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nk lWital}iU1ty ill tl1C Offspring

n:::vie\l' li:;;::iL'1-th::1,; of
'1"Ie","O in ofj\cpring of imxjiated pl1.rt!fH:<:. The ')1'
:mw:ntlofi rmf.'::' ;:n t.he l'l.n:l"c:q>.b,;d is ,irral1i,tted
, r',,'se'H'dand di"-LT,,,,sed, -Ttv: H1l'cil;;,ni,,!'m; mKi Jt
er."icmd "'""",1" W'.' ;J,b;) di:<w;;s;:;.rL
in 1.9..+7 by TiJnc-ft'cj'f'})":::::';:;i)V;;ty <UK)
54], still cC'E{'[sDc;; ,)f T2d.iation bh.-,logy ar:d
t;"nrr"",";oj'], According tD !he w(;d<:rn ir:tcIprcla,loi: (\f it., Lht of
ling1ubtllC'n ')fl llW 0.:tpos'L'd GeE ;w;:-: k' Iht il1j;iui r)NA t.he
dUJt;' c hLl by C,' n:.t:t:e('t;,::d hy fn: f:iJ;(:als ib a
lrni$@ticn water The Elll,fi of tht: ti.q;::':,l
"Y'" lIAi eXClil$iy'!y occurs in JLyt\:tly ;,;;xp(b>.;C'd
:'(lJ1-HT,dn,::d ?d:d nis-fep::dn;(t h nth.;;)' ,,,--'{HY(", eyer;,
'(Lleil,,,!,,,, tle recG?nincu of .. it:-: erair and
md'lClj,<11.lurrelt, in tilt tXPOS-tJ :.:eik ThkH' is ;;<!lotflr:J inrpixum pTd:lctin:x
ilwefY of llldH,,;::tiou if; dir';l;tJy
,,",;Ito tbe mnnm:H -cC :radiaLlO:I .. j OdlKCd DN,'\ lhB:WgC and the a.bil i1:Y of (:'."'p()';(:d
l't(, tI'P'tOf it, trF)aning tJ1at that 'th'D yield of tm:HF.ions iitr'{}ng-!y
1:'7
depend, "n ,he
i':Y.:mdd;{ln of .! <!
l of . OC("U)"
rbg tJJIi:lEnt H,pt;lt DNA lcd. whkJl fl)T rut.. -, by radi<\-
1.1GB 116, lK n:didiDl1 ge'HUEiK dttClkd the
progtEy 1"J nJlslin-g'::'1L;5;l1\ J \ :j,t ,1.5.1- and
dJ{,>2t. an rnutHli-OH rHe: ill llllh.: ce.lb, 'ii'!i:Kh
sirn<;.h, t{Om '(he b:rndmkJ neighhour:" J:;';{}, 4:i-. <5.1. togethe;\
;)bSl'r\iJl1o"HS imply tta( dfech; of ior..bi"i:lQ; fw.!i.1!HW ::-:a:n.
over a priod of tiw('. nUl,d} h}llgt'! 111ft!! preyjQ;l\]?' thcu);ht ,u;u m2Y, th.eri'Wfc hJ
n:::QardD:i ;:t!i ;1, n:mIF;)t'icnl al t!H:: T:isk rwbnj"'l::,, Alzl-,;mgh 1 t;:'C>:n::
I iN'st'EAR CC'iKhdcd th;;1t the ;:;xb::!u, data (::llKiil:::srgcted
\/c"'.,._ ': ,., .. ",;:,.,., ':"k'>p!: ,t"s':i:;t',fm-}"'l'
'a.;:(, w j\::;nl:;; (h (.i)..rl.n! , ... c'.! : .... }" " d,C
d' thjs ph,<.'nmJ,eneB \Vm he i-nrp\:!rt:l::lt;n impro,,'ing, ...,
lbi:s i"AA:l':t!diu!.ry a:fkC!:!i nr i;1uiXlEg 0D
and i-\c;.:;nrding l<i til.,; H..'Bu;'ts of feceln PllOl.!G1U{lllS, It%'se
:lil. ::h:' cf &.">. \,,;elJ a.s ;.n their
.. phenO'm,(,lloll of DC;fi w.rgcL0d .nn,wt::.oTI induf::ihn in th(:
"I . I . lh",}
irr:ildjm.;":.( nnrnmtHS 'iilli .. (7"[,, l.. , ,,' ',< : ' -- . I"
;hi:-; H':Y!(:'W tb:n;fc,n: the transgencT2:tkrna'; cd
1
j
. .' '1',' .' 'fl1"t'J j: ... " htre \'.'(..'1'"
t:) r::n}WJlr.':JS, inL: tilIng 1OiH;':',!),?; nh. h1unn.. ,<.,.U , "'.
to '(cSt tht :h.at H1 thr; gerr:-d d
dUe-;tl..." >tlipnsed could martl.fest in tbe nffspri,1g, ;:iT(:U:ing thdr muUHutl
wteS_ tlredL.;r1D5<iLion awl ;Klp-c-r 2d.'i0 pr<1.::;e,:us
rCgent illll.:b.ni:; v ... bdi v/e,tc not d:\:'<i:US"'C.J in "lUI' n-:yic"vs ! 3. 131 .
12.:1: Exprimelltai Data
'''':1'" b1f !..llfl111il C(l\'iork;;:::><
d1t: r:n;:-qJt.:'ll.:Y cd du,m,inani, Ukd n:uHa.Lw;:i iJi tbe "'g{!J.'J:ntbe of
''''1"",.ld .'" 'J,,:ng "II' :n{:(: ;.t.:n Although the result::, sWdk:::
fIn;!. indkM:ion .tor n!' t!111: F
I , :liCit""'", if'" 1.1'Je' kmg lin);;.: the> had r<:rw.lint;d incnB.d(;::;i\'c-o ({ mdn
'0' Ih"""" th:: bd( uf a Sf:n', iHve lit'. tOt mUhttlori ddt;':,.:r!on in tlK trennhne

,[tad p:rupos<::d tbw,. CA:!.>Mldcd sjmp!e tum.lcl1
k,,;,;( ti.n pnrnde. a upptoach (nr moultn'iIl? mdi,,[;i{)n-induc.c:d
aB'ar,un ,n "D" l:Wms.k gt!rn;lin.;;; d rdative!y iihun
long, md bcl>:mg tu t.he !.:.w;wbk loci in th,>.;; lIlDUi!.(" ,,\'ith
!llcu':lti",' ,.1,;,: H:::
y1lbtlt" ilth :111:' deVC1Gp:sxmt ,)] :itng;:>m,+::cl.lk Pt 'l{: (')'t"'xk:<; 1'0
[6:; 1, GivfH ttw<! t{}-om tli<' :1.<)!;f oj:
"ki'nC'"'B m lnd.lc c:\p(\icd to imdsHlg HI),::! dlcmiol
s,im.:lii.!' m thaJ: (:;)r g::;!l('S [:!-, i"j {B. 19, :25,
':: med th.::m for lilt: mw1ysis (If tc;ws;:<:eneratioDJl
(lei' f"''';,,,bJI,j,c,j ,,> ,ilb :.lVi\Y Ihm mtcs -:.ul:;s!;;:.l1tiaJL'
;<T"'I'"" of and Si:corw ':leDcnitbn oif"prmg 1 . .\
'J1' an :h(t F' . bni,i';-\i( '1g
11 ftt:,;,wigi.n.it:r.utwruJ is n:.'ru.:.nisc';,/ ;Ii,' rhe ilbn'\.,'f
w,1 pbcrll"m""'IJD Lnsl.ariLiiy in ;;;C;Js.
Il1lpXiantly, (illf denomtnHJ;; tbat lb<:' pncncmenor <)1'
i:)st<:hiiily is. 1101 a inhrcd slrain uf m.i.c;';" 8;; mtwhh;
eJe-';Jte-"j rSTR !l'mtm:i;:'B r;:w;' wus .;i::;:ectcd :in tlt: g('!E:.Jim: c:f thre,i;
:;;t.ra.::os ... CB;VCn. C:')7HL/C1J (md BALB/i:' i.':l],
peR. Wf.' ,dso ESTR ;n;;:atl(!ll irl
"''',t,k', hom Jnd In:>!'{:
F ofr,;'jI'ring nf lurtk; miGi'" it' 8 or :: OJ' c,f un::1C X nrv:; U< 7, 3(f
1<1 lile tCC',111h ur t(;cSl' studies, s}-l(lv .. ir;'t:: thut hSTR.
,,,,j.;,," r ii" d:.:.vat-ed i1:: lbe Hnd :<:Of?:latJo..' ri>:SlJ(:': ofT) u:ffspring
'irmdl,\i<'d mak' iIi ;-;cma:l<": mut;itlun :
;"nd I, ::: : .. f !
,. ,,,.);;1,,,,,,,,' u)t,!d that n,O?it 2'onl.p:. ll:ng.rL ta JdJ.res.SH!.! >;pn ,lli(' B ',Ll1.IJh v
t"hPf'ilg "f J,C\.: l[.1J:n ;1H:&i.af\s." ,11 "oloalK
;,){ the pi.nk<:yed uTIi>tabL:: u:n.d,l'.hm [:1 J.] ;uti. at !tw
".,
ahmc(',memiom:d. rc:;u!t,i de;.-!ch, Lrnl-l:V ihM eXI';';"", l(, 11'1'11':."", r' ,.l",,, I.
" ,_., . ',vi.M. ,< .... "'" .;."",. u. 1
,." W 'n,' :indu,:!.iC1:fl :jf 3; tfCHisgtT,eratiood tH tbt: g;:Fn!.inc f pMu
the gefl(,mc:s of thdr offspring .. br HS
(li"iiJ:lplk'al:i Ii,.,,,, H' 01': (hel:;r:' dk,t;l t;x rad1nl:iu!! pro:eC110E ::lfC (';!tlt:C'wtd,
,or how h:ing th;. .. ('x;pns;:d maie,<; ('an 'remember' hi"uW'i:' <;If
h a !::,\)rnl:ier \h1r ?L1h'licD.tJ.nn, we i::"jI,;;: l:\lldre:iir;:d iSHtJ<,. ru ifue)c
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Imlli BA!.Evr,
, rink! !l1m yz-t unknlw,'ll d' pdNmtl IlTad:iaziun
'''",' durir:g the ;-:-,w'ly or dev;;+:!p.
to the ofLpDD_g, desl2,bi1ise ,heir Ow'
that: the r,nw.s,enerJtii}oul eff;J.ct::- rn.1Y il'!1 an
cf tht: sendk: d.-:k n'i md;_atioB. The :;:,mdysi.'; ,A
in the gfTmhne Hnd. 50m:ltk. er F) dL,>;pt'ing of
prcyi;lu,:xl inrCitt;lrt ck-:::s on :b) r<)Xs}tYIe .;,1'
lo"r irrmllurinn :.;udiild ,n ttl...: offspring
i;>.xpDsed rnale".\ <1nd the design
,l.ikp<\'(-!t1 i<t i:;h :F'I ;".! H.!lde::., from the hm:h
.,6, 5lj, FigV!'::::: 12.2 f><;"tdh ()f the- (We) fittldks- The
,;If s(l!rm:tlk' L;l U!'f5prir:"j! of 'irft:tdimtd: n,l:'_e Jnit'-;;, TkJx,
:iT!:] rq;(>n.eaequaHy ekv:::.ttd \{ it\crsions
I.no :m,)c;"e UTI!-lLtb,k both aBeln; fn>m the i_H<lJJ"'d(:d
U,2at
:l'i;;;: itCf::-klBCd '..'If the hprt
mal-:- P, otYspring ofm,ik J:H::ce U--:ig. 12.2b}, GiVt:fl thm rh.:: !tpvf JO(;;':;
;'V illLK'" ,', die n_":suLs (of this ,;wdy therefore in li:ll? v .. lht:
j,lt<'1. [t;" ,ttL ,,::,j.:vdci '(at>:- ;>1:, fc,md. ai, ::he E. ir,c:"E-i derl\;ed
!:k2rly imply rhnl
!!!!iil!!,m"1\1il) ,S 3t(ribufed :0 ,1 g;)lWDL>tiidc
;:m:s<:T:_kd. in ::b-e dwpkr ck1trly :;.,hn\-v (1;,H rnut,atiof; mtCf: lYl
:)0 L:.nd st':nmic A uff.;pring of maJiat<:d rnalcs nr;:: SUl;q;U1Lblly
lt sh.ould be ':li{: ure '-it odds r'iith drre
81\ p,'e(i,,:li,,",' of urg-er JS !ht) of iT!,l:::1if('Q ix,
:TN),'C;(pos,d ;}l1\;pjng, lhluL;diol:l inductl<m in un; d::arly (.ff
::hcrdm:e :he pfleW>menPH of t(l the
the U'!H\::-:riyi.i!S' If;:H\\[:tJl!t':-!;-',linna: It) ,A'
bLcIDtWD' li< has 1)(;,:-:;::0 8Dgg-e;;;tcd phcnmnr.m:m rJ
r.i!her i i! tilt' cLmal pmg-eny of iITadintcd '.)r ill
parell's he- ;lE-;t;ri\y,;d 10 mKh
""'"1,,1,)( l'b,,"\ l<!pcdnJ I,' "10\,' l:kI:l, n:!i!ll! <)V ep.iF;;:tH:!k e\'enh; (,13, 4(1, 44, 4:51 <
)<; b .. ml t:tie 1\\',', "e-.ts of exp,';ri::-nent'l!. daw, .,:;huwing tl1:17 f 1:
itl \.'IVCl geocH;-dc (l\'{:r alnng
after tbe .intt!nI cxpo:iUre; (ii) (he nUmhi.': f of rnanHl,stixW
i \I,lo>"",:! 1,,,,wmicio'llllbijl:' :is Un {(1 hi': !t};j:li::tirkd hy the
of auy gwup of ger:cAs. A:, fal mi the tnln'''psn::::r&.tle}ul Df paltlnal
-rw(:< main \l/hic'h !;>t >
PO)
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aile,;
firs:. in ;:jucldaH: yct m:kucivm. is W tht?:
cdlnlarcvenX5 t.nggi';.:dng "m sigmd ill expnscB The
({ tl1C triw:;rn1f,sbn Of su(:h a tA} dI>;pri.:'!g anet its r"ion"" nt,"
wWf.hcr S;!.;p in the: durifkw:io.n of this :\Sll'>:'. '" """
(}i'vt:n !:nwt eJ.pi)iUCtre 'W iun.ising r;1!diai:i(!lt 11. wkk- oJ:
123j. til(:'; it:; lr;ulsgemTati0!wl "meets may no): ?rr.rvIJe
C'I-'ideI1Cl:;', .for a specifk 1 ypc of DI'l""A d;;unagc an lnf>tabllJty
L :\,1-'1"7:>. Gj<
,llili: l:NU. )!;nd.,;::i!(tl
Strsin
;:-e.HS< it hlS bc::.::n :mggcstf::J dun thur nvJhltJoH-int.1!E.Td
ckElbJ.cq:cmd DNA (DSHsj lTia'v cumlilulc m.ll uf the that
fble r.he ;)f.gcDomic im,t.nbijty i .r7J > "Ii) whedwr thh iytX;
r.inWB.g in:;tabihty in the germliHt'. ',Vt
::Jfuct:i of w ;:dkybl.!ng: ,ih':ent
,)n thl:.' manife:'ot<:1!.ior; of gew;mlc ir-:st;lb;1it}, 11< on"pring or lH.'ded mak:
UHnrH:;!. h:! in;<J:iJtinn expmure ENL lJw.!dy ;,,]k;,btiGr of f)t{:"\
<:nd n:::.,uitiug prt'dnmin<.wdy in. tnse n;.H!cKizrns
k,]d." Ie ;.my in the y.idz; (,.e DSkh [50j. Th.lS wort
!1! il zlUml-:...;:r of griking i.;llnLaritic',. beH)::;;tIl lraw;f:::.:neratii,'Hud eg'cCb (vf
,,:xpOSlixe to mut1gt:ns ising nidi.alim
lXqLJ1.md if:. til;; ;'ile(:t;TE< G': EN!.: ind:.lcr:,d
,<A,{lHn;;iiF, mir dula .. i" ,"It;; clUrlhul::."d ,(., ;::.
""a,,:o;lIb,,"'!rJ UN/\ 'ks::<in
l
;, !,"!!'.Ji fI:, DSJh hut pm1mhly by <1
fb!}\),)Se to (' gcnei'alh;ed. DNA da0:1JPC Our d:.llil c:.re in lint \vlr.h rhe resuh . ..;
Y",,,'i0U'i :hat eq10sIlre tn ;::ne;11.tcn.l <::ard.nug<:::ns and m1.3.til-
io a dehye:d in ra.te W SOEhlt":.C cells IS, ::;.; . J6 .. rn
the iitnl.2< ..,; ni' o.iI.'lFd:ng of rat ... [2&1.
d1<iL tht tcau:!<b;c-:m;.wJioTIal. C;;:Ji a:mxt m:uwtio!l at. m:ndt)Hi
DNA. 1c.}>.::i pfc<;:ln !.;':l}ding genes .and ('bl'Clt1'()$ome' it th${cJjt.::
ibc: p, . crnJd 1:3. variety O[ type ..... (If D;'<JA To
d.m'k1a,gc. in the F. {)-:'fspring and C(H1!rds. 't.'i/>C I;::ioitdcll!xl
ilieidUil{ol' e",r\,bgefltAjs'" ;':U1lJ DNA 1:l!x:ai:; .. n:-easured by the
Comet and H-12.i\X (6}. This aH0<.lysi:-i w>!e.ak',d; un dHjofif:Hlly-'high
75 !fi!ll tlALBIc
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;u:tli)':.W! uf h.{(!iig;-; ;:h't!-' d' (h) oj' DX i\ "q'""i ..
)n !J}C ex 1'h lXHW ;;dL <D;{t3 fr<>m [{j
k'-:'{Ol of l}NA. ihu:::!age in the FI hotit' 1ll>wmVf ?;plteq ceJls (Fig. 12,4<\1
thaJ in t!;'Szie;i \vith II high 'miLJt!c >:t:, marro
v
.- ilj:(Ul
span of comal:ning dl.'iett:rlDllS k;):tr>ns and dOllble-stnF,(\ brier,:" ,,"
r
data c.leady that.
Del.uering: 1..-;,:\ xlnllti:pi-e

the ng of i ;-we! ihtr::::il.


rdkd me-II' L:omprG!I!'-Oed
!TI'!;',)f tld"lf"""W' The: i::tvoh'i::m.en:t nr nr.l.lL""
m!i1C<K)""'" 1r::...'! in ute:> in

[40. :;':\ J. F!',1c1:j


v
e oxyg"1.m
t)NA inckdiflg J\ingJc ,mel
h",3P:r"nJ \'<irk<iy fif 1111( 11l.cditk:a.tif;S I?i2 I,
K';C!iJJn,:r'; l'm c\:l(;, <;hLcLl'C,ir,y the FPfl CO)1:-ct w;'iay tJx: F( offsp:iIlg
ITadi"lcd 'li:1L lUi) al1;;;:r"jLoo 1r: lht: JLmaged
k"',ideqf'ig,, i2.4(1) Lsi(,g rhe C-C.lnid, we <1>':'0 me;l'Sun:d th0 cHh:-iCIKY
fO':1'" Hi ,;i,,"i,), ',!"!lid bren.b; (tod ')thCI alkan bhik the F, eH.'il-o
,Ii"""lh;'m' n;cu!\'W s<tmpk:, (Fig J 2Ab). According 1;) ;}U[ the et'fi:.:iel1cy
{he pfhpnng ui- inadi;Hcd JH(\ cOlIm+; :i:; t}\o:dbt.
an,;:: it! iir;:;! wid:: ihe ::u;nHs d' c<1.rly cornplring
M1d deiayed nrJi;Jtious ai. hpfT JOClh it] Hw
'HT{\d.i ..ued ,"<:.i.:.:; nB}. A;,,;:curding {h:c of ttLs iiwdy., the 'i<jJeCU\Wi
lhc \)(igoin? in:o:.tnl'">j!ilY _"ubsClmi,llly diHen::d
re-[ inducljll mK1. JJ ito;:: linh::, V,;j<!:i Vi J:!H !>w !;pt;;:
("if",;; ir;;r1y that ractjaoiol1-ir,dnct:-;;j ge',nnrnic mil)' rcsuH .im:m. (ie
",ilic'""lOrnentof tbr lH'(!C% of <:.punli.lf1('OliS and lb(,tl'foJ'e he: attrih-
lJNJ\; tt!pai1. vi.hi.d\ would the of
(,;.y the F, genome c(',uli.:l bl":
;n'Jled I;) reflHcatio:::i 'For :;mrw
(ells the ATRf/\Thlr-;.;gubtcd are tbroor:h
DNA the- :n::uhiplieity .;)1' Dt';A utr.c:;,'a:!\ms
j, det:ji"d (,f fh(, pc:i:E<:'s the F, ti should ;;luddak
1nkoown uud:::-dYing tbt ph-E::W,);1:1f;:,110n d'
,,,,b,y,,, ,,,,,w:,,,,'d! W:-}l.!i':::; :;j"H"lYlnr, that th:: o!'!\pring [ pari;sWi Hn.: gesd-
lI}' )\''''i'I,h'. f;"UY represent a 5erl{JUi'; :;i) 11H; existing: jXlradigm in radwi-
:-lcconl.ing to vinic]; fflubliml iLduC'tion in dir;::ctly ,;;xpnsl'd (:db i:,
the main of tbe D.:J;: o:f"ioGising radiation 1\)1' hUUUHi.
. I that in the cJfspd.ng uf i;:rrtdia:ed pntCUH an devEted ():f W:.Hi:l
detected Itt it m.lrnber of fil.nC%-:u:-Tntnl endpoints. indudh;g
",",,"",' " :JJH] ;;t:ne it ,vculd anv:,Hf til;:,t ;bt
irHp;dr ih:- As" i)ccmdhg tn ()ur data [61, the n-wJodt'y cor
01 ifl1lCir:l@,lnde"m,rni!,".i,tran:;b':;rn.1mC;i;'iH3;! i%tabiJ ;!y, th
" oi['i'rl'@ predicted by :he target thco,-y,
,,;h " ctw:ir ,mJ !11oxhidity On! onJ,'t
111Jllc'"qr,)p,-,,"".);J i:, Hu:cibuh:d W of .
deswb!1!;-;;it1t'41 uf th..:'; F H!;;\) ;
a :f1t.m;b;?f cf the: ,tn,,'i<knr..e of t',)nC';)f in the vltspring of '!,oJ;;ll"" lrd" 11l1,'" .,
to P1Y;;(7;fH:<fieti cardnugt;r;f; il.Hnlyi-;cD !3 i. :311,46; :5f?1 'I'he:::f
i:Hcid.en(C: (,[ ':;8.n,:t>.f anii}:l?; c_;u{jnpgtn<haHtngo:j ({'!spring It'ls,ILl',e,(l
mUt!:i, '"he data i:m sLgnihc<lnEy dC"di-?:! uf in the
radia'C\r)B Y'l"Nkem f!'C
T
,l SdJ.ati>;,,'ld nUcle;K repmc6$l:ng pbm 1).\3 (tt'C ill
resuit",
1t 5houk! he sclress;e;d thii::; .furtber ;:liniG;1! '<inpw:.:-: of mln"','"'Ofa'
in:Jl.HbiLilY lJ- cii.m::ntly becHU:',r. of Jac}.; (>'1' (ht; rdi<lb1e e,pcrilllllll.11
dilta iD, humans, SO {;E, JUs,,( ,;. of (li,s ,0'1),,,-
inA lr:,,;ll !:ur .. '{' :.;,,1wJm::Zt:d md theb rG;-sul:\Z' an:. 1:'1': fn)(!1 (x-ing <;ensbtcni.
t:tlH\lpk;, a on zh.<:: traoligeL;'nttiond e[re..;,t:3 (11' y,',,r,t.:'heriil.b,;; ,
p,:ltCrImJ an tt\r:\.'at('j fr>eqUf.':!lCY of
JJJ\::mg ;;J1iJdr('u of cxpos;,'d r 11, whereas '-;:')i,wn .n14 fiiikj
.-:hi111gel; trnong 'the ci1Jidh,lHd Ulllrei' 31irV!V<lUt:S
08 hand, sam,;;: group i}j' rrp0fte
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l'.JHHn:"cn:;ithity in children of d ch),ldhood and cm1('t':f L 121,
Gi'v'en j;hlj' the (};!;;ign of dl.{::;e !,i'Jdic;o,; ;;,'< weH 03 !hp. CGh,,)fU of )'mvJiah:-:d. fanibtt
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highly pmbkt:W:ti.c The i:n\Furtant cf thc effec,t::; (if
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References
j, A, ! g;:l1nmi:': in ebihil'::::l or i:::n<
paw!!;;; ali a if:,;lJh (if t1<;: Cr.,.;-,-::noby; Nw:.lt'_<\! A,;ducflL R;';'J 1:,,1.;)1>57
L }\8Jftb&;.:-1! C, Xi!b!:) HJ Oti'it) }k!wti>::n ii,' Ni!;"rydeJ. ;\n!m R,;\' Ch:1KI. 1 j B
r{. );';<,lk'ova '{E 1.10(6) Tjc;:, ,:::tl&1!()':'il< d !_:'r;diatd ;If::: Fk':

;{ j);;l:'h:::J' Phnnb MA, Sn.::it:1 ACT. (\:!'8f crf, Ii JdiiT!,:g AJ. :;)<!bl1
l
va YE <,:1U)\I) N(}
:')il::-!<;:'alhlt, ,;"nnhuf ml.i{d!Ori lIt 1'{:}x:1 D)ii\ ll.,;dnti{: in nm.k mice
eqU!iNI 1(, X>!i:jh u: Gi$(,,;:,:;!n,
Tn (znOs i i\:.:th:;,tipn tJ; the [''<,Ad&'",'''
1:::"nHli.;;<;::n:::;
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qUimC;-2?C::,
2i( Hr. Cl"!);;f;lar< K
nm0tw P: 11fO,'gUl) of xn,lle, Eeated

29, Ii"rd')/,cl; 'RJ. \-IV, Ihbn"'.\l 'y'E, ;20;:)')) Agf-Jd::;,;,j, iJ :cj}am:.::tbClt< uJ ,g"""",,,;
'(ik';:iillrl:)f:l!1 (lJfW.>IIUHtlJS ,;)n al
in \'ld Bid
W, Ihll).;\t T. D"r:;jd., ik\fL\, Dh;;;; PD. Vim J;:r hjrJ:;:'1,'l G\\' 1.10 E ""r-,'o',,
dki;L; ill' !im 5Dd W;;;;r::Wll p;u ii-ldinti>':<:'H,tlGth;;;d "'''''b,III' ,,;
:nil.%':, 2tA 7J(j;.,.:-.P24
:q, H<:ZYi1i, L-;!JJ 81, C, L:!)dry JU, :'-"1(:.71".', LU i T,,,,,,,,,('''','',''I',,!,;I,,,,
o.:m:im"iil"rbB 'wj,ll (:i5,:Ft'. F.:;)di,:, R,,,l" ,1><" ",i"
32. /tL" LA Tllk ul!:1rib.::::i;n:; f,f (:Hfi'.)g('w.ms D);A
;nu;;:yir; nm-i;;H>,:;)!i i;, ';;;llJ:::?f. 47':7",21
Lca DE ( 1947) AU:1'ms of nKii:llj(?l)S ,)'i! ih::mg ({,jj;;, U:;j';el':j!ty "," v" ..
>L 'y)[)L Sqp Gl" Ddl:JP m ; !97L! "' )S)""", ,,,, ":, ",I'",,,.v," '" nit> ,11"
i\:tl;,;l'! Hi'S (Vl;2)-- .'<.36
'lS. Lehnert S (?O{J7j nl<)xt"Jk:c,l!a" if iml::J:i:jg m:.}i;Jli(',':1 T,tylur & F: :<4<::\<,
]( L C '{, r ,it(:c m, l-b K, L, 5v!W. H Q
In c.am't!f cc{,; by Cnn:;d
('l., K"pbn ML. J\,V, o."\L '"',',,rc.,"'"''
i:,liWbjJity by DNA \:. , .. ,'
;B, i..{ C H. ,P:kmnhr 1. Vt:'!:.'(\;:' H , 1906,i """ ,,,,,,,,,,,,,,J',',"'lO' ,",lie""',,,
:rmf:.w!.:Bi::,\h, J (':ttl:r Phv" 93:! 5;1 fd
39 LB. \Vang i.., .i.;fC,"':;::S I), SA, P,-qV,;:Jil " D. ',""'''C'
bG, 5i;:()1J: D ).99-:<,; T:l,l'<(lur ;l;;lU:;::t;<)Jl D)' pl""""""""',"",1
41
' . wi:-t. .. ;:J({, Hi:;' j ':>.:KG i S:J01 3 i i
Lmimo:;,: SA. PJ. FG ;)O(nl gen():nk
a()llu'::g;.:!u.:l d(0(;;:;.:Jf t);} i('>:uw,'f z,ldm.t::>.:
:!2:7C:;g, 'JU.f;')
L"k;;: GiL Ridws ;\c. PE i 1997; (;"c'r:uHlJ<.; ;(::i13bdity in hilfm;lJ{);h}k(;;.:
',n mdc
:q S:Nj:ll
:dJ, :-(i,H; I1f :Jg(;!lb. MntM 2: i'!:, t " . . .,..,
,l4 ;"..'r;)f;,-{HH WF UC){FI alld (:t;l.'TCC d;'(;d;.; ol h! wrm::ll/ ',,,IUti(";:
iw.iu..;cu i:"".,;;;:>j\\;) <lTi,i 1;> -i,'hm;;:;' ,,, ':i:r::' Rc::.
4:'i Milrpn "\iF (2CC3) '<".J ,Jda.'>cd cf r;:\.{1(::"\!;'( 1.0 ""c!,"""", ji:,
i))$ub@:y ,mJ t/YhtandtY :..:1T:::::::L.; in vi' i), (;hi:-ito,gerik ",,"',','''"'!.
tm'-:;:Fficr;3:ion:;: dru:b, }Ldi.;;; R." 1 59!')
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,p PUr, S. Mult:;: S'rdkr C \ )999\ 1.:111;;) :Hfd f:fk,:,;; l.H tw;j
dmb id Ill<:d1L(] BY!W,e ancJ :'adbtinn >:'.).PG-:iUH; of ;:ygoW!;
iJl;.(.;shili!y! fk:o .. 92
Sudamc+o Sm-:uki 5 .. K, l(nn:..iflnmi R,Do}); K. Nh,:,
i';t::O::.ine mUtiHion a! '-i iJ)'}K'l'<<U'iabk \YH)!;':C min;8:mdiir-::: IOCJ.i<. Int
4V. AG (1 i ?vll)<&::i,-)Jl in mice, A'.:h' R:I.di'l( Binl ,,;.: ;3-1- len

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