ]Iil.'to lCiilk!el's ill n dI 'Ii',(!1 tea IIlIk reeks (2: ~atitllld,e 1115.0i' .033:°1111" longli~ulcle 50.Q9,j"69"'W) and\l~l\(ial!Jk Irock'S, ~a[itud'e 6$ .o7,sa9°N" 11J.n;'ituw 5()',O'!,l ,:S~ 5"W} + g eQ~(ig itr;:aI.

reeks, However. man)' oJ~hioii1C~ilLlck.mlC or fllO~ onh~'lC llC1HlJXlrlCnl'i. <t)l ln Archr."Jl!l g.l?l)cn.~
,,,M;mc

P';~ucla.s~rnITQa,np~lics, pmlk!.d~. :i.i~cc[~ dikc:;; (in and ,iF.li:JlclfOs) ~l>liSlbdmony wOl'ker.; LO ronct!.lC:c ~h<ri~ ~phIOHI\:~'lfcnot 11I;'1):L'C8cn,t~d I!l !.he c:_n:li~",,~ Sl"'..!l.~ oi' E,r:nh'~ hi. . . ln!)l (3 j], Th~ or-dI.,,~"'1 P!;!l'~
IIHl['[(ld cx;.Hll~)k I:;; d'lll; 2 :KIS-~l1i Ili(m-)l\.OI"·!lM Dr}Jlgw{iJ,I;£i uphiolil.C c{,mm)I~1'i in! the NOrtJil C.~lin!1 CnLtcnJ (6), which i:; tL d:~plrtl*1. c!arill:n l lC~mlr,elo' 'Ge!!lbi!il!'~~ aM Depaflfii'l~it.

Il1;h". tho

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HB~ arrd (Oll ~re mD~lrltd rrom (10;, 13,. 15).

lI1Iiips

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Looa.li'o:IiIS 11- ;3
1. ~,ill~lJ!liV:1l

mvatory N'el~o:lt {.AEOr4~a!1.d !')epar~el'!l of Ge«ofJ)lu~ ~di?,n«!s; lImj~~rsilY cf C.. peol'c~~lI1; Qrndi!(b~~a~ilMO Sooth R

~f ~O! r!JI!Sc~n(le. Wl1ij\l1!~iW01 B~rg~1iI1 Bergle", NIQll'i'liI~"~Afri~afllrll"! ab-

tdricii" ~G~oFlJmhtJri9~Z!~m:t~11lIm (~FZ)c-Jl[j~sdall1i, Tli!tegfil"eJfIbKg. ~L1!d!a:.rnJ, GemilH)'. ~ri,p.P'S !lftl;'lilllltloo d O~· I)jlHlpirj, Umvm1ty ~f Ca5fiDla, Sao IDi:I!gQI La loiLa\ (JI '9iOgl-02l5., USA. 5twcir:: Cen'll~ler' l&alih E:\U1.U1i'OO i.IDd

Geotoglc.al
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Muse!,un. lumll~,ewsliI1
f( Denr~

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Ik ~~]ll9lit

U.nii'l'!i.'I!ii~' (Iof

tllbl!r:ta.

101 lEal1h and ~ ~dlJl(lnIDlIIl ~I i6G

"'1'0 \~hom oo~~e~p(!lfnl1fen~e illlIOuI_dI be addlre5.'Sie(iJ. IE',rnait "'arilldJlIInes@jgl!o..uibJIO

1704

REPORTS
imrH~<ltiom, ;loon! Ihe carily ~lon icend ~"Ochern i\:<ll ,cvol~~~i(llt r :Eiilr1~~, o 'fh~ ~S!3. ,,·il~JL<f!l,?d i~~ Si[Ji~.~h\lt.'l;.[iilgrn (Dr~n~ l:nul (Fi!jl •. I.A". d~r.r1~~$\In :m;;'uu'[~ I:Jeh -~~ 1.:.1~1 lo.ng In~t12~5 kl1li:iliick. l.Fig. ~B) ~h:.uC()m~~iI1s a Vllnclyor igttoor;s and !'lcdiflnelll!,Il)' ~«k!'O I( NJ" In. En ~(;!i~f<d, ~hcro;;:k~, .If(; t'ilH1U.'!,glyed f<l:l(lT1h::"-' ;II'I{~ !tiI~lajii10~]hosL-d1,0 :ii!ttfilrjitroliL~ fm~i~. and primm) ~i.:nlu~~~ ~me. The Im,i!i liihol~it..';s of Ihe ~SB IJi.~ :tnC'l!lihfi.il1l~l t1lri~,phib.. dJik':>). 1'It\:w.:::abbr,~ arl!j UlIInII'I'iI!JIiCl'l,~oe~loo with In(:t~j)C~iitC!i. 'li1r1~iI1:.. banded iml1, f'l'lfMlllr lim'L~ (:BIF:<:~.m~d felsic rock:. II). new pre~'YCf'IIed aseselnvcs ""idlill the SlIIIDIJuKiin,s. p;1!tit{H~ Oklk.moq .md Amil"iOCl gIlci:;..<:es~ IW'S. IB}.

lirs! OQ11~LdC'rocl. ]Q In" dC'I,~\'!Cd ITom mafic 1"ion::1 (n}~, ~~1jl h~, been rrcilli~fJilP;:~c" as
~~11~ ~ir'll,;'nmry

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s.j~l~ U. J,,'n h ha~ t~1JiIl V!.

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3:700 m~lIion y~llr.;!I.!l.0 fM!l1 04~."V~1.o1e·if:'Ooek SI'l'ilrI\Jd ~soo'lm:u~::; cHn,oC'tiI.gt.."ii. or 3711} 1. iHMa d rrol'l'ii n'iCna.~:i itIiCHI:S!I!~~~ I'~clOi\i~I.~b'<alben~chidcr C me~"!l.abhro Ur1}, 1ilc hmcr dme 'u~,1~,UI1~1based on ~nm!lli;~ COn!iJC~!X! fr0!11 10000dlton lor~hi~ ud)! \,,~
~15) <lind 3777 i 44 MOl j!lom ,~i[Uo\\' lavas :md

gdJhL'r i:lt~S(l" ,~~cn~bl;lgt;"SI"I1:PJ~~m, ,oo~nio-:I:lkc t~!i.1S1 '~Iat nm!l'~I'~\!,c been obdueied wrlhin an "~'1cJ~!iol1,..ry 'WOO~ -3." bin ion }'~"''1!flo;r ~1,go', 011"a ~'~ul~ uf plll!);.':~~'Cl'i:ll]ic-i~e P]'C_l!;)I;~~~ h ! {J R ....,JiofiilClnc d1ItiJ:1.~ ~UfP'b ~1I~dIlb II:J[I i I1ml s,haWil Iilul Ihe ISH. Ibnla~ bt;:,,,;,oel!ll and

il~ 1J.!OO1JlJd ~~d()l15 1 to':; ~F:ig. 1.(,). u~ltion .~~l()\VS \~I·pn.::M:T\'cd.lmmb.~~tic p'1'~Inwfl1lVo1 (.F.~,g.~}. in (l~~;;l \\ith .,,!l~l~ i.!'i'l!l~1J li'KK.~b;'lt... ilu¢rpiU9!'>v hY~I~t)d!l~itlJ of (fig. 2At lOC1illy,. [Il~ p~]~ow~ ,cm~~3r:1l 1~lsic oQC'Ui (~~g. mile

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1!!nygc~J..~ 11111:'OCk~ are hmli1o~~ l

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!l~TIl,~I}' dL;'~bnn].(ld .md I,)O:ll~ail1 O!. d("...~..~ lll,!! i~ SlilJI:iarnHd W Lh~ ~i'tiw~o~tr:lIJa.a:ycril~~.!J1;lotn1~d O!.TIIi (tJti/gtli:i:i:~I~ s;ph,.;rl:CH~)~'R!k:;;~~~ i ddClm13~L:r.ilj, witlrn S:o k} '9O'~111 l!:h(ii'~H~~\I!l. jl!i:~ltdi:Clllf.ir [0 Ih~ clca\ 9~ liaKi. 2:00 t>D' .2}iQ!h, c:.llcl1:-i:oi'l II~O:il~ Ii

\vc~McJ'il]cd 1[f1~luion phmginl£, 7.10)S•.Al locatkm

1. ~ I ,S k nn ~~Ith of loc.'ItiDll .~ (Fi~. I C~ Ihe ~,uencc COI1Sl5ts ornRbtl~:lr ::-.liJbl1"lrnlici diklo!S: 'with i'nlCf'I'I.'illi'l1G 'c\"iIIllmctru'- to d()cimcIC"lHIla~ck ZOI!J;;'$ of kmlicll!llllI-4~iln.'gubf :-;.clj~lS (If 'l'O~" C".mic !ll"~L,'fjrlIIFh::. 2. and !D.undl fi~.S 1I. tJ:!I1d 1~[uny.UJ]agioJ;l,!"dnih': u~t:'UIN IFig, 201 .m\! S,I!IP!JJOlt~llg onl inc rnm~ri<ll t(:SO[VIH. AI1],'lru~'inmlcly 5~)] In ~l.Ir!h!.'r .":C')lU~h loeaien 3. Ihe miN(ld .• u di kJ~~'ok,uu!iL;' ~-qU~llCC' Ch"IlM.~1:S ~!nbi;;tur.~n~ l

axe·crn:nmw, wiU~~~lllte ,1I1l1phibc;li!~ (!1. i 3) ,IlW ~ri~.e,. I 'pm\lidl,~ ~!!~'iil!!li \'oC',ul ~wid~'"i,"C' or :l>U~)~ll<1l1'il~L,! WI/; e..'\a:!'l'Ii~K11 li!lUI1'txT .of ~elim~\l iu 111L;' L~~· 0,11, Imrus. Al1o~hl.;'r '!najor rol1l1l~ncl1~ (...j.fJl'~' of [he ern pOII'l oj' ~IIe \"'~!'crIIl alUl! <'lnhli:l!SB..rmlpfh..~d as, ISB,,, ,'~ UJlil dl.'">C'!11Jcd ,1:0, '~rOOl.!)lchpd~r." ... ... v;O;ri"'~lI:Cd~hi"">1 (} f}~ .md '~ll:phiooJi!~ ~13,. !fl ~'a

lP'i~k1~7I.':-;n'!.ICllln·~

am!

.'tss~jmt-d

IcI~i.nloel!iu1~ili,."l'io

dOWH\\.'illd

inro 0:.. ShL,.',t'll(;'(i ~Jl1,plex i;Umi~l:i,"g of 100"% Iflhulaf od.i,kcs.~F.ig. .lE ;J!td IJg, S2:}. w~~~ch

[00 l1r'ic WCSI Ls ~ifiIm~I.c!liic COJlIncL~,'~~111f~":I~UWlbM) IDld i.lllrnmll~lc sheets, IlttlivklLutll,dikoes r:mg,c in. widlh lrum 2 In 5'0 elliL [)<ike; have' bodl. oIJc·fll!!d '!mas;l~y) lwo~way 1i1'll..~UilInrlcrl chiMed plauiil:r. InilQ!li 11)1. (Ag. :2 E), Cl'IJIsHlllling di kc,..;: ,nrcll!lm observe i (F~g. 11'1. \Vc c..... ~ni!loo :llluunbcr 1:11" a

~C!iDn:;;

roqUCIlCC

\\Ill uch \\'C :bltl.:U.pl1C~rus p;lrt (I r II ~hcelCd."jikc OOHlph \'IJllh an ,cs~im~K~ prededO!1!'!wllio!n!w idl~D (I r >2,00 m, PoClJTOgr,uphic,~ny, cCI1:!mll)aI'Il,,, or !he di k(.~s; .he ooll1ii!S:1. of lin'Bo~mill~ (.. 300 ~Hn, pbgi.oclm'i~. rul:lpJ'Iioolc ,~pl1l11OJllri!amn), .r~m;lb~mil.c \vh]~ !"dl~c SlL1:X)p1rl~licU:~\lulf('fFig. 2i(jt The dflrkgrrell (oO'm· mllnnl~Y schisros.e} ~n-~rgil1;l~ zones, inft.'lrnXl TO rep~ll d~i~~cJfUHifgiil.'> (Fi;g. lE"~ eonsistof dense
{I

:lC'-.I'OfoS <I.

rdikes,

30

1.0

S[Hn .. ide w

,slIbvclilic"dl

~n) ll'k.ll'K):.l1nlillcrnlk zone'S or runphibo1c {Fig. 211.'" These chilled mrru:gitls of t~le di kes Me
1(-]100

!CJiwrn~~y ami rnillo:ojo£,[udly

simi'l~u:!OII1.c nil!r-

WllS orlhe: pUiows. Manti gl1c'is.<;.(S ttIICllmtC'dmi.'i m~mgaljbros CtCCU r as scanered ()illICrol)S, \vithhl an area 0:[ -100 .I}y B)O In hI dIe :5llulilwC'StoC',1;1l 1}.il!11l"rthe ISB. They arc li!nifmm :l!1llrh~l¥:l~hcs CJlm,lciC'ff;l)cc by
~!.lti!l1!:1C'l:-,:llC.<l,!c ~i5CIlIHI!t~I{lu~ .b)"'CI'.~,,~nd I(;'~~~(,':-;

(Jirph.gioch:l~ in~ .. h{lmbk!rl~1iNIWlart:(:·p~<lgim;:I~t;c ~mlui.'!; Oi,g. S.:», :1,epn,"M;.lIlling ~I hl~h~y dt;ft)H1iI1!.i.,;:i:

.ruld n!lL,.~[,;).l'll:{!'iPhos{;t1!;'lh'b.mi('; k'Nl\un:. nl~~l,j(m-

nm.\tloWilhrfl!)::>t r:iiLl<l'II'1'i1phioo:l.~ki:\, which [yp~'CiiI~ b' c{}l~iJuor ~idto~g,iCflII un~ls"::: I m across :;ilD'1kc "Fig. 2. C .<liU1.d Felsic dikes of thC' /\ml.1roq :D),

SJ1.c.is._~cros,'>'ln the m,clagabbxos and dci'illc Ihc

Fiig. 2. (A) w,eU.-prese-,l'\!'el:i IfJillmll ~i3iVi!e.:dllibil~n.!lI d1ii~~ed 11I1,n!J~:n:'i hll:ar,k sellvage!;J snd poc~m Ijir interpi~ItJ,\v hy~~QC!aslil,e (II,~HI).~B)oOC!eilU·b~arijl'l;g plHCii!M5,.Tlte' p/a~e {)r a~ oc~Uli" ,orig~natlY sp'l1ere~. rJlivea UliI~"'~i!fi@' (Ifth'e d~r(lrm~liQflt!at till€! ~OC~~ ha\li~ sl!fi'@redl. W and! ~B,)o!i!ie Irom tl':u:atiM 1. () [J~k.es witi'! intenreiililftg lla,ye~ of 'II'®~caJnk moe:ks M. UrJ~ Nearly lOO!;tJo sm'ele,teti di~e:s ~~th mriilimtr .iI'iflOUnbS ofinrter:· i(j~k@VOl(gllk lIrI~t~r~L Di~s Ic~n oo~raoO'ld atl:l!1[J Slrl'ke for ml:lji~ tha!1l 20 m... The- Ii'iilili'ti1! len~ (ootl'OO1 ~~9hU ,;:I prngiogroClllilt:e ,~lPGt {q. and ([I} al"lefr(lf]] lO'Cat,lcnIl2. (1:) UIOo% slleeted~d~lkf'!Umplek is \willthLl'red~l!It mm'5 ,Ull amll~ib(lle ~chl~t. m~gin<llly ,hilJlel:l m;9~m,(F} Two cJro:5s{lIItlimgl l:a:reirs ijlllS.'I: !be~Q'Wth~ il!:J!mrnllf h~}. (E) i!nd (IF)ilIre h\HIr! kic@tiorl3, UG,) ·~f!d (!I'Ull Ph>l)wmkrQgri:!pt!sof the c~Pltf~l pl'llnt:~~.t&le dike {Gts.h.owing r,~lkl su!mpl1iLit:lextlilre (planle-r,mlarized Hght: il1lltlge),8!rld chiUellii mllr'gill ICilf th~ ,riib (H~, (cr..n~;1;ling p.red~liTi:irlanJtJlyaF 'rilllie·9irailfle!l:lI[lrl1ip'hiib®~e (pta i1ie! lightim:ag~'). ,
!.l'tr~y Aruh~n
fUll."::,o,hhc ~.SLm.!i.Llpmcfii':;;ifl~:;',

ln~

Ihe~SB. have bcci'l Y'<l:ri;.'llb(I~' ill1:ll15'lic!lllllcd w seI"lfII!I'IIlinLtcsand c,~lc-silka~e rocks by rncttloomalic pR'l!ij~:ilI..~ (II). TI'lCfoC'r..'}'Icred m'l.'tN.!llr.!a11.dic m:;:k~,m;: "&.'()I;:i~. nl wi~! 'tI!L;" sh.;;el~'(ldikc rom. pk); I~F!:f;l. Ie, ,lI1d nU;;~l.Wd)1i1tn\ ~1"'~("tillQl~ wilh
i,minttaTI!!jJto;J,
~.tl,UC'i~~Ur'ilf.l

t I~g, 54}. w~lichoomlt mnainly ah:lIIg the ixllmdMics oflhc 'VCSt~l, bell of
ll1CLlhmll1i1 ~c .~

't!,,Jn~ilitll1i!O

Ii<rnTl l'ayt,tt;;d

IiID~r;II]iillfic:

g;lhbfrn;. {i8~

REPORTS

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1111

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mt>fiillD O(!eiillill

t:
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i!lLlIOII,
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"(~I)

1011 l!J:(Wm)

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'03-3.'$, I03~:3.B !J3~3.,22 O:lH3.23 jU~3.:U 03-3.:39 03-3.40' lA2-IIG·O'5 184-116-08 2A-IG,·06 282A·~:G·O'6

fi~~ijW (iO'fie fi~lw cooe Pi~lw COlle' pmw (Qi'\(l< Pi ~low CO'I,(l<
P9~LOYiI ((fn~

pmow (!f:l\e' Pi~l(M1 (OT~ Pililow coroe Pi~low ,w,e Pi~low ClJ:r,~

I
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lA2A·tG ..06 SSHGF06 S(-U&06

68-161·06 HGi{l,6
fHG,.(I6

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fiUlow (iIJii\e DIke DIkE!

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10

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:IJ'j(~IG"Ob :a6~.·CG·O& 16G-U:;.-06

"f'II1IDt'II!

lI='iig. ~. (N Ti-V" '~!11) Z~·~~N, TH!'r, ~f1Jd1!ti'} If -o ,di~(rim,ijl'lla~i:ol'! d~:g:r,gJm~ ), i!Jh1l gel)Wllmfic.;ll d~t~ (C) ( U
e€!f1h.li ~-. QlIitlil'r-. al1dirlill~ r.'1aw,CtOCUlntC ,(h:H'i'iallli$ ~l!IIi,~in~PE!ifltialCICl ctab(jS:;1llt'S~ ,arerV1Clm m U 7, 19), "[he M\\I' glel[)a:he!i'lilkall data of tl1iis ~tl)ldy!'llle ShOWJ1 ill ~~d. n,~ bOfllil'ln,te dQlta !'lreho,f'n (21). llh@T!N' Ji!lios ~fl(Iii) air'@'ella raC'lErliSlic IJIr~h@' fQ~towi ~g: UI t~ 20. '~:;~C!ndarc: 20l05~), MOIRIBI 20 U~ 3 l'TiIi);)m MO:RB i3lnd is~and iW:; iln~ 10 to SO, ba:ck'o'wt: Ibasins. Bon. Do'nrinit@s; Itll~i:5lj!fld'ar![ th;!JI~em:e; cmd' WP H, ~'llittrli 1iiI- plBlte basa ~t.
1 [).

::I.7A-~G"0'6
]jI'I8·~G-06

rrom~'he

Dike Dike Dike Dike Di'ke Dike

lilt': g~ChL'lnisl"y of p,iU'ffi'O'I,w;;) ,~Il(1dil.:.C;s finnm ~ociUiQ'Jl:." I and .3 (fig, IC)" 1l~ \vd1111i thm or~he least iI.~I.CI\..'ti srn:1llpl~ I] f prev 10m smdics I(! 7. 19). hilsliKcn ploned ill diserhninnnt diagmms wilh various cOfJJlbil1!llil~mIS onhe~llui\'c\. I)' imI1l1l~bi!l,c cleilllcm:l. n, \I., o. y, and z- I[ i J, The new ,goocinl!ll:licrll dna {labk: S~ and SOM} de;Jnol1:Ltrnte~llrnocmlnh: island me and mid~ ocrolll:iclge basalt (~IIO~Bl am~ni~ics (F'ig. J). <Ill p.rc'\I'iot~ly 'c:om::I~ldcd ~/tl). ,Ftu1ht:l111ore, the slmilm.rhie:> Jn~hc oorlc:elllmdon,<;: ()I' lnc:ol~I'lJdblC' C'!1.".InC1l~t~(ri. V. 2:,:. and Yl 'Imllhch mth;;1S (ZI'N} S~mpl!;lJ,y S~~Jim'~1 '~h~t Ih~ pg.~~gW ~"~\'a:ii ,~u~dmhe dik~~ ,In;.'
(;Q~I,'U"iliC'. ~'>\Ippmmip!~ {lllll[

f.}x)l~cn

i~mHp(: dllill

r1tl111l

100001.it"'Ui~~

and .3
C'1l-

(fiJ:l, B::'} ~how~hm t~K: pHllows flre more

~Md

(~'!'~

v.nion:>. ~;~lding Ih('iir "'-p'ni,ul <;IJld l~P~tptlP,11 rel."ItilOlli>hil):;"' .. 11 hns !J,(iCUi (kJnonP::>IJ~~.,;d lh~l Iln;;U1lC1!lbmmJiL~ or Uu: ('cnlil:,u:I! gru:belflt\(!hicfCf 1I~1it Ilre :!il(lodt~mlcil~ sinnnario boa,iniliC:S II i) Iy mt':l;1IlnO~hislITii Ihill. ~1l kes place III ruodem {.I'ig. ~hc presence 0 r bon ~rlitc:> is ilmj)oU'~ sp~dlinlg, riclg.l.'S (SOMl'fLlld \~hoch hills been docememcd ill IllOS~ (It)1ll.oli!le:s of 1'~\l<Incro:mi, Imlt iilllhc evahmtion of~'hclcctolili, cflvil'OfIand ,ProIIC'Jurok: .ages (14,. J.n This ~indillS i::1, ment, bec;nlS('II1l~ arc :gene-mil}, associated nLji.d·im;I~1 sion sludk;-;;:; ,('1,11 with nm(ukm inrM·[l.occflnic island ares iJnldi ~l1e a] 00 ctlll.!ih;Wlftw,ith 31llygdiBics iUl lhe~'SB ll'iHow In~."'Cd.[J.,> that ~]llougIH 10 be related to proto-arc ancll:mck~;Jl'>c imlirntc' 3~te.<lltiml durin£, carl!)' sc.'l'-~oo;(, like '~rrc:<u.Jins j2'o 22). Thill m,ag;nl<1Jti:c progresh:ydrolltcfl1ll~1 mCI~i:li~lOfJ~h~~nn 116), s tol~ :;;l!Igge5~S: 1.lml.0I:lhioHtcs arc good~emkalll.y We pITW [:de :l~~rcc b1.l!s~Ii ues oFc,;ldcl1cc ~Gr m il~"IG:og~~~us and, Illa~ IhcUif Iccb:m ic (rvl.'1<llJIlion iCmllpl~]" as ,I "~$Iige of An::hC,~l rna)' Ili!\'C' invoh,¢{j i11i~i;jl !'C'a-Hoor ~l1remling, ill] lsua ophio:J ~I!:! !;iUp:r<~ ~~uLx1\~'lip!~-::~!1~ Qw.in~{; on:tl~l, fi~. [itf: fullmvcd 'b'Y j;i~!bl:,b;:lion i~ili,~liDf! ;uw n,~!,) or Pl'it:m; cpiM1d~ nrart: spWlilrng a!fid b;liil'i (lpL;p1ing ~l~t~ld-dikl;: cm!'i~~L;'x and ~tlgcnL;ti~ puHow I"""IS, ~cl]u~n~ the i.iJirc:r-em :St:d~(.IJl ora !:Il"~'i":irn'ia ..nllll ,p0,2t).

[ichod ~IlU U'i3l1[he dik.e.'ilTabk: I 3J'idS{)M)" l\Jihoughr the S.I!I(} \fal lies or !hc oolira~ pan of the IlHllows m.nlge ~~~'WCCI:i (,.5mldl 9.9 tflvii!mge. 1,2:~.Ihc d ike s3m~"" , mU:fll,')wer: U',fU,2C bcu, ecn 3 5. 7.md 6,01.)' (a:~g.c. (~.)). These rocks do mot Wi\.~ol'd puimu}' nmgnmlic o:-;yge~~iso~01}Cw ~ues bUI ma)1I't'1JOI:C1, 3Itc:rrati:on b;- () 10 j:! ol"l,lu: 0.1180' value of scaiWmer at a sJlre.1.dlng rid,ge. The pllh:}\~1Sarc more cnl,dlc,1 li~lI~O boCl.llll:iiC Ihey allCtccl al kYI,\~r tcnlrcr.!tu~ Ih:m d~d~hc dikes, and (~~"i.)'~! :b;ol~P;;:: 1'!<lCli~ma1:i(Ul decreases tin ~n,l!ilni~ud~ \.... ill(;'~'U.\liialg wmJ~i[,;n'lUI'C (J;~. i~h ('olk"l;Jtiv",~y,. Lh~~im,I,~r~[1 l!1'~ rdic :s.uboph~t.i:t;, ~i;.X.'~WlU:~ il]~hL: l\h~clC!J dih':S ,II\; (:(mi'iisG~l~l idi w dw !iCll~.v'dt{!'.rIr,!)i).k kl~t~il~.iO:II dutillg Ol~I·noor

kDI.-a .r'lilhi:oh~c. Th~:-li!cc~L;'d·dikl,;' IconlJlh;:;;: pmvii;h."':So cmrl~ni.l.1g S1.l'lJIiCl~ralc\'ud'e·nce .of Il{~r~· m!lUi~. C"i: r¢Mil~ion hy (]~k~ ~nj~~C:lilon,JR ~I sptL""'tidliirl!!\ ridge (SUM I. S{'''COl:"u:i,W'f: n:jccl.ehe SOt1:lfUOO, of dike injc.ctioll1l abeve 3. plll.lllC head in a uon-plate
k'(.WIl it; ~il'i'lli mnli~em., ru"C

£iw'!iTI~he OCCfU1ic islailld and MORE: gooch'Cllmk.d i(;.h8imclcrfu>~ics 01"

uhc Ili~loi\'" lavas ,and dikes repofl.cd here. lo£,cthC:Ii Wittl the data Irom (.19). F~rCi1cJ:~ ooninitic the a;1,"liillily ofd1C eemral galfirellschk'Ii:.I'" 17) is simnay 1.0111m or PJl.'tllcroll"\')ic~lI~jiil ~Lbduct&on~'IDm: {)p'JliGHiICS wllh a 11fOtl:a\:1cd lectmmll1lagilll![l~ic C\Joll!~ion~list(D:I'Y{1~1, .:.n Third, 'l~le o""ygc~~ ISOtOf:!C C<.lltlllQstIJ{I!n::l of Ibe IlmO\vl"V<l~' ~nCl dikes
!)Inld ~lC'if P;:11;{Ii:"~lidctc.."'llll:~S ,I"" cO:!1lp,~~ib~~ with~ ~""I·flOt;_lr ~~yd!t'!IJ~~·.mml H1!.;1111]lH~phi",1'!I <It ,I l;fln..lJding ri~~~. lahul,Igh the ~!"LiP1hi~dULry of ' A lJ:m::;l;I'Q~k:> :ii5 nOI y~[ .~~llm!CiCnllly well kl1(lJwn

n,

'Iop-cn:nit !J dclt'lil~ed reeonstruetien cfl:i:lc .~:>lm ophiollte COl'np~cA,.we contend Ihall'l:!e :~S:B ]W\.~I,"\{CS vestiges of [;1rth 's eldest ophion~eajfld oceanic erua, Th~s :implies that sc~~Hoo; spxcadliug <tIld:'llIbduciiQup~~ ofP'J.mm:m.oi.oiL.':>-!~k~ piJ.ai.c ionks. were (lfpcm~:il\g -3.::; bHlli.(m ye~ ~
:!Igo. as Im)li!t:)[;;cd by Komirya (If flf. !t 111.
IRi@]lm~,I'Il~@,s, ,<lllId N(l~@<s,

1..

1~~u.'.GNL Soc. . IW. Sp«.. ,r;Dp. 37J (2001~. P. 2,~9. ;Z. Y,. (lH~!(, Gi;\!;ll. Am\. ,SPff.; J!?p. 313 ~(I(jill), p. ::ll, ~. I\l ]I. IBidkll1, ~. G. Ui51i!e1',A. lilanlin,. J. ~ li.'o!Z,121 fl9!l"'}. It w. 18>Ifbmlll~j]i ~ ~,: Am iMuy lil 4· ~2003l'.

1. A.

s:~

1706

REPORTS
'5;, R.]. St~H!.rk!ClWfjr 3]" S,S7 (~1S.1. , 6. r, .M. t\Ul~. Ht u, III Ill. ili~DIler. ,itnce' :!!:9,~, lHl1 5

l6. M. 8!:11et et aL. fmllr Pl'm~r;Sci, 1.eN. :2::t~, 417 t2:oo~)"

11.1_ A. Pl)I<!\;, Il ~~" HiJllII'I~rlrn. I'Iol T,.II!:~jl'lg,

7., iii..~~i~ G,,2Il~Q, 0, llimllg\ Stffrlrl 2!iMi, 923a (l~0ZI, S, ToM. ~5~. Ed •• PiIl't"Qmli\!igr;!' fJpl1ifJ~'~ ,~I'iI"Be!QlefJ
Rod::I (Ebl!'d~r" Arn5ttlri.lam, :l!lliI)4), 9.. M. ]1,. de: Wit. ijm ,~W,pmt¥l'fJ lIi Precl!II"f/mm mJOllIY, ~l 13.1L ~ K~~k)l, IE~, ~Elj;e"gr,.L~~n,. :f~4). 'Pifi. 599-.1\ l-!l1,. 10, A. P. tMmiJll. Glt~l'l:r,OOi (if.rll Sr..m, BlihC'lin 154, ~]~8!tJ.). U M. I. !!a;!!19', H. ~ ~05e. 1.'1<. Brid:!J!'Nitef; Ill. '~. T~O!!i~e'fI. liliiIaiOgj.' 24, ~~ U.99.6J. 12', To ~mf~ en!"I:. J. ~i. ~I[. 515 n'9~'iI), n, ,~ Ul'l1f'$, Ptffitmtl~n Kgs. l!Ol5!, 'Jl!ll UOQD. .s:. 14, A. Ii'. I:'M.I'I1a11, v; C. lI!!~nl!u .. C. it. L. I'I'1mnCI .M. i" I!ci~im!i, ,(1jMl., Gerlt. :1.41. 271. '(19971. l5i, f. ~rn!JI.. Sr:It!I'IU'~;8~ 6],!II {It'.l99)"

~ooU"

1lS. It ~.ffMIl'Ian. \1'. II, M{(lreg~r" c it L iFriellld. V',.C. B~~ P. 1)\ ~~"', ~"Ii'l~ Rf!5. 7~> 1 'l?~6.t :1l9. It ~la1~ A, W. &lafnnmo ,1"rffJl'lIliJrnl:l~i!!". jI,:!60. 1.97
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z:ja. (~D6;U.

,Ch~.

27" COOlfl'l~~b II:rt Y. Dil~.

~iIIQ~ ~~~r'$

,(i1Qt 11:14,

lB. RWi~~" r.I.1'Ik LaU!)'tI~n.~lldl~~ t.m!il~dnui! ~'IiI!.!Wi~~ Fill~lidl !:UlIIPDi'lII'o'aS prC'l'iI&i!d Iitj' 'ih~ I!jI:m~h~i'lIi:M~ r,~ C'mmal, Ih~ Gedogil".ill Museuml or (qJf!nbag~ml Iile GfZf'Qudl;ilT1, th~~Ii!\f.[o~ laund<llilitll., ~ mill"~' N;,th'I!l..t~ S!:iII!ii1c.t!!, ~nd bSlim~,~mg, ~rd,eoondl. af '~rI~,

O"i~k,

211.

A.

<!I,3 (ron~, I, Crantcrd.. EdL,8ooi~f,I,f"J:~ljnwiJl

M,

J- r; Fi_~.~.

G!'tl~ ~,

~~

5p~. ~bl:
~OIldCl~,

l1!i'5 is

:~s:m 'OO!'l~~J;JUI~i
CIilII~I'Ii!

1110'.

31.

5~p1Po:rtingl
:500 l~~ :figl;. 51. to
f.i!bl!;:'~'lI.
$4i

Malilir.i a I.

H)m1lfl.

w.wr.li:iieot:tmiuJ'~~'1li!'(Xlflum~rl.lll!']:1~~e:19.!'l!'04.ltlCl

19B9t

22'. Ill. B. pt!ll~n, 11'1. rum~. J. GNd)1!. ::IL~, iS3 Il99W'. 23. lit Mue!l"ie!!b<idi1:!, fhMl, Gffij, :l!i415.i!i63 (19 9at 24_ I!J S. S!i1i.tS. !fI. ,PO. I~,1t"j. 1 GeQ~fl. ..file$. 91, liJ"l3
(li9'!!'2~, 25. (, HdmiJelll ~. M!i!elienl:!affit .Sri=t: :259 .r l:rn !199i». 26. ]l1L :It f~Um!!~ .~lN1tlll .Res.. :112!~, 2l~' ~20[ln

ede.r~~~
:21IOei:~JIlber 2006,; ~~ep!IJed ? f'ebru~ 2007

"t.

1O.11!26isci~m.c1!.lH9li10

IBotlolml-Upi D'Bilermination of' Air-S'08 IMI,ome'Rlt:lllm Ex,c,lhange llnder 8, IMla,ior 'lrepieal Cy'c III ne o
As a re,sult ef nn(iI'",e-.asiflglflrl~qI.H!'iiI!CYillml ~nt<f1nsilY ,(!If t.mpkal (~don!1,'!>, illiil ,mc:~ra~ero.rre(.a.$til1l!!l1 'of' qlcLolll'e evoiLJ'li)[m .md OCtlClnJ ,e'spo'n~e is bemllll~l'lg fv<en mo:r'e impartanl te redu(,e thr'e- at~ to ~:i¥e"l i3indl properlyinl mB~t:al regi)[jns. 10 ~llfI.p.rove pnli11:(li£ll1l5, iHiCl.llmte ,e'llll1lluiJltiofl ef the ,'Jlrr":l],e-a lIIl@mentum exchange is rr!1ql.Ulired.i511'lg CII,unmt lJIo:5ervati@i1S, rec;onmoo iIlLJrin$l .iii lIIiIilj£lrtr@1l1cilll l q'clQne, \lie ha'llle e~'Ilim(jN~Qj this llIIomentL!lm~rllnsJer [rllm tlile: 'DlI:e,'!!'1 ~itlle' of the l!ir~s,eill iIlterro'l'O!l' • .8ill!di 'W~ di~uJIi'S, it 11111erms or tll~ dr o'l'lJ (lle~Widel1t~ Fo r w~l1ds, betwe,;:.n 2,0 am:J!4S meters t ~il\r sec:~nd" thn~ ,cQ~rti'i~nti~~ti;:J~ly ]!i'lc;re"se~.c!1!'!d ptolil..-s at wll'ld:s Qr a'bQl!t 32 met.e~· per
:Sioc(llr'!d

~hr h; <J!11!.o~h~~ric OO~!!d.u)"Ii.~)w ("~op..down·" c.m ,;dso be dO~le rmm fhe O1.i1l<!f •side {I r the i!l,!erlhC\..~ tvith r~1I'W.1!:tcr'-OO,h,I!!1:lIl tH.'C'lln Cl!4TCnl
iJb!i~~f\'<ui.Q'~. Thi~ m~[lmd diA~clb' I;'i\)i(cp:rnvnes dctcnnimllic:m). E~!im;il~kln

or

thi!>I.~"lchtlll£C

mh~ :!!!.©i:!l
currems

~UI

lh~~

ij!1i~~ir'Jl\t'('it!'.;li};~\d, OIl

OeL~Wl

,ru~d prov.u(l("$ ~ \'t:t)'~li!lblc and aeeuratc direct dCD.crnli.n~mtilon or the air-sea

IrlOl'UftC"IitUlnl.mnlii

fe-.r under

~lJ!:h winds

,[3).

1.lo'wcvc; .. lhis~YI)Co:f ·'bouom.llp'· clC'!CiI'll"i il'&lio~! imposes Ihe ,iJlmo::;;t impossil:ak- requireInents of (k:-p'!O'yh,g hlMn.lnlcnl'~ in the ocean

Ib'l\lt~e d~(ir'~aS"i r Jilgl.

d~roc!ly underthe highl~ ~ t1lt'\jol:'l~:nic[JJ1 cyclone itl ~I nm.lcms survive the ca.ulcd by the eyclene. Om i 5, S~Dwmbcr
Ulimrur:~I~C'

m1rredic!.:Blblcpath 1.)1" and then hav:il1g the

en.o.I'1.noIJS forces. gen2(](]4. 4

Ihi: ccmer of
~.n;I'!.glh)

Ivan

~.j)ll c;I~~gory

:p;;~~d ...

of the exchange hence is required for corr,ccDly ro.reca.')t~llg su:mn track !lind ilnl!i!~lsi~ ~y. a.s w<CI i as for .acCLI r..lldy 1J1.'oC"doc ITn,gstonn surges. <t;)CC3Ul CllrrCI~IS. and 'waves. <lna ror mlUkun~ hurricane ns k assessarerss, P>!iI!'1.Ic~II:l.rlJi because [h~ fl.\CqrLlcm:y and in,lcm.ity oflropk:al cyclones arc n::llOrtcdly incl:'C3sJ:ng (n. Convent ilonalmlc~hods of dotermin ing th~"~~I,'-s.e,>1 UllomC'11UWll t~~!'Lt~ aN' lrom the al.rllospi1c:ri:c fer sKi.; !~~C inl:qr.tllCor:'Ind ;~fI1.: ba.. r.ld on m~<I~ ~

lion

hC.!~if"M~fI 1lIl ... ml'liC.ill[Un~ ~dm!l.~.;' ~U1'd.-.cr.'[U n'Oplcfl~ CyC~o.ll(; den:.'n1liltl'h."S I&~eoct:m~~!t I:ieSpOl'L'>'C to.

~nswind s, An

flOCumn!

eaima-

2:0 m s-l .• "HIC' 1£;chniqlli(:SL'1IltJl,loy(;d ~~"li!l:mlCllilc ~11j[\crliwe m.mtlril1hHl'Il c: x rll1lE1gc 3&fL wind !lIre:;:;; .• 'I:~ giV~1~ ~'\I'

~ilx CIi!II"I'('Il~ and \,.... lv(l~idc ,~i"'!ile ~.lOr)J'irlg~ em !IlL; Outer C'fiJ!nim:l'lt!d t;hc~f illl [he IIiOnhC:fil.lcm (.iulfofMcx.iec) ,t9' 12l IF]~, It Tille iI:lSI.n.Ul1'clil.m ion s\u:vi;v(',d and ;iltCru>1II od r n~eorll)' ~hJ~ ocean current \ dadl:y profiles diroclly under t~'l.ceyclose. 'The resulting ami!
d;II'i:'(1~)'OV~'T

SCI allows ,direct '"bouom-liIIJ'

detcl,ll!lil'li;1[ioUDor

or

where PIlir is Ihe air densily. CD Is the drag 1l0C'fti1cie~~I, I\. I is the vetooilY luagniutdc of f.llli:.! tll1:." ~nd ~.1a 1,'C~rcllCC heilg~lI.~!1!isualljj' Uill1). w Il:n open ocean COUl~]~lions, as the wind becomes Slmlllg.cr the rnn~ or Surr:1C"C \'\.'<lVCS, wh ich uavel als]tn,\;'cr :5~cd~ 1iI1oJllllhc' wh~dI, wiol;;l!Js .. !\4;;coJ:dJngly. CD incf!i.,~'i'lC'-"all, W~!1.d~W(d
in~;TClI~

~~'lIrcni!,1.;'l:It~ ofwi!ld pmm~ 'V~I."lf lh~ (1~C<1L11 5~lr~ ~:~II;;(.; of wind Illre u~O!;m';l,i ,tlll'h~~I;L11~~dlOd'i, or h(~wcv('r, c.;:ull'lntw~;1J"l B;luably fo~.·th~ t'!:~lrcnlJl:; ~Cligh"wirHj <eo:m:lilion>; fi n'tf~Or I.ropical cy~

tn. ni:s

!);lu~nJ t'!~~ v~~rj,~bilil.y fu CD

is t,1tlllnmJl~1:y

~~~cl W'~\'{::" :i!UI·~ .• o:tJld c.un;ul!'I" iu!

~)r

C"l£lm:', bocii,m":c
OQ,;i1i1

VI'

Lnd

m(;j}M~a:cl'b1lellll>

nCl:'"lr the:

Interfaoc arc im:v it,lllly OOllhlJnimllOO ~;ro.~TIl ~n~cI!lSiC breakil'lg-w;]vC actR"iti.c:s and 5Cfl

~prn)'

1v[o..'>i <l.vail~lbk: rick! m(;J;;LH''Cmei~i~ of SUI:· lacc~~~~ are for wim1 spced~'Jngjng fmn.4 t~·) iUi.'>ti miC<li.tiom,.,
laiiwrOiikl!iY. 5t~n!s Space Cellb::t, MS

O'~

.Naval ~llh
'"v!!! ~~nq

395<29-5004, U)A.
~~f!~p(!lll~il~e ~lI!)~dj b'l' ~g:g'~~~"~'!'I1I~it ~a.j~msZ@.nd51l(JI~"'YJrnit tf!'e5eril, Old!ireS5: [x:ron Mobil Up:slreaJl'!, R~~:arch Corn..

IP\1ln~.Ho~loo,

'1!X 7"02'l~019!, U5A.

is not gC(>Simphical:!y billllml.ccd antI ,I nm.ch. wcake:r:' baroc:1:ini.c rcs«KHl.SC. ,,[,hLl~, the d~~ci.ly Rcccnl],y, lill'l'll.tccliulC'lemulogicJI £1lJSCli'\'a- [g;~o..">(1~esl}om;e' Oil Ihe crmi illiCntall shell' ~m~ Id bcwd I ae..<':cl'ibcd 10 I,hc nrs~ orde!' 'by the l in!l:'ar ti:OA"ls.~heO:I:Y,mw e;.,;;perilllellls I.~ R~ susg-Sl <I. I,im;::"dc:pcnd:cnl.lk.~I.h"i:~llegrn.lI);:d i~('Iriror~i;cl 11!l{F roo~!cl,ioll ~fl .he Or..ll!l l.'"OCnicicull Ol.t h~gh wind ~(;I;!d~ gt."ncmlly gi!'Ca~CI' d'I,l!l )01 m s -I • fU~thl.~ mcnllWrI !.'qwa~o:ns. &:;d.~n.£ 'ill<!i},'l'i~ 01' ollr1:enl dlb.rt..... tt!, ~wll u,~tL; 'uh~ ,;;Iiir~hct!nl[]!l~c;utuLu (::;';"" vdfl(:~IY~I~~\I!l!l11iCl~tS. .~~~~ ill !he IWr!hl;Ol!'l~,", UITj.l,·~Il~~'!,'~I~ cll<'lllW; ha\'~ 1iit;,"4;11 IlU;~~ l~ilU~gll nU;;~lir'nl~git.ll.l ,;;m [iulftlrMc:~".~cu 'IF Ii£!" 1.1under [11-1 It'lte:ll'IUr,eni'it''I'iI:;;, (lr l.hrori~ :lnd n:lt:MJd:i:,dcvdt.!flol;d .. ~ i:il~rpo:fll'>, .~!!i~ I lli<.SlIi:i1plirn'i. ~pt.."CI.:H~I,ylor lhe' a]!ciii!!ll.

ti:olli 11llfn,.;rLt"lI ~111[l(kfu.;. wlTliiCh iIlO~J.·pnmtti I.!. mon;otoniic inl1.'"!r(,:llt'iC oljt~~.(; dmg {X){;f1:i(i:cl~D wirlh WiDl.d ~'Cd even lIJldcr (;"xnreruacwind! oo.m:lilwi.DlS. Sli.ch ;;m, aSstmlp1.klifl mil.)' rc"S'Ll!!. ~i1I al1, ovcr· c:;~.mtmniOIl ohhL; 1I~j['"~t\ 1l1.~1~,"K·tU!,lnt ir~IJl.~Ii:;T aud Irn~tic! illitlli:1l:iit.ti!.: moddprucl£ctioi.15 (]It, IlC'Ii'lJrup;s even '\I~.ri:S.c,~nOO'cII hnnlinig usi:rlg bad ph~iClli.l

Iitc at r-sea Hl.on.ll!nlmn c....cimuage LInder C'.'\~roJn'C wil:ldOOltditiom., E;:':,Irr\CI1fQC \Y~nlcl of a uopical C')I'CIO-llC s cl ileil, II ~;;;UJlf-:>tl."lge I'esponse when 1~i1.'>:l.!i o\~C'r O~311j n£: W3~CI5 l'?, IJ), The' I1rsuhrec ~t.1.gc~, mekc up dle 'i"o~d, M3.,!;;C"· I\C'5fIOJl..~e. whereas the ;r{lur!:i~:t1t:.1gc is IBN ··r:cll'~'J~iO~1iii~n!!ll:':" ver ~h~ ~pen ~l!li""Hl, O Dhi; fOfC\i'd :\lJiil~ !:1.~::!rn"k.W i~l'lrilu,lfuly b;11!O!;~il~ic (dcpl:~"dcpC"lIld(,.~n~. \V ~~h \'i'i;,~lk bmlllr(J!~.~it: (d~pllh"im:l.~J]Clrl~.h;nl} I"{_~;I!)I]M:; (,"OJi.~i~lin!!l 11 ~n;}ugh in S(';(I, surlltcc'~l~ighl flud Ilfl fll>~OCliil.ad: gca$1ror)l:lh;c~ rI"CilH tibi),I, ,Ire SCI, !JIP ilI.~m,O$l ills'l.'Qn~ly tlJ).(}vcr the cOilllincl1tal shcU: h'L1Wl..·v~'T.ihC'fOit;:(-dI\'.:;lj)Or~:,;(; i~II;!Ililitedi.If~f:!.."I~L{9l It

,I

ur

COII:lidSi5 ur:It

:.i:ro~l~

b.ltonrorllic

cO~'1l!P0IlI~I)H[ital,

REPORTS
IF!lg. l" R~~!!!!t
,jJ

Bathymetry (i1:11 rn~~rs) ls ~hilllwnfor tliil!' IJIIl,nth!"" ,e-aslennG'lIlli or Mexico.

Aoo'l!$'tic l!l)opp;er ClUm!'!ilt pro filter U[lCP} m{lorilill~ are densl-el:llb)f 1111rk dots .. (liU kI Mel). lmile' yen~w ~ri.:rn;g1e 'Is, Bl!!oy 4,2040, Nati~ll~~~ Buoy (E!IIIi-

na ililslIlum~flt

mil!)'

IDGlitiolils.

~-

~. H~rriq:~e !Vi!l,'S patlii isimikilled byltiil!daShed r!l2d liRe", T\Wifl5erts in tfuJe

Ie!'XPI\! ndsd

II'lH'l!l aN!'

oil

l(iolorlm:l il'lhart'lliinge IClf llflulmrkane ,~1i.an L'l~evn by tile N.ational O::el1l11k .a nd lijrnospne'lk Ad'mrin;-

is.tralij)i1':s Ge()s.taJtiCi~fY rQ~ro1~oo\1lllEnviroomi!llr tal Sa~lt~lF12 ;saiteUh'e

~~liMt@r IvI:'I a nc:ia IB~rny M lIJ1Ioming oont.i!iniliL9Ilhe
AD{P'Clllrrreiilt rrn;eitl'r (up-

per lr1gi1lt).,

Fiig. ,2:. DiFiIgmerli~e'Jll Jl;Ir $ev~rr!lL ~i~:nt ~~d\· HCJlri.ifiJts. Co j~ sh(JW!'i as a rUl1IdiiDfI Glfllhi! Wftlds.rp~dl at :1'0 m (W:1Il) :for several lcim@rrf!rlt ros1sL:!T'l1l! ooe<lIi· Idenls r (open I:I~B;tkcirdes - r := 0.00[1 [m$-1; g~een lri',mgles -,,~ O,,OSQ5t:m :("1; b~1JI! amrisb - ,::
1()!,l!)11:1f11 $-1).

;i~;-li 4r'---'__---"--_"'--__'_--_'_F;;=~~::.i9f

('<lTilIie Ilmh for si III !hu.' !lC',M3011, [],O\\I(:\ cr. m~my s

iJliii!Il!.[I[MI'DJIj

.' I.

t[Ji

.;, i'.M~-,~~"""
.' ~,

r.Q:,~_~ ,.MII'1I·;m1i1

(yclom;:s ov~r~hc

1110\"1.: :;Jm,\ I), !Jndsll.'<Ildu~y cn(J'LI.~h s tm~r forslI.dl ml "Pl)l'i)xin~'i6:OTl Hl

,.Q.05!I5lcmr
'.~M_i1'1

lmM. 11]~flif-!iCiiliJIrom(~IlWllll trnlls.lh. cxpn:..~!>{,-d illl temtli ,of 1[~. is. Q;ljm~too fi"oM.l Eq. 7. IIIsing cur OCl..1fl1l, current ohscr\r'at~oms, ln Htmoopi'lcric stlidilS. nll!s CJoIChmlgJ::is il)OI'1lI'l].ml~Y ctr5C~l.S~OO in c.an be <llso delil11;)di 3S 'T511= lil.nirC til \ IWK. where w~ ~s. the alang-shelf wind vdocil}, (101llpOnclla. Thas, for cOlIlpalliisolll wilh olhr:1:" Mudies.we can d:ci!:~tnim: Cp, IJ)! in~~eI1inlg ihe ~bJrmYI;)j Ii)!; ..... :iII1.10 Eq, 2.
I[ ViP:) !C'UiI.l'15

ofnhc

dl<L!l cocm,cic.rIi

KI)),

\Vitld

Silft'SS

The~Hd ~if1e5
(D

rept'e!ie'flt q!llaiiir:ati[ r[lIIrws

J'id!ed t()1 the evahilat,~ :~PIlraooly f()~ each r.

GI.Jl----r--~I)

....... ....... ----..,.....---,.--~ oliO!

I~O' !it ~·'1~

....... -_....iI.
!li(J1

4!1

W111!l!1~~

~'bc~ ,~1'i!1iln,cntLlfn [more discm,sKm concerning.

ih~ ;j~[!.Jlg.~JlL~lI· Ilr~(li:n(·rl~Llllfi babllre i~in given by

{F

U")1

the sea noo,c, and ~

S,liOC1>=-i.

is II~C'a:loflg ...... wind belf

from. cvalumion o.U·Eq .. J inF~g$. 2, and J. The drn~ eoetfleleru was. derived for severn I d~rlere~1 Y~lrLles of the reslstance oor;;I1i,ciem (,-, rnltgi:lllgbc~w(en. RC:;lIIItSproJUij;od;
3~;'C shown

=..2:!.. pi!

rU

(lOmpo.l1cl'IL~, II J~ UIt'\...~[h;,r , dll";p'il't r lil'! ~~C't'li.nsUiIUi:'(~ili.~;jl'i~cl1rJ~flidt'b11 ~~I

C'rOt\ll-~~~~ r,'docil>'

here p isa re tereuee d~'flsil:y( 1025 kg I'Il-Jl,J is til!;. Coriol is p:u·.~mcted0.'71 ~ m-"'!< -I). U and ir ~ Ihe dcp'h-i~lq;(;;;!J~t;>d ;:l!.~~l!]g~.~lh;II.' ,~I1.'d
~'\I

Tlsc siullpJIHicd dYl1luniCl> g,i\'l.:ill by Eq, 1 miliy OJ~'I ern s:-~ ;;mQ 0.1 em s-]. '11C9C: '!>ll-ucs of r noD. hold under nU Qr.l:IldiinOl~,>.Fo:riIJlSlnnCC, ~nih~s, are :;.hnil>1rto ~~C ~!fIC(! in Qlhcr smdies ;inv~ case, (he mmncfllllirn bnlm~cc brc<:"lks owlJl once d lig.·llin~ shd r Cli.1rr:clrD.s l i 5-/.'!~ ~igll:l'C .1 ,cl iil\p~a,.:s e.lilmI~es of CD [or m;Lnimurn, (0.00 I em $,-1). the ,cy..;: I mlC sl.oli1l:1 has passed over I ~u:: oorinss. ( m ml:.JU~ {O:O,5.iOSem s-II. and nm x iJuum (OJ em :rl, and Ihe~",'buJ vceiers mrtd~)' rotillLe l,f.lIfr:'" \Vhen ",,,Iu.cs of I" L~ ill Eq. 3, wherea.'l Fig. :; shows ~his Imralcl1.'l.. the wak'ir ~1o~lurlln c,alllno~ respond ~\dl:.'l rot r = O.H2 em ~-I,that is, dll:res,,,",!ancC' 0.. ](1d,.m~!: di~'l;lCli:t'lll as ""pid~y ,115, the wind, mid ~dHdL;:~1 'tll<lfl \\'<It-. d~tLClI11~inr;d f~11I1 OhSlL;Ml~hq>t;dyrmm;c,~ ~o lon!1l(;lf ]:!OM.AddiIUOH.1.Uy, the lion_... ror ~]L> mII1h\."ii~I.crn S:~df of ~hl..' GWlU' of ,nt":!~K"J1hJrlll. b;:JItiil~c' 11m)' rim hokl 101' u l<[pi.d~l)' .M"'.Ki£o (/5t l11~n:.'iS!I,IIL~.~:;;p;;:t"i!in~ b Ihc ,,,.illld li'i'ili'ilvil'ig IDIX),piclI.1 e1,dmii: ur :for :I C>'C~l)liCWdlh 311

1708

REPORTS
Fiig.3. (lr,a9oo~!kie!l!l>!liS .1I!hmrnall of moo speed. CD is:sIi!iOWtl for' ai'll ,~b~f'U'a~;~ii1· lb\1sed r'E!Silstanl:@ WenD@n!. r ::: {Ji.o.2 {Jill s-l;,Ti1e ,ffi'd open df(~ are the ,~1· lIilte,d G, 'from nil! wrrrel'l!l
,1Ind \ll1mdl O~l'I\\aD!i)I!!s. the S(JUd lied! tine' '1$ a fitib:! ,,!~dratk; ~~IlIV~W ·UiJ!.l· D C
'II~~ ,.,-j,- __ ---'-'-- __ ---'-"- __ ---'-"- __

--+

~. M. III .~1II, 27'!l QQ03~,

P• .1" Vid'lla)'. 11'. A. ~!lif!l1ctil.~(J~

4:22:,

S. E,

I)"

fnmmdel et Ill. f,Gi!\"P~-Y& Rfs. :l.lIJ, CG8,0'1J

••

1lO..1G;!:9/.200SK0!!l304,a, QOO&l\ 1], I,]. W:looIl. I,.'Gimi$;, T. Iflaf~.jI,ilriliW$_ 7. l. A 'T. ~. A. 0., lel'lltlmi~, J. G~,
I _I

S~. ,ilL

zn~,f.200!l~,

k~'" 1~'1, Cl!l~Ql2.o!!

r~

;tl

A - -~- ,,~.~~~.:::,~'~.; ~ ..~ ...,..-"." ...""".-

B. K.. ,Eln~,nueLJ. Ai!rl:I'a", Sri, ~Itl. 1120 ~:i!IOOB). 1 9. O. A. MiKIltLt W.], FI1Il!!1tl~. E, lir~~l. W. W~ I 11).

DD.lI)Z9o1'2005.IC,Olll~:hU UOO6l\,

,~~timatil$, ~nd the fled d"sIleti Li.n;~sa~ ~fl'9S~~

'10
-I

locmlilrlenu:.e liflil~t:!i, 'hJr~'hi!l 'CiuadrailX ~L1D'!ie. Tfule blad: ItitIltecii ~in~ lllpF@Smt 'Wile \'l'ilufO'l'll for (1:), reported in

r~

.'... ,

'.,

~,!!.,U6101OJ,O','WZ,OOOGI.O:230:U l {.2l!iaS}. ]).0'. o. W'. Wtmg. D', A ~~ilcfl@l6'iN" J. lI'~;iJI!'. E. jIlW'H. M, S. "'ll!tib~1l\ 5deJil~ 3'~, 9J~6 >[.2!oost 1:1. W"],, 'fiM!fIi~,IlL J~fO~~ 11). W. \!l,mq.. 0., A Mil!:ni!l~ J. ~ Oceo_~og[ •• in lpre~. 12- w. ].r~9;U~, I~ larg~. m (;l. (~~f!e'O, I!),A, ,~~~1, P•. 1. IiIcg,an, CMI. SMf R~ Z~_ 2~S9 QOO6Jo.
13. 1. F. pna:"T. 1111. S3!Ilf,DfC, G. ,Z. I~nimll.}. Phyf. fX!NmrJ!}f.2I'!. 2H >(i994!t 14. M.!tillii;)t!; ~mil rm~lhod!;~rt mj~tlt~ 'M~~~:rIfng maitllia'l (!if! S':Q'M(li'O'l'!tim~, \1.5. G,.1, Mlt!)hl!Jm, W•.S!uf9e-5,·1,.Plrrs. OtiIQr.lagf; 1.:2:, \l

GfW/rys. ~~,

(6), ~\Ih~rea$ '~~ bbJe dQls reiPl" el5erJt:Co reported in (4)"

i

11'+,

---T'""--"""'---T'""--"""T---T'""--...,..---.j!. ~ ~ ~ ~ a
WTfi!!

s.~~

(iN 1~''fiil.i!;"1~

iloilO)! as a tlltccI19inly fllld Ihe need W e~lcll~ITtea \' cloci[y der,~v:un ve, wh ieh tends iO cnhm~ce nelse. Iluwevcr, they oollsiSllenl.ly show a lIocreasins trend orcjJ tor wind speeds gre.l~cr !han 32111 s-~. thcro"~'C'r Ihrcr.ohold for [I CO'~C,g.o:I)' '~~UIJ'riC'-l!1C Oil I he &!ni r~Simp-.1o.l'l Scale, h i~ a~~o.;uPfl.. wm~h<l!~ thi.!' eu values mtl \~C'ald}i d(..~pci~!d.Cnl (lJl till'! choit~ of dw !'~i~l<mo~ eedfki~~n~. and URi ~.argc:rfor inllCf'l"lrsiflg ~',U~U('S of l". The dr.lg !."QCmc.ic.l1~C"5;tlrHlIlCS l.wa'll:!'a~oo Ibl' r = 0.1 C!IH s-~ m.!lC.011 average, 20% grc.~:tcr ~l1ml thu1\c c,ulcull,u!cd IhI r = 0.00 I CHl :;,-1 frnru ECI:- 3_ 10 prod:u('t; tbe b(5.;l repJ'c.~ifilmirnli of CD Iht eaeh r, a ,$ecob~~~·oJitkr urve (.fIJ fllllCliQJ'I of IIH~ c w i~~.d speed j ,.HIS, Ii ItCtl by a least-squaecs ~fclmiq ae 10 oU cSlimaltcd 'I'.nl!lllcs or Cll'. The ou~ are disllll1yaiill Figs. 2: and 3" j~ditiolllJ.n,., tl~e Y5% confidenee li~nil:-:; lor the fh!oo om'\'C<l1'C :iJlown In A,g. 3, T~!e pat~cm ofille re~mifJTh."';hir bct\n;,'JC'J1 Cl'~d lIIn~ w~nd :i-pccd i" roblllS!!. htu 'the ~\I,·vecrl!.:;m;C'~~m-" ,uI'Ccl~~'!'nnhlLXi by lite v;JltIlJC chosen ~or r in Eq. 3. However, ;;(;11 ~m ..... 1;:1£'<111)1Sho~"r 1!!n~nhiii!.~ inICl'l::lMStonhc dm...,g es (;Ql:llll~iem <Iud !T!QUI{l!QYDic ,~j;l,lcre<!~~~:;, t<mnd b.y .f!;,i~11.8H!ldil,\'i (J ,~, ,dkr v:,'.t1:;hi:l1,g •• u~;~:>;lntum v.d~lL;:"~~-32 :1:1] :;;-]. StI!I't(; nr'[h(~ Sll!.l~~I~~ ($. 19~ ifn,pl.y 'nil~u Ihi,1 d(:tr,,~·u$in.!!l, dmg III 11i!lb 'wind:-o
ii.-~.~ilHOI'llC\dllll reSi:ilil OrU1CfLSlJircmCIlt

iOpc!..'1!Sbc~O'!i\130

~n

tht':1l. sm.aJ1.Iy decreases ::IS d~~ \i!(i~'ld spc.cd m:lI~liTllk~S to risco Ollrvalu;;:;; [or CD 91iC im 3.
[:flUlgc

of 'CD

\'ilI,hj~

round usil'lg mcaco.rol'ogieaJ

16. $,T'" l~mll. J. Phyr..(J(~ffogr. ;M" .l'4~il U!J!9.4)~ 11. 5, .1, lenn. J. Phft. Oce.a.roC,!jf; 3;:ib2H9 ~2001l. ie. 1. M. 1P!r11'1f1~~}.Jlhyr. ,o.rct1l1!~"'fl. ~2" :U!Jo:lI.GlOO~). u. IE,ll!l1d're.1l" J. ~p; Ot:t1llrlll9~; !!4.M29' ~21!10>!). ,IIJ'_ w~m~~~ s, I'll:.ltbM, .1'1. l- QYI~lid, .. lid W. ,I!;., Gl)gd~ I'I\_ ri)rilii«liililli:j ~~ilar:t. w~~1llI 'fli:inl~ ,iIt~ ,OfWi> (illhi:! mtlilclll!le'li5ruS~'ard lblilll~co ~ ~Ilcl III 1bis W!iIllk: was
pilul pr,ojm "'$~~ t'll Sh<;:U IEner£(lIIi;Y ~OO (~[t!~~!l~ Dyn.;i!'llt~, (S~n)F' utld~ p~~t.!lml ,e~mei[ij~ ~rl!5a;~I!IiJ'Q~~Ii!he Miml!l'al5 Managi!'lniIlril S@MC! Er.Mro~liTI!n~l!1 91[jdi~ P:rOtjram~'eliMndogy. and' b:i'111e' Mintr;a~, MillIlilQfmmt Ser¥i~l\' VOOlI'!~(Igil' li!$e5i1!t1m1: ~l1d :~!!'iL'ardl P.rogrO!!I1 Qrn Hi.iI:rKaMI'oIJ1I.
~r1~
<!<;

(19~'U.

no,

oibs.cn'lIti:ons (41' :I:'O[ wind speeds greil[i.!r than 32 III ,~-~ l.II moe h i.ghcr rol!' lo.wc:I' wind speeds, b Thc. se dHlcr1()J1re> lUiY bI: ;mrib~ltcd 00 uncenain-

b)oiheo OfFke IllI Naval. Re5eilrch

Ol

Wi!!' Nl:!~1 fli~l'o'Ir,clli lj}I:iQratony'~

or

tll~f'·~N~~a

ties i~llll.C' wind m~SL!p.':In~l~!'l;md thcapp~ie<l· ~lility (~r :sin~p~~lk\d(]It..nm cl}fJ!;l!ni1::s m (he tbe ~~lWl.:'r w~!ld ~~1,)d...
t. Itt [rn~~g!. flQrl/re' 4J 6.i 686· Gl:oo:st l<iif$>ilii lIi' (lit, iiil IWWd Sl~~ ~ t1i~ Oe~~', II. s. f. ]'Illi!i!'.!..¥'., To. ~d~. C!Jml:rir:l~~ Uhl!!" fN!'~ ~ INew '1',001;, ~Ollo.OOFI' 7. 3. I'M. Ii. iIkIil!!lam i\~'!fl.. '~Il,rJ~,,l:. f!It~ ,i1#t ~l,. 'L,]j3;fil'i, lli1,10'29'I2.'OtiMQ,01:94MI (~OO4IJ. R~roerfDlCe$ am:!' IN(I'~e$

:5l1l1piPi1llitinglGn'l,l'lfI~M .. t~n'i<ll. ~.1!:.1M[tmil!l,~r~~gil(0I1.enWIliLll31§1Sel9Jl!'~71£1t:l

:!i~][~:g

~. 5. E.

ng,n
!lieteretlQe:!i. ta O~I~t ::loa6: ~Ul!f1IlMllliol fE!b:F~:lt)' :1'001 1O,11l261'xiBlil(~.lH6¢66

CRII,SP'R IProvidesA~cqlu;ireld R,esi:stalolc:e Alga, ilDslVii' r'U:&,8'5, iln Pr'D,lka,ryo,tes

.Rodolph,e Ba,rrango~.lChri5~o'p'h,e Fl"elmallllill" H'el~rn,e Devearu? Mldi~5a I~d~a~d~;.l ~ ,P'iI~r'i(Jk11:!,~y.al!/a'~,.~ S~~\!,<lii Moi Ul(ll;JUl, l l[lefllll1lii~5 ,ll!" li1tlomell1o,.lIE'hm,p pe IH,o rvaltl1~~ 11l;
Cll!Jf;b~rit"dr'eguLa mlyintIH5.pC;(ea ~hort pl1l'il1l:lromii( fepel1ts ~CRISPiR) ar'ei' 8i di~5Iinlct\l,!{e ~ea~ur;e of the gel1Qm.es of most B",(~lIiaJ 'l(lu;l .An;h~ll<1iaad illlii!' tho,ught ~,O, b~ ~rn'lJlvedil'l re~l~t<ln,;:.eto ll:JaJ{l'eriophage~. W~ IQllIl1!:ilha.t, aft.:oe:rviral ,ch:alOOrlg~, blil~t~ria, ~flte'9~GI~d 1le<W' SPiiKlli>fS d1eriWiti IIr,oFlilphage gIIilIilQIiTI~C S~(jUi1I11~es., R~fitlo'll~ll Qr i!1Ii!'dilicn !Of ~i!r~icilll~rS!JM!j(ers mltldii"ieid t!h~ phl!~~r!.ls;iiS1i~IiI<Ce pl1'I:.f!lJt'j'pe Qf the'

~m~

to

I.K;

j·{."kn..:;,j to

~clC

~pruy. IO~LI;n, Iln'li

bLlbhlCii; ffiom brc:a8dn,g, w'a:vcs Ihm. l'!.'tIwllac~h(; (iiil~L ThI.PS;,CIFUSP~~gg:ethef with @ss,ociall~dl,'.cl:l 'g!.lllleS". prQi{jd~ n'sis~ili~ .ag~ilm f!hageS\. ,~n:d ' <!rng and <lIIClw the h~lrri,,;;mlc 110 slip O\'Crdl,c ;se<ll·, W~SIi!!'I!o~ 'Spedllkily ~!!i. d~reilil'l'iif].f,l'~bv spa'oor-phag;e'~~Me'f1.(~ :S'irniloiirity~ W:i,th~~le f1rcail.y I~llil w.mcii-.oo~!lJlmn oce;:m C~i:rrent ,lllCaS'!Jftmcnts. Ih~ o-nly l111kuQW11 tCIl11Ie'l:l i'n~he simJ)I~.no;! eq1!"~ tGH o~· mo[i(Jn ~slll~ wind ,;I,C'oCr.g{lp'mg~Sarn :;IIJgl!itlbll}, ~h~ Hl.os<t mJ~c!ioll, ~S!n'Cwgg l~e i~ool!ni!lg[)N,t\, ,""filet lllreSli, Thus.I~IC ~lllm'iQr of Ih~ dm,g cocffidcfI!. systt:ms. Ulesc hl!~~lj,vnrol arb "t1.nuull1f1t blologk"a~ C'tI-w~il}' m~ I,he f,Dlanc1 m'fmtivC' illlrCCtOCl'~1 '~h Tilcir tqbiqllliloll~ disIJl00~~IJ, >lnd riers can:llsQ 'be Cl1IgillCC'l.-edi ;;mdu~mlmirulal.cd~o "CD) C,rul emiii I)' Ix: C5~im:ll.ed l:Or 11r.mge or ~tIim18. winds. DC%Sp~w ~ Ie lail:llhal ti~C' cb:ag:. cooXf'k:~ii!l1t i~ ablmdall,cc have an impoort;ml irnpa'Ct 01'1 IlilicroIX:Ucr oCll'l'l!rolphage POPll~.at.iClIJS, 11. 12, ev .. llI :ltc,d d iIII.'iI:ellilll},hore. csdnl'I~~s of CD bi<!~ ecology and Ihe eV(l~l!ti.(l11 of btlctcrial Nl!;n1eilil,_1l.~ hocteri<l hil:\/oCbeen :5eb:~"l:J h~' dil."!(;ml ill.1".'d "oo~tom~lII!r . .r"aco;.o:n;\b'i}' , KplicOll(; g(:nG!m~';S {.n Con!1ioqjucl1i~ly. h';L'Ck'q;i<l h",~{! diC\ld· hLUllil1!l;~ ~lnd ~IS(''.d c.c~Ir!ms:u~~ly I~ot 1'~~~lW~ kln ~l!c \"III!I!U8ckl~r!1lii~ctI ·'tPp-dow.n··~n Dt'~".Olll opt;~ .. \'1urj~l}' :!li1u~I!.Ir.H~ ,dr;fc!l-~ lJ~c~h~~n i?illl~, .mcl b~ot1,;'l;:h!.'iologJ' [proccs.w';S, Unn.l!lI.l!~lli''!,{;]Y. dl:l~ sEl,ldit:s lJ-7~ Ri;:;.'>LJ~b I'Mitl our [t.·;s~,lar~:h:how S ~hl!,t[;::UH{,1 di\t;'~ ;.Il"p\i of Iho: pll1~g.o: Ii[c ~Ie. nt~'iioI.ie..'11'i,.'db..t1!o:ri:1Wi!;_-,cl i:L1in{,h.J~tnuI1IPJ)1ical~')Jj~ 'i.h:;llt ClJf-!t'-rtk~ a:t a wind sr--'!"ti 1]t::1I" .12 1111~:-I :1I'OC1 ru-t~ oli:~,'i ~u~'PI~ibh::: 10 pllago: 2LU30k. illdLi,d~Il'i.~ ilofi1LIJ~y bl.tK:killg ad.'io:rJiilim'i, P't~\'IL.':ifil~,Ii..~ DNA

or

REPORTS
genem ~Ild :lpc~ie:5\'\.'.tddy usedas g"iry ~~II!ilI\,':'
{.J).

A,(:oro:~n.!ilJy. i:hc ind~L"'lry has dcvi"~

~rill~lC!ii

~~.~b:~gi~~OOH1~l;lt ph;l~ tm~~ OIl ~l:!1~!1l ~" vq:.;.~t:Y. Ll<Id.c;,rioph:.n;gIf ....n:<.l,;'iIl.iili\t.~un~!l.lI~Ls... m'ld i ph"I~.l;d. .. oomll~g p&!iug~-r."~;~jmtc,.;ll1~'Ch:JJ:niSltil~,. .. S,I'itPlt~yJCnl_'-r trll:!mmtJhih~~i'$ a low G ~C Gr.:!lH1ID->1llu\'c Il<!G!~ri1.11'11 ,md <JI k,i,;'!( !>I)I;C:~C.J';C:»a pl(1iil.<::din ~hC' Ibril1tlll~.tion of duiry Cl,lhllliC syi;l~ i ~citl~ IOrlih.c puudWicl iOll },'D:gul1!. and d~Ol-;sC, COi1illparatilv'~!;{·Iii.ClI~n1ic:;, 91~'~.IyS!~s ·r cir}sdy o re 11i'I.cld S. tlw!l·rlwld~lrrJ."strni ilS h~i,'~ p.1iC\,i;Q.u:ily revea 1:«1 'n1Mit geu~cl.ic poIIYlnOIiil~hi:>b'ill Ilrilnmmi Iy eecues aI. hyp;.1rvariab~e loci, such as. th'i::' 1.1).<: mld rl'.,'Ol)cronSl" as 'we:1~ as 1,\\l\Q ch.!lglcrocl rc~ull "'rl)i

,":h;agC"re!lsl4."ml. nn!IWlmli1 were ~cm;r.iJ~d indej1Crl"mUyb)' ch,,~.l\;lngi~g; IillC \li.rT ="ml~1l Wg~l dij.~ 1~~!"M,c 5~,.nl~;1Igli;29n. or "~~I~l!lll]!.',Irnl,~lywi,h £

or

in lI!ile R ISPR 1 ~(X;~!::J. gf the v ari.9'L~~ rh",~p;l~,~t;!m m1.!~m"revealed;liimi!~.fi~'to sequences ~h~!~ i!!hill,nh~ ~1(11Jll.Q.'O ~1~ phlllgcs u.~~~ \\ flo!' in bonh {I !), ~!~d 1:t!~ir (,~ISPR J'O~i W~~r¢m~Hl.y-.t!~, nh¢ C'hahk'lllg~ (f.~£- :2 alld tl£" S] ~h~IIJm;-..ojul]gly. ~imilml~il';l \Vt'l"'L~ ..llN:I1;'1.."<I i.hm1l,ij1wui !It\.~~)img,:, D~m.':n;l1:CeS \~!1C '~llk.i. ...r~n!ily i1lbs~,,'1..'tI IIi. the C'RISPR ~ IQClI~, \;...t~"'1t':1 lo41~i:Milia!1'lill, spaCC!1s. ~!l.(lUlC$. in !oost rit!J:u:tiom~1 m~xM'\,.~ oo~h~n~he 'wen; iliJ~(CI'I{;dnc."'l ~o the 32 ~pa~N r:n;:si;ml in 1;.\"'l(~iJl.£ and .~I:)codil:lg ~m~~d\i. No p.LT!j,cl,br ~l.." ~ht'\Vr s.tr.!liui (Fig" I l" The !lddil:unn or I1~W ql,l~!1:l~,~'i~. (1f f!.mlC'!jo[!<'Ilgmup ~l..".elln.':dlo£}iJc :';:P~Il~~1'S Ii(..";liPOI1M; tfl'pl'iflgt: :illfoe'ui.m ~"t:l'nilt';cl,1(.1 l'H(gttcd ~('iljirn.~ly. ll!I~~ Ib.lihi> R"\call 1Bl1il, oil, iii b~rilt~~flFii~d 1:Ow,lro OJ1~ end of th~ nUSPR I .I\.1~ ~'OO[niilllJ.n:'~i~mJ1il ~i) b:l'!.."~riopllflgo.:-:., theClU5PR~ eus, "'r.h I!; ill; eeusistent wi'nh iJ~\>i.o~)" o~ei'\latIOilS. loeu~. W!ii5 moclmi..'iI by die it:ll~ITu~iCiill o:i' IlE!ov~l, n I'~~1Le~r ~~~)~I''''lriBbn 9L the' l'ad:cr C'1,d of (tiC II)' ~m!lc:r:;, IIj)~rent:ty derived rrun~ I)ha~c DNA. CRISPR IOClll$iJ1 \.'.Elrl:ol!l$ »tmin~ (Q, seSLll1uj$inl~J)', ~~ ah,;;le:rn,:d '~hm ,'.lome s'lf<lil'IS c \\,\.'re l\.'Sis~aH~lol}('Illhl:'1hagc$.. ~\~li.'\IiCJ$ '~lcrs qUC'11CC m'r<l~)'~S: or~I~c: adclili:cmal SI'lZ.ICCI'l' h'l"''CfiL~d

Nine

n~.

inle~tcd

slw11lJm~iil1dmLllk: re:~at!\ (CI'l:JSPR)

loci (J 7). CRISP'R loei lYl~iC'~.~Ir}'censi :-J1.of .\1~\'~

~l~1I~O!!t'{l!'1IiglIUliliS dif(.,(:l 1\.~p;;.';;r(:.~~~par.aJl",'dy b ,~~ !thA.,~· \,',lri,dJ!c' s~\t]U(~Il~L,,~ ~ (llf e,ilk~, ~ac~ ,ml(!
are
.o1lC\llli

aies acljl,treUI[ 10 (."(IS genes ((,J~:JgPR~

a,..;."\.UCimcd).\.I~hol~gJ, Ih~ f~!nc'lion i CRiSIP'R Il.X:i h(1,,,, !1'Qlk~ot!l! ~,,~hl... bJ(ll\],gi~,ally,G~~ i i!~~d ~ili("!l;l'~I1U~)I!ol~Jl of 1.11~ ::>PlI~11\ ""1~"C' :~'·'~~~OO. se•. q ucm::.1: ~lC1l110Io.!!.'y ~~l fUilcign d(:U'l(~r!l~.~l1ctudl'" W
i:11g'b.~c'llCri:OJllm!1Ji!Iil1rlpl!::lSlIniti scqucaees l? Ql', 133SJCd C'Xclll~ivc~)i'ol;t iU1:siln(lo m1ili~scs, SoI.::vcrnl ~))I polil,C'SCS. have been pl!I.I. !:oli"!lo\'arup:roposing WT~.O'$'I WT~·S:l
Si

or

!S~$iI'llv61V~O' $!,iL9i7.2
~f!II'~'ti];-t'.a.\ i:j'lj 1i:i:il"1it1 .~

mh::s for CRISPR 8ml ess genes .. ~,Ih~clrl include irmmllllt)':lg<1~nS! IAl'UI.:'ign genetic c'.['e~lC'~lb" \' iii .~ nlCch:m~sm bascd! on It~t\ i.nHiI".. ' terence ~lfJ)" We rum~y.zedlhc! [tJS!PR sequences of vtlrious S. lht.iJ"I!wpJJifM'· :su:::1ins:,~I'lCI'!.!dil~g c~.o..'>CI,Y reb!ed i~(ll!)i.11ial lmU:lsruld rh;lgc·re:ilis'nm~ vars i!Ja1~ (,~g_ S ~~,_Di~".C!il;.~I~Ct.'S the 1m 1i.!!'!1I~X'il' and in
rrovi~iug ~ypc of SP\'L'CCrs \\'(."I.C (lb1'i.CiI·\,I~'i!.1 prinntiJily allhL;' CRISflH ~OClK Nm.mMy,plmgC SI[~I1.i>il:h'il)'f~ppi:ftlj';(:d to IJc C"orrel<iucd tv-illl ClUSPRI spnccr c"OncenL SlloCdfiC"IlUy. sf~ai:C[ umrem wa."> nearly ldemleal between P.fLr:c.~'.li1 luraiill> mId phage .. resistant dii!ri\'.mivcs. except for a'CidJbio'nml~ficers

WT_ll'l·+'$Il1 WT~~.st

]
IFi'g. :ilL. St;re:ptim;;aOill.$ th~rmap1i'lti~ CRISPRl leeus oveMe~'\I. l'IiI!\my acqllliimd spacers in phll~'~ rr'esistal1JtIIrU.l~ai'lts, a ~d~!l)[rrl!!"flMdil1 g phage s;e·!ii:,>1Uvily., Thl!! (Il; ItSP',U lOCI1$.of lIilGCCli]O (\IlIlIT) 'i~a~ the top. The nllJ'~lt."s!!l~(l!r r'l2'9i!l1'l Oi~ ii~in the nl]ddl!8': relJlllats {bl-illd, ~illm~::llIuf~), spseers hlillmberedl gray OOle1!), le<ld!er 0 .. 'lI'I'hite bo:d, and teorminOil r!ep!3la/t (T, b,!"J[Ik~iiilmo:nd),. (BoUom leW mh~ s~~(~'r ((lillieI'll Olil. th~ lead!er $ide- I),f till~ OO(II.IS. ~ha{l~·re5:j in, 5itafll !1!ll!ltg ~tsi~ de!:ail.ed. with Ii'!@wly ~Ciliilu~r~dlSPl:!OOJ:i(whit~ IbQx;n S!I. tOo S14~" Ulo(tom rigtl!l) Th@ s@~s;]ti!f!ily (Jif ea:c:I1s:trrailf! to Ilihagles :S58~lnd .297:2 is Iref'll~es;elil~,d a,s a, M5toglram ID:Fthe e-f!kienl(~ of ~laqiUinj[J (lOOP), ~~hidili~ t:he p1aq!il@(O'U nt ri!li:o of oC!1 [flUi!'talOt. stlr,;ri1l1 to thilt (([I~th@ II'I'Hd·'lyp@.

p'utSCilt

.illl~le

lm~.cil',These lim:lillg.s therefore

slIIgg'C::lt OJ! 1J()ICnlia~ rc:latl,o.n be~wecn presence tire or additional :1fJftocr'S and the dilnCreua~ observed in Iltt phage SC'iIlsililvi"ly uf'a giv,clfI Sl:min, This ebservaden I1J.'OlllPt~ us to hlw,';:5tiglJlaemhc origin, JIHlr1m.clioll (lraddi~torml :sraCCJ,~" pllC"wm

wr

inpIID~p.~i:5~.mt
Fl;~I,wc

1:Q"unatUs.

~~'~lcd U!C hy~mlh!''?lf!'l!II'_I~C'RISPR

loc:~ ~Lrc ~lj:llC'n:d durtug the n'\l!.I!.·OlI~~II.:~·'Lti(l:v~ of plmgt.r-.D1,~i\jis,t;,1):U. mu·t.;,1:!11::-;. A pll(l~ht:r.i1 l110dd sy~[~,n\'\l.m;M:h;~~kxl,c\lmi~!:u!,lg of l.II phugt.... . :YC115iliv wild:~a.ype S, Ilir:'nm}pJiiAIN slro:i.ll widc:1)1 e lIooCi in the dIlI.irylndlll~try. DGC"L77 ~.O Iw.il~d l:y-po:: (\1\") J and two dl:>l in.cl bUI ,dosc~yi!'Cbtoo "im lenl!. bac!tcrnoph~cs isolllm~ froJnind!llstri<ll Y~)!!!ItlfliS[l.mpk~phlc1:g~ 85:\ alld rlmge 197.J (l.Il"
lo.a:mi~o< USA Illl'!!:~ 3:329 !lI_gJM:ij~lUr~'[)111M. Madis.-on. WI

531106. USA" 2D~iii:Q

[)!'iI!l!1l\HaifilH~,m1ail.,.

de Mlc!'O'bI~eJ

;,rq~~!<:s. 'Wilf1li'!\E'!~it6 ~t.

d~ R~l'fl;1til) '~1iI IEmlt:IlI~ IBlK~iIl.~,ra;[ult~ dl!' l'l!i~d~rOl1; Del!lt!iire. F~I~ d'tCw,llllf, lI:.e~!1J~e ~ntI!I' for ~iI~

FQPW SP!S, ~ilePwl_~w 10" f.~~2(1 For;amre. JID~r!M'iffit de lEio{iJil11iE1 i1~ fawW'!d!e;; SdeQCe5. eI. die 'G!!Qilie,. G~

IFi'!lI•. Z•.5: thftfm()phil~s p:l1a'ge ge-nIOf'lll@ maps with the p(l~nti(ll'l (l1~ seqtrlei!l!l;!(B siimlilaroo tit(l' ~((Il;Jired (friiS PRlspa(oe'f5 Oi[ the jlm[lg:e-rre5i!>talllt. mlila mh. SlNlll::f ~ :lj,h'OIl'l'III al:llwe, [lnd be~'iJw the 'g,enomle l'inalP~

"'1'0 \~~m IlhiLippe,'l'IolY~~IhI@d~msm,,(01'Il

QYi!bec. (a~dlo!; oo~~e$p(!l[nl1k:n~e iIIliOuI_dIbe addlre5.'Sie(il. lE·,rnait

Gil( jI,p,tj

]n,dkalh: ~ba~ th.esPi!(!er FliHlit~heS;i;I sequ~tn(E: Q!il tl'l'e {...)O an~ ~rn the ,-:) :straiild, r~pe(lh'-ely-,Ai11 . @s,tui:sk if!dl~,altesth e-xl"~·ttl.li'l(e ,il! SiMP b!:i:hw'Il!1 th~ ~p,!!(J~r:I@ql~@~l:eil!fqd thal olf~h!;l lph"gi~ o~· 9!1lIUillH~ (fig. :51). The g,erlillrn'es.eql!leIINl~cd ll!hllgll! 29'72 (il~!t:mion nLlmberAY6997(5) and :pnag:E1 ~:I'S8~11e '93% ]d-e-!l!titat

REPORTS
were ro,~i$t;ml ~Jlly ~ Ih~ rim!;'!;!~!!!I;,"XI in .~~ C'1il~~ lengc (fig. l, I,· 1'h~I'~;IgC""re~~:'I"m;~ro nhl p ~>r;~lw~1 oor!'d~1:I,;-dt(l~h~ ~~p<!oor!l;Q:llK'ill~. S~I~~ ~]~m.~tL'aiJ]S\""~lh s.:pnl,;l,,'I-S,lm\'~'ilrng!OO% idclltilY s '~o Sl'qtl;¢lh:)C;, I,;mJ~";r'\~~ in rotl~rhllgt~S liN"1.."'f,L) m.1li::;tI!lIt to bolh pJ]I;1Ige&.such ,I!:,! $1'I;lC~!5 &3. Sti. •!ul~ Sf. III 'CO!l'Ct.I~, when nu!;k;o,ide tpotY~·lotlQdelCmlir~e
tent

\\lhC.nlC'f(,l~~SPR

::lI:l,'l!R;Cr con"

tl~tll1'~ ,:iIl,,£W :peslslmloC'. we .. 1~~'"l;1 the CR!SrR~. ~ot;:'l"by \l~~li!l~ ,,!Ki d";:~CI.ill:!il p,IiI;lL;n; ~ (.IJI an~l iJL""'il'd :>1,I~.I.l!l1I,1~!1b. snnl~n sens..i~i\:ij.ly10 rJk"lg).,';iI .. I\U ct}rl'lii:mlct",\~r'~ .gt."i'IC"r~lLLii ~mJ, in!..:g"!IJ<ld ~Ill'tn~lC .S,ir/~'mm1J1~i!M~ chL"rnoo:;lnmt: wid" ~lqL;'~!'®'md~\<~lflpocl y R\l'!I;idl ;Im:~I:U~~hll!J]~n!;'T b ~;h~~!i'i~ o~~r\ ~d l1l!:.'t'!'.'I.,'(:liJ 'nilC ~p<!Q::r ,nid 'Ilie \,'l'trc (.14). WI'; I\,,"JiiI'i[}\!~".ocljlK~ srm:'{.~, 1~Pid~~OlIll ... il'il the ph!ll!\C :-.cClu""iiiii: ifmrill ~ '[0 15 siJi~I~3illUr,h~o[idL: CRIS.PIUk!o,iliO uf ~tl1l'l.i!i\\fl:¢,&l8 ~1S<l 3Jrndilt:pJ>'iCliitl W!{j\1ymorp~b~11~ tS,P~) ever 19 er )0 ili,~el~t~ 1JJ1~u~1""iilh s1.ii~~ r\.~I\VilhOliJ1l 3 fll'lly :;j:O~" V2~ lide.a]. ~'C':sj'itiCi~i d~,cl, tlQt ~o;~nlo~ro'i!~cl\;~ ... The a\,~llljlfj~ :Y!rninl\~.rr~ .oIfiIS;!&t:[USP.R I was
l»1l1i~1as

'OOt'l\'~'1Jl

!h:~c observed mooilic;ni(ln:l!.~slatdi,~h the hnk the CIUSPl spacer C(lmiC"n~ IImll):tI,,:gC
~,~i$I'l!n~~,

l!ll
\1\lTi~ pinH

the trm;,.,;;:i.s

of!§,,,mcrafil1ill,

WT~I)H8 +ISJACRISPR
''iI'(C!.Gf

I,w,;

:>irail~ cr~ah::d

"'S,IS:I;:p.Jt .. ViIJ:Uan! 'tlli.l! C"i1l1win~ the ~!1t,I."_ wi~hll ...i.!1Igrc R.'f~<llti.!'l"C~l;d .... lx.

l!Pig.

IWI.'U1 !IllL: ,mo;: gencs 111id '~~lt ~\:r~i\CR ISPR I Jklltl,l~ C 31, Unc:lo:pt,.'Ck'tlI)l" :l>ti[lIia'iWT ¢'~~+8!Is::l,:p:r!l; WII~ s!eui~iti\'c to rilila:g~ :-:5t. IIh~!m~.~ih sp:ll.cerl> S1, mlt! S2 m]1Jr.iu~M O~~ ,c'eihrof:l\~ii:O:l;nc ~ (F~g.l).

sistancc,

O:':i~, ~ o'lIncl~l!£.. s.:!~.In Rdditi,o:n. when ~vcml ~~pac("'.I'S wcee ills~I'l.oo ,39' 'to SI4), I'h~ uc-

~1~occn;. Sl, S,;,l,SS, ~mI S~ SI.

:i.CJli:>hivoeto p:hm<2e 8:53, 1,1, hieh imlicawdilhm (tiC I~~l.~;:f.'e,;"ismru:;c or the o:ri£hmll'i~l<li,S.~Ho;!sisllmt
mll"t.'llnl

:'listm1C'e:1:(Nci~were higti~~r.These IindinSo"incii-

cou.JC~h!:!l CR~SPR~. locus .~~ !ihc ~"I.lhjoc~ 10 d~":I'I..'1:l1liC' 0tJ1d r.Jl1id (1\ oluJIliomwy ~hang(.'SclD \'1m b)1 nlh"~i.'; \,'IWlh~1r <lddill~ SPZK'C'f1' ]~~m.'idl,;'flIm\'c~ ph"!1l~ ~J:.pO!>u~"A![O"g~'h~r. !II·iL,.o;s;c r.l.~l!Illl" ['IW'L;:;ul lIwu~ ~,~i~11U!liOL,,~. w~~~~fltX"cl [~1f; CRISPRI J{li;:1,!<l til' CRI'S['lR loci ean in(Ii.'Cd be <!~tlo!'rod uring Ihe d stmin "VT ~~0J12 ._ W I1J1 '\ cl~jml ,",r11~l<ljnn,gCH1 1y ,I ~1{:Cfl' S 1 <Uld 52 I[ J 1) and 1l."SIA,.;"XI t~lhi!~h~f (he .!,!.C'-Il!•.IL".tVOll of ~11<'~C-~-!l~[i(a,m ml.!I:;,J!T!l\1 and ~:~ ('!\llOll.lli"h OJ! link b(~'w~n crusrR cm~tL;'!I~1.mod rhagt.l ~'·n,"i.~livi~)1 'illbelctl. R(in~~rk;lbly.the W~N ~~I,lIlin!J SltEli:n 'Wl' ¢jil97.,l~S4::pSI52. .!lJcin~~: 1'1,:f1'h<lg~~('Il'l.iilil\'i'~.y.111t;~ ~ IldiJlg.~ Sil,lggll,\l>1 mhL;' I.li!:::u p!C-:;ence ar:a CRlfSPR. :;;pflcer ~rlcm ~:1~to a pMgc sii1l.1lUlCe to p.l'Iug(! 85g. Wliich slilgg..:;s;~ctl U~~n ~(l[jlh.. L: provides ':i'I'C resistaacc :l~~nsiph~g(;'S; th("~~ct\lo'lO SP;HX"I"S h::lv~~h(: 1l,bi1Hy [0 POO\i ide comain in£!! this p31ilic:L113!l'" sequence, phage resistance de ImO\,O Fig. J L AimgcLhcr.
1

~h'C' 1IlC:i:CnoC' 1 ors, and 51 U::i;, :n Flmil.c!". 10 <!(k!~ '~h~crili'C••~ (JllClSlian of

(\:vr~~:

+S~~'J!

"}W;}S

~'(libabiy Hllkcd

Ie

Sim'iirurl:y. lite \:\IT ~'P.:~::pS~S2 ct'mSIn'!lIICl 105:1, Ihe i"(:;s:i:,,;U!LnCCIO 1,Jl<lt!lC 2972. allhOl.llyl~:iracoC'li" SiHs,~r~!'lem i11(IrIC Chl'OI'Ii~'i.:i:ilmnc I F iJ:l" 31. TIlC'Sc msults ind iCOHCd th,ll sp;at;iI!L~ alone did 1]/01. '~'wlvidc ru!i.i~tmlCC •• mel pt~ml'~,.lhm Ihcy h.nn,: leo ~~ ]I'I,~ ~l~!n ieu~:!iI:rg:cr!I,,'1ic conic .... to be ' 1:,.~lTo~!i"c " Ail~IID!in!l!:h i'l1illOlI \I!'(l~ SlUggC'i'~OO invOolvCHR'ut in DNA rep'i! f I( .151. Ihe CW!ITI.1!lt hYllCllh~.)li.."\, ~" 111.. 1 . u:~~ ~,L;n!ts {j'•. M~ a~~~wo~~1i1o-d CRJSPR.. ill ~l~'dial~".ocl i[l~nf,uni!.)1(it}). (",O"JlS!.,'q'I,li.Cnuty" \~IC ~Il:" :llC'r~\-:mL'rl~\~1[) £U1\" gc<i1CS in :>tllJ~a1\¥r I1~M +5 ~S2 ,Ill): l'it.d ({ UG35 13 I~Ul.,d{'fA'i 7. 'Ii~'Meh cquivMe' rucm 10 .'f1I 1}(j57/smfm5'l and s'r06MJbm .•fl6lJfJ. Il:'.spoctilvcly 1~6. T.~le t'tJs.. illllCIJVmiml re'i
sultcd iu loss of Ihe 111Ul~c resistance

n,

and perhops C:I!s.:5 S :IS a Im.dca.~c'. beeause :~I. liel C{lmll~m1·:m UNI i"~YI'IC' nucleaso Illoli"f. h~ CO:I'!~ ~rmH umcti\i';.n~ng ms:7 d~~vgrli! a~tcr the re ... ~~~ .. i Jl'ICC to Illwgc &5:R(Fig .. ::ii). 11I;1crc.s~iIlSly. we were' l'CU]cJtodlly mwt'l.~e 10 gener;nc CR]SPRJ
ph<l,!l,C"I'Csismm f1HI~lDl~S!Torn the' ('as;' knock0I._U1. pcrhap~ bcc;llI!sc Cns1 i~, im·'(~h'c,-~i'll ~hc
1S},llIlhL',sis !1l1tl for

tria ...H.

ilrL"i;T~ ~j,!! (I I"' IWW ~p..Ot.;IN :md.

S(;l:is,ulu .. iry
btl!

rrlddiD[o"~!li~ repeats,

Whcnw(; tested lll!'.'

resistant n1l1laUIl!>.we Ihtlrrui

of [he p'hlll:gCthat plaque fonnlmtollt

thM auclilllh-:cI;o

'V.

~JJ]nmnllmln

Si1lIH;I~li!o!'h\l ~Q¢I,6S8
1~'

~f!"~'

I)"t'~J

iDil1~1 1

small r'OI1Llirnlion or hi'tc~criophagc I'(taim:d Ihe ~bilily lO nllrOCl t~'lCtlmlmlls" We rllIrlhe.r !lllalr.rod I)hagc' Vmiani"l deriv,~-d fmmrhagc. 8581hal ~mJ,flcd ]bcaib:i ~hy tQnnllecl. ,\IT1!~£lI =<SIIS,;!. ln pflrlictJlar. \/01(: imrcstigaled the seque.nce of dlC :5~ mK,m~:r,cgtiml C"O:troCSI)ml'd~ IlS io add illim~;[lL1 spacers SJ. and S::l:in hVO v~rul..enl, l'i~l;[!goC "miant~. :~n.IJlOdll C1)SIi.:~, the gcl'l(l~n~ :'icqU\'1J10e the phage' V;lf~ iOlin~ had H!li!~I~r.x,I. a~~,1 '~w'O dr~'tU!lt~ SN.~ were idi~~~~ii:k:d !he M:1;lu~m~1C corJ)I,"N~(ll.~dijn~ in 'to

was dlrmnmicnill,y reduecd.

or

II"

W[(~..sl~

:;;p;u;:~r S~
1

~Ilig..S3~,

!til.
I'! II.

Wil",~·S"I'S;2i!l.CRi SPFil1

WT~o.$i~::IJA
W1r(M$;;i:,.s.;:pS182' W1r~.sIS;2:: :IPC'BS~
1

li'!I..
'V.

\fI1.
Filg.

Wtr:~'·Si~:

:lPca~7-·

Hvity. I, m leila In~ WI; '.,~S;~:l:; lilli, IiI1l uta IiI~WT <M5Bl'mz A(IRI'5PIU ]n ~hi'c:h. OU:5IPl'l1 \V,iilSd,eLetl' 1:11; p...'""aIS,:IS p;n1i or an a~~ivl! III~ m.~t~l1il ~tB!oB.~~;PR ]fi Whl~h C~IS'PWlll ",as~~~1il(ft~'.!1ln~ ~eipli!l(~d with ~,llllfq,~ft_~epe-at hrtl~cr shldics are (J".sil'w p,~a~ed ;,witll1 a ve r:;;iQIi'II(;QIllt(illlnl iIiIgS~ll and 5.2; V, W''I'"¢l6se ",sas:l\;pcQs5- \lfith ((ls5 llilaldilfa ted; \1'iII. WT'llllii!i1l +:5.'JI.s2;; ~C:QS7;'_ mt!li ,~s ilil'il!(t~'\I'iii!t~d.pORI ~Fl(n~!!lt~s th~ilnt@gr>ll!ted plasmid (22). ili1'~P"'@9~ 7 ~~ns;itiv~t.y IHH:fui:s:~rllin .~() 'phflg;e~ ssa: andl .2'972 l!5 repn%'!1!l1t:ed at t~,e beUaml.tls ill his-togmrm Vir or tt1ii1!' efriciefif:Y or plaq ui iliIgl~~H)P).
IV,
Wil"I~4:'iI72 ~

3:. CR IS,PiR ~pa (@!' c~,(:!'i f!@'~r]f!gJ.

cos

gJ@if!eilfl.1lCli'V.ar[ll(1Il,

aJfldl ,wrr@s;p,lt'j!illUling

jll'HIQ@' sef1ts~·

Ovendl. l)ro:kafyo~~"PiJIL';lr m have CVOh'~id I.i UIucleie acid based "immunity" :sySlCIJ I1i VI"hii:rol'Iy speel Ui:.i;ty :i:.di~tl1itcd bythc eRl'5.Pll spacer roll~.cI~L,.\\'hile uhe rcsisml10c is I)roddcd b)! Ihe , ~ ,c~lzymati.c. mill.chine])'. J\:cl\Jillklmily. we spccliI~m...:ihat SCliII:1C or ~'U.': Ca,'ii~~llies nOI. dlirc~~ly pn_widinlgll."'St".,nanlre are 3.cilIlilUl:y ~j'l\!o~\'cd ~DlIhe insert iOll of 'I.i:k!~ limlll ~ CR.ISPR ~a;;;crs illXId P."'"il~lll~I.Ul:··

l'C."fliIlu~sC'.

::p,S]S2,

mu'loT!ml o~~tra~1[1

WiI"!I4;!1'7l~1Ii1

w:tm:1l CIFlII'SPR:llll\la~ dls,lltac:e,ilij and r.c'_

W b~lter ciJaraclcri7.J! d,c~ledl;mi.~m ;U::lifDil and 10 ~,clen~i Ihe ~lr.~ili( 1ij,lnction l1ofl.h~ V,1I·i01l5 ,n!,": g~l.~

'Or

ry

nis

J]!,tclb1L;' .. ci~~--ha."i.1;;'"d5;i~¢ln Co!Hnm~ti. ',Vitfl <!QUlirlo m:id-h;.~~d CO~l l('IlI~I.I1'~ ~n Clilmryob .. lhr,llI,i,l\\h n ;;s \ . ,r]j ie:li H(bp:~Lii\',~ iml:iillnni iyh~ tml lin!]Cll~,I!lMt'"

117111

REPORTS
The lnhcritatrtte oo;tm~ 01' CRISPR Sp;!CCi!11, ~P"~}(lI1S the II!~ of C' ~~SllR '1(Iii;) as m~~ IQf 'C'\'Ol~lii:Rumry,~ypi!!~. (;o~np"gm~'I,\~ g"'lpm~~l\l{!i1,."" ~I!~ ,('9. !9), BiJ~'1~lLw !!l~~ S,}1S,tC;:llfiI~~ ~!M;;ti\·.; tv . El~rli!rePlce~ O1od'INl)te~ 11. U Bte:ilbarl,F. !IW1n\Iilf. r~Mfm~b.foI, :n,278 WXI5). 2, S. CI1i~~~O!e!1n(!1§~ It, ~ ~,",", 10i~_, H.Bdi~ A lJam;tWl 1~ ~617 (lCKI''l}., 3, ]C lit S!:I,I~ •. T" R, It:;iIaeiiilfi~mlli~:, M{j'~HtI!; ,,"rC'tobi~ 4, .~9) (2iJi061. 4. lit 'fl~ .Aim.tL ~ .... .Mk.ft!\M,.,1. ~~~ 2.83 lZOOU. s, lIt J~IIa!1I, J, nl",fic '~n ~Effi'I:!d~n,II'~, GaaiSlra, L~ SdiDI!Ji;, MQI, ,MiatlllfD1" 43, 1565 '(2ao2~. 6. I'i!. Eli;iOOllin~f W,. Nat BIi:J!U~,!]'at',2. 15504 (200~t 1. !Ii. iIC~""in"I!' 'Q!l!ii1l~Yi}, Ii. SIIroli.in, 5.. 0, ~f:lliliOh.
M;Q'~ri1'!lt~l •. lS51. {200U 3. IF. J. M. ffllJiicill (. [lr,~;\l'ililli~mr:" . Garda-M<I!7Ifnt:'l.. T IE. Seria, J. Mlii Ii"!!aL 6~,174 Q:OO~L 16.

fi'.

HI. H'.ar~~ '~lell~J,1II" IE, f,MaIl~~illll ]. (~;;rl_ tJ~" 1l,~60 i(~OI()l P,. ill. A. 'Gr~~ s: L 1klfi!d:HltEn,.
pf~

o.

K. IE. ~el!;an.

n, 't,U'!

~Lli'lgM,

1.11~rhlJl~ cnViWI1Dl1~'Ul, it likdy rillY:' a sii!jailicnnt relein prok,l!dfQuic 'i,;,vohl!!i,Olli'lr!d ~,ol· OMJ a!ld 1Pr(l'l'i:d~ O!!hi~~(ljcic;d: j'i!c!}1f1!;.·cli\l~ ot ph~.g.,n::l\; O~!,iire" <II!> \'I'~I~!h ~iprt,dic ~j'i.'l; IOO~ rOF p ph~gli:' :.c.1£;;iln,llty. The CR~SPR.-LYJ.'!1 ;sy~l(,irn' Imlay lii{lcord~itgly 19;C ,cxp~i!e£! ~!l V;l]Ut': cld\,:-j'ise It'I.eJ!h~ afll h~~l !lnd else pcit!!miany used I!Q :rotlilrenhl.:
di:>:>emiina:lioll ofmobik: ~CllcliC': dcM1CI1itS,alW

~he aC<1,i1il1.i~i(lnf 1II nlksit<Ib'k! I rnlts, su-ch ilsmni~ o ll-iiotk' re.sb't:moc j2.CI1C:ii. ml!.! \',il'lllcn~~ nmrkcr~ .. Fn:llu a pllii1!;lt' cvolutiml perspeciive, t~~(.'illlC,grntcd '~lh,!gc ~'querllX':i:\\'hllin CRISlPR loe~ Lln:~)1
0;1.100prQI\.'i,~ Olddilijo!1~':~;lJlc!1i1f poillil'l
n.,lCQlllbin[llLir}Jl

~'"r. PW~CE~ G. :S~gl1i~"

'G_~g~wd,

..~iicwbiWaw l!il\.

]. D" A !lWI IEnl'bd~~. ,,iiPo.!,.!iAi';rmlli«. lIQ\ 10117 U,9~~, 113. Eo F. ~~wi~ ,~J",r..,]. 8al;;rff~, Ml7, '~?l~ moQ)} .. 19. fI:. 'f'., ruBo.:t, E. I~. MI!In~Qdln.. 11"lB. Elriff!illln, K E. NIlIllQm, .J.1k!(l6ja~ 18e;, 23M U~(I'6.). ~(l', It, 'Ill, 1iI~~~ri\1: !!lI' !:/L, fmC, IIIfJtf, .A!;~ S!'~ uS-A" 9~, 2192.umJ\ 2:1. ]" S, Go~~~, A, 8kiieIWJt J. Mar. £wi. 62. 7li.3 flOO6}. .22. 1j\I~ l!ili,ni<1: L &l)'tr. C. '!1m, ;md A..c. CD!JIII!..M.oo'101!i!n of D.i!nij5l:Q ~!}~t!i~I'!, es ~eU ~~ J. 1~i;.Qnle ilOO o. Ti\Ilim~,.l)if lfni~~W!! l~al. forr[e(h.nkal ~uppcn. ~i!1!I [ ili!cm iiI<liil~1i'I rill' diltu~si-rli'l lIi'1cl mtl~al. moim 01 ..
Ih~ ma,nLl~rnll't. A~" fl'je Uu~Ii:T. lit Kllaifillhammer ('II<r

~S;3(~O~~l. 11.0. K S. ~<irg~,

WIlli. (irii~liin, S a .1!.,5~~1., ~'i" " i!\lOl.f. IE~V. tliacmin.. 8foJ" Dim.r :1, '1 ~2006).. 1t11. (, Lf.wsque f!' rilL, Appt. Emron..liIi6tib.iW, 71, 4iO:~j!
(2005) ..

ilnMdIDi!J1ll1!! ~1'I"~r.i~onl ~mrrn. Ui~ iMlill:. WIII~ ~rlell iI!j' rU.OOllllijl[rom lfooilW M, AL!iOI S, HI, W!:!~~!I
li~nlll a!l!!I1CHl!!:edgEi ~I!~!

W rnei]:iwh,; dWiring :;'Uhi(:tjUl;"!lU 1'I11i!I~ irdt'C~icJll.... 1:111"" im:p''';U:,~rlg till!:!' gCI1~ !1;00] 10 \""hic'lil

!!!lid ~gin~etiIi91 ~!!Id

Di!~QIjl!il'f !'rc9f~i'I'I.

CO~lidl ofC~Ii~,Il'li'.iEJI{)
WI!f~

r«rn iIll1'~ lNai~!ifal 5I;j~~~e!5 iIIl!jl'IlI.'!ih!!dln

~IiIU~~~I!!!,

~'"

1IIillQflli!ltton

ill ilI1'<I\elfials ilIOO lll!e'ihl)l:k lor iIle ,~ene;;,llflO!'l

~~h,~c~ h,w~ access W'J, Boc-oIu..C' RJSPR loci V o.1U"'I.1 fbulld in UI'O IT1lljtu"j~y (If bOCh .. Oo!.I g(l!W"'~~ ~ .m-d
~:r,~ i!!biql>l~.Il}USin An;:h,K'.l (3. U. Ihdr l'iJ,'lu;ly wil~~ pw~ idl..~n!CW ~m,ighu; lnmrh..:: .nc1ill.ion :.uK! codirct"I.oo, C!,Ioi:lltltm bctwuciI1IpmkflryoliCS m'Ri

m,,m!1I'l; E'fl1i~Eeri!!!I!(!if Cfll~PR ~a:oet~ 'm~~, ilI~ S:)}, ~nm i~diM ..iDIil 01 (~ g~n~ i~ 54 t ~. ph~N~~,!!'!t
:t'l~llal!it~ CI"~i!\I'I(',t OmWn~. 1l3~ IR, i( l'i!lestl1. P. 1P,~.milri!r. R:. A. ,G,urett lit &rU~f.li!W; Af(fI~~~', 59'~'OO(ij,. lL4. W" 1!.t ~U.1i. R, KIl:i~rnei'._'.li:i..mm. .I\Ii~

GeflJEl~Jlk"<I«e~lDrn

!lll.rn~~.m

HI!3~>l5a.la'

Hl!34S'0'4.

Sl!Ipporlingl OIil'lilll~ M .. t~in'ii.d ",m'Mj:,1Ma!i'fiIlg,af~Q~oml.emvrilll.l.BlSlSei,W:i!7(j9ma

In

MilIeri.al!ia,OO M:lilJiad,
••

IRel!!rm~~ riUill~ ami

~~ Mtr.'.~~~ ZQ~ ~,~~!~ 10,Tl!2~~i~mf!.ln81.jjO

:H!l5i. 51 ~~ ~~

thDll'prOOMOl1;,

,6'7. 011361 (~OOl!lJ:. 115, lit s, ~'~r@!@,. L P;t~illijjd\ ril" li'_ 'Gtil!liip, I. It Ro'gl!![i1I. IE. V. tlioolrlil\l, fJur.i~ Jlir.1th ~~CI 482 '(lQ:02:~.

1,:6 Fl$m~11Y ~Oll]

A IG Plrot:ein-,Coupl,leld

,RIB'Cle1pt!D1r' 8. Is Plasma IMle'mbr'8,ne IRec:,ep,t,or'ior tlhe! Plant HOllr'mone ,AbscisicAc:id

illl:id {ABA) reg,ul.al~eo5 m,my phys.iologiGlI. and dlell\e1opmental prilc:e~es

.Ir(lj)jfJt¥J.~i~ il'iltll,ivc' G~(,R IlIU~cin ~GCRI)has l~'IC'n c~'mr<lc~cl;zcd in Illants, ~! ,!:O~ ,md no 7 ligand hi!); oo~n.dc:~~m.1d an}, r~.jj!Jll G[IlCR. ror

To id~!!~~ p:~~~v~ou~.~y fy !l!nl!x''COJ,lUilJ;'(!. fi p(_'~

I'lf(ll1Jir!~ I!] Am,hidql:J;,rl:~·. Wi.~sm,~rrtol'd by :si,""a.rctlI~ng ~hc Anlhj'd(Jp:~;:~ gCl'lomc: [md fm.!!'ld J gene 'IGeR:!, G~!!IBti.lnk.:tCc!.~ i()l~ code j\ll,g52920) ..

e<nC'(lding
sh'LlL:'tm"(~

<lL~UllHli\·L;'
Inroictionl

GPCR,
i'ill!,'!;gc."'z..

TI'llIIfI:.:Illllt;:lUhrtine

fhm.t GCR1~s

The ph:I!n'~h,orl'IllQ,ne alhsdsic

in ptalil,!:s' Wh~ !1fIedli1ffi1i5l11il,~~ AIM ~foepti(lll:::.tthf.i: (~n :SJ:.lrr~~~is f!Q~ !!I Ii! de !FS'wC!d, I~~re,. ,~ ~~port '~l1a~.:::.G Pf,t;!!l~fI'""(QtJlploo r~';i~p'ror ~~hCd!lIl~ al"1ldIpliPiSi(:~I~ g 'iii'lt~r~ii;ts ,l,Iith '~he G prl(lt~~!1

rn)uc1l'i""ithl';(:vcri lr.'lIlSiIiI'iC'Jubrune hdicc:. ,!~ng.5 t 1\. and. Bj. The "~l:IIbs.cquct1il(\d. lliillfilr 1oC'i!~i".1iIli.{)n ailllltly3i~ eon Dinnod hs p ia::om8
Iml(;iIiIl~:t.JIillC:

u~l!Itlll!liIit GIIfl!A,l to medi.at~ ,aU mm!lll ABA r~:$PQn~es~n A.r(lbidojU'5. OIofe~~:I:.pr~:silil'g til;!!: re(:~p~r reslillts in sn .t!.BA-hypersl:!flsithre phelilQtype" This rece"ptn.F biind5 A.B.A, \yith hngh ar-~ilnl]ty at ph~ioWgical ,(o:n(:emltrillio~ \ii th 'El'xp!1!cl'El'dl ki~,~tics ilnd sloereo~pe{;ifidty .1I1hehi IIIdlifl91 o'W .I!! BA ~o [he rec:e-ptuf Ileiild~ te UH~,di)S{I(i.1ltiun oil '~he reteptGl:r-GPAl c:omptex in yeilst. OiLir re!>IliUsd,e,li1onmdile
thal

I\1JOI

um:mnbmllc IOOIJl~j'7,;;,ijlklil in the Inrmsgeillic pll;)Jnl liig,. Sl C). Gl'1t2 .~cHow liIuol\,'Sccun 11m-

thisG, ~rOi~eiri~(lIl!ipiled

w,~epticn ~s ,11 pLilSl'IiIlI mem,b.rlll'le

ABA. r'ec.ep~Qr•.
5Ug"

teln ~YFP) is detected in the ~lc~llbram.: rmc~ron iso~mcd froJ1l~~le ' .CR2.-YFP uun:;.geUlI1i:rkum .. Silni~a:!.' to GCR I (/9). 'GCR2 IS, m~tly 0IJ;l1(F
('~[cd wi~h ~h~ nl~lI!bIi1Hl:1;O [!<LC!iOll (fig- SiD'). F~!F1~lt."I·I',10L1C. even at~CI''washing whh dc~cugclJl,
()f <l

_~~tli.ln~glOWl;]l OJInd d~~'.dopn1'C!'~~ 1~,uticWl.a .. Im·ly in n'''1:1IOln ..;.~ ~~) t~ll' IRIIUlbI;:nhd :iu·,,,;....'i,\."l'i env ,t.l .~h $cvlI:nul COI~:lpmlllllts invo ~"cd ill th!.:AUA ~igJl'~1 p~dm1ay hflVC bL't:lli'dcnti I~(\.d(4). T,.V(), ialllj roccm rcpon ... have Slro'VI;,'" 'lh~"L1 the ,Ill Llclcar 11.N A biiJIclilfl,g prol.C'.iifll l~o1{.v¢IriR~glime oollitrol prol.cill A) (5) and t~c ~hlorop~i!I.~ proteilfl Mg chiCl:;]I~"'_~U ;qlb~mit (6) arc ABA I'l::OCpt'0f5 (li},

A
__

b:lCl s.~'c ~"~i'~ il!\, I is ;~n impOl.1~m~, Inoouur.JJ5,t" :sevCi~;:!!.I;C;)f.ncr c,'li.pc:dm-.:nUl, hod ,t A
horn:u;u~c ~l,U
!l1!Cdjah,~Hl'_IU:_;'" <l;±1pl.'(:"~

(~r

:er'

gil,.'i1~ed ~I~l c1IIAr.JocU u~.w 1)I."1(;1;r~{li~~ S cd~ ['I;<!I~'or i ii Al::!"" ~o"l;hi1;",~i!.", il~n\:~io!l!Z'l 7 9). ThlIN. (lli~,..r ( A BA n;,:tl~~;~~T.'i. '~1'lt;."C ~,dl)' phl:;;H~", nh:rnt~mlti;'~otnl:a:tlnceop:loll<i< mil)' Ix-l~cK~ i itI!'~f:!J: p 11,1}'~~r:s~(~r ~)crcdv:lflgc}j,'nrn~lI:ub['ABA-and, ml1dHflIiug ~hc cl fu.. ,,,ic . I\1):A si£nl~ Iling rcSPO:IISC:;. " LigruldJ-.mcdimcd s,iglla~blg ~u\:)~I.~hG pro~Ciflcoup:lcd P;;:OCI~10r;:;: (G:~t'r~s) is ;[1 cOIl.'!crv(;d meCl'mlili~m tor ]he e:li~rnodhll'lilf siguOJII P'~l:'tOJ)-

higl:lc, 11.11bulle'l',GeR.::! ,~~retainCid willi dlC me~llb3.. nc fmC'~t{l!l. !'i.ll.gg\o~~i~g Ih~l~ Gel::! i:;; an i!lt"'~n!l~ ~ncn~b;r·;;J:UlCi I):t't1I.l:ln llfig" 51 D},
Cine f:t-;'ltlUfl;.'i o .. ~l!!.:(iPCR

i~ ill;

.. bi]~~y

'I~

illtli:'El!It·~ willi:! Ci p'ru:l'C:~no Ih·.nl:l1 ~L compkx. fn t cOII,linn the phy:oic{!J,1 .intm"r~c.I:i,OJl b~hn'l\;nGf"R2 !lind 'Ga. we- u."cd Ihurd ilnicre~l. ;)PPlro{l.Chl!;~ICl
dCIC>l:I.lhciii:' iinlcrn.cl,io.i:I, '\.Ve I]lSI, used! »urfacc

]'ll ~ 11I10l1resonanoe srcctrosool)}'~Q invcs~igilLtC Ih.e Il~ern:cli(ln !bctWI.'e1'l GC:~::! ,me] GrrA t For.
<i

~"",1iDnat UtiS_e ,or ai~(!jJ~es. 756l!!lii:f hf~ RolId, IJf~ Xi@l'H]l: Palt. ~inI!i l!D'2~D'6, di.ina. ll.ilbcH~ 01 MClKuLa! and (elUilr Bi~, Hebel NOm'lal l!IlmNet1Ity, ShUiMhua~ IHete'I 0500l6, cm~a, ~S~.te I!:ey
Z,hoIi1gQ.lJl<lnam

L~~

llia~it!J'l

Sdem{e5., Clillil~riadllilf1il ~'i¥. l00(ll!M, Olim. "'1'0 \~hom oo~~e~p(!llnl1fen~e illlIOuI_dI be addlre5.'Sie(il,
'llullg~m9@l1Iib5.,~~~:01

III 1\11lt lttlyo.;TIl!togy O!OO Bil!KhI!mI1~!

('.ill. m 11lffl~Il!l

IE',rnait

IIIbi.ptJJl'OSC. w~cX'~m;~d >llldp!L!n lied fli;.~flmI'dIl.,ml O'R2 ;md OM 1 rrolein~s ill tt.lcteria I~g. S~ l',. Jillis ha vilro~o;:Sll.y dc,wly im.l.icm.ted dmt GPA I is carab [Ie 0'1' i:t:indlng!O O(,R2:. 'whcl\.-'__ sllch as, ",,i'sion. 'till.-;lc'. ,1!i~d olr<LClIOIiI in anima.I:;;, (/n,iIO\I"cVl!r.~hr;: higher pbnl ... Imbi(.ft~t,,"i:~ ~l" no bindillgn'Cth'h~' WlI:S dc~ec!cd bC'tw~l'!n: t~'1(~/kmd h.a~ only m~c cmiollDic:d Go «(iPA! ~ (irA I '!nul h(wr!1~ !l{.~rumOJ!~bllmin(]3:SA~ Oig. !\llhl!l!.n:~l, n" GP ~ubuJ1i'n. 1~!~,cl o lwn Gy !;i1J~Wln~ls, Sl. A !I~d 131.Tlr!c .dJi~lI.n(;iMionb~l'!di~lg con, . '. . .• . -0;1 (I!-j,61, ThR,; ~ign~I~~lI!.bCC r.lr~h~";Sc ~l,.Ibuli~"'<I~n. S~Olm (.K>jJ) to(GCR2 and CiIlA.l I~~, ~ '>: U) M p:lifl'i[ Sy:<lk:lliH' i... pt](J!r:~'y !jIKl.Ctl'loi.EOOd~ o~lly oti~ 1!l1g. S)('~

~I.ollm

I hC'pl<l::'mlm Il!lOJ1ltmmc ill e~.k <JmYQ1I!; orgalliisms UlIl J'hcGPClR'''~lcdiale(1 siwla~illg t]al:h'w<lY p~.ay$ :~ OOl~r.l~ role inl \' ilJI In\.1;ces....~S

17t2

REPORTS
~11.1pl!!rocl, with the ~cmTlll\'ity o1'~he wiid"l},pc seeds (F~g.:m:~,;Ve"lw fmilud tha~~he :'l~d~in~ , ~ glW\l'~h in ~ibltlnn ~l)' A13.J\ ''I.'i18 ~Ii!I:IS~tlnliit1II!}' p'hmt~ (r~~,..20), TI1'C ~to~mla clos... rl; dJ:,f~~~1l i~~ 9f!hc fl,.-d'~Icr;d!ll ~h~ ~I·.J Ili!~l.t"ll.tll" bl,lI ~11'i;:[f,'~'l~;d !1 ~~II.I"!I;-dUI~r1~\'t~cr.wru~q I(h.~~ ~1w-1L;;JI.v~ ~ "-'"""'o!l}' b~\i,;m)~ or~.h~il: !.ll!t;,"mb:t'lm(,;'~QI;;"!ir..@.tlll1~. f,Q G(]~2 nV~~IL""~}~llo:! !in..-:~ (,.."on~p.:1n::-d with th?J.t nhro;; gp-,! alldl;;~·~mlllp;.!rQ;! wilhnh1lt!:!1 wi I~..;uyp~ w~ I,I.>iLi"d :1 S"pt~~"l,Ib;ql,lil.i~~ sy!i1r~un {!J) iI1:-;1,,:'~tdtv k~., ..w&J~nJjJs WIl1J:,~r ~Q;'i.,'" W~1S K-dm.'lC-d~I'l (J"['Rl~~ ill""l":Niiig~~n!..~ (;('IU-(:;';\·~ in~cr:::liL"'1:iul~ y..:~'l."'~, or dl~ wi~,cl. iyp!..~ {riEl~ 2C'} I.n tilt.. ;1h.'illII~t1J.,:: or I.~K~ .iin ol'these ABA, !ht:. w:t:: ~'o,,'{lli~g~ developed !m!TIlla~l}" O\'CI'l,.';1'i:pl'J,;;~il1g plllul. k-..\'~. (lig, 57), ,1\11 Tbis '~"S<l}' conlhm.;d the' !iei'M'l,1it,ldin'Grm;:~K:i'n ~~d.~ ~ndic-alc'!hil!.Kc:r2 r!o!'Hl~~Ul; i!l.,.t1J):,~ilh'c ~n v ~]iD~ 11.1 1I~~!.'li~~h eR~. wi~h GP/\J (/9~, ·;lI1d ~1tL;: .u~d \~J"C ili1clj!.aingui~~hllbk :lh'Hl !he ~, i~d,t)'pc G ABA. wli'I,.~i1~ G(:Rl-cwcJitx:pJt.';S,.~ing plmil"; me fi.ul~k:n.~i GCR2 ~1,I~ournl~ilI,,~cti \vil.1l fiPA ~ %'I(.':d~il]g::.> lFig_l(,~ ABA rQlcdi~I~~phula tk:\'dI( fig. S.\I.A). TIi~ ilil,,:'I!3CliOll, '!II!8i>fIOO~i:;;hcd \ii'hL:~i O~jb1.'l'ii1l byetmlruUiiTIg 11)p."~I'i~iti\it" 10 ABA.d~ualOfili:.nnujl]g the inI.~h,~ t:1I:~~ .. im'i of AHA!J:l~{' l~iiinLiUi~ oflh(![l.,''OC;pl,01' was IJIC!e~t~.hy \!C!I\'CUlil~~ ·of(iCR2 ill am imJ~iL' .AE:I\..n:::Sl)Onft~~'. .. IIU~rUruko,d S~ne~.m~.d we diu:> C('in\-P2iR~ the ill, r.r"oMdt.q"),l.i.•., ~xflrre£.:.~kll'l A3.A. Iliali'ker ~I:'I\:::: h} wildi.ty.p~ lli.ii.iil!i the ~Ibii~iuhi:n rn:i~.tl.::~l[ (CubPLV) !o!he (' 1lI1digu:J plants, Throe VlI.C'I!"kn!..J<wD1 ABA nKla'~cr The 8,['1-2 l)i!li1ts ~:dl~bit 11111 l1.OWtl1 maJeK, k I~mliltus. orGC~~::! ,(rag. S4i-\). "\Ie il~:;lOilbulI,a. Ihm All A dcfcclS, ]'iJ}C' (l ~ii-i\Inu:1 ~ 11iIliia ~ll II is: (xJm ~!i:I1.C$.. RlJ2'O.·~. KINJ. Rad AJ]L'i. \\,\crc h:stcd. ~he C tl::u·mim(:. o!f OCR2 IC:l:.oo-mtl' «'Ii1t'linil'l~ Th is ~n:ll)'llb CO-~l nmlcdillm 1JilC'C'",~~ressioH of <II1roc1c.l~clh.c in ABA-indutc;iJ sma1lo1t;hl, CIJiIcnrn ~he lroe CIOOllinLl~ amI tlte I~n::dr.c[cd Ihim ('yllX""'i'r~asm io~ ~ ~ntc,nc,t~d\~,'jlhGPA~.. \\ h Ct'(l~ ABA ltlaJIkcr SO::HCS. suir.1t'I!~li"lI.~rer~oo ic illS and i;nwmd K* -chanael re£Ullmto~l in gmn:ci W'!!.\l cc'H~ (J,n E.x rlt:.~i{Hl, fl<u~!;:'m ,~IlI<J! ~)'Si.<; .~hHWS in ~Iw!ti.·l'21 mJlm~l:;:aig. S6~. as ~]KO: N 't~I:l'I1im!i.~ {N!_~l'i:?l' (l~' GCR2 did nol lfig, dm! Gf'Rl ,~I!d W:W J .'l1!;1~ <l \I!f)l ~il1[~ilarO'XI Ti~~ ABA-indll,ll!e~d cI{l!>ing and ABA5:4. B O\:J1d l imli~uli'!!il !]~,ul!hi: C ~l>nilhlW1~ ot uri~!ilb:it~'>d, tlpL;~in!i:or S;t\llll!l~ilil .ore (:~~!'i,~t,di\BA GCIR2 is m,le(~""1!I)'and R\JlTk~i"mfor iLo; ij,I!1(;"'[Oi!C'" W'r(-:;"'~mp,uli.ln~ W!il" S~"A .md, B)"v!'1~~ .~L"io .~i('l·n \.. 'i~h G:PAI. I.'...:.arnim.'<i ~hC'~i.!nC'~iollill~~gnifiCllllc(,'! of (iCR:!~ i 1:c.~!j)OJ1S~, \1i,!cobsC!l'v~'d Oml bt.llh J\B,A"il~dm:cd ~lit'lHW~a (irA I inWll)!Clinl! b~ ch!:lcking: Ill!.,,': ABA re:simf.Dl~cl!l!af nl!(I!re$C~'I!'oc ,oomllh.:ml.:!1~ui~111 slml'Ul!~<!closi!!g and ABA·hll!ibi!cd oGlII.)llliu!!ll Wl,.~~ i.!fI:'i('.J!f;!~tin!' in g("'l'l tonlp,ued wilh spo~l~in IIhc g.t'l·lgl¥~I or O'~ W'L.\~rJ:~~lf" of W,I~ u~-d In d~\tl~~ dw hl[\'n"~1ioM bl.'l\li'L;'t."J1 fiCR1 :)IJld (irA:~ in plam eells, (iCR2 m'!{! GPA I ill~ £!t~·l.~~~I1-.,i~fv \\ iI.rn-'typ~, )I:'nli~s i~- 2,D fllld Gnu or GHU ill dilTc:~uu g~I1I;lI:~e 'bi:.l,CkgffiI,lVld iuyi!l1 1 W I!ISilllg s-wrn:B.ml closureas an :l&S<ly.\I!lL! Jcund lhi:.ll E:1. mbi:l [Ill: Sm:rn~m. \\Ii.dlih is larger in g,l'I'lpJanu> ~l;'I1I:CIt~ .~~ .1.bfJln'dQJJ:~i.. prol~'P~US'!~ t Fig:. IAl t!llIri If! \viJ:d-1Y~}CpJrllm:.(F:ig.2E). AB.A regtd~b.c!>. (jPA~ w:rus hl'\!oh.!..~ illlJollh A:l3A·~0J][mll:l(ld: .R.CU.lttlii, of ihe C t(~iJIniU!iJ$, :r (lC'IU ~(':!f'.ll)-'!lII) Ii~ o hnwfoo. ,K" fK+fH} (!hITnlld~~ ooJllroJII.hc: stmml:l]I.I to ~Jooljs.hcd t~lll .imrnu:um1 (A~, I. A and B), Till tiS slormlmi opcl~ing. rrmd cf.osillg {f~g.2E and fI!."l. St(, C to E). 'file defeel. exhiibiK,j by gf'd&m! 9pefl~ Ie (11, 131, To det~nnill.c' \:\;~l(!II~,;r ocr-a itlcli.rotcd I.I~ lite il1~ermYlio.n bct,,\!ccn GCIU ,uw GP:A ~ is ~~(X':Cifl.c nd that Ihc C Icn:ninLls ol',aC IU a ill ABA·an~ucc:d su:mlaul.'l closure \\':1S simil~1f ~('I o'lCdiJlCS the AJBJ\ I'CBulatioll of .K\l C]):;Uillilcis tin ~hc deice! ci'I:'hib.ia",'lI !by g('l'Zor'J!,J'ai mg, SOC). stonmtm.1 :QILlmd eells, '\1t,'C' appl'tcd 1l.111.dHb:1fI1ril1g is 11.O:CSllalY for illsintcrm:~ion wil h OPA I.We also ijq Tmn sgcnic l,ine~ G VCrC1<I'I'l'l\.~i ~Ig OCR] {If (]PIJ! cOlllirvned~heir in ViVoD'i~~K'''l.,c!irlllby a ooirlua.m~ [~dU1Jiq!!c... 1:0 a!Wlt)I;I;C tile ·whoLic~ctU K .... currlJ[lPlli,;lcjp:iln~i[m lII~!Y. GCR2~~~dJ(irA] ,~ ~lC: \\IC~ hypc~c!l."i~i\'c to till1;ABA f(:jlPOIl,"C; l1Owr1Cun~.n llllI;JJL'lJcells from wi td.·!:yp~ and g('}'J~J i ,~'.ams, ABA il1i1li~hl.>d oj.it> eurrems illwllkl-lypc K C\l'Cr.~hceffects caused ~y Ihe 0\1oC'~expfCssocm or c-oilb1mllFl[IJ)n::eiIJi!moo: lind we ceuld dl!l~ct GiP.I\ l in the i'U'mmlJocon1J~lc.x p:!1C(.~pi~ll~oo w idl ;[!_UD gll~I.!I cells bm had ao cfloct on the KTc. 'C:l!.I"l'l!nt:ii GC RJ or J WCI'C crcllClldcllton GPA l 01" C IE), ill M,'r]~f guard cells (Fig. 2nl~R!S Dl(.hCtl~!1g ! 1("R2, [~IflCiC~~\idy Uig. S~, 130, ~.nd ~nd~ ~Jlli~)' ~. F!Ll\:GrWllu GCR1~Fli\G Ir,U1$gC!!~~C pku1t::; (F:ilg.]('1. T!l{lj~, ·L"l:."'l~I~!i. fJ:l::mrnfour di!'iUII1!;;1 [h~~ i!ilvoJ\,~~I~!C'I'!I. of 'GCR2'I~lOOia!l!,Id 1<,+1[11 ~n e<U"n,!;Ih:1I (iCRl and Gop,\] Illm;lirm 1.{JgCn~l;,'r ro j\B.A-.incllltt~d smltl~IM clo_~IlI["¢, We <JI~!\(~ tOtllld II01IlS.uIll!L"C the ABi\si_gnl'IL 'l.~!'i'ly;' fill ~upp[}ncd .ilIe illll,.~:r:m;:linrrlx'IW(iI;'n~ ri~~~,l th"1i.t .'t•.'n'!. cil:h1'billS d~r()Cb, in :II~ Io:.rrlffi\'Jl, lhm ::;;1'1~t11Ue3. room (~CR 2-o\oll;;!t:xPICi>li,i rllmt:. Ilg GCR~ tLllldi GPI'U .. ABA n:::~rKJllISCii,rufldi l.hllt GCRl I:;; ap-JIl:;fam cloe;lUrC To <lHl<iI.lyL.C t~1C: fU!iC>lim~ of GCR2 ill An~lii~ wl'il:chypcr:iCl'iI~itli\'c 1.0 ABA-induoet:l tt(JJ1,~'';s, we: chfllrncterilzcd l~m,; hU]CjJcmknl A.mhillop:.:i:!i Hm:s hnubo:rilllg the Inmsfi;rr.oo IFi'!!I. ll. IPl1\!j!~ii:alifi!t~(iC· DNA inscnioD~:S i~l~he UCR211ocli5. f,t:r!~1. I~on ootlWoon GCilf!2 illnd

The cQlnlroniy uSN JloC'l1~tl.'i.VQ"hybrid system does nQ~ worn Ib:r ttgq '(JCRl.~(iPi\~ ]!lW'!'~~io:n

or

or

am

.p:.cI'2~2.or gal-;llflg. S5), TI.ill1Sr::.rifJ~. nalysis a hy reverse· Ir.mscrirlion poJym.crnsc ch.ain~\l!''[lcliCi'.n n~T·pcRJ' ,acu;(wd 110 full~lcngrUl GCn2 i~un:scri\p·~ il~ of the dlJ:oeC tmHmn:lI!c:~c.s,ln.n any m.l:~ had lhcIJ'li.IIH::a[cd 1I1l1nsc.ripls (Ii:g. S5). A nlhM(){J.~fs S'C'Cd$rcqll~ fe' dc:>iccal~on Of
domulIlJ;:ylo

GPAl. fA) IBil1J'l(!l.ew'[alr f~oor@~oo ~omPrk!;!P'!~fI" tatiolll a~'Sdys lndicilong the- firlleradion of6CR2:
Ilem~~gtlilln'rrin (HAc) wU~n
il

,C:~emhihill liA tillg, b'IJ~. n(lt the mrrre5J)omiljin.g lwr.. h'II'Vc~lij!!g, The fI.\C'Shb' Imn'''~MI "I'i~d~ C·,t't!fMijrqai dRiitloill '00111" ):c :!lt~l,Kt'G(Rtm:~ .(M~~~ ~yp~ ~~11;-dZ'ii~~dQI1l M,Crmin,1Itc; hn'M.·vcr, f:at:;;.hly ! ~tb GtPAl ijll i... o. AIJIl)', il,m:J'''''C~l\;-d ~l"(:d:-;riom !III Ilhl\il,l uUi;k" or S,:'f.:l} tlll.!lOUU til n~:sl[j!!fII(@. Th@ Ii-FUl.Gi AiM .I~' 10: .r-Iwlr·t:;,gt'l"uniIrrulcd wc~:lll,Irllkflh!.: :'<illillC'l'lCmdiiliol:1 £G!iI~12! boar r~rre'Se:nl:5. 2.0 ruAi (iC~ O~ig. 2A), indk:llting tl~r1'i the :;~OOi\. ,1JrO:fll !"J{'r2 ~llm. (B) 1h~ GCR2jHA" :Dex + un'L1li1l11IS :I:oot ~oc,j a.omlflncy, ·nl(:·~d ,d{)r.IiiIl.."I:IIC')~ V fCI:,CiClfrillll:HA-V ~pC~ is nmaiUl~'y C'Oll!rolltcd: by phylollo:nl1.0nC 1\.111'\, a IndGPA1:dl~Y'C -Y'FIP Th~s result :Silllgg\:i>tcdilmt gn"1 11MJ1IL1IniS 0Il,,-c protl?lniS wcre il?~ilf,~S;edctM1.Yllt1'" g,.Ft.Ji..C; dcrectivc i~:I AHA .\i~gl~~II~.lIg. mhere[lQtc OC"iI~ WI.! iilll Ute Ilroto,p~.. st 'Pm0Cltl1ffi.. J.".Qb. sml~d that lite tmIlallimls hl G['n.l leadt{) [I ~i!ilsfrll.f'IiI A~bidcp$i:l: nJ!ll' qKd,li.t (k1!:P.."'3sed .sensilivuly lo /\B,A. We C';~,'lt.nii1oo IlIF(ltIQPLa!s~s~xp;r~SS~F!g ofl,ler ir,J3:Ares:PO~lSes ill boil'll wild-t}1~ and g~rl GCRl:lHA,-YF(c, GICR2IIL:HA~¥fP'c.O!r fiP'A,lxlMy(-,npN weU!e'eXt!i:I,Lleclandi the S'amp!e' wa~ alilal~ed Olil plillllS. Woe Ii rst !Checked the· e:lfect of L'CR.2 a m2%, 51[)y-poly,a'C~y!an1ild:~ ,get IIItocrur,oplfl©r1~i)!Jet G(R2 or GPAl W~iSdl;:tt(lifl~ witll t~~ !,jl!le' of iii nti4111l! 111'LlI ..ttioIlS (In )1Qt-d gerrninOJl[ioll Tilfh~iW6cnoe ' [or 'GClR2>~V,rCc,(If GCR2IL."YFP!:, ,t! r alnli-c-1'll\y<: for (j,PA:ll.,:Yfip"'l. ~C) C.QililllfJlllJlnQPflJ:dpil1l~qoll1l{CQ"ltll a:5s.a)'$ of ABA and tbund 'Ilm~ ;j,II~1 t~lf~1<! J.,~.,.2niUlI.u!l~t' vllrliryiny the ~11[-e'radon bel\ll·~f1 GCRl! ilrrld G;Mliml pii:mti!l. TlJl<Illlwtlllim, Irnm \1Ifild4YPii1 (Col·man!:! ~\'L."~1; in.~c~..,.jli\'C'.whcl'C<toil the SL;o~ 1!;Plll CiCR2 GC~2:IFI.AG til'''' rls;~~tI!ic IP~l1t'S ~.~ ~iS~driO ~ llllvi'fQ C:!;ic~IIP "ntaoo~~e~to FILAG. Imml!f1;Qpr'~(~lPi~.t:ed wlllill O\'J;I1;.Np['l"'lsorlorm!.:.';.'i; Vo, ~rc IiJI'IPCP.lI" .. asil.i\'~ n mlk~ pwlll'inli li'i!ere prdlJedl w~[hal1libodies (~,to GIPAJI. ami to IFIJi.G. An ,]In.'Iii,.FlAG, cfO:5S-r'e'.'lc!il'lg band is sh®~\l1lI~s 'I:he l_o.amlli'g contrr,iI. ABA-\inhib;'~t.cl] ool1i.p;ll'Cdw~.I..h till:: wild.-iYPL:
Il~Iil\'\::nl
11I'CJlltl!W~C gcr:rniml1tio:n

rCl.P'JU-''ii':FPI GCR2·YW

'0

REPORTS
Friig. .2.Thif' gcrl mJilill1!s ll.:l:hribiit .alU. Iknwn AlBA d>e'ffllru. (A) llh~
~(!2 $eEd~lese t~e Sl\led
IciD.rrnl:lflQj' ~1ls@"''U'l\ciillll

th~lHI~ld typ@ (Wn. FiF@~hl!Y Ihi1i"1.1'@s~~(j~~dS '~lerminn(Jtfd on w.a:ler'"

sooked ~i!J.l€w paperr~!" .,

diJyS. The sCilte baw reP'" re'Si~nts 2 mnn. IJ:) The ~(ll' .see!d'i!:.are riii$il!iiimiv,~
toAIBA~inhiib;ited saad '!J'ElIrmlifld,tio:n (mean :!::

:$0; n ",4). The ~em!if'l,$·

t~(J11l1

P'HO~J1t1!I9~ Woll$i

dererrmlilOO 3

\\!llh (mghi) QI'~~ mill aert) 0.3 ~lM ABA ror 10 da.y;5, G(R20[' IGCR2 ~~Pf~ lire. The ~le ~~ ~pr~f:!~ 1·00t ~D)Thi! ga2 fJ/,iii1ii~
1II1I~i!

I~ro""ingl ill 112 MlI!r~ shii~e and SkoOSI ~~S) rmedilll. ~f) l1fule5!l'.'ed ~f10 l!:i~ntin !JCl2 is im!1rnil:illi~bJ, AiBA. Sii!~ were grn'lWl O!i1 m MS

days amter

npe,UlR

- 11""

'"

,._1~jl.~"Ill"

snll·w de'reds 'ilil AM'indl~ffitI ~OIif'I!Q!ail clmliJ ~ ~~I'l .::1: SO; n '" 3)1•. (Ej The !jrt2 a 001gpal pIia[1~
QlfE!'ilil$fillsitive

:r
I
I-

~
1-

+
.. ..
<

~7
+

I
~
I-

~ ~

1-

I-

1-

1-

I~

1-

-'O~

lID A!1t!J!!,
1-

for slomaital il)P~mJtlf!~ ~rIilearl :!:; SO; (J ::;:. 3).

·t .,.tIO~
-Wfo'll'l~U -20!) ~, --WT··.!:!·~U'" ~. --jj;>'.I.HII~IIIW. _~ •• i.l'III!l~. ·.mlt: .JiQQI

'"

·151]1 !I.

W~, W~·~ile~5kVp-.,(IF)
Theinwaniil inS@PlI$l1JilIie 00 AlBA. 'IIw data in ilia graph (oo~.omJ' OveJl'~reSS(H
nl1Cllllbrnil'K:

in

(}C'I'2

,gu.ardl !tells 15

tt

diC!!1m1@~.

01

n~.:.1

.f!'l-l pM

~,

'ntJbtD~ 1.m~1(b~

.~

,Z

....

.;[11"'9' .11Ul!se<ntsd i!1i mean ± SiE. W ""'! 4. 3, S, ailldi ·6·fer m~.d type, \'l'i~d t.vpe+tlIBA, gu:2·1, 'lI'Idl.!/c.r2·1+fl!BP!, of,(i(R;2 {onme,rlI. iii! cl'ea~d ABA ~elrlsm\!'ityf.or S~OIlll<lIt!! l d'o!i1!J re (me<l n :!:: SID;~ =:H.

r~poct'i"lilly). (6,)

release Go: and lhc GIlY diim~r. l.htiS ~1,cl~vm.iIl1l ~hl1m i\,13;A repressed kn.·Ze..\pr~lon. wh ereas d~ d~jt.1SID."'3m, Sigl'lilHn:g eVCl:US 24)1, To pllLYsio~o,gJcaUy~flll.cI:~VC' ~biillll;>or A.!~l\I~ boA!]A or Ir8n~".AB:AJ did nGlI (~1g. S,I m, Asa cmn_ro~. ~CSlwhenhcr ti~e hi Ilding 0 r AI~,A w UCIU cou~:d d ~'I$'OCiiJJ(e dlC O(,lt2-(WAI cQmple1>i.we used AUA did nnt mlTcct the CXllJ.'es:sioJl of Ihe 1'CpI'l1'lot:1i gene in .. l:I~ml~l~.u~W K+ AJ:MS,E I ~K.-\JI} ~I~ ~J1lHt-ub[q'LliUngy~tcm ItO rcC{)11iSli.l![elhi::1, KAT'I pmJl\~ay iUllhc ~m'm:: pIiN!l'.l~qyilhJ gy$~c.1ll S iniaiilll ABA .slgnil!.~ingeven! i~1yc>!.$~. A:pplkmioll Illig. 'S I.!] h in mi\;l i~i(l!1" Cl'<pt1~J .. n {If ~ul0Ih(;'r ~Iw nf the phy:s.iologic'a~.lly active I~onll 01' ABA IIV;)-ABA or (+)1-ABA.J~m;'l~'lI.'d di:>ml*;~ ~h~ I.WONl:"I{:Iri!Cr gcn~~. I-US; ,md A IJEJ'. W:.JN GC'R2-Gf'A I. inl<~l<;lclij!)II~,'I"hcn;;.a:. 1~11; \ phy:.;io:~()gm.1t."(;":I..LL)' .rt~r),(;,Lo;;1 ~mlili;HI,im~ ..... " in 'l, ~fl'I,i";CL{m: S[!.:L'lZlOOfl\lC':iJit<Jty of (j("IU. hind.i:ng t~Ji1Il~ bio:lbg~mdilllll'l. IhG' m.kHlion of A.BJ'\ :;'Ub~l,ullnfd Iy .. Dc~~lIy illla{;tiw romll::; or ABA ..ml~Og.." ill !-ABA . ·AHAI ,lid !~Ol;.tTtI~t the il1~i,;rn;:iion be- rep!rc:iii'icdi h~Ill:\illi ICm!ijQI1 k.dl.), ~i;liv~ (~-)~f\.aAwas tesed in ~mpoe~il!(l:n {lr~:mn,... )'e;:I:lOl ~""Jll,m."'~~g ~W~%"llG('~2 "1~d1 O~N\L i-ndk'1llJt!1id in i1ial,i GCR2 !lI.nd GPIH. bJ;.li ~~idl1o~ <J!n~"f,;t y~~l ~""''''UY-', T!'l1I1-...ABA gild ,t )--J\BI\. !l:r!lI!o~ dk1. Im~ . F1m.~,ir!'_WI;:(}]l ~}irl1/,':>sirl!loIKAT l-·.tu\T.~ sy~lt~'m il~ OOiminl,.lllnrn;:ei.p'.i.t~l~m:l !l5::;llyl Fig_J E). "fhL,j ~~mll~~ !hr' dt~ G(·.~-billl;!ing :iiilelFi~ JB~I (Pig. lE,I. Tllgl,'!l]~C'r.l.h(~:;t'n;~uh:> illdic!lk1l1 L1uII ]l~c ~'Pocili~~ bil~ding oW ABA to r~lri.ficd suh ~LgJlind~arly 'l;Onr.llfiIl~ ih!ii 'thC' hindiubg of A3A 1.0 Gt'R.l ~s:s~.cRXJiSpi.:d fie, GCR2 oo[JI'd IE i'>idliIlllMd \Vuth inL':re~I1.j;i:i'Ig;al:rlit;U"j'lD:;; nile ~n~~t. A.IJA. I.Jfl repori,.;r gene C'x!P'D\,-& of .. ton was ~pc·dlic :Ibr the ('lCR2-G PA1 ~ntcm.otilm. ln the ::.;pl~l-!Jjbklui.tiln yeast protdn :iJntcmc· ofAB·f\ I~ig. JC'). ScaldJafd. plOb mmlly:;i:;; C'x~libii." fLllililcr supporling I!IlHLt Ihc biibdii~g AIJ.A!o tion :t;;say. the dO",I~sll!\,'" In reporter genes a lilJC1lLrp<litc-m f Fig;. ]: D). ::il~~.csIlllg <I !l:bl£.!h~ the GClt!bim.lillS sitetbr ABA :in GCR.::!, The eC~Lli~ihri.Llm (N IS3. ,iDE}. and Itj",Z) were :Ie! iVOIlled ir ~hc (K"It2 n.~uhs ~nthe disSI.XiJtim~ two <l::t.sa yed l1~ntcins i~lcr<!;C'lcdl wi1h each oli~cr GPA l 1.'OI'J.lplex. di&i1ociation cotlsl.:mt (}rd~ row t~le OCR2~AUt\ c'Gll1pl~C'x is 2(1.1 nM If~g, 3D),. "vidch is CJ;m~i5tC<J1~ ~2l).WeL't<!wIi.OO dlal ~le binding of A..Bf\ 1(1 Tocrroslri<ll pianm!'l are irmnol'liiie and im:a['l .. ble im!ulliI!cnl.d ~1c"1nd~· GCR2 eouW rcpJ1~ the c•..;;prCS15in.!l af reporter Q~ e)ic""pi~g ~u~fil\,{lrable ,i;,wn . ~\'hh ~'!C' ril'l}'slu!IClgica~. C'Q1!~C'n!mlioL1 "l~gC !im~ s such as .cl:rQ~g1:!t at l;Iojd. ]m;(J.1'ad. ~~.~1' :n;ly .(1 .~$lIh Gi ('!he di~~(JJl;i,I'~io'~oriCR2 f ABA i~1l~liil:.~ ~II'.,!V'C .he:':!! c{'indi~iom:s. U~.u'KB bindil~~ to the ,(i PC R iadlQOC's.lhie amI Gfj!\ I. l1~i)t ,~rcdr.ctuon was V4:!ti:tkd 'I ~i~. h~,wil)' tln ABA W .... Thus, ho~",' 'Plant~ perceive .1l1lld t'rnnsdll'C'i<.': Ihl!! ,I tl]h}'i:,itdQg~rrn~ I}' active d i~oci,~~io,m (I flh.c G PC'RJ 1'!1~~i!1. OH13J le\ to S UII.''!'lfC' .. dditio!] O
I'l)CcptOt. \¥C e1(flnni:n.cd \.... ~lhcr fiCR2 h is an AnA rr~'OC!r, .For jhis pu rposc, W~ filSl cheeked whctll:1" OcRJ: 'COli td biud AB1\.. Wc observed Ilmt AHA bound to GCra:~ut d~d nOI blntd In dClt!:tullI."!d GCR2 p.lotcinl {Fig. j.A} • .As a .poo.ilivC' <lndlilC,galiv>c CQllunl. AB,A a~so lml.!nd 00 rCi\.~l~Ltm! nSJ\ <lr(jI~1\1(rig.JA.~I. Th~~bim;Un,g ! (lrAB·At[~ GCR2. pn)!dl'! f;) dl;'fI'I-"!ld;illt ()n !~n ,11K! 11111,;JP~i!lfl~Un ~JI~~ b;,<~W~'!l 7 ,() ,~1'1;! ) (fig. Sm, { li~ 7:...'

a«

or

,_s

or

or

or

or

~~n~...,,,i.~

or

REPORTS
W'iig. 3.. Biliidrilll~1 O'f AlBA ll:! G()R2. (A) AlBA lninds

lO Iheplllrih":e-d IGC1F!2 pm~~111 (m;~n =I!; SD; no ... 3}. '~B) GCR2 ~indil!'i~ (J,f
A,'B,Ais s'ler,eo~,p'eIl:Uii( ~!lii~ Ii!:!::SII)), AB.A IilJlfldin;g tel' GCR2 b!!haws ,1lI(:(ording tO~tllllFi!t101'1

«(~

'~C-l"-'" -II! -€....).oIth

~(=~AM

----~

fd netlcs,

<lIndl Ilulnspeclifk

bJnd· 11'1 s are ,repres:enl<ed by g l'Q,P anrll 1~~~f'!I'I 'O,ii!V~, r~p!l!eD~l~' (mean ~S[); II ... '5). ~D) SUPi;lr{! ~t ~Jlailys~ ~rABA t!in;d[!lgI ~5hip \~
,2:

n~ s.p,~dn(
O!i-~o:-. ---~11---~~:":::':-'_''_-='U=::II::-'

:ro~lI, :100_

--=::L:::_~_-'I

!iCIiClU :

:ShUi\l!i <lI l~neaJ mal:iicm·

'~ne'(OI1C1~ai()ncoeffident

tli1e' squam 'llr

~.99',and ,1liIilX!ll1num , i:!~iildiIll9~ 0.8 nL rrmQr1 protml wiiHlil ,iI(jj ~ 2.0:1

mIM, (liI1ieilni ~ SD; II :::5). ABA di~fiUI=Jil$ the 'tiClF.l2!{SIP'AJLinle,a.cllofl iii! ~~ I(]OP) OrW oil$-

(lin

:say. Aft€1'OO' ~~M ABA" \V<lS

addedlro, lihe- me(Uum for 2'00 il!i!in, I1JfC!i:f.linsweJ~ iSQlated The re!~iw ill!too~ilv ,~d Ih~ 1'mmlllimO'$igl1~LlO I!h~ OOFiUo'l is
imHi(atedi byile
iii am-

II
1,1

fr,t'~ '~D'~+'J<.~A(dt)

41

.'

"1'

,I

f'

'1", '"Jt I.,j' "AI ,~, ,u, It", ~ AB. ,bQlmd, I~O GC.R2 OnD~ Imo~I,'1 pl\oti~)

'u'

b E!lr be

[(1)I0'Il

'e!'a(:h ba IIIId •

WB,Wem~;rJilllblklt (Ei(n~ tOLm) '¥1Ul 51: '~re'Mha~ilt.

The pmtifferati(Jn was

:~

7' ~

and '1 mM ~1I1«illniine,

l!:IiOO1oJ '10 ~LM~BA

arm hOO,-

(Db-

~OI'

i\_a~

'GiC~N!libG

NIIJItlG, ublqultlri N ter· IIIlllliLJO; t 13 G liI1iutltiGIm. 'C;;'P~:I~O!Ib Tlhe SC;;ItE.'bar !lEpres.er!rl5 2 rmI. (f}SdJ!llmc: I'IlO!!J,NI.lIbG-U5H

+
U•• UJOiiU.lIColl
I!II."
t~

,e-lfm A'BA si(:gl1la:llPi!!!y j:I~th\~,y, G [lP. 91i1111flDSil1Uli di'phc5p,~are; GW, glJali!ooijll'l~ ~,'-fJipn~$PcI1~~; FlU), phQSph(l!ti~ase[l, ABA !j,iiW~;JIJ~)'i,.;~ iil!Ki'lHw;m'll. ''LI.!~'J';li{m. ltha .. b>C"(n :;;iK"I\'!I:~! d:t;J~ GfAi W01S ul'\i{}h'<ld in AUi~ ..•
ulli:idi;_ucJ ~~(}m~Lt,~I~);,";Spotnn:-..;,f~n~~~t\l,-.d g(;lImij.~ nl;JJ.1icm (10. 21, ;?5~ .. I)u~.ti;~'~ GlPCR GCIU ilill,c.I.~Le1.Cd w,ilh [1M] to Il~gal.ivdyregllllllc AUA ~igllilnfll:ll/9~ 'lfNciIlerw descdbejhe chaa:· 2L,~criz8l,io_l'I (l,Ua new tiPCR. OCR::!. ill ...I'rt1r)i'~ t;~(Jl1.'iii'!. \Vc or.l\ltdtl(h:iJ~1U~ fiCR2: is .,~~lll~Qr AnA ru~1:lPiOrml du,~bHt;i~ of illc :l"oUowing cvideuee: ,t J,) Los·s..<lf-rlillt(1l~Ofl gn-lcx: hilJi:tsrlU k~lO''!il!';nI

... G{JR:2:-mledlial~ldKAI''1''C~lb

:'j!ji!!iiIH~gl'~l!I!igR!i
.. j! II~I.liJii!!'i!!~t!ll! !!!I

"-~~'it' I".. 'ii'

1:;II1'Ii".lll..Iill.1III1

~!i!i

",.!.'Ii III II

1I!!!~lttlll'!t!!!!i!!!!1i!1

A SA defC{1ts; (i:i), au (w!.~'C'!lpl\,"'SSO\l' of ,OC~:u ShfllO'l'S an ABA.hYl'loi::f:!iClJsil~n:' phel1o~ypc: Wi) GCR2 binds AS,;\ wnlh ,J higll:J!lnni~y and rea~om.!l:ll:edissoel >!ilion consl.mt$",und 'I~leb~nd ing is ~ICI\;lOO~~~ 1k':,lnd ••bides by ro;;l;p.m kiru:liC'S;

!:I.'ijgcF~hc AI~A rc"PO!]:l~';;; {fig, G(,R2 di~n~}i.~ Gr.R2.-GI~i\ I in''l'~C'~ion. Nombil:)l. i\;(,'t(l!~ ~l!1dQ{~ mIlllU~.1~:S .f!!i:II ~liSj1lIIY i\BA !"~;';;IX'I:L"Il,.~, 3 f ~.lIna:s. (l1J U" !:';~llll~~ idClnti iil.:"d .1Il AIM rcThis ITmy be; ,~ill!C ltl ~hc w~..i~. n~'1111.r~(Ir Ihl; ~l;;l'~Dr 1u~ld iD:;,I;.Iu:g,,~t. ~:m'[~ii~I.'U 11~wmtl·iln~'l""i,t: G m~lk~l~o.m;' :55) nr n.lll1t~:ocm3~,l,"<illndflnc:y widi Ii!l~ f protein. G(·[U...cbtooprotc:bi.'>, gi",(;ul that [J;IC:L1.i !lilC ['0'1"0 IRe,1e~e(f'IJct~<!!IId N1~te~ all:ICf 0'CI(.1 htll~lOlogoLls g_e.ifbCi'> in tile Ambklop •.,:~~ . 1. Ill!. fiimM'ISI)~irn.,S, L GOlfIiI1ia1~,C 1)._ R9~"I"fu..m'(t'liV L4, genome. t'mISb'ltf~t. with [hi:; lauer po.~ibHi"ly. Si5 '(20102). til£' s.t.'ill~idornillm:lli. pltt.'iWlyt'M: :lnd 1;l~lriia~comp~~. 2- 1. lliilmJ' J. Gira~D~~,. ,Mi!I\(J.fkv. ~J11~~, i!f!1l1rt NW, m!:~'1L"lliO:i:Ior gj,_~~2 by GeR.:! eDNA '~xp~i!)J1 flkJl4'\l, '1.9'9 !l99S,), 3. l. I. Scllroeder; .L M, Kw.ak~,~, J. "'llIe~, N:Ofillf!: UG; 32:,1 (:Iig. S I'~~t>llggC'::.1:> tiJlcpo~ihlc ilnt{!tifcfcilcebc· aoo:U, tVi'Cel:l OCR:! and il IIOfllOlog..-;" s ~" l. K, lihl.i" Ati~~, ~; f!1()rrl 8Ji,,~ ~:\. ,;!~li OOO~, l11!! ronow.ing mood ()'~' lBA si~n[lling crurabe A 5. F. A. R.atetP" s. [HYl~~m.)', S, lit Al!;.ram~It. 1[1, HijL clwi~ged, i\ GIPCR,. GCR2. which i .<1! pl!t1s.ma s NrtJrJtt !ll9, ,Z90 (£OO6~ ~. y, Y. Shm t,r.~I:. N(J~ 44~. 821 ~2006l. me!ln\.1!rollHco Ipc,!li~oo ABA rec.c:pcor. in~~ wid'i

the ,:"",'1-':

(iv I OCR::! i~ ~o~d 17.I.."d~ .he p~a\imil n:i~I]~bITI~~~:~.cSlII~~s,~n i du:utk<l"~ ofuhlO G Blmlcin arid dLS5.o1(") GCR2 !:lC'f1elroCa~ty and plJyllie~~lIy iiIlJ[~~I:tus cialirn'ionhc Ik1Lm~ii1L;"ri¢ cOI''illplcl'4in~pGtl !ilid {H~\Vii'iSift!Llfi ABf\. cr~ w·hh GPA I; mit! (vii) thiC: binld:ung or AHA. to d'H;' CiI3"lf dil'ilt:'1" !o ~ld.i~'lIt(!'

the Ga~~f c(ll1~Jilh.:!\. Bundul1,g

nr ABA

[1;1

GC~2

7. 8. E.,AndefW!l, l~Ward, ]'. :s.:ilIr<oedie~, PJa~l ~ :I;(fl!. 1!.177 n?'?A~., B. 5;. GllltI'I', It lom~~. ,I!/I!flil.. Awl. 5d, U.5A8,9, ,1591

9. E.·Ie';lijjn@n~e~' Ill,. f'ioJlr f,.11!J, :U ~I999),

fl9!l'U.

REPORTS
11.,. A. i\t )llli~~I,L .~. f!in;ll!!il'l\, W
(200)4·~. 10, Ii:. L. Pi~r.Q~. It T. ,remlJ81;. III..1.. leB:cMti'. f.eil 8\bl. ~, 630' {!NIO~.
~11. ~,

~;
~,.

~.

Mol.

55. i1

1l9. S. Pimd~" S, 1'1\. AM.:m.mn. 'l!Xli Cdl 106. :1,616 (2c!o<l~. 20- s. IW_ I\S.w.;ln~ .kknw ~lD. l!l {020051,2.11. IP. 'Cil!irdm !.If'M". Pm.!:. Mat! .1I.Md. ~ci, ~.5'cA :iDl. 122!!1:i!
22.

F, ltiilllllin far Ih eir SU9[Je'S'lio1'!5 '00 th~ ABA Illl1d]f1~ ~~~. lllili~rr.ull: w~ l.'!I.FIl*!ed by Ih~ vru"'i~Ul' or S(t"!llltl! <!~dI ¥!(!nli'lC~~'l',01 (hii'la ('lim l.illi, 200~.M~!1.fIlzj}), IN:lI~oli'la1.
~!i!!lf!i!

12', ", Mil, M. '(OIT!~~k't, M.. ~~!l, E 1'r.g" Nll'rl. A.oIiId. Sll'i l/'sA 8" .~8:21(1'99i1)), n, it Mit, Pwm iIoll:IL !li!llL.26, 1611 U994). 14., Ii\,. G- ~'~!TI, J- ~g'!)~fia, ~, 1Il~!t lI;Illd. 5<1, uSA '17" A [;'I"eil 12(K1!ij. 1:5\, Ill. Go ,fIta~1lfI, .L ft" Bot~la:, BiO!hfm. BirJplip. .Acta lLS20., 1.4? ~;2(lOil1!. 16., A. M.JcmM..i A. li.!:!Il1l1l11m, ~l'b\O ~ s, ~i!2: (2M~l\, 17, '~ fl!ii~do~hDvmQ('" IL)"11l't,I"!1Qo;k, II. H~!I(~~ (mf. iliQI', 8, 315 ~'li!J'9:lJL 18, '6, C~ti~ci,,F,Apoml!,H. ~hm(!rmt D. (1'I.IlImer.s, Ill. F, C1111i5p~15, PfDr., NaIl, #Ira/. Sri. U.s.A '99,,41'7,[i6
~i1002t

smn~l:2~12,2jj!70 (201H).

(2004), x.~, 'Wa~g"H. !J1lID. A. M. ]t;mel', 5. M.,AiW!~n!'l,

fooliindMioo
{~!!I I..!y.o"
I~~;!fd

II~ Ctilil'ia fCif 'Olllilliinlilillg VO~llgl

SiJje~~.I~
t:t)' S<nk

23. II i!.\ W.i!rd.. Z. lot

(!1';l!)!S), 2.4. !i. It .SjlG~

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3'002 5C12,~1 tf;j~ (;~lnt:iIO: iM<!!~on~t f'rcjm {la, W.W,. :lOOtl,C8100'!'OO'J.

~Jl"

!. $~hrael!!e<r" Pl~'1Il(('11 '1, 83;,3 S!!!IP,f1Ol'ri!!!lI1 COI~~!il~ tI~.i:lt~i~!ll

1I!MW;5l:,ll!OO'!m.;!.g;~@ilgilcOii'llenltf'lIl1ill~lla:.2l'llCl N!aien'<l15<!1f!dl ~,leiti]iXlil
:F"II}!O,.

AIIIIM. ",~, Bi'1l~Mm.. I!J;, 6~'i! U.<jI~:1'. 25. s, riinae'j'" I. G. {htn. A. II'. ~ooes, :S, M. Aslrnilillll', Ram P/l~ia.l. ~,:l:. 243 ~;l90'6l'. 2(i. WI!!Ilmm1: ID. 'Ii', SlIn!or hi! ~PPOIi'I; l(. W. ~tJ'l!lilmd II.M. ZlI1i:I~, ~~ 1~~ Q!fI!!!]lN~~ ",mil di:'i~IlI~~~ W. frn!1!mllr an~ 1', OOrdih. f(M! md;ll~ n~ 'iIlm~II~1iI ~~ Il!iilll~mt!"; ]\ ~1.a for iiI'I~F;;iJl911Ite'~plil·'lfFf s\">tl!m iI!I'.ailali:i~!l'; 11 Chili filii! f1is tec!ilIn~1 ai'JiMal1cII! anlil ~d),ij(f! irOr UIi~ il~01t!in !l'llnflGtifll; ~iildlliJ.ilIu~n(i!, 'If. SI1I!~, :!Jilt!

51 ~

~n

IR~ItJIlfI~I!'S· ;I 'Ot1l;J~r ;ll()>ID~ i!C(!ep1!ed

:l!4IiMua,ryal007

:Ptlmll~!ied

Oi'I'liI'I.~ 8; MilI'oChi 2.001;

1(1,11261f.{i~m.1H.s.'Be2 Imd:lld~ thi$[niCitrl:tijQ'lI 'iIIi~n c:li:il'l9 Ihl~ Ili'!.p[!r"

Tunslbillity and Noil&8 Depend'Bnce inl DiiH:,sf'8!lnt:i;atii"D,n Dyns,lmi;,c,s,

'Giliird~. M. Sii~l,:lI. R(!Il]3il P. IKuQk.!!!lr~i} JOillditlllo11ll DlliliUkil'!l,ili jloll"d'i Ga.rci.a-O,iiil h!'o;" .Michael. Ill. U®wi~:l~*

all ~cil1~b]~
SP0I:L'i~. m'J1!,'l.

~yM~!~l

ill w~~i,~h.u Pl.:'hLliw~y :\!"l1!~dl

1lC'-

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oilly ir! .. IkIc'~i~J1 of c(\~L"<lutl

f,of

as OL~lllrs\".~[ha~~i(m 1)!)1~l'Ili;I~:' neu.Ini '~U!n~]k: if. 12), ('ml"~CI!ICC (.:\1C!11L>100ClU, (

IlI]U)' 'be,; dfiv","n by Moi~1; in Wlndl.;'rly[ngcin;u]u~·tlmlJOll-.:lnfcS"-I. '1.0"1..13), T\\1'[) key ctl:;Ull". elcri~:u.LCS of COMl~!l\Ure m).; its .

Tiholl y~mk d
'CQliltrl;l\

PfQC:~S$(llr 'd!ifh.lftf!ti~ti(ilf! d~p,~d~ Q!f!th~ arr1:hit)ljl(tUif1~, .q"hllntit@~~\toI,l,

P1!fQ!l!II~~~t~!1d

Iii!'ois:oe(Ilf ~J1ld~,!1Vil'!g: '!3~f!iIj!~i~ cilril:ldllts;, !i!~~ev~r, il~ reillilill!f!S; l,Iililde~~ ~~"" the~e or;: lelffl,elil'l$. ,;;:~ml:.ii!f!e '~y! (;!e'lhd.a:r~;II~h~vijorr. w~ ani! i~~d the prQbl~bii~j~:(k;;lli'ld~raf'l$~e1iiI~ d~ifr'erel"itijati(lfll of B-odlluS' $ubtjlis (jebil1t~, th~ :'Iiate '(Df wm pete tu:e.. ,1\ fe~~k!~ paramdef$ ilnd'eI,perll;lelllt~~ turned Ull~
freql1l'iitm:y of ~ifllitiat'iolll 111111 d~he d I1Iration (If ~ornlpetelill,('e 'e'lPiis.odf!',5 almlalAmlJed the ciir(lI~t to'aCIESS lCiiHer'elllt. dYfllamk regi liJiIe~~i dudi 1191OS(~llaticn III. A~~ering dr{~l'itiiI,r(hiit.et:.tur'El'shm\led that the n dl!n'allo'li! 'CDf UI;mpete'n!l:~ '~ve--nlsc:aln tie ilt1i!de m~~e p:l'lB(~$e.W~ u$d aiiii ex~erimeli!t.;Ill nii~th{id tl:l' redlm~ '£llo~~loC!elMalr' n,o~~ ;.inldl ~hllj)~~lro l~at rtDls.e le\leL~are corllela'l'erl \ll1lth frequ:el1.C:Yof ,d'iffE'!r~ IililiaJtio:n eVilllili'ls.Teg:~lhr, the data reveal a rtD'i!l.e··d'ellE'! rid illl!l~drmit thalt. is rema rkably ~!I!5Hie!nt an.d tnalbl . eilil tenmi of ~!S. d'Yfiliillmk: l:Iehaviior.

wrob3bulii~' ,('!;ril1i~lLi!ll:ion. Pil~r, r£lm:i the n.It!.""SlI~'um,.. d mkm cf Irm~ie.nt '!.:.'0111IJ:It..-"<[Qll(£ ~'\!CIlIS. ~(~~!P (,Fig. IA), r1!!O!l~dcnotes!h~ Ch31l0!' per cell d~visioll t~at M r I:1d~vidiualcell willbc\,"OlllC ooln~n. Cm1il~ tem:c ,i!\'ClltSCaIl be' '1mllltilicd by autom:[tcdI31ll£,lfl(loSC'

tll~on::S(om:c

mic.lnMXlllY

l!<Sil'l,l;l nmlllL'f!CCI1I,

~:)f1rtcr 1il).\~l~L!!1d\'\rdw o."'lnttul of !he C~"I'InKK :qwx:~li(; mNJG PltIJHOI!..1' .;;u~ 1U10'c.'m~rSpn:mll).'!{lr

(denoted P~jC
kh,nlinr

,md P_",,>, :1i'"C:51'!(,'Cti\,'c'IJ)'}

li1:;.Il are Ci'i:,POC[(,'(! to

(,I).

TwoI11~mI11CK'fS

,tlr~c~ dlC

actions "UI'tOI!.,g ~errneEic c~ci1l'lenlL": qMfmlit!llivc' pamI11l.etcU''118illiJcs.:mch ~spronmtcr S[li"Cllg~.hs: and stodmsl.ic nU,C~Ii.mlioll:;" r "noise," asseeiared o \\']111 the ,ooll.Ot.."'.Illrn!OCl'Ul or eellular coml'OC1IlCnlS,

dY~flln.l~.e: ocllLdar

llI.'C iII.:.. lL: ..TiIC.C Of. g(.~;n.CliC •... I:' C'i~Ui:l"'" CO•. .. 1 nl.m bchflVlflflS ;Ih~ eneu II

flrchil.tt:tlU~~ pilu\:'rll or

or .reglilato.ry

.i]uC'(·

O'Wlll cxpR";.';;si.!iJll but is (k;grod~ by I.he qpp:. C:1:r[' •.M(:cA~{jI(".1I& cOlnp']cx. tf ig. IB) ql 91. CmnS e\lllilpclilivdy i~lh~biasilt&:. dC.!lf.l"ldal,io.~~ and l bU('1J)~ luI C(Unpclcm c.c~Is-.. linmi i1g;ii, 11~.gati\'c ICo:!bock klQ"P'{i/. 10. /1 J, The droll i~ operates 3S,
1

of ~u:: CilH;:uil ,m,:'IJl'1; ~gJ~~ C'xrP.'$S~n1I r,Ii1.Qi,or t'NW.i( ;lnd UlmS. {Ht,;"IliDll,id by .~. ~lIldfl[s. If(::;pL"CI.H'!oIC]y !Fig. I B and Eq, Sol)" To !~lnni~plI11~.le IhCliie p!lreUilfu'1(:[,£. .• ,'C c1iu:f.H~.lf}sfn~,'lfUllyhItcgl'l1l.ad. ruJndtlli~iollol co;r~Y Ch~icr{'(_mt.'ior coml{ under

Ihe c[mlmi 01" 3n~ltdtlcillle promoter, dcrml.cd. P~:l''' .goe.l1Ie.mlitl£ the ,llyper-o:s 8Il.d I~,.wcr.~.

or

.m!~n1!..·nrl. 3)_DiiTt,'i"~lq.iudQIl (2, in1,{l COlI!J'M.:'ll,.'ll-CC" i~ ~lf;111:sk,]t (fig. IA) {4J. ]h:: gl;!l\;1i~ b~l~is fLI.llhis ~dllwior i~~I,c.i.l:t:lI,il .UITD inlg mmK iUiilm,'nS ,,0:11\'

A li~l~clamcnt<lllDiolo,gkal, question is how Ihcsc three 8SI)CCiS of genCl:ic cil:Clil~ls emnbine 'LO dCl'Cllm ilt\!c:cUuIOllD" bclmviolf. il'> v:lria:bilily. and lts potcn[~ ~to evolve Ol. ('onlJ)C~cnCt in 11. -'Olj/Jim.~ 15 a "IW)1S, "CSI)(ll1SC ~ ~h<Ji~ ,1~'I(m;:j, I~ .. tf.I Hlk.e 1.11.1 DNA IU:'l)~nlh(: '~n "i~ ~

O'lg. ~tH,lIl,~tmlJ:;ctiPli(l~~.lil£lo:r'C{)m.K..~

11~()~~

s.ary ;)l1d s~ln:ick~lt 10.1" ifClI'Cfillimiol1l~ilto OJI1i1I~ di pa,Cllcc (5. 6) .. ComK. [r08 i,ti¥ci}! fUJlllrc~H klt~s ilS
lOreen, (en~r Ili~ionror ~1~rn~ B'iQ(ogy ~fld ~iP~I1I.'· memi~. IIf P.hil.rm1!oo'OOg~, U.miwr.sily (lfli~:n~ 5Gl1JlhWl!'!.'lNflo "1l!dicai Wirer. Dallas.. 'TX 75,390.. IUJSlt l[)!'i'i5iJml of Bitllomi!"ld D~~im!en~ (,i' .AlJplllli!! 1fiI~~ ~~fi!:N'm!! illl$11Me tM rEdm1O!j~. P,~[I, CJ\ 9tl2S, IU&A.IDEpolflIDe1fit of rlliluoIbl_, Coll.e§eof ~~Id!ln5 a!i!d~. IGolU!liIhia Uf:!j;. ~'Y, ~I!W furk; m lOQ~2, USA. ~~ent.de !1151U i
I:;rHJiJliyffia

Ttrn~

CC!!vm

it [-oon2

NOO:!?olIr. IlJIlilhrEflilldt If\otit@l:nl:a

d~

C.dIl3'lurtyot

TeJ!ir<l~~)paim.

·Vo \~hom oo~~e~p(!lfnl1fen~e!llIO~_dIbe addlre5.'Sie(iJ. i IE',rnait

m~lawili@{..alt~h.Mu

IFim. :Il.. (olmpe'leml'C!~ i~ (!II p'Qball~U:5.tk ai1d transient dm~rel'!lliatiol'l pr(ll~ess Ir~I.II.ail:edlby " '~en~.tic: (ilr,t::l!it. (A) Th@ 'Fat@ (jl~ @ftil~ri i'lg th.@ ,(~ m,p@t@l1t:s~ate 'rro!illl th@ '!,I'€g,@'taitil,!~sta'~' ~s d'~,not~d: ;by Pljiq~.1!I1@ a D:!,O:ll!ilt of time SIJoE'!'IiI~ tlile ,compet.ent S'~~tei ... d,enatedl by ";';_11" Tlhe (0 mK tralil Slri pllonlf,a;c:t~,r lilil (I)ru::eflu~l'i~n,~;s hig~ "!pink w~gJi(lI1l)' whu (\~11sglre (o(Im~'~rti1t i:Jnd ~~~ (glr~IN11 r~g'iQil) wh~FI they a~ glFewi ng: veQ!!latfvel,y'. (B) M~p ,of the (,\Ire WI'rB pet-ell,(ed rCuiitl)!. K.eyr-e~tllres im:lude positivI? trilni1l(ripti@l1a'l iilllIwregl1llatilllil oiff wmK' and il negaltive JIl,edlJaid; Ino,ll ~nJ ~liIhridl '(ami< ll1&aibit~ {pOiS,~~b\V!ldhc'~y) expr~Sis~~ifl ©f Com5-. which li'l~Um lrloorl'eresMth '~'~.r~t!@tirtlfi of 'C-omIK;1!1lt: ~ glf"lIph:s, !below the p~ !lind IIl'('(lmSpromGl,illlfS d'e,rilliit ,par,]jlill(l'~er.\i, used in thE! text: E~preslii!:m Irates dla Ili,ge f rom ~jIK~cn l~ilIl1'1d ~ to as rl1!:spel!:lirlJe~.~,ilIS'Gal'iflK l::.on.(E'!Iii:trra'lio:n 'illiclI~,,~e~ ~mi fig! ,coII1,pe~e'fIC:e'.

REPORTS
,~tmil1s.fl:s~tti ~"Cly [see 'Sll~~I'lIOfl Om~ill~ 'M.:.~ il!lg ~CI'l"~ ~S'OMI!c."'!I. 'ffl s),I.:5!cmmk:,'lIUy scaa ;11~!Jlg>C of V;1~tll;,~rOr ~'I!;, \llld 00;, W\,'l ~I!lIIw~i!llC·l:iilPi~ Cmi!aili.~I1gd.ulTcl'II:ltt CO;I1""''ijl~r.ii:ill1n..; 0 rjjfl~ imhH:l::f i:;;;opmrw 1-~l}-lh.iog'llac~(lpyl7..JjnQSlJdc (] f'TG) ffig. 2, A 1u~dBJ. We in v ~s~ugmcdhow orlr;, ,a.!lC. fis co:rnbin~ ~\[l,
~guIDlt;Pil1il.\Vhi~OlUt I]Jffi.f1'll nlmo,,~m:;~

\~c

IW;";,l

co:nsiid~«1

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en~!l\

<l'r the

~lC

illCllC1lSillgas to ~~ ... l~s"'"[~111he~: ~r--(lI;5 sirnin 1

~n~vi;;:s of

d'bl,.':'it;

,"":'.JLUL1,~ 9:!ilIlW~~1U

I~yper..ti", !i,milt. liirn~~hi3i~dl :fOOilli lie. Pi!~~ ~ncrerli.~-cli<il"id~y Wilh wsal LVllite ex~~ssion (.Fig, 2L')., htCI'Crul~ c.xfID!..~~ion !',Cam,,I!; O in d'l'C' ~1Y1)Crc,[lH\: $lm~~l"I~t1:d~o" 00 a tirml)':,ii[ioi1 to I an (lS:C~ 11]<110:1:), res,imc.i 11\';.~Ilich eells 1'q~Ii!'lb/ went ~I~ IC OU~ Lhe oCOll1Pa>el~lstmc I Fig. ::!A l" ru of Al (~&:; -2)0 limes ilS. \v~ld~ry~~v:~h)!,~~ t~~"I. ,1111 (:cl,l~ ~n~~'!\;i1co:rn~~t~f~('C(['ig,2.. ..\ <lind C)" CCi!:IJ1s'lu~ CmnK, .il'li m,'(;~~ry fOf cl1iu~p:;k'r!tC'. I. 'hl\\'C\lCIf. c.x,~ion of P'-!I]"..('wuS' Imd :l mOO~N~ C'fK'Cl em ~n~ti~u.Vtlll. .ir!c!i(;'<II!l.iv!g'ln;~fl'J<...2fm (~i .!il, f 2C)l, UnJ"" PinlJ ~"";~ pn;don~i!1"I~~ly R"!ilul[n~dI tl".y
ali: 1".,11]:101" lh~ll
11s,.

innh~ ib< w.i M~!Ylr VII~·

W3:';-3%

lp.!)C~ rll~g: dw ~nflllj~r.~ty ~Q~np;I.c!1tQI,;~l')1rrom ~ rel'lt~th;rti,Qn:il on 'il'Q1I!rtl' Int;:w>b"ing q~ up ttl - 4.,' ~ fill{ '("~ c..11,1j;~~'~L1 ir1~~'l!,.~ 'ro;<.lI"ID' (fig, 2!P'~On~h~ i ~~ ~i.ling ~~U + 5% t'~= :4!M!2~)1of C'O!'lpt;;:'~.'l~ oi'I.~\,"rhll11d. [hi; I-·~Yf~r-u~s~minl.'~ iU!l;n,."Il¥i¢d in o;IL~ ~?;;iix,;~"t:ilM~ S<I}~ rhi,;' I:1;ptt"l':ib3Iity Lh~ w~lh irmt~]liil\g cxr~';S.o;;m:i t)fr~I1'~'OmS {fig" rnullu.raIPem.i1S l"((m1,)11:1: ~s tlIlllsS. ~iI1i[licn1 jtJt IJm~n: .. n."!illlng i;;,...d~Lbilil.}' (" ~ rr>;"Sl.ilI8 ~Im~"" Ih~l 2 D'.lIl11t.-;;. f.cOlllll ~~ 'pralOrni~M'I!.n!~y regulall"d by CN;d!;!b~l~r)' C;U1 be .~'(:lil,Lbly ~l"iJil;l~n-cd (}\ r;r 011 llS m!lrn';U' lih.. ('l;K' Tog~~h'r. ~hl:,.~ an fl:~IIIS ~hm\' tlll!lltPi,jL <InRi TOO(I'\Pe~nl~ hllted i!~d\::pcru;!~~nJ,y y b l~rnJclItU~igc 0 Ils V;l hl~';.>;. To b~w... U~id~I'S~11Ih~, r imJcf!'tltdcnt [lUI1ilj,g !.lIK ~~Ilid I\.~J~~i\'cl,l' (Pi~, lIE). 1l';S.

9r

or

n~

To beuer lliilde:i.'::>I.ai:ld 'th~ eflcet elf t:amS ~~pi'l;......siot'l on f_~1l!" we CmhlnJ.ctcd a '''(ixS'' :;trnirn im whi,c&IP""i'i.!I~c('UltlS W::1L~ C1(I)~essed loom a. Im!Hl:lf.'Y IlWlI1100r'I)!i!::>midl. c:111..'C~ivcl)llillCii)C.1. ll£ .d d'liC' ,)!;'I!i,,'m~'(1 IIDJodu,Clio:l1 rnlC. I~s,by a. rh'C'lJOr of 6 0"11I;,'f ~its~~~ Id-tYli'C \·.. r!l~.3;;,1N!~ UnllkePJti1~"",MNS. 1 bs rOOJlslmCI 1'I<n.1im:. ~h!!IiCglilr;nim~ r«~~ldin [he wild~lYIfX'!.P~m~ 1)!tIlllO~L;r:. includi.ng il." !1Ci;; ligihlte b~~ 'L,'XrJG'l'itl~1 ~.JII,,·, Ills,",l, D~,;pilW'~hi"illt;fL.,'1INC in ~5' c:l!ciu.b!c bdn.wuOJ: \\'a,;'" mainminro. a... 53 5% (1" = 79Jt51 ~Ori;:(l~npCICn[ cell .. :>LQ~~lI~ly
I;lXiKld~hc c(JJ.n,l:Mfl'\!'~ll. ~nmg,:, Co!nlfl~u!tdWilh M% ±
=

oI' Pi'l~~ ~'1HdtQl1I1~P' aswell n~ rcl i:ilbh: "i'lmi~l~enflflce of C)(.CjJfLlii:~il:r. \lVC developed ."i u~lod~l, of she core' hnemcl.iems hl the cmliilh:l.cnr;:e n::£u~alit'il1.dreLi inry (.riS, ~Band SOM teX:I.). \:\1,= !1!~d stochm.'>lic s.Lmulmiou:ii.IO account lor inIril.1..~ic o~ of bi:oc'h!;;,l!1lic,d !r~':Ictron.~ 14), \l!.!'c n ~ a!:so .l!!lil!.Iy-.oc.'d the ctlim$llm~d ung (':Qn!i!1uoutlvml{~~

jI"A'd/.'

iIlYI) 6h;,f wild-l}'iIX' eells, By

tll1ntf:<I!.~,

ti I£~ d't~!11mi11(: p'1r/wm.'[1,;'.f d~lJiC!~r.!CC .uld ~ll idL..'MUi '" b~.u]:o1l~t",ully ,\,,..~rnlilbk: f)' p'lr~Ull~~',c'r n:!1!:inw in which Ihe di!'oC~'>C'[C' rmxlc~proou-ccd re~~d.,~ '~"o!ll~i.(~m ... ~·'ilhCX,!lCrimcr~l",.'WC roquirad dw ~JiIUilnL'{l'lllii1f1odd ~il:I rerm~i!~il~ ,II!~ C'.'i;l1;u"lbi~ ! ~:g;JI~~~~111.I;)~,"PJ~WlC'WflS, "'uril-cl by ~u ~u;:lnrC!r6

Fiig.2:.

Oil(

it.altiiv~t~

an~ Q~ 1~II~l"
d'li!l1Jrg'~ It.h~

,I,
IP~'p IPn s-rtp

Id)'inamkli o,f the (Om· petenu~ dn:lII.it andinl(j,epfl'lllde,ntiy hllll'f1 the p,l1l!w'bitity of frli~alil(ll"l {p!_~ anci mean durn'tiwl h'~!lmJ' o~ (lil'liiPE!'lle,!'I(Je '~ei!IS. (A .and B) ~~nlT sftirjj,ps i3Ild lilIa;mr;:.qp 'I.iffie trace! ffo.rirLdMrlLlllI. ,celW
lOll ,i::ampl?'1Jl!!li(@

1.
2"._.1111

c
1

ij_ '\"_

A.···

.2.

ilK 5

20

HI

!!:v.e!lJt> O. ~I

IClDt[lll1ed iin Hyper·uK

'file IIPfG !1:muelll!:l'ati;D:rl'l ai1dtlii'ilie!; (hours) ii'i~

,,~s(lpf.w~rn'Sl' w~ffl:i .:lI,t

(~~(OO1K),ltld

Hy,per·

'~ fll'l

1\ lii:n:lts)
'IIK

'cllc':Heod. W!~~Jp !!I1iI1li P'm.rnS"yjp a(tivitl~. (Ir~ IcI~t.edm r~lawd g,,~.

~esp@c1ffi:VI!'l1f. 5po m'la,t· 'il'l~ cellsalre 3!eleill in

.~

~-----------------.

\'lI1iite. CeL~ t~<'Jt dId

~pllflLllil!lt1

[I
191 ,~ rF".....~ir,1P

rg~t;

\ller'\'l' PJllI1El' ~o

E) iQIlIarttiirJC[litioolof re.SIiI1H I~bhli~d r~lln11 ~i'1!'ll!i: amil IH~~ stlF.i!iu~U va rio as IIPlrG,(£Imell1lra· t!QIi'I'$

~f5TI~t'QlMard th~ end of m@lli~ a(:(Julisitii:)riI. ( t~

1Pe=l'-Y(p
~'. '",

.2H

'I

cm::: d~pkt€'dirl

red ,1Iif1!:l 'green, lFe:5pec:~ tn\i'ie~y. '(C) ResPQIIlSE! Oil

P~nll (iis(mI88ifl

1- ::.I:_j\_'" . . .. I ~
~ '16
'iiI!
1 .....

.r:
'r '.

'~

{II'! jll un 11:5,'

'.'

ih;

SO), (D) Ri~~P<.!li!$~ (It' ,~ t~ U$

'w (harn'ges:

~11iUK Or

~f!dl (~!((!iTI@C!J! :!!: SO). (E) Indepelmdl'li!t (!ol"ln~~of

P~i!rli,aiild v()Emp,byi(~K ;and (~s ,dll'clil it p.CIHU'illf~ers.

=L
iH.H.:I ..~
la' plinarne"l.er
.21IlI1

.... ~,
.41i11 00 80 100

~ig~t w~ a~ d~sa~~ ~~.odmsti!: ~imulalillrns ofltili1 m.oo~ellor ,c,ompeten.cl' d~allllll!n.

Siho~~rninl li'ghl red a Ila

- - 'iI!me(!io~) . . ~-

20

,25·
i'ctrnp,

301 ::15 (ti1tQuffl,)'

40

vamliDm [iQrnpiuabre

tCll ULOOe!lInilly;!tlti

experim~rJ'Ially. ,tlr:ls~' :5mtlmalk depkliil1lgl the e.fle!:its 't)~ Oil( and rl~ on

REPORTS
111,,"::;.1J.~~ C<J.f1 be exand we fl."'q~lired i~~lOCJmMl1; CQ~lm(lfJ'Qr! to model tFig. 3B.~I~ilSJ.,lh~ r~hn~jas Ihll,n~~: ~nlm ~o~li);!JI!t;r!N(K.!;'1111'j E.",,! gt,.'.I1C'mw Ihe Qmcr'\'«!~!Kl~pt:nde.'~lY!H;lhil'~lY of t F'irD1 .;;tIld '~!;(IIi\", w~ k4~.miiJ!~-d\! p,~!1Jm~KT\~ ·~~fti~. wh~~ ~~ !l!4~~!,l.iJ: <!l[IImlb~r Cml~Sm~~c~ kJ< ~l'i.

SOM l(:~"!l U 5). (o,"j>@telloe! ,du raliorn (1I'c!!Jiflj;i)' (A) \Vrthin 111C' Itiodi;:l.um.::l'!t;;I:s.i!igl:i:K ili.cn.'ti~~ldmh(:: ((lmpet~n(e ,[ir(llJi~. ~ll.aliir~~'ilin~dlD t rruOfLbJUly 1.1rn1ul ~~~l1Ili\'c; udb, n:m::-II the ll1'iing·· il1l~r~dku:ealii a dditlonal tli\illn5(rip.... miU!I.ntQ. «i1l~~'i1r.1ti:ail ofCpmK nCC~!>lla')' 1.0 iUiil.d- UOP1"'llrleg~.tiv.e~~il'dlba<:k Q~l(l LQl;lp ate 00it1J1C1CtllCi::, ~xpl.1i.ii'li~~~b,~ £~OOtl~ e~:li.::,",i. or rGmJ<'~ g~ll!erali.ng t~e Compi'llD,k '(~K on Pil~H lr~~, Iltcroasingtls, O.IiI~'C· fithl!r s'n~!l. (IB) Iflli,l~rn:S(~f!c~ S51. ~1'ii'i~~rc:CI~" ~.L'nd. di.d net mise PlIl!il nfli'iilrm'lily hig,h oocau!ilC' m:lrITIIalized Iby their m,;(ill'u..ImI 1i"alIUIIi!, hom PromIii"ffpin 'Ib,!! COOlIlRiok inhimi{'l-l~ of coml~.r.'1cncc is ]kllli~d hy fllllcm:blion::: S~f.1!]l1ll!Jr 1hil' w]l~ ty,p~. r.l~s i'lilw T in '. ond( C'.xl~re:;.,'<;,ionI~ig,. 2E .amI! 11s- 5:6),. ~ Ibeem.!ligln~lMlful re~l!~ttQ time of i ~n the ~nodc~, w~ 0l11~(1 m1li.l~~1I';('!(I'!',Ct;i!1ip Exil fml'lil con~l](;'tC'l~cc~~lIli~.s Con~S~o~ed~·!J!"dd!.'d, ]l1Iiliil~'ion ®f P:foMIG' ~lI'esl>i:(In!, IF®:r e~:ch l)aJn;el (()il'~po!tdiflgJ disu~u: \;V1Ji.,.III~hL!b.ls.;d l)md~I'L:'uiL1n I::.'I!{~ (I f t"omK is k~ stJl;d~aslk :simllJ~i3it.i!on!),of COO1j)Riok ~hal'1 il'1;tctiu!:t1.>d ~]lUcll!CI]OIl rme ~r~ e: a!ildwHd-lY~~ CQiIl1lpf;!~~!i'I(le dr(~Gi!:s , 'omS dc~roomiQn. <!'!lLt Ilm;~ T.J"'~1fI' i:illU~dr~lC'l1.>d, Cln dw Odwfh,und ;,u1, ns.~"ifu;:~a...oo I¥olll ~L;.m. a re~Q'\Ir'i as ]n.'Se1i5.

·~'iidiabi~i.wru~ ~Iigl'i ~S"illd iU¢lt'plIJrH1..'l1i uJ.11~llbili~y by illS m~dO.K. IPi !iIa, 3. A!r..::hltedllll<l.1..::h,mgeto tile' Amd~i.~ orth'C' n~kl i~~L~".1cril1lCddclail inmh~ MelK$ !ijr!i:IlI"~ mdm::es v.nialbiitily of in
:ilf:~(I(I,lm ..

or

'!,'cry kl\:\'>!iIK~lh!: Nlm],,~ sise 01" ~I(I(h":i!tic IllK!1JI.rtio~~~hl;'re!broim:'~'R~8. n~cll~l~ ;j;ncrE:'llile mh~ \'''!]'I~~'ilY ~Il ~""'" In th\;: ('0l11~)~~C)~ ~.roi~l.

ItJ:r bOlh

nl@.i:!.lh;:n~I'I~l; (lr

I
IHI

r~J

['l[Jn~S ollll~l~i!;.", d~grJd.u:i,O:ll. pru~{jngi,liI~tlJtlpel(lnc..:: Jlinni~], ~ ConSi5Ji~IU \'ti.lnh ,e.xperim<:rlf.:TLI rL':Sli~t. ·~nu s, nile ~uod!!~ illcn,'a. sil'lg 11." switcbes rtil.Cs.ysl.i:m from excitable to oseillmory diYllm~li(;S" nilrlh~r d1i;;!ir:l~ glllisJ~ing at:. :tro~las (rig. 83), Incl\,'a>i~lg Ill!; mkcs !11csy~1CiIlld IroCl.ly from C'xei!;ln.lllity~Q a h-i~~~~e rogil1'C inwhichtlir:ii~ < 1" lilln. once .itlil~ncdl, nlOS~ cclfu.> !1I;~!lilin trol,JJ;l!."i'J i~th~ ~"(Ml~I~!ll sm~~. TIl~ bclmv.ior was alsn obSCl"\.oo in C'xpclimc.1ll1S where .rIO o:"lcibw:r.ybcha.,jof\\";l_S Set'll m imcm'tc-dl:~tc Os \'~_tm,"S~F~g,2:B), As i!'l the experi 1].1(;"nlS" cdb ~ntl~crn(xlL;'Jcan
~I~u:;killll Otn:rI~IJl;l1tL;' _ lM11;;I1 the b;1~.~1 p'rocl,~ICliol1 [jllc o.f CmnK CXOt'l,"1i>1d,"i ~("~ ivak"li U
~>t,;'(Dlnf;

~'HT.NJWH::'~OC,U~ of

10

20

30

40

TEmt!I(~)

p'~nduclioll

m~c (0It:: ';-

Iltd, OmlK

k\'l:L s CfnUlOl

b.c rodlfiXd in compclcrltt. :.mdeciisilGl'X:Inne rmppctI in lih~ co~nrctc:nl. Stille. A sim~lar e~l(l©1 I000J1iS 3~ high ClI.s \1~dues. L'Io!!G."lL1Se new smb~!e it Shl~C mi.9CS ill 'oon~l)ctCI10c-nlil~mai~1 OOltCCIiIill~ ~r.:nuonsor O:NnK ,WiS. Sh lJ-lol\\,'cvcr. iu tillis CllSC. exh 0:1"oomJ)Ci:CltOerellllflille(\ possible as ,fIj resu ~~of noL~, which dk:'smbinzcs, dl~ Ilcwl,Y
~:bl:1]lcd cml~Il(:<1Cllt sum:: ~.'l1CC Scct~(l1'I S 1".:1 of'

b_,C::>
~

W.ltll~

6::::J
,~

~'

SO

[VI tC:xI, ~,This.

is c'OJlsisaclu'

W IIh c:;:,p.;:::'riH'ICII~JI

ohscr\';~~kl'Il'" ;)~ID\''''~ngIhm. CVCIl 31 the

highC's~ induC't ion levels in~J!c I.I}'P~~N[S .~lr-;dn,-;2U%

i;x.·i~QdCfl~~lp~tcn~"i)'.

UrNU)i~Ul,;oI;w.>~i'l.n.ll)'

Tn t.""pl:OlY;,; dw d'lkti:'i orp~rtuabJIJ~ ~hc cin::uil

~l('~lil(!c1 ure, w~; l'l"';flgint:~r(it'l lite 'OOllllpc!tCl]~ circlIillll. 21Jld

IU~
i!i:::

u::;;illgRok. [u

ptO!cil:iJ ttltllt

bjnds

10

P....... R'

0.01

rep.~~ hs C}!i.P'~<;~1U (16). \\le i:r~$CniCd ~ { CClUJ}' o:f mk under the 'OOf1Jtrol of Po:'ftJ~IG•. creating an additional llegal~"V( lC\:dba'C'k IO~1Poeto l\f1111.1(
rf-ilg,3M, Ill, ttl~S "CmllipItu&'" ~ru.irD, a:;;cxpcetcd,

N~l!IIailil!:Q'd1 CIO[~ ni!!!I~lh

~2

~bed

3~~6

,cem qlhl

&

lronl it~ ~1i'ild.'lypc value of on Ill;;: o~her hand, wa.:;;red~lCed. ;IS was i[s: ccn-ocll ,'axiabm~y.:l'n C'mnp'Ro.'k. t',OJ~~ B.9 :t. 3.4 hours I( mean - S D. II - JO l'. compm:-cd w ith 'I'm,,~p = 20.2 + 9,9 hours '~m~;111 SD, n = 3 ~I iu~wiki + lyfJC (IF:ig. 381 Thi::itCdl:.l,cL!l]'J1 iu c~II-Pi:I~V@!ri:lhuJul.), ~hn'Hld 't~1lI1 'lhL; pn,,-ci~ic!l~ .. "" em! of, ~~ iiil'ilpr(j\IG~O¥e'r i;t~'1\,.~]d-~ypc V3~ut'~ 11'11I.he
PiDll ilX."tHiIlincd UI'tdllllUgOO
\.'<iJi..I(.'S

f ~J%~l' Themean

'_!1~'

IFig. I'll, N:oi$~~IJ'l g~Ii!U:!~p~~.n aM IP~6ll:;lhilUy 'OO!i'lilP~te',!'I~ '(nf l.l'1rlrllatlon (PiiI) d~(J~~~ \lIilhil'1!!:f~~ng (@i.~. ~~V19Ih, ' G~n"oolA'IIC!~tijOI1: ln gf.lf!~ @X!P~~~F! .fsh.ad~ of gr~ll') ~ !~peam to doo~~~ ~ etoogatoo W n (:ondmolla~J~W IT1lltant (elll$ rompa~erl wilh ~'I!ild-type celIS. ,qua Noise de~ei!l5E'li ~v.i'tn 'inc~~il"!g cpU leong~h: iD()1;S,IiY;lpi'€Sfti1t flwt,-jifp' ~~ptres'S~(H'I {n~Uredi by m~~ !ll1!d~flgth ()f~f!dMd'~hjl ~o~dif1iional.lb:W fil)lU:aIi!~ {IFlw"ii cell'S, ~nc.ll!ced' \1111111 ,f.IM IPliG. 'OIUIJ ((I~mclenl ,Qf vallilatil;ll'l (C\f.IQ[ 1P~11P' ~~resSitirl as, ,iJrlJJn(lion S of O:1U~.enl9lh illllia~k) (:QJlJlpilr,edM~h diM:re.te sI:«i:I'Olf>tk IIIlOde~. :rmk.tiion ~grilY~. (a O'!lerliay ofphB~El' (Onlra5t p end PCMIJ,.qp ftiJW~OOO~ boo~ ~1I1;;!pshQts,O'f FUiltSOG wUs 'lO'!ilit\hincre~'S~1 (~m llengtlil. m~lEx~r1i!'llli.lil1~l~y d~J1Tlined p~ ~blia,k tllll~' d'rop6 wilhincrea5ing mn rengltl [0Il5istemt \lfilh dmete sbl ml1lstk sim~Ia;t!ioll'i (gra.y IJ11iliid. Error m ~ iii! {(} and (I)) ~l:!n.6t'e 1fin:~511', (See S3:A SaM! te~ Inr details: 0.1'1 ra~culaJ!.i®n of Pru..)

17~a

REPORTS
ilo.~ C'\icr. Ih~ ;);d(]i~i9Iml IW~~ ive r'QL;'(~b.",c~ alllG\,,'~
~,,,,illrom
~;m~'l!!Q~nW ooc~r <llh~~hl,"rCmnS W
!lL,""d~IJ;;ij~~~ !'i\.'!h~a~V.~ypr t:I'~~

11!nd

RO"'oOOI19!,J"]t~~o:~!8,

ltig. SolO), h i~, l0~ klt{]~v!~ 1 wh~1.hL'iI'("OIt~I'M:klto,: ~ili,i!lilO:~1 is oonlmllodOy noise .. ~ iu 'the' model. Tn W1l~!l:n:; hnrmci (1l' !i.OiSi;; ~~ C~~11pt.;:h;;'1t;~~~il~1~~on. ~, ,~~.
l~')'-l1p~O ~loc,h.'1Islie fkn~ltluii~!l~
OI,1l

r~~pgrlsib!e ~brlhc observed \'<!!fi'>lDili]}' iadumUP!). Second. noise !1I!<lY 'be l~:if"!)' lQ main~,.i~~Jo!ic.~I'lbi.lhy (lN~r .. \;o;.,jdt; ]lIlII+<t:~IM~T.lI~]~, r byull.dl,l(:illg ~';S.!I;a:IiW' ihlll~, K'lIIl..;'~ h~,~.h of [·OIiII!.K c(]lt~llht[tioll. TII~rd.nlil)i:;~ appear:,; kl h~I\r!i,': u

~]cli... . i\ld)l ulicl'cffilu,d ~ i flt;R~Jil.iITtiGn m 1.~i!C' s,il'l~k:. whirhlhL: ti':!!W~~ne, wbiel'i ii>iiiC'(,,":O.c'1.:Ibryl:ru' t\C1~~H1iol',\Wi." eell level. 'Q,[j!iitti,tii1ir\'~ 3nruly,~ls, e r!l .gcn~lie' »)'.'i1.~D'i1 bereplac~cl, by fll~inlcl'ueibt,.-: l.'Oi'),. ~n iii iC' Ws~itre il~clHoor. scrlaljo~~Wfi!;, in1hlbjl~d, rf::.u:1 ij~ in cll:m~ yond its :tlOl1I~lllJ, Ol)cm~inlt r'Ct-ii~~C. inelmJ:ing
:-;[m~r'i ~n

to !:llol~!I.~lyl1il1lihlhll~e nlil; I:~IL: c.c:ill. We ll~~!U R suhl;}i\"

alTl!OI,mt

~rn{,'ij!':~irl

ri\,[l~a~ role in OOn1jlGtC-IlOl! iH~tu..ak'O'~~ ig. "'II)} W "I~d l!1'UIL~ ~lo~dd 'lio:;;. 1.""01!~!dl)rocl ~dGn~idL;l~1'1L;[k rm:i~~~'i~liL;_(1;; ;uid ei[Cuil .'I(dlil.i;.'{,~I\ir'C to ~1I'llbip¢'·

or

2;1100 mfill'lC:11tCitIi.~ c.cllt':, Each m~m'\m was o:ml~ ~clte c"'4~~kl.l1 strel'lg111S" cifCUil afchitO::l1l.1re.and noise ~C'veL~. $!tI\Ol~ly COI1.<;;tr.Jiil1s~ ",ul1lca~ mood ~d OrlWLlllil,lc- cell uni't~alii ,~l:l!l'rngc)'ttlpJiasm, els. l'hcoomlte~cnec ~\CS.'L!~atlon)':-;~cmmain" s \Xlilhin <1 IiI"mcn!.. Cmn.ll~icill i, C1>!I:K'ttCd~o cfI.Ccliwly "V(lm~ ccH~"{llr!.l;:nt:,".ro(tL!cing noise .~~~ [,liUlS !CII:ciwbh.: heh..\'iofo\'(:r a bm:ld mug!.': of all( ,m,d 'f~iS ,g~1~~I:!'\;,p~~io,!"!. wUt~~O'lIl OlIK:"Cl~~\g ~tI. ...~~ Ctlr~OI.:·n~ par<l!.IV1lCler v.dm'$;. iEl'I.p.:ri!n~nwny. (: ~alb,!C' P;~;I ,~u!ld Tow,,;pr~~ b!.;" 'UUI!IL,"d i:l'ld(:[l~Il.tlc.!I1l'· ~,I"tillr! .. of ~"1i;lh'!lklr .. 'i. Ctmlp!{'U!~"L" (Fig.. 4.1\) III bl ,~(lmc b<lctcriia~ mu!lal1t!': IIH!:! lra\!~ clonga!~>d iy. ...Uowi~~g~hc s.>'~lem. inlhool),. to udi'.lll 1,(1 indepcluh.ma ~l'cCI~I\' e 'prc~W!fl.'-l' dUli~]g ~~v,oJwiC!l~. f1IlaH~'.I1IClll~ mll~~}i~dJQWc""Ct;lI'~U~i!r .. goo.vlh. Jlltclb·· Th", dn;:ui~ can <ILI~ <I;~"CeSl; di n~tI;llllD dymllui(: oid ~1~lk~tY. p:ln~(;iP1 "",p:r~io.U!.<~ld O[.Il('f rh)'~io-I~

logi~,11!l;h,)u:ale-tL:'l"~\iti,(~ :lppcaJIll{ml1,~I, ~~<in dlU~lg'h cclhlt8Ji" 'I.rolllllm:il:;; gre~u~I'yi:IIICf(.\Ll'>Cd(fig" S;t2} I( 18 l(Jr}. \;llc· iJ:lw~~ctl fin i!Hrlucibh:P~rrJ.&' C"OJI.<;;lru,i:1nd mcasarcd the COM o'f 001 lcngtlil e On! cc;JI[I~U nlll!ctJlli:llaOl'h.... in yeU(}Wnu.OJ'C5IOO11. p'ro!cin OfFPI lC'xpn.'SS~on !fig. 413). ClIVcfound

~\Jgilti~.~u:t;il as .osem'iliO\ll(~n[~ bi~ll(ibilil.ji. .i:J.1· d[c,u£ng its pOlt'mif'Ll it:! C\<{);]",1c' :.a]tiCnlmil ..·Jt: qllmli. UII],\"C bch3\'I:an;;, Rerii:rti!1ct$ and! INotes
'J!. U. A'lcn,An .1Mt~l!IC(tmi ,r.o'.S~I"MI~· 8iarcgy: !I'JIe.i9~ ffil'll!'ii*s of 8itJogJ(oJ (irQ:Jrr;. MaUterna.ic.a'1 and (om.PII!~~ia~t 8jD~ Series" '!I'd. 10 (~.p!8il!'1 is. lli'lIIlCRC. ,BoX;! ~.oll, H,. '.!lIIll6.. ~. 10, 0I!J~l!.i Arin~. i!I'~ N!ior~i~ gZHW:l\l9J. 3~ II!. !)!, G!Oi5ma n MI:i'lI. ~; GMe1. ,'. 41{ U?95~.
~. 6, M. 'iimU

Cerro prateins, d~.IlD!:ed K f(!t" K ~ = . S}I'·~Ii:·Il\1I:l~~ ~kMd i%c para >emg! 'J!! WIIlnittnfilfl!' jJ~dI "!ltl;{~U~led (Qn}K UIM\~1ii~, r~!lellti~~ .. 1!iIiJt:I!Jj~d term IlfWidm rlItgr;ad~t1an lJimog;lll[OOllJ'l!lithot blmdil:l~ 01 C.oro'i(alJl!! 'CIl!ID'~C MKI!;. lbe'~ii!I.i!l lim!! ~eJ!mm.t'5 li~~ irl~r~d~~:n ~I (~m~ 1h~ ~'~!!I~~lpn 5 i~ ti(iol' = Do + ~,rr;-l.,S" If~. U. !rnd fI. IIIltMUrt! IOIfiI!~S!lb ~ti aJm I!lrtulilfl! <I~dI l"e1Ju!!al~ ~ e~ ~ty,·The!hird.a!ll!l fW!'lli111ifms mJl'e9l1i MI!fiI!.,~ m &10 dIl!llfo"!~L'I!!I:I, r.~oc~. WHf'Ij)m~ <!~ 1~lI1w" ~e .~ le~ we hR ,<!lmo ~~ a dr.crt!~, ~. o[ ij1~ 6f1l!li ~~ i1t1lf~~ inllNii: ~ e~llIlad!f.Th~~hn1!ntdr~~1i'II IWlOliIiiIly oIl\!! ,il.I1d """~~W ~.Ihe ~ P;!1l<I~ g~ ~1'I1I11ill~1I'Ie, !i'IlIOOrn' o~ Ill~ Ipn~ ~ .. lb!:. 1iI»lIe!~ ii~drJai!~ II d'diliLln IIIe< S:OMIRlIt. ~6" l T. IHilia ff at. ~\:I1. ~rao~io!. 4;!~ ]S ~'OO2~'. ~7" I\t iB, l!owi.iHr at, j, l!~~~~J8l. ~'il7 '~1.~9\ll 1[11. r, IShi[M 11'~L.}, &.cilirr.fD.I. jJ;j!:t, 'ii~~3 '[U,89"j. lI:9. N., Milki !!o! .ai\.. J. ~U'Jim" ::LiG2, ,4i3i1l1 (~OO). 211J, f..t,~al'l ,N, 0.'.. ~al. .MfCi~I'li'DJ. IfI, ~21 ~:li"2~. 21. H. '1'1. Md!damt A" AD;in. P{M" ~rI. AfIlI1" Sri, !I.SA 91~. S'!A. (l'9m, 22_ ]. Pau!:isrn, l1I(Jful(!! o1(;?',1l1!l j200~h 23. 1':,5, Siomiit,'J, !t 1EIo\!oltz" ~. 0. :5~991", finx. Nil.!\!.AIr,~d. Sd. U.s.A! '~'9. 1.2795 '(2002). cor"ID1iIia~QI15 c[ IComK

:!

~rud5, ~g1~fivel~-

'-~""i~~1

ne ~aHon

iill'!~

rl[i;r-~.,.~,~.
tb~

~11;Lt oise does IIl.dec-d decrease wilh increasinJ.,l n !engl.1l Wig. 413., .in.'>1.'t). A :'iin~plcm[lodC'1 [Ioft"lIl~ ,-<erUIJti'ol1 an~. U;U!l1S'~Hliotl ,(11 14~ 111m im,"('I!1PO" raies ~J.c' con~inlhi!y of m,mIlCIlI01!S eel I gro~\,1Jj~ p,rodl!l.cOO qu{!~umlivc:]y s:imilar 1'C8!.!ir.s {fig. !lB. i!~[ilN. nd SOMI~KO. l1tU~, ce~I:sizc ·Cil..n in Ihis a C<I\iiC fuc U~~Id. 10 m'lxlulgl~ gene 'l;xpl!(..'N'ion n()is~~. ~1m\' doc'S lIl[}i:.t: ",flttl i!K~prob1Lbilli~y of ini~
.~ imim:l (J'.I· 'OOI1lnpi'lt;ltt~'") To .nims\~r ~lli~ qWI\,';SliOll. \vc ili.duo:~d lI~mi.'i(;lililli(JJl i~l Ibe c(J~l~~i[klllful!f,... if l sl.L'1llnm lite IJcgjiiUliHiiigol: or Ix:f~~ 111.o\lkrrieq~lu~ si~iol1. fIInd qLlfll~~i~1cd~I"i fliS Ii flill1iclio:n 01" cell !eng! h. We detctilnlillcd !englll di:>lr~b"lJion.~ of' cel:~::;at tile mOI11!Cnt~hcy :f 11utiailed romIK1iCiilCC" <1...., dC'leCled by PII'Ji';ItGc,xprcssian I(Flg.4C). As ,a compariso.ll, we <1:100 I1lC<lSlm:d leMgdl disuilltli~ liom. for nonc'(lH11)CI~nt oclls m a slmilar d~sl[ri-

9. l!t~!IIr51~Y.er~., .~~ /, "17. '67:W U.9~;OO. t(l. M'I:lIDi, u;~ IJ, ~biilaii,f.&~r~ro.I. U6. 5]5~ j'I'm}, ]1. M.Og,llf,iJ, oIl'lll'l" M~I: ~iNr~, ,3Ia. ?'.l19 (1:999), 1!Z. (, oc[]~h" f/JapiJ,sr,$ (~t$!~ ,(Q;dor,~ 1!)I~n.'_ p~~
O1dl'nd~J:!l99), 113. 10. il'IJOOiIIJ, It WileI';, I'tfol. lMJ\rmtlM.61JL 561112,0061. 1l.II. II).. T, 'GiUe:lpi~. ,. Pliyt. Oe-m. :~,l,;J:34'1l1 {l9]11. 115, Th;e '!I~iI! Illf[[jil~hl!!lm im Fi:g, !HI GUn Iti~ d'emibed, illl <I, toolilljjr'.!~i '~Pilrm:il!llfjlitItri'. by litIQ, ~ooll'1~dl d!in~r"Mth!lL ~Wi!t~~ ilItta~.!lMI~ii 'iN! i!ynamici!l1. ~llii!i(JI o~ iI1,~

5, lit M- I!!leri~ el' ilL •. t.IDJ M'iO'OeJQt~~.U3] WllJQL 6. 111 Vi!nSi[1IIlk!r!1'I rl' ,al, , ilolol. MicMlblcL j,j, ~s {l9.9!iij.. "i " Iii ~1l1ia.. ~, M!1I~u,.!Ilp/. it'!J'ctol/lili~l 5~ 6015 IMS~. 8.. w. [(, .!!mill ~r jJl,. Mol. M'fclooiolli. !it 604 t'!:oos~,

~,I'!L.

/'fr)rI!!;e 40!!iOl 5~5 !lOO&~.

2.0\1. M.1h:!I1lii .A. 'iM ()ii:lM~~i't!ffl, ~l.lrDt~~: StL .!it0t '98., aGl<! '(200.Jl)'. 25. Ii'I'e ·thart: R.. I(~ IB, StilJi!j!1l!J\i U. ~ It !brg'<f1~!h!'1. $, ~1Et, L IIiID, K. Sii~~$ wiifI ~ a~lI1e ~.~. IIlI!~JifiJ ~tm'I'MlrnJiilt!d di~ ~. ~ Wolii ~ b:r !J'iIl'1lSm Mfl (f.01 cw.m'W71. 10 IWAU1.0lfI1li GY.&a.i'li~ iIlI ~ C.eml~r ft!i' IMlidiIMr ~, IJTe :s:eailf' sm:.Iifi ~ lrumill ifwrr~M ~, ProgIii!!Jl. ~!'I!!Il!!er.tctlrn !1!I!!IJj!~!.n G.I!l5, iii 5!1J~ed !tit !~~e 01TedmI'9i (~mer i~ Bi;tJgical (mti ~ it!!i:Ill'Ir! ILlfti~Il!1' Tea!,.~~ fn:I~

.am

me

aii!r~ia

Sdi~1! f'inlg~ml ]L.Go. i!l:kn~~ fna~[ ~ tl1lM iI'I! it;nl'1W dl! MJcOO'M r {Enoo ~;~n..I~j!G f1i!i20ib6· llN~~ mlh~ ~~~t Olro de c,n.awl1!fa. SrL!Ipl:)o:rtingl O!!i1il'lE! Ma~ei~al
'lM\Till:jM!:l!illilg,Qrg.!'Cgi!C(M1U!I'I~rl.illBl~,e19.l1lU£lf[lCI. MaI~rtal5iMld ".l!:tti]!X!~ 500 lf~:l:! ,Fig5i.. 5Ol. til'.! ~~]J

,pi

Tiibles Sl 'Ia S4
l!ief",r'etI0~

13 Natr!fllb~ ~~ ,:l;rol!;ll@d .MFMfIJ~rry ~ao7 lO.11Li!~iem~~.lU]>4!i~

bu! iron onhm:s. W:c ploued l'11e rc13t~VCfrn'Clion of' 'C-cM)), Ilwlt inltia.~cl coml'octeltce alil given ~CI:l,gl!' I..
cml1!pm\!d
\",'tl~1

Ihe wwJnlu]'Ibt .....oj" ccn~ .. Ih;~1 : u

~"CUt'icJ'Q1lW~h:;d~.f.i~. and SOM ~~x.t), A. S,1~l1ij~ 4[) lou' d"cw.li"i~ in Pil.!. \V<l!:;,ob¥.it~I"\'l.ll in llQn"1\I]oorl.d~ i:l'Ig ::;,i~mllmi{)lL'" I~Fi.g. 4D). th 1c,.,I.if Ihl.~1It-.,cl'lCliol:1
i:l!b1'tfi'il

longlh. TIl~~!'I. I:II~~ .... ~lL'l!wt~dliml fii"~ do;~'COll'il.>d.. . .m

Temlpor,alll F'lreqlu8,ncy of SUllbthrr'8slho,ld Oscills,lions ,Scales wi'llh E,nlolr!hins,1 Girid ICle,1I FiellldSlpaciiing
Grid ,~e~1l;in ~i:Ill'lI1!r or rill Il.nLmhil1l£ll, ((!:rtl'';. 1ffireto .Gpatiallm:atillfMI; n n <!II rep8'.atingl hexagonal gri;d, ~\Ii~hr I~ sma~l.er spi3d1'l9 belw@!!111 g,riid ~1@Ldshn n@urtllrk"S]111mllf@ dorsal ,alll.t),tomical ~oc.tI~iol1s. nata 1irWllil'l vitf'lJ fIIiIm'Il~(@U. p~ttill rewrdln!!!ls sli1IJweld ~iHr~lm2n(ll!i ]lIIrr@qul!liI(ycfrubtillr'lBho!~d ITIlemibr.ctlill! p(Dtellltiill m.dl~al;iIlns. ~I'l !ul~(!Irt11I'lallillmrOI'l:llllha~. coml'f'p(!Ind todiUllinmt pm~tn(!lnJs a~(!In,9 the dorS<!l·;ID·'If1entral. axis, sup:portililglll m.od!!~.or plh~io!flgkll~. mlldnllll'liSlmS lor @riid(ell resll0I'l~t'.!L I.1Cel!wrhin .. 1 :oortcJ!.. pla:y;: ;1I_t. imroU'l~nt role 11'1 ,em:oolo..g of :SP;"I<l~ infOl'Hmllon II J)l!Ild '!.7p~iC' U'Kl!l1t"l!)' (4), M.my l;!}~T Un~lllW~~ (If !11iJ ~!Q'lorhWa~1I ~l!;x. ~ .~d o;Hi'i. luing !'\!DUo'lll ~l~~.t i;;~1Ti ("~Ilarn<~y =¥,liaJ~u~i{m~ or .
IOiflltilj,g~U1tc.-XnJilOii!uII grily,Jilhi.l1 lill~ (~UDvimliJj,iiitltL

CQuid be dint: to lactors

Q1~I.;r

I·hall. di~

mninrL'l.I1.ro l1o;ls..e. Vi"C1:)!;;)Jinill,OOI",,'O pro:r1'1Ol..~i1i.P,,;mS 2ind ~~.... both s:tIDll,g!ly [l;gu.l\i:ll:e~ Linder ihc.'i1C \:-o:ndit:iOrls, SpoO./'\ is ::Ii I1liJ1stc.rlflo."guL.1JWii spo:rIlIlHui(Jn~ :I, oOIll~pctil1g .sirlrv~'l.tioll l'CSf0fl.~~" m'Ki ~ligJl conecntmiiol'l';.'i s,liC!O/\, iflhij]ii ot:{'II'ltIIX:" ~cno.: 13). Con.v!"'1'SC~y. I?~Jf~'5: .... c prc&;;ionils nt::'t:~ ~y rOl'col1ilpe~e~loo.Mean C'x:pI"Cssiun orbot~1 P"""Jrs andP ¥!<II'IA was lIImUlbclecl b~' cdr ~cllgrh(liIg, 17~. ""Il1b s!'1~]W'OI:1$ tl.•1ll kiea[h.at .~"'I1~ chj)ressiml Ic'Vct:s. .nrc incl~~~n!tlcnl {i f ceil l'cng1l'l Wlm.kf l.hi<'sC ~'Imclitio~~.. ,~~d~h~lP"lj~ dCp~;Jl~~ !iJ(!l~!Qi~c.

or

or

s:

.~itlii!;"];

NC!i~c ,rila)" rob in]

Ilhly~I~l;~l!hn;"'t;
OOlJilpc'lCiIlet"

Firnt.

IlI:lhc

dilrCl\:n~ n.1lf;~ tuuM be

(57). 1I1.C' 5\plldl'lg omdl'rg fields ill 1i'C gait.! Ymlc.<; ~,,'ilh'!llmomDI pail'~lml ort]~ cdl ",long Ill.': dOlsalb vcnlrnl .;;w"!! ~ of~ntVimimil oo~1I.c."" ;~ 1~~I'!;"i by d~II~<; ftTIlT'l ~l(; IQ'!rlmin1!ll oord:.r l5). i\\;ut(I!tl1 d~T Qon6i..: clbl1'\id br.irdJ]!" ,L:lf cn[Qfh~lH~!.~r1.C]". ~II!'e
~1(lJ1(l1'"di~mJ~IL"" OCu.'!\Id"~.n I:ilrrillg li.tM:._, CtllililJllllnll-

11719

REPORTS
~ional models c."'S~licidji Ill\,>dilj,~tmld~frcll,;no,,"[. in grid ncld ~1l<lCif~g :lhn~I~~ >CQn~i1I1Q!1dlfl Qmi:,r ...
~mQ;,'l'i in~rltrin-~i~
'~h;;¢!~I!-iO-VI,;'"llUr.lll~~j,i!s' lj', ,v~, lllii~ 'OOlllcl jlimtividL! ~rst~lntlliit: wnatillill in ~hL~ Wlin C!.W,<I .rnm'C!1n.,'1fIIs~d ~ig!1>l.~rot pnt~~ in1.GgJ:'.t~ion J. 9). (J.•

~h.~ ~~11¢!¢,~ of

'1~~"'[9!!s, ;lirmp,;

tU),. '~Ve~ooml!ld :'o1J!binm:.stH]ld oscilIm~Qrn ]lum ';'s.t~l~<!!~e~~ls. in !~}~crn ofm\ldi,,-I ~n!.O!'"hi~;iI! ot;:Or!I,':"i, ,( !!~. l ~, ~1l,]!J;:I;-:ii, d.i(JT~!tt si fn;:llJi!
dymnl1~!i;S
!lmn~t~~t

ccJb. 1)1.'l;.(lrclcdill s li,;,::.s between 4.9 mN '7.1 rmn

~tQmdlC
~l~~~\'l

(lif !he

oo;r,;.,,~ sarlace,

~.~-qtl.a,;m;y

by

l;1:s,inlgWillI

IfxJoti'~~ln]N lB.I'cm~lh~ dflr:m:i-'!. ¢Ila;r·:illl3I;~S ; ~~II ~~~di chnl!lp~t'Chniq~u!i,~ (15,"

l,u!ipl~" rc~[}Jcl~'4 fml~lh~~

aul

B\.~"W!$t,} thcgro'yp of ~blh~h(*1 of;l;i~· por~lal.iorl:;' Wig- .1131)

Subr~~h['!M

r)11;,'~lm<l!~:i1;l

in t:'~l'orl~~ti~.1 cnl:i!i~~~11 ~td!~u," (:dl~{J{)J

(:TLiIJ:i:'ll.'tfi1J;
l![mllJ![

pG~ui.d o~c!irll~~il;U1ZOO !;iu~dI ij~di".idWl,dJyrehl!ij\,cto lhedorn,u!!'iurlh(;'c

;H'i~~from
11!!1\'\rOi'k.

The ~i~ion

of cm:l~ horii~ont..'11:;lice
~I'i d.ofSl!),1

W""~ H1.C<I-

~h~b~Ji.li lfi~"~Al,
S;tdl~mc cdls
CDllOr'l'ij,l1al (

or

dt~pcltd 1;Ip!;l1~ 1t~~Ul mCit~brml(,': jKlkuni'1~, tfi!C voIIOlgc,. we perr~m'!llcd!1r!i$ ;,u:!alysis ~';I~,,"mlA."!}' d1~"~ g uIhl;J1L;d <lit dWI'C'[(;nl~[pp!'(l1>i:illil1lnx: holld:iL'lg
n'i(:lirllblth!~!J ro!~jil.iai!ii

r~JC

ur

50 nN mld 45 nN.

iH. ~inigl~('U 1i1.:~lthmti"ltjl iIThol\.'~~'ig\\'il!j~IoI>CIii.";LU'!.P!::

OOr!A,;X

tl / JJ)

ilJiti (100M oolllll1i~u[clt'li

PIlli;JIiatIlI ~VlI~~ummm~,~on S1~elet ~\iM, MIt 02215, USA. ~lIO!)l o-f (o;mJ)u~er Sd:emce< i100 COOllllnffli(iltNln, ~1 IlrI!jMI! '01 TedlliDulow 0100

rc.r

Mi1l1lll'-l'!!, ~nd Brain,

OItIll!ilrtJlil@nt 01 P~~hology. Umenil),; 2: ,Cummington

StockOO!m I!rililll IIn&mluie, ~E/1OO4~ 5Iod!hdlm, ~

5IJjOOfft,

'"VoWhorn C:~!1i~~p~denoe sliliill~d1b~ adc,r~~.E·lliai~ ~as.5erililO@!OO ,eoo l~.tllt)) gioc.~o®bu.edu !L.MIAjJ

th~:-:I~tllll it1~itlbiliii'i.l:pi}b::Dt~i1i1 esei lI:;u~i~)'m;; eemI'arecl~\ u~h:lnw~ Ji1_"qn~l1.cje.~ eel I\i;..l'rorn more il~ wmm,1 enU"lrhi~ba.~ slices I Fijil, 113:). Dm'Sffi,1 C'cH)1 (u )U) \'\'C1lC' d;;::!inccl :!i;":: Coell~L'!Ccmtkdill .~Ii:res ~akc11 bcl"H;\C'I~ J, ~ nun It~lelliOl'dlcr \~'ilht"'i(l;{tlihim!l O~H11.C'X U6)] :!!'!.<l4.tJmull [n:;i!'l1I ~hcdOlSld ~r]'i!l;c (iiI'

s,howfil, hi~lL~

Ii:l'b'iljlIIJl.:a,1 freCljiiC11eit'.ll

1'1 of :>llb-

11'iL::"ItI:(~flli ~J\.'(lu~~lie'Y dOJ1.JlJ, eClIIIl> 'Ivli:;; :l'il,gnHiii ieallilly higher than [i~e l'IteiiilJ rll\.""'q~ICI1.cy iiil.'>'C:Illral

rel.Ui;IO'ii~n~ibu.rcit:i.c'~II;:;U Ill:iQ!jt .500 m.\' (dorsal. ! \,'~lllmi f • 4.23 1. 0.32 I ~'z and n .. 25: P < (MH) ~I' :nul £IS :In'V {dorsa I uneatl SE MI "1 .H; ~ .0.50 [I.:; <lindu -~ 4: 'IoClllrnl f 4.~~ + O.5<.l icI<:!.<Lf!d II = 7: P -c O,O! ,. frc(juc~!eics were OOAJ::!11!!ir!~~ by ,In "l1l'iAl!lil,~K"'~ ,~llJ'lQ(Xlm;~tioH'll:!:!.ll.~·
MleIU'I,f ±

SEM

6.42

j_

0.400 Ib: and

u-

1 S;

r ..

[he bmin.

V""IHml cc]h;, (u =

;r'n

\'Il~'re

~refir!cd Oll',

,A rJ
-.J

venml
.... ,}

,'i\'..1I,T"· f

O'~·SO mV
12'

A.

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Venbral
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i~d~cgJleS~M, ~C' Sl[lbtturi;'5h.Q~d' !oKiIIa,Ij@!i'I ti'leq~e~(y F'iig. :II. HighFill flllq~elncy ,(if' ~Libthr~ho!d oroU~~i(ll'!iSill d(l~~\I~r;';ldS v·eii!t~a~ --451li!V, !;Irr(lr OOIrS; disl;ame hom d'msall ~LJrfa(je,. U.H lUal'l1lp~!i ,of 'eontmihinit! ~ 'mr~eox. (Ii) Tiop,. dorsal. \lie\" !l( brain art€f :slkiing., ,i1!o!l:oI1l1" sagitttll plotted verlllllsanatornkal. $ub~u~5holtl os.dUa~(l:I\~at -50 ImVin ,dorsal r~!mj(ll!i!$ al'ld'llleli1tral ~en) vi:~l,I' $~'Ci!'lving i1I~aiOOIi'lii;(O'iIl to((!Itio!1 olf ~Ml~~~liIta~s~kesA t() Iii. UI) Mea n r:i~g,1(l!:'J,)(figh~). (E)' Corre!ip(l!1din~1 !l!lI!t~wn~arioliis 'l!sed to m@i.!s~m~ ('F~q~tlnqr vir ~s.l.!i~tlur~s~oMo~cillati~):!'!sr~rn~l!n:Hf1Srlf(lrnd~mll s;Hc~. {3.8, to ,~.9 fifUllbre!lma} a.mldlvellltrCll slice.~ (11,9 t®, i.l mm l:uegl1lli3i),i3itt -5,0 liftVarlrril ~rE!ql.lemifY In (lEI).
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Fl!g. [)~'~Fe~i'ln'$ 1r€S.(Iiiilam(~ proilltertle-s (If ~~~r(lFl!linl dorsal w.~!.II'S venlU'iill in oooo.rh~lfIa1 correx. {A)! IExClffilptes ef r.e~cm'lIncJ? resf}OliI$~· (l.f ne!luallS ml ,clorsa~ ilbh.!iJ~) and ~fIi~ral enUlrhii'li;ll oor~. (Ired! fl"l I'eSW~:i1se to Vifs-tiil'li!ljjll~.i!s· <blac:k'I. (B~ H~:s~c.grC!m me-all ~~Ik ~I):n!!fi\:e r'~lI!~f1:qy f@~ lI1eIl!Jl!J!i1s!n ,dor~llwf}il,!j~ ve,rural or (l'J1tQ'[hil10111(~rte\K.~ .:; Re;o,S,ODlii «1' nc:e~r'~U:e'ttc:y pfutted V'ers'U!; allatoD1lkfl~ distant!! ,rirOllI1I ~he !l:Il:ifs.;!l ~urrai!!e.

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