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-~ ~IEN_L-_

Volume 315, lssue 5814

OOVE,IR:
Te-CLuliic [lfactuul;, wlln,~ilf1 the 'Candor Chasm<il regfOri of V.lllles M.Jrine lis, Mti rs, ret,auna
riid~iil'li~ liiIorpllo~,ogy aslhE!' s.urroundul'1Igl bOOlmc:k I.ldtloo'S <!IWly. Suciil flnd~n.gis OniN dues <Jbo!llt 1Ui5l fllJid rtO'lvalldl Qlllochl'mlical conditions 'l'IithiliJ~lle SlUIDsurf ,x!!', TIrDIlimOlgl.?

OIEfARTM
907 909
914

ENITS

916 9H 919
1011

Sr;iem;e' Olllline Ihis Week iI1l5d:~fli,-e Eliiitars.'(ho~,ce, ((If'I'Mu:t Sci'f'J1ce


~lIdoliTl

S~mpl'~
(.i!ree!r~

is about 1 kilometer acmss,;illll,mlir.aliorn is [rom the lupper left. See Pilgll 983.
Image: Lunor ,a114Pl;olfetary l(!b'OTQto~

1012

Science

N eW5I1f'I,akoer~ New Product,,>

,University oj AIil:ooWjPtlNASA

EDITORIAL
9 ~3
by R. K. fuchl1~ri

S1.Jtainolbl!?W'@~I-B@illg s

NEWS IOIF THE WEE,K


M.lT HUllg@r Strike: SDUf GF.i.lIJ@s, l'h@ or Sflter Tast@ or R1.Icis'lIIiI? Bubble Fusion Resealrduf Geared Charges, blil~ Doubt'S Lillge-f

LETIIEIRS
920
921 1i.'Voi(e ove-I'tile Smote for .Ac.ailemic:IFr,e,euom

or Misconduct

r J.1l1!

937

Mew ,SWii nflllenla IDa~_abaselo iFest Promises 5,s.1 of A«1!!s.~ :SCI EIN(IESC:OPE Amid l)'E'IJ.ale, Gelnl'-B<lSM Callcer iFpsl Approved In lEuropoe'~ M,anllJa,y; A Glilssy Attack on Evtlllll.ion
U.,S. Nanol.@chl'1o\ogy H@allharnl5ii1lltty

923

923 924
925 926 926 927

ID~lJatilil!l Evillierlc.elor ~heOrigiilllJr LiKe on tartll j, L 8mi(l' elt at, RiI1!'spof'lse,6. Wu(JiffrsMiu~lE'{ and C. Hulm tu:ta ri~i(iiltIOl'l on ~ ['()baUi,C(~SS to, IDrtl g~A Roy~ Who ~$t:l ,(;Jr. l' ,Fl.,McDonaJd COIllRECflONS AND CLARlfICAliIO'NS 940

BOOKS ITAL
btim::t 'liIiml:HifNi!lN' Zi@a!allid A.. Tel'loysQlI und P. M(.IrtiR$,cm; Ibti'm::tiolili alfu:ll8io!l~og rap lily of Tr'O\pii~~1 IPadfii( B,ird$ D. w. sreadm~n. [f!vi~d by J. D'~lllrmd 941

Rese,m;hi

Slal@'d ror 51z'abl@' GililiiS


Ri!ddiJf~ Historian N~med Harvard Piresid~JlL Sden!e Advir-ser SaY511lal Pnmilrlg Is ~Il.lllKey H ealJih)' IBudg.et
~Q

<II

NEWS fOCUS
(as;es· oU~istak:@1I htlli fltit)'
III'he~ 610 lilil~ Dorn'IAdd A lonelY Crusade

J. 'n!lin.~!'en,~f~rJ,~yP~ G~I"$~'"
928

IE,itl$!~i!'IrS ]!)!Wj!' 1m: R~@ til! Test RrIi)tMly

942

PQILICY F'ORUM
9U.goV'
B. Sll:Iieid,~unrJfl and

'up

J. Pref!C>e

944

941

A Haiti-Century Late. Allem1!li\l\E!'Acce'teralor Takes Off 'Saving


<li

933 934

~05,UCulture's I\\egalitlilic, Jars

PERS P'ECTIVIES - - - - ---

lI"w(i,forll1 eoPnice of One A,}. $cJw!i.i!'la =-> Rl!pofl p. Som:t'.~ke' ~~, Hlot

945
969

S.
A.

KRapp ::..:;.II~M'",

!IItJ6

Wheroe ,Am II?


A fem!Q1I
»FlesearchArfide p. 9~1

947
949
95~
952

Where Ar,i2'tile bempl.i!rs? 1M. MilQrd ~~ Reporr p. 972 On Ute Origin50f Grafliles ). M.. £Her ~ fli!pM p. 9IJfJ IE\liery]o~'n;t a 5, EI Sit \fer ~inif'ig H G. 5'. nJ"l~'SI,f'i(l' ~>, Rt[XJr~ fl'. lOOrl

www.sciencemag.org

SCIENCE

VOL 3lS

16 FEBRUARY 2007

_clence S '.'
:SCIENCE IEXPIRESS
WWV,lS

CONTENTS

cl encexp ress, 0 rg
CUIMAllE CHANGE

P~VS!CS

IBRIEVIA:R~~iii'! Tern pera~ulre Oll!anlY m Hall. Efred ~!lG rapn@!'!;e K. S,.NQvosdlJvel a l. me ql~,mtU!!1r HltII! effecL !;!~'Ii!~lly ~en !mIF 10 ~gJeE'S ~el.~i n, OClillf~ at rOOfliI I.e ~p!ij"ture' \!Iillii iii ~iI!gL~grapln~Qe' ~hellt~ If] ~i!ich' the c.Mrg~ GI~rlems bM~l!'e as ma~we r'el<lli~i>ti( pl.'lrticles.
1,0.1112 61sde!l~e, 113 72Q1

An,I\(l""we 5Iu'IJgl.acia~.'Wi3!l'eir 5yMerrl! imWe~ An,taJfCllra MaplpoEdi '~·mmSpace N, .It Fntket; r. 5tt:1mlms, R., Bi(Jd~[Jj(jillFf, L PtHhrrUl1
!iJrt~llij~ rneiil!illll!mi~m1S m"'!~ tl9 at W'iI.eJ ~s U6\1liilig r~pidl[y und'J!I !Ill,~Arit;lfftt( Ice Sl1el!!t ~Ofrninog ~nd dlr.Jiil11img ~ub9'L~Ci!ikt lJk@~ ~nd im:p;!cti 1:191~~~11!1,1l'J1I~S J of Hs ~lOI!tJ.jliitv-

N EU ROSGI EN GE
CHEMISTRY

J'i'lerm!lJ;e!lect ridtyiill 1I~!)llE!I,I;!lItalr flc!tl;OlfU JIJ P. .fltddy, 5,·r: Jaug, (1. StgCl/'m{],l1, A.. Maj!l1f.1uJt'l{
M@il5lJring the indumd'o'altilg)l]' 01 Qrg~l!Ii1: moloclloo'Sh~.d ool~~r1 gold (.oIil~d(IS.,ill

!Hluma~ IN,eumb laM:5:.Mig,rate to the Olfa,t'~Clry Elulb Venin.::tI~a~b10e nsil®1iI 1M"A Olf'lis el at mig[iIt@

lJi<l

a ILall:!raJl

di rf.l!~llI~·temp~riltum~ rrNe<i:Is.wliNilri'!r ha!i?5 or I!~@(lr![mslCilrrylh~ Ulrwlilt. lillU6/5ctef1ce, 1137149

As, in mdli]!IlIIS, ull hllman fIC:!lJIOO born ~U(l.liIglil~1~uid-i·ilooqj llnlJi'(ll1's~n!lhllibriliirll M S V ~Q' the olf~ [Imy bulb a,ton'!!liii lUoolii~ C'xl:tI'1!~iol'l ollhi! '~mficL!1:,

lO,.lll161sl!:ief1ce, 1B 62iU

R~S~AR!CH ARTie!...!;.
MATEfHA!.S

SCIEN CE

NEUROSCIENCE

AIlI)'lkati®~n j, f. S(,Dr.~

of Modem f'l1lrrole:le(~ri(s

954

PaU'Il,rIll Se~lr<l'~i,ol1 in the I()Eu'Itale Gyrus and (A3

9611

,of the Hip POC<l1n pus


j, K. teutg.(tb, So Lell{geb, MAl Moser, f. I. Moser RillS !)rJd~ slil1lill~ ~il1i'1n'ge!5i11 lfioo.it ~noun ding ~n.vi[rJJi1lm1!'nl by mlCldilyingll11@ural i1divi~)'il1lme cremt~ g~ms and md~ l~rg@r diHmli!iIi1(~~by <ll:liivililinuj mmroJis im ,~n aa],lmnt area,
:;..>

BREVIIA
IECOLOGY

s. (flEw'

Pred.i!!ti'(IIilIFliskMfec:ts D@:!1f!@!yfo!!pilly (lIf Elk

Fleprod9CHv(l! Phys;io!ogY'~!lJd

960

PfnQ~'~m ~·41 fl.

£1 Cnr~'stj(JlIrofl, S, lite" l A- Wi\IJnieJr.

'!'e~O'I'I$I(\I'Itell halrre l~erOn5p'rimg II'! ~a6 in which a prtd<.!Wf. 'the: \!jail, i~, !f!Q«llllenl.iiut demiO<li51:ralill9 en indi~ct QOGt ol a!ilt~pre~<!lQr r6piO<~~e~.

REPORTS
PHYSICS

[:l;perimelfltal Rea~1(ZaI10ilofW,ruefl'iet':s Delal~e[H1f1mr~ Ged~llI1leiFI ,Ell;pe_Fi!iHIiIiU i!I" Jacque! ~ ;\lIt


tl J~LjlJti!lln ~rW1li!~ler'sd'el~~d d1oi(~ g~d~t1~ene:q;l~riIOOnl ...nUla si!li!lle pholOi!l.aUirnlS tlli~ ~1j',e·p.;Inide dlu~m~ ~fimcip1@ 01 Q;UO!iI!IJAl ma(h!~ni~s,
~HYSICS

960

Ml!IUip'~ Ef1~rgy SC~,~@~at ~ Qll!arrlllUJii (riti!l:C!JI,P'Qillt 969 P. Gegftmwrt et at TlJi!eriT!OdI~ai!'!i( fI1e'a!~1J1e!ft1eIl'l1S Ofi!\fbR!t~5i~, a he<l'l!~ r~Mllo!'! J)1e~~ <It ..Q filelllil'!l, IlF!'[eait~ e_!l!i~l\e'i!IOO01 I'!I'IOIl'oe il~n Om~ elll!1,gr scal,e- <1M U 'thus a lfilm dM.S or !JjUM!WI1il crittcalny. !;» ~r.pt:(rillep.. '145
MATHIEMAIICS:

(tuMerin9 tly P.i3lssin'[JMe:r.s.1~,es IBehllee·nV<lta Poin~ 972 B. J. F~y r9'IU/ fl. f)~e(:h An aLgorimm 1hII1.eJ;chaFlg,f5, meS5i1gf'5 abooUhl': iilf1'i~~ri1:!for pailr~; 01 d't1i~ii pm m~sspeed~ ~d~nlilk,iltiolll oi mpre51lnt.l!tll!'~ ~;J;i1rmp~~im ,iii cernp le~ dlam~ ~~et.j.U()~~I~ genC'.l, il'l nJitM d.at~, ,.,. PmpN:five p. 949
CHEMISTRY

1l.e\fie~5iMeC® ilCer~~d L~gallul 511b51~IJu.lur!)n at 977 AhNlnating ,MeMt S~l~S, lin <In It)lt!enr:Jed Soiid o. 8,i"ot.tsh(lI~~, E Waf1,ftl.. M. J, Rosse-u}sky j GII~sll'fllollJ!(liles ill a parous nuIDl.-afgi11!nic 'sollid GL1I b~ S'III'bsli!Ht~dl

,r~v!mi'bly I'm \lIatIW· ~tup to en IHhird 01' too mWilL roIliOOt~. l!oIh1ih~hl! t rI! iTlUii'ling mr1!t1I(1!II't~i:S provid~' 5U!JpoDl't.

wvtw.sdence ma9 ,org

SCI EINe IE VOL 31.5

16 FEBRUARY2007

_clence S
GENETIC'S

I.,

CONTENTS

The ,rawpli:Olileifio ma(jnijim Che!lIH)'~ lItatr~iPhi,

99.8

S.ymfull@l1it
GEOILOGoY

(j,oeI'l®llllfl

MagmaIU(i3lnd Cm$taJl Diffie ~eflllii31tlOI'lIHislO iI:f olf Graniti;c RC'(~ horn Hf·O nsolt.opes, iin :Zi~c:o;nl

980

I" l. 6. N(!wl,()lI el 11 L A di~ni0.3 uwtI'iJ,p'lflk 5ylilbiont 01 ilJi.e giamteiiam JOlJflcI il'lJ hyd r'otherm<l!.'\!Il\i1I~S'IiIas iii (ompl~i( m>Mabdlk repe rIm re <lndi (,18 pro",id~ its oosllMlh IfiIO~lIlU~ilic miltneeds..
M ~CROB~01.aGY

A, j, S. K£tJlP el al. "'ilroi um and O<X!(g@n iSIJl1:iPMin ziocQn (I)'iNlis. iJllip~'fIh<ll COI1Ulilml granitQ$ farm c.,. ~Q('i'dimg uusl<il.lTI~wr1<1i lr. milnl!llNterilr.oo m>llljIDil~" noL ~ rl!rnclaiiog d'~'jl' CflJ6I<IA, ro(~s. ;>0;> kl$~iJ~ Po 9.51 PLANETARY SC IIENCE IF~a,ctulre:·((lI1l~roHeWi ~alelo·F~tI;dI flJ)'1l1:ilf'l 983 Calndor nal~rrna. Mairs C H. fjk,r;ua utta' A. $. ,/WCfP:-,'t'/l Ilie<lcfilillig1) nd roill~ota!iool ~~seen in ImcllIoo$O 1lndi icinl.'~ lh1ll mSfim)s~~he- ~1l!f'Br~d',SU trlltfb-ni;rihra,~ oJ iGalli!lm (1IlIi:um~1 Mms, am
f~\i'eii'li11i!l an Ollrli(ient hydfO~og ic

The Pno!l1pno11li reollline L1jase AJ:livit/j' ,or a Type UIIEHec:lor IFaml~Y

IBade-n,e ~

1 000'

H. U~,tall. A ramily 01 virlJl~nu factors in it,a,cleria H~rTliG'i'~S Ilh,c~phil"~ hom <II a k.e:y 5ignaLrog ~m~m~ in ~13illlf~(Ei:!d hCli>tamlllllemD'i' in~Brr'Bres 'With t~ hast's ill~aw immunity.
M ~CROB~O'LOGY

Ar(IiI~~~fType
(@f

WRd,i$'COs Furtd'iilllliiilii AMP MeIM]iIlli~m

a ;P(iHn~(iy

1003

iI: 5t'JW, N, A/,Olnt iI: 1!!I{1fuJ{ca


In IIl(mphDIrn.ymht@li[ AIch<Jeil, lh@~I~2'jmill' lRuBis(JQ ef)~!; M~ li:t (0.1: as d'o~~in IlI1lmts;illlstil<ld it s~tv"g@5 ad!i!nosfiol! iLIld dJi'llllfts rioolosill'

syslem.

ANI1HROPOLOG'1" St~lu;h ro:ssiils<!I!1Id t;he Dome:.tici1!tic'lllalnd llIi,~em!l 986, Qf liftepper~ (C~'.rk~:t:i'l $IPlj), l..~illihe Ilfll'e'rit;;<ls t L Petry e~, l, s Mt;:i'Or~!iI£ iIIl ~ar'(h paltit[es shoil,; IhM chi'li peppel's ~i\e iJ~ed nol.~Sl, as loed b wi IOil!:ldspire to c.ui~~ne~ 3000 )leaFS agod n lfie. N1I!'I!j ",odd. ewn Ibe~Ofi!l 'tine U:5.Il' (!of patte!)" ~~ ~fJp&fillf P. 94!ll

'!rim

MEDICINE

(oldlh~r~~-l1 SynQlliallU,nriilll'[J hI P(!thQll)g~l'~Il tllr~hri~l S.

F,(mJiloiItn(l!ll oll'1ld

1006

D. M. LeI!\'et at
A Ii'I'Il)u~e '!$'~iOli'lI)~ m~umato~d ~rOi!'itls·COIlI beanlelio1a,t~o1 by
I nl'li'b1lioll or elimIl,natiolil er a ce~~S:UffOlC~adheslon fQolec!i~~ IO~lld

OEViELQPMiEN1AI. 910W GY tl~unip at'!! I'll DroJ,Qphila Inl:estilllall Stem (eUs, 'S.pedfy 9138 Daughter GeU IlFatesb:v DiiHerealtial ~o~cih Si,giilalil1i9 B. OJ.rlstein ,000 A Spr.ruflrng Stem cell d<ltlg tite!rs, in the f)ffJ'>Opfriia ~n~E'Stine' n ta~ on, one 9~ COl UUl!tltdMlm~s;~his (h'!)'i!]~is conlroOtled bj' ~he actwilfionlll~1l1 ela
ool1llll1on ,d~lilplJlloo~~1 reee p~Ol. CELL BIOLOGY

wiUlin jQ1rl1", 51!:ggestill'!9 ,a Ilnerilpe~llic; OIP;D~i)Chl~or hUjl1i1!'1~,

IPcl'~m,~rtji;i g .Actilii '~I'Deor~ Pesltio fI h'llle9lril'll~ Pri f'lIlE!d n HI F,!,nol!le for!l!,dll"l~~If)1ii Sl.t:€~ C. 5! Gfl/o'fairll, It. M,. Yarn(i'd~ J .4, G(JlbflJitli

992.

10 i!1'16~n~ O~III$,<Kliio libE!rs lam im1~r~n"'Q\ll!!f~di pi"!oU'uSion~.lilll;}t are pt1)isedllo ~1'i~rO!(t'lillh surf~<e~ ~II the c~11'5·v;:... r(hfm ~dh,e~ion sfit~s_ \ N I:=IU ~OSCIE NeE

MapU ~~ IFt~p ~~ll~rntati(li'l oJ (dE!~ti(ll f}lJ~(tiJ r 99.1i Orii!'l1t~tionl$i!fl Uil~ ilriili!ll or (1111 lliisect :s. Nfil~'€ emJ U- /1OfTlbr::rg TM ('Ifie,m~~tii!ln ~rIPOlilri~e,d lig IT1t ~represelit@d! ,'!S-.~()olu lirnlilm I:rIilP ~
in I he loclJ PI ibrartll,;-wh idl n]~1 he~p to or~l'I~ title inSeC!un(ler 'lillie open SIk1l'"

'952: :&.
~I006
~aEW~~ ~I~ !M!UI~~tiii'-~~IJI~I:I~d!~J:ll!lr~!"IilFl"lidiif~

i!iii1r,.,~

luii:fi"IIiI~

II'i

~~i:-iiii.l!iili_~

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'" 0'0'''1'01 iC~NIG

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~,"Iog~!\.~, ru ... iI:;,!~1Q;
, .... 'pOO!i!l ......

... rlrl<>",~.""ld,II••"UalO. Fh,ldo:", "'ma~m, -.II,,,,,",,,,, r.11.. OIl'HI<"<I. ~n,_..-I1h "'~' • .o!!!li!O,~;~ ijlUII!~ 10m.~b'!-~~,!.U'l[""'!PII!1iO!Ioi' WliillU ~1!"'-'!!!" g,.~.oIo. _!I'.1.doi""",_~ .. tb,~oIlj onj]"".,__ ,;o,d B,j~oo:ru-'t_, ~.. "u.M.II';!L~ ~Ila, ~Ion, Dol: ~~I1-1( 5I"!1i"'~~"i"" a~1il!IW,m." [!!<{liDh!!mluf .... 10;0~ w'!;'Iiti"'" ~ iW;I\ot;IaIt!';g q. 1Ih"'_\!oot!I'",ioIiII!illl" ",.,-<!!!:.!!!>l_..,I: .. Il!,;i.. ~""" (ji,tI!'('!""11h1,iII;;i~ ~~ W~~~"'1IIIO:i M!1 (lIM ~l'''~~'''''1h! ~tIiJ (1<;;0 __ C(lOI;i' .;ro;)~iIi:<!OI.R,!,p!HI"I~">Qo ~~ Q 1Q;(i(I p:d,III"'I,·Io;(D(,m~.....:;:iIllft.~~,W,OI,9~j,'IIol·~ItI,.,.""m~IoI'~-",.~(OOH!IB_5rkl>ulolh:l....:ilo.ihoRc",~'Gll:IIj"J!!M:r ..... tl_~"""'~',!",,~'lHlllhl9R,
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SCI EINe IE VOL 31.5

16 FEBRUARY2007

Ide -ce

-lliLi~tE__
www.sdencem[l9·01191

SCIIENCENOW
1lf"''1~_s~iel'l~i;'110'''UI

rg DAI LV tol EJ.V~ {o\J~RAGE

fn~ 'Chil[]llpall~.1,l1,l Ston~Ag@' New~lnd 'S!!I99,Qsls chomps h~ thousands 01~ars. Taikir.!I 'S:Lock ,cd Trees,

used

st'm~ teols

Wor

Bank gr.mt m<'lkts 1J1)5slloJ.e. worldy,ide coll1pariwn 01 r'Ofest d:ynal1lics,. 1iru~t Tubular TV ImiJfOVf'd organic LEDs migl1Ueadl to mil-up 'lrid:eo

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SCI ENCE CARIEE.IR:S


\VlVI'II'.s.derlll:enne-ers"org Cf.IREEiI:RESOURCES mR SCI e.NT1SlS
I)I!.I~ Ilmigel"!(

US; fQolira91 Up-lEmr;l[lJ¥m@Q1t

D.iensen
An(l!OrJe'who aCCQPt5ill jab ofrer1mtnootod!oing

a oockground

inveSlig:1.I1i!:lnis lilkirlg a ~eriou:5 rbk


US: OPiPonuli'lities-A IDa)!~nth~ lLife p;, Fisk!.> lin IIJn1likef'l e,urepre,neur ~ilrs QuI. a noM OPfiMunil!l'. 'GI.. I3tU.. MCI$l.eri;ng Your lP'h.IJ.-lJea~'l'Ig \llfith Setbacks O : Po Gos[i,ng(lJld B'. {iJllOfdt'M1J Get ti ps. on hew 1.'0 pklkYOIIJ~~lf IIlP, bmw )'(ilurSfU oH. and mOllIE' en,

SCIElNCE'S STKE,
M""~.5U.e.o;rg
5lGNI'IL TRIINSDUalO~ KNOWLEDGEEN'lJIRONMENT

PER'SPECflVE; 'eetin9 Repon~Targeliiilgl the IKilltom,e, M 20 Years o'r rYlF(l'silne Kinase Iinlilbitor ,R@S'iNircl1 illl Basel i. Bawlk, R J. A.~arin, r. HUlller. B.A. Hemming:;
$l;ilmtt:i~ m:plomUH! pill~~ p~s.mi:, and I'uhm~ 01 pr'(]l~i n killl!il$~
I~Seilrrn

in 5~nl.,[]h;~ro;;~r1iI1llfillPY de'l!i!llJflm~nt.

EViENTS SN~cJil or i:lrowse IhislJ1pdlated Ilist (If signa~~lg·reta~ed eetings m and (Q\nf~r'llllloos.

www.sciencemag.org

SCIENCE

VOL 3lS

16 FEBRUARY2007

'9()7

WIUlW

BV S1rIEUA. HURTL'EY ANID IPH~l SZUROMI

-e

Routes of (hU~iPepper
D,o ml@'stic.(I.tilo'n
A wid!~varir!tv (If chilli

p~'~p~JS w~!"c'c!.!II.ti¥ilJt'8~ ,;:}IFldl used ill (ookingl~hroughol!lt lill,e NleI,\lWor~.,d. P~rry ,~t ,ot (p,. 9a6;~el1! Ui!~ P~~~IP~(Uye Iby I:{n<lIPIl)'idientifiied <I 5ta['( h frcHfll chi~, Ipeppers 01'1 o'lndi@!lilIPorttery mid stolle tools th1!it is diagrlillsUc of gml!l p!'l of (hiili spe{ie-s, .. llhe $tarcile:s \llell"e' fOlm~ <lit 'Varicas .wc;naeo'toglicilli sltes.im;lludil'lgfrQm al:lou ~ 6'500 ye,[Ilrs ag'o ]~ IEcllladm, and sllI'gg!est rnlu~tiph~ d!cllllles;U(;~tedl Q~mI5~edes by~iJ(llIt400Cl years (lg:o,

Delayed Choke for


Quantum Me(hanics
Wave-j:li:i Fltic:Le ~ualhy is, at thE! he3rlCilr quaiill,u lfi uii~(hanic~. P!'il'r\i:des~Ii!l:lpIiQ~[)liis C~1ld~5;pldiY h(]th pm,lJoeuies., aFld whfd'l prCi!J)e,rtllis meas.ured d~fru:ls on 'the 'typ@' of m~SU!l'!]I}elllt !m1!d~.What ir lilte ~p E!rlFiiU~I'iMiI~eIluphanges wrl~n the p hoc !ClrlClr ,p.a.rtideis "In rtiglll" ~Ilid h<ls already eflte'red ~be e~J)erim,efltal a~paraUJ 5? ]laoql!te;s, e"l' ,ai'. (po 9(6) f:@portan a lnW'Sl~d@al,f'@I:IliizalioJl01 slda"~elayed (OOi(,eH eXl!'e~II11iMtas ronfliu'lal,ed by Wheeler, fJ, trr~mer~d '~illigl.e·photl)n wu r(l1!prov1dll~i! met: I~rlis,m fo~ pr@cUSf!tin"ili!9 QJ lhe 8'llleri me!'ll \vilhi!'l l~boratory cenditlens, ilIe b!i!~la\lior 01 ~hE!'phOto.l11 ill~ tinE!' ilnit5r~'emm€t.er dep!!'nd~ 01'1'Ule (notct! or till' ob1l<Ellilatllethst if~ IITIfM5lU FOO', @vtl'l firtm~t dN)t~@is rrnrdt! wlwn th@ pllmnn is already in ll·te syM.em.

gli1f1lesequem:e. Unr".rl:ullatety. 'Iil\!:tr.ading 'e\li'lilllllBJ!~lr~if' \,Qlti!p 1,lIt~t~I;III1~"1I ~nt~lIlsiJ1J~. r;dI ~~n"'~IIl0lI

tional tecillniqLtes mi'ly wu.ri< well with f1u~n.elFi(a~


m~ii!iSlns·

of diata-point :simi~rity

~ndir Ull~ ini-

li<;il g uess is close, F,lIey diiidl D'lieck (p. 972, Il'~.di· lish~dolll~in~ l1.lafHla 1)'; ~e~ tllie hr:spect~1Je by lMezaiid) nOW r'll!jHI:i't a m~lnl)d t.h::lll filial) les lI1Iu(lilfaster exemplar d'ele(ti(llll, Th~ a~go:ridll1ll 'l'.rarK$ b~ h>1l'l.i'ung 'th~ d~la poims ~chan9:e Nm~· ~ages~ Ihilt ~o.m· JDU l1i«(lt~ wnetl1 iill a

palfli(u'la r pcriul
could bI! :tllll ~Jemi-

plair; i~eri!tiolli

or

Ut~ JI1!es;s~lg~-INI~~llg !'In)'!: ess ~ll.a"",'SdnIiJJ~1llti(illlly Jau,err prm:e1:~il~g as (ertain. dM." poi I1tS ~me,j'ge 0I1ii uu I~ r(!'J)re:5e·nl~liv~.

flQ[IiI1 iild wiln~11J ~aFltIh's crust by inlms,ton.or IlleMi magma 01 by r'efl'! i21Itill'lgor i<gnetllls volwielie'5 ,tlr (rf!'!i5ta~ rocks, Th~ ~o~~I.iilJ.rl;;iry history Qr 9 rr.iili'in lrfS, (~II! be rew~letI by exa mil1illlg the (hie,111Ii stny or sueees'>lVe< layers O<~ It'!: large CD:fi,~\ itlleflll (Irr->t,a!:~, f1ota'M~zi ROil. I<relmp e.t ,fif. (p. 980; see tl1~ P~rspectiV'.e by ~i.elr) hav~ m;e!'iSl u red haflf1liru dillie OltY!l en, isoIn I~pi!s. illlUln8d zircon c rystClls Ir,om dne da s:;ic: gr.anil~~. f o e~,st~'~nA!.lI5~roilli~. Tlh;€y iloulnd tl1allhese {lriln il.es, ~or,n1.ed II:Iy the reqrdilllg of de-!llf.! (fl!.!~tal~ m(l~ 1I:i: lli1caln:tle-. RlaJ9f11<1 rliJ~ th r(Iu.glil !h@m', H'ith~r IIhi~H b~ [~rli!dli!'lli] ~f1d@nt, s~al~lClwer I:ru 51:,. as was l...mdely believed.

~a!rge 9r~lrrliw: bOOlil;!~lIfI~y h~ve

UlrnJ:Heeiling Granite's History

Water Marks
s.~ura{f or ilMf5, wheDlit W~5 worm ilnd 'lIj"\'t. TClla<llY, however, all ~hat ~em.ain <I re milnelral. deposits, in dmUn.g sui rates end da,ys. 'Oku bO<Jjlld M,ewelll (I). 993;
~lvYfed
OTII

Wi:ite-r may

ilave once

the

'Guests Move in

Hippocaropal

Dualism

When It's Hot

The fOlrmClti on of dlis(Jft~ Ir~prtsen\oltiQIl.su!1I mel1ll:01]' has been illyp(ll~esi~d llJr refilreu
n~ur(nnal p~U~'IiS~I)@r(lliQIi !:lit tlile etillrly stages 0 I' the h1p l1o~ampa ~forR'fI1i3Jl~O b i.I~ rD, bollh locaH~!l and medl~ni~!Iii~ Ilhr precess Qf ha'lf<f r~mailFiedi d.usTV'f. I.eutglelb fit aJ.• (P. 961, :se@ the fl~rs,pe\.li\f@ by f~,IiIt~:!iI}s howtllriilt~h~ ~l~p,p~Hamp!Js has ,U lea,si. lwo methanlsms lor p<!U'Iilm sep~'rc1lt~onassodated with dinefelll par l~ ®f the h ilpp(lC~ln~p~ ~() [eu i!. lin tlile d en~,ale gyr 1!!5" 5,il,]l1lil1'5 illre sepa ra ted hy h igihMeli!.), d ~Cillne~<lti on or uac.ti 'JIlly pat M:·r,n5. wiLh~1Il it! subs:el, of 1lJclive! c:eIU~.In CA3,. f'u,rIther :sel~ilf(i li on iS~,tlir~".~dbr~' (i ,I iv.3IUo Iii oJ n.[)II1I" O\i'rl'! rlalPpi~n 9 l['Ieu rCHIal 'slIbpoiPu lati Ol'i~. Jllle ~.\l,ioliiie,c:h1ini:o;ml~' ,paUE!'fl'l sepa~~lioul, acsS0 o'f c.ia~ed v.~tihdW~erel'ltIll~ of lhe (irwi I,ry, rts S'U p pI;)rt d istinet IOJins of elf1l5<~mlbl€reRlf'~5€ 11~,alio 1'1 ij iii the ini[)p oc.ampills.
2

See the

r:::()",~

r~ :~JnJ'l'i'iF. veJry detaile(! ifllagif~

r,fi')111

~ ~Ilr(lMar.; ~~Ol'lnaif55.an(e O~biter iihalt W<l~ef 91lCIi1 i5 ~lQw~~.jIkm91'rojl~hlr~lnll~! t;lrQ5'~~ tne ~Ylii:r~ CI G :i depCIIsits;in 'W~~l~rI'l Calf!dOT 'CI1ClS!m~, w,he,mirQl. b ~~iI(lii .. n:d (~I11iE!'il1lldtionIre ~eEliiin 'th,~ 11:9 ! .i1 kadure ];OllleS,ttlal. a~e 5imi~ar to 'Wa~e'Nelaled

i
10.

Micmp!on)u~ meta [-0 rg<l11Ie e?i~e!Qrlediar rays G:I n ClI!J5,or guest IfflOleclu'l~5 a,l s.pecl ftc site:s, b sad Br<ld:sli1l<lw ,e.r at.. (p, Inn 5how that w@lel" melee 1i.I1l@~~ III iI Co 'I;:,,~~ ~d it rr~I~. I'll;! lCo~(bill'Y) ~~S.O~)2(H~O}2l, call, IIIFld~rg o S~Jb'S~.ilUll~o:n re'a'cti[)111~ \'dilll s()U"h~a met!! .. n~ ~ and

bipyridillie molecules

IJIl'OIl

h'lili3l1ng ln a dry

~ P~Q~esses oflEarll1,
co

~ Findingl a Good Exampl.e'


:ii: GtUl!lPlEiJ!: data sets c~n b~ more ~ead il¥ a~alyz!!d ~ ir JeprE:~fl1I,ative e'x,am~le s c:an Ibe ~tI!!!,mifid. g S!.Jch ~e'X€'m;p~~r'S~ 111 I:glilt be !points around \~hidll diHa wiUclune'I.,alrd1,~1)ilpal 'f1lIce!s 1I1'I1I(mg a u ga ~~efY Clr ac~lI.Jili phokl:s, 0:[ po:ssi ble oE!;l;O<f1i5 in a i

i~<e r~ ~tl1llI;lS 1)11 i;lre. ~'~dJt;lge[:l-bend i~gi i!'It~ri!!!;li ~l1spol,iloe two 11~ pyrrid ~ru:~'s, r tW(II met.h'lI1iCilI:!l o nealr ,alt~m!'llillig Gao<'!toms along line\'lf c~a1JlIs L'jl Ihi FI the 5>o~id.Wh e~ t hese iIl~elec.u ~e5 di5pial; e the "'~IL~ ~i ga!1d s, the I:Jaai Ilsaldopl 01 r zigzag ge(j'n~euy. mll'i~r (0 s~tes are :'lp~"la,lOr~ Ihat Il,elll maij~ti3!in the framewl)[k while tlilis

l'e.>di(iUl!pu;l(e~ch.Th~ readi(!!1 (~111! be Je'II~rs~cIby rdydwa tingl til ~ crys,~al~t room


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VOL 315

16 FEBIRUARY 1007

DIuring developml:!!l~, stein ~dls. ~sually 9 frl!l'Fillie [\1,10dal.lg htl:!r cells thal9!t1 on ttl diHereF!lioil~fas wi;!~1 a s a iiolh~r :!:telfll[.e~L OIIilLs'll£'iii'ia nill :5[prad[1i iBgl (p. 9.Be:) njj' ....dI~![filJe Dlile type' (lr~tem cell in the D«m;p~i'ff~ int~sl;ne thijll, pn~du'C.es Uu~eI;lH$pa'iI19 r(l!e~ $k!DI!(e~~ (in ellltt!lro~~~, ~'nd ~I" ~"k!r~.e nclOlrii1e lell" The c:tiloiClE!n ~<lte< Se'eiilU N~d ell'enci m~ [he .a iTIi:illnt ,or the IlMIe,;n, il)etta (a iiIl!'!mbr-aII!i~'i bou!l dsignaling ligiillJlidthalC.1!!!lI actiwl:e Ihe s~gll!a1ingrec.i2iI)tor NCllkih) Ihatis e:l:press:edli n the stem !(~~l at li'l~ time a:r ri:'E!1l Wi SiOlll.nau~n[er (.e~~\\lilil hig 11.a nrIOl!ln:t~,of [lellQ -No~lin s.1g mal~il1lgl d l:Ierl!:liil~ oel1tefoC\!'l'El~.lho~ WitJ11.owell'<lmOl!llts bElt.mne- Il'merMn~Jrine ce~ls, and thM~ with 'the' l.ea<;t amouat N'!ta,ill the ~ILem,('en J au~.,

Try ChemBridge's NOVACore Ubrary

Sticky Flingers
As ~rli!11!alC~~~S mi'gfall@ i!JC~Oi5li asurlace, thi:}' send out pr'OCe's~e~ ~Ilown ai5 filolloclJialll<!t~xplore thesublllJc1l !!Um. '6~lb:r"'HIiI: el (:II. ~p. 9'92) 1110\11 rilrld that ~lI1einUr:a'l;.e~!ur adi n ~ netlffl).rk. d ~rel:t5 very local pmtrllsioJIS thai. cmllaill diu sten of (e!Ul sur~a.~ @'-adh@si'DIIImoi@wtes at their I.i,p;st~all are priim@d to inter act wilh mol.ec.lIles of the e;(tjrace~lu[lar 1!1,alrk 1I'he3e ~s.lid:yfi n9€'FS.~ o1Ilthe I~ooing~dg ~ (If nnijitil~ (t~U;r, ,]Ipp~~ r IlJ' 'Seaud1 for 5uil:ahle- s~tes o~ adh e5ion Hlat. (an h~11: be used! to

help move the rest of~he cell.

Locust Nlavi'gation
The- pli1!!l'!!l'of P(ll<! r'i latVl)11 of stlll ~ighit depends om I;he :S IJ n's posHio:fli, llm~ i1I ~riffty (lir lnsects lk.e PO~llrr~tiol1 ~'I~el1lu to iJuid1;: $1!'~!l(i[llJrieIilUlt,i(lili. ~eiilll!leali!d H.ombe'rg (p. 995) ShQllll!liiIt lhel;lri~nt:a~i,oiilSor 12~ec'tric fi;el.dv.e(~ors 0" Iinl2arly IlO(llarflzea Ii§! h~.{f-veclo,r:s). p re5e~te!:l:'fOln' a!rove the .a ~illlai. a re repn~e ntlliild <'I~ a wp©~r (!r;lh~~il!I, m '1!p i~111)e c~llf.l ~I1S 0'1 tITle c;elll["@1 ~tlml~le;:; ~11ti1!e ~@".!:;tbn~itl'!,TIil~ (:~llitr al (omple:l:ads asa lIlillltern<!l (Q'mllas;s th~~:uS!l5~he 1)018rilaticm P<itteflrl of the blue sky ~:o'(od,e $[~ti;;ll. diir@(tiQns r~ltJ.I~nt W. ,]Illfr:n~l ori@rI~tiQn,

lethal Inierf on
p,]l~hoge Ilk ba(t~ri(J ~,]Il1Ii 1l.~I;l~ t i~lo hCl'Sl,"~~hviru ~~m~~f~lr;;[@rSiO;Oi(! ~tI-~(!I~lt!dllypft II]~lillC!Ic:lilitl'l~ry. the L,},el' 11~.(~. HUla) tiescrill e 11lam H~or balteriali viriJ l.ern(e ract()r~ tillal [I~a\ll~ a lYI~viou~ly II Ilk MWftI ph~sphQUmacmi!l@ ly1!5eacti'o'it~ Iflli1!!lcan remOVE! the pllO~ph~li!2 fr,olm si.gnaling milClgle<n-<!cti'l"ated prDtei 11Iki,rll!3~eamiW melililiJer$ il1l\iolve.d'in illl n<lU~ummuDili ty.TlIli$ ~iliiniUlI' OIletrrerJj}~ isi mp~rtallill in r '~h~\l'i:rl!l~~"oooJ iI v"rf.ely of a~i!11<l[l :plonl ~~c;t@riap~ !h(Jg~!1IS. im:llld~n!l SJ~igeJl(J, Salm.meJlo • o1nd l .and Ps(!u{jomrmlJs JyrJ'rJg,t'le.,

I
;~

~ A Ru IEMsCo with No laste for


':!!

(02

I[.nP~iOIM~ ~~~.eti(lIT.!. 03f'1~~,n!s" UBi~.L(I.~, { ...R. (1I"i.ljU~O~~.-1l'5 ..'biSP~tO$PhMei!:arlm~¥li3i:'O. .., e.[l.xy~el'las.e) fi.... (0.1 €:S til mu gh~lu!·daflk re-antalll' ~o lIIlil~!! orga fIlK nmp mmds such as sucrose. bu r ona typ E!' oJ Ii!IU BI~GO., ku.l Uld iI) mmphoo~(I~)!I1!thettc;11 ~er(lbj( ~f{: a€!(1.!1,.dQ~ not, Si!lit@ ~tat. ~_ tOOl} [QI.I(!Id thalL type III h '..: Ru SiSCo,aU~l!Ig solely as a (.atroo;,,:y~ase, illl (oITIitliflali:o~ witl1 n ew1Y'~~e<lLM a rdl<1,eo3lL nudeostde' i! pho~phorylii!s~~Il:d ribo~~- ],'5 -bi:5ph(lSphill~ n semerase rUl'ldions. C'Olllv@rL~ CO2, ""'<'iter.and adsnesi IlE! ~ s'-rnenep h05'phat!1i! intQ 3· pllo~plmg ll{( efal~. whicll r,eeods Iibose ;lllitCD c;amiJOI1 m,eta tm~i~lTI, salvag,e~
.

:;: ;1I{i:@ n~I1~<iJnd ::1:

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l!:

; FIll~tuna'teid arthrHis 'WI) isa na utCllimmu file c:olldi'tion !,h~t leiWs lil) Joinl, inHalil'll1fl,aJti n. Le.@e;t o ~ (~, UJ06.. ,publ~sh~d 0 1!I~;tll:e ~ ]iHIU<llry; Sie~ t.l'lrll l'Ie,rSlle(~i\!'e[b~ ~ires:I:e~ n) id.entify a I'I~ ~e,gutator .2 ~ of Ihe c~~~u,la .organi z~lion oHhll S!j1ltl'lJijU ~n tha~. might a tso prtlll,i'~d@.~. pot~11I ~ijl.the! ra pe'ulk targ]@t r ~ ~'m in !tarrn,naat~ry a rlluiti.~ .. Mi ce' ~a,c.ki1'191 the ceU .a,tilne$UO.ll1fUelL.eCiU ca tUleirin ns~loj,!jed ~ignifiIe! ~ (<lnUy red u(!!d growth 01 uhe sYl'lo'!liulIlafi d were f'!ils.~5t,im! teth ~ dE!w~lollrn elill CIIf.[f fll e:l:per; mentally

or.

~ body ~o ',adh~~in - u.

~ induu:d [RA-llkt!'~{lnd uli(m. ]I)ilflt i rdl,~IlQnII.]rtiofl i~ll nm;;c@ (~~ld b~ illl hibiledWiitlh,)l

mQnl;ldon~ ~iii IlIti1nI .........

I!"I~,

. e- .. ..,.. ,~..4.c~""le-.t;;'t"".·I. ""m--.'~"·I""""'.;.·-.,- .. ..,".L!;I~,;;;l",_,,::!~""~II'" __ ~ !"II ...... II1II www.ehembridge ..oom


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SC~ENQE

VOL 3,15

16 F[EBRUAIRY.2007

R. K. P,,(ha~nis directo~ gen~ral &~The Emergy


~d JtesCl-'!Jrm IQStlitute

in I~e\lj IDelhi. India, alfldi chairman OIfbh!!l' nW' I


!lo'l't!:mrnmt<!~r"!l!Il. en

Cllmt!ie

Chilnge,

SU~([ THlE AIDVUn 'OF INIDUSUIALIZATIION HAS RESUlTED .~fl1g,elyfrom ruchanees in scknce and l0C.hl.lO~.Ogy {S &J). Yet CVCI~as suCki)' b!.:nc:Hts from S& T dlLtmgh. dt[J~mo, ~hat \~~ ~.l~~¥'~ s (lOln~ to nlk~ lilrgnmltt!i dOCl:o;gmliSmil its fUJ!llru ~Lru ~l~t:ru:::l!£ingJy bel ng quest icmod ::uld scml iIl~zed.The e urrem pmh a,lfeeenemie grm'\ftll. dev hues: from. the object'ives of :;;tn~~<li'll,lbleh::'''e~()pme!ll_ hi~ "_,O[ (lnbi :;)rJ(:~l}' ~tll'lT1:.e, spe-"lIh.e<llded h)' Teaders.of ( i [public opinil'lni. Ehm is expressing ecnccrns, but also the SdiC;iIiI[ine c<lmmulilily ilselr. whiehLili I~ooking far. ways to prollllo~elhe snsl<!Iimll_'de HIeing ofaillunmm:ity. wei 1'1~is microscopic IIU1alysi:> ofseience mill :its appl:icm i.005 emanates fliOm!;Cvcr~li~ Delconcems: val ~hefole of science in the development and e1o;~ensive use lethalweepons: the'CI!J~lin~li'!lg (tiXi~tt;1]ce: or widespoo:ilclptw£ny. with Ol,ler~i bm~.{Jll peopl.e J.ffiI c:l'Je~\iud~. rcnKII[li:rng,drul;;'Uy ml~:ou'Clt~1by l~e benents of,lnodeil:n S&l:1; t'llUlihe ih~eat ef serieus 'Q1WiUOnn1t Iltal ex.tenmlities frrnnll unpn;:Cl.~1el1lkldl~~~l\ofpmcl.llCtfmii ;;:l~d OD'J.l~lI!lnp,tixJn nrgo~Jd'i a~l{1 tit,.~vic~·, A. me,mi !1~ri! I ,t~ lscussion {I f 8&T ~l ~tlti.olwt~l contOl!'l:lW(II'.terlruriMl'Il' .1~~cl. heinous crime cmilIHJ!I.oo (rn::~.l,Ided t~i~~~hni~~::d iu ~p~lt:t:. ut th~ {)L~t'!r two iM\I,I(;.S,d:t:liiHV~ b c.lIR'boratuoul. The dilsl.~lIl,gu isilH}d ecouem ist Kelllm;i.h Sould in::;:. a rare irnellet;."~~I'h~I~:p-;!!~~kld (lfhis til~~~,l'.!l;lj·!1~W !h,lt t\\'O ~~~~1!it:.S w!lilf;\r.,~h~ Ot!t ~ dil1:erem:e III <'I\'erag:e ~11\:l()Hl£bet~\'ee'nlhepooweSl oo\m~lI'}'i,nlhe WQ~k!,

IECONOMIC PR'OGRE'SS AOH[VED

ur

;nu;l themm,t p:rm;~'rom \\I<!S!ilO

.Ilillore

tl~m I.S\~l!t~"

hl1;l ~.p~"i~~~~hl~

eoncern 30 ye<ll"S ago .. he estiil]l'~I.ed li~, i)sbe~n~ 1:50.,.hlCOITtlemld ~ a ll,t&n di~l],,;t:itie5areeven ~J""u'rel!'tQd<!J}'. Dre~i~e 11~~:OW1d.i'!l:g lJ~gre~, g~.{!111l.']~Y; ~h~ s.& T gJiJi bel~"'t_~n i:it~l;'i ~.tltI ilHl~ p t\l.'ft~[JliS ~ro]'iic..'i~)1 1.5 1 wId.er 1'IO\v.tf~.li:s . :gllnll'ing ~:llaSlJlis l~~l~iJi~d~~d, fk"up<!mIlllS t~iii~!f."liidesw.i:I,1 i;llh~bul "no ~V~p1 reverse prespecrs rorel1hiul,1CiJ~g ~mllmmlwelfsre, Ul1lflJnu'nm~~y.EI~eglobQI commun.ity has failed to bil'i~~~ ta:h~~o~(lgiCi;lloPlmrtw!nilies and (i!kills to , IlIn.dBrpr:i,vi.~Gg;;:cl !~Il(~in.l:pov('_r:i:'\.h~t"!JmlinliIl!l1itif~~lt1rO;:o;s thJ.:':gkJlJ~. The d~delil~e ofwidlesp:re.<Id ""'Oli"h:l\\~t(lc~ l~oveJ:tyhas ljipic'lIll.y been 9ddrcgsc~lll.n rolJgh, dol(]:sfl:l'lt1 h9ndOl~~S !'IS ·t\olllv~nicl]l but largely iillcUbNivc ~1<1~ hiilltIV(\','i.. Sel dom Iliave Inogram s ii!1th~)1,.rea erested avenues lor allp~yil1lg 'IInOrefiJl S&l to develop 1[}I;:a~ 1-*iUz.;and cap\3:~i.ly. \,,'l1:i<:111:31IQ'JI~ C:lll !!;eUl!~rzne incon~e and emp,loyrra.l!:::!U 011 a o sustainable basis, A. progrn~l bckng spearheaded by Ihe .Ellergy and Resom'(:es~nst~.huC e mhodie!'~ec~r!.ol[lgi ca.~innovat inn ir! th~ <! Ilied fie lds 0'" agr.ic~~hu ..e~lergy. na~ur.1i ,,~ u-e,'10Ul'Cenl<lll.mgemem, andl~ufal,'g~tion tedmology il'll~a'l.ll[le:L'stililpw~lh ~oc<l.1:~nreplieneurs i I{l r Hl~~"l!~lai!]",Jl,~e\'\rd L;li:llg (]:ftlllr;nl c;:mtnrllll~it~iil'" ~1IiI Asi,l<,und A~~rie'l_nlt\, ,~pproacll, '~hi k;f~ 1';11 i C .huegnM ~l1Ig New and Susta~.nfflblc Teehnologies for lEihll1imuio.~ o:f~er~y. meets a clull.cngc nh,lt 'C(ll~]d heconre ]nsuJlliIO'u~nubh: if j,gno[ed [h!]ylonger. Cu;e,Jting oppo~ll!liliesj:"O'I' the rmdwtti'iii~<lpp,lncm;h)ll ofS&T by '~he[lll{)s~i.!>po:\$csscJ COIl~!illlll:li[ies. orl.~~worl.d is 11task d th .. scienrists ,lll!d [polic}'llliI;lke:rs l'I:l,tL~I: embrace with ~1r<Jent}i: n A~.ltt'llg the ncgat ivc eXI.c:r:mIHt ies created by humau f1t'ti\ij,tliC':>. !~C t C'~nmulati \'C emissions of greeehouse _g.lse;shEwe 1100 bY[t~r fhe most serieuscen se<lueDl{)e :~nthe form of glo'lnl~. cHm'lIte cha:~gt:. ClU~S,~rraellm:US$LmiLlii rhese g;1~~~S rc{jluir",r1J~hnnk,g:tel!1 :un:~i.~I:;:uiv~!\. 5taD:.iliz(; 'Lil(; to !l:om::enaraliol1l ofgl'i:.lenhQl1se· gases. :ffii;ec,m~e I ~'1ei,l,llip'H::~:; rd U:l:Uilm: change wi ll COIIH i,mle~o.l" !D c~mu[ici>, ad~pt3th:m.lmlC'i15~1.rc:> wi II a:1$0 require d~.1.:·tii:tncliy app li!C3li on of 5& T .. 1.:~m~;:e'\fC'. these wil ~n,m u!~e 1113CI:· ina 1)01 icy \~d.cumu..Regtq~:lunyandfiscal measures wi Illil,<I"~ to be p~ll iln .pl.,.cc bygov~:rnm.ell!b ...I~It:mt'll\~d: ifll~~~:;Ml:ry by lli1llltih.m[;m~ Olgr~~IliiI~liIn,;l(l figg~rth~ t developnmen~ f1lliCIawHcEII lon 0 fal'Propri<1.te tC",hlli01ogicilll suint ions, Th~ 'kg~n:~~ for tl!l~ glob[l~ )lcil;ll!l~~fic Nmflnl~~~}' is very c!~m-,\1\h; IRU$t rlW!{lgn~~e <lTI~~ eva ~tl<1lte most ern leal imped:i.D1Ilenn:: to ~hesustainable w~lfllre of hUll'l.,m:oociely,i,1Il clllRlilllg the \'~J[i{]1IS 1'l~1l;;~ltS hllrn.m~ il~ mt~ .glolimJ PC:':'IC~. ,~.~:~ur~iolJs in [he ~fiI;llr~Hu~al:ma~ {If ~:!nh'~ to .1 b m~l.urtl&sYS!'(\!inS,and t he .grO\NiI~.g ap between rich and poor. These thwec sets. of comJlili ens flrc g ~n;ti~nltel}' il!ltedi nked, req~!iL'iing.. connl:im~~ped "ppro1!;ch to solve ~hem..S-cien~i~s mm:;t \\'Ork wi:th dtlC~:l>~.m]·iI!ilHkct.lo devi~1i: :~L[ ~{li1Jal .. j:'iolicy Iljilli:~i:SLJrcs, Iliat. ~1ttJh~!z~ tf~si mbl~ respolii~t:.~ .il'i i ~he fc)nfl'i .[If deV~~(lr:rlt~n~ "d Jep!'ly.rn~Ii.l 0 f suit,~b!~ S&'f S{lh~tims in these .,lrl;lm;:_ m

or

-It. K. P!l(h~I[flri

wvtw.sdence

m <'19org .

SCI EINe IE VOL 31.5

16 FEBRUARY 2007

WIHD

BY G~LBnT OHH!;JI AND JAKE YESTON

16!O,CHlH" ISH'"

SquarrhllQ ,a Cube
Till~ d @t'~l;;tiion III
~,[ll ...ir(l

rlm~ ntijll" ,(h~'l1ig~s i5 '()I~ pij rfllnTUlllrllt inil P I) rt~lriI ee III r

host olr !'Il,e'cha!'!is!mS' ~\liIa >ev(!ll.li'@d 'Wh~dil s;igm!l. d@tedio:r!l is inJ a;ulijJled: to ,an alffiI plir~i(:a tiO'111l'e P 'ili'l OrtlN le in crease SJ~nsih"'ily ,1:1 iildl to reduce $ re.~lpl)ns~ time. 'The ha,c~er]!ill regulator er!umamte;;lIIld nitrate reduukllll (IFNR) ,rfi!i!J~II(lI'le~the lnlJi!sui pti'Qn (lir !fII(l!re Ull~ n 10'1) !Jelfllc's in ~e~PQI1SC ~(l'[h c tr ollllis;it:~l)rl hm i31li1aeml!J;:c aerebic grcfl'lth; mvte'!::lliar ooc~'geontrilggNS a CO:r1I'o''erlilelii ~IS l[lI~eto or
<!J

mk r,obes,aJfu;l

45]'2+ (l~uI~ttlr fnlo ~ ,[2~e~2Sjl" ('lu)ter.l~dling III (Qllifo[lm~t~()'nal (iliang~s in Fl'lJlR:alrtdl mSflCldati:011l '(rom DNA" Cr~d;; et al. re-porl t,ho1t thi s re~ctiQIl InJ@tonly ~c'lecU! di o~y' gen, blult actu,a,~llfuses ~uo amptHy the ~i9Inli9JI,. on;$ull'iI1Iin.g:som~ c ]ndQiillig ~I). ~'irst ~~,pirli ~i!iin~l d~~~til;ll), ii$ oi'i one"~!l~iI;trl)rli (fxidadClii11 of th.e 1[4 re~4SP~ duster that. nalf'lsfo rm ~ it inllo a [3 F,e4$,11" d~$~,ef. o1!11d 01 ~d!il~U,(; il!.fl~ly~si:; !I;Qn~imiiedthe QW!J1mdePilllld'ernt winl:iil:lenlt 1,0:51> t:l1le [4f'~·4$]dill:5ter ,Hl,m tine a p,pe..ral'l~e or tllHl e.i'Iii'"teefe2;.. and~he ~llIperoxide a n~ml (02 _'.). li'h e sece Iil ~5:~eiP ~.!> ,aloolJ t il.(Ii t.~ll'Ie 5 '>~e'l~ er, i3i1il dl i3I se(® nd re' de jlilrt'S (~~ hill"" iin!h~ CjQIlMll!r~io:f1 the [~f,~-!liSP'" duswr 1nt);) a 1[2fe-2Sllz¥ ,Llu~tJ~t. S;1[]periiiKidl~ 'lis killmi.llfi '~[) IJlllidergtl d~$mI)Jt.'i!~ PI Imodel Oif FNIR"l!'filth it~, INA btl'l(i~II g, !ti'QII1i1Ii!1f1 D ali!: tlkllil ill1ito o.:<y.gen, <in,d li'ly<I,.ogei1i :pe~Clixid!li,wl'ii(1'i ,eai'I ~tSieU di5mutine ·tiDp allflillltille ho.n (lI'ed~-lI'1.1I~I~'1l1' ~Y'~Uow) cil.U;'l;t)~r alii: tate irot!) 1))!Y9t1fl .;:!lllIdiw<lt€ r, l!'l ~U!lD,(In!! 02: IIJIQb~(ull@ (Q,n t~i9!~@r t:he bottii!lm'. the- disasSlelllllily Qf fOlll,r 1.I'I~e·4S] Ii:~u,sti!l\rs, GJ( -"fOC. 1II1:l([,A~" S.t USA '1.014,2009'2(2007).

n~

or

or

.", oA

Re!(ogl1lidngl OneseU:
Se.ll-i!mUllj)i!tibmly. a pl;t!nf,s rei!!(tiolll of pellsn helTil il e~r or a ~lo&ely r,ela~ediUl(!ividtla~. pre.. \f@I:!L'Sllibr,e,~dff1!J, which ~ III ~~:ad ~(I iii ~DSrsor h!l'ti fNl"Y{! osMy aM I:l ~,ete;ri(lutS (01ll1 bilJlal.io:f'l5.o'f reCiE!Sisive<!~I.e~es./lfrtJbfdof)'£fs has berome seU(om p,a~~h1.~. Ij'j'hN,~a5 st'V~r.;1 Cr[)sel~ Irel~llfili ~P@(l ~ includling ~j!lic(;I, r@rnain 'S@IHn(oD1pIlItibte. BOlh~Jl,e 5·!ocus rw~plor ~~fi ase '(SRI':) and ~1i1oe5 -lecus prol~il1l n (S,PI1J5CR~ are imp'li· . (~lied inUw ph ~lil)m~U()lrll. 51lDinO salo et a,i, hme inesligatedl11 eo inter..dlon belweer. SIP'l'lana SFlK in 8'roniw, flito pmMn~ of 60 snd JU)Q klJ welre prevl{I!Jsl.y ~h~\1,IntQ bind to 5 Fi n, beth ere f~rIIl S Qr SIR:R, with ttle S'FIlaUe'l one bel H9 ,a tru neared, thOLl9~11 ~l:ill,m .em!,) ra ne-bou nd, form ,of t heh.!~H-e ngth

lr03lils;po~~IlI.e ~lelliJelilbs. UI!!'Se dmer~iIlces iIilU1E!' S 11Jl~!.lr.regii(lll! :5l)1gg tt:~~.hese qenes ha ..e ~~t t roU.owed (mre~em Uajedor~e~ a~lll;!'r tile less >or s,elf~in((l!l'lll~tibi lily, -llI,il,Z
P/,tmfQ1i119, lo.,lJl(l51l!~lO~HB88,tJ9;

lO.1 'WSJ~pc-IG6.04S199 (2007),


CI:.I.l
Igi O~.(lG'!j'

ILilfeWi:tho ld Amylloiid
'T~1.em¥tol (j Precursor PFi)~eil1l (Ap~) is ,8 lira nsa memofalile JI).r{l~e~nhat has been lvn~ed w t So(lllll€ r onlnS· (ill f~ nni'lii'll Ali: heilner's, ,d1::;e~:5e, !but rile n(lrlli'lai~'u IIlctlOI'l Oftl1 e protein, lsa mystf r)'" The ,]eullIClwd~ IIWfm Caq'nr:Jrlwblfiili.srl'~fHm~ poswsse~ a :'ilngte APP're~all:edl gem'.I:enllEld ,Qp/-l. H'DITllIst~n ~t al. i!J~i~~ tl'!"lIli ned the FQ ~e or apl·1, w~~kt.iHXP res:>eU in a vc,rie-ly o~ tisslJe'~,i nc llJldl ng f1ellrQm. 1N-tlm'l:>wUh a d i~,n.ip~~(}pl-.l gelll~ !lBi~d.';ls'~I"\i'a~, exhii;lihlll!l d~r~t~inl molt ~flg, 1@\.QmotiO:r],,(J111U;1 rllOfpliiQgerleg:5. These nn'ulaf'lt~wu1d ~e·rescued Ilyrh~

e~1pr!SSi'Onl of t_I1E!'.~olUlbl~ ,e~uace~lJd~i3,r a ~nl ~om Q, Aft -1 P FIJ'l@1rIIf! n€~U"Qh5.. I n ;,;I"" _ 11JQr.:, Q1l'~re~Jpress]flgthe APH IPro~~irrln di wird ·type i bo1!~lkg d was detrim1l'llli! ~ t~i ~ en@'( I. COli ~d rQ1.1!l !:Ie l"llIiti9'<IIletl!:lv redurilnJ9 lhe t!1!vel-s r: SEL-12, o \~hidl:is ~ C df[jl('l'llS liJ(]mlollJ'gl ~f Ii'lr@s!~lliilin. '" I'l e ill.yme that d,E!<IIVJe5, A!P,p ln hum iH1S. nHIS, it .el ppears ~h~l APL-l P~[I~Hml:;1ilr! eS'5elllti~LJusc~ij[U'l d'Ulr~n9 d:ev~.GlPrhe nl UIIItine ne.rvOli SSY5t:em,

~'
~

'..;'

'i
;1ri

~,""
:~

~ t:i ,til

I'nteode ri~g,w1~h .API",e~pression en 111 odifi catnml~ ~II'P1'lIIienl'5rtlfl"Y therel ore have lii nnn.-"

~.

~ended and une;.;p!!cte,cI ~ol1iseClIL1el'lC;e·5. SMH ~ __ Prt;g:,Nail- Acod_ St'f. 05_04_ lL'M., 19H l2007). :~

« ~:
~.

prOolei iii" The la Uoer b~lld s. ~.o:s Pl1. \!'tit h 11~h g (lUi Ji!~~rihe reas the ~D~lII!er "'Des· !1Q~, SlJg £Iest" ~fl9 thaI tlaey malY'fum::liiiJfi dirh~~E!·Hllly. herS III a,rH3:r'Oo/~~sei ai, h1!i~e ill'lYesti'ga l-ed !.Iu' 11!Io1~fI~en(iln1'~ and d:eg.r~d(i~.iDlll iI)~ Ilhe SRk: and S(R' 91;!f!.fl's. n~!L'y~ound th,a'tio Ull1l1Jlililllfi~lJn ttl th~ ~ilI~ly~~qiJellc:€dArab'd(jp~s (CilLimllia.-O aUession,. oolll !l!en,e~il1i the C211.. ::.E!s~iofl'll"ltl"..~ m untierg{lne e.:d'~ !ilsiv~ tiN! rrrallllge l1Ilent .;;1 rid :;,~g1i~! ka m. pa ns (Ji~ Ihe :sri? 9!ene Ilave tleoelil dele!.ed. pMsiiIJl:f [lim.ugn Hie ~i1ISi~rttiDn~nd ,dE!·re'l.ian 01

lih;fMYSliIggeSJt!> lila t. a rising: (Oll,(fFltm lion or ae,rwo~~shOiL!td reQiUCeW~f!lc!~jl'eeds IxllQW, due 111.a 'te the ina'f;;lSe im ~ILIT~plli~'lfk ~taibiit~ r r.ly l th~t. such l~~diJng ',~o~~d~U:S~. (~\nl'5tqu:en't~1 range rm~ decreased ,~",apQl'atjoflar.d ra~flfaU.o t a mulUlude of soec:ond1!1iY imp.~ds,mdl.a~ :reduc:ed QI{(iI~a~m1.y1 wi 1l,{IerllelrgiY for the giell!'~fali{l11 or 0 ele(uicij~1f a fld d~l!J~asedl1ydrO!1!>l'!cJ~icPQ~.en~l!It l )1icoiMi:in <i fid i?:aLlrmOllillus-e ~ tlnj\fe-d~m.ei1s illfla~ ~,O!1l\~Hlte,r model. al'~ Sdte~~~le d,ata, to e.l(atn'lfin,~ tlw p\ltent,i~l ,dlel:t!l 01 (i~t(lool p~,rtkle-s on the di,slrub!J~ijl)ll Ilf ~'i'i LtI s~eeds .ililld dne IrM!Jhiin g JeedllJa6ks F to predpil:alil)!'l"\'\!ater ~upp~y,and wind @nefl;JV

§
I':!

:~, Ij. -l: t. ~.

il
~,

:!ii' ~'
~'

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~
y,

116 FEBRUARlf 2007

VOL. 315

SCIIENCIE

'!IiIWw,sciefllcemag.org

EDITORS'CHOICE

amnss California. nne~ riWLdthalt poliUlioll tly a~rQ:Sl;Ils nlC'ly h d&r~\l'Sul'Ig to~~ l ~'liiild~ Itl~ up to -

more complex PgU~'m~can b~ tr'~~IIt!d Miith(ld~ ~li(;D"~l rl;lllJ.nd!s


Hilll1'll, and

of motl'!{lJI.es cmUle'
P 0:5

and, wgelne r with IhE!'Soe(OOO iooirel!:l aem'l.Ol e~fed". !lIaoy IJ~ n~dlldl'!grm'l!:illltatil)n by 2~(l5%.
Mi'fI TIlJ~e ~«'~(I:!: av.e ob",ioiLts i3lrJ1c1llil1ll¥elmlfi.e h pra:cti·

of ~;CI p, -

IIp{!l, Phys.. LeiI.9Q, 111.10 &3.1 114 S752:50 (200001».


1

gl"tJ.!.Ipl~ {J,f lik f-.·-m dtu:llm vel.er.,> in u'.h 'J,... ,"J <:1If}ve oj" [,f"l rti i§l,§ ~Nl

(U:';>foww~;~ceilem Wnemde.'l 'J'm! Je<I'r:if~I~S, .mel e{)l:lgC'niai

w.W! }\I!..t'\S I hls

Com~;:e;qJiU'te lht:
)f'f:.'iU·.

W,O{~(/

~I~bu w.W

rrnf.i d~'s(;avm'")'.

(al {:(IIlS'eCll!e"Oes.. II}:lit lll~!f.a ~50 nli~ hl' hepa rut po~ihl[ti Iy lna~.Dy li mmn 9 '01' ~e:duoenflg aerese l pq)llut"!!l!l, C<!tlfomi'1! calli lesser! fu[ur@stro1~IlS all ~15W<itf r ~urliJly 1'II1liwi oo111ydmelfcllri( prnJj~.rgerm a~noil'll sy:'i~ems, - tillIS, Geo{!hfl~'. WI. )3, l2:481,,:j (2:0061
MATIE IHAILS S,( !:IEN( IE.

OII[;I!.'III·S,TRY

[)e~il~ md,e~pr'li!adIli!Umh toward dev,elop'mE1n~. of IlFhllllall1le~emir.mn(ju(tors~'or llarge·a~ea, fte')\i ble ele(uo:nics medi:a. rar ~Il!Sli~tleIltio!1 hOIs been

for nm:uF;:lni !"w;;nm. ('.]Q)]OfC!"5 for i(i(!j!.,turies, rt"O"llf1 the e.'lJstem gq"ass!<ulds

r~xp!m\f! Ti'bet. ",lila.ce orKcI.~Ciim~rio.l,'1


&.

'.I.lbc't.:'U.:t PI.'l.teau .~ulv11-\2,2,200':

~o I.he· heart of·ribf.l!t-U1.o1s;;~

more ~

Illnse'r1,e,dj ~111 isotalt1,on


MucroWMa cl. pri i:II.ifn9! (IJ!'CP')with

the Ilr~l!lla[lile (1[1110ut~i 109 materra ~ requli re d IDr tile wilri Fig a nd con lfIection~ Mlitlil'l ~liId1!iCl't:!lfi ~~5. 8Qth gDld and sHver ~o~e5'S ~ugh
gi~erl to
if lasteme

Gal3Jm}la;O~i!1IIs,~ru1.ds,
~endl!m.nledu:lili~S· on board 11m ,UiV nO',cr v.~.-1~De expl!m·.iI"Dg I.hefllRdn~~ri~.~~ C;Bl!11l.1!ll"g5 ;;IJ1"CihipRI'i[{n \',~H'ln''l \.\,ulc.i iJeabO"L:~lids:! 'ri'lmH $4 ,H)o + i!l.ia·, F
.~[IL~y !U. -:.10 ~(!iO'f 1 DUilro'\.'!Br 1J00nvin's

monotayew!; f5t.1Ms~ onsli rlaces, bu~ it (:atFl <IIbo be uselul lQ(;~eate mh:"e~ SAM~in ~hic!i lOne of ~he ITIrDleltllle-si s, di ~l!'ted \\lith~1l tmt'1 Qther la~iIll'".
~~LllleliDet al'.5how

~t~mp:;

"~I"~!JI~edto p~lt~rns>elh;i:S~~lTIb~~d b

ric

conductivity and opefali onlll5.1i'IIbility, but pred5~ pallt~rlfliIll9~¥itlillh~~ I'liId.llb tYiJiGa1ll~ I'eQ,ulrre5 vamum dep(l5il~ol1, Wu eloi. ha\!'f

M.,

de~5oed

~.111!11...

i'l 5!i !Ill pte

50l~~1~n-bills.ed 11J«e5~ IlJ:r pat~e'mil'l'g(.ondu,eti'lfif ;~lver


r-eal~ur~~ Ilmgl :iubstr,g~~,

that morl'! di rfu'5i'l,l'll! mol!l'(ules, Io'l'lli<.h


01 ffi~i,J11l t~ iJi3 t~em wilh !J.!(Ii', calli be im'IHt@dinto intire <'iFe!)U.e
rI

U~il1lg(ooling,~la mp~ng, 'or ~rilfl!ting tg"ihniQt!Iiii~,they


~[)tl!lti~ucglll-

d~flo~it ~n ~koh(ll

talllling .a 5i,lver(U saU, a ~ll~dro.xo yall~~a no; il.'f, a!l'1.di lOFlglo(hain mfo a

;,., J~y 2;,ll-Auf,"II.'lISll 2o :Z~01 Di ...cm.'f.:il:· me: of ~hc w Iim::SI ~ <LI"e11S :S\vil:L.eI~;I~C'i;d in Jur
j•

An~alkill theS1¥i:s:s AiIls,

~tableSAMs b~ a
lIfu~tiloii;ll they t !lIm mi.c:r·(I(oJ1,la<!

b (t:ql'lkil>!:id. S UbS'llq,lI!Nlt heliltin~ a~ relalwf,ly ki~'1t'i;:nnperi;i'tu Fe etSOOC) f,[)ulTIl$ tihe (lOoouni ng silver e!lemf!,I1it">.The am~ne riU nco

\!\/;l}lIkirlg: Ap!_)em;e~l,m."Id

OiIllp~, Lucerne &, SI (~JI'[!l~]L

l~ngelbeTg. plus

S~

theMgh

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!E!i~~i (jo::im:~5(100~~_"i!II ..al-"lk~ [O~ 1!ml!:~J'PI!_M,I,!('!!l!!i1i _Andrm'M, '5~iIIn,;5&_ !EI!IIUfiJ:!fiHn\!B ]ulm, ~iIIIII:!r'itlrnj)' 1!I.1IP~~riril;: ~(jijjjtjiHliHtiIiI!lliM(jllnr~ AsI\I GG~-o'a: +.!iii ~)))222 !i~~ ~~1lg~,. ~~'h ~- ~~I",. I;ln $. QI~r~. S1~,,~ 1_ srrn~(iII, ~r;lI~r S'[(l,n~ ,!L~QIJ!j '11!'1T9!: J-o.li!~ l!.l~Q~ !I!'IT(II!!;IJl~W!'!I!!~ JUi~Q WI1~li:~;

EdmOflIOO" :5~rlq''m:Ur'I'!Jl (Ill 12H ~,2oil5.2!>.,1'AX ~oII tN !l.::!~ .32.&S12;5\!il'5,Chri~h~ iiarrfi!OO.~IIi~~1'Ia I!a~~ IIIW " •.»'!i!.l!!IJ'lO;uj~ ~~ 1_ ...~J~W1 HIIi!!I:lloait ~1 (0) 5Z 7~~'~,~<OO, W. 'Io1l:l ,[!fir) S,2 'l'~"~36Cl; i!D!o!nT~IIO!Ii ,Ijoj)CllCIlQII CftII\I.Ii!lIi,5!111!~ M!oral1i T,ompI:im~~~lllf'{!! (1Iri5llflo!!b1Jl' AmI' iilardtlistre.: gLffJ" 'rK!{!!5,oI.l1!rrn>.!!]I; PllI!0fi 11lI!d\; Mill)' ~~1Ii
,~, 1Ii_~III' IIiIImIlll1_n.l:5I11W\,aLioillGilbert".limem; 'P'!!;i:i!l!'l!!T ):rli~ ~: !i!Obli'trl; nili!!!t:."'i!J~·!l'.Md !ill~!lfrr~; T.!>!MY!li!lI! !»rid' 'E. S,'I1\i'I!!;' '[MP' l})K1J1I!IE _!alt ~Jlin, I. lo!:!lI1ntlr, I:I!!I!!!I!' ~'Cdiiilo! M.1~lJlJlllJlB;]tilvllE, blJ]~.:lr:nn W. fJKIIJ~t: ~il!Il ,", fiUilalIidt Alia! Cl!r;,t '1iNlma~ PilIL!rd! ~~~r 1- st.tng: KlilhJYII'It ~[U~~

5'.I!UU n'!I!!!!_!lij~i rik.!! Elmr~f1~ IWhn E 'iJi"'9!1J11Xi!l~LJ5:.r.w,_Ii!;Tfa'[lI'IH!lfm6;~4

Ij~j"-QU igry~t.Ti~20H~'~33~ !!!I!~ __ ~ MdI\~ 2m~2iil,61!l~3; ~~~11fU.5Ii A!.li~oo MiU~'r; 21!1!:!·.~~6·~S:'2;!IO!mIEIO.i1i Feri1'~nd~ ]UII[,((; 20,H:M·6i<iiO.: Mmt:ll1it Mney:: 2l(ii2·n'IJ.~57~:

2.01·3M·6!)M. FAX 202:,2)8",,67<1.2;

iM'!:I!(M. UJlliillUHlIII,

il~~ or. ~(oes~

Oii!l!r,it1I~~ "_'!LI;JM;~~l'I!!iwl~~ Clrillliinl.t IPM M,jj'I~ ]nwmte; IiD'I!:i! CoII1ifi~Fii5 _ D.lniel.a~(iJ.W_(j"I!IIIhm VOg;I'll ~ei1'iI!G+II~' '(0"1 ~;II!!'~' ~:?Q~. f)l;!; +II·NO) ;!l!l! .~!l09 &~~S); ~(II ~~!I!lI!!1r!1; te!ll:L1_!i!I1i ~o:;ltca~l ,l'llIl.lllr ~l'-.tM~, M:miil E!lS!!lin~ ~enhlTll.aoo ~oI!Iii;),};41~BiIIliIn110B (iJielllBl' Alii JIlp.n om~~=.I&ii, CJKp!'Iii~(I1.ltil:o id'rliikil" ~~li!lol, Il"it13, ~11r~orrdl~ (h~~O~shI. ~~. ~41.-004~J;lp.!ro; 010&1(00~ 62Jliil ~21'l.lr,IiX.:il «I~6 62i!1iil~m; ~~.'@~L~r.j!1 ,I,I1;i.!!IlI!!~ _ AWlil~dl SWII!!' +&~ ~ .ti62 SUII, ~~~flII@.l!il~JlIljl; ~__ ,,_ iI!i1l:I;!i.I hnrril f>lDlmI~, (oonbibu:llII~ (0II~:e!J!!!1III~1l!, T811111~ l 3'391 rIlb~!J. ~.AA~,1 ,(OJ 3 '$Y~~· ~53l: doorl1llk!@~oII,fI)rnl.: tHillA I!''' ~~ ~}lj0\.;. ~6 (.DO11;1~3~7 ~l'!l'0!l~3!!lr,l ~676,. ~~(ll) 'l<!l! ~~]' ·4-l5'1\1 (n!tf~·om~'l!ic.((ffr. 1~".1I!\Io ~t.l. '(coilIJIbu!lfII,J OOIl~ooWrt '1-91.ro~1122'1:!L ~~, ~b.lgI.ltf1!V,n~(Qj]]~ ~1'.;)~trJ)JJn'g,(c(n1r1MfI,l~~~Q!fi)

~*WI!

-------

Boo ~ R E~ I~'~ IIClHD

tg~~f:~~: .. ...L-m~ ~~.


~<I~
,Il'tlii<,~!"noion'

IEDIIH

I)

BY (ONHAN(

E. iii Oll)EN

SOCCIER M,AN ONllDilSPLAV


11IInnam~i opoe!mfm9llhisl,iee~.Wi VIr~:mJr~a,M~ of Une- kllil ()f 'wliJr~rI·lan!l®u~~c~r player Lma1'1 M \ l ~ Timranl. Hi~ cr.ll'liUlm-lFe1il'roou!::ed IUl;ing a magl1etic reSGnam;eo inni31ging ~(allllllerin ]ilnllilrywm lfIe ill!l 11111 i'IY I]if lWO ®th!l!f FueniC~lmt'l!I~HEll 1:P.lIlliIJlYl~l~illiliSl)!'Ih!!'r Refl,~ II)e~tllIlft~ an!:! ,a {{Imp.<! well-pr'@s@l"!lI!ld30.000"VE!ar-old m1ii11!! C~-lII1,agllloo klssil, (lin disp,'lay [or the~irsl, time, liu~ 1li1i15~Ullll\\i<mt~da lMIiI9 PS!rS(lIil'$kull ]1iI Un~~xlliliJ~it.t. g!iCk~ T!Ullram, a r~~mpaigiler s I !I{Iaiml radsm aoo 5®[i[l1 ililjlilsli~e, as a lOym 001. Qi~ Ule unily ,0:1 hUnl aflkim:l. Thmiill1l, im ,m fmerview irl the F·@bn.Ji!),'iss.!!l@'([!!rSCif1lCfS 1l',!AL>\~ir. ~ys llill;!@dllibi! ~~h{IW~ ~'nj! ~ll rrom ~1iI@ S;]!lD@ ram~l.y, w ~ Til E! c~oic:J!21~ a t5Q a 't:ol1llllIl~nl on ~h!!' m!!!seum '~5eWI1I hislorv, says. ~1rl:ho1!eolo>glis.lN~lha!1 Scgllal1lgeJ 01' the N1~!ioHal ,llI!still~lt,f[[Jlr lR:eseilrd~ illl IPre\i':.f!JlliVE! r'cl'llleo~JjgIY in Pari'!>. The A .Mu'see de l'Homme lol!lg harbored HI@ ~~mailll!i olr S~arl.ii@ 811arlm,a II, an early 19th (~I1Itu r~ KhiS~lrI \~IlIIi1'1"m h~m Scm ~h AJrnc.a k nolh'J'I <15. the ~lhe tilClU.e!'!!~H VeirUJlS" whO' l,!'taJ~ lakell .W1IJlI'HI E'l.lmpe ss ,;;I

dig il'!>"!;IJ eh as H:l23.:1" 'OrLll uln11 ~lPa!S5~lrortl. ~

Tile MlI.Ise,e d@ n"@m!lJ!~ i'!lJ Pilri~ has recruited a supl2rstar. A 1i!@\'11 e.'l!hibilion on the histlllJ)I IiIf

"pa s:;\"d,~ Q r "test. (u~~ers.a~ Iheo wor~ ~hcmlll h!1lp security iii dmi rli ~tr~ters wmbal mass 'l!l!1t()m at@da m!.! ~I!~, ~hie!' m()~t mmliio:1li lypE!' 01 hac~Hng.
w

W'ay' Back W'e',ilth,elr


Three f.u mels d.iirrI9~ermm 11 rolling storm c:loL!d ill! the earliest knowl'l tornado photogrr~ph (be lm\\).•SIl1~P im ::!LaM lfIe~r Ho\'!i~rd, [>eel S.mJ~llD.a kOla, the scene is one 01' se~fall hUlndrecd vintage ima,Qe'S 011 di~.pIa.Yi!t lihi:sgalle ry r'r{lffiUl~ !ilLS, NI~110illa~ ell!tner Service. W

ctJ'rioshy 01111 Wh40:!le5,~e laton, d !Jf nhal:s, a 1'10 b ra~n well"'f on d i:;p lay ~n the mu~e.URl aI,te r she d~ e>d. Sh~ \\'<15. re'tl!Jrrled to SQuth A~ri(a iiii :;HH)'2. ")(II~rljh:! i;~mr;: ill' ihrQ!,Igl~ '" r,~d:;t,c.otcm lal ot;;@JIIleoxt as ill plijl~~rve obJ;~(t ~m dtsiphlY, ~ SiI:n Schlanger." "lhura,rn~s <I mmie-r!ll frem:ih!1iiifl who hap[IlIem
to b~ b~!3Ick<lIl1Idwho aIfirll1ls

The 'ullu~~~'ti:oni1iand pllllJitQ.sdate r,nom the

What '!i~w.uIIGn·tYl)'Uuse as a computer password? IEngin@E!riMiche\ Cu.kieralld rolleagues at the Ulfliii~r~it~ Irl Mal'Ylall1l(j; (a UE!gE!' P.ar,k, soug otto NtlUlilda pmfile of a l'Iaid!:.er b~ba",jor" by IITIO Ilitoringl~ollr Lillliu:q:QII'IJ>U[e:!'S COllln ~c:l@d to the Iintemet. O'll'er 21] days. Hli2fE were

26'9,262. liIadrer a~~.eill1lpl'5. ~Root" was Ily ianhe mosHommon ~U~111Ipt@d !Js~m~,m@, tri ~d il'! rngr~ Ul,]lJl! 12% of th@ tlU'@mpts, N'~d. {.eJm@ "admil'l," "test, " glleM, ~ ~inlo, ~ ~(ildrn,~ MUilysql,~ "user, ," ~admnllljsU!3ilQr," and "erade." Attempted 1Pi3IiSword~ 'M?r@ ,@,V!r1ml(n!! banal, Ilnl·~,ddJit1onl to' Uyllll9 t ~e' !lISf maliii.f, rn®~I.went W~I~ll511Niall
M

e:a~ly 1800~ to the 19905, r'e(ordiI1l9 the 8'tf@ct~a!~UlJO cis. 11.1IIrrki!~i!S. bllina rds, and ,olher lypes of exuenw w!!i31lher. Vis:ilms can r~li'!l~ the dIUS~ Slo:rm~ lhill sW€[p!t Ull~ G r~t Pl.aif~$ in the 193.Qs and vi,ew SJorne !OI~ ~IH~dOlrn.ag@ hlJl'lli Hm.rvc;[!!'!e C.tImill@, llihkll )wi!!IiI[lEd
III~ GuU Coast 3:6 yealf'!; Ib~r{lfe' Kallrilf'ili3I.Tlll~ gi\i~hHY ~ lse recerds ~dVilr!i:@S i n \\!~.~ilh~r'ilbser\l'illg~ lec.lmology aiM l1otcl.~willi11l mig hi tie I.he oldest uis,ting radar ullllage~ or <I weather ~"'ent, which~llm,;, a celd ffrOl1lt bfowin!) lor.ltarll B05.lOl'lfn 1941, » \'A'H~,photo' i b.lwa.'9 o\ltll~Sl(!rLdl''N~~nde)l, il l ~m

Seed Bank B lueprints Unveiled


A 1D®Qmsda,y" seed lIal1l~ desi'limed ~Dpresel";le ~he ,"'oriel's agriiculturai d iv~r5i ty, will b built Ill' suml,l~ tlm~W(lrsl ~@I'W mo::; r 9 loo~1 W(!miimg lor Q a~ lea$ll cefituries. <I.{(CDrdilrtg t{l <lK:iI~itedurall pl:ms. released La5.\ week. The $~,.a: l1l!lilU!!JfII S..,al~md 'moha,~ :5E!edW!1J.dl~U be kKoil~e'd Q!1l~ fl!Jo.rl'l'~gi~'n isla'I1!li1IUiSlWOO ~iklmeters rmllil t!l~ NMh I?'dle ~("i'~l'te'. :23 ]1I1n~ 2006, p, i'f30).lh@ p1il1ll~I;alll~flr two dU;:!IrIllb'l!rs dug d~'iilP'U!ll~J;I mQUlilliIil'lMe ~ 1.30 meters .abfil'e $fa leve.l-.lI1Qr,e thafl,lligh ei!lClllJ:g~~l.o'$tay diiYe'Jlefi j:" al~ t!lll ~i;e in Gfeelflloin d and jlg,~ rdic:.a ll1Ie:hs"The dl~mboers. wiUl be l;o!1lJIIII;!cllld ~(i IlliE I!MJ !$f:dt;:' 'Ifi~1!i l.~(I"riIlfW~lo.ll:g tllli!lIl,~t ~This de"iign takes us (llil~ step doser ~:o.gu:aralllleeing tile safelY' of the IjijIlljrld'~ m®s.l ili'lfmnaf1lt Il!lllJfa~r~ouu:e/ s.a,y5Cary F~wd'~f ofl"'~ ,tMo~ai (uop [lhle;rs~ly Irusl, whkh Wi II helfJ (lund ~he o,poeralicolIIi3JmdteCnd~ml,lE ti'lll a(quis,iil~lIl1l'Q~ S~~S, C(]'n<sbucl1!l!!m!hQukilbegin n~;;l ml)iIIth ~nill b~ S
~@

finis.!heod by 'SeIH1!lOObe.r. Th~ .U(fln~e;(!", deslgl!IM .al!Il'nu.<I'fiC'~ witlilig lnat ~twill ~g,lta!'1llli~e ill g@min Ule midnight sun."

~l~nl1,g

'9117

I'E,W -_M8J~E_~ ,
EDliflED IBY 'It'~Otll'IUif BHIIITIACHARjH

«In the Courts


S ElmllilillEN IED, Scor@!i oi !.~~ter;s C pl@!lId;!'Ig fo r [~rlk'my diru not uop tl ~O~ Amgele~il.ldg,~ t I,~~ m(ln~h frCli1! 'S~flillel1!(i ng tile pionee lringl glene<·ttJeri3ip,y re:5e~r(herW'. fpemd Alnder'>OllI to 14, yoo,rsir! pl!"!'sonL,AI'I(bmon 'i!las (:0:1)'Ilictedl la~n summer oif mole~t.illg a YO'llUngl gfrll (Sci€lIc€. 28 ]l!Il'i 2006, 1)"4137), "'II is~ tr,a:gedyfor everyone," saY5' laurenee lKIedes, di f'~C~!Qr t:ll~ Ir1$mwrh~ G.elleti( ~icim' fo'[ at the Unlil.lersily of South em (allifomia ]111 los; A 11gebt'£" where Anderson \-'lO.rlred until h,e resigll1led last S'ii!Ilt!:m~ber. Arll:ler,5o n's l.a\"'lf,er told!

MOVERS
(Jlil:lIlSiIIS, M:AN.AG EM,!:: N'f. lihe Groha I FLJm:ito ~igllt AI,O:S. Tu t';um:ulosj}, a ndl Mil~1i a has ·t1Il~ guardian. MkhelIKaza~[hk]ne, an tHVJPiIDS !JmfFIl!!!1olOQ1Mwho ~$ (!.!I~rMt[V !Fr~!'!c~'s amtJ.'l:lssdor for HlvtA~D5i and (0 tlilm~ lliclIlblle diseases,

~)'r

will becorill~ diredorr of tihe S7 bi~lio!"!


QrganililMf'On blll~d ]n Gene\l~, Sr.11l!erlall1'(l, 01'1 .3:1 MiOi rch, Ther!.! Illd's!)ll.!It-

Ihe murllnlfc

Ill-YEar-oM

Ked!~~Stlll's. the eqLlli'l,i'.aJ~~"'t of

\lim need
<I

~lif~ ~~ttnoo" f n so l]tal!y ~o'llftil'l~ment.

10 be kepti31lparHrol'lll other prisoners,

making this.

illS.

9011'19di refi:(J'r, British

ptlblk heat!he-:xper'i
R:kllaro Feadl~m~ has b~rI d~ali!,,!g with

AWARID$
MAI~ IIIDGil P~Il~E.. Someti mH~5c~he si mplest SQl!I,JltiOi'i(iSalz..o tile !iflt,m erfe(tive, FQ~r l:ieLa:de:o; ago, re-seaf(hers discovered th[Jl~ O'ral r~hydr{!U'QriI thil'rapv (ORn, ~ mi;(tl!fi!! of gIII,H:o~e, $arU~, and \Mi'lI'ter,(o1Jlld pre,ve,lll llati:l?I'iiiJ; lrom slL!c.'[umbnll:lgoo, cholera 1Iind

other diarrheal diseases, ne low·t~(h breaktl'lrolllgh S.1l\,@$ mOr~!l1rtill'l 1 mill~!!Jinliv~s, ec.'Jen


yeaf, prilnal~ily il'!l dev~l,(lping (oI.lI'lU"e$"

p rize fI~ sm ared by !Ferna nd L~lb rie ,of La" ..l Unive-r:sityin QIIJI\lbe( (Uy. C,anada. ~I!1Id Pa~lf'i(k Walsh of llo,h 11$ !HQpk~llis. Urli!{'~r!'!ty '5<111Qoi of M~d1i',iMin B;lllimor~. Mallryla n d,fm ]1l1lp rO\l'illg the m alli~ge ment of profil:at~ (~rlil:.~r using therapeutic ilnd s,ur!jl(al a pp ro~( hes, 'I$laimic sch()ll~r Rosilld1 R:a'ifuJed,. a forme r resea rc her at Fra nee' 5
Nla~i(ina l C!ellter ~o:r Sdef!iliik IFlesea rL';lilI, is. b~ill1g honeren kif studies 0:11Muo;lims' 'CO!'lI.fIi1bl!~lio ns to bas.ics.(ie i'u:e, palif~icu ta rly ma lileomn <ltics and 0 j)tiC5.

allegatioll$lllat rnon~y Wi1IS I1Ii is.:5p!lrlt 01'1 luxuries. But o,b~rvl'rs are,olPtiimfis:mic.ih[fJl I!flizatd",kiM ,c:ai'lS:~£fr the O~\i:a !'!iz':]tio!ili!'lOO
20 yearrsof e'~p~u~encej ~'t he S!l!Cces$hilll~r:a,i'i! b ag:Q<n(y f.orAirOS naOOil rdlll, {lr!e of the ~:;rlrg>e'St (lr!Jal'ldl~tiOll~ in 'the f.l111IJ5, world," says lain S:imp5.0n,o;pokeosper50n for !he Worl,d H~al~h Organ~z,a~~o'n.
ffit hi!! French national

e'xtTeme~y .1fj(.(ampU~~m] ms physician Vlilih A

$if'ilOQU~eorWiiler~,. "K.!Il.U{illkine irS not

onty

lin

:~ ALa (eremolllyill Baliliigkok'l>Royal Palam 0<1'11 g 3;] January, Kiti'l9 BhllJmribal.M~lyadej Oif TIl~iland b~tolol'l'~d lhe Prim::e 1!i~!r!hild(llAw.lrd ~ 0'1'1 low Ph'Y~i( i(!l,il"S(ien fists who p'layed kie:Y' ml~~,in O~:T\ delle~opm~~t: S,tanley S(h~Hl, 6 Oil"~i.i No'lll!'!, aoa R:!(hard C~s.n ot tll~ lIrnl,~d

~ Sl:a tes snd Dilli j] "~dlbl.fl nil hi s of ~nclia, b The ali\la~·d ]~ named aHe.r IKtf!\g ~ Bhurnitllol',>iilth~lr, i?rino!! Mahidolof ~ Songldo'!, a H{!rv{!nj-~raijr'l(!d phys,uciarn who ~ ~~ kl1io\\illla:s the "IFatiherof IPlJlMic He~ tl hi~ ihaHoiIif'ld_5chLitlz tOQ'kll(li1ile '~SO,OOO~Olr ~ tilt' ll1ledi:dne p:rize; tll~t' miTIers shared.

i~~
I

~ S 50, OOOror~he :pu bli( mr aUil prize. Dmg 1'19 e iii a p'rl\1'at~' audience, IKilllg IElhumibol (lJliIlz:zedi :~ t he Clwa rtlee5 on to lliC;5U1: 11ils.stell1l mi n 91 soil erU's.iot1lto ~ed 1.I':€',lhe ~.e1{el'"iIY()if ~ s~ sQnll'~Ho odm9.. H~~Ma~'(1's;~yknows his i scence, Ca;$h1 s:alys,
e

'0 IHIS CIllt:E,1i)11ilL D'i:lIli_11IY~l~ronL II Sumb s Dakota b~simlss~:n<ln\'I.~ho made a :fort!!llr:r;e' in the. crmil eard .in.dil!l£lry .. Vs d,omlltiH1l.g $.400 m.iU.ion to crelHC a world-class IW~~.H~ eatc 1>YSI£11I1 mtd resc •.r.ch cmU~f .UIfI SiDux F<:db. SmJl,(h Dt;~k()U, And Lle Vl.I"..mrs res.e;lrC'h\~Ts choOl"~ a ,~in~!e area thai to

\,'flll receive a s,~gllif:ictm( ehunk of ~hc gHt Tbeprojeot is expected W cn.p[oy


more tl~am ~,OO rrese~ru[he:rs.a md, S<lll~fordl] t.oi)ei'i i.lwiU "cmw;: up \"~JIIi cure" II] dle 01.

chosen area,
Ttlisisll'"t S,mrfu;!\c.il'S fir.H~OJi"y .i!l!i,tosci't:ln'lil'~c fl,.rnibmthmI1J; :hs! ye.n·, h~ pr~lrn-

~ If~AISiAI~ Rilll"~S+ ~ IEl;~tf~~'hem'~:;t!) ~. P ':i Ca nadlan end ocnrlolog~st a nd aU,S. umlQ·


iffi

gri~t a re

oil !lr!01119

the wijI1.rIi~r$of

,~ 52.00,000 King Fahallntel1natiolli;lLIPrizEs, ~ aw.tircrQcc by thi! King F~i:S<'il FQY!i:d]"lion off ~ SarwrduArabi'iL tlI.K.-OOml JO)me~ Sl~dd~nof ~ til!!! L111!i\tler:s.ity or C"lilfomia, tos AI1..ge[es,. :ii: receive5 the5,ciell(e prize, for h]:I;work an ~ t 11@self-.aslle'm'llly 0('1110 l!2(lIl~r ~;tn~(tu r~~. g acomerrstolle(llf tl,aIM$'cienC'e_ T~e mediidne
t!
l!i

th~s ye.ar's

ised to gi.Vf;l g,o~~U'l[)<!km:.u:i)'70 m:illi{)!:]: Im'!'<.'lroan um:tt:rg:ro~i,ll .d Ifib ~. lhe Nat imHl1 f SC'il;;l~ce Fml ndat i Oil chose ,~ts,s ile ]'111 III, a
!i!jlI.~,[)1in~l,;'m~11~~lili'01~.

BD8p:itu]

officiaht

hepe Ms, lilum ,~C't ofgcl~eroi!>.ii.~. w.hi,ch pro:III,PIJ.'tI [he s.~.tr~:X VllllC:lY I:hll';;p.it!i~.SZll:Id [·Icahh SYSI:CliIl to challige ~,ls :lIImilleliO SI,'[Jfi:~~dHe ...lth.'i<'i n~,I!iJ_~fO:I'mIir the '~C'JI1l~r intQ II :Mliofllii.:i. huil:! of ilJllo:tl'ic-aJ r,!t~~;;,m::h OIliLd p.mie[l~ C:f]~-e.,

~ Goli'i 'Upforthfr£page? E·iMiIlI peopl~@a<la5"Otgl

MillT Hungelr Slrikle: ,SOLIr Graples, IOlr'lhe Bitter Taste 0 Racism?


1 '

both A rriC<'lIil Anm.e.rictu:l.s:.",'Dao suy ~111ey neoune cHmmc 3t lhe schoo.~."Some of my experiences duri n S tha~ time l~ll.dercu~
ILCrc-d IUllfLJVCffiC

toy status and

mpreseUlll.lm,e

kind oha.cisn;, ofbio~(lgy

thaI ... Shcrky is opposil,g:' wrote Sylvi[l


S••nders, <1'1<lSSiSl<!rHIlnlfe~~or

J~1I11ili~5Sher~~y., n a

AfitL(:OJ!Jl;·Au~le:r-ic~uls,~n~

by (xlalllrllliLlrree "OClL11I1osedoW n:U;J:I'nbeffifrem


insideand outside MI.T:.'" At dl'e ~m:ne thne, some 2 [I depm:um:nt
:liIIlt:~nb(;r:\ In<JIVti :S;~.gl1i1.:~ a:;t;;U£I:r'iiIl
111 ""Oll'[t;:;.!:LIiIJ:j. I

cell tesearehcr in the Milissac;hllsens, 1.11I:;.~~nu>e o:t'Te->(.inlO~Qgy j M IT) in C<lmb,ieJ,ge. boeg.Hl .1 ~.llU~g!,;'f:>Iltik", It~~t : wl.:;.ck It) pI'lJlt(;;~1 tit!; :;cllor~~1:> deJ,li;l~ nfili~~ bid fb:r tenure, CIOl.~mil1.glillO'l[ racism P~fIY'OO~ part :ilil his I"CjcCl~.(j:rl.Shelf'lc), VQ\N~ ~('IO main,t;lgrl a dilLI)' p~Gloe OW"'~&: the
provm;;t \>. oIIi:t:c H!ii i~.M n "!ldlfillits"ul1i billS al1ldgran!s hj 1m} tefllLll'e. U~Sprotest h~~.; ~i.dcd ill~ d faGui:l), mtd. shilll'ld
OJ

,I

S~u~:~~ey: ... >JII ~g'l!innil.l'i·'·\1!;,r.,.~ rlu i:-i O'Lh.~t'lHt>ely .. the mom pcr~ecg nml:lllr~revucw'll dQD 'I. think there is sucha ~I~ng," says associ me PWCi\rOSl i Cla~t~~ Canizaret>,w.li)o1>e office ~~sked11

berween 199'7 1lnd 20m .. in a 7 Fclml,uy letter (:in;ui.n ill~<UIInl1g Mn~'H:~lhy,n nne c.t~. I s:he says, ~ sel1l~Q.rwln.ilc p~ojess![')j mid hcrttlat she i'!",cl vio~.'!ted mles by brillgi~ s'tudellil.l!imo tlr!t;faL't~lty~undili:l)~l~n" (Th.l':[C i'i no such n.tl.>.) S,mdie:p,;s leHer ,It-m d~d 'he ~1.'l\:·tt!]~1 of,~ C\'\,~hii.,;o~ICiligueflll(;f n;."turjl~Jlg from. ~Lru:;~ootC :SCLef!l;C :~nee~~l'Ig~"Therewere Iil.Of~O n~tql,'()st~iC'm:~~~~ llrn~r.~, Ut~ the onb' bIack pml~l!~.e! 1

saw were I. IIIe wsiters. Why is th!l'lt'P' he ~Ih.'}kwher dun mng ~1iI clevster ride, Wilen,$he responded ~ili' 1.1~'iiki.n!i: hiuill. "\1ilh" thL:l~ l'IO few Arri.c<i11i Amel'ic!lI1!!:!:HI. Mr.l'!·' "

spOil ~g]'ll{l1l~hedearth onel,lU~ed


scj,~'jJ1D'i!l'lt>i3l the

Afr~~ml·'Alnli;ric~l!il

natienselhe researeh hlSA,ilMt~.Qns. Sherley .• m a~~:,nci'Merl'(lfe.\SQ:r~ ~iOllld hruve bee'.~ll:hefirst ~'fmlred


Aincom

.m~

Aho. :('111 i[omi,1. A 5£cuml stimHu!:>~, nowa ~eniLJ[OO p:rOfI;lS:;Of Qf~he:!1I1i~'!fY ,,~ Wepli1:l!ii'L;lIIHlI;ld etiunic m~[lolril;.,l'l:;'" Despite wilim~ng a $1.:5 un .mOIl, dIe UnivcI'S ilY Urnli.l.lUois, Urumli:;1r C~11ll1liiIlaigl1!.believes that his fOln I~ Directors Pimneer Al;.\~<w:dfmmthe ure lO W~.11 'Ito]'!lUffiHl MTf w.,:... '[ n Nnlio[l'Oll I n.sUllUI(;~of H(:l.. ~tlt rO[ b,~~.cl ean.t~!~~y ~n'~\is~s.'iment01' (In ""{Hk o~ ex.r<uudiJ~g line~ OnlWill~~n his comtibutien teseieucc and ~hc adult STem ecJl~!'>. Sherley's roqUl:SI. ~)l1oEession,'''Jlldg:tuemls are confur ~emll ewas dented [Ill Jmul :If}1 r 21005. l\voimltt'lnnl: reviews did,l·. Purtllk ~~@wst.l<lJmli!s. 5!wnl~~b~irlis eadl nKuflil'lg Q!!~sidethe MIT IPfO~()~t':s ~~1IntlyQ~iJijg made whether you ill:C.m~im'C on'hcl'l'l er :I,lot •. imtl your dnall.brc' the dicci:s,Io.ll. Sherley's Ilisl. aiflilCebeloire rftumil1lg to hi,:,;lab lruhe aftemoo.ns. ~d osyncr <'l{:ilc:s:sta n(~ Olut much i l):f t:~nnrliliiuts~ltdll(k;$ the anl:.g<:l' lion tI~at he ,"",IS,given less, IfI~n;;um'Y space three-person eommluee to reexamine the :IllOl'e :i.fyoliJl <lrebiack t~um ufYO'lI are willul:e'." tenu~ review, '·B~ll It!'ere~w ..s ev¢ty "lmtem.j1t ~~~ys P'l1Iuri.r rhil~ip!'i .. ml Ar~ic.m A!1li1;::r~C;;ln [h;JIn, ol~er"1olte<lgtl!e1'i ,md.ll'il theJer<lirtl1l1~1I~ '\M~O spellU ] (J y:eiHS ~H M~fts c.hemisuy ella ir 8SJI;cdl~ is \\1 i:1:(;"-8. depa nmcnral colIflliHlI.:: 1.0 eond uet Ihe rev IC:W 'IN hh absolute Il~lflile:::>~nd imegril}~.'· a dep;mment. rlilLU~I.1~ ,'mys 'ilie doesn't kL~O'!IV ~e,~g!!e otlen ,1..~Il.edwith 5hedey-l:o who Sherley sa)!$. he hopes h~s protl;.\."'; will. hit weligJ~ illl (Jill tl~etenure decision, lie also "'whall't.<L~lJelted beh Illid closed doors' dllr.ing 'lItd bel~eve~~li t ~(llllH~(If thehostil il)" Ihlnill ~li~ 'thl;: SCh(){ll "I.jk{;) a Hucle;;lr ~:>i;I~I'l~~'lal" ~b!,i!;~ :Ili~ tenure n;:\'i·e'~'illl 1991. ;"'B~llI W~Ui told ... h to e:i\;afl., iI'I,I;;~ he quest non of ins~i~ut ione I If.k:r th<'ll. for. at ~cas:to~.,e faC'u,~.ty rn.u::rn:ber cuUI.:'agucs is jiLJck:d by his opposil::i.u.ilI to eacism, d1l)'5 boe:fQI'e he l:egml his strike. human el1l.11Iry<lnic S«:'111 ,C'eU :I,;e~·.nd, inv.ol"'ed ill the deeision, the fact that ~,wore 1\,'UT rrt}~ud~ru~ Susan ~I<lI;::kfie:M announeed lba£8.,v p:JI:nl:SWl.~'~1'1 issue," "This is ab()U~ ~mlin!:l i.1IiI:~ ilfili~st.tt~e thwt "a comp:reh1:n~i\j'!;" :~igQro\!~. ~y)\'teI1li!.",~ic ,in~ Sh~~ey.,\"~lo:.<;t.} .<tpr9imllN'~~ ~~d~~O JIi.II~le" ~r~~~w A~h~'1!hl\!pJ.:!!'ic<.!~~ie~~i~''S het'ng;e me Sh1(~y(,[ Ille imp~uct of r~LCt: ot~.th.~ hi'.bua.g. ],lOPC;'S~13t his POOh.:;:,,"i" bcg_lIn 5 F'~bru;iry, ,\,'iil~ ~ 1 h~w:c: e:[u:lllft~d hI S:i:lflfltlli:'"" SliyS, Sh~rley. who .,,dVllIliCemen1.1lIl(j ,e1>;,pe['iellceof m:i,:Ilo:rily h<ls\'O\:I.'eCl tod'rililk on ~y\"")~eiI' and lake vi !a~ fOl'l:e' Ihe lll1l:i\leI'SLt}' to c'Onfront stich alt1i.~nlde'S,~t;; . f;!.c:uh:y at time 11I1~tit"~~e_" Cani...;arel; ::;;;!y~ th~ :Shed~y s.,y.... e-'g ~n~1· iX)ull1d~ ~i.lfilce h 14 lJegan- ~, ~iiI~1iI~·~~!ii'cl ,~I~lro~y~~suppIeIi,'i,e!ll~, • 'WI;lre ' de~tif'oying C!lree:r!>.,.thrt'l\:v:ill1ig mll;.'nt aw,yy. Its 1llUJy ":ra:k~rg]~ith'8iilt9gC. ill 2111iml1li.cw~iY. of :I~:i.l1Ig.I~:iii:rcgitlffl.en of .gpt!nd~.migc:t!ry I1mol"l~~biig '!J!. : '[lilt: {JP~'io[llit'i~.lY fui' d i&.i.::u~:>h)!l " t:r~1J[Cd 'by "i.H:o:uJe 'tlttl p~Ji\foslt $. o:ffic~ lJd:[JJ\; i',~tllfnUlg ~.l(Jtlfigh:l•.it'll. rtnl ~iIlI1Hrt.,i!L~d I've ~k.'1.:j,tht:d~odo SherI!!:),·::; dra:lnalk step. "It llas: enel~i:red :;.o:I,nelhilll£, about it.. A dOl)!afler Sh;:r~ey " m hh lab :in the aftel'il:looll. He pmr:nises, to ;~ b~,UIl hi!'> ~1!1ll:e.~~ pl.'OF'e~~()n;,.ilnd'uding [i~l. p!.!!opre (l]l C1~np'~i~'to ~~Ik <lbol!J1~he h,Nue." ~ ()tH~I:illltl.e ·the mutilil~ .in.defiJ~~~el)i_'·Smil1L;lll,liDillil.~ S.h~:r.h.:y'sl'll:lngcr sl.r.ikc has ahm elicited :I,lrn>got to d.1RllgC 211. ~H:r:' hI.:' .!>:'I.ys. ~ ~tli:l>t Noam Chmu!!;ky e.i.l"Cuhlh::d 81 Icncr "';VUli!lI!]llII SHI~rrA(IH!AR:JEE ~, denmm.li~.~ a lrev~ew of the f:Wievance pliOCe'-:. .<i :support foom tv,,'() nmm:1' M n rese,<lJl\C:hers, dl(l W~llllr(:ld
f~IClldt}1 ~n:lIm]~r-

AIJI~ricm~ in ~Ile 1I1.~1N01'I. blol og lea ~engi m:CI:',lII' Ii: c:lcpartment 011 01. canlp~s wfuien~ 41)A" of

he replied: ... ·.An)',o!1!,e c,rn be ,I neurcseieutrsr. bUI~. tilib; be: Mrr ~" S"'!1Id~rr> now ~~<L~~l~i; ~hi:fcl ~mld\.::.f~I.!II(jlil.blicschud. i:1il !la:!'o

nl:e~

,I

RES EAR CIH IINTE G RI'TY

IBUlIbIbII"e Fusio:n Re':S'e,Bllrchler CI'e,B,lred oif: IMiscondlUl,ct Chalr'gS!S, but IDoubts Lingl'BIF'
Purdue Umli\'el'sity onkifll~ last \'..eek .J:(;;:{;iar'e;:d [h,U ,nnL:rIt~rn,11 inqLlli,ry hils cleared 'bt~bibk:--flISion-n rese<lr,hel:R~:oi~ Taiey,1!!rlkh,m of
~LI!~WdIOfiS of['I:~(;~,U(;h m,iM;UI1.rJll~t_ TI:'ei ~11i!~lHUIiI~e1llie'!n pp~.1.~ ~ to 1:Jr.iI'llgto am,eltd illllC'M:liy Y'CiIl:kl'filg r~w th<ll has ::l,p].~I~he l!ii~\il;nity, ~

Tiilh!y['lrkhal~'s work. say lh.m [1'Iey 1:00were never contacted by the panel 3:11<1 2I:r~ Irustr:'<l~ed i~s secrecy, by I~111.l[u; ~oll.UI(~nlill[ld LI~\'\IIrn,n ~l"lilflgon j is insu.<l.eT:lIlcyarkh~IIl·:;; .;aMel.op rlIS.i:o.i:l devices

:re~u;1in equO![.lIy com rover ...i.. L Last \l~H'ung" T<lky<mkh,Ul invited IWOse;par.w! teams to use his experimencal ~,et!il.p ,II Purdue to ~e'e ,\!.I~{;cher hc!)Icould sllol, th.~ relhale signs. of c i\ISiolQ, The experiment I!>designed to use ul.u:o'I:l>mmd 1:0 (.'!l~hJPse b~l.bble:::siJl], <Mil orglimii(;'
:s.oiVe;tll ,!IuD fo:f\.'lC' d~U~:e:rillm~tn!lilSill ~h~ bub-

bles ~ofuse. ~ibl..TIl~,U'Igeid~.cru-h.ilLiill and a pro'!On(IX- helimiil-3 andan eKI'I:<l!teutrol1i.p',It1..,;I~IS


()fl,'l:!IIt'~

B.mwidl :sc~eDl~:ifucc,iail!illS ~Illd ceumerelaims s,I,HI ~:j:wch,el,hlg around tltc conro n.ni.rrl:~i.y••. &i.e::: ~,ii:k: over ta'b:laol) bllbble' ruston
!lio ~igli,nfdyili,g down
~10\"'S

11fl M~,y.T~~: mriirng~.a rhysi::~at Ilt F

•.:~:fi;e:l vi ndica[ecl i1ml esenerated' ·T<!~e}l'rrkl'!!1!!lS<l,Y-:&' .. "~l): been ,'~ ,~a:e.!ISlll'ecooker ['Or <Ibimt a ye;Jir,"
T'lh~)'~I1'kh~L.n lhe, dlic;:f p~uJKlI1eliit is

Le'Ioumeau Uliliversity i~~ Long\'iew. Texas, nun the C:N.perim.lcIIU widl i.\.vo Hi:1Jjc.gr;,~dwu'l~ studems using t\,\i'j) du iler-enU (Yiles {lf~e~~ron cletoc'ton;_bol.~of \1i'hichhe !';'I)'S ~rJnI 5ig:nifli.e-JlI.iilt~)I igllcr levels of' neut ro:l~S ttianl did h c(l!1l~i!'ol!'. II!! Juae, W~]!li1Hl!iI BI!IJ;.i:,,"' physicis~ emerims <II t'h.e 'Ulli"'ClfSity of Tennessee, KI,'Im;:vulle, <'I~_<;(lI!1.'[~tl~~ expenuten1[ a!'ad S1iJS IlL;!
tee foun:di evidenee foreKce~ m:lilunUlS.

o:u sm:1Ofw;,ion, ihc eentroversial ide<1~lm ji(lund~l<lves coll<lJl..~ can 1b'LIbble~ .UI'I~~'way Ihllit y~i£'ikls c.rm:rgy thwl!I.gh nU!::lei;l!lll. s i~l!l!. Ih!1.Le, thr.reS('Mdn h~ds·ol.tt the prot~isc of.~~

Neillle~ atthese e.x'le!l'imemsll1.'lsiJeen.

pub-

clean. hn~ille!l.~ energy source,


T:r~tey;ll:I1I;~:l1~nl;j:lu!iJltlri.ed '~vidC:lm.x;

Ushc>Ll iJllf'lIIl~hm\ l3'I~f" St1ll1.1~.It}'Milnrn, :1ph,%j,., cic"'~ O,[k Riidge;. s.a}'~th~ i!I:!'1 'I.\'ri.~-ll.p(lf~he ,n Fol'n.lnger(~pc:'hm;nj[ Iill~~t he::;' SC(;I:I. d~JCI:rmldc ll:leUI:D'(~n cOllamers c.,~led S'Cimin:ilL~ion etectol~ d

o!.'l~yC(JH~~lo1:ld~lU lhr second one. Nmv,. w'IIlvcrs,iry o~'fid;aJs say the isquiries.have cleased Taley amkhlllnohnis· COlildut:l but thill d~Blih {)fUle:~uli.ding~.the
(!1fl<1~;;S.aUld {,:\~eIIlthe ~llf[l;JOtlP Ille commit" tees '\viHl,k Ib=Pl ccnfidenrial. III k~cp'~lig w:iJ:h the lII'l.live.I'Sitys polfides. "We're dOnfw.Llll it," ~y:~ rWIl'(Mu~;pDke:-;~lli(m j"e'.m:I!I~ NOJib~I·g, !

61") wmnd,,,_ .~I:I addition

Other expens b\)llrl Jl1Isitie and ourside ~hc ulliversi~}' s;~ytill!;: process 'W~ .shfOlLtied iJ~ so IIln'~h. se~recy"t~~a~ hllJ>ossible tiC! Itnov,! \\'1.1<11 i(s HU;l·ir\,f(lrk ,mcll'If~~'d to !'IJ\OPthe!"nftmi)l p~h-, 'th~ reviews e~!t,li ~d, 'hs '~um~geow'" ~f{m''I I· I 'hw\'>' wha~P~lrdL:lle is doing," 5,~y~ lcJi.(;;ri lishilng IFc.5ll.lts th~It contradicted hils OWlli. Tsoukalas, a Il\!cie<lf f'll.g~nJeecli,.n r~;n'dlle \Nh.o 'Other researchers cnminuedm dm ~ itcnge ir!l,~.'~i.~II(:':J S;U~l~ ~~ft!nJ~ !IU~g,Hi{l:rn~ ag~im;t "['lley.,!!rkh,m aa.~);(,::i~i'IIil'icgm>l.l'LlIlod& Utl11 yl;;~r, nQtl~m~a~ said tle spoke to~he otJ!!le erit !ot;-SmlQI;.n~m is~. !i;,~lln~~h s\! ~].~I;~ Tiil~()j'it~!kh,,~., iilitiil~ f'1,;;.'Vict:\\l' OQnilm~.ltm;: last spr~ll.g ~~u.d~v...~ ·of rhe Ulllivcrs~ i.y of Illinois. IJtbI.HU'I·· Champaign-evell 1.'OIi,se(] the issue ![l'W asked to :I:esu]bm ~t wri ~ten .. llegations :~I~ Se p l~ Ililber. wm icllhe smys h ~ did. Yel, p,ossliM.~ s.ci~ntufic f~~ludi~'i 1HlIe>-lh1Ii~IO a T)!lrnil'kal!ls S8)j~ meitht:r ilt: ~lm'!'iIil'yfHl';: !i!~;seh~ ~. Pu rnUCrl::."eillreh ofl'.lc h!l~, hm"~w"'~ct:~L~:i'I[c["ut:\wcllby l~.ll::il:cclind ~. ,.. ,p[[t'due ~ppohn~d.1!I ~on:"~ii[~c~t~lr:~\li~:\1 '''' the m.mt'er.,. a.llh.ollgh JUst v" hal ane~m la~b It parDel, S.uslic1k and S.el]~·PiLmerma:ll. <I p]~y:;k:i!$t Ang~~e~, ~, im~"es~i~,IlI;ld JlI.,l\lli!.e .... f been 1110!de de.'lr, l~ <lIllie UI!I~~rnity(Jf ellJ iIb:rui!l. IL{}:;; ~ ~c ~ Ju :Ilc. th..: ulfI.~ve ;;i;i[ y rcpo rtctl th~it li.t ~n.!Id 'whQllIcildctl 21. De fiC:n-n A.di\lm~ccd ReS{!'~lrch PWQjeCI$ A~en.cy-f~mdedeffm·ll:O L>e!)~ :~c,~te '~ (Omplelecl !!Jl initial ~fl{il~~fY" ,limli~i<ltlm:,l~ed <l
iii

ror~onof~Q sioll drew ,'I!;; ienti fi'i;; skcptieism from dl~ S~,Hf[(SciCflt"t'. 8 Maoc:ll .2002. po, i808) .• md I!."lriy:i,ndepemie::m effQns 1~~Iedwru.HlTol1uce l~i", results, hl:C~.ud ilng(!m~ in .2002. by his fo.rilue:r collc3gtlCS<'Ii Oak R ~d:ge Na[~CIr'lfll L flbau.LlioryillTerrmrlt:s:$'I;1t:::. l~:s1:ye'lr. ,m article iI~ Naw't: sioked !he COIitI'OVe:1fl'ii)' when ull·~;pm'~ ,h~lt sevenil nW Ta leyar.khan'$ co [leagues iU [:i'uin:l~Bc In WeST Lalll~~~e.. ]ncl~,m<'l. here he Imv.recl.f~II~·t~me w in. 2.004. had cQn.rplained thm he O!bSULlCU;:\.I

or

to l.{lH~r d.lt>! (,'OI~c~~\'lfI. iJor.l ~'l!YSho;llJli'k~ M 'to sec l.'VtdCIl~t': of'where any proposed 11CU.~ tl:'(JllS·Me ooming from. ]rdeUler,~t:lm :U.IJI:I.ls·are fU~~lfIg iii! tIlt: coHapsililM buCi!O!es. m:uln]llS ShCKIId hi~ ~flledeicerors .just a Few nanosecnnds ;'ft:lkf tine b!liibbbt~o1.l:f~pti~mn~giveofrabLlfSl of IfgJlt. IBUl[ Milom S'IYS there 'VIIS no effort to
11:<'1.(''k \ludlc{m:~I;:1tior!l1i..

PmtcrmiliUl .. Sm,J.ic1k. and their colleagues <Lnel1ill)~ to register such currelarioas, 'US,~ ng Ta h::yadtll.l<rn·s Wi UI ,el;lgil1;le<el<illg dia~[,~~!l~(i. toc-~m.~t'nJCl thei r :s;~mofu:->io!l 'IP~',~r.Iues, Ina papi:u .:I~.~y pul:lHsht:d .1~l:i>I. week 11.11 l?llYsf,r;:''''! ,R·~ rie~" ,l.€'f{ ill'S" the rese ,Mrc hers wp ~11't t'!!i:!lt 'they de:tc;:ct~dl n{l~~m['O:I:I~ 1100v,e b,l'C:kgfQm~d and ~<l\V no OO!IT~t"I'!~}f!R be~.\\¥;jen I~~t .r:J2i ~l~cs1~nd 11!1J;\11irOIilS, The ~ottom lime, Mi~oir'd says. is that mest :pe:np:le \~'OIii.l ~'iL:!:inm'l;' bub!bl,~ ~b:s;i:(]!ti~ ['"~~ LlL1li~ 'll.~e~\I'Ork.is, \le:ri 1%00 i[}LII..5~ciI.': Ii:'i!Iey.'Lukh~:I1l·s ~~b, "I ~hil'ik1i 1~IHyii1.de:::P~ilI.dl:::ffiI~:re:PJorfl[iC't~.(Jn {If illese results '~'OldQ.~lJIll:nl<'I. kit of ~~ooJ)lelb"Ot:llIld," < did
],l!l;! !><I}'~_ Until Ilmt

.I1.."I!ppeli!ll.lI!e chief fOirull I

(If

CJHJ:'li'gy

p:mdu~cdl by ~LJbblc fijs iOi~ j,s li.k~'lji W


""ROBEMIi SERWUIE

'L'le h:e;ued OIf'~\u'nenl'$,

w'WlN.Slcielf'loem <lg.olFg

SCIIEINCIE,

VOL 31 50 16 FIEBRUARY 2007

921

--

DATA SHARINIG

Ne,w',Swiss lnfluema IDatslb,lse Test Promises 10": Ac'c:es,s


The world wlll secnkaow "".'hctiler dozens of sC.~\::i!Ui ~:> and .Il,(.':3.~[hcxpens mCil:iU b usl$ ness wben they p h::~gec1l0 s.liIaI'e crucial [_~m41 (]I] biln~nlU I:i.l'l~]llt~ fi ,!go_ Beh i~I'I(~ scenes, th~ a satall gl.X)UI) 0 f experts has 3rr~mgcd1'0.1"3 bnmd-newintluenza d'~l<lbnjje:.. Ilol!~ed allhe S\.... lnstinrte 011' :BioinfoU'.ImHics (SrB) in iss Ge:;It'i;l!,'U, d'l,.t rhll'l~ toope .. L~e;:: fliLsr\"''(i~!kof M~reln If) turn t~le:tr promise in~Qrc8.1lity, If .. ~I g(le:i. accord ing ~{l pl.<In" researchers 'iu~d 1~]:5 I. "~l.n~~CI:~d llI.lIfU.t':;.\'iI' ill ~ set he N COLI
d.tl1JJb~lM':tovew'll. iJlrO!i:llH~tiun,~boUl Ihe

t!D

N,etherl'<'l:nds" As J !ligrlmmy 10 ~he Nttwre letter, Osterhaus says Jiles ",colluniIu::d il:'l p:rlw:.iple" rio deiK)sil~[lg ~I~f(lrrnmion in. the d:tlltnb;IS/2, "burl wa!1~to see 1:1ll~exact 't~u:r1IilJ; IVi!'SL" 'IN I.~.IOhas n01 been invd ved :i,l1l, p~:Pln dm but \\ielIDOllle." ~r, says D ..widl-Ie:y!ll!WJln. 1u.'i1!d of'the ageu(;)1 ~ fl u effbns, sms Dir,eclmAm,os Bairneh S~I)'1'i that

orS ..I'\ I to's d.f118b8S<C kicks. ofr, il witl ~~e mle,u with decades"\vontli of
\VhCIlI

nm~e a((01till; i'lave opened a .FIe!\\i a in~~ti en'E!t Ulic ooll:atmr~ti!)n ~tr.'ffllil'JfI E:ilIr~p~and lridr!<l,. bO~5le rl?d ~5t. W8ek at. iI liI1ee~,ifl illl f!4ef1.1 nell' ~ 9 between 1f1!(!Ha',sd@nl:@ mil['lis.terr and lni5i s ((DUnlerpam; frornl ~~l,e- IJ ropeen UiIlio'Jli(lU..I J E Tihe~irst sach g<llheli 119otHsidt> ElIrClpe, tile pa rley fea:liJr,ed India (omm i1til1lg to a S2 5,(1 !Irmil)r1 conlrib utien rOJ the $1.5 billion lFa('ilitytor Anti [lrnl(JJ1I antlll (In Re$ea[ch (FA!IR) at the GSI hea.vy-ton mS!itillrch ~ilbin Daam$!;3,dt (J.flrI1l18ny, Il'Idiaflsdelfllists .~~ ~Icolla borate elll the preje(1, wnldl onc:~ Clolllllltetefllll! 2014 wiU prQdUII;e bearns 'f!l'r re~~n;1lI flnlto 111,deiiilr

physIc s, pI8:>mc,s.,8ml ~nJil:leara 5lrclphysk5

.•

inflaenza

data

frml.l!ulnHJIlI~.,

bird:;" ;11lI.d

virus to tin: world. lmmedlately, with Ihc assurance Ihm !lIQOoo.yc1m use thei r dara to
rmdl!.lc€ papers, drug;>.,. or v'lt:ci.!1J~~ i~~.mm w tlte.ilr pCiT'nli ssien,

SlllIariin,g. feter IElogrier h~5 Hn\!! ncedGISAI O;s ~ff(lri;~ Ili lIl5e(,L

~I

Au OLH. l 5 11u ~.a cur rent I}' sh 1ufc k!l.:Y b~. genetic .d'lta about US; N I in ... pa55word~ pm~~~I.(ld coliiilpar.tri'iI;IH~~'1 [he hln~L'lll'i:Z{l Sequence .D3HIIb~.g(! { IS ID} 3,[ Lo::; A I,mm0::; N,Hvon;l~ LOl.b(l~:m(lry in Ne\\I Me'.x i·~\o,'f~a~
deal, !lIIml!eu the am:p:iiOOs of the V\lbrld I~~ealtin

IVVHO), ,t.~;!llleander r~[~ 1;J~~ >'C'~rfllU]u 1'C!;(!'J:1liChers, lerl by hllll~.2in1veteriO~a~liz<lliua

miry' \'~r"o~('I~isltarim ConPU:l:l. bel ie\'~ th.1~ ] who


e'IfC.fyOItL::-lfHJt jWls~ ~\ ool~cE.!!-ooup-slu:nd.{]

·~I,t.!s'1i@O~~O !J~'\!"t In d~~i;l1I1~O!1l rd.~ :.ay~ Jlol1Jn Wom;i, head Clf the U"K:~ Cel'ltral La!boera~,ofY gf Ih~ R~"2~rl;1lCt;JIullu;illiIFI additioll, India and Ilhe U!I. wi ~lfa en !!:(J)!1Itriti:luteS7.'5 I'llIiUion a nnui!Uy lo a i:tJlnl fe~~arc l'iflli n(lforprojW5 lln, heaUI'i.. dirti.alif, ilnd ii!iIlergy,.~1l.diall1!sde ntis;l;s \',IiII abo i:l~<1Ib1e ~Ci mmpete Ifeu gr.arlls und!l!'~ 'the E.U:s {·year, $7S billion 'Se'!l'>!lllil.hFrameWlJ:rk Prognmm'lil!. i.Milic!ilIJega'lli earlier h$ )'€'ar.~ll1ldiaMll be! ~htl mM~. importal1l and first IliHll!ltlr il1l !J1e Sev~,riith Wramel.iork PrCligiiai'nm~,u~ltI APlllliliU>e S!:ha,Wj n, Germany'!; 11l~ llIi)ler fQr adu!I;~HJ(1n rid r~5~ilr{.h, w~w led th~ E.U. dil!'~E!9ar~ ~iOIll, Indian ~s(i!2nu~rnin~Sler t\:api~ S,iba ~ ca ltea

Ii"!@a gr!!@meJl!t ~hi'S~(lri(;,"


,-PAlLAVA ElAGI~

h,we mt"C¢~~.Their cfi~ici~wi!l~P"lJ!k~d erethe


()fth~ maooll.nlEHmnvc 0[1 Sh~:riUlg.Avi:.ui'l l~nmmZ;)[);It.1 (GISAI D l. anncunced in <l letter .un Nature im.AlLgm;t:Wmi. d1<lt\'I/l:lS to;igm~. t7:~fhy more limn 70 e."'I}e!1:~,The ph~!.l\""'l~ :;;poe<~r~ ether ~pec ies, Bairoe h bclve\'e~ sw~ land's famed .n.eu[r~f1Jhy \~.'ill he! r ·wh.l over hC~lliJ"d bYi!llC dihL cn.n suhant Peter Bog ncr. umi~. ll~e,nunkaown :iu the influenzn \\.'ol:ld cOlll.lIltr:ues reluetant HI ccnrrfbure Wa (S:['icncc, lj AU:gILM 2:0116, p_ :1<026), lIS" -:o;;pom.cm:,;d dam lJII SI<::. hCf~d Carher ine Macken says A :mHI~l group Including Bogncr,. Capua, 1~ndl N<:1I11.ey Cox, hl'!fHI (d ~h researehcrs mily prefer GJSAID because irs free of charge-but she's. OI-UIY w.i,tn that. influenza di'visio.n 31. the U.S. Centers for 1];1~.M;l COIn ro I <and Pnwelni(!I] i[I, AI I ,m!;."l. "[~beh'J.d pn;:ss [SD' h;I~~(,.1e~ved ~~nl~e of GeoJ'gi.f1. has .1iI0W h~11I1Imercd 0.,,11 dcmUs of Ihc HSN I cmn,pAnmen'~ hm. becrn."~. Ilig~Hmo!I~:' she :mys,. ""If someone el:ie W,lnts il. th 1]l<m. iley reviewed fitlai arrange T Ihey 'n: \VI::~.come:" ".1'\ semi!::' of Ii:: iV:I;c elmy" :I,lu:n.ts .m: 3 :lJmHHing ~n Mtmh::h .~ast w,ee·k, A!Qyo~.le ~,m ge~ ac~e~s Hl the .~I;rIV UMOA- led BOM)lle:r l\_l l~,n~krti~l~hl:emi q:e {lpel .•lJi(lrl w~.th h h. owll, nnlninl:Y:, B~i~ cO\r·poli1!.t~ base. pmvli,dcd .~ he.y [eg~5tC[• I.og ~.Ili•. f~!iHj so .~fI~· . D<lckers. whidlhe ,Ieclined to !lam.e. are iQiCCepl an agr~'lememli~ruli'l.lg their nse (If ime:l1;::s.u::d ill II'Mlpillg ()~lt, .iiH~ $l!t~_ th~~I::,~.i';:L,. Bognm ~,~:\;, ~u5e "vhop,rtlviJiil T T~e !ss11e:(lf ifln~u~m::a ~h.;afi!Ti.gm~de d~I'<JJl~<ly'e (J mO!il~h~~o wke (",lire 1)PiHllents fiiC·.~ll hL:~dlitlC's .~~$twtl(;k \l!ihto::n .hldlon(;l:i>~.a ~:nd )Ociclu:ific Fl'ililblic~ltiHiiil5~ !I~1'l~ii' ih;;t.L th.Qi: <lI1lI'lOU~H:ed w{)lI.ld no ~ol1l:ge'r :slilare ,.'ira~. ilt i!'i i~r:on:mH iOIl 'wiU be e~ter,ed i~to~l~ree ~~~mr~~s {]t'H5Nl wilh \lilHO wi~huUD'1 ~lll~~Pll bl~t;d~L~:ab,I~A:s" :!>u-caUecl MiI~(!riilll TI'IIiJ~sferA~I:~elf1cl1t l'I1m "' BU.&lTlcr :ediJ~.cd to ~~r(}vidle d deja iii Ii: oFthe I~llilIit~ cm1lil.il~ef,ci~Ill'l>~ of l~~ vi rillS" BIl.t ~i;iCI~ ~ U~I.:Jt ~i.grt':t':~.nt1:m.whi.ch will bt: .ffill~dl:1:pll blit~ i f ~l'Idom:sia nollon,~er iili.iIi.res vi rllSfljl, ~. !ilhO.Llli:)i. 13mthese r~~ ~es,.---l1lUlldwh.el.ht:t' the'y f(l:~lm.~'];;d-\'\Iilm b!;l CnLcr:i~ I ~Q G~SAI D's ti(ICB.clg:I'IJ,~r ~'Iy,~he hm, !;luman.tee!> llil.u iil _ E:: OCi!:s, say:::..A Ibc:rt Ost:crh,alils tlfErasl.mJS Uniw~U !kc'ep' slui.lri.llg its USN I s.c'QMr.:nee d.iUa ,~ \"'~11l (lISMD. -MARIliN 'ENS!UnNIl'!. !.Ie ~.'ersity Medical Celltel' illl Rouer-cl<l!In. the
91.imn

KI:rlea Tar!Qllets bib Mlisth~,ef


SE~ Ull-irhewee~..set
iU'I\I

50 1I1.h Koman go!;'erllllnttlll]t

last

peDlaltie$ 'for 5ciel!lliffc nnis(iOi'I' d ucl. and ma I1dlated a !le,~ S.y5't~ITI fQr in'l,l'estir-

19ati

1191 iillege~

III ~Wi:n[!li(tt

illl s:talte-hlln.iJ.e(ii

~. sn

re

sciftnce,. IIlrafteil im reS:jjolli~e t~ tln~ Woo S;uk HWilIlg dQn~ll.g st".afldal. 1!hJ~HIles re~lIIir~ 9 onm In>!!! 1111.ahs. UIliversilie:s, ollila resei3trCil (~rnle r~ lhlllt r~eill'e $~t~ f ilIllId:sto Iigl'itefl c)'!I.er~i'!l tl! lO thwlllft5dentiric mi~C:Ol1d nlCl, ~ndudi~g'~I~;gli('!riS!"!1. dala t;[!IJlllpE!n!ng, al1!o ~IUi Ifiida'ti@1'I ®r \'!.It~ist~ebhJll,ie.r:s.Une gUidelIne!>, W:h~dll (onlilill rift\\1\\Iordlr I'rgl {In tr:~ini"!J,c;;IlI 011 An~lillllfioll~ to form ilFL\!'12'Stig a:tive~(onnnilu.eE!s, ![(!)lIiipr~sed cd ;1illll'.'!S'[liv>!l p~r:sClIIIS induding e~l1e.rl~:;rlnd()u(Si{!e!r"llllj)mbe alle!,Jalti()l'I5., Under the guid ~~il'les,penaillies. (Qr lIllis'((]Ildw:t [FI( lude Ule 'g:ClVt'rlrlment l21iLdl'l9 ~latei rlUll proillc t:l; and barrin.g il1l~~itution~ rrom reLif i,m ng $lGte ~undingfor Up' 103 y~iUS" In ·il. Lfte, a fllliinis.~ry oHi dall, 5aid th:<lt ll'te Q{l'II' er!ll'!Thlwt hlJfJ ~s ~hat 5.etti flI;Il up this ~Y51t!m \'\IiI! fClH:e sdeillli5~s~oMke mON! respollsibili ty in their reseanr:hl

NEWSOFTHEWEEK
ClI,AGINOSTII

cs

I~ ~hech:IO~·[C rJ~)~dly and expa~ding field of ''pm1i.(JJ]1u! iz~J ltiie:::dic.il'l~:' !.kl7..liflS, of comp.:,1'·

agency's overs:~gtlt of them, Tfuspol icy 'iwl'lJld ~OYC:l.r~.t~n'y c<'ll~J:~d,HI:l rests, in.dud-

p;H~en~$run simu 1~.m,e(.msl}l.,M."niinl,i'lrl·Unl


~15;e:!> 7('t"glt:!ri(l ~~g!.1:l1JtLl~i~ M:rul.ify women II 'to

nies

<Li~ntl)

crreate

test

Ihllt

read a disem;e's

ing .. pussibly, a method! :foJ' idelltiJyin:g rhe


primary S.Oi;i.fOC of mctastarie C'tL~.lGC:U aund !1 b~ooditest to dielenn i~u;: heth er tl t lr<Hl s" w ph'U~tedh,L,.':r~n bQt:~n!!;rejected. "Weliwd is

with

llil'e'<lS!

ealtceu:~hat

h<l'.Sl1'lsprt,""'iId (typ.icililly

biochcmi.c~!.~!C;a I~ilvcsm~d~mt iic~.p~tc rhe course uwill Hike. The pl'edrC~lom;. \\~lich n~le:lfllre:ly ml gtOltt: expressionparterns, can

one of thl.: gro~lP5idun nl.:ly ~vol.d ehemorherilp}') imto"low-risk" t)II"~ligh·~~k"categories,


{)!nct)~'pe DX re~i~~ ml <~.21~gen€: ,m!l~ysi~ 10 help inform cl:inicn<lns 'wheDhcr p<lllients V\1'ith loc'l~ ize(leMIf(lge!i!·"f!'iIX'lertor pt:ls,~t i\oe C'UlI,Ce:f .. <Ire at risk of'relapse. And ahhough CEO !R,~~n.dys{"'lllt MGI1;,'Wmll:l~iCll(;i,iida,the R~h\~}(l(i

~~,;t:;,~"hnuld. 00 dlt:velo.pt::d ,,~, "you ·h<!~.~ on d ~"HlWl·~dl ~1~vd{lp'ldlll,l,g,:·h~ "lbe:-; not SlOlUlld :>(;~(;iilC~? And ca f'l JO,I~ys iIhtnk the-y ~h("Ylj~.dbe rug,lllm.oo~i:S such. ·'We'r-.e 1101 inject ing anyeians reaHy~n:teUilnell~,le~e"'Ll~ ts'? Last week, I~ll':U.S.Foo~ and lIuiing in~n lite bmly.~' he !:\'!~fS" DI.'ug. Adminisi.r;Lt:i,on (FDA) BLH sOlne phys~dans see sigsteppud fumfo[c,~ into this nirUt;fmt~~ks.. "lf'I \'\I~.thhold life·· debate. issuing its first-ever savi ng therapy 1:I'O~l <J. patient Iil:ppm'll'~d uf <'I. liml!ll.~ligem~pn1gnllos~c;l~I~e Qfa bad: t~~t, lh ~~rl> ewt)' tie test snd .2 d~tys later I~Dldlin~ a b..it as bad <IS i r I ii<lve her a bad c()IlQen~iomi !'ilee~~n: to discuss drug," ~<lYS [)anLe~ [-I<.~yes,a .lHOre stril1i£le~n ,(JIve.lf:'lia:~lofth.b: b:lica sa e aueer specl <IIIist ,II: ~hc Ul.li\'er~il}i of Mi.chig.!;1n., <U"~'L Called M<lmm~Pri~lr,dl(;'l testapproved by FD.A WII.S AllIIn Arbor, willa hejps run an developed by Allilistenlam·:b<!!;'iIed Om:tll~~eD'X ~r~<~I. Age.ffild!i:i1.miJ.cl aims lOPIfe:(Iict Phys~cial1ls sueh as BayelO.• nle,un!.11~ile.,continue tn pl!llnle Ol. bre~$l c,~~l>Cer'~isk of recurr resce. Manlliill9 Pri~t ~:S Cllrrl;m Iy 'Q.\lcrwhl:l:t some progn{)slk I£S.~.S are r-ellUy telih ~g litem, In COllK'C'.i, "v~li~ahle in Europe. SOU!:l:l AI1iI,~ri'l:'U.liLlH] ~Is(lwh~[~;. Ag,l:lirll"it In,,,~y that. ,It 5 y~,lrs" 0]1", l1'1]: IHigh-risk gmur has a 20% relapse nne. d:i" ('l"Y~ i[ ImSi perlh~l.ned~lnre o thi2tUlI 000 •csis so r~""Ir. 5 Ntxt ·generath:m. CanU'~ ha~e long been e-valualetllJ'l' dillel:t olwei'Vat~nJ (lfi'J~(oi;. 11~!.: ther .h.ils a 7:5%n::.bpsc In. lh e U n ite d St mesa I'Id .umm; <I !iI~ lMt uses g~nee:l:pr!!'s~orrbO' j~d~edH!rfi5~of r·eWrlIellC!! Wght oowml'lk nne. but \\i~.m·!):b':Oi Ilf!. to .tla~]pen In iI) ye:;:t~r~'! re we me.\Mnrillg A Europe. di'i!,!:1I0~Ti(:'~ 1;;ud~us eoucern .. he says, as is a "lack of transMflmmflP,dn:t :JI'nd81tother breastcaacer pl'ogthe ehence ef'mctastases, or the liiuc?"flsks p~lreil1ty"t~bfJlLl how the 1,t:m;.fLropc:rli:Jiilliil{;d, 1'I(31S:uict:or (:fIUed OI'lOotypc:: [)X arc n.OI. !,.'CIlI£t· .; a lany N(l[t~Jlt. dCPtllY rinys,ie~~I.n.Jn·e!ah:::f fo, FDA is accept ilingcormnents lIInti, ~ 5 M 31'('11. tQUyapl)ro\'e{1 the 'way dfllgs illind devices trlrC. breast cancer pmgu<lUls at Men~OI.'ial S:liOlIl1r !HFDA't\ meering, Il~uy onmp,unl~e~ and 11m~:(,"'I)I.d Jirn.~hi;.': they\ecollii",ider~d t(].hm~'U.S, [I{~t~e![iln!ilC,II1"f,::er Center .if! New YOlk City_ sum e patiem advocacy gUOtJ1pS argued He aliso notes thm m~ ny of du: genes used lory lCSI.S and are allowed on abe ma.rrl1i.et against :FDA"s proposed policy. They sug\\Iuth hu~e seru liny o~' their science when by 1!I~ese tests are I i~ked~o tumor gnJ\\'Il.h, per:I:orl1lu:d under .:he auspices OJ",l1 sirng:l.e b'"t'stecill~.l11:r.equesl:hlglillore extellcSh:e c:J:ini" desphc enteIf'JlllIgcilat<'!, Ih3~ othel' genes. such c,ll nial H wm~lcl i~.npo~e afi !H~IlC":i:'ll. bwn·de~~ <1:-1 ll1o:-;e li1m,i fTeet inf],1 rl1m,lt! fI J:!" ,,,-,,olUld lab, A:\il~~dua ~'i{lu~btfDA ,Ipprov<l.l of (bilt dlifl.gJ1U:.ti,C:;out rh:. cantl n~r 811dlpesord Mflnmm.1Iir.r.~I.l:~ :i.ii1.IP~rl OfIJthl.:fLg1::IJCY 's ret~!Lllc5!:. a Isobccrli~t:£II. and ~~ paJ:ttQ reasst~JI'e phys.LC'i'lIIIlS li1Ie~e~ si M}' add", cl)-ecade:!; de,~<lY."We can'~ \\'<1 ilt that AL1other cOI'I~e rn. say s Haye s, is t hat dnm 1(1fI~:" S,L)'li Ch!l[~~~<:!\lwl!II. ~1.!ilt'il.l(;iti~i~t beciill'~M~L'iIl~~tly stwJies,w!:lirt': d~si.~:an'!:ld m i... sO[l]:l~l ay'"'the c,1mlIi!p~~ny I,;O-~O"~It~.t)r a1Jld ~ \ ;I~ dw U~iV~fSil)' ojf'N(I'Ii~h C~lr()h!il'~. C'limp~i "llll':\\~r ~ ~i!illplli! q~I!;lMio!il~clid the p<ll~ellll chi~fy,ciemi"~ .r~!f~cer.~~le D~r~a[!;ds~~~!I!l,.~ live 0, d~c';I-sJ[)me of lh ~ d 3.Hl cUfl.g.h~ be I~a~ ,wile!' i.~ trying to corrnlillil.en~i;-I~.il:e a largdy on. rc.-;illl~slrollIil.302 European breast b:re1list ClIIllCer P:L'1ogln.oslk lest ~ut did IHlt I:a:m:er ~t l.enl:s.FDA. e,"p:J:es:5ed oonl"idem::e' Il1I i:id'~:;ld1'1 g. A pafl: ici pitl"ll 'v,lh!ose I:unmOJr i Ulil Mmt)lfli;lPrlinl~ 'lIU~llit}" r{;l~pO!l'ids Ulfllu\mnyw~n ~C) Ch~lillOlht:I('lry. ..u~nd lhi1;riIl!,~~'ill!ilTwo of dle br~.s"t ~meer lCSl$ I'i.l!llare r~lI'- :i:'or C'xtHnplJt:. ~n8ybe Iwisdas~irI!.ld a:; olte :~. But now FDA \'.vi'l~h' Ito extend i.~ over~i hl l.oumuUiyil'!lm'~ :gltii~e:::· bl! :>cd 'tests ,.111 l!ncalaIOlllg. M;nM'i:!trr~ ri~lInd Ol1lt~nlype:: OX. g \v~.th .:;1 :po~:iJu\'l':pru!!!.nm.i,~ flun~dl!.:'.: (li.~!~t:t,]ti i sl]ch OJ case. 5:,1)':> Uayes. a. te~t res:u ~I.\,;',ouh:.I .~ h<lve s£:uled on a ~niddl.e ground: Jille' COiblillflSepte I.llD er. the <Ige'ncy ll.'ele",sed .~ <1mIt d oc!lies .r~ nl~rlket.~n!l the;i.r !ei'i~~ d~Jo!;luWS ~h~ m ~ln i"'~td~,tl .,.wi~li1Ii!.oldi~.lg fhe\l~r}l ,.ge~H that ~ 'U~l~lml s;u.t:ge~nIi1l:!illh.. ~ m,my I~togno~tic basis, !:If r0n'(J~Cl_u\o\c SHid:i.~ of :storod IjSStl~ 'leStS sJUluld be .'G:gUIHt~d as I'IH.:dic;lIl n~adc ~hc'ir prognnsis guoiCI:' ~ ..;JIENINI ER:(GUll N ~, F devices. which~'I.'Ol~~d \',m~.y expa~d file \,vili~e ~o~4erm clinic.:1lI~I:ia:1"nr·em1t C,<l!I1OOr ~

effect. ror example. prmnpti!l'jl: doctors ,ll]d patients to eschew cllenlolher.apy "whell ,I lest suggests lbl <I bre;,lst I,;"JI1i;,I~r ,~:;;,11m\, [fill uf recurring, h 13tU:8iS these Ia ugcll)l mueguiau::d I.C::::Hi :LHOII,-e l 'c~ inieal Wi:"(:, researchers

'have' a ntcmentaus

~J~"

abmH. this class Orp~odllctS. said Steven GUlmiH'I,. FDA'~ d~rectof of the Office of I~ Vlrm D hlg~ ostic [)evice<E'V,I~ ll~ at il~)n ,11Id S.. f~~}'"at :~!i!Nlwl;:~k's]filt)etin~ul1 G3 it hersbu if':;;, MflliY 11Iiild •. r\ OCLH"<tCy is onc

eeneerns'

. 'hy.

CII

n fO'I'~,l:ii<l,

(H1H1I1Ipm~y

Om;QtypeDX,

lhal. makes ~~ien:$, that ~he:-1e

are ;:l~kli.~.l!!: ome s

fll!llilH:iI.<:I.n'iIt:nUl.1

(I~ est ions: A

re t h.e-y

--

lin IE:Ulr,ope:s Mlailllb,8,g:' A. G'I'o:s:sy Att,8ck Din Evollut'io,n


l

$25 Mlimm~ Plrilre'(lr 'Ir'B 85f1ihl!lllSe 'Whiizz:us


and ~d~ellltlilrer flil:h~nd IBrrdin$.Ol11i~ ewi ilgl iMllilt 1lI~ !,jal~ ~'tJq~ Idirg;eSl· ~n !!!'I!'er!idem:e a 001~edml)log!Y p ri.m~ 1.0!iH'ltke dev.elopmE!IHnta SOI.uUOIllto glebal.warmirlg.
1M~-e'flllr!lpr'EIIlI2'lU

fi~odel0d .t.!br the


PAlft~S=it"s the most gQ:J:geous~ looking arrack on e\"ohut~.nrl ~~~TI in 3 I~Ql:lg imc: Thars ~,hcconsest su:; ;;;mwng EiI!f10pe,0!!1 scienrists

·\"iho~.~recent \:\''edk~,have received um;()~i(:uu~dlf[~\e'tlri~5.ofth~ if'/(ls O/ChJUI.1mr, Th.t 76:S-p<lgc. ~<lI,\Ii~h:ly pmtrL!I;.~. l;omc\ ..... writteu "'s
b.y ~~i,l[1.m Y1uhy~ 1.TI!!I,l'ki.sll ;l~lthm: .• \\;110 d~Jm~mr;!I;\sD~If\"'·~.nl~1Ui 41'~Ihe

~!1allledi to the £lev.!! IopFrh"ilt ota. rrelUsatd.e !l:re'Wed f(l(il;.ea iln 2004, h:1 :5-.2:5 nrriWon Virgin IE~rllhi (hilUll!n'!J~ Wi II VI! olWiI rd'lld ~D n,hDiil'if@rr (tin de'lfe~oil (! 00 mine rcla Hlj ""lab ~e le(lhrmloQY .rlllpllble of relll.ol)1iiil1!J ~.~ast 11billlen 11I1lS a 0'[ (arbon d io.:ttdf~ a maiorr gre.e ilhouse gas,. fmm the alir euery Y'!!~r.Curre!lt air-(C!llJItu rt~echn cqlJles.Ci05~ UUf!e to klL!r ti me s more t ha n ~he IiIl arket wi II b~~r. ~llnlll1'k it's. 9 re(il, ~ 5~~5

no milli(!)nPi.ns.1!ri'

X li'rillE!

source 'oWnl;'1.1~.}' cvlls •. in.chudll'lg 9/1 t lrs P~!i'IIsh er has sent huai
dreds ~fnm lhn~L~j'ldb; Ofb!lpi~~{]f

plll-~'st(i~t llrtin Hof'fe-rt of N~w York IJJni",'f~M ~ily,who flon~thelen w.arllle'd b~11 ~IQil'!~ it's te be a tau gtner nu l ~!Jcrack than b!llHdl illJ a bet~il'r :spa~eshlr- (umtlllt L'edllllll;l'log~ lo '9 ril b

thebnokiceeseaeehers U ai leasi, ~ ftlUf cO!li~1ifie:s in \Vcs'i(;m IEmupc. .l'Iiial:!i~ riIlaHililg" 5(h·(]oh in fr ~n~.e1l~'I,I(:rli!oei~~diill ~H~,k·... l!Jm~ Q A source of amusement to Illlat Pl!l!~:pOrt:5to d~S'p'rwe,evoUioon Sl:ielue. :>ojiii~"Itl!\> ~)ook has troubled ~U'id th£iII1Jii,oreth'liL~l 5(J% o~·bio:h.'ity iLcQchl!l.,. in seeoUirngeci mhers----.et;I)t"Ci'ally n Uii Frmt\,'e. w~.ere 01. Frc:m::h tmml<u:ion h.md'e~ in the~\l;,i~oo;i:J;:~. o!t<im"y tlclw.~.n~nr!l"are I1rit sme .'!bout the "Ollijd uly ofe~oilLtion." .:mys Kenee, Y1Ihy.):;: 'boob of Imlildredls of hiSh SCh.MI direcrors and h,l\~ al!;,oOeeu tl!',ll1L~h~~~din~o A:ro[)IC. Urdu. l~b:r'.,I~i;l[l~. i~ <~IiI~S;I)' ..dl: U!~U~[ edu"Th .i~ au ami mhc'lf'I,mgtlil~s. of Ihe I~hmt [1.': "",arid. Ci.l'~iUII' sysrem," slr,1r's, cvohuion,~ry biu~og.~:M ~.l{)\!;'B.AV can a!i{Hidmml.s di!ilrilmtion of Arllimnd de R ~c:qllesof ~il.n: o~ t~e de Fr1!,nae. C books, as \¥klll :!t. 3. pl~d'l{Jru uflDVOs :im~W{lb \\ij,l{~ \Jj,'UrlriI:lS lll:J~ thi~ booik ~nug]iu ~UlJch olf:::J !->i~~ in ~e','enlll .. ng.ll'lJ,l,e~, i~·lIll1;.~I,~\!i!!f. R'lImon; b'1U~e ·tWer the leaf..~h;in!il of e\'ll,hnin!1l ill 9btJUfMI-rUI" i~Sl9Ii1Cc!. abom Saad i or GuropL:. fum!h I:iluc~lioll Mn.liiluSl!cr Gillies de US. ~;~ckers--blllTmki!:~l makes i"iu[]il,g ~aw Roh.~en ~w~~Hywarned schocls W keepthe OUl [~c;: fi'l.c:ts very d:iffklll~. E~i:i~MIYS .. In .~un bmJik OLIl uif p~lr~ Is' .n,u.1:Lls, e-mailed :re~p(m~ 1:0 ql!es~·~o!il.~ spokes... a Haran YOl.hy.,. i~the pen~~OI!meof Adm~!l pcrsen f,~]frYa&lyadcd.lm:d loml'druss flaanees, Ok.u'lr~ the ht:ad o.hh,c .f:01l1ldl~llion.or Sclemi ~.c f Re'~e:udl(SA\!) ill. AnkJr~. \vh~.clll1.1LSpm· He addk:d dmt f:1.'<lrn.cecalli gmher'lll) ~.mdburn " :inO[:~(~ lamic:cre,Hion~sm I~ ~ill(:e l 997 lin the bnoks .•just Iike iathe G,iYS oft he NfllzL~..... yiJ~the eo:bpsc of Da:1:'wi.n ism em,~. (S ci ej~l:e. n: .May .2'00 L p. 12&6), Yinh~f3 is. cn,'t~i~edwli.lil:lhuedreds ofboo!k$: he is.·... mrYre 1 not be prevented by prHh~hi.IUO:iI1"'1lIItd b111l1iS"" Ilib: <I bmnd name ,. l'OtI: a !P;OlllP of writers he French scientlsts :lH:!iythey needtn Oper<lI:f ie~ds.. T;_uuerEdis.. ;;ITu~ki!SiJl-bnnl pi:JY~Gi~l (:;_Ir~funy ~'il) <IS ~'miLW intbune: ~:erit:.~m18w~lh _~,ly_~ France's slzable Mlls.L~mlm~llorhy. iljm {Q at Truman 51.m.c U~ivc[sity in Kirksville. M L'>ool(;l'i,\~,:~o11m.Mllicliedl :-.Imnic I;:L'e,atian~sUl.l. I:l;!!'>pam;eis needed. says de R.i,qle~.if only ~o am'll te·<lchel's. w~~bcrmm:emrgmnel1lts. Kem,;~ 'Vahya.<LOC>epts~h~t ~he "",,'nl!d is. 'b:iIIliom; of M~}'slille ,UlrJ {ltt!e~-:>howe ,:ried to prmno!:e IN{lyeOlI:.\l {lhltHlt .!'eije(:t5lhe[,.'(i!lC~Pt nr evoli!J!,ti{lnhe { ~:l)fChflfl.gC. More d~!ln 5~OO p:ag~s ~.uhc A (HCI.~- ~ud!rH1J,b~UI. ~ys. ht::lllC"¥!..'f m,ga~cs in diJI.:cl S oICmmimt Ollie nrsr~11 >l $erie',~ Qf:sevell \'Ql· debates ,:",itb !;l'e'OlLt ~Qn~ l$, l'I«'ame lh,]i1 \vmlld ~nh,rffilcl1l their Gn;:dlilb,.ulily_ U!rU~~.) ~,refintl~~W~.lh r~C~[Lres off~}~~ ...ls., i M!i;'<lfl.w~~!e,sOfil~e ~"d~r;)1 were !!)'ill~g to <l~:r:~:~~;,ied ~~.,vn~~~l:m-d'~' (lrgS~:j~.~l~~. ,~h~! E look 5'fI~~.I1,gly );1 n1111.3n'-p'Wtl.r "il':1:~~Y1~ ~~,~y,~, fi nd new u,.;~s fo:r Y;lhYfl 's.hoCl.k la~t \.~'Ck. ~; t~l'<l.te.... o~lI~~.ant';eory IS, f.1~se.1t '$ an ..<IbsLI rd ~y Eoo~oSi s't M icllael rI..sse H of hnperi al, Cd ~ . ~~~ L{l'liIdm'l ~Iys. he');; u~jil1g Hi~ .5-ki~:[j~rom ~ Hdll';Llklu_~ ~(l~I(:. ~'!YSG~n;he.!iIiJ~ lOri.g. ori;t! op~=> as;:_:,. Im'n.p ;s[3Ild. l-T.:i.scoUe<l'lglll.cPtlel.' ~. of five lbioJQ~i::>l;l> U ~rechl University in the <It K[IILglu, ::.'IiI'IolhltJ' ruei~j;i~11L:il1l.ysh.c llo:n:.1[~d ~: Nd&iJ±:[.lmtiJl1> ""'!lo ruc~irV!.lld~ m}py.
a

(O~. he aotes, is ~!{er'l' '!Nle~~w ..ifllteFl~i\o'e.


N

~RII:CI!lllAlI!mi . t<:E!!I.R: A

'Gn!!8!liingltlhpE!' Fo~est
Tlue SuniU~l'lr.an Tiropital. ~e,udu hu$lillut~ (s.ml~ Panama llil~r·8ceived ln amS!J lIlIillion !lJr'i!lilt rIF!)1I1 the Lo:ndOJlr-b",SOOl oonki ng gi:lml HSBCW e:<p.aM its Clenwry-long 5lltdies, 01 ra;n~ ~I)rr@sl:~ I:ldlN urnd@irs~iilfld eff~d5 ~f (Ii~I) lhf! male d'lange. Wortirrlg with liLalVardUlli'llersily. SlIRI's, Ce!'!ler l!)f TrlJ[Wi:c:a1 Fores~Sde'lll:e ~\m (Ofl(hiCI en r1IilnUal'(BllI5lJ5arross allnf!WoFk o~ ,il'Oswdy 11)k!1~11115 (CllImlrit'5, as 'i'I'eU <15 s<tmly

~he rarben cyd~ 1111~"~ Irop'r\(a~r(l~Sl'5. The gin is s:nRI's.l1igglt:s~;evef p rtiV:il,teall I~lionaul d I~t~il~~kle ~illll;P1J1liliill~<ieill:liri~qUII~;;tiol1~hal ~ single-site Istudie-sl ~an'ta<M~5." says cenier d~rli!d.Qf5~lI(1ri1D~es. -!1:MZA!ii~1I!i! p,~lf\!NIS!

A BIUilty

1011 ill

Kiilileir

lFilw mlli om (!J1ldJ the B~~l.andMe~ind'a GilI~e5 WQI,IIlII:IatiOIHI rt dlangli iilg ~. $1,.5 bi lHliJn ll;~rrDI ~!liT!DIP@S, that tl~ ph,wnaoollltl(cli in(ill'stry w:iU pmduce a 'I.!<I(CiFle ror tilE!' developilllg world aga~n~t SWE'"Pwm(fM~iNHll1m!Jfliatt. wilirth (iJllIIS.~ji,pn~um,(lnilg qn d i'D@lIlilr'lgili:s L~i:it\\·~@k. ... ~i1@ oo!'!Srtlf~hJI;n1ph!dg@ci to puu:has>e kllllF(! 'll'aC;(~iieSat ill 9luaJr,a l'ueNI prric:eom.lf Ihe p~otlu(~ ~~I)rove;n safe al'ld effe(li~e. Pnennno-

(~(:ci'llillfii!cli on5 kill as. nnaIlly as 1.6 mm~on peopie anil:Uati~~.mn5t o~ Ihem eh i~!1rren.~ Mow
((llt'l(l.a n~es ~n!J.'llh.ll! If they llIalio'e t~hrll)lIiIl.e

~. t..'e\;,~:I~II" n p!:t1mo~'rll!l. cre,ni't1!lis!n, ~<;;;lys~{l]O- f~ey~r~tt~d i~ ~ h was hi~Hle~r<~d,lh~~ !!pd r~~r~ g~st.l'lylktit K(~i~ee f Iht~MidcD.c Eas!.l'ce~~t~(!f1~ c ~l:lbl:L:."he Slli.ys. o ~A,RJINI EN SElliiNIK '~ Uni\ie~i~y~nAllk<Lm. One ro(,,'em~Lf~"ey:tbt~~K1

~WithiIl1l!'LII·key.

BAV has been "quite sue·

his c:o~)~l his, ecol0I:l:Y .~ml]p-"I hOI)!!: to

ogy and lhey bU1~.d<I plant. they C8 11 sell. the v,a\!I; hl@,~ say:s IR!;Ibert Eila~kor the lO~!ll:5 IHop k-ifiiS Il<loomllergS (hool 01 Pl!liJ~i( He.a~1 h in 1Ba.~limor~. M;wv~and. :Blald; says the an<!'nglli" m entca FI a l50 Il!etl lool against oth~r diseases
p~~'lJ~.IiP1gl.lX"'Ir 1Il(!~lom,-MAR'!'liNII~NSERIIN!i:

925

NEWSOFTHEWEEK

nil! llcah (:Il\r~fmil


h'lee-diL~g
t h~

Center for Biological and Envimllmen~<lJ [or increased b.,,~icl"e~eOl:n,;ll NalloH::dl f101 ~ogy. ~ayt; th IJ ~ ... l~ belie \'I,;:!> I.l ill. and mental il1!.lpl ieaspet:ld~~g Juan: money on basic research and nuns ofunan.lorocll nology, the Bush Adlni.tl istml~O" .I~[LSropo sed a $3~ rni H.iol1~;~~yca ;J. network of censers is tUmright loW}' to go, p r

calls

NOI!llo~ec:hrm logies a ~.lhe'\~'!()od~·m",. \~lj sm~ ~


~[Jt",n'l."'I.iOlna.~ Canterfor Sdl!ll:.'J.rl;ill\'Var;,lti.riIg.

mil.

I),C"

"N<ttum::clmology .is bei~lg commer-

m:!\\iOL"\k ef'academic centers to pursue-the topic anJ(J) di$~~ntiwlEt:: Ih{': ri[Jdil'lg~._ But that nctwerk. part 01" Presldear George
W. Bush's 2008 bu dge~ feoql~~esl :.;e:JHlast 'Week to Cungress . doesn't address W~l<Lt lW~l!ly consider 11hili!!ger p robleru tor E.~e

Ilc'ld: the lack of Ills.earl'h tied mere tightly


to the developnul:llt ofn'e\\i U.S. n::gllllOltf{Jn::o, tn2(Hl6. lh!.O fi::di::u<~1,ovt::mltllCliit Jnlllded g

SJ8~nmjon iu re!>e,-~ij'ch ~ J],mnEedlnlo~,ogy O envircarneneal health rm..Q. ~a:l:'c~y" &afl[ is T h kely to gro\'I.' to $46 m iUi(l\l~ In is: ye1!lf {CtJn·
gru:;.~wm .. tl:1';r~"C'('!ild in hike f~.ni7lllad ion
01.1

eializcd in II \ilcry fasl pace .fU.gRn. IH'JW. ou'w Y gQI, to position rue' ~i:eIll.Cle1!lbeml ()f~l[]jt" Ilmlmj,:.d~[J.ologyrem1~,i]rIS il] its l-mrily de\leh}Jl~ Rej~sk:i, 'JiI'glU~Stlli>:!.! US_ regu!:JIwry ~Lgt':nl·' ~ne:lJltd)esl)ite some 380 products CO:nlU~11Iitlig cles •.such as tl~e Ellviromnel1i~::11 Prol.C'Cliolfl ~l'lIl{l~ll<lterk!l"h.,! me <'!1I.·e/!Idy ~hl';l~l,]rJ.w:r, A~e!lCY (I::: rA), mild the: Nmi(lm,~ I ln s! inue For t on Ooc(lp'a~ion(ll Safety and :1 lealil'l,l, are stn~" AI, this st1l,b'e. she says .. ufs UI1JlI)OlULnt to learn gi~ll:g[0 keep upwith the (liil.~sl.uo.lU¥i.beillg more '1ihrJ~I~ how' J.l'lru]]~te.i-~ljt~ill~er;;1lC:twilHu rn ised aboml~!O'Iiv best to regl~1me na liH11teell b~o~"agk"~ sysu::m.s" Ue NSF ne~w{)irkpn:lJ)()o~~ dojt~~t Ih<;;1:~, .p;rod.~lctseH~el,'in:gthe market, And allll~Mlg~l re EPA ,"Ull,ld receive S9llllli.illion ,in 2:008, lip shl:: :-1UY:S. .inYCi:'illi.g:nilnglliow I!h~ Mrut:tun: of b,Y liunil] 2006, R~j ~sk i<I[,~U~~har t ~.wid~ .,~n.~y of diffbT~~t !l,~nmn~~edal~ ...ffrec~1\ ~hl~n $..3-7!I1iU~

"lrs g;1"e:'l!1 11CI~'S:' Co~ \.'~ns'~ys. S.I'a,e Imle~ lilat

theilr env lron m.el:ltil ~ be hay ior. "Form inga llelt\il.nklh!l1IJperm itseve1f)l'0'rle to exchange s~rpI

(;oJlsidembly.mo[(:

is needed,

the fedel7l~ b~,dge~ Ilil is week }. "n~id: pre sithe


d~l~l>s. ~ttdgc,t 'wtluM bOtl.'it ii.lu $59 ~nmiorn., A1 the' comersrowe of ~h'uslile\V pusil :i.S<I l1~~""'O:~ nf C~ll~,~rn, ['l,Ilila~d by the: N';'i[timl;!~
Scicnee Feundaeion a~d modeled .?iJwr

ies ~~i~' m.~tl:iodo~i:}gywi~~.l<:':ally f~H;t~ (\:mm1iW;:e c<'Ilied :I:~')r XI)l1lI"d~n~ the ~'e.'>emx:1l1 e rOIWdrdl th~sf~dc:lby" couple 0 fyears.;" she s,,1YS, ~€cd~d for n::gubiory!lgc;~cics to ensure i'l~1.i: But. IHH everyeuc ~Lgrcc:> i.hc~l baslc sare~y of mlnol1naterials ~Iilthe environment. ~:ese-<ll"ch is Ihe best i,Hvestl1ileuli rhe g(Werj,l~ Alld 5l!1rprlrlJt:;fS h:ave [L";,t1-ii[m fuel ~v~l~g~sI1~·' 1:0 i !niHtl: oo\uld be i.ll;ikilrigtol!llnidoerst<nld Ih~ Gn\·i·, tors wlll.heed tlna.~ I~~C'.J: every yca, sinee i'l~e .~11 11:i:JfH'iH:t:ltHl~lle.~ltl~ and Sl"I:Wcly a:;peN~ of

l;U.l Se[Jtem~ef. bmh Re~}tlb:lical"l and [,l'eL'lloc'r.ll:ic ~.t':~li~.eit:S)~' jill:: HO~I~t': Seience ~

~'.'(islin.t;NSF networks, 'Vicki Colvin. a CI~emi!>tnl~Uee Univer·" silyi~ H~)UMon. T~N:'us. who din:;Cls Ruces

~li!il.lnpndu:t~. N.m.oooctmolo~y '~ms !'wPPl!{j "eim,g a pure seienee project," sa.y:~ D<I".~d RejGs!ki. who diri;;'Ct.,; the rroj~ct pn El1lil~ll~~UIIg

U, S. N~l~iQmJl N<l~lO~ech:lnlli)~ylni t~"l ~''''e began in.:2 00 I, Congre ss hastopp ed ~ he


pH::~idt'ilJ~\;~qt!:e~,t_ -JlOBERT'If.~S£mtl~(E

1lI., ,. ACA.D EMil A S

Han.mrd

U!!:Ii'lo,(lfS.Uly1

search Jbr u I:I~W pres-

ence

,II

Radel ifftL:t:..

1.~mSI

s,~ys,iriJ

au !IUidt':

riam], Drew Faust, 59, wall become I.l.lt::irst f \'!'m~uu] to ~.e:!I.d the eldest am) ~1'C{/Ihhiesl.LmiVC[l;.ily ill the Uum;Lft::dISHllC£. She: e:mcl'g,ed iLS the lOp candidate in the yearlong search .<'I1'"l'e['J:i10m,ls 'Cecil. bim:helliilltlt and !"lIesi.

id:e:nl H) su(ceed~h~coml'mner:si<'!1 Llwre:nc~ SHll':l,llrnll:.1"S ended this \:\A.'lC:l1: w'.it~.dlc !Jpp(lim~ IilleJill or a R~dicli Ire dean and civl lwar ll:isli()l~

dent Howiloo Hughc:s MediG~11 .~nslhIHe, withdrew his name room cGn:il·i,Lier~tion: earlie r, th is :month, Alief he-r <'ieIeq;:l:in!ll wws ,m,n'llJ.!lC'ed Ol~ 1 ~FcbrLlary.FiL~st spoke Ol1:t ::;lroJ'~Y in favor Qflllle>vunilia~iive ro increase int'el1tlisclpl~mary '''''ilJk ill Hall'i:ulr{.E~.exl~'ITi~!'i~ SC~.b"iK"elPtngl<~m
~Sd(;'J~u.~(J J":~~~'lry, 449) andadded p,

or

"lp~:n;:~Jingin the nl{lMreC~~H iss~u~of the Rll(ld~:rfi.'glllU1'(!~'{r.".~l\V"JS dCflr tome .Wro:ru liI'e outset that science needed to be an !1lII.ponam emlllDllmlem for the new unstul.lU:e:'ll [~~SO runs ill Iilc fam iii)': Her hm.bmtd. C'h!lrks, R.m;,1,;IIi.OCIig.liS in histerian of nm.011~~ . ci nil: and science <II: 1'.~.rur\711ld1.. FelliiiOllli~1~IC:;lll~ym~lnben;· 1ilH,~rV;;lIrd SiliJ l'I"cy\vcrc delighted \'\:~.~h appt1JnUlilCfU of the iF3USI to M~"CO;:ed Summers, ,~~lt~CQlltrQ\'e:Jsi3'~ comme.lilts abeur wO.l.nen ':sabillity ~o 1;(1/1,';r.'~d in the ~'iell!oe:-;ctJlllirijhu~{;?d~()h i~ ~1;!:.'1ijgl1a· 1:lon J~1;:;:tFcblltlilry" "TIn ii> 1::;;1I1'lill!$p;LU700 choke.'" So"l}'S [Evelyn Harnmend senior v ice pl1ovoot tbJi
!,It:uhy d~w~~o-pn:U;:IiI.t ~i~'>i;'if'Sityanda hi."I)[:!)!~dl .-illl,l of scienee.whe has "'m.l,,\ql r~m;:~fQ[ yc,~fS,"&1,1e IJm~(IN[morcli:rI!l1)' k:atit:ftihip qi:i~~~ itlies, <Ind e'JOOrrno~s :i fI~egrjl!y." hfi a p~S'S c..'fmr;er~I1t·~ <!fl~r she '!I...IS Fl1I1li~(l '5 . m:8HS.~prfllsed SlII1.Uncr:;;, 9'1~eCO:l,lam:islmul
r!l,

~h~l

'~h'!J\~vmlJ~:>tt}blt:llk~k)\\,~tltllnc bHrrh:rs, ki:tWXn the' sCliences and the :Inun.mili,ies. She I,.kes
i[w~r ~J11l1l~}' Jmli.~ ilHen Iml prl.>litd('lll!rD~l\;!lk 30k.

Son~c s.egreh

co\llUt'iJilLiC'C' ~'llelt~'ber.;:Ii'~m.ecl \

I :~aI'"Vdll:d. "She Iis not a sc~elllisL b~t '~al1l ce,," l~.. t'i Lh~ Ilhu~ h~.1l'blj' 1!1r,d,djh~l!In~ ..'!<',rit}' 1lit.y':; explllliS10n of :i[ s ~d,cncc'elTorI5,."!m)'~
't::!ill S~(l

"

a ~jhysic~~! oi' biulo!!,ic~.1~'i~mi~l for Ih~ poo(S:t. 'btll fli'iemis and cone1l,~ue. . of F::L~lSI: . inti::ist~he

h'lS: 'I.g~)ocl.~ru(:krrl'(;ordl i~~"llp'p()rting ~~'lJc.~~ in, her eurtre.lilt j ub as d.en ~ of the if:b~de~n!T~ ~nMitme for Advalilct!d Slltdy ... s.choo~ wilhin

B!lrl:ia[a Grm'z. 11 oofn:piml~r sciCUI.lba arid Radcl~ ffe s:ciel"lCf a!e~,nw&n.('i h,<lS ,"~ork,ed 01Ifl~~yw~'h,~!u~t "Righ~. rj'{Hn the :start. I hadil'mmy pc:eilllh:: ~"Wto nne, 'Yml shcnlld l;lD UI) m~hav~ ng !>Ci\/;:;:

~~:I~~i~~ I :~::~:~l~~~:~~~c~:~~y~:~~l~k
,1n,!l ~lle .,dkJ~~l '·1 chiUllk; womeni1a\ll;l ~il~ a:Pli- '!! Im~.c ~Q du ~1'l'yi.hn.ng.ilil1d thai, .itll.cludcs bc:iLlli: ~ p:L'esidern't f' i-i nmrwcil." o ~NDRJEW lAWlER: ~,

NEWS OFTHE WEEK


lUI

b<ls!iclfesem'Ch.

The :~e'<ld,er!'l,hil}underam:! the dir(l:t:~or's

s[i.incli;, these

issues,

R.o~tdn'!~11 Olddn'-"'!:Ie.~ tlu~~e (:h'llI'~ng~s. On N.ASA'~ Ii!,ud:tet (]'un(:hi; "Large space science pn~gt:iH'JfI.S fructl [\','0 r!iJlil(hHnt:.IiIUL~ p.robl!ems. The first isthe ~l'IzLlJiljty to n1::iI.kcaccurate COSI, estimates

e,llrly inme hFe oFlhe program, Once th,e price starts togo up. 1.1'i(;a!j}L~IIK")' In l1::i~illc:r , 11",,,,
eancelthe PJIO.gmm.nr come lUP Wit&nlfllOl1e

~nOli!e'Y.And the ¥Imney is !''e~y hard to find; most ofl'he lime. yo1.l c,m'l do uwithout di:g~
~iflg unto other pmgnnhlls.

SeCOindll:l.le programs

.2I.II"C'

susralned fo:r

g.m'Ll 0 r ~L'CC[rai I] ~]I;Ifl'l b 1:.T~}na~mchesper )~e,~[. aad Lho:iie m iss:i:D'u~ z.;tI,\t~.in~. Ihr'1 ,ir(;l to:lfIgcif' period or~nl11c.C~rll~l~ny stmmethilfl!:: U.S. RESEAIRCH

:~on~e~Ill.uI C',,,,pe{,~ted, So~f yOt~ meet Y(Mlr

FUNDING

has

10

give, Do, you ikl'l!JI."

hQM'

many space

Sciience A,dvis"er S,ays 'That Pruniing [I[sth'e K'ev'lio al ,HS'llllh', [Budlg,el:


'flu: u.s, seieuee eommuni ty ~eeids r~gure OtU how to ~:i,\'e wil:iliun its mf~'I,l:;:. S:lyS John
~(l

si;:i~n~~ pl.ador.~b are m,:ti\'e;::ng,i'I~ tm~v"; Ymu m~gh,lgues~ one or .:w:o (Iozen. The ac HHlI.l number b 5S. which it->I !uL We ~:reSp~ltt!l ~:ng f a lot ef'money on space' science=-far ahead (~f'~v~rylmdy ~1,s;~ ~~ltile iI\lo{~d"Bul there

needs

1.0

he

!I better

<i.pl':rediuion o:fthe need


l1II'Iall~,,.,gelfilent~

M~lrb !:l.r~~r.

~i e nee ,,~dvis'i;l!:wrre~~dl;ln

Geurge \V. JBns.~.: ~n,partieulsr, ~.c believes


bicmedic u~seientists need rocurb Iheir fIPI']~'tite fbr fi..ldJC!:OI~ .fi.Uldlillgand s-pa('l;:sc lenti:;,,!>, ust~L;',n! to turn m O! n1!:us~im~ to!1l;!! b~ bli.t~ n,g and launehi ng::l new one, Illi Speaking last week to Scie-ru;(! abourthe pn,si~i:l.1ru'!.;;OCH! lJudg!.:Ot [""(IHeM to Conl gress, M.OlUh:nU;bll::lif lJ.:!!fend~d the Admi!i!~l"!!'OLc tien's plans mgivc a. sizable beast to afew agencies while :M~r\.'.~.llg Ilm,()l!;I: ofthe rest of lH1e sc;e~1~1~ ~M"llb~i~hmelfll (,V:h:'m.:e., {) Feocuar)'. p, 750}. ~:~esays, 111m. "1110 presi:dem l~flS, been ;'I $Imn~c[ MlppOI.'It':.r f scieuce a!~.dJ o toc.hllo:i.ogy ,IS <1. way W bem::fiil SOdiC~y:." (lind he [ej~ct~l1mTlph[i:m.I~'i fIT)!]il science inblJy i~~l\

orr

M<lrhu!L'$er w,~sschedl~~ed ~'O test iVy thi s week, bdol.'e Ihe I louse Commhtee on Scii;lllce '~Ii!.d"I~(;'t:l:!l{l:~{jg)'. kicku:og off iJ):lor;t :r~""r~t:lllg I.mdg:cl cycJ~c, Aad 9,~I]~oug;hIll!.: Dcmoenltil:: Co~,~re~s~v:~111 sW!rrelyrevise the presidenf5 request for science and ever)1 mlllJ"r !\e'~tm nf feder''!~ !Spe~Hli!l!M.M,lrlw!'ge(~ COImnCliU:S•. ,~d~.B;d for space. s,~,~dligh.l. en d~e uamre of the goverillmernl's S55 billion COInlll1 ilnllcl:Uill hnsiemtd ttl l~ppH,(>~ research.
~IIEFF1FJ:!E:¥ MER"J'IS

Ibw s~r<!legi(; rn:OInO!gemellt onl:l(l~ <Isseis.


(),~I

beUe:r prei,ed

.. WI;! need new fimlucilft.1 Inodds for addresl'il,g the tr~llisi no L~rrem 1.!(~la ~~(ln ~Imc
op{:nnuUlil of a major );ci'k:lfI'[ific fadlil.)' and for dellib~ n,,~-e~yIlhasing(']!u~ p[ogra1l11X fl~iH are expensive once they haveaccomW

plished q:heir mi~s~o~. We can't expect ~hel.n ~hem. w~ n~ed 10 C;OlHi~u .. lh' renew n~r ~l'Irra-s~noclllrc to keep up W.ilh w~~.1Iirs alJ~ h penin g iIIIt ~'Ieres't of the world.
lnd lJMIr}' faees a si IhlIi.lar Ch11~.t~ngt:, II \; a ru last ftl~"evtlr.liJlcl:cJ~U M.l
\!f~ 1;: 11Hll '

11lYnn:!

Qintlil@ N,~tiiol'!al Iln5!tillll!lii@5 ,of


'[INlltO budget:

1ifI!I!I!t Ltlil

"".1. think lh~y :have a S!mCILlffl~ pmblmn., and ~. dun'lt hrwc a (IU ick flliIswcr" , .• 1I.~~re h; Ilu w"~Y tllle federal gevermnenr can ever satis(y the
d~m'UH~s creared by Ehe dou~~liU!Moff~(ii Nil-I, bUd.gCl [bcn>'Ccli1 i99~ and ..:W03]. lt's

herd deeision ~oshut dow nl. an


~spmfiu;.blii& fl!UJ produenve

OPCfllllion

tl~m
~I'I

flu~di ili:vcst

dw budget il:eq~les~cllre.:'L~ell'l> heal~h t~C of the U.S ..:;,ciemi fic en~-e]:llri~. "1 tin ililk~,.he m1clr.<lIlWedera:l1 scleral I'~, ~!]terllJ!'ise i::-; weU-r~31~lel~" ad'~If'~r ~}':'i. M '":B!JII. there's been iI ramp-up of ex pensive pl'flgnuns in some areas, while ~mp,o;lrl,aml
l~il[

~ed liQ w h,m:[ call anua r-egt!.la~ed research marker, ,'with ooomm; rutdb~ls~s that arebeyo:nrl the O'Ib11 it)' of l~,r;: ~O\'e:~~ml~,m~ ~~)n!nn~. t,o [At Ih~ sml~,Cli me.] N~ H fllf~d in£, got Wt~y mil ofSlep' \vith n.mding for l~leIlh.y:il~cll~.scid<ls~1' m lh~ ~~ve~ of it:s ~~.~. I~!il!il are ~he rlly:;;ic~ul scicnucs". Am!. dlcru Illf'e imbfll:~~tH!~S withilll the' b~ome4tC:<I~~ese.arch eonlf',(,'pl.·ise ill \\1'hic:1:! it En.at. Gh:-:~ur ,.I,1~pnl,te:f!!.1 a f exr~li;lr,m~ (!:Luucs miltehe~ the im.1Il0i"l<ii,nt:c of ~~nc [Ii:: ~t:!~iIT:.h. A IOl .of iIS $28 bilHol'I bn'll.~cl i~ ailml~d'lI u'eatments <!ind thempic.!! fOIi spec'il1c di~e.u:;,es wther th~tl b,~R~C bi~l!Pg}' cell andmlol.c·cular biol.ogy .. \'0'11ch ral.scs [~'ne ~ questtol] of ~ heiher '!,Ie 're s~)el1rd e~~oll;h ... ing
.~!>.ri:b!ld1:~llllc;:h n

new facillries thm: \,dU be more pl'Oditictive. B~iIit h,~!i b(;': drme, .,. [,[xlk ;;nEil~ Hl)llbhj~ ruo [Space Telescope]. We pmbabJJy made I:~l,e wrong decision, ,Jeclld,e~ago. to haveit mainrained by lite sl~~ce s_huul'e. because it turned (lln.w be much IliIO!.·e;(;l;,;,poe:!~si\'~ ~hu!l 3~lyb~Jtly real ize ell. We ,could hove latum:'hcd many other s.cielltcepl~'1:tfonl1is forthe nrice
of

rr{)gnllti~

~n other ~.t(:l"S am u!ilc;]~!ffil:l'l~,~~ii_" ~J'II;:e.~. Bimt~-dk:<lI~st;l;ln:ll

."'l'~ifb~~IU~~:i.riI.~ ~m~H~thbh':l,
p~.mlr.~Hg C'f;limallCi>

He!lQ!~s l'l~tll~h~ AI~~r:rk;;[n Compil;l'~tij\'~Ii!il;l~ '~: .~lliti~Lt.ii",e"first rropo~-d 111S:~ ~ilr. y atl.c.inpi:fi; to

Ami ~h.e ~)r1:)hl'[!'m~~ '~~"Q~~~<i1tWi'!~~ife~ w


llr1t:

I:yd(:

lIm trin.~qlu~ 10

~ con~e't ~hat imoolallce j,n the physka~ sci"


.~: t:ln1;:e;::~ bj'bool'J.N1ill'i,gth(;l o'WJd.getx ofUi;e
.~. SC~CI11C~ Foumlinion N~I;{]'(I~I~

~NS.F,..Ihc f)lI!p'lrll~leJli.1 !! of EIl.~rgy l; ~ DOE'~) CUrlce of SCi~liC~, ~LIlI.di ; t~eoore labs:.n the Nmiama'il In,~liUlte ofStJ.lUlI" ,~: cll~l~~~ijnJ Tec~mi~~l!:.y.. he SOI)Il'~~I~~l h~r /\~d m ~ ng(-ndc!l C.\1111 thrive \.'o"ilhthcin!ll.rreJn b!lltlgc[s
'&1 !,J.

NASA. NS'F 1m::> e.xpe'rience w~bh.~his,.Wo. bul,~~ il:,;pmj:L:l~tt'i hecm'il'ie ~,,,I(ge-r "l1d~'ime c,mv:q.:.1iea lell ,. the Iinil n.agcm,.;:nl p ro,bl,ems
bmlOlTlc 11~:~.ditr, DDEmiLy do
It&i~

1J~~ljllh, It

h~,sph~ls;ed

OLn

<1,~hll1lf<ldl ities before it


10

b~lildt'i ne\1;.' nes, o \:lle arCAl1IIICri,cfl. We should! be able

by

,~eu~ll.g~()r~t1es lnd p O sliddlll~1,() .hem.

m,ma;e d~ese],)mjec 1$.

927

of IF,oi'cl<illtades" blll~ Ilog~$m wtHnkiin,y with (:IDltalfl tnMed or mils]1d'el~~h~I~1 te~111 R~ynold~oflhe Un~.ver~i.lj' Seutheru C3.HFomia 3:L~drbe C'hild.iJiCr:i,'s Hospital Los 111~nes: h,a:vew:as1:ellil ~ilme a IiIld milne, and [pllf,gllill!ll~!ed s,pulI1i IUS msuls,; Ange~<:$ '~I,u,t!UHe for Pediatric Clinical j,olllirnahl: :amI! ~undiing iJlgmil1ci,es,say III~Snlt.• !e'illijob tl so~ve tllrlIIS,pilloidem .R.~$~alrGin..~vlJlo '~~~l<1 Mu~h~5 !1~'\1pedi atric
IINI Hi!:.
19iUl:5, wllFl!:N1 IH~ 'WA,S PI

p.ostdt1Clnm~ feUowfu tbc Scripps, Rc;,<;eflreh. liilSIUlUI.l:UIl! San Diego, Cilti!hrunb, .R.i:J.i1Ihrarcl Koile'r rece ili'e>d\"I~l<lJW'd:S supposed to be <I
;1 {,~}IJ'lbnl·OlI~()r. '"\:\'e 'CiLlll~lOCd it we cloned genes iI.~iO it:' h,c fee,!!l s, the;ll''[ '>\1e11elllQ~YpOO [t and reel ized i~ g
hUI1l1,~1il C,lDUl,;~,r ~ :tim;lrmll eel
'I. 001l1O meuse," Afl"ei' S'COI'eS (If simil,!r experiem,;e1'io ,\lith m,~side'llldfied cells, Kofl,er ud: his 'CO~I~a~ue:!'i at rhe 'if}'ftdE),H! C,I~lce!" R~!\~'ln;;:h. r r~Si utme in .1 il!UIsbruek, AlJi>lrl fl. f~.o\'\1' anthenticate every line <J:S soon ,1IS itarrives .nthe il1!~tuti;llte" And r~I"UOdDt:.:!I.lIy ;1fi:e~w~H:dttll1lY usea simple, "he<\ll. ~l\!~~k nd reliable ... DNA fliJ~g\.:iipfi~uililg.Lc'dlniCJjw,: i.u verify Llii!H each ceU ~ine conI inues to be ~~ihalit 'should \~S

f ~.h~ Univc.rshy of 'CfJJ~ forI'!i<I, i Berkeley ..amI her colleagues, sho\,ved that less ~!h,ln! 50% nfre.~~~rt:h~[1>~:!ilLI1:arl}r ver:i, ry th e identi lies onhci Ii' eel IIi neg usi ng any B ucbri ng
('I

F:i.ll.lge3:priIlitIUlg. "Everybody is in denlal" abOlH the\'lrud¢~p!'I';l,ld pmblell~. ofcell line

or tlil!.':

cancer cell liae S and ·~1lc;S;~~·pol~!]~i.~. c'~!lc~r .· drugs on (l'x:i.~ling lines, I~deed.rnl<1ny MlLdie~have showa th,u a surpr] si 1'I!i!;I)r huge In mnDer of ceU lines 'have bC:COIl1C ccmamiuated. otten by
(liitie
r""lO rewcljCat1~:b 1ishcd c 1l11l,CC: rlJuf;

sHlflJard

lecl~b1Ii(~ucs £~lclil as DNA

cells. For example, accon~ing~{) iI 1999 ~ paper by R_flderick M,n,::li;;::m;lO'!lmd hu~, col- :t:l
leagliics at Ithe uerm01'lIll Celli Bank (D'SJ\!LZ) ill :~ DI:am1.~chweig. IR% of 2:32hes donered to !m da,eoall,k were ~.n:i,sid'mill.jjl·ied or con~alni- '0

cress eeneamleetlon, :mys Charles I?~urick

~l~l~ei(.L ~/!;umt of th~ p rob:~e1.n"'"hv,~s o Tile ,~ seems to eome ·flS a surprlsc for rH;,"Op~,C:' says John Masters of Ung.vel;sily Co:~~ege II;; Lmldol!!. P'ffis~dt;:!lllof~h~ lEutopea!1. Tissae lI:
Ct~~Wtr~ SQ~!et}'" Ill! And even I.h.tmgh bio~{lgi.$h' :r1i:~~.c:I!irrld
lI:

~.

be. ..~ ;J~'i<!b!>,nlul;eIIUm;tUilm",;"~1I)iR Kof1~r. fs U h;~lIih "ff\~pe<lled ~:t' neouarersprobl em,s e wli.dl cdlliillc Cl)tI't~i'I'iiilla.h'iliJ., 31~d \v~I.hmH Ihis eonstant \I~giln:ce. Kolllell'sa)"s. ... 1 wouldn't be cn!lI'ide!Jll ,,~bou~ll~r \\."0rt>":" {

No •.every bi.ologist i:sso wrury. A 2004 sllrv,ey oflle,uly 500 bioingU!'illi by GerH"llde

IEa~ly wa m1I!lg'. H!I!~a0011$ a,ve (~nlami naled S(.ore~ h of ~ifLl ii~QS lor more Ull11n ~ decad~.

hear about cress cmJtllln~:n<1tion.'·peopl'e '~ just think th~'t Ihi~ i!-lrio! !I rr{jb~}~miii, my :C' lab." Sili:YllReynoMll. if co~gm'lilinal.ed cell ~ 1.~fhHi arc usedill'i~rdy ~Iil ·"tm.t tubes" 'to :I!l ex~,res:tlprole:i,nM. a I~,b'~ work rnay :I~ol.be :~ :;! ~n~ c ~ed. B u C M ..st ers. and o~hl;ln';. g :rescwrrx:h with COlUtL~nilnlMcd lines eemin~

~'.Y

~.

9,2,8

NEWSFOCliS
,s;eaer,He ,I large <lm~m"H of artifacts in the'

se ient ifi C ~il~r~~hl.I'U:, Trouhled by til is oU1l;lo~.n.gpr('lble!lll.


cell biQ~og~s:~and protessor emerirus .. the Cu~ohl:' Vniversi~y u of A~nc:riu irlWl!l;shi'iilg~Ollio D~C.. has Taken iI, lipan hli.b1I.lsetrw beeeme IIle Paul Revere
11

experiment 10 be :$30. "lr's ~o cheap. so cbvinus, sa IrivuaJ. andl yl.:'t ir's nol D~in.g (june:" ti<!y~ M,lsten~,
I!J rliQ:ri !'l!J1 I1Im~~g~y The :roms ofth" 'llmHii~nim~lioIlprolill~nnlgo baektn the begi.nning. studies widl cell lincs. BCl\l!;'CC~.l tlln~ I1l11id~l%[Js end ill(: ca[~y

m.U[!!.:

C ;,1ifn rn ia, Be rkeley ,Ht Oak land found ll'n~111i 40 diH~r!t:nl eull liiliicsr-hcnh

R.(J.I!I~.d! :!Irdonc. N

h~Hm.uli!,und ~LIlIU!l11~Ll--crm':>-C!l!~Wfrlii!1'lu~dby the Heb. ~ lh.c f:i;rsl~Lmm2iJ] ce iI Hne m he ine,

grown. sl~cces~!FlqUy in a iabo'nHory,. By the


Ujin~ he IJubli&Hmd .i1Ii~ .W.iUldiiliig:o.l~.lgrtii are w alre ... y hundreds or papers dlescU'li,bi:~lg d

of cell ecmaminarion. ln a recent whhc: or the Cell p'iilper ehasrisi ng Ihe sci enti fi<r; cmll:lllmllli~y. 19S0s.\1Va her Nelson-Rees C lilt ~l.re labor alm:y 0 r ~I!le:U nU \'er~ity 0 r N1UIllnI1H::: calls for stricter polic~.Ilg .of cell idem iries. He argues thar [ourualsand nmdin~ ~g~nde5 such as the U.S"Na~iomd lnsuunes orl]eah~ (Nil-I) silould lIi.l"'I~date ~l~il:h'iilll~i~<L~io.!U 0 r~1eHI~ Il:~!';. Se-'I"crnll)l''() n~ssinn.1li1 gmupsr-,hl,c:i.!lJdi 11i!fl the Society fm: ~:nVil:roll~olojgY. the Em:opean T~!ii~lII'l' ulture S,odcty •.and the A liI.ler· C iean SUd~l}' for· eU.:O iol.ogy-I.H1ve endorsed 't ilIe will he paper, ~IS h.-rvl'! se'Yc:11I ~ cellrepeeuosies. Bm j<mmds and NH-I m-e 'il1r,IrY {If l~lkil1g en therole nW CI:lU c;~or.mnd Reynolds is skepl:i:c<l1 that Nardone wlllsuecced .vhl.:re he an.d otl'a(.lr.:; ~'nfWl: r~lncJ. "No <llTlim.ml of'passioued discussion by myse:ll:' or Dr, N~'rduliie will fix wh!ll b!I~'been aad cerulnues to be a wideS:l~read problem," he s~'I:Y''''' Mere!)! suggesting \!iI~.mneeds [0 be done. Reynold~ adds, '01,:,. a long way 1~:Olu

or

research uslag the (:OIn.I~.iIii.li:~]metl lines Nel sea- Ree s made ~l lus persc 11i.~~ liilission
IOW;IHl

others

~Lb{]llill tlTIH~an,gl:litS of d

EWI111ne

belllr~ck of pr~senH:lay

C(!l'I{ef

~E!seillr{jh, Ule<NC:~-'(i(1ip.1nel-;al

g~o!1lp

cd 60

([fIKoe!r

cllll

rH~np~.~ !lcw.tUy doing it:' Kol1i1;;1r ~ili1;;!'Ohis rceordas a eautionsry 131.c: "\\le InhliV'i.!' Slalfled doi Ilig this [n:~l;t~~OlIfb' Y~:ngel' rni n; r1:i Ii:n.Cl,;]llfll}, 5, 6. or ]yc:ar~ ago. BlJfur~th:Jt. t
0\\1'.1,'

lius nu!,illlta'il'!@d by the U.S. Natioli!al (anm ImlJlut@ (t'lJCI)ol!l'ui IJSili!,d wid@~y '~or Iboth bal~k Fe!;e-anih <lnd dru.g dnscQ1;'ery-lla5 llio:tes~-1lpl2t1'liTwe ~Cllllrllle of crroSiS tcml.1minalioFl. lin lllie l.at.e
ll99 Os, .Mord@(IiIC!liLi5(OV~tdl of Ihe W.1!!~?:II1!~11I IrtstilUJle QrSden(€ i iii R!~h[)V.mI:..I'srae l, h~~ !\!ibla i fI1~d n from th'll! il~tilllie the in:reaM Gllim1r ~lIle NlCF- 7 ami itg drug· res ist<l:i1Ilda u.glh~!H lillie, mm~ hooF! as MCF-7JM~R: (for Adl'i'cilmycin rEisi~t~I1iI:@J.-bl!Jth Il~rt o:~.~he NClI-6'Q paln@!.A l@w years iI~@. a c.ompi3lri$ol1l off ~111~ ilil ~is lab r~"'E',alell!l certa!iFll lJiIOll:hefl1i(~11iHe:rehces lha~ illU$'Irate<l ttoOw lillle>s ~ ~,ilIl~'H (~ll!s IOOUl!1!l ~ resisl(lllrlt tQ drug~, il'lllfH! years, ollf«lrk liitll n1!'I;!s€ lill,es had !.II ddtd 001 "a ni(@ r "Imy, "sar~S Llmwltch. E<trlll'in 2001 ~e ~l!IiD!TIi tte~ ~ maru:)S(r;~1 (!!lJ lh~ IfflJJ k nco OnroJ)Meiilnd ~~ aWaJilimgi,~ 'pLlblica!io:IlI, ilhel'l, olfl!1of iii i;~ ~lu;del'!~Sli!JlIHtI~.etll!J"OIil ,8 ;2000~~ll.e~ S ifltlil'~1surfJtll f1/,flf! NtN,#:mfJl (fJ'Ii\cer In.lll:i1me. sayi I'Ig tllat. D!'IJ.i!l.ftflilJiflrPfill1!il'Lghad nlwaletl ~ltatM!u·l a:n.d M(J~ 71MrR w~re in [all 'U f1iI\!l' tailed; Lismvi~(h aiM' 111i~liE!am imm@'{liately r~~iz.ec! tlnalthe1 r i l'I~erpret<llrians ~I!Imille upoomil'l9 ~ap Il!r wer'e' fI (l ~ooger villijo. l[)isapliKl'llIIted.H tbe y.ears. or \\'~ted time <11110 !!'!forl, tiley winh d~eI'!Itile Jllall.er t.frore It. w~IiI!.lo ~,rililt~I t !Nils a hig 11.1.0'1'1' III S/ U~(OI.!iU:hsa:rs. tor N@l >Ofll~ was M£f-7JMrR !,Jlllrreilal.edUI MCF7, Illu~ it ah® lurned out t@ be fderrnkalro iilll! ovarial1l cancer (.e~~ n~ a 1~(fI th~ ir'ICI·60 p,mellrhal's not the o.nly case ®r In iISlakell~tl1ellliry \ljmli IIIIhe Iii ~fi NOI·61iJpaneL Tihe 5iNB19 allld tJ2:5 :n.lines. olllce lho!J!!IhllO m.edijslimn (el'll~<llllie.("lffiljS ~}'stelfl il'i flieS, ~re id~IlU]G~1tl) t:(ldl (lith~F .~IIU;I!I;~lmt: from the ~~m~llfld''IIldl;l(ll, AIl~ M[).A-1IltB"A3~5i:I p!'t\l'"liIl:l1il ; i'1Tl!);cMfor mEtlOl!i.tClI.i! breas:l cam:ff. to; tI1emicallA) tine pi1lllie:I'lllillelil.n,eJll1lill.lin.e. M14.1N(1 has tfiioo t~ lr~ce~h!l~ Ihist'liIiIY Oir MO'A-ll!1 s-ij3 5, Mihiil;h \\Ia~O!ri'lllilii;'llly eS!liIb~islled in 1976 at. M. D. 11111:1e'(W!'l 'Cang;er Ce:rIIlitiirin HOUIMH'I. Tt>K0i5. INC] rOlJlnd I.hat the INI(;II-·15,1}il8Inel'~ ve'flllolli i~~he ~iilml1a'5, til Sii!lllIP'!~ tlf th~ line Qriig inaillU~ dej;llilsitecl at a !;~~llbanlltJy M. D. Alildemlli! and as, ill sample givelll tlil an N'(,llrese<l relner ~ tine (af1.Cfr {e nter. "JUt!! rnl>;-liIiD 'l'ji'th l1I'IeLa F10i'llil (ell. lillie 11'114 lile~)j happeliled e~dy in the lrii'StQ,ry \If Ute ~etl ti!1e1," r.JCI ~y~ o:n it:1.Web 'Sil~, t!lloouy1h llii~ NCI"bO P~I~~'S Web sile IiIDWdetail$ til~ hi~wry l:Jehilncl it's wl1ln~,Il<i'ra:d!Bired"cell li f)e~~li!if;!1 Z~'~I~!t~r ~merQf th tn~@rm~litl;n te\IlIl{lI~gy bfi!!'l!;h ol!t ~(I'~ ~~lQIiI!llle!'l~ l Th~rapJ~uli.t5 j\I~Cliljlraifll, cllf1~iliit'rs;~hat ~e~ri~tim) ffil®re ac_c:tuat~Ulan c.ontamina.terlen m;",id~l1ltiftf~tile illi~lillfJte hi! m' t 1!i@11I'il! oul af its "'ay to in:ferm r~~r(,hm who obtai [)eel th~se lines illi the ilIasllhal the lin~-~ are fIi)W sUfS!iI~C;L ~h~Te'".rI;;:$~y5 the ageney is wary ()I~'Jt;.~tililg l!mdW~ (om~~rn,. be(gu,$.e Z mudh l)il th~ 1i'>1Or~ Mlh SIMh liM~,~m:h as iI:IrUlgi te~lil19, j~ ullllikely tel hav,~il9!lH ('jom~r®mised. U~(lIlI,iitc!1 reel~ Ulia~ glFe.alier eil:piUU~Dr lll1£ prln.Me'ifI i~lilIefd'.elll,. He pu~M,~ilIlne stor~ ®1 MH -]fAd rlt flOW ~1l~W!li!S fI/J(I·J\D FtlRES,i 1'1 the 8 jlal'll!ary [~iIl\,erl6'l'lm. Then! may be lillOI'll ~IIKh ~lori~ Tn the l!.IIlllll'@.Th@1"@ S@n1!!! e",ide!lll;.'I;ll~l the N(I-60 pilfI1el's VllrS!Qn (Iii til!!! (()1.o1f!(.;Ir'lc@rcell liS li i)e tlICl-] '5 is In~llhlll! ~1'II1@d'" tine j):ri£limallil'le_, -411:.(,

eveu w~

wt'!'e

la7.}'_"

!Mlglr~lhyt:s t3!W
~.~~,t]iW cell 1~]l~S do assume SiBC[\ll idmuilul]~", .md\~l}' doe ... ill~O!rpe~ ~() (lften'! "~I~~ like Murp.hy's law," says KQfkr. "Ev~rytln.~ng that call go wrong w.iIU go IA'I.'()lIig. h's.jt!l!i;~. . ~

ntarter nfl'ulilu:,," Ahlumgb nm,m~re_"e[Un;::hen;,

200 I, M.';~t(;ln'\.~i~O hOl!'ibeel'l.~dvm'::;M~!~g \

are aware ofthe possi.b:Uhy of>cont~m~.I~.a~i()IU. amllCfJmitilill:S w:hen hmllCllall.g ed!:-;,. ".GC:l:· dents happen, CeH lines gel unis~l.aibeled OJ iCOIU,J[J~ inared ,,\,.U'l111Ib.~t'-:gV(lwungcei IS lh,u can ln uno time til!.:ie over [~,e orii!:lJiIMI~lines. The o(:lly way lO lu:e'\'e!ill cross coltl<l~niiii <l.t km ~sro s pm, it be ~Ol"e ,ill, Sj) rr::lI·ds.. I n

j"o r iI1iC N:;;l.sod, flWlilrellC'$S of the p robk:fI~ l'Or decades, I}ubh*>hecl (II des.cll'i·~)i!iQn ohDNA fi ng>l;l~rri~n Hl1:.:hni:q1;J,e l~mt hlil~ ~et~Qli]liB iIlJg tll~ 5.l~1nd.w~too~ fo~ .~.mhe!'l~!co;l,t~ng 'cell lillcs.Wh'!.:~.l Ii .~inc is CM3blii>:ll~{l .it b; cn~" tI da.l to reconl th.e dOllor's sene~ic;pror:i ~e ~ .and lh~tii >dIo t h~ ~~lliiI,e ftJrtllle nt!\'V~ !'L:l. ~liIy~' 1.!' ~ M i.ls!,er.:;. ~:flhi:> i~ done ;[lil\tl,>[ ~,e filr.i,gcl: p:r.i.ms :1iiilii'i 1l(1~1II\1~1 ihib~cP[Lbi~.d:f. lit \\'olL.M pftlvidc ~ ~H:hel: ~cieb1I,:i~l~ wi.t h"n:.m~~ l1:tIe s.ugrii'lIme: ..i ·t~~verify tile !dellillty 01 the Inu;l.... ReY!lo.~d~ _ ~ a,l1d h i1> cone~,g~iC'"" [c!Cc ntl y csti m.f~t(-d thc' ~ co:::.1for a single D NAril~~er!1'rinli:ll!;l

NEWSIFOCUS
U el, aconra m ~Illat ion. B ur the scient ifi c cumm IlIIlIlib}' mos~.~}' l~.d:""" ith hestil i~y. rcae and Nehmr.-R.ee:;: r;l'~'!1;'i!l1~u.. I~.y g<.!Ive lUP (,,,~~, sidebar, below). Noone was willlnllg to\",'uth, <lmw their papers orlose tlme~~' cl'edibihty~
and most researehC:l[f';.centmuerl moing rhe eenrarnlnated lines. Nar(Jo:m:, Reyno.lds.

:) 5-0 111 mphema-le


frnm

ukemial in es co II eel ed researchers around l.lil.c '!.'lror~d and fhul1l~~ 15(itB of ~:h~:!ln be ~o!llu:unin.l!e~U, '1.0

filed .. Rer1osit"Rie!i. such OI,~ ~heGenm!~ Cell B'::lrak and Iittl Am~[~Ca~l Type: Cultsre Collecnon (ATCC') proWe ever1i h~'e ~.r!.the:ir

Kofler.
l1~C,IlUM::

!HiI.di

dl<l~ h~stor)l'

other researehers arc worried is be ~illS rep e.. ed, espec i:III Y u
of !il."""" ceil Uru:;~ has

mosdy Wilhr9S.~cr-grow.ul:Lg" wei ~.e5u~b.~~:shciJ labs, Btu m,OSI I]C\~I lines are csmlblns.~~d ii,n cell I.ines,~n aleuer ~Ilthe B Febnmry 2'006 ind~vid~.d labs and from thereon are freely :i~SU!; of NtUlW{!. th,llY (ll'>timal~dtll:tl[.29% .of (;xc!'il.,mged ~~lW""C[l labs, n~w.ly .m..~~viflg al! :hn1n,i11I1-tultlor cell line submlssiens to ~he'~.rdent i~ checked . "The se cell lines l ies t~.e ns M Z ineludc cross CO.I!U fuui11 IOItions, iIl,Cvc'[ PM!!; t~lI'D ~.gh OUt doers, SCI t]~cy arc Bec<HJts,eorth,e smnll sample sizes, these aever subject to aeeerase aathenticutien,"
nglillr~s
11UC;:.

't!iJ,~ number

m best.

";1

su.~n~fi(':1L'I:n nderu

'pro~i fereted dnn~H iC<1Hy,


~!12003.,

M<lcLood ",,,d Hans D!'exler of

DSMZ .uHi f~eir colleague ..Yeshincbu


Mmsuo. then at Fujisak] Cd] Cen~e:r in O.k<Lymlu. J<lpfln. ~~lccked~bc idcll.Iity of

e sri mate." $."tJffi Mac lead. E5l~!ill.,ti!1glhe n~mi extent of the proiJ<le.~nn So di Uictlllt: lhe:L"Cllre far too Irll1lny cell iiru;:s b~~n1.~t::M,lbl i,~h~~i Y~1~r.'UI'ldi '!,I(;l ry every , few 0 f I.ht': rn eve r gc •. fl~eiJr klcnt iL iI'!S p:I."O-

S1iys MaC'LC:l{ld. He ami ~Iis coll~aglle:,:.;h;n'~ found that!lbmJlI. 90% (I f sci enf 5[5 ig;I~[lre o I" we luse 0.1 l11l11 b,lrlk~s re(llIIes:t to !Ie!no in !lie,\!

lim:s. and Ma.c:leod a'I'g.~es, I~mt depositing


I~n~t>should'~ required Sf) Lli'I!ll DNA fli!U~~rprul;us can be esrabllshedaad stored for

l~nes rrom 45 dln~r'~lIIlll~'b~H atl)ri~:;._ M~ln~ ""''lt~~ ~I;IJlt~lnl'ill~tfld \~itln 'elll:s hom ,m.he, species, b!llt ~JHI bulk-FIOOre 'than 40 fl'ldivfdlJalliile'~-had been, Q\I''Ilr~o!1lle!i!V Hlel,a cells, "Tliiks SCH~ or scena rl9' hjllil e'nliild' mli!IlY, man¥ lilllle~; peClllJoI.e iHh~ HIOU91i1~th~ We're w®iiling wit" ~rU: tYiile ®f cejll~ [were lalle r ro'u nd ~Q be.lllWlrki 1'19with HeLa cells .•" he Sa,Y5, N&®n-,F!ees, pllbli~fiIed hisIF1E!5U IL~im <I series or p~pe,FS iill ~dt'i1Ct' iFI 11H? 19705, u!'gi!l\ll scientisl:5 lo s~.Gpus.il1ll1l ,0Jlti!mi !1ated ~e:t~ lim'S,. lie"e~i!ll.l"'~.e IhBir pr'B"iiil)U<'> r\e~ea~~Iii,an~ em p~D~ ~irl'iple ill ~i1ilily"(~I1~Ji'(Jt prolK ti!i:e~ $iJ~n a's regu la r~lIwrirying ~11eirlin~s' aUlllunlieily. N~I~ol1- g,!!'5 '5 rll:Vll'llaliOIi!S 'tlir r~~ Ih.e' oommu nil'll i!l1to a frenzy. Miiny stldeswere [.iJlled into llIye'st~on, and! Nels.on-~ee:> Wi!~ 11l11lmi!lg name~. $'Ol'll€ bil!ll~li~'S ['N!l:l€d ~th ihD$liIHy. and t!lalU(l! irl an ,edII orial {~lled I)IJ el~,jHI-R>Il! e S <I NseU'-d PllXlirnler:i vi!JHante. ~ ~f1'<I 200]1 i::@mmEmta fY o~c:e II l~!l(l'iluthe f'lt tea ttO!l1, Steplil@1'! 'O'Brien td MCI in B@l~@sdl~ lII1.a)'lan d, ~ ho f iliad W[l~tetl ~itlli N~I!'>()fI-r:tees, i'8(,allM the lelMJml: 'llultlal1 ~nn,oaiioiils \\I€!J~ ~n ,~d~€!, ed ~a(t$ \',I~r@ ~llIpil'iuin!y f!ll th@ mlIDSl pH!'s~ig~olUi:; l;;.bIlJC!Ir !orilE!s .• and dis(U5Siorn~ o,r the p ~olll~m ~os.ta:nJ~ Sl9'ImM,lIrnce or d'l.l'i.t~ty. I'JIJ slw'n-R'ee S 12i;!@f1 rememb ena!n .i!n(ll1y,nTU!UJJ~ telegram @Uering ~(I ssnd 1fI~m a or1eo-\'IIay lidrel to S@utlll A/fi(a. ~MVa im Wll~ to (1~i3frU Wi a morass In 19'51, <I :n-Y~I"·®ld Arrtc.an"A!n'I~F[:caIJ1 WOm81H \!las, alilnlri!td to jOillus fh~llkilil~ Hmliliti:!1 ~If! ~~tiJl1l~r@, Maryland. f@r ~r~~tlTli@nl r{lT uf1,liQ!l un.(I~{; o~ c:omalm'iflilhoJ'l. aim:! i[ W'a!)n'l e!3lflY,;< he ~ay5. lhe h®~ilal ~ef1t .as<kmple cd Iner (anCerrmJi§; ti~u:e to H®llki FlC'>j$S'lJe (UiULire I The attatKS lJ'm:ima~elv ~ool!:.~heir ~olt liiI 198il, 'Nejjem·l'lees !!Lliit S'Ci· ecxllert 6@"(ugle (iey, woo SIll OO!s!;M~y udll!lr8'd~t ~f!lhis LOIb.IHlellrieU<!i.:i!(lks'~ e I'It.e and olleruKiI 0!!1 i!! rl :;J<!llileryi!l Sill! rra nelsee, f€IT~(i(llU~(<I!il[,€r ee lis sp~atl lhrouglh(lld ilJN b~!liIy1Im]1@1ji~f1b!l<i ~~~ ~~H~d H~La (onliml~51.o S~ln~fI!1!t[DtI~,y. In 20041, GNtrm1e lilu!;!,liuil1lg [If I.ne BNkeley. and hl~r (CDI.leiil'9I!J1e~ SIU r'lle~ed her. And her im'l1:Hlrl.a~~red cells, 11'Ii!l1l1fd!HeLa eells <lf~er Illw.Q.uk~l.f spI'!OIad Uj 1II~'I,i'IiHS'lly 0,1 California, -'t85 ~esea!n:her~[r(lm 48 CO'urJtne~ who were I,!t()rki!il9 Iffuou:gn 'ah$ scress Ihe ",,'or~d-aliid nat at\wys OOGllI<'5f \'I,Ii~hIle,m( (ell lijllles <lmlfond Ih1llll9 were I.lsing ~ res-e\uc'hers h1!d reCll!e5~ed '" s.1lIfTlJl'le ~~udy. lenr III 1'966. Sltiirll€y 'Ga~tl~r Dr th@ JIi!iiwlic:i~lrnTYIl~ Cud~~~n l.in~s~llall ®lhe~s.!lad s.nO\'lj1ii ~O 1tI~ ~olliltamilii~t¢d ~ure [olleni.:!'Il riD!JI1lI!1th.allli of the, InrM.20 hlllm.alll 'tell lIy H1ei,a"WU1'fil'lBlIIt!lu~ng c~md!JI(l>ed a puhIM.ee s.ear(f) W ~h,~s,e~~ bl~sh-ed weredllF@II1!QS.O!!JHl~IY clliJd bilJ'th emkall¥ identiFy the' niUIIli!~r o~ Il'!.!bli:~aJti~!'!s hom r'llSNft;hers idefllll(al tel iiiilla (il~ls" All. 18 lirtei) werll1 kn{IWn ten hli!\!'oe wro.Fl~I'f '1J,5,11'1 H'I1l'N:CUl'tOlll'lin a~e: cllille~ a~ tlml!~h thEl'1f Q wme rf()1111 C~u~sii;ilrn im;livld.ua!li, Y~t Gtlr~l€!r rloillllldtln~l S'lm hiildl;.€!lls or ~h~Qr~gll1iC!lllin~, Sh@ fOUlnd a ~l)I(i11@r each had a ~enelic '!,!,arian,t oran enryme ('Clund mnly illi 220 llIaperrs bel\'tE!u 1'969 and April 2'00'4. And 'the ~he ~m<l!illli!e fQellllh!Jge Q'~ A:rr1~jlil-AmeriC:(!IlI p\lPIil liil ~~ on l1!.1!mberoF pl.!:bH~i:!t~CI!n~on re-sei!r~h using cei~l ~~rles ~~lIDWln to h"Ye i)le~ome ~(jlllt';JJmiJ1i1led !!:iy Hela, had I.ih.p,t 1La1;k.~ h'iiJdllJebmgelil ~ 1G1If'U~f~ubl i~h,~~ h~5 ~~Jildhlg;j; in N<Jtvre i~ 19'6&, lJII~rking bli1enr~t !TeI!!Qr~d ~oiiI5'e in~r~i!~d by;a fadQr Qf ](1 !;I~"'IiH!f! 196'9 o1!lfIli :20011, \'II~~Fteas,the [{Ita'. nti mber 01 plI bH.a'tioflS, hald irncllI'aSioo or Hda ool'd.annnl1·atiM.1l was miil~ Ihe be91ftFIil'lg. A rew ye;a,rs la~~r, WalleI' Nel-sorHtees begtll:l disIly onl~ (! radcH of 2.1. i::jj\i'er i Ii! 9 (~n'l'a 1:llIi ati 1[1 f1~~n I in !'!5, flrQliIl1I~ b(J,r~lmi,~s n 11M p~rll~p:> Nel!imn-Re~~ \\!iU ffilllilUy '9I~t llii:> dllll~. <leras!; [he world, ,A,'t. ~ ~~ lime, ~~E!'wa~ al Uhe Ui'll (lIIh!l r~ mile'J ~nenti5l~ ailE! I1IOW la:k:illig LIP' ~k5!imhl ~d!ifl5t cell lin@ oofitami Ila I,iOm (~eemaii.!tlfi(~~. AirlJl:lnfi20 Oil, ltle La !iiora'ltny 01 ~I1e UnilJe~i~y fa lMom Ea,.l:J~rkdey, al olf CIa kind, Ch:aId<Cle'.rizing!, ~l(lriil1Q • .1l!ld disl rib ubgl taU Ic.~'tm y m ilIliI'~. D'iI eo ~e Ih U lin 'I'l 50det1l' f'Clr 11'1 itro Eliolcogy pli.!~~id¥ ffi(Dg:nil'lelli his {OSl~a V ~ilne$ r ~r lnll! ULS" MatiQflill (;3l11cer II[lIS{itlli ~t~ NO). IO~er f rom H,~Jilrif! Hill B.ilIC ks.'~ ce·r",killl trilllullio!il lQ s.d·~n(@ willJi ,p liJ'~I~lin:e~ thf€V@IlIl~1I11';iI1~ard. -111:.(. HIli'" lha It HI :rears. ~~, (OOIUIIIt@O219 [Ofll<llni rta ted C.ilnCer,
N

116FEBRUARlf 2007

VOL. 31S

SCIIENCIE

'!IiIWw,sciencemag.org

NEWSFOCliS
I'Lml'!'>everi h:..n iOIl'H~e~.lJ~K especially mlTlong y{Ulge r n:!;~nllt:lkl",f; ..~'.n.(l are uufanuliar w i ~,hi IS, h i 5~Qr~'. T~H;:' j ou rua L~ and agencius targeted by Nardone seem to cmbra ee his \M,HIl]llgS but not his se lut io ns. 1m an e-mail. S<l,~ly Rockey. de puty director of NI F.['s ffic~ 0 f' Extrainti ra I Rcse.ful"ch.
rl 0 00 N 0 00
I

Researehe rs .soU'mc~.~ mespuhl i~.I~papers on ir!JJi"id~wl mil-;.•;W1p!>.h(lpi!flg m\'..,J'[!J ~be, u\;<~1cfthe COml1r~!!lH ity about II partiC!idsJ eel ~ line, Btlllhese warnings ;!:Ir,e~Yj)ic;I~~,Y :ru·.~I.dCl'Cd [(30 l>pL::t:Ha~iz~djournals aillcl fa iI ~,o g.nab theaueuticu af'rhe larger sdeaHiHc eomm.l!IUlily.fo[ example, Merdeehai
L iscovirch, a cancer researcher .11 Ihe' Wei.zm,LIUIII[l'Is't:imt~ ()f Science ill R~~m\'ot lstael, S !l}'S. he and 11 Ia b \\"3 sted :3 years is bee .. rse they h,uJ:n"ll1.otk-ed <! rub~i(·i.HiQ!li u re\'eai~n~ d'l1U the lWO breast C<lI'Ic,errlines th'ey \\'e.r'e ~:tudyiUil~ were n,ot ,U,;lU01Uy

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~Lllhollgh l.1coutinues ~l) l~M,;; and d.~~ n-.~butt: ··N [H is awm:<l: l lim ~h:e:cO~lI:;'I!]li~~,I[edli~u~!>:~(jr dnl.~ltiSl'~n!il (!'i« c ~mt.'lm QI!],lli ()!l Id~nt~~Y' h~'I\I.IDI!IJ\.f,illl!JiCrp.rinitinof,h~~ ~~PlI:~ cell ~1l~sh(ll1lS th,llllll'l!)sl, ~ 9 sideber, p, 929~, cell lines is Oil seriif lItOil a'W"<I~eidel1lica[ to! til e (11ronic m~eloge;noos [eulemia Une 1<·562. RS·'1. lor B~mp,1eJ'had iJ,e~lIIth,lJllIghttl) ~~. n a(]fl)~ m~lllog~iIlou S~ellk~iI1ia line. ~ Q~·S iss ae t hat 'c:all ,j!!, b~lIr'thda:y I'!e~sallll.!lti 011111 rez.;u~Jt i I~ ~oss of biclogica] products Ne'i!;:of].-Re'et:: IH<1y have f'li.led tiD srop the 31i1d render research \l,'Ol'dl~ess, ... T:l:H:: they It!!\'\!.: 1lU!.hentic3.tcd their lines,. Sum,l:' ~pr':iLd or Hc:L~1 (.'Clb;. bUi. N~~n:l(mc is takof seienrisrs call be iissues need Hn be worked out hm\~e\~Ii; For i:I~gup his bartle, The ret ired di'l'l~cWr 0 f cm"eell" ,Hid reputatiun ..Jf~~l'L'ld if research ~s c{}ll,dl,ict~d l!Ii$ij.ll.g ('l}{;lliib.pie, z.;hOl~M diJlHft;q ues I. ~oli'iJ6 bd::o[~ RJNi N~L[d~)f'l~ A~SOC'UlttL:::!l. ;l.l:ii'ol.t1:chr'io~.ogy ctHlt!lmill.HitcJ c'!l:II~,il.ine~." Yet. :Rockc)' trainil;ng comp<lllDjI. N.f~rdgllD.e Ilms for more Q r11fu! r a pi! perundergees :rev k:w'J "If !'I paper has £:one thQu!;!;h thepeer-revjew argues: "I ~ WQU Id be imB)racri c,,11 ro th,m 2: decades edt~c,Hed gnl~lmHestndents req Ll ireaut hent uc:n ion, 01S accud it i(I n of precess .md lhe <Il1lhors ,1Irelft!lbie to CO:I~and pnsrdaes al NIH nbout cell cuturre '~L;lcln~ ues, ··El.'ldl }'e';:Lr~l gi"(l ., l~,~~w~:t;: iq Ull award W' C'(;lUlim;~s ,m~ll!>~d rouline]y ig! filrl.1l Itl(;; id~luut)' GiI'~tJ!~oeU ]inl;);, its nol ye[ we rhousaads of baste science studies [h:n C Icm··io!.,'I'Ii2L[ weuld dot !lays Chiilssn. ~f[lw crass c.()a~!Jllm'~~lmi.u:I'." e Mt'yi>, '·AnJ each h Nm funds ... , NIHhelieve3 that profes~ genetic ~igmHlIf~t.~ of al I eMab~ished cell year. 1 !;Cl ~~lesame bla~kst<l:re)1 timt tell base, sU(liiilal.soe~.~lu.c:':S~u:l'ld s,eie.iui~15,dum~s!J~\I~s ]]tu.:s;w-e:[l" ilvlIi h'LlJI~ iju ,I pu[o,1 ie d~L[.'~ :ffillelhil!)' aren 't ad(lp~iI~g th(:~!(jdlil1 iques," :;:huuk~ be dri\'~;ll~lh~ prnf~;L~~i(lnQrm"Olrd ~n 2005. he happened ~o gi\'el,h[i:o; leethenjtwnuld beeasier tbr j{nlm~l\i '[~ste~) best pracuees ill lij,vcddlill.g cell eLI hurc turc QIl,!'n~S, lhb~.Ulhda:y. Afwr the class, n in.ncses DcW~iH. As joumals wrestle wuth the preblems when his son asked him whether l~e hold a COfl~O!I.lU iL~<!lUO~ insread of placing the pnsad 11Ycen Unt; m~x~up~-R(ly.IU.,t)ldls go€:,,, n,:ll>plollsib.ili~y 01] Ih~ fI!H:iJdu.ng gl!lncy." a ~inhdliymiSOIH[Oml. Na rdone I!llal ized thill i he W"$ "':;:0 dl!!!I!ln mo"d",ibout !he reluc~ol~n .1!!~ W e~l!'i!ll<!h;l lh,n j()!l!m~.l~would lou!!f!ml.~ Oll~) he~,<;,it;l~e a~!ier! "nnho!'il}'. to have to retract 35% 1.04,0% of their prcviranee of scieruisrs ro ackuewledgc th.l: "This :~S~~ rC(I~I~r,em,Gunt tbat would be .illDlpCl'secl, by the field, not by the jOll:L'IHI~.,." om.~>'publi shed cell biology papers 1:0 weed seriousaess of the problern.lI1l<uhe decided mit ijnval irl diU:iI-som,~ ot£1!nuZ:!Il.um~are s 'W do mOB~ ~:hillil gi,,"e .'111<'lnrll!.wl I:<,t~k W.,. MlyS SdlJlu'e Deputy E<Ll~.wrfor B.iu!ogy 1(;\1,,' bio~ogisls,. K"'lr~IJlH KclllDcr, .. ~V~do Ilm have an explicit u'yi Ilg to heir und ilfercm ways. The Society IHJI:h::y bm \~,'flll ccrta il1i.~Y keep our cy,e mil [hi:; iar In V~l:itOl3iQ~ogy \villihoM II synllpOsDLln:J, Several weeks later, Nf~rdoneplH at lss 20tH <l]fU'lmU~me e.i ng ~11\vhh:h Yvonln e together a \l,dli~epapertitled Erfldfcarion of if ~tis !foll1le~h~l,gtha,l becomes a stmuJal.'d." Reid {If AfCC \viH ",~lk..b£)lLl how cm:IiI:;;UlliCros:,,",C(ml(:lr~rhU1lfi.'(/ Cdl tim's: II CoUlnl' N.(II!II~ &rdn;iC~!ldy nsaud ..te DNA fil!l~elfIkli~~jl. "Clo.1'll'ly, [h,e ct1.rrem S,~.uHllIi.ol:l ~s, pltlm dlfl,tfl for p~pcrs. reportilLg n.ew IlLnnaUil f~i)[ionC!JI1ibe prevcmcd. N<lI,ruom::..Mm;,tcrs. mul }oseI1h!P~n'(me o fAT, C a:fe,~lso Qrgalllem'il'fyonic stell] l;:eU Hites, but l~l~~po:licy i:m'oJerable and req~lire:J a. br{)oadC'oo~-di· doc~m't exxe n.d to aJ11cells ••,.~:•III:nk '~:e 'wo~.lld ~,ziill.g a confen~noe ~o dIscuss sr,<IIl.d.uxls, and i IiHIled effon :i,l1i\.'ohd.n£l, those who do !l:!J.~.dehn~8th,U I,;n!~id.h:<Id ~o pf10reR:Si~l'"ii~ the white reM;::,u'<;h"I"lJ,nd .re~>e'I!'<;·~ ,11~bJ i,.,h fi nd iUli,g~· iAg!:~e\ ..'ilh. lhe ~e~ti¥!lenr' \.v.~dcl:mlnp~i~LfiI.CC fCl'rmlihc.iukal.~,o.lll. And paper. say::>'Nil(a'Ue[)cWht II !biology cdho:i: rc ~C'fu r.ch .~11lI d cd WI eiH C rcs'!.:.f~:t'c: hCii$,"' at Nm~u'(?, Om "Y(I\! call 't just s~ddell~Y say A.leC. which has fordec<ldes s'o~,dhnes ~e''''II'!i ~'es, \l,(~ ~l~ifi:d io V<i.:lrify ,~~n ~i~l~~~;W~ ~on··t l.l~I\l11l ovt::n!lJk,~1:! [-I€L;~. tectl~lt~y d~~idfi:XJl t>tor ~ N,lrdnbl~'s ··..;;~Ufor ac~~.onr ~wtl'h two b}' m ;:~'nl,.d c h <!QlgeR~mOfe r~gl~'~,I~ion:- .n~ !."b.s in '0111f ofI'ices ... ml we JC';IIl·t ,~heck the fo()m inlf;jIy d iSlTibt~ti ng the I ~l~I:S,e1'i:.cefit t~'lr d ; ~ irH:rtascd l':Jut:'!Iti!o['i c:fftlrts.. N~ml()1iI11.: line~ uu,~c~vcs*M.'1d it's rro:~nll~~mstc:i" ::;,peehd f'C<illi:CSliS rrunIil,i'~se,1~rc'I'il'!rs. Bu[ sa.)" and ami: Fro:l.llll1iOUse:" these' efforts wi~1 hiO'l.ve ~imiled dJect. s<lYs ~.. <L1'gLl. .. <iiIjmll·na,L'!: ,Hid fumJi n.g<lge ~c ie~ ,es N,a;rda~!~ •. ~f julio rn;I~ ~Hnd g.[~lrH~;l.w .. ,[dulii lil RebL'.',t~C11 O'i1~~;lrI., ~'u~,ivl;l edlitor nhh~ :Si ~~.6irmldIIt'illm!ii.e ~tlrl.C~ rui~~ (Hi .ret'i~<ln;:.iJl~rs, .kJtJl'r.'Cll ~}fllie !llrllio,nal Cfl.l!c'{!"I' b.ii~fiml{~ !lg_en.ei.cs won+.: ri.1abHjal~ ,cell 1~l1c authenti.~ fOI.'cj Il.!il 'I hClIDl to ~mbm i. ~)'rtJor:or celli HIl~ b tJ ideun.l:it y ~ It]ng with thei f n'iI~Ulu:s,eripts !I uncl (JNC/JI. ~aYi!l.ii~,'i.cwelt~ :!>un.'il:':lii1'i.e~ r1~is~ e~lluml.\'V.~!I ~. i[}kJgis.I.5 I'lCo1.."<i, hlil L'It"Jj'idlUdl'!~. .Iill:' ~ trail. pro.pos<lls,. II.'especlh1ely. This. he questions about cell~i~.e M,entity. but 0!1l~1: ~S<li "s;t Ie k, l:mYl n~ 111<1i~ you don't do Ihis. lherewill tle a ~'olils:equ.enl~j~," illi ,~ay~. h..~ '[0 be ~upr~.e!l1e~Hed. by !J~l1t'wt;ld r1;lvie'\lin~ N~~:rclon~"s \.V'hil~ pwper" .INCl b: plflin.~ CdlUC~i~OI] clTt,)lrl's to i'~lere3sc .r~·\\!.\'i.[~n~~s,s,may [flk(! aJ fi:m~er .I~.ne.Thc:joul'nal -R IIfUi (IHAli1iEIR:JIEIE ~ o,r the' ,cn) ~~-con t:>lm~n,11,~flnproblem. RMIIi.!I CIl.a ttil!rj,~eis.a sd~(~ I!jri~.w in 1N<!shi ngllo~, O.e. llinS to be~llll)1~ing tll!thors to confil"lMlhOlr

1'<1"'1, .,h~US. Nauoual Cancer ~n~luu~~e kuew and ,Mlemp'~-e~II'O Il'L!blicile,


rcl.f1ted-il

tol d Science !h~~

or

or

:h

NEWSFOCliS
...IIlQ'l!g tim~ (Q~i!!9. T!'vis;5·m~ler·r;)Jdili!~ IFI'AlG "'Iliuill a l tliie KEIK~abCI!al~Fyi n Ja'jian o utjierloj'm~ lF~ld~ti(l<IiI;;l1 !!'liIdUl;Il rans. S

!l.u'e<l!m)~lled with a :;.ym:h mtmu This prow ~yp(lm!~lehinl; WiIS, d~~U.W'llldto test !ll);pl tc<ll ions suoh ashadrcu dlempy. which
tlSCS prorcns 31!lJ ot her chargedper

~ uch::s, to

cleMmy ruJI.TIOI';l., ori ..!mas ;~Isojoined KIlkhil.'O M


Mis:11i ma ::!I[td ,(;'~)1I~1Jgu'l;5.n Ky()ln Ui,1ii\-'Crs:i~ ~()

PIHIYSICS

A ,H,all~Ce'nt:ury ta'l,e" ,AI'le'rn,a'liiv'e A"cc'81'erator' Takes O,lf


1t':SI1!:~~ 'Clui,tea tydotr!@'!iI and 110t1~~aUyaJ ~yndi!I!r(l'llrQ!f!~ buttllle ~ixe<~.ifi'e!,"daH:ern.at~l!iIgl'!Jlrad~ien~sV'IiII(hr;IDi~Jolfl (!!lllJl.o1 U811~e ,il, ~"Il1ote new Im'~efOlr p,ull:idl,~ be'iillliI'Is
When ~IWs:ici~lSsmrted ,~~e~'e~lOa)if!Jg p.:.n~i,cle •.ocellemtm:.; known ,I~ f:i.1re(I,fiekl <I,llel,'n}lillg~ !il:r<~d i'~~H ~yrlc'hlrulWI1,K ( F fAG s), Dw i,gb l Eb;cnhtlWCr W~lSpIfCSid~l'nc, Oof I,he Ul1hctl States. an,d,Elvis Presley w~;s, 01, pwm~$irlg Ul'lklilmVIn, Nrn ....5[1 years later, nt.....'fln:hc.rs :!I:UC I SUlflinfJ, ~~}blUi lU FFA:[i)1 lh~t re..H}'work, Propone,flUs say these aeeclcraicrs Il;l~y bli:ngp9r. ticle i'J.eams to bear Oil! new r~e1ds,md <lpp~ica= lim];;: dl;!s,l.myil'lIg Ill~lli:)n;\'\ii~ll p'.~I'l:poimprod. ] sinn. ~ 1.1!o;hing the ha I f· ii Fe of radioOlCli~ie
ill~Q '.~e acreleml'Of. n<lt C\~~QWj)lH~ OJ! ceih!i!g on '~he~l!e3m'$ illtensil,.l'. ffAG~ ~.Q~.\'~those pr'oblegnr.;hy n~~n,g .lmlH!ll1~'lBc!1d.tllhm su'tys fUl>Iedi il~ lim'!:: but 111m ,grm\,'s !{iI:rlJDl~e:r l,UIi."wd the {!lJI~Fiideof tile ri !'I~, As pan Lt~s gaillllll:i.ergy anilddr,i,ft fllll\WiCd ~hc $t~lll~ f~eld k~ep.~ iile!l'! O!ll ~'01iCk. A'I!.are)1lni:t :p~Uitnc~.('S :ill:l~ il<m~l!'OfCllCfY;ks can all ()lib~l m w f~

test 'wIH:~her an fFAG protrm, beam, can sh.,h tln: h<llr':'l]fe-<l~'Ki Ihereforeulre ~:llre<ll ~o fl!IUfe g.en.t:f<Ltions---o f some h:mg-.~ived mdimLC'~~'I'e \ ..~ ::ile;::_Other ,uft~[llpr~ ,It ··tr'ln~lliiut1~linJl" nt :lrLlC~ear' "'VilSI.,","tnvo.1 ve uslug In ~ru" ~.c~'h:;m~,m:s, b III I M ish ima lh iIlk:>F'fAG:i' have Ihe 'edge beeause lh~)'" ~I.I'C Ch.l'IIPI.'i'.,sma UiI>D", 'Hid mm:,~ effil;:ietlt '·W!iJe~~:;,.. ]Iill~l' ;;;I~~~~I~~'lt{l. w{l~~ld :IiIt:cdlm be up w ;;'I, ki~Qll1,el;C,r~.mlg. ,j1,]D ff.A.G \'I<'01.!M I)mbably be .iu~ <lre'\:'i.'~ens QifmelierS in di.!lutt:l€r;· he ~~,
F,ir$i. however.Ff

AGs must el ear some

Xl'Rtjur l(."(_"hi~~C!I~~~ul'tllk'i. KEKIUl~so far ~O:i1IiCentra,tedion "scaling" FFA.G"$,in ,.\lhrch tile


(lroi:t~nflh€
p;ll1idt)s!l:lIi'iI.'lJrb"
corilz;;I'I~t!1halP~

,I

as the size o:f these orbits increases, or '·scares." \'I!~lh momenrum. Bu~these require l<u-yel1il<1lJill1if lOS aad <ll'e e:-.:trellilel)' ,~omJ)lex , MLI~~hs un~I1!~eT d n'm~'", c(lI11P<lC[. [hi;,l(uisIS ...n SHy...wmll~d be ~ "'ilollscaliJl1l_g" FPAG. i:n whk:h p,lnicle orhi~ch<lnl;."e slmpe \v~lh im::r-e'l~ifl~ c'.nergy, NO(i(i1It': has ~J[I,il<r Ullll yCl.OOCi:llISil it
h;;L~ been 1~ssl!!fIH';ldltla~ sh,O!pe·:dlifling the orbits make the ·be~nm.~~I'IiIQ;,<;1 [ impossible to eonfine, 13u~,ll!leW U.K.-lied consnnium ~r
'Ulliilve0i~i€s OlltlJ nl!d~;:,]tion urH::{]o~ngy o(.l~f1ltl:.r.. . llilm 111(1: beamwill

the ssme t~me, emil~1i~,g FrAGslopJ'{lffi~~'lepmwride beams rli!UI(: eril~rgt).ric ~illllllla cychurQn~ .n~. n!.ol·~ ~Illt'(;'n~limn .1:-;YJ.~ChJ"O!:Wllf1l·S. The idea W:IlS ori gi naUYP'1I1 f(J·rwarui 'by

h!:::h~re~ ill"Ul~oh'e tMt ptroble!ll hy c~!'Culat-

ne~deJ to [I{!:~er t:1Ti e r 'IP'I lei es~ ns,u ~~~ lh~ m'lll~him'! l.imiffi IJI~c clfl~r.!Wofth.c bc<ln~. S.ytiC~[OI~:()DlI.!:l, o",~ro(}IIilI~ IJ1~fl.pliobl~m, y b u:-1inl~\~ri1lbleIillJiPle'ti,1.) lields I:ha~ f'dmp ~~)as

Jepanese physicbt Glj_h~ro Ohlkaw,l in .1953. Re.se,m;h.e:rs inl.fuJe United Slat'l;::.s hlliU lmme aluM :revolmiol'!lize .Lt~cekualm·drii en science .... small FFAHs ln the late ~950s,8i~HI c~rlly iln,gellu:rillL." :mys Robc:~t C'y\:villskil. an experi196{K BlIt~~l~ dm·~.t.ll1tyofcfellIlJn1g t~e carem:emtli. physki~l at Leeds Ui:l~vefs,ity i~l ihc V.I:<. "Every uni.ve~,:>i.~y lllOlildhave one," ~ 1~llIy shaped m.<Igmetic fields made larger d~\ii,~~t> iunpl:';;lctk<Jl Like ~h(;.': yndnmrollis8!nd (:}iclo[L"'tm~ s However; beuer IlfI.agnC'tS and cempurer illilnillim to tooay':> 5'CJ~misls.I!FAG$ use dee~ricf,~e "Is lO aceeleesre bunches 0 f, Il!i!!.~~'e'd si mu lat i.m s 01· Ihre~dh1l![;:nsi{lm! ~ mtlg net ic field;., have renewed hue~:SI, in FI:AGs. lhe partk:le,<; suchas p100rfPI1lS erelecunns arounda the hwe 1990:;, (romp-11m:. tn !'in~ ;;'1l {..'Jo~to lhe "'lp~ed orligimt [1\I! !],ley ~llif:' :sp~ir'(;Iune i~. b!m.i~d a 1t(.'I.JI:dino fa.cl{)[f.'Y, a InlillibMI.i.{)Ili·dol:· fer in tl,'~\""LYIhl..)' guide lite 1)j)I'~iclesareund I",r rn,ci~ilyIhal\"'Q\lldprodm::e e-,;'oremely ~he cil'cu~t.ln (yc~o~r(l\ll's.a fixedllllagneak ir!It~rh~~ .IliUgh.~ll~~Y b~mn~ iJl'" r)~luln:inns_ :md fie Id fmcl;:t'> !iI. ·IiJe;,l 11m. c;:h.. :~~dlp;:llnid'~5 10 of ,n ~mml: in ~ 'l::i!l,;~e.h'iI!~IJ1I.,"Y~I~O!r~i~IHl:!il~ "'k~.~k:;." y(l)1h~h"rYMod "n~ 'I::(lHe1Ug~le!> H~~Kr:K. buo.';1 til:(: panicle!>' sre~dll.!ln.! ltlJt9k~dtClill $pip~Lrtide p~ly5it:s~~lb Ill:Hlrilokyo thOllghl ih~.t I'a~oLHw'l',d, Th,e size of the huge magnets .1Il FfAG 'would be ide<li rOI'<'Iccel,ell:'lI.ing ,2'000, d~,ey bui It Unli1111

~ml. teasi ng ()1Ji't the properucs oHlJlc most :~hlldmne'liltalI1i;lrti!i.:l'es ofm~.ter. "FFAus
WfUS~~"

ingpanid~1> &0 qukUy

uet

h.we ti me to spread sign iri~.mlly.


Tille (.':(]!1I~nlim'll ruct'l1ltty rt'Ct)IVtid: $16, mil-

liO.UlnJ btl ikl a smalll dCn~QUls.mtion It(mscal:i blg d!lCI:rmll H:AG. kno\:vill m. EMMA. ill '~he UK, goverurnenr's Dail,'esbur)' L<lbo:rl.Hm:y and d~siigl"li ,! rrolnlil devke fur hl.'l}(jmJlli1l~rar:r

"There's an.lelcmcm orris:k lnvelved' says R~~eir EbrJo\v ofthe Ulllversity {l1'Ma:m:!hes~~~, v.'bo leads the UK, p:ro:iec~,··13m I:'m, ctmlklen~ it \'IrLli work~l 1> ;'1. q~I~~i()n nf h",vll'il"l.i ~IwiU be 1.0g,t:I EM MA ~~p~mjl. lIIlilllng:." Mefililvl'.hi ~c. F ):ila}'Sicislsat 1l1,'Qokh;;r..~eill Natimlal Laoornl.o:uy
Ulfl lipton. New Y{l~k+worl!;ingw~ttEn ~~~ln~h~rs

Ir(lli!~ ili!dl,l~rry; w~ W ge~r~!Ill~~~f!g I ~olP1lild tl~(dr O\,l,II],n{1ilSG1~~irrig fk:vire. Ir a ~Igoes as p.l anned a nOHlilCm'1~,gr-FAG i

Ib~ 'p~Lrtic~e~ITi~;N~declcTe'<lIle [Ile !i~lLlri91{l&, ~.11 (]o~I[I,rj 1:~l~i~lg p<lllel1tt'i Olt 11 Br~Ei~hlmlOpitld hi ·'I)I\10f 0011· pr ~11iC'i p~e" wli.thill, 6 Oil' 7 ~a~ .• say~CoJll!:M.ilil1i'l ~~1CI1l!ll'iit'l' {k:v ict: 3 iii.d hll.vt:· )i'i.liK.I~ em~~tlfTLlt:'w,d.H lhl[g;l~f B!t:dtllyn jt~lie-Ji,. 1uemtc~r:;p~ci~h~bl ~:r IJllruwfL,iS:t

~ ~he p,lf1Iid~~ ini;:'~'IL'."e ~l I <e'DI!;;rIDi• ono~ the But ~ I1c'kli ii_t;gc9n::d up ro.iI" h~.gl1-cncrgy particles.
~ )1(!~l amw. inject !Mm'e.]!CI\"'~l!lelf'~p<I~tide:;. c

una ch ine wi th a :I':adim, of abOllt 50 ~nel:el's. y~..!C~thi ~ fITI.u;:h ine,~~I",e~e~~,t~d l}l'Olm~S

::>~I)II [OSpi!:21J. Bjm~il~~hlliln,

lip 1.0 U)U IIUimu:n CI~N!lrOn vo~ts at!! rnme of ~!OO ~~ullc::he~ sec![lm~ ~wic:e the max.iimuln a

~EIilWIIN CPiRlitllDGIE 'Ecmin Cartlidge is 11iI!W5OOiIM of ~~ MWld l1Iaga2in6!


ijn B!:i5~ till(

NEWSIFOCUS
ARCIFiAE.OLOGY
(E<~~tef lslaml

)'.Tl'm.:ing the jars' 11fO\!ermnoe. .

Sa,ving a Lost Cullture:s IMel'QI!slliilhiic Jars


I'(onk Q!UiId Ib!afnhillg, InIl<lS51W .:HOI~H~ umsseanered a~I"(I:5s.laes 1iI~<l1I ho!l.dldues t~'lhe'

Cio~hon adds. ·'wow!M of[t:lr V;llltl:3bl~lrllsig.ht i~lE:(l!,;m.::ial (]~l~.liz:at:i[Hl in prehi: ... {)r.~~~ ~ Slluth· (.'2L~1 s.ii~Ii" d~il:i,n1g irs [rml£fo:m~9[ion rHI.IH a A loo.-reenllecrion of Mlhs~ stence l~lI'l1iling '~'Om· iI1Iltl:rI;.~r~ w ,~web oflllluml ct:r:rr!.:ln;.~hm 1!!<11l1~d \~~~IbChina mild [ndia and 31:s.0 imported. I:he:i.r rellgj.OJ.l!l. BtlddJJti.sru and ]-I imillli.slnl,

'r~:sfilofSQMrtlr!eilJst Asia'5 firs~: d ti es


XIIENG KHIOUAIf!iJG, IlAOS",_Yot1 eould cas;iJy insert ,J fl!'lI-g W\~rf! <l.dl~hinl;o O!l!~y ofdreoowU!5. of huge' grny jars MI'e\MI1I, .1(;I10'5S •• wlndswept proml.lmno'J:yin {''"(;,mm;J1 L,lm~nra hUlsb;:uld l!ndl ~\'ilfc. for tI'b111. m:umr. or sevem~ snl:l'Llll..::hiM[~ll. The h~U,ow-ed corpse: of .. king. fir ~hf.~eklng .p'riMme[S 1.:~~Ptlilli[(::~d~1I1:mn~e: ThcgrLlCMll'Illil t po~S'illl~~ilie:s, ,u:e li:ndllehl'i, Alth(1~lilh~heo:ri:!ilili1~

mar. So ls th,eir IOC[lI.li.oll;: There are nearly 2500 jaffi. lids, .,m~ ~1Cl\nedisks at 52 llilowim sites 0111. the PII a in or Jars, defined !IS the di".llIiIUJld.-l'ih1.'lr,'i i,;,Id, 15J}[1Q··S>qJ ll.;u:e;;··j.;IUille:::i~:r i

oflhc p1IcliJis1:Oric ~all.dcSl:QUI{,; VCSSQ1s are .~az:y, d iv.~ning rheir purpose reqaires ~Hlinim<ll.
.l'LllJeir m(}:rbu(~i:~llll.gili.~i:iion" ·~~.I ,I«mnd thl:"j;;€

X iCillg KI'liI')m~l1gl'i1nwim~c, A conce ned efr'(l n ~s not",r U !1!d'eI'Wl'Y to' save ml~ of Al;~a:'o; I.OI'Idiming !1I:Ch;RNJI(]gical !::i:cldle~ l[NESC'O',ln,d ~hL;)L.H~J(~ g~Tw:rmUn(;'iJH rhn~.l~"\~lmI. n X~ell!lil Kh{l~I'~!1I£, Thong ~:hl~ ~ ~n I.~.~ill.?ilSl.')called B<ln Pu.'i!; or S.~I.COn.-e), one of ( IlH~'ejust lasneheda nc,v phase of a prog,mlu H! safeguard the rh~n Jar!'>before its of the h~ggef f:iekIS''I.ritbmorcelh<lI'l.250~~ms" gr:!w~ g,nmh;includin~ glR:;S nmujuarjou, ~1I1~Ucip11~ed .:HW8. fur ~htt: COlall~~It~lrthb-d in

Accord ing 10 ~ocal lore, t h(j ars weremade tiO::OH:H'l;:: iau-le». o'fU'icl;::\'\,iliskey, for a fe<!s~ 1 scm },(!,IT'S ago ~'Oo(.:eI,eb:i<1tethe :I.ni ~itmy 'f.~o!OIry (]:tKin£ Khun ChUoIiLngOyer au I,I[)Inliy ci'ltef'"· Uil ~I:l,Ch;lJOAngka,~ 11, the ~930s;, Ma<lehri.llc Cola~i, a 1"I,'enc~ a~ilaeol()gi8t pieced togelJter a mere m,,,,t:~d:J!I:c~l:ory.Neall' tbe tuwn of

j1US are graves," sa:ys Julie \"m D'efl Bergh. tfNESCO ~Lreh~.tjolog~51. for tilt.: Plaill1i of Jm·s..

She and other expert s speculate th •• bod ies t \W~ ~~~i:tiliJtllL;l ..i~ lDd~O:I:nr,'im;e. 'lI1;rJI·t~ll::],ithe rell~ru~s were scooped out fo.r buri:!Ii, The world's biI;:Se .... morU!ar)'~'e~se~~. ofl:h,e U,S, "Secret War" Vi;'J£.edhere in the seme 'Ll1)rigIM and some tipped over ~n,scragi%O~ !Ili1d1<\~ITJy '70!'i, ~lly~t"olS~r,~rdtoo brmvn ill the \\I~!1~~r ry Se!!· d Dcdph.c:ri'l~g; IJ'nC m~anih'lgoflh(!j3~'S"OOu~.d son, "are ~)!l(}C:~IL<'Iel~hu c.ni.g:nm." says Pe'~(;r b Bellwoad. <In arehaeelagist at Am;:~T<JJ:i,a!l shed ligh~ m'l!he reJa~i:m~!'ihiipetween increasNjniolilal Unli\f~'rSi~y ill e~UI'l1J~ml":f{]j SIl:l:tI!t'lfl\, illg seciul cOIbii.pl,tlN.ily {~lfId m.!.:lg~L~ifn euusrruetion." say~ pOllleo,ml!urOIJl(l~i{l~i RlJs,sell ~I. 1110 H1)e ~ ];;ltm'\,rs'vhjC'h~~ul~llrr,' they bel{lnM:~l1. L. Cioeh.oJl. a SotrlltL:~1."t Asia s;p~ialis! 91. I.h~ Their utter laek ef adorument hail cloaked !.Jnive~i~y of .~m\i'll, :~O\,vaCi~}' wilt] l."Onrp<lres .. thc:m i~lmyslc'lry. The Ur'.I.1S PI:es~ll.nalblywere' .II1N;;w:n r!tOm u.·II?<lJilJy quarries, b~E radio'[~l(; ,lfS U) Od'llt '''' ie haat irug and I:H::t terj l knowninego'lhlliiii: E:ri~aifl ')1 c.,roo:~dwijng :i~a~1{we~:~he _~::I[II"&1I:'I!!t'!'~_Saonl.:l~.cl'Igcand the i'i!lvaf F1Iices R,'lP~1Nui

UN. f~ency~ World Her.ii<lge list, One-las!.; is to poou..ct dIe Xi~l~ .Kh.tllt:an:g .~'lmQ1J~l ' frum an expected ansl ..~.g'j,~l tourisrs, Anotheris o:W ~u pt{jil:~ the ~mlrists Hl~liil~lvl;;S(i-on'l~111i.1;l1'\I)~.oded o:rdlhlillc~ (UXO) th~l c]~fLi~.n::> dozens or victims in the(;,(JLm~1f}'sid!e ..1.Ch yeaf.t! ~es.acy e

and

cOlmdian beads. cemmic pOi~herds. bmll!2 Ib~L(..'{':Il.::t5, ~li~cl. pc.iIIDheads. Jhe Mii s ;L'C5CIln·

m.d ~

IDle~hose From Iron J.\ge sites ~11i northeastern T1111ilmilrJl, S!.ly.~alurl~ High~m.lliil ;1[dj:.\'lfl~)lo. Ze<!!1.a!1..(L who is eo«:av.uin;g ,ITt'l<~isite [n,!n '~hl![ period. I lighmn pt;~ the Laos: !;r'.&\e ,goods to lihe 211d1eIl 5dl t~iltWl!fie-:> CE. Celani aJi>t1i'oull.dch,arrctlboilC3IW ash ill, iii D1eaffi)' eave in a~il1iu.·::;tom: Iili~ 11]a (!OO·p~e I, ~IDII.is~ Uifllill@ PI~~ililof 1111>,s.l:ie'!l'Ci.iJUJood ltm~ ~n~inswe:~f.lI;."Fermlted in the ~ave. hrt"h h .....; w 2t meter-wid c~o~ci;ll its ceLl ing-a natural chimney. Ash and 'bone were then inut!l!'1I:ed in
[~t; j!;lDO.

tist at II~cUlfli\lt!Itl!:ityufOl~O ~n[)m~ooilll. e\I,i N

\vhil,LO Hmt ... .jE,"\l;r·dry, l:Hilt! .0111(;[ objm::l:S

or

Ilrlf use Qfth.e d!ece<!~ed ifllhe .~Itefhfe \,ve!'~ lbwlirictl n'CX~ro l~C jars, Colflll iproposed, SNJIlC disks served .msgrJvc nrJl;kers .. Belbri:: c(lll~aMill~ t'lrdd prone [hi:. Iheoay... laos, descended in.to Cm'i1 nict , Ailer,3, decades I or~.me:mfl~ 5rnif~.I.h:c war in Viemam spilled oeer the border ln the 1%05>. C'el:lll:<lll Laos ~ bore the bruat {l·I'u..S,·b.u;;k~dcurpet bomb· j hlg~iln. ;lIJI,SO]l'L{;:2 milll ion ~o~sof mUIfI:i;· i!!: ;!!; tiomi-w disru.pt 'North Viemamese IE s.upp~y lines, "II's II miracle II~,Hso :!:l

s
Q.
,Cj

<,,!

NEWSFOCliS
Van Dell. Bef!'~i1say~ Nat slilfpr.h.ilil.gly. shuJ), of lhejarS~::!I.lfI. gu ished _ Th i!iI.gsp.icked up 'I.ga in in 19941. 'when Tiholilw;a S~avolfIgk:l.lafUitdy.lilmv direelor or L<lios};: De~)ll~l!1ile.l.H Museums llnd! of Arch;)~t)I.(]gy.tix·tm,·allx! SJln"'~RII.burials near ~he jm"S. His \\\'Ork stlP\1)(l~te[j O:~fIIllrS !"ie\!.,'of C thl.: jf~r fields as ecmctcrie s, 8m the bone fnlg" ments iDe uncevered were not ctml'.wed' .. "\lV'l~.
IITIl.H1)'j<!f!':>l!fVi\iOO."

cn~,unliri!ilht~ did ~hi;;,1' lily cremare Ute d~,~(~! 11m \>Ve lm~sere,' says. B~lhoo{Jd.~ntr~!,!11~.lilflly. 're

d~r.;:!ct~y da~e the SanClslll!l;El j.• C~(l;Cho!lI !',~u.~U!\,. gCSlS II,Isin.g opl.ically £;liilmU~llIlCdhtml~I:l¢K1;;L;lllllce~o d'~H.'l'~U~U:F!7; ry:>wb in s.edimJle]U~ c underneath. In [M~ recent teduiI.tqrue. :I:igh[ H~ erares t rapped electron S Irnm k:ll.lg bari eel c rystals; ~ilu:t::J]siIy d",:p!cl:lUth 0111IMCk~ round l:i)(lioa.cli:\I~ty mt<! durstien oFl'Jmial fou the quenz ull:p~md(Hrttl ~ybenearh t1pright jars. ·the 1<!I!Iitime these sediruenrs were exposed l to liighJr \V<~:;; when the jars \\'t:1"'(':': pUI UIlI pi:u.re," C ioc:l~on Sin.Y5. UNESCO hopei> ~o nlld illll(t0

Bellwood acknowledges th.. he has norbeen u


i:l1!J,~co f~nJ ~.ny "li,!Ilgui:l;li~ dcscl::mtcrlt!O t ;i.Je~l!ifi.,ble- pla.c,-e J~!I!li11~~.inll,'!o!l tt)(jOl}"·'
011'

111rS1. ebiseled, they would have been Vi~;lhe or eream-colored .11ld ::-;pm"Jded ~l'Ilille sun, :->ilys BcrgliJ~,~II~ ~L"dit>lilliu.lg dl(; bmJy OflD;lmm,l.l.lJl V~HilDcIiiB(l~·gh.·'T!uey mUliil ~.'ni\'IlIx.'ttn .~tul'li" 1f:~;;.'ie~·H,'I~,'"l:Ujds fu~nd OIlm~er ~iit~s would . Ili ~~:!il:' Qlys.~n 2:000, i~.higl~klltcl;";!I!e'lf Si tf,l f f!~elYelt~ S{)m~j,lr!->n!!diwen cut tn ~er\'l~' t'l ~he Three, m~lng.'>2L d~)C~inm(;'U~cd v.es~jlg,e.~f a u'oll!;;hs for feediing af1lLLn,~I.sor c·ol.llcctl]'lg tlrl.c o ~fUVC kept (Jut :scfl'Veng..crs.l,S~'H; cites ~L cu~n· pe.~lingrm)(.!em ~~1!iF.allel;'C"moodi.m, L.m, and jar fruct(l'ry; uaf n ~shed vesselsaearan al1c:himt dwinkiL,g w,ner. v,i~i,le ~idsand disks became 1];l.i~I~lmt~¥;..Al1d dmlng tll~ S~':Cret Wru-, uutold Mll!1.d~~Ill~liil qWJI:ry. Th ..il J1.lffi presumably i Wel[('l Thai nob:i.lilyFor cenruries h1Lv.e~€·I;1~~'l~rred in large urns before CI'elllttt i.o.l.l.An ofl'ici;;tl <II ilx')Ued on 100d;s nd perha~ ferried Ily bo ••t I"lrl mnnbers efrelics were bI.O'WI1. to bits. a T.hc $c!ln of war are sliiU plain [usee. Tillailu~.ld'~ Rt))'~LI rfllao~ ink! t,,'dr:-}i(:c? Ii:h~t·l.:fCt:h ,va)l, "lhli:l1i fi.[ilils~li:dat the ~ik:." ~i.yS'\hL~. I)d TlaoliiS Ha:i Hin amI s\!rrouml illig fields, studpJ'eIlU/CilIS kingofthe presel1~.dYI1!iilsty.upon his BeIgh., Eh!l. ~here were noclues to the artisans' dl;cl w~th ~pikr w.il,d1Ikle. ;JrG Ilo>t;;kli'ilarlbr-l'd iJ(:l~Hi't il;:;>_ Nor d'Q LHii,~i,1I'1l!!> iV~rrillJl.d'l 'IW<J)'_ g death, h,.tSiJ~tl~l r~a{l~d ~inillg pn~ilim)J in.., ina o nile roLJ~h,~y2000 \pe'rined jars.] ust one has with. bo.mb cl'a~cr:; Q fe~v 1i1I1ClfrSJeep ~mli )iC\IIm'~e ~uM~n urn fur J Iln'Ol.u]~S. before clX:.uni:l;· ti!;:l:I.l. BlQ~'lilaI'd eviclernC€: loll' such a Ilr.!ctice ~!1 an iLm~geil'l~tT.~l1ed bj' its makers: Ex<{,!',;;~vmin~ er.l~ meters OlJ::~'illS.". The l1ollpfOl"irM ines Adviat 'f.&Ions; .~ I".umailf], N ovem berl 994, E:uji sory Group I~asde<lred 175 ill );;O--:i aeludi Iil£ aae lem :Laosi s elusive. "WhelheJi badie~ were' di~lilk-'lcil il~~ihej,lrtliK ll~nb,lbly ~nlkmYl.~ubl!1;:· N il~'I. <1m~r>t;;lm eo!og~ st "H: K<I~C},~hi!ll"', u Uni'ller'" bombs, :I'(;loI;ke:l};.arti Ulllry ~h~U~.!I'I1QrtO!f~.and 20-llIII.~ II imete r f11IHi a irera It round ~-from, S~IY UUl f~p2ln, uneevcrcd :,]I radc!d bas relief J :!if1)'sHigham. around the j alr~ and pMhs <It lime d\i[ee main The A:si ..rn mle~<~ith makers too are irll<lge of the upper boo)' of a humau n;i!!le I \\1 id~ lIlPl~I.r.;I'IOO arms naar tlie lJifLSC f :lI jar. i I'I.M: !lLml~~c. (~~}~:.q i :;PCt:uifl'[etl thmthe: j ars n Xum:iJg KlmL!m.llIg !>!WS." V~1l Den B~rgl~h~ii> enSlUB'L>d th,u ;,a!!1if~CI;S,U1~ not d"~l~~ed dl!,ll!'!i.~:!il "lheyar~ VI;!!!')' f\n1>l;;'~ionO!lI:' ~"Y~~OlJ!I De!lil V were :r<!!~hioll~d~y .! \'igulWu~ ~m.::iet}'th.u Bcr.gIL. '·W~.lOc\ler n~~~dc l.heAl didn'!. plH~ny unincd s2Ilt. a yal.u.jj.b];~commod~lY. and. was el.ClI.r.llI:lilt:c I"C!SCtJ~.illg :9 d:.ol<!:Cll SIOIlC ~ools. pm.. u~l'y.i~'Ol.l kl~:i\'e:l\.lind boneta:gI'l1iem, ... : si ~JU<I!tedf.,.muillb~yon c,u:av.m m~Ule, Simi!lar ... c!I:~li"'ily :ilM it" A l'l,ll!ldlili of! i(~s ,iU'l(~(~isk:s,hmrt; ili1<1gl':':~. of A II:hol~gll UXOUill;)k~ t~.. SU Ul;l ~Ind.ing ~ large SltlHe j11rs ~],U\r(l[Utlin!!:;,j 1IW!'i:rI~hlO ANSl'lD1.1 (;o~~mr}',.;~d:et1'e;~cheml!!i.. Xieng Khol~,Ulg the rJ(l,vlnce of'nOf~~U;'\"mern .~nd~.,. !->tag~e.';.ing .1 t~ge,:.;,;O!!1d L~l(l!lke)'.s •• !ud ·rme d ~~k,Lt Site ']\;vo si.I{':s~re bceo~ni n.g s:a.r~r for r~sC'!IR:h.TI~l~ ' u poss~ ble eu Itur~ll (lOmlJ:CI.km. HCIWl':YCi"" Vall. .is .%d.om~c1.W~l~II cl!JdmJs, C~:I"!l~ngf 3.fql~.ke Den Bergh.lltltes, no one 11,)$.yet pil1poibl~et1 ~ cl.~wre. large stone j.ns on cemml Sll~n\\~si h1l;sel.ntilolde,ncd Van Den Beugh Iii) hmll rQr .~] bl:.,nd i:rl ~1l.dHlmesi~l~'l:h;o !mvt "n:l~mllals Oil I]i~ pn.rlltef (Iill 1'1 .M,I[(mlild·renf!·tmli.ll~'mtl>l;lr smyey. p:re.lil~sl{]Jic ~"tltlk:lIu:ml. irnl. Xien!); K~:mm'ln~_ fiiJtltcl' afi.cld. me\~'~Yidel~l~tfj,edjar si.tCS.L'le<lf li.d:5.,'·li.ilYs .!kmwl}od. 8(11 the,re us IiIO oll'1er-c'ViThe <II£;,e (I r t~leJm:s. is anyon,e 'sgucss. dl':IlCiC HnkillJg dlt;: E'\"',"()sites, C~ml'C(ml fm~nd jU~I. ~tiHj~1"iOn1leoli ~hem, is. . l~~ C2i:piulfl~. VkmLal1lt.:~ ~ul1Idthe :I;nedrevfu~ .cUl:Y0.1' BeH'I.\®oo !:Jeli.eves that <lrchaOCl~osy 3iO:bli'! .Luar:l,g :lPrn.ballg are whoUy u:nexplo:red. If 4H){)O years ·old. <:lnd ibollieri'Olgment s frOlfll <!I. e for ;:u:ch,lI~o~,()mJl; ~umlll;l'Sk~hej."r I~eopk:. BlU~ 1;;()J1Jila:H~nt'- d11U!S nm e:lillucem .. lill ~mnl.~h I~w bun'u,lb 900' w I(!;lUI y~,U1\ old. T~lC bLl~k of num.::rifll ~s. ~~,,(; ~nU'lg,tdSII()s mighL .P.IiCS.um,ill.g I:~l!)nhe n1>l1k- gis~s. Iln.cj!]rs.~('mdlluc l'tJ' defy an.fl,lysis. When be~weell 25:00 .md I sao yem:s. eJ.'S o:f the SU'll'Wl::-:;i jars spoke <tn AlII:->rroble~<1bl VmlDern Ber~'h rir...~arruved .tlel'e 6 years agO' .. ![lId..AhhQlI~l ~here is noway ~() 11I1lt£lI!i)ge, he' .argues. thoo: :irn Lao$~1lligh~ ~"'l\i-e s.lte adlnuts.. "'1 didn't know Ili'IM,I~ ahm.!lt the b~en A~!SI!!rOf\esi,m e~Cn,~\re'_p~L'hOl~~~lfmn j a 11'>. BUl~herewa ~n 't !11lH:h t\l k~mv:'And there S1:m ilin tt_ l:em:r.LI Vii:tll!Zl.un.l. ,\i\!ltt.1'e I:h(i Sa Huynh cuhLlre used burial pots: Slt~lne .2000 }'I!! <Irs <\g(l!. [hit
(l

e\''eI!1i. tod:il!j' \'i],l<!ger~ ereer !i'!nall stul"! containh.g ash ·of deceased relatives :nem'~he jars. 1INES CO's work~n~ hypot ~e:s;!> i)i th,u bodieswere placed ill tllac ji:Q:I.:smnn and dry oat: .11 ritual de:~ornpo.shiHIl. gr. as V~~nDen

ing fow such dmiqg. V:;m Den Be:I';"~Jh 1}')l. ~ h's evldem thar .tle s,toely gmy jars III X icng
Kt1oi!il.,mg ~wre ~l(;;,\!,i.n ITmn loc',ll nl!Ck. \lVh¢nl

Sd:@'fW~n!:rn~!'!~leld \Virill p!€:Luyo[Jlm2]@ ~.'llflln s()lve, 'LJ NESCO snd laos ;ml~Urdtk:il' IU,( elw.e<I.voril1Jg, to keep Xh:::ng Khu!Ji31.1gS t[·c~sur~S~jlliLet.-bOl:n. for rntme' ~ll!dy mtd~o auraet more tourism revel1m~ 10 the :i.illpo[lVeri~Itt:!:dp[nv ince. V~'1l1DC~l i3el]hs teal111 has mapped l'Ile Xj.cl1Ig Klfk1liii.lilg sites and i!\'~d~)il!!g olfici 1l1~ crOlIl '.~l'u!l;jj!g,em,(!!JUpl~u] fOl"uflle Ph.~lll.orJ<lr.-;. Anew [em':e art Si~e Thsee i:s,ik(;l(;'!pilil~ mllcwttl~ ..\'liHiii:h had Imd c:ii!C:~.csfllro~nldi j.fI1I's" n~bib~ng ag'3 in~tlhem and COi.ll!pf\eSlli~:g the ~oi~ so mu'Cih ~ll<'!x some j• .!!'fi t\Llhsidl"",t Bt:::!bru UNESOC) ~m.dLaos off"i~k~~~,;

.r~

wm

.t~

LETTERS

I BOOKS

I POUCY

FORUM

EDUCATION FORUM

PEIRSPECTIVES

LETTIER .. S
A Voice o'ver'lihel S,mo,kefolr'
Aicadelmic Fre!edo'm
OfCl;lluliurnUlil t UCj bau (1[11 fLl~.ldi,nIIgfrom ~()lc!UCLTI mlltiJp.:!I~i&." (··UC balks ,Ii: ca lalPQls-'~,"icl.e:billl1 'O'.I.l tQ'ba.coa'IillOhlil!)i lo:r research, "D. Ckilm IiIl. N~",vs unhc\Voc-k. 26 Jan. .p, 447) (]llL1s,tm~jlymiseh;;J[~~dcrU;ro l(l!Jacw cOIHp<llay-fumll.ed U>ese"aKhas 3 ··c()lIec~i\'e me' ofspo:~rored research by[h~ n~.. ~~!faclllli;:P; lU1(~diz;;~[ibu~o:D.; ofll)1n1A;;COP[odut~lS ';"~<Iii iniJu!.'i-

edge" ,mcl

I~,U "~ddlres5eS,

the ·()tmcem of p\!Mk h(;ml1h .,. !ie'g~rocl ~!lrg

dg:t:lrt.tl~~'tI,okillg,,"

~:'h.aV>eot decelvedthe I}~blii,c. have n,m pro:l.nme(1 robaeee me . n und h<~velIllnt e~peTi,~nt;~dI ,U1Y CO~,'p[u,nle ilIll~rM:siolIll t he co~~~cliOiL1 i:)l orr anodY5:i;:s of my researc h data. 'Yet ne itherl HIE UNllVERSliY nor the other I(17
"~\'<l~'dil::l;is since 1995 :I:1<1\,1> ver received rhe (lPiPoULmily e to def~nd oueselves ;lI-!ai.llStthese 'Cha.rges, He liIC 13:oon;Q RI;!~n~~ j~I~'t lim!>to ~pplJ' thl;'lpr~!ficip!es; of ;tf;,Iof Jl~rn ic~e~dlom~ g:i!\'t; nU11\li'fiTJ~~:; ~L"t)~,C"::to CU31i,cs,~ l~e~lllcpLi iUI~5. lInd ~l:l,t~~nil~~ faidy if !Illy PLlbli~ d~c;t::p~iojl~mlac~l,IaUy ~~~rrui[t The}' \\'i.:!11 disecver tha.t i,hel~<l,ru evidence S~lPpOil,'t:; t~,e 2006 UC AS!iemlbly oflhe Aeademic Sem~l:eresoletion ~(1 1!SSer~ <!c,~deuni, free-

! CANNOT !lJNlOERS.1'ANlII) HY W

n~!EP!ROPQNENT5 Or

try to 5ul)pOri II pllWi~ dCJ;;;el:liion about it.:;: ]~I'Od.LjCl:;."1['ha~ :1a~emt:m. il1 S the pw.nnMe of the recent <liCt ~on item RE .. 8£Jt~f the U(' Bg.r~d of dQIi'II1a:g~i~,n::;:t ~uibllt'CO fLlit~, I:ililU. Let S il.<ll S{1~mj1dcr Thcrot\~il:t pu~ it: ; a Regel]~l>is in itself'a ck"'e~:t;i!CiiI1. Ile,~hh gainJ.~llgaim;t snn.okil'Q by a~tm:kil~~ I~e f<lumidalionoF fiiee<iol1~ (1~~)' .<1 h'IIi1,~.rU () He If of speech 1lilld iln.tll!il w~lich g1Nt.~rise to I~e iry. "Let's not squsnderths recent public scientists, :im~htcli~g ml~scll: g."J.insill !~'!efir,~~
have eompered su,coessiil!~I¥

~or ~.ab~LCt:O cOirnr~:iI1}' funds


to eouduct, a!-l ,~!<l~~d,illi the [!Jq,UiC'Jil for p.rop(Xij,lii..i gui OC....

IIiln.e:;., "the higiH;:S~CJllmlitYI:&


~(l'I1tt:1l h:l[ ennrribuuss t ~Blhe

hea~th gains, agalins:t smoking by Qtt19 cking the foundetion of freedom of speech and inqulpy, whi:ch gave! risa to 'the Igla~nsiillthe fitrst plsce.
l
jj

r~~,ce.

[)epa.rlm~m o~ BD(lilemisUy and MI!l[.L"{~I~rrMedi(ijn~. Ul1Jwli:~il.y ef Cill:irolll1liil! o.a,~~ Me.dical S(hll1a'~ I).;w;i~, (A 9S.616-1'!63~, USA., "T1l1i:~l11itQor rn({!I1lI~ ll~mJlLeled iI !Phi'~pM_Orul9-'rooded prnj@(t~1) deJt1!Iof;llliOOil1lilillii'!'e' maQ~'I!li~ Jt~nan(e tl:.'!:ilniq,lJts I U mO\l~url!~JlI.(U~~ O_!IiYllffl Il~in lioo~,

ril!llJmV!~m<l1 sc~f)n:rifir: k!'l.ow~hi~:I ormc€:ll1Imltion n~' Y~~~lide vnh;:u c in mic sclusions would rnpk~ly hydrolyze at ~OO::.C (1:1J';! - 10Iii.oun; ill. pH Ii) to formamide, \,!h~d .liIen (lJllidl(f hydlmlyzes 1.0 au]~u:)f1Ihlillld[Q r~!H[R[ ARE TWO MAtliN!li~EOIRIE5 fOIR nu 11IiI'' ·~(2). Hl;liber and W~(:in~!1'ih~USl;ir :~lL~~st ongilill ofl ~:fc em.~~lith: [h.e "piO,lmrer l'L1el:a~Ji~k l~lM Nitf"e-cY.:!l,uniclc prcc.~p:i.lm:Cl>would l'n3ve st;JbiU:i:e~jthe cyanide, bur rob~s~SOUI'CI;:°-:l of ~heory"lu 11m. V1lIC<llllic orl!gin) sad the "prebitlilC SCl~1l theory" {a cold, ~<I.~UC ofa£;i ~,),. ln CY1mide \!'~~Ikl be needed m prodlloCe a :m:,ady~ ij1mle '(,.'ml't:;:~nh'<!t n fO , Ito 0,2 M K:eN io~ ~h~j r Re'J){:d "'«-~_ydrnxy .n~d a-"~illi m::id s nn tifl.d~rpo!:\..~iblc Hadcan, Vl,l.c.~nk:Q;ri£i~N)~·.J.i 100°('.No s~~ch R: ro'bLlst sourees me Il>ml\'I'll. The pnJpoose:d 7.5 i:r.l!1'S CO gill veleanjc solucQm.!i~iom;" (27 Oct 2f~Q, 11•(30)<.., Ht~bel.' and G_ W~dl~eriihiiu~er dl,%";''Cril:iJe relbiotic p I.ions. i:s. ,Li.so iJnlP'll~~~!sub1~l~Th£J Or! cmt!5w,.'iiUJ1Jg I ord!~~,g[iy chc}!~@-~ti;;;: 1H~~:e;:ri.1! ¥ (J)I, ~)'1!1the:;,is ei'4r.~rimemi'i~!]l;,l:t ,In;: Cl~l~m)~cl ~(I moo<:I!o; ~~~~!,\g NOIFare .5uch dt:\1ttod ('{]pM;S.mcs, l'i:b'C~"~Lry; "narrow i.~(: !¥'IP bl:I:i\ve~I~, lodt!..':ii11is:iry imdl b p.revimlS,expel1i~lenls have demon~l'3~ed dml •• Y\o!c<mi!l.: eochemll!>lry.." Ikl\\'ever. no plausig Me' ,I[l:~~)I{j<gi '~lwi,[{lI'i!111~~'i! ~dr:11:Jil~laiin rich 1ISSCIL1ll)1em.ltJf rroL;but~lic al'£;<lllli<; C{Jm('~'I'I COli pomuis l.'j!I~ be ::;jfnlllesiz~d M:lOilllgII v:ari.c~y of ~he c~.~ed com.:I:ittQhlt; of O. ~.[O 0.2 M KCN111 IO[l°C. As i:Wtl!:tI IJy Sd.1W~UltZ 0). ill ~hif.l !tl'ltl;ugy ~ltru~~ fl'0I11 ~L ~t'OC1,dti;l - ~ m.r COlN~ ,~ll1i1ilool)here [see (4) and 're~erel~ce;; th~reiln.]. "eXJceed:i,ngl.y ill.nPI.'()ba~~e" case th ,11;<111 of The c(l!Jnpollnds ~e~~enl.i(,~. Huber ,)lid by I~,ud~;n:! E ..rI11'~ Ilii~l'{l~en W,IS cmwene>d ~t'I C.~iilnide 21,liId diis,'>olvcd inl tb,c t10t:iUl&, a 0.,2 .iV.1 \Vilchlcrshau~cr.!t~ \',,'(!H1l:f1 tl~dr n:h~l.hrc cy.<ln~cle $o~miOIllCOl~lt~ be pro(llQ'C~,Id.S~C'h lltlmlld.rm::e, ,m! remilrt.:<JIbly si'Lni~<lrto t~ose

Debatingl Ev~dence fOlr the!

Driigliln of l.i1e on Ei~u1h

],1B:oviomdy in tm('! "prebioric broil'!" experi mnel1its they d iSll2l.D"a,ge, They CII1!!i~l that !he ~:.ck of ~<lrfoJm,nion Hf()'!~l cyanide l]olyme'lf.izmioH)' makes the~r resds
g'ell~rn.tt:~

di~liIllClr-m!ll

earlier

experanenrs,

bLH t~.it->

.t~

c.

di lfercnee is,e"ilsHy expl1llln;e<! by the remmi.on or cyauidewi'thrvnnaldehyde. produi;ed by metal-eata ~yzed rl.:)duc ri()1l1I:L~;ICli()n~ uf for~l,U~ (~eu.lelf<l.!edi!m thi~ C<!511:l by cy~mide ~yd:roly$j$ .1itlildlih~dire(;t hyd:mt.ioml ofCQ). As for their experiment :~4, wi'l,elJe.h~llley
Gk~ilill
dun IlJQpro~uct~
S>l~CI1:!JI

\ve:tI;: d·~lec~~di. 'I1'e \

sr uspeet that ifthey ~'aad mid lly<imlyzedl!he


products \.... oLciM h1~"'l~ F~~<II~.y,the Ires !J lts repone;d boy Hub~r ~fidW 3.chle;'n;11iil!lt\~[,Ifill t'ii"~iIy ~.ccoml'll~()d!}l:cd w~th inl the l:riiJUewolik of all l~pd.,te'" prehiot:vc sm'-I) he~el'O:lrn])h~~ ~heoliY iIli\\~li ell p)'1'i'le <lind ol:ilI,er metal sulf~!(h~1lU'~ reCtlSIilD~~dl J.'I!!-J,m ' ij,~l]:mnl![ntS!tULrt·~ ' o:f ~h~cl.rOil'li!: !.h~redJlH!t~U:tl orgt'lni: I"t1, of:
lbee:hl identified

f~IiI[!~I. olution, s

j.n.

compCll~nds (6). h~ ~ lH'::11 a m.,ooel m~~e~ll~

WW'!IV,.sci encemag. org

5C~ENC!E VO~ 315

1 6 FEBtR:UARY2007

LETlERS
SUl:i3(:i:S bave lite potential m select, eoucersmne..<lnd org,mi ze l~,eselmm~ecule$ , 7).
JH'i'RE'b" L i1Uli.DA,,~· BRUCE 1~[GlE'I']R,. ~. SrANH'i,i" '!!.. MIL~EIR..~p!Nlm~lm LAZCA'NIO} . It JAMES CLE,A1IIES,I IROBEIR.T M, IM,!li.lEN,~ ]1(j!1hi N (Jil\!dMEIR:S~ c:lc'fi ni,[C!or~giln b~'iilil1.lQC3nl'y I.h: c.<Iubo.UI fi)(.~rien within <!~ol.. vok.miq,;HDw ~L~C(lW(!ci\"'ilim I:mrnsitioll. me~<ll c<[tallY'St:s ~I), The '·,lil.'eb'iol.k ~}UP thmry" {,:I!,~ilns pn;o'tr.'l{:~\,'i& !lIIel;;hO!!!liM:i·, C1'lUy obscure s~~f~{)rgJmiLm.~on in a cold, seemte il@Jlorc ll~:is dLlffcl'Cllct', They r~)pe'3no \1\IIJ:rk ftoom ~he exnenenee of' ilile~~(JlISprebi'otic ~~oupexperiments w~th dissolved tree cY.l!iiide .• ~\'h~d~ did!!!ot y.i~ldpJ{d ..O!;tsmlJes..

,I

is puillJf~es.>;. rS<!ipp~ ,111l!~liuniDm ~! O'~~~I!t~ri!:pIi!)'. ta ]a~Ul~,(A 920i9'l-<0!2::U,USA,~[)e~~r!m~flIOf ~i!TUI ~~d ~lal~~la!)' j'e,UK The experimeets un(Je,y·d:i\li;lIss.uoU1)h,I:W;: \Me used 1b;.,i!{)DI(T,lhl'r:t will). and \\'e Sciem~. I'La.rnl!'talYChmbtlY Ubo:ra~oflf. Wa!hlng;t~m bcc:n d~,~u~ed U) ~t;S'l ~h~ p.nolllocrorg1ln isl'llil discussed at kngllh th£ltsuch CO prr:eSStUreDs in n Unr..~r~Et¥,St. ILou~$, MO 6:3130, iJSA_)D~P:liF!~t cf .lIIeory. and all experisneutalparsmeiers .~.we agreememw id~ rhe volcanic senh~sor tile Oh~iII\l~lrirln~: Ilhdemistr,-, ~,linL..E't$ir.)i oS Calfi~lIfnia\ am a . S Diego, . ...a l~II~, (11. 9!iCI}93:...oS,M,.USA. ~~.a(~1i~d cl'~ lP;u()n~(:liJ' orgO;lrl~.t'inIil liu;mry •. 11l. nrher HU1S. we OO~I] c;hoS€:UiIwi'thin ll~i~ 1~1mlli;"\'ork_lUn:: eritiC:i~mia~, U r;J1'\M, t!lJ!xi:to· II).F, 0" 5;1lO. Mfl(iiOO. ~('iI'liltgi~' cism preSf'bl~edby 3ildrllt'ud,is made l"ro:in1:~'aC I:lsed .110r 75 '~I:' CO to shorten I'he re,H~liQn o Illmtilltijoo, Wasllr.n!ltcm.Ol2'oC!15-i305, U5f).,
~(!fl:lm~m.c:l1S L A. 'N_ .fII;liTw.;Il1z, Moriae: Or.gook(l~mijl'\I')', L iI(_ ~n C~im!'l\ll.1t I11tWJM. (d$, ~Ell!lnel;,bIKda.m'.19iS:U ~p. ~3Q<. 2. !;~Ul)oiitaw3 e!oL, 019. lifef.~ 1'Iimphrn! ~l!'. 95 1

pn~.nili're oeean, in\"'~lich organic cmnpounds m.'(;,ll,niubJlcd over 'tholl.s~:ifld!S (lltiJIIlUllio.UilS .of

Dhe.cyall,idc eoucentraticu .il~\~lI~cr WiLS sufflci'e:lltly higl~ (.):). Therefore, this criticism

r200~. ], l. ScN-ef~~;, 8. f'E9t~f,'!r~~ 1#6, .il62 ~i!OOn ~, $.",1y~~1I\'!i! etl.'ll,. Ptrx. WaH,MQtt. ~i. U,s, .... '1111'.
lA>Li<~ (;l:002). 5, l. p. Foerri~ t"lcL, J. .Mol,. [WoIliU. 29.3: U<nB), 6,. ]. Uliio!la.. A, LlOC.!t1I!. St?rna> 2!lG.. l'J.Rl (2002). 1. A:, ~ ~~~n,;4m, ,M!noel~!_~)1, Ins ~OM)'

of the prcbiosie SQLIP d~eofy. ~: .<Igreem.CI1lI, ':-" ill~ th.e pioneer '(}J'gmn ism n •.l~e()ry, \1T.'e ~u~f,!lcl: ~}t 011.' :f~2' nn c."r,'llyt~t; N purposes, Tbese uansirion nm!ctil~$otm C1Iif u;emely stable 'cy,m() cemplexes, ",\'hi,:h are simi l<lnC! those found iIII YO lcanl c field s:ttcdi~~ {Jj" Th~,~ ~!i!~ th;npI.u;;t:i.qIUy ,LUI;ym:l;~t;:: m ions become fixed as cyano Ij~9jndl'i,w.ith, [~c:: el1ec:1 that IllI;! com.:el1l~"lliol'l of dis501 ved free
PLlr!;[pt':Ctl:'I,I(;

dime. It is. 11 wdll-c,sl:2IhlisJlmd prac rlee eo exped~lereacrlens 'by illlcre,.;,:;~ I1lg a pammeter such
OL'\.pil!~S'i11 L~_ Then::!i:)r~.lh~ {.'ldlit:;il!>ml

uf

Ol;l!U: U~

of7.5 bar 00 is pOif1itl.e&~.\l¥c IfWI~ Ihal Q!.!Jtr use of ~bar CO was 11i~1'cri'~ iC~.7£t~. Th.ere:tln-e. the <tIDO\fCtwo po:un •s o:f geedil~mi~a ~ (,':r~t!i~~~~l~ 'b(}l~,l~[ '.!~1I!'''!Jl1l''1M~ ~{l doubt 01'1 the abi ~it]' 0 f OL! rreaetioas ~!) h1WC ta~en.I)llIce w:i~hin .".volcanjc, hy<lrotllielClm'~
Ilow ~.y."lErnof.e~~rly GIrttlit.

cymtid~
~(l'W

~mll'i n til'iL'i \\i!lleT i

ph;;t1>~ is CN:tr~Ii'i~~y

dli.lle~{l the high 5l1!bili~y of the '(",l'ano oo.[i)lpl~xc:>., h i~,.mJ w~~.I-~llt11b1il>~.~dl of :r~H~1 IRespOIIIIS!13i [HERE tiRE TWO MUTUAU Y IEX(LU$I 'VIE IEORgE5 (""Oordil!'lm:r:on chel1lli)1I'~)'thOll: cY·<lllide ions and TH i,g<!JuJs ll,ule~IL~(~un€;II~~ ~I}Id~fle[ellr em l~ e or~gJi 0 nife. TIle "U,iOilcer (ij'w'mi :s!Iil cy.mo ~ ~ ehcmleal propcrtilcs. [hld8i m t~t .I~Q\ve,rer. .. ~he(Jit;l' dOlil!'ll:>'l mmlllelil.tmy .. moch,l!lititic,II~j'

We uote d~~ curcritics diUerHtlm US,~~~(l '!h~ r~m::of reaet i,t)U ~~mducis\i,,'~l'iflVC fO'iiil.J. From tllel point of view of the pioueer
{ltg<!IfI~.Sltlllin~t)ry. $ib~ m~r nlacli~1:!. procll.~It~I.~. '''Iot;!'

e.g •., u-hiyd!w70:)" ileitis

()I'

OHftm.iIIl.O acid~ I[or

~et, ,639.000! tmeeliilll,g~pam'.the GRoce rilQwmm ruTloliI& f.I1e li!o Ii'lt&est ~ocm e~,tm'!l III A!l'I1!f:i'fea.llor;a.t:ed l'il·!ht:t !heart 'Of dO'Wl1~owll1i HOl£rom:mcl i~'!he mfi:ils:~
S(j[llIIill'e

A:t 1;8-5 ImiD[()n &if'O$$ ~


Ice~
(jOll.t(glJIOtl~

Cia,nv,e:ntion 'Cent=er

IGeorge R:. Brown

Qr 'r.l~rnI!'1!'1l'!em:,

dirrl~ arrd

l"'!iI!!!~~

* GRB

116FEBRUARlf 2007

VOL. 31S

SCIIENCIE

'!IiIWw,sciefllcemag.org

LETTERS
~)ell!ides .. r~s~nr~pherelrom(4. jill, 'l'l exh~bil· iH1gp()siliv~ lIu~nl~~iI;:JJ IYI.ucri.:l~,j'b~lCk in situ by 'pl'ovidi't:lg~n,'alll~ilion Ill,enll I iigalldisfo.(J' Ilig;md 2t~cch;rn~!eclc!tc,fl~ysis.,(6) of earhen f~xmion 'p,nh'W'IY~. \\~lich com:t~t~ltese">'OlvOible rel~m· d~lt:limli {n R(;~H;li!onl1,.rod~,ct1;lmml spill mJlt illlO vasr expanses of the eeeaa .'lose alii.che.ln·
o

:!IhI~lns~rOlIl:~ D·80'3tB Milrucm(ltll.(i@l"l'IlJlf!(. ,~Ilid 20~ Mill 8.,

R,m: Oriyt. C~iiI,pel H~lt~C 27:51<1" l!JSJ!.~()\tliParlrnl!l'Il (Df Orgallif {BI!l!ll'Ii~tfli' iI.~d l81!Kfil!mig:fl\'. T~hI rni!J(nl! I.Iffiit'~I'li,iUit Montfillli'l,' Ucht(!rlb~r~~tr~8~ 41. o-aS'N:;r fj,.m::l:ii rug,. Gll!J11iilny. It.e'lereIlll1ll!l:5,
1" Gi,wa~ilius.e(. ~Zl!W!ll.
f'fr~,

(Wi+D) Imel':;ovef!llrne!Tiwl WO'rki!lig GroUI) tKiWCi~ 0111 Pubiic H!J!l!~I'tlI!.~!1t1mI3[i{Hn. arid ·~ntel[ll;.·ctuil~ Proper~.y. fdlm"",ed by theseetence "{The drug (}o.wlj)m~iC'sElII.d1 ~h.c.l:::umpc.:rLn Conm~is.~ionopposed the Illo"n,)i".,
In fiLCl. the IEUfil:P~1~lli Conjmissieu Wi~!>,,"~ stro11g propo.n.el1lt ofWIJA R.e:.ohuioIlS',U:4iI, ,I [1.1 S I M.:I}i 1> World [-Ie .. ltll Asse;lrulb Jy.. The COnlilltl'i ssien \\'mked \Ii.'i~hh.e ElQl'ope~.n l tJniml fEU) 1:0 Sluppmt lJrleresfllHtuon.

1l;om, :R. 5".0" B ]61, 17!lJ.l

icul pm~lt'i,ll

lose Ftl.r !he origin oFlIi1.e pioneer org.mismn ~n


Frnarrhe poi
IlIt

by di ~lLti()n;Ind are ir!l:e\'e'I,silQ\~y

of view inf (I:I~ prsbi o~UC

'solll) [I~eory_ our ar..itics see ~1urreactionp:ro(~. uets ~15f;,"atering!l~e pri!l!iit~i\'1;l ocean 10 become addiluull'I111 ililgredicms {Jftht'! prebi(J~i.c SOti(p. \"~l\;;n::;inaI1e:~' ~{)nu;:~~l(JUs.<'Lnd or m:ul i~{lJlY~Wl\.., .t'!,Ild ullder .1I~1 anaer of'diverse inflacnces, m lhe- magic af ::;.elr~of':~.m ization i.~ believed to
him:; MJmc:l~tlW' gcJJermed ~m.ItIl1:;pL'>Cifi'l:~.fit.il

~.M, MI.ip:N_1I,oi!~Ilil\t!!S,l" SilIo!1?:1'41. " 3. J, ~ ftl~,W,]. !'!ilS.I!'!]'"r.i!r~,~(J<, lO~ U9!!4~. 'I. C,H~~~r~ G.'i'I'l!idmrH,-!Yer.S~na;Z&1, '6~1O tl9'9a~, S. C,Hti~~r; iN, IElltNticli\, 's.. IifMI, G.iIla(htfUllIiiJ~er:
5~dM.. 'l',;;~.~OOJJ 6. Ill.JI. 1!!~1lil1~, C 1I01m, t:. 8. Slb.;llJlrel'~,An:!l~ (~er1i\

z,

.trot fd. ~:t1,l.~~9 9!j5~. 0


1. (i.. dhtflf:lhiji~!!?l, ~
fill)I;.

Notl.A~.Sd.,

,I)SA 817,.:):00 .

ta:5'aBl~

formonire. lltoC'f\'Iif)theories ~~re{;a!,cgori,t~~lljIdiffercm fir(Hn the~rs.peCI.ive gf e..'Ij,I"~C'rimellt<l1 sejeuee, The rrebi~)ti[; SOI!llP theory i}; reM[i{.1~ to the ~es~il1igor individual aspects ():fa ~ong. p:rotlr,tctl;;dm'~~:ol!Urocess (3'). The p~uneerm·~· p lIlism. by contrast is e;.;:pec~ed to be e'xpe~:.i.· .mI'ienl~IJ.ly reali ;Glb[1:l ili1 I){lto t J p. G,I)NTEJRWMHiI!ElR511i!1AUSElRjl ANID (U.umA I'Iliill'BIER~

.A Cllalrificatiion Ion 0110b,811 Ac'c;le~ss tOI Dru gs


MARTIN I~NlS:~RI!Ntr5 MTIICLc"WHO P:AINH \l,i~ig:h:s. radical id\;l~,,:s"(N~"\l,;:j oflh~ \li;{C:':C:':k. ~ Dec, 2006 ..p, I J 73) IflJiSrep~"Ilb; du: posil ioUil
ufth4i: ~nupotl'l.fI Comln~&~iolil~ntl'i~ ilnpo:r:imlit debate OUil loba~ access to l1lediicim:s. g

~ndtlding the ,eslllMish,l'llcllt of the \;Vi-IO ](iWG, The Commission was n::pL'e'Stn:l!:e,u. a~ lheWI-i{) H::iWGil~ December by offi,cia~s ~mnilrive dil'f:~rellit Direcrorare Gell:er;J~,,, who '~i\Ji1k!edclosely vd.·~hEll Mc:mbcr S~m.(s <!tthe meeti~.lg,md aClive~y illl~en~ned to suppert the prul"eSS. The' Iltlree stmed "rior,i,liesol'th~ E~~mpeall tJUlil)ll~1I. [h.c ]{.iWG were (i ~ 10 promme research and de\fli!lopmebl~ fm:tlsea. on producrs for ~lut;ml~l!l:i. th:Ud~:,~pwportiml~'idy

a ffeet d~liOpil"ll!.oo'llnit1'ies; ~ili, to ensure liilat these P~'\)dllclSm'e affordable ol'lldiaceessible w hhin, UiI Jlic!H<lI.1health sy SleJHS; a'iad [iii I to ~liJs~UD'l:l l!~~ I;;{l~ulilt[it>s 'L;lnl use the ~lexiilil~ i'th~l~ hi [11k;fi\ril!lpt'I.mgm~~."i.thItRl ~~,~ i rd~mnc~ to the Woiild He'II.~hA~ciinbl)l' (WI-IA) reoo:~H- t lesprovlded i III inlCrl1 mi'Oll~1 le.fJilII. sgreetion e~t<la';~i:shil1~ ~\W(~dd He'Ollth {)rgm~i,7,:;H~Obl mears un il!i~e~ lectll"~ pmJ)eft)' r.~gh~:i.

R:e~iaf1l,t

[Park

Reliant Park, wEtl\i

~ts; 2.1

m~lioo, g:I"I):SS

:sq,uare feet of exhibi:t Olillcilmeeolll& :spIKe' is, !he largest s,po:rts,li:rnertainmern.


C~~~"1iI anI:! nded'l.mvcl:lmpll:lx in me'l\l'Orld. IRelSmt IPawkloo!1ses <I major faciliQ!l~ R;,eli"liIt CM't@!' (T06.2.1!) g:sf'

of $inN.1!!1r;r,\Clc:Olll~o;ui!lOO1ibi[

s:p<t(lGl)

~1~t

RetlMrtA$!if'lOd'Qm~.

St:!~i\lmi Re!i<lfltMOO :and!

'WW'!I\I,.sci encemag. org

5C~E NC!E

VO~ 315

1 6 FE BtR:U.A.RY2007

LETlERS
The lEuropc~m CQ:t'nltnliss i,ou :flmcls research ~!lIto Ile!.gloc~ed seases ,11ff1:edt ing (Lte"''e~opi~,g di COl1l~~J'li,es, aetsas J partner ill, ell nlcal I'ril,~~ s, and Ul'i1e~~~~V'~~O~I~~l~llt limdin:g and pulley f re ~.mp.l'Qvc global access (0 mu!diculIIcs. A Comnl~SlSi!(1Jlle,gislali"'e proposaljor c!!Jmlml~ SOI)'~il::Elbing \.\'l1!'l!Ut~opt~cI '.he El~mrt'!mi' by P.;rdi,ummt,und C~~lmcil e.!rl!ier ,~i,~ }'~r l n.,;;g~J~~uion& 16l20061, The EIJ also fll Hy respects the riglnl of~n.d Illig pautne~'S to use co:mllll.l.Ml!ry 1i.GC:lllsinguti "H lJij3,te:r.d rr:e~,

projec~s 1.00 ~ iJllb~liiQiIS. liar.peertevlewers to Ihcw~)rk et at: ii1aspl~bl ilshM ~ll~bc I. csn jind el al. ')i, contaet infmmnmioll Ll),i re~,m:~i!1!~ I:~e Ilmd!'! ~.did notice ~hill ~'f((I, ·j.;ml!!lw:r ~i~es<Lre IX'ltent am] IrJdem<1lrk offioe.lIfo my surprise, .~ !;.rener.dly feurlll or Si.XI~.. ~s !he~ack of nrs~ :fi nd ~m ,,,r (d, ~iMed O~. ;a~~)'pl!t(;llil~N. ,Of IT'olde- Ol~dlO[~hip~h~ re~M}Jlr:1' (fl. ciLo~!\ [llfl:l~eU\'~ n gO\'Cil'fl:l'IlcnJt-.spon",o.~... fitm:lhlg'! H is .il.ot that m 'tI marks. [ gu,c.'iS, L'I f:.lt" \V;aIlJl:S ~U thewerk to remaiM il'llJle ptlbjic demain, HOlM] know] (;1/. ]S an aU~Qr ill aJ 1~llose p<!!pers o:n]y beesuse raust 1~1Hie:cl cd. 3::'lny I.E'Xl rn~i1HJ;r; \,-elll :ifm ~I ~ nf !>lIpr1yirng k~y ru<ig;(';·m~. illlvtl~'l/Clt:ll:'mn ill (If
dl:~t,nloe.M~be l C';;;l!1lfilll:l ~I ol',~ {"X}mt'LCtinfnr~
11TIIm.llO(;l em 2I~

.pi:lli:em
l'IS,

COlle.

The d!;;ws nW,UH~ml1lW'i,c .Hu~}lnrship

Nlll,spOJ.l~ooodl 'l'CSC'.'!rd'DprojOC'[. Am'lzingiy. NIH has l1.olli.mded projeot:5 tlIuder ,~'1' t. SUiI'I!tI£. El (II. 1t",1!;; more~il~~:1 UU}DO prub.l j. a trndle agJI'ce:l:nents. IIINlIIlRZ EjIIIitY:5 oo~io.iJls0111 a :myri!fld of ll:Ip~cs.so \:I,~1IY~s ahm 11m Hiilhb atl1idC:a,11i5~mer "'r~Docti!)n !!)i~,!!l'('Jl!:rale-Genera~, ~Jl\pre:sshtu{Of S~lffi(;i."'I~d'y imli:ru~s~iptij,l;luy)i EurUil'il'i!1l(Qml!Tli&~ioo.IB-·1049IBru~~,e!J~ IBd!Jwln. en{llug:l.ltoreceiee fimd.ing'l ,Does (11 a/" Iac:~ :f~:lG'u!'lin lhe~'~ nr~r ~"\'iewr.'ffi1 A~ fi:l .rd.N

the rewsnl for p:nwiding key COntlpnIilC~~JtS U) a IHoj:ecl aee over; ~mch individuals are' I'l()\V f."{]J1S!gH~d to th~ .w:ikrnnwll;)dg~.ll;lln 5e(:tim~ .• I: e-mafled mny fbnlC!i' pr'OfC&"OTS ~o soc ifthcy ~m}\.ver al., but S/[) I~n, ~.have mu received fa ~~'iply,Does ,mym~.e know \\'!'here W fill1ld et ai, 'l
IlUCIHARD MCDONALD GeflO\1<1r B&o~iemce IUt<!i, ,296~ Cillifornii!, s.~~:t. St m~ir Sha:f!l11,~I 4:BOftO, I!J£A, n)~R.EUIIOI'!IJIS ,AN10 (tAR] F IICAJIONS

HAt.

journal articles dI::UQ10fl!S! w.:uE,hlg l(;'(:!lIlic;'l1 and research abil ~l:i:esthlllJ cross n1."l~y$cienti riot;
fid.d& B ~J:(]mt:s \yml
~:iJ1A~DSI1l:.~lrci'l,

's WORK

1'5:R[IFIEREN'CED IN, ,I)IVIERSE

tetters to the EcHtor


letters (-3-00 words)
in
S(i~rI(~ 111

Olll:UH

thi;' prl;'vinLI} r:.

can-

Q ~l1e fdl ml~r~!i!. T'lf!'~ «HI


!TId

rnetenel po bl.t~h~ I'lc!llh~ or I~W~~Qf b,fr su bnrin ed lhra uq h


or by rE;'g.lll~r

~r d'i:;;c(lver~~s.di!ll}~~~"gl;lO~\lgy. 1!hropoplm ogy, flsi,rop.~ly$i)cs,.i'uldl even i.I'I researeh 0(;1 :m \\;OO:I'I'n called C. e/('gmJs. How does er at: do if? 1 cu~~tl:$'i~fi~ I .frmillh~w~p1g etal.;l:S n~yrlexl m~ll:i('jf. tu!.who is, el (It.'J Ife!' l~l. ilt;,} j.m:;fu.,>SOii." B
a[ a. m1Jii,l:p.>ity. \Nh~lt dtl~(" (JI:[.Mch'? \V~~h.1UI~

th~ 1!'~b (\',w\,q~lhml'l~~II"nl",.O:i:JJ

Il {12(JO Ne IY VllIk Ave., N \iiJ, Wd~h'l'~9lml,

[Joe

2(1005, USA).lf!ller~ Jr~ not J(~~IlQwl~c:~1!d upon re eelPi. 'l'l)r a ~ .J uth a rs gCIl orally CO!'lS ultcd ~'I 0 ~ publicanon. Wfl~ther puollsned in 1dl or in DOIIrt, Il?t~er~~re ~ut;lj!e(t to e-c!1'lng 'or clarity ,lM ~PiJ('~.

_cr~l:·ULtr.lf.v;,tiloo.il so'l~lIi~g illbi~mll!th: ~p~~nglll soDE.rlI's iiill~r<IIl'1lfli'lk Ji'D'OOnlilltwlllil :Nir!(!i"1Iy I), I'll Fnilz ,i!l dI'. f2 ~e~" p. 633,~,tillhe ,"~~Il!JW~d!Jf.1em~ (l7). ooe-o~ nm~rne[lInditmg grou~ was rnistdemificd. fIJliSH~!andil: ior ·u~:e~tanditl1i!l F~~t~~Ii!~~uO!.ed SI~!!.Ic'~~'.!!1 ()fn1!!'IgE's:.'. Iil:~ons: ·GklILMI-!l(i}l~ sl~liJl~rllil!sIII i11lragl!iiil fek!>as~ Ipdt'~m~dll!lrimglomg~rm d~(lII'riliO$m!llfiJ· b:rw. ~~J!gm ell'l. U 9 jan" p,~61). WiUiilm p.~nol1i<lnd Wfrumdl!@L Sil~ r $hWldi lfia~' lbe~!i' li>te>dil'5 ~o-!le~d<i~nUjl'5,

I'j]

• •• Cambridge HealthtedJ Imtitute"!


1111

'Conference
1. IT Infrastlruottl re
2. D f,scover~ InfOFmaUcs

,f:1'

CONCJURRENT TRACKJi; 5. ,e·(mntiaa~AI(!'se;~tioh '& .. TrarnshIJHo:na~ Medliiciin e

3. Compmati anal Bi~otogy Be Data Ana lysis

IE: CO lOQ,y

B1H~IOCl!].e:-l aUld s.colfn desCTill~ig!l~ of spec ies,

The

re~WI~I.ing dliscn\lC:':ri~:; ~.n1It:.~IHcl.C 2:0 ~:;Wer


l!!l~n(l'\'\"iJ

pre" iou~l)'

e::d:iaet .'l~cr~~ new,'

<tn::hmmlo',gical ~UI~S on :P1I.~

P
I

[]I.eo.rntolog~sts s~lldi~i- '


ing ;nl~:11I1rLI bones ex•

lb:ti rll::t EIGI'1dsIlIf :N~w Zecd alnd b.llA.I.!Il'Ilenllr,llollJ'l811,11 Paul M.I'J'lilrsolll


T8 Papa ffe~s, Wallilllglon,

111 EXW,!CUfHl

(,ifill

Bio-

111:.'Um;d:and d.cst:riooill d!olZCI1S of others recogrtized but mH yet named or d'e!'ioCribe',d, nd a Smlillii brg.~ gob'tl.g;T<!P~lic .raflg~ (\x~lln,~to:n~, artl B species diversity~nd endem iSH],prove to ~uli\l,e
been hi~h~r. sym.pm.l"ic
IlitO:le

g,('ogtflph.l' oj Tt[Jpicfl.r Pn-

congcnerlc

species

cavated

Frome;uly

cij.k' .Bi/~h. Da\l id Stead1111 summarize f\ fO,Sli! I al1l a vufamn35 ofPnci lie islands oliiler thnn New Ze;.I!.nHJ <1~d Haw,l:i,i, His .Iccomu Is

ie<llLJen.t, end nighlless sped.es, b more r1llm~ltm:o; d~'LIlI plf{,"'I'ium;i]y rellli7..t;d., Gbn[rnlf)'
rojhe Ibrmer \fUe\!,1' oDpre~im:llls~t'iilll lnunaases t~tdi!1lg Ii,ghdy on ,~dr ~!lVirOllm~lu:;;. hum .. n

eifle .ilsb.uldlsh!li¥e idcmiried


.~mw}' IH'evi.{llusi}l un}mm\ill hil:d

species

Ihal

ev,tdemly

New Zt1!lllang, 2006,190 p"". Nl..~, 188 N 97,111]90001101 ~·B.

b(;lcmtil¢ C'-!\lUnIGl ~'l~XHil ,lfreU"

]],rlsc:;d in I"!Me
Ex] i fi,c:t~ <I illnllE! i'o,~ eOQ r<llpllilJi ~j1

p~lrr (lUIIl~i\

co.~onil:m'ion or every V!.~ell~studi,edlPacHic ~~i~lItd :k!:[{]up\va~ lili,lm\it)d by "I Mi'l\"~ ufven,¢braie C1i:th1.~ti~1i'lS.

h.uI~Ulllfllliir,i,vill. Those C:lI.lim;li(llJJ~ 11ave implicil!,io.m' ~b[ lllUETI.l.Ul h i~<;I:ory in '[he:


(;'\101 miolUllt'y

hio~.ogy., T\'I'o recere books, which CQ~n·

ri!l!~ific. bi![lgen1; Lr:<lphy.

,a,r lr-o,pij,C;O'1IPacifi,c

Biwds

own diseoveries ave:r the pa sll2, }'ell!':'i. He pern(),J,~I~y


~X,t:;I\'lnt.t1 d(lii!(ln~ (If ~,Dt(;i::;'. r'~ri~~~ hun~:red!i, of ltao~-

AU theM: discoveries
!lga:~IfI~,l

represent new d"n~


cond.HS!i.nric.':i ;;;L!I1d CmlllI~L1-

~1JiI,d:

.lib",[l'o~id W SiI'[!!lormw PH] SiS, [:hi COl (), 20OS. 608 P p. $110, £691.50. O ISa-N 97800llilnlt11D, F'lIpar, $45. £2UD, IS8N' 97a02:1!!i7714~l
~,'l;!l[l

v"J~it:h~o h;-.o,;l pr~viom ,Ilxn~~ hioge<l!;'Foirny. e1o;lincliml,

u n iv~ r:s.iiV·0'1 C·hi

plt~:L'I1~nt ~1reh oih~r MillOpared aneeumed skeletons graph iC<I Hy <tint! by chance appeared :;,j u.niJiu n.(;'\!)~lsly. of i,niJl.l:rn l>pc-ciL:s liu[ CU3& p<II,',is!iJn. Thi s mommlle nta I sununarize cu~,'enl knO'\\i~" if;.:d.~!1f ~~tig'ii:"~binl~ 011 :JH r,~Lcifici,~l~lnd [~!i,"';;J:IDChp:tngnlrnwln:~ "iI:;4:;Q!ili,plish~dirrlt)~II~y groups c,w.ept lJ.law<lin" silllg1e"h<jjj,Ulded~.y.2i.g.QiiIlISIlltJSHliCks ~1mobtainAhl!'l Tennyson m'ldPt!u I M.tl"linso:n·~ i [lg perrmts, find i~~. sv!e~. en Iisti n~ eeeperat ion of loc.alpeop,k. <lind,o!bta ini'~J:!:f~,mcling £J!.'lifu't Bil'r!:~ Of.t¥~r:MZeahmd add;!): 10 Trever frOL~} !!,l.n!I.lI!lag.~nci¢~ ~11,1l,dn'l~re Krud~,e!i,td' . , \liioflhy ~lI!1d Rid~~lI"cl.Hn~d",,~'[~''[.; (:cmr]ie;r[H)ok ,t J l' by W.u!!:nlHil1lg New Zealand's 5E,

sands of DOfiIC&, measured the bones of (WeI' 25,1000 ~~ldiv~d:nal b~f'd~.~illJ pre-

nity ecology. T.~lroli.ghO'L1l [~C book, Si."i1.dm<ll.l criticizes conclasioas reached by nrevwtl:S 11 ~It~Hl[~ q e ~,p(:lC'i.1111y Erns t M ny r.Ro ber[ ll. MacAn.,th,lIr. and Ed\!''ilnl 0. Wilsofll ~1UJ worked bl:fo:rrt: L~c inil~d On'n~tinilnH.:a!ils!;;cl e",tunciiol1l in Oeeanla ~IO'IS recegniaed, s.tead~
li'I,,~n !;:()liichd;s t~'i.atD~nrm:'l ":ruvoIUli!l;lillD7.e-S avian bi~ooglill)h(>' (I'll Pacifi:t: ishmds.." Oll1e

mish~ !~'!ererore el'O.pec.~ book ~()he ftd ~of the


~I.'IJI ~er in~~rpre~ut

extinet

bird

species,

A~tDlolLghlhe

l:noM

<m~Jy"ses the ne,.,1 data, demDfl.~tr<llj~,~ of hCi\M im:l~ !,!I~ ~:h~n;b'y '11N;ln.'icl,. Sadly. suehenalyses 3f'C IHI~ ol'l'en~d For mnst .of
dl.'L:' obvious ItILl~I.io!Us... .an:dl.

~'::!:l1mjtl5 (Jr~E:;'1.:l ~U1:1.1l!~gjatu

Il~~lll~"l\ moas,

~~u dy <lS r~n,u1,;~[b~;e the "\'(I!rl.d:~h.u·ge...' \\"~re' cagt!.! (ilIll ttl i 6 kg) 31ilJ ill> :sam:llh:stmgtuless hird (22 g). MO!ILIiI~nson"sgmgeous~.y detailed! p,~il1 ~u:l1gs ~)ri~]gIiiD]ll~ l~JLO [mg~.c~(J&'" nf fur· rner~y hre~l~~ling .•mima~~ ina w,ny th.u re<!~

o!l1ly crude

~~fI,I~Y~;!;L'l

are

OIlliCifiC-d [urotl~~~}ldto~gh dle book oflell provides enough. ,Ui!l<!. ILlI~[ minelll dn the .Ui!.li:s....;ing c,,~h;:aI!liiltio!1s.
FOIi

dcs,{,'t.uptiom ef'benes call1lOt :!lchimlC. 1l11e NCI~1 Z~,;mbmd fassil a\',ilfal1~a,~sby rar
in tlre \.\'i[]i[kl: known IlI'Olill bones ofl~lUl1Iclmds or 1l1mlS1luu.b .of ililtli'it~dn· als, and almost <Illrnodern species h.we been U"t}CfWlOllted ~lS fossihi, Alnollig lilepoiIu~. made in TCJllilysmn 's I(!;X'[ and b~Wonh.:y 2111 d . ~~~(l~dawa}' :i:;; th.ai the New ,Ze"llmu! extineU} 'ii! '~ion$were Ibllmvecl byl'he fO'LJ~dilflg ofNe\'!,i ~ Ze,lhu1~ IXlpW!~,,!i~ln~(l~' <J:~le"s, 16 A(I~tr'liluml ~ species :I.ntesponse to tnLIIWllnl'3 uad~I.Q0d, :lrIflbimi I:l chauges and ro cl:ls1Il)pearall'ces of related f(lrJ1Iil~[ res~~~l1ln~ ~rt\(,:i.(:ls. S!illll'U Australian ~ S\,\r;lmp.l!!emi• ~QI$. ;_lfIl,clll<l!~Tiel'li-:relJl'!~(;t:d ex~ ~jlll.ct giilnlll[1o:~gcl~crit':!>pt.."t::jcl>~ 11Cth,,!£,t!I:I'-cgc. .;;o;~<II.i;mslaek~m:kspread inland after the extine~ lill;)!1 of rhe ""~!ile't1~ri<l.1liFiru~;c;h~li l;llC'k; ~u~d d ~.A~~$U'l:1 i~l1Iptedl~1 ~It:> I~f,o:~~eret~ l'Opul~i:t tmB 3~
themosr

instance, rwo of

('O!Lnl.1~~tdy~,mpled

most 0:11lite c,.,.tim:t speciesere

M3yrs condUllt(mS,]l'i<mt P.,dllc ~sbmd~Lviilmn biflglO'tll!.l:i1phy ','ere dl~t an AusrraI ia[b~{)gcoglt.llp];)ii.c compeIlellit in the ish.mcls' mil inly Ne'O,\' Gui ut;:<!-deri 'Ii~d 1,:1\'1-

fmmas :is ;l;u"ollgJ~SI OI~ Ne\l/' Ca~ecloni,!. ,tiecl'<easlng to


tIJ'Ie east f.lnd north (:2). ;.Iml th~l i!>hn~c.l,e'mlden~iMn iIItCI't.1:l>l:S wi th :1 !l!l.i'I.!ildIf...:fl ? and Loo,!lmion (3 )1, Other res,~ m,:h~I"S~!lv~ ~ll'ld:~ldtld ...

~I!:<l!~h~ body !'l'I<'~"i~" ,nf.m i£~m~J1;, ~l)P :1!lcroivofi.:: iif'ld c,n:nh'ore iucreasew~ll~ is~,nid..!fiI..",~a.lid pffi.:.'i<ulllywi~h
is 11l11ndo!;.>e 3rt1d p.roduCi.ivily ~ (4, j le, SI~~u:!D'i'I~nd,U~tis~~.. nOlle of I'here po:in~s, (My pmlimimu)' iml)~~sion is

I~

'1.11

:i
~
ill

N...:",,)Zea~liii'id black :-l~1 tTIl h:.~

~1t'boJ.

~ TIH! r<llYi~IN~r,~ '"

Lyall's rn iIh~ D~JI'iirlm(!m 01 GI!&QrJJllil~. ~CIJl, !Eiol!leilii'l by r,~!;Sl!l1l(ij1C~!'$. The ~mi1lUeMmglhtL~s Il~r~, Vitelli (Tmvet$t!:'J W:5 I\i1Ig~le~ (II 90@'II5-1524. l!I51'\,,Em.iIl!: j;diIE!lmtlnd,@ lY{JIlJ},d"'<pktro here llil ~I mJeli,~ fi'l fa~ry p'~Dns (PotliypMa irrnl!Jd, SUiviN'!ZlO ~!1 Stephens Island 1U1i'i~1I1a95. '9I!log.u:da.eilu 'WW'!I\I,.sci encemag, org

Ihilt his data cmnfi1'ln

mi!t{'i'

~il'l<movenUl:n Ih~ecOfl('lu·

5C~ENC!E VO~ 315

1 6 FEBIR:UARY 2007

941

BOOKS HAL.
!iJ,{lm;,}I'!;1l l\tkilll~O!'l 6)llnd others 111<\\'(: ( Sllg~ peoip~e i'l~ one diirc(t:(lTY shared rheirisland \vid~.~LIp{:op,lL:: ill th{l ether, ~ gL:5~l.:;di dlm su;<;m::pribai.D)I' of island b<ird~~re e:."~nllimLtion by il1tJ!'n~h,l!l~"1;1 ~!nrul,di!!1'!1 p~·t-'lcln D~_~pi'ii;!' ~.h~~ 100Ck oW ~llQiPpo!1imlil .. mLb"1i!e~. eonm.ors,dcpcnds 01] I)limc:vollll,toilill'~' C1':rrllUSMre m Slcm:bmlill (lIfers spcei ftc but h.l11pl~lmible native m<lmlnal~tm pr~d:ll:or!'>and ,l~ so to L1011l~ c~usim1S:,e.g .. ll1mt L has ltiu~ Of IlD elf«1 on SoW 1 m",rnrll,uiian prmlmfll"S ~uc~. as land cralrs ;U'ld :Pacijfit:i~h'llld~m~.l~.r111111Irnv:lI[olis; rha~ '·kn1.'lr~ Ilmd Cl'OC1lJcj [~es. SI:e;]Jd'twan provides lY:lm:hdl;n<1 island disltU1.ceS ~.ess·m:ana), ~00 km hm,\.'ehad a suilall~e focql..-mlil,a~i\l\c; eV'JI'llat~mlofjbc idea, Iwinimal. crli'lClt011 species richaess": ~l1mj.~hfll S but he does ItOl pm:sue IhemJ<l lysis beyomll i;~ "'ixui!al~y i.lldepelld:e'nt or ~s~m1d for <In)' A all <!lnecdm';!L~ example (New ~1:'t:~1Indt) arid! m;~~mld e':'a:::ee:ding some lhre;.~hnkl. 1.1 tnlil1ging some qllillultllhfC dlisells:sioUl. A:I:lmher mLlch'flll:l\IU ljo 5 kmt in EasrPo~Ynt.'Sia to 5n to i00
kJl~~ [ll.rdl!i;l!" weM). \lV:heL'e do these de;t;_~ OCln·, ~~ed dbcl;[;s:'ie,d~opjcof .,vi~m biogeogrn:phy dml Ste,u.:IIm.,l:l does 1]00t esplore Is thenade-etfs c~usions·come rron~'~ Sr~~u;hn~lml"s ~t~(~d:chrougimllt m lite b()o~ berweeu O(]M~.md benefits th!l.'ll ~e:ild lo~h~ e\,l!l'llIion ~1f:liJ,igJhlkMiilles:s island hi ms, Wi~h Lh~ UStoreplace qllflmiI18.tli.ve iUlflilysCi> with anecill} all!toors discoveries. !:ne prehi~l;rie <1Vir<XLIB1!:!, ~tb~ll e~~lp~es tlil <II he then overgeneralizes, ~~.il£Ill,.; {l\"~rge:::m;:raliz:i9.lim:l~ bt.::li£f:-., aad ~s u:l' 't~~IUl).P~C'i~:lPlltd,r~l: now eTlconiipm-;s tht: :r.1!1'1~Iy !J:ehlbfi]l the bel i~I's .1S cmu,cIU-"Dom;,rbr ~I,ge;,'it !1lJmb(;."I' onli!;lhd~~ birds ill t~~\'rI~(~ instaace. he often remarksthar ,1L {:c!n~i.ua and SI.C',\tdmflfliIi~ts~he .25 bird facmiliics, thill sa)eciesc(ml"u:ne~! today 1:0 some Nn.iclIl<l.r eomaiu ~1igb~~e:os sve>cies,Y~l he does: not disi f I on ~usswtily ni!J,UID.~~&~~be~SiJ1~V~[ !IPfiGaif~1 in ~"o ~~:nge~Iand .1rn~1~ll~e:::llOllilndl ~,:s.~fo~~~ tm~ islands of the Sill:mlle' aKhjlp<:'I~ m..m:y "theil' birdi fa,mili,es, or ,:~i~y iii evolved iln 0, more' ~1111l~.lel'

avifuusas as
q~les,iollll'i

l~llm,!Ui!II'<l1 ,1l!1d

re~~s.1mH~CInteri

prutM.hlll:'L1. [~~l'Cv3Iu~. Yeo£. 3]~1~,\vt:.rrirlg w.IlI.1~

,,~q u ilr~~ 'Lh iS~'nn (;",I~l1erl;!pif;\chve. biorasare btl i!f(tc(1 today. anti hruvc been bllffeled for bundreds of' mimon~ of )fcars, [)y mLIIlp:lt:ls other ~h:::Ul Ul!o~ {]IflllU!111nl~,
1~]ov.l'C\1'Cir,

Uv,~n.~ birds still~.eat. prey. compete, breed, dlspersc=end ChfLU.C:l'Bge Hlndcrrl>iii,un.dillg, our They I"eln.rin~he only b:L~'dS in w.~~ch\'I.'e ean stlLdyba~,rl~ etd.(!gi,~:all pl.,{lC~S5es clilrb'Ctly"
il:er~m!lei ]. 1. iI.II'ICUll!y;.ltN,
IHOId8l\\Ii)'. f!'!e~O!lrl~l'fdoJiliiil'MlM: f'l<ehf:srarf, ~,~ QI He>I!.$1[ORd Un{lialil;l Ulil~~_~~, 1~l.m;lIinin.;Jlan,~o.n; ~Iliewe>!lllllr _IO'II. D Stea~m;ln. 5.ITti1rt 2:99, 2136 t2'Wtl, IE.~P', 1'lo.t:. ~j\!i' p,ui{iC5li" C~, 193'94. 1~7 {1'M'1}, IE,~~,Sck'/I{)~ ~~I j,'58.7(];9.65), IP,A.MarQue1; 1!I.L Toper, EvaJ,~"tJl. It t~,~ {199S~. 6, "" Brurn~d a~" Prot Nall.l1mri So, US,A. '!IB, U!i18 (21l11U}, II.. ~~ [. IIAirI$('<1),ill (tMSI'J'IoTl!/(m ~tJ~lar1il8.irm.. ~~j. MM!t'~ Ed, '~lnta~nlllloiML ,U!iUJi'l(~,r.tIt aird fW!'llt!r\llIiikl!'l, (~tJrid~~,.l'!1aSl, L'lp,~i5-n L O!!iflll!!ll,S- Piml!l, :s!w, ,I!(.i~~~~,~S

2. 3. ~. :5_ li.

,;"

~>oon.

~w:m~ Ptldficrig\::)[Jfi!>~lnd 11iiI(:9~~podlL:$, ou,~y three P:'<u: sOl1g'b:irC~s. and many gmuadil'r~c dwdlmJTi~r.!~~~~t ~m gm~lnd-d\l<UUimg qllll.,m l, The beck s l'IlOS~.g~,uing. oIDI1L.".,ion.~are :)~oti sneu ~ oHm!)!ses of' i$~a~,d,<;peC~CSI~m~lJen; l S .md~heir ooJn~I')~by <II:e", .... <ilist,ln,"e D. alild t el'~V'at~ol'! L-d~spitE; long dliSCI,JI~s~orts of these :llubjcc(s. For all Pacifit: :~shamls other th,rn New Ze.d.miC.!. sl1n<ll~ sample sizes me"'1 rbat thei I' fos!>i I ~'J;viif:lLlIn3::;' an; i~lIllompI~Uol~y k!l{n'!,'n Olnd lb~ their "CUll!!! S v,~I'Ue\l.!lnW!s;~ he' cSl,~rn~~.cdby sca~~:Slic.HI lllnp,iil1Jg,modc'ls, ..1III s j)<Iie()untol()'l~y, ~!cll stmi~;.t~call(alcukllinllS,_ e.g., mt!dSth~ll.~I:l.cotporun:: r;~cil'l1!~11 J:llllnhliln; ,,~drel OLtiH;~pecie!\ <!b~u!'ld<!llw~ rec.;m'ered ~(I ~ date ami kno\ii;'n ~IW.p,Ullg b:~~~!>(t.s (1)-ar:c nmv rouune ami m;mdalOry. UllIfOfl.llunarely.. none al.lpear i~.l Utebnrlk,.~sh~JiId l'ingE;{)~n,.phen;, rout illl.d:y lC<l.'5C ~.pmt errocis .of A. D ..and l. Orl S by mtUltiph:;regr{;~sio:n <'l1U!:I~y5e$~i!fld miter s~at isti.ca ~ Im:thodls. ib!.lt the j]onk olTel.')1 Ul.O slich .f~nl'.t)"Se~of the pn::hitlwlI'llI;:',.vit'l1iI1I1,1,

!lgO, t~Ui:ll eoncl t1lb dlfl!. t h:is :;;LJ ggcJ;b; i:II2l~

mast or 1l!e'<U'ly <IIU specieswere l'orme~11Y


~l Ol!!

r:l.e$~

rnost (lrnm~rly .uil :i~lnm;l~. fli11:l 'Ir,~hi·, o pel,ago. The verbs believe, expeet, predlcr,

,md the Ol,hrellbs I~ossi· bly..p:I·e~lnmibly. p:mbably. nnd mtdouhted~)' <Ir~used lhmllghQI!I~the;bOQk-~_g" on n\ie ~Ive.r~1g,e lOre[hfm [wi.e.c per page UI1(hoe chlilp~ Ie<W em spee ies-srea I:e] a~loll !l-lO precede
!lpetLl~.'lJe" ,Ulc.l. $I:!$petl

PI-IYSICS

,A,s,tronl'O m'lenj~

IR.e Iali', ity


i:11Stci.!I'I·S sp.cc~.a.:~::Ind. ,.gen'i.:.-ii!. ,.1 ~.hc. •.I -.''Ofil.!:!> ofre I OIl~v:i~y began e'llrag in£liJeOple ,u cenru ry ~lgO" A l{l]ml~.II1I.~lU l i!>l of ( ctti,~rges. <lw.in~~the!i!l~,~S nUC~u;Mied frnm ,~!e.flrto year, marn)" return ~.o [he 5j,!IIl'1,c; thelnes. ~e~~.l~vhy is wrong,. soone inslsL It i~uni mil! mlive.. Wo Memm'lt:ieai. S'I}1 oUu::::n;"

sci~nli~!t£IliOmli~l.uy draw ~~5 l1tmchu)li!il~~ f]rom <;lmuly~e:-; III the 1'i1ll,1I .. cha.p1.c·u. ",:ouds Hkc ._[ believe" ~lJn.[I··p:l'O~l3~ bly" become scarce .• lndlthe rOLr]lnelr beliefs
Mal~~.l1!el1ilS [h:n b[l~OLIIIt; ~tiilltid ,IS, f'::lCts.

Why WQlql~d :stlm~'.On1;l~l,1end 12 ye<i!7; e?>C1!. v:,uill!!l. d.ulning,. and ~nkt'iil!>oop:k;'l!ll1y SCfUlhliLz~ ing 25.iCIOO bi.idllolleli: and then lIleglecl moderun
,lI]~liy::i,,*'!'fie
;t'Irll\i\~

't:r elrnk~" On Illig fim P!lg~

oJtI~cITlI:Cfu(:e ilUild~p~ iiep::atedl:y t!vtreaRar: St:c.adtn1:irrl Slates his, disdai~l fbI' mflliry th~Ji!:l$" im::J.~.di iii£: cOIl1IJnU>ers. d~,t<l trnlllsJOlI.1l1at~.olls. (~.Ul'lmIlk~,e~~~,u1'-e.'~umBmh'i, ~\;'ill~rldiz,ltiol'l~,

The lisl goes

O~: GCfH!rd~

:l:Iel:pt~i"'ity Mses

I killed maIJI!,!'l'Il'itt ie 5, h \f,uolmes, Kmmiml ph.ilJmi,op,h,y, One or Il!oth


III n r'~"'.ti.o.fla bly m:lIl1p

"mgiC'.dly c~elllr w.hy Sle<lidnmlil didlilo~. mu:!e;~ lb.€:1S."iLllli;Plimllili"ll' ~11~ ; eKit'1~lld silri1~ul~,uli~![lu~ly_l1ike qu,u~~it,lli\Ie :.'urnl.~s nfhms 'tJ>-'lin dal~~ m.m (;'O~~21oomtew.~Dh oa]iIc:t'S,\I,'~10 cOllldllu~I,IC l'i!,jmp~eBiil~Tetl1n;"so'l'I~ m:l Mm1l:~ UlS~UI'S a.COOllnlS of the Iwdrell hi:~txpto'titi:'i-'li:, fl!lt~boBll~r~lkfi[]~vtl'i Nei.v Zemla:ndl <!lvif:;jJtlllJ~L \wim liS th~t I1il<l'n'y modem s:,peci;es al"r.iiw;:d niy a~, o The dal:<l.in £Wiljclion (m(" fJiogeUf!;n~fJh.I' or ,I~t~rlhemigm!ll,iI $peci~ decl~n.ed. [f~·~s if' qf7l"I}picoJ Oin:bt <111.'1:)' "~V.!i~.lhlefar I.lthe!11l nnw une CDI'I'1.bi IK-d the 1940 amI 1930 M~i:lilhmlLfm 1(,]'!Ilf1idyzl'. Bm \\!~ llhf~n ]~fL¥C toluol!: past S~e;;ndm.ml~d~~Ilii.i:j.sa~s sU'uv~\i],ngmotkmll of ~eh:p~()ine di~:e(~ou'ie:s.<I ~d a~smned th,n aU 116FEBRUARY 2007 VOL. 31S SCIIENCIE

There 1)rc :1:10 g:rniPhs of prehi :>~OIite Sip-Ioued aga~nM 'eithe; DorL. De:;:pi~e<I \\'~lOle cl~p~'ex .. on spede:s~.;n'ell re ~ath:m5 a Ild ~he~,I' s:IIOope (tenned ;;j" the b{J{lk ~)r'e)leml'>olllly .. ~ing~e gmph (ng, 1.9· 1.4) o:f m .. llhs:si I J.a:ta ~Ld0!JliJJ<II!: "\v Ibr C<l ~L;mlat'il1i:g:: tllatpk1t for pl,ell u:swrj,clalld! biI~S mm.on.g ~veUi .i.~I;.ml,.. in, TOimli.l11ie1.f,!l!!oi .. :: y (1:1' O,'fJi7. villilil<l.n}'~mc!lli;m,ged from t~e!il~!;'I(lef!1 z HUJl6, :S~\'Cm~ grnJ1ih~ ir... the- :SpoL:d'!.::.!;"'~lre~ C~al)le:,C!iJD1nbin:eloss:i,lamil rnoo.em SpeCIe.;; 0111
(J

I'l'ilocroeculogy. {wemll~ rmml."W'Orks, mtcs, SI;i1I.iS~ lies,. <ll1ldtheory, He al~o rq:lecrll:ld~y crut:i:ci~s "''OllC~nsiombybi!il:'Jeogra:phers \\~to ~lsed !'Ilese ,~)pm,LCh~ {If C(l!,l~"epb~!t[lt 01'1111' MacA LrJh~lf. 'i;V~ IS()Il. {md Mayu: l:m~ aJso JruW.C5 BmwTh, S~ep\hen l-il:lIM)ol;~1 MtHk [.Qmoli.l1iQ. Snmn t 'lPimrn" R,(ll:lot;rrt Ri(,~k~~fs., DQ~ph Sc:lid~t~r, fiQ!lli),i,!Si ~~Or:!1IJ;lL';§Ind ~~~I!I dit'i..:;1~1~llre)1 e, m \Mth these sdf-impQ'i€J blilildl:fS . .1.1 bt;connc..,

rehwmvi ty <urI;:cor r¢{;lhu't \~'er'e ~hc t~lf'e~()h mmhcL1unic.ufI:l:l· physi,ist Ueuui li\l hl~<lri. the GerIDI1<lL1mmhernl<ll:ician D,/Lvi dl .11.i ~ben. >0:1" the Dmch thc()" ri ~l Hen~tr~klorent7., Re!,~ll\' 1> PPl:!l1,UW re. if), d.ea.nis~ has~c!l'l ~.lllloLLnl{;,(ld time ~md .I1ga:ila. Its ellecis ;jg:o·t e1>liSl,Qr they do exist but .~1Ntl mher ~"p:l,an"luoI1S_ Th~ slLnis (}blub~, ~~W!<'~Il~lsijll!g the pfeceSi~i.on ofthefi~ri,~l~~!On UrMCi"CUfY, Or IhlJ Ilthc[ ~m.~ ddccwd'!by be <L soLl~)ed"up Milcheh:O:Il~M:orley ex~)erimen~, St.adm~ht !SIi.' ~.n~n't' ;:[_~ Eiri:-oteill r~ i{,;'t~d-{lr i[ hi" btU that i~1I~l! [alh!..': atmosphere ofll1t.:
tlu;:nries or
l't~ Iic:~p~1JJdlby ~ 'Th;{!fMI'f~r l~ at Il!ie' l)~pOi!nll1ltlll. 1111 Hlfilt!ll)l a,f Sd~ncA!'. )(iI!!IUI!! Cl!!nMr. '1 OrdQfd 51Ir~'I, H~IV.tI~d IJini'ief~U\!,. ~<! ~bridfl~', 1'1'11 CK2138,. U5t!, E~milil; g,<I'li'lJij.~@f~~.

1il<!!v\ulilJlcliu

'!IiIWw,sciefllcemag.org

BOOKS ,~TAL,
Sun oOT E,~nh nil somecnm'bim~li{)n. Whe~ .1U ds~fi:.li~l~d~ thsrewas :I~ways, the cJ.1argc llla.1 lIle tli1enJ)'wm' too much like i~s .,U~U]Or:
ton Gerl1iiL:ltIil.(J,rnot German enough, or N),!)

!ned ~Cl:et~I ""nruli!;'.!s~!red. In a 1J'O\~'ell'lt.1eonseJ1li.lIs. ob5.crv~uio:u from both Lick and


MI. '!iV~~!lnn ~q~:r~e(L ~~lrl'!Ji!1~ the Arll~~ic,[PI Ilsm:m.OInica~. [ide toward abmad irmn uni-

me;I:-Jl!I'eU1l1en~s.A'toCrelmsten

perU.od:'s, aSlffHiloirncrs •• ne: .... dC~I~t.with llil.r.::: !.:lit deeper iUi!.1pl ie 1n ..io!il.~ uf E,i nMei!i!~ ideas, ne ccssa ry., .. apa rl from the
few !\p'ec~a]i:;l~ 'who work the equar inrrl~. \\"€ diu ItUl Ili(l(ld to

rel,!les.

the

Cmlil!is~£!l1s; r<Ls e~.rnliUi:rlgEf}j,~'IC'ln .\ .Jlil".r: The>Rtle(! 10 le-~:ll(elmi\'Uy traeksthe ~\'3yS in which one par.t.ictJlflr CtHmm.ll1'lity. astrenomers, handled IiL~s~ein's r,elal~vi~y
Jeffrey rht':~mi~~. rongh~y between .~ (I and 9:1. 192)_

Jewish,

(If

too cosmopotitau.

Ahhollgh

GemH11'I and

"rppeOlV. CrelinMen'ji IIHIiI,~ng Focus Is on AmerIc,m


Anllfln~ these,

British asrrenomcrs cenlmi O!!ndmoM ilhDn~i·


3SlrO,n0I1~,er5.

versalacceptsnce ·of general rolmivit)l_ Add~lim:l(ll ~l,Ippo[lrt·, .1 ng.. if more complex, news came from the recslirlts 0 f SjJOO"' n;<lI~I~ne measureraents ror~he g.1it~vi~alionai B:;1.lshiIL There is so:rnelhing under!\'wnd'lble-"!1drighl-ut"{_l~q~

iLi IIIS1rD'D 11I'~J u.D1f The RaCE!! to lesl Relall\lily


jj1:v Jeffrey Creffm:tefll

uudersraud the unde['~ying (!IOl.1{;CP'~s.nor do llUlnlY oftiD,c

experts," Btu is lhh true?


for example, is real,iy probil.lg only the equi vale mcepri Help le, ntl~
Tille redshjf],

d~t\iPiu;: I:~'eritiir.;lof,J

re:m(![k·

ableas Im:lJlom:i.cal cOlnllmtl,li,~y Q 011(' of the fj rMre<l1 ~}(!'i.~r'e'rhnuse$n U.S. science ~.th'ere i
p'i.~c!l;l{l r wnrk., ThmnO!:> Jeflbl·MU] JOlckMJ!l S(;~. qllhe hated !;if1!>tC.i,ll 'S general thc,o:-ry {"t~o :lnet~I):~ji~ic<lI") and did <JIIII oot!ldhe WOh~illiUC~zi.I!i£_th~}r·.i.z:i~:lg. :lH1d rlag!l!_tlzl:liJg,,,,li:W ~"Ullfi'l~

UIi!I~·I,IDlp:k~a!:\lII]n~ypr.::::;;, One

IiUm,ly

'to!:::iJnk E:i,flIste ill's beat. [~ilred bot hi from the UIl~v~rsilY l)fCh~.cago lIIL'lJ L(JIit,,'CI~ ObsewV-olto:ry. See' nonetln,e~,es;~ gOl ., usefal
:ri~e Olllrt of IHn'i~n~lnil>~ JO!lnr;.~ $~<.!I:l&.~\;!w lilke ~l~.y have been Il'lIs.l~ed ~.o~~rd a more :i.~:>Iif'u" IDilt:nn,d il1rte,,)~-ew!'ion nfthe !~e(liry by See's \dluper',u~\'e r~m,'1rb. An,other <Ir1,sry bee 'WU Heb ~r ("1,1 rt ~ ,1Ii1 :;!~.tron(lme:r wh (I t-i. ~'n.3d worked at Liek Ob:;cf\l3wry

,(for example) thecurvature decis iOI1~D ill ~!jVe 15Im 97800.'3];1 ZllDJ_ i.1 is (!Mitonmtll~.~·I;l~ j my de~iver of !;;~l~~i;;~.tunl!¢ is. in ~~cldi"' that its. V<:'OOiet ju the hue i9.20:; • ~~olllm the eql1iv,mJence pl:'i:lJl,C'iple. pWl~ to rhe ~e~! 'by L~gh~ W~!h~:\vo1!ll1OUH~:<I~nS eclipse n':~lLtll> tbe t.."10;tr.:;, O~(l :m~I)~!; en d C,)I!i~IlMi~ of J(l tllllC't ion, So lWO of ~he "elassic" three L.iC'k :'~[1JdM L W~I,~n!i! brnWlgilt p~)we~J!a I experim~~~~!'i d,o~~ut. i~I:'LCt. t!l;lSl the ~tlilile a SpiXt S Q:f th,(; gCl1lcml~h£o.U'y ofrelo:u ivhy,. II.l suprpo:ut 10 rel!l~ivhy,. As ono(; AnlcricfI.UQ a suoncmer {R:Qben A irken] pu~ il in 192 s:. Ql"tl\er to re nder averd iet, t he observat trlml~ IJSlWI.1Ul]l ica ~j ury n~eded--l:!.t1d mt:,~J,~'-1~ "the vuh.llli:s ml'l ill ~llill.~I::!:nui:J~ ~mcorr~ wid~ I~~ f~le!llry,EIliid.. , :1,10 other s<llis.rfLcwrye:'i:p:l~m;la,- sped fie S~<liClilu.~lIlt oh~e charge .ul. is to ad j Ll[ion .of the dijOp~accmcn[ I:> ~."'lli:~~lbk di(;~jtt:.And dimt requi re s it (:olf1malld ofbeth .... W li'aetln.erwe' [Ike' i.l OIT' not.we are obi ig.ed to experiment >IIul ~l1.eory. i :Silit~ the I 92Hs. ahost ufolllL~r (;xjwlri~dUlilil that iii these ll1flS~ ill~~l~K'~~ [[J::ld'~hirl. mentshave £i.ottcn.ll othel"8isl)eCiil,onlle II~cl)c'liillle~.iiQin.starlligbt] Ihe Eillt;;t'ei.n ~he'OfYhas !i:Wlcces:s·lld ~~{lodli1le test Of'~$lml'l·o!ni C,!I ory. Bo~n(;"in.~ 1'O\dOlf beams of'hhe sudilce of ~y cbservanons ... So once the j!uy passed its Vc'rm s in super ior c'()njnm:~ion came .. in tille ·"

ere I iasteus

U nivl::HliL 'I' Pmss.. PiinlUtc 1'1, NJ, :lOCI&. 42B pp. Sl'i; {2::t 95,
Prine 810 n

Or

~i~H~i{.~I. hei l

~ht'l[n'y.~ike '11~ nl~'!iI~em

m,II1J.

w.al:ketl, No double jeopHrdy. lim


I."w. )l'ei'i:iln science .Ifl(l~ qu :~te,. T~llproMem ~!o; Ibm. el mHu·e in the !l!1!,1~II:eT (If EUUl~~ni~1il I.hcory (01:'.

and lhen \\'C'nt O!ll ~{I direct [h~ Alllq.!!,~H:lllly l1~n'~Jmry .. O


One lwmn~!lIeIi!11M~lm!llo-

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lrD\\'liK !lot:so SIl.IiI£ilhu

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~nnl~ld, dl<lt it d~noc~ion [Qfs~li1:t'~igh[]. rrilHl .a grd"It,ning bod}' wo!~~d moved lle ~ ex~s.lS e~ ...erumUy <IS sbo\'m on 1m pklles. ~ ,[1.~?5 ]i,. ~h:i is >does 11:0~ .~_mve'r~r: m~ke :1~le,(II tOW<lird ~:I"elow<:r, elld [) I't Ine specuLI m ,.If: .<IS .~ b e~!eve~ ~.!l. I'he~he-nry of !"I~~;;'1t!\'~.~"y.,.!U :>1] Ait~e!il d"imed. Ule iSS!Le of the I·Wshil't •. r I~ v,'.11£ :s€:tik:d by ~9,2~. Lhen, we would be .~!l rd ~ ~LI'I~rred!.:em."blc [Ile[(:l~c. ~md it <!OCSilOI Inlt l:Qr1lII1dercl>lmH.l111.e;o;.tr.aQr~in.lIry impore ~ seen) :1!l9W t]imt I s.lla.ll~ever S'!I,F,lUOIIN thM I:he" b.'l:n'i;.".lil Lee{lrdl::'XlI dll'l MIi~~IbtHl~r t;:ff~(,':llfIlt;:~15 ~ .. 13 my ~lIfIl~s..~ (,':11~m(lr~}nlledfi nit ;' ... . ~ O~her }\rl~,~ric"mi.·~q~li~d I~sS dn;m~ , iql~ vn::mel1t:ii l~wt .~.v.ro~md.~m! (t 1\. Repk ..! ~ ~1~~iM~~C~ 1J1~~k rdl;nivity. \~(~llhlD~~ \~~-~·i~c~ obt~dl.1Cd i:hn:c tlcc~LdGs. .Iater \.'o'.~.1:il th.~y ; {'<lllllpbel~. d~reciQro:f Ihe Lklk Obser\~I;ory. dwoppedlrighl dmv:n lliie st.'lin'-leU ofl:lal'\ ..~rd's i!:! ~~hin",dlm1i.'1!m;iEiI1iS~e'inr 'i l:hl1;b:;l~i~of n'l~tiC'. Jt;:ffi;lffiOIl .h.bar~t~pry_ E"p.e[:ili'ieIil11a~. c{Jtidim .~ U 1.0 S t'lbSCirVa[~ un. Ai> pre.~cn~cd by ~. ,~ol" U tiO:ll!> dll11'1lg"e: un't:W ~lrm:lbette.· tt.'lCh.ologics ~ 1~~a:gi:L~ I913, Itio~~ lick f,u:;s:iJ~ls i.jjdiCa~,~d in :::'I i.t'iiibe j.'in~5~ bh~ [i:1C' iM}~ 111 ~OI~ a rid i:'lPt:c~J~llil n.ew view: "dl\1l.t li!i~ll is sllilojeC:lto !2,1".wi.atirOH cmkm ofe~!liecls" ~ Olundlh,H:Ei !l-..t~~U!s law ,[If g'raviwD:iorrq i~~ nl0re Th~~ w~.:>e£;;,'~ htlfgerroi.nL Al~U;lJi:c,u~ 9SirOl.lOmcrs Ofi:h,c ~920:!iw(!rc kt:CIlI to jCHi!l5 :l'1,ccnra~~dli.un Ncwto:I,'s,llliw:' Oh ... rvmion e ~ and themy met wit'h great IYfeCi!1ioll- 1."75 !':OUl h~:g~ t Iileo ry im t:,~vor or pr,lgm ,. ~~c

(dlle g@n""!fl!~n:!aliv.iry "equivO~bital. ~ie~t. Gtneral r~!atMlY ;j!!il:p~~in:ed alenr" to magnetie imi~erO!:c:t~ ~helliIillnM ,ofMe1foory's E!.rih~:II()n. p ions i.ndc'ltrrody.imllmlic~) ~l2s~cd another aspect ,of u:ekniv:i,~y, r ies ~ dlo~~I'I't neces .. Ihe :s.~mvinM .[Jfhi.l1iary ~'lt1lrse.xph'ilretlradiaiicn sarUy lfDlCfI.1l th:fl,( ~he lhrough gr~,vi[y WBVCs.--yet :.motl~er.• tlild t~li ired. ff"(ycm. "SP(}CI. of ge:i:icm~ [d1ii.i'\i~ly. em,t: M2I)'l> elost:d" Tah Ihe .~Ol]£,md Il'OlLbled e·''Lrly ClreiilllSI'en !niiS doue 1.1gl1cat :.el"vioc .md his~ory of ~he r¢ds~iJ~. ~h~ d~s.-r;::r\'e.~ OUT thnnks llt'oIckin,\l M~ bt";,nqlifllily [ine Am,er.i.:caI1 f1s~m;llOUilliGa~.lI"C.~ponscto predi~l.~m~ dun Ugiu In:avcHng om

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W&!-il-ha5~d Ulil'llllil ullir.atioll1 (<IIf], prOVidE!' bNI!H lI'@p!OfltiFig 0 n di s;asl>ers, ,c:®~rtlina'lli,ol'l ()f r,~.SiJO'flS@'S. dissi!?Jl~i!'lii!tiiln Qf i!1fo rmatio[l, ';11'1.11 soctat flel~rodhlllg M de~.fiVe'r <I!'i~istilrice.

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to d I,d 91 I, B urwhen narural disasters, p~[]J~ic health threat~. or ~e:H(lri[;:t ~l~I:;,'Il;b; O(.'Ci1IT ~hl.itI;':ff(;l'cn ' 1:I:'1fmsaltd~ f indivh::lllli! is,or more. o ~H1 opewwr.s car~Ulot h<!nd~e <'III the n:ql!lq;:~1s, Saeh Jliii.u;ler,.. nilily r~uir'l;l m,~~:;,i~,~ cmwclilll,nimil uf

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eRGs could be raalntaincd by 1.1iI remer: service pHwlide 110;. ln smaller oo];mnullil:ues. CRUs (ould be run byttainedvolumeers with <I fe'\:v'pro[hsimn~ <'I1~UiJul!.ch VOI~[llI:t:~r I!I:~ depilli'llmeUU5 nov.l eperare, Lool!l~

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filCl.:.'d. \~,1eneed to understand llilello:I.'Ul1!S~nd policies A,'Clolll'lnlll.l!l'IIi'!y ~'SI[iOIi'ISe gln,d, Mmjk~p ot <I Worldl Wide ,Vet> p,~gelI1<ll supports reg· l>rre~ocIla lIilUJ!i.d~ds nl'lIt1JiHLOll!1 of i~te[in£l ollllo!1.lseholds, reP'O~lir19 o~·i nddents, re;[juesUng a~sistalilre" and respoodimg 1~'i~~lgclie~.te~IiJI!I$lI1lie p;:1:r!ic itl:l>e(il;. Th:i S sLlggesis that local, Imtion il:l~ groups ti:uch as ~O< r'(!ili!m~Sl he\iVi~ ipediOl co~ml~lnti.~y s~me" ami :fOOen!l a£,e!]cies could build Cl)f'UlUtllil)! respoase gl"id::; K'RGs.) reports and IH'omptly recognize pt'It If I: m... andsome healrh SlIl)p<lrt ~roll~)S 114) .• Mmw
~.nct:'J~~iv~s ereaee s~~lcce~~fnd c:onl'm!llli~y ilue"'JC'IiO:IIl~ reeue ..I: res udem·w·'resi>ti,ent <lSS~ Sl~lllce. The " p llil whucli! prr.u\o'1IC)' ill r~!oj,pm::til:t! U~iI, It tvill 'l:'llrr~b1Ilt~n:wr]lm and World W]di.:l \\'i;1cb I~"\I~ 'dl~~l t.:{lple kr1JO\Y :abum it and d~"'c:':JDp closer (.{lmllli!llluily eontaets, S~!d'l~c,:;~i\lilt~et-> \\'rHlld p:~j!!'il:llleflh:;ti\'lI:l for 1'l1,ln}, fl~:fpn:l'~. bar gm;·, be i!f:!.p(nw~n 1:£1 ij ~negrO!Lt'e\'I.'it'h exusl'i.J1!,g 'bl.~i~dltlJtlSt and im;;rea~ soeial capital dll~t \\,'HII sceial IiIc~.\mr!dllJg s.i,lcs!lnd ~O(:j}lco!i~lmlmi[y 'L:r.UiII1111~JiI~ agencies have been s,imv 'to adopt gl"otlps. However, develnpling. <l research S(lCUJ~computing fo~ lll,uional see~l'I:iily. d:i~as· be reeded dL!ri:l1g maj,O:IF er!ile~ellcies. Smue t<lIi.!SlOOptl~s ~J!12>~!.~ny~nIf"lstr!illl;',turu. agl\ru.la. testi ng. anlii::l plu'l~.o:..ul. itnp;.Ie'~nL::.mIe.r~spoll~. Olnlc] tlIlIi!Jerg(OncYWlOi ( !L i(;f ~uc~. Ol:~ .!V~a!lfUu, ~.nion eeuldmake eRG\!. " ·n;!,,~..~lj' wi~Ulin 3 CR(i~ sh(lou~d !IHna:elha!!1,~!11e!ne['~rellCY h!!l~i ~~lm~lly~itJ!!~ti(ln!l. be to:) }'~3Jt'!>, rcpor~il!lg serviee as; 1l~.re3.dyp;rop(}.,~cd ~2). c:iilcnlil";li1l.u(l biological snaeks, amllC[I[lp€m. 'ttll:e extremes, do 11Ot. Even \II,!.I~en there is They should also be nH)il'e lImn (11e U,S. R~f~rlllm!!s D1O'p~lrtm.en:r~)Ii' Flmtlebuld Set:lJ:ri Ili (DBS l' fnl:'LLM:.iJ <hurII<I!j:¢ In eommunie aliHI.l[i;. ill]~T<1I' :!L 'W_ K;iii!~~ ~ Qk~;Ii, H. TiI!arlo.,I'lQJ.. ~~fl:w. to, U ~~I.ructll.re. Ihe adjoini I:lg areas need ~(1conil rnul:nfOlUUltatli,on NCT"'Ork lJ). wbiGh focuses, like (lOO~), whell'~re~~ idle!ilt~ t't'lUi1kll ~pn:rl jln{.~id~nfs;in secends, receive ~mc~,ene:r bilforl'lliHiol1.fIIl,d Civic ~1;1l~~P;; could dis,,~L'ttlirw:[(;'li!llfnr!Iill(!li!lu (JI1 !I :>I roc I. by street basis A CReG '!i\;Ollkl be 'nl()i,~~etreoctil\'e is 'l!SedOlll <.!I.r~UJI~'lfbm:i~so :v:fit (mUIi~ CQUII1~ruuJilitie~ n~.i~, ~o
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.nicare I!t) £t':PQr:!m:'l,dre,cc:l.vc: iusrruetions. As the lnternet matures, k5;lIe.~iabi~hy win

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qU.lntUiIirl'iPh,;ts@lr~ nsma 111$.

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b~wt'l1:'1l tire da~se-Ji is relulted to tUne(lllrtgydtif ..

[e~'eril.ce etween the two piaases..Thi s energy b Ji fF~rum"C de fill~S, II. iSitlgl~JJ,(:.~Cl'ID' ~al~e tl1:!L~= vml~she;s,)5: the 'mnsi~ion P'Olln,t is re<ill::;~1C:d (t~ml is, ]t beeomes e~~D~r nne rhil!:S1~ In d'I[.'1~ foil' illIto'the Odli!'o:if'i. lI'"lte~:r.ticl1i3r \~y this S!imlii!il~e e~ea;gy:-;calev;llli~hes i~enoosh to cI1,mu.:~e['il.e each lin h 'ets:oll:i,~ C~<IS8of u'aflsil! ian. This: COIii.~ oo~11.r]f,~'iirl~re energy t'<t;,tI~ '\v4~expoecl~d he· ! l'h V'Jlf:d even IQir se-ealled qlLill1imn, pha;o;e u",ull>ilinn!' (2. ;q. whieh <l:D'e' ~n,dLJ(::e(] y chang:in~ b prelisur:e. Imlgl:wluc nd!ll. or eonmpo:shiolll rQr Ol.marerial held Ol~ ahsolute zere I~mpe:fattire'. I-[m.,,'C'vcr. ~()'w-I.clmlpcrmtmc studies by
Geg6l1~~'<I!r~. {~f.on <J~m.'tOlllic:lnllgnet reponed! ttl onp~gl: 969 ofl.lli~ issue (.:I) d~nHJll1iit:nHe .;iI. ~r.rOl!:S flliluD'e Qr (l~lr ~lIl~L~rs!anding. They fint]

Qua 1iI1.'Um,c ritK~1 p!Jim~

Magn~~(fiellcl

QI!!!!!mum ·f,nes. OJl(l! spec:ulili!ive i!'n~rpr,e."tiQI'! of the w1~i!lpsill~ e:ner.gy S(~I~esQl:.mwwdl <lJCUfI!dlhe qUim!!JIIIJ'I ((~IUi(~'lp.oirll is ~h()wn s~ltl1l'1f1i3lii(aLly.lIll'iigln I.emper~ IUH!~ HUiCWaJlil'l'!lma9ii1~tic sp'il1~ ,[ji;.ru pI line eleL~ '!F@1li Ib,lI~din. (II dis.ell'!~ng!~ !:;riliw,1 r,~giOI1i. 00 IOOClling, lAC spli'nlS may (I~d~F~Q 1'000 <l m~glnetk lI1I~t!lt A~lefn<lli,;,el~, the '~pilUca~ b(1{Qom~lm:;a!~eilltangl~dl (5) wilh IIIill' <sriillHW(luaHng ele~lnm [[uid whliclh~liIelil, at <IIlower el'1l~r'91Y s(ale, is DEt.!Illiied to Jmliil a InN'i)!' cl~drolll ~il:Ju:id.\!iIh~ all tlieSI! r1f'gim'lS $OOIUI.dco n~er!JI~ 00 the IqlJ\3 ntum milical. poilii~i{~ ~t~Fy. tille'
I1iiIml1{l;I~~,~ri.~}',. rB::;~g~l'Ig 111t::con~.i!l.lg

q~!a!ml!n ~['itic<d ~~nim Vll YC[ dJI:S.p.DI.ebt:'illg all ideal candidate Co.l" [lOT onc. bm h., .. oocrgy scales. \'311'1islhing 3! 11 o qm:lII,tllm transitiou. de.1!I10.lfISlrntin:g theory. the .Indi~er~:ll 1~11s stutHed sitioa ~~lnrH:TillLire. The nlillcUtl.H·inllLS ~.h\:,1I! D¥ rif;'gCIIU\lIfIl!lel (II. i$ orne of." !;!:l1l\\fi!il!;l;IIUIIt'liber Qummun1. pfumii;; Imrn~itUmlS h1~V:e rMll\'t:':d un OOOOIWCcriliC'tll--c\I'Cr Ilargeuibllsize aUDtl to:llger that seem ~olie olllSI'de I~lis cst~lbl~sl~ed fmlneroa~ll years fii'Olnmere th.eo.UiCIlicru~ c!lIl:ios.ities re wmk...1?l"ev.~~JlI1.ibildac[~)rs~9, JO)It:llv,cli"l.:::v.cll~d :su~jec~:sof Jm~n:£l.:.Xpcf.i,mentil.1 fUtlll 'lhmB1Cit~.cfil~ ~~!,I!.JtIl-Iill;mllllhey tklmilmllC [['Ie pl1-ysi.e11~.pn:~er~ t~enliSellves di3Crele~y tIIuc!lIgb tmusum'l. oomibi1'0 'be p~opor!i.onl<llliO fn!ereSI "Wherei'ls p'J,'e;o,.rio~bslyit\~I<1.slhou.sht dt:lIl. Ik s, Th,e$(!proper:ties'~~lld poweffi of the ~~ilfe~lI!oelb~~\!,J~~.the tell!lptlnl~ lIU,lt:IOriS c:~i~U""<l~ I;jxrml~L~~sL]n"Hlml'l qjIillUU~ :r.~ cil1'u!l~~8 f ph~t:;;ea~zem ~~lnp~Ol!t,llreW{lii,ll~doo o ofUnte !)rrl'l~IJc...11IIi'CI(;."\I.f~lc;e. 11hasbocOlm; c~e""ar lure ofr~]~ 111fQl~rJilland dIC'Imnsi.uiolll L~UD1IWrnwith the ~~K~ory, \,hRh:iSi:l; dees this and more. In a <..klitiQIl to S,tI<lDll~ po;''''!N laws. GegClIMl11 (:1 I;Llre.~l i~ Ilu::1i!o: t p(l\\~I~. or cri~i"l~ eox p<lHe<rlt5. ~ml the p~~nce of" '11!ii!1I~\m~ ,ruii,.,1 poilltm lh!1'll ch!1'll:tl.clelriLc:e: !J:ll: tI ~1~.vern,,1:U;~y!iQsses. e illloo'.~Ulie zero co~,~Cllle~\tmhel.ess exert <L prncd" ;~'C <i~1."l11J~ cn~mg{S in physk:<l1 p.ropen,ies, 31~ Ibllnd inlh~c\!1(;'''ean ,I nmt~'.i,il OJt. higl\ler ~e1lHperThe e."'telll:i.i'l~of tlli!'>~h~ury to qu,m~W!.m. of wi'! UI:;h e!1!l'~ll..,t~ fi;,om ,111; ~llH.mw In critical qtdte diMi,nc~ room the l1magphla..,e ~rnm;hif,.,s includes ~hecllC:rgy co",i o:r a p.).~nt.11.11 a 1I.11:;liIl,n~U' auues. 1't could l'or ex,t~mpk;, l'I1la~cInagn.cts, n!nctllOltion caused hy lieisenberg'S qtl:mll~tnn ~~l:c Qroer.~llg tF~llSi~~o:n.'fIle ,'I:~'Ilgesdef~ne a lmo slIpen;:mlductO'J;S l.;'i) and S:l,pe[~ondlillcrors ,~,~ner(¢:ml~om I,h~li f~1.m [[!rK:~rtainnly pr.illcirie, Till it> l,Jlti;ertllirnty k'b'1i:lrli irneW ~11i;\IMY ~q!l:l ~111)I]~o\f!1l1111~tillN Y~t lhJil more ~h~ inlll.1Si(6), } ~~".ffls·~m','e ~l1~h~l the\'\I~!~~r<lJlIfI~~ (I~~r q,L',>LIi!Wm~~Olluer in ('Xlns.!UUn m'()tion even 'It II1.Hlgrleti~ t~w.:rm[~[o.!!il'. ~t ~ ~'n~Im- ,no ~tin \'iln" been ~) COlW'c:l1liorrml f~lIl.lC\1!IUr4: for lifl.d!Cr:;t~~il.ding abs{)~lIl:e n:fmJ~'id eeu bCCOIllil~ V~g,ofOi..Is tha~ i!dti.ln!:l zti.t~ (~~lf~MllliiIcr,Llk""lI1poiru, This l(!iUs us t~ml~he lfansitmy <Ippeanm.ce ,af bubhks of it drives the Illehii Ilg of onel'mliH or ord~r ialiO qtmnl:LUrli oCD'itical:ity ill metals (7). anoUiII;:r. At fip;.1 gl;lllu;e., ~'t ~lpp~,W:t> 'th~LI Lhi~ ili Wi~lhill [h~sa,.11~\'ii:UiliQI'!'allr.IIlli;f;i\¥mk.e~'~ri ~~'ii!;eri,rQm~i(u:; ~h~Oldy imilpot;!rullJ:'il rhY."~LlI~ lint ()octlrlriltg!It the tluit~C;l1 PQi~!iU'KI th~!.[1 bcfon:: the trims ~liol1 tl!lnfR!'m~L1~ is, r(~ac.hcd wh~t. istI11'I}pe~jlfl~in II!ic ~mnpom~t.1 :>l~d~,ed )! p~occs~ b b~lbblcs; of 1.11~ ~.t'i!:.'ln~(:; it'i:lW IfJJ'l'IiI, ~:pplt:llir GI~~iri\"m;r 1I,tldCt)"\'ro:fk~t:r1i. Yb[th.,Su.,. i:s~;,dc.ld~to:ry~~jU :~ii.dmn~ tlml,t ~mltC'tLl~ltilll.g tmJ~1' ... .I~ ~ ~,e,!el' be eno uglii 1:0 d11ll .... <'LClerize I:&nese~ralbs::i,icolI.e mu,i n:n:ronliasnet \\i.here the eloc~'ron sl)im; T~eynnu~t ther~~'}te lieOiUlSi(re~h~ lmi.. 'irml1~.e Ih~I'Ii!~lves~1'I ,111tm;~(}~~il~dl1>:rnlO~!.il'lg t~)n~~. iJlf!e<lulll~ors in t!\oe- s,thoo'l of Ph.~~t~ ami j!j$IJQ~!N!lY, i i to ~[~crl1D. bLIl 9;PI}J~illig Iii li.nm!dlllD1agnc'ti.c ni!~.d 'Ilel"Mliil)' d 311S(;S",",'C, h!!1,\Ie k:llQ'!iVI'n dm'c .fll'nd WJniV!e'~i'\Y Ilf ili!miPlgh'l!l), IHflllli!l91h~!11!6,1 ~ ;;l'm:. ILIK. en~anaes fhe qlllOllmm~ a;~mtiol~ onille ,~pin~ E-miliil.:: aj'@lm.ph.Ml~m.,,,[.uk Die{!~liL'e ,J ne .... rphy.sical il.~L'eC]ic~t lhe!%lll~~(;~l!~~~'iJ~lls requil'e el~er1,lJ';, bl~~ the ·'mlCtgy ,cost" for dlCSC nlletHru.imll>reduces to nothing ru;lhe tenllpernnue 11PJJfOO.Cil.estbe 1!I.<I'I.l~

cbnge.

i!J1I.nil 1l!Jlll\Jl:isUiil i:sd~sln)}~dc{lJ.tlJpl~[-eI)' m

3[ ~~llJ

or

{l

WW'J,I\r •.

sci encemag, org

5C~E NC!E

VO~ 315

1 6 FE BtR:U.A.RY2007

PERSPECTIVES
Rl'rl)l'ftIQ:'~ Sl~~n~~I'Ima~el'i,~b; hke these Im~ long been Ii. Stf" f~ ~7.i!fI1ple-, C!ll~~kin<!r!!n"c. ~~bt~~, P;.~ hO\VfIi to I i.e i iii. the bfli !lIl'Ce: betwee 11. [\.\1'0 F.rim:iprtI,1'J/ Ulfld'mmlliMtttr 1"izy'5ic-~ f'IMlbri!1~1? Unm. relk:,,~~w{l(fk O:Vl~[I~I,II',Hi ngp,NfNli ll1!el:!."i ".11~ e3\;~£e:mL;lS (3).Eilher I]il~ ~3liJ~~ I'hr!fi!l ~I!I1 O II'i'e!S, (ambrit!~~,IUK, :t"'~~. z, J!, A. !le1l1, .f!hy~. I} !!.Ai' ~'~6~ n91ril. R>!I\t m~.g"l]!cl.s has shown ql n dmt ~md!el" eertal n ordered mag:l1cti,e SI3:te. bUll'illg tlre eondac3~ P,.~. rllilll~.,P~, JI!i!I'. 848. i:li.EI~nml eircurustances, y'el to be realized in a real rien electronsllione. OT the spim, and conduc'1. 11'. G~~~I1,el Gf.. ::;dtfl~ n:s. '9'6~.(.'loon ~ll!l!lli1:rii1IJ but m::\,Efthelll~lO C:'im~.ruiy pia.usi.b.h~. tion eleetnms 'l:all fLI~'" ~o cn~;ue ,,'IJIlt::[f~lli(! s, IM.ll MiI~hllf ,i!'IIl'J~Ho!IluU' 3194,39IH9a). 6. Ji, ~~~!O!1J~ et ai" P.I!!•• ~ it'l!l.f;i:il, :2.46405 f.2()(i:3~. state oj'<Ippm'eJlIlly Iteiwy eleenens, Usually it exchatiens ~P'~llr .U the critical POiLH th.<ll t. IP,(oI!?IiTI~Il, , j, SdlaliElll../'mille 4B .•li16 f2005j. A bear no .fCscil.nblillltlC 1.0 I.~Cflli~~mu.iolils ~.m. the is fI5SlII:lnt .. Ih:.!H tl~~.second pruCC5S hsppeus cd s II. O.lI'r_I!!I~ ,..,aI~ ~. ,~t'¥. tt!'i~, 85, 626000t:l}. 9'- G" It Sl~r!, R~ ~a'. H.i~ n'!9'l ~200]J, ordered IJil<lses. E;';:lenCli,ng ti'li!sideato ~he m1.d is rQI~IO!I.\l'f!dbY"1 weak maglli.et~zaliuUl .of nH::m~.rnlligB1t ~lIi1m~st[h.,~til i~the fu:r~!ik~npor I.~e re[ijlt1.b.lluln~t;IIL These experim€:UIIls (l[JLI3d 10. Go, R. SUii!ilrtl'li!il: /:lId&..q!j~ 'l8i7U (z.IlIII6), U .. T. St1II~lt A. Vi!I],wallat!i, l.II.:J'IIlfI!!, ffl.1':A, fii:>h~t, the: e I~ecllroll il:st!~fthat ~s. beil;lg reflected in [his. SliJggesl thi:ftt~he qll;puMllliU a.i.~nctll~ poinit is II.lot SO~~~mi3:, U~Q (2004:1.. :til. f. S~H~!'"fU/',.f'hfl. !'It>v:~~~, O:li~!1l(~OQ4~, nf;l\\iefleL]l.Y seale (12,. pf~marity o"b(l~lt ni!,~gLl\~~'~COHile:1To"t <lU bl! t n S. 0iJ!1!M1l,.f!~8~!1. ,BltI9'r.l)' AI~ematu\'dy; this !:Ie,""el1'elr'~:-;.cak cOL.QM r.uh.er isa rransinon berweenthese two dilf~~ u, ,p, COimtdal,. e~oI~J. ~: (om/ws . .Mal. H, 117.2'3 be .. [etliclI;;tio[l the lll]lUIi~Ull! IIl,UHLre tll~~'hoe' !I;lHt r~ulel\ of the :;~:Ulilt'> {i·n (se!l;llh~ ri~u;m)_ ~il:OOn me1~1l.k: 5i:aIr.! in YbfRh:!Su~,.\~:h~.chh,t mi.xlliJiOC \V~mc\ter~hc tmdcrll'.~ng cause, the ~h.C{1r.iSis :lS. (ii. st s Ilalt.elIil, ~. ~jjg~i!ffI~, ~.ILl.mii~, Flip. ~ II ,~!!; i n~l!B ~ZOCBl o:t' I1Il;~g~le!1Lc a~(']!l1il~ I,ilile yuel;h,~l~m l~,~I!I~ed .., nnw havea clear raM;;; U:n!<Qv;e1 .he :i,clel1!t~t)l ill of

A mnmber {)Frocel'!llheOiFe~ iC,l~~ug~es~~ons mli~~nlPuim to t",I~.1L~.\.'I'C have raissed, TliI~o~

or

nlii:~d of mel<ilm(;' tlIIlCI.WII1:>.

The rm.l~of

t:ht:

dn:new

efl!.'lrgy

8::.,1(;.

ANTHIROPOlOGY

Some Li:ka It: HO'1


is p:r(l{hllced~ntheiuUH~ of some Cal',~icum s~cies <IS. a deterrent to seed pl'ed<lwl'S.G:l:eat \'nrj~~I'i(l!l1 ill t,lp~,liei u!~l'J!!11IJeJilt .~a.~ beeV! i:iltro<dJuccd dllro'llgh pl,"l.nt breeding i.UlK)Cli.hiv~l(!:d !1pecies of peI1per)l,b,ut wild srec~e~Oif Crlp,~'it'flm' 1I1.so haw hot l!UfI(i ~.nildfbnn).; (.. ), i Anl~ru~.. t\ aad \;~re~ili!tm~m;:e~~t(l p],.~~e~:>''I,!t;h Humans firlS.'t e_",ploiu~d I:~li~fi!lewbnlile i!l1 [~,c Americas, and Emtlpt'9n c:.\p~.on;n;and 9S [mliiru and Th~i~.:!Uld~r~£.r Em<opcans (.doni:r;~'S1 mer transported III~s and other New explored tile New \1!lou-idi.n Ih.e .~5Ihoel1tury. On I"mg~ 9S:6 of ~.hisD:S~lI(l;. erry ~I CI[_ (J ~ \Vorld p:Jlam'S all ov~r lh~ world, Bur (lx:m::l~y P w'hen .md "",'here dnme~h(;,~lirm of pepp~f)i shed li~lll ml\"'~l~n,md W~le~!'echili pepllen; :flil'iSl occurred hrwl.: proved: diRlcuh to cSIIJiJb~ WCirct first Cl1hiv::~tcd. nam frum'll S~QUjk;5 of ~isl'o ,(4). ill pan duero <l lack Ofnl<lCl'ofos'Si 1 lhisk~ml may also have polel11i,1iI use ,in lite reUII1;lim.I!1J''tIte:;€ 1·lftlpicOl.l p]mll,t~_ almlysi~ oflinmlil1LiiI transport aud spread of Perry (>1 at. UQ·()WSItOlliv that pepper5 were illliV:3sive species, Orpsicum is a gcnWls {'\ol:np:ris,ing a~mll 2::5 Cllhivl:ired ~flJl! 11'1 h:les~read mit; across HiC w speCies (1}. ~I.is a 'Ullemberorthe phll1i~ FturnHy A Ilnel.'lc'lS 6000 years ago. nor on~y as oeea ~ So h,ul<lc;e'.'e.\",41~.{:h C(')rut<~ill mh~r ,~c[)n.{lmi· s !\Im:ll<ll coruJirnJ~m;s. but ~lbo ,It;, compouenrs of i cilJlly ~lupon~nl plaais suehas ~I~c pOllinO. the a complex and i.>{1pJ~stl.cml:d dic~,The all~hms ~OLlm~(l•• mdtobacco, Bm..::i~ is the oelliter of reeevered mk~fos~ih. {Df !!t.1rcl~ gu:,l'il1l::> flUJ.n grim:lil'lg st\m~.,o; mtdcookinlg PiJts in <'I UlChaeO's;pec.iles di.vers.ity fOl: CalMiellm, b~u 'lJ1aany ~ .. ~i~)l .u:e,~~X{~Itmnd iUlthe t\'L1d~, Hli~n>.In~ ~.()gic~::oiteslhm1 '~IleC~!J.~bbea!'JVe!ilez;ll~I<t. mId l~l.c Afll.!{'!S. Th£)' (hHild C{II",~iC~mN>pe~ h~N"(; d!HlIl!.>5tuCfltcd <li.lIdtoday <:1iI.~.~"1II.k :U'u~ ~ ~~S of Cap;:.j('um. an Ib,]; theh: spic)~~J.aVQr c~riot; ~<lu"h gF-li m::~n a:i\SCJoCiali"nlw~lh fl.laizl: dJ~l ~Or.ttlf:lS Ih)llli! th~kIiIlig-d11~inarl~id~ C;~1Ip' and illi,~liJioc_ Th~ir~vi~~n~~ ~.ugg~MS ~lmr

SliI,l i)e:i (;If rnQ1l~I.lypes Qf m~(rllrQ-:s:;~I~,re\fle~l d na'!,!'patterns c.ml c.o'l1Ifl1ections beti;','e~HI human moy·e·rne·nt<lIrnd tile di:ShiDlJl~on anti rn~~em~nl

,cd plarwl'spedes. bGth domllsti(di~ed antlwilcl,

"m YOU i~l<lgine [,lome c.j,fIhe. :~rem .. \~ )111<1 '"lI.isines.-Sltd~ as Inc.iilall, [hlli. .,~ndKt~r~~lIi!-w'it[~{lut ~~hi~ ers? i p~p~ TI~isfic'ryspioc has ~llineml.~n[i:~grfll~ of ,,<]r~ oooking,Lud cluh~!re lihr from irs nllljve l1u)ge. Chi] i plippen, {(lj'i)Sj:'!:tlm)' Col'lle: fwm ~:ht

D'ilrer:sjty e~.pl~OIinedl. These dilf~elll k~lli~h 0,1 C~siwrn, .grOOIn at~he lInfvlililsily ,of Wag·eningeFl U!), iIlu~tr~te lihl! d iNo!)'fsily ~~ shijlpe ;;;lind! Qilloorilni dOHneslicatet!! chill p.eppern.. ~rrrv e~'I'll. U)sh Ow UiI1Il
peppcors, lil;lllle i)~(!'n

~li.Ihi"'~ aCf(loSS I.eel

lil;!! A.n1~ricas

for at INist 600[ll years. mncd:i.d ne· {5}, OIlier ~.nemboe:l"S0' f t h.e SOI<llllM~ch.,;:;: l:h{~!:!l<J;rph: and tOOOA;:.{:{l, .l,u"e ~ <,J,~£oh1Ld chc~~1"fn8tUW dli$l~ribll.ti:o.l'I::;ob::;l'ured.·by Ilmrm:ln n-.,rn<SpCllil. sc:i:eilnific :11~lUlle[)rlwm Th.o:;! , 'i/V;IS gli\o\~1!! [{j, Illf;: tl~mmlrrl~li by L.inmHm!'l
otC,\:,le,

'Jg ~~

~l~i!;i~. S.Qm.e\'<!!ri~!il;:i[i, (Jf the

c~llti"".~

~ecjes

t~~r~~ Qf !he fi~ spoe.;;;i.e:5 of d(1meRli~'Me4


{'"atJ:'iicum '!.'\<'cre (:u.lll.iVllltf..'t1 ~ogether iilPcru hm bo~:hthe· O!;O\<Ist <ll'l:d the hl~hlallds as Im~g .[IS .!10m) .~rs l!.gl:l_ Ai> h~'lnl'llrls moved 1~ru!inllldi fiVel' dt~ 'face aU of [[I'i~ Eanh .• thL::j' C~Li1'~Ii:t1witl1 'thCJll 1.l1l.:'!i:r ·[.. v>o\l·jil,e foods amI. h.el,·ib<l~ l1rledkille~ Caf)... .~kumi~ !1~ll~Jb~e ill thi:);r~gard ,xs it q llick~y bL"Cm1l1l6 j,n~!:gri)III.OH wide rnngc ofmd. \\I'oli1.d c1i!OCi p~ine;:.>.~~[;(l'n~Uldj,m Cl!i~illut to "['I IJet<'l~ I VOL. 3150 SCIIENCIE

(SHCh. 11:" bli:lI pePP!.>I1i, lack high

of c,,1piS<lkin.l.mt tille·se'Dsoll.ionofirml'l.less aUld tile "C::ildm]'iil.u.~l rU!'ih"inl(hllc~dI by t;mlun!il chi~i~, I~argely :f!'{'''Colllm fo:r. dteLr H~iI\'t!rsal a:ppc"<IL C'al)!o1Iici~'i i:-:, I :\p~~i;jti7..£d blIhll~.bl).~;'te ~ 'lhlll
qli1'i.i~n:it~.jJ~

from t~~ Sa!1iM!;lFi~ "dh~~~w:r.iI."b~l~ ,iU :'ip~cijes of O(JIlUd H[e iIlJ f1~t:'t n I"" 1~~d""'1i: to lllL~ ",. oJ Amt:lric."I S (,6)1, Wh,~~is.,. Ii~,:t:i\'t!; .I~ll~~whe-nlliiun~mil\ll:ml'is- il ~ € purt 1)I~ms $0 fillj"liooln I~u:l:il'origRl'II$. ~md ~.llCI' .~ '[h~it~~hrolil£b ~t:I~[k}l'i to S~lillh~ij'rn1l'iiipur.

S !

11:i$I(If'~Mw~eum. (I'onVl'lI!U B!oad. [a:ndml1l S,\'M7 S BiJ, UK. ~~iiiIa'i~ ~m~pL'l@ifihii'l_~C_IJk

lhe i~l!hor

~i.i!!1I~!I!

O~pi~me-nl. af IEOI!!!1y. Th!eNaM,~l

pm;e~J' Tod~~ lXlncenl over hl'V'dsive ,"pecie:> <l~d 'he~r th re41t tirlIl1l1~,i veb,i(ldi·\,'~!'~it y I( 7 ~ ~ &n~gl~~jgh~:ll ,i,ml)tm~m:coflDndC'I'l>lm~ding ~ ~Ju: how 1t1l 1lI1.U~ 1'I10\'eIlUl'rlts and Il'<ln~lOri h Olive ~
'0;

116FEBRUARY 2007

'!IiIWw,scieflicemag.org

PERSPECTIVES
aHocted di~l!fib1!l:~onHof pl ..nts onnd ,'lnimO!!ts,
Src:lC~,1.15 {jrp,~an;t5 i,lillipmhn::~d for ~Lv8in,c:l~y of
.Ulj'

of the sites Illey ~am:~!l.ed. s~m'l.ring~hat

,re<l!~"m]\;hOl\·e cn!'J1L;: to ~nwtd~ et'l(j'~y.~len~s illl !tblir1nill.g~II,!~tyS: witn,cs.'OJapanesc kl1lal\V'0:-d in B,rimiDl ,o:rkmll\! im~he sOllitherl'l (J]!l'i,l:edSlates. n(]il11l'Slic:lm~dl phll:li's ,n'I\,'~ aJW!I}'5 bem1~a!k@Il' by 11umans wherever t h.ey l'nn\O\clfa\l\cl,(;d and
.mn:'~,lf)8,tly

(but ItOC,~,hV'Jy!» ljll.p:mbl~mmk. bUl these pauems czn be mrocmg the :most dir·
f;.cIi~1 ()~. rill

dOlll£5~ic;mLulil of.' c:id!~. eppers ~1~~ necurred p IU~i\;liwf'{lre ~.h~~(;l~i~~ ·~'I.iEl1ie oo~~!.:!piedand [h.l!!!.r .. cu!ti,vadon was ~lUim:. \V'I~crc dO.~ne5tJcal.ion of the fivtl, species o:f C(lpslcmu occurred is currulUly !o;f!~Li.l:ati..'4;i'; mls(;d Oil modem di:s:tri.InJlio.~l i?LrI.d geuetlc aml~)'sis... C. amwum is [iH:mghl. to 11n\'c been dume.l>l.icatoo in Me.;,,~I!.:;o or 1ll0nilleIal Central Ameri~1J_ CJht!r.t.U'f!NrS ~Dl
[~l::

lhe~ wu II al so be U9I.Wlil.ooY'LmJ nhc l;tmJy of 3 few CiU,p phll~~s.. TIliiy I~OI\,'oi;l p()H~nti'll ~!O, he:lp ill ef~t](rt~~~)uader~he stzmd LI~eUn],:s between hLjm~J:l[1n!1llS'pon. and ~1I1i\r.ll~i~·e species thus (fln~ritmlinglo !he dlal le.nge ufhimli\ '€I1;,rry eonserv a ~iUl'l,

nopjc,!I origin,,,, [};m~like

l- 1L.~:J1I)' ~ 01_,

Rt:r;e!'em::t~

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:ns,

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of

(lis~ribllJl,timl~ mll1wd_ NIo""\,\I'\'!I<lyl-i to studyingflndel1~ hLlm~1fl use aUld U<IIl'~p<Jrl,


of

Olriib~:t!Il.

{,Jjbi(!l,rS(! ~n i\l1l<;z.uJli'i.

C
l~.c

brlL'"("(1liWI ~Ul Bol iV1i,~.•

and Co. PUb('.\'!-'(NI:,:illl

a fj.. ~rbma. L de B_ BiilliIdle~ii..~~. 8m. :Ill 6~~: {~.{IO~l J. j, J. T~r.t;s!l!Ji'I' tl Qi., t: (r!lIWl'. fCt'i1.: :3il, 5417

'Oo]p~m1t" C<Jl~ >(',{)Jl!:!iitMe W oUlrk!lm,\'le4g:e

!'iimlth(;lfll Andes, C. b.accafNiu ,mcl C jJ.u!Jcs-

Wlrutio~.

eees 3'1.'CtHiiwnomic'1Iilydislim:1. but the odlil.e:r t~~e ,in; 111~lli1!Ibers f ,I t\.peci~s COrllp.i~N <!I1d o perhaps .nOl. reel Iy ·"w.itl(l," specnes at aU, Umwansh<lve. un.a vefy:siIDrtt.iarle, md~c<!~ly u~tercd both ~1:k: ctu;JI.·m::lif.:.~ri"licsand distribe(lr]p.1ni!'i!1~~we V.i~~I,e, ew ~l\;,n,1 N .rJtm1'l:t'i,k"tt~ C(lp!iik:,~.rm,.r'e\'~.ds L~.ml;·eO!L!IIIciem t imi!~ nf ~h~ oC~ldviitiom <lI.nd\'\d.d,e~pread LII:SC of thls l:.mp types Uk!.: the Slf:fII'c~]mh;rof()8~5~1s di.SCOVCKiJ P,~"Il!I'h<lnpn:"'~i)m;,~y reported It also epens ~!P lry'PerDY"'Ial,.l~"''!; elftol1f.!lfI1!,; potell1~ial, to he~p ~I:t:InIltlW .a'\l1:'m.l1l~~. ()fn:~!lirt:hiI1Jtoh~)\vt~e: r~()p](l~ v~~liga~1.:l dx: 1rl!~j~t·tmDllifi:)tdmtl.~<;ti~~i}[}fi. ,cuttivaliQIli.1UJd trade in a wide' variety cf'erops ofthe Anne:ricas trnl'lSporied and traded plants <J.f (:ulhLml hlinpt)riHtll~,. The' 31ll~()f:i, fOtLnJ .i~() \",,'h{J'~l~i,s~mh::;hilvc mL'lul.~IH.::d t1ilf~c'lJh iu sti!i.l1Ch gl,(l,ins ofwHC] species of G~l'skum in. ~mm\veld~1eh.1theUr lack ofpresen,I;ltioll ol'~Jeil:
the' dymlml:cs or :iw:roduClions. Im:lud ~11,g the eft'i;lcts of illlV1~sivespiil'Ct~_ Il'''':;f~e',~tj/.:s ili'Ul(}Wllivc usc o:l:sum:h gmins, IriJliIlki;'cilen tools, ,'()upled with ~heir eh:gaw Lilll1l'l,(;li,ng of ~!hi;; .~i f'i.dty oftltl:.'l'ic gfllill.l~. ~o s

,~- 'B- ~dlierJ.l!ljll,ejq(,.!l,'!J~8flJ~~M~a~y~lhe' i 'Sa~1JrfQf.l', G. !,(;Jr,o!:e:;, it frl. ~~e-r. A. il. S~eto!!j~~,


IEllIs, {AGi,ilemi!!.II'le!1l, ILoodom.. 1'l'79~ PIlL 6;;'9-700_

S. A. M. D~, G.1p.5N:WI'l iJ~IDr & Fr~rKi!, lomlcm. ;!!1]1l3}, ,6. Ii);,~, !l~. L l'ik!liL ~n s"ta!J~'M;HfJ, J. G Hlw!i~ lit M, ,~:ilillf. r!'I, 11'ft~,N, lSlrad~ ~., lids. ~JLO}'~I iiolilnk:

7. Wltemniwrn, E(,t)~P'fIem Ps~iitl!ml,Ern~~mstlJ'jill'


HUIOOi'lIWtINl;riJ'jg; B.irJdiw!rn~:y S:~;115.i~IlfiQrkll

~NJe~s,. ~, U'l'-Ol~O-

Ri(nl"!lr;lntl..

Sum:w, 'UK" ~99U Ilfl'·

IR5tli.:!rce-s IMHtule. Wa1!~I~, IDC. 200S~,;::1fti lJ!f!'M\.rn~web.~!doa!~!l1}J'dowmiffit.~~~,;)jp);.pd!, s, t:na"" ,. Upf!f_ &\1_ !S:3, 2001. (.2oml

NilE

U,FtO SCI E INC E

ere· m Wh,,···· A'·"1",.


I

Studies in rats reveal tih'.lt. .!h eokey to irlJlerpr'el~n!J !!ipati(il iJilforlmlllti(lln-----wIt~t! \\1~ ,aril: (@FIIpoll!"ed t~ ;~h~r,~we'~c aiflE!\lIdy bi2en-m'aJ~' li e in the' h\,ppClraIFl'lI!lu:s, pkDhc visual or:!llL,cHlOry SYStems. SoQ:lnc'~lffi\1 sp,~lial kn(!~~'It1dge .lsselnMecl by the bmg~, is On rag'" 9M (}flhi~ i~Sll£. lmngla1b (_·t af_ (J) .. provide ~he ~a~~Mil1~ight i!1w hw SllOl:t~.,d infu"m~Iion isC'oJ.tl!Pu1cd and U'amformcd .il~to sp,~tial
,U,"'ffiielJ,e;ss. 001'

\\r,~!~~:S Ol:re 1!h~nlYsn(l\1;'ij~g -[Vll to Y{l1l,"


\1i.'C

I~e G.l'Cckphi,h)SOI)hl!rHcr:at:Ums Famously observed "·You can never


SIEP

eclls. These findings sugg.'C~"d l~1J:n tilL: CAl region IIILn~)Sspat i.di,n~o\1.'m<lltOn ~m(l categu:ru~'_~,( in~~lb l'm,l'l.Gird!; or .s~~Ll1~m caWgflr·.~e-<; Thu~,.. <! rat ~leWl.~i~i~~ i!\'IeU to be i n l. either a drel~I:l[" or square btlll:. aund the 31PP110" ~)riaJ.e :spatial me~l()iJ,"ygets <f.(tuv<u,ed by mechiIU'IIU:;;nlis. 'l~tr.nc~u:rlliletw1:Irk pmpenies... Th.e will~. (:():~lc(:tJve~)cti\filJ' j,n it m:UI':fL~ fl.cnw'Ol'k defilt'f.'S ill~ SI:tItC.An C(i~ i~ ibril.lIm suue, tallied' an mtrac~ ~()l':is<lkin 1.0 a menmry (j'), /\;ttrnf.1m neW-lork I nl"."p~](L1s.e!> ~UlPlilil tl;) Hr~ IO~'llU~to ,I han Oil! O;l. InImp), Sll1irm~e.The ~eEM,'o~kmtcl ~I,u: haU qu.ick'l)' ~~~~e flllto a nearby a~~'L.loC10T il tile lQllI iiu,PUfS change cnmlgh ~oswitch <lHfractors, :~ 2:nQ5.L,elqtg~b OAlldl ,.'Ic)lh:"';;l!.f:l~" (6), h it,ll n { dUric 'CSS'!.>llli,aUy tht: iHU'l,U; cxpcrim.cl1ii whh IfLOrpb. b01'<e~.b~u lI~.S()'r«orcledlb:l'IlIl CAJ. lI'te!iJ:uppO(,:1~Nlra!.re~i[]ll1pWjectillg ttl CA 1_ F'nr tmknQwn r~~l~QnS. th~y got a j1[lI!l1~,\'~m di 1I(;"reni[(o"$uh Ir,mlb i_h(1~adk, study; Glll!bll~

new Hm.\I' dJo recognize a plaecas the SHm~,even when


fi}[

im;~) l!lle J;um:~ river;

it is dilieremt'! lEo\\' dbbl1l'~IJS 1.'(H;ltlime,ly activ:m~ the same


representatiens

know~ed:ge, lilltmgll dis-

ti.JlIG~Iilf:t\.\'lnkf.;nf:I:I~Llmm;ulli I:ht':hippm::mlllplIiIs. U:'iul:geb ~J (JI, reoo:rdlcd hippocm'lllptl I ac~ iv~ $omewhm d:i,Jfc~lmt experiences? Whc:n is ,e:xpc:rh::nce lh.e same tM dilfr:(;rel~L. and wh€:111 ity \fhHe rats l(lmg(;J ln SCv.eril ibOA'£S [~at sy:;,· temOllic,llliy '\'3rk<l in shape be~ween a ,ch'~I!e is .UI. p~<IiI,~dinerent:j just ~u~d ~q~Hlr{;;_Slmii<lr "nl.ol.·pl~ hnx1;l~"w~re 1,l. N'enm~(::uI;!IUu~t~. !)I:~ .~t~iu~~ dn~e;:[ ~(1 00lli'llnDll.g ;U1S\\,'Cffi by r,ooording ~he activilY ,oF prev.iolJsly us£-di by mJ~ersto fi.'0md I~Oln CA I hip· nemoons iu II~emlhippoC'<llIll!p'ustll.!<ll s,l,gal<l~ al~e {4). the inRlrmmioll mMput l:egi.:C~ll

~mn. e5JloI1!>e ro £

'1:~. .

anhnars ~()C.,ti,OI1i, Ollle or these ··p;I~()e oee-l:ls" l J j onb' di:;,r;,~h1L~~ nlpklly \vh~n lfuie~1!1il\~a~s r in a s:poc=ific pan Oflhl'! ~:i.lVirollfl,'i,ellt CUr[1S~ ~PQndil,g 00 the cells "firing riel(j."l'he CQI~ Ib"t'li~i~ disdl,Lrgillg o:tl'rhl(,:~t::eUs il'IlIaws Ll:' W p;redict ih~ r.m\ ~(l",mgon {.J~ hy, il~ fi~~1"e. .1iJ 1'C1.uHng i(s~nind. KI~.fl'\i\,il'ig [iiI iI'".lIf's Iocat io:n. fr(lnm~he",cl;iv;i,ty ofii:S 11.el:Jrons Iis aslonish UI1IS. !l,i\'C;':i~ ll'l~lt ~:ts. luke p~~npl(l, lilo sp06<;ific h11\;~

pocall.lllLl s, 111c em'Her s'~ucly foUIJU.I that a :riU Ibl'Dn~ d~!'iIi!1.ct~euf;;,'lI'·~rr~:;,ellwrioll'~'· (p;Jt~ tt:rilliS, OfflCth-1Licd pi<:!loc cdls'\, hcll oOC'WI:pya.ng

on'ne

either ~ c.ir\:ulm- or !\quar-eb~ " Nei~her the G~lds nrpl!J;l~'e {.\ell~,!lm 'thL;lirfirUll'I,!, {m~li'll.i.D.:jf) ' m~~:;;,!f~ !l"elm~~lI-th<.!~ i~.~~wre ~s ~O!l<I~

ffinlappi[lg ,n.flt!.:lJhJllJiI.1 Heti\! ~i.yin CA I wll.r;;na 'II<J:!: mO\'es, between the two d itlerent hQX siiialle oenvinJll'1liI11;'Iiit~" M,(}I'e't"""er,. rm,ly'the cir,d~ or'th~ ion ~s.ilCi ..i,V.aI.edfo:r ~pHti111. sel~5e o:ril)m:>m~lIiogou:ll 'LQ, ror e"\,:!lif:i;~ ~'llIlmll'(! rlcmal rep:re'sel1,l2i:~ :::'I~ I diu;: il.l(Jii'phboN.li~'i1~)I.:l!i:~ Iboxe.~ ~hIT~m more a Th~ a~'~!~or ~~ ~!1 .h~ [l~p.iri:m;e'!i'!t Gf P~~ili:lto~~ il!nd d IiCle"-li,keacl ivme I:h.ec.urcle re~ue!lentation III ~h .. rm"'~09)" !l.'W~fl F, Furdlg(llU: ('!:nt,~fl!u Nli:ur.ll .aOOI 'the; 111~prrx:.unp~!\1. \vln::reOls ~qW!,u-e-1~ke Lm;o::e:s: B>l! h"'1irl!ra~5(iMte. 51"t~ 1J1Tli\!~f~iI:f rl!f ~eW ¥or~ :Eleti",a[~ the sq~I~'1X:reJYrCiit:liu2ltion, T~lC fOC:'livl'l.·· [)(lWIi'I.st~leMi!'ln~all (e!nler, Brook.lyn. rrI'I'U2'1J.3;. USA.
['rni~l~ ,~r'enloil'l dowi'i!,I:t~:M~ @

I:eLmppul11S occmred ala pan icular 51.. ge in the'


1I'il.orpll1im" ~ql!ll~l'it~li:l~: Quly ~ ~Lhb~l {]f cc3 h (·'·hJlnper.;") illl bo~h !ll~I)POCllll1ilPl!l,1 re~iol1'~. OD.ifltlr li.t::uran~Jl ~1.uib@L:ls. ti<o!I.':alll~'d iUE;f);:' s "~pl c~mn~d by r1llie 1'e:In'IIlPliL~~sy~I!C'nl<llI.!k:<1I~)p il1(:rell ~i ,!;l or dL:lc~>';;1111ngt~~~ch1u:rge H ratt::s ael'Ul:;S 1]1l'!Illi1:1rp:h sctIJlmi~ec whi I~ :mahnltl.ini~}g fi!rJn; rield loc1lilions ~6,. The resL!lt SLlg-

'tiorl ,;,;.Iate ch .. ;e!\ m org

i,,'rOhre;re~uly ac~,'OssCA.l
VO~ 315

'WW'!I\I,.sci encemag,

5C~E NC!E

1 6 FE BtR:U.A.RY2007

PERSPECTIVES

01
----- - ---- -------

clu!.'led

Sei1~ory 'imICllil'l~'ti:lm(~C(ll!e:i)
ill tile ~J~m

hll:ge d\,l!1ges ~fIdel1ltate gY,L'lIi'i outPliI![ m CA3buL Vi[tilllll!~Y m:l eh'l:1.Ulg).:-slll CAl outI If.nll~nCAt

:Hipj:Nl@nlllal

pr(]ocmsin'9

1111$pati~ 1 inf orni,HJon

\JV'
,

I~ ~'" Illiil'Y1. 'inti) flIeuri!~~brlg~rks


D

These hllnpc:u nnd spl,iuc.r behaviorswere also evident il1! the temporal ~I~me~s~onof
fiWC:CiS!>ul1,g s;p~Jtia~;,ll1fOrnll,llimL h'laddi~im'il ~n

C@n1;ti:al~"put to hipJlocarnpu:s

n1.l

Ilelll.tilt~ 'Q!~lrWi~,

tl

Sj!m~ rnilul in,f~~ijorn.: (!t'J'OO"~dijl~lI'fJtlJ~

w
Sp.iI~ij'I awa teM!~

ICA1~~~

~11'J!!' irlf~lI1'i!~i(!fl; efl(~de-s simil<lritie~

CAl ,"••

~nror:rn ~lt"iii1i~ '~Im;erl!!!

~).~m.cc fi ring m~:es. ~he pan iculm' subset of cc~~ p,llfLCCi: cds [lilm do ordo I]u~. fire [:ogcd~cr also 'S~gmds lh.e~Rfs posirion ~.l'),. 'flu: Iikeli hood u1l1!1<Jll'lir or cells w~ll film ~[:lgl;:'11t111tduring1! sl~ort lime \!.d ndrnv is measu red by thdr CO/LCt~\ril)'. The diflb-ence between the clmc,u\'i~y of'paits of dent me gyrus cells STC1,Il'~<Qrger as the ",h,.pe di\;t~J'liCl~(m ~~twlilell~Wob(H~ occupied Izy~he rat lncreased, The &'liille 'WM ~n~efOirpu ~rsofdeHm~e gynk CA3 cells anrl IJllilrli of ~:;cl~s iJhin CA3, with ,. sm:pr.i~,ing w exA.~~ti{ln,'Wl~eIll, Oil !"<'It W'1.S m~J\reu fu etween cwo boxes at the extremes of the morph sequence ..coaeuvityin CAJ was 1lii,('Il;e: simiI~rrC:~l~ln~illai{)b~l'"v~ w~~m d:U~~lt ru:p~m~dly ' oecupied Ol'n~y~, cifi'iC~~ar squase box. Wh.en or amerph bDl\:. \i{~S :~nill:b.mlljly different :fmm the' dl'Cle or SqUObl'e. CA3te~lP'Orlli~ diisclimrge !>~~!liL:.ld: lu I1'iP,. ,ert~ph,.\sizu~l.!,t s~Ilil~II;ITily
~{l

10 etween the t we esv iro mmel1t:>.whe'fJ;;;i)i;'> denture :r:::yrWl~ dischruge seemed QO :;,r~it, emh<lndl~,!;l: the dinerem,"e, The _"",C{lmJdJ~eliln;;g.l:'!.[Dpn m!1::dH!n~s'nl is ~I:()bal renl~Pljin~ in response 1.0Il'il"gc enviromillenl:al '~::tmn~el'.De:lltme _;Y!'l!S and CA3 re~b; a.lso diOl.::reJ. Global ren~ap'ping ill CA3 aad CAl i~ char.u;:te:!i;!ed by a ~e~m~ingJy random SdOCI~OIIl of'thc' aCl~vc slubr;cQ lhllllkl~,g lilkea rat Stil~Q\rv inr omnt!boJll about a ~at'~s!)atia'l envirQJlImenl (dliCldar QrsQ,'Ua'['t bcm Ilr an ifl'llofr· liiediai)l!-~ha'j}~ bm:)f!!~dsi nte Ihrm~ hip.fMJ~~mp<l,lrn,gLDn~ of the bf <lin (d~nlaLl(l g~m~" CAl, ~Ild eM) hOln;, the flIeoc;Of1,ex. Spia1Ii'!'l illf(lrn~<lli'l!1! BOOUt, ~tiI!en<!1t.ure of lih~a!l~tmill"5 e.lWirolflment ~ e'm:oi1!!d ill tl'Le a(lt"'i~y paUel1il~01 iilellrOIll rle~li'IDrk~,n thif!~e ,f'e9 iiiJrI~. i
1

of eel ls:
off

~",~iU~dlhm hippm:a!mpal m::livil.y 'O(jIfl~,IUill~ iln,jorurnathm Ih,m!:lll cl1vironmen[ is, bOlI~ the

s.tlffilll;;LliI.dJ different relative U) a pre ...iom;.~y i ex perienced and rememheted e];1\'kan 1liI!!.n1. SUI 't6'rusposed another puzzle: To ~XH~dCI eidtcr a "same" or a "dii Ilb"'"illl'" sign.:!!. tlte nec.\\'{nk miniM OOrneIKi<\'I,' eoe ....st i~the two correi SlpOlldilligiliU rracmrs, 1l1~S aecessitates mechaui;;,n~s ~eg~!il<!~e tmnmma~ to n ~(:ti\'~'Y imo ~hoe: appr{)~xlil:ll(,I.tmlUPCtr~!fnd SP,UI[Clf' s~lbgr(XI:PS, ~;'oIl. There is £,QOO \!l1lders~mli~g ofhow spa~ li~,! inlhtl1!l~lliori!u~..~~",~,()!;;:i~tt~ stored i~:1he ;J~d t h!ppot;.nnp~!s (9', 1m, hllit <!I.k:lw',\'!edg~:~~e!l~ s
"lidl~:;;efulw'l1!:.Ys. hi thl!l pre~~lE :-lllilldy; LClLtg..:lbrt'l oi. rqpe!lLcel the mOr:l)h OOXcxpcri~ mf:'J'iI. 1.1Ti;i_~C~llI!I.~ r'icl-c{;Jli'ding frum.l th~ ru:l':" dt;11~ t,ne .tl.yms "1~<iI CAJ ,],he del:II.1I~,eyru~ ~s ., ! hippoc;;r~npa!~£i{ln ~h..u project~ ~t) CA.3 ,Uld. .iISlfl.1ought ~o Lrilnlsfm,ln sl~gh~.iy t1iIT'cr'~~11 il1~u. acth'it)' from the n.eoconex into m('li~'edi~~
H'a!)'

lim:tly ,di~li::.reH1lt" patterns. l~[[t~~h als hl~5t et reco:l:di Il@> j)l'OVi de evidence of [WO rnechali;i:'lilll~ by \.';I'fuii.ch.h.e hippocampus S!::grega~es I neuronel ,leti v tty In <lIs&lsslh.e S1'Ililnc3Imd difl'el'~
~llrHlllll:reor

in one e!w~V"omnem 'Cf.lllhl nol he ~~re-~icled from ~.I'"aCI~vily in ~1Jil,mhc!l'cUlviron,mml1[. tile active Sllhset ofgnuuJlle eejls was C)()nSI<lJ]~, The s~:rncgtt'm]lIlIle I~,\.... eel ereactive aCBlt~~ an cond i[ ions •. i nc lud i.l1Igeli II:fereunl-sizlCdbo;.;:cs aad rcems,
BCYOlJd these [l.'ec,o;l:d~l!lgsrl'om a.~i thl1ee IIillajot hippoemnl1l,'1J subregi.tms.\vI]l011 .DS now needed are' r~u brain recordings :[mm, ~he' neocortex durilrng the mOrJ~'hbox ei'tper-

un Al~ltni!ltg.liI.

SOlllle

oell,~ tum

.[I~

and olh erS~UUUl


gyUI$ Q,I~sd.'1:!1.[f1g~

defll.;,tl:l

~h~ ,~limJml\ ~!U.vurm:trtl~UlL The ,fl~lthQrs,confIrm I~.mlCAl place cells respond ~(l ::->Im!l~ ~i'e\'iations in the :rpmial envimlilnlell~ l~nnlli:Llg.ln CHI~el'"""Ortls,lhe stIl'1lile by !1eu~U!l,.1 diL'ilf,;h,~~~.1iJ .. ~~er!il~\wre utl~vecl. i~ tAl rrcgt!M,tcSS.oJ wheedleI' the' r~li was i:il it n~!J.pheti or ~!I'I.m.Clt'p:hed circular or ~1Ll~'I,'l:l box, Larger Jt>·vi1.lrO:I~~ frml:) ~ifh~r ~o!'U\'itorn~ rnlli:m ~1~ISiOO m~A;!ren'l<IIlPing in C' AJ. fie ,(j~ll~ tate gyms '1~~!; (llI~tedi:nif'!.:;'nt Single d,£J'ii::!rt~ mllls:i also :;egm@ut: ~JlfOr.in3ti{J1l lind: memodes 00 that they call he l,ecmn'bi~m:dulil <Lrbj,~ glrd'lmle cells had mOI!e firing nelds than dki

to clun:ac~er.ilz:e line' locati:on-specinc n iea ~iIl~aLl~H> the .~ip~~ n~~!li!ilJUSt 2). ( This win providea cemplere dC:;;Cfril.pl.i();[l ()f .ilmcl.'each connected Pflrt of the hipp,oltam' ~ l eircuit responds toO srat~~11 .. Ch;Ul.!j~~in au ani mOl~ ~!i!'iI' ironment. The 11), in ~he (l1nt's ment. ~iJJpeflf5 hl be l~lCr.il~,c'd .m:t.i.vc subset o:fgrlll,H~~edls, OI1l~Y2 tOo]% trf deni.a~e c ID'Il!,I:SCf;: ~I:;; <l.tte <let i\',~ cmld,j I iupns ( .13 )llil.c~ in ~lfIdi\'~dlJ<lICA.], e~~b.,{j[1illIlUIe ~Us ~spr:mdw IQ :!>llfiiaU mO,rph dC\li!ation~ in!he !<'If!; ~nv.ir'()J:l-· thos'!.'! Hs(;d by Leutgeb ,rtl' til. If slp,atia'~ i~'1j'illl~ do i'lo~ ;:t,cti ...·aw Ilic u'th~[ 9"1% of lhe IlilI~9U by C;~IIi1gi:ll~ 1J!i!th fl.riil1g r~re.,~ mtdfiii'ui'IJg ~ e~ds unpredictably. Thl!l$. the d~l1imt.egy.l.1Is denl.ne synls.'what do es',l Thi sis e~l)eda.lly pu,rj.\~\i <1 C()~ii!,Lli!nl\!;tte infom~;;tJiQI'l.';J'!Iliullr ,uld [lw,:zJ.ing b(;;('~"~M~ the demt.utle g)'TUt; g~l1eraHs mew ~lCtU·OIH. throughout II~Je1]nd Ilc CA3 n,o:r..:: of.2i, IUI1n,pc::r (~~c dl.(!' figure), SmaH chm,~e$ insr~a~i<lllillpw ~nfQm1iEltiolrn th~$ neuro~ene:-:i,~ is Irnlod~~lared by ,~eOlming
~~tI'C{l

iment

116FEBRUARY 2007

VOL. 31S

SCIIENCIE

'!IiIWw,sciefllcemag.org

PERSPECTIVES
Of). Funrre work \\~in sare I)" rocu~ On\~~lY IJillorldt)f 3ppru.~mJ)' the ~al'nc n.euruns seem ~o h.ave,u d!ifferen~fW!111(tin!ll. R!~r~~n:ces
~. ~.01i::~!:.1izy.,N;e,t'If!1.S:l, 7~ U970.',
2. M. A. 'li'!i1s~fI,II. IL..ri'lfN~lJg~t~fI, :'ioi."ltf.ht't ~61. llilS!5
~1\l'13),

4.U.WiLl~, C.l~r;
Stf.tM~~Q8..

ai',

~OOSl

IF.. (~~Ji,~.

8UJg~~]. Q'~~f~

u,
B.

~~,""f

~U.1:09~ ~~'.

~,I.],
Ii,,"

IH~prii~\ Pr«.l'Mfl A~G'1f. U,s,1I. 9: ~5~~ 5d. 1


J405 (2005)"

(l",a:;n.

", fi:. rtt lM.ay!ll3ll.:S, O~~J'.lI~'!y, M. ~, ~]I:le!'W~,I(], lel[ef)', Ell', J. ffewdf.Ci. :iiis., ~~()031.
8. tl:. D,Ha~m.I.'Gim~1!i. H. Hir<ll.i!,(i Dr~oi G.IIU!:l~'k~ 1IlG.1OO!4;Z4, !is:;! (200!3l'. 9. ~.I~~!O!'~{I'I'H'l ~i" Sd.e!!!'e ;):13. H4U;t.@(6)'.
~. G,

J. K. 1!.eLID]~b el ai., Mlrlmr.l MI..

!1!.. l!I. .MiIi~~,.~.~)Jbi~.J, IfMr~fod, I, 1951 L. {1'}\a7~. I!.l H. If'lotim. T. Fe 1iI~~iIlg, A Tr.t!'~s.E.. I. .Il'ICiSI!l'. ~ s, MD:5ef. Sot. NMft1~j:' ~c. 68, 'l' '(2:006,).

~~,s

U. "' j(~ Ch~~ eHrl,. #W;!OC;Wl'lPl'$151 579 (:200~1. t4. IE. GDi.!~, Ii. eeotl~\ p, laf!~pi!I,,r;., fieM"l, ':1'. j. 'S~~, NtIl', ~ruro~;. :2', 260 n'l'9 '».

1. [, iCLel!lt!Jl!b. s;.li!Ulgi!IJ, M.":B. "')J~er; E.. I. .Pl'IIlGI!I', $ci(!1lcl!' 'l:1lS,. ~iOl ~OOI').

·~O, J.IIl. 'i'!11Ji'IJ~d:,A !le)'I1e'11\ ~.

SnLi'l~,1it t ~ar,

COIMPUWER

SCIIEINGE

Ara5t way (If findiflg relN~sfntlllti'oi'e exarnples ln (Oltllplex. dalasets may !be a'pp li(aMe !n, a wide ral1ge ofdiffnmlt problems,

lluu~ lilP di'r~'Clly 1:0r~~, 4· orr-~e:J' ,ntd: Dl~~'1k.~~ S{}\, BH.']lu~ opti.~u,~1 nf~x~mph.u:~ th~ :;i~~ is one IiCl1T which the :sulml of si rnilarhies Qr ~flcl~ and lind l~C best SLll'l"llllilfrry of 11Ie.ir OWll Il<li"ier:infu~lr~efnel1~. But (1:n~Jen!~'am!il,gQhe point M1 il:seXeJl!ll'~~ <l1'1:SI1I1<l1>l ized. ln Il~e im Cr.JIVenlQ!I'e efl'icuenlcomlJtU<JIl ioual m~thml ruCII~i,resHbit meredIar;i. Il~l,m~. du~t>L;lring~nmhqds (2). Ojl~ decides ~L nU!ltl10d~.[0 mllanuze and :I.l'1~I.ly;re WI~(ln :~I·. S:u.ch Inlc<thod~ st<lrt with the COIl!:StU'L1Clioll p.dod on the' l1I~mmber of exe.mnpl<lrs., aad thel~ de,lHng wiil~ lan.go, noisy datil sets" sciereists l I.r~c;l;to fundi them by i i.cnuiv~ :r1::: fi ncment, ofU~tl us..::a C()I'I1iII)~.t~l:iiU:t'm~ mC1hnd thm ~{Jdk:;;. of a :.<;imi ad~y rsauix, 11 l:.\'ible:: ofril~mlx:1i5 th!:!l esrabl ishes t~le reJatron:;hip oJ em;'~ da~a poil'l~ st<J:rti~.goolml <Il'imd:om initia i choice. for data clusters. ln the ease of' gene expres:;.im~'Wul.l~ [€~'it;. Orl~l()u~id", of~qu'l:tliit1i;ls. fm to iSV(;':!'Y ol.&a·er d<!t11 poian,t. A~ W'!;l,~"1~\" iill lb~ Th~JfI1t:llh[}dl 0 f Frey ~1ridDueck, ealled example, rhc ,clustl!::r:; wouldl be groups of SelTillfLntle-S at[lji1ilp~.e. S~.8,. A ~ is. .. mmmber thm 11:ffinli~y:propmgatk,:ll. does net fix the rllll'l':i:ber mea;j\~res how'\\;eU .:he od.m.l ·poiL~lArepire.o.;ents ~lf e~",emJ'I<l!"s., lustead, one mliJ~t choose for s.eneswi~h s~rn ilar Il,tt~'Crm of eNpxe,~if.l!l, On point B [<11.1<1 ir is nm necesslIll.'.iJy equal ~o (\<1011 poiH~ B <1. mnl'libel" i'~ c~a~.'acte1.~zes pOIse 972 or~h:is: issue, Frey and Dueck pr~ P~lS(;;,I ~'1e\I!;lill~r~ocl flit firldiing <Ill o~ti.l;lltll set ( ofclus~(!rs. ~J I. Their ~~ddm~ d\,!tec~s special dam p{'li~llt~ c-alled e\em,plm's. and conuects ewt)' J,~I:m palm 1:0 th.e 'Lil«:lIl~pl:lnh .. '1lllJ!!Sl.rupre~t;;nl.'liL ] IIIprime ~pl~,.filld~.~!il'~1J op~.~mal~ of'(;j(cmrpl,:Jlrl> ~s.:I hflJd problem. bm this algo,
S.,r,]:ood d,Lt,! ~mu rsi'l'om ~c:ie:l~lifil,; aad nm.cdliiCtLi.~?ipc.rim(:%Il:S. resc.iirehel'\S.
I'C:POJIl

··

or

m .:~.'t

I'i~hm~s<lble H'I elricieill~ly and (IU~cldy


"'(Or), 11"~I¥'ll pl';oble:n1f;'t~!!lc]l ~~ gn'l~ipil'lg

hmu:ll1.!)
75JKm

DNA s~gn'l(;l.!l!\"~no i cJ!!!l"':erst A~ ..mu~y~ sis, dhU WflU],J lIlo:r.mJmny lfllkli:llilll'ldr~dlf; of


llOUrrs.of compueer •nne might
.l. It."'\".\' i~lil~lU'CS.
I1{)W

:zm:m

be done in

Detecting

cXeII1lipl!lirs. goes beyond simple

c:ln$lef.~Il!!l' as ~hCCNCI1Iilpliil'l> ·[h('m~d\ie.'i S'lure

COn1,pJJ'essod l nfo n'll,llI ~o:n,All exalillplewil.h a brnad "Ulg~ ofpmi ..~bl~'UPJl1k:M:i!mb is found in Ihe s~.fIIis;tlea ~ ~1l~.ly5:~s Ingllagc., ~:m {J·f
in$!"ln,,""(:,,

take

ym!r

last 5CiemificPIJ~)er

(and

data pO.lf1It. 'lhe s.ill'II~rity between {lien)' two b sentences C.ITm be cmllllmted with standard irlf(]filt1l:.nimll.l1eng:y :!l11L'llh~s qth::.,t ~'" lh(;l !>ili!li~ lant)' increases h~n the Sl;lm1;:~1(1;l5;'! !iig:~~I~~1;l t~c S~llliiIe \l,1'Q:n.h). Knmit.ing 1:1ln~ sinrllnrities, ollie C<1ndetect .:~eexemplary senltmces III the' ~ :plipe:r,.\,flichptflv:ide IU'i (}p~ilt'i<'iI.I~' !l)nlfiditlil:S.-Q,i ~ dt!scrj.ptioi1. If you. are Ji hat::ly :re~.dcr. you ean
1j,~..

no .' ~don't real ~ys~g'gest ~hm it ~si1l.l!al"ge.lJloisy : d,n". !'ieH "lUd cO!1!!'iid~f~m;:h se:UII~e!'lC~ lo be <~

~ ~
~

Stiii:II1Iii~~q~e.aom tilbcratoile de ~!'!r>iqurii '!'1i1~ol!.riqul:· 'e-t is MOOel.~ :Sl.dtin~uel, U'rnjlo"ni~ IP,ni~:5wc. 9l<!OS QI~1. 5 ft~i'IO~.,.E·iiI\:iU:ii'lel!'~i'i!Il@lpliill~.ii·i!l~li!l.Jt ~

']b~ 3l.!II~or ts ,HI]l~

(I:n!re

""t~~!1'i1I~ de:

~~ ~du;:~~~4l'

C;;'Ir.. !.!.~!gg'i@'~ "Voc@:d'O:l'le dJi :5oi11Fi Mil11IeQ\." lH~w tQ (Il:o:os!ll'an (l~~,IiI1:~~ rl:h FQugh 1IIl~~~g!l' il'a~'Singl. Th~ Dll~S" 5a;g,e!>a~e·e;:dl1<l!l'Ig;edin Illlt! di~e-ctjons or lite ling:ers and 01 the gl~ n;c.~s. ~ead i 119t~ the re,mgnilion or San M.aueo <liS 111~ ·e)i@m~<!r"~ org 5C~E NC!E VO~ 315

'WW'!I\I,.sci ncemag, e

1 6 FE BtR:UA.RY2007

PERSPECTIVES
aHpoi ~ts. and .d i points .U1."\U~I'to :a!~ One d::Ii.mpnum ld,ls clli:l ~n1g(:1of c\o'cry {lth~r poillthis r~~l1Ikl>di.-.l (If f~\,'oriloe e~':;,-en~! ~~21·S.An 3.1llg.dte:lls every olhe:r poilil~ tile deg.~'ee 0 f compmibi ~~ of his .~i w ilh tile ty st
~lg,e~ 10 lOlng-ek

mCt\, ..ng.tl ~;:;; evaluated through a s.i.:l.Hpol,e (.)mll!plll!ll.t:i,ollJ on chc basis uf therecaivedmessages and tile 'Sindlmity mairix, Alitersevel'al message-passi~lg I1Jtliflti:., all I.ne (:'harael.'ef); reac1il Zlnil.~~,It.':1:;~mem aad evc:ry point k1.lOW5 its '1l1"~nplal'; In px'lctioe. lli!e:L'II.mni1ig~ij me of lhis a!gQL'id~nl ! sea les ~i uly \\im!h the mJl'lnb€il: orsim,i Imri~k~~, aea At> uu ex,unpl@. 'lIri.IfI~l)' P.IiUP'I:'J"ltt~H1C1~nbe it !)O'I!.;c~rulm.elhod 1.0. exnact I'Cpr,esem!ldvc f~~'e:;,!ii:om, ,g<Q~~ery Wil<~ge:s (seethe second ~ on figu re}. The .in;poIIIIA ]liM oJ nnmcrieal ~imii:;;;I.
;:I.nilg!:d's cOiflstl'aim't;;" Ev,~ry S~I][

lal!:~Jie:.'i b~~. \'~~!Il p,~iJ~ () f ~l~\t.~ p,(liLll~. \\'.i,l;i I:;h

.um~ybe measured, c'OmpLltod or, in the present example.


i~lsrti't:'tiorli

USdllg.

amodel. set by vi sua ~


V!lI.Ull5

~mis:s~iJ1g silllaJ1~rity

~iJ1~lli-

!.:'Ii.ed. ~\full~ q~.estiol1l mal'ks.are accepted by ~he ~,Ig()rithln),Each f!lcci:;: a Jatf~pu:illi that ex,chmlges ~ness:,~ges ith. aU other faces ,i:1Dld w
t~n:li r!l,ll>l'ird;i;,'LI'i aifl~gl
s,

A n.t:t!~l~wi'lBnftions

1 1
1

boxf's) l1aljle be-eii'l c1etNte-dlmm ~ '!lI'iDUIPof fa(~5 lily a Ifilliit~, prnpiil[l" tiD n,

bot 00 lcred

f.<lI~~~ 11111a ~J@lUd.

b:~I11,j;ll\lr~(highligh~~d

Un SieU PI~ilWliilafi~~malni:l: foOf a

~t

,of !I!(es,

~
.~

C'H~(lI'l lhe()ry

'8. '" !

ea

a
,~

~ \lI'~l.erei~ ~$n'rnv at I.hc herun of thebest ,crOOI:llo[l'(':cdoirn procedures, and iater in [ne stlll dy of
ildj,\lmm:t': pmblt':IIIN ~4} MC55!1gC p.~$sillg C.lrrl be under-

.n.

of m.e~sage pas.... i~,~ il global ag;I'eemcnt I!; n;eached Olf! the set of e~ema)IOll'S, 5m:h rnessage-passi ragmethcds I'n,webeen sh[),wn I:u be rem .. rbbly ~fl~d~II'Uv:nn~~tDil}i u &aan:d I)ro"~.clmllllill~l i:lle~ltclc error eerrect h'dl. !e<;lfnrjm~i'nlllaw.,~ networks. c'Ompmel: v,isimL, aad tkm1l[lni !'ling ~he S;l:[d51i~~biHI.)' f trl<gi.clJ~ o I~nml,ll~",,~_!n :~n"'lly (;,,~e~ th;e1i ;,u~. li'l~ be~~ ~walibl"~e3~gor.~1.~1InS,tllil.dthis. IliC'\:V app,llicat:ioll to d~!SI'er<Ln;~~}"s.is I!ook~~ very powerful , Uncler=!Otan()l:ungtheir limits is :I. maia Op",1'!. ,(;h!<1lltllngJ,;,
A~ Qh~],(l'\\"eSlleve~ this m~,Ul~ (:oHtlFo!~iLlg~h~ COJl\iiCr~llllC: propertiesor o.hc (ll1.3ii[y oflilc

~.

3,

..,
4, ''l'

~he'<III~riox,~ knQw~edse ef'how good! poi~ntB is


IJSan ~x~n~plar, :!lil[)~tq:L~~.:; puunt!O a~ ln al] eqy<!.!!y ~\Igwh~e. ~) <l!~the !¥I;m~r,.. ~<lk~the ~jmlt:;'l;jllw; P. Til'l~i;:qtL~UUit:y p.i'uvkb'1~con:tru~ nne i~~i.lh;ly~t1 fiml sd.~bllce 1~~ IJ rliil!i"';:':':'~~lg'£'.~S5i ~,Ig(i(ri tl~I'b':I,'i>f'ol.: I1Ig ~ the' f~l'Sl, figllre), and, 1taims <It m<l.'(~ izil'lg the' bin
net

s~ood by [a~d:iflg1~nlamiluopmYl.flfphiic viewpoint lmagiue YOl!lare ,I d,HOI poinr, You \'lim'll 1:0 lind ,m exemplar Ihm is thC:l1nflSI sim,lIa, lOytlul'Scll:. btu yourchoice is c[mSltrnin«i. [I' you choose some other point .A <IS <lbl ex-en1p!;u:. tt!en !\~1ul!'i~~uh:g,~tclJ.: In d 'Ix: ~t;:-; nWUlt;l'<l:flmplill':lh.is c:re£ll;c:$lUQl: ~ oom;H<li~nper dat<ll poillt e~;mblish~ng<llal~bre
net .... ~'Ork of con:Sl:tai,nh lh~,t~lu~M .!I~ .~ sarisI

mpllOOXirnalJe solmions dl>llllh.e',¥ lind. A more ~()01ilis m ,C'ht<'l~'liWleriw pmbl~nliSi lm~ where they can 'be ~1:l>enll:' eoneept s snd The' iIiIu:lhu ds dc.... ped iri. stat is [it:£I,~p,h.ysic;fO elo ~ostudy eollectlve behavior Orr!!'1i lhe 11I1iOSt ~1rmn~~in!j:p'~rllpe~;liv~ 11m lhi~.IIl;:'~pcct ln physi.C5 It:rn.15 .. belief propagadmll (and therelore af]'iI~JIYproP<I,g:U:I()nl is amean fli.eld-lYpoe'ITlCI Ilmc.il, {.n" ih1lll is, dlC' complex iuterm:t~m!J of gi\'ell (lbjel;t ~.. ~'<!t~lpoirn~ c",,~hh all or lh~ others is liIppro;".imfll\;,'<.! by all average effective illiliel<!l;tion, Allhol~gh ttlis
ambitious

,I

~.
~

'iiv{]:(h.Wl,:l~ I

i~1fI1:I10t\'

qn;~s. Lima), getun:to

trnu-

fi!,l~~.\;l,~'h'G'lI1 the net .~~mi~01~i .~~~~"Xq~li7.~ t:y with. a ~~. Olt'>1:m~Jut~ :s.misl"il.:d i~:iC: set of HI::nl~ll c parameter: The lal:gel' P. the mo~e e"".e'.I.np!~Jrs, e.'«:!1.l1pi.<l1'S ,emerges.
Nmvil'limgj'll~ thalli~1o[tlo

each PO~L~ls;t,nid!;
501~l~0I1(: else

AI1:ini'Ly l)roP~G2Llivl1J ~~ k;nm,v~~ {.'lQJl'I],mlca" a!!Jtmrtli~1191~geIJ ldllil~g whelher im t;all'l·~I.uity_h i~in nlct mJ'IPI1,~i.~;at~.ml, fO! o mednod ,t!~~.lc.d"belief rm)pagal.ilon:'v.,'hk:h \\;;.I~rlvenlecl at Jtea<:t lw~ce:'firsl in 'oonmmni~ ~

11M clilf)~~~'ithll'll point as 1il1 E'};,i:ln~phir 0:[ Ito' .. A ~llllp'pro". immte so1.ut iom of II~e oomp~l, kOited '~reb of C~{lnni~~~:ill!,s,~~c~~l~trtl~nllS i\1 o'tu::.tined by h21vi'i~.g9 H of Ihc.o;c d,1111'~CI.~rsu~lik h) t!ild othe:I';At a g~vel'l tilwe. ~ll.Ollo1glC~S sene] me.~~

C'!ue<il~'I'i.C:;j).fLges. bc~~ W't}r~ um i~~~un1C i has sp!:;'t:ill~(:a,;;;~s 7i.btil. aga iil i IS fu! I nll]:g.~ tJf ,0 t ~ <ipp,ik.,bil~t)' Is still to be libmul ~
A InJjlg:\'\,'it;kul~P~d",£!o.El ic~l\li~~le. t~l~ ,: aUDtlj1ro]~mnl'lrl}in.iC' eyq:'l~~.ml!.~.~111m~;lO~1gl! ~ 0f Fcl,--ISSi'Ll~,11 so unde~1ines it$. ma~~fi~m;Llre~ 11Ili,~ ~

ble '!11f~1~1 !i.~~'I;lmg~t~oc!g.~~(l ~.p'lmwmln- ~ the ~~i.~tm ((H.'1~'lcre SJumc o(JIrfC~utitliil:; become ~ e,<,,11I.'enely~ollg·r<u,g.ed. The al)proprial>: t 11I1odl·· ; il'ic.;:[~~olfi. wh~dl ~ql!ll.in::fo;lISla.lI1I!;l n'lo~ ~{lphi~li.:~.

116FEBRUARlf 2007

VOL. 3150 SCIIENCIE '!IiIWw,sciefllcemag.org

PERSPECTIVES
StlOl.tegy can find .u!le",re~lem aplnoxil~lOJltt!' solmioU! m !iOm!~ of 'th~ most d.iWicL1ll C'tIlTIlpn· 'wtim~:d pr~)bl.'lln15 with O!! very !'iimple reeipe: h uses baSiClmlb'SSH;gCS, hich 31\: cxdlillngcd in w a dis~ri~n!led sysrem, togethler~I!;'uth sil1im~e Ulpt:l~)tt:::rrJlh~sthat :l~tl rlli!Ulyb:~d_ This rca lizes in p,uac~i.ce,~ 111C\\' scheme of cOlnpl1~;Hjml. based o:u. di!Hr.i:i:lutoo, simpl .e de.lUl:~lln!> thalt operate .~np<l§1!!I~e'I,. ihe Sllil'il of nelIroOOln· in pm,nr~.(]rl_ Ch'tf: might expect ~[) flUlid tl:"-'l MJm~' of hs pr.~lru.::i,pl.C5 ::11\C at vrork in Iliv~ O:r~1Ibng
iSIH!> or social sys~enls.E"ch suecessfirl :Jpplil:'~Lti(m C!fnlllf;'Sag,l:~us£i.liIg. 5Wtch ~ 3ffi,n, .. it)' p.roP'l1~ti.{m, h~s 'J(ld~ iJO {Hlr W!!1d~~lalllcl,· t

~e,'"

i're!s.. (i!rn~ni!II!Je,. fflfI" 1 %~" ,.t. jt, ~W'rL P.fO.~b;iJmk~~rI.!1 G\ '19,BS~" s, 1, 5, Yedidia,.1r'\f, ( rR~n, T
~f~ ~~i1i ~hj,'ll

at ~sible

i.o: i~fe:1lt!)tfJ! Sf,iolerm:

{r{ffttJ.,,~,(Mo~~m~~l,!f[J!~~1I,

illg of co,mplex s.y:SI.cms,


:1. B.I, F~~,. 11,10ti«l:. .5e;J!tjt~ ;JU.,5i72j20(7); F'!lbli~lw!d ,~~im~ ]<dllJ~f'!! ,00, (1(1.11~dierl~~J, 1:t6aOOO:n il. 1',«udj~ t4 i~t fiftJi fWttll!y 5fiiljJo~r.om Oil' tI<i~i\il!!mlli(<1l r'1.!i~r:lC';lOlld ~ fIr!ill1Q.bflI~r.ll.e' (i!lm" ~ r.ile:}llEIl,. WI. (Uni~, ~I (~llirorraia frM~ &er~e'i'" (A,. 3. Ill. G, GalLar.l~rt ~ml:l~1;I'I!1'<mi!y(Ii'Mi(~uwr

R~e!'e~!Oes

Y. 'i'l'ei~ ,I'U"HlilM1l. MjrJr, 'i\i~5~. :2Z;fl:2200S~" ( ,6, 0, !)!I~i5., It. MQllil6~, a.,Sli'Im;l1'l, lit, !!ieQl;him~ Eidl"

~Iil~i~l 'i~l4J.,~,~~M' nIlMitiiG:M, Cbmr;i~~ito(1IJ1 ~n If'rlltllfol'!1~ ll'Ifll.r. Ct:lJ'l'l_Jl;56. :zn ~:wttll, "1_ "- ,M~ard, Ii ~.!JtIi,~_le[(hm,a\ $ti.PJ1.re ,l9'7. S 112 ~(!e~(t, ~e:n. 107
(2nOl~; putli~1ted ortlilll!' U Juml!' 2(Hl12: «1:(U:iL2\6J

Ji.~6np, 2oBJi.,

GEOLOGY

Grolnites molke u I:l a ~~r!1l!?art (If th~' (Jij!'lti~elltall p

crust. New ciarta divme fO'fllfl<!tio

re'll:eal.tJ1 eh cOIlll,plec" ,an,~ nt hi:sl~lIrV, ca~'liIng fOIr a !'em511m

of the g'~O'l(l!Ji\.al nist(lri~~ of ffiillny !J riilnit~.

20th ~x'niury ~:',~~hl~I'lgfl" s<den~_iji:c,c~vi:l _ \"';~Irover the or.~g~[iI gml:l.llcs-I.hc 01' coarsely crystamm:. re~(]spaN'ich I'ocb th~t n~.ab;:s~i.che_,,"~·~II:L;1ii1 Idilllg ~lmn,~!'i ml'lj bui kltehen COllmer.~, 11\e u~timm(! losers t.l~ held !lmt g"mitesrrecipj,~~ted from .,qlLetl~~~ 11u:i:d:s; l'lrlilt ~Ix:o]are Ilhro~gh the crust, or [oruned b.)1

~O~QgY ;!>-p~ml:h~ 191h ~mdn1iud. {JIftitL:l

f Cnetlr·C'rLIShll ig;ilk:om rocks (henc~ the ;;1" For igneous) (4. 6). A ;~n~norityd~~5~mh~gvi~w $U~(;st:s l~~"i.I:h,~y are instead 101 rgely mantle-derived 'lI'Id onIy ~'i{)dil~ib-d by ~i'i~.g~ t!fiir{)~I~] Ll~(l 4;;U1U~~( 7), lhe stak(;~il:l;l this al'gulfneJIU are hit,h:
gnl:~~ i~~l>~Uffi!.ne:~t:;0

d~Il"tlremunt,e f[ol1mthe ~e~ ~)f'Il~ si!i~!lle ~!lI!ih. l alldDht:, ccaseqnences o:flhat dulIere,ll~i.aliolil far il.lt:: cOlnlpm;it~ul~, tlfthc mantle .. I.n.typt::

I-~ype !;F'.l:lli~es(or ~heir me ••n!ilm1~hosed r~r e'~odecJ dJerhrni i~les.)make tqp <L ]1<L~errac~ ion of

c T!it~reJ{lre. tfll['~llmu~l:n~ reaClit)f'lofpTt~",i~t.i,n~ fO~'k~ wi~11 'uc;lll]ui(~!ol~ 'tl1(.' tlJ:lt~:~~~~<lJIt;;m:SL ~ reg.ardLn,l;llhc.ilr {l:rigi~lS ilrekcy Iu lIIlth;rsmlldl, the Wimli.!.n; recDgn:l!z,,"'ti thru: gmni~c:> erysimg the 111WChallisn1H by wbich the enntinents I.,U ized ItoClm si l:i:1:.ne mell!S.

en

\~'l, dl~iit:SoIQlliun

on his

11[gnmlBllt

led

liD

~(Jip!l;:i&, ~!I]d lJf)tr,_ ~oK imt'eOlclL ll:!£)' fon» by mclll.in.g l)roe:o:.isl.ing ~rtlMILII'fH:k...,. the')' represent OJ meehauisrn b_y~vh~chthe C'{lHt:inems IliUll':mlll!Uy rcdislrib~llC du~i r\\:!~ri (m~ 5~diunLJ:Iil.-

Sll<ln~les<U'edesceltdecl from basales.theu lhdll' l~mt~.l'Ilim'lrer~~:enlt:ii ni:~ g~}\v:th of~h~ I,)ontinents 001.clltoC'l. re·liHa.val rl'Di'im the' :1~\~UMI.c' c ~e-of ments tha~.lre highly OC1!!lCellltf'.Jted ~!~ erusr '~he (such as the he:M-pil.·od!l'ig~ radioactive iw~

~t:

various mh:en;dll!!!!reINi,[~l ah'l1lo~~as divisi\I~. Me the sHiclli!(!mlldtsthat g;ilvl: rise '[0 gnlllli~.c~

t,~ryaad ignl;;"OW!5t'OI!!'sl~llle!1~l'_
One reasen L~l~origill of grulliJ~ is such a d iffl~,mh probleunl is th><lt Ihlese loeb can be
llxln;ml,l:ly
C{lonpli,t::~Ulld

partialmelts Qfpree.,,;j~lil,g I[{)~;billihe com·i· 1l£!'Ill'l1 crust, Of ~IU ~~.l~y ~:U!iSl:tllfl.d m ~d'II~i> (~f [~ll.:: ~:r}!~w Ili.ling manrle-derived b<1~'ll~~ n~1!~.o~ g.UUS Do Tit!.! brine l~I!U is!cfl '\,ii,fcllCnice freezes out of s<llt)' ,.v'tlter':!1 r gr<milies form by erustal
1t1lf1ltiing. do lilley mlnnl", lirunlt the ~ediml~nt--[iclil l1PI}C',r~r~st OJ: fro.unl preexisting igneous rocks , 11111UdOlilli.11I1UC 1.111>(, ~01iVCU crust? Om! P,}@,C 9~ifJ

(!ii~'" 111'" fugLmt::) ~8).

Ma~y "Ire

nfunin~!O'!Js rep:~nt 1Ill<!! only ~ comp1Jliler~t ft~c n:::~ls fm~n \\1~lkh chc:y o f~rJillOO:some are ,riluxnl~es of miaerals ~.lal
C~ILnpO_\{'_'l1

!;,!;:ruwCtmll

>l!uffl'erem

midiS:

:i!Htlle C{)n~'IUrrl

of this issue, Kemp et ai, l.)') exanune these


q ueslum~s lhmugh lln~ Ji~Ilt\ofl.wo uf '~l~
Il.~.\~t flllfl~yLi.JC'<l1 wols

mnnelmedremn~ms onl~ck sources; imil im:lividtlfll mh'll·:rn.!~ (JIH~n h:lil.whcn,~ erogellrolls d.eunlkal .md ~S"'"
lr}p~c.:;millpOt'l.!llflll)), W'11~rung lh.c cvoludon orl~lcir parem<'!lJ

developed for I_he

theil," crys~.)U.iz;:1~ion . KemJl "/ (fl, (;1 ex_,lIlJj:!II~ lihe'susp ec:l,.looges of Hijpe' ,gl~a!llitl!~, .rew[,!1b1iillglll~(l5e L1~ ~.1i m i net! by Kemp et ai, Vhe-aena!. pi101Dgr.3:p:h (mal III the Qri g,ill,ll~l1ld C''I'O~ a oil (I r ~ deb,\'l~e th.u lq«)per~cnL~:i)1 sedimen~ SOI!i"e h'na:g:e-) shows, llile Sail )000 plulnn {~lid1,cu~), alii H1~e 1'~)iI~egmllitl:'.'l fi.'OmIhe LachThBi~Ih~s.t~~I:i1t¥~OI'S.lYlPe g~"~lit~,.. (4)(\\'Jl~~ .'S" to n~the, or ~lIb4yp~ 01 gr illilile" $Ii,:h pluklll~ il;~IiIlIIiI'I()n1.y lomll 1ml be:JI~ ~n Am;imli,L ki~Q.il1iele;r-sc.<!ll! I)odl*s ~t!'UI~,d inta; ,rocks, 01 ~jhe l!P~f (l\LIst KOO1ij) el (ll, 1;\1\lf~ ,~,.,'!wson seveml ~C~Hl j stmldi> f4;:!f s~~,hm~ln, <ll!!d lhmmm~!~e·,,~~n\'w Q slllgg~ tlitat.1MimiliMion 01en,~~loQpi r®(~~ ~nrlluen(Mth~ (OIlrl~OOiDil»l~01 In~.'i:'wflnit'i.lytkal hUlCW1ilio:ns,. n~ ~ ~sfuits g;i.vc d:;c tu M~.~yp~ g.r.mi~~s (3) (-'M" '>S\ll~h booies~ Thlli i9Sl'~ Sll!1)W.a SPOCillTl ~n !!If <I '5imU~tFtll!n.!lil~ rruln nJ.~ im;~llcling higta~pUlecis,ion. in ~ fOl:I.n1UHle}. Uo\,\1l:ver. members of a Ihi~d :ild 11&"-tille~-lype(4 )-;:ull': <'lh~lm~lkL!nt,wi,d:ety (hihualh!JIi3Valley, Ciltl~roM'Ia.'Thevisi tile ~iI}l1tptlot~mlP:h (ri~lht 1i11sm ~,it~ rirlIe,l"ltwen~!l;:ir'ilts i;l.fOK}'~.!ii rereals dark Ill!lhG 01 ilf;11plnibole and hexag.olQal >CJy5t~15 of bOOiliba il!mOOdded ~d i!>lriblU~d lIn:d diverse" lind I~c.~r Ol'j.g~nl~ 3J'~ :i.soto,pe mli'().~willnll h~rgcin <I'W/nile malriJ: of i,n[E!rloc~in.g Je,lrI~Il'at" .i3l'ull ~uartz, ThE! llrall5miljl~(Hi'!lnr~. ~ tiP fo:1tgr;:Lb& A ~)tJJ'j~ularVi.l=\~' lloldi> thi::!lT~il:l~cc pholOmjc!Ugrap~1 ~~f!iUlsit!fI ShOM !winnimg, ,c.om~tlqoll!~1 f@n~ng. olOler· [itt! ill!> i!:II'imt'i ~crop.r'Olx: 1iiiI{.~~iii !i situ measurements of~m:llhhJ!I.n gmr,'jilll's, and indusioos in lP!la!l~oclMt (,(omple)l ~cglliirl aiild da rt pallem), :! hIN~T ahhlliml j ifl1e ilutnoJiS i!!1 Ihl!! [}i~~i(! m or GE'D1og!u1. lind, fl'IMenilr:ro ~dj~wc·-fI~.til~ cry.stiill"I ~phlib(jle (bWWll}. l'iu;lmit;l'lhll1<1.lyU'-iI1 tedhlilkilll~s, irompr.'~ lI!~~il1g f; eni~1oyffi b~ ij,'ienip EI {J1,i3~m T().3N{)icB ar1:ifads llliilG. <ln~ Irom miix.ilil91 d ilflBr- j (incd \'II'it~m~ iindtl.cti~(!~)' )cleirlC~~. CaUf~rrYlIli!I~IIMt~ O'I1iec>linology, i'aS<!dena" oCA ~ \l'n2:S. liS;!!. E;-:iii,aU:ene'1@gp~.e:iU~th,cedli ~r!l; COfnllJ'O!'Hlllllts 'theS'!!,cDm[pOSi!ioo~l~ aM M:tl!r~~t¥ (omp[e):. r~~~. oi coupled !llasmOl! mas,'l Rrec'"
.:Ii
C~e'dl:.lIltcl>'V!,i'Crsw

""

,e<lrtil

sciences, ~hea,bove qllt'stUiOJ1S

m,lgL~ms ever the course of

... h..v~ bee:Ll fb~~!l(1 ~m."\rimls:~y foor MJme ~eX.iren.~ii;':iype" ()f gnmhlC, l'hc=re .i s .~ iulle

WW'!IV,.sci encemag,

org

5C~E NC!E

VO~ 315

1 6 FE BtR:UARY 2007

PERSPECTIVES
UUliIleler,

Sedil~U~!lhi ,!IU~\\leather'ed or hydr-odtetU'nlllly a.hen::d rocks in tlllt':S'h~l]ow CHI:S1 h'M;;OXY M~1l!i_"O'tope comlu!sit~.<jn~ that d:u]fer suuliIgly from I]~OSC of Ill(! :1.l1ru~~I:c of deepand: eruaal ig:lI!em~s rocks, Uafnimn li~m;tfpes. ~,\'{]o~"<Q: [llr{]!Ilglt limA] d~~ Uti pt'udllLcl icnli. of 17~FI f from .L'3d ~ooc.:lh'e'dee ••y of ~?~LI[q gellera~~y and differ l>'ImllWY between ~hc mmuit.: fllli.gll In

De re."l'~l\~eponed by Kern]) 1.1 ul. rJ'e~em ;! r pow(~:1i~l. ea£(_~lill!m mll~1 i1.\thiulk I:hi: mUlo'! pt',P~ wil ul'!f ~pl"»I'It"i:o!l fJft~ ori.gH ~l."of i'lj'l:le g'f;',mifel; 14,.61 and rl~~nilzc tiM'll:loIwcgwlnl!.csaOC' clese relative." of IlO!J~h M· and aiLe' S'WJ)e granites the
~7} mdtl&f ~lml di5[;l~l cullS.'i s (4. 6).lllil'::S!.:l mll~ a

elusiens ·w.iII~ force .revisi:ml:!>on:~legeological

histeries of'ihc

lnllny

plru:c.swh.cre sudl.

!:l~lflitr.5

ium ~sooo[Jes}. I(ljlillP N t1f'5Itilltk:1 M~g:g.cm E:1~mponioli~ of IJ1tl cnrHin>e,!~wl CHiI[';l ~h,!lare rit;h ill~' type grzlIl.iles should evolve in composition Ihrotl£!h ume: The degree ol'mel~ing of tile sedimclli[.rich upper c.:~rsl shol!l~dullcr'~r~!>~. amd the crust ,IS. a whole should becorneprugn:iSSU\!c:lyicher injuvcnils, mamle-derived r
blIl1l~efinl,Tbispredict
iUDl. CO~! Ic:lbe

I( forexmMple·_ OS!'!l

Lullifr<.ltio alild hlS rehnively ~76nf~ficll)and <lilt iell!. cfllil::oil kl"" ill L u/H f :I1H~.n l <lnd thus relsr i'lleil.y 17611-1 r-poer], TogetiJ.e'r. these geoc~e:miI:,OlI y5i~em:-icai!l ~neOlMUre the '1.)OOpolll.oL1S. ~

are fmmd Keniljp >!!I of ... l!~SCl < Sll~~t ~ha~ OOn1fil~JDl ].IYIf~e~l'ln'lli![;."~rerre.'i-ellil crustal gruwth <1nd 0",1.

test·ed \l\lj~h

of recent m~mle
alld the
.,mUUlIRs,

eX~1.~.'t<;"'e·rslIS

<Indent crust

nf ~h,~1.o\'\[. ~d~iill~I1l'Iich ~

cmSl versus deep. ~mgcIy igt~'lus. erusr :i.1] ~h~ SO~q"QeS Of.wd~.lil-e...,. K;.::n.lp (/1 al_ Il'i~l.h: their uxyg1.::ri1 ~Li1Id
~m[n~~lIn i:sK)l{lpe mel,l!Ml!rem~l~ts nil ;l:i~·(,~m~ .. 0' U'<.'J.CC mineral COlnUII1UU! to most gnm~l.c:;.

.J~m should faclor ruere e x tensive!,y into esrin~ates or lile Jme· O'Inclmechanisms of Earths chellfnucElI dL:ffhentu,u~on"More gel1.er1LUy.~.e t autlsers ex[po,'ie dtoe (.,,{lmpj.~ .• Illiuh i~:{mlilp(JJ]ellir ouigil1i of gRlll hes 1lI1;Ki<trgt1.e ro:rec:ful~y.Ilg21.hlS.t me ~:igiicl.e~her,lnr <'PPRlOIJ;h lli1!l·m e'dnl~&::: L<:lHi:3l<;
IWi'iftitaken
I.t) Kt)'lHj).(!i

hlrge.~'${:,1Lle'l1J(li~~lISiil'llgthe Sl.'Illilt'methods !l'; : Kemp etar, Such work \\iO~ll(~he anSiltl:"aC"~~¥e Ulex.t ~e~li~lone ol'ge(l!n,!;l},jol~l ,rebntes.. s
~i]lrl)f.]tU!rl:!l

!!-

1I_'(i.·IIl~n~[,[~ ~rl(~f.ifi~\IIril'll:"'1.fi1d~{ripli~1.1'J~· #'Ie VmiQ.!!\! .r~, A. 111,.O;pooa~ Ii~.~H",fl)!1.f; ,ttl!l\l ¥tm:,

grulilC d"'lSsif'~l'3ll\m,. de .. <l¥~lUll.ent th.u t he L;H,;h.i~wl.

ZirCQDl has long been used too dererm ~:nethe ~itY~lll.niz"ll~:~ml ~'g(;':s{)H? rl!lrlit~a" ,1It-cl.is ~.riIl,:M~iO~ g i:nglY1lsedi to oollc:>imilll Ihe enmpositicns of

[hC"~.rSOlun::l:'!i {Y)I. K,;:unp (1[ al. show 11mt ill'Ll o.xyg.en and ·~'mliilimnis:Ololle cO:lnpoS~IE<mS of ziro!J:rIs in LacM~~'1I.il:ypL'l !i\ro.!]'ii~!.1s correlaje wi lh one 'Ulot~er ave r a wide ra nge, Ih:m1 cQm·poS~!i<n18Ch:<lli<!cteriM ic of the mamle ~(I those char ..eteristie of uppe r- efl~still~ seditml1ldr. ~\U~~i'it ·th'l'!." j.·l)'pe !ilmrililtes la:rm, b-y in ixitllg: of din·c.i'Cun~i3K1.i nmmltk' melts Wilhnlehs oflhe ~ppeT ernst, Tht!!S.,.lhey OOllruS-el1il.both [I.lcl:t crustal ~ruwEJ.l {~u[ldcorre~j1.1dil]g !illoutLte dep~elim'l) and wdi5tr~l:nllion o:f e.x.iStl rng p;:llr~s of I:hc 00311 ilti,::~ualcrua.
I!flJ;l'lllJo;. The;.~

:~ ·[y~pcgran ile S~I,~,U!C is derived ~rol1~ pflr~ ial Ili1Il:l~5 Ine~mnlle {_liepemls cl:itic.ll ~y ODl their of i:uu'!3ltplili:l-a.t~.{jnof dl(;,:lm~m:d!.:lllIill;lil~ (lO:lml~)~.ilions, of i·he most pdmiiive members of Ihat :>~'~i~.,Th~5~ rm::ks~uru ·brrnl.d~yoof.altk :;;lbg~ ges1hag dmt they oweImrlia~ mehs of~ddothe ~lh~ dornilriunt ti'l;unl~f.]c;:~ lyre). 1~I[lIIIWv~r. .Jllese cQlnpasitiOl'kg ~ni~l1italso be formed b)' hi~h· fee melt in~ ofpr«x VS1ns igneous cile-;g i rods in Ihe 100\i'eiI.' comtinel,u<L1 ernst (tlJJ. TMs a~ter.nmtH~ieC~IfII"II"~U Iblom c;:olllsi!'irA;lnl\~'itlln S is cQmlll1brmil'n:rpr,eglrtiul~s ef'the origil'llS of 1Iyp-e ~mni~es (4. 6), Th!i s question could be rel'i(l~.'IIedmom cun.chu:;;i.vel), wil.1lI gem:.n~IIII~.cl!l.1
11IIe,!... u~ell1~mt:'> lha~!l!reRen:~iri\'e ~J] t]!e liullie at

1971i, F\~ ~~'~ ~- .I_IM~~, SYl'i'fmof lire ~a.fll'l, 11'8'5...rhe'D~r~l~l!'·fl1.J:!h. 11~ ~~NQ~~m ~il'~, 1l:~. ~9~,!~rl.l! Pfqy~M'~ ,£IiPgl'Wllly r;J H.atI'!'{l.II ~liblni~ O:..nelll,. 'cr, 191o.~, :3- II, II. S:. ~~~!1L,.~~.~~, ~OO{~:OOi"4~ 11.. J,. F,!.Ill1]i!le.:B. W. C:J!a,~. (i~fII,$of, Am, Mem.. 15". ~l fl:.9~).

s, ", II

PeJlit, "', ~bler,

6,.

'~i1l-Mim'I~ ~i(l'l 7~.69 U:9.W), A.], t. "\!lit!!,.~,W,01\1!~e(!,]'e{~.'iCJ

I. R,. li!WrretlC~,IR:. 111', ~\l,

A- C!!~J(), I. 1I\Q:p~~.a!,!!i!1li!~" 1. Il. [)elli!p(!1~ .r~1I Sci:. R~. ~~.H7u'99U, 9. J, W. l;'~Ue'j',.1..It. Cl'li~retl~!i, L I'_H'.C~~!~n!!, ~Mlj
PiQ!tog :SQ,~m~;!1~, m7 ~'l,9ll;t),

a.

,. C,~. (Jl~, E!!.$'iP/.~.

,~~, '1 (7)!), S!;j,.~~r; 7~, ~7 (~~i!!<1).

10. :B.W. (OOllEli,. II, J,.R.i!'!i!'!il!i",I, S.WiHl~!'M,. O,!!t~rfl. 11\llt;l', It SIN., Ed!"I!I!I,.WJIl; fo:lf,r!ij .k~~:5, lJ.:ilS(~4J, u, 'I ~hank l" D', Murray 2j1 d tL I ,!;iI~etr'llf u~ ~I' \hi! ~·ootQ.
l'Iega!iiI'e-usec 1!l,~en~,iI~IIl'NMim ima.!JI! iIIHlll Sill!lef ttllt .(I5~O'fll1~h!and ~~l!Cim~l'I~ amdl~iJ1 ~e;!,1f~m~W5lid1 til (Jr,eal~ tile ill!!!'!:;, I ailatI;Ja[;lt IL[ SilYer ~llr 11Ii~ ilidan[~ 9 ~i'idMli'ictl~ m111iii!i1·~lil~u~lf.

\~11Iich crust

.iiS

11m: I..:'xlrac~ed ~roml the m~,mlc

B~OMIED~CINIE ih~r<l!l':r directed

is

lIf1i'lIIHl t.o

fiOin IS ~cHlld

ala nDll'limmUM OE!llUil,ir~ SillPI}J1l!~~rhelUrrl'ilillOnd

~rthrilii~.

en 11111;11~i{ln); Orll,]~iellin.. linn Mll'lbTlTon~ ~h~.nUl~8~Oli.;d. ~"lnhri~.is,s.iltnpk I~j)k_.s of d~iI)' life 8m:ll ~:-;tYI'llg shoes mlu d Imh· ililg Slep$ .nrc paiHflql ami perhalls even, ilnpo,'i~i~e, Sy!npwp[l~ r'o.l:n~ef"wolH 5<l;r~,. stiffjo~n~(;,

W chcm pflin o:r ~ho'.rtlll::Ss of brt~th r~oflm in!ll!mma~ioll <lfolmd~lu: hi;11lnm:td :I~~.gs:, 1'Ile myria(~ of pmhl'L'lI1l:~~~li·11 frUill .1U[I.inllnu~~(l ~}\,!>~gm ~ms hll:icame 'Clonr~~~ ,Ind ,n~~:;< th1\~
1l.0rliil'Ui.~ li:;Silit:S.

Dcs:pit~ IleW

[rc1l:um~fijt:;

for

rheum!iil;toid ., I'lhr,LI~s that foclls


n'\f~liu;:l:i\'e i !)lliIfililllilily.

Oll

con~ro~11 inlS
~J'l <'i.ct. f

j ()in I

desl:liJ,lct;oli"ulid

d i!l3.bi .~ityI,uog res:;. iIn 1'II1i!lI1Ypl'il~e:m;;;.

lh~ ~sp(lllt\e I1Il"CS thf,j'~e "PI,'lm,uche.<;.<I1"e s~:I:" ~n pf~s,ingJ~ sill1:lalo~.c~n dlu:mgh Ill~Y Uil~t diisrin.ct rout-eS!iO Ihe disem;e. such as nOl'l1ilOneHke InoleC~lles. 01"d~Sljrnc~ hnlllillBe (lel~.type5. e!fllpJJm~j;;ting ~hec{lmple~ilY nflhe ·co~diti(ln, Orn,p.!l.g~ Hm6 ~)f tlrll.s iss,lI!;;. L~e~.rld CQI~ leaglll es' I }p\L'QIN:i\ernl"he~'ill>!ili~1ethernpelllic ~'prrom:~[h~u targ,1;tls. 11 ';:l;:l,Irn.tU g~l.l~mlly 'C\ln~id~~cilI1;An .,,1' ~'hehc~dy's~rnnmnr:; ,~r~' :~m1:l!Ju~. :i,5 Ij 1'1 k~WI~o Linej:oint--i:h.~llh~obl;~"~l~ t like s},!llO'viocyte, Su:!>~ptilOi~iEy to rhL:lmn11!:tlid <llrlhriti~ i~ cl,llm:d by bot~ gCI1~I:i.c IlII~d.C!lwi rromncntll II !Ctlmpoiilel~r!i:, Pati.~±:I~1 ~w,l~h g~~ndic I!IHy pro·
~r.,]mm.~ed imumme hYl~ef"~ct.i.\fity oltlell pl'Odll!ce ,·hetlmJ.'l~oird ft!CLOI!~l\~u!munibod ie~ th9~ bind ~D 110rmHl.:ilnti~Qdics-SOl"nlCQimes :yean,. b<!"Im'e the O"l~!1et of(;~:ink'<!I1 dise'll'e. VOL. 31S SCIIENCIE

Pmduct vml of' 'UII~'-cirr~lHiIlOllH;.'i~~ r(;lpt!id~ .m.ibml.i~sis a Ilolhcr ,earl:y cl is(;3se &no D]1Rrik,. U Liule is known Olbom.llm\i lhe prec-Hilical Maw evolves w ~he nlll~blown dilsc<lise .... il.lhough: !ilm;hiJ\~!ic e\'e~~~· s~H;:h as .!'"erewt~ ..Hct i~"aliofl u:f blllmc i.Ulml'ullIily nmjiglu Cl,')J11;rib~ltlC: (.1),. The symO'L"nmn, 01.' hill ing the join!. p L"CF
'irid~s!1nH-n~ilts to c.art i!~lg~1mdp'n;]d~IG~ ~

"I"

~!.1h!'iil;;.mt~ f<!c ~IiWJl;!n~\lVe:~1!l~l~t:;.t..>.e th~ fig1:0 { ure}. Ii nClmal~ iltdtldc<~ ~, mlflli'\f~.1Jy2.cdl~= Itflr sul:l~inil1g layenlmt is sp,lrsely popllla~eCl
by COlli n.e<;'t:ive l:i~i~~I.een~. 11ndiltunnlli~ ~~~Hs c lm,otlQm1.t::h!all'·IYl'i1lpiho~ytcs·~ :ilt:i.d111 Ihin dil~t:olili~lnUi:iS~llJii'l'iallii'iil~,g t',{Jii.'iPU·~

Clf0~~I!l

T~ <lLilhcr i~ ~e Diri~ioPlof !UI~L!lI!!~IO~OQ~, ijn A~ll;:!9'Y~!J~ t!mm1!~oI!Igy, Ul1jl/!!i~,ill' f ICi!L!fo~niO!I ·5iln [l~e'90,. :5cl1$ol o Mec~~jn~.·~50QGiIllT!<ln lll""~,Me: 0656" L1! J~ilii\ (II O ·92093"j},6$li., ~ E'm~~: @rl~i!ru@Ii:t!4i1!dw U

ar

tb,U rese~':n nlklCI"O,p.IU!l,gres 0 f t~e iblIlm~ne ble Kyst~n~.<l!ld riliurJ!b~ ;Ist-~ ike )\'}iIilOViocyl~s, ~n ['ltcml~~Hoid a:rlblihis .. the u.miJmnl UI:l:i.llg i~ hype~'p bl>;;lic. e'xh ibiling ,I mamed im:re<~se~rI

9.52

116FEBRUARlf 2007

'!IiIWw,sciefllcemag.org

PERSPECTIVES
ho:th een types. lhel!"hemn.noid
!1ubl.iining is {]<W~(i11 C3,,~Jy in r. ~tr2l1~dw.ilth n~.{jln.nrMlt: h I(:j!ll" IYllmpll"ltK;yt!1.li' ilildudil1!!;_!CD4~ T ~~r.;'iH", CD2(l~ e, eells, i'I1ilCII'OI)h :ruges~8:liId 3.ntilmdy..-seclX!tiun:g
'p,iasma cells. M.t(.::u:oph,~g:e~hke nlC imi mal lillJing expn::siS .vn.uny of

~lm09Jn:t~.DmiIMtJi

the <l.u[limrs I'equin:!'i pit:ec~!iie]ocOliizmion of illl dll': jobn~~s \,\'C~I ~i~llrni ~tli,[m

cells in theproillilamrmtwry eytokines ~hlltcmul'ilnl:te to the disease, Syu.lovim;::}'tcs produce prodigious amounts or pro lea ses, Ilf()st<l,g~,and~ s, and II
certain C}'~()k.ilt'l.:';fo; ~ mesr n.nlaMyinterkukin.6) [~UU also perpet ~atc inll[1mn11fQtlioJl. thdike ~Y!!lovio(:}'tes ilfA ~1 ~eahhy j,oi~~, t bose in rhemn<lw:i,d a1'lIil1;',lrlis fife irwashn;: an d a~.~i\r(;;:. n:::m~ Iflisc~1lI (]f 'I1;~II]I~lbnl'ot;,->d ~m'di.,~· ~mnt J eel lSI J~,fur illl'>tflm:c. ~h(;y.e:;'{pressn.lflJ.1Y Pll)to·(lnC:O'~~ne{leha.rm.aer:Lst·LC ()f u:nno:r:i.., can ,

~OO\l'illll!l

Syncw:il!1 n""ldi

of s;Y!lovi.d ~IlmL1~.inumunily l .• proi:!l~ll.~n~.!~t~ m.llI.ory mast ceUs,l. disruptillg normal ~,nov:~~d Oll1,;llileC"~m-e .by inloerfe~i ng, \\' ith cadI]~[im-~.I rliiIi.ght [J;!ock e:vmlltN that ~f'liti'mE litH:: c1.ilsc<'l5e,(H~. The errec~, of (''1.l.dher.ln-~ 1~1IIsed
t~li:n;p'y in establ

ushtd disease, howevct, could

adlileriulg [0 Of :>1..dtIMr.tK':.. ;lUd ,\,IfQ\""lhinluihiti:!I!,l ,11"1;;11 C;O!lt[u;l \\'i~h otl:u:r 'ccUSiflomlii:l~y impill:l;S. \-Vhell synevlo.pml:iJE"J:MIC\",'iduJ~JI
1(;0'111e~.,poe-

cytes

ilfAlpJarl1.~tI.withhuHilan cartilage imu tnit~ lBilm~ck an ~,~apriv~.iI1!M.tlllJI!l~ ~~ysl€!;ll

Ik1\''I;! been

iHt~lINAHlIIII ~tul!l~ms S'fflOlillllN'!

DJ stu nguis;~ing betweenrhese po~ibiHties \\·iill r~u~re mMillml!HI \'\!m1k:in ~lniltl~d nl{)d~~s l~l w: miter p,dogc:mic femure5 of n~elllfl~1toid <~~Ihrili~lae~ides <'IlLW<ll!l~ibodr IO~"OlIl;m!j:Oj.l lU jr~intJ;.h wi~I also be JJtliP0rl"U'll"In d:Mti.~rnillll: lh!l:) f;!te;:: of i1iYl1.mrim.'}'~eS~IilI,;~dller~I1..11":~_'it)d nCillliUCim pD'QI(~t'~S, .00 they tl.n1Idi:l;gf11Lpoptosl;R, llii~,gl'alie c.t~~. or,h.e joint. or s.~mpljl Qe<lc!~vme"!' Anr.nil-I!:li' cornsid.emliomi!> tl:ml ~~1''fl[J\I.t~JoC~''

~n\loh'e tuber C'omp~e:i:, mec:hanis~m such as MU~r)iPB:.s.ing int~rleL!!1ki.IiI.·() prodllll!l':t~.mJ ()I' .. s m;e![I~,loproteimlsee.",p:~,['On 'b¥ s,YIlOViocyrc"" or by d~rec! 'effects 011 ~ynr)',rrm,yte iHv;"sive~e..,~, ..

neot

J ua murine <ll1hrii.is ruadels do not dii ~l<1lYlill'Uil)l of the ILllliio~u~ flgg,rc:;i>j,\\: fC-l'ItlllruS, Ofl'!iJCilllliiliLtnid synQ\,jocyl.es. 11Ile,' are geliler.alily .mH imprDlil'~~dand ,,~pid!y d~m::liv,ac.e wheacullll]reJ in ViiOO. 1iillig pt;lten.Js ",~IIIor the potenf l~Ol~ el1'icacy o:f eadherin-l ~ 1~11;'6etedher;;lpy ~ beenespcsed top;r(l'inf~aUlul~'llo~ r~letlfl~.1to~d ~n hUl1mmlS because ofthe IlOs:sihle added bene$)'novi.,1 cylokil1es or cei~s of Ihe m:J<lpli\'c' Rhe-umaiioilill artlilritls. Synowi.DiJf~es, '.whidl e1(flre5s fil. !l.. iu~dJ h~' h~(lckilllg ttl!!:; 11l0~ .H.~~ssi'il\~ im~nl!n!l~y51~n ~'hf m,u\y lnol!!th~~'IIilli ~ 11;!h~y ,c<ldh!l'~~· (Qmp!ii~e.m inn'l'f membrane thillliil~~ n, :lmrviw J.ili libe meuse, rhcum::uo.i:di sy.noviocYI(; :ph(~mrypc ~h~l :iS Itot ~he 'S)'ilI0\I'i,rul,Cdililr or join1S~t:(l;p).Whelfiaeliv<lJled b)' a rea~meof !'i!tOiF!.,1erm tminmll ~nod!el!i. Iheptldwgellie1\i!' of lhi!: llnmnml aggres· l(.yil(l>Hn,~s., I.ynlph@cy~~s., ~,nd ~~'1ironm~I1Il{i1 ~tnmes (htL:lcllilil.ilcD:irgt: in Ull]m"lIiImoid..~unhrilli.lO ntm~ ls .s~. vebe31~uvior by :l;y.iIlmf~OC!YW:c~ is mn \:'I'dl.· ~u{h ii~ ~ffiLtive ()~ygelil S!peae-~ and Ilitrog en], Ht!1r COH1plf;W~ n;lmil;~i{H'll'> Jb,Jm!lthe <i!:\'uih~ble wrdefi~el;lbl!ll[~lCm:lld !>i~l~lly repre'~~l!,,~~m!ml~ pelrlPe'~&l!m\' dlSt<lSe b~ prCi'dud['!.~' PfC1i!lf,jAlmma torr response [0 the rich rhl:!ll~l:l.tO.Dd ~y~.o.kinc rnedlsters and h~ ~llwa.din!ga~tili3·ge '(mid"b, loot- geted dlc·mp'~cs ON:~'L irl o.rlJ,y :I m:illlor~lY of ( O elwi!'ollU:llenl. Olil~he Qi'~,ler hand. pe:~tnanenl tQIIl).illIC'rapy diJ~c:t'Qd at {i1dlh~illl·ll c.cuild ~1l~!I?mlIJ)t panems, U!i-i I1I,gJ series ol'ilnd~ v idml~ bio]ogjc<I ~ ..g~f~lS ~h,.~. t<lli'gel diffcrmH c",ll~im.e",g'!.'ls impri.1'Ili~lg in response ~o lline:: gmt{llO:x~cea~'Vi· t11[S c:a.scaiile by blocl-iiJljJ S!fi1.o ...ioc~le Ilm;(tion. fO!"lm~'~H of d~e melU!'n~~toi :;;)II!l(J'ii'~.I)I!.n~.!n d istu IB (;e~ '(,)d~'~) p:ro~nlll.un~.l'lM)' -y~okine." I~.T (If c OMl.Com£..':l aeceem f\J;1r .I~i~ 1 bch:'l\'ior-~hat ts, C'I<'Ci:llS, such an llnJi!1Ilf~1 mudd .of aru.lluilis, ~iIlc]ncd.in,g g'i,;m;Ii.~ (nnnorll()crosL~ ffL'Ctor) has i.ffiI.1P.rOVct! as sOJ.natk:UllliltJIDOnS aud (J.nooge~neexIJlJ'e.\.<;;iall Ji;i<l~~Y and decreased llilelimmber p."IJ~eI:lL~with modifiedmiee t~mllack the gene encodU[lOUlllmlled d~:Si~11l\1i;. omlJ,il1l111ioll, Ih~miPy,. C {.':(l'U!(~ pennauen lIy aUer tell br::imvior. S~u:;11 iL~!il C"idh,~r1in.'I L cadherin-l !-:.;peci fie ..utib w bodies. anda cadherin-t ],I"Wt5iOIlI pm~.cin, l:n however, :I~as, een fr-<i,ugh~ ith proh~t!msdlliew ,gcnelh: cht'Lm~e5 inl sy.noviocylC'S [Ire rcfleered by dysR:gtl.lmi(·tll11.m'llil·~.clllil)n o:f proteins (S'lac~l ~3d~, sse, 1:11cy e sltuw dl~l ::;,ym}Vr'fl~ ini~:l'I11IiImfL'- Sllp~)ft:~sed host dd:!.::nSt: wi:th[)~ll add i[ifmil~ rien Is suppressed by blockilllg c3dl:te'rin~~~, cllnical beaeflr. One ill.liri&-"Llnng possihilii:ty as stnnoyllmi.on,. defective apoptlJsis. and nm" t t<JtIOI]\l of the r53 ~~If]um~upp:r~!:'i(l:r!)j1i:1iI1,l t5l run.clion. Curti il;l,\l.C:; ~leMu.l[Clion. b'llie-.~ti so G11;[.lhel~n-l.l -di~ted ther.lllJ' !!:'i ~h~:rhe c:;~1~if;;1~'" Abnol:ll1f111 regLJJ8~~.OIl. gene s, lhm ~gllh:ne bo.wl: e~iiQn. "N3S also retluccdi ilflmkc Ireilll.'Cid Q[DI'ld'~pdv~illrlmUni,IY wi,u probiLbly be Il':l illliIII 1Ii~.lienee. cQlHhim~~itfll tIfa~ a~m that ,[nl:er~ wI! sLlrvi\r,nl •. ~~(!lCl~ ; f1EN .nH.i .wt:rm~fl m with dle a~~i1Jooy or ,1lefm.i(l11l lut'n-e:in.lheJ~y ~nlgh~<l.lso comn:ibtJ11l:';, ~~IO¥.iever.in riheullilmold ~onl1rmhlgthe sem~llcl~role ol:·syrn.ov~,ocy!ieS~~ leJreS with iIflUlUIJ1e 'fltllClh:m <lkmg,'w.~I'I:uII110lIter ~1!i1h!ruJ~s,:ooyn{wiOl:;'ytes an:: !lul: truly trO;l.n:;,· tltelhlmi!e:~, 1h,t;:5e ~~Haare ~(!pp[lrle-U by ill v~Qro lh ..~t'lrgei:.<i ~)'novio!:"}'I!i,lS miM:~~~ '1ii:;h:ievethe ehL'formoo:. thC'}' ulti nmudy S(:flcsccwhcn grQ'loYill M!U{Ji:eS sh.ow illgt h,lt t:adh~ri~l~ I I. I~.ciI ital:es s,j'l''Cgoab cr.f Sflt'ety mKl ~1r'9~1];y. S:}~lQViocyte l'IWilSio,n imo ca~tilage~~~ke el';trn~ in CU~I;L)re :<1'lilddQ Ilot be,iI<I\H;! hke l1.1ahgmmt ~~I~i1~n~~ c;:d~~I,hlD: !'llaln" in :m i[l v~tm rnm;h:~t iI;,-elis ian.v~vt) bY!lillel"st~izing In ~dct:~(]tg;I!ls" 1, II), M. ~ee.et ~I".$(]ifJi'{e n5,.lOO6 ,(:tOOl?, ~s(li~~ ~he.~t;l pb~If~~,ltio!TIj'i, i'l\'es~ijp~i)!1i i\ni!IT!mlm(lde~s ..~ !1flt~he S<1m..e ,,)8 hlj!]ma~ il. Go. S.liir~~liin.lWlnrrt4:il]. 21S<f;.(ZOO3).

,(and therefore ClUU1O'i reJect the human eel Ils~. ~yltoviocy:t!jS isolated frum [h~llll'ililtocd~~n.hnitb, pil~'r:enlsi~wadethe :imp~<'IfIitedcMriblge '~~{I,1eLe.a:5 iJ!Ot.inil1.l1~ t>yn[}\':~CH;:yl~ do nol t41- '['his iI~JC~~, eceurs ,e\J\CllthOllgh dlC gyn.tJ\'iocy~t:~ have ~fbl

or

ror

or

un.

!!!'::

.h~we bct:.i1I ml~ibl.c

~om~St':Sti s,y.nov.cocytcs

1i:s,:m

~ t hel<1pe~ll,k:: target in rheu maooiid (l'rtIJlful:is t;bC:lt;',a~\;e I1Kl'lg.ent;;n'1ve bL.~1lJ 1w.!i hlbl(;': 'tQ taroJl.'l1 ~ thil:l (.'CU lyu,e, A~. OP;P0l~~1I11i,ty21~';'().1~,vinli1dIe the \

~ idl'jui!1ail~~ll'i ofcadh~riill-1 :1af;.~i ~b1 ;ldbt:~inil ~ uno]ecule that I't.'gubu>es:fi,rmalicm, Qfl'he ~)'I'I!"
~ {1Vli,~~lli ~lirl~VL!~d fUlilictii;JiH (If ~)'T!{l\fio(..1'~~ i'). ,( g _l_, cc: 2L_ oo.l~ei)g~.c~tL?~_d eral, c_leg.fL~l nil. se..... l IDIlellmcl::~to \t.1Ihdl.ue ~YI1tw!OC)1~-esas a taf~t .I11li

~lnli1l[id$,~~lldili(; ~pplil.~ab~1 ilY .of c.!d ~el'in~~ l"';I~1:geted~her<Lp'Y 10 this di:!'lefl!Se s~i~IIiI.e~d!> be e'\.";!ln~l~ed. FiiJlrfli!enli.m1l,;l.il ili 10 I1tll C ~ca1.·wh ich s;ynov:hocyte l"tmct i,o:l)s i111i:il~ln IJ~ Iii'lkf:d Itl dil~~:Q:J,~ei.i,ti.e b~it~r.t. {},I'i~ hill£i1:'i:ilinlllQb~el"WIl i,on ,~!>thml pl1e\'e"~li,'e luot!Oco.l's ~~ Sflm!ll;wilhal mn~ dre(."~~ thall lrC'l!I;ling ... 'e mice th~[are a]lreitdY8l:llllk:K-,d i;.'I'iilh :lIr:th'.iI.b:.. Elec<lu~e the ~l'tk~~llar <lfthrit i $ Lnodel u~ecl by
rh~il)l'l!:!li!u~d

~. G, s. r1rJ!.~1I!1n.An\llmil:~1l\I1'!. ~9. :17811.19961. 4. IIA I!.Wlle",UdlleJ e.[ or•• Am. J. PaliIJ~r. Jl~9'.1607 U.99-.o), !i. G. S. r1rJ!.~1I!1n.F•.IE[rue~rni, M•.lIIl!D, 1M.].Z... il[.er:. IJ.ltt a Gfrtl'!, i"i<[?,C" !V~r:',Awj, S.a. U.5.A ~'l, wall'S {l9~1t ,r;.. JI,IK, ~r,31'!~ "I" A1lhrili$ i\1JIe~J!Il. 4~. ~"'9 (2OOID. tl '7. It'l!iJ/LHlcij~,tlQI '!' E'1P- MM:';:~ l6H ~c!]iJ~, .• 8. IL. ~lld!j.:,N~1(. 1'. M..!!iIt!:n,OOIJN!ila:r, .~ 3:J:.S t20(l5). ~. Y. 'I'';!llII~ni~hrif.r~I .. Plq,r. Nl(JIU~r;qQ.xf.U:S,.1\ ,~,

1002'51,0001

WW'!I\I',.sci encemag. org

5C~E NC!E

VO~ 315

1 6 FE BtR:U.A.RY2007

]1. F. S(!oU

Lorn'g \!'ieii\lOOa~a liOph: 'iill cilaui:(,al phllS.ic:s, fer~oetemic-nt,y ~ali!ihe destl1 liDe,d by a '''Iu'al1ltLlm me(hanic:al ab ]nitto,d1 ~Clrv" lfh~ n·mm 1'Ii3ll'11o:mlled~",~,ce'stnll:~ulf1es~nleg!rated O:l'1lllO :5'i (ihnlps
mad~ iiluoad:S 1i'1llO'tine s:emi(oilldIIlUlU :5~'IIitdlil1,g, chea P lI'llIiOm·tfm perantl re

tiin~, ana ~. is VOllfL,b',Cacross lhickuC'!':ii d. lIc.iilJI clilfU1!~ ·is 1l1!j,.:ctcd.loosi;.'ji, IfCSlIU iUI a e~flt"'illUpcd loop Dh<lil is ~fll,'elm~(ij .ol~m)Ch;;cllfi,dty" A,Ml("l!~h (mJ; look..., for ~lf~t, ~m!ll;-;';;wd l:iy:4c.n;;"i:; CU,·Vc.s. dfICt'l~ eml beart:il:lI.t:1.'s alse, nmt]~lin,g r1.'ii!nI ,..:lIlIll.l"ditld :nnpl:! [jCri-i. dPIdP in ,I hy~[{lro~if.> '~Ul"\!i.l cannot. be ])(lffuet:ly Il'1~:n:lmt wOlllld innJ}iynn UI'I[1thyS3C':JI dr.clc,olric oCOnsnlllnl 'Df zero, haVE

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954

16 FEBRUARY

2007

VOL 315

SClIIENIOE

\I'V'IM;V.sdencemag.orgl

REVIEW
son, JcrroC'I:~!li'k Ulilfwn~hcs are fair:l:y !'I]OO in-'~1II'1tOrs."V~1<!ut a ~,'olmge is ~!pJl~ktl 10 [hell1l. "[~rubi~ ~C!r,roclcclile memories m'C nOI q~lilC fl tlilC), c-';p'lI'td. OOlllr.lC~. erbcnd ~J~~OC'l:cclr.i.c). F:illi~ fa"brlicmcd by Ilcmfll1itet; ilfl ('oio:m<lofl~tJ ~li~J'.:, Ihe SLilll~~~lh~1ir~ h<l~wil1cd d(;:.... i..~il each Q~i;tcrtlig'f()Qh ihelrrli~It!![ti ~truitl im~li(l:l1iii, iec '!';~Lthlhc :iJL~1'1;:;IDIJlC, Comp!elC' ~giim·a~iOll thill, i~ M~Jiin~ in .Mo~(I1;v. ::-l~(,'hfJil'l~dcnli!m!lubt.;$ have 0.3 l1.willl PZT ~mLY (15) {lin ll?t 1Ifl:ltlJwJre$ 11t;'C~llt!ybO(,IIJ ch~mcl'l;r.lzmJ 11)' Mi:lI:rls,o]l @ (IJ • 00· cncasod in 1iI1C;S,orpom·u:;; .A~ZOJ" These lHl· tm1isr.'1Ctmy ro. o!MnmGl",d~11 RAM memory 1(201. N~jt~Llll)l. ~hc size d,i~-Irub'lf~ len nf l\.tlH:' .~~I~lcrcd )[D·nm.,di~me~er c.[!lPlLdlm~ ~'\'!!ioc:lI ~ vleescau oo;'ICh~t~,cdinhlU'l,lghdqjl>f};S;[tl:OJli w.inb :111('11 .:JS as5~1'1l!b'1c:d l1al'lO-i ...lam;h of PZl' ln Fig. lEI ,It': ffimd:ilbk~ 21.100 el\:lCI!rO'J1.:-l per bit, [md rLl~~Y :!>t»LCt1r&,. :;.~1:t11 $\lbJnlcD'QI]lo;l.t'f t;Hioon S!'lilL:fi\,l:>. Tmc !il]iIrll>t'\.iZC mopol:nizmocm dTocl!); W{'fiC' Iil[)~ <l pll<l~ d.ing~] dit is not :I. S)'s.lml1 ilillh(~rnm~ J'cgi~w:rcdlilumin!l poresare a'!<I:ibhlc. I"~gllm 2: once l!ho1light, hIlJQfliocl:ly. to ·OCl'lJlf at dim.erl\lo:l1-,~ l'qu~llibifilllm). Fig.ure 3, C to E,:\hm~ nue pll!1L\C :s.ho\,,'~ nlIlClR~rnOC'~cr:~aksI:hhl"i,c;urcd .<'I~ ron~ll!08O't i~C)[; ig, 21'\ I. limn lI..FM \\'I:i~]]lg (fig, 28}. ..urn 8Jighlly ~c:;_<; than 1.00 !Unl. .~I] llo'llllli'iplHc\I.u<i di;]gmUl1~ff'oJ' .1.1<IImrcrmcloctric~ W BLII1iO;).. Ilu:; 0!,I11:;:~d~,hcll i~ cuh1c (rm:ncI~lric) $ The. dis!f:ibu~[PI] ~ij,Ulotinll fhrr~~C:1im~~ pc"I: thmllgllll r:OC~I:';ookm beam ~.flB) {.lFig. 2, (' and m!(! 11Ill;: ill'lNlar to;. t /9) l,gI!I'<l~IIUJ :JlI]9, f~rl'l'Xll\t>etric C\;I'I1l1_gc 11' 0 r~~lam~~ ofa cenn uQ. volu'IIlIC F Im:o; I)). SdC'lt!~sl~ ill~ Bdtag~ h,,-\ C~bf.!Calql Cflf~lrl.ir td:n.." 11..~111blingan M&M t',md.y). Ay, dbm~.~ catcd 3D nmloo!IUe~uv~. including ~un PIB-clill r!tlQo~~;mdin,g miCf-ol'i'ngs I Fig. 2('~. :Uld a 5·~lJn rcd\I:c'.iJ~I,. 1:11i!:r."':IlOSj~loon;lis ,III 'C\lhi;e: [rl1df:lOlt in!1ijd~iilm~f PZ']'n:mor-jng (F~g ..2D), §QIU~io:nr r<:l!'rnelcl:;niic. but Ihbi!'i j'lt"~I. <l ::'l!.J!ir:m;~~ dl(;Clli~lry e.~r~1 <l11d Jfml m lillil~ 'C'llilc'l R!;';ill liu~i.(l :;.iizC' ~izC' oo~i'ctI, \,;}jl.hi!1 n pore Clf I~IDJ'oO"!:Il~i" Si 07), eITC-CIL<;,!:nay OOOlm: for diruloc:ICiI':'l~tlal. <!1lC -:;J nil}. lll\ullJjl S)'Tlch~O!mll SOIilIl;i(!i, (Fig. 3A 1I one can ,Magll'i~te~1~{~lIfks M{)S'i. readers ilire :lflllil~Har wi~cl CfMr.OOUl !lam)", ::mociythe '~OU'IUlll SIruCIlJi'X: or ru~hvcry Ihin lilms, tubes, These .;J~g';U~TI.d~~1 glJ!Od cl;)udt~IM. (',y."ool:;, whose I ~~li1Iih,OJ]ufIJl!S t(!l1tm:iltoo 'liilDc:-llli' TIle 2004 32 -l1l1Cgllbi~ (Mb l FcRAM s li-olH (J( ~o;:nlTioon.uLlc~ing, By campara. . wll(l1i ul,g !Jebv ..-e.n ;,:,h,]chie p-olflt.izaii{)[1 P11~tdI S~UiIl~lWg ,WIT) or Mill'iU:>hih.l (Sill) are ~i~~ l!:i1iltCrlite~]lic Jlli,;ado!.i:' !lie OOiii"i";:.ifCiii~ tii,,,&,~,.:tie .li:aJitlO:rl'l !LCCc... ll'leIl'Kl!ry lM,IRAM} .s lEI d!..""l:'lu;pn'l~ljL (40Mb MRt'l.M fi;OHI V31. and Fig.l.!3 sltows
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OJ:51un :;Ir.:lp·;;;('ill ,P'zT <k;,ii§lK M E(" 11.a~ElrOOUJ("'l.'rl 500 million tmil:;, of srrr !Iiilil·fi~m inll{.w.n..rl 1·,.'Il1:!IC~I,)l:'~rit::s!( 1 hil1:irni, im:lllIdinl!;. IOOlrli.wi!~ai" il1~ B'ST-tlll·G:JA~ ~Il}ll~.ilm:» lind ~:> ,H 0, I J1un stacked 4·,k:vel,;D1t.:I:!I d(_~i1'ln. TIle S.)'II1~C1Ji]", 5·ul» aeecss'~jrl1l: "Trininn" ('"C.lm ito Ille H'r:;;!Urn1,. ,,'{~hll t'<.1Cilc 1J1l:lnmy, On 15 Deik cember 1006, :lFllIjiTsu .~Jlnrmm:vd pmdlll(:~illn of a Mat.dnpl'tl, HiFeD.:!!
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dQ'!,:I.'rl" (si!lln~k:rornc~er !~lliKfigtl.1!~hy) and "hot!omLIfl" ,(~elr-as,~mhlj'). The clrC\'cIJQI~llC!n ,('!fl~l-CSOI~Ol'('ilil:-:

IF~!:iI 1. NanOS~HJI'~Ures ~111hi:rl c;eral!Jllic ri~~m~. Strilpoe l!'!'idthsquI~r,~,d ~;:Ulj'~dedby di@lIi!i!li!1 '~Ulhic;kn~.s:s •. t <lnd CiaAs: 11!:il$ \I',erS!IISIitlill t.mc~l1iess: d, $hewi~!£I that. ~/df is coml.ant fJlilreflroOe'~edri(s PbTiOJ ,a~ul B.alli03 and 1I1i1 Oii:n~l~e R Sa~t. PI'C$C'llli!(~ cxpcri~1en~list .. \I;'iah lem,anll:l haU'omagllM2t Go, demOI'lSlr1!JliiFl!D <I uln~'IIoers!3Jl, l.aw Q\!'@r 6 d,e(.tlde~ (9)'.~lll~ PIOIIli 'lfiew ll1IiC1rogwa,ph ~r 3D p.b(~s fur~I'l.Claucr. 11m], olhci' leci1l1'allliodo.m<l~M ~flEllI1iiO~ (16). (e)! Sc.anll~fI!Jele(~rn.ll ll'fIil1:re;oopy (SEM)' Ji'I'Ii!fogl'.3Iph (!ifS,IfI1l'i4:·T\:iI2iO~, Iniln!OlUbe~ .u (ZQt (D) SEM W;)ss,·sectito.!'l<ll vii:w of 11'5 Tibtii!1!2 rlalilOIUra;v !of 1Ft nanolO'!'iJres and U5.L liIiq~1e> lOr OO-i)ljnjll n,lmO\~'ires hf~ve o,bliql!eill'tddence Si, AI·iOJ' (iaN.

tru

9,55

REVIEW
lin<J,gn~~i:anl(lli Ail ~ eretho~~hl.[00 be disco\'~rcd iu FIilJi.cc· in. libe 1910s. ~~WC'!CI:. ~rllCr slJuiiCqu:cm dfllfL ::>.1(JW'(:Jij U1illlil~ L cn~ was l.oi'lJiddcnin the linlnlCriab used, It wasbeJi.cved nrnpu)S~iYl~, Jusilil! e i! hi~h~~ ill·;n~fligrt'CI(Jt.'1ectci.d~y" R&D until I·QH.; wn.ml MO!'ii:O\'!I'gmups =lltITCtcd IJK: pmh11.'In Dzyak:lr.him;lkii prillkrod IlfI:lI,gD1L'l~ooloclricjl.)1 in ('r~03' Ifmpi<~~yoo]]fin1ood by A<;;[~m'. ami :IL\;O ·1~il'iZot1]~lg'lm.~m. quickly Yound by BmovikROlll(lllo\'_ ·[ltlrl!lg Ihe ~97J!l~. a ,,\!!Iroc~ Illlscmrcll l'Or lim"w ll:l<lgJlt'Lor.kclrirn r·:mLU~litcllujc.s,·) W'l~ led hy Jbns Sdmlidin GcalC\~. "[11.e 1:~riIl11111)'
f'{)1~ri",!.miohl and Mis nlaQ,'[he- firstocl'll1!1 il> 001 ~i~flc;..re'\!cr.ml 011...... rJ.rlftl1itllcre .. tile I,DI1~r l'I'IiIgillCIOdcClric cfbCC'[ Jue W 04 Y3U~::;he.~~tlOV·Ct'J (die Neclileniper. T mllu:c) (It T.: (Iii.;;; (Ur~ 1CU'IJ}CI1liLtli"C because J. <iii!> = fl; how(J\ er, Ihc qmtrlmlk lCIil11I due to Il~ Ls pmportuml:I~. ~r.; <.M·;!> :I11K1r,em~:i!l~flni~!! wllJI~ abO\,'~ TN ~!II'llitH T~ 3 TN linl mngm:lI.l'i such at'i. wJ\i:\I'C
l\C'tir&l~ ian.

P' is eleenie

HaM II E!ll ill~~IJ]h::"~ plntlSiMe va IHe fOorthe ana is()uopk cxehaegc jill o)U1JI1iriil:01l. \,\,'itl-. off., dirlg@t-ial f.ll,i inr ~ter MI1+:l: oOOlllpounds) .. This Fm.';Sooumodcl h~I$. rcccinly been discussed rblt
IJ8NiF4l1]JI. Tml[,[~ arc' arti rm~~ . h~lllI~ ill ~nt' I~i::('!n[ p.r.ob J~mgn~'>'tooir;~trk ~iromltUI'C_ Sloon!l, :m:JglJu;:~Oo!.":I· 1,~u:ita:I!IC.odioCl~lcetric cOJ:15lmlU change with ap( plied Inagnt'tic fll\~d) eml occur !llII('!'l;lly thmu~,h Maxwelll· W:Jglll~r space cha~~ III ,my m:Jgllltlrmu-si~liHll!1l~i~m:i,11 Uh This; b not .intrin~ic alld l'\lC]uil:\1'S ne l'emlc~ocH~cily ar.all. The- lOrIcc~s ·~'ml be h~g.~ and 1'Il.:lIy c!(pbilll lne 4 5·~JI!1, .mmn~_~>'il~ ~ndmi~lm ~m IIn~l!n.yl chmn~iu~111 ~pil1d.(In. Bn;nvn. Hfi'rnrckh and S:h~lI'~kn~<rn :'llmwoo thal ~ll'i;l~lbiln: {II r~~I~ 1l.1ctgt'Kl'OC~oc~ric:s.lJili:~Ni. hili~.}' Clmoot('xc:ced 'I!ItG' pl'{l(lu:t1 or In1C doc~lIic

BaMnFil wi~h 21) 1*U1I~1F",pill ol!li)(>fillg). To sw.iR:h ,~1.1mgn~'Ik dmll,lilll \"'iD~]an d,m Iri,C r.t'ld. we IlcC'd tel kn.ow~mw ~UIICh. th.e: mag· ItctizmioillM depends 0.111 .he pol~l'ilz;.~ionr_ l)OCi'i·~\~ung, P illlnutn'bDC' ,lJ by!. ItO, or ~\'kc int1ii"'':ffi \"1h'" I'I!lill.iSl~.~ Iogit;: d(,'IJnCr~l!;; for C(lI11~lltcr bit slaJ'agll) Ih:)!L cmlrd I:Nl gL'L;'llIt'J (11<11] I nO%:e The idea ,.llI:H rmroe.lc~trkil:y cOI!I~d hi!l<Iry_ A"'. mtch, il \Vll;,; hlll'llCr:Ili\:c to ~~I[m .)wi~chr ;Ic~~l<illll' cause .femJln:lglllCl'~i~m ltnO% cff~ctl ~<1 i[)g QI'm~;gnU;'lk:>il"lt~ \vilih il]locuic ndJd:ii <I~dlcw or ! W:L'1 '~-"ll)~ai.!'lC!d IJyFo:.1:;tnd Scou il]!'977 f(!!'mck'Clric :m"tc'" wi~h nn,lgl1c~it;: fic[,cl~ and the ,llldI eXCl'l1plUlicd wilh IhC' BaM fl~ .f:ulllu~y ".\;l il1le~d.iml hct'\.\'OC11 nl1a.g!~~~i,c.i1Illd ,~It':(:lrk: dO. !VI n,. N L a rid Fc:)_ By p~OdLlcil]M- aweak ji;oU'.
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since 20{)O. ~'llmrlmsh::il1g liiM.il:!,!!.m.c:se~lwKl·~rbi.IJJI'b Cll'llljJOlijl.tI::; ,( 7. /I). M~L!llidtJc~eel[ic~ty ;i;\!II(!i~~ ci1ill11r~.izC IwO
[Cnll!:>. wh iC~'1arc: ~i'lil.!1Jli ill Ett. 2:

D0"1 hJi>hi'riSk ~i.Mmi:ytl til\; fm{ielec~ncpoIHriz.ij D Cilli proctltO;: tlti;:; i.Oii effeet i~lI fi;UI'l1ICPODllt symiili;~ll)' ~Upf> (c.J;.• l' ill BaMrIF~)lJuI :rilouJ~.hc:p.> ~..:.g.• l ill Bi.l("oF~), The 11111g!le0 f spin (mning 1.3 n~I':'.J(I = 0.1 T" in '

!l.!IPjl~'lm~~h Ihc all i.'>tIiropic ~:l<dlf~I:Ig>(:"

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"'-'oulcl Wic-m~il i.'ike~!k wrilC QtR\""irn~ ~l~d U'<I,g~lCI ~C<t(1. e'limit;: i11<11' the need rOT ;-~(k'~t!1ilc~h'C re"d i:ng '(.~iI'l{ln::sct~ for [JlJ:es~lq-dJY P·c-Rt\Ms

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c1lndid!lll::. rCO:tli:;~Cttril(;o.tcr®n1l1lgJ'L\:lic Huooo(:;l; :SldUl~ I\J{MFr.h or K"FC:!;f1il! nu;ril InIJ"h Ill~tll\' :>1l!IUY (j~ \.vh!l:U1i: A is S, or P'b ~lII1dM is :any 3d 1TiIc[m,i (['ii, V, o, m Fe); rl:K~i>Cmll'C~ppm'tllll.l~ Ir~'mIcklcl!rOC wulll didot:ui:c iJClnL"m~ :5:50 K-< 7;, -c ?.:fCl K.
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Fig. :2. il'l'B 1I11<ill1ioodellCE1. S,£M ptai'll view o~· maglfilE1llOeLe<.trnc. mmpoositf1 of CaFE1~O~ ~i~lalrs in a BaTiO~ (A)' matrix. U.n.~t9~ INanodomaiil1!5 pr'OduCied Dr .A:fM n:amnl,lfiti'flg ,on '(50~. a.!!!., arbilrary I.!niK ·(C) Hedr'Ofll fill!: iiO gr ilpl:i off fila n:o~or®i~ ofJ·s~i"Igle·!(rysta~. ~aT'~O~ (15). (ID) Microglrap,1lr of uIlM.a~·tllIFIOtomttl ®f ~Zrii"l'!:~~eal me~',p()r{l:Us, AbO~ r;lore u.n, EDX, e'rler,gy·di5Jl'~fs~ve ;N~Y <lllOllys;js"

1m

M~! fJ;:fI'o.::!bt;'U'!:c.s fir il.!n~ liYloC:~l;.'<J;refi' ~rkj:.order wi~h fli..kuri ..... Ilimu;.~ha~ .iliC ~arnllcl cr ;;unth'i<JJrnllel. respcel i\dy. I lowcver; circul~a' 01' tl)l1OOC~11 mUtcrns C:!.Il ocom. A~.Ki~1d li,r:::t pJi!Ucd out ,(If". fcrro!~l~~1.ei;S iii! EIi)~W$i;rJ:: di~ll~CI~l'S will orucr whh fo'!.l r ?lr dQ-

16 FEBRUARY

2007

VOL 315

SClIIENIOE

\IV'I,MIV.sdencemag.orgl

REVIEW
linilins, i:Onrlb~g 8 circle, This kinti 0 r I[)IPOIOgLc~~ SJ>:UIJdefect jVoru!x) was u·e1n~d thcore~iClilly ~v MCt1111~1i (2Jlin~ terms of WllfltIJilUg nl~'I1b:;;rsandis oOlnrnonly round iUnlilnimnmg· IflCr.s. neimlinwb0lh mJI~[mUy oocLMr~ilig lil~nc:r,a]:; ~ and ~J'lJlh]_';~iDl (25). A oomp~otcly d.i.fI"e.OC'Ht orbg~n lal' C'.i,n:1I~3ror ·tOluidHI dOIll:lil'l'; oeenrs .in ln~glOC!look:ctrie.".
Gnbulk. flrl!~ <UluIlyzcrl .ill dt~m~j.lby Gilllzbmg thi:.; iz;. 1'C\,.lJi3'Wt,,;l by ~C'I,'crn~ al!ltiarm; t 7•• '1:, and ~ubject to . Illl!1.y of 1UC'Cn! .,..
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writ~ 01'1811 ~nKl

magnel

if:: I\".ad

(1')0

Alii ~l1Ii~il]@' Theo~y

and hy S~m~i~(1''''"

p'~pCIX. 'blJ~ nm yCI loI~l,ml.hug.~f!~I~yo'll.q~n~d. S~ch <ll ~)"'!I!l~li1ll otr~l'S tbe pt1!'1~ibj.lily of \f~ry hig~l"
dC!'L~it~1rrb/inl~
m.r;:n10l)'

,..nd

~1,{lI""gc with

A vcr,ilable ~Cilili~M}nCC 111 1'I'l'\ioirc~.ic~1 rCIi~ dOClricily has n::illd'oc\d ~mln 1l11C work bylll:rn~,;" 'hmdetb:ila•. R.:Ik. Cnhen. K'r'.ilmltcf, .mt! mBlbl~ on abo iuitio !1l:ot:BC~ Those !'L'plocl:! Lim semida~ic~,1 ba~l.... d-stiek [moocl wi~h Ii.a~~y ll::mUIII.1l n q mechanic!11 C'llcllllal~ml". Th~ have r~l'[wt\d pm .. ~iC'l.u~m~y in!iigI1~Ju iLl 'c;!llctl:i<ltlllg ~QIII]pei:'1l1i\IIIU· ~ ~1.mi!1 pha~ dia~mm." iorLhill ~ihm (.(6) .. illl ,,\!\I.~khdHl~rg.C' Mrnl.lfl<;'~~~111 I'rom coherent ~cpilaxio.~)gmwlh m1.1rl sl;IoolmLc-s w.idl s~ightly

The. Dl,IIII!]OOrof ntoll.'l" till 111t~pmgrnrl'~:'; :lire :llilllflO $lluH C\'C'll rhl" 11.:mOOtwicc .'iimlu~atifm\1_ (h", ~'1 is [11D!:C'a~y ro
wO'rK. i!Salt
~~'OO,

di 11'i::rl'i"1Ii~~lioC' co~mrlt" froHii U~os.c of Ihe fel'rol!ke~,k fHIt! (f:i,g •. 3. C I('IE~ 111£ l~e~1J1iiq~e SllICi"(:1"S rron~ tour linliuuions: .(i) II ~'ii!!!rnHy ~\'(~~£,rorlCroartllied field f.. f~t1roughW'll,t'l ,fl. hj_1"~ done [iul!iv.;lte~d Ci.i.I~~.dmioJ~ ,tii) M!.},'i~
1C'll1pel;:JI~.lJoo. (iii)

l.m"

~.uIl.~o'1lOl'a~!: thC'~mliJGondllCli
fCl'roo!cc~uic oxides ...'!lor ..'a,cHm:-i~" urnI: g~'adients

ng ch:JU!<lc[i[!r
.nlC<li.'

or Ihc

dctcc'15 ~ucla 'l'I m::ytt.._'iI~


~hc electrode

i11[~rra ec,
aile ob"the nanst 1,0 come

~'!!_, ~I' f!,'••'* '" • ..... . .,- .'1 .. ., ..... .J J - ... ~t··"'" ..r"· .... -' ,.,..,. ,.1 t·.... • .•..•• • »: '.
,~ ,.',

:~

,•

...• ._. - ..'..• .. '-r. '-, .... •


• '• .t. ..

•. •.•', 'I.;'
~ ~_.•• ,
it
,#
.

, . '.. • .,•• it·.I~ ,• ~ .;,


: G" - ••
• I' .-.
-

i~llll()rt:.1m ,ql!l~.n'~ili,'I~ IMn~ rmm~Hi~s~ Ile'llllImd~lls; is dl.e p=liC"~io:n slm"lIl il~ fig, ) rro!nl 1),~'tl"N[!\',~I1d T,;!~"I!lm(,"\1;'1 "li. (2~) dm! 0. nc'w ,w P~I~SC' OCC"III'I'S wit!! pela riul~iol! ly~ !itl!l, between the {lllnik ra.ccc\ of

:BoTlO'J (:1)'.$ • •al, H j,)o'm:o;;;l,~his Yl1{)~I~dlmvc


fIP::JCLldINCIL~:!l)()mli

fl'

,.

"Wi

'lIi.

bCJ!t1 new l:lh}':;ic:.. UIlJo:n.u nltuely. Ili~ ab i l~ilgO ;,;,urn. of 'Wliifldcrbiil'L I rig . .lE), l>'howt> Hun MLh u1tJotl.r:ls of Pcrl,'>C,":Iud Taguntse \ 1'1 at. ~.fig,. 3. c and D) am: ql'lfU~iiDmivc.~y wrong a 1](1 dlfn lIicre is 110 t~("
P~I[l5";(.:'r'),

~ ,

' .,. t'..

_."

'

Th~s ::;110,,",5 'Lht:plJwCIt of lIb inai,tio m(Hi!.:·1 i~llIg QVi."TH1~m; pnlm~!.ivc 1h.:!,;'...:iTI..::r,gy th ...:lri,l.'iS. AI· i Ihou,!!h lUte l\:n5Cv.T3,W'lnll!~VmedI'll W~Zi ~llnSl!loc~~fiul ~lQrh"l~lIm Hn;!n;rtc. U 1llLl.t-e s.tltlc~\,dl!l~'1{mnMr~C C

d~~~ 450 UJlgjc 'fmrn C<lc'horlJw IW~aaNe~ or .he


rhfl:!'I1Ib5c JlI~~~~ !11eaninJg
.>3.

is .~bl]O]" which in: bulk has OI~I~' whic and .ctrag~lIla~ ph<'l!ll;!SltnmtIn II, i~ n lin C')\h:ihi~s 1)fI!h au 011 ho"

rohui7.;)tim P lies llt

(,*,

.cU;1M,fI,~;d 1f)n.'C~lrsor

III OJ) ;;m,d ;I r~fI:!nlmbcd! trl ..~ rh~c (f' olo.l1ig r I II H, For BaT~O) .he inhk!l model prulic! lons were
thl]~l) IJ~W pllH,'Vi,:wou.ld be :>Ltlbl~ CfC in iihiiilJ-film fimm til z..::m :SiP,;5~, 1>111 ~ih ililil io {~]'C:lIh~tirnts (}P), :;:lwli,',:~1:11 that I.hi:>i:; IiIl1likdy. Th~ Il1tJ.,',1 exlrente C?iperilll!Jllla~ "'::Slllli. is d!~'l Sd":iO;g.. WhLCh ill, bLllk ~:;;PJm.

",ros~.... ,~I'o:!1~

c!\l(;Hic dm .... 10 a'b:;oh.nc ~0Jf()1 tan n be made fcrmct~x;uli{; ;n. room Icrr:lIHT'.JIltliJi; il:l th~ll-mm rOIll] (JO, W~!.~i tile I£iC of a 1!~lth!l:'.1IlC,,,. sLibslnLl~lli~~LCri!~I: DyScOj.

(0

-1.0

-5

(I

M isJlii:t:S1irain I(~ O'"'a)


Fig. 3~. Silorage cie!j1'(!e"l. ttl) IlftliI!8:fI1f diffuse .x·ray5~a'ltofr~n g rlNd~le5 a MUf1d' the Plilii03d03;:.. Bragg Ipeak at 5tl9 K 1(3'3t fh~ fOIJ FXlite lli~e reflectiens oiIr~'!l@t Bra ggl ~aks, IbUlthr~steo\lld .lIre d'ue .~!;) p;e1n1ll'd~1; ~t~ diQlIlai !liS·~III Eli:! ortlm!jDI'I!lll ~ir~(:tuiiiiSI oj! ndlih:e 9looiiiietr~ ~v~I~~h~t ~larillatiMP li~:s QUIt t(lftn~ P~!II~ ()I~lh~ filrn, ()R:~ PZTs~lfa'5£embly. AFM Kalil ShD~1S P'Zli en SrTi03 (18). Sc;i!l~e.,2.5 by 25 pm h(!Jm leU to rng~l edge aJllid h01'll1 to;p ~o lJotmm 'E!illI'!J~ o,~ pa nel (( to E) HariO} temper a~ure·~:nre~i!: diii3igramlwil.i1li3liJ ilfl~l10' leliiU'll!ii(J'9) (E) and rew ~l!irom f Pemev ,am'! Tagi1lnts>Elv inl 1998 (28)' (0, and 1999 (5.:l) (D,),

lWt-'I"!:tu:"itlJ fNlI.!IUUJlt'IUJ, ferro..:k\~1Irir:l ~m: Ilfm:m (l[m\P~;ic:nL'rlllmJl] fe!rona~gncts bCC<Il~I~C e!cctri,c~ II mCa~irell1!~':nL~ mll~ he m;;cLde (iI~l a S;igt\~wiil:h oolli'Oislillg hVO eleelrodc~ P!\fl"~'~ <1! d~>c~cc.rk" The intcfil:lO;;C is \'CI)' complex, ill\'Oil \1ii1.g 9Cn.,(!lIing: jgl Ihc i1i~c!<.l~(J l)~ftd j!1Sla&lmtics <It <lb~lt ;; nm of eliC!cclific!h~!u~e-;:s (32), The 1lI.~!i!I~iill,g 1cm:1Igc' Ct!nt!ll iii! PlT rUms

or

can be Sdm.1~ky-~~mil.oo fool:eor

9S'l

REVIEW
Frenkc1.riLm1~~t~ lllms can be' fully
our parti aU}' cJ C11~l.c~c(j. 1i'l~crJ!!icj:lL1 phcnwnnem; were rccC'il\t~)' If'Cvkwe(i

{.~,~" Ear.il), problcl1ilS ''''i:LhI~,t~~u..;. <l1ii~ P


'll'Cwv:-;l;;il!.'-'Il),pochhm; {J'4) in fcRAMs. have been m'~I.'OOl1liJ_ Rcpl:llocm(l;lll of .Pl. 'by m:.1dc d('Cutlii;lit~ 1£], ~lIiimim[Le rm.·' ~igH" ~llil\~~U.\d a lOt ,t)r good ~;url~. hlU Mru.>llL"hit:11UIIli Panl1lSo:rlll;r: lind the metaHk P't eloctredes 011 SBf to he vcry ~ntWbclillry_ F!!!l'mek't,'il'h,.slei/1Cril.lftiCC'Ji_ Arti~ Iii::: i'8.11~' ~1IiJ'C;'rOO r~lIlCI(!'CU'iC:Sill[lIC"" IM~iC(!~~~i;:h BaTIO~,19rTIO~~J!! 1L'l.
iIXlII\'crslon

liflily dll,;;tQ

100

'1~'

pmd1ucC' enh:ll1!ood polar.i7_'lliolll P and dl~IcC1:ric Oi.~n,<;l')nl r _Pbli:OiS'l"U1Q~ is.I},,utriC:ll!;:II'ly noJ<thk' bceanse it h~l'l a flear"pt.~,1:L~1Imuicc OOnsmr.n anm:cil ,(JJ}. SlluJlti~ingly. the ~·-rrOJ 'iIDl;iJr~ izal~Utli]~ B~,liO].lSr'Ii03 HUI~rrm'lll Ji~(jliig 100 II 10 II ~ II01 iii tl,jC lreifl..... ed ~L~p,:rMlll"'-e jon, ~~'i~~iSlr.;I,iu eu{16, ergy 'O~j~Q1~'1illg do.:t:lro~I~Ltic5" to. rch:u..'(] I.opic i:;; pocrj,l!die~Uy poled t.T}'$la~$ of LiNf.jOl lind Olil~'If IrlOrll~ IIICflr orlleM l'Ilil[CrUfLh, (.J 8). Here, Ih,,;~u~"TllImi!X' OOIl:>t.1.:S 11Ht (I d i fl:'ctcni: dh.:lrl'li(;a~ (;lm1JJ!aSitinu:; ~11:~"'1 !is BriTiOl!,/SI'TiC}.t, litll {a +P anti P drn'll:;l.il1!!:. prtlv.iIiill!,!. C'flkklm pl ....sem~ldlinl:!ll !i1 J:' I1fmli!M311' optuC&, Tht1 qlle.\t['l,l 111:C~1lI. to llF!ildlHJC':i;llh~ is nnicrnIilH;:!W wa\'e'l!cllg,lhs: a~d to lin"
1

T('1C)

200

d~rl1I,uld df.ltll~iu. ,,,,iduh:;: auo ~ahilU1..y, f,)'lr1';,1IJ!h~ ,~iiIgh· ).:~l,!/,~,Th~ \l~ P QffocUltOd ion h:;:!"l!o~UO CU!llaflCli~·m..
(!~(,'driCl1. \\,'n.~ l)i~~li!cfl,,~ I1y RmlliC$b's

lfi~g.,.4. Apj;lU!l;1!!llQ!l:i, ~A) El~(IJr'Q(;;1lQri~ eU'e~t:ilfl PZT (39). ~IB:)5EM uQI'>~se'1;iQ!1fI@f 32 ll!'b5ams'ung : IFeRI4M .lll!'L1UI' in M:rr:y bud. ~'i:lr ern!], le Hhs c 11'11 Plan vne~~ sI'i:CiLlj'~ m hut .il;,'C~ldy ~.'\;,~!l(lcd!fI Sillgl!C'CI'Ys.- (52). (C) S.EM CfO"lS":!:eClio9 of ftithenlui'rll 1iI.ai'icrt.r,e\i'iIJ!:~es, ~i(53). '(I[I) ,e-lleamlli!hCi£lr,~plTi~. cap1ildtor .maJY (§4). t,lIb or B;JliiO,:l tJ'J') !f! chnmctcJiil".(!: ~~!lgh:~--(:!)'$t<llcalxld~ ntflhocbcS,<; m(;~I;;tlrl.'tl AT= 12 K m 25 VUo!,:rt.X"i> 3jQo 111m ibr M::Pt1Jftd,Ca~ti"'HLimi '~nOIXr;:;;, The C~Jll,11fIlCi.'l;:i;1dl down IQtibu~t6:; urn, lhi:;. im~ :-hm'l'llllhm dr.i~wllt:ldk:;, lin; Ihrli, iilu; im.cll:>c el..::cmri:c::I:ldd::; thin .I~II1iIi> Hf(}). l:CeliS of C!lJcic 1.::m~l!I:mullJ;1i: shin or didb;~1ric q;:oo~iug{Fig,.I~A)i.ri1 !(:rTlJClk:x~~c . Gmsc' rl1li~'iI:OCl1tdkin~ ~lirid{i{"\'ir:c :Il.ii hlilJ.: IIIJd Ih~t peak I:IIUII(1!:1Ii a it! it);lrul1i~ic. t\(}lIi~mry i]]!::, to 11110:' Mlllicil,"l'll. 1.0 dc:;.i~1Ii aprr'otO'lypt:' cooler ror 'dfliJ ~:rrcct if,l pooR:1i ill III[11 11i~n.\.o;liiun in hulk. CQll1l'plllJ:!r main fi';I1~'1~ prev i:UU15 (lt1:JW~llliun!ll1 wi:li.clou1. M:oreu\'(!r~ 11mIma!liIbiguUlliSth!X)L'1(J~ill::J:1 nicxild Ete:e/l'tH.r 'l'mi.~;Nir]Ji firm, folU}{"~~'I1'h'S. TIn: :S;I"I® n-'1I"e,r,m tI!IlIplicalll01il!5 I;'(;:t 'd:J~ul ~~mllC~,,"ClJi{<oi emlil oopi.o~, d~"c[R:in:. hltl5 been a(lCcpDOO Ru' til!.; basie !JllIi~.ioJ] rUOO ..'iS5. M:i'croowpil: I1ro(;htmii:Sll~ii I\:\I1Imiin (Jtb3I;ab!k;, AI[rum Ehcir :stJdhvcr; dLUrin.g ~'I{ildlil1!,\ h!Ji.~bt"t'la Ek['II'(J('c~/(}l"ic amiirlg lm' m~h!fi"r.lllU!S (md mh'I,(J['I(~"ll'iclIMlmil. Ihli fi1"';[ li!m~ (crrocitX;lrle:> knOW],l lilr rmmy y,.;tll1>. Fir:;1 d iseovcred illl uhH'lI.l~] :I.I~~4; off = J01V/cm "m: J.-o.::qUlirr'ro, Ihi~ b; OIJ~y 50 V lK:rm5 :50 11m, n:;n:My (~bmimiblk; in pil~'lt!(lnt::lnm was c~m be eooled by ,LPJ)lyiBlg ml doeui:: :i.kh:l ~o Mil;hj~n by RtIi>l:Ul'-'li~ml, 1.~lifi, Ils!1~riIlVCSli.!,IPI.lwe.1o:.r~,;'I]::;ivdysJt ('c;na~ ELlmp~C1a !;.~d,fI:lm:s. '~h~~n, under ccnailu eond i lio:rh~' is I.crmed i'h,.; '·docltmcsJlmlC' ,.;iUt'rl:t ,. U::;il.1!'l IJLclllllooynlal1.1i~ 1JiJ.i" i~au mlcxlPloi,k~ cl~\luoc 3R.':2I, :1'01" which roor h~ RI.~Il,.;:r~~h,;: lK::k,":lirc(CERN l' ,t 4.1, mJi1d illl N c(jmnll(.'ttia~ ili~""~],[lipmt:'m. {If llili;Il~!IIIJI:UII;hu~hfr.!LliCt:.,Pc!hmd, .lkmct·[lnd 11h~U!lil.tld S'Ula~ The and 'nOO' idoC~~ ~J1.1lr0P'Yclt!CJ~a.~~ llIurlcr applit1~ I}I:II. ikId E, one r~Jtd,,:'L[~mpcilatllln:: cool illlg of 1.i!!l\V(!f~'llkm"'\'.l"C' dINi.ocs c(lll!~dbe made wi~hin Il!ll!o.ty If.!'C' U~ "''''.~ ~och!"Ol~iI7~.•p,\III!;;cd ekdm~l a fe\\' ye. rs, S,1In1'sur'!g C1(p!OIl;,,,(~in Ihe It~!JOs.Ihe a sotlmC' ror aocde!'[lto!'S\\'1I!i\ itl\~\ig<l~oo. inclltllding 111eextension If! n~QciJlmmc!ilbtc S!~I:C~lI!\'S. usc of this l~l<:'!~mCg'Olt fur n~!'!(IH!f'l(!gruphy to] :W()S~ [hiis ctWecl was {.\UTCIHS f1l"tcns ofmnpc!l:s have llCeU OOJ~~J~H.x1 'VI1), MOSlrcccmJy :JJ..~I: by PlmC!TI'!<Ufl's ~~iP ~N ,~ kb1~~ flrr;o!k~ where ,~,ls ~pcd r.l'C h~i!~..~rons~m~tPO!<l!~·7,.:'1!i.L'l!f1 wiilh SY!ld1ro!li7J.'\~, i!l19l100.~~~ le Pli!~~ !cngll~ .P;;!lnd ~ lS I~~c'['4, coollicie-Jli inlhc r~c"Cener~ of HJO IlS 10 .~ ~.s, ThcS-C'M'C sl.!pc"rio'r to lils~o<nl (4J), as a mi!liJ~l!med source or ",.-raj's I hCfmio;llk' c<!tIi~d~ ~n~lill! thc:yllo'l\\l' higher G{1:PI «~T l~)fjl ~113/]'1, o i~l,li .do. 1'1:5 Q lien s wld W':!JIiITi.i'iS. BW,'!.t-rm'i'at-I!If.'V,'If'llfl(! H~floil'l.;,mm.. Bm'ium~~IIn.'!~t dcm:iik~ am! Hrctim~ ~md <1100 have m~'1!i.~l!redrR:UH po mil1cr than di:l'(lct~)I.Elcctrocaloric 'coo! il1g was s~~~~~cd.~ ~U~G:I); I ~n(JJ1., b~~! iII)SWlI1l!UUl-flU (illccm ionic CJihoocs m;JLlire .il! til;;g~aie <h'1"'Cd ocrmnric car~d~Ol"'Sare a 'Il1n~II..~ .. C<!L1'l 'Ute effcci was sm~H in b'!.u~k(a fc\v n~iLHkel;in '.\lllrllHI p), -rile feoodoclfoc eleciron el~~~Ui:'.rs. b1100i.ty (!!.m;1J~o~m fonhe L'ilIlkof'all COll(_i:ctl9CfS ... per \!oIU. R.ccently 1lCi!\\'c\i'~r:.Mijlchenlko (.{ If I, LL'>Cdin c]ccironic5> n'lililiolls per YC;;l,r}, "illcse,1l'C IJ~ o()lPCro~cdin, a I~oor ;'IlCLI~lm. alld n."qLlit'C' no

rzr

..:r~

958

16 FEBRUARY 2007

VOL 315

SClIIENIOE

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REVIEW
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dom [dfJi=2 •.N= t24, :nJldP~{D.{lUOII, ~·leIc.~ \ !'ilm'l1.' .hat d1.eM::~:l1itiJJl:cd;)!Qr 1l,':-1fl'I»1S~ a~ "~Q>o elarod wiril ~ts L1ml 'G!llll~ rn.. "'agJred. b), el.\:Illg~ iu ropmorJ~lclh~ pll~'SiL1!L1g~ dOI'IlQgr.ljlh:r'. and b1 d:H;l G'lr E. c~k '1'l1("l1tlul;uiOl':.1mL~ ln tl'K! ~lIn" m~'r Il\Il oo;:upy duoc:~tc' wi~lwr t<L11~ withrcl:!" ~i\.'cly Ih~II.' !]lDV!,'l1J1C!lt I:.iJL"V(,'icnl fii!OPlid~lionl1 (ill E'lk"\'\IlQlf I'iillim \'my ~l.tIb.\ll~m iany ,~I'OS." willt(:r 1:aJ1~~ (fig. ~). Elk~wfl:l r raJi .... <JI'e g.cllOIOllmy _" It;r...... ill the- 1;:~l~: of ~l\: ~L1S}fti~C'lll and hi.£lh .ill it~ed!l~. to imJlIC'i,b!~ d!C'ICiJ1Sl..$iI1ip~.rn1Jb; ~ml.~qu<l~ic iUIY~u, whcrew~jlr'''llIlhrlg il:; ~;mI1Hltlll mIt! wollfoo.."tIINUJ. Cy i..-;, ~1l{5~. "Ve [~'>:li.:li.'iCt1df,: r(,lil:t.l(JllltU",~pll)li>o h:ol'iii!!:. , N(lJ'iC!~td!.:;si>. 1lIiil!~"lj~Sof wol f;.elk dys ritUnte~ h~L\l'C ikllsctJ {)]11 tIIht"iC1 predMiO:I1!, i~loriug i:liI!JW~ liSHrl~ CIv.)iu'C-a~,'k.oo i]"I'Ull~~()~"II~ OOitOi:'J'~~I" p:!~uliall dl'cc'l$ of rultipm;ilil.luIT bdwlvDor m~ lIt®)lY (ELISA), [t) I11Il~5'lj1Cp~'{~t'-rolt: Wi.liuD]:> I.ilr 1~5 fi.-cu]'~UtpalS frtu'l pcpil.~Lli(~1..'" 0 0 dYUlIIli.lics (l- 4l This i~ lillqJm[llg,b1::CllL!S\! the >cree •. .eJf dliUlgtS ill d[i; OO1tovior .011 plam ooun, li,,'C: wirllC;f IrimyCi. 2.002 200:) WI~, A(1U:ff>fJ!1f"" luilfl1JOl't> ,[1JJfId ycm, nl:<lll r~~j~iro§'(:;Sl'Crtm(; 001' Immitirs 1I~::i,:OOt;U1 diK'lIl'!::i(ld {;.'!;L1C1J5ivcl,yI ~.,gl" (5'1J, Elk l]!(llm~ 1i\.;illl;mU$to Lh";~~I~u:I'w:ohres i,Qr l.1",rlu:~LliUl'I.'i' {;Uli1'l:Jm.",-d! ;o;I)rullgly wi,ih ".:IIk-wolf r.nLtt:;; Low fMo,g~";Si..::.t<m..: wlhJc~~rc !l,.;:,;.oci~ilf;.-tI, willa ~Ill ~I, Sj:l!LlilliJI SOi~ of )lC"v(:ml ki 100lJ1.:it1'5 miitl :II tUI11~ ~'II;; {llf 1Il1UI1l[l(.";S. to d';I~ j {H'i'}.\'Vol\''!..~l'i w~ l\'=ill)O I1du'I'i .... prtd~~Dipn':§UR: liu F~I:A.IKljllii!l'll 00(, lRXl~I(:Ioo;::i ttl II,..: Gruitcr'l{c~lIuw;SOCltl": U'llJl',~LCilfiI It.~~[iol.l ~1 = 0.54, Fil, 1'1= ~;A~urltl p= 0:001 I], (r;!; TIl": .n.icru:iniUlCm. OrC~~'i.~""Sdcdim:d :;Ignij.icruHly ,[GYE, Ua~(.-dSin!.,,' i~ 1005, mIB ~tJ9(i. tbllmvloo;::i hYlapid gIrn"'~~ al1d g~I~I!lic CXflln~ion or Ihe iin Iwn ofuhc:u:: polflllbnicm~ ~iJlcc'local n;loolrmi••;J;.. \V(DJf popl.ll;HNm ,md a GlJIlct1lTetlt dedine ill loca~ [[;mlDy wl'lh!~ j 4. <'fJ, ,md 1]1ng,~ClliOnC'Cl"mlCcr1" ~r<n:im!!:;, w~r.C' wr!~1ntcd ,..'l~h ~alU·recru !!!llC]U illl 0(;1 nUII~t1>l;:rs (9),. ru.~h~ C1allmiDl C:m~~(m k P'O!1Iim~ .o!·IlI1.C GY~ {/(J~ 1m.">(~:il1!i1. ns.ktl'ml'l \;\'i"ilil.'~ Wlril;'~~ !!Il~:if\L~L\1~~I~. }'~ (~I1!11} !!3:.,. j'2~ 0,32:.1'"1 J ~ :s;Ilo1tblly and tcrnf'QI"~~>' alld clk ~SflPfld !p .he (1,00. iUld P OJB2). 11~C' IfI\\,~ (IhSCi~ 1)!'fI:li,. :g~lcro:nc CiL'l!~Ci~'lm',lIiGl'l ;Ji;'\a:;socialca with the' ~\ P"'-':;(';!lCC' Qr wnh'C~ b>' <I1tc!i!lg~~ttcl1il;~ a~~ f"lif Illl~"'I;!i<1 c<II.lf-cpi~\ rmi I,') (1): C<lk~ '1)Oj· I Uti ()!;)\~). g.alijPft tmbil.'"U ~JootiO'!l \~i~i.I~~ltX. r'Ol",g~lg, and whidl was !'he ~f.I<W~,t Df.20 ,calf.c(lwrn!ios mca~1!11.ij..,j •.y t(l C!1I\'i.I'1ImlOl~1 1 cof!dhirmr.: ((j .'rJ!1. For .. <lU of these l.1lClmvi.or:;;"fem,.Ic-ll IflmdlllC'e ~igllm.~ sUlcd ror DhiU site O\'cr a period of 57 )ioCm"S. nC' h~ght'"5I. progcSM"QI1.e ,c(lllo::mrmiOl~, ,,'ere ~sso(:;111111ySIm'llgC: .uujprod~~or l1."'SIl!i)!l$'CS ~l<ll~ mal~s elated, w~lhCi)~ I:cow r.ltios ::llli)','C' 30 UJ'il~k.iil of {f, ~}. mld ~C!!n:I~,.rall!~!tyto \"'·olves.le.~?~n l~J.!l !l'Wtes [ifl 11g, S!, l 1.94,1oi, d~ 01 .ITI.,,¢- :g",\Vi!1gr<!T~I!JmiQllS) ;tr:.1(1 ''!,'jlb ~ilT!<Ji!C<! I~reg.dU:ibl~ d~~lm~c. s {Ch~llgCii i.n.IllOJil)holo~g~ r OOh~"lO, m n:sp{lll~ to I~L'C'daU(mN~rlm ry} reduce :l!l lmI(!i\fi:4L~"'~·S.V~lblL'1r[1bi,i~!y !e} pl1L'(I~~ inn lmt Cnll)' Coll!:': tlmtfie~ec! a.~3i,n~~ dILl'i]' (ll(I'l!L\'&'>i.on \VhCill pmdat~J:l11.~-iilk L.. Eow_ Fol' Illa~lh ,lI1d bw~~~u<lt{lS, tile ~'"l Df hldlJ{:ih!l(.' dclC!11'iCo:;, .h'LII,·~ i!!llQIOl1<illll CIU.'C;,lS OJ] ool1ogl;lpby and omnnl~lllily ~lllC'tUI~ (.l). Elk: (like I1n~. \renebmtC") .alIJ.!'f ~hdr 1x'Il;lVi:or i!!l roslfl(l~~seto prOOa1im~ risk, <I nd !l1>C-~c ant~flrcdmflr ~mlil~:;; ,~I~ :u~.d(lgO"L~
.

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!l1;ltdy l'~}.,L."lL'I1t1)!. d"111'lfll1ll!l :rndinl,lggct~ dk mlV~illI~ Ihl;l G<JiiJa!ill CallYOIl illloll !\~m\Hl'dtl~ U'llIlC Vt.rem killed by WO~\f(,'S U!! Ih6ir Jirnt $~Jn.ni;lJ: a!'Ii:I 1iI:1~ v\r 3{D cah,.e.. ... t~nl.'lIUl~i1ic:s~ A ~lfII.,IJ}r:\',,[11U)re
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di.:ICCK'tIi Vt!i)' 1'Ji)~\,' mIl.:" Or direcl. ('0'01:1' PA.'tJilliOJ'i 0111enlves b::lore ~RT"1)H'~,imcr (ll'I1f:~O"',' ,~~til~,m~. lliK: OC"1Il!i:fil illlIJpretl.morb::h:tl'lfior ic> ~ deceased risk of predilliUJiI, and tl~i.s is inDCOrPC!J.llln] ULU[o.llil1u~ny imo measures O(f Ihe diJfC"a. raie of prot'iruIOIi. III OOnnlu5l.rIlOSl alli~y$l.;~,Oj,; vcm:lIlll~LC lJfl;thl~or~'Y '~yuamLt';S tfu .11(;( it\."{'t.ll.ljl\ IIx Ihi: QU~;:; of~mipl't)ilim;~r bkil~uvtJr. \Vilhmn (l(JI1Sirl ..-ruiiLlIl of Ui~ nlltlirett ,c:fR~ orplt"l1.h~jOlj, il. L'i ~lkd.~i mhs~tHc~ !l;'P~jcu 'wL"!I,I!dllCl'Ili~~ !brl')!)l1nlM~} ~ll1il~trm h}' 1U"'0I;I~ 0iJ1U" da1." ~ltJIW ~)"Ll !fhe !!1"."" pm:.~UIri;tii\1"(N:O.SS!i o['~Il~pi.L'd'[rn' 1:(Ih:OI\'iGrc;lI~li lrug('", be wid~ iIl1!!ML;'11It C011~L\11.Je~ f(lr 1f'IJ:CY dY!l~1!ll ~ i

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16 FEBRUARY 2007

VOL 315

SClIIENIOE

\IV"IMrIV.sdencemag.orgl

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Pattern Sep'8J8t:iionl in the IDe'nlt;a,li8 Gy,ruls and CA3 of 'I hie! Hippncampus
'l'hl'l~~ti'i\I~ HilC'd~h hl~"'e IlI;Jfl!!l!,!olr]lwd to th~ d~ll~te gl)'iFUiS.;IS iJ ~l)Is<'Sibl~ ~l)IUU;!: iji! !IIlllLm!l~l pi;lU~FIl ~~p~IF.jIligln" ~!II ~!Jr'~~li[lelllt wit~ pre~~~tijQn~IlFijiF!ll these m~~'i;lts.,~¥~~@>v~~t Hili~~F!II~ll;h~'!IIgie~ill1J lh~ ShOlj:)!l' @ff t!he e llViJI;) I!1nneot:in ~vh~d!~ts ,)I F'~ c:l:ptofili!'9 '.,)1 nsu bstal1!t~<lM.-yail:cr c~nrcl.jlted .jI.d~!;!jt}1 ~an'@ms a!iJ!O!lAJIpla(ii!-ml;)dl)l~ijl@d glFo1lfiUl@ (@Ils in the d~lntci'!t@: IlYrlllS" Wn~!Il thll! Cllll,1'ironm@llt:s ,an:! m~u:le '"0 Fe difJ'@fl!nll. !1~~'Icell 1)(lIPuII(l!tiDIilSare JlbClFlIi~~d iII (A3 bul rloili!ll(h@ d'i:!t:!l:oJt§!llyrlllS. ]heSA! l r~stlI UsilfT1;ply ;11 dlJla~ m@!!:~,)I!IIiiSml~l)r I~UCH~ S;1liPglro1l~n()ni!l whidh si~IIi!.jIls h(lm tlile '!!1!t!!Jrlhrii!i!~11 (@rlex 'C~1l bl;! d@((Inl?~~l:@dI b!!Jth bly c~ Flg@!i lin (oirlddl;!nc@ pat~lffil')silil~he d@l'It~11:@ '!J~H!iS tll'ld b~ re<m.litm!m~ mo OO\!'E1rIaPlJiln~ ,eeM a~embHe5. hil CA3.

OOlm~1CDiOJlISbC~VIICeiil nllC ,g.r.n'luJc· cells roo rou'li~b] cd~$ in 0\3, tJ 5), H)C :'iC~" rre~mioJi, oriiucOJ'Ul ir.g iUlllL~lS rrr,m~l !he oollle..... ,

lfli:ighl bo;; illClfIilll:(J 11~ Ihc in,]lI!lIS CIHcr I~IC CAj .1OC1~ mk. l"lrl~ pmpo~,d ~'Dk orl!m ~knn[1Jtegyrus ..

in ~LlI~I]1 ~cpamtL'm C[II11: be w..IOO C.1{,IXidlirl.t·nrally b}' nliC'Il!;Uf.ing dlC :ahility Ordcnl~lh]: neurons to di:sal1lili!,tu;IIC sinall d i !terences ill c.mlk~_~ input im1u::ms_ In both 11:112: dCl'Itmc MYI'IIISm)(~ .hc ell. aI.'C1L'l. II!1w prille il.1Ja1cC'~ls haw ~p.alilal ree~nii"'l;~icld!>, i.c., I.ilcy di:ICh,1rgConJ~' wh~] d~c :1II~i Ilmil ls l n Imnk~~ar places U J, .J 6, 17,_ ]~~11m
CA. .;:Il'Ii;'a:;;. l>.lltt~rn ~~p~['llflll;)D1 is t~:XFr>csi<'Cd In

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tliK:: nc:1\"'L1rk exceed a. ocrt:al:ll di'ITcn:: occ~hr~:;;h·

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~r

or

in ~1m iu[pLl'l. ,Dn;:;l.::!Ld, )i,iI'~1dllcilc~fU1ig~-;';. Le~I, n.:~IfiCv1111'iJf :;1i'11 ihn" r~alU>Cfiill. pM>1:~n S~:!<u":llti:OIl proOCl!:tl where!)), ~"oi:li,. that h~L~cal reMy 1iJ,,"i..'II'1 Slll.red ill til,: n:C'lWtu:k ('j:ll,llUcrn ~'Ult~pltliun,} V 3.;5 VI. 'fbI: f:oilmu iol1 C[~~~np!Jls mr.: deC(;lm::ltll!lil :I:; ~JLey '\.n!(.., IlL\: ''-'lirIJ:y of n(lVI:.~ disercLlC ror:rcsl.;lls.ti.U1~$W'OlJld b(;lldi~ sUl.g;esu:f ute II ~ppocruJlPU~ (J .", The llCultJilmi ::;.~I'ongly liitn1f1 the existence of IUIndl:]il irniill un~dl~n 1511:1$ of hi ppoo~liII~RlI pauern sq}f~i:l'I,tiou II.flve ocnifli~too c,lu.~i~e. l1ol.'l{Cvcr. :ftUI priru;ipie., li,IU'llITo.rmlinyc[. llp~t~'l:ll1nnf CAJ. where Mmlll o di 1l";;l1,;ucc;:; bo;:~V"~11 irwmnilll; ~igntll::;. ~mJl~cl. b..: llICW n;'PlU\::>I:;n~lni:oi[l:; ~o!lllkl .~ d i~!ia~!l.1li i::;licd illJ:giI'rilCIlICdbdoJ!.': [he ~Ilf)ui is r!\,;.~,"1ii't..\:1 io i.h~ 'nm~ly 35 u n..":>\II~I. ~1jwK:~or JYilN!InlI"'S ilTl th~ {If !i:>st)eia~ivc CAJ n~1.\vmk ~! 5 7). lnit 'lie-. -3. li:;50C:~1~il"''':lldwork. lhe Ikli'l<Il.t ~ynlj; h:tl:>lool,'1)j I cause C ..\] l!oCTb. :m;: !;I.OOI'~Iy 'l::O:n.lro~ h)" mh(!il' prulll!llioCcl '[0 serve sHell a r!Jnlc~iDJ11 led ll-.ln. On lhc CAJ eel fl:..; v i~1 recurrent OOi~3h:T"d~S. CAJ ~Lrc~1 basis ufanulogit.-:> wilh~h'1! ecrebc]lum {1'O-l .2), the ill has been Siugg.C'S'lcdtll~u. :in the demaic gyll!l!>. 011 Ui1t m.ml nll,IY have ~imirad ahi ~il.ylin, :g~I1:Crntc input liFnm~hc 'CillltL1Ihil!)~1 oorrlex i~ dispcntoo lII'I;'?r.' uCproS()I1!;;l.! iOn! s in n::~.pOI1"~ '10 weak (IIl!O a l'IlOJ)C 'C'JL;iellSive In;.vcr of' ~mr'SlcJ)' .r.~!~.!il :gr:m\d~ ~C'~~~,'iJ'!1>'!h!,h~.fl:cadl g!'[1.iHI~C ~I ~1:0 ~g1Ji GenU.: for die 8i00l0g)! Il~ Mem.~. M~!\'legili!~Umwer~~ii'of (l1)~y .. !lull:!ill <Itld d:is~i!1.cb ~mc'lion or th",!O~,~1 S(JMC~ ami!·fld,nelogr. 7";119lI~oilldliieir!1. ~orwd1, oro ~hoilfll ((!J~e~1I0!ildMQe'~~d lJ.e ~d!!Ire~d, ~·!fIl~i~ input t t). Uy 'way (l r the ~,~~~c level of fhil!:l, ill,

hc ..funn~i.i ..()trl Of.di~~L~tC .. i,rl tti~llirny h Ilhou~I:I[ I,ti

n:.'

.prr'''''S~TI. h(t~~]I]:;
dCpt..'ii.dC}ri

daill~e.s

11

11~~Ii~, 1\..."'>'l1~1 I.h~ i1l1

(lill U 8 .!5,. To csml1Hsih whc~he~ fl SIIl'lil!llt ab.m~y m di~l.iulgqlii;$il o,...crtf~ppdllig gllputpdno;;m::o i.~ I.");.prc.~..:d in i~l~ dcnmh: gyrus,we oonl1l)~IFI.-d uhc 1'[,dli~~lbitTi'iiI'[imi or n:,p~5Ctr'~miOli~ iii th!.:
dem.~le

gyrus.

1l'I1~ C.'\3

SCnJ;(My inp!J~ puul;!nls~n;umr(],.: irli.ott dilli:all:iII.

u m!cr ooli.dit ions ,:vhL'1TC progre~ii\fdy

gJ''[Ijfil~rI'~.~~.~~1'l>OFJ';1I[ll4.!cm VJ.

'SL\ iA'LIS were ll1lillCd [(I 1'il~ll ~rl sl]mm.: 011" CJreUlill" cndO'im'C· s, DUll Ilg nm lTIiaig.,~ri \'j;e:n;; !i\;;:> :>lmlilJkd (n.1U1 1)Lm~ti'''C ~}}'r;lIInidlill ,x:IIl:;. ~UICA3 (1'1 = 11M ;md l;:~lmlak cd~$ un dl!.: ru.;nlUlJ.: srru:;; ,(u = (7) Wi~. I, A mu:Ji B. 1md IiI!!. 'S~ illiid labk SI} (1'5). In Ih'e or d~{'!Oluim als, d<u~,1w~'C flJ,,"COJ(lL'dl '~imlJI"11100~liSJy f~orn the- !'Wo nreas, ·n!C nnmlii:!Cil' ofac~ij\l\C' cells was 'InwICfi' n~1the (felllale gyl'u~ !J~,ln ill. CA3 as)" 'W'h:kh Cflnfiirn1,~~ p~C'\riOlIS lnl(~i,Cali,'m5 of s,p;u~ nrill£;l In ~he

Um~c lfIeU~l1!lt1. tlml wc-m ,u;!ive. !he f~eak t!rifi~


m~.c~WOl~ similar 0.95. d f
il~

14"

b~~

~ll1~1.!ll

!!!~rdJmOS!!lr@nlnu<n!!l<

~hC' gr<1IlluJ,Cl cell

llOalll~m~fJ,1l

(H. 141 and !he

1!'Cl;! iflJ~~ ~l

d~~ h'!l'(!! h~PtH'i!i;<Inl~'a~ 94. not .~i!!.!lifk.n~it~~-s.J ,

Fi,g. 1. Ij)la~e-'\ipE1(ijfic: iirin(Jirn (A) '!.he ,A. Pl"a midal [a!fE! r or 00. .lind ~IB)the .gra:n1JleI~e( ,cd the dellltate 'g~fI!!~" ,~Al' .a ITId ~13) we~e fe(!oiidM flmLlL~:an.eousl~' (rat lLl2 JlS) (SIle rigl. Sl fill' addit10nal '~lMtrl,";f€ po~i.ti@rn~ llrotIl! tli1i> lInil'llP IIJ. 1~e<ordhl(J loc.a'liom (anow)idenrtn· ~ied by r(JIIO'~!1~ th~ e:ledJode tracts
~L@m

'lhrQU!lihseriJal (I\e$'"VIvtQLel~ti3iiFloo~e(· ·~i()niS-DG"d~·liIla~ gyrl)l~. (Mkld~~) hl5tElr .. C dia'(Jramll shcwtinl~ s-epariltiicm ,cd ,extra' ,c~Uu'O!r ,00ctiOfi j)Oi~:enM"b (SlJi~5~ when '~he amimal. ~CI~agediliia S{jiJa Fe 1000~'()r 'l'-rp~als mo~'1 elllEl;r~l1!s~MIlmof :sClual~e~ or \""O!I~ef,o!1f!'!l) 't!\l(l' fIK.ord~!l.g clli<!!iI!ilel'5. on Each ~M represents one '5allilpIerl5pike, Di;1itll'!ct~j;lilk~ dl[l~~~ (~~h oorr~SPQJi!!iI'ii19 ttl 'DI1E! pu~t~ ~E!U) are ~ssign~iIiI1 0 l!l!ll(j;ue ceklr that. is consiistellt. a~rO~li Ilal~ OiF (ki~tIf r diagrams. II1~l'S ~h®w 'ltJ.e :S:p'i.~~('F@JlIl t~~ rilid d!u:st~r Slu,poe,riHilp@sed

~IiVV~;~t'"'\l\f1,/'~f\r\(~
(lJ!irrNL__. M"rr!I,

EI,

JiO l[l1iriill'or

",,'helil

the ,~nllili(ll sl~pl.Sc~-

I - 'YI~~~
'-:-::-:---:-MIll ~

ANJW~.AVJ~
.

'lJilh'fIV!_.

Oillhe rat'UraieiJt():ry {9J"~')' dl!U~lig~oo~mOllva~ed rumrilflg if! a s.ql!l,ue endloo.ure.


dl!!fir!g~@r.a!l~!l gin the b(i!~.

~'IFligmt) waver(J:rm~ of ~'pl'kies rlrom tl'le'lreil!l (J.lJsters (llIi~lllS Htandlarilll dle!lia~liOi1).IBCltlum of eadl pam,!:!I,1~:p.re~liIililtl'lN:!· H.G 'trace' fm.m the ilil!lil~ca.ted rll!coJdiITIgsile

961

RES EA.RC HART! CLE


i(13b!c Sl )~" The or.i.my of 8CILI,\'C'cells cX~lllvhed slrongi}' locaii;!)C'{1flriJ'lIg in ]jmilareas. or III(: 1 I acuve CA3 oC'Ii:;;" 62: (8],3%) httd ~ocfliized finlllg: 5& cells hm:i a single ji!Cldi!lnd (IJ\~Y4 hoo l'ilQl:C 1~1l1ll liri~~g ilcld 'lfi~ 1.i\~ OI'U"IC QltC 44ac~i,\'o[) domruo eclls, 4,1 hm! ,firing liclds. The ,~l1rmJorilyof 'these! OIJaa~fir:r,d ,m! ,mlllltip:l~ l'l~m:t';S, l](J 11:td~blga(' ,~idEi5"13 dOl!lbh, HoJtk !liMia)l(' liddlol. and ,4 q~;tIdnlpk Iit1l.ds:~r~g" 1B arne! fig. S2)1 {H, J'4}. The di:-;~:rihLU:io(lD~lhcmuilliplc or liI~ III!! rl~rm W['l~, gJJllCfiillly i~ruMulaf (fig. I B Jc f'aUellil ~ep!illr<ll~iOfill~1 dlh\ilfil:ges ]1'11 spaUal, <lInd h~llI1lpor~l OO~fu;irdeililil:e", We OClll1P'lI'Cd. d1C' ae!iivity oK ~inHdlmWOlLdy rocmded pri:rlCipa~ .~leLmln~ frf:il~ll diCillta1.C! )'nJ!; and CJ\3 dlJl ~ilng g Fi'!J. ,2:. ella n.'{l!l'£ ~11 rrale CAJ and dentate gyrus ,dru rin ~
rtlisrnDuitJotilf'i1 1!IF:lJ'!Jr~i~!Jfa

mm

l~tC ~ccon:li~ ~i[ll.t trninoo irrl $q1l8~C umi oei,rulylar vcrsicas ;3 recorlHI'I!,ll enelosurc whh ~~ ibte walls, with lil,c .~W"O C',~ ",~,::;,Ll)n$~r:,.:':1e~j~cd ~m:IOIn order (26) (fl.i,:. ini :5:3). TIle OO~ 'W&.. then lrnrn~.[Qm,1c,d.lhmllflh a ~.a'ics ,of rU~'0 il1~(!IImoo lah.:! ~htl""''':ii. from Ih~

Iluc('Crucmld t~lIf;farnlm~l,on
.IR]jIS

c~wiffilmtO!ilt fivc

or

active in~he inllif.hlslmpe. OnlC'1'S inti],", mkldle

WoC[(;

or

fins IIS~Iapt.""S~fill. :1, 13 ana C}, ]l:ICSC di n·eWI.'lJ1tifL~

Oft

~qUI~[t· :;:!1~1p[l1.0 l!he circulal: dmpc {,~ = 57

(£t'iIl.tatC'~lLd170 C A3 cells) or vice versa {fl = M <lim! 1~6 c .1\.3 cells) 't fig. :2 A: !fig. 504 ;lI:Id S5t Ihal~ITI.~s:iimn",~Im~\'~~ aCicmIlJJ~:J]ued by a \ Ilm~~i\~ change in the .~bingnH'C" distrJ111,UIKm
(~0Jl1"1C'

oj' or;Us: ln L16lh CAj:l'in~ d~ltt1il.rq g)'n:J~.!~1 !:h!1 (~C'll.tatC' gy~'1!I~,Ihe m~lIn.iIi1~rc~il'i:l1g lidd~ of eaeh cdl dll~!l~cd .~lntiIQClilc!Uml)~. SOl11CWCI-e Ilfms.U.

ehm'lgl,"'l\ resulte!! lin a $~rol~ (LCCQtI'C ![UiOJ'i[lf .he .mlc (HSu1htl.ao:rl of;im!i~'u(lulli cells berwcen C¥Cll'1 ~li'tcclosest ::.;ilalJ!i.$ of !'hi,; I:notphooii, Tho; IIUe distrihllllinil dml1!1,oo in(kpm~clcnl~y of'al1;[l OI<J~r onh~ box ~Iml)(\'i.[Hill-S6 ~ un liko [hit' Ilring rates , in CAl. which {l:l>nil£'liwdhy:.;[OR ....is trig, 2B~Jld 1'1,~"S6,} tJrl,l, To c:xnrninl()how~h~ dcoom:fuJt~cm of dis· charge [I),aw'\rnl~'.unlhc demaie gyli!.ls might in~hlence (,A3, we i:':;;~im~,tcd Ihe CULnu!~!ti'\lC ~Imngc in Mll~!.~I:ifm .pLlItp~t tFllm the \lenlale grON as i,n~rem'i:i dOlg~~s ng di ~irl1i larill,y W01'C intl'\llduce(~ bCW"'CC~ll lJ1aiL'& of tC:01t boxes,

or

.It.

tion

,el'lVfrooment.

,~r the

ret:ord~ nl~

niS~~I'IIIl'a"

W' >Cells

B
cell !C'1:

~lw~'inr S@l,I'!Nl sh;:lpe~ 'm an eroctnll1JTe ~'iilh 1I.exij· Dr~ \lIa'lls, 'SIMirHJI·~lftillti .iii

\\!efe- ftc®~cled (DIi'i~e(iJI·

:square til) alld i1llr:lilllg \l.Iilh .;;J cird@ ()J 'l;ic@ Vllr$ll. The in,itiat.sha,p.!l

en

W!3i5

:5inu.ltiameom~y r00ord'ed
,(~U.in (~QU"Q!I1.

(18:) lPili'ling' '~~~~s. a 'fCilf re'JPf~tatiy!? 00 I:d (~op. purple bo~ amd a

lfinaillll' repe,rted

(1').

'IEm,
CflJl d:~';

:5lPi,ke- lo(atio:nli:; ~righO {(ld;ed rate


rCorlr~'.5,pOIf'i,8I'~ng ector!1fI<'! ps, the ~ ~il!i!~tI to r,ed (pea~ ra,l,'e), Willtll pi~ ts riot
,color 5.cate'i~ (rom b1Ju.e

U1e del1ta't.e gyn.is glH~el1 b!l;)!liL (ILeft1 Tl'aj;e(to,rie$witlil

vi:silerlsh(IW1l

nn iO'thrtte.

The roi!~ !i1@P:; w~re '!1caled to ~,,~ rma:dIiTlIlUn lillillilg r~t,~, wWlinJ ttil€

,entirel@slll11g S@Cllll@n~. P!ll{llk ra~s are ijndi~ted


to, ~lIe ri;glht The (omp~~~~ ,!)'f adiveooti.sus 'silmwnl'l ~. 54. (C) 1Ili(· rere!'Ilii~l rare (1-i'1!!1ge)in ·ii'lt:lii/Lttuai r.rirt9i~~el~h()f

lhe ~!!ll in '~19'ilndl'l'llg .ad~ilfionO!il r'l!preflentalm I,.e~h frOOll th~ d'elill:a!:e gym~ {hm diJffiifillt an1· ~ lsl'. In@ivi~~~II. ~!I;IWS
l[jf~dn (eH al1!! ~utmnf!d
ill

di(f;erenl

~Ilrt iliOO ~5t·

I.Iii roiJI~1l0liit the !lJIQ:f1p:hs~(J~e!"l~e. (,RilJhO Li~ dia!Jlr~ m:s rOlF Ihe' mean Ilring r.lle (rled (JU r'!l\!i!)' in e<ll:h iru;l~vidl!al!irilllg~ieiW (box ooHines: hi!!ofe the same ,cDlarasW1e cDrres,poooUl'L9firi!li~[ijeldslr. Tile ratas wer,e' liUed 'Ut ~~lFfltoltl, lftn:ear,. or ql.ladra,lirrUIl(tiOi!l~. (HiVe r'ut!; a.1fE!,oililly sih.vWrI ~"i\lten signilic.al1lt {~f'I I:lladk~,Th e' I:le.n~itr;/a'l: IltFbBa rFm 30 fie'l.dll:, sigrmoitlali. ror 21. [].e~tI§,a:nd1
,eiin~ (l)!i.ors

'Q"u,d~C!~klOT ;8 fields

(n=

122). 16 FEBRUARY 2007 VOL 315


SClIIENIOE

96,2

\I'V'IM;\!''sdencemag.orgl

!RESEARCH ARTICLE
bccr~l11Ic n~(ln: di ffcii'';u[, the c{lrrC'~muons rr;]!IC of,winciiJcm liring wI~J1,ial time wiuidl:!Ws of {1~fL..)SC(l in OOlh hi~~pocimll)<jl sUib:£klds. b~l[ 30 ms, l50 ms, and 30(1 am ~15), ]l]lC COJnl· the decrease was IIlI.•iolly more pronmJllccd 111, cklemx: rn~~.a\l C'riLg,ctl cro$.'> ecll PfLil3. W~ a thedcnenc gyrus W~,g"3M, The .adVfIl\C~ of!JiJe complll'OO lor successive Shil~S .he 11~0t1:.:l'l J,_,l,uLtl.: gyrus d~'><lPP:;ilg:edlQ~.'mtll~tc cud Qi:'lhe lle('I'IIi;,'lJi.CC by dh'idiiiug the rui(:s lJ..y each ~~~hCir. or rnorph ~1,'il::jI!lC'ilt~C!" [k.cauRc the imll!1.c'1 or d()'OC!T~ givill'.g a coactiv il:;' rnl:lQ" The ~cli"il'J'mu io OOC'I;(l,h'iOO aCl\l1!'S mo:.L'pillriat~ and, ill the mm~IIC '~hcflI11 .r.1Itei; o'"lhc ,Ierh'~~ d~~\\,\~':!fT!J OOITCiIT~1cl ng o rd:iliol~ plCOC~ i:111'hCcCcntM~gyru~ is ~ikdy to rl:!f eaeh _~)aIi:J~ bi~] s.lmno-d 'bJlhc lWO bQ[!( be cmmm.i ~!~d. y Ihe!' Jlilllliwd 00l1VCl'gIQlIOO O'Y b gyro~, drol'l1}cd :>lgl,l:i1r::J1Jllly even IX'L'''''('i(m mlI S.hllJM1!.~" opu l;IIlirm activity in t~1C dCIIlt~Jte!:!YI't:L~ gcm ule ceilis OIl: CA3nlJUl.mm~ U _'i, ] 7), we asked P jamul \l~1.'<~f1lGji; ~ Fig. sc !I!nJ Ii~.S~)_ Tim dlcr:r'ca~ hCI~1{(,CiIl sbapes ~ and 2: \\ra~~igal iJiicolIl! r(l'.rall~ w",,", highly ~c.I.'t'>ulivc IfD Mna~~ dlITll.g(5 .ilm the \\'hcthcr OJ ~~milm" diroill,c,LKm between mijaCCl.lt t;,ilm(Jes, \1,\Dlll~dbe 1:n~HIl!~ with S"P:H~~,I'inpl!Ils.. Il1rl~CC IhIll:: wJnd[)\\'S II :> 2:_14. P <; n_051, For ::.>hapc of lim CJlVimmmml (Fig_ 3J\l The ~nla~~~I.dilrc:rcllo~ gm'e <I !lflgll~Y _~igniflearnl The ch~alg~ III ,~patkl~ POlllli~mil';!n \'oclo~ :ill lila larger shillpc difre~ncc8" ~hc di:'t~~lnil~Jrhy th!,lls ~ki[lbl~ .Q I'tcr subsamdecrease ira 1I1e cor,n;lll\lt!orn. OIf Ihe 'poplJi~~~hm dJ~ltrrte gyrm;;w'as reached LitleIc~1 of a 8h~fl1e~ ~iSll.ibl,ninn ((I P1 V~I'c!f1il [or Ihe d~llmtc .!:!:t'nll'l({OS • O.mHo.,: pl illg r~'I,,)!I~ recorded t'lc.iII pt"lplilkilin'!.1" Snmll lhcFi.g_ 3fl r~1I p;!irs :iJIClG\vclC n1!'m~ c":'ICT'Ch*~d cifla~~g(;'J> in 00:>:' ~11l:1pt..1' "\L~ml~2 l' led w lIsit[!i~.(Ir (1 ::;:11<1:,11(1 vcr~u.~.2 and '0.37 _,_ 0.004· I'tlW ~h~lh~ 1 d!l'1I1 C'CII flairn \'\'ith one cell 1i'011l I.be deiliale l ""C'!'RI!I~ I': n~:tmll Cllm~lmio!1 ~ SEM, t .. ~5d > dmp in c-m'~lmk!JI iJ\'~I' wh~u Ihe cc~~~<tl1lpllC' gyf~:> and ene i'ronl {'A] (lug.. S~)_ P -< 0_(01)- A :li.iUlibl!r CCL~a~~I":b.'i.not Oblle<1Vi.ld d w,~",rt'duOC!d to ks:sth1lH 10 (f.~~, 1I1ld rig. Sil 313. Will UlI,lXI nsk,~1 \vh~1!hc'f tlr.12 chall1,!c .~u Ihe inl C r\J" whem the poplIlml'io!l \'ce~OT,c:oTld1ltifJn~ illl The J irr~l'CiIlliali()n nf the S:lmtbl ml.c: nKlp'" 'repllI."'gC'IlI;r!kul of'the crw iro.F1I1}C1II file demale lor n,c.i,gcllb!O.dn,g ~hf~X;S, am.! id/;lluical Sill<1PCS cm~~d be improved by dC"(lm:lm,~i(lu ill ~hre gy.f!llSbeiJltS any qUiliilmive "~il1ilihnil;Y to It:mil~ WC!lC8IbolU't Ihc i$iYl~C (Q.(J.:! .:: O,OO~ i~lltd 0,92 .l: pill:!:lIJroce$.';(:s in the (. A ~~c~d:;, IllphliCC ()ell:;ill ~~rllP'UI;.:q] dml~fl~1I (18"1. F Or each p<rur of Sinll'llllha· OJIOJ, Mpecl.ivcly. i = I.96..t11.~,), .A.s the ~fil.rpc.,; Itl"uLli>ly rwll'r(t~'tlCi.:'lkwe thU;i; d;.;i.cJIII]iiu,_'iIIi.h..: f.'A.1. lJud lOt OOlitl.: cxiem in. CAL n:mfIPpillg
Compofj;i~c PoOpllI!ll.llion vectors ~"ICJ'C con:;U\iC:Lc~ Cor cmd) box by st~ld;.il"tglhc GUC' .rl1~psef ai~ C~~)~~lino 8 lIiI~roC'-djl1ilCil$km~1r.)imri.\ willi the ~v.'Q ~:rul ia~ diilutcusl.'&J'ls dis~~lh:) .. ill(lli~, Ihe Ji' ed and .~'i1>;Ci$ tmd id,;rlli~y r>:pmcnned Olll~'II;:; axis !F~g. 3,A)• Fer c;mh prnir elllviml'lmlJllIs;.

~en

or

F1!lJ.l. Q~laiill1tatiV'e a~se'Wm@!lIt o'f '~~a~i:I~~ IIIdll!m:poo ral ,emm:ic:ienoe' redlwti®i'I in the' der1~:.!Ite 9~II',UiS, (.t!!.'" (T@p~ IProc-

A
~';.2~'
€ t1&II
M!llI (!Gi)

c
-

~~U

IlIQ,DCl pal ..

Irs

,eoou;re' for i:akula.tiIil9 pOIPu;~;}HO!i!!;I'~a!H

The

ral~S

of a l~(AS or d'ein;-

(io~n~~atio!1s"

!ate cells \\lere 5itt<! eked i!lJt@ 2:56 p(lpu!alllorn'e(~F5 (PV$), one low !l'olcho,~ t~~ 5, em by 5, 'Cimbin~ thll~.lI~nl r.'har,e~ \ bel~~\I2en the mo:rph SiClIUii!J,e a i'II:I tile lIIlo:rpili (lrde'. iil~ oJ;;@rrel.:!h!,!1 tielw(l!en t!i1e 1Xl~
ulalio:nl V&.lOrs Will'>(O;mpUlIE1~

Ill :' sJ
O'~~I~~~_~~~~ QO OOJI era II mI! llDHJD '1T!Qr;p" ~~rnparI~

~ :11i-=;!:-~+ ..
1

-.!-+.''''*"'~~~
!! 1IIi:1!ll
3(1

:1.4

41:l M

(or ,ear.::h pllir

or plili@ls,'

u)

~Bal~

:Ii! Cof Sllln~

mil51

tom) Meanl popul<il'i:oFl WCIllJ

os '~rl IJiLlIp'i!Jl from 0l!.3, popuren l!citi@1l vectm in 1lhe dertlat:e i!Jyrt!I~~h@'~ dl~i~i~~riti~~
alJ.e~ a~tbe f>matle:st dlange 'ortn~ D@.X !OO!I1rigl)llaUml(Shill"~ CAl IE
!

"~m'el..lti~.!'!ts, for Poll F:SQfilill" ,c~eil",;n:gly- dlve~g~liIt imcuph ,sih!a ~5. Il.Int,ke ttle ~:JlUIIa·,

ID'

! (lJ~ Ii!!

IO:r-__ 4\e~~
;.,.,
-

be'~l,i~e'll ~hid:l)mpiilrnsorl and ( r~tlNJ ~ir;(lrdinJJiS ~Il'sl1ap'11! 1.) (I) IroopuilaJliolllYe(~(lj' (jO~

] vers,us2)" (Gr<!.Jlslipp~ed line~ h~ghlijght th~ d~l\f·ef'em;e

CG-DCl, p.tlN

CAl

-C~pt.lr.

r.~lons· ",ft~r random su1r :5almp~in,g rrom If1lit ~e(erue~

<:eU. ~ mplt!"

Si!J~s!ili

I;Ir

I;C~~

coooo spa~1al (nl)SS"-cmrel:at10r11 matri(!e1: for mmjJOsi:teo rate maps, one f:o~ diUerefilce :shp~~ ~~~ ea.dll p~~r ~f llIl!;lrph ~h'alpoes,.lhll' ~~1.ei~ from ~!Nli (-!-];~ ~i;I bhu!!l' ';;;:(D~;,,~ll!;lThl ,cor~esponds to, 0.5. te 0.7. (I: ,and F) Line di,agrams :£hO!.~ff1l] crosso(DI'",rniatnons ,e,llPlre'.5~!!'d as, '~h.e ratio.. J'(!Or ea<:i1I (eM :pa]r,. of (ooclll!oie 5p1~e:liilil (me ,ell1l'ir'onl'lleru dividedl by c:oodi~e s,Iliked n Lille other, \\!itiliilll tiuee ~ime O!i'S,81 flmctio'lli o,~di:Yt~lnce freUD the cen~er of the!:ro~,~oflrele'(Jr;n'lllr(H pairs o~
(11allg ~inl

\\lere c.l1Oilelli 'ffr'olill 'e'ittler 1Ii@FIi"' lOlle 'gJrlW' o:r CA3. and ~Of~ re!laliml (Ioen~den ts we Ire' ,,~~t;~~~~~es ~!'1 {A), (QrneSpDlildil'l9 ~tal pDiml5. 'in (A) oil nd (Bh"Hiirdetllin red. {O

0-

~
disiaMil (em)

tem,po Farl.(l)fi~laItioo

®f

cen pairs as ,8 ~!lIii'I(tim'l

b~l",,~enml;lr~h sha r;tle:i. The dfoglr'i;l'e

or

Jl;l<ffc;h01llllge O\I~JQSS, Jm;u~h:

\\iiiftdim'll'l: iIil: IiIOll!1d. Tille ratio it!: illJ!er aged alfld lifilJ!1:3Jlytr,a iilSrOFliDe·d sucllllllat lIria Is "'~tlilifil~~a5.'il'lg dis5imi~alnlty ~iii !:lox silape I~on:ellifile [OIF eadll pair; ~I~), r<Ilndoml,y 5,[ill!d~Ie(j1 (:e~~ l!}ail) haveO!I ~i:!ti~ ()t,. Symool'j as ~!l{lit}, fD') (~hn' de!iIlilte gyru~; (f1, (A3.~,

or

963

RES EA.RC HART! CLE


occurs H.'>~ C[fl1l'l1"I;c'iCdmngc in borh i1fi,llg f:ic~d.~ tiQI1.~ mkc- pllacelL'lCaliyi,u Ihe dc·mmc li:yfiLiS or from :lffcretlt :rcg;klnas, IJllltC dClUl:tnJc: and filiillg ~ate.~ C"@JJolttillrerIK!p~}ing" l' Q[ ,IS a lire nt~'(jju~t~d sdccth'c CJlilllllgC i~1J firing roles. wilhin f~11x.00SCI :gyRIS co:rurl'tuu.cs actively to p.1JUly,n scprn'maO:l1. Q r iiring io.cf~tklns j ''mt(;: reifnf~ppI~llg"~ (.;.~4·",;: 9}. IllS$C1i:S~1 i\'.i.IY 10 :smaH .;;;~a n-I,g,'C's ~n IJ'lf~l, plm.cfll:l!, ::;,iIO~ Id: be $uiJ~t::!i.II\~JlIy laf,b'(:[ dian in the hI the prc.~la ::m.rK1y,Ure C Alncu[un::i ' exh ilbhed elllto[hin~l cortex. 'which modial.~ 1.I:tcIlrmjmit:, COJrl~~sfClIllI.rate rellWlpfIlng throllghrllllt Ihe morph Scqlll'llUCL:' (lo), wlklrc-{J~ CiJ~h rm~n~hc cl~J'I't~n~ or conical in~IiIL", ~L'lIJlfl,;, m'~i_ \i\k 1.C:;:[!.lil •.his g~ :rired :;II di Il:br~'11~ IClC:lIllirllb'<; in Iht! m!l)l11 l)mdleliol1. by IIHxnidlu;gfmill1 duuc cells III lim I)(;rto.ranta'ptal.h k"'ll~mLlil.ol!l area or 111m ~!i[JJlO> d.itft'HJIlI ~1!~HJeli or the!' box (F~g. 2H)- Vel, the
l,ul,I~1!ribLlliio.nl oflhc furil1!:,\ :I [lCi!ti~i()ll!l wa<; mmi" F.:'Ul'dOlill_ ln .al~ 'indi'\'~d~a.~ Ik-lds <I dentate

Iit'SpouS'C to corwiroull1Cilrnl ['CCJOutr~~~["noo:JI tilldk!'1t OOJ'tdiLkIlJS\.~~tcre hlPfl(lC.nmpo.l CAl ~mblics


ullKlctgo nI~tc'rrc~lliiOilin~ .(3:). PaUilHn se'l!lillrathHl by recruihl'lellll

'Of

f!lell'il

or

(,U1JJLII~

",hilC'~l1c shape of the ~'(')(:1~'!:Hngox b !.,\'·m~ im:lm'illd ill ttllli.1I ~telf:;; (Fig, 4 and Hg_ S9).

IlICI,m:m.,

HrIng iIlCre;I~Gd mld dcoenJa!'i,OO gmdu~Jmy

acmss

IIlII,Il~ip.le 1!rL1~~. smllt;lim~lhIDl;Jgn(lm d1C' iCn~irc I'Imlpn s~IUt'ncc tFig ..2q\. TOo~tilnillc the C'l(tc~n 10 "" hkll the- plnccC(ld~ W;I:': :N;l1'1~Il.cd. <l~ in Ihe C'A3- ce-~~:-l, CID_.... We' y.cOnc.l ~J!.(,,-d the CO~llc

Ihosc ·~I~ ~li.d grid·m:c ririn~ lields. ~"imil:!If to ~ho~ or pm I.lC~pa~oc:llls, in Ihe medlal ~nuoihilltIJl cortex (30. ill), The!' !'ipjh~.llmay h<Lw:: :ri!!in~t~ . o

,~:n<!i~s;eI!nj)I.lies. ,A.![llO~Jgli Ch~JlgC~ ~l1tbclC sl'klpc or Ihe ~lOlpll 'boy,: ~lrol1:gJ.y iDl~]IJI(~llood lh~ eoarl.ivily of a:lft'ady ~lr.li\lC'11~liIrnni'liL~lhc d~llI~IIC gyrtl.~. d'~r(l WOI£ no lq}hlOOIJ'IClIII. !]or .~tlc!- :l:C'liV(l sub1\e~,of IIlre 1)(J,lJLllmimtn CA~. MICh ITiIJ!ace. I IlIlem c-'.m be indtl,c,cd by [~ll1Ihcr'im::~e;;K\lllg 11m diffcn;:nce-<; lX:lwecn Ihe CIIl.Vimnll.lQllI&.l;uth~I'by 11I10villl.g'd.t-c animlll to ,~dilfcn,ml.l\lJom,ding room
(.?4,
j'j),
~1r

or

di~irni

·1~O'!iliti1:), n~<lfl':'lof Ih\l dellt:1!tc cells fm: C':IC~]p<lir r.Ue . .or~h<1lpe.~:inlh...;: mmphs;cqlICIlCe' by s.hirlin~ theSIfLCt.. from ene of'Lh~ shapes in ::;~m S,L:cps~~ong 'the x HJb'd axes. This LOC:;Y in (! Ifl~P o.r 'UCIIi: I~cd a..(;m~'C .sinn ~~w:ity lfhe pop~dm ['Oil \1'~"C~IJJi>, Iii ~t \lllrimJ~, ;:,Jiii f~ {jui~ :.19Ci'k,rekLtivc tu th~(l1h~r; A ceUl.ml!E>lIk W3i>0~Scrved f:or fIUPili'n. of ~~lUPC:l:l,

rrmn il1~.(iIc[,nr p!lIJ!Cililro;l a~r.Hl" of grid! CC'~~\l O.I!', l'iliU;l1i cl~~Jl~~,{. :-mllki~!t! ~rlunch,II~"C g~obnl n;Wll;~p· ph.gin Ih~ C,!U (NJ, would cause ne\\'~Jld I~CJ}lil~. btlll lC'l'l)1 m1~!Y.~rol11 loe,[!JiIlICIn~1I1'L1mS ,.c!~vaWd bygL~d cc!l~ wilih id~l!l~kal Ii!'i n~ irl{kpelld~Jlt Cl:lI~CIl~mbkl1~)[I be n.1C'!'ui~ooalso in I'oca~iom:. Tille III ~e eel Il1;, we~~ fl,~I'lJod :>iimlJ]~L1.· tlitc dL'n1a~e.!:!)II'lI:;:_ Ral~ were rcsl.i~d in boxes of

V M, 19,

b~' rIllllkillifl

the Cll1df.l!NlI!~rno:rc

13, _ih \~ n;"\kod ~lo'hC'lhcl"

,I'

Or

hut

:shfq]!i;slbC(jmt~more ·din!;i1Cm U~ig. 3, D to .FJ. w~ti;clil :>lJgg~~rod l~lal [lIe rme c1u::m1il)~I~nn \';1jj.'j. i a~~cn:d:iln(}f.)~~iiy,f~ .cImlng ran,(; uclfltl'lppilu.!l in CA.J (16) •. Even iilc :;;111!il~~~-:;l :;;Impo..:,'I.omgc ocausi.:d a db;; iC:J[';!';' iu tl'le O~11Jiml peak in the Ui.:iIll'[nUt: ~yIPl~ ~:;;cc ala.o Fi~. 1.11..). H~i)',C'"pF~~:i(jn of<l (,,·a,u:;livily .•h~oo Im~Jljcm
s.~~~ti~lU pmCX's,,';;' il] ~hC'<!'cu,-ue !!l}lro~
!;Ifu;;e,<;

111C petlk(ilL"Crtfl.'>Ci:J

progrc.,';.:;j,\J;':!,Y as tl'l;':

the'

qlle~iml .or Wll'Clh..:::r Ihe

lInclcu~}'ing

CQJJ1Jltllm·

11,J;.VLISly"...,il.b Qt:lb illl .be g;rnlwlc cell ,i,tli),CI:' liM} olhcr t~lrodci:'. N(mc ,of Ihe grid od~:>,.;,:.;,h~bil.Cd (J,..\iuc Wible: ch~alg:~......nl the tal . ()r til,.; k)Ci]iimi Qi~ i c firilll~ 1!.1Ii1, oOiJhl '0e11MrlOO ~i.:d~o Iii!.: incl\:l,lil!~Uf]1 trm'lS Ihni'lHl~(lIl ,of I,ill.:' roooRl in~ en",! ronlillCI:'i~ (Pi!. 4" B l.lm.i C.1IJJIU rig. 5'9 ~ \iVI~i.':refjl> 111,..: ITmglJiutdc or~hc CR;l~Cm(·lmio'J] OC"l,,~'c.cnp~jD, or CUIVrJOIll~licl:n~ dcereascd prog,:r.."Ssi\!c~y ill Ihc :silllluillflllC(HlliHy:rccfm:ood oonmi.':: nc:mous, L.hcco:rl-cllmi(ln c1L(llml, dlaJlg.\: i>y~I.,'lJlmti~lIy fOr 'U:ICgrid eells !Fig. "In 1Illc:Ji: ruooitlillgt' !.:,nnlirm fJnHct· ....~~Liill~~ rr(l~I'il $Ludi ... wilh 11i11l-~'iT' liUlDibJ;.T:;i of 'S, cntoirhiml~ g,!'id cell:,:. \~'~]ich :i:how Ih:ll! the' d1u~ihlltion a;f act~\I~l}' b,1OlW!,'I()!!1 1i.L1llg fI:cId.~ in clflrm't~hm~ IK\UlllJ'l~ dfl<'S 1101.dm.n:gc (lcrccl.1My in

di«~l!Cm sii'X,col:Or. m:Kl $i1ape in aill\,"1,'; ii~~'1f''''m d ct1Oll'i;; (Fi:g. 5, and fi@;. S I I) ,~m:lS ~~). .1\::; 1,;1\.flH-"C'i.l.:ld. 11ClB. :O;11Lt~:sJi:ici~lIy 1 ilidcpt.~ldJ.:hl~: st1 bpoPLliilliOJj~ ~"'!.TCi'OCru iLCd :i Ii H pnif ,tlf ~~ill~il~rll)' 1iIIUiJ\.'t1rooms lj3HlFi~ 5).\\"iI1l9 nilcl,h~":iI1ol\:l of'tll 114 'only 20 lIS.3%l'[lfrlile ~w:, .1\J cdls c ~11CSlXo.UlcI nUll. or Ll:Ic 23 fLc~ive '[AJ ceil::; ii'eCO~ s;n.lui:tiIIiIOOU:;ly \'I ilii tkllllm£ cells, {mil), J [ LlO".4)wen: i1<:li ~\; in bothrutmil:i, ~tH1Jt.;; S ~I. In 5il'tlk~l1,g e<mi,m:;,t,. 117 {lui, fJFlO active neurons in ~he dCllIlJllC ID"1Ii ..~ (8 ;).IY'A:. ~ were aeli vc ~llioo!J1'i mom 1 :llld .~iIn 2: Oi~llyd.ne ~(l'IIdllfmnl a dm'c!'(lll~ lIo1ma~! furW !i.txdusivdy in one or dm CIl\'iHllJlllllClll:S: ,Wig- )t ThJ:: l'o!::girm1l1 duITerencc
~h:tll.W(:ru:

eeu vc

iUI one

room were

~Lfu;o~e~i~cin

i:

I
I)

~ o.~
IU

~-1

1-1'

O~
0,2

~2~~~~~~~~

""_n~"~llfIl'!

Fig. ~.
2(jin;e.W

IJI na ttPJed

taye.rs. o~ (Mow denta~e g~~I.1$(ra~ U2 9~); 5Yirnbo~s,.illS de$cnl:u:~d 11'1Fig. 1. {B)' Fij~ II!) li~~· .~~ ~~~~~r in W .;:!lnt;!~1 tll~ ~;JlQ!!1 ~ilfl1l!.1~l~rJ~QI!.I~l"r~~l;lr{il!l'g ~~liJtI!~ '0011 (brm'iiill and g1~eer'lb®x, ,~~pe~liiil'e\y)uring ~rogll\e~5i~l1:U(lIn.iSJo'm1<!'tien (h:e! d Oi~ m01]lh bC!x. O'!Op) frajecltl:)ril's. wiU'I slil~e' loc.!3tion~; (baltll!l1l) ~o~or'"uded rate fill 'f~l maps fOlr the same fecoJ'dii1~ Sce~Oi'ls.. Regular gfild~el:ds rec()f,l1et1rmm the l\IediOf i'll1t pa.th dI~d nat ~;w;~ibi.t s~~te!Ji!~Iii'c: ~aJng~io fV~!l 9 Irate. (See fig. $9' Ill\!, d ~d(lltl rnl'~i e~ mples,) {() (ro:$S-(rnlli~la~Gl ['i b~h'\l~1iI rM~ IrrI;1~ lOr '!lni~(€lbin(1le l'.l!i!rrOHlI1t·lla,th termirlal;o:1'I zone, c!3!kll~ate<l.al) in Fig. 3., D to IF,.ami :slhiown for ili1ul'fIle I1lall:h"l1i~es ~n bGX 'slha:p.e. The ~in;e diagr<l!l1l StlOWS!!I1 'c:enlr,at Pe',ak, a's
,e~poec,tedif tliie-fiel.lliI~fI~iinalfl' ~t;3ib~, ndla secv:lllI! p€a~~ a 'il4!.5 ~o 50 'Cml (mm the ~eiillew, \Wi~(h15a (Vfiil:eque'FlLe or~1te ~ mita IT gJid spa(lrn{l (40 til 601cm~ o~ Uilil' reoo rded' g rikll (ellis:. pp" IlJ'l!rtor.a!'!t IIl<!th.

nne dirotriimtfien irn~rld' (ell~ilil~'he pederant-paJtlll te mll1l8tiom HiistQ.tog,y'lJ'ld e~en:IirQ[lhys;lQlQg~ II re(lJrdi~gilil th@ d~1iI d~~dritk lQr

16 FEBRUARY 2007

VOL 315

SClIIENIOE

\i'V\MiV'sdencemag.orgl

!RESEARCH ARTICLE
W·il!.') M~h]y SI,gUI ficam I = 12.0, P <: n.mH, Z :s~mull!/il.uoo~$lyl"C()orrlc{1eclls oillly I. The: d)fd:oc'b':>C IfIC~wa:> nillljnmi~tcdwith {I hn~l~cr riUC'1111I'Cshc {lid [0.2:5 liz; Z = ;2,0,3: ,p 0;:: 0.001 {Liable ~mj and with u I'm,.; ovc.[h:~r inCill~l!Irc Wh,Cb'C. (0;1' each cell, 'he Il.1lJ'1lln linn!,!, 1~1:I:1 the IeRlNLi:I.Ive ~~'o'IJlin!l illl wa~ di\~dcrll)y ~hc nlCmn I1IJ~ lUi ~hC'.nor.o-aeLiv~ :s.!!ltHI]g lJJ) It = 4.77. P.;; o,om (.lig, SOW)]. .M:ofOOoor. in two 11Ilim~]s Ih~a WGTt! W;;'irod in II 'thll:<1 :i.'omn !"\<3u:m.:)~hc Clld~lIIrewa~ sliI!:t<\lami.. ny Imgm~.11m!!. o f 12d~ltare cells tll:il.\w,m active il1l'OQlll!'1 :l and :2 \\'l:R,)'ac~ive;«J~o il!l.~oorll3 {lig, S 11), A l\i1toughlhc ael i\~ ~1.lliiC~ "vas ~11IQ'mlmy ·the S.1I1rllL" ill. ;llm Ih~"C mllrlliil, lhere wa;; no, :s,illlibrhy ~n ~1iQ' !~bl!tiv(! 1f:!C~tim~ of tilldr liriug lickk ~1'I'~)L)'(Ith'e !1}f Ihe ,'C~mi\'~ M~U~'lIirm or the l<;;l3cfl.!:tl. C"~iIJl~corJC~llion H!.1!p!1 tbr :-li!!n\l~,a= llI00lL"!y l'Ccoliled demme ~u;~umm; mall"l~ I and in l!O'l<Crl: IliJl. alia ~h(;:oollc~iv;i ~y or ceU palrs wlAS

Urliq~H~ IKli1IOii:.nIn !~ rate slllill~ rnll]lix~ of gl'llnuk: 111, cC'U:.1_S:p~flol(~ COIIIl-cct:IOIIt..; bL1l'~\'\:.·liH'IgrnnLlk; c~l:Ii:1 gyl'll~~ltd 'pYr.:lIHi i:dlal ''('Us .I,nCA] tim dt\nllUt~ ro'!II,l~ ,md CA3-. arc bOlt! ,1bkl, il1 in d!(!' d:Q'J11~~(' di Ifarent wa}'~. !.O cO]]lpftllcly oU1llll'lgom~IIzc he U5) m:i)' ;IHow the cli:;':lII:1h.igu~li()n of IIr.i]IJ!l L oo.llkt:~h'~ 11r1:1lifl patlcm of eel I asscmhl le:S ill, palt<:1111~ to he translated We rhe C A nt~Ms" <II(' A3~ Tng()lhcr tlley IlL'mlEcll:a 1]()1G'J]'ia~ ll~umm~1 I!lrHUlghlh-e Sllfia~~Cl'iI. i n:l::J'e'Il~ rH<1)'be ~ppOl'iIX~ d :i'i.lilb~tmtc lor di~ambig~I<llikm of m'crll~IT)i[ling hy IJ;vtlClrn comt~leDi{m :1~1,\L.>G~e,~iIII ~hc CAS !OOtwork P. 3, 5 9}_ VI/helher d!,)eo.rreltll~inn in mCII)tldc~ ill ~he hi~\LIJ~p~", Wi'lidi I'Ilcell~~lisll1 i~ ~'crruitcd clcpe~(~~ iO.!fI dac clcl~I[Ij~~ !lYnk~ is. ~~~.slialy 1:01' rnn.e redistrith~ nat!ll~ (!If dl<1ll!~in ~lllr~ll~ '~i(1! ~hc~1~ljpt;l"'~1U'Il= hu~iol1 in IIhc-CA licild;; ~J'mlhu 10 be (h~~~UIIl'lin~d, 1)118, \li.rh(}H QhCC'lwirollU11J:;:111 i:.1 only .slligJlI.ly 'Whc-n I!!Im diKlJIlges in h,ll·lUi !O the hi(lpnqtlllJlll:.1 are mm:c ~Wll~lll~ iat pal~ern sepa r,m[ion .:oj ~~ mooHiit'ld. at ill fux.w ~,I};[II~JI loc;;u!ion, fl"IlI~IH ~.(1]'l!r<ll!'i)(mis cxp'.l'Cg~d il] ,dc;J1ilatC' gynl5: ,mel :JQCrn'I1p!ihdl~ ilIlso by fOCl1I,llLtnr.nI or<ls:tali~1 LOCQIIIY iu\.'oh'I;o'd_ These m~'challi~n\:s" IIl1ipl('l]lC'l'Itoo
("1\,3

h i,:;gliCr!DU~ruJl o !'.[;i. ~Iw !Dlf~~ mil! t1i:,;~i'ibtl.!.on (FJ,g.. 'HJml(]r.g. Sll). [lriSCllllssioln. This s1~d~i ides ("Xpt.:;~n'iCliral SliIPJK'i:1 .I.~r a IT{)lc o~ the ini~i,m_~ li~fl:g';''> tOf !he l'Ii[JIJ!OC<lmlj~l~ ~n1 1)~lilAi~,'mi:'1;,1'1<lmti:oll ilud ~tIl!,l.b't:sI~ thai. a dl,mJ s~ Q r 1IIC'~WOrk meehanisnrs i~
11.01

moo

rro....

diVC~C<l1t

llC'li\ny willlin, I~~ ac~iyc oC'.11Ils:semblly. TIle d~~'Cli(ru I Qr~tilc: rn~c: ChrUl.b'CS~U lhe

<Ii llerem firil'l,!; licld::; Q (<lemmA; Cd~5aCCCIiI.lmle::; i:l:icacootm;:lillkm oi.'li'ic euscm~lil: .mc~iiviIY.'I<'hich a~II~I~"$ each ~'1Iilvil;(mim:lit 10 be rcr~~';:lInoo by a

as a Ch~Jig:c in: IIiCpa~~CIii'l

of

CQIIe.I~lul

ill-Hlcpc:rlacm cd~pofl~dmiol'l hlCI\3 C:

~14. 33, J4~.

,elilSilimblesilil

0\3;, but Irlat iii! dlf1'Fll<ll~ 9y.rlls,~her' li!rg~ ch~'n!ll€~ 'i~the recQri;l[l1Jg€rnvirOlf1lment LA and B) IFirin!]fie!ds off a~l ce\~ls Ire{o:roded simull.lnoous.1'f

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hom CA3(A) ~_rld the derrlate !mY,ll!S (S)in '1111 .mimo:l~ ,expw ringl b (f.'(e!l at va rie-d 5'1.zea nd' co~o:r n iii [\'1\(10 diffr,r'~'!il[ rQQm~ (r,;:Itn2Jl'Sl. W R.ecoIf(Jliil:l~S~i'iucn(~ 'scl1emaltilC al1fllspatia ~f1ring {®rrelat~ rl)iI"a:~til{eeells r~cmded from lielrod'e:s fi'! (lB. 11l~ r<!r~lr:aj@c1iOr'f (gray) ~'iititJ ~ilIpe~if'I'lpa~d s'p1Jtelocations (r~dI dots;) ls showl'! ~,~~ ~~d"~U. Sjl~lltQ~U:;i !ii@l:s.h!;ll,'l'~ W"" ,j!1~ .8). ~B) Tr.aljeno:rifi';!; witJris.pik,e, ~Cica~l Ciilsf®r dentate {e~ls rle,corded simultaneously wi~h the CA.3:ceUsilfl (A). (See~-~'g.>.1:0 f®f (t)'ler·c.oded rate nilajl5..) tC~

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'symbQl:!;)ro:r Ulie entire sample JOiff (jeUs rE!(,ordedi~ G~3 o.~p) 811i11li tI:I~'lfIt8Ite gyrlll:'> ~I:mt~om)' (n '" 1:1. <'! Iill!1i!l ~Is.. three witli! si ml!ltalileOIU15 re(~ ~di~gist(ir,des iQf 119 i1~~~~Q[g r hu;lli~;1IIt~QIi!ll$ rHIIIiT1I~ri<':lls \'iiil;~ 'sfmuhal'leot!!> re(®f{:li~91 ~I'it di!!n~,lil'e gyrtls and (Jlt],; dar~~.n,~ddr~t~'$ illld~altl£! (,~Il$ llila'~ ~~r~ fI~ool1detl '5ellar<!tel~ilil Ol'll!?!brain r-eglii~n" ~D" Cr!;l!i~'"i:() rr@Liltij,olil bel~en rolt,e lIiIalpc~~r crll~ wUh liirijlng nelds;in b@~h r rclCi'f'liliS,eale !1Ilia ledl as in Fig. 311), but wi~h [yri:or FCltM:io.n ~f tlile Inl~I!)$ n~:latk1l'e~o. each 'oli1~r and Mtlil <l1l'!3J lyses pE!r(,ormedls,ep,1:lllr<!tel,y ~nJlji\l'ld'!.].at a !'lim 1lI1s"The lijli!e diagrams she .....cross"!i;()Jret<'ltions· 'ren 1I'ldi:"';d1u I, ~pwiments as a fUlrt(!tllJ<rl,Q!r dii~ia!IiI(:e ~~ hom the ~en~r cd tH'Ui! (rQ~5-r;0l1r'~~I;I!;Ir~,m {on~ tiru~ f(Jr each alfllm!all~~itltl 1lT1iI),i1e' than~!m'!e silFillJltanMLlsi[y r@'(lorded cells ~Iil eil:lil~r sub~ietd)'. Cr()~s,'"i:oH@'ldolil maps (at ~irle bOl~Om) are Of'Ily $hoWIil ('Or ,.lilt .U2'l5 ~:CQ~r~fK)'nLli]!1g~@,~ti.hlli!d m,l, fS~ fig.,S,:l2: Ig:r tn~ r~mali,mtil~ rn.lllH.)
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~KI'\'C--pilr~ic:leca lJ:llhy (if tI~e l~glHficld,. I!.] lUI, dcrsmmj Uu:.'ilr nU:'fl~ll~l~g. consider llile' $h\~lcj:llflotollllul<;::r~;"'l\:lloC' !!:lipcrimcl:1[ skcu:hod Irl Fi,j1. I. II'Ilhcctoo~'(j im~dcrorrn';:ler o(mllgr.l~~ iOJ11. a ~in:!lll;o--phQ[(111 IPLlbc i~ ::-pl ~t!by ~. 11rst llCl'!l1rl' l:f!uit..:r BSil1P.!! (Jf II ~'l:'u::"h~ZdmcJ~'t irirter~rOJll~'\i.cr uiild IliiIv.,;l." U IIl;'IJ~'iil. i.Iti'li.l a :sr;:U1Iiti lL'lCm~'i'I:>plilll!:it BSo:>.i1l:"l~ rcOOllibin-lc;:,; dh: twu ariiLerLicrilll!l ,tlrtliS. \\i1i":l'i Ihc plm •;e :l<hi~L 1Jc1.W~';:1I f!i.e tWO ant'l.~is . (Ii, '1'ruled. inr"x.rcrcuace ;)ppt.i~~ as [!L nlO{itllllai.ou of t~n:: oC''LoDl,h;m pmlmhili!.im'ial.oOIlIIUJ(UL l)OrIS I alld 2.
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~t, ll'IoMlU~I~. ll!io~~ NOO1l!l~~~~rioeurede Cdd1.a!t UMR CNFl1i 6537, 94l~ 5 Cachlilll" Fr~!1.{e. ~K!l~ tab!l.ratolY (!I~ Opticalll!oo Ma.!jne1ic R.i':~iJ.m(lC' '~d:r(W!20I'Y, EM!: (hi~i! ~IniII Un~'!I!1!'lii!lI', 2:O006;]~ SbiInignai, O1~niil. ]t<looralCiile OI.arlfs 'F.a'b~ de- l'!I'6UiUJI. d'OJlt~qlle" ("lmJ!!l.l~ ~e(liImquf;; UMR

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ri"h~ 1~~1~ ,qummmn !i;lf,. m~I\,~ by boOth iIl.l:LjIK~." ,[ 71. ]f BS~!IM JS removed {Ule: OjJCtl coo iig,~1'" Wo.lIJti.oJ~" '~'f~{;h dc'~1Ctor DI or 1)1 OJi the OlUlplil
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of the

[Or (E:OM I WiLh i~tJopcic:JiI~:Ii:is ori!lUctt .ll[ 11.)0 .. from i~IPll[ poh:nizmuous. 3Jit.! ;:1 \VO!l:JiSlO.UI prlis!J:n. The iwobcams of .he :imclll\;''1'Olrte.,;J'. \~lldl are spii~iffll'll}i sc(lfi:I.'flLcdl ilIK1 OrtllOgon~ltypoiarinl;,'il,
arc iir::;1 m~rlfJI\liP'-il. by BS' IbLY~ '-'iII1 ::;Ii II DC ml:.lll'lhi!!!JlIOllldy iclclltliItld by Ilwir pok'lfl]:'!:;llimll" Thcill\.lIm'i:hokt~ 1Joi1~.~'WCI.1I1Hl ilf~'rrilro:m~~'r tWr), coI1.Higllmlicms... dosed or open. ~. realized wilh tllt!J roM, w~lidl ean 00 ~-\\'ioc:lK'd bCrWTIl1Il~wo dimm:::m ,w]]:I.ig.m·JI.iDfI~.with 111. 40 us by :Ilwan!\ of a ImuJlehllil~ J:m..~dri",cl: {ltl}: Either 1m' \'Olmgc is appliicd to tI~EO~1, or iii.'; !I~lf-"'I""rc vdlOl~ I~II:II'> app1kd ~o ~L I'll lih(} fi ~~. ease, dIe ~i~Ju;I~.i:on
COrt1CSi,flrn,ds~o Ihe ren1'Ov;:1I of B~.fl'UI: and !he tw~ 1ll"I!~I~ :I:t'rn;lill !In.crnn~'lincd ,~o~n ClQuligll"

OI'Ul~t:lnEIIC[, one

Tlmfl

d,Kl

photDn h~1!Si t:I':iI.\'t~kd ,maly


au

IUlDll1" (7lStlch

C1{p;.]II:nmOI.~LS;LljJJ)()rli.~

BoIlr\; sl~n(,I'lK'>l]1 th<Ll Ihcli:tal1mvID, of:1 .qu:mtum ::;;Y:lq(,nli.l:;: detcrrnincrl by Ihctyp,.;: o¥nu;l:mm::mcm pcrfcmnrd ml ,I:l ~<'!l MQIfOO\'cr. ~l is dmll' IlmL lal'
,th~~wQ
cm'llJ}h~1I11k:n!11f}' 1llf;'aSUtelllChlli.'; ~1lS,id).

in. ~ ~'OC:hC'm~C~lD..";(! Ehc ~dc~11 {l!lii~nall pTOJlof!i.j~1 111:1 lFi!l- I ~ TJ.ill eheice ~,D' irillen or IilU!:W(' BSOOliJ!.i!L .Iisrandollllly c1oc.i'dcd l~II'!:lugh~l.t[!' usc of:1 qmin.
u] III T<JIl1.d(m~ 11]~llUbel' :!!!;C;]lCl'alm

(QiRNGl

TINJ

~red ,hem, ~lCl OO'lllG~fXlll~~:l:Ig (!xp..-n IlIla:]:ta~ :'1m" t.ill~~ Imltual1i' exdlLoovc: lhat i\. BS_"",~ cannot be ~in1il,IIt1l!I'lOnlll:i\ly Ill\.~Cfr~ ould "OO~lt, [1'1 I,)l{,~l~ri I.Uell1S. where the C'holjc~ ~CI\\'~J~

'I~

the' two

"~ilUiJ]!}<;,

i~1~:IldCl ,~c",~g aJv;m.cc, nne ~!l

cnuld reconcl Ie Bo.hr~~ 'COlllr~k~lIncn1,I;I:i~}' w.~lh E i~1~' .;:-hll·S ioolli c'OucC'J)' ion. of ilile phys ~a~

cxp,;n 1~1.i!1~11I~ COat'li:gl1rm.ifm :;Lildlooll~~ lilen .'1Idj !lI~1 us belllll\ im :tIccllftili:l'lg,ly {91. To 11111..: out UJliJl~ wo-ull'ivc
ill,lolili1111IC!lf· Oili HII::'d~~~i

iTCfllhy.lndct'tl. ~,,·hiO.'lll ~'1lw'QI:Iemcrs 1~C.e~Iflt~f" the R:')furncb.'t, it oOlilMhillV>!.: l'I.~i,\~~~1 ,~OJll~"~liddcu

inh.::rpJiiDl1i.li.tm ,uf 'q LJrulLllI:trl mechanical coullph;. '\i'!,rhcclcr propo::1i~"(1 UiC ··t]oelnycd. IClrtfjlcc'''g,c(hmkc:n c1'>p'Cl1mCIl~. lUi which Ihe choiec o.~whk,tll Iprop.crt)' willbe observed is
Ulkllill3rily.
~Q~tc or by 1J;o~1I'OU[(\'\ afl(lf jl ~.~ alr:c,ady doneill.; t~ .. ~T'f 7t Sill~\""'lbl;ld~I"'!\ ~)JOil1>L1l"11, .~vc.m~ dclay~1.L -choice e.x:pl;'dmClll~ Im.... IleoI')l'l e J-q~d (W 15~, II,m.ce..rer. !1.()IlJ;,': ofUlcnt r~illy fb~knv()d !he

nm.dc d~~'iT Ihepho.mi hH:;~:ii>!.)d BSi~: "Thus 011"':: deeidcs I.hl:'p~iottll~ [Sh1l.~~lfl.ll\'(: CU<UIlC by Om:

QRNfi is hJC.1Icd close to BSOO(fljL and isflrr cmOU!Bh frmnlilc hlPIIU 00 !~~l 1]0 Illfo.l1n:uKm abo1,11 I.hc ehok~ am reach Lh~ ph.oton herbfC' it P<I=s, !hM~~g;h BS:i""LOI001I1"si nglc~pho~rm :-'our'Cc., '~I'C\'i.OWl~I~1 develop;.,..-.J lor q!)l.'lnlllll11 key d~lIrib!'!ljf.ln tJ'6. 17), is based 011 tll~ P 1I11l<:ed. I)Jllkally l".... dled p:ho.~ 111.Hllil'lcsccncc ora ,Sill~le nilm'gcn"\'1tCo1IlC)' (NN) oolof ","~mcr iu & diamond! nil.liml)'smi QI8}. AI ihe ::;:inglc-!!.::rnttie<r .. el, these p~'mto:iIUll'lill~~[lt cell" k kl'$, wll:i,,11 C'llU be illd;vid~mlly addl'!:';S:>J;.-tl wi~lIi th~ use (}f ro[j~bei'iIUjiicl'i()scl1py I t 9~ ha:vc:.I"t!Sw,iIl llJi~lrp~:.;fi:Ccl dYie.i ..".of,,1' mId plliiJl~-'.ubilil)' ~IlIrool11 l~,nperulitlre (10', 11). 111 MlIi~iulll.ill il; pa~~ilJl!:[{I obmill !lirligl'cp.lflllto:lfI.~ \11'.11111 w'cll-dl.:!in~d polar11 iimtlon {J6, ]1). TIle dC:i.:ily(.-.d"dmire:>ch""':J11CU5 irnplcmcm.oo ~'i ronbw~, lll]~<IfiI),fl'o'ljni~ single plHlCOlllS m...: ;:;'I.11H by tlpo~ar.i ;glli,on bc[lniSp~ itt.:!" IJSiil~lOJL lhrull~,h l!IJ1. iliC~.,'d~rolfii.cL\i;;r (~ln~I'I1 "fl. nn~ witl~ H.... $~~!.i!ll.lIysj}iml~"r;.d P,I!]~~ a~~od<Lt~d W~tlrl ~]. orthQ~OJ]a! S <lJ1'ld P Bm'h1:dzntiO'J!i' I( fig_ 2!- '["h~ mClVabk olilput hc;llll1i1pliMCI" BSOOIp.;it 00l1l:;;iSl,'\ of ~he oo:n~bin... in·n or OJ hall::wa\'1;' (lIme, OJ lilOb ..iz ..," dOl! l.'J!i;'anl!lp.~tU~J: 3S'. au c .lC(:Ii&op1:iC'a1 mOOul ..," 1
J

mlion), B;()C":llI~' ~l~migil""~ S OIndP (l<1lai'" iz<Ltroll:-1 orille .wn p.. ,hs .[]!D'e orIt!ut(,d alr:M1:g pri~n 1'iI!')briZ;;b~io]] cigCI'l~lnte:!i;, C:'JIdl "click" or nne dc.C'CIO\T ,~) I {lI.' 1J2 ]lIMed 0.1'1 llie OLIIpt:I~purt., is ilii:iocimod with l.l. ~peci iic pinl] (r~lhI. or I~llh :2, ~lJoeC~ ~\!1dtr~.Vh~"ll ib.: J'~ vult'lSI.: is iiPP'~tc~, Ihe " 120M i~C<juiYiiklii. 1.0 :rl 1:I:.I~f:w~r\!1Cp~lI!1ic Ih~t JiU.'ijl~""$ l]lC iuput ]lOlmil1l![i!J;Il~ IJ.)' mlllutl1c ·0:1.'~o. The pris;m [I1~ll rc,;.(md)hlCf>~'I~ IWu [(m:m:d polati~nionL" lhm h~\nt,;' uavcled alullg dlffrl:<l'(:m op~icai j:l1iultl1. and j u1terl~rc:rlc'C i~ppcnTh ,ObiIhe lVI'O {)~IL'Put PO~I$.. ~VC lh,.1l hUH: (i,1e .ci:05CG iU'[cnCfmncLit,;r O{mli~LmU[~(ln U')~.

Fi~. 11.,. I1toe!s's di~1ayW~d1.oke~!M:IaJl'lken W ,e!!:pe'limem propol>i!LTheo (hC!l~ tiC! irmo· Ii1l!1c:eOr reiiTl!f1i'of be,am5:pl.il.l'elf BS,~ (dosed, Qr Q.I)'IlH "~;mJi'gI~!r.~ticml miW~ ~;HllI¥ ~fter h tile pMsa!J~ ,of th~ photDi'i alt. ESttfl'Ol" 310, 'lhat the 11 h,Qum entering tl1e 'il'lterfierif)m~Mr "(.liiillr10t ~110W~

mirny,f
Single-photon pulse

r....:n~(h of

tn~' t!Nti'

'C:Cilm.~leme-nlillY ~Jlperh1l'ierl'~ fll)<lth diH!lir'~m;eV!N~IiJS whkh-waylMU tOO perf,orli!l.~:

Palh

1\

To ~'I:;UFC lilll: rcl[qi;i\'i~tie :;pacc-lik~ :>'-l:!lIRIlion lx:1.wCl.::u til'; e]:Kli~"l.::of Ill!: ulilcrle[!.':lI~H.::D~"I" orill.a;o umliol] and the I)<l~g,(' or ~hc IIlmlmt :illB~~~~., w\;' oonfigu'!L'{~ .he 80M ~wi,ch~llg ll~()t;;e$"" 10 fue I!<IilJ'ldom!y decided i~] ~I! lmc by Ihe QRNfi ioc,rtQd ch}:'l~ to 'he nUllHllofl.llCl il¥lCrRJmmC1.cr {4i"l In iiunn a;i';ii1j:l1J~), The l'1momn OUlll~' is ~n" ~ml~~1 by ~illl~I~1 ~hQ' ~~~pl i!!g il!eo;l, ~"n(l:l ~l1~~ .of;:l ~lI'hi!e-lighl ~111~" mIDi nQi~e is 'II~ imrin:.1ic qu;mM~ :1'ilJudfim IlITio,."'CS!I. ,Hl~~ i1!1, \, ~lll~,"'~ .. giv<.'ul ~~lneC<lWU'lI be p~ed.i,c~oo21). The li!nin~ ( of dlC 'C'xpC1~ifl1C!H C!!S~I:J'C~ [~u: !'C{luh'C{~ roclnih l~lnc s,r;.1ce-~ikc sca-mmticm (21)., T~~Il. 11.0 infonn;:uio~l ail(lu, the i!1i!Jc~'rt'mmc1tcl"ulrn[lg.uDUlio.n dw'~", '!;.:Un! H:~adl 111(; pl;IOI,OJ11Ix:l~'!fC L1 1.:111(;1:>ll'!.: illllcrli::runIl.Ch.:r. I Thi~ :l.in~i::-:p'liotOl] bchm{illr i,"'.~l"', li,r.:!o\'llc5Il'd
u:cing til,.; twtlOLllptl,1 !.klCt:I,UI,',:; I't~~'tlnllgi.l'l~rcl1~ld ~

.a~ thE1m.npUit~

961

REPORTS
coincidc'ltCe
'(IOJl'CU C"O~mCffi

wi~h I3Soo.,~

r'Ci'llo'\''Cd

con ligllrn~ lon l. \Me 'lISC'rl flU ~11l)rof!£h similar to l~tCOUC (]~"Crihcrl iUI fl'l' und (6'1, {'OIisiJcra If~noom..,,,,poJ1tliligLOj'Vy urlu,crplii:;cs o!1p-plicdro Ihe CiliJilih':;~, Ulh [,\11 «~lID1I$ dcK"CIOO iu w p~u~h ~ or the inlcrfa'Omu.'h~f ~y DI. A~ cOIJI1l5i dll.~ti:\ctod in Imth 2. b,Y D2, and Ale d!.~roilld '1::6im:iOOllc~ ,~(Ifl"(;-';f;31Ondillgl[l jOiliit p]lillod~tccI iom 0]] I) 1 rll.lI:ID2 (fig.. 2). AL,!}' d(h'lctipl.iml ill wlili("liIli~~1 i~~",,""IItcd.,~ 11 elassicnl wa\,l.!, ~m::h 'I!S. the sQilllic-I!~'i."'i.iC<!] dlOOry w.illl ql!l:ll!lIi,,_oo p~[m.. de[cctol'::i: (..?4), Ilrodk:l:;, Ihat th~c ~ll]llOOus or counl~ ~hmatd ~y I.Hmi!~cq{lO!l.liJty

a1.'Iool.ll1c' alr.c:My pilli'llhil>LOI)' of UlfLL ~lhOlL1Il" (7.~. ol:'lbe or~ k\a ~conlP'Ultemf';" The same value was OlilC,;;ntOIC. w,;; Ii tid ~llmooul~ benavcs UlI obla~uC'dl wilCl1ItlilC mher 1l~U-IW'f~ lJlG!ckoo (.c,~., l)fuJlIli). ~n Ie .opelll ron iigLM~niOli.\,\"C: IltLilii,l~.vc ag.rcemc.nt wjih the ~lrOO:iCli()j15 cd quamum tl i llliCdl'lllt-c$~VC.U'1 iii slIIrpti:;;ing ~iJtt!~[i:ous \'\'hC'll~ 3 mll.llim08~k~~t wh~h"~\!3Y rn.oCasIIM'l'ilCjtt. The dc·kty..:d--'t:h(1l~ ~:;'\p;;rili1lcn~i I~ICJ II' i::> tension ~ idl r..:lilli ...tty i$'CCfllSOO f~PiJc.ar {29}. r~JCromled \VUdl IIhe EOM r.:mdo:ml>' swi~[:hoo rol' each photml ~~I"It uno lh~ in1IL'l':tkrol'll(lh~r; oo~n;:;:pol.l:I:lillg ~o' <I :Iall.dom eheico bctwu~n liI(1 open and dmll\d C'O.Il~i!l!lIJ"Jtion:;,. TIle pha;;c shutt (b bC:IWC1::1111 the I\\'() ilil~errlil'OIl'IC:Lcr ';:U'I!lli!l is V!1I,100 by ~ihillg lim ::;ccond pO~.. ri7_"I.km l}C1LllD~Jliu1Jr .BS' with <lllicwelocirlc: 1lCt\1lI1COT (PZ'l'~ FQI"CO'Icia photon, w~ ~or(~d~Dl.e chru~1III (lonli§l,lTalioI1. ~h(l'detection everns, ,;iI!l1d du~PZT pn:;.ilj,l'ln •. All I"J;W drum WCt'iC' .~" cd in I:~nllimC''''l1d 'WCI~ IilITF
OC.'''\''lldonly after ,<I !lUll
\'\,'&~ e{ul1p~cI;cd.

R:il~~:re:n(e5andirMolle5
It IB, !lt1!j!iIILiOil\ M. !"s;~~!t!'!.e~rj_tft':l OIl' f'h~!fiS rJ!,tk!lI4I\'~l'e)'. ~ad!~g, ..~1!i6'5)' 2. ~. Gr3l1g[{oJ. G, t:.Q<.1:tr. A. Ajpe~ fij~ 'NL :;1:, l'i'l
II. ~. ~. f'~!lI.

~.~r i;',;JI.ch o

PIl fl\'li'1Ltioll. d~c~il'lu t'\re!'~!l on I)] :lind D2 oorres.pond il1:!ll to each oo:nffiglllmlifNl were Sm1t':(i (Fl,g. l, In Iii";; do£(;(j OOlllll.bt~ll~itian. "",;; observed \"loliu3.0n of 111, inlXJ~Jilli~y 'L1n!l~ gJlVCSfL qtl;)mi~ .. m~iYc ,cg'ncrJoul IItill charactcrizce Imllclm;::;,ic-a~ inL'C.r~cil~u~,,;w~lh tt94 vi~ilbi:liIY. We mlJibtl~';: th..: lk:pmillln: rrutll un i,~'y lu au imp .... rli.'.CI. ... tlv TJrL'P hdIilV~O<ii.For II :>i~tgl~c"rhQloc-m \"'f~VCPitckc-t, (!iU1In· tu I~l up~i,e~prudgcl~ p;crr~l, iillth::orv.:elillioii I( i.c., of tbe 1,\\'0 il~iClfrcnil'l~~IUi1;:". In Ih~ Opcn een!!J: = 0) it! 9gro:'\I'I'iI~~I'II ith UlJ.:::im!!!il uvc urm~1.: 1lim w Ii"" Gl[I:OU. ilrIiL..:rfCr'i.:m.;: l.ut:~I:I)' d i.'iI1P~l!!llfii.~I$; of u :a shlglep~lfI~k C~11unll Oi.: d.'eICCted sinlt1uII111, evidenced by 11:1,.; ~~bsC'uc(; rl'lt.lt1J;~I,iorl illl the t '!.~,'O(uUplll pons When ~In\; phase 5~1~ (I~I II .UlOO~L~iy ~l Il:te tWO IPmils of the .umcr!i;u,olJK':IC'r « 21. \¥c ntetl:Sllrcd 0: = {D.:l2 ± (1,(1 ~• hculCe we are was ~',~rioo, \1!,1'(; clleckcd [Inn iu Ihe t1dnyal.

indeed elosc III IJ1ePfLU\: 5ir~:ic1)hmo:n The flf.lflklCtlI \dlm~ flftl1C (l paClI!lClrri~ ~'e$!idl!l",ib~ckgmuud plll')[l'll~tiinin.~nee di,ltl1(md ~,~~lJ~I!e alld In Ihe !woOi}1ll'lmm
~ilit('1liug

IC,gjm.G,
d'LI.~10, 1"1:["
d.1C

choice

Can flgurul ~(HJ" Ili!IUiCl.Cil}i.


~61i Ilnl,,"il::>m,1..'dI

a: mu:ll

kept

til~~l 3. f'•.Itl~·mt.A. VO!Amar, I, ~, 1':" iK. ~elb.a~ J, 'fi';f~cht!!J.il', PIl»- ~ A&', ~l'l!".OO ~~O!(ll'" 4. Jtle<i~n!Jllr:, 'ri ~ih!', T_ J~mew~1I" iMc~line'ff'r, No_4:;!,:!, nil G!iOOl5-i s. r, Ai~~~. IIJ_ Henl)g<. Nt ~kITHI~~~O. Ben-;~, 1ilP' GNr/ J'tol\l;. 756. 3~ ~O\Ooo:~, g._ V, )J~\Ie:;; ft Qt, t~, t1\i~. t .Q3'5, 511,t ~WL1IlJ'. 7. ~ ill. !ll11H!~r;in Ouailtum ~tMd N/tmlNt.J1h!l~;; ~ ill. wnH!t~c w. e, ~~~'ek. 1M!. ~f'I'iilO~!t:4iiIU:n.n..fl~~ ~\timootlill. r4J, :t984']o, 1!llJ, lIIl-2H II. rij. SJ).'Ht. in (lml1l[1IJl'l II'lrory tlrM1 Nl'illiiINl'mI!'lll!', ], Pc 'WlIleeleif, '1\1,. H. ZLlr'e~ E.!i~, 8'liiiKelJoo Unrn. Pre11:, ~i~oo~liIl.rrlj, :t9ll:4']o, IW,S'-~'il. 9. G. Gr'eeilu~~, .1\. G, hjlffi(. Tm QrJ~M!I!ID '11~~ng.e ~~es ~!ldlil:a~II. Soo!~u"!,, MA. l~~'n. :!i[i. (O,,I!~lt;r. O. G, ]!arullo~'t w. c. imtl\:es, ~ fir«ffi!~oi l.h.eSK~fJd iJl.It¥MliQl'Jt1l Sym-p~~OOI.(lJ'! rile Fofil'!lf!J~s oj Q,i.fdilttml Mt't4Mks, iii. N.af~~ E:d. ~!f'!'r!ls!('~SRI)' ~r11!Pll~\ '!:'G!:}l!I\ 19a1~, . 36-47. w ilL iT. H~I1rn~, H. W.a,tlm:, Jt (!_ ~joo~ W.. '5~'eil:in,.
Hl»- ~ A,7Z, 2~ill U9SilJ'. 12. ~ ll~ld~~lln, E, l!&lliiler, W, ~r\iell@~m.l:~.
B

the ~·~llJ'IC vil'lm:-:;

LII th..: ~n;:IJlni·

17,

mLry h;:~~:;1}R::.~'\nhld above,

R"ulOlu

Our ~liZllln!01ll of \<\I!u,:ckr·s,


g~~l]kcu
I;l'Xlflcl'iun.CII];I.

ddiljlolil-chtlicc
thl:

H~IC,~"'~lidl bOlh

pFOdll{'lC' I!InCl'lIl\!b:~d

'r:nl1.o1o~S w.i1lh ['flijs~nli;Ul 5't1llrimics (,11). \li,n~~~ s ill!lI!~p~mtQJIl PH.!ses in the open .col!lfl,~.urnukm. ~'\IIC' C~lflocted cm-ch detector [) 1 <llld 02[0 he un<Ulrib ~W~DolJ~I.!lI ,J~$(JIt;i('l,tQd w'i~h 111 ~i,vel!l p.oi!!!~ !fil~ iln!i;'fr~1)L1flil~tGr.. Tpt.t;lSI 1111:s. pflbl1., ',~e c\'a~~ffit~'\~~ill)' kh-waf' ~!lroml;;);til'ln ',\,'h '1~<I!m~lli(lOC)r I (N! ,\12)I(N1 N]I (2'5 2N) il)'

bC'h:wio;. fir tl!lt': Ilh('llOll i~] illllrt:!r[('l'OrneOC!r ilK' (~C'iJ1e~d~:1] tile choke or ~h(' ol)s~;!<l.MC' tl1m is 0 mca'>LIr>Cll C'VCM Wht'L1 !hnt 'choice i'i; 11~~.dC' Rt ,;,11 ~h~ ePl~V~.1~ fir tlu;: photol1 ~n!iI]~h~ i~r~rfi;-oorm:t~r 11>, ~~~~k~ int~N"1. In '~Vhecl"'-I··!'l\vord:--1.~~, !lO ~.i'iilU;L~ Imvcl i~llil!:l~ .;1 VC~OC~l)l k,":;,.~, nti![I!11!1m~ o:f light 1;tI!1 i;',,()~!11.ect th~~ Iwn C\'CJ1ts,. "~'I!"C' hllve :jl S!!r.lPlf,lC :i!1\1crs!fm(lf~he !lOml<l.~ omC'f (lif timC', We'. nrn:v, bYnl1[IVi nglll;C ulirml' in or OlH Imv,c- 1IJtl
Il('l:'li!i'o.nn11.\1 >L ~~I1lC :!\Uch

d~nlJ)"~tmt(.'S, lBI:u

or

il,,11 it is. ~P-;'II':;MC'd, r,u:m

,I

illl!! the emu] I ilugr..uc~ m D.I :and D1, Ii. V;~lllC (j r I ~llighcr. IhlUl 0.99 WlII'f> Ith.:rl'l,urc:d. hlfll.i,U;),jby

IdL'ck~u:g ('I<ueIll()th (c,g ... Imllll~ and Illca:;,ur-

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d";lce~or dl:n:k m:.1U~U ilr.Uj, l"C:;ium:l1 i:rlllpcdcC'IJi}lla IR~lJll~ Q:f lh~ debyed-di!',,~e ,e'."!~riil'ii'e rlt. 1i1il~ phii~ ~ifl r.b (ilil{il~~J~d \\Ii~h .a r:bitra ry Drigi.n~is !!!'ariedi blljtijUnl'lg BS .IE~1h1p~ilfl~ ~oorded \vill1i3!!l!Iu~;itioril

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[1.2 hed IfDlilt~, have belflil :lllll[ln~ra(te dfrom tlile ,date (A) PiJilfUS} allild

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.~I:m[J~ i)i NJ!J!1

16 FEBRUARY2007

VOL 315

SClIIENIOE

\I'V'IM;V.sdencemag.orgl

REFORTS

IMultiipll,e 'E,ne'rgy Scales at a OUlanlum 'Cr'iitic,all Point


{. Gl'j'bel,. :l! If., S.te~lic&l,:ll 1:.
P'. G"'~lirnli'!'§)~t,~*t t, We:st,l;i,rk:aimj:l,'"
AbrralhOlim:s,Ol Q•. Si~"

SLJ.eil

nmlkC's'll'RS
aC'J.{)$~the
IfIi~n~~l,ic

a small uUlgncde field ~Q access nlC QCP SlI! iI.e(] ,~r our puspose : Uti: deterscales requires SCfLl'Inilit phl"l.'>C' It.fHilSi~~(Wl.~nd ,1:11 ex.1.cn1:mll
hlll:ted

Ilninm.lion or energy

~icl(j C<!1l ~

with relative

ease

c. KreIU:ner,l

y, 'fQkiwa?:I: :5. Pa:iid~elil,1>§

and 'i:onlinmmsl)"

U 31

Hll1l~ dloct I1llCllSI!lf'!._"iIl!Umts

We ,~~p"rl~ht~rllTl{!d'~!1(1liTllil;; rru~i1I's,ur~nn~ntli ]n ol f1lIiJgn!lltk-~i~I~-dri~n !q~;:IllIlulm uitv~al j[I()~iT~ Qf' ~ ~e~y Jerrn!©f! lIi!~kl L Y1b!l:~2Sii2"'Tille d;:l~~ pr()vi~e f"'idillin~e ~glr ~1i!~~1J1Y .~~ ~e in Ul!~ ~~Wl:llrllJl']!I ,exril.rtn'Gn ~pe,c:tfrull1i that is 'i l'Ii11rJditDOIli l!:i I.he Oil'te e~peded f'aollli the ~O'l'l fll.l(tuatj~ms of the 'QHiI,er par:alm;eter. Bo~1'I Ii! fle ~(J~ sClllle~ <lIlProillcl'i ,,'lIiU'!)' 1I'!lltie' qUaJliitl!l fill ui'~l(<lil poi Fit is r!!i!ad11et!1,t~elreby pm.'IIid'iillg e\l'ide'lllcl!~'®r a flew dass 'G( quantum c n1 ~ic,alily.

on YRS Im~'(!' :;hc)wn :J hng~ and rn[1 i:d crosgo,~rcr I!] IhoClIlaU eonstant m a ~(N:UlBlCirnLtlm· ool'JC'll<knL m~g,rlcliC' fidd m... y rrom the am ircr· a rn]]'I:Jgr~Cl1:ic l"u~i.tiC!lJ.l, h! the ~cro-;t{~ll.lpem~UR~ Iknit. il!his. CITI'!lW':!']f oC'xlrnpolmt\'i; ~() a ju nlp across the- ocr, whkh has beea i!llII(!,lp~I.od as a large dn:mge 'L1.f tim Fermi ~~[I{~lJ~ vdhillill~_

(leI? is ~Q' n]ll.l":O;I!.t'Clhef!.l~Ddy· j:1ItUi! cflml~~'1~I1:S;. We liIl~aS'lI~I..,,(~ the r~o.1.lu:.yma~I,inC<i!l' magnamic Profl(:'rI~Cl'i:, AIlOI,h~r anlfl'madh i~ to IIlClo.,>u:ic"~iOJ~ t%jj~. INI; whU'i:." t: is, lIil;,i;"ICiTlglh sceond-order nlhifl~'; ~11t1,'1Sil m 2icm Lon measure 1.11"" llucunnlO<l1I $pcc~rllm an cqllli~ihd. lun. I:'or .;.,\mlillP~C by ine.llm.>;~i;tnctlarorl ilICi'lUCr· '~ClfnrCil1iI,ltIrC, n tllldcdic;;, the unusual Pl\lrcl:~ie.~ fLJOIli,!: he II m~ di~l,ion t W,ilh~1l ahc~c~~gOmdl ah p~ali..:.,"m;] iihc lilllIIgii~q,ic li,l:J:d N i~ ;IrDI)lli,".J ill~g ~xpo.:DlImlcng:;" Such I!XJl~il~lJriUll!l mctlsods (Ibs'"'twd ;iIIIa h~l, olr'llIili:llUm anah:l'ii1tb;.A bii• sie are iii OOiUril:>t 10 'Iral~p(II11 'c.'(l'Cr,i ilt~'tl'il);.,.wh ic'I,1 a~OIigIhe ~1~iltcdin.'lClioa;, ,(fIll'). FWll1'C I :.]\O,,,:s quc.~~i{m tlim n;miiin~ lllt'>Cl:tk-d ()OII()C]';ri$ its 11iI..: 1t'l1~n.t:ltll!,lcie ~ilOli W, II ~lluel i(1!.1TI ~)U' the Hn; in illl.cuecdlliy C'I~C!lC~,rulilj;c relfJ.!Xf~litlrliL~prop. pro~Jerl~;,oor'~1,i(!l.d(]~'SlCriIJlit!u, ~liihi,C:~i, 'C~illl~ is 1ili'J,gnl~li~ ~ldd.al. l.emFh;r.'.u~llrc5 atlilging [run;1 , ertics, ..:spc(;ia~l)' Ihr m'l~wuupic :JIINJ multiICQ in,g becflu:>e Ihe f1 U~tlll~nilom;; 1i'1!'bonllcoHec:~ 0.02 K 1Q oO.,g K.. For 'I;;mpc~!ln'C!> below 'hvc and q!J;)rrUHlt~ .Dnoi.\Chi'Ul~ical rL!! class of O 1J11I1~l ~,J':il~n'lis. (I,{I"15 K. tl clear djl>COlll.i~ltll~ly ol):>erv,I;,'(] is Ib extraetion ,of criliCflI Cnlc!);.y $Ci']Ic:;;rc~hool')i. hll~,,;d.0[1 (taC' IIlJ.dj~,i(m"d fO.ru1ulftdou of da::;$iOiLi ()~II.Ic,~jpl'I(::OOI:llCili'lU), 'COli;;.id\;r.;, the (]ui~ Ui~:Slllreflit::IlU;S, IJuou£h 11iuI;;~l>CpS ~.h(; wlll~nl ::;!lI~):I)R..~iugIhe j\f o:rdc:r 1:1)' 11 'l~rili:(;~ij control pm:JIIlfll.:a, ..'If. \ ....hi"h i:$ IlCOLr.1.y iUIIJ}ul>:;ibIC m!lguc'ti..;; field" A[ T;» 0.075 K, ia i:;':i\1 ..n Umt. IhltlUiJllun:o: ~J,( ~. C:IH~;,.~i,,~ 1"·i~ri~lbl ... ~LI.!~lthm·s fi~I;' illJ:h .....tuc neumm ,:>C~Lli.o.."J:illlg".... Iqruri. here iilr ~1, $Ul~ln nmgui~luc (udd, tit'!: irolhcrlrmLiI \c oru~rpl'IIJtnm::h.;r in btL"!1l>pHi~a~~nrlla]:lpOT:l~ ~lIiC1l~llR:'D'Dl~ail~, tloi"tIriClTDliotiYlilLllii{; prup~fI i:c.~ of llIil~~III~l{J~lri.c~ i:orn I in~r~~ depend ~ Oil the li,l!~· dil~'i.\::I~:;iom; '12'- 5). The 5k)wil1~ down {)f 'Lhe y as aCfL'h~<:il~ m:JgUiIllic (lep, (m;h.,:r.c~M1rori1elCrll~tCl~J<lE iDU~ :;tOC·\(lInJlJni~ d1~ !!elk: field. a<;. i~the ~<b.~ ~n ly~ic~_~~m;LlI~ (/4). Rcyo:ml (l cros;~,{wc! lida. hm!,1(!'\'cr. ~hC'l'C'i~, a We ehnse W wmk wi~~I~~1t': l€u,I£;on<l!1 d.h"cmcnC(l of a !ipatb~OO:rrcbJIDI11~ngllh: ;1I~cach dmnge ma lil.iJilh-~idd n-eg~oll \'!I~lh ~ dillhl3l1t v~lm~ of the Ilmil1~ pamrm~tcu, IIIlC 'cq,ui~ibr.i.'I,IJ1a heavy rCH1~iOIl c:oll1lPQm~d YIIS because it 1)~~~1R a c~ea,~and sl{lidlim~w;:tric m,lIcrbl !'>rope, 'fh.;l '(;ITI.'.SOVcr Ii~d dcc~<I4.~" m.~the tcmnll1ill)!-hody ~3le1CIJ'I!,Ub~ colltnIJL"l <l!~ill!! ro-(;:xci~Ol1ion [!1 Il~ ,~~nOC' nf oellCI'JY scale, which v.lIlM~cs at !Jlle qtl'l;II1~!.IJll !!Im~,is ,\'C~~,I:lo)roct~b~ 'I:lCrot~11'C' i~l\'idllCtX~,. To. ~1f!(~~1<'''m~~ tllis (:m~flO\<\I;."'iI".\\' e ~llrp"!'C it ..;:ril!k~.lj Ilfii~~ (QC.~) F,l, ,\n -\I!lOOI~\',~!ti('ln.1~ .. ~ ~:;;I~Wia'l!'li<tgll~~ foc::k!. ylts ~hrnl,.r~ \'~~ W!~~" da!o;sonh~!)" ti 9')" hy ~fll~~I:l'l:;;t.[s illhcrc:nt~j' ,mtifelmllQa,gnclijc ~i\f', mb Oil rt'!l 70 luK, wij~h Ihe Iil)M"d~l101).(~C'nt i«<lolh~I:"l<l!1 ooh<!\'i.nr (Df o~h~,Ilh!;.mnOtilymml.k; <lml !iOU!;~llOf! ~t~1I1ti.ks. ~wall~11111m.:.'Ch')!liC<!~:, iI explkitl.y i!!\lokC'11.~W,~'!r wilh nn fIlUcrcd l!l(l!rnCl'H {ifo'!)ly-IW'\t!nJV~l i\ ,~n!~Hm..g~le~ic I1dd 0:/..., :: OJ~(l T tu~n c:!]t'U~!,l1C'1.1,e!lt c'il~'C~~ \\I i~kh~!~ nlru:l~ fc,;;;l;o)d Fi,,gurc 21\ u!htl<u1!!J;!s the .~~I~~~ari~y or ~he c~s" !i.'li)I~~1to 1h;.1t seC'll in ihe ro,r Ih e ricld ,~pplkd ~~rithi~~ dw C<lSY all Il~... m::, over ill ~hl;lTIa,gf!e~os. ~hmllg,h v~l~is~'!illgC!lI;!!ID'~ii1ks Iron ·iiWC iiuiiid'(Hlion '((I the (l!F~0ssockl1cd with the: slrn:vil!g and HII" O,(ll~ T ~tln~llllthe' hard (' m; is) ftcltl-dCillCIKk:nt 'i:<;('JltllC!1lllOlI rt"sfu;~,h'it)' r~11 IlaH dI)l.1I.'!! ~f Qrocr~I:'CI!m~\C~cr ~]lI!ctlimifl<~~, JhC' 0<1!(mC<li;<;'tl!~d. itll Hid .. w II<iJ I ;;;;QC nicienl was SLllllY.reSSCS ahe If<!m;,i~i(Ul 'iC~~IH;:~!\~re and aocc~sc:." 1,1~ Q('f) 021. The "biHt)' IQ rU~IC' dG;cr,i,bed UJ) by 1m !.:Dnpiriea~ C;I{I~hlH'~r Moe of qlll<Lmtlnl crilJ,c;J:UtYC1l~l~~cl'Clorc bc'~xpli:rilll.CU'~1iI)' dtl.t:id~H ..d by d(.'~I.'fiIll~1] ' iDlgI,\1'h~hCi! ;i>~llgk: or liiiLllti,pk ~Dh:r!i.v :;;cnk';S, V~l]i:ih m, 'the IFmg.iIl. oM., g9~lil("li@l~ d(!P~I1I~I1Ii1(I~of QCP ill 1I\".."IC~INI. oil YIb!Flli!iS;i~. The< We cou5id~r U~(:· hoc...... . y.FL:'rlillionl nlc~,a~ the PIlagrietoS'trkli:Qn sYi'll'ioots IrepreSfl1tll1e Llflear ('{J;fll~icDfnt YbRh~S]<! (YR:s. )1md siww ~haLIlllilliplc (."'illcr!!y ;:>(.':al",";); v~ni$h liS iils Qcr iii; "pproac:h(:d and,. i,n A~U(ll '" c; [FIt trW h\l~e~e- t is title 5<lfiI1p~e.lIddli,l ~(Jnl. ~Ulgg(,,;:;,~hllH;rll U"'1J1 t."C11[{mic111'100,0\;',<; [Milith ~kll1g~@ ~;UO] directil)lil lIIithijl:J, ~ d the'letfagonill ,Gb p!iillll!i!)' verlilJiS, 11 at i!::oc,,,,ii>~wil.h lh!.: :;.kJW IniulC'MlliQm (]It lli.;; l11ag· varioucl; t,emper,a.tul'es.. lMo'lt tna!t A~DD.1<;0 T~K.I 'Illetic OrOOf p!ll:r3nlcl~j". A diD'l:."Cl way 10 pHlh..: all'ld tilla,t ttl e data~el'!i hai;ie beelil ~hiihed --O_~2 the- ~IIIHii1ll5k '..:nil.~ry:y 5t:3~!::;s, in (1:111: C<llliihh.riulll I:i '0,,1:3 by ,jin~r!lll:!l (im,ount::; "'~rti~;ally., Th~ o 02 sh.ar]1l Jiiil.ru,ilre 11"1the 0.0.2 t{ datil ~OJ!M;n: mtandi: ~nstll:ml! fll}1 (JllI!!rniral PJl1j'ti~ IlIIs:did!., ID.oIM7 .0 (lea: [~SP().I1i~~ to iii tli~[)Dfl!bf]JiliWin ), (as is: H.lLc . '1)1<!DIl<, G~iiil'Iall1l'. ~Centeilhllrfflaoorn~i1:l11l1~1)',.l)epa~i'lil!l1t '" 10.5 m®re df'alrl.y Slefl'l ~,n,rile meiMtllfe'~ 1flil9lh 01Pt~·f!:. ~IHi Astromool!Ji'. ~t~~ ILjI'l[loIerslt)'. Pis.t:~.:!wa.~, rilJ iOBilSS" USA, ~Doepinlilflemt Qf Pl\~and A:lllrlllooil'l~. Rioo !,I'fl~~~~ #, ""hlll;liJ ~h:Q!~'Itlii dl~ ~ 1lI9~i !"!~lQP~ 'Ul'lh!.'ll\l. Hoo!i,IDrn, T:X. 7:i'00,§;., 1..ls:A. Ilu~does Mt ~O:ntafil a Fly t1is(;o'Il'ti nIJ1Ity), OJ,O 1,00 '''0 wjlOiil ("(!I~s.pomde<moe<~~~ulld ~ adii~~~d, E·!'I1~ik dl~IllIOIl~tr'atil'lg, th~ WlltilnIU{jil.l~ "i'ltur·~ 1P~~@rn'I@,~g"oo ~J";GJ> ,qm~j@ri{t.td:~ ~OI;~,~ of til e ma,gnetiJ:: U arM,itimi at tlile (rit~
I

Til ~s l'e r..~~~ll~~yc~~n<.idlC;:1" ..~h';arlt;J.ge of

~11f'''"

MtiriDlJ~~h.';lrhcnnadyu,[II!!fIic l~ro[lleL'li.~ in YRS. l'!ll;!cal.~~e ~hcy call 1)Cl cau1f\oamd \\'W~ ~h~irll:!Jl1'S:cfMc~lil!l' I"'!m.nd.c..,,. ~he ~oon.~ maucr ~lllKIcrgo'illg ,I
~pt;l>C!fUln~lcar a

u.mn.lI1l

:f]LIC~U['ljtiOIl:-1 of

or

um.

un

ne

t'ilromRI ~l1d'N1~S: f~ft fItil~(S IIl~itUC~ l!InlWil'si;ty of Go~JingeD,~?On G!M!mrngern, 'G~lifIlilll,)",. :t~ro~l'nl ~!kI~Io!:: bJ,s ,I!liili'i!i~ lr4liuoo.al LaIHlf.a~DI)'" ~.O~ ~lilno~, Nh'I8754')., US1\, -§fl~e5l!int addrffi: ~!l~;M~ of '5did .s.'ti!~.I!hf~~5, Viermlil' UJlilfe~iit:l"(!of Telhnl!llO!l!l',10~Oll'iemrn. A!lI'IIJi::!.

j;~,jJl fie~d af ij,O's.f. S]mi~olf bel'laJ",ilor liS. o,hse~ved at. villn QIUiI: dHfe re·n~ t.emIlElratrur1!!:5. below f),.I)7'S K, r,OI" ,exam pile, at. 0.00,3: II( a ild: 0,(15 K. Till!! so' lid ~iilli,esfoil' T ~ 0.13 KlHe fit~ Usill1l,g dM2'inte,gra l ,o~ tlhe tfO!S£Q'II'er flu F1(:ti,ol'l J(H, "{), wl1ii(ih
1"e1l>eaJl. a cnar~(t~rlstl(
rield~cale

Ho(n a lOll 9 \\IhiietiJ the' mill:g Fl~IIj)s~ri{tIDfll~,nlnlj5

.iii iii f,ilIr5:l.l

c: (l1al1Jge

ill1slope.

969

REPORTS
filll.2. ~n;ergy 5C,a!,es ~!l'r(ilRh~Sil< de~ermlle(j trlltllil tli1emMrrulll1l~(, m~nt:s, fA) 1i1Ii~ ~idd d'@p!!nd6il~~ (!I..ld ,(l'fth:el!Il~gnel~tiol'l :l!.,&1;l.O].
.M ;;;.M ma'lllillfth:;, ,a rild trrall~pOrl measure·

.1'.'---- ---

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a nrl
~fQT

-I- "J.,.N (ll,1hl~re X

the btt~.

tntl

Hait fesistWity PIH for Mlc


~1iI~1i~
\f~~~l;!~~~

"" i3M1fJIJ,

t!l(:t{Jir

be'Em dil{jd~d b~ Iti e <!lnri!!:{I~ro,PY


13.2, wll~(h ,,()mresp,gilli>

th.e fEillio of Ule (rii~1(a1 fie\lcIs, ~n the

~o

I()

HiliU !\Il'Idmillgl~kIst~1;~Qn !:l!l~eiS~remenl$),. respectively .a llat. r"", 0.'5, II{; :sil1lli!<irr IJehaviGF is obs:erved ,at o~Jler 'lPmperatliIl:S., For j'ln(ilI(J!Ld, tliie 'Xlrn~~' '~s,(I res.ldll:jill: ~51iM,ty Pc= tQ"S Iln em <llld Hr '" 0.05 T;for hi '111-('), the $iliillP~ has r.lO~ 1.0 Ji!l (]Jll ,a.n,d H~ '" 'Q,06 T;:and (or I1JHU c).. the s.ampl.eIhaS. P'Il ~ '1.0 ~! em amij H~ "" O.6~l lltesi)'li:tiI ~!1.es cOfliesp®nd to 1113: s.lflg the' ~i'I'Iegtilil O'~. tllle~ m.e (fo~oo,ve~ U

H(T) timf<lf <lIS M a'l high lie1~ S;2').~IB)The I/J~,ro~r riet()s!I;allil' NJiJo" oolcemr>i'di a bom l1lI~n:etl;l~tr~['ti!Jn, M, ind HaU r~istM'ly rJj~iif!gtilE! ~m~ ~mOOll:s in A...It ~~ 'is e(J,1J\Y~leInttto the eft€l'{iiY s.:aleir'ffli). The!' gray tiiamood; and~ia n!l1es f@'rJlI'ewn,t, 1'(!~IJ@C:tiJro~~~tfb~ N:.eel o.rd~ril1l~ ~empllJ mllro (TN) (!l1d thll! (rli)5WV@[ telillperat!ljn~ {Tur0 .b~t(!lN \>ffiidlM'le etectrh;at resistWilY has !he: IFermi UCllli~ rQ:rm p = Po+ Ar.4, ~~ dl;l~rm~IiIOO1~r'~!iD nn~~~~nn~t~ -~ri! ~ ~illlg~ ·i;I~M~.1jh, Pel- 115 ~LQ (1lI1: nc N~ '" onl) 'T. 'TIiII"!'soiKlI a.ntll ~etl:e(llin,es. are 9i;.1i~ tD fill"! a leye; lorllh:e ~tl:lif, d1a~ IIXl~nlsou!sW~~h;~ plotted f~eld regime ffij'lfe b~J11 Ll5ei1l, 1ftle hon1:ZOf'lta~ fmilr be FS Ir~i'Mefit tl'i,e 'fItting e\rror ~a~lLfr !han t~e' widta11 ,cdth~ cr~~'!IElir.

m~.

IUlnctij(m/(H.n " E~dl dati! ~~thij,. b!l'@!1Illorm\'!Ii~.@'d1 b:~itsirIiIiiiaI :S~lJf)e ..For d~riity, the three da~a ~li1 hi3lflie ~e1ll s3111't'OO dlmerent arrKIilliillS '¥ertJ1caH~: Tms is iby rE!'pt'E!'Se'lItetl' by tile ilihireesejpell<lle :Niro ma:rb[!1~n~ th~ WR'i~l~s flJr thelhree quanl'iti!l!;s, P",-Iltt""" \me.~e' P:H ~~ tllle an:{Jma.1.orw: Hall fE!s1SIWi:ty {H), MIlia"'E!~ :simi!awlyro Pill.We a!lil!l. )~,M!~d!is the' field de~lt;.@ or the ml~!!,~( rfee ellleirllY GOiil~ributio:rI hM x .f/); iiuing ,M{H) .loead~,Il), :simMr ~c;l1dmions, .althm.~h the qlll<llily 01 the fit is~om~at I!i'i)O~lilr be~~lls'lil wr~1j; H i~ as MI fll)\

aoo

.... I l· hj,l[ Ihe cmssovcr ji~~d:;crl~"; l/.f'JO) is O(;qLliM.liCIlI'L ~o tll~ ";rltl:~y seale ;r"'(m. We h!wC' a.!'I:llyzei;l Ih~~ rll~.!;!iI1C'lm'iitriCliCln ~lat0. a~wel I as the ex iSI i,ng Hl;QgnlCI kuu VOll d;;t~,~:f J,t.,) (15. 16). U w.ith the smne crossover iflll net len. No i()OI.l~

1'it!lu;aiuIH.I

{fflll(!<)"

1:

rU~I~ forlll}tR. n = A ~- (.... h

IF~g.3. p

lo.flgn~u~ililailm~"

ll,et~mi!s.lsti'l'ilycd'l(bRihc,;Si~as;

::;,po.n(~inli: "1I{MNi~~ks; cal] be resoh'cd ~nlh~ IIIl~:tilDlC! i7.O)t'i.t'!! d<ll:(ll rorff c' (1fl'J, w.hich iL~ 'Jlnl0!!i~llin~ .. r !i'I H Th~ Sll1!iid ~~~rv~ ~f1 :r~g.

land ri~.1\ efl!ffl.·s,pO'!dl(l H!s,of j("lllill' J\:' 1 M I XH aed .11'C<II<l!JI re~i~'~\'ity PI·!. F'ig.ll!IC 28 ~h('lw,~ the Ihroo ~1$ Cif !:1m( n obtained fOOl'll
such fll~, Their oveu~apD'C'pre.~cjW;; a kc}/ emlcfuslon of the pree~tlt \Vfl~: it s,~~_~ts lhal

thc)! ddhlc one

""(11)1, Thisscale i~ seen W be dil"t:iUl~1 Ilurn ci~.HI~r~hc ~ri!m;ii~I,Qu 'tcnlpcTa'.un:: tTN), for I.he1l1llgDl,.;tic Qnl:~~illlg1 3 1( -c H~ or the ~"iJ~e (TI~m_) flu Ih~' 'i!:" ~llbl~hJrl";IU L~h: Lmltlaur Fermi li(il1nd !>I.:.n..: ::J I. H:;. U~.,for ;lIlllhro!.!" q!JUlllllli'ti!.!"i>. Ih!.!"widl.n
Cf11C[gy

seale

dilSlllay,s the phase dii:!!!Iram~ iI~here the gr.w shaded .a rea [~pr~~t~ th~' ran!)!!: ()f 11\0(1.1 VI:'! ~!1Ie.~~.ho~~n ~f11 F~~, 211. Open ye~krw tri~lll.gl'e~ mark tllle PMltiOI'lS ~f tD1e hdl.e,,-

\l@r:s~5, H (iUc) ~t lI~ri'QQS tfimpetT<lrure5. Tnemaxima . , th~ 0,0) and O~()<7IK. d~~ il'ldit:<!~e Ule lboo~.nujlaijl'Oir the Af ordered stolle ~Jtoi'tH'" O~'" 0.005 Kli. The'i1Irmm mark trn>e "[)5ilii(lll~ (if il'll1lec1fiQIiI il'O,il'lu ln p{M, liM ill~!. A

E r.o a
.~

c:

--'0.2

~O,31

1{II'Q

tioo p()illllsin t~~l l(l!n:gi~~idfv-

or

uh..:

crr'UI>IY!JV,ef ,c.",ml~O~"'h";lO

10

zero

at

r = o.

or

illllp~:yil].!l tha ~U~,,;d iflcrclt~.i~llli> Ih~ u'mgll(1W' or ;S1:~icli(}ul. nmg;ncdz~liolJ, Uilld. I.~:~,n ro;;~::;tl\'ily IIIII\'(' u jump in lilw zcro·I."::I1~Il<;[lIhJn; I~mrill <~llppol1iing anI illcu,"~1'lt ,. The r'i.:;;;lIl1:s mis.t' 111"; illl1pCirl1LlllL CJiL~,~ti{ln '~hlll t-mll,,~LI 1t:.1~~1t!.:in '~1lW":i.mIJu: IJil!..1l1l11ooynllmlie ::IIld ck:ClmtlK: lr.Jll::>p,url prnpclli ... Om:: nnil~hl 'll. a~tle (i7) ~h;n !hc 11~'II-C'nCCI evoh!llioo ,ms 11 llanc!ia.!l of tlle" fll[.!g!l!Cii.c Ikll:l (/J) is· caused by

Illlarl el~(~r&(a~ FeSllSlillity. 'Th@ 0.5 -------smearM kiilk behi!\I1ierilil~e 0,;8. (},o4 0.0 O.TI 0.,:;1 j,S(JUiefmal m iiei'WS H ~or& ~~spd)l!ld:li~o ~ p~k ill'll lhQ' '1 d~~lI1d~iI1(~ ~f.x. ih,~ 1000tt'~rIlill~ 'OO~1Ii obS!~w t?Ot th@' (Q[I~jl4Jiildin!J1 ~alt t'~mp@rat~r~ V!?iSUl~ (orr <II H sallilp~e witih Po ~ O',S ~!!.C:1IIl and! U~ ~ 0.05 T i~ dis'p'layed lJ,.y the dillrk blue dia!l1lQrlds and! fioollid w be (an5,~tei1t IWIlth Ho(T).llilll5e't HI· J~p~ar-: the- dern'ltat"Mf dpMH Ve"IW511 at !rotlll "1"", ,ea K snd r -: 0.3 lit ,i!I!iU"(I!,!I$ d mark thll! mirlima... oo.r!'@sponding tllh i!lJileclliionl ~oints illl pWl

'--0.07 ~ (1.,013,

-O·~.I

Hm

or

l3Itl~)'. Mo:t~m'elr; allo.U1g 1.~ll~ ~x:i;.;,iJv~m "",LU~h:l, ,t'

'[~~ml:lil1mtUl'(.'iS. Th,~bmaocrwtl

~I.~pj~kx~ 'L:k'"Cr'l."Il!.~·.
c[lm"5pc!lfId.. [{J Ille

~hc ZCCllb~l srJJ~!ing (If '!he I?etnli $~111aoe i!led~~CI by ~h~ m.:.g!)ct~ro.~i~l {pm! ~nccil!c,~ !!1 the m i!l.gne~O$hicion~. IIQw'c\'c-.r. till:: n!lllgnei.~~i:(ln i 'ilI1.1I)' diilI,,1 iil_Yi:. !l ~neal"cd k~l1k. and ~hc CM'C$potldi~.,;;l Pcnni sllwr,!'oc'c:h:mgc wf.l.'I.!lr:ial,IIlO-SI prodilCC a S!lleilrod k hlk in thr; CVfllLI~ion n.f thc Uilll cocmck:-m; such a kink is loO W'c.JJ,:,oonHP<!it'!OO wjlh Ihe $~nCJlICd jump
S~'ii~

iSl1IIc:m::dkillk I:(.~um:: lis :iln;c'Ht in I.hlil m ....!~IML~. 7..lti(lI] \'CUSliS Ihe m;~g;I'!C!tc Ilcld. rIlS!C~~ it is Hwre mmjra~ 10 \'kw ~lC nOJli)fm~)'tidtks ill bOlh I~~ nlil,glloC'ZY.'>1:rk:lk~~la!ld ma,gncli1.<!1L:il)ll ~ the!"!lllo(~]'.MU!1i.c 1l1<l!!lli,I~Il!lof1s of the ~ ..rgtc

oba.cP,iJd
CmSS10',lCr

31.~:II IClllPt:rdlurcll.,

obsC'rvcd

in Ihe other

pmpc!1ks_

c'\p;crim!.'1t-

fcml i ~uufa>c~jump c-;:m:r.edhI' 311)~kctl\,n local~lion. 1(1 ~.xplo:rc'~h ils ~S:s\LClimher, \'!rIC Ilave ,~I00 $~f!.ldkd .he ~o!lgitll!di,na! l1I'11l,gJ!cooresis!ivity. ]=i.l;ltlfe :; sh ows the: d!i!ClfkoilJ n.'$k~Dhi~I)I fI ;],.'> ,;). nltlctioll Oif the tmgnclic fkld ~&,~ vmiou:'!, m
2007 VOL 315 SClIIENIOE

1~1Ideed.~s show!') in ~!~1'CA~ lhc Cl'l)S."OVcr' A. lie!.r:!!. dlClCl"!Illi ~1 fro:m the mi nim>L (l f thc n (Ieri"'<1~i,ve "f'Jdl-! t h~Qil U} J~~I (ilfl the $i!fllC T"'{in linoc d'e~cl'mined from ~hc m~~1.e~O$lricci0n. njl[lg!~C1i"-<l!iM,and lIall cllcct. ~11 additiN~. itl.'W1 !Jil1h('lI~\fS~h~t Ih~ ~\~dtl'l (If dIe CI:O$llO\'CI" deClt~aS'C~ Ill'll! U;:ofllpcf.uU'C'i ~ fowCl"C'd, A ~ dcu.ile!() a!lIJlysis :;;~!O\'\/S lilJI the cro~sm'cl" withh go~s ~o zero in ilhe ZCro-l~ililP'CmHII'C

9'10

16 FEBRUARY

\I'V'I;VI;V.sdencemag.orgl

REFORTS
cri •."~.1 reg~me ill), ni,ru, because ferroi magm::lic spin ilI.!CllI[uiuntii ~rr,; ii.'lc:nj,C'i.c~lt in 1:1~ f
fc.miug el,HliJ:'g>C.' hYU15pon. [his pklure oomJmJi.c~:1. Oh::'ii:]:'\Nlill(!rrl ,o:i"anlcurl:y #~iflrlcpcndcm nu.io Atr: ~hm~wc:olllj:lf~rrli~,'!.<! !mUifl,gl}' 'ff"de~crtdcm .Amtcl ./1: (10). Here, A is Ihe ooenldt'n~ or lhc
dcstrueticn of K:oudo !:;rllmil!lI'cmci'll.ll1 the fOm.10,f.l()Cfl~ qU~UiD~l!m cdlie.[l~ilY U. ,1l'J, a col· !liP,,"E! (D (a iarge i"cn1!li surface ,1$ If deereases kads to anf~d(jc(!l CiJlcrgy ~cillc ChUI1tClOCtil:il1(!;l

the

au elecnonic :-!Qwing down 1m.:!. i:rl~llcadilkli'l • .:l"ickl~ Do zero-temperatuee jump in the .!·hll!~ r2; cOIlfIpo.nC'nt or tirl('! n.";S,i~liv.itJ'. c~fI1.cj~nl andju the lick! dil1lilDJlll.i~~~ or 'he ~IHJnl!liooymu1Ii.c qllUllt.i.lioK An addi~iomd vanTI'IC d~~;)J IlITSCl!h::ld .Irlt'lru :s1Imwnhm. the ~mi· Kh~llg (;I1Crg}' rS.c:II~ a lID c ....ist~ul] tl]e "dl\Conrnrm m;);g:nmiz:lIlon ~~I= 0) depends .oil IIXl' same FIg .. 4. S'ki~'lLh ~hieslIigge.stf'li q d~~emliefil[e of of fIned" qumlllim critical it}' scenarln rm'.i mmlm 1:11£1 mKkrlyi ng pby8ks ail Ihm. 1O:r~il~ chm-ge H'<liI1S· lh~ 'W,e~sst~mp~rat!Jw~ E}q. I;'I'hijd~ ,ell!lel}<~tw qll~nwm 1n0000lt1::11;. aswell a~ i:Jl il:' c1{:t(,~nSrDn ~9). port, Because theM;;)all~p!{)I1. is dml]~rmt~ hy m(lgln~tik !lUS<OO ~ttibilill)'" A1s H r~iJchl?!o #", i,t systel11S (17,. l:M}~hal 1111:';: vanisihes at thEl'lIlntiih!rli®rn!3AJIiUHi~vave '!I€~lo:r'f;!, Ilarg;C!llI i1~~ctUiI~ lens, Ille I'CS~I,h~ imp~y limt d1:!~ [1J! i(i:mN<'IfI!l ·~Ibcm~);m ~ af!ll,le.d Ic} be .rel~\·.:Iim w ql!l:mtl!ll!l crJ[k~1 hemv), q = (! n1<1gneli~ .t11!1~1[lmiolls are <B. part of .% sl1o'i'l'n. ~n .a,mditi():n~€Ifjlll1as iii s.emrld mlnimum <It 'I!II "" 0'. FerJ(I!"Ill'l1gnelij.,; ~lu!t~!J(ltj@lll~ <It ''II '" 0' overall 11uet 1!I[l1 uClns in an e'>:.te'~1d,1l'..d nmgC' o.U' reFn~ioll 1Il1J;:n1Ik W,O:\"~A'CCUU scales. II. i~~hcn mi1:PfC 1~~1iILr.;;ll] 1<J1 ~~mail[]ilm~r~~rt ~IS; '8,q • 'Il' 9Q~S' to~~r'Q·. assume tha:tlhc dYIKunicn:i spln !l,uSC(lW!tibHily R.e~·~ire>liiJ:e5and! NlIlIl:e5. dim::rl(ln~ \\ ave vectors ohcy~ !he- S<J;lllC tbml .11.. t,t ,(i,Wiiisnn. 1M... M'od, J'~, ~!l, ~!B fll~1I.3). HII]!I {~II)po(l.1.ing nnline ~exl), nmplying a jump. (8 .. J5l' ,liS, Ihal. oh!!lcr\f~1 ln ,1nlolh~r pml.Ol)'ll· :ii•. ~ 14. Mem, PJJj!j. ~, 8 li,oi, 1'105 (i??6l. leal 'lU[lJIItUlIllii crihic~Li heavy (coulon rneml, in, It!he if'CS;(]~nllre• h<i~y~l"ros:;. Ihc nll.l~I't{:1ic 3 •. ". I. MIllL'.;,. .!?1lYJ.~t'v. 8 .~!t7 193 U~93l. QCr. This is ill accordaace wilh I~'ICl.h,c"Qn;tiCil~ C.;{_"lII;,,/WO_1 V 7}: X~q"T!~' - r0~~~ n ll'inr~ 4. To MoiWA t Talo;i!!11o!~ J.. ~Jjys. ~. jpI!. '601, if60 . (US'S:). AI U:~ QCf.,. I~tc ,1\rCli>.i li~kI ill th,,; al1.t~:f~J;WJ'fmgu~'Dec'dlirnl,$ ~ IN • .19), UNlllC:i1UclXi Willil au /: S. W. O. MJlIjI,jr tel Q£. iVo;l~ 3·~" ~ (i99St ~lcc.rm~ ~'iJc:jlil1j.:i('J] tlilli:::il,iolll, netic W3\'~ veeier (~I Q) v~lii~Ii)i;~~ 0Q = 0, AL = 6, ~ S;iehrilell,Qil'mrmm ~.~ f~!irjtmS '~Gant!ri~ !JJni.lI'_ fi!l\ll Ie 3. i~o;;:~ A. also shows the.' 1Clllifll;.'1I'dJt!!lrc the ~~I1~C tili~o.!. ~utl IlIr1J'iike [or l\.-Cl~5.gAlIIo..!. I'm,\ (am.ridge. 1'1l9~)L 7- p_ (D!.emaB, c. ~pill, 1:1_ Sl R. R2ma~slJv.jli, J,. My," sc,lI,,; ~J~ a fUJl-C1on of ric~d. M!tllC!.cd fitml tl'l~ e,~i"'O is very .:;;nl~.Ului YRS. B~:scd tUi the i C (Al\II'd, Moll, sa, R',l2~~ '[2(IQU. peak iill [lit:' l;dcpcnuJ..:]1!cc 0 f~;lc di,[i;:reruial SIt;;. :slll!JIIlU iOIl $CUle M':Cn ill Ule Icuilipcrmillrc dea Q_ Si, '5. 1lliI1;II!'lo, IlL h'!len2l!1if~ ~_L5m~iII, HlJtme .:113, pcndcnec oll" !lIC ullTliforrrl1 mfl8U1CIIc suscepccunibil'lly X = ti!Wi?U: [llC hml.Y. ob:>a-\''Od 6@1l, ~:w~, IC'adklt (2fT). IIC(~;;$al'ily .jJ.~xxl\lllipui,;;s U~C ~itJ.iI uly {J()}Hlld 1.1ilcNMJ)l;. K:lll,gill s.hi (~ dllt;iJ, '1. r; S!!nmiL A.'rli!;i'iw~n~!Il'I, t 18~l't!nl!, ).. Sad1~., PII, p, A. Fi$iler:. 5dtMC JIU. H91!1 t2~'1~. (2 n..:m- I-ir. wo,; ,..;:;t~~ntiJ~,.; • oto be ma~ho,; 0(1 :;1l.l;;dn.i11illlkiJclm ...Lor in Ill,..; ~iUlh";nrJllIl 1.ll verm o, i[i'(ji;i'i,r~~i'(\!'.a,I,. rey~\,R~, .~rr.115, ~lr'l flOO!Jl. onkrol:'O.3 K (211~.\l!,,rh;';·ITI.!lII~ID'>"I.:" tl~iay rrull.l ;:;Ul5 1-1. It i;s i:k~urly M1Ciil that this ,:;.c[urc 100 1~]lhou 111. t!:. I$lii~~' tr ~l.,f1tr!, 1ililV. .!1 6~" UoMOi 1Z0iJ3;~. cul.l1C·~0 or Q. ,Sll illcrci~~..S 10 lite !Ili:tl!cl' ofUilc ' the same T*~Ii} line, 12. ~. G!!!!jmwi!lii: ~ mt. M~ ,1li!V:, ~It. 8'.1', 0564112 YQ~.id'l (R KKY lin· Ollir fC-~ull:il slu;d ~ighl flU Ih~ n.....~I~ phOls~ .Rl!,ldl;lrrna~~Kiuc~~K<l"llI.y.a.~ , ~~00.2}, di~gl"anl of !hi~ dc,1]] :;;tI:lir:hirn]~c!ri!cllmMl,!1'1l ~~rat[j,rl!~ OJ:' bare Kondo ~cal~ [ab(l'lit ~5 K ror q [3. ~ flll&(I)ffi el' .0./,. Na:ll.fft 43:t S:S:1lG:oo.o;}., l4. as, C!li!J1dr&;eil.!lIiIr:, e:, f~..... ArJv.. Pfl~, ~. ns: ~t1. crhlca I! matt:: rial. Nude.;]!' magnlClit;; l'CSon;mcc YRS ~IO)~, .~!l. H'I~JSlrnllCd :inl Fig_ 4_ The en· til.~i]J, (>JMRJ m~!;;U~CiInOlll.~ {1 while Sl,!l;M~iltjgthe haneed uniform Hl,l_gill.ctk ~u.~ccptih:ili!y" the1'5. y• r:~V!il el 11.'" Mf,1. ke.t,~lf. '9'1. ~2~4012 aOOSJ. cO!lcGl'Ilitam ,C'nh(lncC'~,iWH~m~ ratio ,~un" aud .. dQl'll ~1t.ll~Cr: or AI~ ~hlCl U.)Iioll~ ill Dh~ q~lalil~!'l]ll liJl, P. ~!J~~!1.. t Td;;:Jw~1 (i,m~r,l, !C. Geib~!, F. S't\ll91"=1\ J. cri!~k~.I,W~i~l~, have ~.~f:{l n;\'~a~cd ~11haf1Nd ~he _~jnal~g ~ ~/T~TK2 11awrolly ro~lflw tq.0'~11. I,· ~~,~(lc,Ipi'!, :;!'~ (!;~pp1J, 155 {~~}. 17 _ It 't. [i:lmeyll!ll. A. Rcr;rlh, Me V~j,iJ, f: W[l,'!~e·. ~c,j'r.rHl1ag!le~ i~ u!;lu~ljn~li:1. iKPfringll l:;'i~iD. th is pkllU:C. MO~Clo'vCJ;. both :t~o .md /::qaQ h1I~i.I.an:i'I'_ofl]!abdOlll'ld,mati06Qf6H7 {2:OO:~l'5: ~trirJ.{J-.is, S~l1L1I~~ OJ'l !he ,ordc-r (if o.t~'o, !H::.~l,C' sinlB<lrly \\f~'.~! md !h~ ol~cr'I"~~(¥H 11..• 1Jl:_ P_ (DI.eman, JI, It i!.Ialrl:i:DIl, ,!\. ], Sclillllilild,. flrw. fie'<' oil l1-i ilt \\'he~~ Sa L~the carl'e$l]rnldinJ;l I""io rfil' U()f!r ~h<lt•.11;.::2 i~m~:<Ld~' 1,.del~C'udcl'~ i~ r,')C1Oi 72. 20*$1.1.1 t20OS}mal!} ifcr.!ation (I! r an 1'1- ind(!'W'c~HlcmIl'Ilt~, A~ I 19_ Q_ 5i.. S. IlliIbe'lo, It 1"'!I~~nt ~" l. :Sm~h, PfI~. R,eK B' i!li~cracling 'C'~cotm~~. Furlhe-!" c, ~dlC!1ce ror lOB, 11~1.(13 fZaO.n, Ihis: le<lds 10 the ,r:;o)lc'lLlsion th~~lhC' originl oCll~)Ilccd I'ctTfi'l'lQOl,gIlCl.k fh,ilCll!laln,Q~" h~~ Conine .20. F. Ge,!JB~i'I.. j, ClJslm, ,,;,T~iw"" (, Gt;jb~~~. St~,'d" (lfu~~c TO; !nnC'lies in an ckc![)(l~~k s,lowin,:;: n'Ol!l ma,gl!le~i'la~ion !V!!X.llSIll!t".!'nCJ1I1S l!(J}, 1l1e 1l'i,s. R1!y, ~1!I€,5!4, 016'1(!O2 {2.oo5~. Wil:;on HIti:O-Rw = i'lA:il~/hl~~~~ :<: "I/f. w:illh Uil1'w.l'I mall. ror YR.S, th..: ::>Inmg q = Q rhhC'llilll.2:j[, I!;, I~m~~,('1 ~.r.,~, 11,~ ,!.til. Pi~" :~~nol)lOlOO2h, ( tiom hlQP~cli liD b~ II cun...equ";l1co;; 0[' Rh...: Ilml.er ~I~rr= ~.4 J.I:E!lYb tl!) lji S!lro]l~lly~lTIlfnmc(,.i1 fiJt ll .. (!II~~~ tl itI.. i!l~t[l,f~4;Z4. ~<lo1 (.2003). ~ il:>W'l.::H, .2:3. l Zl1~" M. (;~~~ A, JiDlcl\J.. !(~,'5~ fII'j~; .~, Utl.~, all c1'<lcm..lccl Ut:2i:mll {If liic pl'lll5!C di~ltmm.~l j:; 06~<il~~ ~~OO3), \i\,'cnm", l!JImto f~lUr{~ dl:it~i:loo l~i~orelic:~1 ahl."IwJy I~llr~\: 20, fOT Ilil~nl.'t1.ic: nekL .. Urll, :I;';w h ~. ~, f:iiicMet« 11.'," P/l)!;i. itev, .tell; 9ii, ~6640S: (;200.3)·. '[e.:.!n~ alld fLln'l:ler irlen::!N~~ t:t" l~1C 11L:~.d~5, illnIJlic{ui~1'l.'> ,of oorii1t::>IIIII5. OUrillCtlSlIIrelntami, ~'5•. ~S<lhfO:ler ~f.tI!.. Nahiff 4i!lif, 3i5,l UOO~" l : .~'tluc:·,""dI[I]IWllru JJ~, lhl:rulim:, ~. could ~,.; l.....::Ibli:;,h D1mi fl:Ri ...nengy ~J:c T* i.~a.~i~11J.:d ~Iii. .M!!!'e'5pe(i6i;il~~,iI1e d.o!.<! oll!:rtl1~ ~e~.atLf'e' ra!19e' ~f withth..: (XJll i,1~briHIJI Il~!IU1y.oody :.pc(;uum '~t:mp'lill,g w eoniiitkr lhe q - 0 al!m!'lnclic 0,3 ~ 0( ,.~ :l!Ci K .~·ne !ill!"!h\~!ml;i .. tU i!I1deq..~ 'II 0. \\fIeftiI~ IOOse!Cf r Q,3IK Ib'j (J .. :i,.m.a ®-q~. e,3 g,; QJ}. l]iUl::I'lIaILUIl .ms til,.; clmnimml (TiliGi~1 n,liIl)t!lltlJti(J1J (wh.ic~a ,ml(m~ d{.;l~nllilrl~~ 1.~lcllmHJymlll~i( ...', l~_ T. >milfuil M. ~~~<\~, Si!Ji:hdl!v,. p~ Ii\!JI'.. B .. ~. Q~'51n {In, ,"';S«}l,'Ci;~II'y b(X:!HI~"; <1J t!ofHlWlIlli.mmi ,~rro- Mm·covcr •. Lhi.-; :;I;;'uli.' .i~ di~.;illlci rrum th,.; W.mttllLl ~~il!D4~" 'unm,gncli,C' Q(']~;wolll~cl yield a fil:lunc·ls..::n f!Jnn:i li:qjl~icl~k. :i'ir.l~hcc-.. :;c p~y:;.ica~'gualJ.· m l&.r $l!n~m~S- Sadllilel', M. lJol~a\ .~)'.iro.8 .3'5,'9-·~&11, 9 !(':!('I:lIOnclllt (2'3) it'"'\/' = 2/3., dOM:'to wlii;rt L<;, til.il'll rnimli bl. mIller di 1I1~J1tooha ...i:ar 3t'fW.<; ~~~5)" i[jml;!'rved in ¥RS lU), TIl i lJlicl!JI~ i~ problb ... ~h0 two !;C!ult~ {tmpponin!,l anlliluo,; !f:xt), F:i.n~J.l.y. 1.9_ ·W~wooldl ~lt@~'iII~nik IP. '(;II'~man, ~_ (IHlief1!,. 1~ Fi!5~, ~ flli~emanB.. K. llsllilla.. 5, ~i~m~r; i}_ Nl!o1eJs;oo. l.mlb of Iht~~ 5CUI!J:; v:mi£ll;]J111.ll:: (lOr" These lillmti.c for a. mllnbl'T o:Y 1l.1:1."rm:;. ~.tO".\'~vcr,fiI.];I .• !VOf3{Ii1I!1'. ~~ .5(, Wii!ln fo~ us~fuL~il'lll!&1iJm. 'I1'Ii! wD.~ neitheli" tll~c-.dimCll~si(mal (3D) no)' 2[)! 1CrmrillJdill~~ con!raoict ~hc convcntinmlil o!<.krb~~ li<l:m' '~ifI~ l~ ·ih~F~iKI~·d!ar fi~misdief! IKLronlctcr ~1IJlC1JUJH'OllhcrllY iu m lea~ tWQ ! m~:!J!.::~ic jlpin l111C!ll<1ti~~1SC5!lt get1crn~e the Illd!.il~1fi', ~iSF gr~i'll 11lII\1!·04,2:'112:5" 1M RWrl. .:li'ld A. W~ f.Ci<Ul'ldaAi.oo. fmci ['(111)11 C:li.()fI1liC!llo ob:;er\~ un I ~lc~Cnll[lCll('l4lDrc I:'(spe:c~, FJ!1'I .. iJle OIlly I:ow-euei':gy scatc un d1<ll~ d~~!KIe!l!Ci~ 'L1f .he- \.i!li Wo.rtn spin ~~IS~P! ibi I,~IY theory ii), as$QI;!a!od Wi~l I~' !1lIDg!lctk s:~m'il!Flg ~2m" S~ooOO. ICfmf1l<lgllClk Spl!l n~IC'!lI!atlrul,~ (~(l.wnwhidl. fm· H:> He. isT.:'i7L (2 5). Second, SUJliPQrtingl O,nti'n~ l!ltatmlill l",i'Me [en(mt.i!lg..cIlg/(!liltlllJt~[ItifuIlf31~5J~aUW.6.9.,illI[1l WCi!i.itd lead to a d~vCf'~cmlt7'~ f-Ull!. wilh ;\' w~lhifl Iha~ II}Cory.a :!l~'mp fcmullCiln thcmtoay~13and ~(2. FOf 3[)' mul 21) C<li.~cs.. !\.~I'lectivcly} IKr.mks rnlld U<l!lf,fl0l1 qluamijli~, m~g.llt Jri!lC m~'.[lr OORlfM ~I£l!f. Silllnd SO! TN {I:Il~Y, thm i~,n OOTllm"! I.rHhe oh.. i ~r\.'al~on U~<t!I/Tv is .our l\(''Sl!!~!S aJ't~l,s[Cad Cl'lllilis;IC'l~1"" llh mag.approidmau::i}' C(![ISI~fll. wJi!C1J H.c N~\lmmea6 OlilD'oor ;10;(16.; Ololil'jil.tdlB ~jju~ 200]' SUfICV1lCiJl~ nelli,s <!.tLRl('lolmc.d w' ~,he- Qli.mtlim Ite1k (juamliim crhicalhy aoCOJTnr~.micd 1ry the 1O..112&fli:~~c.t,D.U~;O

,l~

.,b.

n.

rhO'

0('

II

or

971

REPORTS

C~ust:elrinlg by Passing Messagle's IB,el.ween :D,81:8. Paints


(luS:leriIFl{l data IJ;~'lD'e,r'llUying i3I ~llM5I: of fI~~~~t'nta.llive ~);aml~le1: l~ jimpoftaln~ tor pro!l:e~;i il~ sens()[y "'·glilals and d'etfflfFl[i palttefill"> llnl data. :SiJ(II1I"e);eml~lllrs~ cain be lUllm:i b~ FJlIldl:l:mly oCliit1Lllsi Jill illlliUa ~ 51!ib5~t of liI.ata il'oillllt~ ilild then ~l.elraltively r!2lfilmm9 l~ ~U1t ·this. work'5~7itLl or1llyir ftgJ U\i'lilinlll'<1,lldhok.~· r:~do:se mal g®o.m ·~~tl:l~iom, We !iieYi~ed a me'li1.od ~allid ~amnlilJ!' prrop0l9<1lliol1;," w\hidl ~ ~esals.I!i1lpWll 1fn~~IU';S of ');iRililJ rril.y Deb."~lnl p.aur~ of dlat~ pollllli.'S, IFteall"va lW;Bd IiT1 ~5s.age", are '~'.'!(liaIl!9,ed IJdl~el1 (!I,Ma p[lin~ lill'~il ~ 1i1191i",(Ji;J(i~itf set af e)leml~lafs .alild (lIr'fi~$portdilig ~~uiSt,e~s 9r·~d'u~ lly '~m ~'rg:i!'S. W~ u..e.dl anillity prllp'.ag,3litJ:ni to 6LuiMer im ~.gE!'5: olf laces, del~(t 9,M~ fn rIlk rQ,aJT,3y diatal,id~lI1tir:l!' rE!iflrEl'S~IfIMtwe ~ril'~f!CIl5i 11 thi~ l1lia 111,15(ript, ~'ndid ~;flItiiry dties~l'I~t ,a ~e ,~ni(iellt ly i!I,~ci!'S~~dby airllfflJ~ tF~j~tt ,A~HliIlty proPJ!)i!llio'li! rOl!!~d (lnJs;ler~ \~iitlil m.u ~I'I ~WllF ~Hor l~!'I tOth~r ml~tliiiIJds;. and it dilll !lOin If'Ss thalli! '()n~liiulll(irC1dllii the amOI:l nt of liIml~.

lilOltI1 Cilm:Hd::u~ \."XCfllrlflf

l)(Iim

LIO ~]iQilll

i,

evidence lorillo\\;' {lJJprop:lime it wouM 1l'C for po~rll. i 1.0 ehoese poiun I; as :i!l.'> c.xcmpillr. ml..irw i!l1iLO ,~.cc:otilmIhc liII1PI~Irl (rom O!li;Crpoinb.l~lm PQjlfl~/; $.~cKmlcl DC an .~.NI1~ll:i1r ,~fi!}~(,~",,((kl :rnc! /Jli.,k) can be
viewed as log-p~!:lb1Ibmly ~.IIU.i,O&1[o b,~gill ~lI'i~h.~.h(!'mJ\'a:i.lab.ili ilic:; are ill! ui~H?_.od~ozem: ~(.f.k~ = O. the roNl)[m!libi~i.nC'-s moe eons-

reileclS !Hie Il()Clu.mllmoo

-nmll.

puted u~j1]g .liIe rule


fs..Ui'l'l

max

{aU.k')

+ ,I;(l, k'~}

P)
hi the fi!'~t ~~em:1io'll. t,.~'II:I~ the ,~wbiliilbil~utk~ b am' zt)ro. r(iJ:~ it':.• "Clto the ~Ilflut ~illli~,n'i~y bClWCCll point i and I'IOill~ k ai"Ll'" C:l!cmp.br. m laus Ihe ·1'm'gCS:~. r the sim j,kllrilk'S b(!:I\\'()C!I~ o
Ctllitlpo;;l

·e1iilt11. lil~iilj,'~il> ~iHI in !,;"IIilt!iail.'lc'1iia~~ ~.,~ [Cn; s, A 0011.'1'1'10,11 ~I.prmo('h. is IIJ ~ISC llmu to lea III 9 S-~~ Ill!: CL!l1lC(s. such Ui:U the sian 0 sqtLtIfCtI coor:s between dJL~ pooil1ls and I.ncir nCar''-':iI.ceruers is 51:ttmll.\\fi;Jcl:I tbc ecntcrsarc ;:;el(,'e~oo :lr{JJ1f1ii 3m~ral dam -polms, IhC)' are Cillklu "ex ..'fnp:i11r:.-:· nil: JX)~JLlihr k-ccm";'I~~ d~'lCrill!1 ·t(."(;lmiq ~I!..~ ~ bcgilJ~ wi,ih an il~i~iIII wt ()fr.t'IIll)O dOJ'I'lIy ~d~lecl CXi.':'I,liplar,:;. mid i~,"TI!li\,o(;'l,)~;;:linc,,'; t ~hiE: sN 00 a~~D d[)e~,l:'lCl d,e ~n~ oJ'S(!I'lli!lmd cnors. k~lllCfNcl~ml(;rin~ is: qllllitt:! ~el1s,ni\le- ~o' ~he inJilia~ selection .0 r clI!c.mplarn:.,. SiO it is. us,l.mny ~C'J'I!J!lI m;l.,y~in~~ wl~h difrCJIC!ll nl]iliili~za!iol1!> in ;[In aHCJInI'll tQ ~illcl ;1. :goPd :'lrJh,l~ n(ln. ~c~rn\lC\"(:r:, thi~ ''WI~!l. w~! OT!~y wh~!l ~h~n~m~~.of d\~ ~

lilI:;,tcrilig dflk! based Oil ~I $iunii'~1lri·LY i:-; Ii ;;:rLi~t.....1 step

.nl.',;l'l::;lJ~ ~ill

ur

);.CicntLliC'

Ihc cllt~tl ~}Oiglll,wi ~h inde1\, .k iii suited to be thc l!ll;cmrJar 1'Olfcluj,:) pa.ill i i', W~u;u th;: geNII i~1.11r.'1i,iri~iiiii2l': ~ql!larcJerror, ~ileh ~.illlll· iilarit)' is. SCI '~u a Ilc,l!mi\'~ ~(J,llar..:d i:rrClT ([:IUcJlidc,lHl di~~,~u(lc:t For points J';I andx~. ~ii = ..k~ ~ ,Xj A'~(!. ~ljdcoo.tlilc ml:'l1.hud dc~ribcd ilcre CllJIIJ.c:apl.Jlicll1 whcn~1I1!,; up~imiZfLtLOn {:oritclliClii i::;,
IM)iW

well

poilu ; .and OIYt'C caml.idmc c1\,cun,plars. This r ilia \''C up<!m-c· ~:sdiL11!-<1riven fmd <.101;,:::;.111m
nOWlilI1L

take imu.

·how u'I(my

'~lth~1" Iwarlls

fia.v(iiit

CRth cRildid:.!tc

CX~Ui'ip~:fir" hi 1~.K,« i[l!lt.:l~iun~.

may ll!;) ~<;oC1 by l~nd. R1I!W'lt;!r 1'11::1~ !'Xf~lirh~~t~~1 ~h~!'I~I!llh~ nof teus i~~nl ~U ;tOO dW:l1Ji;.'C~ are good ~Il",~111 T~.;:ISlI d~I~~~ he P~"f~C>I;i noo. ,,11"li~il.y pll1!(mgal.IDIiI ~~!kcs <t.~ i~ll~U! ,<I !"C'al nl!ll!~l~!" !I(A;.k), fpif C'<tch dbl~:i)i M~h~r ,C'xc~I'J\II<w. (I!m r.u~di.'llU u~i!b~i"-'lliOJ~ b. (:~O~ 'in a ~ood Whcooll:': th,c' :iIIlilO\'C n;'\~p(Hl)1ihili1y ul~dalc ~olllli(m. \1i,,!r;: wkC' a qlL!~!;c. dHn;'Jx)u~ "1~PfQad~ Imi!1~ k ~fI tl~m 0.11.. IXli ~l~~ w illt 1:J;1'.g,cr valrLll::it k,"~..u ,c<lmild<Ltc (!)(cmp.~~!:!;.(:m~lpCM~' ow,,for or ..,~tk,kliitl~ mOl\C' likc:~y ~n be ~ho:~n a~. cxnJld h~trfll[~ucC' <I. !!'lClhod ~ha! ~n!nllh~.J~C'l1~!1.IIj' cr:;;hip or.<I. d.ua l)oil]I, t!u~ fo.llowfi~,l;!! tl\·'ailllhl'l(:{msiders an d: .. a f!oi~ts :t~ 11O!e'~tb~ c·~·m" cm,pbrs. 11I1Cse· va!~ICil ~a'CrdC'ITl:,':d ID :t'l "~rJ.:'rI:DI~Q'5. 3]1 \'k\"'i~lg each <-lat.) Iloint .);,> ~ !llOdc i!~ CmlOCfl." ll1C mun!_leLr nj" Wcmif!oo CXC'!ll~!~ffi it.y 1Illd~IC gatheffi c\'idcm:e foo!!!! d~~a moints 1 ,). I!,\:lwork, w~ de'ili:S\)o a !llC.~IN~ ,J~t J:'(:CUf- (rml11bcr of c:llIswrs) i~h~nllel'l'CLX~by ~1.e \,'~.h.l!~ as 1'0 ..oJI1C'lhcfe;l~h ,";mdh;IID~eex~mllbr wO~lld
;:;i\'!.~~'y l:rml~'iltl i~:s i.'l.:a~-v~lill!!XI n1Q;''';'~I£i~''S· i1long

nmclJrnorc .&~m:lmi, Later, we describe [!J5k:. whLi.1l-"; :;Jlllli r~mili!.;$ m..: derived ro:r pRill}; ·~~f illlil' 1l!lC:l,]MIlliS. of l1ilil!.nmtmyun.~~5 un;ltlcm~ .., P','J~r.s oJ: EIl~li~h ~cn~..:.ooc::;,. IIIjdi p,lInn:; iD r cil~~~. "!VJ:f!.:I~ illii excnlJl~~Nl:IJ1'1!:'.'nd~m. p~'t1h<libiUI·Y InQo[I:C1 i~a..'ai~" aMc. 4Ul) C<ll~1 be- set to ~hC'l!O.g~li kelihO(lii of d,tt:1L point igii velil that .i,IS, cKcm,l):iJr i:-; poin!. ~~. 1\ herll;n ~\C'b'. when 'lp[ll'(1prfi,)!c. s.im ill!nri~ies

wliICU ::>OIl'Ic puinl;5m'e cffceli 'i/'d}' :fis~iguc'tl to ~[~iCr '~x.cmpl:.m;. l]l~'iir av~il.l1bililnC1> wi~i drop bc~o\.", zero as pttSeribw b,y [he 1II1x:hutcflII~C bdow. lhc.e utg1l1i\,'C avadilb;ili~i~ \"ii~ decrease the cl.Yl.:Clivc V'ilitlli.';:;, of some of the JlIlpUl

.... ;~Qi.k·l' i~I dJ:(;: 1Lbov..: I"LI~o;.I'I.:I1lovillg 1:1:1..:~"(Dm:;pu:rK..hl1g [.,;mJl~t11li,.; cx..:ntf~ I~]n::; rr(]lm tt.1mpcli,laO:l'I. Fot~: = i; Uiel\:::;p('a1~il.'!ilily ,1 k.ok) i~~~.to Ih~ inl1l1t fWC'rC~!~J::e Ifll<J1l l:IoiJ;im It. be Ch~~l a~ ,UI ~C!11J1hr~,I'(k.k). l'Ili!~~IS !lte 'l:u~C'_g( of Ihe- ~intihl~illiC'l1 ilct~,\lt'.Cn Ilflhllt i <IImla!~ o1her cm~df:d,lIc 'C'.'i:CllnP~Olffi.. Thls ··8Clr-A:)sPQn!libi,~ili· ~dl(;,~, .. ;Ii~UIHUL1Wd ~~lid~tlCe- th<ll~ poh! k is:;In ~-\~ml}lal', ll;!!l~~ tll.ll :il~inrln (lroi~~~C!?wln" pcred hy llO\.\' iU.+,1l.11hcd ~l L~ to Il<I,):<I!;s;ignccl m
$.UII~ ililrili

!Cdt~-=-o.f 'tlie:

u~'ilnI n ~ood Sli!l C1oi,e Irllpllll1:; und cO:I'.r!.':t;l)(lindi~l.& (!:~miiLL.!r.;,l~r;'t(:r&,!.'5, .As t1..-:s.eribl'tl bu..:r, lifI(~ag,-,:> 31iC LlpU:llcot'i. !:JIl uh..: b:JIsis (I r S:iu1lJ)k IiiI'm ul,J.:;: 11ml :!XiLrch .tbr minin'l<! of .m uppwpritll.cly dh05~n (~nlcrgji 1.!tIlilClnulIl., )\ 1 any p~llnl i~l lime. till..: ll'l<!gul11!llde ()r ~<,'lt'!hnn..'l>l>!I@:'"' r..:·~kcl:; the .t;:u.m;n~ IIfli n ity ·th~t one (hU;1 ro.iHI 1m::; ior chOo!).<;ing :t1110~h~.· dlu~ poilUI. 11. it" ~"Xicmplar; ::;0 we calli {1!11r nrmlih;... iljd ··aft!1I iLy [pmU)~I~at~OITl.'·Figul\: I A i.Iil,Uls.·tr~n.c5 h!:l'~\ dusu.:rn gmlltl~ll~y ~~llnlllrrg.cdLlr.ilng the .!l)cssage-f!"I:'<.s;il1.g pmCiCdurc·.

Dclwork

or

or Ihc ulllIDtprdcn;llc~". iml ~l:tJ'":'In..:~~~ frulTll. .. !.he aitC:;SI~!J.Cilm;iSi IlW proc~dmc. n apna;.Ll. 1111 Jn1fJpo im!> .:;re ~X!lmlly ~lJ~nahl-c ;~. c1{'CllIIpl:m;". the

1Il;~lkct'l glli)di

cxcTl1plar.
O. t(

:l;boukllJe· S~'l'l'[0 a. l"ll:lUllllilml \'~IIU(:lhi~ ·.... ulu~ ~,:::,m,Ii:rc v~ill'it1(j 1'0 produce dUflI,11:I,1U ~'1Ull1bc:rs () r .clll1;SkTh .. Ih~ :.;,hfl'l1.1Uv .. ~III:: ·COUIkJ , b..: l.i:.c 1:l1ooiall o.r 1.1il~inlll'lit sillli~ari[J·t;S (K:5ulling in 1II!110d,""fl!II~ II LIIIIlLx.~r()r Ciul;:l'tn;, or Ih,,;ar mill:imLIIll ~F!.:'.:mh:ia1g irl a ;li;111~11 1lt1!1l1.K'T or
1)lCt'(rt~I'Kl'-":> d~11ollcDi'.

~ (.i',k ~-

Inifj {

k J:)

t •. rOll.[~L'ir} I, (.2 ,

~Kli)d

{V(l.k

H}

Thcr.,.; ~ru
CO\lInt rI

1\',1[1

ki.nds

ofnlcs:;;a!l.c

cxt~lul1g.oo

AmnhY!l)oop<Ii,gmiQn ~akc;..,~s input '<\ cLI]Ilee! iO!Q o he;l.I-\'a I ued ::;.h~l!i ~Cl> bCI.\\'QC'!~ LuiQ d;\t:l
nO!fitS, \\'~1eW

~~c S lTIl! ir<lf~IY .~·ti,k) ind k~.(:s

·De~Flmenl: o!· IHe(~Jfc:.a1 .a00 (.o:mptlJl~r EI!1~i~~ering.

UI1Ii!leli)lt)' Illf T~rmo\ 10 jij~~'~ (d1lege Rwdl TIIWFitG, OIl1:.aJilII M5!l 3GIl. !Canada" ...~:c w'1~C!~ l'O!'~~~Drud~,moe h~d
f~~Y@!lSiYw!TOnki\el!.u 1J,e, ~dd!lil5'lli:d.E'mili~

~.I.t~O :H:dilf1bt'C'lT~ k~l1d of oompc~~iifln. Mc~S.(lg>cs C(l1'I be combined iU ,~ny ~t~c W decide wh"hroim~ 31t'C e,"i.c!nplar:s f(l'r c:...cry (I!h..-, poh1t. whkh oCXCmlll!~.ri~ bc~('lngs IfI, 11~G' ";c:-;pollsibili!y" r'(f,h ~C'!'lt fro~~tkH;I.1(1i!11 f to ( caHdidalc c:-!:cml~l~r pili,m "" renee!S. .I~ aoCUI]llJI<liCd c\'~dcl'icr: ~br how wc'II~Jitcd poim

bL.1WII.~CJlrl~na po.UIlI"'-, :md ,~adl

~Ik{"'....

The ,;Ji,vlIilabilliLy a{iJ) I,", ;;.(;t to 'Ih~ :;..:~t.L't~lJ!IIrl>"j.lbi~lY I~k,k, plUiS. lin; ~!111II Ihl" ~l!o:;i,liv~ i of n.~l)l):rI>S.i,rcri~iti~ ,cnndirlflr.: ('1{"-'llilP Ilar k I'I;.'C>!J~ vc::; .Hkm~ 01lhl~ poi 1lIrc,'o, .ollly t~IC pm;il.i,vcptmj{,\1iL'> {If ~~ICLlm:lill~ r'~OII~ibilil.iL,.';,>; ~ru m1ikd. !xIcmll>\l it ii:;.01111)'llilc(~li!,;,.'I!Iy till' n guocl ~~1'i:(;rnpl:Jr~fil!Jxpl~.i]]
:;OIm: d1lilll poj lilt!:;; wl.~ll. (piJ\'!oiiu\,'L,) .rt:.>iP!{):nl, •.ibi~itiQl). reg;ud I",~ Oil' hfl\v poo'rJy it ex~}~1i~i~ Ol!).\! (L1!"

·,ma..

~: is m ~~vc

iflc cx~m~)lar forf!o~m i. tald ng flilhcr pot(;,llIi~1 excmplim:s ror I~oim i (.Rg. Ill), TIle ·\<!;vfLilabmt~" nUl~. em ~
~JiS,

~ntO aCCOlllJl!

P{litus ~nC'ga!iw ~$[i'I1!~sibllitic:"s1 Ir .he ~Ir!'\'s.1J(l1).'l;i&ll~ It)' dk..k), is l~eg.11~w (!nd~(;mh~glh~~ 1''\1in~ ~"tU~111ly better s\u~l.:d as belonging to k <iI'iO'~h~'ii: exemr~nr ;'1111,\:, th.m bi;i ng <In c:xemrl<lr h:-lc:l.fl, ~hcav~.~~abiJ hy of 1'1'01111 k ,LS .",~~ e.~,empibr'C<ln he ii1lcllC'!;:l.OO if~oH!c ~hcr poim;;: ha\'cro$i~ii\'c re5j'lQJlsi.bi~itics for ~1(Iim t. heil1~ Ihek cx'cmrillr. 10 1 imit l.i!C' inlillLCiilcC of Slrol1~

91.2

16 FEBRUARY 2007

VOL 315

SClIIENIOE

\L'V\MIV.sdencemag.orgl

REFORTS
incmniu,l,! p.l}.o;;U,I\lC re.'>pr}llsibi ~ilh: s , ~hc tOlIl~ slim is nll'c:s.lilol:dcd so that II call1ftOl gOll00'l"C zero. The ··sdr-ow.lilfLbJlit)''' a{k.k) is lupdalcd

dma 'POili~, Ih.:n is tll~ e1(cl1:lpl~r for point i. TItC"

WhCUllhcsc dcci~i;(j'l1s did not

dHU1IItC

J.Qr 10

(H:i'Ji"cIT"'l1:liy:
(1(A.k) Thii& "l~~O

r;';lc(#[l1fkle;hl .~1C'Cl1Illll,lImcd {l\'idt111C'C dl!ll

l[cWJJiolls, mC"SsfIgC"'il)flSiJ..~ng procedure may be lC[l1iIlUI.C-d F~gHr,C"IA shows Ihc clYliam~cs ,of 8flinity f.~klr a. 'i",cd Ii.Ui1l1~Cr f il.Cr.:UJOUl.'" a [~cr chfl1;g~"::'i, o Iii ~h(;:n~ICi'ii&Ig~ ,1JjlJckl1,I\' ;'I iIl1rcshold. Qr~LI1~r p~)ilgmiO:J] Ilppi icd to l:.'i lwo.Jirncr~iDu~11dUll the !tJ,:;;fL~'''.Ci$iom>:>~}'oon:;mrIL rOr );(lollll,; ruund bcr ,C!f il.t~mliOJh~~ Whrn IIp'l:a~l:ng n~mnlC',,~1!.tC'.'i, il i,ur:n.l'l!OL'm~llI lilac. IhiJY be d:WYlip>od 1.0 :I.void mJmcriral 'D!Oc;'ili~Himl~'"hat arise lin some cirI cumsrances, E;,u::h nmssagc is tim to A lillilC'_~i~s va~u{l rrl{)m the Iln:;\'im.l:' i~C'mliOll plus. I A timc."il:l. prescribed I1IfU~aICd\ alue, wh~r'C tile dmllping faetor i., I:;; 'be~,wC'~1l(~ :lnd. t 1.11
all~~lfQf!lr ~xp~rJmcms (J)I, we H~~ a ~k,rau~t d\1lMp.illg r:)lC~ClrL'l.r .J. - R5, "nd cad~i [~l{lti<o'!fI 01' ;(Iiffinit)1 l~m[1t.1.g'b~ioll CO~l~,itlIJCd 0 ~ l!Ill" dJlin!l all ~:-1pon~I'bWtk<;, !li\'C'u~h~ avail:llbil·

~n":!i,~{l,.r({.kH P)

p~)iullz; (.~'I,!lliing

JL'CWLt~vc $l,lL![u'cd

error as Ihe

lMllm 11: is all ,~.ml1J1hr. lliL~cd O:llnHK! poshjve IN;:81l<msihilil:l~ sent [I) eandldate t1'Xl:'mpl;]f k fmm Dlhl?f 'Points. '['h~ :~JmfC u(J!d~ 1ilU1~';l; require ol'illy shnplc" lecal {:oll~l.mtalit:wl'i.~Bl[lj1 ca."i~' ill'lpll~IT~~I'Ib;l~ ~~. I( 21, mid mC&1ngcs need 1}1l1.~b>c' £'Xd'l.1Ilgi;ld be'~w~n ~)lIi~1'i[l:t ]'l>Oilll_~\'!dlh k IliiJWl'1l siHli!lnri~ies:, , j\~ any POi]]l, dl.uing OlIllfinity Ill'Op.[Ig:n~~m,,<l'V.m,i[i. al:>ili~i"",,~ !'\,."!';(l<lIill1ihili~:i,c-s 1IH bc' ICClllll~lned ~!10 :md C idclru i,fy C'1\,q!illp.iiirs. Forpob1l1 i; IhC" v~luc of k '~hm maxil"l~es II{{",I;:I I r(i.k) c:i,III~'Ir klcmHics rmilill l~lli fan ,,..,,cml,lJllffr:iIIIi. = i, OJ id~nli:i:ii~ the ...

!'iimi~al:i~_ o.n!:: advmlt~tgc r.lr~~llill~lj' 'Pfo[lpng~]lon ~~ d}~1 dlL' :rI1,lf1]hor '0 r (!,x(;~f~pbt["" nulo'd nOI. be ~pt.;--cmL"d bclimlh:llml" Irl~w.md. the a~'Ir~IEL! nUlll1.lJcr Or'L!..'I:,mllph!.NC111C!!\,~ timl] l'Ilc mc:>l'1~· pas:;;ul~g ml:!l.I!md and dej.'lIClld:;;en tile input excm~l lar p:mtc~nloc~_ This mmM'{)s <L\iI.olinatic m(KI~1 :iI~Ibt::llJC!Ul, Im.~-d on a prior spoci,fic<T~iriJ]
of'how
OOllUI1.f.lll

P]JI].r~:::dl~~ ~a~h pOil.lt i3;1ii an e1>:!~'Ilil~:ilII'_

or

ilic~, upd:niujg all lI\'llilabiiilie~gi\lCU the n:":lmh"ubili~ tc!l., ITnd (i HI oom hill i!lg ,;)I\',",i I;]IJli~" II i~"S nm:l r.espom"i lie.,> 10 UlOll i~or ahe C;,,' cmpl,MI: d~i:;'~(I<I:IS~nd ten nl ~Iatc til,.; .~~g,{j!fithll1

un

~lj)\V:!'i Ihe' efl!ol;l crf Ihe \P,llue"flhc ~II(l(!1~pII~r~llt]!lron Ihe n~lll'!OCf of o d!ils!L)iffi.. 1l1his :r~hHioll l'i. !lC'<ll~'h' ide!Jl~i~ 1.'0 Ihe ..~ rel;Jlton fmf]W1Ib~'cX>J.c~ly~niuillli'lillg~Ure Sqltarc:(~ C'.ImJ'

figmlt: II)

~a~~

~Jl'
[flC";~u~;inlg i~e •
~I,imdflmd

\VC flC;\l $~w;iied ~h~ I)roblcm o~ dUlsl,c;riulg


~nll~@'_~ ,,~

Op,tlll.lizw·

...

m=JUi,filONI I"

.
I
'

,-" ?

l~mor..llll2

I1liRA:iiIO\l\lI

.a,
I
'

.'~~'
.'

'

'.

....

.. ...

.
"~ '~.' ./:."
_.

...

"

Fi'!!l. 1. How amnit.~ W'OlMg<l!JOnl I!'rolrb. (Po) Affinity pJepa:9~tio:nl lel: 'HijjSlWatt.ai~or ~1!l~m~l'Is:h~:r!t!~ d'l!l~ PQ1!'!t~1;'I!~if'l: I'l~~'~i!,!l~ lEudidiNIfi'l di~~al1oo~5.ql!!iu~dJ~'mlr} '~'la'S [B

.~

!.!sed

tl)< lilUl:i1Stl re

'simil:aJrilV. Ea,dii

1[Xl~iFll ii5

iCoilo.redIarr.o:rlllililgro '~he (.IIIUeFl!t. e'lliiillle.fKE

Se!i!~iJ'!lll ~biljl!eS

thl'!til: ls '" dl/!Swr cenwr (",xeml~li!r).The ~rftm~'s!'io ~f ~!l' ollrTQ'I'II dlri!'(~~d lrom poimt i ll) point k (orJIespemb (0 the Ml'llllgill ,0:1
lh~ 'lr,cllll5liTIlil;l.ed l'lIle:S5~ge' tlfla~pf)il'1t J belmilgs tl'l e:<fnilpialf poont k. (B) HIlte1:pOO'Sibi~t~~" rli.kl @r'll~lillfr!!)!lIIoo~ pq]lnt~·t;;I 'GIFLtliil:l.ate~,{~mpl<lr~ a~ul ~nclka~!i iM~1I

:s~mlligly 1~~tJ d~ lXIilllt f~\f®rs~l1I!! (.a filii" da~e exempla_r ,ov.er o~nfr caMiJ1ate e~emliI~ars. {el' "Avai~lbmlies" ,(l(j,k! are se!ll 'rrom cafildidatce <ei!:emplars to dialta lpollilt:s oilmlindkilll.elQI w~a~ de!J ree eaeh ea mfid!i!Jte 'e\Ile'ITI,Il1.aris a'l'FliLable <15 oil duster [~;nter i,O!' the d!3t:81 po,il'llt ~Di) The ~ffedefilie'll'ahiJe ofttle i'nillil prerereJi)(e ~(®mfl'1)f)n 'rorr illl data po.flll~j oTIiatJeIlulm!Je;rofi~lflll'nM ~mpl:.afl: (1'ltlmtJer ef eJu5.ters), ~nm'!jtiJ. lfle v.3ilu:e tl1at W~ us.edlilil (A) 10;;al!s® s:ltuo\Wi!, '~l.itii{1I11 'r; T w.a~ i(;cJimput~dhiom the media!'! or~i1~(Jalin:v:isr!simlarities. :

97'3

REPORTS
Sl1igC.~ be~wc~nlll~Y :!I ~jny.sL!t.'>C~ {011% or 15 O this OOl1tc.)(i1.W( [l(jdre.~d [he UIS!.: OrCIHslC'rit~ p<lit:'S. r~lati~ c exons W rlll!1 !!!oeiflCs.usl ~1,g~ilC' s:J~1Ir*r; nnLlI,ion) ofdampoim Fig_\u,; JA Ulus;~mle:,> 11'tc idemificfLlion of si un~lflril)' Iliiilri)(. dcrirvOO Imrn mic:r()~rray {!i:u;:tt, oC~~ffi'Ci,~(J li{l'rn Ihe O~~v~t~ fa\),; ('btaoosc (3), mU~lii:pcil1ed gc.lftc clus:t'C1'l> I\nu:l~hc fls..,i:l;lumcllu of :SO'nn~ dalU [['I (4" hll ihm '!Nor1l., 1:5.'0(16 scgA Ili~1 l)roPi~~lion it]' loulld r:.\Clllilllil~ w i:d~ Ine,m:l; orDNA t60' bases klllg} oom':$iXmtl~lIg OCI po:i III~ to 111'0: IlIJI1JC''-;Gn e;>;crl]p~atr" The '1.'4;:"'011struesion CrJr'GIN 1IT~11L' ~it:' p'Mp.:JgalioJ~ [LI1d 11.,an1 much lOli\l~I'sqmm:-J CIm:r d1~n tlTlC'h!.~I, ,of I 'OCl' [UlllatQVC exens were 1111 i.II~1 lrorl'! Lh~gmm~ru:: or rliIns of k..cC'lllt~r::; dm::h:_'f.lng, (rUg_ 2A,I, 'whoch mouse dlDlIll'n'«Ill1IcL TI.1;Cir-nrrn~cr:ipeiOll kwcls centers dLL;;h.Jr.ing, ;al\~ c:m!\pm'acl :111 Hg" 3B_ 'took. a'lxlI;1I Ih(' SmltO :J1]lOlll11of oompl,Il('[' timu ,!.em moo."lI~oo acroos 12 tll':5ue smllll~l"S. :U1:JdIhl:) For each number OIY elusrers, :Jmnil), p,mpa6 \Vc'ilIskoo 'wl1ClhCT a ~nug(' ml[IJllh:r 01' ,lI<Im:IIITIl 5:lml~I~lril.}' b(.;I,W,G1L![TItlVe\I)' P,II!" or 1~11,.ttiV(l C1o:jcmc<: ganon '1;,'.:IS rlll1] encc and ~DC!k min, \l;'hcrea!': k·CC'.IlINS d~ll'ilc.dng \"'Th~ mil m,ooo tlmesanrl ~'Ql11ll1s or k.OC'llWI"S dl!lSledllY mmlld :lch1c\o'c (he (dam 'iJ'Oillhl' W,IL" compurcd. 'flu::: 1.11C'''SI!l~ of r{]lok2(j,~ hours. 10' <lddr~ Ihc qU!o";5IiOH orhow :s31i11C ~'lqumred CI!lUL iFigllnl 2:B:;;l.mW',~ lhc C'.Im!: :siI1i1JUfm-ily b(!1wccll 1l1,ltml.iivexnns \Va.. bused ou e l:hCLr pw.xil1lily in d~('g:CI1QH'ICand the d('gil.'cc of w,;;n IhCli~ m~l,hDd~ perfol'm in clctooting oona ;achiC'v~d by one I'lm \If alTill.l:ty :l:lm~~g.;''lI~i!lJ]and the d,i !l,flli:lIlilian f.lf ~n'Mll,:lic'hiIJ\'cd hy lO,rmt] ooOl'!;!lirl:lliol'l o,t' Iht;:ir Imn~ripl~f.lin I~ds 3C]:m,." lide- gcn~ lI1~PiI110Jlt~, fig. ~{' pJo.t~ Ih~ tn.I~· ,ruiU n.f k«:f~l~ d[l~I_C',ring. Il~OHcd OIg~jll,,'lI, 1e ~h(' 12 lif.'oi.LI~_ Tn n>Oi;;ffilllll, 1M l~l~lmIU'>,(l d cxm~:!'i rms,uti\'~ frf'I~'aJ~ .!.Ig.1JiJ:l:&1 the ~:II~~fKk'llJlive (Fro) i Ihc: Re~cq th~t .[I:t'C' 1I0~ C':\!.0l1t1 ,(tl,g,., U11ml1Sl. \l,,'e 'il1dudoo <Il~l 1!;I!11;l, LL,>ung dlC Inhl,~~ Il.!'(l!\d.;dil'l Ull,mllx~' C1ircl[~s~r.;:_ AfliuiLy pmpag,ltron LUlie d:llaba~' (51- Aml1iuy p~g<Utit:m achicv()d .'iiig,Q'x.ell'lpkil' OIl1ddcK'Jllljmxl the Ihm1lY<lehicvoo Illu:ch ~owcr (lfl'f.!r in~ mDlrc th:m ,tddlitimm~ llnHki.~J ~W(D mdcffi. nf fll~~ni~l!Id~ tW'llC'.J\uf.llh~J 11Oll: ~:i nl~I~IFiIY ('adl o~l.er data llR"1int !O Ihi$ "'nOIl- nl r~C'OIntly Ii! i!!J.hcr TI" ra!(',,", e."l)~ i,1 ~~y ;11 low Ie;ll m of [1'1 ma' nJ:U.imiz.atioll cl,'i~:ri(Dl1, .i s I.he S'LUt.n or <I~)- exon C'xel1lpl~II'" lIS!lIl,!J,s!rnn.isJics mlif,'ll M'Cr the- FP rates, \\lllli:c,]] are llilosl :iIll1l)Q.nm1ll.h) blologl:sl;s, A.i 3 FP RUe 0:1.' 3%. ol.1Iltlt}' Ill"Op.agm,iM c~nil'C Ola~, l!.CE. Ue",-:;uitilllg 75;tJ67 :-; 7::t.Q61 ::>o~ !Il.1:pi",e~ tlil:!ctreIiC~, (W11icl1l better loic:t:a tcs ::;:iml~~ril}' mmri",. ,(3'~consisted !)9,73'Y>) simf1c;hn>c",'~d u TP Ia~.~of 3,9%. whercfl;> Illic b~'S.~ '!;mlly~n~ pixel in:tCRISll,ics). :;(l we :I,~'~mcd the illaritil!> willi Vl~IHe." (I<~ "'"\ ~,mn;:;;ptmdi.n,g [0 ,k·CCllicr:> ,dlu:;.tcrillg I\.':>LI~t Wil.'i :I. 7%. FO<ii~"OIlrl· above pli'OCCdu R; u~:il1gUlr~,;rnJJ 1fIH:i:sun:. A flfin· " .. i di~~lj~ DNA i>1.'gIii~lt:;; Ilitltl;OIl~tI ~'i:m po&s.ihiy .. ilyprol',"~[:iOJj 3gl1ili lIliirUI11'ily 1i:Chi,.;~~d lowC't P':U'i'OOiil. at Ili~ same foP rn~C. Iii...: ~e.~t T.r i'::lb.: be pllrt {If ~h~S[unc g,'CIlJ!.\\l'(;~)plli~,(] a!liu iL)' fur ~iicrurehici~1 a~g l:ou,\C'rl!li\'!1! dUl~le,rn n.~ ~]) error I( 1F1~.1("), was ~9¥.~ Dud the ~il,g UI1l~~.rilig~o()i lCc';Scribl,,'d Ml'lny~l'llXtJlLire roc itl'cmulicmwl1td" 'i.:Xe l}roJlr~laOI,l to Ihlls ~iullilaQ ~)' Illtllri:<:.bn.nlJc· cause HlCSStlg/o.':i1 IbC'OO 1101. be cilIcl1,rngcd OOIW{"I!1:] en,.pl,fIliiS :1Ii.Km;g ~mdy relmif,;d dulli. i.e .• where ~nI ~Jl. whiell. aceousns fur additiunal b~o· rojU[ ~.tutti kif .~(i'.k} = ;t;, crtChil~t.:L1iIUu Oi~ ~ogicmTkl'lu.wlc~c, nnos. sin'lilm:iii~';S, are .,;:i~hct ILU'lkllOWIl or I!aJtc f.tcilic,'oo 1LlW' nne of 'B%. Afiiinli~y pI'Opi@tklll'l) nlbility 10 opc:rm~01111..: ilmn~l)' propftgmifill n;:q([il ut<I e.-:dlHrlgillg rncsandlTlegmlV!C'. Jo e1t~111linie~mlJ~l)' p.fOptll!::i~u_iJ)J11 in bll"i'~or .rtOll~l~nd'~uu UI)~imi~ti:Orl !':ril,",rii,~nlml!.ci:'! ; t sl!Inl.:.1b.k 1:01" !.:1'ip~o:mloty dl:ml~nrllysi:,LI;,'.j,i]l~ 1!IIiiLL"LNJ~ 1~1;C.1i:<:UP1!3 ·tlf !>[rni bJrily. Ulilikc liElti.csp~ce ,~.I i,:;JoI;lrill,g IcchniqUCR: ~'ll:ch as k4'l1ClR!1:.1 L Ol1alnBl Imrillt'll d!JIsrc'r~lfl 01 .~lTil1jly propag."u lnn C'lf! bc ;lip'" ~"=:"':=i plied ~o' Iln:lb.lI~Il1!l where 1hC' dam do rnell ~k ln a A!'1U!il¥ oomijlllLI[lUs. ~,~_ Irde~l it {:<I!I'I !}C' OIpp~i(\d IU ~~==::::-='"='-===;:::=~,=="-:=~ ~O"II~ prnbli.us \\~1.C~ !he Si!lll~~~J~dc~rc flot Sylll~lC~.a Ii'<ellltc ... e!:t.I1llerlll\i:J rle [!,e.. ,~(i,k) t ,~k/I.I,;Ii!)(110 prolll~lns whQf~ !h~ :,:iHllikl!:i~ies (I", nat sa! i\ry thC' ~rii1n.ilki.,~~,\lalhy [i.e .. ;li(i.k} <: ,~UJ} l .~(jj;H To idcmirY<L ~m:;lH

'~i n crherion o

a mil ilY Pil'0~~1[~Oitl :mtl k-CClnCir:. dliSICri ~lg; ~o idclfu i,I)1 ,;,;\cn,p!!lrs 3n'iQI,'I,g~Qn grdY$Cillc imflge.~

of squared crror, We used both

or

=~=~

B,
1$01 1W

,Irc-ee1'll1!fl ,~~~
'~l~'~

fUUl~'C!,,(l'r Se!!~lenCe$ ~I) a

11[0.000 rul'll)
j-II.ii>_ ~I~"{~'. Mnl~l~g

!flat sliIfllm;l!'!I!'.'e

dmt1 ('If tllis umu~."'Cril~t o1her ~.(m!e!lC;;,'K. we lre..lcd each


>l:lr wo!ids"

_ _ _I

scntcnec

<IS (I '9:l~g

1 1~
II :I~ Ir i'I

.~

..,lnn

I,

100
100 00

~Inm)-

lP~rcn

IliIC's:i!~~ikt!i!.y>of j;CJ'lt.mo~ i '0 oomc!Jee k l"):'Icd Ol~ LIIK:I.lm,1of cnK:OoiJ,in~g ~ll;WONS illl :;~':Il~~'1I1tt U;;;iJlg 1 i I.I,IC woni>;, in ~'iCnb::Ucl; k.\'\1;,; 'ttilUlld thai 97% "jf
1.~1!..~~"lI~i~l1~ ;i;il1ijltllrili;C,~(2,

«(j~and OOF.!1cr)~I~cd
1l0~ :syn'ilTlj.ct-

3)'M.ctr

t
'''I:

!four $CnlCocC:S i;s :.1:1["), ...11. in f~,g oleA. .. .. Wea I.:;[D,~pp'lit:d .:JI.flill ily pr(lJltJ;gation Eo C",Q plolt~ 'dlC [nol:!klll or idll.'J11.il)I.~ng .:II n;.'i'Lri,c~,"i1 rmlVd~cr of C:Qlllllldiml ~1Ild. Amc:rlc ~Illcil.ic;:; 'Ih~,t arc 1iI'I~1 c::I~i])' m:IJ::c..",ibk IJy 1:'1 ... ~c :O:1!I11J.s,~ls of Fiig. 2,. CW:stl;!rilnfJ[ f~{~s, bclfTl~l.JrSi nniil'liimiii,n:g the ~tilil~~rdS!lllu~red ~I'r~r m~Sl,Jln~ '!;If ~11i1i~rity ~\\~~ othl.lJ' cj'l i)i;s, .i[n t!..mn.;; !) r ($E UIl'lll'tOO (lrJ:11iIIl.K~'fCi,m.li idemHiea hem 900 f'Kmm:'lJimd force kn:llil;leS (3). IFor <I {O,mrnU91li ll'[eri!1!reli1~e -600., <lUiifli~y Oil a~FI,in~ '~m'ld lim(J:.. Eadl di:,II:Jpoi:nI, \',~'~]~,:J c.i.ty, r;lropa~OIti(ln lQ'ul"!ld 62. dusters. i:lJld the averil!~@s<lLMred er",or~s JiG8.. fa,r complO!fiwl\. ti"!!@ blest (If ~lIld ~hC' .:>i.im ibrhy :I:H)), \WlSi!le~ [() I ~lCn1c,gm i,'IIe 100' [FiJUl5 O!~ ii"(flilllf FS dIl5:1:e~ng ~'I!Itl1[ dH~efeUl~ I"aoo!lm ~111tia~izatlOIlS a;ri11~iii iii \l/®fl'if a:'i''frage lime ~.~wkcs ~] travel rro~n dli' i W, d~y k by S~W!i!IlNI'~ml!:r ,~f 1:1(9, LIU 1!1!~ J.5: i~!l'~s; Wiltll high~~1 ::;q!JItl[ifl,d I,lUi;lf IUT];d~r eiltll~ ~!fhlity prQ~~li!;l!l1 :1~OJl(I\"'CiI' dcj(1~ CjJ. >01' ~·,c:enll1! dU:>l~rilll~are rs :!:howi'l irl the ~MlI. 11'1@ in1~tl'(] lEl ami i:le'tllom ,rMl.I5 ~he e:!lem~~a'rs ;l.~rlblC. inc~lldi!lg CS'li!lH'I~e(1 D~c t(l) hC'<ldwifid~,gh~ tr.msil ,ime 'II.'~ hI iffi~llY ~ss;igned blj'!liIe tm e~h(ld:s,. a nd the boxes show which ,cdthiE! two [f'Flilltlll@ds Pe'.r~'eI1ITl.@dbetter ~(Ut~l image,inl l~nm~ of :sqU'.illFemi error.. Aiffililfrtypropagatioiii '~0Ulild hlgli.e'f-qlliilUl)l eXf\ITI:plaNi. '(B~ 1I11f C'.1.~S diJferont depcl'!.(U~ng em !he d ired ocm of I"WC-J. ~ Ih<it 3(j·% of dIe simibrilie:. were ~ve.rage S,,! U"~ red '(l'~Qr >!!(;hlev!l'd by ashi!g~~ !fUn ofaJrffii!l~t"y F,):Fl;lfl@~@th;lf!~Ild J10,!QO(i rtlJIlIS, of k-~en:l:e~ nsymmcHic. F\~f1ihClI·.I:hr rt.l% ,of ei~y I~~irs i dlll~tEtril'l;9,ver5.ljS lh:e n!JliIllSe1' of (lu~ters.The ,colored bands she'lli'! diUenmt percelll~ite5. olf,'sQLJ!arnd Jnd .t.o, 111Crc was a llih!irrl cit~' j lluch d~iill Ihe ,ellVor" and' the f1umber ,cd ;e!llemp~alrs correspondingl to the Jesuit Jlr!Hl!l (lit) is 1I1(jil;a~ed. (0 The am,ve I riffi.llg~~ Incq!:l31 hy was dQ~.[)K'd,bccil~l51ethe prooedul1e wa~ repe:3il\eilll U~ft91U;f nun ofalli5clUitf furors.as tift,e meEl!>Ufe ®f fim~liln~y. Whh:il1i '~ alL~oa I,rip fi-Oln I toO k in.clu.ded a IQl1g ~moillQ"'~r i!:i:ay ( pOIl'l!l~r opli!ml'Ll!tijo!l (riter~o[1.
Wi
,:100

r.i!1!. Th,e prd~r~ncc:; ?"l':fl.: :!tIjllsk!!1 10 ~dcmi~.y ~llll~l1g 1 = OJli) dlilll~tl;;IU[ll.u:lt~bcrj; !t!1" ruJIrc:;;~·nt;lIj\,'(~~cliIlJ.l!ar 1i.Ciflrcn.ccs (1). tuul the suil lI~iOrlwilil]

L-------------t; '0 ~ liOO lSi) 000

~Ci"

mow

914

16 FEBRUARY2007

VOL 315

SClIIENIOE

\I'V'IM;V-sciencemag.orgl

REFORTS
in ci_/l'

.i

5iLJ;

il,Look

duratioas or~hc ~rips from


WJ,)cn Ihe
Hl.lrtll~Cr

~QrlJ;;'Cr 11i~1l !'ire ~~mn(If the i to i and f to k.


"ialO$~

of

accol;."'l];:.ibi:euies" c

City, .I..us AIlJil~lc.'.) or bceause jheir ilci~l'II.mdlooCil; caabe more e ffi.dc rUI)' accessed lht()~lgli o~J'lcrtlcs~inililions lC.g,. /ui:mHl •. Pil ~~=

poiUIS;" Eac.l1 ]~bc.J iru:lierLtl.:"l> l,iIe '(;.~IC"i'llil'lIarm w.bli.el1 UIC peintbelongs, so Ih:[l[ :.\ili.(·fl' is U~C san ilijjdL)' of <1m:!! POilil, i~o i:~\Cmplflr, q = i

10 be seven ~i:Yy mljllslIng the iup'ut rn;f~r~i11,c(.;~;p~lrop;d ;lcI}'!. th,.;c itlcs m s:I":iOWUi .111 fi~.,11.B 10 .E, were irk'ml ~Ikd_ :~l L~ inlcl);;';SIting thut S0'i~rnl I.mtior dt~CR w~ru' 1101
W·1l!.'> cmlslii1tiine(iJ

acldplii:-l.

fnl(\MillnC3poj~"

cll:go's de$linm~om •• white a 1''lJOri. dcl:!l.y.ll}_

m,;COLlIiI lor chi· m'oidlillg ptflCmifil

L~ a :>(,!cclfLl case im;J.ic~ni~'1g dl fflt rO\in~ i is i~&Cif


all excmplnr •. so 'lh~l, .~li is I he ~~-'fj~~1 ,'q)' 1~l\;f:' eeeuee .fOT" point i. NOI .. ~.II. co:n.tigllr.lln:iDI1~ ol'lhc labclk :.1m \'rllkl~ :ICfl1]tigllmn.lcm c i~vall.id \\!h~l ror ~vmy point i; iI' .iolC!!ll.!! ther 110im th;l~ o dl~(]I] i all 11.<; Clw.I.T!pbr{L~ I',~ = i). ~~um .• ll1lu~1 he .m c.xemp I~r {i .e " ri.; = ilc TI1C Clle:JID' flif a .. valid Q(lnJ]gtlm~i()ni~ ElC'1 = ."(i.e,I, Ex-

~.s

sdcclcd. either OOClIU!lC!' ~K':I.VY i!lt('nru~l;[)na! ·t~.n,«llmakes them il1:lppmprlalc' as casi Iy acce~ilibh::·don'l~SL.IC' destinations (c_g .• Nc\y Ymk

method

C:JJI be ~ t~w~d .~5. a IIml. :1mu·cn.t:Js Ibrulliinim~ Or~]!] ~ni.!r~y FLmcLion ( 7) Iilal. depends o.n a setof N ~]iddeH 1!<Li)C~s":."• _".• ('M oom:sgJoluHng 10 the lV d,"~[II. .•

Art.1rlitl' p.LIDumgnt~,on

I7"1

A.

----

-M"dy~

l1li ~.
-

tlO',t!OO !!!<m'1
AliJUm'~

cb:I rq

<I F.1$D~r;t;·~~

'!I

lfii!ID.3. De'te"(or!g geliles.. Affinity Il[qlagalioo, w.1:5 medto, de!.«t IPUl:aIi- eKOM (dar.a points} OOFlll... e ~5in91 '£!eiles 'rroml _u~, dI~®TI1m®mei. Heile, 5G,lJliIred ~.rlor is rUJl ~~r,o~ate ;15 a me~~ r~ ,~r

similarity.

butiras,tli!'<IId simihulty vJ3ilu@'5were


II'!'1eMI!ui

,m

,deriiw-dl JrClJillI O!I <t;~t; fu.nJ;!ilo!l

llhe puti31tirw e.xmas, in !he

s ,ea,dh (aseo.• affilil1ilY pM~agalio'n <liClnieve<lal s.i~m{icailtt~· la\lIEH recoll'hstruc.tion enorr~'hi3r1i k·clll1~ers IfUm.- I'lc gruuJx.'.dlOg~tflCr b~ ,m ~untior~~ma!(! cd.llsterirlg" ~() A. plot o~lfiue,p'OJlil~w rale \lie'rstU'sralse--r>O:si~i'\!'e rate ll)ir de~etti!n:ge.x~m~ [Uls-ill 9 IlallJ;ell:l; ql!CIlCC flf r'<lll"\\lisc grouWiilll!~, In Pl'j·, it wo.s ~ho\\'nUlm 1hc ]'Ic.l'~njcdmClrk 'from RleJSeq (5)] slto'm: that .alfilmlly Ipr.[)pag!lltiofl .a!:so'~e'rr\[)rms be'~tef al. detHtiing blior!Jg~(aUy verified ,e;I:_(II!lS thalil k=(!?,flll:!n d.l!5terin!1!. k-me~ia~1 pl'ObJc:ri!l C"OU III be rc:1 l'i,:.. ed IiO [oran a

~PJ~ion @~ lh~ plllt~~ l'>i!mP.l~~(3')" W Pi. smaU IXlliilio:n thE! data and llilei:!l"!'!ergm!::e !if dUSrels, dun ng ellch ]tcJ<llilln ~r a~fiin;ty plIOpag"liaJ'l, arreSilhTI, 1m ea,diJ ptd~re. 1h~100' i;!0):~ It;Iu~lii!~~FI il;Jl.iIck (l;Jmr~p'OIiI~ ]00 to data ~ililtJj "\rom a 'lOll'll ®F ,:5,0.606 plIta'tiw eXGi1s1 ami lIl1e 12: (otGfeiJ I:iloc:I\:s 1111 'ii!<!.cclil iJexillidiGlt.\'! the If<!!1~rip:tiOIil ~eh ,(d~he C()rlr,e'51l0Ll'I(ijilil'~DNA S<egJmentin n.2~nsS\! e samp:tes. ~be 00,;': OJll tlile far L~l ,,:orli"e~p(lilld~ t® ail .ani~kial dat~ ~(llllltwitnIIlHil~~e pre'rerelflcf thaJti~ us&! '~(l arc<CIiI.mt fm ilOn.e>:.<CII'I rlil'gigi!'l~ [e.,g" i[lJlr'~III:!), lliFII~~ (;Q!'IJ1 J!.ldil~,g d'~t,ji p@ili!lliil[lJgi~,;Ill: Prt;ltrf1lti~l ~:;.~i!J!IIIITl,~[lJts, wli!'~~ gr,~i~ t~i"'i.es~,n,dkale a~5.ignmen;1S thatcuf,relliliy have \\lMk e\lild~nce<l!llci ~ollirlliJn:es il1,ciil!:.iltE! a~5t9l'ii1lUfiilS 'li~alt c!I!nenUy h<!'!fe :5U®!llg evid@rlc.e. (IB) lP'err'o:rmilm:e 0:1'1lIIli!'lill1liz~!l!!i the r,e~onstfl!l![MOn -err®r ,~r gef'les, for !iI~ffeif\f'I~1f],UII'lIIb,m O'f del~cltelli elMs.te~~.Fo[ ea<c,l'llilllFlilber of (lu~t~I':5., .. Hl!liIi·ly pro;p.ag'altioifl i l(;lok ,6 !inhl, where~~ 1(1,000 n,!!li!i '~f k-<J;~~rs \Ius:~ri!lgoo,~d:; 20a I'I@!H"S ,t;l!ilthe ;~lIi!e l;oQilPy~er ...In

genorneand coeiiil;ll1~ a:tmiS'S 12 tiissoo

ng pJ<O'xfmib'

or

:ICIly Ininhnizing dlC CIJ~CfMY i~ 1:;r!llllP!JII~,trOI1!ilL~y illll.mc1..'1hl:e.l)c~~I!".C iii !lp"eial ~~ ~rlhns 1~1hl· iuniz.;~iml pmblo1.n~~hC' NP·hrud k~nledi1L1l ,lffiib. km (<"fJ. I ~.mw\'~I'; I;IP{~lt~ rules f'O!' affinity the 'l.lfi6j.~<'I;!W~kmo~p!.··:m.dI !11 moo"poh'lt ~lIl'S.ron:.1 o ror J'lTinimi:dl.lfl a B~~tim .rl'<C~t'lrM1'!lY (9) '11'1]11'<[11";" immiml'. Afli11li,'Y rmJ)J:galion is most ,(;a_<;;iiy dip· ~i; {:(l ~" 'll:.IIUS!!aliCC ,of alil;.'rIllt\-81'1m ;dg,\'I.riu~u'll .. ilil~, raC:ll:l.l: gf"IWh ,( d~'1:;~.ribing !;:Oti~ll1Iilll;:O; the (~I~ ali...: lfLhch and the ~jlJO.:rllY ftill,;.;tioJ\ (21, h1. ~'>Ui'Di: m;~~'1Ijj;i7JI';: t1l~;>. D1h~'i.:IfIC~J furictioJ'i l11ia~ Im~ ~flldlar~C miual1lt1 wirth (1()~l}OJirlin~ IfnuUj~)I!.: fi)!;.;.:d poiUl~ or Ute Llpdal,.; ru"-:5,lmrl i:lJlC[;;C: ubily iJf\:v,.;m COll,v<C(g~nce. ]i·or c~al]~pl(:;, if ,~ I ,2) = ;~lJ) mad l'( I., I) = ... ·0,;2). ~hclIIUit: S(liu· ~Hm,~ = .('1 = I mad q = {"::ii = l oo1.ht:l.('(il,.;<;t\:lhc t"i ~1~11~ cncrg)',~n li1iJi~ (;'a:;~, ~'I~lli:ity pm,paytlti:on illla)' ~ilhu,,;-, wilh oolhdm:;t, poinis III~CiITI~li]l~ ootWt1~'jJi, beil'lJ;j '~ell~W'ltllfl:l mid IlOJ10iCI1i1!"'bu];. inl, pmctrcCl, Wi~· rOOm! Ih<l1 Q'OCili.<'IrirnlS coo!ld alw-ay:.1 11lC ;IrI,'Gickd !by :tdding ';ii. tbliY ,]Wi1mIJl~ or rmi~"'C' 10 the: ~imi.brriliCl1 to pre\!Cillt (~t"tlCrnle ~itlml,iflm, or by inCn:)<t'Oing 1.ll'C dalnl~ing r.'01(101', I\.mll~!.y I'tfOP'lg.....i!Pl"l 1~1l,,~vernl nd.\'.an" ~1~~l'w~.r~QII:;Iit~'\~'~~~l1Iiqtl~~, M~!hod~ !:4i~h a~k-<c<c!r~e~clju!l.tC'ri!lg {J), k"n~e;lj1;S c;-Ill!:st~!'i~i~ O}, n!!cl~he e....~C~.I~ UOU 1~1.'i~imb'nLi(l~l' IE M , f a~glu'it~lm (!.Il !ilDtIC'~.l'elbliv.cl.y !l!lll<li1l.-:el (If ~~i· li11:ucdd\I~!crQC'lnc~ a.~h ~Ci\ !1lC-~ Ie<hlfliql.l('S ,~[lC' in"ll:'m'C(l UI'iOnl1)' I1lOlhods !h~~ ilcgi1~ '~\~!ha Illrg.t::' mmlbc!' of dU!ltc!:' i1l!nd 111>::'11 pnme t~H.'IJ~ V 2')., I;l!llll:h!!y ~Ii~~rely on [',Jllclm~l i:li~~lipJiJlig tutti ~lIlak<.; hru"·d pn.mi~],g decisions iIll;U ~UIllI1lM be ii\,\.'O\''I.~'tII .11:1_1;1]:1. llil (lolilm:;il,by :;iJ1iIJia1llliCutl:sliy ml~ir:krinlg ~],I d!1l.ilI ptlimro :~ ~IL1tliilin..: «'1h!11:l am.! pwJly it'kmifyiillg d~L"I:f,. ala.1111~lyJrupm" I gHLltm i[i;~bl\i; [ll ;}wili ~fllnly 01' rlrie fJ'DUr so.11'1tiolii' C'dllli:!Otl b.~,' lmhld;;y illitil.dlir&qIliOliJ:>ilUIId hmtl dcci· :>iOJll'i. M!lfkov eha in MOIm: Cm1olochll~q lIC5

I(),

U 3) rmtdmllillY ~n;h ro.r g()<i(j(j rol:lIIjl!ll]~", bm 00 .oo~ slmn: af~illily 'Pt{,jp[!l&ulion"~ :[lclv~lI]iILgc {If roll~iu()[il1!l ~mmy p>O&~ib],.;: wllltiOl1l> !~II 1111 nce. o 1·1 im1Ll.dIOCo ll :i~IDmi:rntu~'~ ... a ChL"L~T~ng,(J <fI and :>pcC'tmi cl:UiSLeri!l!l (t'l) ~h',.;: Ihll qlli.1.C diH:

fe!'en~ prohlc!tl f)lfro01.lrs~\·ely Q()~atJlntil1g pairs; of pt..'in~. nn Jl~!d Piil~1hk~1 of the dm<l!,These ~e-dl50 l1!iquC's dQ HIlt !1CClY IIl.u ~H points wiThin a Ii!: ~h~Si~crll;)~!mihu !() ~ ~~!,!gleC(!1!C.r <U~"N tlnls I

nl\]~w·~II,-s\~ltc<1ID ~nm!)i~sk$.ln p;!.r~kubr. ~wo ' ~ims IImt shou kI nfll, .~ :ii1,lh~ s;;mlC d~i<.'t'lerr

REPORTS
(iN. i~)l, soh'C 11:0111.<1011.1 s~llisr.ilbility r!pplicu (e,g.• .rPLEX). a mUll), PiOJ'IiiI~li~iOilI cooi,ng makes usc or imllld\>c '~'lCSS~!~,C' updmcs U~t c;rur, .~bkms'.','i~~1 :ou .uni'a'.of.rnrlgllilule iocf'l:ulle ill sW;: {Iff}. oohc llisu.mccs of UlC Nl?~l:tlrd lWOlc .. ~11,(UlillCga~U",C. yntmcrcic. s flJ'irl Sl).t~sI.Yhlg thc be imp!errn,;rlllCti ~I1I few lin~'S,~Jf rode {1'j\ a Ilrobj,;"n ,tom, md '~rirurlillc in~u111'ity. ,h~romm~i1" Ililliuil;y propag§lIiou A.IUil1!Llypro~lnl1l is If(,i~IOO ill spiri~In ~cch· dimiilI!.iJoo~:1 r!.'I1'~U'if\~'row.in~ cllJ/;:~nlly i.'SliUlkil,.; d c from. ~Mil-:;()f :iil~1W !Cmi !."Ii\.;.: I!~ irlIJL1L W-lll~'1 1:1I'l1:lrndr,i,e ~Jjlllil~lIltLc,,>, Il,ifLll.¢S 11I.~lIt~y u,'iCd 'L:O obtain m;;x.·~llil~b~il'l,g
Iwgram

linear

\'V,illl

II OQn51fll~l

'~ioJ]. -rlilCit.~illeilnpul

!;1fiO:r !l!ppro.~il1ib:ilWM iI.'ii>l1Jnc:rl'robe rm:tric,

be

)\,Hinily IJro~~galiQII
;;It;:

~~§n pm\ kb UK.:l\'\l npl'kI.'OltoDh Ihat w(ll'h \'I'i:lU iDi

,~, cml.C,q)'tLI!:i!~y
nmLc~jOL~

:SO~VC:1ll1d

~l.c' HIlc:mr pi,'ogrmnl11.i!]~ 1~1a:~lEk'!ll is ~~ph~illiC'~looSDjh~'iIR' p!ll.d;:a~

W.I LtU\' l'tlm I!:l


11 ecd 10,

resulls in qllU!:i: ~Ub't'l]1 diSC'II)lil.~ (Hll The np· IUtI<Lch ot· 1'(lOUDiii\ld:~1 tlll.'Oraga~jing mIJ:I:mgcs (l7')t iiI! :I, "Ioopy gClph" h:"s heeu u!OCd~oall"

proaeh Shannon ~ limi,1 ill

C[f[1r.oom:;cl.il1!,'l

de-

kn::lw.lcdgc. a:llinllty f~mp:I~l1:iDIfl It': Lhe ~b;t1l1L1hOOLt)msko Ul'it' onln", itlc-a Ib :m.l~'IJ Llro ag_o<oltt .limdb:ll1I~mal l~f(lb~t~lIl of dUI~rC'!'nllg data, B0CnlM of Ill:, si.lI\phdty, general app'J!icabil:il}'. and pe~rhl;un~cc. we bdJicvc 1lrHin.11:J~ .!o OUF til:, i~oc:c

(J?t 'lei,

,~t~,

iIEliilQmp!a!"

unlMCllt:

from dmft ,01' 'tt1!; p.D:poI':


~lll!:ll

i~ypRlpa:g..1.liDni willilTOVC In be oUmml and Cllg:illC'(U:II~g,.


1l.l!~'l!'ll!i!'I!C{!;~

V,lhJIi

In

~'~!f~r'KII~lrW'~
~ge!i!,~~mgl!~r.g~tii,

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:11.

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J. M:l,~~ii, aii! ~fJt,ttflii'j9$ ,~f PM Fi/lfi B~;;Ultt ~~rum 00' mml.i~iKo' Sitliislio: 11M PllfllHWilily,
E. ~ Qm, 1. Ne)rnan,

,and! ,Mo'I:e,~

bul, aM

lromlbI'mHq~,
If!;';Ii! _

pf~Ufie,

3] 1IIi!o

po!!l1Jvo m&'Jl'mlftlilit!!:!I c.aM1d.l'» IlX~:r k I'KIJtvos: 'from cilMr pcl!DLI. '\'IIJ rtlf dit:lnulfIl nwrnma of dYrum, ~iiO
r~~ ~~uon I~' iIImI!'!i.lI~ byll!1il!li)! I!II!! ~1fl1~dU~OOFinglaro

r.a~ll!i!f!yf(U}~~

,~,IG) is Hi'I '1a<!I1!o ~ ~(! ~!JiI'I,iiI!~'

!iii!. ZIll1.-Z97, i!. iW~ilabli! ,iiii ~il!Mit' OiJUiii~. 3. 5Qfilllllre ~mplml£l1iWIian~ >ilf onftinil!!l'1i~-qiI'l(l~Ir.:.Iil, ,!tOO!) \\,~~ li!!e dara '5'ft~ ~1~l!f!lbrnijf'> '!.lied, 10 Qbt<!!~ tiM resi.!!~ d't!(_rj~ed 1!'!lhri5 III'!~ !W~(Ii!pl, are il~a!lable: al
1'J1l1), ~I. 1,

2.~p,p.ilili

seil1l~ltr, GJ!..

td~, Wftii v; 0'1 (atitJ.'ilmra f~~~

mg m!lti!il~l

...

\\_,~~t(lirooIM;d':W<i~iir!~r~pi!!IiI~o!'!,

'l. ~,J, frey el G'I.:. ~fOt ~et 37. ~':ll (200:5,1, :so ~ D, frlllll, V. TalU5'Oiia, ~, ~ Mil'!lbll, ,~{~tfl~Adas !IA1::, of S-'O.ri£i'i;aj Mltlmlll00900~ ,iI'h:IC"~!iil'JjjI tPlllllF\lles.'5, CilI'Il~Iii~,.1WI, 1~, ,_ ~ l_IIIlPl~!I,.I"lu.;;_!lIM A(oo ~ (JSA '79: 2S5~ (19.~2}_ 8. hi, Cl\;iritar, S, GuI1~ A_ 'T:ard'(]~ 11), ill, Sl1m(ljls:,. f" ((tlrIpul, 5~t. S!;li: ,65, 129 (200!2l''1•. ~ S, 'ftdilii1Jo, W. I. FI~emllf!" 'If. Wf!ii!, [EfE r~~ lrif. JlI~!'il. ZZSl G!(]ffi'. ro.. F. It. IK'!dli!tlJa.;!J, II.. j, F.~_~. IH.-A lOeli~~' e .1lff 11.m.l, Iii/;. ItU~1'lJ)''11, 49.8 ~lOOU, III. A. 1\'. il~~t@r, 1M, i!.Io l:J~rd, [I. 8.. ~lIMti, Proc, I'l, !i<Iil'J. Sar: B 39" '1 (l97n. Il. ~ '~9-~f!R, L l, s:.:!l!~!f!~l1, fM. l(if~ (#Of. UM ~~~n ~~fIrrlal\ Sil,~Fr<!m:h((lI'CI!, .:1;(0))" ~P.]s2--1S9'. l~. ~ I<!!!\ .it M, ~~ .1, 'MJpI#. ftif}P1, SM; ~, '!lSI
(ic

31. 3'1 (:20,m3l. c. D, IlI;imrring, H_ :5d1uUE. ~~

C:

IA. ~" !,:, :s.J._tI!~ D, ~he!"!~r; ,Unllo:, ,!f@s, ~I. 6l!~!L3;6, lAilI9 (U~ro, 1'5.. ~5hi, ]1, Maljk, lEU I~I'JS. ~~tffi'l Ailor.. m~t;Ii" rnr-erl ,22,
I~

~~l

c.

a611: ~~(!!OO}, 16. ,hi. ~~rd, $<ie~


17 _

~~..:.;_.
!S~~

~iW~ Pi>i)$lbi,!j'~rif" 1fe,;:J~ Iirl lmeI~~ $~~ ~N'KIrg\ll11 Jr.alJl'rnan, $;1m 1'IIiI~1I, CI\ l~aOOo. HI. D.]. (, ~fKa~, rEfE ifimlt. lilf.. IliI!o._'Y 4,5,3 9'" 1:l9~', 1'1. C. BeFloo, A. ~iWieiW;, tfEf ltllh.!, (Qmrmr,u', ~4'1, 1261
(:t'n6),

300,

lti'SS, r;200~),

a
Fig .. 4. II!1~liItMy~ 1119~r~ ~iFIl'ent(€:'land'alir·Ua1.l~1, ~ rOlUlnl[l. AUilll'il)l' pttl;paglfiltiolll (.!lin tJ~· liIS'ed~)oexplore 'Ihe 'idenliHcatMm o~ eXBmp1.a~s 011,the- b.,15is o~ n'OIilSl:alildi311'd ",plfim~zali~i'I cJil~rl~" (A) Sim1tan'ties I>H\lIeel'l ~ijlirs of se;nt~JlJ'll'lji ~Il ~ dn.dt ,~rU1js IIIiIgnusQJiipt 1i'I'~r'e ';llri~U!u;t~ by mJll;chlln:g \!,illl'rds, f@ul e:l:emt;l~' 'senl'eInC!E!s~'I\ere iderilMIi~ by a(6nily ~r'CJJlIlI~<lll0'f11 ail'l,dlam £ho~'1,nl, '(8) .A;~fii1ilYpro:pagaliolfl w.a~ "1l~;OO to 'sirni:L:l,riitjll';Sd'E!rWed from <lir=tr~,.,-el dfidem:y (me<lSur~' ibyes.timO!ted u<!wllime) ~elll,le@r! the !1!56 ~usie;sl mmmeu:lata~rp~m~, 1mCanada arndll1e Urn~~ed:S!ta~eS'=tilelra:",el tlIii11I!5~'Or ootil dk~d.ili9Jnt5 (s~OIW 11'1 bl~I~} .iI~dliPl~irm fUi3~tJi(nQt S'~QWI1I),i~dl;ltJi!!l!J tl:!e ITlJX!n lro1lni!iffor li'fl'];~ ,~r w' ;;I m@:l;~fI1!J!1ifl Q'~ Qn ~ '~iP 's~p!i)wr. w,ere ll5iel"l ~ n~tWe 5~m~i1Iri~;~5 ). (C)Seve.n eJ«lmplaJfs 'irJ e'lltilieti i¥y alftini~p~CI!pag,n;on, are a 0 (:dloir",OOled, i31r1dth;E!lIs;si~ n;rn.ents of other cities tD .~ ~ ,e-~"!!l1illlarsi~ s41Dwn, Cities klc,at~d <Il!!ul'E!!le~no, ,exempla_r (,ltle5F1lilY IJf memliers, of O'M1erm,O:f-e ~i5tafl~ ,exe-~mp'la~due 00 lIiI,e lack. or dn~e(t.mglill5 bHween lhern (e,g" At~liItl!l; CiW ~:J 100 km rl'm PMI;:Jdetphi~ bt.11~;is.closer h1!ltilgnltimew AtI(!!1~~, ~j))Thili!i!~t :d'ilO1llSloot 1f11lCania:da·IilSti.bmiletr rolUg~ll:l dMdes tn~ lomnWafld Phil.ildelph~a duslNs" due to a larger av.ai!abitity of dOlilll!l'Jlic f~igh~ ,otl!mpared ttl! inl'emaliooa\ m~hlS. Hewe'l!'ef, tl1is ijs not tlhe case on the \lJeSl
roast a~ 'ltI'OO.n1in (E), IJEiraiJ~~ ,e'XlI'aorli!lirnar~ly Irreoq)llent airl'iFle'se,r\Ilce'
i(;oiIli!1ced:S,

.2fl• .rtI, l!M<oor!l. (i..1i'.m1i, R, jleliillil'lll, S6IM:G? ,jI;\I:I'.1I'1,2:~2002) • .2:L B~], Fri!'j:, R, KO;!1ttf. N, 1't!'Iiroo.ott, iii iI'tO~, J~rh OJrl/. NJP:S, ~ III're!~Q~r~tI~, IAA" 200<11, )J,ji, 73:jI-jI'U ~."l. T. .N'!!:It£e~. C, 'i'il!1Q'<ef. V. Weios, !R Pro", lOl!i1 C~}: .ratt (I EE~ ('~llliUlefSodiel\i' !'~5I. 1.05 Ali!1]l~_~iI, Ot. 200&1. pp, ~:2a,-.q3S, We lI1ari~"B,~U)[!'!Bn, (;. Hint'lA lit ~eilef. r, ~eCufl, s" ~5., ~!1d111. Wt~ 'for !1t1prLII ,~!mJi~O!'l~ ~rld ' f _DB'J'.an, Gc HiIlIIlfl> I)'_ iMaci(it'jI'. ,pj'j_ 1Mi~\!.ard, :5, R1l!ll1!L:;, amd CTo:malii i0[ ,oommeruk am iii fiii!ii1Ju~ drall dllflil1'Il\lll~5;(IIi.JI't \'Ill! adm,IlII.tl!~,e hmdill!l! fllOOl thtur.al 5Cie.D~ and &1gil1eeri~1 E~KIiI 0Jum:i1. Di Camdl<l, GmIlf1l.e (all.a!WOmtarig (,momiQ; IIi1~ij~l![teo. ami tlit1! Cilnadjan ~rt5.i1utes (}11-1(o,alll1I!.es~r~h, E:JF., ~s; a IFelfu'll' ~. 'iJl~,c"n.l!liiilni II'I!lilIll~ M!fallceiil ~f(m, fl'lf

S!PJ)porti!l];gl O.n'line i!lt<!l!eriO!~l


~.'il:ft;rw;~gJi)rg!lgl1wrt~I:I~I!Wllt SOU! '~:!I ~i!l:il(iQlOC~

IFiSl~~1 t'll 53
Il!Eire~en(e!; ,21';. 'Od:dioi!r 2006; ~f(~p~
IIlil'liMIBI 00 Iile 11
,2',6

Mfi'lN.n1 Vaillmlwer

<lilt!

Ca~,');ditm d~esin

·~helroortlwlest t~ Seattle,

Se,al!tte

l~oo<ay 2001,
~tllJ

Iil~m~~,f

2006

ro.l:l~.ie1liOl!.l13BJmII! II"dUo!!ei!J~ l!1h:nllatij~

ciil!~~h!s 1~!lItf.

16 FEBRUARY 2007

VOL 315

SClIIENIOE

\I'V'IM;V.sdencemag.orgl

REFORTS

IHe've rsii IbIIe Co nca rta d Lii,g a nd S:ubsliitUition al AI'terlDlaling 1M I Sites in an Extended Solid eta
Wt n~pllrrl a ~ntliu~tl!.: material~ iC(!l2(bi(~h(SO\qMHiO)2ll~Diipy)(CiH~OIH~, {1" wlflere hlp~'" 4,4'~ l!ii~~IfId:yO' tlil(ilt (OIll;airllS; dilS!crete re''Uli",e andl ~nE!rt str~dllliral motiJ'5Iil'dll iLlndetrg:o (i rever5iliL~ 'Slu:lJ~ti~u[ic:n1re,lIt[onin'ivi)l\1i 119 :tIlI'e rnrl~er~~!l!I .a ad SPMi\!i~ly (onl~rolled ~9~rodlJ('lic:f'I off biipYfi dilil'~ and meUll~lnO!llfl1(1'le(lIJt~'5 the '~~di;vE!' ~il'€5..Tlhi$ ~'l;!a(tioll! 'd~ri'rl!~~ths pl)~e gec:mf!hv 'Df 'U1~InE!SllIlltillg M ,opel'!~'fIam@wofk s;IJmcih!re<·<lJ!!deentrels the i'lliHmer]1'I which this strIJct.!Jre SlJ,~bssrnail mo~eC:l!ll's" lih,e' mQI.eC:l!lesi"vo~v~dilil~he re:a:ctio'n <Ire '[)O:Sitiol'led Iby <!I1il amnJ' of \lIe!l-defililedl inter<l!dio'lils duri:l'Ig lh~r patti to bi!'l!ding til the metal Cl!llliers. ~~'ill.di;"ill crySuLlI~ill';; ~ol~dl'l!rmy::> b'1.'lCiIJ OU ; •• t~~C' Ct'Iti:!tdJrJlOlho:lll {) f HIU~ ui(kiilroic (ltgmlic .' ·lli~"I!Iuli$ to inc~f.il eeifiL~r.:: lIlH~prOO~lccd LllltJl'~lnl slnilC'LLlm'l(k... ilbi~i'[)i' I,'), inch.li i]l~!';lruC. '[(1[9], (1- i~) au~ ri.IllCli(ma~ (9, Hn ~IlSCS 10' g!JCtn 5Orp~ion'iamll\;nctu~ iiy ix.'1VoM,.-i':IJUClWUrk fUlcl ~LJC:s1 spc'ie;s al. Jlt.. .AddJI.tu.m~~y.cl",~~o:C (Jr· gall~C oo~ids C~~lImlO!;;I!;O ~lifuci vI;; mmiuu~ 10 'ijlN.:nJll it n't~llc(;li!Ilar IlI~UlI:;I:MI[Q.tl:lliu\l~ll Iml]p'lJmlL~ ;:iO~Ltbu.n h~Jf!:,mjc linkcc;. q;lUI,1L~ rulllhil1",'d wi~1ii tlJ.~~lnic l:iW]I:iI1> irl th'l: OOnt'im..ctiOlllof n~":>~
I f ~I.ch C'X~I;'nded i;)lff(1YS

E:.;

Coo~'dhl[1IiOIl iIIl~hcsc Co( U) cenicrs is QeDa~Jr:;]l. with Ihu!: hipyril~NI. 1igand:.l in the flhl~l~ 0 r Ih(l grid! .mdl IWi) :!;ill~ f"Jf~deriv,~1 0 :Il.OI~ls oc(:t1n}~1 illg ~l~ ;Lpk..~sites; "[,h~ SO~~~ IDIIo~em~!lC(:l!hC' S{~~an:l ,g,!'ids m~hiJ' :iiocmld ~1X\I.cWm:l L1!]it: .. 1D ('"o~bi~~l chain lFi:g_ L A and B}. i,n wh lell [I1JCA~1 <;mnxliuqion is; also (lCm~IC<llili, wi~h N mom~ from, the twc'l'dlflin-

ssien l['o;2(bip)ib(S04hn~:!OhJ, "~'iUt OOI~1I inow III[C i:md orga n~c I'i~ n'lCwork=dli!~il'ij big ligand .. (Filg. ,~~ :I~I'S~ SUlICLlI~t !lIni~ is 3 COlbiip'yh Sl1lillilrcgrid '!hlM is dc:iincd by IWiI of drlc Ihru~ tl)"mIl1lCII.T·~JxlC'f1lCn(kmt lIli P.Y moleCIII~Cl; L~:m!. OO'IN1itLlH~ th~ ov'L'r;:I11 ImIJII~""!1Jf"~_ This ~wcl'(linmll~iClJ~11 l2D) .!lriid stacks :dl:lll!!!l tlJUll 001 dDr~c[ioll in an ABA 1~~hion. vl'.ulh alt(:l'IIl~1(' gr'id~ (lff~c~ hy % b, where iJ b. the repeal. d i81~m;e along Ihe J <IX~ (Jig- S I) U 5)-

kms 10 g:h'c a ~liYv'rid rrarllC'work

of compe-

n,;;

,III

oontliliom;. [\'';;\JI~~-d in

~)il d; bkd;·shull!CU iCl)'S1rfil::. 0:1,' 'I 1111~t ~\'~TIII ,,\<Cdk.s, Sim(;'llml~ IIIn~ly::.;i:,: hy ::;j I~bci)'~,t'il X-il'iiy d iITtitiCliUll HL ljJ~ K [c\i~ led that ~is COtnl~l,j;''1.Icl u rt".... dilltilur::t U'ICliI~~'¥lIIll ilt e MI1ICI.lmJI lHII t;,; Ih~LI are eross-liuked imo 1.1mi~'(: t1il:llcnlSiuns ,,;i:1 bidemate hrid~irlg SU/- an-

ml~l~woLrunl::; ...,'ilDCr' i.mu:> 0 atOlnllS, Irun, the brld~in'ig S04,l'" tli'lkiJ'i:> l{jni,'iili~ lh~ iJltct~ll, eomdimuio]] ,~phete, Tlte Ch9[1'I~ Ihus ht.: ou,,: nnoICCt~ki> .mJ
IWO

Ibmli1~g,bipyrJilyllligmu:li:.

s.qllml:

:lIIiuns

gr~d 10 ~.,.oh.::r \' ill Iw{/!b:ridgi'll!J, s.oi~lF:i:.!!" an. The key clifi[~riee b(.'11.wC!..:oJ~

.vllmc!'i;:l~ ( 1'4},

afC~O

I~

U:01Cd ~lu!.1h(!r !P din;ct nlUII.ipJ,i:!' ~1Il;IU I'IlDh;lC\I.il'$ cfKlipc:rnlh,'cly :il1. Cllcmiqll reroc:livity and nanspon in 0). 1n.!I,mcr 1\,'1'11illi~111 nf ~lliXynl~:;:. tll~)I w(llilll pmb<lbJ)' rcql!lirc OOI11!1iI.e:,x "~}'IMl1il'ClriC ~I'!,i'lg 10 M~o.l~IqKl<1:LJIJ; !!!1~ ~Ci)!,,~(I:!)' ~1~,!~~1r:)' wi~hb~ !h~ ~dJi~L~ll,tl;: ~hflowd!~~ ~hC' crysl,LUine

rWo

:5£iHd ICll2l.h~I»'h,S.(], .h{~hOhl~billy)(C]'~lo.ll} . whl!l:c' b"~yll.4'-hlllyridy.U h!l.~ ,U!1CI:ctC ~(:lij\,C' and it'K'!1 ~'!llI.c~~W~~ !1I!o!if.<:: IhJlIUndClfgo

.. reversiblc- ~lh~liMili:flll re".1Cl ron Ihat ~'1\'olves the 'COltOOll'XI "nd sp(l.tiJII~y mdorod ~m~odl,lic!i(m <0 f!woQ dJnt'WC'lu ~lDlb;:\lk~ <114 !,l;'<1c!ive shes. tbe T!lit-;, ,.mJ!.Xss is m:C(~ntKltlat~'dwiah Ihe 1"\:~i.:iIl)O ~ion of lIo.l~.Hlnl~ O'llkr IlYllt,tlJlICr1J:ti'o'e $IrucUJil,1'I1 n.:![rrulll~~~ll~~ll~';wilil in bo.1h nltl~i fs, TIlIl: oollmulof nilL!'reatel iolil by liI(!: 8lm~Uul'C of .~ eau 00 pree~cly unrl'cr.sIOO!:I fuI('C:JUS~ or Ilre ;:;[rlg~(X':ty51n11 rnmmx: of h(Jail n."il,g(.~n~m:~ plt){lm,1, T.h~ :C11I:;;:rii grouws in lhe ~{Ib::;ln[L1[ion i\;UCliOIJ III!; are locilrod in Iht:ir ]l1,"C-fl."iocliolJ1~~livm in] by a ,cli\, ~'ffiC=d)' or \'1>'t1a'k :il lk·tl1Lcti 01]:>. The :rL.."'>I.J.i1,. il1!:lllllll!k.Til~l.~I". porous, wilh ill pcim"~ly tkJ'ill"'~ is. :;,trucTL~ranj'~ml liiJn.c~i~ln~lly by 1111;) ~opo~~gy or '~hc ~1IIbi;liluliolln:ar.lioll. A OOlllmlllbt:_!!'C<tCI.i(m oFlllctRiUlQlk ,wh!lkilL~ oOr (:'(I(S.O~HlhO ami <I,4'~b"'l}'Widim:: wi~~ a mo:!alllig.1nd 1';I,tio' (i.i) ~lndCil"~allhte!~i

or I,a~

I[)~a.rtm>!!nt 'oi ~bffili!;tI)'~ ~1li'o'a~i111' if Ll~~p!!'DI. Lwer( pool, L69 7ZP. luIK. ~S)'Drl.luQmll' R~lliatiii)1l5!)~r(e, ~~~mdl fm !he (en1!r~l LabO~illlla)! oi Ille' IRej,fan::1I l~lJn(~h 'l)a~eiOO l.aooralory, Warrinlllol'l WM >lIA!O, UK. I)'

(0) E8ch eUra·fra!llll,ework

IFrug. :J!. (II.)' Cioardilrl.ati,on abo!Jl title two sUUIlJ!Jlralty di ~till( t (0 ce:oI1ll:e ill,wbicl'liare I;in~ed by the rsin i!lliI'KlIfl. The metal M~(iall'tld ~~~lIlthe "!!Q!lhcife-'9Fii1 iFlet1.miIr!o;: l~ghl blIiie allil:lll~I'I<lt ,aI5:soC:lated .1t5 witlll Ule :IlID rcha'iifls is pil,rpte. 0 hOIlil tille ~iilllki[ft91SOi-allllllClJ'la'liid the \'lCil:er b>OiJi'ld'~o tlil,e chain m~1;Qlsites ~re red; 5, YI!1\1![U'I; C, '!lriliy; N, (b rk lbll!ll~.{B} llh~ ~:d~~d~dstr~d~lr<c of II ViflWt£i ,i;j ~o:nQ the 1:00 directioll .. Tinebi~y lig'aJlild~ f ~Oiill ttle s~l!a re ~rnd~<Ire li~h1lb1l[.l:ei1llild~hio:\.e ~roQm Ule ,cjtlains alr,e 1i1Il!lrll~e, mp'ha.,lzin'g the rel.<!lijol'! b,et!\leeDl~he< two dhtind; e'OllllpO:rle,llts or lin,e strl.!(tulr,e' linked e i:ly tlil:e SO~2- i31li1i'Cii'l5 yello\lt; 0, red}. Ihe dlafl1<5, pass tilro !lIg~ tile gri!ll~at alFl ~liIg le tOf 1~9'.J~. AU {S, .111tQI'!'1o> 1!!1d 9,!,J:e~t~~~,di~~ ~r~ il;limit~dllQr d;'!ri~f'(l~~!;!~~ii!'l~~'~pe',fll!?lr~lil;mq'~hni~s:J;rll.lI;~II!IFi:ll!ir!Qtif ~~ ~ seen in liionporOllS ~(lI~Ulpeh(SO~h(H~Ohl2t:1ilO, wtlerre bpe i'5 tf(lI'1.f~1.,,~l.tl~s-t4·py,ridY~)ilthil'!ei'lie (20}, (() The e-xtH!4I1'amewo:rk bi~.~ri:dilll,e:s (darlk bllle;) form <I 2D' hydrogm·b®lIIdeodllayw byll1lter· acth:tn wilh tHe Co ~lJipy}!(lIlla.ilm~(",~el,\ie!1 atoiil[l tlie :U)ll !lIt~N:t~®ifI.). 1111 locaitiofl of tlte eha nne Is, e 1J'C:~L1p:iedbyCIFi~Otl !'J!Q!eI;!JItes~!1 i, i~Ri!@r~~d by the giF'~en dr!!;ws, j3,ip'yridyls h@f!i! tih~Co(bii,PV)O! squ,a.ITEl grid'l; alne- ~i~ht bi.lJ;e, tho~e frClIill 'the [o(b~!J),,) ~Iha]ns.m~ pLlrp~e.and If! a'tams are omiu~.

sol-

bipyirik:line illl 1 ,c:lose:lyllilt.er.m.~ withlwo

C.e(bJPY) ,dhiarilll:s.• rO'~fI\IingtJ.'1@


glllE!S[S

'To

w'110!~

'~olfe~l;lrude!1(~

1!Jl.~.rllMeiml!f€ilil!'.,(lt·,uk

~Iwl<~dbe ~dlclle~~li:d.E·~iIi~

l'I~tI'rngen loo'IiI:llI~. ~da~tied yeLWw

UnE!'~}00,til e' m~a!l·boaJmdl water DiI@le,~liIlles.lhE!'{~lOIH ' I'le!!dir. posi~iQn Illy hydrog~f!I bondi!'l~ t(l~he< fr~lIi!lework4'Or!mjl'l!1! SO'!2- "milJfl5-

are

REPORTS
'~itC' sceoud

Uu.~[WO' il;~n);Cttmlll units is n~n lIJ~ chains in nil it COJlWIn Sti'bS~luniormi~Y0Jcti vc


iiIi,ClLmJ: OOIUCr.;,.

The >C'x~cmjcd I'r:m~'[~c .....o;r!.;.of ~ ~u!..~ml

3D pore ::1W1xCCurc (OJ'I'Ipo~d I~Qli! voJds olf Y~5 A3 GiUlcrep:rrl.'Seut.1lO'Y~ of G~tCliIrliL~cll YO~,

or

(}lioZifI

l.!ul'IC. A~ 29~ K.P1Uliill Jcoolvat~OIl "1)'~u'LIsof ~ pjrwc:> one ('I ~\llOU IlilU~OClilc flnd one c..\~ru·
li!iHD.::W{)ik.
SP,I,C¢

or

bipyridy~~vg8nd Ihu. are not ooordii::'li'a.~i:l!inl~W{iln.

1C' mad "1$. S2) tJ 5). nio: n.t..lJlOO" ordi Imloo billyddyl ~~galld OCCI!IP~CS hydlu~n.. 'lxmdDcl poJi.ll.ioTi (N._.O .2J~(l9A) I J6} .~lllCr.mtiJlg wi~h th~ mN:,J1·1>oulld \\I aier 11IIoloculm; lim1ll two nClghlllOtFing Co(bipy) .rll.'lilll,~ tfig_ U)l to form hyclrngiCI1~oom:lmI byceJ: Wig.. 1(') wit~l:in
Wig, IJIIJ

nUlled to die n'lC'l<lllccfl!~1S in this

3D!

,I

rrmllC\'\IO~..

'Illis ill1C'mc~ioll p!aCClllhc

bipyl'id ill1;: 1Il1oliocLlIG' ~Iw :saC{md,I~' oo()f1tjilmin I.ion Spll~T"(,)'o.f Ih~ 1]1>t;:1i!l~ ~Dimk~, in tI~ dmin COlllpKl~lCU~ dm ~tJlt'Clllire. The pooo~liml of ihe CII,:lOi.1 gUC5.I. is ,!IfISfl~!I:}'~U1].1oglIT,Ilhic.illy ~'Ii:llill dc:Fin~ U:'i£!- ID~ 'bcC'alll~t;l orhydmg(C!.~ bQlldinJll ((l Ihe- SO}:' nl1KIIl 10, __ ::U(]~ A}i:iud ;I! uon0

"r

Fi~. 2. The CO~biipy') ,c:ha~IH nil 1 lInd'Jwgoa ~e\IIe'~5litDre !Diml)'I,e(u~.1ir wb$~ituj~i'(!!1lI rfc'i3iClio'lii lIPQn h~aUIiI'!J1 to 368 IK, im~ DiiP1fMirn~ ,]I.n.\I CI1l~,OH glu~s.1 l1fI(lle(IJliE!~ (l'I~rlp'I~) OC(Wp,y hydrogen-itKi,ndi [19 pos~tIO:niS 1m ~he chanlflels Ol~ ~oL\I'ated I" 'T.1li~ tnlcomll'lg9i1J:e5l :species displla,ce m:eMl-iJC!lIilfldlwater !i!1JG!1 he~til!lg, 'ID Ilroduce ac:hilli!ll thal has ili pyJid'il!le~!'Id CH;JO:HIbound to aJloo rna 00 meti:!lce!,!~e!~:S .a~j)lf'Igthe ,cha~n in 1'. WIiI,en :lIl," is (Do,ied back to 29:8 K ,a,lIIdi ~XPO""M Lo the lalbora~()ryi3llmOsph,er~, W,]lt~1Fls li1J~~1II up ~o !!ii~@ ,i1J1F~h,~dlr,]lwd phas@, ]t''''; th@:wi1JlliI:lr d~iS~_;l(@'S(h~5UIb:sltitlJil@d ~i~;llnds, at Ull~ dli1il~nsiite5 in :m' to r~tu rn tlh,e ShU(~U relo that 01' 1.1L", and not 1" liS !J sed 00 demo'te' the' If'ehydlraled 1iIlt!~e.-ili!!L lt~oUllJ'h th<l! eKliril!·halmework lig~T1lch in '1I!"a re !fUJI ~1J,!'UjI!l\r A bo~nd lQI the (e(ilipy) (h'1!lirt5,tlh,ui5 demoflMt~atirtgilhe rewrs'i'hHity ~f ttie's'uiD:!:tihltiOn rM(ti~IiI,. tll'te s:ol\1iel1~c:ompC!sltio:nl ~1'I~1'IE'1 ~Iloalnnels will de~ ~I'fb~' difhl;n~li!t from th~ Ip;JI,r~li!ls(j!llWlt~d phi:!lSA!1, hsn 00'thl~' di)tin\tiioifl bdw~~!11~~~IIP~S@lS t is ,ma!!i~, iI'n!2' in5gt pic~u ~ ~h{]w ilow Utt! tO~[1If or tile (Irysl_al d~ln.g~~ as UII~ SlIb~titUi~iIDrIpHI~$S ii!Wllt@~,

m._,~,l:.2,TG[i AJ. nae


~hu... ~J~c~clyl~l~d: ;
1(\1'~ hour. "lie dill:nlclil'lg
t:llli ~'.

cli:LI!~.ska! bllC!l'(I.Cli:on wiu!huilC' hi I)),lridy! f' U


IwO lUCS!

:nw.[iccllics

fL(C

try rnjnco'(",iclilti]i,,~"t'Le·

~Hm~ willli, the IrulllCW(lrk. '\Mh"iJ'ia :>iJlglc ~l)'~1

on .\',~~ I(~ 36:-; K hC'liJOO

cokiirr rrOl~lIpiJlk ro purple, 'w~!I1{lUI th~ ~05:;; or l:'lj'slullinsly olt p(J~"'Cr. GO afloR! til..: tk'S.oIv;Iloo a'l'il::i!.c· D":Ic:m'liriil~io:rl o f' th~ $lltUlMII£ of ~'Uil }G.1'l K (~111 C.:\P~TiJlli~JtL Itpe'fllcll wilh OCleULic~11 1i1;;;MIH~UIII tlu~c tli Ir\:R,11~,!;:1)";;Jtalb;) r''-'''''(;<I~OO Ilm~

C!)"iSt11 ~ chmJ~1

dlH.: iwrJ' gl'lcsi ll:.tlk'Cll:I,,;::;~vcl\:int:OllF~IlI'<lh;tI inio ~rlC cxl.Cmkd. .I'ram~'w(.d it~lf. w ~ivc a rmiilc~Hnt~Xlnl~}Q~lim~ OrrCOi;i{l'ilpy~(S04l2.(CIIPUl'l

rOJ: I", :n~is~~and ~bSlittlltkm oocu.r~J;,'(1 :;loMym


the Inl;:!~'], C'C11tt:;ffi. :il1. Ihe: ellaill ()Ompotlcilt .of I, when;: nihe-Iw{I l[Ilhj'I,C'm~aHlI'll)I nd w,~nur' lig.1:11ds We!)t)l' ,~isp'J;jJCQd by the- two hl.Qm~]lin~ li,~;m~ls,in {I sp.;!!hll" '~I~!<twd ~la!~!~~r m 11~u!l:hbf.l~inJll

~u'<1"l 'ccnto()~, 111i)1 ~!ilI;r.;~i~lLtioll ~;)!C~ km CLt:'<I~I<{~ .. C:O dl:o'liTi il~whkhtwo CJ h.ol i .md ~~IJ.lO bil1!lis $~Ib$l.illl!imi of e-ilhcw'~\Wl hirnyridil'l(: (Jr'
~WQ

Co

'l~y~idiB~cl])ol;;x:tdes were bound to <I11e~~;)~in~ OOl!C'I~ "Irnlg ~h~chain Wo gi\'C' III (fig, 1),.

ClhOH U10ilccuks <II emch tncta! eceter ~s

driV~'n by c<i>!)~cn1l1iYLty 1:!~'IW\l"'1l.Ihll: inKu.nIl i1lig gruup:;runrl n'lc j nnpo;,''K"!I :;ttuC1I1iI111 :ktllmlOll on n 'drl!.~chain, 11l!.: cXmL-lruail\:\\ 'OrnbqJyridillc :1 i!;arlitb must I1IIDVI! by - 2,:>1: A lID hilluli IOtl'J.OC (~f 1:1:Ic I.w~ m~al.c:.;\ nliCli1~ t'lClll,~n;.which mSlruys
[be :S;}'1lIll1I~lrit ... ~ cnvi:rGI:m]CI:H of[ilc bip'ym.tiinc 1l11lllccl!lk: It<; i~ moves fmll.l i:~u)~ilinll nliclway 'billw",'1(m ~'IN<D ,,;hniJliS,[0, ~'[).';I1CLUk~ 11 w,~.1.C1r nun lu!.:ILI~":•.n

one chain

ml~ml

site,

ClXlJ:KTiltiv..: '~1I!..~r-

pj,md~c~I:IHrlm]iliD~1Dj'DlltighliID;rilJg

1:!~P'J[icliIJC

ner ll:lslLlbS'tk~wt.eialbil.of!wa~ef Ili.gl~J1Id5t ~;itf(iOi(bIIPlf) 'c'ilann 511ntile tra,lils:(ofimatiawlhom a 1Ill.O,1; '!vie<Woo dQSi~ tQ thll: 101 dh~liof'!), ll1w l!;lipry.ridiln:~ tlll~l w!.l~i!llniil!lly hyd~<l>gl~f'I-I\!!Q:J1;;!~g ~rmfT1JiMriC<ilUy Ii!1Il1~l I( 15), TIle coo!1di~m~ed 1l~P1'r.idh~C' ~U W' is hrnII'{~ tc I~'!c JnC'Lalcen~c~, i!'l ~ mOioodemmc 1MhioB 'sei!m. o:n,e ,~f lL'lO in;li~a~ly eql!Ji!,lalent cnailf'lll, slhol\!lfi nn darik bill!! and ~e<d. The extra-r'r.a iilIffi\l'\'D rk (rig, 313 ;Hl{~lig, S,), t lS~ JfI@ !he second ,)f'(Jbi~!{ridliine and C ~.3.0" J1! !Je:sls i!rec:ol.o,red iKc,orl!ii rI'g ~(l whi!:h ~haiin they are ils$oci'£!~edwi~hiPl 1. and male N aiom c:d.eI1~ls i!lioth.:: V(l!id space ID w:hi(lI~he'y a~e lJoLind to im 1". 'The Ilthite II fiI~~alre sho,l,ima 5,,a, gLl~deo to tile e:re to iUll~ua~eU1f l~rt)\'idC' a basicsite. IlIIil~pemjj~!;ld;ar !iJlQ·tiorl Qf' t.wo 11'l!!:i~lh bori~9 bir;!yril!iiJ1e moL~l;g ~e~. I:!I[lJdlt! ~~~~w'W aJIf'iOIW~1l1 il'!dk~le ~ )\si,n ~t. each 1.:lltet:il1gbil,yr.i d illclnllkc~ '~he ~ues;~ m:ol.E1'c!ll~~s that liJlmJ to the chililn,sin '1.'" Pi'£ll.lreS,3 U~) sh.mlJS thE! e'II\()~ilJtion of~'he stmctme a lon-g. t.h~ OO]andllOl directiolls. (B) The el{oilulion o,~ d,e IIl~niliil!)ns OIi the (H~OB and bi,IlVftdinl! (me ,~oo·rdimmC'Ilfl~ld in II~c Irrium~n' ooordJ Ilat!OII sphere of 8 siIlgk' ("0' CilII!OU rolhi:!,~ill~l 91!1f~1molKules: be-l'l\iee'IfI~IO'ii® neig:hb()tllng ,(iha~n~inl tlilf slu'lJstitutlvnl rE!'!IICtIOtllhJ' ~mlllil' rlrom ::11, ~\\'o hydmgcil bonds :i ~ dlC s.::oondmwy iC!\1ordipro1:i!.l1:!119 a(;ha~!1 \'\I'~~h allterl'!<1ti 111;11me'talsil.es. 16 FEBRUARY 2007 VOL 315 SClIIENIOE \I'V'I,MIV.sdencemag.orgl

Fig_ 3. {AJ Tile OI.3QH and e:dra"kO!mework

blp~ridine

g,UI~t

s-p,@de!. 1Ii!@ll\E!in <! (ooper.atiiife mO!!n'

in 'ml,kT~O compkGdy ;f;!lii:liStiu.u..: Two ~V:JIJ.!f nm,k.\(:'Lllll"~,rm dther ~id..: (hf ~ilC elil:JlilJ5 ru ,~13:11'(lflihc Co cCJl1LCIS hlllhc ~cnn~frn;Il~JL1on (mm ~ 10 I' (fi,g_ JA and lig, S3)

1lI1nl.crI!l:I~ laliiJ.:;S, place

REFORTS
umlon splllc~,l)f two eatioas 'I~S, i I,)1; cmn h
pli.":!1C" Sub,i'Ii ~~nio~'l tloOLmd V1"ID,~CY molecules roquire.s ~he hluoducdou of O<nlC ~lll1ljter li:Wl,nd per ra1.eml eenrer: die Gt;Oll gll.c:Slmaicctl~e, T.h,.; C 1130.11 n'J(j!:I,-\iLl'li~" Iflt(,,'~by 4.4~. A. ro a Iforo C'OI1C~.r~l\d ~l,Il:il~hlllliml oj' ~11l ot'the \1;'[[·t('r l.ig:m(lis 'I~ 'Ihe dl:J1:in s.it~ un ~I spat i~IUy (:«~ L'Ckm~d m;lml~[ l.fi!!} 'Ih~ lba1ak~lIIg 0.1' Ih~ :s,)'nml~tl)' or 11m b'!J1}'rid.iu-D C111.\~rormlc!llt ill II plO[lLtoCi'i~~m lroe N fil!lC'llon ~h~~ WCiHklly 1]11.\::,' acts w~l.h 1'111;: nlc~hy II grOllJlS {N _. .H C ], IU jq oftlu: ('[r:J,ou nlol!cqllail oof.!ldil1'l~in:g I(l tiler.! C~lllre!N. .~~Drlj;tCi~m. hnbl&' Tbis (IlremDtioH ~ WOIII~~ ulol lle l)f)Sl'iibl~ becall~ of d~:'*'IIlC~ ~nI'l'i i.I:C!f~ ;!Ilkill;;: ir.:ll ~~'mCl: 'lllO:lcc:ul~ [-q:n1L~n4)d Ib.Cl~n.d 011

or [he

above IiJC ~1liliu ~O' fI! grcmcr '~'i.~~'l1t Ita~n chlrtCr IWd,dyl ring..s in I or ihose rii'l.lJ:S OOO!· t1~mngugto llie TlCigl~oo(iu£!:~ltCI~I~aliOl=Ooulid CO
of ttle

Iit'blyt:8r:n ... P~~51C',~" (fig. 2) lI5),.1 r.~I{I\!il.evcr, 'd


SU:·Cfllli. the eXh"JF I'L'afl!lCWtlrl;: bi.pjil:idini:

SJ!CII1i :1' (Fi~. 4), Thjssteric Ircq14nl:l.Znll.cnu nW~':i!:;may ilil1po:.;es in~lIu,,'alb~!:e on lim lwo IllIlcl~ or Il'IC dmirl bi-

,cJ)'s.t~1 W~ 'C'o()II.!dW 29~ K ini t1'le dry gas d iti nlOO IfCIlI III no lis mig;hml UWSi[~(I;.I:1 ~IPOIi CQfDIliug lnu w.:fll ained IXmIJd 1(1 ihe m~llL~. -nrli":;; dmcre.uc;-<;
~I!i!

illl;rl:lca~C8l!hat I.he p:mt;e;s,~ of ~ig~md movement

:m,.

blih1y n.gl,l~_~1 ~po;ildc~ (IIr a cOIlilf!OIj!Cn~ of the Ir@m()work_ (Iri\'~ Ihe 1:n~l1Iat~rul or IIltc' ;JI~I.IL~mali!!gmrliK!r I>LA'['ON t 171 "":1~.y~i~Qfll r ~~v(',aJr.od;:lJ~ ~hC"l(' sht'!:'l_ C.~!ro·rmrr:!l'wufk VOhlU1C of 14_3'o/it. wi.lh ,u pore ~han (ICJla;J~~y ..'<ull,'\:tiU.iltcd eHmil!l in r. lll<.:lo1mhls Slmct~lro eo.lls.i~l~ng of In ch;l:!lIl]cls wilh Ilarrow !l1m' tl.'Ill~ bo:x:OOll;t ,.~il!l.~tln! ia~~.y pl:lc.kcmd~2';, Th.t~ ~1,hfiliUltklll ,:c-;tCljm~ tmJ1Sfom",~ I~IC naM~ of 111(',ch~iDI from t11m.\v.ilh a~ing'le metal coll~Drictiom; 1Iligned almag tll"'l 01 d,iH:1C~i.nn oom~:rcd 1.0 Ilm:l;c nn ~, and lihe,Y impose <I rilP'" Ihc I)~illl;!: d:islonkm 0:111 arm Qr thc g;ltids 1IH!i~ diC I1rlO$~ Ui~, S6a~1!i 571 (lj). He cross scclionl sile m Ihm. "",ilh al~cl'lu.nillg 1'I1Cl1~1 Siit~, l'he in." creased $t>Cri,~clemmae! 011' ah~ cnatcr~l"Ig bipyridiuc clo;:;cly .~:IJ)l'OO>Chl Il'ie: cha i[l~ (IF:ig. 'I. ~:op kin, IllllTOw~":!t lJoii nit of the dKmnch in ,.' :1'$ 2)11:;: x :Hlj' A. ']11"; nl~~\'ihyl gmtl!, of iIrlCnlcil.l~·boLlll.dI :StrucatjrJ~ n::i~l:I;at~O<nI ixlalll Ihe rcac[uvo,; dmicl ~.I. li~'iuHl" in (l(1I1rl~ari~i[ln wii'h tlmt ,tJ:i'tlilO! u;.:partiul,! wah;r ID[JI\.itlillk: (fi,l;l. $.:1) ,( Ij),~rudlll(x:i; fl~ eloJl." CII"OI I ~\:l~I~(h. ili!.o ti."i!; "Olrl~~ 1,0 we:1lil-;]~ li~jJlI:i~ :>trtIC~m:iii gti,~ OC!I~lrollicli~~, HI~ S5l ilil~li~el. wi,h the L1neoonJIimLL~t! bipy.riuyl N Ih~lt ,l;:fll.iunl oflhelxmd ~nl!.Il1s~~llhc ("(I S~[C ttl 'whid~ OJ") is IhU!~,:requi il'Cd It! 81t.'CI:mIlU't{l(iMl: 1ll1c~lllJ.. iI, (loo;ruimltcs utILI a dlf]~W:: ,of U;.:: S-O-Co ~,~iuu io.n rem,'[~t1'1f1. prevcllIs UII: wlidcnin._g (I r Ih~ da;'lImd" m~he.<;c o~· 'lI1~ water 1];lIJl~· iUlgJlc WnJIlC ~iukgngSlll r~1.C from B5 ..~ in J. m The d i.ft1:ro;;nce umr revealed ne sllll.~mmj~iI puilil~ lIpon'! di$pl~cenll.;lil ~56.3" III ~. (whcJlC!1li tI,lis ,1iUllgic :n:lllllliu$ m Ck'CEWal dcu:;.itji ~u ~h.c cUlllnnc:!ls or .11'~IUj.(l~ K. eotes fl'l1I~l L M!.'fll>LU'CIlfICIUor ('O:l! :S.OIiPli~lU!l ~{_l1'l K onl IJlIlk ::mmp:lt:; l'l'\;hich Ihnn1 widll a 1i1;";. ~. 3.,Q" ill. Ihc I:.;,..:; ·c:ro~vuetl .l:llCllmli,uU-coo.rti il!lalllt1 3 The ::;L1b:slitu!UOITI reaction has thus fbnncd;ll, ;:iUll; in il'). 1J11I: ori!.!IH~JI[Ml.n Qr Ill..: illQ~jmillg bipYfP~):H~!J~ mat,.;riiul wiau:;,1: l D chmm,;;] :;.1I1L101lH'C: i::; dlJC..1:I dUllmt:1 biiJy(;OJ1{)i;.'i1~mli{m1Li (.1Imptll l.:d iatcd idim: is irnp[)5~-d by Ih~ ~lllruc!: grml~}''' ~ilillki,n!l t1i I,"•• ::etl y d~~~.,nil '~y~hc ~~IC.ii :fI~"!C:ic;s,IhiU~ [.(D dl.t:!Ilh.rnc c~y:.[c;jb ;}jjwi:i.iI hen; ~15 H n::".(:~~k1d:[I l'iaN ~ "'~I~h,;d 1:11,.;c;:o;~l"'d-lrt',mi:C'WOrl 5paCC. [hc chain:;, Inlh;; I!rii:ls, ...ila liihun: C- H,•.. O am· Iypc I :ITlnC'lupmt!lIl> i~o1.hi.:m~~!i~,Sg) "5)~m:l :J ta'C,,~ or 2 ..35 a~ld~ 2.43 A nig,. S4) U _il TheThe bimoleeular bindil\g cwnt ni ...dj''lCl:m~ BllIlHI.lI,~A~nln1e~t·TcJler (BET) dcrlVM ~UF~ t'al;:C;Jrea of22 1[3), ufig, W!I~W Ihe- ntIJ'Il~lI;'rin ~lXll~"~,~'(l"l!1.iWC"II.11;!I*' of!l~ pyriid~11(' l~ ~.mU~ I'IlNal ~~nt~rsl\:>-OldiJ!~ ~l"!' 1~is 1lC-'II~hk:_ l\,l'tcr parml!heACS :i n1dkalJ()s the csl ilmlcd 3D ()I r Ihc adj'Lc~ln to the mcl.1~.bi~dillg N .m.re oom;ijdCIII:he cl'}'ll!<l1 was l'Ollllffi'l.'iU [iJt:mll dl.)' N:l 1I1ld a~· l!Ileasl!lre<1 ~w~in~C' <JIro.;:ll. Cnll~istClll!1 willi ~h~t !nwtldm st..lud ·<It mom W!ln~~lIJ:C o\'el1l]igh~ abIY!:!fC.1M' Ih~11ttiI~o~' onhc 11iXlIlom;of l~() ,~ Illa..:;~d \'\I."11J.:r mfit~tdi;:. al)d ~h~~ I:cqLlin;)mCIlI:'~ iiI) :I~bor<Ltoryair" ~h~ ~l.'It<l~ of ;l!L;n1o.~pllCl~ic CKp;x.!Q~ li'Qlll ~hc: ]1~ siz:~ (/I~J, I!ydmg~!~ ~iSOlh~T:jl mC'<l~~I~I~1~11S ;[!~ 77 K !:Ol'\.~ lhQ pyrWy! :rJn~ ~r;lJi.J~"D·to Ih~ .iJloomh~g WOJ~ d~~l~;l~ ·~~c bipyri(ly~ om~ ('!h;OI! li!}md~ il(i(I~!dto !hl)' Coo(biPJ} ch"ill)1 11'10~ \lea hia'l}'ridiuc IQ di~p] ~.CC aJJ!..CfIl~~C'J}' 'l1!clrlw and I!'!;,':\'~al'o/"~I.'!~is I;lPQ~ <re."ilf('l:tioH (I~~ S9~ (1J) b~ ]

Il>'rid:in~ in. m '. mul Il I'CStL~bl in th~ eIoFIL~~. C U •."J I. C eOJlltm:'l<; of C1JlIi\'~kml JL~~!llh wt I:mlh metal ~ilt(5 on 11m ,ehli!] (2:Jn A m l.ha l:Iip~'-IJ(]lI!ld Co and :UG .iI.. m. the 1llC:lh::mo.1· bound Cojl. TIm a!ti:)nmtillg dii~pl~C"Cn.'lCIll. f Ihe o Ilyridil .d~g-li in ihc c:JlllmllQlI cjlhor si:de of !.h~ bipyr.idyl-c'Oon:lil'lalCQ, Co .~.ecf.!fnlil~nte,s: a~~ o-Y
RL CIne In'Cltrr~ CC'll.t~ lIrd

ill!.!!, is {trh'eIJ by IlltHhrnrrmlllly imhmoo CClIl1Pillsili,mm~ ,cllmlg'~ of tim da~mad SIJ;i:lC'i~ :IS ~n8lc '11~f.l..nlCc l'XCC~1l waler ba.ing availof able to (~i~placc IIL'C"uetal-hound Ili r~yr.idilne r in 1~., the Silbsl il~tioll l'C<lc.ti~nn i~ ~rrevcffi.ibh): (/5). TI~c bip'yu.·idy.~ ligand Is lhm; cOll~ml·
alia hi.nd

~h~~mrk cmw~l.~lg

or

Fig. .:1. The b~rdi!lg ,af


(IH~OIf-l anti

I'IlCile<cLtlell (rnghligMlYIin·

bipymdflii'i~

a v~"f)'$i~ll,i,I~,'I.vay t(I~h,~ !'qI'IOf1od ror ~~~U:ilC$. Gif (tlil~l~iliou [Nb(ilh}Yh(NO~l.111 (/91, Cin,~s;i..ent . I.\~I~! Ihe- 1[) 1")G~ SU'UCII!!n;! of 1 h~"'bl1l!, .~;)fI'(lW
cmlsl:rii,(lill)~'l:

;ll!r:mg Ihe

d1~m'el

kllll:~h. 1lle

PI!I~)' M .adj;3.00n.t mEtal :sires olllltie main illl l' makM the MO' p,)!f[Ji!I~1 !.Im~,~ !tie d1afllli-Jomlliill£l bill'Yrid~Qillieq~i.v~~l1t. h ~l ring bound
tDUle Gil W OO(If[J]i'laIei1 D'J ':n<e~ritl1i bipttidine 'is displaiCed aOOv~ ,~Ild bel.®w UQE!'chain, talJ~· irlg poc~fiflg ~nor(ier to bli!~!l the (-H 9rnl.l~ il1dicaltl'd with white
Sl:!ilK'

r.nre~lOC of th!: ""I mc-frra!!1ewrnt biS)}'ridylin II i.., H I1I:Cl""(llI~,i1.C(or 1l11~ InlflPil'lg lx;ii~~VLu:r.a." in

it.-;. ~M!nc~ lh~ 3D po:i~ :SD~hlre I, w{mld L\JJt1~li Ibr 'lIlb;. 10, OCCur. ll:ii.:: ~IJI:nxlut~turrl ofl'b:c o glK~t rllaoJ:octlk~ illl ~ inw IllJ;! fr,1nk~~{Mk in I' i:l:ILL'i dJ::fiJte>IlIIl: ptm: dilllil(''1lsiolili; mid h.t:nre dIe d\:1t1~1(;-rl ililJJ.pli~mnt'Spolli>i..~ TIiJc· ;,;lnliCllIlC or 1 i~, IDdu]m; OC"CllIlll:\(; u lwO
diJ.;tim.'111,'K:I,lIj-O~:g,.1J1IICIIC'Ls{Ill": pcimmy 5111J1";[tlf~II ll.'KlttrS} ;]It' t:onlK't,k:d by II :I]gidill,o~nic lil~kltli". TI~IC cll~:i111l> om UI1:clcrg.Q :'1II1:!¥'~ilutio~~~,t C\!~'I)' cm]:;;li'tLoc:lll. lillL:t~~ t"Cnh:.t, WI1COC11"';Ihc ~rjd~

~r

oc

pre~e(rl~ sterle II:tash'es

dottEd liles. The

'lUre a~Iheoop ~e,ft51i10'1'i'5

~11@t tlrri~ rJ!~~ke!ii Qf [1:g1 d~E!' doiannsindw:,es[I, dis'll),tiO:!1Ilnt ~h.ebilJjridyl


Fig,ure5:S (l5~b .about If~il'metal
It!1l[ ~ IlOW

IOllin .~lmhiUSl :;1;:3{[o~ddmt iGl'L'l!Lllt.;.; Ih~ '~~Itlim~ IlIml illh.!@Illy or Iilm rnm!Jiri~d iis R'Il~,im.'!I. TJIIC dhtincl p:;b.'l:alann of file grkl~\'i ~Ihc ~'IO\\'illg of tl)c' Ik;-;~hk: org,;"lnk biilYrklCl.l. li~lker acOO!llmu" (If @(lrdin~kI!l in ~'!.e (~rn:ig~ rrnC1<!! ~i,~~. TIle Jteaclh ity fu~!t1:dhere (_ic(X.'l1.<r.:;, (:Ill ~he Ijl">."'OI'g"'l~li;,mlk~il

dJLCS ~1C S"!J.'ucwml sfmin r~t1~~!lgfmmllhc'cliang.e

lifllkw~ llfllOlileaJlIiI of ilille lin fNilctBYe grid (o:rn:p!)lFIel'lt011' Mell#elill alanglUle .iLOi1 direooonas illl 1P'1l.!I. lC]i. <'!!~'WQl!~ti~ of the CQupI.w distort]Q!lSa,t th~ grid al1!~ wi~w:li!ll?or!I;'ms. ~!i!dl lel1!gth:i «eI1iWl'S alan'!]l 11m chain <In:! (o-Ndlib 2.'171.0'); Co-Oot~(;ti, 2.]30(S);;G~, .2::039ffi)' A roil' (~~~Otll·b(umdl (e and (O-NiiJi.rirn- 2.,23](;,~; C:CHlJrn_,:L207(5);i3lmd Co--~4,2.B0(5) Po ~I':lr ij}y" b iJOiLIi'lri (0, Wilere ~hrn il: from !lie I!:hallflFrormif\19bi~~yI.llf!Jamj~,afttil Nmro~r. from tM' '!JL!le~tbipynlfl;il

boo!ld to 1hec.hain.

o·r Ihc :reaCl il'iiJ;; n'l'!.1I00!.~k-::il~ IhC'lncm~ ccntcf by Jivcrsc \\'C'ak imcrnc1.iicms w~ll, IhC' fI:(!IJrI~"'Oli; SII'lLoC!UL."!1 Ilf!Cltvf (2m. nLe ti.lrgc~oo s)'!~lhc$i5:.of nWjoi'ia~:>; w.il.lt lilis J:bm~rc will he iilCOCSSiliL')! 10 gencrol.ize (ilis ohscrvalbn.

REPORTS
1.. G. I(. Ifl Shlmiru..

R~~E!~enlles 'in!!! Nl:ltes J" Salid .St.!J~,

[L
(~f!m.

17.8,. 251 '!l

2,. ,It IKiMma, K" $iBki;. (i )U;i~a.. s, Ki~agatm, Atl~ (hl!'llt tll',t f.d. !t2;, AM (201m,

{;lOOM"

tz

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~t~

T. I. filltlOJ', I), G. 11!D., 2186 (1

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M. ~ffiliIIilj ML y,M1i~a;(Ja M. IRlijilll.

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1.,. T. lDi5e-<l1Jf!!1 iII!, (~\OOt fm;,. J- 10. 1371. (2:00il~, Ha'ldle~,(.1, Koi!P!!ni. }. AnI'. Gr.em. Soc_ lll".~wn
(;lOO~}.

cnsm. lilt ,f:d.44.. 215,~ i(wruS}. B. j, L. 11J...m1il!!l. 1.1" B<illililuf .• A, JKga". m. II. Sc:liM!tt~~ ,SdeM~~i.l, 1000 >120021,
];4, I), .1\ Ci'te~~t er &. (<illJ/d. ()'l,e/ill', ~" li.?i3.,:!.9:t ,. ~7 U999~. ~. M3letf.ti i!ld !l1eUiOt!5ar~ .l!1la1~e' ~~ ~~r1I~1 fIVlier1a~ em,~detJ'~ Online, 16, L ~rWcdi. (j, '0i~1)~ 0, ~ ~~rp!o, A. S!rCWi~j. t"Ji.el!l. So!!~ DoII.!tIJ rfa~ '19~~'. 18JlI1 (19\l'n" 17_ .A.. L :5p!!~, I'llAif(lN ~1!kI~~!JI' ~11,I!~hl" !!Jtredin; ~t1jjf!1~oo$,lml. ~I!,C"\'tli"9i b~I"i'u'" ~ "',,'" :I' '('\Ii\;! ~iIi~m Q! ~ rffiyd'rOl'Ied pi'l~lol: ~l B:!¥ tl:1)~'1iJIi: QIt!J e o~d !fI;jj~~ri~~to ~!l>!iJ.nQl'y ,air}, ro[l~edl ~ !ur:l!li~r!!IJt~m!n~ ' ~d '001' 11lJ!l1;l1li1::,. rM<i'~ Ul~'1; h~ ~r~t!'· ~im ~ 16!lf ~ilI~a: ~~ l.i!r ')'C~ 2 i~ l~}'!Tilf!!lI,'~lirfl1~g ~ th.£: IPller~jt;iLll1f df 11l~ 5.'UMitJuMn re<lrM.1l.
R

1'1'•. X. :lMo tI It!, 5€i.:t!rt', ~t16., 1012 WK14h Ha-g~Fi. R. 1'. ~mfl'lOOltl. R. IH~iI!!l~:li!!r. :rubi~la, ], .r:brm.. A1'it1.t'-.r. .• 20U '(1'9'9il), In l.!i.~'iIM~ iIit! ERWAlemlil!J~nt!f'Ij)'!llral.S[jeiK.~ ~:lrd1, (ooocil.!EP5I1:Q kir '~lIIKRif\l] IIlIhr gtJo!fll'EI1'5RUC~:l1ir9~ ,af,j~~~ ~i)t!1el S)f!d!li!Iliri!l!l! f.Hj~'~~ Sil!Jf(e. We t~arik ~\ (M;te;r~~·s., ~I!i!:a, fillr :w;i&l<TI!l w'iLm ~cI!l~HI!'l l'-:

K. ~i'!WI., ~. ~in.. i). le:tig~(~11'1. ~n[tJtl,. So (. '~" J. Am. ,C!ltm. Sir, ln~ lOOOi! ,(~~ 6, 1', D, r. 1i!i~1MLg, ~~'Ii!It", M,. Hir!~li~~. ~. i3tf1m, HI. fjElh~g, Ci'IM:I'. CMl.mtm. 9~ ~2l~(6), 7, ,L\, '1'. tH. :0;. '1(, .I~I!I{!.. III, 11'.!:lJtI •. t, All!.. Q11ll'1!. 5~, :lI2:l!"

S,

~r~

~ptOO1err~~."!1w: ~flt>eUtlf!l!1: Gfil~Yr!e.rnC ~~~ > dim 11'~~I~n ~~mllted


!ol!1~

llftrllrrtllifll

by ti1!e EJ'SlliC MoI.ei:u!l!r N'!o~e~<i~ Cerrlre,

'~I!l',
j!,

~~~~fil~C

Carnb!kl9B

(l~~llo!JfilJlltlc

V~b~J~ 'Mth ~ere!l!a: mWIIIDEH ((OC ..


~1l' ilItd al1!~'o'itfu~Il!'

617200 {1~a:1III(CD<·61i'20i!

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dt,;irll,~i!lll!oWll',IDal["r.ilmLilLIlil'iIaI:iJ_ire;jU~_

e,

,~3?~ RO~S1. C.-L. (i1~!li, A. M. G~~. ~_ 0,. !Smil~"(A. S~,. H'_-C,.~Llr lowI'e', ~ C/ilell'l. rm. U 4o!i, 6~7l
!{200st
~lW(itl
,ej

SUJlIPQrtingl O.nti·nl! ~>illmii'l:t \WM.ICIiPM!1I'Il!Ilg,ll!rgf()g~tenl!fIJW31'5J5e:a,o!!?n~1

~tria:1s

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10, IL Go. B~~~ii; i'II.. P, SJ'i~rl!l:!' ~ it ~!1~, 122. 2763 '(.l:OO(J'}.

~t,.,~~

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J. J\;rJI,

Ci"lem. SM.

22. 'Sil{llei!fl!:ieo2006; i{(iepted 13 !)~!libe~ ~OO~


liUl~.ie1il[E. :iL13M,ol,~

We

C:!HlUlinoo

z iJ:COll c!}'sn:)!s,

!lmniti~

i5l!1IIC$ IWOl!J}$ rucks gllfi:81lCd thrull~'1 wmifllQJ~ !!!.l.:oc~l(;micaJ irelld~)c .Iind~~ylfl"; 1415
il1i i lilioi'l }'OC~U:'S 3!;!.tI j M'~H, \oVli>,Wony 1400 Mil). arild C~L"'I~'O (300 M9). Hie Ji IfRlabyrrlCphflt'lIlS ilfllnLtlt: S:iliuriall ~~425 Mill S.I.yptl:gnmiLc~, ~'.lliC~!U>J1CWI.lY \Vfirry and CooalJ!;'(~ rudks are I cnlpJ ;;!ICOO IIli~O Oiroovician flilCIU$OOr.~'1e:liI,ary !.lIililli 011 II doeptUI of i1rom~d W to I 2 1>.111. Rock::> of ..:ach JjlHuni'tic :;lli~1,; htlw h~llmJilI;'k ~:·lyp..:: ICtUlib in IIIt:ur1ilumdmlCl.: or htmlh.1...::mlt.:[i~lrl iul IIII\ii'l'i~ h i~l(j]' OOIlt:elilnuljmi:; ,of C:J. NII,~ncl Sf Ihlllli do S·-typ;.::gnm,i,l;.,";s s~n1ihlli ~iiiC':J COI1!.(!1l1 (7)1, Samp.I\;o,<;. ,II dtller I)nd .of ~he C0fl1"

or

fixlOIn

~hroo

ill, I, S, Kefll~;~'~" 'CJ,H~w~eS~lJQI1III~,lij, L ]!. ,M,. Helrgll,. '!i:,. M. Gr<l(Y, - M. J,. Whlt~lh@ lise

:~te,r,~a ,A, IP.<lWJSiOWl,,:1 J.

I)+Wooohlea(J\~

,Zh'COIlS W()1t' .. 11;;0~lml·.[iilW fi:mu .. l'IlaJk rnt;;lk" !J:'t~ialcd \:IIln~1each S~lil~.fllCS~ b~illg !J gahhm I as lvid.ely bet1e'l".ed. li'ly~ ima~lm:atjl:smlhlls i!dlives tlle cOIiIP~ed grcn'll.i1I i31nilil!li~le~l1irfalil(l1l'l of Oindllbyn~" sYl1p!\ltOll.i~ (~O~~ri~l; d~k~ ,(Wh~ ot;OI:J'IIIM'tiltall crust, Wn.!ry~.arid dil)lii.c'c~cl<uv~ (Cobarg.o~ The q]!~lic roek\ ,aII'C' CftC\ a~wilh th~ ~$.I)Octi\'C' ilk ."lI!i tcs f,l1,U! al~d ,,,"11>. the- ,~ll'IC di~!il)di'lC' chemk-al fe;nuR;!:.1 nr:~ lhe-~n (7j. '~f'h'S venlCCI!'of cm~li~!Co!JI~ C!l.LS~ L~ 1-'1::rlloC ~~(l1<ni(l~~used to dl'scribcU5, !rfiJ, ~!illl)~yhU~: C(lnS~f1~J!.i'!i!y. uniqw~' <Ln~()l~ghe ~J1~\~.~ ~~n'C$;trkl~ G~:anitcs ('!n~!.YiI'IiIl aJJl~i~iii~ g~nh.. ~~yIl~W1I~rIt, t __ planCi1.salld 'lueSIS I(l a OL\Ullc!mV( !1t(l(lc A comllbl)' or the 1--..0;;I~ pcc;fI:nC'Cp~i s;ull'l1 Ihe111e :,dro:ms of ,,1'1 :l:J.1l!pks \vcreiirs:t dated by i iOilS (Df grnnit~pcflec~ the inn mi.cmprlQl1C U-Pb iOO1OPC,;!!1~lysis u,lbkSn. 001'rliln-Clal)! di neremj:l~i!Jf! ! The vast gJ!<H~itic isotope conlPo.... ~ bmhol,ilh ....end ,!heir CI'I\ijlled <!i!'l~~ogs ~l~adnmi~ cU'llislal mi,dcilCC prt"hislOIY oi' '~lcif IwcoJi!hil, Thisid:~nificd a nli!1o;tnh~l:i~cd (prc-il'li1gmmic) U), I;o;n~~rnh~~!lg d~Q;,1gQ~ld ;Jrc'hiliIXl.\il!C' of th~ jl<iI~;:)on!ltl!1cl)wl ~.md!l1~~$c~i1IJ~,m obvioos '('Imp9l1C~~ ;:1IO!l],'C :l<"!nllk'$ ~I ;i}, I~I tho,: ~i:,;hI cu:;....i,or~ Iio!:I~oll'o' I:otlll~.~ oli tIltitu !Jbl;lin~d fI~nljl .llilmal,i,:i~~lahon of illil~h di ITer!.:rnIill liOJl!, Vel how t1ecp CRl,iSq.l1I~WC~~ Ill'; HLt 'tillo1in~ Ofill:EiCIlII. cm~I.· p fOllrl~ilJ~ !.~pil~txic.... In tJtm'LrJ:s'l. olh(~r SlUUi~";i; 'drl": 1lI1~~1~1f;tI1.ieoni.on Qj' bildl zilrttH~, 0 i,ooltlpc ~nj niE,c.~ rd:Ji[t.! In erUli>t~11 1:N,lWlli piluet;.';SiS~ ~ IIm.i11.5 liJlla~illg. b!:lCmJiS": ~h~ mqiurity or 1lI1(.';St;; :111l!I.'itrtllCtilt;; isol.ope· v.:l~'imtun5 1.0 i.rILl::llllCliOIlI cmllprn.;.il.iom;, ,(l NOlJ~. t;;XPI'I..'i:'i:.;crl.~~ IJ I.~) (/11 .uur;k,o;:.~~",(:. i5(JIlUPC ::.;irgumurt~ 'lhm II1Itdudc ;1 b~Lween okl.c.r ctIJm:.mcl m.:JlllloocrriIV~'rl.fII'I£I;ll~" "'tll\~ {kCOIlnill(:d J:rom ~BI!:s::II.n:: zin:'01li :gmwlh (2. 5. (i 8). impl,y.i.ng Illal. t"!IlliS~] 170wth i:s, zone llI:;i:ng II Camct~ ~MS ~170 iOJ11n]icrnpl1it.1lJc di,lt1Cll1'IlI1ml~C Im~"'5lry ~l-6), Il: rmttuli.gln rlu.iIU H ;;I.lJIr:ltciii illl the Laehiall Fukl Hell i(,i.,~lG>tcmi\.H."· iI1U6!1.~iI:: Hi .gmL1i~ic 'b'\l'l1pi.u(:Gl1l.t:n~. M~",il1,g n.1t:tt:1l~ U.'n (I<lb1c s;n. lJ~]iik(; iJllI~ Ii;, ruck :->rmlpj{:";:>, ZiK:O]] diS, ure COl~In.}V~Thiill {9}.. ~CiIUf>'!..i .'l~(.TI!Lil1illg lhe ~;o;'e"'ITcTlil.dy R:lCllli\'c oftht: Iml:gllil.aLic 0 ii:'(l~~lpC Eruii,l, j" ~hat gnmil L(: nmgrm..-; dCI.i",11: fruln ~)K'" tilll:ill!f:, ~ml llIl;'lgni'Lltrdc ofrl1nDU I~ inJllll. i$ illIiI· mlio (l9), :100 z:in1Xlln~ i~l cqlll ilibrullI!l.1 willi prris. !CN .. i t~llg SDlIK:IC li:hat .arc .full.d~bl1lcrll~lny 'l.:.ithCII.· ..... di fric:ulh~';i;. willi n;:1.rie ...i]~g llie Irct'lm:d Qf ;s;llIpmCnL'i11ll (fhml~-d, or.i~iua~~j' 31.~ll1h's ~Ulr· r~b,y l.iI1G.Imllllh:xkru\l~'(1 Illn::] G.~ ha\'~ H nm'Ti1w mli!j.c maW'lltllie tvuhn:iml [ronl '[)Ill] Ii !j!ITrmi'Lc comlID~i. r1l(Ii.~l· il' iWli:Jel1l~l!td ligl1:(:;tlI.H:1[ock:. IJml :\olidilrl:t1~ O o",i!iIEiilO [5.1 ± .03 :lliCrllII ill ~WH'}I (I.J~. This mu!;c lioll&. fhis mcmrl il;. IllOw~'\'~1r. pre~f'\'cd w.uLlflila i" iilli>~'!IliSiuii~It'~O nmgJlltIl i(: d i rli~reJ11)~lilml. 00· ::lL ullnh} il] dlj31TI.c~X:Ir. :is rulk'lCLOO illl Iht S· :mcl • C1llI!SC Ihe allmd<lIll ri,,(J in lmll .rndk Q"i!iO it! [1m chOJllI~ca~:md jSO[OiJ;t; i'm:l1i1ugr:Jtphy {,lr ~r!;]il], by un i~lCrob (W f 1~ The rotf!.I<SI itCCC&'iOJ')i !l~iFlCr.u1 cf.llnf1!e~ls.; .. cd R~~ ,:m !!1.Crt.."aw h~.r-i,rtiO!l.lliqllid lBii:lli.Qll:~ (if(MJjih. !:anh Sli:j~iiIC:e~ "{!!}~rtfl1,1lIl'i, Ui1l¥enll". it:>: 01 BrlilloL Ilriru!l. I~s:eUI,l, 1iJK. ~dMlo! IEi!f~1l ,md zht{lll has ijhc ,adiv,~m<lgc~~I1!1~ I mric~tc gro'\v!11 e,1!!:O rtaCliOll<i!li{)nlron~ ~o,5'%olor n'a~k nlH~hs ~n~ng C.~~l he datoo bylJ-Pb iSOI(lffleS, <lIm! i~ t(fi "~$%<o for !>~Hdc(h:ri\1.ll!"h'e.~ (11). V<1JI~I':s t1tl" ~rnirmm~t~ S(ien~e:~.~mffi ((KIk !JnivelSitj" lfa.m~!i\" 48.11. ~tr.iil1~. J.S<i:bod,'DI E;ylfij ~~H~ l!Ini'!le~i~lI' of tr.u:ks d'l<ul,gung HICI! d~C!~lisilY i~~ m;:lf!;(l 0. f,1:SO ia :;rj~l;ona:l'li1\,c5,69~ tlms li~lg~~rim",~l its Me\lWlmC', Vi(Wria, 3'0>10. li&tro.1ilii. 'enlJelo(r~to!lRlka~ A ~ il~aijflj'iCrnalCis V 1" J -I). \~~c reporl <til illtcJ;raa..:d itl '~O"C'4u'oched ~lij1:'lmci'\i!~ta~ OOmil(iilCm ill the I~ 'Studies.,~lil!boroo~llI. VictDnO\i if OS l'!w~ralia, 8, ~i!U U-Pb,11 r. <!Hd 0. i!l(l'lOf'IC~ilildy of .droon!l, I'll<l,gma which 1!!CI!~roJil clI'~la,m:;o;ed, ~S,mdi!ltl Ml2!ieumDI Hil1UJa~ Ifi~tol)'" 8~ 500a~. SHiH OS thi s tiIC'hlg dlh~r ,<:cd imcDltrr:ry rock t ~0 ~(l30'M..i fMmngr.mhcs, in!ll~s 'C<l!."Che Lachhm T-!YI]oC;~. t tfl 'SiOOd:lI!!ntm. SweDen.

'Glranitk p.\l!I:oIl~smis t ha priilJtip.a~, t!9lmt ·o;f mJis~<JiI, diiiferl'J'IIlialion. bll!llimlkifl9 Illralllil:e e mpiacemenl '~(1!trust r~:rmatiotrl ~eql!!itres~'nDwl@dge of the liIlagmatic e~ClIl!t~~I1I, 'which iis: F1@~ori(msly dHfic:llIU til reoonsuuc! hom b!!!~krock cempoS'itions. We un~ocked ~'hie pluoonk ,uthille thr,ollgih haftrliulIli1 Hi!) .a nd QX~.g en (O)is<ulo,pe a n<lly;sii5(1~ z®n,edzir~(ltrI (II'YSila~S Irem the classic horn b~e'trlde·b,e'I!r~11 g O·type) g ranites ~,~ eastern A.uf;tratl ii, Ti'liis gr;aIillte type ~o rms by tlli,e Jl!lMIo:rking o~ setJli!lli,eR~ry matewlah by 1mBII'Itle·U~ermag.m<ls tFl~ll'ea,d 'Ilf l:iy ~MIIe\lt~ilg all'l(ielflt m,eIt81I'11iorpdl()sed igIIilN~U:S ro(k:~

or

~~~Ilimml

I'a!lgc' or each suite were ~1I~CI~-

r~o.n'

... :c w1!I;1!~ lllif~~~l;Irud~,rnoe~d ~ h 'OO~,~mp@j~l!.edu,i!1.!

1J,e, ~dd!lil5'lli:d.E'mili~

d1C'Cipl'ler Dhe C,tlS!J~ CVOllill.I(W<lU)' rroc~'S$es

tC'ltding smck:

magmm i~!l'l

al-

Qr

~I~en.xi \(I,IC<1J11(:rock (to 20%0) (1fJ). The 1"!\nDtl7i'~;if rntios w~rc mC;lsurca try laser

980

16 FEBRUARY 2007

VOL 315

SClIIENIOE

\i'V'IoMIV'sdencemag.orgl

REFORTS
3bbniOll nftldl[ioollcClOw imtli:C~U~~~}' COL~I]~c'\l111f~ nil it mass Sj1iCClroJllCilI)' (IS) (ut~i.c 53 },Iillgi~iug.

""h!!!r,~vcr jXi!ssJblc. lil'ie pits lili ienlprobe im~l)'s:is.

gCIlC:I:aI.cdtlly

ion

W~g. l ]

t\Ui,U"KlrlOiII jCfli~m'\,; f Ih~II:H iliffiOp~ data o is [he ~[il!Cctrum ah+lr!',,'a~ulC5 e~hil[!i'l.t\cl (lilp
Ifj,

mnllysis Ms, cs!ablis~tcd 1~11"hrU rolio.-';;~YI')ic::i.lly ,ran rl:'cwneore 10 rim iJ'lIWIC ~"Lu(Jkd iiroous (mlJlc S I). ond Ilted.lf values ofillC il:iiroOiIl.~ 31~ JC'Cn .. :;,)'s~~'lfn:llicmili}1 w~lh Thflll ,li1~,,2:). 'lii\t{;. 'IJ.'>e inL'Ci:lP~t lili» as CV [dc:m;.. for apr{Jg;lX:s$jy.;: ~
mOIIClion i!1l dlC 11°I·mTl1l,1 f I"allo dmiill.!; lhc magrmLlic (';vnl~li{)n of eaeh :."u:Ile. :ts wOlu~d be .induccrl by LlrlC!'nddiljon of:m Uln"tdiogt~ll.i:C (contint\lll~1 Cn.!~1 like ~oompo'll'l:ml. ,[,.l'Ial'I,,'.. r~.

'by ZUrCCill~~f l.im s:JIllfI!Jroek o

HI!?

lIiI~it .. )

011. Su~h \'@[I:nil:nm wili1.i.rI ,:11, t'iilagl~ t1[ul~P!JiCl can olll,y be IlJ."'OOllCilWl!ly the Dlx'lOltif.ln of OJ}(]<J]o :!<i;ystcm l!I'OC~~ Ihill, an:: C"")llb~c ~Ililling the 1'16 HH I 7l~,f '<llio of llre mcftt .from, \\'llidl~lle zl ,. CQn.~prot::iri~,ued. 10. kl.\;n!,i~,yrl'lC~ plDc~~t~ il is 1It'~.f.,~1ry to ~lJd~cc ~le pobJ'ily ol'Bn:'!o!~,l~ il;hangc ,d'l~dn!~ zirorm !lr0wlh. Thi!'l emil he Oleil;O.l~l[\lli~n1e;d y~'Ul1iiuali.nl~ h UIIIl';l>CI)'!'iJal i~ ~I}pe- 2Jmin~ l~'llill; and ~I [)afu.'illlg~h(l ijlCl!{~ll~ V<lriaIUf.I,l1,<; wilh ~wacC' dC'~I'lc,nt mIlo:" 't."-:LICh .. ~.; l~hI1J) Ihal na:c pm::des 1"01' the dc.~ of r!'if.

or

Iow~!Si$O cud nllCi1ilbcr HIiII. j::; ilil cqui,libri~lm wi~h ImmLk" O • For Jjncl:JIh,yn~and Why '~"iCiIirY, Ifhi~ Cocru,:-;f.i!"~UI(~'" tn ziiroo,J;ls, oif ~hc a.~D:llcrI JlIl'lfi~ illtl"ll1lh,'cs. Al hugher ,1)I~:O\!rL~u.l15, tim ar~,Iyt'; ,~jl:)nv~rg!J' 1O\J,,';I!l'd rha .fl(lJdJ dcfil~l\d by
I!Img~llm ic

va~ ucs lIJ~I1@.;IC a sumt8m io ~ S'L\llt'J~r!J."I.l11~1.)m" [he Luch I~I], ,l~~}'Ul'!! hcs, 111;;:isogrim 0Cli1]~ array acfinoo by ,d~l~ OfCilClll $ui~chill.'"(I dbL~ncl i",~' IIr"Jj~c'l.Ol)' lMi:Cil0l'lOd iYy a ,,~iYcrr'I.'ilt1 I
PO:ifICiJ1~ In

Iy W!l(\d ::l:Il.'COllS 111ll1Llliifc';;1cpi:;)[iidic il1pul. fmlll. mal.l'l:i.ri ""ilh hiclu::r 1':~.:I:fv<!lu(l,'i .. CQtJ.pHng th~116HI!,~n.Hf n ulo:;; of lllC zi!.-.
0< UOOIOJX: oompot<;JI.irll1t"1 "e\,'~I!'i, ~het\!l!lll\lJ' onBa~ cllll:;;l;:d ~Illf'){ll!!en~, ZiI'[OIl~ nof iudh'idu~l t';"'Lf:!pi~ i()l(~lillil" n ;B'~' ':;l.Il,!:W of2 to 4%<.. <BI'lIi. n~lh(lLI~h IUl"lllly :!'>C~oh'ablc. :inl:l'ag.mi~l hrnCl'ggclIC'ul), can 'C'J>~o;l I~'\n and !twol \"C-s a ril~C jill Q~~O limn cmc I,Q rim (laMe S2).. r,:,m ,lI'Id ;;' ~O are cOfi7Clm~ed for z,lucOJi$, OfCl1Ch ~ sui~e ~rtd dj..:lil'tc ~urY...J ~mbr.>!'hild cxreudto IlIi uch higher ~l1t"1, \viIJ~I;llCi[

.:tl.r(l'm~ of S.rype

,1lr.m~lcs<lnd

the 'In·

rCHl},d

~~I~f·{)imtOfl\;'

COlUpOl'i~I.ioJ1, o f d~C' ~ItCl!a·

or

Ibrcm ~:11iou. hl CO!S!;''' witcte iSDlupc zoning ~s prom.1tlnlcoe(l, l'it'll r/I"II. :~f g,,;:rIC"ra~ Iy decreases
'towru-'iB zircon rims, II~nU.iligh some gminli show tit!: tl'ppmi~c P'~lto::'l'il. Tille,,; c] !:lltcllL In'lict'6·

ZiIrCif:\ a:rm,Y,~rurc I)m!!~\n:ss.h'c in~e~<I.C"~ ion helM 'CC'J1 l\~"'O CI~d·nl.e!llbcl"' mll~l1!t.l!1iIC !II~~ durillg :.dreo!ll 'c!fyM~~li:l<l,ti[m. 'hose- bd fig parcn1!:l~ lmvI) I ~O m;QMm'I~, ,~nd n·l(l1.,1~",'dl 11J]C'll.:'LI) ... d('.rh'~~ IlIlmcr:i,;!tl!'l~ :S~ch Ijll[c'r;lCiim~ cOl,IIM im'O[1 \Ie C'i~hcl" mi""il~g am:1hylJ:riaizmion wLth crusial flluilaffill dflilrerm'c~~ac:lI::

~dhr:lC'l1l!1i1.V eounuv 1'OC!kS,. , . ·n~OO~'ITfi[]lH~ ~U f- ;n I~)

ing

l~~

o'~ovalues

-4, ' ,-61

-a-,
~10
I

;
.L

,
~ -3

T nmulhtl:s~ of iii!1Xm,~,j:l!1)I.'CiPfiu1l00 {m;$imilmi,(1u) by Itlw.3]~O m~~mm;,. h.: OJ,;' rroli~ mmilk ii'i~lt:o: WiJ;;, ].j. UI~':l'1!~h; ...moo {il:\lO ~i'li of Ihe iOW..~I:~ ~rw~niiiSi$ nOt, ILYI'ii{lLicly c(!'.nt.1nlin~~ hy Illl~zj~~mi iSOimpi.: ~lInly~" The dal)[l, are pem'lis:;uvcuof.:in eu richcdm'lI1.tic hen.'~dh;JCi'~ ~t""; 011f[nl :iIlfhlC lILusDal ~lrogi(;l],i~or. Of (If vjri ;ID~C c(lnnbiumionll> erniC: ilWO (8), A rro:h:mlillamly .'~I:gnIUIe lIlifUlllic ,~~r.ivmioa] fICC!£Hli1i ~,vitll the :nmll1lC O~!I!rllRi~ !;;h~rw';I .cr of 1:11,.. rcm:d Im.... lila ~'\Ill1llOncu'~. in ·~ 1!!!l\VW~ TIiCS..: nm:l:k 1l1;l,!;atm:;: wen; $uh~~ILI":;IlI.ly modww~ i f~edbyCJ.·~Il>I,lI] (;cmlmlll iltt!l~it!'l.1, as i~ ev id';;'nil :::JW"o!jI~ i~1 Ihc s«ncnd fl;f:i:iJ)(:f.t1l f'f~r ~Jld on~O va[;u,!.1"£i,~ ~!finiIliiiii I1!lI1,~ J irl(I"t~}'.oe;ihb~)Jl~~d hy~VQ.rry ~lol!1.'1I[ g W
Q_~~

mehs

'Or lhe

d ig~~~,io:nof

~l!Ipmtn@wl,1

rock

~u,gna urc, •. A!!hil.1llj\lh

(fig. 3}, but .l.j~oml crys~al~h.cd ~~Imdi()ml}' i()1I:dy 1,{lIrcmi III \'U:;;~[!l.C~Q he m'i~i'll"ll iso~opc

r•

!01'!i,!l!; .<:;!J~ptX'oo

p;li'It~yS:Url(l"CJrII'Si~11 ~,{lurce

($). <L m!:r.:,~d DJK~ fnr l~e L~hf.an !,type

-12

+- '

--+----I~t__+_+_-t--+--+--t---+__+__;____1t__+_+__t__t_==_a

.,

-4

-S

gr.mi.'~~ ~II(I\II' inoQl1t!\1V~!1ih~C'. Ma~o,)i!,1procruilmi i~13l"L~!. abo l:ec~ll~y ~n inriC)"'C~~fil!" hu." o~hcr !!'iOccnri(ilcd drclHl1-Pm.c~r:1(: 1-t.YI1C pMol):'; (2'21. The r'CilM~il~ing~a.~k ~, to q,lI~alliif}' !hc

ENd (bll!Jlk, rock),

-7'

!l!1e~(lSot~i.!1~C'!1!aTy i(;m~lJ1(I!Je-HI oj" 'C'<I!dl lachi<l~~ 1iJIUil,.; 1.1111;] W, dl:l<..:nlilill(; 'ch!: ,~omj iil~~)mlii !.meter

Fig •. 1.. The 11M ;,~iOtopE!.oolil1pCilsi~loi!lof meUl"'p~eciIP~taltedl zircons hem salilliltes of ea,ch .5!ilile as a 'hlndiolll (If wnote-ro c:k Ndi:rot,{l~~ (Qm])OiS~tionat the tim (l' OI[ c!!)'S"tiJ'll"ization!WW:, Why W{H!)'~. Err®r tlaws ITe',pr~ent :2 SEM.

Mlidl l.ht. WIIS iocurpomi,_'(]J, ll..:;:SUIn'lillg th~.t a~~.inli;i,btio,ll.I~3~'llim~a~~~ryl>U1I ~izu~ttln (iq'lltlll.u:l1~ {131 ~LI.'eII I"I:lIi9mnlkd~ appn)N:~lTii;~[ioll,t:!f Ilul;lmnic ~l' ohJ.tit:!ll. 11110:;1, zia."C0I15~1:I1.hL:Cohru:go S,uil,c ...

'e' diDlil:o
~1,1i.l·,e,~!~

u!: .... ------"--___.:::--------.


1.(11'

Why WOil1)'$uflIJ

I~TQ100 CIOllIOLba, 'TlKG17

HI 0"91 0'.8 0':7' M


0:;5

II'!(l!\O;hl.QKI'(,ol!

II

CI~.~KK2

ogmD11o

0.2 €I.0C.F~~~~~~~~~~~~~
Q1

:l:;.;he.

ll'A
[J @,,~,

-y.
-0


~I

O~

~ ~~n O~Ip-----~--~~--~----_+----~
~11l1 .,(]i.
..,qi

1li'.:>1 (!I'.~I
OJ(!

o0
-'I'

th.,e -3

0'

·06·

sHf
U~e
s:e!1l5~

di~
ziHO!1!S.

~·Hif

Fig. :2. IHr ~~el{lp~((lJmp®'i.ili®fiI~ OI~ :;:lrmn!>frOIili ea(h~uilt.e p~Otl:e1/il .rgalnlst tile 'I1I1!fU lFat~o!ifI~asl.lred fo r tile sa m!l!' par't of tit!'! (Iry!'.tail.. .#lrr®wed lillli!l!s shoi,;l

of

((IN!~t~'ri!TII

z@nO:!~lol'lf,o~ i!ildivid'u~l

REPORTS
precipil~!tcd froill. m,c<lts C011rlfLillill,l:l up 10 25%
'.

o C@bal1go

OsbaJ)O!l!iO.Ii1,Q

llil<!n!!l:!' rt;!a${lC!

:>l~ac-rU5lnimmcrUall,.

lire cQrrL'Spondillb:

fi,gUt1CS

• 1W11rV

Vl«iff!j'

WI"i!ll' W(lrIJ!l'Sui1!!!,

,~

,100,

ror Jindabyne fLlld 6W/~ 10ii Why \'VflliII'y' r fI~,j <lI ilLJ :;,~miXlt~igi!!l.O,lliillc L\C;:u) • The g'-~PU-~h)' or til,.; {m.~IO:o.urn]'s plaecs 3dclinimm,~ rnnt'llrnil1!>.l 011 the nlili:ing or a~s,imiIal.irm pmoDe:1l'1, IK'C.:JIIltm Iht~, UH:'!t'dl'lIn'C of~ha-NtI C :m:m,yt1 i~ OOl.lLro~ioo by rim re~:Ilh'C H:rcol1i:::t>nn~lion~ ofthe end I1lClIllJCI'..... Par C.oib.argo arul J umhll)}' 100, the IU COll,tm~~. or the rmv.i)HlO IlImgl.na must .li'Ia'\I'CC-..'I:.ocOOed that orlhe CI'lIi'iIQ_~ CflmJ~OJ1.c~11., as woeld be Ihc: ease ir ~hc hmer \\ere a pmrtb~ ~nc~1 with 1I'l."'<"i:"~l!Ial?i~COlli_ Co.n· ~'Crndy. th~ {lL1'Jl'l;',[l\'~lI11 Htm~ dcfi nod by !he \Vh:~i Wa[~y~l;Ii~l.l."jl'OOl!I~, l\Cq~lin_"S ~hc~tlprn~ lh<l~ clU!'lm~ .ingrOOiG'I'Il h,rd the hi~h('r. Utf C·QUlr;:clllm~ ~i[Ml, which L<;, cOi!l!i.~riJJlI \\f~lh hulk a"'<;,nlllilminl~_ PmliaHy disagg.Tt"g.11!.'{~mcn:J:;;~dilllC'Im.I!'}' cndav\-"'S in IhC';5.c iJ,hUml~pJ'ovidc evidence Irtlf ~h.isf'iro.;;l0",4,
I;I.:~;:;(15),

17C:i1.l!i: on..luuKl U

10

:eHr (2Ji~con)
(,(lrreSlll'OmJi ng '~() m.a'9ma evc!lllUlofl lly W.l51:al a!S~mHa'lll)'IiI·(rysti3iIli:z.a,ti""lil U!JC) (2 In,, IDait,a,froiill sampl.e5 ®f ea ch :s'LJii~e'l'llrel'a~l!Jlluld~'hre same svmbol, (eH~H bars dl1'pin t.hearVll'rage2 SEM, lJl1I~eliil:aimJt l'Ie i:l~eO 'ljalu,~ of L,iI:M(:j iii met,a~~o imentary r'(J;(iI: lS~~~1l1 ~rillm (33), aln!ll~e slil~l!tlE!d~Ie!~ .mepl(t$ [ne ~~.oltJpe <emp aosillij,(ll!!s,cd zirrc@ns lrem the lillc:Manr S-t';'iiJe gr,a !!lites:.. lid<s on th,e I:l!r'>le!s re)jlresel!!t:l:CWo AIFC , 'in! ~~IflIe;ruB. and the- ra liloof H~ !l:ClIf1!:eI1I~i.atlo:nl~ the pSlrentall, m'agmal (pniJ}alliid ml~lat (c) ,e'flo ]1'1 memlJeirS (Hfp:nltiI~~~ 15 'indic:a'ledfor eaeh, V.sMOW, \fiefl,~a stam:Ji<wd mealll oeean wa;l~lr; US mHao$ed. ro~~, 1l.aldll~lrI lfold Bdl met .. ~~dinliif!liI~ary rrt!d::.
Oir~;l5o U!~:s

rdro~nchc tilooc-I$ ror


~j);'~ili:l~
1~'il~1 ilIeW dfliUl,

fig. 3. Plot

\I'@1S!I!S£'"1

IDII'zimll1s of this studv., !I'ho~"i"glthe lPu~ti1j'e

theW!c~lbll1 l~tYiJc ~o!ie:ih: I,WG key a5~i>Cei}:of Ih!.: fir.>~. IIillC [111:0 i!1(ltL1pI:: :mTIl~t,S ,~.~iI!~'iI;rl
II~II .:t::ilPCon rule ingcs~iol1l

ro mai:ule.llikc ...alues, indi~i~ug clJ"s'~1~Hzmli~1l1. 'CDlTimlCIlC(1:IIJcaorr; l,\;Il'lpo;;:umure~ Oil' 1li1i.l:lliaretiKII


'!'\'\;R::

of supn:mni:>'l1111 1~:liHcr.i!ll. 1.~.OIN'CV'~. the

illw..ol:!io

iUliti11:~

:i.l~fLgm<l::>

in~vil'tibly

~lrugh!.:r th.m~h,.; li:I.t.·on ~,[ciIbiilily


<l'HDO"C (2'4j, .mcl ~lllI:;.i,1J1 im.. ·or-

1i.~lrl. i.ypiCliHy

F~!!I. 4. PI ~il:~~If!1~ti,

model.r~~ th~rorm'ati.oUl il)f Hyp~ grillnrit@)i!fl ,elli~term AIIJ~tr.aljla,. !basedl


Olll

cplwdlc

~lI!1t1t1

ptDJr.'lli(lU ~ n~(Jired.Tl.:C't:ivc for hQ~ liquid ~t~f,,!!:Il.'ilq::; r(251_ \1Vc- "Il~rormt:! ilnr~rlh~l. C:1lII~"! a~ilHil;l~irn~ l1ud lJi~c'(m p.rCcip.it;rllj,(m 'occu~re.d ;u~ diflb~n 11;: mplT.ltures,<lndluhu!\ crustal lc \fels, fol" pari ,)fthC' 1II1,~!)!;I~'r'H~,C i';\lnory. Scoond. lih(;ldi\'(:flli~)' h of,!" II r <Iud 1)1 ~O va I~~:;;; i~l.(~km~s ]h<lllZi~CD.'lll~ of ~~h ~Il h~ g(ulllrl~~ .ro(~k dilT~f'G'l'lt ~1.~ll ..Hy Cfl!1.lp~ilk~"!l. }'c-lthey ..re !laW j~L~l'''Il!Psed with"

in

11'lC

s:~mc r;(l!ck V(llll!'l'IC'. Ihi.Sl

the j)!lil~YsisQf

'5e,mtllage (29). The~ hybrid ma'9lfm5 are ~x· 'li'o!!cled tQ "'~ctf1d ijPld

i~olop~ ~n :zIrmnl, 5ihcic: mel~si'l re ~e\FLe~at(\d at rtI!i!pUil ~nr,oogh in,te-.raC" lion blelwee n re:sicllu<l~ liqu'tilIsrf'om basaltc¥'" '1:i!llllliza'~,orn,andl iiIU'!h:s d~rfuoo h,om 11h@(),wr~ lyirlg SliP r<:!WI.!S.'lal <lIS-

H~~o'

cess In dlC pl\~hlilic c!t\'im!!l~letll

dCi!'l'Ii!1Ild~"II)r.odbat 1;)111) wlilc

1Ml.~'" Cl)'SI'1I1fil1!l!
re.~l

reNIN
rr1ySro

~(mdin~he mallo\'JI crust


.ziirrom 1.vnme

'~lJ ~talli!ze

C!),),;lals: wilh di~mmtii!'f1'e!.mg~~1.C4.i.c his~Ofks" M'1!~}' vnh:::<Inic meks illj;() CfIompri:!>C <tgg..~gilh::s oi"cu)o':1!1als, mfUled at di""crcm timcsWrom evelvilllg nwi't '"tllUiiX~:;.ilJ~)n:;(lO, 12, 2(;" The h)llowing:;C";::J:lHr;;ilo~s 'Il~PIk'i'-"t.i Wig. 4). aecmni1ltn(}da.i~i!Jl th~ '~i,~w I;h1lL 1l~I!II,OlfISmX: 3~:;.cl~tb~'-"tl inclJTQ]tllC'Jtta~~l' (1 i. 18), We li!Ii .. ~~m~~3 d.yumDlill: drlml!·Ir.;\''C I proeess ~11",olvi,nl!! till.!' ~ul· (;Otl1p.ld~ iSoliUiiflC' .... itm of bl!l'01~'lUC .L1i~lgUIl!::tS in II clc~p CI1I151f1,1~JI. rune ,! <::)5 ~o '10 k.mb, coni>~111ilncd h by al1lc .dJti£uICC i\)!" a g,Ultl'iC[ :s.ignililirG Ii III 11~c: :L<lI.chllll!l I'~ypt,:; J,. w:i~,h bcau:1K';S, di ~li;I'(''inliaI1,.1:I of
ruch 'beilllg c:xnIT:lN~~ W 11l!>CgIU:l, oalesce, nud c cl3"!lla~hz~im.l~mllOw ulil~l1~a[t~l:n.loirn t2!!, 3'01. The iso~.opc Ey...lI:'1l11Ulie;.;; II.mch mdt aliquo.t rulkt"~ I,~I!.:"'dlt'lm~ Inl~all CUlllIlxM>UI,itm and Ihe p

1!\()1:opiJ: 5~iF!awre reoords lhe Ipr@gI~S rl)f S\IIpr",.(iru<>r:a ~ illcorpor,atilm at depth. W.'i !!lg1ing an cI a mal~.ami!iitiOfi elf iiTle~t 'm!Iill!~.!t@ zirco:nr)Wi1tl'l rdirft'reili.t is:ooope dna rae-

or

b~t~h~'~ d~!i!'t9 lp:rQi'Qpili to'1Ii a5:5Ul m.~~ac·

~e'lii~ti\;s. (i:asa~H,; emplace rmm't is locaLized ,i!tt the ml@rf<l:c:e bel~~1Il


'Iti,e maJ1L {eoeaflld slllJ~tta~ tJlJ~ne ea~l~rfl LaCrnla ,~[)Ild l!Jett .a11[1 111'~'e~{'rrl)'iiFIgtllmbidiillf' pl'Loe,;tl1e n

intNra.'Ce is likely to rer;lres~nl '~I5Ub5tanti(!l, rheo1ogiul (O!1itr<'lst

. VOL 315 SClIIENIOE

dcgn'lC O'i\"\llmtCl1ils'~1 hlP~I~, OO~1,n'lCI~c-irl~gID1, .his dcrr~~'! hclore 7:i'rl;;:(i!1S,H!JU<litin~!~md ('m~~ p_g W inui higher ClllS];,_l~levels where zircon bcghlil lO 'l'}'stJllir?-e ,(rig. 4). lh>:'f!ll!;ll :>i.iml~<LUQI'!S luediet incN<!sc';l crustal ;url<llte.xi::: ~"'lth hetzone nl1;fjj~ii i'f~t~(1]'~(J9), p'i)(m10'tin.g ll.i.ghcr rates flf 1'l1oi:wsooi,lllCI1HI_')i rock <lssimi,louion h~i~ltc cl1')':IJlIiililllg bJ$~~IS with time (lS)llInci gl'C'<l1~cl" blem1iug bdwce~l the' b.!Sal,l-dcr1\iCld liq,l!ids ~n d cr~sm I mr.::h$ draini ~~ ~hc h(Ji~ wne. g

98.2

16 FEBRUARY 2007

\I'V'IM;V.sdencemag.orgl

REFORTS
S!i.II::~ssi>,cilnlusioIlS
.nifJiWClJU p.lmofl
COIlL~

wouk!

W~I~I~llghcr01 ~ ","'OOmodby I&IC' zircon isotepe with disp.lrul.;

of ~ciYbrid: nlidL into the HlCrdore rlroei,pamlC .l!lr· and l\Ywca' £U{"nhJ.C.s. &~
~~1';;W~. Z
1]1"1,;

l~re~U1oe~and

N~~eSi

ire0l1s

i~YLopt;,' $ig;Jm~IJI-';::;

jw;'tfl(lo...ed

by n11l1g:lin!!l mld '(;f~ta~ ~};ch:.IDl~c~cl\\olcrDl nl1('~1 bmcI1ih1 during 1)~Ulo.n :l"-"\.C'1I1b~l'[!Ind immI'Ctwr· VO:llf!:'ry!>1~~I.Hq'llid ~r.Jr!:in1.!l (17, .lJl ~\l1Ul\~ w,~lh U1~

'~hercr:h~!Fgt1 mdh would al:so dl'i\ot~ thE!'It;si~m!1


IlI1mgma ,~nldil~ eryS'NilIl i!1.in;g;dlt~nll C1l:[~ lD\v[Jm .h,ighcr 1Si~O and lowl;'1: F.IU \i;lILles. e:-ql~;;linllung the inlli,1z:~rco!1l il;,ol'{)f~ zoning ..l~olOpi,c ~C\~r:;;;)~'~ in~m~ zircons :l!ld h<i~!~ in1ljI,'Clioll in ~h\)'Why

L I, It Ga~~i. '5, It T:ayl'llr; GI!'iIj!!~_1'5..~ ttlt liD" .~fl6·li (U8~~. <1. III J. D~\loJ~, 5~irn.re" 2.09; 6811 (uoo~ :t f: 1" liIarnilJmt It K O'Nh;m!, R.l'. P.a'IL'Iillllrs~.,.NIl!1lIft!' 21117., ~n (:!I.9aO) .• ,I.. M .. 11. IMI2(uIIm:b, II.. III. (hPJl'El1~ £Il\!ili P:t~IJI!'ts.ti, £i!lf. ~, '5;] "1'J8:2:;I., s, eM, Gf4(f,fINN! ~ti, M.. Wt,]O,41 ~l~'l'8~l '6, II;"M, ~'tIn. f. ~, ~. (I]~n.. nOM, ]I!. Soc.. tdil!ti. fl:iNIl Sd. 9Ji, Ul f.200Ji). 7, ~I, III;.Whl&t, iI' W. CliapPI!li, 1~~$l'i'$'1;:!, f

.2:3. a 1. ITIeriiOlO, £ar.l1' I'IMl'r: Sri, W,t, :)3, 1851 ('1911]11. ,2'4•. ~ W, '!in~~, .l,a...~i~, A B. F!J.rIll. HI, f\~~, 'i. SIc WiOO@', 5cJ'tfi~~ 312.. lB9,ii (20{)G,). .2'5. A. 1:1, llUliIiPIM. L, M!lnll, W, 'U1JJ1n, 1. 1'1.1[)1"o! .:I], '100 ~2G!)21 2'6. r:, 1. !ila'I!'!:e-ll\!~h,. hrne~, R. GeC(lJ~,. G. llfiltm~l;

tmn .

P1~. Set t~ :z16" 'i. tlOO!~). 2:~. ~• .iI!, Wiebe, "" ]I. GIlIl!J!l:!, . S~cf. 1'!i~~~ZD, 121~ ) tii9!l'&1 ss, A F. (llHll6. JI, #,\ ~rllE'j, It S. ,OJle!llli)1'\,W. Gra)', K. Z.T~~r, ~ Tod~y '14" >1, ~20IH,~. 1.9. C. .i!tnlli!ll', Jl D. 11!11.mlljo, :So JI- 5fJ1llk5, J, J\erl.l'l1, 1117,W!; It.
~at:r}j ~2~5}"

;$, ~ ~

,~Oii! mJ9',L

~)]!ty, W, ),. (i~lti~~,~, T, "'~lMkIlill, Ge~!a9Y :;:~

Warry

that Ihis. ~\IO!lu~if.ln W:IL~ I)Unc~l<ll!!cd juveni I~ rq~lg~na :N.1pknlis;hl'l'!.CnIS, lJy The wctinlCd \riC'\:\" Qfgm!lirJ;!' g.t,.~lClo*'C<I~~~llrod
ph,lIall:'l ~'lugge~l.

by ~hc~jroon i:ilolap~ d1l!l;l compels n l:ca]lpr<li<;;;l~

~, ],. po, [I~o,;1!1l.:i1, F, ]. T. T~!)'Iey. S~<t ;;;:.15. :B26 (19971~1, t t Wl!,ig~, !il. i'o\;I~li;! I, Pi, m~hL'1!t C~ltft\, .Miltfr. PUl~i. 'l!3~, 227 (~O@)" 1<'. ]. It, V~~el. M,. R, ~oii~ 5rieIJc~ 305.. m. '{WO<1~"
H ~, L GIii~ffill .r!'t

'9. 8. W, Oap~~'~ ". i'letl'Qr, l'f, ,~.q9U.99M~

ilL

bl.l\wils61, BJ' ~~OO:;t

l"type nuilCS w;",.. m::~r 8::'% 1.011' oourgo" fO~4,lIot C J~mJalJ.)'oc. lind 50";", lbr WhyWcnry. IDl,C;s(;

1<1, j,. \\1, Voalleyd r:l~. f.ooltiU. "'fiJ7f'mr,. f'tli'fll1SIO; 5611.I.2ro~" 01' I~lc I~S type cOnCCI)l nnrl i~!>i.lnp~kaHom; for rs, Il l s. Kellip,. m ]. Whilel1DU5'l'. (. J. 1iIa~-1h,. iCfil$Li3i CH)i:lut.o.n. in ro"foCfll:ing the ui;wo,rking M.IK. Aian:Dllr f.amritl .Ni'it3t!fti.r, flemJr, ['S<Il!, 2]0 (200~l, ::mpro~nJ$.,a] nmterJ~l by jlllv"CI1~1c I'i~agm~~. our 116.. fI. Hlf'I~'. IL s, wmfaifM, S, Ill'. (hp~11\. A. j, it W1nt~, ;:;im:1:y:'~~~"l:s th~~t 1.I:Ypc'nlagllli.ui.~11~ Lc~ Imy crii f. GMt.. Soc AUi~t Z~" ill} (l'lM}. iii,\"tlh~:> '~IJJ]Ii"lI:li'[~li ,~I"t]\;,:If.i. ~ilb I:i..:ili~ Cali,liffiiJ. l7 •. The' ~.~"<i ~~~I)'J1ji!iJIm~ies d~.igw Ci~ .iJ1~ m~wJlld ~~i!,ovi!llllle rr'lImvkillll:l nandal\d rn~ OiIB\lIl W3leF ill 11:l\t!~i1 m SOH'IC c;,;,tc,~~by l.ifIC mJU- i!l1;JjlTltl~·m:1,;" pSfR~ par ~haLl!a 001.. isol(Jpt: rn1i,(H; Ill"", 1~:J,lk rocks, 111,.. 1J1!'~T:"I1~~ l;!l Milh!flal~ ind l!1e11~O!;!!. are .l!lf.lii~~~ ~I ~ip.pll!l!11i[i1 pro~l!UrhtJn ufnew mal~ri[d. a.:l\:II,..'tI~}' Ihc lachlrul ~Der1~~ @ Sd~IiI~e'Onlint,

or

or

30. M. ~. ].!Id:~11i,. iii. ]. CilJB~,IQ~" P.. I!.!~Bnon" ). (l~Jr. ". ~ 10a.~l~2 ~,1I. Si. (.otK~ R, S 1. S',PIIIk!, M.I':. ITO'Il, ,illr'Hijif~'4J.1. :~!Hi!' ~illOOU" ~:i!. C J, IiImgwllliln" iii. I. s, t:~mli'\, NtlIr.<J\to ~], :11.111 ~illOOl6;l" ~ •. ~ II. 0, NiI<Il,II. IN. (n~:P;ii@U.I. GiM1. SOl. ~"n H~, 5:)' (inn" ~. W~m ~detiled to l- Cu'i~!!i 'f,g;r g~H!~~a: wll~ ~.M ~ICfJe ·i!J1~I,~~Q1: Vtlt Edi~~!'1 !oo MiC]:"Q,ll'flll!e ~cilil)' .~~~ 'tD C. (Oilil! for ~.I~aJ1ce ~lh H! ~~ql'e ~\!!lys1~ in Sfi:;t~t R,~C~W~, 1M,f'Nrw!'I, 3111:1 , G1~~!1I genfdl!Jlill;r C pr,~ ~cI;,;b a.' Il!e '~MI f ~~f Jtu":i'. (:ons.1n!a!'iJl' o oo!1lm~nt~ by$. S!l~"'s, ii, (o~!rn5. ,a!!!~ IW~r~v,je~ ere '~rillel'~1Ir .a.;~ooWll!dgl!1il .. ~"uruMd I!J:lo I'JH( !J"all1l~ ME!I!'JlJ' Si'OO~,2 a 001 IMm5'1II50~

~2oool

e,

c:~

1'9. iN., ill ~k.

;;;stim~l~ ~lfll;p:lly Ihi11 rh~lli.'!I;·Qmi(;

(';l"u,'H !;I.n{:r.le

"tion m~I.';S nmy hav~ OC":I,l higl:I":iI~h;m hi~hcrl(} ~Pf)rocil'I.~Ota i,rultli :,;Iu{]ici> oil" plu ~o.nit Ct:n::.niC~~ ,(6, .llJI, u:u'!(iirying glL1ih;11 orm1inG'HW.l !lJ'DWlh I(;tII.r'!;:~,~hmllgh~inli!.1,

1'"Wo, V.rl~~ (,I!.I.. Gt.m~m..#Jm. MiiiiiW1" 88" 1.003 ~jlOO3}. ~, ], ~ ~!et, !!1,~ AA'iJN'l~I!~. 319 (2001), ~1 The' E\1l'l~1IIln,jJ!a1ion s[glllil~j@;l:I't!I'ili1Ml(J{If 1M i!QIIlP~ fa1~O ,(li~liIr~~'rtif w, lA~w.I!lt1'Nd~ '!rom ~ d\G!"!cffilk rtfe~el1(j~!!rI pa~~

:S8,PlJlortin,QI O.ntinll IMdll!lriOll


_.~~~g,iill]k!jl/(oot~nVjulll31.IDi!lt4i""98(]1!DC1

1M1~~:l~ ~nilM~'Ulil'!R! ~O»irM To!Il~~ Sl till SS;


i!l:ef~r,e!'1ces

:!~,],. I!.;!~k~. J, '1,'1. ~. li!'t 235,~~5 ~.{Im5.J',

'["n 'tIo!!UJ.'I"!I, V!!~I~, I, @.. ~~Itt!', €~rlil


~Jl1!

M~t

n '()I;ii"r

~CiI'I6,

ro..l:!I.2'~~jell1~,

us 61:541

~~@~

n,laml,l.1ll)'2il!O!'

IF'rl8 ct:Ullrel-1C,o n'tro III ed IPla 1,18 0 -,IF'IIDii d Fllow iin IC,8,ndior ChI8s,ma" M,8,lrs
1

con.'\,is!t'~t w~!h !'ippJ:cs or


It;)!l.g~~ (f!!l,.

-2 W';;

1111 in W:I\,'7-

Sj J,~'!,'i~I~Il~~ of ~~(),'~I ~GHo]~l

o[b5~r'IJ".ti(l:nl$ it r,om the .Iii~ghiR!~'50 lUltiOITl hllla!!J~iii!) S":liflnte IExp~rim~l'I[ 011 100,ud tiM!!' ,Mar'S . RecQn !lIi1i5$;;nlIO~ Orbiter re'll1~<ll ZQrilIil5 e.f l(Jci;ltiz!~~ Huid a:l~e ratirO:r1 kem~l'It;;llti{;l!fl jlnld b~I2'''( !ling)' ~ ~(lIngj~mn~~willnin ~y!11t!dl ~~pil$it~in ~~!lml C!irloor (ha~m~, Ma IT~, ihi~, f~~idl .;;!llt1;: 1 r~'liQ.nJ 1l~(~me4 wiithin the s'Uibsmri!!:.E!int" ~ ,geolo>glii~pa:Sl~lnd has: been eli:illo~da,t the's llrf.ac,eltilrlll.llg ~ sl.IImequellil i ,eros:ion. lihes<~ '~irldijl1g:5 demomtraJ~'~hat Hl.llndfloiw aklng fri3:CU,!!U2Swas <Ii medlalillsm bV whidl :s'uliJ,S'l! rf.O!(l2 niUlid~ Il1Itgr.i3l-edl UUlOlligh tJ;N2'~e layered dle1Pes.il:5. ~Hl!uured llayered deIP'Osi~~ ,t'! Fe 'l~m:5 ~romisln~s]tes 'hHijnn5tigi!tkl~ the g,o@o,logi!!: ~ifl~(lr~ (If water on Malr.s. (o!~u

..C'ti.\'~~ h PCf'!.'a.<;.~~C' Ih~L'lU:g1101JI!. hC' ~1i;lI;)J1e~ulr' ~ [ I~r!in~ o~liIiui;lmll!teJi.~1(SONI) tcxr]. Thcr~'rrn'C, tlilis flat-b' i~l~ dafkll~<1~C!"kL~ is hl~CVp'J'Ctcd as ~lurlidal derH"Il1i!s, I)f ~cdim<cm cOlu~'IIO~C'd of Cfl~i.ll1 :'>and. whll a po:!i.~jblC' COll1lpOtn-cm or ~~:g. Dark !11'1!'l'1"i<l1:\"i~~lil~Ihe umil::utyillg hedroc~. \
])1m), i1bll) I.~OulilribLlU:: W til,.; l.tmC
IlJllIlJi.b"

of

I:~IC d!u~.

.11111<1. gillg SC ic.m:c ..•. .. E1Ilp~rl111G11l ~:HiR~SE) (,~'1nICI<L U. 2}on bO!:lru lhe Marn RocOIHTm i"i>am:c nrhi~I,.'iI.· {M.RO) nat.;; ~-eI.unnI:t1 :inni:lgc:. tI.f lhe :;urrilct or Mar.;; ~~li:.ilha.\'oC CXCi..l}liDlJlI~-c.i ~1rily~mi rusohl·· ~ion. O,'e of !he filfSl il!nllgcs of M~rn oc!lIl'!lod

he :H.igh R.'!,;''SI;)·~1II.1 ion

M., nm,;rJ~ ~y:<t.... r m.


gioll orM;II!l;
(cqui¥;I~,""nL

uf the t111J"1,:·lrmOO :lcdjlln'icn'Li~ S!utly. bu~ ut 1"111[\), ~p.rC:>~·\I]I. wilhil'i Ihe IiIlncl~rl)'iug b~drodk[im:l bl!:call~": IllIGlhil~ u.cclrlu:ougll t:o.l i.ilH (;rosinrl Oil" pCP.ili~t5 in P~tl(;(; sllH:g d(;po::;~ tS.[J'Mfk n l1:lmcri~]1 L1l~ily alsu hm'C ocr;;:n lrom:>pmlvu ~rolli1. 1L -cl istmJI
UITK'I£liTISI.~ljn~'tI ~h~ preH;r;.'IH. hy WLm::c.
Ilh~i; 0:['

Tli~ ~um.:-

In

tht:

w.r.;iil'Lt:il:n CqL]11.tori" .an:- d':::lc!!::I:lhk,


111

II~·
lIocI

l~iig. S2). S~lrftllcc

~~lllR."';S ~O.26'I:n

10 QI'IC pi:llclll

~h~~~I:mpc" '[lr ,t,lhjcets. ruw~voo {It

?:O:r~

Loc~;1 I.G~xl!!lil.-:Ii:!hyml1t~ high $!lJri'lu;::..: nmgh. :-:ck"Cl. ~itll,"".r:-; will~ill, l!h(:' ~igM.tmi~"!i b,,"'li:!·
'10

acm~i; ~IrC

mdk ~LP!:lar\!IlI],y (l~U'[f;i~lin~

Lli~ :lCcmnuhnion

'by! lirUSE in Ihe k~w ,(,2S0 w J I 5 knl} 1i.~I1l- lnd darkAoncd ha!ld~ ,(fig. l .md lig. SI ) [ m~l)p~lg Gl:hil i~or~hc :1.~,ji>C~d dq1{ll;~ii~ wl~hi,n mWi may l'IiC v'ok~.ni'C" coUan.or L1'Ctn:luinlC' in W<;l$t":-l'll ttlildflf t"il<t.'l1!N<l tf:ig. ~ a~1i(~ '!lg. S~.), Qr.i,gi!] (J 5). The' dilrk ool1d~"<l,i1'I~r to 00 ~~atlying i~l m,l!lY <!i~C':l:;::,;lt l~lC' ~I~m ~o:cak~, Many (Ir one rlf the' Luger' cany,om: fir Ihe 'Valles Ihe <llfk-I«i~ed bands C'(~lbis.l of <l mi1<~mc of meIer-ilea Ie- oou~dc~ ofllighHOtllOO m<llcti<1!1 Wl:IIlilr<lnC P1i<IOE!iiliY I..~OOril~my.1J~~'o\!!rsLly oI,~rom, Tlm;D1I, and fi!1C'r-g;rnincdd~§k Inate-rial Higs ,. sJ; ~md A'l 8S,7.2i1. USA. ...~:c w'1~C!~ l~~~~crud~,moe h~d 1J,e, ~dd!lil5"lli:d. E'mili~ S4J', TlLC ptlIC!lCS nor rmc-,iM:~il!crl Q;;im mm~bl COila 11101111' ~cifL'Ve a h~l.nmOoCk.y ~~,nit\C' (hm~ is, (!lJi:>D@(pl.i!.u.c~i'I,li:du

The ~uYClood(reJXlSilS apW'ClIr

iih.'"

,ill tC'~l~tin:g

orlh~dllrk :s~"diI1llUI1!.in dist:incl hmdS (ng.;;;. S3 1O . S5). The dlld:-lfmcd. b;Uld s arc cmmnrmiy fGlll1d

wi~hi!~ tOf'iOgr<la)j~lc dCIJ)~'5S:iflll$i]~lhc

~HKJJ()r-

Iyhlg m..'(lro..:k. Acc\iI'lluhHiIlm; of boulders ;:-~l\o ~t;1 tfl< !!;!IP,l<!rk-I()"c(1 ~d~rm:m \:I.'i III ~!l the t~<L!lcls:

(SOM text). Ihus. the dmivtOll~;cl bands~I~~"'ar' ~lco"~i~t()f 5lC'(U~mcm I~ml hri,S,Iiccuml!lal ..i:~ " wi~hin .he tt,{HI:g~,~ oo(\?,.'een ri.cjg_C$ oflighH:OUOO liledrock.
11111$ri(lgC-a!'ld-tAAj,gil~ nlo:t"pll0log}' is CO!lsff>~,;n! w ith {liiflciClttia~ erosioll. \\'h ic:h crmbe

REPORTS
cx~ec~~-d if ~:hc bedrock is meclmlllcaUy If]),crca, -,iflial is, the mechauically \lIIcflkcrr·Jli}'~.'SOr rock 1m,,!! na.'>lcrerosion niles dl~n Ill!! :slrmlg;cr lilIyc~. .I\$:;;~lming 11 l~ol'nog,;Ilii:Ot~S dl,g.cnclic
hi 51ory, linlc'grai ned
CO:J r~.gr.Lil]cd~Dyrn::. JOIIIL halos rc-qlUUfC$ Un: bedrock ~Ioug the IW~tcllC'S.ur ~OJ~Q~lUi~ht; d:cprc:"Ssio;ns. [Hid ;1 t1)' i joims to IUlVC bCCUI sl.rcngll'lell.cd agli im[ IlfLY~iS of bedrock 111marc dork. The ~acik of erosion, Ridgellik.; s~gl1il(:m:;; of I~e l-roloo ~~1:~,1 t1arkili,,~(;rifLl ~)ohus 10 fln:c.l;lligih~.e !JCCll ntulinlion of O'lii. ~cdinnc:m illolll; ~lIc~ hido.:>, 3;:;, crosscut IYL'Cdil~.'k-~Qlicd bal1rls 0 scdimcnI~ wc:~~as II $y~lc nnmi:c lighl~nii~!,; or auy dm'k o,;spcci~,lIy III<.'CdiObe :.trcl:Il!;I]~Hed because, Ilaj'iJlx or bedrock w.i,lhil1 these halos, pl'~slJnmb Iy, these c1arkp ands he .a~ong

nm.'!Iu:lry d11PI:i!.'lil. (n The t'lL":lNcr&gmlnt1d !aye~ lCl.mdl..! r&"I.cr limn the .rJ.ncf-gra.ined h1Y('m, .~$ubing i~] 11r.idgc,<urui.!ffiugh .~norp~IO~Qgy J.<; :!oiugg.t';S;LOOhere.. lA'IC<!Il ~lcli~mgC'ndl.i.cs i~] ceIIn~'illi!all'Ol1 and ,oh(Cmic.1~ \vc<Huhcrh1g 11m.)" 111:;;0 iI1JU~It'llt:Crock ,~1liCllg~h 1m, '['hus. the ,d,dge...li. ~.':I' ;I!Ildtl",ough~ wilhilllh~ h~l'nck l1i~y reflect !OC.1~ v'Ir1'llion:o: ~.nl d'l11~1'gr<lil1 size or dkl~l'l:elk .hi~.flI~',. ,or 1'l!Gl1~], A~.'illIm~nt are :;;ct.~ of rf".cI~I~ 11I.nare hll,l1drG~h~ or fll>(l!.cml to sc \'tHOml k illome~c:!N i,n k':nglh tFig~- I, ,HId 2 and fig_ :5;,)), Shear di~ p,I'M~(''II'Il.CII~S(J croS$(;utlJ~c.ld~lfIll1. and ot&lcr d i $'C(l<lfniU'L1~I,ic... f1i'C null ob:;cr¥c(l~I(m,!,:l ~bCIc.~ii: I'i1LCilllrc:;. SU'l~11l h.or,i.muliiil di:;l)hlC,,;nl~lli;ii. (~r mO~,1! tlimi (1,:52 111 U\!,o0rkllld:;;) w(j;L1h.1 til!' clet.rll;.!Ubs~r,,"'Cd ~lli~ItC'SoJ ni.on o-Uile H:i ItIS,!! inim.!,;;'C, .til, u T,hetcrOI1t:,lh~-:;c r11LC~!lIre5 are id'clll ~'-icdas joints mh~'1I-than ~IU lis (SONI [ext).

hl~J1S ~ I\; ~trull~Ci" ~llIlu 'wilhilllth~ su mcoc>i:li·

MUlly jOillll:; arc cQnnimKlu:; ··indo·'

or .ii.!l~:r:L.wau;;dbo-..:iliot":'K
(irig.

Surat:.aHldcdhy

a r.1cMlly

lhal

!C~U:s 1I;;:Tt.~); I~IC dllrk·lClHli,."li bands

2~nd

lig". S3~I1.cl SA).. These joi nil, halos 111"1;; ilTl' ~ClfnJlpti,li!;! ~hi;;lmckJilIi:nUlid pu~t~1iT1 of ;i>i.::U in~i.:)nl

dark !11.. t~I'i~~1 Jlll.ocJimlli.ClII~>,W('.:.!kDI" (mol,IL'-~.rodcd) J~yCil's of 'lxdfock. Ihc: oombilnalioll of J. smootli :!Rlrl~.c(!' anrl 11lt]8i~iY,e relief a 1001~~h~ jo ~nI. h3~QS accoauts ror the lack or ,h::Cll nl11u~aliIlld scdimem al1d dllir-k Ji'j.I;n:l'a-ccs a~SlmoQ1.h ~Jt I.he ll.tCte'.I: :>.'CII:iIil. 1.11 ind:k;;l!~C!'i ~~mt Ih~ bedrock w hhin llflU,"<;e halo:;; 11'1 :!oitl'Q~lg..;l:r (mo:re in~hurnl:cd) !.I~~n ~lic Iml(m ~o.nll::JI~I.• !I~c ,~,dja.c'I.c"JU Ug;I1l"~o~'K:dband:s. show Il\':!ndiJ)' er"t"ld\l;:d bocilr.nck amUll,cl il. nu~nlJk_[l_~ C~C'ilr mpogl'aphk ~;.l~I~l:diin!:l is di~!i!lgl~is:h" Ihal I."IUt\Ci~)i~miClIn0.1' ~niincr:JIli' {C-g., Fc."~C'~rjulg OIbl!c i'm:Il":!,,dlx:cf.o vm:iiation~ ·1~~IT.ILl.gh II iJ{1SE'R co~or c~p,"LlJilily (F~g_ 21lnd ng~_ S3 OI!!d :5:41. o1.ill~mls} IJJ jm~~] rh.lid~irou~ .. dl1~ ~\'i!ll" ... Ille )lurr(ll.c~ ;lIong the joim hlllo$ .~~thcn:db,t'C' 10 'I~orc SI:laQ[!-S Elf l~lcmCk a~(mg~h~ .h:ulm.'"{~ illltCl.'lflTeICO to be sllloolher: al Ihc meter .'1GI.i11' joims i!> ,Inkdy '111CC Illlnis Iill. of 'lL':1I I.Ic"ITIDk stl:cn~lh.cnung, These m~nc.r.lIis am ro ~nnCfl~ I'CiiUl'I"C 10 lite adjaC"\:IU ~~h!" I1ml (_fmk=IDli.ed the wa U rock !-md ~hcrc.by illcl\:msfo,; Ihe FOCI;.'::; b1.1lL~ A ::il~m0n1.CrgMmdl ~mlooe nt'Ctll~, ~, I~k resistance 1.0 p:iuilug and cresion, 01' s:malllJ..:,.~lo,!ilOPUU'II)hy illM '~Il 'LeI to tlfllP li;.l:;Sc h~]o~, rlltli>i p~tlul>ibly :Ibli'ii'i'i~d ~lnCit S.:dillBC:lln. Pauems of ttlpo!:lr!lJlh ie :ill·II~.tIiJ1~a ~~(1 :i IltI~= ~lrlCjoil1~s ~IiiCfi:presem, ~,cllad Ilic hales [onrn".:(1 cmc '~h:l~, joi lin hrLlo, ulleu have ~. pQ..<;Uliw Iitsi. as 1]011 Inut:'ul[oo fllrc:'eiUlrliocali:y :SlWlllg ihe s .r:itlgl,.'S,. the j ui ru~.. \',\\Ij'll~cl h:l\'c prcicI'Cllli:dly n;hc.r ....itll .~ucliatC'rl ~Ut,frJ.CC5 Him \'o1(l'IIIM tend prop.a,gil~~d willhj~'11 I,h~ W(1lker rfJeIl;, adjm:x;·nI~ W,ull'lpo;:dc sedimcm attm,llulllti:o'J;l. By Hindu, co Ih{J~(; .~IilI:(10i. rather Ihnu nhwlIgh n.lI\: C--C:rnclt 1;1' wUI:h the I ~glflHOlilcd bands, tbe ffCcllll~tl~a· of I he hI! kI::; .tit' I::; observed. Pn;e1i.i~ lilll!; jeints tiCln of dill'" 5~'clim;;:111 .tIlo~]g ~ht blil(;"C orth..: w"OuJ.clllbo .I:~IC'ili tilt: local iF.::tII.toIH)rb.~~11C h~U~I.: }lHIHt:rlJ1 be inh ibi;l!Jd WllU'::!\: the ~]alu", tlt,e i~~~lrl (lC1J'i'I!.11Ialiun w ilhiu disl.iocl linear h!Lloo I"id,~chkl.:. A rid~dlik.l: IliorfllmloW" I.h.r '[he by a'C~illIl' OI~ cOl1d'l.l:i~~ rUl:r c~ N;: U11Ilill,g Ihlid!>_ III~hJ; absence of fnl>Cw ~·ool1~l'ollcd n!ilkl 110w. d i<l1.'t:llclk <I~I tcrn~io.n 'W'OlIlfl. he dislrih1.Ui!2(~ 11'1,. 1. 11HiL9Ih·re~olllli®1ilenil'l'anclI'ld· ~lNlugh.fl~ll Ihe IT.I!;'k nl<k~" "'!~ler than bC'in,!ll: oooor iilllla!!m~ ~I1IQ'I'lliln.g1!~lmd5(ape .Qi ·of sand dli,lfle~allilill DIIIUe-s aga~ II1s:l kM;:i1~ly-cd in di!lC~tc Z(}11I;'1> thaI ~s\:I)I'. 1Ji,,"ddlll~

The !iC'g,~igiblt· r:rm.{IUlnI

(!ir

:Ilionglhc jolln J mlfi\ j,ndie-.Jroz.; dml. tlm~ ,IK":J:i are u!!jh\'Qir:JIbl~ .for :scdimcn~ depositiDn, A Ilack (I" mete I":i,calc ~O;I)og~aph it" sluding a~(Jll'Ig the bedrock willl:inr~le ha~rlz.; reveals 111<11. lh~'L~

I.n

a back glr,O!u nd of li9lhHon,ed

{t.a n)a11.i:! di3lrk·~ooed (!D~u~' ibal1.d1i:

I(~.g, II ..

ltl~
,11'1}'
I.(i

iln Caln~,or (haslI1lI!3i. ifhiis is <I s.ubsc'!2F1e [nm H'iIRISE ]I1lI<I'ge


TIRA..JI;()QSa:6_.1740" wihi!i:1'I was a,cqunrer:l iOn ]:0 S.epl>ernil@iF 2006

"file- ~)'~te.!l~,~!ioC Ii~~l.e!l ing in tcmJt;l, pf

d.'uk hcdrock l~yc~ a If'i'n~thC' h:t~m IlClill!.'


~l[loCl!CnliC<lI bleJJ(:hi~!):ol' rl~chodrock.

j'\.~

(Mirlrs SOJuilhernwil'it~r)" ifhe imRg

E!

wi'l$I:a~l2n at. tlill2' Ifl~ai Mal5~Jilille or 3:2:9 p,rn" <lliId th~ SCfll:llIe W<!!~ iI,. ~lImii'i<il:ecl Jlfioonlille ~\<~~t~\1itl1 oil

poovi,{l~1sl:y m.cmi ~!1e<1. d~~k 111<l!~cr~.;d ~·n~y cfl\ntr~l:ltne W' !Hle aIlPC"lI~IlC"e- of the' d<l!rk_;hl.1lcr~ 11l.:ulds, C-s;llCC k. II}, on S~Ollejl.1 h)\'!iN~"""cr.d.~r-k.

l]ot.";(hIJeki,s J~cldnag withill die j{lll1l. h11,~m,. 111m, amy dmk lltil1Ll.cr.illll orugnrrn~l.Iiy llrQ;.eu~ wilh ill th~ 1:i~"drtlck u!lliClir:. It) im'>'!.: bc ..n : 5®lar il'lddellC!(,! a t19i~!ll1)f 58.5;0, di~:>(1iI",cdt1,r gcuch~ll'Iie:lll)' :lI~ ~e~I.o't1 i~hi:n ~.hc w HiRllSIE ( .. liIler;;!1 (QI[.~~t~1 imag ~ 111:.1 los. datal ~Inrlhree ~and p~use-s(bl!Je· nil EiIlI1h, l~h:1D ~(."Hdli~]~or ahc rock MI~' grefDl, red!, .ii'OO flfilf ....idll<troe~;4100 i .ro!ll:rLd~ 1II!l(n h:tl.ell~rc j·s 1~ .clem" uud i;c;l~i,on of lO looD 1'I:11lI) 0(1, 2), TIi'I~iiillage s~~D.ei~().2i6m ~~r pi:!i~, \\lhiJl;~i~ ch{imieal i~]IJ;;m~lion:; bcn,l!liX:l~ [11,(; f]uiillo C~t~ cull!~in!l' wilhinlilbat mlcturc <JInd 'WIn: Im~l. ruck. et1M1V<llelll, to the smile (!Ir~he red b b~nd~p.i!s.s;hr!~e, T"eqilh~r tNlm:l Ul,.12~. Addlitim~ IllY. :inacrn.ct~t1'.I'I" ct Wl1!..~l pa~SiE'~weilea.(qi.lli~ed ~... twCi-byith '111;1(: wan ruck :.md Ihc rlLUud:-. Ikl'win.!l lhrou~h lw() p~~~ bi!il!ili~g t,() 0,52 ml ~~r 'rule rlf<l.c~un; jlndm.~ d;'~I1~l,'5 in ~ilil '1tlC1l8~h{Nf ~ pbieL The' complebeil!llilge i~,cern' lhc wan fwdk j,j:-UJ~. F~::;l(;l.m-..::.:mpporli"(lnDW tered at -5,7" ~O!thll!d~: $~.6~IE is recognized a~ ,m ilblR1ifI!1A'1'l,1 pmce~~~h~1 2 t()ngi~u:d'e '~plaftel1H,en~rI(C" l'i1Ile\FFia' loci lilan_e$ Ihe I~~:gc"1'>OlI.IC' ubsl_lrra~e nlligrm i{ll~ s liion.!li .ASitr'Q\oomb:.al UIilLii:l1il2{)OO), or~l1.I:ids and ,!lICIt"! k.ill itI~C~C~ io!~s l'IiCI\\"ecn

n1!~

~].(IS~1'1(:~«'1'. v, IS. M ". ing () f!hc joi ~l1s,'''''all, rock. as \:vell; ,~<\ tilcgc(lchcmic.ll blc.lchi ng flIf Ih is rock. ,~ro\O kk:s, S1rol'~ C\Iidc~lce orl<llb:>.m"lilcc l1~jids h'l\'~!l.ll. eire ulmoo!i!llmgh til i.~~CiO!innof

~iK!se fll!l !d~· !.flcllhe <

The

s.1reng!he~1

~~e layc[:Crl dcr~(!!siT.S. These cpisoclcs ofl:i!tC1ld-

ing and ocnncnl~miolll ~mJLmhlji rel1cc! cpi"'od;;:s

984

16 FEBRUARY

2007

VOL 315

SClIIENIOE

\i'V'IoMiV.sdencemag.orgl

REFORTS
o retihe'llng and ('I1\, idl d u:g ~tl l1().w. as L~ cOI!l1iu'lQllly o!liilCu\d::,(j on EfLnil Ul I.!) lind also on M1Jl1'S U 7-19), These results s~IPpO~1 prcVL{lUIS Iindin(!i1,$ from Ihe Oibs~'T'l"mohfie (M)Hl' kl Mine:ral{Jgi,,... r[lUL~, lcs Gbec:>. c! l'A.cl'ilvi~c ,(OM BGA) hypcn,pcc~ll'a I inrmg.cr 011 bl"mro Mac;: ExpiJ'C[;:;. OM.IEOA ~h~i.:'rv;u[ions or lhiii mIl iOIll

nil!

dCllce for

oppm.renq, (1!, 131. SLICll morpl1O:i,oggc eviI'CCICml,1 SIOOfLe:riIlJI flt]id f!Qj[~' 0( e.g.,

:gullies and :liJ)ril~gulo!..nu:li$ 111$ not observed,


TtiiU$. 'Iil~C' P'J)(;.'Scult.dfL:y CXPQ5UI'e$ 0;1;'I~ii~ haioiXI ju,illl;:.;in.itiaU), :Ionulodwil hiin Ill'; ~tl:b:>uda.et,:lltl!l~lid.~ C'irclll~mlcd~D1:D"CI[jgh Ih~ h~j'cred de-

I)mlit~. ThCf1C" nll!I id.~ ~"lIh,"~qLl~mlly drnim:'iI ~hc~a~l1imd d~~pmlIls~"I:'~.1 Ih~ m~~ray.,mcl the!' haloed joints ';~'Cm I:lXlPoOO>il wt tlw ~mra!)~ 1!II.'OLI~h cro,..:iml. ThC",~rore, l~'!iI~· :s,IX'lC1J;I1 ~igmJillirt:! ",f .hydrntoo ~!)I ~:liu:c, w hid~ s, i~. .i.cw~~ as ,(Ovidi::nec' (I r pilst aq LICOLl~ v aciiv iliy donee of ,goochcm lea I ~nOGe5sc.s Ihrn~once occusred 'wi~!"Iun lhc slll~~llIr.filre .is CiJ.llrrunt~y (?O~11). No~all j~.iln[~ in til i:;;. s,~nt': ha,\'IC ha I,QS ~fiig. ex~cd1 at the sllIlrlb,ce ill ~h~ 1'0n11 oflh~!C haloed joints:. S5), Thls is p.~o'lJ,nh:l y h~aill~~ oJ ;[I. .di~1\;l~11ce illl ag~ fi:l~~ the h:Jilocdl versus ·"ali:l'Il·hll~oed" A !:;U!l""f~lrl Ibclll~of J\i.u.s ~Ul.'~"'.cc~XIl'J'i'lT". joinl .... Ulmlli:l~ d~\'i(llo~d , aml,md joint..;. ~hal tiol1 i~,~hC' inV~ji,lig;b~.i.C!uf,ifareOl~ thai. ~how o W~l'(" '~~O":o;.c'lIIt WhO'll ~lI!h~lIIrJhc:(!' l1uid ... WC'l'C' cv:~dcllcLl' f p~Sil hyJl'Ologic aClivi~y.• with the intent of Ch~r-:dCiI!L'iI.'i;dngIhe I):;lSI. h.<l.bi~ il~~ of a it)' cil'C~I~al~IlI:l th~p.t!~h~hc cmnm!.I}' exposod Icv,e-II ,of Ihc bcdl'OClk... Once" the hulk of 111'C ~he~ nreas and thdli 1).t]~ellli.1~ ~D :<>l,lppml ~H'c. Muc ~I mmiO:llI has been f.lllid OCIclla..:;si:c a , 'vo~m~lesUIb.~~lr:lacc j1~lids dru:inCld rromli11$ 3ref1.lIHy 1"c~IlII..'\1 ~'cm1~m" :O;!IIchas dl)' ;r,"i V'Cr~ and ~1lk'l,)ood;,; UI''':-Wjio~~I1~ that fonued would :rnn have

or

locat ... w itll in nterc,giona I ~.~y"d c:rcdscdiutCilmlty ~d«d.l1'lC: linL1Jirri~}i, COIllltltOJ1J oriuDmloa], fut(] pO$i~ivc feUd of t!:lese: features SU,I!:!J,C51 Ilam I i!IJ..~ arc fr;;l(;~!I!a\.,-';; Ihillarc surroL~t~cd by bcitlwk dmt has been QI,~ull~ai II)' C1;:IIUmlild or onlimf'lvit'olJ illi!IIJ'alC1~ ~~ll1ihJr 10 111l!.l [alms dL'ii'oCriix'(i he'J'{l in C':mdlll'.u ('lms.J-na O:!JPOI'lLlni~y is ocum)'nt~Y:lt VLL::~OJi:J. Cmlt~r. and dCl~li~rn;I sl,u£iit'!S or IlIU.\M.1 ,·idgc.il rn~'y pmvidc~.dIt~lianall il'l.~igln inla, '~~n:n~oh;;lD!ics :~ml(ll d~CIIlili:!'i'1n)' of iJI(Ik().;f1~I[d flo,\'!/' IJ~rn(lg,h 11m mgi£m:J~.~cdil1En~my b>odllDCk.
gr.:lJlPillically
Rf'ff@iIl1i11C@5 .IL A, '5.
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,et tiit}.
~j';50. ~'"

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2. f[t marl! 'i~lxllIn'i!lIiDlii 00 liIiltl!!E. Si!o! 1li11p"l!tliriSe.I~L anmllil.iI!dw,.

~~UirpM;.:d lIrlcquut..: lluid now I;"o:r i )iwl,)rici,cni ~ J.ir.iiD!IIl'il of ~ill~'il.: 110 HI~uw fur 11,1f,; ;:':1WC:ll~lIic~iinl& :a nidlJkneh~ Ifl,!;!, of 111Cw.:all roc'!> Iii {)ce~n-: Ihut>" Ilio h'L~O is :prescrll. "lHaci:etbrc. I,h~ pfol..'f;";UC~ nf
.1iI~ltUi:\may

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CX~tLllli'iCtl

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for rellm.i",c:

[t'JCIUn;

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w"ilh ~ltc Im~oed fl1lC'[lm:::> b..:ing lYle oMesl~nd ~hl: Imrt"~lili:oocl :1:hIC ~Hr(;S be ~Ilg~hr.::),Ol!lllgCf>1 .Hld h.w~~I~ f~ml,.;d lifter tho.: :o;.Llb::;lIIr.l~L(;~ nillit1',~ dm ull~d troll) b:;. level Ih,.; bedrock. ,[h!l.:pre:;i:~u:<,,: u f limi-I:mlm'u joints lhm apl'lal.'CnLly :Ilmtd~liC!' the lmll\'lcd jo:int~ i!ldi~ ocale..<;!hm halo 1i:J~'f!l rlon, and thus eirou~aliofl .. of ~HlbsLll'r:iJcc 11I!1i:d:s, within these joi.ill.1\l. wa~

or

Ilrefj.~ in whiC'la i.tC Ii ~ illl!', I ui lltltli~lJ)Jll re F~g. 'I.. hal,oed. j:OJlfltS. ~]r'!.: oi:):;ocnccl irl (lllilCJ lhIRISE images t)h't~uawf".iul ila)'cD'd.! d'cpOOitii ill Vm.I,Ie;s, Mrrruncrls ;:ut{1 elsewhere, ,lFmdlcr fLIll1J1y::.uC';;; !hcsc liit.m~]roomiily yu(;llt1 ad(jgtiomdin:;;igllt illhl the getlCh";:ll~i,'itl)"

11l1i::; ::;'1iI.I~ly(bil()~i:iinnl.:;S tllf~~ falliI!~ lilt: abo PRiii:iil:.; i~l.!!l 10 ilw~li,~.m.c r,iSI, hydlruh)~.ile;
flm:i

MiG~~ Srfffl', ~ 6Mt ,Ml'tg. _ r-M10 ~l~ic~, ~~I'I'~. [;I;eirrl'~r, CO, 19~:O, '1. 8. K. Lli!:th1I!.!lN .:ll.. M~.!i ,~u.Jfi\i,~f AIlilon~ 11'm$, T!Y~O!IIl\Ail, 1i;Wl~"~p, ,4~~~9". 5, s, Mli!rJjh.l!!, 1,., JKillla!"ld.. f>. ~~ l I,'-"'!mnil, M, :~~il1$I)'''' !wrur. !lA,;!. 4.ij.4. U~', oil, ro'l. G.. ~p:mall, K lll~~~, 1(t;I~~ISS. :11~

~~ N. ~1K1b'll«k, (. L.. l~~"!:~. D, It, Sruoll, Gt,ru'"gi~M~ ~. tile vml~ MQriMfi:~n~M r'I/ tlr.:tN, s~l~ l:Moo.09!I,

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,rmd li'l.iDBwm@tJ

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g.colo.gh::aUyroc-ccl1! eveJ:u.r.ufI!hc[. had ji(lil]iS !'j,l;IllpOr[cd gcolo!!l,k1llliy ~~CIH !!l~ilr'-s!,lr~h~ll~~id now. !1~~'li<l! ~i'O::;;io!!l<ltand d~l~fE~iljOr:l'1~ SIrl;u;;~~m wrllll'd be .. ";5: 111m.
<I

~hc hal~c,~

ruck, DC'I,I u~ed ~l,IfJhoo Oh:.1Cf\'OII in~'I:;~0 r Ihictlill'C" cnntrnl led Jlilil id 1100w !1li1Ybe pm;~ibIt1 will~ ~:hc' Mm's EXl1'lor.uicm Rn,vC'I' Ofll)m'l~llllil.y. 1\ sety.~·m.,e l~iR.ISE umilgc 01' VkliOri1l! C,...cr in 'M,cridi~ni PIa!I~~m rc\'eiil~:;' 11S'Ct of ~l,Ihpam~ lei linc~' fi~I,~c~ ,~Jnu~ ~~c cralll2'r'8; C~S:h~'ITI rhl1
WIlli

the

Gie:JJcgi[~. SDmlc)o' DIl!iiMDII" ltllmDI1" 1'J99}. '1, ~; ~iImlmt &. 'Ji'\Hr!!y" ffu.id floi~ arid rl'll.rl5'pt1rtin l'ii'~ (GM[I<n:iali III Hall,. loo!1lcm .. 1'J'9'1}. lO. N~.k!nllll ~~rm G!lIJlIltiL1i'1Id fm<rul't':l cttH1fJM/Q

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tim: (OiJ~"Y UIIikt.~~'ApprkiUll'Jf!~ ~N~.ioool ACMlcill'l.~ f\HB'!; W~,t.tfingtt!n, DC, l~Wi). rl'I, f4. e~i!I,lll, S.II<tl.W!if'\,.'I\t"T .. l'Ialll)',~. rmi5,.'G., krrlaltiJ, O ,~ Soc. Am, rtlday 15, (M, &I. ,4 l2'OO.~1 rl'I, A. (~~Ji'I,W. t ~.:lfi),.1, II.. ~fii'lMI, Am, ~ .I"llt ,Gt'qI:!lo!!IJ. 1)iI, 1~~1 (2~}. hi, .0\. Cih;lal, IB, g,!iii~f:r,.w. T. 1~~Il!)',J, Orma, G, i!i1)'11:.a1W, N~Jl,!re,"~~, '131 !210@4;)_ B, Beitter, \'.I, 'T. .P~I1f)', ,PI!, A, Ot~m, /. SeQ~, Rg,;!,5,
547 ~2005l,.

15, ,", 1. Pa,rl)',,~, Ii. otlal1" 8.. !'j~i~Ie1, ~~~, h!!. ~ 19'uI.t IIO!I~ 11'5 {2ClIM!l
16, ~,. Beilti.er. M.. A. OMil, \\1, T. (lOO3).
1":1'. a; c (]a~j{e1' (2005).

rn
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sd, Idt. 2~o.. 9~,

~~5;1, ~~OO:5;1, SiMMJI ~[a.I,. s"*c~~ :nil~ {H!Q!Il1. ~OO lli. Ii. (i6ldfll, ef',I!.I,. :Sr~ 307, BET (<W.IOSI, l~, .,- t:'. Siiip1QIl et (;It,. ~~1lI! S~g.e IfJ/ C~WI {Jiwiyje; ((ie~~ISoxi~ !14'I.>!!Il!!~ W1d~, :200;l~, 1",,p,4!:l.C-Sa, n ,Pl'I, !::. Mi!~iil', K. S, lI~lt,. tv. fQ~Iiltl;iI@, $. 11.1, M~l!ey. ~ ~. It.Io;ie Iill!~~, S'o'el1c-e .3:!A1, :I!S,73 'C2006), l:iL We ,~re ~r;ne4!!~~' .l1e 'Ki!ol)l~, '~erO!HQII5,~m~ em~i~ee!li.,g!t~(!'f ~ifUSE O!!lo!Ilh~ ~F,!O li1'f(Ijem:. ,.-.t1~w ~iligeli~ ~Uoos malll!!' lfui~ WIld; IlliIls~IlLe,((1Ifi:rn!!nt~ 111 HiRI5E !JMni, 1Ii~ ~md hm ~1100.~tilrul! rNi!m!I~ .:l're al~il a~~r,!!aatl!!!l. li'«Mn~ !Ilf Ihi!, 'h'Iillk ·were otlf'lIllK:11ed 2{1.
.~.!p"

l~'. N.. ]. lb:;r;i:I ~!.aI,. &NII P!.QII"tf" Sd. ml, 240'. :122:

~Iiif't;r ooiblr:ad WiiUl INIMI\ mtli a~diljooal.~1l1J'Hll"l ~r~n~rro.m i'tJ!I5A's mal:; D,IIi1 Matr-;i.S I?or~Ja:m,

,li!r ,al

Su,ppoftin.gl
SOCIi!T!i:~J
lFig~. 5':1to 56
:Rel'e~i!n(1!".;

Oni[i!'ill' iM.<!l1eria'l

~'l\"",'ir:~e!TI~!l!i)I"9''l;g~!l<!!t~!,!Vj!.!Ui31'~a1,.4.~.rocl

Fig •. 2. ~A to () IEl... mp'l.es ®f ~oj !'It5i31111!1 lJ!IIJiol!ll1tiiimg h.atc.s. 01 ~i~rl1it·toll1iedl hed met S 'lhill'l dawk line<'J.li()I"IS"Tihe :scale balra"ci nOfih .il1".rO\l,I' alPri'~lfroe.ach 11M1~t

1111 jOitn.t!: ~

alre

.tl1~

l] DttdM!r iD,

~f(~~

19 liI~mMr

2006

ro.ll~;emDl!.lB61l:550

REPORTS

St,u',c:1h IF'ossiils and the D,ome:s,li,catiion and Diisp,er~sa,1'Oif Clhiilli Peppers II Caps"icu:m sp Pii t.)1 in tlh e AIl,elri cas
3 4 Lilnd.a. P~Ii'ry,:llIh~ill lli)id;:<lU,~ 501111<1 .Zani ~~o, 2 I rr,e-rll:! IbI1o~st., De-boiraI11i!t'" P'e<lrsi!li ~I~ DiOiclli'e'S, R" t'~;peirrl1:0. lM.aJry)a iflf lit!e'rrrmi!llil,,5 :lI.3 IFI](IIl<lr(ll 6. (v,o~e~ 3, Kiln Radelllfli!lk~r,'& AlillholilY ]1. llilalilofii~, 1 l- S((ltt. R<lJYITiOlild, D!<I.IfI~e-i.!fl. :5.i3Jndwe:i'!l'!ii.,~,8 IFralrlz SG<lram,ell~l.i 2.

of,f';uU'cl:i grrcin il:>i>I!mblr!i,l;,'ts ~romC

lI\mmll1~1 and

Cfimt:;{'ffj't.~aJIC·s.o~~ir:tlil'Mn:l:I9I.~l [ibsClilre,oifu ~1te

KaJyii"arrlble,?'O]a mes A, Ze~liIllelr'1l

diagnQS'ti.c .• i,1~s IbOlpo.'>Sl'bte lO 3s:>igll ga;flim" ID {I slllgic .:'i~l'CCI~. Thcil1.orp.looklgy of.:5Liarch~iulil C, drim'1lc,·(t .~ ~ilui IlIJ; ~.oml~,Il(li~ ic!er,I~i~l~ U'rJll ,of C. to al.lJwnm !IlT C.P'llJ(!Sl:ftml., and the I:IlDIlphdJt¥ICS of :di Ihoo~ iSJtmch~ cli:I:ThrNlgllil:k:mt~y r:!1Cl1IltilOOC filf C "u.~cl'(JJmN !h11d C .rmh1!}if.':t~tJ:~wh[dl,. intusn, \ di~rer ~Iio~n om:: ~UlOlI]Cr.1::tcc.aUl~e R~rl1H:xr~.YpL~ oeeur In mill mlgelll.lric :lpcd~. orG:~rj\"k1m!., '(!Ltlhcl" c a di~gml.oolkClra Imlg/? ald~oullO.mL~icala~.t'f11·
hbgc .isl't'lLliroo I'N<l8CC1JI11):~1>I)lci;cli IID;:uaHication. TIlI~prn;iJllre of !I blllsk gli;[!.Ll~diagI1M.k
~~h, u'I1orphot.y,ptil llIl'C['JlIOO onoo

(liiil]

p~PItie-r~ «(ap~i~lirns;~p.) .an~ WidelJ (lLi~ti'o!(it~lroo(l pl~Iil~lh~1 aro~e ilfl(ne AifrilencdJ$ o;ilfldl<Ire now 'ilfl(lJrpc'ra~~d in,oo(uiSfilJ~ \Oj~rlt!~'litl~,. IbI1er~,~'I'I! repor~.3i gl~.rli!ll$"~e.(Hi(starch mtl'rph[Hyp~ tliia't pr'(]!IIi~~~i31lin!!;:!"'" t'oid~ iltiry ~ntli p~p~~ fr(i'lT~ arch~eo~Dgi(al (orral'eltl;S and lra:t~ Doth til ~,i r dtllil1,~nil:;aJtil'l!fl dli5Jl'E!tsal.n~Si~' ~tan::lilrnioofl'l~i'l!S Ihlav~ b~~!11 ~rtd fOIL!no at ~~~'1lI 5it~ d{ltiilg hom 6<0(10 ~~(Us. biHifir,~ pn~~,niI. to [l!Iml~"'11I ,oon~tlct flllI!li rrangiilg '~f(J'fiIIUII~ [la h'"If1,a~ ttl, :StlutlfJ,~ rill Peru. ih~ st~m;1iIgr,ailll ass~l1IIbta!Jes d~IfIl(J:ni~r.~I~ ~hlQlt !iI1ali~~allld (Jhm~$ ®(c;;uf1F,~d(lgeUl~r ..:; ail t ~ f1d~nl ~ fl.\! ,",ide~p'r~d 'NeI;Itr!llp3i(illl !;Ilant 14;JQdl ~!llllil j;lleJLthat P'l~dI,;iIt~:;I;I~Uery in ~1;I1iI1~ reglQn!~,

Yor 0If'_~i(.1m., i~ Ill'edk!,iI:l~c I~ek(Jif Ilil~nf~ ilolf:rim'l1 to ~l1Im!'11e"

drlC hytlridiz<ltifm l·J). C (1mlmlm. ('~ l-himmw.'. and ('_ _rmtt~'t't!n<; h:ilvc bt..~~l~~<;Cliht.~,~ .~.~IJ!:dcl!1 ' cornpko.x willi" ~illgk nm;e~r;;)! .gJ.~1e:pool! (4)_

Therefore. i~i;$ n{)~:;urp:rising Ihal Ihe ::;m~hes of I~ICi$CIb'l1iIX' $~'Clc:;. .til\: iI]orpholo'g~C'ilny sin.'Iilfllf
1:0 one

spxl~~~~';Iiiill ~pxll~lIivC'. A OOrjilbil'i11liOll or arc~IilCQlq~i(111 ";,,,idcl1~'C. !,!i.'a-ICI,ie .~~1idy:;,.-:>, aifld , modem p~tlm di~u1tiJ!lIIll;{m:. hili"."; k'dl rrr.~nJthclS _ mll~]~y in .he Anlil!:nK~~ ~~I~•. ~.ih!lT(0. WfUoi illilD~1 miy domesImnm~.~uro~l1ldlih~ globe (I. 1~ T~IC IfICk o~·11 [0< SII£'.!!;'CSt that c.wm!{!{lli COiflpir\;llo,;:nl.~h~ arehacebcmnical It;cun:l has li~uJ:.'d iiJ Me;.;ico or :rwnIK'Ul Ccmro~ AIID~J. C Irlill1:lpc~d ~W:ulrruh'; roclll1.-;rmcllons Gr'lh,~ OJi!linL';" (mlc,,1('r,rS iu Ihe Ou:ibbt'au. C. hm'('(J.f,rm,r in ~Oi!iV· dtlin:;;,~1L'C'1Ji:oJl\:i. ~ml di:q]ll;r.;.Ill~ tlli;.';;.(_; pl~mli.i, I!tllltl Boli'li~. C dril,reJ'l;xl,! In uotthern lo'!'.~lmld ill, .MacK'iI1.'~llIin$ t1'f i1]~ l:1;ui~ ;il\;. CUll ~IIlL"tl to 11111.: Anlil11ZOl11 :lI1Id.C. JJIliK'S[,(,:lk'! in nIt~c ll~ii:!-dcv!ltj.UIll ;:.iO>tlllh:l;n Alitlts P. 3~ ;:;~le~ mit! 'envi:mlUil'!.::llb, and n:~ioI:Ir~ r iil;.1.l!;!:: IIIld nli o All fi'IIX: ~~ic~ of dO:iWl.'iSlit:::IJttd ~ilili fK.'iPlx:~r.. poll,"" ar~ e... I:c.~ crnl~rnltm f[:lIbik I ~ \V~ ft'1tmd '~n l}rOOlKl~ 1!:lI'gt.; fl'lI~ti.!n,"-d,Icmilcildar 5tl1~iI gr:;~In. ... tlrl;JJ II wi~.k':\'Pf(~ltl.but prcv i:Oll~JI)' 'LIlrlidcmi:l'i(,~ an;;lmooh1~kail :s.tan;;h 11'1!OI1~llOlyl!lC' is derived li~o!nl "vilh .<1 :si1"'l~kl'l,"'CCil11ml depl'e,~irm4, nfl~ llHllikc <)I.. red <chjl~ 11()(lPC!' fl'l.llLs and is QOrillIlloniy 'I:D~SC'I'\"C'(~ ,on bl!ood C1::lm ~Cl:u;mcc (Fi,g. 2. A 10 C," \V1H;\~' in .mi f~.c1s,We do.."UlllCll!",d ~]llri!'i Illicmj~sU. tmlll l:aI;,l!cd into ~kle \'~w, a CC!l!r.CI .Unt'iirllg-u!le ,ill :!I~\~~ a~h;t~i)k'l,gi~,,1 ~~!~ 1<I~_,g.i~~~.~hlln thr.:- ~1c,"Irl !inl;l aT Sf~!i~ ti~~ wi1ih ji'll"~) OOgp1 !T(1i!1-\~ ["r<1r.llkl ~o Ihc IOI\g :;r:-.;.k;. onh~ gr.U!1. nitlfi.gI,I~C' lJal1!iln!~~ a~hiIPC1~-ro!o }\llIie<Ul SfIol,llh I\n'ICrlc<l ean r.:-xto::odtor thC' cmijre lC!11rtll f.if~~!C' gr.liin or jl;.M '(Fi~,. l~~~l~liti~ 601}1)}'e~~ "go a~hle2}. I) The rive' !'!IlOSl ceOi'olnk,d~y nala~,lc ~pc- a nm!1!of it '(.Fil;l.l. is and f), R:i.l!,gjl1~ tfi~11l ill(l1.l~ 13 !II45 p.lmI~ inlc!1l.,g~~I.!he st,u~h~ flrdmn~.icat<ed dei': of chili ~1~I}Cr>UI'C C imJ.lU~m~.C. l1IJ;ol:tmum" C diil1l.iJ1."i(,", C_ji111r.·.w;,,·m', and C tnjrnt,~"ft/l~- AI.L I~PPC~ an:: elIsl~y dis!i!~guis~'l~b:lc [m~n~~mllkr thO!!.lg:h ~I isgClnC!'<I.I~Y a:g!:\,":()d ~h~a IIlC' .@Clnl;~ \\'~I\:~t)1}CS tn Ihe- m~m~oosii!X.'Co!d (Fig. .2[) ntl(1 Cap. ~i~~~m. ixocu cl.I~li""I1!!,:d ~.I~~a~i"'~¥, h~~VC
Ii i~i [i'CPl"v:>, [it"':rnl.io;:(s,oflhii.:
J!;'l:.1\L~

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(27) {28)

16 FEBRUARY 2007

VOL 315

SClIIENIOE

\i'V\MIV.sdencemag.orgi

REFORTS
we Rwi1.tl d".1H sUlrellCs Oil ~in~ jTIruc;tkOO iu.bctll re@;iOli5! bcfo,l'C l.J'Ci~ilrod~ctiolfl (~f jn .liI..\SQC11!1iouwllh rnrM'c. aJliI.JWUOQ1, .:Iml.llhc p::illJ!!:)', m'l1l~~uisof~!'1ifLLm.cly So'cllIum sp, (]l!Qtmo) Q 17), is We ruiso I(mnd Stfll"chcs ofc:l'iJ~lipcppt..TSaa tbe Thc:l'C diliUl irldi'l'me 1I~~ll Iile l\,'SidcUis of \'Wl}illtlUtli. Thsee Dog $i~ k)C;;ucdm~ Sml Sa1:vil.'dor [$~lifld 11'1 <COtniYJUmiln cvid(;it;X,inch.lIll~'Ig piam. Ni'llfiinl1> ~htc~~Clllmit" This ::;u~r,;; ....,,~ (M~pi,.;."tI ~'Y a gl'(l1lj) of ;t1l1U.s~c p:t't11'i.inti!}, 1:0 ~~~OOJ~llly flood-xl botltllOlill.· were ~~llnva~ing1fii~" h.ib;~.11~u;I WCP1i!1;P.J.and n litlhc,["htOJ1DClLLllllIr.i~L~ about 1000 yr 13:..!l: Relp, Ih~1l11 irl~!.'l· ood. Oll ~iro. ·\I!JlIymm:l. r 1~lId, nndte'dlJ:l'S, ~gI.1rnllllU'e wm. imp::utmt. ill 111c \\'eru p:r~jl~g 111m' ~1;~c:!Il.tillg; I]~ 'I1l:1lmna~ ICl.n1nll~ of ~~~ le:1ffi one mlt~1illOO .['1. ~ommrnia.<l orooth .EctI1Ldm"im ~iL~ EClI!lt.kir i rk11 yid[kd the o:n~)');r.:ll"hmK<i~1nbl~~ II~i~ ... t>1](JCi:~..di:Jg.nDlinj{:lillm'PJhotYlX'_ Tlhr:$IJ ehili pcJ1I)J[1f ~ll]lL<;dlOklk.lllc :;.i.1e M~1~ of .1. IllliddC'n. [\;\10 ;Ji;',. m cmltoiidcrod 10 b;Jnh~Ct'!lI)N' ofdm~I~Jliall [Qi:1IJIY [i\lILy.","'" and a IDW<do:l1c<:.ily (\\ 'CIh... cpl} mr:."J_ W .orlB1f.~ Im~Qr t'()mllOO~K:::SPCC~Cl.O .of chili IlIL'I'lJli!lI)\. .~i\IC ~mrclm,s ~,lJJ)L"ltW 1.0 00 dmi'\fro~bmlll C f.1f.d)(!so_"li~ • F:i.!Iy.ci,gln chen l11icro:li~!18. all~typlC':ll.ofll!t' mOl'l1DGl~I~T.!; tl-c p~'11re{)fdi)fm:5Iikati.1d chi!k~ Wi~li" l!he spo::'ies IJml. inclllldes, "3riClICS SLtdh.M i1re reooto p~l[)kl1gy OOl11monly dcscrih:;;i:l <1It> nmurrn; gIL'"<'Iler' iLl I.hi~t"arly. Cf.!IIllp'll2i:. agricul[U1r.lI1 .'i)'~lOI11 bldirmc.\ IJePpt:ll'~ <11. daili I.l1m.h Clb~Li\f.ned ~~ I1IW"lltitud:= ill il!he Andes ,~.:?~W11~1 ,wII11bfn".,;i. wi.1ih Iman~il l1batl1lr~ IlI~UHS I1m:>t It;.\~ b:;1cn d~lil~'i;lica1cd Ihll:.ocs 08 -'Oll ~lavc~I' ~''i;)~d, .:J~ \~ C\ i~~C' ('fahl~ l], the ~n.I~ dam ilflclirnlQ' ~hrn ah!1 c.crn!l1i~ .~riddrot;:sllml'd~ ~11e l.1li.cmliths }~itJWc(!llm 'G1="il~~ ~Itt"l' "mil {:tmy.rRIF~_ ml;l bnng.ht nl1l.O ~~hymb11;Iin<; or hrllh ~llI~jzcand tmidColllj,licd Cldlh·.:J!Liol.l or lhn."C .11I1~lk::llcd !i:pt;\d~ of .chili t90 n;:gL'OIDrml11 cW'K'II: tlr '!OJ'l1! Of t~lC !'llltJ1lrl. r [~I CJllUIr:I~ Palll<all"l. Ihe Aguadlllkc .Rnck SIiLiI. Ili!lP~ "\f~ mll!CI!~l)[)J"'I~I~.o1l1 IhL1oo!ll!il and ill fOil)!;. 01" !lIlix'l)1, We 1\;"C(l\{lm(i. dilll :,;1<1I'('Ir.c~ro:m !ilm highlm~ds (If Penl .a'I, ,cmrly ;I!> 4{MlIO yr B. P inDhlJ' two fl~kc:;: !hatiJJ!lo CQI.1~ijo~S1<1l'CI!lCg ·ofrnauZt;'\; t~'W \\'a'l ()cC'liItli,,~d from .. h.out 13.000 ~ 32flO y!: Lasl!!;,. we n.'i~V(,'l'lJd mkmfm.~il 'C'IIidt'ncc ri:"w B_[~. (1i(lrillI,l bolh dIe' P1f\,~·<.U'Il le :;tIW EIT!h' Cc-= l;Ue Pf!.'OCt<IJtlUe pcriod. TIle 1'I.~C!'bl'e' .ofnUI~mlli!: oliher ,,;JUII;tuvaN wi~h ln Ih~ a'\SCIllb~lago; flf rncll Qh~ln I1ePf1!Cr QI l.M fV!allgo.... ciej .P~...!J!Ia7..(!. in .r."I~l lc 1'IC[j{l(I!i U 3)0. 11he slre h<l'i. )~(?rdQde\ideocc w lor ~hew1ilvm~m of o;t!;crpialiis nm fliUivr: 10 1:eo";IWl irtdicm(Sl!hlaL OOJ:i.llIis~icmetlllg.riC'liltU!U:C as \l'ciie'lill.'ClfI,fL ial1~ hflhl!mLun she OC'~lLpkd.l'Ihrnn :5{nM~ICrn Ccn.ll·tllll\I.llCrW;l, ~lldL!tfirll,!,l mi)i~" U'Wil' ioc. :tn'll ~.qua~lI~ di.nill~ l-n_maUI."rrIJlit 9000 10 5~OO if~ble 2. Sll!mlml!) or INel:)t~opilei.!l .an: h i!le1Jlogi\:al stil.r,e:hes !!)f C(J,pSicUffl. F.,llaked tOClt G, ylt [loP. \i\i'e itlCiiUil'ir;1d dil lipcl>jia' glflrCh firi 11 gIH1!!llHhkl n~ ~~~t fleramk sher~; So, 5i~dinnel1t 5(lmpte" (, ,b:ITOtL11&lilI1r:: IOO~ 1It..'iOOVr:n.'tJ frum !hc !.Op of the ilf. SilliIiIllie pIT.ocr;:l"dUl~ O::pOSi.l};, the tOO~:llndtI:lLl'; 11:; ssocituc~ ~ S'IlIIx:Jt K1ii.dl1CS 11.~1""'c1 S'I:GLlill!rJpll~C: iliur,; of ROOlll ; .1:"'~ M'~\IllJ05. ~elJez~er~ (-ArQ~QlljimtlQ. V<!J,rloi'd') 5'fiiOO yrr BJP, Ihi~1l(i[illilcl ,fqis.oytelrlCd 5!m!IfC111tlllJi!J:; - 5mHIlOGU) (Z 1) IL,e'!t '1. 'lL iL lS·G from n:lalr.!lc ilm.l d.orllc;;1~C'ill.cd )'1111:1if 31, [, ~ -SOO-:IlO(lO (21) lleiOl' 2. 1. iL The or;..1,:!i.!iJmiono:i·~t~""m~~1l!~1 ~ldl·I~.ud!:k.'n site -500-1000 (21) !Lev 3, 1 il. 15· ~ or ZrlpotiJl coineidcs w:i!h Ihe ~'Jr!y Ceramic .. 500-::1.0010 (2.1J ILe~ 3, ,2 iL 2'2Fc p~mJ ·tlflli:i~.It:~iori,,1'fall1Irilli.l, lOC.llilinili.!l ~ixjll!l! ILev 5, ,2 ·...500-10010 (21) 19,201 Fc 2. 4::':00 y~R:lIT'. ~14).W~roc!l}\''-'«ld snmvC:~-s orchi,l·i ...l' Lie,., 7, 1. -500-:11000 (2.1:) 2 20.• 20'F, IroIl1gl'O~mdslm.c ~ook i!!dkat~ll~ that Ihe 1hr,rie .C)'05}> ,B'wlt:!'r1Ia.l: ~,tt.~far1J {Icn'a), U'l.i1nloe. o;ICi..!t'bi~s (sqlla:;~.\, CmrlWff#U sp, {i:JtO; L'I!!.';;"LI1~ ~le l\.J:o.-accac raJ'Il"'y (f!rumsj, hss and :abo hccn i'CWvcrod lrom Rail AIIO (!{Hl~ .1\
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'lhe rounded le-J1!IiHlilar Iorl1ll1aoo l.;rige:.llat (00,ual. depl'm"ie:n '(JB~ III'dlla~@llogjc~ 'Q;p~rr;,tNJl ~

U~ f19~'1', i9. ro!, j,. ~r:rn~\ I~,;;;. a~lll"i!.,im. Ar.liq, 'it 219 (200!i~, ~ 2:0. I#.!t [)dre~, Am. A!'l~J'q, <l~, ol1l9 U9j15~. a~ fa~b!~.l!\tsil. oo'!~~.Y '[]]k~9@,. W!(i!~o. iLWlO~}' 22. L ~,I, A~wi. SrI. :!l'1, 10019 {;l'OO~,t 01:3. i. P'M), ~~D'~Am, Anr;!q.U" <100 U:OOiSJ. 111. C, E.Smi1h. i~ PW!II;UNY oJ~t: ~!l!(!(lm~lJer. D'. S. Elfl!l!!\, M. O~)l!IS 11!.rm.·. ~. 1!.u~1fiD..l~.sn.!lp .. 2'2(11-2'5 ~ I

0'

:slllrrn grnoole (,"Om Lerna

.A.I.!.1!. {t) AliCli'L'leofogkal :Slarrn granule ,of (,~sf. {tim ftrl1Pn1 ~a1 Alb:'h {m

5tardl grram.!~ [rom a modern specimen 0"1 [I!W' sfnlmGlll'l~ var. mil1~mum."Tlm!;5~:anchwaiiu.lle ls tYlj)iI~:H of t!m~e fr~m ~';i"kI P€ll'pe~. (E) S](je VDe1'1' g of 1:1 mOdl!rll st~m:h gro!!lIIlulle fr~JIi! Cap.'ikuJ'T! ltq,,,,,tttm. NQt,~ the Hrle;:!r li@!Jlfl!" (F) S~d,~view ,r;rr an N~ 19'!!6.)'[).1},~'!!-7f. ~ r~llI~el;l!ogli~~. S;lalN;ihlgr.;llnul~ I;If Cop#r;tlIm IWlI;ml :Z:~;p@tll,. ~ll. ~~~r.lw.e ~Il:ria!l! wt,fiil5i.li[l'J!lied
SOO[ol.) ~ooayr B;IP. (11), Sc\;II,,'ruI,lflQ,;;"':, deep sl(lIi~ ull.ci:lllCl:'!Nt[C seallC.rW o\''Cr 'ntc s!lutho(;: of line silC,
E,'lf..Ciii\tU~~l ,of ~ u'li,;!ldclil ~.h;poo.~1 yic~tk'dceramie ,griutlk hl~n:Isand Ui~i.croIlLlihl.c lh'S .. ilt ure (lJkl~ ~ h a$:;.x~I~(;'tUV!'I'ill] liI'1llIlDoe~'>5ing (1 2')1, i\~. t the a TI'i!~ !Dug si[C, lihc remai 1'1.'> or mi.lIJ~OCIlfC ,con:;i;pil..:l!IU!lIl'ly U~1I1 ~mrn111'1 o.:N(~,"iuui'm ~~IUI.y~;;::k!ed ul1i rli.t1.5 LL'ilJmHy iL,,:;oCi.lt~d wUII~ m~nioc:proO\.';S~ in~ {2".n T~iC~.tlil'tc 'lil'~X:~:;;~ l.ooL.;.~~1I! mj"Uilil'i",~ :f'lltin.:'YUl,.'\S of dlili lli:pper :1Ii>t"1 t1lmL~I1l;U :It!rmLUrl:.. or C m~i.!.I~. [{oot cffij'lil. illdlldill!!! a[!'m\TC!Ot..Aii'.IT)$lfl!~ s, I.. BIll\s., R. G. O~!ell'!I.. S,IIlJt: 8m. ~2.,S·>(1'J'9n.. ,6., Milleri\!l15 iln~ l"Ile<lhOli~ .![r~a!lililabl~ iI~ ~up,lI'Ilrlir.g1 mabl!ri<llU nn S,ri~c~ Online, ,. ]. S, ~c·nd, in ffJriji< lori~ America ir.J Pl..ru~1:l.I)':' [lie fWul~ llj'/in:JIOic Q.fld.i!'"J!!anllril'e' (.ui!l#~. M. Bla'll~, ~l {Wiljhin.g.~, 5~aJ~ ~nr.r. F.reSj, hJl!m~ru.. 1~9}. pp" 'I4'J-1:S,~'. a D', la1l1ra~,. ], ,(I. iMaFOO$, ], A. i'!eidt£F, ".tral"9'1 J~, 1. ~ 1:1977). 9. ], A. Zti~I~. in Arit'lilll!'ai~y of l'i)I:ma!~ frutlilo.r~ ). :5; RIJ)ftiiJr;ltl, R, il!m"Si~F. E~~., (1iI1limbIJrtO:FI CMt!, wa~hingloo, DC ~p,4'81-S,n.

1'5. D. t, 1.I!1l.!l, 111.~. BB;iudlil'"(I:I"~t!, 111.I._ oR, del Agllli!<ir, l ~!llall, ~Gr; Am. Anriq,. 7:, 2-47~19~6i'" ~6.. K lr\ii'&e~lJ, Am. ARliq.. ~4, 5A {~9iM~. 2'~'. l A._ ~m, L S. Wi~\ Q,f.ll~a(i' (1l1r1 ~~ itlo.rrWii? AmiYka1l ,[J.m ~f 8WlrJgit.rJJ ~1Wlm .i~'1I~ ~ lrJ.dli!'j ~1!Mi¥, of Alam~ft'!~\ Til:Sca!ooS<l, z,004), za r. s, '5p.~nt~. E. M.. R~dll!ir:t'!ta, M,. ~m;!reI~ Am, M!i;q. ,!m, S, U9 U'!l\l~J. ,29. ''''']' hliifian, p,,>Gnlmln.. WilfidAr;rhlMtltOl6,'121'(1'l'5'5<}.. m, ,A. J. lltlmer,~. R, 'G, ("~iJ~e', iii flOthS tQ. [.JlJirJ:ofi\mio!1,~ Pr"l1iUflwy. f. W. LiI!1g~.€l!!L OJ!!!'>'. Pr& 01 Go!~ado.. !li\!"t~~, ~M, Nal!D!!~ Jk:Jl!lan\!!ll\ E. [I.e!!)", .1. If>efiij', (!~d ., S!l'i!l1~da. 9. ,sm!lll pr'~ a!m!Ji~nt~Gnll'ie-'1Ji~i!ilmri~:U. f\J~~!~!lj fIN ,am:hil!i'Jllll)gjcaluc.al'alillfli; and ~ldmjh ~ram~1mi~ WiI~ pr,Drid'edi IJylne' hnBrii::a n ft1~~OjJIlil:;aL Scdi~y. thti!, Co:lllI:ejII de D~I!lDIlB (ien!flioo 'I' Iillllllal'lr5tioo lie iii ~riiMl!\I"Si~adl (i~ntr:i!U riI~ veI"ll!2~l;3, ~ (;;00$ :S'l1Penru !\WtrotrGlJ del I!itlUill, !il~rr~mila1iam filr fqJl.ardlfi(m anlli ~e'.lIrran Dlil (ultu-,al Ori~i·D\ Li}l! H~im (Iila;rilaMI! frusl la:lfiii! Arnm~~ mh~ ~I~~ 1'~"O~r~m.. IIhI! NSf, OI~~~Qllh~ PrtlrMt ~t llth~c~Coi.I~9~. ,I'rllgf'llm d~ ih~ AA'iJr(j!ld(l.~J:i,~ra ~ U!..l~[fj:)(,tli!! :wn11~I'IlI1I~n Nil1fi(if!~I.
WlI3i!tiim, 01 f'J:lIblJrr.llllil!l~. tI1~ :5ml~DFIi~i'1 Troplc.'!l~ ~~lrdl IM1jM~, 11l~5!Xn~~)([.Iil~ aM HlI!bI~iI\.iiil!~

:5"p.{gl.mpo~ .. nail member

on.BI~ .Zingij}cl<I.'ccnc

rll.n]ily (gill~C'rl' a ~S{l k:1'l. thei r .~~ilIlr(;hy~c1l1~ains. Wh~~ll co!mbhtcd wilh the dma fhno. ~hc Tih~e !)og ~.il!.()..he (.h iU Ill~~l~:r lli~Mr(j~~iI~ 1~~ I
Los MlIJugQ~,de~P~lrgUil!.'''uS\!~a'lill!'i!~he
J'!iJ~iL1n

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j,c,

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(olfidl jjf ('.an~th,. Tnnpl!!' IlI!ii'M!~lfr, ih~ Ulnlm:!ilL)' iii tlIi5!l'Jtlri ~~~r!(l11 !loird, ,mil! W,enll!'r·Gr'~n.

5~lPortil'i;~1O.n'liiie i!ltateriOi:l
MM'.f.(l.tM~g,.0rgUgli«ntcnlillJW31'ID$1!'!\I"J'&j1OC1 I'Mte:ri~,li ~j'!il M~1h~

·th~~ ..

k:OIJtOO mi.'OC!d ~'~ll~jSl~!lC(!j 0Cf.i11{il1~)I oOl'oolh root ,tnd seed Cmt~ OiCCulli\Xlal!hCl1csi~l\" ~ha~ \:\l~~'C'inir~lUy cmcgor.i~d as l'IiChl,g fi>Cv~Dpk1d hy ulio.1f!iochmticu'lwriSi~ {23~ Nc~jler I~ticrofn~ills~p.id of ~lI'iki~x."'Cics o:r o
~0i'ihi~1

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SOJIifhl
,pIg. 51 TaIltf!S ).1 and S<1

Re!tPence'S
J!lI >01;Qdber2Q, a;~~p1~ 2:1 IIl~BI"Il~l!;r .200~ m1:lWY;~ell[4!.1I.13 ~n>1i

COO!>~.

H'GUlSillOl'!.1j ffll:n'lSf1f'C;(.qMfmm were ~O\'er«~ ~ij'Qill ;[IITY~~C. 'l~ il~<:tlO;:' of don"":;1f;m~1 Itm1:S. 1!I~"'d a:;: C!ltldiny.,u:;: ml.her li.'1IH.mSsrupl\.:: liOOd:l dUililig,I;,.; Pi'L'"('I.:runuc pt..,'lioo Dm.lim[(~i> 1!il!l~,~'it!f!hj:'li,(:)lclrl3¥n;. CUl:lvLlrc~UJ!d COUlIJl:x {llJi:;im:'l>,~ {mJl' H:1Imt:Jg!I:LUl!II U~IC j\Jn('ili("a~ and Ihru lhii:I...'""\plo:iIl1J.1noll ofrfllliilC,. IUJI cltif'ii rurld chili IEwtltl; SJlIt.oo IIifC~JC HriltOOl~

'[iOn-!{~r!}1.k1)'•. lEvid~,l'IX (rulll 'iJ,O[11 Ilmlt'JOOO'Im!llcal p mlCi mllcruholrwlic:''''~ 1CJ11i1~1~~lt1lirul.I,;;:';tl:1U[on;;c dhi~i fX:1P\P'i'fll l>t,;cruJlu: iml::,!I"I;]1cmLloo iluo '~I); rli:t1. i11C)' stiem ~e~lls,OISC~)':~(aUe~r,ed ,1I!.ollgi~ Pl~'llisloo. Apart rrtmil ~hl: chil~ iX~~p."rs.lllil1izc il; iffile1lid lilt Dwsopfdla m~d!gul (onlai FlS mu l~i~ote;ntintesHnal bv amllv.>i~, to, generate both ,enlE!'lO(ytte-s prt'SCIlI~ ~u... '\~]y ~il"; ..: SlIlll'kd.M'~~ and duili.~ b8i~men;t rrlulmb:raIFLe~'alt ltIave ib.eefl slhlJ1M'J1 1~I1.E!'age w ;fXl:ur 1~llm[ rmm lihe 1.I(~hi..'t. uf Ihi:s 1t'IL"mrl l.I!IJIfi~ <ll1im entereemdacril1,e cel.l5. ISC'S (o:ruain~n g high! Ilevel~ (If cytoplasmic [le~taJ·rich w~icil!!S activate t h:e ,canon,ka l N'Men Ipat.hW!3i:f a rid d®WfI-r~gllla~e 11)e{la \\Iitli~ the~r daughters. a prol!:e:loS ~hat III EIlIm;Jl!..:lIU omlllli:l .'H"id, I.l1tl:O •. mpK.... ,,:m all w.K~i.. " . 1!1 I1rogram~ Hle:~e d!!llil'lIhterntD be~olille emtl1'rOl::yte~.,IS(s, that e)lJl're-ss UUI~ vesic:ulate llelti'll. 'or are' Nw1:t'lll0C<i1 pliM1! tixtd C([tHPk.>;" giMeticaHy 'impaired! il1l li4ol<t'i fligf!lal]ng, :>pecUy their .:ICiLlightell1lliO iJ.eCDI1IW ememendoc:rim~ >tells, Rerie~ei'U:ie$ a.n!!l W/jgtes 1rfilus, ISu,mntml. daughter (leU fa!le by modlU'lating N',o"ll:1tI ngnaHrI!g overtime, Om studies5ug;fjest s 1. C. i~.~~ .11';' iIlEot:t(if~ro cfC,,,, ;.rall~ I~. W, Simn~n:!~ that li50" aJc:tiloi1e~y cocudiill.ue c:eM prot!luclioi1i wi~h ~~a~~iss1J"e r,e.'1ll1ir@l1fIerrl5, by' tM~ me d'li3iriis m-" lEd. fulf13lf!lO!n, Wri:!oo" l'!l?'tl,.1l,p. ~~s.-;l!!la.
2. \'II, HI.. [s1l!}a1M.l~,. ~ru 1I'Bw. GlOpS, llamid: .. ]. E. Sirmmll, IEll!5.. U9 ~JII!l'lJ'. ']3l='l ~'J'. 3, ,~ iJ!iI~gill. if! ~C~r~i;m PJ'o.m~1~&.II. 0, s..;Il~, Ed. ~~!'ii!rd l,ilnii .." !\p~'l, (~ml)lIi!Ig~, iWI.,

oc

Mlullt:ilPotelnt ,O,ro,sophilsllintes,tliinal Sleml Celilis ,Sp'e!ciify' Daluglh'lelr Celli !Falles by D'i'Har'Bnti,81 Notich .Slgn,8,llin'gl

Mf!)'. m..... '(0,,,-

:!.~a~l'. ,p..lOHU p ~, 6. 1I'"lli;t,ffig!l!. 6itJi. ZeJilf. llM, 331 U9:6j).

. 'niches, w!!!!)$i;!: nl,ki'OC!Jwirol1.!ne~.. 1'Ci~L\."'SSl..'S, l~rtmmliillC'dHl[c~c'll!iali(m and comrols proliferoVOL 315 SClIIENIOE

spccllic

',(.'l.In:e lb ..

rulf!1j()llli:C:1!~

in adL~1l~.!CS ~L. !~~'~j' ~.S;id'::ill : pos~t~m; km:rwu ~,

p;:nh... a~'s i~c:ltldil1g !lM~J v tll(lTl.e 1l1!Orp~1o~n~'! k rO).teiM, JA:KiS"TAT (Ji~!tUS killa:5<!J'Sigrl>lu Ir.li~w~r O"d,lC~i\ialN of 113IlSCt1plionJ.'Wm. ,1Utd

~iol1! U. 2}. Scw.lml dmCD'ei~' ~~g!ml~rnil~l.c~iD:i~

988

16 FEBRUARY 2007

\I'V'!M;V.sdencemag.orgl

REFORTS
N Old l'Umniii)UI in \NCllJ,clmmct~zediili:c~les su;c!a as those 3lliltC apex of!:Jilc iJlwupln'!fJ i(3-.5~ {lnd t~~is (6. In UiC C,f,t'III""' I'lmhrb'lk'i t'!(?,'lm,{ gon1Ld lfi).. or wllI.h in "-CrL~1I1rnc 'boncllmliiiCFw l'!.l.W} M:u...-t orlllC~ :>l<;;n'l>I;:cll~Ii"C 1 kllrn!"I~ 1,0 I}l!~<;,:iih:dy roc~h't, SiUglllllls, Ji'Olrn~UIf·
Q ...

m1:o~e

.~;l)l!Inding::1lmm~J ~~~~ mh:uIl1ihi.bil d~IfiM:t'nli:Jlio'll, ;:md to !lpntlml~ ,I .:;inglc lypoC! of" dnLlIflhl~.r et1~~ I( whic~] 1n'IY spoc.i'lliro YtlUhCf afu.~r ~LlbtiCqLtlml d lvl ~lolfl'i)" Reccmly,i nncsli 1:Ia~ :~Clll cells I( ~SO;) '~hm Dl;1qJuireNm-ch ~igmllil1g [or nomlal rUIK[ion \'I!1;:fC d~~:Il1>Cld inn~n~:/)nJ.wpilil(1 a~~uh I)Ol1tcnor IlIlidgu' (fig. IA~ (II, n~hll col1tmfO~ to pl'~ v.i.(lu:;ll:l" ~tudiod DnJc.w;~pbit(J ::;;;~~1I1l Ol)'.U!\1,ISC's gC::IIlCCllc ~Wf:l dHlIbmnt C(l~~!y~, ~llIh~rocy!.C~ I(E(\~and e!tr.::~llOOeL'ir'!el '(Cel) L;'"dk Wilihoul illiter'!Cllin.!J,d id:;;~(lIIsaile! are not assndatcd will! :!i(j:lOCid1(1I:I1OlIomica~ shes oonmin iillfl candidate
::>tmnlwd l'lf01UCffi T'i)i]1i~Ii,ga~Cill~

ii1()lcru~ru- li'Cgu]milJ:rlor lise niRlliiiXlitcl1CY. ~1IlliIl",'dilud,~lIi:;; wiih muikxllic:; we ~JiOOIliic rQ(' llie Notch I~ttl Di:i~it n.u-;:, :.oc1ldyU $ltitC u.llri~imi~1 OO:ntllili.o:I1.",(3).iSCs m'C JiJL!I~(J prririil'Uiil!y WilillllJll r;m~]1 ch~cr:> of 1M) er 111~ dipkJid aclls, n;..;11 m:';S[S""; filJ!, ~Bl di~ 9Irruu~ t~1C nl)tio.lay~'1J" of po~Yrr.oid EC~ Iinill1,g ~~ICgl{t,
H~~rd Hwg~ M~k~1 ~il~l~IJ~~, D.ePiUlill1~~Iof l~i1iIl!l~, ogJ" Caroo~~ Ilns~IUJiI1on W\util~gtOIl., 3502[I S~m M:llltim ill
,Drl... , il:~Ltin'lI!Hl!, MD 21:li8.li':;A,~ ~

on

Wig. IE~, w;; ,ob~cr\'ctII DiI,li ih;: ISle Wf~ aII!!'fIY;:-:, [)ch:i.p<)~itivc. wh\:n;~IS, pfCF spi.:eti".,.; EC~ or ee ~clh hJek.ed De~t:lI. Thus, iii.
opi icn~ siacks

rell tor VI!t)' rrudy Dclill (F~. I H l'. lAIiltOligl~ lilt'!! DellijJ b-ct Iii Ill"; positilYC cell 'Vmy {oomp~re f"ig. :1. C andr)-}. ,W'lbeiirlgis, ~lJ'OnIiIJCilU kl t(ll~e 11Lmct_tl~ 5t~iCh!lI\,'l>~i.'Iinbl illg ";:rtdY"(;Yli:c vc"i~~" trig. l, C ilnd [)1_",imilm: 11::1 obRelrvcdin allt,-,!, dlClSt'! ct~ll" WLlih :Ictivc D~!l1.·NQ1.ehMguoling (!4. i5~ 11m cell t'N(lfC'l.'liUlEl Ddt,u was ~lOW1] 1:0 Ib.:l nim ISC' ilygC:llffO}1.i nill m~rk,_oo SWill cell clOlJ('S~ 1\01Lhe (lh~itT._C!d time, mcs of ~lC llflpmpJ!,nc _gt'llooypc !"\'Cn:! hc,;u:·~,hook.ed to il!li~.ku!C a R'JOOlllhimUliQlI. mcdlfmcd iJ)' IIlC ~l:z}'mc- .PLp, lfil;ll. ~.t1iv,<lft()jlhoI'!I"hle. '1aeZ ~l\p~ion hl :J"aI,(', fandO l1l dh,jding eclla I H~ On tbe hasi!l~r f~~t:;I'i(nl§i !'rtlld'ie;<;,(In ISiCs~.1I] 1:Ieidcn~i fiodiu Sltt;h dDlles: as ~he iHI" tia~ ta~Z"pO.;'i.li~· ceil, and IiIIC' .~SCIlI:.II:;I~lI~ is kllO""'ll too 00 Jnc,ut(ld c100,'leS'~w ~h~ ba.~~ll1em mcnllmi'lllc. Guts; were i!>flil!ncd m "'lnf.l{l ~ times H rlcr cIQJ~C im:llllctioJ'l, sHl:illoCdlhr l.ocl :lIrid l)ctL1L .-mel tlnfllY-ii~'d as l~nc~·c3iI1lCll:ii:OIl1L~ l oon~;liils n~ill~l,;

EflI.~ oclll1es!

I.lcaring lhi!l;l'epoocr
".:
C;".~).I'CSSCd

WO::foC"

OOIllIlIcr.;~fciu~'l! for [)dt:a.

l'lilO cells) Uml. cxp~

Illgloly in UiC([;[~l!gN.erodl Ualc:cntcrulhl8sL rEm! is\_',,, whh high 1J'i..'1~ :lc'\!~I\s. (,rig. 'I(j} [.UI W.fL'> ill t'N~:gmm:d :I,",;;C:I~ tf-IC~SC gi~ ~ (IF~" lU'~, ln In I!:!Im CCU, nc:sL\l \vjlh ~.\VOl)._o.:Ila-fl!'l~il:i\c mn~,IdliaQl" I
.c'll$Ctvc(l ~hcl~l, the NOlet) P;,lJOll~1'wt'I."

or

{]fIe ~'i,tJ1Cmgly m{pn:,.::~t'fl,rl'K! Nl'l[ch 1t11:;n:ll~r(Fig ..I, ~':~ and II'h '11m ~!IlHlItg ,III\"''R\.C nJ~a!iom;Jlil~ i:emwt'cn I);_!~a hNd~ :.lI!idNrnlCh ac1.i.v;LllOIl ~llm,~IJ]dtha1

1SC':;;~lgllal vla [)C1J:3 III adh!1Irc Notch dl.ugh~CiI'S (fig. .l I~

in

Ihcill

a roJ1mlsill!l?: ce~~ mrni!.::qrl clrn~ (I J~ or ~;m'Cml dillct't,'lll Not:eh Im~lway ~GI~~ 10 i!J"lsI.8.ll:ol ~'I)IfI'fiI!h. C'ib- The- gn.. ~lLl[l~OI,;r!! 'c.n 1)]\':iIdn {G~IF':~ nmi:1;I' ~!li,) and Imllnnt ~()1'!C 1I!'OOui"A,;d ill such ~~rJ;;,'fi· !!I1COl<; sqgrugltC' illilia~~y iUlO j!n1l-~ d1e l:se orjll!l;l il:;;
dI.ugh~cr, )lidding IWO ]'lOss;ibk' Q~l\l:lml'CS (fig.. .21\), J:SC Cio:i~ of illc Imlli illl~k:lJI""cpm {O'} [.If!JId\JWcl ~.hc:rCliuliSI.-:-;>;pt.:Clooi (Dd~l ml ~ml!iill is ~!pll'KB ini;;::-timlJl Nul,;;il ~i@.ilwlialg, .\o·~ruJlyJ)dlfl l'IjU,DliI,IU, cl(II~';:;; grc·w ilj~!o !!llIiwr.-! ~F i:!:;. 2131 imji~ili""lj~lliIlhl:C froll!1 U~ ~1il.:lflik"tlplfC~iml:o:liy by ~il.liloj (J" /l!w('h [Ull'lc:tlull ~U ~ ur illlo IJIH: EC' and 9 Ilm'JUJ.it Wig, 2C). because ()f'lIflJlJsi~,'1~ pi)n:illlmnol: of

We

g.Qr1('Cllcd

MAl~:.C.M. '(mQI;lIt:

a:1ls:iII)f:;;~ wI1ih

em

ur

·n~ ~liQlin ~~~~p~md~ii1~ !iI~~d ~ ~P\HI~~~w;r..dlln~@dl''!ellib.~~

~·rn~~

llll: ,~.d1l1lliiiitl,gJUi, Dc:lla b'J\.pn:.';:;;[m~ seeves .:BS,:!I, Iligl,lly spce.i[ic SI.CJ.TI cdl.un~1Jrkcr. All cdls w~~-~u ~illc :l'LC:>I:s C'~It."St> N01Cil. Oil (fig, I,f~, C:~mSl,;(~lK'lIltl:Y. 10 rlctclll"int:: ''>ihteh gJJt CI.;lb Ht:tLJall~ IUll:i~~ NOl~h:iigmds, ,,'C lM;..u:.~, ili'lll%'l::liII.: ruril~Iillinlg 'illrldk:,r.Ilt bilJjilig ~ii,,;:; !fir cBh.: ili:l'l]l~riJillimj IIIC'I.OO> Gl11in~yliu:~cl1Inltll uppn.;:.."i-OI" of S Ih~irl~, ISU:(UiU)! Inl<L!ac:~ ""~~ ~l"ilh'C' ~l"iJl;;~!'I<;,r or [ Notth actiqllioll U 6). \lIrhcl! gull!\. n!(ml adlll'l~

[)';1!l'I, protein. Pl.TdlmUlccprobrihly ult-.(I c:.pl~Mill$ ~'h}'. like ,v- lJJ~n.o:rs lie Dr UIIIi.'J)fS an; li'l.,'('I~L'i:Unlylllltlslic r{~I; ec-likc ocll~ '''_1\fl'[(~i]l§ Ptu:sp;;ru and lo:r~SC·lilk:;._~ t'Clb Ihm. bel!::. P'n:l..~'ItI "F:i~,113). hi rl'llll~l' oIHi,.,r J)J - (;blril.~, ni.ii.irlloo ~"!CIIII l~..: difr<,'r!.."II1!i~cd ~ 11. t-"oillgk EC ~F:ig., 2D'l p1JL};umah~y 'l~~I,L'iC OllI~Y1111 IEB \.va.; IIlLIt,"ltOO ,iiUrd il rot~ld ~Ii,~~ ..1

on,

Fi~. 1. IISCsmually rll'lfid:easymmemtically '1Nilh re~ec:t ~o Delta .~xpress;ilOn and N~tlh l~tllIW~I~' g'C!i1l,;ilthm" (A) UI:l@a!ll@ ,i3l11011t~lii~ (11:)' shCl'«S that ]rII~ivid'ulat ISC~,gli;ve rise ro d<llligh~~n '~n;. 1J!2~oone tliiaJt 'e.ntefQqrte~{Ee)", n:dll~ ~r>e~IJ!e.rlt~ ly rnJter()endlXril1~ (lelb {Eel,. {~} .M;idgtl~(jells exp~m. Delta ~rerl) a~ hlg~ !~~15 l"'rrm~, ",15.0 (01 .or low le'O!'e~sl[all'f!l)'!'!II'!.e,ad, al!se(IDXi wilh~!ll iI;I!il~ Ci~U ~f I~'al:h diplQ~d ~eU. !'l!l'St. Herea:n.dl inl flilla~es 1i'1~,u.elll:{1,iJr,es, D r4Ai~ 5i'h.o~m iflI Ilt-Jie. ([I 3D r,~· mMIlf'UiCllO:Fi1 a 4-~af lise dOOl,e' of '(g~~oo~ S!'hQwi!1'fl~'h!ij~, the- '[le~'~~IlFlM::'>ii'l.91 (~ed) 'i~basa~r ~1i;'1 (jeU Dasliie~ <lrr!l}~lIilfltli(:alIf5 cmller (Jr "Ilon-e ~r,m. ~f) INQ~dI!(red)' is ,eJoljlF5Soed \ffiltiely. Er.~.ir~lpt mature iii!
~1I!1)\l,!1l l(ii!l;~ltiOIii 'IJr ij(~,(ilFrQiI-\I).

E:C~ (!!r >e~ c@lls, ~6) N@tdil r~@Il~iQI"l


IiICl

~g~'e~I)15 'lIJNe!JI!J~Medin Uil: [ju~

in Delta'!ilositwe iSCs ~olrrowt (G1 f\l(i'tdll recep'l:iolfl ~epofter .d~l!l1n)~l(white) aL@ne. ~Ha,rldl HI' A r~n~'!ilE!~ wl~htl'l'llD [l~~"pcsi.tiil.t!l!' ,c:e~!s.:®wrs as in {GJ. ~U Mocle:l: A ( :5ig~aling ,(a rrow) in i~s dau9hter ElL Scale b.ar in ~B), 20 !J!lIilI.
Deilta'rich
liSe oi!(ltwal:es No~clil

I~

N ~i,g~id

~@)
Iilkroo
NlIirt'lifC

.~.~oJj~

REPORTS
[,."(!CD,",,; II NObeil Si:b!ll'l.'11rem d'tC ooj,ocerliLwitd-lyp.; ~ 15('U'~g.lA),l1li~ l~li11~li\.'S D!!h~1 is rIOt nCl.."(]oo Ihlli! Q~l~de Ih..-:15(, jin' 1::(" tli llillCilllalnOIl, oonsi~Clal \rith Ill,: fnpi~ (ljo~ViI'lFl\:glUt{ltkll1, I:Jclca [\1'I[lleill or
levels ~n ISC {i1mghllO.YodL~ h~ rui\]:tlIffi:. dOUM n'ILJ," 1:1111.or :I.IlLIU alll~1e Or;S;t'j~""Je ,(,Si'jftYM ~ eOC>t:JdIn!l [ 'nne OIhl"J.· nliajrhr NOold~ .llugmiitL s1m~\\;':d \\,ur.d.ly~

liI:'OI:.Jun-a r(Jf' dhu~lI.cr relL~to 'C''''' II.the l1liltll.k cc~~ cycle mid Jilrrn:Dl~i;]lOC: imo E 'is.. Otl1i;:r NOIih pillll.l\V'o,I)' ~1:tC-S\~rc I1.'lnQ'IoIC(1in Mf\R('MdD:llc~ wi~H!ill~sc.'l 10 r!llrt!/lrn' lIn:3ly'[C' IgC s~gJ'l;;ding.J';bil~di~~/d I~~!) elu::odcs ,m 8 V
fO!' D(,~~

OO\'t~ClprrlJm j FIg..lE~ ~rhl2l'i(l c1qlC~i,'lill:'!w'~ s.lww t~l;Il a!lJjl~ ~IgI1H'I Hum Ihe rs(' .~ lilJlt1!imml~:y

ul)iquitiul ligase IlmiL is. fLJql!lirod il'llhc ~.;ignmlilllgccl~ (,'Il.I~IOO~ <Mid :IDClijwui(m {H} [rs['
U!ml'll'S lllQi;ilIlc ror ~.ffi!,']1Il;!fl"

C~Ofll;'S OrIJ~Jjj ~lt'mIOO

(:xplL'SS.ocm (Fig. 2F~anmv) 01' else .Siing]c EO~ trig. :2F. anrow~'lcfl"H. !liC !C~poc~clt\::StI~, lOr it i!!lCnc !;CQ!I~d in U'lC TSC Wi~ .2:A), M:Qn,,"O~,,;:a:, cllfl D p~1l, ~Il:>lII!:h hm~u:.;, w&'> pJ1C(lmni~liUll!l)' nt3j8l:t;;';;'~1 'L:i} Ih..: p:lf~lIl ii'iClltilmlJiC Wi~ 2(1), OOui.'i4:¥.1J;.l1t wDlh 'hlric kmw.11I ui~,ml"."ff~ ,ror unl iii Dell:l clfKk:eyDi~ ~ V8,. i91, Ch::~1'lllYI'l:11~Q'lllg Sr1l.fl),. HLi!CSL Crilll;s,.. ducer of H...: c:lIl!~r~wl fo@L()11I Sig~L~Uy. W~J]jI1"'<LY V51<. iillso gCllL'I'JtOO ImlOP.; n'KJ6li11c for r~l!L1'() {1Fig.lll). Unlil:\.>C" clones {I:fll:rc ~iEllJa~irlgec:Iil g!i.,11(S" <!OOLII ~lrofwhicll h fII~x1W1X.'"~illgl'cK~ Sli(Hn:SC : dones, :IiIoc Nmdi- :lSC d:oal~'l; fm. ~1~CGIiIJJd only ~mllm'S, iUldttati];~ 'that 'd'l!.~ ~,~C::; .~[i(: n.:qillilfil ~~ tlrwEB ,(.Fig. 21~, Tl)IIlIlor oclb f~kijl!g lPlUi'\IM.m iCOI]lain~I high Ii:!~'(ll!<; oe11~ (fig. 2. J ;Ifld K~. of

brane then d()l;$ liS d~!!gI;IL\Cr (1l'Q;. 3. E find r-, TIne ~Iliking llh'O;FJl1Il1.lClny in DdlD. C(mtCl.l~ wren).. prcsurmbly as :J r'lliult of' lhi~ di""i"ion:':I[ IK\t\~'i1~l ~SCii: arne! Iheir 1Ilru-IT' dmAIr§ht~]}i m~gl.l~ or.i(;'I'HrLitlll" Divi;;Joll ~w.:J>' fmnll l.he ibmllCmmlt tiro 0If!9(l hy 1IS}IJlrUIIlt'lIIilc Dcha ~~n;lgalf:on" Cm~iSC!' nlL1Il."lr,lm~ 'IL'II.'l'~p]<Iills \,'.1:1] Ih(,IIILlrk\L~ ~:s('~WI~fJK~\irl!I!L'i.ly{)ng;lJ:\'~"<I[(Io I Ll:'! nlll«~ ' il:l... ll~y (~IJ]e1l.t~y, we ic!t'llli:fkd mitotic ~.SC:-; !II \'ariC!tliS l.irlanalb:!' celll[ Inn:;ki (J'!)l No com::bnioo IxIwoo~ i:'i![IIg~by s![1Ii ing with ,1\11 ill inand :IlmlylOO the n dis~rilmlMm oj" I)cila bCI\1i'CC1lIha d~tlghtDf cefls, spindii! ol~C!~latr[m and drnJlghlcr ac:~1 I'i"nc was :Sil1JH~1''~1Il0tlillIS of]]&1a VCl'i1i.ck,,,j, W(lre ~el,\'ed dii:'lGoy~d, hmvC\'CJ~ We n~xt co~~i(l(l['l'il wllt'thcr Nmd~ :sigrn~~ in both dHlllewly JOlJl1in.g [ltl11;llll~~B. I;.,\'CIII ~11 ~ul<'[fi im 1~lnplmt'ic ~.Fig. 3A}" Ilow~·'cr, illcdl p,lil'& dl,U~ reocp~m' ~nigh~.cliITcriu nh~~r a!Xi .~~~ fl~ ID m..."('(I.ll11!lee ooll... \'V~ £OIlec~~ I1rr.dlj Llo;tcmnplel.(l~1. ~lIliHl!Ri!l, as C\lidcucOO h>' !.he ; reuplak(' of Al]il~iilll~ dallg!IIlCr (,1C'II~udaj. [)liCh:a· !ll,nkQd ISC clm~~ oflwo m [Oi,ru:0:::11:.1Ird &1r!~l'C'd n lhiml for PrOSl1'CfI');:1Il:!! Dc_iL U'ii.g, 4l Tile lSC in ' fie!! v(!~ide." du;;;'!e~cd in Ihe ]~C no;;oar Ie' fi'l!11lt;'r d cleavage t'lIl1Jm'll.wh~Jea:;ouhf~Y \\rere clj!'i~1Lllt ~1)J],~n Ihc;,'C c101'L'C',:O; co'l~ld he dl,:~l'IIlIlil1.cd by il~ ba".1I[ ortlS IliS~ Ollie H) lIirt:e the l'Urrro'l.\I ililrnl weaker nn111c EB (fig, 3 fa, 11'K:sC' ~wl~kH'I;~'~I'J:;c., thoe ~;>;;p.;:riI1'tCnl::;:;.Ilow Ih'ml Lfi;hll a."yrnrl'l~iY d'o~s (~~ dfLllIgJ~l.T:> {iul rwo.i:G Rl!In"Ocli ~ktJo£, I~Lh n ~I~ , Ilri:9~ s:illflply by di Ikl\:tl~i~ll ,~I\(;g!~liull Oif D~Jt~t ~xUII~dbc dcdui(;-cej ml '~:I~~ilsi:; ~~ Pro.l'PCru rnJlo.:iri;t,jjlJj~r. [lciDII 1(liljiC1lfSiO I:'!!;;: rupidWy t1t'!o''Yll· e,;';PI(Ni;it;1UI. JSC,; 11i11~ luI jtJ:>t '~r:ucluoo~a p:ro-EC reglJ]hn~'tI ill '~heEB 3Jlcrlini1j)!ii~~, in""ri"iliIy ,C;Xplt.'l>..~ .Dertu til C"Wpl:ll:;r~~OC:~OC;I\;;:S, u~l1~llly rn: hligh ~\ els qF:ig .:lA, N ~ IOO~ hi !;IrikDUg To iU"'l~~ig,m(;" tlte l~tcCllliUljsll'l or Il'li~" tiPC1l1.11Cy. 'I!.'C !i1S~ COUiSDU,;,'!'Cda r6.1c .~r IlIii.tmic t.."OOlnlSl.I'SC:s (hoo had jLI5~ proUw:cd fJlo ee pu'I'"CLDSQIt COIm'urrui :Ii:\y or no <:y"lt;pl1LiU1C Ddlrl'olcslc.iCs (fig, srinti le onemfLliuul. Milm~c chromoooll'l\'::;,were 4. B 1(1 D" ~ r = JIJ), Thiswas mlC 1M ... ~~~l :l1IX] :SlflirKll wU11J pho:: .. :p:I'Lnh~rQulC HJ !tnd eemresomes wL'Lihy-luf;JilIlin {FiIg, 3, C ulld DJ wld cdl:;; W,,:R.: jlll~ ~W~('~llrlln-mll EC to \,'0,; prnill!lclion (Fit. 413) Ill:> l!un h1id jlJl.... OOllilJllt'fi:,,;J~L~('fUliidl Im~]y;lI.'tIIin 3D iI\,"IDmiructirn]~ to· dcl~TIllilr..:: DBIo": ,,,..:II m, ~i~ mi!:Jc- lQ) bcl.~~\)(!ill lIii! ~)irl{i!C mid Ihe bl!Nilrll.,',l~ OOII~'Culi",.::oJC q F:i!j, 4\ C mit! D),. Cfil~l!II'Lir~~ prumL,ndl!'alle. 'Ilrlt'~ exl'll(l'il!l1(lllhl l'Cv(...I~d ni!~l tS('s d!JIclion of c:o:~tly 'tI.\~.t'e dal!lgill1l:)1N <II)~ mll(l tlw.

which SL~~IU!,t~ICOO thatLl'C}'~' SlCl'ltl .cl~-m~c. c ,ti~,ro~pe.ro.~;>;ptcssil'l,g Ill!ltn(tl~ ce'lllis eorusined .... ery liUlc[klm ,(fig. 2. G antlL). lhlls .. mcs s~111!J1 theurpte"I~C dmjjJ,lucrC5 viII ~lie' Cf~;ron~Cfll[ NO:Lrcl~ 1}!Lth~"'1LY.

Ix.'l'W'teill 3° and 47" lillcwn - 19" ~ ~4°; N = :501. Usil~geyOC!lpll\srl'nh: nililllkcn'l!..... room] lIim the' tiC vc (.ng. 3E red) tYiliCl1l1y ooJiluim; fllfIl.udl g;r:'CaK'I"

sm1fLCearea of ooltlm;~ wtth the iJd,q:l.t:lCUI~11e!1~'

or

~:sc

ime

~sc

~:su;

d~vidC' n:rn1];TI:l'octrnJIIly. wiil'h W!htCl1 ,rnf ill' l::mging

mle,. as.

\\'C ~~'I"\iQ:1

llo:l COlUi!t~1IR1J~~

il!lil!~

IF~,g'. .2. A~ym metrit


~i'i!l.~lin9

DoeHa

flilluklllailly
Q;lint~~@f

liraili! ISC 1'Q ESis

re(J!l!ired r!Jr EC

p:rodm:tion a Ild mit:otic (!r:!~1 cyde I!'c)JjiL tA) &pecrtedl OUl'

lad:in,g (0,[ r1en};' N em:h 5l;gr1'<li. (NI'i~ 0""10 l!Jy<iln alr,m~'. 0511' IFm.lf~da,y IS( dOIile'S (greerl) ,m ~!rJ !,!~'r[u~ral~ iI llJlliflQr ~9) mosaic: (orr Pl'O'5pe~O' (r;ed~.(C) a l11lllntlr pl1!l5 a ~f19l'€ E.'C, ~r ~D') S1. ·f1.{I,l.f [C.~.~E~.IIf:OO.i3i_Y... ()1011:e r-~.'~ . (g~efN1~ tlile 5efNJrl{.u;rJg nl!!~1 Qr

ISC d~!iI~li

~gr~~iIl')

allei.!:l :llhowin91 flO;rma~.

[15«

(a,r'FQwlw;:rd) ~ 1II0[ :EC prrl)dl)l~" tiol'i (afifllJl'\l'S)', Aifmailil'alkl, (red) hlghiliglhts c:elllll:ltill!md¥i~s" fJ

Sei!!'e~-da:f IS/c ,r:[(nne5 (green) of

n~"u fQlrm ~lImor~(~m;liI,li') mmaic 'fiJ:r Prfi5pe;rD f);iPJMSiDi!I wed, rmdemt o:r s;ing~e rI'!~~~@
[0Cs (ai'll'O'i';'~), Mra (I'e([\ (~I&
ffifirril

plasmid" ~G) The red (hal1lli1~ lA; ce,~$ la.oolUjl IP'rOOJPElI0


l[),e~ [15<C

s~mt
(~-)

oont3111'1~;£OOldl-fl'lembrali1e

tal.

Q>I!~

d.llIilE!' (g:rE!\~fl.~Qf tn~ $'1J(;"l~l 1l1:!~1llele fo:rmsi3i hllmor mo· a :sai;'c.for Il'r'OSIl!HrO (Ill ul~~e.a . Ired~" (II) Expected IOl.Il<:omes of [15,(dones (~r:een) ~ lackfllg 5!i(W (,orr!II}., Ilmrpa'ired N,gl,(h;igllli31l jecefJ~i,ooilrn tl1l i5 d en CIted! [oY03I blo.r:kJ?<I.mow. (J~NCfldr done {~fee!l~ 1:101 ~r;ns.sinrg rrOs,llef(l (m.!d~<'!r,

lack

fi(pre~siing Prol!:pe~o
("~

c:1.of!e!O

Deltal.

Twe~ve·day

red) ·e~prre'sSiesD@~t<l {cytopl<lsmiil:. red) like ~S(,s,.(Kj D,el!.] {rr,edl a f'lIdI DNIA. (blll!l1~ hem UJ Notch- (ton,i'!·(gween ~eJ:plie:'i:sirngP'r'Ds.~em,(f1IUi(ieo.i3lr. red) ~old::'jDette! {~~o[lil.a:Sliflk red).

m,

990

16 FEBRUARY

2007

VOL 315

SClIIENIOE

\o'V'IM;V.sdencemag.orgl

REFORTS
C~ln'OK'" [[Qi;\,\::\JC., rSCS Ihdl had >leccrlilb! rom· plelOO an e.;pfur and 'I("'1(.11 j~i\.'l [c'IUfUOO ro ,IE,C IJro. d!JC'l~i COIiauairtcd mllllrplcDJdrn "U>icics tf~ 4E~

These

ri.'Sul~

S!ll~.,:o •.,. II

simrllC' n.'KMld

[QI"

111(,'

d.l!tCllflifituit:"m of ISC dmjglm,'i cell [ate" ll:lug1lile~ of Isn witl,1 high IJIclm ]i:\:c:ls cwouM I\.'Cdve .iii.

Slrorl!,! NOk:~l si~n;lllmd bccOl~ ECs.IJflUlgjlIL'I$ of r Ise!;; w~jl1 few or no dcooct:liJ.l!1c CylQlllilSlul:: Ddru

vesicles VVOLild receive 3 l'illic.l~ weaker Notch andbecornc ec eells, Ii." N{1~d~s:1g11~~hl~

:si.!;llil~ v,'CIC

Fig. 3. [le.~ba, vesltc1es, are dhtrihmtetl a'~rIlmelJricallv morrtty after mioo~is. (A~ (,qual dJi5~rillJiJjt1finor nJleliti31 1.i'5iclles.bed~ dl!lir!~ IliSe ill naphOlSC. (yto~i!1l,e:>1Sr'urFGwissti'O\\lll b-~,Arrillin stain (gF-een).. 'IB~ [)cnl,ln-rleg!.d<l'~~n or [)el~a hetJ) 'ir:l
'Ule- Irte~!dyh]:rme-!l!I IE!l,f!JlJ!l!I tralrusi~lIi~

as,fl'llIlll@lrk POS;ltiOlllil1l!l!Oi~ vesiclE'S wiitt1illl th~' IS" ~iliJr,ow};AniUlll1 (g reen ~ :siholHS m,lI:lei. ~C:lln d D) Nli,to~i( 15[5 'inl 3() r,i!:~OIi'i~ttrurli:olli, W'ho(sp,110hi stlJlne "3 (~re,en) and y·wbl!~in (red), Sl13lin·
ing ddiiru~' ~h@ diillisii(;1!11 (!IFI~ t~

,(~)

re~trl{e to ttu~ bas:ern~li"It l"IIlembrane (b~u~ilnd red! ~~. TI1~ nS( ~1l1 l'lii~ m) a l!!lo s;!ilinecl JOT a .:tubu lirl fred)', (E ;ami

F) An IS(~E:B p,ai rsta~!l~d Ill!" [lelt.'! (redl.l and: Notch a(tirlora ltOrl fe<pcmer (g~~~)' r~l'l~l:>U1~~ ISu IhJ~'I,I'~pl'll:l~~r g "®l'IItad '~da~'i'leiJlline) 'lli.1til ~i'le !basermml ~~mbr ane (bm),

di$n.~p!~t cmircly. as ill NOlch r;rdlti:'.w{ly :1mill mlll.!l1.t clones al'lDr nile o.r two divi"!DliS. u111,Y~'SC-likc dmglnC'J'iol WOLIII{] 00 produced. Noil~~r',~ ~l'Cll'so~ 00]' :~SC'·likccd~~ WfIUk!1 ct':l'iC divi,..;,lrm'~\'hl'n NOlch ~i;gJIl!l~l:il]gi impilIimd 'liVe lr-iilod ,hie<; nork~ s r by ~~lrcss!n,g .un! ..K1hra1l..>d, Notch re.oq;tou (NOClj in MARCM l:SC ckmes Wig. 4J), ~'!:.e-.lJ:.pre$Sing o;ll!'i. which ,.Jm!ll~d OOIll~,~~ ;t higl.1I~,,'d of Nrnth ~gmlililg.mH bo,.'Call1~ SC:s. (fig. 41''). II] ad:!ilicm. mile on~~ cbn~ ~l~'" 'InoVC l'MQ ~~I!i, in:;li:li;:, whkl! !:hm~d1<l1 Notth .. c~i'li,"'ti-n!il,.'U(ll~ll hc-IS.('~:.1 \ a ~Jr!ICklin to rm~i)'Its di.tif'IJ101tiallioH W <In EI!.~. '1l!1\,.'$i2' i;l]"llCl'jmC!1t~ [II'Ovidc .~!]1';.ig,htinto ~C'\:C'I:'/I.~ IDn~~!'lIldiJlg is'~c:;;, in stem celli bLo~O"J}'. r. irsi. L'hcy iiil:isli:mc ~1m highly speciflc stem ceilll IIUiU'kCC; (:1111 .be- oo1J1i~I~...:J,m ~~I rlJ;r !llPllilliClI;i~:nr :;!i",'i'UI ",-..:il tmdcr r<jiti)!.;ul~r ~1J.mli~it:m V''Ii.;:lJrr~c •" . ISC~ rulil~OI ci.ut..:rr[ly ·be i(kIHiii~tlwilh ~x-mlbtty LIiC'~ic""ct1 in (111)]., DUn N(I,1.e~'1i~m'll ijig i~~i~nli lil~liy s ilfIVOh.'cd in SCCI1CRIry'oeil versus EC f1tOOu.\'llion. Co:rl;Scq~tCrnly,lra;li@,3tL be possible loilkmify i "'Cril.'Chr.il.~e .ISI(,~,i'I;;, ;J :>'l(bpoPllll~~ or odl::; inn Clllru~h{\d ilfl NO-lcl~ liliglIrldj; wi:th[rl~wom; Ihc 'II'Cr[o{lhrm!;;,g~l."mt ilrC known to ~,JIIlaill ISO.

or

~lm'l' Sil.:ll,l cdlllc~ii\lity w irh lis$ll"; it"Clli ilK.'!I'1k'IJll'o: ~f'i!;!, SIA), lOOLI rC\~IL'Ill!.d: .~i~n:lll!'l toomi)'Njg!in~ cc cd!:;: (111(1mhereells Inay lllt1lK~nct'! :In ]SCs N!:l~,C:h signal il1.~. c;;]p<!ci!,)' and BlC11CCtfle~YDlC fir cell II

s,.;"CUlUI. oLIr!t.:,';lIlb;. irfw)' be coorrIillllll1.id

~U~I

lm:lil;lllocdln~h~l ,i~(l ~.§'lkm, JJl~l;;!lpJ1i!{j n~u~nbl~lJl-o;; (IIrld !'ICU:'Inry org<Ia~ '~Ci[li'&llI'S ~' 1l1'L~l1!I'I1I!?(~ by llfil~ ~~~][llC!rk:
!I;I;.",£re~'3tlljJll ~[mnlroldb~ d1a1 ij:.i"''i:~h,c ~lltlS'CqltCIll

fiif Noich :'Ivg'nm'l1 ~'\."\!iCl>vcdh~III )]. DHf(,'!l))mi<cl Notch si.3!!~lil~g !lC{llujre~ dl~ ,L"Y~1!lJ!1Cuic Cl!!.(~lic 1r.lfHcking of Dc!!'! <lila Nolcl~ a Ocr ~C1.I!'llO:l' cell l~\ i~i:lfl [n:viC\voo ill {is}]. I~,ydirl~ em:JfI.SO!m'"$..am C\lser!~a~ OOfllpoll'!!U (l~.' Nu~d~ i>igJH~ill!:!,I!ot:i~ia: ~OI)(".lCl:iil~muie L'l:,;g.lLlIliS tutti have 1J,(~111 ~I](Hlcd 10 ;s~'&4It~ dnLif;""I\::lllji1~lyill R Dn;x;1}j»rircl ~~~OII)" t1lwm Ill'ocUffi(J( mvlli4ion."l {n). 'Oltir !>Irnl,ii:.i~tI~'C.'>'I, thIn ""dOC,),lic mllfk:ki]l~ (~j' DdD:l, Il'fly :(150 be iuwh;(rl Erlls.c fUlllCLLurt \l\'c tib&:wt:d 11!Il1~ Ddllil.-n:i:cill!tS:id.::s !L<...'UL11!l!llI)urc nOliI-

tmn~i.",;;ku

IlIndOlII~I)'

in

.he rsc 5IJO]1~y;1il1l~rilIuttJ...s ni:.v'lI:tC i

Fil)J• .I.l,.~A to, E) D)e·~la exprl1s''£ioI11l,ew:1s in 115(~ dellEl rmi,n:e Elf: vel"Sl1IS ee!'a~e'. [xa iffIj)tes o~ lWei-tn, kmr·cell liSe cllo'ne~ ~gre@f!) stili ned fo r Ii'r,o~peN) (1IlIUde'<!f. ~!l'dl) illl'ld [lelt~1 'cy!ioRl~.asmk, r.ed) are :Sih®Wfli. AlnrO\\ll'team denote IS(:5~afiFO\\iS ~elilOle Ilro~pe(lfil!'e En. (A) IISes IllrmhJ(ijiilg a I1ms:poe(ti~e E:C (IPfot;l$~rQ o1!lJ.:ienU ,il lW>ilYS!l'~~,r,~£s,~~~la< ~iE!) i:SCs thOlt, 11~'I,I'~just: :MI~ll;h>edlkl ee 1I;e'lI,{prosp!l'u) pr~$'~nU ~nitlll,Jldi'(ln ~~:F@~S. ~itU~ no O~It(i. (( and m IS(~ ~r()duti"91a sewnd ~o:nS~(!lti\l\~' ~~ 'OI'r ul~l 'e"xpnm liWe o:r ne [lellar. Inset:: Clol'I!O!J1mar~er '(or (C), Uj Ii5Cs tha~ haye jusl swutl:tled rlr~m ~e
t(D leC prod'uctiolil eKp[~~
l!.'CiI'l

De-Ita .. (F) ISC ('lof'l~ ~glf\eeolil} eo'}!lPlre~~~ .flrt ~efler.ateI1g


ell le

Iile

OJ 'l\'l®

H::l<,

ind'iCil~j Il~ thaJls

~SCs, .are !'lot m.ai!'ltC!Jilled.S

bar .• 20

~lml.

di\'t~iw~ p.il~lll.c, Such UPWCI,";S;.' mi~l1 n.::::;Uil, lium 'nl:l\.~ jllh:;rilmlc~ of !~Il ""ildt1Cy[i~ ~1.IIlx:Qll,1p;.utnlcm by the ISC bl!la not its dtlulg;l.tLlf.:1r. l~uLi]l;!l[1) dit: kl'tllK:~ irn .Dc~I~"'~llbility Ix'L\'\,'~n tht: h'o'o eclls, F~m~~y<. (!.Ii~';~n(.n~~III~';SI tha1 Dmw)p!.I01lI ~ ria lSCs p~:Jy .:JI.IU:Ii;l diill-'rt,u, mlll amu of5lt'nl O.1'1:Il! ill pn::\'io'!,I~L'" d11l1",c~l!ri...:m nklli::s', hI !he M'a!lPn ~1n11iI!C S;~C1n ceJl lOSO nidK" (i};lhe OSC ;xW1.Q'CS 1,{1 at~oo:m :\!IKllnJiJce~tsm ;II~,i'!ml'!OJ1.tical·~Y It\ed ~~P!l t1!ld ~~lds to k~1 ~igr!lt'!~d):[11. bleck diiic!l:'Ilti<1tio:n Illg. S 1B~ OSCchu,ght\!~ (,lHl~rallk~c~Jl~~ ~hc\y arc RHUI.';(1oo'tiide Illis rone of itlhihiil'iOJt In OOhl~!<lSI. ihe rsr ~)~y$ ,ll~ OCiIi~c role- in sigl~lhD,g Notch 1.0 induce C'lihcr Et (I, ,ce di fiCrcmimiol'l. ~I rC'lliIru!1lS lO ix' d'e[Ct'mil1o:l w~'mhe.r. cXlCnml s~gnals m'l.'l ~hil) iilill'U-

"'r~

991

REPORTS
'[1111,~u ltliiim1l111 bl~ ISCs, ,in !Ii i1Iimmcr ~1I1~llo~'QU."IO 05("s. ~J] 3IhJ' cr~. sI.i:Jll cells ~lC.f:I as ~'SO"~,hich
7. ~ l'iig,tr, D. l.
s;rltiIC~'

sense iGC<l1 C!!IILllar


i!

req~Mrn.';l1i1Cnl~lii'l.(l II'I~rrnlSC.

produce

a
'9.

.tlPf.tR1flrilue dil.~lgl~c:r celliti

~ik~y

rl;~~)i

celllil;;ii'li I\_~"; in III,.; plll%iok'Jgy" klJ~,-~ifY. fU~d pallm10gy oftti!t: ti~ucs~ilC}' Ililmirrbatirt

Re~e~en(!e$ ~n(!l NIHes


1. '1'. H)i~ttit C, Spr~l!n9, Cim, ~Il!, Gl;>fi.t'r; Ut<i<: :iiJ6" 46] WID~_ 2,. 1)_ T $!:admen, N'oI~ .U'l.,~m!;i '(mo~" ~. T_ );ie. A. r, S~l'i!>!llill!lJ. (!!'II ~l1, 251 U,9WI)4, T_ )J~. oil (._ S~radlill!lj1, 5fjeM~ 2~O,. ~<l(l ~2oo~l_ ~, x_ :5oo§. 1 XLe,!J'o!V~r .. no, ~:t~9 (200;1). 6. r.I, illllijBil" ~ Ma~~Ris,. $(jBrl~ ,2,.., 2~~1'i, '(.201101)"

s:

W, it, i2.. 13:. Se~ ~uppoolil!'ttl rr!al~laI en SC!feM~ Otili!lle. lA. N, Mar-f,!~~1 ~ll!r" ~. CWI9, Hi (2'OOS'), l~. $, 1!r{il1/, ~l. ~, MO.t t"ffl aNl;l~ f. ,ISle 0(2:0061
11)." M.. ~urriat\ S. Bray. (iJt:r:, I!IWl 11,00 ~20!ll1l"

JrllJel. c, S~ItU:II. M., II, RDgiti:!, M. T. f'ul'i~,r, 2§42 !200n S, II,. (ri1Q~~tlm, A L1:CIi'~~d. n T, B.~d.]. IKim~il?, r" NJo1. tlJ'tJ1. (til 17, ~iD51 (:ilOO6t L.III, (a1101i (it 11l.. M:l!IlIIi'i! /l,n. ~n.{200':31. J, 2J1'~!I "J ol't, ~,~ 4,z~, 830 (~OOM. @. O~lmin. {). (, S~mdLf~g, m:m.rr.ti ,~~~. 4·70'(200!J,)'. (', M!~IMiLl1, ~. Pe!'1'!m!l~1NiJlI.j~4l9, 4H UOO~.
~'II,

,20, ,P''1. ,BjIll1kilH!~, (Mrng, Am. J. ¥o" 2:B'1, i(].]Bl H. ~lliiII51, illi. ~,Ro~i~, y, N. )<lrl, (00" Op,in.. GtIJ llkIL 11.. ['15 ~lOO4), zz, A. 'GlQm~ ,~ ~,,' G:,-J~l:il'3., 7S 11.·OO5;~" ,:9. 'We!l!~r'!k ~ ®.l5ola~ ~" T. ~!f;!'uYI ~'(I<f c~rne~~ >00 tI1,~!f!1.lfl~it.Ii.[!!I~ool H. !W~o~-8il!o)e~, ~, !!'i'1f1e, C. ~'ie{d, s:. (jiN~r,.n, ,andl !. JleliiJ.)!' fIDJ gift~of !W!~lI<liia~. S~PIPortil1gl Online IM<lteri<rl _.~~~g.CliSif(giil:!oot~l'It!fu~l:ID!i'I·t.4fWoiUl £'d1t11i~,11 ~nilM~'il'i!Mll! IFi~.51:. '
'il:E<r~rief'j~e:i

17_ f: He-ilter, f: Simpsa.rl, Cf!IJ ~l'I,l!Da;~ (1:~911 ml!, [, 1P'a"tk:Jpa~iDS et iirL, ~v, (ell :I" ~7 t!QOil), 19_ It ~e 1~:lITgne'. Remaw[j, :5, Hamel, f_ ~RWI!isglltll, fI{,o1; 1)'i£!L :I, !I!~6 (2(111110)\

s;

~3 'Ooidber 2006, W;I!pj2d :~li!Ilua


J{l.1:l~~ef1~.ll~ 6006

I}'

~OF,!

IP'D,lvm,elrizingl Actin Fibers P,o':siition 111.nte'gr'iins Primed to IP'ro~be fo,r Adlh'B':siilon Sites
Jlltigw.nin'g (j~'ltsenend IP,:roUiUIs,101l1s,. pr()bing th~ Sl!r~@ l!ndij!l g mato)! in :sealn:h of Ilerm~ss;i~esiles to kln'!i1 adhes.kllils. We rnlildllh<l't a(~rrI fibers P<lt~lif!erizifl~ <!long!he Ileadlin!! ed'!1ie dijre<ted liJ'Call ~ro'trusij~:ms and dir'C1w sY!'lCihro!lo~ss;ide!,\!<I~s mOilferment. of fll imeglrin ad he:sl0:n1nl(ell~Qr~. f1hese I1IlO'\i\emel'IlS: lead ~o the dt! s~erirlgl <l! rid' j)~nm!l)nijIIlg Q[ "cl!rI~@rfllloi!tiQI'l<lillf adi"l'<I1ted" blMi: II.llnl,ig:~ted, r1", ijllJteglrins aW:rl~ ~~e ~eilldirl!l ed',gle 'Ilf fibroblast larlH!Ua e ami !l ~O'w:th 'CQne fil,Qpood~ol"Ihus, Ira [lid actin-based !l1i(i\le'ment o,~pnimed ~Dltegrim,akiollig ~'he le.ading oo!lle Sll!g,geSl~ oil "!>tk~ ~url!!lers" mechanism to pr-o,be ~or ne~1 adhesion s:ites ndll(!! dired miil1!roltiolfl.

UIC .~>q,~OiIl.ey or rip~k,," cilCCfl.... s;cd i~1 m":rN)I1SC m


tlitc ac~h~=dC'pdJymr!:~zing a~enl cytod!.'bIaSlll D (Fig_ ~f' l" lihe <iIS_'ille;i<lIQ[J aCI.i11 dl}IIlITI1~ icsllI.re dir.. !letlil l~ 'V i$Llfliizc w i~h g~il I~LJ(~rcs~n'L ll'IUl;eiJ~ o[G.i'P')~flCI ina 'bC<!flLl.i.C of fh,.; den $i~y i! nid mlli(] turncvcr 0(91, We· 'Ul!;J..:!bb"!: ~"~~)~(}it..:d i.heiJrt1fH:rtk:~~o:f ~lho.1Ol.lC'~:iv!llflbk: GFP W;[I(jfP I (U~ lO c$!2Ihii:ihDhJ;! ulIIl1L';i!,imld (If "'isi.bk IIC1tln ~IDid to li'Ii:"slri~~ ",islmllilill ion to ~ ,Hl% (Ii~ the CXPIfC"Sf;I,,'rl PlliGFP'..aC'tlin U ;;). Liru'Iitiil<l;! 3C'Li'l~u~o.l1i tcv(;llkd kiICfllhoo ; ntcl\;!lISlCS, .1 RCliil [1l1(;llsi!y ItlrU nlu')~j(:(] UI sirlii,"way:> wid; 111eripp'ics (fiig, m,t'lm1i Irmwic S I), 111":: ill(;T(;'ili>~" di,ll Il.O'l il"'ll)P,;i>~'UI, lhi~km:;:.;::; ""ariil~i.o.ll~ or n1crnh:rml~ Ii n lng, hccmll'l\l Ille)' PCI:>~::-;,,"'t~llk...r nennalizarieu I byd1t: c(lLnui5 fCcI~"t!
!1Ilflllnllu::riC

rod

ULlNC.«l~t1pmh:ill {u~R.Wl $ig"

iCCn ula,lr.mtri,

,he [~1)11 s!l~P!l nor 1l1.1ImJl,di;l1'l oc-1iIrnigmtinu;;;l!~ ~1'>~Il~jfl~ onhe 1c0000~l,j,ngo,gl:' r


;I:n(~ "U"~l1'1l~~! to·

p!ll.y!ncrii",,11iO!1.

U),'E1!,;!ell~i!:lll .jg d~hiCl!by aCltin

~hC' ~\[~! I~d~!l~~,~ITI-

ilh .he ,I~ilij{ln (,;1).• sp..~d" af,d pe ~ij,..ieuee (4') fir th~ I'cil!.di~!~dge ~.~W!1l~d o hy >L\;lio-~h'l(lil1g, flm'C'~!)s. 1~L1\~. (;"Xl!.ensUrltl h~ Ulfit 1Il1.ifomn; ,u;:ljaocflt rcgi mlS, of ~lliila~' 1)]Th1d"

en ~\

!l,iWl1~ s~d (0. W '(HlfllJ,lll/!'l, n - 34) lh~. {:!in n network 1!I'C~dmmhl!l ,(CU1'lN ... 0_013 Jllll/s. l' ;ill (9, ~n(lvl;.x1 ~~~kl~")ik~ m'l~' !il\:)p~~1 Ill'C~ I.r !9!ii l'(F~f§. Ii) ~l~d1l1W\,jli;' SII U.5), C:.J~!'!; 100'C').':JHy (fig. IDq.M, III. hls;lcad, !he R;~lriC~~llg l~liero.giotlnf 1~hnln~,cci\i',~~i~11f1er !'iPiilcil !V!l11\'Od a! the :{.;'b~nc ~X"ICd Ilw:1 n(:li~l w<lled !l!!ol' aclha~cd Pili.GFP-<t-c! itl 1L'1!C'",~i:,,"NI~l I~C)~Y!ll.t'I'i:i!.'ll~jm! elo.ll~<rt~ gmwd,. Cilor!C' ~~~olmd.i", UlC'I!i'~01',1 !!M'lcell~ ripple <!'Ild aceumulawd <I1(l!,!:~ t 12),and mC'ke!~ LM~'''f{l ~lJC"))!'i<l! ~13), A l:thrrughl tlllC G'klngmif1,g ripp~e t F~i.~.:2 M. In cnn1m~t!O .hi:,:

nail, ,u1(l(hco "l:!Pies !XlUlo1i!K.1d\\f~dl i~l ngicl)IC a.S;i olMi.cal ~ctifl:n Ujl'- T~IC' illlcn:;;hy il1J:::~~ moe Ioc<l~ <tel Ull 1'10]ymll,crri;miionnt C~~$ of rill'p!l{)s umvern>in.g ~1G' lieadi~lil~t~ {g~% on!~.liw~ in111.

IP~g•. ill. local im:lrl'ase,s,]n <l!!:tin ~(!!~Ymer· i;~tiQ!iiI OC(llU at51'11lall rip,pioe5 ll'avers1F1;91 the le\!.rJ~iiii~edglE! ®f mi9railO ryiibro:b1asts,. (A) OI(i3JlJd P~,(jF1P=.1I~tiln: ]~gi~$ $It~l,!i n.91 ,[.'1.6,,, Qh:s..:rvaliol'h.';, ma prulrusi,tlll i:lj'Il~Llnie~ fci1 INIH31i3Iiocob'las;t !arnett.:! <0 <lnd ~",rme,m· '[0 ~.lh': hypt1'UI~'Si1>\ Ih:1l UK".>": [pToUru;i.tJlb \I:Jri~t~OIl:,~ .lire ILIl'i(-d too se~t[(;h IO[f!'Cnlllissi ..,..: :sile::; lo Ihlim Il'oilliumr Up~.Rleg~iiI o~ .rei[ '15 ShoUIli ii'll iiii5Al't~ atld bl:aJ~~,bo.:< il1:!1iica:!.e',5 re~i®iii 11'1 nI~W mlhc:sim:r;;. 0( 'I, m. Ilow(;v(~f. ClO !:ul.Own ,dirc.cl r,..:1.1'lItiouiA"1w~"Cnl pmll:u,')I{J1]. d)'lUlmi~:;; rulli '~:Ir (O,~, SGii~~ b~r; 5 }un.(I~ Hisl'09r~1IIi Ilf 'tihl~ ripf.Ple d !lIratieiii orr wave ~8'n~dl '(n ..1;06. <lbil ity or <Ie ~1!..;:;i~J'll 1'l.:'C'"'P1tW::l Ao p~k I.I'I!..~ nm~ri", lIceUs) U.5}. .Kymo!1!raphs fr,(!!m DK Ilmlio m~n idl;."llijiH~'-d, iima,glE!'S 51101'1 tlir,e' d1Llra~oinl ,orl.hle ~raIil5" \,V,,;;81!l.u:licdIltoLlllIsio!l dyl.mnHi;r,~ duriil1g '~.1r '!,II~Sll' riPl~~~ (while bro1l\Ikii!tl ;:Il1iIi lo(a:l in i,lj:1i nlHl:ris: ii'xploJ>lIl.iDl1. lJ~illg lrIi!:h-c:lOil:ilmlll Ie Maim p~etrt!l1>i®iiiS(hll:a~'k br<lckEl'i}. '~or" dillli:::oofll.ial im~1'Rr.cnt."t: mmrr3~1. (0'10 t.1f!!~ic~. ilro!'!ta~ sea Ie bar, 20 s; "l'ertit1! ~ 0.5 ~Im. we o,b,f,,"i;1 ... ·00 Mmdl ript>k ~J.[o[ru:sj.on:ol lrd\.'C(xing (C) C)llli1(Hla!~as~rIII) (I)) decrea5es. 'I1hie '~hc mUcmwsl edge {If Hb.L,,)1l11lli~.hbUlld Ijlpm:li!~. ~~!l:q~ll'm:f t~,~~~IP'~R~" At t~~ ti~~ ,of '!DJ. 'Q~ ~hi: le,l(I:l!lgl 10 ~ !JIm oi'ihc ~;;-!l~lc~I(I ~,rig.L A rippl!t.ll. ~h.e,rl!' alN!' locaiilloo '~ncr>ea5es if! .find U). nco lirrl\$ (~id~WII,!1'ltw<:. sidew~.ys; at the P~6IFIP"i:!ctiIFi f~uo,r,e'5c:ellltirrw?lils;ilf (red alrrowhMd'5). 1iin~ ripples rnove sMJeways INaliooil1, ~l'Is!ihJre~I IDenlal a:JlC (fiJ!licl!'l,i.;!1 R~5~ITiIl, lals,t'~u t~~!1! ei,th·er·lt!~ M~f·..dmo"'~!iII~f!l~f ; to!~~iooJjJI. 11IlS1i'l1J!l:e5 1iiI~'lh. l!elll!lSi:b. M'D ,20<392, IJI~ 'of ~il{lp.od~ hr.~rti~li :~~llo.\\iarl'l;li~~"'ds.l ,:;n :l~aIioOiII. Ini'nule 01 Mealr.til'DgiGtl E)~oroM> ilnd Slrrn:~, the rea n."i'alrd mO'l.!'eFllli!,l'It ef Oi;;:tin' (hmironMalool'llal.lni!liMes illl H~mli, Boe~ht!':!da.. Mil 20a92. UIlII., tal Y'elJjJ~1tanoWlteal1ls'l diu)!:! ~o lamellal ... :c W11C!~ l'Oi!~e~l;lrud~rnoeh~d ~ 1J,e, add!lil5'lli:d. E'mili~ I'll't~¥!llr!k tr~'adr!i!illililg •. lOd'Shlld lines l'I'!oiIrlkiniti1l!l. p,oo;ililofk'S of yellow (lH@~1II'!e;:Ws., Sca~e' !Jar, S; ~Uin, l!:_"<1!1'!ad<l@,dir,micl¢r"nij~"gov

11<11prol:i\l;~~m; slilch 1IJS !lh:robJ,t'lSll ~a'llC'lbeand neuronal ,growill cones advance aad !'t!!,(l.CI ill'd..:"f1!t;!!KIt;:!l~!Y. <tfldwhkh ,~gil(m .!r1QVi;'$fO!"l""aiXI h. 'i,xHui,lfiim'lI,ly ro~i!filloo by Imnsv..:.Dk nll.t'.!~!cnficlil

992

16 FEBRUARY

2007

VOL 315

SClIIENIOE

\IV'IoMIV.sdencemag.orgl

REFORTS
;[lsyrnn:lc:uic: ~L~!;Uml~fLl~(m of f.aO fBl-flCl in. P~IO'[oo:c~i\'mim IJ r ,PI.lUFP alone IJrodtlcoo ~ sy,lIlCilti~

Illc~,ial,e,I,OUgill toll nklug Ih~00.:.,...-: Oif ~L(.h'3iiiC-

ilug pro1l1!S~OIlS,

iiUClIirICsignal Ihf~t pn."8d t'hrotlgil:Out the cell aud s W·1l!.'> OIfI.~~slem WLth simpie (Ii mOlilS i;Qil ~lIg,S I). O We ""ioo fOllllid that the 'I"~orodi~al(l<["stimLllfLled pl1Al~pla0l':rdtdl1 (VA.S,P I.. \'iIhi~h 11rott:ciI:~iJ!1!rood ends rrnm C3ppl'llg (·i ~ lilJlCa~iZlld~o nClin elongm,ing aemss lh~ ,~L'ading tldg.c (FIl} 2.B and I1IU1VOCl S2), T1m ckul!l;I~ingVi\SI).tjppt.;'rl ,tel,in ,ms],lrtlxldi and fL"iliooClOO~~I(, larger fCg~mh~ of ro.~\,[I:rd pmnu."joil (mmfuc S2l. Thus, it Si;C!illiil I,nin the 'tl'mL~V(,Trnl; r~,lpks rq'l]'C'~all!l ,~0Cl1l lros ofprc~er~ s

To ICSI W the loo@J,i~ increases in fl{;lir.1 Iw~yn;,edzaliion arc inV\liv~"'.dh~ nmrii~ I)robh'l.!::. we om'nrm~d Ui,Cilfl wi~h~hi: !:Oi;,,'llti:on(lfint,,;,gl"ill. mlllt'Jl:lol1. roccptOI};" lI:sin!!l a 11I11nl;cs~cn~+ nnnIlO:r1.mbing ~ntilbi:'id)'.~wob[ol~2n.'edl e b";'lC'rin!£!, Qif Ili In~t'gl'in8 lilal WlIS tclnp0flII.~y and . ;;p,:nilily ~'ynchmn~:zoo \'\I,hh ;JDtin po!Yl.1uJrizaJ:i.fm :lit the Cf!."i'il:il or rillpl~ l(I"'l!J,. 0 to F. and movj,c sn 2, lila p~ill~)i.;,'-g,ll'UIlS m:1in liCnmi m:4 rniocOlI lzed and dlJlrJng~aI. -3-mil1, nlOUil(lf.i,U,g pcdod, t!:;om;:~;UKm

.f. £l. 12, l,r = 211, IIYe cd~), ln iu:tcgrllus dii:d .i'lOl co.~oci"lljzc, \\d1il'l actin bch iml [he 1e;:q.ailllJ!: edge {oorrc imi()~'1 cocili,c Icnn U.Oi ::: ~6. I,r =~40. live eei,is} 0, {F~. ],. (j i~rll~H.aad movie S3], {l5~ Similflr co.~lal.~o.liIii \\'(!ft' oo~aim::c1 ~or positive and loog."lti\·~ comrels, w.l1l!!X:I.iVoCI,y Uig" 503)"Tbus, ~:I' ~11t.qgrblS nt[l'!"~ ,SYIflr.ibrmm,usJy wi~h BXl!ymiOr~

coC'f11denl0.50
~:II

COlll[7.ilSI,

izij n,g ;tcli~1 !.Ilbng tim kad Ing ,(!(lg'(1.

To 1.'Cs~.• vhetber IhCl\c ~limcgL'i n:iO \vcre capa'bk:

orb.llld.ing: exmL'r:;e~I'Ular llJalri:ll.~~ ~OInlincd Ilile;:ir ronn-U'nm~iOJ11Jil smtc" and nll)lnd~ti;\'a[cd, ~j in~irl;~ cfilQ!;,-a!ized to 111m I if}." of aclirll allollg 11m Icad~r!g ~Ig,c loow~l!al inn QocfficK:m OJ~3*" (L 10. IJ - 14~lng- :;,i\ :I~~dIB, Jij~_. S.2.to S5), To vi:'U;;II" i2C <LClivlJltifm andto d~11I1~inc ~\'flcri;l I!Ile' IF,U~:-;i" [ion !o a l1,i,gh-afi1n:il}' ~'L1[C'{liCi;l!In't'd, \~N~de ..'t"loa~~ a lii\,~ren .;!s:\'oc"l}' using :II "LI{I!I:e.. ~nl mn1jbo{~}' .c ~Lil'lg h]gli~illIil~l:t)i ~I imc-gri:1'1S(91::fi7)1,"II¥C in"lj~iall)'labckil all oflllC!Lcii,v~[oCd [jl til~l'Cgrirt> on dill: 'c~11:::ndiu.x,: :;'0 cimt an)" .o;ub:s~'(;],H"li, h~llillg w~1liiklllt"'o'cal iK"\,yly ;!C[iYIJ.b.. 'iI J, inL'C~ril>, These lilJ..'Wiylflrn:;I.:d igtlJ!!~rim HP~~OO o:ril~y m Ihe k-ilc1iug, cage: (F:ig;. 3C 3ildll10''o'11i: :54),

r~

The liC'ti'wned ~ I umcgrj us


l1.brotl~C'lill

I.\'CP,;

IWI hg,I~~"tI ID

bu~ ooultllbi rill fi~toooot~1l li1Il!lt~1!11L$ (fN l20k,[)). ~l'CCi ~iClII ny. Willl Ill.\; o:s~ I ~m.cgr.i]~ ~lCl;'::R)diiuwr Wig, 3D uud fig, S::n. which wli::> Ct.lIl:.i I~,tclllt il!h ;m j !IC("Clq:S~'tI~.\' it! ~l)' Ibit fihw· w OOt:l~11 lit lite I.~·;rdin~ ~-d&C (J'6.. 17). AllliHm~'1 unlig:Jltoo .~I.~gJ!""f1i.l]il.}'i'~\,~J im~g.dn:.1. whk:h can hi!ld '''11'r1l1n(lctiV! HbfmmC'lVIl, !.~.(lIa1sit"lld~'lOCal. 1.1:(: iz.c 10 Illf: ,~tg~ of spre<JidiDI~ cells ( M), we- di(~ nnt lind ib intcg~iill l'OC!IIf"'...iJ!~ion tL!!!,. S:nOf int;:~~s~~ Vil!rl1r1CC'1.in~.\ id ity at !h<l ~C'<ldillgcd!l~ ofmlg!;l~:i,Iq~, ... {/7, !9" Mo!Wvc;'!". l!lIt;!ol:~r\II:,,\~ o;:ll
Sjl.lliiai wlJ':tci;liicm hCi~\C~'Hhi~h-'1tlinil>,
~il

io-

WgrIi~~ and nc!jlll 'Wa~l)Jll re."trk~cllo ,I !':inglc eel ~ type" 'li~Ii1JOdy. or C'-'\;,tr.1!l;Xllh~ :Ul'!,rix (Fi,g, 4 ~'Ir " aad lig, S3it !'\..e!i~"'iCfLblill tII!l~i~,l!.cd,Vibmm~c~in
adh~irn~ !.\,"OClpl0fS,OIl thel,~d ing! ,cd!!g,~ my thus n I~~\'e a di~ct role i~~ ~':m.lIJIi!1gIhe 'C?i:U'acelll!I~!" IImUril\, dmingg nligni:[:i.Oll, We h'::>k.'tI 1.liIc lilruci~Qi.oo~ :i:\;JI~[iOn OC'b,'I'\.l"~l ac'Llli ptjolyll.1\:ri;r..aI.i~1II mid ;.~i\,;lh,"tI, ~,u intc!iiril1~ 11, alll.\.:rjn~ a.c~iI1 c:lyl3t1.ll'Iic~ {lFi!£.. 1111. lig, S4). :mdi D;:C'1t:1I5 iJl;g [he mnnlJcT or ~}(I~)I rl£"Ili.l:i,ng aeun u ends lb)l ql)p:i.lIIg willi cylOtlmhlll>iirl [) UlX:ulIIS ... tI r " ~1iI(; Liens,tll' .0 f li(;ti'!;'<l~{~d iml.,.'gnil1;:;:. whi,ic fl:!, inlCn;<1~illg~hc namber tJ1f ~}I}~ylll,",L~LLtngcrwtl '!N'i~h n~n; :-;m.hii~i~~:r.jm:;pbkirm~ido,; ,W) inK;~i!I$~1:I d,l!.:: :.1(;1i,\,1I!lt~ ~~I in[1.:griilJ dens ily ;rI~hc ,~}c.1.I1 pi.'T.iphCiJ)'. 1\,,1'oc(~!W('!~ int:rt.llI~illlg at'l iill l'lIlrnlOvcli" Willl.O'lIrl.climn!li~]!l1rl10 IlUm~bcr of barbed "'-11.i:IS. ~~i~1l. 'tilU! m}fo!>ima H adenosine tril}tm~l>hm~l'i.C ,[/\ r,l~) irahihi tor hlcbb.is~)! in, ~I J, 2m did nut T

·,10' -:5

(I

[)I[s!!ancel~~m)

:5, 1(1

Fig. 2. 1& I nl~g riliiS IfIlfll!.l'~5-y.rl~h,r.(I!:Io:usLy ith actln polymerizing ,~It tlilii! !C1r.~lits (),~ n1 p~I.e'S" {A,J w A:c:llYated 'l"aGIFP-.a n muore~~en(:e) 1rlitiaHy lo~i3ilijIJes,liC!!tUi! tillar a Iila~c:ent. nipple ([lI'(~and c:ti <'!C:CIUll !.)Ilal.e) ah~nl1!~h e entitr,e riilPp~e ,~s. it !etong:oltes.,; Arrow5sl<'!liI~ed tow.<! r,d tii reetlen 'ilr lFipp,te iIIlowemeiilt :S(ale ba r, it um, ,(B) VMP, w·hicl1l ernhances actin p>oLYll1l,enZallOrll '~4}, irs Il'!I~a~'ii!edartlh~ tip~ Q1r~din :~ib~r~ ,elo IiIgOJti!i!'fJOJI[o'[Jgth,~ lea.di ~ ~~~" S'>1l~~ b.;ilr,. 5< jJlnfi. (} HiS-log r,]lllll o,r lFipp\e 's[peec:i (n .~. 3 II, 2 "I Cl~~l!l~" PI. ~ieQ lIefilee from ll'HI'Ij!ie )3 ~hm\f:s @iFlhan(,etlllJreef! fllilo rescMt protein ([I) (EGfP);=.acli n and ihin~eg~il1l mO<'IIing :!lyru:hr(lrU:I!.~'slywiith the tilP or 0!!1! eoo;nJ~ating rip~~e (<!rr~~s). Scah: harr, :It jJ!m. 111i1e'~im ~Rila'~e of mov~eS] shows [GIFP,a,('lin and ~1 1I1fl1egrilils !lave oJ;;Qrr~~W(;l!ildi"~ ~Ilr;;r,~~~ ~Pl iT1ltl;ll:lst~f' 'Tth~ y~llkwrI il;Q Iilt@tlrilildij~ci!tl,l~~;!T!e ~iI'!9~~P'~;I:,~I.-\lll.~~Uru~ !!!~~OO ,obtalin theintElIii~il" 'traces sho\l,IfI ilnl (I')" SQ:alle lila~. 5 ~LllIl. ~Fl' IFlllOf@S:C!!Flt ftnMn5.~ty 011 EGFP\,acllf'l (redi)!!Jlld lit inID8!jrill1l ~glree'ril forimlilge- in (IE); auowhe'.'lds t:Qrre:5lIDnc:i lolh;o~in (E). «(0 Cress.(Orre!i"Illliol'l ,an,a~5i5 SIil®l\iS,i3I stl'®fiI'g (Ow~!arlion betl'l!e~1'II the 'inten$it~Ual.es o~ IEG,FP-a(,tin and

on

b@h~el1 EGFIl"ii(hri iQl'idl ~:l.int~9rijnsl:O Il'I\iiIiintai~'u~dl~~I)rlgi 2,0' lAnn tMgl1i Qrtn~ ~dnng ~ge ,Oil 1inelll,~ho,1.!t (he -.3·lfIl!irn mOl1iiloriill g period (cD:malatio:rl (Ddiciernl 0.5<3 .;1; O. B ,i3Jt the lead~irnJ!! edge, tI '" 54, a ne'O, OIl ~ 0.1,4 belil~ mlUhe L3imeHipodla, ~n 11 ~egi® n d ewood ,()[ ad he'.slve mll1,t.u.':ts, fI' "" 5•• ). 1 ~,A, to, a are INm3H (Jells· <'!r:td(O to H~ tllr,e human fore~ki!"!l fibwoblasts;,

Ih inteHri!1!aJlthe

~dge all"ld ~~'S~nthdly IlQ ,o,.r,et~ti~;m behritJ1!d the edge. Uii~ A h~g~r;;Qmilartil;l!'l

ch~nge ahc .).'c!,lvalcd [31 i~1te,gni!l de!lsil~. Co~~Slcm chtu~ges in ac~~va~cd!ll imeg.dn d",,!siW W';.i'>C c.ib...~rvl~d w;i~~ ~:iRNi\ t:5I'l'l!)l11 i!'I1.er~rin~ lilii\) k~l(1okd'fl~',1l (If till:: (Xllilin plm.~~,1it1iSC HIii1,g:;lhol and c.~q'iJ)ilflgproldil ~ (fi.,g. 54), D~"'\l~~lC ~~1C,ch:::r_ng,e!l. ~fl d~l1Shy. nile coi~relm i['l;i~ 11!ClwC'Cn ar::~il\"~L1.OOI~! ime~l'im; and po<ly uWl'kodng
OC!~1 ucmillh!oo
[iJ,1 o(l,54 '1<'i~l

h1~gh 1,('J!)I'iI'I:IIa~ ~OCft1Ciclll 0.,49 i'oJl no $&ati.~ica~ dimfcnce (/'> O.05J

993

REPORTS
filll. 3.l\c!ufI!'itW<i

but:

!!If! 6gallld

Pl.illl"
aC'lwated 1\11

'~gnln~ are I()(a~d at tlli~ tips; ofa,ctin

'l~OOFS at the l.eadi~g erlge.


~i()FI(illf !;IdiY~
~l,

,(A)

(Qnfforma·
!r1tfltgri:n

ili,tiNJJilli~ (ir~ ~~li!aUy O(Qn.'Cl!llliral~dat 'the tip's IUr pdlYlilleriizilllg .8(111'1at tllie- lea.dln:gl dge! ®f hnm <l1Fi1 fQ re:s1k.4nll~b~oMilMi. scai.e b~r" '5 ~LIIiI. ~B)Flu· ,~~~~nt. rrI_~~ltty ¢ ~'1IJ;:~I1' h~ctI,,~ctijv~~i;I ~ ~rnt@gnnl(bhmt ,and telRt (acme + irlcllrlliJli1') ~.~. inl€grin (gr~~) al@ft9lltif yello~~ IUrtf iii! tA). llil this exai!llptif',,~f lI;iI;lrl~~til!!m u!eflffid~nt ~~D'.,69~~fNll kll{!~ b

r~l

(II ..14}. To,ta l <llllid ,acdvei F1[egriIII distribL!ticm~ i£!~E! I!:Gmpiilrd in rig. 52D. Scale bar,. 5 ~.m. (C),S,aqUl!fitii!J1

~:lI.itlitll'!gril'l and f.,8!tfin, .md !ClJ'5 biltween ,actNat.edl III ]n~~grill! Q nd IF~a(tinl.The .aveorag~(o:rrelatilon (®e'~rt(j:t'iIl'ts'ror lliUllttil· pte (:!I."ll~a r~ sh!l)lll,nim f~g; 53 {illl!d Fij!J-;2 [I

rrtiOI'E~c:elilt Hop) aln.dl IDII( (looUIClm) fmagl~ sliIrnlJli'I.'9~hie a(.tnVi3l11iO'f'I of 11llwgrin) Qn IN~IHllB (@,!~I.s. [IIff(N'cnt (OI.!I)f .1r~(I'I',Ih'6()ds dliflf,~renti~t~ '1I'IdMdu<ll antva~io;n;-l1pedlk a,f'iilnbe~y blndnng :sil'e5 a~ the Lt'adilllg ooge' Ihat

rh

5'uiooeql.!u;orrd.y lIllove r'El'al"l'Jtlrd. ~erli(al 5l!:ate ba~, :t! ~LIlI'II. (ID') Boxpi@t snO!Lvs ~{tp,.ati'ld ~~ -inl€!Jrin.s d@ l1(1t (JCSs.-«ln~I~~~ Wiith <E!ll!d~,'~n[)~ 'ffiiJ,rmlimilil '" '" 8)

but do

it@JH!Late

with added FNl!.20~DVl "" 11) imd can be blocikE!'d by inhi:biwlY

ar1ti~O!!l~e5 (a "" 8' ~p.:; O.OO1l}.

Fig. 4. UH:;1~i~lfun
'~E!jJriflS

Qr ~di,v,~t~d ;ir1!I$,~ at the teadirl~ ,edg~ depelild'~. ® III

.avi!ll~tabriUt.~' ,r.r ~otyiiDerilingl actin efl,l:i5. (,ru INI H.3;13 (e.Uss'ta~neo r1:l<r acti\ia~erl [51 ir:rlegriinso (gre~!II)~Irnd h!din b~d ~rnd .gl'a~S~<lI!.e) f1ilow .. diE!( rtla~e'ill1i aniljla~lild ~:!Liru~9r~rl Of!IlI5<lty wnellaOiil i:S ~ pp~,d (c'Itoc:fiaiasin D, Ir .. 19} and ,ell'l1111(re.asf

adh;;::;,ioll :rCO;p'l~r;s m rippk rroaru~iO:I~ WCiI'C Ifnore Iii~dy !.o tc'gllllah.: rnigm~iimll fCliI't:c {R:'.I%. I.P = I 31 dH.IIHiOil1i'l~liO:ll.~ ftHllllcd ~[(!J~Il'.;nc8ijul.'ls aMllg ~.II<;;I~c'fItlin.g edge (23%. Ii = 'I, 2). This SLPgg~"/i.'1S ~hm~ nClIV<J1I.cd ~'I :111 tC'g~IUS a~ i&:td rt.i,,;'i] ai, (h~· ..:::rbll:sof(li;;,till ijb\:r:;.lo probe (Dr li~md::;." w

w'.n~r1! ~dil1.~ ~l{lbi~il~l?dl'Q hm~o!'l$e 'lilt! Il'iIllmber ~rril"e'i1l il! r.ds (ja~,pl)3kil'l®lid~" 1I '" 22). lin liiblti I1Ig myi[}~ii1l III AT'P(ils€
(b~ebMr$tall'iFl, .iIi· '"

un didll't{Jlt

(iha:l1Ig,e the

,mnceliltratiol'l" Scale

barr, '5 [.1m" B~~pI0t I~f ~~tw~lted ~1 iJlJl~!lrin d~n~i;t.y n(lrm~l-

I1lI,Hnber ,af~dirl ends or th'eim~!Jrilil

iz.ec! t@ {{J:nltro~("'''' 27}. (B}A(tivil~!'!d rjl in:lf?gJins are e:!lso l{l<cO!lliii:edtol the tips (If polymerlizili!91 growtlll (one fil.®IH~dia (NGIOB (eIJs~. S~i1Ile bar, 5,fJm.

between cj'I.oeJmla~i9 D {Ii = 2:1 i. j:lsiJll1Ikim:J!ock {II h.l)l;bb.iiSlntin (.l = 34~ and ~"O:I]lRd(Il 25)1 U ;'I). 'iIlhe: al.llUg.:.s in uch\.·;itik.:d Il! integ-fin deL...... 11m;:> ~L)' amL'tipomill<1 'the I~Lm1b~H):I·p6.lynll'

=:rn

lIitirnml.ly. till.! loe:llizal.iolll was nol ~ll~J9g00 by k.l1l1ckclt)w'~1of (j[ouctin ~Il L ~nmhCi.r di n:.::t:~ .~ilJ.kcr (2J), Ilmi. is I:I!)~. i>.irnir~Jrly e<m(lf.;'111.mKXi a~(m!.llhc:
i:;;;mlilng <;;dg<;; (Iig. 550. These dam ~tlgg."1iI.dmu

<erlzill:glRClil]

'Cilllt~.

'f.t) rLmht:r test whether ~n ~rll.cgrin" i:rI't~t:1 ~lyn!il)riziil:1J;: m:lin, we ~k,'I]liiCill1~rJII]!ll'Jcl cdl~ with 'fI.11011.X-IOO 10 reI11m~\." ~.h\."'l(]11.m!:r~1;t brane and IlItlJJl~br.liI]~I!l;:>(x:.i~~lrl IlfOlCing, blill leave hehind 1~'lC' c-yloskdotflll aud C"},!ookdciJ Iliy .)j;SOCL:HCd I'l~d~ ,( IS)" l\C1.i, ..ated Iii iOlegriUl~ qL!~ml~]mil\'dyC"oliX(!jHzod '1I.'ilh Ihc: ~i~~f; of
w.ili!

~11:~in I.l!:ml:ribu~(,,,. 10 ~h\." :liUlku.gc; OCIW\lI;.J] ~'n ill' h,'!JrlnS mx! polYI,'I1criziul,g xlin at ~hc I..:ru:liingm!g~. \i\k: ~]JKII round activated fl.1 illl~.:~riJki' Ioca~· iizml ~h~~h\."lips [l'I'grow II! eOllll~ lHollodill (Fig. 4B). Growl.II OCQn~~Uliidu H1igmlion by ~m]p~ing g Ihe nl.1Hcix for Hgauds ilnd Id~~i"c S:Dmh:-ss;(UI,

gl.lidc n'ligil"mitlll dircc~:iO\n, mEl 1.(1 n::gl.llmc migr-.J~.i,mi lolfCc. PQlyu.u;:rj/.;ttil;m of artill iill~~ d'i,~h' the !!llOv (l1!]lC'Ut ol <I v;uriC'~y or prmru sion~ r"ll~ill!g fmm :I'(l!(:kc!:in,g UU(';fi,~ (.27~~C!ekll1~<lni!lg Jlilopo. di<l U _~), Our dOl!a :inJ(ilc<ll~C that prcr~R\mi.Q.~ kl"aH~dl 1~(lJYI1l~ri:l::uO{I~ aun;1 cl(m£l,I!~{lI] of ~ a~h1 "km~~ll!:t> or HI~1"l"'i,~.I.(lngI~ !~.aoi~g 'I;'d~~ steer if'lCnl to.fwan;l I~ro' nJ~j D!'I~. The pol}'~ ~11~ri2inJ;lac! in H!~w~d cll!$~~~c-rllil alw ~.~,~e~~i !1j1 t{l i<JHlI!i ~ I~o~i'\i(l!l .~!Heg:ri~1.<\ ... ~y Ilri.nl~d~o p~ol'lc the ~n:ur1:1;,;).:t th~ ver)l !m!m ('!r~'III'l'r:tl~U1~ions, cre.l~il~g"s,tkky I'i~lgers" i1101lJil ~hC' k"dil1~ cd~. Transverse \'i1ri;:lIim~s in a'li!~ Ilolygllcri;aLlroOl11 .!long the atlvan<::i ng p,.;:r:in!I~III;r allow th~ primed nll~c~[ill K:(;.;plO:11i. 10 ;snmp!,; UIDn.: (If '11I!1!.: Imt:..:·lmulu inlalr.i 1'1: Ihan simple liJrwaru ~~;,; alld bilck.~I>';lrdn;mV'L~lth;n'[. This ~ni.~hl.allow edt..wiilh wdl.di:vli.:l!,op~cl adhesive comaetsto 1'11;;lke SlllIl::'!,11hlllllgc['; illl mig;r:lllioll dil"C<:I.iofl c wl uliHml :ievcr.ing !(Ix! 1\::t'-;';"iIiJ~~~~ I.heir malr.irx [rig CQiJ'IHcc,tioll::> IIIm:!crl]lI<;; cell body.. TllU:S lhc: ~:'I>~I1::rIil.dydynam it, 10.(,'1111 arlat iO'n::; ~III <lociin v .~ Ilymcri~,I.io.1Il probe the mmri:o;:fur ad hesion :;.ilc:ll by rruilionil1~pl"imod .l1Irl!ll~&i,on :[1.~'C..::p. UlT:;iI.I,rln~ Ih\." mlv3.n1.:in!£ cell peri meter clu ri D1!jl 1lI1i,gim~ iau,
R~ln·l!lI1!/n(I!:~.0I11d1INiOIIP.~ lL A. 1- AAI~ ~. 01, $~.e ]~2', :mot f20ro~. 2_[ I[)_ Ma'NJ\. (i,e.. Elcri;;r. Ct:l!' U2~45~ {2~.~~_ ~- M. (il'ill$ll ,et~-'" $",~ 31@11,7<1H~OO4}. 'l•.~~ BeiI~',~' ~f., G:,Ji!1~9., ~ ~0!001i. S. ,Pit J!tlii!iUQiiibi~. J. E, Ht:J.~~fiI~lfI., M. I'f!gruiii, EJip. [t.rJ .5;. :ill:'J'. ~'J. U.'l; (191m. ii. ,N'I.I&!!i:IRai:~k, G. Da.iliJ~. BOO~.; f.. '!l~I,1.!B5' :
~~floIII6,}.

pnl:y!m~rl?iin.g :lJ.Cl!!l ~l the ~cl!,di!lg ~~g~ ia th~ dcnw.mtbmnaieacdl!:1 {fig. SM). lllC 1c~~ti;llg eog;:: klC,lUzalioH (if ac!ivatcd ml:d roml 1$11 in,tc,gfil'lS ''''",s dcct'C<!..-..oo by s~RN A knockclown or ~(!jlil\ I. J cli[il;.'"C~ physka~ I i~lk~r'lO.:'l,:\,'ecnacl ill and l fj,! llllcg_r:i!ls 't J f) lllil. i~ asstliCl!iJtCld\\ i~h Ii lilS of
;[lelilll Ilbt.1fS 81ol1i~ th~ ~~"itding

oo~'C ~.1-1, i\d

<lnd !hcir ,(IICrigl-ll"lt'l'>'Cd kI;pod i~~11C' n~ccssilt)' for Ii oorl\.""l. f>lcel1ng ht \·'iV(!! 1:5). '1'hdocal i!:l'.(IItion of <I'C~h'med ~'n if!wgr!ns .~~mocPQd i...~~psslJ,gg.:sts <lI, role in g!~1h oom:il<:!lh-.~llm:!ing. i\dditifll1<l!H~', we ~estOO \IJhC1.l1Cf th~ aCliwuod adhe~ion I'CC~tltO~ JI Ihe ce~l ~:w.:rhnC\t~f !'Cgulawrl rni,g,Ta1iol1 [OI'CC'W m~~ch !nmri;": ~~~fb'~lC."$ using ,m by 0PI ic[L~~~l fl!.'l.S~ (17. 24, lfll. \\{.c. found oollItec~ i:om Ibtrnod ~'lCI'W()I;,'1t jjh~<OI~ect~nand the
2007 VOL 315
SClIIENIOE

t, ,N'I. MieKrIDiiibi~. J.. E.IHt:J~~fiI~lfI.,.s:. M. I'f!gruiii/. EJi(J,. [.rrJ

JIltI. ~;t ~~

(l:~n,O).

994

16 FEBRUARY

\IV'IoMIV.sdencemag.orgl

REFORTS
a, G" Giannrn ~ ,ill' ~,,' (('fIn!>. 01 ~Zll(l'l1t 9, N, 'Waiaili1!I:!~, T, 1. u.n.lhi!oo\ SdtMf!Z9.5., '1(1:113. ~:1Q{m" :m G.. [i\!Mleil. It. 'OhiMlxi~~-g. l3i~ J. 'j"9.• ],.91
WKll1),.

'i2, P. MaU~r~!.!" 'T. ;,md!~o!l\,t, ('!l'!!I NaI, :tq6, 1i!Mii)' Un9}. il, 1'. A Therrfo" T. J. l.!i!k!l'!50!l!, L G, lilmey, C, A~JI!O~, NqIlJTf 3;51,. 25:7 tl.~'92}. H, Go it. i"~~er:l~, 1. LI~p!J!wll-S'lln~nl, Sd~,"~ ,z~i1,
1&73:(.200,21,

u,

T. M, 'S¥ilk:im l't ,rJI•• }:, (J!{r tltar, 16(1. ~ru9'(200lJh

m" Ncr!l" l'ItM!:ilo:o"l[I. T. Burnfnf, ~, FOI'9.':rnn. NmI. C~ Il.~. 8" 21~ '(.2006), ZL ~ HOIfl'iit!.. tt IltI!ill\!ll,. (. ~1J~t . c, El~~rlf. K.SUEll1iIiQle., M i!ltl~ ~ll!D,s;n ~lt5'U:k. zz j, A. IDI!-ralq!J~l'e.. I(. S, luata. j, 0/1 6ifJ1. 113, 1B51L li99U. ~31, C fl, O!t\l, F. ,rlt ~~~lto, K.. 6"'f!1dg~, 1. CtiJ Bfa.!. ~J.:!!l,

rtrIil

me,. tar ihe' lSM ~SLM ~mllSI'In, DUO" we ~ 'In~nk z, IwIn~i,'rrlfCMfflal~gf~ IE.5humlk'f,.

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~, .)c, ~ )c,

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l!L iMa1>;!1ia,Lsmil, m.e1Iood~ <Ire CI'I'.ilil;ltlle a~ ';tJ1ilpi)m.i[j~ , ma~eri;l! 111"1 S(:I!'l1Q' 'l)BLlne·. 16, T_N~za~ Q, 5m~ M. p_ ~'-.tz, ~t;. Nl:ln. .-lan! ~l USA '!ii'. 69\1 (,2000)'_ 1.7. (- 'Iiio_ ~lbJ",illi1" It ilL 'I',II1}<1IJ;1,. He IP_5~11', J- (~,!tW.I. 159, 69~ R(!IJ.2~. 1.8, \'11, R, ~Cl59!~ S. ]. :5liaiJtil, N, P.ampori.M, A. Sdl.... !'U. ~ NfN. (fiJi 8itJl. ], 3:l!.6· (2'OOll. 1.9,1), ~, n:tsenlelil.. 1', II. Stim.lfb:ber1lO A. vemegili5:, R. T'5I!'. lot ,Po ~m, Ntli', C~l,fJi.~,. 1, 2.00 (1'.9~!11,

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8e~l.'!e:'I'. tI:. T!NI!I1B"-!t.3ot;mINld, NaW""ft: 3Z3, ill

,E~nldm.. ,~nd). m~ ~~l$~~e-; !j. ~at1m.'i1iandl f(!f ~ liilPii'J(tm~~ fOOIPoIG~f ana ~J!UJssirn~ <iilllll ro!, Ha~miI~1 ~5ina!1ile~tl ~'e\lerlelra~~lioo'f!!lf' ~ef;m ,dli!l'l ~e«:ll(ifl5!iw:l!~ ~i!9~tl; !lrwoi:d~~ R, T'S!eJ!1 ~~ji~, j.1#eI11ii!l!d (e.~F~lO!S~. <!f1d' s. ,!,~~, (!"I:!I!!lUlI~es!;'iB ·~~~.su~~ r pro'<1d~ !;w (tie hllilm~ilI ,re-m ~~f.lml of NiNiiJoS i!f1d~~ tru.jUlle .;.l·IDeRtl~ <I!l:I Cr.l!l'bIiOB! Rm.ardl, rtlt~,

t198~~" Zi!... 0\ OlJJI\1~e~,II ~.. :~Lmlle!:rll, ~_ P. She~1lI.. (!!III' ,e;s:, :l:~

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,~i!l5. ~tto Ss, '~rer,ences 1Mmle-.; :5,1[0:], Sil
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R.. Booj~III1\1a> D, P'afl1a'!mnii,. 1;1. F,. Cartier. ~~u. (U99). '~H

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211., m lhamk (. Sffii1l1 afld 1. ~~~ lew :!IJ~~ij!ln~ U;'IItI!? f.1.~el1i1ienl!.~ lid nlart.lJ9;lJlM Mld M. StJr!:'tl: Itf lJ~a~ OJilical
lr1ip 1lI~1~ toIl«lM tl;r C&!G. v.f1if! in,lii~ ~boI~I~. Frlii'U~ mtfllo laSllIl' ~nm~ mi!lf(lS)ql~ IJWwM:lr.ti: 1t>M1~1.

IMii~ui1!·of r.telJljolil'gi~~ 1JIm1!'~~nd SI~

(Nl!'.1DQ 1i~1

,2 r,!\l'li!iIIIl:>er ;t!Q(l1l-; iilCfpudlS ro.n~r;.r9.'M1l1oe, D.B7'l'Oo1i

r~m~ilf'J 2'007

IMap llke Re'pr'esenlaliio'n olf CS,lslstii,s.l1 E-Vecllo'r Or'iis'nlalti nns in t:lhe Brain of Bin lnsset
F,o~ mi1!I'!Y im.@!!:I;s, th~ p~ la riZia'~(lnl p",Uefll Q,~ '1Jh~ blue sky ~""@'S as ,~,(i(l!'!llP~:S (!.II@ for !Sp\ltioi!~. F!ii!'I,I'igalion.. EC"'~K~t:U ori~l1tatij(l!1J~,~r~ ,~M.Ed8d by ph!)lur~W2IH()rS ~rQ ~ dIJrXll rim o!!!rlla(lIr~h!! eye. P(ll.lri~@d-li@hl si9 f]I~lls; rrO"Ull twlh @ye5,alr!! IiJlii!llyinoog roilt@d iilil th~ ,oo.nt~;Jl (om P~l'l, i! brain ~Irijll ,c:~nsiS;~l!1~ ,!)r t\~O s!d:nJlnijts, thl! p'rorlQc;e r,~br,;!l bFid ge ollld thecel'lhal bod~. Her@ WI! 51'l~~ t hal ,;! 'l(lpographi( rejpreser.taltit:m O"r~!ffiIilhal, f-\I1~d~r orie!lt<Jliloli1~·lUrld!!rties the (oll!lUJ!!'lar Q['glllf1liliOlliIJrlor H~~ prlltgoc@lFeb.ralbJijdg~ in <ll Ill~'uiStf1he liIlap'li~ autll'lgem!!nt i;. Imlghty :5!.1!iledl~ S~!l!ll1!~head oQ!ief!Mh~1!l' !.l!f!ld~~ t~1il QPi;::!1 ~kYT

lfilw.d to il~inJltec6lumn
HII: ,PiB.Wh~'i"l P.H

in each hemi.'$.lt<:p: of

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l.l~ilh,:ndit'to:;. colu ~ltt'i iii tile Ie: 11l:i:iii i>j:'illCrc of Ih!.: I~B i ttl U (Ul!'.Iist liJ)~dii~1 (loliJl:ltmill, 11].;: I\;:I~ h...:rni...phcn:.:) and, flcooLtl,ingUy. fum .1R.1loRE in

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bilt.r.;.. f~i'h.es. cer-Q1r

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mO!OU' eo(mHn<lIion (11)!(1~

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....'.... __ ity W l~i'CCi\'c .I i ~~r.l),' polariiOCdngh! {t, 21. 1111e ,"hl~lC'01' ~ollari!'ali(m (£-VOC'tflr) v.. ries
~C1!]1!~Qic;)lJy aC1'oS:$ ~1.C hll!": skyand dtq;1!CI,ds {IIl,lIhe 31tn'l' pn:;iLiQl]. ,fo:raVl.mi~~lYof' ilfLWC1.'",dli:; p,m:":rn Im~1x:<.,"'I], shl1JWIl.il) g~li(k: i>p.a~ial orimm· ~iOJ1' I( 11, In iOClll~Il.~, puku:OI.!:.lc'lnc oric"i"tlLaliol'i cle'P..:tJKl.~ on :.l :;po::~i.t.diwd pIBn ofll~\:" otJmprn.uud c)'lO!,

~~r.;!,.~d imnh"C'J'IlC'!!!i,n vi,sual p;!ucm leami !lg OIl1(lrcoogn!itioo (J.?t I n ~hc IOCLI:';!. :'!C\'crll~ cell
~ype-sQir~he(."(' are scnshive W the oric~H,niolfl oJ:' ;a:ui!hlill, E-'"OCHm, bill'[ dl<,;: ('''IJiiI\:~rUl'l' tlCdtC~ (]of rei I morpholo!iY mid E·v~dor ulm~ t:Ig 11a.~, rtlili~in!l)cl ooSCu!i"C. In Ihli:. ::;h!dy, we h~N!..~ lliK..u iutmc:cHIII ~mr ruNm.l in~~ tou"ibiin..:d. wnlll dY!.1.Uujl!Clions '[0 :lllml)'".lc £-\'of.::cto:rumillg in C'C It~mmL~ ,of ~h(;kJelJ~I. with (;O~U.II'IIlJr aroo:rizll.· lioll iJollmi rl$. T,>\'O nmj;or ckh."'~,""of iXJ~nril... . 1.ti:(JIl-~m;ilh'c tPOl.~ rtt:1lI10l1~ wCIl::cnwl.llnI ...rod I"Cgllhirly \.\111"':111

lLil1!C1'dc. a iJ.a<lin ~l ~rIl111CC1.~di:l <Ii ~"In!lellmp.~~ ~ i~l~lflC' QP~k; flbc.tl'K! "oc~~!"f rncd~I,II<lI( J 4}. I l1ile Im!~mQr \I,uico!S(l .. roori"~1I! ifillS; rul !he

I(Dl:l~IIU1~ (Fig_ lIE), The ~!m~ wi~IJ "'.;;Ilicosc fl'IlL'ces.~ were free of l<tJnifk:aliOllS, ·~lI1ds;ixIn d~l1: O!.UIC!: colUl'rms v.'Cn;': i.nvndc:d wilth Ii~ !'II11001~~ ":Ibmi?;tttntli;"\. FUl'1h~I"IlO.l;c, <III nn UC\lmus :]loxl \'ari~se :lr~~1Z;1Jt:ion~il'l~!~~ poslel.'ior opt!(. 0'!1>e

~n

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oJ 111'\:1 fUUT i~mcr

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PH oflrrc:'ljlOndl;{l~('J pllj'sinlo,giro~ pm~Jtks of 1111(:' 113~ncumns, Fo, ill.1!J ... cdl~ll~r n.'Cordiug.~.lhc ~llhni)ls ',\'Cre~1ro'l;W i~~ dlC':!:'!OCord SCltm' ,~J1d i~,g ::;tinnJial(l(fi Iro~1lI I.HII,; l;1';Jl~UI w~~h ,I Il;mu.ill>g E-vcdJ~Jr. R... ~co:rdilllg:;,~'I,'\;n..; obHLlllOO :i"r{lnn ~hcitr
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:ill 'tli..: rB. EiLCh

11] ~ IlIi.'lLllt.'01lI :;:h[}W"L-d

!Wbr,it.,tlInmIOpiffiilli.:Ucy. i~, £·:~'OCWr mi.~'iJl1;l'~h,.; dOJ":;:!] nlnl:llU'l QJ)\ aD,d in\'lI~v~ sc\rel'd~ (ii:Jt]o I.ions le~.d:i]l~ ~o jI.l il'ICft,\3re it:) !!pi"i:I"lg:.'OC'~iv it}' {t:.;\.ci~;~lio]]b "~II'\: Ilricllcm!p.crpeL1d:lCtil.~1J[ly w ur,lll, proc."C~iIlg tn3:gt~i. indl Lidillg the ccmr-.d t:mll,. E'.v .... "CWP.l. ~GiNlil"t!,g , u dearca~ ilil spikinlg ;"H;I.iv· m piC)< (,i-i).. The renllT'dllronlplcJI! !~CO i~ gltmp j or nl\]lilIDpih; ;;.a:rmmi:rlilJ l~l~ mid:iialc ohh..: 1]]\:;(.:(;' iLy (~Ddlibiliml' (P:ig. I, C nm!m, 111.1:E·v~'Ctotr lnm:llli. 'S~lb:;'trocltlrt;'. m-c II'll; ImJILl;iICc.I'hI",!3 br[dgc ... lunil11J1 (e-vc<:Wtr OJOC:lJIlIloOlI rt~lIlilillg nil m~;:o;;ia BlUm IO:xci.lmti,ol'J, dlmil.. ~W;ll~ dcl(Trnnno:l l:Or C-adl {PEl} :;tilrl~OC ulllK"r tmdl lower di"iis.ioJl:> or ill..: .. VI:: 1'I..'t"!)l'(kd fm:lliI ('1(' nocun:mt;; {~l' .paHiclIlar type aof ~;'tn!;.L'Ilti.~],UCll..1mil of tht: PB mid. (ii) cL'Ttllrnl bodl:-,. An m.l1l.,>l.!:.Ilh:l it:Jg l.1IJ1~litomic;11calun:; f IlICUIPJlI] b~!cir;c1l~11ut ~12uti51.oc[';.R!I.)lh.:-j~l'1.Cl>1 ~.r )'1. ~ 5 :sc... -crnl ryp..:s ,(lfrotLilUllmr utClIW.t1.,. T:[u'I~nl:i!~1 E.vlX:lor Hmirl!ll ,ofIHl 111~urudrl.;; shm',;,~'tIi.:3..liinl.l':lli ![j r Ih,: CC ~.~ iu; :rq:.1I1~r mid II ~I.dy :st~p~li~lic~tb..~ l o i!1.i.oI!'m;ll, ~'!.ellmru'ch~!cc.~uWC',,8 N'), I n :o::~olJ)I~f.~d !lCli,[!:(lJI't" f'lf ~hc .PB,~t'fIt]loo here TB1 I1IC1,I!lL1U!S, rekl!im! W the."l'IO;Si~ifl!~ nhdr \'.~riCfl.S'C r.ul~ifk::1Jhave Ilot l'llCetl desli;ri~~ prt;!;."\ i~IS:~.Y in ~he l'oc~s;[ ~iI1Insin d~C' PI) (Filg_1 Ft lhc~hy. " rnl~ge of tern'll:>:, il. cflus:i:;:r.s of S!;;Jcit<;: ,flf <tJrr:lYS:. 'IXtCh ~;(HIl" or any ,oid'l'Cf hlSlX1. A~o!a~ !w!ahcr 1~ of IhcsC' ~I)j;,a"\L1!lingBof IH2~ ...71~ c~IC'Hds tl'l1.\Qllg:~~!he 'j:nosC(lof tl~inear a!'raflg,c'll~I;.'Jlt ofl fl OO~U!I:!I1Swilh cigln colu!t"!Il:1 (II' ~oh hC'lnisp~lcrc Or~lC ~~]3'elgl(l. W-POf,lr.lpJlk imC'rhemisphC'ric cfll1.n~!iom;· ~'.oclls ""'ere" a!la~J'~C(I. llH:ir nm'f!~llfllQ,g:)' w~ !:hl~. m~'!'ds 10 th~ wl1(1\l:1;!,ll101!:C'flJ"possiMy ~'~reenco:1 ~Inms. bo~] '\~, ill "11d bcl\~"(r,::n <.Ii r- fCV ..",.!.;:1i! byiiflll~l,')pllQ;l'ctlcI.r.\"cil:'r il~~'!Qjl., his.[}~Ih GCC'!.nlT.illg E-"'C1Ci~]1!" (;J!ri~nj:!itiom;. l:ercl'lt arra:ys.~ IYl'lalhe~ ,on the flllm;:~i.(I,~a~ Io.gicalp:rocessi!~g. and ,cmnera lur.;kh !lCc:(!~trucBCCJl!1~C' Ihe E-'I'OC[Of i'Il~l~ill thcl"iB Ci"~rrelion ~13 ~ TI3I I~Ctiro!1S 1"111:0\dc' <'11 CNHtOCI km i bC'LweC!11 a ~'!(IS'liciio:rbrainl)C~ion. 1I1~ Il{b...Wrior . SI,}O,1'!d.~wid! .he :~ropos:oo 'l1\L~rnll regjol"~ of .he AlfIim<ll ,Phl%ioLCI!jj',[)~ar:lrn:I!I11 o~ BitJ.lf!9'l, !Phi~i'ppsUlIliTilj I naumns. \.\'C' a~(!d whcd1<:."~ cilndid~[l;!' 1l'0s~oplic ~ihe:rdc, <l!tul ~le:p1J (Fi:g. I, A and En I~ch t 'I'Niil/i'. 3~5i032 MJi~btJrg. Ge~tm~l1'. 11M l'!eUfflll h~d w,'(]; (!on1ajlls ohmicosc <1100, syn~!J~.knlcul.'I,,)!J'1:S SlYrow a simJ!il.f repre$C!l~1HiOl~ ...~:c ""'hl;l!~ lll'!"!~~I;I~d~.moe ~~d 1J,e, ~dd!lil5"lli:d.E'mili~ thus. rlutf!.l~vdy presJllmp'ics!I'l'll(lril.1!1hms COIl- Qr E-·\lcc"'!;ox~mnhl.bl. COkU'mHU inC\! rotl5 ha\'1! :lrII!I;I!1!'berg@~ti!ff.'l.!!1i'·f1!'.i!1l1u!llI,d:e'

or

995

REPORTS
00.1, ioes in silllgic CQiLlIlfUl!': (I ~1'Ic: I~g and 10.00 40 I lz, C(IImrl<lnron ,(l:f~~ICirn~~wrn:~'d serul ilxomll projcC:liollS to an firea QJjlSkI~Ihe tlll'll'l.'> in ~l'lli!I)Bt'U1d (Jillmc'l, I L.miny '~ili.I\.."'Cidcd ,:1 Cc. the lil[~1Iil1 f!.OCL'SSO'i:)' lobe (LJ\ L), SC\~~iI1l~ Sll8lt:if]1 ~~II:c:senlmi0J11 (I D .... ~tor ori.cmmio.n::l, ·
botiZl.lI

c,;;ii

h.r.'>c:a(]d Ition<li anl!Ori~,~~kms ill disl HI'l'C1 oOol!JIlIil':o; or I,litc (;';'I m:ijl rody. Tltrcc IYilJ¢~ ofU~"'~i:
'L}'['I'-:> nCI,num.
aL'C

:l'llj)huci::!lII.ioJ1•.~n~a~uvc:

tF~g"n We

c\'ailm'L(,;d the data fmm :If)! rcron:liugs fT~'1mth!.>:;~ cdlly~. MfIIIDOCPU l. CP~. :imd (,P2 I](}I,IIID.UL";;. CV'lll lfIa~lroJll'l h~\'C smoolh CIIldillg~ in M Ilgle COILltl111~ Qfthcc .PiB, in 'COhll11tl.'l on~x:, c:mmimosl d b~~r r the Uf~PCf dIivisu!IlII ohlrh;: CCl1lml body {CBU )•.and an ~::o.:OJml j~~r: w.ilh vmi~se ~nd:ing~ ill !lli() tAL (Hg. 2l\,). Eachr-el,m:m connects !I

or

::;,illgl~ column

ocoluo1l1s {lI~·lhC' eml, fo~~rn\ling;l

or ~&u;: with PcB

h'!,\n

ITld~.hlXllrin!:l

wijrbl:g .;:ill~m~

d~ribcd
01~'I!1I1.L'!l.cy

.UCll~O~:s. (~(]

jhl:.DlruIJt,.MI(I{9! ~.FiR.2G)..AIICrW l~Ofl:i.i~~gsl' s.ho..~'~~ tlo.lm7..'lJtrnn


11:11'£1Imd::gUXlllll'id

11C']l1I~firhLlf'C (Fig. :3 ~I" r.lm ~Iop:: or the n:gn::s~io.la .Iint.,sn-1"l'm and (,pm .lfIc!~lroJR"i not difth did t;:iglflilicmn ~yW = 0_52. alfl:Jil.y~iso r om':uilanoo (.1611. i_nrlic:uing 1'1l!:1!Ihc III ning range !I~roughIhi] I(\, cnlurnas of I.lllO Ib'J~dg.cnmJ1chcd :IDI' I"llh Gell ly~~I-lm,,~¥cr. thc('PU 1 map Wll~ ~hi I,l~ hy I(H~ ~Cl!:]M ivalent ~:o - 7~)") 1''GI,~~ii\f'C' If.!'Ih~ limp of lB, ~ I1~UL'[lI!l~ (iligui~-mJltly di.Hl~tTTIt~1~"rn~iQfIo.Y [~~ioll' lir!c&'P <; GUMJO I. ),. The ITIlll'liniJllg ~~~ro~~ ~l'rc\(ll~lml~:r. ne1;lffin,,,;. ('PI 0l11d CIf'lLlC!IIrolfL\iill~ci1'lllrec'wd Ilh:c PH wiU.!

eeross d'.cPIJ. which coeered 3 mnge {)f'22X""::: 7J~ 1J'IR'mghlhc g :l?B eohUllI1::> iOI one brnici

I~r{)jccliens were con ~ilncd to d~i1.cy o '~\~'O 5liJ0C0nlJ)fl!!ilteutts. the nncdiiIJil Q~~~C f("Pl} @f the
~LI0l1ii;~~fI[lgk; ICr.:!), Bmii types of llCln'On l"~ Sll1.(~Lh iltt1()tll,;mi:oJ1~,rll the PB [ll1.tl 'W]moo:;;c Clbtl~l'I,g:; inllh~ L.Al.. IU11n iliI"COI;dil~ III = 9), CP~ and CPl neurons shm,vcd. IlOl~ri;QLlk1fl ~~PQO looney, bu~. h:Jdlm\ICf b.:lokgroolldncln~'l!.y ~3 1.0 ~S ~r~:t~' CPU 1 Hrld TBl IK!U!I'OIUL A~ fQr lh~UI CPlH~ltd "fBI nClilrolls,. OOMffi'ok~ijQl.1li1m.Lysb of pooled WIAl rmlll C'P~ ,md{,Pll1KILul'{m_~ :l'C\.-coakd a UI~ClIU oorrelmi[)D1 'betw.cC'n I!mhlllel:\~Ltcd colUIII]]] illl the ~FJ :md Ihe £.vQ!;lnr ~~l!Iillg ·o!:lhe !IHl!lmrl1~ ~ !.~ig_211.1-"fhe Ji r!ctUr [Q'§l:iJ:".lliQUI jm CP I and ('1>1 COVCOIW ,I m!l~e of J[l{1q ~ ;(11;> ()'vt'r~mC' l;lCmi~.)I~re '(,eij!Jht ~IUlnli.;;) ~f th~ PB a,~cl 'W;,!!'; !'I(lt ~<;;]gnifii;:",m~y dinbrenD Ih"llu IJ'tt;llil!l~;lr ,'Cgm$~ ~on for Cl'[J I l1CUt"O!Il:'; IS:10I1e: P - OA7n. dc~r.lli{m: f> O.3S25}. It wm;, ~lO\V(:l,lC'I'. phiL~e· shilled by I 1'1,'" (~luivflic.m to 69~1 tlgJlilu$~ Ihc IIlif!P ~d 1"13: i nICYM'IS, TIlcptC-:iCnl! ~Lld}' • jI;IO""'~, ill.dC-po.;uK!i,;utly rOil" . ulii~ dif(e~1iI ,eell 'l~vpo..Hilil. fu r~Nirr'()( :Ii~'!IlliI1n1111 E"II!.1ewt ,tlricllla1i:oUi> um:lcldics III~ col uuilJl,lt (j~. llIi~niun of m~n.:~OOllM. PI1 Tlus ~rli]~.ifll 1\:ipr.'C-:.en· Rllio:~ ;rudii a. Icv.,,;]' 01' ("UI11plc;'I:.it},t'o SCU:><iII)' illfOCCs:>ililg in Iht:iIUSCC[ brain, hilhcn~ 11]()!!J.!,:Wn to be achic\ICtI till~}" :iu v';;f!tbriuc:;, hi umM neurons

slPikhll,!;l .~c~h·.ily

or

dm LAL

(Fig_ ~. S, C. ~Ild .1 ~ I

millOJ:D?:JIQ:iom;. in (liIC ccnuaillbody_

\Vilhill

oot

~;rck!cd ~ilC' l.AL.

at

kll:;~11IIU:'nnkXim~'1

il£Iivalim1

m .. ';:lIIr1fo~'t!OV~'I",ln

'C:

..

3C

II

I I
II!

I .~

[Ii)

..

E-'!f ... "C~tJrmicm~utit!'.ll. 7<1.118'; of i~hmjlt 60". nnlplyi],~ 1 roll:,i(l.r.d:i~C lll\,'I,.'fhlp Ibe- 1~,.;~~hoolfil'l& cOhn~'I'I1i:'! and Lile nil;:~;il.~' fl;f ~. l'1Ollul<1!tion cod!!) (tw~~
j)

iI'Q 10 !O
(I

10
(I I)

i!IO

'!7i1!1'

10 2C 3D

1iliC1'

~ao'
P{I

»: 211jJ

I I
[IF'

60
00

"

(
100

triC'",ill!]l 'CX1tClinfft[1n:uii:m from 1.liK!: firing l';l~S of tHlCilC' ,ceJlls<{ lil IJ:rock;r ~le Oj~ sky. the O'Ic~ivity~ll dnC' POL I1lCi.UI.'QI!I;'\ (1~li(;lib..~~ h:~ is dil.\'''CI~Y l:e~:ltcd lO Iloc

dip.,'lCt~fI!'<'Il nr~.1;;I~ion or~hC' ~Cl..;il,~fs. llA;';td. IWL~ vidt;.o(i (l(.k~itiol'kl~ !i1e::hauisrns: (co~r cOO~'~!,!.
1<1 dhnillIclui~~ U-.c' ~I",~' limnUilc' "'nti:<o~rhc!~]~pl1creflllC iky. o ('oto.r"crJtili n,g pro;J'Iii;'!iI~$ ~I i~!c.d jo Itrlfi ~I thi,>;. ~qllirenl1CIll h;lvc ~X'lC'!1 dClllOl ~u'['l1.(:d ~nll}' torr POL nanons at all ll1.JfUiI.stage to the (1C, the WlIll;oor opti:c M1Oi,;R;1r.: UN). Neurons ~11~o<IliJlig 11lI..~.lCj din.:;(:!liifin i1~Vt: been irllJ.:ll~,;J~ sr!Jdiik"t'iin ilI'wI1'1il11Jiih.'l 119,.10'),An :iiliipiJrtlllll tli 11~n.;ljtli:: or IltC IOOLlli'l Il()J~lri;&It:imlmm~yil{,"I',s 11~r;; ,[C ~~1l11Ji:ili in glDb!ul llillure {l r 5iigll.:J1irlg. The IU:Umnil Fire ao~oruing 1t1 ;!l(;J]ili1'>J1 E·VtJC:1UIJ;, wliit:h p:ru\,'ide i~llC!lsily C<idinj;l)~.~~...tl~omim",1
inl(OJTiI]IJlim~ <lwn
ihr;; S!Jn:~~zimtl'h

,h,,~

10

2'0

4501
3B{I

~
~
1; g

r:
e<

2ji'O 1,~

'.
'!!!I
!!!Ii

'00 '0 1.1

arlit IhcllC-

I'!DliJ

~~ !.!i
'y!!~,

L7 IFUI R.6 R4 R2
~mi!li~ [II 'fhII' 1F,e.

!bIt, 1'I.(,"SC. n1:UU~m8 bch!lIV'l.~ I!ukr;; [m ill1£ml'l.i, IJOCI" am1p;J:;S. Iu !lit!!>, in 1(.1~)IIIL1!II:>t,. hJ;;mJ·dirt!(lii~)I1 cdl:;
:lOC'

Fig. 1 •. 11,lt01rphQ,I~~i'C<lll<lInd phli's1C11~Cl9n;(,al propertJ,e~ ,of 'il'B1 11i'!~ur(HlS. (A ami B)' ~,rolllta!1nKO'Iil· :str\>lclllJm of t"'' f(l is] II1CUlrQm; Insat shows h~mtal vi'~iH~rlJhe kr<I;)UiS~ br~lim. Sri;~~ ~ rs.• :100' jJ m" lmod (C <ll1Id D) (in:ul,a r p~~u {~r H!!:'I1li Hril1l(j rate dlil ~ill~ E.I!,I@(.liCiirmtadcm:r. fCllrllh:e< mel! rtln iFI (til, [(C)] anI:! n ~B) Ulrm tn' '" 4, bllfl ii\iid~11 :1lO~; '~IlrO[ IlJilr$': sm, S[)~idlllinE'.s ililldi(<Iil'e IMd:groun(l ~(li\l'illf. ~E) WilTing 'Sc!h,e-m.e o~ ~I'le llBl neuml'll ~yS~NI'I. ~ach Iline rep~e5Ml'5 one llB1.neLlmln ·1111he IPEl{ibIEl(k sqlil,a;~:l!: t ";:QI~IinI!l:;wHIl vc{!~i~ors'~arloo(r~~tkI!'lS; 'fjf!1JIY ~q~,'I~S-; ~1;I~l!,Imfl~with ~'lir].~~l~!!rbQri~>jI~lQPl~; wtri~ o '5C1UllIr,es: ,c:olllirmu w~t:holut .. ar:b®rlzations) . .A5t~rid:s indkate TIll lil,eU~CIfll:!l~hi)wl'i im (II) amid (B). (r} (o,~~,~tl(J,n b~t~'I!~·~t1 l.o(~t~onl of varimse ~I)ll)lmnl~~n 1M !PIE!~,nd :E~v~lIl:r tUln~lilg ((~'fii.!i:J DI TiU . neurons W '" ]8) .. Ille'r.au~e ,eaJr.':9IfIfl:lOOFI I1lels v;al'h:o·s:e ,arrbori ••H10fi,s.in :5,~Filgre' C!O~ummsin tine M!ljlill .a!ild !gh PIE!hem~spheJl~,,ell"",,,, vo1l~l.!:es w~r'~' r;lwn~ lwfi~ffQr tli!,~'right heIi!11sp'~~~1l' (open drdeS> and HIE:! lef~ file mis:pbere mll.oo (nrdes). L1111.earrre'g:re:s~on sh 0'W5 s;ignii~kalf'it (orrt!latn,oill~r ;; 0,$1. S;O ;;;; 3:5.&"', P < ij,Qmn. l\l,Io,.tailed t ties!:. y ~ 24"2~ + f7"O~. Ai-sC! £how.nl are 9'5% CD:rI(iti@Il!l.:1;! barrn1,s
r.o:IloE; (, p05te!ri.or o;pliir. tulilerrde.
~Tl). tlL,i3lmtermal

It'lC~liibf'db,;'(1

[0

V051l11i11 l~llIdall>JrkS

ura('''l.Ichll:t:\\;,

and me .:JI11p~J\.'11ll~1 Iml. ;JIIT'Jli!}OO WlliO~p'hic;III)' l21, 121, /\..~ll·L{lLughsome p:mgociii> hrl\ been made ill w1ll1Iy:;o.il]glll-C \virill!l. WJ!'ind;pb rmm \li.'hkh crnl~~1l\'oroJl]w(,"U]1

pm,u~(hIl<!1 .maps ~J:Lveill vertcl~~tes. t23. J.n. n~ ~~,pt-clmlh~s of I(lt;:Ll~IS offer .be OJ'IJ'Iorrli!~ily lO

,()e'n~fail ~OIFli'lPLt');;Mil" mUi5!l ro@m 00 [J~; PB, prr@'tJlJ(erelJra.r

br~dg~; '~li,j,

iX.t{h)C~ th~~,qrtl!~ti(i1l.l.t the !G'~'~I f s!!lg!C k~n; o ~ifloo nll:~lron.~. Wi~li!l i~lC' [3, Ihe spill i:.'li: n'!ap~ of P rongr;m~'liilU1d1c(il'~!:fImar t!cli!Km,~ Ji"C o!.!t flir I~'a.~ by aoo~u IJ!lP. JI" 1111 1lc!'~f'(UlS :!ire (]~OC\e!lyOOHnccwd w~hC' c(l'h,ammrnc~~i'flm;. a ph[.~ sldrrt of90~ !r!h~~llru~QSI asi~y result rrom iFlhibiwry c cO!nHcc~i.o:i"ls <0 f 1'.1:1,1neurons to I:he clCi!ldritk

16 FEBRUARY 2007

VOL 315

SClIIENIOE

\I'V'IoMIV.sdencemag.orgl

REFORTS
'~rec~ ,of co~tJmnilf neurons. Do<ubl,cHahc:1b1Ig cxperimems siloMrctilh;Mtscr,OIOIiIi];;tntl nlDJp:[l~Ial.o".'ilmiu~rclml)d Ii()L]rop~!klc are prc.'lI."'l~' in Til I Ii';LlronlJ, ('Ilg;. S5}. For iloolh :litlb.SUlUCC.,>" inlii,bil(lr)'..rl,'~ls hw.. e OOC"JI dC11110l1SIIill~d iu OlilCr- Hys;fi::n~ 115, 2li~_W!Ji~'I~ m!inn.l'till:i! iSpHce is lincmly rup~lllil1d in dlL~ l'O~l!Imlll<;; (If Ih~ [lB,
the w.irirlg ,of CPU InlCUiffilli1 shou 111rc.ru~t in
ill.

[IOC mC~~.~~le

alimUlilr~'lI\I\"Scmmimt ,\1;'oul~bc
{I

supc;pcl£ilion oHhC' tWO ~80'" represemmiotl$ olf tbe 1)13in. niC CBU. btl. with a I~lcml Sl'ilm Oi.f uomll 50" (rig, S.H.l1l.c iunCliou3~ eonsequences
01:'Lh:is ~lin m:c prc~Ul~l:y
voctClr lUIDIilll&~.
ill 11.1.d!

.rrOOlIC~-d '.0 ()i:illty allJo~l, ~56" of hilU!li1 $1)ltCI::. Uti dle' basis oftllC dam preseut~'(llu .. ,am! ff

or !Ill!!:dghI

or Oi:JI1'csponclung

~tukm")''1\

[I,

Ir ailt.;

.iI,

Cli. (lJ" a OOitCI'l:UI rl.lli.C~ ~iollml ibrlhc 0::: i~ rolc e~n.crgillg;. In ,l)ln~mlJ!.I;I(l.
li'CI,),;:mpl!pe:r by l.iLI
(f'!

CPLII
of

m::lImn~

do Libra l.lCj~ llillns

[he CBU

t:mge:nI1111 lIIetl ro:t18 .itlUlC'J'Valing ~spm::ifie hl)"'C1N of the CC"JlItm'I lx)t])1 :life Ciit'>C!nli:iI IQr roC0£il1i~jllg fualtl~ of \!I:>ImJ{}'Iljcr:[~(d!J\!:uu.ion nil 1~1tlpmm· I!<mm: CotU.oLlf urientatinn]. C{]lIUlliUll~J[ neumns. like the C~Ul ueurcas, OIl'('; ideal c.;Jndiil,mJI.e;s; 10
Wl'iocia1C' n~.csa \'1SUa~ fl;'.;;lJ.llm'i with in:l'l/lI'I!llil~rDn

Ill' I.,1IIi!i

,~ I:'
..
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011)IJ~ir :iltiml;llilml dk>C~tit:m (lig- Sot)_ :UU (7 aJ'. U _?) m111W.:B9yhYJl>tJlhcs.i~-d. Ihat~hC" wicll.1fr of 11m CBU rcr!l'!.,'1ICnl"~zin1.'Ulha~ space. Thi~ fu" H1:l\llll~~y ,..;upa)Oi1cd by Ci~tr d:II~" I.ml. til is: :r~ln;!£lCil1.t~]ion !!Il~ dilf-cr bL"i\vQ()1l eell typ;.,""fi. and CC ~~lh~'ll",IC',· tLl~~ ",,0; PI"Ilutl:ld out ~ho\'u.
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:liil!J~'ffiI

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Thle ,Ca'IYP,tog,e'n'IfJ'm,agnifi'c',iI ChelmO,8utotr'opihiic Symbiionl GlenOIDle


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essential 100INQv~:;;inliiii1g
:;;y,ll1Ibio~is" occur

~)r ;uuil'llJ

acids UI1Ihe
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amlfllonia

w~lidl cuter

1.l1C1ell \'i:1 c

lIIi~rUllCl Ir:JIl~'1I}OIit'Q!r~nd two ,mmu:mil!llll] permenses, rLl'C reduced via 11.~Irn~end nitrile a

(lternJo1!lul.otmp Ili( end o.~Yl1llh~OrIt~, ,1Inelll,e m.Ha Do~l( (,OiflllentOile Clf h,yitlroth~rm1!l velll'!: >c:orlllill!llllltl;e:S" ,pr,ov~.minglin'l.l'@rl;e~lr<il'e n(Ji.st~ '.'Iilil near[y aU or~11~h Ill!i ~riljol1i ,. lihe Ca.IYpt,ogt'no mG9'fl',ificff (~illaLviia;Vesi(0Illiyi~<le~ 5y,mll~onl. Cal1dfd,Uu~Ruthia ma!7trif,~jj,ls tn~'Un,t rn~ra(e~LLlII<lr :s-u:lr~~"o:t:id~ziIll9 '~iild!J"'fmb~onlll!l haveilS gel110r1lie 5i!G11J~I1!~etl,f~lJ\e'<i Ungl a ~'UI~~E! of melaio.oli,[; 'L~p'~bJI:ili~. iI"lTie [jeillome '~I1(jjdef; major (hem~a~d.tJt:r,oplTi'i( p~ll'iway~i'i~ ~\'@U as paIihJ.~ysrjjr Dio~yll'UI1~:5ftsor vila_millis" >oo~aict(lr:s, nd flU 2:0 amino ad~s required a by tlfi@ (I{lm.
1

J;'Cllll.c~a"'\;C<lml m;:[oiin1il~ull(i 'V.i[l .~Ul[a t11 lne 8y(n~h.e. rase- and g;lumm,ti~c l~ymJ'lase... ~'Spq:li\'dy (.[i,g_ SI},. jid:llmllgh nitl'<ilI1;t iisthe dOllllimm[ ninog.c'l] lil'l]11I ,~~ 'vcm~ t Hh tim s,:\''Illh~mu !lllly a!oo as:;,ful~ijlalll)!' aJTImnnia ,lI11illlO

,,'ia ~".Cj'C~~.Plgor th~ hO$;['\1

,C'!aZQr:n~~;Ill deep-sea !lydm~~U~tf'll;l! vents ~l!Clu d2l.Il w~lhlhe SUPtmrt or _. _ synnblot:ic (:h()nmilu~nlmJlhic h~lwi1l! {.f}. An,,!ogfll;L~ !O phot!l):;;Yllth~Dic !:;h~ornp!~!'i!:S. whi,c;h '1.'VO~w~ fj;cilri ";J'fJlIJobH'Ci.crifll ,~ln~~::;;Iur;i J.ilid li:i,.; ~li,gli1~ ito;;rg;.' h) fix t:11mC!ll fOr lliii:iJt r~m~1 C

!)!,C'llOnl(l encodes 'r:!i112yl'l1i~,

ilncllllldil1g

,II ~(lml

~:ri>l)dfk l:o~hil~ c!;'dC'. II II'iblll'I'l:ol'C i,5.!lJi:,;phrnph[Jj~ic (RuB


i~CD}

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Qi~

acid WOl:<:!C 'Uul ilke any CI'~:i1£'r1(l{IUe-I1.'01Jd l Clld!i)syu~hiontgLCmml(l. tiL m(1:.'~}j~lic~J I1l;:OO.C:.1 C comr~kl~epnth~~'~ysrQl' the biosyn!.hc~k<: or 20 amino ndcls., indtldi!!!l. 9 il;!.~ntbl mil inn ackls on ~lCj Ii PIfCClIU':OOr5 Iill,!;. muglllnw fli:5.0
lRl~ eOinph.)~..: b~o:;yrll'hC>1ic paillway.o: for Ihe nli>ljority (]I ",ii~illl~iJ'I:; mIt! COtfl(;IUt:o;(SO/lij text), T1i~ gelHj1llilC cneodes a ool:ii:rlel~ glyoolytic p::nhw~w ami ~h~ uUI:KI.;,>;idmiv,;:[Jl'lmdl oW III..: pcm()i.'lI.! pliiU.~TlllaK: j»uhway ~u1lcl. ~rIlCrn>lliulgliy, also encodes <I.lr.i.earinol'l},h,~ acid {Ie M c yclc lackillig a.kctoghu.arut~ (lchydrogClfIilSC. The Ia~<k ~Iili;s e]\io/U1C has been $ug~':;I.oo to il:ldl,Cllt';; ~ilb'lilg1l:t..: ll!l!tOlLuphyin olller b:8~~K;riu ,t I ,t~. C~j[Oon fixed viII eill:Cti~\lin cycle can Chli~r !.11n;TeA t~~d~i! Ihm!ll~J'i plJ.{~ho"'I'io.lpyruV:Jh'; Jllcl~lC~ ~1~II~d rollow baffiyn •.I:M;lIk; .mlllies c'i~hel' !i) fUtl~;;II""1.C 1.0 Q."kil'01ilh,lt~II1lIIR or I\~ wilh rnher: illllrno;;dh.ll~r Sflccics, g<:'IlCii not blJld ill .he R. m(lguitrl..Yl gCL'lCD1D.;': l'iC"l~.1Iim-

s.n

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3ndm,iWLI hlJl>~. CttCll1t'1fHI~t1\1't1p1i it erulosynn,. himm.; use U'l~ 'C:~k~I'I1ic8.~ CUICrgy uof lPCilm;,-d Mil rUJ

enl!IIIllU iJrlg rrnl~v.cm;,;tI) 'pro\'~d~ lh6r hosts W.Ud3 carbon fmtd :I ku'g~ Ilrruy of nddi~,itm:li nee ... "sary lillllrl":ilL::-. l[u n:HIIil1, busi. irlvertcbml~$ iJridgc the o:o<i,Q.,~~lo;o;;i..::]]ll.,"'rlil4.'l1.; pm'\;' ide :;;yntbi~)l.ic ~:mcl,"'" i 10 'rri:tl w.ith ~hN.:illmy1mic l'lIb:simLiC$ n.(XI,.';l;<:;1!u:y ~o:r
dl~lil'IDIlIU~OiLro~ly,The

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or

~imi'~ cI~llii, ('a~'PI,tJg("Jm Im~fl{lim Boss mid Turner (B:i,v~ll via: Vi.:'S,i,eo. m:.y ,[)xiidil": i~$ MJ,lJi.!Ir g:mllUh~,""S'Ii:.!. mil" hmml'IInyid::h.:). '\lH,,> ,~n~ or eh!! fln:5~ o~rmi5U"~ de:!i.e-rilled .. Her the J ilj!CO\'~J}' of h}'idrm HU~rtltl~ 11r.i'~ whcnl CXI!Cf!ml s;~IU'idc is l!ac)ldl1._g" as OC'c~n in hrl!~h(WrjcJ:tmm(~amfj I'fnn'i'um rind v~n~s a;.~[ llol~a rcd~oc'Ci:gut <lind cil i;~r)lfood ,('htomMm,11 NmirrjJ,~ j 7" .'if),. IIQmo~o.fl1~ CIl.,groove lInd Ls n~1!i1i.(ln~.~~y dt;!p~ndCill.!l'lni!.~ r
1'~P'ft1~i~Iil"~I~ri.i:i! ,~yn~bi[m!$ (Ble~ 11;111Nd (_'~m(li~t{IiM~ RutJH(i NJ."gji~l~c(j) V, 4, ,. 'Ij,!'/c pfC'SlCilllI thC' colllpiclC ,!!,leoomcor Rluhhj m,~gtjHk~ ~F~l:l.~J. n~$lliw ~Kl"'~llga rcbti.vely ~m<lll.ge~O!!~~~~

:;;y~jfu~:nt. F~w~ll~~r,rI(;-j,TiIQ~Uk t"'i1Sp1,1!1~ hL1!~ .~. S\II U!dclQl1Iil1n~()' oxidnood"~II;:I""il~ <111(1 ~'!I-':I~ fourld. 9,JIggc-l'ilj~.,g~i~~ii!he ~J·mhjrnr[:o;..we '111~D r:ilOf:hmcsc lIiJi'lC' al~ 1}n:::o1Ci1~i, ,,\!iihU'lcd,~.r .~nd, '~H!d :!o'~JX p~o!d!'l;~" !hc:il~ em',YIf!l~~ ~ai! oOxi>di~C' k<Jky 61'~~!<U tl~ ~)Gt.<,t aClivel)' digcll~~ -"y~nhi.{'i:nt hl(llccd. Rwhlti'~ dmcst k~!OWlil rcl.uh'Cs. iHd !ide' or thins.ul rmJ!iC' o ~l!!1 ~lw (f~g. S I). s,,11~f1,tC' t

Il(m;lDl~ d~~aiL<;ah{l!UL Illc

ill~~m'C!i(nn 1~[w~11

10il

(1.2 };1M, R. fNuf;!NltlflU i~ l'IOOdic!C'd 10 encode (Ill !lire' l1il~:~hol i'c 1ll00~BI\"'.~Jst},pi.C<l1 or n't'C-lh'illg chcnlooll~olropll$. i~llKliHg 'cmbm~ n;:o,:m km, S\M~ir o::dcbdon, nil~'(I~n ;;-~slm~i~~i(m. ilnJ ;)mi!!lo~,.dd ..~J Cfl~;;,cl"(l!r/vit~rln!fI b1(l!sj'll!hcshqlig., SI :md~blC' sn.
Iilfll'llC

'[he Ihlltliwiul,t M.lClitIDlS, ,~jUlI~~lle.~IC~'C\m)o 1 ;:;In.Jceinn of R. INClfjllijil':rJ'S ~ho.::ll~mllllo~ruphic .m~1!ltmJ,iiim and wl..al. lh~::.; l1Ilighl .. ml"::JU (ot Hn: bi.o!()ID' ofil1> htl.'<t. Ollt inaldysi~ P~"~l~lid~ dirc.cl 1C'o'idClitco;;L~1lI1IJliIS ~ymbiom 111\,,:; C[lrOOrll "iu '~lr {,uh'~ III cyock. 1 he ,[km~ilJ:JmIbnn G f l::JIrOOlllil)m." lion in\ <!.~IH:;')1 ~nhiir.!li!..--:;~.'I}, hy 1I ~in!!!.~1.cI:gY i[1c1J'l,,'~d.I'mi'll slIllf'lI[, '~K id[rLIOI:i, The R. II1{JgrliJi(_~j
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Qlui.tm!1C in

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O:;pF o mlll.,;l}; Ed. 3n~ IlUCHlIate,'; :in,:o; p~to.'f.Ihorylln ion d~e tl IprO'l~m (.(Im:oll1litanl \\li:thdeanlle ·01' the ,r;:,·,op ba'liId illl the Ipho'sph.'Othreonltle residae, lnhis SlIIIIII:;;, ufl.El'Kpedee em;yrmatk acti"ity, ~emn;ed Il·trlo,~phot~ roeQnnn Ilye~Ie,.a,ppea,redi splllCific ~orr MA PKs ilIIlild e G i\ 1.::11 ih~ dim:\,"C! ;~~OCiilljOJlo OCtW\.1C,h O:;;pF \\las!iha~El'd by other Ospf [Ianll'ly m,embe~s;, ,1110Erk2 3I1d~I1,I' ~hC'~~!1,1i1 n]n1.I!l1 ,ofplll(k1p1H,1. a ErliJ.t2 (jp·Erkll2l' wa~ dimi!!ii':hcd 1:1)1 n"pF nw.1'I-\lt..:g~l.li'1''': lbaCll..TUal 'f?lItlrlo~llIs. QfLl.:'U I).:gul~.I!!>t! ki nas..:Ji, I amI:1 {Errk i11} mtd c-J ulll addi ~:iou iutile cell=[l'o('( fl;OO~~nj!J!,utio~ ,~!lS<lY I u:;;e I.ba type.~ ~~'C~:ct~Pfl. s.!1I~eLn~rrss ~ N -tenn hial k~l1.asc ON K} ~ign.l~ il1.~. \.,'hlCll1:l<h.<;.it II (fig_ 21\ ~ we ~(":lwncd uh:n O:.>pF oolll'd h~lf)()r hJ ~inj~ into,I:JiIJ:;;I.cell~:;;effector g111l,h;::i~~ did 11.0t illht"~ NFKB .:1IC:li,!,lat~km (Fil!} ~i\ 1- n..,pf' (l} that i~~tl;:rfct'e, wi~h hfll~t ~iglwl tr<1:ll;i;(l\.IC1irm abmfli<LlCd Il~ln!':p!lOry InU.nn of endngeno us .. 1~11.o!Opl1:lb~'l'i:C Otc.i·vi~y 100 l'Cw!'St'E!'k rhos· .ErkHl, JNK, aud p3K kLml!.~ «~l~. I,B ,11!1dn. plilfjorylm.~n'.Il_ W h~u ln V'itffl l~ho.:;.~~ho!I.'yla~lX~ Im~bwa)f~ l'll:ODll(lw.p<:llhD.~en hnfct!ion. MAJ'K In (iST-Ef!4. JNK. ;:if\OpJ.~ w~ ~1~c~uh~1,Q~ i~~ w ~~ID'.;I~ ph)'~ ~Dl h~pt'lJl~n!. !'(lI!Chli;i~~ i",~th~g i~g lh~ n;:.gtll~,"~ $~IWjit'h~~ O!3JIP hllrl~-:l~;;i! ~dfk ~pmbiJ~;lI~1 o~pr? dcphn~I~ll>f1ly~1l1k~~! irl'l;l~IX~ ;~C!j\,j~ dM1>'!HC!C!}11 of ~lii,~g !1ll.l1liplc MAPKs hLlt hl;)Sl ~~O<Iltf)~.nlHu.llue OOSrolL'l~ [I} boll'! plants and (l!Cc!'WI\~U (F~g_ 2, D .mdE~. F~!1!hcf:!l!ore. OspF $ 0 <lUi!HOil!1mJ11disa freqlllcul. t;wJci. 'L'lf ~'I,"lh(i~C'nk !lOI N F1ii:13. igJ~b'lH!lg '~"~'1!i;11 \ C'!'C'.'<,Pl'C.'-llOO in mnn~cJToctrml. (1 4). ~dC'nlil:lcmirm hns, !:[u'~c,s fir m~~i;incel~. To delin(!~~le(o:,1)OO~nC ~cp(s) ill !he- ~ke~ilV'C'~Y !:'e'lllCWOO l'h6'$:Jl'I]l.l~C ~!\lUlloll rrotn !he n:ss ;):f!Jd cluckLlt!r1l1o thC'i,r biflb ;VI/WK pa!hwJY bkJ!Ckcd by Ol1Pf':. wCI~lmcd to ~h~eollhM;! ll!! no·t frfiln Ihe-I!}'m$~~' residue il! !he chemleal rllll1'CIOClIlS are cri~~cal~n 1IIldCJ1'l'lMIi~utiing the ·(;<Ul(llbI:cOlI Erk Il<1l1'1\.VIIY tl1;;1:t is: nc!i \il)hX~ by 1lI pT-X-pY U 3) m(lt~fin p-El1k2 Wig. 1, D and F). Thus, 05PF h~;rbm5 an in \'i!.1:OO eliRYmmlt<lcl:i\,themcchil!1i~ltl and tl~e C\ ohltiij)l~ of 1"1c1'C[ia~ IllmslP~!QI)'~<l!io'n caseade of Rill: MtWK kiOOSlC' i~y 'U r $pccific!~II)' I\:ll~oyi:l]g pI]~llt.il~,,; ~TOLiJ~::; (MEKJ'. ~!1d Erii ki~a"l."$ OJt Os,rfblo('ki;!dF.rk 'l~mho~C[lesi~ p, ), 6 J. pT-X.p\f ult.14.if il'Ci.i,·ai:ittl!uin 293T ·edll:; IniJlSl'..:ctcd with cil.lricr lro:rn tl;,,; ~ll'tUlip'htllhn.ullill";: in ·~h,.; 'fhl.: ITS'S ..:fif,.."t'lor Oli31IF ~~p;rcscliE irl11M R;I,"'V~2, .....R:J1i' ~c.·ol~:'lillllive ~(;'Li,~ Rid), or '[hI.: four kUOWIlIpilIlJ1~l(;il ie sp(:ci,,""S 0 r Sh.r~JJii illl MA:!? OC:s. ~Ijc:rtld iUi!: Erik Ill,. JNK, =mt! pJ8:. UillF m:P!)~tlruI!I:~ fUI1CI i:uu IIi; II Uln::ouliUlI:· IFi£;. {7-9)< :J ·t:IILL';!Uli 1I!!~m ![;If IYm:ullm)' dr!;(.~ih:lJ)' MEKJ -!ED {CtlfJ~E~Lulliwm.c!i,v~ MEKl) .....: I Dl, McrlIllwhiiII;!.R::t~V~l-il1(hic(!{! MEIK.II p~lO::;~ ~c:jl:I~C MArKph(:H'~)il~rnsc. ~~IL1Wi.!'V~r. ~m). O:>pf. ~(j~lhcr wilh Sp',( mltl li()pA.lt. ourmndy.i,i£;. rurt~"O!Jt 111]',,; as5~bilil}' p of;) {:J~,i!l:a] piI,u!";'in reul!llil'LiXi. iilLUC~,tk;spi[1.: l~!~ t1i:>t!P'" R.ll1C.'>C1l1lS :J Yam.ily ofTTSS cfl\;;clul1S ~"OIl:>~,[vccl plml)'l.lt:ioll, in. oothp~;nllnnd un iHnn~ Imctcri<l~ plllhogcl~ I( i!) pIIHJ. :phmill>C of O:spf i!i Uall!ln: l'Sulpprmi:ng .on... pCllffin(lC of Ed; III p1101f.Ihoryhnirm in cdl" liute M~H .. (Sm• r"_"iI>t~.'To nc'\ol(,,"al ulilc Ghi:nltc~:il ~i:ll, ." GspJ' (I~~. I E). 'flull'., in~]ihilnllrl oJ {Hg,. S 1,. None 01.'i.lI:: O~:JlfIlui ~r l ..:flix:lom; ~m:;, C1\IIF(;;:;~~lJg Ed fJ:rlU."llh.Qryhu~imll Iby Oa;lJF .i~ dm'!'n:;1.Dt'i1II1~ Qif IlIiIUlII'C ~)fO:;pJF ~';;liv:illy. Wl: ~\lb.il,.'1L"[..;d MEK1<Ill C~~llblii'Jl{;d hio]'[)~ilt~] 01" hii(x:hl,.n'licall rllncEll,m$ipho;ybIL\j.~, M~I~-Erk2 ~p':IEn..:1C'0l11ro11)• a8 . linn. To dt;[jll~ the biod,'l!Jmi~al .t1.1l1c~ion fOsp.f o MB:(~ .;'IiI:lJlg the cla."iSic:'d~ Ilt:a:/i-RaJ-MEK-.EIk 'Wd~ .:iI!i. M.BP p-Erlk2 rlln11:I~r Ir:car..::d \...olh O:;;pF alld 'Iii,.; OsrF 131l~i of crui~wrs a~ wdl a;;; 10 c!1scmk. ly Edd/2 phm;:p.l~(lryJ3Jti~m lJ;{o; r~oon~ljTLJlOO '[p",E~k2 asp.f), w [31ttL~mllml15'~ ~pi.-'l':I:r(!ll1~L!)' c:m idclllti:l.Y dlcir I'KIl!r:lnli~] jn, .....\/oi:l l::ar.~ct, \'o'e ~i .., ::nnin~dI..:m~ct~ OspF on 'the .host ~Ulllllll.mt~ sigl. of illll ~hc cdl-froc c1>ltmC!lS .... sl!J'Jl1 by adding !JIll" aa:ll)'l1.i!l. Mor:e Ih~l ~arw. niF !hC' Map- EI'k2 ~y qllC!1CC ''''a~ oo\'crod by oh::1C1"'1!C'(i iandC1M pe.piSnrt"am ~.iglmlill!l ]1lI0lQC'IIIC;.>;i("~}_ Additioll·(J:fbm::.rmljl1!l p::uhwaJ'K. im::ludull.g D]1\It;C o.f n~tdc;llr wi!'Ll~Iy e){1~!1C5~(l.mfld high~}' '~lIl'i IJed DspF (rl,g. ~kle.... AM of Ihe l:IoCp!i:d'l\'i.[illl,1m 1:I-Erk2 cfI!1'ro:i1 I:~c'orIdJ (N f"U} mId MAf«'s.

The Phesphothrecnlae Lyase A,c:li:vity of a Ba'ct!e!rial Typ'e 11111E.fl:lector Familly I

.G

cn~~,~

or

or

"C'n~ {!m(;iCl~llyb

i'I:gI!lStcUl

~""'11ri\:'.s::;io;n {lfOsrf ~Il hU!n[)1l

1000

16 FEBRUARY

2007

VOL 315

SClIIENIOE

\IV'IoMIV.sdencemag.orgl

REFORTS
mOli:!' W~, JH~. CoJ'lSiSK·f1t~Y.' wh.eul {I s)'tl'lhctic pltosphuP'CP4 ide OOJlliliiljml~~hC" TXlJ'ph()sp~lo. ii1lOlif in Errk2 ~DIITQ,i'L-~H=E-pY-VArR) wss Ifnolii~~'\1 by Ql;p f flrKi ~~'IC ,,-estill tlrlg P:J'O(JL~'
;t1l1J1Y,ol.l(j by Il;W:';S s~lrollflC1I)',i!t, mass d:(ICi-Cll~i: or 'g~dalll:lnml 'lIb!: plm2l.~ll1I'Lhml[lllllil!le n:'J";0CIL1~

oompm:cd

1111C ooul~dpcl'~id\(!o !N~IS;, 11100 In, 11l1.HS, l!~lOJlI (Jsp.f lOCi!i!:rllei'l~. til,;; pllOSj:lllmhroon hte residue iu .hC' 1,){Y ntmif ill b(l~i"lP'"Erk Dud '~lcrh.():>phO~l!i:p~idc !IIbs~~e $ IJIIK1~r""''\':IlI,t~, d:chy<1nllliOil I"CJCltlfm t I~ df]I~(ll1, Cit
iiI,' itil

grol1!P ,(80 d.a:llOiU, 1]\i:rerrotiOtl ,ofp,-Eitl

~!;')~~

ob:>cfl."C\(! (Fig,

OspF probably ,rC&lI~IC"(]J in il C=C' rond OC'LWCCllll q~and Ca in IJtC side cllmii1. or lhel~ll1J."Ouiuc Ii'CSl'tlU'; ~. JD),~"Llggl.'Sttng thm Q;pf fL~I1;\;'ltlnS W as <1 ~Jho.~llI]mhJiCOGI~UC lyase d,;'it cleaves the
CoOP homl wllh I.im ccmr.03l1llrlm.lit ahRIr.tel[o!1 of i['~ 1)1'Q~Oll ill lim IJ1h:rulJ1iml:hracminc resldue in 1~·Erk2_.F'UrnHimnunu,thi}lL;~l~mkJllm:!(H ion licm or r:p·Erk2 ru!S!lII~lhlg rrollil OspF Ilho:o;pho'Lhrt'll]> nine !ya:;;:c 'IC~i\.Uly It'd, 10,1 OOllllpk:h;!y irl<mth:c: ki!lllivt: bmh in vilfO and irl cells (~Ig_ 53), In] ..dd~~ion. O:'Opl~ ,~·d!Cpho~pllOT}'~rI.t~·'E[k::l oo~ll~ !OO 1001g~ be rnphmpho.yl ~tcd hy M£IK l·.~D ,(rug_ S4 ),.TlhuIS,{hpF r~mC'liol1 .~.~ 'p~l(k'iph.{l" s d~~COll!hl'i'i '~~\a~c.':!!~hi:!I'Dl~UI~. [D~lf.I, ~I!IllI.l:'OC', ~ l ...
dlHJ

mass

IMS)

in adeliR'! m~

'to~Hic .I'IES or

a [Iltm;;p.h[!l,l,i::"

F~'£I. 1,. ~n trri~iti®1'I or imU'[·' lii P I,@ Wi~A K p:!lllil W;:l~,~ ~ dm'i'i1s!Jeallll <!!r MAPK ki~ iFla~ IJ~ O~~F'~n,29'.31(e~~, fA) ll.ldf~r~e .a~~a~ Ol~' ,~'Jfi!!(ts (,'If O.spi= ~X:PJ~;ssi~n (11'11 E~kll2. ]INK. alndl fI\I ~"B ~<!~il'ill'<!Ys;. IEfkll2 <! nd JINK .ac:tfiilf.at[>o\M\\iere'dJ~hieljJ~d by CioQ'~'lJfessiklflo~ (ooslil'lUltw actn'!/!e Ra:sVll1i3imd RacUll rie'spe<otiwly,<lllId rI\I ~,,'EI 1.l.i'!I$ st,mluliilleil! by '~tln1{l:r !1@CfOlSis 'f:~d(i:r (i 'lreatm,Elnt. V,, f, .a nd J denole Ve(!tOf, OsplF,aJllId Yap], r\fspecthrel~, Eir'rolr b'i!rsilfldi~le !standard de='It~ti!]\n" {B and () PhM.Pll:o~ il:pedfk i;lI1itlillilnlo!blfrU:;FIig ,a"l$:al~'" or linni,Mliotl o~ MAP~ acllv<!llQJ'Iby' O~!F, Ii!-&k, p;JINIt o1I!1id~iP3;S

A,

,I

Ie

ivruvc~iil~y ln~cthmIK': MA PK~'ro g;tin fUli!illC"f ~mdC'!~;uld UI]g or the {l~l~l~


FI;!_g·I9IA~SpiIi' RI!l~L!l,1

Iy.l~ aCli.viDy. WI;~mUMIlt(:d 10 served in <!ill tl~c Ospf :1~rIi111y IIIiClllil~r.; (:(lg. :5)::;,.<\.). These nlllHllll!S hmj, :>ili.lilillf Co'i.p.-c.-;:;,iim mid pun nCij~ion profilio!.~~o iho.i'1,; uf UI,iC'''' i l,a":LY~IflIWlJlcill~, :;u~,ailil;j !Iilu. lIIiey \\,'1..'11\:
rrcS~dIJC8, COli

pllmspllrn~l~l!illH~

lul(kd

;:;,mrecUy. MOOU.1,fThc

I11IJLi:lIH.l.I'Ctfunc()

Ih<.:

ID

a nrlbodie$ recognize- the

dual IPh:~1'n(l:rylLa'l€dl iXY motif (1l1wp¥)' Wi II:rKlI'2, JINKm~ and 11).38, r~~i:~ 'lhre\ly" IH\.A, hllFllla9<'9~ulifnm.
(II)}

lu)(it~r ase .ass~ys of

!nd!.!c:eg E~k~ctiJt~tioo ~n t~~ p!~S~f1;~e Qf Os.:~F,"!'-R<!f arid MERl ED olI~ ~@!1.!,lhrti!{~<l!l;Ii!;!l;l "!'mii'i:!!1~~Qr ~f a fir! MEJKL ~rro:r bars ind,cate~lalrHlafd devi.ttio:f1l. '~IE) BlI1M~h~poedficimmLinotll~Mliii'l9101' .a(~.atiOJ'i ~f .MEIKl! (ll'Id ,EnkW int th~ pn~5!i!iIlOOQ~ OsptF,
Fig,. ,2.. lmm ullflol!Jr!J~tiIll9 !lIneally· :ses, ,cd remo\la~ o~ p ho:spi"late 'grDup:s.rrom M., ]N&(,and ~3B

RiJ,~·, Ra[·, and MEKl~

Erk1 rohospJwthroolllinc l~i~sl! ,:!Le~ivily,_w~crea5i irl1.u1:11JI1S K ua_,·R'un (I< I. 02R." I.I.I~-A 1041 011 04AI. and KII~-A I:M (KJ J4A) were IflfgC. Iy ~MC'LIV~ Uig, S5, A. f11:ni B),. These InLJI[nlt8 !,;t1\ll~d:>~i ; biml L1:IEdk {fug, S2, A.lfIilrl ("~ :-tlg· n g(!5cin,g II i.UIlCl:iQ!lWI:i :inilpOJt:m(% [lor K ~O~, HI 04, rnlid K.l34. A neurad c.uacth'c pllJl:;p~Il.l.,a5e~:-;5ay ~\'Ol" u~d In ql,l!!n~i~mi ~'Cly mca.;;urcoC'1i1(l (,'Ill:tynmt" ic ac[i~lit)' or O)1PF. O~IPF drH;knl~.ym~n['woo pllttl~phlllc gmt;1 liS I'l1(llll the Erll::2rimllpl1npq)ti(1c witl~ .. M i~1Otc] is;-jV!cIl!OIl con:;t~ll1lKi1~) ob"95.:i, + 1.4 J.lM and, a (:ai"ly~ic rnlc 'C'OTlst<lll~(/':~lill of 0,70 ~ <I;tM !'.,-I (~;l~I~ si ~T.~~ K IU2",.1 ~.~.04A,

or

and K L14A 1I1UI(lillt~~\'(:~ i~mclh~ i~~hcIliI;lptideOIAA,T)"


l~lf1re.~h~c !llU'ta!H::1 \:v~'C

{:n~-S5(,~.Fl!!1hcr-

"~U11~~

rontp!C'1'C:~Y

alsn deficient b~

~-

dllMI'!l-lXg\4I;1~i~~g ;;"II!~PK sigl)lI~hJ.\l \\'hcll exI)rcsse(~ in 2~)3T cells (f~g.. Sill,n~:;;, K 102,

H UH. and K U4! are


MA:I?'K~,.

j'I!'(IlJ.)

bly esscnti ~I, for

o."pF' pho:.iTlhoUnx:~mil'l~ 'lylJ..';c ilC'h"uly Imllmu


We U:icri nl..c"o;.Si.'[] TI.'i!:: rt1~!.: or lhe ~lhtl5"pJIiI'C "

~nRilSe'S

.~

OspFilr1i

En~(~~ ,of O~p~ !J'1l! tIFkV2

vitro.

(A):

actival'iio>r1 reQOIfl5tUlll!~E!cIirn~he

oe(ltl·free

1l,·lf!af pm~elil1 \l{~~ added to ",ctiv~t~ Em'k ~'~hl,\'ffly illl tih;e pre~fl.(,e or abselllc! m Ofl,p~, ~B) IEH!fClS ,of" Ocsp!F ,olflim 'lIi~m reoo i'lSliitll~ed ErlI::2 pho!S;phoryt·
~~i(lr]J dl;lwl'tStr~1jI1II! ~~.

e-:<l.iralct ~lem, Active

m~m) 1!iiI,

Osp:~ ~,.JNK .liNK ••• '

(m the l.ylU s IIIC in 'pT•.X"~Y in ll'ic MAPK liltoJ;. pllKJlhro[lnilllo! 1)'~li:<c aetivi ~yorO:.pE \i'\'ta.euErk2 '(.pT.X.pY) was tyro:s:i nD-ooph.u;,piwI}' I:nc:d by p:wDcin Iyroi;~n;;J1h.oJiipl:l<Il<lsc: (I-lcPH). subseqLEJn &.~ph{)sphu.L")II~.Dion O~pF ,",',m, LI.';:\:> by

ct·

~t Kl

Ufe:ctfl ,cd 05JllF

'0111

~llor1Aatj(!!n fly MEKl i!'lvilro.. ~D'JS,eilectilvof femM'al of phesI'!h;;r~ '!Jrotlp:s f~~1II! tli!,~ 1j;lIiJOS:pllotBl reamin~ nlsidue ii1i lne

E.rk2 phes-

l1!:.<il."JI't~Ug. S 7)\ ('on... il>[~'I1;L~y. ineie IIlmly.si~tl:;iJl:g k the :;ymhcl tc ~hooO'Jlillc-n~Q.lTIlJph~)h orylatcd Erk1 p!.~p~de .!fI'\,'~ 11 i ~1i,~ncrKm li'alULC~'Oml~l]n\.-dI wilih ch;tl ol'lh~ dual nlholl~lhcLylac.llr::i Erk2f1!C'p. tide (table Sl. ).,Thus, ~ht1pholilh~H: on IJ~'lyra· S~ll!! ~Il ~he sul'l$~r~l1.e Ilml).!!bly co:ntril'l'l,lIIcs·to
~-~bl'l;S1rn~C rocog.~it~~mb)i

Os WI? MEI('~ IE:D 'oil!GS.T.Ert&l .......__

III ·)\·pY m!olH l!'l Efik2i!'l r

Vi~I'OI,

... ."".::.~. .:..-~..:...., ~~

'G:51-p·l::rfc2 was UBMe,11 'O~p!r air MA~t( p,tm.~h,at.l:le' 3 (MIKP3), Ioitowed by p~~r:k <!Iild
oil

,mUl

lP"Erk2

====~
05p~Finl

031~F. flUdlC4" killctks snalys:is l!si!l~ dJl1:"c,cm rcp~:i,de ~~lhS!m'LCS sl1,(f~V(dhat ()spf \\r~ sp..~if!c wr~her~'l~P!l(Jt !)h~cflnhlC, Ul the c'JJssi<C;l1~~\PKs twil~1 TXV nl10ui 0, withlhel~i~1t'~\ enz>,mru~i.c:-lCl i.... i~y t(l~\'a 1.'(11l3:~i~~bI!e ~). S
SprC,:! u'l'CmilcrQr!hC" Ospf {hl'l1i1yomsS
U~01~'}'JjillOj:d tib.ll{m~!tt! ~'>tL"~i S ns.,

ptJ Os:phol~Qsine-sl,pe<ijfk an-

'~lIJM~ (p-1iyrr)' ~mmufl()bloUlft9l. (In Re-rrmi;ia~.of tine plnosplila le '9Ifeup~ I1FOIfl po) NK arHjl P·fl3B s~ vitro, '(f} Il?'hOllllho~r'()si neiml1fuJlllobtottii!'l!j 'Ilf My,c.tErk2 If!l{ld~lfliedb~ OspF in 29311 ~eUs,

cliFc;:::~orsfrom

is

t'Cl.Wirod lor fi.~~~ina ICiKIe ~md sY50cmic iUllcc~i(~~ v

1001

REPORTS
~~~@~hu: ly{!~ ~diivitii .of Os:piF. 'A) The mM~ :s,~clr@m~tljl' ,<lliIiI'ly.si~ 'I)r tile molecLllar mass ,(If (VADPI)H DHiGH-pT -i~p¥-VATl!:.l' h~11T1 OspF''lreatetl ~Hirt2. I[)frIW,elE!tll

Fi~. .3. The lJinospho-

I~
-

~r-------~----~~~.~I~,~~·~·~~~
2ilii ~ "":cd

tl1ellFypti

c peptide

I~

11~1l---~~~"'l'if!I'I"ETc"""~...,.II....dO~-:OfIPF~'
O!m , 111"1.' \.'

11,1.11'51

ru

pe-ak'S(()rresj)O!ld ~oijhe
lS(!~opic. peaiks ,o'~' ~Ih,e peptide. (S) O~drQ:s:pra yiQIl~~~ltivn (IES~) lalldem ma,:iS spedmmr 'el!r~' (MSlMS) '>Il'f(~nlii'd
rl I~~ f111~1 I!1L.

Qf tir1~ t!Jptk ~~'~lid~


'Sin.<Jwn 111 (Ak b<lIIl1~y i(!ilils ar@ m<lrkedillllihe :s:pe:(:trlllm. lI"lile' oil m~ITIOI

.ad~ seqUl'ence ,o'~ tile


pl'lpnd~i~
~il't;IWlfl

left, and Til marhllhe

jj1lkiiP

OSlpf·modl~ ned phospl1,ofhreo;nlllil~ ~es.ki!lle. "I'1iJ~fr;P!ilme!'l~~il;ll1 ~lc

'lern$

th<llt

9Eme'~tJ!l ·lthe

'CH3 I
I~ •

OH13, II

lillils~mtll I!l anti 'tIDffi .are lID:l&trate(ij a [1)11 g~'he pel'ltid~ S;~,U1l1nll;;:t:, (IC,) m5mS ~pec:lJrum Ol~ lihe 'OiS~F-liIlodified S1jJ'llhetli:: Erik,2 p~n®s,ph,oll~pllide ~IDIFI1GFl-pT-E r pY.~\lAlrRl MOltaiinOlU are tllie same 0115d~cribetl ln (B). ~nIe~Sh01.llSthe motemlar mass ,c.:I Ule ~lrk2 phOOfJnop~p'~lictif! i:l~ror~ (CiirfJtr(J1) ;plid ;;I1~~r(O.spF}~r~bn't!mit~ i!tu", a rlb'itra iY I.iIfI~~> ~D'~magra m ohhe cllernic:al. alllfl' ai~lVfl,or Hte!'philSpIilOlJn.re®·

_ii,.""IH_'f~~OC-H'-' ~O~\S~PF~~. .••. · _J~. '.' ....


Cr;C-~1 n

",-NH-C-,B-NH-

~HI

n~ne reStdLte!in I:!rk2 by Osp:F. 05pF deaY\e5 the (-op bOIDd !Nitti ihe ~on~l)lIIiilAlotP,Dst~ctgOIiiillln~ a pt()tllli'l, ~~nti;lhi ; I~l)ilt: in,~ ("'( bllm~ D~e~n tJil~ (0 oil ltd lile- (~ ifll thE! phClspta()~llirE!tln~ne fesi~lle Erk2. 1m

Fig. 4,. TiI'I~pho:!:~ho'lhrM· Iilili~ Illl'.ase a'cti'llity of other


OspiF faliJ'il~ Hilell1i!Jets, (A)

~dlcc!tm'

f!'f:l!l!l

Immu~Qb'lQttililg ~~Ys, gf remONing pIiIM'p;ha~~gMliJl5 'rrom p-Etrk2 ~~ iflldi~aHfcl 05p!F '~ami~'l m~lllib~r~ ~n vitro, t" time. (B ,an d (I PI100:pho-[ nk '~l'IWuoo~Gtri:ng assay5 ®~ dQl,1I1i1-i'E9ula~ion, l!;Ir Eli~ (If!). JNIK~ iUl(f Pi:i!lWilYS ~. SpvCtr<ln~fec~ '~Ol'l in 2'B1i cell'S. The aSXiY '1\1as pe'r'WoFmed 811i1d tl1E~ r~u:lts a~ Iilrmnliffil ~irmi" tarlya,s mowJ1lin Fi~jI,1, Il

~(mtn)'

EIil<21~~;iiii~ t!Q
~3!l1

so

3iIl 3tI

p.3a~n

C
~Hl1\~·~
.!L., "if'

Fil99"HA.~ ~d.i,l p-JNIi(

JNK~~~~~

direr~e ,a,SMYS or en'e!ts of :5]jlvC ani IErk s<1!Jn.Jl~!'I0 oldi, VaitM blj! ItJir;V12 in 2~Bir (!?lls., !Error ba rs indi(<I~e '5tallciam deylal1oii'l.

and C, re!>j)ecti¥ely.

([Ii)

LIJ'

I( H. I :iI. iRCCl('!m1JiFklilt SpvC ,C'mcie~u!)'~cHl0ved P~lO'iiI)·hl)tc' gmip~ [roU] GST p-!Erk2 I(r~g"4A~ SpvC (millyzcd rhe s~mC'cn:tym;-nic

lk.:a.. t~H I lUIS s,-l <lind Kiln 21':.5 I ~iMj. Similarll.!!, ro U~I\F. ('I<~'Crexl:ll'C:><'~Qil Sp'!"( ill U.r mmnmrnli.m cd L" ~bro:;::~ncd ph(ISllIKII){~ati(!Pl of cOl1vrnions or~hc Erk2 ~VO$phopcplide '~'llh mIdIaCI~!"~nio!l JErk ~/l.JNK, and r3 s kin ases ki UCltics crunpambk:w ~~I~l observed for Os.ltF (Fig. 4.13 to :[J'y, In f\\ldilion. nopAU. a WI)!;!~n 16 FEBRUARY 2007 VOL 315
SClIIENIOE

ns

cbecrved in Ilela cells in~~wCJ wldl the oS)'"l!!-(lc!\:~ioll !'>U"l!10r sf{~l;ii!{~ oomp!:)~clwi~~.he amoum :i~! eells i~lrectcd Wi~l tlIie' ~"'.ilid-11I'fi'C' slj'"i~l ~!ig, S~A', Furthermol'C. re.i'fltrodw;:i!1g wi I(I-I.~!IH~ OSJ'f:, hut n!Jl. theIWI4.A. u'mlant, i~Dto·tbe U •.pF-<Mc!km ~rnin 1o"""Cred Ihe al~lC!lm[ or p"p3fl ill iI1Jicch.'d Ilel.n celrs{fig.. S~Il). T1Itq::l..M}\,I)Kis a polcnaiailJ!l'g;:l
W~Olm! '11[" ~llJI-l \\,(1:....

rOs.~<;>&~cd. 1.1;,': $<I~'<;l in \' ilrtl u ca1:o1!ytk i1C'li\fi~y as clC'!'tCI'ibed .for O.s.pF~~l(~ Sql\ C, ~I~ilil.l!l cilhe-r p-lErk2. {fig. 41\1 or Ihc Erk2 r.n~sphopcp~id~ as~he aJ,!i fi-ci<l!ls.uhSlrmc lk",,~ 0,,7Q ± 0.0.2 ~~! ~ml. K.~ L32. ~ .dU ~IMl., = ThLJ~. I!bc ~mii;C o'if-!Fllimily ofTISS ~:lf~I.tJiIi:;, h~::;[1~'1 in v il;mpho:;pliiollirou'nillli:' :Jy~~c [leI ~~ill" To ~1\mDlli]1N.': wh~llk'T I~II!' typo;: I n-s~!Crel... i11 OspF illlci!!Cd li:J~ct~dlhc h051,MAPK..~. we gt~n!JrdK'tl an O_:q)r.lId.clitl!! ~u~:[in Shigdlfl of flt'xlU:.'ri J:arund 'pcrfmlllOd all in.l6cl.ion <I:ili.'ty {/61, II] (lmU:I1~-;ll 1'0areecru rupu'lt lhm allalYil ... "tI p-E~kln~t J hours JX~~liIliV~:ltolllU 7~ we amL~poo p-p3R at :m miDi po;sii;rlVa.~iOJ1\. bccmL'>C OsrF W.3;.';, S linb~"s.111[1(,(;' acuvc I.t'!"\vard th~ pJ8 pl:Hl..\P]m~IJlL:idJ.: ,~tmilp!l~"d wilh ILh~ fik2 plmlipl1{)P~I'~idll:ll.ablc S 11..1:1'1 arldilD{III.1. :scv0f:DlI: 51.1:uil.ics 1,!il\"CS\llggcslC'd thili~n~F is onb' tr.lIlSlcrm:ICd Rur 2~K~U!'Sller ha.c~cri" hnom:iolfl (at ; 9)"I\~ig,he!' i
,~p'fI'tW-~~' 1(/

n.

I'~[]i)' I"'l~hoge!f! P.~,'j.(b,moll(J:\:

\l'V'IM;V.sdencemag.orgl

REFORTS
orlttC phosphothrc'Onin,; 1}'iIlS\: I!:Clh'iDy (If Ol>l)1' durit'lg SMg.;ttN{~ Il1ifecti:on. p~R~!.phonin."Onin,; :lyfliSC 8uivily has 1101 Dcc'u described PJiC\~otls~)'.lhc do.'\I."SI.clIlzyunc n:l'i~~~be t~1C dl~0I1]IlC $Yinta~c t EC 4,1.3 ,II iU'll~a~rillfU~d pbnll'!. \\~]Jirh LL'<l1'-S ~ pyci,oo:ml pJlQ.\lllllll'l:c cofllc:'in[ Tlli~ t'lIZ~'I'I'1C {lon\'CIit~ O·l1tim!Oflloo.[_-IlC![iflOlK'I1iIl~ il'~(1 '~1!'rn:mi[J;[_1 lhml;lg~1 an iI'LCm]Jl!d~alt'!Ihnt .h~:.1 ~
l. S.,MuUMllj~ ,~' Q/,,, Sdt'1lLt31~ 12:11 o(2:00D}" F.. R. 6rmn.. ~~·w. M, WJJrin.,SDimU 297, H, 204:8 ~200l~.: pll'Mi~iisll ~fiVinl?,<':9 Alllfjli61 ,2002: fm~.1261i!:ient@.r073;lIJ!I,n. 5\, f.'iI~(I ,ill' QI,,,. Sl:ii!lllLt ,3:tlil l2:30 {lOIH). ,6, F. S!~, 1". ffl. ~,ffil~ z:. Baa, ~W. i!V1e"1" I, IE. i}i~a<!!,
,ii, j, M. Pail ,"" (I,

C~I :1;.09,~l$ aOO2t l~ f'!1i1p~1t s, Ei. 1(5!~o1ldi!!" P.]I, s.ar!5llI~eui,. rrt".O~

C - bomlbcl\:\,'OOJ] 11mC~, m~d ~illC('y ..Ho\'IIC\'cr. n'q)f dOC! nrns~mw 1IillYScqlICIlccidemiry w:ilh ru]), pl!ID.\pb.n!y<l'i~ • .im::luding 1I1~;I,nnil1O ~YDlthasc, run] .~cql,liredno oolJlClOrs. This ~lIIggJ.!'t.~ ~ml~hC' O:'ll!F faJr1l.iJ)' ph.o3f)lmtl.ll:1.."Ol.liilC Ilya~ lID.~ g~igocd llIl I,lFoHl',U1! n:lCclm31Ji!lt11 rOI." (mz;ynIC ,cmai}~is, TI:F
gclh~l~ O~pF" ~lwC. OIlld 1.!npA:t~ dellnc.3 Jltl'l1ii~l'

Iimr.!~ l~, 4~ {200tl. i!. $, l i(iI,riUd~ e!at" i!'MSOfi~ ~, 736 120oiil. 9. H, IKot!lff, s, 1". Ro:I~ig~i, B. Ii.. .N'K.COf1T!ld!,Mf&~ IlI!it)[IJl. 7'0, H50(<!OO~., ill. I'lt O!liroYa, 5a!alCawa, (efl.. MiormMcl C 1n (2006]'. 11 x. tu.e! ~t,rroc. i!l~tJ~d.~iUSA ~JO:!, :l2~9(i

a,

a \1'. Z!JIr,lJW5Ili. (, Mit~ui'!~i~l, t:: ~. MaJmy. II. ~ r:tlormock, ~ T, ffliiJr~LlIIIl'rfr;~~ ffiilmrtll, 7'4, ~:9,M ~z.tlIIIIi'>.l. 18.[ Ill! GIi[! ,ilt ~1" ~r&-rMlGrogy :l!:!il.'.l.~l il200§.), l'i1. t. n K:t,r.e, R, SCflIllCh, W. A. 11}A:rlr" ~ T, ffl;!~r~[Ij. j,. ~(l~ 'is4, 4400 t'l~~l'. 20. W~~hari~" IDhOil ~r cri~(a~ .1. II!~ J!lii~ilIKIjWl; R.~~II\ .1. i!)t~oo. M, ICIl~b, M, s.ot..o~, II, Uf\ t ~ste'lin, ~ I[)a~neribe~\ ~!ld tl. li!w for p~dil1g r,e\)i!lent5:j 'W. i,w, for prwldi'41£1 lec!mkal ~ijOO~llo!tl, ,~ft!ol!tNaeOBI ffi~O!1i!i~al!O!lI; oil!'l~ mE'l!'i!>t~o! 111~$h~o 1$ !.:!r j}el~iul.disrus!iilm~ ~nd 1l!!!lrnil:aL ~~i!rt:iln[p" Ti1li~ \WJd( w.!I!. 'lIIJIIKIiili!d IiI}' (ilille!e Mimi:'fr.¥' of Sdefl,e 2f1d ;r~hoollllJ}' '~6~:" ~OO~/!.;~,'tlm95al F.~) ~n~ (
17.

ft~,~ ,~

12.. (. I[k]~!i!, III. [, D.;I'ofu, Ill. .A. 'PI~II,

COO5-t

ZDIHM2111000
M~folR'>i;'V~ I'mIlllUoo.I,

~IA!"'ZJ

g~_

~D.'5S 'tZOll2:),

B, );iit!!1~ILe1RQr .1btlrM~iooli'l~for
.!I'!!

.of

ca!'l~f\,<ed, 1),!flhog",nk~JTcct~1S \\'ilh a !!I1I:iql!e 'PJl(n;~1'Ihml]l'Conl!l'C J}'a~ aC1~vr.ly th.u IPmh':I~~Y "'1U'gels ahc 1m:>!!\,fl\r~Ks in l'D..lllmg.CIi~csi~,
Rd'E:!'E:rulli!$ and fijotli!$ ]" ]1. E. ~b!'\, A. Cc!!!roe1. Srk;n(>tZi'!.]312 (l~91, 2,. IF. lit A!1Jllllbel" Nat ,tmmlll'l~. 6.. W~ t2D1l15')o,

ft, A!.a: (. ~ 0". Alp; ~,(jlm; f. 1"11~; iG.\ Gi~Ill,I!I~ I, lilt. t:i; lr~ l.. UII; M. MI?~ rt Alii; f., 1'f(J. 'll GIn.: P... W9> s.. Sell: 1. Tlftr; \1', \M~ w. Till, :r.. ~flI" ~mQja O!!(KtI o!ifid '1'. [Yi'. 1~, r~ Gu'I!g,.l .1, (:)0'j\1~. llf~rt (l'I'ImlM, 61, 5$4. ~1'9\1':D, P.

a~ fOLiM1&':

iI'iI~

amifllO acid

~"!s:i'd!Je1

:SIfJlIPDrtil't~1 Q,l:Iline maL'!!riil~l


__ ~i;E;tK!!m'!Ig.JiJtgk~iJCootem~ui.l!'31:5~aallllOOiIIlD(1

IMa:Ima:is ~ntl MelliDlll!


SOM 're:di

'~I£I~Sil1{! ~ talltf: Sl

150. S, ],. ~bb)' ef .:n!" M~(i. ~~m~n. 65, 1786, Ut9?'}, ],6, Mollerlilh ~!1d me-:lh(l~5 ,!ji,l\l1!.;'I!h~~ ~;p,l)ljrli~1 ~!'e ~L\fri~~ ~;m ,~dfi'i!'~ Qnlint,

IlerlC~~n(1!l:

1S: ~'t!Tiberl'OO'~ i!i(t,~ lU:w.rgi~(O!l, 11.3

e~~o

12 ]am!aflI200'l'

Ar'cha,ea,l
lilD

Type 11111 uBiisrCOs F:unlc'liio,n R a Palhwa,y flotr .AMP 1M eta bo IIii sml

Iog.~tl'f sc\!cm~ :;!li:p:; in Ih!~M~l ~i1LI"'iIl,l;;O;;: 'P.tUll w""y. iftt.::lllldj II1g t!Jl\;(; YkJrS !J(Ii.UiO.ltlgS. .;,wc PnJ'!.'>CI'II. lfkrX artdYl>a lll>ltlOlog::; CilUflliot he i~cmi 11.d]
Gnl Iii;.:; It;(HI~llulmel,r,~i. gi:11.rnllC.

lllil.: ilb<.;I,,:n~,.; Qr II(;OJII\fI~"'!C:: M'1, i:>\iJ!V.igc Il'iluh· w.~y~n r t;oclcJml'm:,:n.W,;_\;wa.~, (,"titrl~m1Ccllby ,1,.".... .I111~·
ililin~
;1

1it'iUltllUll:wl, kldb, lilc :lliiili,ly to :;~lIllhesilX

lh.~ ty~ U~ nbukl~1,5~isph[)~hlilte (albL»lyklI5~xyger\as.e (litult!i5.00~ p:r~rltili1 ti'l~ o)Ir(lha:e1ln 1hel~(~kodakllrolWl#S w~!O rol,lllll~ ~I) ilJarli(i~~ in ~;d~1i11)5nn:€ .S:o.moO,n{]phiJ~t~r.AMP) 1In1)'~~lJoIi5m.l .p m~ that is d~~n(t tromlDalt !l)f (13'Ssk:a11 RI;IBi~O~ lofllil~ ( a I'o!ira-l1h~lliS[)n~E!~il<'S~am .• G~ffi (~

<lm.OItill~ as tlwmidirte ~1!)sp4lOryl<l~ (dOO!l) and eL1'{.illiy.ill~ra~latim!'l ~l'Iltia901il raa.or2~ 'ie2b2) \\ler,e round to encode AM'p phQsplliaryilaMil!il!n.d ribOMl·1,'5·bi~tl!1l:Sp~li;!te i5@meras-e.. r\'spec1hteliY.T1nesi: elllO:ymes :sll.lilP~ed !he 11hn1Bi!s.\:O s!J~~II'<I!t,e:,rib!JleSlil·1,'5·biilPhQ'Splita~e,rtrell1l AMP and Ph05~Iitii!~e'.AlrdtiillNl 'wl'tih

'lype [I~ I~ll Bi3C'os: alllitaribor I!»1:h IDeClA ilIrtdihe rorrespo.rtd~ng E2iIl2: homelog 5. 11.111 palihw<l'(,ad'oo1ne tms \l,lIllS rele~~ed ~'rOO1i gp' and the phosphoriiOOG.e m.tv err~red ee.Flltral.·cl'll~oon mel<iOOli5llll.
i

novo, lw~ ~U.;IICC'HtgJmC.g ll~Cl.'iIlnl1l~ ~ !1m COl~l?OnCII~s of Mel s;yJ'ltl.lQ~ lIJoL'tEJ (l1C144rl)a!ld m~1£2 (TKI44n \"lCIIC driSIJ'Ui1I.('ti;~ i(l61 ICs!.l~ling MUlin. iJm~1E2I"I. cli!1fIl<1}'Ct~ ~lC cx.1~1:d g.cr~flIYilM;: ~J),~,J,;' !.i.11~'tE2j:1!P:F I <lu:d Q(li.lik~ .~\I ("~I.'I!}' !!l Uli;l '1~11?I'W. ~1r ~L'j~~IS M'ct.l1Ic &.miil£ll-1 9r.UO cOlll~dnot gmw~il~~ Q.ltoDd~

M~ de

nc

'll>!.: Ih(~mm,[ ~~bLlml.illll~ ";lfl"ym~ (]in! our plmi!.:1 (i~and is witk:ly !knmvlI a~ til,,; key CO~. li:-;inlj! e:1'~yl~ti.: ·oil:' Ih~ C311"'~III·.i3\.~n:;tjli-U:Jl",d~anb ,tUm) cyd...... wilil"i] ii> pr... ~~m.i~1;dilPtS:lllllS, ~IIgW':., '!::y[U'wlhm,C[;IJQ!1l. mK11111:JIny 'Cinta~r mm)I~lli'c rnLCb rim. ~2. 3], .lki;iU:(.';S Lh..:dm'l'ic~l~ lype .~ :lIld II Ruhlil:.{:Os of ~n~CBiBcyeJ,c. two .!latmp:; ,[,:I" pro11.:ii1l5 'll.'tiUatt: :;'[[lmltmdly 1\]~n:LCd, II tiiK:'~Ypc' .1 m~cl 1

ib~I,~[}:lC-I 5-bi~l'I1os:J)ha!C .•

(nXY~C!1;l!SC

cilrl1!!)xyl~!!liC:IIlrctlolHHcd lil

lB.lIIB~CO) is

h;wc b!;:t1l id..niUoo ami di::>igJtl.noJ ;Iili Type Ulll4. j)smll~ .IV RILd3!i"OD.'i <6.. 7}, TyPe IV RuBa:;;COs., !11so C:I~~i.XI Ihe RuHi1lCO-!ike
prrn'lCiUlI;. b::k ~~\. ..'FnIn.':SitlIES nocc:l:;;~1I)"rbr COlliVllJlIIUiOll!ll RuB~£O c:mIIYlli5" TiIlJl:o;, 'lype IV RuBis":'"O fl1L,~ll!hJ{j/~ls ,m!jjjlt~insIC<K~ cxhibjl,~

n cn'o/ull~

2.3-4iJ;:J!'o'-5-QIK'1.hy.~lhio]l!cnlyl-l-pll~h[l~C' 'Cfl.(ll<lSe IJClivllj' imd OpcmL~ tn ~hc lneUltoHirtc (MC:t} ~h'~~o;;: (m~~'!~'i.iiJyln'fJ, .'·nh~lh (7).
'D~~l'I1mflit 'Df ')yrffilli~ti{ (h~m;~1iI)'iIflid Bio!~i(.il' (Ilel& i~U'l, Gr,iIOOitte Sch>ool 01 Engi IlI!leriJ1Q, KyOCil i:J1U'l'efsi!)" Katsur.a, Nlishi~lOO, K!f(iIQ &lS·851O,laj:lilD.

'To w'1101~ (Q!~~5IpI)rud~rnoe w1d 1J,e, ~dd!Ol5'lli:d. E'mili~ h i!m!!i!~i!®ibdli!e!1l,!qroto4J..a~~jp,

Oi:') R~IIBi o gene l77.:-dld i~!;elr a~d ,<:.ul\~¢.. c qllCn~ Ily i!1o(awd tOle" genc,-d k;;~lll)lifl,Ul nlll~,U1t itlnhc {~.,r'b£,-~l\AI. \\hid~ indicated t~);)1, Tk-d~' is !1(1.1 i\n'e'hac1l!. <IJ~ld tl'lC}' h;lL'OOf an of till:: residues nlX:(.1l.~)'fm' nbLlkh'i(:-I,:5'-bii>ph~Ji5(lllfUl';; U~:IIIBJ', (~Uli.d fur gl\lwrh, \1\f>e: Ui(],~KO~ycr.o;Dub';i;:.;~· "-~lItly ro:.l~lrt,l,,;n :;:iigln dlXmJ."\lJ in 111,,;o::U yil.::M "jf CIIIl1h!lxyll~~o;;(~'-!:D",.)'gcn.a:;;c[leia '" it}'. .Prulcil.'t> frull.l Tlr~'Tmot.;oi.'('II,~ kocrtw,arflt."1'1S.is (4, 8- ](I), j\,jelfmfJ,}'11O- ~AA a~cIJJliIJ'mrod l{'i1 the hlli;t:;I~liuIWJl!.~l 'li!Jmk'm."&.'C!1S ftrllm1sei1ii. AI"{.'ll(,U!/;;glufm;.~ fij~gl'- (~IIl! Wt.U ~~\IU :ill nullri ..'rII-rieh Illtooimll. 1I1IS, mul O[Il~,. areh:Jl£:tl(5. 11)0 ~1'ihibll. RuBlf>CO w~ ~Jlll!nl sI:dtcd OU.r all.\.mUOrl In ahe Ihn.'\: Vk:~S hUl11()IIJ~.pJ!t';S~I.'11 ona Ill,,; r lifJ'fklkmm'I,1.'iili :l'C1;iv.ily, ] .!owc'\:J;.':.[,:J runeriull!ll~ eBB p'1~]way ha~ gtllOll,lC Ililltl (lm~$it!clIVd .... mitw::; ilO:>::lilJihl:i(;:; Iii:JIf 1101 beell :i)ticu~j 11,,;1'1 in i\1t~1!1\"1. mt<! ;lttimcn~ &'C. n~lc nlk~:;; Ih ... ';S(; Im[lloJog.;, 1[11(; l.hllX: :110111011001.1lC :;(;qllu:;rwlJ.,; do :ruo~sll~p(m lhe p:rCSCII(:i.: Io~, 7:k-E.lb I ~45% idl'!1I1 ~i;.'jJl w ith ¥I;;~S fi:mn of .. C'.BB ~1iI,fI'IW3y, In p,lIlitiCM III I;, hmmil~)g~ of l/, sllblirls~ n-:ElB2 132'ilf,'"tkmit'3I~;'Illrl Tki rlm:.;plmrihullok.i'l'ttlsc. ~noIDt0T key 'l~JlizyrrKl ot' the cm~ ~)mlhw3j' thai :;\lipplk~ R.IIIH" lor {l<:d:2 tl,% itlclII~icnl). ~Ut:l:nCl:Il~ o'r the !.:u.Iillrylltic 11"J1l;,..;1:1I iioll i~1~lialiou ~.hclo:r :2 B ~clf2B) R!!Ii3iilL"O. carllml. b!.: tound Oll lh~ miha ..ilil 1.'1:. ' rnliYottlmily Q1 '7 !9l. IH:m\!I1'~,,"~r.he l:i.m.clioJ1.\of t 110'J11!l:5. Ik:ncu, w,~ IIr~l iI1Vt~lii,g3Icd wlacth~~r tliIC elF2BI'tQTI,n;liIJogS hi lite J\J1'C~l;1ea ~. not OI~chaC<llIHuBi".c:Os '.ve~C' ~nIVQII\'(;d ~ili!MC't ,~~"<i.ge. Si!lniLJI' to type IV [ilUll3h;(."Qs., k~mw!1. 11)(: 11 .~·lfbNli" YkrS i::» a !1lC!nbcr of this I~IC' i~n1lCrir:.<l~i{l!l of MmlY 'OI]t~ni~'Ils usc Mi;:I, .<IS. a rrr.x:uITIOr for Iillnu [y. and it Ca!iiJ~ rl::l~y;)Jn inc <:"i9!tg.!!!i(1) wi!h wn.oomi\;I11t ,g<:"llCr.lIfllo:;~hyl:!h!Orihfl~ H~h~rll~W ~;V!TR l-:iJi) ~L1 ticm (lfmelh}'llh~ood",oosi,nC' (MIA}, The Mc~ FI1ethyhh~flriblii lose l-pllOSpllJIC· (MTRu I-P) ll!C1\\'e..'i!~ )1<!iI\lfl.~'C·I'Nl'I~.\'a)'i)'i~if\;'SCiU in son~c'h"dCri<l <lind I( 7). We nOliced ~1fU('ll!r.lIl s i~~lil~.ri~iC'.!i c~l~myolC1', ;:);!1drocyele.~ MIA 10 Me~ (fig. S. n MTR~ -p .<lm~ :nL~C'~ .5~hi"Phmq:n~l[lIC' ~R~5]" til /II)" 11~I~toIDg.~of enzymes i!1\lfll"'<cd~n Ihc and b.."'!WCl.11 tvrm.u. I-P am.! RlIi:liP, ul'l(itcMing COl~\'C~~cm Me~ to MlA are pl1Csem ('If!! the' tim!; dlc.~ mthclC'al IP2B {aIF2B) Ill'OttilllS n1i~1t liICinr ...ohred ill ~Ilc: gC~lcrnn:iol'l of Rlil:W: T. Jwduk'lNlcw;,r;is ~CiitOmC (15), Ahh".}ugh iwnnoTIle- lyfilC Iml RIJIB!sC{ lis arc
rfltl!ld on~

or MTA. \\fc~l~f' dbJt!I~OOIh~ :r. J.."(J{/(IA'(<N~#;,w~i:.~

MI.'!!.

WR" :'l!.!1'illlillUJ(X1 \,~,~~ \'a~oos CfiJl00~Lrntiri!lS

or

or

n~'C

REPORTS
iOOmCJ~l:.ilIIOn of R i5.1? IOCJo Rulli? 11100 coo'C$('ItfII~(ls tetbe l'iCact~[ll1. :p:mpo::ed 10 be n."Sf!Orlsitr!lc: b 11,Oii'l)~Q~~vllt(lh includes tk,I!2.l.'!t2 911(1type n[ RuBls,cO.

B.uBlP s)'liI6lesis~1i

~r.I:',lB h,QTno!o~ rrofli 27 mch~ca~J..:-.cI]OII~~ The I:KlI1I1Jllog.~ 'COJ"I"Ipn:l&lSC'Hna~ mbgmLlIjlS. ;100 W~ ftXllld almlOlhl immcil t~~!:MIad in .Fig_ I) ilJ}dl~ldoo lilOnm!og." {mly fm:.I.TI. rlrgrnl~itll.m hmbrning type!' lll IhI]3,i~O;'i. 'willI ~tim Oldy c;,'OOept~rm lJ,cin~ ,1\}\Gl0059 lfurnn f'ill!'(HHldf!J'i!{m spc NRC- L TIle hia~i dr~lribl;l~i[ll1ll Iml. con-elale ill. aU '\viah. I.he clird play.lngell!.ltic 1~~rtiQn:'1l1ip~OnB1C mg<ltli"'llii~ d~C1!lle :s.el",~ lh~ ro~lali[ll!~ pmmpwd us III o]O[lIminc il;Ji"fK'I~lnliinmll}' ~~'hcLI,~'Ih~!eW,I!'; a rlllncloc:m:a~
.~(!hLlIDIIlh~Jl bt,,"!;~'oCen .. thi:.; NubgITI!ilp of' ai 8m

m.etliful.lOgclflic A:rdi<1tfL 1,2m, i\ pllJ'I'(t~rrlC'[~ U'!CI:'YilS COliSlIlJIt'~ W~dl 49

llic n~eJh2 ruld


1}~s&Cll in £_"drf'rl['lri,rj
iiuil ia~ ,..... f!rlil~llfltioll

Tkcr:2/,J

genes were ex."

cali ,;1l11dwat-t!lt..'flIcd. l

(Iof R\IB:I?

From R 15IP w~,". performed . lioll CllzymatkaUy prepared w.ilh ghloc!oo [J).bi~piml"ph!llt(l (G 16[1t ribose 5.plnm~lhm~ ,[RSP1, :mdl pllO!lllllDglucmlllLl.t:l'itl_ EXli.'ocls, containing ftzElbi ~U~I!ayo:l OIllly ncgligibko level" tlrl1(> ~hlilynhm \\Icr\;loqul~\'alent N.. 1]n\C I:OUlld ill exbact.. from E.. cali h,lJborin!!<L mQCk IJiht:;,mid. pml :I( + 'to'!fltll to 01.002 fl~lnl nun -I mg --'I~ [Ill oonnm.,r, ,,1tI'1O~lgn '[,.'l:I~i:olt !cvd" we~ !1U1~ ~.<UlilLia~~>' rllWl;\r Ih~a~ t~~ or n'"8bi, c1{lr..cts

!'\.lil S)"I,llh,.-si\li,1~li v it)' LL'lUllg ,Jl], R I. S P willi·

containing nt-E2hl displ,:JIyed. levels of ~luiJ:PSJln~IIi .. 'Situl1g .[Ie! i\ i[y Ih,n were over HJ Ii niles ~li,g~lcr I.I.'WU ~hn.'lC'!I)~ U=~::2bl. P,Ll!li ~IOO n;E1b1 gcr~ro~cd RuB:!? (rolll Ih~ IU:5P ool!l~j(m wi~'1 3Cli"'ily levels ,of 0.4 ~Lntlul fI]~Ii~~ Il~-l. 1~i.l,:tl·
li'CJLlOCI chrnn~togr.Jphy '1m~~I~'l-!. 'C(!UI· limwd IlmJt i.1IlllLy R 15 P; and not G 1. (:~por RSIJ', ~lia~tim :s;uhMr-al(l Ibr IhElP ~yJ1llllClSi~[.~ig" S2l, 1111order to :idcn~ury a Ilmlcilll thaI supp,l~c:;; R ~5[" we: tlcalcch0Q 1.'Of gcnt:~lh~l W,l'reSPlI1;:mc.rnUy pre:'lC!l~I.. un archae a 1 str<ll.in::'i .R<I Uixll'il1ig rypc ]! IRw.~is.CO_ lhis kd OCIIhe Wcnlitk''!!1ion 01" a rllt.'lh~ GD1)'i1l11idil'l1l: [lh(l~h(ll)rJ(lL'i~ gene I(d~"'linTI'bC' i;)!1ly C'XCCpl.i<l.!l W~I!S..lM~'JiI(I'Jt~~ tJ('1.1t~ IIJrJI'i/J(lhjdj.,:, '\\ hkh hmbm\'1, ;;3. ttt!a.<1 ~nc bttil Ilot
'perf.i11i11l1uoc.c·

--

.Ifl!: (oJ'iiosous AAL.l!il599 R ~klWJiJ leM29~e p iIIIl1y.t.ml CAB~OO

r ~itarwJll.~$

BAOO~7J6

AH; N. ,,martro~,.Nl1tr~nas pho~ DSM21160; p, iOfri'd(1S, .P'ihophilll.:i iwridus [),SM9'790; P. ,~erophirl1m, PyYiObrlc!i\I'um ae,rophif!Hi1 IIM.2; Po ailpJ, P;yfl:WXiWS ab}'iSt 'GE5; P. /lJri05lIS, PY!"Ow"u-", /llrWs,rJI, D5M3 6. 38; P. i:rOrikVihii, .Pyr'C!'Qn·u~ h"rik()~MJ' (l11f'3; H,a/iJW'aAfti ffl'QriSJ'l'WfflJii\liCC43049, M. ir:mllti$dffl, .liwn~il'ald'Ci(l';l('(!f~ 5. s{JU.(Jla&:~, SlilfoJ.oi:NJ5~fJtf(iJa;""t)5 P2;.:S: to/.tadr;ni, Srdjcloo.r.;$ tlik,(jd~fi J; S. aci'tiQwJd.arius, Suf/oiooos addacoJd'arius IDSM63:9; t. ~d~~(}faeM.s. i(lfifNlfich~ DSM2,66l; J\1Il lwrtQllii, M:etilrJfioaNx-oides tJurtQfj'ii DSI'tUl242; m£t:mormt,lJiS k,MIlli:amfl'lSis 11<01Jiil; T. o6dvpml1Jrn., m~opfosm(i acmophJ]~w M. marifIDtOOi~, .ME:'I!'MIl!!)COCOl'S umrfprJimils Si2; 1t1 IllN:lldlM, .Mt'tIilGriOPp1J(~ k,t;.ndteri AV19;: M. Oocetiw,r,on,S, Mer:haf!OSOfOOa acetiLWf,(ilfiS QPI; M. b.ofJier(, DSMl728;[ ro{ca~itrm. mE'mili.Jpbsmt:l LWic,mium GSSI.

Ilalllk elf )apan,anicll iii ph~togeiiielk tjlfi!l W<lS rn~!jiiU(t:ed Wl)h tile maximum,titBhO!ad method bit mearn of PHYMII.1[26. ;17). The shaded looanrn Mr~r:sthe .aIREl hoFI'HII09s fmm ,ar,diJa.ealstr.3iiM 'lliat plSlle5:$ a t~ 1IIIII\\iIIBIsCo. A, pemfx.. A~JIl',lt1iil ~ern~ ~J!; AI. JUfoM~, A.rma~tobl~ Ji,I/9MJs; VC-l!6; ff. fil~nrnartui,

.a1f2El i\1lrrtity,all~,tHI~rnri ly pFotm$llreentinl the 2~i vmpleteardilaml ( ~~qllli;!li!t;e!i ~r,e -JUg!'l~~~'!'!Wh .t!1~.OMj~\W 'p~r~1lII ~vi!ii~;;!!;!lelIt the

Fig_

]1.

IP11I!ilogeDleWic u,e@or <!rcllli!~~ lPl'otein15

<!!Ilr!Ol<11@d

<IS r!1@m~rs ~ff the


gemome ONA ICi~~

MdJ'w!jQJ(ilfdrl~'

OCMlB8;

1\0:'1.

barker! :str. fl!!s.ar,o; M. mmei. M'e:lhtilflQs(lj!:Ciffll' mQI,ti Go,ll fjlJt?gtlt,ej, ,Merhmlo:ipiri~,!.tm iruJJgotef D Ht; A\'I" sradrmaooe:,

Mrih~~(;js'~Jl'mli'r(;re"

IDSMZ309:!; M, thernro~tQ~,~nklol$.M~lll(lllKl-

;~l;jiFFrn{)oorl4;':r thetrntJul(J:rDphJru~

16 FEBRUARY 2007

VOL 315

SClIIENIOE

\I'V'IM;\!'.sdencemag.orgl

REFORTS
;[I Rld31iSCO rrilb~c: ] l.[)OO,iI\ is ki:lOWI!I ~O re~lse '~ile HIYlfll'ilne base fffil~l Ulyulidinc while ~liltQS. phory!mill,!,!; tile 1'=Cf.Iruoil of 11,1(; ribose moielY {.:n~f Deo,!\. c(ltlldpIIO$I~1M_ytme n:LLci'oo's~c

cxpue$SCtJUiC'

nnoutJpho.'ijJllfllC::>,

'thi:; ~W~J.I:d

gio.'i11,C~;;MC I5,P;Wc R

lk"h'liA gene nK03S2) iu E. tl)/f. ami f04mci UIllI. Ii prood:llM 11Iigillevels ofRI5P in the piI'C~n(c of phos1)113lc and ~.d:ento..";llC: .'l'.".mouopiw:>phtll~c (AMP) (Tahl;c: :210, Tk·[iooA di:zph'l},cd i! :;PQciir"icaCLivit)' pmifi.oo lhe
l)ft)'[cin.

or:::!] J lJI.u1'(i!]mi ii-I mg. -i for tl1.erell~'!iOl'l ANIIF' ~

PI=> It ~:5P t edcni ac, "HIe stoichiolllClnc m~io of n,c oonCenl~~lllani of AMI~ OOll5!J1111CdU AMP ronl· :>l~rnlXiil[iInd [a<l~J,lilieptOOlIIwdl was >conl~lI1ricd as l: Illig, :53~ The ¢q!J il~ibri:llUn CO.lll':lfml. K <1R'1.:]rn~dkl'JbiirnC' J/I.,M\(t:~'1I Pi!) was 6JEl )(m~!il t (t4(l >: IO-~'ITt flS"C U..illg 7k-lkoA. w,~ w~m ablll tO~ltnlc:;;izc 1:llgc "'ITlOIJI,'~'; R~.:)Ji'and III ~e:,1 nf ~m R l5poo~Il.t'r.!!\(! illCli.\il:y Qf 1~·E1l;.2 nil il.1C p~"C.r;,t~ncc r'lbllnd~lnn:;'Lll)..;tmlC'. We orne'rvud Ihm O tt«~~ ,tH17 Il1DI of3"Ifl!m_fiplm~~;l.yccmtc (3-IPGt\J WCt'C prll(Il1.CW, for ~\'i;:f}' Ililelle;: 'o.t" R I.5.11i oon~1I1e.;!~\ih~n T,t~~2b2: and Tk-RlIBi."CO W!1<fC CQ!I!pk,<1 ~5.o;J.C, t'ilIDJ1~.!l' ~!JII~llIOl1ing. !lloj· . chkilllCl~C milO of lRJ5P OO!lt<uulooj:p.PGA pood~OI::odl - 1:2_ Tk·E21b2 exhifuli~d1L spoc;ilic: ;U;:IUVll}' of 32.3 !Jlllwi mi !!-! n~~C~ the rror~ ion ror

Tabl~,e,1. Df,Slrnoo lio'lli o,~ ltlu8nsW,

a rrhaea I. g;eoomes, Shaded be);,e$ilfttl~Q;M.elll1l~ presence Type IU J!l,JlB:is(O

"'IIF26, [)@oA, am:!1AD:P.-dle;peFld,elilt :sug,a,r .K]nasoe h OIlllOi[oglS on or a hOlllfHII.ogl,1)111 tl1egefilome,.


~IF2B hO:m:lo1Qg~ ID!Il:O\A. h()mo~!l)!!I JItIDIP·d~~@Ii'IIlI!l\I1I~
:SUg:aI~

kf:nla&e

,u~

R 15[1' l1ilC

.miolll oo.W¢.:11 Jil:.I5P ,lmd RtJi311?' in M.jmmwi(:liil I; ih~ prOOlJct ob' 11.11000:1,but ~!t;;n:cHltibiUtUJi,pruo;:.b(1 oo.c~ ~i!JI~';;fii, 10 be ooivciJ1 "ilm (Jll). like~~ii~1Ix: MJJ(Jf,tl'l ~lVlliOk~ iai [, Ar_Jfklktnif[{"J1.'·~\: Cn.-·t1rri/, TK04~1 WJS "'"'lp'ns~-d ~nE. t'()/itIJIlrl, again., .:JICliv·ity~ as .nio~ tL;tcct~"tII, Ihii> i]itti,CJ]i:<,;iS ~11'aI m iCML in, T. kmldktJl'(l('mis. Tk·E21l2 is Ihe
crll:y.n~~pu:iil1\ilrii)' l\.';:;ptI1l1$iblc rOt .lRulJP Syul~li:;"
~:;. (i:1)n1

pn)l~'in

RIJ BP (TabJ'iJ 11, :lll,'OpOO-OO >Calf~~~a;; i~ 10 I

loouncr1'

.iU:5P. showed thib~ill To lu~dc!f;;af'(i(1n;oJs.

M'e~hlW·"!tletm()baCl'el'
,thf'fIj~a.tI.t.oII'JQphic.u.>

OlMr K':Suh,o;
,III

3 rLMOOI.,.!.: ~h.YClj'iidjli.~' l:tO,,":phtif)lll"''iC i p

:i~IIclluall~y

AM.P phn8p1'!fllyl!H1'C',

WIH,·W·iiL~.(l!l .:dFlB

~KlmD1og iii. ~nl,lny aR1S],)' j~OIl1C!<1~. [k!l;:;JI;I;.;,c ~lC J\'lil~.I;·<l 1~:;si.n.l1l.ll)lllC IH RlIBiC<:'O <t1:Cair!: OlIllt~mbk;. j~ is; 1'I11OiH liki;)ly~~,!t m~'ly the c;u[J1'fi";yl· a'(l' .~tlh;il,y ,of Jtua~co. a~l.ool. dlC O-Xy~!l,,"~ ~i-~ ily. ~~~~Cl~~1gJ~1!y rel~I(W!~ ~] t1u;s~ !<Imi~_
l1!!.ls.

Tk-Uoo.l\, 71-.E2h2" <LIld!l~ R~Bio£O


AMf'. l'i!m.~'I1~(i.~. 2, C(J
O!lui

COIl"

VC!'l

112;0. ~~f.it."C-

~h-c1y, ~J):tdtmhlC <Ii!ld ~""'n nlQkculel' of 3-PGI\ (Fig, 2), This padlW<u)' h£!loC'~<I!I\lagc);~hlc~d~lfIi~lC' Im.~ of AM,~J a~d di vert:.:; tli'C '~ihl1sc uloiCly
imo CC'l111!.d-C<la'l'llOH

~ltCl!a~li'5~v!.

MW1Y ~)rli1l; RiI:l1il7lll1l1'l~I"O,,"''i';.» thil:. ~)~1I1Wll)' A tm: mgtlni~ltl:;; i!li.li 1I~ ~tusit.K~ .5'·-cli~ll1):;;pb~l"~ , ADI"}-tkp.:mh.:nl ~A.Mr.~i:)IiI'I~~J:I~) ~\I~IIt kili~~"';;::;

litr ~~yrolly~i> {11,

11},

lllCf>:~

,.;nz)'IlL~,

,.multl

plf(J{[ul"C ~~ 1l!1l;-'{~ il:lln'!!.l..:III:LIII!urpool Oi~ AM.r as compared ~'i~~] n~~I]i:-:.:m:;; PUl>l";;~[[Ig elassieal .ATP'Ed~lldern kiuases, Because Ill>..:prOOLJ.t':i! f o

uhc P::Itl!\V'dY. J,·I)GA. C'1LrI be 1J51.il lih~.- ATP glJll)l,~rlIliml, ~~m ~~Lhwll}' Il'lil)' enable mmcrolJit:
Ardl'~ca to
!lI~{;

A,\lIPwh~'n

cner!:}' ~.c'\!d~ !In::

ab~en,l:i~

iIlaM.e 2. SPe(nFi!l3J(;lll!O!ities®f r~·D'eoA anfil rt·E2b2.Sryn ~hles~s f 3·PGtl. o ~r pFQte~n or :S!.!lbstr~IW+

\l.!i3,S

not O,b5Jefvetihn

the

(1;~li'I'Ia~mill'l..J. m£l

:Spec:i~i(: aCli!;l1ily ~t IDr~t'eil1~


22.3
;I;

knv ami i~.·inl:rncr:IllIlar A~;1P' is ill1 excess, .h1 A".fillgitills, \\bd.l tb:::; mn li:!lrbt:!i' Ai}P'-I:Lp.'lldcln ~g3Jr kim~~, ,(1:tIl· pflnll\'\l~L" IlJl!lIy be il1"'~~.voo illd'!iC Ctlln~OI' or dlC AMP g~lCr.uOO b;'ffidCl'lQ.;_,;,ylP~IQ$I~1OS'llmllC roollClooe (1-,n c!\irll1g s\i~jiuc

AMP
'GMip' (Mil'

10.2

(Mill

Ij,MIP' TMP rk-E21:i2.


1'i:15P~

rU4 ';1; 'lHl'n. ~106 ';1; .cUl!


0',01 32:.3 :!::oA

ttd\l!Clioll. (-\m)lJ!Cr ~'iiOOI\i!ilili~)I ls ~lal AMI) is ~xlllWd 1~'Jm .5-i1i!Kb,;"1l~K!ri.'~I!i1YID,I1.4ms~lmc (P[{P'P), b)' adenine phospllOrioosyhwl$[~~. whose hoOnl(i~O; io.: 1):f"cSCi'lt in r. kJ,rlf;{tumf(;lr s;is. This !!C<lCi ~m \'\I(lIjM recycle- t~.c odcniuC' nlo!ead;;: 1iC'1!C3soo by ]l-.[)roA lm~1 illw oomNcte a cydk: rmthv~lay oomposed or ~i} pcntose $)iil~lCsis. ~~~ conversien oH)RfJP 10 3 -PUt\. il~KI(ii IJ l

REPORTS

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(lQmee:rnIO;]of ~-:l'IGA to .l'mclJ:18c 6.-f~~l.Ol'iplmL~ ~illh.lom~ro,gi']IIU!'~~~. II" the 110l1~o:.xuclnti\'c bl]L1l.'Ch or III~ l~n~ooiJ flhO$~ll~j[~ pl~rl~'ic\'ll)' ~.J:.c to be W
.L;~pt!lnsihr..~ [or

pcnlr.ii'ID ~)'mhc:!'ii~. jhe cydic

11lJlh\\'ay wmJ~d mr!~'ritLlhl:J CO:l lijxlll ton pal]];o w'ay (fig. S4). ATliholigh their gtill.Ollil~ :o;cqUlt"ltCl!.',s, am [f}[)~ aV<l:ihthk!hCl'C' an::: ~cVCRLI llu1.Otmp!]k Ml:a~I1,"" In IliIll("n,.'1]Wmco.m IIllal. ~1\hibll R~~Hil1m actu~'it)' ~~ ~ad;:jng phf.r!\l:lh()ri!lIl1~Qk~111L"C fie> t i"'iq_y('?5). ~)pc in Rt~]]i$CO~ seem ~(l 1m",,; a r~mc,tiou dislllnICI,1i'O[U Ihfll of !lie: cl~~sicniR~IUi~,>COs : in 'Ute ( EM. p.~I~lwa)' o( jJcmt('ri(J ,~i'L(] Ejtn~IJitj, \¥~iCl1 (Oliliiddcri~lg thm ~hc OiJ!:flli sms harOOr1liY i '~YliC~I~ RlIl3il,(:OS n.1JJ'e~cm rc:rmi veiy deep'llu(l shon IllI';Ilg,c:s in C>'()l\nlt:lIl. ii, rnflywcli be ~'kll 'til...:: C::lrOO.xyl~~rtCiu ...~iy ef RlLBi.'!C'O onlWm1Uy .";O"ol,,,-cd '0 rliltC!i:OJiI in ~lih paiJi\"il)~. :t :t
Re(iem,m(;es ~ niiil Notes It], ElIi$, rmrdJ ~m, SrI', 4., "'''1 tt5'19), G.. M. F. W~t~ill F. R,TOIiIiiL:li, ltMS Mkl;M.!Q1, i.t,ft, 14·6.• B (1:9'97),

:1:, ], "'- ~n.eL), G. Ili!I'I ~eule'l1', IN_G. .t.J~~. R'eI'=. MIt;r~'N!lII. 5~ !!:~1C19~(»" ~" S. I~~ N_ Mo!ed;I, V_ ~iil1iffi!ll'Jl HI, .~1!!Il!~. Im;j""~ ~'. F B.to.I. Oi.!'.m, 2:74, son {19'.l~~ s, G, D. F. wa;ii!DII, 1.-1'. Yill.. F. 1I:.]~tal.~. 8tl~~l:rmJ, li!i1..
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~200.5), 2)1!1, ,t!;. '!itlh.lffer" G. IFlIiiz. P,. Ill, IH. ~rooi!!lk. IJI. ~Irn.le~, ,1l'.ioc,lmtmlsrzy"'5, 2%0 (~OOM, ,2'5, .Pi'L Hrl!JlEf" iii. Hufl.!I·, IK, O •. 51~ier. rio FtJ~M. Anti\l', t.li'(\!IlI'J.r~1."'9., 160 ~iOO3), ,2tic s, Gu:ilil!kill, 0, G\3i(l.Iel Sylt B.W, ~,2.•. (>9.60~:!OOl),. 2'~. s.. Gu:1!111fo!1, f'. ~Clh!ili:~ fl" tJli~Ll.o:. 0. GaKilJe'I, Hudtk

M .. W.. fimru.. F.II:. TalOOta, I,. Btlft~'1j'!IIl 1115,. 30W ~~~l ]. 111_ 'Wr~!f. It H-Iige~J $i~~. !!7~ U47 ~19~5)', l;l.. II.. 5et~.a. It. I).aEl~hill,. toor( ljli{~jar.;:!:, ill QOOI!I'l~" R.~. et fMiI'( Mmi!.ik!1. 4(, 9 !~(!~, 15,.K ~iJkJJii ~ i!l"L. Gt.iM~ ~ 15, j§<! R!J0':5~. ]'6" Maleri\!ill~M m~Md~ ar~ a..ai~~ a ~s ~~li"Idiil91 U

;K, rj, ""~ed~ ~I ~,.1. BtoL 4:9~. S1 (]J.~~'l. MDL !i. k. t!Jt<1!l];O fU!'.I., S.rrllra~ 9, 4H (~OOH, 1(1, 11, fflIed~ ~. !!.Ii nil, Ij. P.t!mi,. f. [iN~i1U, J. ~r:;_ Clttm. ~7., 31.65' ~200z~.

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Il"'~ l;;:a(llhcri~l ~1.1,lliI:ryol"l~(:d~YC~Opiitil";lll. c~idh,.;rin ~ pro'o':id~ ~h~ basis ll'or (_~~.l11 SiOr1jn~. tlggrog:3Jtiol.l. mtdl tissac l1l{!rpllog,~:]1{~~i['; U-J). Pmtl.llIl.11l.1iY, eadheriss provjtk:cd.l~w-t."eU ;!d~k"5i.Olll
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Th,1i'! mn<l~ ~yn@vil!mf'l rOfllllS ,8 ll1l,emblrall'lE:! at the edgl!s 'o~' ioilllt5 and previdles IlLJbr;(alio:FI and m rr11il ~riel'its -r~H the c.m-Halle. In rh eIU m1lJl(io]dOJ rthritis., the sYin.OIVilllll1!s the si~e 0'1 ]nflammt<lllicm, and! i~ i pauill:i,P<!tesin an org:aruize,d th.'>!JIiI! IreSpClf'l.~@ that dal'ltlilg~ <anila.!le a'rl~ bmli.e. '!Nfl' iden~r;ecl {adlhenftl-l] ,alS,e'~&enli!l3iI. fo [ .!h e del,lelo\ll'melilt ,af 'ttl eSlI'l'Ioviu m. Cadlheril'l-ll-d'efidel1il IiIlic:€ ha:l,\!1l a nYPoj}l.astk S>Yl'lovii'lll.ining, di~plaYill discng<ll1it'ed ~Y1FlJololila'lr~i3,cliOil to, ililfl.am n1 allton, alntd are l
F~5i5W to ~nriatmllll1~ltor~;Hthri~i~> nl O(lIil~rrifn~1] therap~nJth:s pr~~at <lnd re(jul(~ ~rtlilriti5' mod ~$. 'TIiIIJ!;,~lf1l(lvi<ll ("adheri n":ili ~'!ll~!i!ml1i Ul~ iJernaliiillF!)[ sYrlQovlall.!l:eRs in U~~h 1lf!:S pr\Ql];nf~~mitr1!!1tdI11 -!lllldl d€!~tr~~~iI,Ni!' lii~ue rlE$pm1seilnl ~nfl~lntm<lOOFy .D1,r~lr~U~,. ijn m:O Ulse

~i~Ul~' inl.i:£lrily .1lld IInt:ili1.OC1!lrc U. '/~. KIlIDWill;g nl:H.~$C ;Ji[,13\'~i~ (Ir C'dclJlIJrin prolCi:rt>, we IIIYlx)lh~i l>iF.l~'(llh~Jt;j1ll1~ndh}'llla~C'mUlcrill mtly limcti.on to rJ."O"'OCI~ (lei ~acIli~"5jonto O'WI1l1n: 1h..: :')'IlltWnmJI
1II1i1tilJlrul.t!_

conm.i!1 'nvc il1lumlluogJohl!l~il~ (ig) ~ikc cx~rnC'dllr~iku:fc.:tdh.eril~} dQma~t.ls;, l'igi.ditic(1 3S

·~(~h~.'~~!~"LIjh.C$. ~n(ik.O!']Ic.:-';.' a ra.mi;(lll are. II}, '01 IIHc'gro~mc:mhrnllc 'rnrotcnt~ lhal

cMended ell ~in by Imcrr:IIl)n1oin Cil[,c.il.il~l L~~nd1-It.'Vccwcd in (i." .?}J, ('adhcrin~mooi~!:C cel'l"e':Ie~~I~oo, w[1lq~bulfild 1;l~}1!illly tQ~ll!i .. wor~, lular ,i)dgl,C$~Nl b~i bJ~il1~ a (Jahe-!!!1 ,or IJu:' flo ~i1'M!ta~~po!id~~Iil5bo~d ~. i!~dr~~ed.. E:.rna~: ~,,!lil1Ctype on an adjacem cell vi~tUnicmcti(irl.~ 0\( IJlbrern!l~'®rit5.hwhJli!ali\l'avd.edu
aUl

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Sii'k)ll)-l<lu.. yO'jlmd;i~tmnmil.d1~ IKyoto 6ll6,9501,litpiirn,. lfiIK~N Cem!e;rfar [)we[(J[)melltill BioImg)!', Miria!ojj!1'!ilMil1<1mlllJ13i[ni. (~IKr'm [{(1~ 65(1-0047. ]~Pi!lI,

Ifl{!~~l1mMI {Ij' Mool~n~ ~lId flWi~I.f.\~01' RJI~Wi1MloLOQY. 'Inln1:wo1ag~ iIII!!I.I'!A~~'!Jj!'. IBriglliiml and ifI1om,1lfn IIiOlipital, lli:iw.lIrd MI!d1~~L S(hocL, 1 ~trmm\!, IF~lId W1!J1. Bo~1.OO(. l'!&!l. o~n;,iJ5P,.~j)ep;iflmll!li1t 01 Cell m I!ll!'!'elOPllillM<ll. 'Bioogv, Gradw!e StooQ! of Bio~iIIi;[ie~ KyQIi!1 Urri'ilel11t'}'"

16 FEBRUARY 2007

VOL 315

SClIIENIOE

\i'V\MIV.sdencemag.orgl

REPORTS
consists II ~~llingl~(\f1Cr e ,oon filbrob].[l~u :!iOOi'llllCrophfJ,gCS 1.111 (lVi:.Irat lie.'> a :>~~bllilll ~fLycr (]If leesc oonncrnlrvc li:iSUc, n,g l1:>ullg blurmnohi,5IQCh~rnkf~ sminili~.w"C lom~(Ii '~hf!ilthe ~tllldcil:;~-d Ilni II,g lay,;;r 'u.f sy~lOvi.oey~e,,~ 1111hC'llhli"ly mice e.l!p:I~8~"1i cad bcrin-ll (F i!!!,. 1, i\ and til ln 11~1"[IIl or Illie~ tlmt (l!cwJlop!'l SPQll!tmllJotliS :U.llnilll1lilullC •.nllhril:icll {!0t!J,;.N} (5. 6) • .UIImkcoo cadhcrhr-Ll 'axplm;sion W1Il'> ~OOIl '~m cc:H~ wil.~1 fil:noMm;D Illiorpholbgy III lim :'iyrl·
'iii}' ~i(wi~Il'll

The

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.Ihl~ng[lll:{~ill the 1'i.Yl1o'l'ill~ P,UlfItll,L". <l. P~~~1" outgroW!.!l sY~l)[lvhll IU&'HIC dcriv(.Id rmlfl~ libr"il1IlII\ll'i~·~i~,q ~yuuot\'if.lGyl~ (.Ft.S) l, (' and ID)" P.1r1.iCl,lblrlly inr~m,e :'ilauni.ng of il;ildlili:!rin~B w~~ ,aho Ol!ltc-J hi FLS ~,~'ilhll~!hc.rmnllU1il adjaQQ'ru.lo bone ~m;l 'C<lfl.i~agc_

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or

cn!l.

'['0 dlJ!fim!'~vhk'll ~YI!Ovla1 cellnd:lll" HU~·;l!lle." (:xp~ss cadhCl'i]li"~ L we- used m~ll~kolof cyto~l\l&

'OIflCM'icSlairllrlg o~ ~hiy di~fJ.ggfC!.!ffll!.'t.l j:~:i'iI '~i~\lC.."" li:l,ill,!!l a n.:wly ~'I~i1~ H~0I1OC!o.ltil~


cadh(.:r1u·11 (rlll~i,cfLdl••:riu(fL ili'll.li'kCr of bOlle liI'Iurww-dcrilv~'t1 ol.':ll~)c. Co:rl~isLiCIII.willh :it1> expression OJ'l~hc F!LS lillC:lg..: ill h!lU~1JilU~ 11), we o~r;;"r\,\Ctl Iilllil CRdl:lCrill·'I 1 ic.knti:I:~I;."s. a s)'rlJo ovi{dpOljIJllm i,(111 d iSI.incI, (ro~nUUII ide:rli:1fied by CD4-:5 ~Filg, i.1E fUtu [F), lli,,"<-iio: ~m1l ill~'~ were 'ctm,i'irunoo hwell!.:miealmy by lluon;I>I;!I.':I'Lt)1;.:Jt:'livHLl.1tl ttU :>Ullin!:{,MCS) ~:mhjiion 01"(,D4:5 'Or ~:3uhl'1T"ili- i l-~p1t.';S"iIlJ,l; 5'l11ljJoO;I)llir~inl'i:1."'!es,1_. (l11l] hlnlnill!l, ,or 'iJl:'{l!>,'C' Sfi!:Dod eell ~ d~JllOl')ilml.Qd UO dCL("(:!~hli!: cadhCi!in~lI cry,:;pfeii:;:iQl~ the on CD4S" AAI1:ipOr'Lllblliml(r:j~. 1(1)- Prevkll.!ls !lJlu{~i,-,~ h1b\1IC' denl(lIl~I~i111.OO tll<lI ~~bm)ln;;.m Jini~~ th,-, ::;;Yllm~ U~] ,-,:>:.p~~ CUSS (i'i} ~~~ mnt' ~1l<Ily:'i~3" :Ilulibody tnV\b)

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oc<ldheri ~I"II + ccl:is i!} ,wiinriik ;""fiiI1L~(jig, s.,n , C:tdllcrin-II C'!l:p~",m.m rm cll.l~l;Ired FILS. wi.h ~he lJ,;<,C of coli ~llrfhce bi:01inyI1l1[a!l an",:1i!n~t\lnflrn~OCill':jl"lOn willl al")!i"~U'!erf!l II" !"C'I;\C'<Ikd!he' "il!k~fli..~cd p:mmc:in SFilCci:es nli nnnle>clJl~"r wejgh! ~. .ufIln from wil(HypcFl s btu nQtIi.'Q~n FLi) w d,.TiH'd :l:ron1 {:'iKlill'l;i~I·ll-di",'fici/;;I;11 I~lioc (Fig. I, Ii to J.I W} h~ ~ddili:tm. b";'C~~I~ C3d'I:Ll.;liirl,~ !i:-:;.ue!lUIO: willi t..)"Luplill!>mit ~-<-.Jk:lJlil1" we p..;r· l:t:Jmtcd, :l;ilLlhicili.!or illl]lim~o:I:ILlor!.';S.tolmio: :Initro~ cup}' ~lIlclIDu:rKI Ot1~tK."uLiz;::uiml o~'(:1J(iili",'uin-11wid! ~-C:;JjL'Cllill iUl ;n.llih':l'Cil~ jlll"lCllimJs. sp~"Cil~C'.Jny !.II ;:;~lC1; of ecn .. .ocJi C<mlltcl m.~ m W(:l1] lFiLS (fig, '~. K to M), By LiDtt:IJ~b."" ul' <Icctl-uo-5'LI'b..>;Ir.n,.; lIaoc:;iml oi!$llIlUY O}. Ihi." !llJi.Li,ly (If r.m..:s ,~k~-i'iA.lcl O from cil1hl~r wil~cl-l)lflC ormd~lcrin ...~ ~ -dd~.ctcm mice wm, 3$~iS(\cl tOr bindilill '[0 n:::oom'bnrtal:ll cadhcriiul·:l1 Fe, \i\lild.typc' FLS t1clmJI1:;;Ir:!tlil~ rohL~I. ndIK':l>iQHGO t·acllflllr.ia.l,-Il. fe. whereas

<-T)S;l

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Oil

,~I.!O"~n<l!r or h

IF~g. 1. Syno1.riialfiIJrol!ll!a~t!; t'lI!Jlr~h.ui"[lio"i3lC!ldltenr'\'·11. fA and m INormal :syoov!ulIIlsffiiFled ~'i!lllheither (omJ'o~ (A), or Ol'f!ti-cadherin-ll!. ~B) arn,t1~ra. Ma~.l1Iiill!aljOi'l", >6150;.:;"~C:a:n:dl lJ ,Minto!: KlE<lIiN~fl'l(l1luum.~~lfio~MI [ pamlUl5, in,vadin.lJ carlil~~ staii!led ~~tl1eih~ (Q,r1ItM (C} OITaJFlltn.ea.dh.emn.11 ([))Qlilt~r.a; t :s61~~~. nl~r~ is ~11~ellS~~J;!UuIaI' fE'.activilyi3,t tihe bl:)fleI' IMnlil!.!~ imed;;rc,ll'. p, prJrmu£; C, C<!!njll1ge~ bOFle. 1!l'~,l'Iiri:tatioo, .!lOOf)(.(E [B., alnol F) Cyto'ih.JOrmtlwic 5Jta~n~ftg of drr.aggreg,al\e~ atlli'lrlitLt.l:tlil'ltIiS5!!lf. (26) stained '1.ffiId"l nsoil'y~ ro.ritrols (E) (ilr al1lli·cadherin-U and a,l'Iru·(]I)4S id'elfllirii13 distilird ~Uui~J 5!IIbpopuli1lthlniSili!joini l;s,s,\!u~s. (6) Siodil!1lmn~l -.' 1;IIi'i'l1IiiI ana~is Qr Fj!!,CS-sal1~ :r.y,no~ilil '~UilIpo;pllhniOl'ls. Dis,[)elSed <!I'llorirtk i~int tiMu:es ~'i'@;re labeled wiMil e<1ther illFi,tHlJ4·S m antil·Gltll'lerliil-11.a_n.dl Sfil'ited ~ means o~ FA.(5, 1::)!5a,tesoJll'iese ~u'IJp(lpufa~loIfl5allil!l p:res.orled ~,e& IO'I\erE Gin.aLped b~' Wen~m btot. 'ro~ ,e:adllie"filh1!t amd 'g~e~a~detl,yJjjiil'-3"JP;1to$pIlale d~h~roge~~ {GAPDH)~.pr~l;:!;i\;lrn"Thl!nl ollrl ~b~€fiH;~ d~t~di3bte (~d~ril1-DJ.~W~sij~J1! i'll (045.~ IJIf expffiS~il1l~ce~_ts" and U CytoflltOl'Omelrk :5~il'lll"1g af ru~ru!l'~; ns. F1LSd,e,rtrecl hom !O'i'i~d4We< ,~~~ (HI {'Ijlffl and[ [<tdh~rin'llL~l!U ,~Umic;e a!l!d ll1!~ged tjJ,oe'~~meswere stal!1!ed with an .m~i·(i!.dh,,@rifl=o:flli!1_Ab {glr<l1j oCm:lh~rin ..11 dcl1dcll.t FLS!'ai~li)dqo <tdh~~ mn, Is,ol)lpe OOl1tro1(dadiled I~ne), an ai'i~lloorty W ma,~®r lrIiSl~oonp~t~bilclly ~OIinptex da~~I (m~d ~liIe). (f'~g. ~ and fig, Silt, 'lnu'$C s~lldiCjl demonN (]) !!l!IrooJ1!~pr,~~ip'lto!1I!iQ'I1 ~~~~n1rr-::!J. ffl;lJT) ualttl!Plilg fiLS, @~ ,j!l1dCo1lghl~,~nl-lJ!...-f!I.!!Ufl;5, wepe ~rr,w~ slrmc ,,~dhcr:ih-i~ c.."i,prCSS~OII in, nnmril111;l :r;Yllov.illiotinytalN, lyse~. 1I!ittlimmllrl.'Opredpitaled \'lith imt1jJe ~Oinll'o'l ~g (lgGl) oral'i,fi-Gl~l!ie..rii'HIIJ (l5Ii1an.d .. n\:l!'Qh!;'!$"'''''. i~~ekli:;.;i!i".;nion of,c<ldh~'ill-I! 1('1 513:2:H5),(lUoM) AS ,e!!:ll~esscatihierin·Uilil adh@rem j;uru::liolils.M fLS,were Iii~@'dl sranned wi!ltl tml;e ! am;1 ..d~ler'C'i!~ juacfions, and !iDe role of cadherin-l! (a;tllleriI't""U (00 anti i3ii'l'tfF[Hatelililii m <1.1). fIiIe (~~!Xalfzalfon of ca[lherin"llL and j3-ute1illn WW, in h01.nl'l<phi Ilie .~_dhe5iklj'l by synCJ;\' ia~ fillf\L1!L~I:~$I!:>, 1r!l~~g~di!m!!lrl1!t sites Qf '~tAW~arijrrterir~~ ls ~FI ~ ~~iterJl! "!mS,1~~lill ~'fl:lih tllat \lit ad h~~n~ jUl1ctiOf!S<... The Ulmc~ion<!!1OOfliri~"lliOlt ot" ciiirlhcl'jn-ll, Mag~mrntiol1i, ]{lIQ(Ix, (N) Ca.dhllfl~·U mooi31tes hoon.op-hiUk <l1liI1l>e5<10n FI[5"WTand ca,cIli~rin·n-nI.lU i~ ·[(I!i\.)'!1O\'~:!1flhroll~~..-! Ol]JIlJlitr~tocm illl S,}'llOvitl~ !is~LS- WM!' a ~.elM!d lO adhere to' :5UeSlr.at'll ead h:enFl'-ll~P,cilli31 (el~-itO":R!!~tril;t.!!' ad~esi(l!na,s.say CUI). The ~ues w<!s ~~Xl c:>:.pl()~1. \'MllC~1 oom~.red to wildL Ito:m.oplilHic atll1~'ilon of WT H.S. to, Gadherin·1lI_...f( 'is ab~fltt in catilileriiiHLlL--'i'lLlIlIM. trlFOf !Jars: In.dlc.al;e SEM, '~ypc s¥IH')\Iilin~. c~L(lllerill-1~ ([dklc~l~ $ym}V"1ij'n L

aoo

REPORTS
VC'illcd ,[I. ntcl!SlU'Ilbk res~SIm~~' m '~llilliCi"d ar[lulllS ill cool1crhHl J1LIU miee {fi,g. 3. A fHid ti,orl all(lr',.~«~uoc<d mCl'l'lbn:.uc laid::; (F~g,. ::!. A to Ill. IrtdCl.."tI. pooJcd elmo 'lOr IiItC e.lhlic~lla1dcx I'i,'()tll these rniec denllOn:Slnuod an average C)" i\. p.dmfLify r~IUC:IPon, of f lSI Slito(; eli.lr..~;W% wdm:t~O<~1 dini~1L1t 'lrlhri~Ic !1CI,ivil), in~1 of '!J[illiOtl or c:1>llftlceUl!Iklr rmnri", lEC1'>'~), U£~ug ('adhr:rin~~ ~ null mice. ]h:i~ 1'!.."'{I.Lm:li:on .. ~lIgg~~ Ma8l,;!lU~\ ~a:iehro:l"lu: s~aiin 1,0 I.liMh~igh~ lEeM, ·~wr6ulml I.h:n 11K:! d.'cn,<;)(1 EC .! or l.h~ ~!llImtd:.d ~hat C8i1JlCrllI·I~. ,~."'p],m3cdon l'iibro'bhb~l~ IC"dllB su b.. ;,Imui:J~~y < jrll]ulCl'J1{lL) ,~'OC', S('~I~rily of p':lIlmlogy lil1i:u!ll, npp,:m:ml :i.r! ,.~rUI!d"lY~ IlIllt'\[!, w,3s.mmkoo)', in m1.hl1lli£. Iy aUI:.IlL1att1cl, cadberin-lt nu~~.mice Wig. 2, in J\~~irl rhrum[llo]dm~lhril;is, mil nine models of D and E), 'Thus, we OOllCll!ld'c!lli:n c<lflhenin· I! hi in 1l:IIlU1mtol:)'<tl1hritl~ dl!!ip'J,ay ~~},!JCrpl~t\ia.;)lnd , llIoc~~ary rO'fcel:ilJhr or:g<l!lliznirm. C0l1ap13Dlion, of syn .. <l.!Ild IIlm,dx. c~ab(jlr;!l~ of the ~Y'llovi(l~ tlill~ng {)!II!gmwrh nr symwiall .~issLl~ ~11l~ lnn ovial 11bmblas!~ tl)1lJ]I'I\~ t!:-i9.lc}. Ttim ~}3UlUlll~ ill \+\10" '['0 f~n1!~H)roonl'il1n dl(~ ml~ ~r C1l(ij1crtll1··11 in lb~I..cI~ 10 .md r.nig,,;(t('!:,'l'cwtCr oon~ ~11~lc:Ji11i~lgC' ::<ynM~:d I:L""l,Ic f:iI·l~;al!!i:-:<I!tirnl we li,.;,od .". ~hro~ ~lll'fa~, loc.;llll)1 .illlli!1:ding ,m!i:l d~lm'l'i!.lgl1lC!~ •. !l1"1.cll!l~ (1trll. EXiilmiU.lIlinn or lnnaJll~ joim dim!)'!ll:iliOIll~J (.3D) '''yno\'j,l'IJmk:rconna.,''5 olga!! ~i~~C' :llrl~itcC1J1U~'(l fJflmw.ikl>lypc~nd cadlmrinculnue ~}~'lC'nl in vhm (ng. 'S3A) Ull Sinfll~ II ,nu~11mire clcm.o.l'l'OJln:allXi tlll,lllh~ -.YIl'leal (Jon· c~IIs;Ll"'1lCnsifm!'i, oil' wi~d·l}'~"lCiTI:nd c.1dlilc:rin-1I. J~l~cd hy~j!(;Il}~asli(':s.ynC"wiiai Hnin(!,; archi~,;cll!lre fluitFiL.Swcre dispom¢d, in, a S!lIlrG.@iUC E>C.WI, ill iii 111 fllllmiUGry an Hllfl.lis I:;' ~mr.l;;"';~)' f1b~l.:Itt in and :,i}'llovrocy t~ cfllmC"i:Ly li.lr ~~ lfIing liI,}'I,,'f rbrnm· 'iil..m W11;i::1Qi."~~Ii!l.l \I\."ikl.i~rp..:- FLS ,,;:;;~b:li:;ll a C{HH'u,::nlll"U -u.;;:-liciai,1 gllic~ f Pig, J, c !1m 0), c(lllidciis~"tI l:ililll~~llike );I1lt\k;iill:\:[lIq, th,;: ~iUI!nN:' R~lht;r., iltc :sy.fIDvi~'LI areHlil.ce~m;cb~ cadli.~ttll"~ ~ ~C ~noolll linlocrf~lecw:inh I1lurkctl ~~nililnl)' 10 111m tlk:fi,ei~IH.IlillCC: ~'r~" i1i~O~te an'll 1OotlSt:ly nrwmi!t...cd~ seen ill 11Il:'~yua~,'i:l~ l iili~tg :II. ch~ syn()~ ii::d ~lluid TIlC 3D mnerol:l1;],.o;.. urg;1JI cuUUI'C :J}rm'idl:." ,t1I, .. intcr,lilc~ 1]11 \.' i\'Q (fig,. 1f}. In ~'Om.r:!I:i~" (n'l1~tCrin· mClll1S of fl!l.tal)'~llg SC'liXlJ.."d f~lClO[1'hm mod· [ ll~tt1C sYII('wial lillr,o~~3$~ lilIIl.t1Lon in a $in~p~i IjM ~ ~ nuH fLS mieroJl~81S:>tlllllin:-:. ~i'lU,ctU todevclop lin~1 ''!)nltlcil$[llaOrl Or (1(!'Ilnpft~lnc!'1f1 r.fig, big Irwde~ olt' !Jll.':(1OtllP to;;." ;:iY,I]OVJf~i micsocnviron1liI~1lI. Wild-Iyp<.;,uu:l cadlLi;rill·tll-mllllll ,i>JI'JIJvial 2G). l'h1l:';, 1~li:> in vill.u mood tlf.i>Y:UloYi.allli:;;.'1u,,; micmmass (I'!lUIll C!ll~llll!l)""S ";I.1,;;i:i il1.lill*l~~.,J, W : willa l!im~tion oou'l:illillll~i~ml li~ling 11ry,,;rt'{mlpm~tocm is lNF. a pm~iit 1!l1fl:II1'iifli;U[{)I~J cl'lokill"; ~ljll[illt.l1l~Jk.-d !ill ililh..:rt.'ll1. 1~1.I1!Un:Of~)'iIlD ...imL~ [ibroMlI,,'>t ib!.i'hrw· ill rh(,'l,ll'll1Maid mrthrilk ~Il OQ'I'i[rOl~ 10 dlC' iO!'~h!li re~ilCS on c .. dhC'l'i~l" 1.1. W~ I1€X~cx,nuincd the role of c-"l(~h('r:illl·ll I~atk ~~~llOIslmC'HIICl~ha!lgc./O in wi.[{lJ.,typc Cll~" C hllms, which .<ifC c:hi:u:oc,1J[ori:;o..ooby stlrr~e.e ~in ing illl S}'novi.all tfihLTIbl!asl. ehavlor un the KfBxN b ocHlIlar a~~mlll~atinl1 and ot:l l1{icns;:ui,oDI. nUl)' :;;efum~li.l!asr(:r model of J~I"~Oinl!'llun{) n<llllr i.u 1In~~(,]I)' 'Irlhl'i~is.l:u~Hlii~ nQndel. p~~sivc H<lnsr~' IlHl(icSI (:hal~gC!'i, WC'I:C ,~rll'l!'enu ill CR(D'!.ertn-I~. (U~fk~cn! ~}'111(W~! !1l~~ID!11~!l: ~'11!! qll~'~I~ of <1r!:hri!og~,.l:k: ;t!l!w;un~.ip(ld it;:~ i!s ;[I. d 's! ..~ (Fi.g. J,E ,1l1J F). ::<yJ.~ln~ric cfi111h'c Ilolyanilr.his: ~,\[l~h p~J!ilml!C1g}' The n;!~tl!B t~llII:';f1l!' $lllg,_~i'i·I.«~ c<!dl~ri~I-1~ dm~ Uted~l(i. ~lY i!11mUJ;iC -d ... ~1I'!f'i~~lhI'!!l1.1Ilcm.oo!11fik:useful d~c!"IX"Hlk tm~C'~. UICllIl aC1i,wllti'm, nlycloid IklClI~C (icl~..C't~..'mf<m" migbt !be apllIcmiaUy \ ~~d ;!l. ca;;OR'ri~l" 1 Fe ~i~~ln 1 ;jHld 'ci~bo:r:~uio!l of cyl,Ok.i!~cs:inH,"rI~LI~drHi!.nd To c:.,;mnl!~~~1.is.1I;'e PffilCiFl (l!nmni.C<tdl~crilt III U1Ah.. SiIi11~~!;Ir to ~\lImr necrosis. f:actfll" {TNP} (;i., rS" 11 1.5), ~hc c,jil_dhcri rl-l t n~ n phen(lIype. ·(l,(8nirlistfOl.t\dUlil1listlnttoJl of .~tlhrilogcnic iKJB,.,;:N 1um It'!iCl! dCiilK~lSlrmcdmm"cd hyroIJk~<;;iil ontil;; sy.rLO\i·ifll I III I.ng. ,:~i1illl decreased s}.'l1l(}Via~ I Ul1llUg O()U~INlC:-

Diemof e:lt~lc:r rell.g~1t1 m]lC~~ul:m~>dK/BxN serum mlln~l$fcrartllri~l$ ;[tCII'''''i~y ill wil.\1!WI'I!Z nll~e U:IIg. J. G ami U)!, To C;I(fJillllae :I role [OT C[lclhc:r.iui= U
in UllJ,;."Pinl!;liil~l~nu~lfll(),~

drliJ KtB:'\N :;,crum tr:m~~1r model o.f ITI.1hntis is mpid ami .,.d 1'·1Lmih]d. WL~1J1S(;"(1a. \,ITIi~l~o.1lof dili~Dll.ockl. adlnil1:isll"l:ring .. lthri~ng>~~jr ocnnn week Iy 100 drive p!}'1'SiS~CI1t~YI]Q\'iti[!i; {fig. 30]). ,\Ilcu[lJI.m"illg aI lemn 1: wccl;:~ of .~.t1.ivcariJrll'ili~ we admill~slm",,-dI mlUi..c'Ld~1l1rt:!l.II 01' imryper;:Olltml. In;'mmLmL~ rID' 1~pcr.liOOQj" I {I O(lY8 ;;]119 CKalI'l~llo;l :Ji1'1~1I11.is, dinkal r:J..",!;;~'IOI1:;;.c;~fig .. S, I. and JI, Si!milar~o our ob{oliJ!"\l,Jtimas. inp~voCl1~.· ing ,~el,ltemlilirili:.1. we ~Ind that ,m~i~<ldh~~ll II di'lpbYl' nn;l~!mc <U!:llclirn:',MioM oW~""L'bI:ishcri ;Iltll~iI~. IIH".~ oHl~CI'\."L~i('l~~~. provide lI;l\ludawc tillal. ~,(!~IIttCri~l~ I, COllUib1l.tffi~{I tile 1'C!_llJ!'lIirlHof I arl~l.ri1~c ~i$Sll'C ~'C$POfl$CS lin, both acuteand chroni~ ;:>yno~ itis, The ~Lbiili,IY (If F:Ui i!l! migmh: !lnd invthi1c u~lro~~1il ECM i:i: 1I jX'Ilh:i:ui~lIy ~l1tpMmil, ~b~= ture Uni~$UC:~lbOd,;l:i 111m, ronns IIlt~ hy;pcr'phl!.'>lilc lini:ng and ~. cooliJIP~i:sill!';;"j;, . ero:;.iorl JIfIW cRruj,Il\t!~. (';:ulllC.rin:s and espec ial ~y 1~1.c'S";lleli.}'-

hs f1bi~:iIY rl."{hIQCe:!01~ll~lisll~(j w di.~IS,., :BOrull~

3nhrliLus,

\~'C

3:>Se."'ii.-..J

i'~

am·

nml c~dhcrilll$,~il;:c CIldl:1erri~1I·1L lire knoW.I'1 OCI inHlu(.;nec c(.;1j migrml:t!'.I:1;!J.nd ~IfIViIS~OIli (l7}, To !§tl illl fun'lll,::r irl:iiug.lnullt~1 til,.; 'Ct!d ~t!,:ein- ~ ~ 1"Cg_t1ltilioll .FlS FUllcli,!'";)n, W~ Pii:lTili.ltlI"Iloo migrution audinvasiou i~~:rl:l Ihroll~h ECM-t(l['IJt ..1:I tmLm\'C! f&. AithmlgHI C'adhenl]" ~ ~.~1lI1.lRS~is:"

or

~n~

1~lilY ~iJni b1r Hlllfal llli~[)ry (tcLl",i~.ytc ~l~ of wltlcl.rYI'iIC! ct"l~ ~file.ir illducihl'c Il1jgrnlf.ll)l 'c;lp<lc!ly \\'<L~ <IlutCll~"lhXi lfig:,. 4AJ. 1S'::<tllnfJl»!inn of plnl,dct.d:-dvcdgm'\f~.l (1,01:01' WOOF~ \"lil,ll\llm~ SCM ilfl'\,~ii\le C<Ill;l<;1~}' !1G\'~.~l <11 d~111<t!k (~ri.t;;il iH

aoo

caclheriu-~ ~ ml~~ PtE, wh iel. d i",pliayO<i m~~y -15% i.1Irnll~' il'l'i:~shiC octh il) (If \\f~!d.... '~ 0:111$ } (Fig_!. 4[:l:), The,o,c ,il~O!~it.1ns. Pmlllr1iOO us ICI c".aml~C' 'C:<Ui!ill~c <H'KIl'll!ll1e ~'i,i'on in .~~ inIlmml1atrny AAhci~isHil{l(tel in \' ~,,'O. I!~clce(l :>:)11'1O\liilJ lisstlc aUi;tch~1C~li to. ~nlgmtron m-er. and

..
:S(l~jgns.

,I

..
I..,

Fig- ,2. 'cadl1fflil-U ~llilalM S'yl1ovial tiilii:i[J a~(liIile(llIFe'" '(1\ :md B} IH'eiil1atm:yl., nd e!OOin(H&t~-st<lilled a '!;If s,yno. from Wir ,tA) ~d ,~~thtrilil-U-lllU~ (iii) rmLl)~,The h~J;ll.ar.itk S}\~'o!iaIl tinio!:! i'IiI \OtdhffiFlU-f1tJl~ mice iJ;[iks ilihe d'lara(ll)eristk SJ{flo\liiai ~ilillgl ceUular oomptl;rlion (llo, 3 (el~~l.hJid) Hlil~i~ oE!vieeml 'illl th~ WI' 5<y.novUtJlm•. IL, !i.lfi1oml ~ni~; 51, ~wal Si.libl:lllIin!i\ Magnif~biOtl,~OO:X. OC an.;i D) QidhE!Jiln"U
~,

..

..

,ca.dl'lernrHIl1~l ml1re-at1. ~OliI'1pared to Ihooe ~f !be IIlffi . ill\i!gJilfu.l~fl... ZOOO-;:,(E)5ynil!'!iilllijnhl~ ~I~ de'f!~ily.ILiliril'!g l~r ~el~ Pi<lf high tP'O\'$FIi~W (HAF) (100, ,""wR) '~~ enum!l;Fi!Jkdi I'll WI' {~quine~} ' • ~n(lt";lIdh~iiI- U-I'iUU {tJri~n9:l@s.} I'lfii(-e. IndilJidu.flll~l,i~t~ Ilri~o, ~iiI~ SIEM ijlf~ provig~I,. !1' '" 11. nlIice pergr(Jupi P <: Q,OOOtL '~IF 'GI Mklrnmi~ (]r~ n w~tuJ~ ;lind lining cemp.:lclion. WI (F) and ca.dh,~rin"ll-n!l!ll (G)lfLS, M!re p[aCied in micJ'om~ss ,org~1llwll:~lfe ~'er 21 days ell) dirudl were filied, ~ediol'l~d,.andstai!'led rNith H&E 1I'he il:elh.il\!ir (Ol1d~iilQtfi!Jlt <Ind iil1lll1lJ rmmaiMil behavior e\i\ldel'it inl WT FILS [aflre;!lilFl (f)] amll FIJlar~ly ,8Ill'efllIl8.ted llil Qldhenli'l"l1-iiUill rllS [allf'®W inl '@::i)] are strown. Mag!i1ific.at!rJ!l\, !\l00x. DOlt;:)1 tire rer;resent.lli,'l,re ,of fwein.d'.egl!':!'fLde:rn 'experimems. 1S;.tlw!1;JiIl ati'e

~ulatf':> ECWI. auitectu~ iFlWi\ll, M385M'S I(tl1no.l"I1e'~tains ~I' ~rl'OfiIl Wl1 (0 0100 'G3.dlileFilI'l-1I1-1i1um f[l) iiIla aiie'slocn.llll The ~bfJl:5¥lQViat celllllar llli1ing ~redI antrl m1Irted~y atlewa1.'ed ~oodemedl KM (Dllttie) In

1008

16 FEBRUARY 2007

VOL 315

SCitlENIOE

\IV'I,MIV.sdencemag.orgl

REFORTS
m.!hc-sion and CVI11P.1di.ou. as ....!!II as a C8IPf~iLY w,~roCh.!Di!EC'M ,(filg. :n. III tile co-nteM UIJIlamm;]!ool)l anhir&Lis, inl WhLth I.!ItC :>ynovlulLn lJU(ic:rgocs ofgolni;r:,cd ~lIlu~wlogLe changes, 11IlC$~ cm;hYlol!fl1 :lincm,~ Fl.:) i;:llJl.tniblllJ.: 10 til,.?,": dro· ~d.~~'tI,; ('j"lI'1ila~ ~C' {F~g. 41. C and D. blli!d: rnaticcih~g...~~l. Iht: s).'II[wia~ lilling (1.1 _:I'4),. Till ;:]nl:l\!'~l.C'adll). QI;I~I1I;i.fic!u~1o.1:Iby hi:.'iI.CIIllt:l!~phmllct. c:lldhmn&.~.~ dcrICimL mice, lih.!.lsylul!Vi~~1 ~inil:lg .ric methods COlll1nl"lC!rl::llH .~Oil..{,r-cdl!lcL~(jn in OInldi t1)'a1()vi~l ~I!IDm~smit".':u(l :IIp~n"il ~Hih.ill]C· cmilll~' '('3Tl')!o;licm eadherln-ll in del1cilml micc. HWdHy abc!!l"-;;]nl. am! ch:JOIic Wi!l- 3_ c !lIl:ld D). By eon1}1l11S01l.'l]l[)llC(>m!lion~. which are prinlLilrlilmk:rS!::mi.n.!l Ihe· mle of ,L:;;d~.CriD"..1111 in IiblfOo Ily di::pemlcillt on OSl.cocbst rillllctOCIil ( !8 JOI> Wt'I'"C .c..'~d~J1t 11. ~1 1!1T clio:§I'Ii'e ill ooth wi~d4YiJ)Crun] b~~t'i!(l'l'Illk:ipatioll in~hcc)rg,"nized padmJo!lk,ul 10 mi :i'iynovi:~. Itlling h}1Jerp~,,~b <[I'd P"'IIII1~1!,; ti:;8\I:C cadil~rirt·~ ~ mil]. m.i"oc (F.ig. ·4:E.)t. These ;nl~~.s ~~~lit t~a:n. 0.1 hmhhy mlee, d)0l"'[ll'4IcmioDl Oif em'! i.I mge irl in Ibl]]lllutmy anllria i~ c~~('j'irn·11 d:iro:lN the ~~~,th~i:o;!mH::lIl~ :rmlli~ and TIne data Mlgg,~~ 1.1'to.11 cadlmJ'l!l·l i~nIlUC;llc~ ~IOll~e or ~hc:Ii'Ol'l1il,11 '~ylln.\!ia~ ti!lg by c(jIJ,f~lf· bo!hntliC' d~gl\;'Cof ill fh~oll'llmtr:on (.Fijg .• .:;, A ;I~'K1 lin .ri:ngl11! RS 0lI1l ailmt)' !DRmll ,cell·,~cc~~ B 1Hld fi OlIlld IU and th<: o:rg~!]i~::cdp;trtllOl~i:c iitCfldlu:tifl...~ ~ deficicm. mlc~.'i,l!Jikl'lypcsyrlovilli pfllili U.'>eaased ~~II~I-thicl;:ncsl'cfll:1ililg,; erosion, \\'"~rol$ c;!I..cl]ta1rl·11 m.1I1~'»'ucw~alfliI"nnlu.~ $!Oppi..'d

inrva,'>iou illln car!il;lf,'C \".~ mmkJ.'tIiy dimiulished

or

re~pool1£e

artllrilo£;'Cnic .K!ll\N SCTUi:ll.[lIJUlSfc.ii U;;'ig. 4i. C UI'ild .IE~.In t1tcLnl1iitoi.d llr1hr.in.is.~~lc FL'S popI!llmi(lll clCl'llOI.·L'!!lffiLC'S .~. m~aboliaIJl~y active. ::')C'"Cl:CtcilllY ~11Ltc {,loS}. S~im:Lllragioll or IFLS een ~mlucc ~ccrclocmoJ a rnmlOO: or p.I."O]nn~urull~I~[:I~' ll~dil'l1.Om IIlmn .h~."'c; been IJo!'!ol'iI!U~81edloamplil)' ami IlillIJJe~I,I:JW "'yJ'KIlI'it~ .i.n:I.l'-mll]~1Iicm_ O!;ir fiml~ngs, MtggiI.·f.I. thai. m:Jhrn:nn·I~. til]nLJCIl~ fLS

or I1ItC ~ym:r.,;ifll[is·suC,'

in l'J...'SI){'II1SC m

paniciprurall ill 1Il1CS}'IIO'\iia~ i!l:fl~Jnl~m{Il)1enor lion. ull:MiJiti"cm to it£<; nul'e. in Ij~~lc I't1IMc!GC'Hl1g and Ca:I.1l~bgl.1 lnv1L,'iinn. These ~1~jgh1l'iidel!lnily pre\'i[l~~y 1,I0~pplll:;dalcd meoQl~niS"mtl g_ovml1ing Ihc ~1ICi~~.vim:or .Ile jt"lim liDli!lg ~md rI;lV(_'<I~ the C'Cllill':d! role ".~I1D\'lOllI eadhcriu in d~r.cf!!n;inil'lig IIrIC ~11t~\iif:lf or,.hc l'Cl1idC"nt ~lm'i~nvn"'_~ldIY·

or

JIiI@dulalre$ ~.r1~a~. '~Il~u;~ r~p(m~in arth.rmr>. fA. <ll1ld B) (adhel1l1r"lil-nuU mir.e are r5is:tallItlO illI~Ia"UnatW"f arlTInillls...man!!! (:adher:il1r-ll:-f'1ui~ mice LlreFI1 ililied!e~ wi'lh aFtlil~'l"O-9il!l1ic: 1{J13~~ ~Ili!J!ID", ~1iI~ diIilkal fn.d:~ 'W ~nl[i (iI1allgl's. W in ankle lhickml!>s (B}werl! reoorded

Fiig..;a, .. QI.dh.er.inrJll!

A·~
Ii !i:

'-&'~
.... ~Ii~

II

mblromfouf' ~m:Je:pelilden,t ~lOOOS) are s.llQll,l!1" {C (;I!1'dI .D~ :SYfllo'l!l1lJ1.a ~(l1h~t!Jjr·~ iFt lnllilmllll<rlor( ardlll'iti5.
{(adlternil·n~lin.l1D
1~:!;p~.riIiliI!!illS

,over 2 weel($ (6). IPvlill'~~ res!!lt1S ·rmm 24 (WIj of 2:6

.rp.

<:

rNi,a~n'~tridii riJmMt'lfneti alil~.lle ~ctioos fr®nlI .ar~:rili( m (C) <100 (a;tllterIl'l4i~Dlmt (D)' m~ce d~[I;!I)!'I$~,;r~ ~:Q~l Ulilin:gl 1i!~rp!~~!.iI b~d) .j!liId del'i~ [eM d!!,~jrnon (blue\! in M rn:~.111111 rnnllaM. "adh.:e n11'1·11-i1!iI~1 iflfltlme d ~issues demcm~liri!JliE! dltaotit:reUll!a~ or!)!llil!1a!tiofl~'{i'lhlril miFlifnllal [eM. a

·_f~

...... '~ng

:SVl'Ililvial.eJfusiC!nin .i@i!'lt (:ario.nl 40 Ox. (IE aM IF!

~~'J: is ~bel~~ •. liIia!;JJlIifirIFlS tlii1in9 !3JrMii'ec:tu!Fe in 'IIirr,!l, M (EI or udhenrn·n-n!lllll Of) as mrioU1OOld5~. organ (ulluFe'S tw we.re e'i:.pmed 1:0 INf (1:0 I1IgfDlll)

:1:'

!Ii

rI,

lor .2] d~vs-lhe ht1\h d~r'l;:e (i!f tilling oonden:::..ltiQIl .alil(:lmi'liinglla~-er ·r,eomnanooeliMent in m as (ru'is rnamadly .ab~rran[ in ~a,dhel1il'i,lJl-lll1.dl FILS (Fl.
41!lOx. Data al~ re'p'J~ta.tlve of lIui!!e i!'l'depelllder![ e..xp)erimel'ltis. (6 to J) ,cadhenn' :ti. biC!1lh~rtlIil"l'UJIilc ;}!1II!i~I;l'H!tehdlltlmfll~tory ~rIJiJr.il1s_ wr CS7ilU8 mkoe \l,itlfEl inj&ted 'Ivi~~arlhlri~Clgenk KlIB~IN5~rUlm ~~Il!Jl[!.tr~al'l!:llil[!~ in ~111.~ (~lrtj~(~1[i ~rild ~<G) ~rOCI ~~~.~ ~vithrontr,CllIg ~Q~e'fIl S"q;ul1!res.)<ll'lilJ !Nill '~it~er iCadhl'riin- J!'!L-fc WEt s.oUd tri:lri!@lesJl ilJr o1nliF <:adhe'J"in-U [(1fII), :lielidltriangtes.] .adm!Rlisi,erecl'IlJIth ~ Ilwdiln~ do~ (D.S nlI!!!} f{lllllcn~ by l~int~{I~~ do~ing (0.1 fIitgIl '!Ie'l'Y 4:8 heurs, Pomed resul~ frem e ]:8 (C.'!dh~rin!·ll='lFd~pdl 10 '-~lll~-C"ilQ;Ulleriin~J!JJ iIIlkeare ShOiffl'll rrom {ollir and l'I\'eoiiildepelilQ:h~if'lt ,~~p~~~.~ ~~pw.ij~ly. P <: Q,OO~. F'~r ~~IUn~~t :Slu:die-s [(0 ilnd 0)]. Ciootro1 I~i ~J)en s~lIIares} or .j! ntii'(1l!d nerill-lJl solid tf~al~g!les~ wel"e .a,m~fiI~~tf~d i®r' UI da)'1ialh.e~ al~cr«iI'I!J <It IlNst 2: Maglliiilil(.al]oo,

.!~ .~\!: t~

<Et-COiM
·.~lR

H .._

I
.0:=

I i
J

eCMn!! .... ~.n

mo.

lliM1
h ~

jD

'14

'I!,I~~~~

@ffaTlhr;1tk ;acti'l'ily. ~l,Ilts

two illldl:'p~ndellit ~enimerll!;S

.grolilp) and poo~ed lJie\ilbTnem fe£u:tr.s 0)'

~~rese;ntii!t'iw I;I~' (n '" 5 mice pel'


'~jI'j ..

-lEii-to;N;QI

10

_~""I~

*~ "~1~

mite ~iJergiiV.tijJ; P '" CLOrn:m<lFe SlieJi'iiil"ErliOT bilrs ill 00, ~En.(!Ild [(6) t~ (Jairld~C!te SEWt
II

I~~~

'II'\I'IIII!,I'.sdencemagl.or-g

SCIIENCE. VOL 315

16 FEIBRUAJRY 2007

1009

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