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Expe iment reproduced, discoveryerified

IMMUNOLOGY AWhl),Le-G~nlo:m~ A5,'SlOd,~U~11 Sit~dy (If ,M.ajor D'@t.e~mlil'l~lilts, f~r: Hbst (OPil troil (If HliV:-l



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Sc.II~Nelc VOl31]

20 JULY2007


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20 JULY2007

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With OI~E!r ••• :P!l,lr:S.(inl~lityTyp~5 Bl f1/oofdoffl ond R Gosfill'g MkI1DWled~ii'i~ lilat oti,e,r ~~p~~opera:~ d iilfE:~elit~lylihl~n you
wAn. malke Ial:lll11~mo~ prod,udive:.

:tj 1!i'~~'SPI~n,~VI!t; K~~~~lllgt~e ~ll(nn:ase) IP'alrty'Go~f1,g,i:! ;:;lhe1 ad~jl1:Qrproteii'll SLp·76

.~ S~P\./6 o"llilld K Dan(~ U ,~ Q. 'Qi {Jnd A Al1.!!,ust.

d10llna lileulll'a l1lidaptor

St'Ne'S 0lI5,mare

UlSzFrom the Ar(hi\le~W,nti!,!g a Wi~ li!i"'~1(I)Q(M IL~tt~r J. Borciwdr ILik~olln,~',~(lOOs~les: p~lC1~,-{llJf rover I.e'll~~SI1(lUldllll~cr~V.:Lt~ il:I1I~ y OiI~~mIU '((I w:~lilt to l'eaJ~n f'li'U)r~.

:~ during Tcell aclival.ioill. .;; 1I1'ERSPiECn\llEi Ch!:lm(llads.=Na~ig;.a,ttrr!g by .~M.!ll,tJ1pl!!! ~ Slg Pilall ftg pa~~w;3·Y5 ~ z E; P. J. M. Vt:UlH(1(;lsr,ert ,(JwW D. M. VeirJll(ffl .~ MUI[ti~'~$~gnollili!~ P~!Imt¥ Ipr{lm.o~e(,el~,!lilOl;reilil~I'I'I.~hroUJ9lh

I dlM1o,a~tIiaCli1.Je groHlienl .




M'M.sden c:emag.o f9


200JULY 2007




Otfalctolry IN,euron

Pre'CUIrSOr !Div'ersiity »
th~ braill 1~'~O\lidielS aJ st:eadY'5l!~ ply ofril'~wlVSli'll1erlli~ed Omhl.[tory lr1eltlw,oti!LWh~se (Eilts; .;;irrt! generatedlif! ~:he:S'l!lb·\I'e:rm~ritllJhilrZO!1eandl mij~raloo to the' ,ol.'f!1lctlCry Iblll~b. Me:r~le et {lit. (p. lin, pubLis:hoolo~lnlililE! '5 )1Uity) I'I~W :si~.QWthat 'oliff'er-ent re~li!()ns 'off the ~lll:wenlrkullair zOii1le g~\IIe rise to, dUfeiren~ ~Ylpes of' otfad(HlI' 'IiH~lIroI1IS. li~!J.ts;,
1"1 tihl£!adl1.!U:m oUJ~,

ind/iYid'lmlly v@rs,'lllit@(Ilnd tire bener ,(h~I~'fu::terize,dl sa smniliillg s jJIofrmt cd,tUfiI alre,m:lll' druel'se IPOpU'latio~l.

tl~e::ltem {elms Qf~he511b\l'e'n~~'C'I!l:~i1Ir ;t.'QnJe'alne!lot



Borrowing IPower
from Nature
Meth,ankil~ ta~~ ,a~ .a(mlil1l1li~hei:lin the oeQl d~fQf!,j)gha f!, array of 1l10~e':(I!d:ilrflf'!achfl'!~:S and lher'l1 has beemi illlle;r'e1illll e~plo'Hing tlilu) ,rrl[ldll~lIl-

~ilYin il rlilkia~ nil ~()iSc:a~(!,5tr'll(t~J\M, VaJfI de Vl
I'IEm'llcl i!11~dDeU.~r(p .. 3:B) lreviel;\l'the rN'fllt p:rogr@'~ (lUll Ith~ 11IS~,off ,Fol.ary a~d lin~arI1IlOl~r prroteim Ifot ti!~ks ~~!!:~ as, facilitatilllgIIFiH1!~I'!IClIr~ OJ p~werif!ga devic~, At~hol,lghsome d~\l'er .!lP!)U(C1,· LiQIIIS; halll!!! ll";'ol.wd!, lh'E! i3lU1lhDfS ITHlll! thall1llany

ra:{~ ol'lhe liI"ins!<q U~mlw~Sl3 r Attair., D:n@of In@' Ilright.e!>t ~a r!i]1li tlil!:! nig:1ll s,ky,. to a r~o~utio!l1 ,of d. mliRi~rc$.t:ol1id, Altillk is Iii llul~i!ia:l as U~pi 115 velry ~pldl.y,~a5l 'r;lnol,lg h ~ha~'~ 1 ,apfi'e'ar~ ,elon· gg le(i1l~l r(lil!i'gh C,(!!1ii,rihJg.1lll rQfCes. lh~ a mou!1t or di~ ~Dr~iol'l lhl1 at~eillda ntc;h:alll'ges in surr<l ce illild t~mp~~aMjUe~; dmraderlz:ing ilrn!!.l:ul~r mOrTII€l'!turn lran~pmt with in the S~,Ir, d i'llerge '~rom!he pJiooictrons ,of sia Il1Id ard f!~it)d~ls,~sp,~'da~ly 1IrOlm!:i the 9Cluator. TilliS.• @ldiFlI ~i'roc:@s~~, uch s .~~ di r:fe~ ~li.;lILrolaliofl .1lJ1da~termllive II) ~.!wulY
d.ankening [!3!Ii'II~, ar~ IIllallde!dkl 31!)]prorarl(E' oJ rrounin,g stars. exp'lain th@

dil1late. Tile 1~I.oge'n dilefJflls~ry I)~'1.1l!!ro~elil hf:gl'l f ~ati~url~ has p,mlfelil to. be partk:ul~rlyinter~tUliI~, not Iln~t II:i~(.au~~.or thE! ro~C''Ilf tlleSill' wtgiijEil ~ as h.arbilllgers of glcbal d1 llila'te !l)llange,. but a b~.uerr uderst~!Jdifl;g o:~i+l~t (~eimisti!'Y11l1l!i! be~l hg"~& PliMf!d Illy lild; o~'data. Sai'il·tgpe:z I~lal. ~~. 41n 3



at the' j:DrOOI~,",Clfl. p,ri!lciple :sta!:l,e.

Dating Differentiation
EucrU,es, (Ire lII'Ieiooriiecs tha.t baoe ig:neoo~ ,l(tM~y OlmSlilla U Ibedi!:S. :S~l'lila rte the' Mte-.roTl~ Ve~!:a,t:iil!rl,y in tine SQia rs,r.;lleiFll'S hislury. DalIlJil51then'! cen te~l !.is ~ b~l!it g;eophysica~ processes ~u work wn~n these bootes '!ffll!l-e dmeirl'JiI~iatill!l 10 form <I: mel.cillt( {{Ir'e ilUQd sil.icare ma!fUI~.H~fv.e;r,sliJ;ch ~lU!nl~IS 11'ii!~~ beemi diillkuU bf!(ilUSie ffiJ:r:ri)li8'" lend to Ibe dilangrl2d by ~tel' Iluralifl!g, ~lIIld 'fraclulringt. 1I1lid' a'l$~, the isew,pii!!: ~st!1fI15 ,a¥a~~abl! fOlr' diatilf19o"Jlreh1.trd l~ti3 nbraltE. Byalil,alY'll11l9~in:on:s wi Ihin eu eri 1.e5., Slrilrl~v.~san (p. 3115) have dialled their {~tQIl~fif,1l't~Q~ ~o'il!ii~hin ~ J.3, 1lI1iilli(l1'ly.e'~~5 I(i!~ !l1et~l-

Strai ned Relatiol1s
Wilen '~~llrn5re 91rQWIl om sm1fBce'S lfuirOIl91h a vapQr-~h.arse aep'05,iliQn. c'Qmj1l!i!\t; he~tir(;l~~rU(tm~li C.;I1Ill fQI1fIfI Ire~!llils.e'01 str.an 1II~lfllat Qlri5e ~Iuou:gh Ila~~(:e' rnf~m~ildles" Rl)!bil1!l$~:n €,,' (II. (p" 3;55) $Il(r,~ ifl 1!!iO~ution el:l;iirol!lment ~h<lt ~U! (lJmpr~)(5ulpedal· aces can ronn SIl(l!f!tiililI@O!!!s[ly inl ca:dl'!iliyrr~ S~llnd'~ n!ilnomds~tinmllQlItn [lte [oll1llrolle~ intrmll!Klion or ~i (alio~~, Alterl'h1llillg lalf~r$ of (ad Ilfiilldm 5-'l.I~. lver [kle and! ~H'I;'@r suUiil:le,lorm ,a,'longtfrtlt'! a:<isor ~hl! rend~ec;alil~~~he lla Ui!!:E! 1m i5.maltdh~trail'l th(!t hlJil~s up diJrillg $~i~!?ri nfiltral~lQn Ilimil.s tiile
glro.\!,!th of the w,I:r .suU~de

bleef~detected. nH~ ~p~li'lgl~fne 10 levers 'they roundl ilIirt! ~he h[ghes.t reporled iln)!'fllheore'ill the almo.spl1ere" an d 0;11'1 aplpare lilt S'f nergy beliveen 110 ~l1d ElrO~ugg~~~s!:I!il ~nkulQ'wV1l al.og~rN!di\l'a~Io1fl h i!IlI t'leJaallllisrn. llihe;se llevels of n"iogens abo c;;:IiIJse li~e rapJd ai!(ld~!,i:DI1iof d~ll1ielhyl SU lliide (lInd mer(~ryinJ MleAntan:tk OOUfidary laye:r~

Alllti3irclic b01mdl.uy t~r homJ![muClry .20011~O! F@bru,,1iY2{) 05. Tn~ 'Obs@iNl!ld hi'~h oonc;U!~raI.ion~ and p!N'~1SLll;Il(ll' ,Qlf ~l@$@' l1f1log !!lIiIs!.nroug'hOllt U;e slJlllil ~ri{!d, 'CO!'llrraryto e~peda'~jol'ls afild ill II1lnlk~ tile' ~~IlIt1~iWlinllhe ArC:l.k" litlher~ I a has n!)l,

~~e'wt !ll!e,arsurem!!'!m EI~O alffild or

~o the In

Gradually Becorninq Dominant



s.Hi<ate diif,~Ji!tiaJ~o~ 001 'the:ir P!i!iie'!iI~,body. Ttl~ 'It'l're abi.eto anchor the 5Ilm:r1I·lived ittf·W Isotope' s.ystem ~\Iith '~~il' s.W.1e.r II Pb '5.ysl.em lQ t~e dm.n the liming ac:uratt!ty.lalEr m~talilIDrphosiis o,ilhe I!l!crit!!':$, l(l~k IP~awe' a~l@r i!!f!®~'h~r 9 !fJliliiolilll€a~S firma WflS lik!!~y (tIlUSed by hetilli Il!;i [Forni 1lI1Piu'tS.

domains, U.e
cO!iltjol. OV~~

gmwlh a(h~~e~
lJ,y l(;Jaltll1l<QimlQ~tu~ oo~utiQl1I~!dra,rrle-

ql!efl!iO!!~ IltllV! bt~1I! r@laH.. @ly S(iillFl:@. Irmis el' lliil. (p" 358, :see the (I.'l~r} nOl'l

lers and


~5mo;, 'nlUS, ~50ITU~t a

il, I~(h,ros&il


~I a S pi 111 rI
Imagillg 'th~ SUl'ra{e~ ,of !>~;clrsothl1'u UlCln 'the Sun, LliQl,Jld ,aH~~~~~ro:n~me~ io 1lIl!llp'the physk-a[ p:r,ol!:ess;es I!'i'~rk Q!1 the 1'IiI, W~lh ii!dv"Iin(~ d optiat cal inteJife,r(lFlll~tlri~ I'llqu:e$, MQnlll~i~r et (l!l. tedn (p. 342, p!Jlbll~het:i ol'lline 3t M1I!Y; ~et the f.e'rSlpe((i'lt€' by QliIl[rrelillba;~h~iI1aJve fe'S(I~vedthe 5ildr-

'wiw dillllE!HSiam WiiS, lJi~edto tune lhll! n~a r· if!i~l'ared:e1mis.~'fM f~~l Il.ese lilaUll'Or,oos,

NIlW ,[I;1e!l:im dai~,ln9 1.0 tiw I!iIte iri<l~,oclti1,ai~ ilu~lIde~ bOIUh ~ii:'!osaurs::li'Qdth~~~epIHiai:'! Ill're~Ur~

Halo'Qlens in Antaf(t]Ca
Tr(lpo~p,herk l'biliogens. a ffedtihe



Oili(Dn;~.~;h!:1 Q:!iid~zing C1I!l",dty oflih!l: <Iitm01Jp nlll[e,

alld a,eli(l~(I~iorl1lwtiOH, aU of which are Ht'd:.1€(!10 1

I.he pr&llIrsor.s pE!f5~Sted mmh l(jfi9~f IWlaJF! bli!efi thOiJ~~U anel exiskid a lo:i'lg hild with diiITIN:1S1I1.I'1'S !lIillllpol1s· f ~r~, 'The~IQ:~ils f~r o s,l!lp,pOrl;:;l modlel O~;;II 9rad'ual ri~ of di I'losa,urs il'l 1:1111l!3~1l1ni<liS§k 1~1I!~ PNl'(tldl?d the~'r domino'llllt:!llily Ihe beg~nlliIl19I1)H~eMassl:(,


i~ ,~


R~ce!M ~dIol'QIf1(fSJill r.equudfl~ OOChilOlogy hav€ in(flN)$ed our J)ow~r to' sUlidy vollrialion Ii\iUin a sillg~~,organism,.C.~;'jlfk etoi. {po 338) f@S@qJililinli:@d 20 S~Ji1!UP1lS or Afabidopsj~rtiCJII(ma wil:i11~ljglil-ct~niSi,lY (lli'QOl'!mlleolid,e!' ,j3iUi!~i!llI!d f~:LII!nd exlefls;i\il~ \!';:l rialion" Tille cmnprreh l\l'Isi~ei Iilv,e,nLmy or ,~,enom!ll-1~ 0 NA pol~Rlio.rllh~sm$il1l ./'J,ro,bidop;Ms llMlr4lt~ icle l~8! e~Jll!!nl ,of nall!! raJ 9:~neHc \ralriat1lo~, with lrnafly g~,ru~~ is(!! bled ill dirr~rem \~kI :liUai 1'!1l,. w~ll a~ h't~ h Il~vells 01 polYln(l~pllf,sm am(l~g d as !£Ie-ne fil'l'l1iLy ll'Iemlbe;r~.• lrlc.l9difl1g l~ciSe if1ljiDI.\I'~in dfc5El.1lSer,esi~taIflCie<.
III nJdeolide

o:r b~e: hi,1''es are \Vl'i:l~kill ClIii'I1Ito b~ d~~~lyr~~ollt~d• bu;t.lhirv~sc~n Qh~nl e~hib[Jt mo~ . ge;netk diversity 1.llIan might be ill'ltkl,pilted hom theori:e-s >01'1 Ilhe l)eone(iL'5, oJ w(]Ip.eratio:namol'lgl (lo~ ty ro'l<lilie(l 'umdlividLJ!:I'I;sM(!nil~a (iUild Seeley (p. 3(2) s,hollil Ollie reason ~Oll'~his is Ithat mlo.re glel'l ~~i.. cally di~~rse iii i\115 (Ihm;!! (J,rigTfI.1lltin!lI!'mm ~ Ienl'<! ~e m~t~lIlg ~'I,Ii[h l'nump,le males)' ,~erf,(Dr!'lll bElUer iiii the ~i1lte or OOIl~b ~!Jildif!,§1,!'~.ragiflgl r a~e~,anUih~ftey pr~dl!l(lf(m U'halll th!!):Se oog if!~I~1l!g '~r!(lffila ~ift!Ylle r13RlIl~.!!' andllll1ll leo.to a d\!'errti~!!her p,res:em:llinll1e (.o~ony 011nel ~o e&'ii1rl inHullliU:e O'lfelr il~ me mbers .• a ~i(li1:ey:boo queen p.rro ucesa (i!!i!'Ipl~ b ~~~d ()!i' SilIbsta nces kIl!,Ofo\i~as qu:e'e!ill !11andWb ela r IPhe;Fi)!~Hifll!l'. 'Ve-rgo2; er at. 'IP. ::I M $@i: tll~ 1P~1I'Si1l~~tiv~by Galizila) i[ound that @xpoou f~ to 'ClU€ ~iI pherom~n~ tll'ids ~n a redudkif1 in ,avt\r~ive lellrlllil'l\11 bldt Illo!1to <! ~dlJ(ti(l1ll nn aiPll~liti"'~ tearlll'ng in }"OlIIngJ hll<neybE~.. lhi? qJ!!!!E!I'i $i!!b~~a~,ce miX! i.!la'~.e~the iJoparnul1I!i!.rgi( s.y~t~rJiI of b,et!$, \~Ili':h redlU<;es~ihe ca !Jilcu1y ad y'mlflg wmker~ to fQrrrtn[W~r~We Imemories,
Th~ ~sid'E'I1I~~

What's the Buzz?


:!IOU rceclcou

location" l.ocation. location
OeSlli~e Sli b~~.antia![~flion,il has. w,Bmalill!ea Ire~alive l:!llmys'te ri OlliS. how the prot@ifil kfiOWrH! s H@dg,~Ifl(lg '(IH'h.hcthl'1:lU!!i s~gna lin g patliiwa.ys tha l re glilille \i'a,riou~ hio~~ogi{a~ p'JiOil:E,)SlE!s,in.dudrimig ste FIn (en rnJl'lcHoll(. devet(l pmeflt, !l!lld ~a ~()er. IRlQhatgi el' '!'l~. (p_ 37.2; .~Il!'!e!.hll P.er~lJiE!dif!ie by [hrnS:hl!l1sen all1l,d on) sholll' ~ha'll~lammal i'~!l'(!e'Us tlse'their Ill'ifl1'3ry cfit~llI!!Ji!as ,a~ ,a~uetl !liar that sllmp,~e5 lilfe su rroolfidi 1119 nvironment fill r the pres8<rl(lR of e Hh.Willtln Hhr bOllifidl 00 its reCeIPl(U' W'a~d!edll (P let), lh,e !:ec.e.j;)tc,r l~rt. th~ !l:11l~~. \\lhere (in the ib~m:t cf St~lfll !ldo:!llQIl ~ it aeU to r'e:5'trairill!H h :slgnElIIMu,;il by preventil1Y a:u.I!HI1lI.lII:alJorHif l!Jies~glnal11119pr(Dlelrl STrl~~hel1£id (Smo). A«lIfT~ulaHon: OHITHI in ~he< cilia oi!'~limlu ~Ited i()e<l~5 (Orresillonded tQ! ~('t,'\i'illlon oJ Hh sigll'l;;iil~119. FIJrdu::r .UrI de rs ~(mdl ng the ITHl'[et,Ul.:t r mech an ~sm:5 ti1l1t iniluelKo!!! u!I.lula.r ImaliM li'OIlJ or flt(l a II1IU Smo wi U irn[pwve 1I0clefs.tall'ldif!lg Qr l~'~ sigl!\i!IIi!1g 1JI~!~n~C!y nd may ~~<i'!d Ii! rtlV a t(J
ruh~raIPBu'tk lii3r9'~t5.,

Long evity' (In the Brai n
S!li''ef.1lll, iShldi!~ snow Iha~.losr!Jf-hJll1ditm mll! Iil'~ion~ inIh! r(:J~uUIll·I:i!le siglrliJLf ng {a~c;1'uj!! 'e!'xt!!llth ~h~ me 'Sl~an ~f worms iilllrd Jlt~e~ h@"l,!e~r,.:~ l;,IiI,I{I~efl~ m ~ta~k)<fl:S an;:: .;ii5s~r;:i!1'I~edwitn met~bolic: dlf5!eil S(' a ~d [alai. d!all~te-:5. in mke. ~Iilcontra!>I:, Gl,t(llri'eore5ltrid~ol1 Qr gi~nf~h:: fit!ral~.e9i(l:5, ~11rnke that enlwmce iifls,lIIf!]!1s.ei1~itivir'l' hl\~e~ 'the risk !l:llr age·w~l.altoo disea:S~iilftd 'il'l'iteP'l!di lire SIP~f!. tagllld~i d ai. {p" 3,fm ~es,Cll1iE:the:s~ CQi1i~lirctillg reS-lilts by p(!ill~ill91 00 the braT (fg ,11<'5 I,h~ s-it@ wh:efie' n~,d Wlc!€dinsi!!U!li-lik~ Sigl!lill.ilng (11m eJ(lJe<ndmouse ~i~'~ spallll.

Applied ae:nOlil'tili~5, CQlilversant He!; Exp!JIes,von Ga:!lletiic:s; Extlr·emol;)'llllilo

Getl!a~Q Eliomedieal Pl"odY'as
NektarThera,pe.ics INew' Century P,",lal'mac:eutic,~! Olpenl Biiosyswrns

Mirl'imcd ExposLlln~
Tilill f'ElfJE!'nt.d~Sro'i!iBl)llliIdll(ellitain vim~~ expr'~%'5 micT,oRMA!; ~miR:NAl;) f<J,lS'~" ~lhll qUiIl!>tiun ~ wheli~er l~~e P!iI~ho{!ens might use !flJ~RNAto evade thli!l r ho~ts" S~erll·'Gil!i!o:;iS:ar et ai.
,~ ~h!E!'Perspective by CI!JJlllell~ fiml U'IltI,t fm hmal1 (yi:oIll1£!9<1llovil'lLls tllis al"P'Btirrsln imi'e-ed bE! 1.lIIec;a"it'. ~ 0 i1~ or lh~ li~n,i~' miiRNA!l \I,i,acs r~dld~d ~iirgel, lihe 3' jJln!!r.ar:iSI,~l~(Jre~fOiii!l; or b~,(i ~rmiiiilu nt'i!-~i~~ted p to '\I~.n!il$., which b~,t1:me: adiYate-din fe$llJl;ln~ t@ vilr,ail fnf!i!c,ticiln. &;pfes:~itln Q[ ene ol~~hes!il prQteimi ~ was ifllde00 dam pened by the viral miRN.A, Ij\lh kh reduced re~,og 111f Qill by {H1!lMrallHr~l,Ilra ~ kmf,'rce~ll~. I '~ ~'llrelllllllim.~o!be seen if IIIiIlNAw~lllliLJrn Q!JJlto be il WUd~f>,p,reo1lldf'n~~tul'dexl1lo~ bYVirlUS'@1ilQ tvad e :~ fl!(lst iml!1fUJl1li~y,.

'Opero." EliiomtCh!If!;Ql'Qg1ios Serin,a lMr:a:peui'ti~s are als:o proud Ailjo€iales or
the ~Lid:Soil:i!-Afp.ha htstitl.ife

'p. 376; see





M'M.:sdenc:emag .org

Sc.II~Nelc VOl31]

20 JULY2007

MieN'S yo:ur linllt m'ICSJlreeradv\a,nc.Eliment
y,(lrkSi1ops.and :st~tegk; adviQe'~hfOug,h S(reoceCareers.org
:;li1d our C.eoter W5 i5~llhe foliefmnt: of adiaJildng early-ear1l;eir rreseafGheirs -!!lIT~~ing fob s.ealth. grants aoo, f~Io~",~hi;ps, skiU-builtii~g

rQr Care~~if5im Science &: TechJ1()~ogy.

NI~ nile' NilHQnal Pij,stdocJIQ:,ralAswdai~o:n. is. prov11ding a oa~!l!'lla!,"la1(E!.Mdseekin~ posilNEldlal1lgE< ('011' postdocsp<lrtnlenng iA!i~tliAAASuflC6Ir:eer 'Ilairs" semloors, and otMr events. In [Rct"AJ!\J\;S ~~ il1stlilJlIillent.a~ irn,l1etptng the NM get. swrtecl ~Ind de'!.ielOp inl'Ol a grolMng orgp l1l~aUQniIniLI <'Jyjt1l~ ~ rnk 1.0postdOt~LK:CJillSS. Iffyou'i'e a IX!'rSI:doc 0 r~d Sltu:lent, ~'O to lhe: AAA5-IN PA Unk romdQLJ~~ to spell !ca~er ~~(ess,





TARGHS (oLin, Ola'iilen is



[1.1 ~t'~ if'f[tey'vG b~ef'ib'l~,et [!'iil~oosli!llg~,}i"mud~

d~.¢~~g~t ;'iiEilWfm,~uy!:lut Hw~kmgr;;lloUgh


member of PatUilf1I1l1rll: oll1d cihair Qf th~ I\!Jl f,nl)1
lP.. r~i mell1t!1ry aill1ll~te .. (h~lnge Grow,po. wh~dI h~5' lilUl1dll>O alii ~l'Iq!lJluliY nl!l'to tbesetli ng 01' gfteniWuse ga,sredlil{tijon Ilafg,elS, ·ma~~ ~ (oollem @P'<IfUa~1iei1lt.l;lk

U'[IheCi Ki:llIgdmn have iI'o,~~d ona, a, co::! elml:~ssg.(n:IJS C'ul of at. ~e;<'ls! 60-%. v/ill,ich is i.[1Jlemled to Iri:ldUefl averrragc;:' gl!Ob~~. w.%r'.m,ing by 2:"C (JlliJ.c Juue at'! :!>l;Iillllmit aiSol."lspob o;f~'l'Q. ~'m,is~oll$ cut of5iO% ,gIQm'l~ly_ h~n on~y in the OOlltexl or,ex~j:~m:illlgsuch <I gm.d fIIml '!Lvith IlO gtrCGmllHmse .@}11O SlAbilli:lfltiollla~N. iii! :m,~I1:(I_) W~~a~are-the chanc es of meet ii~l,g.the 1'" (D bjeclive"! Nat I.ik~~y.~c(,)!1rdif!gto M,ll'~~~ Moei~s~l.wsen fth€l Swi~~ Fedo

;lboul t~cldil:l,g clim<ll>e ch31r1ge <JI11i~icipales~1a'!;,lillig achieved !:!ulu:,thing b~~]~e :1l1I,1.idJdll.e il~i~~mni"y,\;Vh.n·$ the ~i!lIg.;t'! Qth~'h(;j EUl'Opem''1 Un~~ln B fEU) 'll~d.;xi:.11J1lution~I:~leve], t~e


~nllll!1i&litute ofrechr:J.o~!)g)'. w:~o p·resel!!l.t~dthe M:i"'lfIlif~lc
~n a report oflbe 200.5 lExetc:r I:;lim,.ue ehauge eoufe renee <.lind. h(~'s been q llo[led si rice. bnth b)! UK w government eCOltmn.ic ~dlvi Sior Sl r Ni.chQ.IH.:!l Stern~lul tlt~ l.rll:ergo",eti.'1:~Uel'il.~~;Pil~H:~ Climllle Ch"l'ig~.Hi:s amliy· CHil

sis or ~, ,climare sensitivity studies (}{the ell1,;i'C~ ~ ofgi,obai C01f!IID(L"1)h(;;r:k 'Con~~nl!1l:l[~!W:'i (llfl 111i.milp~f~~Uf~l:!ll\,\;'S $ that selfUllg for a 60% cut in .umo5p.in:el:.k: CO, 'lwhich oorrestwod; ~(I 550 ~rtS perm illion byllOI ume)~~e<l\les, a ·p:roin,:bi]hyboe~.wee~~ ill~d 99"l1iiof Illissllllgl~,e 2"C tar63
g~~."Both the UK and EU proJlo&Sllsilldkatc tluUllbcir


em issions n~dw:lion ~;jlllgels]uighl be wl~ghellied.rer~l;J.pS. like an ::J!lhl~t~~m~I1~luing h~~~julnJI. are ~\:llI~n~ing til.;: \I,'I,;! '~qp <It 100!i'eif hei~hl!S firsr, DUl sc.mll evide~liI.oe suppor-tslhal"
IUNury. Na! ('I1I'nly mus:t \',~'(l r~~lc~ ~J1Jthr~pogiil!D'J.icgree!1ho1iJ!$e" S''i!l,f;::misskl~S•.\~''e !t!,r:ed a ti:me~ilble U'hl~reduces tiH~ risk !."ifpo$~~h.'e Il'e~dbil.ck~~ind I:~!Hs,~rik ures t:liai'l! eould l!eaJdI to rUD3'W<II)I' c<lit<!;stIrQphh;cl inl;1te' ciJollllbl'C. 11m, d(lml)I;:'I1!i;Y. iti8 din'ic~gDl to CQI!iliVilllcevot>en; that d,le,), ~~,!.')u~d a l<lke ilcl;i(l~,l~e$p~!Ci.l~ly exiP~llis.iVCenes, to avoid @f1il:5yet largely l]f1S0ell:!llld u~qual~I:~'I]:ab~e d<U;lg(1!r. HOOf do ~'Oll base a pol i..:.;ylft1<J:lis Iike!li)! to 1111.wesi_gnifk:<Int economic n mp~~'t::l,UI'I :n,1:oth:.i .J'l~:~ forec .. srs t~}l(.'i:1 and smne u.nught regard as glu~5"iom;'k:"?We OI.lmly need to recall du: fal:se ,eCOJJ1l.0ImlYOflUl't !s~lldilng ~;)L)(p.~V€:fS· doll~1"S; filill huHdiimg up lilt: N(:i\v Orl{;!llmsk.':,r.;l~ t~.lrealize hO\v;)('liO'ffiI~ tak~mi mdil}! oml~davert :ilIQlUIIg,~nm~~ problem ..De]ay•.oomhined wi~]n.~:llD.e tkat skepltics nmy accuse the riek AI '._r~lffiS·(lf~hi;~ t\i~1:r;dof"crying \\c'Olf:~(:t1lQ.ld :lmlflkt; mlgh(;lrp~)lid~ ~ h'lror;f!n .tldtlP~·1'He~. : I:~: sett:ing <I LlIK t3rge'L the !liffio'Cl.'Ilmem. [Im:LIsralso ask \Vilm~!~lhe ited K:~ngdmn'ssi:121reOfltJlt:' U:n blillrdc!m is, U.!l nijliol181~ U'Ilrgct 1U'I,I~t.nec~$s:~dlyr(!laJt~ to li!L'!duelio~:$inother eou~nri(!.i;, i~.d~~dli:~lg dl.e cl~;e~:opinJSwor~d where il1l!dl:utn,d glwm"'it~.l~o aUevijJle ]povel'tyi~ urlc[em.;il.lge~uissa!Cll]ls,. as
C;JI1IIl-olm ..ike <I

shcmldl, IIl)'OIlC sign an ilI,gl:ecmclll thflj'~cenU~li1i,tJ; < IthcJr rn~1n disad\,lflnmge':l

n'lo(re incl~'lI1Led to 1J~ld:y,ei r dlweloped ooumriesliJUy

Ibr~!>I~lIdQ\wdin 1.99'2bj' the U:~i[~d Nalkil~. F~m~~,ru:rkOJIt\~nliiPI1J o~ O:ilU~ltt; Cha~g~. rd~Uh1iliill mllliml:<l'l C.1lklll<ltiolill ;uullhUH'!.' i.t~nto the gldba~ pot oftilmge'I:S; :IruUn:I'.\'I\'f: ~V~ ~[) d~tl;l~i[l~wh~~ th~ gi(lb~~!t~"l;ld ~ldl·igurGi!llit l~(l\1!i' ~fl dbtribu~e it~~lc~I:~~lii(l:tm is. ta~{,'i~ UlIiLu~t t'oln!bi~e !'>Cien{."e w~lh j uslk:e. A :i.'U!,;t¥:5:sI~u[g~ub~!'1d iil'lrlte c1~OliIlgefr.Jme~¥l.)rk ,."ill :~u\'~ ttl .~' as, mueh ilI.u>elllkmto the l.n:l:er3S tuthe fo:rm£r~ GQ:ua:~n·ics. such as Chil1l~ ;<Ji1m.dllldlifl~.\I'm be


embrace [heir Oli,v!1ref!'.pons~bUili.e~


The UK gO\'I;:TlllTlem is the first to ul.ke enrhis c]'1(JIIe:ng:e, \,\thh pw(bI ic..tiO:iiU unite drdUl eliim.lre Cl1i!1n~eBi ll il'l!\I~;lroiilofthis )fe3f. .Its Ile<lder-:shi]~earries l~rleres,p(]'IiI'Sibi~liltY~D t!.I,UUss:iOIlS ye~ t'l:~I;S. ~ght The fimd bi~~f'!~~~t1;to m~e ex:p:~!CiT l~e 'm:nnuhl u';':;b'd~t) OlrrV\F(l ,m an,,'!u-~~ Ihflt glJ\~~i1!!D'J.~~m sets, "fIun; fornmul<l. !ii1l{)1J]ld~ellus net only the size oflbe caJire btu: <lilso .ilI.ovl,i~ve'eslculate O!1llJ!il~are' ('Irk T~~ d:rol1 bHI prepeses a fi:g1J:~ 1~:Ut !I!Il!1;t'ub~~'l\:pb~in~d terms ·~:f~i~~ler eriterien, .~ dkL C ~f'I fil: ~~li!1~'1i.lDu~durely OOOS[ confid.eri:J:eI:~OlIIl the restLh is desiglled~.o sgh'e the prol.l~eli1.1rn,:stel' th;HIl s w~'rcc~<llillg ~t.I suspecr] haw s.-et: l~-nyS<i;':~f,~targct onivrun!§ ul!illti~jl"~~l toO ;!;;~~ w\lr'mi inI'lSP]~~. 97 -(d~J]n (11a.lmEhn

M'M.sdenc:emag .org

Sc.II~Nelc VOl31]

20 JULY2007




EI,@{trh: Aft!e rshoeks
,E,~rlihqua~ ru ~tliJ ~I!:S:;lIrE! expec l~,d,t\~:;g,~n~r(ilte ' eLeiCJt:mlililagnel11cd:l:!iivily wi'lhiin lthe'!i.EirmUlml~ iirrlg m~:Il:s, but, d~irect ,evid,ellil:eff,~u tlilis efied, h~s been la(k~im,~.L~bor~torv 'e'xp~,dmeI1U OIl'!
Ireall'lo!ck~ ~!ll' 'generalte cuueillt!'i dll:le ~,oFluid mO'l!lem~,li1t aln~! pf~lOeL-e(~ri( ~HecB. but they oil re 1.1i!e;;llk Ildli Iii ~h,e gl€'~ t(ll,gica~ setHnlglh a 'is, ij:J~rd to d~serm~flgl~ ~h~l'fi'!hl)m !i!'~hlr~p09lE!n k a siglnt.alls Sf rlUlITie! ambient e,lledtrOlni[, nl]ii~le. Pal'ik el' al. rE!'p~,n pa$siib't~ d@~!!diofl of a eha riH:'lerisotk !li'!edriieal 'I!]glflaL 'LI~ilfl9 ,Hi1 e~.e(~ ~rod,e arra" ~laJud !D'n Ule 5i!!fI Andlrea:; ~il!iult ,n ParMielcl, (a,~ihi'r[b~a. ,Eiloed[ical disturIbal'll~~.li las~ll'lg ,;I hl)lJlr~ we~e pil1:~'~lll,lp with~!'I ~ 50 I'!ll~f thief,![IItlU fJl'!llme,dhll~ely ~f~~ ,aI'!i!i!'9191nli,t~de6,oCle,alrthtmiU(iIb~ ~~alt (lI1:C~~r~~ ]rrlSeipte IIIiI be f 2004; :;~Ig f1ids,off QPpOSnhl' polla rilly wel',es.u Dlltq llIoe'f1Uy ,~'etededan-ef ~1I\IO liIiIag nitude S. (I ilrt)~rs.ho',ks. Alt:hQtlg hi :s~:,~Ual~el,~"ttr(lf!fl!lgl,llIed~ (hang!!;::; do '~W,lr iOl!il a dlalil~t bo:'JS:is if! tM~ awn. ~,~etea In all~guet~gt tl'u:: lOl!:allb:~ti'I;;;!!l, ~qllil'l~ flg,lIlld Ulnm"lI<lll p01i1lrHy ef th!l1ilr5<ig nab slI,Pllmt <l1isocial~ on ll'lliilhi tlll,~ 'lilartlhquaJiie m,p'l.lIJre pmc;:eSIl, 'They prOIPSfie n9~!liI mOloierilll1lmh as the lIIl.o,n likely ~<lILIISe t '~hJ,e o 'eledrka:l. 5igni!lb,i!lltl~cllllgn tlheyalre IlIliIillbte to expla~nt the rapid onset. No 1P~1l(!.Irs(lr .s<i,gnats Wl.U@ Qbs@rv@<~I" so '!hlls tEldiHliQI!.!l! m~y net ~ltirn(j'l'e'lv Iill~lpwith @Iil!irthquil!.ke' pr@'~r(liom.·~ JEI J, GetJpbys. R~ 112, llj(),]0291200S,IBOM196 ~2:o[m,

IE: c 0 UlG VI'E \1'·0 UJlTI! 10 N

Sma Her Harvests Than Expected
iLeall-[1i tli rgg a nts (Df th eo'gen us Atla~re ubiqulteus f'e'S'ideii1t~ Qr ~eouopic:.[!1 fO:rl!$t~.lt!~v ('QJiIs:lnJ d Ilarge :sui1ter ramH~'iln (:o~oa1ie~i3inil jOILl rnney on trab ~rCliS~the f!D,~t ri.CIQrand: il1lt~'!Ih~ f!Dfl.lsl (iHmpy. ~~her~tmey hcl~t lea~ haglmielf1t~ that ~ ~ ('.g rried biiJ ck lI(Dlihe nest. n,~ fr.ag ifnen ts IIlOUrish a IlImlll1cllisti.c rJJngl.k5. ~h!l!l in tum j]mvide.s p~~:teif!i:md (.arb!!)li1ydlra.~e for tI~ aiM wt(!ny. te<l&'[1i Uer5 rulV<E! ~een \~delv <I,~um,ed to, be l~,e d{llnlill'l~mt lherbi'l.l'o,e~in lh~ rQ~~ts thll1!yiJll~a:IJit, I)!il~ 'Sup,p()rtiv~ Qi!!all~nalive data r,or this: .aJ~'S!!!lIrIi,p~io!ll a re s:palf$~. lHiE!rze' at ri r~t II,l~OO (! r .i"lpidi1lll~ noolli:le'strll..!r.:I,llieIm,e!tilCid. 111vQlving U'Q'E! :5i'1llT1pling (lJ !iehi~' depi)~it,oo bV!;l fi!~5(llJ t5~de ~heir f!J~5~S.a~ ~ prQ(I!:;yfIDF r~l'~ S!!.J'rinQI the daily hilrve:<;t or leaV€),. Tih~r1th~y adlee ted dalalmrll l1eauly :5.0 nests. O'!oI\ef ]LS, m,~J:Illh:!l in a F1~na-

l1!!af-tuUe~in til Br.lIzil1l1l11FoFes,l irll::IF'Etasedi ~t the marg)in~ '\I'e~sl!ls. tit!'!, lF1teri(:us) o'~I~rem ~h(lt had be,~n hagm.e'l1It!!!d by hUlllian dl s ti.nbanu!, '~hu~

ali 'wa:teris lost WhI!!I11 water b@C'om~ aV'ai'lalb~~

ampUfyiflge!'lviroliimental[,cha!l)ge E1!dge.-AMS



Ii E MI S1Il:V

a9QJi~, dehydr.a1red ~aMe 1ll'!dielN_l(l ret~,~r~lkm and reSL!lmll!! th~l r deveklp.mel1~l. p~'Glge~imli ~ a!lLIIt I1lo iinjd~es. K~blj'i!ooa t'l' ,Q/. ha'lj'e!:deil!tifie~ a ue1iI00Ilosellran"ipor~:er (cOIllloo TRE:Tl) 1n ,F!: WJ!l~l'er· prallki. lIhey'5ho'l\i~haJt ni~spceJ!:ilk [Of trehalG:se v~rSiJ.j~malt.o:se., s~m'Q5e. and I'HJas~; t~ey abo 5hO'W~ha~ ]t hilndklniS as a lo~aril flily, hi:ghctIi'plI(i'ryiacitfl!.ated ~Htll1sporlErl~a! can be' er)ll'llre~t'd beni{l.Iil~l! ~lfllII'hilmm;;llialnc:eUs. - G.lIC il:i I' '!!! , ~ Pr(}C, 111m'.AClW. S.d. U.iAlIl04, lisa!) ~2:007t <:l ~ C ~

Survivinq a Dry Spell

me '(01:5 we k~OIN ~Ois; ba!>t!d, on earben, and one r,otll!!iOClu$f.act·or is lIu compa'tillililV of sug:airs: and Wdter. Glu{(lSeis. readilly 5(1~ b~ fait 'mud, lhigher CO:lloenlr(jtiDn~ tl1!l!nthe bul'ldlng Ib~o(1i:5 ([i~he'r IJr bloklgi{;'lI. ,polYll11ers). ~a5ily bmUed byeilll'lnile~
via i~s d)emk~llh;II'!r:U(linatili~.~!1d
(and IJ'E!r'haps: e\l'efl bel1l'e'~cial) in its in~:e!:a[Hol1'.i Mth

Natural and ArtiUcial Flavors


(Idler bi()(Ih~.mkals. If!
Gm~id'erll'lg the lT181ji(lr d~LUlalifig sug~arilll if]sec:tr-tr,~l1t1lo~. which is a h~d-lO' hBtldi dillT1!lr,of glu:-


CQIW!!~lItersdeli1(~stshave l!;):rlt;lW(lr,ried th~~ ~heir i :! n~ldsuUers hom Sipllill p~r.~ollal:i~y disorrdler:::is H

do ma~ht:!!'lalic.s, ow elflgi!1!eeling? Tfu~"
1lI1:'i S LICht


Ihey work {In preble

as wrltil ny


rre ~

10 co1llrry OUI C:<ltCil.l~ation"" Of! 0lI machine, «II

bUll ~hey

mal1~afl fo rest all!d calml<lted th<!t lm,e.alnts
~'l,it'.~e actuality ~es:po~~i-

(iose-=~he e:<uoilo~df"
~alry tOb.lr:fl!1;C:~Oif

tile 'fOoronly a DIJ!.d
0, 7~ ~~.to~.a~.Ie oons~llI1!p~il)n by aU fol~vor,~, ' ..~ (jn!l~d~ alnd Y~fWbrates) in lftllii:(wet Evn thOlD ~h these resuH~f~d i(illl~ 'tIlall t ~e d efQlhraiion by ~~af-(l!'!.tfrs is m~r'~ ~l1Illde5ll.h(ln p,re<viQ!tI~I,y tholl'ghl, 'Uribas el oJ. fOIJ1'l9 lha~. herbiivory by

PCJpPM#UfIJ vander· l[iehyd1r:a1~oo (hlfJl:) aad planki Il.aFYa~~.o.essid r,~loydratll!i!lll;1!r'oia. cation ((Inl~s!ID'Inlind. Whu 'h~ poQ[~ rod; lII'here th!i!~El' ~1IrV'U!' dry up, the 1<llVal r:tlt Ibodn, ~Vf ~yf!it~€siz€s lrif.!h.a~os.e' Jie:lees.€sil filIO~lh~ a,odl h:el1llolymIP~ 1n mde~ to Ij:uo~ocl tisslie mnsli hleoflls

als1J 'gN1JlIl~~ ,\lith th@ JiIIIoslo1!lbs:tf<Jd allh~nllltim ~ :poop<ellies or ()(!!ITlll:Dll,llallC)'I1.l! ~ proced tlre·sameli ::;; I.he' Ilogk (J~ i!~gorillmm5". So,rih@,debate has -' rag,ed: Is the rteld a sdence I!)ff t he ~a'Wrah!{(l{rld .~ or O!1llyascije!iM~e.of ~he a rlilfi(iall? Deorlil!19 ~ al~g!lies thOl[ [Olll1lpulte'r ~ieme ls decid~!rl'ly a nat- ~ i!!f.1il.$ci~I'I'!.I rlf:Ofm~tioti ~t~rage and pr-ot:!SS~ ~ illl 9 !lave l:I~e~ rOllndl to be fll,ll1ldal!Jl~n'ill er~~ g


H~~d:!i. hQm til!!! bJ el[Q9Iica ld a~G ~ :s;tm,e!dilfl DINA to the ''I!lHllnl!il1lJ1l lrdmnlo1l!i:orl that ~ is tr al'llsr:rnitted arfld !1Ilod ir'i€'d a(!; parli(i~esiJi!lieir,a(~. 1111 m,m:f[lrM~. 'Prim~lplles~lho1ll. trilllilS(eollld mm0
ilIiI ~nts O'hlll.:l rlV


200 JULY2007


pu ling machines ferm aset

{If q ul:!sti,tm:s...II)1)U l

the de,~:pstr!.JW.lro of c(!nlput~lklnl. these questiQursl ill U!m, elf@ d'f~"lli'IIg ~lrlll1o¥i.niw \\iays to


mmpUI~nl!1g, SOl'ne(imes ",itholl,l[ u,5,il'lg! !Hlphlisl"icalJ@d (<lmpL!tIH gald9@~ry ai, alit Thi:!'

InrlcrOO~i!!I1~sml$).~fd1lis PQl~{f'~ €_);t~i'!clsto o.th~r ~Bmpera'te JOfl'sts"lhe mllHguril~iM ,o:r below· grol!!'! dJ roocl webs an ~ iJI~~!t@fml'~If 'firM III nu>: ( nli'ght Ihllve to b eo fl;!!DllIIs!idered. - ANtS, f,c,oI, t.~rt liD, 72:~ {200!L

.ilutl'l~r '~~flch,ld~s l~,~ltthe r'ield en~~~p\gQ:ses a
$iciem:@<{:If li1rOr.ITIl1t10n: ~roc@!isirlg illl IbM~ natuI'iiIllllnd1lir~m(i]al ~ys'tf!fIn~. - Di\f (Q!mm,lfl. /1(1'II5iO', 13 t200'';1).

IJifferent Routes to a (~lJst~r

FJrM Tum:!!' Amhors www,'suomllt'2:5t!lenc:-e,Q!!
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research with impact, turn to Science

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study water wav'es'?

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s:pend the rest of tho ,day sllr:llmg: the waves.
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erosion and natural

d~sas.ters such as, thet::H;mam~ ~n South lEa,st Asia. A'nd this research can beapplled to all.

S,e'ing a. member or MAS means I get 'to learn about areas of ]n~erest I m~ghtnot oth,e'rwise


encounter, It gives me vai~uable oppontl!J.nitmes to eX(hii:nlg:e~dea.s with colleagues ln otherflelds .
. nd [his helps me find new approachesto A " -

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!Dr, K:atheri'~e Socha is aU'llassistant professor ,of mathemstlcs at St. ,Mar~ls CoUegE, Maryland. She's elso a rnemoelu o,fMAS ..
See video clip s of thls storv and others



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~F in'il,llimzil 'llirulie5 bllrgililQinrlotramelll H~lb. To, lD'ilU~ dnlrlgesinr ,lII1IelliiclIi knowledge andi PRlCli!()~, browse the hiSlOlrkal. (,[IUe(li(lIDl" I;'I,In!l)se~(ltdin.gSi ii1ldlild~ Irari~ie~~liIch as 15,tih cuwrry Chlne-5!E! <l11l:atQmllcal dralrin'gs <!Ind! aJ1l9,20E So:vie'~ pmpag,mall pmil:er 001'1 d'i! nl!lJe~,,e:1tvph!J~. thi1!' Ii YOll!r il!'!lell!,li@f!5 are PIH~ (lW!a:t 115, m!,oflJWi'!uli!~~daRyOy ca~ dow.nklad~he ~f!il.~~S il're·e. »0.::image ~wellcam~ .i'H.•U k

Mo5qflli~~t;!5'Ci~1'! 1,M~llk fi1wa~~ a5 well as il!l1J' O wilt~rb:u:g. o[stid, tea w<!llll~lke Spadermliui.

Cfuiil1E5:e bioEngJl neeir5. are r~gming 0 ut

what m!3!bs !ihem Siuch ve,'Sati~~ pests,
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[11'1¥ scall'l'!~, eaeh 'wil~ I,Ip tQ a d ~~l!1fI ~.ong-iMJ!diilia l ridgll!!s {Ollllil~t;t~ g:f (-illl~ I.ra llI'Sv!!rSlE!ribs. 1 hI!! sci~J1li5·l'S,sp~li;li[.;lJ~oo lha~ .;li~ uaPr'ped between ~Ihe ralbs may hum "1ill<lno(IJI'5,lhiol1s"~l1al cOl1itfibl!~~ 10 lbuIOY~u'l.CY. bUl their eXpEri melfll'S alsQ il!ld~cat€til th@

~mpO.na~H)eof the a Fl9te or the IJeg if! nell
Itlri!!i!k~lng through lh,~

~!lI'loot of lW~idh h~e- Iil'll\l'ef bf~1'l 'J,I'i~ved' by htlfFn,an eyes lJef,ore/ il((ord~ng ~o a s.tal~;eme;nt 001'1 l~~Web ~he. 'F'alrddpi1!lllts'\Il~lc(l'leg,or;~@ IN!(h ima.ge as 'Spi r,d, elli plka~, ~l.aj/dorr!'1know. a:r Inefgm. [he' spilrail9!~daxies ar,e '~l11e.ll ~!,Ibdh.l'idell il1l~o dm:twftse. illI1tid(!(kwiiS~. i! llIe ed'geo,oll. "the hU11Ili'!11'!1 brah'1l is a!itwa1lly bell!er
lh"m a WlnljJlj ter <!Ill pi1lUen~ r'E1'wg.nfll~oa! l~sk:;


i revealed lhatlhe ~

insect's l~:s <lIreequ ip;pecl \Vith

S(a'~~'sen sur{.a(e.As Ih~ .1!Juthos r mn05qllll~~O I~gi. ~otll,. m.o:sqllitnes .aN! ~qi,NiUY :U h()m!! oii1 dry ~~n~. IIItlJi'~50iJl dll1!l their feFj~.t:Ire eq'uiplPed wuu, t1 rIy hOOKS "nc Wiler!!d in (ldhe:~!""e hairs simllar tQ thos~ (If!1 al


lik~!hf~." sa.Y!lO>:ford asuoopho/skin Kevi!!l S(II1~lil'inskl, A:strop hy~i'i:!:t Bob Ni (tlo ~or P~rbmou'Ul Qdd~till;at geUillig U~~'g.alaxies cL1:l!S~iri~d is "as fundamermtall:!:S ~I'lQW~lFIg if ,ill ~!;.I1lTi! ~IFli:s rna ~e or !eFlilCiI.e.~

B'e!lil!! rs in t~e' s(l!.lllh,~rlf'l reg~ on of U!ll1lg'lledGJ(,. S,lephi1ln'~ M~itllne, wil;h M1e Frem::h rese,1IIr'(:h i!getlcy O~ RS.!fIi!. Li!'~t,~. ~,aY's~he wililery was dEla rly .. big busllllf'5!1. A j[2:.II:nY'·50"'liIU!~er blli1ldlillg o()!1ta~i'led 1 SO huge 't~rfi!'CO!.l~ '~~nTl!I'.lI~ vesiSeilscalllil,d ~ill ~\ia," l:alll~nl IIIiI<IIny sflIlIalll.e'r iilmphoriilEl~Ora'Q]rn'g WiJll~.• iilll1d so~olle5llpportflt rlJ(t.u~s ror w;ineilll ~~SiS~Si.~ It ~~l!S (Iii! ite as(I,p',hisl Leanedeou!l'prise, lM~h HI rllnrrng walE'rrm dearnil1iQ' I.h:e [j,ugii.]I, ~ S.[f¥~MiilulIle. Dallilll"g the IfSitatbllishiment wa:5ea~y thanKS tc;l a (oin 'hm a bol,J~.20 CE. rQ!.Jod, 111'!!he' a ~e.a,

A~chaeQ~(lgi~h ~Iidlla~l we~k t~a~ Un~ [had dlSCIWEr~d 'ihE '(plde'~l ~iflownl win.erry iiii Frillil~Ie-, a t a 2000'ye,ar-c'ld R!OrflilrmviU.a n~al"

M"",II'i~ijI1igs i(lO ~hJe WjII'lE' 'v~s.flbi'Ill{Ji,cate '~~li;!ll ,aJ l'Iler(~alrft hQrrn Piu~eoli (1110111' PiJ<lzuol.i), 1II,!!'3r Na.ples, 'liniilll1im ~hre 'will'l!!ir)l'. Maulm!)ilIVs ~\ICI:rkers hallie fOIJ I'!d 'nhe ill aline's (lr ~ dO'~en CE!r'gil1lists (!I frI1oni1lh~ wi [J1efl/'s estllll1il1ted ,eo le-:mploY'E!es. Arr(hee,lagl'5t 1e.an- Pierre ,Br un, d~ rled!Dr !!Jfllhe ]eall'll 8elra rrdl

(enter ill'! N~p~es. Hi!~y,~ays

th~s~nfdte·ct:i·II~eo EHi!'~rIflllOU!S

!Jlrmli'~hol (0 IIIiImemda I.wi'nle m~1ure ,[IinI~ l'xPQ'rt. dmilngthe (irs'! !illiflds.eOOIil~ 1tE!,IiUlury C.E. "l'hi:s ,i:l!~~(j ~'WiiS, [h'e 'Fair We~~' :ror the R!JmalrliS," he 'SClYs,ml1~i'I!I,g lillcll till ey ~'®re Ill!m~fB ~o GallJlllby' (h~iillPer p [odl.U clion (@SU. i;!Iffi!dllalil~.

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~~!1l'lli~ bl,lre;.\'iJl,ICml

became lhe tQ~tofm

does a hog Hnr.rn.crurned g(l\'cm~ t ~3t~. all furl~(: grc~t~r glou:yof

S1:.~i.enoe·'![t ha:ppE:ned kJ..'Ol week iu SO'lllth Dakom. \'~~n:nrn, Na.teonal1 the Sd€'rtoe P(i!ii!~d~ti.O'~ (NSP'), ChOR~ ~he Ho.!1JiI.es.~~~ Mi!1~' i:~ L~~1ld [~(l as site f'olIl"1 PiL'opo~d $;)00 mi.~~jiQin eep Undlergromil,d Science ,md D Eng i~le~\i'i"gUib[in.lOr)'. nhs'C:r1,'er~ 5<1.}' dun Dave Slil'}'\'-~r {h::n)I,~~e62-·ye,n-okl he<m QrSmlth Da:ktJita.'il SCh:::Uil.a: and TCCl'U10IQ_,~' Authority. and :~is s.ta:W'Ivorked tire-

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<1.:Jllmml\:',oo "iii 'IJl~eU1l. f,,"Og'l.~p!:littoiil s,:utc

iaJl~ ~ deal with

('OiIWOrd.~O:I]I. liar slme :p:ffij,ecl~!.hcLl the stare apPo][I[ed ~l'inl."&1i~fSP~mick(]:m\>el,". exeeative .. vice pr~ii:dM~ .unl g>e:f!il.er.d c~,U!ts()~fllf B.!lltriir:::k Th~ pr('Ij!;1cl.aTs{) benefitGd (oo[U a $70 :[fm,i:lll:~on ~()n~~imiIJ.mm ~~lil~flthrn[pis~ T. De,liluy Sanrtml. ( ··Da.'(l hm;; \lilU[!ked Iml:ir<lcles.~' 8~~yS KrwimL Lesko. a p:nysici.);[ frolm,1( t~~.C:· ilBiw~t"llce ~h!:rkelcy N~ltiIOlIiU,,1 labor.'llO:D"Y. W.~l.O ~lc'H;ls the 1~IOI!n€,~~l1ik~' ~[c!1~Hic e\[)ll~ool"l.itij;}~l. Snyd~'!" says th!ubisf'o!\: hl!lls b~~n !'lilllp~y 'I>W connect t~.ledots."HerutS:'1 few n~.C!:re~u The science £'0: t~U!iln must ~tl!l)pl~l~ :I. CO~.lC..~hLfHI d~.i~~. ~ndlh(l NS:Phlls ~fI 11!~dl~~ ]ilmOJ;u~}r .furthe IJ['ujeot .in iii. flfJUlfe' b~Ld,gel.

in 2004 ..t<lsl:ye-m:r •. tbe M,iffil(:'Sprc;:vious(l'I:vmw. earrj:ckGo~ld O1;;\1m:rship Or!~e s;.tc-. "h: \'14115 tlnrl:l!]ll!:'l pUl'I~ illl~h.e .. 1

.~ A, W ,A 'IUl :5;



:~ :~






.~ .~ would explain a stighta:symmelry

IPRWIIOIVE POWIEIR. Theor€liGlillPhys'ldslS Makoto .l:<obay.ashi ,(!if lhl;! ]alpanese a((eterall~r Ilalboral:QT!/ KEK in Tswrkulba (!I!I'!d Toshihide Maska,"ol of Ky>olO Ullf!J~F.sity hav~ 'WO(l1fth~ EWJrQpea~ Ph!,5lieal Socfety's Hngill Elle'fQlyall1,d P:ar~i.de IPhy~';'c!i.PriiZ!@ 'for Ol'll~ of the mnoll'.a! ... rlispil'eli 9 lle~e~ f n ll1ie hi~'tJ)ryaf ~delii(('. ~n:11973, pl~yskisls. had onl~.y r~(i€rn~ly disc'l1~eredl~lill prQlQIi1'~ a no neiJlll"'ons oofll)ist of p.ilr~ldes ciliitoo II po qua rks 11no "OWl'il 'qua rks . PIthi rd such p~rtide,~he~"lrallge '1ulal"k, WJs.ImQV~f1, anda fourth, [he (Wl~)lfm quark, IP~ooiic:too. Biil t ~@J1I b@]ore. thto nonon eta qu.i.ark WJS 'e:nlirely (!c:cepted, Ko bayashi a I1d Ma;s~aw;l! a rg Lled~liflnhe ~xllSler(,~e 1\'0' me reofthe m of

salVS Helen Quinn, atheorist oiIl5tanf:ofd '~niv~rs.ilry fi'll P,1l'loAtM, Callifomiltl. "u 1II'<1IS a Ib:riUia I1t step to mflak€, b~l neta diHii(IUU 01'1e01"1 ~Ol! C~



askt!d Ih~ M'gI'r~q it!!@S!jorl."sh@· says.


1(]iAJIHiIEA1iHARVARI~. .Efi,diflgl a 9'-ITWlf1th

se-JH(h,lhe- Ha rvar.d Med~{al Schoollas;t

week picked a new dietJlrl
from wiU'linit:s. moks:
ob(?sity exP<N't ]ei'frey ~:~ier.!=Iner,S,8, jClk~edl ~heHarvard famLty ill 1978 ,QiHer5tl.!dying ]ni~l!tiio;s. role if! m~l,Qlbo~i5Jl1l allfld ·dlrotl~ tn~he Nal.iJornaLIFIIS:llil!ut:e~ofIHefilth 1mBethiesda. Marylano. Hj~ rt!ceIl1HO!;u$ has Ib~ern 01'1 hO\'jI the ho~morne mpHl"11 affe!cts!he: Ib:rarlli'l,. appetite, iiJll1d ,~be.sUy.lFlier~lso has been itilV'Olv~dl'ill e'Horts 'to make :sciell(~ ·ilI Ibigg,er Ipali1! Qflihe !,Illiderglr~duPl~e olJlrriCl,.llllJ~l, He st,;J1"1$ hfs new job 'O!"!I Septe!m.ber.


:~ lITI~ll(!r t1nd al"fllimaU.er caUed (IF!v1o~a~ion, .!it which Illad been otp.seNedi ]1Il 1'964, :~ PhvrSc~s'~, veinl~llly ~derl'l:Uioo ~jx lYlP~)of ~ ,g qu,arks •.and! KoOO'yashi i;!llld M~slkawa':s'~lOO!)' i~ IP~ecis.ely des(rib~s (P violaltiorll seen f I'iI •~ accelerator experi·mf'Ii1'Ls. Kudos. to them both,

SPIOrrTilNIG~NFII!)Uln A Mkhiglall state Iorens;i(sciE!niliis~ ~~'hoi!lrnCillyzea IDNA salm~le~ from her hl.!lsbarld's uoden'N?alr- alner :SLlspe('~ijllg ~i '111 (11G1iil'~ilf'lgon her ]S ]11 hot wol'r"rrOI" 1 (I'r m"'~ifl[lll!sed go'tennrme!lt e{llllIipmel::rt to (:ondoct h~r tn'!.l'~s'tig.tltiiOi'il.. Ar!!'l (hamberlain-Gordo!"! fOWlnd another remaLel"s :I)NA in lhuEl~lPl€$ arid surbmiUed Ih~r ~il"ldJiln0 as evh::le,rl!ce· a 7 MiifchdliVQn:e ill! Ih~~d~g ][( Illlghilrn.. ~llt.1Ih'e~hew hlllsb.aml's ~;;rVri!yeri !I1fGrmed a ~thoriti es i1bolJit~h e test lhi@' Mfdl'igli1lln State IP()lice (MS r). \liIhich mIlS [hie' LiM'ftsing lab \!IIhen~ Chia:moerla~n~'G.ordor.1i wiD'I"iks, i'nill;1!Jte<il,an invesligill1!O ni nro \'Ih:et her si'tE!Ihad i)rokeirl aepalrlmen:t IfU~Et!S,. infl5.1l1Y The State Jou{fm~ (llH~te-dl hiU as t,enifyingi ln a 25 May he~lrifilg th.ill slle ih;;r:d d1on@ the'! analvs.is; on hew Olll'iln trilliN~' IlIsfJng chemfca[s. I~ at .~ ~e'~la t@d fm dll!ilPO~.tiI! . AI"! M51~s;PQkf5,p,erro,n :say!! the d'epa1rtrnelmt is ilftile-stig'atilf'lg the rn.aUer .

~lp ror~hi$ pa9c7 E"m~~i pe<lpk@.!:lil~s.(),r'!J

B(ROAIDI!J~IIIN:GilJlIIit Mark A.bboU $a.ys a (~roor of '~:rJPloriiMlg t!1~ rnYS~l'liri€S!of !iJ'C:@itl1 ~H~h1l5 proj)a rmt hi 1111 r[!J~:l~e s 7·45 miUf:!]" GE!!mciences IDireC~(lrrate at~Ule lNii~on:al Sd'enc'e FoUil to fi1I(iaMIl (NSF), "M.y expeneru::e has al'fN.alys been ~~(llogt(;al. ~j;IO~ing a[ iflli:eIa(Jtj'Ol)S.OIi Jl!~lural sy.s;tems. " say::>~he prlOifes.ser ~f bioLogicall O(,@iljl1lcu]ra;phy alt IJ ~~(Jn Sttlit(! lU.rntve~sity (OSU.),in (oNa~.ti~t Nlo\ljI he' ~I.be h~l!plngl to oT-che5trate th.eif!lt'eradolls.oi a I'! 'ermlires!!:ie!mt]n'C Wrn!nUlllil,y~ Bl!ginning on 1 OdQhe~r Abbotl1lNJ W00 tal<JingQ:n bigJ"!ii.':~~I'1ICe. p,rogra f'I'k"S ni1fV() Mmrt~he so~ld ~tllr(~,. deep·s~ Qbser\l'i!'!~ !'Ieh~o'rk:s:. ocean drillillg, a!oO'lher stll:p'il'llille [ontn!1ll1ed bwoadening of h;,~ and ~peri!i$e\. HliSGhserl~lionl exallll1inedl the Q<cology of La:~ Tahoo. but.lle lamr!acklOO sa'l~llijteQb:;llr~ vati'O[l oJ ocean bd:r;l,QOgy., AfiI!<tlhe IrnO\I~QYe'rsee:5 ... he' Gab "'~he ~hQIe gd mut 'Of 'e~u~h ,11(11t s;.:,iern:.e" a:s iihe d~lIlfl o~ o.S;U's 'Coll.eg,C! of Oc:eank (lInd Atmospheric Sdences. He'll ilM b~ mLinqluishing Ms p>ost Qn IMSF'SOi\ll;'lrs.igIJl:[body, t.ne' N~tiollliil S(ient:e Board.


OF ElrNlIElfH'3Y

C~Ul~~dll~~U ImJgll'il'1~ the W~:ii: ··'.~lit rh:J ... ·l'~.cy ;

Nuclellr' WeSIP'OlnS M'i:lle:st:o'nl,e 'Triglg,erSU,.,S,. P'olicy' O'8 b,8t:B'

dk1d dillla)'$ tn half l"lnh!!l; rou!!hly 4!0 11.u~cle~r CJlil,a'~'UlriH::t.urUIIU1l;g pracetlures 10 be f~nrn.ililized . •'1:v.~r.)«ltle(uH.k::re~~i mated hO'Pii hind i t ~li.t.m; going to be," says; former DOlE oll;l,a~,i:I::11

M.!.tileiYfI ('~e:t:d.on, now." S~nOillle Hide.
ln r,espolll,::;t1. l~.e~;a,""~abdi recree John I3n.1wILmereplflc~d t'~(; headi oflhe program \\I.'ilh eo-leaders, o~u: @!i,l'ersee'inSpllrsiC'.s.m.d U~.(l~lth~e:!''!iveapoJm.s l.~m!.!facltQrij~~g.M"!Id~li\g:i·' •. ~
cl.eerli,lIl,g. laadditien, s<liys.Ric:liD.ard MM,~~,.I < termer Roc]!.:y F~~l:sl.nCiHI~ h,ng 1St WilJ(l! led the revamped ml_l'lrn!Jf;l.chJri,,~ effbn n·1JJlll. 200' I to 200'4, the :~!lbbrought i~l '·'.sO~llri:e old hands

As \%'ith hi gh st:~rnoo~i;\~~DI~eo\'lrl:~ ,mCO:ffii I1CCI~ ing:n:a 25th reunion. U.S. Ilim,;le~lr"..re'.pons sc h~ lsrs h;))\1~:f~1i1I~1,d t h1H mCO'lpmri n§ the: In 11,1lJg:i:c f the past l:fIke:s the nii!;lln kU,lllull ofpeo" o
l:lh~.a~'!Iilli.~lgneuto ;J,dl!l:pl" ..nd a huge C~~ptu:i'!,yor d~~ayt1id gmlif±e.nion. f I n a elassl fied c~n:I]1;{)ny c!ll'ly lbi:s IflOllth at los A lames Nali <lna:11 :l.abor;l,tOity ( t, A NiL )i u New M exieo, Depa rtrn en'l ,0 r Er!~['~y (D() E} Y'ffh; ials celebratedthe belated >conlpl ctio I1i of the IfiJllIuy.lilium tati gg,e:r of ~ !1W': :~ear bomb Opefil[ ion~,lIyid,elnka] to Q~,mS~<I51 ~1118 yc~r.l ago. The slar oflhc btl

t~rne 1!l;t'YIIIernment asked: LAN.l to build :a set of r~p]~lc~m~!'IlS within 5 Y(;l,US, \\1hb one
caveat A 1993 bsn Oll nuclear tests IIlll,eonm that the !ldCl1llist~, WOUMll·t be mblelol:e:Sl


!lrn.eiw verswom,.
That rcsrrietleu ~cjffened the ~1.1~~I!.dy-

iITipOSillS ·t,ethn iC<lJ~:~~l~.,lelil.ges c OfDl,~.l:l,chli ng the R9Cky fl~ts sp~cifica.tio,~s. The re~I:rI-' I:hr,~~y small plutonium research f~H.~mly.f!l"

,i;,~hohad dom'Ae8()J;He of[his rkali ()11:>ul' th.cRtlckyfl;;(I:!;;


By 2003.

~"~~rd1t;r!>·h~]d ~'l~~du~d pby .... speeith~ leal


had the requisite

~itS ftwo-

and g~~ve boxes. ilior ex,:rjlnpk~., but
darion would notsuppert

cer~:u)ny was the so-called pit,.....", .. ~1'Wered
piece 0 f usetal l be ;<Jppm~'I; ~.~.]<It·e~zc lJ.11l1 d s shape of ~ F.~b~rg~ (lgg .. with a hOIlI)\'\1 c;\(]!~~thal: DOE certified as ready t(n~the mu~I~~r sl{lol;,!kpil~.The {le~Sitl11 \~lS ~ milestone for the 'I'IWld.c.lf we<l~JQ:IJs P~,!:la:;1 III <lind paves t:hif!' "jay 1:'0[' raere ilmb:~ti(llm;WOI'k, indUldin:g bui~d~l~g entjre WeilPQI1S :WrQi,H sc nne: lfuiwh:.h'G u t co 11due ti ~.g mrclear tests, U <1150 ope.b the door for lANL. tT.'ldilioll<~Uy <1, research Ilab.. D() OO!l,~ tder '~"(rl!!l.dIiug i,mo IlIll,anUrjJi.ctU:ri~l;g,.

l:~,e:he~vy=dmy p~,lUonlum forndn,g to{~ls used ~llRocky Fh!~;!:;.~.n's,te'l(jL\!;'O~ile.rs, ~l.;Jdto pour urolten p lutonl u ~n~ut Q S~,!l. pe d Jn~('I his ~~d 'IIM~~d p~eCles tiortether. To oonform~Dcl,e!'!\! env~':L"o:mi." !1JiI~:rtital!.'."I!I;;Sol;l~gi'H,~~~ down O!~Ule use ,em of ~~I,i:lliic<liIUS;, used newsolvenss to c::1C:'<In i1lld metal ~LJ rfaces, Even se, ilm 200 [., DOE,IUdiiwn> CQrI-

The pan'l,Uel~ Ill.:mugemenl structure helpe>d~tle lab ver~61 lh.~tt~le lm~Mt ph would ·work. :imys MiI.h. For ,ex.!unph:. fem':.:; dun <I dilferem metellie g,rnill size cOI!.dcli hamper performance d~ ssol ved .. ficr velific<J:tion

c:xp~rime·l1!is-,'\ihi~l~ h1Jdli!!ded non.;nucl!.l~r e.~.p~oS~OE'lI!'l. numerical Sil'lllU~:!Ilio:ll1g, and nii!a.n,;:ri~ll!lseleuee !ol;wdie~-g'~u~V(ldi bY$,[p
dSit~ t~ia3t tln.e d~rr'eri!'llIce W()Uid'mI,·l d,egll:.'1,depit

Oid,es ~1rUS,mlcl€'"r TJO]~:r.;y.
hewever, sa:y lh,al. bu,iildi,lml.glJll.e:w p~.l"S (!'ol1tr~di~l~. 1lJneenuntry's

stated i.lnellinio:~,

reduceits Th~y a 1:S1j believe rhatthe $1.4 bi ]lion "P~!l' .~.~~ prt)J~lCt~.1!i!~\l,i~tlmi t;~);:
Ml 1l1ii!,~I(:iH ~!\';{i~H'it







l:~].e words of

performance, It look untllthis year for L.ANL t{) oertify the pit as :lItm;kpile--n.:ady •. u'il.e>eli~g; a ~()i1I. et il'll 200 L s A~tllUUI,£!h~:I'le Rocky Flats plts had been :p:mvtl':lm1:0work i1'1 undergrou nd tests. lA.Nl. researehe rs r~'1J j!;~d the y were If I perfect, h L~Yl!'.red[lfllet~ I S!!l.II.1iCIU.Iil.ds thc::ir hoi I()\!;' 'plu~o~ !~il!!n shell, 'J,lihici1lwlderJOli;~ ns~io~.l\.vh~!1 crushed by,c0I1'l'e:nu i.onOl:~ e~'(.plosi!;i\e s, S'mtuires :;It LasAlamos fl;J!Ul'ld Il'uu [he ar,igin3~ pits centuined "impu.rities thll~.<1.I1.II'fiecl c ~1mtich;~nie~1 l~r'Op!l;rti~j;:' ;o:.ays DOE 'WL;~.POOII'~ uffid.al [)a\r.id Crnnd<lli. At I:irst."therc:, \Y<lS ~U!l;,'Jjlt~~~~,Pf m~tk~ p.I!J.~tl!~j.i\.!~.linpitsf!s ~f(l to as ~)IJi~ibl-e~" he says, B\I~"'II\."<iiJlJn ~ ~C.ir;::lglist1'i made more cred~b]e progm.ss when ~"he'y

OIJ,;~uvi"l Greg Me~10 (j,f the Lo...
A ;Iam,os S UJdy GI;'Oli p in A lbuque[ir;~m~. Ne\,vMeJo!ico, could be "~O:l'i:j(; jn science" by d ivert ing

cllr.:d,ded P]~~"needed the


be as muchas

poss.ub~.e HIk.c t~'nm.e [pre'~'iou:dy][e~,tcd. i'll1ci.lIdiug llJ.ly j mpUJ."il~eS ~mtphno liIilll~'ll." eral]jd;)~1 s..ys. tl:Mil new WS8 [pits swmv

Ilv~ds froV1l~. res~li!rd~

~il~~u~h~ COU!'!it~~tS mtel~~r\.\I'eOllpn~sc:omp~ex
esaberh manitnr eurreut bemes <lind build

'11989, dl,e ,g,Qvelinme.nll fuund ~ll'dro:nmelflwJ find tJlher ·\·~C!I!ltim~ s al 1:~l.e'mI,atOil~'.s.olil~Y i sm.~e of b~.i:Id.~ngp.I~,h)n~1tI11'! I:l<~, ;t, DQ IE,C,n: itilt)' at Ro'e ky l~, FI~I$. o;r~ido. h '''ra s ~~~~ ~lUi (',o~ r d{)·\lI,'WlI, :~'a~,lin,g ext. oml a batcilil, of w phs t:ov~h~ \V88. III 47S-ki:~own

oo~li "s ~h!;lm: 'This was ~ " v<lll.m'lhl.e i:niti<lltion ,into ll'ne processes: ·we'IJI JllCOd for. RRW [till;: R!e~ioblr~ ~~.'t;:p~aC~lllil;:~U Warhead1:' The !tRW p:mg.nml seeks 10
i!J.of;i,:V !l~:-l li.viIhtlm O


'l~W bombs


scnl~ch 10 r~pk~c.;; ~

asillJg Wl:Irlu:<l,c$s (Sdeuoe:.9' Marchi" p. 13·4S~. ~ r'~mletan' g4;ltrp.~:ysid.s:t. R<lY J~a~:loO!f the.. ~ o


lJI~iv,en:,ilY ofOdiror~ia. B~!:'k(llcy. !>ays tb:n ~h~ ~tfllgs~~ m~ki:;;rhe new pi~:> highliglltl;, H}
tTrU'l:UUlporlmH::e Of:li:I,l,<I:~llt<l,it(,1illig;l, weltr:\ind''-ld and experienced t~d~ni p~o] t~~H canres po nd qllJi,c,!dy t o eme rgenci e S (I r
~~~VI' de,,'e~()p~'illl'~I~ 1$"


pn:::lgr.aUl by


~p:pl'rip'r~,~I- ~v.;l~!}d di.~l]y f'Q:n~c !tCW B~tihiD~l·Urti.i.!"~rsilY


$11DOE'~ up"Qnli!~g

2'~IOS. bt~dg;t.;:t sev-

aff>:'Cl p~"ml:JifX~e:. panx ~'Cljcw~~e LQS AhH1!ilI!,)$, ;u, The fine of l:lln,e cu ts is uncertain, :~ov.rever. ~iven d~,If~u,'1;ll1it ver:s,ion::;onhr;: sp ~ndi,i:ig biU
th<lI:IiI,m5~ he recaaciledaed a th!i'ii;l8t to \lmo li1e everall hill.

ern I, ofwhid,lWDUld sion !JfplUlnnium

i~~wHlil~ge~l\i~in Pf!!iIl1!~!'~hip. says t~m:~liliidillg; morepits OO'L.de!beenme om] b <ldditi~IHlJl blil::li~e::!5 for lhe~~b, His fC<'If is
QfC;i]jfor:ilIl1l 111~lgo\l'~:rl!lirne]l1 oIJici.l'l~'smighl


~~ilcll~r~llg mere hi,ghly thanscience, B~Lt a
131> spokespersun ~<IY:l1"dl,e,lFe's, no pl ...n te make l[rIiIlflnul~eulfw~I~:JIl~,~ pr!i·~lH.ryr:ol~ o:i" [L05 Alam,o:!>.'" -flU lONr11liS(lIji

Semuo(F. Pe:r'e:: Dmnelliici



White ~flo~~e

ing mJl;line,:2 July ccilebmli,(lll, used the :IIn~Ue~ t s~o:UIle <ntack some :l:l600mtiU:uo:IlIi in Cl~ts to to

hi the meannmec Mah.wlro



ln ~ ~.igJlI:~~it'Lt~~s fQr ,quick Jn~d~ ic~~1 betlefiES frOm:iit~~l (1i;;US ft~ I'ad'ing. ES CeUlntern<J:tiom,d (1::5.1)-01 compauy e~lab~is;hed with lalliirr.o;n',:~n Si n§~lpoo:re 7 ye:rurs i,i~(r-is h""U.i I1Ig work ow llllID:1,il.1ll emb:lF)1onic s.tem (h:~S) ce,I,II:her<la p~; 1nvesto r:!>lo£[ i,!l te r~s;t ~S. because "the Iii;:el il1oo[l of hllvilJ.1:g I~ro~hjc!::; th ... eli!l le ~nth~ s'l~j)i!it in term\v.ls'f.Elniush~n,gly s,~m~,n.'· s~ Alan CQIIll~ml•• 11 stem cell pianeer

rc!'i~~~~(:r~ •. AI~1l Trotlnso~. fI ':;,.qy M~I!1~shU!li\'L;:~hy stem (,1,0]1 S(;~.,

cnatributedto the ESlwflS '~ryiogW H!~keH) marker, s~lys he hi "prDW[I'~mdly diS<lppo:~mx.'rl'· th"~'ll!re c:Olf:J:p<!..oyis gi.vi[lg_ up. W3iUl be says lUa~~tESE



pursued "a ~~.igh·risik $l~l~gy'" in
:foc:u.si~,l~, H<uffl)\,;'ly Qilm,1:\11'0'petential a pp 1iC:~lia~s.W ilh t ~~firl;l hi still yO~ll1Jg, Tnmnson says, "the p~.·.irnl"Hy ...i m shou ~:dlDe to e~l.]bli,slt!"l broad phufon:n of rohast and rena.b~e~ciJen:c(l t:hal 'em) uuderpi nl:r<I,n5~j'lti Ol,~, to

\...:~,() uurillasrmcnth
ch~e f,e.:.... eutive, e


~s I!Srs

ES.[':-; setback m,IY d<~n~lpenm unV'eS~ofS' ellilimsa11s:mfof SteTllIlcell
[he n.aJl:i:es. ~":!.~iS :~!)b~!l blii'tf.-fll, viee

'C!illUC,11 ~pplit.,~dol1Js~;·




~ ~

~rvi.t!g \\Ve:~s[llm,~r.l'I•. (i stem cell Flr!!llm Ib~d$ildEl' t!~ ID~!fIchl, S~ggillig inlo',emu (Oii'lll'il1li!'i'I,(e]1111 stern ceus Iila~ pl'Csident for R.&D at A dvanced researclser at S~lnTrord Univer:sity !1'U.o.mll,ted Alim Clllmlfl ~(l t~\i'C ~!h~ (Orlll'l)filt~ warld ror.J lI:!~'5i( rils~i:ln:h lab, C~I~.Techn ology hl ,,\~~ rceste r.. gnp.do Aho. C<l~if[}fliJi.<l., ,<lgr~~s; MillS&3i::hll&eus: '~WIiJPLtthe Field I:l.:ld~y needs is "I:S cellresearch is. for l~'a.enms.t part, stiU UtJ"echit. Nel.l~.e]'I<I,mi s, Austral ii:'L~l:iiIlIVCSUl'.rS ~~\~luHlc di.scoII,'cry l'Csearch:' Ql'!le or l\W ~lJC~S." ii'Wrics;' ~nd al)invru;lImmt~rmt~ of S~Ii!~~CJre"s govColmm'l.:iiI me~]!berofdle Iie~lm dmtc:IO:lled .Allhofl.~h E Sllii~, out of the s;.Ulil.e, .'I[ least emrnelliJ.l pLll UIJ seed fflrlidin:g .. 3:~,d ESI had l'Wf~Ujl!1ii~,pa~ii.;s ~~)!. litey~~awh!;Scell [ht;i"1~ the s~t:1~ Dully. \vi~1 bOC:~~l~i;i~e~df[h~ Sillo r~~~d $24 miU~t:iI!Ji!!l~fhlEiJ!Oc[{'ib~I:'. ~ceur:d:· o g~tpooo StCil:1l Ce~I COillSO;[lt i~Iml.,,!Al~chill nds pies i,tlli. the :~):iipeline. Geron Cerperaticn in illg to the comp'lIly"ES~ hilled Cclmaaas ro~~~.~~h!It *~ii;titut~l> ~n-~li~l;!t~d Si~~with MI,;:!'ihlr.f~rk.'(,~Ii~~ri!lia. 1,;)\.~Ci$ t~) itflr~ 1 elinichid' sci i,';:!l Qj St iu A.pri 12:0 01 ~~.~ bee ame pure's A:ge~l~Yfor Scient:l;l, Tet.:hJi1ology .md c.d trials oh :rh~r.tpy fur ~pimdcord injury CEO. in :WO:5. RC'SCfi,n;:h (1\ ·STAR ~.mn.ti,also otlers gJt<Ints. He T~~.c a:mlp~I'~~\W.'l>.i'LIIClU'pI.i.IIll,g tum ,~liES e~If:I:y .i f1J 2008, accord.~ml,g to Sl~oIl •.esp (>'TSOIl to Davl,d Schull. A ntl by ea rly ~1 x l year, e (leUs imo iIDI5u~i:Jl-P,L"Oduda.lgcdl'5 ~o ~rf;Hn'dia"viii .~~so IUp ~. lab at A"STAR''''[r1~ilule of 5.0t M.oI.~c:ul.iu- and ('eIII Biology. Mmn of (he betes ~f1lld ca:lid~m::muaclc (,C~~S1:0 (':mU~ler 'OOIlll,~ AJdv31lced Cell TG"Chl~D!.og'ylm.o;pc;r.;I.o l1~ea I)C'~M ge:sti'Ve I:Jem'l: 1~]i]lIre, Both c(lI]di,~ltlru rep.~Sl'1Iu tnmg ~pp~iC' a~],()n [.elr OJ lremme~u for macalar 24.sc:i:uUisls \~'Orkill,1g?,Iil hES cell [!~I"dpies .\l ES,~\\~II.IIl:om,u IlIUI:! then resesrch wirh "ammre nnJiiljor maiJr~et:i> l;A(irli.l unmet d~rI,i!l:;alneeds. blllt {~:eg~m:Jr.I~i(m. UUR<I S<!LYS, ESt umh;.!r n~l;.t1I€'~defsh~.p, \l.fi]l tlfi\li.1 n)C~'s ~!I1I(lli!,g\1i.t~~~-fU!lCl )[J:!ing: insu I in-prOOtic ing g{' ~,.;~re g,O\!~mmel~~fU!lnd~:l),g"~lA ·STAR); fH,;\'V lnstinne o:fMe(]IC:<l~ BI.o]ogy. COhlltl,an Sl;i)1S. t'ells "llrovedrelllJly du'fficu!C' CQ]I[Jf1~1'Isays, en [pi~o\'i.di'lJ1g ~:lES cells <Jinddel',u\fed Cl'll~:s liar ( A "'~TAR I.li!Jl!iJm~!'!:c'.i;~('(llm~lSm{W~ ~]t~~)i, Bod1 ,1;l[:JJv~sk)nr<ld1w!"'';I)ies W{'II!":~d!1:~d~!l ~~t ~~ ba~iC' !i~st::!ucl't .uul, d:il.!g dl.'lvdopm~~t.. (Ill Colm<!.I~ !!lay.:;. He: I!.dmits to ... "dngt: or dis~~ILn.na:~, dQ~. PtI'Odmdn~ ESI W3:ll sCIWlI)m 2000 to comnm.erel .. ~:tlZe biliion cell:!> for ,e'<'l!ain. !:nore Ild:::S (;!~U .fiJl!~ingl!i' l'fJd1,l~odl by~. coU~borap :;'l,u~h~~J.m.bcrs~.tlillaern!!l !Jl.iredp~tI':uly "b~com~s ~~ppoi.llJtfllentt~~! lh{: f]e]~dis: mml~~llig
S,~g~v.ly tharl I: had :ooped." CQh~H'rl ho:poes tiO ~p~j th.en~ld ~~fI~.g r wtlrll~,h i:s n\l,'n.reJ;c!il~h. a1thm~gh he' dec! ill~.esto d ifiC~l,t;S{f.etai Is. ...jDiNNIS N:DRMIILE!

Colllm:;m says,. arid meel;:~ng I\~o.~~~b UI~iver~i~y i!~~'71~Y: veryexpe'liIl~i\!e:" lhGSC d!(llI~e:llg(ls WQ'uhl h;)Lwt ...[k~11!,i,nll,gel' Nal,lOn~~ Un.¥~:rs!ty 01. S.IIlg.a. ; pOfre;Hadass<l~, Medkal 0:II',;pmi7.;l:1[[QIl :in ~~<'IJ1lI ~'nvestors, ha.vepmic:r.IIll.O;: b:. o J(;)'l"IJ;$al~m: fmd~he Hubredn bb(JJJ1'!lQI')' h'll 'ESl's~lbadr. tD6~!l!Ol ('Olk'i'! a Pf~~~OIldu~t1eld
~ ~]Olillitn:',o~,Yi,l~

if ~OIl• .t\~tSI.~~I,~<,


R,~dp(! ~@r rll(i@i!!lIW/. IRduildi!l,'ij 'hh !ito(k~ wHi r'e;ql:Jiir'l1!'ll'utlj .lit ~E!'asla riflih of iHlawaii':s willter5UMell' 1'19 pml@tliolil ~ike Ii1an>awillaBay, §il~;AJl.;illil Fll'iedmaloo~e'r.
tnure ilI.Jli!.k:o~lS~IfV~lli.(l1l I.h~1l~~l}'

b.iUs dun were imredueed i-m:;l\:~'~ii '$. rlighHo-f~:;:h [han.il CQil~i!;rv!Hkuil bmii~ th~ Hou~~.]ii1
helped d:lra:Wi legislatien 20% 'tl f ::it <~te waters for 'co nservauen. L ike other states. Ht.w.]~.i cOI;ll~f'O~I~ first 3, I~<lmk:,,~miles lh,e; (6 kulomu.lle:H:::Ir.'l) :ul.'l.I."Oasl'~" and rhefederal off gOWflllmi:l1i1 co:rn!mls t,l~e ~st up to .2:00 miles 2003, frtedlatnder

of ~he other i~l U. S," the bi.11 got fureher

t hai: woul d

~illil!Ve :i!~~: ~'l~id e

(3il'[I kilometers). Tlite~b:r.il]e Reserve N~l~'!.\I'{uk t\CTWtlllld have nmd~ Hawaiithe
.~ei'liderlin miuallme censervation ill the Un~.u~d Stases, where :~ess lIu'lJ1J.1 of CO<lsl;a:I,~V'illeJ!;. %

are :pro(ec~:ed.But rhe hill drew fheul't:: of
H!:lw~Ji''sf:i,s,hi n!:llobbY~In,d W~t~ ~CIiIUtled, ,iFIISHERIES M.ANAGE.MIE.N'T

Cons,e'rvat:ionisl:s, ,Bnd !Fishers F,aee Over Haw'8Iii's M,8lriina RJch'8S
HAN1!JU'MA BAY; IHAWAII- Th~ school of bigeye jacks "Nas Ililgint \\'1I~!ere'AlnJl] Fried~<lil'ldel: ~Jft!H;;l N!;!t ionai Oe~a~lic ~nd AtmospWD~!ii!;:
Adm il~lis:~~'<!u:i ':; bi.Q~ogl.<lphy tJrnnch s<'Ii.d on it '1vo~l.1dbe. cun;.I.ing :s~ow.ly ..t the :n~~.o~lt~]. {)If Ha[ll.mnma eilY.• 1 pretected areajust 15 kilorest,' Fried l;tInder
S!I.YS •.


:stl:y, isa C;HJI~to~~ry ~:2I~{:of bow seienee smn~li~es :is. no IlKilCh. fQiI ;l. pow~fful bureaucracy lil;,id!lo fi sfut.il,~g :lillt~re~t:s. As Hawai] '5 tourism gre\v. and 'C(l,~:~of Uvi ng skyrocketed-the state has the


n,UlOIJI'S [l[gheM average [enn~fis:lili[lg became <11:1 in~lpon:<liL'U supplement for

meters from the S!,;YiiC[!IPC::'I:S of dowlJtQ!wl~ HO:lmo~:u:~ T herem ~~:St~il.rltC been c~o,s to !l!I.•. e 200. ~is.h. each abcutSfl !l,;e:~t[lilI:1elers 10lllS aad cUitte-ulyl!'!whid,IS I"£edi<:i~id~§•.!,l. mt~ri n~ l b~!Ologis~.gl ided lhrml~h lll.e.m ... "You hardly ever see (Ids ;l~YIfI~~ruil'! H3.wai:il,'· friedll;illl1l.de:r said after surra.cing. Jm:'lb;a re p:r.i:tl)t;i [)~a'!m.gkrs J;lllit~ such I~rgc SiC~ru.oollsi1a'Ve ~ become nlre in iilnli1l.,.bited :p~JI!l;; onh~oSi!,c~.ipL:l~~go,.h~$!ly=-. IFr.~edl.mdelt knows the bll)' better than

TI'o~1iIl.!il. s:lm.ore C<'l:st i.~~,g,aud speOl.rfiish ing are unregulated, aad the st<ltie's estimated .im:gerlislill CiltC·~,. 260.00U ;nll,g;~el1'l<Ire :mliPl ljeensed, Ol.1~ydli.~ The appeal itbli I,H::\1\,' c(Jn:\ic'l'v~u;:iol1ll ar~:s y,ear wen: resrrfcrlons put on giJIIUQcu,. ~)Hlmpted a reacticn ..101··'iIa!lrt:h. t~,e s~O\"ne'::l ~Ild:ud n,!;l ,J ban O\~. thei r use 011 M .• l i. l::>l.md i m House ofltepa'esel]l~nive5 3~JP[1Qved <'I "right~ Rlild overni~~le.!se\"~~ eee, [O •• :fi~hr bill Ih1!L1\IOillkl reqJ~dn~ the i;Q',fI~~ 1:0 \ Oppt'l!le!Us of '~heM~riffile Reserve N~~~)irQ,.riti:e lllll~IU;;Jin:!lbJe data, such .1IS stock work Act gained momemW[lll earlier [his Iin.cllrem;i.~rng p[1[)~ ahie

smal]. ··ITyoll W~Il~ 1:0 f~buUd fi.d~stacks, you need to SlOp r:U~hil1~.ill.1arleasr J:O"*i of Haw~ifs \'\tlU'tlrs ~lI1d reguhu~ f~s:hin~.~!l the: thetefore.would .re.s.uh illl

poor~rrres;;id~~~s.. he U!iit:! af'glllners, '~~hiciil T snareturtl es, seals, and 1.1OU food fish :u addhioll ~fltar,g(lt spaeies .. is wid~sprcad.

a sscs

&~1I1!iili1DS U~ro ug


Sp~.H'; es' l

e~M re

yearin a series nfm~(;(ing!S dl,l;sign(ld to
increase lilnpul fimnml nanve ~i:a.w<li.ian COIli1l~lULlli[ics. The ITiIClClings were ,!tIrgaJ!;lizerl by lhec Westem P:m::ifg:c .Re~i!oilImJF~.~~hfnry M',nl1l,ag.cmctu Ceuneil 'IWi;:~pac).~ull~of ~i~1n


j:-Ie r'Uibl i$t~~ ;.I. ~i:!)dy i~l April h..s.t~~ ~lw fE ,;olo:gicnl ilppficmion,~ shewi 1lg.1 hat

~.·aflges,. before <Illy new :poo~ccted area is C[!L'!ju~d,.The i:Jli]i "'WtlfUW lie: Up!lnf!!>hi~~ .r,egui<ltifOIITl~.HOi: jtl~:~m..'IIriiillle re.-;enes,in endless slu.{jjes :[;1m,!ed ·Iifl~. ~mlld~g h i:~npQ.;;sib.lc r fo:ltllil.e state to 'pmpoer~y maml:;t:· ~he puihl~t:':s ~n!;l!i~:mt>e~ SoS!;;:ts," S~ly" WUI h.lm h~i!lir,nGr.

director of ooe.u!I roHcym: the M.~rune Conservarlen

~(Jlid fish biomass in Henaumaand 1 ~.!Qlhel: prlJ~ec:ted ..liens 'iI\'0]5 2..7 Itimes glr:emer than lilin.ebuol1l<lsi> in cmn~:l<<Irabl£ unprotected

ltute ~.nBel:lle:'mc.

sll,d. fesimm~ couecils lhial advise the U,S. Comm~r~~ D~1P'art!lJ1lI;:r.lf. WI;:SiP(IC"~ chair is Se'IlI!~, M.~u[n. p're5ide~:n of the HaWilii Longliners Ass!QCliado:~" Sl1Ite offi-

~~',.::!hi:mL£i:On. ~~i5re!:ie[ H,n ....\ii1i Sem,ne To
~dgllufic!lnlly modU"iedllhe bmJ. But :;dc:~,risrs •.n:KI Shiite omcillJs expeet the fi.. to ght CO.I1I,l [Iue' in l:~l.e·l1I,ex.~ Iegis I;J1:ir~resesseo n, i



are,<l;;~. inthe UIIlunluibitL-'d 2Ot1O·kii!o.m.en:i1And
k!l'lg Northwe;steaml .. ~.a'Jii.Ji:il"llfI ]!>~<'I.n:dlshaiJrn .. c a !l.ationai rIIillJn~.~uen~ since 2.(U)6,. th'L;;ro~:s


start s ~!'IJ;)!m~!'!!ry.

fi ~hb~orniPISiSon3vcI:<lge t~.l In < h'l C()llIiI'P~!r;'ihie h!l.b~nI!lS----,r!1l im.1k~[.,to~' l ~~:;;!J i
6. 7 t~n~.es Illore

A~.!hollgh :s.imilm: :rig;I~H:(HI'i.sh


Roell sed \'~esp2Lc ,r~efe:mI.d~ narrow I1s'~,nl,g ]rlg in.dI~lStry iEl:~:etre~t:il. We.s.pliiic·S influenee ii!s supposed to be i~ml~H~d to :f~del"ll!~ \\"~~~~r.::. acli~fi&'~s<lind bliU !\une off'i(.;:~I!>contend. th .. llle olrganiult1ialll, ..




fis.b s.lOek:f> in [btl m~~Ui!, ~;lL~\I'JJi ian i!!lh.mds to a'b!Qu~ 15.% of 1 \\ihat they Q]iI.ce \vere" To F'ri~d!la!ld~r.,lh~ nlcssflgc is $:irl11'p~c: Jibe mil i'111. HHW<I~,i lJl15bnd s.' reSer\~e5,. wlhi.i'cl:l < pro~~C't (l!dy 03% M the COO$[U!l~, ~lir'~[on
Inuna.I:I!i] hm'\lc reduced.

be~nl!lpp:m"led i:~JU1udels:l~l1Id,(!nd Mt,rybllld, limy h~vel1l.Ol impeded tile ,ere.rt.i.{m o:f prOleCll;:d .. :uc~s ~n tho~e S:I:!lt!~S;.s~ys S:t~!!h CI~rk S~mlg;t Qr~he'C(Nlst .. ~ Ocean Co.,.1i~ iOIll !!11 A.t~~nlic Hl~th.I~l!lds •.New ~'~'me:y;B~inlis¢ the f.iO:\.V'.,.i i leg isIHti.oJ,l\'fO~I,ld eft'eclli:\~e.~y encl ~JI fi~hJi[lg restri.ctio!ll'" sl)u;).says it "is far

~Qbbijt'idmegli!ll)! ~)of .he riglMHu-Hsh bUL "NtJme:rous lin.nes dua:.illg I he' Ill1oce~s dUll ~ produced thL';:bin,. [ !>!.!w \ijespac e~~p~!oy~c~ .~
o'p'en~,y Hllkii,ng Hl ~e;gi:sI"IlOU

us~rL<;Keiko B,ank of t:h.,c;Norlbw~~t ~ H... i ian h,~:ands N,eh'o'ork. whid cmn- ~ \,,'<1.

3.bmu it,"


p,aign$ ~O:i nl<tviue: cfmsrerVOlItion, TIl'e bill ~








passed rhe HOU5C. but a Senate droli 00\'\,1 all';,'lJIjti ng3ct i.m~WOlJll d e:uwoll rli}ge C()[ImJ1mJl,~

th;;,!!:[ \..~s drmfi~d by fisll~~l;~Il"" i

"The sc.nl"y~lill:ll!l, is lh'lll~.heb:ilU couilidl pass Il.ily~~ed prl)~eCI~On ellb.rl:J;. next )le~~r,'~says PelC'Ii YOllng. who recently In M,ay. B~llk fl~)ed complainrwirh tile e omp' leted ~I rer 1lfI1I s d ireeror ,{I f .IJ.I.'lw!l~ i·!i. a Cemmercc Department's irlllSPCChJf' Gc:~~ [)l1!'p.'IIrtml):'l1il: of L.i;'I,n:d:;md N~uln~[Rcsoun;:cs" l>\jh~djl.mI~m:l.t;es:he Mnte·:i\\,\'11:I~r$. t eral. claim in£, that "V:eSiIMIC 1~<Id ViO~~'11ed ·'11' iI passes," 3dd~Will[~Ull .. AH~. an sratutes that IM"t)hib~t fedcl!"211 c~~~p]{.l)i',e'~!l



'The United 'Kingdom !:l:5.k:s I,ttgginlg belli no f 11
space studie~ H ~lhegQV~mn'll'rnt does not

imr~ase SPCI'wF!spef!dlng,I~e H(lL!s;~~f
C:omIlll.0115, '.'l.Cienc.ea~d te~llli1 olngy mmmiUee

fn'j.lmllolbbying :;hH~ legtslatures, lBiilin'k ai,ciliwl-1l.11\'\It'i iian i1jsh!.ir~i:'id herbermester, (;<1 ned. for aui rive sti.g<ll,~:oIJi <I rid. cong re s'~ "~,t.':s.gO:il,lgto fliJI.["t1ia.erdeplete eurmaeiue resources, That '~t]rlillcc~ptab:lle for our Sitlmml he;;ni,~g".\~'bpnc denies it ~n~ged in lobbyi:llIlg. The lli.ighI-lo-fi S:b, bi I~:"has 1l01;liiI,~ rutm~ ge:UIl,emli ens." inS l~l with us," <\l1l.Y..,p.,ml D .. ~,t!e:~iI. do We:-;;-C:HRI:no'PtIilER rAilA ptl,c's Slen~o.rsciemisr, addiua£. "A~I] know is, ChiriMDlllher flalillli5 a '"'titer Msed: in Hon.oluiu.

Wilrm in a ~pon~hl~ iJlie€'k.. Thl! parlilam@'li1tari~lll~ .~uggesl seUfl'l'!J up lille! N,i:I~,,:nallSp,l!i:e T&h rmo'l.()g~ IP~rl9r(liif]Jme ~I),provid.e~e-OO(!ofIf'l hlllld in£!, ,ilIlthou gh n 0 'to~:a is~wJ;g'L?s~.ed. ~ lhe ~.K, 5~ll.i~J5t $4:!!!:i ~!'lillio!,!ClF! $pa<:e' .au rinQ ,2005-06,. ~l!Ib~t.alllrti'i:lIlYo)5lhanlrls o EuropEl[lln mUI1~;efpil uts,l~f' repc!fl r'e{OII1l11emlds bolsteti'll'llg Bdtish )h1e.rlglh~ Siuch es p~ nelm ry
t'xplmalimll and ellirl~ obsef!ll'ati:(IIrI Wlil'Uft (oofiid-

[ku~~,iliii. an ungJl.hdYlltnlil1:lS11II.i'I of FralteC'! It {IS: 11~l:~ghl.h!l'll~feOOe~Iii,.I-\l somejime in the ,geo,iogic rill~'I,.t~hn()1>t (:~rt~i~]ywas. i BiI,l,tlonga.go, ~!~~~e; l(:Crc~$!I.l!~~~!~Q\,\i f low~'..~d~. hig.h-S!;M"bd~:rI,§ Itidge from [)OIi,~1' to :F:U:-dn'Cc t1iD;<I~ \~'{mld be dry iilnd ~(JdOl}i.A g roll" of ge>osc]e.litd&t~ :h~1.5 ew n evidence aflhe 'C[~iprlt::A huge gushing of

~~rQ:U~Ui~lgdow~stH:i!!m. ~nd~re!iC!l,Hll]ike :S:CO'L.lrs. All dl!l:~ele'alutres spe<llk of e.",tte:lI1I,lfl flml;'s.. 1~lle gmup :!ii¥'.

@ril1gli1@/ii' @H1oflsin IhulTl!il_rlS!Jl~(E! 'rli'Qhl. and ~~~ undl-ef'5, The ,comrl1li!tee~l50 'cal~$(In,th~ ElJrop~aln Spac~ AgiE!ncy to ~o(aOO oru.' '(ijJ~s radtilijesin l~e U,K•• iii topk of ongqvliIglllegotialioos w~~h[SA., S;ays Ric~,ardl H(lkla~y Q~'the Rul!herkuld Appl'eitonr L3bora~my in rCh,lllo'll.
-i1l.A,N'1 ElL (L!E:Rl'

Gupw and hi~

('~II~f!gJI~S ~ll'Qvis;~i(l" ,<I.


into s!i,)~id mckliCl fonn~;hc Daver Si;tiI;<libelblfe rllshi Ilig dO,\ll'/l1 th~ l~{;\!l~dry lgl~,sJ;i Chf~.m~l" B The ~inaiH:lIl1~:m1~ meg<'l.flomtuf it·~",:p~ jj€n~d.tmu~~dllthliwbL.~n ~he ~ np.;~'t~rLh{l hl~t of it:; kilnd. The dll'l!jsi:a elil:,mnp~!ebriike own of
aneiem Lake Misso~I~I<Ii about 115JIOClyears


W.,oo[., Ul!;:)' :5iJlgge~I.,(;l~l down

llu:m:mlu:d iJJI by glacial ice \\'in.ere the sOlllhm"J:I Norah Sea is lOdily•.l'~lelab's w<'!lu~rncould llulve eeencpped the [)O\rer fudge a rew~~.I1dred H]OU~mibd ye.m~ •• LOWI;)rnmg the ridge go. :md illlcl'e<'li5ing ~,he flmv mUli 1200..000'1,0 I. mi i~

B~O'~!il'Qi~sts IOililgl (J'IlWIAJ UIiI,der' ... G
La!;l '1Ii''''!1'~. !hI! EIlIJOPI11t11ll Mol!2{1ultllf 'Biology

Ihm euhle Wl~'i(~n;Il~T ~(:om!d 'I\f~ro stre~unin~ (We:f tile' r.xdge. The m~~<inood \\I'OU Id 1~l;1;vCm e !oc~~ th.e r~rlinS\'~rld~!.ri!1g li~'M-;;S {lf~i,gJl~a
.ic:ve:llli!;:,c rhepresear,

I:sbllld Udw.il'l \\'OuM haveboeen

the groLU~} sWI,ggests .. bern..

Laoor,~~iJi'Y (WEIll.) spfe,alv to I;~~ antipooes wnei1 d'el!lglJ~~~h~:11Il gHlu,il's; 1.9 :m~mber the ~<I~Ofi5· '!(Ot€cI to, el<Cfnd an as.~oda'te lFIl'll!rnbershu!)' to Ail!5tra~i1t. 'nl~:;ye:arr innral ten1ll 1ll<iItIs lllexlYlei!r. wilen ,AlJ~tJ;~Uawull hegin sellidillg fa.(lllirV llI1im1~er~ amd! rr~Seiln:h: i'elw.~!lt,() [MEll.' S rwe Europ€ran 1~i3ist(irt!!i.era:r.dll l<ioori3il.on'~IIi!hibi!
~@~vfii!g EMElL,IIl!~~f'(h S~p~!ilrLSpo.ii!ro~, indlJd!l;! ~~lFilll,!!lmtrali,m ~mwellSitie~ and h! ,gov@f1rlltlliilfill, ~,!I!iich Wit~Si,p~rudill wm biMIlil OS. 2' l1Bi~lEOI'l (und till!! lnilia~ tErm. "With 7 to [Ald~lrali(l's] 5pe(ial.e-x~iuti~,roil" eX~IT1'p1e, in lhl!' lneldi5 c:[ mJ!ldii(.tlil '~pid:emioltlg!!i'1tnd ~l'~,m

a~d'Cl:'c'll'Ctllil£ lCi'rrured terrain of the Clmll!l:i,d,ed Scab1~'Dl.d~.,11.,n ooq~ired a :ilm'f of U:I tn~lIi,ou~Q 20 ~n~nk'l!'l llb~,c c meters or 11Il.eake's glacial l
tQPl SI~lC
~1I~~U"V~~l,ii,;lr0flch ocu~1d.o!'50 o

eastern \\!<:1Shi~,g·



the now of t:Iln"e

A:lilHlmn Ri\~er"

dence in this "iv~~k's issue n r ,~III'~:" III 1"~lI~We' tor ~"l:J1'1.mW! 1r,..l;ll1;'H~Un
~ ji;'. ~..

Gool~ist Sanjelw Gl,~;'lt<l f o I.!fi,,~'ri~i Cllll~g~ leil~dC,l~,aud his eclleagaes preseatevi-

ce~1res~rillr(h. it ~l be ~~ e~c:rtUenqtQiI'!;P~~ ( rm:ml to ~IWBL's rlXlUi5, QI11I basic Fii!setm:.h 1'1111 lI1o~ecliJl.ar biology," 50111j5Olin MaU:a,j. ~Ilil's i d!lroclor~,el'!eraL -BEIN])AMI'N IlESl1ER

,., Itflleat Rjs.llln!g
The ~.t~rl,dl '~n'lCirrord ro ~tall"olli cO!l!hlilting dimCII~;e(l1i:mg'E!',. says <I If€solu~i(ln J:na5~d ~asl. w@!.ikbyth~ rmlt~m1!!tio!1l>1!~.UrlioJ'! Qf 6@oo~1 1II'ldG!!oPhY5k~, an [j~del1~.i~1$ ~hOlddFl 't IEll it. The Sl:H1I1!liM5.l:i·elllir., IlJ!f!h!'E!'11a (I~g"!1iza !io:!1, W:hkh IndilldE!,!; sev@n il1~€Hli1Ilioni!ll. ibodl[e'5~ pass;ed JiI~!1e-pO'iiilt lreoolutao~ rOt:usin91 ,on t~e ine'J.l'it;alill.e ml'l!>~qLC~m::e,5, r wa r91~ng and o urg ing 11i1~il)Jn~t'O~promQl~ad.;;l'pta llD!r1." In addithm to p!.6ning!Qf roore "l~llaJte-mmi ~mng ~e'ar(~IT!.I !l~tthe I,i FliQn'~ !iIl!embers, p. rorn~'se mQre "Q.u~re,]lcih.,~whicih its QLltgmng 1P~Il5;fdeml

~ dc:nvn:s.tI.·~,mlilt f lin:. [)nvc:r o ~ S~f'<'Ii[. GUPl<II, amid 'coUe.~,£.m: ~

i 11~il1!C~ tDI~ook ...

for l~n.dlr v[dence e

~ <Jl Ul.e bOTtom of t!'le English ;;: Ch O:IJlnllel.which nu::hill:lg i.l!:e ~ ~~(':(;ts fi II(;~ \Vidl\,~''al~r i):~ th~ ~~d, f~lh(ll~;;;t o ~ ~Cle~~!,:re. USlrtI,g de!l!th~f JUde'a" d~t<i li::ollected :~m' is ~avw,g~li(J~I;d~i.'i!ni~~~ c tht;)' !'ru'!iprli,;,ld~htl bm; tom

A ,d'ay's 'Ii'i!ork? The e.~~lIlgale<d1!5lam:h'" ami stroeamtillied ed9l! of dri;5, n~blil1'a rine \f'illle~ ,()nlh~ 1100r 0 ~Ih~' IEliIgli$n (han flel. :!illg.ge~t llilaJ( a hu g!! Ill!lt lornl2'i ~Iood 1!;I;tJI shecil oob'l@·~.rI 9rililil1l and rmru:@.


"Wb~elmym~p\U~h~ :ilss.oC'huio~ofl~'~dforms tlJ.gelher. 11is vel)' ~im~I<Ilr ID wh,u Vi.ctor B,flbr h~IS ,dJ~SiCri b~d~!, the Seiilb~~~ndll:' $~y:; ~@logi~1 PhOJip G:i~b<lrd. o:Fthe t1.rruiwrsity o:f Ou~:b:Jiicilgc. "'I'f~~, e~u~dG;:d by j,i.:" But B~~Cf. p of di1.eUuiversity of Ari7Xi:l1I';'1 ilill 1il~CSQiIm. ~.y:8 Pl,'{Jtirl.u;ti· ve ~,~e<l de~rvetl ino:w' ''1JIl:IJIDioJm,:. th<il:~ ...:!tlICiIiiARD' ,A,. IlER R

new cl.eHIU. They FOllndkilometer'-

~ SiCl!Ile, n~t-'tQ\PIPC'C1 i.s.lrnmls in I~~¢ saitll~ dis.tijnc~ ~ tu\"e tJollg.ued :slntpl::S ili':! the cms:uQm!.l [!:m~ ~ ~~nt~ ol'l:hcChiflflfleled ,Sc'flb:la:mids" Th~yaiM) ~ ~aw b rmu! I)! S\\I't!'CP:U I1ig sHe" ml i.lnecd\f';!I1 ~,ey o 6'tlg,e.,,,,. "Iir!r<lid0d" ,Ch,;iJ!mels. ridg~s, ~mdgrno'il~

'''if?:, not

~1$!IHOlkiJm.g ~Q'I,

bm t'~l~s ~1"'('ry ~~.

l\!khaet Mac(r,a(k.el'll says 11'1dudes geldfll~ the word ~t about ii1ilp@lldi~m1g w~r,mifi9-1f'1:llat@d HOOOSOT dl~ilug'hl!5. -MAR~S!S.A. flEVALlliOS


Bo'b ~l!ld Ritfl h;l!v(; p~rf~~ .lbre·sig;hl.~~lt:'Y \\'il:l <I~W<liYS' a stalemate in whkh nil:]tl1e'1"f1n reach, ~ fill ish the other of[S 0 chec ken; rcsersbles

IProgr'lam, IProvesT'h,8'1 Ch'eck"e'rs,. IP'llrI'BIC:tly IPlayed, 1:5 81 INIDI-Win Siitluat:i'D'11 am;! n~ith~ !~k~~ a i;,vro:llig,i:lUW'G. then the' game "",]11inevita,bly end in ,I dr ~i\!,r, l'~,;u 'B, f~leresult (I·f.:l proof execueed by bundreds of c~mpYler::; twtlrUiLe'Jr.ly :2 decades week I ww\\u.d •. mceluag"or~1
cgii f(;Ollfie mj~I:,,~ tr:'.u;~l:~ ~4"1(79),

~K:k~t~ck~!ne lka:IU~li.'n.I!\: "riI('yL~gl~t!'a[(drosses" e

B,riU)iu). the gmrm;: ~!l\"'!hich players ril~.1 2


(",,'0 p:~:r'~rsf~(,lC off m (ll'.~-ekt:;rs

~l1ia:ree·,ib:y-ahooe grid with X"g <nld. O\:: ifl~o;pes
of .gi.;m:i!1~U~I'Io,'t:in !:'rn:~'~. Oi'~!'ll t~@ltt~J.;,"~~':(i

andreponed nnli meby Sr:i(~ne(·lhi:'
The findi~,g gLU~.rantees that
an <'I,ppmpU'ialt~:.~y progralml1med ~u~lclrli wi U ne ver ~,Qsell) a ue ~:lmn.u'l"h:..l;\l(l, marks O'l! persona] l \'.ictory for Jon~lhflll Sc~~dil~[". a cmnpuler seientisl <'It the Uni\ie:ifs;it}l ~)f A]bc:r[l!ti~ Edmonton.

set Qui

to "'s{llv{':'"

'H,op~I.f'55,. IJIrnahle eWlllt'lJ\al~!f m\ll~a

~CIbealtlheo ((i~lfIl~uter iIJl~Slil~

IhaJi leads toa


prog ta ifn, a hUlflnan willi. the l'Iladline,
lh~[ C<1!!l

checkers :in :~9&9, ··h's ,[I ~~u:g.e ,flc,'Co!upi bihn:~nrt'" :$.WS David Levy, presidenl ofll;telnterDlI~niQll!11 C()J;npu~er G<ilffieS A.."Soci.lAku1i ilJa Londouand iIln e:xpert all ehess-pl <1)1 ig.lg machililles. "It's by f~lrthe ~~lO.!ll o~~~ple;.~ o gamG sver sol'l!'~d:'T~c loot:;; and sl'rmeg:iies developedro.r the IPrub.lem might rrov~ u~ful fnr afllfilyt:Un,g ,~netic (,>Od~ Oil?' (IQIliiIIJ;l'la:tfl'ized '~m,sI<1:I:ioo, he :oays, The pO'.Ulll of checkers, or dr:;nlghL:S, M tilu: glJlHle is ,,!I:-;©kn.ll'i,'I.'liI, is to get the jump {Jill Y9ur 0ppollent. The game i:s pl~,yed a:n an eigJn-b:y.. eiighl; grid (Jir~;;e:dOIl1Id black SCll~are:.'l" The .c~.ec;k-

<areblack and red disk!'.

sllde for-

wanl dhlgomlUy lrem black sqlmu'c to blill.ck

square. The


Bob ~imdRi~a,

~mrl 'Ivithl2 checkers, e<'llCim in the rows do... '<eSil
~o tI~(lir s~(jJ~c:::; tb~ baartl, ,~ ~Y~'I'&:nii!.fl'lf~~11 of ~llrfD1l, .u1d Bob l",J[D eapuae one of Ri~~] 'sdll'C kers 'by hopping OV'~r~liml;l'f!n I.;'lnpty SJ~ll~ juS4 beyond~.iIll~d vice ve.rsa, Checkers t:hlll cross ~:Im.e board ~on~e "·Id.ng»" tha~ C..,lI1IUO\Ie!' ba:cll;.-

rough~y 500 bil]i,M bini{Iim. pfl&,,~b~1;; iu·r~uJm!~.' :m:l:nts of c:h,e'ckers 0111 the oOiml pro'V;~ng (;'h~~~e:r,\}i~,1 ,gl~<1Irill1w~d dn!\li/' iij. r~r :~,H'der ~Jmm.ll:lrovin~glh"'l t.UCk·~3,{;k_;to,"c:a:n'l be' V,,'QIilI, ngl:; reseaechers beg<mby t.1~.ms~nu,;ting.! d~]ttlha::;e of ul I 39,000 bi,I~li:OIl arrtlD1lIg:,enm.cL1tS wil~. ~{I or :r~\\lierpieec,~on I.'!~e(mrd,~1:l lh(: b proeess, !~,l.eydert:e:rm]ned.\~!he,ther each one I~d TO a w~~ fgj' b]o;!ck. 1).W ln HJr red, or ~ dlraw,. Thc:'y then CO:l~ sidered the very beginning ():f lhc:g.<ITIru~. pcncd~'l' il:~l, move by q a bliJ.ck .• U:ild ~h,en Wised .1 specialised seareh ~]~,gori!~.~~i tnt!L'€ out s.~lbse~~u~m to IUOVi2:!> and ~ho~1i,; th,U',.•1S, thel:\'ilU p,.h.• ers lr,)' ~o maximiae y !h~ir ad'Vflnt~lgc. l~.(:y i~~vit.:~b.ly steer the p~~~ l'O une of t~~ IO-ch~ck~.! ~Y'nfig~~fOi!·' tlcns that leads to ~1dr<ll,\I', Sdm~I:~~r redits impfl)'i.',eme1li!r.; incomc .. ~nl:1:eI':Sfo.r m1.lk~ng the I't:suh p~ihle~ [n fa.ot. he suspended t\fOR, from [99'7 'tit) .2'0[11to ''''"lit for:i1l P,I r~i,cll:~~~rejdnl[dogy-~ t b(;l,M·_:bh pro(,:i;'&~r'-tlom~wre.. HUll M~li!:~Y Cmnpbd~.
.1 C'OIllI~ut,er !ic]enlist at I.BM1\ 1r~(J'~~l,IS,1 Wmsoll :R.~~~h C",m~!I'~!1 Hf!\\'dl~ff;!~. N~w YOrk, SilIYS t'h!i1llthe 1I,'ese~n::he;::lj's' ~Iligelm,~,ity WoIS key. liOo... "Without a lot Dr theclever ideas oohi~.dwh'll~'~,e)' didl ihillk itwould h.,:,,'e Deen ilIlil!lil.mbcr ol'ycaTs before t.cChl.10!.Ogy

ward, The s.~me centinues until one [J,layet
capn.IIJCs all of the oLileri>pi'cCCs.

Sch<JelJer ,nul: ~Ii:-;tet1li,llhave ~lOwut~n't


cculd have solved checkers,"

:-;.11'5 II'

TOI~~·i.ghtrcs;c'lreI~c!l!l> ISIlllS, ur.lh'!cr.~~tk!>,. the mllion needs you, Thh; f~U. U~~ U.S. D~pl'H't.~~.cnt ,(1 f Defense (DOD') Wm~<'I~I.IU;h <I grants pro-

eh a U cnn.gc.!> lh~le lit ~~:ro:ss severa I dit'iC~j~J:liD1l,e;s."S<lys WiUi<l~rn. Rel:r,S"DO[;Vs de'lII.ty

Hnn d~!l:S I) f w~mMh::rs will he ann (; anecd
ne~",~. sprii,mg.

g rnm to :rt~~~dro$l;l~i!l:h~1l'~.\,\I'hi!l~~v~t i'll\~ it idem;; fO'f wd]i.ng iillport,lllt ~ec~1!rily clia01Ilenges, lt wi.m be medeled on the Nat.ion:rul. Illis~i.lme~of Health Di!'t:cwr's Pioneer Awall'ds, wh[c!m support b~ uc=:.ky. interd~,::scip;.lillllilty re:se,Jmdl ilit bimnedkine .. DOD
p~.;<LI'IS~O IHake :<lbm~1 10

under s~e:r~l~ry of dd(:riI:;~ rtli!1'li~bfi:r~tf~iiill,::~ and bas iC $I!;; i.e:mtce. A ~thOllg~l the re search ~lt;:don~i,;:d m~d~irfL~~ pr~)gr~w'n.\i\!ol~ldbe

V. S. S1,Jlb':~hnl;f~r:iliil:~,o:ll
l~SI;"l ·1~.l:e Uno,V>f:rsit)l




Prlrk. whose

r~~~.~rchi$ p!,'lr~~.y fUl1Jdr;;d by

ml:c'~~!~~il'i~d, awardees

""lUlII daeed it ~e'CUI.·

I,·.ityc~,eillrnffil,'a'l'l to be briefed on tine c,Im~ll!le,rilge;l they ..re supposed lO(lcMress.

DOD" ;\lOJys <'I1.lowing 1-e~(mn,;'i~erl:ito~{)Im~ up whful proposals in respouse ul,agcncydesi,gnur;:d ch11iIh:::tl,g,e5i~ an "'outstOlndins"
id'~m. "\Me iUC used re h~vill:g l:ies,eamh lopics defined ~op·{lo1j,""nby DOD," s,ays S.nb:II'". ~nnal'lli3D11. ,\1,1,10pi<lns W <Ipply:,. '''Whi:lle .

for $3 ft)]iUio!l o\l~r:5 ~(ljrfj.. Applieal'w, for
Ul,e N:<Ituon<ll. Se'curity Sc~:e:llu:::e<lnd E:ml,gi~ ne'~]"~llS f~e'lQhy F~~kl'w~l~ips mus.'[ b,r;; US., llltuzeml" ~ndlLIrefetrellae will be giv~'rI 110 ~a~·.ly-C,alf~er ~:s;e,alld~.crs. Ago;ncy oflici.~.I:> hO])1: the p:mgrmn wi ~I

e,m::iill ,goo~

Tile ch~llcnngeSi. 1101 yet tCllo~e:I:L~l:Ue Hkely 'to be~:~milar to tl~.ose idelIDt i.fk:dbs~ ye'H by DOD's QIll<'ld.we:lmlliii.<I1 :[)efense Revi~w. Uslisl ofprigri~ies im:'~l:!de:s bh.l'· ~,nel:rOC!):. social. cl~llur<lI,.<lind beh.,,,,,~.orOl]1 lu{)de:l~n~:~m(~khlg I;;n~my W fg(;:itS: ,cou:i1!l'· (11' [eril1lg imj.M10vi!;'I!d ej.... ~.osive devi,cc:ll: !llild p exJ~et i.~g~~ :[ol'ln:'l~i onabo~H s~ s.p~ci Ol,l S

mi l1.!ri IIY"~~l.U'k:s~,li~ll. oos~~K~herf;k!H,)\ll best\vh.'ll tleir fieldh1.l'~ to ofrer:'~}Ie also d'ilvnl~~he: :r~u(W!'~:~~PS: i,;'r~.ne Ii "C(I:rps will of ac_"d,(:'mi,(,l :1I'C!scl'LroeThelis dedkat.ed to ai (Jl:{:~b~s.e lind ~H~tiomd .[;!;,'lCuri.!y," i1

it ieS11I:lici

eNe:m1lt'8 frOll1


dm,tOl. SIi;l'~S,

foster r~:$~a'!"Gb Q~I.tskl~ ~h():bO~IIl(!s of predetel.·mined 11,"esem:\:hquc, ...ti!O:I~:iL, "'We do nol; h1lW splilci fic fl~~f~S;if! mi!ld; ra~h~r. \1i.I'eiw~v~

A.g~.I],cy offid<lls P~{!.1110 i~v~!~ Olbom 20 I1lppl iC1l,lHt, w~~,os;~,nf~:veaJI:lilIi11:u~i<l~c ~o m ~u,fike l~re~etH~ljons ;;It tl~e: fe]lt;Il!];!Xl!1 .• Tbe

If the Hu"St nnuulggeswe~t DOD 0 ff~:-'".~.-ci(!!I~~'l.or~~o.e\l'~lmudly s,~prorl.~~ mglflj'~s; «
50 :lfeselln:her.o, ~



ba'l.'\'>bellle!'! Ti~sl~y."SC'h~H~IT~r s.f~~~S. ''Th~ pn~:rranl Wd3Y would neverlese ~o"'~ns!eyor JI:1YOJ1C else, per~odl." A:und b ec~usc Jimm~ns; ,even!tu'I.~.I.y makemistakes, rhe prlJ.g:ram is C01iill'iid~re;(!1, bc;!;~p~~yer !!!;\1{!'r ~:ho lacst Inl~ !;]Ild should [m:vit~.bl'y prevail .~nas e·.ries ofgnlnes .~1mles~,~rom ~: to 9:51 :••gai·mJ!'>t :!InyperMlifl, (."'Ile:D1.r~.:D12S1ey •. for whom \'.m den Herik, ., cempmer SCi6l1it~!1l OIl Mlt'~· o[l~y three l'Cllllf.riI~~me·lli~ ~99 I, d:ie d fl f clIm: e:!i8 ~uQ~fhs~~~c r, SO~l~~ &h~f1i:r say..,.h~MS "'g~t OO~P(;Gl." tricht U!~ivcrsily il1it1l:1'~Ne'I~~~,alld~,. iflln~ pl<:!}ll;1fl);~O(im~dSe~:I:lell~ii". ~le $;;11)1:;". ~d ~~~ Varuden M~fik \l,'Vuit'li. ~h~t :s.ehm~n"e:fli. ~]'~C~~U!l.~g.ru!l'ldm~~ll;r 1'$ ]"Q~~~m" p I~y p I<lY~ 1)1 eaclia. ether 10.;) drnw. Hut. he says. ·"i.fyouh<li\le s.oiution win accelerate the dec<ldc:s-~.Qllg cha:rged.lha.t the stress of the !>ped<IJ1 title ~(ltpfQv;ed ihi.;l re$t~1t then e'il!i;lty e'xD~Ci)f~tim 11!1<:1i~h had kil 11;:'lI,~ l'i!liil~'Y' Chiml~1k d~f'eml~d ·ded:ine of~l;IunmOlmem checkers. M!:l<lll'l"V.~.ih;!. its C:~WJl ~,[I1:\.'1,1'0 snbsequem l.lmtcl1l~sag~ i:mlst s.al~i!I.cl]er is tulllJl il1g bis co:mJnlternto I:mfker:. .b:ll is worth 11Oll.liJl~:' prl,m;iple.l:l.l;u: giuln:e C<]I)'l be w~\'ed'_;bm it St.4:1.1en~rSH}'!:' he f-eels YiJm,d~c.n1:Jd bylhe 11l1el~ex~-highe8!''';.JI;lllll,klf:idpl;l:)rer •."'To this (I;IY. ] pl'Ol,)iI: r.n 1994.;] 'p:m£;mm he d,e~1'C~o~dcalled ,ronm<lke you ~.~:Olof mm1'l\.¥, ·-ADRIA·U:(H9 st~Ugel peo:p~k.:~yil.l;Y lh'tll }"O~.\..nuld never C~mpbeH,
\i,,:~{) C(l·\"{~OOO


dle D~.;:p Bll!I:(l pre-

gmm that defeated cUn.es3cI:mllfllp:UOIl ulIrry K.I1Isp~mv i:l1l.. 1997" Mos~expens expected lha~ dleck.e:n wou~d '~'iiClllU!lfI]~.}1 IJ~ pm,VL'(i 3J drnw. ~y.s; Jaap

Chi n o,n:k pla}'~'d 'I::~ed~~!l··1I'(l.g~,]ng \'\,~{)rM i channpion. ivImio:riI, Tinsley fO a series dI~:aws bcE'ore Tinsley \v~'t~ldm\"" because 011" hea!tll prob]eJ[!Jl~ <Jl!iJd(l(Inc\;lde,t1. Ti!rl~~ey\1,Ii]:lo ..




A:!fij,S~:r'~!I!l"-('i.1d NASA We~[:~l£r i'i::'ltdllliMSIt::> improbi'ibly atl~.e' center of Ihe ~: ..~e$~scie'Il.· tifiJ;! ~i:onn ridging iu \tVMshiJ~g~iiJll.D,C,

NOAA researchers ~,ave !~tlde:d its data, \'Ii"lilich is Ij<JIrl;:~c~Ir]y USeflQ:11 for de-~~ct.~~g .. tfopi.CioI.1 At~.~$1tie ~tCJn.m; c~ldy ~nd piI'ovidIII th~ lili-a 2: wetlks,lwY' ';Oif!gr~~~;<;tml'I~. i.!'!!g vit~.~ ~;:{nit;i:r;i1,g';;~~\'~f colder waters, p!Ulcl,silll<l.\fe held bearings on events :SlW~ incl.udrLng tilLePadfic. A Hawail-based r>!)IJ;mdi~,£ OUS:!ef o:f\:Vmiam Proeaza as U. S. N,H"Y {lffici~~said Insl year it p,.~~!ys d~e 1;11 "CI'IDlical m~e" gnP.~idfic forecasting. N'I-K director orlill:f:' N<ltion<l~lHurricane Center (NBC} on 9 ju~y.P'r(lenz<! had repe.~lbNill~y f(loo'r.::iG:S~'(;lrs mosttreasuee th~cndt ~ <:lbility cri·~;i.,c::i.z:ed hi.~e·rnp~o}'er. tllae National [0 see de\ie.~opi ng t IiOP iCOiI (te:pressi ens Ion!;! (NOAA~., for lai IiIll']: to III,Ul. for the impe'n.cif. ing faiJur~ of Qu~kS('Al". a $!;IOO~lite launched in ~. aad ::;ye~n:s 999 past lts design nfe.PI:oellZ<L. a 35-ye'lf NQA.A fORC'):-i.ler 'who bee aIUcNllIC :fuJ,~!Id in Ja IHIUl.fY. ~'my.!:l loss of the c:mf~'5 sensors could degrade 3~d~y lia.ur:ric,me [['tie:k for~C<ls;l:sbyl ()%, e i~ ;ru ilm.g ~tm]y i~. ress. t]J;i:llanJa]yzed rooo~ p easrs f(ll" si x 2;00) storms. Scientists Jamiliar wuth QuikSCAT"s ca p abl].l Ii e S s ay 1':1"0.1:,' n Z![ \\V~ s; both "right and wrcug" ~,~ his

qU~l~.[.i y hl,lr]"~ca~H:~ :~lld speeds greaeer f w 111,111 IS km, can't seewell throughr<li·IIli ... H ~ndit5 pol~lr o:rbit Ine~m; Q'ui.kSCA;T ··~ll~y nQ~be ~'l th(ii ri~~tplll.O:;:I;,i.j;!1 tht'; right Ii~c;::' sl~e said, Eun,·o·pe<Q.u IlIlJidU.S., N<'I.vy satelHI~5 p rav ide data "'not q ui re as good" QS Qui kSCAT but couhl pi ug holes. ifl~le NASA craft f'lils ...... sOlwd, 'UJdl!lg that be NOA.A:'s o~1ll1er :oo.~!ipick up stOifIlm$once tlle)' ~ seem hcm:de:dl:~u ~. Iand faU •• , W(;l .af~mJO[ b:1 ad" :ilr Qu ikSCAT dies. Ktcz.1 asserted. i l\lle'l~ow!t'I,g,1st .u~d respec f~d wem:lherll' blogger Je IT M<lMe:IFS ;agrees. nDti~ngl.imt.the Ulll:)uhlis~.ed ~h!dyPrne!1JZi'ciWd inv()lved ©n]y one of:rougl:dY~GVCfl ilC,tli'llC for~I...sti Il.g modds. Fold ing in,li]lhe . si Inll ~a~lm~!S. IIlIs the rest of the p d.1)Uu sources, creates .(i··.g h) bOl11


1111d Al!nosph~.!"i.~



i()t]n.~n:\1h;~ d~h3Ctcd.


year, NOAA

forecas~er HWI.;ll Cobb ,cal]ed


Qui.kS(, AT is

b~! one ele'l!!i~~tlt. l ~'I~IH~Jric.~n(l e·x.pc.rt Gn~g 1,'[oUand of tlse N0:iio!l;l:.l1 Center fbi." Almosphcri.c Re~e.'IH:h iilrl Beulder, Color~d().
So PI)nC:[ijZ~1 "\:~'~lll:. ri~u jIj.lld\vrm~g:.~'

.~ canes,

acerble ,[lhl!ll:"i!Cs"




~ ~le<1'1'ily

~iadar01.' visual cloud





HOn ..lld,eN.pl<1liliJ.s. 1 Ajfl.inl NASA-NOAf\ ~md).I •
dUI:: n~l'!!.1 ye~lf,

wi.U *'I}IJ;:·~Iulullhlil
But Iawmakers

~ ~'IUI fl"?Hl s.iuell~tcs,lypk."lIy :;;;:N o.A A's Geos:~,;.n I(lru~ Op,e [(1ry

~ ti.orml Efiviro:I1lIllC:I1lHI~, S~te~ liJ:~. ~ Ils il:lifO'rllrnl"J[iOIlll a.s bo Is~crecI by.] ~ rl~.et'!,vol'k ofbuQ'IJ's. ~lu:f'riC:<II11I,e~ 1;0 . . .~ ~ lUl!njl~~,p.~1Jncs. ll!1dco~&ud ffud:ol!l" 1,whs (0





May, R,ep'r~:>enl::J;l~vc: :Roll Klein (0 .•L lund ~(l"Sporu;![l~ proPQ:.. -.ed ~. b:ilUm a mhorize $] '1:3 I'IlI:ill~,i.on QuikSCAT. now openMing 0111 b~~kup w In,dM a Q'lHkSC AT r~phH",:'~w~,elmL its Ilu~'lp .nIiI,odel;ers m~.l(e comnputcr !>~,mu~ations ~ril:n£l.niuer. '·'OUI:' bl'e'ad ~Ild bLll,Ucr.," "The 10£5 of this daUl-'wbe'lher ~.11i.riI:ul;e.or of de;v.elo[ping s~orD"l~$.QtQ.ikSCAT !.1illdtld ({'I :liSnn fic'[I!1l-,cou]d ·t:o:'Iuse !'iI;;eon.$e·· di Bm for~C'fH;t~rs don'~ ]iVi,';l un bread ql~te'a:Lces."K]e~,:~], the ,t:olllilJ1i~iuee. "~o.se tol.d 1 thHl el'lsemb.!e by bOlJIIiI.c.ingnml~~OO'W;i1I'I,Ie iugf1lmm.e. La.st ·'!Neek. i:lt. 1'1 Semite hearing in I~<l.l;f;oW O~e~l n w~lC'rs OV(1!!r U gOn"~.ilorl1uilt~r .•~ whi(;i~ NOJ-\J\. o:lfic~~~iswe,re !~u~ba:;ted for hL,.~d&,~ml\~\I.illr.hove ~HH been ~m~luded in appropflintiol1iS hili.s moyi[lgtllirollgi1l COI1isw,nh.rel:lOnin£ :;~'l.rfacewind$pee·ds. by 1:Ii00t p~e[la.rirlg. a.deqmlliel.y for Qu~:kS' 'Af';:> OlIli;i).]y.zln~ ~hc ~~I:le~lion!;. By I'oHQ\;\:illg a g.r.,;;~~th,U o[il..;;rwisc p.rol"'id~ ge:fiIcrOl~S; d~[,s.c. NOAf\ :l>~t~nit~ bro:UiIch chicfM.~ry EHell K:~CZ~l. ri.ed to poke ~]oles i·lI1. t Pme:I:lZ.a"s i'I~.cJFei!lsei'l NOAA N 2008 h~LdS'eL to pohu ()l:~~,t,QuikSCAT ~over:$9'0% of tbe ..jfbll liIlINiJ'IISCliiI ;rurgnooems.l"he Sf!teWf~'s :s~nS(lrs dO:ml'~ oC'(:almS,m !1ru'!Iny ~lr~!.s f\\/Ioe I.'! d<f~·.
dir~( ~,iolQ '~ins!?d.

It's ~. brl!~z@. NASA's Q'lUli:k5CAT :5<1telm.l! musu rl!sg~ob .. 1 ocean


to push No.AA. <1Ilong. In

:!i.urr <lice M'rim cis, ~ndlldlilil 9 sp~e<d":11 d


Wel,comelo ··· I.' S E.IIIOIPI8 h F'lly F,aclolry
has an a mlbitliot]s: pillalllllio el~i~miin,at,e'~lhle t.se~s,e'~lv.
IBulSOlDte sclelllt~iists s,ay irs :3 w:as~e of nm:l1Iey
KALITII, ETHIMJiIP 1,j\""""";N oi sy.rnt 1I.],It icolored trucks lumber a.iong the busy main !load :un It'san eleg;Hilll arid en\'~,rol1llmlelltally ~Trh::ndl.y m.et:imod; birth eontrol !WoP' insects, ~~Oll!flec.,} nit The 8 teri le ~UiI~~'C ~ech~l ue l iCl {SU}. as ]~'s o'l]'.ic lil'Illlyk1nO>\!Il.l~asa I.o:rrngnd ~ ::;ol:~d track receul (see sidebar.p, 312~. Over ~he course .of 5:0 ye~rn. iI, :h,elped s."!i'l<eep lhe j;~re'~'iI\'II,(1:tll1 fiy.whiICh ft"£'ids an npe:in.wounds ill liveste ck, f110~1hal r lIm.eWes~em .He:lliil~:~ ~pll~rl;i. amidh ~~. be;irllg ~! ed to tyM!E'l(;( ~\1~fys ~h~ng nro]ilj]: OrlilCilll appl.c:s to DU1C~1 onions 10J~.p~mcs~m~~O]'il l'rmn voraeieus I~~gti>, Per~l<l:p~mme impO:L'ilrull. it hell~edwi~ out the ,euuirllil tsetse fly popu]:,'1It~o:!li on .ZilJniZobiJ;r1; ma~lni:s.~i1:Ii1.d iii! il~,le'I 990s. a 'proj<l~£I:lrn.e-\\'OlIld

lOne' tO~time PlltOtlrest OOI!lIII1l~!Jiiies: the 'Wlolrlldl iillfl

have been much bett"fJr spelJil,l
~t'olJ:yi~:lg. SIJ}'\'l Gi)YI1

on cheaper 3:lJiId simple,u\vays to :Wi.g~~t tsetse,


f<lf suburb ofAddi,:; Ab~b~,. belching douds of smekeand heukl Ilg ~l tliae peu:el'lni,uu; t:hlcnrwd the road. A mudd!)'. blll,]1m,py sud e mad! h:;fId spasr a row of

"1 h<l:te lO see


01' tsetse r~'ie<'lre:h,iIlZjm~!biYTl;e.
"~Ipoor 'cmJlltryW<!~le

so Iml~~h




~h!:il\>. llnflm)l;) (0

factory ~ioU' pots :.:I.1ID.d pans. Then ~I gate s[idel' f!P~~l, :;;~nd.1 brand-new g:r<ili)'bui~d,· ing the size oh, suceer IIf~ddle:m!m,e.ge;s. SllIJmu!~dcd~y a !ilca of ~lrIoo{haf:p;hml1. h.'s .,lllIiI]O~l ~oo clle.,rI <I,lIiItd,or,g<mi,zed for its ch~o~ic~urr(jl;U.~ldi.!lg!;+ Ina marter of menrhs, the \1,ilSt I:milcl;~ng w~11 be b~tt:!i!ilg\l;'id~ f!!~ti\l~ty-i .. itl,;':r~lly, Here, EI~lillpi'l is d~;,we;lopi!":l:O! ::il1phi,:..11i;,;IJ,l~
\'o'e<'lpO[[D ~lg3 install

~~r ~1.iil~'l(] ~!l ~!np(l;)'t~n~~ f~f'pr~.nc.~pk, ~ !I::> prf,lOf

E~~:~ h~pe::; il iopi.]


become ,"'iIillo,k::~



~s(;ll:.'iielly. "'\,~lich!rOlIrn5mil~ u p'.f!r'dsilic live. stock ,dllsease called Il~g.~r:ltil rhet has long (ripp led the c>tftl,ll~ry's, run!! €;COm'lltly" The scllte~m,e SOliHl:ds; s~:mp~!e. Pmduce m, m~myasf!ll!1!i ~ lion li1lH.~(l les a week .n~I1~~ n dU,;::I1I1, slel',ii~.eby blss,~.i~]glh.em with ~lIdi3tUOIlI for 'H~!.mp~:~ (lf~ca:lld:s. then ~h~i'l$tlthem il'l t~etst::-inr€~~,ed a:remi, llulI-dlli.:J sure they om1~~1l~1ber\.'!I'ild ~mll~~,.~ to ~:,H~p~ess 'e'm~les 0 wil:~ mate 'Ij,ri~[1m the :Il,llb erirters, buttheirrend~z VO~.lS ll ]JiiiOdlJJ'~~ no (I ~fspring. !Rcpe~lt w[ the procedure several tiares, <'IIII<iI: thetsetse
r(;lptl~,flti!ol~ ill w

the Afi'ic~m mainland. More limn 35

irse If by shewl ng 1~]3the sam,,:: is P01iSib:.~C1: t on



Op.i nions d iff~r ,~bc,1~~ ~o]minw b1i.Ut nQt ~ tlH~ for :EI"~riel~y ofrea~lls-s:lJI,ch <I,S :r<lclll:i!at tim <lbmlll~~e problem, A Illmlvst n <!I,lm:tFre,,"m~;~ ion I ~. t:~lu!::~are tsetse $pcck~s l$l :Etbio;pi~--oiti s •. ters .of '~]'10sllyfellit ile \,<II.~ey :!i1. Ii kely ~o f.ri]. And besides, ~ fSIlIOt'J :MJ sla:irml.·· sq l:I<1rek i~rO~l!e land in western and ~olil~:~l.'l!\'es~em B~hiop:i,<ilis ~ ~b~G:-;oh~lio!1 ~h~yS<lY. because I:l!~$ : .. ~,ay rei:un.I·csllCd \,f]th tS~l!iC flies. N',f!~Jil~,. 'Ciil~l.sod by inr~st the' cmHlltl:y. The Iltuoney-Ethiopia\', <I llll,1icel I u I;;,u :];I<lmsite of the Trn.HJIiGSOfI.ia ~ g(l~r:Q~~n[ SP~il1~.l2mmion (m d~eja~t,ofY $

~:rr.ic's, ave tsetse, all1d i111 l~nillny,~~iDey tm'2I!11Sluit h notjust :JiI<1,grUl<1ut,}Is.o si.eeping siekness .• ] b dew,s't<u il.lg humsn disease, Ami }'IilU'h~ Et:lliopi~nproj:ec~ is a!t thG eenter efa divisive •.often CI!I.~IJHk:,. debate ~~:ill1tg!'!gentemeleglsrs, CriUes believe Ul<!l

m.oJm.ey;"V~l~!e adds', V~~~ril!l!.'lry~iW~~lmol[Dgi!il~ hm M~udlin of ~'IleUn ive~."S,ity r Edinbllrgh. UK .. cal ls S [T o Elllklpi<l!'s "m'~!l·o!'!the-moen pn~~\a:" .. These critics blast the hrremational Atomic Em..t'!rg:r AgL':ltey (IAEA). whk:h is .\lllp:JfImt i.i1I,ji: ·the p,:wrOjlecl. for sedue i rig. Ethiopi.~ i~h] Ir~ing ~:t~r~:lcjn~L;lc;t~.-:!f'ld '~hey"r:e even n~,ore dismayed dUH orher Anic~m c'u~lntr~i,;l-\'l·!;tru l"olluwil\g ~llit, BI;':.~~ kno\MIl fudl:s. wl:O:~I.,g~.ing ~~pif.il,g n~ldle<.!r '" ''lith P'O\1!"eI:S, ~]~e U.N. agency. heildqn~ml:red .h~ V]eI1H1'1. Al~5Iri<J •• ~~,so~J:tU.l1Llot,cS dIe pew::efudl ::;; use o'fQtnllUic ~Il~'rgy.im::lmnng the creation ~ of s~:eriI e :h'llYeC'~~., irs l<1Ibin Se itlers.d:otf; ~. [lnd
~ O'~[$:ideVie[lll:m,. lis, theworld's premier S,IT 0 '" re$~~r,c~l eenIl\;lf; ~ 2 ~




pfodtu;:iiifS. $ 001 n1liiUo}l, a ye-ai",.according



Yei by the 1970. • IAEf\. had all 0 COl. it .:.;igln:;: on rserse, The .eibi£'rs.d:Ort lab ref~l'ied
the tcchnotugy o~:re3rin!!l tsetse fliic·s.. Wln.':'f>e~;s.• 1:I: :fI.llfS1 '~~u:y were: ,ed Oin live' mbbits Bind guinea pigll ..cow blood ,is used today.

me- u.s ..D~pM'!lInI;!:IH or i\gl'ic-uIUJJt::.


ln ihe mid-19:80s. the! "'Seney and the

I.lfsel.'S;e Uy I'egion
one-,fihh (lr IIHiliopia-i.o; ~ome 1.0 dCJZ~ns or $pe,lili~$

'Ianza Illiangovern

mesu pi ked U m1glljl<l. the

main b:'I:gnd of Zanxlbur,

(:Ora test site. It

Fl?' belt.. Aila~iI or i!1b!ollJl:i!O' milliolllJ :iqu(lfe kilonDllteI5,.....ill1c'lllldimgl of ~e~e


look almo t WO ears to build a n:)'-reari~lg faeilhy and train Joe.. S[uN'. say Andrew 11 I?'~.rke·r. tsetse ex pen <IliA E..I-\, ..Hl"C1" the a ITIIi~sl'l~dl IirSI: been anaeked lIs,in.!:l: in~eClie ides "ph1.llii.es s tarted

de h veri ng 'we ek Iy
811cseimated una:1


leads of male fhe: . across the ~$Ia~d in AlIgl,i~l m '994. By '19'97. Zanzi bar t\IO!~S
·OSI. of'

tset e-fh~c.'ll

genu " make'

keep/in!' Ih "JI)ock

iJe~l'h he gavethe

raw data to entomolo-

1n;1.~:S to

hill means f \:\ ani'r
plow l~~eland, ~Ie. .s

gi:;l, . Ilirii!, afti· of d:ie Londo," ·,ehl'ml of H:y,gi.e·nte and Tropical Medic~,lle.\\'~lO

mllk.and 1~l;S manure-e-ln short .~ve;rt)'. A I<a mye S\'1!~atiii Off\fi":ica has '~tl~ sa~n~ probl\ilU, The U.N:s Food and f\y:ricu:lltUire Or£"lliz a~ ti:O!l puts the bil 'IIrorlllliss~d famm,i.l'lg n.'!Ve,fll.u.~ in this "Green Desert" across frica at r.bUUl 4,..-1h,miOil al]Jnnally. Th ·11 (here's the hum 1 III 'ost: Sleeping si ikncss, or 1IiI1,I!lm'i~trypanos omiasls, is, belie:\ied,1:O infect SO(I:l:1e 511'1)00 to, 70..000 peepie' a ye1~r. 3.!tllOWigh hard d ..[3, ,m:e nOli w... il~ ~ tiM!;:., N~l v.~ci;;inc exists, ,~md drugs-il'Qws;~ more than :;0 Yi:ars 0 ld=-aretcxic and

was a SWU;!i,;e;~5.

i,:rm, ill

2005 book. T~ile~dal

But ~ ]T diJIl"[ many rake o:ff unril tt:llCIi the successful U,.I.lght il] the '19505 <I,!t"rinst rIme seeewworm.v b:ich \ ~iS s'Llhs(:qll~ntl rolled hack <!II] the W'l)' dO\l\1Jlj IoP~nmn:;). !' it turned Q'LiU. it wa n't hybridization 'but: radiation Ihf'lil proved the most effe 'live hl)1 to ereate sreri Ie in!:>I!·.is,
~ ,~U'. he Wlu'Ijo rity (I f S IT IJfO!;lr.Uill,S 0 t have ..ddressed :Jgr.:~cuhura ~ipests in richer countries, h~ pr,fij eets '~ml~v~trens uf mi 1lio1lJ~ do ,II an. bm those costs are ol~len of I qyick~~yrecovered, The screwworm eradlcan n, rom: instanc . saves U.S. I..vestoek i

nll~ .ex,aml~,I,e p:illl1cd the i.llii.h;r,eSl If the "thiop,i.m government. says Temesgen lerna of the Southern T ,eLI: Er,ul:ic,ni,on 1?~;uje:Cl. ;;J,p§qgr<ll!ll uhllll,; _~1~1i:i)pil.llit.. Ci~!lC~ and Technology OI:g(miz<l~iol.l that [ABA supporrswirh sciemi expert lse andteclmieal advice ..And :m ye<lO:S later, tluruiks ~n '~"IIrt lu funding from tllilcUN _.tbcA fr.icaJll [)cvc~~ opment Bank, ,lRd the :go\1'emmem ofJ.~p a n, lhil~gs .are \:\1\:11 uf1Id"r wa • '\\~lItl,lrs. ~fe bus unloadi~lY new FdCIi:S and i:n!llallin.san ~tItOm:u~dI r~\;;difl:t? :>)lst 'm in ~p~lil"ldifl:g elean i'eiiliil'ig hl.l.lls.fi old buihJing 91'1 the same grounds. ni:a\1 house a. colony of about

5.7 million, h still is to !~lrY.


de 'r~a~i ugly e ffecrive.


r{~htr::;;UPfu.l. .il,Il,

arseme-based drug, k:ills.between J% and I.OltiJ, of pat iC::.I1I.1 For colonial pou.'er s, tset '1: posed ,I foemidabla barriertuthe de elopmcnr of l!l~~f' AlfiC"JU ~ ssets, and Illey a~11 stiJrte<il, pr~,lI';lrns 1,0 deal with ihe problem, Tl~cy d.id Jm3N~ some e1llli:y!o~l:ete:.'l~s.. Mo~t f~mousl)j,(he :r\:a'l1l~ueRe
Ind Ijhc·'I:m.l.U W~'rA:ffii~em~ :islmnJofP,liincipe of tsetse iii 1,906. :~.~Iy by etluippu I'Ig p'bl'libt"dlion . '~ ,II, rkcrs '\'llh, sl.icky backpacks s, 15 O~.Olllli~ll. concerns .11,'-0 in~pill' d one of :1;( th,!:earlies; bur 'least known 'llld:ile~of':IT.

ion.OUO breeidil1llgfemak~ lhat produce a 'II,veekly harvest of ~,O,OOO :m~3h::$, ]nl:h:e :1fI,ew 'b~Li:~ding. those numbers shmlld £,0 ~LP by a facto.r 70 re ~. 11 A~.enn.l.cxpn:;rins. 0 _he praject invelves Illlucli, l'IIiIure:than I'I, . lernu says. ouvenri l1a~technique



~I~ treps

and S'o-~alled t,.trJJ~ls-bliJe er
sprayed with in ..eo rieide and

bill :1-: sheets

rhe H:J!40s. in \.. h~~was dU1 Tal.n,gamdka. i now T'Jnz3.lfI.la. 11D'UI,is:h entolliliO lo~ 'iii ~ thu,l ,,,, giS-I. F. . V1JnderpJu!:"!kli,sc(i\'ert:di 1:1 l~l:ossilll.gtwo d.im~.relJ'al spel~je n tsel..se:s 11k~!!lre:su.lu:d iin h~~"r~d:S wi~1l vert.' 1m\;' f~r~ '~ J''''' .. :~ tilky .. Tlli' gave I'lilin the ide.~ for .. lri"I~:~n
~ <I,II.1c!is:


baited! with ~ow urine or' m'I.:ifh:iIlJi anracI.~nts-mte curremly used to d,n, . down the pepulatlen tu less l~~U~ 5% of :i'iS l1'r~g~!:vn~d level, SlT's role '\'!'~.:Iilbe '~o r~lll.is~ iI. 011 .. A1ellllU .%'I.y5. because s}eXU<I i <lUraetion can do '1II'h.~'~!l1ec:'l~c ides c till 'r; reach and kin gs even the very last ny. T,ile 2S.00o.-:"al:l" '\,',a'lley ,·ha' has bee iii. selectedR:5 a first ~ilP~ct is protected by moumaun . redueieg chances of H::~11r~stal:[on. It halO 0111 on.~ sp~cie:s., G/oJdua pl1l1idijH! '. \vil~i·h i wh<lt (be f3C= lOP')! is ch1!mi!~Olll at li:le !ltOmelU. Lal,e:r. it: 'wi.n :ila:!1l\ '" to start pl.·odm:lug the l"ulUry's fhur (ll!~l~r '(Jlo'.'iI;~·,~na spec.ie$.f.sw~IL

,,,-':Irtch I:tILeP'lIP~~ ur

'bec1JlLu,etLle g.!.xd is 'to rid alii of' Et:hi:opi.J:50'!m;e, of plt'idle. Pnl,jed cooljd~n1!ltCir llemes~,rln Jl!.l,e,mu (rigbO ,and in,seC'tf,adlily rna lIager SOlCmHllJ'1
Meklll1li1ell""""lPos'illg \lIith ,a 'gamma Iraf 's.o'I!IJce l!I$ed t(l' n~rilize HIts-hope Ethiopla's t~'L'Se'lfglilt wi'll seNe <lISam eX:lImplefor Africa,.
1:1; .ri ghl on lh~ 'I'norlhcah,,: rn cdg~ of Afric;<l's '~e'tNe . elt ~:tile flies. To iPre'lrem dli;1n from ,carll:irn,g back, ncigb.bor,ing I,;OlIDl~ie 'wiB h~v'l: to auopf <lg '[!:lsive ,olJtJoll prognl[ll!S <ISwell. Alemwl, says.

Q!1~ !sets!;!' sp\i'~:il!s 'w~r~

wh id1

,~ coUected m~d llr.msponed by U~.!Iilll~o a'l'Il '~ <lIrea 0' upied by .ulolhe~·peci.e .. in bo'pe ; .. of, 'r''''Oltil'ig :-;(eliil~ Qn~lnil"1g, '. ilii,d~rplill'i·k .~ I~.e:,(er publish d tl~e result. bu,l b fo.re llIi,




or n~e most IDrmri>tiabl.e ~'On;.e~rn t!hll' wor[d1:at~Htnk elH'~r9Y'end ~ej(.

Pir'Dlven TOIDhmlilogrY M:ay' Glet :a Mak.eov'B'r Till€' s~ri'l.e' ifllse[ Iled1InliqIU~' tsm bf!"~ng lie5~ed i~ o,~i~ neHe s on IElhi


~Ia!ifillmgr~~ii;a!i,o~ Ile~p,s maike rnal~ iItSEN:U su~rih~., a~d stxyall aUrae· fro Ill! elll~sur'e~ Ull~l rel~Rased en lllIil 'ii'!.li!', 1.~!lY II\Ii ~~. Iiind J'E!!II~~es eV!1l111 th,~ ~ i III !!IU5 t ~'~ii!Mlte Iltde<@,UIS. AI~hOl!ghi'!s us.e inl tS~ts8 !iln,~ malaria CoIU,FQ[ i~ 111glilly (IJJUro,'!!r~,fs;ial, SIT has alU,owed se1,l:era~ 'lriium'Ph,~ in l!f1s.ed to,n'lflol,over the Piil~H SO yeali'!), ,;!tl'lll u~ riillligli!! or jijpplnCa'~lrlJ'r15b 1t'!:Kp!l!liidil1lglevell1!!odl,1l!Y. S!iI!I, serne ~elneve th~ rl)lll[lr~ "Hlr~ be: ril, !lew, gel1!elic versiQ!JI o<f SH-Q:ne tho1l'~ ~Iep~ tlile ~el'\ but ,~I;illlillinla~~:!ihe ra,~ri~tion. 'o.fII~j)d1\1',ant!itge]s that U t dees mlOll rflq eire ttl e IIJ'iie(If gil m n~ .. ray sm.m,\1S, Ii'IIthidn terrarlsts i(]OId.!rl illS!:! 110makE ,di fry bOifln bs, 5,deni~h·~.~knrelll <15'ec3lr~yas tine 1.'9.20:5tha l x-ra¥5 IIl'Id iOlfln zilllg radio C1li,(Ifli pr-oduc@' dotni ihl!IU IE'lh'il!l IlI!llJ'latilms ij!'ll Ifl'il![!!l fF!'S@~ts I~hal,,~H'@IC:lirllt!'i, m'CI[te' t.helrn ~te,~ile. The idear tOo III~ ste rility to mrrnlrol ~@lPllJwrti(lJii ~ was; de'I;I',~I(lp~cl ilildepudentlyin thel!93r(lIs af!ld 19i10!i ifl1 the 1B,(ithh {!cdn!llllJfor Tall1gallyn ka, lh:eSov~ el Uini CDI'l,fIrnlii thl! Unit.,ei1 :Stlltes. . lin the i 9S0s;~U.S. plO!l!~(S IEdw~rd Kf1Ii p~in,~and Raymo~d Bl!lsl~~nd pUll 1,1iI!! id\8ill ]1'1 il1D'l'ildi:cE! to fig I1t tin e ~(FEW'i\lo:rm ny, ill rnflr,jo:r pest wlilOl5~

l::!.nJ,a;~ ie,oo©:fI (h~ fle:sh or ~1;I',(ln(lck il!lF!d(I~llef ~ Ifliml!! ls, Ah(!f ar sU((€.~$hJl I~t run ~:rl!lh@' is,talndl or C!ll~a,~al),~h~y t~o~, on flllJl'ida, ,ilIl!ld later. all. or l~e ~.5. S,lQltes where 'h~ S(if'eW!;~or!J1l!Jei!l!~oo. After vt~klJV was decllatrOO in ':Ii 966, '~he'battle m,oved SQIiI~h, iII'~ewe~~rolJglh 1l1l;ellrllltiolilill (ollP8'r~Hon, the rtie~\Mer,E!! roUe~ bacik all '~he way 'lhrn1l9h Mexitc!o al1d Oi!rillr,al Amenca. la~ll. ')'e'111r" a 111 ~I.TeM'l.rOHI'I-rEar~ n91 ph! 1iI'~ Wil,5 (llpelned in i!!W Pain~1ITiI8i t hat. [!i!rodU1U!~ 150 11l"l]llioml \lIeElikl, [0 glltlm ~he CIII mr~IiI'l on· h

l~er, dootE!,tr!J~he CQ~oI1l1biaP1ibord~'f. SIT 1s also wi!l!l ely 1II~'ed to p revelllt or sruPIPlress infE!'5~<lI[j,onr:$or ~hre lIe-diteHo1l1l1eaDII hunt my. A ~I~!b.ai peSI. Me'dUy i50 C!ihl"'~ill to rel{er!l~hin9 hml1l, Clglprh~S'tm!~Olill.i!lt(lle~ ;;irnrd iP·llrlilu~gr<ul.i.'lrte'5; bei'nrg ~Me;dny;·hee~ briU'ngs Wl!liilU~e,~mpl)ini:lriifl trade' ~'eirlefin. Medfly l~gJ('lori~s. fuiave
~5prrJmg up !!!1[O,tlnrdU~I!wo!ldI ..1i~e ~o'!rgest,.]11GU'<I,l'~mi!I;fIj, Ipmdlll()e1! more l~.ar[l 125 b'~I~lonl ~~j~s ,a y,ealr rOlr5J~e~a~ ((I~HltM~s;huge' 1fIl!Iif!b~nare I~m ~ p~d '~'II'e'll,\leek l[Dver the p,ort dtj as oJ lO:5, All'Igele~ •. 'Ti3rmpa,. <lind Miallnftl) p're~.~ni[~~,(lwa~vays !ITromcalJJsi!:1,!ij ol!t~reaks. 1iIr!lQ rllOIiI.1 his. 3 grill ,3 III@W Mled UY'-r'U r~ I!I 9 I) ~iii nt INa Ii nli"IlI@1IIiUHllll'i l~ e S,1lI:~li!is,~ Ip,ro~"n ('e 01 Valh~fII(i.QJ" a IFlH!jord~rlls-e'.'(I~(IIlrll flgi r'E'l!lio.~. J\~elitlll\\i~lIe'.0] .s:nproglF.fIlll ilIlso hel,p!!cl elhlliill1~l~I,he melti,nr nV fwem i~la~d5 nlf} s.otJ'!he m jaj;li;lln between 19 nandi19'9'3,; aJndi~ th e ~>e'! he,f lamJs" a (Omr!l'.Hlyr:;<lrlleod The Gr~~ Fly ~1~lllti~lWi1lronmenta~ly c;(JiIll'lici~HJS ,


tUg,liIr ,co5:b\l YIiI'i;)ert:a,i!i!1 outcome

The erities bare~)iknew where to hegin, A lecllll1iquc 11lJ11t drive down :I JPOPl.!~ c~:~
sid of'the w~~~l~i!li.~g 11Ii~s;. s~ys Swph(':~ Torr' [If lh~

tsetseflies .in!Ele rn 6.I()OO·kln~ Okm'IoIII1br{) Oe.ltEl t1'l 2: Y~flrs hy aerial sprol:ring of \I~ry [ow
1l1l1L~(H;!imS ,of

poim. E~"'lf;1pol.mim.g froJilJ] ,:~e expedem.;e


issecncides .. !~ '!~'Meb I;S((IS€: are

95%, or 99%





i!lilthe UK . "There's

II~Dllmirlig ~;ullgi.c-.'I~, 3lliaul th~1

:Ievel:· he SJlys, "lnt~p1lstexperiooceprnvc5 iI. Tsetse Weil1€: wiped mit Of.;)Jtl I 1.S:OO"k:tll~ area ~n thewestern jP'WtlY",ince fL1nlbi'l m:;ill'! o odor-b ail:cd ta rget s; Bot$ ~·II'~.l1ia, l rid (11 F go

e.:U.Il"~lr1.lfly eusaive, ("nley OI]~y s have to leek m:wt w dirop dead," Edi~lbul:gh~ Maud i~11s!.lys..) And tmu~re.Iire If!!(IIny Ot~lC:I' rua.sons ",",'hiy S H crulliio~\\'Ofk~lnd ~)1,the \v,rollirg thilll,lg tOI try in Afi,ic3 .. cri~l,ics,say. Ap,~l;ro;o;;~Imne.!!yl OlJ:lmilll, ion kin! ere h'!lf~s~~d" lh~1\;1l.'We2:9' ~pl.'i!,;i~ and and subspecies, Of\li,fbk:h iii! least seven moe iumpal-

zibi! r's :~ 6001 sqlJ<l.roe"d~.ometers. mnfes~ed lOy just ene spec~es.i[would mle J S!Qn eenImiles<l:1i1.c $67 bi.~~.ilf.'.)l:1 do the ~~<lm;e ~~I,~f to in o A:fl"i~~. D',!lvid MO~~I~~UX of lh~ L ivt;.':rpoo!1
Schoo] of Trep iea I Medmcine sneered in <I

tam ftmn

.]0 ·~cQ!1()mk' rpl~b! ie b~~l~thaando

2iO[).~ !:,)OV1!:UHI~!ll~rry•. Whll'[:s. ~~nm. ~X~r.kl!lel;l ~hQ~\'s tl1.1~t'IS ill. resu It poi i ucaI ] 11I,~t'lhil:u~y. poor i:IlJr<IS!tm01J{!!I~re. an1d.bud gO'venla;unce".s;~lCh complex op erat ions .Ircll't :i!US!,,] inabl e ill A:rtL1,c~.g~ys M'a1,.lldJul:l,



;\l(}Jml,e::.H;~y. Ih~imfes]]Jil;l~rm:t:ii needed

~r~' [00 hi.!il.i:I g],vc~.[he lU1C{:I'[~inourceme, [AEA deesn't ftUf'IJ(I 51T plfojfj'C(!'';; however; iI provides H:ctllniC::['1~:tsS;ir:st.illl(:e. with cmlllt~,es

pickit!g up mo!\'[ on!'!e: l.lh."·


you i;!sk
9. fac'!oiL"Y

1E~,liIIli.o:pi'lto spend $12 ~nJ,I~,iQJl01]

if )1(1[1'f~ nm evemJ ~ur~ the lecifll1 iq U;f;l,\,m \lilQrk on mai nhnuW A fric~,'?" 'IS.k~ Ban Knol~. ~l ftlrm.el"rnAE.A i>wlr~r \~i11)[)i;now ~u 1;lrle ethe:rru.<ti:llds.. N "TOm!:l~, fh~fs ~n ~fhi~(:~1 '~lij·~stj.O!L·"(Th(l wt<ll CO:sl u.s mm,klffiQ'IIVIl but ",fiU bCllilllm;h hi gber r.1ro.iUlI, $,]:2 uilrMI ion ,. be:c!luse the j)m$GCl ~,s.expected '10 ulke dec~]dt:~.J
Wag~nil1:geu Zimh3Q)I'll\le~~, Vale s~ysl.hflt IAEA. [un.i.l:s ze;l] to' promot:emlWlde<l;1f ~echnology., . .h'l~ lO:Sl" :sight of m~ I.DJ1.itRC pmbl{;'lrls. "·NO!tS~Ml~~·'f!~lSi;.'l.l(':rs S5e!~ M~bmlll. 01!!'! A IElhiopi,J:lrn ecologist ~md Orrlfl of the fcl'llmlding


~,1uMrsof f~e c!t)~j mry:~SIT pn~jt:ll~l AI! A ~ ••~ v inrcecl the go".eTrllml:ent it~ And the expen~·e is 'vc'I~, \\~rlh

~~id.mI:l~eU~:~le comnry 0[11 "mythitng., he s-<ry!l: it ~. '!,~I~lS El~l~.opill.~ sel,eiHi:.::to;i;,.dlO g"JW S~T's ~
]Klte'mI,l ial<lilm.dcon tiO ~ ij,l if




C1:'I:mI b]"il.l,g

20 JULY 2007 VaL317

t']bo ~t[,1 pe:rFu:L~'<III,lelJit educt ~oIII i n ~ r M'ebr.u¢ d~p~'Orl3~" f'~cnh~( dl~ !oJ' f~.\i

hlilitfl.!il t~!illlt.IF.a(I(lri'e!lo\!rollJ1ldJ 1:llIe li'lIolrld pr'Cldu!Cc ibiUions of stli~lFlIl~ Medit.eorranealf'l!ruh mle~ iIl~'JY weillk tQi p'r'~toct ~h~9!~1l!~1 frrui~ lll'!d'lIJiS:t ~~


(i~i\il" farllfl1iHs :ste:uile male
onrion Hies.

:S.Ul ~!3t.ely, the s!neadl 01 gilllllil~l!llI rilV :;c,ur~e's sueh as oobiQIM·60 ;!lln.d '~e~l'um-UO to, poURicaUy '!IIoialtii!e ,wulIltril:!:) hills, 5ipilrlk~Irl'(;ontl€'rn. ifhal'5 (!f!,~ rea sen ~~I@' srr la tl' gil ~hJ~ 111I1~~r:l!ldti(mi! I A~tJ<rnk IElne'rgy' Age IlIq (:iAIEA), nreil ~ Vienlli3l, AJU(s[ria,iJ5 now ~i!p~rimeni~,if'l!il' l"rays asa way to '~t~rHi~~nqalEis, ~~itl1

r@mQllt~s;. A_s iillles<ult, tl1i!'Y (~1'I1 ,or!ily !lav@ mal~ ,onspring, ~\!hh:h ~nltum can 0111 ~~ pr.od.u I:e Illl!!! r~!'i•• md s® 0111. Mcnd:ells show 'bill ~i1i5, c:a!rlw~lle (H!ll a pOiPilil1l!.ioill jl!ist as qui(k~li ~s SUo ~~h@y S'ays. The ~eC:hm~logy, IIIm'lli III diewlo;pl!1l!imt. Qit i;!I (0.1'1'1 ~a.lly GlUed Oxit.e(~ in O.doitl. U.K., has .illlroflOy 1:1I~!rll!!l!ll'cl to (r@~~,~IRllOt MecUli!li:ls, Mexjl:i3Ir1 fruit IlilE!£, a mndiAM'es ae9'YptJII'I1I'DSq,1I]11l'Rs, 'wlllic:n traln~mit. the! ~erl'9 UE! "fbnJ!!l. I!litom(ilo~i:Sllfl~u,lR~i~,er.', ~l,iho's i:I)Ir~~rul~lestiUlg it hebe h,aivicu ,ail:ld ril~mlfi~ of A:lpi1ley'~Aede5 ~limsqIJHDe1'~ ill hi~ P.aUellJrF Im;lIi~ut!1' l.eIb in Pillis.• nWts R~:Dl ~,~V lP~f!liS<ing.'~ M[lny ijth~r Iln~~mQI'(;!g~st~ lil~'lI are us~n:g'g~l'Ie'lk trick!> to Imake ~1lI1l'cnSQIUlU(l,e:!i nable 'to [ra1l1:511J11H dis.!Ildls:e u l~.at (~'lJ1d .~repl:a~~J lla~Ulra~ pOIP~IQllio 111'" '~dem:,e, 0 Ma r(h, p, 1777;, 3 but Reil-er belfe<ves wip~nQ 'ClUI~popu1a:~i MS, ,1kS RiOt dOll's" 1$ rHOlr'e' Ililkely 1.0w~r:k. H:(lw~ver, Rml comes wi~h SOlme of tllle ~1;Ilme pmb~eIl1lJ~ bee mil'illl text) as cLa~5-kall SUo hrlij,e' IAEA i!1JS'Il'ct la~\ aJ ~~tvj'flg force beJhil1i~ ma~y I(lf the ~re,~k· '~~r~1Jghs, Ir:3id1a~io:rHl1oo t!:1chrliqu e5 would speU the end ,of i~5rais;o.n:

A n.eVl' me!hiod (~Uedl ~IReledl:5e I:I~ Insed'~ Carryinrgal [)Jorni,fFIdllll~ d'e~re: p~0I1ruILin:!lI i!"~,im!1'rltll <ioo¥leratim111n IlIUcl,ealr ~&hnology.,. Bruit ]m:g'E! l@tl1lar (RII DU fIll!lly prov ~de' 311l,liltl1ii!r t iOF!.• D@v@lotpedl ~y OX:~'Cird U mri" :s,CDhl' H@I'I~rich'S, \!in1o h-ea:ds the S~C:l!OIl!,i!: ifI(!It wOl11'i 1fI1~l're:!, Ile<>au:;e j:tll[I Ls.tHl ~as 1,0 !Jir®ve ]I.s. Il'II1eiUe, ~'ll,e prolPon.>e'Ili[~ ~he 5@ mI o~e(IJII.a!" !'!Ipprfiadu~~ Ilir lJIersf'!:y eliltollIiologist illJ~!l' Alphey !lind (olllellglU!''s, (),kuce" 311 Martih iilfide'f'esi!imat.i!! ~he step l!,,(ilm a ~1iI1l.1I,Hc:ate ~aJlbe'.xjiE!~im~i1i~, i!i1F!lopilira· I~ :;!Q 0 OJ, p, ~4? 41),~h~ ~edlniq~'e dQesJi!'~~1I::Ulia~lly~er'~ii~eve teas,ed !Il~~oe'$i s tion'aiprogillilm, ~ he wrote i~ iiIIl eflall toSdrl'DCC'" -MI.E. but instead eijlJlips the~ with: a '!ljilne- lhar[ is len~L whe-I'I ex,'Pr'~:l!>ed ill

voeal eppesirion. which he describesas 3 "eu It:' has made donors shy of f~~I,n(jMn;ag srr in Africa. The head of .IABA:" hisect Pest CQl1Ilml Secuen, Jorge He:ndrk':hs., decllncd to be' im:ervietn:d about tsetse t!lJl(:IIl1l'~.edS(,j{-"I.P.Ct:' ~oin~n)'~,d v.jri~~a:b(]l!l SIT's S,~IC'C~:;:-: lu Ul¢ fiL£ht agaiastthe c::odlling Moth. a pest () r pome rmi.t :JI,nrlwalmul tJ:ee.'i.Bnl he di!cli send

lh~u T~mz;rG1l,iand Upnd~1 C<l1!I 5lvcc'0edfcv[dl a S[T, because they arelarged lli,g areas that are Sl~UOl! nded by iafested ;I]'e~~~, <:.1ft': thu~ ad mech rnose prune' to re:~:lill\".ml~OJl,. PATTE!C head JO:h~l Kalblly<.l, a Ug;i1nda~l
bj)oc:hellll~:~sl~vlm speilin (j yem1l<lS <I. researcher ~n Ilh~ IAEA ~,;,;Ib.&"'1%., ··Pt10P Ie U~,I;,': dtibll~L;l lO th is :il:s!ul.eurrti I lll~.e: ~()WS e () me home:"I.:Ie
lrie:!>!O a'\110 id. ii, he Sio1YS, because .it's d iverting >!tteIH~:QIl H:'!Jmthe realwork.Tnseeticide ~prayillg and ~!lfgetS wt~] re:ua~Ja:ill PATIGC~" mil.~n[{lols.l(abayo says, and S]T U,S "a


a $i:l1lli~Ol!rCO:~[fO~r:;y ]5 sim-

S rf"s useful ness, .i u co.nm,bati~~g

!l!"m!~~Uiil.\Vilh [I-\£A S[~PpOili1. SUdZlr! has just eI!IJ.lbm:ked on <'L p:mj,et:~·o fight ~'~.eAliOfJ·he:h~" t


I~,'jtof e-ntailed

qlllesl.j,o:l1lCS. "The [AEA h:; PllShi,mI.g :P'n.mhillg•.



:~ of mii!ki~lgp:!'ffljl;:t;:ts

to demilm,d~liwo:m,~81.nembef i l-i]crn,drk]a:s ':~'I!"O[l~.,. =Thls is, a~, bac:k~Lp 'Opt iOIl," Edliopi~ii1I r(lj~clluf!der theOM'LETIE p oolll~rol. of thc:' Elhio:pi:J:U1:!!i:" Irs 3, "f,lIl:~ICY" to th~nk rhar ~~I!1\1~n~iol!1all~c:hn~(ll~~S~,aID'l 3hvays kill off <IIOOPul.alilo.un•. he '\'oI'lrON~~. n!CJ11 ~ IABA believesin a rele for :SIT'!ivlleif~ ~he;y cairn 'I. "It is :1i1lHmdly deJte~Jtilble." he ,uided to cl~i.11I:i til,1It Arri,ea!t'!!~ shln"ld~~am to Uve \vif~1 ~he' " .. p'['Oblem bec:;!:Usc' they <Ire ml~, c'<Ip<lhle'

mO$qui:l~ I~>o~~ I~'ilc NUI;l 'II';)!.Ue:y in i'ts Nortb.ern Sl,ne~ ccnsuuction .of;<l special mn$(lU~LO f~cllJr;yis pI3~!m.E1dfar Khar~fl:um. KlJiols, ,\,,'110 works, ::IS <I eonsu h.<lnt O:I~ rhe Ilroj,ect. S ;IYS ti'J,.H at ~A EA, here peated Iy q]U4l:::!tig:,llied ,\'b.et~,er S,mhln.~;Og ]!Q!.1r IQ buy I1lfli"ria d~gS'l1ia,d bed aets, ~llQuitd]'>iuyfor a :Fe~~i hi~lily !'>tw:ly. G, iven l~,e II[']c,kof q~Ii~[inl'ld sUliff. In:g~:sdca~ ~~ghim.!lI.n:;:5. a~d the :sua~fled [~!l~1():ilSW~U'l :h~'Sti(htn~s;egrfverfl~ R ment <I:S,I result of tile lOa;rf'mcrisis .. the OOU[l~ uy ··pirobt~llIyw"!IS~fl t~~k~1 pl~~e-" ~) $!lIl!dy 1:tI.C ,)ppro<Ldl eiljimcr., he adds, F'rm~: e ilew ohb P,u~I.;~~rln nne in R sri far;~s calls tile idea lid nu~kle m'lhnri<l.in A:i"k:'1 '\l!;'il}~ sn '·'cO~.nplt;Ekl ph:l i~,he !lky:' t [11 K.~li.t i.~;b:e debat,e tlloes ['lot seenrnllo


bo~:~~r t~fUfll llt~!~ng;~ng tilL;; ! t1w

:~ Pie' intih~ ;s:k:y'll


has, i1hiO eng~1,Ued Afr.ica's, ,P~H!I AI·d(~al1 Tserseflnd .~ TryplIl~o~ml~i!lS is, ,Erad! ~t'l._tiem ~31ll pa ~gn .~ (rATTilIC il i~ed lmo h lie llY Ahilc~n ~!I!t!d,·

~ The:: debate

;~. 1'1fgerprojectihe

muc:~,. TheY·lie I1mifl~)' e~lgerliCI gel QiI!I,~~.1h I:~,e'poojoat. Just recently. Lhey 11~<I,ve sUl'Uted rel~i.!I~ ins :sma U nnn'!L'beirSofs,leri ..e ~.lOl~esin l l~l.C' project an::a .. a d~y"s dr.!Jv,e fro]1m, Kl'I~,ili

ny ff~il::tflry t~~)

TliLe'y >!Ilretesting

\,\(helW,el the slefl,le 1n<!Lles

i er~ ~13


.Il:'1cctingif1l Togo .ill 200ft PATTEe M O(~~~ool i'1 t

emn slIrviw: in n~lllrl1! an~d ar~ stUll ~Ur.flcli\'\': N) wild fem~I~5-'I:t:{;;hhil;lf 'of":~llmieb is ~l~o:!J.-m~
I.eed aliileli"~O ggnecmliol1!!> of i;ab~i,te.,

]:: ~divm:;:~w:;. [T S

Wlil c\o~rin~.f1i~p<tfl of!se~se

:~\'\,':ide ban:.~e.~nd,e,ed, SIT

:1 p!"'~lg ra~s


l',h¢ first


.~re wry prorni$ in:;. g~S

co,~r di~~ 't~.d b~ PJ'\r':_EC h~ :!il T<ll1Ilz<l:mI.:I!<I----'w~~u;h h<ls the 11)1 ~a~wfy s,I:lIII~J :~ rl'omlhe Zal'll!:ib,1f ~urnrHI.i.gl1--;(1~ldi:~1 g Ugnndtl m~d ~~n K~l1iy.~. oh,i~ompl.]n too both bu_i.~do]];.e,B HI '~.'!I~n .~1~br,a~e.\~',hO fi r mlly y bel u:::ve:i> dle E:ti:1I OiJ11'1In pnuect, ,bs dou blS ~n

A~enl'Ll.wtl~olis, DOIwinced lb.,1 the I~foject-


Bb!!Ild b~~k.

To'I~!t'dl HetM.!fll~,(l'~. ,wwb~@Qd.



'h'lle by" toc,d ~l<ll:ll~htetrilums.e. is

$Jttrll~i~~.d.f'l100:~It~I1:d ~rlffiifll


'!,~h~chhe hopes wi U become ~. ~olirce ll.n·itwi<,,1 pride-'\r~r,ill~, e'!'l::I1i~;~iI<l~ty beo:uiuit "\:\fe .lre \~ry 'c(mnde!llt~l;U. we ca~ do h. otln,d \~ muM; sm:lCced:" .hoe5,rys. ":L~,lhrtopi,~ ~"llnOI; li~'e \lrith ~l~ fS(1l5e ~]y:'-MIArR1i~ N ENSIEHII NIl<





IA.VlNtilllJiSH U. ,CliiINIA-;WhC:1i! N~~~11 Wurit ~t wasa boy :iln th]~ vi I].~e~n ]nner MOall:!:o~ ia, "the gm5k<; grew ,I:!!: ~aim<IS, a'~l, ,uluil,"lft1e !M!lys. ln thi2 I 961 If,-, !toW:' gfo:l('J3ld y~~!r-mu~d never ~nd went hillllll,gry.#dl:er N<llSf'&:I W~~:liit(1 reached •. d~!tlh9'('Id h9'i!\1;'i,ier~thirol~g~f ~iv,~stm,;k ~~ud 1 o denuded (:tl.elaud•.forcing him and ether 111~n!defsH'l :sp~nd prl3do'lnJ> C<l'Slllii OM. a~~b'illa~

and the dle;.'lnUl~tiorluf more 'tlmn414[10~,ouse~"[1 d 2.,3- Iniii ii:o.lIlI, l~ec~:'.I'resof C:~O];lS, nccordli ng to !~~ Chi!ili~.~ Aead~i1'iY (ifFor~s.H'Y Sci,eiil(:l:;s. TiII,e ecenemic tell 1:11. China al.one :us. <Ippm~.i~ 11iiIill:e~;y 6:S0 mj llien <:I ye~!r; <iy:5 Wmlg TaQ •• ~ S ~ pbysicilligeogmpher w~.Q heads <I, narieual proj~c~: combat dCSe'rl:il:1cfIJlianl, to
loss: of ] 2{LOOO he:,;Id of Uvcstod,

ctny of'Seieuees (CAS) .ill L!lll1zllOu,esltlH~tes ~Im~northeru Chim~'s arid ~lrn~s,lolllds ~:lfe
1heing, deglL'3deci




Qf 36[10 sqwu:c Ik:il,o~ thU!1!tale US, ~1al~ ·of

Rhod,i:'lsl,a:~.dr-e\'ery year, \~fimg predicts

~h.Hit S, n result, dust storms, whie h ~~we ~ lnereased il;l nmVlibc1nearly sixto~,d over the
~~~ 20 y~t'I,rn.will be~'{lm,emore: frequent. 11liI;o;JJl-eil.n~n::.;;e., more de<1id~:y; ~lnd I f the 1~:;S(HilISf~r BlIyin,lmshu con be appHed. across the 1l51SlSlejppeS once :f'tded by

lieed. Hfmd

i'Dl, hand

\'/j,!i~Uathis c risis

\'>'<15 <'I ri511'1(g ~t

Ulings <Ire likely

to gel WllJ:se

be'Co.re tilltry'

rreq:~u"mC:Y~l1din~~nl~itydust s;lonns" "P,~nof ph: cl(lu~dn'~, gJCl ml~s;]cle,.;md we Iuu'[ to ~jghl

ca~dl(l.$in the m~,ddl~ of Ih~ f!f~i?rnOfl~."ii<:l,y~ Na"enWlIlritl.l. !lOX 5r1. .... 111 B~yin.lii,us,hu. t~1i,os.e' hard times have passed, O[U ;J~;neSPiF~I1Sd<i!y here. lush hiJ]ii
rcll t{I""IIl:rdi the :I~orli.z<:m t!:~d

bellCf"\Vilng, ,vhtl isbl~,,~da[Dl~(: CoM :lIml Ar~t1Regions EIlIVi:w.lUWClmtId. and EtIl~i,l1ee]f~ i~.g R(;~1;'[l(ch ]m:.tiM~ ofth~ Chin~s~ A'(,'<ld'-

G~1g_lmisKlm1ll!. d1151[~l'OmlS
~i'I.imd.deland practlees


But there are chal~;~;m£,~w imp]eli1lu!-:I1~mg r-us",
~:~Chl.in~E nOll1:hcr.u



des:p;ile ti, ste.icly \!.Iiid. A~telt ,iI 5~ye,<u'~!rIJort. the people of B<'Lyi;DiI,hmdm,~w'id~ llin.clp trm.n gJlfid~d5 ~ndf!.~,~,rn~ly o~~ec()i~'lgi~~ .., b!;)li~isls, and 'ecO:I1I,omliliits~,~.'1VC [t:'StooocilfuJ.e pastures. A]ld dus! storms here .b~!i\ie ~!00i1!C>d. Il<liyi~llmshu [~Si:l.rare hrighl Sl101 ,un a Meil:k 1~!md,.~~lrc" 11 the ~r!d gnlss~~1I11ds;of ~J)rth~!1I [ C~,~[IJ~, and Mongol ia, ovel~'graz:i.l~g. OVCIi'cult iV~li~~J,l,. ~1,.1 f!1~dcr~llg II)~~ ~~dI ~1Ut::~ water Iroe~Oll:roes have created .. massive d~LsJtOO\\r! wh~!i~ wind:> S'\l€'~r topStlil ~w~~)'.Du~t ~ll)ml:li, reyu~~~l'ly ligh~ e;~~~tem b Chri:mi1. KQr~. iII,llId J,rpa.!]. closing sclmols; d~m'lgin!:'det e.:l'IIgimes. Rl1ld tdgge1fi~g n~siPlil:<~tory ailments ill slM' ,i\WY illS ea~i,ff.ll1d }\ p~Ulictul~r.ly [l)l'Is,ty 11" Mm,1j,Hi1'1 N,lay :~ 99.3 R5.u~ted lin 85 de<lilh:i. tUu:


p.rcwioceli.. "I~ok)s:iC'.d Iy.

is e.l.S;t


«J~l~ro] ~

b: clear

~t~l"l •.Be:ollm:llic~Uyj.it is diilmFieub." ~J"~ s.~o~~s B;Jyinhi~l stmproj,ec'~ .~eadelf imlg (.i:'l(mliln,g 1 J .• ·ltmt cco~O"'i,Sl m:~Ji1leC' AS .~ml:Slitl.ue of ~O!<II1\)' rJ1'[h;:~ing.~501~.t~o:,~s be mik.lf't'lditp lh:~ m~~st needs of]oc.d res,id~'nJts,U:ld ,ecQIQgk:~1 cendlh:kHliS in ,ea~h r.~i{)!~:Comp~i!c'Hing the picaire, W'I,"'·. C~~:ullese effieials still llrn.ew du:o~ tiO t· crcdiu::t!lpoHdC$lhat <lin~ to subdue du!>.l ~MlI.'unsby conq[~erinag llite desen ...... , "We have ~~lot ·of C!(}.Iwj.l~.ci.!n,gl.o dfl,," j:ila~lgsays"


.. Iiii ~ ~



'i DUE! ,!!'rf!@d: dl!Uis~ :stOlflnrl ~. The ba!i:!e 'H~~tHinlY[if Ea~1Ju.kfs d~~I~!itor~~~s ts ~~!ii"ly \:\IeU ~~lab:U~he:d. F,ol!'stai!l:tSr~. the ~. common ~em;!"sand starms' :is <'I mi!mm.ner. ~. =. Taking roa~:. J~ang wrlming r,t)M.s.l:Iold Sl)ii~Ilnlp~am. show§

dens.e graS's

SMiiI.dpm"tid,el'l<irei~) henv), h) g~~ lin~d h~gh i:rrnt"I~:~.e OillI1lUISp~;u;'!re. Thus, lirlde' of the ,(limn

... '" s

20 JU LV 2007

va 1:.317 sc~ i!N:CI!!

wWV!.l'.scrcmc~m 0119, org

~A '!J! ~'~i~ln!,~,¥(l hl,U,g!fJ '. Ii!~S~J ilCt i1'l9 g ri;l:i:.1j,11 !II ~ (it:l$e,!) allowed this pastlne in IIMer Ntml'9oi[ia to r'l!'!~r n!:it"lr~U:v.


ev~r'"\\,~id~ningrial!; of

d,~~1jded ~,f!fld {rf(~t1.i'l!d

th~11bUglu~ E'I$~ Af.ioJlcomes rrn~nl deserts,
where eresien

mrer~:~],elllil~'emlu:1hes Clllil,r~:.ii:':d n

aw,lY Wl:0!;t,of ~he !'lluli,ler p!Unidif;l~. S~~A,d~e~ fuel. Irrig'I~:ilon .~r<1,dll~lly dllried 'Up~Ull1ly ifldic.Me that tile dust origilill.neg in d:ry lak~5il~d Ii~v~r~. he !,,~S1i.dt. Ji~ng !l.~ly:.';. ~~ T

sertlements, The !o'l~nIIme~t Ui.SlD e~couraged H Si~ Chi ncse farmers N) 111igrn te to n{lni~.elrn :llegioi:1i' 1.0 "tame the; de~~~t~" whh 3!1'li~ic:13.1 eases m~dirrigmiml. The' ~1Ug!.'fUU:S de'lt[:'ed land for f~lr:~],:s and cut brru~l~ for

reglons, ~l~ s,ays. "(klC~s not help combOlll d\;:st:[~if~curim'l:~The- gqJverrHl.'ilerli[Cmn inues
1:0 ~)our money into ,an~o.r,estatio1m,.regardless


resources, Ul rOlllgh .~ blll re<l:l~cfacy
iS!liplant !'I'Ccs."lt.1
50 time

\\41fl!;.C!ll~:is,!;i en


I~e Ch i1140:S!egoV'e[II1:1i11en[ .". tO$iC~lem:ifh:;".lliy e\i<J,I:I&~~le ~lol'I~~tterm l:lol ieies," Luo says. Slllst~ii"'lalbl.e ~'iW[!1Iig ffin 2000.,. CAS applied

13~~b~d!S ~:!m.d!IridlaUlidls l~!l t~t;:ii':t ffi~g~~. ImI llil.<lt901% olrCl1.illa's g.ra'ssl<lUiilds,. <I.Il area d these re,sim,ls, a I:TW>~fonns on lUndis:~:urbecl ~~lrxl~lpm;$,i rig 4 mi~l:i!Jj;] ,..;q uare kilvI1!l~t,~~.• ;"(]ii~. givi~g ~lml;ln:$iS<i:u!!!Ci;; ~~IWii~d ~ru~~m~. aJli\': egraded. d Bm iu~,~prillglilll1u:. that, cml!~ is brok.en IIp by Amll~oridli:s l]i~Ve long rc~ogllized ll,H~ p.IO\~dng and livestock, which


scic:li),ln,fic tlp:proach


problem" Lmt <luemp'ted

f~xes have been

h.l[ldofnl~~'1i gru'l.'o'l[irumld pound sui I inro ,

Meflnwhik=. the tlemper~l1ure difference between <I, chi 11~i".atnillOS p.1iae':lre"lnUi "" su:rlih:c:e

\~filned by i!1~~lls~fyil1g:!!:pring:!>l'~~.lig,M c!'e~ ~h;;~updmnf' 1~i!l!1 dl~s~~mo rhe ~i!r. As ~,ir lift S>U''C:'3.I.HS seuthand cast frem Siberia, the'
\",ri~lds ])UD1~PWlP'lSl'li:nsllm.e

rUli ..e U·:rm.O[ cmmlerrn1dUClh'~" Sil'llc~ I '97a, l C~ilKl has!Spel~,t :al least $1 billion planting tress in arid :;;lInd~()miaJidreglons ~t'I 'CO!l1b~I de:sertif]c[I[iil;.)n., S:liWS LUQ Y~Q1i an ecologist . lit. l:~'neUI[rn.ive.rs.ily ofnkJa~lmni!. iu Nor 1l1iIJ2L:~.• who W~[!l1loUe~g,l]e~'l'lt~.tl Co.ld .nm,d Arid t

by f1;Jtnding fiv~gr';;l;;shJ~d-, re!l~Or,ado,lll pi IolPlrCljl.::clS. :~llICh.uii ng J im,lg·s. Jia!,lg headed fO:1i Z~l!!::',lgl!lllil eO~U~I( y., 21 suhlti,visiull ofXi~.ili1.golLeague, 'p.fU1~ybee ..use th,e
~:IlMfHU£ (liWBot..lJil'D.y a resesreh 5t!ll.iQ:I1i.there has

dw,~ sronas

1h,1[ had doc ~ll'1i1.e.n~ed lass of [2 C'e.11timerers the ofi:o'psoil to. winderof>ioil in th~ :p$t 24 >~:rs. J\m],th~f .re'i~ro!l:Begjim!~is(Iflly ~.s0ik i~oml;~rs

II;m(~]III deg:Jj'JKl~ed here'.I.lime d List

:mi.~(l~I~.lrain rangesthat ring .I:i,o:rtll~en~, C~.hul and M o~go lia. fb':rg'ji 11 g i

w in;l~rr:eet Be~ji!,g." Ji;;'i\!.!; Soays.
Rf'<II~,iz~ n_gt:~lal. lilekey [0 wllvd:1:JJ'.'l1: pmble:~~. is i~lvohril'i,g



dUSI lnro the upper ;,tt~ffi~,os.p~,1ere. aste rly wi nds B


the people ,..1m live 001] du::, 1:"IiI~r-:<] big task ,giv~!l. .i~.~g fi;;1)':>. "'dl~h~ J
pov'eny 31[:ui,thei::r level ofcd:m::illid(lu"___:lineinvhed onro hif>~~a.~~ social sciemists <lITIlclCC'Ollmni SIS as
'IIIt'C.]~ ~li':S ecologists ~mla'~iom~1 husI:mnmy .~pecifllUst1t goo.! "V.rdsm the

sweep the panjculawm<lner


B,eij.~I1ig. Seml11'0i'kyo, and samedille!> !lcross; the P~eilic Ocean to Nowtl~A merica, There ~rc good years a:~ldbad years, HeOlvy !-lnQow~add moisture to the seil, d!lln:pe'lll:~llg dust

irni.WIf'O'w thL': ~hlCs ol·".i~lil.ge~,"S,,vliti le

reduc[in~ e'llvirmrn'l:m;:'~llll de;r<oci~-

in eafIY~1Priii!lg. Cri!lv~ii'1>eiy. wi1thout snow cover, s.oU dries out d~jilin,g 'wilil~~~ j$ mere prone <ltld
to wind erosion.

(i.o'I'D.A~the st~m 'flr ~h~ :5-)1~~r. S6IHI.!uOUproJ~el. e1Jyiiilhu~hl! censisred of 72~,(iluseho.lds. with J,:~i'l people 'I~d:~ tS60 head of ~,iV1eS~Clck-75% sheep <lnd g{lalS,.
1M rest C<Jit'lle'. he vi~ hl~":e :m';'!.Il<Igl;.~ T 733.0 hectares ofbum.d :mlMlcnl ofil:
C()m!IilMIIl~~ I'11."!\lrB"

This dyu~:m:k;hasperslstsd :~b:r

tim storms


ha'ff: dU~1sl)OrmtS, 111111.

have bee:mI,'i"'fllrs~n~i]'!lg"UO~
bears the b.mnl of

m~:!k~! Eh~lflr!1!J gels; bl'!tt~red by d:~$'I.i!"l this 28: A.1;u'll~QIQ;5, ]!iI!~9~


(a?liL:II~elil IMA;SJt's by l~rlra sa~e\l~it12.

.~ ,i
.~ .~ .~







'II! iii a

.. ..


Ea.~~: A:;::~!.I~ dlH;'~ ~lllnn'S. suffered '-dust eveats" ,on ,23 days dl~fil'l:g. the l ~no~, Reg:~onil Jns~jIWU; has stud led sueh 4~.day.s~n~J~c~980,s. 70 d~yg,~m]ln.e ~,990il>. and irlfor·~Sl<1i1ion elTtJiIl'H;'. t.lt), d<iYs $:0 far dlais deet.de .• u::eonlins 'tol,he A ffo!:C:.~~~1itl'[I'[j, nli~gui.dl;'>dl, ue !l s~ rI'~" ill L !<,orea M~lrnrQlogiC!l.l J\dmllilli>t~~lionl. "iPe{)ple proposed the ide,.. withoHt considerThe pl.·umlt,!)f reason for this enslauglu, ~Jl,gi:':clQlogic<l1 principles," :ful.e ays. '·"[hey set s [most m::ienl:is~sbdic'\l'e ...is d:egr.tdladQIl Qffra~ tmllo 'J~te Miore.~ru in region~\\~l~fo~;~I'::l i:~1;;I.;iC (l~y:))t~ln:;;. Thi,'; 1iI'(!!PU! ut ion ·~~·f I XiiwJ1IgQ I 113UII.':'IUy do l:I:otgmw due tOo llirnired, p:Jfocip.iLeagJll~ the disl~:Cil ll1m i[llcl~L(~f:sl3i1yalJJl:m,slm, tmu;m~:"The tfoo {If chol:ce h..~ been~~e poplinr. i!1!c!reas-edI[rum ~bom 2,(]ro.ono il.1 [helme if wme:l'le{i\porlillarn gmw ",~pkUy. !but \l;,'~lh(ml ~,940s to [1'D,lOrelh:'lll 950.000 1.1111000"JiU:'Ing [11t~~lsiv~ oCar~. th~y di~. S~kb; pr~uw(,U~,g ~ys" Over 111m p.;;.riudl. herds of grai':il'Dg tmi~ f:wnffi,b<lne~le'i!lrl:~.~de<:ld popl.<!r s:.."Iplings[[11<1Is ",kym!i::.keled frO:IiI.1 <lI'OLlnld ~. nrruiiJliiiCIl1l he~.d [in~ ]7Oadsin Innerr MU!l,gOUll.\;l';h~re rO'p~,f!iI' to n~or.~ thtU~24 n~im(m, '~l'i.lhi]~thlSgr~:d!l~ grove!> become es.ulbl[i;si:led. Llio !)!.iys.,llle areEi sl:mnd, fr-om ~ hectares pcr.mi,~mll. to d~~ply r!:Jo'led tii'ees h~n~'Of!ihagew;wJr <"lb(lUi ~!l~wmh Oif!l :h4;'C'~il.re. l~.r{;iilugh Hill'i SP~If~twon ,kl'l,'I-eri I.lg fhe\'1,.M.61t St;nggeri Ilg ,grOt",. d1Jsm; hr.s Ihi 5GCC~H,:['~d tnble fUlld 1l1~~ki~g,uth~rdc:r for. !mi,ti.h~g:~IS&l;':"" ilI.1I aa.oos...<;.llo:rI:~'n,e'If!1, China. Til~.e I'Inti.omai gm'~ .mds,hruibslo :mrvi~,'e, e~,"nmem encQiu!:i1.ged Ilomudi,c berders to ('hin",-S l~~-p~a]!).Iing ca:~lp(aigl1i toltCb;)1l: sen Ie ~,Il v~ Ila.g~,~ Inl,lll iply ~liae;:1idls iboOSI and 1:0 ce::ssfll Uy rcftn~;st;ed arc<I!Il VIi illl <Imph: .r<li::[~, ino(lIL11,eS',,:ii'<lY~ Ji.Ull;l. L~i\"e.stm;k cre,u'ed mill SilLyS Llio. B~tl p~j3r'1l11l~poplars illi add

Jianlf:s team c;I.lt;l'LI~'lted dl'J.t vim~fug~!Nloluld boof-t in~om~~ ~nhcy C reduced sheep
UI!ild~(I.~l numbers

<lind intro-

dllced an 1~11p3:ovt::db3,rod Qif"dairy cattle, ~,'J,I'hHe cllrr."j,1 ili1S'1pi!11! gll.<Ii';jglg.lhV".1S no[ elll!')'I){) '-10r1'Vil'Dcil':!~cln. hm.vcvf':I:: MQ!l1.,gol.~ t~O~ls.[dcr t~~

size of ~he'her.a


oOf we,)~th. To he;llp

doubts. local (Il1lhor,itie.s chipped

",ddi ~iOfHd ij~cemi v,~~; Tb~y d~!~ \1!.,,~n~ ;)!nd
exte.nded tI~ pmvcr gr,ud to Bia)'i:~lliuw[s.h~[, 0 nm 1 pumps mmd e~,~c!rify lrtowse:$. Tbi,;; COIJIilty !ll::;t) impm,red lhe din nm::k l..l.1nl1ccdng the \fi:~~ilgc



"[:h: vi.H3.gers<lg:[reed

t(l bOll) g~t"<'lzill.g Q1n





r~!lg'~hmd to

liI]lklW vellctatllolm lOreC{loVe'r. H;".rvC!l;I:in~i1J!<Ii)' ~roMthiJ1: ]~:l'ndin ~Unlmrll~~wi,dL;;:d~~oug:l~. (l:~r ~e ft)i il slHn'~~:er mlmber~f ~hie~'tt~ckmii~g ~I d ~:ypi.cal wi Illcl!', e Ii III imMI.l1g i:~,l!ee"'I;pe IJise of COl'lll1l,~e~,~i.,.1 t:Q~ide the yillilge:rs (Jver feed. ,1!I1.l'iil.e :~anCireco,'~ro'L th.e j'i~IlJil\;I:eaml plafiil.od C{)i,IT:UU, an !:iiev~[:.I'1 dO~C:~,l :I)te~~.,re-s.



lLe.ss'is Ini,O·re., Naselii WUlrn'tu

SilW hi s l~cliI,rrre inclre:.a~e< .atfllE!iI'






\Wm!( H)

m~dl!' so!n{;l :Jro~hqi!J;lt$l1[Hlg <

the w<'ly~ j\/,iOI'e than Im]foflheunilI11] budge! aerial grass seediglg and p'~u!l'ltin!A
willldbr'e<llks. BOI:hl PI10Ved "01,

p~$Wrio;lS. Stiil n~~f!yi[fllnl!$h~ experlencc B lH<!Iiy not be easy to replieare in plaees 'Wilh
less f[p,ltl:lifibl(; '~l~gll.),g~,c~!1 ,~tH'ndHlou S. Jill:~l,g notesjhat ~ ... i nhushu had suffiy

I;l'rdl..{rf.ld ~0,5000 people

~o \l!l~~[o(;IIii!rmsin

rrees to form

Ofmgjley." Ji~u!lg Si}YlS. The trees dh:sd, <ll:tt~so~,rn 1)1101::. :f,u;-edno Delh::r than these le~:I:

e lent top~o


r~p II;;tewith



to recover g]~.t:oroUy.

srQ~H1d1W<1I:e~ levels had WI(U beenaffected

by exee

11:5 ive


worked, The vm~g.efS geew ,e~l(a~gt~, to com
fei:d: <lni,mtli~s, wi:I~.{)ln gFdz~ng in~:1'aecOJ,mnO:i11l
IXl5!lJUe, HC'rols~vcl'e ,oodlwed LO :5 783

By and huge. huwevel:, the sn::lm,l:plephUI,



~]tlleover 1101 Ifofwh~dl\M\;lr~~h!;,~p'-'[1!J
M.~lk pmduc~i,on doulh,k:dper~,C1!'lll(t

g_{XlJI'!t Byl]~C'

degradaricn Ilm,U\Y require "human ff~dl~i[OiH~Q~ of the resrerarleu process .. says Lu Qi. a deserlir.cat.um,l spe" 'CiilJilil ill ~J1~e ]n~t;unue ofForc5,try :UIlBeijing, After .<;lludyiH~ restonll~(l\lm. pmjecl,-; on the
Tibetan Plareaa.where

a I OOO~krnl.l;il area :s.m'luolllndin.g Min'quil wid1ii'l ;3 Ji f2 yt'arl>. WAng says l~I'U. resertl ingthe farrners d,:o;ewhere "mlly relieve ~Oi!iM~P~'(lbl~ms in !:h~1 .1I:i'!:<1 but Coni se aew prolblern1lu;ull:l s another ~r~I),~"'~'I would be'be'u~r~ he ,1I1lIDIC.s. 1:0 intruduee warer-couserverlon 1,cch!liqtu.;.-s., such as [ho&~rio~H~~n;;d srael, in] whidiil m:~ghl allow smsl<lliilll.<lMie tlumdllm~ in
I.~'[e· area,


At Bayi~fuu~11U .. ri<llng, cent iuues to measure tile Cl'i:.perw Ine!a['s results and

end o:hhe I,hif'dl smfll11er. du~';rm~j:,>~iglilid fecO~f~ eredto pn:w:rde :m,mre dl<l'IIII, emmgh IHiY fel" the \!:i~~,!\g~needs, \l Fi ve yeiill:slat:er. Ji 'lI~~says, the ~l;lIldIQeks

as il prnhubly

did ~

Ci.1!:lU!1iY ~.gtll.

A:D11mJaJi illcomesinaV<e iucre<lJleCl46%. [tom $315 U'l $460 per C:<l.pit~l. 1:11N i!cSelli'liVIJ rh~t 's, ~i\iin£ room, a fr".ul~;ed ceramic 1fe;I~er of
Kinl!l~]1 Ilflll~smm.

the: \\/aU ..A.~arge"

screen TV mua ...satellne dish in the from yard pir~i~ pnw~ou~l)f nim~gilll~d (lr!:~C;l:r.u t<l:I!ilm~m .• '·'WE;!ilsi.;ld r,o jll'''al~h<'"~tthere 'V""l'
II]Qthi]~gto:1' Mo.umg,olls to do m I:Iliglu Inn slee,p .~nUinmke b~lbi('!l,:' N~:senIWl~rihl ~1I)1~. And the dm,! slCi!li'~m!s:.whi.chLII.~cl to drive people

created shifting sl.Ind.dUIJa,C!K. Lu fomrnclthal <lh;mds·~off <Ipprm'lcl:Ii led to() ,I [olIO'!l\' ~Ii!.d !j.pI}Ul'ru:v~g~t:u~on ~md JiulG ~Hlb~li;mti!oII,1 oftlile dt~:Ill.es,.In contrasr, e:rec~iDllg :).~nd barriers and p~~n:iilmJg soi.I.-~l;;l,bi~i;::~ng shrubs pr<O:m1ffi.oted lhle hcaldlliY rc>covcl:Y of native plants .. Becalls>(: shjifi:~llg dunes s;rnolher new W;ls)etal:ion before it can t::Jk'il: nun, Lu arguesthar aerive int~!iV~nl.iun~!5 needed to reverse desertitlcaticn, l'h~ (o1!1!gh(':!;l [;TIsl.: may b L;'i to w'Ddoth~ h<'lr~~l reught by artl 1:1ch!l ~y expanding w eases, like OI~.e31 M~nqhl.l}e1,wee~, the


vi nOIse lncumes, The' ~UOJ~~l is I{~~vi~!i! f~m!, unexp~eHld .~~gaeY'.B~rtlre t:~~ lP'n'ij~e~'b~,g~Jm. Nns.en 'W'lIIwitu says •.v~illllge youDllgslersl)'p~ iC<ilil)t dr{lpp~'d Q~lt Of~'CIDallglld:~~l!tbe

explere w'l.y!' ,~ofurtherraise

TenS,~er Desert aad rhe Badaln JI.UM
'ne sen]

indoors once





m;>C;lsl~OIilICl~uisances. n

BayinlH.lshu :i;.;,.. "goad ~:>;3m.rl!(;" ',~f grassland resroratlcn. W.mg says. mn (1'1111 ~~~c(l~n\gin~ S~!~l" herders i~~,eu.rby vilfu~H!'l!gth~ e:!l.pfms:vO!1 of the 1~~'(1 deserts, 1~lges <Ire restri Citi.~l.g gr<'llz.i ~mg 00 rnlnn.maI Ead iif! r l'.i s yfl<l r.pi revi Il Ci,31 ~lUlh oriti es on 200JULY 2007 VaL317

iGa:~ISI,J1Prov:inee, \VCSI: 0 r ]nner M.OWi1,goh<Il. Begilll.ll iill; i Ilt:heI9.5 lls, wni g.ilio]iIJ on a massive seale helped: establish thou sand ~ ~l:f,Ib,r:un$ hUlt: ~'ie:!mhl!l!!l.iy dr~e:d up natural rivers <'I11I,d depleted grommd'!M!lf:er,.

p~IIS:Qory 9 ye.n.s. ~ut the scieatjsts whe SP~D!lI: :rin1le i:l!1I, the vlllage expased rhe )'Q~L[b£s.~eU"~~ to the ~nl:el]le[ and ~e:XI:IliiI,eSs.ilging" '''MiI.ny youtlrn,g p(':op~i;' real ized the impeerance of aueducarien," he say.s. [xhibH)\ is Nlls~!l\lVmiHiI.'seldest ~o~. now' study iII!!;:, to be' <I veteri na rU'OIn<l!1 rn nner Mongol i II. U n[versi tyiu~. Hol1.ho l."I.Ie ~:V1lllns, ((l,S!t,,.y in 11rnech)'~o ,~rsl!e u beuer ll fe," N!I!m n '\1i;h.l.ri1ui>~,Y:ll,'fThm.t.wouid mean oliil: lessperscn ,ekillg out iJ I]villg alii! the gll<lss,~andls'-<llldl a greaser chance I:) If'ellmjoyi:~g ~~.~ ~!wj,ro!lm~")~: i !:!~f~"s~"!i!!gly] i berated ~l'om dlHlt,


~~ ..•






~~.;:qu~n.d!'ng or~m~nyf!~lh'l~lal sr~cI~s to help il'lllef11l'lre~~~m.e hmmnrn gel~mnf'. "'O~Vilrd ~I~l OIIUI. ceraralized d~n3~ses, ,sm:b iii; IEnsembl, dC:"ld~


As.tilnlE!' IrI\Iationa~ IPla~tIGel~(llIl1!e! InUia.tiive'lUrns to!. it is beefing ,a.nd its: IPo,rtifoUo, €!If S'i!!,q1u:enc@d (Irop allild IIiI~O~IiII,(jrop g'enOIill'!l!'S





'tiP its bi~hlformatics

eenservedgcnes and p;aJth\';t~. That. h~snl'l

i,"I~:lJ:lCn&1 in dlepbm werld, 1\'':10 .~ ~re!\1l1l. "dam ~O'lIIrCi..-;,"a~' bfu~km~,izt,'d:' 'OO~t~pI3Unl'>, in:(:u]n l Sl1.ei~'.~ hjoin:rUiJifi'1;.i!tici~t i.lt'Cole! Sprin,g H~!rb.or
l.<IOO.f3lo.ry ilm New Ymt: st:1:I:e,. For' ;IJrilitlop,;;.fs Si."\[~~~l(,:e infQm<lti"~lt.re~tlfCl'fi:~ g(l~O~d~t~i-'

1.1] the genol1li.cS\'~i'Ql:ld\rl~nlls <Ireseeoad-elass citizens. R~~~~.rC'l~~,ft; :h~VI;l:i!~(I~ll;ll!cetl b!: ~ DNA luu:tdtrnds olF l.ni.cro1.x:s 3Jl!.d dozens of

rO.f :mI.lore CI::OP genemes

, E:r.ic W;J,rdi, of

anirrmlli" yet dIe), h""'eL~~ipheredi11lu~ ge<L1!uik~1~
three pl.t1:nts. J-.lm,blch:JfJ6:is. r,ice.;md rapia.v-four" itF)!iflU Cl;'mnlChl~j,,,.~t\(lrtmmlm~ .. an C'o:mp.;uisons betweeu :ri.l]ned.~e~ed


~~.eT"\'Q IBlmJ~ pQl;jfl!iida(l~lnil1 DIiII:rlmm. North
CI:ro~ ina. wmn.ob
SUppOIi!S the deve'loIPqmu':nt .of di::iiei:1.s~·-re:-;i:ii~,mlerops, cited lh.e- IL1,~cd for specieslbal represe:~1.l ~he plant £l'OlipS. all Ol'h~ arg~H~d ··~ql~IH~i[lg" species lrnm lor dliJferent phtces'l\r,lm$tl! genmnes <lll.oe .~Ire·ady known-say. Al'c{/Jidop.'f{l'''''lO g(;l,t a se!l$e of ~:~.e I!l<llUl:~II! 1 varialn.on,

and r(~a~h~f~d speclss have yie~doo lrem~· d eusi ns i,gttlJl;s iutut he evohu ion of the se a:I:g.>ulisars. Yet l",hulA biAJ.!(lg,us,~:!> Solin .19!Ckthe
abi.~lity[o (;~Ht\lpa"tl the gel:lioltl~~ ohhe]'j l:al!,.tlrile g.iP'0c.i~s. k,:[ ~I(lll.eb>egd I~ to oon~ll!1.tC~ a .~ ceherern h i,si.Qry of Iilaats, NQ wonder 1i phll'll researchers !lr~ ccm:~iPb.ining. :~ A~ a 6 JI,I!.ly'!!lllwk....I'Klp ·to. ~'If;;~~~IOO dw a I,O"ycllrc-okliNa~iOn;[I1PlilIlll Geacare .I~~.i"


\~tork!>hnl)pi!l.~.i.cipil:mll!> aISQdeeried the

sen.om!!; ini tg"Hi~re·~ lack of progress irI bieinfermaties, F~~mling~g~lmdes ~ulP;po!l'N'l.lllhe

base ealled TAIR, b1lll.~for corn, [l~,eyhead to M<!J~<iDill "TYml]: can't go and ~e.1 compr;:h~l~si,,'e cClmp<uisorl OOt,veCI] II mbi(;lopsi.o; .~nd n:cc." 110l!es Ward. "U.l~ f~~Jnral:i~~g:~ S~eilllill1d o~:Jiters C<IUed rior theiutegr.nion (liF ~h~\~Jlij)u~pl~u ~l(Jm~ d~naha~s;<l~d IJ1.~ , f~)r ~tJI1Lis,IJlnemoflinifo;nn Sl,m;cl~l1icl.s,liOr,[;h~:<Ic~ tl!1:i::dng ,gen.cs ami olh~r DNJ\.. "1 fyou dQffi~"~ do III is. )"tmr 't;O['l.lpiIDl ~OIl!sbet\,,-e.en ge-n'OItI,e~ are WI.~.teD'i]'y 1ID.e!ln'unglcSc"," .S:;IYS: Suzanna. Lcwb: .• 21 bi!Di~lformilti.cisl :<]1 Lawrence B~tke]e'Y N.alioll~1I L1ibnCThlnry i.nil C~Hfl;lmia,


NS,F.DOE. ;l~d its co~l.ahQf"ti~~
GI!lm@!me ~iZ!e
i1lge:mlcies are
t;1 k ing :S~1l:ps

~o adcl ress

~:nmivc (NPGI} .. expe['i1,.)i ~nbioinfum!ilmics,pl<'ll1illY~~e(milrag: and hiotcclmol.ogy .il ca lled f,~llna J\~ p l'lriltgell!Olnes. to be r .~ !'>e[~~~eillct1dlin:! bllleJll:ed tire de~lrlh of <


these {;-omp,~,a~~r:;,. llil



aW:'Irded W<]s,hi.~ilg~'onl Llniverslty in St, LOUli s, Mhii.{)Wlrl. S2<J.5 lU illion 11(1 :s·e:{ILDe:L~,ceCiOlIl1. flOt.:l'i(l.~Dm<l~o. and

;; compul~LkHl~d~nd ~!l~~.Ylic:~1 toC!~s tQ 8 1)j;,1" .p!",J~~dhe prQ<Jmm It)1" !t.; prog~~" t to dlarte. O~lf:rIhe P<lst docalJl:e. NPG [ has :~ !>;pe~mt $7.00 :UiI1.ill.iml ~illdif.l'g g~nes and
~<l,].ua~:e.~e!llomes.,,'{ct. <l.lt.~u,~ . .Sill,

soybean sequencing ~salso under
way. DOE's JOillfDtGe:m.a[f.~.e.~ns;~iulle in W.!)~~Ht Cr~el\;. C~lif()rniil. :p]~!'lS tfl devote rnl1creni ngly more otf its
c<lplI,ci.ty lOP~~1'lmtS mJid ling its work 'with ani mal s, say:; JCi l's Da f.lli.cl R()·k~tS~r• .A.~]il(lM, abuut" t1,,,'O dozen ~peci,c!'i .n-e sequeneing




DNA. Th,u sa drop ;~in~:lia,~ ·?"'1.C:kCI.,.cOm]¥lr,ed to mm'e.,II:mn 'S $3 b:llhon,n,~<Ii~I<lbl.efro;~u the N,m,on<'l~ .~ Fll;nmi!1l 'Uel1()I'nl;C R~se~,n:::blhlS'lUnu!Clhr :fi! d~".C~)d~lllgfl~e i;~nrnnes of lu:n'l1!l~LrI:;:, f~!1d ,~ '-;: other mnim~.Is.. notes JefflOang~, of t~l.e ~ Ul!1ihl;nj~}" ~lfN(lnh Carol.inf!. C~<!I!pd :~ Hi! I. "P~a.r:LI.genlornlics [;esearclin~is a bug.e ,Ill bww,' f{);nl~e buek,' ;:,n!;UI~ V'IIl!!:!. ,"~m .~ ~ ~ ~ :fi! c;h'i~rstbeN'lli{Jllal R.e~e<l:r'Ch CO!unci.~ :~ p:lu~1dm~,0e~wi~~ro0Vi~'\v.~mig and NPnl 'f( 1~1OJnmelldi"IIS IlI,tuC'e'dir«[iom;:,
~ ::;eq\qtH"lCIDilgp~,.Ilbl


in the se'q,~lencill~ hoppf:lr. Rice·'
NSf ~spu~~!ifl,gfrw better bioinfo!j~ Im1.<U aswell, ]t is ["C'!.fiev.riug pr-apo&'1I.ls ies


for .1"pi~Ull' (}'l~rinf!'".!l$tl1!t:H~~:· w11ich
wiHbJi:li\.~!:lin.e t'OlnputeJs iI'~d kllO\:\I"!11m~¥ to me::M t[l'~'v"ti~JlJI$ sequence, ;,e!lll-' explie-s~iIOmi. fWLa:tClicnal g.e'l1I,omi,t:'s.'J~l d :~~.u~~1Hal:aba~:-; to mak!ept)1.;~fl}iL'; ened :SlDps.tmppiirls, ItCi" s,emmnics. NSF plill;lS t~!!:~nd ~IP to 0 miUi(lll~ a ~~r ~br



C(l,~gret>~ kicked off



.~ ~~lilrtki:ld·'40 mi IIioufbr tire Nat i:mlOl.1c iS

~g\mt'Y[)~'~In g f] ~99~, \~ith p~n.~:~ from u.s. c:or,~.gl'O\~ ers, It ear-

:; ye,lrs-IO




; cnreFmnuiali.(Ml fNSF} to uslmcr p.lallls, !il' ~n jJ1I!IIlicu]<lw l'Um .md olher crn;p[S.. ~miCI .!;! th~ ge:.no.mics er~, Nn\\! I n years ~~ter~ ~ .ilS· N SF. wi tll ;)Iddh iO~ fI ~ S ~pp on from :~ the US. dcpill1me.~,I:S ofAgri,c~l.b.l:'e '!l1! <l!!'I!dEflJJ;Jrgy (DOE) and nth~r reder ..~
.~ iIi_gen,cie.s.,has> S;1~ciOI.1SO:rod l'Iu,,~dn::ds of

these gt'!iIl.amic reseusees i:II""oos~ible ~l1d to lr'''l~nreseu rchees ~.1.0\'" W WI$e '[bent Trn.i~Tingis ktty., :Silly::> [)~]'I,g1. I?G I has N "'bn:mgbi! tliLe mi:nui [rarne I]fgemJlmics to plmn sy:M:elm \,,111l:remhe~,'e\'\\'"d:SIilI't mllOC'h

:5 genmn~..;rel~n~pmr,"-,e~.

;~ 1m1Ooc;DNA. f\~ .n~,~;eling, the


B Jut reseaK he~1)re

d ",:modllg for

Erik l~g

:~ O£S'}'iI.lgent~l, w~\iich is b.o1sed:lin Re::>e.m;:h :; Tri;;l!!~g~tlf ... NO~~l C"nD~ .. c",U0d i~;il


oofQ~" h~~l(l~5> lndeed. N .pm ha!! b ecom e '''I:~Jle 1Il~~jt)rmts;ie $Ciem)..; prcl£'rum for P;~"!'I!l,," ~s JeiITrey IBenl~etzen oflhe UIIn,i\'ersily ofOeorgi~, ;.\t~.u;:~~~ illi~iHti,'ilewiH Th~ lliI.e'I'"« h<Jve ~hel1erou~s of the NiI'fioiUl...:11 l-l'liI:rn<Ml (k~lQ!]1i~ R.e"%"M\;:b i:llst~nL~~, but it is Sk~\~lly Ufl:ilmg~l}lam8 trnil,u 5eoomldass S'!~:m,l!-;;" -1[L!llAB'E1fiHI PEN NIlS I



~UHIj fter 1']'lIH1.~dlf1;1~ect~tllli'ii[lr a


insula, One could smdy



IIIYu.g '~~,eill!)l~-

1;,lr slIrI[m::e. ~nd:Ir\'~dlm~; eleetredes csn be S:linmhued. 110n31l1i.s~elltly Idock the Function ofjhe area {.)filil:~ierest•• md the ,im:r;xl(;t'offocall dfsrlllplin!l <:~:~ th~~ ~){l: ;iLSSe:s~~(]- A Ilf!tam ic',<l~ THE VIERYW~S~E FORM~.R (ONIGRESSMAf!4 SHIE:R,WIlO[) IBOEHllEIHOF llH E HOUSE SC!I.IE.NCE Commill.~ ~icl, '''If.'>Ci,emis's.are goiil,!!: 110be more e:.n:ecli'\fepllf'!'icip<alll~irllhepolic)' .~rellil.lhey h~,ve 'to .. , learn mere aboul the po;~~,cy 'il\·'Olilk'·~"S&"f :FollJ.lil~ii: St<ltes. indJ~lsl:ry pi'l)"!key role i[l~ U S,innmr;;!t~on d!iiv~:'AM:S NI,;:\vs "I~d N9t~~ 25 M~l¥,p_ ~~40 J. For O\''Gf 4n ~~~ [ ll:1w.;' s, 'OO\;;~ ~11i];ing the scientific societies f~l,ke AAAS, Ame:ri,ca~, Chemical SOCiel}!, American PI~fSi,C31 S{J(~uety;aad J\ilal~'ri,~~ti e~a:w.;h &x:~~~)'.I:Q ''l.i~id'! [ h~]~.I!,g) ~u %iIlOP R .l'1.~.ef(;~y. flg<;'1i.nami ~~g.:.bl .• berdtUngI!lie public's ·":sc~elld:r.,c ilihern'Cy" snduanall dil2lll energy 10 fil'dnglim 1mbe~!lie\f:'db:lle pol[c)I iIliter<lcy of sc ic::mu~);ts, For ycm.;;,.i!l~\lelc:s!:ed this.,~nlal.k:sm: larglsh mldf,e~lCes in soc ic:ty ~].1.eel:il~g:~. vcr90% typic:d)f ~u:lmiued ro not be:il.g <tole to 1l1lD.He O Ul.eir ttli,\'O lie.Ul''llm'S ,mdi congn.:!'SSpo.!'Soltl .• V~ry. w:ry 1m",'~(),!1lnizcd \\i~~l the "House Sd(l~1C~ CommiltGlC"Wm;,Of did. ·'.rbysid;ilu tor I~hysic:isr, 't:llern~~I:. io~o,gjsl). heal lhys.ell'!" A rie£;LI[.~;IlF. clever qu..:rrer~p<~ge b ~essm] ClI1Ic:mnempomry "eivies' i.1II1 Selene!! Inig~:~:bea st<l~iI:~nd a [f!cIUIITi nrg l:el.lll.ndler of seien~~$~S' r~r(l'imsibi~hit!;S .[1$. d(h:!;!:lI'ns.

[ilnfol:lI,n'll i on (M R.I. CT} ShQiLl~ld be supplemented wid1 aaassessment of fl!lncl~0I19:11 iintegrhy lllll'JWLS~.l ~[[J~,iJging (NIt. funerional 1\IIRI)! !.I:IIld~.eurollt>ycharogi.cal ~e1>1:urI,g'~ I), Tlu~ hislory of ,'lddic~i.glllreal.mi:iLl:nl is pl~gu(tcJ witlnl examples of scientific evi-

de,nl!~~~!'IiLi~I$I;ld JI.i!•.,dfy 1~l;ia;tml;'i!nS wh'hoUlJ '1(3' 3iPP roprla lie ~mfety.md efficacy te s;d:~g. fi rmled (i\2lla lead to. unfoundied hope 'lIm.dhuttel' di~'lippo'i[JHlf'1!em~ fur desperate patients .;l1fadtheir f:rumi ]ies. MOJie: than ~(JOQip<lliems ~Il C~dn~ arlcl JJUs;~ia I'~porlool}j ~nd~r!i\''(:~ll bra i~!SlIrge'ry fOlr <IddiJC l:ioIm.before' the preeedn~ ,,\iilS SI'l)PI~~db;r[h~ f~~~~l i~ gO'lte:m~ menrs {J. 6). Ahholllgh Na~viel til. do not
3d:voc.~;tesurgery, '\~ caatien i,rlgill~ul!1, that the study

PremaulJe ,c:onC'lu5.h:~ns based ou uncon-

SQ~id ~)tdte [m~rilllu~il! fr.Qlfll:SI.;l!:Ir of 5~~en(!~. lTedlno!o!J~. ~nd SDd:e.1)o EmeriI1l5 .• Thie nil Pe~1'1~11I;liIIii~ S~a~ U'ln;l.I~fsi;!y. Ulmii"'euil.~,P>1l~.PA1,6~O:1:. USA; ·'tis1HlIligJ Pr~f~$MIr ,~r M~di~in~, tlmnw!J5il~ of .Ari~oo~; Oisti rng;lJIhllea Fmflilsso~ (If IMrori~~ tuilCIRiI ~tiJ~~ !I:Inillore!5lty,. iE,"mail~rllJl'@pilLedu

Eil<iI!IPugh "rol~~'Sor oflhe i

:5,hmuldnot be (mis"l<lk~~1 as. evide.:ncejm~tify-

:surgery to aure a.ddk:'don. The 5~:udy hasroe ~nany metlU)doh)gica~ !laws lh;n make "~I finn cOI~clus~;[I:Iia.OI" ven endJm:[~ e Dlamlage s.;J!llP,~~, s!1m,oking (~S:\I~UO!1Look p I~.c~0U11 ~s; m prel'!'ll@'tlire" Wr:, are hop iI'!:!:!, UmfH fiuwlf(: conrrolledprospective stud~es,and appHct'li~ avernge:8 ::i'e~rs c!I:rl.ier ..wTh,k;~:l introduces the proh!lbiH~)1ef'a major r,~c'1i~~ €,~~jd.~Uy lion of lumMll lab~nnory (l;lchlinklll1~S can bias, ",'he'll :sHldy iu!l d ~lnage to brain areaswi lITh iiln~Jnwe~hc\m Iidity of derived conc~lI:si.m.ls. :S;liANI:S!lLAlIIlF,I.~OR.E!~,J· AIDAM [BI:5A.GA,l IN THEIIR RIE:P(lIH ~D.AMAGE liO lilHf ir:.JISUlA Inc In oX)' n~nel.ion, hu.:~ ~Id.i,}gth~ insula 113l GIIilli'lj'N\CKHANN,,2Iil1iERBEIFlT D'.KllEB,ERl di~nlpt:s addictiOl'l1 to eigarerte :i>llil~o.~ing·· i'Dladd i~i on •. retrospect i.\'e.l~se~Slflelll~ hlle o (26 J~m p. :5Jlllwhieh .• i~bfl~~d QI1J:;I re'Mio~ i"Nll!"va[ between thL\l: oceUf~~H.~~ ,of the l~.~l!I<kriirflll~llIt01 P~)'(hillir)';.lDhihioo oI!1J5w'b~am.i!· Ab~,~, Cdlili!iI1!:!1 UiI'Wef5ityINm ¥or~ Stali' [P5yr!ih~~t!il'c 1it~I~ruiil~ a ~ • spec'l:ilve study ()f pariems who had ibr3[~'I~1 .. d1l11 ~ br<~i lesi O.UilI and iIs detee ti on oni a u N!i!W 'i'iD~'I:~NY 10032, IUS . ~D~Pilll'lll'l@fll r4@mlJ~!IIgi(<l1 A of ~~~~o~t-O. . H. Ntl,(!Ivi 11;'1' cd, ~o~c,]tid~'dHl~ di."gf!;o~t.i(:l>~~~nif!) N d~ifnc~! Th~!M1S~I~cth'~~:. $w ~iiry. Co~llImbja IlJliihrer£l'l'llJNew 'Jerk $!lilie P~.chia~tic ity of bmi alesie ns ~]1,,'lU:5 ~hf;linterp:rel ,!t~on I'1iIsUWte,N\~ Y(lll~ m 10ooZ, IUlSI'L dn~n'jgl;;l~Q~'~~ wasresponslbleforthe iUil.'ml.~ disruptiau of nicotine dependence hill some ofbmin she-related kISS of'fuuctioIJI pa:liI.icRef,eJ'~Ci~ cases, The cI:rliLui::. COl'IsJ]stem: w:~dllrUl earl!~.~r~y dinklllit The m!l~lol!i sed MR~ ''1nd u 1- l M, Milli!l !i!llill., -I, Pzy€bJQNy 16:31.186 ~2G[[M. ('"[. scam; [Ilmill: may not be sel~shi"'i::'el~Dugh ~i.c:r report dC:llcrib~:ng a ynlm,gM!BIIli who lost a s, M, !le:r~o'd~ [):,S"Zm.i.· N~$d. ~5"U1S7 ~o deteet po~el].liaUy oo]ev.. lI[ 1J([;:l1 lesions, u i[I [f]lle:rcsl: ill abuse of sabstanees urter a selee~OO51 3. t Fujill er ~l.lJjf~ is, !i(J] ~2(}[)i2i We !m~es!t t~i'I!lbefore any furm conelulive bilateral stroke o:hhe .~Iobus :~Uidus i!. I!:. J!. r~rk[fl5.M, :Slil~er,C, le~mn. Pzyl'hap~cfflil~'oIO'9Y 11). \"Ihicl~ is a!1!im1POrhin~ !iu;;la:rli!~ [[Ii'g~l of ~i(l!il$ about the i~sul.a:Si~volv~m{';M in !lieurBN1.) '184,. ti2Jl (JOOoil&. Ul.e' :i:nlsu In 111). 1 ]owcver. I.net~lodol.ogi.ca I ti nle' di.eJjlendielliDe .'Ilre est .. 'I)lishe(]. li:le resalts :So 'C. OrelJ~I1~, '~i'!'(,erN'wJol. ~. sss f.ifOO::!~, ~i iwt i,OllSi nherel1 (~n bnli n I~$inli!l $h,dll;.'$ af U~is «urospec ti \to: s~wtdyn~edlu be v~rinl 6. 1,!!', Hall Md!.tji!iIj~(I):!'.1I. (:ilOO6), Ublldermincllu: \''3I:ltll~{y of oo~~c~m;im1S, fie~: lI~,ing pro.s:pective s.ludie:sond 01,mm:e ~Icdvl;:d lh:n1Jn1 thi,!; ~nu(illy •. riigiOl'DUS ~~lCI:bodo~ogy. VJiI~d;;tted h~m1'l1~ R,espo:ns·e the lll<'l,i:ml. o:utcmne memmre is a KC(l~11 of be h,wiio;mi :~<IbQrnt,Qr)f~e~:hlm,i~(ijJlq,e!:S c!Duldbe WE.SHQWED ImiAl SMOKIER) WIIHll DAMA:Gt: S-111loldng ~h01Vio]". c$p~d.:II Iy the: di I'F~~I~li u~d 1.0 ~~C~'SlUrc c.mvj;ng f1S \,'\I~II. a:5 siUhjcc~ ~O~ht:lll:Sllll~, ,~'Cre 11I:IIorc::l:uke~,y1m::m !>1,nokeTs t ,niicnill{.) "''Ill ittirl,g snlloki~'B\?i ..idl d.iffic::u Ily" f ti\'e amI reiulbrcilJlg l:'IJc:~'~:So:f,c:::ig.m~Ues {41 \\I'ilh da~~.f!geiDlll!1)I~Jilf 'bmh~!Il"gio~s ItO be abl~ fn:}m ~q~.dtti.!m.gf;1V!okin,g:,." l~ rh~ pN;)~"e!n [" $;~rok~ pat iel'lts, oOfi fU sl~]gic a I pa.t~~]ilt!:, to q nil s~.noki:~,gf:'~siIy..~Inruned,i.a~cdy:. \,dl~.OlU

IllnsU~B Bind [Quilting ,Smo,kling


20 JULY 2007


[l~~ (anr,er (o1il!!~ I)!' Mlldlcim~ IIr.'r.! O~y,IAS22it2, IJ,)~, ]lDcnrfli~i'f~ Cllgniliv@ [N@uros(~@m~ ~m~!!Iliiltg U\nt@r~ IU~i¥E!rsit,y ,or S>l)uU~eri' O!Urorrm.la, los .Alf'lgi!les, CI\ 9i0089', IUlSA.~8r.ti~ ilOO (rro~ivil)' [I1IS!i~~~ lLI'ntil@'(:!;ity Solltll!i!rl!l of

ImrisialfV or Cog nit1;re N e~rvu;i~rt(e. [)eiP~nm'lelU101 [Ne1.!irc'Log~ U:n~!fe'/sityolliowa, C~Mr (o1lege ot MuUdne, IIOW.~ CDI!!'. ~A.S,21,1l,2, U'5t!..~l.iImorawl1J of ((lInp~ila1iio'na:l [N~f~rme'1J!09', [)e~!tm~l'lt or Ne~~I,og)'~ Ui~'I,ii~1ty el

Cali'loi.a, [LQ!i ngfil.es, A


90009, 'UsA,


'"lie wihom "C!~espo'rui~nc.e~ho~hl be

ad~~i:S~IHj," ~,'ma


rel,8ipsung" ~.nd witheu; a pIOl:F,!l].is:t.cnlurge (0 smoke, Om' COIl';;:I~lS~Oml, from t~j5 result. cO'.mib:ined ~lI"i.~hl re.su]ts the previous luncl:~o~!t!l il:1Iuigi~lg Sln~dh~s ( ./-3 JI .;;lilldiHl '~KUih~ Ij:s.h,ed theoreti C~'L~. fnmlcwo:rlk .lor :U.U'I,sUbl nlllc~ !f~l!l 5),\lI'~!~ (4. that lnsu I;~ d~!ln.!tgf"lim',f;;rif~~~ W.iLlia. specific "I ps:yclll,olo~ica~ precess thf.ll 111Inkc;:,,~ it diFnCLUlt t:Q qU~lsmnldm.g :;lind d~~1 pml11o~e~ relapse, mlli1lM::ly.~he com:cigm ~t"'g~ 'til smoke, \\l~!ltr'C ~o~'I'idle~l[ h,~ltl:~,e t a 1:Ji~,I;omi':lIl"lldbehavi or~I~~:ecliil n~qlueS"\~' u~d \\c~n~~PP!:'il)}'l'ri~i!il ~1i.)r !j.f &1t~ f~nd adeO qmnlo: to 5\.pporl thi:-; ccnelu ~iOIilI. Weelldre~;y agree wbl) Vore[ el al. then our nndi,~1gs do not jill!$lifyinvas,ive I:IX<ll~rn~'I'ilts for stunki ~,£acid iet ion.\~~c lrn.eVJCr s:LJlg~ gestled in our Report or ~l:ii1iy'!i\,l~tere H'Ii!L~. else s,urgjic'II.ly dm'l:l:OIg~iJn~ l:beills!u.!I.<I~~to'uld be 3. vi~d.'ll... ~hL;l!:~py for smoki~g a,~~.JieliM. tndeed, we jo.inVoffl'l ,el til. [in !>ll'Ongly eoa,(!Ie~min,g a~l)!'sl~rlJICi!il m~!'!iln~ latlen 01'Ul~ insula to. aehievea 1.~,el·.,,:~eHti(;·im, Apl2lrl a fron:~, ·t;~~.('!his.t():l'ic~] excesses of ps.ychQ~ smgeU)'.. ~hi::rei:l: clinical evid~n'C~th~u ill~~]ad~VI~~ge: 'Q~I) impair a '!"'driel:), ofruncl iOIll~"


SI.in cx:i~Is...however,. dUll insula da~ll,~gc disrupred memory fo~~I:be errl(nkl~l.d experie:rmc'(!of qu irti ng .. ::md~.as l[rnemory tor hew dUTiC:l~h i['l:Y'aS~I) q;ti.][ C1llid for urg.*-~ih,n~\!(;'ire fell <Lfle~'qui.iUing, nl.~spo:s~ibil~,t)( seems unlib;I)'. f;s~ciod~y gV'i!~f)th~ vivid desertpuens of the experience of ~]1Itti.~,g previded i by sn.me 0 W1our PI!llicnls" A I SlO. wjtn.1t!lund :;I
~r.I '~



:U;" ;l:l~, (200i2l s, A.IL.Ir~ (,!I'l~"AtdI. Goo, .P!;~hicI1y519. Uh2 (~OOZl. "'I, oil [It 1},U'ii<l(;io,. ~ f;~1Jrng!Of~! ~,fJs: &dr oM IK. [R. ~.n5m1l,~ al.. lI'i!Uf~ydii'lp;IMI.l'mll!rnll'l!'l}" ffIllojj'ttJ!11 in ilreMakfugof(fJ'~s~ ~aK/IlQn; OlI~J~al 2000). 5, oil [I). Cro'!~9,l!lWl,&~; Hf,rI~(f. 3, 6S,5 {~OOll,

IR~~~I!I!!K~~ G.l, W.mg ~ m"


Sri. M. "J i.S


t[f;nd for p~.~.:l:e~.1t.\1 wiUl.~nstllil. d;"m!<l.!i:e

morc' UI<:~ly thaul I)lnli.'~nl$ with d~ufllf~i,;: ~lI'Iolliler :rrerd,ilOl'Is~o be.abstinem at Ihe ti,lI1.1.e IQf f~~ s;W.<~y '{i,c'.. ~n h,fIVC' q~~~~mok:ing nnw h:iiion Qln,eO, ~.fird.llg thatwas fg,(lt suscep[ib~.e H) recall bia.s. Nonctillelm.s. we agn~c Mli.~hthe need iWorp.llo:ll.pect~.ve~H!di~ and
haw .~I~l'endy begun ~ucil1SlWldi~s.

,6, A. Ard~jJ.,.IIpmiar0.U1l. i'IHl999t 7. IF,M<!lm, S. i'oIra!li~.,~. A. 5I11rliM.lH'. G. l.ilmb~JlI!f, It G.

liIalllil'i, !l.llal'ildi50, It U, Ro!±iiluQ:n,,}. ,i!/M'"N/.t>n!, Oi~, 18", 1QJ (i 999'1 9. IH\..PIt fl.a1.", ,'Nel1lo1ogy 66. nas '12'0061. ,9,

iJ~1~~6 m~'!1.1.

such as hHlgUiage ,[6).fl~~G~,HO!l (I), and nworj (('i:~, and can cause Si.~li.ficallil. ,c:rrdi.o\'u(,:uj:~!rm(lrbidity (9). AI~bolJgh our results do hllli~~e

Vorel 1ft (Ji.po~n~ out certai n teclmic "I lim il~ll ions t hal: Olre~I:JI~Jeref]tliO~~U 1ll]~ImlI"'u ~~sioil1 stud k~s.W!~.QSI. !)l~bl'y thll pro'bleV'l.'n.of n ' nmlsel.oC:lil.vi[y oflesioHs,'!oVe .. ddressed this rrubl~;:i'll Ih,~ough an Itn:;llrs~s fh';;~l Iook~d!i,t: : e Ilee IS, in reg ion s su r ooll:l1I.d i.~g the hH;~l]<Ii. Through this ~]Il::Jilys.is". ~ot!:l1Id that the we insu la was the o.IiI,ly :regiQnin \\~I~fchl~sl,QjlS~ad~.~dgn]ficanl: effect mm Somal illg ad(!lliclion, Byinchltdi.~lg ~ larger muml1e r~ f su bjects ..nd i



the! fiilrst

UillE t4 EW5MA KIEIR5 ITEM "0 PE.NI N G U f' (1Jl May. p, S~. I) .:ii:~:ne!' ~:h;H"GeJilmll,Jll l,bys!i:c iSit

ofVi.;>mli(l!hl AID MriOl. has been me lhefi:fji~ .IMH'I~Ch inese
lIrt!~,,!If;l:r$ily '~e'I'SOllto be named seience dean ata Chii'i,e51:.li~iv~n~i[y:' hi f~lCI. {:;~~Qf!:eli,V: GroW, a :Penll S1a.te gl:.]d~l~lle1.111 .ho.r~icuh~1.re,.

RO~ll~m('!R:li.1pr iOfUl(:

was de <II'! of 11~:~'~Ueg;e 'i;lfAgr:icu lture <It

Chrlstian Co:~leg!.t in OU3111,g;'l:hiolll,



hi~If1. fr(r~~ ~92.2 tol9~ I Cn. Cant'On Ch~is~,i;.nrn Coll.ege beeaeae UngUlan Uua,~\f,er= '~ily.,whos~ Co:lliege of Agrk:ulluir~ mr;.'irglL::d

rherapeut~c~llilrp] i co;l[ion!>fer example, the de'\lel!Ollmem it) f d!'l~gs lh:u!llrSe'~ hliSil~ la fmru:::t ions. lJeb1r\':iol .•1.1 the r,OI~

pil;lSih1!t Iilddr~;l~.~ th~ bod ily/vi,8ce:r.~~. co~,upoJi1l,e.~u:s

or sm.oking,

le:'l:ion .m!A.pping ~~ehlll~ques, r~L~U s.H1di el>rn>m~ re be able tn de~ecl elffe'Cri$ o,f :Iesions~n other regiarill!> lJiII<I! pfi$~i My ph1-Ya Ii'~')h~ ~.~.!)l'!d~:ec~ tiou a nd ~n<ly be ilI.ble:[0 ~r3.(;e ~~·f~~t\l.w~tl~illi:mb!i"~gi~)~~ (l,f the in:illll.~.
O~~:1igoo I ~nr pC:liforl.llillg

\\'!li[h thal of SlI:UI. ~I, Sen Un i\lcl'sil:y in ] 952 ¥ f~r:11!'I S(]~uh Chin.:! Ag.ric~llh~r.!~ O(lnl;;g,~.

50mh China Aglficul!lurlll


with .. CLlInC::lU c~:ro]hn~nt of )6.[100 SUI(!liellits. Gtrra.rr be.g~1n te.u:: hi 111£. hcrt ieu h me in Chh~a in .~ 907. h;igi!lfii~lgae'~.m!tli1.f)l'

mld flJnarilHull

of ~ln:ml~ tU:::li\f~ly to

lhi,s ""wd)! ,\r,ISiiliO,t '~he imm.edi.·
:u.~ disoov.~ry ofa "',cur,e" for smoking mkl:i.ct~an, btu I:<]UIm.er to 'sliiLecl Iighl O~l a blrd.in reg iOIl dl~t h m, boo ~ '~il rg~;d)fignor,ed

llilrmli~r the rr0!illiess ofireat~

ments-'¥e never used the
d.escr.ibe m:illy f!!SPI;'el of O'ur fi~ldir:lgs. The fact thm our s;~.lIcly W~l~rel rl.'l~peetiw;; faWSe!iill
~C'I:ln"'c ure," 10

of par~:nership be[\:I;'.een 1"1::'111] Stale and Scmlll.l C'h:ina Agr:ucu~:r,"1.I,<I~IU~i\'\e![,':S,hy th,u is .,;ti.n thriving il1 l~l:e f~rm [If ~ruini~.l.g <lUlld reseallCh cona'btmltioll~1l p~,ant b:i:oIogy. illd:'lI!(UliIlg .] jOi,[[H LiI.!JO'f.fltOty o~'ROg'l Bi()·, logy~nl'llug~l.nlted by lite p:reS[dCnl'S of Scm{h Chill ~ Agr.]euIllJ~ I lJ!1 iv~r.sily~ttldP~UlO

MillY 29

.of ~h:Us.ye~1 r: Rupp i;1;ppe..<ll'$



m lib oU't.rec ..~1
\",~~y \'\I~G~""ci.~!d~d

§ p<l.tueiil,~:s who~!i1Id
.~ and

biO:l~, Thif'O ij;

impa,i r~

in the drw[.g Ud{Uc.:lion Hlem~ ~Q b~ "~'l ]I,;;ast) 85 }i~~fS ~[~u~[I) mer]l t1l~ti~k; lUrlol. \ll,ll:: hop", l:hotl Oll r :!ll1d,~ of "firs.l ~riI,cm-Chinese Sic:i.em::e de~n .11. 11 CldDltl~ ~nliver'sily." ings ,s:~,ur frlrthe~: n~se"l!'Ch (m ]ONAIDHlAN R LY,NCI:l tbis ~op.i~.\vhi.c~, ld[i~,'l:fU'~ly could le.,dlO beuer t'fe"llllm.el'lts [D.~p~n!'l'!l!lill: 'O'~ lionk uUu ~e. 'C'gUeglll' !:If AgI1(u[ltu rai

2!11:1!~m~~ (If lonjl;-t~r.l!i!ll"l1il~fII]!)r-y

for ~n.liltlkiw.g .~ddiC:liog).

obt~i~.l!(~~d ~::UlfOl:malioiJr

:~ (rom cu.Uml;lra V" !{,~'h\"l!l~:'~r.it ,",was i_l\l.<"U 1<.1.1:1[1;. Tl1epo,sslbduy

NASiR 1i'i.INAq,VI,:!. O'AV~IiJ)! IRUD)RAUIF.l.,~ Hl!rNNIAli)iAMASlr,O~~~ AmOilNE BliCHARAu,h

S<i~n01S. ~~fttl!lyl'o\;;IIII,i;!l'Si~1!iIA!' lIrni!;\i?,r]ily, Uln'Ti~~i~y P.a~1!,M 16tliO<.2:-420:l!.IJ'S/t IRefeflllliKll l. ~.nqqmA~/iic. R1ew> t :l! U1iliW,





NIlfi'iS F!l!CII;ISl ~~laij(m liJl!'Ji)ilt[(~ nn~'o\I!'~dltm-esiilL9le'gei1ii!" hi[.·~!f!i5iQ2 tl!!ne;,p. 16900, mn~,iMiiid!Ja~, pictured ,''''' tPli.!le.l.69l. aJi'e from Sibm.a oot.INo~tlfl ~ild 500m AIIll1nt.a. '!iIlIis W~~ ~lf:'I5ae,l!'1~ rOQilolt~e p.rQ'ble!fl'l.'"'(i3.hme, p. n9'~). V~e~!$!,~nteP'l'cE: ~f !!'II:!. ·n~ neW! W<i~tnC~fH:1.Mthoo~llmOWi!sr!'lo eIi~lbil <i rel<itl~1ij prl!illtU~e me·$'\~I~. iQO~ ~~. bam li!aI.lJLo~d mew~'iind ~ dlilleid qlOHlpJirl!le plh~
."A!i1e\'j lwl GI! lifaMlln9J teache~~~y .1. M.E'f-vlis U l~e,p,12'O'). Ue ol!tl~:l-i! miM~lllr!d 1.Iir;~lame' (If laserl i!rrn!i!r" al ma~t~r It&l;[h:~1' lii1 !hi! Uli&l£h IPro~!F~m at dill! f


lIniiv!1f!iil)' or T!&<IS. Austin. oCEm5itY!T'ianomteM~sIOf~rrayS" byf. PiTi~C!15lyf'~m. ~l5 b ~~. 2)006. p.l:llOO), The lIJaiiE fer NW2 in tFig, 2iCaoolhe ini!'e~. flli' NW''4!9iill FTg.~B Wffif1f,OOiJlreot, Additiirmili~y, me [( 'tr:a~e in fiig., $ati wa~ ll1i1prlO~)' lltoU~ I)m !he lime ·alis.. TlileC~r;I)edl rtg, 2C i~ SIilQ'i'lllto ~h:~ wiIh IMI NeIWS Fo>c~ "N,Ungllte ibram M ~o.i3t b~ C .5tJtrmlt (l.8Maoj, p. 9161.1he Ibraffl~can~ '!he Irep~ml!! rig.21B. "n~re a.~ mf1!O!' dlf~eJI!~es I;)ei~ b origina'l and ~ep~I:1:edIo\elr>lOO5 af sdIizapmr'~liTi( palil>n~s '51'10'1'11:1 mil P~!1I;[! 976 'I'I'i!1l? ~TlI]<I:rnllO m~~ffiI1!~Ihat~~1! brfl]lll~ '~@rn .or 'Fig. 28, dlle to f~M sem~ in,!he< ~~m~~!lh\'a!'e" I1g,4B~ stI~1IlM!1l below Mili liIle ClOriliangill]J r<ll!l~dly, lOOt ~!lsugg:l!:rt lhaitlh!!: 1h~r,aP'J' desorib@d in Un@' illTlid~ "a~ld j:!T'l!V@m. ~w'dh rtc~!i!din~t .' 1iOO r(\ptCiIt'~D~rac~far !Ply, S4A h now ~iI!iIlllIbl~rn Ih@ 5!lJppm1ii~HJ onliJlt ~iTi~l. dl,!U'Iges, llIe paUell1S 1'W!11(fd medica!»" I!Ef~re and d'ii!lrrl.lligl.lilel~rtodI\1lie." SC~5 were!;mn, rbrlhli! R!lPi:!d~~n the 'IJrrg1~ Fig. 54"', !hie'.mriQl'iS, showi:mg IJIlIrwml.im.if'!LtiQ~ ~re rm]~plIl~ed; 1n 'lIieco~reC1!ed '~~r~, ~~Jecliolll u indi~t!!d IlIyU~ rl1"gioo,or IT!!l!O'te IP~tivf ~_raftt po1:i!.!i'ili~L INI~li tfocus:: ·(!(lsing dli~' 11II!lon 1[00Tr ~ IB Ie Slim nL... __ ...IIII1L. _ mO!ll dise,)~ '!le,i1)~~·1iy ). Couiiii' ,i!I~dl. 1Ilb~ !!!'~r.o ~rmed d!]ringl '!li!elin~U ~d\)J~i!!Oor !lae f1[1UIl!S, d ~!'!e 'of 'llie ~sn!li)il ~r' «I~ ~~iom ~ Uile~pe!f <Ire <!f'~'led. 1Tb~aut~hO'JS ~~lotiJire for 'tllle~~el'!rgrs ~lfIlh~p~liilih~c :~I?er. Kaiser n 1. .M<'!,y. p. 820tJI. Tlrie talbr!! O:~ _jl15nS Inilddltiol1!, tille ,aUioors\¥OlIIldIlllC:eto,dilnry ~everal :potrilll:, '(I~ IDe!<iiir~d~rmlnjJana~1$ ~",done pa'lli1S22 ili1(!ouecU!I' li~~ecllhiil 5ailfl~ SCI il'l!lS.llll MOO!Id'aMllilel;in tIOO:R~m (\!lI;Id!I!!IJirs, I~(., IIo:rli'<rm, a:R)lilnu milJrorn ':OIlTl~~d1~[:gDlIfi. ~ilt'QI' iI 2@(!5 'QIWDn-widle <I~o~dliD.n Wi)In Fig. 10, ~le,jmrOl(t!llutu a~ iI1iU'Mi!I,'ilre ~giil<!i~,~re iil)til~r,ed"on ljIiff~nt ()omp.Ul:efs with ~I.IlJ:dy in IIfIll(Unm di!genl!filHlln. 1h~. "in!!!~Urn~ ,clr~t~ ~t~~!1 '!tIe ,d~.<!I·~ O!nd .~~ ,!"Iat iiil'f;l!1I~dl~gb~ !¥oo~el~eli;JliileHms;1~!I'i~lud\ocllM; p~o~lI!. oo~ ]1.7011 ·rlli!l. (qH)In rig',21~'~me tpulbllillM~C.aJle bar IifeP1!l;m lM~caL'e or e.aCl, fmivicUllllJOl>re; 11owi~; INiews lFows: "lIhymoiini: dlllk-al protIlr '!hell a_rea!b1EJ~IiI:v I!)Hs:fl.lr,e~ti!"e to eaiJM ,oaherror claiity. (iiv) 1!lIFiJ. 48.. Mtil ~ere !IfI_f(!$U~ 'is~' ~Ir~r ~ dJe,c·.u:!,!~·IJoI1i[l$1'¥'(1ii"' !IV ,~. :wq,u~mliilll'" ~flli!rlrl'll.d~~p1jl5IillTlUiirmolll'''' 1lI01~l1TIllitaD@Otl$~,. M IrJl fig. 52. ti1c 'g~tim,@~imib~· M~ roc 1(.4, M~~, p, ,6~Z~,IHym.oo ((1~~mIl1\\lH il;~oolJ,!l.ri@n NiI!ll3and mY) ,~fl!' r~i and' ml'!s1i.li:k€'~' {.3mOO hy looulll1ing 10 g~nflImdiooi~~ k",l!S 'l!'lc(w!lCIIy !d~nHt:~ ~ifI!lI-e oiflid.e, 'lU~U~~ aiild: ~he U~!!ol~llI'!Ijkwl!xk'li1!a!!,!plifL!lr pair:aiPet~Js.I(lr d!'!~~!iJ~Uon. (~~!J'I :F'lg.'55;1\ Sh~ ii ~I~ornu!r ~mi~ramwral.sci~nli~l.ill '!he r.e00il1li I\'M ~l.ij~latl1d mrerihe ~Our:ie QI lOtJf 00ur!l. H~'I\~, Ut~ c:.t<~w~im~~d ~Ufl W thE! NaHolli!l ~ ~mtilUl.I!' or tD@'Iil~I.<lilt'll I( tp-lpi!tt~ol1~lfdUllillgillIl' lim aoo '~t ~e~'Ef<rI, ifii1lifS a-l'ihe IIIl;pe1fi~lli. 'teo(",n!ihmti, nf!!JJ'oml m 1alO mtG Cr·~l\iofaci..~IR~eilrdl, r~0m.s~ ijIT d wilhiliLy. EdileiT'!li' l(ho,1<!'.!: 1!t>dmjD!lI¥tiIl!r~ (21 RepQrh:: ~Oli!'tica~5igJlalur'~~ 'cf Ih~ Aitrar'QiT'IO'l1"Eliolirrn phil:~ ill~irJg,te'~<rlledi carbon na ITr CMN, ~ ..,.~ ..mJ ""f .i .•. Apn~ till, 5160), TIi!@1lIiI1P(lf ~,~oo in, Ih~ m !jjl~~~" b~ :5. Zane tl rd. m:l.1ItIy J:OQ.4!, . 112:~~. ~OI!Tii!' p orlh~ dal:.'l~mdeoodlu~ioRl; tP~~ IUl!!l [C,111, tKi!TIel',rf!,I,_ (hetJl, ~(; '1 w.nWCl oil!; nOf!,J'l!ljn l~.eFcI~,polt ~~re ~.~i!vi~~ly IPre~>enmdin S, Z~ri(: ,tt I!!~, 5up!!'tlmti(J!'J

Raifid~m S!am.~e$~ "C:OUi1I!IY coo'Rf!ii!!i~ (2') Ma~,p., liIie lut ~eJi!teiiee Is !ifiOO!fect !II slr;culdl wild, 'Th~ Oli!"qoe may nOl (llIt dawJ'l '0:1'1rood mm~IITiTp1101ll,bul: ~!t~ ~!I.rsl Uiwill v ma~i'!US@ ur Wlr;ll@301% of a wood lir'f'!; ,~rl@rgy, ntuKh ImOI1!'IhirliT an 'op@nltr'e's 1% I!.'iri d@Dq!'. ~


l129•.1!o.!0lZ!J]~06~7 f2m:m] ~lia'5i'!OW tE\1Jr:a~ Ibit Ute' edL1:Ord' ~t iOLl!llliL


Re(p(iri!s; '"DetE!Clh,wn, ,>tlrnul~t~oJl" .. 00
~lTihi]'it[oo cll!1e~l1'Ilrniill.sigilflats wmll thig!rr

MiQl(l"(J!JI~~!JlJe.:l 34,413 (2C1(14).I'iIhicli 'WilS·put Il~iI p.rO(~edingl;i 'Io!1OII)II[Jl~ ~h~ 6111 hom Il!i\llE'fn~lijon·~1 (O'~fefe!lce< oil! NieW PIii:~!'1orn~,Ii!3J in MesosGO p.1c stmuctLl!res, olndl Lhe 41h Inle!1f!ational Gof!fler;enceon s~~ra(f'~ tnl:erf:aas 'OJ,rwMOS~.o,p~COev[~es.•lihe au!bon, I'!!:ffl' amd re~II,~~ lilorllhi1i ~efffelllJ~ dU)!lJIll:!h~)vebeen iDliitiil:ed. I

fig. 28 ar~1IlIC.

IN "R,edefiniililJl the .llge of Cllolwiis: ImlPIiicati 1:Ii1!sfur 'the PeulPII ~I!mIOifUI e A.,e!f'jtCas"

~4!0 IlS

:2 msec:

Gillii'Yl"ayi1l~s •. iDavid G. Arnde!C5imilii!(,C. R~ld !Foel'riifl'li~,1.S'tlil'~~'tJ. Pi:ede,I,. ImJli!DliI,a K, (haV5iill'liI~ ,c.. Va Iii!i')e lfIi'!Jynes ]Ir.,Va iIliCe, 1. ld H!oUid<!lf,Bril.lt-eB., lfItud~el~ ,Mal~cieLl(o:mfe.Ld. DaJlofid J.I. Me,itjl)e~, ~Ulli',e Mono~, fodd Slul~CllilfeU" N]Cioibe M .•Wagll!l:l!s:~<1Ilcik, Pe~ier :Wi9Ia:rnd~ Roillua·rt M. Y.aiH! ~II
Walel'UM. S,taHCIfdI(Re.pen5. :;n; Fmual'f :;;U>07, p.1122)ll'm~elil,u~iUl,~nI'Clr· ma!io:n atloout tlii~a~ I'Jf 'ro~ 'Qmo[~ filM Mhiril'E! not ~ennilIDrel.~ ~btislhe~ th~ l!]m;pora~ ~paJ1 or thi~ I~u~~u!(ll ~[)mpt@",. 1m lihe I\lDI!n1~ Only a Ic,on~inuin!!l P'lOiJ'i)m of !r.lcilometJ(c ~tr1li9 ~1Iii!l~OOirui S'lira~ig!Ol;phi!: jjCin'el!Itioi"!~ (<TI! <!dd~ ( th.@lin!lE!rin!1<!mbiguilYi!OOut !he ,emeJ~c.e ~nd SlJ)faiid oJ (]o.\!i!. 'Cl!!!!UIi\e. MI. ~~al ¥mffl'.M:iilntl!ff1;~9!,(lf,~q;flrontmllJh:lllB ][J15:8~6.il'3:.2.0b



F~!!J.48 •. •


on "'IRedelin~ngth,e' Age Cllovi s: hUIP l!ilcaltiolilJs .,nhe P'eolP IiiA'g oUbe


M.kh'!:!el R. 'Wa~(,!!'s,!!i!ld lilhomals i

w. Staff'(I~rdJ!~;

Hiaynes~f d, mi;5!!fl~r~enl sever~l~ i!~P!!(U5~' !!Iur SUldy. 'OU! ffi¥i~ di!ws 3.f!d Qther;:'!rm~e(ll(llQit:<l1 dOll<!inm~ thoat (t01li.sd'Qe5 1TI0t JeiP~l~ne wM~e5tQcw· ~oo Qf tJt~AmeIliGl5, arne! we .oUeff!'Ci bolJri hillman Wl1g;ratioo and lecltmBW dillll!s.iG:fII a~h!fp~~e~~ te, ,.a~, the ~>:paM1onl ,01'(ICI'I'is, We stand Iby lihe Idata anAiioon~lIisMin~ jX@sl2medi1ir 'ilUI"Q;~illO'l rTI!p;!Jrt. rrull ~tal
~Mie!!cl!f!'I~g!.l;)!,'~(9~OlI!1!:f!;iillJll!lLI3 :ll7l5B3tV3i2Oc:








ill deep-sea

lI<1wlls" revel im.the unsuspected



'fj ~ .~ a



sea hot s:prillg~~, nd eold-water corn~ reefs, a D.mnutl..~cdeep-sea discnver.~e!' lln~o~,d jfellr ~ncr ~~an:'. K"f)sl.nw ~~u~:~ld,s t~la~th~s bm u~ ~U!.ij.vns QH1~ebi{il:og}~ I;l!f~l!$ d\'~p S~ from e~dhV']Kil '~o ~~n(ifi:r., '~hit~iimdmilii~ "pri~in~" (JoI;.;:iiil wi'~oom~ss b~!~glf~m~pllf;;'(!1 (h~ ( is tesd tOo focliS 011 't~,U:l: el,nrl5wlndel,·'_ e'piic preci.CJus. mi.n,u:tes, of s~I,b:rnJers~Me ,re~renochadd Ib~ithe iinsid iOlJS "hmml'l,l footpri nt across the 1 h~k~ni:4l gai~ a~ce~ to thi:;; h~nlm~~y l>P tl;l a p'i~~~lIUy::lm;,!!:~~I"\,'(;(fflJ (JI~otl.\>!l;rVmiQIil (Ill deep sea"; '~[lle :5eN.'bt;d $~.Ir\:r;~ 11igh~m'lrli:l.1iI ,I i.nllia,osp:illabl,c eavieonment alliu! alill. <II:riLllIlliII{I~, ooslls,llffil:f'up.icaiUy ce~,~ :iega~y of tens o:ftllom;a:n:ds OfD;UeJTiC~ljml ships tered onpilmpu~m I{JC',fUI~·in 11 ~,it- sunk ~l1!dwtling on the .'51;\1Jbeu. :~;ml~dg·ii;d!.'ii!!)f enthe Mm~'e .1 r1iIlll.d:.. tb.d Iive lh,e SU~Q;!'IIIIIDI~I8:IPi there, Th(! Silem .Deep IX.)!S eral sea of ~.lOJ1biJ(]t. [hmus<l'lmds of l()il~,5 ofrni ~itallY orx:tnJauce !:lCl:l~llH! Uiscuverv. O~U dli~lemmj~ is l>quarely belhom0g.;: tOo thQ~ ~1.u1il(t themes t:~ed ~IITi deepwater; Ilmliill~ol'ls (If curies of ECDlogy. and I:nn stands aparn1liuIi o,t1inerboeks fure us in Ko;!;~O\'/sdmptreron eliII"t!lJClioRctivewaste and n l'I'lu::le':lr reactors Co Ilse rv aiM ri or d~l$llp~d .fl[ d~rthw~th~o ;u~(:mp[ ;!It oontf!inllbol!i 1lfe hll the abyss du~ m~'[e (:h.fl!l,gl': !l:~d~h(': ~~p ~~. d the [Jeep Sea to TOllY Kos,IQ\:v's lhm1ghdtd II~n:',n,t,. ud residual a DOT O:lm,d!IPCBiS ac:[m];!u~ As the d.illillJ:!l'e VI'<I:r,Im.1S •. deepw<lU!if by r~.r.I}" Kos!.ow :I!u.ing i~~. d!ecp,~(!.", foodl ehsins. oooolmllsofdL'ep-·\!JljItel'n1i1~cr.if!S. cin:en.liltio:n p<Llli!.~IiI,!> c:.h~,llige. inUniversit\l (If Chi,cago mi n~il]JI e.x:pl()i~ati~Jilt~.ruinl't de: creased C11lihol1 d i,Q1I!ii de levels Ko,~lo'lili oilers a lit.my I:D f new ve:mi:l'u res PfIlS,S, CiliCilgO, 2<007. ~twmioJrl. and ,c,'JO!~la,l'I'!Iil!:1ialiol1i of ae.i:diry the ace::l:n,.I~Qth~lli1~s .of 1~I'i~ \I'"ith the ptucnti~:lli)ehll~~.~llgt:: IUlId im.uh the 29~ PIl, £35. [:22.50, the deep-neeaa wilderaess and Itlmyprodw::livi[y ,altiJlf' surface deep ~<I ,eVel~ l:heYD"ei1fi tiCClimaL'lllic <I!I:ldsaciml: ISEiN 9~a.o2 2845'1 i51. ~~a1b,,;\~c:fits;: Ju~ [hl1;:~ dcca.d.es~ft~r th(l' fin;~ his calli for n:c\V slrill~'~.~ie.E; for rco~niz~. ~~d iI'n~~I~~Il,It~~~ydrai£ .... Uni~(!r~IWCirN~w S~lLi.h depQ;;i~::;; biH '~IJi~lew~qu nibrium ~ discovery Qfhnr s~ri,r1lg~on th!;.i~;il.U~){lf ~wre m~f!;1!gl~.!\lJg YC~f!!ia ~~Q~re~'5 •. Wales Pre:>s, SVdmIY, sl:ate..~,. There is, lhrle doubr [11m .is a 1.'US.i:!, s:trip~m,im~ thdr go~d"rk;h OI:~S, ~o A:llt:linnugh :~]islJ1l,'Ose lacks the Aus'lr <110. Aus Sm,!lI5. c poetry rhat R3,chd Carson these and od~er 'clim.ue- iaduced F~eLdsofmJ"lmg ..nese trlOdluics on t1Ibyssal ISHN 91M8>6B404 lSB. ~)lla]ns.<Illcl:ci()b~.lt=:l'ich crusts on the side:.s of bl'oughi~: 10 her books: <I1~~LI.t clliiJIJrnges wi H affect deep-eeean ]U~. bur t[le manne rs lnwh ich :sc~~mgl!nt:5 i]llipire (:"lpuc:ity buildJill1g illl ootioiil marin~ li f:e 'It the edge efihe ~ !'lea. Koslow succeeds in :pai~:HinQ <I p:ir.:lu:!re effeets ''1'1 U be ,exp:re,:.;std~re: ne·<uly l~mposs,i~le M~les \'>fit~:l 111;me~ic :ots,wel~as ecenemie im.ler~ tl) p~d~.cl or I!) docl.Ilne-l:It bt:c,~us;e~M~ h;~vc c:sts~1l ~xp]'!li,ur~io~lof dcep-neean rm;OlUlC~. of the ~~cp S(::('! IlS an ~llviro~lIn~~M \vhh i,:~dle,ren:r;<J:nd~]jue.ltened 'l'al~'Ue. scant lllldersl"andil~~ on'nuw cleePFse., et:oQ!S:yS~ Ph<lmn1JCleUI;ica~ ,emnr:mll'ues prospectfbr new Tltl.;; ,~~I1I:~~lI"S ,erl.,'{!~nliO:'lisa. ~ propcmem of s l~nlL;;flpe~l~~ ~h~n~ r1ac:.e. \Vh:i~,1;; ~ t~o:t,"C dru.g~(I~HI ,geneHe re:l>nu.ro~s:i't1mt.i'Dg :l'€,~. A in w 1b'll~g,:e()Il:IiIl.£ {";nno:m, eaOi.lrn.obilrn;~,)f prID'i.d,eg iilrnCien~ deep-eeean oortl:S(,:'l'VmiOtl:l e:Wfolils ,<lIre .~mpeCl"i1I,~ 'lUb~ll aiuJl1do:~,txl tl.e view tlim:!l't eep-sea d ble, For a d~caclc, K,osn,O'W :~ed deep-water the orga:~.isms, sre el'(;quii,s.ilte~,y,~dapted, to <l stable t~v,e fDr COlln]l.e~c:u<l~:illlerf!.'its te experimern ,.'1 e i:!CO~g,gy pmgmm .]l~jhe AAli~tmlian C'QlI'I:i1l.norl!~ .md UUU,Vill:yi!'l,g en\'iron~~l~liIt;1; ~]<JI,\fenly a with unchecked \'5cheme~i~)~:'ertili;;::e ooeiHfI \\':t::'~~~lh Scklltlru:~,l1md I~:dustdal~ese~rch l1ilI.ad,C'SI unde'ffital1d ing ofp.hys~ologi.{,,!11 tolermeadows @f1d! to dll:'1rel.op deep-'lValc.r C'a:~bt."II1i, OI:g<1I:1iz:<1I;iu.lm,slmli,e!:; ojfbi!l)duw:r5itYilS:socn;~ in ances oforg<1:r1i:::;~,n:il and ~cologic ...II"eSpumil:~, 'I~~.e!:i.Co~,mn~:ci'J~\H~J.lnll'es. ML';i:I'l:SIi1Le~ WI~U eonsequesces on deep-sea bie.ned\'~~lh seamolllnl~ COIM] reefs lh,lt a:D'elluf'<I!t- ofpop~l(Qtio:lm:s and ecosystems, to c111i'll'lS)es ~In ~~avem,u;e~taill1, ened by the (:nii;lc'tts; or e(l!nml;:!"Ci~11 tf;lw:llhmg. ~i'~i p1Jr;:,]il'M~[~r~1ike~(,l'I"!r€'r.m~:!"(t. of'lxyg~n d iv.;'rsi~Yflool the ~u:S;~I.l,iiil,;)!hi:llhyor d~ep-(!c~~m C COIl~e:~1. iQlIr~emregimes, and flJiOod supp]y. pH. e~"O.~y&te:l1m. services, He has I;lld.:li been engaged in using om ~.i!nil(\d k!'!~\~;~!;;dgl;l ~~(1~lU'lie,cO$y~Wm~ H) of \~~ 'l/\'i,mdr.::r;u thit.'; :,;Inlwg~ ~l~ma~~cap~urod Ko~ln\'v closes Tht: Sflem Deep \'l.fit:fu a ~nl!hr:11III1.i~lliermUlQi.~~.lpo.ik'Y ..m a kess as II hey Ill1;:sc;:l:il>.IiC lsws tOo 1~~'~~1,age I:h~~~a:f> for' t:lJlcbenefit oflmnmnkiiml. The d~;ep OC(:UI'ii, ~pi~~I1J~~[lh;~ ~.f~r~~~l v,~~,~ ume gf I'li,t:hly ~nhabi,ted biosphere on our Th:e! Deep. Whe 'E:xlutlrdinary (rea~llIr,~fi; or I.ne Aib'Y:S/ii. (Iil i~e ~or(ilii.jl 1':1, Uf!i~ rs1~.y(ilf (I1'li(.illg(l P\Fess"(~i~ag'(I • p~~j!1~!.. :1<01':10\.\1 nill(.t~ .., ~Hp~.~Il~~f ~!~ be As :2007. 2S6 PiP .. $;'15. ISBN 97,802,2659'56.61). ..'rri'~kybu!'!i il)t.~~toO olfer inlhn:Ilf."fJ. M;iellict:: of snell ml t':x~11i\l1lJs.hi'e s:y:stem.,. 'VJ.'e .fui2li'll~ o:rrn]y the AbYSiS:es, riY~lfd, a ris. 2:QO,6.€'lO. P H;B~ 9782213.625137. b~llre~1skelel;on of tm::ie~i~,e km.w.'Iex.I!~ Rud 1nl in(.'ompi.c~elilludel:stand ing of 00lffil.ponellls" Weirrllliliess and llea'lJ~y are ,oU.ellil paireain the 170 SP@(i@iS port'Fol!y@d '~ra ~I'!i S'lfi~lHI~ S l!rv,ey 0 r d@@p-Si@.a. prm:;es.'ie:ii., ilI'l.d dJy.rnnu:i!c~ im, deep-s.1:i1 :sy~1-e:U1L1S. (jive,r~il}i. NOI,nrialt:!1 ~ro\l'ide~ lh~ ~,(Dlo:r ,ho~o's. m:@I§'t p Dc;:ep~s.e.. sc:.~em:li.sots \,\\'ork in :!In C'ruvli!:mIllJ'liI,eJllt taken by rl!ls@~n::n'L?r~. Hhi !!)';t@!ndl@~(:ap;liio~'~ th1l't w wll(jr,~~~I;hicl!f~ of ~fforf l~&bi!~ll. sLUb:~Ji!~rsi bX~ dlesc ~ijbE, the Qlflil1'l.;l~~. I.ih~s.~yiloe~nd h.;l:bita~s.. For a orliel:liLered \/I,;:;hicl,e 0<11'1 >cOSI nrn,a.re thaIl examplE" :the~\~immnJlj,g :sea (li(lHlllb:e~ f~yp'f}i(l5U!'S SSrJ.OOO. AHhnt.gll l..\i£ h!~"''e b!.i('om~ i~llld~lfftt <III: ext:mcl~:ng the :IIIrU;SIu:n[OI'Tl1ii3tiQIl pos1>ubh: !lxi/nil] "lin du.~ale5 ~Inwly 1IIl1ldgrac.ef I!IHy th:r owJ!11 fhe wat~~, wm~lii1ll~:S ~~;n rat~'~,F h,om thE :boU(i!i'i." raJ Shurl tapical t"s:says.hum em; nen~ de~p-sei;l b;(lb)9i5l~ 'f$J,e re~e~r ,tis ,in tJfJ]e Di... ~~~"lo,f Ma fhf!e!''5c~t,fl((l, .:II" d ~ (inclt!ding IKQ\s~ow,!:.Iodi 1/@!IiI OQ'!,''Ilf) C,Oollt:f I",~li!~t!l C,g'.n~~ii'I,Ia~i~~,. T,(!h !~~~~hg(l! Qflbe el'l'o'i~Qi!lrn~.n~ ~'r1)(j Nc ii~lM!d'5of marine sdtnt:e., hi$~Qry. and cQfl5.ervillion. E,artlil SdeMe$.rllli~e IiJ:nhre~slty, J!3!S IDu~ MiI!rliUiI1 tab,

Fralgille! Flronliier':s iin the ,Aby:s,s

diver.;;iry of Ii Ii(: dwdlilrlgun 1]1.<: co~d mlillds of dUo: M."<tnmlr; and ~:,"e~£lbrnl~~:1!ae lJe';Kny of deep-

Rood" lBe~ufQr!;N(2BS16. ILIIS:,A.. 'fl1~i~ d1l3@Jh:l~.ecru. E


<'fh~ \iVili'1WOV'\i@.Jd!."


\irdd~ d.(J!~of I~~J;:gl()i~I. deep ooo<lmlin !la[~orm] and
n~.cmmlm~r~ l:i.1~m;;~i~1it ~~a~h.we the :pt.Jtt1nli.~1 ies ofcxerlillg stlbst3.Diltia:l1 .~mPlft;Ctj;on the ql~La.]ity of deepm:e<llll ~'C(}syi'ii~ems: there is only <I br:~efwi n!dQ\v of Oll,'fu:Hlnilty fCliiKQ:u.in~ poUe)' iaplaee hne:r.:mwuion:a] W<lters. We, llire omhe C1,ISPfellgilging o

A roruWlfurd-'~:llinkhlg !;l,pp'iCt)~chhas Immense

eKpell't ol'llin~em<l~ ttmudoceallgffilell"n~nce. ~Ild Crnig Smnl:h. II

GjfiJ'de. arespecsed

:e1.egmii.. ~iglg ami ~JllIpl.em~.I~Jtung

p.f(.~mi'ml~mt deep-sea [biiologiM, Tlle"ut'ho~
Om! policy

'pohcy wildem

~~h':I.'r.~~nd;fl[ e:iiri

eapiral isinves;~:cd reseurees consumed, <JndIwbiH'lts




desIr'O)'~d. Ko:;;~ml!J' provil(t~~s wH~].[I repanen u

~ltEt~I[leSS~]] re~~l!i! :~~'ed by

:h il~h~se.~sb oman

,we eurrera .\l~~nn~ onlu:'

de'p~~ll:s. nw is N

tn"w] i:ung.i:l'i!.cluding ~.g~~) i :!I'iI~m:a~ori 'ln~ ~ ~t~n d~p·loi!e,ati\"OiiiWli~g:. ~'g.l~!~li~]!l d~me:r$al ,of rlis,b.e~es. <lr.ldesl<lMi:sh menl. of marine plll'O~ teeted "lre~:;;.They bd!i:l1y tack I~ t~e bro~ld!;;r ~m,.lfcy needs for s\lsl:aillilbl.e use ~nd conserva-

I~(: li!~~~ttl chal·I.~.pathHj'\1!Iii!:rd~tiOl~i c!()~ ..%':r-

b\;l:fyreh~bim~:;. ~ oo!r!lp'i'l.,mi~~cl.Sueh pi.'iIIey, ~,nte:(,'.m<lil~,on:E1lly spnnsered 3111.duU1lte'm3t:~0I1"! iu :i.C(lfI[;J,c'~!l~ holistic and prnt;m~:ti(lg:l<lry, t"d!~er thalli, a :m:ea'!Jt:~oll 10 eny.urou,n.e'l:It~1 CiI~,,·slrophe.

Vlltio'n. s~~[l;:~i.cs ~mdmIS'lairu!bll;n...~liWL;: uses In.lit <tc::kJllll\\I:~ed~e 3ccommoo~l;elmt.e llliIlOlny and ~\'P'.1O ilJ1 n"ri.nldel]!;~!rldjng pfthe deep (lWMt
1~Jn.'6~et~!'I~!i\,1:J;.il ~46!ii1


B IOLOGI¥ re,u.ling. Tile Dvubl(JHdj;~: {,2').1Iilu:1J11"! ~Mre :r.~lts

b(;l1.\"'~~~!1'1~.dW~thi~ I~bs.g()~~P, uml dilf~ruw[ ~
cuhlm.res.but there are' also lI:m1ow.ledg,c arlul. e.N.hi~,amt~iJlg pfO£l'eSS.

iced t~p by F'o.·'(s. wil: iUild SW[)"!trm~.l£. ti:l;~. sbe glv(:!; pnthyb~ll cfli;lc~hl'l;: . made ~l clear that he \'IrotS~~otilU '\lVll~"'1i!'n.~~g h!lv,~ orig~~l :~>1·e-m Cell t:;'wu,!gr~~'ii~!l!JfCh ~l;u~n"!l OU]; ~~llbrY(]llUC stem .~~Hy b(;:~~,~'heelanex ~)r ei'O.p];l~i~lklr!~ Wlrt110Ut b;tumi~ d .!Iv .cyu#riif roJr cells, Thm August he ~.!IIr:d o:Wm,;<:eS8. jn loo.~ the nov\,': for seientlsts, the book is the book oceurred WhCl:l NOf!Of!, New V()r~, 2007. eral n.i!ilru :Ibr reseereh Uri ~~II lev;.' [Im.O\V a smooih read becanse FO:.\ oes bura Hw.ag:'!gbecame th~n!1!l 10 d 5~:BIl p_ $25.95. eS:l'!i50. s~d)OI)tlml! I) Iim.'!:£ of ~~:u ml~il,llbl'YOIlUC stein m jJ>lII:bUsUn, ,voil cli!I.h~~illg the Illiot dmnb d{)Wll~ scient i flic~ermh LS6 N '9700393ll5anl. cells, Hi-; action not un!}! caused steL'il1L ~o nO~:(1!~.:y. The kfi(l,wh.>d,ge she <JlCce:~~ genemtio!l of <1. li!u:m:t1!11 em~ryon~lcs~emq,;eU I iae frUI1:1. became ~t uati enal I~o~it~c:.d issue bm:.Jls!J) q:u ired :~nher jO!illJnejl\'ii!>":f.1mli~1~ cmb('lkl~iIII~\(~ OPlli!llf)1 th~t \\I;l!!llOO U) (:;011111'" any a c'I!):n~dM~"'I~y~t J)l, SO'lnal Korea seemed { iIlg: i:l'l! I1Uli,g:C .~ nd &Ill~,~ she i;: i[oes Il~p~n> 1l:~H~, pe~e~~'ith~~le Uuitesl St:ne~~in lhms research, pOiSleli. Hl!;vi III the racejn lhef"ol.pE:Ulac: t:lonin£. fr{Jl.n~.1i1.e~]IJ.in~ Iil:ernlure "'S n.llligs OJ.1the ladT~,~glob!l~r~~~lQ ~swlhU~h dflm.i~;:H~C'~ in a d·i.;iit l]~~r OV~r'!,I i~v. (1l1~ hI;lOki!.1dud~,4) t s but thL;:tall<; oo-eflI~e:a G~'(lk lr.:t,~OO;y llj:' hubris fi,eM wi Iiill euermeus scientifia, he<lil.lh. and jJiilI.,!!;elll ofiLif:licll,-;enoes: O1l111d imcf'II.iiew Ilotes.)i "'I~d,dQ~'!'nri~,t lies, :It~,\ s as if iil~the space raee a I~m" ere ..tes i.ncl!e:Jrilb:le'ilmlg_m;,. Her :ff]y~ml- Neil Amls~llOIlghad faked l~e' Moo~, land.ungs. ~mnmel'cia I poss.ilbi lities ~\r<'lS. •.iuLt. \~uth us. on the--Wldl deS\,'rip~~,{)n{)~,Ik.idney!:I'ml~'P',I'~IiI~ m!Ld I~\",<mg'~~ii1~.ing.~ri.tU¥l!p~ mllf:iJveled :;klh,~,:ly'Mt '<In icip.lllt~. deni.ed li:dem~l1,tllidimtg. llilJ C,~!I jTdls. eYUlthi~ F'O;\ bJiillgS he:::v o firsllL!:}d ever flllsl.or as IH1l11.yo:Fhi s cl~lliI:ns~.~ chi.[lm~ria stem O~~~ opcrol.ioll at Mas5i~cl'n:l,lreu:s iIfl1PJies:-;i.\'e~dellll: as OJ. science ,\l~:iter .md jimw .. aeni::I.·<tIHosPiit a ~is .rivettUl.g. as. is the ahnest !IIH.tennirlm.ed .. Bec.u.lI:se Foi'i. '1\'~(Jte the oook be:t'w;eJt::'.Dn, 2003 .,nmd 2ml,}. ~1~.·e P'ffi1bah~yhad to smd~ aeeounr ofe. ..... u·ac~jhlg OQ(.'yl~S. fO:If tissue Il:tl~:i'sl:o tell ~D1glhe s'tary o fl.hLS race. The heifl)~ t gl"'l~LCkimd u(!rj "llwe~~~ m" or ·\'1p~rei1l1If· dOjilli~g from p~~. In J~r!)' Y;lng'~h~b.she \'\,'itoook olfie!:'!<\.'i: .!:tI'~lW!read r{)r"'rliYlm~ i[l~~~~n~d ; .i:~.d]elopic. Fox nmlresl[l.e s~OJ'Y<llladvemHre. to ,e'V,el:"Y reference to Hwang's resuhs and nesses t~.l.eStm: W~]rs~lli.ke diil<'l.nl~ ren:lOte.ly to l Sh~ e..r.Il~$us lf1 un]ilh"ly pl.a~~". [.u;:gil1ilmdng til)rltnl~]iug 1]~Pl;;f~~ (i·~l~dti.j;~l~n~Dcytes f{lii lil~'~ cre~t~ <!Dl.e\.~·aJl;~ ehapter, "The fa]l~,f with a c<lllle.~ rude ~o the .fymmids wit~, an Sl:!mLI 211i1.dl~~e~:ise of S~.ul.Fril!~.ci:sco." She sOIll3t:i.c ll:!eUnuc.lealr ullInsfer. Sbe t.ells the de~ Egypli:f~n S1:cm ce'~lIrlese~iI',r;::he:r. Sll{l'[iten HiCE: [0 writ~s. ·'I.hc Woo :SuI\. l['I.:~'.ang f!i'l.'ludi!i: tbe ~·e~u~~:t()ri:csofpI!Ji,ell,t:t::. w.il:b he~~rl r~ilur~·. hll<lel to meetlbie sciellli:S:llwi·th whom I:~.e .uLwi D'liUllume t1ii!lili::tlllie.. kid ney ff~:il~,mi'e.. <i:ncl b igge·sti III sciell;ce lll.i '~ory :~nterms of ·the s Egypli~n '!iWt~lt~-dit~ eol~bt)~l~ (ptllilkR 1U~.d~ D~eh~~l~l~~ dy¥;lrt1ph)l_ Sh~' ~1l'lO !l~Lmb~!i ~u~~!y !l!;'liU'>:" of p \W1m~afth0 il :impooslble J.I$[<,~cl's Joseph hs,ko~(it;~ ..1IE Idlo.r t~]P of mle·tJI1ica~. lIIr.1,S'C~enli steill ,ce~I tTeat~ • file BlIsh has t"\I:i~"C \'eI~oedcOJ~gressi!oIli<lll bills provid~d fl[J~r(lOr tl~ Ih!t;lli~¢S ~Q Jf!~~iG Tho!Vi!.... n'L;i~~"i!lI~"'ii~'J1Ii!s "u~h f~:O;MO.l;QffiV. Ui;~in\l'. ~fl i!~c!l;~\'1~~'d@d limd:~ng f for htun~~ ~;i'mbrySOl'U fori:~ir ~mi~~~ p,lpt;:r un thi:: figst lllqli1.~.,n un~~th!:l C<.!l[b~.m. onie !'i~Ili.~ cell rni"-"iut:h. Cell (~rCHl'iinLl;jllfllll~l'i 111~lsis nat a book 10 be .read '!,\','hiL~e m~dli- the C()i~:;;equC'niCCSlor ~Job!II1>Qi.Gm;:e,sImes that cl.nbryonLt: stern oells 0).


C,e III ,t:I~ Cen;'ll h~'Jfijpres.itlent Gro[f'geW. Bu~h took oll:ioe ill) J<JiIl1.U1lI1)i 200 I. he qtq.iddy issues
Itha~ :>'t~~l1Ieell

Cell ofC(1U\' isa


.. 1='01'lfio:r'iscieu-

TI1'8 Global Race til} Capture a:I1(1 COI1tr·(II ~he

j;e'rl~ClI;II:l>book, sp




journalist's eye.lbx

!.h::laih he.r

n~:!iikilJi,g. flli...... explaira'scO:!~1:p~e:x concep13 OQI'Ld
f1Il1mCIX)lIS P~,flICCS

Fund thek O\.~'nresearcheni.


dl.e dm:ihed

inten1'ic,wt':cS:' olli.oes. II~b!5l,nllIll1t:risIllS. ~Ild t hi.lbi.t $.-ew;::Ill the \'~.ews lhey :see c<!,d.. &l)r. llfillme d:el.';li~$" illmlp()s... ~,b~eliO oblTIiu ff(n~l a sc.:i~ I~miie p1.l1)er; JtiI~:~~ th'\;;r~arthi.;ns cll'lln~.~i!~i\"G" f O Jovi.n~ OI~~O S:ullg,lp-ore 1:0 d~ribe steml ce~.I1 wor'kV!1Iiliit;! b'fii.iish ru$~IH'ehc!lY gfBhJfl~U5 amI d'n.en on lO Austrnii.1l, .I~p<lil~ o.lu~m. a:mid Km'!el!l .• .• F(lI;; m;:cun1t.dY n..'VC:::21~S t~lC s.rn;:i.oIQ~~cal. ~uld
Tihlll:lr'l'i~Mlfhi!t~il~ G~d~',,'e In~i!~~.S;5w.r~~i1I I KIId O(iy.e. OliremOi!ll,A '9'1711, tl'!iIl E ·mali~~ilflJ!hi'lll~. ( k!i.:edlu



There <lIre p:]emyof Itmiiil ch<l!("tIJ(;ter:li~in~ chulullg mrv WeiSSm<lllm.. Ronl NIt:K<:iiy.S~~mO'1iI1 i ShlWill. }\h:tl'll C(l]m~!l. [an Wilmut Sl~"I,Ie M:iln~r. Wi:Se YOllng_, I[)oug Melton. Mrnl.le:mdra Rat), ~!!ld d.lel'lltj,\"disV.lCl.~ WtlO Suk 1H\il.F.!I'!SIl.nd some otppe.n.rirel~eaH:(Uy.

hopes of dlOse \:\1110Il~cd potC:::~lillillru:·."IAl1lellffi. r~xel!Dquel1ltly chrolUides the cm1sequeooes of
Lhi s isobtiorn s~ JiXllicy ;[~d sq[l.iiiIre~y i!,dJvoc:tl~ejj; 1l ~tiomd a,proaoeh~(l ~~!nd~ng f(;n~(;l~reh oolh o.~ lIdu!t ~mdembry<li~,ic ~temil.ll'elk 1. J.A. IIi1cl1'!~!'! ~~.t;I1,. S.!fn!!'e,2sa" n~s U.w.Bh :t J. 0.. 'iii'.aI~Qm, '!tl~ t100bJt fMIx: A .~MM'A(iC~ of Ill;! £)~-"'OIIfj)'(t/l[_h~j1.('Il\I.l:~ qjMA '(AlI!~~I.iLm.. rrlewYoIt, 3. 'th,. S. Hwai'l~1 t!l'l.. 5tl~nt~308, l1n' ~2.000l, tl


lhe: ~1,I.thor!s 1:~~i.ni21tioll Wlfh "'se~encc:
tmum;;il1ig sciellice f~Cli()ll.'" the pO~lni .. ~ of

s~el]''ce:lls, ..;Ilnd 0 l~rdCi>1~ to .~!$li1m\vh~thappens W!1'l.l: makcrh~~ . .a n.lI!J"!l: '<tn'[-piU!l~ il'~dgwrll sd.oencl> book, [t oomii1ldsme I)f~h-e Ibn oHin;l



SIJ!>tai iI!li3ibi~it.y ~';)IJ,elj ill!!]1~Ilteg r.atedi nt!o hig1her·,~dl!(~,tiio nif1 SitiluliiO.rl:S.' lIIliSStOifJ a nd

p!la!llliliil'Q. wrrnwla, .ami ope rat1l:.ns;.

JeSJ~'W. stu:d~nt lH~•

U[ilI~ii'i1]bnit)i lsa I~n~ dU:'Ol!ghwhic-h g.m:re<kli~:ng numlber:; of 1.[J;(Iividwd c,o~.~ I ~.f£:lg~1Ind un~~".t;ir. ...tlt::;:, O'!:; '!N~ i tmtioril,i~: Qrg,ilI[lizlu.iO:I,lS. are ccJm.ecti ve]}' cXlIIlliini.lllg and



:;;h,l[ll:d wurild :i.)ffitt1m~ ( )" ,].~. In the Un]lI:d St';;l~es,• t mniomd trend has I:le£un. .
"eli ug UpUrlI O\!I:

bm much mo~ 11eedls In be (Joliile, C!!lrueg~'iI!lId iUI!!li!re!'s:ilty AcUIi!!ills: Susraiaa bi~,i~)I lis be:i ~i:rue~,mti'!cl i:ll,lQ U.S. htl,1.itmi.olls· mi:ssion ~uu! Il'I:H:I~n~.ing, CUlTi.ClIifl.

l(tiviid~s (If ,Natililnall O,rgiillili,~~tjl@~~ Allerl]mcU.t1i lted Ni).tIons d!edm:ed a [),eC:i1Ldf: dent Ib~s ,and ehangi i]g(;l~e~ p(J:~~.c:ies to l1\1enl:snnd"."'en action 'to nlOiv'l;: ~~]\,"'tu:d ()f I!dtLCi1dolli fur S~151;<1]m.hie Deve.~opn,wnlt "~lr1xJI1I mow' '!:o renewable seurees, 1ilIe'~I~:ml.ityBliiid to e~:fm]n~liC greenhoase gillS, (:2005-14~. fI grassmets el1;cm From [higher B US. business. <!'U'C~li~ec'ntre. ep"'iooerr:~ emissions, Michigal1lJ Stille Un~v'elrSily. NYU i.llld e.dJ~lcati()[1 developed in u1.e Uruled Stil~'S int the .!5 ~ '!ij: ~J~ sehoelsare ill the fOJ.ermnt of susta illmi!lb:il~'~ (New Yo:rk 'Univer.;~lyt Un~l\rel:sli:l:y of Cali<!.osellOf:ofa fedel7"dl g~vmln.ne.m response'. The <lim) ~h(,l i~y!;;duca~ion. Ar,eh]l~eU!re and ~~,g~~.C'~dng oon~ia at :Bi;:rk:c~~)'.dl~' '~!1r1s;ylva~i;l S:ta~~ NatiU!~~~~ GOM~en fur SC!i~J1i!(:'iI;l '~ s,chool!; ha\le' C'I'~leri;E1reI" ~cr.ed itBlli,O'Dl. lhm Unlverrsi.ty,lnui ,oLi:Lel'Sil<I:Ye cOirn<1lm:ted sust<li~,~ En'IJi:mIm.~lf!'IliI:~~al:aed.I:S<IInrlU1d,confcrel.lCC1!~ll ~ i 01. ~ reqt~'ir'l-l sti.J!di.;ln~s to be; f~bl~ Hl l!l'Id~fjU~~ld ~~bi ~nl!d ~'l~,d I~ty h report~. S~~Sl~l! Hi ty- :~lIIm ('II'! Bdt~em~{([![f~)r Sm,,~~hKlb1e ;)r!ld i!l~1b :~ ~ ~ o[iemed rilS.~de:l'llli.,]11ivinl£ Ilru{;ti~'f's, ru:e ill~p:l<u;.;l SeCUll.'efmul"e. Om ,[jnt:l!£lilm.e.el~Im:g,. the Us., ~~ !Iud .~mp]enm,clu slbgt~ilil~OIb]e iJes~gl1. NOIli~ <f profi,i, m.'gI'IIlli:rnl~!oI1JS$Ud1. OI:!; Engjnoernlor ~ ~lt Bowdoin. CnQ:!I1~gic Md[on, Dilrlmo~li'h, P~I'tni;!~:iihip For Edt~ci!~ion f,~r Sust9:ill<;llbk~ .~ ~,~larv OOl, 'fillfts. U n~.v,e~:::lity If \k[II,~lIOrlit. <In(ii Devek,pmellll(~) \\,'.15 Cl)u,ted to catill,IYle a U.S,,, .. 0 Va:~~, R:;(l!gers~rtd~l~ Na.t!ouno;!if\ssocial.iollii, f.:w resfJ'O(l!J.~ JQw~ll.is d~~~de ;;ul~1'oo¥ond. I'll is ,I'" Prg'¢l,d~;rnt !h~ IlL'~. P~r~nl1l~hlp fm ~.dl[l:~~;!iooh!i' 'Qif Ecl:m::<ll:i011011Pilrocu:lJlemenl ~,l~,v,e fmm5'ed mil 1I~<1Ji(Jn~I~: [1~.et"HJ:rn of over 300 o;~~llIJizatioml!lll3s ,~ Slmal!'!i!llb'te Oe...~~C!pmlliln!. wa~h~IiI[1I.tK!, OC 21)~l1,USA;

r~ellJdl . .'5tu:td'em nl~.pe~niml:-J ~md pUliTCh.,::,i~ o [~l!h ~nd cOlilunlJ11~ily IUlrlner:>hip~.. Sl~l,di;:t'IJb' <lmulstaff at hnncl1red:s ~f c''Ulnp~il ses ii1fe euiWJ:ged i~ Sl~s,t~i!lab~Wy CU!1l1!1ll ittees ~I!ld ~cti.O~lS, i.m;:bJin~ dl~ foUuwiJ~g; l:.f!ming t~lj:o~m~ en m;:qu~ring sus.t!l il~bi Ihy kD,lo~"ledge and '1ppln:~ cason s:'k m~:, i!!laid! ity-uri,ellltW l1~ml\e~g· SU'~H!ill[.bl~e\'lo.~d and Engineer!' \\lht\lOl!i SUM!) Re'~ vals speakers, and Oailf:'r CmUpLD:5e'\fent»~ Borde['~ 1m",!!' developed, T~I.'eWorld si{)C~ally "!IJ!dt:mfinJl:lmel:1mUy respon$[blc cri- SOlIOCes lrl~i'Mte and the A~e[l IIDliSi:iM~ lrna,,"e leriafl[lrp[~rdl"!'li.llg and endcwmeais; .~ Ili~n~ ~)!,'(IIiked with business schools to develop case siQl~ 01" S'Usta~!l~bi.~~lli' !ll1Otlh.;: &~'be~ ~~h,I(.';]i~ ~wdi~ ~ml bU:SUIl(;&!;:, eurrteula that include ~ lion core requ ireJru.lem·s, CCmfSes.. dI:~sc:iip~iliI.es.. su 5i:al.ilrnabi]li.typ:riDl,ci~)~!es OiIncllpractices (5). "\!I~ok;colk;~~s.. and sp0Ci~llijl~)~d~,~~~t'~ [uctv.f""ing]y. i~'[e:rd~~i;pU~~ry ]w:r!1litmgexped and reg:uo.rnii'll and .~ljJb<l1 <li1~proHc1n,eso sust eienees fOCLIS 01'1 our sUS:~<L~mllbi.~i.ly dil"'~I~e~lge;s. ta:inalhU ily iI1coUi<I oor~]tiOll\'i1ilh businesses •. The p'Uwhm;:i.llJg110wcr aloae o:W ooUeges government, tl!(ln:gQ\'eilTlmenl;a~ org~l1ii~.tiom; <lnd m1ii!.il;1l'l'liti'es.• as they demn;p'gd more eavil (NGOs),.,<lIld kUlldelrganenlhm!i!~il high seheel roo!]lel:ltal.ly ;fLllDd :sm~u~~ly re:;pQl!lsible:pmdm::ls ~ K-~2)edliJlC,niOl:l, .md ~:l\rooe!i:s-es. com helpm:ove :-JlI5tailJl1tbi,lily from its p(re.::;e'mu~ nidl.e mm:kel::!I:O beco:~m:' the Core :roqWlilireme:lli~s<n many universities lH1!~d~~rdnn pmdi:tc[~nd pr{1t,."'>et;;~ i.gn- This d~.. aad co]~eg~ te.g; ir~lrlliil)d S!m:e Uni\1~ffiity,. lVIii'lmi D~lde (~omnmni~y ('o]~eg;e •.U:universuty C'-'.1~be expressed tllWu~ilI. {;o.mm:iune:t1i.t!>1itJSU;i-" ~~fMiml~"!\{lt.'!} il!!ch~~~U~.!;l cU'1!1!~l!ITl~IU~~lf ~., ~a~I'E.lb~ :-;t! b~lliavinr& .lill!] policles in ~n:':fiMi(lJfaal tEllinalh.i]ily edUmnlim1t, ,evel:l. ifdu~~ \m.m :i>1IJ,st.,irlIli1I~,s:siot1i1 plilll'lilliiua,g; mare' e~ua.el'gy-dfic iem <lend 9hi~ity~s.~ot~l'edHcany ~l!iled.![)~greil:Sin S~llS~ ~I!ld g~~l~]" b1,.aild.iI!lgS Hind l'll'~ralitl~l.!l:; $UbSI'Bilil,. tllin<llhi]i.I)' h<lvt' sproured IIp;rut dflzens;of~n~:i.~ ~Iii!l~pnrehases and in::~Hlll'll-UOrlS, of renew .. ble .. m>ergk;!> and ~1;l~-WiUi[lnt:nr::i ~ilrb[)ijemisto ttH.i[m~ [see (31 fur~. ]~s~iijg].~n [hI.;: ('iI·J1I:iPUS. ~ sious [-edltC;~'ioalS ilDl.d lIl.etll:ndity! :ms~<1il'l1Ibilji~ Clinl"~I:' Clml1en£,e. !'ii11Llde"ni~s011 over 4Q{h:am· Pli!S~:; ~re~,~1)'rk~.ngwith 1!d.!m~i~is.ir~h)rs·1Ind ~ltidits ~!)drepmting:nnd ~lJs'~~n~b~~ Hv:j,ng ~udJ l~uale~H)~~1i'e~'IIIl!d redI~ll;:e g:~enhOl~:5e g:m:i C<Jl11~lui~1\"il'llhe wsidenli~~i!!<ilns. For I:.x amemlssleas (4) ami MC VOI.lDni~rUy rill,ising sruple, aver ::)OU' prcsi.del:us have signed ,commi~:~

of suslr.i~<rlb:iUry (6, i'), SU1~ford Unh;~rsilty


d cve~IOJIDd both l::t1iIVU.I~'o:t1i1lil~!ellt!lI<lnd ial soc screens f(lrnlil.e~r el'K!l}\,vmem'S. A st:I~:em~~n d:ll<lfted by [he B~!lSi:lme!iSecl:or S Tesm Ordl~' US. Pal!'lt1i(,!'Slhip'~115 up!D!lMglh~ edut:<1Ii:O.1iI1 make SWlst,l]nab iI i~y educat ien to are(I~lir(;:l'l~.~!'l~ .~n fiJI' u!lde~:mduAh~s- Patlici~C),i:1til1l.g embers m

eZln,1.1f'from bothi sma Illmil,d l

hlJfge co.lI"p't:nl'<]tiolJ1S -from
;,t~~ '~Q e'1iI,~}'


such as Dll~ Energy

'media. congillQi~~.er-



j:looducts such as :S'l.il'i; Bees.,

i~te'rdiS!: iplilii<tl)' apJl(Iiil}<u::h. no~ olmly the speciiiriJC:.J:l nf nur ~ilIS'[~ im~bi~i~)' c~.nlit~nge..~ the. pOSl'ib~e and solltnli,ons. butalso lin: it1it:erpC::mln:~1 s.ki],1:s, the
~:yM~~1I1~ thil'lki:[!!g and rhe chmlge ~~geliat skills to e£'femive1y help t.ocl1e<ll~e a ]1:]ore sus~aill!!1blc fumre. '\~e aile lo[):I.;.il1lg these sus~<lillj~bJ;!'ty edut: •.ned ~m1(Jel.m.asllq:~lu;e bllS~Ile'!iS i

''AJl!5~;~q~l;enb eed to le,nn. duuugh n






\:mpl.U)I~i;:~•.a!>OOllSilim~~,hm~]\IIdUllvesliOI'!;" (I!G).






t:I E·UI1Oli~ n(g~~~lalflld1cudu.

dle\~~I,ori!lgreWl'lre(;)-,> for dl\l purch~"il1g sid!$

S~B)r~~!):!,l;]S i~f,ni~,. B~~i!l~$, C{}!1'l~mU!l~rles.


H i,gherEd~lcadG.'m. K-~:2 scjllJ(lI hlg. ,;U"Hl ~'h~~l~. Tillie Ill.S. P<ullte[$h~,p It::OJIVl'l'r.teSnlJi.'1~llSLre;un !c~'lJdeI"5!I.lmd Ica.tl:'llyzes lleir OOmm]l~lc!ilt to edl~~.Jitirlg a Sllt~t'I~I1l.~bre future. Wi~~l ~l1r:1~ for pews, fiom Iml~li:ple sautees ~nlC~5T~d ilm SlJS~ ~:JiIl!ii~bll~.ity.three: mOljor en:i~rts:,emer<.;J,ed irill the II~ighcr-~duJ,;~UQf]'fl "t;c'!or; the Higher Educ=:ui~ll1! M;';~'idati(i!lJi;. Sl~st!li!'i"bi~ity COil5iO:it illm (:~[EA.sCJ, the Discipll.iuury Associations Ner\'\,,\]['k f~~rSUM;nirl<lbi lity (DANS).ilnJit~eA",,~o~I.iom,for the Ad'~Ulie~l.lelfiilo:fSnsl:aflll:lll.bi !.tty i:1iI1 I HigherBdm ..~lign(AASJ1E). Thel~,~EASC I( lO~was fanned te a(hr;;~llict: 5o'1,lsmilm;;lbHit}l in the mflil1s'u~alll fuli~~€iedllc~l~ hOIl assoc iatiom: :.:J:lildiu tftle sy:Slelll.1. of ~,ligher .M!1I:llj,Jilg! p!(U'w!'u'tdl

K-12 level {2m: and thL;;~ob(l1


~or reail pIiOj;,lre:ss.. the f.:[I~pie'.DllenlJtll.tial] lms to be hl.'OOd 'lacf{X~ all higher t:::d)~lc~t.~on ins! itulin~n.)a~.ld [horou,g.1rn. We need to !ll1IlI<llke MUI'e thm Ilont:: o:W'I.he courses C'tUrrelll~y bll!l.n,g U.llJlg:~lt in the l.Jn1I.ited St,.u~!i reflectlhe' old. limlCl'll.llr.lte pa!l"~di.gm~ such 5r." "cndlt\:l)s r,(l:~t~uD"C~" 3~d "~l~Ul COl':iqu~Ir$I1!,\'iTJilre; T;e~tboob need to, descr.ibe m1l.f slIsl,rimlbi lit)" cill2li~.I,ellges,<lind the ~,:~m~:Ii,b~~~in1!il~" dir::;cirlil'Y!;1;CiHIl m.d\:~ l(ll thi,;: ~.~C:h sollmio:l1l!s;, :Fm;KI.ers hW'\fe U-" ~lIPPQ!it such VI'Ork,

rew!I.I.I'Ces,plrooLJI.a:dl by F<l()i~~~leFlItlire (21). i.lm.dud~ngr K:-I. 2: cllr.n'icl~]!l .,;:onumllliity service • ~LCI:ivi~ ies.aad teacher prep<l!r:i!ti:tm pr{)gltdll,'1:~



Scienoil: FmJulId!.rriun

its grnllJ~;s

i::ilm.i[Ourugea slls~~lilmb]l!i'ty

~o STEM l~projClCls; ~O~IlCE3S~ S'tlldellilts'illter,w est iJ~ s(;jence.~:ech[loloSy.. el'lgi~l:eering. ~:D:Id l,.'!!~tK·'Rion hs~lf_ HEf\SC ffi(tmb(;;!"S (~Lqfoo!Uly mmh~m;ui(:~) .-;!ndothsr <i!oo~!s~!ld s.h!)~I.diil[uJ I~;epre:se~[abeut hlllf"ofthe US. crClileg.~ <IIn.d intf'l'di S!l::.~plilmOlry ~oLJI.r.se\\l'or-kOlilmd resec<u~1iI, Other g.{w>erl1l'n;leru!~I.fll.ulding SOIUlIr"Ce.,>. Ibll:~~d:lli~ mlivcrSi~ly lClI'eslidcnLll; about h .. Jf aJ~ll.of the' tiens, otiild eurporations rh::m lQ~ii1der~nn:uJ ~O<l!rdlii of trusrees .. 'U'ldn~1Ia1y. iflil(J[ mOld. lraeililies dlrecters, bll~~nCi>S: olr~cers" ,t:oIJt;':ge nnc;l &~l.PllQrt this lrn:~ldl. mid lufiri\1ersit), pl<llllieilflii; purch ..sers; and ~tlrfl1; Thxo~gh siltlr:iillg st{lries of~U!v people have ofres;i!d,t'ntia~ Im~l>h~,!£. :[;tltd~III a:ll~i!'$. ~md !n~:t: a, dil~~~I1~ in sot,'!tety ~iQd by JO'FOv~di!lg th~libc~ls Y!1 !$Olvi"~'S~l~flust;)in~ caiim~PW';)'~~~""]li~s, rooj~'CtsindWlA[~the .Fligh~r il.'\.."Igr.!rffi1;'lj.U:i. Ed:~l(:aJtiQJ~ O:i.m>1lil:e .Aoeliol:l :r:al"~I:lCrShiLp (.1.1 j!i!J ..ibn':ly issues, educatorscan ,e~l£i ges,ruli.elllt'S a ..nd help k!S.1~lWi()ru;, end d'!eh:nger stl!Cieuy uun measure <lmJ reduce greenhouse gfl~ endsIOM~d more ~Ll:i>ta.imlblc be i1,lViorn I <Illui po~icy sions, SIJPPOl.lror tile Amer ]c~,111 Colll~ege and IJlliLivcr:sjty f're:5idel'll~:S' CI inurtie Com:mtlli!:J'oeru L'!O:li':I1l'nS,.Si~uden:1S ean ~ea!11!l1 <Ll1:cli pmcti,(~ 'I,fi.~such

Hewevcr, state standards and assessment~ pri 111m mil), 1:l1fl1'l1]]l<.i ze wri[il1l.g" ,retldins.., ' ~nd math, on~'!ldonO[ relate to Slod~tal pfoblel1li!s end :;o:hlftioil:-J.muJ C!'t;;!U", ~r'lie!'8 10 le.. rn i~,lg ahout sustai 1l13,billiy, The K-~, 2: t sector l~illi1l ofthe U.S, ~nlfler~hip h,l::! ereated (h:a,l~.su staina bi:1 i ly ~d:uc.uion sta 1'1(ilo~rd~, as ~Ompl!~.l;ld h resou rces for K~ ~.2: 1:e1U;;~.ers. ..d~~<'Is.jmil begwl,ll aprecess simi:m I~r 1:0 whil~ \,W~ done ilm, higllD.;:r education lTD

eeuvene the ~l;[~~mm~, leaders

:u 1:11

K-I.1 edue 3~

and to share ~nfoml,('!liM



w slls[ain:aabi~i.t.y.

slJ£t!l.illJilI:b:ility is treated by 'ldd~(Nt .ts~mofheri[€,'i[LlI ·1111 U]1 H~,r'~I!i.d:)'lull plate, S~l.s1:.!I~n!llbilily Ile~d~ to be ,I main I)OCHS of om' e:ffon:s li,:IIii, edm';'<l:I~!on" OiVot,,:n~hii,: !,;1dltC.I!~O!na~. !ln~d e::i'i~:~ ot.~~pa~~t.y,. ress I~~ e:i'> te:l"n~Q[ f!ers~~p~.""mld dl~ pnsi lion n,. pflrl hi glter ed ~1,Calion ~-\S'1Il in :ff1m~'I'It I yoke ill ia society, lh~l"e is ~mpl~l)p',porwniry i~br highe.1" edL1l.t~alion[O belp s;Mft Sf.l'C.:k:lal Olllr~,lS n I.u'!il.';n.rda lleuhliai:e r env iron mer1l~,fI~. ial, sac
mllilly 1I!r>tl'~,


fo:r e ~,e<lllli 'r:te~y. c;,uboI1iLm:U!T<~~'i:a~,11pnSe$, .<I!!i:si!;!iliJm~msho!''''to be more emfirurun'ileilltm Iy, e

ZUld e(;.oliliOm~,c: s.tl,:5l~illll<1:b i.lwty.
Re<rerrlllfl(JeS. <lfu!!1 NO'l'e5
1~ ,A, i((Il1~~

oomnom~i~~lUy.and ~i.dly :~roll!>ibl!~and 1:1.0\'" to s.tJI'P'Oiifl:~JO~i.eies <'Ind.~'uS!lmim] thM SUPI)Qrt .~.j;Wc"I~im!bl,(llluil,Q:I";. \\lhM mngl~:tl~~p~ pen if stu.dents \\lere :reg!1:l,I;i1ldy O1ssigr:ted aO'tmll SlISI:<l~,:I1I~'1biHtYp'u:oblenl:S lll,li: Wf:!re lnougllt to h~:ghe(education by cities, businesses, nonpfi)fitmg:II:Ii.ziiniO:I~s,. "~ld other insl.iHlti,Qus .. ~: f cI<L>;s.l;\Kllh 'eJi:err.::is~ pmd~lced'Jliurk<lb~e comri to d iscllss e'd,c1in. dliscD,pl iue's pal:ent~3i CIQIllribll~ butumls~o sol Wlll.oM.SmdeJ,1.t:'> 'would 'LJI1m(h:rn~m,ld [j(l1:lS;~f)a ffil~(lre~tL~ltlin;)_b~~ fUIilIIl;;_ 111~e !im~eet~ they C~b!l hflV.e [I. r(l~u:li;",'e h'l"!p@.cr~)!l lil\: t\l'Orid through their <]c~·1.{h::::I1I.~jC~e<lrulil.lg"\,IJ!OSl or om j UI1lg:'> itl.,dltdOO:I1I"'I!lfuOl~1:ls~oci<iliOIlS fm':p."ycll.o]~ : O ~n.dltdril s~me~ agy, ~lcio~'O'lY. pJrni~o!!'iophy.~lijg iuw!!.bi(l'~(l£.)~. h~,gh~r OlLC!ltilll i~$~~lU!ti~m!' chemistlL".'I~ cngi!i1.e-enl ng. <il1ilill'O(pOlagy, po~,i~i.(,'td \!i~~e'.re' lhe::ir mis. .. ~!oru i~l :s~alemelJll:S gOil,ls fur pJ':cllQriug. sluclclUs. to hd!~, ~~te ~ be::'llcr !loci~~ ~ seien"c, ~',~th. bro~nje21~lUg.!llIt;:h:~n'CHbr(~; i ~Iy.yetI:his,ide.d ~;.;oien 1101illll;y impielll]e~;tt!d,. 'i'",onten~ :s'[lld ie.~. and odu~~~.V\hrki liltS gillOllpS a!~ fGiCW!:ili.mi~ {Iim. ifi1n,iI~il'ilg ~lHtil.i~b~my i:ri!itu ur~ GUvcn th~ dlall,;n~N {if SUS'l~.ilm!iliill.i:~y ~ ~nd thlil Ileedfior po~,icy ;<Lml be'ji1.I,viorn] modi ~ioJ;1s, Incuh'L,. pmfesgi{)mllde\'e:~oprllleliil .. Mmrn.dilj:;'ds (i!lc:.IUird,i~.g !~~,P!'i(~~fl,ikm. and fLC~~d~t{l~i.oll W W~ n~G1I to ,cl!l!i1ng~'o~~~~plt'i~j~ Irfl:~~~ri~.i~.1 cr~t{;lgiula~01'tvo.!tm;:: !j,\!5til~nabu~i re5~:mrdlu, iU'Ld lhir]~dl,g ,;!~O!l~to the indll~ion uf etTec~iv'[;: ~y l:I.ctiOIm). oe[Oss".d ISloip.llliuuy projcc~s. ]egi:sl<'Ltirve' d]Wlge-l'II~enJt skms ~IU(I0pp;llr~~mi~u:es 1:0l::lki~ briefing.';. and ''''''['\)of:;, 1:0 edm; ..m:: the public "ll;xlU~: !.1e:UiQ:fi O(~. Ctl111pnS ''Iiml off. A.mau::bim'-'l.king ~iro\!,/ to he:lp ,cre3te a sm;;I:.:"IIiuu!b~~ ruture (.17 J" \liJeb site ~:iis:~i 1re<l1.~WCldd S'lISIiI ill~~bM hy pmJ~ Illg AA SH E II !5 )ISCr.r.feS. (1IJ11e:y,es <lIld WI:liliver:si~ loclSlro'lil] blt~iooss. gove:nu.m:u,t]. m:tcillOliprofit: ti:es ill~. the U oited S~a!.e:!> 01 riled, C;l[llada. ~to:fe:fS OIFfm<lmljZ3fiO:lm:s: lalble W stlill,(iiel,lll\l. :facu Ity. aud .)wi all >~x(~~'Siv~nH;OU!IiC~ C~lI!!le:r·of SlY~l~!:~m!b'i]hy \'\),I~~r~6'!;l~ffi just b~"l;!nI~uneh{\d (l~" ~O\'!Js ']"0 .hr<~\I'ea !mst9iimllbl.e nH~JII-e. !;lII:s[ail:l'ii1bi,li~ uu:iil:i<lli.ves ,mcll pol:~cics. di:sn~s,s.ioni~!i:~:S. ~.nm" p.le :syl hibi ~~Q\ilt~,!,g.IMV SI.~smirn~lbilily t~~nb~ ~d~tc.ui~m h.,:s~t'llXl mph~m~n~~·.d K-~.2 ~ Illtht: II J)r. oJrnd:p!fOf~:;sio]1l,d devdQPIUC~Ui!lili,~ri'i.I~, 0[1] Slillsll!Ii [l'<bbilg,ly, OANS (I j) ~,Vf!S l"anl1l?d afler th~ US. P'.utners!hip asked the As::sociatiall of AmeiJictrlnl CoUeges and Univers.ities {M). the AASflE ( JJj, "I~lddie A~ociatro11 (lfU~~i~~itr Le..I.~rs l ltor ijI ,S ml:a~nablc: Fuun:c {.I 6) to (l,:[iDo~ Int;'et~ nl1l~ of 111(" th~lll :20 d iM;ip! il1l<WY<'lS80Cliniou5


liIi9n~r ~~ioil MiG~~on;OIl:5S~~"',i~I}iUi~ C(Nll!~rSjjrn tHEAs:n ~r!!!@nbMi\:in, :1 l.!li'lti~!'j':1.0[)7" 1
~~e!i, 1;J1It:iI!!:~~~J'!gl

.2. rtfil!d'dllio.Il!.~1inlc!1I!iJllo!l ~aFliltl'!!l ~'!\Ii!ablf'




:J. If'Cf ;lllh1i~!l. se~,AilO(~a1:ran ,Df UOilll!H!ly lIlildef5 f~r a ~u~~ilimill!'l: ~u.tl!.!r,e '(UI5A" W!'!IW,uI5fliD!iglr~t(~011~1;_ des~5,l1lm" 'I. (~m:PiH (1IJrul@ (rnU'~mge·. 1!MW"c.lim;!I~INlll.f!n~Mrg. S. ~!MI (j:r,1l't PimC!l1ipe~,W!W\\I~ifi!/O!1ll~rwpm~n~,al1[l, 1\, Il:.ut~~ httjri:l.lliliitdl.li~'ngdrIilgt.r'$.IM"1If !lJ~Mliniltdilb'i'i11. '1. ~;il4io~'alAsl.o:iil.1jCl~101 e;d~(atiaf1!ill, lPiNx!LlrmlMt ~W!J:f!ll



1!!I.5, ~~i'I(!~~i,p r!il'[riJ~~I~io!iP ~~~ Si;i~!a1iiialil~ 1ii!\l~.qlfII{!iill,.'Ii!Si1ingl.oo, IOC. A~II~I:ullil: \ A, ~. The U,S. ~i!lntrslli~ ~11I5<piII1nefi'h'lp.c~,





1!!lJl I!!£M;(. !Mm.:1Mllo'!;.~~C. lJ. Ihlt;:l~I!ll Ed]G!OOlJl <lii~ AcIioo Pcar:lllmfrp.


'll:.!. 1!!I!1J~~mfry ~Hle!!!I's U,

[]if!1illil Cli.rnm'i~mML


El!W~~~~ A:i~QoI;iilli~fI~rlelWi:!!i:rQ.rSLl~itiniJliliJ.itr., r WJMii.aallhe<,n.e~(1Is:. :H. J!i.S.!;OI:iOlion oIl\1in.eri~:uII Qm~i!'.! ~nilILJnl'll!i!iili~5. c
WIWt,~O!{~:.oq~. :li$" A~~odaliQI] forn't@ M"',~lnOOO'i~i:'Il SUlMlfnatbiUt)'in of Higher E_dluC31lDn, wrM'.U~~ilIQrq, ']1,6. I!!Il'Sf" Wi\IW,1I1sf~DrQi. '~7 ~ AAAS, 15 pant o~~tlis me!'wor ., Comtlct Salr:~l1 !I;jI1o~~ .

~b. . rll ~IIPilllidP'lt~. '118. '.' iP<!!i1•. WI\MIJP~:KiI!I~'e'£!1~J1··org, :l!:9.>ustaili'lalil~ '5dtHi!il1s frOj~~ ~.sil:;!ia5i'1;o'M!sOhQru .'Rr,cj~rlIl,afiJIind~:thlml.
21IJl A.ifiti'ocli lI)ii!~rsit~'iIM,.,\t<ln,nOl:lliFI~,~IJiI!!llJ;~it«IB.~

infllll,scd inllOV'i1riomi. 'l~alll~:5!e£. I bj:C~~~li31C::Oml,~ !

llel:t'!ili;~,a!ldrmr~$io:m'li~ dlR:Vdop:~H~ru!. 0Pllor~ umilie:s. AASHl:: all::ii:Qpub:~isbes an dec~m~c~ moJii buUcli:rtI and om'1]1!vnijal d.i~@'J;t.\ivilh cal:!1l~ ~)U1\ sm,l'l i:lml1l i'~ 1:It~'\V,':j, bi'l y :s~o:wU,es.,n::;.'Oourc>e;<;. ~'v(>m~.;Ji!~d o:ppo.~l'ldl ie·lt jab

~!ml'eI~i:lS '!Ive~,LThere are ex'l'lnpi.e.!> of i.m1:lm>~ vatuofll, i.Il~llIdi:lm.g the ~hl!;;~:!ji~l~h!e ~boGlI~ S

2.:l riild~~ liIlI~ F!l!lU.!l~,'lW\lW,f,ITC:i!\!l.1I1l~ut'llHM'rg{, SLlp;pgrij!lg OIl1i!llt Miillllrii!!t TM'W.j(';M1:Ml~og!l(\iln1e~Ll'MIIi'~!l.;1S33jji3:~3roCl

.f1'r-ojt:~I: sPQ,llI::m.red by Shelburne f~trms jgl V(;l~H)nl (19i'; l:hc I3dllC1l!tin,g for Sl~!;t~.i!.l· 'Jb'ility a:LU:iter':s of edlQ,cl.ltioi] p:rol;lmm ,It A~uiOC'h U'~hi~rnhyu'~i!li~g l~~ch~rsfor rhli)












ASTIRO INOMY RillIJ~!!lIV rol~lliltg Sl/alr:s, [i!? 'rI",t;tI~IlC d ~p,hefi!~$ ~

\!lith p()'le:~ t~1Ii~are hOIU~r "-hi!ill the eqIJ ~~or.


m.ovetowa:1'd tbe eqmnor.g~.vl.UlIgtil:u~:rn anelljp-


u~'[.[!:;. c~rol~$~l ri,dL;;!'l;Would :;Ud~!lwmt
[fI"D. J~.l~jlrneCenlif: .. df

:.>liU limit~d 'by d[n:l:fletkl:~ ~uthe tl,j'll;'~,o(j~':io
3!perl~lre. Even the l!3ll:g:e~ CUI.'"I,''er.lli telescopes see ~11 but .~ll<ltl!dfl.!~of lh~ very l.lrge;;;tc(l(}1 Slm:!i as I;lu:oopoi:n;~s ofhglu.

i~W'flA:~tlg Qr;;I~~iOllboOOi':5~et'ld:i ~\)

rt Ol ..

he.Jd i:l1. p.,.~"ut: t.~l!!:: . .•e.

Ihi;;rohllinnr~(~ and vil,;;\Vlng g.I;l()~H' ·~f fry Vega,. it w ill be' pos;!!.ibl.e m improve iastru1VI!;l'i1t

~olcl.!1bulgi::. "hi~ e~re<;(is well known f()r
E<arlh. whi.ch 11<15 ;IH er~u<l~ori;J1 F'Jdius some .20
"m~1n'lg('r lh(lmll lliDe ro!ar radius. T~e SI!,II1I, s i o:I~ly :i>lig.lilltDy);uitened because of:ilS. slowmtJ~ f tf~J!l !)H~.wi.Q~~l!l~q~"llorjfd ITtd~~!sI1\\:c~e.di~g the 'pcl m: radius by OJ~~y oae part in ~. :; .. 0 .other s.1<lrs. however, l~tate'!JCry rap,idly .. Crill. jJ;".g.e 342 ohhis ~i~!I.]e. MOl'lnier iff (;1/. U) prc~~'mI,llima.gct, of AJtmi~:'. !I.I::;o k~lOW~l:;IS ,01;

AqJ!JIilae, one onhe tbree Stm:s of~i1rH;:SiJlllill mer
Tri.~~],g~c" hcl':(:: :~ma,~c~d(;':a$'ly T
t;~~l{"'V tb{;li

cal ibr.:lli ()iI1~, Stellar relation also ~l~,li Sl'IDll.g implicaTo (l,bt,jirl i.nHlges of Ah~l~,r\ !'mrfUc\'i. li(ln~ for out m!lde~u:$linlldiillg ~fdle fOrrtlinion l\~IO:lJirli.eref al. h,-,(~to (;ou:illhil1e the .I~ghl fmL~.u aad t!'vo'~'lll,~i:olm, f galax ie s, neca lise dI~~sw o nr bUuid~c~copes ofa stx·eIJ~memlt flrl"'.$y~ocfli~ ,gal::lxic:s CiUMlOD be resolved lnm imlividtl,,'1ill on Ml W]b;;on near los Angdes. 11m, such stars, l:lia.c.irobserved propetries cannot be th(l m.(;):n,fli;I..::I]al!" P 0Pl,il,· irn~rfefOmtllrh.: iurr:ilYi.>, lh~re...ol~tioll l$ gh~n i!n~rpmredwi[hom by the; sep<llI<llii,Ol] bet.\\eelli~;he in:(Iivid[11l:1Itelelation models, These II'llode:~is,assume ~:h,E1l' these g<tl~~~je!l (:{)IlJUI.i nil. CCWIIJlJ :~ni:ii.0 r S;ll'I.r~, SlOoIM::S.not thc.ir sizes; Monni.er ~t at. Bcmld d'll~~ ,.1 "Il'eie!it,;o,[pe" \'I,'ilh in"! enecti'!.~ size ,of \11' ith jJ preserfbed di ~~tibuli un of masses '~rld me 265 In by ~. :m. lhelm is,,~mwv~l", a price to ages.: I.~~~C stars ~J:~ (X)n~ct.iveIJl wes:poom,i:blc 95 ~)o:1rhe Igl~~l"en'i~ued by the di:Slalil~ g<]I.•t~ y. r ... Rotat iOI~ ~ni:ct!!: l~c;l'Q'l~h,\!ardapp~~:~~~e:

:S~ieUj,!J :SihOlJi)es.{Ce!l!ter} Riapld!ly rol.atillg Sitars de'l'elojl .In ~ UI~J~(Ir~~ lbl!l~e:, whid'l is ,t:ooler (il!IIli!.l [11us reddcr} th~!'l~h~ polar rel1li(lHS, as seen by Man!'lier·eI al. (lI!:ef~' Radiatioill pressure d riiws a fast wind] rmJl[~U1~[pala r r8gIDIls ir ll1~Varefillflldllntly hot 'I'lhe~!Nisd~sts ,nel'rle(jtUlll~ly fo:rme.:l in the ,e.q1Jatornalp1,llne., n~t9li1UStarn in w\!~ioh il1~ Ir{it~t~oo, ~ate d~re~St~ s:tro~gly willi! dl~til!1CehQm th~ ,jj:l=i!$ !l!1l1~ l1~w YI'!!H!.!!.1I. shapes (wnltt (O!1lVect;ve r,egJQ'''~~had~(n (di~gr;aHl i.ld(!p~d rr"m; (8')]. Alt3i~,' is

the p(l~ ar regh.1!l i~ 1'I1i~I,chbti gful(~rlt,;;u1J the

ntIlu:ened by





f!(I]U~:l:Qirial zone. This ~'lgree'S w.~lh the theory (],f !'teU~"Ir trl!c'~ur,(l;.\ ... ~ . h~d'l htlldis Ih,u \I·fl riat ions of the gl::!.vi.ly ann ss a ste Um surface ~nus::t bc!iH::OOI'lilI~>an~~dby v~:rifllioni>in t"C'I.1ilJ~ l:le-r.Il'ure (2)1, For Alt a ir, 't~leoemrli,!1Llg,,~ fbllTOIl: Cu~t!lllC'nM':li.fi,g [hc~lI<lvil~uio(l;ljal ~nlfiii,~t~on ~~~lr
the eGIi~.lil;t,or tlliietdore causes coot ins-a Iii.dl

and e m.iued specrruea of stars,.dl:i: rate ;at p~liy fbI' suhsri LU~:ung<I 11t!!ll~e: [(deSl'O;re w ith <I ]nSl:~l;ld,of wl'l,kh they ]OSi~!11ilI~S~. ~uid ~~~irifilfc;:rioii geuing ~11I,1iU,1:!!t~1II: d.etllLilkd innag;e. one obtains struct l~l1e.,... i~ll iaturu dele.rmi nell' ~,lO\V ,~: h f<lSl a :;;;eel ofD"<'I:rhtlD" ,,.b!Stmcl d~t,l dl,U d~1 nell eil~i1y the S'~lIil'~ burn rhelr hydrog:~ll $uQPpl}tal:ld ho~:v tiley evolve t1IVC'D' time. To avoid mis;~~~,emll~~""msL':hrcs0 iutuitive .unlcrprel"uioll. 1 I~rn~,rct ing gll.1~ct le :!:l~~ I.ra, the assumed ee ~llo~ni£rt;:'1til" have UO\\' l>ucecedcd i.l1, IlrO{!H!}~ng s)!nthel:ic i:~l1.~ (Df Ah~illi'lhrollgh age"~ comsueU~u' Opll ~<I~ musttherefore eontai 1'1 IJ I~Oil pute[ rlrOC~~~~i1g f ~}~[;(ihlt,.;rf~rO:lliil,~tlik' d~Ut ~ Ih~' COl:lroc;;t~i~ix of stars wl Ih d:i~Tcr,~~~liI'OUISuchrn,t:'tbo ds <!recaasi ng asrronomers to. I,~on rates, aswell as [the COMet t«:ilpes fUll"
slm!i.'i~Ii'littll'1i~'x;r flf~]~~6r;·si~'1.ill o~~; r~ttSc~~_",", ~)~n~

thtls~dl:!1!"Xld brigh~!.l!:;,.;tr-\\'~~h~spe;::ctiX:1 th!)' !Julies {.~r;: rhe fi,g~!nl. t:l;l~ref panel), Nat IJ,L~ly l!Q(l$~J~.e wor.k (If Monnier ei al. ~t'!Uus some~hftny ne"M ..bo!JIt 'rhi~rJ:~rli,cl!k.robject. i~. :.~1't() shm¥s d~illl dct~lu~ed ~.mflgiIli£ of 01 11ef'm,m,LU'l1

,tlf~h~~XI~V~!~~i wisdoml, ,,~!'i~~

'h~'i $tntC[IiJ!'Ii; .md~voi~~ r io!'i,
Forilldi\':idWl~ll ~Iin'~. r{JIt,n:iull! has <I dramlltie influence an mass lossand winds {free: Im;e riglllre. ~;ef~:~]m:I). [m~il1e splJlniul~ fllP'~, st~r~I]~ccC:llI~mil:!iiIgi!.I lorces ~/oI'Ou!d d~ mllU ~l !i!O'l.~.le jJoinl they (I\ier~;,hd m (he gn1:viuu~ona1i 31:wm;I~IQIli ~10dt~.le shu n li:c:s apon:rt\¥e do not knmv ~~(l.Wdo~ly s~ar.;, cm~f~p~b this:cri!ic,d r{liOnion m;I:e, but it.!lp:pcB!W:!l t~]jjn ce,l~n'ifLJ1~ ~l~ fo!'Cil's I;:'"nl r~einl~Telhelo"R~r fru1I~S ~lnd 1111.eQ:mTI",{IJ(J!I~ diisks ~n the eqll ...tIlli<'l~.t)~m1e. r of

st1J[:s \,v~1I be po~si.lJ1}e.
As;~:roIlQmitai. inl<lgesl~lkie'll

inll:d·i;:rl)m~~.ri.c O~~c(\,r;;~lio:l1I~ o:f\!\~ga (0: :Ly rae. <111,50in the Sunnae r Trlf~8Ilg~e) ,h~vc r~("emly shtrwg]~,h;;H: s:~aris OJ r"'!lp~d~~a~ (hi::. tou.IOO (1. 4). U~~likc Altlll.i:l". hm¥evi::'I"" v.m see this star nCi1dy pol,e~n. Ve,g,l had: ]nllig ooe~] by gl~(l'Lull,d~ lIsed ...S:I sl:<Iimbud sl;;lr :for tile cn~.i[)r<llion of !Il1m.d be

l:J<ii.\~(;d ~de"~c(lp~ ;)JOO!l:orm1:! I~~'rh.l:g!!':ilOOd 11)1 tliXf~ IJlI~lem;;e .in E<Lrlh~s ""Inmlo!>phere. Re~eaKllil:1:S havii.l r~t~n~~y d\;;~iell['lpt'd !.:\,"l)'~ Q,o com~ns.;;~l~ ~~ this ~u'llOJ;~lhe:~riic: luriI,·illg. \!\I.ulh ada~jt,~ve opt.i~ b
:~ ~a~ :S:li~~';;:I~~s~ butl~ie

groq.HldJ·bas~d af!!d sp!lc~bf~r!'!~ (~lel'(':op~s im;;~:l'lIme'l:Ill~. Sonne .of the ,asS:lli1,ill'P'~,~:OImIS l!ndl;;r~f~~,g dle."~ clTIi~ibnrl.i(HIS lIiItl<tVO'!pp~'I.r Ittl
Ol1l, sh<liky gl.'OlIIld,

as they



~SlO~~~t~o~!lchielllllbk <



,~ 11!IIilmmil/l' or Heqd:ellierg, [li·~9'11,~ l1l!ltlelber,g, GeIm~Iii:V.
t:I tE:'lII<Iil~ a,.quir~nl!l~dh@!'!!'I'li.lrni~h~~d:.elber_g1ile'

T~l~ <JJ!II!h;~!," i~ ·<!t!th~' ~,lf!d~~~!!!fiIw·~~~e .!{(!n~g,~tllll!,

;;,p~leri.~~I~·~Y s.ym~llelri.'C sidlflf. mod~I~. Such On~lc o'J~C'rh~~]4 fad lh(ltU~c pol~ qUl th~ IOQdd:s dQ nO!: ulke :into m::COllnt ttl:1:: :~·liIlnU· be ::;tlb.st<1nlt~;lUy hOi:tl:!f thmlt~le eqmJitiori:all !f;)!liDce f mtOliI:lofl ~, the struct:t~r~ of lh~ l>'t~r., Ngiofl& C(lJ)Jg~v~!!i~ t.o ~. s~m~.~ ~l,li!ass-'~oss o ru~ech;m iS:l~,l.mu.nd,yl:liu: gt1'll!enltuon of <1 fil st or\i\'he~b,er aile :sees Pl,'~&o~jlllilil~~ntlyl~,e ecl'LI<l;· lodal or polar r~gkHls, No ..'!;' th,a~ '\i\~ lkllu'w willd l~,1J":Ot1,ghro.d].uio!'l r~e-%ur~. :~:;lik;~liy h



~hnt ~qlWi~rrD"iOl!I disk:;; <l~"d polar 'i\,li!ld~ c'a!~ coeX~stU'1l1 scingle objecHs: both phenomena lend liIu::.l1!ll!';ch-es to i:ml]agi:mig studies w.i~h interfcrO:I1~,ellS"ffil2ffl'("" 6). A solid bady Uk!e Earth l'b':!I,H.,t. reuue unlforml),. wi~:hall p,Uf~f; om~~le'~in~ <'I revolution c ill tb~ S!!L1n\;~li!IiDMl," i:s i~~m1l rue .Ihr g!J~OlUS "Jib bed ies, \~ h~v,elo:ng known [hod n~~1 U!Uta1,"'\ Ili<l~ regions of til Ie SUIm aeed ~boul.25 d3yS. for one ruvoh, ii~]~l.\~i~,er'cu$ h~ 'po\l"r reg i(!lns t u.I'ke !ievc;m I,(JIaYl> l,o~,gcr.l'lmmgh~~,e illtll<I]lys~,s m>SiJm~S u~lif{)\~n\!Jro~';H:in~~. Th~ylolllt'!d sul,[le disc~.'epilBdes. whlkh could \\le:]l:be c'xplatned
by a. made s~ ,3u;nmmt ofl1loFrllul.i [(linn ro~i'l:I.iQlEII.

Mod.e]~ (.:f ,:.;;I:<Irs. ~t[Ollg dg.fferemi.~~.ro~a~ wiith

nOM,!o.subde modi f:iG:lI!ti.ons of the
ia:l\ e:X.ln:~.llii;: C<1!!!eSthe

mfG'ld I ~eUm:, stIl.ICtJi.lre~

OJ) kU()m(~lrie btt;;;elhl'liis 0oli!ldp!:'O(]u~:e much mO.f11l: detai ledimeges a f stel l'I,1L $l~~fi]Cles. revealing s~~r ",polS" c:rupt~ml!S.!)nd (;onvec~ lil[ln p"merns, ;lIrnd e~.l.ilh~imlg studies nf eomplcx. pl'iieno.Olcn:)llbat tod~y (::lIU bemeasured orlly in the SUDi!I.
:L J,1l Moilillili~rJ!'(0I.;S~~'31~. ;f;~2~Q(I1)<;

Sl;;!ii'S Inighl :~!lV~ l,a!lCOml~,,\

i;~13:p~:li IIii ~If\~:r

b~I\'\1W!1Ja Kpher~ ~nd 2. dOilliUl (N). ]1 h,!1;fU kno""'~a:l1 whether such ",exoti:c'" stars e..x iisl. but


th~t could be disco\,f~n~d with p~;.il,;lm·dwy


of~~'ny o~cill,niOrl~ of'the Slalfiw:~th methuds .,mllog,ous lO~hQse used to i:mlfeUFd'ae ~WlJ'llI,cture
of EArth from =-en~I~~,~c events, ~l'i>trolilflmern ~:la\~e ett:lJ'n'!l!illeci tliie :licmniollm.lrelte for lire iJnle~ d : nQr fl,f Ille:~hQ~1[7).
Nlo:Il1iIm,ii:er etai. cornp<nred their observadOll!!! to the si.m,ptest pos,!>.lble 11ilI(ld1:'~,. whi.c:hl

:E,I.ucid'll!:itl!g the drmn.~tic effect:::;of rotatinu is but one gh~,e c(JIJiDlri:b(uu:]oll$. lin!er~ th,u



caw. mskc to stellar :k~uophysics"

W.ilh telescopes separated by several hundr~d me:l~rl', current fOld Ii tles ~U'~ 'v~~l !'~,i~~d
~o Imlemmrill~ the "glob a I" properties of stars, Aliuun!' ma.·~~y \Vil:~, 20110 3'0 teleseepes

,6. 7.


onll!1 e n M~200q (ro.l,lUooeot;~" ·llLH:OO\s'~. iI. Wln ~Jl\ilil~tMDn. Nrlt R.MJ,Ii~III. S'!X. a~". i~5 U'924J, . £ J, p'JlU;Jr~~'rI ~!;-~~J, M~, iloPllG~OO6l[l. M. Il'eDff~Jl e.I' al., N.rmne 4~O,a!il6,12006l. A. tQtiirl'>!!mIMdlI8al.. ,w1DPl'lfl,,). 479'• '1;17~jJ;99'l• f. ~!bel. el,al .• M~~~ n7, 37 ~,0(7). M. ~]ltIDmI!l':!iOO,ArWIl\, Gwphys. 45\ ,U (lOO4t I( 'Il, MatGli\!!~P1; S. J:xHom,l!i. S:wmanid'l, t S, Mel~I~,Ii.S'lIiQpj\ly$.J. 61$3. :5.60 W:IO'7t

Quelelf] bee

prevent YOIIII'IQllbee~

the PB2 [love] l1mm N,(lw I'Ibrld. f\ldo ..~:;; U)1i(:y .crc~U!;l:d, ~'!l~i(ltywh(;ro H a fetuses deve.lop lilil bettles aad alte tre.aled wi~'h e~,('mi.e;) Is t~ modi fy U~~ir e di~SfUld b rnelillill;jllies tl ).Later. ,ebiMrt'illi111iif! Sll~ee'p;oolli~ diti.muil to theUrliJ.lu:re' task run society ..This

hom l'~rn~n:goo ~tili1l'!J l:(ntil tllil,~ l.e,ave~he hi\l'f:.
lIl1ilke~the! I1tves,lIfer.

. · I

gus !.nxten witll1in:llill:emf:SI:. im'e.lrnm.ed~!<n~:.-sized i!mdi'vidmd~~ search and !coll~el l~\I!3$.llh)!,U the fores~ ~o feed thel\mgus."uui hn:~e i[hdlividmds wh~ ~tro~~mfmdibh~ d~fendlh~c(i\IQ!'IY. HOJ1,eyooes have evolveda differenl caste system: 1fhle .illd iv idLll2li worker bees (a~lwilYs I
fenl<lie~. 'b~<m:'lem:;111;: drones

~im"'lf!vec. t~,le ife of ~1 I hon[!yboeeis> \\,·iu;;d. Bees ~ ~~~glyi~ll~lli~(:ilt
and .Ie:<ifllla lo;t,llooLl:t Iheir,envummllelll!.


:mliD parfUll!

lic~.I3f.lb!",J.~t b1;~ l~!1Ii!color. edar,




lPe<J1~,h: \'-':~o

have ,ch;:ar



reles, pm.d~gl~,e:l1.~,in castes, r.ulgingit11l];l <~lph1JStthe :Ic~ten;) to ep~~I()!111~ drones], Uh~

~ribl,i[cto soclal IWe ilI:~rt fi'·oln !nadn,g W,ill'n .. que!!;~lS durimg mlp.t.ual Iligllns) r!Oerlbrm differ ... iti,V'I: I~:jj.,e fo{)d",f\!·hned} leami~g <U]dim:,L~,lya:y" em tasks in thecourse oflh;e~:rlives, The ~,'Ork~ ,lind the lia.oJrneylJee has become an ,~mpO:r1i<mt .Among ot~~.er I:hings,. lower castes are 'pfiO~ ro ets Sitilrl off'fls !ltu~ ~~$!Idi~,g [he hi\l'L;;righ~ model ;;till~m,I'~ Ihi,~ euuj t5l to grolmilmt'd !1(]~1:0 11(; lf~g:gre""s ive f\gfihnit hijgh~r caste membeis, A tf1etl:Mine,lml,w.il:h :uaCllf1oloxic :dte:l'cmergenoe. OO'unlnm:u.e with chemicals {~~(~~udlillg Oi!~c{1boll dl~Ji~g develiasks l'ucb as n~~t.. 'L~ildi~g,~~ld b OpnM~1rH lead s 10 the ilIpprolni~l:e brai n ~;li:\iedefel:l!S€. and li'lellll end t~.c~.r OI~9_1lJ~s.. pa~384 ul'tbi~ iil$u.c, Vorgo2:t1I'oi. On ii'llC'.ll as Ibmg'Crs b:.~ ,t~]~!~ctillg, (2) eluc id"lte .sOJm.be' of lhe ehem iC<ll. cues lhm nectar O1Irlcl, pol~en ~'O 1Tl000iw i~llu'~~lc~I~~r!'llh~,g,dsve iOP!ml,~jlli~l 11 il ~1O~ti.o'Y~li,1Ic~d1!I1~irSiR~t;ln-;" ~uld <1CtL.m~ .uiiii~md ~:.,,:ste SySlel11l,l the lil(!I:ie}~bees.. ,BO~,!1 is this de .. e.~opl1u::nt<l1 ~ \;V~thimlth~ 1ui!im...lki~gdam. :;.ud~1i!m~~~t> r()'·~}I~H~i~"ll Qtgu'li?~d!A~ b; h,we e\tYI\f~d ~hl: gHo!'j1iSwbir: c.~~e f;!o<;i:e~ie:l,. ~h~ roO!ilfa rt!ctor thm~ ~le;rM~ny <linl spe~Ies~\:lj'\.'e- 1I. wide r.1:lmgcof c:.,,sh~S.. In1nes ~he t~tsk tlmrnt allind~v~'(~~ fro!1l\,\iorke;r5 w fq)n~.ger~. I:torll groomets I~O ui1U'i'i,inp~lf~ibfm.medi<ll:OO by soldiers (3J!. ~'nd~'IIidllals 21111 develop ihl>lU I."gg... reglll~l!on wll.h jmfcn ih:: hor~ laid by tM: SOl~le 1fI;(j~11er-the l."olony's'(IUeen. nlone (4). bl'~ feedbilcklRl:liill (~:el11lcrnlly. due food supp.liled to ol"ach egg,isl:iill.e ~he hive :~:s.. ~so inmlpon<nm.l. . lbio~t~gic~~, i~l1id ~hJi!1 S I~~dl' d1i...; I~mbry(l HI Much i!1JfornmNt:llJil is d~li\;'ered dieve\l~op; into one C<lSI:eor Hlliothc'r. <l. situmion b:y~liiIef1omol~f!'s;. Fa!' exa'l11I.lpi.e •. rem~I1~!~eem ~fl1u.'\;,ley1: fiC:lklln~LIworM. Thu~. \V:~.~na h~\~" lo:;.",s Ih~ {IU~~~L if v.'Clake m; <lin CJUIIr.u,p]e lhiC le'<If-c IJII>er.' ,un ~~e'rq,lil.een ma:mi.diubular'plu:m~ Ati~,{{:'l(m~t, :>rtr14'IJiI ~.ndivjd1,.l:ds. Ielild lo~l]~li~ll~ i ~~lOlle~ QMJ') wi I] also di'iWllPeilf. 1,e,ulinlilliO 11!p id clill.lnges :io be.~f~,dor :;9mo!1!ig \ii/f~rter IfJiI@liI1iciilll(O'I~1Ji!!1II.. th~ Nlul~S'~' S'mrrmmd~he' queem (kl'~l1lified ~y lIil!~ bees

lio~1 of necl<lNkUn. IlmYeJiSlllld U5!.e this in Ibr~ nmt.lon to opti.mize Lhe~r haevest, This Ci~~ity has been used :rof WIl<!lW re.ns~oi.e'IU. mere ,a'bo~:u bade mecl.bffi]li:frn.~sl,Jl.~.d,er.iyi~lg the apf.Kll~


".~~hor i~ ill! ~hI!! ID~p~lr!l1l!~nl Ilf tljgUJ:Q!iljg['Q9Y, Il1ni\!m~fI! ~(J.m51.3m. 0",'845 I Kon~tll~il. (il!irmalll,y. E,mail;'GaU~o!@!Ini~:on~laailz',i!le


bees. \,\"ho sl<lrt r.,j sillig




ma,Mk) ,erid glKlOIi'll Ih:elr.. 11IIIi:'lboehaviorr ~~ impo J'it.illilt 'for dlis~lIil:!uliiln,9 lliu~ q,i!lil'~"s ph~r'()mJl):n.e', 'Q/l~lp.wilhiiil ~h,~ IMUr~iiliv€:,

~lO~ m{Jri I~ lind aggresI~~ir ttl:'!t ire Ii!e Sr<l:~+The: m~nli u b.U!O!)of p sive.W~l)fwmdd aversive llle51rnillg be brain ac:tivity lJ:y the !(]I~teen. n'ocKh~h.tling learnblocked ina :yollng In~lr$e' 1Xe?W~,t~li.n the iln.£: C'lIP<lC.~ly in young beeste makel.hrnu mere hive. the sl:iliil.g re~lex t~;l[l (lili1.~y h~'fe neg,<l~~\f![~doci~Je. isa tHfI:erel1l~ view of parenrhecd, As C'lfc,ct~,. Th,us •. pl1~~,.e~uiqlg 1l1iJl,I'SC bees :lhl'In t~'1csa.Y.ill!!): gues:.Wile~ i:~]ik:lren are ym.un.g • de\ielopi ng;}ve~:ive memories <I:ai n srrhe give them :Lf<:'1'{j~S;: when l~,ey ;mw. give ~hel1l:1. l Odl'lfS h'n th~ hive, w!~~d~indlL(lG: l:~l~~11~1'l1~~'~ \V.~ngs.the bG'C:nulth~r ~!L;'~~lii:,it'! h!l\le i.wol'!Jt,'d tereer lOt i:mu~!.is 1U!~ uneend ili()n~d ~limiJll~~, own odor, nlf!kt."l> ~he Cti~ony ~lo!'·i;,'j :-:ee~.H·i;,'j'. c::~ac'~ IYI~is sm:~t~gy far h~ frunBy, {US} such <Is.gWfetl~ectjlr or <'I :ua.Ol(io~ISelectric With .increasing age.. bees atartto lle,we the ReJie;Jr'i!!!I!l~S $t~()l:kl.The: US ·eiidnl, <!I r~SI,lloC)I1~. f{)rr.l~llrw.~ C'i;l.I~ny.ny lo di:iM:~!m~:iJrilgil.g sires. ~1If!Ld:pe;f.l(, tI, :!!fW~, ~\fIl..-e1tew-Wc.!\!i\d·~I)Iil~f!lW~j],. Qrd~ ple,t:llae exiension oft:lia,ewllgIlIe to Iickl:ll,e IJlOC~ fO]l".[lffil tasks O1Ms:i.dca:hlfl hive where Lhey need (it:y, IHY, 1 ?32:il tin or l1la.e I;,'i;'\,le;m,iol:loHilLe ,~ting thr d~fe:nst:. to learn iliil,!!Ji!!Jlly ,,!bo~rlf:""I"re;e~ necTar bHI'j!~::K) :t '1;1. VefgDl" H.A. ~l1Ir~ufl:'A. It, iMfl,re;r, S'detlre 317" ~s:q. r ~tOOin. Aller cO[Jd~~~Olli~,£.the CS 310ne (,e.£ ... d:te aboll[ ~l.'.ISty d~~ge;li1'.O.h is useful, under these 3. 8, H!5ll'ddbl@(,I!. a. Wil:!iclJI.ll'te,,,tnrJ; (BeI1;~~il' f.ire~ '01 odhr) \'~iII ~Hcit tile cmndilio:n~d response ..ThG' ei rcumsraeees, Ihl.'ll! the ~lTeC'~of QM P 10 Har!l;im Uni\!. !!~:1'. C<!ml!fldglii. Wl. i~Q)'. !l,S;, E. IF~~m.adl, (i. IE.~bin~n, ~ N~rJfl!J~i. Ill',. n,0'~ neural ;'5WIb.s~·]<I~e:s ilI[1Jver'Si,,o;e liI:Sm~dror a bku::k . .:a\lers~,vel,e'lll"n~:I1,.S wanes, Tbis jsa Ilor


R~~IU~:Y, il has 'OO~I~ shown tlllfll bG~$;a150 learnte assoc iale llilOXiotus st i~.mIH. ami (Ill expef~menlill paradigm ~la-s.been developed whereby ,UI uder can be ''1~~ci~tled\,,'i[h!h.e .s1'~!.lgrencx. (.6). [~aslOClCii!m.i,,'Ch::~U[hmg. 3 sti IliiI:· ~,~I'!J:S wiJh,OI!,H 1l1e<1'~i[l~ [n. c(mdit:~oned $;~imllh~s K'S) R!,lel~~. an uder] is Ul>.~Od~i~t;d with ~ r,~i'm!i"

~fv~ gJjndbecome


pO$,i~~'!i,Ie:US dilll"if in b(;l~bralas, l10llli lmenns
of the neu mm, im10i vesl ,3 ndin terms of th,e n~uroln'Ullsm.incr' tilley UiSC. The ~pr'~t;~li\!~US ,e!mli1:li1et l~lJl"S tl(':Wp~l'IlIi~le,m; fl ~r.~nsmi~,rer I( 7). whereas Ihe a\l'ersiv~ US u~ dOp~lUi:llL~(6)1, Istltie leaw.li1iQg c,llpm:: ~tyef 0\ bee lre] ..ledliO l her dev¢lopln(:"ill~al. $lL~'&."F;ioH oftas~s'?~rui: f!el:>ulls ha\'\l ~r~flted~.~. H'!p~'Ct;;d~!w;;dI Unk ~ ti.'O'rnlullil,olccu~s. ,III t'l'a.e Wd~f to' cOlllpl!ex.ooha'lF
iQ:D". '"!Ilil ~)me(lrnhe ~H(lII;XlldQfeiJiecrs

\i\lo:~d~]"f~1 '~xamp]e: i~~ \\,'~r~Ch{~le ff(i~'~f.(l ~ o
releaser pheromoae


il::iimbe riQl:~owed all the to t~l·(;:II'ClJfODS, lh~[ arebelag modul~l>ed.

5·. Iii.

v. ~oz,


ll'I~m~e:tM ..IGi!.lrlt 'ilf!'l~Co.gn,Sa. .5, 6l R:OOU. E. RC~:!'!li!lj]'C..$~iI.dQZ,'ill. GrlIrf~,~~ O,tvf ,

and Ulen to be~a\li,m;:'i:l~mod~fic'ltioli1.!'f, l'h~I£, honeybees: diff-er :rubS!:aIUi,1IiUy f.·mn ~he beiegs ill1li-iuxli:ey's world, because .il'iduviduals are 110t tl1l.pped withi~ tb,cir castes for

2~iI!':ll eM~OO7), 7. M~liIi!rl'lIn~r; f{~J'Hi~ lI!i!'IiiO$(~ .il~1 2:~.'5.'!'1~'7} • I. ~,T" e:llgg~ f.IIlI:" ,f':{(X, ~'tJ.rJ. irilt Sd, U.S'A :111M. 2iil60 k '(200(1).

of OM P

'fIIl.'e:Iil.O'\ll,f undersreed, It turns om duu QMP d1iJilOCtly m~]ue[I:I::e~'~11eC'henJil!i~[)'eftbe brain in an ;i~gcN:lepelldl::'mll manner, cOlurulbming ~l:)
dj~\.\L;l~f)P;f;llGIiIW~, pol~"i&ttnl,



nent ofQMP is hOlnl1V<L!11~lllrlO1ti:."OIKlllHVA). a. $t.ibl'I~!:'!t'iil\\I~tha ~,tliking d~mk"'ll simil~ty II) tlln.eb:i,ogeni!C .Uv:llilnedop<lnmine. the Ileurolml~:!l~
If!i'ITiIII.ing. i:ndeed QM Pi <lCis di reedy O~ the d!tlpm'l~,i~le [%ldlW;lY: levels of dOl'i,i'I!mi.m:.illl!ll.C b]:iil[nl.m:c redM.ced1 ill j'tltlng bees e?i.plll'>~J In QMr. aud this eflibct ~s <IImp~i:ri:edby 11. oo~.curre:un re,ollctiou in Ihe lev~r~ dOp;ill'!'1lil~B-~n,~i[ rooop~ms (l'il Thas, of ive exposuI"e of~ IIlU!IFSe bee to IbJe queen'!' oo.Qir d!(I\Ml·r~,g~l]m~ thebr~1n~~ d)rlpiu~~ill1i~!~!i,i\.~!, and reduces 3c;r.ivity levels, WllliIl L~ Ibl.i(}:ll'lgica~ C!l)I1~qU~~UlC'onh[~. the miner


Metabolic -co(lperatiQIJl ,e.rlabl,es·sofllf!: mim:lb~a~.partners
til [Ill neither partner

l'Ol~ rive

on t~~\1'e'1lrtrgy '~~rb'OI1ISOYrces !


ld 1Jt.iliz~ on its


dJo''''':'lJl~regWlhu:ioJib'JVergo,l fli al. 1100'W .\1hrn.v~'lm~,:~: yOU~1~ be~~.; :;{,"(]~~d [1;) QM P' ~n;liIflt ;fIb'~~ttl ~ I.em.lii noxiolls M'inmli .. Uis defwt.is nOI:1Jgeml.~ g.enome ~qm:m:e ol1the anm::rn!h.ic lJ",cte:ri,mn c'll:~Y:1 ~he lilel'moo.ynmni.cs fot goo\wh ;i!llie er.l!~ ~~tr!ling, d~ficithL;l~!iiJ~ flr~tiii~~I~~~,. S):,.mvplufs (lddirr'opilit:Ns. Mc]~~n~~ t:t .(1{, f~V{,H11b:J~ '~)!lly \"'~!~L:;!il ~h~ PtirH~¢n>~ng.:;~M~ i~l iliilg .i5 ~i~~l Oi.n~cl~d.F~nhi;lnm:lI:e. ~h~ u\;::ficil is, (n ilfl:j)ot!A;'idiI1~\r il!isighi~ ~:n1lO MlJ~lJ[;;i gflh~ fimlfi~mbd ie 'C()upt;:lt'llkI'UII. ( 4',,3). Umited to youllg bees.. Specir.,cilI,lIy, d~e" d ii1lJllent<l~. machi:Iil.C:IiY 1.'OCJildn:odIFor livi flIg i:tl TIte b~llonm 'seque~liloe o:rr ... (fciJitJ"flpliiclfSo \ I]fe5~I~ermodyli;flimi.cedge. :!ll!lhOfS 5.~,Q\,\i"\iWth .'l\il::tl\.~"e o(lndi ~io:ll.il'lI.!t!lrnat (/) p:ro\'.ide:s n:e~"'ins~,gh(imoth.e de'tailisofthis young bees exposed to QM!> ,CilIJlIlOl .~e2lf'llto s. acidi.tnJphk.'m grow ... ~ml.in1~y y fL :sy.m= untis-uill an!lcmb[:c Ure..sly~~".f\.S Icxpoctedl. the b eXle;lIfid!heir::lri~,g ~o[Jn od!mr th::n 1i1" s been b~(JI:i("p.mcess b10\~lnfls S)fllllropbY-<l metage:~oo.le eillMes Iiu Ie po~entu,~I] Ot <'I ,re;spird>r pl:esemt!ed ~CI!fel!;l,er witb <In e~:eClrj:c:: ::ohock. bo~:ic coapemliOil miuaUy u:l.lwJiV~n!I'ilj,'O ~my:l!iIlue'~l'lb()~i ..ml~~s.~ng ex~elFllli1ll.tem~inil~.e.I~(."=' am).~robic mic!'{lb~~, hi) which e~,eh rf!~:rller tOOl)'! iloOOptOrl> lib~ \1I.Ih~roa% yOl:U~~h~es tmJ)T ~l'l,ro!'~d QMP are ~t'l ,~xyg~ll. nitr.u~. :l\.Jtn~fIliW,t~ ~dr;lI:e.or iron,. 1.1 illS{) ~m:b Hlrul)! olhh,e geillcs very good :U this tiI:sk. Aversive lemnil1~ ~n d.epends m] the oLi:Lel' ~Of ~owlh on ii, .t;pecific lllL older OO~ is ais;t'I~llW.CI. sub~lr£~[!e(2,. fur ,ex~:mp~~. Clcidi~mph~c.il'~· req uJ!'i:id iu,r ~:~g'mili:m;.ui dle (lIth~r mai i1! s. TIllis,. lhepresem.:c Orl:~]e qneelm. through C'~ln efic.i.e~1[1Y' degmdtl f,u.tym:;kls;or bem~zome 1)<It~lW<iY bOl~ancillg o.dd~zllin,g and ~iedlleifllg fa.r her p~~eromon'f;. i!l~.nuem;:esthe be'h::'l!vi,or. ~uld in th~ nbS!enQ~of QlXjlgl;:ll. b~ll 'Oll~.}rwhf!!JlI •.1 syn~ Iml~nli~1 i~ th~abs.em::e of Q!Xyg,'CIl •. ]mlead. indeed the brn.in ph.ulH<l!l;o~cgy; onler~!i'i,1e. tmphic ~rtl~er (l}1;)i;c~l~y a hydrngen-cQ;IiI,S..a;ddi:,mphklls is dependeu:n on~'neel:eclronYo~,n.!1i \ii,'o;r:k~r.~ :~nl;;\il1in l,he hive'. ~r~ cQ,Uijsumilmg ';lct.i\~li~~ofilS pl~I1I1~:ffi a~ h~pr:im;Jll,'yLerm~:n!,~1 elec[mn SiJlk. docile. '11l,tlI d.isplayl,e!:ls motor activ~[y, 'liI'I~ ilIUlhorris: ,U UgEl 1l\a~,aOnuseus lilflstlluli ,oif TeCinmclogy. where~s older llJjt~rds f~!ldfomg~!"S lerlve IJ~,e C~!f:'!bfidBt', .fi!IA 02139. lJSA.lE·mait delcmg@llIliM'd't! AJrhfflll,gh S. (ftidlin'phict.ls !c.;rl!l!'Olgnrw

r!Cra.b.:i,(i~ ll:i. e. C.i!:l11 .Pcr_~.f~t. 'III.'.lld.~.r. S1,lUlli$lg m~,th3Pl.ogell or suW~I'>lreducil[llg bae.. F physlc{}Chemi~<Ilc{mdlno 11S that ~C:I:.ium) :iis <IIrmHiI~:~o eonsume .its metabolic ch~t~:~enge the 'iftry Coon.!.:: ofhiolog~ waste IPlI1Jdm::t~ (3), SY:lmlhiotl:c hydrDlgenl i-C!ll srrueurre fI!1d IlL1f!CliO!l. l!tllii.g!bh;J!n; of r€:!'oO\"a1sh~lltsthe ('ht:!'IIl!k'fl~e~IU~~iMll, Iher~by extu:me lempecmll~:lJIes, pH'S: .. and salinities, y[e]d~,ng SLlilici.enlm.e~llIboU(; free elle'rgy for miercbes fire On(lO 'Ihe sele hn1ln:abi~agU1'.Bl~1 S, actd#roplu'cw;, to g:I"QW \ii;lE1Ui;;i $i!:rl!tt~(f!nJj;mierebial .~ifc >I.I:so c..'I;~,S,IS; <'It another ~ype of olls,ly feed~lng it's, S:Ynllu'ophicpillflrler w~.I:h i:':':ltr~m~: under ('cmd!i,tkJQ1i> t~'lJItyi.c-Id 'balLCly ~l,!rg)I.¢'h:h gIft1'Wn'n. s1,lbs.I.~~~ Illij.ke hydrog~l1J ellilivugh flTeie energylb:w ~eU I'I1II,rillliel1<Ulce. Of' :Ibruml"ni:!!(see 'rhe f~gulte). A.~oiLle. neithi~1: I~lnd~, gro~:vth, lw ;fI ~"OCIU ~Uldy OfUl~ lilU p~u·tn~r'c~n g~f~""W{;'~~, bcfh:o;.uQtC'1i~ljjJii:ruhiI(:AA oil




Hkti ,s:~~rch $~Igal!'.)i. h CO':UiIOJ" ud llisibu If dir:Il[C.Ui';:II[ (~-;liIill1yl(if~genes eillomlin~ "m~cif.ls. that hydlO.ly~ starch, Per~\i~ps .it not ordy feeds hyd[()g~l.l. t{) lllhe.f d.o"~111stre<um !'jyn~ troph:ic partaers, 1[)lI,Ualso ~Il,,>;sisffi :ilSWt::rtf!,(~nUl~ ri\,·elrneig~.llbm.1lfunher up tile anm!!mbic food
O~l C(lml'l'[)U~.lds,

e·hll~~. i!li! ·t]~ei~linhd SI:CIH 'of pu!l,yj;~u~c~~a~id~
~~ydITi~}'S~!thi.\i.eo\dd h{;;lp'~o easure d~O:'ltl~'iuy l add:s-dJ!e p.refear.red gllrowtlia.substrate S. (:!:"Mi1rophj,l;itor-;n~ ~~~pp]ic:d d~e !(\iotste as prod lIet s 01,' it s :stlg311"~ferme:llllii:Ulg co~,orts.




nc r= H!I

Sud~lKll~Lltialfur 1h~~Jbo:~~coo;pemlkla.l bOlh! c

and; dO\.\fllSll,emn in theanaerehie





[ood clUlin.el1lphm'i7...~~ll~B

perhaps m.o~n m ic.robial met ubolID.cClnd biogenc'h€;mh::allr,\lm.f:~Jrm.'ui 1.111$ in ~lanln;)'
llleq t1i re !t::oQa.'djn<l~ed,sJi'llmergist ic i.nile:Ii:::IClions. w.ithhl :~aic:robbi~ COmmlll1il.ics.

Adf'l1osi[f1! trip~l()b'JJh;!!e (.ATP) is d~e eom!~[,eI1!Y ,c~l.rl"!e~ey IfI~U cellular UfiC. One O of lhe!ij,lIic liOI:iS of ellergynle~a 001 sl1l1l, to i is
lI~lllU 1,o'111St]r(:

WillloL!liI.pii!lrtnl!!r:ool'l~o~I~- -1H}:"".' 3a~~ll'W" 0+ iiU::O~- 31'1-3cfl.1 + 'IIIiUi p ..ltii~r, II ~!W~~·-+ :;lsQ.~ + 1(;1!1./)'" :u .i!~~!<It~-+ ~IHlW~-+;lIfIS-+!lW

lJ,(j'" es

~(i>. =

a:~ad~'!;IU~I.te s'Urp~yorl:~is 'biQch~I)m.~~~i1Ii li~e], S~lJPfisii]l.gly,. ,r~,·idfrr'Hpllj'~·"~,'i ~he 51,. lao:;;l1\:;; Metabo!~l(; coope;radOIUi. itlYIP0IilE'IiiGal e1.ec:blt:lrl tran~:r~rb;erweeFiS. fJddi({Dphfws al~ a ~UI~~iille·redudI19 !J<K:'De". ! ge:lJies. for acetate kinase, O.llC of t~~eeennal ~olllM'rIIn~r d U:rill~ U1Carmefiabic Q.)j)jdilfiG)~,erl b!llf!zwl\!.lh~ s:<h~li'!lollk tfil~1l'S ~h!~ krtmpecii!lsIJJ!'I5.fcr of ~ledliins el!it::yme,~ ~m:u~f·(lbesIypically use 'to produce (red) l~m'i1'tlile :SOblr<:e f1Nill(l!<lf1t (belWClat~~maothel.erunil'l!il l m:idallil (:'ltllfd~cl.. Elfidi!1lilt fEN'Illwal.or hy.dmg!1li11lliy (I,e ATP V~f1 II. substrate-level pi~.ospho.uyhldon. $i)jlhll.~i'!tIdu:tiii1i!j acWial pa rtrler [1iI(iif)~~ Ihe amount ~I rree1e,f"leftlY a\l'aiLallJ~il'l line allM~bijc,oxidat'loFi ill b<e[il· b HO\~\1e\ier.thepresence of nUlelil.eitlt.$ that code jlO~t~~D.Q(~~tli!·.IRi~r. ion~llr@slQ(ol~ing~OClr(l.fl!~r"n!port (!~m.ll~~ IFd.~~,nlNjO!l:ij"; H~~., ·twd!mg(l'!iJjJ~~ EiC.iIl!l~· litl:r ~~le ellzyme i!l;c,ely l-coeuzyme A Sy:IiI,~ mm IlilfiSjlOn chai n;8.G"'" GibbsilFl!ll ~IiI!lI1lV under ph~IlI.ogiGd sni3ll!dam S't<IJ!! colle'iliomii. [MaWt!!!rl~rnll1lU.. J}l[ 11D,I.;lL~G s~~-Wl$ i:h~l S. ~t·.f:di.~liOl!J.hiC'lt.~ clm~p!ia;.t~ . •1:111 m~l:enl!(iJl:i",e reactienalse used b~ acetatemiml~1 a,,~dm:II~N (~i~e·sll~firuleamll1littme), <lml the carl IJe 'OOllple,(~to the lieductionl, of sll~fiHe formi~£.An.,'h~e~!l!ld by ~!l~~nlbic Euka!"y.a 10 (6--'~l'. ~t rce~Ctjfm lhOllt no $in~l~ speeles ·~'1!1 imlig~iIm(]~~~ di"~'n'ily ~fluit'~l:'Ibi:;l1 ph}i~r.llfl~ies.. IJ:I,il1(1U!J;."e A:TP. IVlk:l'ob:i:'li enel.'ge!ie ill:llen~cti.om. ;Emep:~o~ peili'lbfrnl"~o ne ..Ah1'n.ough tll ese asseeiarinas vid:i~g! ~mrol.rli!I.lJjlil1sigh;l:S i,nl.o lilg.~~~ kmbes m depend 011 h'nte.factuous. !ha,t o!Oc~n· il miereO Th~ S. (N::idifnJ:{Jhic.!lsgenome sequence scopic scs.• les, ~hey dJivei:ef;]gr.,rIiI ( ]O~a g) can e....tr.:1C~ and glllJ\v m,illg V'dl1i:shiuggly sma I:~ also provides evidence~~r various mechal nux~ Ofn1i!::t:~\iillle aOii1I\li1~I~Y 3. global &c{lI~e ~Imm:llts '" f~'e ell~I'gy,Tin: deli!! Is ofl·~.e-se Q(l'n, nisms to ge:ne~te 'CCU~~llLr em;::ugy PI)tl;:tlti<l~via (I reae ~iO~.l:\' how they ,rweoplim1nz.~,d ~b[ effi.md dep~lmd.ou cl illere'mlil types of i{)Il"W!IJ,l:skl~'lIim.g pumps, 01' ~M). Sm:Jn:glob<lllf".U'l~;rOIil~~;:iti;om, ~pe{;i<ll.1i.nl.e:lFen is the pl'51l:noo a membraneciency through imI;terspecies iIlIlerm::li.olrns are .. mi.crobi'11 C:OlllilillUlni[y in~ei:"actions. inc:l:mBill£, II;.li,!\.t!~ .l;p~cit'S bnUl'ld i,nru·I:I".Jf1I:;;!.OoI;:'(it~l,g ('f~tnpl('.; ~C!.lned njlf.. s~i!lrrr~phr u.nlimil~el}t ·t;I~e'~1;; ~nl~ra-cl:iof'l:!i.prfl- we-II\:i.I'(lrt1h l1Jn:d~r:sh;'III1:di!'lJg. A~ (!~I~ 1<IJ~es<I sober [00],; ill. dw.im:l.~]n.£ hydro·C<1Jrbon tYI),e) lh']l is lin I ike those ifull.UI.dinm.osl vide the cheeks and balances tlrat susiain ~lilIcrubt;,,"'" \\'ith th~ ~);:ce:P!i!Jn (JFn~1m.ef y~nnl- gl!obal bi(lg1;i(lel~l;,~niCllll)id~. ~ (lra~I"';Y !'iupp:~i!~!i".\ ..~ m,!), \~~~bia~ .~ few Il'~'· p~~i(lllni.cr{lm·gumlism:!!. M.dllu!·meyel al...pro~ sons in cne'YY en"~lciertcy 1:0 learn fmm IniCi~ Another i.mportmn aspect of mierobial pase (.I ~ ·~l~IU I:ihi:l!:omn~~;I~'_!( ~hG: lotcut h!l,! s.)' I~U"oP~lysthe p{)i~sibm.t.t~ 3 f]ptud~d by ijl:ll uses I I bial S)'~1n'ophit:' pmeesses, en.el:~y :i\tQood il.1 ~.(]\mI. ~r!ldielll~ to 1C:~1l pie the ~nhereL~t moCllll.1~ri~y:Ful?' e.x."mple. ffi<lill)"synOIKidaliql.Llj.o f N,AD H nlk:~~tll'il!i~1~d~ i ~'ide"flJi 1~ lmp~lk h~d~ogj,:mpredueers ean :~f)[~1~r wi[h ~ Rie-rel'eftI(!e5; 1. M.J. Mt!!1erm'l,elol .• PI« lIl'al'l.~~d ..S'6. «:SA 'lQ~, dim~c~eolide.n:dm::e(ilj)~()!I~~.e reducrien of l:!ilhe.l1' hydmgen~cculsunli·IIiIS meth"rlo~elils or


",70.6. tJ~mal b!tIilUl.ilI!~'/~l 'b~<I!:e'-



;,!reroq1[q.ir.;~ for critical

..~~I f'jtl;'~du~~r~. d~~n.di ngoJii!. ~h~~!lvin.·I!l~

bi~lbYi.uh~lLt::re.]ctioiilli. The t"hllra.c[edz!lti.m~ of :undcmbia.1 S.ylfll,~ trophic i.['IlLemcaiens hots p:rovided impon;;;uu clue.,> <IbmLt theffil~~t<lbo lie inu:rac lions Huu dri.W!mtlt1:w;:rand energy c}!Id i~l£...e.spec~I~.ly iiii i:II1...emibi:; habit3ts. One ~tI:Ike=:t(;Ollm.e ~le!J.SOIl 11m,. b~n thl,iiimp0Xi1:Olm::(.;; y,f swdyi!lgt1li!lkmb~ml in~ef'dCliO'!Iii!S rnixed c WI,ltnJ!re.!! i.lili ~ULt~~1l elHli~ the 1it):i[Umle~(,. iiil.;M]jrJiliottl m Ji)l!:rt> cl!h~!rog" 1Vb:l1J)'



x~".Armthc;:r ~~..umple is ~lro\·'id~d .

Qoc]dlEltioJ;l of methane 1.11 fif('shw.'H:er ,~ediml;lrll:S.which appetlfs to be coupkdl to ~llitin.Jltc re:d.Llclion (./1). Very dOlOe

by the aaaerebie

llOOO f.~OiJo7h 2. 8<, :s:ctJ1~k,An~t1t1~ ~~ :~c~ 811,257 ~:20o;z),. 3. i;f;, [ ~<!~n.. 11-~- !!hY~~hir.!;j).i111t s. 1[an!!Eif; (. It, VI~~ M,]I, IMdm~ii1iII!)';.Jilr!h.MiLtobl(1.t l:rl. 'tol ~:lt99'9). .

>'I" g, ·E.l.!l~

re!.:i~tive$. of mll.u-illl.e menffiu!:Il0tt-oplIic A:r-chae:l
o:.... :iduz:e lnelha~.le
illl Dh.eseli1li~b~!lllL'>,bUll iIlfs~ed ..


M. k1!M!nerl]~!f.l!IaMteo11~



of~e;)~~:illgtlp \'Ii~h 1>l.dnUit.:lrediJ.tX;n;. th~w hm,;~ cOl.np.~ete.ly d:iliferenl :SYI~ltmpbic ·p,artnels th,u oodtqoo :iililn!~e (Ill. Tlis f1~.KibHi(ywilh 1I~1C'lt!11b 3:~l(i biogeQeI,lf!mh,::",Jm~.1I,l1isfol.'1Iml.~ r-espec:t to who J1<lD'ltne:rs·wit~1whom Q[I'lel1Js.IJ~:I" ol,i~· t~ons.\'I'ili onl%cCl~r i.n ~beCOI1iI:Gxt ofth-e i]1:t(;r~ w.udlc W<lngc 'IP(}~sihle l'edl,h;:t~l~lts~md of (lixi~ dal1i!:Slh1lll cmlibe me~<1tm~imdl.,nd Ofemli'~f10na ~lIec:i:es ini:er~t~mls. Il'JI .tmpbic l~fll~ractrons b~'t\~i$~ .!fI@rine !n~~us in w1:1ich th~~ i!ncr"lct[oos "~f!OCClUT. ~ <In;h<l~tll. methanJD~HJ'piiI.s .nmd: sulmle"ire~ku;;inil@ lih.!: oentr.d dJFu.ven. ill'tllhe {lye.~lll t~,er":nDdya ~.;t(l!~{l§iOl!,I;hea!l@~oobic oKidOllio!lOf !l1i(ld~(l['!e IUli!lllicca~&1:f~i~,ts. ;l!\f;;lilobiUty Of($lI;:OC'f'· the

5. B. :s.-dIll~k..i'Ilia:aliN;!,l. M!"~ Bltt.!. 1l.ew: 61..26,2 U997}. ·6- ~; B~6u5 e~ ol, lVlit{\!t~dQij!, (i~3;!200~l '1. 1(. IU. H~rlridlu. ]. III Hqes. S:. F! ~v.a" r; G. iBlt'W£!~;~.IF. [l~.Lo~g\W~~!.I1l! ~96, &Iii! !l?!l91 V.i" Orpham, c.1'I!, iiI!;M~. f'{.•• 1I. lfIiruklts, K.V\


Md<I!l!gll~, £.. fC•. [)IiLi:Ogl.S\:telJt'~ ::l:~.~,.q;B4 ,~t!l'I,ou. ~. v •.!" Orpnam, tIHl ..HilllJl5lt 1!l1ll.l'I!inridK, Itt Ill" Mda!~llm, E.. Ii. 1i'i@\~CfIg\ PfOC.Nafl •.Aan! Sa..(,I.sJt 99,



1663 t20QIH. JlIJ, ~. S.IIl~I)~~h,.O\iemi. Ii'.~ liU. ·1Ii6(iQ!!I!m. A. A. RiI!il~cu!bil!~~ffl!lJf~ ai, Na!Uin~440, 91.M2006). .






Human (ytoilT!.eg,gl.~ovifYi'S,'~xp!'l~~'Se$ no ncoi1Jin91 IUJliIs [h<llt. r-e<p:roes'!ii ho~timmuU1e r'e1i:pon,~l'~ ,(I ilinng i~fcUi(!!lJ,


~<I~~l1t ]Xln!j ;:.;h~iI1~il~fed~()\~....AJle'r ,tli iq:aJec:ting a host cell, a hcrpesvir.ll, ,DNA genoiue enters ti:~cnucleus \'1 :her.ei,l.b; Unit'scribed to make both Ine.~se:il~gerRNAs


!l';rp.:sviw!;il;l!l. are pfilhogelil!ie DNA \iirm;es. that CHIl estaotisb leng-term


M:t'i.rmmdsare W'm~


(.InRNAs), whi

ch encode pmNdn:S.f1lld


cCldJ]:~gRNA s rue RNAs) ~i) (see the flgIIJt"C). Somc::' qflh\;,:~ v i:ml !leRNA;:;; ~!r~ I(),~g
I[ mQ~ timID! I00 aueleotides (:l1It)j .\'\~u:'I"e"s eihers, ,t"3.1~led mh::ro.RNAs (miR.NA.S:).I!11"e sharI, (-22 nt), Ahho'lug:~ q:!ll~RNAst"1l1'1 selectively

<l1I.1d'ld.1I1~t~'C· ~ntiviiW'J~ defenseml,1.ech<lIl.1 isms. .. hlii]i1,Ue ~~Ptlll~~:;'. such as rhe Pfl.1gflH1Ifl]i1~cd death {3poptDS~,S~ iH~fit':ctf:dI cells or tb~ :pro~ of duct~onl ()f imeli<:nl':lliJi'i thilt ."cl"i,\i~te ifi'inriIHm~ res;peo:~ses. are ,oute.~i~fi£s.en..odby molecular derermi n:fllnl~I:~:2l!tm:ciml;flri;ltrlD cha,ractcri:Olks of '1!X1rticu!nrp;<ll.l:iio~en:~. Ad<'lpthre immune

a ~ng~ of h~!ill~t~ d~il:th h; iijdu~l1id ~a.Wyi!iilU~·~ vlrus ~ eyd~. if~ Despite ~:h,e:se i.l1Ilsi.ghlS. we are left f~lirh
f1~!lIiil-:rtlU.:i Ilerp~~vi ra IIH.;RNA;o;,.indud



forw'hrich no fJ'l]nClio]1l15 are k.I.lo\.vn.D'[l these ncR.NA.~ prm~lQte "ind

r1Cp]icaluon il:~.d/or pmhoge'm.e5is·?

l-lCMV ~I

in cn~~n'!s:r.wcogn~~.;: alow~ viml

3[1!igenS:.!meIJ.eF;:'IJm.:y prolei:IDlS" tim! are I'ImtlnV'.niii~

an!.iml. Ile\.ren~lci~ess. :e foreign lo~1teinfeeted m org<1!llh;;m. 1.l:IIltnte rel'>'flml~~s ilrer.JpidallfBd em]

at. and Stem-Gincssar

herpesvi rus [~!iJ~H C~11 cause sevese d ~ sea se in l'Iewbor.lil. infants aad li.:UI. ]mnH,m(lCOmpro~ mis~d lndividuals. ~~l!cod..,s <lit J.~~lST n:\ii() long neRNAs: and 1.1 miRNAs .. Reeves et
Ill' (11.suggest Hl.iU of these HCMV IlicRNA~ are .~l:Id~N;ld

Inl,~(tl!d noSi co l[ ~lI'Idvld"1:I5 ~ur""'~


S h'lllttiillil,g ,d~WIIIi~heil!llllllllllilrul! ~leSpO:l!l:!iiE!'. Inriet:tio n by HCMVindilHJeSa sla·t.e 'o~ moetahCiI.k stre~s. lhil'~. wo·urid flo!ulr1'i'llly nigger Itos~ imlfflIUIII'I.e< I'IeSpO I'IS~S:~ft~!lJdiln.g(~[[lly:si$ 0 r <I.~QI1I(l$is. H eMil pm~rmlS.lhE'.se 1mmulne i J~SP()!lSCS; ~ilil~llwCl Vif.ll. nc:RiNAS. Th~ !~2.7 Il'lfUU\: bil1!d'~MJIKC-!I OO:rrlpo· ~2, J), t~JC fU!fI{;liou of ~:uost;v;~r(l~1 RNA.;':l has been ~lncb;:,n:~l 'm~5 I:L'C been sug!l"e2>~ed th.u "ilrill nc::.RNAs migln

Inh.ubil gene

reduce virus spread wbi le the :.Io)\"er. bm often :mlore

aents 1Irld Mabi tizes

If!ilO(l1io ndrial. '~ll1ie~l1/ rodlJ(li®lll; p

mHiHJ L112

illI~ FINA

blo cks,lh ~ I?'li!~~S5io!fl (If (ell sllr~~c!!l' MtC8. ~!1d h(![1(~ IprCyeliUs< immune ~e:1!lshat rtC;ognnlc iI.;IO. t



etl1kuD.ve. adll~li\'C responses
k!rem(Ji;liiliit-ed" :Sllc~~... fu~ pfUhf.lg.~mhm'e nlh:;~, ~ ,evolv·ed] :1Jil.lecl~allJi:srnlsdl'J.t 31'treI'llUm.c these has,! ~mm·lml;i~~m~;~. Although mnueruus v~,r.dly encoded inr<;let!1!gOlSl i!n~lnunl"'·ifliisp~~ i nsein h ibirors, \Vitlmin hours ofHCMV illifeclio:m:. a -2.'"j~ ttl viml !~cRNAf.ap~lY called [j2. 7)flc~u!1n~·· 1:lIles. re'2I:c:~~il1ig ~20% of ~OI:<I~. irsl RNA (8). v t !nt~IlJOsUflP~,st>,ivc 1~1.IOI:(.':i.l.1J!> knm!l~l.rr:c.gl~.lm'- lRiIlC\lcsl!UII. ir~p!nlhat ~,;t7'binds Ull;)Olnpoare tory ~!tcRN.A's o:Aer viruses ~ver;)lpo~:el'liti'll nelll~.s Of~l'ill~.rodlond·fial resp~J~ltoliTy chain i)dv~mlltg(]'~neRNA,K db ~!l1: eed h'i be transn eo!u]p<kx I (M.RCC~~.) al'i.d db~f'liby iOul~:ii~i.zt:~~ lated aad C".1Il therefare <let ,,",~pid~y.nley C,In MRCC':[ flUI'IiClii;on. [nfec'~:i.ml.of ce]l:s 'i",,:uth an !!](>(J' b(,j' qU~I~ ~m~~!. \~i~~ ~1~Y be ~,!r.!IJ1'IO~~~!~t HCMV m~t~:<mll~cl!;ing i~~.d~.~~s relo~2,7 '~[l~ ~Trom.lite mhodu:mdll'ial.m,(:lil,l~bl<lnt':. resu Idng re:ilp()n~. Ghr~ll the pre(:;OOemofce!hd:;;~rl'eg~ irl. rt'eluced energy [in l:lefoFllu of ~~derlO~irlitl ullmolfYl1cRNAs.,it ·usllliCrelhl'l.': not sUI'prisillg l.riphmphaN:: lATP J] pmdm::ti!Ol1 Ilmd lillie thm herpel}vinlses. encode both klR[g :iilCRN!A.:s intdw.;I~:(l1ia Oil' apeptosis, Stwiki ",gly • a Si:L~.l il~:r and nnjlll~.. . i\llA,~.1.1).. or e.x,Jmple .. Eps~ein~Barr I( F effect i~ seen when ceUs <~I'etreared with virus ~pr~Sl?i'l\'Il,Iolol1£!, neRNA~ th!!!it irlhiitlir ro~~fl(ltl~. Ctlmpo~lnd u '[11mi!'lhlbi~s MRGC'-I cellulae ~J1tefferon res'~lmlSes. <I1~~DU!:lh the '19).EJ:p.res.sio.~1of ~\2."1'~~ w.il.d"'t:Yl~e llChllV by l!I'd~!ly:iI~ 1I1~~llt!!l~lSm~rm.li,fItt';~nc~~rv·'). A pr~vlOilu~~h~ relel!;:~ih::;Hk.'!f'I of MRC-~ miRNA ,ellleOOed byh.erl)es ~.illlplex. v.~rllS: I. d~;ml."ll C'MV inWeCfUo.liI, or l·otenOIlit:II."e<ll(l~1 S.V~~.)p!'ev!2!11ts~pop~.O'$~s.by b~.oddng till;: u'lcni .. A.s <l res.u.lt, ~2.7 m:lit on:ty pJ\.."\!'e~.tsd'~c ex.pJiies~,()iII1. f two cel~lll<u pm~e.ins. SMA..[l3 o ]n'em~t!u'lre ,death of infected 'cel ~'s;,IJln <'Ill so andllmllitSliormirilg gro;\"ft;~~. ~CI.or-ll.whic~l call f (;f1s.l~reJ>·lhe sh"lbh~ pfoduct~oJlI ofAT]l' durill.~ lr.agger .<Jpopto.sis U)..i'\~~optQsi:> C<l1lIbl: m.l I~.elilCMV :~i! lief.~cle.

respm~1ii~t3)- Evidence ~ulP'po:l"t.ing~:lln.is ]Iiypothes.usis: no·w pl'ovi.ded by R:~~\l\~S€rt fJJ. Imi a IfGeemi~~lt~ ~~~'S~·il?m:·.r: (4) and by StieWiI~.-Gillossil:r Oil !)ilge: :3 76 ilm.~ll~.~.s is.'>ut::(J)., Bath sTudies :s!inrn',v lhm h~n1iiHU1 ,cyw~ 11.~leg~ r !JI;':l. HeMV )e:x prtt!l~l;'s neRNA.s levi { th~l ia~~m.vi~riL;\lClil;d ~i,J'US tJ) evade ~!IUUU~ im~ uume re~.po:rtlses..
i~J)SI imJml~lnle


W here a s dl~ f~!lct ian (l f me] s~ C\;ll~~111!1' reg!Jl.rl~»ry lli:.::RN As is ijlJO'r~y Ul1:der,.;,ulod. ~ollJg naRNAs are knowllJ to direot ebremesome si Ie:m1cil:lg.eg l~1 gene ex pression ..or r aie ~~la:~ffit!ti.'tirnl. chlliOmO~,lInc unle~li'"hy. 11m. ~ddlilf'(}ll, etlktlryotic (1e~~:i! express nemereus nuRNAs th,1,1ytli.de a r~.bonucle[lproleiu complex I[the·

given lh~c\l'il~:pHcl~i~ ~f\'i~::'l~ gr:=ULUl1iil~ RN~ !lire <I~SOpoor!llIrg{!'I'S for im:lapti.v(:l· i.nnm.um:

cal ir..~ti,lJn of M !lee-[ I;,'{IJfill~J't.U'l~.n'[lj


RJ'>.Jif\i !~dw::G"\I1 s~[~!'I;cjl!,g CU!1l1!Ji'lI~x(R ISOC)1


~,nRNAs l.h~1t l,isplay sequeaee ,aoJm.lp]eme~.mr~ c ily ,13).,. IUSC reeruitlTht'!U 1;;'11[1linbibit mRNA : '~ Ir<llls!l!llklrl QlJi [ndm.:c· mRNA degra.d:,,]lll.on . ~ o ~ ~ Th~' ~ulhtlri;~imllhe~m~er fo~ Vtr~log,:v ~mdD~p<lf!m~rn ~ G


.~ Mille.c!illn 'G~metks ilild MKlr·DIl~oIDg~ •. O.Ulk:eUni!ollmii)' i!:I Medic\a I ('~IfI.te'~',[)u~hall!1, roK 2H'G7, USA. E'maill,
.~ cu~e002:@mc.,d~U~e.edu

[fTI'r(l:r~~Jmthm;ne deli:fls>ern,f;)cl~llnismli

f 'cell

Mell1be~ ·ofthc herp~\Ii O.lSlha.llUy encod~


!l~Ii,!ll~ro:uS m~RNt\5 but.w~i~

the G'-;':;(l~rli(]!~ of

l}iSis by mlluml kl Ire!" oellf<;This,




one liSV~ miRNA ~i). 1'10 frmc:tioms ha.~it::' ~ bcr:'~, deseribed. Stern ... i~ms!>:H ei (il. noted thi'l~. G oae I:ICMV luiRNA. [l~iR:,~UlI12. di~~l!~;}yS SU"ltL:ellce cGm~.~p!,elml!cnWIl"iI:ytn the ;3. '"m~l.m!ms~ liruu."<!, region ofcenul~r lli1l~RNA ,encoding MHC c~a.~!ll-fdmoo dl~i!l IB(M[CB),.~ lig~t1]dfilf ~. re~eptor fO\lild (]~ :~<'lu.tr'llki'II.;~r ells 'of ~h~ e nn~j,mllu:' s:y~tem. JVUCH e1li:.p:I.es. is; nOfilmlilJly s~on

p:lli:si:ung ~'1l"j~e the HCL\lV lJL .~ 6glYCOJiliro'tei~1 ~I~so .illhi~1!il!!MIen eel l~iSUrfmJc exipres.<iionby Ca\!SiBll(gi~~~nlirO~ce~~.~~ :oeque.s!ralion (II },ir IiiI' dlC):'refore :llppe'an; thm [I!CMV has m,:ol\ ed 1'\,'0 d~Slin!l::1.~1IiI]ech'''Ini~1Il.1S ~~ prevent cell-s,~.lJl:l~,oe

'~rge:~s for C'h61n(lt~1~~!p~l!i·ti:c i!~10'r\~lli!d~)!) iill berpesvlsus-induced d iseases

:l rrl, I'.. (onlO!d ell1j" (~dSgriiigH(tfl#~~.

•~<;liv.w~cl"'h(;1~cells .~ru ,,~~bj1;)cted severe \ to
stress, inc~,mli.ll~virus infeeuon, Cell-surface

M1CBn~<JIj',k~ Ihe~ oell:;.; ~'brd~tnlct~U:Ii1I, y natb 1LI1I<1~ ki:Uer cells {I0), The authors show lUBa~
HCMV~~,~,m:~UlI12: inhibits Ihelml'll[;latkm of MleB mRNA. n.le:D'e~L!hi[lg:llli..'<eI,1(:e ofNUCB
pll'Iw.in pf'(l~I;)\.I1~ ]-[CMV·~nr~cmd

("$11:0 ;)!g.'!~mll

1\/IICB 'I!.,,!pre£i"km-0!1~~1IiIi;;'tI~m~'t1a pmldn by ~!1;dthr;;oth~!' b,y iii ~u]RNA. l'ogether; these 1\\'0 studies !iugge:st. ilil....t m.1!lY 'o:f Ihe long I:'H.;RNAs .1ndmiRNA;;; ,elm~oded by hlU~liil:Dl~Il~rpesvirases ~~ay share the <!biIi:ly ,of'~;;,ilu:,!un ~lefJ~e:s",;~ S nOIi1l~lrueW lUL':il~ pr(ne'i:~.s to inUaibit host immune respoma~s i.I11i1i!, hence, aet as p21lhOlJelITlid~)i' f(low tors .. Viml ncRNA.:s theretoee clearly merit ~~(I~ASoan~id~~ol:~km rOl~!Uk~~ly a.fl ~~ur;)cli\l·\;

2. a, ,~, ~Ietl\ N(tl. Gene1. 3i'i11 il:5 (~OOi$h ~ ~" ili. P.ihl~~ (I1Jll:1~ 2~S:],(:;li)!O;1I). ~. M. lB. Rf!Il'i'~!l, Ii.. It ll)ililli~~, IP. [!,lj~1I!flr~ G. lilt S., W!Ltim;c!1\ J. It Si~'!li!il:.S.if,ll{~ lJl6\ U45 ~OO<l~'. !iI. N, .5.I.t!n1~GrR~;I1tf 11ft, S.r:iM(,l 3:1:7', 3M ~~OO;l• 6. M,]I. Clfffiem~., 1.111. J" ,~. (ei'J 8j"a~.38.164 l(2iOO6.t 7, A, """~~ ~1~,I'l'~~~ ~J!~.8;1:tlOO~1 8. I!l:. P.WSli!llrJr)l otl.~L. JG'tn, i!itD!..9i\l1, :i!5,1J[1(21llO)), 9. ~. u el' a!. Jl tl'iaL ~7,al 8516 '(2)00)).

BIIlI.n,ln ~:mo~l.


1iI{Il). El.;iw~r,~


c. '1JItm1l ~

!il" S~

28!l\ 7~~ U,'J!!I~~

a.!..). Eitp.I\1M. '1'111, 1.-1127 12.0031.


Prima ry e Ria C!O nla,in the ~t'c@ptti,r lor' a ~~(Jleted p'rQ'~'i n (r!,u:l;a I (It)r 'cell rl;gti,l!1ltl(lii1, strengthen illiigl their potenli<1l, as~i~ln a llil1g

centers il'! the '~~lt
,ny peop~e are tI'!I:n1d~.iru'with motile cHitJr. the ce~ilul<![. llilvdike projec~an,swhese lbe'iti'l:JiY:~euer.lte;sn~~id 110"11,1 lh~( .~n~(lI,!;~s pa~t.id~ frum ol!.:!ro$pil!'a~ r wry U"'dC! O[ help oocyl,es~ss thnJWlgillliner.al~ t If~ll]aillrub~. I)r~ci3d~!lis A nnw ~UO\:\I'tng fl1r' primary cilii<li .. the n03ilmotih: CO'llIIuu:rf)art,S,. p:resf::n1t us 11 :singl.e ~Olpy 01] the surface of
11.~1'(1S~ell type::) c
illiA O~~lt


i.Dlg~he e:xpoossiclJ) ol"spt':d!lie gene$. This g)en1;" pl'Otdns [I:ela.ted W Iht: i~:nvel'mbl'ate' Hedgehog (llh I proteinJl~;Ol~ bnnds ll.) tbe tml]$~nembmml~ i,X,d iiCnelJ1l'[eis\'ve]~ ,esl!t'lb~]:lihed~ ho~\'~v~,I;lm.er~ proiem Parehed (Pte) allhe eel ~slI!r:f<fCe. Upm:i .~re :L~~<Uly!JIUll~tll ved <ilue!itrol.1S O!ilOllt lilm.ere.g!JI~ bb~ding. Shh :l:bo~~s,h!~sth~ i~hibitory 'L;lll~ct lat~ i!1t()iI'<I~lifinbili:~\,~'~~M,1¥~rnl!e tCO~~;puJ~61U:;; ef Pte on .U:IJQ[lu::1f l'l'lnl!Slfleilffi~b ran1.e prerein, ofthe pa,i1ia;\·,r.J:Y;' i:1 is: um:le-d:r how Sl:IIh·PI.C' and SnriU)~ll;hG~~d (Smn). Tirl~ relfuL:lf[mm i!1bbi[Utl~' im!l1i!rac'~{)~nC~l~~ Sl1l0 si.gmdi:~~:u::tiv ~ 1ty, .Previloos: 'IA'Ol'ikhilS Sh(w'·iiI.lh!l~, prim,my eil k"l :Il1owsSmo tD u".msdm:e a :signall to the' rmeleus via gHonm (GI,j ILrdns~:illb.ioll facb.OiIilS, t~iggem'~ :!Ire essential For S~~~,s[gmIHrrng, .M:ulkllioDliS Wil

bod}.!, D~fect$ill


~rnlilryi~is ~l~ve been ~~~Ilikeddisease . .and '11'0"(:. c [1(1 ~,lm\' know thaI; t'~~ie}'liLlnCl~ons mli(1]ue a ,U:lilenlll;I~1 ike S~ir.'lIt'n'l,re:s., p~'(Jbi'lilg the: eXlrd!C'e.l~ iul1ar ~lnd!Dl'i!il~):ltRfli m.()~e<,':Ul'~Sthill! flre f'6'C~ eguized by the receptors they rem.': Tlns sens~Hy nlm!Clkll'l allews primfllr)' cma ~{D ,ct1(lrtii11,1iMe. }lIInC1101l!> ~.nte.roeI~ ~ uhlll' sug Il>mil. il;l~Pilll~,~

surviJV"JI.~JI]d dif~ lierelllialiO:i11 of ceUs: dllru 111; t~.illbrYOllic d!;w,el~ OJlUlGf.!Il 'lirl.d in~imi~el~~U1e~of h~~J[hy lis~'l.De5;
l!Jl:'ljlS l'hiijt~guhlt(!grOW'Uli.

( J )1. Oliia'p<I~e J 7:1 of Ih is iils~II,e:.lto.n~llgi(!'j' ,


(1, f~rf~~~rdJilfi~l;\'~l~~ Qfpr.~!lUlg.'}I' dli~ i:~ rQI~ regl~h.u~Ii1I.g the rl;~pU:JI1$!i: f ,~~~U:; SOrlie o ~o ~:l~dgcl:mg [Sirl:~;J.:Isec'rot!e<.ll pmtci:n1, that con•.

stit m~ one of the most fllliidumeClJl,dl


II~ighty censcrved sn.gna:lli.11jgsystems iI~ verteb:r.:l~e de'llelopme.lill (3). R~glLJ,~,med movement of key p:rote:in S, :iilllo <'Imll,eut of the i(.'ii~l!1ml ceealt;'~ a &flphis[te~tl;;'d S'I.".jn;:'1:!by \";~,ic:hsells 'C'H~ tum th:is pO\\reliflll s.ign~~,illl.g ,ml:lnul off.

S:~ltli, \Y.ElII;; ,of lh:roe: p<!:mloSo1Js w~neJb<It"te:
5. '[ (1wi~~I1~n t!; at lhe IO~Fln'Uml (tf J!\oDl@rnlii'l BiolDlJ~ lhru"e.ril.y of Cajl~II1'h.alg:~m.., fopenh.agli!ii EJlt;-210!l [te:llm..art :E'!Il1lIt 5 _"eM. on is ~ml lhe Cdl Elirikig1 aod .IBr~lIdh. ~alilmil1, ,lmtL1Jlilre ,ei Child He,altlii< .. mill Wiumll!JfI Oe"'~klpnnll,j1Jt Bieithesda.rlm
2l(}OO:2, US!\. E·ilT:Ia~~ ~mI@M1<1j~lnjlt!101;'

A. ci~~'<!y~wiill:h. Jh~ ,Ilrima ry'~ilium t'mal1~te s ,a:l;lIl~O ~itilry o~g~ne!~e'iiro llil mOlil ooR~ i~'OI!J Ilod:f. l~leU} r r ~n '111'Ie< b~;nCJ! (III' Shll, IPtc lrran,-sIO(alleSt(:l the ~rilif1~ry Cilhll1l11 andl b.[od::s dl~~loc;a!:~.l!aU(1flIII Sma, TlI"iln$crI~ltlon ~ ry ( I~Wm; (Gu) ,~r~ dC9rMI~d or proOOS,i~(J t.OI roprl1sro.rs. (GliRl. '!lRi!iJi~t)Upon bi flIdilflg orS!h~t(l :Pi!ch!!hfl' d~i!!!m. IPll: '~eiI~ll's...ill Ad S mOl emers.. tha dli uml.Thf,s ~wiltd11IIT1i!!j! 00' {i(lIllIJol~f.ed ~S.tl'['Q15 rniE?J3soohom [be dliu m by mellillbraln~ by ~t(.G li liSp!'()(eSiS~d '~(JI01I1i! ~d~l;\aW fClrll1 (13liA). .

ge'n-es (lllCOdhll_g prfl~eil)s e$~efl~~al for d~h~
assembly, such i:'LS, int"'Llf1agel~iI'Ii lmnspon the: pm~e.nml~ (4}.res~I,1t:~IldysftnlCli.OIMm] Shlm S:ignal~ ing andsevese ,dtlvelopmelll;;']~ di!'lOllden; ~iln mamllll~i:!; (.5). Furd~.;:r. ~'!.'C'r.l~. ro11lp[llle~]!:!; of 11113Shh~ltll\V'oly speei fic,1 Illy localize liClthe li'Wl or I~~L;: of lite pxh1~~rtei lil.Ul~1 t 6-9).i!)dl~dh1g iGU If;;jlSl,;'riplionI11ielo!1; ~nd pro'l~iij~ tha:~~glJ1~ late GH .lclijl\'ity at the ciJi.~lrnllwp. ilLoc~~~Z:"IIliolill ofSnm ~(l,~~IJt~ pri'1IIl~y,r;;:~I~"m jm;;ro'ii~;S\¥lle:n i~ cellis s,tIJmill<ilied w.ith Shh {7). illidi.cali~g~]ll~U:


imp!)!f~~m fol" 'C~II~!I~!! ehelesterel lrlld.llcking (I! l Oxystf!n:ds m1li)' ilI:fte;:;;l, SnU) 10c.,1 iza.d(m. d.irt1Ct]y Or' .ind:~ooc:t~y. y Im'ls.k~ng or umnEls.k.~ b il1g ;)L ciliary
H~r-gl'l~ I'I,g

lhr.~underlle ~e Shh ct':lhd'(Ir S"\I:~l:C~. d.~re$to mine hoo'·Il.I']mflIiY cil iii, rWl.llctiO'J~ as sP'L"ICi.di:rod
orgmncllc. s thElt ~:nlegrnlt1! pa.r;~.ive!ln~.d .11I.cgm.ivc

i:rlPU1'So~.llrnn$cnplim .. fm;tor'~Gll) ••r::l~\i'ity. <lind d!l1mli1j~~of Snl!,')'. Oi!ffi.e11l1J1~tioo i~II~'ni!> demafu ~tl di1>!:.'rnreli ,~l.O\vthese mcdli:;JilmiS'l~lS ImliYinrc:r,lcl PJiieveJ1l"K ci~ioru Im:;"I:iIZ<lliorl ;mdthu~ Sllh :.1i,gI'I!.~I- \~':~lh o~:!ner~~i.;Iil::;~I~~ t:v~mN. such as \'\fm s~~r..,liug .. \vhereas a1l1otbcr~,~uun i(lrrm:~nClf'C'~~ I; i1- ing. i~hum~. hC;li:lth ='Iuddlsease, i~ryl~1 i~~tijUil c 'td {'ifS~l(ll a:iu! ~iPpgr!5Cgn!l!t~IUR!ef'ili!f~IlI(lJl,~ ilndNorb!s rive ",c:t~v;:lti.o11 of~IJm.eShh 's.ign<l]il1gparli.1W~1:Y (i. 1.). T. (_l!.fiil~l'!<'l~ t ~, lr,e,;lli:~em. Sd!!ru:~r. P•.snit; t 9)., M(l~~Q'Jl~r;lipi!d n~ft"'_mern h:m't'le f~gitlllS rrofjfc :81 97 ~2001t
enriched ~I1 me'mlls-ulJj' p~omole tl'il.e continuom; 'ci~ionr targ,elio!; ;mldlorre'tell!l,ion of speSl!l.C~1


ShIHnedi,,~led gl;:neml;i~nof active form5 uf ~h.e(i~w~T.lI,1'J(r:lp:tnon :ra~10[Srakes pl~ce in the
.izes IC~~lep:r:inmry eil jmn <mddun SI~:lill. billd:::.ro

,(,':~~ Ro~:atg] !;~ 'lll1rr~~' ium" show Ihm r~c !ocal~

ei fic si'yna~ COIllI,~m~~na'S ll).

asPre and

Pln.:wh~~lit i$i~ '~l~isT~lC.di()I1.[!l ;;\ilMJi[io~.~h()' (y'\YE!I~rols ]11 n~£pril'm~~1 d~it!ln

Smo, .~nrcspOlil~e ~oShh sign!)~ imlg" lfhe,concerred: m:ll.O'IJI ofSllm mo~eCU~e!l.,;lIld

~lIt&a.or:s show t~KU: the uuique Olm,d'r:::onc;ened I\IUlVe~~~ellil IllitC' out of: ,UTIdSnu) :ilmo, l!I~cIli~ of rnilr'yci'~i'llllll:l COl'IsJt~[!lIIte~ cel.~lIi1l<1!r Sign'l'l Ii rig ,<I !ii,~'I,!~I~J :~~'p'Q!Ulsive0 S;~l.~, d~c figure). 1 {Sill~ W,~y does [l'I:c locilliz.e to theprillillitrj' CI I~ ium? .~.f(: £~em! oLw~o~ !ld:'~llt~,gcS, T~£'i d~,i~lm p:Wl)\'id~s a m~~~h !SJ':l'i!ial~r.;:iI z.i'l,ufacG' mea O',(:f which ta ~:nI:egrl'u:e<I sigrn~1. BecallJ:.u~

ceUld~r S\'!.'itdl bGl;"!I'ee'1J1 ~cti\"<Lt.~(mand imlOl.iiwniol1l of I:he Shh 1)<IlJrl.MIJ)I .LU·iJ g de""cId n

z, 1fI~,Fl(lD~\9i.l.Milen~C;M, IP. SC(JitS!:~(e:H!l. 3n: 1200'7), 3;. Y. w~ I'!.. i!. !I.'~Mn, B, t, Mt~;rI". w; GpM. (eN 8i!<!1. ( ~~'. '1)9 (2GI:lI:n .k J" L iiij!~mtJ~~m, G, iI. 'Wii!iIrI~n. Na-l .I'la:: Mol: ,Crl'IIll!l.illl ~, :B:n G!QC1.2:~• 5- J. M. S~JII~!~,'It ~errs~(;], Cd~1,2:5, ,n~ titlOO6k ~" if; I~~I)'. (. If" [arki~~. Ii:, 1J; ,1!JI,ti~ii.. ~i!Ii!:. (~Ir1,'2,
II. (.1. H\ilYlI'.:I'fl ~ ,"/., !'loS' ,Gj)rll!!'.:I!, ,~3 (;!OOSJ. 9.5:. II:. ~'1 ~~DJ" ,9~ 8ia~.:ZiI1l7,3711 ~200;.)). ... 111!)- 'I': t'rna~; y, IO:~; liL H iR;il:~, NOl~~ ~;J<5, l:j1~



c, C~~bl! ~~,tll_,.lV~>!'!lIl'i: oli~;7.IOU! ,(:;)OO~~.


ami cO:Luml of tissue hl1imt'ltu.~asis,.


'~'h.eciUum p$'l_nrllIdes frorn the ~uailil\e~l b!!)dy; (
PIC has,a gre<lleli o;ppo.rwnhy to seil~se gm.dients,

Anil~~ogmL~iy. the proh~il1 il~v~rsrin,. '~l!.Ihr'(;halso Im;<l~ji2:eS to r:liil'lril:<1U:y ci~:i~ .. lbl1C~IDoIll1S it meleeas ular !iwitch bel\,\il;l~!ltwn s,igJ!ffi~II]ng p;;ttbw~~ cluntfOn~"d by th!:;:\j(;,lcM~dpJ!ln~i!1 \V~t.ful{jl:t~r lundamental and conserved :systelm.l t:liaat cenuols Lleve;,~opm~!l[ llJ). fne.'I{l ;ht1!llll!'l-!~ ~Ire { to defiae l~.Cpree ise moh:m wll.a:lirn.e()h~m.1smfl!

~2()On. ~1l.lY.I[jIii~ln.~, C. Ot!iRg, M. ~.!mi.~;~.!iiill:tiKhl,MJ'lr.t, C. kev, ,O!{I ,~. 8kiL 2;l~ lZll (.2000.~,
1!~" M. )Utigiij e1'~. Mdt Gen.ei'.l'1, 53? {~I:lII!)). ~l!3. Wf!' ~llmm'lol 8" '~d'''I''!I'~n and 'P. S.ollir fi!!f hill!pfilUoDm, L fiijJ!iIIl!! illiFil ih~ma~(jIiC"pl. .


of S.~,hnJilll~e£u~e~ ..:we;:~s :in rl!]"~l~etU' remc],\fed from the y:eller~.1 ,e;:,11 surface, For ,ex;Jln~.ple. du~illg 'carly 'L;'imbryo:uai(': d~'v\l~!OI)l~l!CUt. O~~ ,(II irected DlU)\'C~liue'mof ShiID·conll1lii.l~ i1'IJll ~n,p()· , pH11~in p~nid~s i:f' U'.oug[rlrt Hl bre~k le f'l-ri~hl
bad,), !'lymm,etry in vertebrates j I:Il.~I1~s seenaeie •.rolaJling ci:~ia. p.r~s.enl on cells b;a. a



t>f;.\gim~Cill~~d the e;l111rbtyolli.~ node.
l~n'1II'all'dftow of

ge!tet'tll~ ~.

nukllhal d.irf;lCIS, i~~(WCllleln of'lhe partidle!'l lO the neft sade oft~,l.ede"'dop-

LI,arnllilng INlatulr8~s,Way: B,ioslsn:singl with Synthe'l:ic INlalnOIPOrei:s

il1lg emrhryo. The pt1.ni:c.;~essL;Ib~t::qu.ellidy frn,g~ m~nl, .~nd released S;lmlilllml'ecl!!il.;~ Ke~$~d SYllitJl!I!~k s.ensclr!;· Wi!! rIlll:I'~E!wli!r rl!c:O~l1Iitionl e!jl\E!nts]111 are l1Ii3Jnomell!r"CS~iI~ IXIn!!~for se!~~tivE! dE!t:ec:lhm by pri:rrn.l~rycil ia, wh ieh aetivaresa .s.ignl;,'l,1ng i of p:roti!lllS, rU,lcleotide:s, ,gii'HI dr!.lg~.


the .apprgprhlJw


(hal ~!;;.Ip~

diefi ne t~U~~t!fl-I'.UgJ.~t Ibadypli!lll.
AJ!'Itltl)[~r ~lnpOI1~I~H
ql,l~i>liQ!1m ('{lnCi!r~:l'>110'li,V

~liI.eIwellJqI.tl~d tm'n1c~illig of Pte m»{i Sma ill.tp.ri~ 11IlI3f.jl,y dl i(l i~~OC:'hi!",,~~tF~r P~c. 1[l\i~~~~9fi\.~r [~, :



Rohatsi ,fit/' til,. den'lollib:strnl,1;! !h<it~ddWI)!l of [lxy$'G~*" whi~h ~~!i;jl~~~
milc!eijlll". hLlt

me :1ilI.illlil.omt'!~:er~:SC<l:~e c~eoLid,es. l'Jia.ey :~3W: done s.o by ~l~~.ch in~g 'm(}]oc~I.lar~.recoglll~li.m.l ag~I1J~s; 1M RA:» that p tllI"C.'s. t:.~a]~. :.pl1.0tt:i. cha.l1~h~:~Si .b.i'.Q~. .. ('1d n ~L.t':., .senmr<i. I~oli'e'J>i.TInple the h~ml~Jit<ll~:ed bind I:hese ~pecii.es 1;0 :s:ymhel:ic OF biologi,C<I;11 .. l]f~n(ipm'Cil, iOllli ~h~IlU1I;t~ hum~U~llIi~1 (II ~,pan.the h~!~.t: m.e:mnlbrn.lles of fU;i,,'e ce']~s,Hird ;Ire dl!o~d i~:i Tih is ~en!lins ,concept W"<IS first used :ilnIb.e
Il~U~· .c~)t~h'lZl!!,;o1,.mt~f.~,\'hkh l'~l my,lecu~'m.·-H;lcog~~li.!J~~ !'iZ\'id

b..ring systems

~)l;'jr ~~nM ~ ~[!t~'+ IB~I~~ \vhl;;:n ~ s;lX~ilk

do, the Sflme mo:I(!;~lllcs n~qluDrl1!dill, :ml.1,uch $lfiIH1~II!ef pore. ~U!I,d bi,oiogy I~mv,~ded an i,de.a;11 Shh :dg n:1Ii~. tn:ilIJI,sd.ucth:rl1ll. w~He l?iiCU Iso ca.1I1idi.d'arlJe.:lilre n~helno~ys~n" tbH L} ~!1]nopo:re. rermrlm, illlhe 'c.illi.t~11.t tliais finding supports (JDR'l:tioJ] fmllt.n: dilemi~I;~yinto it IL~.l~'.~$.m"jbh: ,o::~[C:ILk~ ~l '1J'.if(llleln 'tlmt fO:f~ns ~ n~HII(Jpo.lfe o ~:Iu:::'il:"Omldiusi:o:n,s lh;u ox)"Su:ml s make Snm c clIrre,IiII[-i:s the e.ssclfI.ce o·,r b~oseml,inlil. :~n Imwolililh :llipiiclobi,I<I'JeF membranes (see~lfne ri~~ ~~o;);\I!;! us~dlhes~ id';,fls i~S(;n~liv{;~iO lhe~!lhibil~ry(lff~cl~ ofP~c. IU!l.d re(~m },l!0'!1"S •. SiCh;':!illl']~llS Uf{;. I)~n~:~). Ttlf!ct ...,." ~n$or. the In\tmoJ! ~h~l:re le'<kse of oxy.ste:m]ls, frill~,nthe cell:1 :~.u~l1nr ~D11l1lkc arlifici<ll to~.·"t\l~<U:IiI1,c~: sensors fbr. alma-~ llili<m.e is :plmced be~v.'t\elll:'l'\"O S<I.~l S!(jIUlliO\~:!!. bnlrntl reS~d~l~~:-: Sm{ltt'dffieking. (~ili.tryll'l~,(;'!W;- lyn;'S i~d~ding, dfl~!b'. 'Pl'(l~t;~oo. oli~(l!:'du- ~!I'!d~11 .~!J!'Iie·eurreliil(~sp.~ss~d l~'irough (~(l and b.ra~.es a~We1'U!i.}~ i~l,clm~e:!!teroi. t~le precUlIit.;or of' : dm.:tl'O]:yte~fill.ed~1l'1il~loporl! (3, :}." Wlnm ~ln oqs!:el'Ol:s" Ilia), 'C{)!!ffitro'il ih~ rel~se t'll.· p~ ox:y~ (. at. MiU~in i~in Ihe lD'l'~ntm~n~ of (m~lTIiSlil:Y i1md th@ ~illal:yte s.ttch Ill,!;, ,~l (]rug mC!'~~Cil.!~'~Ililc]'S the e (eri~ flbr !ll:es!01illlth .at lhelli'olN>alio h'ltler!a(e, i!Jliwei5l~ IIOr sterol:;: from the chol!esieml~ri:clil memhrnlile af Il:<Illopore.it u.'IUsiendy bj.ocksi~,iscm;m:n~; l~,e ~~a<!fll!l".u a'lnesv:iU!!'. ~llZ61:lL USA. E·;ma-ll:(r!l'1a'!lin@ G Ih~ pr.i m~ry dl iunl. nl.(troc~IQr CQfn.lfl~n:sa CUrf~rlil .is r~Slf.ln;:d whC:1l th~ drug ~",its,. As <l ,~her!lu,JfIL~'u2, $, S1""'i i~ in!h~ 1!)~21r!rn~J1It 1ItIys,1i;~ ¢ lII~d ste:rol:~SeJ,1~liil.g domilin .1Iml i:s ]~igMy homolc,," r'Cs.\I~~.01. string oCcurrent pu~ses is obn~ill~ed IUllfCniCllli,)t, Ufiwersil~ af (;ilj;lf~W1~a.Ir'o,ijlfl.e. ¥ilfi~', 0\ h wj[b ,e{!Chp~l~secorreSp(l~ldhlg to ;;t si!1gl!~ gous '~oN~~m~Hul-Pic~Ct prnt:e]n, 'which is 9'26~n.USA.

re.sP'U'IiI~i~'enel\!lJttl ,sill:! ar:lic;l"clh~.I~ [c:nl :il!~md.i:lil,g (31. c.a~1.~Sma (hm:oored i.nlil;l~mc:e-[h,diU·

t.rtun~rl\l!~tu:::r bind~s


to' C(Jllllll

ves[des) ro[I.~JJ!!,'C:into lliwcHilun

<II.nd .i!cti\r.t~e

:s<it~son I:he c~3nil:lmel.it opens, ;fIUo\'>'ill,g ,~~.l iO:l1liccutteIU ~:oHow. 111~S.p:D'Oce~.-jIll,lo;.h:cu1<'INieeogllit.~on ch:em~s,rrylb~lowedby tn:aIlS:~




U~~S!~mic~~l.~tr.:rPlH'!id..:s (.n To




rt!1(1~(\C~L !IlN[''@.!1;$lm~~liO~lc;):\f~!lt

me:~'~1(ldsfor prep;i1xing

111~:!'oB!lStlf coukl dl3'~~c~lmlb;p!!comoh.'lr pc~l~iill

ana:llyte molecuk,:s:lll~e eouuted one ~t a time ~Jl. Ihey pa.'>i'i lhl1.1u~:' Ule 1l~~I!!I,{Jpo:re ). 0


Ho.w.c'vcir. II sensor bared
ml[ ive


'!Jtlu~dcOl,lm .any ~~1ok~t;:~LI.\; t~~u



c~~ e'ln/!ef~~d p.a~sthi'(iugh. T~l 11~13.kehe pore selective •. <HI t M RA th,~t blinds ~h!i,lUii:rg~'~ ~~Illl-' ~y~e' as be~:1111 anaehed tOo the h

siilrn:g ,e~l,l~lIIopor-e[lm.el,1.l~ comu~enln,u:iallis I~i'n. ~ braues oft hi:; ty'pc ( 12). nlf; second MRA.. sYI1iI:ilcli.c..~ltumpore se.Il.~ :SC!f ( 111)1 deri\~e!'i frO;lfIl <Ill M RA"b:oll."ed o~ig;(l([ilu·and rOlf elchi!lJ8 cenical dcotide-lr,,:~:spmt J1I1IC1!'I1~b:r.me {.14). Hu~JP'Ofii! PQr~$ (se~ I:~.C· fi~lJm. W'd:8 moc.~rur<llbr.iC<lled ill .~i lilOlli diQ)c!.ide. \!,;TIich ~lJI;lI!erI)'lrte~.ll{ 13~. ArmJ~,oje ~I~e,)ci£sCl;1!1 l~ (If'!'Ul~cd 11> e'H&)I to ~:hl1ct iOl1uliz~ us;ing :Kilo:x=;Ll1:edlGJUS(lrv~d as th~ one ~~11i ~hlll~ they rra- i$~ry. A~pli£:onudwltd(i I.';:; :attached, MRA •.31Jidlhe t.rrget OlIIl;lly~eV/,13S d\i.e V'f~ I!be:~nU!.v oa::elliing 'l oriip J' of II (!(lil~Ci.1li.YrlQ- Qligm:gw:l~tllidl.;l ~l;ItIU(;1~{Vth.~ti"C{lmp~iilne~f pare l SeIlS(y1'Sw.i~illqp IID'Y 10 the bound seque:IlIJCe.l'hi:cUll:'iren;t pu~ ses
A peptide· i!R~~~e p~"es til f'Q!lJglh Lhe ,~IoHIL~~11I0pl!lrti! (4',

in:-;i(~e S!l.Uf.fi:I4;.'I;: nr~hc nmmp(ln: (.1). N m'l'. when the aIli[I~yte
enters, il. binds to the MRA and r~ri()d of It!m.~.e: d~wrmim~d by the' ehem istn:y o:hliDe MRAhlna= iy~e: i.lThH:i~7I;t!t The .lIesu~:ting ia~l.. C'llirlrrent pulses call! be d i:-;\'i agllli.!;:~.0d from l:~.o,~e~mdllJred by ml:l~ecll~!eslh~ll do 110itbilll<B.oo the MRA. and a .s;~i.~ct·u\~~ :S~!i!:'

opellillW' of:2 um beea ,described ( 7).



~ilr lh~ ,n:ml.yle 0 lig,(lnuch::otid~ coul d be disliIJSJlJishe(~llrol'n pulses produced by oiigO:Iil.lII.d~e(lil.id~s [h!lll diM Itot bind ·~·C)ll!iellIuached


Ute ~mll cur.bieilU fDr a

R~~iahJ~ metheds
Ilrepar ill.1l); Yllithel~C S



il\iaH~b]~ ~1:Ir

:ml.:lD1,O pQi.l"e:s.Siflot

ods faX' ilt~m::il,ing I he- .MRAll ..~re II:ILO\\l'n. but several ch<'lUen~es l'em~ui:II1 Fin.;'[. because .. I:~~.e· o;~i~I. n~n.opor~ isprepsred by pruteln se If-,Issem],bl y. J?Or~ fiJ.bri.c'I~:imil ]S .... N:rj·
sfiU1OOI1l df&J!idil' memtiran,es. {S~

OO,V' i,~ obl,dn~d,

IEJ.ec.U'Onl m~C1og;raph OIi!.n.rft !1'imnpowes IHlitlil d1~lenml: di.une;ters. in

.rellmdllci.Ih.I~,.[t mustbe dei!illl!onstr'.. ~edlh.al


This 3.PPI1O<I;ch hss been used ro !rli1:1!ke !O'.:-HL-bil~~d sensors Iltilt are selective 10 l1lel:l'l~ lens, drugs, '~llnteh;ls.. and 0.1ig:m:mc leet ide$, (J). 1-]Qw~I,'t:r. tile .~ ipidi ~,lle1!'l~.brJllme hoases rhe flJ[lopo:re is \lew)' that I'r~gjil(1!. dilEl~!ljshiIng '1 he' PI"OSI)a.~1$for develClpii:111!;l :J1l'rm:~ ..~sell sen fronIil '~his~ed~:11 I~ ic 0 o.g)'. SY~ltbe'~~c ~1Ul!~pore ~,ors. :~~~Ih sen ~ ic:hi
the pore :u.~bo\L"Cd into

!);.'Ilii,OpOl\l~ can al~i!) be pn;:pO'lfL':ld reprodudbly. Secol1ld. d~,e:irloll!H;e~:1ll stabUllY i(ow.'lrmJJm'h:5,) mus~ be explored,

f.intl~~.y. ilh a~I.~l,gem:1r[c en,ginecl"i~g ci'lin w IDe used liO ....~ w.clla the MRA <lIt ~r:uy desi [led lecarion ::J~ong Ih!:!illis~idl,e: ''''''.IIU~ of the !1f~oopure<.A srep '~oward such p[lijl'Dt~$cl!,lc-

li\'~e M .RA fu nctiolllanZ.,.~ion :b!! r-eoendy ~~!1 dltmO~ls~td wif~!l :SY!1t~.~[ien.[!~lO.lXIfe by cleposi 1:1~. a. nl<ltedallll.3t n b:ul1.ds rhe



mechun iC<'l~.ly s;labk mem bl:";iI~ernJiI~e rial (5-1'0) .. offer .1 P'Oi'i5:~ble50:iuti,oi.L Current reRe~rch in this area .ro~l1£e-s fin h¥oke-y que:llI:ILol1l::1: H(JlW cau 5yUlI:~:letic pore~ , ... ifh, ills;i de di<lmet,ers apiPllrOild, ~i1~ mol ec ala r
dim~nsi()gl~ Irll.~mlm€:l'en') b<i;; pr~~r,~d'? And ~~.o~:\t MIRA:; be rlUflclffied. SOO that sdet:'tive cmm,

MR.A <IIIoldy one aa)ening of the pore {15,. Sylil~hei]c na~mgpm;e SIilfl8HfSC eu Id reveluIJofli:re buosCf1Is,ing, but l1l\lic:~, b[ls[c seleace
mi(j[ograpl1l r'DG~illlg 'c!:'DWoIn C!l'liIi~~ a n.'Iin.Oj)ol? iI1i iii po~l'i:a~lwn~t~ liliteililll~l<Ine", ElemGA



be do ne: before

l h i5piftJIl."1


is: reallized ..
R!~r~rl!l1!(~ 1. a. IiIEltQ, ~i!lr.1JtllItiI1"l'f:!:of.~UWIi!'M~!lJ1illtS
~~rnd~"~~I. !\'JII,~rd


Two h':·C.~11i1:1Oogi e SHire' I emer g; in s the frolu n~n1mers~;orI'rep~ rlng liiJ!'lliQ.thci. ic~ 11.1.UlOiP01'e se~,S'()rs. The fiBI. die·ri\'es. from ~~li~ml itheigro:ph;r @!tiidi i.mwiills us! fig .a h~w~ of etec~mn:s (5) or :~ons (6) w' bore' 3. holle~:linroWlgh:


Bfd~micro f'''IbriCIlI00 iIInd rraek-etched mlllrl.Oipores havebeeaused assensees W Co[~ll'llt 9l!rl~lyte spceies. i~lcllld~ng lurge [)NAs.. pre~e~Il~. ~m.I,lIrn,o~~ec!illles{3-nn. Hmve'l,ier~ and ii' most or lh~[;e ~n~di,t;ls ..the n~nopure '!,;\!!I~
1101: sel~ec~ive.The fi~Hll ehallenge





~I~~!lmc;'ll~bnl~~ ~!~ i!lorg~l!lk,: m~t~rj,11 l\JIRAs ~o c'~li!l~ ~fU1!~y~-1'GI~-c.li'V~ S;~!~S(~fS~ of !ioucl~ .a:-; ~Hic-un diQ.'!\~clt-.'The ele\:lwfI-bemn l'h~reh~n'~ be!r;:n~\\iQ roellOft:5 'Of "!'mking
.3 trilll:SUl

[m.[ul~jibody w preparat ion (SI."C the' :Figure,. luUdicl;]e pand j, lhe pm~l'!.ill b",.h~l"Ii;dllj \\H::re.auached 1:0tmck= Ptirtls w~lh d~m:a.l!e~en; ..s ~;alil~\1J.:n; ]D:I[Jl h01i'l,oe ,e~chp:r-.o~ein s~]II.sors bm~ed on COIllic'l~ ,!!pl.d bee:llp:repalied (5 Jr. 1l3IJ.m.'ubes (.8). T~e MRA!>were'H]<li~'n.tld tothe In~he ~c!1nd f~clmo~ogy.~~ [hinplasd~ .gflld lhR)ugh simlP'l~ t hJ[l~-:bflood ch!(;m iK\ry. ll,nembmr.lt" is. Ibombmmecl with Ol. helJ.m of The !>e.~:so.rW<1S. l'lI1,oull!ed betweellll:\!I'O .sH~.t hi;;:ll-I1lL'i!.a:ry:y 'p1intcl~g to c'r.r;:~!t~ dumage: s,oh.u~(l!mlK.amd ~rI iomie C~f~~uwm. pasfled ll:iI.C.b ~:hIITmg,~,he rnle:l1l~b:mne, foUmved b'y t lhlJlOugllilllhe: .n~1:1!ii.!Jtllbe. TIme di.a.meter ,of Ithe che~nic~1 etching of ~het;(')'t~ilLC~S to Ill.:a.k,~ mmol1lJliJcl~.pwus fhli:,:4U~1~d10 mal~h ~hC1: ~iz~ ohhellll<dyte protiein (-S :iIiIlJiI.~,) boullid by the the na nopon::s U Th~8 met~.lodl 11015 been ~lscd c{)!unwrci~l]y for d~fld~s to pn;p,~[~ 1'I111A. As ~1I"(:!rLl1]l. lh~lip b~~~lil~G ocdud!i;d ~~."lln)lp{~rel~;ken·'. j\(lol:e :u:ec::entiy. G 511 IU'Ii!, upon :r1wte:iin b:ind~ilg. ~lmuinJg Oin~du: klncm~ re!~f-l;)!l ··]!T'IIe:rse·' igalld·~T ... i(]l.Ji11 c~;:)j!llbel. I -d Dl:1!nn$md~'<'G~nnfl!ly •.~~>l!S d,ev~]o:~d simple

issioll elC'>clron ~licro~

1\JIIA~f~l.natmO:Il31~zed Se~cSQfS. In the lirs~:.

wope ...dl~, ...irag~J~e pore 1.000 li!O,lg,ed {t~lriu~~ pwtei~~. ind~l,g MRAs,(ML'Ch b

9. C ..DilHifr,liI.::rl .~!I!Jmh, 2:" :!Qi9 (oo01). ])0'. '~.M.l!jbil!II[:I.I'I!!i;ri~. ~5~ir.il!WJ; Mmrd«h. !l~iliO It

:2. Il W.. ~jrn~, im fMllitl'.t ID't·AiM.Iy:dl. Wl. G. Slnff!'J'WlI'lIi!I" IH_ ~ilile<s, Ed5. {8:(Jfal5otie!~ 00' Clt!:tI]iW'y, 1~lgrid9~, 'I!/, llt::,. :1l~'92), p~.31i1:1-3?1. 3. It Il~Ley. iP; S.. CveJ1'!llf. ~~flffI!4n" 2:.l6 I[;WO:U. "I. if, Ill, $utnerlall!!l ,~Qi!:. W!1n,a l,~~. "', :J!li3 ,[~oa~), 5. A.]. SIxw,mo.1L Oien, )t S. li!'!9,. !iI. W. :z:a ~dbe-rg,t!'l, H. C (lIi!k~~,1II1lt1, I'lltu. 2" 5:37 {~O~l\ N 6. ]. U et.~i:.I!IQ~t.Jrf 412, 166120Cl:lil'. "]. 1;, II, Hililn!\. 1..5.Siw)o; L Ii.. !I~er. (, R. ffla1r~!'1, l'la.iiJo ,tNt :5, 132<1 {OOO~. 8.l.!iiwt e.lQJ•• /.~. 'I'rc~. S!.!ICil,::l;7. Sooal a~5}. ~2oaij!). ~. I:. IFILeisdhe'f,P.IB, ~~~. I!t I\t W~bl~, .~.rellfJiam.iJl S,ollrh: ,P'ffnr:'ptes ondAlpp1ir::aJf.rm (iUnil'. CamDimi~ I'r~~,
8!er~I~. 1975),



~ ...
~ ~




1;1.:5i!KIl1r. IliIt!m1~n iI1iil!l1l ~B .. ~~ 2.951l)'1l. (2 tl9'83}; '" 1!!I~!~d Slille:!i. PiIJeIl1I.N'~.411!r:.93;:rO 1(:l9:e;}. 51 1ll. I?~tu '!Ill [ ~~ ~d. Z. Si\w. It s.prJtr, Mo. Yo!ltl:l!a, /!1m. Ii'lflrum. MWliWm:/ltyl. ~8.Seg~ Ii 1M •. Hi! {lOO'ill). :][11, 1'; I<AjIli1j ei 0'" ScrM~·105,.9M t2ijOol}. :liS. J. N:ilJsl:olI, JI.'It i. ILe@,1'."I.l'!<Itto. S.. E.li!lllml., Arlwmru lMm'r:iI!J!j.~~7 ~:;:(t06~~.

! m





IMI'oto,rPreteins a.1 W,ork 1:10 [1' Nlslnot)8'chnol,ogy

I~i.. or in~CP;,,'lVtlo pgIWC'fwhcthc'il: w~ ron (';l~lp~Oy!hl,":{i~ir,.'jrl'JiC' ..] fl;;L!1m!l1~hi!l~, il~ rtif~· h n I;;i::d I(;nVimIll1fil\'~I~al'iQ!I!l$iw ~rn~ I;:~J.~ to ~rfOl1"', m(l~k~, ~ha! WL~ dt~igJ~~o a~r bc.ti(;:l~l{2.. Oit. III !hc \'el)' least, can 'Ih~~ l1!1lJo~i!i.s pmvidr;; I)!~, widl ~hc ili5«limLirmlo ~11ill~i~ hiIJiCO~lliXl'llC!.lIS 'or dt.~ si:g!'i~irl;r!Ci.:HII'ilOI'OrNOl'i OOll\pambrc g~<JI:r,{.1'1


Tlheb,iol!lgic.all. eell

]!l: el\ll,1II1:IlLPed~'fnh i1I variely or mOI!!C!l.dalr miH:i1Iines tha'~ lP,ed,()ru'Jl cCll1iIple.x mechanical. t3isks)lIIcfh ,as ,ce:tl di\l'i!i~ol1 or in~r~celhd::H ~an!ililort Chlll! can '!!FlvijlS]Cm em,pllo'yiln!!l th€,s;@ b1o~e gijical. motlOrsin, .ilrt~fic,i,,'l l!,nvilr'~!I1Ilf1!!flJt!i. W@ [!I2VI'il'\Y the PtQ~I~!?~ th"t lila5 bC'~fI ~ll'1l!de ilfl ~:s1!tg Q'I:a)t:cr IProtfi.li1~ror pci!!Ii!!nng i!JW m"n~lll!.!~at.i!ll!!iIi1.af!os,eale ('Qmpon:~nlts;,. 1!IiI paltacl!!tar, k]n.e~n ,alf!dl !T'!J~~si:n bfomi!Jllli'S tha[ !T1IillJ,lie ,alo!i!!!j ~~!lili~~' bi,~mamerlts h,,'11~ beelF! \lIi:d~ty e)i,pl~[II~as a(l~\I'e 'COIl'!;POill,ef'lt$, (!JI flI'e!'l~1y ~ea~~;!e~ ;;IIPPlfc;;I[io!1 $ ,al'lE!!,ii'ne,r{1~.y jprO!1Df-of-prim!dple ~'eli'li1on5.~r,~t,lon~. Yet. th~ sh'~er ,[I'I!'a~talhit~tfIllf ,an iIl!tlt1 re l~eadly·Ui"1lJl~elQ(I~b~)i: of I'Ilal'lo~n:!~·d I Iiliologk'all .JrIiI,ot~lr~;15 3 'Ell'leat '01 pp!!J:rt~i'lity thi1~ (a ~l5rQf ,~:.:plorat~o.r! ..

N!tI~lI! re'$ 'W,swll(h,orsesfl1l tliII@ C,Hm. 11"1onli'fi!>~ 'tf,i rnOJ:!!rMcopic:unachim:.r.. ib1i01:ur' proe telas opemtc inl ~, wmild where Bro\,V,nl~mlli mOlten aad \',bX:Olli,~ lerees oom!imIlC. ll1~c're~\at'lt CI~s;ca!e here is ~$Jl: the 1l\I\o(:tiJlcl,(!rBohlDl,",lI1n~s: rol1:;,t~nt <lml ~~mrc....,~re. whk'!1 .nnOllflit'l ~('I 4 pN·mll.l'hi~, :IIU)" be ~01TII]mrod i!:O' [hc"'~O ~I)N'm'l:l of !Cn;(:!rg)' dorivtd Ifm1!l l~ydroly~,i5, f ,I o sil1gk:ATP IJ!OII;",Cufu O;'[t p~l)rniijlogi(;".(i.1 Clll1.d~ lioJl:i.1!IK'ilnm'II, ~1011~":I"l1'lini~iC:"IJO,tl(ll~i~,liUI!S t m~ .i~nPQn:ruu a~(."e! of ~hi;ldy-


ri['1l~fl!i'~Eionnlliu Ihe: Ii\!~il'ig,eell ('fu~1 be vi.C'I';'cd asa ~n1.ll:iilture 11.clOI)' ~1181. ,oonroitlS II ~a rg,e c;O:II\::el~ml .of d~..akJ~cd prolein b11achim.'$ Con •.,[dcrlilc oomplicaioo mSks !hal a ~n~lc:cC'1I C'anperlhnn :~I cau CIlI."aW.I f~! CQ~~~'of il.'l(ll ri~l II less Ihan an hom: il. ,c:m~ rroof~ .re!Ic] tlI~d~p<I~k CJII'OJ:S· in i~" O~,\fnl DNA. ~'C'J:L~ its C:llviro!1mC!~t m1d



IogiJ"...~ ~'{1IIcli ill R'C!.:l!lI . ~'"'(lldcs ha·, S!Jl,lnOO iht:

".I1I.. 1n:U!~~


nm~li~$of !JUllnfl;'lro~~ill"\;. L~1. '~b'rildly(,lCIlk"";dt'!'
(',XaitJph::,'O lill)'


'DrliJijo~no-m~,Tle m-





(rig. 11'\.) SYlltlK"!~i21~ AliI? rrom il'dCflosi'lte dliphoSI)]Ult-e' (AID'!»

~Ild I)hospilmte


1.11e ITIm\1 of

akmg an dlccll1!.1!l:'hemic,d gt<!dl'cmllu:nugh!l,u;: n~eIIntmIl'!1.-":

boend Fo




f!,,"'SIJl!')I1.d 00 iI, c~~ml!l!.t i~ sh~gx;: m.o~~ph(DIQM.')I. K] 'lJhmi~~~!lm Cl~;,Y Imln rhO'l(l''')'J,'1l1u:,\);il'i me .. Oli nnbolit'il1\ WI:>ing priuClP~~\llIit me ~iml1i~f1r :m~f1r m cells 0:, WJtI£r1CSC Alrllhfs fLlI'iC'tirn~~lily derives flll:m1 11~Qt15i3nds 0011' sophi5~iCil,ted pmtelns, OJl~il11izcrl by billlons yeaus or evolanon, AI Ihicl1l10>11l.1e~I,WC'C;H~(1II~ly dremH of rons~mctil'lig n~a.chinlCs of simillll'


ef lhc Fo ring ~td dre l'cntrnl, .sm~k !!:lOJDK'l1illg I1lc Po ill_lid FI l]lIotQN>. Tl~~ Indlll:ccsoon[oll!m,.· ILo:rU'!~ Ch>lI~~Cii (Df Ihe F:II ~ll{ll!llr liloll dli",,~ [l~\.,': e'w!)'dyl it [(JI'I'lrulltOIl! of ATIP_ R(.'Ilrnnri-Labl),.uhc OOllt'" Ilfcrx can also wo:rk in. reverse, l:I&iu~.!!l 'i!UICrgy the AIl) I~m~. ys>i.~ w drive II.I~c reW:fSC rolfil[{m, Oorlhc PI nlO1Or2tnd i>LlbscqllCI]~ly


IrLU~lp ,uulOnsagaill~

d'Cir elce-

The rotor,)' 'bfL\}lCri[l1 :~lflg,ellar

• i~ Inl..'llt I'IOtlSC$S j'Llst a rmc~h;m ~ dw .limclijmlEllily of ~I!I~ Il(l!;t~!uld wonders,


mmor {Fig. U3)·hs lIsal by bac~ rill, such as Esdit:ridlia (,'f_lrt 3S a

O.~~C~:~;c~t1;:l~liU' o.f pJ,flCrl[l~5 letn...i!5 IIJJn~~;d,by !l1!oILit~II !· .. UlmIO:I.- 'l.,~rl'£Y.,m;'t. \vlil;~h ~. (;111., .dy~iC'rroteilL~ dl~Jl c~)n1.Hirl~m,;vil~,!; 1M~,i:\lind IIIS~ .fu :;oLl,uce of ~i!t'.(: CJiIC.rg)i [0 dii\i~I. t~leir me-

the- il:elL fA) R!~PJ~sw~tion of Fo'r1l.~TP@s.e [r!.lprlWl'l:edi pllOpulsion :nmh,misill by :~pin~lIi~h permiS'Slonrr:o:m (45); cillpyrigin 2006., W~lizy·vClHll. ,liB) Ilitep~e~nlla'ti~1'I ,O'F'~l1e !vitlg a he~ical m,~:gc~luml:))_ "111is ~ llaJc~riaiL~C!!g;eru~rII!totClr (image ,~~~rUl5Y or~~kM IM.ambll, O~k~ IUlI!i,\1ElfS1t:y). hlS@t jJ'Cl\'I;'lC:di.111 ~l~~. ,~"iCff~b~ r~'Qlll IFig,.l. MQt~:r p:ro~;eirnin
SRe\1I5 an 'eLer::IirQrl mi~r1l:!illepYiITJai~,e off~hE!' m.llh:J:r Ireprint:e~' wiilih pEiiFn:Jlls:simhtll1rl {46~';' ~o;j):yrlgM. .2001, IE~evil!!r'~. ~O' COrlv.~J1lion:at Idnesilll' illiU:l: d~einJ t:rampor:l:
(~9!!J' hll (lIPRlOiSitl! dh,ocl'UOflS aWIiI9I mh:rl.1t.uJbul.@s [adapt@d h'om {l)]. ~D} I~I!1~n ~s .Ii

lUPOOIlsiudyija'lg t~~ l1i1OlIIts, to' flIlflChines ~on1.~ nnore W~d itl'tOfC d(;'ill:.


111[(U:e:5SWe motor, (t(lnliii~'l:lrl!lll of '[''!lO' Ihea,d£, that \llil ~l;;5, in 8itl€iFlma~~~t:e~, of 6 om alounglt~e n,if:crowhlu~e [adaptedfm:m ('51], (EJ Mld~de (:Qntnmt~'m is ~alU~d, by '~he :sUdiP1~1 (If i!iltemig~Ia,t.ed (Kfilll t!!1I1~1my<JSilil mW'Plienrls 'ifill !1! S:i1!JC~!i!ll~~ 'I.!f!it 1Ii11~ mlU"I,pmC!5'[ill!i1~ !1I)IIosinJ IIImo~ordet",des: after ead pmver ,streb so as: Hot to liliiipecle theJ~ illhew :sli.dii"19of m,e .u:lil'il i~ilam~t ICaw5,ed ,~~her mYS!i)~ii1s, [ada;plilld b~
from ~F) RNA IJl)tyllliief~~ m~w~ alolillgl a dc~b~"Strandm IDN'A,'!:@liIllillat@, traltlS(lrii)irn.g oil RNAoopy ~imiilg~ oolll~es~ ,of Illlt ). 'GQ:()d:s~l~, .s!i:ripp:~ l'I~;;Irch ~1I!s;I!i'ru~). \eVe 1'i.~111l1(illml> d",,~, atl: [ID"'~'eDU:i by ch~u,k:ai C!lc!'l~nr. dct~~~d hJldrol¥7jl'~ OOi;'~~le~· rii~ll!!!~e {J\TP~!iltlr(;'(d~ 'i~h~ ~r:> ~111<Bjot'''.i'h;'t~y f:Um:,11CYl, mid ~:rI1~tC!1N OOipk1!l' a. gftidi,.1tt ~:Ial of intl.", ti~iliig boih c~cclric: l!li'id ,,;ilihujJiic rO!l.,(~

nl!gl~~l<m d~J1(;-p;;:nl I~wt~ill.~. 20 ~.:;: nl'lli.;'r! by 111l iuwul'd, In(~1.{m d i'lu;o,; ~hfll i~c~Jn\'el1ll.id; c:Y ~'II('[,HI 10:rqU(;'gcncnHi i~g, :stmorn imo a rotary Im.oli.o.~l the cyor ~in.dincal Iings~iil<! centrol sha n. l·he lillotO:; gCl:l,cmt'Cs IOrq'LlC::>
molt: timn

\,V~ Ihid. In'i.ruy ll:Kiton;~hat oom~ f.1l1SC :;iilil fis and beruiitJlg.<;" m" \¥c1I <IS ,~inear n~i]too; Ihal IllG\'C' along


m) 'PN'nlll~, (250 k.all


<l:wd mi.atcs 31, :.~-d!> OI~' m,(!'r I Oil] .11.<':~5t Lill,ear-rnol,iol~ I'IH1tt~rs are
fOlllnd 3111:0111; ~lC'

~rmiks in






Kl3iJii ln~titu~ilol,fij.unlW::iElO(f'> Mt UJirrernty (l,f TN rudlogy, I 1, ,26.~8 Cj Q.e\[t, W4e'~1~~00:s,. .


the ~\Iper filmi ~~ gf kim,'siin,,cy,ndn,;md myosin piJo:wins (Iii fFig.l. mE). Ih:;,'sc rno~fl' ntmre

'"To vdlO<ni1 'C4Irespoliideucce'

s~'Outd be oi!di!lres~. 1E·!'I1~n t

in, dn~cwlc M~Jl(i; ;1~Qf!,g '~u~iC'bn'!llidt~ of ~o~g Ilm.h:~in 1'iIolym(:ts hlcti.ri fihimt:~I:ll 1fE!(~ll~·osili.
iiI'ii~ml~illlbuk.";!;, kifjb~li Iflr lilid dYI'L'l::jll~) dild fotl'l1



~he cyt()~elelOJlllmt C'.l'\b;:ruIs~hrol!8hmll~ eel] jhe


l~ ~.l(.'1,"~n\,':!l1~ m!;l!!on, ;do:n.g i2!:il,;~rtl!""Qk.'i_ !Df
"'~J;llJi1i[1I,1o'it')1}; ::ii,ruiiIVtdV1,"d

5ri;n,!Q'i1Jf;m!. P9!aLi1y ortll!~c ~'ilrnlllc!li!." {de~Q.wQ ~, p!!;iloi: <lIW mi'l~~'lot;'j:lCl) ,I!!m,~""~!i!ni~!ir«~iO]l;;i!
C)"iQa1mnl>t ~'\r~':1)'m;pli,lC:~.

of tg!llr'Qi~ed !1!lJ[}l~'on ,1I1:doloree gone r'dlio'n ~'\1ilhil~ CliM~, s~£h .~ imnl(;",II!J!I01t: 'mn!;;pon!1" rI'i<'li~cn~I~~ (Fig" [() (11, celli di\lirs;L~ti, it11'~rriO\!,I<CJlial~ ~Uikll~C!I!ie:rl!lIIgd.l~. allt! ei li~l. The co:iiilniCUol'i ,of.j ~'ii!li:ml,e i~ ,d'..rilvl!l'i'by lite oreh~lmk-d :;!idi~~,g, f o series offLctiln 11~~hlirn1cnL" rc~~pect te !Irrayi!. wlith ~f nmYM:iJ1 Il1!1Hm;;,(Fii£,. I E). T:rpiea'illy. a. ~inellr ~noUl'1"'C'mn!:lel1.Cl<i!e roooes of'up 10 - W pN. M~:my liCi:1Jhl;}r~:oti;lill~C,'I(~::1<t (lIIal com usc i."iIler.g)' ~(l ~rlhml \:\,()fk, !liU~JI OJ!:;' ron chmncl~. DNA- Of





IFig I F). '~~!T!.%'.

I[}f~ighl·pm'.,~ •.'~. ()ol\'Ci!ttul ~JiW!mllS.. but [I\CSIC'rlll1 (lul","id~ tli!~!'i~"t1~1~ pfl.his,Dt'vi~"," M'1Il5CI,e; :liilJw~rfQr INitlllflot:e'cl~[liIol~OQlf

Chli,; $mki'I!!ld'emfi~!.\iI!rl"~O~~ of


~ioml]k~liIlb ... Ifig., ,2:. Moro:r p:m'leim,in nilnote<lhrlll'~Clgl!I. (A)I An Ir3,-AlPI:Ii5e--tP®'Ilorered nanop:r,op.eUloe;r I[am,ptm 1irro:m (6)]1, IFh1fOl~Qe~'l:e imag~$ {1ll33-;ms iml'leO!'a'~'a;rI!hmnl the 'eilr~ier eKiP~rimmt (47,) t~a~ iij~~dem~n5tra:l~d ~k r(li~'"!, _._",I!!!Y.. - r ·11~,1 4c ~"fIP~":__.,,,.L·_ k" _~C .. !:II ... ~_.".r~;o .. ~·,,~ .10'*"111 __ ''--.,IL _ ," ._e"l,,1J L ""1,,. :,~...l -_ ." ",Hill .-~!J- - m""'''iili 1;1_ ~ .Ill . ". ~·"'·1'. .,,,~ 1tfti"'\[U' "'.1 u'-irn- ," flu"-,.....-,'"_I,,\L rfl'''m ~f1It ,o'\lliIrlFP"".J >'1ilI "I; .. ~~!!!<l. [.adap~ed \~th perlil1jl!ssio~, 'hm (47'); (opy.ri~ht. 19~1, M,umi~l:;m lfIubl'i~n~r!> Lt~: Natur,e'J. (I') .1lI. Imi~rO~(I:r (2~lllli'!i1. di~m!~~r)' p~'It'l~d b:y baJ:'teri~ that gd~,~to~he rol\(lf:;l~d: glqd~ Ym!~dlire(!tj(li'i!aUy ~hrl!l'LI~h U!@ iIf.cu:lk. Ph~loimag,~!1 :;hOcw!he dQekwil~~ nMi!tiolli ~~"~h€ rot:i!l'f [aodaptlld wii~1h:p@m1[jsiolflhl!lnll (1 ru~ (opyri9JIiI~20[!6. Nal'ioll!,al Aal(i!!l!iI'II' of Sde:n:c'e1i L1SA]. (0 Schen'i!alicol~ ki~e~1iI iffi!(I!kI~ p.rol~i[!'Is
!l!11::i!i(lfb@~ to,

1:1UI1A\1llr\IlI;~l,I~ Llt~ C(U~tpil,ll."Dio]] a ~ ;Q1f .!lkkc~.mlllo;pmp;:lI!;r nh:,umm~!""S 1J1rough IhL:tl~it'll]1

lL':nigi:n~U('iI F~~'\IPas:c 1.1fIOIOr (8) (Pi!} 21'1):. The direeted rliSSem.bi}t ornhe devices \';f,i1,'> COlli", Iroilll!t1lhl'm.a.:m.h:C'I~cl:ic 'tlgm~lCcrill,g oFhis.'lidna'Cmgs g Ihm. !llll.dktth~ F~-}(['I)aliC o.nw llickc:III?OS~ .... with ~ts

cCIlllml stillk 1~lUlrudiu'l8l1:1~\";';llru.. cmlllootw to 111is a nickc~ Pl:O;IX.~ll'r or ... 1lI111, 1!.:ng1h IlU10:UI!lh biD1:il1~ 1
''illll.il1ttl\'i~t~1il bonls,

l\'ddi.~o~~ ATf £If kg;~iclli~f thl! pro:pcUl!r, A llilClal-ibilld~, o



site \"'<1,'>

rfl~f.nimled 'm~n pe'l!I'Il~f>f:01il


surr1l!(@p.ro~~~iin:(Ja m«c~!lJtull'l.!h~ :shl!iW~1 wnl;ch lbll'1~s ,ca~(1 !ii!!d! as il IDNA m~l@(!.!Jtil'S J!f:lJI11 (12); !liopylfl1g[~t 2:00'3:, Arn~ri('!rI1 ChelfniGl I. Socie;ty l'AiCS)~ .Vhl?

C'ilgill:lCP.>d into ~}C: motor Jllld acta.! ,M~l t':\'Cf;Slblc 1 CllWJlf ~.. ~il.th 'b:r ot1rS~uc~!1g~he romlirnl l1J;i!lJJi~
lJant;lil'lg or [1;,0]11(: Inl] ~9~, s:in1!~I_mrrrl~he <Ictil(ln {Jf Plluil1:g ;:u SI~ok ~X1,\,'I'():,:'11 l\'II'(>()og\,"'~I~I:ii, On ,. I~r ~lk;,!1J;dil1g becteria lmvclL"'1;'U1 used rOoprnvcr II liIiIU::mliI(;-eI'llllltiCfi1~\~ee romfIJJIi;;.. iug .~ ror.\\'hre~lnapccl IDI.Ol of 2~m, ·dii:tIJJl~lcr rolfu:nifDg in a silioon !rrJcl,: (/(f~ l'aaclCri!i adl:lCi\.'dlO lite nJl.Or.lu'~.ing it wilil -2 rp-m lF~ 213),11lC'in". CI'CaSC in site lce~1s cOibilllill'l'l.i ,>\([tI!lindividtlOl IlfI(}[CIl1!."i.)i aCCCltnpmliod,\\'ilh il~nyCl"lorqucs. lO~"Ol~lcr

[r,~pnn~ ill ilout L like

lDlihage shows '~.l:ne~liI-pmpel.led :m~l:trl)itlllbulif'.'lI!11H~v~'ng Hm:lIlIgI~ 'OlJl!?oI1pdYl!lreth,aJl:IE~ malflnei.s M~ perlflllissiolnhQI!rI U:6).;c@pyPight. 2.003, tlCS]I. 'The w!~(!!I;,ily'of mibJtll!blj~f~is ly;pi(,:dbl jJ.mf;5., whe;re<3s.attin nm.tmty cain!readl :lipt!ed5: up to :1:.0 ~LI1ilili_

3A-~1 1I'Il~'I:~lbrt;'"Ol'llt.~icl('FoltlI1c ~rrn:rlh[L$ ]x,;~I! dilllccll,'il f~Del'l.ltljilll~ oolrnr.lcll'llOI~I~y by t'ilTlplo;,"';ITi~ cill'W.:'li' diOOllt:;al Pfill{',I:nillgof rtdi\ilr; rlxoor pro.eu!ils taehed mrgo :iWlch ;,';:1; (Pi!i,,:;!C). I(/.fj or rtlllrirnn~'tI1~1p-hicrt! ~~~n1'i (lPil} 3A-[I) (I j. 161 A di:-.oni\'1ull)(~~ ()rlitlJ!,';~y dl.).l,l~CQ~,PH1!Ltnl.~!i; h~ ;fin fil1cmill~vc gooum:lry, l'IiIot{)r.(;f)li~M Cfll'gO i'> 1I111:~ liku:t~lI~ I,,~:lry d:~It'il "urn d~·lnr.1ckN. ,..,.hkh .. Clln,nnoV'c O]DII.~ cyloskc1.c1311 IiInmrnClins I!Il[il!8irt ocru~ w.Im~·t1'nile ~C''-Idin:g (ll.dof tix: lilm'l'K'I1Jl {"w~Iif1t m:lso:mai (lillO, a SUb$lI.'ilIC, This roquUr~'S thc ~illl:d a lin' bllOliOt lobi:oo IO.W~b~M IIbptireb' l.1o.mro:~kd I)~<lre~n;!nlor rilrnlll.l'lIln::> ornoa S'ui»1r.:U.c' wilh$C~]~'P3irirlgp~ics, Crurlb~~ {h~1n and IM'COO!ll,dl~g ,oft:he cargo \...ul.hn:lOm;rs, 'Jlhc :til, [O,rm£lfflphk~IIILY sr~iU(lllT.\,-drurli"J.rcs lite sclCCliYilY 01" 1m1~lejl Imvc ~'l 1II~, to drrh1C' a ,scU'~~lllbroo ~i1IIK:1ioJ~~ mmor ob.'>O:~J~ion 1001. ·tis vctiled glidiJlg gC(I(l1l!uy o.tlm bct~.~I' O;I~mn:L.IIlillics. hoWO'er. Ihr ~1t!aliQll. ll~lClk)rmli~lkm.","'\Scmh"liilicmowaikel' 'Q./I), TI};!' oooruil1<Iled conln1:C1io:n of til. combimnf.on ofIOfl'O,gJ~Q'~lic<11and d~C111r~ail gciltC1ml. Il1R1;!mi~ {] 7}. w:Ttl~ 1Il.'C :111010:1," ~1Il0lCrinS on~)' :Il~hc Im!$c!c~tmdles" \:ihkhwcl'c ~L<;$CIllblcct Gl1ttD ;11 Iy, I.md comro~ and is jhus IncrlCm'd. -0'.~·U1HI la~c tw~,~ggcc~ 1slruc~(!fC;.d!uvc :I'IS mLlh:qllc :liIlO'tOl1ll .uutcb ttl u sillg~cl1:1'oultau,,slnmJ~, Si0 'hmmn oflhe ll'CILdhC'.sWig. 3A"Um"lms :rrO\'\.'n~o ' O;_J!n:dbim:!h~ ~~ of ~h aPJ~!1!'mdl'l.~ !1I.~d \\fi~l M) ~;m'~ ]~irgCfl.l"'~~I!»!O ~ll\'1C<Ulbe g.c!1aO'lt~'JC.I. S~"'p',\~~eITl<1\H.'l1Ill,nl \'\fuLh ,;I, Np;lCct of3~· "un's. ~~lic~~lll:ln~:Ib> U8) C ,!l~!i!li~l~ty III ~~IIk!ia]g .ntd cmr~!n:"'f1il~~~Or Line,,, \)'oo..<lk~l'cq~l:ki!.].~..,in n~y\l,~il~~lc~t~Pl'i AIlJjmh~:'ld\'!.'II~~'Jgci:;; flam th~;vj:w!.ul~k·:;; Hl.l alld Wld>lC~~1l (lQ), T~ ~ceil~; \119': of' mclosed rI(~idk: b b'I"~ dm'lllrl;,i!l~ the i,ii!ll;(;"rgiUI!lj ntild t!rpn}1~iu.- bt:.~ ill!li,;r.lbn.-d ~!} ;,I. y,a~i~ ~(cou:gp ~1~il!:I.l'l~hi; imillI;lmm1'cls (JI]: J.l} ~ be ronffiidell."1.! :OL..'l .<lI'Q~Il;<1!I :.tf('P[;;I\,Jdin I.b:di:i"lgj; ~1I'-ID:mail!;rrlHlJ!ab!ldi~. J:1!.f'lnTCd dei\l~iC,'i;:Soc"C;nl~' dl.cy an; ~I.dfltiveb ~;(lblli\U ~i~<IiI'~~. g!l thl,i ,Oc\~!'Qpm.clllll{l'v.:!_~ CQ~~iIJC!~!1~ 21J;ld n:;;}diily f~vLliL~abl(;, ACI.irl fLnd ~Llbtd~1l IC2In tltC KiillllH:ilil· iI'lItWA~~c,sifl~Drhl',en[i:1IIllSpcrl ,1',l:J1 (hillS T I;)ff~ri'li!ll !~ll l~~!L;Tpg~x:!i\l(,,'5 1~~:k~~!l1~ col1~l~roiaHypuiiChiliOO •. wh~s the motOO' ~IQ2i'J~ brad for i~~ !ld'dn,.~bilijly ori~~i:ivid\ltI~, One \!~skl.n ii>~l31. I'I~O:to.l'pJ::Ci1tlns. wi:M ·be tl!S(d terns. 'Cilfillx: purified fi:o.nn ~:IIL.. (If ~~"1C,5Sc:d iUI 'ihlltlllgh dt;-otci~'icldl;or 16~¥l;; l"CCoo1bfnmnbiilctcri'lll :s:t'sten~ ;nd Il[l~e<l .iu lor '\1Ii1~mnOO C<lO'go 1~<ll1f\plll;]lii(-lfl on a chIp •. u:ilfll:l'iK~I~S For l>OiijJtgappliCfuDirn1S,.n~. Iii desiruble: tlmt lrurg,c qLlmrt~[tr'S.lal i1~irrilt~ imsiegt;'UII,k::UI)', lih(~e \\Iith I1ppl:1cl1licms,En S{i1riirll!il, iS~ruiJOIfi, Pl!lrir.G(!i~ I~lilliolloccur tlJlidi:r,ooiiO:~lijly. ,IJ.ccJtli.'lC thio:: l1iilOiC!l~ tiCltl, or ill>.-lCuribl.y orunm:tcr.iklis {.:? JJ) .. Torrnch IlloIDrl>yS[e~n_;;;, m:e ~m~~~"i'c~ in a sfM.'3~blg~kiin'lg; OOlllt>d ·to a &IlrH1CC m:c mirllio.m~ I,y m~.xib!e. 'Llll~' <i$$<i!.Y. illl \.v~lkh the cytaSkelet,llti ""lItlct,,!:.,, (llSlIIIiJillly (ilais .gn,tl, al1.<:lleoo.., to dcv(llop cOI~lrollloo.unoliOlI ,c;m allw\I'I1 ro 20 ~tnl ill k-Hgth), fflrt' [:DD\1pclled by a:iclng ,~~ilic a:OLII.e8. dkeclkl~JIiI.y,. oCOllllling W direcliio:n~:~mm}.'locm ill, glid:ing ,i:!.0011lJ.:t~,1a;: only .I)e ach;ievw IhlT!!~gh IlCOlie~llm&on orthe~~mncllts. CJ~ c'x,'lerrml 1!;l0l'11ro1. and slo\;,~n,g. .slillrfao.~-bmllnd lnolo:rn (!FiS- 2('), The :mn.l~iOl'li:!l1 ne mcdlOCl e:qjloils. OJ~Ylnll'enrkd 0I!",lIW"S11ii!J~ .. A ;preli:,>ql.!~te of mill' ~!~K~ Ir.ms:p!J~S)'S~t:l11 i:il nC'.xib:~lil>' of d.: mnlllr 51,:ijJk~ l:5.hIgh c,noogh tel ~Letmt~ (Ii) (.F:ig. ::m: I,nlml, rely Oil Ihe p:riJ1dp~e d m.o!~~ 't&i~ rnndounly oowld n~mrn::;; ill1!olhl; OO'J'li\;'C1 1l1a..1'IlOllol'II aud tr.ul:1p:JfI canbc (LlJr!i-~ it ..>ctiol~U¥ ~ o!'~~I.'iI:iIl!n fQr binding Orll{l lI~~(llicn~l_u'l1uh;or J,ll(BiIl'Xil. ;u[(IJ~g Pl~rt..'-..igIlIil,;dl 1l)~~~W;(¥S "~fi~Ol 111~ .• 1113~~h~~mbahilily Inll¥~\ ler::;e IiIlClit'Ctil'l£i1:':lil1'li~Cllll'i;! dcpcn:JS ou th~di","C~iOJlroom whkh the lilruncn( ~1't{,.11"~,1Ll'!:;: fth;it.irk-d r'JndQl'll!y rr.'!Ito ~ !iL1b>1f<1~i;'.d;;e Oii,CIUITi ~·il'~i1'('"J1.. PiU!lo-<L,"rld-clJu~'l;;~d aliJ;)lll'l:(l;itviU Ihorn cnlICr.:;:,nitjimnc.~'JI'l· melilloo <:<!n.u::hkwC' VI) '~('I din.r~j,ml of O[~l) tn[j;i_"fXii'i.il.; r.;lllclll~U a_.. "'1CJJ (Fig. propd Ill~ 11ibm!'iriJilt;'willi their :iuilH,Il:l !ti1cl ]~clii1g.
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sll!L1UI,1~ th!~D.IUIII~Jl

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'g'2% dllckm ~~i~k"j]titJl,1 fPCU" :m.1TQ\X' {22J, A diYl~~m. m;:lh'C·Q,"}~!r.g.], ,liIct'l1QO~ touse ~"",!crn;'ll !h:n;l~~cl~"'iith~,t{~!ld~ld ..B~l !IN" .~... ~~i!.~g11~ Qr ~ irl s ..nSblS >!!~prucO!ljm~:i!. A slmHi.v CN<HJ!Hp,I:C 1$ IJw : usc miCl\)i~!bu!es C9J~LXl \:\'ilh $h'!Jl!IK:-.~r.ll~cl(,'!l;! DNA iI)~'~nllll~lc~kl~%, wlriC~ l;:o~lk~ hy~rij&;r~\~)' s~itfi~.lyw~thi~:'i UtI~1 ONA ~l .8ohJ,tiol\ wiBl st~!'iu~iv1~¥for ;i :tl~:lgk~b:iSl;l,,~ir~lu~n~d


'\lcJ\Xiiy orO!C~in[ijl;nncm .s ~J'4), <JJthCl!.l~l ihcl1IDI~{IIl 00\~ki! net be I;l~ti~l)'.s!!.lPJm 1'I.!I1Qlhn Il~~hoo


¢'\I)bit.~ ~~I'''rm~p;:tl1c-.i~pt,'}!JnwH: on ,;I, Sil,lrtl);;:~ .(0 Gf]IJ~Q!I~1 ~l~ !Jw!i~~yof mk!o!u1.J;!III~':> {.:.~jJlh~~ rtllyql'i<,;'~ ~b}l.utlJO:.l~iWl,'.I::~:IlI~ k~~lc>iira nlt]~~1C1:iltg;, i whicll el!i:n bot i,lUccDric (J j). I11ijig,lk'iil::tlJ ).m :no!'v f~~JJd'l t141, By :suk'q~le!i.t thmlioll gf~hc fi:I~.~ j\n~lhtr .i!]tI;~ti,~g <lPlIl!Qach is lhe ~pmt on ,Ire in 1111C'X!e!i~b!onnfigll!!r.L1i~'inl IIJW~{;'f'l"~lm~ <I~ll!!t,.",- \!,ihkh !D~l ~.cri~lIy P~\~fU gMi!1g Or~hl;; nly~i!i-C~~~ n.u!~!t or .~-,lkll'!i!UlM'rii~ ,tj2~ -,;;:.,.t ~1~r!1~ 111(; rn w!d",-I)I~'lg ~~lPl!.oirn llil:S:!Jl£ '" ~h~:il;:1d (Ouch ~~ Ml!LIIJI.,".f.'d~dchyd~~. Ih¢cm-pc:'1 (""Ill ~~!\,C as ,;;I The IllMimv~['{~ ~\~~cn;ilLl.'d.by !tflla!ly~ie cn~ 111i:il,':rnI!ll!bld~f'~.3D~ Illt~llii~ll~ Ul; 'LOilp.rollu:(e ( d ioc-tti:ciI)I1!llymiligllCd ~illf~~ .IbtiJi'c!~l!}t.ptOl.dnnilb.iJil f gph:l i'i.\UililjJru'liele:>ooWilid o I.l],acliljlr.~"nrtt:li'l."', ~hivllnk. li1.lC f,"olYJ,l'J:;l1j, 1Llit! allowliimhe nniCIDllloolc:l C'oat"",'tI C8~'f> (25, 16), Plll,ttCil!l~Dlg wh~!C lt1ivirn~ ~:irie !l;CI~n1 I:h.>c t.o ii11'cFl'icl wilh lite 1.0 inllAlrnel. wi~:l ~lC Il:'Italfil'.';. By 5J:m1i~~ llne oo"llipl~i~ of C"dl'Ji;"O l\i} I~rolein sJ'Ibjl~k~ i~ 1n,)ll~in~'Oflli.'iI Siili'.ira4..;c. ·ni.iil> 1:It<!lhod fi1Ult! (J~i:Cif1], 1l\L're friO~Yi1Ik:~~hi,,~"C:~mique enn be {'if usc ro.t" ~Imi\'c~y stlll:igl1tlill'!!\'8rCll, Ilile llit:i,'ij)lcsl, OO-l111tyllla~ Winy of~sellmblil1£ !>n'lilill ~r~trirnJ e~t'tiilils. .~lllil'l& tlilc ,CJCl1silYof mQ.tile~nicrotillhubs., AI1.'OI.h.cr. Onc wa:)! ID!lcilicvc u'C\!~'jj'!>iblC' :mrlin£, mId mctll!.'X.l of roc.cl~y s<ning t~IC moc:mlllb'll~ dCluil;y s lio.l1 reli~ un the nOJ~"'Ci:l'ic elcclllC'Gt<U1C 01,' h~ ~llii~lc,ldso~im~ O(r~{I muo kimlsil!l.\'\,'ilk:h \"'.... :\1!OflJ)i:n,goJ rtJ~C' ~1'tOl:l>Hty 10 oontroJi tt~~cCOlb!::CI1- hlli, C-1'!:p!oitt.'\I cllX~rI'C Ik:]d~Ii:! mHmtt micro~llIblJ~c." ;;: f& IF.U,km [If' AlP or 1J1.!h~rlllX~l)' ,ooif~crors ~n onto :lkln,-~ n"""-~nl!.ld~old ilurfhCl;;l {~rGl'" lIS~ for 'LmidilOC~ion,~1 1:im1'5llCf1 (1f 11'i1I1~ds :such ClJmm~ or~h(: ,d:~~'~!UOI~ {l:W indiv~dwm~ mij~ suiutiml. UghH'IOl~~ltilied.S\v~t'Chins of ~'1~ me01!,;";' ,gold. [Im~¥l'l)rrei~" :md ,~ru:i< (1~'1). Bim in-


fi~m!l!";IlI[." r~umIIM~(ll~~ j·kikL



lll!11u.~ IWt!} r«e!!!~}'~.;.tun(l ~~ilil:Jc i!li1!'OtII~h~h~

u~ ,of do(trit:' or :!Il'l1~gl1.elic. fo:n:::e15, B.Y lI!S.i~!!; !;;'m::IOlIi.'d 11!J~dic chtu m!i.'lk on!'! can Olllf~:r ~h.\C~ri'C~~dN \'CCY ~OC<l~bitim! .1t1 on Ihe k..tdl'ing~ip n ' of fl nliclTIU~buk~~which ~~'~I!J(:ui~L"Ulytl~'.(! h~1l tlus \V-a)I. i!1di\lid~.~i!ll:litnJl!'i1)i,lI~ :lH)pJ1OlllChin!g ~ '{ j pcoifln C.'1l1lbi;. st~"Cred imo lhe (iL'Sirod, dt[(~CI:iQna by ~n c:t..."t:lric lick! "klfUI~ rapp!~~d.c.hmllgh .rJ1 ~\Iil]!ieli ~rpi!:ildiCUJltlil:t(); (ill! jL!mC'liOll I[Fig. 3E) (2n Ue mmIJinrJt&ofi (If li:~ectric and :~llli :lC'clllml~i(.'S; is idle ;tc.!.\1lru1I\gemt.;,for OIHlhip imcgrni~ion. M3gnelk:


rield." .have· been Ui;~ Ul [tired (he IMotili~y of m itl'O]!lilq,!I~ 1~l!1lJt~"''CI'C 1inw!i(}lli,di1~ wi~h !l~1~'1 m .. gal1J~ic 'Pru1~C~~:;; (13t TIle latest ad \!!lIJ.ces .in d'lC r:[C~d or biomol:cculil'1" ~!ll()itO-l~ iii] l'l,mo.tochll0logy ]tmvc made it clear lhi'!l \','C com ese l!lIDIm pP;:I1.C~I1S dl~ ve IOCiI
I~,mo;;_~~e con~pom."llb1<!i~d Dh'~.\\'e C!l!1i innco~filCc pm]R;:;.in~ NCIl;~~i,,~~I!}' !ID di fttLil.:,lt [I111n(-ri'i!:'i~ \~ ~III! commlilii; j]h~nOJ11 or Illu.110, IJR"liocillfi. tluirn.ugh sel (,;,m&'lC!~lb~y,oolll1nc ~:lCnr Iimmion" and CXOrl dootrrf.cl1~, !Olk'nnicr~l, or i1~lysl~,1 cmm:u!. soirel.iI'I1ICseven over sin;gle prole.nrDS. Th~ ,iS1]lfLl~"(], detl1ClnISlimlll0nS h{lvc"l~owc¥\er. 11.01 yel. been inl~"alcd into IilllC~iol1>11and u.~I.ill devlees, Cmret'llly. dCI1O!.1t!1<ltlnn of p.rotein::; lirnhs Ihe :lif~.h]1c' (I r 3 regl~lilr glidklg, ,a:ssayto scvCJ,'i)~ dla}iS.. Il.l~ Ilmugh scpamti:Ofl 1iIe1.\\'eCIl rabtl:cadon: "mel use

F·iig. 3:. i3i:nmo!or·c!n;VElll tlranlifl'Ort. tA) ~l.utniIJII ~1fI~e cOl'IIfinfnlmt ef motility" ~irnOn ~Jait5lllrrae~, '!!he rI'i.otiklfl '().~ 'in~menl$ is iT! ~~dQn;!, dkero(llFi:>~nlill::n:~~iII~ ilf!~g~.al~lOOIbrN. (U) lQ ~cilflfin~* m~~l~!iI, ~op~ ~rrlMlly t!~ (wp) d100i~l I~Ui~fotll (If mQtoo:~s!!1c1~~ by ~d x's)()r ~mij~t~)ooi!l!!)9faphi(,l!l:s1n!!\l1!!m~!!l1Q'U tt~ :S\J:lJSlia~ES. (B:Mk!ni) S~iJJnl1irngeredrolil: mf:cJ1G:SwP'j (SEM) im<1£JeSliJOIvs mi~lJf,aililrimted dlillmcl's: lfll SiO~.(~IOfJ.. ,oiid'nbinaMcH'!~f Ib~~h noolhMS :PfO.ved m()~ ~ff~ctill~. olli!J:r!'I!i~e ~~ B ~!li!@"if!~J3:!!ld~lluQf~:e!lil:€of3(W!
f1timEHIl\, \lIlli\ii1 ,~ma' mbile a"(illiIsiimy inl tlilt? !l:tl@f..snil!PIlIJI ~oo ~r!liPriritedi wilUrn p;a~;QJ1 '[roll'll (l.9); m~,pyrigliTrl: .004,. ~Iilstitll!.e of Phy5lcs Piub'~st1i'aii~i. UV)ThE! u~, of :MIDm'iool1l'le!.err i~lJlIidkd~illlllli~ oHtlrll '!liim~ 2 d~iiriu~rn&iol1al (Qrnfiif;lt'lif'lieflt. (8Dgm) sm lm~g~ of a ~ an) d!alilnet (BlAlm:rL"""sI1~ped sim(llni'"e$ re(l[aJ!I the liIlotilfty. lllilili1ll~; Uile illOOUmit OIr ,llIIil':ro~ubuff5is eql!li!l 1m beth ~esefll\mrs:, btIIt aflter 18 mill troo)!: mlcmtUlil:ult"5

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driv~ Ihc gW\,,>'I!I.l of mo1!11Y C'Il]O(.'iff.. JI'l'""L'iI.:iclIbr!}, ~h08{' ~n~Jtilll)'il]gc1>w!1~h~ dil.l~UI;!I¥v.i'l~io~!.m~~ st~ldics (if !'It'l111"~ lytnll~OIl~a~;md rt\ik!;Quia.,; indkaJWDh~~ ,m 1e'll:'1 CCl111ia:'l 11ib~I~§liilirtC.lDl f~l'I.i~ 1iI~1Iii'ly lho~~ \!'~~IJISiLih~I!Ulli111Imil1f1~i:1dlly) call,

S~L"E~in~-.d'~y ~.rn~







i11l1ii.iirlt!.."iiUCI!.~(1 lb)lflre~,j[Ti'\!ie],y



!!ire i.I]U,igUed .b~1


eell 1I!~;.r CCIITlJ'!!CF ns tlml ol11,Y!l ~nh~ute ~~rol:lOJ'lionor IJilC cci:lii:~-II m61 in w~ne inmmn aclM[.C myc'l~id !cu'kcrnia (AML!, ~l~f}l~ am .>'(!oo, Il~mnfg;l'~\'\'!h \)i,ih,",IfII:~;Ul;~pl,lnwd iu'liO ~~bk'ltll().~!)'~miIdi,I!\''d, nO!f!(l~Cli1~b;::!ie '(NO!)}
C(1J:r!C'C11~"1r'C hc~~m@i:'nl;.\ol,""


,~i~ionofccutlll in liI.llllOf!l'lJ1ddu.:' flndi

ru~d i~jOCDcd I{]I lei lZY; oelt .. i~110 nO:11imldi,~IOO c(m~cnk <mimls;, ~itc~1l'd~e.~ oJ·II1.e celli m.a DutW.'ir'injoc!oo. sl I ruci~i'::n~ l~llcm1rJl~b!.lnd li.~lith d~.;ciIni[m~r;:.d IYrnJ~hoHm within ,35, d,!~, (T<lbl~ I.}. i\1IhQll§h the m.!~uoor of injocrctl ~C~~!i did nmnmio."'tlbly !lace!. uunor b.mitcn. rmg.rnl infilnrmhm.er di~ Sl:,'\"cr~rj!. feCipiC'ms or or 100 lyml~~on1il1 oo~L~l~"'l!'~·~Y devc~opocl Wl!mgl'~,


,,. IO%~OrlmI1Or,eells, lX'dlOPS eventae Ilt<ljofiily. .AlthOlI£i11 mou~ Ilma hLlI~'U[nlunID:1i'S i.li'Ul:cr in, ~lo1~blc respt..'OL",.~hen'l.'l:L·kc(1 di~'I(;el'[y \\,jlb 1\,'::>\lll'i. rrom. hmmul AM'Lc:en~ (J) sU~$lSdlm:



1I~ltL~r~~ inK~t~~hc:pC'r~



~nbilTl~'d in111ll,1l1od¢f!~iem (sdd) Il'lkXl is OOI;lIIp~ierltoo by alIiL:: crill:crill lilllil~o:rgllQWlh 00 nn:lcIDCli(ms wil.ll~he'mi\'!;"hidl

U), 1]ac illil'C'qlr'C'[UDionof !i~ehx"-~IJmI'fLll!.'WI~.~tlmiO:~1
!>Dt:l{liCs,. liIOWCW)ir.



I(.J~ o;)ll~. lm!i.i~~eiwdr~m]1 (rOf,c).wmp~~,C'lDl33i nhm com~~ Up'I~' 20'¥ti or!B:iC cdu- (4~6~ and ~o:.';lO!mU': R:nl~~'IL~!'1g I.lUS C I},~nphon'tl iJ1 d~l,l·ord~lhl fl,lci.lP'.icl1.tI!; "\Iid]i~~ 3.,3to 1,6 day..s (ease 2). ~li~lI L~ m diRhn::'lJni;:Hti(u~ S1;lg~ tn:l~PliOfUlbie I),y ~'E: m.oUSlcilllic~n\'iro[ll1ilelll TI~c; reponed A. Siirl\:1l,1IrrfL1::l:im~ h2 .~()5%), uJ the ce~J:s in ruri.ty or c."Il[lC!erstem, ee:I ls il1lI\M:L 01 mIld co km: pa;,ll1Iiil}' E~l~'~~!lymp:l~ollJ'l~~ 'C d~slllfuyctl the charIHt'lfC! [hg~~1dUl.~p.'C~\v\lil'!g


o.;'tlt:.'1§~ ~lIn1{lr~s~~t~i!lUI:!il of C(lU.,;.\\iiah oonlln-CJIl hUJIl!(Bill $Ii1li~ !!1It:!tJDN Uhf' Cl'l:;.n:l1plC',. k<ll:~rl.opl'm~. :lU~d bl\."'rulll'.lmm'pl!':ulIU!kllil lllmx."N thc,~umQ:L' 1lJilJ'''\ I~ Ml~,.~ining ~~!:;: wi'11:!i~~ .,,~~~gll\!!alj(J'Jil'ii


~"I,!r.::Il.12f' lk....:tb'roi'fl ;,i~g~~dl a :


are mctlhu:cd by



ael(.'t1S1ticstcl'I1I cell ~llffi.t'kCr:s SOl,. I ilnd/or AI\4.1.

!tIl~ile'i;l!IJdI~llCI:11'bl<l;nle-oom1.dfi!Lcton> (:21. l:Uably. rI,lmlY~ld! meusc jSU::I{I~ lllltllOil cngage the cog~ ~alC' }ulIH1anuc!Oc~ll:O:1" ~md vice VC!:S<l {3t 11ulIs"lhc low Ih.~uell.cy enr h~ ~~.lillli ANI:l cells Ilrodw::ing

II:o\\1C\icr, \'\,llum SIOI1.l>dSca-~+ AA4, I' - or S~:Ji-1.
-_ r,;,


AA4J . lyrnp:ilmJ:m cells WCi['C irJll s:I~Ilan~cd. as Dctcml ~rli~~gwhcthl!:r the glOW~h of v.ru'®,!!~ I~w :IS UI ccll!f,:of c1l'Chp(!lim~:ltionc:liciwd £in;mJ Iy~unphmn:[ wili1in 17 W .'iij d:li:ys ,(TaMe .1~ Sinn· !lI!UIDfS i~~!.I:-i!1!~n'i",,<1 mn....Or'!h\J~~m{lr ccn:-; !!IT ~l}' ~ illlrly., wi~h EjJL-N.RAS lhy,nniel}r~llpll()nw> illl.d 'lily ,[1 mre subpoP!lIlalion has i~npo.L1.mlll :rnll1:mc.tl!'!JIl1IOI'll".in NODlsdd 11ta1.C'e migJ~l, l'CJ!klCt in '~3:t1tlhe liollsfor jhc di.'Sign Or]IOVC~ [lilerllp'les. Thctefbrc, .ronl}' o[hunmll Milar cells thm enn ['e<1d~]y i[!_l_Ii;q~~I. fom Itldcpendel1l, cases AMl eaased by PU.I dC'lick'illC)'. IPCCi:lpient'!! ~!~sp!an~d wi~.has [c\\' a.s !.he eaneer sliClncdl hypotl.!C'Sis merits !mJ!iC rig"10 gR."I\v!h in a trn'CiW1" ~rnouse, mi!~'Ci!J" OOOlI!l, K~~'(I._ fori! I!JIll"U~ lYn1flfl\, 11h~ I'Il;'l!tl;:'!Y 'til is; ~Il om vil;;'~'\!: he rrcqUC'IlC}I (]if (.·dll'i~lml call! •• I (I 'Clillls dC\~I;:!r:Jp<.~d~ll!~l[)rs, "~I!JOIJ!!lh (~n~r;:;~ was wi II n.'qWlht'[n'i!n1d,,'il' (lit' I~U:nQfl~lIs inrtl 11I1i~".l.!' int d(:ll.y~d if! nrn~or;,; ~~ii~'uflg (ll'il~y 10 QiT 1100 AMl '~WI!'imTI] I.mltt~f.~iII:}wlh~ t~MdJlllL~~l~\.,':~.(;n.e!ri~I~ Oif I)' ~,1IDc eaaeer stem ecll hypt'!;'!hc~i:;:., em11tf{';S1 be [csrr;£! ~nuU~l wid1 rr~11ie req~lii~ioc; d 1IIJIlllrlim supipnrl.odk eel I~ (Thbjc ~l F{JrlliU lh~iJ nmlU~I111ll1ei('ii.du:~celli ln~t~y.bc~et'tJlJ.'K: dtc[,,;m~, ''clUlltCr ~lC~.n cell' f~lso b)'trnn:.lh of lilllil!mdnllillt~bcf:iiof nm]!.~sc nJ.lla1t'!:r ~urli~ccm!lrk.cr pl~,;nllmYtPe(:Iiig, 5.]). the gem~c:xCJJn m'llt~y d~.i.i~nmcs the ~lQim\rn.1ell dt~!tI~)lntldl:ld c cdms imo umdmtdiilHzi! h:istomlnpllI:i.br.c IICCI;picrn pre:>S(iimpro"~'c: ,Oig,. SJl, mtd ll~cin~'I!llsivcn.cs." of ~nkc. ,\Ie isol[tlld p:rinlill) pre-BIB Iy mpho.unfl th(!'ll'fmsp~mJ~r:rl 1.llImo:us ftr~iIlO!UI!::dlhm of Ihcpl:iDhc~~mor;wli! ~Llgg.cst that IlIlCoclis :Sll~mill1li~g gro'W!~lof an csmMishoo ~LlfI'n()rbcreferred was C'c:Us fi:o:m 111&'(; inclcp;:l1.dcmEp~NJ'(: ulll1sgem~~c 01!ll1f'jo' Ilumm. "1 urn.o:l:'-propagalillig cc:llls. ,.,

mnct.~' ~4) migl!! flCj'J~~.lh>i:' n!OC'd!Q !llQttrtinli'if~ran csscmiJI il uman accesw.ry QC:I ~ {we 1l0~C' t~~:tI11:111do111clillil cli!l~~JlOgel1i;tors are nllto CD 13.3'''"),


1.• A ~rg;e IprO[p()ukifl.~J tij,llrlor (e~ls <a1!1~i..i~aiii!1 ~ gMwl!h 'of l['iuI!IFii'ii;i lymphoid aild ~lold ma~igll1ar1CM~. ri['~11in pnm~!J' ~~L-myr QIIs prl!--IW ~~;mp'h(lma~·j EJIL-IUM5 thYilfliic i¥mJliniiJ:rl1~ Dr PU:l+ AMIIL, an from rmke em iI (S7BLJ6 {~'5.a" bac~g~ol,J!'1d {>:15o bad:'cJoo~s)', lo!.\ere uansp1all:!t~d iiliiu~ iI1'~F!]mf.a;di,,~iRI ,~m.!,!:g~niil: C57B!J6 (LV5.:J.+) .V'Cd,pi!l'!iU miol? To dr,c~fffi!'em lP~ob!ems ~issoci"tI.!d iMth ililljediol'l ,td Low oerullillJlmlJeffi, ~'o'@ rniillld ~hehllm()r'c@lb Mth Jl!0i~ ~!llI1l~lllilk(C57 BLrlFl.y5,. ill"'1 ~,pls@;n«'!lliI <lIS rnrJl!en" Shown al~e the frra [1n0t1ll 'o~ f€(llpiemlts that developed tumors, ,Cir:ld the a,Vera;§jftime h01llil1 ~r(lnspllllnLilt~oll1 ttl' 'Il!JmQrde-mQilJlllI:nt N'D'rnib (Oj24}inr~cted wi~h ti!rri~r spLe~lII~ells akllilJe dewtClped

,ilInry tumor ®\I'!Ilra lOO'-dal(perhxll"

N'D, not det>e'rmi'noo"

lL. t:. J, IflliP!i'. !.. lin', ]. i~ ~ic!(, N~l. .,",mr.llJal S" 738 ~l; 2. 0.. Hi!~~!1!il!'l~ f<. lJ'!leiri!i!~!!!.'dll!!~, 57 tWOOl. !l. ;1_ K. '- At;li tI' I!l{, .M~u, ~li:. !'lNld~nt 59'. 700 ~1~:9~»>_ .. oj!. Co A, O'8i![tn, Pi. i?!::Ill:tll. . 'GoJI~llgl!'; ]', Eo ~'k" W~Mf' 5 ~S, .1& f2.G(I'~)" 5. S, [_ 5iiog11 ~~ r;tl,. JilrJMe ~';I~,39'6 fZOO4». 6. i\l ~aj, M" ~ WKm." ,Ao 8eflljjl>f!em'IIMh~ SOl J. Mor~1 M. f:. (wt;e, fifO<. mil(. ~ xi UjA 100, 315'63


ICase 1, ICa:5~ :2 ICiil::i~ 3$ii;a~1 + AA~,l hi S4;(ii!-l '" AA~.l ~

R!~di»i!ll'JII!:~ tlii~td~¥(lI(lil\l~1lI1!lmor~~d'~15. IkijiU} t t,() ,~~~m!ii lIr !ICIe1Jl~ 11(10:l;elU;s


1(1 "e~1$
,Z12 (35)

,E~i"imvc,B Iympilo:lfla
3:J3 (25)



3t3 (:2S) 31'3 (,23)

~1f3 (3Z) ,3\1'3{24~

~\f3 (21) .212 (:i! I)

3(3 (2,i)

212 {2;~n
PU.1..,!'- ~t 212 ('t6.$~

IN I) 212 (26)


(24J ~13 (::i.l) 212 (40)

SIlliPPOl'tingl olll'l,ilile M"~~iri,;d iMWr~,i~fi'ilIil,aiMgil\:Oi'ilti'llil'iJrllii3l.ij$1l3~331J1OCll fIIiIlei'M15 ,ilOO Mif!:Utiad~, FII!Jli. S1 ~!b~ S;,
l!!d!!retl~~, 01001fj,0!~

is: Marcl!i

200,7; acc~lll~




tWiN- AAS, T ~Yn.~pn(l'm.Q .313 {;;:a~ 313 {4:n
,Case 1
'C~$~ :;!


m ~!!'I4)
Vt ~!!'IS)
'V:J, (30)

m (54·)

V~ (;a:S)
2J~ {~2) 1Jl (3,7)

,m (n4)
:212 (M) .21.2 (;;~~

V.2. U.9:;!)

'C~$~ ;I (a$e4

m (U4) m (90)
.212 (B~O


IWa'lIflU a;rnd EliM Hall liulitune 01 ME'd~1I1I1 R~i~rcHll, Meloourne ~QSlO, A;I.IHralia.~Oe.pari!lmel'lt ()~Mtld1,cal, B.io1.ogy, tJlm~Ydlr~il.yof MJ!!lb~r~~, fMlb~mn~ 3051(1,
'''I'~~ .<IuthllB ({ll!IJtniootl!d ~quO!ll~m d~~ sludy ..

t'['o ~h(lm (OfU'tsp:uder!(;e

$loouli!1 :ooe ,al!!d~e~~. E-I1i1ai1i:



--I I



C,o,mrno'lnSeqll'B'nce P'oly'mor,phisms Shalpin'g G:enetic Div'B'lr:silty in A,r"bidop's,;s ths'lisn8
Rkh~,"dl ,M:,,tlaJr\I!;;/l ~a~ri~le S<hwe~k~rtt~ J,2.~~ ,0 i1'i$t(iphe:~r!l'i{iiliilil,jiiil P'I,~" SteIPh,(!Iii!1 O$$'O.W5~Vl~
'Ge(lt~!Il1 :Ziiilll~r,,:l!,~·5~ P,allll ~ hii n 11,6 Norin'aif'l Wa,rthmam iii.1 Ti~'I.i!1T., G'iil!l,4 mC'1Ii'1'UI I~u." ID'o'II,viidA. IHi~d:s"i' tlIllII!alii~~n91(h,il!!l,:~ Kll!1lll.y A. Fraizetr, '1 D.a,IIII~e'l. ~. H11I:IlOIli,,5, BIi'l'rnhaud SClho:mk.opf',,3 MaJg IlIID!> IN:o,u~bGI"9,'~ 'Gliltl'llnar FI.a1ts(I'I.,·~ ]ll!seiP:1iI It Ed:elr:,~a DeUef Wei"ge:It.Jt

~~~!I]I;l!!;1 wi!h SII.npiPort ~!.Ilf

\~!c :Ll!ood~'CkJlli1:'d ~ n~dl~w '~'I:ning; {ML) nJ<l!.clril].C~ (8', QJ!b:r


~pk!;i~i1t~~2m(D d(lia, 31d .~""il~~~~ v...; ~i~'tI, ~dhml>1liQn rnr;!U oJiigom~]waid.C' pro~ COm.'~l

icl'~ltlli~(,;Ilio!1!(71 (F!!§!'-S3 i.o SR~ Th~ 'J<Li~inlg
poi<i~io:r~, ~'fj1hlin;! 9~bp \vim;Bn'lvo~
~U~1r, We


'!e.oo. Oil cal~iWt~


lih~ g~nClme-s Q~ilildlMdLlillhhQm tile ':ioij me~eJ;,ie-s, 1;\!i!wyin: 5equen~ as a r'~lII~t or diU,e'ftntl ev!JllUllltlnl'!Jy IIlr'Ooo;sse'S. ~(1 e~:a:min~ tl:1~ pill1xlflllS 'ct(!lnd~!h~ f'OIn:e~ shap,Ing, ;s:equ~n.(;eV,tJiriaV!OIflin Atw1dr):pSi'~ ;thrJtiolW, we liJ~rf,omrted h~91l1·d!lln~lyCiHa1f [!lt5:eQ,uen(iI19 Qr 20 ,a~,w'rse :>~rainli h".i(Je5:~lQIIl~)., .M@re ihatrl 1 miltl.arJi Iil@medh.llnlllialllt siIil91~e:-',m.l!l;l~t1tffde Q'~1ymQrp~~:Ftmls(SNIP's} WeiF'e h:Il!ii.. lilri~d !ill mDd'erat:e ~iiI~se P !tUsoe\l'~rytat,!!,~ (1FID:Rs.). nd -4~ .of the '~~l1o!mtwas idmtUied as Ibl'ingl h1,9My dissimiialF 'or deleted a ~,!.lIalwe'~~h~ n.lte~!Ke g~li1~m~ seq~INl~~, PC!tt~r~ ~f polymoOrphgsiilIl are highly i'I@mOl!!ildQm ,(!tiIIl~i'lg i'1l'~F!~f'!!lmiUi.i!:S. 'g~iil~ :med~~lirl~t 'i!l~,!,lr:1ll(lt(l!1l wri~ th~ biOOik ~PI!:vir(i!'!im~~t hi!\I'ing e;li~~ptiQ.i'I~~ ~~h l!iI~lym@rp:ltiSimlel!i'e~s;,At th~ c~ror!llQSolif!.1)ls~~t~. r~i()~,g;l. va~!§I~Qiil pd-YID!)rphiisml W~5 nla~ily ~~ ,1I1p;!)!;Irelil~. A,5i(~~f~r f~o~rlt ~~,w1I,1~5\~~p5 re~~~d ~v~,~!'ll (i3l1:didIPte f1egilDlft~ il1dllldi1'lg ,;;III!lCitable leXilmp~.ein whid',[lIIi1'IQ$~. aU varialian W>!l$r~mol,ll;!dl i~ ;;I' 500·l!.:i~0ba~e willirlow,~lyzili!'fjlh,e poJlymorpt1i'lim~ ''!Ie d~c::ribe iiil Larg'Br $ets of aue,,>si)r,ms. ~viLlellllllbl,e, ill dBtaiLed ~md\e~tam:l~ing ,of'Forc~s ~hilpln91POJ1I1da'li'aI1·wi~!l' :!lE1!qlil.eme v.ariailiiDn:in: A. lJudimm. 'l~

~lIc:ooe ~h~.cril>1:ies 1mC~ ,ofpRll'i!!.'S ,~!nb1I.'SS:S and S6}, 'nl.': Ml m",nl:rod 11i.~;UgIb> apl'dlMlb.i1ity to !..ooch l)tredict[1:)I],and ''iNC' !;'C1k."'-'i1uNl44O,!o 1.074,055 l]mlll'>dmKilu U,SNP 7to p'.tldoc~iCfJl.~ m ct' :loorrespnlldins 1<I:~gC' ol·FDRs ~1rorn.2 1:0 10% p. itJ',. [~~rrClrmi!!l!Icc ML rm.1hod was i~~icrigr Ill(_! M B mCl1;hcild for OO(~. ito iug t-"ctIIIIIC!!{1~ blli~ SIIIIIIoC"ior for :1l1O'Hcoding sequ!m~~ {Fig, t. A.1~nd ,md mhl\; S3). B, W11(.'1]dle FDRfiol'lhc Ml :L1li.:ilhod "\,,~ .a a 2%. .~\'C' FDR. ,imi :rt(,"(Ivcry IbrtllL.lMl aad MB H~~I!Od'l simii8Jr: hL'l\\IL;;"\,ttl~ 1!11(;l~W(l lli!!1tb1Od~\~ (li::ln'!W!~'ill~.nI(lJY,. will] {iO'% orllJIOOtc~iu.m;n1~ willI only OljC of Ihe ~~.1G1ltod;;'Wig •. ~Ct lllls ]'('SWIJled, ill

roJ'YI~'iO~hi~'i!;i Ii}, h'l1Ld'dilnm iljcl~l::ll1c.1 !I:> illl'iUL" S-!.1flild gea'L{i!llfDO.\vide repd:i~i e-


~"'' 'fl.'


frem dim;ril'i.!llperlbn~:mncein


(I), D~itlJ .~ \"'ill:l~·W !J>;~n~okl!,!~. di!1.."1;:! 1'0'''~lll~1~in!l tlr!Il(~\lidlil,I1 ~Kmbi l" mJil y(JI. 'l"{~




m' ~n-

Ihf'~hc sy,,,tcl1tlli: id'!."'n~inOftKmof se-

~amn~ ,[~nrt'(1i~~ 1~1tlt!O~F($

("Ilb::~ivcIDl_' n![l~1 OIgIl!i'-:nlS 12), Hig'lKbt:lil? ,ofigorluck'otidc UWiJ)':ii i'Mi!)ll;'idb ;)II, n ~rm~~ivc;)~~d! fiT po!.y!l,..,..hi'illlli ,OC'1'lL'!;;1i( ] "md I";l\'~ ibx1] ~I~ to idL~...
~i~flb~ fr-dCUitmofUlt:
SNf\'1lLtUH1iOll ~lris,

phi!.; n:-giol'l.'> ~Fig,. ~.Dt Roc:iJilfOr SoNP:s Iloore llmnl ~30 bp lkMu iillothcr SNPo:r ;ill1~titi(m or doC'~ltkm, ,(im:l'cl) u,'JS higllcr .tbT Lite MB lucd'lOdJ. '~h~.t'tas diversity :(6, 7). In Jdditocm, \'~C chose Sl"!'C,1flI1 m::nHI lor SNPs s.Cl~arnICcl ~y 7 ~o .l{]l b,p foom a cOJ11!!l1on~)'used !\ul<!i~:-;,. sl!clll rul. u'1'-1 (I:dl!c S ~carl~r ~d,ymorphisln W:!.'5,al1!Oln 1\\'0 tin'C." :l" C'ol.(l',~hc refuro!'~O\;' ~iiol~" wa." illdudcd Oil' 41 .hilh fur dw[\;lil ul!:lh~LfO!' V~' drn;cly I~]'k!."j St\J Ps (<:: 7 bpl fCCiO\'l:~'Y was ~O\",'IMhhllodll1mctrlliJds co!'!I!i(lI. for 1'9 of d'!.'!)20 ;!C~"(·$,\~on:,~ (11, nu. r!'<~grl!".ns or ...5(OmM~ Imil);; [bp) i~1(:1~j:J1I1I.wJ!k11 ,[-3%). ffiiRsthr both methods :pcakcd ill regLolls • CinO~'1h)'bridi;t<JItiOll quJiJ~l}' ,Wig. ~is). an «.:l1bc~ or were SfI;1re~d~rol!!g,hol,ll~1C: gC!1.O!TlC, ]md pl'I.'\' t{)~ scq~rC!"ICC di\'\lf'J,I';H'CC b,\Ul <lIS!;) or otTrnC'f J!1C'[{lTS (flo. bl 1;rIt.;"'1;~~Srnl!l(l!td ~]}' diklox).' \1~l!t;I!c:ill!g:b:.'twei;n


ill abChiJIlil::IJJa

.1lml ~IC







or I'l lJialfmw, [or wlliClla ,!!t'I'tOIllC soqU!ltre Iffim a sillgk: ,!iCic\";'~OIl\v[1S .g..1Jl. C13JlLi'cIill ~l! ycflf 200[1 !t5~ m,.~n.~I~:tU£ Inl~1I1ilOt~ i pM;;llii dan.9i sell' 1,;"tp:tuL-snl.ltl'dl Otr the OOnmlQli~ SI."qllCllCC vamllm in~lC: \"-lOtlldwidc .<1,flmlitmtl I'OOJI}~il.1n.iOfl. We u!lerllihis, ",!(nnmtion m ~1CIl~g~~i -ddx'fITlinc ~hl:'Lym]($, ,od\~'l;qoo~~.,and ~'C!~~hm (~~er 20 lj,\(uld~~im't'O





!{li'1P1U'IItdi::,m, .hi,gIl"JL'S!JlWItiO!l~

d~ptk:m 001' the ~nJrc--widc d~mm~iot::a <of 1:!Olbt~ rllOl'llhh1ms in~li5 nll!!~i.ce~"~br uclCl~t-.:;e O'IY<l!ll:::ll,ll. S:aJm~'t~ s~ltdiOi!l!"i1!~ray d!~,ig n" iilnd pOlym~;rrpMSIi!i! d'e!tectrQI!il. farpol~n~f.IfI1hR:;nil_ di~"a (;1)'. \\-'():~k'CIl;'d !Jf;;l,1;.>:\.",IOU!t5Wilh m:llL"ht;;i ~gl,!illL.'[ic
lDt!iJloi1dIlI,M {II fOO'li!rnliIr 18idlogy, Mail! PlillKik IInsliilUli? Icr tDe~lCl!iii'iert!alB~. 12(116 VLi!iiligeJill"Gmm!iJl'. ~Mi1dJ1ien' Ml~!l:liIeI' I!.abo~"'l~ a-f iOtHe Ml~ Planc~ SKfet!!', 72076




m lrn:pinqa~.Infm!OOtt ,~

iP,i.a!'d ImMMelCf

BI~ogka.l. 'C)bernelics:, nrl16 TUb'i!!l!l,er!,

Gemlilm~, ~M~e~ulo;ira~d Compu~~~QfI~,B-icfogy, IJI1WEf:li~ t o:~.1lootliH!1Tl1 (alif'Orn~ L>D~W'!l~~5, Cl\ 9000~', USA. ~(illrlll@r 1

~or BiQ;l!1hlm:lI!Ci'Wllli~n,!lii!ll_filoefl' ~1fI!vem~ir!'. 7l076, G~~!fiY" ~~~i( ~r~, ~OO~"'!T'/, 1I~ S~ ,IfISI!~l!Jl.@for &l~lOgl~~. Sl~d'"m~ D.iI ~{ilal.Gl 920~'. USA. 7Pert~gt!il SdeMes, 2:02:1 Stie,!'iLi!'!Coo!,!, ,~ntaln 1{;1~ !CA '9~M3, U);!'i,. ll~t.I!:lD~ ~11l}', 1h~ ~111lnst,ru!l! r~

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92o.~7. UIiA.

mullm f6lli1riliH.JtJ!clt 'J!~)!J\illly 1.0 nlis wM::. flO M10m (!orri15P'Jndern:~' ~hlJlIIl'd be- ilddrfilied. :E:'rTlai~ 1i'jil!!lf!l@,_~~~MIiJ!![ll.Ic~Ld.or!i

MIl r[(,dlit~l!ion~ wil.i~1'!;/iL ~Jrodletions ::illlPpol1cd d.."LTIliL SCI, etllk"tl '·2010:' nllmllloo. wr;fOM:;(~S lite m. <'1.2% FD.Ft The 111(;S'II~l~~m d~iBr~ "MBML2," ~r:I, eoesisted o:f(,.18.570!lOltrodmldfllll SNP", (7. f(Ij qlll1fllnryo[",oor 'PO~n~rph~1i1 r~redielh:m;." \i\flllo~l'-geuu;1inneaii"l~plmed DNA .fl:om each !Wig. 2). on mlc~~ of One po.l~morph~che per s aeeesslon w3s,hybridiZl::d to f(,",se<lJilI.cncill1.g miUi6 1l.0I1rtl)clith/C posi~bo:ns bliliae gcnome. ~Villli= cl"OO,mys :iil1.teit1!Og,\'Lti~l:g ;:.~9.'99%efbases nll~lilC in. MBML2. SNP:s ::m:ppo:rted by lilOlh methods 11i9,-M:b 1:e:1i'iI.'eIl~egenoI'l1Lc~qliC~C'JC 7). Each 110( tmve a l,'cuyl:ow F DR of -tl2 Yo. whereas SNrs stliPportOO by oiil~Y enc 111i:ilhod have cQ:rn..'Si]~n(J!. si~iOlil \\',a_" ql!lJ;'ll.1.cd whh ~bn"{ru:d" ru~di :re'<\Q~ld ingly higher 1,:nRs (l·a!hlel and m!bl.e S3). l\ j)ro'bc qmtrlf1~:),oo!1SL'i~i~~g ol.igOll~dOOI~d~ o:f caveat lilr our 'en:o:1.' St:h11lll~l;:S, is IhlU ,201lJ &m. C leH,g;~h25 ui~. s,·~x\i\rhhl~l pmoo :qUiMlI.'!; ;~n(our ".~h ~"'IC tJlsm ~bir$f.lIi."Cmch) ,::nui~Illsitivhy <15ie'l:! ~,lll!clooHdC'5'\w.e rqm.:seimiXl,;u, Ilw O::n1mlllXJ\jii" flO!]. and ,d~~lO;."$ in 1:cIaHvc hllell.';~ties ~t{k~ :'l.".Ssnmel1t or'~ile~wo Ilredlctio;n mcdmdti, mv tJln~ ~l~dbl: ,qlllill·tc:t... indk;~~f;"d ~wmi",1 SNI:l.s_r'Oll:~igh!.· di',Jm..1l!'\.~m,-icl, r:or !]~J~inJil.scq~L~nc~, :llI;P~""~' l iy Ih~k«i SNf!\, .111,JWI;;,\'l,.'l:, all p'.J.u1~ 1:wIx:ll' ,,,-I I",,;.:'it wld ;)equ .."J~('~ n(:!~ ~1:mH~u:<Olhg; .llCrt:'~n1;;c (6, 7). on(" !'n~ua~ud:r. hyb·tdi;aJ~.L'01] L., MIPfll\.,,-~t ,111111;1 Ap:'!rt Jru:n SNP.s" dl'k1:~nl\ nr :.;.;.-.qUI.11.l,1,'.Sill.ghly dissil!llilklfto ~'I('~j('11'i11{'o; lire dCtoc!f.1;!I~ hiigh· SNP OO:CClL\l!1], R.<; 'ltmjbWllldli'tl Uig. SL ~ \~~ iIL..oo twO COlifipwilmionoll n1atlr'bads to U{11l:;'C[ ~l>Ury rum}'::> {ill> fC=g_rfills ,of rodU;Xi-a ~.),triilizml(Ju U I Hllg. S de~;;;Io]X'l(h, 11l;,mri.'aic illgQrlthm to SNfs l:ni o.5i.9:W.272 pos~ttorn thm WeJ'C 11m highly kk1-ui~~~ IIll1C1S or ~l.Iccd hQlbJjj~2lltiro:F;I C;;Y;[Cooktg ~ili\'C'l7) liable 52 rmClIt._g.52). In;1. llIt:IJkmt!. ()\'\,'I" i1il~ dJiII'I -lOO 'bp (i') "I~ SQ rulclsao~ l1tc the 9.'qlllC\llCC roffii~JOsilioJl (,i:e~. OC OOfIili.'Dl) .and In-.'tI,m, kllgjh, ofl3A 'JOpolyt.llrn:phi!:~itx! p:ru-~ow polymol:phifilt~ ~evcls typ~,c;]1for coding dK;~km:s (PR.!?:;) ,!pc~!fl(l3t;Ii'O:lS an aoo.--s..;;iouls.Wj~l, ~~1C1:'S iml IID"Drnb~ fOl" hy1?n~i;m!1i(ln"'~<lScd ~li-> ;JiIgori~hmw;t> S:-.'9bp; Ihe 'b.n~:S1 \\i,E> 1I·1.2. kb S;N~.]' dc~"ctiOJl l\OOOJd~,gly. rocovay 0 rSN Ps with a. model-b<l!$e{I, (M EJ.);)1~'rjlllH:t1,~J. 4) \1{;JS (J{J). ~nti~ :201:(J; daIrI :ilC1:, \\~lkh W,IS a~t.'ii.l'Cd b}' polYl1lO1':tl::',eh.. flWction lK:R}. I1URlg OOliMIDIU"" 'lin h ~,gIu~rf'Or cooi ng~llmn tor l1oncooi Ilsregi:o:m; (36 VCrsLl(.';: ~51i%) ato! c-om::sJloF!lding FDR~lllIl Wi!:ii. ~d 111p;ut ro high!y polY,nlOlpilk or (klldlw only ol~~~hilicil .. h~£h (lFi.g. I A}. A!I~'\I~m.g.¢ a of rr.;g~15. COIl ...~renl wi!Jli1iSll ~'t'md~~'tw PRPs., i a 96,~ 14 SNI\, ~,\'()~~dCi~~i i"Cr,,~'I;o~~im! i]u; Oro by 162~rokl m~~~I'!liuior~ ~""4S bd\\<Wl PRP:>~rlld ~d~1'i1!OOl:a iri2{Wl W,;; ;.l<;D .1LU~l'iP1C1l l'llm~'iI.'Ui~o~1i, I(]i~~ lQ~1.1f 4S6.936 DlMI'Gclkubdgltt o \oaldlinii otlrJS2 ']tP..~1 KiR ,ru'!d. ~U::;1icinbg. fibSNIP's (Fi(g. m Mtlill!ribk S3 t

22M llInd ]94,9 ItU,"lootL'dC Nl:1~'l'l1i!luli:cm~ p(ct ~"'" r.!ol'l IfC:hnii\re~r)['!'!I.Oh!lct been i:d(;11ti~iL;d (6)_n~:.;

For silllb~llti,)lJ1l~~nl,dy~,.\""t.:

(lOll11biniDd, ,,~'I







~I)xl! oolnpklC (lfp..;i!l1i<!1 ~tl:!flQC d;:n."Ifor 1711lID"ducts. <I!ld id'cniil~cd !24 d>c~"{io~~rnrlgi'~g f~~m 50 bp'~~ n'lllfiC ~~ Hl.~ ~~"", m!!Irr ~~. PRJ's ~'sporndoo:!D d~lSIJ;'iI!; ofs.:\'!Ps a: sn!llin~~I~ (~~ S 11.~ Mnnyd:l~ntlins or dgJ:;;t(:r..of iD~ynrDorphi.·stth'" C'xb.snclcd oo)loW!(i PRP bPl.!l!d"'"~ pn~nimJ~r OO!l~ ~riblllm~ nl~ I~~I ~lU~lIn; fov v..li~~i~ "~.a !O rnle h.,n'il~ (-55"%~Wlt{lt S~'\1!JI~i!~ Wli!t '\M,~ a'flllh1Jb:k.; 9K&'{\"Q of ba.~ in r::R~ \V~~.ru eillk~1' dd~i;-d [if wh~linl 6 ~p Qf 9 SNP OJ' .~ldel j]t)1:Yml.10ijJlh:iSlil1 ~ 71. Nearly 4,.~% o.ftl~e rer~.nre &~O:il'C S!..'('fl~itre\!i':fi.'" illdt(l.ccli in PllP~.wi!h 'lml~~oJ'1lind p~ctldoo~,t!I~C
tlI!JI!1S WoC],\: 'pn.~ icted 1.0

be ~h.:&m'lC 31> theN~: erence i~1 ~c dit'brC'!!~'1 accessions (mb~cSSt 'ili.Ve Q~l¥nll~!1~-d U].!,.,"W!l,;fG!~N1~b!I'i'i~ 1,;<I!Is.wi~~MBM'U

~Hvgh! difl'cr

~o W!ll~rn!~


cacb pi' tl~~2!Om)CC!f"\;litlOn~ ~1 fJl,

EH~ds: I(lif pi)~Yl'!~iill~~lhism$ n 'genes, To o


":S:.:I~'"td ~fI~("ls ofuhc:

ill 11.


SNfsoo (l2~ r.iii ~1:~fiGii1jliO S!\IPs ~"ti!till!il d~1l(JllJ.979 ruli,~~~10 aed ~1:li~l.Jj;"''S. kbl'ljr~ lmii~y SN[Js Wilh I~ we e lTecl:; 0:11£;Cf1Cilll~~ity. II~ elia!>.<>, i227 iilllffidtlCC tl~is, 1 :roglliCH0C5: O\~cm.1)D\''S\,'1uro J5-lolcli [Fi£;.2 ~ "reIi~'mllllC ~1tJt)ceders, i5rl al[Cr~lli.t~nl!()11. f~lC'IJrlio" ~pJfce To cmllple~m'.nl, IJiDI)lnmlphb~n IUf!i.'<i~c~iol1~. nirc lL,..,-ioolS, ru'1(14l5 k'ad t'Jno:nrtlll:HlOlloll ~"''l:!de;vclk;i~}i;.>d a ba.."l~,all il1~ .dgorilillmlo ide'HI il)! dm1Ol' er "~~Wl!' !:iL~ (HI) (t.~!~ S,9). f\li1o. 1t)3 ~oil1iitioPl~iclcn!~c<lltn ~ht, r~rerel1cc ''It i('ili'l' fDR!'. SNh 1tl!11.tW'C mUl{MmOO ~"o;1Pt.'OO0IJ1S. ro:>l!~~~l.S ~l I( n B,jJ,'!:~\·~'¢.!:l£'03 ,md <J2.J%of,~"Oding pQ(ii!!kl!1~ tlJil!g!..'f (1pi)11 ~ing r!<~lJl"'" jh'1.l!1~i1'Ll kl!~':-d'ft:t! ,iU1d 1X-hn'!l.'11 39.7 ;1114 ,6,L2% [I.ri.~Ut.'rg¢jlic rtlif;~SNPio; ~. I;}:.po:.~ W! boI,:' ur~L':I!lI!:lI:1J1('mi!1 II"&.! ~'I£:!!11:e

:iJion~clUr'i[11[Uil MBMl2 1!1.1c]ld411UilD1l[ijtooi p-DllI.;il1.-ro:iitig g¢OC"'~

l;n·.dorni!l!IHldy ~D:~l :SNP' I~·{llls,. a.,,~y~ ')1m \\i~ pllt;difiit1M~ d~"l:i1y {mMc S9J, Oid~Jx'y~!J~TtCi~lg ""dioa1.oo 6:50 SNh.i~lu;l$r!,g 41J n.~i!l:~I~ it] ~"Hl;nl,l(:!'; ~Ofi oodom:; (!..'lb[i¢ S 10 I. At 1.J%, ·th~RJ)R Ihf~~-L;"ft'be~ SJ'\lPs i~Ili!'il:;k.'flil'C'ly

<111SN~s" FIlI.t;;;: ibr Ihjs. SNP \~!lOCl.]s.o;;; rm'nd~t rbr J\IlBMU ~ "wh,i~, lh !1i!:!1?011~ tI~l~ibu~t)' !11~"!~IJ1.~lTcct SNP_<;~1!1,i'!;l1




l1iglJi.Yt~Iai11 m'i :lv(~rn~ SNf' [or

~'i MBiM.L2

( -'~I'i

0CiI9lI614~'I!.~ ~wOO:rru.lca;j. (u:iC hli~,.,().\:jrCd.SNf. hI atldI1~. lll:! codii1! It,'YI01'i!> I.1fl t'!H .~~ ~Ii:'e!i1 l"nS'l pm'liaUy iltdtKIcd; in ~RI:Is: '~liIl is, .~1qffi'I! hi,~lhIY~~l'K'il1l~hic o:rdcl!el'ad. 1iI~ c;'!,'!::I'b4) l~tvI'OC'Ill .~.\'C' l"\'O dl~~ ~. ~!!lCil' 'lh:Ul ,e.'!:~~cd by d~1!11CC

,tt .. 186.1.eJll"", 2.1"<

or .... IIJ(,I~i(;m~pr{luin-L;::(lding L
~UPllOr!. (;(2 = 2~92. 2 ~ti,JS (X = 25ti4.\

I{f~~ TCfJetJ'er.. bl~~rrCCl, SNIIP,;;'l!:10 PRPs"hcuwniY :I:'l:libm:d ~ !WlliJ}~)1 as ''nl~~Ol'-eI'lC(:l(h!all~~." .. mb,,~~'{! 495. or 9.4%. 2

of ~1.'Ile,.,:;hrntWlfh~g: m<lJ'OK-fIbct

0..00 '


dmn~ \1:!ri('.'-1·~igni[it<lmlyaccQHJ~.!gtoo1l1!u;Malijol1i elf= 2, P< 10-2°)\ dIl!3)li(,'<uijO!'l df = 2. P < ur::ilO). :iliad ~'\inc UIlmi~y (Jr.:! = 3 U,6.. (If= 12,.P <. I(r2Q~ lfl~!}3A),

lOeOO 0,00


AI C<:otlOlg

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,m):d OC-peluli ....e ~.'mlJ]Jn d4Jc."



appreci,LlJly cI'II!~!gc:ihe ob..';('I'\,loo p3IDC'.ii'lli:, (fig. SII). By a~lIIwmlkill, !l.!inCS :k1ll:riV:I1 W be ,exp1:t'SSCd bn :Iacld~lg rmtl:li;mmll S'LlJlp<):l:l or i~~£~lhcn:tology 1,"E:;(plll.."\l.<;cd m~knlfiwn'), as \vcll ~ ~'Jl~ wiltimill.
C%1".t1..~io:n SIJ!I)JJ!IJllI. f'N(I~C1'!I'!llI."SS~'I!I"")" .rl~ O\;i!r~ rcp~e!lt!.lcL ]n~dli~iolll. of~36 A. ilraJit~uag~n,.;'~ tll.at cUth.cr hlCik or have OIlly nrodl::rnlc ,simiJJJ11y



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fiig.l. Gcpmpeni&)liII of !iN JiI ,de,t-edi~1l me'tilPdl:. (A) F;[i),R,·rle~nder!t reoo~~ ,~~48,.7,00 ~!'Im'li'l SN~ fi~ 2'{)CLO r1ragm@nts byl.iJle MlBalfldl MIL _tnIilOs:. iBoo'!l!~ ~rs~1~ S1i!mlPlc- 'mef~r lI!!IIbia~lal~d [legion MR)SrNJPs 'in .lOJlJO, 'thi:!i '!\lroup 'wa~ (.tIf1fIbiliu:d \~ilillllil!rotl ~lJellce!>. (Q~ mR.depe;n;rlen,~ .rE!().overy i!lffQ~ltIe' en.tRgeoom:~


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bell.i'i'oon ~nome"lrme MEl ,and ,JiI"Lcall1s .I![r(l~ illl.aa:~s,ibn5.. U.'Ii) Rl%:OYe~ rlrffJOOn~ for :SINIP5,as,[I fllInfil'ol'l Oir 1~~stOO(~'WI U'!e ~rle<$t po~~rpffi(; f~WI1l',AniJ.I~~i~ W.i;'lS b:!~~1 ~!Ilzli)mo :S!1W$wii~Nffkhmt fll!!nlk-iiFI!ij inf;orrniltb:m 'lQilJissi@n bin liI1I!!milillrmip.5ilmpiJe! si~~, per Ib~n tlre~hD'l\ln ;attn!! 'Wp. ~E} IFDRsfor MB ,aDOCi Ml ~1I@dktim~ a.s ahmI:tiOf!I of t~e 1lfL£Il n quality SOl:re fG)~ ~eriCIJIl'j\[lJd-ami re!,fe~-5IJr(!fid pmile q,l1.ull:!l!..

melhoo~ iO\ii~h pilll:~nlllBdi'!ilat~ f~m 2CI0. 1~ 1~Jl)R b all SNiPss also Igiv~FIwilh g oo'rec~!!lii1lfor i b(lt~ ~tood5 t(i) amumfoJ lite diffl?,rents;e~l!!eoc:e cl)lfIIIposttklllll of 2:O!LO (jjrl~tli1e whole g.i?,rI~lfIIIe. IQvci'lap {O'

~y both

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~Ii'~ !!HiM137


F'iS" 2,. :[llim~'!;!ll!U(lJil I~f SNP:sirl MllMl2 and piJ,s:it:ionsi mdl!Hi,ed ~I'I :I1"R.Ps I:g,m p~nllcil \'liit~ :!ll1!qU@IiICie:l] tit@ti en an<\!{:!l.






.PbpurJr,~ l,i~·h'(j(m7Jr.r {ij,lhc C:~~SI ~~("Irnc t(:l ,0:1, ~h.aJimm UJ),:;!fl,{JII4

i!O'!.roo,· n~j,~T·~!l!;lc~ ~hw~!~'S, o"ml""~i;-d',viH~

~f1ln other gen~. P~f gcmc I:!.!Illml<ll.ioo ~ik~l}' oo:n~ribul~;;; t(l!hi:l' 1?f1!:ct. b"lJ~ rnp:id gCl!le CVO!i,J~ipl1 V!!;ill' Olif,O U~I;uy ." 1~;Llt
8, 9%

C'Clm'~icru '!;tfl,tll ",i~w rNll!~ryffig ;;!Cwkln lQ~c ~Jgll,,~li ~i; d~{lrurrnii(ln (l$~. ~n daw!ii~'" 1IR' .14- ~I-Q I.,ji'·.. m~~~ .~

,1.. SNP~ ~d:el'!tHied

per a('~~:5~!lln 'il'!l MBIliU:2 \vithl ;rlll;l'Laml n'!cm.rerv a5i5~sli:ed agi:liiosl 20UI.

M,e\ililil litO. SNIPs p;erac~eslli:'1Ili by lIil.e,nlmd [i!d,e<lilil IFD:R: ((lJ~ll,Mean reco'l!\e:ry (~\Ml (ooiilllg

!1~jPr-t~Ot"d~!1111~('i>n;~1ii\~ ~ifJl ~'g~ll:;ntr~ly~i;.IpI~(n~ :rnd 1t~~.up5t11iOO:gt'l~~~tiwb' (Fig; M). &:~w"'lI<illy dup![Cl!!!Odgt,nos ~1;l/hil/i(Nld ~~:trod


~i~,'!-' ~Oo\v:idc

dlnl~~licl!'IS (5. J'5'I;~:~







29,39 S 1~~Ll:20.'5.] ,6'1],478 1[3.5: 211.4.] .

5..762: [9'.6:: It3JI 2:2:.395 [13:7 .7JI

A~l .11.

1~3,S7.2: 1[2.8: 27.7]

3'9.536 [6.S:iJlJI

21,8.3:3 [0.1: 24.}61' :ll1..fi52 [(L4: a..8] ] '7 /],16 [0.3: 1:112] S7.~,,6l [02: 13;11'

1.4-'188 tl,9'8iJ. .20,,101 <'l.6,,57'S,

[4.8: 12.1J


[Ui: '6.sJI [3.7:1'.8JI

h,uli«·maf!li\-'t-ly J;""'li,V~I'llj.W"dR"ti! cii~Il~~ \...~~h ii:'i C(li!iiiNlailwith t~mi~\.M. Sl,i.*5'C>ll!'ig~¥ dl(j(iem\.'J>~S'IliI~ ,flVCi' king evotwiJ1iOiJ~m)' ~mVl!c ~i<m,"""Sme ilii~'fSnroll~ptiiiiryiltt ~'1cctitliilI.l3, 11/1. ~~
Ant'il~sis of [i~lcl[ivi:d~a~ ~rIC· II~miJIi~ pro\lfiledi 'lL(Mi~io:mtl ·ulilsighIS. f.am:iiliC'!i im:olvoo ill, ba sic lil&{)~o;iad ~lI~Sl'lCS (slldil ~ ril~J1~1 It1.u1C~ml). as wlii~1:01." r.1I'111Wiesmn!vcd ~n U,t!1"~ripli0I1'11 I~~ i !:.!lm~iol1 1''::li.I!I;h as MYBtl!ld h:!!.'lit' hcli'x,",oolfl"ileliJ!>. ,OlI·IUI.) IHl!k'>t'riptiuil fOOoJ:l'I.l1<1mM 1~~mi'IIC1}'j~w

(fiJ ~11IIlS1!d! I~reli!led



·~1I;,Sl35 7.021

!JIl.~'~r-(;m;~~ d!:Ll:!§~ tF.[g..3A~


{li l'lII.IIc·I'cOlklQc!h~ldi.l'l'.il: h;;l.!CillC-"lich repcm ~NS.

III eo~Hr<~~. Irl~y

N;)I (r.qI(~~!:I



~iII NilitdJ~ 1I~.;m Qr~I~!TIiO~L

'1~.7 ~90~ il'~.

M'f6 tr.i!U0I!~!Q!) !It!t~r~a! I!IfILH tli!~ll!tplJOl! !"lJrnr GIj!cOO~III'I!iI9Ii1Sl1' ~EO!I 11M


!LII~R";'!l'!le:::. (7~ rn~ '15%Qr~"(:~l)t~tikc! kj~I"1M.'l ! (RlKl' g,",l~ hmxbor Ul k","L,<;t(ln~!1 ...~nr~lfer.1 ch,m~1" The m!~y IUl1oriO!l :L'S~gl~d Ib 1'i:!.Ilil!n'b!.'fl> of mite NH ..LRR g.;".ll(: fillr!.i~y is il'l :oml1l,in.:f"X;-cii1e fi.'SlSnmlr£ Ifi p::lhhQg_tlrlS (Ull, fInd t:L'C'Cptrn'l> Of~li.s class 'C3l'! be 'C;".C£OOiiilgly ';l:lrtialil!c,. !IS presence IDld ~~ce po:l.y~noIl11hi.wSfiJlC OOll1Bil10111 lin A. lfwlitUlfj ruJd o(~lcr w!lml:5 U ;-if)~Our d3UI. i~ldicrl[il!' ~lis dlffil CJ!!lCJl~IO tll!~rmj!!'lly oFN B.-lRJ:t gcl~ in~h~ A. fllidimw ~lmm:, A I!hm~l ~lCY hml'c diverse
ID l.iJI~iIHl! SNP .0 !'RI!' .. IcalJ!lfHia:1ld: S.H" SlOOII"RP

·!IGI1&9 crua~:&I' Ae:riili.~"l:iI~Ii~!!«l

fiu~l'C~io.rt!i, (.w~ R..LK!J)l"c$ luve

J~SO ~.



!3~,*,!IIl"I)Ittroll3Se 'C~omg


::S' ..~

Ql~.ed iJ~ ~.,a.spccir.c p::J.tJ'OJ:,~l. d!:tCtl:il: (11). ntIS, ~1:'C li.nd1t~g nJll.ll RLK ~!:Iles Ul'C·m"!!:l'1'q'l~g1!l\!dIil.li' !1J:.ljor-.cITOi::l ci!;)ni!~ ro~dlC possibility 111m~li~


ii, :si.Jl,1~1"T' III NB·UtR

gl;,'II!~, .~ COI'f:il'q~¥":c


11>400 F!!IIG

~r.!dt>orllr.\\ ~'iID1'1".~rI ~t!~(l1.Iil;~1l d~fuill:4; :tIn:I,

~\,,'t]~ ,t2.:? j.

RQI:~~"':~~.!I!i!I~g .1!'.1»;t






All ~!*I,






I' I



~r:r.rll"il)!!::;1(94I,(1<!!;(l I..i:IrriliHII~t 0!hIIr ~O) S!I!ii>s

IJ NQi~g [] ~~(IjI~i


l'il!lfiiII.!Q!Jr;a S[;ep!II[l'19l



We fmu~d mf~(Jr;,en~CI. ChITIII.~""''' I~i,j·IiK'Ut· in 1Xn> of .he: F-ho(:(. SlupcrJhmily,w!akh cOlnprise."l u~lliiC [1miii 6f~tI ~lll!S ill A.. #wiitlJla {21) (.Fig. 3A). g Tlhlis,ri~ld ing. in cOllflbimuiOln with ol.her do\1,tii (H 24). s'ho,:VSlilm F-box genes 113\'(;undergone !mIlia. bir1!!l Ha ,d.c,~~]~ the ..1 llmlimm ~'cenOlne. ill !\Ithough F..:IJo.,,,, enes hove boenl Ilm,posC(l m g CVO:INC quickly hl respense 10 palimgen Illi"ii:S:ii!llU'C 1~2~4). x ~)erintcll,ndstq}P'Oft for nil is bYlrol:he.sis is C' la"king.l'1~e poly InOJpllllslJlS we dcsl;:li.be·I~IUVidc ;)! ~OII!'CC tb:ra~qlbig]igllki~iC<l1 rob: 1.0lUClllkilCfS S'ign;aJhi~ of :~e~tnQn by .s'NIF.lYt~ and gene Iy.• To ;~?"\\ tl!1~ i,,:d,"iI)~.to \vI:li.ch I!I~ 'l"rl,ili(IU '!.'.'C Qool.T\ll~h;}8bt:.jCl] sli<.!jl'OO by ~\::lt'i('lioJl. wt: (",1\;filnlimxl.~II(\Ii\ I~"qucr)(:y ~li~ribLlIJ:t1t1." fordi IT(1(.":IiI1: classes Olr 'po.~f11ouphj~llr1L.'l ..'(~OOSi»tCllt widl g<:ocrnil, CNpectmion"" ~ilr )1.ck:\;th'c OOfl5lrni1llS nlll cooklg :>eqLlCIltes., dim' .is ;iii skew t~'rut! ~o~.".."rlXll]n~C)' ,,"ullifllttc:iat 1I000lsynmJ,ytriUll..:; re~fli.i,!!.: m t.)IIDltym~li)U:; ::ii.it":> (6) (Fig~ 3B~ This sk~'i~ mo:il.nlOtlblc lor SNPs lhal, inl~' 1);fC1J1Ij1!U"C' SIllf)oo(tGI1,,'> ,trLti less ~lmnc' ror al.h::rmge-clTecI. SN Ps. '[1'1(; tt:m1em:y orSN Ps c~l1ig ~luamlJlm;: slo,ps tobe r.rrt':Sllgg~:5t'): ~l<1t .,l'I; k'iiSl. unclei.' 1li'!~Uli!1l !!cUli!'lh~ Ihcs'e dianf,'CS ~ Gih;~1~S5OCiHtl."d wi!!hnb'le'lS .m.'Il:s. A~~hi[)!l!.Il~hlIh.lk froq,\M;_,!lC'Y dislJihi!l!lin.L1~ ~crG::'s; ,


(Jf~t~ ];ugl.\ )'<,;1 !l";I,t \'\;,iIJ..c~~I!:a.;!~.i~I;;,i~ g~!C Ihm~y.






fiig.:lJ;. l[)tbi~liI~iM !)If majo~·en~ct ,c:!ilo;lIl!g:~ ,aJ!i'lda [lete hNllJlel:!.d~s.,(A) Fr adiiOdl of gel1Es aflfe~ed by ~a~g~n'@ct5NIPS er PIPs !by 'g,CJi@c,U'e~(liiY. HLilrg;~@nocl5iNP (!ndMlfI'~ i(]t)ladid ~not.@~a g@n.Il' h~fbofi!li~ bo'tA 'IP~ I(J~ polli'IlIl:G.rphi$m, either ~vi~hifl ruhe~m;~ .i!c~s.'!]i~l!ner 'hill d~H~reli!t .a;(jc:e.~i®:tl5, !:oli!e~ \'Ier·e G (hat ,~nti~n~'l1i!'asikedl J~.p~~tive'. gnd fl)'F wi1j;d'l Il~ SNPs; (~ijW bill p~dkt~, "!fer,~ ~ilitJdoo ~~m .~illa~~s. i es RING, ~all~f II'nOO~lin~1 New uel1lll·. "B~ M~~or (lUet~ irequem::y' by 'SNIP tYpe and gene r~mJ]ty. Crilly p'(j~illon'S Wilh ooill1ll'lel\e daiial rQr at. l~st t6· ~ff t~e' :00 ,aJoc~~i~I'IiS~~e ·as~$SOO, Ihe I:!Uililiibet ~ir po'lymorilIDPc III!@S~~O!'liS ~~d\!id~t!inld11l @~llfSl!I ~iS. s~m.n In th~i!l~t r'O!F ~ r9H~feat; SNPs. "Otller" im::h!ides liU)lIIfun;t:tiQlIIlIl~p6n! site tlililn9:~' ·5HooLih.lltiitm.llin]lli~Jatii().n melhiolll~m~lim;l~,lIlildl ~bm~LJIii:l'ns '[ha~ rl@m~i;!' termnn·atilli1i !(\ocibm. Eirr®:rba~5 ~@rm,te''!l50tg (ol1fi(j'i!nce ilri~va~ Il:lr ~in;ilmia[ e~L:atf:Dn,s.

[,II'1j!l;~i~ <ff]!C b:wJ.;rcily ~hni~, N B-tRR .~c:~¢.ll ('xe£'r1i:[Jl] fR.g.. 38. 3lwliig, S~2). Merr.

'bo.hnOJli>'}'flQrI},lllQ"LI8 !IUi~ SlIiOl!gly f!\;"!.1~~ Wi~l

and synonymm;!~



s!kt;WOO, ~Q\~m h~g_llii rrc¢ll~!~'!1'cyCt1m~ t!m~ g~!l~!f!~ i!V{;T.!_g1:;: (Fig, 3B), 'n~i1'i

wiilh high 11C\~n,.; f po~J',ll~llli5lrgl {\:..g•• ~l\\'CCl] 21 o mw :25 Mb QH dl@l~lOOQ!1'1e ~·I,
h ~~d!nk~III.!.p ~~i;';l!~mlil~c ii!.1~MJ .. 'ilOT!,~ !hr~h~

,~f ~O]!!lf,! tYl~' 91'" ~~:I;.!.I1.ein~; r<JM!;.'f:IJI<>- (Ji'\I\t;Jl U1lilth~ nil\; ~\'iG('I~~ fOIi lxlih ... kdiml (p.!rI!!.:iI[l!;;~I'I ~hc:ri;Ilrm Orl8;,~~01Tl:li~rro(bp- s,~n[I:Il)11IH)[I!i, 1iI1d inj~t:~tilic ~IY1rn:lrllf!hu::\ifl:l. di· ¢ r1l;;".Clt li;,"(t~(I(ll'On .ile p{dYIll1~·flhic sites seems UH1" ~ ~i!tklln)', mid "lgroe.'l ,"...illt C<il'~ic:L' \ii!{1lrk., 0:1:1 dle basis
.8J On~7ilq

nlml :Ii~O'~I~d b.;1~O~ ~·~csri.lyGdOOlC;"d ~I~ .:;;~I¥'iflg otgm:li:;']lI1S ,p.~,IH'ulih[~~ l hn.", heen rcporic(1 ~illt ..I ~~iuli~DW 2'~•. 3(1), Ci~l~ (12. l\ghl~Y ltrdiitil

li'kelly C!o;ll_ikm, B<!l,~rdg1S ~cti(lll ernl h~ Ql;.11~lhnc;;;tbrR,'g~!5 linkOOtp foC~LI\~'y ~~l,ti!ml~!l~~o!y~mnph~~~t'l(27). " l~h~!wm\ll1'Dn p


1~~i1!iiy~'K;.liiiOO!";>.. !;Iug~~ed!:hi~ made !J)O!!l


i!]:thr:U,JlgJ'i ~ciQ:;t:!:(l!!i IlVl



of ~"ctiQJ~lti, I;M; ~10:~ !linli;:On1iinlDll Ibt. NB·LR.R ...t- Ilm:li(Ul(J (J 7. 19, 22). Ai'iii:cl,ditioi'llfil ~rctlk:I:i:Ol' ofb.!'l,kmC'i~!g )i,C~'CI:i;O:llahig!J~ei".[h~I~ iii .• :!:v"1i<ill~ ievd!lrpo.ir_ymoIJhl:m1 because oflilc: ~n~im~l1mt'CC :rclatiivdy alteic:m, hi,gMy d~\I'Cig,OO of ~,lIck."'S"Com:,i~cfll 'willl tl~is cxreClmlon.mmrc dum ,SO% of N,i3:-lRR ,~IliC5 are all:~',!]jS;1 tl~illi'lij:ln:r ind;l~j in PRPs U'lg. 3/\ I. UlW1r ofwhti;h OOI'I'CilfNJIoo ~o~li.gidydli!'ir:.imillr ~~ 1!I!~11C'~. A.hhou,gh l''"'~!t i ~"'~OC'j:'!l~.~ :;;i!1'!li~'~f[~IIW'~o £,,"quency .... ~!1'd k'1..'lw .iligl1 nl!arrnbL.!l'01' PRPs w~~ 1;!1~'i:.lrv!Jd fur RUK. gQlll>.l,tFi~;;:A ,>lIld r~. 12). A~lholJghf~bo.x S gC'-nc.s~ma:bol' the $i.)cond·hi,r,d!c}l! eccurrcnce tlf .rI.l'ljor·C~r~l (:'Im~~~. Ollw~:~ ~qu.(lnC'y d~mri· hl!llio'J!;'; O)II'(! siltiii~a~r 110 lh~ ;J.\' ~rog,\.l (lFi~. 38 I. Gelil;®me·wi~l:l;plClt:~eWIll!" ,®if ll;flllYlinol'phriu1lII. Tm:tti:ilg m l'irool!ter pilk'li~ ofvari~Lli:un. fnuuKi n we m.laU&;~cll~II'ID:llrull .. rl(mil distribtuil).il flfp6.iiy.UiI.l~himi le\·c.ls 3£it'OS.'>lhc· £C110t:l'tC' (PUg. 4), R~ioilS of high pobmxl'lrhr~'I1 '~I.!::ald Im11l!hc' 'CCI~I[mn~ to

cilhtln'U1C!'f;:';l!:\Cat d~(!n.'tL~~ytu'i~l~i'lnl

t Iij.M 1,I1.11Iio]1



mue:;; iifl~L)' a I~o d~~fcr ~~WC~i1. dllm':irimOiTI1~J regiOi'b<;; 1.l!.."C~hDS~ r d~ 0 ~;li::mtc(,s~l~ base ~Ol'~porl· ~itton. Fnli'll:illy.n11~i"C31:\C' Ulrn1i}i;t eci1aiiil~Y 'bjrus,..":. in the 3'mly.bill$OO I'CSJ~1iI011c::illig,-~ .g., due to I)cgimKl:11 di rrerenLle5: ill roJX"tIlt co:ntt:n~ ( 7). Vtc believe nhtll~1iI~ dwe {:"~l'wmtLiO!lS OOJ'IlI~l1tl'le to·llI~e olcooU"i. ii."drmu..1TIR. In 11 l'I1ul1:ifile ~~ml, the

b1Il;lr.!ci~ ~o\""1il:irl. S'udL11i'> l3,.l~R ~'S (_2_i){F~g. l\ fi'iil,Y ~vt;ri~ ~l:i.~{l;;l'jE of high 4 poIYiii.101ph~lti $htllila:r 1.0 wha:t lifl!~1b('"CI~,obll~~t\,tldi



f~f .~lC' ~t'iIlI::




~c:> ~3'J. J11,lb~' lOi1IhOOill~1~....J,~7tj~i ~'tlj}jb'ilC ~eI-..e l13) \\liIII W! nl't>'tlUlf~~lal~iIlrulo1lyt:fIlg ~1~e



allow ,dh'!:~cj]cc 1(1 00

bcymlt:l :ti!c ~:nc:ri:Centl'mnedc regio:ns. Sim:ilnrly,. dll~ of NU~L RR :BetICS (l, 15) ~ <I'... ;lJ!i;~111td < Fl;g.4.

c~y ~.dlilliOi'dSl.::ca1cs.. Uti. 'Wi~1 ~lilaI fo], IttcIminillgd);.': reJative' im" be c~il1lm1('tI 1'iiO];)'n~(kqlhi"n1 1c'l,;'Clli-; ~l'!.;: !'ig;n~ntil~~~~l:1J'I!l!1!.u!!Oc !lrSCI~Cli'on 'I(''l~l~ mUUil~!l";;!li:! v'l,irilq~iO:I1. In ,rontr.:ltil !~i~l.s 'ofl'm:~ pObf.~!lmflltils!1Jq. n~glll correklll,'dwiijli\("II~ ..<\l 'I!.lIilblll!l. im:lu:JJiu!J;: lli;;·IX'<.r ~ d~'O.~i!y. :[iC rol'l~nl.!F.ltn!tliJ~ tl~' nli;;.:.;;iin~ d<,w~ili .... t('lk-e!n,:~!l~l;i~ve ~l't~,(Jiu!' or "~I~ci,,"(.~""'(:(i1.~'" . I(M,. .U I dmmclu;.'1'-1i:ro~1 'l\lyext\)l1!\ivc' haplol:ypc ~CI..:~roln dlcCClll~m:In;lre, anddm"i~¥ oj' NI~Ut R :;ll<!.ling. A $11!d)1 wilh 3D I 0 aw.:. ~~HId ~~~ g.... (tlb!le S12~ lh~" IJia~K"!l1f; 'H'O. 1!000\IC\Cr, nO! '!l~ l.'o,..i(ic!1t'C liiir '~\'\IO :;.l;;~!"lb,:p~1 ~\1\1"'W i~\,'tIih,'iqg :;'!!Il~ply w~ rutii'ac!t of lih(,l ~,q,IIJIl:."lld!l!l h:.;clmobmr. ~i,;lCIi\'Ol~iouf FlU. ~LIt!<Hjordi::~f.lluill::am. orl;k~,\ l'ri~1g o !.liqL,')' OlI.~O :)It;' I,wilbll ~11re.%-qlill()i~~r!lg ~a1a otl:l:iI1~'d nIJn.; m] alll~lJl~u;l, Ilt)fJll1ft.:auiO!lis ~l.Ilul/illlm {36l~11lc of \\Iith OOlcr !oolmicnli('S (6. 161 t~g.s 13~ r'O:r Oc rugi:ms illtllJuocIN:B·Um:. .gtlIl1: c~ IlO~)'molillh~.n w~",~~~ro even \"".h~l UrICNB =. Lru1:.gJ.'IJCS ~Gl1isd\ccs \~'\'tte CM!lllicd trig. S,M~ ruld 11O~ymQl'a~b~,tn Wfl'i al.'iO dC\'<iloo i~l fmC'lJ,:,'I.'lLc DNA, 1\.T\'lt~h.'I(.' m,. ~1llt!ilm£ sill'ZliOIl ~:fng.:(}n l.h~· ~~;a'l ru,~ted ~~yll:c It..""ldr.<;, in Fig:. JB <!Ix! o*r ,~IQ~~~ {n),is 1I.

LlClWQ.,'lil 1I:l~IWO

or n!lld~ll1de di~ AlI~gg~ ~~~~.


,~ Sy.nctll.:l'IlKIUS -.llnt~nk

GC~o!lle!'l!t R:e'PeliUiiIoe, j:irQbes

~~;a~~ ..R,~.~.~.,R.~.~ •• •• .~.~.R R R.~.~,.R.~,~.R •• R,~,~.R.,~.~.~.,R.~.~.R,.R.~.~,.R.~.~,R •• R.~,~ •• R,~

plrlttedfor bot. rourfima de~ll!eT1!w ~QlflvmClll~

~UId.'Hltid~d~\I'@r:sil}' is

!a.ooli~1iIk ~fles along eseh d'lwlOOool'lle ,!litlil ~Udin.9 \lrij'tdrfi.~ ~f 250 I<!b (ctllJfllm from ail ~ffijj) wilh all of~eI: ~f 100 kb (7,1.. GC oonlmil In ,eaxlll
wiru.ilow wa!> Gliklil'lrlt.ed r',l)!'Ils'!tes 'CiI~J iillihe (lO'l.c' sampt:e' andl has bem [~J3iL't>d ~ 3~% a,tlhe boH.{uTI, of e.aclil, ~; ,andl47.5%i~ ilt. rIfi1etop. liw broad pe.W [If ~~~ itWe I)f-o'be ~ity @Iii ~(h



00 tfule, 'O~tromeriL



~(aled~ ~

fit~peal content

n<l'i bes1









is atlh~

'!O.P 01 oodi plot an~ 'OW:O ~.5 a:t Ih~ Mtt(lfl'l ~ of p!J!~rptltm for butill fourfoM d!![l,llneraffi i!ln~

Q,(!OO ",-,_ ~--.---""'oLlL!!U<,-.!.....J"""'_----,,!!""'!.~..L-"-.------'

inlerogertk>ite~ are ~mlili·
,c~lnd.y f!eg'ati;v.~~ oonrl"" ~ta:lmlti!l!llle mffl:.:llil~ 10

figl. 5. R!~illn~ or ~x~Ensiill\E1ifJairm~ 1l8:p~GJtYiPe ~mtll'ili!~a~o:ng '~hlnOmOSi(lllf'!e; . AI1:a:~ioI'lIP~k$ are 1 !S(I:Fit:ed a.I(Jn~ '(h~ 1 o!!);-i~, H!Or1!zl)!'U:a.l, ~@dI lil1@s dem:ar'Cat.e Irfmlpa ri5:O!1~ lIr5nng'(me'a()C:l'.5~i!lI"l,Md:l1
lPoosiblepi:l'lf1i'lfi~€' B~a:c:k tiflf'~irldi((l



.i;lii!d ~~

lMly wfrel~ ~~Ih~ IIIl!Irriilier ,cd NIM:RR gOO!sin IiIOn~rl!appifl9 .SC-FWMmllS bi:ile S'!l!2). Poo,jlliHlfJlbiSmis ~MUl' :S¥l!lOi'!,,n~ sill's:" \l!'hic~ is ~rlty dlj@tol to!..'!~r ~IY ~f SNP;iir! inm-g,etmfc ~


(e.~" rig~W.

at ili1ltelrgenk

be~1I'ee!i1l 1 pa'ir of a~!te~@Iil~~The ~egflOfl ~l'7i>ee ~ .1' 20 ~nc! 21.M!b,~hib1,ts ~~tens-~w , h~p\Qtfl)~ sh~ ~i'l~l
'DVer In~;111r~Y500 kbil1la~l bl!lt iwoac:ces,'Si'olms, C'OIi-O anlli LQv-5" ~~111(1iI shown at.lhe 'b~t'I:oril1i. aire

,c()mPrari~~!'!is ~e mQ~ons (!If

Sh~Wfll 'Cl[f!'lV 011100.

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ene di n1en::;;ioilil than, [he (jil]~er; a ,cl,b· Cl"ClUilblCY too Iilrg,~ to be explal eed by \'!Clll r-d\'ilJI\'ely r.m:r,c hy m~iliibcr in~}le lim.:), W~y Zdpc~ll~cOl.y. Expiamlliom; FOI" I,hu$ h~dLldc G<l13X)'. thc)' havea di~!~'IOI)(!!r;I,i('l!rmtee[Tcc~ o.~~ shiO~lg ,d:iI~TcrerlliaI rouILtkm (IF I,h~ star lZ'fn or gotlm:~ic: cwlmiou due to 11u:;~ilr high it~milto)1~ the 111)i;!S~nOI! of ,I I~O~I' \'>'ind 3). ddlCI' of whoch it ic.'l.~h(:lr i'il\m!n,s;wi nd~"m,nd thei f nmd end m~, h'Ne 1'<u:~rom;hf!l~ C(l!1."iCq'Ul:'~ICC'l' Ihr our 'lmOIlJ'" :-;nmdillg ~f $I!tTh.r ~vol~l!io!~,']':o nd&~ .~~~ '~UPIi.~L1l0\"'''~ (for d~JC I11D:;.l nl'l~si" ~ml1'). Re' (It'!l !b',. 1t;;I;p,idl'tmili(!~1 in .~i nglll ~t;JrsI\UI~ b(,~.!1~ i!i:."I!I~, w~ !I111!l~~ .mlC}ve bc.,},Qn!d: ~1L;" ~i ~.lptL;1M ~ invok,I.."!l to ,~...~)~.ain",l k':J!S'! one :Iilmjol'type of mgdc~,~ ir<lflid nmuing ,~taff;,'. al1~llli! ls will for ~y W!!Jnmu'IIT~Y bursts (9') .md binary C()!U~.'HXln{:l\1 l.iI.lqu1:r-c.~ ,corresponding jW!.m1'l ]n 'Ii~t': ql.mli~y of ¥!1"'~~,U\'~:<1<U'l' iOrremll~t.~ ~I fJl. OI!ld qUOI.ntity of .in~errCBDr~ll11~ry d'U~Ol,I!ld!/,;x,dI. Th~diS:liru::li\'~ob~"n;llion,~1 $i,gn[u~IlIrn:l~ of Ol)~ 11)n:ivi'Q!!!$~!'i.~IS w'"~ 1Ja,'\("d ,on li~lil~d i~1" rup3id rul:;lltij~i!ilt J~[l [i""l tl~stifi1rloo b~ VU~1 t~rrcrornl~h:1[ busdiilLb':}l:lilf;"u[ h:ll~l<~d Itih(l' C.:IIPfI" M .Zetpell 01). !]cgillning, ,~'il.!itl~e ,(!.\pccl,miuni Im'i~r

~U~JroD9~~ofl1"i[l(hlro~I iflglllncn;. i~1i;.: stc~:I!In:IIbld.l1dmtres, I(~}. A~~lOU!lJ~ ho~,l>L:m,~ lire



d]O;s,t! of Ihc~r )jl~wlyrol,~~in.!!l

sUB" Aehenl!li:"


\1i1i~~ ~neEbiu'Cd


b(' ~%%

li~lu rroh~' rOli;t·IC'I~oW!'C!; of~hc C! 1.A.lMint~r,lila\lC ~l'n~h~sizoo an c~· li,~li~f11 aperture with dim.cl~s,iol1is. 26511' b:y t9S'n~ (f.lg. 11.~U('lwi~ us to roCOIl~II'UCI. im~i:ro:InClri.c riflly,w~ a
OlI~C::1 l,h!l:l~ro1OtYI)k<l1 rnpg,dI r<Halo:J' A~I,f1i:1i (:'Jl""oc~14!!1 '~YnliCJ..\ TV') w~lh an <1H~'l.IIIill' re,~oh.illion. or -,0. M nm~. the: diti'ra,CltOn Iumi! defined by '),JID. ~[lf:ob~llJrv.i!.]g '\,\t;~"!.i\ti:lII!.;'P1glh ,divided by 1\'l';C~ 1~1i;Imlg~~"! illl("ll''f(lrol;''1i(\t~r l)"'$c~i!Jw+ The


n::t\~lll~yoon1irni~k,.ned ~v!idh j~:!!I~ ~n{'I'hlt!dComVII o(.~wmij:!L1Ibirth is work. odIO\'\'llm [Il~ li~l[ Hum lh~ CIIARA wl~p;:.'$ In< 00 ol:llmbin(;d :sunqUII:(9ill\;'Oiliis,~y inlo l~ight :;;~"'<;" II'.1I.ichanuels sp"mu:ing l~liJ fUi>lroW)mic~i H iJfhllld!
brl1l..~ fM IRe)


I:i.mt ,ooluifu~91

:folJCCS; ould d~Slo.l1lhe piLmo. w

sph,eric shape ilmllhm the resa.l~!ing o:b~;mcne.,:>s woo.ll[] ~nducc lower c1:Tccli vc~eml)Cr.lllmCS m. ~11cIXlll<l!OJ:. Tl~ is Inlier c:1:roc~.kno,wu as Si<lv.i.1y

fig .•.1 fourier ...

(til, v) ~ov:erage: fer IhIN~~l:alr ollSAilrva:tiolflt 'I'lh!.lfi~ ~;:.!c:1Ii POlnl~Ire pr~!!rI~ the Ilwjedm ~~flllrtlti'm bebliee n one pilIir

(l:ilu:kC'~lin,g" wi II Gll.!:.'>;(ldi~0!1 iD'll!1,in (he (l,hscrt,~oo li~l~ ,~mn~,'!l.01)1, Wli;U a.~Ihe O\'()~'~U .~pccH""lencu;gy dislribl!lthm. PIlCC,isc Ilroo.lctuons can b<c made, but these 1,'Ii!ly Ol~ l!JlCcl.'n."Iin~. a!\:mLllpi~iol1s, inpm'toc:ul:ilf lllC d[s\ribulion of ,1.'!:lJI,g:l!!~ill' !lThOlll.Cn~
~llImin Ihe s~ UI~irill']f1 ..r: :roWlioll, i!'i On1.;'fI 01\,.. '~1lII1:I~dfor 'SiiI11p~idty. 1111(; m~l. basic pn;:ducliml::li of \"0111 Z.e.ilpd n ~ha()ry -e(ml.ri~.·Wlgtll di5mr~k"1J mldgru.\'~ly d,llLrkenlill'ig -liIfJ\IC been em! !lnnedi ~o someexIcm. .I"a~'mllllr Testbed ~m(;,f;;!rollt(,'.tcr t~'l])I.


100 I


0 ·1M "1








! !

'~f1er'OlJir(HAil'll!; '~Le~pes; S~ E2, Wl~ lIndW.2 Olllhe dadJm 'e\l~i~e! s&nowslihe eqUJwalen:l cnv·l'ra'!,I1! lv:r a III eltlplkal aper" w~~f 2615 UIi!. bV 195 !l1IJ ,ori~.nl!ed ai{)Jlg gl pos:lti'ooa~gl@: orr 135" @~[ of 1fI~:rlh.






"," -,






... ""




mCflSlL11C111~QI.osj)Ju::ih; 800 2.00,

dm1iY-lI.ion Ell a rnp[d rolluor.~buru.i~l~e diall1.Cter orlhc 1'IC<I_I:by A~I.ypC' Sl~lr A ~tair to be - ~4% larger in oae dh:r.le~ls:im~t&mn the o~l~eL (11). The NaVji PI.'()Wiy~ o.~Hic.,"L1 Ifl.lA':ri);romcler (N[!O:~) andmhc CC1l1~Cl' :rar~:igh Anlg~:lar Resd~LlljQn As~rn~~onIY l ItCl~ V\R/\" ~mel'lcmmetdc almy liol,h, mc@iSull1.xi


CI I~a$l{m~

~1j()O ·2iJ)iQ.800



:)'!roIJ~llm'b~:ken1ng protil~ rOJ .he plrolmlll::I.~ rIc ~"!lIlKi,lrd Ve.g!:l,t I J. J til,. oon.'lj[Slx.'J11 \,vi.h a mpk! .mIU~(:t< \'i~'\,\"(,1~~ !~e"LL~}' PQ:~Co(l~'!' A bli~htu~ ~:;'yl]nlil~11)' I~r AlltOlir W,Il'i ;~I~o rq,110~Ji.i~by )\lPOI
lDeiDar:li~el\l 'of AE~foMm~. UnwEilsU)i ,of 1!\lcilrlg!l~ I!iI'In Aroor. IllJ 4U009. USA. ~Schoolllf fhrsks ~nd ~Jf'OOOIlilY, 1Jl1ifIM1i$i~ 01' '5'1. I\milil~ IFUii K¥169'Aoh Sootianc, UK. J.A~t~ph,y;I('~ G1fiOlI;p, COVEnd1s1i11labora:tol)l. (ambridge 1L!lmnllf!B,iiy, (~lIlbrid:9~ (iS3 OHA, UK~ ~IDMsion Gi' ~. lQQitil~ ant:! Pl1met.l1)' 5(ijeaJ,~ Glmn~a l.milUill ~f
VechliQ!log(ll'. PiIJ~di::tllal, CA 9:n,2!S. USA. ~JIIiI10i!c!!iIl1 IDepilrilmlffilM. (5aliliEll !:.irn...!1!ml,)', UlihiilC.J, Nf!j"n:48~{I, USA. ~U:borMOiPl' liI'Aitllo~:Iii.~illliJe d'~,Gooolile. Ob:~t'Oije die

~ ~








G~~!bW. ~·3~Ci~1 ~n@~e

,e~719. 'uSA.

~i~nffi (l!fI~r; C ~fomi:!l lrn~il!llt~O!IT~i'lI1IIl~ogy. IfI~oo· ... liIal CA 'f..I'11tl.'S, U5A. 'Cmterf!Jr Hliglilli ArlgL!l~1Ir~~1uliDn Asli'OflOifl)l, GE'i!I~lfa, ~Eale UjjJ",er~I\y, 1i~l<!fiM. ,Gil 3030'2, LJ:5JI.. ~IN<Iiti:omal i]pti!til~ il5'IIOOOIl'lY Obsefll<ltGl),:. lflJ{'Som. tJZ


Fr~Ii1ICIil. 7M1~he~!'I'

" of:j·' 1E::;a~t (fIi!ill[~l~oon~)







E1:~s~ ,(mlill~.ar~~onds)

'"To whDr:n oo~II'~~pOflden~e mO~ i'iiei'@limleh~cu

.~IIi ilddir'E~. lilt


IFhll •. 2. lA.) 111111~~iil:S~lY of 'th~ ~lll1lia(e ~I A.ltaJ~r {i,. •." :1..6,5 ~HY1~ illn\ag~ Cf1~~led: wi~h tlii,~ M#!iJCIWMlM imaglirlg me-tifla.d lII~~g a UlIfIU~m1r1brigMrile-s:5 eli]pl,ll(01t prior {X~"" O,9'8'}'irVlllac~t phooornetri' eWJlr~ ']n th.e i m..a'gll UlHl1!sporndl1l ",,4(t/Jl)ill intelilsity, (En ~}I1'~@mtfllldedilmage (olnvv:t\feci ,Vi~i:J,<I Gillufifiiian !;Ieli!rm~r 01,64 ml~~, ,~©ne:sp~l'!dl!ligjbtl th (Hr'hl:t~~!l U!lfl]~r CHA~_A~!!JiF(h~s~~b:lef'lhn~OR,~.f'~f b~~h: o ~..a Dlells, th,)2. s:fledril::~ ntlHl~itiec5 ilIt. 1.6S pm 'VI! re ,tj~lH~vi!'rl'edi ~nto 'l!he ~!om!!iPQlildi Dlgi Ill;"!(; body ~ r~~t11ipeirilitl1Lr,~~; Jl:O'.nltlilllli!i ('Clir7000, JSOOI,<lllId 00000 K::ue Si'hmllnl. NIQlrl~i~ 9p ami !~a~lis Idt


1(:\ ~

'1,$0 to 1,'74 JiAm~"The Ah~~11diii~ ]j1lC:W!ll!OO 3! i\l\!,g~~1 ,md I S~p~~noof .2!rJOf:IllIf): I;:O!1l;I~!~W O~l~l;Jll~~!~"'! i!lfom1;;Uiiol~ i.. ~'I~ilriM~ tJl)', ~~l !tg,ditim~,. ,\!If,~ i!I~ld :;UlrIlL;'K.b,u~~cl = :t2 ~[m) o'lll;;~f\l3[ierl~ by (;. ~lD~ P'f~[o ool'!S~rni!'l!.h~8hmt~bl,""clintl visibi!i~k,~ I i:ra ~u~~lq~u:mt,hmd}'~i"" t!~:lil
'iNC!l;: O(l~~'OC~oo m~

\~ ell IICs(J!lved U;'ig:. .21\). OI;p;[)c;)ring chJI'lg.<B.ted ln thcnQr!hc~!-"g(!~lwc~t '~~N(:!~O" .>!! br.~§ht w~~h domimllu :rlr.!Hl!re ol,Clv¢~i!!;g~h~ nonh\~sl q~ll;td·
~,~11!I of

~~l'VC}r, \1!,1h!:1l It".1ng

the grrnvil}I"(IiI.rkening


",f]!l;I.1£l.1l :l~tl;fG

1,;11I.V!.;:k"p~-;;;. 9~tt


!ll:!lf~ To~d~I'i.Xl

~hl; ,~I~n~lI,;-ru;:i,;: of

p,ri~~oC"~IP~~tl'l ~i!1r~itb1g r~rd n.r \I~t:.~'\;'~1s',I!!~d,~ru lh~ ~lpmuR; ."yn,~~i... IJU(:Ie( ..;iI,J!('!S,lhi!l ~~ri~~io~ bi",,,,,,~our inmg~."g <l~Uijll~t fai~.1I. ~ni!lf\km f('"I!I!II~ Ol!Ji~in~ oll!!l.'i,idclh!.l !dqo.o~)h~~ iUlWC\I~r. \,'lfll; c..kl rliDl ~,'i;a]{,"C~ .:m~~cin.:mllt'd!<Ht ~I!lli .. ..... i~ A!!<lI1f. ~on
which i~ rdOl.l:ivdy i(X..1O~ la;(;kiulg $i~I!.~ of W"IS ci:uiSl>~'rn'ilor SIIlU!!!l W~!td$" FlJrt~aC'tdCI:liliO Ol!llr i:i1'I\:l;gitig ]p1lOC"Cdures,:rulong wihb resulil" from vclidol.kli'lllCS'l.'>. ean be foom:1 in (21). OMI' inn~C' shows lhc' :i>1.clilarpholOsp,hcre .AJtffilir 10 be

\Villi llill; WI~~01' rNu~ CH.ARA l:ck;;;oopt;,1'. il1l.tl'r~mim'i(.'tric ii11\li~iiig ofAII:tiiur is lioW jK!5i1,[bk, :!,h!liOl.u~h Lhh. il!.'qlilire..~I>~ci[llit-cd in'lllH@C' iieecun· l>truel len ~","Clm~qll~~". I.t~~d I!a,;: publiely 'We g;v9:iliable appJlical,ion MAnM '~M,m,1.·k.o\l"(,h:l.!i.iu il1lflj!'Cf Inr' OIJtka~ :r.I'I!erfCn::nnctry) P'~) Ihis :111 wo!t.!!iPI,ljiil1gd1e mrnxinnom, ,enlllCJ!W method a",~ ,~.Ul'. "lYe reslricted tile ~Id 11!!1f EM) ll1l<1gc to f;d~ w~!hi!l"'~l Cllllip1i(,d IJil;llUIJdO'1!)'..~imi!~lfin

ro& ..ib~c lbw-kvd ~l!iirlicts.i2!mtaLTC lJ~yolfld Il~i: d inhle.ja!l ]:imil of out fn~crfi;,'$om~1.cr, ~"'I; ave h rd~~~w~'>cl, Ml!lnd;;rnl~mCl.-durt': (251 of OQJ1!hi;;" 'Voh· illgnli~~nmE n;ur.:::tL'cl iiiiiagc w,ifh O!l G~IWI~~~nl jjcmlib iltiM~~~ l.lte~:soIILlU(1I10 r nhe uUiIli:rl'(omirn'iilig c~cr (,Fig. 2:11.).

{f;IM~~ !J t~all,1(b(;'.;;l-fit ~~mll~i~n; P~lI,;'fl;o~] ~'! (15) i[]:iJ1 tia.;~ ..~i~ \'~~ilJlc ~~.:l.i:fi. (II. of

to ~

= 0.25



\V~n wit~

{~riV(,)1;11 :p.~a:,lmi,;:~I,1(~


of l ,79, whiichs,.,rgg~Ib'" ~. !lL;l;:d rnr :mdiHo:n~dclc!:¥,Ct" 'of lR,"i:k!l~ in 01,1, mode].. '[:0 ilj~P.II(M~ our :Iiill.~. 'Ivc e.q'lIQil~"dan C'll;:tCll.... iC!ili ~o nl1c \ICiill Zci~l~nooc!, ,~H[l~\'ingllll~ gmv:ill.y~Iu~"d


bC~1-lh model reached o:nly "

These i1II'k'\5'-!-"S cm~lllfilll tl~!!b.",.ie pkJrLit'l: of . gi!'"dV,~ly d:<lrb:n~n1g inlclil1tloo hy IDpiid rol!leion. Wi:: :iCC A~Jta:llr's "hoto.... hcre (0 be obll!lte wilill 3. p
b3'i£~n re~ICi:n idclw~fiab.l~ as Ihe );teUar p()~<lr lJ~gion.l1~c inlCn:::.it}1 olr the dark !>'Iq,ll!;MO:I~OlI bmt(J i~,hmn 60 ~o 10% or!ilc bri,;hl!l~'i. at the p.ole, n b:ro..d~y ttfin,~~fi;li:m 'wi[h i"!)';4'li1;.;'(;I;uio!.1@for !lrl~ ["I~,lr~il1lhuR;id 1!'~np.~v.i(]l!11~ Itlt~d!>. .Aldli()U~l W,I_! see !i(ln1C cvicl'cll.cc· tor (h.:v,~mion:i" 1~C}m a..... i" Sr~1ull~t~i 1:'I!fm!! exeess crni!1!ii'o:r:l pnd1(:' nolii'll" Cf!fIUn'lib~.'lhi.'J Ii;)'muro I:> f~!n11~ hll!.~l n I~ouri~mijl(l lid~~i~y ~ud \\I ill .ri(lqll!iro~ jd,iliQI].-~lful,.II~i~r ... OO\~a:~I~cItJlimoi\!..~ligm.(! 1!I.J:ru1cr~\M.:: h~v~ also lined oill!' dam ~('I. winllr 3. rall!id. rrm:iltol'"nnQ(k:JI, iOJI~fm"~ll!;lthe Ilrcscri~JI.ilm :;,e~ QUI in Aullh::l1oo~ I!I Itl. (J 41 ITmd rcferenccslhercin. assuming 3, Rm::;hc' pDlc.nl~a~ '~cel1tmlpoinll. m~~sl'ml(! solidbody rotmion. The m~i:n pammclcr.;; or the model wr:e Ihe ~~lJ:::lbw rmj~lUsand 1iCm~lC1<II!.!I\C at [he" l'i(!!I~.~lre <J:lIl,gu!!l!'m!,<J;~iol1 mt'c 0lf1U fm(::!.ion 011' ~l'CaklLP r( rot IJiIC gmvil~j darl\.ciIlil'lg ~()C~lkicnt (~). and the \~iIClving !.lIlSIC's !incliml1iO:I~ and I]Gl'OilioJlmlg~(;".). '!,Vc lIS00 dlC SlICI!ITI" u~m.o."phcre m{ldc!~ of KUlrtl¢z (.26) Ml dCll.1111linu~ d1C:~)~ciJ!ie illi[(.IJ]S:;~y nf ~~d! flOilll 'n.Il~h~ :iil.llfi~c:e'il! IIfi.nncHOir! ,n;f locf~lgrn\liny. 'Q:m;iCliv(l IClll~pcnJlll:r('!, find ~hillh dflri!:.OJlir:ag.hll fldH~tiOIi 10 1i1'1.i[i~C'hblgIhe n.(;\\,' h~URD'CHA.RA linln"wl: .loiICCdbhc mode:! 1.0 I1lil1Jtch IhcinCtIL'ilUl'Cd V~ !lind I ~.·b.allcl phm:on:ui!~ric l1'1![lgniu.lcts (0.7M :±. OJ)15::md 0.135 .l: '~lWl3. re.'i"pOOtivd.yJ! dcri'I!L"Xi frotH til b"J\Qna IitmtMrc

,~ ~o be 3 r:l!i:e~a...meier, "\~tbmtdnlml, ~ Imlorlel with ~ ... 0.190 Sbg;J:lifkJmiy iml~m\;'oo lit!:: ~'()oolla-:o~' j~~(Trible .~). ru~d I hh, in1Iu'O\ eUlCI11, .f;,.; \'L)!lmll)' aplp<llt'illt wll,;:n, ool'l~pm; ng syn~'h~~k mllc!el il11:;lg~.lo ~~CA Ilau, iI1!11~g,~iom CI-~A RA ~F ig. :n I n ;JdcM~:tonm .. lO"if~' 11. ~!e ne;w ~rni(h:1 prefer.; ;J s~ughlijy I~ti iu~lin~d nriC1]Ii:iL'ti'tm. iI em~l~rpO:~f~rC~'ip<:lro'U~', [
ilUH.l a fas'l.Cirlu.mktll

r"';''l'u~t~ oint p

BOlh om
,1:1< iql'!J){lf!.lIl!

~n1"-lgil.lg ,1IiPlt] !lloocling


cl!l.:rrki(.'11~~~ in!:h~~ ,(:urtlll!ltly It~OPU!n:< ~1~1}; to~~ulJlkl rnl;'1![)Pi. Pro'll'i[1lUswo:~",n; I~'il~ also l'lrq~~.u~h.~rudpruMcn:hs cXp!:lUl1blg hugh ...
:inl.(::rR.';rofill~l~t [jOU:3. ''!.'.!i:lh :Slal;utu:u



for l:ol.lnin.Kl SiMS,. In add~ian


nlb'l@ l.B~sHit~aJ~am@'OOfli f~r R~(ih~1il Zeip-el liIlIode~~ of .PiH:.il'ir. IPara IiIlIflier descriptiiol'li5:

(0" f:; (XIle"On, 90'" is



I!l OjiliQJ:n~Il~Le (d~g ~es.e-..a!5tc~ IliIlr~h) dE!:Sl:riw Qt!f viewil1g! alng~l\1', :r~!:Ih! illild R1PIllie. ,de:5~ribe the I()ne (;ilIln ,!llIl'S{rib~ tl:Let~mperah!l,B@ .and ra,dnii at. the letJuatnr as. t!q and R~i1I)''Id i:o. thll'ilIn:gllll[arr 'r~t:I tiQn r ",U@I'lS i! 'rnc:tisll or ,clil'liiC:i:I~ ~nlli!k!llp tate.~liId f}~; ~he gral!O!i~.y-drk~!!'!lIin"Q ~oefficie',nt MOdels a~lIImed .. :S:!eUi1!lr Iii'IQ,S~ '" ].'i'9J1i Me {lS'}, m;etalli(iilY 1[lF1e1l11) '"

'I.CmiJ~:ritureo a!l1di



,pew. (aU:cmatlveily,

J.\chcnmr case 1P~'vrolllsly d~t1 . .P1.":~ct'SQnt!t at ,{.lJ) were lHl3blelO find a sllI,tislnCIOI)' til. ~or: .I\lulitr assmning it ~"tmJ[JliIl'd Roche \lOll .zci(l'!'!l nlC}(ro~ (X~ HIt co.n:-;~~lJ.Cm wi~h [ire need ro:, ~dil.,iDml~ Mel~m' pl1ysi($, L~cc~Il[re.~ult)i fw J.\ld~Jm'IIiil1lUQI''-llso s;pec:ifi.c';:llIy r:l\... m:odels or wIth srn~llc'r f}l;:. in .. ~inc\'llilb Olll!' fin.dil'lg.s. J\l1JJ~flll!gi! ruoclc.l l1:u;il'lglm'i re\iC'dled l.hhnoons
$A<)lmm'(] 11100:1")'. OLI.f il1!b,lgiH~ ••;Mow\; !lew l!t'UIUIWC"; '~("

1'l":lIxk!~~lldc;l;x,':..n1dcUl I'Ioe ,d ~[i;Go'\101"()d



lliIo(tel p.tmd~,glnl:li.

-(11,2' (32)" i!~dI '~iiSt-!'!r!II~

= ~LU


S'I.rik.ir~g difl'cr;e.met.~bcl.Wt~n our Ci lARA ~mn:ge mo.d Ihe sjlml'tctk" r.nndd ~1'I1f1.~s (fi!il:. J) i. ihfll ()tf, hnage :shrrws stro];lg.c:r &d.H!iD~n£ :m.!mlg d'ic oq 1II310r., illi)o:nsls~enl~dth alliY v(m Zdpel i~kc !rnvi ly-<lmtCl"I il1~Pl~ript ocm, aS~!Lnnil1lg~iI1lil,brm

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2$. ],.A. OOgbl!rrn.. A!i'lwll. M~ :15 ~~W~,~ ill? {U\l4:~, ~6, It t:lUlru~ Al'M59Stg,\~t Ail~f ~9tWm~ ,(It:id l k~ ,G:tid I$nnilMMian ji,.sIloo~Ii.~ic~IQD~MI~, camllffd'gl\,. 1lIII., l003l~ 21. S,li!d~., K. B, M<H;G.rert):lfJ, A. S~!I1~nid!ll"AsIfiQf}hyt. t: ~ppJ. S~r. l1Sfi,. ~.<l.~tmdk5l, 2S. 4. i(.l!.ar~1. ,i!s[fOI:l" Al;rJ~p"'JI,3~9, 289 (20010}. ~9, F. lI;~pD(ll~ I~r.!!, m RL~~Ili. ..iiUp1lan:W:.~b;lailto. IllVPiQZ2 !lo5 '[~001). ~O, C. 1!!J\!!!i;ill' .•. ft, ,~. I)f~~!ff.,C,. n, 5ih0fl •. AID'~J1.1, Svp}'l, Si'!:; ,64i3, <!!:60h'lOGM. ~1. ]'. A.ttll BWI1lIllIlt:u!' I!'~ !l'l~A!lI'l'J!hft,}. r628" 015.3: (a005). 3~- 0, ~!l!'e~'. IPO. h(!!1i1'.~ro", .M~.fl, ,;I98,.1l~]
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'The 'CrystaUizB,tio,n ,AUle of Eucrite Z,ircon
E!!I!uiu~s aI"!!!a grQUIP of l1lleteon~'!> tha.t re'p:l'le~l1!it the fir:'iirtt plalile<tclll'yigneolll~aC~l\lHyrl(lllJ)~\liIil!ll met ..H~i~iC<ltedli1Wie'.r,eniliilltioli!.Olil an r~arty 'p1l.aIn.e:li"'Slrn[l~fiil'lllii~r to A5te"oij,~ ,~ Vesti!l, ilIlrld, 1illJls, help Id.i3Jtegeo"ph~B:k ..~ procl!)se's occmrillll'9 01'1such bodi,e-silrl nu~ early ~oli3irr!>yst.em. U5ilrl9 UlI,e sllol'l:·livled radirOl'lm::!li'de :11l12:H~as a pelatillle (hmnomet.er, \Ye ~'E!.IITiIDm~rate tbt el!li(ri~e z:imHil 'l]l:Sto!l~i:i!iEl'd 'q!l!~d:l.y' 'lIIIfrl.lhillll 6.S rrniUi~11i'~e:ilrli ,of ml?l<lIl'i5i'~{at!l! dlirferemtii!tiolli. llhii~i~ml'llirl'5e thllt. 11Ti11I!rlUe difhmHlIl1>t!ltiM ,(lrl~l1@ @l!I.n:riOO IP~r:e'!ilt: bol!lly ilJC(Urrod dl!llrU[I'Q tl jle<riod Wh!l!ll1iil1it@rIli!ll[ h@iJIt !rrrOm~'he

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~o~e.~hC'decay ...r Hl2~:.1 1 S2W h;u>been used fta !o dc!!erlni~e time sC,oJI~'s l1ilet,ll~smc;!le ,dH':' of ~erenl~1Illion~c..w ii,£: 1:0 "(WCrormalion 01' I~~.,n~ t"ls (5 I iiJld 11llml~c d ir'&".!Ji:ntialicUl roSlI!.ilirIg in t"f1J!I)~~On,of b~,dl(ol ol:iEfU· 14. 5l 1~lo\'!III;,!(I,)!!,. Hr ..W ~C!I1iQpC!!iil.i{]iir~ of J.]l!iUt ~i~.. t~ It]im:;r.d, ic of ~hurrL·~i\>tl[I~Lu:li(l. :lind It:icfi:;lll ~CPiirn~(';S lin, {_~LlcJ'i~t~ IrellrL.:ci I'i'ici,::t·

WI'l\:iil P:1:filllclsCOl]~II~.!lb,g bulk ::;Orlilf pfOlmr~iQJ1sQr 'CIC.lillC;ll!S:lfC Illc~tC(J ami dnll'cfClllimcd, UlhoWh ~1l:!111' runJ :sid\::rophi fe'll\! .<lre: chem~c;:d I), ~rnc~ iomned ,md r~islrib:lIIl.1il~ imo t~e ftilii;;i/i.tc mmlllllko:' alld l11Dc!alllicco!:'e, resllCclively {ill), Th,Cl\..>;.


r.lJtilJ Qf-t'7 10< Itli/i. In 1\·:8~I:;g~, i!he highest exA"~~ 138~\vinh lU'i.','~1lI]1IfIii"~V w.'I:ues ~l(1!'\"Cn(lr~ 1 ;t9,5~ '.:: I 0'1,?~ {2Qltl)1 is 11~~\V ·~f!O]Pflic cQmpm;itin;n, If one "~1.!n1;l..,_'<; Q.."&" '0. &:1:\11;/,P;d1.C ('J<f ~ ;'.Q l <I. SO 11 1;i~i!'''!1;l~~~!13\'IIi~l ~. II, fH/ 1>6\V .. ti9 I,lf ~2.2 Z 1C ~»li~1 ~'!r on~1~!lr.nl.i(}n -~% fDr ?~n;:\~~.~iu; c of l O~... nd dID lowi,.'l'>f Q I,S:2\?ll'!~lIc.: of 3,4 (;I:~.~) :.: ~tijal!9lJ:;,l@ W ~bl)lllcljnt~c; ill ~"'n:\.~iritll zi~oirn ~:> ~11!Jl:bri:~~.la!:y ~!&n!"~IlII~~r}'~A:i1Illizrui~mr~ ~hml]!l!0ID' l 0:10{21Ji~) is ollJ...o~n~1;I i~:I22 ...., lit. I oolHrJ1 ®~W . abo'~i 760 !11!pb te 9J! P:P'~~l. \1I'h~~K""ISI~.l(;;h;m.iuic of '~~I)Iil:rih~~. 1"'&lio r -9.5 :.;1Q~• lhc- hl~h I',~Om r :2%) and W'::!)I' ktw\~,f .ci!1:ll!n hills "bmll I 00 '~o 76(1 r~,pb En 1\... .. ~8.1<!(W, ~'X~~!';!5g 5:l\:\f \'1d~l~of 9J (-2} )( The (lh:;l~f'\'ecl ¢l~'f;L'i ij:llt~W I:> l;:orJil:lil~i..'d c'Qnl;;tn~j;.tion 'l'IJl'I,!n ~, subpprru) ir!;>.:iroo!l mllkl;: i~ [h~ highl.~1 lingi!'~}I\",ithuhJ(l I.UAN' 'II,~~II!)IIL:,';S, ill zi~ojj ~Dtt! is (JO'm IPtlI~Uid~~!~ k:v(;U i& o~'c~i ij~ l2 all id~~~ nlii~.\i'OII'~'f dc'1.~"tnliniIlMD.ht' 8l.I~~.r0ilIb!,l~.~iQR~~~_, 1 J.HHr,iiJ;~k~1(1'ube in. ~i~Li !te'3y ofl8:!1 U' i!! die hel';l c d~~tire_ Tltc rr~rcl lorpril'l~m)' i,!jTiOOllli cl)'.~I!I~H~ \\'iDh jj cuM."POilt!ij.ll!!l~8"Jlill~W ,,,<I[io (I r -1.19 IO~, ,mId the Itl~vll.';SD 3~~\V V<I~llC of IJ l±o.6) ... jlIHiS~ The ill,,:Ii;i:reti ~~,W· abmidimre [ulMe S3 :Lal~fi~1 ~h(ltiJd ab:o be b~~t~i'jJrcsc~d ~niiro(i:n. IOl·'&.f2:o!l'l~ is o1:bei"\,ed~n.z.l \\'.~llh a i:;;~~.wI~W H) ,mtti Fi~. 2] ~l the i.irll'l.c of '~ii)' ~hln izat~,M of whIch isii'C£;i~tmu t'On::Sicltili1!!\lo:li lJ·[lb isotopic systemru.iC'$ UllW It~~mm!l)J:j}l~ic 1li:.'11,PCII:':.Ilure:; or <l ~(' The i,on mier..aflrabc :~ ideal!l,)! ~~uitoo lilI!!" fl1i ~i[LJ 1lI1kIXlal1~llr~bl r sma~.~l.'YCv.crnl 0 IFig. :l!.. 8a<w.attl1!mti e!edron ~BS6 1 (h1, (If micr-omClL'ts orless) ,~h,Ol,;'>Cl' 1lL'1;.';:I."'Cmin ]l'l1ia~ 0:1' a lY!Pi~l ,gfilli1l!1lilk regiilln 'inl Ar H<~C~ abul1!clRnOO, Using lhi::-: mClnod {U). \\,~ :6S1467h,p} and eli!~~~d BS!E: ~~ of .ililll1lhe!r lypketl !ilr,~al in A~B!n467 d~~mlijJ!~d the ur~w oomJul~i~ionorind~\'idu~d (b(Jm!1f!)., All' R:i,il1!erall$ Im~til'l triPle ~II,U~rilt;: !,jlfC(I~I'. il~ pDI.~]l(.ld.lhiu ~~~i'lI~~ ~I] mcll:;r

II"IlQij~~licI:ff'lt~i~i,bro.li.gn 0 r W 'bct"'''CC!1 l!hes,Q pJ~,,~,~at 4:541 ± 2. M}, (9},~~ [~lr·Wm~nt;'wl i~~~!rp~.~~. ~l~!Nqrpw !)(W~~:'l~!)q.] or ~~~rmini~J]l: f






W 'OOI1."lUi~1he chmllOlO,g), of igm~oa!!' er.y~ru~~ . I iY~<l!!ion of'l;.ll.!crill"l' ,md .iH1Jl."!.C I- indlt~>d IPa;rti,~1

'Jh\'!! d]l1l~b;,i~:ltic (iut~'il,iI;.';S, 'c~msen lor l.!hi~ "'~l!Idy :Ire ~he A:'~lJk;l m~toori~~ A-:>I8146'7 :.uoci A.-83 t3~R ami EI.qphom. rvl.orainc nillcloor.ilb: EJET90Ct10 rron~ Illc AI][ru;,C'!ic Il~eloo.l'i~eco.llcc!ion. T~1i: ,,ic\stlkiil 11'lt('lcmiite.~ nrc mlbreCCa1lLlctl mld ~<lve' 11 rocrystal~ izcdlrock.lC'xu:.I!~C til. ./3' )1, P lil£ioolmSiC cly~tl1h, ate l'I1Io.'>~ly al1hoo:rn~ (s~i~c: ab(D~l .05 mm In 11 mml iFig. ~.}"wi~b a H:w 0 1:lllill·~IOlIJ,lI\.>dIcry~t..'lL 'i !!lliOl!I!ll nlll"ll )0: 0.15 limn)" ..

Me .s1i:e. (frlOm c~tJiJiLrt ,O/OSmm up to lJl mri. as \¥i'.ltl as mJ~n~ Igr:i~'S ~,P t(J' 0.4, mllJil" is p~Stliit MilliU~ra;1s Ii~d fifWlfl

]l!oclio!1ls. ,A_n_h@~r1llplagio~~ 'Cd' vaoo,.

da.~ll:ll lignter gJ<lyAoffiite' SSE re'1PolilWo SiOol, SiOrpha:re; Pla{J) IP[agk:idpre; (Pr~ u-ri;rfui p~em~; Lpx" Ca-peor l1}'loxene; Urn,. i~me:nit.~~ !11 mild Ou, Or'Qmlitet ~B(Jtt-©!i'!i!)Zi~l)m! i~ka~ I~~~) bV a~ M~h <! brlli!llhl SS!E. ~@5jllDnS@ 11l'DssOO~tiiDJI a ~1li~h an 'ifrnri~ Wgh~~.~~ IElSEl'end dnomile {d,jif~r IgreWB:S!CJ 9R!in,.lIlt
]lIIdMdually pnoloo(KlImemitedl rmie~onle :rirql!il5;a~' laihleled,. '~.g,. A~s:m.467-Z3~ rorid8lnlil[;clIIiClml, tIIl1d '!liIe isoro~k data U.allle :S;2),ror each g~ilm are rOll!pmtedi as;

OOFJi!~CjIOO 1cr!!N'i' iltl'Q.F 1e1!~~t;: fior terrr~tria~ a nd w[tu~~eorit:emiDl-' el' al Iph,a,'Se-s,1I'hll!dg51l~~ tin~ (~n tlil@ mflifl figl!r@' fln~iI1Si@1:) ru~n~" TIL!;; (_~liIllHJ:l~dI 1lI1e:Ili!nlIo:rphue OOIli,per,MUII'C Il!~~~ semts '~heEWR mrrr'!.ltiiltimn ~illl!!&fi1JiJ 1 O.n~ I.h,cCa lh(;montc~er :us -;~OO~C:' 161'. ( ]mt~~l ~!I'I2lHfll:BoHir = 7'.2:5, (:i;OS) )( lhc I ~:~.:w cO~lpo:si~ii)an of W-rm:!al; N t;n~ I0' :il~' (20m). as I'llpolfl.~ by G~ Tille ~, .~;.r<ti:i" ~ilic<1i,te glass st&flmrru; tcrrcstt1lal zirecns (91.500" llorizontal5~l[jdl ~ne rlejl1~enttl'lgJ i! • Me.f,~ '~7.stl-:n. and ,G.,z:il'tflU); t\'-M tMllK 1\-.8~,~,467" .U7~{I llionn~l WiS<lWiPic OOm,pDSiltiiClIll m'lh and E£r!IJOO20 iil<XJl~~ imd y.vool'i.C'mu,:: ~;j)~IT:" D 9"500 3:!i11i~ = (I (J.n 'i!; r.hO'l\l1ll 'in the rnaliin • G·ti!'tlO" POrl!.litl irn~ilble S2 UI). ami Fig, . .2. ,,-\.111ana~y~-.d !ij'~ure (,111'11(1 'ins!(l!t l~e~erre$~ria~ o 9,"/:SIM't ~crrc."'1 riol:l, zirt'Ol1 !!:l,m ill::;:()i,~el: f~ \1i.'idc r.mg.c or zl~f1:; {G=li~coi!1.9:1.5(110, an;d 9j~ o N!sro:r() ~:stlI.H'r/lS6w m1L'OS Ili~~o 2.4 I.:ms~ have no.tp::: SUS']-2) Ime!iloumal \latUE!">, 11"11<11 IS1lWtbS6W andilS3WtJ~V iSo.'lGpic !::mnpo- ~toorioorlf((JI'l;S in A{!8,!l4:67, cA..~iIiQn.The al'mi~A:mk3 ml:.1eOO Ii:z:i~oon ~mil1.<;; 8:81388, an~ lEEi1i90020 \~th hJ;jh i!~~'IC.... '~ cr:ss· ;SiX2W and n{lll'lllil1~,~ll3W U i" 1/'1)1, lMHfJjft'W have mmdatm eoooosf> EIT900'300!iJt;{l!ll t~ri!h l1i~h jlllOlUla:~6\,V ~ .~ x ole;O!W.,lNh~(~ fi~,1I!tlfibl:lrt~dl ~d)' t!ll~' 1n~~~OWiii \Ie')' Tow ~X~'l'i~ ~ ~flllW e'PJtjJ,,:i~un)d\\lifli~ deGilyof 'I1iH!sh'!lrt,~lWed rildillnLidid~ 'J.®2Ii',H.'Iih~ diltll ,i1I~e' regre5.S!E!dl u~nl1i~ 't5!llIP~O,t to dele"m'lin!! th~ (218) ~1iI'iliilILle2Hf alt the tinle' ,!),{ ~Q.rmatilll1! of tlile~'iirl:oliiS: (~atl:l~eB u.:n). A;l;li:K:azirnon. P,yro~eili(';S loolili A-iil~14(:i7 :nilt!

pig~rm~ ~LIJ'C l'llme conllno!]' (fii~_ I ,. bonor~ll', M Inl1:l' '1fJ'~~!!8(~\, ih1lC'Jl~1.'C d1:ltnni~~, <Iud OOCWI r t1~ 2010,1.0 50-~lvn gl1Jairrb~,wh~liC~s the trace phfl~ zircon, :l;lldfr:dc:;.. end silicfI are s;unoMer" .l.ilttml gmm I~S ore Iypk'lll.lmy a~OCi!llL-d witlil ~hne:iilhe oalld clt"",m~[e, arc lri3l1lguki:r inl sklill'{,". ,fum:! rcsemb'k b<mkcn f18glm:ms, A few z.iirccmgntD1S fIIiJpear W bord~l:iI'rncll.il.~h:roJnlil.e IHim~cr;a1 !ISSCl1l'1bill£(:: however. unHke hi(gh-grade m,emm:or~ phh~ me!!;=- U·n,l]iICji dOnll:lI,~13.v!C II 11111tbasein di:l:'CoCt 'cm~t."l('ltw~th ilmenite. E.ET9U02u (po~iS~ICd~hrrll seetion no., 2~j has g,mflulil~C~C%UmC' and ,;)j In~ner IU]lonnt (I r cC!<l!l"Sc"grnined lire;] I .l5l', The gmm!~ulic ponlon is comll~d of;;!llhedml P')I!Il,'\,\l!\'C H2SU ~un)r:.,~ckJ\Wd bylalidike tmd ~~~hhyphl,giod~~;o;~ {-60' J.U:ll.). Mh~l;lr oxides, trudyrni'~.Luldl:rnilil(.'! • .>1bt_1tK,~~U' i.u d!.i!ii:;;~~ti:OIl,

.P.~goo!nli~cand augil!! gi.<lills l13) to 11.4 mll;l arc PlrcsCIlt. L"i:soh..njIGll bmclll,ac wcu:e rOlmd in b[lt1l~ pyro"",eJlI~. l!~!lKlIJgh ;!t~gile ]:mld lae in hOM

A·'881467~23 ,alnd ~~i~ (ll1abi~S:~!il the same as ,A-86U,67~.3 repeat.

IFi,!!!.,2. lhe U2\IW!!~ wllll~s are , IpIJ"~~ !1I,ga~n$~JJlliOlfnjl'1!B~vahll~'




A~~ ~D8.'-: Z~liOOil1:i1i,~ I I1 :1t.(1U '7, S (~O. 9) .. I!K;!I r10.-$ 112cr,~; mcO!:ll s~ ~wrc 'I.\illli.gJn\>d deviaies '~MS'i,VD) = L~~ n~~dI,rot A-:·ml·467 ¥i!.li;:oIJ~~. il'


:S1l1~:~iIIW~'!f= 'l,f, 1,4)..: w;",,S {2cr,iltl; MS,,\.n = 6,0 L8~ TI1II: nt~)dC:llillpr;:t 1 imi~ ~32:H r~,h~l'Ifor dOlUlCC dlC time of crys!1!1I.ir,ation OIl IEEl90020 2:i~) i:;; D:8llL :18Q:Hr:::; "} '1(10-6• lfhC~!1i,i1d Irl~\1I.f/II$.;j\Jlt) Wil'l!~ deWII':~nilitd fmiTn ;d~ U1L;Cw!-


aminimmn gf 12.0 My ;:mc~lh;jJ.t QrA~f.:~"l146'7 ~y a mi'llmwm of o..S My. He Mn."",r i1~. d~lJ;;m1i.,]\,'l~, rm~n who]c ro~k. PY1\OII'\·~~':. "'ld itn~i,mii~ IIqill~m~ ~~'P!lrnt!J;S ~fk~,tn;~IIJ!li!l~ of it'io[opic ~.tgl]~iJ~fi,;~. byahc m~I~:r.ll)[)vph~c C\r~'jjt ... ~h,11 p!'t"ldIl!C';1d ~hi<lgr.mul.~liiC loC', ~\lI.1JTC ord~~~e ....

Ihm. was dirc'C11~ ducwiIl1J~~t 0,1' W bur,,,1 under hot I1mu(i,;'n",! CIiIC~\,;:!!,tod ~, imp>!I;;,t is CQll;)!dI;,'11Qd f~ l~~ -'])J' mQ....' r!mls;!l11~m~h:;)lni:s.m roOr ~~I:¢ril~
~ll~~ill1illOflll~i~"I~l. !;i!e,fer,liliFllOei .. 1!I.lfid N1c.tei. It, !')i'Ii~~~ 5;, XY, Sd~w,~ :lI~Q,~:;I6 (1'm)'


Ih an lL'li.tn.'ol1icll:ylow

Cfi[¢ 1l'i):MU'C~..,;,i]lW.m'C'elo~c m the i~~[lil~ v~~lu~ diCil~'i~iii'Cd nem eaeritc w~olc.;rock lEWR) :;iU'i1I~

~Ie.~,~5,. li;Vc !lIs~d .Uii2.1 U' a':l!1 relative ,dhmllO:lib1C:1Cf O~l Ihe,lSsumlllion dmt it was 'LiUHQrI~1~Y dilSlrl.llli.noo in litcresCI,'oilr rro,Dll 'wllkh comd~ucn!:: of ~ri~nili\'c mClco:ritc <lnd E~B 'W~\1l: dcr.u\\·cd. The

!oi 1.0-6.3Ij,d it~ !nooell :1 If·'l,V age is <4SB.3My .. The ~Dmlil!l· ti;ul' t~ifn'ic'of M83 1. 26, iIIl:y inl:em.."'tI lilIon, 1.~fiSIW. 1l1ht11i1.dance ,~.l5~,and dtelow I ~21 m r~blii ndan« of EE1'900!2U sIlISg;csl. :droon c~!7s~llmza~i():i1 3. ill

~~\I\!"~X:C(;;iilli~~i.~ !~nr/!~~lnfor~7

,('1,U1~11l~~~~lfl~~(SSt ~I01I_f :$)IRklUU ~ni~iall .as d~W'~minud fm!l1~ri~li~ivli;~l(,1~Co~ilCi5 is LUi ,~ to,Un x 10~ 09~,111.C,~1b!.lJ1d'I:lJINl or 1t.1:2UfiJ~_ fcrrod R-.olll EWR ~m]lpll"!' is In;;::!1 l~~1 ~IIIDii.'R.= m
'J"25 (Til. 50) x

uldlki£ ,C!'i,/cmwi~c.n ~~Ai mId '"'"'Fe had complclciy d~"C'qccli.uK! ceased to 00 hCtilI. SOOIt\.'S .. Zimon in EET9002U ]~.nmre ~i~dj' a l]rodllC~ of l!o~!e Ili"!1i ..~ nl~Rin~ (lIn EP]B: U5) ..~ <lft.~l!I.h <In


~IJilpa,cir·~~.l:t~dlh~.u~.l1g C\."C')It.

m-!!' (5).

Ti!~ !im~ 'tirr~L'itnoo

IhO'l1!~l!io!l .wd n~o!::,d· (m l!Itt:l EllU .~~.l!i:. LI.3 IlO~\I'$ Of fllml.hn\1 ivurusinu:ic lltf.l:y 'iJ.'{~riL)rl~d M.~'. arid ihe model Ur~\V ~1;g,C lOr Lh:ii> evcm is: p:ro.lul1gt~d l'lblIUlli'LD:l.phis:rn and ilt'!JIXICI. \\!bk:h clismrbc!d [heir iini n.ctalogy. IC,(lu:rrc·.iS;ouoW'ic 4!S63.4 ±I jM.y ~5t nc dHJcre"cc illl:li;211f abundance OOl\\'ren cJQm.pos.ut~QIiD. flmllmdi;01U~lric ::'Ige:;, l'h,cr:unm.l AS~lkn zireeus IIIbdth!c EWlR {51 $Illgg~ts tlrlflt.• me1.;m~o:rphbil]J ml!ly I'CSlJII. froft~ oorilill under re 1m.!ve to!ir!C! hue of l1rl(ltllJ-jiil:iealc di IJcI"Cl'I- )i;UO::CSS'~\1C lava [lows or ilmliXlC~.Om::'possibillil}' is 1:1131 rco;:~.atcd ;lIld Inu:m~:InytnC'L:il111.{lI1l~imscd h !ijHim~ (wible 84) on EP:B. A-$l·~,~3:i'i1'i CL'}'Sr cw:::ri!Cf. ~oumcd at~h~ C'()!lillm:'~ :rone... 1l(!l\1i.1i.'(!n ~aUizw 0.4:t. 3.5 ~lly ,mdJj'\.-Il·~1467 2.4 :t 4.4 M)'. B~i;:!nIIS!(l oncrites fOI'!'l1Icd>t." " n .. '15l!1!l,1of crdlror \:I.'U~I!'; iJmu hot imp,!!;;! mC~l ~1~1!;.'t~< J 1), A~tel']laH\idy •.euerites CI.'~iS18~]i:l'lOO mld W'CfC e.'1irly .D'lulnll.c dincrenlil:uiOl~ fo]iO\\".ing nU':I;m,I-,si~i.Ci!llC SCp~JII<l~ iOIl>lfChu.i·\j~ ages of zi roOIJs prcdalnlllg u'!.eilJllaOll'h('!~cd during iQ.lilnill:im~ of en!S!. on tile ~JcuJJ"'Si hc~tc di 1TC'11cmi.n IOJ! 0011 lEP:B ·~rc phys~ Il'!JlI."Jll oocly (1:.)1', 24). If cueraes lhm'il'd by P<!lilimJ il;: "lly ~H1~l.O!m.~brc" TJ~i.~'C:f(JfC. ~h¢13_'2~1r 3lliul]· 11'!1e']l~!l:f!,':. :lne!~!>\,'llOy,ld l'I1Ii'Jill!"u.~to Illc ~lIrltlCC ortlilc j):II'I;'MI body 1111(1 ~!!l pilfldlUce 11 W;1S:lit i(;C'n.I~~ Ihml d'1.11t;~ i1110ClHCl' [lil<l.I. A-8f: 13;88 zi:ocmll (blll1il,~"d i

Ba.~lItic ~'UCtilC~ l.b:m,C(j a,. ,I fC5.1J~t OfSOHliC of . ~he IhM pla!l!e'!)."!J)' I'i(:~"l!c ign~'Ou." !l!'¢I;CCS)1c:<,. Th(;:ir ~¢",rulJ~ ~l:!:!il,~.!.,;&I~l!<I!~rfi~~'Y I,')rWf!t::tfii ~f l;lv;1 ;:ire



1811,. S8~ (~.ooro. it 11. l'\Iil\1m~> II'(; .Mm~!!r.C. Mlin~@1, IiI. falm~.!. A. fI~dlof'f;, ~im. AC]I:! '3, ::1.935 ~lOO4·). 6, K illii~~~. 1\..'I'illl1~w!il1n, H. ~m;fl" GeWlim. '~~~~hJ'm. AI;~ ,~'il, ~~~7'(200~J 7. It W. ('~IIMI" G. W. lluglt1\!~r, im ONfJrfl oil! ~1IJ'r.JfJd M'WII. I(. lIIi!iln!f;F .. n~ It; G!!~~iil', [iIJ~ ~lhililt. ~1l!iii~(Jma ~. f~c.. Pil. 2L10!!O, :l.~!l4. ;on, .P~. !3. H. 11a:1Iri~, w. jgm~', Ilr«.. '.l!'lWf s« tM/.1'2. 9~~ ll9.!!:!l '9" 11. KI!im.eo, II( Mt2T,l~r, HI.. l'a1meo, 'E, Sl:l1I!rs. C I!.I.Wt~. iE(I'n,/Il'lr;m;N; St.;. {<"It: ~::t, ~l WMDS}. \1!lJl l.tt w. ~. I,. Wil6~.1!l.,1 'Eti.is, N~re 3~~~ 11. I'!MtM~&!; ~n!! ii1I~11'Kld!; :I\':a'i~mr~,~~~~~I11ng ~ri!! ma~Ila'l ~ifI Sdrncr'Dr<lll~e. U. H.'<'!koed!!i.If. !i!ffi, T. Ar~~M. ~!l1~.M(fjr{.nN:

2. G, S(Il'IiWl~II'I,. J. N, ~ml,. N. 1Il~.!m!!liJl~StiM~ 284, ll3!48 (19'\1'9). 3. G. w91Pa,ir. A. '5~y~~~, ,GiLlJ:lMim. (OS!I1'lodiim. ACr~ 6Z, :O!l;!63: (l95\~h oi. G. rQliilt~l. I.B5rcl" tL.J!. ,!l,II~f.~, .~'flfll f'l~fJ~, $cf. Lm.




l!5" nom),

M'~Jt'lUm- KJ!'S',]l!1J\~0'1 nnil~, 13. A. 'folmagllrllL 6. 'ilaIf1Qf. t:.. Kf!IL M~/(I'roIk MNoe,:Ifil:i!'

1,7" ijn~olal1d ~!I'w .a~~de'mn~d IJ.)'I~!!: et:Il!Jatii)1l$ 1('I&!lW/t*'W'"",pJ(~JWf'U~II_dl-i~ )l ],000 ,:wi [fSJllI!"I~"iii~.!(IlII~~!!~~'-11 'lie S!i.. md~nI .... . h;i~ lIIZWtlB~ = !l.9·lo:!l3:e:;! "'!lid, u'YlIj~ = 'O<.SOO3~ Mrl! l'KlIiltiJ.I::IlerllrrM1'

flt., 1:~ ~l!!!. U9971 14. It B5nQM.. Ill...~~III1.oi!~l1, M. V;hll.tI:II~iJM:, J. ~tm;.. ·II,Z, 3i5!5(;l:~Oll 15. ~ YOImagJ,!!!.hi et u.l;, ~i!J (!]i-''f, ~l\im. Casm«,iiiJ'!!. .4d'l:l ,~!li, :~n (<loon 3 16. O. !-il. U~e~, AIm'. MIi'lMilr. 68, Ull 1193'3)'.




i'l~ <:JU Myf~n~ ,1\·88146,' zillCOlli r(MlflOO i~iJb <~i8 1M}! iner tm~m~··silic<ll.c di.f1i'eremi'fIEioln O:il! f EfH. ~r zirool1:s 3.repr;inUll)' Ca:y~~I.2Im:rn~OCJifl p;rodm:ls ilS ()bs~.l'Voo UII mm~c>utl1lmrphos.od

OOlII!U grtll!i'i' IlHI Ih!ickIlC$S

of 10!;;m ,(15') or up 1.0

crupt~ml~wou~d bu~ tliic cadror lruy~rs, which enuld Ihen be I'h(!iiI1Im~ily

25101m f 26), lmelf

la, It" :khl!~!I'b~rg. . 5. K~bef. K. OlHi:IiSO!]',0, ~l~r. 6 ~;m. .r::jj~llTI. .IIetll U, 3:151 j:i!OO2J, 19. V. t3~m·~.i!i.M!!~.g~r.~. fil~ml!'. !E. SciIll]<mr. C. MIin'I:!Ilf, ~ • .r;a~m. AClilOI69, !S8~ ~2.ooS)',




(6), then rnanlic: d i:r:rc~el1la.Jon, 0111 l EPB lasted [or a m~lxi mtrll~l(I r6.8ulliliioll )Iearn mJtcr~llet~l-si Ikme dfJfrerclUklt~(JJ~. IIi'thcym.'C

now.s wi~~ (1e~ WI:I.Ol md:i.onm:!lkies t(2(>A~ ~m.d MFcJ arc: dcp:l:e~cd siglmlfirnmly \\'ilhlll Ihe ~.il.st t1;:i!.vmillion ye<!!~. :>oooll!dill'Y InetflmoliP:hicp'rodLjC[:s,.IJ~cn dtlirll~ aner' Iltcml-si n~atc din:Cl:'Cn.tfilt~o.[I of~he EI:iB. The ::c:ii"OOll of' Asukll cl1l~rutes. \vhic!l fQrmed ~n ~ion ,orlgnCOl1S iMlii \I hy on :E P.Il WOS C'io/'cifl silm1Cr. By lIS]jlglhc nwdcl ~II.:'\:V abscll'uic <LgC <!l.tl ~vly aReranetm I-sillica~e diifl:'creu~1i:ath:m «Ill forr malllic d~ ftercntia~ion oil:" ·S(J3.4 ± 1.5 j\4" {J~l. EPB" cOJlSlmfnsdw Imt"'lnuun dmmiOJ10rnlomie 4 we :'i.~K)'l>V the zircoll IInHt ab~.md.mcc llJ:ldei ]hiJiJ. d iftb:rellli'al ion on IEP:s." whicll 1'CSlII in [he lis or:,gc (lfA~S:·I.35li8 ~s. 4S'()3 .,~ ... 3,:-: My OlJ!ldth;lt Ibi'~nalkm oj' c~lcrit(.'l'I. 11l'Cl),::thn:, 'thclJm.~, ']lCI,i' A.}l}[146 7 ~s 43M .• :±: 4,{5 My. O ~n{l.t[!i~i"imd~lc 00 ,~H!'«I ,,~nWCl \llnlU:] h£)l I,w'l flmv.~ will Ct::lU;l,~ ~1U;:ii.;1'6 A~ ,IUd wl~c 'In; ~.l\i. lhc ~~I r-W ~nodi\;1 .. g>t.;i~. or A-:';IRI3Sl'; .(LL1d

hem,. I kIt IH\l3. l'LCfli",producin,£ s:h.olt=l:ivcd

~ ~~. 'Y_ ~ Et W~~~IlD, (·Y .. Shl~, H. T~ked~~ iE'(fth I'IMrM; Zli.4: Jll (~~. ;!l, G. :5rimr"'an,tr.rn~r Ft(MJeI Sci ~)lJ!~ 14,B5I fa'~.)'

j!1J!. l

'Sa. ~

22. K. IilI!Il~r, K D.. Bcti!ll. 1-1. f\alm~. lB•. )~~ttl,. O. ~Io.fr~r,. Pfa~e.r;¥O,e &1: .~i!I~ U99Sl, ~99 ~3. E,. S1lii~~f,. G~m. C!:;!~m" ,o1all' 41, S@7



i!41. tI, ~m~!J(h],

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A-~~ 1467 QvC1flllPW:it~i~! c~'rors. flltd they f(mml~ wh(';11 :MiA.I {djill~t'llallCe mal~d ff.OJn 2°1\.117/\.1 ,;;: t xW-lloin cli;eril,C:; :P.iip:~iil Kl.l'I1l11l,fI1~d A-Si'l1.394 tt,l • .1tJ • .1 J )10 <lI ::>~.gl1 iliI.C'il1il11~lower ~bllfldailcc
of.:;2 x 10-7 .. The m()cde~ -(>i\I-:2tiMg~ges A.-8S146'711i11"idA-s:u:n:30r . !$6! .My UI))at1(! 11Gil:;lIl.'fiCi:C_I~i Iy \\11/: II c()n:;lrn~lllc-d to d ~ ~.inguisr~ : ..
bctVl."'Ccll.lj 111~,eoir cry~,>!a]:1ml.i.'Oll b



p:hic llVC'1ninl$.

ancl lnci:mxllor~ Th.e mode~ Mn·Cr ages of

1ili54i6 tQ 455 (l M'y 1:01'A ...l:314{1im Id <4S4~ M)' ' ror .I'H:.J:R~3!l1t .1!!1fC ym,mge'~' l2m ;md POOrl}i co:r.ns~rn.incdl b~a u~e (I rUl18Im'll],oa:p:h icn..'l:lJiO'~[il1g..

;):\ h(':{u ~UUllrc~, TIlcm''I{LI eV~l1.l!!.!ilnll ... rn.odels fur EPB (7) :ilI1~,C'ii1 layers .> 1.0 kill. bCfI.caLh lJ~e ::iJurll1:ce an.;;ml.(':ll.lpcmlillres >1000 K allc<L<;tI6 :M)' ;]Ilc, lliQCml·sllicatc ~il1('~llfimion 0111 :EPll. inlt:1I1Blll ml.:l:alillarp.~lisnm h;:acl.~flg ~o [ccqill~ l~brMi.{l1]of r:~ f.:\;\f i:lOIi:!pt..~ !E:twretilbo1lli1k m:ctal am:]: t;iliic<lij~ phasel> ~.Il.e~crit~ ~e.g., OUinld .l)OIl;g.il.) 31.4547 L 2My ~9} po.~ndmescry"'k'lili. 2:lliOll or A:ruka. zkccm by <'II.t~sl -9.9 M.y. ]j" /~.:s'lIka ziilN:OllS. :repfl\.;~~'·nt hlSit ve,sl.igl!s of tlte igrlool~s ct)"!II;jJJ~li7"'li{ln athe smJacc' of EPB, o

.2!l. I\\. MI.!I'1ImoI~\ iii. ~<I'kNa~ Etlf.rII PlrJfJN. sa ~i!lL i!.22.. 3'l\~ (1'!l~~. ;!i'L l. S, I)e~rney.,. !o!Hlllr ~tI SC£. ~.3;;!9~1'.il9$127. A.~. H. 'I': Ml.'Sif,.un ) .• lcaroo: !134, 187 U:99ro .. ;li;l. t:".Fl, tlJdw!9- Ql~bn ~l1'I'l11Q1i 51~i 1.6, ~li;~!i~ :![(!. 1l9<91l 'il!F~Ii!'j' G~hromolll!lil' CMI~, 19!1llJ ( 29. SU!J1l'IlIi1M lb;v <I Mitl!I.Ir\l!ISdtlil~ Ol~dl fi8gjlletr.irlg R~ir,~ C'I!l'!,mtiL !;If Q~ad'a g!iiint 1;0 Ci.S.
(o~lTIIC!!irnemilslfy LmfdlOO;i)" Co:MiTIm~~ionM"


N'4~d~lm r.!ICilit'j bn ~lcdi:001liT1 I:! (tpt1I1llI~ 1J.m;r ~1iI1 1Ig~~~t irrlDng ·the· r!l(JntN~ 11i}~~rd1~~!ilOik:..hoe Ged1l9iCllt SUM)'!Ii~ ~Imlllnd, Mdt~~ S'1\'edim MIi!MrI'Il af M;JIjLij:\l!~Hmory. Thl~l!!i; ~~rd~i;m('OII'r1~1J1ijoo 00. 184.

~G.5J .,di;ll ,fI" !J,liClmuli .j!lnclc~rIlenlli .hi!lli'~ WimffOlli! Ih~ m~ii~~np!,
Clil1lriile Matelitlal.



~.5(~~miI!J.Clrg{C!i!iI'CgnhlnU\rLl!ll1.n ffl;Il~fi'dl$ ·aDd ~.le~lilil~.


Tlu;: H r~ Wtl,l~il

rrom i(in;\rnl "Did M~1·C'r ages of

~he'ill tli!C' me."'Ha.orp:l~ eV;;;;nl r~.lting Ie in I.~r·w ~l\.:'t'tliJi l~bF<u~icm ~,I'('a!1tel DrI'J]g;JI. In 43t17 ... :2 M}' <

:"19. Sl

Tabl1!:s. Sl!


U MoW 2007


il'i in'iL~"~~ :.'lq~lt~~[~'sS~IOW E11,~tJliro.ll~i Cr)'$lall~.iza.'

Ih'}l~ rrec!..'tIc(1 1'i'Q.('tnnil'io:tphi~m,of A-8813 S& by

CMJiot ~l!c due ['0 lav~. 1100w I;II~ lh~ ~l~rlhtc1."Ji· trgi:i!il'ti 1J:y ~lL"iIl';PlpdLtdil.!!ll6AI ancl'r£QFt'. hTi:~uiiig

.231filllUlil1jl' 200'= ~"((!I't!ld

l~lUi!g.rllCii!m~e.. ~OM~· U



Bo,undi,Blry Lalyer Halogens iln ICIU'ISI I Ani rctiea
.P!I~~O n®® S:.a,i!l..l.operz.,:l!,~~ Illi'l(IOIII' 5•. Mallil iilill U1,.:l R,h'ilallil A. Sal~mollil,.~5te·p,hi:lilille ,P!lliIlliliiI E•. J IlHilles-,.J H:IJiIl'llIl.r'd K. Iit;QSC!CU!!',;;I; ]ollilllll M. C. Pi~<llrn,eil,~ 19l1lW!!Iens lllrLlU)entl!
,a@lFo50~S:, whfd'l

CfllmlJ~ed release

or 1PhQtQ~a~i~,c: b:rwninc
.;I!~tI B!fl


li'){lIlrJd~imo the g.'1i;'l rl1ruJi; (4)"

Th,;; cQ~~~lnn~,ij9m ~rlD


~~l~"'I[a!\."dw~O! ~l~~.Ion!g~pel!l~d;rr~~~ial


]1•.·:8. BilIIll!Jguitt~. ~

the o:xf:di{;inl~, 'Cilipil!CUy o~ EiiI rtn'~ tropoS:llnllre,

and! 'iedJilne o:(i,~e:s Iorm !!I1iIr.3,li


:ilisoqlti!m. sp~"d:ru~{Jlpr i...,~~!ni(w!C: (l.P.DG.A:S) VJ~.A k,d p;!I.i~0 4 km {tom] pillb = 8 'kl:ll) W<'l~ s('! l~P,II!I :i1L;'ight of4, 1.1'l'5'!"l1 .. br.},,~Ih~ s~owp~'Ck ("1Iryi~iM 'U:aooughlht;: )~~r~b'; fl n;;;'i'l,l~l Ofl'lllHW 300Ul'i'lr1il1~liDnl. ml~ li~ht ~t.ti\N:W;L~ !1

fnWoo. o,,~



may ~ii!'l,iiC,ilIll imlpaCit on dllma,t~. W@ U@PGtty.iNlHOIUn~ 1lfIf.li!SllIr,!!mer'lts: bol!.!~ltii\!ry lay@!' ;'od:if'!~ Q~]t:il~ ,and bromiin:~ ox]t:i!~at the flf,aJr~I:()<1!rS'l;~ts;litE! ,of IH31Ilc1l"S[::u~(I~.

;;:eilo:ti~nmp. \~VhlO~ b..,"9I]~ Wil~ rolded baekte !l ~kwl£ji"iI!~l te~e~oopc receiver lly ,ji]l Iifm;), of
'1l1t1l1tzrolnCtt:..etjbc~efll!C~(yr'S, 'l1ljl;:p.atlHll,Vcrn~d OOIlOC'J11I'iUiohl!: ,~rm ru~dBr-O \V'C1:c' Ohl:EliblOO by ~I~tin~ blbca'atiory-hlleJsuood >C'.r.tl.I",,> secl,iO:I~~~f Ihe nlO'lbcWl~C8, In OLtntospllcric .\oIKlC1f'dIJctomoo ill dl(l '\.'i~ILbIf!; ,lnd near:-ldlr.n iO~~I, r~UV}. rosjXIcli'\.cly 0'5) (fig. 11.EI(~ ,.\~'(:gl Jlax!l!II,[:!:r;!' 2004 ,md FclJ..

Alr!tar(:th::a. 5!Jlrp'.ris~f!!Jlly, hth Sp,OOElS 31~ p~e~~~n thr()!ugh~m~nE! SIllIllII!1t. p~nl)dg!i!dl ~hil;i~ts:im~l,af' S1~,S:I)il'!g~ e,ydes a!ild' ooli!:ce!1u:at~~:n:s" fhe~plril"l~~tli!'t~ IP~alk rof 'ii!)di~r~ 'Q)[ide (20 p.grts pew trini~l1) ;is ttl~ l1i:gh.~$t(l(lf'll(eliluatiil):n f~(iO!r:ded i'lrnrwhe,~e ~if'l 'Ih~ (!i tl'l1l~$P Iil,eli'i~. Tih~$~ I~e~e!lso~' Ihalll'!ll~I'I$ ~all,ls~ 1>ulnt:;u'i!tio"ll o:t'!~ne d~ple'liD!il, o;g~ \VeWaSl the rapi(j r[llXid'o;grllolil IJlf rd'imethya~I!IUitlie (!ilii,rj .rJi!eir!Llilrj1'ill! th'" M~''IIr!Lticbel unda ry laye:r. ~'n~h~~ Anmrclie,,~,,'orkO'Il buun inc: spec iC!1. h<lli b~t~~l !m~in!y roli!~cmcd \",hh the h(llog~n,· ~ (;hL;!IlU~l!)!~~.lC';)Ui'!ng roP!cl!sl,Ir[aoo (lmn~ Cfil<H.iY1..oo [n::nlov.i~ g·f ~nl'!ew~~h(: ~~m:t{l8pl1.Cro cl,c,pkliQI] (,>iI.~{lIS r(ODE.."l inEh~Aire!:ic $i:uirlg~iI:lHl (/). O~lI.:nf'.ltiol\.~. i;:I~'L,:r AIl1:(![-c~i~Ol of th..: I'm,,;. U)" Two, Siluro!:"$; f tt",Lcai,'\: l~lug('lfl$hll"'t:~cJI o gu:'3mctl lmp~ll!l:)f.ic. !;:o~lI~l1n cle:J!.~hi~ ,of brop;ruiPo.seJcl: ::leklilicd scfl·snft SUrfD(lC'S such :JI~ mitl1.coxide (UfO) !'LJridu~:Hm~' oxide (tOl h1~\i'e ~'crol>ol. Ih:}Sl n:owC'rs, or sea S8~[ ill surfaec snITW been lrCponed.m. lit ~OriliSlll.1 site (.8, 9) and, illl 1'111":: a.oci.mu,lilrnLed Ihr.oug;l~~loriz.(im~tl dnlling I:roi'l~ case of 13,0 O"~}I, [rrnfl sp.'!I'c,e(I{JI). UQiWC\"(.w', re:fm~1lr lC'il{I~n:3S 300 p~mlooC'g.mooblc' Imlothese m~iil1tc!lll~ed melIS!ilreI'llC.n1H': There' IHlVe C:lrlllQi~~. Oml)Omld~ I'roll} ;mT~llI'OllIOgc'~lkand O bCCllllO d kcc~ measure 111C'.tltS of 10 or UrU ~I,mmd on,girls 0),1 b~ogcm nre ;!~~(li~wo~~~oo !till inlhe bounda!)' ~'[!!~iCflll()v'c [he S!](lW p;;lck. w ! i~!~hc n.,,,,Ld.u~D!] of O(;~>!;I~i,C' dinlC~h}ll ~lid~ict..~ It i~ al,'1() nO~i!~I1;'Ol1hy d~atun :l1cwc!m!Arelic . ,(DMS), whidllmny ~llnlJ:Cillre' ,~ml'LtiGll orckl.ttd IOC<1tion.s, ceen whc:1fI elevated B.rO was jjl:1cscltt c'Omlells'lI'IOll Ilt[:c:~ei t3), :md Ihe o;.;idation of .in ll!IJC'. boundary layer dm:i:l,g ~lnODE.Q ""vas, [ gil1iCOiJJ.S cltmcl1ta~ IC~g i~dlC Arclic toolJ(lspilC'!'IC not ob~f\'!(.-d ,t J I. 12),ii!!Tlhough llI"c:ro~I! 11lC','~· 1(4), ik~ll!lly. it \,V<J!i. S]~OWll Ilml (:mld~I1s;[I1il~;L: !i.!Jj!lconu:nl$.or 1!~~crOl~]Jc iOl~~ne h~nl~oocn !!C. i(lt;liillc o:xid~ 'I,ipms 'c;JIn1l~IC~crH,", '\",ry t:'f!Lt;~Lnt!y j'norlJ;id(l3l A ~](;~(I C'mllp:ill~!1l11 'wli1>pi:'r.lbllIned from ~o ~Q[]InJIlInidC'S, which lTIuyllll\'c rhnl iiflltpoc~lUI Jmlluury .2(n:l4, TO f.c:Dnmry 2005 atl ~!rcl.ky Shuion cloud Clover andhence 011.d~ Il~me (5 <, .6 J. (75"35'5, 26"3(fW), w]~ieb'ft uS s;litu<ll~cd em [1iIc Bl'iJmtIcc'Shcllf. Ant.fl]'>Clica~ olxml35 in ~ooV(!:>ea, lSdilOot 101<JhFmislIy. lImii!jl~Jsi~ of I!ml's, lI!C!d:~ 11;529]1; UK. "$(11;01)\ O'rEinWoOOj)ei'i~l Scleooes, IiJnwersilij o~ Eac'Sl: Icvc~ (ilig. SI). 111.C'ice edge is smne 12 k~11l011.h ami3U km west ordae .stalion: 20 Ion S:O~ ~h,\!CSD ~:l Mg,Lla, I'i.IQrJllji(Ii1,. NM 7J~. UK, 3Britis!ln IUII~au.til: SUNe:Y, rN<I1;lond!. E!n;ii fCllrmm! IIli@!i@.m:~, C~.mcil~ Cilmluiclgeo, (B~ PR.'CkmsEhy. msC'mt.lir:!cm;t1ne.IlDI:I,y open p:J.lich of OU:, UK wm(~I' wilh h.ll'gC ••reas. of [re.'lh !5C<ll itt where' IImt ·Pr'e!lEfiI!ierdlte~~: Eaili1lr .~nd S~a: Sti.eooe Dhlkl On,. Jil~: I1ffi\~eli"s.,grew hI \:vn IllJC'fm~d spring. Se8-salt ion
Of Ih~


~~m~ldl1iJ:<lbh:: ;u!en~ lion ~~. oc~n gi."'~I~W .dl.. ' r-ole: of ~romin~ ~

G}:a.-;~2. (k'>CII{_l"!1,

2005. 10 w~t~I1lihc.[b';l!U!("(i OIl 41 d;,ay~~u~ld 13rO o'n 125 dmy,~.Thc'IMI\O s~'i!t.""C.ic.:s, Ilot raeawere :>\~~i~ l>umull4MI.ool!ll!i.~.Y lX."'JC""l~~i~~\y "h.<;(Irn 'in dil'M~


~b!li~"lOJ)',. CJ1ILilomia lIi1\'~tjW1!erOI Tedli!lOtGgy.

~bf\'ml.fil~!fmi n:g~{J.!L~(Fig, 1), 10 1um;l StuO \~tcre cli;-r",,-C!J.,d.,IhuH,~h~ OC'[t(1:~tm limil wh~n(.,\'\.'I"dl(l SoiM ZCillilh~[nigr.e was ie'S!> I1K~j, --921!1~ic".1\''1lmg~lwId fiJll d3)'~i.WII), Thepeek OOIl'CIl.'lilll'iltim~ ofooth !in;i!;:sw~t! droll!. S x IO~ Inok"{lu~es ellil-l or 10 pans, 1l!C'!' IriUian by VOIIlI1ili!' (ppl.v). oo.'I€'rv!Xli durillg su"'n~ ~rlf!! ~Ocl\aOOr). I~Q:r BIO,~hi!i. OOUlIO~'1:r:nIl0nis shlli~;),rm ~1c! 1C\:l!ls cb~cr\~cdili1lhc ~rringlim.e AIlC~ic'~ 16), In 'cumms~, ~lCIO 'OO~'lml~olIi[lT! is <Ill rP1Uer 'of n~gni~ude I~' th.m Ihc: 1II1JJ;ilCr .repo:lt~c1I1'Om d'le .An,'eIi.c limit UJ. 11) mKi ~snc~lJIll!l,y d'bC'.iJigbc.~ OOI1(i(,utmlton ~tocJ rnl)'w~l~re Ur! lllc '1H1;[QSI~1L'K". One i~lp<!ct or &rth high kwh; (]:f ~odine G.l{~{_k."S i!i. lfu~ ror1rn1111irn1 ul!r.J~1111~ :t1.:!th<iol fmln Iil'l!(; i"Olj'l1Iliri!llLtOClll! air of 10 !ll1d ()[O ,t6)1(0'10 !Orms.fml1~ the I'l:.'fL'Ction of K) ~/fifih ilse:!" or will'! B10 J. \.... hieh migh.t. explain Il!lI:1:ick n.ttC~~1jJtiQn(:vcrll~~ ,ooJienl@(] in summer (lit'Cr SICa ;l()~near AnlDrurolica (1 The 11lOO1Cenll[;fuioJls of both~O and 13.0 C~.~ ~ibitcd 3. diurl'la~ cycle. cssclltial:ly Imickin~gdu: solar ro..diati;oll [figs. 2i'.lUlcl3,). l1]b, h~[lkmteslhm, both srcc~cs OfCprOO!LlOCi1 ph,Q,l.:Mil~cmie3~JIY.f11lf1d h.wc moollndary ~ayer :iIHl::!nlc or IcSj!; tI'Wll .2 i


h~scl@iI1IlI• (j'\ 9nO~. .

fTO wham IlclT!O!Sl!KIndem:e! 5hO!lJtd be addreMed. ).rw.(.IF'!!!~~t<@'l.e.efiic..l!i!k


OOlwcllt"tmions on frost j;1m">'C1"S .anel :neru.'1)y s!lIrl:at-c :mow an;: ~I, tl......lst Ihrec .~ inle5 glt!iUCJ: Ihan dlQSC gil SC'lw<u.cr. ;md 1,1iI has, ~lCr:1l Hnkcd 10 [he i.~

hours again:s.1. tl(mll~lOIfl!Chem~c~1~ossproca>SC'l!. lIte wnCl:lntl"al~m~ 10 ,~n(~ we~ SII'OJ.l:gly .orO co·m:iI';J!t~ \\'llh ~h<C' ~O;;;:-;ll"..~ nril du wct]on .~ well ;~



w~te$ (!if


rit~ The fO~d i~lIliE1 00f· resJPOJildsoo~h~il'lmlF spherl'cspedml1fl .[I]mr ~enHI!/ililgaU kill O'rrIJI
itlib:serwr~ e:u:ep'~ t~,~ ~~E!d!1!s 'O~ ;.nt:effi~i!,~Vitih ~he O'If('rtyiO!J ~ef~Cl~
:spE!(~Uf! (d'Qtte~ U!fIil')r ,c,alltuiL!llier] rifO!lI1! !iile<ra· ~yre' gb~f1Pllj(lf!1 r(JOSS

'S!ocllon~ llO;~asulr@d .at .298 ~ \27, .2$). fh,e rit~fJJl]pro~diJr,~y,lreld~ (A) IJO.l ~ .20.5 ±. :U !lpt an.d CD) (BrO] '" 18.8

430 435

44JQ 11m


Wra.v~le!,!gth /

Wa.\,!',~II~!lgth.1 m n

1.0 P[lt.

Ille cmnlXllgn ",rM ebser-ed in thCp:r~lbC~ ![II· !'l: 1 (If !OJ {c~!Of!Cto lhg Ot;;,to~r ;J\'emgc of 19 in $imntll!ott~!~, IPP~J. :a~\t;';!\~ l!I~c~i~ i!!c....lJmQ;:i~liltNO ... il~ ~r.rtiol''''''01' bnl,h I::;Jjdi~!~ ""''I;)IJ~ Oht1~'l;l~whI,;11 iihL;; h"s; !li:I'! ~:! t'O!l~N~'d 'RJli\..~ij~Jr.~~J~ lhe 01'01 ~'P1~·!ijm!.l~!$K~ t\11i<l!l;;ii(;HD1;ip4,l§'1111!Jtl;:. i li:tolJ!1d,,!y I""J~~r p:rnnmlJ;l l:lhOlrn;J.lili,;'iliIli~;l.! pm[0 Willd\~I~ tt.:mIl ~..~ Op;,::I~ i~ f!"om _iII,K':t~ t .",4,00" ~, The '~~"Il,miti !~.Ciri.ng the c-.-nnpaiglll iOho\\,i:oi. ~~I:Cl~iII" (Iii' O:!. thl..'ru·m-..: hw jC!t,:!.:;,:.;~hk ~:plrrl" 4!5:1I~in !lit tI"fliS..lil"~i1:wthll~ ba::1~m'ot..Y ~~",[~ir.; Wilhi~i ~hl!f!~Slll)l5,1.:;j01Uial drf;'PI~lio!~(!"'~!l.1}. ill S~p:tl!nl~r ~~miO!\is:Ei!li~i,;" ~]:u;:halogeJ.l:5. w~w.fl"l;'¥ihly ]!~lod ~he P~\li(i!WI!l 24 11(1UI1l, A~tlmu,ghtn~l(;ve]~ or ili~lh~ OO~J!cr,ary .!ki;yct nir less !l10l!l 1\ hOlli'S h1QI~~ {11!>!:d~ ""r;'R; Im'!l\;'f in 'IJOrw~i!iA;:!tl:;d mU'" 111:.." and Oca~t 2U04. Dliri~]g l\M (I r I~~ ODEI:l, l~h:i:(~ 'ii'/,)t~ nO h~~ogt;ll ~h~i~ 1~:C'a:;'lfltljii(;lia~, IX.:lhre w.lc~ing H!lnc~. Ml'nhUl on~.y !I~liii~l~l m!~d :~!mbc~~':iOVI,l'*-o.::~ ot rO:ta'tl~f~'t J da~IO 'Llt>d ~ jjt)eaiJ:;~ b1i.cl v liSibl,HLy in. ant: e~b'iCRrr'lcl in the qli~lIjf~iry .oJ (II~ thrlli ~ 2 ppb IWu!ii, dl::pk:lcdJ of .BUO ~r~re l'if~I::~'i Io~t:\'~ sli~ rn i~:'iitxili:!!ir.lll~!ll.'.. IPlil'i Y OOeL1!li~e OOAS\\\2Is. ~;jl!il 1I~'!! to abIhr.~ !~whriC'h i~ s ~ t~~n 'OOih. .. wi[h 1r:'LjL.""Clmtl~~ C:iifi.~il\g Ihe -,fi, jlpt JttLll1g t>lJilJlil 1A.~oo~C'I~ru'I)1 i1_00l,'l:: It'he o1~~C'i' i().."'r-riiOi~l. s.cctlU sueh Iii;; :.I1:OWDI in Fi.~l'(, 2 ,alile!l.3. .1 iiJrwi!\lcf., cl!liriiii'lt [h~ immu1I1e!11all ,th.'lcctkin limu~of ]to 2Plft (Fi~s " .2 s.c":!O! a di f1crenu. ~"'C~,. third ODE Iilerc w"re IOI~:1ca'>'llrcm.eIlt.~ lhllLt IOf)}, 01" ()~ 'wat:; ,el~us.iUlcdflm~, lJih~ Fi'CC1roiJ& and 3). Thiis;~tro,nlg~y ::>~W5i'l:., Ihm hll~o~C'nl <1'(1h ,uiml hl :ilmli~hl r..:. w~desplli.."OId tbc co.'lI:!I<i1 al)llCar to berullic:oJrelalcd whh OJ lli~" SJA I. sphere "LI)id~)' rCl1.Ou::h 10 on~c,t ~,i~~d i~l Ikanl: Ama'rolicOOlllldary 1<l)'CiI'iiI~ .dhl0t roi1~riC~~'<i dlb.,>'C' ro, hl~CI'CS<ti~WY, ~IOWe'l.CiI:, ~Iot.., O.rd1C' devimoon of 1'Knm(31)' 11lr)lCl"cJcI)lc:LiilUi {thb\·\'m.l~d req~lrI"C' a f-ifO.'(imil), ~o nJilCioe rmnl,[Hli:l\~III~lbn.'illCC is .sup" [IOJI Ikll1~it" :il;1!1l,s,"I.lCl'm I1lC.aJil n~'~II 1101 or ~1\'m,S hlllfose~1source 00 of[~t the ;sinmltarlooWl~ .. 'l<t diJlnion of 10 m1d B~)l. h1Ckw. dl~ l'oOlimcll;uy ~I~(:d b}!)l(I!lldlilC G~:va~ion.'l lire BrO ~ [lBuO~ dt~>lot ,~h{l~r",'~ :ligH~!k.a:~1t , emreI01!Lo!l Ulg. S4) .. IUlclb~,d,. rM hugh!;.",,! 10 C'{1'fK"cul~mli(Jin dui!11il I~)\;,J''J! Hg~.i~jIdlJri'l1g the S!!I!.Il:~il peJ~{~d On!IJ!l sph~ (;'Qhw,'!I!~ di;n.;;j~'y0\\'r/\~1IarClica (.If,!~, hiili! w ,~h!l\,VI,;l.I alOi! ~~II"£C'OOblUI ubu!ld,~n("C'i ,Ulipfi,,·8·cru abm'l.l"lIcN~..;m' areas oi' Amarclica l.x>~\r\,'l"Clil ~'hc !i!!ir mass Ql':iginl'l"OdiclOO. by b;;Jck~':<ljec1QIJI

I.llc (_1cP,I:ellm~ mill;" fl}S\iltirlg

fronll b;rocnilwillld



1 and J}. '~1ilimnq;;C!!:1OO~OIm·~

~I"!al1!i~rtl:)'. ONlpa.oo O

\:\'!i~] b~!1I!ij!"N

.. !~,




Augu,"'>! mld No<\t(.'1ubl.~r. wi~h mmUcr


b~tw~Lin D(;OC'!l1bL.~r APfij.~(J'O). .md lhe ~"t:(udof If) guul~ B:r;()et11it.:i.:lYm"mio!u;. m'~r 111C(,'ffII~re ~f{.'frris d!i~l'lary·c"!tl ~ljJFri!} ~,1JIL:: (b.~[CJ;.




lUi~OOirib<Hli!i1It.r;; liIiC

qllit~ SOlllli:r:OO

1iS·81 rc5lllilt

d i:f:l.crenccs in rolzllrl'udial ion fultl \Vim:1lIiililCdiOJ1!.. IlowevCf,. ~tcm.dl!l.y nJ!llning 3,~ OOllccntril .. !IOf15 (roHrllines in 1:~ig.4) slrow~hiili,d1.e' ~aoom:1I c'Yd~ Ono[lWU!] IlrO are renwkab~y:simii~lr, li>otlil in limin~i.'tlld :lhsollL1.C' cono~rm:'!!li(lll. "fhi~ C1uillC is l!n~,~:;H."(:wd. ,g~w,·..n Ihe diftb!i(;lll !\OtH'Y(,;\~ ~r[h'l"w.l)' imlo_gl.""'\I1S (:i!C'C below), There is II di:!!tin~cln'lll.'dnlLllill ill sp.ring [nl~ SCCJlI in thcim~trt.""d~~'OJllOSPha:ic: CQIl!Jlll!~ Hl.e<lSIIJt'H1CI]ilS of mn t9). whichm-c ICOIJ~ ,~isl!::!11\'ri:!Jr! (lll!1"n:re<ls!.!runil'C\ll~ iJf,~nlhJ; 1.0 \l,i.I~ iu~
~h",tlt:1!.llild~nY~;i)lcrl_Uis i:-i (:o~IL'l"\N!1,;dby,u dt1;~l!."~





II '" Ii !iiI iii
iii ~


r "',

-70 -60






[he ~ulI'l1lmj('t [~ndl r~P<l",,-..1bl~ ~("Oml'dt~r;y


SIiI,mller .I:C'!leL~nlfl IilUH.II"I1In~ cOililld be a ,oonsCA1~~ of~O'\ycr mite'So:f hfilOW-""'j;l acln\nl:irm fir-could ~{S!liJt fn:mlfi k&...s;~Ll)~eoundb at)' Ifltycr nll!{)~!(ing ·IThriSl.er mixiili£, if1rt©,~hc Ih"C IroJ)O= ~~pl1.Cre.l)ul,'[nglhe Imlnl,' nightlhc halo,gcl1 oxides were Ult.."'VII.T ob.'iel"iiOd a1mvc~hc ,dcla;I~OlillhlJil of'
IliIlru."l~mIill11l. inlmlltn[J1I11,TI~c

Ill!! hISmnlill~~q~ ~lcrvle¥Cf,ulil!: ~1rs1 rrpperurall«: ~o and BlO OOC"lIrs very carly illdu:" Amffi'ttlc S~)riug, dmins twins'lu: Mi;"in,g ~mlim, tip to I}PI, '\VeliC' dC~(i,'!C!ed s c<u1yas 9 imglil:i!~. Tllrsim.l~ a an ~tttiVturm~ nl~clm!l;f5l~l IIl!iJa is c.~"l'ich;:nt i'nlh:c O!.b!'!CIl~~ (l~' din.'ti!: ~u!d!igln OJ' 'wh~"l,;:~h.;;' 'H;I:1v.mJijm~ (:K,~l.IIn"d r~n~)(,;rnollh h~di!\.'C1 ~IJlli~h!.\r",i~:! i!lr!~



b,d~~1Q. o:>L;dr;,.'"SlhC!lp!;:ni~tiul~ fOr ~1k.;~1.O~mllDN ]1

while being 'tr~I{~Nflmt",-J to 1-if.1JIr,(..1"<. Vle lllOCti ~ .ph01,od"u:.'\lnic:d b.rv;: nnodel (Milre~ r.llliisand Mctllod.""to CS'Iirnl'ltc ilte (;GnDrJ.bllilio:rl~ of ha ~ogc.i!Itihctnistl)' to ~o!Jm!rlrry layer iIlZOI:IJC loss in the po iii.!: s«);r:ing. With usc of Ihe observed B.uO clillima~ prol1k; we e1:\1iilll[~[~ d£ ,diLLlnmUy aV~ C":I'llgt-dm~(;of OJ 10&0 b..""'CiIU.~ of mmnille oIlcm~ islJ)' alol'tc'lo IJc 0,.~.4 POIl'IS -p...-y billiO:IiI(ppb} hour:"'u , f\ si~Hi~ar1.1l1!cl~:JIID~n fOl"i.ocl'inc chcm.isGy .. kl.l'': a )I~!eMs amLe of O~reilll0\ ,1I o!I' OJ, 1 "'lib hour:- , '~(lW~; ""he;!:. iodine and bromine' ch;<lmislJ}' .ac~ ~o<~ru,;r, d1¢ II1lOdcW 10iSSIUteh!cr~~'l'ro OSS ~Ilb htll;lr"";l l}(;''.cmlllc l!IlC:~di(Jf~!O l-(!ltO Br + Olni~ii:rcfll>t,'S I'he r.l1,,~~~ whk:]'"! UrO i~o:m~·\e~"d burl{ ill E), (J .'I'~til!;. S2),'lIliil'l. IbiJl'ifokil i~~rcilS.Cin







-oIlC! -20





fiime in '~IMil (h:0'~1lr"s) ..
IFi,SI., 2.,



dil.m~a:l \!',aria'tim'l oifthe U} rni:<in ~ ratial~lJ~idk

POllFllii \,,;iiJh :2g: IlriliOf ~i:ilrs), 'the DOA:S

SQ!l.!IF ]l1'adliarnre "b~ar(k IlIFl~~. TIIu~cillln"'€spC!!id]n~ 8=dz!,y alr mass bile k traj@(l(ll'y ~rl'hn ~t HaU~y (25'0 nll FIg aimve ~a level) ~5~~m'l'r:l .. dj ..(:enl~ loeaid'il dilJlrl1a~ plot. IE_!!clilcircle m'l~hetrali&W'~y rndh:..~:e~ hcmrs. , .241 mr,aj~rClli'rie'S WIl'i'!ll cakllJ!atedhoml 'the I:.!i!ti:sh Altmoopherk DJlt~(iflil'lt~ W~!b service. whi(h I!.i::;esthe ;El!lropeo:!r1 Cen~n~ Medilllm·lRali1g:e- We~~her Of"'ecas.ts; ClU:ih'ive-. (A) A d':IiY~r1 :spring (2lll Odo~r 2!004) 'r(lr F dl15pl.lyilllgl Um l1iglu!fit. mi~]r'lg ratio, orr :;:O.~ j[ 1.2: ppt Ob5e~\I'I~d ,duringl the !G8impai:glil; tlil'lil air IFl1liMlll u@~ecto!l ifldif-ca~~ @ :s'~ro!1gCie'afli(iJ!1n~:I,llilc.e, (8} Pi ~f lrl ~pr~ng('lL~S o (I{~~be-r20(4) 'IWI~llith~,!i!Tr ~s;s m Ilold rHlt been OlJ'er the Il(!~a"rn ifor ill t lea,"5~l3days. (('~A. d~y ill 'wm m.er ~29 J.1I1ilL!It1ry 2005} wii~h 1I1il Doc.:ei1mk .;!iir' mal~:S~(I,H' A rJ\i!y 'il'!l~'I!~liIter (4. May 2(04) \l,rl1:E!'1li1 ~O ~~'ai!lil@ki1!1 dete~liam 'liimn~. ,ailQ'IIIt.2~ppt. the 'o,{

detllction limitr(lr ~O ("[,gl~k ~(PJarl'les\!lit~l a dillUed llimd, \~hiich v,glri!l!!cldepe'l'ldilii{jl on vi~lM~i~, (llli!d ~he



"'llriR~il iI~'mJi! ,[cs:nol $ironSl)' CilPl1lt,x! (.19). Thes, ,.Ilh~gh dC"'i!!cdn~~Qgcn
c.'l hl~l['t~ ~"'tWl1W

25,-.---------------.-' A 2011"1 Oct '0<1 I

~5, ~O

Iii l


, ill!li

Q~i'd~ !lflii: ~!r!!r.lfill;~1i~!~y ~PTQ~q;~li,"i~~ rClr I1ll Ol)~. ~l~ d\;'gT~~~ 0;1 i;kpk~iQll. !Ijlil}\,'1!IHl>il,ll b~ 01' (!O:1!A['O];k1d. byi$liC biill.l:idary l[t~rU'i!l~~~IVIU<£:y. These .h~ l\j\'d~ of h,~!og(,'i oxides win. gh



r ,~






"..... '.





fllOti,t i Hlpcl!'!mlt o';I;j(Jlrmuor D;'vIS, OM\OlUFld co;~Ull )\:ni:alft;:lie~. A!lIu!li¢ 13:1' i~ .d~ hi~hl~' rooctivc ~o\",".)[[.'.0, (X.'\!t1~in.or!l1nlic compol,lnd~ (21) and ~I,,,_ mlIl~iullll.g:at 10'1,\1Ii:mlqpc:t:li.~uR;'S I,);n, The Jr~nl,w!l:I: of U!il ~hQili~~be i"l,IlilJJ'H:ir ,cUihmlcudb,y the:mmfliie iudilll: !i1131 mml. (!l!I:iIf>~. in steady smli:' will] dte obscir\'ed 1>0'(23). A I1l1a], IIDifll 110·oonskh..'!is!he source of ·

(:h~~gc~h(;. o",idiicing c1~p',cilj' of UlI; l\Jlwr.:;~ic IOol,lnrl,uyI~ycl' l~!l'i]jC'.r.L.'llllJinigdi,~ NO:l 0 ~[~O (c"g.,\! is! .10 + NO --+ NOl + I) ~:r1d, i:' dc-CIiL'!i:;,jiiig I nl1c.~lo,jlI'O] I r.:lJ~iolc.g., villi rn 102 + 10 --+ HOI + ()!:i:: IMJ[:Sl~~~tiel~l~y li~de~ek!S: loss ~!ilf~ro.~1:1, or phol;ci~ysi::; to ou). The rOI11~mionQir~s~phil:le !~rONOl mlti !:ONOl, .fallki'\,\IOO'b )! fast deJID$i~i{)1'II fO the .~lo\VI}'l,ck or acro;ro~.wi n n."I1ID VC NO~mld accc~crou~lh'!.: m:~l\\miOH or b~omidc ould ,ell1~orUdi!! ~h;unsc;~·s.;dl ~<;liJ!rl~o~ (20).. The ml.c DMS OXki01~ inn by n!'I)' ,,~'ill i.! OlflotdllJF ot !lmg;lli~lIId(: b ijil.~i[ ~I~I~ !h;lllJ.)~ Ol:~ .~\:~nlg~~~ni}j,il1:!:l n~'(l!S o;L'! .... (II' 4 PtJl<lntl 0.01 !}~ll.e"rec~iv~I~. nmk~ngBrO dl(: r




ladltilC', Uigbo('ll'lCCulm!im~ of pil!i't~p~,mkwll. whiclh colonize~h~ m~dC'lNidc:or SC"d ice, cxti.,;·uld 0\'01' T:Uf!g)c "R~ .(If ~hC' v\rC1:idC'~1 Sc<!. ~24, 25).
P.hy~op]!rulkton pl'oClducc i:OOOCill.'1nOIl>SSlIl!C'11 as OI2;I~ illt(](l12[B:r llfDd prooob~y [1. These pb~ !o1i!bilc species (oo!lli~)~'dwi,IJ] tiro rnueh k... ss I'll;ll!~O[,-dlil~C'~~iggcl~jc bm'1lI1OC[lri'loI1S) ~hc~~~mv~dc ~ ~. ,"<ml~ (if ~Ilomtin!c LnduIlCi:J]~hcboiIJn:dil!:Y In)l(~r.. 111(: i~li kveit:;;oHO l:l1lhc b()Ulndm:'1'llII~t 1h ml.Ulst th(~nbc sustained .eil1 lrr l!hroU.~l [1"lcp;lioto.llysli> ali the ~~!5he,!od~l1c ol!i:ldcs le.g.., {)[(), ~:l'O].bOil, and IJO..,). wtlkhr l~biJel)·rol]ll Ilrotn]O


1M oa: 1.2 1620 Til iifh'~'in GMT (11I0~lr5)



-00 -dO

20 IiiO' ko;rn9~h,tde (De grees)



(lJ [hii1! IBrQ mixililg: fall;!) aJflld S(l~,a>r im~ldiiali:lI':~ (iill)t;'!lt!Olil a:$ iii! F1ig~2t {A~A :spriUlg: (20 OW)~@1'2004.) with,~h@ n~~he~t If!lIh::illlg f~lli~ .Df2'O.2. ± '1.1 ppt (Qr ttl!! (;1mpali'~IlI. Iha alir 1mi31~ orig]na~e:di fmlll tne OC:El!!!.IiI. 8), A dlilly~n spring (10 Octobew21[).()4) wUh a mnfinenll:<lil ( aiir mailS. ~c:) ,r;., dlay ]Ii1IS1I11il1m6rf1l [l'&ember2nOI~} witin an oC:flamlic ak mass. (Il'l' A dalyinlli'l!'ilil~r 00 lune 2004)' whIlI1IB:r!il Win mOlt detected aDiIJ·W the detec'liitlnl Umit ilJf "bout ,2 P'pt,
F1iig. 11;.D~u~r:!·al\lC'lri~tn(lill

!fl6). or the ~.cc:yelingor t11e.iSC sjX"Ck:sdmo:lIgh aerosnls or tile S~lO\\IIMLi(k The llpuakc of ~eties stIch a" :~ on Imliidei'kh sul'f'nces willi <llso I(H nigger lh.rc:rele.zisc 0 r bremiae iln. LlDe I~nnof U3r. S~]ch.J rol~pHIl£ lI111y explain the strikin,gJ1y SlH1F


.• ,--

10 A\!\9Jag.e IBlrO A\!ie'i;ac9@ ~O



I::iig.,4.. AriJlual vafta,fi:(!i'1 QI~' the hat@Qeli'l m:l1dE:-$measur~d at. H@U,ey St.:l!:ioo, 'P.1~ :!;~da,!!, oo1Ni;1;J ~fag~, ~f [IrQ and ~O aJ'e imditlmdl i:nj' ,ill d~snedl ~r'l1! and a ~lilill Hill!!, trf:!Sill!w!;I!!lY. 100 ~ailr!Uilil'lill 10and B~ irl'lfu(l['I)!l rattos ,(!WilO-OO m Sp:mllgl (OO~bi!fi. wh~f@~~ WluflJl!;j w1l11t@r {May 00, AUlgL~t'I ~il~ riild~c~ls were mn~5,tenHy lJeilowtli!e detecliell lirllitt A ~({Irnd ~mi!lJi>,r Ipaak 1111 ~he anrtillai~ ~a!ag!;!i!"Ilmmflig ta~o. cyde maVall$o be- P~~riit during <llJIrnmll1l ~Marth to April]>.





Mcmlll of ¥e~lir(~004~!5)

~'!i!!riattmls gf 1,0 ;md 13rO (Fig. The Qb.s~rv<i!,ti[}m;; QJ'rg(lmbl~IOml" B!tQ ~cvl;t'lin ~Q:!wi~io1'!:!t;l~a if m;l~~~-S i'110i~~ U~a ]~;ll~(I~~:jJ~ ~J;:~i\f"d[iol.l! Il~i!imit~ i~ to ~mli!'ikll '"~~ in;+ SL"l~




lind fm!Si 1l~)"!I"t:r Ir..gnll~~llj;l;; ~,Hi(-d ~",'id!sea !l"~[win .~ w~nd4"1Om~ filJm lhe ice frol'!~ itnlp~]~ ~nltrirlf CQ~!'~ill~'~l.md ~r.cpoo~,~un I:h~ ~'!iO\W'~lilItk: S!lIb·~~llli!,;l~t. li.ttemg~li(jO!,:l:' 1i,\,'iJ:;~


~'. Illl. CIil~II>o!l\J. [. ~~od:.III.IO, Andre!il~. S. G. Warnm. M\lill.rl~ 3~6, 1655 ~1'1l8-7'l ~. s, it llttd~rgi!\I·"~., f.nWil1!.M. Sd, rll.ldwD~. ~~, li:l!.'I~ (2!OO!2t 5. C, (I, O"~ ~' ~L.Ngnr~'U7, 6iJ2 ~oo;a:j,. ~. IRfl~fth f~ool: t(jdn~ ~Inlll mi1rtn~ ,a,efOliol'l,.fI:. '1100 Gtiliiah'!:, €d .• &1 ~r~. ()\J~,rn; 2: ~nll). 4)~ 2:<II3~331

20, It :sandl~,. 'iI. Rudich. R. ~i:l1I GlisiM. P. J. OrWell, ~hfl" ~ ~~, .!1~ ~3,$7 ,~],'1l~ !1~. 21. it T.l~n ~j 61., .tGlwph~ t!,~ ij~. 253il,5 (i~i}.1) .. .22. t'JoI.i. IE. GoOl:hilf .. l. M. C. lltI,n~.iii. ·Stw. En~JfjM. Sll, T~ 38"l'nl.! ~Q~~ .• .2~. ~.. G.. Clll1.!ilrt .s. 'Ei. U;ndb~rg. iUlllilJ. &lv,lh,IJ·.~8; Sl.O!i




~il:lits ~",,~III.helfl.l'CC)'el~ p~aC!IDI~lbukn!llo,gt:n." liOnh~ ~a."p'ha.'\It. 11. ·il.I~~t:rcfOG"l': \!e'i:)' l:ikc:l~ 'th~.t .10 is ID> w.i:dI!:;prcnd ~roLiili:l coosla~. Anl;'li!l:'e:lk:.[i. as ~!lICml~
~~1e<11S11.1rel'l1iCI]~shm'!.' 'Le!,

be lile ease

Our resuh~hld~eme~llm~l!!sc ~aigh!md sIJS~ !3~l1ed Icvc.1hi of !mlas-en ,oo;kk1S snmllild hlll'llClI !'i,~ofmmd illlJ~3JCl on Ihe theln~~try of Ihe An~:l!'C1 ic ool.!!l.dal)' hI.yct. Fun:~I~cr obgcY\'.mi{ll!;,'l, Ot!ld IM,ol,uto!)' ~\ror+;.(~~ n!l},.;,,~~>,d l'{lllll~fmlh~ ro p~~~ !~l!.IJt k,,'1dCo Ibf:ll1!illioJl of l>lIth .~~ hl!fl~~i!L!I'l:oon {Irrcm:~ive iodino ,lI1d I.'l:rouniu!~and ~o '~'SI,~bili~1l dIe (wif;.md~ inl]w.)llCC·of ~1W: Ch~~ll!li:\l!)" (H~'h~A:rmm::!ijC'Dnp¢l~piU.. n). ~

ror BrO 't Un.

II<. Krelllll, F" 'ii, ]llIi1m5lalJl, S, W. Wood.. 8. NtlIdi\ U, fliU~~ (i~y$. ~. !el't. N, ]O,'l ~'ll"'j')'. 9. !J.Fnl!'lll., '1'. W:!l!ln!lf, I. I?umd~. K. 'Pf!liil5lld~fJr,UI.. ftiMlt. ~~. ~, te,fJ'. O!iEl;.l9U ~2001!l'. 11.0. ]I., 1H'~I~del ,tt ~1.AdIO, S,m'~ Rfs. ~, ~ {~~~. ]'ll. G. IIiIiMln!!ll!r. 1iI,.I!i?I!iI, O.SI!ba~~n, 11. "alb, ~ft,.
lQ'. :11].


7. S. Smlllmi~"I,. N~,rl! 3~1, 3'1 U95\i]}.

25. l.

ID It!: ~il~~.

Th~!t!~, G;. 5" !)ii"",J~nm, S~ l{f;;.~ {Jl'III ~


OAooro.. 00~~. 2



ffl!!!!Ir,~. iIIl It Ab1X\iln.


ifes:. lOS, 28::ltl!9 Cl!~. ~ 2W

(2:000) ..

26. !l '1'1, !!ifIJf!d~r'~,.],M.
(005). .27. p, Spil!lt

e. POO~. ff:lw,w,

~!I!lt:~1. 1!G41U (:lM4)'. i!.!L Wmll:.~m>WIo!',t ~~, r!.\tlrl!' ;,198, ~,H U9':i!t A, &irr.:lil;. I.,. A. a:affi~, J, fJ~~, ReJ:.1I0'l, .lIS')!!


III IR.mijp, ~. w, W~lff. 5, ~l1:iD; t. ~!i~~. ':I!.Oi7, .M;B~m..1.02\lf.200l~OOO~~9.l t[;!oo::!). 115. J. M. C. P!li!rne. A. S,il.i_1·lqpl'l, il MaJ'yfl(.~ ~~lirJ.fq,mJtI[ ~pllmc mtrl~LirtmMi', 10.IE.H'ea~~. dl, tlll~di),¥em, ~ CMilm:I\2006), clI. ~.. l!6. M. !i!ii!Jimilf!~" UL f'l~tt Gtop~. Re:.. '!!9,. Z5,~!I'Il'
]!I" 1\;.


i.(. GoI'll1l!NIIaUlill, .I".·BG'~ J. hJof«fJMl .. ~ MalrJ.~;m. A 1.76,. !i0 ~OOS". .~8, O. i( tFll!il!dlm~nli1,~ 1'. eulfl'lW!l!,~.Ort'>llal A A\lOrDlh~, MoMl~;m, A 1!i1. ~2i (;aoo~. ,9. ]1l'is iiIU,p,pDl'led by ~!'!~ UK IMiltUr.a1 :El!J~rtJI!rnMI II!~O!~l5rn C~ngl"s Mtarcti~ F~~d5n~ IIIi'ia.l~, W~ a~~ grli1l!!M. 10 '111 Uni\oM~I!f~llHd~ ~ 'f,1:W ~ ~~i1lh 1t'llcWlSlip 1)li~s..IL.~im!I <I MNr·m nwdw~llIip !(!:o.5.M.}.

we"". ~



Maltmls .aDd Mifi,1Jiad~. :Fc:ll!lli.. sa ~ S<1:

5!1!1PPl;l:!1ting! Ol1lilfllli~M .. t~\~.d

111. 18. 1JA\il~OIil !!I' ol,}. .•
~f,~J1el!l;;e~ OII11J~ r!J'OI~~::; L II.. A.IB,m'f@..]. ilO!Il.!:ffl~tiln" It(. Sdm~lI,i'.~. llWiH(, C It A. fiiillilT1lm~n, Nrmrr~3!4. 138 2'. F,l.~!l Glall!1W, P. l Ci:UI'il:eo, r,~t;l'I' on (iie«hfmisir,:i' IL tC]IIf,etjai'l. H. 0, !'I:DI~nd,.:d:s. (lli~~r, AA1~!llalP~ I


11«. :[,01, .22li,S,



{li:5I:i!6t :Jis.. M. It ~l'It~" p. A.lIlI1fijlp~~~. 11,BiJ,~~df!\~. A. a:. ~ihWlk~ril, S. 'S:IifDI'I1:OOI, J. !fry:J. Chm. A :rot '55.26 f1.'Il!m. 19" Ii',. S,..Al'u!l~,. 8!,l~l'UI'a.if".ttl,.w .NI~~r.]).Oi1, ~.~.;I

fabll!5<l IRd~refJ~~



·4. r;JJl.U~64.

(,2'0013) •

16 1F~~mfZOOil::XO~~ .~~ Ma:i' 1O,112(i;!'~i~~~~. u.~ J..¢OIl


lntre- and IllntslrmiOlls!cUlIB"rB;and
Diislparsiio'ln in an Organiic: Cry:stal
l(h,>'Iierm~Iled~hlrou'g h il mllglt'-resot'l!le~1plhlo~oennissinn s.peclIl'OSmlP~· (ARIIfIES).S~tmi~i3w sh~ dif:es ef ulElljlllgated IOrg,mkrrm'le(uILe:s orin~e:"estit;l optQe:l.ectrlJnit:s ,He oien hampered by dif!ii!:UllUel~illl '9w'mng weU"'IIlrder'Cd ~l'Idl weU,o.rfell'Qted 't:ry!>t;:l~s.or ~ilirn:5" We hai'll\e grow .. "ry~tall:inerUrf!ls (If

c~cnronic band
sus UbOmC!UUIn

!\~[UCll!llrc,. do.x100H



E{k),or t'he conjugm.tcd :rt systc.Il.l cciilllL'SOOn:h dtc elcctrenie j}ropCfilics rnK! dlC
IJm~'lCr!l\'~or 11K: ~l1U'cd ofll,n.J]ic sentis

(om~lI~~o~. m~m;: we rocpOI!1.IL1~c¢;1CClfOblk:IItiI'C



The niSi ill (IIjf5t8iL~ilnny O~RI1\a lIIyin,o:r Siillnic m'Clterilall5 aUmlis their ban Iii slmlcu~~es tD be

or $icxiphOJly~ 1(6P}.1I mdli1!.'!c :iittlrlr'~\lk \,VD 111, siN P~IC]'i~j.l: riil1g.<;;Ullh'idlogCLhc,r in IheP'!RliPffiililOl~.obI3:i1'1t'tl wi •. ! A[,[PES :fruiJIl !I (20.3) !
Crio:!l1JOOcrystlflllliuu;: IHIlIDm!d cemparc the n::sLlI~

t! II'Qf~_xJi~lly (lri~liUe:d se:d,ph@liIryllirw~e()ldl?s; arid obtClllfled ARf'ES da~ct SlJPpo~td 'by de:rlsil¥"r'UfL<:li!On~L !CC1kiJl.aliom, \7ie' si1lllw!ih[ltil'l the dliredKln p.<Ilr<lil~etto lhe pnm;ipal mo!ecutllraxi5, <II'qU'<lslkm.e· td~rnemiO!n;:l~, band slH'll!:tUfe O!r!ll system off 107ieI!l-dellll!lld~~Ili~eslH dewlop5 @ul. of ~·I'II;mtf.dlllil~. IIJiIdte:\Cubu IC!r:bi~ ts., 11'1"@IIl~roi!st.perp'endi(uhllrl;Q.th~ IIJiIQt~lJIles, the band ~1!r1.1lctYre'relne,~~he peri'iod'iidly oifthe IiliIdt!!'Gular 'c:ry5t~t tind '~oif!H!lIi!WtS bl1nds. wi~h ,(JI lalrg;~Id~sper:li~lTIlI'I'~fll ob5~l¥ed.

witl~ >BIlb inil~!O c<llC'lIlmiol15> ~8). This ortema~ocm urn be SI\')!wn on l,:','t.'1lldyintemC'rltlg ani'IDu'OI)IC
SUh~l'lntC$;. )ilOC~ ;11.", Cut II O}'p{2iX l '0 or TIO~.~IO}.

!lLSQingk;-I:I;.'7i~~vcd 'mk~lI.tt


he bulk b.U'!d :>tIllJ;;l~lt'CSof~~l~nyI;;ry~;d~ Ilnc inorwm~c: ~n.J~~ri'll., Imvc I»;;n d~~'"
,vi ~h ~n~llmdl-; :iiLl~:h

. !.I~in(;d c:x.pt;.\I.'lnm(,mt:ll!y

threJ>: ~;l~ic fl~'''t*.il'~l~ :'im,~11 ~,~ ~imlt:!i' TIU:'k sign!,d, dl~S!;l l'nill,~,,~L\l ~ltJ~ Idghi)' c{H~duCli",.e OlIl)c "l]cll!hW!.~ dlmgillg:, 'Illd t&11I; (ni~ll~uiil~n; '~\l'<1Ii I;:iibk~
;}re~i~njtcdIQo I,heir cd(''fI'\!IJgcpJ~m':ll N ~ltlCro~l:\ IlHCII~pt:-; W 1lj.cfIS'l,n'C 1:'mm1 ~'11'!lJICI.~rc... n iu sill! O

whk'h lmia:xil1Ln~ a~ilg.n ihe mo~,C(iglcs pornJlcl '~O ~he <I.toml,c com~gotroill of the :wlloormcc (~-Lt). Ute Mnd ,C!f~l:Simls were: um:.rnf>u~ l)am,lIc:i: ~o Ihcoof'lS IW(l:I~cl!~ar ",,',ds, ;;:mc] in Mel db~tr~ns perpculclicl:!'~,~r OCllhe molecule. In the drJ:QC~km 1"~m~le~. 'i~~~n(ll.~~d~. ~b~~r'I'ed d~J;.~IliO lhc
siou, ~h;~I.~'" th~ "m.''[~y.~IU:1,,'lJ~~ r ~'Qbt~d or' o 1'!~:I'll.. f,;'Oll~~!ihu[.il1!il;1. b'UlI(J,. i~~ dO'[~irminf.,d by the d~.speU1:i:iolli ptOv~des
,:;O-CflUC:(!,nRr.lmul!;.C'u~'1:r i ]C'.\[oook cl'lmirlplc ur ~h~ ~oinl1atio:U1 olf d'K: 00 IlJd struotUlreI~nl d~,screte ol'ihiw Is of <I ",lllru:i -!()f1,Nl.itnncll skl<lla I system 0 f '1'\1ell-d:c~illcd I],dte s,~zc. 1,\:rpendiclIl'm' to the Ino:I~Cliih!:;' IMg ~txi:;,. contim.nOll:> himtlsacnootill£ the c.tys,j;,,~ hlilicc periodJtc:~ly \:vcre observed. 111is al1ri.'lIOiropy. tGl;;"ctltc. w.~lh 11K.'co'nil:ib:tll,talll ~m~ various1t orf!.il'll~ whh di m:rcnt in~ern lile Ualo:ii!cILIIII<l1' ovcl~ap. is disoussoo if! mg~lt or Ill.mel ~.!'l.!cll!l:e C<!IIc,ua!tions {t 4. 15). ! The 1001 A!hick mm!'i. ii.~\'C~lig!!u.:-d consi:s~~d of ~clt!'lll,l1l,,1.! ...r 6p(20-3} f;:ry:::.t,d~i~i;:s



ulwl r;


~pec~ro.'>Co,y (i\R.lJES}. illl pari. oocaLll'ie ~ingk~ c-ryS!Il:i1:;; with il~mm,l IUI~!d~S11f\C'd OrJ!Cllllm.ioJl tlT;\~ ,


,orgflf~~.c tUnls

hill,>'';;; tlad



;)I,\,;(ii~Jlblc ami 'lVcll..de~1,noo :>ur:[ace tcmlinol"
~ions c>lin!be prqHlred in sil1tl. r:ol'l~'lcir orgmllk c'Ortlut.~~rr:i:i, ihis :;ljtliiiitiOiJIi~ .UOl ihJ,: cflS~ IJo.

S.ii~glit': cry:;ii..'lJI~ of


11!I:H~~l,![te ~~YS!($, S~rfilli'OO' !lr <If'!d h,t~1~~iQl; "'I~'!IJ, ~r~· F,ratllrelil~ I!.Inkier~i~ Gr.n. 00'10 Graz. IlUOlrl\al. '~Ciflair ~f ~rnk~'Uc MOOeHj~ alldl tDesigr!01 Ma~tals; U!'l~~r5i1!1!(If
ILl!olmo, 8?00 I$ol!i~ Au~~.

because of inhcl7C.ifUdisorder ~nnhl(':Sc 1111'1']8. nle p:mpcn.~il~' or the mO~O::lllC'S to cl)fflmm~ lenas t~) mllhi~Hdl.y o:f clY~;nm~eoriclli.fllioo::. alld r~ m!JI·lflha\1.ogk;;1JI1~e.,;;~.w:e<ll core l..lnlk':"iil. T!!Ie challellge' Ik:s il~ gro\li ..ing lia~n r.lms OllOOltd'i.lC~" ing :Sllh.~IU3teS wilh a. s;iJ1glcllIl1ltqlLlC C'I)'S~mle
GJ'iCI'II;IIJ:o:n (i 7).

~c,ijVt; i;~at)~nl"


1'10 ~1fIo1'l1 mnes;pondelK!e sh!MJIdlbe· allld~ellli1d E-rnillt r·Urn .raut (GJ;:J; mkhae:LraOO@'lJtrrlHlr~,gt

~~I.h~~~11~~I!II!M,J~ed! ~ql1!'~l!)l WII\!~ '\'i!!l'llk.


~l~n~iI~l:!s 01' co~j~~'L1(.'.d oL'gi.1.nic cnt~ril'lgd~~ lmxkclplm;c ,l~ 1Ji!~

~.!id~nlWlidiiiM (j·r I!1:IL~irclcct«:ll1if;: :>:tnl:CWn:. C[-IJjCiiillOi:lk1r li.iJi(:'~iOll,.~hu:li,ing. The i





It-l00 n~l by 500 .IUfil). \vhich ruli'i1plc~~ly ~i!od nh~p (2:x ~)oxYgGlj)-ru~Oi1!i.I.rueiccl Cu( I ~()) su'b-


The sdlel~~u~!; ff"ig, tA sllQW~,~ ,(\:r{2~j:) o Qrimood cl)'sm~.~iw ,,\:ijlil~the ro!cvrun dim(i)lliOt~m;

the Qroiml:":. Le., ~Iw

\V Wlh:s. ofl'ne



fC!'~hc lir,;;l 13i~'ij'l n;>;n;nc OO~IK~!I')'. IIlOOllllF<!:SI. ~(I\!i

!l!1.~di~iI,,'1;!~I""" indi~\tl;ldlllc(4a·3) pl[l!!g;$.P'lwi~!cl [~~hli;: :S;i!I~rn~l,);, h,IW,:: ;1Pl,;~'nl"lll~~mi~ ~~1]1 54,.6 A by I~,l A IIl1d '~l~i~i -(;Dfi!!I.~i;,lX~9~ty uri·


en,C>dIn:I~~l!~ win)&::,Lfm'll;~ic1J'I1~!!l~ <!XC' <!:t ~5g to !he s~~~r..l!.C. A ~c<nlni"~ tWl1tnclin,!ll, rI.'iiem~n:r~~1 (SlM}~imUigf=or~he 6"'~¥l.noJ~ycr, C!l~ \."':.~I~ca'i I~ e~!l\IJaI.nil~"" t ~\'I, ~\Ii~j [liD!.'. 1\\10d~nt'io..,'I1SiOJ'EI1 (.lIT:Il! i.mil. oc~~, ii"t1Lcat1XL ~~ slKMi.'oI1t rn Pig,.Ul The rI'Il~l:IlI:I.~inolc .~.diii\,~lon ai'C' elesepacl:.cd linCQ~lJ!lm~s, '\\'~lhlhcir till.rulf!gic iilteliil'lFil~D'I,!;, bcl,weciln cmmn~~s. nll:IC eli)<'Slallill!.rcrpcn(Bkllll~r ~


d~!~(,,'nni~'W(j b), wh(l i.nwr-ril!1g «jo\'>i;'lrl"p. TJile !Ji[ln~ {)\1I!\fJd ~J! !h~ ~~.m~~imH!oJ~of !W,\;:U11; ~ or~iw~s wiill !lw"t \,~i!illl~ nt~~l~ ~.il~k.il1!g qlrbt;lll 1I~')llfi",t,1OJ1iOilitDII!:!:l t~~C:illru~ Ungh,(,~i. <llIlii<pf i l;IifiP1dillg and ~i(;~h~ ll;l\\IC~ ixmdia.lg orbi!:Z!l~,

~~. = O. the grM~ of!~~ h;lc1!!li~oo!ld k ..]: is If:!o:ri~lI'!:Y, th~ Wim!!will i,~ov<iL,~i!l~rlcr:gy ~~d ~in~nI.he;: rlPOi~:I~ ]~ du; ~hCl~'it'lt~, ii'm.~,a:r:t;Jitlll:;:
ru. the
'~X'~1'iLl1l.1tc:;. ,y(

hms !h(l hi~I:'i~~OVCf1~1l b~~'\\;\~('I]bc!'!7.(;l!lcriinlg:s. ii.nd ~)~~['~ ~J.iiJ\;;~d,,~d \.v~l~bt; 'L::.. ~k'd dil: dt;:loea~i~~dbal:'i[t
r!'i.{;, oai1c['

I;)rb~I;]~::o: be rOUlidin .ijg. S U: These have com bI..'C'.ri c~~k:~ I",'~~ by I:init prirl~ip.li:;,>l dOfl:1'iily fU!1Dlio:r'i;.j t~~~'!Ory~~lrnddi l1~rrm~'it",.\rp;.1t~~lio'~i~ fitl'Jil



d,.~pi:~~~d (fl~:I:




has l(j!\v~riI1Icl"~ringoverlap, :md 'OO~SI."Qlh.:a'll:ly
the slx o:rb~u~h" 3r.isi~ng~;iIQ1nii. in 6:P h.~\·e
!O\'l,'Cf 'J!~IC~'ID'd ispersieu.


~mm ~o~he $!Irfilcc (= diro::~im~~'.the molecules <l~' ila'\~alt'Cnlmin~ dh direclioJIS, kadinS 1.(1' <l of imcNi~l~ iJ'I1'Cr~O\pi~~mw~,~II)' dCi}eLuicn! ,i,;ill dl'C'tl!o:n CIi~lissT(mmlg!c {a" in Ihe direclion <l~ml~ ~crfooici'ly ef 7."'1 A or lwic¢ Ihe im(j~l'llll1<!r lthe raalecules ~oT:$IDnal angie. an I, ~h~ '~l!1ot;.'!'g)' !hc I1w~ool!brm..is, is ~hO\v'~ in fj:;:. 2A. The d.i!Sll(:F,;iOin/lil' Ih~~lie bmad~~\im 00 ~O\II\C'Sl i:nlli(! oroJnd ~mi."'1:~on,~ ~xhibit diil'!itlClt mw... ~n01OIl ,cl~ ~ ..• ditfi!'~l~~.
S~ .....pi!0.!lyJ, ~{lU~;[';,,"'" 01' .~.~~ pllC'IIyl i

c~~ersy "blon'bond~l'I~r orbi~,i11I:: \1i.'iiU be refc,r.roC] 10 ,:!i;<;: li~e IOCOllizcd l;,md. The dcyrec



a ~ibl'i];'i~i~liclitii;;.11t t1\ni~bhilll;tjoP1 of l'k:1if,(t'llC or~ C himllo beesuse of th~ lii1li!c ~in1elfl!iim1of Ihe clmi~l. willh t&~eeleceen dCIfI:.ily t~!l.iliijM O:fU'towiiml Ih~ Ciliul:;:eif the n1O~reldlc; A ~e! of valence h~UI~plIDI}('iCnaiss.io!tl ~!lOclm lhc :n' o:roi~h;.. memlilllL'd asn ~lumi~Jnof


,"Ill iIllIL!'f"~1n,~ sl'm::in,~o.i'

.. 4.35

A. In

r~l1ig~ the

ItwiL llC,u giL" i!/lll ~L'le!illDke 1I~C'S. ...

(.[\~·I~EI"'alu,",".OCC~~i!II~ clill~n,;Ul~;d~Gon.·O!I~:!!jl.~_\l

II)llll:!lC. S!t~~r~c: i!1:du<J:nc~orl.]~1IJ' h)ldm'~.n h O'LlOn..<.; ,or :1f!(l~gJloorvn,g ri'~.g;" 1~.d.1;;!!O OB. ~onii{ju~1 ~Iligl~· of -3QPhL,nw!l.:Cill u clj~C(lI1JI. phcny.1 Unil&. SUi;.Icrulltrrll :Sludk'l> lfl~H~tlolleh,llr;k'(! timt. tti!.is l.jW~Sl iu~. d'LC l'BiIclhlC'lIk: isrel1~(w!..~ on, pfLCkin!il ilil [be ,erysull. :m,d [l~.c lI:ing.,:;;bctOonne: cCf'.pbmu

11 Ie b,~I!1,1 $1I11C'~1II!fiC'. I' VC~U$' Ji;.• of an l!'l& lI{k ~m~ld.6>P molocull::' i'l ]lhl):!r.Jrtcd 1'iCh:Cfllmqil;..Hlyin Fig;. ~ C.lnUw rmmncr :i IrDi'filmm! (19). ~..ewi !'k~ng pJ.'i(lC'i>;;i.Ol] :ar (;~~ir~[y. !6' orn~I<l~~re l Hi~ p hest ~tc('!ptUi::.:~it~(~ US ;), 1[fI~Jt co:~n]}h):lI:ion of

~ml111.~ dirr"rcnl
[0 !'IUlmll.:!]U~ , m





~h!:.:1t b.'UW~liltPof ~1!i:iiphe'nyll 16l·,1(o,:')1.~ Qbwi!1td (fij~. 1: B). TIl~ (fci!l.. i~)1of :.;t.-uk'*>.d~rl,~:I,';'d i!1II(l~Cjtuil~g Ul!l.: d:n:li:iIo.v(l( kxo [rom ~~ inc~lldcd o:r~ tilt::· side, O'ITIdd~!l fimil:ilh. of [he



06). The e lccuun ie


of 6P 'C'aU be L'!!!::,:'>I o:meqlUJ\l:1:izcd by cOllsid"''iJ'i ~g the l'I1IQ~,1:.'CIl! as <L ch!li:l~ Or~;U.1I17J:'l1ie ~c rings. each o)1,n'~I'ihmi ~~g lWO dc,g,cllern~e :n: Oroi[lI~i1 On buildi~!:!,l~hcl2 .. :r:I o.bil!aL~ of'lll~ ScxhTI.c:r. Ihc~.""'o dcgcm:rnl:e ~ ()rbi~3Iis IToI'm in!:Cr-rilligbomliing~~d aillibondi n~geembma~im~:s n. MJd so two :It h:lI'l'dS, ,co~sis~ing of U ,~i", mbi!jJ[:;; c<!:d~, .,I!:1\le_ The ~ntrg}' S~:C;I(I {Jf

or 1:11..::Upp'!..'r.'


~CIOCI1~.b!cd band r~re ~~hcl~(j. sls:rlh~g \\\'i dll ·It Ihe: Ho,M:O .. The cmi5l~ioni!n~'iI'L,>ilies oJ d1.C·moh.'!Cll~lilr orb1~a:lsiln:' dearly not hml1log,C'uooMsl)~ disuih!illod in k/r> and dis~inlc~ ~mLv..il1im .~1It' o'b.s~n"-~. U' I,he Cl)iSI:IIT~m:Ii)\."'iooit:i[}I o:r S4llo Ain .!his di!t'!l::I~,n Wt.>;Jl<! dOlnin<!lnl. the llrvllouin ~11i;;' lj,\'(lllid L~

\\·h.ere a i,~ :ill~iITi!l!!l~pm::ing M~he bCI1d~c and :zen!C'rolliU!~ o:]"herf!h ril~g.!For [~1Cdclocalizcd Inmd ,m k - 0, qll} .. ~,,~ i~lhe m.osl ami!:aondil1\g OmiHl~ r~hchighest OCCIJ(}~OO molecular O'.L'ili'!lld ULJOM:O~ll\. wbCI'C:Jl5 !:ll. Ie - rr./(J. lilll"~_ !J ]fx". i~the mo~1 oondj!1\!il, OJ1bi~~1I l[]OMO· Iii). llilis ool1d d!Jer:f."iLL~ill ~I~!Y)' ([n11'1 Ir (Ii: = 0 ~

small ,~()r.12 ~:9uc.l~ a small k.--SUlilCCpcriookA-I 1111'\¥Oll!ld vh;:lid :1 111O.re or kss 'OOJU"1!nl IliJlCI~il.Y ~"C'j" d'!:C3' A-I k m~gc fCiOOri.icd (151. \,,1!idl ~ mi!olcv:idClIlI, in~hc d""t.t In ll'iffii~,g~nlCC. Ihe dmt9

~'iig. 1 •. tAo) IRep:re~'li1ltali'm ,of

~ $E(K'iphe"!lV~20·3) I)l'iu[,.ed ( ,crystaL The ~,l'Iacirngi5 (in 11) inl
the lihl"ee' Oll1lh:ogIH'I.;ll di~~(-'

tioniS '~](pt~Fed !lfeindh:atoo. {B) :Si!iMiml!lg~ IClf~h~ mi~ iPh~rr~l ,~Oi't<iIl:aJ)i'el!' h:mpla'u'~ lOin whli~n 6P{2~3)r 'J~tlJL~ ~i~l'~ 'g~O!i\l. The 2 D u nit ul~1 (JIJl.l A by 5,tL6, A)indi(.i3Jt~rI is ~Oilil1pF@Sc)'t2d 11i1UII! Y din~(' lion re!atille tlO~hi!lltt ,of !he {20':3} p~aJn:e'il'! th€ bulk ,61i1'.



{~ :s..::helil1lil1iic:of ~e'n: !baillid $Uluc.!J,ti.re of a !iliSQiat,ed I~lalrlar 6P' iii'iiO~e(il!I~, ;sh\lWiuq~1 a S~· I,!UI!IrnC!i!! 'Dr ~:rbltals defirnng bands d~\I'ed f~1i!'! {he' MO !(i@g~Il!~iUI@ MOMOs IQlf 000Zlene. Immil'e! antWI:mnding!r. m~ iliMis,ill~ l!Ile tl!l;p of e<ldil Imn.d, ~he l\l,ItIl degeneraten: !llmitlll~ I~f bEhFli';ene"fir,om INnklil IDtl,y l(lrigi~a~,!lrreli'\!~U 'e'~~re1S!i;\d,


! ! !

HOMO 'of the 'local i~@:dlband'




-T 0' 0.··.



arc wry ~imi~<lri(l l~mlc;ij,[)oci~! for all i~QI<!!ioo

froU!l ~ll;

in Fig,

Ie. The

k~splij"~c ~;\fidlh:ii of e<1-cI~ ,oibitL'l!I is r~lprQC<l~~}'

six ~bi~"!,,


of !hl\): ,('l~lQc;ibt;'d lmm;l ,,~

n!~!n~nJil, dt)\,\r~~

Btil'~'~I~n um~'~J1t~~f lk", = ())lt1 k~ = ~(J, mild ~lt" olJ~n (..d~-:SplJ(:~ p(;'IJ'i~d,i(:'i~ of t A A-I rcl<l!ll;",s dl [0 [h\;'! in~C'!"·ri!I.g 8p~!)cillg w W]dlil1 tii~ !l1!Q!~'C\I!cof> = 4.35 A. N01~th,u~ (~ I:h~¢ ,tJrbi!<l1~lt:Usi;:inns (in::: lil!lch Slml~l:i ro]]~i~lg iLlp. 1~ilIi If =rdo to 2rJ,{i. ""41tc:&~\i,liU be di.:,;rn",,~{"t! wh\'i :the fotnc'i:er j1jWy:;;iiS, :HlJlw,cvcr. ifl\iJ'i'C lea:h.i r'e:'l lire seen I.hml \!'/\elul:d be CJ(PCCI.!."tI 1« I'he !.lalfld ,o:r 1~le ~1.11Ullir ~nolecule, il~1Jl'iiif.1ular tl~(! ~itlCnsil,Y [[~ooncdI'l!t~ k fOflhc nOMO mud the UOMO-L The

to:b~C:L""'ccli ~)(riooiei!y
2:1l:h~ blllu
m.ltl:l,lllilUCr ~'lC'!'i{lclkillY

rmher 2r,jl(~. i.ud,r;a!i',~g Ih<l!~ !he

in vcc~llrocal '''pace is nol

the :ii}'''~~:f!~,\If!~!~!1I~~l~~Y i I(l!!l~ 'PoQ~y~~;i· rhf,)<l~y.~~!K~ c;:IH,il~.\vo1,ltd n,';S;~,dtifi a lJ!;IIn~1 ~Olt· A~, \\id I a'> d~,,: di;.;i illcil'c'UIl[I'l"5. ,IK~t1(:]iik,dt s.isbil~g of m~~n'U1i~iitc 1:iJ~ljn1Xr (!!I.f mnitii]:;. l~m~.11ft; witli~ these dcl~,,~izctJ .It m'blwll:1, lJ:i~ nQntml~d· infini~dy Si"lfJl iUIk: Tiu.",.~Ji,cFGlJricr~Ra!llStbntl (I r l,i.,1¢h(u'bilal, should !'~n~el k ~f1',L~~~pl\.'<ld in£lt~ "iJlJlttu- .Ul <l!Ul,d~l~'H'(lntim,ll!tlLIS iU.l~(m~i!.y i,l!l (II" 11l ln F.ig" 2" ~loe rL,";SlI,ils of !iI:litf01.lri~T '~mJIt~· b~lq£11Ilu:id. il~ ~l(:[(llifm ;I!DWI!1d .1,25 cV. d1i<ll ( IOiiln of I.~~~;ll: dl':II~!llizIX.l r. umil:ifi,~", ca~. i5,mlli.g,1'iI~lDelo;\I0L~mhe l\\lf) .1iI1il~ibJmn.lt!t;>tote c~lmctl ffir a twisled 6lP mtl.tot!ul!c ilre MI'~rla~d illSP."'CI.~Oil1sl!l~~t" 1.]111i~. 12hi~leeslizcd OOi.'K.Idre>l: on the experi n~1~i~Ii11 bandmep, Jlhe~~ul~itt j iUX'I!\dI 11m Lijl rroi~1 tbe F IXiJnl w~.lh iI t1i.'>I'l'C't'Stoj], w ill~ C'xp'cril~le'jU: i~~ remali1kabh::.\\.il;h lbe [po:>i· or al'lO~u 0.:2 cV:, The omimls or Ihi,~ b1l::~~drure ti,OJl!>,lhe k w~dl.I~. rmd Ihe re:liUi\,!! i~llii~'liIsilics nC<l;, deJ:l~lcmle,oo expected i'f 'ti~c r~mlccu~,e is: bC14'1g lidly ~)c:pllOdu!::ecli, OfIXlI'iIK:I.lI<IJl'h1ote is: (he l,wilj.~OO. lnM~. ~ivcn~hc 1II1iCCliCillJlI)"misill~ Iron1 11l.'!Oci!<c agreeDUR"'u or bOilil poGs:ilitm ami rollnivc the c.'{peri~nC'nlal ~o~utioll.;md 11K':1~1.\(lximil>' in imJ;1lJ'l~~i~$ Inr U~~ HOMO .~~gd[he: I-i.'MOb I , ~~IlIl.~L'g~'Or ll-IOM~). no s~ro~lgOiOOlt;tl.!~iCln win:
b~ dm"!,:~~ [fo!ll~~+
S'uelil W11,!,tllllJlijjl~IQI,I~ ·~.i~O,i[ion O~\"'I~J:I i~llr.~i.-



its «rmliiIJl CMm~

:rurId tiDU!5 dw l'Pni~!:

clect_Hm. such ~~1iC'<lng ~f!k1s b}l c:i\'C'11 ;rn1i;(rJrnic li')otemi:Jil: and <I wc!rW'C;I~ ~~miw,~lQr w(lldd gQh~!~l;' !:h~ i~trL;l1- (i~'(IUJ' OMO PIt r..~~ch .. iw H w i.liilot ~linnd~Ll;:~ in d].l;: fQ~ri~r b'Ul~r(JL<tl,


wij!.l1~nthe lH(}I~etll~e i~~o, !h~ \'Ihi~h ii!!le~~lic-.d~)' w~U ~olvl.'d lrom 1'l,tiK'f ~m: I sp'teil~~ bu[ !h.e ~~m~r $11~<Jei~ljjl, lJ und,.. ~m'l ~!il. S1. ttho (;'NI)cril'vq~n!~1 .i!1I)L,~~~ily~~~I~ wh~cb leads to a folclin~ b.~ck {I'WITII.1'hand, 1'iC1U1.">a[ ~1C' !feSpcclh-ernbi~~ C\ll~rglc!1, are f.il{l\\'11 Th~~, periodic it)' c~un be rn:!io!1a~i,;rocl b.Y~h~ in con~p~uri.von lO dl~ iFourit::r lfll!1... rcmns of [he crl!cliJJm>oo o:rbiic;:B.~ Ibw bo!h '~I'l!raf4!' ~'!cllwii;!~1d tiP. ~lok\L)u~~i"~,,,'i'll:g .~~ V\\'iMc"d cO!lfor~lH~io!f!. m 1i.r.ls~ i.n Ih~ 11!Jm:r.:vurrac{l n:lgi,ml ~)mbtJ.d by J'\..gl)iu., Lll~ b(~!Wr ",n.pl(11111~iOI] ,or 11i~L,l dM:ul:iy AR.PES.. Wr:.:h.a\!t; hj~l'Q{t~l(\c:d tl~~ poss,ibi.~itJl t.\Yi st(ld 06:' is ~1T!!P.h~El.ia"t:l. of such fi lW Lsl!:,j Cion torri'lllJtitlJl for 6f) ilt!ll the A. syrp.riISi~l1g nsp<CCl on:~c: clJnnp,:[d~:m, (lIf [lile !>olid SI3JtC 1111 carl holr work !:rol'l1l liO:IfI IZl1lio:n F our:i.er ~!l<unsthr.UiI.t"'with cClI:pcrim.Cnl islll'!m tlile energy romid'crntiDt~s (2it 2'1),. Fro~11 a bmld OhsCM"<i high imcl1silics hl!:hc second Brillollliin ~~IUlLIc:tIllD'ep:lilr! ,o:t' \ lew,~he d~IOCil~il.Ci[J band ~oru::m~' repmdaced, PllCi.locrnj~sim! ir:ucf;ls Ltics should miller Ibcun~~t)I'C[OO in 1I'::;J1l1)l. or tw\), c:m.oo dctennincd by!hc tr:Jllsiticm nmu'ix \vi~h~llinib;l!1d." nem~d by the (.hlding bOliCk Zlil!l in Femli ",~Gl)~den Rule and o.!regel1.cmll1l' dit~-;' fk!ll!h ~() t<!k:1J!~;!~J;;'.I (1\vc\~~r., nmlhf;lm,uiC'.~ of ~ di!c .:nJ!lihrnlil:u,,,~ bmtd oo1l....;;il'>!i!lg '01'1 IOMO, I ~OMO·I" the .Fou.rier IlIml,>fo.l:1U or lite ollhil.a)S is eqtlhlttlld [rnOIVIO·2: ;uul .~ l:IO~di~lgliJ.;ml]d comlmining alcnnt K'I a call.cltl]mioll Itbr Ihe tl<ulSitiofl. :lillID.tr~x UOMO-9. UOMO-IO. rulid nOMO'"1 t wi~hlhe rro'Ml lhe i!li,t!!IlJ SIlale oibillll W II pJi:l..!lC "\,,,),\1'e flnmJ <iJ u'l!Si BriHmll!1. z.O:l'tC' bomKI'.IJ)i at ;It' ,_d. phillw Ah~m~lgh the In~1l1l1<C1: orn'bitlll8 ~n ;.I! ~;;]jnld !i;1;;)!OO~,if)Jlll~i" <!n~l'l's~ 1'!iII.tll!'1!!lyU~cgk'i';:I:;;, of !i,l,.;n <LillO di~l'h(:l!i(m CiJ~.!xIi!i, (III' [~D~ 'rl\u~.!!.oi!!~ 1)11011:1is d~IJ,,;'Tnl in~d hy ~1ta~ !~LUn~b(;r I[IIm]il~~n&,Ihl;' of
InQ~on(QiIii!l DFt:l.itaJlS -

m,oiocuhu {lwi]uJ, di~;lI:~:~io.nhas been ,d:imcIII,ll t:tJ; ob!<!in.F't'f1: ro.laI!lvciy ~ng OI'lik;y1 ch;;!in:,l. \,\'ilk:h p1~k \1\tIrt,1C'aI.ly ~ll ~:ihn..... ~~ blnd:- f~um floTlT!:.d i.,;mis~uo:!'! (i'_~:pcrim:Ci1i~ h[L'I,oII.: b;;"{!oll jJIOU!l.lcl,. but because of !he l1.o~cs.;!~u·.vlyllfg1i: l1ulln~r CD 1lC',.. h r
P(\fLT WHits,

~he' di£C'relc oli]l~fi:)I,ls e,oiilJd ItO~. be ebserved ~4, 6, 13), Siimiltiilr I'IU\ru;u.I'CmeJl~SO~l :l:1ih~.lS o f 'Uprigh~. 6~P' ilSCl r (J) ootlilcl 111.11, resolvejhcse fC9!liLlre\because, ifl ~lC {\~lOOn ork'.ll~1IlioJl,~h'!

rnok,,(l.lIcs are m:tlll:llm)lm 17° Ih:ml oonl1JI. \iV~ tlC1'1.'II' oo~sjdf..'f' lilc:!ngubr bch:rvi;or ol'lhe [lm~!:I~~ni$f;io~s ]!l Ithc di~(::!iDn 1'M..'!pcndiculltr ~ ~o nllc ~l'l'Ok,'cul[l" ;flL.xis.b~ this dired,ion. d'lC In{l.~COl! Ics JO,U!I:::l1CIl1I.hCI1l .. 'lCIVl"S ru> IlClm u'!I\cly dCIIll'lcly
p:~c1'1Xi;n:s!.:!ck_s.\\~d1 ~l pcriOOici~y 0[5,,6 A (Fig, 1A), dlC closc, ...1 in~ll'.:lipbnIO'lrs,}.[!ci !ng Ioc the moJo

!$iuk:':S illl iI (IP' ~ryslall_ AI'!}' nl~rOI1lT1n~lIln~!to.I,oI\,O'f-







17 '1-6 IKlna~1cErieJIDl' ~'e\I]


F'lig. 2. (A) T~~S€t ~:~ Hie ~I phottlem~ssiOf! s.fl€(~r,,, off '~l1e'!S,P n: banld's rnE!'dl$ur,edi fOlr t~le(;~r(lWl em:isslolll ,,,ngltes from ~5° tOI 70'1' in t~'e pla~!2 tCQliltalil'!~l['Iglhe tDl"1lQI molecu~i1Ir !;Nd~ all'ldme sl)i:rf(l(I~ lilQ~~t (B) Ute set of ph~~(j1eiifl~~i(m ,ti!,;U1!pfes~fl!let! a,s;,~ (ontQ~r ml'lp'. Tihe !l:,(!l(lf s~!l!ll'e:I$ U"'~lr \yith n~'!Il1es~ inlt~rnilll' diadt nH~ enel"'~y MatE! is H'ferr!~dl t~, ~he Fermi ~evE!l t()flihi!! nlils~ra'l'e, wrnfrMs the momt@l'Ilulflil ij::I:) ~ll,ri\i'edl 'ho'lill thle 't'Leictro:n

mQ~te(liJlle ol\1!i!rh,it! OJ1llth~f};p~rimlerll.:lil balld m~I;!T A fi~(~llr1(1!l:ill :laHfm ,~Fe ve(il:or {G) or ~rr.M tri'!~he; th'an 2rJa lis oos.ervm. 1r11~tWOt dltl,~t1ed 51nlll'ooiids on tine leU. glJiide~l1e ~eatler:'$ eye ~o '~1lt!.wo Ifn'illl'ibami!>., t

o1lJilgOO(ill, kK '" Ir2rn'~Iar/lll~/2s~It(!l., ~$ ghl'~1il i,1'I A-~,11l1~ d~l'IsilY ,()~ sta'te~ mO'S)i!! ol1~ne left Mt~arul tD~~ilil!S c'nll]m~ratoo. Th,e FlJluri,~r spe'drari[:~mtlrl~ IDH·~kl)itla'l\~d Q~bhal$ olf alii isr()'lalt~dhvis:t~d6r


iOC<lI'i,mld~lro1.gllmll the (~Pnml,~~!lai cl1'~i;]~. Cf:nis.'l~(l.n<liS [l liJJ-'l'01.~gll (If 1~'hQtml en.crgy I4n~o The direction in wM~~ch Ib>Lr!ddispersionrans l~e 110 '1::1./ 'lll~, SS}i,Uc :1'Cl1~!11.";J1lC q~<lU'!iI!lh>cly sin!i~r w ~I]];.r~P\iX'~iol~t1iS!;;'~I~'iiOO,.,,~~!¢,i~ lI~t , !!l~9 ",..",~'tl~"nwrcl:;in~ ild~T1II!~i!ion, U' ih~ m~~c· Ih¢ ~l!him~i>< f !.he J~~ot;:n~~~cl. l'i!·Jb..:!qld ,IN \"'~'l1k o f<l1~ or ngul!;!;'O~r m~g~,.i1hIJ d~~OI;:fdiAi-d m'bi&l~ls ;q; i;:t~l'c;c1\''\:~:;;~~U;;~I;;-d(1roi;;I;i-QrI. I'l~is b!~~~~ "MI!I~d ~n '!i,,~l:Iii ~i!t~~ il.~tlClk;.iuy\mrillmt~ilrI"n~ ~,rn:itUL1!1c.iUlg dO'ti"'D!rol~lr. 11mi it ~l"lJX,~Ilit;':;, LiP [is art:Si,lh of ~r~W~~. ~ftlld the '" ~al~d ~mi.... ~io:ti:;. ~~~ d011li!pubuwls d isp.lay ;1 sll!l~~n\C1t is~i(l!1 d G im!!~C'dby 1!.h~]oc.!Ii..:.cd bmw lJ1mib{H~dling ,~t· [i~~:hlle !liolrtuhu' plilu:ro inh!ll!\.'illi ill'UlC'~I~n:i!~g· ~~oJloondi.~g QnU:li ¢V. Th~ orn;crv~ld ~p-c:r.iGditiIY is 1),1 A:-' bil~lls (fig. S:UI.ln qmt!<!c'il. tQ Uie A:>r. llll1l,~h~ k~, ~GP1'" ~~:~,cturc d (11~. 54). The llI,."Su~t~ the l'i.ud ;l;i!tLl~~!Ii!'~ cf~leol,l~;JItiPl]:-;. omcl :tcfll.~L~ !.hie ili!!~lJ'ila.ml$ disbUice oij' :u: A .pr cllilt';, ~huw Very Iiuk .hllL\Crhlily \r,q'i:,ajti[]i~wii1i~ un Il'ii:; ruJthc.r ~ja'i 1.6 A ;lilt! i~iii! :igrcc.ritiI~i:ilw:ith IJm'idi ~:ri:gle, and th~ ITWijor d~li1gt: j;~ Dim11~JL:~ilID'1lY lot lh(~ PtY:,~aJI oFp.I!:illliiC M)iiilO~~ll~~ ;!M.itlCI.~lre ,~.lt'li,Iku:iml~ im ahh. t1irt~ iM .. J~t\I~lioJ1 ore o\,len'lil~d ob1.lh~ C'Xpcrit~lCin!l1 lima plli>ltiOi1. :1: the ~loiloon~~lil~g, 0 fealilJlre. The daID are COlllml:l~~dfillll1 ~'Vdl Itlis 3]m~\.VCcl IxYdl, pi\..~'f:I~{I~ii111m ill~blly 111~~ In r-j~.l afler OOrn~ I) r F'i~ . .1.. Ef~e1ilmYI~lIilt1.i (jiruiml 0 r 'tIilC.i Stl~"tI Ihe ~nlrn,.,md in'Iien1~oleeull~r l~ml.dSill1l.CIUre of meleeule gives rise 1i0 l\'ll\O ni:ar.d~ncmte baads vcr..iol1 [m1l1 0 to krlllc only de:1oca~jzcd cib.i:tar!!> of~lhe ~~lO:l.ee'L.llc ean lite lruly ,d~SllUgUiisl~e.d b~calllt>e there nre ul.'o6IPnIlO:lcc!lll~!>rer m~~1 l'I;lm IIi Ilrotctl)'fl:i,ea~ 'It: 'OOlljl~~lIlCd s,y.~I.cllll In be ~III are ] [01\1[0 and I ,IOMO-~,. T,heir ilUCllsIti:C$,are dirocll), dCI)l~nllinc(]. PCI"I]c~tdi(:ul;lrto lite mo.~in d1;C''crys1.ll!L1. The luoo!icllOO diSi~n;iOl~ or (he c(:LU:tC~. tllC observed band diisrersr~n reneets: :spOO'd~1a]mosl unifO:Ii'J'I!~])' ink)I' \\'id~ n ba'!';:!'b~ dc'localuiOl;.'i! r.:~, UOMO and HOMOc.l i:;....Q.1 cV. whicb is (}f Ihe giroc:r (~rho c.'I!.'pc'nmtmai ~ lh~ ~imN(:1!! sp<!clng (I r lh~cr:.1!'i[;d. I nl.eros.li!1!!l~Y. d is~"t"!l'l'!.i.tb!c ,e~~tlrg'Jid ispt!f:'licll1J. Such Ila'lbi.u'!id!l. • o ~l) il~lpl}' 'Ul'''Si,,; oroi!... ls ;n~ local~-d '[('l ~'t~ Im~~ .~\WI !~~4)!l W ,~l]roor!l!t.\il'l~mIUfl: 'lOW C<U'I he .~~ (l~i~ll~ d~~«u ~ij~cdv~ •• h~ nrl,~h:lC'u ,m~ :S(;~!l Hii .~ InG"'ll~~-dl('lnl)'iC rnok'¢:f.jl~~ i,~ ~he t;'t!.ltIJfi. n~'(' orbutals hu\'e li~!il,~ (j\'L;'rlllp be1J... lJUly Ji... ~enK~~lllhL) '(l~perif~'iL.:rru"r-of !li(; C;J:y!H<!I.whcrc'~!I the (iJd1!~tO!!L'l~(lcaliz.ed on [he ~~~ICJ.'1' m{l~cr.l!k'i' iII l!h~ ,crysl'i'L1 <iI:uclare LI!1~ikci~\1 localiii ... band" <lI~1 .od ddining p<lr.ll_lll'CU.'f:fi. Ihm i:-l. m(lll;:cllkl,~ru pcu~odtc il.y.. <lnd siae o r I'hl':l '~lhcny~l: bt!.~I:~I:~ngbloc,];.;:;; or~iw ~o be dilt\i;.'~11I' in\lohr~\d in $~.llkd b..'U~d [r.i!n:;i· ~hc din\Clkllt d~loc"~~:-';'\,lld in~hc l;!)I",,~;(l.1 .wd fO!TIl ,..;utmgly di.;:lJX\r1ii(Ut, 11I(lill1 f:WOd~!ll~(,~n~r..\m \;~'i~h 'L.\.'\p..."fi· ~oll.h1n11!i.l E{:ky) rl'~Up,~10 ()I]~~:.' ehank~ iJ;l~h~ Lmu:d~lnbl:l¢ di!\~("lio~ rmlOlUd '10 A JI.oll'PcJIIldiing tI!bim~~ \r!l'ibly pt;!u:kidi:c in k. It!i(!'Jilt, l lilll r POi"l, dw b~~!II.I~ !!i1q:l~IJI~~m~dily di~~~ing 01~ nits ba:rld .rul~$I,!;,P jmmr m 0 zOllcixlund:l1)' WI &h~r ill] e:DC.I1lY dl;llli I.I~C'C'Jll~ul!lm:~oul. 'TI~~s mile: 1Il.1oh;.'Cuks, th.C' inlCn:i':!f!kci;J~:rr tg\lcui;Jp i$ seen ItO t'DC 1lJ.;gil igi.~l:c.andll1c result U:ct1lb'('ffi, the cliscr(''p~uICY 'beIC~\I(".ClTI'hl:.~ry arid cxper:imcm I -U.l6 A-~.whit'~ implies a rc!i~ S!pacclfllt~~ p'L:riodici~y01' 5.6 A, wh~eh is dre same a. nhe arises pllCSlin:ril3b~y bOO:ILlI!lC~lC' mokruJ'cs :.u;,cin ~amill.iQr:I.or 'dte bmid stmeture >Oili lin is.(Jll'Itod s rri.C1lnO! pbnill' (1.11. A twisl in t~'ae moleclll~cs, moleculsr Spil(ing Qif the b!ll~1k crys~,cl in lfilis IHoicl!Culi:' Olllt of i'lldh'id,lml omiruils,. ln H'l~s leads In [~ooiliapse 1.0 w:o,al!' dll!gcl1.crncy for tlile direcl.io.IfI .. eII$c.l~n.' imbidl.ml orbitals con$n~tlltill'lg the'
1<111.and IlIm~ ,;my cOJl;<;:iQernbl~ lmein1i.QI~cl~lo.!' d.i~p;t!f:'liolll.\""QI;!!d be 1;l?ii~~I.'1CI~d 'l~ in ~li:5. dq~ ru N(;'~i()!!!, ~fII. i;!~l;rh~~~~~n~ll!or! d"l" Q'I~r.~.6(liP


~oc.llil~ od~h:lI,Is. hnelillIDlCiCLl~~bQmi basa disp.i!n>L'01lof i VCr)1 lItn:g~'ftJr a 111m wi~hfl ... We have alsn il'wu,"Ug'II!1;ld!he hmlo '''IJilJctm'c (:O~lPOU1"~l~(:'n. Sue!]! a b~rc ,cli~u:1I'sio.~~r()rpcJlcl~culilr lno!hl;!' ~Iil!rl;!o~ ip~;ro;ll~~,f,(k~JII by I. ['CCo:lrlrngdl.epi~otoonisskm SIl!CI!:Ultn glonnal i hnpJieslJwlI d'lCSC I1IC1l1ioollding omimls ared\::-




bml,d<lI:c~e."ol'!,i.[Jjbolc. nnd il c~n b<c t;OI1i:l~tII,CJlJd ~hm~h:c n~o~,iX:u~e8. I'ztC1i ~dlJrl a IW~ll.!~1d Ul C'O!l{ommlio!l. The rcsllllt'l me n:~ i~~lponmtt for olJYllhic I1l un t~lm fabricmion.whcrc. .fo.r c.v...'ll11l!pk, dlC

C'fTOC1iS on U1C: clccueuic :slmCllM'C 01" S!n_teuJi..l l I'h:<t~c ttr.lIlsi~ion~ or din:i,;re1~~ thin fb.1 :po~Y"
<1 )11;1;1.II .. I

IrnIuph Stn.L'{;I'U~~,. ~O:lnirlloni.l] ti~(; rg:t'illit.'i, arc u Llr: c·xjplrlrt:d. Tilt; dlfllll~l1flei", lO crmlc,tl!l mhe orgfuU1~c'uhill, mrn gmv;,t~l, !is d'ollchore, ~O ~llk;ve~iigh:ly o.rh.~t:m::d mo1cculflr lilms wilh fI s:i,l1gk crysmlli"~ m1C11IulInclJl.
!;i)e·fer'liliilltl~~; ,i!llild N(J~e~; 1. S. N'aroh f/till:, fl'flrs. Rw. 8 Sa, 2'1'2~(199.st 2. "I. Yi!!ll.i!!leo elI7J.,. ~ ~ •. i5i tl!~\ 033IQ;! t2'OO3:~. ~. (. ~!Ii~, ]•.(<10, PI!.. Gflo~~ ]. ~ri~etila5, 'Sr;tf.


4;8~1I5. ~4' (2M]),
.4, D_ Y~!himl:llia E'~ at, filly$- l'i'!lV' ~ '~IlI, t(DJl6


[band ··9


5. K. Gr5~PIi11,~ ",.t.}:. AiIJ'. Cht.m. .s:~~.lI;ZI6.,iISB,9' : U004~. 6. K. ~;,i.I)!j.~r~n, H.I~5J~~'~rd~ II ~o~_,. fil'jy;: ltll!'.. l.Il~JU tl'51ll~l, :;!. S. l'i1lSf!g,l""ilI ,~ !fl.• J. C/tt!rn. Ph,s. j'm~1 6%~'
{l'}'t~~ ,



;8, lilI!lY1ma~~M an

mll:1i1!;~d~ ilYi!it.lI!ll'.!' ~n 'Iii,

9. G. ~l[ie~,tl oJ." M •.~1!1.!>t.r; 16, ,'l!59' ROO4~, IDOl $, 8!!!1tlei1ill'! £t i1l,,~tj.. Sci, l!:'Jr;6!rno, []en 120061.. H. G, ~llt!~ ,N' 01." H(JlIO~~r,r; 6, 1207 (~~ ..

$lilffl(lf' O.Il!~I'!~.




'(16 k'j'l[k~]




B. M. Olllrui~

filg. J. Wile D81ndJlilillP as mea:Sl!lrl1!d perlpmdiimilar lo~ne miO~ec:lJil,u <lIX,es. The mO!r:teli1l~lllm Ky is giw1'l iln A~:l!, ~Ild th~ e)i.Pr,ocl~d !3;ri~~O!J!iini!OiVl~ b(j,~ndalry ~(lr b ;;::;: A (0.5<6 A~1ii)~.lndic~t~d. Whe i)g!l!cl 5,.6 '~trlJJdlllIN?;irl this, dire:ctioli!, ,GlIwul[.:],ted 11.0IfllI the bu~k :s~rlJJdll.ll'ie ~,f6Pis 's.\Jlp~rilil1l p(lsed Of! tht data (Qr th~ :f1ecOliU~ Blrilhl'uin jlJorne. Tile 1heo:r'~ti~il~ r'e:'lult m~~he' '~poer~m'~nil: \w:~I, repua1l11dllgl not only fhe ~'ery i!\i~;\lk ,gi~Ii:J;s:l~!1 1~ the biln'c!:; d~nv~d, rlr~rn th'~ Qrbltii!l~ d~lo(C!Uz'!ld (b~!J1(l Ur'!es) Q¥er~h~ R1Illllet:ule (HOMO d,nd HOIMiCHll), but ~1:5Ill' Stlrilll1£) ,~ispe'rsion Gli[ tlle orbi~ol'~ Lucollizea hlffinte ~]Iil'es) the ,Ili ~ the phenyl. ri l1I!1sil11 the l(Sol,aJ~ed mlli re<cl.l~e.

t~.M. Oeru~~ rJ:tll" .A!dIi" .M'IlIS'. :Ul.. 2A1ii6 {:!OI06J. t'j' al.. ,(i!jMJ:o PIIth GlM]'", :il1i Ilrtz,; . .!I!'iJi!li'I)lt!, anljn~ ~1.1'It!'j):tf.!l~dQi.oJ'g!1O.100.~llht.~:O(!l;O(1:S5? 14_ ~11>!J SlJruCiul'e (;jki~lli!i!!'In5 "Ill' getlie1'.i!lI)I' ~en1>et!jin hl!l!1\ ¥iJli'I1l!lry dirl!{llom~ cj' ~11~ U)'~~~, ~iIlith ar~ nf!~'m~~iilril~ Iili~ imlllllfl:mn: dirm1M~ let ~~I!I1Il'1~.'10 ~~O!fg.e'lr~filp!llil, Gf: Ute ~~enm>e[]I;!11y a~ible


d~roololi1~. ill< IlliI~ ,m!l<l','I~I! (;!Lruli!t~~ 1]iI~~l!~.r~~


[ror iM'

mNWr,~!iIdirec~all1!i~Ji!llli!l ,~!'1d pt~~~dlc!Jlif!C (1999h, lll. K. M. B~j.:t!tBt


[5. P.. I'u~mhnig. C. A:m~I'MdI.[Iri!:d. [!>,!J)':t fYN. 8 ~1l1. 71391
Il;!,,~i!'!! .i'!4,

1571 tt99n


17. III G.

10. SI~iml1UII~ef,1l'I. Scilatmla~~

M_ IKi~l(ijmOl<",F. I!. tl~Lil!iir. Cfrem- .Phyt 1117', 1I4.? ()!:~:!l;o18., ~ ~. M:!i.~J'.D. !W. 'i'urm~. FMM"r n~~'ol'!.I~Mfi ,SM.'. (

54, :l.ol~ U9l.:!!, 19, p,... Htllfmlliloo.. ~

(lnrJ ~~

BmHtfll9. [9 EmnaM Strudlii~


A 'hfl'flifi~ Wtw oj 11I!W'Voll:',


1.9Ut J. i:"'~I1~t;I', '~'il1~~~. ti~~~rr.,." G. R:1!I!I1~~, Mv, :!.I" !" P"
Mot!~l'.15, ]j12~(lOOn,

n e:. WilliHI' ,fl' Qt•• Appl. Pill!!. ,Mf/;

83, 2:511111t~(I,6), 2Z, E. ll..Si!ilite'l'; I.~, 1!e!i11ll'imlllO, S. 5ii!i!oo1" f, ], Hi!'!!pie~ Ftl:fl. R,I!'Ir" :~1, ~61.4 n99~'. 8 1

2~'. 10. 'I1'lII5l1imm;a oe1 e.t. ,(l!jtmi P"~,,~JOI l!OI7~S:l 2S:.'ilhi5: 1j\1L'(~ 'W\j$ ~VJliPIlfloedDJ Ull~ JWslr1a1l Scie!IC~ Furn!l~nwF]o. a'OOll~" 2A. m,~~~ slrutJ'l!urill, ~di~, nil'r~ ~diuili!dl 'fuM ~p i~ ·mHl'l'l!irn!I~ pI~miJr" illl thi! !;i)~~ ~till~. ~U.l" ~ii'!lJllI!rDI!IfIU m SNJp;poJi'tingt CIiI'I~"i7Ie Mate,n'ial I!f! ~~Ol~rw~ ULI!!I~ ~,I(j!liel(!'l;1r 'iU~~~oi!~~, <ll/!'!Ia¥j ,~~d Ill" ,@I~!'I~h!!i.!ln 7I'5B3613S:ltOC:l! Iltit lCl!liUlliool ~all!<rrli~~ (II') ofilll~ '1n'fi!;!iN! (OmfOlIlfl.!tIM ~e!'! 'lIlriimcl\e(!.!Ii:e5 <Ire' on'fnredlpar~tlf~ lolllla' ~wl!m,i!lIe'. :Fif.!5<,51 ~~ ss ti~ ~~,r<!~, f:if!m1 !l~~l!ri~iillt!~~~W~ I'WOO~lJ<ll )Il~~ ~ ,~efgren~~ liightr !'P. 1mbl!p'lI1!l1 il!liiill1:l1 Jil~jj\!lr mmfoU'flilillom, Th1! oo~~\!!ldi twill. ~' be~j~~. 'i'll, !h~ ~~U cf iI ri~ri r5!!!i!l!ce rew!i5tJ~cli(ir) Dr ~e:iiE' Iliale-tlll!!r ()~ :KI. Milrcli! 200i,!;' ~a;~!l'~~5 hm~ :wail 10,,11!2!l1~i~~c~U."12~9 lCl.!tffimgl 'l.oliIl ei11'~JID'. ill!!:


IF',o'lrm,alion ln Naln'D'lrods, Thr,ou,gh IP,artii,al Cati:,on E,xchangl'e
B~yte' SildUer,.'''' '!)'!!I'Ii:s 0 •. Derm:.I1u~@I;iJ~ (aml K~ ErdOlnt!IU~.z.,~ Liin-Wi!liIi!lQIwang,,::! .It Pa~l Alh!'i!ia~ilIl!OI·~t
Rkh: a~!t!l1 Rolb:il'lsol'l/ D.

SponlalnSID'IUS S'Up'Brlll,8'tI'icB

d~ooc-l ~.~ <!!.d.... t'('1~ (9H, OJ~ck!ll)' diffi:rcnce I o.l!lIf lli;;lw(l1.:1] qlli{l!~l~Ull drl~~ 'Li~~!11J,'I!~ grm"~l UI1 .. ,~lIy ~Lb~Iit',,!Iltl:.: $<1 li·~~I.UIl{l.iH~!!.OOUoid;J1 ~lIO)n~tWUl1 dot!i;isihL': pre.'k·Utl(]Or~DX'dill. 111. ";p~a,,.illJly gwwl1 sySli~nll&'the imCl"r~e lliC'l~"ItX'.nlllm suhi\lJ~uc!!JD')tS.ml ,~ild ql:lru~e.U1m 001. ,Creille'--S fI ~~ion 0 rS!tt:1!nill, dtc !ltllmtJg1dh~g ~1b' dOL Ing,c-niollsly. 1;~lis.~OCll~stmin
til<lil> 1!x~ used ltl cre31C ;:alll(olIerg}, ofillllCtaC!![011 1be1.!;~Cml osc.ly ~~t.cctl dQ1S;; tbis !!Ise Or'~lt"Hin d ('1J!i!iilD~";1fI),l;" hes !cd, ill uun, to qUWI1Unll-otiQt ill"~ irJ.y.s tlmt ore SpalI31.~y 11l1t1oC'l'IKl:! .in L'IA'O ',md CV'CI~

LaUi~ntIrsm.U(h:s.t,ai~$ ,flr@ wiid!~:r Ilmm'l.il to oDntro'll!1~fiQS(ale iJlitteftf!l [,Otm\\ll1o,r:l ir:'! li1@ter'Oi@lllta:qr. bl!l:t :~i.i[1iI @ff~.u nCiYe !ll~t OO~1l! l~pIDlfltE'dl ln !)OIlloi~oii!l !llan()crys!:al gl!'l)Jwtl1 •. W@ Id€m(lll!:;t!'!~l@ .111(on~jd~~ Ir!Jl!u!
00 SY,iiU~€S!l!Z;In~1(dlS"Ag;:S Mf!orodiSl!.!p~rlanices til!MlIlgh IP~lfia~ C:ati011'! @);;d1a!1ge'.• Strainl Uir,~IU:Oes ilh~ :sp0f;UalF!OOldSf~:i'1':M~(IInof pefi~dk~U'1I,.lcUJ!lr~5. Ab i!iQi~io i!:al(ul~t~(I!i!5i;)'f~he l!'1Iteff3ld~lIM~rgy' ,~~d moo@lliflQI or ~traill! l~rI~r!lj~s ~hm n'lliltWh~@ fI!lK,@~ I~riy@ Uua sli!'lf~~9'.o1!mzoi!tiiDn :fll'l!1! s~pcrlat:l.i!oos. 1i.I1!lJ O ~anorod s~pe,~latJti(€sil',;(h~bit higlil staibiUty aJg~~[ls:t fi~f1iingt .aJnd lP~ase- mixlngl. Thl.lse :mal~i!!ts aFE Hl!i!a bl.e nea r~~",rr redl ~Vli11tt~t'S,""'rttl IP@~fI~\a,l lPp,U(a~I~i'!s 3IS f!af1l!'Jml~OOj-S(atoeop.tol.ller!:u(;l!lIkdiE!vioos. a g

!.lIme) dkm:nsiolls f 2'4l




~Plllic<I~ioJlof SI min ,enginoering .lllkl (olloidlli qlmnllll rHJot syslem by imrorlucmg a mctJimd lim!'
~)OI:!I_<lllOO'llil[)' CJ'~01,I.I.~ reglli;:llx]y ill
f' .~gll:m~ll . (l!"




\'I'1ilhi!l,~ COlloidal.



.. It1 ~1i!L."" qUill'I.11.Wll~.-!.,'"(jI].Dir~(~d '1J11il;>l!l:I[l!~ .w !lU,~C~!~~to u S:Llb$!J~m:; OJ Ctlt1J1J1.'AJded i.nl.;'LI ! id .~.K-dililitl fI j)_ w ~lfrials 5deOOll tDWiisioo, l.awreoo~ BE~l~)l INattooal. A '!!.1.rg.ct 01' ,coHnochd !1l:n1'OC'r}'$~1 [1'.:.lC',;)I:rchis 10 ;I.ol'borattry. ElieiI~eloe)':, CA9~7.201. USA. ~e-par!iIri!e!u oi . 1;1'(n1li,~ Ih(.~(~~,mil~ ;\lII'l_l~~~~11\.'S \",hif,c, :le\'Cr<lgin~ ~lile (]hfMli5!l)i:, Unj\l'E!'si~y or Calif~i~, llelkJ;!ley. (J!, '9'~72(11. gy Ic\,'(~bi_ S.~I1J.II;;tU~ stith m.. qWlnl~!lJIl1i.d'm ;,JITIIY" LISA. )[:I1,mplitillioQM~ R't>.l&Iirdl O'iliisi~n, li!WJ'~I'I(@ 8.llfil::~lf~ and INII~'t['J,wiru 11(.'itQ:Ii!':ffi[Il!ICI!llK~ c~~n DC la.hrit':fuILi"ll :U:IW811l2l,gCS, of ~ulllI1itm.'phfl~eHhblri:'CmioJl, such ,IS Nalioml La'boraterv, B.ed:llley. (.1\ ~4l2:C!, US,:!!. ··,unlbHilY
~!1l '~iUt:]ll ~m


A l~tl(;.ttl]lT'ilY n.(,QllIHlt1I1.UiIl wilh~n ''ll1if~no· don.<; rod. cftieethtcly CJl(:l1IJtc~ a one-d iilillen$~on{l.ll (~D)

~h{l m~I!~

Icd 10 ,l wi~h: tJllgC oil' ad\'anlCctltl!n!]l1ei~1 H1J!l!crim]!\, 'with ~o:n~mU:m~l!CQll!ll1wrn ~1,'1,.':r'"

by VtilCHullIi· !'Iml \'~po[.(k~po$;[kl.n~ !.CcliniqLllcs ji~~h ss mok~ll!rnrbcMn Cpil~tl':'JI (MUIE), oalld \!:Ji{POr.liq!Ji~d.5ioHd (VLS) pirocess~s. rcsuhi~giUl


s,y.lTirh(:.~is oOJl~pnuibj1:ily in di$rJ.ililil~e !Ui8CI 1~es.tauthoi'i OO!l'llt!illMlled e(J!<!lLy to U~.~~Fk, Ic,~., lor IllSC inl biolo!1;Eca~ ~[lbc~ inl)l 1][0 whom If(lln!SJlIDi'dell(e ~!A!I!I :b~~f!dil'oe.~" t6. 7) ami SOI~Hio.U11'i1:(}(!l.'S.sedli~ht-Clflt~tt~ng ([j!M5@IrUN~~Ley.oou h)w~~.



filg. :11. rE_M ~iiIi!laq~ olf Sll~l~rla~ti~il's ronl1(\ld thrrOllgh ~::!rtial ,cation e};(;na1nge. (A) nil? orig~na~4,8-b~64-nm (idS l'iIaJnlo:r,~d:s:,~B, ~l'Id C) T:r,ans:rl(lmled Cd~'hA.gllS $\Jlpo€'flaukes •. Uf!S~O HfiSlo'!i[.l!iTtl of Ag;lS: sre9m~nt Sp i1l!c,jng (ciIi!iflt:e.rAI(IIcel'ltetr). The aVl!trage5pii)dn,~is B.8 .;!h; 3,.8 nm •. The sa mliJ1.e


U!E hiislogfiilnl'


gl1~t~r 'til~rI 25'0



;<;\!rCJ'~m;~jQe-.1. prmmsmgfIlCW generauon < flmltcri .. I,~. I{}.IJ~. SUCih '~[)swper!f!Uice~ e.",hihi~. ~QJl !h]~I~lL;l~J~ ~ff~c~~ <Lrld ~N lll-ll.l,!Il'i~!:!~.b~tll~C ~11~~r ;lb~~~W to!J,;-CilI~ .h.rg~ .:'Im~.ml,!l,~t::;of 11lI~t!,t:'Ll to



w· A§:l:S l Nf) .. ~Llm\fli!\,(!I'.whcn~hc Ari"





d~slucali_om: ,anl(l
(12, J 11. S'!ron,l',l

;,l[rllro1>i:hlni!!.tc~y ](~ cx~h:I!l~. l!J.tq f1I,,~i!!~.!ij~l!i!:!T!~lr.ifOOS ,Gl~p~<!!y.t;-d<fl. r~r.iodil; rpi3u\-,m of Iigil'll- ;m~ ~:iI;rk-~\Olltrtli~t ~~gioniS. Fig, I, U :Billd C), The 1I'\.'t::mgc~~dng t
19118 !lIW lilfllli!oo.iO

I\82S, (lI;;SpeCl~vC'~. E!W:rg)'~i!:lf'!'C'l1iiv.r: x"'.my lRl,,,,,,opy lEDS) im!J,~c.uK."'" Umtnhcst~~I)li;\cll rods Ollt~rmn~ l1ll,)lllV~~ Cd-Sr;IJ'~~ Ag-S r~c;h rl;)g~:(ln:\l
, Fig, 2A) (J,li~, P,oWdJ.;T X-;rdY {1!i.ft"~]~·1im~ {X1ll;,DJ
(!H!~l 'ooJ:lfijn:n;;;


i.1Jl~ JIDt~~llIiJC

or \\ l,J1:l-ti'J;;

ecilS mId,

dli;1'.I;lil.Mi·lI~ d,,\-'kvperfol'lfllla!'lC!1.l

COUj"i1i!1,g orclL.1iclml!1ic ~,Iiill,."'l'im~~k~ th~[j'~inlL;I';. c~[ilj!!l fo.r ~p6iet'iJ ll<y~t!.tn'i~ ~I'id good. c.,,~did!llC>;, fOI'P.~'OH':iJ'iiic~lppl ie'dl~ol'i~. ... l,iI1iCfi~'iojlaij :,;lIliOlij~ c

~t~nidll)tJ cli1l\.'j;1h~ioH IFi:g. iC.~n~I).The ~paCil13 b~I~\i~I'iI'!'Ll<ri,odie~(~M;r~'1C~ll:>; 'be can

between lhe- dark re-,gjons is 1.3.SV1n~" wl!h

.or ~r+lJ


~mlHflcl ini~ ac;n,_thi~!!l\g;lS (fI.g. .2 B) (:!J). Peaks Olppt;;';unng i!lthc Qri~i!l:t.1 CdSm'cl~, '~X1 be ~1lJI:,k~K(jcltn wurt:zil~ CdS [Joilll G:'jI:l1lt!:~Lkll;On Pa,tv!:k;'l" Di~~j;.d{'l[ijiQJ'i SI,ID:(:II:la~:S (jCPD.SI 114,1t04~']. al~d 'Ilim~ in ~jC lii.!ll)! ,e)(e'h.n~ged rod:;: (,:"lifl be iitd\::~.e-d1.o!i'!.."'Iln~~'iil\:(.!ICPDS HI~-oo,n.). r\."ab: ,ihible inlhe s!Iriiipcdrods c.'l:tl ·be!l!llrib~lIed


~.lI!:ootr"iulteill"-.'.l ~hwd~t:'fllt1fiel:Oel]'k:
ionie l_fIIl1Sj)M

colt'tl"ollcd (llfil;l. S]).

by Ti'aedim'ilC'1ii::'( an~:i~ii:'ii~i~,1CdS RXI

dc-vice::; mtdlJ'ic l>ludly of

i~ I])

~nation ar eXli."iIuk.>d nrum:H:vili~ )1UI)C'riaJtlke!',[c.S.,. a ~~,efll~tlI1IS Sl-Ge or hll\.s-luP}oollL.'1iaillg h!lmdrods of rep~.l!t unhs (N If'l), To ndlicvc ~~i!),. IFi!lJ.Z. (11~ r,fjJd@rf!u!t:i~fl ~hc '~IeC'!.!!'iro'f.\; 0I,:e .ancrm,b.l~ ,;br' the gfL'I~"'th of or Cd:S.A,g2.Shet.e~osIJru(hjl~E'S.{Al' EO'S SIPKlra (If C'fLdll,'~)'I:Cr.;Tl~¢ (om~.m~~l01'~. ~LI'lfleflmli(:~',\1 by D the- ~1I]ped rods oil '~!h~ ~]Ii~ $(~nle Til.iJ'liL;..m!~I]C"]~'·J'I[.'O)II'ii~ili(m Ofll!!'t"CU~l'" \ C'o:m::cl1,I!r.nklll g.'lOI!(:t of realen lor Ilre~cm col~ Uo;p} ~ighl'- ami (boUmTI) d.a ~k~(~If!il:f regi@I:I,$., as.t 10id:;ilJ ,gro'\\II,h l\CCilnique~. l'hc'l<!r-gt~t Illl!mbt'f mrres,pondil:l:g to (~~,SOJL~[~~nl!.ari!l~ 1<ly;t}17i l]vtIthu~~d NO' rar- i~lhro(l, .md an.dA£I-s.-nch. rrE~OI1!$, ~~ yi;"tlh~ $J:.'GII!I~m.c~of p<l!IriJcmtiio!l:' ['(,l(IU i:11I,;1ji~:I Ihm I SP€(tM.l~., i:l.I,!!~, arbl~~

sy;:;lOJ1....... V.lS ~l'Owlln has dcmO~t"Ir.Hed Illlc :tOI:~

E"'liDl1[~i!lMi!()i'1or rt:!1ICS.\! l"C;l:;jio:fiS:>!tffil,d, thm. [1iI~ ~i:ginl" end da~..e;Ol'l~msl: lL--gtcmS, me CdS lind


ill:>~m.I1CCWf6" :l!_t~eJ).diy~:lX~ taimplemuut ' Ilg

Cmiul1 'CXCh3l11:!,c'prcwides


,(1 r:.u::~ IitK:Lhod lc


1J1IiI~!5., ~B};OD~8Ctr[l


for SYSI~11ij)!k:d]y v8Iyin!J the proponion or !:wo ehemlenl (lml~I)QSiti:oaswuth~t1 fi si wgk~ .~m;noct)'sl,~I. W,~ have prevkllJ.~ly :sbmw Ihm, cali(m 1C'~dlml!l~ ean be !m$cd III flllly (moci
re"'CllNiUl~y)OJnlV!!:11 CdS!!:!. CdS, m~i!I edTc~ulnt}oCit)'~bll<;:to [!r!C' (o~m~d:~ng, i\g-chil!~ggC<l.li(k~ .~lmlllOCI}'S;~811 by a cQll1IplC:le .rep'!la,ccm~lC',l1l l:'Co:1CliOl1 of the Cd2+ canons ror c<llkl:Lls I( Ml The resu ~t;jlm rnm~Ii.!llis jhe A,g,,;;m~'rnlJnOllqg of~he ~!;)Ini!'1gmnI~ri uI'(ilJ,;:., Ag~Sc. t\:~S,. m~(] A,!,l;:lTe)i. The si:i'lJ;_~<!ncl sh~l~ of Iii!\.': mUlMf:lCrys~~uI ~l'(l(: pin:;;scr\"lo-d! w~l(;nhht; ~1l!llllJCrysnrJJl hilS 111111 Mlmum d il1lletJsjo:l't~ guc<lllcrth.;m 4- Ilil]] I( Hn..nehigll~ mnoh:iili1)! of ,c."ilti(ms ~n (he CdS~ se,Tt! ):!all][lC :mgg,c81S tlrm~ rf~rtinJ. eatlon c.:«:h~nge Ima" lead WiIW ... 't'iCSI.il1i£, p<1uicms 0 r scgrcga •.cd dornni IlS or I-"Ig clmlcogcni:d'c withill a O:l-dlilikoEJcnidc ~t'l:Il0r0dJ.nis ~edusto im:e.,"lii\g,1lC thepossibi~~ il~,1 of convening [I prcvk)l;Jsly ro.ifl11iIOO, nanorod of II. s!~lglc ch.cmlcill Q)l'I1lll!08~lkm inw II :>~~~lltCd pilI,t~em by a s.i IIgh;cslcp PiJIflia~ch;:::III1.Ci.'lI.1 tr:;(jm,.~ ~;br.tllmhlll.. ~11: tl~c 'CJlSC C'x;p~,oH .. here, nlhK.'1!r xi ~jnnmgc:l,ncnt of l'eg~:I;urly sp,L'Ctld AgiS ,dm:;: ~'Qm[li!lWwi!h~!~••'(,dS rl)l;l (brm~ ~l'lOmOlnC1;l!:Jifi~ ly,~1 -3:1\'%, ~;,QlliQP! 1,;'.",~I:Q'I~gl;. ll'!~l1~",! .. ,i~fi"fT\;"d (N Ul) b'll'1dW1fI of Un,~t\gzS dot~ i:;; ~~I1~br;:dd(;X! witu~in lJ:ieIllHg(T g.apl)rlh~ eds., C~"!lIUalg ~t ty~ l !ief~ro:;;m.r.cIJ,IIUC",'ilh iU!:CR"';$lillilil'Ciptl:Cil:llpmpcrt~. Shrdics ofp;runifill cm.i!lll c>;.ch(j}fI!;;~ilor4.(l-by-64J-.f:lrn CdiS to Cd$"i\g~:S :lJlanQfCd!; ill',e :shown iUI ~mjlsli1liss~Ofl CIOCIIlClI]n1~oo~C!pY ~T,EMj iil1l!.1g.es

CdS, rods Uilad: Uli!'~), stJ;~~edrods'Ib~lI!e ti Ii~), and ~UI H.y iIl':?:C h1lrng,ed Ac9.0!) rQd~ tgfml ti !'!I;:).
Spechalhonill l~ocils!)hmY



!Il'~\I' peak!> Cl)rrespo~dilllg .lQ ArbS illlld ,[I lII1Iadirierl (002)


lFI.iptiiOOl ,~fthl~ CdS

indliCi3ilin:g inl-er·


alolilg 'he rorJa.i(is.

IFill!!. ~. Eff~cts ,~t
]!FICJ~s:iIil9 ~INO~
OOIi)(~l'1llrati(l~, '{A

I~A} lOON 1()lln,{l8i11~ tra.1nollil I(AgT,I!d~24!-


-0.2:).. (8)' IliIter~ l'Ilooifate mmm!i'.a,'~jt;m thai: ~~ed

illl fi~ L

,~Iii I:&~~(! ":xpo.!f.irilCiniS,,

~h~Uflili~J CdS

rltli~mrOOi> (f~g.. !A} \~,'cre~c.~LIQJl1!Jlly

l'Iis:logr a l1l1I1. :J. OOnm. The CdS colltoklailliOJlftrirOOS. were 'lddai {E !1I!Il:d A IP~~r·dlislribl!lli~'fI[ ltis~(lgraIJ'l'l!l f[lr Ag2S re~i(lm$ (1m! ~Ii'!dividl!IallGdS4\.g~S, nail! "rod~, (E) 1~[lW ~o "looluttQJl ,[}rlolu~ne.l\gNO~, an.od m.clhm;ol ~~I al1d (F) ~1iI;tElrmed~.:!te!!:ofl(ierttrath'lfl. Illltr(l:red distall:ul?S b(?l'I~en elll(i~ MzS w,egliolll, measur'ed, fllJr200 , ,660;!1 il'l ,,~ir(l9}, ne conwJ!l~lio:n of }\gN'Q~ Ila iIilorr!l)ds ifl'll'<lch ~~:the ~ m,p~es'€lt5 shoii\!Ji!, ]111(A) iiili)di (IB~,.S:piad~g$ \lier~~ f'I~'ff!Ila li:1;edl 'by th'e !'!iI,Ilf!'bl'lf 'of' ~'\Im:,~cQ!~tmlh,;.d rtlll.i;:l~!1iI'lf[lw P£ll:u;;e!ilm~-ilt;:in of AglzS 1",~k~f1,:Siii!~~~he~~g~h ohhE' Jo:·d •.lQW C'Of];(!efI(tJCltio!ll(E) silr'ow~ !10 ,cQm~~t:i~11beV<ln:d:th,~ f1,~fe:ll~ C'd~ iO'i~ PfiL.~:tu nn [liqJ..:: I;lm1lin.g 11iI:iI¢r1i:JL ]1i~hL;: 1·1i!;!'i91hJb.~r spacing. Il'Ite,mTI~diiate clJon~en.tratiIjm 1~11F) shlj)~~s !it pEtfiooidly, ii'lihkh extern:ls oyer s;evero1ll,
p!\_''i'l~lj,c~ ~?iC(,'li:; Ag {if

s,tlmoilll and th!!'rod diaH1lCIlef as l~g:I,.I~y m Ironed (SD "" w a t O%I.whCl-:ca.<;1 he Ic:rDglfiI ,""",,,icd between 30 m'l!d

Ilat~()2!>(A~f'JCd'l'''' ~o.,9).5cate (A. and 8);, 20 n rIt OC and DI) H~OOgr;ms. of ~~ m.!'" Q'I' AgzS, regiollls pelf WO!it '~Q low and (D) llrlllerm,i!!dia'le (joooerlffii~on. MIlle· ~f'll50!U~,







W~ffi' ~mt!1;f.ldl f~r

" A~.S fegTicm5

T. the :mdll ~


ml!:CI re:s,t l'Ie'ngih Do:rs,.

memst.'bblc :ll!<!!~~; I,~,.. the iCQIllJ)lc~c.iginil!1g ot' Ihe : dlr;ecti('loll {<lY'if\J~)tif;. ~l'lo()r1s.ible r<lf ihe ,oo:~1Ilbjmlt.i,m~ 'Ci'r~bcsc two j)~~~. No Ag pc<!k~ 11!l': pP:'''~1l1. F!J!r!~~cmroro." :;4 mli!~1~io:n~ h\'Q AglS rcgjgl!1~ i:-; alw;;!;J!s a ~QwOl,'"~ne!'g}' ~!J~sh'~' c~m5tk~llW<fm;~iQn(F~g:,4(,), C'dSntmns Q~nfig\!ir,<u~i(l!ll,~)1Ji'Qllli! th"t (:..nlm~. ~tgi~y h~ 9f~h~ XRD ~)~~.~ r ,,.. !]l~N~l)~ A~S jlgt,g 01' Ol~· Irl,l:ilhi,l~~ !!\"'~:Y lrol]]~hc ~iCI~C:~! i'\,ruS ~l:r~ll,.1(!l1!. ~~~ni~~ Q~IVI.;'.l;;,~I.~,qr:!!"~.atomie ~31yot;ilN., For ~\VcO CdS Qy$U'lUil~~cl{]!]1);i,Uut~ Wil~ di!l1~~~io~~~ l:!iIl'I!d~IIc(:('8s~d by z;,i~jl(pm~ ,~~m~~ic~~e~q<l'l'1W 1;;\'!O;:I~h1, A S~"I,;~U!~ Izu:tor d~!ii prPJtiHh.':;SU~~lfCgJI~ar Ag~, ~gmi.ll~ts~PI:lrmlc:hillg each oiitci';nil.c .: illg ihQ:~ ofdte ~,,~pl:c .~~gfl,~ q~mlib3lii,ldy with ~'JC:ing or~tilc :s'!ripc jlil!uem is lhe ~I~~ Lt:~ll!l· ~lD'"C'xpcrU~I.1C'~11.dpilltenl!{, i!l't(;rms !Of I:J!c relative dlsfl~"CCmC!lk' in the CdS i!!Ir-c-in (l!:I~po:siu; ~i(m n..:l\lI.~'(jI~ lwo A~;.'!S ~.gJIlC'~l~ h(;l.;';.JI.\L"~ If ~~ie d~r:~'t!illn~ I~di!!lg~{l' !!Il ~ru~mL;'ci(lf!t1.;'!'1nlx;h\i!llli;;n ( itI1~~n5:~tit.li, AgzS pc<ub lG CdS '~tk~, 8lJG]~l'la 0r i at il1M bl~~C""t~"~ oW~It~ el;mvclNioJ1~'(jd i~TE1I.,I~ z.;:~~lil]n lite Qlll(ir\r('n1il'i!!l CdS regi.oa A 1l'icldt:1 mli.~~r~dclg tll~m! !!li~'S hj:!:l~1itt ~tr...in 1;."i:'i~:~1 lot InK: oCohC'~i~[ i'I:[.oI~:L~ (O~]~,"~iGIl, lii{4:'O\'t.~n tlliit i'l'i'il'lhllj.:.';S, (fl:g. 52), Xm our (""':pcnl'illell.lal XlRlJi 1)lIl ~tiU~ !~riSt",P.JmUUOlil>PSII, The li:nodc~ h~ COii"i~kiil Me! mlMC~ii~~, ii!l ~hOWtt1hll1 Fi!;i. 4A (,?lit To Icil11~.t!i,c CdS (001~, pelJk is ib~db'.' ~Ulcl,w..;aker wil:hll.he exj1Crii'n.~ru:t1 lilttli!l~ l1uII ~rn...,easi:ng Ihe rI'Ul.teh 1.!'It: I:m5il1 lat! k.;: OOOi] ....1:8Ll'It ror .cdS (4.3 Ai. rod dianne •. r inere{l~e.~ the )i;P'icIf;I~ between e fIT, lh~ £lripeoo, roru: liJfm lor iJ~c' init~.i CdS S!ii'i!i~




pic, n'IS indie.u.cs t1lteelL"IISC~,C,clS cryl!il311lilC sise <Olll::> I'nlle ~'(Iwlh. a.d .. 0 r the rods: (J.2)] .tller Ihe :~'<IrI:i<lI~,~1 (:;xc!um,sl.'l. ,[)e:b>'c-Schcr:li!.:r .tmd)'sis or llIC':!k \,\licllhi....ror ~\~ml, :.. 1il:ilC(,hod u ~m~:Jc;s ~ndiC'.l1t~lhm IIw CdS gr.:li~l $:~ akm,g

th'C' A.1,liS llooy-cciIlI.ctl.ed-cubie

I£ij.uf:ce in tl:l~

:iegm.~nls (I~~. ol i. Sin~i l!lir cllOCl!... {I E'>ponl:mnoo'u::: S

p Iltme of imii!rritcc has



CXI)JJl1iJ 'by .:I%iUl nne b)' ~.5% alon~ Illc

(lmCri~l~ p-rodll'CC,j

,clil~S hl~WQ

1:1)' MilE


dimcllsl:om;: hm~c been cxpkliuiJd

l~crIK'li1diiellllbr dil1l."eltiOUl. There is <,.rel'lul!'ivc c],,,"',tic!nltc b~\~'Ci.:n !'cgmel:ll('lof ~ik(l n,..~~riai !11~ ;:1:,<-1:;. hot.,," d~t~:;i}d fiui'~n~30 'n!'l'tW 11 W 16 beeaase 1J1rti~~ .fti;lIlh.Y~!g ,\oo'1",~i!il RI.%WI~~, ~I!I!.llOr '~~'1L; ~lirip(l~ mdl;. Tlli(~d~~~~'i.t'iJ! W ...1l ~z<l i fro:!I!l '~'~~I~lCC ~ll:~(l 1l",k1 It Vfn r.unood ~1I1!l-(171 shew dlm lh~ d!!,Slifc cm::Ig;' ~wrod in the I'I1d m,lov~g thi,~ difl.lclkm is a~1lI1lbmcd ~Q lh~ imcm.l~ ~inn or (til~ ~OCH: 'f~~,'!llCiSb;~ Ih!i',lA~S ~.lliJ~(,,'ri<ll. im:Tca.W$ n!<.lrlkod~y as 1\\'0 Af,l,;:!S .~Cg!ll!Cf!Ui Itl1,;'Cml~c l~!C' S]IIln!"o'f k\lIl~th ,~"(1'!!5:~11t.r~1[ l~ f~'i~h ~h~ Olprtrtl u ch ~'•.!.ch O'!~wr (.f~~, 4B). BfYndl :;.m:.~n~n


'with c\l)rro~I"tOnd~I'Silf',d.lllmcnt~, ~c':ilo. (1)1. W lowCfI,'cr. d1.¢ lID ~tfomcl!'y C"~I~~l[Iredhere i:mllQ~es .. ~mng~con:,"'~!""~ll~ ripL.\!i~in!;l,p.v~~>("i'i. ICtu]~Ulg 0'11 tn. ;JI~ ('5JJiol'CiidIy lir.rul;l~! p:uh!(1 Hmb~c. .CCg!.!llarly :ql,..,L'>d q Wlml~!.!Tn dot" '\.\·lllrln <I, rod. TIle ~lln~1.<~nooor ~~ !in in ,lIwlnv!'Ig~h~ su.. IX'rI!!l~ltic~p:a!l;~~rlI! h~ il h.I!~~'~HL;"dbiY n~i!] Ulg C".I!l e;l.... ~

<l!;\~(\nLg~ !fIH!~D i!luhi~ s.trip(;domd H!!,g ~1T![lIe. TiEl\.!~ i~l\:l£:L."SSh:CJw nlm•.iJ1C!AfuS lJ'(~gim1S.wtlllch ha\,l(: a

b<mad runjii.e of F;Cpar".'li1:imIS~!low c(mccl1.mllio:i'i~ !(:Fllt,..3A). become incma,'>ingly ,oMllred ill, s.ilig,hl r Iy higher ('oilccm,mti:GllIs {,Fiig..'i,. 1,13 IIml C, mld :38,)1, Ute CI~mlgCin l~le nmnbcr and pcYiodichy ,(slucin,g) ·oflhc J.\giS I)cgim~ !>LJ,ggG!1~syslf,lllatie orgl.'ali~Mli,on <IS,IIKl volmlle I'mction of A~S il~C'L~l'l'l!"\"Fig. 3" C ~. F I'. ]1I!rnro<l A~S spac~l~gs ~ were ,oorre~m,cdl'llroll.~l 8rai,-distl':ill~.u len lintc~ion iI~ \,'I'Mch the distaaecs OOtWI(.'CJ~ 'C'<1!.Ch\g.2,S J regilaH <1nd ,iiJ!1I mJi~cj' A,g::!S regions 0.11 a rOO \,'I'C!'IC ~ll~ilI!\.\.!rodl1l!c nJ]pn~it:.illli(m,ol"lhc Ag,~S II!~w(m!'i imo MII'ilC'Lknlio(.~'l is ~%~Jlin~ lli!~ p,:'ri(ldi,(;:i~yol~hi';'
,(Fiig.. If ), c)\~cndin!:l I~WC ~c\O\l"ml It mlC"JrlCSlolliC:i~h'bor dr",r:.m:~,hllihc ~~pcrbu.iC'CS,


Illc J\[l.'2S i'C~im~s, are spaecd evenbl o~mig mhe rod, wJlC:r'C3i~ulopcrio(Ji:dlyis, 5lCCi11 forbh.cr.owcr .1'\.£+ COl'ICCibli:':lil l Fig. 3, E)'. ion ne mcchan ilslrnl Iby l,:\iinf~hlhc' ifliaia~r~mI1~~!c~
rI.li!liU or rnndomly dij]~r.ibllte.d sm;[d~ i5;1l~d.,> of' i .Ag2S cvolv~" imo a periodic, II) pauem is of r."t;liiicnibf imcrcst .Bccau:r;c 1he:rc C;,,,,i:l"S a posidye (,dS~AS:;!S ilfl~erl'm;lc Ibn1illu,u,ctll c<tlc'rgy 1(-1.'~8 eV I~cr Cd-.l\gd.S cl~IIl'C1l1~t)' ~nL~r,fi:u;~e ~'rllI, rr!ln~ our ah in[lio c~lcy Im1ons), .~~ !:'i



Cd: orange

= gJlI"sli




\'I:2!1~ 1:50(1

Wave1lerngth (nml)


C'J.l~rgcl.k.ally rll'!!<nlib~r;: III u~~~!"~~;iilJ'!:o:d:l, A:~2S isl.t!~d.lii in:~o I;JI:g~r A.g2S St;'~~Il1,,'m:s. The fi1b";~ difr~""i~] or (:HliL'}ns~t!~"'~!:\IU [;;;i1:I;I~,~i~l'nl ~\ WhL'J."~ O:s.\w.ald ri~)cn~ng l1O'lWo;.,C'lilIl:II.: inililliity rOl]n~ i::.;kllfld:. Qr A!§;2S eml ()C('.tl~.M)' d~m :~fu:g('rblun~ grow m. th~ C;o;~ISC o,~ UCLld.'!)l Slillr:ll ~ OI1.(:S, D'.iJIYl.I:s;i,onof lll.~ ca~iol~Sis~no1[~d ~lIS1:.' both

t1i.i:ld('cP+ aT\: o:m,sidcrcd ~ils~ dmilsers tH-.2..'i). AII~u. A1!l,d~liJlt::ogcni:Jcs CJolhdJ1J;t :su:pcr~
iOlilie oO:lldjjctivi~y

in th.c:ir


p:~a'SCs (2.5), A crilicill junctllgu~ OOOLIi'Swhen, nile re,gi.om;. or i\~S :gJl._"toVI. ~{:I'I:hc pOll1l~ where tlilCY .~paillilc di,llInctel,' or the rod. lu Illis ~1!Oint, ~i!lthcr O$I~~fJMlipenlll_g is ki:m:~['C<I~ly p:t~ hib.ilnl.. bec'.u.!foe.~n "W~il·b}',.u[OHle:!l;di.mgc . Ag"" ,~rm'l'l'!g~glill~nb wm~O't !'oo~Ot; 111c~ol~d i:tUJ;afnci.d ~~~I. 11JI1o>k~lcl~to AruS :!>.'Cg111t'JiJl." of n1f'.3:rly t;,t1lm~ size t Jlii~. S;,. T~ie mJ h. ill ,Si


and e'xp~rimenti!ll. QJPtka ~ ,c:h!i:lf~n'eri~<a,tkl tAjr ('IJ'bk'i: yt:t;l~t ~t tr!prfSl!!r'ittlti!ln iji( (fins i1!s:ellif,ijr ;iiRl·il'liitilJi l!<nef!iIY C;;(Ik:i!!~aJtilJin:s;A dis.til:rted liIi!:modirnk A9i5 (100) pllaln,e ,O®lllI'lEl(:tr; 'MtIiI 'th;e< 'II';!I!J ~iiill! GdS (OOjU pta liIe. (B~ EI.asti,e: iUI~~S!)' of tille' rodi as 11'rtlI'U:I.iClIII ,of se'gll'le'JltS:@~ilfi!t~QJiI (('.e'JiI tet~OO-(:eJilt@I'), {C) ,l".a:l:is ~(aJi!1 fill!' ~Ih@'(.as@' of MO ltillsmi'J!tchlHi s@gmc~ts oiIt ill [l!liIle;r4Q"02'ra~er £eparatioH dlistillilce o~ 1'-l.1 rlm ('I:tlp,) ilndJL2 ..1. trIl11i1 ~bQltolll1l). iI1h.~ elar;tic ]l"IIlera!!liol1l ilehveen ~~Imen~ i~ 'grlei!l~~.yred~!.u:lerlfm ~eparat~ol1ls >][2.1 iIIm.A1rrllWlii sl1®1i'Ii th~ IpllaulIlH!lfal Oir mi5r1i1al(~~e'd segments. The CdS rod5 'lIs.ed (011 \!'FilF ,ci.lk:I!I~aticns {IB and () 'I'clerl! ~,,6 ntm ]n dii:lml'e1iell, ~!.IUIll t1'10 4.8I-D!y-4.oo,nm laltliceomismilt~chtecl segments. 'tH~diw elil:s;Ii,eulIlsl:<ilntl:1l~Vlr
lr:ijlllRPL. :spectra alt i~= 400- o!!iil~ 5S,O'fHTl e)::dt~ti!D!I'I. ~cSPect!~ely. CQ~ptfF!9 b~l\l{~JiI. ttillCidS oinll!l.t!tg~ is evidie'ilit il~tine (Omlp~~te qli.l,~nc:h'il"l£'J1 '~he vi~iibll>e I?L (im il"l ·lh~ he~eros,trli.u:Uil~e$. niil~ shin ~n NIIR Pi~ i:iIf I(E) is dl!l~ t~, It! ~aliUI!I!llil (!(!IlfQr!i1I~iill,~lf!t ~f'lJhe Agzs" ·

IFig. ~. 1!'he>oetka!tIifii,(!Ide!tili'!~ r

,the ml5flilatched


l,\'i'.e-f~ riJ'l@11iI

iiilb iilili~

(alk!.JI~~ri(IniS· 'r,@rlill'of!,Qdili!:iol;

AgzS .. (I'l'} Vbilbh: ~tild (IE)


",imil~r sradies


Smlndc~ 1,/' ~i,h,.\~ ~:pc;:]!~cd~!l!~~(~I~Il !il~a~~ ~Lnni~l:!ll~!I,lCl!m l!DnOl;:ry:;;t,.l. hl§!.$m~hCiiIolf\.'i'i, by tI,~UiG:ing, A:(JH iO:l1:o< on1<1' lUiA:s q~~lrilul~l {lots P9}

~~llll!()l;D':-;i .. l$,





M(lk<iW~ {,'/ ai. .m,,!

and CdS/Se

Il;IIII10t~~ l.;lll,.3/).

Bot'au.\~ Aul+ has

a r1u!lI;:hgn;r.n~r d~mn ;dfinity I:h"n Ag +.. :m-. dl[qeaiol]onhc i>:!I1,UL];:;es Ipli.'I.cc r.llhcr· tlilOlll:i .. a:i¢"'~ ellamg.e rc3~!i[lIi1, lhc ~,~t:~\\t: ii:i~eif.."Lti3~~1IDI1\Y IntC'LW~.ul tille l\\I1f.l irnl~[('rtl!"ll:;dri",~ lI.p!lil.SC ~gre.


A..~S~(·dS 5j'S1X:m. ICflilii1.!l ~c O...wllld ri p!i!nl!I1:£. 1,lo,*X'V.~u.. cpim .. t ximJ ,:sInJi,inl does ItOI,P:I<lI), a m.aj,orro1·e in lilt: Au Sro\\~lh. ''laW IheS'Chc!cuim1~nJllC'tll re!';. co.uuhme to lI'i~}e111 :into ,~ifl.S.[ic"'U1.1C'oaldom,~il1~, cidwl' <It lhclfp ofdlC rOO {Cc_li~ Sc). or inside l!h~ 'qll!.mtl!l11 o{rt (h~i\!SI,. h~
C'(l~!uro~"!. ~,hc'~piw"'l;l~l"L;!mJ!iol1.;;,hiplix!:!w"'Il.:n~hL.1'


1.0 ou,

8. IML Alil'lenT!ilfl!l! f'l 01., /f.afJJrf 0\1.2:~'. 642 '(2;004lt 9_ It ·~,If~. NI, ~ Fromer,. M. t. ~~r, A ~• .t!IIM~I~~ SdfiliC'e 31.0, ';;6z (ziiG~. 1i.O. 10. y, I.:~ Y. Wur. R.. IF"m" IP., I), '!f.-mg, A. M<ljlImd'ar', Appl, fih;p, te.r,r; ~3. 31196 t2(03)o. 1l'll. ~ S. DJItl'!o.'!lIib;tLJS ft al., I"hyr, S~li1d SID!f! 41, !!J79 U;99t)l, ~', G, I~~~~r;. UI. G~~~. PMfJ.'j., . .'J1.!:19- 8~, ~Q(4). ]8; IE. frl!iHn.. ii'. It Gr~~.[I. (. (lin~n,r. 1[1...amd~. s 1, Appl, i1!yr.;. 9:;1',114~2j ~;2005~'. llA. '1(. Y. Wu, i!!. ~ilI!11 .P,. I!). l(iI~~, N:gtJoklr. ~, 8~ (ZOO-li. M. ~ s GNliilsen,. L )1" l..lI.utoo;n. I. VhIl1~" 11).c, !imiU'i, C, ~. ~i~!l~r; .. UI"','15" ~ 6'17 (200itl),



:I!l'. 31. 32.

]~_ mr. !!jork .~.,~~, lI(plla' ~1!'I1":!l.e,7 ~j!(}O~~. ]if. 111f.. t.!IilJl~~nj!t of" ,mJlti~ 430. 190 (200i1). 1!.6. III 1ft So~\ 's:. ~, HlIglm, '1(.. Y1in" fJ.. P. lllIjvjj;jjlC!$, Sdm,~
306, 1009 (201M}.


uwo il.]h;l~~";\l thl; AgzS-CdS "'1!II1)~rI,ullti~ ~thl(;" ill ~IUIl;,'l\1\)'f;l!~I~·n Sir.lin fields lrom Ihe lanice ~Ili:il~ i

119. fflalt.r!<1b a~dl mll1!lli~d~ ·aft <li!l'illl~~ ~~ npPlllning m!flierijij~on ,SciMai' O~lil!lll. ~O, 1\, fliil'll!ri~1 ~eg;m~~~re Ag'ri(ll 'MljlilII~I~ or 11\1 ),' g~ j)i\ilb!!IiA.:f d~~ lli 'Ih! dJ!omm;po:~li1:iQfI ~r .Ag.) ~,*l'iilfllllnd~ II)1lhe' eMwn Ib~rn (lSI'. litis ~Imi da'm<I;e ~l!;o
iIIi!'l:o:FlM th!! Ill!a!li~and I!iFl!\n!I'I!ula.oourale, fl'l'la;gM 'rmrn lI!I!ill'~ ~(j~u~r.flliI vrili11 Il'iglH',E:lalilitioo 1iWl, IFI;f~!i!~r;:e;qJerifl1~n~~ '!lMfm~~~~ Lbol the O!Ii~iltill OIS. .!TOiI~ w~ 5J bJ S(! 11m, ~rn~ nj~~iii,pedl rQd~ fiili;Id~ lll"llm ,1Itt!~ tlad ~.~l'loftli'l' CdS ~J<lim~, X. ftng e1 aI,., ~Jffl!. ·404, .59 tn, MMfafl'i'l', m 5:ti!rii'i1 'ill'. IOHlm, B. S~D~U, ~,i. :Sadig(!~, J. Phy!. D .~l. Pfrys. ::Jl, rs U~9)\ tHo :fI. Wc!Q4bUry"P/lJS.~fl', :1l3~, ,i!o}:t92 ;~6411 U M. 1{01!i~:!!ltt s;oJid Slipt fi!ii!lkr ~?,.1l1 (1'9~. 11m OO1i1l'cOOl1I'Icm po'l!iliIC!rpm~ d .A9~5 (w'blc: ~.lul mwoool'iic,t t!ie~l1!i(!n m~t!irfi .iI&~um;es .~ I!>Jl,d!fQlIiItf!r~'""'.Ilblr: rtr[Jclull! mih om1), ~~gltt dl!l1JBrlion~ im. Ih~ illlonQdl~:ic pba~t' 13'9~.·)~er.il~ tp~at !'Ilr.a'iD!'1~!'!i# ~~ 'O!l!iliide,red\ ;!Jld1l!i!~ ilpi~rl~. (OOR~~to:m with minirnalli!\11i.cE di~w1i~mto th~ (001l (d~ .ri:!a! ~.IIw!'lllod~·~mllu,OOJ..cI!iIilIl: (.:l:lID I~~~ lWoof:IDt!link (1.00)0 f<l~~~, 1iI..]. WiIl[l[!ll'ISall, L W. !ii\:liil'!il •. A,..lumgl'!r, ~f1. ~, B 6~~

34. ~. I'. it:lijma!!',}, Mj't,. QIem. !!l11@. ]6S21' ~'OO6l'. r!\; M~,.r!!" ~~mJUw.. M, Gl~I-S~. [I, S<:11~\ I'll, lIleU~r. J. .I'myt. Cltt!m. 911. 9'31l(l9~).. ~6. C~ D. W~me, V. IItlllid,rl. S. IDlurrmure. j, rFl'ijis,
f)App.~. ~

II1!neeatDrnK lili~rii~eJl'lfiJli1glhe' ~mel1l~, (leif1mo~~ p;:)i"~tin Figl_ 4111,h~ n~lIl~er '01' d1i~)):n:~ !~~~ 'illl!h~'l t dn[l!dic~, i~ ·~m;!ll,. whiclti r~~ iii! ~ ~1i'fI~~I~rIr~~I!l~~ imtl!fi'lcoom" .Md~liootlli.)'. ~n~ il1irer<l(oo~oIl!1l~ nldi~l di~c!ilWl1!~hi;i!llI~e ~'!'!'i) ~1'I!~IlU .:~~ {QIlIPefc1Wiil (u!,!r,kll' iIlIl!!2...dir~t;lian liIis1Dnli:fll(l. billlotlilll!' tn~ !lull iIlIe·.illDm !!lIllle ~a!m_!!l' i!lirecliofl, T!'!~ a\1e-;r~IU r,~ull i~~ !l!!w~I1!l~ of W!~ ~IM~~~~1"9)', t M:!ik-a'i, It M~l"!lfl'l, il. w.opa.<" Ill. B:l!nin,~, Ottm. fnt. Ed. 4iS.• &001 (2006). l ~ilri, IE. ltQill1e~~ I. J'Q~. R. (cst, UI. B<i~i!i!" ~i!\\l'~ 104, 1787 (l004~, f!.. l 5iil~nd~~ 1..1P'v,pt1!', u Silrn!!\.}; P/1p. ~Irnl', ,£I :!.lIlIli', 25:",U j.200~h 0. Batt.!91f:.1,. It 81!Um1'!1\ .x.. ~n!1l.). AlII. (h~m. .so.r.',. :127, 'l108a.9' ~tlOOS)'. P,.iUI'IOO, HI, Hl!di~~, 8, rJidiOl', J. WI!Jlb::li1'I~~, flmt"~ Mag, iliS. 9~:ll~'19n).

U11<J!~ch.\''I'hk:h ~u'""¢ In,~


to lie~JCI


ol~h~r'~~"(~l!ijn~ 1i1i!'lher r\PI,:JniJ!~Th(l rc~l~m'll:g Kldp:;d rads dil~l~y r!tI!:f(ICr~ G ies e.,"p!l~ of:l type] ~,}I of Agt.l:S qlu(lJ~.~ GlilJfn d(1itd; sqp;lnlw,d,by loo]j'l1ni[lgregi.{iI.us. or ,'dS, i:n,~C'nm:ni[ wild.l our ~.IJ i:nilio clllcliJI~ IlUilOllS of (l:Ic~band stil1JCI.un;:'. The 'Visibllc dS pimllo.lltlmill'tCsc;cnce {Pl) is "luCI1iC!hed, indkali-




tl:992}, ~,. L [)~k. It. ~MB,~~mp. /. Wd ~lt:l'J~EJU1:rot!:1wn,:e~ 1iA.2: tw(l!l~, 311" t, Mgii~ ]•. U:rb~ii1,J. f!hy$~ Clr~ .iii l!~~', :i! U'1l9' ~2005l, J9. H. :5'drm.!lzI'rtrll, Pl0g; Soild StaJ.! (:Mm. 13, 1:1:9flirlilLH. ,t1l[J', liIll~ ~k w~ ~'UJi'J~Q!iI~lJ the UlS. PeiP~g~~ ~!JI' ,!1lf (rnil<r(l!f'~ni!~ f:ait1m~~1 M. ll:l[oJ!I.C!lJll!·ilJ5CIiIU2~:1. Wil lIliilnlol(. Neil5OO; C. ~iiekl~ij"lIlie N'ii1iilm~1 ~(:jtllr lor
Illeci!:on rMi(mcr:lP!I' .!It .~~I1ID~' B!e!i!i;~l.ey ~:iiic!'!~1 L::IOOI;ai~:I)'. ~ ~l$!ill i!lIl.af!}; [l '~Winl T, [~.a.gIje... .:!M 1),.",1. Son.R.Ddl thllmli3 ~I!i~IL~1'I!fi(J: lilerk:tlfl~ ~tiOO1i11 ta~~~J 'Ih ILa~iQ ,P,o:M~!aI !f"~I(l'W:jh.i~,



24. 25_ 21J..

i~lg. cQurlhl~ between

um~crk!ls at thil: hc!!.",'()-

j'!!l'lc~ion (Jl)" ;If![j Nm rLf~onll~ll! A£.2S SC~ml!)nl~ is ['Ibs~f\",>d (fig. 4. D and E)o.The o,,'lJ1ndg~p or~he A~S; scgmc'Ht:ll dCIPC'Jlds ml ~llcjir size" mald~ing the In!!]k '!Hllue (33) ror

Sq,upiPilJl'linglhil~~!!I!! iat'eirnijll ( N


.ilOO M\elfiQd~

[l~]]y cQ!]vcncd mmO:I,iCids. :llJd .sbjHi.Dlg ~ncr-gJ U] !:'l1lIDil'!;;'f d{r~:s. bcc:..use r



(I qIUiJal~!.!1ll C'o:nfim;;!'I1\J~n (F'~!!l,4El In. 'ttiU) l'ue~!:O!n c<]l]!'igl!ll.~ ronion. Gllw Ag:;!S quml1tUiTal dOls •.11(; {mliy v¢ry wC,lHlk~ycoupled to (''fIC~l ml'acr~ IKIC;}[I~C:lhc CdS


26., M~ !/h ~ ~~~Iffi ~r!. tnt! oi!l!:i~tit:

1196.3: G!OOO~,

fig!<. n~);1 IIMerenQe!i.

~~, ¥"~!Y

d(!!~e.l!I '~!!(ho!fu~~,


'!li'iUjj Oi'ity

15 Milr(i!il ,00(,1; ~~e~!e'4 "

M.1!:f iiOID7

jOcgmclllls im:cresl.

lli:lr cQII\)idrul




solar cells, where ~lC S]liIffie dcn,~itjl' dCClronlte SI.fLte:> Wil,hi~ a dot (,llay k:nd 10 1lflult~p:Ic"e.x.ciw,nl gc:J1crn.liQI] (.NI. T.I~c lilil:U1<ldon mlllo.md :ml'fCr~aniC\."'S lhrrougl~ 1)31'1 ~[i,o:n ,ex;cilal:l,ge iffl,1 call aL'>O be apjJ:I'ied 10 ather Imil::> 0:11"scmu(;ml!dll.(i~lJIrs.)'~ckHng a broaderc:la.'!;S, or qtlillltLll:l'll(;<mllllJcd Slnlc~t.!!I.'!CS. "l.u~{lIt~'Ij,dm~~gl: reacah::m:; i"lm:vc alreooy been. .~llOliJ:cdill I ~g;.'5.I~S. SnS:!, CdS. ZI1!S. Cl~:S. l:l~lS3" ;au!d~b2S3 {3j' 3th T:"''!I\O->C(U11]X1n~Jl[ cQIVQbinilidoas (If U!~~e C<JH~a plJl.lucl;:;; 1/,~(:iJal !~!ud'u~~ n:l,u~l"ruul~ \,'!I'i"~ r~lnOtiQJa,il ~'.rur(':n:il,."i\ r.)ngiU!:lrrorDl l}TK: [ to.g.,. ZnS-A~S) .ii1J1dr-vpe U o«(;.g., CI!I\2S:"CdS) wnd.tll i5l1Inrlall ... 00' 1:I~cl'ifr~c'1~ciric,po·WC<r JIJli:iC"bons (e,g.,. CdS-Bb~;h).



-D··· AL'T"·' _rl,aSSIC ..lnoSaUl!rOlmorp_ h.. ____:a'ie Ass:emlb18lge Ir'DlmINewMIB1xiilc:'o and the 'Ris,ls' 0 IDilnllol:s,Blurs
1 "

,R.. ndalm It
Ahllil IH.

11~lI1Ili:l>, l"Sl1£lrti


11I1~11. le!li!biiU, :i!.:~.. Ii:!e![li ~ . Dallliel, ~liJ,~,®dy,,6Am!!,,'!; [),oIi\lIIllS?


Padi:.ilIlI'I,:II ~alt~~<I,1iID. SlI1Ililh,,;tS

~re!IftlIil"flS alil;r;! NiIJi~es l .~ &!hal A. MiI!ilI~~l!.ar.K. C. Illajikumilr, .~1 •. P/J;'i. .~"w; , S;i!, ,'UOI ~::!'~~Ol .z, M. So, Mtl(~r,~ 01., t A(i:pl ~~ si[), ~l~>(i, ~:il9~1I), 3:, 'iI. A. :SMn~~n" I). lBirn'b!l:rg. Rfl'. M~d, PhY!l.i1. 'm~~ ~U'91~, 4., It MIJI!~. Si!mICtmd, ~}, rtlrhnal. :1,1. n~1l ~'199~J, ~Il. ;:._~.~I~.... 0. nl!;l.:i'. m'!C'd il~ l.hC' t~!c' TI"'i..!SSIc.. O ·So. M. I~, .~ GoIdbef'ge'f; P. O. 'ri'.a!1g. A!I~l.!'.~; lIDI~u. Rt5.. [i\-rj{ld t I,. .J) (Cmlmn·No!'! ..n s~g!1!.s. ;,~, ai3 f~.qII;I4), _,uoo(Jt :no ~l 200~lm,ioll '>'~~ ;Igoi. 1.., lit. IBrudI!l' Jr',. Ilt. MoI\ilM~, 1', Glnl~. W!i~~ \vhl~i1 ~hC)l n;p~3et'tl rituil:i,I~ domi!1<11TOd b)' a Po, P,..Ali;vi~iJlilll:!." .:s:dmao ~e~l, ~~n nOOB}" iOIarn~I~1 bfl!.'t1~1 3re:]~!IJ~!tlln;, IUidc}th("Il' am~~OK~ or ~ 7. 1#. C. W. (im~n,. S. I'lril!, SdtlJce 2111, ~OOi6~;Jl99ro.

~t il s gelfl'e'liI'lly be'em thought that lhefir!it ,dirmsa lin. QlIkk'l~' repltl!!::e~ marie arm air!: l1llle' i1'riaScslI.:: h (ililillilas;. @ither by IIl!JlOOm.p~l'irlg tfuelilil OflMnelii till!! m(!JiIl! ilrcnai)t~l!Jn!'ls suddenly rboc{Jm!! I!X,Hn~. Fi(!!).~ilsbDm ith@H~.~~1'l QYi3ny, in the Up:pe:r 'lrl1llassill: CIi1I~iI'll@ FOI'fI1IilIlirlfl ~ff N'!!W Me\Ii;ioo, arid 111i11~fl<alysl~ of oth'~r 1"!llQio.!1ail UIPp@rTn~l( i!mmblag@s;ili!!i~d limp/ly' thillt!@ tti!~'S<ltigrl Wol!S; ,gfo!ldllla~. :S:~!fn~ dli1~SflL!r ~~~al~~~'~ pr~~fi;fed lit:! this r(:tlfF!l~ i!~So1i!mblagr!lb~l~ng ro gro~p~ l~rl~iCMJ!iSllyI(o(l!>'jiill OJilly 'from Ih~ M~drjl~ l : tll1ld lQ~nIllOiSt. LipPe!r TriO!.iSStc ouUilde ,['iJ'tlrth A..llTiIerica.TI1I!1S..1h~lrt!n$itiiolil mall' Irei:r.!'@ ~lIU11!II!!d' ~~Ir IS. 'to 20 milHolli li@clr:i lind wt'!s pf.Obab~y diil!(hromous at. dirf!ma!'lt ipaitool~tUl!iliIes •


,;~. 4}, f\Li::hGMIJI!S

<!iredi vlded into two p~mmy
~~.which induL&ephy-


[h~ P~dos~~h

~l.Clo..'i.~Urs, ""Ll,li'~WII:;:hi~nl~" ,11:)0 t:1i"OCndY!c:.!I1.1orph:o;.., altdGbC' Omi~trw;lim., whidl i.uadllldc:

bii.~3.11 di Itl1~1Ii,ilrojj'irCD'flh5




{.aSlerplotlt.JU am! MtI'l,j!!I~t1Iir;~, and dinosaurs and Rioan"if)(mK lu/:,<: '" IUmlfl(!n.~~'i:, indi(;Juh'lll or;:t {.:J'. 6). By the ~~~~IminlilQI']~~C.J~IIa_~C, <III QI·t~e.~c Nori;m <Bgc (1" J'~~ dm;l)~1'l ~X9t.iP' ~!t'\i di.,~~..urs, P(~¥~~~f11,. jlgt,g Uw UQ h:,ls..", d:~w'fw.wtmpod"""''\l~~lbi!:lII:!§:I.i'; in .m:~.(3;.I.iOJlt') v"I1'iotl~ IlJpjt~ ... 1;:,tt~~l.i (If phyt(JI~ ~J:nI;:odY!olt~iD:!l'11'l h~;~~:lU;i,;'N;lq1~t ~11 11~ of [Ulit;1~1l"1i11~unrm~ is; IXIDl'ly rtUJcl(~IO(l!\l C\~ salll.ffi,. 1I,~W~f~!iilrs,. ·"'mIJ.ij~~luchilfll:lfi:'· .I!t~ od~~\r

~ho\lgh LH(", Tri~~c ~k~k~."I~ mtd l~lr'i:llI .'l'1 ~r;!~'!bl~ ';;1l1',) di~ribi,lt'i;'d ·w.rnkiwkk PI). M·~ t"iVitk:llcC i~M SI!IIIl,Poil!;.'d.I!JTn:: ~·Ijipolh~b... IJ~A ditlO~i1ll~ d~i.v~r.>.i[1tJ i[l ~1(;:L~lc. 'frilfL~~~.e a.llcrnh~ Mi1Qi"C a[Ch~.ic limmis were cll~lt'iJkt~.Ied.C'jt~'l'C[ hy mpklly OUIl::mn~cti~.!!.the IlrehHic fol'i~s or' 1\:.. ~13Cil1ig~hem qll.~Cik:liy !llld op'po:t'I.!I.U~i$lti.Clllly ~~:l~r Iheydlied Olilll Ib,· otiICil.· ~"I."IDI'S. O. 1. 6, 9, lOll, rnlher dmll bji m~ gr,ulmd lu'OOe,.~C'.~ eeeo~:


the IHQ


dil1'os;,'IUf I..'L....a, limld.l!.lldil!lg [he

n.""" I ~,,!.!!ri~h~n


r>lI"~~I~li~~S. Add~li{l:l:mj s:I}C(:imells ''''«'C fo\ll!ld 'iNi~h~n ~'crnllbct or·lhcl'!(I.~(ltyp;. A righ~ f~nlJr, 'Gil. 4:1.9. ,,11\1 ;;!I.!~n ,ijbi"" G~ 2JO, !l).g}' 1x.::1Qr!~l~ mh~ s.;Drll(,) i~ltiEi.\'id!.l.~~1 . d~ holomyp;_ Ad~iii9Iwl, m ~~lai~ri~~indlld!e'l> GR, n I. f~. ril;UiI~~lll k~fI. ~~Ul~il~ CiR:rn'. cO!H~I~tckdllib-:h! (Fig. 2. E to Ht GR 223. 'I"[']nip.l'i;;~a<;;~."'!J';jiIlaqLkOlf.!~l!!m (Fijg. 2,1~ ~o



h~~Ym~/mJm midooc-Iopti)'im:i.d d'i(:IU~d: .... Til,.;. sali]:ripQilmi~mlphai1d , omil1ilOlj. clil:irun diil;~li~ 1mowiii. in:·Oill~ hi!!:h.\oip9k'Ol~tihltic

K); fa,~dGR 234.

:iI. 111,;,,1'1)1 ('.CIllIipl(ltcri~hl r~i1'ii[llr. RelI~i'lii\e41 ~]~~~l':i.1i;j1lL e;;,,,, MC:Nk:o~hif,CUln of N

~miill~. H:i:itO'l")' {N.l\llJvlM.I) P-3::B?9, ..[1 OOlllill)!c<tc

sites of E~ro~ ami Goi:idWliiI'm lire ,abs!..'iill. at-l. they IH~ Wrmn alii OdlCii' NO:lil1~ fUUalC311 Trilli.~t>OC:
a~~n:fu~ (1j, Jt}).


logical 11tl,lacc ~nC'U~I.
IlJnuiI roo(!~l'!b'. ~h~ only c'I,Iuci:enecof diu~n;S<lW!r

!;~'as from lh~



1~!L~m~lrj~.J Ch~n1!! I\)~ F(~I1lWt~'Oll:If ( A fg>C'mhl'l {11 13).\,\!i[luiJl!l1 !.:\fidc!1(.1c rmlH gl~ ier cQmiw.mll& [he SOll!lihi\IHl}liic ..!u f'Q~il~ocom ~WI!!li!ll~~dd'!i(il lite Mi~~~IItf; Trimllli(;b,'l.>;ill din~ ~uro~mn]jll1."M.UV!.,l way [0 Iht,!· ~"Ilrl:~-:.;~ dinOM~IiJn;. ~Mi\'e~yal:lnIJJtI}' .ol a.h.~oog[Il:r:lillg of Ilu;: L:nc nifli.~...,ic. T~lbILlnru::r:slfll:lding of r~dy dlinOlSfli!lir

(l .. dini>.).Dl)

cvolmioU!bcgru~Eo dmllgc '" hh5icvcfi1I] l1.ew disC'O'Il'Cri~:: Sr.t(!:'ialtNI.~' Oi!JIJ·I.t.'1.,st.~ ~ 1.(1 ocmiOllstmtoo ~]~m. a5.lll1, ~l.ill.asITturulhmls. survlvcd lil11:Otire WlW h T,·iru."si;c; ~ re.l,pp:rJ1i~all of the North AubClI::ic .... diw I1ID:;:::.!llIl:~mrl ~~!,gb"CS1I1f!d Tri;a... ic dinusm!l':' ~hm s ~hou.g,]1l (f.. am:! () 1.'CC\.'.~lh.!otllion orrl[n:~IIfi.vc T!1.'i.5I: assic nmu!ihischfan (Ihlosrml's showed IJ]gl rnnst were nl!isici'cmiifk'Jcl ( 16). Here we ~liibc rossils fmm~!]c l:!~~~\'ICr CTr!il]lc'FOl'nl<lRjrm (Urpcr Triassie] ~1 yd~!1 QLI<Jrzy (HQ) un N~;w M!:01\,i(lo d('I'n1I~ ... o.n~l[inillg tt~i!~ ('arly Idunm1iJur ~;ml:t;d rr:p;lneel:l'Wne Willi l1J(oiiIDuciJ"nbrupl ncr 8:\ shUfl1e ;IS jJl\::V iillU",ly


diverse and rmtll;r~h(!,n ~vio.1!~I)'

ll.c I ~Q i~hl ~hc IO\,'C'f IJoOIitKlI1I)ffl~e !Pelri I~ccl ! b.i.'I..'W!l d~n~lI.nllbI'l1llslmil31'~O SHf#:(!Un~,: OlIJ~dOJ F{'i~~ ~h::l'Ilrel'ohhc Uppe, Tria...... IO~inlc: For~ sic new !mn"dino.~u":iibnl!1 di.no."'i.!m'{lHltllll~b~il1i1~h!r Illation. The :l1tlfC'ITcd NMMNU ~t1iXi~I~11 $ rronl ~ ~he M~U{~'Y S~b}i.d1i.WUl,lrry ~lt}h ,lii'!)wiitll ill! li:l('l Q [0 i..t1§,.¥I"luwm (.~ m%;on knlHvll .(ml}l rro!'ll! Ult! l\1~ iddk~Tria........ .Argi(!~~~il~~11 ie of Un..'1"'hid~ \'1i.te P~I]'~li(-cl r:o:r~'<'l M~'1I1Ix;g:. ll~~P'i,'nri ii",,cl FD~~l d!.~rilx: h~~oo. MCH~oor in thi,~area i'l of NOl101:ll "~c·. :tC''C{ll'd:in.~ A!I'l::h~L!rl:! C~j)(l 1~69 ::~"~<::jI Gm,l!lhicf !!!1d 10 rO!'~iil porl.!.:!~ 1~nd\~CH(:brli!1!C' b.ioMlrol!:i.w1lp~y P.Wi;.!lll ~9;';::h Dill(t"'~li.I!mm{l!'l~'h'~ Bm1.'On 19l':5 (J 7 19, .210 22')t_ niagll.m~i~.Di~J~r.;; rmlUl :LU Ulill('r d~ltu,~"(,'.P.'iN S~n,."no 19,9I :/JIWIJr.NI1{fllm IrJlIMi, ~it.;1'n- f.:lt s(1u~omorph:fi ~'XCCpt lilfJ;l'rrJtdcJ'l! dM~·mll"(!n... " L &P' .110\1. E.IE.Yi:I~OWlllgy." Dl'(mj~m.eUJJi, li'e!'!~, drj()'ll.w,~ in. poo;s.~ssin~ a hoo.k-shli.pcd :r(~mOnll bend! (Ol:'!l.'ek wmd to.r nm~lillg) ;I1l11ld IJjt'i'l'.k'> (GfiClCk I(rig. 11\.), 3. tmcrnJ CJ11~.l].ilrmt~on \ c:ulral to the \\'oru :J:Oi rt'tllIUL(f}].iiffl'i, ror Alilh:!d :S:1'Ci'\~'QOO ~.c·l1I'Iorn~ head repig .. 2: A). 11111 Cidrurgcdi pestemROH'lel". ~~~I~o descri~ l.init thc i[]inOS;!IMf procl:ll'· lu~'tdi3:1,~lIJbc~: on I:h.c pmxill1l1HI Imnio!lfD of [he' ~cl1lmr F~g. 2. B IIlI\i C), all. C'1l ~m,£c[lcrista ~I.'l\ rron~ AI"bTCmiml!,iIlCh.l~H~lg .i.tI!,!t'l'IH1rm. 1:1!(1!I:·(H.VI)C\, 'ouuplic~e leH fumu.i. ~]!t;.'Cinl£n libioW1bulari~ on the di!1H!I 'C'D1dof ~hc' tClnm." number OR :2 U~(Fig. :2, A w D'), repOll.~~ ill ~hc ,(Pi.g. :m~,J. pO:\it~ron1.OO~;!I Cvi;!-'t O'~~'Ih:~ di:S:lal O end of the: llibua (Fig. 21]). 311.d an astrngall.llIi\: collcetlens the mllO~l. Rllnch. R.I.nh Ilall Mu;"lcl!llIl. of f'JkmuiOJ:o:gy 'lGR). w]lhJ j}oslcr.im <1~oclldil~.Ijl. p.mo."Ss (f.iS. ~,. 1 ~"'lmins !)flwn.<~n.Ml!!arilJ.ll reIOlli.v~of dl1'~;mll"': a


UQ !ll~ cCii1tailb the

l.Ot:'3Iit}! illl:d ih.OI~~III. Site 3, :HQI. l(lhoSI: ~~M~ch. :tiQ J.\rriln ('ourU)!. New Me",ioo. USA. ~


~m.'v~ {Ill' e~~l Q~ILCrln\lI'!\.~n.~1;l di:nl'f;'CJ~l bill i ~~~rli,~ny O'\l~lnpl.lill';; <'I:-""'IIf;n~blng~"fi (T.d:i!Q I~, Th~ HQ ;'~~IT]b~,_geindlllcli;'iS~~icl~F'1Ioueill~ I~hyti)i>tI~ln:\ .:uld '~he a,,1:a'kUIIi'S. 1J'pm.JW/~L'; (·Or.O}i(lllllWi
lMLIS-eum &f IPa1.eoolOl091Vand De~rllliimt 01' InIEl~Ja~'Pre Sil)'log~~,lImj~,ersll.yol calil~n"I" e(!~~eley.CII9!t1nO· ·479tl. I!.I~JIi. 2~!!I'IIQm·i!.lIlJhffly EaJlh Ob~~~Fro~y, (o1;urmbf,a1 'Ul!li~\'e!,$~IY. ila'W$<!.des, INY 1()96.~. US.A. "O'i\!l~iM or
Pateantal.Jllg)" Ameri:c<lt MU~Uf1l ot N!~llJni Ki~tlO~, New 'lId, NV 100241, 1I5.iP\.~(amrillittf1!' INI E~illulimmy

il~ 2006. aud s,h~re (I~en our I~dd cliC\.. l~ruscolleeted and cil.mloliiied ncal'lyl3:llD vcrl.cbrn~c speoimenls. SmniUdiphk31~y, ~}e .II!Q ll'CstlPpro:>:~ itn:ID~cly 65u~1 bcknwelu: f:u'lllOus OI1.olSl. It."llfl(:·h ('{lrdQt-~I};'I:>'Qtnan)j, I $lIln bC~~l{ Ilw Snyder QllIrn:11',. .<111<1 12 m belgw llhe Cm~M!Unl c;.klU:Y I~rig. ]1~ I( rf). 11iIe5C :t()lW ql!ll!TIt.>s ~ll: ili1<iJll sC'fll!{;:d kilO"w

ell!ntagliu. ~lld also lh~,tfi~lidl U!·oocurrctl ~n NOiu;t]1 of Amcric3. Ue I~10 wns ~l'Cn!od to largc-sca:i:c C;';CilV31ll1lf1


~~ Ibiiii'il~.i;!jU(lili"!$





11Ji~~. U!II¥~~~u~!j Ch[ci!g~,. Chicago. ~L6{163:i. USA. QI (llicag,o. IlL. 6@6!QS, 1jJ1511... '~Oeo,p,afMl:~ma: of G~lo~tal Sl:ience~. l!n'!A':Js!r~ 01 C:01.or.<ldO,. Boolder. CCl803 09·
lit3mih (on1er!~m[~. Uifllh!rr., Ilibtqlriu,

~(j~o:;.ogy ID~p.irlmem. Fieild f!.!i~um llil' Naitufal tI.b'tO!Y.

I!J)A. ~R.u!h H~U .M'~oom of Pjlt'>!~ll!!(ogf, ijn~ MM .1lI,7sm·960:lL us.!'\.

"To wh;crm to~~~,~'pIlOI1~i!Ge ~utdi I(S·.PMI..




ce add~~~.[·lTIai~


REPORTS w K.};dinb~ rnnn ,l.a~rpr.tf()1~in p(ls:li~ssing m !lIIlJ!<I;'h I~·g.,",r cri~~1!!tiilj,(ll1bl!~"r~)!; ..nd dilTI;lr.; frO!ll!l .. I! ochlllr ~~~. dil~.{I,~<tlJwl1i!.orptq\'lin !h.~ ~I)~~l~~~ of :lrQ~tI:h tKl~h;lI.~k'.!t (Fi~.2. A1h!l1d j],), i.l!~ ~}rc"iCI~~ 0 r a :;hmp ri:dgl;: otl i.ll"; ;Utoc't(Jme"dlldw,gc of~hc dism~ end ef'jhc 1:C~mJ!r~fijg. :l: Pl'. ,m:\! " b!(g:!l; Ol\~ gill Ul<f;IInl~rorw,]l.~i;l1 cd,g~ ~r~h~<I4'l""rM~I~I~ {.fi~ {;!\~~~~!!f!Qrphil,,~ J.
DnwlftN'cl'OIl' ~i:!,I.'1,,': lW" S}~~~,p\.¥m(.l!il~hi\.~\\,j~.l .L('gC'r'J"h,,~i£Jr.~:;I Phylog('mHic ~13~yS~{i; !:;:0I1_~;~'liii;ltt~y 1f!1~~c di.~!~1 ";I~~, 'Wjtil ~po.~~~lliIQ~:i~.1 eresl ffi~,~~~11
d!m,rm~~Jl.;~i.~ l;:.lnliliJr tifl


'~I'\g.' Tl,w referred ~~b~nc r " o

fC!' !ill~lIOS<l~!fl'i/IJn>d

psclK!oSIJ!Qh ~J1S

tt'l. tt, .1.;.q

l [f!.,.J(!11Jf!Um


and llPOsl.cfDlfltc;:rall ()Oll~tl:\'~iyf()rKh~~~cp!.im~ I;)r!~le :1?<lSlCl'a.of ,u:I\."Cnding ptOl>~5 of ~h~

1i'albl~ 1. m~tif~pocls riF~1'!f! Ch'i!l~~ F(ln'rl~ti~f11 'Ql~allai~s ~nr~h~ Ch~rn~ ElgS'in.,New im~);j,,~,. layd~n H 'QIiI~HY, HIQ'; Call1jl]~.olilOll,~rry, (~; Slily~e~ Q~"HY, SO; (oe.f(Jpby~i$ Ql'Iarrry" reD; l~trij~ijBd f~n'~t Meiliiiberat Pi!trifi~d I~(ifil!s:t Ni1!lUoIMl p.:;!wk. Ari~o~"PF. S~e s,uppon.imllg 'o!'ilii'ie iIiii~u:rija~Unr(ir \lQu'il:ifI.@r S ~~dmen 'lfUli m:ller'S. Ta.:':o" MC!t@IIlO~lulrlda;~





x x





roiI.G!"'i~y II\o:rl.1i1~[\i'Otp~ioii llf '!h~ ('.ifil:l!i:rg~dr :mlcromt-dja~, CfIL':l>~Orthe 3~tt9~tllx. el'CaLing I.m run~~ruflll~:iii~~~1lCii.'Oh:II.cRLI!;it'iJoDl\'c ~hC' disnllt in an.a ,or lite: lilbfla, I Fig. 2. E al1,(~11I).The ~:,;tmtt<lllW: ovci1~I~S: he calb,mli!lIIl~l d~rs~tlly (F'ii1,. 2K). The t ealcaeeum r~~1.·hm£ul"lJ· ~n dDI'SDII iC\"".wjdoo~~; \

aM!O.I·M.;;\lu~.The di~t,il (;nd {'nf~h~ libi" of Dmmrml?l'Tm ;111ii0 po-,')~~~C,~ ;;{!1I(\il~Iii1~,dj"~Il, ~I".

I( roCQdy~cJmill1PliIar

pS~~'(jpmotus~pp. ryf1Q~'h(jrQ;K ,t',t)I('(i':nGWm Fl:iQt1Mila$UcJlu$.hama~iJ~';s l ~IR~,ilIisili(llifa "

x: x x



1)Q~~~iDrbJ W!,s. 21). IPh>,ID:b~11C'~ic i!!liiIly,~is, r] .. t":c~ DJ'tJ.mrm1l!l·o~!' a'l !.h~g~:;~~r ~<BXOIlI~o i..(~gqI1J'(!um ,rw~dJ!'ii.!(O\1IC'f~ U1.~1> d,II;!~ a"i;h\): ~i!1:!~F ~w"mlCrl iill cu.ll~r dim), .Uf

M~cnc~'I.~' o:u~d :nruroo\'viv\g

Dl'lIm~meriOJI mme# Sil,il'sallJrus·l~ke taooolil

1< 1<
J( J( J(

:-1mlro~lo'11hs IH!lil. :n (i 7)1. A~~lmW!ghnlOl'>t. of the' HQ !l~'H~ri ~" ,d~v. ..~ Oliniw ~d,. sev~!,d d;i<)!l!!In.~!ij~ ~rcm1;!!'!t~ ,...ell (llf tlLl\i~m ~I,WC OO-CJ1 l'L,'(:o~~~-d, Miull.iriol ,!;Ii" Ih~ SNe~1'JW~IS-i~kc dhlC!~!JWi.ri:riULnifu~tLlcl!1;"''' :)L 1%lmj~.:I:

d(!iH,~iY~(jR 224) (.Fig. 2L"




m,um (GR 115) (I:ig. 2M).min.d

,(OR. IQ5}(fug. 2..


pm,""l1rn.'llllcnlUir O~ ni., mill.eriall may be'

filg. ,2:. D~n®:Ililun:ilrllUlrphs 'ko!ll1! Ulie ,HO'. (A to D') Fe,mlillrof D. rOffj-eri (GR :t1ffiEl) '!Jemll.et~p, nov, in ,glf'l~rnOir (A). ,p(1)5l~rno~~Bi), 1i!~~lma~(C~~li\ldJ dis~l ,~[)~ vi~. U:. t® ilii~1II'ib~a D. romc;\rt'r(GR ,2ti:n 1111 ,i:ln~Erio:r (8, ~att!r:i1ll~~), pmKij mal (G}, and {If dislQl (H}'Il'i~. ~II'00 I!IJ A$l1'a:!lalkl~ah~amI~!Bm Q~D; romeri {GF: 2~2:;Uin pf(:l~ii'[(!arl {OJ i.i!nt~rIol" ,~fi.d11l~~ri~.r O(l'll!1ew,(l!.lo ~"):$Weu!w't.!s~l'!l<~ ,d!.lfilto:!l1' ~GR2241) inl mooial 'vil!~m, ilfil..nn {GR 2:25)'inr l-tlil..eFalvie:\'11 (M)~arid, I~no~mal're;mlli'r ~G,1i! 195) iF! pr,o~m::![ '(NI) ,~~d: poosl'er!o!ii (0) ~ew. (P~, Chfm/~5(1,uV::!(S' b~R's'mGl{j ~l'm:Ulr (GR2:t16) ~n p®s'lie;nm vi:e\'\\.a~dl (~) ~J)el.ophrY!l®idlli!~ropJ)d fDJ~d 1,bia. ffiibula, and 'ilI$~r!l!g:goocakan~l!ml(G~ .227) li'!~mlIml1l(lIT vi~. Abb:relol'atn(ms areas ~o1tO'l!.lS:ill1tfrlOllf'1ie'(!iai~ ~rre5t (a rnd.alllter®medi .. l. p,oc:ess {amp), .clnt'Brml1l!ooia I. ridg:e (amr~, b~evi$: rb:~1 (bfj, coilkane~m (ca'), Cln~rrd~l cres,l b), Cristal tibi~ft'b~l.@ris(,ctl. gW~"Il' (gr)j, la~~J~l '~m~~!IlfiN~lw:n 'lillIel!:~~~Ii!!'! gfQ~e (l1I1g), l'Iotch (0), pDSte-:rigr il!i(~J1ding !lr"@(e~'S (p,tip), post(!,J",eml!id~o1 ~ u~(pmd. po~!'erlJm(ldial~iJb~f {lPnllt). Sita~@ 'b,,~s.1 en,










clmlg,"l!!e, slllb~~;jll'lgl:tl~ar r~mD,ml head wil h 11n~.t nl(x_~~ or.mp~d OClm~!'ilu~~e'Qf I~:w.",irn~! sllIrmoc (I.~) lPig, 2.P).One ~""RlIglll!r;~f ~'ncm<Ill: !W~ crur«l. f~Q'II'~ro~~~ophy~oid :~ml,tOl:!1l!! n,lIj'iQ~ tibia" iz.!...
I~!w~<, ;'Illd
!ltil!f.t!,g:'lloc~I~,:"~l~t:lll~ (Glt


ro:r!t!~I);1~!$ fCrlJ~CC"

421') (f'i~.

2:Q ~\b11tt:

rlJ,Jl;i~jl1i of

tt:k'llIl,('f~~ ii~p:~"::I!a

codopi'ly~gicl .md !i,(!QCer"~Q."il:l.!r lhcmli{d~\2.5~. ;u~d ~11~1~'"A:;i~l~H rml1!~til~ !~r~y S~)!d\;;!' QlJlmy ;ll.)l!'l di~I'ilar:l. lhu!. dN,ltr.!tC'['qt (16, 26). Ali llln,II!111'

1. It, R. lMigel\ C, c, Swi~er !UI, Il, 'e. SePl:P!)o, ( •. "'- f~m!::r, A, I.W..PiI{n~~~. S~~t 26'1:1, 19>1 !:~~9l~, 2. P. C. Strl'!M" Annll.. ~. f.Mtr Pltl"'~t Sa,2:S, 4 ~$

~e~l!r,eflCie;s.ali'ld Nio.U!S

l1:ig1,iOlii> ~oc~o;pjJly~oid s,y 1iI!lj'IOilIIlcnpliy

pK.">C.i'i1 iill

s, ~.f. Bom~p.l Fl., jl
5. M, ~ lB!fIItr:n, C~~atifJfJ'


GR .22:7 is. 9. ;)iill;KIIIinillCromooj~aI p:t'Di::C:,:' l'm tllc di"!fLI fiho.lillniml i)"I!~llap.slhc as.o~i~(liingprocess

~1l'. Jll:lJIIIM!D1.2. 3~2. 11l!l1li21. 4. N, f,ri!iBf. l.lrm~ oj ltl~ D!,rro.sai1l1i i!a~I~DiI,IiJ~Mlfiily 8tooml~~iIl!l!i1,1!~I:il(lQi!iI.

J. M.

ofllt'C !1,slnl~lm;~Fi~.2QI t25~ , l1~Ci11Q a.~nlb~agc Chrul~" ourpic:tl.ill'C of th~ 1.'2.ly cvo'J.~ltiml of d~I1.o.~~lrlm~linm;(!ls ~n ~w~'al w<I!ys. Fir:;l,. it ,dacum,tnt:;: 'dlat n mixed 'l~licmbh.g'C {lrm.!~ d:~~IOO,.u:r";biVld Iheir l,),ulai d.~U!~l~rcDlllt111)h rchu~\~~il;l~dtog!i:!ltCf <!~{lll~
the Nori1!!l!l. Ou, in\,m1t~!il"'~L'OWn.>1 ave romld Ihe same h ce-eeceneuees i:n M:!~''l,!',d ether Ch~!1lt': [fomm" 6nn and DockuDll GrolJlfl l!r,)cll~ilie~ in AvIftlOll<l, New MC';:!.()o, and Tc,";;J:; (15.In, Thi)lf(!fult'!.. Ihe llQ 3s.q"'tl~M:J,gc (ilI:iI!IIOl be e.\J.p~~:oodby tnL'rJ;~. Il\lc~~ing, of ~cdiIIICItTa't~ml er oodi::posi'IDon of cft~J,cr fo.s;sls [,:v~'i~,~h is con I1li1.ltcd by om i !l.Cdin1.ClttQlo:gic3~ :m:l1Id,o;;lmIlgrnl)hic am.l.llysis of 110 (./7) I. 'l1lc W I.QiiI$SCInt)~3,giC also reinfo:ra:s the p:mcm thll[igJw-~lJlcolal.il,llIdc [twniL"> of WL!!hcffl Nonh Alneric<! dt~i'll' rm~l I'hclt~~h" r..t;l(lhIA~!lu,k~ i~W!n~!s ofE'U!1lre. Grccnl.mdL l
wi!}! n!!h(;.: l)lpic~1I1 T~i:o,a_,,~(:' OOlr.'!plJl;l~~n

,7M:~ .M. ], (!m1!oo,E:d .. OCl~m·~~ ~~ OI:fGl'~, 1900). 1,I!lL l,. ~H1iiji.,~, 6, MI. J, IBMltoo.ln ]I'I!~Drnru.aurloli'. I), II. W@i~li1amPl!I; p, I)O;!!~, Ifi. Osmd!!ka", IE!d~ (1Un~. 01 CaliIcl!IliiJ f~ !Iie~el~< (A, 'edl .;!,. ,004)', !iI~-]-l'i!, ~. K. ,F;ild~i'I,i'fIj a_ r.{ ,iJt A!ilt at elrnIM"~ lim tJj~ AmffiwJl SiJullmt'M:" ~S,G. IlJJGI!. A. p, Hun". E~ (lMM M~i~o M.I),~!!iIJmr NallIli~lltll>lcry, AI~ruq,IlE[q!!il!, o ::t9M~,pp • .:IO:l-'l:ll4. 9. fl. E. 'QMm e1 m., ,Scfffl~ Ii%', 13100 t200:2)'. s, ~ jl. I~~~ Q f1~~,fl.~ 5:e, .:!:~ (!!~a.3~.

r4 tUJll

C!;J[Koln Jfte PnJl'(lgt~ aid


]![]l M. ~ 1I!!f1trn, Sr:i~na' ilif'jO, ;l\Mi1 {·lL99U.

H. p:, C. SfJ1~mo.A. a;

A!HI(C~i.. rMNi;

Pi1I«mrai 1:3.


','"""i>1liItmnllfitllml \
Dr~~11;;1 OMl~hod1ra

.: ,


n99~}. 12. p, C. ~[~I1ID., A. Ill..Ji.l'u::;d,


1ItJl!bt•. Al'lM~l'!rll!: 14, ~3:

:H. J. il.'tit.}..


U9ll-"'~ ..

-'~ ........ ""~~!l1<IJ


S. ,W9 000.1). 1(;, It B, Irml~" !til .. G;, :P<l,rltt~,
19. ,~ t2:007),

N'o~t6, Am~il:liwl(i ~, S~ t19S1'L lI!'ilo1trllf. R"I~D.r'Jfu'Ilil~, .~~D '~200Il. I, Mf'i!ilHlt R B. Irmi!o W. ,e;, fIilr~r. J. ~; JiltJ14tet:JiIfl«.


N!t!ib[1Jt. JI.. Llu, Hist; Sia1.

Ifiig_ 3., PhyLoyell'l~li[ posiJlilon o~ [1 NiJm~ri 'g'~n, $P', nov. 1IfIliong .mhosal)lr~ •. A, Sl111'gLe I'IU1~' pa!rs:imcmioIliS 'W:~ ~~a5Ir~()'1,i'l'!~d :~Mm a ~Ifsi" m®l'Iy din\!! tvsi~ ef 26 'ta",.. d nd 12:7 d'liUa['ll3~s (171. .Jls~l,I:d(l~l!Idilii3l, Oi!'!i!ith i$Chia. Sa,u~op!!)d(D·

Sowh Am.C!ri~'ll.aJld SOlll!lAli:ica

in lockillg

til. See ~~~n~ 'rm~e;ri~lofl~re Qfi1irn~ 113, !,;, G. ljj~", PttJ~~MgI. ,1"g/1lM(limalD.!. P1111mJ~I'JI;
143,. ;~7' {19'l!®}" 19. !j\i.. G.ii'.a,FMl~. Mus,.N:.N~, Sull. U,,0116 eOO6)., ~O'. K. 'E. ze~!l1~, fi.. Ii..Hll(il:tJ'li, ~S,G. LWiIali' N.. i!iI. .M£l!l. NtJ~.

ro..re omilhischt.tln

d'inos;lru'S, Thi.~biQ!i!oo,!W a phic disparity

m~rph€!. aind mhcfQ,p!'Jld1JJ h€!w iJ@@n (O~~,ps~d

11m iJrelolilll'. St<llrsimik .. te t~.il p'J'e<5e\ntl:i~ th.e IfUJ',For ii~eggle~lh,gt do fll;),t ~nl~iI!d l!'!t(llhe J~,r.lIs~ic, Ih~ lellglth .;d lie gfillY balrimll;c!1t.~:i s'lra~.gwap hk~ rI1lPlr~ds:io.n. IliI,d, II'ndl~ltIf11; Olll1n,. Ot~neldal!'l; !Rhaet, RIiti.a~liant.
re:li::<mble I{)' .fm.:odap1tJ':i.is tJ{IJd!~illi. a Sfkw({unl:;~ Iike dillo.~1~lif'itI:'I1':1in l'liown :fmlllih..: same s;tmlfi~ k gmllhic ~mil (15., !7).lhc dentm)' slmres IWO "'ytlOllmnolphJ:cs wi! I~ Silesaurus: llCngte,1 looth b~"c.\ ]hatp.m],d~)1 11:!~c 00 tile j<lW lurllike m:t:!iU:rr~thi~n dam;I'... r.:o) and u MCcketk!l]gamwc <>il!1iI .~!!;; r!hLi' .. 'i~nlr.doo!l~."or ~]!C diC'J1!;;,1Iu,y U1<ll!:'''''''.~ ~l,)iI:td.~ th~ '1L~1i;.'fio:rl,ip {Fi~, 2L I. "illit'; [('\J;:11'1l,1Ip~ ·to j;1~JI:ll:rijJng.~~.mw.~lh i(~1ila~d ,deu'!!,icl(;... l:~ ~flbifll . b\lliSllII l>W~m Il~ of dtc Cl'OW':~I, ~'Jld a disli~lCI, ~l~m')l~\·ing I:h£bru;c ofdtc 8[ C~"U ~,l1mUlar til) Ihehcrnivoroilisl:ikc lCdh (]Ir othcr l,1Ioc'hos<lur:; {Nt. 2-11, 11rlcy :I~ck ]ihe 'aiSliru;:~ SI r.iil!,iom> fOotlllid low~itdd.l~tiporSife.HIW'UJ;; t\c~cih(/4). Tilt: !Ilium ::>har~ wij]~ SJl~(;jlml • :m. el.~iai'ged blll!v:is fo.'>!ID ., <1Ln~ shelf. \,'~dchils, openl:lh::raUy (Fig. 2M), The' :~cmjJ,tml head iSI:riangII.Jlm·in prox,imal "'lew O:~g.ZN) ami bea.l's:I smi1i:1lnoO !ch ~ientrnll),' >Cf ig . .20): 'both (I r these chm'."clc~ ;we found Ifl Si I(!,~,mlm~, ~i(..·fj~'''J1)/~!li,'(;;.'i" .~n.d P,<;(~'H/fJ~ £ J(J.g(l,'mchu,~,t J .5 ).

[1I.~lk'(;l 'l ['I'.'.Ol1 h~DYC-SC~~(! d~m"lic olr~:MiM.lim] CIII.;"i;l{27)., ~tna~k~l;'lilC1LI" cmllm~llililY mF !;_"k"Olog. ic~1 din:t::~~mlce:.;., Il~~rdy1l<r.c~i.(';i..dq}Clldcnl or Sllmplil1lg biases, The m 10 ~~e!il.lbl!~ge end ethers or \Nlt..~1e.riil North Am.i.'it'11ClL [ind E.UfiO'PC document theper


sistersee of basa ~ d iiniOS2Lu:tro.l1l1!Ol"(ih:i well into the L3~e Triassic, ill Argcminil. La'J!;eI7JeltJrr. J~4tll"u!l!:udms. and !?:H'I.{{la.t(1g~Jsudlw~ are conlilted ml~le MiddIe Triassic !(ut.dil~~i1n}ClmI1m"(:s .f-"o.m~ation 3ml nc'l'!,';I" ooctmr wi~h ,Ciinosmu'S f.l •. r). 1.I,Q\'<C:'iC'!:, the LaIC 1:riassic (Cm:11 lacn .;md N('Iril'm} Ibl~n.l~~ilfl;rrml,n lhcir MIddle Tl'i,ws~:crcl"lil\'Cs. c indi""ting aoontIllli.l!od dh'C'l1>iiTIcmion 01' thei>C gunul's, Til~ <I.gc 1<1~,gcnf~hlC!,c~Q IhMil:~,I!ld O1lf

gf'(~I,i~~r a,!~!I,;n1b~'iM~fi llii.;ir 0lJil>. ~n .md ...

North J\ml;rk!lOl 1l1!,.1~~1!.UU ~':{J~:k1C~inJls inl!ply ~.•~.t d1£iSCdi nm:smH:i; Ilmd,[ltmecl~oru~1ill riurl ,diill~~Lum· rn\l!i[[lh~ "(:tC.X~sl(jd vor at 1c.."t.1.'1 j1:0 20 OHm iQJI ~ yems;, TIleI\.':arct()() Ii:lv rudimal~tri,l:: d<L~ orL.lIl.C 'rritlssk ~ocanli.e.·, ~:o penni!: a nwn':' ]pn.':Cilsc tii1l1C cal ibmtiOl~.b!Jt h :Is Ilnlile~hclcs,<; c:l.c3!r~hm the h,1c 1rrims... iC' r~mJ~. n:;.·p.ltlCCI1~":I~lWllI;:;'nol .filliJrupt. R.nhcr. ~i WllS I:lro~""dCted and possiMy diad~u'O· 110IiJlS,. as c~lklcnccd by pa~collaliliIJdif!.1~faunmJ diiTC'L'Cn:ccs ,across late Trlassic !>angae!!. T.he appocmanc.e oFl1l1'! lim! IIIilJ{lSolU~'S. in t~~c l'!;.chiS·l.mla"lito :]i;~'lc~llb.l~ge ~ ,dong wilh~ilC ~ll'C (lcc!:.!!" {I rl\CUICCl'iof mlf;a] dinoolnl:[\['ln:tQrpl.1S Irom IlhcHQ ..t'i~(;nQbl ..ge,. '!.;1o;lcnd~ db!; I.r"!!l!·l~ilion lin'lC fWHi!
..~tiCI'iUbli.igc..'"of dj,I1J,')j\;;Lurpn;c:ut$On>.
b~l'!-~~i(du~~'Vd)1 till <HS:>~P11' o:fdlifi'l)!;i!iiiIDl'l> iiliil il~Jjtc;lIli."J! iiI'im.t

~!~ 1I'liI, U. 9!iu" 22. A. B. Hec!k.~IiL. :S, (l, liu~~,I!. M, SII1Ufwl'I, A. II: !llIm1. [, A Si!lie~M, N...01.'1. '~IlL :[iOl:, 6lrJdMo.oJI .56,30l (2005}. a F. [. ~~. ).. ,~~. Poakol)ru.!. 16,72.3: U!.l~6}. ,~>'I. !t'iI. s, 1>~~kBl.lit Ill. Irmi~ S, I, r!I~iM:', J, '1'1. Pilartz. L. 'S,. 8romn~, Prot ..fl. Sac, l,llrociM S"" .I!i! :zn~ R.@ost 25. It 5•.if~~I, f. RIllt.!.ill ~ .000mtl.u.r!Q, Il B, W~~nilpti, p, i)oo!;on, iii. 'Ckmrut.a, Ed:s. 0Jm'i!l:,r:II'Cd~1riiI1 i?rm" ~rk.e~. Gt. ed. l. 2~~ j).p.~l~l@' .~ii,A. 'B, HtIlkt'1ii; IK. E. Z~igl~r. S. G\. !;uta!!',. L F.• Rinrnarii N• .M. ~~ Mdt. ,f/lit Sd, .I!JliJL 24" 1;l7 ,t~C!l)itI. ~7. t ~ Si~Dr ~r&, lVUI!l'I~ 41g:~,389 GOO5~. .~E;.. JliQ'lci.m~~ a r.~tI~lrdl 'I\'@~f! fjjilldi!dl bit '~i1eIth.,noo~1. M !(ie(]~1il~I'If:(; !i1J(~et)' (~JI'UII 00. &0::14·1)6)' (tD K,!P..~. !)~dl ru"!!1~','h';: ~(I>fj~, R~~5'e~D~i'!Ie«!o"!!I Fcund U~ R,II!,1,1;~Ii~ l~r.l:.i~ IHjuii~a~gn ~o $,IJr4J ;i Bi)i:lii •. filtttffoDn iIllllmDrrilIl Q.il!Rt Ol'l'U.ilJ,ilR Irm~Gradullj!@ ~ll~~mip '('0 RJI,I,. ;lIfIgl 5;,pij'J. Il,JC'''~~~ey ~ltiillEnl ~111r.II~rali!;1~BlorG!;i~ OG IR.8;,I,), w~thiW: 111f!

l:L 1l·~.lliWim. A. n.3\r~JW,.).,It ~hi. iI'I~, I'q,~(;!t,

'sa. Bc.Ill,24,






5lalf "'00 r,J~teQl1~

~emill!ilf J!'!iIr1il;'~r!I,~. ilt ~!1I~51.

~!Kh~ II\, I~.


Bal!iI1i!i,W, !.'iN{, MI. (~1I;:11\,1t'!I. G, Par~er. G. ~mdl M, ~S'KIdl~r (l:I,f <I!os~~~jr)~~Wi~1hil!~di,mi!t,. r

Gnoiil. II:ilmdt\ 'Iht 'UI'1i\1~1Iy ,dI ri:'lil~I'OfIl1a Ml!UlilJl


f!:all~QJlI,~g9:l' '~IJ(I!,l,[»,P,f!1uii~n MI~!erum'of I"'~'w."l HI:!iICltl)'. ~~d ~~~ MlJlJ!uml'Il1I Nil1!lual. H.l'1UlfI!<I[l rJ(II~:<llfl!l Ill~ Flr'epill'lIlicn Q~ ~ecrmern. 1lhT~is 'IiJOO~ ~oll1l!iibwtiQ!I


5:L!lPPOlft'llI'!j1 CIiI'lriile Matell1'lal. ~.5(q~miI!I,a!W~gil'cgn.enU\rLl!ll617\!5:B36J358i\OC1! SCli?lli~1II If.~ 51 ~t! 54 Tabl1!:s.Sl! '10 S'l



~'!lIt.'Hf'Ol'i'~OIphy of

(GR') )6, d~[lIhIlY;,<! ,lUI C1~ill(!t'<WJ~lfVli!f l"·]/t!II,"..m(dt~', ~Ui

/{pril 200i'; ~a:r1P't~ '17 ~~ lO,1~2g.r$(ii!m~e.. U~HlS




G,en,etic D'iivers'ity iin HOlney
B'Be C'o,lo'niles Enha,nces Prod'u,ctivity and Fitnes,s,
~\!hh ma:RY 1[Ji;eiILes" (l!Je-al'iI('!~m..~~r(JillS PQllrn~n~'s Mthi~ (O'h:mfe'Slh~t cltr~ '!J,~ne'ti(iaUy ~btJnct. 1:he eU'ects olf glellll~'l.k, d~\t€~sity en (lioli!!Y i)rI(Idl!!ctivity oiIiIld I_():ng-,tenn fit~,~rs,are
~e iiI~tl'(;aHy d'il\il~r$e c~!kIl:liies, (15 paltri'lnm~;!l pel' (l)~,Ql1iy) fotllilded Irle\lI 'c:olrmi~sfa ...er d~i!l1ll s~~i!lrmShCHll glel1letn(a~ly unHorm (i()~®Il~e$ (] patrnillie pt'~ t ullony). Armrnll.l!ated diir'l',e-rem!El'silii 1lo,ragilrl@ rates, {,ood smrage, arid pOIPIJ~ilItnofi grmllth ~e'd to impressivE' IbQos~silii the ~HIlIe05~ (i.e" ~:rr,ofleIlirodlllctiolil arH~ ~\lirl~elFsm'!lival} (II 'geinetkally ,cii'll\~r~e "OIOlll:e-S. nese re'f>'I.Ilt£CI!Ither ollm um:ler-sta lIu:llil'lgi of tine: o:ri~lil'ls D,I pot)llllildiry ln honey C'ef5 <I nd lts r hrl~lril'5, ~~lrc;~klIllY perfO'ITlliall1~e.,
~!IiI ~n(l'lll,fl, We ~flQ'!1 ~hoilt ~WaTm$rtrDl'!ll


~ar~I.g., fOf<l_ging <J!(:livi~y, IOOd \"Im<lg.. f!OJ~~l'mklfl size, find weight :g;;;!.in :[l;'f,,~bf inl~Tm
\,,,Is (14). ~~Ii~~~~,~nf""l g~f!G1.k: ~~.vc~iuy im1P~i;,i!;,.~ ..~,.:l:;1,in.~~:;ur~oo (!5). ~1,,::~fQ~ ~olo!i1~~ di. '!.""...~ lltedica.!.Cd d1l!iOlISltoui. iJ~~ :;~dy ::>0 dll'll We

OOWlliic",ml1ine~h!J c-l'Ib::l"'ofml:.dliplc p1!lril inel!1Oil pmf;ku;;~.i""ij.ly.~~d1~1~n~~ ,ni~i!i~~!1 '\",~i'h i~~~rt'ii;;ft!l(:C ~:ro~i] c~l'~<l'" c!'ilia,~C(-d rt..';S,i"hu!jQ~od.~~~~~. Ihc 01' ~ l1]en:\v~tt IlI}~lMc diif~n:;I'i~ ill lib.:: J'i~fe"ll
of !!;l,"'li1cl:iCilll,~diverse <UIId ~hlticai~~ iJll:blbtm Mloni:cs, dilldll~l]ile ellr,l.y :>~~,i."'of cciOl:iY fm] ild· ing. Co1onies: '!'.1m} &Cl1.e~ic.alliy dil\"~C: \M~t,' lJ'iOl~tII:lmiol1~buill -30% 1l.10rreooll'lb Ihmn oQ~onic;"

liI,olley bee 'Queeos mi!~


hlilo W<B}'S; by ~Jrod.~Ir;;:i~,g rtlw(I'!I;luot1v\l m..l~ 'td~'O!lr..~.} <Inc.! 'l':i.!}, pmoll,cin,\; :;;~'I'Um4"'. \vhcn ;,u .f(ipf'ldll.(:tivc fi:;nm~,C' l qi.!l!;.'i;1I1} ,IUd ~"'"\'CHill ~:ltlWl,· j!;}l1id i:~~~'r1!:a(; ~(;-litlfllk,:;; lwo[kc.I·;1»' ICiIV~; tuu.ld. (''Slnbl:is.h a new nC,'l!I. SWflrtrlliulg, is c'~';II:ylind

111m p.11CIil10rnt."UlL:IJlof fOIYSlHldry, or illuItiplc 1'n~linilil wilili di !fcl'iiilit ItIlll!:-s by q:LI>L'CI]l;; " J ), Selecticn Jer polYlllldryis uDe;.;;pcetoo ~~C' it ~'K.nlc·s illIrnCO~ul!li~1 ~~lfIc~icdi\lcrsily, whkh I,~itlr::',drQrngil1£ ~cvf..~:5l1h3n. were 17' to 7&% hjl~het:' sw~nll.~nlgC'I'Cllt, E~h !!\l::ltcl,iC<IIlIy d lverse co r.ony erodes l~i~h levels of relmwIF:!.&i> itlllOl1g: rel:nJIC' ten (n -~ 3:) Imd a q,ueen lIml \~',L'" i~lslnnm:l1ta~!l.y limn :gu~!e~ I~yU!l1riform [oloui.cs, nathree {I r live' ofl~~ril~g. thoreby him.!cring ~hc cvotLnion ('!if mJ~l'\!Ii:'HiiC bclmviorIO\li.'lIlTd kin, NC\1Ci11hclcsl>, momiflg~ I(fig. 2}: p<liin.:dI tl~L"wilill l3on.lCoolli, il'h;'i.'C'millmcd wilh SU}cl:1Ill ~hun:~ l!n~UC sc~ of ~H~~[hl11n .Hdju~tm,",!1t 20 J Il.!!W; 1{49l = 4,1, f' = O.{)OOI ~w!c1 ~~[lilY'.u~dl)' L!P.' r~lX'{IK'dlyi~~ ~ !i.OCiaJ In..ttl.;, (.2) fHh,lcn dillHi.'$. llInd c~~d~ g.i,m~~ic<!~ly 1(49) • 5.7. P <: O.OO!{J!.I ~2 t J uae; ,~(Il9) - 3.2. po4 ()ol.olly (II = 9) Imd a qUl)"'lli~~!,.1mij!!;~'~-d \Nii,h a ;,!!:t1d W OJIU C1\:.!n(;·!l!~C d~ro~ in I,,\,\i~'ry ~IJCCL~ of a .~lmlCY l»l (FlllllS, .,:~pio..) 0), &wcwl hypo~tl;.,~'?; ~imilav \IO~U~Il~: of Ml~~n fmm a ji;iDl!ilI~~ drol1~ . ~l.OOO: m:! 10(4.9) 5.5, P <: OJ)OfH ~ n j'li.Ilrc: .11'1(49) 5'.2. P .:::. O!OOO[ <lIla 1(491 - 5A~, p « 0.000] I. Df{U~~'S\!rI\Cre ~k.>(;lcdl m ra.. .lOf'm from>! pool uf m Jimvc IJtoc1!lil3'Oj'llOScdw c'x:[]~"in l!M\!, tl:IC lX.'Ilel'i1S of M.O'Bcovcr; ,.r~I;,~f' 2wcl,,'k:;;· in.heir new :nt~~ :o1~~C, OV(;[" 1(100 imDdividtu:!I;,lcollected {r1l!l:l II drone<IL gfo.'!lCI;i.c!lI~y d~v"'''i!'l1ew'Ofk .fo~~ CQll~dOiWl\vcigTI! ~lilJ.;' COX~~ ofutl(hl~~d,lIl!tl.ismil I\):WITtill,g [iir)llli ~ow g~n~~I~,~ull.y d iv~rsc 001 onic:.!, i\n~k,pl k'iB39% more !ill'Ur(:C CO~!I'In l'(:S. To n.:p.lic(b~J,;' ~Iil~ C'XPI..'mi(.;I1(lJ;.;of Ihoo~han !ilt'illCl.i.c~ill)' uilli~bllln,oo~,ol]ie!i [m£lllli wittl~rl-co~o~lyacl:au('lIIlCll:i; (4),. A populflr hrypo~b~ 1(:~1;1,M]~!I'ITli(::;,. ;S\'II;lIT1lbS W(;'U'i C[('tL~()(1 b¥ 1i"H'ci:rl~oJ 1390. ± 1200 \,'CtM~ 990;lt; 145 Ctlr cOLnb per esis su~~s dlll~ gCIil:~liL'iJJIIjI diverse \\!O:rk [OK~ queen rUnld ·~kg 01' her 'I.~lorkiCr oH~rilTlg 11-1700 001(11)1' ull1cd wi!h food: ,f IC$I~ 1(19) = 2:. L P = bees) 10 dUbi~(!r in 11screened G'~ lor throe days. mllflly o;pe.I'IlIC more c:fliieierIIJ,y ij51 and, Co.li1se{juC'nl~ when:: Ihcy 'I;"\'([C I;"d :;UCfOSC soil:mi)iUl ,fld~ibilmn (JI.MSJ. Tilts dHIcrencc was OOl because rome IY.pl.'Odllcc oolo:nic~ whh a ~IUtcS."ad.VUimJifl'f;! over celenks ~fLe!kC(1:spaDe (gcltCI.iC<Llly d iverse ,::Iilild 11~oscwilh uum:mn ~'tC' l)(loll\>. ~:lowl;,.'Vcr. ~hellCliS to simulate preswj}ntrl~ng C'1~gorg;crncnl on honey. I~ru:;h sW:lI'Ill was sllb.sequ\:n~I)1 reloc.u.c(J to a llmifoJm ookmb had ~Ilcan 61 r I.~'%fmd '66 ± ('Ql1nic;tUllg ci\'idCillCC dmt ~enl:!~icall)' (beusc coler 3.2% of (mill'! C'Jt~pl,}', respecdvdy):. in~'leoo, it mli:e;pcrloml tlskslX'llla a'> a ,oo!llieCil~.VC tll:m comblcs.s huve Ihillt \"'<l!;~ ::>inllilarw thm pre'Waslik!CI1y il CO:ilseqll:(;:n~ C!r~ncre.1iscdfbmgniJI,g gil;,,'Jletk:ally lmifiml:n ooion!cs. do ,~6 1m <uoc!. liC:w!CCi b)' colcmic.:5, llall!!u:i}lIy 02J. O:Jrlonlfe~ were bundoo 0;] I~. Jhm~ 2006. d~ri!lg [he l\.~tOn'il, &l~""i~ ill £cne~iC'<lln}i,c!i:\',e~c !l;Qion:ies..The magliuru~C'~l]){l!1C. enlm!1o~1 prrooWlclh,hy of li~C' \'.... Q!'k ~lesc dH'l'Crcm.l""', :i 11gf(flNlh dlllJ.ing Ihe !i.\lrI,~rani!!l~ se'~SOIl [jJ). O~~~"r(;'S\1orac.rl1S\~.!'C ~n uivude tote" hili ~l~\'uril!(lcr!liubxi c)l-!~Iicilly 'wi~hro!lJJlya ~hdl."nC"<\' :nes1 :1:~tes.\1o'C d(!iCmucntcd CO~lil)" deInItial ~ \\«ks al:beli colouy ;~bli!llding is :imJ~reslevel n~Il!l:SS g;l! in~. sh;,c. Cllllsidcrhlg I'hal WOrlK :tbl'Ce:;;~n gc:nel.i(>!I~ ve]OllpnCl~t by rne$ur!l'Ig comb oou:Mnlc~ion, A hOlllCY bee colony rlropagrulcs ~Illg~nes in



r the

t::eI1~ r.J.le

hill ~ell'''!.e SrOt


III bi.:'t;'1'. :J[1.1l~, !Iud WUMl!A


'l::.'l\IOh,I!".~n.~~. t){~.i.. "lit\.;" (O!flk:r ~~Iyn~l,~

with gClleliit~llly llnifonn~IJ!.I~m.tilJ;n$,llJ.elore 00l1il'cvck'.d olT ,dieL'::! wC'cl;. s ~Wig. Itpl.""l(lc1 mC,!iS~I'CS ANOVA:F( .~,,19, 25.7. P < = (UXml l('Ol.OHy~ype}~1'119'.342l' = n,~.,9', P -c (UKlOI U.~!~Wl~: ~9J4,2" = ),1 Jl,.P ..: O.'tJOtJi R U~H~~r..,clit'ln)l. DlHiu'lg ~1'lC!lC()('tlW week (I r ~Okllfl)' vi" ctbei r fll'li:l year U 1 ~.l'1iOSI. {Iou, Ito~ w~[i!er fmmding, we Oo:!T1JWJ'I.>d rorngj.ulg ~,Uil'S l!llllml~, " m .nlt.lql~W fOod.~CI< fhd U!C.;olouy ~hrmli!h~!I!. Ult~ or 1i~'Qrlw',f,';l'h,l!f¥!il~!ll to hive 1"'.111: imu~ rllr a n W{li~\t.;ffi !l1i'd Fot (mly I~~~ curr)'ingp!ilIl~llI) be\\Ii,~~",r and di~ ors~an'.:u:in!1" l ween dim~.~J:Hrorl1bil~ltifm~f rundmnl.)' ~ilir(ld! o \N'ilb Ihe ch\9]~~llge(i;of S\I\:(lt"$f;fWiI colouy OOioJlies,~ (UK: ('{l~OIll"«!ill eachueeunenr, Ii' = 50 f f5l,lWidirlig, illl 1¥I:lnd,. \\'C l'()il1d!i.mlm] 9 IOiflg.te~n smdy 10 COimlpare [11(: dC\l("I~pl'm:nt of £l..~l!,)1icil~.1y p~il"S, per ,dilY IIJJl:',Q'lI!Jlhamllve om~sccu~:ive di\~C'~ and g~m:1icJI~.ylimmnm C:O:l'oi1.1.es after 3. rn,rnn~l'Igs. G;:ltetic1l1y diverse ~t:lI~o:nia;nm~I""




F'~g. 1. Al'Sil! of o~m~ (mll'<llnli. "" .SICW (iOrl-

pt".lik"111.'>.; w~th ~.imil.cdrcsoul:'OCs
:S¥.ii1 n:ll muSlt 'OOI1S~iliCl FL¢!W QQrnb.



,di;vers@ (.)

~rm:ted bygeneti(aUy



r":""ci'VC',,m'KI L'qlil1 l1C'ariT~g'''''OJ'ker.;liCI repllacc' lhc <lgtlll£, \'!,',o;r:k, :tbrcl:. In l4;I11PCr.M.li:',diIllUHCS" !lewly f(lllndoo C(lo~1fIe;> I1n~~ o;pcrn~ce m,dcntl~ llt'" I C;J!lIISC1!~Circ is i;ilmiliCdlime to acq,uil'C the

U!!S!'lUI:'CllS~O s!lIIll'IJ!I}utdl!.'SC m:lill!ilna.. Colony lomru, h,g is ""0 dli :tl'icUlh ~lalonly 20"4 of SliWa.J:tNS :sttr~
8e:fulliWor. CorMH '0

'ka IliIUy tim oolil,b a:t'e'a

netic.dilly unHtm'l'l (D) c::oloml'ie~ d;e.r iXC~p:y· ~ng11m- n~t s~te~ on 11 June'. !i)a~es; '~h,en g;~Ol!WJs.dijff~r«l signif1!rein:dijcaJ~d by a hQr~ izo IliItallillile (lop).

ilInd ge'=

DI1,pilTIJn~;ml !;]I! fljclJlroli!i(lolog:yand Ui!l1lijlllirr.ity. li!~ca. NY 1~:BS3. UrSA,

'"To ~~!t'Oi:'Il(or~spomdena' sho~ld be oi!dd~d.



9 i['!J

:s '"' s













~ !1l § '!!1' ~ '9

m' ~,



.." "'






'" .. ...


~ ~












~ ;;;I,

til ,d!






'oS; 10



1)1 diverse ned gcm::li~ll}' ~m~form coloules I m~;mdl tlt~~r !h~mdi!VJ,~9 J!'dy ~9mp"r~~{I!1 Of\"ITII:rk.~rA1QP~ ~ I~i iml.~; { ~~~t; I ( .~ = ~ P = (lI" .12J. 9) A'i, OM~ tlIlitmtl~ :all:~r ··l>\'im1t~ullig;" ihi;l~ \Wlci ,m i~~llloo .mdlbricl" ~tiod wl:!e.1'! ubuJ'ldmi I Ibm,~~ bil;~i~ll~ O!vailOlb!C' l-9!CI' 2J JlUIy). Al~hc ~l!!!!t
were s~iU s~:~~~il""r~y sized


this IroniC)' I~q\~,~. gcn~IT:CilUy divcl'S'eool.onies were ;Jk~'1)~ ~\:",iN as h~<!vy as g:~Il~gcm,lIy uaifbf!I~ C~l!9'~ies IWi~, 3 j; rep~"lgdIVil,~5!ltil,;" ANOVA~n~, ~9) = .J9.9. r < O,{u:ml t.;:ol(n],i' ~ypc:}~FX ~6.13,73) =237.0, P <: (M1OO I {t:b:uc)l; F(82, 1.:1:;1]:)= ~:I'A. p « 0.000 I (i.nlemclijon~l. Th:fnu!g).lOi!llll'he now •.~i;n~!ic,d Iy di ~~IW ,eclo·

11~:es Olincd an average of 0.3 ± O.'OSkgJdJ)' 'P'C" ~ co.~tI1Y ,mel increased t~~~r Mghcr' ~nitiilloolQtny '''{eight by ~!I]r$%~ W¥!~{I~ gt.'1.1~~~~]~.y \i~~~.f9!ln I;QSDlr!i~~~.~i,l~"d u!dy 0,00 O.Ol Iq~f,d"uy~t
~l;jkj!TIYhr:Jdi:~ICrcll.ji1~~hei!i.low,~v~'u~OI.~y ciM'I~i [ w 'by 1001~Y 1fi3%bcl'o:r~ resources W;:l!II~d rlFig. ]~ I;;Q~l~p<.llwis-o~(1 f ~n<m.11 &.i~j' \'!,I\I,"li~h~ .~,ui!'l ,d\l~i!l1g now, piiirccl: r !C~t:. II~~ 1= 1.9. P = Rom)".!). The I il~fIIl11;;' (if ft"liiJci s'p'I~kcd ~ t'1t:iliU:J1l~llc-e illl ool~,b OOnS~C:I~Oil in ,~i1~ljic.'IlIlyl~,i\lI::i'l>CMki:iiliL.~. ho~\'(;'\'Ii;.'r, eomb m'!..'9 ~r,enmliii~OO lin eh!llilg,OO, irn, !i.t~'m(;'I~mlli)! Im~rOil1m 'i,,'O~(lIb~"S> {fig. It Producliml of newwerkers om gerlctka,U:y d~\'oC'Iri1C eolon b )lm~r<ls..;.:ecli thai. of gel'lctilctl.~:ly !.mUbnn()Cilo\~I~,cs \vitlla in~h~ fi ~l month (I r,(;(lI(my ,ii!:!\!eklll~ncnl [tF:~g.'I,: re pe\1ltlJd 'nlC<l:>lI!rc-.I1 ANOVA: F~1.19) = 1635. l' < O.mlCH (oolo'~y Eyp~l: ",,'II t.~ 7:4) = 37.4. P <" OJlQO~ lirnL,!j~ 1"0 1.174l -Mdt I) < OJKJOI nnlC'~:aclion~l. Brood L'(:,nir!g bywo:rkl.:!l' ~ncrea.~d CI'll~t111W!,.i!lly


r'e!~lilmfng workers; .: 10'''111V\lQrkers ~~Irl)iing polAerl~ iii!1~d ,g;e;n.~th:all~N
~nJif·or,m (0:
OITII ~V WO~k~HS cal~ryijllg IPiQl,~ ~M) rCOIGll'lIi~,. A'liltl'":!iks

ic.a!lly d iver:5le (II: {iI'll

~meam,1: 500 of ,gemlel'•

IFIig. ,2:. Fior<lgil'L'gr~e~



imi~ \'1o:rbr:s.; 0;



mark d~y5 wh@in dal~Y





ingl.:!WI~~dly d:i\'I.'1!'l'e·t,\Oloni~. wuil Ih~I,,\~dI of

\\lh(:I/L,,"a." glJJ1,-,u.L~llUiy ur~HQ!l'u (:(H.oni!;\S
S~i1l1(: pcnioo





21 June

22 JUJiile

l1iig. 1. W.ffigt:lt {1I1~1fli~, ± SErlO 00' ~~ft~tka1lY lIe[~ ~Jl)~j~ lfinu'!' tn :L2j
r .~

![:lPlg. 41), CUU~~(~qli.\.'till:ly. gcnelic~lly diiVCiTSIi: c-o;l'onics h~d iar I:q,cr wcrkcli popu:lmiollshylhcClld oil" AlIttiS'l 11i1I1Cill12ti.700 ~ t~JI[) \'CXSUl> .53001t 2MlO i~ldit'l'.idur~:s er ooli:m.y:. p t U!:;;I,;Ii( 17~- 'iJ, .P< Clom I. TI1elC dilferen~cs in p.~\.:]'m~l]dhlg Llc-vclopIMl.:'nl Iike-I), 1t"~~llcd overjhc


aJfI;d gef:!etki!~ly W'lir:orlili! (dil~~~dI~in;e) ro 9) Ic,olon[~ i!lfle~ omllp~ng ~1e'!Y nest s.liOOS IOf! l1ll ] une. IDal~5nen grol.lJl$

~i.1:~maoomhi~!8!~ionr(lr ,m (>,nh~l!ltCCd i:~J~~[~Y(If'
!m~'l:liJ)Ie-palil'i~lne colmltCSW liOllslUiL1C'l 11C.!'l materi,,~. til IfC.mx 'brood. and to acquire rood r(gh'cn
OOl'!lpll.J:abk: \'Ii'()!lkcr p(}plll~ltiOm-) 1.!J!lc.1lil!e~lOmcn· ~ l!lIm [hat this len! w!li~L! pace or 00.1101111' go-rn,'I'lh" ..


.I·U _.I • ill" ',1\.,. 1111 wcigl~U ir,! ilridica;~Ecl b.Y,ilt

1!..I~li~lmro " "~lih'l :\119'


~m MOliltal L111l1! (tap)" lEach


colooy ..

malf~ the d~th or
,:I. 1ii ""')
"""'JI ,1111





1i .,


{."J :;;:1





i' .;g ItIIJ
N C>!


~ .-



,~ :z









..;: Li ...: ..: ill ftI Q,I ~ iii. ~ ~ Q rn ''

... ... ...

CL ,~ «l!: '<C 0 '0,







s.il1gl~·pmJlrillilrl~lll(.mk'::-i \\'t>Jt: nn!. ubk: ~ des;pi1(;lliJVnll1g :;;~milar OpjllOr:!Ulllilil;_~ ~:I.lcl' ~L ··J\.\VttI1il1rlil1g"c.'vcnl. Colony size is doseiS til)cl; Ito fiml~: ~ft~~r colonies liI'Oducc more drones. ]~flI'IiChigher winteli S111n,'~va~. and iSslIclil10re swarms (IoU]). H-cu'C. i.iillraco~on~31 genelire divershy resu~~ed in ooll~[d~a:bIIYlnore: 'IWl)~~ous <mG,:res(lurcc-rich l
p!L'tt I.lml. ntil,llttl


~meafiiS,±:'SIEM) or w(lrk~ (!lQllfd lil'lles) ~Irnl

ltii'9 .. 4.


116 .,-------------------------

d rol1!.;rs(di;l$~~d

pn:u:!lu:ood ~. g~n@ti[;1l [~ ~, d~v~~ ('II;:clrr<er~ 1.1; droml{!$; .), aml1 gene1:-

kall;V IlJflmOrlil'l (wOr~eJS:
DIdrolri~:0) ro~o~.
C.'l Prp\\!d~PIIIPll(!

A. u!~rmSJ~sof illdl,vid-

uals in


dla'~ 1Pl!!PU;~ool.!mlt~dat

:mooe ,on each


t~~s timer eme!!lig;edas

vat b€!hlf@erl1 dlat c:@fl'-

ri;ldull5 durirlg~fule inbllli""

~s .almdlile' next tI41. Hoo~)Dntallines (wo~





,ers: S11n~glelii'i.€!I;dmii'l'e<s: doolJle tioo) mmt perlodswh,M

brood r,eijrng dilil'ei\e:d :sigi'IJiI'iQln~lybeh~i"i

mlMLcs. whleh in IHrn aili::crod I.hear 11:'1Jile:>:li. e-. G li~ctUC3~J~' diverne colo1nics l\.'3rOO :sigliliI1C'1ll111:y more droJl~S thm~ gC:iIlCliC1lII~)'ll1!ifnrtn colontes bclbrc t .~ :l.'e;!rlI1l!ll cll::c'l!i~l~in &'p~enloo:: ~,ncall191(] l: SI'l:IJ 'iCI~US 240 -t H~9 droncspcl:coklll)' IF:"g. 4; ~" ~I.: ~119) = 3.7. ,r' = O,Of12 J.. L ~,m:~!!;er., ,g}~U~C~klll.lly di\IlC'J:RC cQll;mi(i'l'i ~~.~~O colk':C'tcc;i und] "lofQc:i: I!ml(: Ib~xI limll g~Il.cDiodl.y u:J:liih:m~ cdnnu1l,'lii .,>tnO !lI~J ::iJurvillcd~ 1!;:1im.AllgLl~tcold p... ~rrod th~l ~11lirYe~ and killed 50% of gClJiclk:u.ll.y mai:forIlll rolol1ries{Fig.. 3). The rel1lu~i:l)iinJ!l. ~noticlJiJly unj~br~n 'OOIOIli.C.~ exhausted iheir :lIOodireserve and dilccl by mid-Dc~11I100r;. whcm:<ls 25% g_enctic,[;'_~l,ydiV1i:r:sc eoleuies t;HrviVl:'£I~t) M~ytFig,. 3). \Vc.: hJ[J,yc d[!"IlXlI~l:fi.lr.Ju.edl ih~1 i.he ~rod~IL'I.b\li.iy and nini!;~sor~noncy bee celonies is en;I,IfIII1!CC~ hy iniLrncolon.ialr gcIN!~icdiv~,~t}r. Our daia conllrm .md esiend tl\Clad.<: to~\'m:d, increas(;Id ~:OWI h IlC:POJ~ed in shO:I.1.-telm slI.dcs orpO!yo.mdrollS oo~()n~ wi!1i ~aw ::M liupubl,;;TI. (lrpi!!rihn~s (fl, i). The bt;1~cr'hs of ii g,.;'~~~iC"UJ div,;;n;c ~"'CI'Pk-G'1f .. ~f.!~11l!~iQI]am ~-l>lioJ.Ici~d)' t;Vi:d~Tll dln'in_\l CO~O:Ul)' I





,.~i~l1"". ~UIfv:~\l3~d~rP(::Iltd!>eriil:i~IJII:y em


~~o~lt!lly accomplishing <l\'1]r!cty QI'I'lI"l:',~sin,g ~;t,_<:kg, Gi \len :5<i~~li~.u' n~liIlb;i;r.;(Jr \"IO~Crll, enviro:r11~~!1I!~!cQ'.f!di!ions... ,,!Ri I1(')~,'!~W~ hun\l~ (:Qkmi¢s h~lilt i;:opnlb !iIl .. ~¢[.rOI;1l1l1-d tiJD91Ji,;:, !In'ld ...i,{m,,'d gM~)('t t'l:llll1;lJ,;illis of ft!Qd \.V~K·D~t~iK~irV1;IiIk \ Ib:~s "\~J<"~ Orl\luJu~ ·of miUlJI gC!1iC~ic.;;!ni' disIbe rmlge or!:~PQII$C thrr:~hK)t\ls in cQrQnil.~. il increased the ~~I,Dbab.~litr {lir~m\,~.[l:g ~~umdi,.'1]t W[l.n,,~11' f!J!'1Cli~:mi!l~.;IL'l tor<'l~:r,\1"~ldl(!(r bn;md~~~.J. dt@:r.Hf]~ of t;'.{)Hdi~ioD1~ 'O\~T whkh Iim1gm1i ilb~llCCiL-d, ~*!(~d, oc'Cmitooin orWt::fi; r1l,,;'J;:ilCliJi!l;\d/rcm;~ii),lOJJ1.'crllrilboore.s~\m;::!!'.'!. Selectien lotpa.iyandry W(wh1 bil; 8Jmngif '~hc:g~JI.c~ic
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dh'l;if\;,il} [hal il~-.\l'~~V~ filii ,c;nIGliii;,~ ,~n.~,u~et,~, d~ t>up.l~i~B~.'Cil~,d. ~ik~1'a~iIlisrlil~ofl1ar'i~)' b...'i'."rhr K,'CI(~lilli.l~gik'. .. t :~mlt~I.~, food. 1l";'~lL;;e 5LLCoc~:,VC;j.'S~ Ofi~O:ili~~ t i.e" 1)j:o.tltliCtioill of dM:ne£. 'C'l:)k!Jty j.l;ro\vth, <Mild surviv.al ,. 1'I1LL~. we C'x:fI'X'I i!nl.C',It.~~ lull colmw fOlil!dlin~ by hOi,"-)I' bees depends S,1l) h~f1\'illy '00 r:aHyhlg fomgers. m~,d Ihe S;W.i1n DCseleetlon li:lVorUI1~poly~jJflda'Y lJol..'C!lIJSC inlrnooklcum!i.lilaa,kln Qlft'C::;O,u~Cl.."l>,Ih:is C(ill!~ ,e%p~l<lill. lin ~iElI~nClk du\'cr.;l[y i~nproves (he 'Prndtlcli,vit)1 conee Ul! wfthl[llhcil' belle'rill> unli~latcd!o 'wrn'~cr of the ",\!(wk loree asd increases ookm)¥ :r1tnl"S.'ii 11lnducth. it}' (.I:i'. ]<1 )\ ~ll! wlcL'c~lm';M O()CIn I'IfCIlCC dun ~Ig~IC' I:isky Ip,r~lt (!it' oo:ron),futmdi ~~, U ighcr oo~lcl';,tivc IlutKh.!C't iv i~yof gcncl.iCO!!lIy or e,l(nrem:e po~y.m.(bJ' in ,di h(lil~' bee species, d i\'~r.;.~ cijilmli~"N nmylili..'l !lO{1~1:,;'dill 11 b!r1J(U;icr or Rerere~u:e~ j}!'!d' IN@te~ Uil1tl!1l)\ii.:Dt..;ili~ l1t;';S,pall.5~ fn:J.!l!:1 "mr~(;r IIWPU~~l!.IiI. iii, (lmier, ~. ,PBlllnh, t.'I'lli:tltior'l ,af $Q.c'al ffiSM ~ion)1 'to Ch1!:l1J;li~l~cOlloi'.IOI1."-. The ~}robabill~Jl' (~iet (Oxfordi Uli~.f',r~~ O,o1I>i!rii'!, '199~; ~h<!!~ 'worker \lIJ:iH ,~~!g,~~in ,Ol 't,Jjo;.k 1'la'iool;'.ul ~ 2. 1\ S~lI'a!!manm,m~d'~,5: 5«. 43, 1 '(200:n. li~koo fC1I'lI. l;,o,d.!y M~~!QtYlu.: e.g, (5. s. .i9H. ... U ~o [ ~. III IR" ralJl:f. ID. II. ~rllil!;M.. Anti', lIUtl',iliW," SM. Mt.91~1 SB l(ltOC2). [" OUI,~~ ~lI(mU.y. i.\O~O!1 i~wi,!llifll~~lip'~~ l}.(luil il~~ ~. III iii. 'li;!~eJ. I;. ]. IPjje~di~d. Aim, 1(N)J, ~~'Q ;II!, W04,lld be l:'XpcclC~dto li9V(lwot]ker ~puIOlio:i'ls; d ueD~vjl)" <In'itJ~1 iflt'd gmwlh I1ID.~"$1j\ICI' tiLl~ anld ( I(lt! 'l(1!d:t1U'i'iMi~ 11mc.:-.. ~iilliillii: l~f~~ili~lie~LHy ... dij~

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'PD.l ,domains hav,!l!! I.IlDli'£i beBnlhllll~9:ht ·ID· ,dlll~~e'J'fnliO ,d~~m:!te h.lrI:r:tiQ:nal ,daS&e1l de<fi!ned by Lheir ~eptide,.lJJindJil1lgll1ref;Etlnm~e!i, We med ~rd.ein' mimlO!llmlYS and 'QIlI:afll~itrJti!ll\e~lit,mreuel1lu!! pO:LcilriM tiOIll till clhtlfillcte'J'ize~he!! ~indin9 ~e~E!rtii,vilJj" 1.57 Irlln)!iI'~ :PDZ dom'aim wilh ~spl!d tGJ 2:17 g1E!IiU!lIlilIl!-em:O~E!dl o,{ !li!!p~de:s.We the If! tri!~Il!E!dl iI mlurudlJ'lfIlainSAl!L@~l!lvity IiIlQdel to p'J,8ilid: IPIDZ dOIllilOilill!-p;epliid!! ]iflte;r,j3tticHI) physiologic.1I '!Pp'l!I';I'USes or e..\p'C'!Wncc)', A,UC1~ lij,cr>o):S th.e!i mQU~ pnJI~IJIFJU!l \¥llli all ojjI,((ura(lj' ~hat e))~@!~ds mlCl'liiV lafgHcilt@, '!:IIIJerimental il1!{es;lig~tiOfl) til@ .~!;]!.lhrcly,.h!,""l1hoki" ~m¥ be ~l Mgh for oo~m,~ of !lrote'iiFI"lPrale'i~in1~r ,llCtiflfiJ). tG(!ll!~falry 11)1 clIlrrool par,a:dig,m, PD! oom.ains d~' fii~t (lill r 1100 diis(!',Me I :!lp1lce, whiich, ~U!!!gies~ Inllt.lhey hi1l!.l'll 100sks Uml~h,~vi(llfl'i ,1fC ,cir~"~lh'fl;b' l!f!i1'i;).~~g r:rom eJlal!;1Il'£;irut&1lI!l:i, tnla'}I' ,tlr'll evenLv diistriblJlledlhmllgh:O!ltSl'ltwj!,l'ily Ib!i!!i!rll'Ilptijmrii!edi!c[oss the ~roteom.@' 1Q minfmii'le ',::rQ;5s'~r~e:lJlillily. We p:.r,mid tliJat Im::u,siIl9 on Itlmm@!i ;)I, w{l[kC'r"l\~~rl6il'l;,.nh~~ mul~i'p~c p<i!~r~linl;,,, i wOl,lkl (uutriblllC lit.) bz diVl.'iffli~}' ~flborllF in ,~,C'O~~ of ]FlWIaCtiOfl dioNiniS .• wl1tdl f,~d!it~l@$ ttJ~int@~I"(JiI>iolii!Jf ~lP~rfm@1'II tiJt1100 illil!l!l IfIlIOd@~in.g, wm. IP~aY'fl!lJ Qlr ]1i10"@1ts;ilflgl¥im:p~run:l: IQI@ iF! ~lfll.llr@' in1,l~ti'lJ!ilIi(Jl1s i!)f prote1i1fl JiUIIfIUi(JF!. Q!~ly.mblicr dlflndiviisicm. of IfL'borfll'l'llOillg, ''!'f.trt~r:s, A kcyadlvantag.e or hnl[£l~olomlit'll g.trlClnlC' d iveusitJ, \\lflS i;C, ..ca:Jed during :inFrequent perioo:; whcl~ fOcod l"I:SI(m~s. WCirC [plcnl.iful (-33 days d ~iril1g our ::>i.lI:dy)., G~lwi.iC'a.My di.WfSC coklni.C"S g.[1l,iuu:dwdght ai:,,~IClid\3t [lIf ~:(ac\'*dI~lll.J~e 01 g,ClK1 k<!~lyUlil~ rGm~OO:ltmk's (!ti-ig, 3), who;~c s]:ug~ gish Ihr.~g~ll,g I<I&CS. suggest d1lil[ iIJU<lCo.[Io1Jal i gClJ,!jk dh~eusi[y enlmnoes lhe discc!\'cl}' 'lI1dexplo.u'<ltion {If Ibod rcOOUI1;C~b}' won;; rbllf.1.1~, t".'l!">"(:i..Uy duringlltC'.L'iods whm ~-j;Q!Jrclo'.sli.;'J;:'O~i1c l s~uJdcn:I.Y"!!id .!bund"l]l~Y ,~v..iIIlMi;;.lnlm~o]o. ~i ..1 g~(K,'tit;: d~v!,1rnil}' wo~llc;1 ~";l;lJItlill ~itOOI~ l1lliPid m'iObalizi!itimi pI' riJlt!i~"t work ~:hwel:~ if. by
lJlkill)'lo:tiC prolcim;, ~. nu",:lHiOir by lil~~Ul'\':, C'O]nprl sin~, l'l(!~hintcJ;:iM;do'lil amlCili,tIIi ~yti' d)OJll\1r'll~ (/.l). On!: oi"lilc nm:s~ fllCqlllCn:!lli¥ cm;{'I<\HlIJCre" IIl~crn.ctij(l,n (_~mJhl.<:;"!!w PDll: d'o~ nu,~n,mediru.es ~:mJ~i::in-tlro~cin intCrnI.clto:tlSby bi:nlcl;~1lg~olhcC I~n!lini OfiISk'lt"get p:ro~eins'QJ il). Previ'Ol~ S~lndics, Ofp~lide~bindi~1,g sdec~ivily have I~bcccl PIl,Z donmill.'" i!1~odi&:rulc' rUlllc.;j:(1ln"

~lnC'Bhu~these 1'I.l..~d.~Qouldrmt hove been gUXlru!e.f, ACI.iJmi re.."POilSC thrcshold.~ of workJi![S lire nol wcH docmne:nlcd ~20), ruad Iii. is, diffictilillO know ~IOW Ihey mt: relined to~he p1.UdllC'~i\:ily ofindivid~la~s, and tlle co!m~y as Ol\\'hok For C\"lill:l~p~e. workers may vmy gellclk'a~ly 'til 1.lll!r:mme ~twhich Illcy l]oCr~~rm 3, tm.'lk o]~~ thciumpollsc thJ\cshof,d iSI'C'Jchc-d Oil: ~hey IlfI3IJ' no~,bc"goOO'" 3~msks. Fer \\'hi:chlilcyhJ'!,'c Mgll, rlm.::slrolcl:s (i.e,. IIlty ~ac'k

PDZ IDlolmlaiin IBi:ndli:nllg S:ellect:iv:ity' 11:5 Optiimiz8dAcros:s, t:he IMII[)Ulse Pretenme
I 1

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domain::!! l1':i:I.':~);:rI\Spf(!lu~X-~--COa'[I (5', i). .Mo.. ~e","!Cent ibil fom1!:l~iolfl hils :>t~~~ted dml th~sc dCS'il~nmkms a!'CIOO NSlrt.t;;1ilVe anc! :ilO tldklillOl1.al d:il~~ h<lvc boer:n Proql~ (8, 9). TJ~c W'C'<!111"'1, dQ:I1\i]im, r.1n ii];!O discr.:clc cill.cgori~s" h,'{)lwevcr, I"ais(:'s qllC'stku]sJhm.lt nH~cl,iQ\~lail, ovcr~'ii1p:: Do',n;mins wilhinlhe· $,n]l(: c~,~ are :r.nore likely 10 Cl:o~~"rc<!(t wtth CiICh o.her'~ ~ig.mlds. '[:0

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9.liwd 10 out :~l!~ ~urlv,'o, T'''''llS-rnts, first, 0\1, ~"Pl~L;~(,;qu!Cl!1Cei'l <l!re d~ivtd Imlf!l Ui.c M-~r!oni!; omd '11I~!IN !iO't.r.mn'KkIJ'il, SCl;'i~l1d. O~If~ol!ll ~.~to ,un;

i~g<J!!nilw <td~ q ~t ~~ill:Q11 P .~ the :ili.li;l:5(i{ to th~~ dO~!.~<'Ii~l UO). 1l,is ~o:rill~ ~prn;:!~d'l i:ii ~:scfi;nl ~4r fn;:d~i~1g illi;iP~ icl~ th!ii hind ~UItil ;a~ngk dolt'i1aill!. billi. itis nm.itk~~lIy

le:"..'f~""'~1f;1~ is delined;i,,<;: the p.i.\iJb.lbil illl' (II" Gp,~


Qr r~pt!dQ8!lm~bind

~U C .term,uDal i:'t'.l!~dlit.';S of ,U~l(jtli:.:;t:: l~ml~iIK ~:IJI lui!!l, we ~'ynth\::;<;i~(j(l217 s!lac:h pcptide:r.,whic,h \VC Icnl'l1cd our "lfIlinin& ~I" (I!libk: S2} ,t J S}, A~dlO'U~l OOUll" lrnilnill'li: SCI, is not ~lImmllt'Cd re (ollB.'1ill, Ils,oU1:cb rO:i: C";-'\,,\I:Y ,PDl: !:kmmill.il pc~:~lliuoo !laSto oumin a bro a d view of bindi~l,g, s-ektli\ i~)'. To ir!\~iga!,C' b.ioph~iC'al ul!'\"m~~ii(JI~ be!,W~1i.,;'!' IS7 \\idl·bc.'~m\I~i1r[liZ ~r(ii11lQ,liUlI' ~h-c and

151 PD:Z

Oy5 ,217 1~,eIillGmlc

IPlrote~1'I l'I1Iioroor:ray,s Kv1.4 Ep.hrinlB,1i2

each oflhc .217 ~hmrosc'~'fItpcptidC'l'. WCl tl.'cvif.l.-..d <i!.Slmrc-~ 1!11!i!;t ~'U11!1Ibi!M,)'.oi,~heh!il;)ugll!llu[of,lmt'ein !
~:li~!'l).(!!frolyF; ,1I1~~1~ ~d!:.:lijayof n ~~N;~~~!~~ .~

lmiil..<"ulion (B11 Wi!fll iPr't.ldic,t~~tcl:I]Ud(:~iltg (r-tg.
1A). M i(:'roo:amly.> of PDZ ,dQ~]!i(I:blJl;'!;.. ere prcpaliL'ii





w~ll'Iill IndJividUll.I\:vc.J L~ or Il~icrO'Elter rl:lltl::1I. lUM p'rob~-d, im tf'i!ll:icilW. with a ~.pM soll.nh:m or e!tch PCjp!ide {Fig" IEn,~if5~, Inlemctiom.will~ 9 m.efm ~lIDrUCS(;I:!'I~CC lhal \'Io'8S m,t h~;ill>~ three tinlCs.lhe med i.m IlmlllC;«:cl1.(e (I~] rlle HlTIly were $l':OllOO. <IS "Ilrmy~~itivl'!>"" t 17), 11I1~P'l'OO.'!iS, ic~{.kfd .~30 ~ y ~i)l!I<!Ii~'!,l~V!h.'l1!I:! l!]S· i!l\'ol vingl2.7 K PDZ do.~mills. lit~l.c can be concliltdcd alOOut the 30 domain~ lor whkh. ~ID'ru:r.:ty p05ilhrcs were: £{mlld, For dotl1rulh<;U:h<!!u bm!ud ~t 11.,";i'!l~ ne rcrti{.lc, o il(l\~'i;1j,'c<t." ~hc ]1l,lhil i~y III) lbiind ~)o~hcr eprides p p!mviclel;, ~nlpor~n1I::II1J[ ~~ll.1il:nUII[UnOlI:These ~i'bollil1l.([~
aeuonswere :;,co~.:i IlS '"lli!ITlH.}':I~c!¥uli '!,,'L'.S_'·



Ialse posl~ves

h.ighAhrou~hpUl tliiL.'1:lt{)d"hore I >3I:re rror m~es 3&'\(}ei~lctl w0111 odemifynirl~ botli e p,()sithrl...'S 1II1.d I1cgnliivcs" Toe1.ilT~jinfll.C Qni'1~fot..e
p'0siti'VC'S. we reresl.CcI .arid qllamilkdcvcl)' aml!,)i p'0sftive \?o' ill~ a :so:l.~ltio.UiI-plmseF,P 3:\lS1lI)i (Fig. 1. O. which served 35 OUL" "gol'd Slandm:tr V 7). ~3,y UiI,l~S1Jrfu.g ,f~ ,m. l:i: ellflocluIIMh:un:s of P:DZ doInflill.we de1C[11~,ill~d Ihc di~<;,<;odalh;1n consUlm

As \\ruhh

' '... . .....•.. ,mOO9)
............... '

p:redf.ct arliay laj$i;l n~g<i!ti'll!!l'$



[P'[:i:Z dOliminl.~ (~M)




lor each of the 1301 mmy I)ooilive.~ (!.lb~,C' S3)"IUillcmc~iorL" IJl!!!lt shewed wll!Jratimg bindling Ol"} whh a Kd 0::: I~M'J I.IM wer.c l;:on,"1Idi;:11Cd "~lQ,'lil.i\'l;1l'i.'·; ~IK~~clhmdlrJ!l{11 WClI'C co~! ... 1'c!1l.~ h ·~I'-ll~iv..l'," [)¥! Ihe ~f~ {lr Ih~~ ~rit~ri~~. .. ' 85 PDZ ,dtHt~~\iu.:;; 1b'llJ~~dI <lit 11~.;J:;;1mu; pl,"WI'~id(;:~i i Itl':;: ~rilulli:[lg :"'\:1,. A.hl.'1ough Our K.d cWlI.ufl' was !li~'1, -i)ij'Y!!1 t)!nhc il1[c[7.ii:ClklJl~ h'ld oK .. <: 50 ~LJM.,
and -60% of :iilfllC[3Cli.Qlls hllcl ·fl Kd -c .20~.M (Ii;g., Sl)" Inoodlkio.llI, I:'P assays f!i!C¥ca~cd the la'l:ilc"
Qr Ollellllis:llr}' and (lil!ll!li~>ill B.ill[~' mI~fIfiI'Ild lliliiM!mty, 12 (l:!;f,or.dStlff~t. f,aI1IMdg~'.MIl, ,(1;1.138,. USA. 2[hei~aJilJmumtor MD~l'IlIlar andl ,Celli!.!lmr l3iDlOW. ~arlj'~r,d '1.!1,~!~rs;~. 12 C!;tfQr.,g S't!!ee!~ (oi![Il~Edge. 1M rOOiliit USA. l~jl~li!!lIIel1!


oonfO!,et a rr'a]i' I" .. .,."..../;
,i, Fl~~On~moo

~ a rray ra~se
(I (I




l:P'!D:Z dONil!'l~(~M)
:I). (Po) Strat,egv (g:r ()~J!strucriJ'!g a~~tddklm~i!1l, S'l.!lill(~h!riJty m.Q~el(il):r !ilIIQ":!~' P,[I:Z dQm!aif:l~, Pf(l~f;']1f! Imi(J'Qa~r~~:s wcr'e u£~dl to test a~l IIlCiS5nlJ~~intemdiiDliis [lleh,~~1'1 :!1$7 mtiu!Se: II?'DZ d'Qn:iI1~n~and 2:1,7 9i1lnom!N~IIHiooed Ile.plii:ie~" Airra,y pOi!lntivt'~~\!ere rel~Lm ami qDlarntUiecii ~' fP', t~.e~Rl:l~ oorreetilrlEi ,imay Wa'~ PDiSitiw.~. Th~' ~ru~ti!l!gl d~Ll were l!S@~'t'o train a preolictivt'. rnooEi'l or f',DZ: diomari!ll ~e!edtTI'II~ty. 'llie mooel highlighted pul:d'tive i3 mlliy lslsa liIega'lhtiil'§" Ii\Ihlch were te:5tEN:ib,y fP', an d !he '(j®rJ~Htedldi3lta ~lJ!Hll used t~, wetrain~he m~d~t An~f ~hree .:ydes ~Iff p~d1icli®liI. ~eslil:!,g,allildrelll'ainffing•.~e' rer'ined mol:il:!l \\filIS 1L.!5<m tID Ipredict POL dQlllla~ll-PrQte-]n nl'll~ra;(;tiiom ,a'CJID~lIe IfIiIlJiUs.e prot.esll!Ie!'.(B~ IRepre:5en~~livei mllges . of p~"leilll rnkir,oarrraY5. probed \ldiith~lui oreSlC:enJtiy l[f'be~ed peptid1tIs. Piilli. d~lmiJilll!> were s'Pott~d[ ]Ill ql!ia.drl!.lplk~~il'l i!ldil,l'l:d!JI@ ~,we\lts!!If '96,"IYell m1Cf1Ollf:il:r ~I_ate~.F(I~r \ltelb illiere req;llir,fld l(l·~~~r;lI'[J'l~Qdate ( Ii!ll Qr~h~ dl)mffFI:S,)Vh~ J\I;ld ig,ges (<:),5) show ~e' w~~tii!;lnl Qftn'e POl,gD:main, s~cts, The'lY~1~fllmo:!!g!lil:i Sh®~\!~Uf.1lYSp.~obed with (I p~of1ril:c~u:i'll~ Ipept,iide demrw hQml lK\lilLiI, (I.@n) <lndas:e~e(Ii!o!le ~epitkl.~ deriVEd f!'o:m~:ph"1iI 8112 (ng:ho.. ,~() I~P tQ~,~llio!i)WI!'lie~ (lbtaililMrtlW tlN~ array po:s1tij'lll'.isidefll'itni!d inl (9),


• '1111~ ~'II!bmrs (l)m!riitlmedl 'I!QJ)J1~Uli' loth.is MNik, tPI1!~@nl: addniS!i: Mi!rrnmillilk P~,~rll11la(euIKall;, i1!. ~mdaU SquaJll', Bwitdino 7Cm. Q.iQ~Jrdg~. t"N!; 02]3:9', 11J15:/I;,. t"~Mlill ~g1r,e,s; TIl1Pll!ll !.ile~i1!d~ C~~~1Q~. 550' W~ A~l!m!l~, Slj)rn~m'cl, (II" 009012., !lJ5A. §VO 'I'Im.{;llfll~or.~P'Dndtlrn(f' ~h!J!IJ'ldbe addrfilied. :E:'ITIai~ ,lIla (~dtlil'@[llI!mlslf!f,it.arv.!r,dl.0cru


ool!lmti n ,d~rrm, rliOn1 "Iloiher, {L~,. ~~I~c!ij\'hy.~ ~J~i~\,~d),TilL<;inlbmmfi;on .iwo f!9t 1;;lptt![l'~. h~... t!<td!ti.o~ll!!-I SM ~~,~~~C pqptidl; ~,~idru~s lhat~oIrnDrib~lh;: :lJirQ.rlg~}' ~, billdil~~ !lffiuily, :;.~dii i:ltt (_'~wm1ilw] ~jd~1t;., diClll1liJla:~ lhc~mCfel even irth0}' ,,~e impor-Iml
l~ru"!l howone



PDZ d{gn~im; 7%)i:lOIliI1lIln 1:0 Ihe Mffi:!.Il~PI,iQl1 t!);'.Ilt lhe cOl1!r~1MklllQI' ~l'!C'h INil1lidic po s illon til ~!~~Ih'l; hil~~Unil f~ .1I,~~i~.'\~~, i!1!~ lnlIini;d the H[Ii\ l\'~PSM, t",!; ~~~cl ~t)pl1,~ici fill~~ ~lI,."g!l'tu~~~n il ou'" ~.niiC:liOI1jmIY {Fi~. .~ dM~~ 1\·1, Prt~~,ct1,.1:1:lmt)'

OOI~n~ted 00 rDIl!



dQn~in~ (trnl)!fcS6}, We ~)'IlII1e" In'hQJ©~ ~plid~ OOITl~\ilXlmi"g

fi:'l<!:I'g~i!"C ~l.-·'f~~ini :;tI~)1'ot;,'l;1~hi,lm for h'~ndi!l1il:'!~ ' ;;rm'l
mh¢741'02'; dol~,liDq~~!llO~ilrMDSMwilhh.h!l;:

~.m~in d~Ding\li[iih~l.1~ O!i¢ d0ll111Linfiollli ,~Ulolhtit+ To CO!~,,!n;H,;it:(i :>'UUl~C l)tI(ld~~Ul:!!l ~llcJudc\'i m1a!iljl :W'[)Z dom!!iiri&. ;,....;d!<:\lclopedia. cv,1L1'1l!lIk!ll of


PriSM jili which~~P,li(le PDl oonmi i~ i ir

ls ~~rudic~ed i 1l~i1d U I~'

1:",1sc .l!1eg,,!h!;-'Sw~rcnss ..yodl C'.'1Tl1<·]'jm(!l]r~J~yby ~Wi,"1!1!i',u~ Fr, ,Uia. IhcMDSlvl ~,\rO!5. ~!mill'i.i'd i\le i.J:;in,~ Lhe l!Ii~~~Wdii1Jb!1iij~~tim~, l1i~ ey~k: of 1 r:md,iel~OIt, (;).tre~~·'i~'1i1llii,tiGil..~i~clIftlflljn'iii1g \VIi::: pcr.liomloo lihroe timl..'i'I"ll'i tmal,wc t~I.l.'tI )03 p:ro(Hel~~~i~ lill~e i1:~.llli VI!.'s. Qlf which ~33
(M,'y'~) were famtd 111) belXlSitivcs,. )Iickiillg 8. h i~h-"!IlIlilJiily. qllan,ti,tat:ivc intcii1ii'tuo.l:I maRii 11:. for mouse lPDZ domain.'" 11.F"vS. 2A and mble :&3). Ow.~r.l]t CINCfmund lh.u the ,lvcra~~ '!:Jg ndin; al1ln"
it)' (lftllc m:m)'

.rp a~slIy 'P6llUlL~ ][t"'-~}li,dIC!,'i! in Uie!r.!rniI1J~5t"'t.uld so 01'1~f" Sl.rillg(;U:l I~l of QUl"lIIi(Krc]:_ ~Ii~owl,. 493 ~.c-w il1tCl'<I!Ct!ll!il~~un~~n59 :"(1n:i~o:iA.'!(iIe'liQl'ii~ 3 \v\''i'I',;:~dl;lt" or~1 srl'lgl","poim
~"'Ci!'i;; Iml. im;luiiJoo


QUl'i1ti1dcl rrudielM 4!,R%.~J37~ofdtt na,.;:W ii1K':rat'!li:tm~~b'ftti 83'::1, {26S0) of lhe ItI~rl:il1t":iI11!..'_



IAJ,II 9j.~1'l' :>


n} or


lio!!s WI\c~lliJWaS set kJ .5 i~:Fi~. 2E~ wil.~ ,::IJIIMepelS:i,ll1.Ve1lhl~posill.h:~ (TI,JfFP) rstlo of 0.63 237.G7o(», The 1,",PlFIP rntio or oiCIliIf"mociCl predk:l:lQll~ c-.xcceds, b)1 a ractm ef more nJrlm~ Ihe 20 TPfFP maiQ or:1 Ba)'Csim~ model: llilm ~1~t:cgrnlCS:

where ~ is 01bi~'!cli.nl;l~ool~. A is OMl~nditO!!lmr 1P:(;'1FlIlid('.; ~(\i\1u~"il]~~ {Afl';~ = '~ ~fth(; <lJlJ~in~D"t'id 1U ptll>itJol1 p ,or tlt:t.;p:::p'!I~dc is q mId liM .,.(] (I~hef\!,Ii~lJ}. ~u!C1 is a ~O(1ol'i!lg t~lm~~hQM. flc .~ ~e'Ci

~ocucl~ d(l!!l~,~in,- To !1.>lW!e l.h>l1~O'l,Irmooi;i 1'ocu ... 1;..~ POZ: d(U~111~1:I ~~I~clij.v ilY. \\1(,> cOli':ln~iru;d

1.1:)be 0 fnr ~",'(~I)' po_~i]ioJ1 !~ m]-Id 'L,l\lel)~ amine acid q. Thus, [1;,['4 ls po~it~,~''C PDZ if ~'~~'fi'!f

domain i pucfclS IIjl:1iillU} rrl:eidq 91,po:;iilkm P' mooc ~llffiililIhc olltcrPDZ domaiu~s. Ilcgrllivli:iiritprercrs il Ics..., mIdi () il' .ill,has, 110 bila.~rela~,i\le ru!l~c ,aliter domain.."'_ 1"0 tai'llcu·tlilc!hllCS~aold ilIIPpoop:ri;tJ~cly fo:r

each do:nmill'" Vl'C' dl.::l'irlcdlrj!o he tire mih r~'l~c~~li~11J <jl~',~ tor;mU 0:]' [~~C I'i!C';Plj,d~:o; (I!lil!f III !l<~~f!in~~c~ 1.~~<!Jt ~mMnd W PDZ dn~miH i. Ernpiri.cJlly. \,\~C ['(lu.II_lld 'dml setting m .5 provides a :Qoodb:.'~;~lre lnc~'~'IOi!Il t.JJ~l.'1l!O'Sili"c Ined it.. ~i[1lt!i and Iib,l~\-;llcgati\le ~]retlic!i'o~l'N. Bl."t,UWl:SC lillis nl1L':1dc~ i~ cl~gn(ld 1(1hughlighil !iI(;~O"li'llily ~~~m_~


inrom~tim~ f!tHll tw,tjo l.u!il~" u I~ ~$ll'W(~ s'C' r.·d"'~~lc!:lmi'\.'~'S sl,jg'htly ~ow~r W;!)Il:-> ami)! t~philiwiil. Th~~li .~lI'ib11' 11:t.:hrid ~"":r~ri~ll!..~.~hl nd twO ~·~tf,[;."'l'e,~lc i!l' \I i\ll~ . n Iri.ons (I~' bi!ldi]lg,~mn1iti~. howewr, o~~r~aPl'M.'lcl 'lJ'Ililll-do~\il~ ");;pt:ri~'~~~u::, &1L'C/umw1Y(,,,,w (,'(!J'i~l cOI1:..... idcmb~~ fig. 53). ,"visia!;!. \,vhi le I1fmill!lili~~ing~h(! .~rnl~c~ru""'1lO\"j live TI!e' ren!ll;,'d~mclc~ ~l!Q¥fil»'ln'l w~~n m~ [J)1i:l !""JOO;U7~I'Wc "uriinl!n"e~he ;!('lCUroey OI'Ol.!f !l\iIDS~'1,11 i~ f{'}Cl,I:;iI~l <Iffila~~>d fwnily ofdmmu!l.'\!".!l!!1l.L."f .. upolllo:l d,~!f~ N.'~.~\'il!h.. n!(!·,piOfii~i'lll; 1I'1!1,\;l of mlum flU !l lm1o!d cQ!~uemi(:ln OfIPto1cil1,'" 'w~lil] Icl~ ..... 96% l~iL ~'O'n\i'CU)l idl;"vui,~t,;" 15 of 5]6 :r-p!. 5 IWl"JJt~ p:ropcill.i(\S,. TI1~$ ;lifgU(.'S !\I:m!llg~y [(If II S%oOl]fUll1~c~d 'p{l!>~[~V($) :ll1d :1 f<l!sc;:·~~itii~\C rJlJiJ of Wnmilie iblln ~cgllll":l:1~i\d ,elfmtfo ~_m.cov..;:rp.tt1lcilJ:l!I~J (ilpp..;:IkIS f~ui:[j~:C[fle~ioll, 186 of 1229 lOr prot!;:in ~aucmctiQG'IS;by fOcl'ls:nnlg em Hamme> of FP·co:nll1liimetl m:gm.ives} when mn!> ~D to 5 (H!!:. 2 B ~ The 11lI[ilI~leICii'S ornhll! M D!SM are dil'ptctl.:!d il'l~:eI:rL:cti(m moo I1II1':s., W~ also ,ohSC'.r'l,'~d apo. ...hive oCom~~lI'iOllbeas a heat l:n:JIpin FiS. 2C and 9ro~~~ndcd in, t:l1ble S4. ASi'IJ!~icip<lit~"!i. l)Qsll~on 0 docs nm oonu~lbu~e ~WC\O'll (~Ie nl!.>cldJ Ollt'pU.t (qJJ} lind binding <Imnllly mg. SS), We rbumi dml ~lml[1thinlg over oo!h ~rl{lJlgl}' 10 d:i ~cri~li n:1!~vc ililll(ling. but ~1C' four P[}Z d'om!!l.hl~'i!ld ~mim) .tcicl."; iSli.Qb.xL:ilIJniiillly eonOtl1~ 1]I(l~~,t~QI!~j1 !I;lDlltt'iblilc :;ub-.'OUUI! ially (!"ig. 3:' t uibl!ltcsl.o tIlICilCru:rJCY anile nrocll::~.~Iflll!!llhe To c.x!:lliC~,b.io1~IlY:lllic!lJ m.oolL~cs out or the ['iCSu~.ling lintcrtlClioJa l'lCt\\I 0rk.wc: dcsl.gn.cd. a TPffP U<llio by 44% O''\''C.l''l~IC :unoclcli OOIlstlUi:llOO \\ritlJ'm1!~s~loo!jhi!1g.whilc majll~ilJi!1Slhcll'II.fC" modi rioo version oF1:J'!;'C' Mnr:kiov Ch'!sll!:r<ll~l)l'ilhm f],;)' ~vlo._rcd~o Ihe special s~r~;;)Itirt!l of hip<lliIi~C' posi~i\'~~ mll!.: (,'sjijcn!~!ny na~ ~~~l~ ffvg~.2 E). J\I1o(iil
Unl!:Ili1m Dflhe





mmlfl)' 1tI~1l!iI~


f!l ooumun




~~f~r [0 il

<l!:s. fl IirilillliaO'Jl'mil1ll sck£~i \1~lY model


in,lo OC00t1:l11.Ihe: I~ ,\,'(;L" DCi'llililf10]'esidlles ofdtc' peptide ligpm.rl: po$~lkfm; r -4. -3, -1,11 •. ilnd (I, Even \\~i~l2:ndompoJms ~bli" ac~ .ammLin. Ihore is iUS![gj;lkicill info:nl1!illiol:Q c
Moocillli\:cs W Irojn such ~[li~lgIHJ i~lCIiISI(milJrllodcl Tonvoki ovemtdng. \1I'C imJ~IeIDI1C!nliCd SI1t1Oothnlig lech,[I aliquc; Ir~wo PDZ cklHmi!1~ bind '!5hllila:r i:'i1.!hs~1 of repl,i(te~" il is, ~~Il:;l~J~ to C.xpCCI Ilml. Iheir


(12'). 'flil~,1!~[!~~:1!11 ~>i~Il1IUI1rt~1.l"llI:11(;]01]:] 1 \,,'.~Ik 011 U1C'~ttLph.rlJnld IJ.flIs.cd Oil lite lobilervmirnl ill that mndmTil. \.VllTIc~[cn.dlo IJc onnlJin.oJ Wil.iliill "li~ht clusrer:s" Orrrlod..~, lhca~gm:l.thiml.irlcmi Ikd lour ti!hlh! ,ChliSILCrs of .PDt: d'obnf~in$ 8mlDl~dr b~lfIdillig lPa~I1G'S,tFig. 2D)., For c:li:.ilmplc, [lilC' cil.t1udJins i(1.v,gJhl jm~cdon protdlll.;;) chJIs.tcr widl 2.0-1 am.! 210-1. whcre~,I:hc N-llmll:lh}+D"H .. "\P.wlic

or Ih.:

,\.,':!ThCI '''''!I!;; dcriv(;'d IrmlIil !'intrnl1.hunlg 0\"01,;'1' PDl. dOnimLin~" billA.S1l1iOot~!~ {)\'tr1un~lillo ,ncid~ rllil


cN.clude tlie possibil hy

Ihnl. [lte mooel pelio:mmmtcc '!,:Vj}S dlilc 1:0 chmtce OOi1ll:JmioH. we pcni"o.rmill ii ]r.;~mdA:ml~'l8~1oof1l tCSI. am ilm 'I\..hich dlC in.tcmctio:n d~1a were' shl!l mcrl. "rille [(sullins nx;c'j,vcr opemlins. chllrnctcristic
Cllrve W,;]!S


li'Olli Ihe


acid (NMI)A) :I"¢ccpwr :il:tIb-lI1lli~ ~SOrO[ms NM,I)A~2A and NM.DAR2B, as wen.fIiS sc\{cmi voll~c--galcclpot<lssimnl dllrumcls. C~tIiS'lCI' widl lajil,rl's m~ m.1lM:l SiIDIli~;lr. lml~ss the dma. :SlJlggcs~, PS:D"9S •. till\. PAl?'. Ma,gi-~, '. M ~1~,2,alld Mas!-.3. o;~henvisc. Ukewk~c. ir t\\"{) amino acids h;ty,e ErU';OL!mged by ~fl~cl(l:'ic ~-ilroelllelil o:t' om shni~,lf ph:,¥s:i~he~uic.ali1f'Oix:rti~. tt i;;j ~1(;'~f!Cm~ ~J1.{ldd \\'h.h ~he 1.!1lI:i~ling"'.~~I.lbt.]" \Vl; \L~cd ~he o1!.b:~ lO'-lNlw"IJl;;1 ~h<l1l. Ih~~ir e.p·\ \,,,iU 1'!\,; ~imil;L!-, M DSM.~(I Pll\,10iCllQ whkh I}l:'~,m:lir,,'i in t~~()' 1Q~I:\.C n Simoolhiulg 1\,;qlll]IiI,':'i ;.II, q WI,luaiRlllh,'l; 1IlJ,.;;;l",-lP;; ~r ~ll{J~liOJl1C' ~ '~'l.~;hoiflhc 74 PDZ JOJl~i:l:I~ ull..,,,,'blc III b UUI~t lnlll'lt~l, \\'C Mln"Irey,,-d 3,1:,30.2 .pi;plidc ~(i" Jf!~,im'i!ile dlisEOJltto:;.\li,'ilhP'D.Z d(Ul'l<iill:l. cl~i\!ru]~(l qUL:JlOCiS cor.uc::;a.]l(~!U1d:il]g, '~o 'tihe C ~m]il'lli or a:lll :w3~ d~j]ll,l00 ~:s 'tl'le ,llflll:llliI:illl:g di:>t[unlcc of their trmlls:ln~oo open, :iiCadill;g frru'i'll,Ci.'i (.14 J. \;\'(: ~13VC billdiJlg "o'C{;IOI'll <!.cro.:ss theIULlinHiilg..set~ptidcs, p:re\'~OLlSly ShOWilIlllill. our (tmn<lill,~bfliScd ill, ",11:1'0 Whh m~~illli} <Lckls. 'we 1t"CI~clI em PiUeV],OOI::ily r..:pom1eCl .~ sc,ules~' 10 ~t~lheir physi:C'Os:tm:egy lilfthrllily C1lptlnllCS --t~5% Qrt~c prc:\~01.jSinvoh,llu1,l ,M)Z dmmill:i ,lhiCm l:cal rml~les. \vlwN: =, IS COOlskleg,\.>d il ~y I1CpOflcd i!ll!.t,:],H:ctiollt> d.C;1lCr~ph]r orh),ciro;illii Ikiiy, =:l ,i.,;fII dcscdpt\u· of (,/7). We lh~'I:cfo:l'e ~)in:'l\'idc Lh~e I)oodici iomi. ~no~ecllkllr VI.'CiSin :lind surface :JlC'd. and:) is <I (I S,.I ill) !PDZ .(JflmaiJl-~pl:td.c ilnt(!u.actto·~ls~, as Sl1ppla:ilcn~a II ill :Ibirmai.iou {talde SS}to ~'IC~P dcscr.~ptor of [XloI<M:it1(alld chl[ll'!c t2.l). We reduced till~ eqt~h,';]jcnt degre~, of ilret'dorn in guide fiu~mlll biolGSllC,lIl hl,!esligalion.~ US). We ~m~e. ]Dm.\'\C\'~r? dlIl!~ nol .d~il]WI~ti!;}ns. lij'ml are O~lrImx_ld by s'lilool.~ing over PDZ domatns ml.'d o\,er<l!lli!i.O m;::id.s \",i~h .. Gm!s.'li,Mki;)f!l()~ durinJ',l Gb.L~r;.~"d in ~lro neOl.~iy {l!C:(;UJf in \'iv(}, rug¢s.\lion (11'). To HI!rU'iCF ...~"'~" ti'!:l.1' ace~l!:m::y of lJu~ii1o{k!!. \l;\L;: ~~_OC[i:tI ,1~ "!l,.'Xl ~I[;~f''lf 4!l IM(l!tCnuns rm~'i !.'hel We \~ Olbl~ !.t'l~jn.dd 74 of nhe g~ llDZ

d~sClri;m:i1l m iOIl in,n,e O:1Ig. $6). ind k:milng, ~hc. c1:TectivoClL~SS,OI'CfLliLU:ai'liIil1g ~md•. SC!iii. cst II!lI\1hlS esl[ablis~ed, thm,ahe ~l1odc], aC:ClIJrnlely aJ.;p'tl!lU!s, :iirllib'l11131.ion ab01.u tile bii,ndillg sclC\:[hi II)' 01' PD'Z, dO:lll'ilUIl.5.we asked which ~}hysicooch~l1r iC<ll:~~jfl'e~iC1i !,'\~!;:h ~~{,n!~lUI~~~ p WI:1(;,S em;'!:! ~Xltii~ aI tfl!lJ fo cl~i]!lc il:' ~~Ittli.\'il)'<. For C"':'IITlJl~. if \,\'01: ~on'k <It mh~ 1UII!]11.0 ,1L(;:ldp~~.C!t,nct,'..\;; nfDi~1~3 (1111 ..I pos;iDiou "h w{; ~i:Jld IllilA ~lC 21J 0':;, <'!l1i: pnsili vdy oom::tmoo wid] Z.il bUI are nul ,(."t!!nre~.aliC'dwi~h . :;~('.r ;;1 (rig. 3A). ~fI '(;{mtm.~I, ;;;2, Illl! fla~.;;1 or ~]" OO~l.ilItes wi'lh ,d~:lCriI1flif1"lilvc bifld:~llgm posi~km 4 ,~hr[\,IlagH, (416) trig, ..:;i:B ~ ,,\~hc~s~. bUI, Ilot

or 2::!,oo~~.1l1tcs ..", iilDt dis~r.i:mill,tlli\!i:Ib1'I.lding at posiliC!n ~, ~Ol'",{UP]'! 11{'~o.r 13) ,P~lg,3C'l', 1'1u.::sc ~h.roe '(!,'I;,m;lpl:c.,> lliI."CC)iU:cmcs: :ingcm:ml, :P'[)Z dO:luj]jnsut:i)' m~ OIl1lhl\."C:::::. scale...:lhr di.~J,imilla·':1


bindfllg. To


ill~:On.l~!tion for ilH

whi~1ii ~li,g!¥-"Si!J.l thfil

ihL;: ~nllJonily of

n;lG!iJ~epmICo'in;C tl13[w~li(."prutlil'tt(\cl!.n

be .hig~,ly

POOZdmm!Jins at OJInpositi('lIl~. We cmlsU:m::lce " OOI1Il31.1I!!ionnlatrb: bC'~~v~e~ mQdc~pam~il!;tel,'S l:he ould~hr;~lM H~~'e :;; ~"t;'<!!~ .Clf~~ll1i l} ;;tI;;i~l~ (Fig, n J[)" Bttal~~ tlii~ I;;l1nlnblitioil ~.o,clj;f,t[inlijmlli~lC: t11nlil:!ili~ fllprtJSiTion Oi~, \l,1.c>JIlk.~.\'c 'QIHill~-cl t!ii~

P{)$fti91!l frmll our <ii.ni!~YSis il:l ii,'\~o:Td ~i;Ji;'Jing Qli!lf fl:st:!i~~~ ,\Iiilh .. nific.i;ll~¥ mliill!i nf,,'1;1 efl'~~~, To Wl~~'l1l~lld ',hi;" Q!I}'.nr"'-'!!lRlI~or pt,,'p~i~~ billdirl~ ~1(,X'ti,i~tym~ :,H. .\lk1m.d k:'~I. \,~ ,d(,\~lI1~ lrui.,.'!dih,.:';oVFCIm:ti[}J"I'~lfllri;':i~~ln'!~I(gbSiui!;iUbr-v-.d'll!.l ( i dli.;'OO:.lnlflOO<i~ion'!ll,d rOl'!ndl~i,~~~h~ di:iMib1.!t~:n or PDZ domain biadJng Ill'!!:!I1:li'l:'nC>lo,"S caa be h:lj:gch' cxplil.in1?(1 l~'y Ihl\,'!C rr.indp<1I.1 (l)'i!i;~, Ih~ ~rm;e
dc~n(,,"od ~y !11\,.,-,,!.!;!1i'\i,;-)1 ~<l!~.'~ !!IIol.A~ht oh!!'i "POZ Jo~mil~ ~I(}~iiv~~y s;p;J;:(l."Ead~ qF !.!he fir~i two 1l.:rIOL'l; ¢Np.i\;J.il1!:; -30% ~)ril1Cvw;ilano; ~:an ilrii: corrclJilti:o:n Jml!~iii..., ~""hi;'.re'l'5 d~c third u.,;i'l ex.

w'Jaim, -14% lF~g. ~mJ.The lirsl axis tF.g. Sf) can d~~i~gy,~1il c,m~]~l class ] PDZ :d~~n<!iH;~. ,''':hicl'! !I~~f~rro;l by p~pti~~\1j,'i'.~ !I ~!l~,i!t hydfOph~t~ ¢~;dtle :1!I.t ]il>OIl,;,tim~ _:l,rmnll~n{jl];i~~ .;:1':1:;1;. U dOr!.mil~~"\lI,hich ~Jl\i;: ~fem;(1 by pt.;'}J~id~~",'i:th h.~~, h)'drnpllObiic ~idl!l!e ,II ~,~ • ,I


.Figl" 2:. lA) (ii'glPhical ~:!l~ ,~f tlle 'trainmg"llet dalal. K!lJ ~r.of Ipp-(jDnrir,lfI1Ie~ ,~@STItii]jies awe Ir-ep;re~n!tf1m1' b,Y (:ot(l~. 50 ~M 20 ~M afifi'[JRY mght Ibtu:eJ. Array

ranqinq from Illigllil affinilry '~rre~'to tow

afl':!si1.O'Ilf.FIiJil bwcik; an'CI If,P~Q:rdi~~ed lile~1Itwe:si art shcm in dark 1l!~1II1l'.
ffrJiullililerical vatue-s,arJe pr!wijded l'n t:ab1e :5.3. (B)~jtorlilll;\iIii(C ~' 'mile MDSM OR, '~I! ITlIinrill91 .se'~ wi!h .m set tGlI S. irru:e ~o~fi~we$ dire sho\~nI in ret!,. f,ilILse ~~~~ ~J! gf,~n lli1!!~ li'l~ativ@!ii!il' bl)Je.i:lllu~lral!1ene~JatiVli!~'iii! yell.o~~.«() 'Gr:.\Ip,l1iocaltp,n'Sfrntatil).n ~;!'rthe M[)SM r

• amw

<: 111iJ1M





!I' ~


!il •






! ".IIU .11


gf~dl!;:d h'OOf! b1a(k,oo~lww,

d@m 1:0 diKriiturlli:lltirw Ibinding are

f,ositive ,c~!ltrib!i!:al!f!~


II~ .:1111- II -I

1iI!9\a.ti~f! (lcn~uibLlt~om$are 9radc~
'rmmi black t(l Light l~lIe" Numerical >C! r'E! l~rQ\lidooillltC!llrli! SlJ,.S-iIll9:l~ leU'eI' ab:tme\iil1itiorns 1m Iih~ amlmJo'ilidd jiesi~l,IE;"s ,i3it"e a:li ful!oLli'S: A, Ala; C. (~; (), Asp,: IE, Glw!: F. Rtie;; G. G~~ IIH,IHit I, ite; IK., I~ys; l, leu; iM. Mel:; 1M, Asna; P', PllO; 'Q, <G~;R" .Arg; S. Se'<r--;[,Thrr;V', ,

II ~. l1li1 II
, ••

I 'r"ull ,I

L .... ••





Vi:!~ W. '-'Pi il!~d y, TYr. '~D) nght oostem. elfib~dedil'l Ihe biplill1iile 'irolei:aCl1olJ:n rI~l<!\l):rk belMi~n 'lJiIe 74 POl: del1llaril!lso alld h.!' 21.7 ~aihn~~~fse-:t:
,pept~de!i:, (I)

Olaoollii2: CI<U1IdImi.t OImx:!1tiJ7 CI(llJltl1!ll8 Olmildll1i!91 CI~ud1lm15

ClmxI!l!i14 ClmKI!iif18

:5ft1'1OOihing 'Olver bath

~l!Ir\N1!S forrlt1Tlee o'bliiflfled MIh ~e te-st5l:t 4,B pt!ptiides., be)''!: p!~r~'orli11lallm~ \"11[> O'bl<lil"led af'l:e'r

of Ihe




.a:mi amino 1l![ld5.1I1he perrml'tl!ance o~
V>erst0i11 Ihe ,M!'[)SMlMth i)lf tt!i 5, isin<ik:.U@ic with an ,UftIW.

I'I[);Z OOIm'-i!i'I'iS: .m S~


N:3II'P,I OC:o.1lfafi~, T,F,If'C4 'TiF:lPC5

TRlPM6 'O.71~~~~~~~~~~~~~~~ ~~ $flJloolhing over P[lI,Z'$ aJlid a:tIiIirao aQidls -smoothing OIt~1' D\Z'$· OJlI~y P 00.6 .~~ no smoolhlng ., •.•. rlCHiisorimiii'i<lilion

CFliIPi FiI1z:i!r~!




Kv1,S, Kv1,~,
{I. 1

K\I'1.5, K~1,7






NMDM2B pay'~


HOI] 2. 'TI1I;!~ ~~edass I dQI!Q.min:sI~SD ..g5 I(UJ:) 1lU1dSlnmkJ, lVI} n~;n !ihe :n~fct<J!iivc tClIld (lfnh~ first 1)!in!;~I""~ <I;'!;l'l{fi~, sri}, wh~fI;o't~ !!1:1~da_"s; U (,IQn~!lin,... PDz"RflS3 (1/11 ~u~cliGr~u)l (flt7), 1'~ a~ d1Cpo..'liiivc: ~It~tEm in (]II ),.~vihio1rl I:i~ OO~nl

slio\,vn 'lO ~~indboth d,k<;!l, [ .uid class n j]cp'rid~ 't2,'1l 1n I:~", k[\~~nhc h'\lo !;,?\,g:~I,"~'Si.ni~ ~ond and I.hird prilte-ipa~. ::t'l:C'l'l IWig. 3F) O!(w
IUIi:1Ij.C[ 1\.""i'li,hJil~on. lifi 1)lIl1i:[;'ulfi[,nhc IhW ~xi~ cl iii.I:ilil:!il~ji~l~"~ c~a.~:>.1U dOli'lltli.in:>, toii,Jch as neL"'o~ml I1i ili'K: exide iO)'iYlhtls.c'(t:lNOtS,) ( III ), fprercire(j by PCU)licles with 3, 1K'~lil\ -elydmlll1-'tI n::skiu,(! m, !HlSltiml 2" rU(mmllle o'lI'lCI' PDZ d(lm:lii1~, nlC' d05'C'I: a PDl. donmJin lles ro Ihe 'lXls;itivc end of

dl:C ih~f'lj WfllIlcirm!1 a.'\is.~hc more ~!;l~111:ii. the ~111.\)' dl~cl\,~1i! but lll:>tead lie Oll a cmllinarum. c:!lli..;;q; ][1 de-1'ii:g'~ii(ln. ~r!clcCld, the c,mrmiC<lI ~'~~ he Qnl~l in ~.k.~l U~~l,;'i ~+I~.h.o'\,\rqr,,'qt. 1.W9 in")O!1!;l!~~ dill:~r. JiXKrq;~Of!~ rliqij~ C9!Hil}!,I~m It,.,\. "l!l~~ g ~J.>~~~ll~~ ~f ~nQ~ 1li,,'1t'i\'!;".!1ilil,.:: ~iflll:d~'d: 'i,li(;'1j,\' (!If POZ d~t!llli~:I dH~flrl1it ,~I,Jlh~!:dptil!!ci pill ~~~ MOh;~w';:ir,ih~ s.t;~I''''"C:i.i\'~ly :H!l:Idih~ ",i:c\v~ha. ~iTiIlC'I:!,'1,,'S fmltli eur PDZ &tlK![i~IS tl,:P"''S~'litld ill nur ~l~otkl arc b:iDJd ~n'il\~'slig~!lion., i'I!r-;i(. :I?OO<i~i~ll5. F 3. -2, e\.'eL'!~Y di8b'iill'L~~ed Ihl1t:luglr!~Ul :;lCICl;~i'!,l ~ il)' and I <'III I;;O:!l!libul~ <"ub,"nl'~~~i,~II)' 1!l1i;l dL;~i!li. '~Q (Fig. 3G), .~uTiIlP;!ft·1 al. p:1.t;\Iig~ll'b' Sh!ll!ll,\<WIIl:a! !tT'fipi ,or our Lhli.;",(!' rri!1cip:~1;LXl,.'t-l. (.Fi~, J F], ~n~~:1. mite.23 Sre lio,ti)j]l~y 3 dOJTf!llillS lill s.. I.~JT:li~'iklf: il1i;plicsnli.ru. lijGlcc~ivity ill d~ri\!~d. ,fmiiu inl(;'ifm~. l'ite iiJ]lnl~.'in~ In 3.'Void Cf'OiI.:. I'i."m:iu'V1'ry \i>'il:l1 ihe ....


tIilelJiili:li~1t pCM::h.'11.,whi:rea~, peptide Hbrwt;y sereens hove iiihowrti Ihm:~1imlnily b;. derived b:'1dy From the lIL"Cogt'ljtiol~ ofmnino addslu~~:itiiu~S, -2 and to (i). SOOOlld,!linl~1 mare impcrlmHly. Pili! dcmmhl.~ do not fan :~n~o
~ioill~ dlr~IHl!lI~

nli.n!lJ.Cil-3Ci~.\1IMcl p.rOI~~U kilfl~ ~r.,giii8iiilg ilR).. leiiii ~bs2 (12). I ]crc. v.q~llllind,on Ii mmh broader ::I:c~letimi. a::;~milllr p,t1l11cii~JIcis: iu. cfITb::l. IImo~g ~nml;;e PDZ ,;ijDmafi'l'l ru~dI~heiI" lig<mcJi:.;, A Ibholl~h Ihe $('!~IC'C~ii (I r I~rol~~ vit)' n-'l'll.1tcill inlCt'Th:tioll):
IFig •..3. 'tA~o0 (mre~[il;;(ms bel1.'JE!el'll SUl1@;s ·':!rlld m.od@i, ptlrnlrl1~~Eif:S; ~t j)oS'ilion ....flJor ~Irel! P·O'Z dk.m'a~lIls.
{AlZ'l po$i1il/lf~


IElI--4.q po:;lsIlOl'l'~


fo~ IDI~h3tllIl). (B)z~ ,F!!!'g~ni~ly((I~~ relail.~~ \iiith G~~f!Or M.agri.:l[ ~,~f6~. (C) 1'3 "li9~ij'YiiliLj!' !~Clrr,~~t,'l'S wi~h fL."q r~'r MUPP'1 (10113.)'. {I),) COH!~lationl matib::Ib~wee,ilth~ ~od~~ paral1il~~r~ ~'ofil!~I. 74 fDZ dDm!:!ll11iS,~t posilli!J:rI~ -4"



1:. v '!ill

~J3 ~.6
+0.4 'I!,(l:2

~m 30' .' Q.




,CD ,!II 20: ,0 ;I'fi .;:>










n-111 pmincipaJl al>'.is





0. -CIt:2



Z3 ~

!i!SOO tl)' derme ~DZ: dlDrnain $'~reuhrilys,pa(1@. (61) DishiiJul!ie:ll o:llhe 74 POZ dClmi31ifllli in :s~t~mily~pa\ie,Selec~d POZ d'oVlllain:s .ne s~nn"fII. ,repIN~s'eIlUl1lg class i dornaill~ [PSI}-'95 (illiJ) almd $ilIillllk=: {n!/U], 'ii:ia ss Illidma~~s [Gri\[iil 617}amJ {

m!'E!e pri~dpat ,~et

-3, -2:, and -1 '!rId lllleli~t lti!ree l s~al'e~ or lihe'.alllil~iIllClI addl!l. ~E"~r(ent· aQ11l Qf \I'.;1ui1l!l1\leil'l tne ro[re~~tii!ll'l matriix '~lhail ls ~xplailliedbf lhe 12 Iprrinldp~l iilX~:S ide:ntHiedl ~~I"(llilghl sin'~~lilir~\I,il!lll!edi!(imp~iSitic!'i. (IF) Gr.<I~l1Ij(al rI;m~rmtatn~1li Qfdill!! rir:ll~.

-0.8 -~.O




domllfn, Ul'S be,1,\\fel'nbliie d\ass lalild dlas.s ~II d':(lm.a~nsc (!Til) ClJ"1.'~~l1l!lIFlbe~e~ painM~ seq!']e!'.rJOIl! liJi\lergooee ,of i"O',2: dOIilll[l~ns .allilld ~h!llir piliiMii:9~ -11

lJ~fl d1escnb@d as

PDZ·RGS3 UJijJ, al1d d<ls;~ IIII ~mlilains ~:nNO:5 "llill)]. IEMil'! (lJlI)" ~\lhi<h h~





dlS'tci'll~~ in s~1ecd!.1ilW~(~, Seq~~~oo dive.rg!!'ru:e wa~ ob~:aimd [rom p'airwi9E!
ali:gnlllill!!liit5 p~iiI®:rrnro \1litJili V@lC!:cU MlI Vf.l!l'SHonJ 8 (In:rvrllll:i!x, l,"vilJ~1i!1l! me

th~ b'k!I!ilJ1lm62ml2 m.atrb:.. .W'gi~:. Wilj~ Idi:sta~(es 1rI1 s~tt.Cl'i~ty ~(@ar@' [u'CliidMf1I di~tafl.'C;~.oht;;rnfd(irOIflii [he th!f!ee,jiilll!eli!siY!li!~lplotif!l (G).

Sdem:e Seiw,uE', Ffedl'ni:r:~ MaIY~[lIiU~!).

'Q~!Id, in" 1l1~l~lioell~~,rn' gl'gm~i~ll. be co,utmllcd
at l!lJ\llc\i,d org:tm!;l~~pp;:sjjirm

<!ncllll!;*:irl colo-

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1~lt!.'LI:!I~S 1~ ul""!t'ro~ 'Wi.i us..;cl &C;'f1llij~lr.un;~~!lfk lMdparo.l'I1C~rk' IlCgIlI.":S:l1km~o teox enn~rol ~:h! (;1fw.Jrijm(!~ and CVll!h.ml~ dl'C dn'el, or Iw ol). COi'l. :~g~iOlln\M,,~lW a 4S,:..fok! {P' <m-Q) rndl;Jr:,cd. ri~!I;;: I)" dLi':nhi"oT ./rs.rf- Jillie,;;' '~'!!1'll~,p~~~ wit~i\Vr oolWlUIs I FIg, ID mW Whk 5:). By U.~nl.!!l piilrnn~mC' fiI1HZsis: "II: [ouoo llml lite l]rI(~dk~.I:ife "'1N1fi ~or 1;1ll~!l)C \1I"3S ~7'},~kutgc'f [[o:r \\r[, mnooiflll "" 780 d~)'s, rum:1 9S,'Yi!! (~QnlndC'IOC imC'.t'vul ~CI)7~'5'to 'I':.c,.~ tf'- ,"',_Incr I' . It"'''; ~or i,_ • lrun- \PS ~II. . _"' uiil.j'S '1.:1 m:lld 9:$% ] S!ll;7!o 940,; r" - OJ) I ~(lablC' S4,), 'f.1~e

ru1i:~ .In,'i"" I~lioo. i:rlslX~tf:on or thc reliull~~Jg~ ,~'Stcd~lml [hI(:dm~ 'e'fhinlh IDOB}, 1~1.ml(jj1(pm) O1I:no rml'C'n'lm~ (M[D} icib'nDi!3', illld i'e.",iU!nU~lC'OO

100l!C"!"il), (.j~ we Oo.l1~l;}M..-d tlile 1i:I~ :;;}:In,~o:fWT

UiUI)\ C(:!lttml flnU~1i~ d(:'\,Ic1o[pl.'d ili~ulin t~.:~~!. ijl'iC!.' li;u:'IW(itra .2 mid " mo:nD]~~ or ~1,g,C'(fi.!} 2(' ,. Uil,ili:k~ dJd ill'lli.ce tAg., I ~ old: bln.;;Tr miee ~md )lOi!l:nw end old Mi's.r~ mioe were in· :!Jul illili'l:sci~ttllmt (fig:. 2.(:). Alii the immlin.re.~~S1Il1n;l mi(;c di~~hli)'oo m~ldg~{lco~ iUIJO,I~ralli.:c·(Fi~. 2: D}. I.hl~",'cv!.'r. d'f a betes did ncH. de ...elop :in these mi(:~ b~u!'C inslllh~l 00] Ictntmll0ns h~creaj1~\d 10 c!)ma'K.u"'1:['~ !hI! IHmpher.11 ro;.i~OlJ!Ce. ro.t-ching

(ixm}!k1nJ~l~~;i:!ld l~iliU¥



~igni l1i;illnlty :~'»~ICOO c(lmlJ.'llwd ,:vidl dml o rmlr u lRl~ (rOf ~!irst" " 14·rn(lj !i!;-QlJc~io!1!.P0;;: Ilr ; "; ~br bln.~ ", ~ \',H",ful~R;:'1;I~~ P <. ~o-s) (Fig. inf.!, 31\. ..ndi~JbI~ S3),\V~thl»lrn~l1,,1rij~ rcgr\i~~ilQl)" ''\i~ ~irl!iia:ld t~mi thlil 1~~~'tI~m~ :Iifr;: :ipill1iS I~ll."lJlr...2"'_ ~~irld, bfn"TI- ndce !,,\I,j;<re I~O/tl al1ldl4% ~(Jn~!", resp~iv~ly.dimi lifu!lp.;nl~ for ~!:u~lmL~ {fot_lk'<1 .• Ml~;all = 79.~ d1i)'S ,und 95% ['1 '}'J I 10 ?Vb,. fot h1'j"S:rt-,m~i!iii = 916 (1,1.)1$ lIi]d, 95% ('19'1) 10 94.5: for blr:!irf- .median = YOI days ,midi 95% ("I
8S!!:. ~f! 9): [Ille maxiftliittt'n .llir~~a'l!i (90iih jJCfi.Xi11,. 91 ilc) iuercased silUikLr~ly (I~lg. 3A muilllhh::· S4).

lire sQ).(lJ!

[h~ <Ul1~:una_w! o~d.n~re ljll:"l ,~~md .'. IJksT m!~ IF!!!:"IE), ((I!l:;,,,"!L';nalw!l]11 lIJraC'oh!'lCrv!:.'dllypca~nli~ lincmi<!.P"mcro<u~c islers were IoUJl,!!f in oW n'I"~1i! MI'~rfOlI,:ndu-a= r:l~k(l









nidi cf10mmb (~i.~. Ft 2

NC')(JL. W!:.i coua:lp:::tmd ll~ li~t ;;p'Ul,,, of.flnll, b11~~·;rJ-., bh~~TI- 11l1~IiX~ ll~iflg M:;·fn,~pm.'i"I' fIItd by


metric {C~:ii.} alidpilIrnntemc: regroi>."iC!!~ W etillllm.1 for lh~ Cov.rui'iiUe£ (l~bJc S2). Atd'l.O~gh hb~;T~ ilfld blrti,T~l1iilia." displayetl 1Iia!1:1n1OOlk changes. uS'lm.lly m~roehM;crl ~"'.itl~ illmrler Ii Ie' spm~. their 3 fisk ,or dCJJ(h dC1cnJllined by Cox ~:cgICS..,im~\Vn~.

To dClellnhlc wllelher bt,~in. Irs::! al1Jf..>cl<: ~y:>111Ct:aOOJsm. \\'e ::11df:cdYOlllllll,S .'Iflcloki mice i u ill a romJ~Icll(,m~iv~ 100b ,mulll;l1 mmlitm'illi,il. .-;)~ !~~Il (CLAMS). YQun.,g~omrol iflnl2~rni(ie were !l110~ 1!'c,i~!~ .~!!d c(J!m'\D~nOO 1l!lIJro O,w~",n limn did )1O~1Ulg b/rsrJ- or hk~TJ-~tI!ioo (fig. J. H a~1:d['). (h~igcn C{l~$llHllf~km by old ,m~~2 and bJi'S2~;~nicl.: ,d'edlin«it~ ~1~ ~!1";: 'lJ,l1Cll~m,\\'ll(li:e.'i."~hl.'l b!n":1~!l11i~ 'OO!1.'';ll!aTl.cd ~~~.Iilh.(ly l1'XYM~l O"!..!illess :m), A~I !ll(l !lilLt:e W(W ~C.!", ~lI;:lh'~ at .22 .tlJ)l~lh,lO:! hmvc\ (it••~hi;luld b/.i",.l'il-a,nd birs2.-1- ItI.LCC \I!lC'ru :JiliIDl,II 1.\il'Uoc~s ~('~i\'i: '~hc a1t! cU!ll1mL~ (Fig. ]C1, a as Next W~ d!l![oCm~n{l~I:h'il reSI)imlory qlllol.iclnl I[Rq"" !1COYl/o'il. winere V~s\.oililmc" I!Q('$ti~nnl'Lt~' Ihe d!~i~y~nsilkm between i;1!. (R", .. O.7}~r!d C1lrbQh~ldratc' fRog • I) ox idm ion (! 19). lh(:! R'q 1m:

life .<:P.1U1!. 'C'~t~malil.·dat~hc g(n~1 im;:~!.'iGdl similm~y (Ibr \lVT. s:r; OO.}rs and f.l5% C] ro l loM5: l'Orirsf!- . 982 da}1S '!.l.a Q5% (lQ3S w 9!},R) ([fig. U) and table S4)\ 'h2 is e!'lane3scd t11!L'm!gli!out~hC' body I(F i\lil , .lA} ,H1d !l1mn!)I [l'!gijml"oflhcb:m~n,. il!ldlldi~glb'l: C'Cn:;illllllm. lhc ccrebt:'~lun1l•• md ':h~[~I'CWI,IIC and


~!f:ll\'lI';nttru'cu~~lr mu~:l:d of 'ihc hypodmlmw!.ll'i ~Hfl,. S2J (.11. 13), .RcdllJctxJ iUS!llllimi,·iiikc Sigllll~:i,rlglni

B~dy wigln [i!\I\~i\i!!g,C'(g}::I::S,EM, n ~. 6] at 2:2Y.l ,l'Iumrth s. (() s.e'SSI'!l'!!l!oij ~Iri~s,uliI"lSiil'VlSirivity ~Is'~ (,a,vernge ."" Sm,. 11'= 8"IiP .:::
0:[15). ~I)}SlilfVh,lal pmbahmt)J

~f lit:r'- mi'oE!. (A) Ft:I!lld irl!:a~ ~Qj'll'er.ag~ tgf2.4 h,cu,llr) ± SEM, fl "'" 6 mit::@] hili mile or 'relflll;[!'l@' mfm al .2 ,dimJ: ,22: montlils of dlg~. (I)
{holil'lleo5ta~]i~ If1IGldel <liS-

Fi,g,l. M~'lQbg)'lfiSm ,~iFid m~SPi!ll1

!l~llro;tlS incf(xliscs, tt~elife srl-m.ll():f C eJrt:&'imJi onrl
soiit 1:> pOSt"ihlc thait rL'd.~.moo ime!lii~ mlll::lJllrs2 ,eolJj~ extend Ino.u,'>e I~rcspm (3, 14, 15 l', To tcsll~fs hypothc.sTiS. '~'J,'c ck'led one (lJ:h":s2'"'_) d or both "bir,~TA) IOld)~I]mlkcdlr$2 alleles 'Vk:sJ.lJ1"fJ,WfJir{/(I..


<lIJh;;lcs~' ~Dl tI~c bl<l:in by im~reli'O:>:Jjin,g tTl'S} a:nioc wi1.~l neslil1FCR: Iml1lt\~~ic m&cc (} 3 ~\ Polyn~I;:Jif'C ch3jil1:react~ml U:lCl:t) :mfl:ly.'* ~onl'il'ljncd Ih<ll~ .lr... RNA WilS ~'1lIlIinedin allt les~cd tis:luC's of m1JIC' 2 birsrf- mJoc cxccl~~ for ~lebr.i'l:in W,ig., 2A)I, QUIl!l.li!<l~!""C !\1'Wrl5~ ~:<!I~:f,Cripli{ll~:pel fRT-.rC'R) c(ml:i~n~d~ln.n II~~J :R,Ni\ \,\"lL'i ,.',,,,"dll!¢~'ld.;.hOliJl jOllA'. . il:lhl,,~2rr lm)liU[~·:;IIal(ilnlil:lLCGd:""~lrJO'tQ in hk,rlh.rui:u:;; 'L"Onilll;nfld widl~hm of ctllfllm~Jb~,ll1l:k~(l ~fll!i;, A}, :!3y CmaLr.lst, .lrxl RNA aa]Ca:~,j il]U~e 2

~by Cox r':gr,e<s~iO'JI r~ir e.lcill 1M {'.)'.1:llld lrsJ: ,"'"- (." Ibl~!.le]' I:'IlGUise. SQ~id lillll:!s ~!ldiicatethe (ox sul'\l1lvalprobubil1!ty of M and 1fS2""- mtc~' wnmr,ol!,e(j llr oth€:~ ~ (l)vi!3Jri.ates. '()ee tiil ble sa r(D:U detal:l!s:). li1atl::hM Lines .aoneSUJlOnd to IKli,pl~,rq·M~~r (KM) '~ti~liti~ oIc~taJined Iby ~~Ull~li!i( r~lre~oot~rmirl~
~Dn ~ til~'~e54 {or dettl~l!;).

hil,gnhfll, 1JlCSI.m·Clf'C cl'I:prcsf':iolll did rnollai~.C 1)llloc in lP"IlICTCatic ~ «Us {f~g.:2A). C'ro,s.::;e:> IJc1WiX~l h.b'Srl- mice· p.nuJilioOO O!l~lciillg al a lti1!rn~la~ rr:cqllOltq)'" W&~~I1lSC'ro~ ncl\1;'U.111 bt,'S,]""/- 311d hlh2""'""1I1icc ~ru(lLlcOO fCM"Crgnsp:r:ia,g: ollSpJ'iIlg~"''t'I:c n~\:er ,wodLlo."id b)' C1.'OS:1:iimg bln.':Tf. mice (fig. SJA). O:III 111>JiI'c·a:nd ~cmde hll'iiI-Y-1'- mirerolm~nled~he r~105~ .. rOClld ~.ih d~Jb)' c(m'qlmsnn to Ihe {l~hCl'. ~o~ (F~,
How(:-v~r~ b), 22 IV!OOl!li'l~ll; hJr'!'!,!

or l';I"o,:r1- fl:IfIClMi'$rf-



• l~rn2"1-1~(lrsl"") - IKM (11rs2.·~~ QWT -e~l('Nf) -IKM (WIi)


m g h.t,~vicr

:(iu:d /ih'};;2


tl"llll] (:(IU~l'(iIk






iocn.~.cl, ,~~l(~iEy i~ig. 3. B mjd Q Uie S

tSOQ lir~Span (,days)




!!rIin'" u.. ~S~I~





OlJII~c }'9~~ng mlce ,cl.~S'plll.}'Cd~'Ile 'Y8'~ml ~i~ml<ll,



rhyl.trim" \\'hid1 <1pprooch,cd t!~c !l1Ii!/!>.im\!ffi


~~l.ri!1!g~~lL;' ~~~ ~y~.I'~l·F~~, 30). ~,y ~n!ro~iJ:. okl, _t1rsi IlIi!ij,1~kl!'.! !~~ dUI,I:vrm1 ~hyl~.l!~l~U'i\;'RCIJ i;.\I'I~ illdii!i.Lul'Ig)IJl.i:dl(lb:l!e t~lWi.,.~'TI .lligl~'t. mId ~~Ja"'i ~yck'i! (Fi:!i,l. JD). IHol~,!~v(;r,!he Rqfor old bb~.Tr "ur!d h!nGr~ nlk:cill~~I~~cl ~igll.ifk~u~~ly dlll~;ili;g Ih~ d~u;ft< Cyc.lIC. r(~vt,ali~1,g"u 1'i1.orc jlou~bfu~ !r',imt~;dlCii:l
1X1\','(~Ui I~l ~]'i~ ~~LrOO~ljldr.l[c flXid!lliM Ifig. 3D i. .lneJ~~. hl!;Jil~b,~J!on~.J!i\'~d~n:im:lLtil.l~b:o d~5J:'ill!!y :i~liWl(!j' Rq Llml. is C~OliI!:'f. [l':! I.he '\,Itiilui! of

heallJ;ly~nid~lc~.gctl m:lItull.,>10). ( Ol<>.idiAh'C' stress is a...>OCimi!d ';.\lith:1 fL'.hLccd ' li[c sp3f1" mnd n1!ilH)' cll1.yme!' I~Jintccl, cells lrom i,))! tdrn~ i\i~ !{11Ul'.'.'e:<pec:illlJ~y Slll!(lro~ OOCdi:illmlaii~ ,(Sodl' O. 4. :lll, In miee, [':r..xOI l'lIDn'!iO'c.s dlC (."(pl'1.'S.'liO'!l of Sod2 •. ."0 w~~~vf.:-l'l~ig>J.l~d \7!"hctJh~f ro(lUct"d ~1I.ll!,lm'twl [~2 IlI!.~gl.n hdr t:i~,lil!taiu b,,~il!t Sodi2 1;'CJn1Jcntmli.o~1." during ~:eooing, s002mld F~1',i'i:'o1Ipm~Ctifl wa" ITic'I!,>lI.u~db1Jl i!11ml!nQbo]ou i~g h)1lJ01llmhn]Jlic ly~:r~'~ liio~n }<'Ollll1l. .~"~~ eld !(:!.li~ ~ftlru ,nW 2: ~lol!.m;; uf[1,jlf r~,]ii1lg: IF.ig- 4At The ~ ,ool1cl..~lllrnJ:iO:r4"W(triC Ilalrhrln~cd by j~dL iug t~.c ~Q1,in1!l,iJ~icc;hO\\~c'!.IC;'I". 500,2 deercased 91. le.aSl 5£1%111 the oM foo cO:l1Itl'ol mltc~ fri~,. 4, A. aad .13). .By coul~rnst. S()d~ was Iwt redtteed hy fe:ecl:[ug otd ..M~:'.r"_ .m~e]Mf,'fr~ mi~ l.Fig, 4. t\, aad B1. A $llni~u ~}:!lucm~'\.'::ls OmCl'\lOO lOr FOi'l,Oi h:\'c(fui; in )'QW\g andotd h)fpotllii!~!mlk: !isj;~~ IFig. 4" A and C), 11I~ui'l, h}f~liliil.~lmlic So& re\il'll~S;a more YO!1llhihiroSJ)mlSc to recdirng l~ilg. 2:. Redu,C!edi ·U;!UIf®'lla~. IlfSl (a !1Ise~periphe ral lml'stll]n Wle:si$ti'llllce. (Ai nO,pi) IRT ~ ·P(R of /152 RNA in m()llS~ liS$lIe~aiiiid (bo~t(liIY:I~ 1!il1iI!iI !l1t]I:a'live RT-lP(lR (li!(lfill7latized ::!: S,IEM; n "". 5,~ ~p ..::0.(5) in I:Jlr,!:I~nf'!~ g

paim::f1~a~l{s2~- irldk~les~ys;teV1llh: 1r'$,2-1- mk:el (18) If,ood r n1:aRc [tl1;lerag~ {gI,2~hQIJJ'r}::t: SEM; {j' "" ' 6;~,P < (U)5~ ,[I~ :2 ml)liI~I1S,and .2.2 IlllImll'h£ Illr <lglll. (C} HOMA2 of is (a1lllll;r[lg~'Ilr ~. SEM" n =: :S; "'.P'.;: Q,O'5). "II)) AV~f,i'l!g~!w~C! (.tS·EM,: tI' = 6i ~, .z:: '(1.05,) lillld~f~1l bw~dl gity,(!~~~d~~f~'W~,C'!.!rVt. ~E) ~iI:5t~dl in5-uliil'l OOI'l(I!\l1lr.1~~ilJ'I1S.tSEM,. n ~, :14 10 16,; "'Po:;: 0.(5)" ~F) (l.ert) IF'iliiiI!:FrIla1l: ( sec!lolil,5 fr'~ 11111 .22·molilitifj·oM micer ~~enf1iI!I1!mlJlllirostained with ntil:lorly agafin:slimuHn ~9r~el1~ or' '£llwcagofl (red).

in mroewiilil ooLluced brilil1 hs:2. Tog('~jter. OlU !'Cl>U I!S sl~o\r.. d]:~1reduced lni:2 . <,iigl!lillli~'\g iu~ aU ti~sl!c$. Ur_~t""_) (:If jllC'\i~.in Ihe
Dmin (bli"S]+I-) iucn.~~LI\i~s I~ ~mll>n.f rnice t]II'; mila! mlli rlcod oil. ;J high·CMID1k"Y die. ilNlUllt 5 1lI:1011~h!li ruttilrulIDlil"" m(~u in blJ:vTI-~nkc' ~'13bk:54). diS


{ri!!)ln) th,e ami)liIl1t, Qfi!;let ~ ,cell glr,~;:. ~QI~~~age ± S~M. n -6, ~~cti,oniS; ~p ,<


Son~~ :mildi~":ilm~1 d~fltca1ntl:ereslric:~io:n. :i:'l".dUQl~ bod)' si ze, fLnu:i inereased peripheral ~IlSiilr~n Il, sclls;iliv i~~cXilcnd IiIlmmmliliilln li Fespan (5. 22) .


Cqx (Ol1112")


COx (blP.il2""')

.b!n2"" KM~!:i'mtM

~.*.~~~~~~~~~~~~~~~~ iii m 600 8IXI ~m

-'~J~ - ,~~M~_2)

IIL:~rDSp.;m 'I~p)

Fiig. 3:. L:ijf;e span and 18fi;~rlgy me!abtll.fcsm. (1\)1 O1n.erWlcl sL!lrvil¥Bll pl'Otila:lli!J1ties-' f"'$2 ".), [)1rs2#~ "., b'~l,Je). ;;Ifldbffirl- (.&, lled)-ca~\mdatedhu:mt~e COIX: I'i(!g~,e!lsiort Sotidlllf:lI1Iesshow ~he 'indktllred 'Qf!it'a(l!eu,(!(15 c'()l1Itn:illedfor ,(li~~er [OVodlli]t,es (talJle :53:). Ha:tcl'iecl ti:ne~ are thE! (Jlrre-.spomJ.din~1 KM I€stljm,gjl~~ Ib~€d' Olm'~l'Ie piiHam;e't_m: relgnt~roml (wble 54). (I) MJ€dTalni 0;2 IconS'IJm,p~~:Ofll (soHd horiz~flta I. IUne). '99"t~(] (n;otl:h). all1ld tile IliollWr Clind!
Wlpper ,q,U;;JII1t!1~es~~

= 4!~S flOP' each genotype. '~p -e CUl001)' bV rn~le mke in d!lrik bhiJd:ed OOJ;~) and light qrcle:s (Qj~f1ooxe;>.), (() M.ediiiln vo,iui!ltalry mOV!ltnelflt {±95% 'I~ [Qr mille mioe Ull00!ilJr~ dllringl:al 5"5 ilm'tf!ilVal m the ,d~lrk,(yd,€!' fn "" 42)for'ilIach g:ooolype. "p <~ O.ol., """p <: (I.OOOcI). ~D') A1{!lrag~R:1lI (ifCiDzAIOzl di!l'e<milMd dUlriiiilg 49~ouif:S: from sl~iIf1Iall!!mice of U'te1 i~d~c<'l'lOOage. and ge;nolype •.Medliai'il R~ der~nninedl fill dark. (DC, h~;ck bad and l~'ighil:(LC) 'C:y{~es: I~KfI!I~k.li~"W.i:d[i5; M.nj]tl"~U'llle,trit te~to'P <: OJW01; ""'p =: OJl3!t 'I .

Fi!g!. 4. W$ ~f brain 11f52.

5!ti31bi~f!zes S®dl2ir1J l'iIyp.olha!tamk

p~tprandi!:!~ mam.'A)




,~~ mate SiibU!1lg!! of t~ 'indicated 9llrrl®lype' be-

\w.re prepafeirll Wrorrlll pall~


,-hoor rt.emn,..g 'Wed), r~Dlv~ Iby
.ill iii d <lllli·

SI!)S IP~l~l.ai!nide
I(l'lel!:t irQI!),ilI

lmm L!fI;ollkm'ed1'lI'rith


.SOOt! or

·4" 2mOl,·~

1~1~~~ii!mJef1l~ ~~l"IirU'm ·w IlNJUlrvalooj~rnooijn1g •. AJ,~:MJ~ad1ogJ ...phs 'I'cre:re qUiilll'ltified. and ltihe~~rlo .~~''U1/te1I'1Sm€:~(~'~Si) fo~ ~ 50d2: '~'" .S)'Of (C) WO:m:>l. W = 4) ~'!'as('~kll!~~tm EI~e-s sh~ liiie mooiilnr rat» i(s®~id non:Nntal tillle)aoolhe toww y.~, Qllllll:U:es; tbe ~rud<ait::W<ll]is·lIl~mp.aral!flt:trfl!; 'ffisl!Was: lillld IDOOIIfIpare ~he gm:u:p5iiICr,ooli aUL'g~'HI'tYPE1!i: ("p'.< OilS}.

F~:r.O-l (t'I\I~ f!'!~~:lX':nde!flt ,e-JqJerilll'lE!ljJhjillie ·~hG'll{fIlI. r&'tubulin (sIilO\lI,nr~ m.e 'll'~ii!leri!TIe!f!!j was ~!iIi!IiI'I~ ljJ,oblottediroraUlhe

B, 1 ,.r-rII~~



ROOSt la. M. Sth",b~!t et ¢.,I. NWf.ri!w,;!'3, jl(!I61! (20'D~1, ~-9- Y.·IM<!mg ,t! ~1 PhI"'" I~Qrfpi('$ .'2.'1. :U~ 1(2~t •• 21[), M. III, itlrm I!\!' mr., J (fIn:. EJodcr:rmi!tl. .m.ralt. 9101 4·0~
~Sl, :21. H. ~!!bi1, ~

IR~:t'e~f,lifl~1!'5~!ild N\iiJt!!,5 1.. K. A. HU!il~el, ItM. Ill!e)flll·I.d6,Amlu.iIlel'• .fM~omi!lLse', "!~1UQ(!:S<), 2. 1t..I\.. Miller. },. M\. Gt~ft. 5,o1!:.~~, SllM ~MO~, ~" C. ~~II)1Dn..('('II :120. 44~ ~_::!.o(m. ..:1. 0. P" '5i~~liIir, Q!'<!Il!l"It" Sift Am. ~94. illS l 5- '"- $, Sl)nkgw~H J- S" Rochal, 11\. "Ii- ~Wim:liR, t1. 1\, Al R~i~~, A. lIin.~~, Nen&.Aw.r;I, flr«. . !),,1A. 1;03, 7~QI 1l(l~6t 6. '"- IF.Whi~. Scit-J'lr::t· ~iC!Ol, 111[(1~lfi:OiJJ. 7. 0. 1. Cbnrv f~ ~I.:, Sr:iffi{.e':~!2, ~o~ f200dl .. 13. o. S, H~g~. 8.. 'Gershm<!fi\, !W. ~p,'T!!!, LM. P.alme·,r" '"' Ti!t2lr, .W/I,I~ ~2.9, 5!!'i~ f4®~~, 9. 0.], Wi1!hel> el oJ,.~Me ]!9:1. '!)'oo tl.'~!lIal ~lJl D_ j., Willhe~ et oJ,< ffl}t ~. 2~, 32 U??~.·ll n, D, w. 'bl'l'lf!1' l~ iL lftrlI~dbn."., iJpp1/M Sr.lrlTl.'(lj Arll:li~ .. (\\IRe)', r4~ 'fait. 199'~j, U. !l W. G~Ui~ e~~!.,r;;~~tllIe!~4r,~~.67 {ZOO~}. u. :It 'um f!J 01,}. Gin,.I1'1~~ :U.olI,908·i(2ooU. 1..:1. l l IPi!~ecS er ol., NOI:. ~t :,!:9'. l1:11 !1'!l981'. ~5- ( • .ilI, W~kglli'l lit ![i!- ![ilifilJl'.~ 1lI,-'S, ~, G·. II:l!>'k~!1TI,. ~M~ 2"90. ]~·10.000), 16. O. ]. Built!; ft al., NfIlure 401, 377' f2m~ .. n, ,Il" t Cll~dhv:r:r ,IN (it.,}. Cli4. US, (MO'



increased! (i.;ht:u:l.u.i~lg lillsulin might hove Itc·g;l· tll\'C c~lQc!sQfI dl(!'bl'3,in~ha~ canredaec life span (4, 11. 13), By ,dll.iC'O!~y ilu!;!Iumting brain Irs:! si,g.. lived s}'!i~moc:- 11~t'1-- lf1il1::Cafli:l n~o.l)C illsul:i~~ 1l~lilr!J!. ~mr!!,gin1! hrain elm [1!C ~ht[!'~oo .from dro ,~r:!~~ili\'C' alldglloc~C' l(l~crnnt ,lulI1 WT mi()c; IlI.'Qo:tive cnbcl.~ OifhYI'ICri!k';ldl~lC1nia ~hm mllil1",," ilo~lICvcr.Iong~lli.v.::d b.mfn-sp:x:Hlc bIl'S]""- and ny devdop v, .~thovct"\'!,~eight ;['Ind ad~..ruJcing ~lgC. hirs,T'" mice .Me· imlllililfCsiSlfflll. hYWiCr'h'tSlJJli.- Co.~f!!tnnl W~~l tlnl:i•. hypOl!m\lls,. moderate d~g])" nt.':!uic" and ~~l!OO~e ~l1to·k~'t Tl1~~m,,"Ch;_!!~i:sm cxeP:'i'>C', ,cillJoroo re~>1!!1:clion, ru~d weigMn loss rCSI'lll1·n~:s]brt,;ror this d~:spO!ri~y is WJllknowl1_ whidl rodlllOl.: ciri;~II~~ing i!!~{uhl] mi:glil!.illlc","mliC R(.>gi;'tl'dlbis,. 'Olll'li'!,,~ltlN poiMO ~n~h~ bl:l~illl ;J:lilh\;' ~]I;;:s.,pml by n~t~·llutiltin~liN s;~11rllin!g im. ~ilc:: hl:'il!m site wimIT: ~dllttd illiS!lI~ill-like S.iWllfllU!'Ig eml Olh(:"I" ~mu:1lie.... alU'it hrlpm\l\l:.: Ipcrip1KT.:tl irtiiulin tliJ1VC II cm~:si$I.clln .c~Tec~ e"l:tc:m:l rI1lfUll.ll1li1!Jliml ~I$uuhrh.li". ~1I0b as redueed grroMh IwmlOillie :>irgto tlfilinJ;j, cUHIl:Jlmvclhc Slime efi:ttt ~5),llldG"l'XI, life ~il'l-fI!.." i~docs h~ C: eh'g>fUI,' 31ldO mdtll'wga",,;ier(1.3), hiiimml OOltcflllllWlS. dis;p~f~J' inCfiC{'l...odperip1:aeunl /\5 nUnlilll1la~Sage. llOJ11;PC,USru[OI)' hyperin.~M:lli~ nlilsl1lin ::U1.si1ivi~y::md rud~n."d dg]ming inmlin ~lclnia WlS I), dcvdoplJ.ID nmi fltnill. Si ucose OOnOCbm'ilItOm (13),.1 I.cn.ce. we SliiggCS! Ihlil tl~clr:s2 IJItl~ ~IOmeOS13:il.i. amlllreVCI1:l" Ihcpro.W.'!C'SSIOH ww~m siglJalillS cascade in Ihe llJmin lntcgm!Cs tl~e effocis. s li~c,.·dIOC'lll.Cl! h'V£~)iPC .2 dio1bti:a.cs (6); hOWC\ACr:, ofpt..~pi1&'l1l1: IrLj'[ri~m hmnoosk"l.~s; Wi~l ~:il~.spoo.
l~o\\IC\fi(;'I",Olaf kll1g..:l.ived mice m.'C :sl~ghr~~y hnger and c:mlSUIHt!' ~baut d'lC s[m~1i: r sll i.yhl~y t:IlmC' o foool:l1ml t~lC'sholil-l~vJC'd ~om:rok Imkcd • .IInl.!:l.l

,'2. 1t..It Ml'IteJ, J. M. !'Iar~J, .i!!. Gale-Oki.. D. T. ~k~, Algj!,\~ Cm:1ll~ ROO~}, H. M. IBarb1!!1li t· al., &p~, litmn!llJ: 318..137 (2003:}. r 2·<1. li'l'i1'II!.in'k Ft le~!i!il!n, ft M.nij~, A'!ibiIl, ~. :Fuji~, .ii~d c 'i,I'. ""~~(I'oI'~l\i!fQr t~~liIil;;!~ ~~i!il!)t;~·~ (. ~~, l..E!mqui~iI,. , Andmoo".oll'ld H. li'~dmal1l '~I:f IleI~M ~ OI!M~~.!'!'!~~'~vlk VilllS ~'IJ.p.PIlli1edIby ~!I!iI (gfil!f!I~. ,*SSJl6 ~~dIIUS',]2 til M.F.'WJ, 1'h~ ~ap~nSociEty I j'or ·tlil@· !l'rU.IIlOtiOIl or S:(iMC~ (A.l)·, ~i!lr:ilIf1f! 'II'~\IiIl\ild~I' 5!;ie!!l'~ ~1J!lmlJiIl~ ~jII"U, Mif:W, 15 an ~nmli[Ja~~ <It iI'1e !TI~ardll!{YI~Ii1~s fl\edltal !!iI$!:i1!Jli~.


rart~e:, ~ G.

1f.n;!l1ltlllfiOlD, 1Jiqg,g~t!iliW


5l1!1p;Pfirtingl OIllII~ii1il' M .. t'~iIi'ii 1 ..
~;l!:ijMremll!J,a~g~.ooltnWlillil31.iiSB~6J369tocll M.aIW~ .!WI ~'l!ii!lfo~· IFig50.Sl tit) ~.~ Tiililles~:lla Si; ,1te1'~810~

,6 M:mh 2007,; ll~~ep1)ffi:I :i3111~t .20017 1O,1li261:l!;ie~~~. n~lH'9

IPiB'Iched11 Re'g ulatss H:edgl'B,hl'D'g Slllgn,aUnl,g at thai IPrimlsry ICil!iurn
R~]at .Roha:t!:ili.:I:·2:~ .~]liljoi!ll1!laJ l(ei!lll~rnri(,.:ll", Mamlew p, S{<lU~ M
Primary di.i3i1UE1 ~si!!ntiill ~!Or trilnsllll\m::~~;Cln~ '~he Hi!!dgE!ho9 mlh~ s:i!lJna~ 'in llI1IilIFlHl1Iilrs:. Wi~ o i:nwf>tigal~~ '!hI! IFole (IIr 1[:Ir:ll1lary ciliaJil1l r'i29Ill!"~:Q:ne~ W'a.l>c:n,ed] (P~d), Ute U!Cl!lllc r for Senk H.@d~!!l1og (Slhh}. Pitt!. ·toe~l]~ed to em.!! alnd lnlnll!l,Uedl:!illrlocth@n'ed (Smo) bl:; p\r,ew.n~iin9b al((lIl!lflIl!~"ti~nwillih]1fI d[](!. Wh'A SIl1~Ibound Ie Ptd,. IP~ll.€rril:~h@ dl.ia, ~!.1lldlingltij) aCCUlrTlIUlillitioifl a:r Smo ;::lInd aCll\l'~liolfl olr~iglrt.1!U~gl. T.h'U~. ~il'im"ry d~i"Si~ii!s~Shl1alnd U'a!ilsd!Jo~ si~nil!ls; tha't pLay ,clnlth!:~l mt~$ilil, d!f~:k'i! pmen1t, (ar(.i!'10gen:~S~s.a!'!d$h~Pt'l1 c~Uf~ !'!c:llio!i:).

Or,~)1~IKi 1L'lJ!l;;'\.mcoll1IhL'3nc: I~rmein. Sll1IoolilcllCd "SI~Kl}. The "r;lCO;:l~d pJ'otcln Soulioc 1:1I,"'dg:l;lilqg ,lSbh~ bund~ and iVH~Clh~<IW:H :r'!e I. .,~llo\\li~~
:;ICtU\l',~~ of Sll:1P. Sm(~IJ;II.;rJ 'lr~~",n;: '!,Irgl,;i £CW'; ~m mr,]];IM.:rUp~ion throlJ;gh I:hc (iIii n-II'1i1.ily (11 r 1rn~1:.!;dipli{)1] ~iI.cl.ors. The mi1cimnism. o:},'I';ilirich S~lh

il1hilb~ts I?tCI a~ld PI.\:I ~nh~billS Snm, ]s. nol
Iilnderstood ii III mammoils., III Dm:wpMf".Ptrc irn'hibiliS ~lC~mOV(;H1.ellll Sp.uo 10 the'p:lasma IlI1Cmllbr.Jl'le .. Bin/cling o!f Uh Q1\I:ll."S~hc inICllrmIOOJi~(tJ.1 on~~, lromlhc plasll~.


il11J}{)l1;I;nce {D Hi h sig]lml in,g in 'nmrmn<lIS,lilcre .h.c. F.~e<l~el!gog..U.-~hl' s.~gn."LI.~J!.g. . P"lhw~.'j. 9005, IJ5-A. .20I1ipan~'IU!n~ of OmQlog(l!', :~mr~d iUrr1pllaysml UI!lpoOlil~u~~role bo~lhIn !;imbiiyon~,c arC' g<l!~ In o1Jr~IidCIll'mid~"_g (j·r iwr]y c\'~nl" ~n 'II~!iity S(lhoot ·or 1WNJidn~,. Slillilford, C1\9/l JiJIS, USA. __ .dc\'t~QlunC'nl.aatd.in .. duh !)!1:i.'i!tiJtx."1I r~u:u:;- ~hi~ P1!I~~.~"'ay.. d~.t; .~Ib.'lr.. !I,;C of ~ig!mr. i]u; '1~ese authors ()crn.tliibuled i~ally to U~~ ~rk. 111 .. ~im~ (1, 2). n,)l:sn;.~UbIJt~;I~ ,oH~(' I);!jtljw.~ 't.;,I~lSC~ [r..~n:>!TItn1brancpr{l't:~ It f".dc~'K-d] tp[(: I.) IU';l;-p:'i f1l'o ~h(lm (OfU'tsPOrder!(;e shool'l1boe .alilld~e~~. E-I'!l1ai1i: birl]~ del~'Cl'" !ii~d li.u~]ii3l'i ·it31'ie1,':[" (2). [k'".,pi~tlic till; rilthMIYUli!WiCcl off bjl ~lll1ibil:illtgth~~u[IC'lii)],i l.mmt!@~I~nrord\.~,d!I


of De:v~QPlll~nl~i BI@tOQY.(j~nBLi'l:S, .~md aiDel1:flrneeriifi.~ OIliId H'lI'IliJrd IiUQlles Medical ImMiluie. SianJorlili U'm,I!;'eT>i'ty Sdt'OOl~1 M..edlc,'me,. Slanflll'd, (A


a ~~I~~!Imdr~~J!c thm dim;:tly~lru..1s <bnQ tl!Cth.<l1J.:1S, \"c:lok!~. ~~lQ'\,,'l,ng Sill!! lO~rnJ1s.l0"p/.(:../ is it.~U· ,..uIWlsl;rh.tKlf1<l11 qmq. to the Ill~:li'm!!m'~n~brone1!illl{! ;!~Iiv;ni,,'l S:rn'Q' flO, .. B~~~e QI'{l'\~~!.ft;i~!l !l,~g~~,;;1!i!lg3,'n ll~!l: d!$Qt;)v!i,'!)'~!>ti~ ~!!'g~ !Jf Hh ~!wm!i,~~" in~(ij.'<I!,""'~ :!';i!;: I IIm~L;'i!1 l In ~'\'iI.~z.;,~,:;~~~,,~ a.'i"l!ll'1,;...;:ri~ fo~r!Ht.h'l'!;l'ty~.I;lti\'lllim!_ p(~id~1 ~(Dllflp(,lnC'J1IS or prim<l:t)" ~i~i:1 Ii\;'WII.i!fl;-d "t~ p!~I ~:iIrokin:k~v~!~ t~lI;!¥tti ~o d~~b), 'I, Itollill}l lind f{)[ lIh $!I,gn.laij.il~g sill~L,,·;t~'(1 lhrui l;ubOl~nlIIHr !oc;:ll~i;.!lllion h~~ an im;po:rt."I!~u.mlc in :nlii!.ln~lb'!li'Lf~ CO!Jit:~!lm;d 10 a!l1.:'re,1IS1:. lInti I 24 h('l,\urs a It~r <1(10i·

Illese ,C(;n:::


Hh ~igll,~!i!lJ!1: (jl fri.~!.mr.Ji' ci;~i;l ,,~ od! :-;li.IIr~ a ce pimjil."Cl.rOi!L'lruWI!'ild m~ J~'iOll·~vC'tlelbr:J~c ktifi1~~ ,0;::1 IUriie~ocllil s\'''fi'>o.J.y "~i:lii'~'mnile" rOil' :o;.igilfll~ (6 h Severnl OUlblip.ili1Ci1lU;, Ithe ~HI pmlilwlI),. jllil~kttll" 01' lliI,~ Sma end lhe Gli pro1Cins. :r[;(;(lI1nlilat~inl p'rilml1ty oeili.a. mtd Sm~,)is,enriched i~~ !111 ponl ci~jll. 5Iil];nll!ll~kllil w~lh Shh I[ 7. t'i/). 11;,* c...... <l:t11htGt:! tile dYllliJ!~nic subcelllUlar ~(!o(.;a~-· il<l~ton of".Pu:'1ru'ld 81l!10 illw~.Irnma]i"n ccI~ wIth du.-:use ofl!!m cl <1I!1aibQclrol>!rI 1i'tC lWO p:m'l~iUlS (fiS'


60:11 ell'Sllh Wig, I.J\.).J.\lkr~.limlillalijc¥n ~~]~!'i, ic:t1 Shh P't S/I,{it C'i:10'O:!¢1101Jl:!< SIY!JJ\.Y"o.lS eYI~ie-i1(;·din ~ir:bn.'li)'citt.1. lFig. IB ~ ~11l1~ ~]l,('llil'i rlllo. ~~'IlN! :ijUC'rI~l~yOfSI'liifi lin ci;l~i~ ~il, 10 ii~~~~ as car!y 0.<, I ~~our llifll..'i," stilii~t:iIlIltum of relli> widl


Dn;IIl~ .E). Rd!1!~«!lIClt~~~ I imp !h",'>,1Q Q~!l~ by mg.,~!:s of ,;;1~tro\' inu;;, ~~PJ~~l:;~S9G1, Uhll),rUn\rdY lLl;:,tivil.y (.~!}S~) rul~ p~'\'t':Il~d sIno occu~m,lbuiic!l~ il~priJlIIm:y cj~i;1fF~g,. I, Do and E). llms,.lhc rogl~l~ut(l'~:1 {]j:fSnm !OC.<!J!i~liGn ~y 'tei iil oO!1~crvcd fliQ!w! ~1:~1' 1j]llQl<Ir.'iU'];;1L~. To !,Ilj[kl1'i~ltd liow pre I. ~'iay ~'L;'g!i~iIIC clll!)'

ijnDllI.:,,'b:i:en(lC 'of Shh


or rt,~








(:N~l'i~Ilk'll d'ii:'


Shh or Sil.:Ci (Pl£ .. l:C'mld fig. S2). This li~ely klill"liCntcd rel'oL.dit!nlkil:l rrol~l II ,cy~('ipla5illlic

,dlmwd dCl.l.l~iaUI of eudoPtt:l aDid Sm{~m old!.\~U(\d !110~~';; bl<l:<:l<>(N nl~ 3T3C4.!'1I:;:~ Wle. 11tI(}1l'1CCCllb~)'(lv~ic f1tuobiln~!l'i, (~MEl'.~}.~wl) Ilh.ro.."l(Ill8i\l~ eell I~ !( 9}. 1111~!~D!1dil].!il \lI;I$ u(:~h'.uoo in NIH 3D o;,.~""
~(l ,Q1UlI~[~ gi\,;1~1J1,L~

s: 1). Tlu..


b<yII(O;]lllll.:.:n[ Wllh ~ilfltt,\'IfS~d:i !f SAG O

(S.'llh . fll,!l(m:iil.h

bCcal~ Ihe I01\i'LI!U1nmmt 'Ciif S1l1~~ro.tc~n im:OOJl:;t' ,It dlis tin:1C point en!:. 1J.\, , .. "[:0. d(:~cnuinc wl1C'thcfP,~cl~gld u~~ the loc u li!'''"'l~O:!1 ('If S!VilO. w~ IJ'xmlml:n!1,\d 51'110 ~oe;:tl· . ... -. ~.'-1-. _ W","~IIQ:!I1 m MEF,.:: ([4111 Wei 1ll1C(' i9'J. 11~1,;$ cells :<h~Wt>dI,.Xl~g~!ill!th'C m:~i\,~m iL)l~0", ill mrWl gcnelr.mM:ril!llim~ Ulg. S3t ' 'om;~"lt'\ll[ \\'i~ha "oJ~c of Pl(: liJl 1!I;,~~~~.,u~i SmQ [!':<lffi!l'ki.,!g. S!'IIiJO U\g \\,'Ol." ~;!JISli~li.IIlijv~dylo.co;llijltidIQ priil~1i!1'tCi~i:J ~n



ii~ME.F's fl:l'itl IY!OU.~ cmhryiM. s.\d{l~nOili~ Pre I was present 1m 511U-ull !I1'1.10Ui'lt .. iu. M_EFs, ~bC'3rlhc Iim~l. f deta;liml by ilmmmo.. o ~lllore::t."'Cncc. We (1IDCiI'CfiO'rt illflC'uca.'1oonilc OII1'IO'i.ml:s: . on~·[d pro~eju b)' sli~mll!.1!tinseclls ~~Iit~ AG. S Under~hc.sc I;ondil[o:rl<;. Plel W;]j;,hiS,My !.lnrich~d in. ~~rilln,lg)! eil im, FvS~ 2:A~. TI,1(! eili,~!'y 10(.~,!!i~~Lt'i11t or.roc I wa.~~ord~:rrllOOi~~ OiI.dO:II:iomd ~~';!ys. ij!rcc Fi!1'l!L.Plcl fllC~ HI yt:l~"\' W11utln;:.:c.~.ll~m'~il!J ,(ptc.I":'fFP) W,,"" ro£llmd aOOlllnd.lhc base and .~ndl(: :;aw.n Q rci~i<! IlllHsl'km.lI!il!i;.'t! 1'1(.; r': IO(.~Is in1i::t1!;,1C!: i ~'...il.lii ,I rolr-Q,,;,V!1l.i!\; eIHcooij.!l;l;l [111,(:,1YFP ,t f'i:gs" 2B
iz:ll.ttomi ,or PIC ]

~1cl fig •. S ~ 2). s"''COJld•. 1~1lCI-YFP


0- UiI'iIl'fMtedi

.Shh .SA.G F'iig. 1. R:api~ 1[(I(a !.il!:afll011l of Sm®

primary ~Mi~arlie',act~tf{ln olf the H n pa!lll\lIay end regulatioll by Ptc;L M) Ifl1lmlLlfl,oblollS ~\!ithr ,;lIn!tlrbocli~

1Il~e-dlttl assess allTlount::! o~ ,e!1d(!)g,oi'lnCiltlSp.r~l'eofi1~ in i!:dra i:lS r'nonlll NIH 313 oelts trwU!d wiU1 Shih or SAG (100 nM). {B} Enriidn\'i~re



and @~dn

!lIi!e~t ,~~S:mo,l~ primJY dlial orr

NlII~1.313 ceUs le:U umtre-atBC.i (mJl' lr®~)or trested \¥ii~n Sh~ er SAG

{lOOlll'lilJ' ~'!l;r24 ho'urs. ~C} M~1l in tel1l'~lty QIW$;1il1I1li lil!!lO:rllKefllC'eill ,dti~ ~II~: teRs t~~~~~~vilh @,~ ~:U Shih or :iAG {:.lJ,)O filM). EiChr poii'ill:
1lfI1'tI.rI ;it

:SE:M or


~eS(~rIt~rro;m 10 to 20 cilial, ~[lI ,~nd IE) G(lI1!~ltl!lti~@1l:r'Si~n[! o,'f
SIiIT(lin~hE! cilia or uM1iimLlla~e~

ptcI.--i- .M Us and rleven.a [ by retrO'lliraUy ~rillrndlLJ:CiedPtt:l~)(fP'.
11'1 (B~alild ~[I):.cOllilf'(lIcallmage-s

,glootylij'l'fild ~~ lilu'lil1l be~)ilindl Smo'(gree,n) \VCfe: de'te(;t~~ by ~!fIiimliJl'!~~tiJiI):re~:n~~; ~u.!d,~1 (IjWi~) M!~~ ~!a~md Mlih 4',6 .·d~illmlidlil1o·2"llhel1ylindo~e

~hedWi~~y ~r~r


i'n Pfc;l+ oC'elll~bY~!;;lIlf>I:~!tion s!]Q'W~, dear ,i~~ ~(I~I!r-.'ui911lin both~i\'e ii1ml nxodcd! .. O~,~,3A m1d n!}'l, S! I) <l!m~S ~.4).niR!<Lml: mn.~~ i~tlIPqt"UI!l. '~I.~d~W'll(l1~~ rIC~\v;)I!;, fO~lld il~~lil,) C"illi~, r m;QDIJ~ CI!flb:I)IO~~1~:td'~f1l1 ~db;n;~ol~sive 10 Slih (Fig. 2.D ;..m,d.ijgs. 54 <Hid SS~ m. ~'c t ~t;;ii~~ in cfpss. ~I;ldi{]!~nl.'~n~bl}'l'U~~;C iq~

cllil)' 9',5 (E,9.$) e~n~:Il.}'o:-; "vasd~l~tll~iiii c!d~~



neura I hllll!t.\ ~ol:tichtird. ~rl[iJ!lCh~i~()


and pa:m~.ill~ ntc:;odentl~~i\.'Cb;e~y~hc re~iDn:>\\l'it('I'C I,.Ih si~;n:ill~u~y,i,,, kn'lC!~m1 10 ~ ,a1.!~hr,c: and Shih 1ru'~C'~ £,~tci> sueh :11:> ptc_. J .fI11l'1:: hi~'II~1 ("x~~n..'S-.-tcd 'WiS. 2Cm~clI fig, S4B) I n"We iool):!l;."<i on n1e~clc;nl ,oons because Ihe)' are ~ikdjllh~ (tUs IIl<lt gave .ri.-.c lO the MEFsllml we h,w~ a:mll)"L:.OdI in (!Uillllr-ct E~~cltlID-'~{ll~ flC~ i~~t'I<WN!

a."}inrlm~!.ric 1000.d"''''lioll

lit,'!" oo!]l!<li ~~M.'m)u~~din.s

~1!L;: b<1~Cof h~' ccililll:!1l ~uicli!~[p(:~rtC~I~~ 0111~f,lgmh(l ,~imfli Oir ti~~>tilh.lIm ~fig. 2D and 1'1g.~.S4C ':!Itd S5}. TIl is .~ot'IHi""'~~ '1f.!"<11:[Om ion <Jf(mnd'~II~008C .mlCi ill .. !)(af!~ell!blc p,lu~m ,dO!lM: li1(:' Khlln of Ih~ c i~ilun is siniliilur. !.Q dl~n .(;c"![m~Il ~lllllurnd JiiJ.r~ bla."'l~i'!.:lOrlliIJilll'C Fig. 2D !l11~ fig. S LJ ),In Cilllbry;(l' ce:lms,nh~re W.:JISn'i!JJ:'(lvmri::tb.ilhyil'i nhr: ~lJJ~'I:~iln or Pte! in the s<1'imJl 191,n liilldlng; Iik~lyre~ill.cd to dl:f.ll:rem;C$, :in the amoU!l4 orShhsigmlll.'CC'C'hl~>d by cells in the COllfll}~x.lnn leuof cmbf;y(m~JC: lis.'llllc'. The COllCclltlcmiol1l of r-~d m db!! b",&!! of p.rhnarycihasugge$ls :!lmlC'(hll.nisHl ror~!mv il nm)' i!1hi~il Sl1lO' ,rc!i",,:mion. ·1·r.!'1.~~1I()11Orf'l~nro~1.1S ' in~m(l 01.11. ofl~.rinlmy dliia is dwtl,ghl to b<c .regult!l~d Il!tlhckm~c. an.d !PIe~. OOl!!ld :nmctiollrJ!l. ~l'li$ I:OC-J[~O~to il'l1ti,b ~t a :pfiO~c:in~Lrank!kjIlg step' cl;ilic~llof' Snao 'ICli~f<ili)on (/ l). Sh;11) COLlW illl[lCIl:\".1Ire rlC I: 111)0' hi.m:Jin,g lovr !lllhl;' dlirt 1'fl1I lind ~t!ditM::inl.~~I'" imcnl~I~lizmiOJ:I, cll(~ilIdll~ 110111, r Il.lto'olCltll"ii'lD. m mllC(uc!!';ton;>i of [1'1(; I:illflk'iiliil[~ O mll(!unb-rrUiJ1e.lh ,dr.1cm~.ilne w!lClhe.r. Pie 1 tttlhe dlimH C81'i bind 1.0 Shih. weprodneeda IniJio:t~'CS,. ('Cntly ~8lx:kd 'vcu:okm or~hc N~~i!m'lh~rull s;ii£.m:lii~'I,g,


Sh~l (ShhN:-A594), M innnc ml1lO1JH1J1S those .L'l:'1LJim:i m ~ifvepu:!-icc:llL<>.ml1iSI~C(C(1 wi[h, flo~~YFP,mld 3 tlnrtcr :tOr dllil. imu'Sin~lti$e,] 1.0 'cyan :r1~1.0IfC5CCl'~1)J,'()tcinl !(im'crs.1Il.-CFI~'l'(11). Live cells well(! t~sec:lbecause ~hc :il1ler.lUioubctwO,Cn Shit ;:!I.m! Pte 1 Q{IoCS 11.U~. ~~~!viw Fti:o.;;a'linn. ShhN"A594 c'!JI~C4.'1Ill!1l1.~ ,'~~ clli"


of S:hhN-AS94. ,t'tllC"hl!lIdred~tI lorL'Glh'are s~£lm~lij'lg:..wcucaddcd


l;(lnUlini!llll .PilcI.~Yrr .mcl Ctl~J~~

ur f!c~ -"iFP





whi~J:lun~ !'I ...

57), Pf£'1


ff' 'tl'ol1c; did !ltJI. biuldl Sh!iN-

A:594,. flllt! rl.1~esecss of lllliliabeloo Shhl:-4 prCV!.'lll00 binding of ShhN.A594tli.g. S7~ 'liVe 1fIC:;:"I:fliSked \'I,1hed1.¢rlhe ~mot,,'JmoC~iO!mf S~lh o W~dl r}lc I, :iJl~;luenc..~1e ~oc;il~i~!ion OinPl.cI. I'tc~, h Willi Cmli.,'Cit'Ii:rul~(j fti. ci lia aIil1.'r'lrefliinraeI11 of CC'lIs w.iidl SAG lI.1ml': bu~,nol.f1H", i.I\..'ilim.cnl 'W~i111 !lI:1 S (lJr.JC()n~bimLtioll ofS,hh <and.SI\:G ~Fig,. 38), lIhis sl1ISgestcdlhat SM~ bindhlg migil!. II'i,sgerlhc remo\'3 1 oflhe .Plcl"Shh compie"" .fl1omlhc cHhu1l. 0:; fiml,m3w :lPtd pliClduc'Od in respcase 10 Shh wm: ~l{lil IOCldit:Cd iii ~hc d~iunl. To disti~~ll~~i1'~~~ :~i[(;i~ij!il;"$, ..... cilld~I~~d[J:!i;.~ 1'1101

IFig_ 2:. Loc.alj:zati:oo of Pl.diriprimaliY ci1i1l" (AI (J[ln(il\i1'IJ<rlii(lf[lof llooogelilOlJl'i PiOClin cililll of NIIHI 313ntlll; stimu[a,tEd 1>'IIi'tll ]00 ~M SAG. (EO Iloi:a~~1io1m or P-l(PI'IIRPin 1'1(1.+ MElhirdiecleri wilht al M'tl'oIrirul5 Gamryilil~iillil em~ty \f€CWr' ar lh~ 1*1''f'fP ()(I(fii1:g~qoon~~. 11'1 00 andl (at dliat (roo~,~!fg!!l Jlll::Il (grt~.r\I \wre \i'is~lmed by ]fIlIr'rI!J!n~n.uorJ~~~~ Fll!Ictcf {blur) WE'f! s!alll1ledl1MilIDAPI. fa f'U :SIi.o'!irling in :SlillH'espom11fe celts; ,td the

d~eli.Ol1l0:rll!~~~ 1uillou~]L~f nrc I in 11r1i.': cili~ of o 11.1 3n~ll~ Wlrill'i S,AG l[iC3inilC:~'i1 ,aUid lhL;.Ui

!'I~i,jra\Wibe ,~~t),1!l~~h!1i!!id(Jild', plal~ Up), and para~allrn~efm Cr~I>S~OOO['l~ wildoli;ype H~p row1'@I' wntr@l p~'d {Ib~twm rm) m~~~ embryo:;(E9.)~weJ,eirnage~Wf:l.h a 40~ Qib.iedMtli') .Asyfflm.e~!Mic. +c:iJiia'ry l~ari~!iolil of Pkl~1l pi1I,rnXl'alme:!iooem ,[l!lls.1rh:e c~m bGMldl ~Iilwhite is l!Il\ilg,l1ified fl1l~hll!biJttam ~n~~ arr'~iW;';tliidicaoo Ptt:l sta~",~~gl~re~j) .ai'O'Lmtilhe ~~ a:F!~ ]F! the shaft ·of dta "gfOOii),






3" Illflte~aC~].HI'l b~hlflNm Shit:. alnd rtll <;lit !primary cil.i~. (A} iii mil Ptcl art the dllh.l Illi shQ~~1l a :::oJ1flJc.tJllilllla'~!! ](fI i()f 11 !:WiEl {Jtd-1- !Cit'!iA!lr,.lIurededl ,lJ1it.i1I IP~'{l-¥FP (!lI'r,t'!I!IlI~and in:(ulbai~~ with .sl1hN·A$94 (red" 300 li!~tlmUfQir 4') min. Il'!ilo!\~r~if1l·U' {cya~'d! V'IiI~rl!s tfr1'~ dlli!liil1l', ~h@don~d lifl~ ,(i@m€!~,c~U~s,lil'~ (@U oon:hiir,. aJnd t im.et~ sh®\lI rnli!l9Initne,~viel;\l~ ,()f~ne ema. (B} Mean PiI.c1 'UtI®reuenllre in d~i~ I)lf HIIIH 3113~~mls, b"INU~d'whih SAGi WJO ~M), Sl'Ih. (I:r. bl)l~h. (C), Disappearalilc,e of Ptc1, h'Clm: Ilfhm:IIIY ,e;Hi1l 1IFtlU Sihh lr'ea~l11imt. N~IH 31.3 cells prei n{)uibat'edIN1!ih $,A,G for ,2.4 l10u rs w~r'll' teh tid ~o cOfltml. V'lilii!d1i~ml '~untml'€!d) or'imiOO SIlI'i-cOi'![Ciin!m1gl iifiedlirl.iiiiil" rh~ red ,clili$iiled b,alseline slhe\\Wi, the ,HflOlml oh:milry Pl.rlL in: (il!~l:litr'e.1!'le>ill \lJij,till Sh'h for ,4 houlr~ witl'lllut aM4IiObu SAG PI!l1:11i~, Fiig.

Co ~ec(lballio'l'l of:Shh




fime ~H~~~


D E .~
.7·0H .3~lOH




Time I(Hol!rs) (If SI:!rIQ iil'ldBll'Cm all cni~oiff 1r-4 ;In 'oelh: e~p!l):s,~dlt!!l' Ui :iN)(~~hvd Joxy'hClI:~IAl'rO!t, (.A ,!lind :C" w'~:I!'nZ;~Uq(l1l of (m~ (Ii~) and S iilII'~ or Ptcl (9:re~n~lil'l ()~Ih '~r,eiltedl\vitih 10 ~IM 2:0o,~hyd'roi!i:;!I\c:h,ole'fite'J,ol at 7(:1;F'iig.4. AWJlf'!1'!QJ !atnon ~ydm)lychll'lil~terfj,l far'

hO!lHS. (IB) nm::e





~.~e pMR1!~lryci~~i;l!ili!i!i! NIm n.~ !l;el,t$~r~alied c~i:lIh 2!Q!~.·hydli'1o;l;,)f(;IlQlestef(ll.• (101 In(IF'ei!:S~ ~'riP't:c lmiJjQre$(@!fiI~ie ~ifiprim~!V dl"!~ Clfl'e{ trl:l~'!mll!nt. with: 20o:·hydlr\oili:~.hll,le!iter'DL In (13:) am:!1 (DI), ell~1l Ifu1i:rd.:5,holl'l'£ the mean ;:i!; SEM offLu'Dre.Siceiil(!~r'~Om 10 to 2:(1 ema.


swvtched the cells ID ~Qnlrnl 'ullCdillinl (IV' mooi.mn ~Qnl."Ii!1i!1~Shh(F~ ;K').11tt:! Tcve!~q!1~h~cilium
Nm<cill(l" ~,~l;Ik~ll!41~

clKrogcnm!S S'm~n


th~,treg\!laI!: Sm.Q

l\qn~~~9n. CcI~\,I~<ur terol CQn~~lU~~~9n\'i ill'"C 11'11· s

oC9riiR>.1:, 1l~1! S.!llh ~IiQ.;:I!'W!!i,e'f!~. ro~:nt~ dIl,.>t~n!~l.;:I!I!tl~f.lr<l~.!!'i" ~l~!1i5.~~v~~~ 10 Sl1:h. ,1I1d iPXy,'i1:t';~h~ ~Zll.~ ~thl"'iI: ~Lll1:o;;~gI1~ling ~Itll'i{;-d '" ~j~~.ll,.w;lqp~~ld~lml ~h~IJPL'i1IIT1nl~i,;': .orPii;:·l (.14~16). WhL'lll w~nr¢fut",x! NIH 311 cd:l:. wid I [[nrn ~hc: p:riun<"[ty (i~l.l,n~l.Fig. 3<= alld Ij,g. SI'!h (
di1.ic~"i.\l(~ill IOI~d Ptt I iPm:t\;i~ :r,c'V~~f Hg. S III ,~u~cl't~I~J~in~p~i~ 1i'J{Waa'i~UiI. 'of r'~~1 It.nn e il i~. III f~tol~l!C'rt]!C!lInf.II'i in '~le cd!. ThiiO dc1o1 calli~aJ.ifiil wm; tlj~I'y('vitieb11 \.",ilh the ~'tcl:O~~ilOllii; 'IN]d!

lR~I~'(!(ri;lifl~1!'5i!1rildN\i;rt(!~ 1.. P.W. rlll):bilm.. 4. P. McMatJoo, !(i;ello:lj If)w:jl.'Si .• lI1JS9' (:200diJ. 2. P. A. IBea'hr. 5. s, ~ilr.h~rlI!Jlar. II). I'!\. E;eHl)il~. NDUK~ ~2:, J'.~f2@0A> .. ;I. A; J, liltll.;. 14

Z1J~mg. It(. Suyama.

rill. P'_ Soolt ~fS


The 1~1' of Ptt:'1 irolll cilia ~'\I,18 nO'~ OL<:..WCi>li'~

m::livOIlillg oooccnln!lioos of d1'C'o.",y~*,:.ro'! 1!OU· h}!drQ""yohol~~L;'ml. Smj]r;,!!~]idb' lr.Bn'l!nC'OII1cd to

17,1l4!1 (200n. ~. ~. I)e~et O. Neli!!uJ~e;r; l r-enn, :s..IMi. ,wD!i!erI. CfI.iJ :mil, S. 0.. H'LOngfu m' o,r., .mtlW!t! 4:2i1!o., 113 (;io[m. !S. II. ~I.a a .1. ri'. lreiter. Sr:;i'Ma'313. ,629 ~0l(j;06). :;I. K. 1(" COl'~1: l'r' ~I:.~tJJiJtt! 437,. :m~.8 MOOS). , B. c.). Ii'lili)t'lillt er rUl.,.1l'!Ia:5 G~. il, ,,"SoB {200n 5'- ~.'I~~~ e1 i!1l., ~rfJ~"~6u l!OO5 UIJOOI. ]!JJr. ~.Itt (mel'll I. '~\p~Ilf:,tt Eo YI1IU:ag. . M:liLii, P. tLB~OIw. f fi'{!K. Ntlr/.: ArrtlU" sd:,r.rU',A. 99r, 14071 (~Ol). n, l. l !!a~elTib~1I'!. 61. IB.~i!miJ~, Nffl. Iil'no: N!ol.. Cf.l'l ;!ilPc,l.
5iil·l ~OO).

prolcii1l and Ilot uflGln ~mimi.Ti!Jn ,of l!iolilSCC"CIOO I,'tc I~YfP. !I f:r[t" l1~m~ !Itiultdmn ~)rol.ci~~ S7.B). (lJi&.

O::~Sl'. '~ and




Sll1.u :locali:mtiolJl

3;) il~ Iile s.ru11C C-"'I)(!I~H1~nt Bcc<i!la.w ~11e~Qc.il~i:!l'!l.iO:I.1 d'lllL11~CSfor fu::1 ad Sma. dl:~

were each


in >!':O% or~hc

vii'IiJalil'Xd. :t~c~vd=, of PIJ;:~<iJ~d Smoi~~ (·mil '\,\~L.'!fi.i imIC~F.'icly CQn'\t.'I,L't1.iI. TIh,l ~(:ip~,d ~hm: C~Llfl!l.'" of ~e'~ di..;;app(;',<!!mm:.c S~no '~r"" and pj_>::_Ir.IJ~!OC' <Ill ci~i .. ,f~t~fShh,addil~Ol!(Fig.'l. Ie unlCi Jel' ;.;u·PJ1Ort,~ :~nO(rc.]ill \'ihic:h Shh 1:!i~1;'r.> ~h~ ftr!I!!Jva;1 of P,rel ~~tU dl~ ciUil.lllH. ~nm\'irlg, S1lt'IJI ~o {lule_~ (UW Ol,crh'zlC' s,irgmll ing. C:Or[l,~j~[re'J:11 widll

Ith-tprn Il'i~u~ c~~I,ilU"lWiU~ kiI1~li~ l!lIl;;JI1 \""'tl.,~ idi;YI~iC!il loiho~ M.~I~ ini ~d 't[Jt~uoo \vinli Si'liG or I~ Shh (.Fi~..4, A afild B,~ndl ~!} 59).. TOC;jllt'l.~ll:L wil.h ;ll.I,,)'drux~ch~l~~:I. mn. oxy!llerol ~lUlT does Imt aCli.vm.c Ihe .~ paLl~""!~y,i[l not indMcc Ih d tmm:;lbcmion ofS~no" T.hi!l;~l'S\lTI ro\iidc;" <I ::ij.iC~ 3. 813 (;ilOO2}. ci~c m{j~ooula:r' moc'l~m~u~m-Smo lrn:n.sklc<lI.ion 1::1. II W~;OiIfl~~ ~j' aI, ~IifJqpmI!'n.t' HOI 1(125 1;10031to ell i,t 1.0 'e:O-"Wlklin~m\~'(I'''~l.Crolt1It!Sl~~tcl Ul B. ]..~E,pilti!:, M. ti'.:. CcKiiPl!r, T. M:il~~ A. BNCh/l', MJr:rl~ Iii. sign,di ~!g. 418, a.~.2: GOOn .. C~II~ lre;)~d \V~Il:l.40fl'-.h~'cl~;l},,,ydl01""'1~1 1~. M. 1ft Cbr!J;,p~rN 0'_, Not ,GieJ'li!'l. 3::3., 508 Q2Oim"

1.11i~ j:dC.ll~ c.:::II& oflht: '!.1('1Il~ID]i~Cur.dliUbe3nd floor cmllsc they ~ikeiydo 11101 illL~d~! ind! to SI'I1ICll.l6). d b pl8ite. whid~roccu\ e lllr.!Yl 3IliIOunlS of Shh, (),LJr results Si!i~t!:hm 1.'i.cI loc:lliUl~i(i1l w shewed high :revels or Sn1.O mld raw :l;C\,r cis of Pte I ill. d!~ r~ {ng. S 13 ~ Jill! l'I'~ovcm;Cllll of PIC~, prillliaJ)' cili~ ill.hi.bili." I,hll:i lil palh'~'3.y by C'xdudand Smo lilt H~1i!ihum is anll~{lgou5 U) 1IU::~~1U'1l- il'lg Sumo and ,[Il-;;o 1!~loV!'Scrliia iI.(!!. ~il~clkm as ( ~inn in Dlw';,aphiltj, \1I;:~I!:,rc'MI!1l1\l;';1}' ;n~v\(~ltio!l is chCmmCl1!;<;f:Ir.< 1 r~rlhc dl.'l/1."CtiO!~~or c.'\! r.u;:C'IIILJ~,~r Shh. Bindi.~g of Sh h In Prd w 1'IJ?~!lm!)'cilli:! i!l, <L...~imo(j wi]b $Om!":! JnOV()mCI~t w~h!l:r!~'i~n" .~m:.Irn.lbrnnc: nbO!\!e~~;lenl p~c a\lf,[l)' (J). illld ,or coupled w pmhwfliji ocli.'.flllicl'll by ihc reciprocal Pld :I!nny regu],~~tc:1~1O ]k)!l,.Il~rn.'1!lion!!hroul,gh H! m(l'\~ClllC'JiD!, of flcl ()iUl oftl~c dlia ~ml S:nnoimo S ~he ciI~l.a process 'dW.l IIlay be moo i[li~ooby ,iHm~n.molecule (/.1). EeCilllliiC 3[1[0 tl:lltiil'ocatim~ ~o I.he p.l11T191)i ci~ i~m <IPil'ill.'m'S m be ~~c!'i~i(1df-ifC]) oXY::ll!Cm~~. EluciotllinJ;l '~1C'Ilm!ccUII ..r 1!11i4!Chi~lC!'}I'

ci!i;lu!!D ~~ :P"Mi:;itn friil'!l!~'~'lrltl Uilo.la~'t!nin·~I~ lliX,~lcdwith SAG IFi,g. 4.(' amI DJ. llln.!r~, oxysl~roL'l aPil]!(>'Jlr fL!I1Ctl:On not l~kc 10 SIl'h. l1y Cl!!Uing tl1~ remm',d of fu;d rron~(:i~i~!, bul aia Il1!O~ (In..\n:AA,i(l·i!nn f;[",'fI I.Q'riSl1':~ Smo hl«."l1r;ili\~ ~ IIi~inlll~bitoJY CrrL}C;l\~or'te l.. H,owcvcr~~ys:1.cm~ rUJlCtiO!ll d~.rt;C:~filtlyoom SA.Gbef
011':;0 !1."1):l~ll(ld!Pte~. i~l

is. it ,9. C.QKQrtlm, ~,ii. P. $cot!!, hor:. ,mtll, AiMd. M tII.SA 1O~, $Il@S· ~OO!>t l6, I,· R:. I~rel flI•. J•. ~ , (ih~. :ll8!1,8~SI}!f~II01h. 11. M.f.l.S. i~ i!nlE<5<iig~t~ of t~~HO:W!I~d: !{Ulgheo;>Medk~1 ~n Im>liMi!' <lInd i~SIIRPOiilOO I!!i!' N~~Mn;ll cancer rMillJ~J!; grani !'1(~l'OOBe:OMt I!UI:. a ~bert Blad: Ff~~ of.he h
~U!ild~()RG 10NI6)- Wi! mI~lI!rQI 'Il'Mii!m, P. BNt:~li'~ul'rni"" ~el!ll;:iIIHIlrl:lrlbiili!1!!l res~UIJ belCif\~ JI!I~~Ca~ir;)n, II, C~e;rI fo!' $,AG,. 'C)'. IBr~r,dm~!1I 'ror 111I'I<!~~ ~o~'I!MJI)i[)~ (;am~~


lI1iilln~ It.

(«C<<~iI1I fi:!r di!l(lmloo~ !lit

~ilI~lly$k O'lm.l~~, J. H)rmM f(lrjf mlcm!l~~t:p)I.,d!dU. It JIlllrnoo ~~!!! ,;, S!l!)liUJ1~iOJ ~l ~n'!i5.u~!'!\

H. '!ilama;hl fllj il'llrl'eFcSln ~tJiij"1l!, Iil., KiI, for 1~1! I'!C'I.;,YF~ ~OOS1jlU~n, Salk {01 'I~i! !inn ll!bJ!Mllgl ,A. ~tr.llt-gy. ~ilIdl 'If. WIBn. CHe .•A. ~~mlar.,aJlcl iB.almiJimCC(' A

:S!IlIllipo.rtingl {l1il11~:I11~ Mat~irrial :r.WJf.iilijM[l!m"Hj.ar9l~llil\:ooleiliVrllill!'3:1.7fS:e3£;r~7::1:!oc'll

i'~~nQtivtJ.tiQI1, tl n..~lIl[I:to:ry slina~ I

wouldbc p~Nl:i.cI(\d: [0 ('olllro~Ihi"
occumrillg oxy:;t,(!iiOl:s f.ulcgotxll

lIlId STiI~[]' lmnkk:il1~ ,1lU II:lrUlfn,ll), :rll:!!1:]~el!.l~'~ [l-IIHtmlln:~I~r!c~ N.m:tlill"d.1lly cHill wfl!i ~ilk~J~' provide !leW 1i3!(gcb, for IlIiI(x!u~ac3n.didmcs fot tilnll o:flh i~ h'Jillli)ruifi~pillthway.

llialerii;!(j .i!O!I t.'i!:,1li1od~ Itt9l' $1 trm )'14
IRde.mn~~, !II J~~ ry :t!~l;"Wl~!ed ;10'~'r' 2.01iP'

l~l Uilm.rSCiE!iil~~~ 1.097·110

·Y,ar'g,etinlg .by

Host ImllmUina Syslslml Gene

Vir,al :m'iRNA

N!!J<liIi!~ Steom-Gim®·.s:S.ilif":li,,, rl!J,i!I,HiIliIJ I~~elfan~.:il. Albe~~:lilllli!l1l,eli!ll~·<linlil~~ Dalila {],. WoU," N,i:'II'il!llSa,lef~,4 ,M~.:I1he Bi~i~Hn~:i:·H ..d Mf®!rwitz,·:u. Z.<!ifml.t P\mkocilJiler.~· Mark pnd~ ItiI~~ s ,(j;allb:ri,ete tIIl.. l'lli!~ 6,t I .. DeliJ:tii3J Go'ldlrrIli3JIi!,-Wo:h'I,:or {a rylil G~eelnfi,etd~' S,ilmclll<!, Yilgel," Hr"lflmll!lt Hellt.~:el.•~ V'<tlel ,Aihu'IIi<ll,;:. liillllilih ,Millr'~lali~/l~t (lifer ,Milndelb'Dilnn:~*

is kn(w,'I1 10 I~,vc evolved cm::C!UVC lm~ mnmre C'o,I:<!:.'JiOll sm.'Ltc,giC'-s. cIl£ooing m,m)" ir:llrnullloml;)(lul!'IWU-)I 11ooidns, [h,ll Illmn]!miale the im''ll~!~~C re~('IQnfC (5. 6)r. ~l is ~lll!'" OOIH)c!",.hie dHlI. 111IiltNAs clloodcdby UCiiV~V (1) m~ghll, be CI1lplbiacd dll!rh~g ill.mmrlc Cl!1,~Oa"To iIlCiillhi!:> hypo~~C',si~, we oo'Llghtlo .i(_li:Hlify put~lllliallnmmn ~'Ii~C'[g~IJ4'-S (lf~hc :I-!CMV .1~.li~N.t\.."ii hi' llL<;J'ngmUir .~!~iwb \'~~op(:d tm~"'1 Im;,'dic'[ t:ou ~liJil!;;uitl/Q:m. d....
·Hebr>eWU~il~ HI~dl\l~h,MIedltdl.%I'!O<)~ D~lem, I~OHi,L ~l'1JIiIl't[l1iffilt IiMdll!!!lilaJrG@~tks aool BifiWChoology:, HI!br!i'!l\l

i(I ~jCMV)

aJf!d !!ppflea ~tt~ hl!!m~i1 cy.t(I!i·n~g(lll(li,llili1;!i5 m~RNA.t ~~s!.ll't~I'!!iJiJllI 't~e ideiFll'irkaitl!OJIi ,(If ~e' !major ~i~c():mpalibi'lily >ooml!llex dass H~tate~ dlainl IB·IMlun gre:n~as, .allt~IP GaI!'!did~tel:ar[l~~ I)f ,honv-miRun12. MI:C8 fl5il1~n~"il\'dl!U:::Eltllig:;IIi!~ alf !he iIiIatural kill~r (INtO a·c:tiv.atilii{jl receptlN" INlKG2DI (lll'Id i~ {~~l icri:tie<iL ~D,r~he N Ii:: ,~ell ki~lil1i{l Q:f vir!ll!l"irdede~ c:eM!l. nd lllllillH~r [e~l:s. We :!iii! 01':1' lJI.at. Ih~ll,vl.miR<-~ILH.!2 a
~peciir;{.8.~l1JdQ1l'j'IiI~9ulate.!; M~,CB eX.~~5sirllnl during vir:alin'remilln, lea diirl 9100' ,de~re~5ledl binding 0,[ NKG2D inl'~ rredLioodl ki~1ii1i19by Nt( ·(ie~Is. Our re~ru~f!j;rev~~ i! miIFlNA-lb.as:ed llrtlml!lnaevaslion . li!1~dhi!nli~m (hat a!l~ear.s to bE!l'xpLQilw by' numalm c~omll~1!I(!,virui5.

Virally encocl~dI miCfQRNAs "miRNA5» hl1l\l'i!fOO~.iltly b@~n di$oo\,l\~l'I'.li!lin, 11~i1Pe:wijl!is.€'~ I-&olml'i,l'f!r; thiltu ~riol~gr~cail'lJlesal['@ m~tt~ l!!nt;:;!'I~~I1,We d}E!\I'(ll(),~~claJll a,lglilrilh!i!1 f~~'m~pf\ooi~o~ of miRNA t~r~ets ! '



lor '~ne",,~ a~ 'VwnQ' ~!'fm'!!.!J'!Otogy,

'l1Iil1d!a~!IiI MedicalSclio!il lN~lejif,i. ~~i'aeL ~ v ... ,. H:~b,IilOJ1 IHe~me n~rs~t)i~ IMGZ2S u OO~etdolif ~rrnllliW. ~'D~p:!rlrnentol: aimk.JJ Mi~rolliD1Q!ID' Ol!'!;d ~n!e.liu [lii~m, tlad\1!s~1!i U:n~W!~lt.Y H~Sta~ jl;!~rn, Isri!e~5DepMtnrum~ QI rledia~IiiCli,. IiJnLW!rsl~ 01 G)o~Ql\l!;ry,.'bd~~t! H!l!!r~ iluae'L "T~.I1'5e authors ({I~:tniool:fd!i!qu2ll~~1DtI~s l'tDrk. tPfe5.ll;mt a~reo~ KlinUiUm Ing:oill<li,ili[, 1i'!~m.ulh1f lilOO· !'@1_Qr~~rnedi!!:i!1, IDM~' 1!!'I9~~d~, Ge~!'1l!lRj. t1l'o Iylmm ulflr~oo~lK~ ~n!iWI ,mil! aMr{l_~. E.JI!I!ili~ .AJ; ~1I!'Iahtrl@!kmiI.l'iuifi,ac.u 1l!!..M,J~
7tD~~!'fiIl1l'! ~

t.iJabarrn~ I.rmililjih.illll,.A1. ~~31. U~,~IIW:m.rn hil11nl;;tller 'lmijun:. Dl6pMlI!1~iIt o-f \Iii~.Da.r(ll36 Mumi!:Ii1,G~aF!)!"

~.rr~iI:;t~ng Ii1RNAdl;'g~tdwiolii
~)i biL~t:-pii


11!I11Mnily Hes,pUa'l rum S~.oIU. Jelll~~" M



il~NA$ ~:·on..;';liitu'!c <lb.!~ r<l!mih' (If (3 'ILITR~~· (i ,. The sm,d~ :nQnc~'u~g RNA(o; th"~ r:.f;lll!]~1.C cnoO¢k.~ mi~:tNAs,

I'lllceJl~ 111IDS~y



V Lt<lU¥

in Iterp~\·i!ll.u)1es~ tn·


~I'rd 1.llllt~hJ!tPrl'l

~ri.g~ui!1gl¥~'iilllggl;;;;\~S t m~RNA';. :!~'lay lih.. f~l:!.etij~~l ~n .ifilt:C~!J(.'t.ic;ll n._~lIl~~i(.ill .ill\,nIIo'ing \.i[od n1iRNA:>
;il'id ~1"Slg~lit,:'i, f 2'-.J

ir~I:U!!liO\iill]llle3' U[Urnli~!alI;d rcgtOIfiS,


11Iillti'iall c)'['C!;.I1.'ic!!l1ulo\,jru~


;]dd!i~imm siflgh: hj nd:ing sites by ~!:)ing the ~ l~,culTOl_:r (7l Bfien~i, RcpT ..r lro.:ilu~s ~i!iSls illihe ('lli1~'1!'\fmiOJ!l~h>!1, ~I1!~\:NA bjndig~ ~ti;'<l can !I',~~~~ inCQfI!lmtioo Q'buJglttld by !l!cpr~r \7). Thts ,,~~g~ ~"\~rol ti!l~~i~ t~w(!l~fl'i YUTll It lt~~~r~t~ ~i!hn~. !,I~~~!w '!IKl~ o!hlJl' .~!gorinhm,'" 'F<l). is ilt. <,~l1:;:h~~fQ!" ~p;::ti!1~~ ~k~I!IIi;!!I:\' ijHl Q,l!l!';b,J'llTR JqJi,;"'itd~U!1 of '\,'VOI~!ti<~ry OO!h,t,'<r\"~ltio:rI of ~w M,'q~1(lnt1": ~x! I;;YJtLHt[1;,';S dU;;':i1;: ,~I!J::!H~:IL-;; a_" P;ll"~I~billJiHlg t\flt'S" TIt\llfl.~frut::, tii i:. mm~ SIllut;lhl(i li:nr '~,i;dmiRNA bi~ltH(1g~!;i.;8.A 1."'Ql'I1pleir!lU!IiI!), niOO~ "rtdiC~!~lg~~Uf[;l~iS Gf~h~ l'G"'S cvokllimimy O~!1· ~Ie (l! flh~· iil,g>otilll'!!¢'!., 'cRcp 'f'~f,. is'C'ro~I!~ rOF ~n'Qd \·ir.d Q}:ciRNAo:: (2).


\\!c ·ii!11IdhilIQ ~~eI'lTil!irand !'>l!b5Jcqw;nUy d.~,q~~~·to ,,~~ hl!~m!n 3'1,Jl'11,;;;. ,,~[1;;hing Ibv .... JI)OR'l1~ii!l!Jil1di .. g "it~ Or~'N ~.IHC'MV miRNl\':~ li;.;kKI ~I~ mjI1,EI;I~G 'W {9,1.. M!U~.n~ il""'lw~~!llnlil,;'hlih,:d



Lh~ hig&a~stn.tnkiUlg

prcd!c~~-d ll<'II'W:::~ .,~~d~Jfli;wr 'IFI!];ldicliQ!~ fOil" hcn1v-rl'!~R-ULlI2 (fig. ~A~ M [CB is ~ ~Irt~



mIB.!UIL.11'1:2 3'


At] GCA atfru.A





1IiI~R.~UIL112: :1" 'C::mACCWMMmI'1i 1 "l!HeA, ,5,' G~OO1I\1JC![!~ 'I:IA!I!."


.....g;.... R~Cei'i1m"C¢irtfO.l




~ 5 2 1 NKG2iD-19I1Collt~a.191 ~1!.Ig), O~



R~O·m~Ifi·CQI1It1Q1 3IlI1~NKGllill






IS 1:0 S


F~g. :1. ,hil:I!rI\i"miR"!JIU1ll2~drli:ct!~ty ~own""!l!Qli!l.atll';~ MICB ~XPil1!S;s(OO and ue~U:(jes,NK c.ylDitox:i!:ily. Fo:rall. p.i!nle~ IlinE! rejpr'E!senti!ti~ ,ej(pl!r~'rnelmt ii!> !!hown 'I)li!'!: aill~~'$t: ~ree IP~rll(j'n~d, (Allle 1il'I'Wktoo d'l!ple~" f ~Olf!\I'"fr'!iR"UIl1.12 © ~r~d) a m~m tlrgel :5~t:e~b~lIIe)~11 3· Uiil'frl or M~CB:{kip) ,1IIne M~CA,(bottom). the (8) Ed:opk e)i:pres~~onl,of: mcm¥-rnilR..un::l dQ~'\!Ili'fE!9Ula'II'S: M~CB ex;p:re$lillclll" V~riIjJLJS:hillman c.elJl ~ili!ls Were tramo!.!Cioo With te'~tiviln.iiSes '~)I)pr'efii!l'lg GI~P ieoi~he!r w,)!h thcm,\I·m,lR:..1Jj LU.2 (bllack nlli'togr8ilil1l) er mi'R,-c:cHllrol (.ofHm S]iray ~i~£ilra m). E~p;j'\~$ii(lf'l ~ev.elsI)'~MtilC ,e:I\1$$I, MliCA. a !i'lcl.l\~ reB we~~ a~5e$'5edby



@xpr~sS!rlrlg Irn~IR·wlillrol hl:1f11'lr~rJIi~IR..uS:s,.l, er Ih[mv"'l1rliR~~lU2 Clo'iIiIiS t!:5Sess,e~ b].I' fAl.G· lJisifi9 NlK62D·lgl and !he (]orltl'll'l W9'9·~gI~Contml["'llg~ ilil varioLl!> (O~(fliltratioli!$.(E] The rooucedl NKG2[j)~19 brlldf['lg $5. dl!~ .t© rduCioo MICB iIlxpn·.si5~olll.llne e.;lj:pr'e1i5f>l:QliIll~lQfthe'V'ariol.l:1lN'~G .. :D !ligand5, W'a~ a~es5i1ld b~ 2 1~A.CS in RKO ce~~~exp.res.,~fng mCl1lI\I'"'fIliFl·mU.2 (o,pel'l red hrii5~egrram), miiR;-

(o~tl'ol ~op@nl grtliy Ihiil:o~r'am}, or 1Iit:I'Ii,~-rn]~US5,·1, (op!!~ Mi1!~ik hnsl:QgiFa!lm~. The' h~sllO£lrOimp~a~ aN'l :glillm only (In! ~e' GFP'·plli5ntiveiCe~Is .. The ilad.'JgmlJlnd ~~as measl,l~ed !liS iii! (S) !l1i!t1 C<:1 (Solid 'gr,a,y hijs;~g~anl~. ~dl,l~ed kimlllg 'of' (F) FAG. () Ecil.Opk @il:II,f@~i~nI)f il.®1.Ji"miR-US5c=1 ,~~, 110t aH'@(t MI(B IRIKO 'iC!!lls '![.'(p~S;ilil~ h()mv~m!IR·UUl12. Bl!lk NK l!:@~1s Wliir,~ IP~~lI'!Cll!baitoo erilhl'~l,\\Iji~ <ll'!tij·NKG21J mAD< '(~!.!hliite') \JiliUhfi!io'W~m'atc:h ,cill:nlro~tmM (grra.v). or 'E!'xp.re~in.!:l."liImal'l (~Ul l]liles. 'L!le~E!' UilIfllSd:u:oe!l \'Iii~h b~'IiI[Mrtlise:s !E!'}i)J)re~in.g 'G~P' and eiU];~;r h(m~{f'rllill?U55~1 (IJlack l!tiil_ngr,am}or illil1l1Hlontro1 >(open, 91F",y I~~lbel~ RI'{O ceil:j, exprre:.s.simgl Q'lffiR!.(~I1IUl1ll'(If hCJIi'llNnnIR.!lJl'L12 wel'lt thenl ~i~iira:m). The exprl'ssiol'l leve~~ ,of MICA and MKB were o'l5se:ss;edl by FAC). i!ld~'edl a nclimllb.:i ted (or '5 hQul'5~t ~helJildilcallcdl e'ff,e<!lQr:lar:get (IE:T) rol!tios. Ihe l1i~tD9rillm Il~.otl!'of (8) and ,~,q were gal'Eldi OInt!!' orlthe GfP·po!ffiliive lQe!l[s" 'Whe' din'e'relll(;e.~ DelwflElIII !he tkn~tilil£l oiff !Iite Il:KO' o:llis e",p~eS'5~ng miIFi-Jl;o.rrlrol Ela~k~ml.ulldl ~ev'llls(i(lf(~~i!ftd (0 we:re me~,\!!t'~ by ILls,log only !he' ~e(.Qngta:1Y ;j!1fIJ!!j UWiS~ I~Xpr@Ssilillglh:(jmv-m~'R-~U.12~rl th~ p~s.eIflW ·ofth'll is.oilYiP~ I 0 (y5~Dn1il!lg~Md Ab {gray !l!ll'litB hi~t:ogrtllml .. (D,!' 11Ie:dILlJ!:edlbindffing or NIKG,2D t:o, mi'!~J:hedl ::onbcd mAlb'~n! sign'iriirillnt I(P <: 0... :1, ,t lest). (rmr bars r,epres:Etlllt IClel~s: e:i!~~essin.g,h!l:I1i1Vl.mi~·UUll,2. Bilii,d~IiiI'g ,of NKG2D';~g to 'me RKO 'ce:11s !It,a,llim:nd deViialio:n {if rt1ptic:~te!l.


j:ndyo.;C! :~~~_nd(If NKG2D, ;] mlllW_rnI~;iIII}er(NK) D'if(;,'l;1b.hn! il'lg of hcmv-mi
~J;:li",.ning~Qcp.l;(}f ~"\pJ\,~cd \lH ~!m{l5~ tdl hll!!~ .J lJTR or M 'A and M'~C'U '",,'i ..;; Sl\~ icdby ~~[\:~I;L;'~~ rq'l>n!W-r ~"''3l1ly.~ ~11H~l=1 eells ~iclli· ~~>'I!~ NK 00!1:;; ml " ~y,to(rlxi~ T !,}'!11,phg~y~~{crLs) ,( UJ)". Thl<lhtlportJ~:tCf; oFMI[':j3: i~~ ~~dly~~p~~inM: ~~dU;:f !lC;:Qrn"'''n~i!R-vtJr~ or ~niR..qoJlItrut ]~:I ~1t3~'C1l1-C11ilW.iiththe st3tll'litlg 11;:il'K~in~m~ulC: ~1",ml:;t':1uw~ill"'>II ~.cMV~nl~cii9ll i~ (ilI.!I~~ iFi~. !Bj •. a m",,"h.."I.urnMe dil;!cR.'<!.."1Cin !Jj!~ :lilil!b!ltinlti ated by Ilhe sredne clowll'"",-'gU!!;;(~ion of M~CB ~!)Im~c ,~.",pn,-!S!!igll via [he UL ~,6rm1,l;;i~1 acli.v.ity ¢r~h~ h.lci~~r:;~~"p!l~r ,gene ~~~ otlof 11l:CMV (II, Ill, MIC A. 1~,~!j!~iN;t ~!!ft,~~.i~~ s..;:rv",d t:l1l~Yill;'li'I"tiil'!""i.i~ n~IS!.."d lonhc 3'UfR tlf



I~.)I!JL 112: ~Cl111c

n~d~<!tcd tim;mgl1l Ihe Ill\;>dic~ed b~nd.~ng .-'iutc,\'l;'(; ~Il~ml~d two ,d'Q~!'l1I,c ~~ibfi'iDlII~~~1l nU!~ltfQI1~. dl~~p!~ngih~pr«lict'l;-d~ifi~g ~~ti.R-Ul ~iz ~~!lbd l'lJln, 'ill~


be¢w~f~ !lIi;:IV!'JI.


Both 1~~lk[ti(l!ll:> ,nb!lllis&licd tl~~1\.11n;&"iQl~I!IicdilillDd, ·byhc~~'~'l1I~R-UU 12 t~,g:. 2A). The bi~~~ing ~i:t~

diliCi.-dIVg."Uid of
IJ~~ 10jl'


\~:..SiL~Oliili'k\-'tI fuUnOJlg

pilt'.d:h~tC'd I:iIrgc:ms: of bCl1lllV·i'irliilR-UU 12 't.F"i~. ],A), Th~ hcm\l~fI1iill-ULlll PUI,1iti\[,c: biudil'l,g, ;,tn'CS of both gCii1C$ 3re!tlf1~o:M idcllilic!:Il 3ndm'!l: :I!oc.uoo with,in iil his~I~,Y :ibniliu b!JIl itO! cvol~ltimwll,~ly ornl~~I;'OO ~7) 15·O--IJllloClCmtdc fltl,} regina o:I'nhtCir YUTRs. TO~~~'$islhc 1~!lnC'~ioll onlClm,'·mi~l"Ul! 12, W~~JI:~!'i.:-,&YI;;"d I:~ai~miRl~A ill l;'a:rl{~'U8 h\u~lm~ !JU~liOl' cell liDl~S fhm cradog~n!pu."ly (i';ql~\~" !I;ln~i\iir-<J!~ \r(!-'CI~ lhcm'!,'-,nliR"W_1 n mtd lWoO con!rol ~,'('o(;l~~ O! J~O!1l-!11iRNA~'liLI~~~100 (miR· cembral~' .. !t<dher~w-nliR·US5-1. 11m c:xp-rossiml of hCJlil.\f-:Jui R·1Ul.~ [2w!ls ,ootllfin:nc'il l'y t(luamlJ.ti\leUl'4l]~[:~tllCIJi}lym~r.NC eha~.ll. n:::lcl:ion(qiPCR) r

~,Iil'd ~iud6(jl:idc in !.."O"~pan.MP'I. l:(;!1\,il](J\ ,tF~:!} ..1A !ilid IB)~ To t\.~lwhiC'rh(:r the di'Rctc".i'lcG MICB i"1I:]dol]ly in. cdb'i.5(prt--55ing Ittlt)'''·~Hill. M:lCB wa~ cl~tc ~o Ulll2 (.F'i!!j-, 2.1\). To (h:,'lI,ojj:;.l;~U~ .he {fo\Ytl,. ~bs:eiiV~dOOtWtiL':iI1 .Ivl[CA .m~t~ Ihlil iiC.\.t~llljjHultlof M ten by hC'iml'IiI"i R:·Ul ~12 ~:;, bil!! e-hai1!l.-c im I;hi~ s:iUlgl:c-m.H~loolitl:c'. we ~~C'I:'H.ed

'h~3'lJfR nrMK'B h~,'Olll; aclditlnmd





nnd [!!.IUCHw~~hIhe



!Lf.I1 ~i




IJmili'lrC~ ID miRULU2

120 100






-- rr-








01_g..s:~.).llu:: \'ec~ors c!ili'lto,nlflo:ib'l"001 nlll,Ql:'eSCl.."J1D. f;!'rotcin lGIFP) for nnof:lill!t'J;rin:g. the i.ilfocJi)o(n ci1:iGicncy Pl NQ dBli!rcnre ill~):'c Irnn~dllCtion clticicncy Ofll'lC' diiTcI\.1I1t I'Cllll \'iI;<I!1 vectors \\1<15 ~le<l$uroCi~l1,g. s'2}. f\~l;~!)'si~ On!1C~rmri:ous~l!~lOf cclh1, lm!lMh.!I-c~1d \'\1 ~dl hCll1iv-miR.-UL 112rcvt"1Ik'<i a Sj)l)ci ~icand cxl~ll,>i\\'cIlCdltc~iml of nCB and lin,lc ill' no l!lJdUOl~OI!D of wI[CA (filg.. 18), The dO~\'JN"e,gyJ"li'O:Ilw~1S :specific to M ICB and 0Cl il~11\'··rllil·1IJ L l !2, OC"COllI"-C!~O ChmlljlC' Ii 11 ~~C levd of m.-jol" hi!'i'OCl{:O.Dl.lp'llibilily mnlilp~ (Nfl-Ie} daM .. ] was O'b~lYV~d ~F:~ ~. <1111,0 iB~, '~r;m1ldWlcllml

~. (Ij, -;rn,

~ ff.





, I I

G'IlIG». ~ CAcm!:)I'




w:hh hc:~ml·nt'iiR";USS:-1had I~.OceUJ("t"D.,Wig. ~C)" TQSlli:dy whelhcr dlC {)bs.cl"oiUrl chan!i!es iUl M KlJ pl"01:C~iilr.c\'l::l~ a neeDOO ilS illl.tciI'<'i.cttlfl widil l NK cc~ ~activa)ting reccplfir N:!<:(i2D. VlIC S:ta:nmxi RKO cd]» C.""'ilut:S~i~lS ei~hcr'1IIliR-{;OJltl'Ol hcmv~uiI~_:USS'-l.o:r hCI1W-lUill-Ull ~2 whh N KG:1:[) ~,i.JSCd [0 ifl~l'l'nlllbOg~o'bIlilJlial 0.11ngGI). asprevlolilsllY dcsail'JoCd (n )"l:hJo~m;;'J,>acl:i,vluecli cell ~or~in.g If A' Sl analysls revealed 3, me'l!;~n:;lb~,c decrease III N K:(j2D~~gsmining of cells 'C1o;.1)1;'C';;;S~ i:ng hcnw'lniR"UL 112 oomJ~;u'Cd wfruh thnscex~~~~i~!giJ,1;:'ln':·m~_[t·U:S:'i-~ mliR",-lOnl~d, ,~.~~g, Of 1IJ), TI!~ n"dilll¢I~::m~u N KG2P bi'I,(i~n!il wa", !i~K,'ciUlci.'llll}' du~ th~il\:~ll;i.;cd ~c'lcl~(i!1'MI:C'B. 10 bC'4;{{u~ the cxpr~::;..<;;i{lml ~)Ih(irNKG2DI ligands """fJj,~ not 1li1'l'Ctcd IJY~I.etilv-m~R.l1L1 1.2 r(:Fig. ~I). ne ri.i1llC~iOflil~i~nplk:iunO:lil O!rthch'fIl\'-llfl~R~ U,1.11 J:-rnedrim~>d Il!;.'dnu.km un N:K.G2D biilding Willi dcmjJ~.biml~(1 lJ;yl~,'t;;'-lsu.riug NK ]ly.;i~~ of [~KO {lells,C'xiPac&'ii~I~hCl'lm.v-miR.Jl.JU ~J rliliR~ or o:mlro:L CcIIL~ c'.l>press:ing, lK:m\/-.imi:~,UL 1.12 were ki~lled less el.nric.i.cl1t~y thml cells e,'\;.pl'!:.'S.·;iulg miR,crfi:nlrol (Fig. I f). '~Vhell NKG2D illieractkms


~ t::GGACCUI'l.aAG'C(H'l mlcrM.S' aO~~ACC' CA~3:'


'J/' 5'

CGGlM:lCl1"HliMm~ aO~~01C.l('

~);·tJrTFIMICA t~,IB

m~·UL 11:2 3'
MUll 5'



~ AJ!l 3'

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~\'ereb!klCk.ecl wi[I·111I1IDlmdomdmllil:iQdy (m;\,b~ a:g..insl NKiG1D {;;J!rni NK:G2[]t),km~l~g levl;'!s bnlh ~II!ol ;,l,oICf() sii,,~hu IFig. If~. iJ1d.i~Ol,tillg ttil~u~ llii~ rod,l10~ killiJlMW11L'ii ~xpli~i[ly ,C1I[>C [{iI't'dillltiL"d


~S:latJiiSd(~l!.y s-1gl1l~~kg:!1i~ dfHlt1!r'~flc~ I;i~tii\lee;n~ruls, ,~pr~S'S'lllgilflil!i~(ooIUol OiIiI1Id! tl1(1:sll! ~_p~e~f1g 111;"",1,\1'miR:,Utlll1ll2 (P <:; 0.1:10 5 by t 'test!. (18~ Sm@'lI1IIillth: repre~IUI~:ti~1fl (jir the I'Illlttlli(ms ma ~e ~11 MICBalfld MICA,:r~J'ms (b~uEl') and theqr bil:1le·:pa1ri'ngs rrtSlh !h(D'I'Il·liIi'iR·~L:tli!,2: bil1dJ.. MIl~aled p(lfliili:Dnl'l ar~ y~,~ef~!fl.~dI,. r~~R ,!l!I!!!l!~lysis;~~: MICIB, E;(p~mtll{!t~ were p~d\Qnn~,d tt! Rl<O'!I~lls expfeSSi!ll~ '~~~F (I m

m I~'ve,e:ll!periment liS shO'l'l1l out ®~ two l~err{lomled.(A) ~G:I'IlVJ. :;;l a [miR·U1ILU~-iIIiIedh;l'led r~pre~i()il r(lf llJrn'fera~' Jep'I)'F~[r 9~n~' ~(tlvf1y,Th~ 3'U'I'IR.ilf' MICA, (30:3: IiilJilIl1d ill 350"!l~ seglmernt Cl,1th~ 3' Uii~,cd Mice; (hllc~uiJij'ng the' pre,dided hcmllif"riRHRt12 binding :sii~) W~1'e f!'!s.ef~~ d~fiiSt~~: O~d---~~~--~~ of iii JifeJIy Iltl(;]ftlt.l,~ rOplNI re;iil;dimigl 'rr,arne. In,eo ~~9t!re d!l1l1r1olll:~trates: hu:if,era,~ al~tivityartBr tn,eindlic;ate,d mlJlCM~R'o.1 mAU.11 ~2 ~epOIffl~~ pllal1lmiud$ \V,~.re' ~taJlWrle.ct~t1 lli!to, H eLai ,~~UseKpre~~'g ei~er ~:Mlv~miIR·Un:ll2~Ir,ay') or mil(oiilttrol ooMt.e). Firf/lJ ~l!dfiilrgSl:! aJC~,V~tv wa,s no Hi'la:t!Zl'ldi 'to R'e:m1ki ~li!dr~r-as~a(ti'l!'it~ iilfldl t!1@!'I1 l1!!:lrrn1!l~z.edlte'~e' ,allIllll:a;gre ildivit.y e~ the (iel1!trO'l weporter. Va~LJe!i are mB1!f1 -:it: SID f-orlriplii!::at@ :Samp~es.

~clific:arulY linind~to MI(]l:·3.'lIii1R g liId1l'ilh1bil'S its ~J'gnsJ.ad~:f!. F(I~ g n paf1oE!is, ,C!~, !iepr:~ll!ta·

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.!U'lD n;cognilim'i,

rnliR'~~l1trill~ or hiCJIiIV-lIIiIl R,.JU l112~. Tlil:! IleWlLs of 185 rib01iillllirl1ll RNiA were (liliiuntV:a~Lles, a~e meii'ln ~SD 'rm' lunpl,kate !>arnrpl.es;,





or M~CB "vlllmuaits 3,'UlR ,,[I,U B) in IPril,nrn,,)' JDOim :mbsllll~~ti<m ili~~clmns;c~~Jl~ MI'C'f-\ hmm!!1Il 1,bD,,".Sk,~n~11J,_lroM~_.s U IFf) o:lh:, Ec~~pil; 'binding: si~c w thm ('If MlCB (NHCA !.Q llf.vg, .;m)., 11~~!'i !J~\I~m~!ll illdt.'l';~ ~~mSOOI r~;;d~c~~l~ CXrI~.jil51~~~.9f~~!!.~\'·m]!R,·U1J ~111,,'{l~~n~, a r~~ 1:2~n 11,Id!l~~~~t;:~h·i~.y ~Kli. ~b"iji!J!k~mtLm~!i;aJ1L"dam ~~~!~~~~n do~¥,f!I.~1i,:~~l~niar! 9~!~ M~CW·J' b~n of y ~f~.54,. indit'liilil~g UI~[!I1.iO:::i1i,':~lli:; wi'ih iiJ:I!.l 3'u..rm of Mlf:n, ::i1~lppo:rtin!:l pn;.~\'iO~il'i. UTRpml.c;:;n :!ilil1ggQ>~(l1l5~hal ;ld~ilio'Hl!!1f'ICI{I!'8 ,~ iH~'(:h~d in are ~tcl(;,lq,lIa!tc' lC~I~ng!!he I{]f dlb;;l or viml h1::m'l'!1~L:; ,k:i!.I,;l'!'ll~H;ntiG~ jh"ic~i{~'ia~ (lr,~ bl~'idill~~itc,tn h ~niR.!ULl t2 OIl M[cB ~"'[ln.'Ssion. TodlC'tL'Ii!uiilC wliClhcr Ii;{,Jnv·miR~UU ]2 .~~ W~,~I~"'.dthe Hff edl~ ~Xrn."'&iiig ,eiti~er d 1ii(;t1:t.I M 1['13 (~N:ptt.';S.:;ioti by .im.l ~ilL';~!~C i~s n i~'ig MIf.'U·J'U1lt ol'Mlen .1IJidinliY:ltell~ieifi'i wilh

~gldmioll nm~'i<l'tC'd ~y 1-ln.1:V U.ll'~~ I)mtein. ,(t.n \\'C 'i,","'<J.min!.>dthe c!1b;l gf l1!clfI~v·llniH~ Vl...~ I,;!! iu HFF ~~I~ ~~IIf,~<,;i!~Il ,~it~r;;r (iFr. 'i;iFP f~s~dlto tlUJ 3'UTR ofMICW,., 9r(if,P f~~d, ~r)n.hc .3'U1l1, nfIVI]Ci\' eQlll'i.isi,L'mi~wildl ihe ohserved M ICB-3 'UlR dmvn·.reg~ulation (Fig, 31\ I omd the qPCR ~$UI~~~ (r~g,.:.5.51, 'nhC~lL'¥.~ ~~I.uiJ11111:edm:;linr.i iml fifP r


AD I 69..wuld-I~C \'ijl'u:;: alld ,o.nly ~ii1, '~IMe cspilC~iil£!: t]~1i:GF,P ru~d 1:0 3'LIr,R ofrvUCB (Fi,!!.. ]C}. Nt} ,c:hmn.~c,-:; in~hc orp levels were o~eNcd ill.cclll:. cai"ilj'l~1B GFU?alene or GFP n.lscd IllCiIhe 3'U1"R or~inC'A Wi~. 3C <Im.!t]s:, S5), To tilrth~r deUlolIstml,clhe biCilo.sica! relecmwiiL~ I[~~ attu~l'Ldmi{1nongc~m =miR~UUi2 vaaee {If Ili;:U~l¥·!~iR~UU 12<lud l(~' e"1('IiI.!oe Ihe Vro IICc"~ c.",drn'O!:IU~~ li!illC!UUIJ1(lfhcm\'.!miR~ ~ibi,~il)' !IHI! [h~ d~I~!ijo!]~rd1.eUL~1'4Iwolr:::in~ UU l2 dll!ring n~tl1~l1t1c UCMV injlx'ljCinwi~h USL:; 3· day:>: "Llft~r in fOC~io!l ,~n~. ( I J). ~.r!COlltlW~, S::;) f.lir,,~\\~~k!"I.Yi~H~V~ADh'llt) ,,~u~d ~Iiu !l1~~lMV~ l'il~tc·B do\\' u..fi;~u,~~n i~l !]~~:1~4,~niCC of~]le t'l1~'Od.lllg.gC~~ "Lrre~l.~dMICB dn\'\IIl;I"fC~g\lbtilu~. iU:ll we COll!-llfll(;'tOO 1;\\'0 Ht!W mLll:llliLCd vil'!I!!le~ in dlC ADI~l}lluj!!lIlIJ11 (14). In (hi~ mU!t<l!I:llnJlc UU 14 3"UTR ,~ "Mllich \~'a$ rnedi"l:~-d by the 'Vl.~ 6 IDeMV TB40!i!imin ~lik ••ro~~!1m1 'vilfllJ.l,iin ',~'iak:h gem: Ii' dcl\:lI~dl,Hradi ~1~C'I\':ftHI';1 ]~lC' heI11\'-rniR~ !'lro~cjn) \~..." !\i!11~~I',ur C\l]lls ~nr!:.,"c'~c(] with [!jli:l I!] UL 112 ~im,," tll1t! p;:'1'.i(iC's O!l Ihi,:: ()O~.lpJcm!i,;!t![l!')1 UL'~ 14 i~ ,nult!l!.1X.-dm~t l1enr~~"'!lIli!lVULll2is~!~[:!)f.Cl A D I (}9~\'.ird-lYll1.:' ;:Inc! !:tlll!:a:l'1,1;dj vi I'll,,,,,, I~fi~,i;~r:.ut(!is ,ul~ab~rll" HCMV·AD~.6lJ l~ulI:!(lro:~OI)' ,lA )'.Th~ clf(;"JI:;:t of ~lo!m'-IlnjR';Ul.112(ljil !VIles· (lJ34o.:lIL I :1 4r·ml!BU ;:Inti ~1 1l1'l1!(HL;,j ULI ~.4 \l'ims ~I~~] i<;MfurK-d to iU:lfi.'(:1 pfiil1l!u~! 'fiilroM3SIS mbd 3'IJ.JT.R e1(p~-:;fiL'01l \/I'mi!; $pm::ir.e. ~e.[~usc lim" !e1f("1 il~ \.v1ilich h~m1\'';Il'~R.UL[I.J W~ d~tt~tc-dnHAO. ~n~R~UU 12-:rmnl, lhelli·MV ,],'I34{), ~rnln.l1 is I) r i\l~ u: elass ] rbowl:! [(JOc' dQwl:I.~gu~m,ooby I. :11, lirl.'~~OO uumbsr eell lilies. E;'1pr:c.~i:O:ri silnil\or In dillj;e~J s~r.lins and iI1~:CCI!S('I1dQ~he~i;:l:I: M~CB[ll'Qitcin was iilffi Q:b8CI"I~C'd lllli sc\lcrnl p:rU~ ~he virus durin!£!! inlK,'Cttcinl 06)] MI1lS, similar]), reills, .o:l1C Qfth'C~UYll:l mll:argcllS of I~Gl.ilV \' i\~Q ill. ~lm~ fi~roM3'!1S, :;1i~!dI lell .!Iines, berore or~~:tcr € rcduecd by both viruses (Fig . .:;B). ,t.li). .~n:l~tioJl of hllll1l.~n!iIImb:ilic3~ vein cndethein:lfccl,io.n ~j:iiQ. 'S3t We !ll~(J d~is~CI ur rellClil and o ":0. s.cgreg:mc~hc' 1l1iIRNA [XIS~tl'illl:iicripljionm.l C'x~]U\.".lSCd r..UCB ~'Vi1J1I1::13'UTR fM~CB,=3~LlTRJ I!cgl!l ~,U!jl[lll 1 M[CB 'I:loom lilc' 'Po.sl!t:l~msllal'i,Ollal Ikil cells U ~tJVEC,~1'Ih:u r:!1],clogcunoll."l), C1>:I)I'I.~S
mi'iR.!",!J,\, ,cl~~d!llioj1. W~ te~lCtii M[CB IlnRNA

th(: ADI fl9·w ild.:t,yP'~ or \'!;"w~ [lie AD 1,69· k\1~~ jn [tl(O cc.ill" C1<I)It.'.,>Shl1g tltcii' uni R-COltlrol d n~Ljtrlli!..'tI ",i~. Thclif'oleiUl CJ';;PIL'S,jl\.'tI by .M.I.CB· cmnp!l:et~ly dQWIll,...rc.~lIllaJi,."tI or hCJ1tV"llliJl-UL J, I :2: by usin~ qPCIt, No s.1jp~ir~ .; UT.R was 11.1'11110::;1. icantclmnl:.'C i~~ M I:CB mRN A level W'm$ ObSlCr;.,'OO on day 3: :I:I:krinfec~km iDl the AD1I&)-,vi~dcl)'I~ inr~c!!l'deells bw nOll,." t~~e ,ocllsinlbcb.'d wi'nh(he {,~iS. 2C}. sllS!eSl,i"~~hal 111C' C]owal-regllimiml of AD ~(j1l<=:~lUJI~3tcd ~'JrIlllSl,(Fig. 3 A),. n~jl. COII.'j.i'i~= ij;. M [ B ,\\fru: due 1.0 tr.!ll)1kltlllJl1 inhib11~{ln.

I;h,y 3, on Iy \v&at"i~llie t:~lt"

c~;pn.~i~'i\""~!Jl o~n'I:.".d. v.,'~~ inllt'IC'ti."iI 'v ill,'i






A!~ti MB~a



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HFFltl!W MIC~3'urA 1W~6'iI'M




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IDaf$ a~ii;:r'illf!~I:on [0

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IFilg. .3:., Ihcrnlv-lII1I:iFH.H112-rA@di.al'e'ci! dO~~I1I·.regul<!lioiii or MICB diurinl~1 illuthe;nlk vira~. 'imlection. (A anti Time (Olll~e e:x.~r,e:s~iicm MliC!!i (A) andl ,()f MIHC (1a1S;~,~, (EI) @1iI Hlf~ (ells e1l;p,re~:ll'1g ~ith:e~ MII{Bi-3'Ilm'R (If (Illty MI:CB. Ce~~ M'~ ~11f,e(t.e~'wi~h ,efrlJh~rth! AD169:"~nd~lyl!l! Virlu~ 'iJ\DL6'9-wm 'Clrlh!t A,P~'1.l'·miIR·UI~lJ!:!! 1!l11[l~ i'lt~,r~$ tAiD:I!~,9\.m~'~~UtU:2·muO. The 'el<pr~$SjJ):n ~~els alf MICE! (#It) an<l MH!C (Ila:ss III (B} ~'lif!i1l!g~s~t:d by 1~#I(s,:S:!a~lni!il9(redl fuiisto!iram~. 'Tin!! ,gRiy niswgrarn s rll!preSM~ stlirllilfl~ o:r tm!! «lrIreS:!lonrlinl§j1 l!flt!i'!re(~,d (if,lUS, IBackg!'<llilr1d le~,~'ls (Ma(k h1istllglFam) a~.eo th~ ~oo~(lafY Huo,!'@sOO'il'l i${lt~~GCY.tlIflI!~ (IRllra~ooJiliiL!Qa~,@d Ab~" {() HIIFF c@IIs,@iXf.lf!!mIilQ 'G'IFI? I~r,GH' Imt1!ciI. to ili,e:3'UTR off MiCa: ~\len!!inlrE!d!1!d ~vitlil either A.Dl69'-w:1ld·type

wilt! AD169'.m~!ril.·Ui~]l2·rI1lUihim~s:, ami the' leveh (If GFP !!XP'['e;5slQIiI, \\lere meilulIredCilofllgl lillie[O'Ufse oiin~ecli®lil. lfII1e'5ym'l:mls I\e;paesem '1111e ~eK~J1ta.ge OIf GIA~ ((!m~a~edoo d'li€' !i(l~!i€~Ij)~lldil'lg u i1ili!~'e~~oo cefu;, i'(I~ 00 ((). Ol'l'~ {A} ri~:p~e~~t!~i!' '~PIi!ri'rn~l1t liSshowfl glut t'hn::e p~nflJIDlTled.~D) IHUIV~Q; were ijJl'fe~:edwith ci~h~r lB40 U,U,$4'i[i1il~i~ntvirrlJt5 (U!4{HJU14IP Vf!1,I!:J or wilh the TEl40hciIilIif"mliR=Ul"Jll'1I!2 ffi'iluUilnt ;;iruiS. (TB4,l}m~:R~U~U2 mUi~,ai"!d! the !IlXpr'l'cS5~0Il1 MitiS 'lliii!S m,@\uursd(r!!i!i! 'o~ ~islo!:!raDil's). 'The- ~9ra.y~isto~ra:m!> r~,p~e~f1i~lhlil' staliFl~[ilgl,tdlihrl' 'oor~5fJ'OJi!.d!~,g lIo!rdlEl«~d (~nS, IBa~k'9r'@!.lfld k!w~ (Il~k~ IhisWgr~mJr .!:IF'@ th!! :s~oo,ndawy 11F~1I'C·((I'nju:gil!lt'@dl :5how,fl ~s(Jne A~!i. r'e:~rl1!~nJt:ative 'expE!rirmmit cud. 'o~ mo perf'Orlm!1!d.



M~,:CB 'li.'ilh Ihe IB~O·U:UI4P"nm~f!:,.",!ltcd in [!lmQ."~ CQIH]llcM ,dQ\VlNCf,_!p~niio-n of M !(:'B q;';~)g1,.>K.",ir.!I~ .. ~rJ d..,;y:s, whl;:lI\.'~" MK'!:J C'4rm;.!jl~ ,,,iQI~ w;1!!l $!!i~l evidi",]l1 iJ~ c;l~U1iifll~.;(1 with~lil,) TH40-UiJ~R-'lJU~,2-I:i~Uli'!.,i~s (Fig. 3Dtllt~ k~vel oJ MHC c.ia,.",,,1 was nn:~ redl!C~ dW!rillg viml inll"1;;ti~!~.bI;;'t'1~i!.I~(;;!he TB.JO \ri~\I.~ W", U!it;-d hl!t;'l I:h~ vil!'li~ US,2-US6 ~ot(~ t~~u~ l~tdf.!mI!t:.MHe d~'I&"

~.dO\¥,r'I,;.regtil:m:lun Thi'l'i~~l,ih :~Lig_i!C"SI~ ~I:la:lt d uri i~~ ,el ii~~!('Im~ i~'[I1in1r~ti(m., ,~ndoge:noilisM]["B v i~,clo.\'U!.n:~1iI9!cd lJy hcn1ii\l·m:iIR~U.L·~2. 1 We IIC:.<t ~rli'omlodkimnc;a~ays ihl paral lei ~


m~H."IJ11, in!~ \li,Q!:! .~il~,e<! more etl~I;~i;!nll~¥ Illlilfl v thQ:5~in~Cloowi~hl ~c AD l.f'l9·\;\'!!d·ljr~Vn!'i (Fi:r,;. ~, A ;md m. Shn!!~Jr!Y. HUVE-Cl' i~~ r,~i,I;;d wii1~ ih~ IFI3'J.O.;.~!.li.R-U'='~ 12"1!f!~1 "'~Il!II~W1;~ killed I~lot~ d)Ic:~'I]lily tI]DI~ thtll;;~il1Jf¢i)icd wiih ~h"TB40~Ul U4'·Tnl!l virus tFi,g. j\,dt!il!On of;'iI'lti':NKG2.D mAb ~ni>l,'ili~hcdDhmtd~trer:t'~ (rig. 4. B ;Uid C'), The 1ki1~i]'ig 1L'l'llCj~ tl:ic af bl1foolru .1iUVlEt':s \~.relk~\\i. Cvljfi r~l. higiier c 1T~~h~r:Utt'gj,)tfflliO$l. j]ro1bfublyooea~L"~ af tlilc


The di:500~'I,;'1)' l'vl m~ ~nil~NAshas raised Ihe e iml"1gwiUQg pfu~iMHW QI' their i!liV9~ ventent in im~mJ!!~ 1,}'\'Ol!~i(l'] {'It The HPll d!~c~ (,;\'i~I'el~~ fg.;r ;Ill ~~liR,NA-n,;-!alediltnn~1Il0CVlI:ii(m atu;dHlI~il;m \\I,I~ (ii~(](J~·oL,~. ill lhe SVo!!l,",'iru:s,.wb~ ~I. vnrul, ~niRNAn."I~!:5' \,iF.d g,~n1.1. ~!..!~.Iing ln en. C\i;;J~:O\!l .~J.n O!,!l!F 1t;-1i~b ,dClll['u~nnilC 'l~tI!ltC:!.


ml'iiRNA-om'lt,.'\!Icv~,"~irm ~W1utL;gJ l.iM:dil}l 1'ICMV. in, \~'I'Iii~eh ~ ~ Vom~ ~iiJjRNA din;:ct]~ dO:W!ij . eg.tilblte,~ •r

W,U~llhc s~arllirrlg,Il'IL"l1.cIWL--d FilJ.. 3, A. B. and D, in On d.. y 3 or Ihe inR.'CtKl'n. the O~USClIll)l:essing \
M[ B-3iUTR




bigh levels ofMI·le elass I. nmlecllil\."-~. wllichil1h~bh N K ~Hotoxicn.y., Tbus, dtlla:ing alllth.clftic Vil:<lll:ililfcclKm. hCll~\·m iit-Ull .~ dO\\\'It-flC"tllJa!l~S 2 M leB. 1)C'rI~H;bi I1S~I:l,binding: ~Idlh N!.K:G2D and cOI1;;wqucnlly aidfng in NK .m,u::k evastolt


s ~

3€jL ~l)

AD169·nn!FI~u!L'111 ~1iIlI1iJI

1t0.1.1. ijmnUilte dcf:Cnt~e gettle'. nCMViskllOWll to 011 seveRtl rliilCLi,on~J~y rod~jttl[-jjlill. irmmmoC\!2isii vc pmteill:>thrn,t can ci:l(l;pemt~vcJiy U.l~1 Ille S<lIlf1C I)~'OCC$:::: <OJ' even Ille same il1ml!llll'lClPm<lcii'li of ~lIc ho~t ,~6~, Our 1lC5la:llSe....I~mldlhis,\'icw. dCnUjll,..:lrnttll£, a COO~r.JIlt~\ 'C mcdla1il ism l~I\\,CCI1 a \']r;;llmiR!'\,IA Ihl;!J1J\'--In~R~ULl 1,2) <!no ;lvi ..11, ,~,!'O,tC!1] II HCM v UL HlJ It J 1}[,. l1ttlth '1<I~~t;l~~ng !h(l h~~ M[cB pl\~in.. Dt~~"'¥I';.!1.~gul~titl~l hO~l (If gell~~ 'by ]ntl..'f;llC~tOn~ orlhllir mRNi\:s \""~lh '\'irnl, rniRN}\i:lm,~ been ,~!:so ~!i!l'lil~ortcd r(l~elll '[by ~:hl.dU~lg;"in h!.qK:'.\1 ..-:im ll!cx \'irus I WJ). The ~WIllIL~~g.~ of :n, viml l.Il.iRNA-tl'.-..~!i"ld. l~\!,~l!&klll mtrl(lCiiru~i»nil arc muhiph'i" Firs .., I.hc$Ii1· Il'IOlce!.!ik'$




me snmU,

1l0,l1il'illIliiUntO'g;cllie. 1IIl.d~K"CmC

(41. Sec-


ond, lrom,,![n


(l~--.----.---..-----. 12:11 '8::1 IE.T' 00::1 IDay3 HIFIFIiMII,C8:·3'UiTIR

E:tT !i(tl

Q~c----~----~----~----~ 6:1

pers:pt..~i .. ~itsholaM 'be' Sillt;pk"l' W d~\'e:IQ'P 9 ~"gul~wory tlmiscll~n~o~ eeule rnllH.~r hanl a regMI.~ory pl'o~in" Lm'I],),.lit'! ( • ooluhinalion of l~rolc~n'-'medi~lI,ed n1~RNAand l1)!,1di:ttcd'~t~rn'!."Cri!JtiflnJlrc.gl!lmklllpl:o"ll!dc;;; i)
!i§ll!~fcvm,'i.j(ln ~"I"lR'~y, wh~cbi!\'l1!lro'l1 r-csif1lmn~p !the i~os~ d;crcnscnuc:cll!ru1 lsns ll!!:Cm.lsCI,\'I.'O in'll-

i 'ti !U .&r


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