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PLANT SGIIENa:

lid 3a Piro~e,in II!;a Mobille ~!ooe.ri F1'!] 5i'gnat in Rij(,e S" Tl'Jrflaf.d. S. M,I.I(suo • .H" t WO/tg, .5" Yako;, If, .~,~iml1m{Jul un u ~rrh:!l1(e,:1iti17:53 'F"!" rotein IMIcr~ement (onmbu~e'S ~CI to ngd)ista~l(.e S~gillaiilil,g flnl Floral P :llIIdu(t~oi'lQ~ Ambi'liopsis t. (orbesiet ,eiat.. Too pmb~im P'fadlJ(~s ohoo gllRl!S. Hd3ai n ric.!! an.d: FYi n Arobfdapsis <lIFe! tbi! clmi~~ ll9rig~rn .si;llrii!15 ll'llill mll"~ lrern IMllo !iMl:lt ~oioou[@ ILmo,-ernn g. 10, 1126JSr::tl1!flce, 1 B.417:52 MOLECULAR BIOlOGY ID!:~ lQlI;lIi!1~i1WUfll!~!l l!!~~g ~ ,j;I~ll:~ ACWI~~r~ Imp li~~l'e MIU in V'r.a rj~P©SQ:rl

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G~rJ~~ IR~l!!uiredl,~or Mit~t~( Spindle .A.:s:i@mbl.yi!fl Dta~!?phUa' '52' (@ll!j; G" GorJrfm(l ~L<lL A 1\'l'Id(!'-genOJne '5(~001 fd!en.tifr.e-~,lihe2M geBe'So iIWtll,;,oo if! ~s~em.b~in9 th'l: mlleti( ~pindle in Hle$ and ho\!!!tl\ty rntgh~ ,~~~rib"tlil.Q'Ci!Il!Cer {lnd QIl)e~·(!bnQrl:lii"li~lif}s.,
PLANT SCI IENCE

417
GHEMlSTRY

An hCDI:lltio:rr,alriily (,olfl~ened ,Mi~(ha r1lsmDe'l]m~tlI1l9 421 SHIl: Mov~mffi l [)efltm~':S ill Sln.9:~lLai~~r (lr IEndllljl~,rmB·

Fadile :5plinll19 ,(If ~~'dro!,len al'ld Annntlonia !by .439' Nl:lcleophl~k ,i'!,,;tiv,a.lt~:m a :5fn,g:leCarb~iiI Cent~rr al 6. D. Frey @It. 31. SI::lbi!!< cll!(liran-nidil t1lJlOOoo II1!lI'IQCU~~ Gm a(tivo1t~ IijId I\DgmI a nd:'s!llil
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In P·lants. N, C~/di3l"

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I'ndi'lllidl.lilis \Nilll iJ.1lit'i5m,il~~ rrnmeW:.d'l' '1.0 ~'hIow~ari~tion'~ in thlll!lJmber 01 (oPI~~ o~'[erta~n §,'Momk mgiol'l~ IhiIIlI are tM~~r IJ naffeclJ2d rd~Ii'o'e'S,
BlOC HE MISTRY' Ji:,fllllIIfllrn-:A$si)Wd (l'}'~taU(]grilphy R€!leab Elild"On449 Pen:u;M~11'I~~f1llfld~leS il1l~ N(lnh~liI1~lrofl IEn,~,:~IlII~ G. Kutunu el jill I nWI1'l!ld~~I~~1ill a ~,;lnrfir'l ~~t.:I~~(\~ I:Ir ~upil ilQddi! tedlJ~~~ ~dClopl, ei'lolj,ofl i!.(iInnlJilr~UOf'l5th<llp~more ~~·O bond! <!L~'f<l9e..
:>~

P~YSICS Ne'llaHv,e' Refr<!d~on at Visilbl!i2 Freq,uE!lIdes 4,30 H.J. tese«, j, A DiM.re. H.A A~froler A thin \\'/ill\reguid!;: oompg~cd 'of thi n li!~~r~ of gold, siliil'cn IIrtrid~" amr. .nt'l'er prl1ld'u~e~a nega~tljl~ i~de',i; oj. r'efriK~i~n at blue end '9 rl!elil
wilVl!lerig~rus.

Poer:sjllerl1vO'

Po :; 19

MATERLtlLS SelENe E Crte,p-iResislOlnt,.Al:ll0l "fomI'LnJ~AlUliteflm(

4,33

r. Ya,i7Nl'mQ~'a'et a~,
18)'elirnlina~ill~~'t.amllirn amd 'fa l1Iadfum IftJITl ,~ ~t~~ illD:.', teMugh a~~mpnurli (all be ailded 10 el1h.I!;~,e re51~ilnl.1~ 'to cxid~~iic!l at h1igli Ie Il"Ipl!f~tur~~M1i Ie rtt~ii'iingi!~ ~~~! h.
MATEfI'IAlS
selENe E

Starill'lles.s Steelis,

BIOCHE.MISTIAY Cr )!Stall StnJiI;~Ulres (jl[ FeU m{JIlt~g~naS!e' SIiIjil!eroxo, 453 AU;'~i!Jo~,o. and' BOUlild 1P'r'O£iu,[:t IiH.erm~dia~i1'S ~ E. G" l(aifOleLl"(1 amJ). fllip~r:r:mw E~{h S,UOO1!UQ! ,~ ~li'iUft~ri,. rlilig .<i~,~iil!!i ~l!l;t)!9lem<li~e hOlf'l:iou .. d;lf~en t ill~rlllet:li~I~, ~~I!lm'lin 110Wi! \!r~1li$I'eI5~1!CI~S 'rr'OlI'!I g ~ubSI~l~ 'to ~Y9~r'! 1.0 i~1~1Jl!~c~:;W.llge,
:-~ P.:!"Sili'£'C~1V!I!~. j 7'J

Sylfltliesis of mtil~·h' com p~es:slile SIJP~rhafdl t IRhe,mlJ iItI mbo [i~e at Allflbii~FiItPre5:~ln~~' fI.·r. [flung
Il~ium

4,36

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,MEi'rillllry'

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J,~.ff. Han

Form'8li:cm

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.. IJ_... ~¥(,YI\IOh~ ~~~.l'Hic6.>:JIiJ II;Il """",,~!oo~!J. ~~!Itt~ l!:IIlJ'ig1>I Q KQ~ I!t !"'·~-Iy.I,;!O<.'I!rton !i1"I'II,!I'!!I:_ .1~ ...ltooiiilo~mN(E1 'lOlktti... ·,_ld .. it",.,,J,L'_ik irld"""''',_liImhip ""'~~511 ".,";"I.ltl' .~!~ _~'~~li<!1I.~kk1'Ii!'iI",,,, .. ..,,~,t~,~ "'_." !~jill r.. dg!'~OOl" ... C.riIDor""~ .. '" 0","1'01 itl N,G '51: IEIIICE,. :~~I'\VI He ~OI:I~:fY "'~ I~~ <lflorOl'lrlrh:>'illr ..... l:dolloaJ!'·ln. F1<o1<lo", ......m. Clm<!!im_","h~~l .oiI!!l!O,*.;~(i:i! Itlil'lo!!lem'.~~.I!I.;'"~!_~ W:;JoI~\ U.!J'Wi,.l" oIl.~diI oni1,., .. odot-,.-.d ~<ilpoo:WlI_. ~.. 1IIiIlIi""Sfril.,...d'.od-.",_ II'£i_1!O>! ~1m.1'IoIhIoijion,Dr ~!1.!l SI"":":,,,,,.~~I"" I~~ ~(III!11I>I""',,!UMl ~ m...~~i:I ~ ...I ... Pl"1o;i~ MI~''''' ""1__ !I!I"''- ..j{iIfIIII~ .. w..!liI ,.~ ... ,IIIl!;l .. IfiI,'l!o iliI <iM ~ (ji1h!("~~'" iIw;!~fI\",* ~,!,III~O!<u-wo "'" (fi; "1i>1.ot5il_d'Ol" ~~ ~ c.r- iWJ'Tr_~I~,!,!,"""~~ p"."f'od ~ .• ~"'(IIIf*"~ II

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S··:C·.·IEI~'CE VOl316 11'1

20APRIL.2007

An old friend ~ with some new tricks.

For alief 20 wars,tile
fhe MJni-PROTEAlIfl
p:rotelrt eJeGtrophore&s, Now

Bb- N1miMini-PFCYTfl1II.f' system has set me standard 'or exre!!enoe in the bEfst m{nt f]tactrophofmls system is even betlm - Introdi:Jdng

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LJI5, 011 U,e Web, at discovar,biio-roo.com Cali loil tree at 1-800-4 810Mi) {1-BODc 424 -6723); owislrnlne US. {;O/lItm;;~)'lour lecalsales office.

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SCIENCENOW'
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Bri P1gning S'e'x 18il<C:k ,l!i;l!erm~I~l1s,of~ears 01'CeM:iiKY. oll.lfin Li!lin en iI ~ i~ES se\llU1lIelMlut~n. r No Twisting Out ol Newton's ~,aw
N'e.v expe,riml'lll ccm:firms· f!llIe-a((@INilliom

{onoorliol1

'for ti niesl shoves., (himps, Are (Jiarnp5, or G6ntlic Cl1anges; N.llIllr.Jl selection ~Clmd mnf~ hea~yon (himp<lll'lzOC! gcnoli1ile !h.m h U mil n genorDll_

GiOWil'i£l po4Ien'lube.

SPECIAL

SECTION

Germ (ells,
SO EINCE'S ,$'TKE
~M'o\I.5tke.[)rg 5IGNI\L n;:A~J5DUCTION K OWlEDGE ENVIRONME

SCII ENCE CAIREIEIRS

W\\,",.5d~n(e{aJeefli.org CAREER RE)OURCB FOR~C1ENllSr5

IEDITORIIA.lGUIDE: ifocur.ls,sue-From Eggi ~o EggGeU Siglnal'ing in Germ (ells E. M. IkJft.>j (lila N. fl. 60ugh SlQlnals from tlie S;UIIOU 1'1 ding I.iS5I.IfS mn1!r()1.sUi\'iwal mlid
mlglfi)lloIHl1 gefm cells,

MIISCINET; Profill -Wilhegl;l CQ,IQIl A. S«sso A biocM.emi5l~! RPI Yli!!lcomes mioorli~t5 inlo hf<s OOpai'iliMll .!Indl rtiaarch lab. GILO BA!.: Mindllll'alttJ~fs-CuIUl re 5ho(~ I. S.~eII'fn£
IKriOf!~"g INk-ill"
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ahead

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mecha nisms 1'0 guide mlgrillion of primordial and elimi I'!<ltth ose !hal rilil tl;! mi-gr<ll • P'£RS,PECTIIVE: P'clhm-Pistil in lhe Pi'Stiil?
B. McClure

gierm

',eUs

Signalling lin, S@U-IIl:(iomp~lib! P'oj:lgly-Dc s II Allow MOf@Elfie! 'Ill R@!iouw! AJloci!!liofl

De:ltrl.l,ction 01 if'l~mpilitlble pollen hliw5 ~U~ IlrNlI~ IllOIIe ru'5

(areer skills need to be developed before emplo1~rs can reroglnize and rely on them,

rHQlJ

av,},l1,;)~Ir;:rlOr S\lpp-ortin.g]tM gWMh QI 'IQmJN.tfbh~ o~~s.

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CJENCE PODCASI_ .................

liulle r n [10 lihe .20 AiJril Science IPQd(a~t lQ heer ..blluL N IIIH' s budgd cnliltll, IhefnRilellQe 0'1 tem,peratl!lre 01'1 se dfl.etmin1lfDJiI illl replit!!1!.

dllmestkallOIli of marine .5Pecie~ Olllld more.

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sciencemag.org

SCIENCE

VOL 316

20 APRIL 2007

• __

..... .I! -

N!HI! !;NHI~,L!!I!"D!
Bil'DL..ABJiS

T4 DN'A Ligase from New ,tl1lgllland lB~o~abs

WHEN YOlil NEED UGAS IE, TllJi1NTO THE INDH SJiRY SiAN [lMD
~ ew eqg1aTld 8i !Jlabs IS deQ!~a~e(Jtill IIlwvidJp)!] ouf ClJsllOmern wil'h guaranteed ern'(l00 pafrOrmiinw. Our rB~rnnbin an~.T" O~A Ll!la~!1!is thl;l rrnDit ~~Bn~wflly used Itgil~B rgr dgn lug exparime.ni.:!;, J, ls available at e~De~tr(JIa~' vall.JE!. and an e:ven greaote1ita111f! iNl1sn pllImha:soo rn hilFlJE!qoonlilie5, 'Ier hig11 I.hIlDll{lhpui t~htmlogiet. Filr cooeslye, blunt, simple nr c[JJllfllex fe<l~tJllIns, make 14 []INA Liga:;;El fFtlf1l

NU yom nrst
II

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~~~~!fI

EDfVIED BY Sli'IIUA. H U RTL:E'i t\N D !PHI'lSZURDMI

-ce That's F,ar Eno,ugh

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,1

wiith mOore S(R i1Ivilln~able01'11 demand, 5HR and its devel.opHll'ei'liItil.t si9lla'l {Oil nmlat miIJIVil aut iIny III I1her t haa tlh~tflrS"t
(J:II Lay~r+

Mantle M,elt Mixing

The het eftlg:e Il'l!ity o~the con\!\E!dil1g 11I1ilnt'lE is .a result Dr re,r;ydUr'i9 Gl( (](e(illli( .1Indsome {!)nHliIe IiIla I. UU5t wflhin it. So:b,oi!!v,!tI· "I. {Il. 412. pl!!bti sh ~ (In li fll? 29' March; s(!!?Ihe Poersp!!di~ b~' Ilflerzibllf!:l!) haye dewlopeda melhoCil fll)r [.ep. gla'ting out Ul~ re(y<[~d 'crust (ompljnent.~ Til biBalls by cilmbin'irlg m!'!<I~uremerll:Sor a VlIr1@t}' Ilr 1!~l!mer!ts., irldlu di Ilgi Ni. Ca, (0. l!It!], Cr. and
At TBlE!y a tlliil':i~eda targe ~.amp~eor elil{im~ 'P,he!Il1lt:l}'sl~

G1ii.m'~it (11. {po 442.Iil'U lilishll!d O'f'lli rIt! 2'2 Ma fth: d See tine P'~fsPetl]\i'e ~ BiiI£lflfli<a~ ~hm\i ~hatmli~l.o~
Ih~ varFatRllfl Q!5I:;UlI'$ill !l(l[ll(uL~r ~OIi9iti!Ide~ i iQlld
~1!1! ,otbiHd 1111i1! oon ~:nt~ladLl~. m wl'lt!~ vapor b als~ ooil~g l!'ljl!~l(!d. Theal.dhers propose lilal !WIn Cll[l1;l&lionl ,c~L1s re set up 01'1 ei'lhBr sHl.eDr !hE! a 1l~£lna dis-k tha:! hmllleJl iom.llrefererl~oi!ll'Iin cme dir&li.®n, wllt!:.11 ililu~a~.s'lhe ~t~Ltr{ll1 derl5i1yand radio em~~ofl all (me llJl'Il].ilJUde. Il~ar

r~\pI!I~s 1rlt!""di~n s to phD~lJ;il'1dU!: l\!d D(I;r'itll-tl;lIlIIe {oppe!r ciha~g~It9Iil1s.(er ..

B,endi n gl V~Slble Ught Ba[lkwanj

.AI liridal 'l11l1M!rlaRs"@r melama'truia ls, wr~htu 1'1a Ille ell!(lrl'l an<d ma{ll!1el1~rE!~ons<es can gi~e ri~e to a liH2gab\l'e imj'E!X [If r·~~radill;n,i Ii! !!fll ieh i1[,e(lromilgn@~k radiiltiolil is belli! lnths op~mitE!! d~w::!km (.f)I!lm!h>!lt ~~pl:!d.ed 1~'Iur~l.mil~~fi~~r als. Other eff'EKt) su~h al) pede1J ~ePlsi!1gand (.Ioaking haW' also bee n dt"mon!il["t~d st longe,r wa·vel'f!ngUI5. L@ze<: et (IS. I(p• .1130, plul1li'lhed onlill ~ 2:2 .I\hlrch)' Il(I\~1d@fIIl(l.m5tr,)t~ 11!l'g.'Jti~e re:frra:c.i'l1'e! 11dec,," i1'1 nlle vtsillle regime, prOl1liding 11 the potelml.i~l of practi[.al d~vi (,IB, such a"$ su pe r-

col!@(Le,d horn b\lsaUs hom

111Jd-D(!~aQ

ntlge~.oceatl islamils, and lar.ge iglneou,s r;m!!vineas, 1011 Illic:ro:jlr©be iJn!;}IY'5~5 'Were ~~rJied oUl ~or17,000 9 rai n!i rrom :2 32 ~s(il!llpl~IEly .. .al1i3~\lzil'lg ~i1.BiHolnp~iti(m~ iO~fIlly, 'o!iHii31ltons \'J~re seen ,'or ~h~d il~~H~·ill~.llJi3J5al[ that csn l~pe5 bl:?.amib:ed to crusta ~ mb illlg .. RJ2c:yd'E!d ma terial. is· dd«t.~dlne ll'll~§t a It liI1e1till'!Jle rI11ironm ~nl~, .8ml its comr i bu ~ioill III th eom el.l. can he re~aled
10 poren

Cuprate Contortion
Advam:es illl both x-ra ~ .md electnm-di Ullt!lcllo n
tedmli'log ~'lTia~eol'rE!'rreclani rill:reaS:!r'iglly deta H~d vie\',! of tlU!! sud clel!l s.~n.K[ural (hilng'~s. that solid s C.{In LJflldergo upon Qptic.{l[ @:l:citallion. Using eleclmn ICIflk<iCliorl wiUu picQs.emnd timtf rescl'uticDrI,

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p~d;:lred light-il'lauc.e d e1;p1amiooin ,ill cu p~a.te (~fami((a ~ul}erC~fidU(Wi' beilow 32 ~!!'lvi!1lt SpecUi'" ~UY.·" pulse I;Ir a:oa-Illii1H10~ler w(I!;'i1len9lh p,l1iJtOI1lS abo';'e ,8 {.e~ta I'l ~liril1!~h· ~
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pfi)~,lerrnl:iu~. 100 i!JtJlen5iv~i\D~WiiiJesP~~iI~ are i.lSll. :5tlme sle~ls hi'!'o'ilHlifftt:ierlt Uei!"stf>e119~h,. but ~l'ieir prDI~tH""E!diI rom ul;lInil O);!ili:l~ layt'!t5 do net have Sill ~ridefit oxidati",,~ resis lance at high teliljl E!t a~Lires. Alu mill lim ,o.:dli:le jlf.[jiJijJtl E!Sb~tle r Cl..:(id<lUvE! res15l<'1l'ltlii!. boLllal high AI (;Qn'tE!I~t, ~le1!11 rllml~ (llWea~er rerri~e lP'has@'" V:i31m~l11!oto ,et. ~t. (p.1B3) SllOW ~lnal by ~.umina til'l9 1.i'lClnil.HI!l.a tld (M!itJ.Lii1'd tin pt!9~ 317.

o~dirllllllni'l~imj

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s~lid ~1it!nleC'I;I!l1~r.. ( 1:0 IJlia 10 -fllll:l sllllM'fr HIl1~ r scale ..Th~ ~\(pafil~tOIil ~ale~ lii'lMr~~ with Ilhol.ot1 Flue 1'1«', whkh isoonsislent with iI flrelimi flil f"II nnotlel [d Iii E! trral'l~ient £IeDIm~'lry that a ItribulE$
SC.I jNC~ VOl31,6 E

www.scif2.n(~ rn~g_org

20 APRIL 2007

SW'Il losing time amdl samp:les. IExoSAP~IT" oners (Iulck. one s,tlep P'GR cl:ean~IJP with 100% mcovery !I)'f both shmt and longl PCA, products. Ex,oSAP-n
'. UHlizes IExonuclease I to degrade leftov,er prtmers & Shrimp Alkaline Pl1Gsphalase to degrade unused dNTP.s II' One easy step-just ack) ExoSAP-IT. InClIoote, & heat lnactivate ,_ Directly use trealed sampl'e for SEIqLJencing, SNP analysis, etc.

BeJ1e'fID~s,
1k11realJld H an;:! E1lI1SAP·IT 1Jeawd I'~) PCR prDliIl£1iS were analy2.e(I bJl' gel illectropllOresis_ A vari~fy 01 PCR.lloducls or tMfarenlll!flglbs may be tre,lIad Wllh E:iroS-AP-rr. With 00 sample rCS'.O_
'Il Rapio peR clean-up--15 mill'llto treat. 15 min to inaclivate ,_ No sample Ioss-PCR producls are treated &: then used directly in tile ne;(t applllcation '. No tedious columns I lJolvi}(r -more rree tune ", Solution-based clean-up'-compIBre~' scalealJle (or manual or automated use

Tills Week in Sci'eru:;e

'll'al'la~ium lrom the' ">lee:l at~oy. and !!ising !lilly

nioblmn to 'orln1irhe carbide preLilliLa~e.~, a lowel 'rra(' '~~onor A:l ran be U500 and mIl lorm a prOiI!!t:live' sl!:ale, TIlU5, a ilIig!ll·t!!JIlIlll!!rOl lure, ~ower·I:.osL nee!)· rf~si~ti3 ~t~! .<Illoy !!:<l11I! be ror!ill~d. nl

Ilightweight Cleaver
ThF! r«ld'I.ity WII!h wilkin lIIIosl tri!lnsilf (In mdillHlIlli d~1Ii'if~ tiI~ 1ii1l1s, b~~n ~~Irib I,II@dto <II pri mill'll !rlon afion ol electron 5,from IWdr~;e n to m e,ta~,LOuplM lfi'i tI~a secenda ry donation in tlu~ oPlID",ile d ~~(Hol'I., Fr~]j' et Gl'. (I),. 1'139)5hI)W that. iI si mihn IJfOCe5:S prornol~s H;zaddili on tQ a 11eleC:1on-rich r <llkyl("lJlIrn(lo~ G3rbeille., like <l tra l'l5,itiol1 111e,tll~. the stal:l~e sin'll tel GlrVellle has beth an ele,(tronlpailr .md vaJcan l 0 rb i tCl!. b,1,J l ((lkul(ltV()1Il5 ~'Ugg est Ul'i!1[ the priruC!lry p recess is n ud~ophnll(I?ll.'(trQll dlma ~io n rrom cerbe ilfto h~rlrogej1. fhn~l'IudeCiphil1( nu(ilafl "5rn a lso a ltCWI~ lhe c:arbe'DlNIHleaVf the rotHil bond "I am nwniil, 11 mu!;h rar'errre<lc;li!O'H at ~le\tn)ph H[, metal U'fIIliliiJ~,

Color-Coded MUotk Spind le Assern b!ly

nl~ abl ~ily IosGr~ef'l lh~ whole !Jt'lflOl1iH:!w1~hi DroS(}p/JUtJ ~el.~5 lUl 1'1 w RNA. ill~~t fer'~ nee tf(hnl)lo9Y lilfOvir:l~5ili1~ IlppCHlUiIllly ~o el,am iilli;: ilSpe<Jts or inlra~eUlJ!br pJo(5£es ~111 unp~c:edelllied de~:aLLGeshitma d'(il. (p. '1.1 7. ,pulblis,h~d onll Ill? S April) p!'ll':>'lmt <II 'i\.'ha!e-ger!o~ 5t r~el'! thaI 1J:'l~ Uillp li~ali()Da I a'na'i~s1:'lto aid Llli:! P f1~ i'li:i~ypiI: SlfPf i'ling 0'1 II i,g Il' Iimmgh Iilll!t 'liIk'ros:co~ im~!lIes ~CIi ~QClkfor prtlte'i m;involwd in mV~lJtk spi nd le' fUl1u io I'll, Tn!! m ililtk spi;mH ft repu~l1m!> lhii! 1illi:i5to:m~p l~x illllr.Ke I.~u r51.ruc.t ure to bela 'Scre€ll F,lt! I(J,dalto_ a Ildl the cnJ~ollJ!"'too scrr.el"! i!'lg t!!(h !'ID[D9Y and dF,llai~@dill1.@notypi!:. iJf]Jatys~s IHtI\I'~d~ huigliU. ~i'ito how spi i"KIl~ rni~rotlJ bulle~ dire g;~fiE! ra[eli a~ld InOLlic~nlrCil~fjme~ are pM~h(l~I~ ..

lnsiqht 'into Autism's Heritability

JlltllWllfj h lwi n ~Uidie5.llaVe
5.nIlJI\lf'i tllala

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slug'gE!1.Ulhat MritabL~ risk ra!:torsfor ,UJlis mare rI:OlI'Il:H~.RaUl er tln~n looking al pa~t-~ms .or iIll'leri· taliO~ OIl~ focus o~ trsd ilio.nal genet~( aipp ma(he<s}, :S~'liIat ~t at. '(p. 4~,S, pu IJli~l E!dlJll~il1le Ei Ma r\!:ln) Ila~e 'looked for Uite OHurreFice off sponla naous 'YilriilHol1 in HIllY Rum bE!r by m~uoilrrBlY mrnparillivE:> 911nOlillk hybr1di :roU()I'I, ill, ~'jI [![lUll d ~ lLO-fold ~n(,r~S'Il!i n th ~ treq u~m:y or (Opy 'flIlilmb~r v~rii!!n.tj in ind i'o'idual!; wijthi'lllll.ismJ lre!~liile 'to mn~nol:5. The cluorlilo~mal ()1Ial'llge~ we Fe 1n sm a ~ll'eglkli'ls and

dison!ers. the millorily of cas'!!:!> ha'ljl~110 ral1llH~Ilislmy of inni~m,fer reaMlTlS that ill!l110l ul'ldenl.ood, Lin k3l] ~ and !'!ISSociiltiQIl ~llUdi~ h"'ofli! J:iG!lunsmbiqaou ...y id en ~ifi~S!trDl:Ig g:~I1I'~',aD1(!id<lt~. whid'! l

included cllkl n:g:es Jill sing ~e'gen es,

[ust Enough Oxygen Activation

lllia I1V IrQ n· cen ~iliIIi I1Q '!mzymes ae l.ivlIte 02 ill I1d ham em, the Wliaizill~ Il (Mer alt he. readive I);llQ. r~uyl '<;;,Il'edes, ~Vh~rl1!J:ii Qlhelr~ IPH;I~~~t.lh~ from <iiI~thl'i;llt!:'d cell IJOWg<en. 'cry:il;;;llhJ9Hlp,hy studies T~~ p[Qvlde i!'!'sigh~ iate the mechilni~lIQs gf bglh 02 al;tivallio!1 al'ld !:ie(l(;tivaJIi.Q!li {'5ee the Per.sl)ediv<e! Df Wi lmQ't). l(;o!,l'al~f'ljlaJ a 1'1 dl L:i pi!i!col1llli1 tpJ. 453)' re i1IlJ1t Ihe SI.UJI(ture5 (If 5iup~'W;t;O, ilIlk~lp~roil:t1, ainu DOU ~d ·proe lE,t ullter mediatesin the O~ac:ti\l'atiQna nd i Jli~;efUOIII resetlo n of an e-l'llradio,1 dFlg· de~'l'ifl~ di", ql]'iiini!!~, T@1J1lhe·~, these ~b!,ll\.h,nIlS· defi fl!l t 1fI~ m~j @r ~ helllli,~l ~t'llps Qf lh~ ~ro~Y!1et nase nH~challi:liln. 1i<.ilt!(I'liIiIet' ot. ~p. ,Q,49) ~r~llped ken p~m(ide irnterll11looiat.es in 5tlp~roi!:fdi:' rE!i:l;ul;!.a~e and sug!)~~t. hQ\~ Ihi~ e,rl:tyrn~ rmilils the- rifa~tiJOr1 prrD~UI(I. HilO~, r~Uil~r tha,ril Flrllduciril~

re,a(tin ese-I ~rryl i1l~l:1mi1<1l!d i!illrts.

The Winners Recall M.emories

'i! Ele(tfl)phy5io.lo.g,j(a~. an:d C!e!lIuta r 11'1~lIgi1'l9 5Wdies !>how ~I!al oilly al porHo[!1 of neurolls a ~~.in'lii:llvetl i III
~ a giver! memory. Why is 0 Il~ n!!urolt fa ~hi:lr than lt~ ri~t!Jhlx!r, irdu dI~ in a pClr!kula F lIQ!!mOJy?Holll'! ,e,t ,(1/. (fl. 457) rotund HI~I. n euro.l'i!~ ~Il the ~a[erall affi~gdla la tl1.. t 'C.Ol"ita~11rlle highMt ie'l,J',eisor hm e,tiel'! t :3 ohhe IfanIenp tien ra~tor (REB a t l~e I i mil oHhe em:o-di I'Ig of 01 ~tlitCifi/ fear lll!mlJfY .nll! 1110'5i!! ~i!l 111]i3,t ~ p~renmti.ally e~"ro$i5ti)l;! ~cU'o'it~-reglJl~~lffi g~l~Arc iJ'flcr tliie 1'@(~lIllf I'tu:! m~,no'ry. Thus. l'IeurOril$ ~ {oll1lpele CilUiring memory form~liori, i:<!nd (RE:B hell)s to clel@Jmil'!~ 11h:<I! wirmers.

'MM~,science

mag.org

SCI EINCE

VOl316

20 APRIL.2007

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HUIMAfiil IEGGSARE n.J1 'SHOR1l SUPPLY, BUT MUCH liN DEMAN!:.L j'jrm~ MeLllen i~ill. lfue Y,'e~k:OIil1~' TlrustlC~mer
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MANY INHRfll[WOMIEN

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s.i.tumh:nn:' m3kifli£! hmrlim cg~ from, sten1 C;c'C;~ls·.j The Ui:'ii~.ooKiltgd0r11alJ.o\'\lS\!,'UIi:ICl'i und~crgo~n~ il'l "ilm, ~;';r~ilji:mlioUl f}don~l\:~1:!~1 t fQI",dlh~it el iniC'dl or researeh use. In e'I:JcamnJ:l!et"or fill'1k'te', lre:m'lrlem. "A hru ~t le" dnll~ll ion is also pel"lln~ue{l :un. ~i L~ j1n..;sibl(;l h~l~ t~. ri!ikl'l. with l.ntml1:t~Lry C(Jmp~n~O;I!I~OU1, Uni~~d: S~Ule,~ TIn,; !Ilim~(,,~vu.rnr.'IJi to sci ~'r:ggs; 101'~.IilfIit:at use <IS we,lL SlilL Ihere is 11m d>Jbme ~bam iJ:.C ethics .. ForresCflreiTlll!ic. ~1C'h CO\li.' OJ.' rdbbit eggs have been used .IS carriers On~iII'1rlm] ~u.!;lei because amy stem cej,ll~m!sderived trom tlnese VI'm,ldl be c~selfuia~~y

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E1ifJ:I.li:r tlTli!s mm~lh.<1 UK Pill'~~llUilirnll<!ry 'wmn'I.~ltl~ said :illlJl.lrestr~ctimlS on cybrids Pl\lpo~d b}r the Human rerli~iz... arion .11'1111 Emhryolog-y .ArLII~unil)·(H FEA) would delay ,i mpo~l[fH~l. research Oli! diseases, The 1~lfEAr~an.~ to cnnsult the rmblic !lbo~I' such research llili::>rr1lUmh cl~"Cid'~ O'l:llt$ pnlicyin S~p~.ci1.1 t\.s ta II <IS nbt'l~iIlg i:ggi' frem elnb:ryl()n~ I: steilli (ES), 't:,ell~,,\..'e· ve Iknmvn rllr more 111 lI'l:W )'(:.r:ur-~ C lh,ut "l{]lU!i;e sit;:m celb ~uixed wi ~h em'bryo ce ~~s(t:himel.1.s) pm<iuce mice with Wmlct~mm I grnnele5l. 'Why [hl:':l1 w,l~ ~I SWlc1:1I:'1, surprise \'0'11.1::1:1]11. 2003 1!l!1d 2004, lhrL'1O gmUJJ1~ IIl!porh.c:ddCl'iv~~g egg::; ~11.d i>1H.Tm from, I11ml51: ES cells 11'1'vitro rnthli:1' ~hm:] from chim~vicembr},{l!>?Afle'~lll, n!li.!l:~Y o~he[ li~lue.." ~Xlldd b(;:def~'!'~dfrom ESI,..'>e11~. On!!;!;;! dl.i.: t;UCt:I.1:lt'l r~lli::;ij'~m,i(iln 'of ~ilImmjJl::S c~~1:1 I1.. tI bocn I"CptJ:rtl:diill. 1~9~. ~ucmlot!l ~h(;iJ~l:dIon thea.!" ~ pm~n~,i<l~ lh~J.Olpe~u'~i;;v'l,I~~i>!', ~no~J~ nlrKiii;:r~r:;; Peg;IDl HIl~ki]]~ \~ uy.;~ ur i:rI~r.L!ci!1£ ES ~1(;':lb in "XI~~

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ti!m!Jera~~.IiD'l\ln to ..,so'c II ~ su.p' pooing aD sas. ~tal!ldar.d (If'IJCioced :MiPs !!I,d [:I!M!J, COI'I'l'binro WIth 't:ln~UIIIIQlJie.
lP~m:ted Ii:E!MP;yi:)e T"'~hn~C9I'~,
!~

~imll:es
~~Ie:!

U'~,degt:ild3!igloheluab!~' CilllstJlii ~y frt~~etlh!iw cyc.iE~

.r:leaml~ 0!1I!!i! ~I(:l5 ~I) obli:iIltl!l!'!1~ r~~!:i!e wl~.~~(cor )'Oilf' rw;:~rdl,

l1!r."! RE.M~ Sample Safe''' i$1II2sl;Jned Ia !I~, I.lb.mj ru. ~~ 1:0 grow -.ffltlli yevr ~!I)i MGragllln~cI~

Charl~s RilJleiroiflElrs a serles of wattdisease'

nnodel's use(iu~in Metabdllc Disease research. data is also on ou r weill s~te. .

Pllease contact us fDr addi~lio:n8J11 information, iln.diuding baseline, dala and 9!vailabiilliily, CQntrol

-e-cCHARLES IR~IVER

~'E,W -_ I
E.IDIlfHlIBY

I;

_'

vuouun

.IUIPITIACHAR]IEE

On Campus
and grnJmlte

[RIME S,UStE.RS. l.asr spring. ~OUfllplLtt:rsei!:ul.iS! r\i.i il[!llt9.I:l~ Black gmrc dE' I ,(i mu,~i'Yrujjlll,LtL~

!>tlldenls ill his, cOI'lllniler 'visiolJl class ;IlI1, mwn University.~ B re;;l1·lif~Ilf{lpjem!.l [0 \\,'Ork Oil: <Iii! nr!~I\<ed lluurd'cIF. Their re!i..t;.loch hel~tI 1'l:lI~rttY\\' tk search fur 11, Stl!>POCL. mt~ o!~ ..19'April, .B~<Ltkand his class .rucc.uli'Cd 11co~mleI1A1ali.O:ll [rm~l the Hellrico County. \i[~iJlia, police'. J\ll}ou2i]~lle clilpr.ut is siUl.m ]argc:.. Blacks class cX!trncroo ~~fid mrurmmion a'bolJl~1l1! cr:i!'Ilt:: ('riel!'ll h:hmry ~md,difiilClItOO "ideo t;,!.:elil by 1:\.\'0S'Lllfv'ei~~'lniX· ~-.,mems. ··rve I1Itv!;;r t~fld ,U class '~hl!i I1,Ulluwwd" S'dl.)'s E3lm~k.\\ho JeamlJd; UDl)!.l[ the C'1SC Ih,~n I.:olle'l£llc,the Uenrko police 11311OI!I~;ac!~dor help.Ille a c [ d"s:s used 0,1 vllfj.~uyQrnl~lhoos In de,,;,uI up the video. whid] slimlfs l,h!ll"i~1. tim, getri rig :imo the ~isl~ec~1;C~lr utsMe ;1c(Jn\~'1l em:e~t"re. BI.!ckMYS the o i S1:ud!llUIS devc.~oped aigori'Lhl'llls "thatwent beyond l~~ currem J;11l'1~~Url~110 tIIr!"IQI' dctermlllling the ~al:-:lire dimensiens ofobjecls .1111 scene. 'The.iJ' dle ill]~ Iy~is:;,u~~~I~d dl!U't lh.e ,(;,U' ',vm'·~Tny,:m. C.unry. \'im,~~1 992 ~n ~. 94. 9 and Ihi:lt~hedriver was on 'lhl: shcn ~idc. H.bulll 170 {lll'fltlim!wll)cr:;.
·,\V", huve ~ :;;~E~PCt~E IlUI\',:" ~,~~1,11V(;~l:igl.lw:r f\l'IdB~"''i{ Stnlll'l.OOI;g

I h~l'lrk\) ('mumy p ofice, "He'S w<IIl:lw>(]lloranother crime, and we 're trying to ~I!lt;h ~~~!Ii!l ~s;!khim whl:re hew~s ~hi:!lt ni:r,;l~t~:" ~!iId

or Ehl::'

II N Tii IE 'N IE'WS:

, IHECI OF A fI,lHiHRI P. A. Fre!lCil soc:~oI09 ist ~ returned to faln:s on 1,3; ,A,pril after Iliivin {II ill~~gedl,y entering ,alforbiddl'!r1 mHilary lome. ~ St~pha,m .. iDl!doi'(iiilM. ail expert on Cenual :; tI5~ai3iml Isl;:lmd~Ule N aln omit C~nt~r fm '" S(ienlific F!~)e:iJn:h ]nP'ilril~, wa~ iilITe:Medon ~,,"3.0 lamllary ane-r~illtill'!lg p~ctll'es of i1I reu$ grious prO,C!'.lSSio!1lill 'Sivs[<'!!1!-B€lIi!.!cllll'sta!'l,

i b~~n h{!ldlf n Iran fe r

g

ITI 0 r~ I:h:il 2 [I

mont hs for

(ol'lege [I,.all cneeses

fi rst state~il end illS!! Km'a's late'>l idea: O''oI'Qrritii119 t h~ ~I.~CIO tal
U.S.pre:!;idellffi. KOla tir ..l t.©ok an

~~e fOl.lrth presidend~~ ,cai''!(lida't~ llO wi n tnlle popula r vote but los;e<lhE:'e lerni[lI'I. In 2004, IKmtI--W:llo \ll~l;\I~I:h~ cul'r~rn
:S,y$~el1l'! I,lI1ir.;lir-I~IIlftH!d .IS

profit rafrVole 1,0lobb,y surte le9'f:st~llirl~~to allocate' th eir el.ecl@ra~'I,!I(Dte>5 to the l'1Iati:cmal

UI~ with the nOfl~

lT100'f'estin elocliom in
]966

as a gr:a:rn st~de]lt

i iI p,rmtime llorderni:llgAfgh.af1listai.!laru:l1

I
i

~ Pi!ki~!.[ln [htl[\ home tl), a sn lCI b~e Su flInil l'I'Iin()ril~ and has; beci'iI tile scene or recent " .mtig Mel'll ment vi,o~e;n(e. :,'. [)udoiglru:m 1lI'<I~ r~lea~d shorl~y after hi~ : ilrr~5:f ,jJmi 1,lO\ilSnot dl[lrged, but ~ralliarn ~, il!.llilorilies held ~ls pals:spo~l. He lived ~Mlil ~ hi5 ~nmi.aI'1lINife·s f~mily lntehraa unlrl~ the ~ {IiO'if.lmment senllnFll horne, ~ Dud 019 nen s[Jlidi nair! i Iltt'rvle\'i in f Lit Fig,afo a wee-k~~for~ his return th ~l his, resear)[h 0'1'11I r.a,llll':5, 5unnli ()omtllllJlnity .~ mOlY be ovtlr. "The ne:l:J lIme y~:u W<lllt tQ ~ (~me ~o ~r~,f\",hes,aid an Ira1lia:1'I Qflidal ![j ~old hun, U 111'1[1bUQ S.~1l YO!.!fln-~.:lWs~ ill1idnnthinq else."

'"'

:o;~llirtg

til board g!~me I:!JS(:dllifllh,~ cI~ct(jrat

(l] llR~Il!. ThaJl $l{.5l1] n, whi(h ,ag9r'E<gJt~sthe r IPopular vo~ fi1tOS[ilte'"b[l~ed ~el.e(lior~lll;1"()tes'"

and aJ.,'J\alrds each 's;tate b.toclD tile (i!llldid!!l~e wtHH'jj rtl th~ 111I(I~I1Jo~e-si that s,t.'u~. C~1Ilelect n a !pr€'§idelll who mOlY nen 11i~~ \'101'1 ~he most \!i(U~ ~nlll~ Ilalion" 11'1 2:000, AI. Gore became

W;llfll!lF cFf the populalr W~l? 'TIlE!s;dleme wou ld go iI1t(l effect ol1l~Yif elto llg h '$l~lE;'S :sigl1 on, ,al 'Ll,'hklh ~oiiImUf1l!'! eol.e:ctor,aloolleg:e w(luld bl;!(ollae m~~nh'lgl<e'5.s. La~I,\!,Ie~k, Mtiryl.:md'~ 9lov~JTllOr signed a law a doptin g !he' p roplosa l. Wll~,cll is u neer (1]n~ideri!lil)n in ,!\O oUm5;~;a:lt's. Kozals optimhli( that the 1'!lI(Ii"'~m(!tll nolj\i~,"~c oH~"Wh~ big gl~~~ wH[ :5inglt' ~u~'>ltol1 we'~t! goUen has been wh!IJ else t1 as done tf1li $i,~

Movers»
QSTWiA.RD IB:OU~D, lril ][5 ~ld
~(I

become a

lliIiljtH

re~efm:~h~,

~ PCH.~1II(5,
,<[

.~ IBAN 15 H~LN1GTHlE: 1(0 LLEG E (omp'l,jt~u

a.

Snmlg<lpore has I'l<lb'tlled ,~t another higlh~pwiiH~ W~5'~erl'l S(~flltist IE;[Idt~f this month. 5'O\1edls~ 1Y~!I)C~'~l'I'!i)t BiU!rWAn,a@r'SsCin '~;lve up ~he post of (hief e:r.:emtJve of the [mnpe~n 5del'wE' fOlJlndation (E:SF' to bi.lc::orll~ prlJvo:!l't at NiI'nyang TechnQI.Qg~~ill Uf1lryeni~.y" Ai!iId~rnon. 5S,. who ha,5 5erved es rectcr of Lililko:pingl Uiflrr,rersily ]f!1 Sll\'~den .a I1id~:s CllCilir Clf lme f',l,cgbe1 Co !'JiImftte~ fCl r Ch.~lIflrslry. sa'l'S h~ waffledl l'Q "be Pilfl or~hi~ ~defll~'J~(el~w':ii"'l1 happel1ltngin Asi,al.~ He found IN.anytli'n9',whkh has ~'S"OOO stude-lilts. arid !;1Ii(~ of the

'~ sc:ieluhl]ohil'l

KOla 0'( SUII'IJ!nrd Ul'ii!oi'eJsi~y in

~ :Pil~.oAl~o. Cd,lifmnla, a JJi;oltt'l2r of ~9I!n~tic s pro g~am m in g, ~ made his Ir! ark i'1 the \\iidle r ~ world b:yilll{entil'1lg~he sU[lt(h·orff kiU.~ry g tkk.et. Last week, Mary1~nd bEurne Ihe
~ ~ ~ GlJl a ~1Jl f.llr lilis pilge? [~mail Pi"~pl~@aaiH.or9

-------------------------------

largest engli~ee'rktg Sil;;hools in the WiiiFld,a~pe~ling be<ill!!s<e ~filS g!rm!i'il1ginve~tmel'1t 11n f.adli~e.s and S]ngapore'!'; risingR&.D budge'l. p.~ plrQVQo~~.a U'I ew pOi~itil)lll ~'lIiIer 1,1 tl~1 benei1J th the pn~sid eflt, • ~1'11der:S.1ion \vi II bl:!!in char!) ~ 'Of till ilGIOl1mic progrilms" ~Bl!lt my malin l1Il~s!iion i~ 'til I:ilJlild up top' level res~~rcll e'UorltS,~ he s.r~ .. Andefi'S,<in':s SIJCCesSOF a't [SF i~ Jol'irliMart.:5, who !'las s.erVi!d as: IESf ',. dirl!c·to r af :scie!;nce a nd ..irat@g~' si I'I(~ 2004.

\l'!llAl'w.sden.cema g, mq

5C1I:EN:OE VOl316

20 APR Il2007

PlA'NT

SCI!IEINCE

LD'ng~S'lougllht IPII,ant Flow,ering SignaIIU1n:rnaslk.,dI" Again

A~!;liU[i.i\.'e OJ;:> rhe top l;!lIiiIOlIrk.tthC sign!:!.i ~h;;:ll teH s p lams to fI oWl1:r has b e:l:'~ddi~ctl pila ru

biologists

for more !hu!'I a century w'it'l~

y~ilUi:>

mm'l'y l:al~t: li!i3~b, ttl ,LI~~dl!:tltity. Tv{o

<!!il(l.fe~ e,II:>;h~ r.~ Cre-,lte~~ 'qlL ire 01Mur

wi th J;IlI~ bll.dic;iU~fltJ:! I h ~t lIds ~,i gn:llli was messenget R NA lmRNA~ rhat traveled
St)'.,C:;III'L:1;i

PI",)c(fedi]~g.~ #w: .'V:aUm.wlih·riNl'em.I' (jf 0.1 Sciences, Elkl.cr U[sl(;~jcz of Tcchaion Israel ][lsli nne or 1,ecllmoioQJ ~!1 J lai fJ r,l_:f),urwd. ~H) !i.i~l'i of m.R.N A fro m ~1il,~ ,FT-equivalcili. gene ill l~C 11iJ\Jl!'c~ing shnets nr!Om"loe~, And i!!l ~h(;'drreuuctiun Dwtiee.. I IlLil1l1£ l'O:~I."bQ,mtors ,l.'ep,(II.I:t hartheir initi a ~ 's r ~I]:IjysJs ~xchu;IlLPd some dm;;,l and gu~ '~Nt[t~ 'weiglu 1.0 mlil()1i diatl!, Whcn Iltcy :rcdl.,j the experiments, "we- could not detect move:tIle::1U uf d1:1O:tlralll],j~t':.UliC''FT 11ii:R~A.," S"yl' Ow Nil~:iio!l.i!'l "'~lo. ..e hLb HIi!'HI.g did: ftl is 'work. Huaflg. I'WW iiLt Xiamen U.I1k!ol:iSily

T 11;' g ene ]n Illa'l:ll leaves h.l the gmw!h ti,p~ ...h!ere flower.;lam,'ring
!OClli'i

i1rmn tile

in (l~~n;J. h ••s not <!grel;;d 10 Ille retm.~~ In.,,,

Tt~lf~bllU und fie::ur.~~ Ct)~lrhllld of the: ~lia1o:Pian,d. II'I~tiHl.le Phm.~.Brec·di~lS

Ih:st.:tl.rl:hi n Co ~{Jgac. Gel' [millry. cwork ing '~\,11.th mbh/f:Jpsi...:. and ;lIlot Ille!' IC3rn $1 !iJJd}fing A rice, h~ve !fi()1,,!r IProJlo~dt~'i~LI Ill;;: l'i)JoM~~ 5;~lUI] :i~ Ih~ FT P mt.c:i n its!.! I f IF<LH1(.'r th.ml:nR.NA. 1'1 rice, the eqllli'VOj~ er!l oft he gene is

~ng t~kCtOpI1'lct:. But tho:;;~ author:;; ~I"'l" 11t1\'1' I"~tl';m:lillig 11\,1.1 ind,tug (p. 36n Instead, two F new n~l'i0tl~.'llLld islleclDti 1ine by Sci(!.IJ~"t: I
rhls wc..:!': ~\'i.'W'w.~(,'liC'Iiii!.lCn,UI!ll"O~th:!,l~f

r"r

corneF!t ;IhsH!Jcr/ 11411 JI' 52. ,111 d W\II,''i,l.'. se ie nce'nm~.oI'J:!h:~JJ/~ollterntt:lb~t m~~ f 1.~41 rj31.h:.~fiD1~~rcdll:lile "'"TlPmll;invb;;e~r. "1'hi s :i£ smnc,t I~i In.g W~ hil\f~ b een wai I ~1lJ~ Ibr ,I Iml! lime:' !i<!l)l!\ J. A. D. Zeevaart, an !lmcrilus plmiJl p,hysiuIu!lisL at Midlig:Hl SUH:e U!'liv~.fSir)' in EUl LOlll\ling. 'These two p:IP~rs will bl: t':J:cIS~IC)j. ili'n dn~fkld fur }1,e;~r,;; cflme,~';J,ddl'i Pbdip \:l,I·:~gge. a plant W
lJiu.~ogi st ,H th" .llohli! ~JHiH,S Cuntre

c!IUed Ud3a" Ko Shdln<lmOhl (~~'lhl! N.,r,~ Int>.liwu; 'l':Ii" S,c-ien"c .. m.d T~,c:h~HI~0!ilY i~'1 Jl:i~n. i'D.~llsHi.>L.k:nt Sh.tijiro T~mlilk.L and I'hc'ir

/·r

.un

coHe.,g~e~ fi'fS~ measured Ud30JmRNA in vari OLH, t issu es, Til lily fOLL nd t hut ill r i t: L~ :J;lJmvrQ.woit 11!>hnl'l day~ ~rice req u in~ ~ shart days to i.k·\'c~oP lil,QwcrsL lhl:' mRNA increased irll~a\I{!s bl!~W'<JS pre~em {ln~}' im
'!I'i;l[')' 10\\1 lIIlli:mm:;; ~ ~htl~~lIJ{H ,UpiC,ll nteriIII ~a~li!l. the :growing lip. Ne~t they m.~d~,u m~f'i'5.gcl1k .rice &Imi.n. by joinilliH rhc gCI~t: fur green fluerescern protein {GFPl' with l11n~ for Htlla. wh.idl m!mJ~ alllY Hd31! pwn:in

NClr~

wich. UK. Olher,~. hm'lll;'ver, rhinkthe evjJl.:fu!cis Ilo[ )/l:1 ~onc.~usiv,e. "They haven't takeJl the stery ~ny runher." SflyS \ViUiJlIllD Lucns, '.1 r~mn [;1)"11iD~~l;gH~l'lllhe b Univer${i[y OfClll,ifolrlli.fL. Dm'~s. This storyhas LIS, mots in. "-I J9]Os slm:ly by Rus sian p'hm~ phys,~.olo£is{ M.ilkh a ~1 C h.. iI;JIkI,ly~~Il.B'lsedl l[).iTI ~ [<1.11lIg experi~ ml~HHS,. (,11,a~lakhY:l1il roposed thm whc'lII p leaves sense the npp,~l'Jp~ile (I!my hmw h•.~he}' a send J mobU,e s~gml~ "allied !:1o.dgen 1,(1Ihe 1)!,nn~g~:o\\'i~,lJilli P 't{1i nil i;~le11m·,\'er.iny. BUl ['lfQmi$ir~~ ~.. ads hal (() J"'lId .,;:rl(ls. nd "I]Q.iI~~ a gen r~)ec~mel the p;Jrl~h ofhot;:lny. [akiuto]
Bug Font Qri~t~U i~l)m~ i?X tr~~I~r~1!5tri:.l1 life;'

PIEiIi"ipa~e'U(: ,l1Irot.t'ili'l. 1111 A(Qbi:dop~i~~top),a lea r

prateiJi nH'Ive5 lrnme

IlowE!r~l!uJl!il"ilU in,ill

no ['!fI@~cMiI'! 9 mUilta m, cau 5i.ng a !ilem 0l!'!1iIb;lassoms !JJ ro[l,rm. 1m ri(l~ {biOthJ.m}, ilh ~ @qui'l'i1lem praoleoi n ~gree,n,' ~i10\1l5, up illl tne shoot aj}'i(o'Il m ~r~~t(1m"

visible lil'ld.C~ a eunfceal laser )lC~HU1 ~l1llil :tni,cm£t'op~" They ~mW the pro:tdll~:I:I the \1a~ CLlk1D' riss ue ohhe 1,(:3 tmd lite LIpper siemas r well :liS, in the ('O~ oft he ~ IJUW~IlI.gtip. ,0, They then attached promoters [Q rhe E ccmblnetjon GFP!IIdJd gene th;n caused :1 :;:
lite genes 10 l!1]rn OILin due l,eDlfbull ,I.lO~ the ~ in :g!'Owin~ tip. flm!l'IHi~~!l: ~tiU {l~iClJ!T~. they ~ ,r.,;p~)rt."Tbc OIllI}, My [FT1 tO~I.ld~Cl UICfCii was gf~t IUO\'ed:' e;.,.p'liaillis eeVi.ml1, Z ~,

Sciences in! UI1l1Il:L~. 1:11:;; mt~ cdh:~:!:llL(;S pUJ>posecl. dmt mRNt-\ was ~helill(Jb~~e signal in

A mh;dopt;;ls., as the), :>:3\11'111 R NA f.ort! FI' 'btui W ,up in hmhlhe le;:lf ;md 1h~ grm,\ling lip. !\;I}'S Brh"mA'p~, <I ri<:!~M b~91!PJil!i~ ~heU!1~· 1:11 V!'!f1;ilY {lfNon~l Tol'XHS ill De::muiliL They C Oi.le! ud!Jd t.h~l~ FT m RNAwi.l ~.pr<i· dltl,.~ed in the' ~eafDll1dtroweled to ille girmvi.ng ~n file pi! t=l(~e-c~~e. l1es~'<m,;hel:s.al'lned d wI'til ~laI.t..>c~.klr~m)l!> libr Itial1~riJblLing ~~ru;::S. t.LiP. 'i,II,'het;e~1 \;va ~t:u:'lIl.~]~l~ed in~o Lh~ !:-r pruoff ntHVt:I.'ill1i!:J, lind \li~ll;J.Hzil~~ pmtdn:; in ~iiV~ l.i~:>u~'1~3ve tc~n. whi,chi t~(:nldckeJ [~v.~vt~d qjli,C'~,l.Tht:y iflililllCtl.d,OWI], lilc tlll.t' ~Sll:"l'eliCt·.9' St:.i]ll.:~rnoc., 2005. p,. 1(11141 .This I 1l't!1~(UI,sec'rn!.!d "'mm enl'lnlOllIs~y e.",dtill~ FT g~ne. the 1.:!!lfl'IIl'llein lhM tlJl:n~t'T on. ,1110 fl,[jwt:lril'l~g~~le l~,1!the FT prm~lli! 1"."(111-' bfe,,:klhI!UlLJ:J.h." ~(;",Llt"Co~ ii'llW!!tI!lbll~. ;l pi t1!]'t ,bI I lrokl.1~~n, i1l2UtJo5. an Ht1ang. .:l!. t'lu~tdot: III bhJ,iO£,is~ ilt.IJn~g'ial Colkgcdill \~. U.K. T 8m it ~m."!lot held LIp. ~n 1l1C ~,SApfii I 20[]:(~ file S,:v~di~hLlnh'er~it)' ~ll:" Agricu.~luml

L ik!e;Shi mmtl,l,)'to. C011]p.i!alild ;;lnJ Tambul] '~ Ibcased O!il the IT r:r(ltl;1i~ m~d 'II,! ~ G FP to :iil tr~ck.~ts.ra.tCt i.hi~ I~.mle i!ll At(1Md(JIJ·'II;·~< Laurent Coroesie:r. ill'wstdiol;; in CYliplalld's ~ .I.,D. ,~H~d~d lh~h,lsed. FTlfiFP g~mlt'" tel ,I \J .i' ,IUm::lllll ,·hl,bitHap,~i.> Hrain Ihllt Ir~(!'k~,iJ tin~ '0' rf g~Uie.lI~le"y (lb~,,"'TiI'~t1lh~ p,mt~;jj Ilq;;.l iill d~e ""$CUI;illr'li:i>,<;;(I~' 'OI'lh!! st!!ln, and .4 d!lY'" :~' j'lleU" •• n lht!ba. ..~ of the g!r11,.dl1f,l. lip. g
IJ ,1;j

i
!~

In ~fI~)lhcr cxpcl'~l'm:nl.. Ilti.:' [~::il:lngr-Jilcd phillIS. carl:'Tin!l.lhe rWl~'f;.I~ ~;I!ne til ImltaD1~.

~. S

350

BoUI. not everyone r~E.I',ec~~, U!sch~tz (~~I~ I!s so it ,co~~d have turned nUl rT mltsiJ~ the the evldence III both repnrrs "uircumstanleaf. L~C'<lI~, ~'H}rntsout. Even Ayn;:is ~til ~ •ial.' He" N iI[sson. and M ~glLd BI,[lO'Jq ez or cautious, ··flmig.clIl has, a iOIlS hisUJryor u dlo!iM' pi;;JllIlS. no nm,\,'cts 1(l:rll1lu::d _ Thll:~. du~ Ihi';.'iPoiyt(;ldmk l1n~versUt.y OfVil~~llCi,l, d[~;lppO~nlin!l pl';mp,i~:' h~l\;;Iys. <'\Vc'r~ !;,:e:tresesrehcrs could rule OLII both RNA :;UH.! the Sp;;Jin_. !ltlilU Oll:~ I,h.u nehher group tested dl~g thete, bm Ihl.: m~c .lIS ~mc~sc, and we existence of <! 51~n;!~ uC't~\'t~tedll'J' FT. 'wnelherG FP nlOVe~ ~hroL!~iIll'llepbm on n ts aeed to ~eep cool heads." -EU:l:A!BIEiliH I1lENIN Ism "The t:'!' idancei s I:'u:nv.~n.d ng,. C::'PCl"1 .. Hy own accord, Alld Lucus d!m::~!I]"1 think ~:he

P lants t I~at could [lot rJlwkc fT ill 1'L]I. The FT/G~;P' protein, but 11.10 mRNA.~n(Jvcd :.I'CL'OSS the sraf! juncuen antllhrongh the mutLlIIl't plant, ~ll~Y report, fumdly. \:I.~lC'1fb atmched two GHJgt!lle:i l~le~,' ~o [he: FT gene .. the r,e::iuhi.n:r,; Ilr'l)le~rl ~~'!!S too big 1.0 IT[;lvcl beyond Ihc ieaf--,md in

ll~!e£\rJf~i.UIIgexperenent s," S;)),'s Ay.re, And. srrcfi:;g{hc'IfiI ..ng the C'.:fISc, several othe r i researchers a re prepari ng HI I'll bl is~ :\~ ~l'IIilu r resuhs,

:il.lluho:rs adequately

demenstrared Iha.t fT gets .in~o Ihu gl:'uw.irli!6 ti pJrum ~hc'It::) r. Fur exanjple, illl Amhidops:is" enelearprorncrer ld:;t.~ 'tlUm~ on gt:I1~'i tl~e\'\!~le:.1l: in ahe p~arH.

OH.

RESOLIACIES

The wnrld's prOdLJctiori of oil wil) peak. ~'"~ry!)iiU! .~~I"~I1l~.'" So~~eti mejn the cOIiiili I'll!!'
t1ec'Ld~i!, 11ii!!: m,;:lzin~ nrmehi ne ry e r 011 pro~ daction l~ml ~~ouh:l~cl worM oil '~lL!fPtl! ~,:ew¥ decade for .. century wil.i ~~'UU~I'" utput O

75 experts rrom ~ countries to 9

C1,UJ

sider ~'he

woi1.d.·:;; ('Jiltoe$OLill't"t;_~_n~~:1i:l rre~~d he thL:."il'i 1"0I:" t ~I~ ir b~st (.":;1illnll~e~~ ~r ..:\'..::ryth~nt rrom
h(IW much ~j.:jlllni~n be lellto di!iC!/wCfH'Ihow much mi,ln be \~ rLml,~ from ,exi.~lil'l,i:l. oi~r~eld;:: a!l!d h(lW nrueh :lnil;lhl con.~~ fj-pm ll'll~~~,!iI\I~n·

\,';11 -~U]pri~~I'l!i:l-~V€!l! ~~ den'l!l~ll~1 'l'nl'll~iJ,llUI!l!S. to gmw.l'lh~~luC'~~iunl.lIS when,

Ferecasts ef'peak PI'UdUClliOH have m:ilgcJ~ruUlll T.~..nk:sg;i\'i~]gwl~~kcild2[m; to somi.fl.'II'J;e11e beyond 1050. B lit Oll tht.: <ln1nua:! n11":i.1lli ng u r tin: A me ric rillifl A ssoei m ion oK re!n~leLHll Geo~flgh~tsI A" ..I\ PGlin Llll1g Beach, Cali fi::! f:!TIi~~.i;;,n'1y IJ[~Srunnth. Il'll:ll~;t. ' e~~ i.I.!li.~!;~lJ(;\r ~nl~U'g:I;.'tI: Wtwld 'o~J pfm~I!.ICliofl could S,lOP !!Jowing :IS earlyas l020___;too soon ~o;)\'oid a crisis or j,l could hold olT mll.il ~.040_ "The peak ill \,'ifn:r~d(li] flG'tltim;., thm is not innln~ nen.t:· oi ~ ilnfOf.ilnali 01., analiy:s!t iehllnl Nehring r)"N~hr~tig Af;sm~i;.Ucs R Jn Coloeado SrrIng~. COlOr"lJiJO.S,lid .11: Ihe !Ii~e'tli ~lM. l1lU~it is '~!l!e'!,'e:nll1(;:lltiN.." ~hre:~~ilble:'

created three' sceaarios with successively higher pe,.h l:H~gil~ll~[Jjg 2020, 211m, ,Ind iill 2040. H~ th.i!11i CUnll)l!flld tllnl.!' aruuunt ~Ir e Hhn if would t,lke tu <l;ch it'V€: e.1I:i11seen ,Ing with ~lhe w{ll'hl oil irn,c~u;;;try'::'1)3!:>1 perfl'l,n;Jmc::~_ 'Th~ ~ ('In~y s.col;!·~"LriO \''l;l"re q u ill;l1>ll[lHl r

on

tionll'l~ SOlJI'CI.:!~ sueh as (:.8JiI:lJi".nD lHr saeds, :Fl""om the meclijr!g:~ di!'iC'lI~~St~Vl~. Nehring 1[!:JIl'i.C ~p\"'±t.li1luw. med~unl.al1ldl hHgil estiIfIl<l[~ uf ~dIlh~ ni llike.~y ,~~"L;;'r br;.! IlffldWlcl,ld. to [h~1 as h,i.: sa it! al. the meeting. n'w 11 iiro18i~~ It

is, [he luw om:" I}roclueing 1l2020pcak,lw

:S'MY~~.' onrerem:e !l.m~C'j,p"l!1h'were cenfidmJit.that ut t~c!>l Ill'iLllow \!SlUf1,l~ll~ oruhirnl!1[L~ oi~ roeSOll~'ce i-~OiCl~l.dly nut there.l'ltl !>;'ly~, And th~ world aHimJuslry has mm1i~,~d I.u add the needed productinn !.:OII)<ll;ilY;!;llS r<ls~ resouree ~ m)l tile unlY' 'c(ln~lrnint S lines ~~ ,'~::20.20 pocalk .wu ld rGltllui e, r Hohl ing the IJe~'kn~f until Th~e Oit P~q)k SC~D1!,aJ:ri(D$ --------------------------------------------2:UII:I,ilJ. however, WtlulJ retjJ u ir'l: both ~Ihigh -,[IIH:I mw.:l'l ]ess

ro

C'enll:i:[l--Hn.U.~ nil resource

W~h

~Jiidium
Rf!"QIJ![U'1

.!l'redictions !lJflJ1C Ibmtng ofpb"ilk oil h~,yc been ;111 gver the 11111,1111 ~ Sci~']~<?18 Novetnboer 2lH1;;, p, I .Ii 061. So-called, p<:a kists rowor gllt.g~ 1 [:anh :;:t il I tnd.d::; and h-uw nml.t: b has ~1,lreilltl)' ; been proQllced. lh~y come up With a !'leak ill!
01

i" ,., .'

:2 ,~

~o
~'5 :!!.G

Wii!h
IlMi

and mldulll;,! mOir~p,rodllClioll each y,e~1.1" than c\'cr before. despije h;!v.ilug <ll.lfe<Ldy prod~ilc;:cdll!ie worM:"lIlm!"i~. ~,l~il}i exiracrable oIL "\Ve (';;;,m't

behave now like \\,'e're going

HJ bl1l~foC' I:h.e high scenarle." Ndl'ring CO!llctllde~. Nr:;luillg is g~tti~.g,MHn~ auem ion but llI;<!t mall y con ~ "en~. ··Ri~~H!rd ~,i~ ,l!1 good

U~y!'ut~Qn~

g
I~

~J'{ld~'.CI:in.lI]. j'L.I~.h;::i:~l~ by ho.wUl.l._u.!:,;h

w :::II

Ii:

s
0
"

,g;:

Ii [he l1;i,.':Xi

yt:laf.j.,a~rA;~ti:!1ly boef.h~ ~{l20,At ~ ~heQI~~ife:,"ltrerne. nmjoq: oi ~~L1iIHfi~lni~dfuw on in~~lutbC~xpL:rlist:' ~qb[Jo1J't 'how :liIli.it:.lloil. ill. r.::mmns lIml how f3:st II will be ,,:n:uhu,'elL
1'1::1.'1;'

:~~l!I

i
a

S~~;m~r ,or I~ ~~r; Th~ l€!Is Oil 'leU l~ br! .p urnped from tile grl)u n d,. ~he .. f!o1lr~hi!i WQrhii pH;I!;i!l,a i(lrl fii!'o1l~;hes .~ fi'~ijlk. IIIl !I n~ ~ n(l~ys.i .... oJ1lI~ bh~

eail1~e-sl.~ClI·Nesou'rc:e p'~ak li)Ob r~lf.ablE!', ;t'InJ ;:;:a~il)~ tll1ir~~I, ~ml Ihflit how :H1'I:lchu$t! t l.:lN:pitli.lClll. ~.~!i:. .lii~iprt:iiJi ng i_.; 'tl'day 1:1, 'I'/cnc,zlloCl!I, lI:"dq.• <lurid Nig_o.':li'i.i.And
.fti:;'J}i[lIl."e~,~~ti'il1e:

lli'E'j' t';etl· 110 ,~ndlln .ri,~ingJ'i.mdut:lio. 11.'" f=li!"Olll ~~~

lh!!,Y l~ook,L!ils:;u~lI not kyol.U~ 20.:m. ~ ~ NL:'.llr:irig 1.0 ok !It~iHe rtl iiJl l.m: k, ijn I.Wo ; U <I,Y s. Fir:'>I.n t: clll1.dlJ:"'~i!J !In informal :lUrVC)1 I
~ liS ~ of CX~~L;'IUi~r m~ 'I<!lij;.:in!; ~i .mJl~l.irlg. a p~)\'~u •.

"e[vioo i'n h(l~l;Iing!h~~ Hedberg COIll.rcrcn~oc," :;uy~oil ~:%t:1i;~~nelllt spet:i:d ist DClmald GOAuti.er C1fth:~ US. 0::nlo.1lLt~11 S\lrvl1:}' in MiIl:lillo .P'drl:.. C:~li fQ,mr~'1. But ~h~re·~;S(l.ln~ch
iJll.L::'L"rt!li~lIY~ GlI~Hicr :>lII.~s, from \\lhl"mJAi't~lh::: ke l.nisllllfi.el~ (lLIloFIi'te \'\~y M 'when ~a>Joo

~. !,'!hms I-h,:di!)l:rg Cm~lbr":l1icc, uHn(h:r 1.1t!.!' IlU:S:~ !'lices, ofAAr'G b~t ~h)"l!l'mber and il1vil~n~

t.C'ch!Hli(l!,! i~ut c ~OlJ~~m!ilCl!';.;!~~ Ihe lit ill· ill kX'~JUUlJ Arclic. Cllii! ~l.O'!N C)(lim~liL!i!l~S well. S(~for h:~s~ec(llldin~cwal~f:ln. Nehru.I1g
SCmENCIE VOl.316

n~w ~1;,·t"hn[j~ql,!Y bt;"d@.'dop~d ~h~trre~!il'l

diclin!!:l th~ p~uk ~ul)y:!lOt b~ "vurtl1w&n.ilc. A dee.,dc' QI:' ~ cOI.dd ICn, -R:ICHAIi!O A.I[;!ER:R

ViMI'IIII'.sdencemaQ.org

20 APRIL. 2007

NEU1R:OSCiENCE

Ast:r1olcylesSec:rete Substance That [KUlis 1Metn r' NIBurons, in AlS:

AS'lL\) t;ytcs,...-all1Ilmg (tie g] i!)~ or ",'HIP port," .• cdl.s, i.n rhc lIelUl~I.Ult:I:VO.uS S~!l~:t.rc:m-m~iye h the prhnt1ry '1.:',,11 I iu tlli,e tieJI h o f I]lJ'IlO:i pri
Il~LIIron1i'O.in atIenst :;;m~h::: '~"SIO:;; nfuruy-

Texan Mi:s!h3Jp' Not: Reporto,d

Texa~ t!.&.M Ul111ver§ilyt'l'AMU) llnl(oUeg e- .staho.1'I w ru FIlling a!~Lll of rE!d@r<l1 bio~efl'or rul.e5afte;r I'm em~~eE!' ILan year conh.;lHldmiU,ed tr<'!ct@d brtJ('!ll{ffii5!, tin sn im~l dlise;:ls@ li1<lt's high on ~hf H~l or pOlenlial biolerror ~'apons, in a :5laleme1lt to tlllE!'(.ellll~rslor lI)iseas.e ((In' uClian d lPle'llenlion (WIO in .Allafllt.il, Geof91a,. made l<lSt 1h'El~ the ImiveJl)ilyadrnowledge!'d ~i1~tIt should have flolliled (I)( 11~I' <lfj;O ab(!!!J1 11111inddeflll whkh came to Il~ht ~Uml~ h a rf·eM(lm·(Jr·fl~~(;jmlil re(Jue!J.t from Ihe H(liil SYllshinil Pfoje-ct iI will~rndog grQLlllil1l iW~tirl. Te~~" A ClDCii'l\itstlg'.pl!orn b~ul'I~hb 1iI'e!;!~ (ouki reslJ~i lf1I1I1~S and tess of federal funding. To!: rl?s.eC!r~he f ~~ diC!g!lQ~ed w1t~ b IlJ eellesls 1H Ap ri l 2006 arter dea ni ng an aeresel "INim~r (Qn~~irlilrtfJl1fu.c~~/a b~'!erli!1LI1 !!

coaxed the ES ~.:e]~to become motor ~l!etls rons fllfDdi did sernc mb:ing and 11Kltchinj,!.
ell ~Iiv,m il'l~ the 111o,lOrneu reus end astrocytes WUIli! and wuthDut the rtuumli~lILThey Ill~l'nd l~un e VC.UlJ~onl'l neurons [[lid trlld Iy in d~c Ell ptre...ence ;!)'i·tfOcytes wi~Illhc SOfN mlU~<l-

mmphkllalcl1Il seleresis tALS,. reseerehers at Harvard ;md Columbia urnversjues report, The [lew rnllding.s undersenre the pmv'L:l:r '0 f rexea rch wit hi e rn.b.!f},Oll ic .sI~Jn f~S~ cell s 10 elucidasc basle disease

I:iml. showll'lg a 50'V~ del; rease o~{e~:14 days. R'~\'CULll!l, the (: tmdi[ ian1oi.~.hc;:y tilundl h
when m luam motor neurcnswcre

or

sur-

,I~

precesses, lite researchers 8;1)',', rnut:l~ted by n~:nn:mla~tl'(lc)'ltcs,uheir losses ALS i;:.; .m iIIIl.ttre~J:taMI;; disease that prowere Illil~l£h Ie:>!> ~evt:ITt~·" !il:re~si\~J,)'l:;m~~llUW~' ,~'~LIB.m!'i.F~lrye'~!1>. E.~~.m S1UY!'i I,h~ wnrk. "·re'I~ly v'Llid'ite:;; M:lelill~SI:i, tll,'i,; dcbmc:J w~l,i,;thcr ~hc motor h thls !lOlJlg 'W\C'n: been ~ingiflg: lf yuu CiJliII neurons themselves aredefective, or moke ES cell lines I;'Jfry~ns In.e gellle~.Y91.. whdl11;r ~mt~t'~:'I;J'er~ll~.~rUClO~ '[::iIIs the n~" ~,ln slUdJydi~Ii:l.!~~~il1. a mHl;:~1:Il,ure s.nphi~tiTwtJ papers, published o.Uliil1.'eon I 5 .Aprri~i~ eared W<'li"" He phm5 10 do the work wh~ Naiure Nejw(l};ci,'~jce, lru~"::l1i.new evidence I hrtJnmm ES cell poptl~l[tiurl~ cl1!l,till.~iJI:iillg tit(:

DNA ,trolu ALS patients. [he C ollunib Ha lCllm..~c:mdeJ h'y biolo~i:;l Se~t.' Pr~~JboUl>k.L to~~k Ihe't'ili!dings <1 stell Jurthet' by de I1U'ln,~lnHini:!: that !,l,.lIf()cytes ~.re ~oleJ~i re:;.ro~l~i1J~.~ru!i thi~
toxin, lind thi!i! I hi;;,~1 L1l010r neurons. They fOLlilld d~~i[ gro,,,,,i.iJlg morcr rlJ~Hll" rO!l~ i!l aeulture in \\~li(,:;h OIst!'(1o'particular

WU'lsrnis,~i[i1l lOr lh~ p,a~.llIogef~ e-Jmem~ly is rare', S'UFlslh in>!!Proj e(t D'~ri!c~llr Ed ward

bio~illri!'!lylevel..3 la,b in F~hruafll.l'AMU 5<l~ ~ili! rese,an:hI!J rei:lwere.d i!rt~r tJe.a [Ill ent with ailtih:Jltk~.l,ne~ <iji~ a f·~iitll'iliiild~d US Il:a,~e.$ !!)!f i;m,lic.d~os·[,s~achear, but h uman-t(l"h!lln~ n y

HamlTiiirlilil say$ !h~9°ver'iJirrllE!n!.IIIE!E!d::tDi devi~ .a ~Y5l@ml!:I <l:c:~~velya~~ 'fIud, acddenlS. u =.MAIRTiINI ErIIS<UIDNII:

INIMH IE'I'esBI B,ilgger Sitie

,~ONDON !l;cmlr~rw TtheupcQlillTl1g C!1\'ail~bili,t.yijlr,ij
OVef

onlymwg>et

l.S-hecn:are sHe ill (Enlr,[l ~Lcmdon (ouid e F1dthe

I!N~llnlnl~l!ll, 11)lil!·1!l5 W r'eIOl:oiIt,e

cy tcs bat! bee I'D. grow

111.PI'~:IIJM

eed

time ~1~U!' de<1<1lyresulrs .1S C'XptlS'" .in.g [h~mn to '!J>~['!Jt:ylC'" ~inJc~.~y. rrz~dl:m.Tsk ~ ~U}'!'t h;u lmvVll..; '0 <! roxie substanec emitted by ~hc ~~i<JJ ~"e.IJs.The team ruled out both SOD I mHI gllllll.am,uH:-1I
~ BaJd liI,e~!J hb~lrhooo.

~~Ie iMallorlili lusUlu1,e ·of lilI.eclr(.a~ Res.e'm:t. (N lflilru IWiII1I 1i:s, famous su DU roan cempus in tl,e ~Ill. H~~.afea to ill O.4·~leclare 5l~e in d'owmol'jl'l II.ondo!'i, Clirl1~~had (IJJIllpl~il~fd jll1:t!i, ~le
platlne~ ~lte was
sg

~ltiian It ~WJI,llallin1il IIM.lf) N

.~ (glfeen.)illl mllul'l' wilt!

AslrcKy~e$ ~rtlcll) till f'l!iI!.e.n mol n1IlI~;liI'lI. !icJD.l gifne-,

01" n~u r(N'IS

~ ~h;;u in smn~ AlS ~~lLfr<i)'r~Ir:'i-~lu!M ,\',i[j,'I " ~,~mlllillU gene ror. :5ltP~[01>: ide d is.m,uHls~ ~ (SOPl) the aSU'{lcyte$ em it <I t:(J....in !ha~ ~: s..:Jecdvely kU,lslflllmor Itctl.ro\l1l>, A LS re._"l,nllrcl~cli ~ J ·1'1' '0'" . ~J h 'U'. 'i~' '~~."

~:

~. ~ ~@Y~'~~~l~'>'Ie!!l ~:~ ~.!lS~! O.V~~.~:~~ .' sn)'.till Sa ~1.lnltli1l:. Mary laultl. s.ays I,h~ piilIp";;i ~

_~~~lV:~':

~. '"add ,\'e~ghllo a growing body 9fdmallla!

~. ~m:s:S~lggt:'~tl::dthm a~tfm::Y~I?~ CQi~lribuh:: to 'll f.\ l S," 'Roug.~.~y I U% of AiLS !c"~'~:j <lfe .~ famiti3L wi~h I.h~ ru'st bl:i~lg .lubddl, ··s.P'l)· ~ "'Jdic.'· SODL th,e only ~em: timt's su fllr
m
~' 1J>I;'~1rn

bmiua ,di...C'a~C' 31~o imp.~it'mctl in ALS-m, tliac ID:S:ic substance. SE~'weu~fi oldman, ;;J, Me m ce~~ rc..'i~'lrchfj:r at ~l.1£Un hlCfSll]' of R·ocl,ms:u:.:r1\,~c cllh.'al, 'Lt"llterin New ''n:'brk. S;MYS ~heJ('e'!ie:lrchprovide ..... "very va I Llab~1 :i,ugde II~' ~:'or till is p<lnbc u,~::rr' e typ\;'i r A LS •. Th~sll;l Olre P{H~'li!ti;'LI~.y qlu i.le c:>:!C'ilin!;papen> Ibr i.i,um~u(:ing ni,i:\V a''I'\;;rlm;:s I'or rese~rch:' (my!> Gol,clllmm. lfle says ideMti'lnajt1f IffiLlbkm3kcir.in
(l

'!.CI1en(1l! fSdt'lJcf'. ~, fei:mJ[lry 200S, p. 6S2~.In .. Iifl'tl\l!r i!llJ:iler this m~!1th tl;! NIMR st"rf, (!:)lij!1 IB~akemgre,direcltiJ o~ tile' Meilical R:'I1s.!Mr>th
('lijuli~il (MR{ls;;Iid 'th~,a~elic)'.'lI~~ril;Ii,!t (~ruidertllg with ~emln.l~la\jm~ a larger sile next te Ih~ IBPitislhPllblk Llbrary ~lJ~.:tanolh@r g(lll,l'trnmem OI!Je nr.y ~I.a nJ III ~t'lllt's ,al§O !!leal' rre~l'a~~hh(ls· P]~~!~.·rli.lfiUing M RC's desiilr~lor ~ 1!'evfll1!lll,e<l INIMRto 'orus M lral1%1litll!la~,es~rdl. f:,JIMIi:'~ Rllbin Low!!U-hd~e, ~ crili[ or Ihi!! planllM rn'~, agl!i~e~ rl13llilE locatillai ileal' Ille lijllrary is m(!;re I'lr(lmfs~ng but wa r Fl5 Ihal ilS ~05! ((Duldi s~jU ,E!l'Id up' ~11rilll k'llllg MiMR. ~TIl~ devll is. in III~ del.ail~, " he sa~:5. Btiill«lmore s:.ay~ 'Ulli!nh~ nell,!si~ Jl!i:S d~i! iidvilirItatg,'i:!), altliough he 5lNSSes lhal ~JH~rigina~ one,. which u~ed to, o h{lu~ [lli@ IN~ti"llal.lli!mperi)n(@ H~SIPJnij!, is still <!((e[::lti!b~e.lEIla~.em(!ire MilL's ~haJ'lce!s Si!~ '!letti!!1) thE nell! sit€'are undear and ~o ,<lf~ nlS (OS-lIS, till t h~ lnope5. o hilv.e NIMR's,fllllJll re t re~dlwd b~tor.e h~~~tps atlwf! a~ MIK diH~l:wr

rY~~l~I!l~ agli:rn~ bl.:l:itll!jite!~aM?d by ~i~lt'oc}'tl:s .. IUld ~lh~:rl:l;.Jte!y .\1a:,ein:g "·w!:Lelhe:rlhi:.

type o~' In,.;t:hanisun ito;

Ilim,itCocli to

d~ SOD

1,

'.. _, ,.' • , '• Imked ~o the {h~~a~~. ~[.; .lit.;!ll>l;'d u~ m

~. IIboll& ~

2.5% ~)r~mU,illl

t.<1b.es.

~i'i I.h~ 1!11.~s;t. \i,,'ork" ,11.Harv~uI'l:1£rnllp

~ by K.;:\' i III E,g~., liD

~. MIl~c(lt'Y:;;~!;; 111h.::e br~d tel expre~~lh~ l1or-

'I,r

bj ~~n~:I'''ooCi ceil Is ~'rorlllthe ES

'fjlU<i~'

~ ~~~Ia:l human .~,()D! Cll~ Hili.! ~ u tin: I'Inmmt .",.oJ)1 g,cne. lhe

m~i~'.C' ~'itJn

~IClltll$~~

then

mod!:t" m ..y he'llm unlock Ihe ITIi,Y$leries of !oo'PQ:r~.d tYIlf:."S of A LS ;IS 'i1\Il;lU. it: Th~ Colmli1hi2l, ~MUp pl~lfIi\ 1.0 dl!.:l1n~ 'trr~~ basic djamc't!L:'ui~.E:iC's t)f dte pobQUI a.ndi El'ld lISC I,hal. ~n~wli.ld~e 1.0 :>C~f:'1i'I f'o~ nt:'I.\' dn~'>. "fnr'L ~i!ieOl$~ in ol.:led ofdwg~ Ii!;::.; I~w oth~l}j,.th:u would b~ gn:at:' s.2Iys lE~un. -(:(INISTANCE HOLliE]N

or

~Il S~pl\fmbN,

"';JIOHNI 'liUV[S

NEWS OFlHE WEEK

A Il!!W spin M~. llile ~uolniccI<H,;k cnl~~dy.~e~d MlJTlI":of [IICl1llJM.,:m ..[;isH measuremcms to date of fundamental physical COI1SHU'Itspotcm ..~.~~yIP.PO\'.iJ~f'lg eruciul expcriruesaal i
j

hl"lrs sii:b!l~lly more energy Ih;~n

does. (lrl ~no.ml1iclocks, c

~~'e'

o!hcr
of

:r<hifll~e'lw~!;mthe

hO,lll

states, The collided

Um fnrq!l1l:lm:y

lil!!~H emitted
bCI.WCeliil tli'lc~c

when cesium
TWO

aloms mow

"hYIJt:rl1ne"

~C·S,lS. of~.

number of "thenries nhwery·

,IlPclmsylJ\':3I1:i~l SI~IL1.:

(11Ing_" !Four rhysicislN

U,I,livcr~~ity

h,]\,e' h'lli~l'tine ~~r()tnl)1~e f'a "quilnnu1Il SCiI[e

leri.llg fn~e:1,1eliOmeler'·--a de\ ice capable of re1'!~:'lerrin!!l d ilftre'lm~~. ~rlJ the; 111i,ISSand ()'~ler propeni es (I r a ifn s W~I h. unIH:3rd~of s.el'nSt~i\'i,ty,.1"lile eN,re:r:ime~11~,described tirlils week ,~ll IV"trU\r!. C'llUld Ic,ud to mure-accururc ,I w~n ie cl nck~ ,I IUd het p phy,~ ic~SHi sl~~dy
C;(Qt

(Penn

Stale)

ill State College

P~ml State experiment, the bring rh,e two clouds togf.nh,ef, ,,.\s the (11011115 c'om de. I he l W() hi)! pel' l"ine stares of e,ac.h eesium :IWllt<eC'nil di:rre:remly. {:tQll'\!:ing thCIiI.l to :i.mcrr~.r(l wlrh each other in ;~wuy ~htHc reares at im~ lag or phase

vide's the 1~IiI!P~l f!.! o ~I,l[he researehers

standard
seeend.)

11l3.t..:'S pre for dctt.:nnill~ng the

~'tm:l1\O sent ~llrull.llh .<inlli~lIUwavl.O l':1\i,mll~ are ber a second time a nil then are sorted Lls,ing .11lscr pulses tl'l.al. taUy Ill.:: phase gbufcs. The l'eSlilils provide a >t!Iet~~led pO:rIn~il oflhe colli~Um1l. (sce fil!.nll'~t "I]' yml .rei.Uy \\1~i:H to 'L1Ii1,dcrs:lllml etom-atum 'illU(rflicthms very

prociSJl;lly. Ilai s ns ~he mel ~od:'S<~}l:-;Kurt Gi1bMe ()f' I'l.n: Il-lllln State team, 11'1 ~.IiIe'fiulure. the seienusrs hnpc 10 :l'lIIr" 'L~c'r rin':~~1Hne t~lC e:o:perillnc.n.'l by [ISIng nl;~guelic l"ielcls to coax t~e arems ililto sho rt -~i"cd C·OiUl'lX: I iOJ]:;; ell 1led F~l>hbach :r'e:mmUil;.~~.\l,which r-e:s.euulhle moleeular
bonds,
Bt::C8L11.:$cl~1t: ;t],tums ,~l'It!l.:I:.lIl;t

il{;

~,t,OItes of matter

::;,~iC ~LS D,os{",-, ~

!JIveI'

Einstein condensares and degenerate Fermi gal';(:'i"'.~~'ysRanda:!.! HlLle~ of Rice IJ [J,iwrsuty in. I Im~s.tun, Tex a s, "h't;a new Iligh-prcdskm
t(m.~

for

:tnh:a:>uiliiliig

[he dIceis
says,

~)f[.,toltlli,t:]

iflnger pericds, researehers will he <lh~eto .k:nm I1riO~ abo~~.h~:i,rp:mre;:: rties; Th€ jfC!'Mlt naaces should enable physidst:; to '.P'p:ly ::!tUIli'll~c~dm::k ::Lc~umcy hotl~ to, 1Otud,iCti of
the coHisiolls and W meOls'ureme'Ul Is of rliHI~ d~!ilt;JjH 1~~i1'hy !lie., ~ C'OIiIS~,l'Il:lls-su,d'l astil!LO' rati.t, onilit: l'I1ial:\1>onl~1! electron tJofh~ maS;.,> nr'the I1roton" S~lme\'el[~·~('Ins of '-;Ir:~flg~he~ (uy :<1 nd ot I~,ell: m nd IIn i. ftC,lti on t heori es ~ :prJ:;ldicrtil,n ~iiJlchC!CI"_~!,m~~ \"1 II "''m'y~.~(jwI.y owr 'Lime. ~J Oi bblc says. 1m u hrase i"sisu, uve I"w.ure versien Ihe quanrurn scatterillig iill E~rrlJr{JmLJLJ,J[ \\luu Id be ,::Iblc w rucas'UN.':the \'<lriutiCl!'JS U\',i;,lf ~he.s;p;i~ ofy~Ol:~.

~nteml.:llens,"
cljtHld~

Hulet

Thill' resaaruhers

~lO! t by 4;:oa1]l:i!;l I.wa r

uf cesium atoms to fr,!Ict.~tl~H J1j, ef m ie red e£, ree a b'n'!' e a l'i:st) hue Z:C'fCi, o ne cloud! ls then put ilJlw, its IOWI!SiI-enoel,'£)' ~n!l'~ ,[ COlU ~d l he- !il fIJI 1I~.hJ 51,;Jit~) ,W~ld seut n'lrou~h II ,~1l3Ijnb~r where raiernwaves excite fhe nl.an1\" into 01.b:f7.arn: stare '"OIUed ql.Hll'I~.u.ml supcrpusili,t.1Il'I .• in \libic'h the !nUll!t el'ecrr·r.lfI il'! e,'eh :lHl!TII h a~ Oil pfCl:Jle.Fil:y (;<l,lk>d iI1~ spiua pointed bOI.h upwaru ~~tldt1U\\'Uik WHrd,~t the same time, Om," :!1',Pil1 stnre har-

or

sp!l!(i~l~yilr~pglfed
()r

M;ilillil tine 9 a p"Tirne lOll; (a used

1

1>11'

,tll!snUlH

atmnk (OmSiORS <lnd

atems c:o~d btH)st Situdi~s P'hysk.tl (Olll.stilntt.

inl:erfeJe;nce @f

"MiARK, AINII!!IERSQ'N Mi'lrk AJ\(jefS!ln i~ wriief .. NOfliwnpOOIll, MM.saC~~l~> !l

PSYCHOPHARMACOLOGY

Studl', Que:s,li o'nlsAnlt:i dlepres:sa.lntRi.slk:s:

Rcs.;:areh lusrirure ill11 Oltiu, W~th Dsvld Bn:rll of W~ stern Pi>yclli airie l 1'I~lhlHCtll:l.d Cli nfc; in Pfru'Sbm'g! tI, Pe'lfInsyh~lI'Ii a, rind Il'acI,!:' >colh::.!g~.es. thC~H1UIll ~e~ nuttn mimic HJI\,f;, ~l.pp:rolilch \\'hill! im:o,rpor;III,ing some liiiI;JW dHa :md n,::pcllited benel'il.~ ol'lhe '~nlg.". As I:OA d id.1 ht::.ltM.l.4ii1mlmrs eXt'l:I1l1 un1;,'d des '\!I'Orld,,\,idcwarned dOO10r:l 10 ij)~l:g:ltCm ..g\'~~cQmp~lI:;;i\'e ~.H~UI'I rf!:~t;ribing tb~ d:~lJ!il:;; {.~lil~.ke'~l'l!!~ Iril.li:s gf d'~rr~s:;iQn, ob!!i.e!'j.. to Lt;ll'lliil!:\l..'-~ btt~lus~ d1:(;',), mig hi ~~~,~.,.;: ';'sllici~ d isorot'![, lind. ~'lh~rafll1'i IICty d ~soUd!.:ffi. bri.ngifllg wgelhr $3 ~{)I s~~.bjlecl.~. Dis"lI"der~ (l'll1~y:' SJltCl: then. oomro"'ersy h!j1[,~ ffi\'i'~ QI.'ef g _,;;p~cifioC' :,p:mly~~sdid Il{li. yi~ldi st.m_i:'l:ic,nUy \1!il",1.mer aJl~ riljk~ ha'!. '!!~~~n ~'l~r.:lk'ti. ,~igHiFi~rlnl re~\!ll~. !'1m wh~n !!II d<!t;;;[wer~ the <!lllho!,!, ofth~!lew~tl)d:y. rl,.lh!f~he-~ Llmis,~vc,~.kin tl1lJ Jmmwl (1du_"..J~merit·tU~ rtlo~"u:d, i.lilol: ~\ilit hc] [:s, fo und i I~ ihu:-:~ .oIl ~tt medh:al,~()n '11Ma s!Th;IH bu., s!~ltis.~i,c'lIUy sig~ Mcdinl! .. ';\~.f1(' fat h~'n(,i,lj'J~J ,! I. un dell'look b .. tine: il!, ,I n~.ly!!:i.~ ,LLft~r tile US.fl:ll1'l.l :lill~1 DrUM I~~fi.c~Ul~ inete<l!!i~r:usl< llf suicidal. d~ ki!1~ iii Adm iI1i bit ~:~u tF DA)' II ia pp~ d II '''b~~d. .and ~,~'1avi or 01"0., '·Y(I. Thh \ 1I:sIJ;;:~' lhm] orhC' ion. FDAr[;'il'l'clI, 2% lI.ii'l'k. TI'l~ 3m~fii"J; al~n found b!»;."\\r.!l.l'iIlir'i!:j Oii'i.most iJ~1'ti.d~r.lt:~s~_ifil drn!l:i ~b1 Im,le 2mJ-t IJ ut that Wllril illIJ tI i~Il' ~ take Ih<.'i.l.. ilil ~encrJI. t~eIUL~cl"~ w..:re hcll~!!J by m.u~yufth~ drllg~ te:;;~:ed.D~pf.e~~edl t"hij~;MCC{1U~tl '" the~11ed iC.L~~UI1$' 11 ~nefi~~. :my.s 0 ,}e:lrrrey Brh:lg.~. :alll \!·ph.!l~llinj:u]C!gisl who sluJdr'CI~. :IS ha!> h~~Ul Pi"r\'i(lUS.~y repurlctI. W{l.~' h:~ teen ,~~ic~i!.,hi: Columbo'lls (,h.Hdln:n·~ :It hl'!~ped onl,), bi' Pror.<ne, "Whm lhi~ :iihow~is

An 8!lllliysh. of 27 el hl~.!..:;)~.[I"i[lls nfamkk:prcs-sams ill1l )l(lU~g."'crslm:; Itllmd IlUil~glligibk: rillk of Sludda ~thou£:h.L'ii.md SlJ rcide am:n~lm. \\d till dte ~~:re:d grol(l!iP~ S1ilm\,jng (n~;.,I;lreOltl;:r r.u~k : L~81l panielpams given a placcl:l~),.Tilf ~Uldy ~"!Omes Ol1ethall 2ye;~ alber :I:egulatul}' ,I~,en~ m

IJDill

IJ~C f~",cran

the everall bad eflect is very sJm.lm .• md .!;t10dt::~rtl:thi mueh hi ggcr:' s;;~ys

t'I'Ilill"les Nemcml1l'. d~,lil" of'the der[lr~mel'l~ ~, p:;;ycill~a~r}' :1IiIt!1behaviorul seieaces ,~tlt ~11:'1Hl:ry Uilliverl>hy i111 At la_llm, Georgi a. ::. IL~'ke Ule;~~ly<Ill ~()1-~,~lIe>d met3-ml'lllyses. ~. I,he J1.1 AlA :mld,)' iii a:s it scj, II'wwba oks.. .~ re~~<lrdler~ 'Igree. T~~ twu,111,';q~ ~~'~Il~~I1e:d ~,

or

VI/i:n; .iWl (k:::;lgile.d ~tI ~~~I:se; 5I.1i.{;it~al bcha ..... '::1' l'iJl· ;md e;o.;c b:!ed illdi",.idtl.;,1ls \.,noW'ere ~\IIicidJt t\Jiiln.H1gh th~_-;e ,~~I]g~ lJ1ll)' p'rl.:.1E: ;l 'rB sk. lO :~ Ci, j!OrlT!e. ~lIYS K.eny :r'h~m:'!". <! ehil>dpo.:,}'~]li<!tf}" ~, fol~l>t:~Llreh~r 111 CohilllilbifL lJ Ilh/cr~i~>' \\;I~Q ~ ,m .~isled fDA iF! lis salety <JIl<lII)ISis. we 31retl1.cly !<!'

i
I

know liTmL "~'111~il~d cll1lpn;g:a.~ion I~whll!t ki.~is peop,k:," ru~ln~r I:inJ ,0I1,~i'S :!In! ~tlri;\~I1Ct:JI.Ili:n ~' :Ii! yo i.i't.h Hli~~J..: dlill~Lnow 'L::'iVii(jr!1 ill!'l-~l .. I .. .. ilt~3:;:e ilillhe U~ri,tedSI<llt.'S iI12flll}.j·and ~ W ~. !'l"lilg~~ing 49% il'l~r~;ISIil i!l1 l~~ N¢~lle'r-' :!il
~'91)PCJ1S\'!lhcilaillhj(!;I)rt"~ ~:tl '1:1.' ~~nl dmr; WI:te dl\:linti! .. "', ";UINN!FERCOI!J!lN 0

lmt'lio;-s!tlM' \'II'hm

'Hum;aIi1~,Nmt.M:ar:Uan~
ElrIJ"fillt ICilled "~;m Gl(lbalS!J!"!Io!!l{~f' I(M'GS) W@~lt silent I" st Nove roller dlrter il,rMn nlg the p la tl~ ~ for 10 yel!rs., no1 (rom 01d .age I:lLlllx'!::,m~~or a Mlft· t.1I<l1iE error, Pi NASA inveslig:a'live bil!ilrd repart has (om:luded thai the' m~ss:~tm IlG!l!!rallion:s
tJ:!'alJ1. s~nl.a ~ortwar~ IJ pt!alli:! 1tit1l!1th
5·1:1

If

A,war'$

~iO~1..ThE!
oil

-flaUij"~Glim~

rre~sN!ht(jrty d~tal "

d'ehil~~

~fore- ~o

(J'i!Q.ilh'~nllll:tl hQ[ ~iI vI'orefit'fiH~,latEd deafliii, iii! ,

E PI DE: M IIa L,OG'Y

~hl1!wrong pan oJ uhe spatCeJ:rart's .. H@r Sur\l'@'{or rec-eived a rouli

(omlJl1l11!~ m ~iIiIll)ny. 'Nr'lillsl[ rig hijvOC un I h~ Soi'i!I oe!!'.: f1Ir~

fliIH:O

ITlmand,
~y$-

Alii ,an!iqulll,~d ,ontmllltfd falllt.·lprote(l[Otl

IrBI,q M'O'rlBII'ii'ty S,t:udy' AlUlhio,r:s, Rel'ea:s'e Data, but Only 1:0 S,omle
TIle mltho~ of., '(l1IItro\'ersia~ S'llild}' 011 cflcnllictp:llat:~d (nl;adt.;illiniq] are ~~~.king tn diA;I~tl erhieism by 1,'ell!::lsil'l£, 1~I'.:itrr.l'W d.lIn. 131.11, ihe mm,'t; :I,l~""h.":9rJly ~tbl~t:l IItei:!~bilit:. l'lltiics, 'S!J!f the ;m~h(lli'$have \\iil~he Id key o!.ietl.ih;,needed In
tl-ie~'k lilt" ~~li1idy. Allid Sirni1~ !Iife oUl~g~ by I,hill cundiii,mt:; ~C'l, Jor w~o ean ha\ ~ the cl;Hlil. ~Ilclu(flmlgthe r~ll~f~ "ooj~t~vily:' The 1)"'I~er. pl'lbHshed ln rlw ["tn.'", :ll!i~l Ot"lorn,;r by ,'~US. ,md lmq i te ..~Y!t. tlst i~'f!i,n!l.~ th~l ()S5.0()(llltOn: P!;}!J<]'~ch:a..~ di~t1 [Iu.ilfl ~11\J:rm,~Uy\\uukll iIl,we since i~e MiilU'ch20n~ U.S.
umm"~i()n~r11Hl'f(!l~hm:l W

te'm 5U b~eCl;ufntly mhin:ter pre led lille- s~lua· ~iolll!,and'~thirl 2 hours Sl.Il'!l'eyor had di~!!i 01 ~nsumdem Ilauery re(in1lr~il1g.

li.~(llgH~r !NASA rni~i(ltl5exlefld~rar

been a p'n;l])(lfl,ell1it of the street-bias idea. Bmnha!lli, '~~.ll1illi1g th:nl,l Jl:lltdm+], d~d~1iI6d Ltl
co-author

explain ':<I.p~ah.:h ail.e-f.ia Spagat did not meet; Let> Roberts, IHr'!.'\o' H.i Cu~,1I1,nbhl

bey(md their Flromlseti ae-stg,llltfethm~., J'IiI.GS had ~uI(~red r-edll!ctiQf1!~ il's QI)er~li(;lns, in l:iudg ~tand $i<lffil1'g. ·Woe did n't ll mllliliit any det;~~· !liIir'edly ~.iluiSli!dlh~ ilfl~m,~ ly, ~ :Si!llid board chair l)o1iy P:erUFtS or Giodd<lKl51li3(.e
1Fl!'\lIht C!l'l1ier Inl GFll'e"iJ.!::U, Map/land. B!Jtshe sald, ~~t l:letl~ll(]ar ~D ~l'fpbark and see whal 's ri5h Irtim ,ag in 9 and dlilrMg,e5. in Ij)pli'ra~fon~ w might !J~ de-velopimg.That lessen i5 b!'!il19 appl~E!d1.0 all MillIS mvss;iillnsas well as an1'5 !ar,;.imJ ~l1tebelte. p~Tt (I~ a ~e(.ilde lo I'BQch largel$ !;Ud1 a~Merru I')' 1IIno PILI to. -IRI:CHAIRID lit!ERR #I.,

llJ:Ili"'l!!f's'~ty, says he w.<I~n't irwo].'lfedil'l ~he ~!~i~ioti but thl'll Sp!lW'1 ""\H1UMIUlI. qntlell,he I)r~il!'r.i,~~' I1nlllil~it!' :1n~".l'I~'tIi:e b s ,"

i;)Mifll'I~.

~imm" mb)1 tllr~~i~1I The mL~J:!I:m, !1.:tl~~hi~resu IIby 1[.'X~~p-

SPIIS'-'H c.!II~ rhe policy "dl!--er~y Ilnwed," ;}tldi I~I!-."~J we dl~ :iO,Lnc~hi dlm~h tlr nonng o[bjec ri ~'" w ith ~he' U,Il<l. q[ t!.0I1 ined Pt:lOp~ ie sl~cm~Ji:M.:' t.lbk lu c:>:~o.'il;! uHr Si.UipIdIHy."l~~ l decv!1ion <lisa 1~W';;de!'D;_wid l(,mc.:II reU~'lw :H the Hmtv:l.rd h:t.stUunc fur QWl,anuha'livc
S('H.:i:,.:I. eieneewha S ~..!> rlIc-eivc(.i the d~rt.. :;.t.1
C\'1:n tho,ugh .&It: s.ayli ltL:prn>lcd ~mllliim::IiltS on a \!Veb log lnst 1;!llllim~F'.-I~seli the ,,~s!1ibility f fraud. IJ ~n)'.vl'lll .'Otnlll,1' cri~ic~ acc!lJss " is

:SenSOf' MOVie, lilJleellDed S,ensiilde!

INASI1! a r!id till!! Natf©I1i11 OCeClJ1Ikilnd :A1mos· pheric Adn1fnfsIr1litit:ml~5lwe!;!&:: fl!'sl:orfda k:@y erllli rOOP1'E!Ill<!lsensor t(l al kIJl!l-~~ It'l!d 5;lI~@llitl@ (iel1tlCmstrililiOIl nli~iorl due 'to be ~lIum:he([' 1m i!00 \)" But resea rrhers are 'givir»g tilt' II'IO'I'''€ ol'lty IIJrteothUll11:1 up; lhe a~eli!d-es- h~~.n/t d !!(id\t;!d
rMlltth-er [rill re:sWTe Ihe 'Sensor; «!~led lh~ Orone: IMI<'1;1~[tl!J Pr(l:fill~rSuil~' Lilillb tOflltPS·Urob). and
~CIslx p-la nned So1!.elnle:o; that malkie' lip 1l1f trGUb,tKl ilJ"lit;l~a1IPtlL~r-O~iting OIP!lI,lltiQlIliil! EnviJa.mllellta" Sa~ellite :S~:;tertl (Mffi:ESS~. OMPS-Limbo, t,!Illr,h \',1m IlrQvrd~ O:~iIllloed iSlrib ution ,~ala lo:r envi,rO!t1mefital and di I'lIH!te :studi'e~. W~ ~1Il~,ki1d oJHh~ N~OESS d EnHJ and Ihernaf rl sa~e~jlite~ S il",-e 9IOJI;-ey to ~dem;e, 161une 2006, p. 1580t IBut in i1I

ul;".tiflgl!h:mm l'Ilolim.~ity dan. e~lllt:c~ctl thruugh drlo!.l~li!N:hwr sLlI'vey=>. Ol'l.le'r academies b,nn:

(lllCst;OJ!lOO,a.~pL"Cb;;OnULO ~Iud}':.l~"{ll1l\ll'hell1(Ot ~he·~!lti;lf'\r~e>\,'S t.:(Jlddh"",'~ b~n dt."]n~.1-." qllidily

LISC laimedto

(I

r~.dj(;!I!~mus." K,mea~ld!o;,
OllC tl:!'udJ.::.l!llio:togi!!>\ apan

whether [hI.::IXS!!l ~I.~,,,\i.;ill: .lll:naH.'tI by surveying only households near main streets \'ul ue ruhle to bum DiS ,l nd stitnmi n.1:l ~ lSt'i{~ffC1). 20 OctCJber 2006.. p, 3%J.

asr-.;:es t~at l1u: condit ions

<!J1'e

Ilfm11l [~1I::ftliY unusual: "l am

\\,ary ()t'lryillg to liruit access haseel ml the predi lect ions u r these requesu ng le," Sit}! s

Lent! ruuhar fiilben

B~lrllhaml's

team

1U

Johns Hopkill.s University i:ll B;:]himol"e. M"r]ihu~dn<!d [e_ i~wd (:;dt~ ~o:mlc,;m_~~ .. Ih~rmlll'

D1bVid Snvit:z: of Momtl SUll1ui SC'hool uf Mcd ielne 'ill'! New Yor!.;: City, Btu /-\,Ilen
\Vik:o_'i.. I;:di~!!'I[·iIN::hi~r ofE'pich~mioh)f:.Y.

dmbl, c:iti~l~ P05S1 b.l.c:d3tlg'Cf 1.0 ~hC' tmqi im~rviewers tlnd~he ~lLV'n:y Ilanicipm.us-, Eadie'r ~bls Illomh.. ho·vre\'el'.BLlmhrut'l and his I,c.nn P():>t~. "I note OIl! lh~,ir Web $.Ite :';11)' ~!lg I:h~' wouldl ~.'de..iSI: ~~ Jmlt sct ~\ripP'Ud of inro~" tiolli tllJlmigh~ re\'e;)I!id\ent~!ies blU (l1I1i~Y to q[l;,lI~f!C;':d~![ie:~lli gnl~Ups., Sl,Idll gfL1up~ln~;:;l t~i;:'

Jcfend:l> ~h~' c-on.dhi'!Jns seilly ihc- Mop,ldns gU'{lUp: "'I Ctnl ~aHny blmne hhem]t1tlJ' behl~ cau~n~ms in tbis case"" s<IYs Wilco1'l., beclmse til Ii.:: t'ClpU; i~:;;~) pol.hk.tI ~yi;; l1'l!lle,~,
Od~~f.ii lire
(;u,i1'l:l.:rncd

'tinl:1,~ lilt: g.,ro'Up'·~

decision

wwilhi1tQ.ld

inlhmuuiQ:n 5u'ch as

1T!1a~n~lre1ti n;lItL~=,. and ih~';s'~lmf1lin_g rro'(:J:l<.

imr(,)s5,ih!~ W dete'C~ -"~!ree~ h",\\e~\rer;!i!-iei!lJ him'i~~tij~1i!l;~ ~!1'~ el)~dfinio~,. co~ h~!'i ma,ge i~ b~!is. 0.1' othc[POi.~[ltitl.l p[l[}bl~nl'~'" More ogy, i:l'i~ tim~ ~l]h"" and mti:;~ ~Lb;ob~ "withui:lI det<lils nn the inler\,'ie\'vs "'-Ire necess<ilU'), iF p prill] idy st[ll:ed views th<'ll "'"'{Ill ~t1ctlm::e dmdjo~ lhl1:' O!!-u~hor.~ Ilfe In hry ~o r~sl !.I'J,lim.,.:tiohul' ~ ~Ibmll~ilmi:r nbj~ t:ivity il1!, ;ln~dy;;::inM th~ dnt'L" ·H111.n.ic~~cd' J~U,!I:'" S,1l),'5Mlldi!.llYi11 Hich. H !l! T~.c I-Iul)ldt'l~ loe~u:ll SH)-'S ~~\'!:.I'I'i.1 p~ha\'~ ~rutl p:sy~hdaJU'i~t !illid Pll,blic ht'~ihl!t [tl-l>11:3rullh1:r m fi::ccivt:oJ llilt: daln. i But .n ~eaS-t~:I'Ie ~,,~reher h~ bt.~t Lllnl~,(~ K~n!:l't:('o~h,:J::1! londoll, Bumllu'llilil S~~'S, his I;lfUUIl "~f1~'i!'~o~}[:;;] ~l(l.uddlil~o~~ullt~iel!l!>!(! dm,'it1l1: Much~~1 SpI'lg.n, 1111 (mnu!lili~t wild ~ m~ui.:rials,." _;mO'CE:w.YN KA!iSElriI. E:: c:xpcr~ 01'1. 'm1lliel sllldiie~ !II. Rl'l)'id H('lII!O\''f<~Y, t ~ Uai''''eL'Si~~i (~r ullid,on. il'l Egharll. who h,~~ W11l11i L rejll:lrtin9 :by 10111'1 l3oM;m nJ:lI!i,

~~n:h I~t~!'!rKi lil~ Wlllit~ Hoo~~,House S(ieJ1!~ and r~chll'iltogy CommiHee!' Ileo1d~~hlp painted out Ihiil ~.Ii:f5el1~Df for Ill£! dej;Jio illca{['.. ~F~aldy bE!@1lbuHl and that i! wouldn'[ CO~l .any IfIIO~~

~(')fly il. Or'j 'th~

d'llll'!l).

J!lel'll}o~.8'-~E!!'Isi l!»li[)l!.n Beril'l~n Moore 01 Ilgi t!1e LI nfrv!!['S it)' 1;11 Me~... Ha III'lP!lIl~fif', 0 Ulrliia m, aw~a:[HI!i!d Ll1'I1re~lor<lltion ,01 Oi!JPS-UmMnll

i i

also

W,iiJfl~~ Or! it

Ilui! NPOESSUitj hts

~.c!$ ~!1

op E!J,lIliilF1al ~et'lso.r." ,1\ Hou~ Scie'IilCoe .il.lTtlflil· C

!e~ st~nl;!f.s.a~S

or

meml~[~ \\!ill p'liIsh~oma~ lha~.happen.

~on'lil'!ue

IheLr

Klll~T

5:VOIB,O D,~ KINIE.W fll1!:OM lli)'J!IYONE Ihm ~.cijence meaut ~acr,jiees: 'P uIli.l,g I~lng . f
hours,
S\'!,"lJ!ltvng OViI:$

r.iil.e d;iUI, und suuving

his Ile~L'M,n,!I!if!:! f~lrrr\()f"e~si[')n:11 ~llc(::e:;;!;:.

\\{i'n;H the you.ng assjsram professor ~u Louisiana Selle Uaivcrsily (LSU) in B~II()n Rc~)lL!J,e idn 'I rea 11ze \v;:J~ lhm Iirue sp"n~ un d grflill pro posa Is. nm :g roll ndb 1JlC1~:k~~][!: exper~ mcllt~,. W()U h:IlJe::

Ih:l: b~g stressor. madel oncJI~,1

'WlIS.

Ih~y hoped wt!,ultl sw~~ thu re~'ic\wrs" The ri rn:l~'version the uppli'!;;<'I:tion com;) incd '11 d'llj r~!:l~ln:,s. S\.()b{!d~i:-;ay~,. "I've l!1(lV~r ~t"'el!u .~l':;.a:n w(lrk c'il':llmrd in n~y .. hr~."y~ ~ Svebodas LS lI' cO].IC~Lgli.C., n~llmb~.o.iug ist John Caprio. Fr nrn ['n I i rOLl] ia w Ln u ~.:.; i.1U'H1l 'W M 8 ::;;,Saehuset ~'S" tho u sa un 0 r b iemed lea ~ ds researchers ~'~ndthemselves in a ~.inmi~.;lf

or

N~,doM ~[rnsl imtl: of A~I,t:l"g}f ~Iu:llln r~cli,LJ'Lls D~~c'<lses ~NVUDl, !lillieof the largest I['Ir Nnr!!; 17 in1O~i.llJte:s; ;lnrt.l (;CI1W~S, Thill perC'~UlIWg~ of research

p'r(llms,~I~ funded by

NU-t 11[11; dropped from 32% ]I~':;:Uo ~ ttl, a p rojectied: 21 % tIl is year. F undedgrants are
rulu:indy

Svoboda, who ~urnedcJn LnJanmlrY. \1I'01m~ to '-;,I.udy h£llw nJi(:ot irle;:: ." .. t:;; br;u in

d.c'llelapm.":mtim.'leI:Mi!11i~h, as
lexpo:mre in women
ll:lIlcy. ,,'1'110

n'~c

<:l.

smoke during pllegblll he h·n~f et to y

ij,

Ii~[Ie:mm:e than

Last surumer, his ~en.urc rev lew

,I )'l.mr'i\qy.

l"inaneilll bind. This !I'C' ar will be the feurrh iIli ~~mw lh ,It the hu dge~ n f NIH. tlre wealthiest research age He y iIII the 'wolr:i<i •. has not (~i:iiiQ!Ilf Iil\!!ng!iloo;i!jj ke pi 1)3 C e \\dt h b i0I1]C d:iO<1l.J ~thlt~!lJI: t!y; J~lrl.~tio~. The ,,,IQwdHWJ1 j:j
ofl~ of I:~h:lIlJil.gl.:~iyt:iil"r~'Ii'l

,~ six N~H ~.I1Miuw::dirce- '.~ ... tors. and 'Lgenl.:y illeadF] las ZerhouUll L ~
Sd('il,(,,[~~lungwi'[h ...

Do

em by 10% Or ml'mre_ zens 0 iIn.yestig:nms :i ntert icwc:dby

€ '"

de;_~iClr~hi;: cl ~m .. ~~ in
YCllln!:, scientists

,I

\\,h ich

SLI"lLg1J,;i.t! 0 1

laul1Iclllh,eir careers and even :eJ Ihe most s:enlor are ~Iii mi 1'1£ ; m rri.;;:e winner led NIH;

I
:~

$C'(;,IKC IlIlrl UI1!:;,pokl:::liI PI\;n.;"LII~li:;lit'l::

rcdcu.ill

~h~Lr n:.~!!:~fch pr ojects , ". H3:r~}ld Va:l:nHl,:l;,11 Nnbel ~.~.'

\v'hQ

gram. 1!\v;JSIii 'l~O:r lack (lftrylflg. Slj mee 181ld· iris, his job ill lSU iUD2UU.2. Svobcda has
been
1;l1iJ :l

ccrninaous

h\ln~WL'lr money, five

lime:;; :>L1bmbuing (I[r,,:~ubmniu~ng m.::Ijo:r .iJJrl).~ir;:atians ..OI~ongwil,h 1\\'0 smaller ones. L:I:ro:;t. S;U~IlI1l.1CT. he W,1L'l if;1Q\1,r11 to h i~.I:lrni.rd ;:i:nd f~n,d c hanee Ihr <.'Ippm\'<ll of a rewi sed RO I.
thu' N alim'i11~ I [blj,~ utes
(l

2:006; To make marters worse. there m,I}' not be much syrnpHthy~or t.1m~~~ :jCiCi1IlLsH 011 the outs idle". nuonle~1id UlIei s t~OI]~id~ted dell SJn(1 \'1111;: U-P!'OII;; eted terrilQVY, lndeed, tooO'!y'~ craneh comes after ill h~aJy SYlm:r:>of grt)wlill, ... '.IlL:N m-I. blLd~l ~ deubled b~ull,eelm 1.991':, nt:l.2003, BLU pen~

scient ists can reca II!,

r I-ll"aJil h l; (N
OU!i~

rBi>, i

in'C<ldi-.!md-,bulll!l' ~T<lI~n, Wilham

S",'ibOJ3

~~~~pie'~!t;d \vould I{l~ h~sj(l,b. h~ HI;! and his i~IU g,radu3i.c' students \Vol'k!L'l1i i 2.~.lull!lrdajl'~ t"t;lni;.:in.g OI,H iBm:1

ph;! O~l~lte il'lsidek~lQ\\1 ~heJnCi!'l~Y eruueh it;, real, "\"0111"1";': II hVl;OS L ill a d ~ rent 1TI10V' Ie:ITc 11Im,v;' ~~iyS AlUhc!l~yFaud. W!llJ! heaUls the

frol'lill ~t)~'3 to .~ 9\19 and is :! !ilQW ~~e<id the Mern(lr~;d (If ~. S l(Hn ~K.~u~dri.~ C aneer :~ .ll: ." Center in New York (']ty. 'tl I~ h~LS 11wid. his ~r.;;~tlt, euj, ~~llhou~61, he ''i.'VOn~~i~ S<lJY by how rnm:h. ~ Sl~~ Lvdtl'i ~tb~dJ!:!t'll(l~ !~liil1g$]9.,.2 bm imj in ,20m, Nm is !n,m:!!y ~i p~,lIll't:I:,WhM i brmlgIn bim~u.lclil;"'l~ I'I;l!-;earch 1(1 ~11isrl..~ ~ c e flln:ailid~ll'Ln_:i:.i\:'lt! lhc dou bl!iI~~, .I:1w.Jti:L:d :~ NU-I w.utl"il!billiuns ~'itUf~t!tlil ..rrl>. ver :.i rela- ~ o

:1

:356

NEWSI'OCIJS
,....1 Ri,S~n~ ~~I)e('I:~tiOln~. ,lik! ~'~,,~. tine ""'" Wli5rnn~m. Mildl50rli, I~ ~1l~M~mglm the
U,ni'i'~fSqty of binsrlences.

ri.v~l)' brief'time, \Vhe~1;l"~ p.Di'll'I!;e I,:orpor;a" ,I E.i!l.1'n J:ll~~;hD conserve SO.UDll;; nfl.l is, ~"".immIJl o by ia~vN m fillUM spmtd IIlL:u:rly alllt~.'ll monu.b·Y

Sochotdrs; lsired !lie:"" faculty nlember~ to Hll the bllil,dln~, i!:>:,~,e"'lin~ to recoup 11'n,ei'r' iIfW~:.tmc:nb; f'rani! the NIH 1:rr.mlll inw~lig:n.tors wm,d d hall I il:l,. "Un iversi ties and L I1dr le~,d,efllhip did \~'h:ll ~.'WfH~ have Id
s:~~yl!>ZmhmJ.ui. "Thegevcrn
don:e~q)Ll."

u pward, i'H H's re sou

A n~,u. ()"'I,. increase in 20ft;;, tilt: fi rm~)lear ~ fJf'thc dOl!b~i~l1:J"NiH r~":I.;'in'~~ Ol)%lbom>t in
:WU-I.
311

fCC S

begn n

U"I

d..u p., r

abrupt

reversal

Ahhou£!.h lhe g~llew.d

it receives
voked

i'll.,e}'C'IlU" it receives

.ilt.

Thut 1)1:0-

:. m1lss,i \,~ ex pam.]r:m in binme die ..~

resc3r,cl~.and ":;1(pccultiom; of fe-denl1 sup· port surged ~o 3 level thm could not be SlIIS"
~8iil'Dcdwhen the b\ldg~1 s~opped l1Nowu,lIlg.

The crush b h~niJllg Inhs. careers, and the pc5ydl...:;s nf sdeuu HMs, with fl V~lIlgcflnct':. 'Tillie ~Ii'!illru,!llru,liI'~~ NiJl~ ye1llf.':; :ago, Congress set lilli, I,t) JOl~b~!I;: the N 111budgetand w i~11inS y>e<1l[,ll senl iI ~oiln'i.riIg from S! 3.1 h~Ui01:1tn $2:1. ~ hill iou, Bul, everynae [knew I~'a,eoltlt!1il ,kL}'S would g ~ol~<I:.'iL. n Ckloi:Jer 1000. e~gt!l ~eniHf I sci"
;md polieyrnakers began raeeung I)' di~~Wi!:iS ho~v [0 ilI'i:! iUil13~u thoi.': lfnuln~n;lUl1il. In 2U02. Ih!!gruL1ll I~LIIlI~~hctl ;<j cum'!n"mliiiwy in Sci!.w(~(t.pwe~:nti!1g dvff~rer!l bHdg~I.L'nod(;:1 s and their impuers UI1 reStlOIfCh pl'i[loriti~~ ,~SC1·t'fll'd'. 24 IVIOlY 1:mJ2. entists
i nfOffL~,~L.~ to

l. tli .1006, wa~ rt~l: isin~iC::~lIII ILI.S, [)el.~,unn~.e"'t.1'tTol1m~er«, ~ll~ ~"Osa I {II" i!l~ MIPlm.rl for th,!;lstl ~Lc~ivili1l:!;;.'·and lh~ IJX pa nsi o.[iI W,}S, "cxIlCily ~v.I:1m. gouds~md services ~n biomedical research and develC OI~gres$imc'ntied" Op!JJI,t'I1't r{lt;t: 4_,%,. Tilt: I1n.l.llill~ TIl [:i ii P'P(:';'H~ u) h,we r4Uiml!!tYo~ oDppli;(;lIiDI"~ ber of com pet ing gants Nil i hel red ddve 11I10<rc I:nPP ~icants, Ipa ~~lI~lIRli: r~1I11{1t: I r~~,lo:.edl I'll 100 ( i 03 all d to NnL 1 It T998., f~'!,:\'Crlh3 III l.B has becl'ldmpping since, The 20.UOO scieruisrs sought de-dining VH~~t~ CltNll~>:;. dnlre:>IHL n;h g rants fnl m dl e 1.. 6 ~ 3g4.:iIlCY~ 20(J(). that uumber in hm. and I:.islng. demandwere
W'1:t;

rule of inflatiou 3 _l %.!Lt:cord i..g ~.O Ull: UiI

or

rOI"~

une.

mare than 3 ~,OQO, and

3ceol'd~ng NJ 1'04:1: Hjorecasts, tl1(;: 'llLUillb~r ()fu.p~'llk,uJ,tl; ~s ~;o,;fleCI1,.'>d W~(lp J 5.'U{~O,~I:I2() 07 . 1.1.1..:- llI~iflm u r ap p~ ku'ls ber kill has g!rpwnm an even ftlsler d ~I'I.;11$ sci e nti~(s,~mll~~rn.~dI ::Lboll,lt their dla1JlCC ·of g~[t ing ~\IIlJ(!~o. ~llbrl"]Ji ~ti!:m P!:P1J5,,,1 s !liID'rfl;: 0

fr,.;qu~liltly.

'l~

Bio!caJu~t:,

growth

I.'I.t IJ1Ilio!(li,t:1I1

schools 1<I:ygedsemewhat beh'ITld t~e doeM 111g. ~n!lllly insti luHm1J~ ZLre sti U ~",,,i:u1t~in!!\, At Slu{'j~-K~u~,illi!:l, for example, I:lflf,i,dl1il:;;
(Jilll)' i'~~C'~HI)' b~!,j11~1 !1mlllg
~L

P'"

.I,

40

I).

[[I>

most pC'~:tdm is ue p red ie I1.011

medeled annual im.we"$e.~ (If·4'!,~,. s.<l}'~ VI' il:h sei ent b~~s. hey ,el'i]'I;,'e~,[0 increase IIlI,ei'r T David Knrn, :.1 farmt:r S[.[unrord UniVl~rl!ii~y r"c!IIltyby ahnost 500/11<. ~ay~ 'VUrmtl."dean now at the A.S:S\:lC'i~tiQnor American But as re~llil::sl::;;for NIH I:I.lm.lq! t'li!')cd

new

bli~ldili:l!!;

"~~perfet:1 double \'Ilh.nnmy." says Zerlw ~1'Il i" Vel the ~umbcr:;, f'1il to I;;O~l\"~y the gmllw:ing II.Hl...:m:{: l!Lll.al ibrcbodil'i!'; expressed by sci eurisrs aC!:Q'Ss dhcil'li ines and <It every ;;t[lge' of d:',1,dt careers, ''TfuJ~ rip'plc e fff:cg h Iilf'~ is ;;!m~·r,; ~~ g ;[I!l dlfl'~~'"lyzing." S1.YS. Stl.c!\'~nDowdy. a c;l~lt:Ct bio]ogb:t M the Unilversity oWCaUomi<l. Sail Diego, t\.t 0l'aw~ Un:~V~~ily. ri'i!('JoI~clthr cl:;iI ~i.ol,osist Susan G~rbL who hd1~.~ eversee !Ar~l.JlI!lh: I.[~illling, c~Lr~V1Is:>~{19 Filt'ull)! it'lc-mlE[:I, in 4 dS ht dep artme nl s reee nt Iy. as sh I! do es ~\'ewy Y't;l~:r" m !iie",hm:v In,iny wtadd t,~kc on a gradu:He smdcl1i! Wro'm 1l,";lo!.1 Ic~lr·:. pack, ~

:vh:dic:,al ("oll~gl::S; (AAMC) ill W9~him!,ltml. D.C,. who he~ped l~ring.the grnL!p I.{lge'ther: ·,\v..:; d~dn'[ model Increases lH;!.oW 01:% OJ

'Gr,lIiltAp~~i'(~ml:itQ'Ni Ii, :fit ·'Pre,t!'i~

s 9~2007"

ye~]r because the tl'adleof'fis a~d the sacri [ices, '~h,nwnliid have 'o!;?em caLl~ed _"_ were 1m) d iIfj eult for usto deal \\1 ilm injhe model," Al NU~l senior o~fi,ci~I,ls. ft]ltlnd tI'llIl ·'riI.f.l ummel' w.l~J.L t hc'rc w ill b~ 1)8~11 <tlkr t h.c

"I n the pasr, i~ w~~"J rIiI.,jor~l~'." arcund O IlD% ~)f~l~os(!' w'l~o respnuded, sl~t" S<I.Ys" "'fbL~ ye<!r. (l!l~Y"bo!ll25% on,he Ir..iners said !hq' 'would be' interested ... because they did rWlhEl~ie ;1!£,l!;Jr<IIIHee'of rlmd~U1S 'hrn~M f:lW' <·W~nrt"s chi.II.i.llg" abnu~ Ihc I.!.nllllght .~~ L~mt'\\Al:'r,~ gcHill.g ir.no years, 3 ;;md 4 WW.l

{hH!hl ing.'" ll<lYS ZerhOIIHl!i. who biftc~~me N1H Jir(;C'[(lr ill :mo2, To sullen. 'I~C blow, ill 2002 ,mcl2QOJ. NIH tri~J 1(1 'lc·('i'l'il1lralie du~ p.we or olte-t imc oCNPC'lild im:lX:'ssl~h m. 'con, Sln~ct ion. lO rr« up mom::}' for ~he ({ll h:n,v"
i Ilg, yt':~~r.l\i.Hul c,"'en "'Ilm I:!JC\\·tm;l ::;.'Ccn<lrio,:-;,

end in sight." says I(j\\~Jro Benz Jr.. !.hI;! pr~~id,~ni. mni ['EO ~)rDmll!~E[ltib~r C,mc:~r h~sl.itule il,' Boswil. Many ,rc'sc3.t'\chcf:;
1'10 8J!IOO l1otU.~t1 l~l<I~ llilc pn;~~m~s;
I~ ~ I~ 1- 1-

un I.h~ fed~ril~

~ooo

?cl,')'ple reilUy dItln"1 lhL~k th3~, Ih'~ Nm w!lJLudl(]l,:lO below 1f:I,11atU)il,. says ! " .ze'rl~'(Jtwi. .an flutOOrl!1lle he .ntrii:lutes lfl the oL)il I mmd.~. LhL: iltS in Af£lmlili~t.aIl:l ;Ili1iill W ! ~r3'1. <md I h~lTi!.:;m!;! Katl'ina. ~ ~Mea~.wl1!i.~e, n;,"~e,lr-ch ins;ljtutiGf!~ every· ~ \\'here\!r:el,"!! breakj IlIg gl1(11L1nd ne,"" ir:ldl~Ies ';:)ll i ~ ~lmd~p.i.l'ldi Il,g th~i'r j~l:L:~l 11 1U.J!OH2.~lLr~'I::}'. i!)'.~ .A AMC fOll lilt! th'l!ll !l!::\!,' co 1liS,III'lLe[ iorutt ~n~J"
i~

budget. il~clml:ing tbe W3r illl b'lq. ~e'~",e Cu:n!l:r~~s~inJ~ '['00111 to ~xp;;Jn~i m:"e\'~n sl:abi Hzc mh(;r pmg.rams.
(l'iI!Il1IIche:d '~!lllt,:ash

~ bLlJg,cl;

i?emmttlg:e O'T P:r(lpos~lls. :Fulrd~. 199:8-2007"'

'~lI!iIImd 1I~ 1-

_.
-

I\Jan ScliHaeycr.

OJ

5:l-yea.~oM ,reprodul,ive

eUi!dol,;rin{l~9giii't. "i~O h<!!.'1 ~SFl~!Hlilis ~~reef

1-

'i

.it

i~<ll SChi.l0:1~ hil'cl~,"'rlodli;ld: iFl"{]~l

~dtL1'O.l ~ ~r'I\.'c~k~J

I, ~}9n til

II-

-1-

1itMjfi,~:~i1ch.us~u:;;G~n'~fl:d H()spihd in Boston. <It nl"~l w,nm't too cOllocenH:d

,vhc'JI hi$
~
II-

gr'illlt

<lPlllh;al,~on F;lned

hl m~Ik!~

r=
I-

; 1997.

:52.2 bllll:io~ lin iii 1.-'\\.'

f-

lheCL!t Uke S\'obm~" ,Hld ,!II &ci~!'l~is~5 [;}p'pIY~I~g R[lI.s. St"IIHh1:YCL il'> ~llh)\:vcdttl fo.1" s;.ullim~t t'W(l ~1!!'\Iision (lj' .m ap'p~tc'll.~oDlin
hLlP(;." sol'

~ 1[,'(']n.,"u·ut;lit111.;0E11pllr~dI ~I:I $oJ_go biHiolllWoi'!l t :g '[ i);()R It! .2 002 .. B Ln, t ~3t I'J:I,h,:J iUD'!:OI'I'IJ1tlJ:if.>O.1l

peC't.ed 57.4 lli'l" 1:: liul1i ~n lfhl\,!/ COl:L~lrucdo:tl It!Jm .200210 .::W01. ~ ~ A.ANiC 1'13~,1II1[l~ et cmln:raru:.:J w!l,elh,;::r tl~es(! y ~,~p III.ns ~""I!rec [JJrl'i.~d ,()ut _.
~:(I Wh:~l~">;I!oi

:e ~

ee-

lO i"."OIlfIIC::nl ex

II-

,~

('Ier~lJ:ol.~~.i;n viewe j;l~ run-dab] c ~C{l reo Sclm~)!l:I"S

!:If

r:.; I.n gi ve ,~

1=
I-

1i_

,-

1'=

1=

r'L!c'-~ru~)' bl.lbi:ll ~a'I UIIH~:-;.p~CI£'J BIII'.n.

dCC'<Idk!'

~u!ldyilil~.

w(ukh~d Alh:t~~ how I,\'Y'O pw~ehils

..rTI;lC'~1'f."IHOo!;hu;:lion. h~h~d •. em In ~ni!Cc anlli hit upon. h

t;'lin1in'L~l.ld
II

sllrprisin~

\l'!llAl'w.sden.cema 9 mq

5C1I:EN:OE VOl316

20 APR Il2007

NEWSIFOCllS
'Sul!J.miuing grant aplplicatCom, I.aK~511i1®re'amlliJ1or~ iinne, ~a~sMu ro!iiologil~~ K~ i1 S~oIl;O!I!Ia"
'O'in UUl: ih!!!Ilt"

N:niona~ r!ye lnstlture .:' ~ fee~m.;e I'm

doin~ alot (lfsocia~ work." Comp-olllllill.i nIl, the ,pfabll;.llliil ~ ~h,1l n~!CI~[ ... u nivcr.sirics ,[fII dI11l1cdit'aJ. in!.~L~U,H.~OI1S rdy on

Anc'r J year)';
;-'UU l:lrn.l1its renew a
{]Jl

nf IfY~llg

fur

NIH

I'lHlIl'ey

for the bu ~k of scientists.'

3(;1<1

railing.
RO.I.

inelud-

iQllil~\H' 'L1nS!;l!~!1,;~~~ I 'LI~en~pt S 1:0 Ih l>'(lijOJ1,~ Sc.!11.1t:'YC["i>

ovt::rl'lt"<'I.ti costs and IIO:CDH.'Il Stl'l up 't() Sl~rpon 1filC!lh]~ nefuber:;lonJ;l,~t~.rm., ~ Tmditi.ullaJly. "il:rktl.!:lt: fmuJillg" cOM~.Jli!.k

lOU h:lII:iC!'s II mi.

lah hns swr~nk from si;;opeop~e

to

researchers

over for

<!

few m onths_ Btl l I1:0'I.'~'.

e --Ili,l:' works there hy h~msel r AItIlH'Iu,gh th~ N<!l:i(m!ll Institute 0.1' DI.3.b~K:!l11111.d Di!:!:l:sliil'c aad KMrH:Y D'D seases m~y ye. m~ ke ~1U1exeeprion ,1r1d 1~lItd hiIH.he':s mal t,f~['dng. any chances. Next ffillol:lJt h, Schneyer \\i in lJC£i n a job< [b.n dOCS,n'I, N~Y as 'h<lvUy (lUI NIH n~mll;y, wm"dn~ ar Pinneer V3~I.e}' Life Sciences hl:ltituu~ ha

amre scieuris.ts lhml ever au: bavillg to resubmi,tgl'3rll ilpp I icm. ions, with a ~ap 0 If 1'lillOfi.lht; UII mere in between each 1iubnm:i.s· sion, AI N I A,l, I), the pel\.jen:tage of proposcls fimd~cl, {)l1dw tir.:H IJy hi~S ~ofle f['![Nl112"i'%in

5pri ng.uei dMassachusetrs, and ",mnn,utilllg rnn'l.l BOS·tOll J hours

~~W~ d",y., "It \ .... ,~!)'

2llij 1: ~a 11% iUl 21]U6, SO me schouls are b'eefing up the if bridge Ihnd:il1lg, DlIml-,ra~'l:l'eI·. f(lr exarnph:. i~ ~tting ,,~~id'l: $,] IInimm~W $,4 millhml! tt!is )1~;)I r. Hi SI ~)rlea lIy. I he ~ 11:i,l ~t e reserved m $sm),.ooo!O $1'Hil~im.l ";!;ml.<I!im('ls.t never
~p~nt

i~," 53}":;;

,B~IJJ7..

Daml-:r:~wbl:."(~ ("EO_

f,','1;11"1 "lhJl;: S~!li(l!, ~ n\f~S~

i,.
fi

rt.:1II1 i.llnaiytltn won't ~e'l <I, gran L eve 11 with~uti i.d~;1;I~d

gat.or is tmllill.g alit to be

Oh'llll':l liil bDl~rll~llIr!~ 'am

!he! tntll\,

(;11 Uea,geFor n

1:1 S

to

SL~

PllorL"

ill :2006;

r~~uI t.~Tht!

.mi~n~i~[;. l]~~d ~.upe·rior g 1i;!!L:11i ~~

20 ye;I!~~ r ex per.ieI1C>e,md 0

O!

abundsnee of panereai it' ee UlO that make I nswlli,l1.. "l r )'mi 'n:~'tryi~'1g W treat oJJ i•• betes, lh is \II/>I,~,tIII~Ibe ~'c I:fue t." ~.~ys S C'hl]l;Iy~rSe hirll::yC'r says he went dU!J,ugtn 3. ·'SC\·~ eral·ye;lrs~e'ilm ins C'LlIfVi;" mm ~ilLlluinginw Ehe d~nlx:'lt'S field, :~nMarcil, he'Icsrnedlh~l. o:nh~:-; ~hin;l =-!1(j .~a~ cJml1r.~e. hiLl'd Ini\~~4 !lli~ Yil!in"·s fund.i iilg c [noff by 3o/o!!.

I.fl~e I"mlCC II nd an

~i~,$t'~~ffile'

litim~~·t1"h31.~,j;Jtlic1iil~:;; U~~LI.~ l

~p,plli(;1!'!t

Ilnent for ,di<ubc~es::' he ~tly:;. NU:il nWficiuJs ~;,!y ~lwy'rL;' ,1'l~aril1l: n'om mUII'I)' ~c~c:lnu~t~

Establff,shec.l in",ec~llgll't(\r

wh o.nke-

SdulJl.:l),c.r .• ~~,~a'l·I.

B!;l!l Z conrj.nue s. Fu nded grtl~Hs, 11i1fli: !>ubj!!~1 iO t:uts2-'1% en av~IiHg~ 1il Ih~ Natlollal, Cancer lnstlnrte ~NC] )" W(1i r i;!~'i;,I~nr <ll'ld le. 18'}l~iH the NmhllDlIi.l. lnsu.,me 011 A::,:i ng, I fI each ~ll'lh!:
:;:::!:t~

'keep their h.bs rUlndl~g,."I get I)intllte c;!III~ r!'nnlp~~CI'pl!,l-""yil'l!,l.. 'I'm I~lti~lg nliy p.l\Opl~ go. what lIo t do UlOW'!" ~ says 1\,!~ichaul

l<l!sl. )I,e-a'rs. 2

Benz. two or I,h,'ee D,ul1a~

Oberderfer, :" P'I"o_gr,UiIl director

III

the

Fi;lrb~v 100h:;;l'u!i\.'e ftl~ ..uMii lh~n~$e]\'esh~rl· dreds oflhoiI.Js~md£. of'tlcllats Shur.l.ufwhat ~he'Y s~y .hey need ~o keer'running

\!II'i~ll'IJIllm~IiIi,'(.. 19ralnl ilPp~i(iiitt[I!IS at an all-time nigh,. (~lIiIil~tiliom IS p UI~ti il Sll'illl11 ~I'I!!sysoom ~B1ii!t I'!g on [lidS lih@ wirmers. Arter 'Simrin~ 'Inl receflts~w:l~lelliO,i1iS--<!5judgfng Iil<In~ts known aillile tNiillt09alln,,~· ar~ lLl~es o~ H e<alth (11411 to-many If"e5;ea~(hers oes.cril:ie the suess ilnd ~rU"ilr,[ltr Il'ri Qr hafl.l'tUig ~® malk.e i;lrb<Hr.:lJ!f d~5Hrl(t[(ln5.. ~'rt'tI u 're !\In I hI!' ~irp~~1lI~ f~~iFig h01m,~ thil'l ki rig '\tI1e werl' really rUpPlng (ooi F1S011 dlose last three thO!t~h el!ll'S oil lal at st1l ~li! h ere,"' :iay~ Mich\lel grallts; ~ s~,ys lII'Ii(be~ Mai!J~, ,a ile'i!JrMdenHM ilt Illf Universily ~ne:ta~, re\l'i~ers ~re very ilo\'JIiiIJJe ODe rlloller, ill P [tt9 [am dtr~clO r a:t 'the fi4i1ti&mall,E~ J n~1ittll,~. 'Yo u 're' lal k· I PWl!tin" wtw (hairt; iii sl'IJd,{ 5~dton I)il l~HJI1Ihll'~ iilnd IEIlIKH)" "The pro~ 1~1Ii! htn !;lilY li"'il:~'g~~tl1!i~Wilt i~~~\\Il ""~ 're I1!QlIi: U'tl~!I1g ut Ilhe r~t\!Ie- 're ing ilbl;UJll the :5~lJvi"'~1Qf' a lab," Thesul;4;~~ r(lt!l'S ra, bre"t:hlln~-bl.llter w a grants ~I'IQWfI as ROh ~e'Vid1.01lllCrease wilh each silJbmission. ,Acr05S, NI 1"1., i mUing ;flm ~he. eal." m ll'l AWhngll ~he s.y~'t,ellJl ~ml orks r<ll~rly weU Q'!!'erlIll. ~aY:i 'Oil'flid IlwslI\!;es:5 l'itte Hlle [Ill! mber of hmde:d qran ~sdivided b'{ all'p~i,,,~qoflS~ w 20(16, ilillll WE!{Ifmlillil ~k rm rIr~~' tilti!e Rm, ~pp ~i~tirmn s. t~ 2BiD,\bh;u SBOOOO 8 .. Perk-e I, ,a nemolldf! ntts,t at~he Unio;1er~~tyIf Waslli ngiclFI, Sea U~Iii, and ( sub lIIlis,....iom. a 00 tG ., 7% f'Gr tlni rd auelnpl~.In 1'99'$, il1l clmtra.sl, 1:1'tIl tHrIi! l;pu~mbe.r ;I,IM~I!!~'~s~iJ!dr ~~dil;llll, ~t ~Q~.sfl'l hil'lfli! the r!l.!sQll.!!ll~1ii tQ ! re,rI1'~U!~~lwl1e.n Hrst anlil thrrt! applirntiii:f1 ~1J(1ce:!<~rah~s.wa~ ~maiLle,r: r pred511ly r<ii'I~the~!Dp <llPpllCi3JlLlilm tn ,a g,i~,n pool Vet a!O,NIH PiJ~nE'S 21%~~dU%. iligaiml.ii) n1Jo-gN)WUn b!.lldgel, small di~L'l1Id~(I:ru ilm@llg th!! bes:! proRes\'!~ll'd'iffij\"~lld lhe !:Ie$! ,<3 rat! ill ngliiteslllJ lhdr (~,[~e.a~ (lj", ie~lji ng u~ Ilo.liail~filHIUer ~:n.[I'El'and IIHlOre.'·'Il'~ plil~5ilJt~ tiCi giil!UO d1,ekip ~Cl'%and ~ilied Ki· do it i" aI d ~ii'berClli VI! \\i'ay," Per k~~ ~ays<. ~BII!~ ~vh,aJlI lh ink .~~r~o1!llI)' 1,I~ir .sI ram Firoplll~al!j.. 8u'! ~ocking ~tU9y :'lec.~ons wh~ wf:lIl-l1Ill1a enlhls has:goU!!lIllh'~Jdef re(en~t~. Grant applk,aliollis to NIH illr@prlllpr!Jllll.em.ul.c. hi ~i.ilidlvl(!Jii'li] U1M to!!' Inhh. ~ For ex:aliUpl,e', Per~!i'!'IIii.a~ ~eel1iwhat he [,Ollis ~figu r,e-~~atil'l!l eUeltl~. ~As i FI thl!! 0 lym!ilk fi 1"1<11:5, jHie!d 'to; ncre.ase 65% belwee!l'l 200£. ,HiO 2: Oill "and in re:9p~!'IM! til ~tt1e lirs;t !ih~iIi!r 11.1i!~'~r 'ge,t'S ~ il.O~ b-ll<l;alu~€lh~ hil'dg~~ win'!I.~1) illi?{I~i? 1lI1~6I; ~l'Qm overioodEd I!l!WilrC;nl:l PS, NI H has (ut Ihe [11.1mb~r (j I pl'(lI1lIlS;!3'i:; IMd~ r~¥1ll!wer is .."ked III remel.Y., A~ a result, Ute I]GO~ o:f ~e'vi,~er~ hil:\i rlJ(!III1: ((I r,,~i l~-morl!-'!mp rl!ssi'oll! ~I! rro ~ma !1CIl!'~, iP'!!«,klll s.ay~. 5imulilrl~, in

hr~ ~U.ldy 5i~(~Iol1. the beosl§,mres alnlO~~. alw:3iYS:come ~.alter in 'tile !Ille~lfl'lg_,. IPed!~ r5<!Y:!l: 1i3l!C~ use' there's tbis IJCOfe uee.11ldun IlIQ ~he d i3i~" ~ II. il1l~roolllce~ S(lm~ ineql.li~n~. ~ 'Uhere's a 1fI()~her UifiliP oken m le, resIN! rene rs, s.ay: gr ant o1pp~it:ation~ b~i"g, 5ubn~iUed r!l;)rthe (hiJd ·elna ijll>l! time! orten recei'll!!' pf,~lcntI1lHiiil I,rei!litmel'lt, UJ~H orlly a1iml{1; 'till ree~rie5 J Some fevie'l'.le{'S a tlmit tn feeli IlIg 10rI'l bli!lW€~n a d~~r,~ ~(I!IO)5 ill1~~lil1!~~I) a 'IIuln~r;fl~h~ooll€algu>!l and tile oblig'atiolfl t.o ~ore grant j:![lOposals SlriClly acun;Ul1I9 ~!l merit. "'I Lh~fI~

20 APRil 2007

VOL 316

SC~ ENCE, www.5ciencemas,;org

NEWSI'OCIJS
snHlo'thlj! money finn DOIm.·F:uber has just t~e ·~v.,~' do. hUI I~'!enumbers do:n·t ~'OL! ~leked li.f1I. "The rull iml:K~cl hil~n't been bear I,hat mat" he's .. ),!;. Sull, eenrers hllve f~hb~\;~II!II~'~ int'>tiH.uin~l' h..ve pruvided OJ :I;lrmvn snmewlrat, In P.l9S. IhlJY !lfI<Jdeup ::;IIL uf'tbe budget, compared Wh~l U)"V" iU! hlIJ[CD", bu I. dll~ Ill.llds tlmt prwDdc for Ih9~ ~~,e 2rmR budget re(]luc."! fi~cd i~ Fcbru~~ry< h~Lffc rare d iMI~~Peating," he s "Y s. A~ .Brmv-I'II. ijcfll~i is rlll!l~Uilin~ {Jut CIf nlmn.~1' The ext nllI1JU[ a] ~[am ~ 1J11ilI.g~~. devore dI 10 [ ;U~ J wi U !';W!bf!li her ~~~ofl,d ~hot 011 her RO 1 i!l l R (I 1!' ;a~~d 'Olh'!;lr ~!'1 ili\'i dU'11 g.mn ts hJ:Ls iJiPPiild Jllly. r nn. esperation, she 'scast 11 \.",idc net III sl~ght~.y. room IH% ~.ill~9g3to 7S% in. :Ult}u. d her quest for ru.nds.;1j'l ~h' ugeve ry whe roe But ut"s not deaf that this is compn!JndJilng i ofimli,,~dlUi!~ researchers, :.ays fmnil d~~e'I~£ ~{lllmf>ll~ions to the D~Ptll'itln[;HlIl ~I~~tnnulll:e_~ o I' De fense, anti has aboutl 0 grant app liell~i![Jllis pcml:img iuidc and o~1l'siJc NIH. In t,ht: IIllle'OlIll nne. she' S<bys. the un ivers i[y nflere d 111,00111 her some IIlmn~y lh[l~will run out th~s~mrn· ~l~T-b~H onl.Y because she reee ndy u aderwent lr~1ltnH;:nt l;or breast eaueer :lmt {Ill except ion was. ~n8.dr.:.. ~fsrmlll moneydrn:~n 'I !(,~nm.e~h1'0!JM1'!wi l h~u mOil ~hs, :t! he MIY s, she'l:! be d()~~.ng Ihe S'lu~·kecmer ~hl!,;keeps 'I1'W"ml[e l1y species INl]lljled,~f«r'Oiiln(_l in~he
worst ease ~~C:iJmri(l; er h

.04. stra ~mCll'ltihle yOIiJ I1lgl Nuwhere does I,he Ulmdliin~ g,<!p seemwider lhll[!whc!l ~O{l!k~~~ lhroll!lh I'llI.!'I~nt' <lM.c.. 'I( .. h 0;5 j us: a bOIiH incuncej,v~)bl:L! I:or abrandt1~\"o' a.ltviI;til.ig::rto:r o gi.ll ~.n 1FI gr'dlrn.~lriill.:d t N on thei r ri I'SI. 'sul,ml iss ion these t1;)y s... ."::IIl's " ]).;!vid S'I.'I:eUn. chair (If the !l~lIr1}bioloyy c1cpanmt::I~~ at the Ug}iVa1>~.lY or Alab<~l'n;j.

or

Birmingham.

Sweatt h~slUred three YOllng

sdeUiltisuilli l~lepits'! Y!!I)r [md worries ubout their fluure. "I see il[!s this d.rrk _~6iiadm,\i I'll) n.g~n,gO'llc:r r(;op~cwho :;;re ,1 lei s I star ~ ng i oW.~he'irlahs ..· hie :says, "They're ~mvill1g to · !\,p~nd !\OflHlcli time be:ul'l~ o;L[J~",ua~l:S over rt!ndli('l~. to Ih~ detriment ofll!<ldll~ lime to think {.TC1U uvcly ubom d\idr researeh," Al V'<ImlerbihUtiliversLty in Nashville, '1enU~!i~L;'ie.lle~ i,Jrri c .~i!IW~~~i{l\l~~ i'~~'I!'ie !oipeciallst Johlll \ViUiii rns l\;ccflIll:y Ilcilrm.::J ,hat

Jab

as

M~!~}I' il,;':lt~i~l!' (:ilmpt<1j~ ~~ tm!g!! f1,l iJ1uJng·d irll:i:rt'l.: is t:uccrojll!.:J by ;,jJll cxcc~· i sive Wocus <II N~~.I on ,>osli}'''blS science,' such ;:I_~ [h~ C~~n.cer Ge:l1mif:!'L" dil:;'. which is A -,,~ ':1~ '00 '0], ~ 'O~ ' 0<'0 'O~ ~ LJ:;i~~t~3rg,~-l>c~llc gcnl!lic .s:t:'qll~neint to {h~empl1~r~l~mulcnlJaf b1il>b {!If cmtL:'l1:t ~1.1~.d t N(,,~ D:L:I:ecMr John Niede~l.mb!!r. :li.1lany bi~ whose J-yc-ar p~ I~h"$c is i'ila(l!:1:,eted at lot :$ ~00' mill 1.tl!ll_rrt)jiftlCl_~ ~ike th.i.~ ont' _ n,~:mly ~I{:!a!'ir!illlil im:t.ivt;l;-;.•~tNet liI;L~:Jr~ are f~.nd il1g •. l>Qm~nf'rhc !lame re$l'~I"iI;.:lu:irs wh~)\!,'(lLiIJ hi! se lent isl S say...are ClQIW ing HI ~hc' !::I::P ensc of grant" that sustain illldi"'i~tlmi labs mui have a pplyi n,g. fur 110 Is. •. h wo~.ld be ridiculoustu I~lnk we i been tine seuree of much i ~HU.'l\"'Hioll (wer ~iqt~ulcl1 b~ dCl~ll~ thi!l1,lS tud,!!y ill2mn ~XH'l:U~y t he ~'t"11fli. t~(: '!;''lI/'we did! in 1970( says Nil;!(h;r~lubcr . .It:·rhouLli titj'li,!.!'~ [hat [to Is ~~rt:3. Imvcr
pf~(lri~1i' ~h.. n th.ey used to be ... ~ hear Ih'l! "Science IlJl1I"" ch;!Wi,gell~rr~rmmc;lmlsly."

\\.1;:1 I. ~h~tlll~

b~~ fiw:lil Rm. upplication nl~led to make lh~ cut 01'1, hs secend ny, A..lthO\lgh Nn~ insrinH~S !);1!;i~~W.;l! Iy g iv~~~w iI~V~~l.~ g;I.I.l)r~ " bmH1S, bY~rriCrC~il>ing the pan}' I iu.c-fmm, 12 % to ~4% ill }HA ID. Ii~ exampl e. or frmrli I" 12% to !.M"/,-, .u, NC 1-[11;.'[t (lilly hdp$ H' th,,; gl'an~ i~:r!.~ ;llnmlg~he <If.lo/~ ur so 1im~ are 5Ie~ r:isiJlI::wlllclil. I.htijl ;rc '·lr~.t1~'t.:t1.. 3l'iJ .~t1: n unscored by c'I'<ihmtvrs:., This: happeued 1.0
Wi~li"QllL~, \\/l:Iu hrft a bl,~uer·~~yi n,gjob ;!s an

emeTg,enc:r'rnom ",h:W'::I'llf n~~w :;~mli~i' n ,u1d rL!~pirL~Wty vi rus, ch~t :S1!ti:k..:~t:iIIu klrea1i.'"W!.: leve science, um~ we
WIII1t

hi:, _~~ys fJ f phy!> ic ~a!l~ ti kc' hi !Il5e 1f. "lrur W~ ~[w..: 1.0 eat," WiW3ItH;. who 11111> four cmuldrctl orh~!i own. :Jpprcc~,!J[ws lh~l NIH IS

to do researeh,"

near l~d(l~1ill lellil; III tile I)f!~l lew ye;;If~, iJlllld ~~Il'l' qu all i tv i!ii lfI~l ii!~WaV5 Ii:!~le. ~ (;on.O!!d!H .An~oll1iQ Stii I'pil, [lj re(~Or 01 N Hfs Cl!l1t!~r [o.r S(nlE!nH'fic I'te'll'iew ((5f{), l'llllidl ooo:rtlin;ilte~ tl'le gralll~ review IIIr{l(I~s~. M®reo,ver, m,lnV ~f)fil 5.dlil!fI!i~b!" 1LI1111der IIlIr'eSSlm!! kI k@@'il theilr ~wn lilb!i pnochH::li'l'!?< dedine t® 5~lve 0.111 ~tu~ secUmu. Tile !a.rk of iL!rop~rf~n(~~ Hi~nli!i:t~ cm rt!~l ~w~ 11f'~ ~s.ell(lTlrg-~l<Inuin 9

hs'U'e that's beem incem'tlllilted i:!V Ihe currelllt hmi:lillig squl!eze, Si!~Ji IEd~'i'arld Kr;alo'iU',. <I mIJrtl~(iellll~~t iIIll-li3f't,t'almlM.eili,e<ll Sciloot ~11 6M~on_ H~ ~i31!j1'Se':!; thE!! recent !ficfim (If Wll!3i~.he claims Wil~ aJ poorly inrnlled h r@\!'i~I'L:AHholLlghthis group h~'5 pl!Jlbli~hoo fOllr p"p~rs in Ihigih-l)rlJ.fiIe jomt1:11ls s:in(e NOll'etlfiber 01'1 Ith~ glelll~ll(Hiasis o·r agg~eS&kljle be-havim in hli LI rli~, !l'ds month OJ pmprmlll IlJ r:(]ntil1l'U>~lh~~ wl:uk 17I{IIsUtli.. g'f!dl~r!!je<c:~edwi tnollt being disc.U5~e<d by [he JllIli. ~;.Ludr 5 e{ ~ijof'l. ((SR 1~'1'5OI~ I ~ohd5lUdy :seuiofl's, lhl abO!Ull 50% err!lra"~ CI!)p'lk,nioIll5 shauld be Uia@.~~ ~() si3Ive lh'l1f ror dt iilti solng: th,~ ~~t iilpplicati~n:s.,. ~tiJt Lne ru t~has b~en !!'On:! stri(Uy elllhu'( ~ ill ·thii! 1(I!>lyt~ F (lr t.lN'(l, SC.!Irptl S$~:d "Thii!re's s®ffilellil'l'Il~ "'Ny WiG FIfI her~. says J®h III HUdeillmnd, iii InetUro:sc~el'lti:l&at ~ the iJnriY'efs,vl,V ·~rAiri~ofli ~Il 'l'UUOIl, wh~ des.u~tJe~ Kr;nrHz's. fly reseaF~:h a:> "U'nt~ft~ fIOVElI.QJI d r:J'~.lille, ~ 5,.31rRJa:5ay~ (5 R 11"'5~El'\!'era~.IIIiilW ~ntili OIliv,es. to re(11I h qu a.t[jned r~\I'ie-I'j,~rs,.illldllciilllg ~olllcl1fFlg JifCOnHI'U~fI,daU~lfIsram sdel'lWl( ;,oci· r etie~. tapping! fIII!He re-~eart:illm fr211i1 9gl~ide ihe UWlH'ed S;l;1~e5,:and ~~~,Un9 W~1rri- bil~E!t!I f~vi~1'\t$ llrlil~ ~9fi1'l req ui r~ J~sl;:!lIr'4;II1J~fi tri!\r~ l 1,0 t.o
INIifl

he...dquarttlr~

NaUonal A,ademy !Jf' Sdem:!!~ rK'erIl~if agrt!~d to ~ent'i! Oil sud! olllLilll1! ~~udy sections betBlu:!i1! ll'lev wolilldlll't ~iI'Ve t~ (r.we:l from (aIlilOFlllia, s,.car'pa ~.!i1f~" I-lil'l 5e'iH ~lii$ ,n .. n 1i1ll((:Dl,.lrafliI1l9 :slEin. "Thlt revl~w i~ good a!>Lin! review@1r~11M, S~ar~il ~ay~. SD that. Ii Ofi~ cd a~ ~ !:IlU r'irs! p!'kHil.i~~. 4i.M. PINtD' ~.C. b~li
tl

or Ill!

illl Belill~5da.

Mil ryland,

T'hrER' ~11[I~O'r riNlli~t5.,. nllilm,· s

\l'!llAl'w"sden.cema mq 9

5CtI:EN:OE VOl316

20 APR Il2007

'l1~e' Ga[rdlnef rouudafionis;pl,eased 1I:Ilt,e([nOll'iof~.d Avr;l,:ITils.

!hi:
(;.I~ruLH:r
0,.;

~o n~~IIUHlill~C:.e \,"]mu~n of the lOCI! G.~~nll'ul'i; Ilhe

A'\J]l.I~ ,lTI:' ,mw1ll.N the IIUM prnll!-\i~lll" J\\.Lnh Illf 'H1L~L""lIdi~l~ Jl'lo"l~,{"lil"" III h nl duo' (~,lil.,"llIl;"r ,,~, ,IMl'L'" ... 111 i"L'l d~~· .,,_\oJI~III .!I!d .! M,IIUC. "I .• t ul:ur~ , .u lh~ .1~\.lrLb,-1!1\_<l]J.Lmy ' hL,M ill Tllf"llhl ndlillot_<r ~;;" ~(III{ ~m",' lb~Llr ],!!'~lImltl~ in Il,?i7.,{1MIlj 1M.."l, {'dtnl"i1~r \"\.Iin.I('o,:, h.lH· :2;<)11': rm L.l wm Ib!;" NIl!>,d l'rlft; ~il I,:'Hilot' I !\h:'~I'I~HII;' Ul ('!l~'l~ll~[r} III ~hL' 1.1~1I~\I; ) L:.lr.'i ,.lnl11;'. <J nl" II Nnlll."~Pq ..~~ III \ 1~ licuic h.l\'~·l!' m~ II ~ p.l~l (, ,I] nlucr w ILIIWr~
nl~...liL.11

u-tu

1;", ~ •• ~

i\~ r-~n u( il~

JI~'['-d.l\ N.nl,'~l!.ll PrO!-!I..I1lL ,]tTO"",

lk.hIlleI'
tIl

~2-~(,. ~r1l11LUrren!. .md

p,l~l ,1\\.ll"tlt·C:~ \\,iIJ

~PL·.lk III 1k ir.ulrmic Ll:IIlIn

c'm.LLla
,I Ilu",

([,uit\,

medic.tl

.II1J p.l!l.l}!:LhJU.Jll'

~lLLJl'Jlb_ ~Llior
\~m'll"~ILlm

hi;.:h ~,hLMl

~!Lld,,;'l~l~.~l1d Ih~·l'llhli, 11u: rU~illd,lti.lll

nlU IlL,~~.1 L\\'HI.I~

11 d~r'I<I!ld Ill}.!: ,Ill{j

("UI1,illt:f

Tiv, iuuciu .~lId lli II;." .uuer" ""~l~.I~~r~ uiclude th(' 211r1- .lw.lnkr:~, .I~ \\'dl ,I" ~~dll!:')lhlllli:f. _Inlm I)u k , U,ri.111l Iruker, i'i.Lpo]n1ll1.!" lcrr.lr.i, R~ll~n Horvuz, I.lk ~!.lk, [~h;lu,.d Ihn, lum H'Lltjo,;,;~lI• 111.1Ruhl:OfI \~'l!'lLI],l;'r~ •
m

,.l ! ht;" U!llh' r~Il ~ HI

-I,L'I'llllll l

l'l! •\.ch

liIt' U

C. Da:.. id All is, Ph.D. r

JIl,~ 8: J!:!Ck 1IF151'lfrnrn irufe5!~!, P Head, L!lt:lQr<jl~ory of Cl1rom BUn Bmlo.g". Ttle Rocke/elrer VI'lI~er.lJI}'

H;;ur)! F~NoUer~ Ph.D.

iRii;l!iJert w'uis Sll"Ishe[mer IProfns!l!r 111" MI@lecuO'1IJjBf:i}lltl,g~ Director, Center 'rar Molecutar BiolQID' of RNJ!. l/t1J\-t!(.My of ~lJfor'l/l.l, $ilnf;J ~.'!
.llhl

''j~)!r,a~i!",is.f~)~r(' ((J.ie.

II

Dr. Kim N~Sntrtli;!!,fh, D.

Wfilil:lo.~ CIllOIlr

Tho!!uas A. Stcit~~ Ph,D.

St.erUrliG 'Pruf'e:i>:iCllr @f MQkl:l:'ulil ~ IBlD,,,IiQi~C5 & a:lQOOemI5~)r
Inv~tlga\_{}[. Howard Huglms. Mooical

Departl11em or
OxIom

B!OCheffilS1ry

Unl\l'ersrl],l

lnstuute

'~/'ojl' dtl'(lU,H't~~Utetg~lllm"t~,u ~ in c,rU .di,~.i i~m . $

1.J

'mit!' UnIve~it,.

'f~fi.JJI ,t1~t!st r~fd~"~le .Jj~ul:'t'imJ tU,I'ii!'

,oJ r/u<

Y'lb(JS{uu('"

j'i

Dennis

Srnau'1OillI.

M.D., Ph ..0.

Pr:ar~s.sor of Medicine. Chle~ trF Hl!lmalolCigy/

O!\Coltl&\l' Oil1d DiE!'~ IJlMl \llGe Ch.ur for ReseO!rc~ For me Depal'1.Irlenl 01 Me!d It:llle

'~J~w ,tilt'

UClA

del-'e.l0l'J!Ul:'ut

t~fHl.'iJI'CI"'lJ'ii~'

ill ,burt'a~"1 Cll'rfeer tJ~C(.l,t"m."ril.,!J The

AIRD'NER
Fo',rmdatlo,r, Vii'Ww"gailrdlner.ou;:

,TOR.ON'10
Iil
Nfll!IJ
h.!l&KIufl:ODEI:r..ni.

~o,Ii~"l'i''I'

'c.

w., "'CI.!irJ ~kI:i Ill! {ldJw;Iwlg~ ch.1l ~lItIrlllil UIL, UfI;..:J,!.ily

'i:lrfrltl::~ ~

NEWSI'OCIJS
tfY~~l~ help, 1:0
dOI1'~

Bw "the' elfofls bein:J;l made

~IC,I'yeffective,"

NU!i-Il1wi"L New H.. nll,~hire. Cone-ems "bout n

IhlllC~ [Ill!w~re a m.Hj~rli'<Lc~('II' Iah is decislen, ~ Rck:ul'l s~lyS+ Throl,llglmUI. bis gn.!.{hmtc seheol C!iU'!.!cU' ~llltl the fir!>! yC~Jr of his pusrdnc at M If, he'd ~,cen menrors and

~!.;,'enrn he 10

he ~lIYs,
app'~:i·

As,ut h ,~ppeUH. in 200(, fi r~~-~i Illll

cams, 3cw~,:I,ly had ;0. better S~.ol ehanesrabli'she~U ones ;It scoringan ROI or e[~uivaleru 1l:rarn OJl ~h~,dr fi rSL try: 9% C'mbl.p~Lno!:ll to 7%. ;u:'r..~Hli n1l Hl ~he Nm Ch,!fi;l(:to(~ nff'j'J;;I;l_ B m NIH efflcials nrc Il1crca.s,il'lgl.y worrie d

ruspccrcd scientists
..bout th'eirfund i~:!i!'"

C'ml~l.am~.y frt:H~.ng

h'lppen~ when IWinding p:llterm. shi fI <Ibwrt~)', "Osee you've c.."r<mdle(~ Ihe [fIIIi$C'<'Irohi COlll1LC~.~y, wim~ ah~ yo~ J!,lo~[lJ!l [O~tlT' OIl-ikl' Nuhel, "YOt1just ("Hn't turn off'[hL: ~iijgul ~ukkly and keep Illle oeillgine wnrling m 111~1 gear." S~.ilL, ~;:Iu;.dds, decisions :ilJou! 110w to prioritize . bi>L'II'li!:l;Icii,c;dr~s{l,vrch
"liIfC fC::LI]y

uhout mi d-ca reer scie I1,lis~s.~ho se seek i!lid! [,0 11011~\,rthei r fip.\;~. or ~!,_~t:muJgmllL "That \
p'ro~bly where the pain is maxims]," Slll}'S Z~rlimm:i Lastmcmb, NIH 1i.llJIU)lliilCedt,ht: .. D'lreClor"s Bridge AWWlrds." p'rnm~sinl,Qupto

"That was 10":111:')" ~.o me :'. he lll~yS. R.t: ka rts :H~\' iser;

llil.adc at a

sodet:l~ level,"

!'mT neurusuientist l\i]1urD ['''iUlan I.i IJilC> ha Pawim. Sit}· s he carne '$ 500,000 to ~~d~ ~f ~eh:cNtd r~se;)l~i1I'ln;,ror HI her 100b wirh cutm a year~~l0 jus\lJHi ss ~he l:ul1i!.lilllg ~Ulofr.B~~t ~mll1idillig I"C'CrnUmlCDlsuch llI'-;SU~Wllcei:-; the ~ullle ;IS J;;!l1ding ~1 d onion ~ :1r1d t\ num'!lot grl1ra. Vl.ihiocl~ h;,"[l hkg elemenr HI Ihe ~enmii: bel' ofpubllj,calions,

IIJU hopes I) r intlueillcing those deci-

sions. S.dl'l1iliiH!L convinced that an ~ncrea sc in NUl'_'"

1.'!!Lj,dgct ill w
ilmlJ~rovclnelill

mark an

in their
alao alierna-

Zwdl~Il.J,l, "ice president for rescerch 'H~miFlis~nu~.QI1, M. D. ~~ AnJcDo:n C"rte« ('enler in HQIl$:l\ln, Texas, ~jh;c~m~hot$rl'io (,l:udy iri their ~l'Ir~er~ are ~~~lfIg ~he :;.it~;;Il!t'}n ;;Jnild \'iQ~dI;iTi [If: ''I.~l~!h~f t.h~y :S~.lt;rnnM!.!WIl try i.flfi,u..: ii out, ·'WC·'rI!.: losin g some ve W)I la ~ ented peopil.e 'Whl~ a roe d!t.ddi mi.gth .. 1 t if!(\'lre'~iJiOl '~!lPllgh ~,l~d)ill i~y h~r.'e.·· ays ThC!I1HI:> lnsel, J~'rcctor 'orth!li: s
}·bli>tl:tl;:!)II il~ljW[~

-cle(."ilstnl:!:' ~!.1yslLe{l!l .. nJ l

"'lhi s I~ll(U the kind ufpc.u:;un !,'u!iJ,'dwant to lose." she say~.

NHI

~lornLIlC;fl..• are lobbyi!l!il C'rmgre~ he,~v~

U}"

SOll.h: MC

consldering

r!;!l~ ih~
t

;ilfl'patL:'IHly ~<!~l~ w ..y.

ilWMdil.~gRckl:ll.rt.

1.

fa hni fl,g. grant last y(\'laf.~y uhen he 1l,ld

IIllr'!;;'~~d~d~dd~d ttJ move tll1. a~~d fuJ·~ ha"
11Q 'rel!l:I,"I;H~,"I
10': 11

I:iw :flJrld~img sn~na~~., ''T~e I'©~! qtLest!9!.1 to rue." Stlyt> M ~iry Hendrb •.•presideut

Olll:

~!\

M~i'i'l[l~ H~lII.ll.h. Jell:\"Hne RJekJr'l, \\'.110 was a 1)~1ts

or

~HiJd~dt'~liH";c di~c~ i,oli" 0 r l"h:i ~J lien \:

MtHiihJi'iilIIRIC~~1 fch Centel' :It j'o,l,(jil,'tilwes:l-

d oJ<:ltl>r~1.11!Uow Oll t~~ M m;;:s.,u;;blL&eu_"~n~ti· I t use of Tcdnlu logy {l\ii In iil Cam b,rhJlc u nr ~I he' lelil I.:l:sl July ror Oil le,!ch irig j nb ,II R.iviel.' CoUc!::.C', a libc'Ii"<11nrt: s sehnol in Prop,m1si,lIls IFill G1l:ded il'l2006 B1; II1Slfitll!lt!l!:
N~t1ili1i'l>'!l1 liIiJi'fIam li!nOtll'li!! Rl!!!e'i!Il!lh Imdll!l:l! 31:1%

eonrrul

teach." he says, '''I i(ih~n~nfin.Biologj'i!. SU~11 Geubi or ll~iI'I Uni'o'er>ily l'hk:llgu, llliaois. "'is WIiIrnies allllU't.lo(sing her ~tl)cks (II Sciara, <i orar,E fI~, hnw W reeenst iHIW don't ~O'I\"cC'onHol the ~O'~l rm~di,tl~ uver Ill"" N II I blJ(I~;;:t :.and im\\' m anypi.1;~\l!S; pfl]~4; pi.L;l~!f(!<!wlil~btE;! wilhol.H relyil'l.,g 0111g(wernrn.,elH suppert," and whclhiir ur 111m~.l'fmgcl. O.UI.C,," Hendrix, ene af ~ht: leadership ~r.uLir who mel~'(.'ar.o; I:!IJ4IO II,l)!,1k tte~'O'!1,d the NUJ deu1,0 Wllli~e sonre senior SCi,!!L1~i:-;~~predicr [1iI,L~ :u !:!!~I1lJroH ion of yOUI1i.gm: OillQ::l>\'\I iB dis,. bHl'i~. [<L!mr.s; bOO.l!.1~l!l pncl'li(l""p'ri \r~trl1l~n~lL:r~ i r o c appear, olhl'ir~ rc:jec ~ sue h glT~.l'f!rf!~1;!(: asrs. !iih~p'~uld\\'orking n~ofol~k,s~lyw~ Ih eli~.,"..e
"There ~sa sense 0 f uacuse,' S ays\l';:mnu&, Inn .•[ dkm'l till iI1Jkwe're lruing youny peopit: ollJright. yet,"

~HI"'\' w-eU

~H!

Ull~vl!r~i~y

i'l

.!hLntlih~liolils. SI;)tl":S. Cut example, arc IllcrtiJs-. ~n.sJysitepll~n£ iii! to finance humau embry-

onic:: ~~I~m-cel]
BUI.

rellm,nt:'mL

Ni!t~.aI.I!ilm~wie
H~~[~ ~~~>Qe5

or E!ilvr~rn.em"'l
21%

N~tP!ffiill rmsti~wtil:uf Di~tlf1It~ ,;tRill Digil~~ve .:!irndll':'fI!IImll~1i' ~.\'lse~ N~t~~1 i ~ffi~!II1:Ei! of iW.~!g~ai!d ~nJe~l!l!tn Oi~iM~

,LQo:king ahlilld M.H (Jlficl<'lls 1irt:~ln.LMgljlll~, IllC<'1.iIlltNbHc;o t !.;ecpt he pe reI: Iii r:lge (if ijl rOIlOS~1.h,rlmd cd fi1:)!t~.:>inkirl!il eVeDl km;eT, B~~~ u!h n'f,) oft he w rt::;c'drcb budg.et tied up Jonllng,oing pmjt;\:'ts.
il~c.l.llld~.UIIggr<mts 'th;n kISt."~~·~r.d years,
t lilie

lIitilmllely. COl1llgr'C$S w:i III set the puce", mliJ l.hcr~ are' g~ imnn:~r~ next years dun

budg,elm;;;,y reverse I~le years of 11m fil ~di I~,

Ah~lOugi1rrC(oi~dcnl G<enr£l.'l'\V. Bu:.h Ims

imli-

Nn!clm~llmm~lIite Dtrn~1<Inc If .c1',am~[oc~IIR~Jrm Nilt~ill.lmml)wtiil ~llIdl5;Ifil'~

l~% [)i5lll,~d~!1

or U~()'~Kill

di reclms ilInd ,Zerhot~m~ &"IYIhcy have uoel:ni\'~ly liul~ ~~I,f~ni~e,M"ny i!l~t~U~~e~ IW\'1l:i t~~jnc\L:h;,dor Je.I'tlycdprogr'::LIIl::>. part~.cliI.larly costly di,n~c<L1 ones tSd~'nce".2 Man.::h. p. Il(l2.~, ,1\.1 the Nl1Itiotlill t-h::II'i. Lung .. and m (ladln!oit~~ut'e. Diwf!!.:t.{J:rr: Ii7.;.b~th Nil be I s.u:t~ lohe h~:lOput off fi.Lildi~l.l!:l;ln1.."W hyj:'icrtensiol1l trial. ,mclm }HAm. I:,mnei Soa)':;: he fre~J 1,1]1 S ~~ ll1!i U im:l 11)' del ...:ring I'W! rt~~:r devd(lp'menH~r.,m Ehola '\I"H::l.:'in.e.
Ile~,'I'

fr-m:llrllm:mlcd t'tming Ihe lou!; NU-.Ibudgcl1 by 'Ibol;l~ S5(JO mUllmll, ['Olllgffi&<; ha~~ lIUg,gcs~ed II. may pm,b. the ol'ller \V'ily. In rL;lbn~,~f)".S\'obod,1 :uLSU Je~med

th.~tUj), hill ~ h.i ~t ry. ~LIl,J to hi~ 1::,["':1:11 rei id: his gr;JI1H 'wo'uld win redel'il~ 1111 fliW, He O 8(; ore.! i1H he 17~h perc em iIe and ,,~i:llIx: recOmm1?n.Q!ed r')f ftun,cl.il;lll ;Il tl1e N,niomd
IIIslilltt~ of ~ril\iinHlltl(ilm!l~ I~.~caltS()j.cnKI~~., ~ As <l res~]~I.S\'obocl~1is l:ee~il1g a 101 11l011e orl.ij.11l1i_~li!.: ~bflul .hi...t~~'1urt.'~viL!;w ndl. lhis;

N~tf,pm~I.!'!1i5ti~!lite ct.md I'le~ttn"jjd gf

ihlrna:n ~iI~lo~lliI~nt

At thJ: ~~~m~ tilfi1t~""\\I'C e:il1.'1 [lo~h!!v~ :iI'liy .rl1ilimives.~· ~mys F<lu(:i. "Seiem:::c' tI.llCl'o"es il~ ".way rh.n ~'I'U h.we 10 p~~ il !>o!II1H;ll~!Jne:s_" ah.

r~w

Today's,

~1II~i!.:lY appt!3l'S

to

foCnL:Cl

what

,Il IhOll!;h. the of..oxpe.ient:'e len·him d i,)li~iml,i"D1ecl, "U ',liwipl:d otltO'iy j:'iilffi~)ljIIlJ: IHit-:"'i'lt: ~a.l'~. "h';:::just been;l1 bl"lll!,l,IIUtlL~S)i," NI_~wheh'ls,~ )'e:.rs~fm:1I;! ht!'U nL;'cd ~tI if.\l'In1·peh~ tll!' filJUJiii1j:!again. -.1 ENNIFIE:'1i: (OUZ.INI.A~D 'GUG MIL.lLEIJIt:

\l'!llAl'w,sden.cema g, mq

5C1I:EN:OE VOl316

20 APR Il2007

NEWSIFOCllS
BackeD trOick? w~lh tile \IVO~llCi Mili~r>ee:img 'fr>em il'5 2003 Ibo,ymlt.llf the poliIlVila:inc-, 'Ul>B NiI]eriillii slaltE r o' Kane hiliS re.st!~C, massl'\l~ !!TI!!TIuni~tio n ,~mpaig;ns.,.

k."Ch!1ical d'l:lI len,b'C nftl'yin,l;l w \~~t:'I."~Il;ll~ e"'e'I)' ,.;111 an the planet mult i,pte times, Asdt;:!!Id.I.ine'5 Id rove ('"Ome.ul(1 MflQl:l;!'. t~lIiM~U;l a per(:Wlion ;I!ld
of dim.unbl1illg naurns h,a~'(;guu\!i.'ll. aed it 1l!:!1£

become ]li!Ic.re<lsil~g~y lOllSI'! to raise money,

R~glilllll)W. saysA)rJW<1ItL llltiep~l)gomlhij fnciri.~ Ii tsmrk'.I (.'fUln(:.~~ fml{l~ngsholf1fOlIl e'>'e'[ GiWl' Ih"",'\Csetbttt:ks.lt'S nOI.~lilTir.il$i~'igthar

q~lesti(llilhl£ whethereradlcarinn cencedes Steph~1l O)Chi Oflh¢ globfi~ 1.I:mmillilZlllLioin pl\{),~lnln <II.[~lIe U:S, C{';I]1.\.Th~J:r D.iS!,l:il:-;e Cmll[O~. and mVl:rull iotl (CDC') :iIll..'\llamll, Gern:giill. Oneoftlte mloo~poolni:11Qln ~~C!JI~hr Q·~..l-~!>e~~~em I ~;~lne .in ,in 'Irridl; .in &'i(.'fK'e !i:lm.y<:flf by 1s;'"JIJ'Ar,~~,.L,,'1en'I,W. or both ~I to aod ~1l31 ~po,. erndfcm ien C<lmp.'l.igns, and x
31.e

people

11r.I00~e!'i Sll;:i\:;;~.

,I

INFIEcnOllS DISEASE

'Some eXj:llerh

~1ij1,VePlmIP®I)ed iilllg!illlido:lllillll'~1 eff;auts~o

Sail'S

'eradi'(ate~he

'~ifll:sil'l

fa'!ll'o:r of

wntfolllil1g

U~ buta f!ew aH1ialysis

tlht. o®luld be more t'Ol:sUy'if! till€: lo:lrng run

~Er.ldie~lt~ilnil'; thehl;':"~ hlly.I[!rn;d,.e'~ nnt jusl ecennmle bLIl ~nlln1l.n~I<!f~,m sense," S<l.Y)1
Bru{!c A}I~\\I'"Jrd.\!"htl \

co.1le<~i::;5.. "We he~ne!l!e I~ t ime has come I:()T tfil~ g~~1 !i1r,1~~' for pt;l~ij.o~rI b~shin~!tnm "c~,tdtcal.i(rolt ~o ·dl~"l;,'ti\.'c comrur··tln;y wrote ~Sd!tfl'\~"12 M.IY 2[ItlM'). 11', ~51). Skerli.c.~1 lh~J 3 ~m!!r-~{lst, IOt\v-c;jS~ (I pl i~!'l ex iM ed, Tlmnrn psrm "Lf.! d Du i rl ~j ell Tt:bb~ils ~l~!.t'Hlit to in\'tl~t~~~il~, w,ii1g t11utdli
rn'lll't

the ~hUV81'd Kids Risik P'r\~ject:

fW

Lma y~ur .. ~~\'c:r<11 pUb.liche;Jh:i1 expertx brom.::lled <II.I~c~·e~ idea: M.!)iiJ;e il: i~ ~ill'lle to iC":~1

gi\'~ lip on cir.ldjit'ati)]~ pul:io. ]n51~aJt~ftryitlg wipe out the disease, perhaps lit makes mllre ~Ulsell] seulelbr (;ol'l.trolli.llg it ~!f!;~ l!'~ d~l: .rnoncy ~h<1[would bl: saved fur IlMJ<i"C
It I

durm;:tt> tI~t: g~flb.~~1 polio-

p:m~~;ng llC,ihlll jJmtil!Wlu;" sucaas 1~lar~a O[

[llV/AIDS, Th(! .klta quickly Wl~U!!cdadvo,c.u:es.bul det.lils \Vh.lla (o!l1lnui str.meg),

,[)'W

ern(lil.':a~jolll ill~liatli\li: rmnl theWolr~t11 k'.<IIt&a Org,omiz.nion f\li,'1-[O) in Ge.I1e""~. S:\!\.~m;nl;ll'1(t ~1Il:(j is d.C;Iltfly dd.~Wllcti wid!. the new ~Lna!ys:i.i>. .. TllIonlrsmt~~'II1;;n~k,~'~ in Ih~c![]lrln f(lrlhc:: i fIlII.il ideu lln.:lt there is a clleil P and pm ialess \WL}" (mI." Even c;:ri,tics;uftlu: ~~lInnll<luf,l'l h!i!l\'~been

~LLl1pattor t~is study eame from WHO. f CDC'. or uth~.r I},l!jrln~r~ in Ihe ,(;mdiciltiol1, ~.m :ni\,t" •.ThuJ:ilPSOIl .~;ly!O. hli First II~e~'k10ked lilt \'I,.hal would hilP'l,c!1l ir e!1dt,l!i!liCl:"aI!llMri~, ...we:re to :'i(;'~.~.!,l back on ii~nlnllniizr;n iUIiI: To \~I~ill extenr, and bow (Iluick~y. ,\"fluid pniin C;ls;e~ lieb~)mu]'? They used a modcllhm 11ili~QJ.'S v~rJHblcs such M j~l l~e b<!~k repfOd~lc;:tiv.e m!Unhl'!f nf the virus, the level (If populmion ir.mnmility. uad OUl~

milght \."I~~fl~i-flm'll \\It~tL~ ~[ would '~O::"I-inavc

been 1;ICkillgl. Now, researchers n.t I lsrv 11 I'd School of [l'ublic Fle.dth iUl Bostou ~,~ve d~vei{llU;nh nlltl:th!:m,mE('f11 modd 1:0 explore the op'l.ior:n;, Spe(:iri,c>l~:ly. I:hey al1la~.)'.;::c almost a dozes ~mnJ'OI ~'~Il<~r1(.m-:rn:~Il~ ,I~!;~!>ive e:tlbr!s l{) keep the mmnbeJ 'QJr 1Xl~[(~ (,:::lsoe~ver)' 1m.",.o t 111OP~ I'll. in imal :L 1][ ervent io:m; ~ha E. WO~1 ld ucuept a ilIlgh,cr disease burdlen-and rhen ,~.:o;,;In~:i ~le !~~~~r.I!ls;n l~fIlm clf·cO~IS Vl'.:if'-"1.I!'i Jl!C i
piJ:lio ('I)SIL'S
l:tlI

cm"h.

El"adic~lD0I1, ;!s~il,lmirlJg!t i:i. fl,'l!J!>ibl~+ m.ighi. yet cost b:mions to achieve, But il wCluld ~~ill~e che1lp,er t~mfl ~n}l cGn~rnl
opl..o.n-'t:' .... dtrnK" ~1l[u\i,'<iuM leli ~11~d;t>CI!tr l ~n,

U:l 11m, nil 0<.:1 )'ear~t ~lel.e~nli1 J'CI)Ufl~, .. It LS not inlrLIhivc. ColuroJ. tlo~s not IIppellli to '1)lrclT mlY financiOll ~n\·~I'I~s.: !{,!,~j~ K~lnbe~ly .. lllwnlpsull of [:I~lr'l'aru, 110 led U'O.!:emll~l \'1.. wilh l)(ls~.dm.:R~tlbOlnd 1. Duinlji!l~ 'Te'bhel1ls. irhdr results. we're pl!ibli~hctl UD!.lilM': hhl. \\'&:I: by rb~! J.tmnN.

Uttar rrm~t:~h aad Hihar; I.'IIVO :;;talTC!;.ull nnrthern ]!ld:itl where polio ~rulm:oimi~simm rem,~ilils ~[ilh;I'I_'lt:: J~p.UIC (~IJCil1Sj\lt': WlccilUltlun. The model Clol1ifirllili!d whilt Ihe' experlence of the p.;.l$l few )'e,ir~ h~:;,shown, !\"iJ!\' g,11lrn 'lJl,lll~~ ,'ilu!\hl:d ~;;,n;L";i~9Wm.tnl!lil .,Lixl~!1 l'hom:p:son:'\tr y{).U 1M.>e1!-: }'OI.J! wHlllil«':~y see hUgti' mit liireaks," J U$I ,I :1 {l11\"j) dl:ld~~~ iii 3;5n,l}[lQill ~9iMH W, alXH!I 2.OfWI in 200t'i, But 20 y~~Jr;o;, mul $5 biJ~;oll 100t~[. IJ:mL Im;t :iU'lnn!:miz.n.i:mlIIJUelll~i\IIY l'e3ul:led in 1110'liCihmt 11"'/[1 remains e I~LIS At~Cf cl'1,\OJ)pifig to aU!.,bl!- 50{)lu' addhion .. 1 e;l;peCEedlll;:<!.s,~.~ }'eOl~'in i,v·e, <I tim~~rnl!'{lr'!·ll) in20[)~. g!ol~l1. ~.:!.~ .. I~\~ h~~"" I ~es.\:(wo :tltdimll states fllgJ]~. \.'f~ ~:~nmdnou ~~ lor i.1il1i: P1i!l;t ? )'Cl:1fi>, Then !hey exIl19rJ.."\I the I:mINt]!; o hruio~b':> mil~~rolii[jrel:;I,~tls :JI rennc i~l~iS ho I cI i~l rn~~r C~1nlrQIand e[,.fligki~ti!Q.lrn scenarios I:~;"rtile "mm[l:kiS-lndiLI, Nigl.:ria, i\.Jghiilnli~tan. anti world's low-income i;;o~!mri,~!'l. here the w r,~kiS:(jm-I'rom \,vi'lit:'h i~ period~!;~IU}' erupts b:~gg\l~llJilrtl~1l1of (!!l;SC~WUlllJ .~ikdy t1~"Cbi[. III. ~ ,1LfilJrdl'i.I~ctJ> p{jlit)~r~ct: cmili1lri~!O. BL:lWl':!Ilf'iJ ~If'rd io is not an iss,u.e iIII \'I.'l!· .. hhy ",",ouillri es ~ .2003 .lIltl 2006. the Im}l,J:I:am spelllt SH:SO nrul\~11iL'u!!;!'tluli:1lI': \l"Jit;;\l.i,IJiLiLOn rates ~MIft" hi!,1h.! l ~ lion lnMling omb:rt:~"$ [n 25 (lOUi1.triCS. CnollllrO~mc...lll!s. i:1ll g,...e. . ~ ~l)I~mtlng II .t:."ef- ~ Mast e....per~ ugree .h.n e~udic.'1.~i(m ta'!h· i... lail1li.LnTl.be:r of pol io ~~iS~. a~;!t'm'. bul.th~~n.:.is .~ ni(''1!H11~<1.intc.gi\lt.n lIJ,'llil:!'1iwd ['il~(ll,JlrCt~_ BLlr no :I'Igru~nt: ml \\1\ii~1, an ·'ili.ccI!l'tl!lllle·' ~In~ I~l lUll!: '§:! Ille plrogrnm has bilen, s:idelhwd by bullig,Cl migiM be. A rit,~, ft'lf im.;;t.a!1i;:~. fit ~l~lPIll f.lo:-<t:d ~. :til~o:rl:lillls, nl.n'nlOrs ubmn vacc.in~ S2Ji:i.y. ruUkeerd illS I'lolio C:lS~ 1.0 fewer .han son .~~l'!12f, ~ ; nlOl1'1~)tilHlamd DuiD:llj~r T~b'IJ~Il!S cla:itl!Jul ~ gi(lll:.~'lWI!d PCl<h.tk:.!I oIlPJ~o.~ilion. ~"Onl'~~ct. :md .h~

km(~ 10 Lhruw in (he tuwel P(J{~ioisnne uf'a \/,f;,'IJ'Ym,lI~ Illimber ofd ii.'l.e<lse5in .he wuridl.h~. s fife' even cl:!l1d'kllPttc.s laT e~clk3Iltm, because ~l ,QlJe;;vn"l hID'e.1n anirm~i rese~F'!,'O·if.. rela~ve]y ,,]flcll cheap <LIld CIl;''Clivl.:: wl,dn.Ct\ ilir!.:· "Y~~IIi1h1e.h"$ alsa h;Jl'lliI,10 a~:glle 'with ~Iilesuccess 01'8 p~lr

break risks, ph.uggl.ng in eeal-life t1aw rlrOm

on:

~i'

20 APRil 2007

VOL 316

SC~ENCE.

www.5ciencemas"org

NEWSI'OCIJS
data ~rom lmlia's N atirmarl PoUo sur1.'lj!'illil!'l(~ ~rojKL ~(I h.oelliWy d~i l~rei!l who had d!!'II\e!'o~9td bype 1Lp oIiomyelilis, wi~ll QMel 01 p.aral~· ..is ~eu'i,le,elll ]997 and ,WOo, U~i nog adniriuedly 1ii1d!ir€(t FnrMMJfle5, lh~ lltlel!l wmJ~rfod how m~n~

..
, , ,, ,, ,

n~J!! r~r~1 dElLa f~!)JIiI th

u;!ll'lSmissi(lrI.

rn~W I! ~n on iii Ii!~W~f!Ji!POJl U!i!thll (i!llll,l;IlIi~1:i ~!1l 'It e[:;3iJj ica~!:' polio: If'lol'I.ov;!lle ill. oral P'Ol~ovi rt! 5 v~(dllii:!(mOP'iJ), lnuod,UC'E!'(J~n
imJi;a in 20050 illl lI1,~ hOp'e ,(If ~lnlppiing '[irle ilfI,(I~l jr.U8If11!;igli!,l'Il nat.lll or d

ThE!!g{l{li!j nelNS ls It!a1 tll'e vacIlne '5eem~ to IJ!i! ~IHe~tim,e~

b!!@1ii

mOhil @tfh~l:mn "'e' llilEi fIIlil@ ~riVilII@!lt \l'@r~l1}fII hhat hss

lI~ mainslily

Qllh~

Pi)lJ1oe<J' adic<lticun iniHallv,~. B'IJl til ~ ba d IfteW§ is Ul'iiill e-\l'erJl (he new Me is still a prett}' blunt instw menlo Wi~llOUl qu,eosi!ioTl, ,AlI;u~rt abin's O~r a li'llt' 'O!<luil'lE!'made lrom hee S
a~[enualled 5!ralf]ij;
rnisapPNIFillfice

,"i"" ,
,
tnm~l.lI!I

~li1"llirl"'Id'_·

~J1I

,oci1

()r POliQ1fiW5,

Il~ 5 'WOf~edW(lndi2i'S, le!adil1g til llil~ virtllJell

or polfo rrom aU bm a lew (OrilfrS of the wllrtlll hee Ililain t'e);t). The liiH'~ls hllflging em in fl~ruhern N'lgerial, whe.re aim arr~hl'aaine bOYUf~\ hi 2003: l~dI bJ 011hl!lQI: re~!IIrl:l~m;~ (ilI~~~,ti.lQI'I!J the ITU'£Iged Qlf' btl rd-er gf ~aki~t~fl a nd #l1~hi!ni5't~Ii!, iP'Setlil ~ty ind ru fl'KIr.s a ne h"QIIperill~
!:)c.e:>~ 'trlll dii,luren. hn If!dia. crowded,. 1I Ilhyg iMic ~eolill I.\! be pll5mng the bi G'I.-I).yic;all~mlils Qr \I'<J;i;l:fne. Illli!~ !@f1!i been ~1lQ\~1lI Ull~t'O:P\! i~'l as erf!ll;li~ in poor
\lBliI!CLrl"~Clf:$ 'a

ClDlHlili~fIiS
~~ri(.ii~

ranrty reHabl~.~ The},found l~a l H~ enicaq per d~se of mOPV wa&30%, l~ni'e times year, w11ell N](hcdas, Gras5~ or I"np.eria~ (oll'E1~'~ 1i.4miJon ilWUJ cCI,IlBag:I!I>IlS aJi,al of to I!IV in tl;~ salmie seUillliJ' ~H i~l,[jii'i'er Ilhliiilr ~(l'iJ ~liDl.Itd i;:(peCl and IOWoi:i,r~ham yew WOIJId lik'!!" bm i~ movilllSi 11'1tile ri;I1I d i~ec lion, ~ [.ay~ i~ f'l~iJ(IJtl!d a Il'!er~ n % '~Hii!:acy per do~ ~n li]ll;! :sllates Df U'ttar P~dlil'Sh a"d GrM'!iI~, l'eadl <iu,thor 0111 a !>tLu:i~ !l'Uhli~ltIE!d on lifll e' I~I. ~'i'ee-t illl rile l~lU:el. m!l:u f5ciI!IICI'!, 17 f'4CiVemlJer 20rl6, j'L 115l)}. NOWQllidt'!r.ase~Dr ~illti:i af'~ oc(l!rr~n~ li1l!ri~~n tn dlild Ill'JI who hil'lll! retl!~ved as m any as 10 dClses. of Paul Fine ClIf lhe lCl!ld@11I School ~1If HY!lI[81!l1! ~11Id111r'opil!:iltlllll:edkine agrees Hi1it the Rind ilil~$ SIilO\\i Ulal "~nOP\f i~a bUi~r tij(]11 fiilf lYp'E :I.. PQL~o. QPv,_ Gra!.1Oly 5~, "We A. 2006 ~u'tbFl\a'k in Uttar Pl'Clldesl1'pr(l\l'~ded a !l:ham:@' to CIISSi2SS Ih~ Wll! would! expe!ll tllat. ~ Iliu'l ls it g:oml ~ncmgh t.o lltillplJl'ammissicll'l! don't k.flo'.w,.. s.aY!' riFi9. -UR:., eincacy~r 'tine 11@ii\i vacdi1~ Olil lh~ 'gm ui'i~. Gras~y a lilt! oolleal!lu~ ~Qur&il
ptlpulati®f'I I.Ilr.i;l ttan~mi~ill!ll. B~! ~ l~lgs,whm an~ flladeWjuate salfi~~<lticmCGlf1llil1~ HI ~III~~ 1~~~~iit'~ dear ei<,atlly l10w bad thil1l[l~ WeFe-u tltil!a}~

set-

will;] gQll\llmflJlefili diil'!11en t.ow miny imm!.mi~atitm rounds hid been C@!1IdW:loeil in eadll area, anll!l W~i(h \liKLfw.~,. riH'ilit[)ijalent or liti vs h~1l~ had been L1~g~ .o1t'~ 1Il1;l~ ~ rr~~t ~~y~ fjJ~sJ)1(If lille !1lI~,hQdjjlhn IB, "b~ I it seerns G
IT

IJm ~. i ~~~.& 7al ~ ItI li :l~ .0 .. U Il~~e~. wses. 'w vacdn~ L~H! 2076 u~e~ .ijlfld age-Ilnal,hed {cmtro1s had r~,ef",ed. liihey relie1J en parrefi:t reca ~i, ifilltdaied

d~~ly

..

d~;Jli."'~~J.1Iiilllyre IlI:'Ct~ mudd. Thlllrt~ an: OJ lot ,~r~J.,~~lln'['IlilJ!l~s"'· she ~lIy.s. '':A..Hmodels ean b~ cl"Jtidl..oo." eeneedes Jdlm S~l;!nmm. Ol!. fellow mU~elL:'rOlt the M,,!c\\;<;<1'd'H.H;.C~t.. lnsuuue ofTcchnology's . S~.o3.~ ~ a }'(;,IT: Abr:nn S4(m milli(~n nfth,J!I '-."{l\,!:'ft; the Schnol. q)fMm~."gem.enl in Cmnbri.dJge. But tlrlis is. "the b~~I. Hln31)lsis. mit th,ere" l don't ~ costs of ~~fvc'iJla~l!.;X:', rouaine pello iImnLmiz9~. . 1Il t~(l!l! md l\'I.I(:I fOIL!!!Ih nf $L!pp~(;lm~f(Jw~imn~~~· .: knorJll,'" l'.my {ltllCr :I!la~y~i.s dew'i~ed ,an!d o as
UJ SItaR

\\'id~

a f1illn~l:!iL:t

~ \\~lCre We al'ii: wday. ab(Hll 13(m ~o~io "Is:eS;L!I .;ii: ~"!f;WIi;l .kn(:}\'" w~m il LOlke~~ Ik~!;!'p ca.<;CS<l1 '[0 ~ [~M rJlc. :saysTlwmpSLl!i.l: :lbtlLll $6g0mmioll

mndd (;N:li I~}ll r \1".i,t1dy !.ill!':!:1.1 ~,S~ d .. poae It1:uJkouion'" ~lrlimn~: swppiJ~~ rout i IIIi.l
LmU'I1

.ili1.it.:.al iom.un alllow-income

-coRnlri!:s. Allothn::r

~. S2RfliThilhoil goes Rv!' Ibm' m.J:t!Lliofm~ imllrm·

i~ ili1IZ,~limll ru~ml:-; in Ittld~1lIiI il: IoItel'lil," Thmmpsnn. i ~ .rurn Dla i~ltier Teibbe'lilN defined! H!li ~ orti,on,ls, d :.. •'~I~gre:osulI\r.:"1'1" ·vc~y hi!:i~ CUili1n:rul., PI"yililg il ( ,.

,; out over 20ye'3I'S. ~~s Olnn,.!] e,WOI'l5 \\iOuld C

~ i1X]]l~i ili1w~irldleri

!lul;eiy"

l~d tD ,,~\~"hqpping'~G:l"1
1~1c"v<,].ri ..

!ui~:;;io!lof!~!;: wild virus ~DIIS been l~iilDcd ~mll.iml~flg iml'l'l.tmi~trOll \'\r~lhor:11 rolir'l\'~nls v'.accine, ~~nliml.il'lg . with orv and supplememal rounds, nnd ~\\Iih;hin~ to th~ mo.re e:.",pen:'i~\'C~rI!a(:~ i\i;u:~d pu lievirus vaeei nc, These ~C~ narios ranged CIlfCfld1y .!f!mun:k:ll in '~~ll: rulli." d UalLlds dOlAn. thewiil1Lle(;' un ~I;!'m s of C<13eS .rmllllli()t!g~llysao million to S5.5 billion O\'cr ::W ye"I,j"l';. 11I0l UITIdudill:~th~ co.:;;l. In :JChicvt:i ;'llldl cmrs i!'i emd icmirm_ sn)"~ "fimrtlrprs11ili1"Tille er~Ldic ... :in!1l lnli ll~f: rim pJ'n~e. \','hich l\yh",Ol'1i'd ~sJill"lu~~ CoLlltl lx:anothL::r $2 bi~1~QilI. t The i:lulI:om line. ~~ys Jc's ('ochi. ~s n

~mmuni.l.:.Ut~~n1 after lr.mli

,. there IS .not

..

'Vi~hJ:e- tlmlnrtll

upliml.

ll"~ 'Ln

i
I;;

~ 01'$ 10 bmion. ~ Tl1"y l~eli. fi,d~tt:d w.ith i >C'ome up \ll.'ilh ~e';:>iHlggu~ssi'le

in.ul':l.tll'lil.at~olll

bks ~~) ',,:.omm~ SCeTl<:rt""

in:;., Und~rlh.~b"m~.!b(J!i!.es (Jp'li.nn-routi~.lifl w.lllh I\~)i:\'LIprl":illenl.~d nJlIlIllld~ ~ ;j)lml nQ(I~ubre<!krespOll~e-glol}JllCm:e:s ~ \iliicldy )lI(l<f1I""--xilto ::WO.OOO pc .• yc;m,r. SQnH,~-

ll~ll~ioll Ll1I3t}'OLI (,fin l,'tke your rOOt of!" tile g~:!l pt:,Jliil. 'll'1.d emie up ,and it. will be nn,[lt~h Ic~s costly and there W0l11 bc-~, hnge resurgtnc.-e," He adds ,h..l this .nl;!b'~iSi !il~ows'"\\~ n~d nIl ~in~\,':lbU(V I>n:orts., 11101 :;.hLCk tl.n~.\1I;re nc!.>d to SC~ the I11QIl~ <lnd oommitm~m ..ltd n rdsh:'· The '1'1.'01:SI. poss.i.bl.e optioIn. whi~h .~o;Q:ks
:iioorm;l't~i~g~~"'e \'\Iln!!t j~ ImppelJi~11l~pd;;!y.i5 wh:J.l 'f.l1loll1,psm.l t::!)lls '·W1i\"d~ng COI1],milm,enl,'· As she dii:fiCT.ibe ... ii, \\'hen cases d!l'Oi) ~:o ltlW Itl\'1l~!!'i lhey d.idill.20n ..~
they
I.

:i:\app lem~m~<!~ j3rumH!s ,.:\ '~ry 3 yCll~ :1i1.d a.n1.\]Jcifl~i:Cly l.!~gf!.:5. f ~~ emergem.:}' I'e:iflOll!;e \Ie might keep -:ase~ to ,-I·ll.Om) ;1 year ;:Iif1d~~t atmCl'~t:S~ biUi!i;l'n

~ .~ h!!1gin I!lern iddle:~~;L}' •.~\,\'0

!
'0

iI'l\'ei!'

10 }ror:.'IfS. inb.el1i\le~!i1:· :;.;.~r-;Thmv:!p~on, S.he thli:ln:umbr.:rs !)IX' proj,iJe'tr.:d ".~JlICS

."n;: ~pe~dil1g

r~Orl~m~ly ri;;l:'~

Wo :(:lll!ch on

polio <!ncliu:m t

C mUm~ C'~s.'i::li'Il~1i~ 1'ilI~Sill"~ "'Iow ,t'!i)~ULnd I~ ~ hiSh ~a~c:>.t)rl~igh ,"os:!. ,lnd Il.tlW ~!lSCS, ~lr
~ $tlHi(lL;lI.hing ~ ea~titlllt.~llu.il

~ ~LD1d:>htmld!iDm 00 l.:Jk!.:!'IiI~I~ g~1i:O~1,. ··Lt'ii. <I big

c\lnlin!l~sto' dr~~hl't~ in UUar P'ra,d~h, hllillia, desp~tl!'th~ flig~5t1tac(ini!lljij)f11 ral.l!'s in '[he 'world.

Entr~rlil:;h~d •.Th~ wtiOll'irulS

Ihtir aUc.lil.ioit Ii:':Jt>~wht~IT'. C::r:~~~ :rlt'""[~fgl:.'" ~rnd the \\ljirld rn»h~ ooek iIDi.·'Itl: e;'(pel'lsi\'~. ;and we bm'~ J110rlt kid~ P'lml:Y-I,l;ld.'·~h!.) ~"JP.'. "II:'\\~ do il. r!o!pcalcJ~y. 'Woe~y unt.}!\: n,oncy Lin an if w~j LIst ~lrlisrn:,tl~L ·ofr"-1U.1I51LIttlE R.O IlEI;lTS

NEWSIFOCllS

MIEE.TiIINGB,RIIEFS»
,AMIERI:CAIN ASSOCIIA.fION OF PHYSICAlAINT'HlAOPOL:OG~SrS ~ 28-:3.1 If\A,AHCH

PHllADHPIH:IA.

PENNSYL.VANIA

E:urOlpe,sn Skin Turned Pale O'nly R,leclentlly., Gle!n,e .Sugg1ests

Resc~i:ehcr.~hu'liC disngn;cdl ~ordecades ubtUi'l ;m issue Ihal, us on Iy skin-deep: 'Ilow Cll~ tckly did~h~ :rvr;;tnlOd~trrl h~lmlTli'i\'I.~m sW~Pl im.{l I::llI'ope ~cql~j,re pale sli:.nU!'~ No\YilI mew repon on the- evolurien td'a gene fur Sk~Ulcolor :;Iu:g-

:.ucUs of :;.1,:; in. such :IS Ilmlids ;!lid fa.t:cs .. -;.a,s 'P;deo.~nliumrol,[]i,gij.SlNina Jablonski of rsu .in State Col k:;b:';. Such receru .C"l.um~es il1l skin color
SI;I(l~V

thal6mliJiim1~mt:· Mil'! ~"¥~)I\'in!,!" ;;~yl'i I¥If)~CtLlhll' s

A
~lf

ituun led by gcnC'ltd.s~ Kchh Chenl!;'; uf Pel'll1lsy!v-~rUl[I Stolte Univ.t'J.'sity (PSU)I ('01: lege i

Me~i!O:~ nein ildei[!\hey fOHITI.(.i two vari;11m::;or~hc gene ~hm d~lli:red by jLl~I. one amino acid. Neaf.~)' <!~I fi::ic.ms <Inc! E~M A Asians had one ,ii1lh:~t:. whereas tj~l)li, ofdu:: ges,l:s Ihm E\ll·ope;m.lls Iiglueued lip quite ~-e1;;e'!ll }~. oe.~~.l'~p." ~p ()n~ ·rfool) to 12.mm )ie-',.I:r:;; t:WEm-nr""OI.~s Ehey iitiil1di~d hlld lhii;) other ago. T.h.~~.0LJm.LlLd!ct:;tL Il:lfll~.-slllllldi.l'IIg h]pu~h~ (Sd.r:·t,rt·C'. :lH October 2005, p, GO 1 j, Nonon, who worked Olii Ihe (~heflg: sta~dy eS~5 that modern humaas in EI1.fope J;;rl:'>'" :l'IS :1 grud~HiD!;_: Slli dI~(.It,(ll.'ec id:lii:cll9 fin.(I, o~n ~I~~r ab-rlU~4[j;~lOn)~al":;. ;lgP., f~'" seen as til!!;;}' '1~~.gr:HlI:ldili!tnncrthem 1'[I~Ul~ld~~"'lInd~r \\~~ th~~t fln-!~~iQ!l! ~\\~pt !-h~~u,[,lhE tlro~n~dil[k..:r skt~:;;,P~~l:;:ik~lll:Ib:;.u:rh~moru sUf:llight Wmldng~I'" a po~idoc \vith!jC:llidici:01 Michlilci Hmmnef<lt the Unhrf'irs:II}' or Ari7.(m'l. she
f DNAi,n the SL 2.;1..15 g~Il\:: ill"l1. EHmp!.lI1iIli~,. fricans, A As;~_ns.~fid .AITI£ricmli I:ndiarls, tL"~~llt.vari.nion,o; in the g-ene that did not C:HI~>I;' [l,J1i ng.shl! 'L:'.'Ik:~Il;IN;ld 'the 1J~~~k1: I1d roLl ~murili,lln nm:: o:fSLC1.JAJslfId I]ll:n:bydl:!,.;rs~q wU~Ii'G~d\) 3,00 li3\;~
j)[U

Tf .C'Ulry 1 ]-I;lll'pel:l(ling of the Unli\!c.Ihi~lyorVI~~l. Stil~[ ake Ci~y: "\l!,\_:; h1L\re all L tac illy assmnecl rOll" ye;_lrs dml modem hUIll~11lS showed up 45,Om} YCi.!r~ .<Ii~o~Lnd hnve no~ changed .l~l~l~~srnce, \,'!I'IlDHc this ami ether '~l,c'Or-k ~h.(lw~ ~!Ilat e cominne tn ch'lI'n;ge. ofle,~ w

illlthropoh,g]Sj,

,u

1'1. \'el:y

I'm>!. fOlIC ."

l

IGolrilUas Hidden HistlllryRevl!'al ed

,,.l\lthoLlgh goriUas are mil' clesest Hvil.1g eelativ(_~,o[h~f limn c:~j~1iIP.tmJ7~<.._. lhl!:ir t."Vohltim~ is. wauellli.~g of .. !1IlI}':1lery. There '1r-.e~{lr{l~tiils tl r g{]rjJI~ 1i~ iil~d IiHI ~ DNA ~hJri' w iM unes, Nt'm',:il new iiil,lld~ of'nuelear DNA from Ihe tt'I'O spl"'Cil!:S ,tJ(f\'liM g{)r.i!~~". nl'ri:'l1"5 ;1 !ill in'lp-_'it;

il"!o;

CI

o~~their mysterious

p~I!>1

:mll oJ:'how

U1.t.:'\'>'

~l~ned th.u ~&,000 ye:lfl1 had I~Js!ied since' fh~ Iight-sk iIUl.li.ell;) \'''3~, fbled 11'1 molreanl'. B~t E
till!' errlff !li!,~u-'~i!1:;;w'.lreuIl,c. h
s,p )i~1e .L\;{:J.• ..

SJj~~d~~ iltfpdm:uH_':l> arise,

ma-

Hili ike I he iIi IlOlilKi as I.he ·t;;11 imp~. h ese t ,,,1:1)' hdbi'wn;~ turn O~lt lD h1M') l1i"'erge~.
slow~y into lWO species, ~ppur~mly 'I~Lkln~ th~ beuer part 01' ~~miUion}'e'ilrs. 11(:col'Uing to m. M ~kat the rne!!ling, I.r;i' ml.)·~ecu~ifu· Jnthl,,(l-

lyzcd varistiun

in rhc DNA Il.alilking thu

gene. She fnun!:! t~at [!~m)'p,e'lils with the :ulkl(J ll~uj a "striki I~£ Jar:k 0 f di \f£rl'~.(Y·· in ti1l~~ nankin~ DNA ~, sign of v~ry recent gCill:D_k ,C:ll:lLmgl:, because nor tituuglll time IU'I£ passed for new ruuuaiens to arise, The dtlt.] ~Ll.ggt'sl. t]-ml. the selective ~:w~er nccurred 53{10 to ()om)' YC0I':'"ngo. btu givC'1iithe impre-

'PoJ[).Iili~[Lind,~ Vi!i!'.~~:m~f the M;lxPi~md o

l.mstilWiI.t:or Evoll!iil.ioll~_ry Amhropo~.ogy f illl. Leipz~l!. Gemlllny. "'.Il shews us lHJII,'I.' liule
Wti"\I~ kllltm'n ~LbULII

speciation in

J;!_.tl:r:ilhl...,"

stl}lS .!lJnlbropolog~i{;a! ,~enetici~t

Alllle Stm~J::

d~,iml OfmJ'ICDhr.!d.[hc real dt:u..:could be a~r'lIr back <IS, 12..(J{Wye~'rs H2OO, ()f~\nl ~;ltd. She N added tbur other, uaknown, gem~~ pmb'lbiy alsceause pi!llifll,g in ELlmpCiltlS,
L1'ghten up. IlLII]QI1Q (or p.i:!l~sktn ~.\Iffl~i lhn~U!l1tl

E,Jr.Ctpt-am. r~lat~~~ rNMt~~. th:m~thlrlk skin, 3.1. ~owilflg U ~.t ~L\'.i.ol.(;~ ~ to r ,my l'l]'Ot:!~!cemore \,fU!1;HlHill for bone £m~"'th and :[)

C<!l,ldllf1i'l aboorptiofl, '"The revolut[on 01:1light skin O~cl1ri!:l;ld !Q!i!S"Ih:!r ~he ~,rri\,;:!~ fI!'mQd!I;jif:11 hum~lt1i!> El1rupc."nltJk~cul:.Ir anthmpo!qgist in Ile-dthe~' Nortull oftl1Je Univf 1.'Sil.y (l:f Ari2iO:ITIa.
Tuc:Suli. s,OIicl! in .1:J~r Illdk_
1'il11'i'i1Ui'J.skin C{1·~m·ll

The ge:~n:l~(:ori gill of the $fUH;:t ru m Q:t' halS. !JCCI1 Onl! nhhe: big

IHI.:r::ll:les t,F bitl~og)',R~~"e<lr"he[~ I1IHJJ!e a ITWljor b~~akth~'{111!lh lOtl:l ..by di_~~(W""--'li!l~ in ~L !;;l.:I1.C~SI.C2.J.H. [I'un :l1)1'!i1u..::inly C:llllS~:!> pille sl·dn. .il:lrm'l:ll'lyElIrop~~LlJs_ b~ulMD. i~.lA!oinOirK

ies sampled tlflO fewindivi duals to clear ~~p the COlll fusi 01'1. s fMy~eVil iiJl io·na ry bit) I ogisi Mi!1:I~e~ J~~~~~p!-~·e,;;um<'!lI1.,o:f O{L\!Itll;!;o;n.e U~~i.. '\vh~) Wll:fl: ~I-!nlef- g;)!th ~ver~. J~~n!e <!lot! rs. ~ n~h~[:>. S1lfV~\'cd on irCaJY'i1tt~utl:enil~n~(1:s![if YCrs.i.l)' illl Piibibmg,h, ,~clUt..;yhrMi:ia. s \'itamilil I) In thdr d~e'l. 'But ,vhen fm:mil1g V~~iIan l.;lI1d her 'O~ leag~le5 i,:so~.ned DNA It-aliI'i ~Ii~ OI.1d. ti,\ler, tU li;:t,~S nf ~.~ ft 6i!L:' iii :spl'~.ul in th~ j'i0-!:;.'t 6OUO y,~o;JP's. he .~u'gl,Hl'd, EUfllpe.lI1s h.td [e;,\'ef $.{lW!IFCeS vilam~!l D gf ~A.Ua{JI \,vihlgowiU~lS len nn the phu'Iel. I ~. iltciud:il'ig thl'l~ ~astl1rn gtJ:d Htltl fi'nilll Ugm~d~li iii! thdr road ~lnd.IIlil:t:d1l!:dtil:! absor.b irnl0j'~ ;iJ'I'ldtille C'(ln~Q: ~he WC~h::l:Iil ~'!Jriiht:, \~CI·t.l .~ SLlf'Ili~~l tn prudu.c!! the villilm1111 ilil the~r $ki n. CUhLI mlr.LL;~ars st~t;'h 01-" h!1.l;wier ch:d-, chiif;!'ny from ('anll;!rmlJl_ Th~h~'Illll ~eque~cl;)li ~. 14.(1)0 ba:J>";: ilitrs C!fnon~odiI1~ ~and dnl.:rr~- :~ P ing ~nigJ'n1 .r~Is.n h~.vl.:: r!lvurla:d iner!.!'::iSl!d liJrct :pre~Ii!!1l:l.bly liIol 1,IIJd~r s~k_"C'~il"Ji!1]1 ~:llIdcar ~ ;lhl>orpt~mJ. f !i.~~:lligl'n lUI Ill!! l~wl.!'1'irm~l!!cl u
'.:1'".·.

Eidll..:r 'I,,\'''y. lhl:limp~iciltion it'l,lh,n our I~LJ roret1(,n aaeesrors "'ie re bl·m'>'n.-sld~l aed fuu- leu:" (If Ihnus:mds Of'}?e-'.W5-;1 ~~!,lge'~-' tien iinli:ld..:.3tl~dr~ ago by Sto:!! lI'o.ru UUiIiverU ghy geneticist IL.luc.n Cavalli-Sforaa. He aigll~lhal the ear~y limll1ligrams to EtGT:upc •.

(lrAirizolln Sm~~ Uli1lVcr':o~L)" Tempe .. in Pl'i!'\'iolll~ s.~mli:e::;oFlhe "mel'nal:ly inher~ il;ed y ~hromosomi;:from J!:;mUl'I!i sUM!):eMed lhat the LWO speclcs-e-castern gonlli~:!i 3.ml WI] ~:tll;'r:~gnriU'l'~-i Il:tll<rbn.'XII uati I race mly. Bm ImHell'lIaUy inherited mhoebeadrial DNA suggested ih:Ul they M~ff)jJ·mted~1Iore th111i1I. miUoon j'!:~Lr.'iagu, Nl.lelcll.L DNA stud-

{J

20 APRil 2007

VOL 316

SC~ ENCE, www.5ciencemas.;org

NEWSI'OCIJS

RiI!Sie!ilHhers seldom :!Hi!!l Darwill'li~HI I'!ilh~!o! I ~eleu~ol'l Iiall'li'ening in living p~op~e. So phy'5i{ill.alnl:hrCl'~o~CI~i'Sl Cynlhlia Be<all W,Hii
d@HgllltmJ in 200<1 'I\Ihll!WI !ilw !!iis(iIJ",~~'d 1J('";jl!r~ 'till a l bl!!l@~ls.
11

IIIre lee ts a g:1!i nst mil ~iHla a fild hal.a Ali:ll"SS r.llill Ii) ~ 1:4166. SUI e~ilulll' how do lllrl2,e
WI[JfIJI@nm'i3Wl1l'!lI<eto urry 1(1)l~r.l

til 12 surviv,at ~f some nbe!t.i!IFI

rh il~ f~,n,a.p fI<I r~.nU~bV r aJi$i 9 111~ l!rv~1 orr rII

O,i('.l'g~~rL their motheors' li:m.l!~~-il ulIliill ~ ,~d"'~l'Itill!i'~ I1lJ1rfng Wl@gniU~(Y 01 k:iklm@~~f5

.atlCIVeS~ le\lel. r:Ilow!!leilll has U flliI,a IFd ~e[ stud ~ ~ ufllmi9Q' a pessible mecitlal'li5,iIfI for the a.clal~ta~ioln .~nd by OOtlllrllll:! !'1~,ing~lva'l Uill:! ad all lalrolf"l repl'eSfinS, same 01 '11:1 mOiilg,e~lne l.tl'rlill :se~ectioI i ~

exy!)en in their btl!!oo? The:y dt! nat prO~i!J(f' more' 1iI~'moglobin Ule Wi3i~' Al1ItI"<ln~ l1'l1'irrlga t IIIig h ~Itil:udtt: d~" Orll~ PQ~51ibil~ty i. [11al [he \'.iOl11feFlwi ~h hi'9lh C)xy" gt!1'I h av~ anadaptatinn Ihat Beall. is~ x!'DIlOr in 9 imlie'~,en Ii" er!lW iflln'Ese same Tibetilfil villagers .• She ,ItOll00 th,all some
'o'md~ef~

ye~ me-<l'5lHeti in Ilumaals, tier te,i1Im is sh@wing imw 11g,~nelic trail can (iti3i!1liltkally ll'nprOVe ~!Jr'fivo1ll in real lime. in ~i,!,j[l1Q Illwlhrs, olllid
ba,bie~... ~ love Ihi:s \.,.~rk, • s.ays; Mark Glaldwin, ·chief ® I tliHlvascu lar medlidne br anch Qr the N~ti~'f"!i!llHli!ilrt. hllllg, i!l1d ~~I;Il)d Iln~,!l~ute il[l Bel hnd~, Mill rylEl nd. .,Ii 'h OW:5 yQU ll~'lii d a '~tr!;lI1,91 ilI'tl'ElP~jv,~ re~p€ll1~e to Ila'lt~dl i'llilr~ilr sur-

uh<ll.e eJdr<l nHric o~,d eoin lhE1'i r hrlfillh. a :51gll11 Q,f

iilllllOllll1ls, Df

~ddiliQI1I.;l

th

!!;i.i!S

'i
'
•:ijj

i ..

I i
it

:Ii

L:lxY'£l1!!11their ti!lsui1!~-wi't~· to Cn 't~p clf l!lie wii:rl!t:l. TibHailis with olyrg,~ri'J'l(1'i IlI.iloOd' l'Ia~~ womel'!. arul ,[hi Idn!!'l livl fI'!) in mer@ I ha,,f! tlut pr!l!ltlt.ll!:IIn.!!I Ul(lrrIa h~!lmJogll(l- morn ~!!IWi'lfll'flg d1i1l!ire.n, bill IH rai!ling the; hl!O~d 1!Ii'~$.90tCI l'lI[)IJ!"ehollh I,n 14 1JiUa'ge!l 1111ihet A!'l l !>un~ifllheir ~!JI!Illt;l!,;. ifhat':s ll~e Ul!lEHU' nd whaL !l''Ier. !h~! ifs, qlJil@ pl)ssibl'.l' thil! thl'libelarls ~ilrlorthi! study, Ih~ ai;J;olrackedIhesull1l'ival have e'o!@I'll'EHI more I hiln {1I'I1Il way l@ boo!;~ W,ml!f rmm ®~y' Df clli'ldrrtl!1II horn lo mOlhen who Ilad Il~gl' happem w~u,n mounlail1~,E!r~ h lood IJil:Y!lI!1Q.a!lld t ~t t[h.@,~ ri!I r'~ n !IId'll1J@41~i!1iIt g@1l d@'fi~iii!IIH:V: The b:loQd .llf~Y.i!J~e! illl the-if ~Ifld I@IN !liCV!J@ ~alura lite!'! irI ~!lei r b'lolJd n iiIIdapta [iolll~. (i la clwi II :5 Llg9 ~!lt~ lhat flea U's l,un!IS rT !>~$, til ~ Ill.(jj) iii ~.e§~'I1lsUIl! s,lrT !:;I. <lmd (!5del'lce.1iI0~MdEnrL)l1iitlag.or'9/~g,ilulii~ellillf u III lea m (lbo me.J:5,ure nl lrie @.ifi~1! and blo@d flu id !bui lids UP. 5 un ®I!:ilti rilE!' l!ill1lg~. 1i1~ 2001lM'la14L BeaU rel1cnne~ lin Lh~ me!"lin~ in Ulil't ,""OITl@!l wi~h high ~!N@ls[;If oA~g@ni!1J til@if illl! nul ~t~JII, sa~i E!l1!o1!ll, h til tf~ lill~~e pres5.l.UE!' the lungs filll pregnallu WORIlell, who ~tr~Mle'J' l~@5e' [\!,!O lin!!!!; ~d reseran;ir1 mee\. Sill!! lIIf1! und ~r t hll!!liJIoSll!llny.slollj]J!Dk~ l $tn~5$ ill {litib~tJ~d had mlllr'~~lhan lwke ~~ IflH11lW sUMving tum e' an d pre-smDa 1J1~WOtlk! bel1lrefil iIfIOSt {mm ],',Ii!ll'!l!l1lo HI1Iil U~f! uml~D'lying !!I~ne bl'hi nd lni!! balllie~ as h!!liJ.these with 1M !l.'(y.g;eille'IJe!ls----a this <ldOllp~iI~ion.;'SIU'dy l~e pf@g,mmt women. ~ wo, m'lfl'll \ &1]'9 h· {IX!!, g:iJ:! n bl(l(l(l- am d S!1 e riltio of 1:0.44. lhi,!j ~6 a '>lilrWn!llv~[ml1Jg w!Q@clh@fit 15rl!?lilil@,[JIl gerU!1i thi,l! f'~91i.'Jte 1(!'Il- hr(i!Si:!y~. "h,[ri!I UiS€ lhat 's whiNlf yffi!~'Il sse eve~ 5@l@dil;ln 1;DWE!$5lllrl!?,.sh@ Si!lI'S~1jf?ns~rllm!J@r eni ~t\,JG. eh of n iUDI!: xide illl 1.11~llll~d. ~he notes, hlinW lu ~i n atl:lien, ~ o h lIn1l,:1l IJMl 01'1 t.nesiclile (ell 9'IiNle, whiICh 164.0(10 m:no.onu yC~lrSgo. whh males a m(wiJ].~ more llunl fe~l~le~., M~yS Vigil,mt lhi?lfs surpnsing because today "he I.WO !iJl~i ~~~i Ul,()O' k iIQnle't'efl'i .,p;;:J.n a~d~n()~t \i~ ~ori II ~s 111C\'I!..'T YCn~ LI:rC 1[~11" from horne .. ".1 rind i~,iI!m"r:::i ng.lh""t g!:Jlie n(ll\'~~ persu:;~,ed t9r IllmJred~ ofth(lmailld~ of y,Ltar.\L'· ,~h!2: nys, ~ !-Ier st11dy QlfC'~ " mr>e\'\'i~dm" (lH .~pe~iation in ~tr~:;+wh~ch ~~nrclO fll> "11 Itil<tldcl~br

vive at high altiwde." hail, Qr (afOeWesllrJn R!!s!!f\I1tlI Unl~>ef'S_i'ty iii (leVi:! lan,al. Oh to, a ni!l ~'I~r,coUeag lIi'El~ have! ror 1i d~C~,de IJI2'elill ~l'aIU'li~Filn9all ~lJrts Ilf ramWj, bil!!',llIglul,~rr1d f~rliUly ddlta fum men,

in thei[ b~nd. In these Ti be~i3iJ1f!i., itric C!:!iide ~ilille~ n th e b l@(lid v~n.~~$~Q t ~ey 1);!!I11 liI!.!m~more bleed end @i"i1i!l~1l to or!Jilnl~ ailid th$u~~. ~~ measurlld by illl1,8'g'r£l' of heart ,HId hmlg bhuld vessels, The T11J~LM5 calill bOO~l 'li'le;r ~l(Jod
",[fI'lun'le-jlll!i:I
$'0 p,ump 1lIi!l!ll'e

ore hounc 1.0 ilfHh Chit'llfP5 amI. gtlrm3~, Ahh.tu!~h chim].l'>ln~l.~s sorted inm 1\11.'0 species mpidlly. gOl',i,~I.. s,took nml;h longe.l: IG; odi'rCI':l:i~y ~Jtchim,p.<lJil7«s.bu~ tWIce <Q:5 high as, hl ~ til ~ in hlU1l1;~Il-~.\";Ilf.l <l~·C[rfm1,lrk~ble fu:r~fiu;:iJ,~m:k '!rId Ji;.~(lntiI"l~n:ldl fl1;u~.d Iow ~.~ve~:s ~n .~ 'fJ,:f "'-dri:aliolif, ··n~s'~h~\Y;:;; ~:.; ~ 3g~LLI'I, k~~rrt<!,dI hmlil<lllisare. :.1<"1YS Stone.

i a~ 79 difl:en:m
I
:i
:

DNA. :I:'mtu each gur:i:l12l, :[)ispa!l:'.II.ics l.ume~ 111 p :;;iles. shwil<ll' ~o ~he genel~!c

VUMi~a~H's
C~I~cu1.OIte

~ ·~t!rnbe.

or ,g<:'ll.eti ~ ~~i n:r,'r~~'o~~ w

iI :t1lILl hli.kro.l

l~,l1llm

~t'(H~

i.h~

! alim\oed them
I~

i[],£, ~ 1:U'~~ni~illl SPL'.C'i~llu~n'ihJ' Ilbpu'l

.0 d<'1:tedl,e timin,!:!

rale:':, whi.diJ

~ \l[lU.OUO to ~ nmi~li.olil y!l:~r8 ilgO. ~ T ~a~];: j [l!J;l W.Il~gl lllL:: ~'Voij 1ipc(, il!); ~ {),f chiln~ ~lzt'e!\went Ihei r sel};)r~w ~vtllu~ion,lry '!."'lQ'.s.. lO~~gg~~tIi iIlg l Ita I. dUI il g'l.:S 1f1J t'i.il f1iUI[~ ~ '~n)ke lip .he' dense Foresls tn:l1

undenj"md:i ng human <t;lYoiUlio'l1:" ~;lyS Jel'i~en"Se:ull~lIii_ Utilil nm ....,n;::.e~m;:h.ers h ~vc CU~t;t1 rOn t ~c ;q li k<k ~I)i iL b eI.wc~n :>,]fI~~im,of ~l:dm.p,1~llZ~~ll In, f1it'l, s~lys ..

r()

Jell!;~n-Seaman.
I[ast m,e't we5t~ (as~~ni

c<ou~ins u,u~ roec::e!1l~~.

IIIa,ri lIlI,~ (ri9h~) bred wit n ~h~ir ~B51:erlll

~llI1!d"lbl''th

~[;'l.n~

n(j"'~'-'~ ,drOl~"'IiI"oU~ IUllJ,g.

-.A[f!IIIN 'GI~BIO,f!IIIS

~lniU<l$' "::;~opp'Y back-

pnloL'li.:!!:s," -mii)'

i:ic [~kflO l".In.

\l'!llAl'w.sden.cema g, mq

5ClI:EN:OE

VOl316

20 APR Il2007

The Simp/,est and Most Effective Way to' Analyze' and Graph DatrJ'J

T
Exact Gr:aphs far Exact .science

1'1'. '.l<I8IIIolII!!ir pi1II\IIlII"Ill\. tut i'la'I',U'II'!'tlY' bean I11III hi ,~'t!><> ""' ...... ~D!~!g( rbof!.!>nl!:a! 'IIf'I1!fut -.:I,,~ .. ~
ClDlfII'IIl1t.oi,'idJ'i

5iglrAPlcill.._N.

IH)1IIIIJI:lIIm

I THIS ISS E:
,

tlal, 5~gl1lm(ant ted'lnical h~lr'dles st~LIneed tn be ovelrcolr:t1le'. Illilth~s~ea,tur,e, e iI2:Xi3Imili18some w

the recent sdel1ltHlc advances

tem cell, :re;;lea,och Is movliliig

rorNaro

rr'a,pidly. es,pile the ongoing @tlilfca1la:nd d

sdel1ltiillic: debates, ira truly tap 'llh,eir peten-

orr

future, See page 463, rn this lssue lIP(OM~NG FEATlIRE5~

wh@m the tedlrHlI~ogv pack leaders may~a.ke'us in the

at

and lock

lune

1,-

RNAj

eel[ Signaling 2' Auglllst 24,- Mi cfOana:y Tee hnologtles lune 22 -

LETTERS ~ BOOKS

roucv

FORUM I EDUCATION

FORUM I PERSPECTIVES

LETTIER.S
(NUl l and the ~3~i.ol'lalSdelll ..e foullI.datio:n "

rhejrunwa ..rering ruorul urnJ fusc,d Sl!PflOIil OIl'l~lIr tlme of g,retl tcS;! n eed.\Vi!hil1 d,lY5. rumll"LUnlI:'aa.O{]:rI W.lll; e~mbltt'ihed Wllh 'Senior <'Idl11ill1ii SU3'<lI.·~,ar I:he 1'>4.111:. indud:i ril£ W,E WISH TO ~ETRAa R RESEAIKH ARTIClE '"m E MRNA. OF TH E AJMBJD'OPS.fS· GENIE Fl1ll\OVE5 Di~~:tor .r:~.. Z~!'hmll~i'In~liDI:~~~!J' is Ui~'~l:;tor imll ~c-.,h~ 1>hO!J1 :ip'l.:"~lfId .LlldLL'l:~S flrnvl;rJ lng''' ~ A n,er ~~C' f author (T;I-I.) I~ dll:l U:n<!."i'I. Norka ,RtliL-Sr.m~. afHJl~ithe NS]~ Dlrector Plant S d~lJ.t:-eC'liill!lre for ~mmhn pUl'iiti.{m. ;Illllllir-is arhi!i unglnal dntu rev~' led !;1ii:\it~~:allllloiUl- An~1;n Bt::L'lIhml und Dl::rl,l~y Di:r€t~t!Uir K1Jlhh:OJI~I1.llolln .flg.eflide$ esmib4 ili:hed points of 0.11]alies, his <'iPPMe.l:lllrnlTIi these files that data 1~'Ol'II1Ilhe real-time RT-rcR \\''e~ analyzed incorre;::ctly.Cert a ill d"'~<lP'li~ts. Wl;~ ~mmr~l \\:!~i~eotherd'l'~r.rJi!1J(~ \,\~~ ~j\"e!l~n~;;L~Wqigh~ iV! t,[l~t ~n~~pFIC~!.lre-~ to :Idd!ll;1:s.,,, ![I~lr !le~d~H llm~stali~it~Il.~ mm~t>is.Whcn 1111th~ primary real-time [tT.PC'R dai.i:li!.t"csu'iJjl..!Chxll. it) C(JflU.::'t 1>il~· F ive scr~.t!.I1tN.~ ,jjm(:~als\i.i sik'tI, 'fulum: U111· [ ti s!b::,~i mKII:ys ~3. most oJ Ihe reported sign iric"n~ dlinerenc~ between Ii 111e POillts di s<lppoe.n:. ,!ersil)' :md ethe r se'Ie~l.y OJ Ilec!ed iI~slitWl~om;, Bem.ust:' ofillh;:, WI! are ~~1rm;ling I],'~p~f)erin ,i,t;;~flitrdY. h~ ne'<\1experlrnenrs, \'I.'e 11:!J\'i: repmdll.u:ed 1:~leIlomt indu1;:tlrfF! caused by a heat-shoek ~!l:d~ti:- 'i~Th,esllllliftantil effectrue litlil elf FT ill ~ll£in.!!l(: k'a.r; biJ'.~"1: ilIa'l!!. 1~.i]~t1lud!:'t~t'[ ii.1tWt'iililriU tl f I.ll~ 'lflltit'ig~tijc FT IfllR.NA respDr'I!!l!i,e of tlh@ ,Blcaid:slmic [1'('I~1I1~e;lrlO shool l;1pe,,:.We ih~l"i)rCil'e 'I,'CM:<lJO:I cl[ll'ldll::>iot1l that FTmRNA the i~PI'lr~ !lfihf 11:1am! loom mill nirty and gDvernment i!l1d~l(:[live ~i!l,~.d n];{!\'il'lfl ~Wllm1L;',,~r u $h~11]1.,~pt;,'I>. L \Me deeply re,srct .111)' )1ciemific: rni.'i(:{ln(:€p~i(1!lls I~'ml h,we re:i1.!I~ed rlltln;! the rllbh~a4id!'l 0:1: ,8geracte.s made it possible
(NSF)
rOIr

au

r~r.;t

Lil!ul!&i tRaila.

The 11M tmthm ~n~e P111'C;- ~TJLl sU'olll£ly obj~lj;; tn (he'I,·~tl'<!~'Li{)nthe ~I~el' am.!hm; of lli1.c[llllbredee lined tn be ~l!i1.. ud']Qr otlhe rulr:.. c:~vo:n. Our reJ~tl~J SC'ilJflUf Jh:pui'~ u.L1Lht: COlrll'i..\!;lulmory moolJl,c in trees ~1l isll'lot 3Ik~l!Jdl by liuiiR~Ir:;tl~itln"IU:l d'lis P'Ip'I.'lf. T.~~]"\\I"d£ i.ln'llI~'L:.'tI.~nli1('lL:'m'il~ul'l~'iJJ:I arid unla~Y:i.V5 1 ohhc Pff ENLl ex peril:llClllt\j, rt:fX~rledi iUllhe' SllPPOfILi ng On~ ina Mm~rkl~, These dm8i htlv·e been ree\ ~ IUfLwt!llln.d
Jhu!1di to be Cr.Hf.I:eclly 1.·~poL'1!;;!~L llii eNIRltlIBlijH~EN U!kS,i SVIEN Elrill IG$,OlllJ,1 I'RAINiI\OIS PAR(:Y,.2 ,(I'liE NILS$ON1

to preserve much (JT the Gompetibcve reS881rcil

enterprise

at, TuianE! and

provided moral support

for t'heiir colleaquas in

New Orlleans,,'" -Levy

oe","!'1Imem\ or FO!'el!1GeJrle'lit~ 31"!d~~ nl. !!\h~iclol!l~, 5wedi sh i!J!lLwr:silr '01 'tgri(uH~r"l ~'E"'Ol8l !.I'm~~ S~n. ~l~bOr~t~ire '~1i1 r'W~iQlQ9i ...C~'lljJlili re ve.g~ti1~. De~<Jl1ernlii!lit IR~.~rn<e. OyJiil111Ji1W1: ill Mauch 20Uh 1('1:,l;:5eSS tine cffcr:th ~IlCSS ufthJ.! e:~ CIi!IMill~ei jIlROOPCV!!.IU!I~[~ M~~le dill Rec$;ert~i!l sua ~~l!WU l~i~t !t~54!a~th tfJ11tl.(enir.e; N~ltonil~ de II~ IRE'!:h!lIr.:It~ 1'..1'1H response. A. ~i.i.~e by i13~neJ;ll ill Aflli~ !.i~il >c~~rrtiflqu~ 'tCNRS}. C~lItmls$~rliit. a fEl1I'!rgie llt1l!rnlq,iII! W:I(I, Ilm11t11!. N<I~iOlhll.ilh! ~a Redln!r(]ln~ A~rooomiQul! WliftA}. JO(l6 W.rr1:sf~~SO vcry 11h;,nuoill'lg ;IS weil"t: ~hl: fi:ll UI!lw.e'~II~ l~~.nfI,~,:n ru~ d{!~ IMi:in~. wl1m6! (2·31'10'5.11. Grt!MI;[e Cll~Eil: ~'. Ftilil~J!o. F!IIU
f,(j....~rE<~t~ Ce!ltre,
)~ie~'!ie1,

l!Jme~

RIlifIi!~~n~~ 1, 1_ H!J~n!J, IH. lWihit~lliil,Jl:, S,IWiIt!ol!ll~, Ii, f.a~ Q.~, :H, ~h~!'I!I!I~ ~ tl.!" :S-d'e,!r.ce n'. ~ lO~(!' ~~),
Ni~!il1if\,

S!J'~(~ :;I'1!!9.~6'~ ~OO5t

A Thiank YlOIIU fn]lD1 'Tulane Unhf'ersirty

n rlAS ~[[N 19 MONrrHS: SI~(E Ii"lUR:f:ICA1I'H
K[Lir.i.iIIa S1.i:l.I:til'l'ilL~into thi: luub;i;Ull:ll
Cotl .. t.~~na,

grut~'tud.t: 1.0 lh~

!icau;cuni£
WI;

l:il)mlli1l!ill'll~ty

lor Ih.i:

streng tmilPorf lh ..t i

t.!,;;;

received brillil ,~lUl'iflg

~d.\';~~~

~lldl31Ilt:;.nh~ storm, M[[nyit'l~:[illUi~iOl1S

"oum.ims widc!jOpre<l!Jd~slnJI~t1k:m and Pl'(ll!C!:n~d

110lldin!! i:~ New Orlel!'!:JIi_ GiuKf' o[mSl~I'l_"lill!ll~ tiOI\:;'l)filrLiume ttl d~!I~ \~ith tlIclIfldmmh o~.·dk:
~'ii.t:tI'~lt'I:ioc. ut gn~l pmgrus:,;, h11~ b 1k"t;'fiI1i:i;td~ iln l~e;~I<~bll~hil1g, .1 vubmlilt (:om~~!:.'lili\,ll! resean.:-h I;tmrrmmity. At Tul:l!le Ull.ive~ily. Wo; h,we

,mdi ~Q:m:l;!n~:rresearch I~tllty. witli1 :" p3Jr· I.icu ~t.jr1y~l:trgcIlluulEr loc~H£d ~flj\milil~'f.: of Smlt~lTexas Health Sciences Cersers irlsli,lll~
ofT~)i.l'i:ll

Hcnhh Sch::nce Cl!'IIU~:I·.\'I,I. HO~I~ltm. T~,.~u;A&M M~:lIhh Sci~ncl!: CcuU~j' &-h(}ol

Mt:dic.in.!!,. Unil\CL$lt.y of r!!X~IS Medical ]3:r;uu;:h Oll Gah 'e-';I!ml ~ W!.:'(miC' a s:p~cli:'l.1 ddJl of gnu.jHRI!.: tu our
or IJ~

tions I B~lm" C~Ii~ge {l r :X~l;ldod lrlii.i~ ~~. (.i1iily

many Ki~tl'[I1WHda~.cd ~Wlppkm.eub1.s rH'm',i{~~"(ill\l N II I all'ld NSF gmnlt.'I!:.aL;otfl!Jcmd iML.ilUt~om. fhmUy.we <Ire \I\t;.... y grJLend to the US, OmiCe orsc;i·~Jlcti [ilid T(;cl'illohlgY rn~icy and ~I.n:: ()nice· of Managem,cllt and Budg!.lL,whlch i:oi:iued ;,I. Joim Aunmm~~nt::!ll ZOi~!ll uft~r the Morin eswb~ild!l ng "'l)eci;j~ prlXern~res. for td: 1~[{;o.JilL~Lil1,l~i~J.Il,';.,. irlti.:lltdirng.Ii'_\lI)~~l'iiml~(lfHjJp:I.ication amJirepl)rn1~~li.i,l1i:mem dC<lldJ illeS, Tl~e !>'!Ivin m~d effective response of the ttc-adcmic {~mltllllirlii)' 1md guv~m1.nL:ffiltIig,Cill· cies madie ilpm;iSiM~ to pre~~r\c'e ruaeh Or~~l~
~i!.llnpet

uiiw re.M!l'Lrr.:h!o:nltt:l"pl"i~~' Tu~ ill~J .;,ii. 1U1C

jjl:rovided n1(lI1~~ .s~upp{lrl For lh'eir '~'ol'l ..gtles e i~'iN ~;",'Orlt~3ti!i.

lA!ll!!lA.S.•LEW
Pfof'l\~~~ til MI(lrill:iilllog~ '~I~d iitiil.Ui\Q!o~~. Al.~d~le I :SeniorVlc~ l'I~iiI!~nt 'lor j:J·~~~r·tl 'flJl'alie !JIfll'lEr!li~¥ N'ew •• Of~::li'l~. LA :1'01112-2709. iUSk. -

~:x~rh:nc~t18 ~~vl:lof ~buml~, 31t~ t'('I1L"\\I[I~ heyond O~~'I'expect.~u·k'ns. I write w e;'I;,p~e!{.s

lea;m.'s

Oil 11')01;1' N<ltion<!l~Il1s~~Iutc~ of! ]e:~1t1l1

LETTERS
egg.'i \.~here\ll'f the rules a re mo»'1 k1'i. This prn1>pL!-t:[•. :1i1d ~Ulllcrg~rng ill!;tlL'Isis I~_n· I;;'ie~.1!.ri!ll:lrl~ lit~~ndard:.; f[K :;:t~!T!J eell r~~·;!!:!1C'h. polm 10 ~hc .IlIW~ for bimJill'lg. ru~es UJ CIlISLI re 1"hc NA.I[uro~, and Ti,.::tin FOCLi-!' 'Uro~iP."hill\lC are ;~~~(~ I~mvid!i;,ldl lIIllln! to mission 11~;~Ulm.")j$ntclya th<ll stem cell mal ol~,e-rhielcchoologic{; ~ i.ulg In,tmf: upporl1l UII:ili,cs_u~ld ~il.lt:A~,rmi!1O:iny iJm'ClupL:d;:9.l1d 1I~. in \V'ilYS lm:lltndy SlUPiPPfit. focus Group has provided 311;[Ina~y5i:s 0 rl~'nl!! mther than aCLll<lI~.y undermiue, hl!tlhlu.end \~'CH.oo.lng. [Jsnobiulogy pdlC~.lti3] of the J:nrn;s 'Webb r;;Oll!ll'lnuni~' inp!,q~to NASA:, ~ll.ar.~ Expiol'·' :,J1.iUi1'mgrn:[f!l An~.J:ysis Gro~l] lMEPAG), 1'\"'0 NA~ f;l~$ hO!w M;l'r'i;~d! ..~ MEPtl.fi Ch;!.liL

HlIlHisruws
I]!l~mfor

FO(~SAIFlTKU

~ASTROBIOLOGY

ill> lifc"" II ~J'Jan,. p. 31 S). A. Lawle;r deseribes NASA's r:l:~;;,trobil)~o!!ypJ'ogrm'l1 3S tflll1);dy tliSOUJ.H1Ct:U::J Irom irs Sp1LCI.:~.mgh.1 missions. But recent cornnenrions p;)in~ ;1 dif1i::I:em pirt~llUre.F()r elHil'lilp!I.'O, Bruce Ja"'t)_~ky_ thePri Illcil){1lt n,vci>~.~g<lwl" PI) { the M mil'S Anno:lH1h~re .;md\i{ll~tit~ EvoMion l1lissit)!'l

Sr:.!ce Telescope.Rather

rhan being

(Ii,~col1~

MARCY' IlliAllWil (DVSK!{I AIN [I SY$AN !('Ilti~er

Il ER.KIE FOG H~

or

l[\.iiAVEN).

-t:IJ.ndkll<1:tesfor Ih.e ml,Ci'li.l the PI 01' l~~ NASA As~mbilJlog:y ~l1cSIUUU:: 's (Nt\~) University 0 C'lli(Wdd'l team, Mf.\YEN WQuM tlh!d}' amm~
oft\.\U

one

rl.oclL"<ifrmiJl NASA i]i!;!h~.{JiPprllltlln1i.lue~_!lSifl'w biology uoJecthtcs and Ihe asur()b~.olo<gy comnnunij ty are repemed Iy to IJIno ,.u thche ••rt of NASA S 'Inissiol!l5.
'(AIAiUI. fil.CHiEEI Dlr'Mti F. 1rt.lA5A .flslro'blinl0Q~ IIiI~H1Ut~, I~MP, I'!m~sft(!s~mh C~tl'r; ~11'~tl field, CII9'40l), USA.

r~r !';jre",~ks;!md S!Xil'!Y. 113~ 1.:11h :S11!1l~'~,SiuUe j!!iO, Oaklrliii'ld.,(II 946:lil. USA. ~:Prc·Chcic1l' 1\IIIllaocelor R!!~:f!I~I~I~ RI:!~~:rt:ln, lJ:l~,A~~~~, Ci'I,9:Jl401, m>.A.

]\1;1[\\.SCOUl

ut\ ;1~~O

]. ~iI'IIllSSC:B.gu!delillH.at
lnd'~;i;,h!m.

R~~ret'ef!!:~ ~.~~~;Cifgig!ilrde!'i!'ie~

2. A. Ptar:l:!!f!'. ·Ne~!!1l1imill~i!~ '~!Jlt!!~ll~~ r\~rnell! crll ~tie~(t,· N~tl1flist,oom mew~~e!:liit:e fm 01007 ••

(~eE 'I'M"II',a1~D1l!rI,nsl,[00il',aJilil:ll!"M.~id=dJJ1l100A:& (eeGJ!I=IlI1~Ile"·_rIl:.J,s52a~"

~phlCril: g.u; ~:;t'ari:> frlJm MiI[t\ KI Lmd:~L":;t;md \\h,~~ efn.I~1 !l11l1m~ph~r.k:....ol,t~irJIlhilS h~d 011 (,. , tilt; plum:!!:> cllmate 'LIn,d~a b.ill:l,bIIii~y.pI3~,i,ng lh;nol'le piece "flhe Il~.!;};le into the 1;.III:ger con~~1'>t tl(rlh~ pl:;l.,.t~t.s biological pnt~[ltilll The N:I\,J :st i11m'IJlcs til us kind of inl.egn-<1ti.\'C th ink~ng, y bri~l!j.ing i.\Jgdhcr Inuud I:ntl'lddisdp.li~ b

Oocyte DID natiionf'Dlr S,telM Ce:l:i R:esean:::h

:ror Slcun Cd I

AS ,INI Dl:r~Cc D lIN mE POLICY

rOR UM

'1HE

THE NEWlV ISSUEC INliH.MATIONAL SO(IIEiY RC1i!:an:h ~ISSCIR l gu.~(k:~ ilfl~s

nary
\\I'a~~

g~'OUriS or hl.'l\.'~ IllL;

sciemists who m~gI1tnot

oll.ler'"

r~pp(JIr'UiJ!it), 1.0 \"'Q.t'ik L{]~L;'th~r and IelIrlli I~O\\' to uo ntrib lI~1! h~ eaeh tlilii er's re~"f(h. M,\VIlN is .111! exiunple o:fwh<!t cun l:lrw;~'rom II~is Feni I!! s,rmmcL f The M:l!1i SC~"IL!l ~e~~ct:~o!lr\O ill(:'llUjL;ld ~W!:l
.i11l,Sr'LlInC;IfI~. d~'iI'e~tlpl1rl1.:'!I1L L 'l;;ffUrls ful:"

l!SA.·s

E1<oM<l'uSmissicn, oo!h ofwillich are direel prodLJ{~IL~ ofllil.t" NASA As.HOb.u{llog}' pru!irrJm. In ~ddilio!il.t~''l]ri il'!nn~~nel1!ls on [J,lCJm}~} M~trs Sc~ellcl~ l[,Jburn~ory ItMSL I r~cein:!.I A ~l'ohiiL1logy sl~PJlml In cumh~e 1he~,r S!JIc:cl~m] hl:r.1l igh.l. NA I ,r;~rtt"I][~~Lnl fonm.lr r]~ .u'Id C(a·l.nv.es~.i.g,a~.ClI':'i (:CJ!~ra 1.ly in;U:~ volvcd in op crm.DtHl.~ ~~H!I Cienecenalysls S rDr the ~VI(!lIl$Expl(lr~uion RanTs" Spirit H~d
0rpOIf[ Iillrli't}'..

r~b._.p" flO];), V\:or:thy ~ollll~m1.t1 ~jJny ~~IIfIl:l:\l9.!l,(: ~bt:i~~t tfLHh and tli'~il'ispall'l\"ncy ill! b:iL)IVllt:dit'~J re~l.'IL'L:h.d i,~~>Cli1ilil'lIFi:t ton llf reseureh bene fiB "!U h.Ult1!~llilY "ull.;;Lrg~ on jl!rSl .,nd OO"Sitl!l,lbl~ It:'miiJ>.~· lilt! tI lse ussien 0 F cnhanei n~ 't h~ a il~l:bfll!1,~dcom em process for Ihe PI'OC'Ufe'a I1ilcnl.oni!\'SLne5Iiim.l~]e~l!:> It l'.
Q" Dul~"yf:J.cll._ [i'bltl:'Y FOIfUITIi. 2
iaelude

~orhlll'IIl."Ul ,embr),olli!;

sl.eIiHcell feseiu.\:·h~G.

IsseI{ glJi~fulli~.l~ lil, hum~l1el1l1b!i)"~)l'Ii~ Sl~in !,;dll'eSC'lfd~"!G. Q. D •.'IJ.I.!Yet ~~r, .2 Feb., . p. tfCB I. couatries wih publ,ie po'~ic ies 'non ~h~ donatien of surplus embryos I()l' ste In eel ~
~~LfCh brm~ly 'Igro~ 0111 I1tl;td I.anl fliity 1];11;: i~ll~l:'med ~11I1~1li~ .!lJlt'i the /:1\ okhulre!l!r ""tltM~ iet.. ~lr i!1teres!. QU<I~itj' "L%l.m~lC~.lhempeutic PUIT· ]iIQ~.lD"'dJ:il>pl::il\:l'ib)l, \:\ool1:til,.:ntialiIy"Li7dt:~1hi ~~lY" aad ~thi","!;.teview ar~' <ll~, t:orn~nm! i:l mlll1~~

~~ide~i!l~~ w'[]llld in flier \~'l',ikie!l1 !:thic:'d.1 :Sl:l~l:danI.5 lin3t ~1:lVCatrt'f!itly been e=>t;lhli~he(j. \l\ie are r:!lnucuklliJy eoneemed aheat t~'J(; rucon1!'llendmi fiffii. ~h:ll ecid
Silt Ih(!

d.lIIlporhl.tli

Howl.:'\.er, I~.'II:~.'C less ~i:mi:ill b; u:I.'gilnlh~ liill.! Dil>;[rie~ .,!>tnclu.'ie nfhumml ~loC¥l,~rtlr stem ,~"CI~ rui>Cl:treh"The ru'C~'(Itly 'oc::lcm;,,'tJ ISSCR g,lli&·. lil1cs eue Lnpl to nil, Ihis £.<1P,btLt. in,m attel.npt~u IrC'd~~ cuascnsus. f:t n;:JOClnA,:d ~O.f~ VU.gli_~, l1fl.cll!Iri .. · •
fied pL'{'h~bil'ion Or·~III1(tlie~ Irnduq;ernen!~·I~w.u-dI· in:g t\[]~.tt'pUllsat~o.". of'!J!JcytIl, d,u!1u~_ The ~S'C of Ilrmm:1.ilil lnceatives ~()obtain 1~1lI1n:)[I eecytcs tu b l: used iu Slciill ct:n research is. a ool1leliltiow:J. issue. Arepill.tidp~t~
ing wOIT!en V'C.,.{!,[}fS. pnwidt:ts_
(lI"

siens <!bOM.! payiJ'IJ;,l.\\iI:'1J11en .1i,1ln.~l~ir~J',!.gs :;h(lLI~d be .~u]l o n.msdy 10l~tl.1tM-TSi!!J1I.'1.'tJmmh.t\:'(;S,. t

This is .::1 comllie7i social and cll1i~al questiou, M~LIl!Y who have ~N,um,~'rll1:d the i:;;[>iLl~,

(~Orlior:;;?

The NAI {il.!s,ocmmriblllcs 1.0 Ihmn:: missions through fOt:Ui;; Gn.lLlps ~hilll tnubHiZil: i:,.;:pe'lIli:l!e rrOll} 'ICrrJ5S, tho(; Instu!,me arid the wmde:r st;:icnl:iric c;:mluliiIlmity, Tl,l!.~~'kLn; H:M;u~ Group began by p~.\1)',hl.g .~ scmnlilfll ro~e in
l'eSHlLC~Lira Ilg

c~osdy. i,nc.iliJdiln.hl, olll:'Sch'Cs' have (;olll.dmk-J

IJllin rC$~3I'Cht::ffi

:;.:hmlkli compcliIs,.''ln:::wmllil:UII

d1C Ma r" Pwgrom .;dkr ~he ~OS$of NASA, IIIldss.it'ln~~ ~allm;lted in ~,'\)91i" ~I 'H;'O(lnlipj.i!'ih~~thi!oi by mwa~l7j[!1:..~tr\lbt[lln,gy al:~ifbTn!'l'Lnd;,l :~lLI!r!hero~"'..g~ln~i~,

<Jill~~~!~ i ru:t'!JmI11lr.:~lJlld i:iIIlh~ US. N<'Ltion ij,1 AC'w:il1l!n ic:\ gl.li~eHn:e:s,hi (lth!:I' "~'tTfCl~,he ~:SS('~ ~sn.0~'" I ~1J!!Lg,tt,;li Et t:hllll g.o!"~mlt1~l'Its iln~ "Igi;llll~i~~ n

mll.y 11'01"hei~' d~rocl e,xpf;.'uses. [0 <Il\'loid illduc~ t ~I~~ ~(l[l:no(lnic: i;iy \rulll!euO!lbi~ WOm~[lI)[l'U;:C;~pt •• I ~lcs igMi:nt"am rhiks of egg reuic\;~1 when Lhey W{)~hjll'H, otherwise be wilhn~ 10 do S(1. lUlis rel'spe!l.:~iv~'Imsheen adlJplcd <IS ,klw hn

That dCI)~llds (m the type: Qrcmnr!'l,::~1smiolll, Cunemly. then;: 'lfe fh'e !ll.odeb gfCompeD1~l. t[o.Tl': Fr.ecm;mket p~m: i~L fI.xed compen:.n· g
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LETTERS
the donor. ~Q~i,j]eCon.[mli~ ~t<lWS tl.e ...need]. ur li.ilI111(lllloLi i[u,::m'iVCI1~Ctll~~" Thism.ood pre~
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wuuld ,,1:-;1.1 include reimb~!r~em~n:r IOf time

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lur pr{jcuroL~n.lI.lIU, \VC.I.'J:J

indluc'e~n~nt5; OIl;;,'~ prim,lry

me: 1'If'.ll.i and IIIiIII imizes f~ []1l.da,1 1'!')S5~(I ~l~1}1.Ihc ::nld~.tj,cllul issue uf rO.rilll'lm shoppi:ng o.n Ii1I the donors. hl~~ ~~nore:: i!'1d~,,'idui!l1'l;'"'>;.pen~e)i.. i~elecling ;1 pro(;UI"Cmenl site on .~eoo",is of llilJ~ particular laws. in. clT(lCII IJlmrullrh;IY b(l A~'thDligharbitrJI)". il.liiu.o;;rll!: ad.\...rll~gC:l of cerlessened, The lun H;:mfll.io.nall S ~cmCell fOfll.!.lb1l, mint.y ami. in. II iSt~t:k-rol.SeJ1~e. fllir.lless. The fthics WorldnyP~ny .ll.l:li.I'n.llI.1t1S that I.his ~.nini mllm';\V"Jgc"PJl'I"O:llC:~ takes in~l]I llt~U um ;lpprO<Lc:il respects ubu:isilil am] sohd:lII'ily. rhe number arbours domlt(ld" I.BI H~ik.h hood a pmvidililg t'Lmirilimllml\'v,.'lg~ \Vtm.l(] result ill .:, lhJt: :tllli11tlUrrU will '''lin~nrgdy remain H sym~ 001ie reeegn iI ion, of the true V~tlue of s LlC~ high.ctr rate vir 1)~YIi1l1c:l:ltllb..'lnhe fitx.cd CO,IHt 111 sterneel) research, h r!iL)~ ren~mk~[1 model, R~i~11bllr.;.l.'01I~mof ~re~~"'!'i IJ',!J"licip'lli(lH \'j,d es .a feasi ble so lUI inn to an iSSW! th<ll seems.ro be fII more illll£lividllaI!lY~,I~lo:red apneeds ~o be 1~\,(In1 in~d: w:ulhi 11[hI: .~llrgcf COIlproach, 'Illl.:> depc:U!d~'i.hfl'l,\'~'l'lr. onwhether reimbursemem is for I:nC(lllV~nh~11C'c., Ihne. 1.C'.x~ ol·~I~e pnrljc~pmkm in r;m,c:IethiC's:1 overp;~in. nnd di;)("mnlhn. or is lirnjted 10 acn~;!~ si gill 0(' biorn.ecl ~c,):1re;;e.ln: 11. 'FcO~ fH~~Nn::~M'!~rM!~ It::t~l:l:iplb~!,;"'p':: U::<'l:S :;'~H:h.~~ tU'O:I!vcl.IodIgi mg. S1i'EM (lEU trolUJtM [nIleS, 'YOtRK!NG tPcAJRW, I)'u.-kiml~,!'l~~,d:;" ,HId of.hl)'~;.l ;e _There ;,!we~iJnit~~ u BAJRTHPi M)!I:R:IAKNO PPEflS} MICHEL REVEll" i Ufl.CCfli~~"'(:ii UtHU.l.liS ~IrI d,~l,l P ~lCll'L'l.:. I It\: GENtE'lm:1II R!U(IMAtR:DSON.J~OSEIFc KLflRE;4 adm1I11i:ii~rnt"'~I'ImoFrequired i~b~!rrl~n&lllile. I SAULA.lbnONE.N. ~ ROMIlI (I !5.ASV A :!imr.ruw~r receipted ot,>;>;rt;~se:;;-->i;]'Ii.ly rC11it:y ~llX_~NDIU MA_lIR!lN," ~AIN'WAHtLS:rROM,~ win in the long, ~enn ruri,~el" reduee ibe <lv,.ilB!R;ACHAlto\GER.l tPO.G !LEIE HlIN ftEMG,~ H~,~Iit}' of thl.::Sc nbilltt:ri11~:>, Tihili (jouLe! I\::iuli nn

~W!~h,m

SllO~~dlPrnhibi l '1 i~';'lI!e!(Juri SIVI:· "file IS SC'R g,uill.!.:]i.Ll!:5 di.rC'cl.I:'1 addruss these Cmlcem!O..
11~~ innml;h~(;"lia!]IO ~~c~vm~I'~ 1·'rJUol;:'~Ifi;l·J1le'"t ~lflfl\::L[crials") dearly states: "Consistclll w.iJh \'IIC~I~e~~a shed pori nci ples 0(" ju stie e in hubli man ~lJbj~cl.rest:::ut:l1!. Ull.:lRl 11I!I~i~lbi::' an,::iSOI.t· alLi,J~e rel[lliQnsh~p ~Ch'\ICe'nl. those from whom sueh .mtu,l;;ri::aJ:s ::Iren:cdvctl ,jJlld I.h,t: popul~~

o.Ippl!'a'l~h

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tions most iikel)' to hen.cfil rr'O!l!1Ilile research,"

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~.5b rend~: "There must 'he i1'im1J·

ito.ring of recruinnent 1)I1ICLk'es to (;,flSUre liililt Ull,vulncmble 1X11)1I.1';~!io!:] example, eeo· ~,r(Dr
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em;;,n~r .. ~d ['0 pllntcillut~ Ol_"i (lOCYI,(;' provklernlor 1:C~9r'{'h,:" Furthermere, the ~I~idlelinel'i~~n~~"I.! late~he ~Of!;; p.rim:~pl~ thiLt thttt be 1.1 "rigorous n:VtI:.'W tn en~~lr~ t h..t u:eil'!lbLir~ifi)n1:e!ll,t ~lf diu';:Cl exp~f~SI.;~,ur r:i ilal~ ~'II(Jonl::>lider"uitms of ~ny l-;iu'iI.d do ng~ ennstiture ,OIn ~Ul,,~ue inducement."
It~~"'ltth, th:1! is ~[lbji:'C~. H.rigomus In

oVlt(:1;ight

impor~ali'lI!Il nr ,J black m;lrk~t SG~'i~\"'Gli].d see l.liJ~ eurrent "exeeptinaal" ease liMil11~ UK., of~!,\\ dol1<1/1,i(CI1l research fur
cnnHllfl,d>li ICi1i~ ill' 'Ii1!aUtt'ltIlU 1(I, <lV\ITrbuim11 of the reili1ilbLlffiel1l1ieillllliiodel. lhis ;lp~lirwC'h~ ho~'1·~~ .likin til pn:widing a~"Ce~ ~r,i~ to drugs ur tn!i.'!i.li11c:l1IlS il1l dini~a:1 mrials m~iJ does not p;nll!.ll the .1~e'lIthyvolunteer guidehes In hltc,I'IlIPL~~o~ml J;Lli,delim ..~ ror biemedi:l,,.]re~1[n:ll. .Pet,l1apii> iJ 111Ujrl: ~(ll.1~'[abll~solulitulI,wl.ltIl!d
~I'I,t;:O::t:"h.:ll.tI!!f~ for 11t][C~ to ,

l(jllllll'l'll d~ !!!e~~'lif~~~ en D~"'lil fiublk, !LIn [".e~5i'!!i ~!ii MOIlHW'!. MQl1illl~i. Q;C IHe 3]',\', Ciill"dt-i!', ~llI1IiI!!11 !~jlU!1!! g( >ri;en(e,. kru~lem.15ra~lISdiOgi g! ~,Krn(~ C~We~t Lillfld:M, IOnd'OiI, YI( ~'Ii.!n ili'er~i!'1' Cil!Mr~ af B5~ Mh~c~. f~'~lyk. 'Ufiiv@'(~~~, EUnD, (~ec~1 tit~plil!}11c.

plPl'toCessat the'i<"lC<I'I, legion:!l, nr nauollOl ~b-el: Wi Sl.i puJ ttccJ i~.l th ,.;ISS,C R ~rtiid!c'I.~n":""-'::lfiId conducted inaceo ticl<lIIlCt: with these gtQ ~dng i rlrrll'lt"ipleswill. ~1;,'C!icl mh€ ~'Xploi tatinn 'of \mtii~ti
t~ili:lt i;~ m~II" ~liJn,'cIll.1o.~'u.~rl'i ~ ,

~r4aIi(lDalII!d ...n~ory

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fiil!ll!lil.ful. ,t,IIUlillllt 1I'~lhiqu~ ~ioolil'ldLEaL~. Un~1,!'i!IJlt)1 or GC'~~-"a.G~RI!'I'a.:'l'Nitzl!d.1llld. TUllillt~il)' o~ CirnlleoolJ r@, Gotll~FiblJllg,~il'dI1fli, l8i1n GlJri~illUnr!;'I~J·,~ty,ik!llf ~5lI111!~:J, l~r~@I, ~~tij~ool 81tx!11li(s COO!lminl'~ SiAgaJlO~'.

fl:eslplllllll1!!3:e

DARNOVSIKY .AND rosa ,RJltJSE,APPROPRIIATIE Inixell mflde~vn wh~ch ~1 eonce ms tlWll oLJC}'1e l:!(lniU iOI1 shnukl !(lot ~;IH (j,i!;pmpuniomndy ,Oil i[_"'-"UllnUl~.c,dly v~~~n(;r· M:lQu,hJrd mTi.mU:Ii' olcmnp(';l]s<u:km wtml~ be det~mrv!l(.ld by ~l competen.1 ,u!ti'!.nr.vt>,', b~u ;~bloe\!\lllrni;,\11 ,IImd t h;,[ .n::'~~'lrch guidelines be to develnp
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20 APRil 2007

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':LiDJ

ijni~m.~;a'll£l hOi'jlil:ml. ~10l}'lIi. ~!11l

ct:'n:ilif.l bi;;h<!vi(Jn~1

devol-

El.i.IIllIV 011 !U;!ik TI1e Un.cert.i:lin Legac.es

opmenml {ftItoomes,.>lnd clb!h,

lbe.'II:: ~i.nri~~~K1dtB reader In thillk ,IOO~1t 1'tl!! !;;,tl1;_,"'-'f {{Ilceii.!:ion: \\llmo LtlifiniHLidy dcci~~c)i. ~rnl!" care Ifo,' these newborns? :Sollle ofl:h~ !;;[mt1;S.aR lr.tgr~ ~i sh~-lretu'-o!d eh~ld bormfou!r'

or Merdiud Mlr;;lclers ~o (S 111D'I as. F11I11l i II as.

and SOCI·ely
.!J·.V Ji\l'u[fr io,Vf ,fj',[,j yell"

at ~eaS4 th!l\,"'C mOlMh'li premall~~I·<;,lt!!:Ind th{l';>.C:chUcl ren boruwith ~hm:m.Q,!;{)iill~~~r gt:o !l~ric' d~j!;;(,'t~....--'~~.le~~:. nl!M:h to be k;am{;di :l:U'tHll th!..: (;011"1: preto these ch ildren.jhe ourafthlL,:":ur birth, ;j11{;~JilOIi'Iulut

vided
OOinl.e_~

'11'1:.'1.Ii3 S ire~tll1K..."t ()nl~n. du}~r poiUb, ·(yul thai our poli:i.k::111.urad Ii IiljJ'm:il~pal iI::j~s. ~ll."iI1d ~"ol'Ifi ict-

.0

Callflffil EltHlks, Smrling, VA.

2006. 1St pp.SI..2.95liS 81~ 97B 1933·' CI2269.

ing

lil'i~s~~L:!'~ 3bn,~t tht' value

~an"ll.dlQjl_edu

I'!IIOn1h,~rren.mt~lre~.y

who ems

dntll nllnpOrit!HIICi! of these ehildrelil.~I1I.l1e''''e:r.'l1 We;.;.tcm eoun-

At I ,as mlllr:on gross s~reJeel ;6:39.00<1 ::;q,rore' rell:{; ~.ttguQUS meeOll: ;!;p€OC.e,iIle: GRBCC oow rnnks ru'iflong.d;,e 10 I,":~st
oonvm.Uon'l rnnt:el'!l in ArrKl:1b loQlOO t!'l

CO(J1{'rliention C:elllt:er

G,eor:gi~.R. StroWf1I

heiliFitd do'lNlloov.n HOllston <1I1d in the midfit of eflo~inmem. dining and ~[ghdif~,

um

* GRB

20 APRil 2007

VOL 316

SC~ENCE.

www.5ciencemas,;org

he<l~lllcare ullvl1!1:nciy coml'~bl!te~ tQ II~ecare ...mid U IiIU::al1tL!1> Ihw;~ chi Idr!Jn ,. Nit'Jill lasof ~inO!'titHl wi!h 'the "lmlir'li;"i~ child" I.'llu}' ba 01 \,v,ay In iIlCf:'C9SC nHings ~JT viewership, bm ,it 1V1lIY also relle(;'l~he inherear bcliefamnl'lg
W~swm secicties l~l::l[medica] wt:~:mo.~ogy c.:u "cure ~rlythj,n.£." \:l,rhen ;flI physu..::iall euunsclsu ]iLrnily thallhc'rt: is.a UJ"'o ehaaee ef a good euteeme, how malty families b~]ic\'i.::thm (bt::~.tll~l.d wililboe dun nll.€ b~lby mit of I.en? t I.ow Uillany phy sic ialls or ot her health care nrofe:;.si(l!10l15 believe llw~ .'IIlM,vef(;.'lJ 11m'!/'~he ... ~r: pe{:lh"t:s ol'I'll(;: ~rc.,n~r
t ries, l11ere are ~-egulatin" s and la\\'S th,u CUH..uun~gc ,1!!1:l ressive tBo<:Lbnt'im ![l th~ lTiolll '

this child will be differcl1t"? We have not

Ihlgih!:

Jl!;;\\llxlm>; Olj!{)~lncl the t~ll1~ ofbi[1th.

rcgLJ 1'~~I.imll::; a,h.lp[(:J

For

during Ih!l;!liteag;rll i\dminis.lrndnll ledm rhe dlsp!l~~ ~m.. of ..t~rN tll!iu n';'".1~~ ·'Dj;;cbltni!il;.~my~il;il .. UI:e' m !et~l anti "are ~or :Inm:d iCjJl~p!;.IciI,inJ'lni:r> in 'thb lucilhy is rruhi1J~too by fl:u't:nd !1l\\,."Ancr

!!.;'Nlln1!plc, Boby DoC'

the conelasion of une l:OI1lilni£UlilltJJ.' ill dt1.: ::1-004 :Bl'ili:sh :BltmJ!:'I~ling C'(lrpo:!"'1tiO'll,~jm;lun~n!UL)' H'h,(w I~tlmdf:JOll_l" / Gww,,, VjJ, \C'!'h~c:h £hl:: ~lLInmilri~ M "(lfII)I (OtmtfY that ~ rich elW~q:ghto >l!nQ,t'\l!'leo\ll<l.~1 s ilttl:lnsi\11:l ca~ ,also 1m;.;uht;;· ~lO:i1:IloM~g;Lnim;i ltl, accept resp(jillsi'bHUlyfor i'l~ecsre nf injpnired d,l~ldrcli)Lhwugllwul L~t:i['llvc~, bQu~.. neue of .
1l;lIilll1l,. end{lI'Se;oi.

G uycr

SQch~IY IiH:mr lmo these "~ry persoua I discussiens bet ween :fllm ili es ;JIHBme" ~(;;~~ rop ~~S'~i;!}ml~:;; nor ""h,1[ thee ..ru oflht~.~t: t:hil•. ~h'en ; ~1d k.,u-e:-. ;!b{l~lt lh~ ~e~ ~,hmlh ,.rld !1'"f .r~ilul'C in our sm:icty . .l)",b,- m msk sitU<tl,ld stimulate ::I discussion <In~mt difficult tepics

~ thege inf~'lfli1"S leave ~hli: 1l1e('iH.<Inili inlensive cuee lIIl 8 unit, t~,lo!lgh. liam!llcinl. ed~~~llim'iilt ~undiiil~" ~ ~ iC1111IPPtiD"t roo, r~ljni~k;!; ofthe1lC ~llilJl"t!l'I HII'\: If' U!l~I(11'~he COf].:-1;nn! th!'!o'\m~f.cutb<ick;; ll!, I!~~rni .. 'w

thellirl does" ,AJiJo~h~r q,lL-1,;S~im~. that rBile buok nl~~:;'. btn C<'Iumutanswer, Is ho\, twck:ly'i; underM.mding of the risk$nn.d ultimate bem;ln~~,pr

rL:'.!!~u"di.lilg rl v~~~Ilc:rubl.: pmb,ion or om INlrp~la!.:hi,l:vi Dlg :1 bJI ance between risks rm!:! b'!:llr.::fit:>. QV'l.!'rbr~aum::tlt m~d,underl,'e"IJnlenl, ~Hl(,o~ reasonable Imope~ versus ·'rt!lir.iC~li::';·wU~ mutin'U~ ['0 be il~pooIl11,m~s a Iri.Orc imd Inure child.fit;:'n ,Hrebor~ '·at risk."
]3~i ODl. 'For tI s,

l~. J..j261'5Cieoo~;1lA~~~'lI

[Reliant
R.elill!'!t far!<. wim !u 2.1 mliliQi'!gr'~ square' I'CiIl'G of 'ID<hibit and mec;cilll& :l;pa'c,C'

P:ark

~sdiie

11~~st spart5,ent:emU!iime,nt.,
ReliilllIt

~Jme WQI~~,

~}ijpo~~i;ODli :aTldtr:ld(!~iloIN

Park hm,mes 4 nrajor

!:QfI""'~

ill

~iiiti~, p,~tJan~Cer!ter G'06,.2i3 gsf of ~111g,kl~I '~O!lld&UOILI~ ,~i!lit ~~ei)

i%e'lilInt iScatlililm, iR.e1ioant: Ar':e1il0l arid!

1%~1i<trn; W'CidOf'l'le" A

:3:75

ENV~RONMENIT

T'he Burnliolg Issue

Ne i tiler gClvi1!rr:)mel1l'S ",or civil :society Il:lli\l'rEi'

be:en a ole t.IJ m ilijg:a [~tI1e a nillua I shmlllcl of Ilez ~ Ul'8 l b Iankens SOl.!the8sl A~i 81.

muamt:llt~1 C'lt'liS~iWpht:sdriven by perverse Iland-usc proc't lees ill tl,c I:LTIpiCS. Har.e poU~~~ in S('lU~l·· iora east Asia is an exaruple lh at Ilms imiill~-d~;lle and ~.{m:g.I:!.>fm.m::q,m1.h~
'OOIl~m:·nOCi>.

he' ilII.teffllltiol!la~ eommu-

nil}"

11~c5r~iJ,lJcil.I'C.P.I!;Jl.Cll!Y . to PF~\'e1lt ~aUl,.~(',";lle cnl,'j;~

"'"'
.

'11.

" ", .,.

needed toclenr 'Illnul wi!hmn f re, 1I:8aoo I!rnnro,l!Ime:ntill mlanagemelJlt Imrr~ slWl~d i'w;:' made tn edaeme IOc:'dl r:.!nle\L'S on tl1e medica], ,~I];\~l"!;ll1nlilelnal_u~d pul iti.eal rusni II:L'I~ nons or buming.. {1~ongWlith. ahernarc ~nethodli or kn~d de<lri!ij!,.Vnh1Ill;eer (I!rgmdf:(l~a(nni. ~.ueh .
1.0 tlre, J"J'!i:-frlher the IIli~tQne-~i~H1ove"u-nmem nm' y 1X;'gilJ(ln~tl S:t::AN pnlSl\~1 re .It•.lVC been ~,bl'l.:W A :;j:£Jijl ,1111;: haze U fJ. II). OI:!!1lui!l~elfonl> by the alt~Tlmivcl>

asihe u.s. ~t1(;eCorps(lmlk~ heJ~):;lllOp i'i1!i!S,(ln the glIl~illd ... Ilem;b rural Cl)ll1fllliUl'lit~~ Ilbmu nd

The us-e' of fi re to c 1e"~1I'" .11].<1 1 is ~:;:n~l'eltdllt;din

11l'hi1L1),

&mtll~.u;t

k<;j,Ul '~'ld'tlI«iS U). ;nM;II~lge I;;(J~ll·'

ml."iX:iall.[iIi,"Cpliljlmlii,,!)fir-pani,C!U~ Im'Jy i[illl~@ne~f" and E;Jl!~M>lliiI!Y·

sia--haw he~ur.~ bU~nJing{Jill 1iI.]fI
,!.:nOI,'.l10HS~1~~e

Forest Iilltd Agr;i.~lI~tun.; 0 Ugi!tl~.;aLI.i,oili th..: of

IJniloo Naiions ,,{.il. indt~ing.",

recent m~til1g; buth

'.h The

fe.'';llli:ing

and ~~. wn:tr~shop lim!h~dolile$i.. h) {"lin ~t1n~mumilti'-biilsed:

on :;~t'i:aiI11IM~ Iilru,.;tI:!:l~mg~lWe~[

S~~l(Jkt:,'nX "haze," frtl~n hJ1:iIlJrud:> ,of simtlltmleoLIt:: Ilre:H3J.)lfe'dd~acrl"lS.~ Ih~ mgioiil ~:iti~li~io:i~l!:rm,0il1M!1mild d ry s~'aslJn. Iii1tIgr;: !:in!!: Fur weeks, Fires to de.<'!!rland Illi!W hum l'llIt

11~ imLTI~~g":f:!,Cl'm. atklfC~~ill'!.: probkms

al.

,l)F centtol,
Niiiu

!ie'

~'}3I'li,",,11<lr~rIItHill)J

EI
(lif

gi Ll~.Tbese exaet 'llil'i:ghMI~I)rIthi~ ,o!.!i1!tri,ei r~sli!~lil1lgh@m ~il~' h~l~ (ii'pjsode ~Il;il,c:e~b~~:;:d by I h e hOI:l.T:lCI~:~biudJi\l~~i~y ~J.3l [Jur· il.997~8 EJ Nino tBJ. Tl'iesliooilti 01ha~ fr'Ol!r'Il'iJ~y :m01 HI'f.~ cdi'! lJe seelll ~11I1M ba(kg~ound S<I~ellite Image, ing 111~Iu~~ EH Nirm in 1997-q~,
tires (10IBSU ~nc~tl\c'r!:l mHi LlJIlllw('-

y~'Ii·K, m,!,d o;;OI!!_~l1m~ 1:1['!l!,~ S\~I,[Jhtl uf 11:'oiiCl>l~~9'll~(!d by ~~l~- i!iletltS,

~iI1mlk~ haze. ESlimiltEd damage ill I'ndn ne--sh!iI nd Q;1h~.

!yO\'II:'mr:m:nt·Jl.undt;;cmmnmLly levels, ln <lddition, 'tiltl Ul'ii~!':'d N1ilio~s ot th~ \Vcr]d Ballk oould help gel1)1;jr-.U~ iltled1;,'tIlllmd;lll~I\)LI!il.h dclnIOf'-nOllI,Ure ~"'Olp oif'C<!ll1!Gtnr credit o~~e~ fund\;, :SOllL~LOI'I:): lA:i Hlt! haze iluohlem!ll'C seeded b[!'foDTL;'h~ '11i!15Ctof tile dry ~.eaS~]n in JUJM. <L" t 2;(X17 miIY b.:.M(ll:h~rEI Nili1oYl.'"drj i}J .. la~l11i! . le:;lmed 11iI.'liln liscO!ll1llstt".l~lhem~y help.,me.li~l" d f'J1~ s:ii111iar !i.1l1IJk~.lkmJ llpii:-;0i;,1c-_~ l in AI1I'I~~(!nJ~ .•

AfriCa. ,,!n~enher parts of A~i<u"

(;. G. Mlrlilii, Ed.,. rilMiti~~~l A.lIrirofllll't' i~:lQt.<rnfil.Y 1Illia: AIHt.i.m~R [,Ctlfl:iQy,Pl:!f.fjJ'tGri~ WIIMt'lii!lr fr~~ C. v. @a~l!!!!<r; J- S.;hIr.'eilJl!elm, ~h[~!tly l(Jl~~j~iI'(i' ~!IItf f~ f.rg~ll!J' flJlky In mdd!'l~i\:J'~ fll1 qCfi~i~ ~ Rl~ (!l'i\'ln(d Re'tCl;!w~ l~iliUw!~. Witjhi",!!~@II,. 0<:, 20000ffl. ,F,~~rd. 't Go 0'!;:~1l, (qf.l_ ~·aI.:1Z.~5~ U!l9& i}" Gi.I:J!II!l, I,Jiessup. IEd~., !rItd'OIi\!'~Nl>'sHIes lInd Hal:!!: I1to!' C(l'J;~oj(I1r,~'ltJPhr" ~lmterfij,!l~lJ(Iai.llle\1efulllll1.o!if1.1 fl.!!:;ei!Rb
Cellle[
21D~dJ I! ~1il5~ibJ[~ III

.highlig~'al~\ter<ill priorvlies 1'lC;I~HW. :1. ICOIUl"!;lltrat.e' 'CD:'" dil~l.Vfill'e5.," Tu IllIlifl~ffilli1!.1,;: " Ih"Ul.3 t!1uIUol!I.knl:1 al1d 10 nlliJIUua riUQJ~}~.~ pol in haze (2. 4. 51, tVI;,lleri!!~ d~~m~ge~~~i'~l1!~es ri)l;l grl;ii.ln~nn~e ~;I~S ,~~!1d ~lrtion ~~)(!CiO!t!:!d 2. .ffil.lgC'tl From USS4.4 IJO' $9.7 bi UiIJIl ,1.4) (sc~ with fires, pn:wll1ingm'l.d.cxti.ngu~;i>h~.ill~ "dirty I1res"' Jiom peat green vegetation. which 3btwe r]gl~U: C0:lD!lld s.oot~iber;!lCd froiffl burning pe.n and IJ~Olllllllll;:-; released alII ~lJ~t: more ('0, mid ~om.lh:m t~ri~dhi.Lllll!i1~. 3. ,esliml~lcd0.8 I 1.01.:5'7 ghgmons of'earbon imo should gl.:~.hig~'k:Stp:rior~lY (to). 1'.1. IJ~(~ anncsphcrc (6). In ilddillo.ll to ilt{;~asnnll!l (!®(lrdilfla~e ~irre~lfii~hl:illll~1 ,efl\olfts.lhc ~998 hOSJlital ,Klmissiorns llilnlllglwllI: t~l!'e,gkm. unI J\SE AN Ab'lfCe],m!~l 01'1 Tl'aHSb(lliliull1r.)1' Iiaze he;;liatTY itt:.1Z~cm]{liliml~ increased :~n[l:rtuHty~iii Pol illlL[Y~l alls (hI' e:.;1.:Uh115lrung l'lju.rdil)~l~~!il c ,I !ii, ~, h Mal!flYS1;'l am! ~rnve~d 1111al1ll. and ICtal stl.rviV31 cerner for ooul ~l1g haze, wh toll has 11101 een fo:r~ 'l, in: I~(lnesi<l. 0. in IIm~Uy OJnMiiJu'r:d because lndonesia has 110'1 ye~ IrmiWiccllile tngree1m:m, Slre'~n~lil1lillg governUsillg ilile m clear .~:J1I:lcl hM been iI:Ieg.al iIII hlt;B[ln~i."l !ii~'f,': ~995. ,uad ,I ~rn·bl:!:rl!lu!tg j}il'lUil~nt.,.1 Il'r{lcedlwe,; fnr h~!1dl~!,1~ f'i~'e~ m ,~)' S, 1(;], \~'IS .I'Hii by [hi: }\.S:-1m::illlliOill:Il·SmlllTI~Lm. ::ipc-ed :1,'i.::Sp!m.~" i 9, P'ro'lli ~!l: ~1iI(~tliti~~t:~ $1QP !;ill!! ~l1ing. I~i'll~ J\~i1lnN<llklnsti\SfAN)ifi IOOQ(4,9~,AlthougJil m,my M~~~~Y:i~j:DklnU:!~~.{la~ alii h::JIv(l storp'pd ~n~jal ir!r:."I.'\lll.i",~ '~mkl ~ if!'Tw"id~dI. to d i'SlTicl!.~ 111 tl-'lP..lgi]re (4tf:el!I"!p~~trin!:ln<l7~ m-er Somt:ie;;J;S't or vi I'>l~ll~~l;;lt hiil.....r.nl i~ed th~mfiif:~\'~s ;.!I:~d .n. e 12, A,,,iil every )'C~lr, ilildll~ii1g WI}(i.Ln.dicm~~ 1.Ili.~L I~rnrca'foi<l.'<! btln~i~lg during IlilZlJ1~lt}util0tm.J~;·· ures on Barnefl mxl Smn:~tra. hl;lIlke!i'l~g murc

Refere~,t:es ami
i1IIIli~~.

r\liOle5,

co. Ion! M.

ns.ure.

,onu

SGiIIJJ@1I5iA~ian !>fudil!S"

U~ 91~!:! S.~,I'ilg!Wal" flr!aMHZII. 6U,0031, ~ s..1:r~~lMIfo1n, "Mr qu~1j(1I' ~l'Id Irrlilrrt fI'WIl1atiL'J dil.nn~1 ,'"
ImdltJ\t!iili'.~IYIMl<llt cw~ldrim irl1.9!i7" (li~il, I'tjlOFl u 'i'j, 81!1~1(D:8!lif~U 1M ~l'!f\uOllln IEfCflomrt rl!.fIlll~iI! cjl ~ :~

'5i11.~poH'. Wil'J.I. '" Np:~ICILAlmI. ~n~1~~ ~l.lllD9J

ottawa a!oo

nc~

il'e~IJ!I;ImtlI!:o (am'~Mge'. MI!,. ~05~.Q

"',Sa:5!iI)'"O~§r~Jllhyl9,1(;!OOO).

LI.~. foe<! ~~E! Agri'~~t1!irt gr!i~ tii:.!tiQI'I ~M!'Jl"ftlil'M!ffl~ellU\Qr-Gr!!J~I41~~i!i'l~2M {FI!.O', it(t!!M'. 200~,1.
r:I, B'j\roll, Agfi~, .f~, ffflihm 1m'll, .5:1 (l:OO!!~. !. ~Ldwt ArmD:!; .fmllrol1..;!~~3~11 {l~ !jlll}. Mrilll!!'m'IliiJ5;.2007] Year 01 EIINlifitlrh\a~iJa, ~.SiIroiI3 ' Jime-; •.'''~tilm1l007" Iii',:t!1l. 1 P:. T!I!mlC'll, D. LT~omaoll, ffl. ifM rtOllrd\'lijk,~"

i
,G')

~ '§!.
~ :~

lioll:>. Stmn,b'er im"elltlvesfi1u' ooJpnmliolls that 1:~ml11e'S ihfOUginolit L the :~iall .md p.liml~lioRS in ~.I];[lmne~i~ b~non~ tlbL;l an. fiiVel1 till~sm~go;ll !l;:1~._q:r hmd I'm ri~l~t'llimil\i. ~l;~hl. i:n1d~~ep;;ty. b

B,

a
~

t~r.

Emo1l'l:1l1', ~4~29

~200~1.

:I!I

find ~eolill.mmi.J:COl1iillllo:;;xi.[yoflht:: m~unm~~ I~azc prob-Ir.m. llil~ru:""'u~1 oc Ui{H~{tlo)' ~ALlliHti. blll'!.\~ Th!~ iilllli1ors: <!ire~ Ihe IDep.;lrl~~!'I! of 1.l;~l~!tal :5Cienm,
NiltIDliIi!Il ""~eI1iL)'of :5i1lQi!:JlIlI!O'" 5'f~~~'!i:' 117543; Hm~l1; lilll~l'Ili!!l1@~u~.~du,~1 ~I),I,UQJ dbm-5$Dl:!S.eGiL!;$9 N,!i.5J ~

~nt:nt {Jfup-rhlm

P'rlM,fide~iWIJi!irnati'lles ~brulrl1li!il!JI. rire .11\'1;';1),:;' b~ thel:'h.e.~pe~1nl~l!'tud h1 ~1~0l!f Iud in lhl.: uupies. 11m~~n~llkm8 ~aid agtn.clc$ :l>.hOlI.hJ R'I!;:'us on pnli ..~>di~.; illrml'i~a'I.I~tlll1r: hil'I'dwHre <Illd

nru: ii'iz.;u~ul0L:" um,

el.cpu:>~i.DS Imm'L.bntliry "fU), Qr

1'1, limnb IHri~·-OOIJ<-210-112Icomlh~S~lIPM(lM!Ilk!lry

01&14H.tlioll Wjl~Fled Ihl' ~ijlliao.

,5
'~ ...,

~.

Wm

!iUPIHIifUng Omtfnl! M~l1!rii~t ilM'r,t~i~c!llla9:i!l!!l!1l:9ar(DrniJ!<i'lb'1u11.i'11.&r5

823137&'1)(1

:~ :~

37,6

IPILA NT 8C IIIEIN GE

11'18't ra ppln~

o-r ,iii d ifrusib le r!~l!!llatelrv j:lro~eil1l Ill/it!: bi [nH"91parll1c,i:; (I (@ntral i((llln pl)ln~rll ir1I the de\lletopme,rlt o~,as!IlI(tun~ iilll p la nl:!! 'lila t ha~ thetlli(kne~~ !Dr a ~ifigJE cell
l1:lcnt of another b}'\:l' of ClldlJ d.cf.nnul. cells, It

A
,I,nt;nl

precess dUriJl.!1,~n.1C dcvel(11~nrnel]~ rmul~iceihDku mllJ~SI'll:S is o till", ~llYlllg dowu or I:xHH1(tITn~r;. bet~'lX::ngroups» hlil:fclJ'IC'llD cell types, fh.e ,"1'11CI:g,e~K-e el'such btJf(_ledim.o::;. ~uany require~cell~ u on e~lher side oHhe OOi.mdmy to COBIl1Hlmicate with el~~h ()ther.lll mllrny orgllllli~!]!;.'ii; Lhe~ s~g:na:ls may hi: prohlillS lhilll~U()'\c'C lrom OI1~C(][I .[,lJ'(lUp tfl f1ml~ler. cliciling reSPQwms ~h:111:>e5ull in I1.NO d'i:;,[im:!L'L:H popul,ati!l:lm;. ]11 pbli.t:;" n Mll~11nu~.nb~lfnl~p~~I~~IlS L~H'cm.nml dt:\'!,jl\:IpiIIl()<,,"1: Ihmm

!! ..impui.'l;mo.

mlld,,;us. l~U:1lp.tJSlt1 vely

liml

m.ohi.1 e S.fl RplfOwi [I, j IIiI I.~k t':mJ:mlemil!~ eel l, Tbis fonliils a p'1silivc .lccdoook meehanisru ji;):r SC It pr"tJdllc~i!ll!m inrCliiill(11,1~eto SBR tr:'dl.1SfH.'n, Sill R therdor-e oct ~mles .I~e p:rodlin:tion of its O\\'Jl Imp. ene c~H !"yer .w.ray. ". S~1Fi.~~g.Vth(IJ1 may accmnn lor thl.! fU:rJ113tlOll a Side surroundeo by ;I. f'11l,gle I;zye:r nf enc;kl~eelTI1i i~in the

or 11IIlOI'eSCit

fl':gUI3Jh;S

[he pl:'m:l~c~
U'JPS any

\''rillich JII1Wl'l'I.

,~)r SHR

\\"~ij;Ilfe\'jlo1!!'1ly

:i.B'Dduc!l::S (!I:,velbpli1il~m () r l>upE'rmu· Ihe

shown th:I1 ectopic eX~1re:s:!lioPl

mera~y ~Dldodt:l'l1mi:l.lI!i)''C].'$ (j),I(;ldt~-xi. Cui (!I ell. r(]lnu~!:11'<Iiplants with kS:ii SCR~:rIJ~f.lill
develop slIperlJllLlnel."uy i>lYl!l's ofendod:t"fl1:ris. ,l'UIl~,is:Lelbr\'IIi~h the ruilOcl~llll"t SC R n::s.~ricl~

or

JL;;v!..'.lQpiUlg

roct,

rl;l[ll,!P~dl!.~:!mlt;Celewun
L",tiaJ; 1~11J1·'iJ.lisl.n\Jd;.;L The ;ml~~(l1:Senetig

pari:

ufthi:;; !:e-Im!l1

I,hr;;

iharof neighhmillg 421 of thi~~~~IL::, ["Il~ e:f {d_ ~ II describe a I1r1ecI12I_Il.islill ilU~. any d t;p.11i t,.;~U1.c.:b; the mrnrcm,!o:fIJt. nf ulle

c:'l'IJpla:lm~ ul' Oill.: Q;:cH to ,~II s, Tn a rel1flH on pl)~e

Ihc nlObi:1i,t)! of :i~1Rm a ~ilgl~' lay(; i,' or cells, i The proa)(t,~d pos i~ive f~~clb;ICk~oop th<l.t ensures :>ufl.dt:lll SCI llU U"\P SHI\ ~n Lht': endcdermal cell n:,'iq;",i!;~.~'1.n1 in it;:l1 de\'el()'~l' merual :;1;;,ttclin ·""chl.~~l there
is a
~O\V

h<lckgrom1(!~eve!

'L)I·SCR ,It»lpm . .:~:tm~in

I.h~

such r110Mle p'I'Ole~n beU"'\iet:Il~'~'o

;;iIdj~lce!ilt gr(rup.:. 01- cells in p IlInl reots, lhis I1may(:ol1iilitut~ .. ~1'k:1f'Iii!
~l)1;!'.m~Ib~' hich w

~nd.odefmis,,~ljlhough lhcre is nodirect t'~'~,d~ll.tclor l:his yet II <I lse re'I~~ ~ L'~5 mb'IIi~[~)I+1t~[(t>[00[1 Qf SHR
in i:h.-; ~ade..whk'~ hit .. bet
III

mllhcc~~lul<rrnJT~ J;l;_U1ii:\m~~ rel the WQ.l11l~tion.of !:om

tk...,,cL;'ll'liOWldllfi:t!', ....

shtJ.\'1.'1'I The model of t3). C~U et tjl. m::~otl I'IIS I'or l'l~e i _';'<Lbil~;dl'l~ lh'~lil;'! f'ul!ld=> of
Ur'i:XpUC:£!l~.~,m

li~ Ilhunt ,.:J I'ahil'ir!r).o;i s tJwlin~ lirma. SnORTROOT (SHR.) is." GRAS 13.LrniJyrulll>C.riplionu.i rugu~ t hntl:r Lh:n m'l~Y If!m'!'-~,,,[he di:!>~,!l!nce ef a ~!fI~lc cell fl'UIf1ILts cell of
H.t)l:!l i~ S,IIR mm'eml!lll t~5:rriC'~£d lD l!~is.d~.M;mc:tt·!'h W<I!<;

\dmhJ~I'h'i:d

1..:on~am bOlllUbLes 01 the rid

~'v!tl0J:l11l'llt'U![,of [I si:n~c bycir of enJod~runi~. The cha:fuc~L;lri;c,llir:1n uf Ihl: Si~tl3Iill!'; Il.cn..tllik duu leads to ~hl"Sl.::iJ~'ilfi:JJ )it;l~e:01orlll~ 1:\'.;0 cell IlnpU!:ltirlil.1Sn.~l~\,'n!!ledsII I 00 t1$.Ohb'd !JUt. The Cui ~.{aL r~~~dl~ :~u~ges~ '!;tl!ll:g;ol 'hil~ of'thc llillXuiurn. 1lInili.,disU:m(!..; f'mevemcm of e imercellelar fre£tl~:t1tOl.'S is detemumed by tl1~ spmi~'1 di :;;tr~blnimt oflile~r h.iind:i fOgp1uri:nuelJi_ It

~1rigiIl12),

pr~viflH~,~Y show~ dl!<ltSHR ~1'rn~ tc i~.l i,s &YIlI;th..::sile,(l in 0he iancrmost tissue tile roor, the stele, and nlH]r\<e~ If] the .nLljacl1:rnI. ~n~

or

dodcrmal

lL~5.LJe

wl~cre a s~~nililr

nf~leun c;elt<;, ln Olddi~ ticn, SHB. mm1J~ trom tile :sick: imn the eado! dennisto pooitivcly OOml\l:~ SC'R exrnegs;i:(lll.
CO!mn<l!};.;e't

~ ~
II!.

~ ~
<Ii

I
....

ilrlg ~).,rmt:rr in, th.e cell 10 which lhc' reglilamr ~he s.um:-!lnel;h'l.!1 i=-n~ 'cmdld l)p,(;m'i.U~t1u!",ilflg anirual Jc:,~clu":ilne:llt cal ~y!Jill ~irn.I.:~·.mdpliilnts ,..... d l>('rc~lfl>t;d exi,l)) ~ rr-eS:il ion g f SC' R t SCR"iklIl""kdo\'\~lIl iI,lams I, inw!h i c:h ~!I';m~cri~t iOIl;.'iI ~-e:gl~ <Lt(lrs~m;h as c lh(:' n!Cl~~II~rn:pn~'Ll by ('ui t;! (Ii. ~:l';rttli[l~ Ahh01,1!;~thl? SCR-kVloc"'dm:u~ pla~ts ;~~t:~~ chi!(_~k 1~:gPlih;:d2 1.1I!![~\~~~mll dt tu cdll (/j. "\Jtl~re-.,scl~i~~ 81groiled::! rr.nteil'l mm·~:s. why SHR Im)~~i nn~mresVlO fbnher ~h!ln 9m~ 1l1t!!n~~ le~ .. (If ~Jle SHR~tr;;(_ppjng prole.ill :Irotll the c,.;H ~.~wh~_c1Jiit is s;ym:l'"L~-sil'£d. io i~s tt)i11~UI~RLIpbtliS.. diC}' rmd.lK"t: cd I j tl,ycr. On.CC'i:l1:ItI:;.poriJ~d. tu tIIil ~llcl.C!J{;'fmul ~SC R 11\:~~rdrv,~ 1100rm~ :lIl1a~lflls of SII R. U' SC'JRd.(1cS indeed n.eigilhorins cdl vi<l,,) $,«I\r:tion alld reup!akJ.:: I !:cU (by a cuue'l1l1y Lmk,l'Iowl'I mechmaism). ~!.'que_~I~ S!~] i!1 the '~ii,dod~f!TIi~. ~(;'l~1 r R ~ d(;l~ [I'il1;X:'i1;J!Ili!f<irn. it i~ul~cl~l!:thOWP:~;l_til p;roleillS SH R bil1ids 1.0 SCR.lbl'll1 i[J~ [i tT~lrI~ r~pli.onw t:re~s\ld 11i.lJ1illJlIIJ1iO$ SCRill~ the t!ltdOoLic!l'Iul of ~Uy ::leii\l~ (~mnpl~1(. th!il ~1l11i l'l:lng~r l1~o\~ ~1~\ :!II"Clnoving. rtmll1~'I!i~ ce.lrl. 10 CI:i i l!1' al. ~.b,C! sligy::sl thll~ Hl~~_r)]r!1j)o~J e!1:l1~S11lioLLIJ li(: iIl'iMi'!Tidcnt '[(I iJind ~~Uofdl~ liln1.n ce:U to cd I. Thi~ ,t;'I(](iUp,i.L;X, !O!:':ul!.'It!iii,th~ I1llechllni~m of i:ndodcliiln"II bOllndOl'I,'Y forma-· i~c0l1Illirn£! S~m.pmtein. COI1!!>et.llI~nl.ly. e:'oa:e~, nonfli:ull'P"'"cl SH~ could pa-\''Sthimuglil.lhis h~ve~" lion n'Q,I)' '~. ctln..;(;'rl,'ed lilrougll ~\!irl'~tLlio!1r. Th!e iuthli!r i~, rnl~~ l)epar!!I1~n.I 01' 'Olll.and Di!!'!'el~pl'!!elifilal ofl:J1dt:ltli.!rUmllll OL.'1115 prrocc~ tlil oLlN:r c~'lh\ i:M::cil.LiliieI_h~ ric~ Qii1:htlhl.g:s ufSCR l-lil1.J SHR. IIml: ll:ielQllj)'; llijl~~1!!!Il€~ (~r.:, ,~o!J!!'!li~M"F!4 ,Uti. !JK.IE.·!nii~ I~a!J1o.da~iln@ blm~.iK.I! k ohlile !:oor. !'lol~-miall}' trl,~;;::r~l'IJ,! Ihe d::~''e'iof~- hOlve iimi!"'rexp[~s!'i,o.ll paUierns. The,endooef~
n'1I0\'e,,_ [n print: iple.

GRAS lumily 1.~Ln~YnplkJrl 1[ltlGJ' (i'D lilt!' ~iulrt!h~'r. ~R pr®t,~in i~ I 5 called SC'AR.:EC:RO\<V €SCR~ is, '5)'!1I;h~si!l:~ d i!fl S~I!~~ ,t:e~'sin p1<1F11. ' e--"",pn'L~j\t::d fl. 4) '(-'1eL'llhe ri~~lre, .• Vlml;pdop!i~ tll'l7l.iall'c) roO!:!:. alliiil moveS th~ distta nClE!@J Olt~ ,,~n T~Jgcthcr, Sl-l R aad scR. pos~L lively whe;li~ vt. biliids ~(JI SCIl pr,IHeiln, The (0IT1pil'!l remailrl~ trappeiil ill L'e;,g.U r<!!l-e d~welop-Jm::nlof the euda~!;tQ~ c12-'115,trig g,Q~ IlJOOI!U;Ii@11 or motm' ${R, .. nddlirreC~5 till? (G"~~ III dermis a~.ld.c{1J,'!iC'<I~ tissue Ihnlll 3. develop into a ~ii'lgi,e la,yer 01 ~nrdode-rmis Illai SUrrO'lll'ldS I he stele.

is lbt.'f,dare possible o]mllhc observedmoveii.'~m of (]th€rr~1~IU protein:..., sueh as the n'jJII1~

sCl'ip~ ional regWi ~~,tors, PRICE (i u jhemot ('A -epi¢le:nH~I'i.) mtcl [{NOIT[DJSHOOT l\.'lER~· sT.!.: M Lt::5s. j in} [be sho O'l apical mer! siem l' 1.'1.oJ.. is deten:nim:d by lbep'I,~sellcenra biad-

ViMIVIi':sdencemagl,Qr(!

SCIENCE

VOl.316

2.0APRIL.2007

PERSPECTIVES
rnis of rlK1;.;t '~mKlp] 1I!nt.. is unii'iCI'i;nc I!!.'lllnpdsUlg
!JIlL:

lay>L!r). aUld

irS-HR

Uii:l!,l S'('R

o.nlluolags

~Hl:tro~ d(;'wlop.!I1u~!1l in L~e~ ~i~s.. ~bis n1ll\~h:illll ..nm m~y 111~i\l::C\,<Q,ivcd I..:ilrl~ in 11l~ h~s~

~~"ry lalld p~al!lts. But there are some rare of

~'CJInioru;;_F[]irCXaJllp~.c.xtalll,
TIlUmwl'S 0[1111::

l:t]!JIlsetales (hoI"SCmi Is], whose forebears were ill~pOitml species in Carixm:ifcrOl:ISfim"5l.s 300 million j<'Cill'S ago, h::l\'';: l~'I.'O1<JJ)I(rl1 r'~'Jltioder:. o
unwin:. ~t"il"rfi!~~ hllP.Octrlilsl'la\'l: appiiil\:lMly ~inkel\-:dwil'lll:la~ir SCl~!llld ~'l;lm genes ~()ollll.)w S HR to esc,!ij)e the C'lu!iC~es 0 r sICa ~n ttU:nr.:ll ell!jod::i11nEl~ bl)'llr. lllilus eXlil;.,"IlcJ~ iH,!i!:lilt;: 011dkx.iemll!> III Ihn~r hl)~rs.. Why is [I~>cStnWILlre of::l shng le-I~:ycrcd endndermis seconserved across Ul(Fst :l.mnd
~tiJi:;, il1J ~lill~

plml1l:b? Al~'otf{lphi~ pbm~J\, [t:ly 011 the Wlp!<ik~ Ol!l,g~O~~ernu;, {1]l.n""'eri~l!1:p~Jl!nl'"~mnJ .dso pj}~!l!;"l ol'l1U1:ncms .3i1il.d \~lI~~r [rom ~hl: :lOoil. so tLgh[. to ~h~tu:l.gcs: I:h;'vd~pm~I'!tlU! in prugr::J,~.l!>th:J~ !iP'<~ti~1~~i1!imicJn of e;Cldl[ulL;lrlim~~ ~~.aflrri~nJ~ lIuary Il.~vl,l OI,.'lI;;'wu'red r dl;l.. d11ri!'lfl the :p,~5t470 rllu~ lion C re rI eets il.~ gl:l~ C:lI1("C'. 1'1,k end\;, dcr rnls is )iC.?i.US orl3lt~P~r.lI1i[ \,'tlhlt~{lll. S1 fi highty ...pec~i'llij~~1 in ~Imt inle~l.lelluhlr spm::t'!'l itfffl'!'!:!l!:~ OOr\rol:n I,ml~ culls ~1fC' ~~<J~,~d, pwhi.D:l~;mI!j: tlilic 1, IH,(u! tr Ill/, S(j~!f ll!6.. 4U (20017l'. free dliITusfo,Ul of water andions lhm~l!Jnn .L 'if•. H:etalTiiulia ~f ct, (r-ll J1j)l. '~5~ (i!:OO~'_
lhil:

intersriccs.J

r water

anti ions

.IUI::

to pass
so in

llilroLlgh (his mmJbJod.

'~ley IlfIitaMdlo

<l

re..~uhm:d mm~ll:rI~r_ i., rht>qf~npl;}:\nm ,r)f endnv dCr,In!IJ, cells, 'This amonls strict eennol over .r.:ln:5p0!1 and ~n;lY '71reU have llIDVided., MW·' \'i~'.d acll'3'I.1l~~e~the highe~1 level 0 r I.:mltrol over:;,'l :s,~i.di:rn~i li~~L~ of v~ry~i,LiJli~i,.'lJl~!7:t1_ f 1 ,;~l:igMy fe];ulmed
SQ.,ih~~d~:ocovc.ry miily e."pI3iIilIJlc!o.'Vo~lnLC!'ll gf d(;welflrmeL~trLl pU'q'~rmn ~n

~"Mall:aj~mill, G. '~1!ITl;I,"I: liIiliWj, 'Po tL 8!i11l'~, HlIaJl~413, JC7 ~20(1:U. 4_ L. [fI La,unp,ruill ...Im.. (il'l :1t6, lHl g9,~L ~ :5. T. Wit:la tr, Qt", ~1I\IrU:9, :;~!JI\! 'lOOlI, 6. \\I, .11~ui:n ill" :)lflil'U 2]0, l~BO(1'II'5'5J, iI! 7. A.Mailtl. 0, IIW,~~di?,.A.1'1llruJi~ 1It1. A, IQltil1. i Pi!rtl,opt1tMlll!!~. nll~ ~:l.9'!l'llJ, 3. 8;.

IF. '. !l~, 0 19~5I.

himilire ~

NWI!. ~M~l!!1j!~~. l~~.

G :EOCI-I EMiSTRY 'Chil'lJiji{~.\all1>!!!·Y~~~ @I !aVoil ~'11 n 11~ ,F~~~~ ltl:l~ natll~~,Dr ti'te reeks deep 11i!h 1~,utl1that t meLted a rid ro~ ~IJ g;eif1Je.ra~E! p ~(i;!'it;VI) lea naes, s

'OI.C1illih: ilnl.pi~u~S. hlll,V!I::' thcpewer ~o fie':iru~peEmtb'~ I<md.'iCi"pe.ui,r:rdimmc. . and an"ect ii Ie, To llllder'lll.fiuRi h~w I hf S M,~ukli~u.irel'i 1.11.11 ~l' d~.", ~,U!"ntile E;ul!h ~1'1 \,n; I:;;anl c.\acl,l~ wlmtldmll of 11tK:'k nm:i[s. to prodace m:~n'u!i, and Ihe chl;lmil>l:~Yof Ih is scuree reek, These ;1m fU.n1lI31:nern~~~ pro~I:l:!ll\; illl g~"OI~ o~)'. OlP1d ~h-ey~re Olh,;1]~lWlng Dln~ln['JNl [~im:cul! to understaud, Oil pa:!,!..: 4 12 in 1.11 issue, is, Solml~'!,.'1•.'1 flf. ~I ) describe .1 meihod 1i:11: id:mli~ fying :';OlIm !In.ll~~:so!lllftl:l fl.dlic\i\fId c~u:illlli1bd;: of l~m ,~~fond for "'P~.m{l~ in dl(;,~n.w thai lht:y mdt 1.0 lPfo\lid~ the magl1l'!lil$. thai crILl ..:r~tpl

to the surface. To uaderstand ,,\11!lilt Sebotev at (1/, have dUIT!~ .lm1: th~r.mrn ifit'1alionf; ~hal go beyo.nll Earth science, 'W'C n~l;)dto 3tan with ~
n:frc:;ht"fillt geology. ~r~h S lliiI,mDleolil::>:i sts mosdy ofperk!otitc'. c a m~{; dt'h ~Ifldle rJmil1lt:r.a~(ll iv:iJlI;l {Mg. FehSiO", \¥ hen per idrll he pilr~i3l1y nm~hs. ]~C ~rl~~ids I col klcI t'O magmas Ilml. rise to the crust gj ve oil' i'lyrm,\:i:'Il: [Cfi:l ~:Ig.,]i:)Sii'r;1 ;:Lindp~.ag~m,l::J!.: i>' HCa,NaJ(A 11.;lSia'J )O~l, 1\1rti~1Jm,of~llese (mteu; 1'\lI)!~~ enn 'bi.;l rtl~ydoobnc:r,;. Lillo llID!illa~lh~ m !;;~bd\I~':t~PIl ";:Qn~ ..ml b~~.(wv1~liA.'kenedt'n~~i" n!.:l.l:t~ I~~ecthL: figure). n,!.: t~yct~d~li:s.al[~£
CfllSI

Iike SO" (0,. and :~I,O.and sdJidilY to b,~:-,,~It.,I !'ock, rt~ll in the m~Il~mb; cl i!,l{1'~M~,

the milall'lt~.e<II~ subd uctlo n .i!(lInes {2. ii, 6', COl'ltineii'llal CWSl (Ilmwn) !orliMat. ~!iIMIJ(!.iQI1i ,tOn~ and Cd 111be' 1fl'qN;~~dlI;'Ih(lJ] illni~~IiI~ ~y d!l~~ill ..~i~J1(5, t;5) ,AU Jt;1I1l~\ ~brQwn) i$ ~ri:lln lor~~1iI tQ pyft;l:ll~~i~~(brooml s when recye.le<! ..Grt-e.n o'Ii'IIOW deii'loles. ifiBe~I~i1g p~Fidoli~e. Ft~tlllrrn'i'l d~ii'la.tes ~l!lli'lg p~ro~ei1li~e. R«~dei!!l e:~iJSI m~y be- i!lis,1.iri !J,ulied uJlifo md~ thr@l!!ghol!lt ~H'!emiJlnllle,. ,or iI !11i!.~ b@ c:o!lc:et'!Ualoo in Ci!l!rMin heF!1ls\p~1er\'s 0' depUl:s.,OuM.ai thitknes~ is e.i'!<Igg~rilJtecli clarit!!', hit rang:es hom ~fi Iv 40 km ,<II the present t~me. roOr I!Jll,~c:UlIg ls 'expect~ 1'0rcdu~ iI;!f\lS\ ~i;Id~m~lIsiQ!'Is r!;mgill9 fro [11mla"liIl1Ie~ers to k~offilel'~rs_

Models of IE,arth'!lILJrliliti'lIM:I mi'lIIUI.e. (ke'ilrlil; U1iJ~U[}ruMl) is solidiim:f:!d 'liquid tilat fCWi1lS brlPlillfti a t 1i1iI~lting or m~IlIUQ ~eil'idcl1il@(gW'llerMj<lItI(Ice.mi.c rid~~s,: ~(Io£lll'tnler witilsooi rrnel!lt, oc.eal1ic CI'lJM calli be recyclc'.d back il'll~

~stm11S~Omle,1
nm~\y pil~

~1i~111', 5o'Ci~~~~d b~l1!ll.~

to" !:IfW ro,k C<1i~eclpyl,'('IN:~ it i~ ~udilllf~pyto~ri6Jl

~I'kl.,h

Mil'ull[.w-Ih 's,!;''OI'I!. (ir lllfa:Jclull.::S

be

u:il~x\"'tI

been des.a-ilboed as lHflfi'lle cake (.J). plmw r~d~ and lir<l'Ce-.eleme:1'Itge(lch~m~$try(lr~iiJ.\'>lS;iIt the "lUrf:le~+ N1?'v in~.etp.tct:I.~t~nl:!_~ :'!l~i~~tIJmt ~tin£: {J~, 'P'l.Ml:Jk"tt~ ,t41. ;JlJd .g;lIit!bo Ol {~~the: S ri:gt!feJ, \cbk<l!1o~ li~e thn..w of ~~"I\l"ilii ';;;In.. wit rmnw ~I!nic i:,;I;I!ld~ m~h~g, from, rmnnh;l 1PL:t.idotik ili;,1.t hml 1b!..~c:UiI ~.t'iocljfictl by tncll.iS, 'lrw~n SOlIi'Ct:rocb ClJ:rI:iil5ling (rr~.riJutnt~ i andklf pyro~'{elllrte fllCiml'ie'Cyded cm:;t. SODol:e" t~ml. ~1o'!;,!'edhrol~gh it t 7. H)c<l process C'<I1led ~1~'la!irll:ri~!,tisIiJ1 ~ 9)_ BI,."C;;J,U~UE 1li!;I~oe~ rio di I::' el (~t d~db~ .~~PlltlDh[ld lor idt!ti:til}'QIi,g llilil> mc~ Ii:reltee '101m illom of Itlllith::J~nll1~,for C:O;:~IBlp'le. OOrodrtfil.IJ,~ ,~lk:-I\'~~I t)'l1HIll.V<lS oil VOIc.rnloc~.
s.ubo~~ t!l'al [[li.'W'r.i:'D'ii~k'tll:h~ttnalli>' vul.C!l~
IH.'lCS ml!l~,.,'tJ I~O:ln fie(;)icied. c m~l.
I~

lJa&.::.imr~ll1~ iTIl:lilUh.:: \\'ith

Ih~~l.I1!i\\:

'I'.'il'iI.:'lIiJ!:r

ic It:i'i&. liij'i t"tli1.cICitlitr.!iitt:c1 in

the t:fill!il or

Th!e ,~~lhtJr ~~,i~ Ihe' DIii~liIf~m~.!!1 Ge-oiogi tiJl~i·MCet I!uloger:o un [ye~il:l'; ~i~(i'l~[I~"y. I oaa'5~, US;I'I, E"I'!li!~I: ~
he,lZberg®lti,lfTU!Q~5,~

or

l!ii' '1~'h:m~~('Iimatiz(!li lle:l:klotiltt::. us:~ns it .. s ,~ <~<.'Ondll~iolll It!;I~ i,~ nC;l.1I" 61, 1-[00M.:!v~r.'th~!1;,! h,~~ ~~L'11 ~h"l'llY;:~ trolccr CM h~'~''liIbi~lI.m~~,md .ruul:uni(lu.!;I 7.1<1} A ~:i\:3kthroug:h (,:I,m'!!'whim S-t!~mlcv l'l tj!. l'M.:'l!n li/OIli1iJ!.: inlli.gui'ly wi~h past fli1i,c~llo(ls of 9 I( /O~ !1h(fI.\lI;:dtlmt IJ1C nicke~ I:flntl;.'lll!S ,uFnmllY idem it)'i'll~ :L'lX ycled c m:11il<I,'iCd<,111~he ~~owpc

378

20 APRil 2007

VOL 316

SC~ ENCE, www.5ciencemas.iorg

PERSPECTIVES
oO,livine IT)<;''t,~I)1 ~n I:~..wai 1m} I<~''''\'il$\\~fe higher thlun Lho5.e tl~",pccl:cd j"Wlll 1.n.1llIt~of :p~ridi!ni~(;, ~L!I1d d~ey pru~rr~'d ~o ~ph.jl.l this with ;l!, :~'. eled !!.':n1.~t SOIl!OO!.! tllS[b~U.l.B~1tII 1:~l1Igcr.ingml1ibi;.suit}' i~Ih.u uhighl11ckel eomem innl~\~m: C:;Ul ~LIMI ~r:ii>'!J wh!.:ll pcri.do~i!IO .i~ 'l"llri.C'iM.~ iUl PS'rox· eD1i'1!by Indl-rock reae lion U 1). Suppo niD1ig evidence for ihc lX:cyelm![,..;C.li.lsl.i.~[crprcnniul1 ,(W~ comes rL'On~ ~he caleium corlt.emil'" 0]' the
HU'!.'li'::Librl b\'as_ whieh .1r,t' ~[]O[ll(]w lObe t::nsiIiy cx p I;I~ by meh in,g peri:dm i,ll!' l/2). N~,c~d sed and c<l~ciull1 are !here~hre telling the same :SliOry.ll1Hl is, lhe main shk:1d-lmi I~cli'n£J"'.... rs al
HU\'li;;liu w~~ rndl;edlh.HIli ;ipy[(:t~T1:ile MlUfl::C'

cn~s~ i~ :lfl irnpertant

H.:!I'I~~1 ij:m

seurce rod Fowthe

i!iil:.'mm ( i .W. J 2). iwltL:rca::~inLILaSl)~ Ill"li<'.!~dl p~rid~)ti~~i:oLh~ ~';itlun:~vu~k t~c fmr Ctmk ishmtlls ,t :1'1. All, olll.smJniLIing qln!tllion is

dm~d aml made OI\'Olil<lole fOfempti,ve lleedhng, mllJ Emlh's, bill'sph..:n: coulJ 1.1a\'~Ilvulv!Jd .. long d iJfi;1f(."!lt put.I1\Il'Oi..'I'~,
!tell,~rienG:e'5,
1::[

whethef pe.~idoti~e' srmrce~, beccrne metasoltlIal.~ zsd b,y ~!ildr~ .recycled CI':iJ.'iIt ( 13 I Of lim so:me oiher wily ~7. 8,,,
Fntm'c surdles Ill.ight. aHC}\~'us to lrJIlSfuJll'l

1. A.'I.'sooo..ey

01,., 5r.M~,! 316,412

jjl1Vin~ 2 9 ~i!lh

pubMIliI!d 21]01 (ID,'lI2,f.rsdel1~I!·.:ru,f:l1:11I,

{2'oon:

our picture

from hypotineticalmnodels: recycled

to

act ual 'thret;'"dil.nerll>.t~llf1l.'L1views ~hr'l\yililg 'lime

sizc~md dL~lri bll~i.ornof c mst in th~

3• .I, ~,I&rg!!!l, W.J, il&;Jr!llan.r~I'1.Il~I. Sol'. ur,!. ::!,.;I'Q. .?1~n9~9}. ,>to W,,;!!b{(! ~1 eJ ,~" Ml~u~ ,~l:4,(!;~:I.;l'OOSt l ~. 'I), l $IlICffii!lI\.New 1lt'eot}' ~ !1rf!~1! (Cam brid~'

z, C 1. AI~r •• D'.l JurooU~'.l'laloo'i1' ~3.

12'l1(11'O\1I6),

rrmmh::. Theirspl icmi()!l~ go fur bt.j"Ond l!leol(I~W. For example, It '[Im,y be no sl.!rpr.i sellu'l:t Sobolav 1;'/ fit, U} klemily rYmNcn~~e us the

rock .hm required] IJlc JnYO~VCmilcn~ol·~C1::yt:~cd c In'~~ pn~pflsed by S nlmh::vi;.'l at. U The a~ ~LLilJmu3g() I'tH1:ht:r in th;;L~ [hey ~x~unintt IhlL:: .pl·nbIe;Ill tll<ll.ui ~~ \\'hen llIil!;t(;::l.co,lcilun, .uld

m,

rock lhm mehed m preduce

ma:;;!>h1t': in :;iL'I,I~ ~hLl,~. it !H,m; eX~imlCli{m ul-We

TnLp:.J. This W:~!i a :I.n'lgm~tk flood

i,h.e sibci"i:li1D {lilian" se

tLigg\i:r\.':Ji
nil

11u: brgl!:M

IlJfiw; f~~ (,'.'Ifijbrid~ iliI. 2. :lOO]), ,rio. A,. W, HO'froonll. W. 11\ \!lhilP., faith flamr, SCi. WI. !i7" 4l1U9~2}. t, ~,l'il~i d al.. EMIit /'Wilt,[. Sci. It'rr. ~~~, '1<16{2lHl5<). a y, r.lll!.'"',I. O'I!Idrn,l. ,~p~fi.Il'f!l,10B' l2C,~ ~2'003:), \1_ j..n_ Bl!)dil1ierl!lqi" f. ~IJI. 4$, 1 (~O~}liID. 11<. 'l!_ '5Ilbolel! fl Nll1!J(~ ~;l4, 5:~ t~OOs.t n, 'p_ B, ~I~met uL, .€l!I'f(h fl~r_ $(f. l,eI!_1I.6~. ~ ~7

i!l'..

mang anese arc UM:lCi. iI:m:lthey C'j(,~I;Il.d Ihc :;;[~lysis 10 a 1<I:vg1':rp()plll.'H~\l:n (l f volcanees, a Their re.iult.., w~l:h(lrwith mhef n;:C<GI1;t 1>Hj.dil:':~ ,[ N\ ,I J). sh~lIiovhii!il .iI, uS ~mlikCl.y thtll a ::;.imlgk i, t ruck ty~)~ wi~1 be an ;t,ppmlninie ::;mlfl:c for ali IJcennh: \!ok:mwe:>. Filii" example, reeycled

E..u:th, :;;ome 2.:'iO mi,~~ilJfI Yl."affi;:!gu l/·n A.II.h(mgh the (:;.;m~l C,~III~J liinks ~m>Jin poorly tl~d!mlioo"irl detail, ollie:~tll1,~asomlbly ilillt~~ililll ~IditT~reni mnCmllle ~fthe mantle cl iet I~ad less P)1I'O):l::n il,e iH1.dil!'J(l~ p.;.rCcltld.t.t.:.Llnder il:h~sc circunrs~am:es. less niHLgrl'l3~""nlqM h.1.vebeenpro-

U,9'M!~•
10l. (. !iln~ r'lor:~4'14. OO~~jjI!lO~~,

H. 'II:, 0iI~~1~ ('J aj", J, f'l!.frfil.<ll. M, ~~&" 1.4. II" t'I,,('.amjlbcll RI)/~ Sd~'t ~53, 17~(Hl'il'9.2).
l!l. (. H~

~r ~I" fO{.lil'i).

MilltmJ, ~I,

8~.'11.1%'.3),

PiJI)1i!1'I1!d !l!mliiil! Zi' ~Iflim ilD01: ]D,:U:i!~I!:i!fKi!,n410~1 ~mdl!ld~!his imftlmmalioal when (itim~ ihiis ,(:lilper.

BIIOCHEMISTRY

An A:nciie'ot and lntlmate

Partnersh ip

I'!\Icll'lhe mllkon enz:r'"u~ ea n G'IL:a lYle .~ wide villrldy !fIf &i,enni;cat J~(tiOIl $.. l'W\D !itl:ldies Shllll" how ln111 '!lie r:l:.alUil~ i~ 'U:lriiIlVEl!lil.

11IIO~t mxie by"pJ.~Jd~lC~. saperexide, by ~,.I0l1C~ mill lilly ]11 ~~s.Fet IU ('i<,,,~di'ltion Slate, The <!bun· UIi!!!:il. rn 'the less tOXLC ~lJ'd[{)gl:lnl p!Ouu.,(~dt'!_ thull. re{ n} soaked up lli1l~t"<lI.liyrnolseulur oxyKllVflkv~ and Lipscomb rcpon ml 1.1l!!'baegen gcncrseed by p~lotOSY~ldl.csis.Comp~c:l: mrLJltltX'li:Lli.:lr (ytgt'lill i;;.mllli t~Jtild nut (''iI'ohc lIn~i~ l.erilll C"1Eti'l.airll ri ng.el.ciJ.v in,g d~.(u;.yg~naSC'.iI key cl'lzylVtte in the mel<lbfll~c (fegll.~dnlio.~l of subr:mmtl:;' I <Ul1mm~sor ON:yse~lbecdlme ;wail~ able ilfl E;.lL1ih!.; O1ummptle~ - 2. biltiol1 _'l14;::,!n; :_.~o_ <trmn~,i~ hydnx;O:Ubnm l:ru:mdU~1ro iDiln :i;oll by h.llUIIlSJlilS., The crill)!m;e comaim;; II miC!l'l'mmc:~ear ThC:D1Idtc k"lblc.ii w:rre UII'It~: o.rgal1i1iln~ used

- -B

flFO:~.11t~lIl\_ I;i\,(l~.ln.u.UimUf p~Clla.s ~. 1I.Il¢:\ .. ... ~ ..'i~

... .) lnlholl ye;11$ <~:[,l!fl. bani!! S OlWUf,· J sphere \V<lS retll[]cing. aml iron was,

!'"JllidUy, 01[1 p'l,g,e 449.

bu:lligbly loxit' and ~.lmJtI.111i be removed Katorm (.>1 ~d.,t':~reveal details of how (Jn~(llu:y:m~helps tu OCllM:Nc lhc
(:,11]

Chl.::lllistri,L;!> ill. [IlG s y.ntiE:...si~ I[] r I1mJro!tl:Ul~~nit~ H;'~;~!ltih(~oti(,"s. phm!. hflrnmonJ;:s ~3t fhe mld aerhors ~dd~dI,aIi'D. nefficicet SubSlf.::Jilu I with i ii slew.' l;onn~:lrS.iol'l r,ml!) to tile CL')"stJUizeJ enzynlle un:ul O, .• ~ll~.et.L"-cl"'lmilOsphen;\. rapd B)' illly frel;!zing line crystal, thcywcrrc able w sim.tal]nm.t:iCIusly trap L~ro!.! ,~mi(;ffii.usieps il~ ihl: c:al:a~yzedl U'eilcti,oJl. Jml.l acts <IS ,I conduit m

en,lble e r~~mn d'ens~ty tu rIiImnermm~ll~ ,~m·

D:ilmic mlimrmc
~I.:es lWO 10 tilioxygcu.

ret I~,)Ul m,-..;:~!'>!he oxidative

pO\II'l:I:ormnlec~~·

tar !)xY£'CIII..~hus, d:mmat Leal ~y IHoreJsi ng, ~h,,;: ,me. 3. Ii1Il(mr g!l d.itlen;'llIt eIJ'kieiflcy of ~lel~~mik elle:rg;' gl::nL:lJ;<:!!ioPII, Two l:epuliS ~lil H~l>m:: i:hi~, hig~.~ighi.th~ 'O:i.o· 1!~iclll intifll~K)' of I hel!el<lti~'ln ~elw~cu Fel·n ~ IIlId do m;;ygml,. Olfl p~lgl:l 453. KO\rd~~V".J :unDl IJ fl~O~.lh ,I} !\.!;)ow h(lw ;!ll er:lrym~ l'i!l~'111!):~ dlC' prl~p:rth_"S 1::1.1 to ae~.i~t'h; 01' tlt1..'f~y .II} eUlJbling the oYlidiJ.tion of <IITmmtic dio:ls in~' li1ur-d~t:LroUi!11~llloo~ ic:ptol:;:~ll_ Hm~\I~r. ~he by-p,ruJUCb. l'I101~cu~ ~."'ygen n.:JtlCt:i,LH1 ill'

re~ ~h(l(L~dtot:\.\O II

1h is

m!l,n5r~if'ct\\'·

hi.~iidf Res ;1Il'd <I carOO.."9'1· Ihat has evol\'e<l !>e'vc'r.altilVtl!cs

eI17~ynl~2i. lh~t

~:ut:dy;-:~ II;:ru~~ijOlI

1'e<t<Clive s~l!et:ies: il Slllpef.'O~Wand a sub" sua;ue .~i(;JI,. The ;supemxo species is bound side-(~n to ~he irfnl, h~~~ int'~"n;:~~mi1'l W\."l.I~ the

or

or

11~IIl·dijUV'9i~,!il r~t~h~ns.IMpi5, Mon.Oil'LUC [(lair no n il~1lI'iI! ",ron e,nz~m(!i til n <lOCO rnmodate 01 di\i¥rsfo! Ir(m~e of Th~ oi!,uthor i~ 'i rnlhe [[)ej;!~rt!f!enl (I!'f 6io,llne(ill~n1, geoil1lelnes I,'jJth SLl~.Slril:~S and dio~~ge,n,<I~ i!hJ5t~a~eti here lb~ uhelrdppHi cataiylic il1lt~rll1lilt'[IIMe~in an 1l'I~lecu~rB'ir;Jlog)'a~d aq!l~~iti, lI'!1ilfE'~~ cWinnli1lOl<l, by Mi!'!!1!!:'<IJHl!!i~N S-M5S., U5J!, E'iNil: wi~I1!!ilOO4@llIrnn, Q'*~.dt(jl riiflgi·dwvirtgl diQ;X}'g~nn~ (~~ iI~od ~nt~r) I:VPOI1~d ~"'lllw,;l ~1'II!ll1:ip-'.oo~b~rtd jill sup~fQrld~ ~ ~;d:1i! redl!lr.r.l~C 'Irighit:l reported by Jl{a~!JiI'IiI elal.

ViMIVII':sdencemagr,Qr(!

SCIENCE

VOl.316

2.0APRIL.2007

PERSPECTIVES
Cry,\\t .•11Ilou:king [[l,IIOIyed 01 erueial role ~1l1 :,tm vfled by the~If Pflf',ph..yri!l [~uftnen:;" TI1is rac~ Ix:cal[qli(,lpf ~!el'ic CIr{w ..\ll ifl.\l ficm the II,,"e o~heU' tu.~nJ~ III IItL:m~"lli~seethcflgUfl~.I:~fl pallL;lJ, tt':!ippin~ these various species. rill, btll.~:SIuJil.:'i>. Wi!tlIJ'IIJ~y~xp~ai ns Ihl'i gr(!aL clhrt:J.'Sii~1'or sub! $tr:I~~ chen~i~,rie,s th~t C"UJl b~ ,l;".;±wlyzcd by ~hL;' bl !h. 'l:rrdn,~""m~I:I~~ ~lpeW;'l:fll ~:;; [he lPL;lde(;~~~yl~W~IW IJc~d In g~mJcr"lc ~.11~hL;!ilul,lnl~~d"'Lles J:.'Ilmt:from a single;crystal. In each Cl'ISC. dill~ nonhcmc c:nzyme.<;. 91~.gncd 'm mta~'k ~hc !illbS'U1!h~'rm:lic.i:1jand fOllll Wil.~:e i111LlfmlJ!poeLldtet~! i eopies ofth:e ~crh~e sire The' :dllmrn:mc:e or iron flH ~nh heh:ll~lck the <llky~pero:'!\n i'lllermooia~e fsee the figure. in ~ll!l:Di<LlI.i;t' cr}'st::L~ b~lildunJ:l o~tJck.~INlMillg tlru db'Vt:~!}Pn1t:liU C(HII'lP~.GX life dluri.ng the of eenrer p;~nd l. ~l'!lum:Sl.i~lgly.[hi,] only OUII,:r.t"i'yi>" t~]~ SU'LJCl~lfe of d~.o.."yge~lll.illl complex v,,,.i.th.;Ill di i 11:1JgcS ~hcact ive sae to be l.:5-bm.iI!lIDi.-ycar msting event tlmt kept atmo(Ibsen-ttl sA um:l.hl1It'!..'Ol.isly. di Iferem species The mOI,mirnut:.h:"1Iltllun:l'lcmc iron (11ily~U(;r;J r''':;ics,k(: .. u spheric 'il:>iygcn k';o'l:~s,~O'I"', .It isurltt:rust~llfl i.u ~m~y n:mnifew.'d ~he;JllSe~Ve~ strucin \','Ol'Ickr wimt Iiife \'11:0 LIM be like m this poil1ll. ill dioxygenase, with oxygen most hkely ~!..S' ill! e;tl,:l!n n1Cl<1.t:I.i\/l! sites, lrnl~ru;}!\J:}UB ",,~ydWg,(::olo~i~time if iron. was less a1Juru:~lImlillit! II h:(m)"r~rmw species] <11:'0 11lli\,mL~1.,,}'g~n, Im:nlty dii.:oOt~ li'erentcrystal padkiug enviromm!llt~ s~abi~iz".'d the t.'Vt)~LIlion,of eoreplex I1[]JUltli.eeIlul:li r organOOlHld S~dC4Jll to IliaC ili'tl'Il (Jl (~irren::nt~r~ck~ is beyond~he reSf'i~lIlio!ilof ist'll!\.h~ld )1~'!'I.1edl bi 11 ion ye';;'lrll eariie:!!: WOl~ld K.uXJnOlC" al. :\;tudied Sll!~J{!(!; tde reductase, ;m ellZj'll]e~illlt IS ,fmmc:] ill 8 :r.mgc of ~QCleri~, .liIese dk1ta;suhlle slru.callrn ~ e frects· '\,"Ii thin. .~c iml'l slill have such ~~ plUmim:nt role in the F3b" ;limd u i't'llm~'1'15l upero:.\.iJe fI_"(IHt'W~l;::cOIWI;?[~ pmt'eil'l pmb;Jbiy cornml d'l.~chernistry, Th~ ric (]1'lul~? I [jt:~i~\'1;: Dt\\'Ou.ld. Th~ chemic.d S tiltldiy by KmofUu'l {It, shQ\'I,"liilhm th",ooll1lbhlnri{lI~I~CSS, J UllI»1U11Ctil by 1~.iS \110: rsail to L11l11Sill:ilUll supcrolilJdIl ('O,~I 10 hydro~11I pcm,,,ith: (1-1,0.) ~Il~.~u~e~eJecl'i1J~l reducuon, Re,!(,:t~veG:\:yYel1 metal 'l.nn..:I,dbe cl iIIk:u lt ~;nreplaee .• md ~ife lion ol'hi.gti.-reiloiLltiO'Ll .'i11CClrmcolli.c experi111I:lilt~,(b~nh in the crJ~L<I,I. O1:liid in !;ulm~~m), wUlIlllIcc:IfIOlUlJJlyb~ 1t:1S:.'iU,UiI~~lPl!' ..ling, wUI:llol,JI~ ~1". " :;pL'Ciil:~\such ,~"0, "-:,p.i<IY OIl boy roll: hl hLilil101Il ~~mp~lt1Lttiam~ nl~thmjs" OInd (;ry~!:II: :iilrll~'Iu~ Im~hokl~}' tn'ld ;;Lg~nM' ,lUbd tht:i~ d~tn.t(.':limii by iteor8eii!:e:5·!!!i!d Notes CililZ-y[ll£.~ such ;J.S ::.upero.:dtk rucllJ";:~~iI.:s therei rup:t\:liCfiitS a p1.~h;:m ttmlbolll. th all pnlm~:!'iC:S to ~.. L G..~[ey~,lll.I~[ooW,.sriefj{£' 316,. 4<51 f2C107t IbN (.!if great ~Iliter-e!!l,IK~torKI to'f (71,lreoi:ze~ t!l!lh!ld; m,my secrets in chemicel biohJ"~)'. :2" ~ KairJIE ~'01., .xiej'J(1!' ~;'il£.. 44,,) l1!mm .. -rogC-lh~r. th~ two ~hlIJi~s pnl' v~dlll:iltJlln.~[I,~ m!pp~d scn~tr.1.1 rJt."tOxide coffilftun11Lt~m'ls with :3.. K" 1:1. I(gEfl~IDP. ] .. f: Eml!'rs;m, I!... Qu~ !r" , .. i!tillll, fu~ .. (h.wt 1.0, ;a7' 1[1':OCiH illlsi~ht~ inlo lite dh~n;c ::;1mclur<l1 rel.Jiiol1::; F~~U ill a c rry~t<ll of an cnz)TIile mllt,mL~h<li 4. A. tl:"~Oi:ll ,~i.oJ~ :SOMa tl:99, llH~ uoon. Ll~alm0ii10l1~H,::lcar ImLlh'f..11l.~~f\[Jn ~"tIzyum::~ een :>~~ li:tts till~ Feo( .I)~p~roxo .. ln emurast t.!l bi H ~. \I. Nn.IH",-. M, IHlnltt'al!l!, t, IliIbl, ,Mtll'f. di~m,9, lI:ll9 broke'a" ;;Jmlll1g (I1'!:Y g!:n. (Irolei n, oo,lvent ..md the snuctures r,epm1~d by KO\~iI:"~I~'r t: the a {~OO4~. clttI.!';Y!!lI.'.'U i:siioLIli.cllU the ~Hnl ~li.d"o:UleOJ'Ifi~~~ hydl'.oc;,::umOI1i :!;LLIw:;.;mll~s" Tille irOJ]l C'cI11C!:p; iPl 6. ~~pp~ bj' INII!! ~i'.ltIl GM·665!S9. IJ~C:;CCfii1.y~nes~re ltuliidn n~.(}:rc flexible IhoEtuli I~.i I( iSCC the l'igHl\':. I'1ighl ~rh,;:II. ion The di!'tLn~l names Ih:r the three Q.\id;ni~!l '.~o:->e'i;lhh:eir~eme 1,.'fI11)lin~.whkli! are cen~;R!es or di~l.xygen j 'Iml()~ecLlI:H')~Y,l!.-en. ~"lIJ~eli'~

~um rnel1cnt

or

ide, omdp<I,;'frn;id,.-l fm:-u.s OIue!lliun ou eleemm 1t"'::In1iS~:~1i cv~m~~,. liOlU·CCl> and H'ml sj{:,I" but
1I.....

oWtwo [N'ntonil,<IfC 1l18(llr~liired ror the redl[qc'~iol1iJifOlill 02:'~Ltl Hl:02• T~r.J PH:lI!CJ1!> ean Ci(]mu lill1l1 ,~~mi ,~t:'~ s. t:'o folt"L{lfl'. ur 'I.'I"I!I;l!f. a!lld no d ciu;ymncs I.:omro~ lht: prmlJmll:ion SICPS in pedecl s)"nchml1l,}' !with e;k'{:t!r~llfl Ir.m~re!:: Ahhoug~:I'[iIIll' iroUil"!l.Ii.()X)'~LJIl ,';;lIl1Le~urus ~)ru-

PlANIE.TAIRY SCIE.NCE

~n~~d hy K:!u'n~:! ~t al. ,blfe <111 un "poeroX:iJ F~ slP~ck .... d'iH:Y J.h;;phIY three J.~ ~ ffcrem Crnlcl-on peroxide oonf()J'm,~lio:ns lMOllgli. .~UlJto.1'lOil
lll.,- PeOn) pi\,("l~. 1.l1us;Iin,lki!l:Ig~h[;' pero"idl: ~o \!'1l~C1' .Ino~eclllcsam~ t1Ui11hm [IC~d5 nUilI~ll;! 'lCtl\'( si.l-e, TIte t'tl-llC'l:ncd rl1ItWcm.cmiO onhc~ pl~y~

M@dulatiiDlu ,or riU;]l (I ell1lission5,fronll s,<llLJm that h:.aVel' been L1r,edlto FI'UNl:S'LJ ret.he rot,niCl n D~ lh~ pt~H!d:sI'flil~riofn"w turn Qut to be {;ol'lJ~~d tllll ~:d~;!'lal.img of placsm~ ..fh tru~ int~m~t by r

rotation

eli'Ssuggest p()ssib~e Iwaton IrnlilsIt.T 1~i:llhwilYS. :lllcll~ i" study .WpT~M~ll!t~lhelllm!\t mll~pi~u;:
,~nliig~ll1 :imo p.womnift.tkm dm:mgh m'l~lioLlU)

:r~i.cetdb. lL:oi how

'

(li)l"It.i II LI-eS

to

Doe

a .i1tIy5tery.
il,IS

~DC m'~lkm rille of 3.plim-:a is~non.Emlflt!i~

il'lil~milmm. prupt:I'IICS, hlT

s:ohilil
Sllr~

pliln:;t This; mise:> It~\' llltooio:ns ~Ihout ow \w h fiJi: In m.enSm\; ailDli ~Ti'l(IlL;iJ'$mm:l 1111: .nJtmi(,lIl (If
t~'r!! Imye

planets, s~mp'l!eobsef\'<LI~OL'l of tile

ra..l·I:~

gas p:hi:lll'CtS,
~h!l!Jnbf(lllllldi;.'d piun~[~ry
fl~l'Xl-

{lbjiect~~, ~l1lnl1illl!} Oi!1

SUtlUiiilnn;t riSI.S, ", ..ito

d;lt-e [::-;oeeth'eilloviein (!)]. It $hml~d be no-ted ~hl!1 crrys~a Istn~c~ures. of plJlOte~ll,:;are I.""m:lyJI a high~!l.{lL~gh ~·e.'i(l~uian l'O d urecl~.y de'l.e:rlllulloe L ei[h~,.l" o;>;~d.~liolll or protml!1lilJl1I ~hllt~ of..:: I:nml~ [C-<I Ig IJiOllp$, nese ;:n-e illldi:r-ect ~yirnfeFlre"jl fh"11l1 ~~om£l+u~1> c,ltrl-lp.~~J \kilh ch!;':n~.i~~i~..:nm,\'I t~~~g;e. i~lh.e Rt:~dy by Kov"le~' ;;i.!i!dlirm .. SolXlmn .In th..: :;tLu:ly by Katofl.~~i at,. this lum:eriairaty is .emm'e'd thl,'o~lgh the Lise or
RIIQn~!fi~fJ'e~:trm;cnpy '0\11 Ih~ !i;;.r.)"~h!.l, Wlli,L;ii

lIir~cdy '!.:.'UllIlIi'Jns Ihm ond,), F,.;~In ~-p~ru1o:0 SJll'Ci~~ pn:~:ru. DCii'l~ilY liJl1t'~ionall :11'1: ahC:Clry ~<I;lcll~;<'iI iOllls PCilfolr.me~1oal:ilc stnlC! LIre::: l~~ S ~I.;l~t t],l,1t .n II;';I~I LWl:)~~re 1n.ycl'mlperol'l.idJ:ll. ~he
sing~;y p~Olol'i~t'l'~ int~li'Iinl.!,t.li;!J.~ or pL!mxid.c dulfin~ it~ ('l'I'Lw!;;,l'l'i:on tn IlftlodlH,,1 ~:liO~.

~~ .. p~iflncl'S st,lchHi>SmlUm, or J~np'~lCr.,h~~vt·li", .he \'~~ibJeOUK!T la.),el'S mm"t!' 'u iWfcrellitly Ih,m I~e IlilleriQ.r: So Itow do VI,"(::kmJwl~ow 1:'.'I5~.~e g.~teri{ll· gl.!~ plilllOf;::l i!'> ~p:i I:Illi!,1~'! l1.-e I,.IsUOLI 1 I.nck is 1.0 1l1IeaSLlfC l~1!..: .;:rim:IJcu'ty (1 rRLd~oi;;ml~ p s·LQrn;.lhilt ~re ;lnod~l~mecl b}' Ilile plrnlle't's. illt(!uml 11Ii!;a:g~.eti!l; r~~~~ Iill~l:hocl !s!;UII1~tlmt 1:1iiI~ ,Tl~i::;. ! [w:>I~ne.ic field i:;; til~~~ ml!d th=lt Ih(l d}'nmJ~' ftghm \\~t~-re lh~ fldJ i,~ !,.."C!nCrutl,j :;pin~ t\ [}1JiJ.: lit repr'!!51:lmtath,;: of·lh:: bulk of tln: pbne~. Ilt~tOlm p!,1.~.~ 442 of Ih~!?i. i~~[oi;'!,fill[illeU ill ~/J. ~ Il ;~rg.u<l 11~1I1 rec~i1rt d:lIt~.lron.~ NASA'~,C~!i:ini~~lC1.:~"il1 o.U.Jit'ah: I,h~it 3prr1U~IUclll!i.L1.Ws in S1Itum\, spin cOLdd be '~'HUSi!ii.l by ~~l'OOesse~ 'C.X'~U",ll<l~ i~~e ~()

o:r,~

Vh~ a.'LUhCf i~~ll h~ i.Ili!iv.ef~iL)'III C!Jklri!d~ IBou·ld~r, ()O 1l!il3:[]9-fi~512;,U$.it ~rn~ll: ~~9~F1~~®(i!l11I~.;Ilfil,~iJ

study d:ymuna, madell> by ~.lfllVhlg. ~ III .ghly :;.ynnne-truc intern:..! luu;~!1'e~k field. A ~ i:ielod thill.1 :is S},1Il11l,lCui{)i:lbmu Ihe ronniorn iiIi<';j,IS ,'it)~Jt~ a ~},I:;.iclh.eoJ·~!n 'of m,l~netic d}r~1r!ln.<; (:,. Thcsccond pWI:?:zk:-~"um; w,ulll tll(;dct~"1:til~1fll of a sy;~tenWitic rouJI.i!{lIKI~ Hl,odutllim,l orthe rnclno!!lltli."",iun si mil ~r Lll :.I Ila:>:ilin!:!, s~iIt:i €. b \v!hie!:! sho~ W I1l)t fioi;:~!.!r~.~ ;)! ~Y!l~!1JiI·~r!(: n1!in.gii1~ti.c fi~lt!. Mt:t!:iJUwln.il.c .fm~io nho:us,UJlUlrillCllb Ill",,!:: l"evea'iJe<d Illal ~:]S1 year iJlaat &tlmrn·s {j,lj' ~;t~ be-cmne Ollbrrul (~ trl! g milm; IOrni:gl.:':t-il is 1IDm!.'r{mghly ]~ ~l)U'n.iUlti 4,; mill1r---'!>ilti!c tht: (I IIJ~[b \;,'I'lt:n m~;hLli'C'd by 111~VOy~J!5~r mlss:~{)m:I.n, ,I.\, IDid ~he :spin ... ;;eCIl:i s l>l~ kot.-..e]l' .J~e c'iII:ul!;ing. [U1 tll~ir i'!ElW !wtuk. Gun"len L'f (II. :~ltow !ttlW :&1Jturiril'SliiId no i.:n.,~~sOrr!. the m~gi Illel ic fi~~d mC'<L~lLwedin~he BTl,~m:ws~lhere"

20 APRil 2007

VOL 316

SC~ ENCE, www.5ciencemas,;org

PERSPECTIVES
and Ihe dt'n!sily of rl,~rm.lr.tlJ1ped ~D1 In,~~· t ~he im~tie field !J.n:: :I~~ I'Ilulill lated w:Uc.hd:ne £.:.lnlMJ dnf!ilfl!;l per~(l.Ij.And the pl<lS!llm prodll~~d lJy S~I,11m'~moon ~1il~c.lml~1smry hO:lJ tilt.! k~'Y ~tI I the I1UJ.J'Je [I'Ir~he!F'.!!c1io e<mi~sill11<;.
Ca~sini ~IfD.l:!g(!:S. of g~}'l;I.::!I1> SlIiUlrfl's, on 'Sllla]1 Jiloon I:n(dadus (., JI I riggcr.;;i1 great C'Xd~i.::m~n~. prin.1N'l.ri~y because the)' i~npiioo Ih~11 wunn, polemtially lire~l~arborirlg \\Ql:e I' migJu Inrk j l:I:s.l.bela~v ~he ucy ~ urtaee. 'Bu[ ~u un3,gllcl.oophcr.ic S(! i~lui$ts. ElloC'iailus'gc'yscrs

Imps lilleriii!',Hfe~cm;om!~\'<I~<l~ijons.If! the

~Clatc1<-

spitt!'ru (a~~i"C'[.ingrhc tlll:ciIrm! y.[l1Im~ e Co ~11" pl il1g I t}'i~f SO'!.I!I~.·ll·:;.lQ~}'e'iT {l!)b~1. t~U~Iii{)ll n::m~jl\. .~:WI~mC~Wi'L:Sd1CPIT", W'!lerJ ;I"'}frlm~e~il.:'cOrl,,'ec~im'l~~;mem'! .• he ln I.9~&" researchers lmlml.lu L~pl!linl \'a.ri,ct~oUil~~.UiI the Je pl3Sfl1il 1D.I:itl.S ( 7) by 1U1\'oking 3.00],}'!.II!C-

n~

[~C!Jl pmrem tluu nm:llcdw.iih lhe pland. But t:videsce of such <! now pauern in the jovien m'lgnelIJSrl1~re It:mitin'li clllil:oi!i,\'tL flUill1~l.lnOre_ recent mmlel ~Il~ ~fth:: 10 toeus ~815t~MS d1:Pl:[ Ilig!a-energy ekct1fo~!i. th,H n~mke up ;J mere rr;lc~iOIm the lOla ~esen <I lIluge effect on. .~c of

A flmdlum,mUll is.....te is '~!.ih~lherthe' rmlJ,!!1el£J5phc.ric ub.lO!..~n~l&ioru;. if1c.~li.t1UI:J.g LnUl radiu L,ll1ilvli:>iuIlS. !1,.·q,llire l~~ Ill"'~!lelit;: field '~!I!Ia!1u t~~:gi"ronm t~iJC inl.crior (If Saturn 10 bewi:>yunmt:1r rie, Nerrly 30 }'ei!1f!o.! :lg~l. S!e\ii:m!'J!~D1 ~ll~gg~:-lted tUI,1[.s.I:ru.l1g s]oc.lr m{Jl.iUJ1Si~1:1 COlldlK'tiUilg ~ilHJl~ sm.rtJWLmhmy tlTiem}'n3'IUo unig]~! impose symml;;l.ry tlJrnlll!lld (he' ~JUiliClJKII.axis ~1 O~.

evoked Ill~ ¥tdC1.1nic pllu!:n~s,of Ju~~ii'e'(~ raoon lo, lo's plume~ spew [orth g,!!.~es th~t UJ1e iOIl!"

Southweod recently rroP'O~'lI lIm! currems ;i u 11110 ri~l.!i.S III ight caWIl'iL:.1 d irml~ Ii I[ 10 I)!..: obscured t Ii), Thefact t~1il[ dle roL~lliol\j)~ odm ulation of !ll;,l;gnetospi'Jeric phCnO!il.lef!t!I . .-;;eeulliS to be ttlirJyl;.ullSlHllb~ wUlh mdiaidis!'1nce an:glles in Ilwor of ~LI'Iextemal r.arnet~ uu an lmernal SOllU'C!.':', Yi:llo""'i!li7~d magnetic :111nm ..die~ ~i.e," h~g'hUrul'lr nlllllifi'!)I~l.nhigfuJ la,lilud~
it 1f~~I.i ng

n::ill:J~11J possible ,;md Il1IllY be the t:uf'f'CiitlS ~hil.t c(JlLpl; the m~mtosrmere W lme nl,met As [h~ !llySt~ry of SUlU!"I'! S

1'lJt,;llk~llthh:k,cl1 s.~llc Ca ssin i :>piltk'~ral'i: l~g.at~~s ttl m-bil$ ltmt n l'e<lCI'l his~el' lat[l",f.J:s and unexr~arc:d J:i(lh~trugim~"oflilJll m~lgIlloJ;!tm'phere, If NAS.A eXl~ln.d,b 1'l~l1Idinsfor.~ few )~r:; he>'{ll'ld rhe mf.JI-lfmS end oflhe prim,c
micl;,:.;i.m:l, Cusr;.i:rai: UJl g;J[~lL:lr dOlw o Ill\lillSfilaJ p~5;~;ibJI~ties,,, Ge'l's!us on :Saturn's ~(m Enil:eladus. eilut ga'ies 'Ihatfo.rm a IJta~rna I.oM. The cenl rifu;g,al i n<$.labi~[1;\I' lM:'r ri:!..'8rly mlc"t~I:Ji!IiJ1~r rSilittrll":; ~eilI~~(lmll cycle. M;agm:tos~lhC:fi,~ th~t c.'!JU5~S pb~ nl~ Qu'tllaJ,lj ,~{)uld b~ slr'Qngi~r Olfl i:I!II~ sillc an;;! n !he o1hilltF.,Gu 111I~~l l c 5ug~sl Ul~~ ~8Jis il~mrnClnill; Ilt;l~ ~liJiIY m~i1ite<It a J:iilr{'r(!nl. rill!!:! lh<J'n SilIll!IJllll '~Inlerl.or. plfl.%ic.usis wiU hU\loL,:lU g[:nI(ILillllL~ d~ljll'JtIi!Ie nte<u1il1!;l (lftl'le C'L~in:i .. iled m 1!r.'t~c' aboUl :1 Hl'I1 perr seecml, :1'01"1111· 1)la~m;:!l. ElcctrJ.cal ClIIl'I'Cf:lLSl~al t~OUp~~ .~O:5 t11l~tu. MC:lln~iJ~il; l.h!L!8iI111i1tJii.p:ilcrie JYIl!tUlliIA1HI.... ~(l!he .ionosphere IU<lY per!mh cists :Ire I:ryillg ~l1 underswndho\'I.' the 201l!t!~ ill~ .1 dense !tllllid:IJ dD'll~d uf p~U~I.n;1 trJfl['!led pl:~~n~;~ [h~ db-C'twns_ Sud. p(l!f'l!Idmliun~ nriJ';'~y[h111"1 ,ilOn< .ilJ:l. ..~ ill JUlpit~r\ ~~lrm'lgmagnl':l:it: fi\;lid. Tln: ~tI[~lf;of Sillum's OOlild'mJ .aI'lnus,ph!Jru IlPPL:,iT to ,~e'J:"iJ;Hd v.lTu;;!!:trrnl~ if! i\,n~,~r i~iu~ p,Ia."!lUl em;m<lll~.!'lM from Em:eludus .~)l !l~tJ!ch l~adto.!S~{mm~mi~lb'l.~ 1\.h:em'lhvoely. :! ~}'~. ~h,H~!,':e\\ij~h [tlUII ub'n.cLllrJI w.~nd~ Jl!.apilc:r·s annosphere in ruination (Ln, But I)cfun; [hey can Slil)' Itow ~'t;Jkur (:It meet :.J couple of hu uuJreJ kill.o· gramsper second) Ihu that oflo.\\'~ll1.n~os~ of could ,drng the iOIlm.pher~ aroundwhieh then i:rust the 'I.~.'irnd..,are b:iowing rik]h \'I': to the intedt:'Ctrmi)'l:JL!rnK:ally sri I1i lip tiM: mugrmmsphen:i_ riorufthe' plunet, Lht'!}1 nlllL:.;! sit t\.'!,ri(knlmg their ~ht: go':';&<; rerml~ 1iI:1 m neutml aHIln~ Of maleIiIg M~1!;:h:l.HtJI.X: 50 ahan [ur Jupiter. II~c unmgllc-- lhtil\l!lS fmd ",:nll, to be 1.00IJ th.e true mmtioll eulcs (5, 6)" Cassini ebservariens show tho. w~,ph~rcof SI:n:tll[il115SI~.''Ollgly (klffillun:lrt!Jd by '~ht: l::n(:elmlu~ plas.nl<lltli'WIS :.!o\.vlyspllCads. alII neutral ;}lOm~ and mcleeules, The IiIUfllberto filii Smurlil'sma.g;lle'.ospllacl'l!. !te1l~rienrei i G uraett If! Qf.,~rg~l~ th~:r lhe~~H~~!; th, n del1.~it}' nnio OI:'ll~lUt'nd!> WI .~ml~ s BU In. ~~~ 1. ,~,A. GIlr~m tli:J1" SdllKf H~ ~il2 ,(~n. i::I!I(1ehu1M torus cmnp;:u¢d~,r,i;~h ~:.20 ~n ~hc ~if'.at.l$.:rmrls.pll'l.l'lTIa radi81~}' m:n\~'3rd'll could he ,t T"(i (1l1\'Ii~'.MIMI IIlw 1/, Aslljj\!]\. s~. 'i4. 3'9 ~l!ln}. stwoLlg.tr (lUi one side ofS;uul"n ~han lhe other I~ lonas. Thi5 dilTerellce has Ir~~H~Y heeD1 it. \\I,!j, Ktirlli. l'i.li=OOt'iJ;(,I. F. ~~~Ii1~rnfl, B, Giilimt, J" exp1i'1.11led hl [ern15 ~lr~henmC'h S:1:mv:er plasma 11),A. CW rr.~t~ ~ff, j\!f1. W~ ~;~, lO'Z2:IlIl, {see ~he f:igmel. Tille'), SlIgge:s!, ~ha~ ~hls,drcul;)tMC2I!)1~006(j~O~!!H6, t200~). :o;r~edJ «t~d ~[;ln.;;c ~tm-e~'e'J.1i;r~.y of rj?eK~. ~'tnL1o;~~m ;",IM)IJ~'{ldlli!ce;.; hi.\lhcr pl,L"im" deB· p il. C c ftJRIH!Ul1 .. Sorirnrtj;n. 'B1I3 (i!JJOM. i'~~i:;in .. Ilmvi.Jl!; p.U$t r::n.c~buJu)i ~3t ilil urbii.al ) $:i ti..::>in tint: ~~I'I of"siruUl!FU' uLlUl",w; GLI i:1J.cU §. ~\ ill DiJd!J'hiBri:r ~ Ilr" !i~llr£ ~::U.~.40i> {2:00M. "'f ,d.lllr()PQ~e tlmtnla~m<lil'F~dttcliflD1 StirC's.'ieS d~st;lI1ce01'-4 sangmianr:KIJi} o.lmJ?ilredwh~ ,6. J,!<t. W~ i~.~. tr ~I,.Sd~r:f :in. 141~(J:OOrl~. d:t~ pl,,~n~a nm.\f p,hil I{],,~m ~6 j(mi<l11Jn1dii){ 9), ·till~d~lltldytmmicl"oHpti:rlH bt't\lre~V1 thL'l,nag. J_ A.l- !llSsler, R, It. 5~nd~, '5, 1(, Atr~1li, .f'J'ar.ret SpfN:'~$(1; 29'. n:!i,h~n~ Could ~ml"II\~ri;'lti(lJ!:ii~l hi.w.·~m~Tgy i elec!Jfm ~~t'''$ll~~re;1m!! p!<I~I~ C'~\I!'iil1~th~p.;l!u~m ~he

cr.

or

<)

r\'\,ulk.!.:rl:;lrun~i,;lr

011i\'W.1rd :~lGWto

S:1.OiiVIy'sl ~p

I'mplIi~atiwl ~n dlll'~[lt:d1tdU:i ~(if~s ~'e,tq.ll~lilg

,~, 1\. :5!~IL 1'.liIItiami!'J\e', F.III~ Qi!~L fwimiS 1841',lZ~ (~l)~&~.

ill pl:li:l.'Sere~<ltive1.0 S:llur:ns

inlel'na Ill:Ota~iolli. GiV~i1 lliJ~ ~l:}'n~rl'lic ,C'hO!:(:i~e;;.sh,ll (Jct~ur in t c:ut!l'm.1 ~ p:ia51l119 !iIt.rLlt:l~tf.\lo!,~aooun~SmLlIill. ~1'iP~ lIB'bat~cHl lar S alJ,l m s ~ipiPSlt"

tl~e dl':mmlic !!:h;mg,es in rh~sm.[l d,ensity

()b~nr~cl

by

CiU!TieU ,-/

at"'~I r ~(l, hlrg!1l·!;'I;;al~

\1, IP.ffilelilmer'~ Ifi. 8a!Jell>ll 'IJ; 000.)', ~Ii~

fits"

.. ~D1dyerl'mij~~p~n mte S~ m~fmr ~nol'i! I)i;msibk ~ th~:n penurh:'iti{l!lS i[JJ the ru!I~~~si i.l:lleri(ll' of 'II~

the

~1I1 tl;r;:~t:rl1a.~

~ ~,~ l~j,~illL!'Ll?t'$S;~I:IIJI ill,; gc:~~rs "m EnlO.ldmlus ~ aDiemoreac:d\'cllo~r Ih.m in ~ll;!: 191OOiS. nr pelT·

cum r.!~lionj).all\;I.'tl~ i~~ n'iJl.1.wl!;.'l:~hl.'rc 1'I1i11)' thc i'lnL bl:l r'i.... "Ct'~"",;m'y.jmt mlinl)irmiIJJ~fm~C!3'iiS~nll.~~1t: c ~ectr.iC<11 IJl'ri!Iil~S: I hat f)(lW ,1~ol'l2-lhe 1113~nl;."i~ ~ fieUd be~we(ll1l~e ,t.lqll'I~(l!ri..], 1'I~.L..,n;1di,..k ~IilRi dt\:~ l'hul~L~ itm~lsphc:['C' .•bringimg small Ilu~s ufi)[mi7.ing hi1!h-l"'lu~.... e'1iC'{.;trolil:;I{l the ~t'I'Fll$" .4Y

10,100~i!ZO(l,;!C~Oii!9H; Wl'll~. W. !~,], :S1lt\'~IIDI'I, :S'~1f!W4I1!16, ~~!l :98J1l. U n_ ,I)-I, S;lIIJIt~l!IiI.i "'- ~ Da~~ i\l,r;;_ KiIo04OQ!l, ~er

n.

jft!mll!H ~Hh~AlJ1i!nWi ~~ra1 Union rnl!O!LIr~. 1 1 W:15 De.:emlitl'.loIlI1I6. .5,:in rr.r~cls:~ ab:!IrMI1;>521i-1J8.

A. ~'Ill!!~-ll!i\!!ig~S~t!3iQ7,

m~ ~~1l61',

PERSPECTIVES
ECOLOGY !)QmeS'l;lt~lii(l1l boilShad ~igher 5!i1tteSSr,l~es in, !TIelie,; relatjclji(! 'to lTi:o:s~~,e~nin Uti"! long

i1i~~o:ryClll,and sllede~' d'ornl!ni~ali@n.lihe'

'of aqUilI(ultu r~ hOlI~ ~()b~l o nseq 1!I~!1!:e~. 9 c

ris;e'

A
ture
j::;

bmJ1t"

.rmo yeiiffi age. ~lUlmms j)CgI:1U1 doto

Ihr

r~llUue

domestiearion,
~ he gI'C;t'Itcr

Sccoml

~ _me~li(',<ue p~OlIi1I;S and 3Dlilm,lis, (/--,J]<. In t~1e ne~1 9(JlJU }'~ .. ~"S.; hlll1d!\lcl~o:~ I~nd~11~i~l'i' ami. a few OqU.lL i~ spec ies\'>'C'Ui!;

~"l:.".~tw spL'1l.;:i~sha'l't~OOm J~lilxl

dn!flle~lk:OIted. ]11 o;;onll1H,t. the

dnln~.tica!~,L Si 1fI~"e ,hem. howJ!lJfil,cMic,llion of 114Lial~(;rrn:u.u(I:rni,' IUi!:rn rle-S"~ies has

grtJ~VItI ,~"poll(i:lilti~.lly. A~llm!';lI~'-

dh'Cl':cily or~if~ Ibrnls ill the ocean compared ~D tho:;;!;: ora lilrad Illl"ovid!.'l:.i [~ breeder runge ofoppom~lIlilies fn ~ domes'!icat ion. ".lI::I~nilik-d!i.~lt'l'lO!~li doin ~~n~~d(;',i gl,u"Qd .1U"ij ~.!n.,~ ~LJ] speeleshas heel] auributed ~olhe p,)U<c'it)'nf suitI:lbJ~ ~p~i~s: Malbj'

b~d

~'vd'L1i,io~l ~n~lgricu:lirur~ of global imper~I!><'i,

elne~liIg

'pbnl;!!
tnl':it:to

~f\~

poisenous

or

.~WllliJ1i1li,.

~uidlm1:llY

lam-e 10 hUl]mraki ncil, les the breedi ~1g,. ..~!'C. ,Ind letldi U1l;:: e o f erga 11 is: m~ are coat I\;1Ucd
by
hUrl.'il:lm;

DfllTiA;'c'1'licmiolii'ol\.vild s~'M produce food l!n'l!~I1~ i~lill

hmcllllliillil'lil spec tes h,l\'e slow ~j:'owlh :lilld:llcng life ~j't:.~~~. ~p!!ci!li l:il~J

diets, or ,~dven;ebehavThird, ,,~idl;d hy ~hi;'! ~Lpid !>1'I'I!!ui Clnl:~lmk':l11 and !;Cienlil1J; I\nm,'!'ledge. dOnlc,'ilic:u ioi'l or:,tl] lKLDk~

iomi units ~.n.

,!4t .By 2000 }"l'<I~

an cstimilwt!IJt},,!t) of lh;: species presernly ~l!~ti\,~L~l;ld on

IIg0,

II[lm~hlld
dl!1'

btllJl~

tiOrlJ1i(JIStiell.l\."I:i"

Sinr;:\f;l thJ;i!llillus~

.. l rev{ll,~tion. increase in dlC mnmb;:rs [If

O~-~~~~~~~Oo-o.I!!!!~"""'~=~~1
1000

=~~~""""'j~~~~~
100 HI 1

_';lJl~ciL,l~i,~ dL,l~~~tlfliP11'lglo-

deraesticated ~~md 111;ll"lt aUld ;J1:lirn:d ~lJci(!'l'> Il;l~ b!.:lIJn. modc:'il,(""3!JotJ) ,[ ~c~h~figu~),

TilTle ~Elrs D@lllIIe prl!s@nO'

Land 1jO@r!lll!l water: lotos'! IlilildISp!1(iC'5 ~l'il?

ln 'COi:ll.msl~ the ri.se (1 raq uacuhure is .~ c01uclllpornl'Y phe-

dGlIl1l~stiC"alNll?a~liE!t

than ~1[W~tic 5p«ics,

but in

'the past lO() ~e~r~.man~ more aq~.1!icsp!!(ies Ih,m i.iln:clI s~ecies ilaw been do me:stiMed (6).

billll)!;.~,,111~rul!i, !ipL:,,~jCS I:aml w!:!'re clQl]:le:\ti(;'a~ed in just ~t few r~gi(l~lS U- 3l A )1 ~milHr gll.utl\i! efl'On ~:o ~ donm,~!;i;ticaDC m.lllll.i'lionilil. land
;ul!im<li species

~spreelud-

aqJmLt~.c: :.pt'rlies am'!.OJDOOloA, nf'knowu land aaimal spec ies hElvc bc~undernesticmed (7J. cmnp~r:cd Spcl:k.ipre:sc:IJH~Y ~n euhure (sec the liilbl~) hav,e been d{m~~stk:1L~L;Ll since the: Slur[ tlflh'l: WhJl (U 7% of know iii n1{~ rl~ ~.l~3m d species end I[), I :l% of Ikm_Y!,~'1] m~:tiJle auimal species" 20l~ cen~my. ~H1(1<l11i cstil1i11<1:l:edW6[ilquMk i species h:.\~ been dOlllliil\ti.c.,l,ed M'C;:r dn~ pa~1 ]Efr~~.in::gt::~'1~~k:mllu;.;)\~lu~rnt prngnl1ns. for decade (see 111,1(: j~lLrc) {6). The nllmh,;;r f f'lll31'IY tlqt:I;)~u~.~tlrespecles ,181sl~otJl~dl e ~~1CUib v .ulu~lic ~ped~ ,f(fmesli"',!]Jed '6 ~~ I ri~fIl~L'aP'" "\lt~d b)' thei:r huge H::pr.,odm:-t:i e ou i!Dut and i~ [dly (sec' aIle nguli"C'~(6). ~\'\I;.""iIl <I IiIm,d ns~hm the shan gcm:r.ul,OI.l thnes ooJnllarecli ~o those of n1':e~ for ~i~y Jlonle-~lic,~~~:;;are estim '~le~. d'l:ln1e;j\i~;ned ~.m:nd 'llli ~n1LI~ Til!,;:!\: l:]l\; I!Kw~i.l11 fl.1liS'QJ1S for l.h.1 '~O:IlI.f1:~st S ~UILlllilc J.u:ll1.1.::$liCf1:ti,Oll i rates ~I.i'i:- H10 I.im.t::s as fast >IS therares of domes !iC"a~ of p 1<1111 between the saceess ~I'l~hedemesricai iOIl of ion ~I!lml ~(1 I~.,urine 1ln~r~1~LI a species, First .. Iand and [~nil~ml ~Pt:CUI?i." n hmd o\;(¢:r [~.ep~rjodl o \\'~Cnd9ml}..;,~i~"l1,tiqn s r..stest ~~ the t<ll~I~' ,r(lrne\'1~i~!~iL1i~l drawn ·~;!!~b! from ruam\\·,;!.. ~<J5.

nomenon

(5," A c.tmt 4]0 (9~/1li} of ln~

or

etB by 1.1]~ p:uL1:il"}, of ci1nd~dll~1;'; species !Im~ ~Ih~ lack ofll~ill'kellklmmd., Th e dli1t.lU;S~·t~fllj!),il f .ml aqumic :;;~)(:,e~ (I es typk~lIy in'l'oh'e!> about a decade ol'sciemil'k ~~-ilfCh (13 I. Current s{ul;'Ces..,;;in the dmn~S"t i.earlen of 3qliiU~~ species resullsl~om the .:iDlL.ill·

~:emllry rusellf~n(lwJ.edgeon the 'basic bkllo~:y ofaql1ll1:t it: spedes and the: lessons learned ~'mnl P;!SI _\;~IC,;;e_<;:\, r;.i l~'re_lhl:dOIl!1~ti~u~ioI1 .~~clI nf ~L({JmMt: species abu in~~)~Yc8,I~" ..~[risk!> lhiln thnt l'<lndJanillHds. \\:Il'ichwri.k "' Ilar~e wll ill
~~1111al'lli\!~s thm~l~l '~~ise:>

or

lttlr!c~l;;r!b"!l;l rrmrl

16~.D~spik a vtl~,d)r ttl(llH'i!f 1liI:i ~~or)r..the

lio;;31 ion

Jtlr.tiC5~

I1liml)OtlJlldhi!rds. \vit~ 'ri::\illilm~'rr~!.:bL'H~~ (;!i,H'C~

~erds ') j;~mli!\l!naf! p;!~o~n.~ (If ~omp;I~Tlb~~ ,dlrukiX:c h~H,~ ~tl.:m!.."~cl fruml :l(f11iclll.ill~.

li,l'lli1'lUll:i~1 marine ~pe,r; i~~ n. p!u:ltil;:ull;).riy t uf t ler ~mi11m als: O.. il:l'!tn O ~iIl0"": III hmL:!planl

ofl and sp.:c ies has been less SlL'r.:",,"'eSSF

usbees and s'Il3ilsH:Jmneslicmed. In COl'ltf'Jst

di\'~n;e
~lJ~Ii:S. C-I"lISIll1l:~l!n s, Vcliit:b.li'::ll!!,".

~I

'fIte' ;:;t:lglll'lti.OIll iIII ille \';'{)rld's lislaerles

(l~.'t:lre~.p:t{]'UI~li{1f.1 tlr2{lCtl

<lDld

or

c.. M.

Innil!.!to

uib,e;M. Hc~m~r i5 i(1 Ill'!: 1~~tjtli!l.';:of !J.·io~ggy. Y!li:v(:~~it)' OiToUil1ern [k;mma~~, 5230 OdL'fl5le !)en m'l!t:.

~porkl,

[)1Jl~r~i!'illmd ~, lildrlM<lfl! ill IMEDI1I!. (C!jIil-U I B). ffledilerqn~ d'~ E!ilu!\lio5 AvilllJ~"dJllil=,. (l:719@
~~I!KiI., SGl<lj!(J,·

E·m,ilii:cMmwa:rle@lmedfil.

m,

l !.l~hil1.~li:l!!r~ni>, ;',;J)~ci~l>!i )ha\l~ prov~.dt:d illId ilitll1a~ impt!l.u:> jc-1I1ri~h. and W(l:i:1'ii1:;-ha\.'E:: bl'!.."il. JI:li'iIiH=,,~ti~ [u d.tlm~'di(,~I~~I'nHlflnt!: t\p~ues, jLiSt.[~i>,o\'d,~· em..::,] (9. 10). ~Iany l~l'l'I'ewHd.lIlt!lnlle ::pecif'~ 1ll:1~()iMHiol1 hmd animll!lls I'r~wi.d~d the ~11~penf "ire !,I!'ied ,I~ f~Hl'cl [,jilmre th.,Ul m;lI!:i~ii;;! t\1~ fi:llr ~ht!L'll.rh' dr.fJn!i,';Slic..u:i~Il1pf 1:;,l!l1d ' l'iJ11;'C ii:~ t>~(;c'i!!'&e~;n.np::Lll:tIluf,wcr than 200 IrililJ (1). Atlu:n:u~ilurt: ]lrm!lll~l~.tmhal>m'iC'1lI !\>ru~l.i.lrn~ )1~1eCge~[9. n. 12)]. prt'lvid~l'Ig I1lrlh'~rsC'op~ 1 utmte:ilor"7 wS% rer)'~'ll't5i c.rlmpefl~ming

arr,IY Orli)lr~rjll1l(;! l!~NiI-im;~uJil1l:!il mJloi-

30%

'OI·It!:.:l!.rillt::

li-ll1

:moo

20 APRil 2007

VOL 316

SC~ ENCE, www.5ciencemas,;org

PERSPECTIVES
for the ~"'I~gmniol'l risheries. me! i~ ~ike:lytl;l b~l:l;}lnl'! Ilk Im"il1l SOUli::c uhl'imrilllJ fom.l for h~!m~lfIss d~m"llcls (;"mll~lI1t~i; ~ ll'll;lIUW The bJm1~ r.lIi1g~ uf dOlltcSli,cmt:d marine

ur

flfumb~r ,c1 '5p~!;,~~9fIJllP

~JJ~d~s
tl'MlI!~Uf1!ti:!d

Time sin!:e dume5ticalilln '!yean b~'fore preseliitl

!!Q% ~1!t~1t!: ~omi:5tlc,nltill'

species helps
~It:n)~!:;

10 integrase t~e pmduetion trupbic l'lO clJ;.t~.lti.rlOby .n1l.:2L1I:irnu1':i lUg v

,d~m~~i~t>N
~Oli!(l,

~"Il~r~

!S\]m:iri!l:grllwlll in II'!~ "!"lIlll!r or di!!~i!~$!1 !~<¥~~ ~fX!lCl~

(o~

IN!'arl~

c lltput wh ile deere ~s,illJgenvi ranmenJal~,~npuct UJ). [t ::JJso Increases 1.1t'!.: [,!1Il!,lc of marine h<lbitn~s·where ~lqll!:.!cuIIHu"e c.m he cnndm: I ed, '-1L'I'll diiv,t"I1\D fies ~~](J(I~HUt:'C:.S ani! , d g,efl.~.ra~e:s wide mnge of masker prices for fl

l!!-~II!'~fl1l5;

~~f'lil~nim~.t!!,
~~i!5htio\al~r iIIlimalli

25.0 44
l;IMI

41)Q~
SOC!];

OlO\!60DM
O,D14

U6
>li

~2

M!J1ll"'B ,tu\l'fIliI!J
Milliim! ~lani1l

2~
19

19
3i!

II!

..::!UI

OG.l U Ila~ 3J ~2~iOO l.1i UOO

aquae l!:hm~ rUtlducts. ~'I evC'f.;aquaculnne 01i~ development ,.'1 has Ile~ath'e consequences so for dn:: !.'l~l\ iil'Ul1lm'Cl~~ ~ndl biodli'l,le:[l"!'ty. ~l1l~hujin.gdC'lCl:k~171J.i() n c f coa:i~a I ~~():;y5~,ems by <I(IUm:lQ ~un:e efll U'C:IllS ,1I!rJ'~ hw.p1J Cl~(J1;l wi ld ,i;pedt"~ w'!..'ld .;Is((':(;'dl Til .. continuit y of

IRattei of 'Clliliil!lii~e. r1l1mlirm or prMe~1t~~ TIle iI'lIome~ticate~ spedes, I,he tilllf~ by l"jh~ch 50% or ~OlDtgwe~ dQm~sJlij~il'tNi, allltllih~ 1'iI1.~s f dQ:rtnBS.ti~~()f! rllmm widely bi!tll'{i:!f11lJ land 1IIIIdi'!.q'u'i!ltt. sp~i~s;,. ,linI!lll) t~~l ull~ Q . urn fl!, Hl~5P&ili( gr!il'i'rtnin Ih if! nlilrnMr 01 dalTie5~i("led ~pE!d~_s,iseHMdi3~ ~h~ per[~nt im:r'BiIS~ d in~h~ numbCf of s;peri>il'S dcnn~ti'(Jl'Cl!i p'£'r ye~1F ~\f~nhe PQliociiolll!s!llst (j,omes!n(iltiOIll r.lles (6),; ll'L~ number of Sp!l!(i!l'5 , foil" w'l1icl1·ifomMIJ.:ali on dates I'jerea'\!'~lili'!ble is gi'o'eJ1In pmreI'lU~e'ses,

u.n

.ilY

tlJ,TIployI1Il1(;[Jt

und HVl:litu:mJ p2mlir,Il~'uf

[e.
U.

P~~e'.lltra~~-" Ur~f~Hm::~li~'I~i(m the ~..paeaud 1I)" 0 f i1{llI<OCU hu 1ft: to m{:c~ til.l;; 1'1:.1111:; I;;il'11U'l.ds J I'm se;!lo(lgl ef .. gr{I<\'I.'i~ghuman 11C)IIul:w1o'f! :fCqui~ th~I~, by m~_~im~Liillg ~.nlivinHIIiI.le:Ul~ld

th{l!'i~ in\'tll~'~d i~.lthis iud~I~I:r}'-The g~.'(lwth ~!11 the lkl(ffit;;:;tli{;iiliun ij)1·m~lf'ifl.cbiodh:cr:;.i,l.y Lhl~::\ renreserus tI tlIPH.hnn~nulil change in rbe "'''';30'
I.l~tlfllrul~rek~~~ dl~ ~o
O~'$~I:lS,

Ois::stm.llil#ioo.fJj GI.:!b".r id ~.o:J1 m f#l1tIJ' 5!~;!i~ri~. fI Pf~r~w:mir~"~IJ1N foo" ~jjd AgriCIU'tl!r~

P.I. IN. mi~. liydrY!.rJWtfJ9ia <11);2,I1S U\l9~}. ,FJl.'O,~At4~iflJl!r,!#f tlml ~~~ ~!]~ A!JIiitq~1nr~
Qrg~n!!,]IiQlj\
~.QI'fijj).

(!1~!'a'Ili:iil,

imiTll!, lOO~~· Si!i!' WiNlj'!o.!II!ifIiId~alAJ.Mp.

imp.~ci.

,I

sm;;tai'll<lble

11.1 Mll!l

be achieved,

shoul.d ~iUll at clused proolrl". t i011 cycles. ~\11eU'e Ie ed is, pr-('l ueed ~n the d Il!:rm o:!ti~~~[le 1~lrg~l iO:rg!lJiJi~IliI'iS ~""prnd~c,;1l ZLlild
Dt~OlL!',5iiciit~tH~

~rQW frmn, cg~

1.(1 aduh ~11 culture,

n:dtl.~:inl!,
",0111-

4., E, lit IIlli~ iii MHJ~..kmf tJ/ ~1ltB.1i~M'~, liM~, EdL~BaiLl~l1!', Timdla'!L ,~nd !:;<ISS~, Ll!Ildw,

R.clefi!l~iO!1!~,~ nd IN'cllN: :1. lil Ntwmite.. 5,~im.re~~7~. 309 (l'l''Olt}. 2. "', M. ~r1rri~1\ rtrJ!!f. PIIw, G~gf,. 2::1, 37 (l"9~]o. 3. I, l)i~j'fi!j)M, II'NtrI't'llB, TilO WII)]).
IE.

llel:lutt, .!!p_~ 1,t 'R.:5, V'_ Fc',idli~,Tn BiOOiv;e:tsny. S:d~lI'('~QM~ropmMI.;

70!i'!l!rd '" ~ PM'fJrnMp,

2_P,; ~IH""¥I'.f~!l\!~;JLfllQ,QI-gi
F. VI Cm'll1l..l. 'II!!iiint:l~. El!!!>.

{tC~!'I1Il!iII~eU\'!ino.l'\r5'!' Carflobr1d!l~, 'lm, '.I.0j!~6~,

119&n

s, E.

n.

~fe~!S\lr>:s tl!1l"Vqld Mo~k,j.

11,.,::,iI~CIPt;."aSC in aqlJ<l!CuhllI'C 'Ims g;iobil:1

"~~ll'I;'I1;l"t~C!o~h~i~'., Itl()L!i!Si~lt:t!' ~UliJ i.n l!."fn~!; {)!f 1..'fI\.' ilrom1.~;.:n!a il1'I.pli.e:MitUl s, The developi

5. ~;~O, :s-ro.111 ~/1f.n,.fdAlj.ut:lwi/!m!': z006 • .FALlf\iwne"> !"oWm.irni J!'!lp~L l'IlO. 5,00. (U.M ~[tool ,amlligriUJL1IJ1I!! Of!ll<!i!1k~Il~1l, .m~, ~06);'5eo~_'raur~ ~

PJl,~~~

fiid~ .. uL1.oIll.'j)..

ment of:tc!~"C'WlII;ure is bound to replace fi~· cries ~I.S;J:l1imill hLilitmndJry WphlC(II:! 6!lLImil'lg ill'! ~ laud. This ~hO'uM bt::~p're;i~;I~ pressure .un
illCIT'fL'>hllgi,y

~" HliIj_ftii~1s ~nd mellwi!1i!l ilfti!<1iaIl~lJ1~ il~ ~"I1l~rtln!il mlfloJ!ri~U'I¥l :'!oi'Cf.I{e' On!ime_ 1, B. G!iljClrnlintl!li!:, M. D.J~k_trI:!, (JwjlJiriI1Jr.t-ni~ WI~ fm/l"! li~illl9 ~llI,IIlm; i~'rli~ o?']'~ (~twy j1j'OOrtd

pp.~OO~2:1., :l. (l!rrj''W~~S !ll,.,4!!l~r.ru-fe' ~I);!:, 343 ~OO!!i. u. A. Moon' d tIt~" ..t~liIarullltoi1' ~'~;1, 31>l. (2.a~~}. l!i.R.I..IlI.rylQ! tJ' 0.1., .Stli!Il~1282, S Il.~ tl ~9:aJ. 1~_ ~lJp~ 1±rr!~Il.r~~~'S~a~mlll'la.s~f ~I;I1NiOrt al E'cellM!ct', ru!!d(lt!l ~y 111of!lI!'['DP~!Ilil C'imli!'!~~O!"l, '!\\!! Mil .!r!rJdlecl tillY, O~I'I ~r'MIliI" ')!l)~~loa ~nll~l5 .t.i!d: C1jjl[jmmlS tllat !lawe ~re-<ltl;i' imllmlo'l!dllili:s ~,J. (ilJietl ~1I4f !1II'tp!~ ~.o!I!!iI!!!n9data, i!n!I~, A~i;!~i.and S, Za~1.tI'f~r 1\ii!r,¢1J1 ~w!a!l;l!:!Iioru.

"1JIl',pII11:!i'1I!J QFI~m!!

Mnell1i:l1

searec fIfC5h\\[~[~r~!.:~un:1,$ Ul>OO

0111 ~;j!Iru1:u,d ~'Ii ~nu late!1::CillllClo" It'Dgic..:~~. tk,vd()Prblrn~s., TiwSL~ch<!:t1g..::swiJ~n~oll.· I t(~ pli!'ldm"elhoo

IlJ 1(, NlOO). a, f'. S:~!!:loo:!=, ,WWlrl ~':fmlilh!~ ~Q. n,u~:m"

COtiseli'l'illioo, Ci!m~rrii!l~e,

Mi>iffi<ilS ;llid W'lI1od~ T.ab'iI!!S,ll

9_ E~_O.Fj,sIj!ll!tl!~ fAO tFiYr<~ry lrl/(Jrnttll~

!l~t.:l tll':u]l )1J:jtJ'ojtJa .Ufl.il·(FAQ'oJ'ltJI', 00!l'_((Jrlu~ifm, AiJoi~ 2

l1lerer,ell!t~ ond

.Agi I1g ofma til? Qrg)[!l1hms and

Clr tneJlr Sll'l!l'rm roou,ce~r~rtilny ,mind embryo viaiJillnly.learli.n9 to evo'lLltio:n 1l!ry (o,rnfli ct b~lwee n tile ~ext'll"

__ ~~et;~~m,;LH~d thM~eni pL'{lbl~II!''i e!:lpehI), ,ficm,~d' by o~l~·dn of l:mJplt=s ~;af1l b1.: ~Ltl.lib~ i.r,d

many h~!m:;Lnsoch:nie8, men lire 'becoUl}· n .iJng i1:l11~el's inC'~eas.ir:lg~y I'm~'in I~ie. <llltd h;

nil, the evo IUlqt(l~mry c:onUi ct bet,,,",'een 5exu;1I

l1<LrLl'iel"S,knov.'fl asscx.l1<11 cOlil11 iet. M 'Itr;l Ii;~-l~Ii~iu!ileH"i.;:iem.::y j s e~lle~i,iU}'
~imp'U(l"IHJ1jt.1111. :;,pci;;~i,,;:>who;;.ru fem~~'~$ matI!.: wil;hlllmlli~l,le !Hales OlInd~he ej:['l,Cti~<1tes of din~reru nml~~ '~mt!J)etoL:llt)r ~~rti'liz1it.i:CIIjJT ~

stud)' dem{)m,u.m::~ t~.1!.\ ihefenl I i7.i ng "lbi:1ily of all eja"uhHe peaks m an imenncdime matl,1ue spenn. :iml:ic<lting Ihat
I:l~ale:::~

I'll,de ;~.~l:, ost }i{~un!l !W,ll~s ''''(']IpLLI''~L;\'d I~l M wi tt-=rnll ~c".. bHt tJ iii ~ 3,r,% orl:ll~l'iil. had d~Y~:hJopt.:d
I]iIQ)lt}'Qmlg

Uled to male fm;vn:'S.inclmljl1g<l<g;lng

AgiJ:!.g

lll,

G:'LIlillI!l!;'C'r m~Je: rep:rod~lcli"1lli~K'Pr.;:~S ;

thrH~~gh tlll.'(l fl'ro~ e~s-e:;;. r:.. d;j\.~i~hi~t i!Wf d rfOC\~:S;Sl ~;,'II!ed ~"~IITn !;Om:run it iWFI, One mt't~il[ :;tuJy oftl,k hiot!~bOtl:d.~~v.t"nlles~f'.~ ~ CVtll uti O1i1~Lry imp ~iC~ll~O[)'IlS: t h.'L: 11gc' t:I f dl(: manl!alf'.~1t,,:") re,,'eals Dh,lI m.lle aging lIIl<ly I,mlle al1ld .'he <!:Beof his sperm. Reco::nt enlplr~ b~ Il!!l impon:,;;u1It J~ler!.nun~:l1l ut' thA;:o~t~mti,e ic,l~!'itud~~~dc:mnlll:"ltatt' (h~t ht1th m,~Lle ~lgU.L1~ uf :lpe:I:ln ~o'lnp.etilioll. 1.1~~~tUll)l i.l11:niyzed ~'l:ll~ ~pcrl1m agill!l, Iml~ly hll,\i\'! .m tI r"mU'u li.e IiU'I1p'It:I, iULI \.~h~1'I (lJl fil1:nililjl. ~mb.~ vi;,;biti't1/. ao\d ulil.il'b'I.SJldy Ictt i~ iml S~lil:C~g; iI'i lWtl ~c~n~!Jti,[}_~:
The <llIIhl1lf5i1!'e,i!t 111e I[dW'ilrll G~y l~~iliUlll, (l.epa;r11fl1;l!1!t of Zc~~g)!, ljJoil,!~nit:.' ~ O~1~rd.otrorn 00;';1 Jlr5" 1Jt\, o E'lniIil: 1i1lm~a.~iu'ari@lOD,0J,i!t.~~

.;;I!hho~lgh .. u old m"lesp:rod~c~d,

:;p~'j'm" ~olt1~ Ofi.llt:itll I~Iiilcd
1.0

,,,,e:n:: itO! s~;u~aMy nllltur,L:'_ Sin~,ilarly.

rm~r~]~'e

~r,am ..ri.:r Spo.:liIH

during m3t illS u~spel.·mk

ii:OpU~

ation ~"Jvbl:t.es

iJl lheirp.ICu It'i~ (i IIlI~fJim!~di .. [e ,[~gl;'~. th~ {lIli !Jt~er hall.cl,. Illt!VCI' r:d~J'to il1!!>~~lIliI1al.':~ ;

~1:"L"rillalc. :VII]~(I~ ":'f. i I~ SP!!I'I~ 'IL'Qi1'il p.eli E.i,r::II'h. IJu:j r ,ej<l.cLl:kl~!!$o~'tr;oUlll~el~1!:ithose 0 f tlhJmtd }'l1 ~Jng i'!lal.i:s i:11 I~rt i~i7.i~£ thell.ru:~ n f ,I lin~ cj.ar:l1 ~a:t.c uf Qi] 11 Ol'~ nl:,J 10.:: \!oms i.n !>~m~:" J'eumllIt'. Tinc-,f;t: fC,~L~il~ CO:11ri.Ii'.lU c~~rIi.~r '!!,:oil:.: on thi!i :specie~. ~:hffi'l'in,g lhm <I felllmlc' .i m;:tI~ n2l:~t.ld(~~l;i~'I1,r:C' f )'lpe I'm C'CJlUllj)lJ[ on )1. The o it~ l!ile ejt!c;L1~atit!S orl,W(I m [I~e,~ Vi;'oi!il'C int:cm,i n<lwCI .r!l!~n.1. f~Tmll,e ~Sll~r~l 'Dt.'Ill11]'lr;!! ilion~, ~lI1dlwhL;'n SCIENCE VOl.316 2.0APRIL.2007

~;5dencemagl,orC!

PERSPECTIVES
ler!ihty

eests \'\1"hen ::-~e' Imne)l writh either a

YOUr'll.!:!, or ,1~n uld 11'1,~~~ (3 )',

ca~cs

This recem hidelJ'\;.\L;l~~~ !'i~:udy ulsn :iml:ithat ~hc pm'Pcrl.~II.y of a Icmale'ttl III m!!

with a pm ...ic~u ~;IrJill,)~e wuU depend nn h is ;!gco.

The !Igl).tcl~pmldJtrn .ri~k of <lspIOIJrnic: copula-

liolll ru1J asscelatcd ~ermity costs thl.IS F:l\'or ft>i.u:n.kpre rl: r'Cf'l1IL: Co'! parmers of imermcdi:.I~C <lg.ete ensuee rerljl iz:.!~ ion. Simi k!l:,~Y,we \\,tmJd t:xp€rl,.J 1(;;ItHI~t:Iv be nmn.re~:ikdy to re~lillN,lea~'eucopula,t ion with dthe:r a }'Olmg nr an old male, Whenms the ~jacut;'l~'€'S or un.term~.'d i!::JIle---;age' 111;[1leshnl ve in sperm comt retit ion. u't Wl.UU kii be p:1I11 ie II iari:; ildV1mUlg,COllS, lor yO!lJlng a Ila 'QIJ l11e les .. wlto produce I:e~:!! eornperitlve eiacu lares, W defend I};M:Wnit)' by pl;"t:'\'~~liiling I~nml~~ fIDlim,,ru~ltlming wi lh !nh,er n.l.,.I~~, litl"'!; imp t>::-;j Ilg thti H~y eusls orl l'cm,II~~, A '"h.{:!~P \.v~I,yt u if'IIilI bi I ~
lenmle prQpens~I:Y to re-mate I'm!!}' be [prechl;~IJy to P{:lti{jotlll l1splOrlwic Ctlrll~mit1n~ (4),

whkh could esplain why ~pC'r,ln~~lcl)leti:d yOMg ami uldl nHl~,~s s,~~11,t\opubul.': wi~h Ie mules, There fore. t he 'e:i~ec~.of .1.113te <Jg~Ilg
un

1~lrtihtuUi~ ~n~.h:::i'~llt:y ~Hay liL::' !il:rUliC;l~ rll.C10:r lInd~rrlyj nit II~m~I~'!O: psrmcr choke ilnd

Ag~ IJlaiite:rs" Spi!'rm 11%1)' 5Un~r Inffi uCEd !~rmlizi ng el~kien;[y o'Il;ul high m uUlI'i{jDl~~ to~!:I m, a millE! <I g·es, In <!dgi~on,im!lMdual s~Jm c;ell~ wh~n slor·el!lfor prol~n:'9oo peliOll!ls olliMII~ il!'! Uw male before eja!!:ulalion OLg~ and 'i'lh~n slOr,M in, the r~male a fte-r inSfnli nal.i()I1.Thi~ mlil.y IreGL!lOf f~liIilili1gl efrrriMr.y ililld of[5!,p,rll1gfilne'$'$. 1'!e\Il'f1t ~hldi~s (6, 1) in ~~uiW<lkes,sI1OM1I~ r~~fl<lllh.n .. hilllf1~~ mOly b~ !lel~ted tgbia~ r~rtili~tIOlli In f>l'i'Of of ( mGfe rMent IllcSerninalions, boy !her patln~l 1.0 oodu~ ~ts associated wi1lhre~IiI.iz.a~~cn by old! S!Jler~n,

re-msti ~,~ ~:twli€~i'es. J tim .. U ,.tety. llmt~ <lgin~ hell" pan:nof:O- declines i~.nel.llfl}'over ~UCllCilSi\!~ sets the scene for C\ft1hnio:n:uy een I:H~I ~weil" J~JI Slit" o\'tlltui en a~pl'o",cl~es. $Llg~e~ri,n,!l. l~n~ 'fcn:lall.;\f'i1'!1Iay .1J,e. b~11.!ii I~g f~rt i~ ion iPI iZ11l ~Th;,!tiiilgbt;l' ... ~n I.wn pm,",p~Cli'~'I;l parlnt'n'i, ~j3,eu:la~~:s1~6).CunSi~{!:IH In :Jd!t.! ~'Li!J;ll I1Ililh: IIg..: dl;;:"C[~lLl f~rti lilY fiwor of 'frr:e:>h.t.:r [0 rates, the' iml"lc~ urnmle :a~1;;' ~mete Ilu21~h with Ihis ide a r.er~ilir.,~ti(ln by sperm rr">l.~m on !.J~~rlytljacuhnu:s (lll~1 is, frunm d.clcr lipL:rm), ,dS/1:!r.,clats into liil~p'rml~lct~W C'O$LK. The DNA ~<J,vied by ~pill'.rm ct!UI~ pmduced by old !1~ a1e:ii· n;sl~lt ... in h ..tchi ng III ilum :Ind wt!duL;"l;ldun-"'~ lar lemaIL: m.:ilJI.\l'~ fr.ll:ll:y eccumulaie dc~c'lcrill~l:lO ~~fh.:lil: u:ilUti'l~ springq ual it,!!' ~ 6. 71. Si 1'I"l,~ tieas and ~Ims eanse de\'elnpmenl,~I~ defects Sics cccurin crickets lAdww d!J/,ul.~'li('n.~l' ... 11;1 1 reduced v i;lbi~~.'ty~ft~~(:lLm.lIh[yo., for in"ru.! J'ru il n~es Ii"J['t}.}wpMI() fJ'IL'immgash.'",. \,\i1!:lre' fenm ~~!1.~1ppt"'lrQo b~,~!, (ll~~C"'Dne of ~he st:ln(;c. ~lrO!1g links have ll~e:n nrv~"led besperm ca.mnp!.!~itio.l'I. by Ill'crcr'(!fltkllly u~ing tween male ~gt:' uml .frct~oc.ndC!> ofsperm with ]f,esher Inseminst ioU!s Uf. Q i, DNA rr;lgm.enl'-ltion and g:eae mut;UIClllS
"M(K~~mcd \."ilm. :!chol!1,dir1{lpim\!l! .

Togeli1er. I~,es~ recent ;;>wdi,e$· hijgMught th:lt di;~linct l!I~illl!:l. prooes:>es m.• he O~:U1iSll'll'Uldgi:..!illG,t~h,,\ ~~r-p!;iy critic,Illy dilf~rC'UO!. roles ~,Iil sperm c.:mnllctilioll m~d 1::'t,Y tm~ d~tlen.:mbal!l.egro~ml~~or evU:~lU ion <111' ~~on11it:lbL.;t'i.\,~C!il ~iJXL:l>. I.hll RcJ~lc~un in l,iJi'lv~izing ab.~lity dUI;; to !lli!.OLl~ Ol~in~~:;;e;q,)IX:~i~ldtu in:I!1.m:n.cc .~J'I:Ji~C panner cheiee and !,'l~n.l:linl: sexual con.r1ict [1\,'l"rVllming: l";dW;liI1lH in !!lIT~T[ri~l!J:\riiMliHty due 10 m;Jl~ ...1:1011J\irI1l1 .a!!jru!l~ ~ an the olJler h;.l:~'d,g~nem!'C.8 ~xl.!~1 c{lldl r(;I,..; over iic:.iiit iJ.iZ<1.tiOIL LJ nravcl ing tb I;) mcchamsms

1f.eIlUiilcii:rllth~ Ii Ihl'll:l if'l!-l1mal ~ wi III .he Sa Ini! I l:id:oru: a fcm~le shu-I:; ClYL.dali~~~. Hh: j:'i~iil" i~ Kt:'i:l,ll.'iI,~. nMtclilr~+wl'it':n lhi!:: risk j]f spc:rl'il!. [~ian ;dread)' ~oplilalhnt. mil..!d1e r~rn~l~ d.~sl~~a)o'~ CiJW11~~1 ~.s: sull'ic:icn~~)1hitlh !Uti.! :Imuin~ Oil ,U'I;)[':lgC a 0.'-1 p:rnh,1 bi lily u t ejc<: li Ilg ij_~ CO:Sllyfor the "'lOlI~.~l[l~e:; ".... ij.!ll!lpi(l.~tl feriU !.ilil.athm by old ~pt:rm wh~irCas lC1mll~!l wiH ~lm';I.iI:lb':s'l~ortly allh~r C~)PU Il1Il.iun. 1lt-L: p:lt:lbahililjl llf afemOllc ejecting ~he ,ejm:u.lnles or be 'sciecl:ed!o avoid il.

disorder lC3J~nE ro d,r.:i,s;II~illly shortened hones, l11,~rerurc" in ilt.klit~.Oll IOmCft,1 I~IJI CUSts. Jnseminaiiens by ol,~~l'IIilflle~\"~lC:1J they-do ~~uit in lef1l:il.iz'~I,ioH-llmy ;11:~oreduce 011':. sprii:lg qWlilhy. '::.re.<1tmg ~fl;ath~J;" s\1ll ~c o:f rcm.-' met: ~.um ~!l;:onll~~~I ferti UZ:.1l:ion, ('vcr ,[}mnage to s~~etiln DN f\ :Illajo' also ('I(; CU[ ~Iwn~!.\lh sper~l ~:Iili~~g..,i.~d~re'll~lf::nlly Uflll;~~>e 3g ing. III ~,t:Vt:ir;J,~:spel.:' .(:s.. I~mf.llt::> :>lul'(; ~ ~I!lenn (lver prolonged pe-rirnJs Ix:lbr~ re"'l'1ti~~ ~z\llii!lnt ~11Gn;::m;ur)g ~lu;,:nsk.o~'~crm d1iIl11i1.~ by 5\1~ft!~rlml it i(l~s a~ oll~d:Il1'icll1.'For ex,;nHplt:,.. in, kiLt.i.\\l:Ik~ (ilis.M Idd-aCl.!'/l!). ~l culo~ Illal ,guU ,(see th,e I·igt:rr\e~. m<rles race OJ :Iow ri~ku:f l'f'Ietm C![1~np!tL~li'i1n mid c{lp~.h.nt; fll:~ ljuc'l'llly '~.'i.lh lh~ iIi !!i0iC i!l~ p~lrUl(!r: Two ,"",'!;!ckz.:

en. :1!il1t!11i~.t~.c:

fh~H

underlie

rnnl!! ;Igillig aad sperm

{If

Sf!~H1l1.agill1!il ;uj[fi:;; ~,iWlher :'~JlJrt"C of sex-

ual conlli,eE. OI\r,erlcrlmliiltio:1fI 10 lhfllt gencrm~d by spt::lFnm tI'11,mlflgc due UJ ll'1ah: 1IIgill.g. Ferl i,1 Z3l ion by 01(1. sperm in Cre.1SlI!S 'the i
~pm~hllct~"'~ Sl;.Icce:s<s nf llinesperm donflr bur red~l,ces Ihcre"pmdLlcli¥c suooess of~, rcm~,lc

a~url~ not omlly p'fmr,ide'~ 11 d'''',lrelf picture ilir)\l,' rel)rodtlu,)t~ ve :;1.1'l:11,c:gk$ C ~'Olv,,,,d "lLll1l~

al~} help us to bener 11IiI~Lllill!l~ problems in ilI11m;alllsocieties.

:l..

1'1i13

rc i1~~:;';n'~1i~y

.;m:dth~ filll(;.':S8 oflu:r{jn~ril~g r-elativew fer~ l,iliZ3ticlll by young S~)!!i:,li'm.'fh,ese sm,diies j 6-9 I~ln::rern~ ~'~:vr:allh<ll ll~n nJT~lll:~l,l~w
nUlg doc.:; n~)t rcd,(Il:cilh!.:laliuIL\:
SWI o;:e~s
(If;]

fl., P.Olf!ulIIg,;. fI~ffHiJ, M,pd, S'l. 1~9 (:i!OOO>. .2. l'. M.1~m~, .it. f~l1eJriblil'i, II), 8, l!l P.1P''lIM, .W!.lI!" ~Irj(ar.
8~M~ l!'irar" lB, Z)l (lOOI'i). 3_ T. M.l~m~,. R ,~, EI!J~r, Prot:'.R.. ~c ,M;'j~

v. C!la~

R",I'ef~n;~ .,111111 Notes

r~pfiHjm~t~.vlii:

8 nt,

male ,(th"l~~~. i 11l:g u·s relm fo;,,'e~y rmn n~~le_~ sel~~ec;i to ~!1:l·ma!1,'l \\'it~ ;I~ (~'e ~,II]Ii!~r~mtl~e ~otrlr un :spemil re~er\"e:;; wit.h fn.::.,h ap'l.!r.'m. ~ltiwcv~r. willen IVti;J~t: 11I11Ltillgcosts ~",>;ceed <'l threslu~kl it is nnOl'e ad\''''!1t,~.geO!,l:'' I:hr .. m,.~1 to ij,L1.\1elt!:i.llmte new ~
Co:~i-tl;;IN;).

nu~OO\1l. .:I. 'ft, l~..tl~. C. t'... C!:ffl!J!!~IfI!I,:~" i'imlr!. Il2\OO5~.

CW'f;

i""1,:l~. 1222

I'i.. J. WjmJlleil ,~I ~/_,.ko'K, N~tI_ A(QJ:s.;i U, SA l~, '~6!)l (.2.alJl.fi.)'. 6. R. iii, waQiiII!r~F. i'I~I!Il$l)~ijj. t l~fI~mifl, Pro~'. It So,r.
,. II" 9_

s-

,Ill

10. ~:e~~~~:~~:~::~~~~~:I ~~~~~~~::~: ~i

~!!de'tl.

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20 APRil 2007

VOL 316

SC~ ENC.E, www.5ciencemas,;org

Fresh angles on the science of education

Aw,e,aUh of kjnow~edg,e
~Ine SciencE" Ed u catlen forY m is II dyn[a m lc SQmte oifi nfQmmlign and ne.w ideas Ol1elvery <l5peCl or science educ <ldon, as weH as the :sclien{€' and pel ieV of edllr<ll~ on, The ffmum l~ PILIbl [shed inlhe I<lSl issue (I Few'rv month ~1!1d (lnli ne, in ~:ollo:!bor~ti,on wiIh the HJOWtl rei HughesMedi-ca l ~n$titut.e. Keep up-La,·date wlththe latest de'l1elopmenls <ll: wl,'IMr,s,denc:emag:.org/educaHom

What's '\(Qu:!" '~~rs~!l!c[iv~?

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Warming Island"

and Ethics
113~1II AnnlLlllal l[r,ai Hller-,of-Tram me 11'5 Sli1Io:wmass, IC::olofado
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A'\t'tS1'1'fD§~mtt:hDr!~qlcdI~0C!<1I15"OC:i[Ulll.

SPIE.CIAlSECT~ON
INTRODUCTION

The
6ERMLIN~
gWUl

Iransittoninq Germ tine

ous tml.1~itiolll~ and d(;(:I:;io:l1l$ d~1Iiy ,.;Imlll.·o,gmph..:d fur lh.iO WJIIlGLC. .. • uniq 1iI(: <lIbilily in mitllLLl.'to U,.;vt11llli 1110 comp kx ::J ni rna I mu:!: p!1~:\ on g.1lII1~1.icin fiXrnOLlioJl1, TI·.i:. ::;PIi'1l:l.j] [!I, :>ct'lioll higMighis ihl.:~ransitiCilllililg g:cml line aed otherwiseexpsmds an imnglllirl~ i'~P'l:li:tsof lh".lll! eel Ls oK hcr."d il)!,. [n Ilw ~~I'ty~mbt:rl'l. t~] lis dt~ iiJe Ilt:'LWL't'.I'I IX:,",(JI11inglmlna [lIT ~11ll" lill:!!. ~M",,"l:ml[Iili1lJII'" reiect g~~rrn ,LXIL., fr1DL1li :;o3'11a1ic I~J(!. j LI~I :IS; ~olln mie c dh; ruquilfC in::;'l~~:lI.iml p u loom Ihi;;'.!,!£IlInlinm d~.lI.l.m.mli[!~.ionp!l~hw;Jy .. Sl.rmlll~a:rid Lwummn (p" 391) outline ;Jt~ii\'il;i!Cl.\in Iht Iny and W~1fln. such ,~" Ir,cll~ilcripLiOlml qll~tSCCilU:C and inhi.:I[Ularu.]~~r :!J1.."lU Ot.,:.1 I dl'tCnll1 ill:II:lI;S. mud 1:1;~ya~1J~I ai, rp. J 941 wll 'ntthc murine -lly .... cm •.ummki l ling ,1!I1li induetive signall. AIUlO~I.slllmHid :;;ysh:rm L1:lCdi ~T(:r'ml J;!tml.qj.it'li. cQ'~mll()n

cdl ill:liC1 is ruH oL" h£111@_I.:.The trek to unrl rrmn r C

(ON5,T#lN.Of

ISA MUS,] FOR 5PH:IE5, WNTIINUIT¥.

lil(:

Burr lfHE LIFE. OF mE

germ lint' inrlilulcs numcr-

Germ Cells
(0 NTlEINTS
Nlews
388 3 '!I0 Mettln.glOpposiitlofil ~ClIFrCluu'E~g$ A Clos~ Look. a[ UrtJi5~lilali~y

A 11.(.... ~ dOll:lll or 1'I1L:'!'~cru ks rq1l'l:r~.ccll,:c ~1111.i!ripZllcs in ,l!\1[Jml ,ITU dt",(;'JlL:'!pI11CI1~. U 11~p. G J (J 7, di.:.scriibccl;[h~ ~11~r:dl.i!lJll,:,!lJd ftJ,k L1-f !l~nl1l.i.nC·iilx:ci ilL: i:i1J:mn RNA~,: 1~lilw~d pi.IR.NA.~,. FUJlCtimls ~b'l d.iulcru as po~ltF.m!\Criplion~J fI="'g!JI,Himllln!.l (~p~b"l1mm~ic 1}fTIl!!,mrn,· ~11ill!,! am:! (lggfJSlOd ..f\.rl_o~herepigenetic event de n(_1!{{j, DNJ\.. mClhyblion ,'ik~IICI.!:;; illnprillh::xi 1}' rIC.... md ~JK'iJt elemem s ,cl Llring \ CTl.CiJmtJ:: !lJllm'L~III.~ k .sch:.mfi.:r rtf tit. M1)_ 3'IJ~) dl~ La il IJ, h, hmila ble ,,,i:l:i::nciIIIgmGch.mi,,.rn and its :>c\u:d djmorphi~ mi. A.~lhl: germ. re.~~l.rr!ll1<si~ ion:;;;(~, 1.11:.; nmLll1FC rDl"'m. more dCids Ions .i!.~1!mndc: 110 cllte~ rn ilD.sillor Ilil~io"is II 1,11:1 n ,di I1iCrellti~w ;b<: SPCfIJl cg1l,'Ivith Pf1ssihk lilllk:lg~ Ihe.g~ ~ of dIT~f.liom .;1[01 r:riJIP~j!OtJd by Kl m!:!~c lind! Pn.gl.!'~ p • .1I~ mho S.lem .r.:C'~11r-~.f1t'w·,,~\,ersus ,di r:o:d rc 1~1I ii~lil)ll :iHllh(!' Di'I~~opfJi'o male nutl f~ma~~~I"!il Hru~:si~ i~CU:;Ncd by Ft~~~c.r d ;~nrli :Spr~£I1i~~ ·4:t11~_Sw iU;hi Ilg I.a \'(ln~bmu~ gC1.'1'I1 in~ Rh~,!ln cd ls, Brj nsrer Ip, 40.!ll Ip_ ~ !FC ports methods rOf !J}C rooovery. cuhure, aud t"~Il;;:pl;~OU!lioll of ,'i,I)(!,~TllalOgoni~.1'mml . celIs: ,md Dalley ~p, 4(9), slIl1mmr.i.z.cs[l£h Qnc't'~ i.1! crllhrym~~cstem eel I ~ESC} di fifC'l" cntaonionl IJ[), ill. g.u Ll1I]~ im;: E!'I!'~:Im:!Ce(h,lF~!'i,wil.B! re~C:'<l a nd cl il'lk;;lt ;~~ll)!k.ujon~ Iml ' ~h but

dillb'lnltS

~1{jl:;;L

nz

R,levie'w:s
Germ
V,~rsu:sSOlll<!

[lecis,iol'ls:

n4
H7

0,

from fli:es ilnd! !likIM1I5 5: S~II'I.m'l\~and R. l,,m.moJ'!'!'J (l,erlTil (el~Sp~iffcal.icll in Mioe K.HQ]1<i:IsMIUI. lP,i:I;!NA~ the Germ Lin~ in
iLc5~1}1:1i5

H. Un

398

ua
.tIOZ

Epig~n~tk D~dS'i~n~,i!1M~mm!)liOilI'!: Germ (jeti1s CB'_ .:kJ~rJf!jl!rIH al. Tille M)!S'~~ries Ol~ 5I!!XIW.a~ IdeI'lHt~.: TlhcGe,n'I CeU':s Persp'~l:;liV@
I. Kimrlle' ond D.. c.,

(~l,:;.%jl[.~k~~ Ihe dli,'liC'lII~SIOn J'lI1111cl.·. hiflh!i~hlillg elhr~al and ~Qdal issues s;~I:rm"'l'lding ~St'·derh.'c;,d germ c;el]-~. C·h-i!!nge.~ ~1:l11IinLl~asthe oocyll;llr;~I!,~il.iml!i1 !Dth~ rcrt~liZl,'d ~g!} Sli'7cd O]!.~ld cjlS d~~LI:r;; 407 ~ d1ctfli I Ihe Ji}(}~y~~·!lib:'~'!l.~hJ~ ti~!{iI il~~!. \\1i,I'1 :l!!~.:::odl!r~'AJ I'I~. I ,H~'tt'r;l~ iorl!'~ iti~ prot],)'! 11 .~)Hltb],)'s~~. IH'011],)'i!1 a.nd RNA ~.k~l',l ",dill.! ifln. and (ll'g:~n,],)'lIc .~],)'I~lfiikll ing~ :l.nd StIllier ~P', 40>1l~ djsc!!!~'C'i! ...ygol k' lllCflO!llC ~icn~i!l~ and 1l!1Jten~1l1 nlcS!>C'ngc.r RNA I d~r.:t~bl Iron .uxlLImlllhc n~m~rm z_yg(l~ trnnsit len. I.• fr:: ln Ncw:-l.."1<1\' is (p. 39() I di!l(;l\iIS:-;eS "u~rhisc:lai!a~ it)!'· with .. billl:"gisl ]'DOll deri l'!.!ill BIle eV<I!lutino~ of germ e~U ~I)oCd neal iOIl. Illld Lt;sl ic j p. 31<1 } dCt>L!~5 I M hO'w flocy~c INef!i!1g hq~ b~C"Q'i11~ mere crr~lh'oC and "i;(;,C:fl(;Il~!c(De~phc,qr\t"';:I!flll~ ;I~fll1il~he l{:Illlv~cn1l! wiLhLl~t li~ll~':Hi'l'l:I'~. In an Ei;~[loriat McLII~CIl

Male j,lt'id f~mill~ Dro~r)-phifrJ G~rml;111l M.1. f~/ertmdA.

.;o~
.1106

Sl~m Ce·U~: TW~VeuiCl~\S of ~mmorl<llily C 5pr.niIlIl!l Mate Germlff]!!, Stem C~l5; hom Mk~~o M~n R. L. BrilMtrtt
TIle M<lterTf!O!~.z.ygctic. 1i'I'a!llsilno!'i:

Del3i!b 11m'! Sirth: of filNAs

A F. 5,hi~f 407

h~llih p·f Q.~'S~tring_nmIU:y. ill :~-kju-,,+s,STKE ..:'I!I~Cluoo disc~l::;ses h(lJ\\ l~flPp ~~-:.rejc,cl ~C] f·pLlll~l'!. and U~~IJ!~ i.~fllir anti J~;I~ &Ii:gh~ighl ~j~~~i,hie~in ./Jr!l.~I~/,Jllhj ,mu ~n(loiJ!;i:C Ll n!cchanjs!l1~ im'ohC'd il'llll~ I1:ligra~~unor pdnlO!·di ••1 g>C'nneells ~nd el~mir"Hion of
!!,!C~flfllc cC'~I;-::. The anie lies nere give nIH ;Ji ~~Iunrlnl!! of C'xdl il'lg omgniml resc,[!rch O:l~ Ihe !>1ccd:Sr i o IiiIe o1Ilen15d\,'l,,~, rl"OiJ~iCi-S of'cha nge. Ad vanoes be'il" n l~~. i!~ lu~dC'r::mnldi'bg dev e~Ofl'mem ;lind hs ~i!t(;kr!Jyil1~ gene-lies. as ~\"C~I:;I;. ... hll~Cfl~liqy.
_;!,)~!!!W!b !'1.I~~mtllili!l)'IOIi!~'~:;

Reg u lalli(!f'I ill title Ooq!'l~·~OI-.Zlfglote"fr.UlsVt; 0fI 1M. L 5UJte:l.a'fld G.Scydt),w G~Jlll1'tfSlirom Emb",onk Stem (Qlb~. II Cu p ITl\aUE flI1IjiJly or HalJ ~un? G. O. D6~ll'r

1109

,il'jJj ~

Se~ Il'lsQ .m.(JI«i €tfi~~itl~prl!M' 1.~ S~ 1iIiW~I'iQJ!WI' ~ m.lm'W..wn~WMl1.~!'lj!IKf.e~V,q~UJ.!

\I'!IIAI' ...... sden.cema 9 .orq

5C1I:EN:OEVOl316

20 APRIl2007

Germ (eUs
.... ' B:I!Uer c!!lh!~ ~rl\i:!li!1g a illHllan Il!!m l,1{ill~ou~
.,. [f<!lllilging

or~iUnn

gl~

i~ tliIiciqr.

L'O 1JIl' Il:illaWG-d y!.:ars

or

~vt\1fQdl."C3d~~ lal.fu'f. \\~llJn

r;h{;!irnb,,, <LI'C ~,;:urc [If ~hey'v-e~~r!d~h~ r.ighj~ j jpilfl.mn:r. Cryo Egg,_,,'s dOlld:!' bask opcnt:J ~h~

same yea!': Fertility dillies. now offer

,-~stm.ugl'.

Ck'rCl~'itt!

Eiul!y fnllll't .oocyte cryoprcsfT'riatima S!ide~1ep:5::1Ome'01' the pn:',L~k:~~ nrJ t:thictl~ pi r!;'1!~. .. oflVE S<~)!l'Je:fi'fL...-y Boldr. an cmbryo~.og,[<1i[ fLl.ConMHLmity I-Iealth Net\\{)at~ in ~'lldiaq~'l!illli!;,lndia!l'l. \\11·0i:N ml00;1 p.1lnner iu and seiemtflc di~'t('ll' of C'ryo ~, Far exaraple, \villilit 10 do with ".~P'!~" colbf}f(lS ~h.. t. ill"en'+t Jll1p~~nocdposes .fl di k:1Il1inlfli,. s.a:¥.:;, he Mmw couples ere loath ttl di~m thern.aad ilS
ill.~ga.~hl
~I:;~ th~UlIl

Fm ft:d~[nlly fun·cl:ed

IN EWS

~s~;II"t~h ,.M~1n;!OVer.d~ \'o~i n~ ~'!JP~~s ~ [~~',~ lixlglu Ru- c~bl!OOl)" ol'frutcil 'b'l.nbryos Llll.imtlM:. just as they would over ~hildren, 11{'I~esl innC'd~ c ~!1llb[·yologi~l. Mid,.ld Tucker {lTCii1larM.i~~

'Docyte, cryopn:!s,ervaUmrnis s,tartingl to, mat.ure :n.OiWthat r,eslean::hers: have, developed a faster, better way oF~ree:zil1l,g the vulffilerabh~(Je~[s
mH1Ht THE HOT DESiEJU su N IN PHOENIX" Adzona. )1~~rne[hin\~chillin!;: ~$ govllg fHI.
1:f~lllI!!,ln ~gg:;; ~urt:sl o\",ly beil.,!;

1h.1P;(:Qd~!~li\~e reci;dis~'5in A~hml:1;!. I;'en~i.~~y S :h~~ly ~.nd Ge rma n~ face <I d UTere nt ies,al J i!Tic LIlly: E'nlll)'(l 1:~L;;rj!]~ Ihrbicld!L'I'l. i-~ Frozen oueytes obvll:!.l~ smnt:' ofl.hcs~
~r.h;dil.~~S.ts ~OLtIHd~:;stich m; in {xmcil!.rnJl.fur

e:x,~·nnp~l:. P~OI)I.~ IiI1!11Ye I~ss b 'sqlH!'mtl i~h <lihmn io'ssin£ mil :1~,mtlC~C than
<J~ ,~r:nbF~o. :Bokh Ii"Y~;. lI.U:C 0.111 lire.

Me'l1 fou.::i!lg c'an'lI~rlrc:..~m·f:nl !hOlt mighr ren der them )ll'e'r LIe al so 1m\I~ ml adv:lLm :IS';:

hc adds, til.:! etors :s],llJuldlli ·IIJr.:Ct!

Anti if !>p'oI!'emlqll(l~"
1u

Ilcud dmVU<l1

wll1u has been calleel tt!lc\'vu:dtl,s fin>,tbuk uffcring fruzen, tlm,l::mJd c'ggs. for ablHIl

S2500. C'ryo Eggsilllerna! nrm.\'l.I, h.ecomp;my ~

wl]~~n!l- lh~ ~):ll]k. ill seU an inJ:hlil~ \nUll,Un w 31'1 ctlg fm.~none of its eareful) Y screened dnn{l]"s. One eg.g \\,O!i!'t be renoll,g'h women

lmur use with in Villl) rCI'~iHl:J:tiQ'1Jl UV,n~SL}Jm: pl~y~~ci,u)s C\~!li nhlUl ~:o,,<!o.r~ :-;p~~.·m ,,,W!1f!Icells ~oom pn;pljbc~cm lJu.ys S\+~0dH~.t:lj u.rsl ~nilar C
!}wr\\'U'm~~l" TIne}" t~1I'I. .it'l3·z'B &ptrIT:!l for '!".1!lll:!Ql't!wmpy {S~~ [3,U'l

create as m;my emh:ryo~. CU5lOd}1 b.nl~Cl'> 3.1:;0 !'lit01.11 din 'I e'UilIpl. s.ays 'Iueker: A \'!I,'OUiI1ffUl's
1;;'ggS

are' her own.

Hri:rIS['~rRc1fiew. p_ <I!04!I.

i\:S:'A'I!.;h!o.unc r:lSl:.!(!~ r, fh:ei!:ing is. I.h..: triek has been l-'C~tiVl!;!:it lQwork. The firs! reports
0:11~jrt:g;m! e ies fir'um l~.awed one yl~~ d,n ~ n b'<!ck In I: 9~. ju •..r: ~l yean; ,Ifterthr: fiJrst.hinh Irorl,~a ~rozcn cnmh.ry.(l. Alilhuug ill the use (] f

cg--gs hmJ¥r.! been d.~mcllit to f~o(!ZI!!"Ail

~e'~lO:hed new s~.~ge.And some £w ifs ,,!mul .1 li n~e. given how lInf'1LirImi ~ityis, J-\ tyllica~ Ill:lnr h'll:> ,al.rm.lst a ~ ~rn~ h~omoi:l ,i 1~lh~'[,[nll ~ ir13~
~[lo

the price dm:su'l imli:::hldoe crystals, "The pJ,1)i:tlc.mwi til eggs is that the}! i shil)pjlllg illl a I.i,~uid I1lil~elfl-U)oloo cOl~~,im::tr; nrc sl.:rL'li~i'll":to hoII'\' t~mp(;l:<uure." says Ju.~rgCIi1l whkh can run S200 lor,~ U,S. address ~md tieoonn<llm. sc:iemirfc dh'cclor oflhe' Fe:I.tilily Cellli:P.oi of 111 1i1'IOI:\,in CI,lli.mgu. i111'l]illlicl $1 000 l"or dcl,i'l cL)' ~o ELlrlJpt.>. ~, m~'Cvcr, reeem techn [(.'a~relhiemems [0 I Smile ·ethvd M~ and p by~,ici,lns dec D')' ~hc sdl.ii1l!:l or doml!.>clI ~ggs OC'CrnJiSC o'1:lI;;;i~ing. uhcn.l e.~!,lWr~e;.:i.ung" ;d~o known",) {l{)cyle; CfY'O~ I'l"tuill a wnrllm illlv(ll~\'e;·h(lnnOfLoe ~n~ti;t<lm;;"Uld p.~selfv;Jt ion" may soon even out some: {If I~.e ,I :;1,n-gil~;dIflJ"K~-dun: Ib:H l"lnLlIl,Ji...;,e ~ud£,~m,,'Cu;, reprodm~li\'~~nl;.qm~l~li~t> OCi.W~lll men <llld But (;~'\) Eggs":; business p~an is sigfliticant IIlOI '\'roIlJ1lel'l.AlthQu£i11 some scleraisis fi:inlail'l cautit;l~~~ bp\ll irs e ft"'er;;:liw~e~, arid s:lt,~~y. a (ll~~fl' b!,;t:31011::i-'!.> i~ raises IK.:W t:thi,~<llji~L.llcs--illh;r II ~~, ~itglti.: that cryoprus..:rva'L~{lll i:;. t!.!nd},ft}fwidewomen lIl;Jve been .Qiouali,ng eggs Ihr decades ~Iuj~ad.tl.~~-not just to J'udr C~!Ii~r p\al:i~IlJ~~ 11!1cl:~1;!.1 b(,;:G!Tlg~u~.~g 4iCl-bl,lt ~qlll'''~ P .il ~i!:lnah; ihi'iL!h~ p.r.~.(!tice' ofr;!g_g f~ez.il1ig .I~as ticcurelltdr fertilit)' bUi. illso to g~\ll!wom'!..'l~
r:re:M:rHll11.r:y-ant~

lypicaJJy

erderslx

[0 e~ht lOOOOOI

the eddsol'

.!1.1:: 1:I!'gesl. cell i~l tlre buman fundy, ·;H~egg brillm \.",i(h \'i'Ilh.::u- l!tW. Cilill lIimlil dflIlmuging h::c

embryos created byl VF boomed. tc;:(:bn.ic;:"l ~rl,lrd~e_.. la,we I<l!r~~~y~);d~~cloo CIf)'_ ()lPre~nDd eggs lroiml the cliuic, Depending ml whu 10 C{llillinillg.. froz~!l OIJ eytc_ a(~cmlfllt .. for a !iI)l~J0 f 30 0 to eOn babies ~i(lI'Mwide ..
[mull

C{lm.~K~r~wiUl th~ IrQi!.l~hly 300,:nOO hinh.<;

o ae reasoa i;"Q rt he s h')"" acceptance o f l"roo-en cl{lcy~!es in llile il 9~ijs am! II)yOs Is that !;:: ~he !V F techn iqII~='i ~ li1~a usedwere f.lHld~ :li! ~ IC8lOaliCCI::Sdul 1i.::rLi.lizn~lg tha\\'l.:ld e~g.~ thun ~ fresh ones, I III mhlithul to ice crystals, rreez~ ~
ing ,l1IiIeM.~ ~Cll~gh~'n~ ~h~ lIHil:! p~IiLU,;iJ,~ .• rliL':" ~.

from Il'uiOCilI ...:mbry(~$,

IJln:recontmt o~~I",""'en tln:y Nt.liFt fm:nilies. The

ltl'dllllolo!:W h;il'; :lIJ_ru'1Idy jlumjmd b~Yl:lilJd 'l~1l

mem n:r;l!l;e.. rrHH~ddl!: egg ~h.,tthe sperm h!!ls ~.'"

bl~r:t'u\.V'throll!JJ~. ~ An[1th~r ~iilotQl'{lUllgm~ lietlP!e pmSl!w",,, ~ conrem !!b[l~~porl~.. ihtL;; bi~th Egj;ls 0In: ~ al~o :ilus~l!pl~bh: t!LJ cu~!.Ib~ellmrL::I mm.UI"l" ~ tKK-"yC~i~Sl"i~ k:do.1 tlu; Itl iddltl, 01' n1iiiiiJs,is., [hl' ~
[u

\\tlJmn:n'~ n:~rrodu!i.'ti\;":~if'~iViI~ ~ii~l$ ~ya I.h!~d(! ~m

or ~('
~~MI~!lU1

~d,""Oim:ide:, with the cnli'-<lllime lor

eclucmtim~ aJ~dlei'>~,)bli$~'Din~ ~ ~ C<Lrt.'Cr,

la:>t 1 yem-s:. E;d~ltd .r~fli I~ly..biL~d ~~ \!vbkium. l\'!Ia~chu&!tIs.. hill!: 1'I1,I~keliedil:S :i.el'\'ices ,IS .11:1 opliol1l fOH<'I~f;,'e""
11IiIirlid~d \\ tlllm~·~.l\vlw
\'l'U1tl lUi ('Illl ;l"~.d~ {lOl"yCt':~

rea hn 0 heseml:h,

F!.1l· Ihe

(~r~I_'"

20 APR~L2007

VOll'lo

SiCI'ENCE

www.sdencema.g,QrQ

SPECIAIL.SECTION
double d:i\'isjufllh~[ pm\:~t" I)~n the ,~hromoSOIl'M:S yh~~datnl'!'l!(!~.ih!!'C'h.ruUill()fi.omCii are 10 ~

.A~t~U~Jl orlJYOl.'ImUl :~ ~b:i~~"'e~P1 Of)

~~d{.n~c

hitehed W rhe $nini!lk'.:lllle~!I,,,'nrk nf'micre[001:11(:". Ih~,l will J.ldp St:j.l:I!11W tllw.mJl .. iHiJnilirl":ltim! O![:~U~S. E~e"- :.! ~,,~jghir !'o:p b(;li(lw bondy d tcmpenaurc 1r.~ggel'S ~L ""dllllhlg inj my" to an "-"gg, nOli.. S!l'IilIe~1Leibo, 3 c~'Obiologis~ ~t the 'N lJnivlOfSity urNe,\'" OI'~e!IIl~ U:lui~ ...Oln!l. The in i spiadle falls ~lPllrl. s ihe ~nic.rombllles,dcpoly~ a merizc, A~I.l110~ghthe Sltm::WI'C oclhrlfls~vlhe"n temnpe 1"UlQresrise, resea Iil'ihrn hU'!;'CIn't n1k.xi 0111

lion."l~ re,'jle\l,' cryopn~s.~.f\F;H~.ml. Ali, Licburmann ~lOh~,~, lhm.fbl:~gl1l tflllbk: \voJlt.u:.irII Ie! .IiCG:3.in ~hllir l':g:gn,(:f.!!lit SWl~li(,"~ SUll~!;;)\~·~flat i~ boosts I,,~ surp["i:)d~l(;i"~ cap;H;:it:y-I~ill,g O~'!,lfi,l!l ~is~ue vi \lll~.IHU] fen ill izat ion over s:IO\l,I fr,eclJinl,!. ;J11~.n:i.rnp.~~n~lil.n!1, afocr. tffit1JmClll-.b: ~Lill. for i~ reducing Ih,e <~t\erage number noeytes research oilly. Ibe' guideIiTIes oOiuchKIe, M ore1!,g,liirlls~ Il.l!1king ~lL·ctU\il." teqUilrudl LoproLluc.f:~'~ive birilhh'WIlJii nn'l[l'retl1ll1irll ovcr. }\SRM ,1~~r\i'is",!s 50 to beeveen 27:J1n1l30, eoo freezi ng comnterei ally a"'a~lablc. ching ~he hick !(In[l'Jl!:,:,4(;r'ln .iSltlJ.dit':s on babic~,'i'nl."tuith,

born fmlll \1.~G:rifkd c'US, rome II'CSI.'.fLrohcl'Smt~ 01~ready c~lIliLlgthe I'I"Ieililll1>!j thel~!llI.re of Otl<C}'te

\.~iOmcn wilh

it an~}' fa'f rresen:in~

'l ... . ltCCI'~~,

~:l.l~r!.(slutimi board slnpcol'''I'isinn, Analt(mll;I~~ve

::I..~,

lon~

fen.int}, in

or

Reild¥ .~~ Ipri1me dme? r

HU:irlldn.'tli'> o~ ~'I.>e:I"IiI.ullgiy ~e,allhy

hl~~ing ,dl.!Lll"~~!'rmn Ih~ di~Htptumli,l'iu~h..~ ibunon113 ~ mUinbers, C~lro1J1;OSOl'l'n,CS jn ~le egg

or

b!"biL}~ "~J'urld

lJinh ~fdectfi.. r-reez~ng sari'S sperm, ~ClO.:kfliling abm~1 jO'X. of ~heln. ~I)'S .~l;:Il'J."oclm.1Q v~ end~~¥i~wlo~lsIKllIl.luk O:kI~:Yt;lfWciH M",clIk!1Jil. eoul.':g'!.: oil"
;Climclll:'oIfe

Co;rnel~ U1ll'iv;ers~IY in Ne'\'\.'Y(l:rk

'Pnl(lli,g;~oll:~ fi~ll~h~

iii!, a :ot;"'1m~1:Is:;.1fi1irl~ llliw

('~!y. hll E

the
~~

~hc werkl prevc 1~l:lmoC!C)'~e C'r_'ll{)I)OC~.rva~i.oi1n wnu;ks,i3:lll is it ~t'lfe :mdeWf'iciem enough to' become 1ft, !>131Ufl1lro, pan assisted UiCpilDdtlCtiull; Inan ~Lll~lly!\is publ~!\h~d lOAM y'l;!';!~'i!l1 .f'(~'r·IWrr rmd Sn'dilly. OkldY and ~olki:l-5.m;::; pooled resuhs 9:r,',26 studies I'lfll ~)OC'J'\ie cryo-

or

OkLJ:Y.who headed the ASRM eomrnlttee. rl.{m,'l'dlUt Ih"eZiln~an ~ e::n~l~"'¢l~yi.!~e-~I~:) or !} years" So urdcti...s II w'Onl~UI,l p~rms co \,l'lJ:il ~.oillger li1an ~hil!to try tn g.e~.p'lqUD.mt~he is better nlf
u:i'il'lg !ncr 0\.'11, fresh eggs" Oliuer f~L1~~.ity expens CQllItflLIQd tIl~ t i nl~ that is r.IJ)c :I:Or egg ·~rt~:rjllg, gllv,~n Ille kK:k or a ~I.(;rnarives, AI.~h{lllgh elllli,i:il~s railed when EXloellld Fi;'ni!ity b~1ll ~ldwnisi.liIg ded.u~·c: ~rug !ro~l;'f!g !.oyot~~~ \\;Om!:m.~he pmciic\L." .~itt~spN;at! ~'P~d ..~~y in Ihe Ullli~ecl S~3tes. where ii'ertulity diUii~s ar~ 'U1Ifi-I'!1!MI,d'Lt~d. Huw n~JiuliyfadHll~'s effer it new b,~l't cl;."n~LiUi. ,"~)!~ O.kl[.jY. l~b; ,dill,k: doesn't, ~~hqt"'''nuls:on.·:; SC dimi:c: U
dO~:-l, "I dU!iII"[ f~~1 .~C~U:l,iJ~ny

~~\5!.~reIImt e:r!ffl~gi!l l~;.>!~dW' nll~ s~r'l'i\~ dIm"I"ing i.o p:.::m.'1~t I:;'.'r.ii lizadUlJ. he: ~ty:;', A :!l:him!~ili1 ,egg 'm~ the 'CirJjl0Pf\I!SiIl:l'ViI!tiOIliI The 1It!\'ili1;Ce th!u l~vi,\'t:{1 c~.g Ih:cz:i~,g. s.i:I)ls ~p'rtlducl.i~'1i: m'k!ocrii1:[]ll.'igj~l Rie~liru .Pim~~ui1 .urlhe Unlvt:rs:iw orSo~utlleln em.li!')l1Ili,~(USO r iI'!, L,~ A!1gell;ls. W,~5 vru rm::y[oplu."m ~~ ~r~rli!l il,'JL:c~iun Ut'SU. The hic~n:lliq~w. which researcher'S finH rested on fr't"l7.en h!lHlUlIl .uO\:;"Y~es 1995. ~~l~lol:;'lne in sperm dlrol~ line
il;o:[d-ll!<wdoo~ldJ$1'!lI:.! prfJhM;; ~&diil'l>l.'lly 7 into dl': ~~. "rm surprised dH'::Y gOl. <Ii."Y leni li7...i:!1t ions at ;!I~ with~ {nil ie s,L)'S r,tfLI~1>01!LIn 19~H.

ac~CS$ICI thi~~~l.ldUl.tl~.o!l:l'~o ",;c'lI~ nllbrl!fled parients, he says, HI!: plr'\R:'.:.clLlre "Ieveb:the lleld fOi.mcn H:!l~I,WI:ln']!;!JC R'P~H{;lin~bt Bu~dt also :J1rguc.sII,1m~h~ evi-

deuce backs Ill!!:: use

~I'I flJrtilil)!

or ifrv,en

egg."

~-V't:a'tnTlICirllb fop-moB,::

Il;!l~(!;nK-The IccltnIDUig)' is Il~cre to ~.(Jok. rC!tlil OJ1 b.omki 1l£.1 [;IS .1 fi.."ah s:Q.c opt ion" he Ad!ear llIiUerelil~ •.A Imm.!i,11I0e(~le B It! tid h r i"bo~H5 lu II ,oh"Olte, thOJlt terms i (C ~-ay:.;" 0 I. [i _:ii [at roze n Qucytcs, even oller advantages (r), sta ls 'l'l'llelfl fro~ell $i.o!'ll'l. fa ster ;l A une'i\'l.cdHI~.qUC fur frc~.~in9 can mi"un1li~1Clee cam~~:~; a over rfl:..""h,(mc""':'l11~IVFpJ'tlccdlur:c ~ fl't.'ezilllg ei.'!!s m,igiu produee :saline llO'~iJ'UCln 'Slowly Imlien na$ a hn ~s silmp~er because 1:l1c,egg donor ~und lf~iC:ipielJll don't need to he ~ just ~'~~,n'!J1<! !Jl impact ",S ~ ~I iCil! (i1l5!l'{.. lejO, wil(UII!Oi S til ~ 1,1 itrifi!l'di solul.i.tlni j,sdear {i1l1!l'fl', rfghd. ~ ~.'cS;I. l3<llhi ng eggs ill ,~ryrepn-ld:ucliNcly sYl1dm:mimd, iii: 0 p ro tee lmll UlH.:d \l:l es 5 uch fl S ec Res.e3n;:hefs ''l.gree (In 'IMhafs o!:rr~ervruh:m dmlil)g Ih11l1119~7 [0 ::il. P':fi1op,m,medioL cd~.ylell.eglycol:, .md III is£1 rro~n our ~J:nd~rsmndhl£ ohgg l1'et:zIng ~ s~~<;f(l!'ie-~'~!ll s~.~e"~t.~[ ~~]l~ I'h;m~ d,lIf] ~~.~ ':WHS .• ,~m'~rin~ 1.)7 'cllHdren. O"~ro.lll. s~.ow- ~.");l;,I de;;Lr p~cl;m:eofth~ h~thll1 anile- !:e"<;\I~tirl!il the. ri"~~ZCUl ~ggs 1\;11 ShtH't of ru'~sh m11,;;::O;,t!fIJ li;:''o'1(;r~! (."hildr~". Ahh,tliugll MHl1t': !>dl:l'Ids,l~ hin'~ .cI hll:~. dOCM11, 'elun~.iII,1LGl,::n. 1rhl~ solutIOn. ,illlany sc~ellll is:ts· Ilii}\..... gay. g S <I Ill£thgdfrlv $u~)eliifiI:M meas.lJre. iUllc:luding feniliz;,uiQIII ra~e and Qn'er~pKliminary reS\IIt~, IQ;r :l1man mllnbel'5 C(md~[(,':led, .ilJr{-mfl~~errI)!. illf~rl:m't~u.iollt'.JkL rn.l~1t:': W~reli!'l e'irIIOl.ugh cltl,t;l of c-hi Idn::".!:I.tm mu;;: 1~~IS 'ill dinillg klilown "s \ri~rific~tiouiL ~o ~ 'flHl .~1:,md"r:;i h.W>!l~lr'~~~ngnwtho d-~ Ive Of!vi~fi~l!qfln {M;l ~o~np;~rei~ Sk)Vi' freel.iQg, ~ ~omlJ'lrehe:(:!s;i"'e llotu.d}' (If b"bie:- ll(lr~l frnm, t.l~~fi:~;a:::~:rd'wfSL.'"tenl1l.~l:lli:d.1oOCli s.ttys ~Il:,'lt Jd:rm;tli:(1 ~g.g~, d P IS: te::(Jlf.li'i[~ CtY{i E~1!;~lk~i<..,~with ~](I,m: m{ldi Fi· l1ml CGuM :ooUiii ~hlillgc" l1\iI't'Jjj ~ , . her e:":::pi!ri:ence is. svmillJ1r; Abmnt 3% 104% of and ~(l~ !e:O'I,I:!lLes ;,w.e \\1('I\1:kn:g 1.0cl'ei1Ile'.m Illlel'" i :Il: '!:•• ~k1n5 el1li,li,ls gr<ldWlOl.Jly c(10inllSlhe eg;g Irozen eg~s \viU l'~l1!rdII [j;llby. \,-en,l,I~6% to 8 i£lVer ~1Pi:lUl 90 millu~lI;s i:!'l ~.i:(miII'Hlletriel d~ll,ulw.t<:ein ''>'liiich ~luniC'imlls I.;~ui .vecnro ::1% uffrl.,:s.h 't:ggs. :5.h~ ~iY::;, elmlro.~lcd fret:'z~r. By Cl.IllIlliiJ>l. r.,;t;I-freezlliig ~nfarrml!;i:ClH. ll~e ~ il'l1!i.n.g i"~ The aMest oI'l,f1"" OlpL . i~'M}lv~ JUllikir'l,~,I.h~ l.' b,,- ime! ]i(111 n~I;~lJeCri: "l!!, i!:l fm·r1.1!il.~Y 1;'ih)'1.ill"hms t~ildj ~lhici:;;l:... ~ht:: ~ta" l'il'ir~li~ :it!: l'iJ1~.\."rnHg llii~i'r·~mJlI.t(;livGyt"-!In;, ~o ._ 1 ._ t is.t i(."s blu::k Iimiih;:J LH~ t) f cry...J-[)Ire:i:CI"V<'IlLcm. ~ it vi'U,·iili~. ~)I" m1n~foI'l1li'i!:>lbtD~ ~Ias~y I1ffl<itcrfi;il. i ~,t wi~I be· po*"ibll! ~o deb! 1'I1fl incv.11el hell th.eir Tit!:} ide.! ~hillid vitrifie,ltii:1J1. ~~y~l~ibo•.~~. tl'l rL!irlil:~lyi... ~(I!TIl"u~.Th<Jt rl.f-~ul;:h tnijilht 0011 u~ Gliiddune~ ~rL;"I!iit.>.cl by lhe-A~lwn(:"11 Soc.i~ly I'lu R.C'proU!uolltive Mcdidn."::' ~ASRMllasi. 1'311 m,ure al:lmLi.Wh~lhl::r 'L..'g& freeZliillg th:~l.'rl,l'Ci> .Ii] "·oumll~eh:iJlil1lg injury:' :rl'cel~Ilg th.c ~~g r~g!nu Ihl." ptuced~l.ru ~5 I!xIJeI"Um,~i1H~il ~LnJ W~lJ'nlm.~ptcon. --Mlirel!llU.EflL!lE ~ ~It)re ic~1.:~"'H'ry.;taJ~~ i;ru"
m.e·.[N!il,r;U florell

oflh~Ulliuf Buiog.na in ~.il;jlj' .2: :ilUlid 'J;;:oUe~lle:i\ .repcrtt--d the .; finn birth frnm !CS[ on ~I l!§tha\\,oo eceyte,
\'Q;n;;,i~y

limnju~l cancer

i:~

--

'I~

l
Ii

\l'!llAl'w.sden.cema g, mq

SClI:EN:OE

VOl316

20 APR Il2007

Germ Cells
E:iliH!\'(lur cml~ id~ rs the ml S\!ilerS N::Isuch questions l.9rg~ly unknowable .. l)p:iclIIlly. eve hu iun;lry ~lio~gi,~tspiece together the o lifes~ ric of a long" goue or,£f1Il~~l by scmfl~ S

A.develolpmerw~a~. biolog¥siI: takes aim ail:understandling~he ,e\l"'(l'~utiona,ry' mighlS o,f leggs amd sperm]n om ,600-mimon-year-old ancesfnr
F(~ a c rC'a[ u re [10 cue :1:'1<:'1.50 C r seen 1:1.1'1 d r C'\.' never wi'l ~seeslive, Urbil>lleria g:ener,m:es. <I Iial. 0 r pass ~O". Scient ists have Y igorousl Y debaredwherher ~t :sponed legsor amen-

lfI~nll 'th{;lnmd~u'! ~~Pg;; ()f·~vohniu!1!;Jry trees for physieal trnirs u:tl!.i.;'tt:.lulJ:Lm::mtll !,';.clliclte

nae. '1".h~t her iI, hall:!

ll. I rue

haarr,

[md

whether its bocl)' was $eg~lI::med. Such 'l!'~~unents QH,IY !leY'~r Dr;:: ~n~.ed~ iuml,'~yel·.

Urbilm:t·eriOl is :1 hyporhelic;d organism th.u ~in-d 5:50 Dun tt) son mHl IOn. ye~m; ~Lg;O

mm

duc~km: the o.rigin~ uflhdrspcrm .. m11d egg. llc'r\'\:or~ "'IS. ol'igirull:' ~~ay:s aleobiolog~5r p 'D\) ~g~as Erwli 11 of •. e Na~ h iona~ Muse um of N~1.uu.,,1Uis~{lry in\'\hl~hi!'lgwn. D'.C..\ ..iho ,,,i h le, BOIh e~g I,wing and viviprnfY, ill \'I.·hieh ml. em brye clt:'ld()psiln,~.idie ,~ parent, has wriuen '~unu~icl"[~lbly 0:111 .hclII,tL~uU;of are' sen :t'iCI'O-SS ::1.:11lades, I'm' e:>;~:lInp~e.so c Ur'b~:Ia~er.i:a. "_h '11C\'eI: dll.w~ledi 011 me 1;0 COIl!'i~d~1;r lbi;~e ~orls of issues," Wilkimi ;Jg1p~el>: if~h,ud fur re~t:,u'(,:hcn, tu t·eU w:hich: Slf'~nI.:~y e\folve\! n~sl. "A~ far as lknos v, this is rhe first serious
W~tU.lpl re lcuk .n lh~~[ n::p.m.dI,.iCli~ll.1," Mo:re!Over.[::'!.lm. our'sconteraplatlon (I tlp{r.i~~~~",!i.~y h~L" ~~d Iller to relh~~" the ~o~ hI ii.on ,Iii)" oml r!'L.1t::'i iom~1l:!l:L\\'\::t:nrrb.norti ia,1 !1.IC:f:~l C'~1I~,.''1ihichi!Jlh~~mudy make SpL"e.flI:!

aetworks shared by II1Imsl or its descendents ..Traits in C()lfIlIli1I,tlrIJ t~)II 1iJmad spectrum are l~'kely .•ncestra]. [hll so Immy i reproductive ~rraregie ~ ~LI't: strewn uc lim;,'. .. every b;il~nelfi<ln phyla ~~::Iilthe idctl1tiWuC3~ t]OIJi or~HI <lru:;~~'iIJU:iiI:illl1;:i ~ prub<.lh:~yi nllpmi· ~

and was the ~.fLSI.'011.m11011 ancestor QI";.1I !l1e!l~~~~,~e lh~ti L~c!~I~~S l!lO! ~.~i~k2'l.\"'Qf!'H;<;, Ili~~+ mice. ~1Ild pcopk.

P.icking:a mort:'

llI:m::l •. bl.\:,.qUilO~I[~JLl , LI~){mt

l:.lI:bis.ex~IJ~ EXI;3VO~II:' conecnrrated Oil i.~y.,. has \~.~l;Jt g9~ her th i!~1.; inil ,I iJ.o~H Urhi 1;;t(er~[lIin

tlre n.[st IJ.lacc.: primordil:!.~~fm

lIloteslh;rl these germ-cef

cells, ~Hill:
corn-

!Di'~C~I~~~i.L!J(iIll>!fthi~ t

mote t:l~arljnt ~nd e.ggs. itImlihc stern cells ~h;[t~i\'\! rise to the end of .,1 Inns diJYin l~r:lalm~l~()ry, ~,~y otheiF tissaes, rraethtoi'i~r=, of ~\I(l··d!;lvo. 1.1 l'id.d1llll~I.I. u~~.:1 th;:\'clo'l~nl!o!nt~1bio~n.!~:)1I study evulution h

[iny~bucal.~rc:~Hu:r~ an; pub-fueled spt:clLlI[]tion~ a.i

p~i.':)Ctl rsors

pei,c·with. one ml()t~er Juring dev~~Qpment

within .t~'~hl'or(ld~, p'rimordial germ The cdll~ uUkbil.ari lOt: in UJk~ rettion u ftlltl: l:J:nlJryu I:'i~d mi~rm:t!! rel<llive~y luug (iitst<l!n ces 10 ,~1l<l~
1b~C:(Hl11;'!$ .. ~"m:!ldi! 1,~L':r~

IInlldct:

versa .. Ahholl~h

sOnii.C·

animals

Uley devtlJ.op into

hH'I,'C':li\l OHl'-lturn ic <l~ ~Y!111lii1,e~I:Y Sl}nolll~e~} or ! j

Ji~:p I u)' r~!I.Iial £yI1U'J'l.~u"y (t:O"d~~ mul j e~ I)"~ ri!\~'nI. mmilli III imal phylr.1hilvc' bi I m.e'n1i ~~y Inmerry, :md U!,b~J"tlf;lri,! would tHlv~ been thei r foreru nile" A~ sueh, U rh~1 :ll:el'ia offers
:I

fnullc'w{uk

fur lh~.[lkin!:l :Ibuu[ hnw

I't~ntdllivers~ly bilaterians emerged Ihurn the h'l r i.... IN ~ml turns of evnll!t ion. .~n 200'2.. C;,v.;;~alldm E;utlvol!J[. a P!XslJ,tM.; vn Niid~"1l1 Ak.,nll,'s, laboratory at the Umivur-

or

the

eur-

eggs, Bw studies had ;-::iQown Ihu UJ1I.y s.om'llof ~il]~ pri.m~ru.i~.lgl!ni.lI. cdb, 'I'.'i[l this raee DOlite gOI1lj)d;tllIe Im~COllllt:r~ \I,milsh fmm tlle ·~m~ryo'l~ so-eallad germ line, i!~ cgg~ ur sperm, This I.:.'l:.mnph: survival of uf [ih~ ntl!:,:,~[-Or lit~I~:sJ-I!L:dExLU\lou:r Ult:likL:J Ili. b reader Darw in ian look ill p ri 1'1110 Irdi U ~
Of

sperm

s.hy of C91l11bridgC' .illl the U.K .. realized !i.{limelhing ~'\'<!.~ ~ssil1g fl"mn all the nuus;~ m
i:ngs oil, Urb.ulmcr:itL. "Sn minch or t!V(N:lCVU

geml.t,1eHs. HI sraned ~(l thin!!; .,hmH the roJ~ of germ tt.~I::>in evohu iml a nd bc:g~n \\10:11· (h::r]:tlg 110w ,ULi H"cn:nt <lin ima I S mu ke genn lines," she says.
[\,.llllchl~a~een "I.'rincll ;1bc)llt whe~l :u'Id b why ~rlim'llls.~ln;1 CVOI'l,lltl a gC'nlJil,line. Smmo researchers have even ~lrs'Lled [11m the emerg.e:nce ofsp~lI!luzed repm~~b!;:t.~\'IC'{:'eIb;~i'iiti~c;:1t fmlT! other tissue the sorua=-was .~~ prelit'!~1lt:;; ire to r UllLl, h ie ellu] .. r uli.gu n i sms. i n EXI:mvol~.r agl"'Ccs with Ihmihhnking. So. a%'!.lIt~in:g ~h;~t"g'em~. HI1Il:: p.waletl Urbi 1;1!J::riH,. ~tlc ~:sk{;-.dhow !;:l.:rtil:i.f'lt'c~ i iii. an U rbi ~m.,.;r.~.~ Is I!U1I.lb:r}"o~Ildi:;dLLp OJ),; aJl!~ '1,nJ.1l1!ll.'k~er~ll c;:(d~s, Illot fl heart or <Il1er...-e cell.

over thelast :20 ye'llis. eenrered on wl~.al Urb:i i.31erd.fl looked i.il:fc." ::HI}'S EXl~.voLlr. .. "But ne ane [WOlS] talk~'Ilg onhoul how this
jill; lTlu

~ft:pr{JduccH1.;·

::;,o.:in t31~s a •. till: Society Ibr ~m>egr<L~hVe

and C O'tH ~,r;~!i\.'~ .Biolo);;!, OlLlfll~ .. 1 meet i,~.g Ln J~lIluary. and ~Il bonk dU'Lp·ters illl prc~".
d'l~::>de\i\;lopm~n.tO;d bi()~.og;M. 1m::>be:gl!.m ~o

disCII:»

\\ihfL~she

deverly

calls

1~l:itJm,e iper.m or 'e9~g§"MU'lmdl~!>lIIJ, a pmte~1'Inll~ilI r V~~'I ill k) rev~OlII ~(h~ JelOlllively :r~w ,ee:tls. in lh~s
I;m~ta~fOl n. @moryooes;DliI@'Id

··~.II·b\il$eXLla'I,-

'p

19,be~omf! 9~rm I;e~~s.

~lY~" Her OlllPl:"o;!:dl rIiI:<tk@ji Sie"nse, ~:J'y~ Adbm Willd:tI:>, I.:lditol' UflJiOe'iY",;~.1':'! un~d Ii g,.:rndj.~ ci,~l 'ii"~Ul s.t ~d ~es. :s~x d~t:i1:rn:!i [~~~ "\:\1'.~ ~ O!iL ',JI

.~ikc h.l knuw \.",l'I,nI~I'!: 1!',,"~I,iest bili!li·':r<ll Illflii.-· ~n,.1 I~,oke·'1 ~ike ;!!lId.;;Icled like. and th,lI s ~n:dLide.i tlt~i r l'\:'plI'tl{lw::t~'.11: h i~tc!I'Y.'~ Tlw s~:.x lives c·f lht;;.~~ anci!.:;~t c:r·L';:"'ll,Lr.~li h9.~ dUth::u Elti:l~\lOur.lnil. sh~ h~ll> iil1rt1J.(! r:'·~gre~sumle f~! .md ~I'Ig (In,e tl r l h!! In,'re lu II'IIdlll'ttlt'JlTOlI ~SPCt'l!!; uf Ul"bi~H t~ri ,Ill ruIJWO~

~il~erll ~rg<l!i!i~I]i!~ tYllic'II~j'(lse (lli!e of twom~th 011:;.~,ogivl: !:',crull t~eltIs tI]c~r liIi[i iqlil: 1'Ules~"ill?t>:t q~~~~lO!il" a~l9~t ~:rb~~~"~"b~.~ty idenlul~l. By {lne ~O\lte, rreRm,uation. p<lrl of obviulI.sl,y CtUiit:!.tnl 1II,b:ill'Heri<l.'~ ~ex liT!!'. em ml:fcrti~ireJ egg's. C'y~op'b".sil1l,cunt<1il~~ ,I !HiU mlbrn.E! ~>I!9;1fI n11D119S
\V;_l s the ,me iellit .,m~lfIml he.m'~llhradi Ij,e"! Or \.'O'1:lr·,: lhtt' 5CXC. r LJrl'ii la·tt:'ri ~I~"I:p~Jt'Jk:" a~d i r ....0 !ol 0,. d idl :m:p!l"od lie '[ iO.f[I r'L;'lly ~1I:i. e'." levliJ:11 11 r i11th.:rn al :Ii.-rti I i.zr:l.li oun';' ~f tilt: I~u~:r \hl1'~the.: C:iSC'. then Urbih.~e'fi,l"s .~mmds: \.vould hltlw! oo.g~'tilli i~11 S~1i'p1m~~ai~a~t:J,

d~~liD1c mi'>;. 01' prole il1:-;:. chemica 1:-. lind . , i'i\'e gt:' 111:~ i11sU'u,c 1~.Ul'l~. 1r h b'i1'i IW:: ri~L is '[~t:: '~ ic ~ SCi-"";lil.!!ll, :b'.f'rnm~~I~::;nLA5 the egg Ji'l'iJe~ ~ ;;uradrediv.id(;'~. hil.~ of g~!'!1Ii!P~~L"r!il ""'in!.! lJ]1 ir! 'Ill >J j;O~le een:5. hm :IilCit other::, Ins~detho:-v.: ~e Ills.. :l5
il ~C(~ oA'.~1gt~I1!.~lic CI!.<:c'"tI~. GC'l1e£sud~:l~

20 APR~L2007

VOll'lo

SiCI'ENCE

WI'IIW.sdencema.g.orQ

SPECIAIL.SECTION
PROTO'STO'MES
~ ~ ~. ~ !~ C!o m ~ "5 it "" 8. '" ~ §l' ~ '" ~ 6 'E ~ '~ -<= '''eI ~ 0 ,~ ..., ~ '"ai ~ ~ :::i .2: i -"= ;'" ~ 8 .i ~i t '" ~' ,ItI IC 5l a ~ "" ';
IE

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CI

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..:i

:19 ~ = a ,., ::5 ~ '2 = :~ "l! IE" '~ "

'lEi' <; ,~ 'Q ,~
'0 ...J::!I <II

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i ~.

"! ~~ m
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6; ~:

I ss l! ~
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IE

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~,

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... i ."" '" '" ,09 '<:I .... .s ~: ,j; '" ~ "E 't:i '';$; ~ -iil' £i,..;. e! ~, ., t: 8 t: 'iE "" <'!I' 'II, ~ "" s:
.£-"' Q!'

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t~.

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-""

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----------------------

Moo ~ e r (;.~fm (811 S(Jg{ i r~(J I ion

Eipig~n~ili, Ii'mf!!111'1i1tion

(h!losin~ germ !C~ls. Of Ihe two meltloos OIiiilrm,al~u5f1 to spmly Ilhl'!ir germ cells, {>pig:erLE~i:!l i~;m(lr~ Iilid~

spf'Nd a rll:llikel~ to b~ t~e' IIiiD de used b~ lirDil:!lt.eri~ (~l1isl"~ re~i~@I'l}. th ~ ~rtijiL'S,t bil~!ef~I.lIlima L
~'Q"'1a 111i.d IWNO.'111(!!e,pJ'i'le: I!LC·liv~. r~te;:\;:al.tiJph~. ;!'ind ~~d'pIHC]llll"<C lf~~ 1!'t:11:;fur II futur,;: ;IS : egg:>, or :>j1t:·'I\~. hil lill iul1:l1.b r.dy img ol'l,Pf'~" fQnmnian. thaI £e'rmplji1,:;.~n is retained ~lr

l:aI !'iplgl!i'I'l'5~ ~lWiI pf~f:ili!ffi~~n

DiWilkil;tJWJiI

r~_"lnd~d wi~1\iin I!:I~W ~oc)'u~~: '.hu~.~'t [s if:lJ1Crited :rrom,g~I!I.i.:'i"'Jt iUI1J~:(I~C~lC'm ~~cm.

Where lhefei~ nng;ermI'lIOl)lm. epigenesis

eorues lillll.o ~~II!I,)'. U the eells Ln th(." ~m:ln,'O A .);!:<llFl ml!l.Olpparenlly eq I,I:.!J. B!.lt <I~.dlc !;.!Itd)~yo
arews, ecrtaln ecjls release chemicals th:u
l~lo;

~ three their nei,ghbnlr!' to become the germ

I
~

~ .liJlJCJ_ it: (llICs;~ion H

for UruilaH:;:fia~

Do

cells tll"~decide to he~omcir~ prim.ol'dil!.~ ~ ,gcrJIU cells ··.illlilcr.it ~omdhin~ or J;CI un ~ in;~m~(;1 ion?" asks IXll!!\'0 In.'; 11;1
The n:ln~il:i!L:relies

in~id~ rrimCl:r'c~i;:!!.1 gt:run c~R~ that clriv!l1 th~ ct:Us:' JHTelf'~Uliti[fjtiL1un inte :!lpt:,IiIb1i or ,e,M:> !Ji';jill.i:;.ho.Ulic~lh.:,njob is dtln~. IElo!~a\!tHiL t ~1.n!.:)'.;th",t,~ t :l.",~mt:poi IU P(l;i::t-Urbil :Ib!'I'U<1. a It)lutO,!!:tioll ncc ~l:ltrecl th.~t ,I Url\"i~d thO~l;~~H;HIU to p e rs iSl,ill an ooc)I[ II!-'I"O i~H.preferrnarica. It iCm.lM h.we been ,I !>imple m"~I.te'l! 'l) r ~'I;."I,jIll'- ~e nes sueh as vasa <! nd ,m1'lm.~ ~lInl;[;N~ on i~ f.lOOq~1'instead o~:'lutl:!ing dll:nl U:IJ-50 simpti.:. :i.~ll::aCl, Ihllll E1(m\U~1:i suspects thls h,lppeneJ nmhi,~le limc~ Him d iIf ~f(ll"ll evolut i,tlrI!IryDr~Ltldll.'i s. Ift h11'[ \; true, she Sre(;Uble~. then;;! aremodern animals able ~o\:Jl'I[lI.loy both modes of gcnn eell specificatien. Evell so. their embryos
I ikdy I'd.)' ,r]ll onl},
0[1(:';_

htl!aHI['[l~l: Tl1~Y~lI'rvi\"e:

!:Il'lci divide

lor ~i

ltln~ tiI11.::'will~'!.:)lIt ::;~i;l~a'l.izing:.

NIOi"t:oV'Gu. !'~~t:;j[J::'~k,:r5 h.~u\'c l>howi'i db[ the rd:inlord.'f1LI ~ot;;'nil cells of sm1il~ sJ'i;,"L:ic~ •. indudirlg Ill!l1'IT!;1I1S. ~,iI'l n:~as sO!l1i;llic: ~11i:!1'I"!, " ec IIls, g:i v ifI~ rise tu more 1111111 j lila 'rL:"proc.lw:nhlc cetls. An,~ ethers are !,;':oa:x~ng nO:ll~~~'m~iI1iC S[,t;n1 ee II~~ fmUlil '(."3 L'Iy ~'I.nbryo." tu de\rtllop iJHr.I :;,pL.!rr"H'I Jl:ld eM.g~ r,l".f~".!'·(! O •

23 September 21lO5·.p. 19~.1). Cell hi{l~ogi~~ls~~rw.! eonsidered $om:llic SlU:nl wi b to bIJ d~"·dopnlll..'ir! 1:0. spi rll:lJf5 (Jf ~ geull 1."e~ls lo~c-.m~1;! or th\;,gr s.imil~~rili~s in !!lC~ll.:: expression ;}lltllHrr~r'l'!mimk~l'i. T\ri.iS!t.~ ~ng thar idea a bit. Exta\'OUU'byp()thesi~es
that germ cdh
1mHojt:as ,I

tlnl 'I::r:ig<~lltl~3~: CcrkLill

subset

'[]I:!:"

stern

~ cmbrymllk
:3pormdnsl.lilm

eell:; nJc!lsc bO:llc-nlQl.'phQ.!:leln~c

!,f:tun COll}"

tell other edlh;, DO become ~ cdbi. I3l1Ji I,in.e U10I'e primitiVoe 3uim;.ls :~ lIiIl()lrd.y SHlduoo in th.e I;;!b-zebrwri"h.

I
I
0.;1

i1!cs. mem,alOJcs,~rogs-illl depen.d

!l:[;tiJnp~l;'I8;tn 10

, 'hhdog~sls Iilad <lss~~m.od th,u pn;:l'orma~ioll •. \""'.=- ~11e'l~lCe~u'I,I~I.He-~n revie\v ~l~' ,1200J ~ c:;>;i:;I~.n.!;Jat~, 01112S ~Hili:mal,P~Y.13,. hOW~YC:f •

'''''

~'Ofall the cvo-devo sterles, EXlavmlr"s \"Iurk b affillu~g the mO~I. intciiCsting:' say:; Helen 'Whil.e~Cooper of the Uilivershy or
O:1I.(onl U_K __\,vfilo ~IU-cl~~s g,em1-(:'e:l.i dle;::'!t'CI-

cells, She notes tlfl~H pr~Il1I~JiY;;: 11Onibi:la~erfLl a imUI'I.MI S suchas !>pcmg,(;5Inld jc Illyn s11maiulain :HfIH celb~hal,Ct11l
Ilic.:W~ml1

t'ruil
011

sp(;dfy t.hck

gtlJlltli C~US. sO

h'l':r C'.lse, txtfivOur shuuld: lit}' ~o confirm !W1~ethcl"<lll aUlhna~ .kI1lO\,rn m I'e~y lqpan. p.ref;mmltinn "lli!i~ht be "hie ~o~I,~e :,Luoth'er m(;;dnalfl.i:.m If i"o,rccd," ·Wh~t\:'.C(mpt::r ::;,ugge::!:!is, .~ncleed. IZKI3~·(Hl tl ml ~.::.,,")lln r l.Hlf<l!ClfS

OIHlll,em in :i:'mh. nics, 10 strcngl.hcli'!

lh ~ s~C'm C~ I~:s':1'1' ;spcmgcs allli. Jelly I1tl~~~:t"C dl sPllffiCd ~.hra~glil(mt. I:ht:ir bodit:~, :;.h"lii~g-

'~i.i{:' C'~n~ or

g.Lve 1.'JS.e either 10

~;,'Llrnleit(l:;;,B~c" lL'ie

ceHs, "",'ere sim~I,~.rly !i1;Olt~red. A~ !oj(:ienti:>t!'i d~n~ih' i !11(lfe

gCS~S~~l~U Ul'b,U"lel'~<I 'sg\:I".In

!j'l:'ilCS rhi!l.l i!I.rC'.•Lt;tiv{; ill bUIll. pri mOHlia

A.kal1fll f(lmld Hltle cytoll~,g:k~l. ~ cell Uil,eilgc. Of e.1I:pcrhnellil<l1 aalato hndi.~ oC,He prdrln~~[Itio.!iIin mos~ :~Wi~~ie~- They
• 1;;o.:l3i\(lUr Ilmd ~ Ct)i;u::liilkd dl;:[.t ~pigl~l1t::$~S V;,'~ll; ~r.y wid!.!" "
I~

~ '$pre;ui ~md that

oJ

IHlI'lOO,bl)' ,u'ose i::n:st

F riiim

st~m (~U$

'l~' gili!:flml ,(~U:;?

13;;:11~VOi,ltll~~ t'~)m~ nlmtil It! ~Jntll[~s!S da~ sri,p· i 11

~.-." 'l)l"l ~'11g ,h.lt !:lI rpl'.i.:::e· mil dI h:l;;; ,ciI ~o de'l,·,t:IP ~ ~1pli;lcl <l model. (,H!.LU1I' rrefbrn'!,,,uo'lJ, I:cmld ~ ~ ~~V~tl:ti ~Clil n £m1':c ~l,f U i'bI n3~t(:ri a's. tl(;st:el~·
~ ifJ,t:tLl1;,

doingjust t~;)L I~l!ta"om w.n!~:5 mtl!il~~!"Man~ C\'ohitinnary ~i,ll,k$b~'t\\lc~:np'hl.'wrdilll gC'lrm !!:e~ls "md th,e S{lnl~ti~ stem celis ~hat 1li\'tif~~~ ttl .iliU. Iher'ion.~nnline '~$SLLI!lS iP1 ;Ul o:I'g'll1ism. The hvo ceU lype~ !'i.hare mil n.y i.n,o[p,ho k~g i,t:'f~r~!n s. ~jl~d I i.i:ru certa iim £,ClIles 'lila!. gu idr.:: som m k :;I~m cc II ~ro'\l!,'t~
fiI,o\V

are

Next

genn cells <In cI ~Qmmk; sllnn ce II~,andlf;,,,,e their hhuol"Y. 1i::!'!:.t3VOWlf hopes 1.0 conFirm, this ~\'ol~lti.on;;nrysC'erl:lfio •
\l!,'L~kilb;, \,i,l~lOphlltlS 1.0 c:oU!i~mll.kwi.ih [ ..... ~avour•.c<lll~ iller 'Lm(;on:V'entio ~ rev! s,ion mOl o hh.~ hi.~lory tl f g~rr!:lli;;dl £i, :!!lid ~teni (."CHs.~

"b~,g ide.,:' Andllile thinks

Ext.wrU.lf·$ i!lvt"sLug!itit!h orurbi:~1:1Hi.;lI.it.y \.'iI'UU.drmir ml1,L"r ... 1,1],) p,ro be t I~,t:~\"iJl:u t iCNl 0 gcuill ct! ~:I!\ !m d

r.i:I t1pl~It~51,S,

'!l'ln~W~rehH~dtn !:len!:;;'\>,U;li'\'~ in rd!lillCifd:i:l~I!j~rm ~d k Als,u. ::!Ii the l'lrUl;St !:Im.it::

nl.allY U!! th~ m,o]l:liCul.~s

IC:': "~ l.out

n d~U;l>11'li C! r ,t'~1 .1iIr~d h

5ClI:EN:OE VOl316

r~'prodllc'~iv,1i' _"y:,al;m~. Tht.: ~UIfl,iL;:. ilkills W 1aughs. "wi! ~iK: ~ornc .. I. e~un [CS.iSI, i!:3yi ng_. \
~~" i(lit

{I ~,~

~o

lltl'liHi.dy:'

~,JOliitN i1'R:AlImS

\l'!llAl'w.sden.cema g, mq

20 APR Il2007

3:91

Germ Cells

G'erml Versu:s; Soma IDe,c:iisiion:s: Lessons frolml Hies and W'D,lrmls

lihe' early el1llbl¥ois rom1:ed by tile Ius;iolll of ~wo g;~rm ~e'~~s,~h<lll m'Ul5t glmerone not onl!y all Ci~ th.'!:i lloJne,prod!l.!.ui:\I'e ~m<!liic ceillypeso ~fits body 1t1l.!lt,i'!.~sIllUhe 'gemll ,c:ell~ror the Ille~t.
'gener<!lliiol1. lUlie.riliiror~, 'Iii m~IYO' (;~~~s,I(!(e (! enJ!c]a I d~(ij~iQIll: whe!her to d'e~!Q.j) as '(je'rm 'orr ~'Il1I~, H!Ow ilS tlli~ rlUmi~IJ!1IeJlf~ill clKis:~oon ITbildie andl ~I!rlill (:e~lr.ati! mainta.ined dun Il~ d'ev,e[.o~IIJlI!f1lt? :S~IJdie:> 'in ttJie nem.a~odl! ,",OrIll! ,r:;~eflcrlN1bditis e.feg.ans!8!l1Id rrL!~t.fly .D'{:o.s~fJflita'iden'lify sonu of

~ :~nd 2). ()m~ likdy ml~ i,. 1u PJlnt(.'Cl early ~CI!iIl! ccU:-> ri'o.111C';'1I'~lil]g ,<,Dm.11lc d~llIel;l."'Il!klt[{IIl!l,c;Jl~;, 1hu1S11;ri]'l'li>t.)llilI quiQK~!~ i" a,hfc"oo Ihmugh a
~)iiliOil Il~]'1~~;I!1
.~Jjfl,l~$

,h~ 'CflN'lirTI!~-

N..-rii'liim!<l1m<!£hii1Cfj' ,~" ~wdJ as dln.1flUtiu :'1!;It~ TIle ,~~~k'lllf!lm;;~ or rl.!~WIi.'il$Cn'l it. m~'d~u~~ aL d'lC lc:\'~1 of U'an9:fjl~ i~1" dOl'~"lion h~ bnlh (lI~1I1~i:-;n'L~ ~~ 'he l4'~Jl11! ~~Il.C'u!ld ~~ll'lQ ~re ~injg scp;;l~ncJ linm ~tch olher (4)1. rmtdns inv,[)lv.c{~ inrnrC'l'C!1ljllg ifNi\ IJIOlynrK"t'Ok">C (B't'I~ II ~ elmg.1n i~1:I ~n g.,;Il'I'l l:o.:I~, Im~!.." IJ,;..,.,1l id~llli ~,.."t1 in bi:llh oligi.illlli:m1.~, zinC' ~illb'Cr prut!:il1. ,~l.E-1 i:; a liicy The Ii\;gui:.1",t'lt ~11 ,~u.n~ ,t5). In Cilli];;t)/(IS 'lImt it!!:::!.. PI E·I 1I~1.i,,'II.y• .Il,.;'wlj' ro:nlh'::(], ~1TIlrl tid!:.; ~{ojuilll in c~~lIIg:lll.ing.lJlo~ ~I and p.ruLlu~ :l.lIUII.uN:il1lRN As .
(I, oj, 5}, I.u l.hc:~!Jrn!lmlrll:..,
§C:nlll ccli:s is ~h,.;d",vc IoPrI'lCl1l uf

Ihe detis.iol'li-'makung strilte~ie:~,irldudil1~

lr.alnSC ripl~Ofli3~. regUi ~i3;hOJ'l.

Sl!!!;Ireg,ati,on oJ

iii

s:pe(~ali.l@d ge-:lm pia,YI1.illlId ,g~ob.al

'n

c_

L1~"l(lll,~ II.'lnln:yo

SCJ')JlTI""'~ HIl~!:ll,'iIlIil

l.i,l,i~~,a~,~l:Oll'l, ',~tm;~llic_ I~u(;, 'I.: ":I~l::; prn~,~' S:IVoL:: nhr!JlIIl;: h :!I &mt';i, I)'. 01 :ifiy III"

d~\'isitms (.II. During 'lillch of tlmMJ

fll'!:lri!:

roo.•

D~,,~lC Ih,,;ur diil,l..:-IT'ni: ftllmmlion t.icu.c,gil.';.'\. btllb 'wl1L'Vn." ami:! 11[c,~ It:ly tJ'I1 Lh,.; p!L."SlI!J,i;; of ~pl'.Ci::J~i~C(1t!~rlli1 rilbl11 :li·omlllilJ: t'I>t.'l::yll.'t ILl·I.h~

SillJlihlr. [(j, Ihm of ~h.llir ~l'lllMic ;;i.sh:lt ~~(;i:I~ n:!.~lIc !J IIlbl)"oi! cl ie. Mc{;hallil.sl iem lilly •. Pi E-I. ap['l~I,p;;IH ~y !;OI~]IK~lt";liWLlh thejni II or Poi U 10, the <Calzyrnll.:~ 1.&m1 mod i ryl'!'o I u Ib:r ..::lml&ll'liIOB I r:i i. :Ill I~i~, the pt:pl ~d.c [l!lC and dll.!: [r'.u1t;!uliDn~J rl<~rl"llim'.:; N:illJO.~ and Pmui.tiH

cDmrnl

tI'W~~r:i,plitl~~.~ sih;fldtlj;,l H, i 9l_ As; i!'IlJ.ft~! !'llllll[!!ltl". ~rn~ C~U;;: Ih;L\ lack ei~her

dhi~km,·;.a ~"Ci,ulill'd oCYIOI'Ilasm. or "gclU~ nl asm," i:<; fIl<lr! !ti.f'!~H,.,tJ

~fI Qnl!di!t!gl!t~'1l'~H.

the- V cell ,IFig. I )'. t\ n!!1" the ~~"11,[[i{l~1

o rth~ ~~rmmul~m;)

,8'111d S,e!'ml

Ocu::~

r~gc-or N~lr~!<ii;iR;:~r\' iiY ~.. dong;_lliug Po'l ~I . hfl'\,'~· w.~1<.'.s r.::hmil~.cri~ll'Caf !1.O'lnmkl;'0;;>1I)': I 7. s, 101 1 (fig.. 2. A~(,. D, ~'l11~F}.

U('Iivi.ly !l!n~1e:l!fi~~ll~

is

OLlI1l1pklC'.

!I Lie I) -!

cc~1serves as;[he so l~ ildmordh:d yCrl'n cdl ,IPGCl\. 111c j;Cnl'll'lrlc lill~i,;' (lor P4 I'H'(lib;Ably IEB~I¥ I[ PIE~~) ~C'li!!~ ~'I<~I OO~h inl~rc IPGC'$ imoc..: or germ p~3:;11~ I!.OOk eb1si3q RfIIj~ Pt!I! ~I (;OllllpoJK'1II1!o; IIn!l :\l;Og .. aOOmllms L"C.g::"Ilioll (I f smua ~ic QI·~J,ve di,ffc.r~'1]miulion ~b.etDn; ~ma!JiiI <lWIIY ifu;mn the 1-'t~ma l.inca_gc. in Dn",Mlpil. ita. ~~ClJ£nlxm or ~'),:[m
cdl$ iu n.h~~lnhl.'~I{~ b Jllfl dl:1y ~]JbH:illrd durw

These "Irnf'lsfomrod'" o:lls,ci~li;;'I" !i!I)(I\.Yglrl~~'!:\I-pll'lii~ ;1 ::>!Jtmlic cell,

~.< 'I!:,~
~~.!

PG:O~

de'dl1.h p::!lhw'dy,orm:hJl1, :4jmill.i~ ,1.:,.:]1

ili1JC'.... ,[ WJ!.

1Id'cM1II MI"ilYItic C!l!Ib

Iby bud!:lirlllJl

PGC5lorl'l!1J

11~t.' 11J]~c'inlll is 1111 ~ 01" 1]]ml.nl1iSlT~$) by \1ihDdI rgc lind the N~.f]Oiii UlldP:lII11ii.io 1.1·:lmIOl-

tJrmill

IlCpl"I,";!>;smlo1

'I/'I/;'I:U dcmg."'I[[oll

pn:;mcl
be-

'the r,d~JUOmlhir
me um:,liL'k1J:

n'l;~~~J1I.h{'i,;~~!l)id:rhll'l\

in~

1iXlg1'JliC'j;is.. \1!1~u.;H!

uteI'
P,G,(is Aoi:lilvlitu
!i'ltrisctl.p.1iOn_

U1:Llt'ln:lII:)'synLi.~irot:l
~C;IiIIli1

IilCnts resauWlc 31 (~ll)'

I p.ac~m

COI:IllllO'

~r!awa~.

~"~ pok L'If ~)C OCl!:yt..: W~ I l. DLii'iitg~lll!lni-

tifll ~~~~

or ;,.'mltj)'i)1It1~

.b'el~~

n:.y ';;'11I'ibi),l dh ides b~'iIl.i~"iI(llirrnlla' ~t'wncclhJbrdh'isiolls;. llH.15<! ruelci Ibm, emerthc 'PosI~d{)r ,~~111~plasm beeeme PUCs (21,
!D~p a rtm>ell

IFi9. 1., I~Drmatntiinof 1iI(].Cs; a IIIdllalllldmo'iJ·l:; stag ilsin tn~r d'~.etDp:m:eifiltifl, ('. (!fegaM and Dmsaph~{{j. (J@irm IlIla:smis repreSiented ~y red ~H!!!1I!!~es. Key ~egl!Latofs mentitmedin~h!e text illr!! llI00'te!ilin ydl.ow ~!n:.ei, Blue a rrow in i'iln<l~ panel ]l'IdiC1l1t,es II:I~Iif(~ionl olf germ ~eil lliIigratiol'l ilil fJrosopllilrl .
('mlillC J,icfilnC'l!lIs.huILtcl ion. fin !~llCn1,~li'Vemode

~ a1ier P" divMc:; to ~..i1 crate the Z:! ' .'Iild ZJ ,edl:.,. ,IU Ih~ -1~lO-I.C~~ M ~g:c pf I ~I

/\. dlfa~lJ:Itin·b~l'ilOCl ItllllSC' of Imi'lSCri~~ion::d I'!:pressil1n kicks in

\Vf,rUl (;n:lbf)'~.I,!Cn!.::;is.
c~\J IClll)J E·· ~disuptxlIJS

01:"germ ceil s~"Cificmiojl',

g"1i1~nlll. inct~t!in,!,;

op.;mLi:li iii olher ar~

tJ).

'or ~oo.gy; !rn:!li;!..., 'lIniPle~~t)!, I;llOO!lIi,illn)t~rn. Ir4 ~;~,Q5.1J$A. llfDWll'id j.lwgll~ MoI!clJcil1. Ifi~tlllll!. ~kjrmn Imtilut~. Dl'Pill1m~nt cl C{!1l Bioklg~, fi,I{'f,j Yolt: I.!inivll<r.Illll'
5d1100i oIMl!dkirl~. N@iIi Yorll:.

lini~

N,f!l!i.'IIly f\Ofm:edi G@1fII11 {!~~ c .Areilll

a1iran...suip'"

Mdl"rE-llncdkll.;)d H1Ul:3Cnfllkm::d f'i!Pl'!!!S$it~l ends 0 nOne llldicmGr of this Ira~l.~h~(ljJs the i di.~pp;.'"ll!~UliOfo: (m'm Z::! aud:O ~dhiswu~ Im~i" li.~l~i,()m.th;rt ,a~ iil:S::>l.:1iI: ialcd \1>' iill 'tr.lll~"i'ip![i(1<u"
,.11 COlJllPI..'iteJl.CI), stl.d1. ,Il> 1~11.'11:h)' ~~timl Hi.' lYl"i1I11i!:-I {lrh~,..LOI1<! JIB 1(~1IJiK4rn~h Oll~ k!Jy IT'gu!mrnr of d!... nr::u]:;;~tio.n i:-; N :rm:ll';'; IIJK4m.e ,o;;1:aJm .i:;; pur.~ tUitod in 11 amI ZJ, dc:plel:cil {,rthc U',[l N:t1K1!l

m WCli16,

LlI'i1!t.

"ro "'110m(<<~e.~~i\iclM~ :~~ilIld IJ_~ aiXdrel'lerL ~·m:l'i~ ~bnme@ind lanuali:l: lemiill~mn@'l.illUflfl, fi'IlI!{l1Jlll!! .{!dllli

l... cplmOOIl r!:l:]WI~ ur t';Jrly ,grnl'll Cl~~S iu WornliS 11IIt! n i~" i:s. tl:tcilr tr'[lJ~:-iCripuimlal (lllil;t;;;CelK"C (Fii&'i~

Ilil:rta'Uy l'I:>t:pressed St,r~e

39.2

SPECIAILSECTION
I.hmoll aCli""il~ nad [II].K'ilHH~ ;qJIJ>J;'rlrII~ g.t~Dm cdr:... ilililiol;e ~hdrmjgr:lliQIlI ;)Iud I~t (t~pp..~w..ll!)'. in wom'L~ 1111[1 nics U!J !?l (ie 1111 I;~II ~ :II'1PC':I1" t1] ~c l~r!itll~:;L;ly d~p;:ndc!!t on. RNA",mscd .~lll~d~>t~rmmld 11:l;lY bt:lli:ilh li:al~ CJ:lIll.S'L1til"i.;u!ing n.N!Vp~L::{'~~iUl~ ~l>t~hln~' i.ll1t1· l;t~~go gm~wl':lr
~"!m1 !;!,laiUlllk,;; WlC'

,B,~~rtd~

O~~!1l' G~~r5us
~~\I

SDmjl[)jni~~~'oJ!

BL')~h1J:'~al:.cli,rLio:""d

i~~l~

,104

IJ<B!ls~.~hm>1l

('~nun~ IlITI~~ ~<;'"~I,cJclb frrnn C':\p!'~~ ... ~il,lg::.tln"ll!.;: di mrentiatvon ~C!~!Jl'i. (iC"mHfl-M-'ll~;;l troUt>; i:lr~ m~ti.oH_"'. hkhl'l'i'.ClllhlJ: w 1ll;'t';iJ~~n!a~ "l!I!lm~1K in Ilrghlli,ghl the n~od linr ~rnlll~lklll of ~tl1lirlC rellro. !'!1}n~ oo1di~·ifl)lS. 01:' .fil,gll;lf5 Ihm s~'f~d them (1m",!'! a ::;om[lf[iI; dI~n~n;llIialiml piJlhw.uy. In Ihe adult W(ln1h. germ lines ].Likil]g 1!l1...:: 1fl[Jl:sllnliOlmII'!.:Sllll~lt~1r.;. GLD·i la,d ME};:·] 10M' Ih.:ir ~'mll WJlllll~~:1lJ.1d ~s,~'I\"$ii, 11I.r!rio~'\'lo, dll~nl'1:I",'ri~1i.c!f IIM.lLt!llto: :mlJ .lllu~ t ck'£ V 11. eUJl.".~.:r.·;.cly, SOJlIlLiIlH:~'II:IL~ llIo,y ""\1,tl ire pWLI.'lrlion from oomlilimlS or si.gllluJ:.: Ll:Wl Si.;'1lrl 1hem cl(J~'>~] n ~,"'I1li1~JIl"; dilli'i1C1111miou ·I)lllh,wy. l.oss of some!:):!' l!h~ C !!;f.1."m.~ :;.ynMIII" B
I."ltnommin
tomn
1,'t'!!~I~mUI};. SJli:h ,1.';

assembllcs, llu.'OLI!,!:hfl!,l~,Ill", lifc ~ydc, how(;~v'f. .. ~1.:f!1~a~lk i~lfi1fi~~ll[)~v. Ctlfl1jill p!Q;'i.itirnl. al1d RNA .~l!lI.lI~'1~i;l!)' l~metiL'ln. r~ body Iikc':JJ:'!iv ltles mrl} !lll!:-l 'Pc ou~>' 'l1~1~ ~"I1~1 !hm
f.'ej:lL.btcs;

gem,'

i Ii!~

r,·It~_

$lIll:!'!f!i!~~~!!i!d Hies

OuU.ook
In es-

.ruOO?'<mtll:> :lhrui~ .il.'VCnd:'-rnkgi!l.";:;.

UN-J:J{RciiHUh~l>-

I1IhI~h lind: ITi:LIlllllill ~1iI ~II(,; 31](1 prtll!."t'~ Ih'-~11I lroml sOlnmil.: 61.l",'b,. BOlll ~li..nL" ~~U(;' !ifl!<'cii:lliA~u. ~rn~ Iflla:::m m, Ih~ PGCs. The RNA.fL'Clh g-'.JJlllij~ ill !Jl~ll plasalll$(;]'1;."; mit'S in K1{.1iJ m.aima. pn!ICC'[[G:n. IIml 111Ul:>llIIlH:m(1,r 11IIR.\, AK A!llmu.llia 1I11:t.m:U1]~'Ul gCnn o.:fl~f; IIf1C nm ~p;.:;tilll.ld by iiGgIit1· 1'lttOO ~rlfl] pl;Jt;;fla,Ihcy 00 Q!;ml~il], 1~f11l ph"".ll
rol]~pm]~b. Trun~ptiomll K'JX,cse.;iun in rn~~y P{iC':. Iis.a ·l'ltlrliJrmn ·(I·It ,);; 10 f1'l'L"'~~.il. PGC~ rmlilli .. . diffL;wlllti:lIlin~ ill ~ht'SlIiIli:' way :lI~ limiT oormLttt' ~-p~ghtlO!'S,dilmL~~ tile ~LlliUlJfli, ; \,vo!1nP1E-1. 11~ r-~-..mnd I'ImI'l1J111111Iiml Bliu\pl dm~r bl!Lwrol

amJuul'tll11lixm; or ~lC 1~LH:],,'('iwrrl1!.:' \;,nodding !

!md Blb,>L'tIl.1t (~'lcl1yI1l~1C' ~NIJRD, !:o:rrlljll~x ~c~. M B[I...I JI l'~lIi:\(~,\;liI1JIIUItc ~I,hs ~o cli:{pl:JIY gJ.-'f.IlIlliu!.!'
I!rOli1-~,sl!dh

g~l1 J.lj

gm!luk~ arll c!111.1!1.ccd RNA

(Fig. ,2:. II and I'. Thi~ C{.1[)~1ic

in1l!rl'oo.:ncc In.

exput's:;oirm ofg;.'flnlinc !n.~il'" hy 9L'll1~1ic CC'~fis. d'cpcrl<L .. Of! tI'C h~'lDllC ~ J3.KJ f'll'!Clhyhmi'!."llct<l'lC
Mr;.:~
<i!1d

tl~~d~!l~~!l!kliiQ

I U 15). T1~C$C'chITlll'l~niu !"li!g~d"hfi~.• u·~ knnwn

tL~ be- ~ri!lk'al lOr ,~nlililic d~\~lol~mCI'l!. but !'he I'IK'C~1l];l1b1k;d~aiJ:;: in IJiC'~o;omllarc um::k.,:r. R,Ql'~~ ''lr!' G~rmG~,]I')!!~Il::; J\ dis.'lincth'C' LbufWlC' ,ofgcrm

r~~fI!

MR(;-~

~'"f!Cd~ '-fl. Studies. OCIXISS ~Il!¥n,;;-ie;1'IfO~li~c In [1«Ivklc ~llfIl'C' 1l'ltt~Hml~tlc tn."li.YI~Cf; haw gmn lmo r.1"~~pnd. !f<!f!c'tg'ipt[o.]1>111~'Cj;:,L~Iitii:io!l :lflii;'l;lfy !;\,i1'Ill cdb ii~d1'C ~rJ}' Cnill}l<i a! ~dpmtec! #!UI celi~ Ifuc~~"iItn!gJ'fli.!1. lile.
R.c!'!·~iF~"iC(!:l!.an~
C~rrml~iI1itl!',
~Qi!~S

!J_ ~ Si!~lfTlf. ~!'IWQml~.

11]~

:1r",.'CinJizcd Fi~. ,2.Gil!rrn cell ,tril'~ CIl. tD m ~riO'1'

,gcI'In

ecl~s i ~SSioil s o,f pLlii'In \~~lic~1 m1!alll~ ~mi~t.!c

~J:!, !~@nl'l]@lIIlik • ...........,.i~lI1lIi<;I!Ik,Dr'!I. ~OO<~.

R. U'hr!'l~nll\ (1.111'.

r_

~~\!Ir;1~

.~~\mh
11;518 (2004'1.

2.

1!O.l~'So'l\'flnmb~IIILI,_9_t.
A.

ly.poe [(p,) lal (0] <100P9C I1I'!l!Itant [(0) .~ (fI] Drrua¢Ji{o DiaWOOrll1l 'M'Iiltyo~. TIiIf fCD (aFf,'O\~s) ooi1itainlilie ~~r ~~~@IIi\PQIller).\ ~~ [~r~~inIOO !!Il~ ~Dl], lh.ll! f(lIDl'ltiC nudei aln!.ail'l etoi'l~rimi~ RNA Pcllm [red ln (At·(a), '~D), .:1mm.l and IB!I{!'II1I'I!2'l'1lreJ!!11I b:!' (16) arlli (Itil .. ltd produce' taille:tr m!tNA I(Dlllieln (C'aM (In IfiI

end of ,!.riilrJ.

wild4li'~~JIlb!)'l;li5 to ~Ol.IPGU'~it~ {W
'ekln~b~lg .' M

CI!f~lll'!'ii~ ~k

Rf!4APal D, lfBK4n]E'!:,ami raillf?U mR!NA. rn ernl:i~ [~D)~o ~F)l, (white <lrJ'OW,j)' rnmttaiilllfl~ tlve~ Niter;' OIreiilap or greenl arlGJ.rN il'! ~IDO!OO <lj:lpe<vs yelkl\'f.lg~I?iia;da~dkllm m (lJ' \'\illh P~Fl'I'l~OO ~i'l)1lI1~ef. {>ill E!eruoo f'I'1iiaograph ,~r.a P {lraIlu1@ (deI!I:rolit<OOI'lM!mm labeltdl IP~,

mmanl

PGu

i((i11l'l:.l1l~nginta'Jflale:lErtrentdm~ ma~t ooli::M ~ an ~i'lf! <!rrol.'t) Il'!1!Idtilr p,l~es OJI<!d:: ~r!tll~ifl the ~diil~tC el~n;:; !Jet:m ~~. 1~lll'il!lf.dl (rom U 71 wilh permlllmolil fram, IElsevier. ~HI~Wifd~ L:t C degrm~ I\3iNaI. f' 9JdI1Ul~ ,arE r'E!$Iri~led~o the ~

o~r~~ r""e

PGCsZ2aooB 'ilrrD\~.mm~l ~la!nlU laM. P. Hr{lI!lI!~ liDiU~S (>eJ]-i PGL-]) are ~1Dp.l~lI1ye:.:~

('ytorJ11~mi:l: rny.."Ihd~~!J.,"1DOO("d~Uy reRm.-.d Lon." W=mllll?'lItl1l:.~ am! :;poc lirka]])' ealkd polar ~tlj)o Il1lllb 'ill UOo. .mild r ~lr.un:lk'}. in. 'WjjIlTh I FI>jj,. 2. G mn! ill. Pobr ~nuk':> :lIuxI Il' gcmub ':':I;~ITfi;f.I]lI.C rumfllOOn(]f:.~~. ~ld~ ;.l)i Ihll ID\lA hdic<I.9J Va~. jn l1iL.. mid 1lI1~rd~llc-d ~1lInlillli.': h.c'lj(;ll~ (GlIl~l jl:1 ... \'.'01:111:;' rul!.1lh,,'Y roJLI:"~'1mmKmL5 '\'f'!u'CiCi!-...~'t'm,;: prNdmi. IiiL~~:h G;kur in t1k,:; and :~Gl· ~ ill Ll.'i \\'Om~ ,~1. . .;"AlliJIOIIIgh Ih~proc.i:;;cmb of ! IJu!,:..c gr.:mudD; an:: ,.,till·lx~il1g ~~'Ol'k,.-clout, it is t'klLf tl1~l1Illl'Y m'l~ ;nlillllnd~ :N'Ot'JlIi[lU Yo i811 giJt1l1lliu~ r~~~ ,~~ ~Iril>rru.. iUi..:IL.i.li~1!J. Illoo:l ~ilII1i..;nh'" :in 1!l1;:r~iifI'l:"eury gmn l'db. ~ indur.tiw hfgJt:~S (3)!. ~Vh~LI e:> iLlo~"m gI"Jmlle:; pl'a}'! St~l~~C:S .in M Ill,;:. ~~ld \'\!Ol11l.~ ~ .. tfl!lL a jlnm;!I)' role ~ irl 11fI~llJI in,l; RNA. nCftn J;;Iml~!~ i!1I: gidli!1 RNAiM'Xl p:rcdklcd R1\!j\-h~I'dillg r:toLct!..;. mid l1~tIYRNA:::. m~~lrn)tdi~. !.Ill! du\::i[' ~);j fical~' kJot!ilizl.'!.l[(~ (If 11I'('I!J).'1.cd ri'flil] l~wrfj;Lnkln in ~1~~ gltmu.lk"", 1/61. 11L'C: h:lti;;nk1il J!,IC!illl ~Imjl!!' (1\!-'1I" I1UC~ pt,."!Il.'S 1'IoO$j[ilJn~ UCIU to "lli.c'miiucr tl'l frt.">:As. cllinl1ig dtcit Il'iltCiptlit (mm tYoc welC\lS [I.m:] .rlllY oonirilllilc to

3_ I[ 1'fa.!'iI~hi~, '"' (1lI1I'i'adl! s

Sdm,1!' ~Ol~. ~~. ~M(l''f.I,

.c. ~~nltl!,

l1if}t. no
~

StnJ:;a1

Lnp~ lIt A_ ~uro!ll~

CLS;i!ijtI~~. 111. ,Ii. 1il1l1lf1\ DI!!.i£J'~I!'.r.!r. U'l'I!, llU {l'l~ J}. T.Itt a!~~IL ~. """ ~~Oell! ~1, 'l'<:i~ ~~n. 1. iii. 'Go M;:ifliJi~Q., ~. S. "IMYlar. I.. aQn~, R. Uhm~mn, Qm; ~ 14, l!S91.2004). II. fi. ID!tshihlild~ ~. C~lIialJli" T. ffl. fulIks., A. Il'. P~ilanilM>
(i, F.lh~n[l. I,IL ~rb\H:Ii:;. P.

s,

<:i. !j, ~~~ltl:,

R. t IB!.Ji !fl]. CefU;~1•.891 {2006l.

,~'~"i~,.

$tiIlOedL ~~

.!)ii!1l'. 122, 6ot~

MOOSI.

1:2. R. (_i)5!i, ~.lJ!ihlmi'!, I. It ~e%S~~,~U. es.l ~200~. 13. y~U~l'Ial"iI~~ ~ aI.. CflI :!!·n.~l(200z},
14_ It Wang ,et at llI\Npe iII~,~,~'~3, ~l!OO!iJ. l'i[ Takasiitii~ ~i!_. lllwe/.u;;!lmElfl 134, '51 (200'l'). l,6.- IN_T.ai!lrDl, iii. ~ Li~s!Jitz;. J'h¥ Dyn. :!~:~, 9;1 tlOO~l 5 1, _ ~ rot p,jn, Jl A..5dIllia, lilt I~~ 1)",,_ fill{ 2:1'. )15 (:!OOOl'16- 0- !bclMo~. A. Ephn1s~j, Nat,r.me' 42ii!, ~S'JI ~O;04)_ 19'_ P_ Il !loag., .1'1_ IN'abmllra, 1_ It !I!al'hl.ell. D.!Tffl0jllJl\l!'1'J' 13i!~ ·4'N 5 ~DD§~. 20. M. til. Un. ')' J. Foii1, \\1'. '5;. M!lW, [ .II. {iMu. .Urn (tn ltl'. 6:llIll:i!llall~ .. ,2'l. It M~~IlI1~,"" ft .,Itm!};f!(.il, ~. !li!J!~ S. fi:~I'!!'.fl!,

9. ,..•. i\~~"-i;l!llfl;l\r. I'll., Yall]~3!, Il NmnwJ~,. ,1':: .. H~R'J'1l', ~ K:D~ilI!itii, iIIflt Ce!18Jfl.1. 1!, olin n!l99]'. M~il1t i.iI. tia:ril!!1~ S.. ~<BIlt WG<: iIlr1fi. Mad. Sd. U:.SA lOll. 10;13;8 ~lOO¢). neE. 5dE~e'f. G; 1il~1}p.!'1I:1~,,'. I}, Srherlli. W. '6 .. roelli)'.. , .~" C~ll 5./.41 {OlOO3j.

my.

or

lflso'matic

(ens.

OOiml'!~ ,of mR:."'f,l\.

in

1rtTlI:l,')lffiifm ~1l1d.iooa l'il1lLRlll whlld II: !,;YI(1I}~~~"l:n :, MJ Wil!1- :JiG~. WLIllClgli.ioll1y, "

~r~!:f.!~

!;!,l!~l~gr.iIllq~I.."I,n;i}' b.:1:J.:~li..xl ~

r ix_Jdi~. TII~ hil·

NOS-I :md NOS-l Ilh~11 Ili~, I !3Ko.Il1.l.:l ill ~v!'.C'r1t frl}HI l!u'hfyl:mi.':' !,!etJll] tdl» fmllll t~.[![illL~k~y oornl_ Thi~.Ol\,,(:1K" raIL~mrc- Pgc ~ :1m ;[Il1dNrwln\ (7. U HFi!l. 2, B tmil H T["dI"l~cfiJ"
IIOJlII!J!O!;','"

H."J' hm·~ h~"I.'II..I.u:mrn in /'""a.,,1 and rmwfllllllllJiml

it.ip.:Il~ ~UI .nR:r'lA. OOl.~iY. RNA i.llwr~ Ir.Ul:.o:!~jUH 1l1hih~lion by l1lirmRNI\.s .. MmlY P.!IJOdly I}JJlII,)UnS tauJld In !!,CmlI ~rdnU~Cl'i lire
tiJ pml
rl;rt"J]~.

ooH,..

,md

iii. Llfl, Cloift. ~!g' .. l~, .1flllJll{lOOllli. :1,3. !lit ~i~nk Uilr oil1!NJ!lIM!~ ro~e!I!:UmOM. WillI1 ~n U.·~ JarMtlrt iJ ii;lp.~l~dllly 111~rfj!:H. {I~d wOlk in El.L·1i 1B~1'f ~ ~1l:p:il>lii'I~tI llir rUH Mil tli!J 1i11~'M,d b~ Hugl\e~,~ef!lic"'l In~1iUUi!'. ID.n~~em~. :iL1A'D~ 6

,2·;:!.II. iI. Meg~~,.

~li:~~ nn (~OO~~, I). N. CII)l, Co ~I.

WWl.v.. de ne:emag ..org 5

SCI ENCiE

VOL 31i6

.2.0 APRil 2007

Germ Cells
piece l[)g.C1h~' ;"P~i liC'l!~ion. O!ii:gifll,of PG(
thl" key HlCU'l"\ tlmt k';lld ~o rae
Pnl!WI~5Qr5 he m ~:~@ f'llulripQt@nll. Poo~i!ill'~1 ~pilbtaiStt(~:n.s; The l~lur~l"'ltcnt rrO\'!i!~I~J. c~ib~~:.t(It!lb ~~ptllU.~ to $i~IlID.~~ rft1n~the' c'l<Jn'!lc:!nhr~L1nic ti~)i~c$ ,ornd

l'!ICgin 1.0 <!'xpn-..'l;' frogIIM',jllij.;~ as thcJ' ~t:q !.IlI'e

celli: ~fi, mke 'eKe urs relia l;~ely~O/l;ein ElRi1~r}loil1ic clevelo;pment I,li~ 'initiated b~ signta15 that indLlce ,expressl0fll o~ BI,impl. a ~~. ftE!~lItal(H o'f the g:em1 HH, 'in areIII' 'epib!last (ells af ea rl~ ~05:[ijiIIIpllalnlt.atio:n, e<lIiib r:)IIos, Blnm pi re~m1!~~~!~ lI"~ill(i~iellt sornallic:: iii NIQ~,amin
5peciricaIDl'ClIII 01
{Ite'm1

the nhili,!y

[0 fOm.1 r~(i('s,

,:]1!Imugh 0111)' a '~m[llill

.as \l,Ie~1.as

llu15li! cells and IlrcHiIlol~ IProgre~'!lil:!ln~ClL'lart.l: ~erm ~~n t~U! fa!:!," IBlil!f1lpl may also ~,'al~e.aI ro1le in lhe mai nten'a l1:()e®f ea rly ~e rrn ce~~chilira,c!eristks by 'emu rillglU, ei~ e.scape 'rmrn tile 'f>Cll!flalk fate
p051>iblte

reversion

to

phuip

Ollill1tste mI

cells.

rilllOl'dial !Jicml ('d~;:, (PGCS). 11LcfOllntl:Oj' . 'i,;~lisnf'thc g~rm I..~d~ 1I,1IC<j.![!;C,Ort:'L~I~li!y . .!.1SllItJ:.1 i IK'd ~-.Jrl)i dmill!!l 'I.'1IlI1D:rycmit" des ·\,dopmu.::rl'L. SIX'lCil'lcati{])1l ~r P(;C':;;C11n oceur ..:ith~ir tnltlulIy&i 111,;rnh(:rj!~n~~ o.f ~nn cdi tk:t..:rminal1l;:" :1Ir..~ld!r j'1:[t":.~m in Ih~ IJ~. :I~ in CWlwf/wr~JiU,~ d~~lm_" ~nd, DnatlpMIII, or in rc":qlnI1S~ 10 inducrive s;~gn.~L~, il'l mice and ,m!'; rroh.-rhly aU rmnumls. ln .i!lsuu~C\.'$h!J\~'C\'e Ii, ~'oCnn cells arc u'lai j;1!,~i!1cd, 111'(,-..;:hlUl I~}~ L~n'L~ ~h~~ 1/!~C'~'J;I~t them frni~l ~l,m~~llti:!1l,iflg il1h1

'p'.

t..:!.nI g':'llcrnH:

an inJnllil.c auraber

(J rlrGCs

whclII

n::ILII"UI~dw I i:Ki M~'lU(;y~1 or when IJJ'0' etherw iSl: roc..ivc :oi!)~'(]i ~i!lnub; ttl irtdLK"C g";.IT11 tell Ilc rill~'_ \lI!,l·could therdm:c: 1t'1IH thisjhe :;,~m cdl rntl!lJd f[.'i:r I~G(, ~pcci I:leal.imn.

minL1'ri!yiJf~hC'm become g.c!1~l cdl~ in~hC'CiI'!d V. 4) (Fig.. II. V'.'i,lhill Iln.. Inl.gj/.i,~-I)Oiiilh'c "ii'I,; cdh;. m C'l1;ihryohl.'!.: ,00-)' 6.15 ~E6 ,1 $~. IiOOiJ~, :;i::". cd]!> ~UI 'Lh~ plt!~!.'C'lil~!: pu...h.:rim' pro;o<.iL1iiil :;11,.;: tin ~i1: ..:rllll:r.f)'il aniJl \lirel! Lly in l~l.lrm~ ~ W ~~Il ~.hi: {l\ ~riytillg hE begin 10 5how I!'Npl'CSSDOn 'If BHmp1 IPnlmI. 'Exp,,;tinll,mb It.tting gcn~lit: limmt,'¢ t!l,;]iIOI15U-m:t..: ~jhm1111 of Iht; fJlimpi!J1'i.pr'Iu"Stii:ngcell ... hl~ mig~ll,nc in 'Lilliproxirrml1.lIO~I.. 1ior cp'ibb~1 Oll.':~:h(~ ~i .i I:It~"gC-Ll;;~1 ctlu-tl !lGC [[
rm,,"'CLIIl>OTc-dls ~3i, 'n,i;l'111: i~ !"urth,.;r accret i'L:m'C'! r a.rir[lJ~l-I}L:!liili'\II..~

,~n

1\11d:I~:mr.u",· 'an~ "Fl'llmllli r~rmn n LI.liut fl'( i(_' !"l'Iedfie.moll in !11 llit'q;~ klC ,~n~~ Ihem fmllil n CfII1timlbng dr:m: lmvm·d a s,nmnttk: liHC. ;;l!~dIII i:5 is ,coulPkd w i(h ;0 ch t\(llla,Ui a-based m~'C~la!1ism~hm CJ·~~ dl.i~ !!'C~ld.~hL.'J'L.'h~~~~lh.fC~lIl~~1".~

cdl::, a~tb't ~hi':; I.im~. Lillu..\ll~C-lml.·ill~ b"xpcrin'!e~l~~Ill,d Pl'\!.'\'·i.~lU,"~}' shmvn 111m )1f.1<n1'" s;i~~c cdls i111:hc fWO",i m:~ I .;:opibl.llM 'It 1:6_5 could gh'c

'~rI!1 ...

ntie «lis. M1:ce

()J'm!tJp/!llt1, fllUrldcof l~( i()_~

.~i!.l!w

The Stem Cf?II~M.(Idei~ ~tII~ ~GiC

:S~edUc~t~illrl ijif! I!! C
t!1'!!;'ftjjl,\ !Ul~

r some ph,~riJ'l!Clite]JK:}'A~ 11LeSOllif!o;! lime,'G{'~ mU)~t (lcc;qnire ~.ml!'l1,~inlmin Ihcir li!l~a~f'H.'eiik ehar!!lg
~!l1'9:.\1JlJ\,~~~"1l ..

genes,

"'~.Y

rise toilodl POC~ and C:":Ir.l'cI~lbn'{HlOC !'I~C'!H'idcrmlnu never cxc'I.I:i. sh"(:~)' 'Ki f'li(:s (frl,. F ~on~lCrmore, dL~~;I! p!bl",~ ~-c!I/:;from ~'",,:;; C'mbry(l~ e
wh..:rl li<I!l"'J:IJ~nloCii 10' ~hc 11m1\. ~mal.-g'(I;.'lterifltl"

regl,on can

CtLlfitriimlUI! !ei

Ihe

~';!l'iilll IX

n lilh!<lge.

'l'C~,i!>ti,,;)1., IWc;;."nI 8(1\.~~1~l> :1.~ b';!gi Iming

[f.l

~\lh~~I",'<!!.~ W1.I)I].'I.imr~I-«'!;!;.slcrior ells I~!~U:'I~b~iIl~I",'d e

U)" IiOW'b"\!O:[. i,o rniee; .... poxi I'iC~liL:l:I] 0 f PGf!ol b; d.!.'llcm-d umM aner inl,phllJllll.iQI1. ofbll:JSH!lCY:'~5_ The C':"'_~e !J.utlll!yo,Uliccel.Ocl~nTl ~.[;'1E ~ :JiJ;Jd ... :i;M.'oru~ cmlodcrm (V E). will i,cjHitl mnmd 1J]c cp'ib~<!i'll1d b. of

aside at the outset InliVl a trniVOlclDI Zy~(IoN;;and prevclnctl fn:HQl di:lr~rctl~i~~in,g imo ::>OIfl1;l~ ic: eells bJ' rcpn;$$ U('I<II 0 I:· I he gl('l~o.~
SCi

are

PWlo!!:!iPllil.!

tm:l1:.~r,ip~illrml machi nl:ry

ifiilll~IBS1: ~ -

'"

".

DVE

r
Compelence
1"[;.

~r~ignilis dm~. [1~:lllmCI ,~ .snml,~ [luUIlbl!J.· okpih1a.~t cells 1.171 become l'l(!if's> !.h~res~orlh~
MlIIIUl::t"
iCCU~

~h~

[lI!Jt;~illlp.imnaiion

cg~

C'yliru:kr. 11m Ih",

oomr.nb"I1cr: di mtN~1Il1 ~atil:itJ1 ~nto ",Ol:llialie

~L~llll:o..

diviuing 11ilOLIf;C ~pibi3,~~ cI:!'IL" Iy equi "<I.km 10 tilt! Dlm.l1:pJs. ilfl '::M (2~,llow(;\ er, whereas ill lJm.~'lrpirUfj., ~hC',c:lcl!:fl1~inmltS (1:1: S\lJI11 tic audgenn ~H s ::In: ~ 1l1~,C<!:-dy $'\;g~w-~~¢d ill spec I :Ije P:J,!lflJl$ ~lr I he
:P[C~ JevdlJli)lJlt"lll:d
'l~ll!!,;yh~"iltJ;$lll.'h

Tim mpidlY'

d~l>Ctmin;ltJ.lb~~,bl in

the meuse

'i.lIX'Y~C'.rlifth~r~Ii!lfC. O("s ~lrl,l!i rlilUIi: fi'lli11 Ihe P 'rn~), i~lHIIr'1I~! r:iji'lo!com CW1~hl:I!."1 C'ell'~ Lh:U !U"C' n 1~!.f;!iI}' tmli,';JCri('l1iUI1.dl}' aclh e a n!l ~('J,~oll'le C.\liC'ut hzli\'C' !!ll1 ba[k~,j li~OIl (1 SJOJu:uic f!lIC, F~yrlhcnnorc. plmi(Xli.cm c'nlhryonic stem eells,.

which call b\:' flrop:al!;m:d hlll:e:11i1li~.ely in vhro"

'M!rI{(ll1l~ irlllit C.!lmi[~ R~~rclllJ~ ~d!lll ImsHl~f, U~,'!ill'lsil1l' of (.;j1Tii~l'IiJ1g~. TMni:~ Coo It R.OilC, 'CamtIl1d~ (82

,stDNa .8l1mpt

"'Ge~

1Or4, OK. "r·e "",ll0in (j!f~e.~J:liiiclM~ :r./ItiJoillld

.ta~:10021@mOOe<_bI1I.GilIfUl~,i!l~

IJ_~

aiXdrel'lerL ~·mlli~

IF~g. I.. II),fVfi.o~mfnt or early p ootll'l'lj)la l'I'~atioDlemb ryo ~f'(lm E5.0 ~o E,],5" l'ii,epicl;ilIgUle formation O'~ PGCs" 1tN i'1f@;J;1fTt.!!!1. 'e'pibta~t ,c~ll'S re~~~lliIw :si9~l1;h,t;!!ir! tli!~ ~f~~rn~ry@!i!~~tii~'!'lIl;',s., whf,dl ill@!!!~e il1XPIll'~l>iOIll of JtogPlisinthe eJllI:iLa~t, antil 01 Blimpl li'l the PGC precursor ceililsaJt Olil'~ en" of lheshoi'iil a~s. 1:1i;'~Ii:U~ ~5~1I,ilal"i~n. g ,A:~t~'r g<;l5itrulafkm, lh ~ PGC pliE!l;I;Ir~rs I.o(ii~~ 1'Qth'~ pO:S~l;'ricn p~o~illT1~,I. r~i(J:nI, wl1ere~h,ey Ilmd:er~() ~p~di firflitnon to, [ofm 1iIl1e-f,Olii1[Jer po pulatiofi ,of .:5'/fl!'ra·pli)~ib!,ieIP'GCs..

394

SPECIAILSECTION
embryos, which disordeu .I\.~[!.rtxis [m.l1!1:lti.m Dc111illed :1I1:IJ1~L~is:;;u,ggcs~~ ~hal. the IJ.!'G{·· and rc:;;~iI~ ~Iil ~hc ~1i: 1~I:C;i:o;iQn0 f ··I'IO.:\I~r~ar~." C:OI.'I'IfI~'f1C'll'.pm~.lIMa~ CiI)~hJ<1Islcc~l~ c;qlfC'!\.:;;:ill,~ ind iC-oI1C•.~C' pcl)1~:;:lll:nce :iii:gn;p~~~h<lt call ~Cll.t\'" inchl~~~I'I!l ,'\'n!.t(~l<l!U1~ Fti'rf [n~hc entire li'agil,_"J/j.tul'; arc illitilll.i.y dc~ti!!\!>d ror a ~~iMi;L~. sJl()W b1i~IlY cx1.«Ipk C~l~l~~ ()rr{i('~ cWl'11:;i!k flue. "'\'~iXIH!ill!llb'T Q~.r!y .fllfmp/(:(jl1liutJ:!! lIy i1l{II.ICC fl'!1'1<",im:lkpo..~1~r~of ~Il!hbsl il;;C~ I~;HJ(."~~~ !.lIptp (:;()5 i{D ,"Orn~l! it Eo l~l~ pn(' c\\l;r~'l':",ill~ cell:., ;~.DEiS.7$ or~g~~"J~bl~ in the I M. l~J). h mpil-CIU'S Ih!!u .w (l1'CllC~trotjon 0r !ilffi\\ Ih f~CHH'l>, wh ieh mny incl~!.1e N{~~~[J.~ aud 1~~Ll'Xi!1l,11c ~~ibi:lm ~'rC1I~~xhi~)il (l:!>I~~Cl:l). fif ~m ~11!c. Con.~i~te!llwith ~hC':.~ ~ng.~.he jl!,Ulllind t fg nt crC'~1(."~i!lU iWmn'l::J1l forPCnC !lre'~~l\ H''l.;\'/l J as wc'U 1L;~ other I1lC,.;.odenmJ, gene«, ber of Brlli11''/ -(."~IH~S iHg (~I L~i!I~C"rc'a),;c~froll! about 6 to H:, b;!1W~!1 ECl.25 nlltlFA5 (.'i), CU!'SCl!)1 1!l1 Iw:. :-1r:g~~~cl.i fr{l'lu ~O!!K!.t~c l.id I iudlilJing r: rgltl.. and SJwU 12' nl~~ g\!,!l~:-:: IlinC'Jgc~. T!!ilkinJil in~n acc.omll Ihc- pnss;~hilhy~lllll Ihcrc h COin~iaU,le'to he ~llr'-I'Cg~ll~alcd in the: nCii1lhhtll'il1~ me5oo!Crmai sometie tissues, Ul1'1.\'i!w,;:r,thcy Icn~lbcni!'l~ of the cell cycle ti~l]e if! llite" roc 'tJ;;.:!I;il.Wnr.: rc,IJrr..~ ... iJl 11'11:Biimpi"p.tJ::;iti,vl.; ,] (..1,;U:\ IJ~ 1IIP.'~lJ~ IH1{n ., hou L1S '[Q 'l~b~l!l1t ~(! HIO!lIP';;" .Biijm:li!l:;j',nc IK€)' RC!JIU~~~Qr ()f ~P'G(.Sj)@d~i(~ti(l:rl 'rr~mlilUncni of lidIll~titJnill pn::CIllI~c;. i:- ncecsin :IIi on"k:dy manner alollg W~11:i th!:pmg.n::~. A crucial pm~ POC' ::,p(:cili~llion in i:[mn~ ~i:ol; {i r l?GC slJ'C=~ifit:;11 ~t!t:I j MIl)' ~:!J 11!.'!::mmt lor t1ht'1ut -10 S'[cl:I.:J·flrl~ili ... ~ TI1!.:: r~'PI1."li::;i()J1 Ibllll~kr PGC:. Dli::l'I,:Irl,! I,imdly o[I:'i!'it1.!;)d :11. I]'ll.:' mood o.r~tlllit-illh in..::huJ!.'S !\.l1!Xil\::,iO:I1 or '[hi: uf :;~rlllti~ ~~II1.C~ in IflGCs ~ ctllll~i:>l(!l1~ wilh son1iklic· prugumn. 1111 nnee, u l[[liqU~ g,-ml1 ~~dl 11111": ~lhl,mmrt~'ill,!:JII] of repreS);~!jrLl or Ih~smlllilic pUi>~~rrim cull of lin: ~~rin1iti ...Cl ,:.~n.!:'Ik.m E7'.25 ISpl:ci fie 'iTw]~lTllpi~iurrmllJ.(_1\\'tltk :iCI;."HmJ; H.1 ru!ll!ll:.uc 0.:'1. ;~l, IJwgrr1l1.1 observed in C dl.';[l!fmS and in ffi(' speci liC";uKm" E:lIICu:;i,'c 'lI1f~y~il-;or gcul.: Dl'O,frJr:;ilila .. c~pn.:~j;il[}n proru~1;; in iioin!J-ll:tdj~ iiJI!"W~ I.he O:JlIflk:d \\ jl~l ahc rcpn "];:>lml ofU:lCl.Itx:ilCn1.1l.!1.. or~, :molg;:L1lm: pmg,mm during !l1fXiCi'l.i,,~ .!,!!.:rlCl>, Ih(:rc i:; IL~)-I'J;;,!:l'"nul i1JU U r ~H~K"r BOlhl the E ....E. and VE ~I.l: ,",,~i:11~j~ Ih~ III';- illvclwlIl,.;rl~ I, War gcsm cdI5pt.:cilli:'1lI.itm (3. 5. 21 •. !J)[Fig.2~ 1~'11~, ifldudil]~ Sad l5. 1.?;. .,I\.Lmtilcr _gcm::. quil$itiml or comG)~rlOC 1Iind ~GC pn.':CursrJ'rs, NWl(~R. I~ aL'i.O 11."C:i\:pn::-:!lcd I~GC, {J4~. Thus, i.n 'DIJI nUl. Ibr 1'(1(; ~pt.:cirit:31.iml ib~lIl' ~9. WI, AinOJ'!S 11,.: !1fm::;id:Cllllifkd ~o I:hr, Blimp ~ pml..:irl iJ~ a kc-y lr.uul~riipL1QIl~:I n:~LJh~tI:r d,:.n i:;, amtm~ all 11h~ lin~3~':i l:lIm J!I.~\.\."iajJ frmri th!.:': Bmh! mt~lf'!h~edH.1il.!' I,w'wi'll 4 (BM.IP~,). whK:h ~~ is, l~m~IL~C\'d t~y !:xE, i,s ~~fl~b.l.(." Dr i~ul~ldn.:J: j)oolv ~p['l!'l;~il~.li!!' for :1'C(lflC:'ising dl~ som~l.h;, cpH~I;t;<;,t k on b' germ eel fu;, rcg'li~ (.tWl'CssiOll f",.g_ili.,'lJ(i,tI1lj ,c:!;prc:.;~ion (3)" f;.xE n nd VI: 1II"C luogrnl'l:l in r){ iU, whb~e ;~ll,mvin,g C:'lllllhihhl11~l'It or fillI1'ipme~lcy-ml.'loci(lted ~C'I]CS du ri tlJ-l: !he ;[I1,~o d~c sources Df6MP:-;b and ]:MYil'2.. ot' g_.;:orllll cell d~amcl!er iF! IhcsC' cells l5. 231. C0l<1 ~ thc:ir !It'Cd j"~ca'ion. TI,.c re is nlso exdnrnahl. ;~ ri;<l}CC'~hcly.Loss of ,iii!!)' !hc:;~ l1.i~!lalitlg U.~~mpl pm1cin 1'!1!l:;;~~l'lvsIT .mo,reCtllles ;lrro',W!tl:s ~h~ (;'O!nlX'l<:'r!~ tfl give rro!i!'le-rkh regklfl. fr.·c {'il! ~l.~nc !l~~cn;. <l!tnl .tcklk di}!if!,~i~l. rise to ;til Ilf mml or ~hc [-'{1 {':!> U i J J I. 13MI~.~ ,. ('·-tcrmi~I:~1 ·tf.igg~r~ril~ fil~~~I~&I~~i)'lmi1i,'i" ("if ~hc l:nlnMIl!ccr SmaJ I 51?:. wll iel1 tr.m~J~lc;lltl'~in~(.Ihe nue I<L'L!~~\ il.ll I th~ Crln'l!~'bf}ll n~cdia.or, SI11:ld4_ E3,6 Loss. of 5ll'n~Lm~IJ Smad5 (buj FlOI SmJd~) causes severc rc9' • duetion illl 11mmnnbers of P'GC~ U4 16) • .:1$ does I.h~ ~:unditiCln:.il lo:s~ MS:lllItU~ un. T!! ~ l:f1"/~-Smad ~~Il~ dos:J!lll.! + + + i:;; criueul rOf I?GC specitlc n +~ SUx2 + +1~
tl) ~hc disml rcgi~m of the l~piblaM ,"IBnl ml.l:)! gi,vc f1_W ~Q _~mn.:lljc .;;dls lil~ lhC'S1l: :l,ntJ:ii:'_~

or

or

J1"

or

or

e
+ +

tiou.For

examptc,

in the fhJ11'.J'·

o r I~GC~ is mhm;I hah'e,d. whlch

lilctcm:ZY!,"DLL'i llmliOJllA.:>.11'1eImmoor

..NlIDOfJ "" "''''''''''+... ,.....,.., ..... ,.,,.. .,.,,.,"" .......

8ffmp~

Dtib:.n

:lC'Lwvmcd Wig. 1.1::kle (fPIlL'!!

~h"i

double' II~tltmZ)I!:_l.oul'l SMW~lJ i~l1d Smlld5 OS). indLocd liw .P' l{' I~m.:ur.:;ol'l; \1 r,IK~r!l1J rm!lll Ih~ mm:;~ rm}:\ iLll al I:liy~;r' 'Df ~~II! e~lihla;;t. Wht'l~ !lIc' I3MP-Sfll~d $i,gn~liglg i~ slmtgly
:1.1:;0
C'::I..~

is

fire

il1l]t~

81610 !¥8f1INs Na'flOSlJ

+/~
"l'UF

+SCF +tlFGF

TIt>;; dcllolQ~m ·(If lJIillu~t1m~th'c PGl "pM"i n;Of::;,~t IlliC j1QSI,mlJlf

';l'1if1}

+1+1· '?
fneeo.xpr>E!!I5oo depoal'HJlg. 1l<ii~I!I;,~!Oi!$'or ~
DaJliIlIB

cmb~l! iildi'"t"';~
(A,-P,

till.:' .um.crilli'11i1},!'>liCriM

a:\i~~tlnn:ltloH

may r.l~.lt' iii JX,h:

in ci'clcrmirlD:lg Uldr n iJil1bCll1' ,[lml

l(le.~n.iol1. The: ~Ilt~ rio!" \'1Sl.:Cml cilooucml {i\Vtl l!~l r'lr()du·ce~ N 001]<11 r~IL"1 "Vr~1'l1n~g~mi:li~:s,II m~
~~~I1J~til~gNr411 il.l1d WI~8Si~l:ul-

FJrEillmll1<lJiY Fi~l.

IF"*"

Dmlt88

iug

'III

nhc POShl'rior siclt·

[j

Ih~

1ClI1:1~~'o., IIllt~ 5IL!J ~\;I ~'11l

'po:!<.irt~~J l IJI!OClIP,;OI'5 !Ire d... !· tC't"lt'1d. NOlllMy. SmmN_!·nuIIHlrn

roc

11m IlIimp.J~

Fi'!:l. 2;. A ~lImmaJry of lPiliC !l:[)!lcif'icatioln. flog'resf>ive !l:h:a!n'!le~ il"! g~H~ el.~r~~iolil ·rumlllll~''~p'itDlast in~rI~' @mblJ®~ {gr!ll!lli!} ~(I IPG(~ {red~ a fe' fmli(a~ed. E.p'igenetk dllHerelllres betlllfel'li PG(~ ~rld ~@m~til; Q\ll~ (~l'tffl) !lJr'!l ~1h9M1 olll E:U:)S. NfJ'(!"r)53, Mvh" Dmt, and !)azl are alls.o germ (ell.-eriu'Ldlil~11I genf.s., I)lNA

rnu!lh~lni:~n ~S,",~'E!qisrased e

in ilmprilltil1Q

c}olllulll.

,el~e<I!I1EmI5

a iild 9~me'e-rlli:~diflg re<girOrls, <lr'~eJ·~JlO,5 03", WW!,'I,.sde ne:emag., org SCI ENCiE VOL 31i6

,oomil1e~ ..

3. (A) P~~ential roie ,00 IJ!lil'1'\p1flPrrm:s, compl1'x il'l P6C ~~(Ui(atuen !,lniliil~~~m~g~.ti(l" of PG(s ~.nte 'lile g;enital. ridge-$, ~~helillihe !(II ~~le\( b'olnS lm:a~e5 JrQi!T! tlillll' r1wdtlJ!; wl~ ~wpl,um. (100TJiJe tQ~S. 1;)1 Illfmpi m1il1 rli'I!j d&lul~er,ei'llia,lkill of P(j(~illll:i ~tlllrilXlt~t EGlI;€lts.l)hdS, i's ,;3 tn!l~,t r eli mp]UlfrmtS, Q

,2.0 APRil 2007

395

Germ Cells
I~n:'~"if.lil~ of cllher. mliqrLt~ :!;,'Cn~~in 'NIC~, incll;IlUUg ['1'lim N. ~flCllU' lh~1 i~clo!'icly !I'C~a!Cld ~~

m Ih~

liml;

()fPfi('

_<;,'Ili..~ifi.cati(m(.?8), eomrib-

,/lilmpi1i'ni.lm!. wMch I'!II~Y 'Ilso ha','~' role h~ !)I(' :';iPo;;'rl;:1!~Pl~~'i1 ~!J),

rUDl(::tlo~1;~1 imllGr!~nC'e lll,imp~ in PGC )ipl.1dliCli,~b:m l~~'JI;J)!H~lC ~,.'~d~tH ~ILI!'i!1~ Ibc Hle
'Hl'I!~'

,I

l~!t:~oPGC ::l('lecifiC(;!lioll

in

COI!jllnu;:l~p.!1

will~

I"r~n.S ,ewIII iHnp ~t W'111.1!;)

COIll111:CK.

rh~mlc('lfPrn1l5 in 1ltCIUf.lljl~!~(]('s n::lutlin!ii I(l! be r~! cll:ld,d~~!c1:I. h ,dl~D. ~I1l;'i. thmb3lllU('Il. Iy S

pm!l~IiJ..1y h~!p,,, 10 diro~lP1Flllj15!o~I~';I!'Jr:t~, ~~Id~ f]JuJ.V., l}u~ a !'nol~ Cfll11.!lrcI1c!'I;'ih'c ,!l~ sc,~lI'Ch ror mJ;!~rt'IlL'g,rct... b, l1.clXkd l~~ un ~\ c~ its role Iill0f~ rully in c~rl}' PG{"s, Fl!!mhiJf ~!we..,I..i.gation:-; wil] dcepC'f1 fmr ~n)1ight~ rI!! ~hc :-lpocci.lk:niml of gcnn L'c11I:;;, lh~ l1l!hl crilic~ r

lit:;; im~grjty

Ihl.'rO\lflh IhoC' im'Ll,h'Cl'IlC11t nf'Tudnr,

or

:Po.s:~P@(irWj(:(llliQr!I~~!1~bU:!lhlm~:!lt of :P6'( ~.pigi~I1I~~:k S,igniJilu m~

sls

Llf

the- b'lim/~I-nw~,~mITrl(lll~I.:' cm-

hryp~, \Alhich res~!~!!<in ahc rmn! dc'\,'c ~r'Pm(,']]! or !ol.ludcr Fji(lCs., ~f1 Ihc- ;lhSiCncC' of Blimpl, the mn:II:Hltce~islllm~ tj,!;!htpnC-Jike clus~ef. ~I Inll Ih,.;y '1.;IJSC II~) pru:liCre:rnh; ~md Iln;y ~h~l'W Illih.: ~" i(C~I~t:'r:or nrignniml ollil ,ofti1cd,u:.;.kr, f They ,S 1~(1:>Ji,uw inC'LlIli!l>j;:;; t ....lTllli.:pl1.~i>ion (11 r ,lJfJxb} , whii:.Il i~ II h:J~ln~nrk of IPGC SW1i!ciliC:.nilUb. '~\hi I..: fila lilll! to ~hIJW c'Oll:siiSLI!I.'llU FI,[1J!lLllalion of' .!i{dla urul So:.:2. :i!:. ovscf~'Cd in llonrmlPGC. Thus, T:Hi I'll ~H phliy$ n c:ri~ico~ ro~.!.: in I.he -cst<!bli:;.'hml.:l1l ef the n.lLlI1dc:r PGC;ro. The Eliolle o,f PIFllillt5Al1!Iil1lilil~ ,Meth.,."la:'l!e in PGCSpedllicalliolil R~ccut s!~tliC!S hCl"'~ liI1J)wr! ,I rl'C'\!i~lu~~ly uni:C1cogni:l.<!'(lBHIll['lI
I~~111,15CflllIlpkx

An imcgYlll! I~,~nof ~hc PHC ~!'l>\:C Dh...n~LjI'1 i I)!TI!,,);!,""''' l!ldlild(~ i'1~11~~~mi.d rCp.i.gc~lc!.k !llO~Ul]cmi.nlls. which occur in the s.rdhl-l1osilh'~ Itoes. i\IFJUl ijhclevel rt11'IHK9u1C2 l;IJtl cp~g1,;IJi.:li!.:'Iii urk ~1,};~{!ICi~l.,;d~'~Ih Ifllii':.<ol:ri~lIi{)mil " I'l,;p:OC- ...... ~rJn I d illilirli:;]l!:~" wh";l:Jr:a;,> 1.1.3K17tth.:3 (mjd~h !.,(If!:.'(Itt..'>~iv!: ~p:i~l~i,ClLi~ til,l1IrK11~~(II;;.i~~I.~\d "" illl ihiJ;h .h.."'i' dt-: Eilz2 j b~COl]ll':5 'Pm:111I ith:m I 29gFi!b\. 11, These dltlllJP..>·Ur!.:" lolluwl.'lIlJ} 1 upn:_tulmiullIor r.BK.;[I]I~lf3{Fig.l)"Ut: r..1)~g~· 11~tic nmiks IIJKHmd mitl n.:nC·111'11(:11.3 lire ItULldJ'lc.I~, Ih~ rhcuilllli'O'\': 1J\ilrt~of'g_.rtm; ~ilJci t!llut 111:1: rqln..";;;I.'tI 11111 pl'Uripolr;;II'L 1.'111 bryonTr.: IOtCI'l1.

~i!1c~'C ln ,rLU speeles,

R.e!!e:ro!;f'!(eJ .and! ,rMO,leS. i!.. S" Sirolll1f; it. IleI1!m~m.. Scien~ )16, ~nU(lO;}, :2. ,. Iii. O'F<J~IL J.SA~!'!111I~ 1. T, 51.!, Cuff. INa.!. :l,Q, :1t~5
3_
~:!iI)l)4J,. PJj, Sa[!)j]w, S. (. 18milorn,. I.t II. ~raf]l, N~~418.. ,193

~}r

~:!:,oo.n. 4_ .~ S,. 'I'~maia. 'I. .l&il5wi..I..'!e.:h" .i!Jr!K 119 (osup~L It :5Zn.

~2·00.2t

eel I\:;"

'i '~"hm ij!~ el ,0.1" Natrnp.' ~36. ~i:l1 (;l~~O ti, tI... A. LrWSarl, W.J. H~~~. aM 1'il:Jl!.~t!: Symp.U2,

~!'I

Genu {eUs, <lll1id P[u rill!lllel1j'~. .stellil

r(~!~~5,~

ln gem]

trnns iOCHI.ionL],f nli Inp' ~ PrrntS from 1.I~1l 1 1 LI' dell:' to Ih~ cyl{llplll~lt1 at EIl.S. aner wh,idb t,hc' ~::;,;.pr~::.;~ion IJf I'Iluriptlb..mcy-.ll};.so~.i.:lwcl gl<JnC"~~dw hcgil]5l0 bC"lImHl.:r~guhl'LrJd. Thus. B lill]p] 1[11Tllt5clJlmpk:?< ..unay pl~.y all cMcmiu! rok in l1l~il] ....ini~lg the fGC Iint~1\!!l' d~Hil]~ 'tile' mig!.a:LII,ioll of the iCcHs illH.1 tht' gonm!l}i {Fig. JAI. NC:H:lihly, ~C;Cllt ~~.l!Idk~un D~vi;;(¥JljlJtI InJi· iCalJ.: I.bml~ ~I n'l1'l1t~~tl!.1nl j,n rilL:' .PitnitS hnl11alo~. ('IJP.~UI&!lIMl..wtj,a i].rcl'i ~t'lm l"rliI.I.... pt"Ci I'k;Ji~irill il1l f(]'lMI~lJ,Q':oi mu.:!. JOCO\'t'lopnllm {) f :.pt:.::l1l11l!lrIJIC)' I. ~t":S. ill nmks ~2'j. ~~J.l .B!)th tl.lt'! r0n11l'Llio]} ol"m:ng!J in. ~llll'$'C' c>C'l'b ~Ltld pole ~'~~sm,inacgril}:.'Irc af· , I:CCL'Ca In Clql,'iliu:'l·tldw75 I1m~mUic;>. nq~'.ukl,;nf l!nri.1. haslhc l'I<otcmi;lll{5 m~lh}'t'nJ;; prolcin

cc~ ls, ~)mu5 ls an :ur~~fl~e-speeille hi:>.tone n mc~hyhron:s.Jbra s c, wh~C'h ~~tC\hlrC.~~yn1mclrirC~~ dime~~!ylminl1 fir ;l:rg~!'lifl~-3 111l M..l0nc U~A i1rldfbr il41 a i~ UI:!Ame2!':.ln410~nC'2:]>:}, ~\hichi,~ dc!C(~eJi~ go!rm celt. ~!n.A fc'l.\ Er(iec~b can ,r)n!y ~o,!. derived lh.m~PG("~ berween Ef-:5 .l.'!nJIE 11.5 \Aliterl 113K27nl~J. ~!!!I'.b'Ct~~}rthe- Blhm('lIJPm~l$COml}lex hll'l'!: {3 b",'(:nid:C'l~lilied ill gt'nl! ,cC'll~ ind~tdi ~l_.:!l/)hx.l~, amI 113K4~nC'1 nlUlrk.~b~(lII1~pr(lm~nCIH III P( i('s, DhxJH 'i.$ rcp~~scd ~IINI ll.hOW[l1~II~ j!~ldl ~~'I!c'tl 9mC'~ is lIb-~ ill 0'0. 3m- t\ t I 13K lime. l~liCs ~l(lr~v Cx.rrC:'l$~O'!l of H4RJlne2s C'pigerlc~k mur3\ until E. ~:!5. lIS the ;"~~m~ ~~III ri porcncy-assee lased gcues (Jet,'. :'irl,r!, expresslen iii lil:i:; time C'(lincWes with the
lIIuli Nml'(JU.

AIltif'!iersfllie Pil'lenotype 1'!(l{·s I.I!ldcrf,lo cicdilTcreu!1.Jlion ill'110 rlLlri. Ilotel1~ C'lUhfymli c; gel'lIl lEU l' cells \,'hc'lI Ihe)1 :1I·e c~~h~lrcd w~[h basie t1hrllhl<l:ingmwll,1 r<lc~m: tl~I"(Wt ~~tlk.;::milL iTlhibit~")I1' r~lCwr Cllrl ;!Ild ~~J;m cell racWr (Sen ,t.W 32!,

X. ll'lou, lM~, IJilIl. 118, 12.4 ~U5irJ~. So A. WllrlfM". K. If,, l~i'I,. ~1fmiNi(lrNm73, os
1. ~. P.fMfl,

~n!Ml ~~OOM,

9. 5;. M, dt!Soo~a l'tlt:!

~~~~,

{~' (11" G~~

fJ.t!,I"llit' 1836 :!,~~,

10- T. VOlhi'1Ii~1J, ,~. (I!iii~~~ y- ~~I~~ llM1'~~

,Bli~n;p~ is. down.;n:gulmt:lI dmllJg th~ dl:iri\'l~litm EG cells, \\>h~'re;b.'" nllll5 i:s, P (!~LI.:':r:h::d not tmly in. EG ,~e.~l.,but also in. ;:iH ol.h":fplm;ip[nC"lll ceH5, Ttm 100); tlf.HUimrpl nmy n'.~tl~'t i~~ tlrJ[1qll't'~:>im~ of (,l'Ol1..i.!inJ!l~mc:;; ll:.:u[ II'IlI<linA.l1I:in Ihe g('rnl cr,;]III,i:rLt'ag_t, ~I!t.h,....Df!.'I[3,~ {Fig_ 3B "." l:IIrg'L'it ofBiimp 1, .P'mll.:5 cmllpkx ~ 11 PGC:;-; un\~1 E 11.5 ~.:"1~, R.criproc:'dlly, cctapj;c cxpre~8ioll of:mi.rI~p I in p~l!r.1polclli cl:nbryoni!i:: t:am::inom:ll !Eelr c:cli~ k[l(9:. ti:l' 'the rc:pro;;~],[l:rl .f o D.lI,vj,'j,. h. '\.viH UI<1' or il1l\lre~L to ll.~lrL~1'i11l illL~ w hell.lll.I:IBm EC' l'C!~'~ il1.d~'I.!da] Ipll~riPOLc!lrt and ~wm llCIl:s. acqu i fJ.!' a."poc ~sof ~hU'.PGC dllil!FlIC'l('r lUrOI1 expression 131imp I.

or

oa.~ M. I!,\, M.ruuk, ,Mifcfr.. i!Je.; ~, ,61 (:!iOOU, 1'5_ Ii:, iffa!lil~hi a I'll;, MBrl'!. ,OBI', ns, ~9 {~maI2j," Ui, It. D. 1fr~'bla:f" N_ l.. i)ulIm, IE.. ~!!r:I1, ~ lM~i:!p.!I'!Mr 128,. 311m!! 1)2)0in:~, 11. G. C. (nlJ" N. R ~ni'!1 0, (. Al'id~MJI, L. O~~t1, [. ]. ltatlfflJrlfi, .l)l.M~~~ Ul, ,5tH. f.lOO'l}, l&. $.]. AA'iolil. :~;,NWctL'O,. A.1~~m, Eo r.:.II&iloI~ L 1, ~linIlm. Dl'I'; ~ ,Z'il~,l.c'l r(20005I, 19'. ~. Q. 'TliI!mIi~ A. IlifitWii\ R,~, I!ri!omf!~illljlj, ,~JIi,"IPl'll~I' lll~. &~ ~'l''l'9!Bl,
],.1'1. It

HI. [!;. ~ ~1111'~ er rI?l.. GI'~ ~. n. ,4i,<I u'.l9~i. 12. V. l'it1g, X. 111. ~I'.I!I Pi, 1M>;!rb:!~. ~, It, l.rIO!C!r\, G. Q. ~A, ~E~arA. ~.~053(;!00J~. 13- '~" 'l'm.g, G, Q. lruc.. 1J'e1li. 8iD1_ H2, AB4 (MOl).,

~81~~(JI)j"

~~. W. :11,W~ldrl~ l !l Bik!l'll, tP', IL H~~!~s,. ~. ,l,. WI'Oll!ili, e:.. ], Itoli;Jef1~~.(tIl ~ 797 U. 99$)'. ~,li. R. Ano:eJlir! ~~ ~,MJt; C'eII &'01 ~ 6~ (:!OO6). n.~.rilfJuta" ~, !h!!OOw!~ '(. Oi'ii!lat,t, '1'. $eij, M. 'S.aiICQ~ filar, ~ffJd. :1~i 70S 1[,200!6), .2'~" 5.. D. Viinoenl d ,o.r, lMlI!1fopro.~.nI1U, ln~ , ~;::-OO.5\l. lA_ ~ Yama~"JI:l1i, IH. KimiJlJ"a!., ..]iI!la.. rt I'lal:allu.~!. ].TiHIa, M fi~m-eE.~ p.mA'J'NS S, 63!l GlOOM,
.2'5_

l Ann!!',

or

lJl,

OulhJ@~k,
):;pt;.'Cii1c'lii(m hi ~liI¥ci"l:";ntlr{ gllni:-ollri~ (fC'ilhln~1 mtc.~ bOih tlH.' d ~ IlcreilCi!li a rid AIl~l~}'~i\lo ofl.)Gt"'
C(H11i'llUlil

n.
til.

lit OIl!iJ. rIl E~lJf1II5~i, 8, ~ lI'.l!dlll!1i, lM~llIp.!Ii\~nt 13<1, 137 WllrI}. G. III.G!:ii!~~Lf,!l. K. RIl!~~~Ol, liJll, ti. G. 1'I!tl!1I!ili, I. It. ('AN"'. rIIi«. ]<6,. 10'n '(zoo6). 5i. 4, Ililfbill', t" L el!\:!n~, ,f)~, 8jDt J:9:1. H2. ~l\Da'~.

~.utJ.~irdic:>. .
n~hl?

\'\r'hkh I,.~w' role j Ii Ihe inl~';gir~t}' of 011. p grun lib iii [h~germ ~:1I5 (if C dl.'1:.o(J:,~1171

ffl. A. ~U1~n~ It Rll:r<lllil..

1'" !!a~~

t~

121!1' 1117

grumllCli: ,Inti mm.b'C mtRN.A

rich and!

c~~lIlmil1 s~r"'Ci1n~proldns, hI CflltSuil'4fnidtlf",.l tlni 1::'lU1t ni~"S,lhc loc.::~Ii1IDlioil ~:bf Tudor, un C$-

~·['1h'l.~li in t!lll:il" .!>-pcdl1c(~[iO:I]. Al~ ,~'Ss.l!mir~1 ltoocs.~~ly (iJ.' thc :g.!r.m 1im': c},ck: i;s to 1ili"C'I,'cm
(l

smnc-

(~~uur..~ u.!' -U1e~nJ;,'Chml~nll:::,il1Il!JJn] t>0DtrOl,.;n<:)'.

i]$ I

semrol t'ornpolflclil, oflhc pol\:: I)~mmilami nu.~gC, is f~1xlHshcl1. [~ ~"ill be inl~mrLtilllt ~o tJ~:~;i"lninr;;whK:'tilcr Pnn~5 lb:I~.,m,wlro: ..:ill riI ir.:r' illl til c PG C pl'I:cnn;ur:;.
eiL~lt'r Ihl"!iJl.l~~~ ail:>, C'l'Loph~nli~ SLIil!· .:>1.u::lIk'l:i or~hmugh 1~~:!I~[lC~:JLlon wilh :~:U~~llpl, r[ Fig., JA 1_1.1 ft.'111!1il'ls to il~SOCi'n w:il(ltht'f 1~lu{llr damn'l1 p'fot,e"Il::i" :ilOclTIiiC o..f whicl~ :me dCI()~ll\d
!l1.i;~t'.

arc tOOl (mm ~()mmIe ICr;;Ih> lllHicrgo cli 11:C1IC1'IrI ~;tt~orll.

lhl' 'rowl1.ti:!iI

Io..~

(I

r :~ lliripll_~I";rn.:y

\.",hicl~ hey IJ,;:g~n10

(200H. 49'. 'to 5er.:qf! ,~I., ~~r. 16, ol.~ ~2005~. ~ ~ ~Q. G. !;I!!KO~~,,!~!ll, I. Iti\l1l~:fl1:;;' .s" ~. ""f'llo'!l, m. A. SWol!~~ ~ ~«eg, Yem (!fj~2i1\l!.4ollJ.(~M.. ~!I.. ,!". ~Ij!li" 1(, ll;e~. 8, b HQ\l~!\ '~,!lt~ 8>11

~n~~}.

A,u r!ilnr..'tgil1g thr..'til!.: ii'i pc; C ::'p!."I:i lie-ai kllil i;:;

in.

:rlJlk oj' Lh~ :1U'~il}il1!.! l11euhyJI3t"l" III nli .."S. LI. .'§t1l!'m.~1.0 h~~\!.! [I. ml~ :I:> ,[] IHO~!L!'in!m:gil1.i.r1m !I~mhy lase fl.ml :i]C'Ls 011 emnpo11~nl:-i(jfth~ gt'~ll] phnom ml(~]K:"II~~ mOlilltlJill 10 Pnm:'i

3:2" .~ I. l7:e~di:, L. s:.. Dii!ler, L ,00M", P ••~ OOn.il'lilD, l'II'a.rw~ .:lS:9, SSO !l!;1'>l.n n ~_ liI;ijk~lI~1 ",1, t.lr!l!l'!. ,Q,~ 1.'lI.7" 15 (~!OO2;l. a .... 3''1. ~lDli1~nk "'- Si!ilrmr.o1 [J]i~r.al Illll'TllMrll!i [m Ihtr! rn'ra1~5Oi,piI: aM !he 'I'I\eL~oon~ Trllili.fllf il I'Iilgrarnlllllt'
6rnIlL

SPECIAILSECTION
d~l\'c
Ihe'

:5' ends of
1'(1

,1\.gt"l3·"s.'iOciafl}{~ ill

l~iRNJ\:ii dIC'
:gCflll:
;11i1~

(~ig,H;l)..

pilRNAs i'n 'the Ger'Hl Liine

:SmaU .noml)d~ng RI'IJAs.have 'emer~ed as poL1l!liIt nlg,ull<lliOr~ cd gene exvrefiS,l,Qr'j at llolJtI 'uam(riptnonll~ end ~o~nralil:!i(fi,ptionalleveb. Rieceliltly. <I class of 5mall IitNiAs ttiatkneract \\ll~tl p'lrwi ,prcneim. Im~ been dis(,o.velredin the' mall1il1JlilUan germ 'lhilt' <l1'I,d.Drosophila., Tlile5l:l Piwi·il1ller<lClin g RfilJA'S (pjlFlN A5) Ir>ellreser'lta! diillin C:l :lirn'h1U RfllJA pa,~hway. 'h~' i:II:I[::lmlaU!,::I.~ j,m~lIl raoUltctllH!l KNill liS r<.:gli1lilll!O~ ll,r I.rlilllll'cri3)lion, RNA SlIIlJility. rJil'U:I U11111;lmi,tHl is, be!Coming. IncI\:1~il1lgty~\ id~I,1l t j}, Th~rui>'Cnltl:y dlseovered IPi.RNA:;; Jiflh r~nl'l til e known :0111:] H j,rHcrfcrJ rig RNA ~ ,(si RNA t'i ~ (If miC-flll· RNA.~ (IniRNA~1 in ~cl;'~r1J1W.:i)'Y"" 11 6). fir.;,I., piR NA~ UlIl":I",Jil.::L with [h~ Pi \.I, i, Dlllt nlJ1. 1i.t>C A,·~on.l!uH.· (.A,gnj, .sUbIaH1i~}' of the f'iw ioj\gLl ft'llllny Il'·fllid''l~. Mouse' I~il\'\r~ ~~V~iwUL" I:'e. qu.ircd. 1:O~' m.Nil!. b~L)gellC)lb, <l!1dtor siahHity r whcrca<; mouse Ago~ i~reqllin;:,11 t'or the :1>l (\ p;1~!~\'\'Q Sc~l1'fl~. pHl, N As ~~ ~4~(I .~ t R y, n~lcicotid~ ~nt )~nsi-Cuc1 i(i r - 21 Ill, Til in!, ri Il. NA!> ,COjt~f~~~)r lt~~,:Ihan S:f~,'tI!lU diffe ~~i\1 ~ :!>fX"cic~. in ,-\Cmlt;1!~,! W ~~'\"C'r.:l~l~lIl1.df\Cd peeics ~ or IniRNi\c~, hllr!h. I1flDS,!piRNAs nm1!c-h tn

W 11(1.1:i~C' the

les or 11iRN 1\."

!il'lC'?1h~ goCf!~m~(: isllib~t.i~m orl1iRN!\~ d

i~n!'i0!t~I~1 ,C'hIlC8 Ihb ~l'j. Hl:!P 10 1Iu iq

d'IC nmc~iL1!~Qr thcilr ,~'iwi pmtd1l-' pm\'i.d~

(ItliC:.liol!. Mt)::.~plRN/h ue ~i~~:;: ~!lIh~ !J,t:! larln~, im:Illdi Dlg

imcrgc n.ic,i n~m~!i,C, and

Fm C'"~.~mlll'r:. oilly

n 10 20"A;, tlrl'Il~l~!rn.\lli:[IInl

~XLi}ll i (,'

:>~'!!'I:~!cn.;:el>.

or

in Onl.JI(J{JbUu •..:!IUI lnJ!dil.i~:m:ul ··.P'ill!!l~'tll1g·· 1TI1.:..:i11UlliMH might be invo.~'I'~-d itl f1(~dcn.Jlilig ".hear pr,oc(;$~illlg !from preCmli{~I1;, (.'I, rJ,), t( fig. ~ I. Thi~ ii~ possible heIJ t';nht;/; III,.;'l\'lIb· lmd Piwi~llII'Mi'(;i,II,~d l~lRNA:s 11I1I'I(;h tl:r.c ;mli:;cn~·~ .~~rnnd orDNA, "':l.mwullg
t!L!r:i,'~d p:iRNAs ~Isll'(jn~ liiua~ r(l.r~, umr;yl (U), at lilt, .S' cuds:

,(n.

yi.ll i\.~tl3 -,I~:HI~iall\;-d piRN A~IImll.· ~I the ~i.:'HI~;~ !l~ran!d. and r;:o!nl'l~mcw. Al~h- flJl,d Piwi"s's.t~ei~i..:d Il~RNf\S in Iliu;:i,r 1in;1 ~0 nacleo~ides. sho \'Ir'i~]g ;lI ~l.lI,sen cd ,(!4;: Ul~nc (j\) <1/[ Il(l:;;i[ len 10. ·r1lU!'> •• he A.Illo;i;-pHt • .l\. complex mig~t g~lidc ~il.d eleave rhe S' C~(~~ (If l\.~lb-

kHilldnl~ Thss. piRNA~ rnay have d~'I!:'r;:,,.; :fulTlclioll:::> lI"w.un cpi~n~tit' pro,gt..lmfn in~ ilru:l .r;\:pl1;.';!"i>illi&'tnm~rnmll ~tDII tLl Ilt1oSll.lrIll'Isc.r:i!'lt ion:1I r''"llu lin ion. fui,CIILl f ~hb'1S\l'specu l~tL..:rl roles is $llllpOr'IJ.;tl IJy ~h~ k:~IOWIl ltl.Ultl ttJ'11 or UII.:ir p'LIlLII(;r PiWL protJh:in.';. F{JI" il'l'l'lmlc~.riwi i:. !Ill qligeuh'::l,ic .rugllllomn" (.~t. ~~"o,lom.iil!(;:..'> \,.,.ilh P'Ji)'JC:mnb gHmp (P~GI pwnl~il1lli ttl dU:;I.I.l' F\;G rt';S,pUI1~": ~qll~f]C'~ in th~ g~I1IlJ'Jnq ~9}. T.11U~, 'somc PI\Vlolll~~ociali.;-.d 'piRNA:. ri'my be Invu[, cd i~l c:pi!l!,('!n~lic rl.."!lJli:rJLim~. ~n Iht't. ID'P*'Y i'iRNA~ in JJI'.fJ-"!1f'Mh~ have ~~~cn~I]pw!'l :[0 dm~'n~C'gl_ll:M.C' scnsr: g.vpsy Lrrtnscri"ts t W)" ~nl l~iRN/\s m!lp
tlflHl~JI~j;';OU:'

10

a~~n(u.HC\drcIPCaIS,

untl

111I,;~n.lla-dlll:;rtl~[m~ ~2},

,uld

t\ub-I}i~\ i f'I:iRNA C~'~lll~k:x might iSuid!i!! and

Piwi-t'!':<t<ClCV'~I.cd lliRNA~.

whereas

the

addi lion. ~) wi l\l'e\lC!ll~ retrot I;';5ID1,SpnS{Mltnlll'~j p!Osh ion hi the tesilc IJI~r germ I ~IlC i( W~I ich Si1ig.g~SI~ ;I s~{I"d role 0 r r)jIA l-assoc i:l~C\~ pi!l:NA~. f~I!i~lc'nnflre. Piwi '.III~IAub app~;ilU

m.

to fi!i:'>.Lli\'cly r~~ul;!t~Irnn~.I:1!li{ln hi carly OtfJ,~'!lt~M!a(.'II}~ bl),'Ci'I> U 1. t h S~ i.lnriy, M iw~ m

th~ gcnmnc in ,c-I~"!lI~rs of:W If! 91] kl'l;ll in n strand-spec me nmnlllcr. W~th ~3c1~ Chi~N:1' likely rcprr:scmh~g a lo~,g :>illglc-sln1""tr:d RNA 1'I~"C;~1rso u or. ~n(:lrfiC olten, t\\,'~ll irJ<~IIDvi~rlappin~ and di'll',.;'r~nt Iy lr.nl:-,~(abcd pr~"
II:: Ul1-Ifll1l ~Fi~.

IIlui IIII IU[jJ

in]1

11111"1111tlllllf[lNA

-'

pNlJn~"!e~rhe )1!.ubv~ity ll!1id •. Ii kcl},. transIlniou of its; ~a~cl ,nRN>i\s t i5). lilliS, ~a'I!IC IliRi'l M Itla)'h;w<! a Ihirrd rotc in pt:.vsilivcly rre~~I:ku U:-~!l~~m [1m;! mR.Ni\. ~Ul bi I'ily, ing kin
The discovery of piRNAs reveals
:1 11~"",'

u.n

IA~ t::'~, ~n cnllUTI:S1. :s,i.RN}'~N arul mi It."{}b arc rJt!r.i, 1!.1c! from dDlI6J.il~l.mm!m~ Jlru:l"'~lOr!,.ih!Q.i:rp.iill.llN A precursors, respectiveIy. Fimri,ly,. some pilR:NAs mu.~h~ lm,..ili'l'i..':I,y regLlhl.{· mRN t\ $wbHl I)' "m!~rua."ial ion (~t"C bcl,,\,\·'), ill cOlltru."I. I{l the !legal lvc (:flee I. ~)d'l:ill.1 ~iRNA alld mi.RNA~" .. ln D.I';(~,·t)[lflilc1" ~mm:' riRNAi> 1m,,!: been calk'tl [(,,~JL:"'1~ociaLL:d t':~RN/\_" ~·'r::I;siRNAli··} 'l"t oJ', (II. UOWi.lUlr. I,IIlHkt:! ,:.;jIlNt\.~. I.h~~~ .F.!I~IRN.i\~ bil!llJ ID ]liwi .. Al!lklL"l.!liilL~ (Al!lb), and A,l,!,o) nll'cc Piw~ subf;am~~~protdlls.bm fUJI ~('I ... \go :>LJlblaJ,11ily r~eii1:';. M~o\,>cr. nhcir prodt!,zti{IJ ~ l-.cqul~ nc i~~'ICfDiv:.r-~ i~(lr l)i~'Cr-::!, wl,iclil g~l1!o!r"'l.;: miRNA~ ,~ld siRNfhl:J.!."f"!!J;~iv!!I)' ~.n 1l11tj~.h:v dr.:'!ii~ Ltt(1U. iJto!!5'e' J;ii."i RN A,~ .fire rriRNAl>. U~Witl '~P1: plHNAs pmu.lJccd'!' Til!::,!, ~~ li,J,:,dy IJffilhieed from lo,u~ iSuligl.c·.sm.lildcd pl'~cL!rso1"$ h~ yC:~-lo-h!!"idcnl ~ ~d i:n.uon~lIf! dea$>C;S,,~n. ,DnkWrmil'I" .1P~wi. A.1.lb. ,Qn~ A!;!.u;l: ~~)~dd :;;.u~h IJ\.; I.·Hd.m)!.IId~N~i.'S bc~mAiS," tl'~}' h;lve ~~Ik ,i'Ci:i ,'i~y N. tJ), i'tlr ~Oln: irdJl:-po~~~tc ~rlg

dimension CflJ,Cflt.: rc~ul:!Illioll. Fultm.: ~tudicll un the lhi~~l·nlQl:.j:!.o mac! the pi~~~'n1~aIlJ d iV·iJl1'ii.: liml..'L:iom or piIRN'A:;: :.ht].1,I1d :>lIIb!>1l111IiIIUy udvunee our u:m:l.!.l;n;~ ill~ () r .~I.1IlC' rt'g!llI[I:L!iu~~ u ml
~iml de [IIIC-:>ihegerm l iuc.

rui'!e~li;j1(e5 ~_~. Ii. II.. ~11I5E4!ri:, (~ U4. an {21OO6h 2. II. r.t Kirnl, ~!m.l. 2>0, l'l'.B (Zm6}. :~" V. 1,1. 'Ilagrn,.i!'I ~r" 56PJlc.~ ]:i3, )2(1 WIO:G.J'. 4. It. :!iiJnJo ,et at .. G.rn!!i~" :!m. 22M Wl1l6h S. l IlI:l'I!n~d:.@ tiL Cdl ] M3Kh :2IOOi:/' ,~ ~:l!Il.lllJ:!fJ..uILt!OO7 ,Dl,!J~~~,in P'fS$, li, L S, 'Gi!in~'IM!fMIlJ~ tl Il'!: •. kirn(l! 31;s., 'Jj&;l

~~1l, I. 5i. r, 'timilllL [, ~r~., llfi'!IIn~,H. Ill!,

HOi,) (lOO(j, II. Ji'I. PJ'l·~lj.i.1Ja,~

~J:~.

IDi),

,,',. ScU~, ]O~,661jl O!OO>l). ~i.,(llj' i!.~~,957 ~~009~ • m lI!. ~Ii~on, ~. S;¥!)!, (l, !!i~~'~!IW1~!'Ie',#l ~t~, I, VI!~~m. l'Q:5l u~~i, II- )l. 1.~~Em.)'~~, M. S. ~~"" \I. /l;. !3\Io1J!fe'l',~~~~e ,4{id~ ~es. n. 2002 (aoo5i. Jl. II. ~M!i~!I:l~, I), ,It CI!1:, C, (MIl:I!~!, !il !ill, Q,Iff, ~"l

,. c.. ~~~

t ram(irif)lioMII t~\lel:s f'l'lot ~l1,(lLl.fIII).N~,'the 9th : 'l'~le S1b~1I1(~U(:ern~Ef, 'i'a~ Un'nii!J:sM)' Sdloo'!. of M~dl~jlni'. l'Iudeoth!l!l;!' '~r()mr Acto lin A!J,el-~iRNIA;l!h~ qu~¥ Meu Haillefl., (f [t6S~fi. USA E"ii1\\lH~ lialfanlllll@yaLMclll tiM mark. I()~hff IfIUC teil5~.,

['e:sulltirt~ p!:RNA.-prot~ilil c.omp'le~fs U,en r~lI· l~l~ '9f:n~ e~lpr~~~iolil ittnll' e~lg~!1I~k Qr ~~t;.

IFm~. l.. ~A~ b~t.l1rediiOnallPiRN.A [lln~ter.,([B) A ,Ao ' PfOI[)O:sd pilR:NA b~'i1g!!'IH!~i~ piiUnV13Y, Ihe

16. 183'l '[lOO6h E.. Wil!!r.rn..~ E.. Cooru~11,. .1it MaullJnallll. lN~l7p.IOOn.r P ll!2:, 163 1 ~19I1M, Ill, 1.11" 1R1iI,g, ~L lill, Ow. Ci!lf 2,E!1.9 (lOOl). 1'5. 5_ T. Grima, l. I!)flIila, Hl.1L1l', f'fr1c. MIll ~Jl! sri. !l$~. 1(1"', :<HH ~2OCI6). H

J.

m,

WW!,'I,.sde ne:emag., org

SCI ENCiE

VOL 31i6

,20 APRil 2007

Germ Cells
j):Jchytcn~' I fJ¥ .M.e;thy lal.ion lI":mcms .~~.ahc '~~n[ll.111
I!Ilulili:!cr of p;Ltl;"l'n~lIy

mclfl')lla~(,Id El~\!Ut~

lII'C

E,p,iigen,s:ti"c: D'IB!ciisions, iin M,omDlali'an Gle!lrml Ceilis

:Spedflc SieqUienc~s alIce ,de,(g~Fla~ed JiDlf d,e I'IJii,iO, IIl'NA M~Ulyla~iDn .a~ q::l'G iliI~i1IlU:I.eDlide~ il'll mammalli,!II!i Igi'!nl1l cells, "lie fE'5Ul/t g~ the long·,term trans(till~ioni3~ sii[e,f1.(.ingI(lf tliie l'IiIel'hylated :5eqw:!,n(~, most 01' which <lIre retrotrafils-pMoiilS antll CpG,·i'iclii ~etJbl~n(l?5 a~5Jodi~~ed wHh ~mpril'lt.fd {lfFles. Therei:!! pro lo Uf'l1!i s~xua~ dimor'ph~sl'lil ~ill both Hte 1'Ii'l;~ulre(Jfll'1e- :o;eqlJel'l;(les that IilMerg® de FlOwO melnylatniJlrlilii 'g!Niffl (ellis anlli ilnlttJe rnerhs illi:o;fl'I 10,11' whicn de r:JIIWOrnethyiatl;QFI is r~iul.atel:l. The re~U~(l'iQn efrluhlre ~ejiJif e;!;pn~ssiofll ~~ tile lllllp(!slttOFli of heril:al~le me~Iu~l.atioFl liJ<itllfriilS tn Igerrfl 't~ II '~e!i!!:)m~ i~i!:h~~i'!dcrhtitef mi!! ITlfllals bli! is r,,,~~i!loth~r ~ ~aI. t

thlll1 111~ 1.6ic prohlCl':IIII.C'!J1 and ~. liIl;igrn'l'i~~n or~:ril1i1(lrnia~ !l!;t:'n11 cel)"; ,[PGO; I :lltll r~csmbl j",hcd il1, ~t·"'.-:;;Il~t:illc
1

C'llC!1n~(".

m~-a_h~ n palt~.~~ lin: Jm!ld~ l~ltil.lj].

1.:1<~'I.'dI

pm~ClITk~

Uk MOS:Ide

durill,!Jl

:'pCll'I11:1Li!lg'linrn>~s

~mdeogencsis

DNA llK'lhyl~tio!l is dh'C'Clcd ~fI ~rnnsposons .. nd~h~ir rcn~fl;i!HS and ~,O d~I:iiI>CrOO I"CI'llCtll$ lprin,"!r.i!y ~ri~ml'~ ~i:!~(:!,lhc DNM. with lesser mllfl~lf~ls al ~i!lglc-cfIJ~~' ~~~t!enc~ ,md rl~ J~ITC'~~tliJUj' i1~!hylalcd
Hfl\'fI

:;'F(.'Cil'ic I:rnll:i.eripl:;, that :llCad ~IJ' the IProdm:lilO'n of" I.mm:::al,cd fonn~ 'Lh~ iH:OIJl.~ili>". or 10 ~lLlqll1.lnc,"":o., uU'Lran~"1i:r~d mR.NA~ I.h:u, 11m>.' Imw ~r:<J!l!llla. :J I:r~lt"[~-d l'iJ1C' 1'IJiLnrtHI!!d~,()'xLlal dhtlL'll']~I~ismin the [t1'r.vprepenics j f'i!l" II. The ~lurl.ci{!!,mL~ L c';m· omog~n)' or 1'I'II~>;[hyl<JJtio!l 'rn~tl(':m;;, ~u mnl!;!' m~1{~ sequences Ihe Ultt;rI111liHI m.RNA~ [IUd len' S.!:!:gCf!ll ll I1CS has a d j!lCe~consequenee or! p!"\lil~dn~ ;-L!"C t~rgc1y Unl'lF!OWn, The oocytC'-

or

hm thcllC' is ,I fh~h,l~({l tlli\::lhylalc IWlmlF;i~sp.r.t_<:'oi1-~, "nN DN~,ln[_ p.!"O~ci:f1 ".I?" ~.a1S I{I bi: c~:s.cnt~<I ~dcl1i k·.rl iu hOl.~1 Il,'l:!fI!n ~~>, li~lC~.l~lIJ dl~ ~i!ni"~ C'1ii.pl'e~km~~aUIrJ.:.,\Hy ll( Ji IlcJ'(!n~ iH~~le two sc:-;e,<.. til e ff!!C'!hy Ia~km <I1u;1 1~I~C~IN~pes pf DmmJL-mnl! l~l~lC' ;~ud1'i::!1bla~c r.~ relics nre n(.':;lrly mirror ill!l'I~S.. Tinis; rm il11plros dUll PNMT~'lL f1Q[;;:-rl!i~.lI)i iJ~1£rprelS. a Ilr.'I.'C;\ L:;I.i~l$ Inilrt llilm i:s I.;<iimh~i.~h;;;dby ~}Ih.CIr liL!.;t.!lffi !~lIId k'l(~utcdi m. tII~.I:K'fC,r1t !!"'",Tlimllc r'~gitlll:S i in ~ltlllIl~ rn:lll~j,t't~c mnl 1:'t:rmJI"," 1Pc'!\.'>tn1!.:iw,tic ~j,'iml ",dlK; llil!.t m~Lllrt or I_h..: nnk m~.)'be u ])I1nit: IIlhlr ~m[;ll:t'dIL~lmimll11 hi:l.!OIII"," 1Ilm:l.i 1 hic:nion Or set of modi ric Ul.imlS.Th~5 W!lI.ttl ,;,xpJlIii iii how f.li:\wlilc nraodifil~mions lwhich have not i;Jc!"'n sl](Jwn In be su bj:b"Ct 10 'I~):I]~·a~~I111 wm5l\ L'I"," in!l.l:ritmu.l,;;j mighl. h..: C(lJw,"rltxl ~lfltQ1l1.::rtlabk p.alt("IITIS 0 r ON A 1Ilt:I.l:ly'1 ~1~lmli.h~ACl'LI1 iUljJCJo.'jC Imi!;l-h!Tllill uunscript iODI3.l1l> i lcnci 1iI~ em ~Iu~ aJnU'!1j
!lOb'lll!lt

or

:1~cdfic

IQJ'!'ll

flf DNMTl

~DNMrh)} h(ls ,~

the gcn{lmlc characteristics

r~,gi41ilS tDMR~} FlI)j',hll('lril)~(j

i~~IIIl",L<;.Ilf.sfIoCimoo ~\ illl
i(j ~

or

iU1llf~nlcd 1(I<l.iii ~)'I. ("1'10 pmllU'1tC'r I"C'Jil~on~,

fir

II accu mll~.Hes 1.{lJC'I)' ~I~g.hleve !:s inlh~ cj'I{I\ r~a:-.m fif (lot! ytes ;f! Pld~.;:~i~i~iPl r~!:i ml~lanl~lkl~~ cmIH'}'n~ w'l~cl'C ill 'C'l~lcrsnuelel m the~~ghl-C'C'U S!~gC' eo In~in~aill .~lnnrililCd rn~!hyhltkm pmlcL'I'~~111,

h~gh~ ~,!,,~'H.i.~}' r iC9mnilNd tfl I ~'le.\fIoj11;Hi,,;: Fflml.

associated

wlt~l p;lt~m"Hy ;lml

or

I~H.':rJM~:;: .1~l. .

~r,::rn"llly

impri!r!ed e'Cnet- ~HI. [:).:llrl,\,'(fi IWlii:!hyblion eeeu rB .~~w!tl)1be I~I"(! fl'\1u ~Iti(tn in grtl~\ illg
Ql.DCY~~,

,~nd

n'~Il'lyli!ti{lnl~M~crr~ f\UinlOf(Jkl~

erased lnthe
J,;-.cncrol ion.

germ ,elL~ (~r 1lC'J'i~ '~Ie'

Uiml rulPC'in]i:th~ 1-

<1.i:C'

(>@on

~ue(l ~r1 .~.c gcrlll~ Iine arc nlilid t)' m Ht;_l~d lD DNMT:3l-11iJ'(c'cd de m)\ (I mClhylm~on Less (I hlC~h}l~m ifili. or a wi h.ar..: Hi csmhljsh (lC'CU1'5 in p(!pl.!~mions rl(I,ndivkli~,g ~II~ itl lrti and l'r/\ diinlcllcmiJC'll U.,I'h ,md because tIt~'il h~ iOil p;;uu:ms. reanimaies Lll t'C!fOIf(LllSP{),,'Vli:nl~.k g~~rnn:Jiulc!£?,I~11QJ1iIC:S, jH'~rtlellh!l'lmctl to'r '1111$ in IOcrm J:na )i{l!m,~~ic ce~~$ and C~H!!$'C'" b~'(h mill I" ,und fen1l1 te germ I ines {rig. 11" Ilt)wever, ml!lhyL~iion ;il;,'ql!J~iL i'L[ni,~ <II. fln:l1lCi(~ti c mlt~! 1L sraull 'iiLIlt1ber !)f ~d I tli ..,i"iori::.. ~.hc hi ~.ud ~,)ir~~;(1i1 or II; "~~:;t>tun r impfilnlca ic ph.;jl(ll'l1eFlfiil in! he ~'i'l~ I:c-l:icnij~ Iine, ~\I~ re i~ il;lcirl!.l!ll.c(' Lll.' c___.,.'J rI'Il!IlmioIiJ' ,:Ill ~tI~I.HI,yrl:'lJ1)","tl ~etl~~S ki e.ll1bry@rli~ I ilt-:'Li'!.;~II·' II'i I,. E\ errl In1 ilOJ'

_g:.:tlCS ,lire mOl~~!I' ~Hlnnc'h~'!!lled

("p{~ dh,u(;lloolil..l.:!l.

or

(J

di~lllIlniml OrmliC!llIyl:lli[,l~l pauerrrseun be kllmt :urlll u Ii:hmiJd mll!li.l or di~vd(lp.1'I1,",rnal, abrllJrlll~JI· ilic,~ j:;. :;;t,:\::I1\~ hell ,:;;pliXi fic ~CII()JII it roegioJl:S Ilru ahr-.mnu!l:ly mClhylmcd I;t 'fh", DNM,1"3A B ,Imd ONNITI finnilic:; or DNA cytOlline-5 Il1IIlIJ1yltrn.L1:;rij)ffi.'l~" am R'" :,poL~ibk rm the ,:~aM i lnnt:ll~ ::mr:!l1:mill1.~" .. nmlcc [ld"mil1IJryblLiml P':ltt!&rEI.;;'. fl.·."P"\!: Ii \ ~ 'y, mli:!l .!! ~~ ~.:q'lre~",,[!t1 hi IlilO!o:I. d i \ idling (leU lYI)C\l< l2 k DNMT3L ,([)NMn •. ikc).is rdwtl'<.l ill :i(]q!]~llCe ~ to l}N~nJJ\ ,Llld [)NMT3B~ml. llm:b l'illr<:ym~~.k .[lC~i;vily_ Il ill exprcsscdi 0I1ly .in genu c~rL~ ~11.d villy ;]!!~ be :l2t.~~c;s,"",hC'J'( dC' 11('1'\'0 1T1~lhyla Dioll jl"un: ~ ;Ltld .i.!"t-, diS .L reg!ll.~ior of .it DNl\.H3l'\.~mdDN'Mn:ll ~ Mn.;>t /)111111 g!!~C'.~ ~,~, (!{lrlt:.1 in ~~,\-$.""~ lfie gcrm~ine rH'tlrnOICI'!> !hm ur~ nClh ;!~ctI at !ipi.-cilk SUIJ,,'CS of !.!!;jtnC!vgl.'ilC"lij~ I( V 1J), These pro[J'l{licrs y;~ !.:: lI'isc:til) gemg oC'~l

oeeurs in plCll':JI.~1I pru5pcnumlo;guni:.r. ".,'llillr~a~ m.tlhyl:llit:J1I i~ a~'~!IIirr(ld 1]I{)~1J.T1J.::iolil':;lllyin.th!J rl'lm~l""gerlll, lim: ~'IIIllruwing oocyte:;, l.h"l. HIi!..: nm"lC(i 13'1!ldi:ll!o[cIlC :st<l1\'.1> ul('ila..r~~(l, J,. lm;:; of O:-.!MT3L :r('~uh~ in very diill'c:n.::Il[ rliic(Jo.cYJIC1\ depcl,ltUI1I1J UPOIl LllC' ~m.;. eXITI:r~ Ddct.L(,!H DmnlJL in r~~ mioc 'Inc\ Cllls ~lUlk l~st:!bli..Il!l~C!m 0 r: I1lmlAmmi 111l~tml)'~:IIillll impri 1I1~. 1111 !}~~}'~[!"... ,'V i.n1101Jl1 I1Ilm,rk~,j C'm'Cb 01] r'Glm-tml1spciiSOIl IllIJlh},l:pliul1 (.'I). The ~'l\ult j:; u m!llcm!ll·cflb:;l~("[hai ph~11'OlYrcil1 which the l1C1em~gou~ olt~pr:lng I'Ifl1UJlilCli'..YgL'lill~ I)mndL· m~~11:"erl'IDles (which ;)II"C' H(IJ~m>l1rlllCn~l:yrc ~ ShtlW bla~~dk o,;xpt'l:S.'tit);il !,'Cn~~ i11,lt ill\: U~Nm~llly ~tll!h':i'milly fill~~hyi!lllcd~md re~;ro;~~~1. rlti~~c<nlli1, !(~ilIblwrm:11 cl~\'Ck~Flm<:llt {I f I".":.:~r-;.l-

.n

or

or

moo.

,:,;il.~:i \iii i~I be tow. In 11'1'1.: male ge:rl'll, line, ~.hc elluiru (.uhr.WlIUif p.m.;pcrnmlUolloJ1iu lDldel!!,u cle Illo\',rJ UIiC~m]y1mion ill [bt: n)i..":rin<ll;~], IJL~rL(Jd.ilild 1'>j)cnmltogrJllll1 1.11usl Ilimulln~rll 11IIi.nhyhn~ml p~~Ul;'msfor Lilli: large ,l'I.mi v,Clr.~lblc II~nTll~~' I)f

CllLIC)t~C' d i,'li~~Q<It<;, 'IIml r:IJCCOOC ,"~p,,~rj :311~ tor I.h~ 111fiprin.!l, f

~n1:ry

into rudo.-

:-i:." Thi,~ illere':;I:S~ the h},~rl()rC'-.T:rI1Iu~nliflf'-".

or (I~d.of,lr1111:11(::.,and
r

~i.. 1111ul:Lllmm.!Ij)I'i:.".":>lIlr\.";.'; h:l\ {! kid 10 :J low . 'l dcn.'i.i~y (If CpC; d~rl1lck~!irl\:>~ ill pialc[~lIy

r.1l.:~h)llai£(]LJMRs rela~~".,;to mmcm ... IJrvl.~s 1

ntHI 1(1 :1l1 l!!!Wcrr~ll"C6.cm(ld(ll1l
(I

~~r
~~r

1'ltC'lhy'IIIHlZclt:lMRs

m.:~,hylaiH.:~1 DMIts,

n:l;!th 'e !(~ m<!tenllllly Fur Uiil>.l1~~wn I'..:;hm~~. Ix-

[I<11iCm~111'

1'I"lt~rIHlli)' ut~~thyl!ltl!d MR~ l~MiHll1 D ~rmnih~ j1mmli'fICf1' im~Til1~~\~ ':':""ti~ whC'l"l:lI:> 11mi:rIIlflll},

I'Dt1P;llf(m'l!tl ~:I fjetilil~k$ and lJiie~ti!P~i'I!, (g~~e of ,p~~H;illm$.and S~~([IIIi~iI! (o1ulfl'l;l~ IJlii~l~ilY, ~.ew 'l\ork,

Wi' lC1(ll~. USit iI~USlEfWj ;'U 1Jn~i{fm~l" U Pilfi! 7. 'i5:1!i1 ,PdJM~. Ci!!'dr!:!: 0;, Ii.rJJla:!'. '''i.o whom ~orl'!!!itlOiililemce ~lDiW 'be< add~~d ..'ifOOIr lI3/UI, < Zl!m~ ,~~ 2: ~~ ]iI!'!.l!1U.. '52Oj,1l htk, ~i!IeI: 1Jii), FIt;v,!1;!. E.-ma'il: ~~dti!@'lim:~ll!l!Jlr

cmbt)\!;ll'lk SIn.ICl~H'(!~ arid Jcmh or I I'Ie i:.:ml'lt)l\! hc:fore mid~J;!,cs'll1ti(l:il. TliC DINNI}L-l111M pltcnmylX' hli'"mdc inl!~C:C is mlllt.edl:y di:i1icrem. Mall,; .!;,crm cc~~s i~UIl :1 ffick D~MlrJL ~huw 'IYh¥nillm.il'~g C;\ll":::;$utvrl n;tmDrull:;;rH~tilb ~~t" thc' UNE~'I Uo:tIg,iri'lCf. ~,p!;~~>d ck:nt{;'ii'[:.,~ :Jlid lAI? il!lllild~lci'ml~ A !l~lrlii;lI~~) d&~ ... . !SI:!Vcrl: lJ!l;ymll}~i1i,\ ~Ind, rl,tll1~ hiJmoIDgoQ<~ sYlliJpiS.il.ll:Jl rlli.:il[}~ic ~)rup~la:;:ll. :.rnd C\1l'nhllll~pD~tosi~, or ..II gctm ccH~ bd~l'~

11l~~hylm~clDMRs
in .he

or

C;!ll

1:0('i!I'H.'!tBm.iII'lY

Io:.ilooru:cs (1W~y ~~Y!m[he 3f1bctcd

1)~\'er.;.iIY
dcl'Si,!,;llmoo
~(I<p)' '!!'C~u $

gl!ili!S.

~1ifILurc {Jf U I,l,'; S'Ctllhi!iflCCS 130VO

Q~[iJ

or

for de

"crsm T..:rcm~J

~qtICiil~'r.:$.

ill ii:I':ip~~cd

melh}'I;-~~iou ~sill.liih.}cI~ll>l~.~d .. : rl ~:a.lti(!n,li~lnkl.!':O: it

lik:~l!>,~h['ll n1lJllliplll :>i!J~a~:. di.[C;i:L ,,j~ no\'o r.lll,_~lhyhilioll 1n !l51..\fl:ll cells. bu~ aim ImILW.'i:luf i:J:UJot';l! tiigmLl" i~ Ultd(~J!:.. RNA·diroc:l.cd .[)iNA 1I11Cthylllliol1 {Rd D'M I ~!S II cml1.uidaru. mid ~lK II h

398

SPECIAILSECTION
IEllrih ~rmal~W

...

-T'

" ·P~~fIl~
. ""

IIISpl~·.~!Ii PaJIl!flell9.
-t"~ ...

$ilIl~'

rcpeat~:!I.C'll,~il1l:l

8,jIamatlil;

COl,ll~cl,[ll'lenlte

ill

I'Ilcclil~n~~.m l1ummd:;1ll
~iI'

gcn~l cd~~ \'\Icn •. ~il~

~.rro.TIl iIlcslabi~:;;!,!'I1~.nl
!1UiUICU<L1J.oc
{l~',lCITll !

ONl!!iF1I ~b DNMi'iI"i!J'
~Ei1IB

I>JHlImwym~

O~l D~l
Spa,FJ\ajjil;

Or>l_ll!Q~
~f!'!l~

_-=---~_--_~ ....... __----:-.:,,-_""'

I'""

..

~n

uld~"
h~UIWl~

)'I!J~in!~f!~U~I:1'L~<:~f1~I~C

...

di...;,ca~'$..a.~in

ffllUC

eases (] HI~
:Ilild

impri!'~.iI1~diStIr!(I<!~ 8\,'!tkwit.l!!'Wi:C{~C'nna!t. AlIgdt11Dm~.

Ol~ DnmULin femaJ!.esnd (i!II~. A l~g~~l'J(:lJ)sld.fl5~flellmy(ilii f~~i~tai'IiI:~ mlilf~N wa~ fUlied t() tilIe prom ote,r of the~ruikl~e!"lii)'l.Is Dnmt3L IfJl!ne. E:i:pJIf;!'SSi.OIil (b~ul!1l)is Si1l!!!"lilil gm~ll'i iil91 Cloc:y~e;5 fill m~ adu1t

If!ig. Z. IEx.PN!M;iolil

male

g~'m

I'"'

;4;,..._

Somjjj;:

orrrunA.

~C,.;;' --

__

--....."-o

........

-::::::;::;:=-,;;.....""""'OIiO.a.o!_ ......

M:n,~ltl,fidrng pr,ecursor
!!Gel;
~.
~QIilI1ll'

mOiW ,oval)' ,al len Sl'

.a "olin

l!'~fIij

~kl:~8,.

!!i!ftlJi II~COOnie'1
~~

c~Us),iff} l~~ testis

P[&$~iirnGlt@,g[lnii,l (a tQ'5,peoo3,l"09IOf'l'ial. Sl~m hom ~ I'lcwbom meuseet

..

..

,++

right II)nm~L. ,~ I!'xpressedin teMis frl:!m ~7 day.sb~iij)r~ bhlh to -3 d~~~ art~r birllrl,
Nhur ..'i.I t!~ IJIil.: Dm'~"I!MilJ' mel·
,m.fogli!o'h'l"

Pmder-'!ill-rUH ·~YI~d~{Imc'So j i). ~,:I'igh rl:ll~ dii>t!t!n1j~nI"'1;; in mtl"l:Ml<:y gurie ~;w1~1 l'!.'lJOfll.ld rOil ~ ~'>'J:.tJ~~t11iiC'tI i~rclo:."l1;; IIm~' nri'l<l.Iivc mt.thylm~[In :~lJlill}nntlJ~i'L[c!>lI!l~t urI1iC I]] (;.Jrly L:!I~~'liIOiJJUrnc . ~1'j.'lm'll.in'C: m,Lllill:i.o:ms laiU1.OI 1C;)!piJairl 'lIt\: di~Cl:m:lJ'I.'li.:ll lind the piucmllil. .~c e-l"lt;clin ~chTZoID~hl\,'lJliLi. rimy

'~Jr

i'IV(,I~lL~
nanec

'!.iBnl!}> ill

the

Il1lIinll.l:-

(II' gcrlomicmt"thyl'lIiion pl'l'Ik:'IlI~ clu:rilllg lho: '~1llIn~'~dJI

nil LM 1.1 nd~I.'~n ~jTOr:;;

div isio!l;.~ [tam male !!t:TTIl. .:dh;

as !i.p('nl1.1H11l~1il

Dkcr-I, \!II !lklt b re-

i'f~..Epill~!lctic

~1~I,ri ~!Il!

Oliolmin'g,

=~
~,
DI:!I1IfIiI"!lL (!3e!'lll_

quilil;!dror uniCI!'ORNA vniRNJ\l pmcessl [QJi,': and ~lot ll!"fld~lc,.io£l!l (If' ~'i.m",n illlC:l'fct'ln.,g \:s, __------~~~~~

gc'm] cel'l dCl,leh111mcuH arc Ilikel.y W

c;fIl1~rn~lIlC

D~M.~----~------~--

r<lmit~ clf(.'Ct {If'I 1r.'LrIS~On me!hy~.members. Rlediildi~~es g;eam [~U-sp~ific exOJll5 ami :!lpti(!~lo!i3ri<ln't5 mklll and acth il~i. 1:ludi j;~H.NA.:S, ,p~ Idt; aflpm~im~l'e (iEi;!I1lill1pmi1rrtal ti_mirllg (If ~pt'~5~i(l1l iNIch J h'(lC'1iC"(~ w •u:fIlil$PO:l.~ble clcI01m ii5 snm\llil 'll r~tn, IDNMUsand [l'NMUCI' lire Ilaive DINA merus have m:1I been idcl1;~illcd melhyllr.m~Jer~~s;O~Mr1p fI'!'R~Awntains. mllultiple s.\(1~(l'Jd(l!"l::oin ~.!I, II app",-rno; tbat RNj\~ C01li1· I .an .a1~errNu;¥e S' eXQIil and is I'IIJ4:~rnnslaltedl, DINMlf3 L is ~.q:lre5s.ed in ron(l:nIL~ ill 'I.TI1mirmLlli:..u cells D-lJli1' rrQ~:\e oiIllII;! fem>lt~ '!Jiffili c;~lls Dlit bom diU~re"t PfI;IIIT!DWIf$, (lrJd m:i,ybe u:IDd ~xt'lusih:~~ fur a thiirdatt.e'lrr.lltive m:nm:ript.,. whi~h ,e.il aet ,encode f!1ln(tijon.il~ n lUi.Ito\li\ pnxllJ{~litln uru:lk'r norDMIll1iI3l,is. prese,n:li!"l :s~~rlll'liillil:l!5. D@n()~(1IllE!lhyl~lij~n O(~UIIS bejrore M,tJ; in ~1te en~ire m~oiit. >1)[ prOOf)ermalogoliliaiJi'l mates, br.n mal condi niOl~ anl.l IImt DNA d~ !"II;IVQI IT]!$tllylili'tm 1'fli ffliil1o)l~s; ~~~~~ !;J~:r~ 'mll!.i!'~:qFl Q'1,I~T'U1(! mCIJ.ylaliDIIl il1l rnanuuuls Iolmy
I[)jf

Fig_ i. 1~:(J;lr'~~fl QI: germ cfUl-;:p~dln" fbrms c,lOf!li!lt

qsmNt\~ s, """hkh Pl1.W~nt~ th~ .""I(!ptivc- RNi\i iI'e:lll!i.)~l;'<C'). Loss of :Dk...'r !leli\'i~,'in n~~~ C-1fQhlYtmk cell Hn~"Shas n mi F1im,~1

n.N...

to flJ:l!~ly ~~her disLlrdC"!'~i n ,~'hich !1dthu ~,nclic no!' \;:11 .~r{lmncIU<J ratt('l~s call acC<l1.1~ \ I n In!!,:lU .is~,

R.I!!H·ein!'IlCI!·S ;and!

:i. .~ ~'1. l'1'll~~r.fitprtJd.

2. ~1. G. Gdtl,

~. It A. ,Ro!loiij~tf ,ai" G't!'lMl~ '5,

~~oos:~,

fffl/J, &~,'Ie, ~ (2006), Ii. iIlMi'(ll,.A,IIII.[!i. _~~. ~ftMl. 74, ~81

NioilI!S,

Re.. lJ6" :tel;!' o(Z0006). '1. [. Li, T. II. ~IO~,. j:, ]311l'llst1\., Ctll oJiI\l, ~:Hi ~U'9,a

D., t;loor(!ri~ r,H_ ~, (_ ~. ~~_ B. ~'IIn,i!~, 1'. H. [le~J:I~. Sde¥lf~ ~94,2S.~6 {loon Iii, D., ~I,!r,~'t!i\, T,. I'i, ~i!~r, .l\Iarwc l!l!, 96 ~OO4i, 1. !(, 0.. ~()!M!rt5l'Xl, "~", ~~ ,Gt~~. €i, Sl) 7 ~oOO5J. !)., ~5'!!.~k~, F, :Si][J1~~~ [, ffli.r~~~ H, ~~~ilP~, s, fajimi" }, ,9.~I- (!i'fm, ~'t9'• .2~e~ (~Ol",

reum'ldl.iC:live Hr,~ 5pan (I[ 'the flE!maile"A;15(), de nol,lO, metnylaoon '(Ic''Cllr1 killgl be:for1ll! sperll1l<!t@ge!lli<! '!2ll1terinlo ml!jo,5i~ in mO!~es. bu!;
deoo\!'Ol m etlr~la~tcon occurs

!lei. I"nlillirc

JllpLlI

r["lcm~

.m RNA
ul muM
eells,

cnllll pOlll;'nn.

H QWC\lCT,
[('SIJ!.'}Ut'lil·

mei~i:i iin
'~!Ith\wy
nOSGrl:ii. oC iflCo:

fl:JT!\i!l!.il'S.

only .. ftN ~Ilie (fo.s~~flg-()lIlei' :nage- or PGC, primQrdi!ll i~~r:ml CitrlL

pbL'lI;:';,

adaptive R NAi
0(1'('1"'"0111.:0 il1

ir ;kla does

'J'. L MMJ'inei1 el t;i., D~mtm :l!l!'ScIM:~' U~~JlJ), 111 T~.Chil!m, 'Y• Ue!.'I2I, :'i .Xiif. E" u, J. J!l'ia,!. 0'!Ml. ,<l77.~S:rMI

~l.oo.2J. n T, C SIulil1lil"l . Q1• 11m, ~.

~6 ~OO~j.

mamrnnds,

hl,VoOlvC'd iin!r.;l1~$'Crrp.tt-m;],1repJ)cssio]l of nansUO\VC"Cf.

docs t'xi:>l in
,all

,\'ht:l'ru I~ is

rm'DJl

1'C"~IPOIll~

wh~~ 'Ih~:1h~n~ or [In in,hJr· \~'OUW :111m\!]'or !.h~ ase oJ n

be

:Jcti v e ~")I1Iy .~In g~nlll

flltprrJr/. 2:!~ 4S7 WIlli]}. 12. c. 'fi'. HlllIielL e1 ai•• fol'i'l :1O~, alll 1200'11. 13_ D. !lam,['M .. T,. H, 1les:tIlJ. (y!~l't. GP.oom.I!' ~es. :113"

1'1. WI. L G~ f6l;Q, P. A. ]CiltM, Mr.!mr. II.ts. 386., 107

t~9~'l'),

U~NAl) i"(SP<l~ ~'\()matktl"t;lJlmri[l:1 mil i,;db VI heR: mjL~bl'_~irllt\'dcd RNA {,!i~R1\1 A.I rnl\'l;k!:,~ .[l gCi1C'rH I inhillul k~n fir ~mu~.I,ti~i:~luviii ~~ll~illk~ua.~~RNA-fllZlhlllcd Ir PKR ~ IIml d~c iiltcrl'O:~l'~~ n.~..,,!lll!,(!' (IS ,. M 001
orgmlbmr:s Ulill u.,,,c RNAi 1'1,)'1 bnfh 1rru"1$!.'"rirt· 't i mUll mid ptlsunJn$l!ri~'ktioru.!I ~HclldnJ; h:fryr;;~

<ld;];p~i\'.; RNA imcrtcb"ISC' :teCil~ \!111kely w c.\is! in i

l~s:RNt\-.~:,>pn");.h,'c mcch;Juism ,Ml A !!Cl!'m cell srcci lie ,dOls:> of S in;] ~IRN.f\s h~ s re~ m~y b"'eJl ielenl i i~ed. flfld thcl ~ pOlenl if!! r~IIIJ(;' i(Jon~, ill the .g... l'lrl Ii U!! (I"';: cl j,:¥t;: ll~.'>CJ dSll!whcl'1': r I p;:\ L~I~r.:!I ~!lU~.TI1oC)1 ~,(HiI~ i'iM tdl! I;'\'~11i:1I0CIhmRNA .i~ irflpHell~~lI ~Il ~e~llJi...ncc~f!(.'I!;:HiC' dc' ~lI(lVum,,~.hyhHi~lu rn;m'lmiL~iil!,' ill

Hi. ti. II:. ~.lt1:" ~ M. !'(tjj'. III II:. !ItiIl'I~i'Thl, FI, H. ~lloIflnwl, ~. ~, '5re1'i!!reiDl:f. A.nlfll. !'lt1(,1iti«ht1ll. '~1, .2'lT ~HI~m. t6. It. WI', Sl"ierorr:- ~jI..6,l?7 '(il.OOn

i.,

11. C. tl.!!1"I~I~Y!oo1 e.~,~., rlie!Je;o t),tv. 41,9.olIllf1)' Wl05l', W;, ,_ S;~i". ~. :i:'eQ!I,. Hl. P,an. Fl, ~ Sch1Jl!l. De.v. ~ ~~g. ~M ~::!(!I05~. 19- .,.,,11- (;!m1!:!htoO~.~ (·dro..l~~de\I:;e""OO7,(1:3,OOl

ib\C'nll

Di.~cr~lik~ r~tl'Lcill:;i1!!:cilic '10 oCtlch rrOO(o,~l> I rI.:" i..:w,_-d ill Uti,~. w~u~c:h dC'IY'I.::;. dsR N A 10
~~hllmi,2 i~ 21·nu~.I";l't.tidl: d:sRNA:,. ~.IHl'1. hibil tu ill tl'.m~lluti:L'm or illldm~' oiJ"~r:H:I"l~(joI~ lIonmloIrl!r

D.: ~IDVO' ntl~~h)i~;?IJtion Cflll \')c:R.NA indcpr:J1J..:m. TIl!;: 3S(:omycctc liil'll1,uS Nt'.rtUJi,I]H!f((
fmd ~1li1..,1lliylimcs -q}~ilicd ~ ih~~, i;:;. IILlI. dock'd hy IlilLll::llJOIiS 1111 il:ml~IiM1iIllL'I1h lh~ td· RNAu pmJ1.!.lw.:JI.y lllJ J lh~ It[y~ llolaj[)J:rity !il'.r 5o-111.t'(1 ~yk:yl.l:l"iJlC;i; in [~~ IlIIlllIMU.l[J~i:ll~ ~F[l[).ll1~ .Us· iill (jrsr.!C'J~rl and l;and~1r1 rglit<lLS l}fI, IIl1d m
'·HI,'!" ....J

cd Is ~N~

l!;..:uonlf:

~.""i~~

:;~;;ILJCIl'!.:~

JUSi IJ.t.;t{"lr'~ 'l.,...io$'i~ ill :l rn~

(i!S 1oIdrt!Il, NlOn ,211 N'I. Hel1i3gJ ,r!t el',. 5tirmc(' 3m.;!, 1"13'9 (2!OO~,~ 2.lL .~ Ii.. "'ocI~r" L P. Wa.lsl1, T, H!. 8t!sill'l. IfI!l'id!: Gm!.'t. :n, BS, tl(:!n), 2'2. W~ili..a~~ ~ U S:a[l~,a~d j, [raster !oo ~elpfu~ di!irussiaD!c 5upJlIIliied b'f !!Irdnt~ from ~ho!! tUH O:H,8~', ~ LU.l\lllle'all

FiialtauID~ '6101.0(1)'Or~likj'Jdofllorll[l""MI'l fBjlC!t'I'I'hlp ~iK.W.l J!'IiI ~ eiirt!l:Jllilarilr~i'I!lI lin~r.I!r~~.or A'wmd l~riI

,~~tilf'lt !rom ~I~ ~~rw:e IrflIt~l~

R~A\i..Maml1l~~Il~I] .!!',i!!1.[)JUK'::i t'llI00d",' on!y iUl1~ .Dicef prokli~~, whit: h .loc,iJ~ i.l.U!'> 10 aJlr.l c~t()pla ..n1. Mtll11111aliilm DiOL::fis mo~. d(l<!ldy
goo,~

,pour LlliillChmllle'

~D.8.}.

WW!,'I,.sde ne:emag., org

SCI ENCiE

VOL 31i6

,2.0 APRil 2007

399

Germ Cells
'\'j)ell'lllle~

Tha Myster'iie:s, 'D,f'Sexu,al Identity: The Gie'rm ICeIII"s, PerSp,e'ctilve'

Anill'f!1algerm ~e~~sdlneJlenti~lle as sperml (If 1E!!!J'gS., dependiru;J! on du~irse);. So:rmlllLt:)ignals ~e~t Qlerm .r;;ells whelherllhey r-e:side in a mOIlle @rfemale bCldy. ilut how do ~erm ceUs inte!'J)rd lho~ e~t.e-mal cues to <lcql!!ire tl1eir !:!WI1 se.'IiIJl!l!lidentity?'A ~nll''!:iill as[J!llCi of a ~erm ',ell's~exul!l IIIU![Zlei~, tnaltl'ie :5p.jurnfegQi dieds:iC:ln its cil.!:!selv ~kliked llli the cel~'C:!ldedeeisien betlWlIll mitosis <lnd me~rnis. Mole"ular stucHes have be~ulfI to tea:Sl!a1part the reg ul!r~~r~ of i:lc.tli1 ,dec:isiitms. an essent.iia I s:tep lCwlilrd Imderstallldi'llii)! 1L1'Ieleg!!liLiltory lo~~[ of thls f!l!lfIdl~nll!!'nul Ci,UIe5tiofi ef germ (eU bio~(!!gi!l. r

dCcl!!i!Oll migln he' Ico:l1:-iid~10d <1\1 dhlinct fmnllililC cell i;ydc d~h1iQn between rn ilD:ii~~ alld!11c~i:~ whfii;hfii;lk~ pl~C!: iu b()1h
\'II,.';o.:~"'I.,.~:

:!()~\A~er,~he

l.iifL~il'lJf); ('Iflhc 1~1~psi~"PIWto~!:.;

dCJ:;iiS:iL1!~ .mil
tt'(filllh!i~"I~i(ln

~l~c!Jfi;·!il-, ,u!d ,"Y1~11~11.'Mr '11f dhhfo'l.\,,»' <l1'C o~n ,.;cr.-~'oojlk MIg:g!o>.'li'fliJ;lacI~C' ~lnii["l.I1l"hiJ' l'!IC'tw~n t!l~ mitur,;;Wn~iol1~:s, <lind li'rcrnllc~
Je~.tl.ll"""'& flf u)f!io;"L",

irr[:o the world of ,:>U"(uml~ity, :s,i§TIa~~r~ nsduecr am:! .:JI~li~lI~[)r of One t&!'(."it>i,tJllI (;n~r)' illJl~ ;J ~rmliil1i::. ;~ ~Ji.~ClITe l~ LI l.!'yckJ;, c~lIt"d ml,::i!.'l>i~. 1uiil th~ ('!!hCil:i~crllnmi1l!l~C'1n to d ~rr~re.mbtc, ;11:-:; ~'PJ;'ml or C!ll~. UJuri!lij;l emroryoo'ic
_ IH::tIUIIT: :>Iot~

~ml liJdi~ COl.l.ritm L ~wo 1.1rlmjor (1~n·r.LLC U!.i't'"ilOiDll~ as l.h,,}, UHW\i rrom an illn,.

vari~J]1 IIOOgdl~)g p;:II'iiw;~y III Cat·lrorh{d'tlili.~

l:.'hgmM·

(l,. 1111,)AK·ST.O\T J

UIlI1LI~ ki'lm~,.;
1.1T1II

nL,>crip.

dcd!!'iio:n:\. On~ll'll:'i1 I hcn:::trwc ask 'Ih~c: fjLte:;;· ~kntl5: qi) Wh,-.I!I'!fIil~l.II~ar [lldl iLIC!), if!.,Ide j:"o,cflrl m o.::i tlli iJIT>;t'Th U]~ir di ni;f1i:mi~l~iolil ~1I.'i.,:olJlCrm or qm'! ('I i,) Wlm:i nmkcu'll1rUI1~l'Chincry !lPVIi.'TUb 111~tn.ln· ~'[liOI' frull'i nl1ilo;;i~, ~(l 11lIo:'io.. Il,;,? and ~i;i \\,1131 i:s . ~ dill: rel:'lIj~iJLoihil1'bWl\';',""l.~nI1:.,\;:UhlUII:'i> (!if lil.:: sperm! (.~!,!. mHJ IlIiiIo.:>isiullcim'i:" dc(!'i~irnu>'~ We t1!::1ll.~id<Jr t~~ mOLIeRs. 0111.;idc~ is Ihat the SPC:lllll~ml end 1!11 imiis,'JI~,.;tGs,~$kci:oio:lb ure i\.!tIlly enc mid Ihe t :>11111IC t:Fig. J, "",. A{;,"'!m:liDI~ R1 Ill1 s liflttd~l. II ::.-c.'\;~l'll:'i fie :r.;:g.tllOllltlf iI:tdLlCIi.';:\, II ~I.'X .•:>f)\;C'~ iii:: 11IICr.[Jlil~ cdl cyclcllml m 'iJlL::.:m~~t;imc.cmllmil" th~gcnln tX"U '~o dilli:l1tnl]l1.~j"m as sperm 0, 'b~. The H~jl~'r i{ii.,~ i.~~Imt th~ u"'~,, d,"".Ci~(lm.: ""t'lC'lJ,oVICffioL'id by l~i~!i~~ roJilllltum.~ in~Ht:;; I fig. I En By Ihis IntlLlc~";1gcnlk:r ..nu~'lI~1!tI11

gcm1J cell.. fn,c~!;c.,: ihelr cell !1UlI!llb..or Li,~inili hc '~1!~~f8rd ~ m il('Jl!c ceil c;)'\':~c" ;If!(.~hcy !l1@! seme ocdl ,ydk: ill ~duJbi{lr ~all! cell Hmil'l'l(!n:~t.cC'.TItcnl. <L~ s~~~ 11'I!i.lure~Ihe) eel~, ~mb.J!ii ,on th~ !1'ICiNic' ed I ~ych!. \,,'h~~h dc\'CjOlnllCnl.
~ol~1
~I~~

mechanlsm

,!;lovcm:;; c !,ltl)' imo meiosis,

ill

..--~ r ~/~ IWI.' ..... s....•..

;n....

~F~f~/

bu~S(:':-$p..~i flc rcg~ll~moffl \'\'o~

PiU-a!-

~d\!~

1I1C' m!illlx-r

foom U~C'i)1lkal !Wi) Sf:!S tQ !>:itlgJ.;: ~I in both sperm ,~ldeggs. Upon lenilhm~ j,on,I\'\·'(l ~CI::; o r cill'Omnsolnes 8'rC' ~'1:ol~d ,;Wlll tile III~I, gcnCru1!OU Cilll be
,[<j

'L~r,"1,ltll!n0l1.0nlC'l!

melC)lh:l' 'I!nllli)l' IViS~pi,1l':lII!"'t to ~he~peilf,mI~9 ,t!l~t~t'On

~ ~

-----

!~! f!':.1I(1~Ci~~d~ic ;Lj!a""~,f~frnC'ifl$h: ttl ,mcl diIT.crcn!iiJi!iflll <lS ,"P.:rm fir eg;g. To ~I~!.ing:u ~~h i 'l'ilCL\\'CCn dlC':>C h\ e idea,. the mpl~c~lklr m~clul!!li~ms contrnU il1~ h(l~h m~w~i:o;'meiosis and i,\flcmlfC'~ d!c!')i~ to!l,G. !11~I:ill be d~crtl' i aed, Thill I 'U~~h11liillC
gordhas no[ yet been reached IQi' all}, o<rg;;mis.1n. bm progress has ~ccn n'1'tdle. :runa IIIJn~r.cc~II~la'$o~lItiml 10 ihi..I)lI.zzlc it'! on the hrn'ilmn. MDie-!illlar
RIl!g1lJlatorl
g~

InmlCht"d.
Ilhi"

or pmti,mJm ~al:ipm1mro~ ftjr ru'l,!i~", 'dl'l~ ":'jJ!.'nn ~t~ [~l{1 miwsiisl

:$;p&Jli'I'II

rn~'iosi~ d~l!:i:;ium arc dC!Sl:':~y cfJlllpku, w,hjc.h$UlmH~ ,1.'0 [J .li:ulti:.ll:llc:rmilJ.~(,(,L ool [ or !lCIllI11 ei:a~ rl.'~lIIllHio]] Ih.1il is only I:K'" gln:1l1ng Itl 00 understood,
'1:;1Ii11~.5~iOIiiS tlloou
,(iif'rrlil'l

.8

Gflli:'iiIl (elll Fail.'eS

Our 'lJIrld,.;:r.>I~lldug {li."rh", Inli~L[Iti;i'i(UI1t~im;il\ nnd sperm 'C'!i!\', d!:ci~ioll:S (JOIIIW:; frmlIll :SlttdtcS ill ~!I1J",>!:· ood mgmlu:;:;ulL'> lhe m

~ Se:Kua,~ ~1:!:emUtyiill

nematode

C i4L'gami, Ih{~ rn.l.h Ily Dmlake

~I quick

(jell~~
:!

:mpilfla mj_"(Ju~gtL~,{~'l".. anclIlHl·mmL~.M·u_~

mil,',(_~lhl.'1. '\\Ic

!:I~liIl1 ~d~ Jin~·It·lr~i:llil! w ,1]:0 Sj'lt'rnl OF ~1!g? One imporKLI1L nnd IOllgi'ilJ)IIIlHllg ap:llm~chm ihils qLK... .;JLUJl
\'\'1'1:11.(j,iJML~

ij}r

leek

!~lM}TI1C

'dw !"J..'~ nlol'lll:ul:ilI' ru!.tu'I:lII~ i~1 11m l'olJowillg t'!ml'.r: I'l'gub:Lnr:.1 of milm;ii,,"- uf l'11'lry j'n~f) m),)n~ils.. and filmlll}' ol"iJiIl'tnm·

I~.~ fOCllSi.I(,! (lUi h0W ~mnalic lis:"'l[icsinllacnce ~hc sC'Mlalidcnlity otg.el'm cells, hi i aice. I he qll(s~iml i.~ 1milyelPtu;]i'>OO 1IISli. r
ttl 'I!...k wlml $'t)DI'l;l~ic '"tICS ,~in..'Ci .!c!~m ce~1h: 'ttl ~rnai~ilitlUl I"I~lj1~min,t)~i~ nti!dtl"[{i

tWlkm as s:!)O:!il'nl ilr egg, Genu (;d~5 iH\C' llKL~IH:lin.edin ,~ s!mc or I;m,~ifl'cr,;;nlimed mi!mp.," dh i~i(111$ by
fil!)'.!. Germ ~eUf;ate d~(i~;ons> As germ cds mature,. they len~~f meio5~s:a I'ld din~rref'l!tia;te as spenTIl orr e-gg. Germ (e~~5. IP~pii)5ed awe to be liIf1coll1illiiiUed~ree'n~,~'fma~e (hil~hia), '01" lliia1e ~bllle).

~L"'; IUIi:i:<ILkiIUilimi:1.'I'11 hi, aU'It:Hl:g the' ~ir.;.l$igill.<;' IilfSIC:li.LIi11 !.Iimorplli!'m ill the tnutiitC (H':.1ry j n T~tc 1ll0lOCWlir l'l[lllfC :!'>Ol1Ultic :>igm~lj lg dCIlICIHi:> Oil tJ:h! o 1'gJl IIIi $n~: ~ h

or

ii~)11 )PfLHlw~}'

'~H~ftl !f!U]'liI!!$ Medl{a~ II~tiHM. Ilepa..~oi~ Jill Big· ,hernj~tl'l", lInn.~~t~ W~{l)Ii.~n·fA<I~SOn, t.'I<idl~m, '0'111 r'::I.lh\\i:1l'~ 's.~",O&" IJ~ ll~QI'{otn:l HW!1lhe~ Mll'dk:.:ll lIutllLl~.I!:. ItlIl'ilti!" rnoJot'ul~.r l.l.idr ~~xm~ ~n ~~!ln;Jl~rnl oclll~ :.md itullwtllClc itkmtit),. UI~iIiII'Irnslitm~ tlnd [)~flroonl: of 8Io1og~,MaSQ~~UM~~ h] lhJ.:i 00\ klW. WL:l ~'a" c Ii~cs.nma :lind Ib~tb\ IlUtllUtl!! ar 1'~hil!:ilDID', C'.JInllrid!>l@,o MJ\ 02 ]42, US,!!" On \\ .hllll lit"" d.01i,\11f;lromn SU]l!ID.tks~~l!Ib to "ro IfillDifi ((i!I~e.~J:lrnclM~ :~~~tI IJ_~ aiXdrel'lerL ~·m:li~ jeli.imbr~@ffi:!u!du mn:lJol gcnn td~ ~t'Xll~'1 idl'l1tily. A I}Jioor.i"I.he

g.

j n lJl'Ut~IlM'(,~leslcs ~3}, atKi retlumr:c m:id s;~fla3i11Ig in n'licc (4", j). Ilor ~" :1mpi!:. The !Common Iheme :lcro$s~hc~'c ;ji,pc'C;i,csiSI]~lt ~hcoonm tli$C:S ",.. d~-Iru Ie!.! vo;

~cqtLCito.'"""'I~drpc RNi'\-bkttljll~ W!h11.;:ins or Ihe \\ iJdy ~",l~~ei'\ ~Li !)Uf ii"~Dll1ili~'i Zlnd FBFl Uiul1it~yin hoth wom~, m-Id Ilies ~6 N). Itll1iIk~'. ~W(l pur p~olcillS, c811c-d IfIlHlll il.i.ld 1\~In:!.ill\'!; bClCll imp~iCl:UC~ Iri I m~lint~'lIt~tC~ !!,:fifllhtc Slio!tn cells. bul Unua or role h,JS nDl ye1 h~ll ~lll~1nrl~tl(9). inl C i(.1~W;111,o;. fill?' kC'CiJ$ ~Inl ~-c.~b um:li.IT~I1;."IVl~b.1]

or tiPJ!i:'itk:
llllkrmwn,

tubd ~li.Yi,t!iug

mitutiClllly by

UI~t! ~n..'I;i I\.".~",io:i~

!Bg~II~I(h1};tlf

11iIR~A" lJ:llU ~nl"Od~

or

lXlllh'l"lltf}, illw [hl! nne~olic Cl~3Icycle and .5~XIl~Jj

iLl!lmi~y fL'i.UJ ~ .~Il Uil1l!. lBlLl m!..'ch:.m:i~ln.~, J!'lrgcly

400

SPECIAILSECTION
[tcglJl::tIorl'l
!lUll

JiU'CCl

g.~nn cells

inH10the

eell Imil"fl1L'11CY ~w c.~p:tC~il.y I{II 1'illpprn1 di 0:(':1....

1I1101C'ClIlnr CXphm)1li{'ln I(lrl~nw Ihc:sc ~.wn, ilL"}'

fl:;'g~1 !alOl'!'i cmllml~h~ur wrgcN. RllRII~il1lil!'lg 'Q!!i i!~li(l"lS an,d PIW!U~~$

UIlJ'iOliC ccU cycle II!~~ ~lCt,.'Ill1A1l.lnd ~k:linith'dy

h~ l'IliclC' and IlClll!lQ,,)dc~., A ~lO\'e1 cyt~JJlsiInk 'Prot~n .. ~:Jlk\llSnl.t\~ ($til'ml! .. t~ by ~"'I1.hilOic

;j!C'id4i), lll'~'!;'l;'n~~ ~nitintifl!1. of !l!w,iOilL.,in e,,~Ilf')'mli~ ~!W~I:;'~ ovnrie;; Un. In C. !.'/4."l."''!Ilt~. Ihrcc '~m,.iJly conserved cy[QI'I,Il!~mk: pmtC'in.><:,wocomcpUsh this real 1,"'"0 RN~\obin~iin~ proW'in.s ,(GUJ-I qU;o1ldng Cliud « ~LD-3/Bic'llldal ..(_'} and

all Ciel.! 15b~c;>; dlC illCXI g.ertCnnKm i~] ( l ~ I. ~,~m'!l' Iht;s-c .~~ 1\ rcglll'ltal;l. t!11C1'Jll1;C\\ ill!! ~pi.g~n~lk ~gillb!~M qt ,h.1i!' DNA !~'(:l ~!1fl,"Ih1-.<;'
.~~! ql!lI:"~ioll IliJ1IC~!

C!lt~11 I:on Qj"

.lnd au

~lltoe!1i.~'illg ch~.~~~ltg~ few

I'llture :-ludi.;'5. A
M,@I~e~u Ii:lr

llirn k

oiIl1!d SiP~r~/hgl

ll;ie~.wll~I!'llh~";it~ll~;M.eiQ:ii:s !;I~~~:~1!ii'l1S

The in~r:il'!"~icn:M~d;11to~nor gel'll» cdl ~/1\. I",,·c km,g been n~y:-tery. il'lil~n,"'vght), dm~'!fn rrC!u~ w'Ori: irl, 1110dd {I!:ga~]~!'l~n~ have !lfl\~ OI,~C!lOO a t;!!.ut'~I.i~inllcrock i~ l.h;l~ l;l"J~' l~1 Pick 00;1;, O~Ir fi.rsl !,l! imps!.:' insldc re\'cn~!i. M~mC' lIl1.C')o:.pcc1lcd

,I

OJI'!,<;\'ilC1"ll ~md.u hos~ ~CW ql~SLious. Willi W~lh ~fl(:d~lc;cgIJ11atofSol'the I.ll:iw;:;i s,mcinsis r r\,;gul:,'l.Il~H'I t1omurmh; ~h,.; nwk'~IlI:I~r ilntll "'~'lim.l""g.y d~i~ ion::> ill hmld, tlll.i: ,(~<lI'1 RNA u:;ctl.tf) OOriirol germ ~-.c:1I 1'111';,';",' nil" bi.'gin in I!:XP~{~~ their m:o:I";C'L1.1::1rn:lai UOlJ:;.h~l)' '''It'dtegy "fhl:' ill i'tu~~ id~lj'Li,:I~t:'lItiolll (II r'~(,n'l;li 1'1 rC~III~nor::o; of will 'cnjll~~ril)l.iLJ!ml.~rc~l~l!Jlt~, Yo hidl liia\'c :'0 lin" 11;.:1."[1 1.:1:1 LIl<~\~. prow 11.1 00 t~u: ~ k\J-y'J .,1\.1\: rI."tI the millt\.~i~!rllui!J!>i:, !d~"\'!'ii>itln '. <:i L[) pmIc.t! proli.:ill ... unsversal regiLl~:JIIJ[5of~\:m11 inc ~l:im"l ilil.lcl ethers 01' Lhll' ~fI,.;r1ID1 I;)gg. 1l,.;~i:;9'1.m pm~ l:IlC.r1il (XI b:'?\Vhat uhout o!in::r g..::r.mcc II kul.: contclmk II C{'IITL~,"T\,OO ,n~t~imuj~m. {c,g .•. FOG·I} ~CIlI sUPiJOn 10 ImJJt12 (10 IJI. prolcim;'? Do However, m.Gr,e in-depth .$lud~:S. rCH~<JIkd III!u[ r.cglllhllOlW IiIke chI: fOG und ]t';~idl.lUj"', I.he :spcl"lnfc!!!,: ,deci:;i(']t1 hrw,.; been id~J1li:l.k:d I]] C l'i(_'J!WL~. M:GIlOIlm~hl,~ lire I their V";IL~bt".lIiC 1'1i'I11~O~U_~ a~~m'Cnl]LftJ~ gcnn the GLD IUld !FOG :r!Jg~llmon; IJC~tIIQ~~l' inil:u· cd] .i1nli."i. or 1m",.; UJI;)Y oor..1l, oo·o'jlh; .•d fllI Ihi:; ~JWC both d~'1i:'i~~{m!'i. cn1l:tllillgmll~'Eiph;, lrrmk:~· r rOG·1 fii:lniIlJ:alliul1 {,r~m] 1i11";~ 11m.] rOG·l, ular linb bctWlli.'1Il, 11.1('11.1UJ), M:o:,,,L mJlahlc is ""I:i:i",h bot!'l ",",";Sidl;) m 1.i.K: end or:l COliIlJ)I!..:X task inn~lIllatudc;:;.lJy ~onlL:: qui rk. 001" c"ollil~iou'! FOG· I., ~...lidl aclt. lJ~ lil,.; ":-[Id Lhc p.aLhwl~y And: ho'w tit, human !i;L'fillil (:db ~ll. il1l.o 1.lu i .. ~ ~dl~I.o'iy~l!dw"my {:on~m~llin.!il~~m'l t:dl M~:>i b U3~.. f'OG·! i~ .:1. IIOl1ltllo~ (If tilL: \(t"rhtlll.~mz :nltJi it~ thQJf!tfim.1 lilwiy '10 be 0I.1C ~}r:i ~~, mw.L l,icllI rc"! The l]JllOVl'Clb 10 Ih~':'i~~lll~iI.' ,tI'lbt.::<I1.itlirli;; or !l;Ci!TU ~U hiology nrc' !lOW withi!l 'C'xpcriimpor!<mt !leW'~::no~" FO(j-l t\~nl:;: QUlIO fllT~t C PEa· 'I,~y !opla,.;.mk pOIY1l(rcDly.~~niil<n ,~~tl.~ln'!~! li1'ii;'~IHII,each. r hb1{~il~g' ~rn!l~I~lifln:1I rc:gllbittllr, and tl~erl!ffln.~b. Ihe 'two dC'Cis:io!!~ in .Il d(ls;C'-dC1l~rlCl,cmnm,nl1cr,

the C'a1(tJ~Hk SUbLII1.iu 01' an

ll\:;;:;jdLIIo't\ o dl~ ~

CI1")'I1lC'

Ihal add.:;

3' ":I'ld of :II!IRNA:;" a j;lol:y(A}PI'l~yttl!.:r.JI~~ ca~ktl CiLO-1 U!l. Ap'~ ~ilil~ ,COliliTIOn theme i:. Lh:1l~tbe ,1l:~II~h~irl> :111: (;:-'tori asmie, bU'I,wilhmn ~Lbeuer 1IIltkll'Mmlcli"1l or how S'J~J\,S fllni:1 i:on~. ~1 hi, 'll00nUll[LIl'J: Iu
,ud!.ill~ill";

or

or

ot.n

or

ft) 0-1 homo]og. ,,"",lied n~r: rH\~mnlil:l, B. l00.1IM5~!PlI\atl!l:~.a4!.1. ::I; .Nt W~~il!; tim •.Nt:t~IN;f~:~!5.~6l (.20«5). preSCill m (I low tC'vC'~ nn~B pm~rcs~in[1l lllOOllgil !'U!C'iosi \.\lhC1~ more [~tl\~ndil!rt <I•. ~ KrniJOYol !i! iii, l'iI'«. fflt!t. Arnd, xi, ,!{,i,/li. :]]03, 2-\7!l ~ f'K)'fn1I1lII.)' dHTCll.'nt!;)!C' as Sj'leilln ~ !lC'l!IK~~i}'l!l!l't't(:1100\60.1.. Ibnned imo ~~OO)l(oCS.nCilCr(lJ'C, .. OOlh FO(i-1 and urn. ~n Dm,wlpMla, Ihe FOG-I ~lo'mo~og, '5•. ~ Il~~ t!l ai". Sc,ir~ ~1:2; 596 ~<!oo6)', caHed Oli'b, hillS been imrdi,ca~d in ·C'OmrtJ~,(If r-OG-~ IwmOiC rhe sperm f~!IC' m ~flc~:.'ip.;11SC It II. llil. A" C 5pratllim~, l'JI!WJtl,p.mi!l'l!' ]j,;M, Z.II,6:1 (:1.9111), bath I;)lll:r'Y iruo meiosis and g,,;rmlii:k" sex de... oOogc.nI!.1~I~, 1, <'!. ~~Ile~ R, ~an!l'!1!f111t~veI'opmt.Q1 US, 6'9 L Tlhi: kllO~\:h;_~lj{' of Illoitrn ti."S lP"',"T1Iil1~ '11m tl:miillmluol1 t2'IJI. ~I b, I[m c~rJy 10t'Oll,C'lluldt.:: ~ii~t ~19~~. I:h ii> dll~11 IliIL!ch;~n_i:.FnliIa:.-o ITI~1~1:n~~tv~-cl. . l]t'Jl:! bll[ ~Ji:milA:~ d:L1i::ii>im~ in 111tJ11:~~ ~'lIUb:.!..';i; ~" i~ 8.. s:.. l. (fI!!end~i"! rI 1fI:. ~~ '\117. ·MO QOOZ), ! :;:Lr.i ,:.;i'b~1;) ~(I J.l:;k '~'ht1JllL::f gcrm l1dlb:, are tllmwfl>i6J~y lh~' s,ilnim:J:rilUl':o.kUi.'!' knug. [mkl"t!, II nlnk~c:u~:Jr ~'. L Y. WlI, f.ii, 0iM!'!51, N'. A. $a!l!'l!lfl. iii. A.lttll~ i'ffil. MQI. iQoomlillHrl 'Loi:llllJir ~xu;11 i[klltit~t 111..'11 Ll~li(Jn q iS~I':1I~!lYilu wilidl :J :g!tDilli ~11J Sl':?iUIJJlli!'\Il!lllwr ~p!H, D.t!~. [-Pilt! :Ilr:1d 01 ()flj:fl,ml~"mm~~!J~!; ~~oon IIhm l)Timmtc~, n~'1iu i~may ,.;xphli~] Ihc: L1nnsu!ll ..... 1:.,]1;boen mldro;s,t(1 by ~lImil1g k.t'y Di·.~I.I~mKlTh ,on 1,0- ~. ~ lbl;fll~1!Il~ .r[.I1~~ H2~ l~ll or orr IIlw in. w6li:da !~ unnc lII:>i!lg gcnell:C trick:;, UIII:IlOn; lyp:it;~.I, of ~)ul.niv·c g('Tun~in() M.::X dClcr· ~~OOS~. i~l~ :h. ''0\ dlll~ ... '[e.g •• Ri\;A·nmxlimk"d il1A!l:i.'t!.:I'!.''Ila!C). TJl,lC~lb.'iWcr l~ IllIhm!j,rm gem ....... [)l'flwpM/(j u- it. E- iB;!~aJs ~J !If. ~ot" ~t. 31!).~A~O t~OO6)' !tn'll ill~ imrnrl,:ml. In'l'f,~.~i{'nlion~ Ibl" 1,l'81.CCUm.:rr 'tlml :-.~mll iJr.mllay L~~~MIIt'!: iltiJl rCll1!lak,:.;ln;)kin~ a 1:;). ~ OCin:iti!~, S.l. ~1Ii~efll;le-",im WQ1rn~. C, ele.!N~·~ ~llli!1a~ Ym11l;Jrn_lI. '~.!ILI'W(IftfI!B..r~ rtN~.\t~rm!J1!i1)101" t:;umcrs in mmllln,li Ibi.• 1C.!,!1!,S t:".1n oc swil~ht\d ~IT[{lI ~'TlllIlflgcnQl,i ... 'I J 6)., DII!j. ~DrJ~t lO. m:fuI'taI(mti1W!l "L.1I3.1 •. Sri, IC'ml \\',l' d~"t~lIgl.l;i.~1lil('hvt'\~J'I Iiltl~ op"" .md <ltluilrIHLkj,; I~rnrlll,.:in~ _~p(mrlll"allhl~_';WUlil~I~!;o'lil l3. 11:. EJJl$.T":~r!d~ ~r1I tllMfiBti&l;, C ttrijiMl' RelGP~h ImiS,~ng. 11.1Od(lJ~ M11,fmll.h 'l!'dI~'!iL'I" o~w a[!l~hillg Ih~~L 1111.0 ooJ¥'.nlC~i~U51" l\':('!Iab~}'. mn ~\:'1I5 :Jd~lpl y Coonrnmllit1. Ed, ~lfflmfl&OO:k. ~\·ii'lM·o!lilm1J;OOt:.or!J, ,2m<l'~. ~Iw l11:i[a~.i~'DlIIcin .. i$ :mll: :;pemlA~!!!~ dm:i~i.o.n:.; ~h~ir ~L;Xllilll lille :J~ uoo.n dl!.: SlJIlILC~jlllLC d~L'Y J~U\ ~ :~I],:lllS5fr.'1l1~m~Qtiic.UI,.2 ,1. 101'10 ~ nHN1i~ ~ Wi'ii~ms" 1. tiiml!.'ie, iFI !iullwlimtt;ll arc' Dhr: same aoo Ill!! (liher thin til,;:,)' are the mitotic tdl c)lc~e:md i:lll1iC!' andtlsi;.. (I71l1l<11 [oolitill~ ~ iNiiJ ~n~. M. II. MiiI~ di 11cn:In',~ lu r~nKl\odc, .., lhe I,WO (£C'Cis'i{IJ~lSF~i.rn ~C!np.or;J I, i;Oll1cidc!1c(: U i:lilc~ ~h·c COI1!leciinn N. Sof!ef1~e!i!lo 1,· W.ID, ~'erl#1t!t Ell:. ~.GDld S~rf!1g'Wlior Ol~t \(lo br.:' g('I"C~IUI~ by Hl1:my tl!' ~hc S;'Ilnc b~~weel1 ~he IWfj decisifllNl; tmt SRY)i liulc~bfl~1 l<Il1>lI!3IMj1.f're55, Wcodb:IlQ'.r4Y, :l1Jc!11,~, tll..:- umk'rlyiing uncdHmi:<lll or Illgk l~tdJ' rcgLlJlIi.(ll'S" which '~ll.iIJI:1c ~u""d to ,'l'Ultp(m l'S. 1'.1.Ch!l!l!.,Ii. $l!l!(l:al1, K'lmI!l.'f, ft. E, EUfIr" li'e\I. 610', ,U. n ~2000)' m{),dl1!~ LiJ'li I fOf,~-1 innu,:n>c~.:. th ~ tl,\\:! rel;'l~I,lil~:::iI1 'r'. 16. r!'l, milri" 1'. K.i!J~hlmll. M. 'fa !!'@'!O~ Mo:lt.Biol. CttI:l.1. b i'I. pl'Omi~ItCUi 'U1Crlh:C-Il'ter..lill!!l Iro~ll. 1~'UIc~ 11r.:"cV:-ioIlSy :~ tk)~'-dcl'1clI~k:m n~chllnism, ~147 t'Q;~~~11 .. ls til!.:' l!l.'!.c of RJ~Are~lll;-~[{)~ l)ro~~jl1 s !'Ill which cou:ld be il1llcrrre'.cd ",,0;; sUPWior~~ug n_ PiI, I(_ ~:lOO6}, IBar100, ,1_IKimiltE; '~I!'tift U5, 2~' mmJ,;:l 2. low FOG-l ii1ld IIi~hl l;'O CH ~nliglrt U.'I OUIrt~J:!,Iei'lll cc~1 ~b."S, \1VIl~ RJ.'<.!A re,guJ~ltiou C ~1~}" of is,'liscd for liu~ l'Cgulm litl1. in ,g~rm eells remllins fI oIT'cc[ distinct mRNAs. i);!CiUJSC diIJ!!r.c:m!cs, HI. ~. '~i~k, M, M~,Ima, ]~~" !!.ril!;l~ j'ci<'f.I(~ 3;11, R51 ~~)" III bindi~lg :l'I'1:'hl~LY,.or ~~Ie~wo FOG~j leveis rnmlClllo, spccuJmiOll, om;~i~lem is abm dll~cn;:n,c 19_ ~. Mein:d'~ I" iii. llidll1er, lIlat !ili.._ Mor" CfII8ia!. 2, 5<n Ildghl ~)l'b"Cot;:q Ihe $llIlIC u'iRNl\:;; iill difll.~~I~lJt ti;)j~iono:l'y,.nu tell:; 1~,~ll";l1n m '"1!!.1!!!. is iI~lIr~iy II ~~U" wny:;;. fi.lr CXml'lpJJc, liJwmo;,ing htm$lm~(I'ti m htlfi..~;Cill.phl.,''1l0mClltllt ihrn~,liI!iI~tl.ll!: p,;,.,-~~d ~in,;r I 20. J.-It. HlI'J'M. 0: :5; ]oWtllfl. ,rl\i!I't'..IiJp.mi!l'll' 1lZ~. 21~~ out· k \'CI hlil. iIJhibil.in:£!, i~.:1,1 the ,mhQf_ Arl..~11 rl~11~i4l1irin.Ht~L1I:!1!i~m:!I[ LI,,~ ~e..d ,tlj' lu:t~NA ~oIlIOO). 1II. ~.; i()Iiv~r,Jilt'. lire!'; Cp"l•. 2:1\l, :1 Q.ooI2j, o~l::JbHily '~'f Ir:!Jl!llblicm InugJu. t:~cilitatl!' lim I 1JJI~dt,~T>..101Jl1di.ng or IlhL:l ru'hi~.im!Sihip bt'~wl";!!n the hvO <Lb_ll~is:imls.ruqLl.il\_~ kk'mii1iemiml or the .ruv,~I_ .I.nlde~::d, RN A .tt'gu~a.!i.D.rs tum::! nl~!:l mrgJL~!.:ii0 r hmh FOG~~. ,mel 1"0 Cr·], and m '[limed mIl n"bc cnll~coll l~lrl'l1:liIiHliUlill.g gt'!m
~11. fi,;'~I1<lhl" lI~lch:lI~NI

~!~'la1scd['ld{l:!1<11C\'C'~ (/.?, N~. I:(){i-:O. on the ~ ~llh"'l" lW1i~d.beers an Ncterm~i1;t~~11f),til·~),!liro.l.of th;;:'!lO~lrJy u!~~IcI';lJl(loocl \,~;!dU'lle Ttlt.!nn Ill'iltei~'l'l.:ilS I'I'IOk'>1.:lll,tr role lTl Cftnt~t~IHrlg J:lt.'fl1!1 cd~ ktCldf1g dth.~1! FOfi-·1 Dr Fmi-:;.~~liIl cclL~ tllr..

HkJct~,!o crn~lroJ

gC'JlO!

eJ>!ll~klll

at a po'S1"

pmn~lO •.ifl!; ~~~hJ!~i~ at JI;'I"I\ tc'\'dS~~lll drh iu_g: :gcnn ceUsumo Ihc sperm raDc ;11 hJgh levels I W!, ~s this F-{>(l- t ~l~9i.lh}'.~.bi..arre ~fll~ititli1 ~,pc>cHk to IIcmOil!ode~. or Ihc h:u'bingcr (~riiI . !nOile' 1m i\'cr:ml rllC'r1fl~Il":I1Vfl'.) Iql X~jm'l~ij,~ .. the

R~'r.~i!l!'J'lCI!:~

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1l<"500111W11!', D~1t Bllll: ];111, lOll' 019!}~)_

.(;"mMI)' 1I;i}.ln anlin~b

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~v.mn~. C

Creg<l\'l~

Aeoe-ardl

milnsi:;; when

or

~ln

or

WW!,'I,.sde ne:emag., org

SCI ENCiE

VOL 31i6

.2.0 APRil 2007

401

Germ Cells

[Male and Flllal[l! Drosop'hila G'ermillin,e Steml Cell's: 'Two V'ersiions IDf: Illmortali'ly
M;illrg'H~til'. f!!!U,e,rl and .A1,~an >C. Sp,r,~dlilll'I)Z Du;mJpMlc ffiliid~ '1I"1d f!WiHlle!J"iilrml'im:: ~t~m !!:~n~ (GSC$~~re sustained by I'!~'oh~ o!:In.cJi tregUillilt~~ Ii!"'t~~y~ w~~,~ I;QIiT1~g~ j)ri:f1Jdp'~:i ~i~rl,!'~ ~I~ m@~~Is<J'Qf tlFld~!};'im.gil!'\l] m!'!mm1!lUo!lf'l~,t~m ~eU~ D~5l'1lte ~t'ri~i c~~lli~I~)r nd !Jl!lletic s;mibritics;lli'lat iii!) a :SUog9~t iii c!Jli!lmO!l ,~'!j'glJ,llli.o'~ilJry ,tuig]l1, however, I!!l~I~ ,~lI1dr,~ ~ 'GSCs a,~ d~splay ijm~l;lrklnt dmeHmces" ()omparill1~ lih@':!jl?'\\lQ stem ,t~~1.:s ~ l o!:I 'l,hQk niCihes in d:tt,ail ls ~i:hfi!~.~' revNI h(lw a (om !lPOIFi hl!rit~!l@ has 1Ml!!!f!.i!!dilP~Q·d kI the rid 'I(l di'frering' r~CllIJfrertJ1ients. ,1' nnalei3:ndl (ernste 9J.. me~e<pro.:!ilIcMoill. 0

n'lb of BAM. f.l( ieN, and BMP Sig~l:ll. in malt'S. where I~AM :1~u;1 .OOrN Illal'C!.lOIl t(1 cml mJ!!lI:;r R1~i;la~ i1aili;llt~ mi~\ic eJ-"t d:ivl$ial'L'i;. UMr siW'ill m· ccp1:~Il. l~ "'lillre.qLli~ !ll l]i~jJl~~ GS.C::;~n~J lrl reJl'~'~" lltWf [!'.u1,':Ci!·iI~li6l'. lritn OBa b C"~IWC$i~'\1 iF! ey~'"cC'l~ :lnd the' h.uh, ,L~ld bam i~~('I~~C:d ~~ ,~'lni<tll)I~SI!i. ~1iI> ~\~H a~ GSf.:S IFi.l!l.m), E!"I\M Illl [se"'rm~S!ljml can be !o.~iC'liD !nI11.:: ",~S(s;, 1!11h~1I' ' t11i.!!l j u.s~i1JQ!1ll,~i~1~diff'cUC'!11~;-LtiQ!l. i~ <N dOC'S in 1i:1iIl;!ll';::; (Un 'Ihc rrilrtl1lry Ul:,;k oj' c."ommHing GS{, Mlar.. [;Cgi.cv.'111 iiii m!lle:. i~r~rfiJg'lII~'tI IJ~ :11 d iffo,;l\"lIlt ~J;lli:d, 1I1~ {'Y lID!:~I~-~i.ki: ~i¥mliiJtJnrliir.,.'tl r J?D I i VI,5). lJrD l::xpt~~d by I.IlID eells 3C:liv'm~:s rhe Janus kiumsc sigm.l Irum;dlH.'Cr:> urul ,1.'{:~i\"311'lL'); or ~1nU1~crip.ion Ur\K·S1Al) ruth.
TliU:i

in~

,IL'e ,III S!.lI~I<Illl!i...~~y dif:lbtmt

:Jdul:l.:'IW!Il1 l't'll~ :i n \. uw~_ M:llly o'!lllinli:mu; pmdul"C':opm'ln nnd Ll'gg.,> I~UOll.~1.~!JI1 thl!ir ru·pmdm.'L.i.\·!J ii'o'Q;, by{;J~s:;;ic hi,yh.IJ,lr'll!JIgh,j1lJl t;k'Jl~ ceJl S}'$~OOl~.although the ac!ivc (Ofnli~Uli{ll~ of ,germlinc ~tl;:m cells ICiSC')lnO O!lgcn~jsin ::!d\.~I. m~Jffi!~'!II1h;"~'o!rI!Ji!l~ dfl~~\I'\d. Above <llt (i!lt'$

~l.C!?'Cnrlll.ine .1:' C!i.lt:~i ng ,~S, l rJf ihc LK':>I models tor MIlIt!~1 ~r:I!i. IllI.~ bml.a!J;Y or

1;lSidnatc <!~ the l!1i~!l"llc mrM1.iJb,"~~k'lllCtf~tc.mc .~~C~rilulalillg a ~p(,'('ie.-,,' C'mi~1;' repcl1lnirc

ceJlubrdi\·cr:-;hy.

tl~.'-~:: n:milll!!li ning dli,.~ on~y eell i}'f\'i: ~i!I~lblC' i!.lt'l~nril~l!lhlg 1(:1~ll.,; nC1-l1 gCll'i.!'r.!!~ i~1nrmd ih~'iI'Ch~

fir fir

WtLy in GSC~m 1!.-pt1l::ify ~II?JIlI(;cll :;clf-!l.:."IlJ,;WilI, WIiK:nl ~II nml.;; GSC dh'ick:;,. lh~ d~ltlgin!l.:I," rentm.clnll!.:n[ 1.0 nicht: c'~U:; .~fa mlhq;oiF~~Il:> j,lIlJc. tl! inrlll!l conl!1c:t wi ~h tl!J,; huh 11IlL1.i.n'l1l stem HIl~ uons mid by ~'I.i't',:i\' m~,:lotll~1~i!l~KtI'~t1l1l1. I'ai111'0 cd] ~dt..:nti'ly. whi:lt: tli~ dml~ilcrdi~"P~Hcrcl ~\\Iay L slimLlJ:~h! ~~[lillct'nS loc;l~oo even om: eell !..~N'f'M.:rilm~",'.Saw!..~k~r :o;it!-fml ;md inil.int~~LJlir· 'contra!1.I, rC'm(l!l~GSC~~lilvC'm) lIi:JIllJt[iI.'[ U\\'~y [ra:m [lim [lithe. Bt:H!l l1Ii:chlJS 1~:c.lUiil~io!l. :~rn nutonomous rcq~uir~ntC'1I1 ftlf Ji\.K-STJ\T :!i.iga Il!>LI IlOlII.'iC :SOlJl!.U ir stem eel I:; w hose d:.Luyl;ll!Jlli Cfldn~ c:arJ;1 germ 'CCU3..~IliICr!1 ~H)' t i~(':s, l'!i'diu;g U. Ii); Thus. hOIJI I~m~C' al~~1 felil\;t!c ,. ~lkhr;5 m~:limaJ~ USt':> IUHJ :'>~.i!l,ub.oCa sym:1 flue fI~hi;'J: Jist~n~d\'C' O:<liI\!re~ im:h,td illg~hC' r Pll"'~rk J;! !v,i~i~PI by .~~!:n.i~.hlgO«ICS)i!(l;'! 10t;~.~ :';;P¢C!f(,!~L"fl1C': ;l·n ~g~~~w ~!ld(lpl;lslni,(; relic!'l~u!n like\,.. id~ i,~I~ri.~<J!$}'mmc[ri.'[;:~ll}i ~ ~~gujl1, but ~hc!, ;c,I.yon d~ I1crl:!!ll s igll<lb Iftu d~ b: lit!~CI ifill. lli.JIri~'g mit~is.. Male and t"cnli.lh: GSC JOlugh~Clh. bl(fWfl ,~~ gOil~lI~blil~I;" 01,11' CY,~lflh.r.. re-

fir

A Till:~@f Two, N]ch@s

h,,;:sl-Uli!.I~n;lotno,,~ ~d~lh '~~)I.':mcells V, .h il., k:.;:-.[l~] 1'i'~1Jt11 their ~tli{iy b ~J~~:i mero.rt:IIlt'1.l or lh~ mi~ro~~lnf~ronml!r.JI 'llhl! S~~'11 eell niche)' in mgLll:uin.g dceisiens ~t'IWmm ;sdl~rrllJ;;w,11 arul dilferC'lni:lti,cm U 5t 1h\O eb~~Grv.n[tm ~.Ini11{;:lf-rcrac\'\ll:d depends orr. ~hol'lI.· ~
Of'(l"iIJ1Mhl

osn,

.1~ '~!I"rl:nll.'l ;Iln!ong Ihc

p!:.clh ("~" lll1.dc;go S;}'~IC ~lmnOlJ.~ ifloomj'!.I~I'~~ di v is kliit~ h) g'cmrolc !JflJ'logOiJS: l(.-ccH g.Cf11~linc cys.~ f~:i~. and ]3). I\~SO. inrUI\'l' j,;CI1C:s. ~.A :11:'C Ile{!U ~ro;I [11 ~{lmm,on, lnc Iud hlg {Ii,,-i, f1((Jft. b~PI, :lind M'~"', P~liLC U1Cse:l.imHfUll ies, male .utd (cm;11",: (;5(' niches c.\hibi,l cOIl:;.ldcf<il'id;;:
)1

~I."'.

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'~rimlil)

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aCCQUm

~igraill:~ rr1.mll U Milprmrti:ng 1lI[c-h!l.~ c.m tor Il1nIllYp'rorc:n~!.l8 'Clt M.(nn ,eel! pop· UIal.iolfi Ii, iJ1eh.ldlil1~ th{' ir abi Iit)/ 10 I::xpal'l.cl ;L~·
N.~·tra:I'l~p'I:m~atil:m IU
.l!. SICilI1I

r['~I.~ -dt:!lll.('.tlld
:':1.00 dlJ.l~'m:l
1,ILILlIDe~ tina

'1110.:>1.. Dti~·.dJ.!'I:Kll!'

!!Ill

lliCil~

Ilni.gln

:II~ailm ClllK't~[. c::!1lt>1 n~ ~ll'm

cell

pl1l:~f'l.!'l1iln,g~Dlc~c l)o~cnt llilldi !fcrlC111 imcd prei c m'llOFSfrom :'lear·mlll('\l,I ung Till the w ~[)llg. Im:lI· tiOH'[flt
~Il IllillCf>,

tidj,LtiClli ·to :l lj~ht~y r'lC'k..::t,'j f1'lLi!!i,Or:>{'i,iYUII ic II~ih celb ,HU1C'~~~!.~till. <l n.:gi.(m ~~1'~~ ~tl ri L~ nci!!nt W aCU'Hl1llllotlnl!.: if,. tv 1.2(iSt"~, [~1 R:._ tllU! 1,(;"$,the midlc is 3~jj(],(:cnt[0 tb ~~ro SC:'i,'Cli Ctlp' t cens a nd SlIPPOI.'~::;· tVn) ')~j .JlliCe (jSC~ {FijJ. ~}. I u bmh :;;C;"'Cf.,. GSCs arc:m ~iunll11' xI by dilil:c, ....

tht

11i~h...: I~I

irui ~~

10 U1C

~pacC'

ID~p;lirlntI!OO. 01 1!l1!'ll~'laiJlfl'lE!nta!.fl1olag'j ~md G~i1Ietl~. ~1>!1nlrllrd JIIiI\!e~11y L ~5d1oo1Ajl Mi!dhl'if'l@.. ~tilli'lflHcl. '94105. LJS.,Ii., iJ~P.Ulrft!i~t 0.1 [mb-ydllIgji' .a iIIiI 1f<'l!lia!1d Hu~Iie.~ ? NlI!dlQl ~ri~tlhJl~ fI{HMI), C':I'1D~1~ In~tU",!r.nn Witdi!mg· a)r ~j)1I\, ~ltimull'e, 1Iol;D tlH8, LJSJl

ol'ie-ms Ihe .S1,').~lldh::sor tni~Qlk (JSC)1. :to ~llijl~Ihcy 1ic 1Xl111CLUI iCIIII'r~r H'I lhc 1.1ll1:'l. There IS tFiJ,;, I t·~.J;;UT~ ;lctt' "....i.h i.;5. fll"lrincr. bC'uiJ,;MI I'N 0,· idcl1cc~hat I:'l!!ch ,m.:'iy:llcm, <lCIS ~Il ~hc f~~ I ~~mi,]1 Cd~ih:op~i.!~n~ ~ I](lCN), OJ lar!!!,: DExH t!1;lh: nichl!, wth.:n: ~rLC!i! ~imiiatkin~ mOlY C\ln:.-Lmii11~ d~.ligluc, edl flinm e.'I;,~iii1g nil!! iiti~hc. be.x pW11.dil,It! clri"c (~S(' dl!lj,!i~'il~n. .hml~~.vc m;("~,~~btl ~m~,rin, tlle :I~!ca; lh~ ~~~ehC' il~i. i~b~~uil'"cI'oC!H i~:ti!)n by M ru.l1- Mal,: anlil rClll<ll~ !ll ~iomIfIfthc' ::1pCClm~flm;,;~. which m~o/ rel1eel ,~s IUi(l",- lti,I1C~~I'mir;i.li. TIle c~H-·rmc switeliUl;IY ye ~ uuch~!1;icter.ktCtl diITci;'iCui,;cS j u ccnlmoomc he- S[:lbilizcd by 3c.i~'i:lllo'll ,of (/-M~flwf; ~lJlIEJ 1IIhiq!l!~.in lil!a$~ thai b]~]c&;s. re~ddli:t!1 B1I.'11? bchLlvlor. $i~;rla~in~ by ~:lrollro.h~g (_lcl!:mdmioul (l!r ~he {.... ~'mro$OInc ~lOS~lklllhlg rlays a key rolcbl CII~Ll!riil!; Ihl;; !.ior~ca )ipiult~jc ori,~illm~f111 in uMic Srv~}\I) f"I'<:rk.u l.n. F{!I«\J C/!iPI1;.,~JOll ~'I:'BAM illl GSC:::: ni~J.J:, ille ::wtcch imd !:au.~:. GS{,s ~ 'OSCs ~!, E,~rly il~ intc:rphmiO>:, ill>: $iilgic !2~. Or:lll<1' 14)C alli..{':l\ j!I~'3 r nhl'i coliI.!J.. , .... lIi If!!I'C'i'ilimc. Gcrrl1 O!I ~ reSll'l.r.ul{)l] l~}' t'itl,I,11,IL i!: GSie' 1i.'(·nlfD, .... .8J'vI.1P :;l.glmls I» pml:nlhly [11] arK:!!Jut IllI:!Ch:.llli:>m whl![C1: II IImLf;hcs [(I Ihl! hub (f'i~- HH. W!u.'n
n'gulmlJO:r ~hm j:-; nool!ally H!rlll~1

DHfere,iflt Si'!JI1ii31I:s lhc'big!lW:'l1 knmvill difftn;l1rc bCl.\'o'i"CI~ the t\\~x.(;'l.'i t~ inthl> ~h(lirt.·rllll~~~i!lJl1Ill$ l!I800 to :;pcoc.lfy GSC :!clr·nm~w .. 1. Female GSC:;; arc ~~ncl.ly gr>vc'lTIC'd lYy bone ntorphog~m ..ic pm· t ~cill I B MP Is ll!,lml:1 rrom Lh~ n idt.:, nK:d ill'llL"d il1~ by 'lht· ligands ,iJ,"~cm.penl;:JIpk!:l!t :.md !1:1a.,,~. baiL.mllm:! l)Lmr ~liB]!H C!xpl'L"S6ed U1 ~mm~ti,i.:: Ilichll cdk; t 1). llMP :s]g~mhll!:! ffij)I\:'Mt'.s lnn· !icl"i1pli.rm ~lrbllM,r I( a b:y diffc:rcl1! i;n~iUll

m~I:;' 1l!1l:crc_tl•.i~.IIy I:(CC h e Ille"i:~nQ II i o.tkr ~I~m od~ clil,,·i~ilJll. Thi:s, i:;. n'Ccml:lpl,i~h",-cI by oti:~n~).;d t1ivi8!Cm: NornmHy, only {litH: d:!ll~h. '[~~ eell IT!1II1l1 in:, i1.1 '[ill..' niche. ~ I il.mth :>~XIC':., r GS:[~ auuelrm nidl~ cells ,i:t kIIl"a.lizt,-,d adh\H<::Uli j'll!11C'timls. rich in &-C:.Jdhcr..in ~F ig_ I.• (' un!! DL (]lclllt·~iC'.. lIIy di:;:rupli:rlf,§l tbc jLlIltliomll1 C(]lJ,1lpmu:mts IG~USClii stern (ldl lti;st\ ~I. 2t The 11Ilcc.~uni~1l1 used ~('I' dl~l)J:~OC ICN'I~I:I.y {me oHhc dIUgll~cr:> [mn!! .he nkhe dIini,,-n;, !lei wccn :r1:l:1:lc~ und Ji"TIII.:!'I~. hmviJ"'~~r. I~] 11I~.kS, mmp:I.~lscI~L\Dn'OII.in!l m:Hf1pktx.::l; ~AI)Clj mJ!i]j;·aH.ll!d \,,'i~h E-C'adlmrin III thl!wrll.J1Io. whll~ru GSC::;. OO~]""lC>l t!llJ .h!Llll{Filll. lJ:n rnlild.i;lI!.!"" II key polCl.rlty CUll

Ilro~rnm ter cell

For flith<!:\ll~,~d {lUI ~OC'1[1s~gD1mlb~g re1i~hl}' to asynllllC[ric Itu.<:i O~!licomcs. OIlC drn,,~h-

an ill] GSC's

d,l<lt

11. '9),: ] ti::!wc\'cr:. BAjT.I.i I:Ul(8 BGCN {lllhoJ)[~g1i.~B!J m~kl'lm.vn Olubilll!' IJm.slJl!/Hhl ..

tl!m dllpiit:1ll.odi

(l~nI.N'lM'I!l~

SI~p[l.fjllll'. ~1.nlll':Lmlly
~D Ih~ hub" WI'I.iJI: ~hc

cru.'ly in {:i~.ml,e ~I~y I'It'xl. ...

40.2

SPECIAILSECTION
rmUllr 1mb.or cap cell», TI ~~ C lnse phy:>i,k~ Ii

,Il'<_",oci;;tlilIlll or ,'\{IIll~~ic "TlO g;cr:mlin~ s~cm I !i;;i;1I~ wiililil1 'COll1!11flDl !1k11C pmhiJbly l:(ftdl,~iiJjtbl.ly '~Ol~rd i[!.~~~, PITIII:hiii;; .iOTl • ofd ilr~"i!.~nli!!lilrtg d:oll~J,lIHc.r:- !h... I!lU~! work !
~(l,~o!!Jll:.r.

I!m~~"th~!r

,I

g.Cf11,Hnc and ,.;,onl~!ic C'C I~~ .fi fonn cy,.;.t!'1 ~Ubl\,"" s:iwr'o.~s processed ~)' the g,o;-rrnI1Ii1C·

~n beth ~x~~. d"C'fmlflcr in'I:",.;'!i~, of

sredne

r/ltI\lWJmi,d~chl.ss pmteJ:.'lC S7l7 and

e
D1DP/Gbb,

NCl:iy,,,,,il1 ... il~ ~fI'itlt;liI1~a] gmw~h ({lClm- rethe rt'I.:r'~orirl the !J~irn.::l.:'lli cy:;t ort.:::<~ol1 ce ~I$ ~.13'1, ;mJi the I'd3IiOli~~ltiJilI~~~n"l!..:11 nl~:;c wr:m,linc :md :;oni!L~~C 1)jllrr[l~r:. illl II ~)'.~i3ppC:il11> to I'll: D sign:alirng JlI,I(·SlAlf signanitiSI
I~!i'!~ilil

li!i'I~I:f"i!!II

ill,jlil.'!fiI.3Jlrl .for proper clim:n.::Ll~i1l[ioll.

JAK-

STAr ~Iilrmlill!l IlnflYIO u role iin I.hi! ~:i.;,111lak IlriChl.": ESC...ocq llIilfi:: S{jlJ '[0 prcseree niche rlUlt,;'ti{Jn 111)0. TIn~:;, a~llloo,!:~·i 1:101.11 ule unrl m ft'lnalc llit'~I~":i LI~": B~"m'mid JAK·STAT ...!?· i IKIJilig. thc~ :>igr.m~~pl3J sex-spceidlc ml~, p!.'flIifJIk> U Itlnml<!..:Jy oct"tlWJ~C or l.1l£\I!''tly 'UDr· f~r:."J1l funt:liolll; t:~i[da!lut hy ham ~~I ~h~ 'i ~VI'O
!ll(:1n

ccU ~incag~,

lilt!llIte.5 SU(I~~:sf (I:n ,g'r

(:l h!(:n l~lCir!!(ite
as c>c1I~~lar ~C'IICI'.... !i r;;:.~1 \WL:'l ( :ii~rrwrl",inl!ll te ~~m Ihrut If!dh'iJ~;1,1m;r.lc :Ifi(~ Icm;~IC' nSl'~, arm {l,,:er i\!gl~brb' d. 2).t~ is n~'w dear \h('jt[IC'Wila~i.,,~cnt GSi("~ fnr tile

Fig. t. "~ale and 'female germli'n~ a,nd SOma lie stem (e,u~ 1m Ilhit'ir g,ol'iadal. nfJdlles" (A) Ovariol[e lip ~U, per 11;t"j]lry~ ;1!!l1d Unlfl'sUs tip",Gc:rm ~~Us,. dark pi'ilJ~,(GrS(~) O!!ncilight Ill'inrk(!l::fsb); Slolmoi!tk {'C'lls, gf'fllrl, b[u~ {eSC~ or (pes}, hJlrqu~is€,grnd y~llllJ!H; :S1P~lir'QW!RiI'~ ~S}~ml :fUS,Il:m~(fl. red, (AI Fem~r~~. (lB. qI'~t~bti!:5t; F.~ennin<ltriLamen[: (;C, (<liP T (:e~~; eS(~nt ~e~~;fS(,loltide5lem K ,ell: ~(. Id]de n~U" Hi) Male, Gil. goni.aiib1ai$t; <C, (~st, c:el~ 'bl'ilrk iiI$l'erisk~, centrosomes, ((: and Ell) K<ey reg,ullatllry pathways o~ the n kh e. (eJ ~elillal!e" IJ:MIP's;ig:n'arl,lng ma~IiI'!ai illS GSC.'s :by repressing ~pnt~510n, Qrffbam. Aclitl'~ltillll @f STAir !Il!.1Ilrltai'h5S(J'nlo1l'tk ,~'st(lrt 5;lcm c~ILs. iDpp" d~ii:ilp~'tapl'~!iJi(, (0) ,.~,ijl~" Hub cells (to expreoss Upd", which .m.tvaJtes: SJAfin GS{s. '{eUClw, ~(lcilli:z.edAfC2 and ~,·{atelliinl:bl"Kk a'§.'t:er1 mothe r Celfltf'!l.s>omes,; 9 re~n al>telf'ii5~, c1alJ hte<r (el'ltro5Ii!nes. S.~5, 9
other rU i~lt!.::;. 10 lihe OptH>I:i] : side m

,!~'i;x<lcd ~Iil.iI'I.C~ tiln srise b}, lit I~ast ~W{l di~dI1CLmc"Clm!1!i~lllS, Nc\'\ !~Hm~1:Gsrs mc
rrodlle~x!l!lncr GSi(_'~Ll-SS, W]-~C'M(~ p:>i!1~I~tr

GS{, divides ('Ii!rnll::~W the cap cd Is. ,rnm:ill!;! txd ~l dm.!:glncrs~('I rcu~rl ~n dtC' n lehc :J1~I'~ ill
b!.~m~,~~m cdL~~j~ ... Wig. ~). The IlilJi,iil:y10 undcI!.'(l :-Judi :'j'IIUl11lC'truc; dh i~jQII:oI ~mdt"n.h ... ri~H Cj!lt.'UH1Kl:lllo.:-& im ... be .... IIUt..ul:tt..:d! ~H u , !':l'xpl:l.i,n 1]](1'llxp~usirnl ill ~~w'nl eel] numbers diJ51Jr¥lU 11 Hcr If:llll1Sphmwlion of 11IHlImUiJ.I ian riSe:; uue dt"jJk~l1d :;iiJmi~li!'i,rullJ_~l.lllJuibl (.1</)" UO\,vc:\'('r, new G SC:;, can 01.1:;0 nrise Irern the .1X'¥c[;s,IOIl 0:1.' Irllll,l;-liHI.mp.lir:'\':rn~ eells ill ~orll.l· lil1.C"ry~~~up mal k:asuhc cight-coll ~lag"~mo iii U:-- UU.m::1 ionul ...t~L11 n:~:1~ p 5, Un ~!= i~ R~\ crsion or difl~r"1l.liming 9:>1 edhmay

~lr thl.l

1:':1.:'11.

"Fh(! ~l.iJruo:tYI]!IJtIpos ~tio~] or the cC'lrllo~OJ~u.~:o; ru li>lhly' uri.cntl\~h.t' rn iU:IIDt ,'i!I)~ l[!I!~ pcrpcnclic. n lIl~lIr 10 The iilllcr.r;I~(, \!II ith ~hG"hu ill. ['crimp" bccauM:: Qr Il" gr'c"i'I~!.1'rcllpm::il)' to, Imld :b....tml mie'r'J:lwhllk,~.lhC" 1II101.i1cr cenl.IT.I:l',{)n1c rto,rrmIJ· III rrln.1irn; nc~1. hltiJll': huh and is I'!£'L!~in[-dby the GSJC Illil·m.I,!lh manlY GSC' ~('noCl""oI~ioll~ trig"
I D I ~I Jt

u remain""

:m~nl~l'l!_~li!lg qjl!,I.!~ti{m

IlIJrrI1llJl)' be prunmh:N:l by BAM:. HU ins u!m.iDlg

Iu,ycr of C:SC(l[1! or MlIllmiIJ.: l.'Y!iit

wbethcr l)rr'li~K'I1lbl inhcdlllJlCll or IJl(1 m01hl:'f E:~mtm!«m.leis impol1Oml. for stem cell fmc or is

cells,

or

ol.hu.r

a side c~ycc~or ils ligh~~)' orie mcd cI ivi:;.i,(Yrml

rro,Slllm,

Steml G~I~ PaJrlneir:s~lips Flirtl'b!n"~1JrI!'. illr Il';J~lilmrli,llrl l~t~. ~llw,Jkdt.rt~li~ u n!J'~t 111~h'Ilud ~~m~llC' II ~11Ci'> aI~{l mId m,~il1 u( till.: }\!;ilm:g!.'.~t IlltJj~nCIi!!i1~i rt~-cl:~r'CI'II1!I~~ngl"llg!ll: C)l'S'I$ ~ t~m!lgJ1t ttl 11I;,,'pkllishusn. Irn, Thus. I1w!lllh~)h~,!!icall}'~irnilllr i'«l,ll1iuie stem cells blt:l1e (md ft:rtKllc G~Cs ~Ilf')' Clilpl'oy dl ffrCn'iill ~IlICrsp;.':t'S<.XI~!JuorlJ!! tile GSCs (Fi.g. n. IKfiUWIl IirlC:>ol' succession ,...hen ~hdr reigll i~1.l!GS.c~ i:,;'iil (I,,> cysiph"lg,:nn'tiJIr cells lCPC's) ill n'lfJ.l.csorcscon IflC nlchC', :iI,terll'l C'C Iii:::;;~E'SCs~ in (~mak;:;. 00, II r-rodllc-e
I Fig. ,2. T~o mOO~,r'ill' ~~e r,e'p'a.ce mel'lt ,or 1.®llt ::;qumn(1l.!J:>UJ:m:lkidillg dmlg~'Ih;~ l"ySI, L~- es~lJfl ~lbll th;i~, cudQsJ.:' th~ GSC d;LllgiiM- rill.';.>.- '[oTI!C[lu~fJon :stem I;ell·s, (Pi] Eqju@l, :5t~m (;e'll dtvis;iJ;lr:l.(U) Fin~'f ;1l.~I)c!CLi or 'r1l.'1~llaliml !.BI::'I!clInt>r bctW~\"''ll l.i ried f@r!:li i~~~I:Clllia~it'lJ}1llid r!l:rSbt I!l' eiWI:~'iJr !le\f,erS'i~n'o~ lra m:il-ampIHying cells in g~rlT1i" IlIml~ and rllillm~~GSC:.; ~i~bmm~nh oonum:m iilS pro!:!!,my ~1. ~,In Ih[!'Jl~Chc.lhin Ilro::r."scs n lfm~ qrsts. tClark !lin k, GSC:s; Li'(Jht p;in'k,diUer· Ieifltiall~ngtrilfi~it·a mplfrryiftl9 (111.l; g ru,f'I, ~Q.matnt

lilc~ors, S},lml1C1ric dh. ilS~0I1is Ihc l~nl.jll1'ICchanism LI~.uhas been ~ho~tn ~Clr,C'p!ilcC rCnQ;]tc (lS(."s in ~dl!lt,s;, In (;QJllrnt::!. tlle~,. is .rIP <C'>'id!l;:IJCC I.Ill'lIC:<~n1 ror ~ymm~tri"tl~;111!OSl' dhi'>tt'I1~.

u\ll~ lh<!lpn:l1ride

the ijlidu~,

rr1iln~ I.he crc~or ESC ....'i:O\'t'r nm",t GSC SI!n'· r,n;eli_ isolmliu~ GSCl'!. lrom 1.:1Ich Hth~r hun. not SCI ENCiE VOL 31i6

S"'Ill."r;p1. IballlK'll. sm::h :L~ adh.c;s;uClIl bN ~lmJ]I~1.~ tdls mid d~ilCm!mlCl~011dOli~ range :-':i~n".1 \Vc s"

WW!.v,.sdene:emag., org

2D APRil 2007

Germ Cells
em] only '~CCl.drule' uhmlL the selectlve [fIIU".~ ror 1.11<:: C(l,11U~IOIl la W~ C)r ."tCII1. cells, \W must cOl~!in8.!C to expeet the i;mw;UlII ,~nd even the ~hm[may b~ m:-opm~~.i"~ctor dW8C dirr~l!loc:;;. The I;u;:!. lh,n ~hc~ ~ ~mIy I r! [;;;hll per iestls Olnd ~n)iqu:e, ~() !1!k:l~!I. p~ OV,ilI!)' fl11) ~:\;p.!ai~ the l41r~~if IRIl:re~~t!e~;,god ho~e~ UI,)nlibL:r or GSC1I. Il!l:r nkhc' hI l'Ilalc". The ~11f.1l'\;' L L u r, l'!i~ AIm"tI. ~I' .. CtII 1Jl~ IiIlo'.Zl. ,00,'j ~~)bli!:iitlnC>:'~,~uil~1f)J fi~'~'irWl~c IIri~11t,ui(1!l ln G!OO~~, z t M, 'l'.'lmMnll~ HI. T. FuIDl!r.D. L JOmM, t. CeUS~~ 'us, I.talc.' (is{.·s ~]KL}' bc a C'~ln:;:(\IUe;'~C'.c flnhe JaI'JCiI: sss 12!J1I'5~. nlche, TI~ in ~Urll, !U;~y ca~!t<;~(L Iln;r~lt'~ltia~ i~. i. 1'.. Xi!!', Ii.. ~!il~!iIlJ!I, S!'JBll!re ,2"i11D,. ~M ~20!J(". use or ,cy~~ r,:v(,"t1'iion ,.. !I)Cr Ih,m s;ymme~I'1( ,~" t<. A. ii;l~ef, D. ,L ~onei, c. Sthuil, ffl. 6. ~_ogers, GSC' divisi~lIl!l '(11 ~en~r:ralc rel~!(II(\clf!1ent ~~C'ii~ ~ 'if. fi.!llilf. Sriffla!Z~, :liS,q~ ~lOO]). 5. 1\11. h'lm.. E.l&!ihmk.Sorjil"~ 29~, 2~4~ QOO!l), cells, N~~ITICEh,_;I:I,.":;' i::; pn':l'oohly ~I!.; il C:\l~.,;cl 6. I'll. t Oi!"'~ ,M. 1. fuller, C~1l~<Ii. 111llI2006}, 'th[l~,""~Leh :.tC!I] cell ty pe hus bl,."C{~men1(lr": !J!l' 7. D. !lim, 0. 1.'i~Kf!:mii, (w'i'. ~I.U. lm~ t2!Jo.~). k-:;::o; ;).p;.:cill~izcd W I~c pl~rlic'li1iart\:(jlJin:nl~nb II.• ~'.~Dffii~\ t. flr!l-g~SDI\. DI!ldOpmtri[ ]1$1, 11J.II:1 ,(lOM~. ~Irhe t1db it ~u~lptlrE~. Thull. whilli:'i>'Carehil!ll t
~~j

u..

flWf.. ~>'!, 98<1{aoo~~, r; '5dw~ ~',~.!lI!~i!!l'~~ 1li7. ]Ul I:ilOO'l~. E. Oefl!l!!@,. Ii.. t. SiJ~t:!IW]g',~v. ,'~ 9,S'O~ ~;"1oosi. ~.~, ~ ,!,~'lruiilii~~,~_ P..~~~~f!l, ~ ~ ferrnll, M. 1. fLil!~r, 5(~a ~:iL!5\, 5111 ~Zll!J'n. E~ c. '5i:IIW~ [.WOl]c, n 1L.1&tl~~ s, TllllJk.~, M. T. Fliller. ll\!mIlttJpflIi!YI! :JL0!l~'. '1<)23 t2(102~. UI. R. L lItim§:ie-r, stii!fl~l ill£.. '11M (lOOn. is, T. Kai,. A.. '5!1i1"adlim~ NrmIrl! <1128.• !i~ ~ZOO~~. U;, C. E'.taWl!'j'. 'E. l&blrl1ii,WMIl' ~O'l, 133::1 (:lOO;t~. 11. t MB~~\)" Y. ~~~lI:ilmJ, S. r~nrdi),lft; Crlllt i 19~,~20t:l1t II!. lMe,.mellllly if(~'led!'le'iIiJIflOFI ~ro..m rttI!:H (51rul'l~~ roU):K53014 "'Jld ~Ol~1.'JS61 h:l M,U. I~d rt~ffi 'ItMi 1lH!W 10 M.S.

9. i". GUm .It. lehm~rllll.

'~ff.

REViEW

lMa]o 'Ger,mline St!em 'C,a IIIIS': IF'ro'm IMlic'e! to Men

R<l~ph L Bl'iin~tCir

mci:l:imilt ,GJ":'1itOJlii>lnn~:d Dhtu GO;.,rp J5 the prin~ny growl.h rtlrtor:>IJ.ll!,porling [n;O~,,; sse s(;Jr-Itm,;waJl
{7}. The ocv!fJIOf}uI~I]~,of (;U~Il.m:.~ DlIIl,.1hoili;
~l ~. \,,'.iclc rllilge

roo..lc!'cC'Usa$l:;:~,mdsor clllnlll'lS orccU",. liffiiJiNl= i.~ rcrlm',"oo. .he du!Up cells begin 10 gro;.'I' ln I ~I('": illil lal f'L3ge5 (If =>1J;;'1ncell, 1I(I.... ..... in I t)!)4 ('I ~'il~!QlJ)l(mM!I S1cm "d~ a.~~.y Chlliu:;;rc:;eill1JI,j].,g . 'l:\ r ,~dl:gcumli rIC' :>1.01.'1'1'1 eeI1$. ca~tcd ~~rI... ...In:Ih9g;:mn:J11 stem ~elb iSSt"s~ irl po:l1i- \1I'Ii..'i reporb:.xI in mice diml, iOOllilkrl SSC's 'by I!K~r diffart'lllti":lliol11,::1", S(''1JJl in ,. w.o tJ, 12B~. 1i1m~, li~~;11 11TIl]Jlillll:!l]~, iIIFC lilt:" 'IULliIL·dlllir'!'J.1or ~Ibili~y [~ MlTh'ml,,; ,13c~rnly or ~il1IltIIo~n~ilS. GDNF :IPPC;fl':', kI bot a I"rim~!IiI;~uI:.IN Ilw: sel r-It:'III;!"WoIIII ':!J.lMb din;''fT!."il:Li:JIli.Oll f::'lll!.: d~\:~i(l]] ;11~~'r...1Il1>ipl.ml::Ilinll 10 ~he . :';('liltiJl~r'-'rul.!Jli lublilkosol' ~li:milrntlg:~ILC:-i~ llilll.· proeess lor spo:rlTli:lIt1m:L 1m mouse 1I11.tI mI. S'SC:; (7. l<1" und il i;." prt:!b:Jb~y ~ ruciUJ!llllD i11.d..:.If:l SUmc'iml~. number or SSCs r11:!11Llrlw.n!' and, 10!ll1iJcrr with lI!llCyi,Q'S frn:rl1 ueonservcd ~J t~l\!'n~w~J:;..igmi.l ill a~i m.c:Jll.1l.llal J WI] li'l!lmk~ •.ill'!J ~~'111:i;111O:rsfX,:i'el> ct1Illlllllllit,Y. ssc~ "vc!Utr.:I!l(.;plllllt(;'d. progt'ny di~l)luyil1gruJm d"Jlmr .. L"CliLdI: onthc W:il::"I]l":lll 11Il1.1Ilbr·ul1.c hhc :n.'11I] rlir. o hf)plmy~\X: w~,,'t: rrc.:ilJll::t'il by 'll'l:t' rl't'ipirm ~}I, SS('~, ~7, u. lj,~. Sirnilar h) ~'lIlbnyClfliit ;s~(;m0.:1:1[1 (E..IJ(';.;}. SS["~ 1!Jffi .... in !,'iuo OH rct;"(lcr oo.ii:;; In lIEllldie:;; ~rn)'I\o\l.1d linelil SSC~ of all emus nlblJ~k:in LIlt:'I~[i~ mid .~fIT(,'" ,d 11I1O~1 on~l)k'l,C- SU~lIl!m o m~:rl"lI1mUlml SPI:CI:l-1; C1IL1Imil1:'1.Icl{t:".g., (;;~t. ItIIbhil., Iy :-;umlumkid hy scm.uil:' S;cr,wli t1.~lh;;, ,,\'hirl~ rcmn i'ill.llld~ dUI11p:;. ami Ih~)':oiIail1lXi'l'iauvC' (("]f rov or ;[I Inlcrocm nmfll1al~lln. !J'r, nich:l:', \V~lh~11the niche, Jummll d(l!>.l~ J Imlllcrip'liolJ' la~'IOf'.1 to:.)!' 3,.f~ ~f:1;,.Elilll.lJilJ~\I. bmho'lIl. :md: linnmlll ~would rorO]]irM: and lIlk!lilnll£: P:1.'bOl'.f"ll~IfIl.l:;;C' TI!}l~ ob:.... ~:;~ ·:n;;tLiom~ ~ {If .JllRlwlh ftK:I_ru-s, liml cxtr.!tcd hurm ~g:~Il.I~l-i liI:'!lul~1£ ~hC' toiClninirt"Rm.. tllll:m~11'~ inul'lLlrLOl:kr I['Lt"nl mine .<lIiIMl~I~'tI ~I~l. ~ ssc:;; l"lliight ~ prl[I~lIDtL'nl I ~O'l!l.mt! g~.l1L'l"Jk' oolurl~ or Sll;;"l c~~~~ Id ~~II), m ~h['fl nik~r~l'I4' d':,{'.i~i.oIlS'll fScs:n: Iltllh~r ln~J.~:mll<l\' or [0 fQnllll.b,LlghLl;lfCL!I.~, Ilmt ~\ illl'lC!.Wnrb~ oo:mp]m 1.;"\ ~[. wlun::m; E..~.~ ro.:JIdjl,Y!J,.l'IH·ru~JJ k~ruli'H:!IInci:J1jl~'Ko';lrud 10 !:Ie carly dibl,!.."f.:!nI i:lI.ing ,dal.lglll.er lK'~11<Il!; \\ .~I(!n Ir.rn~plrnll.k;d .~Il.,·iw.J. 'SSC's do nol !'~x1nl]ntogrJl1 i:1 (I. J]l Tll.':,'\.I.:!' ITrn'II.lIl.l~ bl.dlOC::lll"(]~illIl di I:li'romj~jtion pm..-C:;ll ,(IoI.·.q:w..~m'Wl.o.g.~JKlldis,l'l-"\ltI!lt· in~ in mm~ire S",,",~11a10~jOil a ikr ahm.u J 5 days in ::1Jgnals il'i .he stetn ,ce1~.uld ulkhc mus. have bc~~ lrom ~lI.inl(irs umh:r $in~~tar (;'"J"I'KIitil:Il1iS '3" 7). ~h~ mm.!~ l."lJld f'l'i· d:.l)~iH Ih~ I'IL.milll f i. ~"1iJlnH: high Iy con:fiCI"\',oo m~roilg, ulmnmal!r.m ~~r~cd~"';!, WllCl.hcr .he n(lUii'll~.1 ,[!!!Juln ssc.'ca!l l~ in~It!~l y<:UcJ, Slll,ce [\~ l"'"iXI!11C r'lJt.lri~lo!)1"c~l~ fI.:n~;]iI (;i, c"{i!ntr{)~'cisial, t irni!!g of :>I.,'q~cUl~ii11 :steps in SfICmm1A1j,;CuC':5is~~ ,tLJringlll~' C"OI:!!}i~ ..:.r'rne- HN I mil~~('Ifi -nil: ;~ ;jJ1~bilil:ty til' a nm~iim~lbt~l.:$I;lI'i~lilM dh.ag,.'1lc.Ii!urmiac and hUllw~ ,ti~,htl}' "'"g(llllllld by" .b~n~ of II'll!!g,...'111:1 ~dl. ;llJd fhe rll!}'lllg1011.1i!ttc ~I~i ,kilid II Cli!tliUl! :>~~lt:II~~Ik~ .IiIllrM ....~)~.,;,!.'riitl NI'L:Yi;."(I~'~r; ~mli!-;i! ItlmuiC ~pt..'i·m~ltutN~ whidl ~ S ';:'i'It'ili c,:1 bo, ~ilj'lj"!{ln I&II.'!Ji:lr"rellit~lIi(ln j'il\l:;re~':i. TIi~ ~i~1 $1~f'I :in ~~nlliilhJ,~~I]l;'siJl; ,b.; ~lb': ( IIi nieuh. I'll C~VI!i\;'~~~'VI: :lnd ~ctl,lIire sl'H-'Cic;.1- i:'C'fIlicmkm lIr S."iC:<: n'lill(I~ ii ra:sihk ~l1',:X'!.Il1itllC dc~i~k,n ~lfill] Ss.t·II~l!llit>dm;~dmJ~Il!.'f 'Cl;.'lIs'il:Hltc !ip.'C~lk' ltroe;:,j;Li'iU'>, sse!'> Ihmtlu I,~of Iii,,: SJiX~k":5, il"il~~n~ilrlf =1\:,!;r~1~s hiUin n!!I.'I'1C1: ~] f-alC~J.;:w:-~ilnd ~ liJr loJ.lJ,!l p;:riuotilli di If<!L'Ii![lIjm~lI lit vilro ill il rlJo,'UMllii mrull1,.'I" I1mt lis tnllAh"d !O dilf"''I'II!~i~i~l:iuli. Tlierc rut' lID ki(,"(l""l~ c;':'~jltin.t,;'d c;]jll be c~o~l"i!W\'cd 00i. ;[l'io'r~bblc ~o:r 11:»001 IJll~ .$!Ient,cells (16), Ucsc uniquC' bitlch~i'lltca~ or P~~l,O:j~]ic i'l"lfll'ker.;;, ~OJ' witlli C0I1WliiOll lI;."C'lH1LqlJ."S '[!lied I,hr SOJliltilk odl:>l t j. d~'>lklgLl~~liDl,g.SSCs I'rofO !hcir iui. i-Lll d~aj~l1.Cffi. n~a'o'f!!ilaiJilit)' of ~i":: IlfIDlspiammitm OO:MI}' SIImii~ dcnnorlillRlIIo'd Iii;.!! Oct .W Qjld SRf ;'flX' 'i;:.~~IbJ: !Il11l1~11:'i.:n;Uli~ltt;:d utt.!,!,:lUI i~L ll;i:; lir~~liJ- Ir~a~tch Il(Iii>.~ldc~! :>1:ud}' s.~ni~1Ulli!ll~ t4) ;~Itd rrHildiml1f;:, Je,'([m.' ! (.'lf~ :1), "~I i'C~1 sp..::111 ~":;!;lIb.eMmrlg, 11:ru; kd 10 nil,_:- ~'i.l11liilrLHm~ !\.'fIltc~litm rlK'lUill":t it~~ hUllllj"l!:.R"d siud;~ 011 til!:' hltl!llgy of 5'5[\:.. has an..: ~'l::1)i:Cl>.","'tIin sse...,~Iu,""'~"T,.VCI!1'Ig, 1]1.:" kl.)' SSC~ ill ....i1:W IJ), G~~1i>reU1[1111l dai\td IK'1lfl}- O!Lt'!mtirll:ml ,~.r SC ."o.Il.~r-.mlll:;·"Nlil E ,mOO rlmip(l.1~n['y. 1'1~fQru, tlw ~mllhic: IlLcmr (GnNI~l ~~'.:J1'iiOO]lifk-cl as .:J t'rili,c~1 i~ m.1I!.(l<:XpIlL."'k'il,d 111 sS(':> i)E'ilh'Hlrn~nl I!IJ IInlmal IBI&'logy. S~~ooI oS \ilIH'1!ffnillF~ I~n:~min vivo lor Dim ITlll,i~.tIi[ln o1"\'fK'm1~.I.ogol1i~. si!!II'lli~ll rllCclm:lllisrns lI,lintnl(linLlliill.-.clr~:lim'1;,1J~in ll'I~k~mil!, lfull'l!i!~~ of ~iI!Iin~v.ilnll'l, F\lilhd~ptl~ol. PI!. SSCs and ESC ... me dillTcncnl. l"k-.;,t.~ sE!ildics al!'U 191'iJ1lI, us,," (,lit III \'i1TIJ':.tlKiie; IlIs.ing '"<Cnlln·fruJ.: clllltu~
r -

The procl'u(tiCln ~r rIlJndj(lIl:i'lll~t~ g<ilil'l.t1l~§; d~~~lldern ((Jlthe OOFllim.i((!U!i:<I{tilJily (JIr g~nllfflli! stem 15 ce~~ Tn!!' al,iiill~labiljlY or a '~i:!lfl5flla iliti3JtJI~n !I~.y sysl,lelli! 'lEI ulli~~ui~Il~"'Uy ndfFllif'!j' rn:a~~ g,elltiililli~ stenn ,ce~l~ lila!> a~lo'i'li~dl:heir in 'I;i~itJo (!Jill:!Jm~, (r)'CI!i!lr~l'II'fI.tiolli, diliiOg;~m~i( lffiodirirnliM. lI~oIfKlwr"h~ ¥~€1lI1 nas, ~nllb~~ th~ i(l't!ntm~tiCln of LJondi~[)ffl5<llld r:"c~or.s inwl~in 51~rn o~1l ~if"f'~Ilfi!wa~, th~' IOl,llIldiitlifJ:nto~ :i~Hllil'la!!Jerr~~l~, ;]I nd ~h,e prnductien o~ ~pe~",,"lt~~iiI. The i!l~rejistd klUQ!~ooge abaut ,th~~ ~~I$i~ ais[) @rg~al pQl~n~i!ll pr iJiI;tl(~lv~lu;~, f(lr 1~\'@liDp~~.r,or 'the liIo~s!ibl~~ryop!"l!"Sliln,r~tiQJ1, !Ilr ::ltl'lm 1~!i!lb from lIiiJr.; ul1d~I'~[linfl lui~oMme,nt f@r (a l'Kl;!roo sar!E!!1iu~j{1 ~h.t!frr germ lilne.

of :\luJ~,,";S. n S.S("sin vilm lR. 9j. o GDNF bjmlin~ mil.! :;iSlJI~ljn!l OI]t'LIr 'dm;w.~·1 (JDNF·r:Jl"l1i Iy 1Il"Il."'I.'lplfH· oj j GFRu ~) [mel 'U1iL: l~(t ~p!Or O~I sse's aed LII1.diflb~~!! ~i~~~~I/lC!lI!]';1tl'1~nb. Allhoogh 1ilC!~IilOFn,r!II nnil r~ct me c'iJlJrf!i1ec 1il101:",'l(ll.aI~.~hcy ~!al;!cf!flI1kctl ideal lo:!' I)(nl'if), ifiR $,'its, rrom 1."C:'M;~'" fI.""SU la,il~ h~. ~'ICll" :i I.is-~ol~~ I a:I enrkhmem 'll". ~eILXU\l!i1 whh ;l~1!~0I;11e~ ~

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1'Nl:!du~l:$:~ 5- ID

~U-rdr!1 .:nrklllm,·lll nfS'S{'$., J;l..:n~'tall fI~ !!X;CC'~I~tt cell ('It"lpulrul0n,o;. rOT :;'l~Ldli~ Oil s~('~ (Ii}, f 1).

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404

SPECIAILSECTION
p.$;H.l1nm, Ihe bindiing oflh~ stern cell lit«nr Ki~ ligand ~l-Oll1~t'd!D!icclb m Ih~ ~'Ki~ ~OC'-:I"to~: could ~Ilj.l.~mc !~Iu~!irli!\g dlmcrcluimui)<1'1 Ih~ rO~~I\\.~y,fi"lis ~~) (rru'!111~lifll~r~';!11Ih~l.r~1dini..'f' .:n~i;l1i!~)enniJ[Qgflui" arpcn~ ~nhea crilkcl ~ sse .j!~nys QlllfJ!'Ill1.<'i!Ihe imJ'liI)rt.i[jlct~ orlJd.r,b .: i:1!i~ i~ i~ n'1rLt~ntn.r)' ~'!nlHt b\ "pt..'l!'Q1!ah1g~1¢ih. n!~ limin,:,: !) f ,wl~cqlilem ~'Cf]!1 in clm1."'i!il'l'l.1i<l~iLln ~. Ct1)ltL'rI! Ied t~Kl!d~ hikely !II!!!! Em.! ,1m!l.h.t'l in ~LlCIlt\e S$t" scJ~l'C\\'l1 061, £/W! C';;:Im...~ionirl s..'IIDlic~i~s i~ b}l g~ cell gCU~'ll!!~, ~'''<,d.h'i!lu.~''\!I,'~il'C'a ot i\n ,11m 1l\;'~CVl,,"(1M ,~nCCl niche ~I,IU,"'lion I fi), Ollt.;~' !l11C1:'I1JlillllJg.<:"I1C~i S 'Il:gublion '1Itvnlw" ~!n;11I~llIcrferiu,gRNl\s. 'e"\'nL\11krJ1<1riiy 'COl'!.5C!"I.00 Ill~ ~!cc!llb, ~1w.lk's, judic'me .~l1iI1ll1J'lOn.1!~rotc in S~C rolf tha1 inhibit £,1,;n,; ....lp-II\.,,"'ii>i'~nl ' lIl1d ilfC pfl.."1' .... ill. li~ !;c1:1!.:, ...ui lfir 11FIfIr_n:lrwm" k-ifAemia ;;.im '·:lh~,,'t'1' t(~ N genIi eel I~,liun:ruglilDllIi ~'Im:tlJitJg~H.·.,,>i-;.I! I j. 1m' (Pl=/ ~ TilfJMm't(iingll1rueill.a1i.mci(1f("(l{ih..'mr ,,11'1 r"F4fll', Hnd pcmlmp;;; j'l,rull{).'i: 2 ~r\,()S2HfNllh, TIll! :Lbility 1.0 :i'Ct'tl\,~'lt,. c:'1)'tl'~~I'\.'~. t:'LdLuR:. hi.u GD:-rF clo.'l-. nmn;!;;,IlIllf~!:: l.hl.;"~ gen ..~ j 161, mit! '1:t:!I1]l'ip~m';L SSCl> 11:1> SliUlltillrul-d, nt.:\N apllnm.:!'bre •. ~tii~:moollJrli<;nl,o,f'd,lCir ~I!:liun is. unC'il::ar. Ilru:Jdl",s tu 'LJJukf}j~ilOOinlg.Ill'!.: bilJklg~ of Ihl.'!i>!.!' .in.1ll1m1~lm l\:IB.. Il hal; also p:rovifkJ all (IP!X1IUi· Allhnx:J!!Il dll.: 0l~~1J.m.:: ,Hid nrtlJlSplmllIllion :-;y.s;[(;rFl~ h,TVt: m1l:fe ii poo;. ... ~ihtc u, :>lli1dy sse ~clr. nily thr proe[icn~ andlTlK-dical <.IppJ.iCOalDO'Jl.'l o.f [\;1Jlcwal. dM ahsu]ll] of' :;imilmJr i!l)':SIJ;;IIl~ for g~iil1l SSC~ and m~klli n~Kli"idlll~ ullIk g<:ml~inl;,,, rmlcJ11idllly inIl1I~lrtutl. The Cl)miIlLWLJ~.lonfl"'u:nn ![;d~difTcn:miarLon :;.mg,~::; or ~f"I."l;11Im~O~nn;;i>r:i' Iln. ; . im:n:mc in. I.I,W raml]oor of ss'('~ ("llJhLJR:'IOC~w in !Ctlll111lirnLIJd 'lJtIJ C1Hl1l1l1lilmuian nfl't~I'I~bfOI~' rncch~dhnvs, fOIT LI:n.:: illlrndu(;'Li:o]] or !,1;L'Ild:ic lUodilim· ,11l~l>m~, fm Ihi~ pn:IC~":$. Moroovcr~ 1.I:u.:: ~l]lri.::~lJt l,iol1~ 'lhmlllgJ'I 1]1.{'~nn iim: 'l'I"il.l:i nlilthm.b. nhTI:'·'G'~di]]I!,n."\Uom~ ~lrut:llllm~ ~:I3~~ni"l.Ilii(lll Dr hp!:"1i· !..IC\d0I"-"-,1 mrSlf'lmaluc c~II~,a!1~~ES('~, Pot~!~~II~. m<l~l~!!t:."l\i~ h",.;, '~'OJ1~flOUodC'f~ lh~ probk'!11. In nckHnirm ~fI:\IJI1fL1IDndil~,!,l;he SlCll1Jl ()d I ~n provide- [I .my ~CIlClK; n:t1l1lptulntif!l1. nochaditlJ;llarl.'Clcd gl:'nc t ~"UI;~~L>IY !1!chc. iJ~c:i1;:!1n.1.i cell ,c;.:,te!1d!'i: -rin ~ knockout flr eorecetlon. can be used il! uII mJHU1);1Iku} S:Fieck:.s tor whtch C~dL~I!1C' c{)m!i~i()r.re;, nUIll .hc' bascmem !'l'lCln~ram)'10 Ihe huren nfthc ~H~miICPI~~I,~~l.lh\l'!'c. «I.!u;!l;:~~nglInd ~1,l!l"!'(l~!1d!ng <ll"C' OC'ld()pc~l. ~Il)\,\,wcr. frl h!!!l~!l;~. di~ic;1l! and " j,l!i.'m!1 ~'\:Ih in m''''1~I}' ~la~cs Jin1:l£nli;.lim~. i!:nh i~4.j1liu~~jij!li(lIl .. will, be 'CI'ili!.'llIyiJl~ilClrult!t. S1,.;b~lJeell fl.1C(i\":rj lmd C!)'(~J~I'C'll!:'!"\<l!illl1 rna) F'1i.!r1.llo!I'I1'~11'I!. 1mi~lh idU,l'i1 ~~'f!ncell m~~:1"''>OClc icatJ~;: 10 !!~~ l1'IIUl'l'llilliml S3'Ii.X'ic~.!Iimlt"ll!.dJ .DIe with "'lO~'~hnn one Sen:oli cdl t]. 11}. NC'ilf be l!W!~1 the [l<lS1;;mCnl fllr:!!lllmlne- [Ire ~mi}' s!lji,ge:-; fir be used \.0 l'I)(l:1!!l"\.~ I.hc nm!e :g~filn line- (lr"'[ltL~~IC' ~'I(: ill1.'ltflJ,;'ICiuli(l. itlCh.!ldiJl,!!l 'SS("s. lind tll~d iff'!,· I!hcstock :;.rl'1iill~[l!!>:. cnl1!fl'1H(oil Uf!inmh, nm! ~1llai\g£rc~l :spt..~k$. J?crlimfl'S til1Cll'iOOI pj'OvflC1l~i\fc clllilUiJd SllCl"l1'!.l)t{\,~l1i<1!; hese cclL~ are tiDNFT ,md IJoI11allillll) \':lllL~MC' mooical ;:Qrt~licmio!l ('I'li r~s'florl$i~c ~ilL!L <k. nm dhqplay .he ~-.K:hlyms~lC' :sse 1C;~,m:d1, ~:;; far 1m.'I.llI'1x'l"Lal, ulldIC'I]lOing ooy:> kim=~cpi.or Q~ Iheir .'\I1Irf~~ I '1, N)! Ii.. C'1~I.irnl dlb'll,mmh!.T.trY ,tlr irl'1ldintKm I.or Q[oc .." ('!h Tn ;!o.~P' ttl in~l~Ih: 1l1coir OtUI\ lThiol:l 1.0 d.rrtL'f'-''flti<IJlin~ many ~':IS~'S, Ih~ SS('~ l:il't;lruyrd :1.-'>:1 ~i(l~ :In:' dlh1 ~li:n1ilrtltl~nil~ i..; proh:JiJ!J:-o a :;i1!llml !l1 ~Xpll.""'~
dcmL'8l~!r;I1!:1lJ ~~!"t ~pn."'\.'liLl!1. of gc1Ile:.1lin: Ilm;e the tr.!l!n,~P!iL11l r~Clm'S.lJnlj ClJ.)/J'lllI}!rmm,r (J'm''lI!A,,'r }j IJJdfil). rl,~"r~k~{C_il mlJ';'(utr.: (lrml~ alld UM !umlt.wl(l;rl t/JJ,'i{ [t;; hls,h~}' reglll[]t{:d by '(1DNfb~ "!'lfL1. FLI!.~!.io!","11 ! n.,!1~5;lIill1l[!1inu of'in;.IIIIK:m, arid ~h~ pmti:tl1!. is k~HiuH;rI!ilc.
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could t~~~.;,. LIt .u,~' e :!!1l1!l' I!1ir.UU\CnI. l'IoOto.:!!lial~}~~ !'I!I1!i!!g ill l'I;:oo!o!li;wcio!~ nf ~he tCji'l,~ :1I~dS1~'m"
[tlP.l;l Prf.l~L"lt1ioll~Fl,!,!.

a cell .~I~~12lkm pmd~IQcdl"mlll1 tl'I'C Thf!'l ~U ~1."'l~Pl,<;;br!,cflll1;1Ji!rling ~SCs; nh~J1 be ~iml~lani~,\~b;lII;k ~illQ ~le Im!icf!I":';
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(i,~ J'Ltir~tll~'!' a·Lc dL:rlniti1.ll'l oflm;lro mid.h Ih:ll, !MPGJm1,~!.:"H:n;uC'\.ru] prssC':., h1:ni"I.!' Hllli!1l>1.: Ih:n il'li~i:JU:diITl'fl.'1Ilcimit.m illl urdt.'f to '1~rn...ide I.L OO1.lWf ul'lduOilrlal.dliJlg of' IIJu~ ~.;"tl!' dcciision; (ii] 'IJI!C exL'(lrll" ion of Ihc- I>1.::l1Jill,l:r'l'l: C'LJI[LJI:Il: ""Y!i!lI..'iI]l m OiJl~'1J'

;miclmls.. (i1li!l'1I'i~'(j (\Jlrl:jIIll thtIl ~H~rull,"wal, si!!n~ls nrc C~JII1:il'rv cd rurmJIIlg 1i11lUml111L~ :lIIId thir 1lik:~lml ~ipl:iCiL1j'Dra:-;;mid fi ii), the ,tk'\ ck~Pllli~'11of 11.it.ldlICKb II:!I al.l'ow ~11 \' itm di IK'lt~'lilO~ll.lurlui" :.t!tnl celis II!) Illrwidc n};[!Ll~~iI1~~O~~ whfch \}j,'0'!.I11!.~ 1.'fIC'elllfl!nml~ly \'!lhJClbll': ln umkust.'mdiUlil Hl'C COlllP'k"X rroc:~ fir SPCM!~~W~UCS!," ~1A1 \~m~LI;~
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These advances \\·.i!11

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Mk ~ II!. 0i!1l!:~, 111.11.. ~~~k,

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'TesmilS; e,iopsy

Cltlemol:he.mpy' 'o:r Irra,dl[:a:~O:n

Fig. t. M~~€ '!J~rllli'li~i!' stem (e~l Ipr'E!c~ffilatiilm. B~!tlr~'lr1!'a!tnln~ntroll' ((lI'I(~.r D~ dle-lr]otn!l;!r ilJlY ~r il1'mia'tnon, a boy wuWI IIIl1derg/D Oil ~e~li'l:uuw blC1~ ttl, reC!lVer S!.eI'l1 Cie,us. The stem ",ells (I1u1t1 be i(IYQJP~IHe!'l!'etl:'O~,aflJer tl~~'e~l)lPiiflel1lt Ol!:tl:ie n.e(e~sary '~e(IiII'1't(JU:E!§:, (®uhil be ,cultured. Ane'~ 1!r)~lCItm~IlI~~U1e :stem ci?lb would belrltnsp~ nl~dto the p.aLi.r:!nfs teste) fo.r~he prodllldi:Q!'l @[5[)~rfl'la~(U.a0!1"

11. Co 'CJMin ~' tir, .• IIlMl1!:4:16, :l!.010 (<OOs.Jo .. 11k Nt ~_ waJg, 1... lin, T, :lilt ~ l'IlK ~!!t MI" :1.73; t2lID5Jo. 1'. i!t.. E. fillema ,~'tir,.,,~!;Ity. 1,9. 791'1 fi200!i~.. 21'1_ SIllU;i, HI. T!!lna '1'. Saga\. ,flI.~!l:i113>'1. ,Il 17 (DJ}. 2']. It 1:iJ1,S!ri'tr.I(f' iliA 391' '(2:00n. :2'.2. IliIIar.'l:. R. ~l1gl!rl ~ [l,iIoiig. 1'1. ~a', j., Oalley, 2I1D .L'~!i.!Il J~f'IIIf~ lOill-n!l/!l!J: O!'! ~0l1{Or1~ulfQl! D:I II!~ IMI\1lW!p1:: M, ,~_1Il(,1J.D, C, fntllfiM, ill:M.3PiJ!fl~ij ~!l1d 't:. i'o~ 6iY Ol!liil'llin(t In ~:p!:I'imMI1; ~, H~.)!!ir!~ 'for Mp iI<if1ll,~01Oilj'jl;plfl' .~nd f~lft.ll .l'Ii'!:I ~hE N~J~~~ hUiiIJit ~I (hiMI !iliUm Mit IHII!'!~n [l(ltll!lqlfnljff! Igrool JIO~.I!45~ ~!'!d ~pg" ~,Kte!;Bg ~r. ~ !-le~ I C. t:Ij;I'l!erl! Fci;l',mli!!l~!'IIgr ~O;i!ltilUil.l~~
lO..n~~emDl!JI.13 nil 1

WW!.v,.sdene:emag., org

SCI ENCiE

VOL 31i6

,2.0 APRil 2007

405

Germ Cells
concluslens arc umin:i1' bmS.(ld m~ ~hc ~llIdy of il'ljlJctoo e"ilmchrnmMmn~~ DNl\.llm:~ i[!hi.\~"

Tha MII,le1Irnal-Zylgo'liic: Tranaitien: ID,e~alh and B,:irlh 'Oil RNIAs

C'i'!r~Yde'lle!op rilE! III 'wIl en the 11'Ie'!\lly lerrned embryo is, lJam'( ~iplhmii! Hy h~aJc;lhre. DlJ!lring the lIiI<!It.e rnilt'~l;lcll'~ tr.<!psitllon, 'e fiIlbryonktraMU~Il;lic'JlI is initi~te!li! <I rid mOl!1lY !l1I~terf!lill IllNIAs ,.fire de!"lf!i!ded~M!! ltij p~e rneeh anisrns r!li~yk!teihe birlhor~gQti~ Jlltaler!l<ll gel'le prod'l.!iCl~ d'rille
RNAs alldi til! ae"lh Il~ m alem~ l RNAs. G~nome activation aplpealrs to r,ely in part on the :seqwaslriitklril (IIf trsn sniptionilt ~~pres.u)rs Iby the ,exllol1 enhally 'inc r!asin!llla,mol!!"It or D Mil duri neg dealtage ci1'11isiorn;. Matenllal RNA degraJ!ilatiol'l is, induced Illy th,e bindingl of p'rot,e~n5 and microlftl'!JlA5
'~(D

LID'!;: 1"e!J111lhll'; unclear .((1 \\11liu "''''iJ\;!ll! Ill(: lia'l1litctl (i11~hlil~:ry mpd~~ [jG'liP~~1,;"S, (,1l(9o'g~!!!n~l!I. to ~~!w!'..

Th~ rupid ~"C~Ic,vc~C:,~I;_'k'in~(il .. nd {12 1~~1.'lSCl1mig.h! ~"!L'1~!~liIU!c~ thil'd '~ImI~!JY dm~ inl~!Cn'\S \liiith :i)i'l.HJu~lhC'I,,~~cril~liL'lHduring early ~k4V,U~~ !'ltp.gc,~IIh~"raj[l~i ~'II c:liclc IillO;[II::I'l I)Ni\~crli:cmiO:II can inl~'1I'Jbre wilh
tm!J19Cril"lj(nffl., 3114 mimjs com lea\l·t(llhc all1iJflion i {If ml::ot,.1."ul tRm~~:r'rn:; l }. ("~ll1l{Cr.;l:ly, ~1)l1l!:1:i· F.11""jLa'l:ly induced cdll.-yck kl'lgt!:icnillg run kacl 'ro pn:I~'Whlrt:' tr.JIll~~til)til1n 1,'1:~, A ~llm~.J~h I el~ c;.;c,,» r·~111t.'&>(lr, lim iklll rlllJI"-'ll~nL:I)', m'Ki rupir.l reI! c:d": model s can ~~i1Um
lor SOII1!J l1~llct'ISoJ i'VlZ T. IIh:Ylli[l not explain '0\ hy :SC'\ .em! g?m::..s nrc ;~111(.11dyiI~li'mIl!OO dtlri]l~ ~'ll1y dl:4l\'<JIg..: Mr\gt:; <LIlli \!lily .1.ygoLii:: gl.:.Ilom.\:

the ].' illiltlarrsl.ated

re!!llkifl OiIta rg,et RNtrui. th" a~mil1g of ~~IZ'f(ah,;; ··C1'!CI:~~ repressor

Pl'It"h il:".... th~ ;-;trikillg illC:[\;II~ in t-1.~a~ mnn'lxr !'Ilkr ~~T1ili~.uiGl1. most mTin:mh db oie Wilei r cm_d~1 >i.lL"\ldOp]]l~n' w· rup3cl :3l1d .':yrlirlnrtllmlilli ~dm t'yd~~ ~/I. \\,h~"f.1..'lll. ~.~ ... O'\'ll'roU, :Ul'loon~. or !C}'l.Oplasn'l irl th~ cn~~rytl. ,[TIJ!ImjJl.~ constant, tim J1llllllibt;r of nuclei aJ[liJ 11.HJ

ocl UV:dliun i:;; a g.rm.l!d~ I lmr.l gii.~an:*J1lI."Cjli(; prOOlu"$l'i i

rtf DNA i~lCI"t4J!~ c%po!JemkLlly, LJ~lrillg ~his period, mltNA~, ;illd PI;o~C,jW'~~cl\'ided W the (g,gJ ~1' I ~w m(l,h~r dp'h e dcvG'k'11fllCnt Bjl '(;-fl:n1r<~t. ihe eml'l1}'Q1~~c gt.'110fl1'<l i~ ·~~f1.~",ti l:lI~iO:l1aHy aCliv:L1Cd (!nl~ a~ iuler AN~, eell c)'clo!~ This ll"llm,Wflfl :lhlUl 3
anltlU:f!.l

model"). TI'h~k~y t~ll:I.{!Ir:>D~m~ uecd ~ohe tUIl1:ltt\d 1~9 II I. TliI~";:jC' OhliCM~i:o:n:> :Sll!l~i I~"'I sonre {If tlre m~'clmni;S!llllf> [1f'l[k[IYUII~ .... )'~!::!'l~C ~noJTlI.~ m:t: still ~ULlSi\'!t.nlJlt i,1 is conceivable lHmiBl hi~III be di[';Iu'Ovm-,,"!.L Mon."Ov!.'T, ~t"J:il~~ ()tll{~r dtrorml:tin 'W:llIlX'TK:IlL':l 111ll1:ia1t~il~a!:'1.i'l'~llroll reruuin or S ilcm:ing muil a critical :Jnll.:mnn tlI" ON A i~pll~~'11L ml1f!lnll~l~iLC~h".h~ ..,ygrl'tic- !:!1U1~"rirtfil!l m'o'eL'}' c-.111~' dCII\~:brc''''[fIg,.~ fc',!!.-, 1!1C' mnu~C' ~L'i11mnlC is; (.,r. 5)_ Gt'll(_1--"'Pi:"C'il.i~ DNA .n:wnl'lyllltiLm 11:11>[~SQ been itllpli:caltd ill repres:-jem. iJepl~~j('!~ t!1i::' jlC'thatcd ~~tIhe two-cell st~L~C) j It ls COHmC'l,hyhmll~~.;'_'lTL~nn111~1 ou.pfK:;tlfn, ~ Lr.;IIm~rip'_ celvahle !h;1l ln !1:l3nun~l$ there migln ilc !\ill~lic~m~r~l~eo assemble !rn!~!lCf~ll,i!lfl t cflmplc~C'$ al .. i}, ele ...vr~gc ",f:a.~. bew

fir

n.

mil!~~'L,'111-fl :li',:}I.~L~~icnn(jidc cl!;,'V,.~ope mcnl~ Irl[lS 11!I!C11 ~ 11100 ~hcl'IIlidhJ\rL,;;tul:J c

Matom<ll·~
tl'lmlliIQn

ltm'l

~lllns;i:~kill INiZ.) ~ .itlg. 1). II ,anal ,,~ililCide:; w ~dl I,;'CH !';~i>C'I,;:ICl1J.1~fle!lhl~~ and i1~ d~g,!;;_~~lat~m m:wlj' rni.1nl>!l-mkl ur
R N A~. U~re

~ musi ~iol1

(2)

orrill:uem;rnJ.-zY!icot Ie

ullllkn;l:JJI1iJiDlg 01" IlllJ 11Illdmru~rn.~ "Im~ ,[l~ul m.k' 1l:1!: inl .. or Zj lllotic RN iU and b [hJii dt'<llh of mmlCnml .RNAs. AI ICib1 l~II"CC Il'IliXhani~I,1l:;; ~...ve '~c~n inl]):lil<lhXI~n the ::;,ilcm~in,g. [lr tltc :I:}' s,m ~c !j,eIlOlflt~ duriug cllrly dovt'l.opmeDlt:
~L')]'i.':ll>iL TIl.

cm~th,; !;)';'U C)'!,:Jo!:> an: \'.:4)' 1a~1g. hi.!! Ihe esact 11H,,"Ch~~s:~n." elusive, ,~I'C' \;l;nK'reas l!}'gulk mfilN' A,~ are ::;yn~l~s~cd duri!lg 'Cll'lbuyog,.~es.is. 1n!!!1}' nl!![CriIml mR1'>1A.~ dcgi:rndClL iD~,d~~il~"I arc .1'L!n1(1"(.'S, gene ri\1!(h~C1S Iliku mlgJn :im'CrfC"r'c wiLh biter OCl,'dOPll;1l1111, 111'i}O 'Ilt'gulllt~d l~llIl,"':rrIll~IIllRNA. d",';S~mibHimtitm is
111100

review

eur t'i.Jmn ~

u-::Im:huiI!:1 rcgil:m P' UTJ~) RJ:gula~ol}' ,RNA$ {.')f proteins sueh as 'S:~lmlJ@.
()I"IEDEN':lBir

iali~t1 [~)'

i>L::q IiIllilCJ.:....

:in

u.n

IJIi.~

J'

1:1 n-

induee

I,~I~

bim! W 11'tc:;;C •• ..._.qIL·m::~~ [lnd d(iwji,;Jly~,u'iol:l of w.rgilQ

l,hL:11

!.'>C'riplioJl m:wchin~l)'.. om:! ,(in~ ilrnn:!>,tripti'['!'JlIl~ rul)m'~Eoll

(i). c Inol1mljn-nrl~j:! Hal,ed ,(iii) dl~li.cil'llCi~';!" ill th~ nnrs-

'These RN~s.afe degraded d'uril'l~ dil~llenli!l~:.ia!ges. olremhfyo9~mll'Sis.,. llildl(i(li nUl b~~ ~tl,l~ ~nrlgoil~truiaJ ~~iI.:g:E!$~ nlil~ ennb;ry.g ni'< !ll"oom:e is, ini1iaUyU'.ammj)ticmafllyirraadiw IIIflm!he MZir, l'itl.e;n

f~g. 11.. The matem;ikz~g(f!~'i[ lr.m~~on (/II1lli). Ma'~E!m\1l IltNAsare deJXl'Sil~d fly title m ~i1~fiTl't~ th~ '~!yg ~nd dri,'l.i'el"arly de~li!wp:rnC'I1L

1~11RNA~,

whi chnre

r~loo

10

dt~gl!ll-

d:lllOIl by 'ulocJea~cs; Reecnt ~U_ldlCl\ In .l:.C'hmlLJ~ have i[k"l1.~iiljli;:.:! mtcruRNA the IIIl_jR·4.3f~ as 01 ~O'll"~1Ii:J11 lun'" b~lWi\!..·n :Z)lg,QliL::ltl:l'Jl.Oln~ m;lU\. aturIll :miJ dill u.~cay
ol."m:flt;·rrmJI

zyg'i>tk g@f1l"S <!r@sl<!r~llllg 'I.llI b@ I'rOlnJtribed_ Of .:J [110[1 uon IJ.~ ,,'pilh:{;'H rydes, T~n:: first l~v.itllJmc rOF ~~olml iq'i'fCSSOr hi udh'lgtQn1c.ltylillc<1 .[JNA. tqlJU:'ll1;r.i1'lii VIliS! p:r'O\ idC(1 .2:5 yC'IIIti. .ogo. 'wltLiIl i!1 Ne"'I1fi'~ fluid Kil~hg]cr fCJPI'Inetl thai ;1 PJ~IlI'!!i1h!!J1: I.~~'~~~!l> II'lC jlL-ellliliture ~C1hlllion flf ~.:>ub::;ct i im;:IiI:'.j}~~ij1 ih~nl.llulx.'T of IM:ld m- illl'i!all tG!Lmtof (Ii'" glCilJ;!ii. ~61. lllc~~:>tU(hi.e.'> ::-Uggl;,-:;l.l1.11 Imillipl c !Cili'tm'illi in-i~wdiillliJl.'rl ~.t\.'Chaili:.Il\'~ j:we, !!I1~~JS(k1 i'" DNA re."'LiHed ii1 r1:em:Mlllrc inhiJiilifili i'if,ecllc}'de !,l;L~milll.. ~~1i vlllkl,ll, k~lfIg~h,i:lflhllJIIUt! "'}'W1'~ic u'lIl~rirllti(111 iu ~\~'fjf1fIH~ A d!::{It1 II 'r!lriU~i.C!ivmi~invf OOi.llp~lU!~m1'>of!!1 e t:mbJ)lls j 2',3t lb ... r'~~I::'ffiI1!.! n:iDh.-U:;;mtlie.'l. ,,~ ilt Dm;.l'fJf11li/u :md ~e'brorI5l1.Im~'I!Sl!.I!!!g,~.l;;lCd illlli lJiJlll~crlr~ ImLcilillCt}' Illiglll b!; ~l ~'>CcotKI i:ol1 ~h!! llIClco"cyl.Oplf!i.',mk: mtio !lii'KIll:IC I i~~n~i(;ln U of!l :i.'liiltCgy tor g€'J'lC :"licndJlll; l~I:Ore MZT (d1:C' u:3nsc:ripti,onrli ~pre,'>l)!Qr by IiIe C-XIlOII~l1.U8Ii~Y '·1hllhcd Iltllci1 b1,Ci)' iJl10(t~it'l, :lfot CXnml)I.C'. IIHI!L!r..: ,.J\.rlrc.~i(lJl TATt\-bimli~lg PWtl:ill, in"l:C,usiu,!!l ;mUJLUn 'llf !;,I.:m'llulic :DNA dc,l:;.:nnine ITEP). a c·tllt1lr{~~t~·<rI1 of n,he imT~~1:irl~uj;ml

mltNI\ .. U3t Mi('®lL'>J'.A.. am shon IfN/\;s dm1 blild to tl'lC 3" un:, of ~rg,cl m R;..ff\s 1~1 rcPIICS$ ~hcil' lral1818~ion .~Ind ooccl~toc~heii:' d~'lY. miR--430 tl:!rgelS ~'!.Cl'-~'oI<;i.11 I i!.mJr..'!!lu\i_Ylemali.~' rttl\'i~li!d rnRNt\.'i IlY bbltlillg
~1 Ctl,i)lfi~~nC'11l!i) ~riCS ili'l~ i'C~r J.' Ul1l:~!Ild
Ilh,:ir t~~tICtlykLtrOit

[Ttliino.tbtg

In

Ihl: 'il~I.'IIlC.C yf

MiIlfffii'l~' ,and (l!!liIJ1ar 81011;IIg\l'" liarl{i!rrd 5i!i!iI! Cl!U ~iI:!~1tlJi~(etI'bM I"~f i'II'Ollnl 5dl!n(l!,. il!roacl III~tntiti!'. lia",,~Fd 1lI1li1i{(!lslt}i,ill!\. DlVi(;l\ty lI\11!ntl~" .Roam 1:0<2'. CaiWrldgE'. fIVI. o.2:US. U)A ~.i'i'I!llil:ltnlef@ld~,
.[lEfl.8 rllfl'lefl,to~

mndllJli'f}'.

Iililr'o\anhcu

III ;:>~I~i!ll1;ll:y, rlluliirr,'1!m!.1chli!fli~n\.'> n,'gLlklIC RJ:\lA. !>ynlh~i", and d(!gr"Jiilat~ml durjn~ l'.::lrly ~Ul~jJlJlof ,gClIl".S ~..,i.• j~.Anmll1,ElIltIJi·I)'. ul1!i~Im'!5. cnlbL1'L::Ig.t.'um;i~. h tL;,'llmUn~ Ill! J[.'~,"1:]I~ ~H:!\\,~~ ·LT, s:iml uf ~\mn~ lr.ul:;;Cf.·[p'l~Orlld acti\,'~.tt:lr.;; c:mll~~ ~~ caJl hlJlhl'J IllDS,l '(11' 'ilK: ODIk'l'l;l(ld jllh.uI!ID~1w:ml can InC'oodou~, 1['[1111 ~It"rillliollin~ aCI.i·\ at~OI1" Th~c be explailled 11)' II If;',w key IT'!:!.~~~aIQ.ryril1.cip
~!I

ur

rp:~
~),f.:l

miR-4JiO nc>Ii,'I'r~. tl1!.'5.C' ium'>lA!i nu;!1!muf,m~ lind ill'C Iilt0!J,l,!11t to hu,-'1'k're ~"illl ~.tlill.tr} uuk 11D'P~lll~r~b. n:~gR""'JO C,.\IWcssJol1in i1i,lI1!:'til iI,·]ZT. lIlllS iirtll UI'g gCI~('lIll;e ;flc~h ioll :Jnd ntI~LlCm~~ milti'\ti\ m
d.·~I~lJmio:n,

.iniilu!;."C :pretn~ll.'l~ acli\'.L~thm

SPECIAILSECTION
ples, su~h :1..'> the 1Htc:lco..cylof1'l:ISH!I~.C ratio or UliC1tlRNA"anduccdi rn~{NA (ill;:!!~oo!in!ll, FU,III~ :su,lIdics rn~y :11~obe !\C~cvan!m tl]cf1~ld of ,mi~l1q.l::l1.1lflr~~g..Clm~hu~~;o,;IJ'I~.rim~li!:.~ rely on Ih~ (
:l:. l ".~~ IKijf!l!;hlll;f. (!!If ~,@, ~jl~ ~l!)(!~'_ PIL s, ], Mmporl. Hi, :1oili~5cllll!1. rt;M ae, 6B7 ~'l'ilIlZ~.
~, M,. ~I f'lli~~

l~h::~~(e5, ,and NO~e5; L P. HI. ~n~n~.~ swmpn. V.I, Sl:!,C!!ff. BIa,I, 'lA, R3:§ 12«1<1.1, 4, N,KlIyo, A. ~, WolH~.J, (ell sa. ::I,l~. n ~2000~, J, lili!tl, :A. S!ffilel'1~[, m. ,~~~!!. t"!I'i'l '11, 43~ ~U'AL

rer!TJ:!!,r.u~nniu.rJ or dO;(l()r ~~ucll;lih>' ~nl.!!(;k"' ..lcd C!l,1!R Thu,..... lcmili~ ~ d1<I1t ,,~I~OIlec.:,: ..he "')'!~fi4k1;,1)}replllgI"[L!Il' !musrCin,:d uli.lcl& ~. ;>m:c, h LRl(k~llldinjg t~lc'lllocl"'mi~n.~ ~hi!!~ llu!'h"JJk, 'Z'jga~k g~ool!TlC' s.ik"'c·I1'JjJil will i~!fOllf1 alu: design .~'PI'fit~nll!u'ti~g SlI1.ltc~i~5 (41. [klVlll.l ,II~

7. ~ W.~li.ffiftocfl"

or

,6. Ii.. fill!!J'J' ~[ i!rI.:, Ot'JreIllI,I1I1l!'llI :13,1,~l~~ (;!OO~l, P. 1'1, 'O'FiUml. Cdi ~7. ~~; WlI'i''lJ, II, 8. A. ~:GI~~r, (j\,5druil~llr.all 44., sn !1l:S'lWi}, '1>1, ], 'I'~iig. C.T~D, R. S. lIJ~f~~n:, p. ,._ Wibilii\, 1'. S, I}tm~1'II'Ml :10'l9. SN] {2tKI2}.

~~an.

l(l. ~ ~ Pri1t;!'Jar:{\ G,. Sc:ll!.'!!ll~ff. Getil~ O!:v, :!.~ J!'lll tI.9%), 11. So. M.l1~~!'I ~J Il'£ • firJS Grn,,", ~ 2~(.IlOO5}, 12. 'Io'l. T,alil~Ik5' 1,1, 11. l.l~il7, ai!1': ay"" :!:~:2,59:1 1200S~. H It. J, Gil'il,ld~ f'I" lIt, :>~. HZ, '15 t200!l), UL I ql1i1rlk~_ [1Qai!I1'. Nl r~am:i$,. D_iKitrnilrn<!l~,1'1\, ,Mid,ael 5.. MangD. 11"_iiaileti, ~ Se!'d~w; ;md memlile1j~or In!' ' ~ r.t!r ft1~I~fui, diirus~n~,. 5~~r;I tir gr-a.rn'I3, liml ~[H. tl1l! HmI~ it Honl!iu Sdmct Pr1l~i\lmh ~Ild 'Ih~' rM:rnl)ht ~iidiIWffiI!jjI f1j~i!l f~ N~fIlKl~n[~.

REVIEW

Ololcy1e

IRagul,Btion of 'the -'10 ,-,Z,IO ote 'Ira nsiti 0 n,

M~k,ilJ. Ma~kinhind IJi!.: c.tlJ'i~l:iindinl;: 1\1t11~iH ... elF·1 E. !}rr.:~~"lJ.lin~ Ihi: reeru i:l'rlll!..'111 ,['jr dl~ tIIII!:';' IUL~c!11.iJlll imiuni IlIc'LOt d I'4G. During. ~'!OI:yw rrmlf1clYUlll.'1~llJi.!iou run flI.'OO.IilIbl!..m{,r f)i)11YV\.j.;bur~ill~ pro,dn OOOIJttl to c~.f4Ci. .t1:I!lOflliulg ",IJF4IG dis1~1i!~C::; M<r;:;kin from ~F-I:E. fIIllo'!lllllg 1~lnlmK)111 of dl~ nll~L~ni:Ol1l OOD11Jllb ,( II, Jb, IlirsL 11.'(l[lIJ!jniz,,-cl lUI clam lXICyL~ lH'i.
'Ir.U151~uitlllml acliv~lirjlil
Uli:ltioll, p~IOSI)hm)'l!ali.ul'J. of

'[PEB, ~1.iuil.II.1~Ir,,;:s

E":
oID"
Oi:'CUnii

feimi!ll~ germ (e~l$, ([III1M!n ,aU tile messenger IUiUI'5 1iI,~,Les..~aryOO' ~t;:lirtill n e'l'ol'IHe DlJt typka!lLy wait lunlliLr~rmjizatIOll1tu' be9~n ~1f'olelOpl1ll~l1It. n~tr,clrasition lrom oOC!!'lle to re~!]ti~~~ '~~J!J(z~'9ote)' lHl"'D~ve5 mcal!l~ cJ1.alllg~5,. i"n:duillJlg IPrQ~e'illl :s;.'n~1'IeirS, pr~t~'in arl'Cl :FtN:AdegradatJ~n... and lJo~g;meb r'~liJlod~li!'ll1i.TheSie .dh~m~'e5Gl!l:'lUrc:oru:urrel'llly ~'i'ith 'th,~ m~ial"it diltij!iia~s~hi!t luo>tl!.!ce tlrle hi!plo~d mi~terml'~ '~e!'!ome. I\ccruITIulating elfi(l'eflCeil'ldkililies that the ~d·cyde reguilators that ulll~rolh,e meiClliic divisii(m~ ~h!l regu[alelhe t ma.f1V ,th.mg,es that ,u:Clom,paFly!he ,oocyte'lo'l~gote tr~ru;ili:o!il"Ij.'!I!!! sllg~est tnat tne meiCllijc: machijnerv rUllocIDicln:5 .;n;al~ ijliJle,nal pacemakE!r lh~t. prop els 00 cylie5 oo'W<!rd emb ryo~e",esi!.. ,rmrli!U.' .11E\~~I ..llingri.\1nH~n

Oo~~te-5, t'he

~lr 1X1C.~ 111.RJ\'A... ~ ·(jllk~l Ir

~Ilk!">d 110 porIJYlAl-t;lil C'\WU.'>ioll. lr:Jut the' ~o rn!~ Ilrsn (l!CCi!ar ill:d~i'lill'd(;'I~I~', For C'xill]lrl~. in f}ifJ!iillliMi(f C!M,.>;!, qdin }],rnRNt\ i~kcpt~un..~~[km{dty silcnwd hy Ihc RNA hJnd~l),g FIfOIlCln.F'inlli,i:io

Oh
{-r.

<.c~,\I:b~,

P~!i!'l.l!!O~(tl".m~n.ltI~ ~ JJj\N i(l'l}' :an 19lk!W'\11 1l~li.In~~ml

ou

k~~a~

'C)"diu B

~'.O ~,

!lJd~iJ1,g !\tml~.\!,k'$ 11:1" :gene a~'1'i(~tkll~" g~'lC' .., ill:"! h1UiOll,ilIod flrgrnl.dt~ .~ClnQ.jd~t~.
:I!ill wUith: th,1l' New, ..

mR;>.JA h':i:in~l~li(m an~1 1r(l~>'~Mllli~ C'.''''~n ion ..... I h In l"m.r ):ill nmw:rIl,~.I~rcoo ~'l;.pre~l>n!1f o ro1YlJ\ i ""lIYl11.'!!!~e is: ~LIOki~u 1.0 re~'\CLtcpt1Iy~

i h..l'IJI' docs (m egg bCl:"O~no "e cryc

IhinJ,!"? The jO~imCjihc:;ins \\ ~UI0i1lCof ~h;,:most C{1I'1'~11~~K II I 1:1)_11$ ce r(lm~mkltls in biL') I,~
ffilllotKkiling
01':1

I\.::'n.illillll-a ,oocyli:' into

::I

tOtipo'!~11Ll.~'~DIc.

:i 11 lln~~h.';(·I:K'!.1' of

rL1lru (kpo.l'lKl5 Oil. .[Ict"llm!ldm[i;d In ~Ul!!(I(I(.)'IC

Rt::m~ukaMy+ Ih.i~ n.1lI1lNhitm trdllf;i~rilJ'liul1 :miil thltl""" m!..'l'O:;!.:.'n~·r RNA~ lm.!lliA5)

U:LliI1n~(iIUIl!CTlC.'l]:;_ .r!lli~y IIIl'Hy Qr RN/\

,gM ..U ooryl~ tOl1l.uin'1 cka.ying ,

nlt.. ...><lgc~ rom~~fIO'l]IiUI1!S to '

1"'m~ f2.-H and 55%

gem;:> PI! These tml"l'lCrip~ ~ick (J>OC'ylll'll d{lri~ two makeo'IcCI>;['In Ih~ 'O\~.Y to bo<"t"m'lfling zYb'o[.a;;· otX)'k I'fmlLHn~inn :mdl !..'gg "d.i\ :I1irnl_ Durin_g ~~bc: rnMI.r:J.llk'!l1. t\,"lCl~llrllllar ~gr!~1]l\ s;tlrnilr~n~ otX:y~.t'2>
:um..."il!.cU in pmph:1~ oJm,"iffi."'" I
rnllOi!.'i( [Lill! ~nilfmc
[(]I

or nUl .DmL'i(jjnlii(1

:m 1.045'1Y.,oraU

I]IOUi.'H)

3(~:4i'}'I[!I'ioll fir l1'e'li!l BlltUI FK~U~Jill;;~uttn!' ,[.J~ ]JL"~C1iou ol"l.mlili~h~ s ij~ 'Jill'iO;:1ilcc'lr~ct trrlns.lalim Q~)'IC m~l~iiJrmiooil'l!;"(,Wirli.":>lc~ml~:is d ,of 1J;:W t;;ttSlOIOO. btu ('l(llyoctaWkLli(lll is 110'l These ptol\i]ilu:>.. 11'lL'cr"d.~ndClni lr.m:>lalim~[lJ8CI.ivmiolll ~1T\<;",i(]ffi ;s[JW"lit tl"l'i'I ~tl1 pOl'Y'dd"'I~I~mir.oul' ~"crrL!;; ~1S1.1ilt;' ,~.~I'I Dm:lIl~IIl'"an: I".mIlQ,;ti In. all!::" ildblXmkrn m~d poj1'1II:1l-I:IylatLOIMfqx"id",'U1 mO\.~h· :lIli'J1~ .:I~,~J{~ Ilr'.Lllsltnul in ~ltt)"lt..."-and a orhaliL'l1~ OOm.,\::lii\lL'I;.",,::>:~ioll I;:'). For t::'t3ll11plt\ 'b~~1 dmill£ (lOL,\),'W II.111MInioll,in li'[HJI]~lt\:, 1.!"lliflJ>!irtu[!nI or ah~ 1m nl'h: .iinuR! wi~~ be tn dod'lIll di~ r<L.'1[luirun:l'II:'; [or each .[Il.t'Ch~mi>;']TI_ Ant1l.l.11'r iL11rK'n~ullt (Itlt~ ion t'ytlil.l41Jo..11Il1.m.L kin:llro ~CDKl ilindiJl.!J. I)rorl:lll R~NLiQ,"Sp)' ,rrivml:<i lill<lnll1L1i~m p.rollfJUo1jng j~rlt!r i" [I~L';'r;1"'K"~.1 W~I ic~1 mic-roJ{N t\;s; CC!lllr.ibl.ltofJ1.0 10 tMPF; {'UKJ ~c,yell[I1BI, er.nplc:o;lc i\1::1:iv"c MPf U;. ll""olm;l::!t iOllIl! 11\1.l[(".S.."liou in i[)[JC~ 1\. roccilli ~ul'l'CY uirn :-;Iirnlilll1~ Iillil:LL •.lwt;;lmkn {)j'prolcilll~ .I:It'!:.ii:l·d it) {)IFrLl;.Irl.y I000 .l)l'r~.w~piljf(J,~'I{)Cyl1l:;protl:illli fOl.lnrl {!fl~y 4[>/~winh i.OCR"ll1llctil abundancc in Djo,'l" 'UlTU' mmnnmIL1 1l/Jt"~"fIt\C I.t nn Ilw 1111111m::"J:iI ('jocyre O:I~I~::t (!Fig. ~~ [luring ~~ atil.i,~IlII;icm.adtiitlomd RNA.... ::!Ire 1:.aI'Il';',~I~~'SIill~ dmt I'llteroRNA:. n;'\~,d~[1.:: 1\;truitl"i:llN ICllliSl:llLtlil. A ~nldy col111).:IIing 1~1l11!'I.XI l~liL'IDrity nfmRNAl! in ollCy[~ Ill).

on

~'l:'''''

oot':)'~
[I]

mid

~.!}ff!Pt('lo" .L't:l\ !.."31~\1 ,dml]lati~clilfllMu"S

Cll[('nHciotic

~.I

po.~)iSl.m1C<lli.'J!._'IC.i:llill
b..,."weCil
\~,'C~

lite mdOlk dh'i~iCli\S (5),

thinJ 'Df t~l:ilC'ri:P~,

lrun:dillkln llo!)~~r)(jl.

R N ..~... wilil] l)C'arJy 0]1" .. t~~)%l ~h~J\',ling ,H!lere~gi,,1

1iru~cril~L" C!~i~\l,;

••.• A,IfIi!l1(I,UIt. w~tlii, th,e OM

TYP[;cfflUy, (l('l;CYI~S ;~rc (lw~m~ :md hi.--comc '\,:fi'1fI1pCh:U~ j~r ~~gl ..~tkJl1 bcrtlro: l:e"d'iing ;\ ~">'X(~lJill1'!i.."'11 110ilUI (mci~[:lhag,: ,of Inei Ol51:. II iu m;mml~il~.J,I:::gg oc~i\'mi(j~.ifl1~i'I..'tl by ~Cinl \;1111')' bll UDW!if:Io;."'Ck"ii, QOtnr,k'lc. !l\1;,' Inll'l=>thunmlil~'1 1.0 1}'W,1l!;! hYi"I:!!il~lillg Ih~ ullnrJ>!.:llflal fir mdosi:s" the fO~'i~~ti(l!l protillCild. am:llllt~ Ilr51 IIIitOlic:dh, .is.ion(6), I~llih~, B.C'\' l<!w. \Vii! ,ai$(l!l~ the !Cimng'CS ~~I~I, ,lOoon~lruly each cyr~hese h1:~'1il,i(!~k>, s

~e. .

LI,lC ~WO ~1l."lb'eS (lit Ctot.]'tC

or

11)~p.ariJfliem I;I~ ,Mo!eC!i.!l<!r 8~ ~!dGeJIll,tk.'l, ,!l1l'II HoWillld HlIIgh!!s tlImkiil. r.iiStlli.lN!',)II>'lm 1fllc»1~I~ ~(~~ III MiI'liIlclo; 7:l!S ~Di'i11 WeIr!:! Sil~l PClI~ 1~, 8:I,1.t1i1i1Jl'e.o .MD 2m;;. uSJtlfr.(idliltl!illfal 'fr~lrning ~raiJI Inl HwmMl .Gi!1II'!!ic:! and MIl!1e.:~B~ •. ~Dhrn~ EiliJJ11:1~,~, S~ 01 I'.IIi!i!Ildme.. Morill WO!r~ ~~;~tf'£TIi, ;roo, Eal.!lmo~i!', 1lV2l!2GS, USA. 1

ns

prol~Ii." iI1111~~~cd ill cclkl1;i( h"'li ilCo:;lm.i:-, whr."t'C<~ ~Y,b;utiept"iI}",Oi'l'l~ \\~ ":nri,,,'il~ li"ll' iJ'f!ll:<Ciijti;:, irnr~i~,,[!i." ill IlUill;111~'lccuL1r 1:liu;:ynlh!-i)1. I ~'D\\ ttl\.' YtX;YI~ !!tIl': N ,'\$ :D!::Ii~mccl fr.rrtrnll~Umioll'! In 11IItiny Ct.'[:\l!S, ctel!ilo.'miJlJrl tll.:pcmls Q~I Iliixlfllin,g mRN /\s fronn, Q"Ill'lP~i:.'..tCsIImlbkx:k IfiH1$l::uioo ill~t~lkn ~7. ~). ~(lr c.... lm'Il1k. ill Illlnl-se, d[lm. ;md. .\·('rmfJ,m 09CytCll;, IIlRNA::; l~'I~I,I,,:OI1' ilii~lI ~1'u:.l[lJ;N~lIi.c f101y.L.k:llyJ:L~L<tm..... ' IbI1Cill~ ((,PE~) ItnlJtciu" 3' 1iI:nlF.hrl"lmoci fl!glJl}Jl1 a[1l:t h1lOl'Jc-d \\"u't1. :; ~1D!11 Dl}' "I.I:.J:ny]IIl!.;'dllmly~ A 111:li 15 :Inti b~111L1lId p by :I lr:U1:;hti.CIn.~I'of...1ID"~ing C'oml)lt\x COlll@Jinillg
I.hc CrS.!JinJlilg rn~mcil1i (CrEBI alldil~
p:~!;ftDlt'r~

C'llI'khcd lOr

Ooc),le mlu~jrnlkln. lind c~ ,L!!eth'minll ai:s.o 51! mula1.e mRNA dei;i~d;ILIDrl" PHk,;-npcNem ~1f tmn.. "';g'~r~lSI'C" ~kgmdoo odi.!rillglu~m11:miPil ~ ill mkc V l Il.:~ll,,~itln i" .o;.ck,;ii\ ,r; .MW lm:ic~Uii~~*,
rc~I1!;l\ ~ lni~l~crJpb

rcq~I,ii'l..,\~ !I;'[j'rroplw!!

Ilfi\o.:Si.

(31. Pul1l1l:.:r ~&!grucJ~I,'itm (!I,,:C~!1'}j af1l:'r ~.enij~iz:il1i(lu usher the ~rualslllrnl i~~' l{1 Zygotic cu-mro~ 12, ~. i.. ):I,MC'dl;-mi::ut'ls nor RNA dir-gmam LU]'l,LltJ r!l:'iiv ~ii of zy!;.olic Iml1i:lcrll)'[itm loll UI-C d]scus~ ill f:Ioll JCoonlp;!luyirl,g .R.C\ ~e- (/41, ....
III1;(J OU-:::YIC' m~l~lt11l,1i1m

Pn'ti.'il~iAI.-C'I~~ Inlllctoo 1{("(~<1tlnlimn

dir.": '1oOC~1~~iU-~;go~

dLlrllJg

1l'.JU1i:liiiml.,r{iiti1r!!;.iil~~llb {liill\: lh.iquilan.;pmt\.':h~'mi2"Illlh"~I;' r!II,'Il' wdl m;1~liLO:.l il~l.l(!· ~ylC" IrEU!l£C:Iiplnnl.:l • .!~I ,111d f,(l\ ('I.'ail... 111.Qit:> L'uvc J1.:~D1Od ~!<:lI!l1Jllb; or Il!:Ul~iill IUmLwt'1" b~ 1I11OLL';C ~r5 .I {~ Jj(t'Wf1H~ (Ull.:li1~bT ilih ,r !9), aud .d .

WW!.v,.sdene:emag., org

SCI ENCiE

VOL 31i6

,20 APRil 2007

Germ Cells
ix'C!1 dt.1!CllnlCnI~i In :'iL:!lcra.l Org.1JIIL'i.l1N. 11'1 0[111.'1;) Ell-rich mitoclmll.driil~ cloud ~o ihc v~§CI'Lkor~i'i.I1<I!,~bcctllinkc-d to 1~li)'lllion or~1l1.I.ifl~ ~~i111iIKl.rl~~~~). L'hm!1fl O~X.lIk 111;UlLI<.I~il.m iu -,\"J.r{11r.I-~ ,'1l1~~~11f1ollCl1(;'. ER·tim"! ,rdc~b..'lt.1 abtlH~l I~ln i- 1~1i~Jly d:",~doo ~~eri'l,ll;.,!~ .. t's.ick-;..~l'id aligll uuilC'l'1C.uh II:~ fl~~!illil!!i m~milrofl~~J3 "U1Jlbrcdj~lrillll!:irl'n ~11<l~ I'. (acili1"~~ dllr.in,!!.11li:'~~i~ I J HI, Injoctrfm of :L dCgI.drnioJlC~ l.~C' .m.~,"tl!mfonn nJ (,PEH i~1 .\"!Nl,lfW ~I"~ iuwrc Ilmll!l!F!.tioH of I( '...~ ~V1I\'~$. alfu!tilizminn r~l'I:Xl \",.i111f;.lfNIa!iill nfc.ydin B ~ :UU] mgn.~L1u beloVII} b~' 11lL1\il'lf,l en2+- ~fl.l'e' do,~cr 10 1ho::!i.itc OiU ~ [0 meiosis n QMl S;ig1.1il;;1rly ill C d~'~m~. :ll!,loce;;sr )lf1Cl1tl c~~l)'. l\i"icr egg acti ....... iflU. cnrtkal ER luI Ir.Jlh;.';itiLIIiI (I'mllll'I'I!;itJ(-i~ quo :rm~lt)iiii:; liI,ljuili\;i; I,;hl~r:; an; R.;·irl~rniI]i~cl p.i). Pcrl'liIJ)~ IiI ... m4~t dr.IIn.ILt~C'r'!.3rr~jlg~'Tll!.:111. 1:~l\TI~w\!jn~ ia~jonit-;, kl,ii:l <kgr.dillirnll C!r Ill.; nIL'l.::IUlJOOIlIc-::... ..., ~-ring: mTllrJ~ MEl· i M EI·2 ,Pi)!, MEl· ~ ;"lId M E]·2 0'lIl: R.'tILlUl:1.id I.hl: mpjd rUi>Km 'Of ror~icihl ~llll1ll.';:i "~'Lh A.hi;;" Ibr 1II1Cil..i'(;ll>hili. ;r Ir~im;lin~d durnl1¥! ul~lt~i::, p~~ISiI1<11 1Il~ll1br.Mllil tn pJf!.:\'~nIp(lIIY~P'"'rl1rry (1.n A iflL'Q[thc with gnm'O'lh or Lll.:' lim n"Ui~otic ~JiITld~, Sillilil:Ir tiJ."luru Cl\'I.'I"nwm~ ro!.!~]!ly di..~ilio~ in C 1."ro1~1[1dq:r.rl.ll.itl1"1 is ajw ll:!ii..lI ID ["I.';Stocl tl1lU1.cim;~.ll d..gWlS irwoh'"iJl!1. \'{:;:;_idl:iSridl in ·c:J\'(ulil1 ,!JT~, ' :.:Po'ci rIC [U(;~~or lIlC ~lm. For ~~jlrlp])._~~Il D!'().\'rJrJ1' ... Arnd il1imJlii it ,J1,nt Rigid i/o. C dt'1!.~IfI1i, ;lU:K.I zdll1d.l(.;h. gt.TUIl1iul~]('{:ilk'
nwmJ"'m/~{j!N d(f!"wu' (11)) ~h ..S fll1~l!O]' ~II~)I ' cmhl:")ii:l:ii_ 1'J{~t-ci~ d~grndalrnu ol'tc!~ :iC'!,\'C:S ~~ hl;lClh i;!llC pm1-ci!1S lh;lt ,Ire Ht'I.':dcd COl!'!)' ill Ih~ ~r.1Jn,~li(i1l 1r~rtI.h;t! w«~dd I~ 1~~!'I~l*1jl~_r.fI,T ~ ~,,,:u~~pk {':PEB ~l.mrtit;:.!L ...C\lrly U~~l:X"y1.e :l.nahJr.1110\ 'C'

X!.!IIi1PfJ.t.lx:dis~rihl,~lunn

UoN}- Simil~rlJy. IIlIi:''i1m.HFL,u.ion or ER. gmnuks d!Jrbl~ egg ""th '"-inn ill mice (rc~td<;, Oil ceU

c}'d~

1\~l;Jnll~klll

(14~ ll~~!l' ~I, ha C ~'kg(.I~".

1ll1lit:l!lJirJ!l:O;; 1f~ ~\'CI~llh~ ~ t~~1 l"Il~in11~ ~1;i\'hi:l"n h1o.;k M EI-I d("'gI'n(lai.io.l aud Cll\'cnlin \:~~'i.;1c ~

lil;:.ton 1125.3m. nl~~\"~dy,

II!:e-!lJi;I~UJC ~!'l'I:!)'

bm

M l'l~c

hYI\!tml'l."~ llnh",

\VE!!-~j!!(]lfl':'I~. rille- ~1'I'~liwihk: .dCj)cnll~tlCC: of ~wrnJiL'lI'lCC~Snf~~~' CIf.!C'}'1.C4I}-")'~QtC' uml~Ltion t~l mcJm;c rm)~:,;i'~~l, l>L~t.~ Ih~1 111,,; '(;cll {;}'I.~i: irl1l!c!hln~ry ~~rr..:,:. m... :m il1iJ..TllIJ~ ITlllli'il;r liiJll.cr 'lhlli. d'IH." snlt1'OOm'rlil1ilt!.:;;. Iii.: 1!tdi~jlirnL E~ ~I I~lil,)' i, olreil cill.:ldl,IL~:I ..:nl~ill1 ~llIr.llll\.\1l.!1r L s lot Sml::~og; in a~...i1>l~ n:,-.rOOLL'ti.v!: U.'II:hlllLlioyi(:;;;. .. .
00IC}1C'!' (:61y!<1Ih~ nlLdi:icu~ill pI1tICQ,1...::iS
ql .... I:ily rt11"U'lnLH

li~~~ MEl-I '~I.~I<Ldnipn h~inu~1!a!~re

il'l~lil)i!{l~'

MPf

E~ acti'l'Ili~m. in 1[10fitr.'lt'ot. dtllX'IIC]ls s_i·~nrn.lira~, ilil!\~d..: Ih,,; ~~~ h] nmnln'mls, {'.ll~' sjg!I~1i I1g m;lY be ini!i~t~d b} :l ~cnn[O~hm~C'!lC)'in Ii1w "-Y~N~, I( '~1111IlllC~l:n~fwpmll.'Ornlc IIRllm:td111;'.5;Wm be needed 10 re\cal nhe C:llICf~1 spocifi,c pl·K!:;;iIlhf.l.1i~)iC' (F'LC{l'. which xt~C vrues pho;;;phoiHosiaide s.igJ!aling Oil IcrtiliZtll tt.1~1 .of I'lmlebl dC'~'Jd;,ukHl d\lri~l~ thC'o\"I(;jf(tH~.~}'gm~ ~r.~nsa!icm its OO!1Lrilllltifl'llL1J one) ~C'!fe!nQdel~!~, \('1 gel)C:lllt~ ~vCfilI ('112'- wt)\'aI Wh RIC!Th)ek;lhi.}I, mld ~hC'egg ~P'~r:;l "co~Drn" och ~'i'~V~, ith ~,uly e ~\ R!?!illufilll!l! WII~~fsTh!)[@ ••., re~I}l(l~I;.~so'utic.al g~m~IC'e.~{)(;~t~i~ 'U!.'Iq~lltring t (C\\W I)~i ~~ukm~ l.I'!1an kLl\:'r encs (U(.'$li!11p1 iLln 'l,r {·h;u~g.!t> during fl~ ~l;,_ley-1.O!-w-ZYSCile trufl~ilinu lh~ $l..'CLlild rnc'ifllk di ~~'l:im~I I'll. j .ore'nul ~$!ricl~ to l\i"1'lCifl~ ~!lIl! Il.N}\s but ..h.L1 ;II ~F(X. (lC'II,uI:nr ~l,l;g.lm'lIcs. The mC101L-c ~II,\'i si(1n.~ &\,\c'ffil lillCS '1;\'i~k'I)KC SIJ~~.thil4. ~l1i!1' [lmcl1i~l<l'Y cHdl m~tllY,ch~'IiI~~"sil~ UI~'ilC!C~'IC'n1Jcl cus, H1l!1 11)C:i(: il!itb1 R!'i~"'CII"'S,IhC' eelrevele ' j~nr~ oomrol:of slLl'1."CquC'UI. evens, hll' C:limnQI]!.:mi!lk'S, iu Ill.;: CYh!I)I~'>1n [lj1rl ~:n ~he cd~
on {_';l"

l)Cll'l£illt> 1111:di.:gr-drli;(I, from, Ih~:>(JI1lL1rit: :r~gi~ln~of !.'Il'h,.,;gillll'!, !.'I"~}ryCl ~.: 1 ~!nIhi:!. :rt"Spl"EI. :rJrncin J. p d!.'~l.aLiC!n I1fllY be v as ;1 "'ill,Y It'I, p!'lI.1il1d~y IIlml;!;: ~mlJi: Wlfllnlj.ll.ll prnJi'frann onil~ cl~ ~) i"mml!ill.:'
'~Ih:'

r.::~"tI

IllIDtUrnI.io:n

W~tin ~'Utllll1~~nli[i.Ll,l!i; (h""1i1: ('~l'lrrlg~! ~ Om:y~c l)' picillly i.~~l;arlcd by C':>;:Lmcc~1 ular

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a;;~I!JC~"ftbi,;' t!nl~ttio:n rn~s,...."S o (~LI~Slij(!JR" abou! ,h~ pl~~IOClC' l'L"n~m,ng Ihe' 111.d('l~is. :1 spirld~C' rlllm'l or i 1

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:2'.~L V41lmln~ (i,. Pit L fiLrr.~;"p..1la Bia~ !l8. S3 (lOO3]'. ,2':\'1. fi. fill:Harri~ 1". MilraIJo!lO~. . C~mlll Mal. Bful. [oeMilil. j <1M ~2I!1031, .2'5_ It '5ar1D oe!' ai. tIIal BiQ1. rel'{"ll. ross ~2006~. lo'i, M.l. 'itiIn!I. J'_l'EIlI!tli!!lil r.. ~D\, £iIl\!r •. 8i\!!! U, !i6 (!.IIOI,~, ll. ~,M.. Sc..~ulU" fI1!ptMcNrm :lH, a~:i C~"!i), ~3. S, .('oj!), L l:.I.'ILtmm tfNi~ ,~mi'~.l'. leo, ,29. iI.. t t;t;J(iy. f. ~o, '5. fl. I'IIll rI.!"tl. L.. Yl!.Dm" l. ~,kt> ~

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120. Ion

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pnnnuve
'L0111b.rytl:lliL[;

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.~l,,:rpllf.lg~n:'>,10'lfI,l.;lnGll, d~GiI!1!k~ls, anc! Cl,lh\ITC

condiliD~I~ c,~nIl~~~, or

~mll,U!l!;C' 1:h~fnflll'ltiflll,

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.i~~I<! llf im~rl.~t .D~&J~~te dl~ Jif('(l.cd di 11cl'C11lilltiLl!1 rc!~"iit'l~ rnm~

{lr u

\'rulll!.!' ILl I'l.~h IImt would r!':ililtv.v from .111 ill viun ."J:'Il~m f[lU ~lL1IC~ l~mdli1tLUm]. ami thl.l '[lItJ~~1111<li for :[lppJicalim~s in :lI'Iirm~lbro.'1:1il1~ uud somatic ,,·.di mt'plltl;gr.unl1lirlg. makes il lrunru] DD il::mI8i~I.'U" \\ht,~ t];~ ikld hLlll'i Ix:n arid wlllm.r il io,; _got;).lJlg, Is 1.h0l: Cllp ~u] f CmlIlJlDy or Imlr bil ~r! [nk:~.:.1 ia~£itll:rlCl!J~cnL\,:I~frof;l I:S cells \,'ila~

crn'b[)'llflOC: dcvdnpmcilt. dcr~vi!1gg:;~rm:tc~ sok:a,g~~ate 10 !bfm >-dilnc!ns!omd ~3(J I !l~!,I!JtiliU'CS, gell!: CCfl~Ta~ W rh;uirootc!Qc.y, U'I CX:[]O'CSiil ,!lItlC!~ Iy ill ,;hl'O seem s ;"~ d~Hl!l~.i!lg dl~:lIi~il~C',O"lll~"I~$ can~'(~ e!nhr}'{lid U:iQ(llcs (. ~). To v(IJ!}'~n~ 11'!.!orc$'Ccm 11{Iuwin (Of'P, ill ~l'In C(:I!!'> (~'i), illl-11!~!~ i!ll .i;! uiche wiil:1-~ :;;p..."'(:i<llllW~ ;~~I~['i![lrtivc (~CJ;,T;]\.~s"$\!Ch 20 J.!ld 3D' Ilri'l'ITt;I,Ch~ yi~rd 'U~" ~ho!~ ':;;l"\1~9i'!'di ni;:~'!ll~l!~d eoclls IjI,1rbffl"i:njg g E~ .. ocdls ,Ul~~,~ lIi~~~~' re~LlIa!~ hrlml(ll~~! mi Hc~, flrg,;l!li ..j,,-';~ il~\IC masses ·n~.a'1'{I~1$'C1y n."I.:'~I~i"lo!l!1!l'C t~)C' ll.:(1'oUiiCl" ~J:e i~1 11) I.:IJJlute:l, and dc!~icd rr¢v,I].lcn~OFP+ cell C:lu,'I~" ~\i.;;gel'>U~w ~.ltl· the tis~uC' tr.l!h~ltlm~[;jti(;in:- l'if Ih~ gr",'>lfuLll iug end ~~ ~ili;."i!~I1S il1~t'lI[ltl:>ih!,:! th;J'l ~\!elrl n ~J!Nattlre ~,l~llild ~H'-JsN~m[l)k in"'~II·n, I kl\~ et1o\i,I~~w 1,l'irno!'d.ira:Jgerm eell rOfll1<1!km. l1u.'_' ool1~,(!I.>U ~m~ryn, ,[m~1luuhul" CI'~'illlCnli> or 1111:' ~h~C\~ mo.st ~1'lCCi8Iii~_(,')(1 uf c~IL.. 'I'M:: '!.~ggmlJ :;;~1iI11 arise hl I} ~1!ii dish'! 1\ crl1k:-a! IC\,icw ,nflhc S,pOi'llal'ie~U$ Iheramrc S~lO'\~'!1 ES oecas ~I ~P011 ~~lCc,trlk~1 I~I~' .a(;tillMloll :sl<!gCS {)f geiim IinC'<lJ,;c jimllmtolt in ~H ,C\!tmrc '~~!1do Ilfii~ 1'l~l'l~I~ I)' J.~IIO:i,;nc r~lm;;~i{IJ h;:l;['1I(1id 1<111 ,g:1I1ll:l~. ul.lkirlg c1inj,c;~ ~JlP~iC-;l1inn~in In fo.!uti I:ii}' Oocy~~,Jij~e EmtwycilniC liIH~ul~hlC' I.lI 'i.hi.~ t~Ii'k'. !,:\,i,;fihck~, dl!i:il'l'lUl.ii;e;~ N (ells stern ~lls, POljt'u~nogmelk

reuNty.ilem:c" nlo_:.;.~ Iubn~':I!toric:; WlUrS1.1 C' ;!J C!PH1l;in~d dir:oXliC'llll-:;;~lloc!iL1n arJ"lfO'lA.~h; Ihe' cells i!U'C 11rsl tlf(l(ltkxl M d~n;Ct'e~ti'mc w~lh as ~nli1C1h dil'~1;l it.m B.o; poi.,.$i.ibll:. ~h,.;r. III..: d""in;:d ..,dJ~ lIn: 'llYi1e'HrifSllreportoo 4 yeElrs Ol'go, ~<!me~Ggene5~5Jr(lm ell!lbJYolFllv( ~1em ~,ES) ceU::; ll·romiS'~d <in I:\:tnm'tXI Imni ;j, IiIl:qo:rogPlll'tnb OCUlI~L[U<i! illg ~ aCi\!lSs;! lnvit ro medel Ltl r,~dlh<!Jte mG~eclI!l<!ra rlalysis,af ~he g'eUI'lI inne<!!jle. Formatiolll O'f ble al1l:ilxidie:; ;Illail~li.~ ~ll~o.:l-cdl i>tl n;l.tlt: .~~lti!.'~'j(ll-;, i,Ollt rnimonliat ~erm ~:eb l:~'ro!;I!.J$t.II;mttermin.:!l g<llme~Clgelllesi~remainsin,'il~fident arK! ®1!lbl'S by vili1lU: of rort'l,i:l!b'l'.~ll~ i..'"....p'A'*'l!t! frOIll ~i,o;,:,U~2 abollt 'gilrrneterllmdion persist AithQl:Igl'I'l.]~fl!l rl;lr re~;ardl" dil'licO!llIlse of E5 tell,-d,eri;~ed !ipL'CiIicprnU1U~ii:R;. ,gametes tlIlJU'(lIFS a dis:~an,t prClls,peet UiTIJi.: mtrq;pClfbllcmII. lw.or::Lltlri.r$ orj~in.:JHy eth Imllllilm Inad ii1lf1lh:-i(; ,.;mbryoll.Lc; i:lI'-1l11 w, form (J(JCylC:S Ih~Jt arc mDluJiI.: (:oolJ,l!h to :\1111"'" n::PO::II.;tl(he d.i:liwnhm 0r g[1mfu~ rmll1 ES (ES I ().:iI b;, han' pi:llrip('Il£I1CY, I.he U1lnll"k· pen 1;;11{:(;'t~"'cqmlgnllmni~n!,'. "f IMI!:1]f'-~rL"f\:;rrt~ o;llls un 'Iilrn lIi>i~1S diR:("LIOll·;:;d{;(.'Liml :\lliIIl1Jgk~ r {Fig,. I) U ':l'J, Alltlu\:,.; ~~l'ipkli~l.xI Ulmrl..."I':> m ::;onl.1.ic HLlClbll;. <'Ihll: o['<J:~liK:i,t'i tll Ibml ell H~l'illH.';S ir. the ES cdl~ c1~I:l;':I"iC1nim~t::"pOnlmloou:o:ly ill vitro hody, '\\ih~n m!JlI:.c ES ~dL~ lIn;? l:ml1.'ipInJI'L'l.1d prI.\!lcri.iCi'\ thai di:>iiilJgui~h £otml, cd~:;, [[(11m undi f'li;R"lyti~llcd E.S cd l.., a dl~jkll!lo;; ~i~ Ih11t I:n 1I1Ii~..:;s.,;cliJtlllrod in k'l.l k.:1II1i!1 irlhihilnl:) la~'Lt:lr into early ,.;mbOY(l~ ES eell ~ ean 5J.Nstai]~ '~ll.: I l.lf'], ~ 1:') !okjn~ that Intliniailll.~ ES {."dl i>l:lf~ i;'J.'IllU 1:1.111:>.~h~11i.:' umln~ ~0:IIUrl~, illl, ES cdls w OL:IHiw~willatit' 1Ulrl ~'IiIY1~ irk' d~Vtli~jpl11';ll'L tlf a ""), tn .hi; 11~t C;]c.~C. S('hn~~r ~~I ~~)U~g~I~~ rnorn;;~ . ..I\II;huu.gll I.iln; lfI~m'L~lli.m ES cdh,;. Il.lli111 nmI.."VI':JI.All1cr UF :It!'ll~!!Ha],. ES ~dll. run dir. g~1lI1C[CS :Jl>Plfl ,~jrl1llm >l:mll,pl:c:\ daM'Ogmphy oJ Iim.'IJI.i[nl;.' en lhc surluce or 1111;)r~1ri ,cliJ:Jb or CI'IIg.inccr.:xl ~ Iltlll'Clltcd 1~l'otl!llf)1l!r tb r Ott4. n

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'ta:-,m~tf ISeNl), a HI~.imd lim! ean '1J,c IIISt;d 10 f d(;civc [Seell':;; rram :mim:;J~1i<> wilh ~":fX'c'i..Ik ,I;';;F.~.lcrem~!IOCS91: ~i'orn :tf1ccilioc hlJ!l!l;liUl nmicll~"i, 51[' NT-IZ-o:;;. eil .. .;'4 ~uldhe 1il~\.;'Il t,,1 .snl~Y d i.~:J~ c
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fi'!Jl... ili~'}~ kH!i!C;~tei:'l.':jeclk'l'rl-r1flou hb::-1lq~ [oj' iy ~h~111r~~ hl~"'~i:"'~,~ liIl:U!lr' blfLl~eT~-ek IiI( Imn\!Ul . nrrrliilnimls is, a ~L!rr..-c' or ~YICf;., If ES !CC1L~
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,Di\ohi~~ liil PWialri~ 1{e!Ti!it~~lO~~01i. CIh!ld're~'~ H!.'l~t1aD BI:i~1Dm; D!!p;I FII'fIl!Iilt af lliol!lg1cal (l1!mi~l.r~ arid IiIIlI~ulllf P~llrmill~DlI!i'!D'; H~f\iatd mi!d~[~l $c~ooI,; i)lio!S51~m 101 Hf!mail~m~, 'Eh'lQl'tam ~rlill ~Y\:imI!'JI'~ H B!pjll~L ®mUm-iii, 1M; H~iV.:lI~d St~m C!2~1 1n~lHlu~, ~jfj I!lfidg;~. I'I\A, U,~A. E.·maiL:!!JI!!oFoge"~l4!.f@Cilni.ld.1!Il~ ..ml!lar[L~.

IFwg, 1. IIiii \!11!r,Oi '£iailile~o!lieilles.is;rf((:m ItS cell~, (loP) A :2:0- i:lli~llili'e yieMs ooc.yte-l1rt:e clfH;; <ii'H:I j)~nt!~'lIlog,@.rletic bl~sro(:.'stli(2) .~Midldl~) A !IiIi3I~.germ (eil nk h:8 ~'lillhin !EB:s: .Sl!PlllJfI'.ifQrmi!lliion (If plim'ilt~ll' ..perm (], ii, 13), O(l!.yl~'iil.~i!'nlOn, ~ll1tl'l spErm p,remr50rs 11,8;.l:ieen, i\eflOrted to !J,r;merate QHspr~l1g (11!'OU;onnl) 'ri!IilQrdia~ ge-rm~el~ (lIn bliii~:la~ed P fmllll 1EE!~ lrL<I(ILllllllred~,~il a retillQric: aCild<lllid ~ytckine~ 'l[) gen'~fat~ embryo~k germ (.eLl~ (1).

un

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SCI ENCiE

VOL 31i6

2D APRil 2007

4009

Germ Cells
.i!M~g.i!li'S tin>' Q~II"'~ t'lllILLII'L"'(l Ihlrm for [I Iilrt.h~]:' sc ~I~mlweeks, ,mlil ohso'\'Qd n I"C'mru:kablc~t:ma1S rOJ1l1l;UtQl.l: ComG'llC'» Dln~l!i'CclM;u' :;;,trU~U"I~$

or

dev~lOj1<Ctlih~,~ ~.'>()l~lhb:1 fI\'=1~J.n ronk~~,,md !"mm tLu.\"oC''C11~1,.~>JI nrli;;rlNil.;e c.cU .. )iUlrnurtded

h}' (l :#l!~I~. ~lhi\::ida IJk,r ~Q~.li!'li;l.E:l;t.c!!.~h e l!1olccubr. inU]l~lllll !i~ILW:h. ~]iCilI.. UID!a\U'UC~ ..~ :ll!'ld tw:al dn:!:a i!lldi~!cd ll'!>u tl'tCsc !:."C'.IJ$ \,.:~~'C'indC(ld ofK;y[~S_ EVCfL mOTe ~1!I~ni'i.ing., cul!llrcs; COIl,. ~I~incd~ys_~ic :'iU.\IC'tllp1C;,llllll~t rl,.~mhJoo hbi'iIOC),,'\!5 'liml thlJ .~,ul1WI}; ,l-p!.:\.·UI'ILlCcii 1Jrt.-.::;,.; l'i;N~lspmmm,_" : ous pmih~llo.b"':lll,.':lic ~Wlir~ ntion (If ihc Otk.'Y~~.,>, 1\.,~I.'lhJlJd;;; (iJ·rotIt"yl~ lornI1Iait',ll :fifltll, ['S, 0i:1.b Im~ e Iml, 'j 1o'1l 1~~11 tlf.iddy ~.xpJ(JIi [,,:d by ah~ t"lUnul'mlliLy, lnu T:ItJU UL<;{!Il'S !¥illip has indqn~DU:lc-n~ l;y IfCPOI1.\.I!J d.il~i.;!CrI[iali!oJll1f ES ~"t!~~!> ~.'iIO liOc'yL(lo~~kc {:o.;lb u:>nn!!t ll~iollbr cd -cunrlilumlCtI l:n.txlilIJll. 'VIhidl [lK':\lll1.l:ID'jy llmun ins Itu;mn; uiml :-'1imul;.!lc bmh >egg and :;;pt:nn d!(;'ifr.;I!u"m~ul.1 Ul), ,1\ III Kmt:J1 'tilllilalizil:l$. ~litc:>c""':mhs Icll~'C one \roJldoon~ W~H:dK·rsuch ES~krl"cd oocyk'S ,3I'l.~ flllllcti.uI1!~, Cau Ih~Y' be r~rlili~",'.d·,' elm rhcy' :;!.l'I'W :I::.

Sl)crrllmidl!;~ \Vhen irljeclcd im.o tlOCytl1.:!1, II~re 1Xl1l~ ~C'I'cr:mtoo dipfoid bl<!!\'!O('y:;.L~. ~W! never !lUpPOU1~ nlu, l!ro~I:'iC'~1i\::vel(npl'll.CIlI:, TI!~~ ~:-I~ly n:f'!Ort~ ~.II~iloC!,Jn~('!m I\1!b~I:;;!for" O I ~~tiou (;It' Iwhl~Of(tbl ,I;,'Cnn cells, bi;.l~ lil'lli~.~i(l!!:!'i, ~n ~C'hic, iu,l,l nU1c!iol1,1! iil'UllC'lC1.. U::-.b~j;ia !,l,C'.nu
''''.lXldfk; ~td~", ool1l'!1e.r. SUr;Lllr:ii bb 1m... in!JJ1?jJ~Dld~m!y!i:ll!lrifll~~g lo.l·liIla~io:nOif priHwroi<!l1 ~Cfln cells frnrn E.'i cC''II~ iln viMn '?}_~~] lmll1:~1'1

Ol

injc~1l'{lilllo (locyt~.:;;, ~~: anim ~Is \VC'[~ boml1l.8Jld Jb""~Ii!rhol'ccllihc lmJ'\1~ by Sou1hem lllnl all(l.lV<"i~- AI! hD-u,gh ~1~."I!! OUt(!. are I'lmvp".,uivc. g.;=Mrn~\\idA;! fli,1lyrno.JTlhisrH ;l!ln~y!'.is \"'fI~ll~
~!!!\'~1~1l, h~l,pli!J ll'il'W!ml\!~ !~~ d1~>A!' a~ Qflspr.ifijg

eell

carry ,~ fli! II ~UI~If.iid"QI'lrlhL~lij~J'tl limin the ES.... derived !llWnR

G~v~ Ihe .~n~flilCknc)'

ES O!.lllllll'C'&.~ftll ce!1 1'Pocii'ic gene c''''l'rejlS~lhl hm, been tllx;~Tn::tlinan IIPJilWl'lprjO'!l\: 1r;;l'npoml :>,"'qur.:riL.1~ tlf)') 1lI1JiJ iiJHl\.\'iTI to be "'iTllittlllh~d by
illlClliMli:OH \1I.ilh imrl~ rllm:rh"J~Iit::11!;

pmll..~iil:O:

Ill). Ntm,.:nd'l..~Hs.gartlr:l;'';l, ~~l\ol.' IWI y~1 bt.:\!,11 L~[lI~lOOfrom hmn:m c~.dIHIt~, Prill'Jo:riJiml g<Jml t\:~n:.l~lCk ~Xll<lil d~1].oJ:fhiiOtII. S~1l~.uion ,of ~ntl tcli Iill!.:;. illl'O ooJ!!OI:liu or
::;p;;nllllL,\S(lfl~a[)(~rur:; i:iUr,;;:i. ~li;

w]lhiu

IiItc

(kwdr)pi~\~ti:>-

lh~ !1(;t~J Ul:liUcl, uuul,crmi:n.al g~mL,'[(Igv.'l· g

hOTlllmm:i t,)[Mlllirot The II) ES ~LlIIIIIIT' {,'Olldi~.iQn.:n11PIQyt-.d by Sclu:i.lt:r":;; ,grollp :i~'~~~lb [[!! slil11l1lalr.: ~I!',i(Dlt!li'ii, whrJJ~t~ 1J1Rl3D

~i-;. IeqLIi=ligflt

t'i~lpicms ill Hudcar ~r;msl~r ItI

cc-Im reprowm'luning"
1\\10

~1p}1Of! ~ln'tni\;

other L1h(W1llorje!l1 reported dCI'h'i !\g male g;aU1C'Le5t!1lm F.s 'Cells d,il1'c~C'u~i;mxl hue 3~) E!h i[ I. J). N ece <If!d~~k;tgtaoC"':C1'\p!~illed ;a re~r!<!r f~, the germ ~d I ~~I!C l~MW ,~nd :-.!imllilmed ~~rbl~ eell r"'If'1ila1~OIlWi~h brJil:1;iC
cmh!'}'fI~!ic g{]I'~K~,and 1~llspl;:um~rl testes

r~~t;l!'lJli~e ~1~r~I~~OOIl rHhibi!g!\g~h$!£1r:t.W I M IS i.~mk",'\l.w~ !._~..r..'CI~'<.I is cXllre~oll M in our I!Eh;.lJ'!Crh'~jb s~a;~!;'ll ~ijt!! r(iftl~ilili(m fif a btr; ~Urii'W.~]~~egerm cell I1kl'!\: lOr nl:Jlc l,!,mn~~lg.t;11C'Inl11f111.~C'i!\ctic '1'I1,lUidn-4IUMP4j. Thc1"isoImedil'(~.~"f~·-[in" cells, mixed ~Il .i~s~c J.i'01l~ sis; I/J\
::;r'ti".~~riC

Eli, cilndijlijoll u.. hy J~e '!1grfHlpan!d fllU' OWt:l sed IlflllltXLIS 1o r,1vflf~pcmmt{l~cm~is. fnr ul!~k'.1r «"<IS(ltl s, CiCJ11l cell mal~~mtkm alW~<IfS 'Ol~lll(l'!.'" oog~!cllb by dcl:ul~~. ~ll1cs.~~ilpre~~ ~ by 111l~1c-

r.~n(,':!;i!lJI'Orn P:'i cdl\; i~1\ hm, and the r:UJc:ily of dJm doclIU1Cn~i'f)g repmductij,'C' jlu~c~inn. ern[lwyil'log ES ~"t::I~$ ~ H.l4J ro;.; or g.1Im,.;lG> !a 1ftll:1~, 11:; in!i.'ni ~aty l!~m:>n d isi;tl:l~ lJr,t):;pc(.1;. r~lr l.i.l[JI:i,\; \ii'ho ~~ I]i.\: m~l) ;r.~h:1l]f li.tll,;mr ~urilct~ri{:;;;i~ i~,:.. 111.'U"!'d. [lifil~nc!l~ia~itm ES cdl:. imo primerd~l] g~iI1ti eells :Il1lp::J11i ruibu.s!. mul @intll.mllhk to Cllpc.ri IIIlCUlUlliIJDI. ES l"C~~::' thus pauvid..: :JU:1C· ("C...... ilb~cmol 1'0 :;Illtly ~~m.~lImt :.-pccjty ~~llIn o;llls. Ih~p!lllhwnys ainlU t'OUltt.'IiIgcnn ~l(,"11 igm. m aiul:I. ami ~h!: IlIItlklC [JII!ur Ill<lchilli,;,I.)' of umprim Cfnsllm:' ilhm acb when gcnn cdl~ arrive at Ihc cmbnyornit' !-'mmt:hd cid.~!::. Fm I~m~w'hi[) \'ieW 'dil\l l:uj1 as h;lIf Itll1p\y. Iltlll:ia wark I'l"ltmilJ~.

or Wflinil1lll

tP~llet[li'"

or

1~'iJcnc~ or ..

fl1C'ijo~~,<!Iflt.i 1.\,!,r!'lliJ~! gll!!1:C~ogc!I~!is

10

h ntlcl1 i Ilg_ ~l~11!I~il1!'i

he !\lcccnwhclhcrtcm'lliln~

I,

or

.'f..:m~ii(l~115is.can QC~"I1.1f ill a ,""'tri diiii1~~mI whcd1c.r OOFlle;ts C-;U~ 'llICOrHimimd(o 1~'J;.l\'klc3 wdy ~I.IJll~~Yor {lQ!I;y~~,'i: repwgTILl1Ilnli:n~ for ;...~tdi'e.,,;. 11' ~ldjlil7n;~ M)Om1Drics 1\.i[)lk'liC' !he re~m~~iGlbl<! .:!chh;,\~mcm ~lflh'~I'!ig ~~nspri~!l.lrol'l'l E."'-d\!n,<,cJ gi;lI~!CS. Ihcn 11il~~Y 1\C'\~r avenues lor genu cell re.'1.cJreh mlC f!OSS ["~i;,Pn:!SIl<!.:!!!.'ts

.I!lcm'icell dcpklcd un~(X,lllC')' I~:mml n'll!lrker[los.i~ivc: eel b in rcro!lS~h~llcd ~ct.niillli;:q'(luS

~111CgQh;b:~:!!ldi.!nJ [or gf!l11CtC !la I'IC!i(mi ~ !il1J(,' fI)rt"'''a~i 1l,Q:l~lrC'!iml)' 1'I!,,'(0111';:.t3. 1l~O,reR1'l!lllsltlk rrQt.1l.iC'Lion (If(lll5.1~I~Pl,!;. f\.U of Ihc ca.rI.Ys,MliC'll, IUOSPCCI• anti tile C'!.IJ'Iw~~~ . ~Ildeed ~cem 1l'll"la:C klt~cd IIlL" enlci:!1 r.whico\C'l'l'lC'iTII." ~nEngel (lJ;li;! Lhml 11 J,I r rliit h
Rfi~'~ll!\l1~1!'5.. mil

'~uhtl,k:;, ;Ihhou!j,hrm ~l1Ili1 I.imct~ml 'I\:.m..~ dkJCiU:' 00 111\:3.~iC!\ :rooC'JlIL~y I'L:fl'0rt~d :;mx::~<" U... illg ,::I, GFP l1l~r [or SimS ~::;IDl'm!ilal~'.d ytetimic b rn~l:Il.txl (.H. Folll:w'-lI!"i Niudi{-;:" ,1itl!L11. nhl: ~!Lrt:lt:' ,. .!ll'tmlll fNiI,ru lhm.111.l'i:li:ll .. ~I!lI~a:;~ t~tjni,;~_nlllr:niL"in :L't'.id..:.."i. :~ ~'lloC b'-"!JlTi.~~'tI ~n r:lt!,.'iIT!~iDlicg.~nl1 inlhrcm::,,-;o. ~~ml ~:II IbD'mm~rnl. :o;u~,l,,1ill.g dUll oclL>. Lhc EngJ:ll !/l.'UUIJ t1i rIIu~!:Il'_ua1£'cl lile.l pllllli. POkl1ll P) IcmtOCl.lllt:.inollw ~UllC j:[~f:!l11.omh,~ arid '1.\ m!;: runailll~ 1.0 Upl.iD'l1~~ d~ Ili~\;1.l1i<Ititm I AI~ho!ll.l!h il]ili'~i.lII:y~'('king ooc:yl.l' d{·\!,~!.tlPIJt{;IlI, id~nliil.ioo GFP+ cdl\:;., 1h:ll. '7>'IU.~llITttm;pl~alk:d inl.n g,lunn tc~~ '~k'p~',:lt;dIt';Sl'IN, prodlilGoo l,tl:JiLU[C Ollf Iilbol1llol'Y illdcJlc]1(k~IJ!.ly ob:sC'.r'I'oo nmlc :;p~·l1l1. Ahhnu:.gh H1CiifJ'l.~1lI1li nnmli fl'l!l, :;,lmclu I~II g.ml"-'togt...'11.i.il>[~ frml E.'; (ll!lb (1}. "V(.~ ISt'kClod cdb cxp~_ .... ~,iJl.g; Ihc gr.;'l'DlfI d~ amigcn SSE.A-I ~ lI~nOTl1mlitk.s. and :tl">dllCcdU1ln11~il.y, lLX'} :l1!Ipported !..lnbuyo l'bnl:mtiOil ;llk]' beirlg itrlJcr:k~ and lKalcri IJOI!:m wi •.i.1 rrtil'lOIC ncid. whkh in· dll.t"!!.~ WRn C't..~I~ 'Ilrn1il[i:JTIILOI!_ TIlL'll, lIl"illl~ iXl!ntlilinlo !lIil""yl~"S U.::' I" A:g'Uin. l.h,"~ ~d: of Ih -.' orr· t iOllH l;';S:Iablish;...\9 I.br iM.Dlu.il1g C!I'I Ib~1~m~ic gt'.rn! :;prtng l.;u~~ts, dmt ruuctimml !i!,llrn~'IQ;W('_R!liJt l ,t EG l cd~:.; ]rnm mnhl'Ya,~ (,91..we c~lllm",d .00· rnrrmliJ_ E",t'H mmt' .mrl'lil!l'k~L~. ~!OW~VL'f" j~" .bair like cd'is ihallm!.~ CT<!sedthdr m,c~hylalioil marks ooccuil claim 10 h~w'':: ~~~r.u.JCd li\'<! alke rrotl1 E:s-oerh N !I~mc~~"S. ,ill ihis, case" dlC g1'(JLm m. key impri med !;'i!ilC' loci, Il dk!.gnns!ic (~'3M'C' or IlI'im(lrd~,;:iI ,!,;~nn '\,',,::JI~ i'ro()m ~hc ..~'I'Ih.l-y(lnk ;:1~ucl'lti~lIy~~~"!C~cJ :SlrnHir-~ cds, Ihjlfr\"'~xI b>,"dh C.\lm..~~il1g ,~ J:;..!k~Jn;;1l>Ort;.;:f dri\'~iI 'lJ;y .l;!oua~lrll r:id£io:. Fro~n EEh ~1:.I1,ill..ll\:t1for all

Maim,

n,

:11. ~. (jgj~1!n ~' lit, MilUM! 4Zl", :lIlS (ZDa~l. 2. K. ~b!l!er U 1'l1"S'lrnc~ 3~Q, :1.251 (~O!J3}' 3~ ~. T~l!Ok!l,. N, T'5!In~~ R. ~w, V. N\CI:<Il'. 1'1«. .4CQ;:I, Sa. ~,S.1l j[(l0. f 4. To P.~~,l. A. Th!im1Of\ DM:J~t ~~S}. 'S.V, r!I(i!'(illOff N lit. ~~MW. Cil.rN.tmt' ~:U',300 {20Q]}. ill, O. 18:1'!~!'!Ho:llpl~m,'Ii. Chr" fj.,froun!;M!J, .h!r.! "tiM ~4.

n'l<~ czoon

"~tr,

:ua; :m

Mo_t Reprod. !!ev. 7~, 4137 ~OO6L a_y- M.a~ui., ~. ~~1lI,. B. L 1fiM.!!!IIl> Cell .\0". ~&1 (1~2l ,_ m.P~et' e~ ",1., ge;'Jg~i:;"'~> '15 (.l!O~)" 1.0_ .k T_ (!;ir~"r 01., iII!Jm. l>l.!;Jl_ (;Ieme;t. :l3, l27 i~~~~u- ~ t:.ee, l- r:)Qn~l~~ A, T. 10a11~, It J!eijo I!e~a,~em Cetl'~
;,

26~ ~:.lOa6),. ~. yjll)m~ ~I ,f!'l.,

"'". 11:5, ~2D06I, [2. K. M~~~I"ia er 0'1;" 1Ilm:i', A'l11l. Gi!rjtL ~3.,14s..11 (lOlJt), :lot It M~~!fil1Hl m;, i!l!il:, f.allll 12] (200M_ U. CiJ;), f! 00 Ih~ ~ti~tiffic admory looBl'dll1 :1M n.D:I.d!i

au

.f.!ddi~io:11<110 10 .", ~'Ci.:k).\, WI: ilI~d an utihO!J~ 2 dhil.:'Ct~'J 1lW;1~l~, T"r1 ~nliHvr: ~(X'rn1.1W,,")'I;.'l'lei ~~{IIMe fjll\!~l~YI;nI(I~d

i,he ~~ljmm~ I"t'YI'l,m.. i, ~'li~ r"; " id~i~li relb,.1i ~ ~1lklg

11!'L'lC, )1~~_g..: bU'l'Icnvli~,go!ncsli~ (IJ~ 01'('15 Ctl1hl~lfl:!1

!!IlJllIi." illl Vi;J{![I, r.t1\ii:h f~ru5l!~ ~f1 till! IR ~f :11um:in (~I! toll, Iml dllea~~, ~lId MI'A\ c~Jliltat aJ'l'e1fIlw~e'[iIIlilili, 11111t1 fflllHbln!l in ~~1Wr!"loro' ,~r!dJIlf!Ji~l,tf<!l!fll!ll!lg~,

~Hs

1I1,H rcNI,.~'I.'Ib~~dr\lllml

f(mnoo 'loIh!l!rlIh~ rMl.:lltli~IC.d\;;~ctI~

~l::~~Sw,",r~'

410

TSlOp,sralufa S,8X, R,sver's,sl Illm;lpllii,es Sex Gene' Do':sage in a R,e!ptiile

ALexa lfull:er E. Quil1In.~·' Anh:uw6eorges,~' S,u~phe.rl ID.Si3I'lfr,eo,,1 fimefil~!l6 Ill.arimo,! T,i:1II1q I[",a,l,," ~leI1iUlaer" A. 1'I.iI:a1!'5hClIIQ '6I!'a'!;1~52

]fl 1~tl~ (~7",f,~ Jri.ciilm.lcti ut ::!:.:~C Me.! 00.1:1Olrd~m. ~ plli;,nm)1)! um..l ~'CIl.o:typc:. IhJIiI,';:\~I;,'I'. oi111y I;:': ~~fj anil1iltlil;; (j 1%~ from llllC h i,gh· tel Upt.:r,lLml,,"ln;ilill ICnU \,'Ioop; .;om:Gruam. All ,ai~· cordni ~t atlimuL .. 'W>L:1re ~m::ll.~ pi!:: n\~~{,N (ZZ) Ihllt dL", dDpctl m\. [!'iit'i1J!1lyp.i.<: ~i!'n1ilIks.OUlf iilitll tr~'i'I,1· OI.'l);limlL:lIhat. high iuc:ubDlio1] ~t'lnJX'!;IIUR'~ 1iU\ (l~1 sex I(Il;~bl~' SlJ. This; 1'i!ldi!l.!l el(tt'nm:;; ;1 pn..'Vi~xl:;;
rtll<UL'II Inw'K'1Illpar.1IW'C' NC'JJ:II"CVlThlJllin a skillk i(l]J b:o' cxp:Jicitly idml!.i.r~·rng dbc:oruJml.cc 11liJl\v~1 gcIlQ'Yl~ "lind ph<CI1O!.Ypc and ,tdl~ cmpiftc~~~ !'iuPAmn[o:rthcc~xl~~w.c '~11"lSD an~1:lSD, (

d~t!il,Ulli IICld by tC'!l1W'C F,lU,IIn;l d L1<i~]g ,i;IUbl)'oU lc

~ S

or

Is cll2lJ;lnl]in~~

by

.!_l;~~

0111 sex ~lllTIm~

:u::rru.!I.

:.onl{,";r;.lIUII,"IIIY \'(4~bmtc~

Igc1l101·>'lI'ic!le'''

c'vr.!,lu 1~~n'i1:II)' ~rnlh'\ii!OIl!'. i

d~m~iI1;lIOCm

1(1:8011 Inn I11"Y abm b(;'

1•

~lJgge~l:s. several i~~d~lW'n~~nt i.n ::"~X·d1;'~cr.rni!~ ung I'IlC~hnlli,'U.n_~ 3t blil~Ir;iru;.i~h'rilll fonw. have l2. y~ t{l bL" d()'I11.{1!!'~IlOl(cd.W~'~ho\V '.hm hig~1

~pl.n~

·r~1.eW

cbmn1l:1!SI[lJI1c

i!';,lllUr;" tW~lk~:I!)'

rl:ll"

de\,dop'!ll~nl ~!Clnl'~ll!.lre-~pC11dmn !,e", d~· tcnni I:mliolD OS I)) 1 II )'. Iu repl! les, s,c:lt. (Ile. tcrmijn(!!ion can ilH 0.1\ cOSD wj~h Xx'l'!nd X'm' sex Chl\llllC)<'iO!ll~ (JlIlIlc he~C~,)gaIJ~~lY, "'~ in tIll1D"I1mab;), G8D \7>,j"lh ;Iiltl ZW .:>!.::'I! ~hRilZZ tiiJ'Iit~1ie~ I( :fi:itmJc .l'iJO:[eri1gmiiJi:IY. ;i~ in ~i1.'&::"" er
TIll) ~J .1), Uc d:il>~ri'butitln

er rsn
'DPmiMlill

~lIld GSD

inc!lbm ionlCtnpt1P1:tur(ll1 I'C V0J'!iiC' .!J('!lol.ypk ,(ZZ~ ~[l, ,lhenot}'pk' f~I1Ui~l'S in the A U!ltroJi U cent r.! ~ lllC;[lxd4'd d.roji!~lU li:,;:,n:d (PtJgml(l t'INh'W.~). whkh, ~!k~' hlrd:;., ~Ia~ US:~)wui!] :r;.:mnk h..:tcillgmndy ~4l 1!~m~JoI:f'd' tu Ii: th!ll~ tJvcaii1I.1 ...:. £,CIl\:(!> I inl\ ok!.1d iti 111mI!: IIi ni.:renti!~!i(1r11. \MI.: iill!.:ubalOO t:g~ t'iIL Po lil1rnp.v ll~
maks
(!(mSIlm~

,1ilT~cmi:lliLjll .. whil-h :-;1!I~CiJ,~ Illal Ihe ~11f1.1i.'ClJll]r m~e:b;P1I 1.'<111 dlin:c l~I!!l ~~~ !k1L,.'I'u1.bm· tkDU is the dL'Is.;igl' of;p Jj;J.lIiIC on dIe Z C.hQl)!lmolll'll I:.Ilht'f lima1 1]1:t;~ '1l1\~~~r~ of;1 rCl1ltl~c-dcLCIJ11ini:n.'il 1'(:IiC on Ihl:\<\: Thm I~. UrI:[llc JI ni:r;;!lltl a~IDJ'! Iilj,-';Iui~ 1111'" ~(1<pic:;. ~J( ,I Z rnJIllJ,; l1,":UIf:, Ihe !.:<\pt'C:.<l>itmor uct~!"ily {' f w!lidi i."i ~llllci"'rli rOil Il'Il!L1~ dCvclOj:'iIl'li.~iilt'illy III ('I;fliLilll~1 b.:rrlJ:'h:~I,r!if1!.'::> (Fig. I~, W'~ hml<!.: ~~C1tli'll'~lT'dl~'tI~..:X re\,'!.~1<:I'1~It
7

1~:lk'

$WJ!OI"f;fMAi.'

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:,

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!Enilllil19

tCfilIl1ll:m~wc.~ bo.:b' J;!aj 20'" ::1ud

JT''(" '5). No c:rubryo::; ~lIT\1h,~tl[{lilm[ch·

Jll~ m lW{'" lih,1.W~t 22'"

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SCIENCiE

VOL 31i6

,20 APRil 2007

4111

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This AmoUlR'1 1011 R,ecycled Crust in Sourees of Ma,.nlle,.. erived Mellts D

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Knri1(l. Ar~!". ;md NOl'iuh AII(I~ltioc Pm\'i [)J;t': k~m"-t~i!c.,md '~~l"llci'IIC{~ j]kri~~" 115 ~"-mrkH} fro'm !h~ Arc'h~~, Abi~ihil ~r!i.'JC!l$~nllC bel! ill {'au;J{~n ;Ir!cJ the Uc;li!!g\\,'C'~11 in 7.imrdElw.i; ,lll1..! Snutll JHrica; I~cml)l!(lk ScI{!nol!S, I'!ill!"ifOl!lam ~lItl.nii1JI Ulftl~BI:ty, C<l.fI~liI'a, .;!Ill ~1nl<1!~i iti,ch~)1<1lts Ol():m<Hhlfl()UII' Mafld. Cllli<lda~ 10,200 Aml~~ll1a. ~L.. lior.atoiru de (j:.iQd~MmlqlJ~ di!~ U 51J(1i1~i'ne~ .Jtlplri~~< IJlf1lrM.~I~ d~ Glen~bl~ .1S4Ckl ,Grenilb~~ f UUI" II1c~lmd8is i~~ h l),)~i~ I) tlu! C,l"'I1'Crimem~~1 w·ud k.ulilill iilG> uml pj(;Jil~";;mm GO~'tlJln bl[rn(II. i 1Oi!di:!:t IFri!lmw. 'llf'parlITl!lllit 0'1 ~~J!I:!i r4<1~!!iM1. famili1 <lm;!l1h~'ret~~1 p!'!l:dft! to!'! 111>J:t ~gh1:tj'e~1!! ,:: {I> .... Cok!l1.1hia. Mo;:o,i :>t1mWlk,,>:.'In,; l~if;,d,~ or olj"iuc h n l.IilM>FIliy, r?osl. Clm:!! h n,~ Talpi!J l~, T~IwOlifll, 14. of 3Jl' {lP;!i ~~~dt~l!g (lrl)'1lkal fOCyckd (H."e<'!nic oo~dl~ oonn;liUlilllS IH~ :1In~L;'Iwll~, rn.~h. h1gh't ruJSI.-~l1afl fi:>J!!li!.m:hlowncik Cea'itiro'01 ~a!Il~IM:e f1I OM' ni'lI;;lJb'l>illUin ....iIlC"rll~i1fJ>f,;I)'~IJi,. Th~ ~mrb I'ITI.: tlli crui>1 ~~1l1h.: ll.:inlll t~r'b'!,i'lo~LI~ ilh:J sC<jJ.imllCSLO~ ~ Dif~ .mll :s(itIJ~1. Qj !Ef.!lIlh S(re~~ Ulniv~Hil)' of T,ik1~imja. f!ri\\iltll' Elii!! 79. Habal1. J.a~rniuJi<l; 7001, ,1'(m' l)h~IM,:) sntl 1t:'~.t1.ion ~rthis undl wilh pc:ridolil,!'! l>lIIbdiviu(.'dimo tOur gITlLIJlii.; lil' MORB; nil ilrOlli~,![)~Jl<Irllnm! 001Eilllb '}(ien~e5\, Quttn'~ itJ)ild, Wilt plll!LllIt'!.:S ol.iv il~,,;.Ih:;cP}'It1X.tHil.~· !j)_ Will )i1,IOW "'I'ithin pll:11l.; 11ll.!!!,I11Il1;O> (\I\.{PNI, m~I,gITWS cnlpl,-'tII M~fOl(!fi;iil B'IIi~d[m!l, l!~1lM~ty "f ir~to~ 1~1i~1DIaSI!IllJ!'J', f,n from plHlt bOlllllclilrL(5) forming 011:1 UK, ~~~rIJ1llffit of E.a:rI~" AlIDO.~~MC, ~lIijJ ,1l1.anf.'li~1)' ~hm rim.n~r ntdlir~ 0ofthi:; hybri.t~ IlilhoioID' in U!l: Sd~n~~, 1!,IiOI,~~(h!s;e!U 1UJ5!i'llIt~Qf u~dlllo!~!I' !M.ITI,n ,lbMir.ltc of I\':siu'IJ.iJi oliVIJIC il", 1l10li\: ,"OllHll.iIlDIlI::.' !.illlphl{;(;d {WC. 11"]11 ~i"'liKt;.plJ,.;.rct.::I'O iml Lhid;,), WI~M-THi N; lLi.i} \.VPM: {O[B iliad U PI th.m Ih~ mdtull!J, o,rpcrh.i~lliLc ~~n IlgiV,""l pll.'l'J;ur!.: ~~~1iii!T!~~~llj l\"'~~I!II" Q:IIl1!;J,ri!i!ge,. I,1IA (DU;!'9" USA. 1!Il!l!!,p,;!r~!TIeml Ge!:!!gg:r :lil!d G~I;l~h:rsk'S-, Ulnr..enhy of af mull kl.1lpcmiun::' IIlld l.i.tI:Il ·P!,TOi'il::illilo-d.:;r.i:v~xI clllfllm.::ed ()...cr~h i.clk J'~llm"'Phc.rc i;;- "I). kll1J 1.liIid;). H~~II, 'lJiGSQ ~,wJ;'~t F!l~, ~f!g\J!J!~. Ii!~9~;O:~~I,II)A rnt:~Ii> ,l]1iC ~h;Ul"'.L'i,;Il"ri. ~it~iU~i b'jllricll~cl in Sci ;mrl .. WP M- TWCK; mul j ivl' :kt'!I1l:lI.iih'S. [[nd aSM!u5Ch~ gI Eianh, (I(iJ!Ilt~ .. ~d lPWiI].I!IOlJ)' S(ie'n~~ Qlidkff N ulDlilt (kpklAld in Mg, ('11~ mltl Mn CltIJ1lIJo:L!lt'cl ciia~\JI m~W11:!~, .l(OMATJITES. [klau~:; or.:.:IItIl.IJ!i~~$11.)', !II~rjl l;Iyllcll~. ,r~r~ ~CJ;', c~lyjiU cFJJO lVE:. lit: .. 12f>. P. Stllrdlmf Ili:'I~l~ut~ 01 OuaMlog~ 01 AJjJi!5I~m w illh 1.I.K!Urpmid milC-d"''Ill\ cd. COUllLi,!llXu'tJ;" Th iiS pic Ii:M::miom" l.uli::rum."Ct> f[1r ,"!lfnpk d!:~i.::~lp:Lwm;" ACi!ldll\m'l' -of $tl!l!~!~ N1a}:hi~ il:ro~PI!kl.3:~, lHt97 Jmd~lleir gJ'rlU!,!Co.1TeSJ,'l0lU"il:'ncc ,l'l1lC' I'lI\.'Seulcd in lIi IriJ.rrt.'.m:t: <lri ses ooru Lk~t~01i~ im; pr.um:ip'.:JHy ,MD~~'II!, flilil!sl:iL llDr!~rlm!!m ..,r :fI!!Cl'$~. I\;JruLty IJf L:~hlcS2a, rDn1TOl~ Ih", U\Jmpc!l.\ilLUll of mdl pmorlucc,'!d ill 'G.I!~Ck!I'I:Ci!'1 IJIr~c~l. IJrnw.wsllil. IEiud~J)I!stlain 4., U1.ifflilri, Tille nlQ;.'iI;-m~llc;\i<:I~1 olivine c{lfn['lOSh 00:1'1:;; IlC1iclo1ilC. wl,!Crt"as~'l-'rmcl\clll;~illly C\'nlR1.ls the Ni:!tl!ifll!:ln~s.I~~l1Iom~Ml 0,1' :PelJol"lJl'" ~c~ll\l' (II hn~ {d~HlliNl~~j'~~Ih rl1e;~ iPh-cfil1~'Y"'1lS. \vith j'!'o ~'!. hhln . ({m~fK1-silr.(l:l) of m'i:ll, ~mln o~i"i!lC-lroc hyhrid oIllld l11e Sdernc!!t \!'r6J~ iJ'mw.Jlc;r:o;llelt [)e- 6o~ladn loas, lQ81 HllI .fIIl'I~t~rdall!" ~!1IJ1;(lrLand~~ ~Depail'lll!leni of BaI1il1 I)yro:~cnitl! ti Jill, Exrcrill~~t'iI:~ d~!~p!\''''<Iil;I.~9) ~ 11'l01 % Ir'i1!ln, <l 1l'l":!oo.im1.tfn Fo) for ~W':~I5JJoCC'lmcn oiI!ld ElIwicomrnenhll Sl:itnOO5" 'lInLW.~;Ly Mu!J;i(il1, a~rB) th:lt. :til ~tldl i~)·t'l;'Jx,~~ti,!c--d(,fj!" U'l'o:!~I:;.I'i~e ~~I luwaN ~\ !o!~' ,~\ C!<~~~d(I.[!hk: $2J~ fQir ill!! H'llab~IIKl\\ n in MI:!mtll., GI1rimfl'll, ~~li1i ~a!bnd MatIJU'~' 'Q1IW~, £lriilldlrE1Dur l thJ:~lI!rtace. Ute dC'C~~e ~n r~~lI!re' e;l'LI~ Ilil'Cir Fi~ I, hH.!i\'idl~li t~lh ine~d~;~s are I'lre.. .Cu~cJ '~Q, j/.l'S'tmiIIIJnillf)'j)Y, IS 900. 1rf'ldOO\. 1'10epilrlJTliffil 01 EiirtJ1 ::;mumdLJDl In {I~i\,im~. This (~I~dnC'I's UtmMIilUy Ni ~tl !ig, S4 and 1.1hle$ S:!. !,;' W :f. ~00n~, liIni~ 01' A5maI'iJ,~mar.J, &~r.B;l. If! ~d(Jj~ iOl~ to Mtl J)nri N i oofl(;'Cmf(1tto!-!S, rich ;)~t! M fl ~m! n 1'Io[~or. Wi~~'1 or (II ~'\ii,':\\', 1!~ '"1'0 w'1,c!1J (J!li!~e5fDrude-moe 5:I~L!ld hoe add!~~~d. (-m~!~ :llJ'l1de<'@geQl~i.,ru whk:h ;::;Iroi',g~)' Cl.ln'el;)lC wilh Fo tPi~. I. A ,,!f1d IIu;rgc drn.~ :let (IIi hig~11~n."'Ci:liotl ;iI1Oi!I~~C'S OIf

illl!l- SilTllib:r'l:!". tl~J1lISC or iSiCD1Dl~mlio.~ for m,1k· ilng "~ur;:h q~;H~lit:uh'C cstirna~[)'S,i~CI'llHI~m~IlJ~~ Ibe ig)1oJlk~'a[1;1.h~Ut)' ~\lbdcllCccd nlah~ri,I!S im,'(lh, c(l ~i1 !~ l'~cy~!inIl pr~!"":!'...~ ~ '\V~ 1iI\'oi..'J!1 :l!!Il ;111em~i~C' 41ppm,u;:b hawd n~! ulmbjll;~lirn' n,U' ... m~j:~r ek!11C!l!:-a~~d ,c(ln~J"uihk tmce l!ieW!!.. i~l 1~m\."Jnr;l.~ ~ndl>;" 'llICcal!l~~ 'Ih~ arc IlltH\l' ual fa['tl1 i~l ~hr; mlm~c :H~d.iWC' :-1!roll~b' (:o:r~!ITIIl~d I)j' theI'CsidLml p'IK1SCS in ~qllilihrium wilh flJ1ni:J1 ~TleI~s

"!II

or

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Sm-I Carlos ,o~i~irl.'C$~Ilch~r(! (J II. In 111C'lollol,;,i]~ di;}gnll~1::;' we use ml~~' hi:\lh·p:r.ccit-;.ion dilHil, We 11J1,\'c lllKLl.y~d 1]~jJrI)' 17'.IJOO~r;i1iu:; ()f oli\' incphClllOCJ) ~r.:; ~rul1], 2,9 ~a1IIBlk:.. of ~Iii!:~k:i2 . iLIe to 'lr.;lI~,i1,gm~m~ t"l::mipt."I;S i~;i;tllit. t:U"'Crill~ 11. !OR!]::.; if 40S~1l!P~CS ~ rmlll l\.'lkl.-j\.t~~tllLklliJg!;!. furi!i1 Pldlic His.e. SOLlthaE:lSl llldi:!$l Rkl!l~. mlfli Klilillov1.ch Ridge: om" IIi.3·g SIImpl,",s) i"mn! Hilwa:i iOin 1,~I;llld s :lnd 8npcmr 'S~I1.l{Dli t~. u {'lm:u'}' 1~I~nld~, Rcr.,,~inn, A7.{l~."'. J:1l.dl~ll'Ilil(1.:

"l*

UI:>:'~(~(l ~pn~pbl Si.bt;ri(!., Eiuchdmu,

~ro"ll Ol]lO!l.J;l J;lViiI r",I::tt~.\1I.

41.2

RESEARCH ARTICLES
~i,gl1if1~IlI. n.eg.:llivc' cLIf'll!C'I~~i{m of N IMg vCl1Smi M ilti'F;.: ~ linc":Jlr cmrclallun we r, lieiell[ ,. is O.{l{l Ii;]r 23l'l siHnrk'S) ~11 ~C'~)r s.roJl~ Si)i kon'i[Ui acs lUlv",:: Ihe-hi:!!;!:l LC$~ 0 ,,'alm~ ,md 1'I'i[illlCl~ s~gnli,lkrmll)' ':Ill ~h 01i". 1m! fi't~eli()llmion (:;;~ !,)!,)irunpui'>it~OIlS or IJlh' i~"!cs rm'.il1 3 r;pin..:l. ;ll~id dcrl\.'I'1i!locm:~ o~NIIMg en .he- deg~,; or oihinc model CL!~"""S,~iPuncI arid Fill" :2} bLn ti~"','CniIC!l:i\l:; i!ill1mCI-~rc.,; rli:ihl~C(H)' ~";5tLI.,; Wig, :1Fl, llwy Im~\i!ll ~11lOll (:)LCt.: lr.l("1 tomJitlcm m~~m-[i(.", in rll]igo;.; 'COll;;idcf"Jbly Wm, 11.13 i!!UU D~ Mosl (,![,!lJll'dll&l~fDTCt be (~ri.... dJl.Fctl ~rhill hi~l· Fi.,g.lA L "Jbs corr·dnliol1 imprm'L'S (I' = [j',31'!r;.'lf cd ooUvinl! rh!."oocl)"~~ fh:llll MDRBi> ;md IHlmy fmm kmllmiilcs 11;.1\c MI\ :mr.l Ni CI,)[I[(lIll~ simLbr J~",,'e IndLillg of lKT:iJolI~~. 1.11 G1:hltf g~~l~ll!h~ W] ~ll'l1paC'.\l~ for lh~ l>uilc";'t·l. >Dr ol.h iIL~·:swil.~1:J ~ti~ 0 or 01iv ~1I('5. Cr i~l1.l;JrkOOlytmvr.r I~IDn ~Jt:a)t';cu.u liI3IlUW{'f 1"iI~]!l~of ~~ C{1jllti:!lIt~(Fog\) 1.0 r-09 ~} . ~o those or PlIrWotlgoL"'"I:!l'f~n,'L!l:I1i.l1l"', In COI1Im .... in. ,"1qui.libri urn \"~Ih [IlcridOl.itc nt ~!i[gi! p!Illil;l:IfL\'; ~10RB o~ivines :life ~lu_'rm;!",,! iu :-.ri:;rnliJ h iglK"t I .mo:~t o~i\;iJbl'\!llfrom Ib: WrM·TIIiCK g.roup are in M"n,wlwre;:", oilivilll'i!> fml'lllht' \\"PM·TUlCK I see l!,ll'ill::ri!li1C!m~1 tI~~1'.l!elll 1IlICI""LOlitc ~Ileh lng, !Fig. ~~ignli~kanlly d!.!:i11r-UX! in Mn nnd curl.cliled in Ni. I f), 111C'.~fiW(""it CIi,conH;:iJ1L,. ;[rlC"tbUlld .ill MORI:l Thdr COl1cenlrnL~llil[l:r1J not COI,I1I'1:itibJ~\:'{i,lh tJ tc gml.lj) IIITC 'l11e:111~ghr.1ilin >lu <LIm ~mv~~in MI], wi~h o.livin~ lrosn Ihl): WI.~M.·'ll ~N group beil1El I Illd~lng r!lnd.I,IC!:;;Qr'COll~lnUIl p!}J.idmilez.;, The oO.IF o~h 1I,lC'-l';, Indk"livcor uc_.:;.k1u<L1 ''']lilll~l. C.~ '(,~Il (Flg. JA) shows Il~[~y !.mi~rom~ ~()1:'. ml~-nnCid ~11.c. lvines JioOm I.~c \l1,lf'M·TIIIN:!!:l~lp IID\'''' in~:er· :~nedi;ut.j;~ 1\11;1,Llld :-..Ii Ofl~~[ml1.<j. oo~,Lti,i:lu WQlh [~a lOr all rock !!!Oup.1. ~".i1h 1lO!It~ihly To l'Ilil1ill1i.~~ ti.(." I1'flcc1~ of 'i:lli",~I'~tmclimm,,~II;;~ ('OJ1L:C!ltl'O!tiou~· of C.:lI :iI~fi ~'m\'i~lL:l ~L1In~ lm~~1 nlim:'lr {'Im\md 5%~n:~:Ilh'c e~lIiclll~U"lli~1 ~IDI:1,. :.illl:i\v paranl~L'!> Niln.~('jMg} ,IUd C:!IFc dL~("J~lbl~~io]l in ."pilL,!' Ll,r ~lC' gro:;]~!.;', i.'lH~rI.o~, Wr~~TI ~K'K aad WPM:1lI1JN I[W!:!!" MORS n i!ll Fig. 3. Baud c. Thi~ IH'~d!llf(! ~L'\{)~oou~ 1~~inl1@~\.\dr1Dm gll[)l!I~)[i(.\~t!:,!P n)iFc or~;bI:L!' !:hc scaner in Ih~tll'dill<ute ~i!JIIi !k;llotl)i. tliLI:01 Mrn;! olh'il'll;!s rmn~ LlloC' \VI~M~TmCK ~m~p:J~ S111.1" Doc~II)1 il1~ oren rind Nil }'i.c1d.~ N UC.n m~i[)l'> 11iJ!)!lijiginillf!: I I'M::di rr~I).:mCCll CLw'cC'nMCfIod)ln~mic: I h mo 10'111' in C:l: 1.0 hnve rrocif1i~;]t("(1rl'om i'!cridnlilCof immy wpM-Ti U(K iJ\ii\,ini.'$ Ihl~l, ::Ire unSi,,'\'Iil ings, Jeri, edmeus j:;j\(IlNIl as e:x:r-~rmCillal.ba.'llC(!J rno~~J, ~"4;~n IJ)Q:'I ~ion~ ;)111.1 1r-llCrill'JiL~~:i~J.n [ruji."Ctt). 1J:O;Ul!ll}'ili:lIh ,ionh.,; Eal!lLh m>1l1llc (Fig. ::m~, Fat!;! 0" recyc::~ed '@C-e<1l!1lllc cr~'5;t In );ubil'LlC'iion rri..:;s, I~~g. IE}. Ml1lfc is il~ p<Um~iid";ir l,.;.;:n;s'i J.,;p..:mJ~n 011 at P --.~ j G PlI, the b~~~lhi:c ~al,l gl~l1"Ili:o(li~ ttrlLtlll~ p IJf thl:! uc~iIlliii.':mi:.1 !~rclir.Jlj~Iortii"''d ,cUillllpkl,.:l:y CllrnniiLtlii i::. ~lmigly 't:rnlll'tlilcd by ,I,!"IIIitl.!1 (iii\1 itlC 1'ii1~1ioaill~i,lll1 (;I;,.S~il'\' iI i~... 'U1i: til odd g ~ll1dSiJ,urloCl ill p;:r1dolicl.":.~lIld .hUli. nlighllJ.c u~ri!l~ fm!:liunlniun CIlIr"i.",,'S il1f~ 3~ nIL~, it i~tI~at:-. I!u \.:cr.tlgi.;.: (elilnllp)'ffi:-:clt..: :~mJ, ,nntl,.:[~ willI t;l 1n:."C ILtl:>lit: of ["i1In:m'u,l :1tI;~.iTI~LCNlllKl!'i ~iO:Il~rIdm~r· SiO~ phuse I! J 15)1, Unless ~ilacn. 1]~iSl!!i;(.\rl ru clc'\:il"hcrrrnJud~ ili!,!iI-d(;gn::..: 1VIi:I1.augf o (")" we
ill'S'!,)

pl"t MTilfc and Ni

t:',lt~OS(fig, ~. nlKI Dt B

\'C~tjS, i\i!.l,l!f~ These .tCJ!lioo d(lITI01. \1ffi:1:)!

1)c:rid~l[il~,wh i~rll"'l1\ c rcsidUll1i!' ~n."i>li'l~ i

Cir·rich

rJKJi5.CS

'~i!t idue'S OJ

[r,um Arehc-:JillI

rre~

t'llf

t1YC-Cf>.. 11l\:I"C ~~ a

or

I~rth~

Fig. 1.

'YiII, t(J F}Average (lijmpasi~ons nlO&l highly !lQolg~il'Sia!"! olivil!ll! phi!il(l(IlS!ll in ~ad. !>i1IllI1jlIl'. (!nc:en·

.A

'~&«! .~ """".'THIN

'Iraliol1s and HtMr ratios are gi'r.lli!llll in Ppml verslis r'(lr'$teri1!e C®IiI~ent01'olMne 111mol vto,.O~iyijne group names iJr~ as :~ deliiledin tt'Kt 1fWI, liter.!illoIl1~1in!lilica,tes w!rr\po!l5i1iol'ls Il;d olfurj ntl i[1 ~qllJi~i~nurn a~ [I,:1L MPa.llti~1i1 metl oogfllally gell· '~ro1!ledi!il 3,Qw .51,0 GP~ Ir@JJiI f~fijle pe nd'otile (1~, catml.ated by Pelii'~~ 'SOrb.'1il~~41)1rtiF I;\l'I,rn~1l ( fU9"dly oorrcs jXlndWng to qll,,"tL~rayollntf·magfjje-til.e (Ofllit) bUJIDf;er I.!siJllg the I~er,lber,g model, C (4)" IPM, I<;nil:ler, olivine oompasilions :simflaJr t.o PM, HerMerg !but wi~ Ni oCIIkuta fgtlliy I)Jsing lMilp:3lrtiJtionin!J 100" tween olivine alild melt frem Kiinz~er t't 111. (28). Fm:~ 1 islln,e treml or o~j~l'Ie

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oC:omP'Mi~jCi:n1 during r~a.c~ioli'lal c:ry5lam· ~liM fr(J:rrilal rmililt d~riwdl frol1nt r~r~le

f}endotite <!'t 3 GP'a1 and 1515"( (12'). FradnOni!iitnon ,or ol;'I!'i~~ moi1ered l!ip t" 21)%fof 1llll~9!!1lrIJl9ac:'i'lV 0(0 flesPOlld·1:DOO 'i1'l9 t~ O~M bl,llUc~ l,115in;g1 '!h~ H,elliiJ.e!9 mode~ (4). Fr:a( 2is ~'mi1.ano IFrat 1. b~l E.~,
<ta1lculilled lOJ .35% iClry;till[~ati(m or melt dernved at ,1.1:1GPa alil~ :L630"( ~12tGril'Cf! IIilli~indkil~ fj'dd

+-_._ ...... ,-....,.- - ......__ ...... -...----.,

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,olMl1ie COmJ'liD~iMnS compa~ibte

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~er:ido1Jtk SAJl.lfI:.e.ln ~A. Hf,!Z, H,e~oor~ :s:ran.d5 fiorr rn lr!llilal~d (empM~tiol'1l> oj IQ~M!'lIe h@i:fI ~p,ifll?~.' ~1i!1li! 9oi!I"l1 ~,t"f,r'il:e namurgjtE! JeSlile min:g (4); LH U, Waher .:ill'1d~II~Z, Walter iuidka~e e'xp'er' ime'fll:at ,ofilliifle5 JIruilnn lllef2om~· and '~ii!!!iIl ~-[r,~e h!'llr.llbu rgit-e lFeS<ldlJlal O\I~mbla'ges, respm:TIvely. prodycecl by higil· pre5:lU n! melrinm aHwtile peridGlie' U2}, Bll:arl e~j~ liDarl«1~"ol ... ilow [,a.px
1
N

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g.e! ml2l'

~nd 5pi !'le~,;free ass<~mbtag eoof oti.virqe"Ol'l1d kI.w'C~ pyr(l')l;~.n.e.

SCI ENCiE VOL 31i6 ,20 APRil 2007

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RESEARCH ARTiCLES
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RESEARCH ARTICLES
au~ otjilsi~1~'r~t willI lJilhilrlll oli,'o'iim; trilm ,t Fi.!I. JIJ I, t5. fil anda decrease iii 111J:r'Jti~~ ur II'tc h~k of Nln :1i1!(j Fe !7~. Thl."'S1! .Il."mun"t' Ill!! r,~:Jmit~lll!;;[\!\',""Cl1 ~:I.II/Fc: ~H]~toodooalH~~n'"S, occur because ali\ inc is LEI!! I'Illljor :.il [cfll1.e ph~l~e unid l1lIixh'lJi. 1~J'liO:Jl;S (1m was usedto eomIltJU~ ihe 3Jl»mll, PJm;«'"il1,i,l~criv~ eompoin, p.:ddrjli~ COlJ';;ri:il~l~milugNi 3J1K'I ~ he rtll:iY5iH" O!;:Ht,.; rl1fl~;1o: injJCrkkll1it(: hm ing Kd (ur I::""gf\:<l!'-'i. ncnt iii, individl<l~i $ilmra",,, U'ig, ·n. 01~~ ~1Ji;,';ii. fmm Ihe \VIlM.·T! HC&{~l.Iil) {l1·b~~[I.J:l l'i.121(1 ~m s, thIn K~ ta Mil I i), TIle blllilk.KoJ! dl:Il1~ \'\/i[1 irWCfl:3SC 0I1lJ.1Wl1l' "he MnoF'c I'lIljQ{l!fI"lYIll..'I:.!milJ"." Nii b ;IH'f:d.!lP 'wI" 61 ~(,,% l:.l:mdari! devimiou ~
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'iJnh.;l UOml:JI]. mlL,;mic: erust, _OO~"lUI tltf' d!<lnvoo mull tllJ!mpa.rnd wUlh pt.::'riOOlltNjL'.'I'i~'lld the W(}JhJ'~ n]~r suilc,,'l ofllrnx!. b:ts;!II.~(UP~1. rnehs, ~Haddition. nlld~i~]g or p}'m'xm~ile }'idJd'~ dJ~iI'iv>l,ldfrom Ni COUIC'nll'i il1 Hawaii;!!1 Uldt Imver ('II C'OI'IlIMI\.-d wi!h peridotite ~i'i'), I\,dUIii'IlCI.LIIs:IDr!S nd oliv.i~1l's ollly <[.5t The oh\ UIlCS SL.'1iC't"'i'I.I, of 11mm:~r nruxlt':111Ilunlk pIUIII1C,S; a ']' il~ ,~tJ1D'f1glY ~ilicatil1Jlilll l,mcilOC:I~d dluH(:'I'cnoo"" ,~I'C. .h:i~hcr ~"I1Idl [rom some COnluL~lllai liP:,; r~rccili~ sui!.~ rl'(!11.l lind srrl11C" konmti il~. l fl:lI::'li,m~ Ibr hybli:d. P'Yl"ll1'::'C1l1ItC !Bl~1[I 11C~idQItC' Sfucrj,~ alui [«(1:1'00(1ndic~,[c a'lnUlS! IUm; rY.I'O~· ul:.deli1ull(lmle,d<'l:;::fg~dlllip.rt<;; 110l oo\·c.I.'1:ld here i mcl[ldr. O\""l~·ninemru rm ~lIh!il.llIm~}. ~:mlll~~ and higher Si :LP~, !n\'rlcrl" Mg i~ p'P'~!,(ill1il~ (l~~lk "'ill1l~~" C011\"5.pflr:!di~g '~~I1I~t.. 10r the' bud[', OOlbi~li~l_g mnfilly o f .d:k;a~ih~\...~, dl.!'t'1,\ ed 1I1C~1 ~_'i. 2.5• .?ti)_ 1}1J1t'!'WOOI~ ru~ low WPI\iHWN. 3'U 13~,: OCe;IU i.. Olnd :;rl~ :;.on!L} bf!!ld'~lllL, 1~llIJ~Il"!1. Lii'n :.;uch 01:'; raj' An::hron kl)m~ltiilC.. 21 W~,,: and fnr TIh~ l)l\:!dktjol!~ W~L~ !0!'tLXi by ~\f'l!Criml:.'iflMorU!!,. ~":'a::I!.IIdj.~lg one ~n~lml ~uwl[!fJnf11 !Jh~ Pi]e<ci Ln. Ttrhili.!UlJ ('O!p;;~ \k!d:c i"lmd .... Why rn.'C t,~1 m ....lulI1.f!. or a !1l1Lldd h~,bJ'id I}YI.\f.~xclli~(;' l.5l E.",p~.d rn~,,'1!!L';'\'\'l,'!J!:! rUIl at P 35 GP[l ,l1l0 SO(lthC"tu l\dillJ.lil: j:ill,l!:!, ~dowt 17 I::!%1~~ill1ibr S1!u:1! h;l!'4tl.~ !.llill lUJ[! h~gldy l'Jmeh~ ln 1n!· tempernllUt!ll l~twc,"J1 l-lmJP mul 15?1}0(_' in :~ w'l~I'I.".dicLiQm:; ~lr C'rcnnl'fl11!l.nrul ~hcrcfhT"::: pft'SLUll::lbl:)' LJ5)]_Bcc:l.L~eof IllC' tIIll[;C'l1ann" compmible &,(:I1C"IL~i.! by 'I'ery h)'I., d~~ of n;~ltrJ1l,:!.m:ari:y tics im olvcd ~Iil csdrm:lIil1ig the end-member CQnlv~llliL)fI[I,i 1,.21;':1111 I~J~IDn~'ylilltl~1I' appilr..uw; UiW~IY:;1IJIticr&~tllF~~i,;J,ll ~i~ Th i:;, ohlicr.' 111 ion i tca'? i ...n ul Ihe AlI~Ugillji~n :,..r.~"iml~JUni,,"n;iL~' VO. 1(.1), "~)UllpCll'illD:rJ.:;. th~ ~ifcn,;ucC;:\ bcl~.~'J" g~m.lp"" ~Jpo.:'jlD; to ~'Olt1!rm.lic~ LlotIl" tIln(kl; 011": "!oull] Iii!.:' IlU]lIl~n., These l\:.·ill.lIl:>.,klgdln~'t wilh Wl!~!~t,,;d ".\pt..'(i~ Ill'!.! bi:~i:\'"H11J~1l:i;miu~'111Imn ~1b.'>U,IUq.,. ~po;,."C1 'lImL :iilic~I,~itiJl1ll~'d mJ!n~ g.!:n~'1I'idC(~ li(,1I'i'i A ~lht'lLi,ghl',mJRI3:;; CUlil:tljl1 ~~ICh"l'\iI'c.... rmpOJ1~i1 tll~ii,l:d d;IDIi foil' IItdlari!!l (}f pj:r,i,a(~dte V2), Ilylllrricl p~l:Ui\:cl;i~r;.,. hJ ~ pre\l~ Ictlt ~ci,;ifll. ~hlJll c~lfI~l m ~h;: N i flml Mg~ rClmill'lIz;hnp.~ as ""dl IL~ ul:llIyfi'Xi.'l,i!~ ...{kri!'l,;tllIl~H. Ih,; ~~J'Cali ofMORB r I) m \''':1}' WW nlJ:U flriLCILUi"t:i. 1l1o::r..: urc )l.Cwir~'11 dam is, sigl:lili~aru. ;1~1dilIHU.'!' slimp Ie.:. do Cilllmii'l l:nd!IJ1,g ml.~~ lJI1.:~}idio.1l~OO''1.:' ,milk: 53 mid rig, SUb.:;;!.llIlIM :JrllOlun,~ 'of pyro("";I.lil,;.u~J:i\lctl CUUII.55l, I.~wnnIh{!S'.: dmu. w.c' (.·ak"ulm/ll;; I,."{ILlihbr.iUllii piJ.'>14ilJk IiuJatlulis. ~ (i~A 'V-olLlljk; lllilJl."Uy (_"Ol!'~ , tlliggcted fnc:lljnl~, ~)rpt.:ndo~.iEC IillilY be the 01 ivinc COl~"I:p!l:-J£i,lithl'l.'>u IoWi"rt:~llR: 't -I. ~-'7" 18) 1:I('IJ~I\t I ~~I(";(:'xucrne i~the ~Ilr.ich.c-d iil~ilic,l i MORS :;wnpJ,; (hm:1 Iile SiJulht'1l1 I\llalul" (~9~ OOrllIJll'Uilt rm;,;;hrrnl~nl l~nJttiJIU!l. sLRmgly sil:ic3in order LO co:mpmc them wnth Ill,,; IfWlliTili ul~KI~rNI~'L.II[mli."lI aUmlin,; nlllugmw; .~~,'l:nw~'--.r.tLlr~::; l IJh~il'or..~\ d,lla H'iy. J),. W;' Lm.. 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VOL 31i6

,2.0 APRil 2007

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RESEARCH ARTICLES
G.. M, Y;nm-,., n 1'1, GI'~!iJ., ,St:lIl1l'it. mmor.rW, ~if:()gt. Milt 78, 2'=:3, (lJS'911), <'0,'", B. 'tz~tmi!ll, s, R. H;!!r~ S. III~inm:ili\. ,f~lm l'toiMn
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,2.0 APRil 2007

RESEARCH ARTiCLES
1;."Ol'Ii(1('1I1 ilJ'i!llgiJlg ~lit' NIB I'Llnl,!I!r liI.;.'\Ic,otl;cl thin l~NA~ (lrng~"$iJ; DglS,L\.x1L~ MT d":ll'lsiiY im,'kk:the of '11iCSl..: genes ul:;o pnxhlcOO Imlg. :;plndle:>..m :>rilld~~" iocl.Ul:>l!d tliO'lH.'!ptJlar :ijPluuli: ioonluiorn. l)hCl'imy~lC Ch[tl1.l~[.CrL.,.ic or 'NuiluiinRl"\1/\i. 111L1iil and efm,;;'cd ,,;:jlJ'l1mo.'i('1iltCI'kiuC>1.oo;:h{l~ nl~~I~Il' rncnl and l1Iit0~ic u..:loy Wig. 21'; ~nd '111O~~eS~ (ltl" j'mdl,.;rii!!lUClill~ig .['I. mtc in 1'-1llb~lliu rlilll~tll."I~, ~n ad',dijl~mllo C(!!TIt~L:'iYDmeloc':]:i'il'''Ltion. I~ ['ILlbo- D,!,U5 R:'>.!}'\'i~.RI;n."fllly, 'l'l\dl(' '~'i'lb impliC'..mtt"d in Ii.VI!' '1::t"11 illmging. mnl.·o:r 'l~'Ji.lhrlijn,-"d the d¥ccl ll.~ ~1~bldl"Hls:>!.;~:nh~y dll~efq~()jm {SAq .~iguml~("L nfY"'IIIIilIilHll kJ,r,ali:r.o..~o the ~plndll1 U,;. pt}l. g~lll~ knorlrotlwu on ntiil~ ~OC~L~i2:3J1.ima l.rne1Oof ,v(ll!.:.'1'l'l ig, IHight OO!1Lrirnll~ to J.... IT I]UI.':JI.!31.Km II;huL1[l ,I.] r' llC!'lll.IDl\flII11~ In'Otl~J"l'" ing'LlllOLigin bimli!lg Cd~rl ;lIld BllbR ~ tllJ"hul WI' W,~lll!U !.he ~p]m]l!l 1(16), !~.tm~llb}fy.~nio]] !;J!([I! Our semen ii:lm~jrw;d .. ~.s{D 1lI1:C1(pGeIJ,:-d OJ 1l1.'i1ldts.!liIIlgg~1 Ihtu tbe los.-;, 'l~f (lmll'!f kiR~WI!.."lo MT ~1111,.';1;:LCliLll~S. IlCll:lD\rnr ohpjl1dk;.localiZl.'(~ an~r Lmi{llO'i,\,1il gencs (l'1:u pmdlilc~:'(I; mneiic :RNAi ),TLIIl:.(, S1lbtllnli~ i.~ roqu irud rbr spindle IDC'-'I.I lzapl1m!lJlypa>.. t'<'Id of which W!t'i cm,timm I'll' ~ tkm. of 'I'·tUlM.I.I.i1l i.1l 1'Il~a'mnlllli:m cells j 191. hLIL y'fuR.C m~y COI,l'i11t1UC'111C Ill'! mary IFca';iOJl .tor failure .~I~:'i:l.Y Ihc SAC. to 6 rt'J~11 i:'''!:'l)C[km.'1]!~ ~I".ig.ID~ h~ Ih~ ~hllml"" otherwise, lialc is ,km)wu abmilithis popu ~atiQI~ 0):' !l,pi!ldJ,(lo.,~C!(:il~~.:iT~ '("~u~lll~.~.ln L':!l,Jr screen, itl~ see![OIll\, \\~~ ,~CSI;:ri~ 111<;: .mlb II l'O.Uhsc!~ 'rlili~ ~1119:Y SiLl,ggC:-1!.~ ]11Mh\\~~"\ ~~or y. l\M -nllColC' !!l<!nH:')1 in ,!;:t'!lnm3fhl'lle ,~Fldy~!!h\.ll in rime! OCI]l, We' id~rltiliedglt:'n~ ~h~~, ~mc'd,~d kIL~'IJi""I? luhuJin ]oc;~hm.t~nninl '1ni~n~z.,{.F.ig, .2n,..Al lhe 10' 'I'c:lI"Ihulin Do the :>Ili,il~i(!! ~IIU!L'Llllt!~ pale ,tFig. 2D !:lilt:' .'i.l~pc ~ L:!ylwmic:> !i:lfthc 11oO~e:-.. ,lIli,! ~)i!~dl~ "pindl!C' fiO~tJ... eore ~I of Wiroldn~ bllli!dt.:('n" L .md chmll'~n~ <I!]Jgr.unC11l 11~C'IJm'l11~kL~ .md! lig" S.I..I\)'.~ 11 t t\nlIjIll~ ~h,-,!\().. ~~ ml~1JXIrL1.\Jl~~ tl"iL)k~~A~l!1 ;\rl:l2. S:pk, DSlliM.~lld SoM!), whit;h !lnwi~~ !'£lUbJdll SJx~2. I"elm ki1~6..;c. li:..1~f ~CIl~ WcUIW ... iu ni!C' :<iCtOOl ,.-mo be rmUld "j unhl:! l'TuRC ~])grip71 . .Dgri~~n. Dgril) I ~K MLI ilil lrilbk' S l as weilill ill'i, ~~Ir WclJ SL1C,\ ...'hkh :11s£) lJ:grip iM), !RNAi, (1 r:oi.C'\cr::J.II}I~\'ifIl.l:-1~Y klM.-aWII rmd (:"11IlOOlOOI11'i10 1n:::n.I.i~.,'4uhLlI.in thMLlg~] Ihc n ) un ylhS(, s~IJ'Uilj~S. I h)'!!'i'c~'I!:r. Ih~s IXLLhwa_yj"dis. gC'Jt~ 11)g;lJi ~I('I DgllM '1f;;{1 dimiUDishoo yctliltnillin cOtimii',",,~rirnm_q'Y dam un ffi\J ,I\~rol~lfiqCL". h("lnop (",;;111; isi!.m II u S::!C'CI re... j\ ~'COJld div ::;~Icct~,{dy~, jt~l,i)l, 111\:: iJidb: 'O\JrnIlHl"'tl! with 111. pi.'IJ::>llblc $P .. '~n1ll:il<,i.'tJIGprn[l;lu kr>Cldi;GHlkm Clig, 52 i11!Kl hUIlJf trlllli,J.m,[cdLI.lin iiLlSpihlllc:9"."tIJ"'1'l~ paUnvayith oil. ilfl!l~'Il~W:)1i.~,orrrIJkill~ j DgiJ.£J pol,.;:, COU:.~"L",:nt'" utll >I TIl!..:' ill iaq"..;jj nIl;: y.tlJlblJllin Gent ~osl)'me!!·and ~~.,tL']bl!lli 1.oc;<lliz<llij·o,ll. ttl< Ihi: :.rla,dk, GFr7i::J~i III i t Dgt4. DgIS, i~rid D~~fi, ~IlltillJi: IJlit~T yT LIRe l>!ibLil'i~bR.">C1IliI h 'j''''LlhLlI n ttl "rilldh: l\,n~, Sl~'l~ri.~in~Jy., thi~~ih: or y>l,Ul'itUI'llli 1'{!O:::lli:tetllLlI'lilimi'l~y~~f:1 ~illllllC' Nil.:.;, ,,,·j!l!l no 1<."hD pacr~lII1I~inli' r!'lr l11i~IX>~i>.UI~ c':l~hiit:ll~~, dUlll i· tlm..:~;in;n, is ITIoIi'I:: ii,llpOr'IllU~ rl:mll nil~ C~ml\)OOIil'1lC ri{l1111n~'n1m Ih.: Nlitr{l!:mmre; lh(.: ~iullllL: :;~illliJig ocmc, C'rea~iJ1'g l,wt)'I:.~ubillliln ro;l1mil'lin.tg 'C~irltm:rorul;,;;li,U:iiIl nlJ.(;~ml;; MITs aad uLlljnwdy iJo;.::~1JJ:nc wa.s IOSL ~PUII MT lkp;!llyul1cliiLllLiw:l :lillI \.~"','lCll iu ll!uild~.1ga ,nlUIli1il~~J,,;n~,1h lliplJ.)a[ iSpjl1dk' 1i~'ilih t plXlpool' ;:d 191:IOO d~ lil}l]IOSinlfle:;, '~hc l)Orc.~ lhi: milOl.ic l!i"pi~Klk;. IIO'l'\ioi;,;I!~T. t'Jl:o cyoClc th::pcndel1l. trl!) ~n ]OO"lltiZilliOll in irlll..~ln:;c J Tine- shape ,000nd dynamda. of tine' p@~es .• (fig,. lE Itl1d Ii!):. S4U), ~Ji,!:il-lilroughptl~,. ;JIJIR1-o ~riOkiS, arc mN. uct;.d~d lor 1:11.0;:,1: ('~~ di",""I.olli:;, in M L1lltk :;plm:1I~""nennal ~yImVi: 1~'tl wdl,f&l;LL..... 'tl 1li1......,;L'> M,T m!ld~Hiul1 aru~mdcllr'llmNnin ,;;Lll1k:~ Ilmb.;..u, :imil.girl!! ~lrJi;v 111J;c ..'1 n... Gi.lll,~irll!ll1\('hl.ll.)'. Mruliil'l [lind I [!B·GIFP ;l!.'>w'(;l1 a:;. ~llglill.;n.';:;,c:d~ltiOl:l pelcs, ill our' o'iCi'h."II:'IfI. w~ ilil .. nti ri~tl II ;St:n'C:> (~f tC1r hilll!:llilr:;fJjmJI~ kml1iiii~lIJ U3. J6,P'~. In our :M:'Rl,,'FI, d:CpkliQl] {'If prok;ro. iJwo~\'oo ~rl c\::1~hi{)l~ dUI~.I.it'lliOll' wiJ~~d hv L.\.'qliOC","ocl 10 p:rodLtl"ll! n C'~.~i!t~~ UlL'Il~lru [)far~md ~lil1dk~ (no Y"'lIIhlllin stainiug .III. tilll:: poles ~ all'll m[lnl:L"tml hipoliIl' !1PLrtd:lC!'](m~~y 1.2,7,6 om::i'iI!)Ik': will! OJ]rI111~'f,(uo.lIlin ~!a[!l~!l,l;::l..liIcmUil~ we d~.crmifill ... "{~pn),lJi!ili 1(l~'ffilit.;..liu(;J.~~ gnclo.'il
~hlc

pnneln (Gf[)], l;)lJ!,!~1J,l {3S !,1,l!n~, tn(l$[lylitl~,.:d m. iw'll1 N tutti C ~~1ililil'l~ to be c~rL![l~1Jof ~6u;;1:~JC1Ilim~iOli p;LHCm) (1;lhic 51 ilm:!! li.g.., S'] TO S., ~<I~ll~~Y-"'';:.u NJ\ i piMlot)'i1\,';:O by . R
OI1J;.'S(Xuc:c

'd-'nlbLllil'l i'>'I.~ililirl!b\WiIJi~!tI! ~l'lrcc.illg ~'Cillr'll!{'1lnirl I;} k!lW'ili Y-1ldJl!l~ill l'utealiit;l1iul1 filCKlr ~ JHJII, ~!N ... \j

or

or

or

"0

'IJ~

l"~~

ror

ror

or

'C~ml10il~In:lllll~'S

"ll)~Y

gmdu.l~~Ydimiulislwd \\'ilh

dil",l!1ion Ii'lwoogh ~~lil;.oc~"ivc ~~~. ~}'d~~ in ,m~<lJ1

d ..RNA tw..lll1.Cul l.Fi,g. 1,.,\ ;:!I!1d li!c!- S3Ad, O!.!r !lcrcCll kl~niHk-J sc\cml k!'~lWU l~ri)!dl1.~ IlS"k k~I'l'I~'. DS.I~4. S...,,·()l a~"''lC1I ;1,>1 tllree aui!' . p!oo'l'ioLlsi.Yl~llikn~l",n g~'n!;l!\ 1~1!.m~!rn1 ~I'llndl~
j'D11;::ntatypc It/frml'J. Cm~S;iS'lC:lll wnlh lhcir RNAi
p!hC'u{lI~j'i~.W~P"Ana i a~~,j-Am1l2 c{!I{)1,,1llhood during J 1'I[..;;11}11l~IImli lIli~J~l:;;P';"...illl ~hi; oJ:o!.::lllri,(l~tlr nMr,,"~'lni nl1tFII)-S~1'<'t'i tr:i~ ,::.!B and i'lg, SJiD ~lI1d Slik :IIn.:1 DSl.t:.:..oI ,tr~" S.3D~, GFP·AlTm ~ \\Ia,,~ 1101, di.:t~"C[~'rl ~A. :iluStmJ ~l}I_lldk ptlk-:; afl\:'r S:tk RNAi. ~lni[1Ri\IA~ d,,:piclitmtJf Alm~. l\:"~ullL-d in :II ~llbst1'l11iri.l.ti!.1CiFI...J.i\l in fJFp..sm;!) nLlur"';~Ctlltl,) num :>']Jllldlil.: poil:::;. MI~[in~ .1In importli1l1lJU 10k inC\]IlLLioll:lDrmtll,ifm Wig. 2['), TlrnH, /\:I'wl (md ~!II~

}\ uj;J:! (imcl polJi.>l'iblyAlm3)

'!TIm""

limy

Ix; COI\; l"~UI1JIO-

ofnllm

,o.:mnulic: thm

lIJ'cl:K".C<.":;sm)

luI'

iC:L;'r~trnlt.::d'up~iCaliL:lII,.
RNAi I!ll~ known ~'Lm~""Spd~.2,Pi:i!~, cL'llIlI!i:li;amirl, . .Dgrip%. r.md Dgtip91 ~1ht~lDgri~lli. rn;lkc' LIm ~fill~ SI:lilk-co:!'c ~yTlIS() 0 f ~he'l'-II!!~tdin

ur

r!llg CfI(!l'lpk-x (yTuR:C1<I dccre<!,~d 'f-ll;Il~ulil~ ~~laillillg ~~le ai, l'I(ltc \\'~Ihout .i~ltcrJ1;J1 g '" il,h CC'l'l!n nh;.re fI~rkcf IfiC",~iqAd(J~ U'ig, S), F' !'111~(I ~ i3~ O;:1I;J:mJnjug lh" ~W~"b:of RNAi (1:1" IhUl~ gio'UIefi. ou

IF~g. :I!. ArrIim.agem,basoo,g~l'iIonile--I:Jild(' RNAl ~(lreNnif1:g1of metalpiila~ s.pln~i.e munpl1Ology. ,!A) Dl'OmpJl~tjCJ 52 'ool~ W~l-e h~~llW, 'i'!'tth ~R~~ ,!;If llli,J;l2:S; ~~Il~ '~~Q:Il,gw1~1:i (,d,4:?iA.pd d~!l:~.A. 'W!l.t;)I;l.IlT!l.I!~~ me-Mp11ine ce~ls, .Aher I'l. cla'!f-i,~ells wert!ll.iliIlsr'enedafid adheretli o:nto (oIiiA·c~i3iledi !liIliiI!fIcs"l:irJt~Clm pla~!ls. Cl\1I5 were<imm!Jl1~st.1!in'ed for [INA., n"~ubulirl, 'I'·tulillllin iIInd j)mi1llld ~mol~ed by hig h.ithrol!gn~ut
ilIU.tQ)HiliMfiI mkroSlopy. 5c1I1£balr, .20 ~Lm, (9~ MiM4:1~(e~l!> ~'l\ere al!l~OiillMi(:aII~ SIll«:tedl by a c!oiiDpU!.e1' (llg~,rilhml~t detillds p~.3 ~t~~n~!'lg •.:rn,~se:~,tw -200' ,"iWIl'!; "ds,(m@~tly !IJU~t~plll1~) \'JIW,~ dli~played 1.1'1 aga ~~fY Jar af! O~5e1W~ to s(Dt~ p'h.eiilOlll' pe'S,.(() Ph'~n~l~peswere aM'il iilaly~ed by rnlif1llUtefil'fll:1r fm.ag~ segmellllo]lic!<n (1, monopo!a r: 2. mul,Qllcl:ar.; 3, ilXl'le< d~tll!cl1me'.nt;!l.. kmg Sl'ifldle; !:i.. short splnd le; 6, , misali~nm~l'iit 7, ~i3:fgl!!' !""tubul!in <lJE!-a; fl, ,[Jim y·tLltDulir:l~ (H}. (I)) lweM Hilarj~r Ilhef'HI[YPe1: id.enl]~ie'll~ilil
~hl~ 5.crllEl!ll,

IOC;llil;;,~i~lU.\l.'c :i\lg.b'C~1 a n1.<tl!ceuhir IWlbh~\'.Eiylca~Hng[{l 'j'-,u~ldin

>O~1.1't't1l0mill,~m.l 'r-'~\!lTL~~iil

1~~I~~.ifm[j11Jl1c ~ioole I'llOle i.rig. :!!GI, We

;~!il.l

fO~H~tiI\\'O Iwviou::;ly ~inkl:lHw.r1 .b'C!lCi>. IJgU a!~d l

Dj,:t1 tdill1 y"tuh~in). d1,li~dccreJ.q;:d s:rirRite f"Ole

S~ate

oor. :10 JlIm+

RESEARCH ARTICLES
.:.:elit~:;Hlarunn~c~ •h~ IH~rt'iliICr or nrguili~1lion poles. {I~.bll,.': S I ruu.J fig. 55). Uu~Ocusedi kill!!WcJmWl! IIil'l!;;t'S,~K,f~I1ei'$I.i1I mI"C pl'l(:uotype,. were {)b~ai~l~(llhrRNr\i (If ~llmod~l~in K"aM I, a bro.IJly flnlJe.li.tllliIlg CillciulU ;;;ijl~~.m Fig. 3A l, ,r B,Y ~imli1·tllp:,c un1~Lging, CaM RNA.i c!L~M'\d K·n~rd~lc:Jdlln!o!nl ~rilln 1l~('nlm~ln.rs., :L findill!! timl mnl1llkkl'lG ~(ll[":Jlnrln.~ tor RNAit:'!L Ahm:lll" .n1Z11 Sp~lcll~ prowi" (_i\Nf,1) (11m\, i:c 'S11 t.:? 1 I'. Like ,i\l'i:I). 0dMloc'ilhaxl m Ihe mim~.. 'Luis Qr K lib~, ,C'\'Cn when tI.It'SI? ema~ V,'t"LiC di:sr,o.mlClC'lCcl from lIl'CCoCwm.wl'nel:iy NC(I(Dl1~ RNAi (Fig,. 3B). rl!lrth~m~=, r'.IiM [Wl(9 A!'\Jl II1~rI!I,lnlly depended I,II?m~m!l!',luL'1II~'lhrlOC;I~i""llio,~ 'l'lll~~ IrnblU!!i end or Ktil)Cr.:;: ~r~g 1'U,I", MIJ1, '~'I'h[l=h]ta;; !i1lI~ S5fl liJtli;. a1f\;{lhloJd C..r..Hluding 10 nw!il~ ll'~}, .rcqUil'Cli, CiILMr;.~l' il" IDCZlliz:[tioJ] and IhnC1iotl.

SIJlimJ]c

or

S i.uprii>illgi), Ill,,: sf!o;!Cilkil)' of 1~1£(·3:\'] RN,I\ i r1tC'Ilmyp; S!.lg~l' Iiltm('uJ\B mai:1'I fillIetMI (huin.ll mil'tl.~~ is SC[\'~I]J!,! [J.') rL ool1lCtOr i~]A.9J'm~'dimcd ImlC'-'liliCusing.
An
UI1,1;.'\llI."Cl.i.:d

IlllL<:nu1:1'I~ W11..~ illl,;~i.tlioo

mn:nlx:n. of rlim~::'II1!m_~ b~Xllar but 1]01. ,~n,lL'iA:r,:d~ !-:I)imlle:. ;I~k'i!' RNAI til" Irl1ttl:lip'~ pml.l~OSOm~ ~~Ibunlillli, Ltl!:! E2 ubiquitin ~",{lUlj,Llg:nrJ1£. 1:!11ZY!'lllLl'
[Jbo:[)b, tlmlrdDfl~IJLocmt~ctor ~{yh. ;1Jl;l CI M)'binlm.r~il'l!! pru\clll n\v'i~1 iFti~_ ];['L Pm)[['J]so]]1C' RN,\ i. shcl\wd ndK'I" lll~~].oIy~ .~cb ,1C'i ~ Ilmlil~r:.n~{m dcrort«, hi Ih~ IllJOll'l>;lr'd.! bll'Klibr ~IJjl);lkp11C'~}~ \v.J_~ 11f11 d'l~IlliCIClio;lic o.thcr

dr;.'1lmlllbLC \\ 1ili.'lIK'f' i.'l.::ll!roSOU'II.:mmllJll!IS were rothlc,_'\1 (F~.IJ,3Dl.ll~t~w!:"\ Cit'. ml~PfLr..~bflc (p:lXIl.CffiOll1C<llld lJoclJ6R.NAl) Oli irll!;:lllCiI' !~lyb iIUllillwil RN/\~: see :lLi('I ~2JlllllIlYi' .~ l!f PffiJlltiil.'>C ~~IlIRI.''''." ~ Wl:P: ~bl>\lrvaJ n... mrr~P'11fl1:l'~'I'ia:1 (,'ofi[rnl.cd~~. :s.Uggc;.JLillg 11m! I!]t~
SC'I~~i: illr".~1I1'Lo;c In

RNA.i

Ifi".t!b1'ltmJJ;

Ct1\1:>I::"tI ~mlll'ffiolaIll1l' till'l'ial1

~k'r

NEB In b'l.ll1t'GIJl'~llO:1]or;tmJ bipolllr ;;pindl(;'::i. To ~""I this ~dJ..'41, W(; perfonnod ,~lIIDJlIl~~OOt. !i\'I.' iru!Ig~n!l, (>r cell!> (OCxllf;t'1\!ijng "I"luI:iLU~in.GH' om'l
111nliOflry41.dxJ'lin ~FLg, JE ,:md movie' 5~ ,. M~ IU>J/U cells Init i,llIy rWll1l~ Dmmlil~ blpfI'.I:u' tor

.RNA lIIJ1J'(LlUlC!~L~, I.h.~I\ldlilf:fd Cl;'iI nU!1ll~", M'o~ lfU'!~. R:'J'J\j. or Ulx:lnfi did InL'il irlhibit ~w/'nii. To 0J;:mlJf Ul1.d~t.i:md dlh rh~,\!lnty~, we SCOlI.od Ihe- IllWlilb!.'l of l'!I!lI~l!Ilin SIlil:iJlj1lg c\Cn~

or

ocrnfiit;m:lll}, 11l~ I'tipolar, !lJlhld~~ 1I ~r N I~.B_ I !cwo'C\~I~ ~'I'I1'II,."qlJ.Cll~ly; ar nU'!!1'Iof thL: ,L::NIi:m~'
tID.~1iI1~$ dt'IlI~11L~ t{l~\. . ri:m~l till' ~~[!Ind

\'I'OIItd~~J1l,{~

[rd

till' !l.X:n!(!'~' Ii!! OOIlUOlilI !fI C',QJl!..~:.d 'Coi!~~m'£O!I~lC'

~\hi':h

lJ~va) fl'mn nh~ Sf1indlr:

nll!JI!Ili:;lru~ bipoillf

~~4a,I! I

dYIl.;;lllR.~Ai. ~fId.t'l.(Ich and mo~c

!i.O'II:IC iUl<;UJIlCCS,

:;pim:Ur,; ;Iw,> ~n nil li;.;.~l1 eells, This wha10lypc Irl~y n:llb:i l!I d ifi.'I,;~ iltliml {~f My.li !I] Ih,; (\"'U1tlMliirL~ or ooLlid h..: nl~di,J:Lru imlim.:x:tl!' LlinYU1!ih il!>mil.: tn gr.'III!: Il'IlU!>C'rJr1i!JlTI, hl CiJlltrns!L RNA~ 'uf l)~Dfr illi.lln:~-d (~~IIItt!i>(lml:C lit:Oi~Jll sho:litly 3.1It~1'N EIJ, .!!l:J'Ltt'illi lg .rnonopo.luif I spiildi~"S, wlilirh ~i1~11 rom'Ct[lrl irlm Ill!.1nll1st1'll: bipolnr spiud~bi!! by "hrontLlun-dcp;;:Il,lkm NIT,
g<;:t~mlj(ln.
TIiILL'i, WI;;

Cl:Ulf'l).'>Gmr;; (tl\i..i!m was

oitiici.'\'

ed, cncmnrlg

(ih~f'I:cd

h\

opl1-"l'itlU~ly unreported
crl1tim:>Olliir: dynitllilic~.

r1fil::lioly""_"" t'ClIiIXCriil\l.:

LQ~~ro
cornplcx

uhiq~~~t~u-{k:fl!,:'1IU;!l;IllII!l('jlCu~)'~is l'I<Inlll.

~v:J)' iurduC«l~:..;n~. CClllnl'OO:!U~

tlt..[OIl. m~d M},b

d..-g)i.,1.iDn C<I'\I!iGd ('JC111m:,ml1el ~11 ~t1!Ci· ,.;;ili(m~mw:rd LI~~ ~ri~liJk interior (r-~g,3Fl S'lJlfndl~~iz.ll a p'ldl1:;h rQ:!i!IIa'SQm·~ ,;i ~~'glrllJi1~'IlI~~ h~ ~diniOJ~ in .r.(r¢~lufyi!.l~ k,llm\'f! j1I,,~II'[jI.f~ ~~f

S2 cell s:J~il1dl"'~e!~~1Ih tJ5t

tll'li!

~rec~~

.[~,,(I

IF

. D'l'I

iJcmiHcd f,I;\<!ral W!~\d:i,'l'!.b.ly Uq,kllll\\,u wnoC:-. !!lim. Pl\x!~c~~;.h~~-!>l~.ndk RNj\~ rillCllOlYIl!Clii I ~hPl1! s]lincll.c' ~.'lllpl gen¢;.],. and dulC'e Oof ~he.\C Inca~izcd 10 l'Ilitflli.c spil1[llcs IF!g_ 4A .• ~Me $1.

I1g. ~(\).

ICGJ3UII

1I110;,.i"'s.:n, II :'Llrpn~.ins fl.JI~t 11!.'c.[lui~r: ~I;.:'''l.:ri]l~ i.~ :r.m.::l),~bM~rYOO in UU.U;il::l~~~d. 111~ )],C\~I;;'fl.'tI t"IClb., MT.., Ol~willromilllNi:l tu gr.m\ ;'11 [I~~ilf p1L1~ ends umi !.kiX'~.ym":Iia:: m their llIIinus "::Iuis.. cal~iII:J!!, ~~lCm [0 ~~r;."tdmil~~ Ihnxlgil. 1.11": [[[ipht.:>1llt~6}. cy Thus, :il.TI4' It1~UIuIr;.iS MT ~\'rtir.IE:' ,mod Ih~ .-;ilwrlc:r·spil1l1:UIi l)h"'lmtyp e~ llIIighl be due ru Fig. 2. Genes reC!uir~d for llocaliiing y=tll!bu:lurl to '1Iht oont~o5~l11Ie '!JlId lnl?spiJ1ld~!i! tAlI (lert) A!l0l5~r<t1 <I~ Clllmrw('rl .MT ~~'OO Ilg and clcpoIIYIl:K'r.l;{J]liml ~[ •. \'Y~l as mo,nastr.al, mP'O~r :5Il1indl~s l'\!~rt (!bs.en;eda:rle.r RfI4A1ito s:ak OI'!hree pre'llliour$l'j' Liin!rnqjWfl tI.11 a,gel11'es ~Ii::poles, ~th~ Mwr b~@:s~m~JUi!1 fig.!i3l', fIl:i[JnUJ (~nt!!lS(l!lll~ nu Iilbcnll~ ~!J!nt!2~ (1)r b'ip©l~r ~pfudll"S OiIWlilr r~!IIr Omf!l;.·r~~11 alsu Tt!l:llII~.ixi ~'I1~ ~ll...(}.I\'~!tIin r·crulf1ds IillWA:i treatment. (10)Anal ij~far m ~) WiU, ~ot(!(~1il..edwilt! ~he cemriall! m rker mRIFB'·5a~6,'(~ O CllrolTl10~mC !lligmmlll]!. jf!:lblc SI. lig. Sr. and A~all filNlA.] (;O!u,s,edthe loss o~ G,F:P.Sal~6 fr,om thi? PQ~e regl0!'! or ,!Jl<lstr~l SVill!dle:s., \!,Ihe reas SAK !1U\lA~ movle S5), Orrtlll1!1'lS(lIl)C mi.ql!iJilI~mC'l'tlrl\.'qll~H'1l1tNj ered \l.!iln 'G'~P-,AM :Il11Joea li~a:lion UI polt:5. (Ii)) (Le,ft) Variml"ii i'J'i1'll y-tub iJltlf'l !tNA~ pnenol'ypes ..The t Iy oohtciclcr~ \\ lnh ;1,!~i~,!£J~C'i~ s,p~l~dJc IC~~~l y"~!,J:I;I!IIlhl ~igli!.al.i~ riildl!l~~d ~t tile ~~li!trQ~Qrm ~11l;)1iI~ (C!lll),,;:!t ~o~~ !1iJ~ ~~ntr~~@IIi!~ ~li1dl~~i!'lgle COIlShStC!lt with ;1 nllXlcl in which W!lIrlpfl<'l),IH o.nll.y al ilh:~~ilil1dle Wgt4 ilrli~ [li~rip12~)., ~1P;igllit}Iri~LmS.1tV <It <[.:en'tr050m~ a~and $p'ind!e u,'1~le ~ l~, ~ de 1);;t;1;!,,~ dl"~m(l$f)Pl].l}-MT ill~Crn\;.~0~~ L'l:';"~i~l[1'1 regiCln (b) was ll1Ie~)ur,edl ilnd lJ'Ie lrelativeilFlt!llsily wa~s(i!!J!:iu:[ated (11 i. ~E) :Sp'iindlrehXillirz,nitllil 'ClfGFp'·
1

llh::1'kt~l!l 01' (lHC 0-[' these p!f.l.e~ls ijl!:nm:d ~IHl'l ('~l»iW Im!~HlLl!rL1.'''''tI MT :>CYl::rrin~ in illiJ,:rphlll~~ ((.'11:- ~ Fig, -IBrmd

~1iK'nfll.ype$cncock' rnutcins dmt 1~lilt~1 :I~ Ihe rni$a'Ugtll~d dlP(lmtl~liiieS+SihrI)Mlis (i IiliI~lI:lriiu Iillilillell5it~, pr()lecttDrl tJ:f' fo ur ~pl~(all. ~iCfls o:b.t3rilil'f!d a'l .$ O.2511mintp.l'll'als Ily :5i1111'1I11ing·dis~ ,eollr'oc.al micNm~ope. Reat antilood:y t,O CENP'-I\: grl!e~"l'ln. SC::i3~.e khlCltlt,'!1"fre. and t w[~ ,(niIlli Sf~ lind C'( i5 ~4i'! , Jl~S() locil klcd ~t (';'C!lll"fll'l1lCilCS in iUltl!tT~l ~:\'A!. II lIdC"~ 1)<11:. '5, !JIm. Hi~ M.oCi'!ll for f~rui~.me.nrt pa,lnlHay 01 y~lJLj'bullil:!o the (!l!iIlJI1r05~ll1Ie:liM sfJindtE!' M1I''!;. t (fig, 4(" an(! liy. S7. Bond (), ncSC' ~h'e :genes Pr;e!,fl0:uslylll:!~f10WJ1! or II!lnJ~~ue!d genes <lr'i? hi~,t:hli~ht'edin Iblue in this· iJnd other figllres.
WW!.v,.sdene:emag., org SCI ENCiE VOL 31i6 2D APRil 2007

r~(!uired

D9it.4 i$ lost afire r rCollltfr iflle-'indl!l(emi fl~1I' d~~olyiitl~r.zatioli1.,. RNAi o'f iD!)lS, (as well. as. other gMes ~JFJ.

..ll I

f,or ~lPindw w,al~z1lith;1n !;)f 'i"t:ubull.!1

l)

prflldll!r::e:s

h;lln'!ll ~i!llidlle5'

wHh hlw

~n dill'lll:ii~:~ and

in'!t"'al"~tt.'!: ·(1f I;',roc~

act~llg 1Jl~)ontate" ::;pimil~c

'I2'J~, ~h ~ c .b'CiI~~wilh ~\'~"m~:"1I.i!:iH !l~~t R,i\lJ\ ~

RESEARCH ARTiCLES
18
llill(l 1'J1.'o! GJ1lm~

l:ilp-llm~"CJ ~I nOR Hl~vdm!.'tl pro1.:in (R,~~Loti ~JII!IJI. 81~o prt'lCh.u:;OO un-

cxpecicd

RNA i chremosorae

mis.al'igmncl'lt

p~ICUOl.ypcs (tabie S I). The misal i,gIUJ'I.eUL,;Ir~ts ItuPKJIf uad lUrcH 'RNAi as w~iI: mi· dlO~ pmlll(:(;-.d by Iht' 'fOR. inh ibi~.o:r I':!II'lZLln)'C'inlI ~:i~. Si! I}. '\\~'im ~m~y oh.,~(.·J\oo 'vll'hh CdcJ!7 :JIT~1.. llt'>.JAi 0.[" mmnl't!"ilLl~ ~il~lioooID!,nL' l1[!1nlp;riIlC'.lJ1.'ii also euusod Ch~)JH10!':!.]111~ rnisnl ig11111 l\!JIL A I· though this f1 nlding i1'i h.II!lc~y lln1.c~~Pl:"C'~JLd. mu~~niaJ1R in ~pJkil1g !'aC!Cin\ cmL'1C ~ni!illCEil'k'g.'1· ~km r,Ir ml]][-chmrllQ.SiG!11t\" ill ye:1S1 (3':). RNA ~,.~[Cing migln ~g~bi.'h,~rrntd f1l;~ .ill' ~h edlin ki~~loth~ s~w~'tureoL'l~' gL"I'!C!~ut.c RNA~ Ilm~ ll!l\~ "'flJ~Ll,Jr.Jl wl~ \','~llIiLl 111,,1 ~Il~nd~c ~3JI. Tlu.l~ Cill'[!flil:JIli~ ~ruC:lUl1;' .• RN.I\. and r;.i;gm!ing I~Jdl\1;'4YS 'i;:~n inrltl~.ooL" dmntno:.'Dm~ OIli~l1bI!Jillt. 1J111l!r.u~h Ih~ mcdmniSllll'i .on~le!i.C clTC'(:b remaln ro be Llm:l,;;r:sI.OCI!dat a molecular level, (o'ndl.sio~. Tliis nmrphulogjcal ,$\:f\:lo:JI ~f

~Jr

IF

P,~tmwaYi5klr
l'!i!';,MJ p'~eoo~

th..: Dn-'.Hlp!lj'la

matoii:c

...pindle

Wi].-;

g]it~!!lc

tK~..ib1:~ b)

C!,niipi.UllO:'r",jN;i~I~'1.I~tkilti

Ifit:iLtirnl

td"

l]li1:oti~ t,>;,;ib R,r \ if.L~.d ~~lr.ial~ mld '~lIIll1lJlil:ili,,~ OOIl1PUlill,julllfll irnl.lg,'I.: ;nl:I~J'~i~.~kJwt.::~ ~f, 11-11.: ~" sWLUm -1:&~.g""'l"; RNAi ·'.tiiN·~i~~, lJy il:.'>(;~l:was
in.'iJllllJicicliu m, !;,(uin new ills~in illU!) spbilI[l: for· i:lliUitllll" l\ Slll~'i.'! soo.mlllilry ~1s&Jy~,wM nccd~..:I 00 d~~iflh~f tile ~il.,,:: I ~1"1iml am:t 1lI1 ~ ... -clmlli::;ml ~)d" flll1.:\ inlli.~j)' IIltkllll\ry,on, pmteius, a.,.;. w..:11 II:. ttl d'\':· !(!.!:Iop aTIiillk!i1td~~-d L.md!ciSl:1IJnlia1$ o.f how !h~~ 'llrn.tc:ill1> woik U"lg~~lcr.

or

••

,.

\l!llMIIli'l-5

'\

~~'JliE
a~ntrosl')me ~epi!lrationand iHMci!iiit.lalrl W ~pil'l.!1I~oles, 00 l:delrbtical p llIii'dl)ct!lsingl ami centrosof'l'le-dl'tad'imel'lt. IPhWi~1'l'1!les WNe 'D~se;rved .a~~e~Asp and ((!Ilmodulin RNAL SllIe- mOll{f;e 52. 1(18) GFP~G;iM \ViI1S 11lI~lwli!~ at :SIP; ru1~ fJOl~~n >i31lA:;,Il"d~pel1dl;!l11'l manne'f, but wa1istiill d@tecte:dillt liJiI1foms~1 poiesiml!iu::ed by NJ:d/D:hoo.4( R:NAn. (e), r,!jlma5~ri3lI, tJif)Ol:ar :<;'p'irll:J:le~pliUd uced by UIb(IIl6, pmte<l~Om~rubI.Ul~t ~15, fir ,f1,11.y1J !RNA] {p~rcerU:agE!' lor ~eU5~!jjitlil p:h.t!1II0lype 'il'ldicatfd1; .3% tl'l ISOJ')J IlIlltrMl~d .retls). CeMrosollne d'el.Gc:i1l'1lerlt aiso was, ciwNt.oo arte~ Mlyb IRNAt (iO,} At pr~f;I!iI!'IS~, JTi@~t P'r@~~@~li!le(PrQillrMl~l11~ P~~J~5~j ~,bo!;l[)6. Myll, a,!,~dlT'wi1: IIUjlAi cells (rj ;;..,65~ ~!'Id

idneoo(hore-,f'iber

Fig.

]I,.

Gene1: reqllifMfof

~QI"'~

Olddil ~[1, i~l1Jl~iC:I~iJtf! mil!l11)' I!!U!C!xp;.'t:IOO nhi" ~!lldy hm'I'C\'~ a ~cd Ult:IlllkbpnlCu 1)1'iCi" oesses iuvoh cciI in )ipi~l:dllc "ss~lJIhl.y.F(lr exaruIlk:, ~hc- MC!l1:irtc<ltki'll Clf .Dfjl 'Ilmid!l~ !od 10 .he lindhll!;l I,'Ij~~ ,.he :tCIR it}' (It' ·~"I~Lh~~I~n \, h:hitl ,he
hl
,1,.re11cs,

ron

rl)~

~"Pin.dk· L~!~~

cril!C;l1 .I(.lf )1j'Iind.!c a:rchih ..Cjl~u:c '

;::.J:. centIO"SOmie-; Sak. PI!n<!1. tCf,nltl'11o~l!duilltfca riOIll), and PlV ((~Q~~I1e:si3J ~emd as con!rot RINAis '!l1al de(Jea~E!' 'Qfil1crE'.ase!(!ffitro-SOrDE!!luml:te~. 'l'imf!~lapsE!irna!l~liIgor y·tllb.mn-GF,P (r~d) ,;;i IIId m(h~rrY" 'ltI~lilnl (greflll) rmRii prm.pl1iMe to metaJl'ha~ '5ht:J.\vs, ipo~ar ~liIndile 'Iolil'l'l<ltlof'l"a ndi!belll Cfn~roooon e b d~'~dhrrtlllll aft~u Myb ~NAi (130%) W = ,~Cl). al1d immedi~me ~!ffitrQ-;S(;lm~' Ius:t!lll ,~f'tl::J iNU ~@r Ubdl)6 RNAi (251lJo)(n ;:;;;2!Cl).. ~I.l m(lvie SJ. Scal.e ~i3rs, !S Jimi. ~IF}MiocMs ror V.ari!)lI~ Sr;lilrdle·!)O~e pI1!l'rIOlypes 'ifil ff'SiiliDnSJE!~D \!'larjoU$ plfl]'t~f1I d8plpti®n~.

on

and chmmnSi'iIl'l'C a~i!lJUlli~"'th;m il~ ~cr-kU~l\'!,~t ! 1i.!l1.;~iflH :It Ih~ ~"Cn~('I'~'!C. \VC ,~Lja) UJ~m\i~'lJ U~liUlli.cjfl'\tOO. R 'Nt\! pl~~L11j'p~.sL!Ch nc~ cX\X'So she CCnlrosmll'C 1ffi.lsiou U.Jho!;:lJf! lRi'>JAi). CC1'11.msome ck!3c1lt1lR':I~1nd mm [oj} within the ::;pi.mBic a {,~'I;.'l) il:.NAi~,imcl m:tiv'llWIl ~r MT ~"::H;ri]l~
t(S:>p4 RNA;),.
IIImdj_im~ lill>l!

~m!.'!rr~ring"ith

&t"u.mrl '!:cllu.liMir

~\en: mmmWI1LteJ :11IhlJ ~:inl!Hdour sercen, iJ!I.Il sequence ~Ji£mnenb. purlbnlltxl by OLIT M~ll!1.11l' and {JullK:r:; (] 7i idcrui I.'cd C'G99'38. CG&90:2. <Iud (,G is ~56 ,,,",[ly I1mnnlog.~ or Ndt'&Ol,~ h:,,,~ Nlll12, I.

.LInd M3~12.
IIW{~

,1,"(;':;p!..'t'livt:ly.

ddttl

gcrl~ CG5 i41'1 ~dlr(lIOOroi1ilc aligrmK'1hI ~['alii J mu:ICG n4~ Wall I did 11;01

disJJ1:~y :i.\.\qub"Jlcr
Jlre' in'lCgrnl~~ alllli

tiil1ilil:!lrlty

~o kilillWIl k.trk'1D-

ChOL~ prnCL.'ilki" Tn IJlncb".Il\l!lmd IltlW Iht~' rru1C:illlli.

Hn!(IUCm iln Ill.:: mok't:llbr

;psscl~b]y :I\;1Ihw.~y or the !Jnl''r:opl,Jlfl k,h~cto'Chore. '1I'~ .bl"c;~igmcd 11;tC,kIoc;l~w.;mifll1 (k!~1i;-Ire <t,.~l;.1nb!~I. f\;'ll~~ 2 <!.TKI Ih~ Ndc!<iO CI\i(ef d~K}' 'If lh~k!!)clothnl't f!g1IJteiru; 1J~~g (If'!' fti~ion Iml!leit~ ili1d RNAi. ,!l;NAi of Cal:! al: I,dn~htll\;' !;."Omr~~xwe ~lUi100 in :~ 'ligl~'U Icetoo n~ih"l4i~ I('ICilli;tfhtioj'i. ~d·NJeJ.:() :md Nl~l2. IMhw •.!}'. ,.,;jmil'~Ld H.alJ~!!1ih~ rnr (". t"l~f.il'iji.~ I but i1i)~ 'W!MC'"Oil lilitli!l h c' ('il.~llrom ....t\" l'iDOC in~~ ~.mthl')'{t" (.11~. &vcrrd :kI1HWll g,C'll~ illdli~I~I\g tr.~I1~riPIH'~l can" Mis t::!. rnld C EN P-AI(~J. !1.uwHurtnml!ll'~

wilh Nr.kl':O (19)1 1l1.·I~ In' Dl'mitJl~ililtJ SP'l:2:5. RNAi or cu n, en L1:t!.: 'lllJ],.;r lWIH.l. [1[;; r!."C[~d IOC1]1u.'LLllIll '~I.'" IIITlmtCCl' le:;k~ l Fig, OlIn: midi Uig, 57•. ~) 10 r l. Di m:fI:'nt rronl (]I1~cr ~,y~" tenia t30i" G.:1'llnmilClli: k'\l,Llizutirm:;; ~JliCENP-A. CENP~. m'KI CaH WCIT mmLJ:l:ily dependent, bCC'~lJ~~ RNAi dl:pktiiDlII cd' any ~i_n~k pmk'il1 JiJ;rupl.~'tI ut tIIinliL1i~h~'tI Ihl: TOC..:l]i~~~n or I.h~ other Iwo. IhliL~. S2 l~U im~r k ir~l.!X:h{l[;C r(liI·1l1 ~inn !m.~' in,,, flh:~ J 'CfI:--L~c!l1h~y 'f'!f:'iCICCM;, .. or «ENP-A. -('. <t:!1il tllC rrc\'kltls~y unkrll~W~l Iln'L<:'itl ('~~ i ,Fig, 4LJI). Once dQ~~C: ~'li'C IPro\cir~,
:J:climl

!~e ni:>!.: 1.0 (]i)o,lil1~L :,rim!lk~ v ikll,;:cf...~:dh :IS 'dlWiC seen wirth n.lI~ 'P;m~",":Jl,OlllliC '[llImnmilr-dILJipof;n" ~pii ll\.":>~, It!,,{A II)tfly IIm.'CIO.'i.C m ll l ~lL1tlg "'l)iIKlk'S). HI:WI [1]1.; ~plj,(;~'O.';{_.[r:,,; t~hro.
1110:'101];1\.':

mi~~lignin.rn~t 1hi:; wmk

art;o

proof

"il;k", II :sy:;t(;n'llll,l!C' analysls 'rlf

tlK"

'l!.'i..'!(i111b I)'

rilusol1ilriu

ct:rllrul>!lmcs :lI'ld inm,,'lf kllH,;!tK'I"tUl\,'i."\,

mrlcrin.s sevcml prm.cun:. irlto Ih~lC:pal.lIIWJ!yl ... .Mlln~,' of tit!: Il:IL11CX:1J :.md JX~th~.Vlly'" a=rii:l!..'1:1 k.";i" in Ihie:; :,;rLLclyare rit'iil~ w Il.: wrloon'Ca in hursan rnl~. lh.·r(ll~m;the alk~lr.n[{!Jh-";in ecmmsome IlI11C~ 1Iiud d!I'OInLl.!ifllllC' aligltlln.:11il OhSC1'1i\Cl~ imll in ~lis SCiI\.'e.rr1 ~~y IlJnvkk i!'l\~J4ht Illrl ho~\ ~h.. ~ ":f>C Olnmlf1ltl~' o-l):1ct'llcd dctCC1S arise ,iII h:-JJmOH C<I,~(;e~, ,3'11 JfI~
Re~'~1I!ifll:1!'5 ,MalIN an~ 1.l!jilCi1..ilol)~,IE. [I, s.1'lmrnl, ~tJt, (til ,rlIor, "" E:1.1 (tlOOI). 2. ~ It r>'l!llltiJ~. IE.b (flil!1clflul(, II., II. W~t. Alm~.~..:. [iI'.I\1 ~, 81~ ~JI, l'!l'.~(1OOl.l, .. Lf.

Ir:I 420

Rlf S.pc2;'l, RNAi ill lieU! cd IS. (J~). TI~Ll"', al.:! C 2(bk i) rrtllCibl with knoOwn :lwo-lilybuid h1lC~-

filctt'I'l' I srO, c-lil'01l1MiJ11-bll1,'dillg r·t'!i~ICj_gl_,_-';; !Drnt! Dijllm~tif!lI). J_m1~.gu(1l,iilg p,OIeins I.llrg~t o.f

3. Co l.. Ritmr. A K!:I~J:d!~':o\(, .s:c1fR~ ]00, 91 (~~}.;