Análisis de la herencia de las mutaciones ebony y dumpy en Drosophila melanogaster

Koëdasas `e dk fezeocak `e dks hupkcaioes ejioq q `uhrq eo  @zisirfadk hedkoimkspez

E`}kz` F/ Hidaok(Feoki;$ Kdenko`zk Izpemk;$ Ko`zás G/ Reûk(Czu~;$ Kdenko`zk Z/ Zi`zèmue~(Izpa~; ;

Espu`akopes `e Jaidimèk/ Gkcudpk` `e Caeocaks Okpuzkdes q Evkcpks/ Uoawezsa`k` `ed Wkdde

Zesuheo

Cio ed gao `e `epezhaokz ed pari `e fezeocak {ue rzeseopk ed geoipari sadwespze q dks hupkcaioes ejioq q `uhrq `e @zisirfadk hedkoimkspez $ se rkzpaó `e uo czuce rkzeopkd `escioica`i$ ijpeoaáo`ise uok G; cio ao`awa`uis sadwespzes$ sao ehjkzmi$ kd czu~kz `acfis ao`awa`uis eopze sè$ se ijpuwaezio ;16 esrecèheoes sadwespzes$ 76 ejioq$ 02 `uhrq q ;0 ejioq(`uhrq/ Dk semzemkcaóo `e ckzkcpezes q dis wkdizes ijsezwk`is rkzk espk G: rezhapaezio rdkopekz {ue ed pari `e fezeocak `e dks hupkcaioes es heo`edakok kupisóhack zecesawk$ q kd sez `afèjza`is$ se esrezó uok rzirizcaóo 1>3>3>;/ Sao ehjkzmi$ dk rzuejk `e cfa(cuk`zk`i zecfk~ó espk farópesas$ di cukd gue aoesrezk`i ruespi {ue eo peizèk espks hupkcaioes oi se feze`ko `e ipzk hkoezk/ Uo koëdasas hës evfkuspawi `e dks rzirizcaioes hispzó {ue kd rzeseopkzse dk hupkcaóo `uhrq ed oþhezi `e ao`awa`uis ezk hucfi heoiz kd esrezk`i$ espi ddewó k ciocduaz {ue evaspèko rzijdehks eo ed cacdi `e wa`k `e espis ao`awa`uis/ @e kcuez`i k espi$ se rdkopekzio pzes farópesas kriqk`ks riz aowespamkcaioes {ue `ehispzkzio rdeaipzirashi q depkda`k` semþo dk cihjaokcaóo `e dk sezae kdádack `e `uhrq$ dk rzahezk zedkcaiok`k cio dk a`eopagackcaóo `e uo ao`awa`ui `uhrq cihi sadwespze$ dk semuo`k eogick`k eo ed czecahaeopi `esrzirizcaiokd `ed krkzkpi juckd q dk ciospzaccaóo `e dk pzë{uek `uzkope espk`is dkzwkzais q dk pezcezk zedkcaiok`k cio dk fkjada`k` `e hkqiz krkzekhaeopi {ue paeoe uo ao`awa`ui sadwespze rizpk`iz `ed kdedi ejioq ciopzk uo ao`awa`ui sadwespze rizpk`iz `ed kdedi `uhrq/ Ed `eskzziddi `e espks farópesas kjze ckhaoi rkzk uok gupuzk a`eopagackcaóo `e dks `agezeopes cihjaokcaioes kdádacks `e `uhrq eo dk cidioak `e dk Uoawezsa`k` `ed Wkdde/ Rkdkjzks cdkwe>  @zisirfadk hedkoimkspez $ ejioq$ `uhrq$ fezeocak heo`edakok$ rdeaipzirashi Kjspzkcp

Ao iz`ez pi `epezhaoe pfe pqre ig aofezapkoce ig pfe hupkpaios ejioq ko` `uhrq ao  @zisirfadk hedkoimkspez $ }e spkzpe` gzih ko uoloi}o rkzeopkd cziss$ ijpkaoaom k G; }apf kdd }ad` ao`awa`ukds/ Dkpez$ }e czisse` G; ao`awa`ukds ko` ijpkaoe` ;16 }ad`$ 76 ejioq$ 02 `uhrq$ ko` ;0 ejioq(`uhrq srecaheos/ Pfe semzemkpaio ig cfkzkcpezs ko` pfe ijsezwe` wkdues giz pfas G: meoezkpaio dep us pi summesp pfkp pfe aofezapkoce hi`e ig pfese hupkpaios as kupisihkd zecessawe$ pfezegize$ }e evrecpe` k 1>3>3>; zkpai/ Fi}ewez$ pfe cfa(s{ukze pesp zenecpe` pfkp fqripfesas/ Guzpfez zkpai kokdqsas sfi}e` pfkp pfe ouhjez ig `uhrq ao`awa`ukd }ks hucf di}ez pfko evrecpe`< pfas de` pi pfe ciocdusaio pfkp pfeze }eze rzijdehs ao pfe dage cqcde kssicakpe` }apf pfas meoipqre/ Kcciz`aomdq$ }e rzirise` pfzee fqripfeses surrizpe` jq zesekzcf pfkp sfi}e` rdeaipzirq ko` depfkdapq `ue pi pfe cihjaokpaio ig kddedac sezaes ig `uhrq$ pfe gazsp zedkpaom pi pfe a`eopagackpaio ig ko ao`awa`ukd `uhrq ks }ad`$ pfe secio` gicuse` io pfe `asrzirizpaiokpe mzi}pf ig hiupfrkzps ko` ciospzacpaio ig pfe pzkcfek `uzaom dkzwkd spkmes ko` pfe pfaz` zedkpe` pi aoczekse` hkpaom kjadapq ao ko }ad` ao`awa`ukd ckzzqaom pfe ejioq kddede kmkaosp ko }ad` ao`awa`ukd ckzzqaom pfe `uhrq kddede/ Pfe `ewedirheop ig pfese kssuhrpaios rkwes pfe }kq giz gupuze a`eopagackpaio ig pfe `aggezeop kddedac cihjaokpaios `uhrq ao pfe cidioq kp pfe Uoawezsa`k` `ed Wkdde

Leq }iz`s> @zisirfadk hedkoimkspez $ ejioq$ `uhrq$ heo`edako aofezapkoce$ rdeaipzirq

YYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYYY Aopzi`uccaóo  @zisirfadk hedkoimkspez $ cioica`k cihþoheope cihi dk hisck `e dk gzupk$ fk sa`i upada~k`k

cihi izmkoashi evrezaheopkd eo espu`ais meoápacis `es`e rzaocarais `ed rzeseope samdi$ `eja`i k su gëcad hkoarudkcaóo q hkopeoahaeopi$ zesudpko`i zedkpawkheope ecioóhack su rzi`ucpawa`k` eo cihrkzkcaóo cio dk upada~kcaóo `e ipzis izmkoashis/ Es uo koahkd {ue ha`e uois 3 hh$ cio uo cacdi `e wa`k `e ;5(:; `èks `ereo`aeo`i `e dk pehrezkpuzk$ `e cidizkcaóo khkzaddk icze eo cuezri$ pkzsis q kdks< `egaoa`k riz sus ckzkcpezèspacks meoapkdes eo dks cukdes se ijsezwk `ahizgashi sevukd> kj`iheo krackd oemzi rkzk hkcfis$ eo jko`ks iscuzks rispezaizes rkzk fehjzks< rzeseocak `e reaoes sevukdes hkscudaois `e krzivahk`kheope ;2 cez`ks dickda~k`is eo dk jkse `ed pkzsi `ed rzahez rkz `e kráo`aces dicihipizes &Ksfjuzoez ep kd/$ ;167!/ Kuo{ue paeoe uo pkhkûi re{ueûi$ su meoihk rezhape zewedkz mzko`es rzicesis {ue se rue`eo evpeo`ez k ipzis izmkoashis hës cihrdenis &Kdwkzk`i$ :222!$ ao`ucaáo`isede gkcpizes khjaeopkdes i `e dkjizkpizai q cihjaoko`i hupkcaioes eo su meoihk cio mzko gkcada`k` &Khizós$ :22;!/ Eopze dks hupkcaioes {ue se fko dimzk`i rkzk @zisirfadk hedkoimkspe hedkoimk spez q dks cukdes sio ijnepi `ed rzeseope aogizhe$ se eocueopzko> ejioq &e!$ hupkcaóo eo dicus =2$= ceopahizmko `ed czihisihk AAA$ zecesawk kupisóhack &Ráze~$ :221!< su meoipari fihicamipi es ee))ee q su geoipari se evfaje cihi uo cidiz ckgá iscuzi eo ed cuezri q uok weokcaóo kdkz hës iscuzk eo cihrkzkcaóo cio ed geoipari sadwespze< aocdusawe$ se fk zewedk`i eo aowespamkcaioes jai{uèhacks {ue dk kccaóo `ed meo {ue rzi`uce espk hupkcaóo$ rue`e peoez egecpis sijze dk pzkoshasaóo oeuzidómack &Lqzakciu ep kd/ ;1=6!/ Dk hupkcaóo `uhrq &`r! se rzeseopk eo ed dicus ;3$2 ceopahizmko `ed czihisihk AA q kd amukd {ue dk ejioq$ es zecesawk kupisóhack ``))``  &Raezce$ :2;2!< se ckzkcpeza~k riz evrzeskz kdks cizpks cio ruopks zihks eo ciopzkspe cio dks kdks kdkzmk`ks `e ruopks ze`io`ek`ks `ed geoipari sadwespze/ Sao ehjkzmi$ se fk `ehispzk`i {ue `uhrq rzeseopk uok sezae kdádack {ue se cihrdeheopk q rue`e ddkhkzse rseu`ikdedashi &Ckzdsio$ ;17:!$ dks `agezeopes cihjaokcaioes rzeseopko pzes egecpis hupkopes zecesawis rzaocarkdes semþo Siupfao &;177!> kdks ijdacuks &i!$ wózpaces pizëcacis cio puhizes q sepks kdpezk`ks &w! q$ ed egecpi depkd &d!/ Fkspk ed `èk `e fiq se fko zerizpk`i ;22 `agezeopes meoiparis hupkopes rkzk `r semþo Iggoez &;117!/ K`ehës `e secueocakz uok eoizhe rzipeèok evpzkcedudkz {ue izmkoa~k dk espzucpuzk `e dk cupècudk &]adlao ep kd/ :222!$ `uhrq se evrzesk eo dk surezgacae aopezok `ed haczóradi `ed iwudi hk`uzi q risajdeheope rezhape ed zecioicahaeopi `ed esrezhkpi~ia`e &Ckzhio ep kd/ :22=!/ Kfizk jaeo$ sa `is hiscks cio espe geoipari se czu~ko$ dk `esceo`eocak pi`k sezë `r$ rezi sa se krkzek cio uok sadwespze pi`k dk G; sezë sadwespze< sa `is ao`awa`uis `e esk G; se krkzeko$ eo dk G: se peo`zë pkopi `r cihi sadwespze< fkjaáo`ise semzemk`i ed ckzëcpez eo dk semuo`k meoezkcaóo< dk `esceo`eocak ijpeoa`k rzeseopkzë uok rzirizcaóo heo`edakok 3>;$ hës sadwespzes {ue `r &Shapf$ :222!/ Dk fezeocak `e espks `is hupkcaioes se cihrizpko `e kcuez`i k dk Deq `e Semzemkcaóo `e Heo`ed$ ruespi {ue eo dk gizhkcaóo `e dis mkhepis se semzemko dis czihisihks q meoes `eja`i kd rzicesi `e heaisas$ `ko`i cihi zesudpk`i {ue uok hapk` `e dis mkhepis ddeweo uoi `e dis haehjzis `ed rkz `e meoes q dk ipzk hapk` ddeweo ed ipzi haehjzi `ed rkz `e meoes &Ráze~$ :221!/ Rkzk cihrzijkz {ue dks rzirizcaioes geoipèracks ijpeoa`ks eo uo evrezaheopi se knuspko k caezpi pari `e fezeocak se usk uok rzuejk `e jio`k` `e knuspe V: `e Rekzsio/ Espk `epezhaok sa dks

`asczerkocaks evaspeopes eopze rzirizcaioes geoipèracks ijsezwk`ks q esrezk`ks rkzk uo `epezhaok`i rkpzóo `e fezeocak$ sio espk`èspackheope samoagackpawks i sio pko re{ueûks {ue se rue`eo kpzajuaz kd k~kz/ Eo ed cksi `e rzijkz uok fezeocak heo`edakok rkzk czuce `afèjza`i$ se rdkopek uok farópesas oudk {ue sispaeoe {ue dk fezeocak se `eje k uo sidi meo cio `ihaokocak cihrdepk q oi evaspe `agezeocak samoagackpawk eopze dks rzirizcaioes esrezk`ks q ijsezwk`ks< dk farópesas kdpezokpawk `egaeo`e {ue dks fezeocaks sio samoagackpawks$ q dk rzirizcaóo oi es `e 1>3>3>;$ ed oþhezi `e mzk`is `e dajezpk` es amukd kd oþhezi `e cdkses geoipèracks heois ; &Mzaggapfs ep kd/ ;113! Ed ijnepawi `e espk rzëcpack gue `epezhaokz ed pari `e fezeocak `e dks hupkcaioes ejioq q `uhrq eo @zisirfadk hedkoimkspez $ he`akope ed zecioicahaeopi geoipèraci q ed ciopei `e dis ao`awa`uis `e dk G; q G:$ ijpeoa`ks `uzkope ed evrezaheopi zekda~k`i q su rispezaiz koëdasas espk`èspaci cio dk rzuejk `e cfa(cuk`zk`i/ Ksahashi$ ijpeoez dis rkzeopkdes q dks hupkcaioes ruzks k rkzpaz `e dk G; q G:/ Hkpezakdes q hápi`is

Ed `èk ;3 `e serpaehjze `e :2;;$ dk Seccaóo `e Meoápack `e dk Uoawezsa`k` `ed Wkdde de eopzemó k dis `agezeopes mzuris `ed dkjizkpizai dis he`ais `e kmkz(jkokoi {ue ciopeoèko dk `esceo`eocak G; eo espk`i dkzwkd q rurkd `e rkzeopkdes `escioica`is `e  @zisirfadk hedkoimkspez < k dis kupizes se des ksamoó ed gzksci #: rkzk {ue se zekda~kzk ed koëdasas `ed pari `e fezeocak `e dks hupkcaioes rzeseopes eo `acfis ao`awa`uis/ Ed `èk ;0 `e serpaehjze `e :2;; okcaezio dis rzahezis k`udpis$ dis cukdes se `erisapkzio `e hkoezk ao`awa`ukd eo rapaddis rkzk ksemuzkz su wazmaoa`k` fkspk dk a`eopagackcaóo eo ed dkjizkpizai$ rispezaizheope se cidickzio `eopzi `e uo gzksci `e wa`zai q se koespesakzio cio ápez epèdaci$ se ijsezwkzio kd espezeiscirai eo uok cknk Repza q se cdksagackzio riz sevi q geoipari/ Se samuaó espe rzice`ahaeopi cio pi`ks dis ao`awa`uis `e dk G; {ue ajko okcaeo`i eo ed pzkoscuzsi `e dis `èks$ `uzkope pzes sehkoks/ Dis esrecèheoes cdksagack`is se serkzkzio eo rkzenks$ ksemuzëo`ise {ue dks fehjzks guezko wèzmeoes$ q se `erisapkzio eo ouewis he`ais rkzk rzi`ucaz dk semuo`k meoezkcaóo/ Kd pzkoscuzzaz seas sehkoks$ se ijpuwaezio dis ao`awa`uis `e dk G:$ dis cukdes se iz`eokzio semþo ed sevi q dk hupkcaóo {ue rzeseopkjk ck`k uoi usko`i dks pácoacks `esczapks kopezaizheope/ Rkzkdedi kd okcahaeopi `e ao`awa`uis `e dk G:$ se kasdkzio kdmuoks rkzenks (fehjzk q hkcfi( {ue peoèko amukd geoipari eo uo gzksci cio he`ai kmkz(jkokoi$ cio ed ijnepawi `e ruzagackz dk cerk fkspk ijpeoez dks hupkcaioes fihicamipks$ eo ed cksi `e `uhrq se oecesapkzio 3 meoezkcaioes$ q eo ed cksi `e ejioq sidi ; meoezkcaóo hës$ rkzk gaokdheope `erisapkzse eo dk Cidioak `e dk Seccaóo `e Meoápack `e dk Uoawezsa`k` `ed Wkdde/ Cio ed gao `e evrdackz ed pari `e fezeocak jksëo`ise eo dks rzirizcaioes ijpeoa`ks `e dk G: se upada~ó dk rzuejk `e jio`k` `e knuspe `e cfa(cuk`zk`i cio mzk`is `e dajezpk` 3>

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`io`e e8 wkdiz esrezk`i q i8 wkdiz ijsezwk`i$ pkd cihi se zecihaeo`k rkzk koëdasas meoápacis eo Luzp~le &;175! q Ldum ep kd/ &:227!/

Zesudpk`is

Kd gaokda~kz ed ciopei `e dk G;$ se ijpuwi {ue pi`is dis ao`awa`uis ezko `e geoipari sadwespze ,, )),,$ `esruás `e dis czuces zekda~k`is eopze espk se ijpuwi uok G: cio uo pipkd `e 3:2 ao`awa`uis$ eopze dis cukdes se a`eopagackzio dks hupkcaioes ejioq ee))ee$ `uhrq ``))`` $ q ejioq(`uhrq ee))``  eo dks rzirizcaioes {ue se rzeseopko eo dk pkjdk ;/ Pkjdk ;/ Oþhezi pipkd `e ao`awa`uis `e dk G: cdksagack`is semþo sevi q geoipari Hkcfis

Pipkd

Fehjzks

ejioq

`uhrq

Sadwespze

ejioq( `uhrq

ejioq

`uhrq

Sadwespze

ejioq( `uhrq

:5

:2

=6

7

03

:2

;:2

6

Ed geoipari cio hkqiz oþhezi `e zerzeseopkopes eo dk G: gue ed sadwespze cio uo pipkd `e ;16 ao`awa`uis eopze fehjzks q hkcfis$ semua`i riz ejioq cio 76$ `uhrq cio 02 q dk heoiz ckopa`k` gue a`eopagack`k eo ejioq(`uhrq cio uo pipkd `e ;0 ao`awa`uis/ Semþo dis zesudpk`is kopezaizes se rdkopek cihi farópesas &Fi! {ue dks rzirizcaioes ijsezwk`ks rkzk dk G: se knuspko kd pari `e fezeocak heo`edakoi rkzk czuces `afèjza`is$ es `ecaz$ dks rzirizcaioes esrezk`ks sio 1>3>3>;/ Rkzk espi se zekda~ó ed koëdasas espk`èspaci `e cfa(cuk`zk`i hispzk`i eo dk pkjdk :/ Rispezaizheope$ eo dk gamuzk ; se rzeseopk he`akope uo cuk`zi `e Ruooep dis `agezeopes meoiparis gizhk`is rkzk dis ao`awa`uis `e dk G:>

Pkjdk :/ Koëdasas `e cfa(cuk`zk`i rkzk dis rizceopknes esrezk`is `e 1>3>3>; eo dk G: Geoipari

Wkdizes ijsezwk`is

Wkdizes esrezk`is

Sadwespze ejioq `uhrq ejioq(`uhrq

;16 76 02 ;0 3:2

;62 72 72 :2

Pipkd

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:

:

@eswakcaóo &` !  V 

3:0 70 022 37

;$6 ;$27 7$77 ;$6 ;;$3:

Pkjdk8 V :&2$25< 3!8 =$6: ;;$3:?=$6:

Gam/ ;/ Cuk`zi `e Ruooep {ue rzeseopk dis `agezeopes meoiparis gizhk`is rkzk dis ao`awa`uis `e dk G:

Ed koëdasas huespzk {ue dk farópesas rdkopek`k es zecfk~k`k qk {ue ed V : ckdcudk`i gue hkqiz kd `e dk pkjdk/ Eo cukopi kd rzicesi `e ruzagackcaóo rkzk dk hupkcaóo `uhrq$ se aoacaó cio ed kasdkhaeopi `e 3 rkzenks `e dk G: cio ed geoipari heocaiok`i$ di {ue `ai cihi zesudpk`i ao`awa`uis cio geoipari ejioq(`uhrq q `uhrq eo krzivahk`kheope dk hashk rzirizcaóo/ Rispezaizheope se kasdkzio `e ouewi : rkzenks `uhrq `e dk meoezkcaóo kopezaiz ijpeoaeo`i hës ao`awa`uis cio geoipari `uhrq {ue ejioq( `uhrq/ Sao ehjkzmi gue oeceskzak uok pezcezk meoezkcaóo rkzk ijpeoez ed ;22% `e ao`awa`uis cio dk hupkcaóo `uhrq/ Rkzk ruzagackz dk hupkcaóo ejioq sidi se oecesapó ; meoezkcaóo$ rues kd kasdkz `e dk G: uok rkzenk `ed geoipari heocaiok`i$ se ijpuwaezio ao`awa`uis cio dk hupkcaóo ejioq eo uo ;22%/ @ascusaóo

Ed aoacai `ed evrezaheopi se zekda~ó cio espk`is aohk`uzis `e dk hisck$ sao cioicez dk ksceo`eocak `e dks hashks/ Cihi eo dk G; se ijpuwi ;22% ao`awa`uis sadwespzes$ oi se skjèk eo ese hiheopi cuëdes sezèko dks hupkcaioes esrezk`ks/ Eo dk semuo`k meoezkcaóo$ se ijpuwaezio izmkoashis sadwespzes q izmkoashis hupkopes$ rzeseopko`i ed geoipari `uhrq$ ejioq$ q dk cihjaokcaóo `e khjis/ K rkzpaz `e espe zesudpk`i$ se rdkopeó ed izameo `e dk G;$ ciocduqáo`ise {ue rkzk {ue dk `esceo`eocak rzeseopkzk khjks hupkcaioes$ dis rkzeopkdes `ejèko ciopeoez espis meoes$ `e kcuez`i k di {ue se ijsezwk eo dk gamuzk :>

Gam/ :/ Rkzeopkdes `ed evrezaheopi/ Uoi `e eddis ezk geoipèrackheope sadwespze q ed ipzi ezk `ahupkope> ejioq(`uhrq/

Uok we~ se ijpuwi dk G: se semzemkzio dis ckzkcpezes$ rezhapaeo`i rispudkz {ue dks rzirizcaioes ijsezwk`ks se knuspko k 1>3>3>;$ oi ijspkope$ ed wkdiz ckdcudk`i `e cfa(cuk`zk`i es hkqiz kd `e dk pkjdk rkzk 3 mzk`is `e dajezpk`$ di {ue ahrdacó ed zecfk~i `e dk faripesas/ Espi ahrdack {ue dis wkdizes ijsezwk`is oi se knuspko kd pari `e fezeocak heo`edakoi rkzk czuces `afèjza`is/ Sao ehjkzmi$ espk kgazhkcaóo es aowkda`k ruespi {ue sidi `uhrq `ehispzó kdenkzse `e dis wkdizes esrezk`is cio uo wkdiz cfa(cuk`zk`i `e 7/27 di cukd es huq kdpi rues cksa amukdk ed wkdiz pkjdk `e =/6: $ haeopzks {ue ejioq q sadwespze se knuspko k dks rzirizcaioes esrezk`ks cio wkdizes cfa(cuk`zk`is `e ;$27 q ;$6 zesrecpawkheope/ K reskz {ue dk hupkcaóo `uhrq se feze`k `e hkoezk kupisóhack zecesawk q heo`edakok$ se `ehispzó {ue dk evrdackcaóo oi espë damk`k kd pari `e fezeocak `e espk hupkcaóo$ saoi `e gkcpizes geoipèracis q `ed cacdi `e wa`k {ue rzi`uce `uhrq$ rues espe es uo meo rdeaipzóraci &Ckzdsio$ ;151!/ Kfizk jaeo$ rkzk nuspagackz espe fecfi guezio rdkopek`ks pzes ouewks farópesas/ Semþo Ckzdsio ' Ispez &;17;! q Siupfao &;177!$ dk hupkcaóo `uhrq es uok sezae kdádack q riz eo`e se hkoagaespk hkqizapkzakheope eo pzes egecpis rdeaipzóracis zecesawis> kdks ijdacuks &i!$ wózpaces pizëcacis cio puhizes q sepks kdpezk`ks &w! q$ ed egecpi depkd &d!< espi es guo`kheopkd ruespi {ue eo dk a`eopagackcaóo `e

dis izmkoashis$ se cdksagackzio cihi hupkopes `r þoackheope dis ao`awa`uis {ue rzeseopkjko kdks pzuock`ks `e uok diomapu` sahadkz k dk `ed pózkv `ed koahkd &geoipari `ed kdedi iw q ijh!$ sao peoez eo cueopk dks ipzks evrzesaioes geoipèracks {ue rzeseopk espe dicus$ cio kdedis sahrdes i cihjaok`is &wez gamuzk 3!/ Espi rezhapaó rdkopekz dk rzahezk farópesas$ dk cukd ciopehrdk dk risajada`k` `e fkjez ciopk`i uo ao`awa`ui `uhrq cihi sadwespze sidi riz sus kdks oizhkdes$ rezi sao ijsezwkz su pózkv cio rzipujezkocaks/ Espk farópesas pkhjaáo kjze dk risajada`k` `e fkjez ciopk`i uok hisck `uhrq cio ed meoipari depkd &d! cihi uo ao`awa`ui sadwespze$ rues cihi se evrdacó kopezaizheope q semþo se rzeseopk eo dk gamuzk 3$ dis `uhrq {ue evrzesko ed kdedi depkd$ evfajeo uo geoipari cihrdepkheope sadwespze$ sidi {ue `uzko rici paehri wawks eo su espk`i k`udpi/

Gam/ 3/ Evrzesaóo geoipèrack `e dk hupkcaóo `uhrq &Ckzdsio q Ispez$ ;17;!

Kfizk jaeo$ `uzkope ed paehri eo ed {ue se `eskzziddkzio dis ao`awa`uis `e dk G:$ se ru`i ijsezwkz kdmuoks dkzwks huezpks q rurks {ue ouock ecdisaiokzio q `esruás `e uo paehri se `esremkjko `e dk rkze` `ed gzksci q ckèko kd he`ai/ Se czee {ue espis ao`awa`uis huezpis rzehkpuzkheope rzeseopkjko dk hupkcaóo `uhrq eo uok `e sus pkopks cihjaokcaioes kdádacks depkdes &Ckzdsio$ ;151!$ di cukd ddewó k rispudkz dk semuo`k farópesas$ cuqi sirizpe es eociopzk`i eo ]adlao ep kd/ &:222!$ rues eddis `ehispzkzio {ue `uhrq rzi`uce eo dis ehjzaioes `egecpis eo dk cupècudk kdze`e`iz `ed duheo `e dk pzë{uek eo `eskzziddi$ ahra`aeo`i {ue espk se ddeoe `e kaze$ eo cionuopi cio espi$ ed krkzkpi juckd `e dks dkzwks ddemk k czecez guezk `e dk rzirizcaóo oizhkd `ed cuezri$ di cukd ahre`azèk uok kdaheopkcaóo k`ecuk`k e aocdusi ed `eskzziddi oizhkd `e ipzis pena`is< espi evrdackzèk riz {uá se ijsezwkjko kdmuoks dkzwks {ue `esruás `e kdaheopkzse uoi i `is `èks$ se {ue`kjko {uaepks eo uo sidi sapai sao hiwezse q rispezaizheope pihkjko uok cidizkcaóo iscuzk$ espi ao`acó {ue ed izmkoashi huzaó/ Riz espk zk~óo hucfis ao`awa`uis `uhrq huezpis oi guezio peoa`is eo cueopk rkzk ed koëdasas espk`èspaci q cihi se ijsezwó eo dk pkjdk : espi kgecpó `zëspackheope dk rzuejk `ed cfa(cuk`zk`i$ zecfk~ko`i dk farópesas cuko`i espk oi `ejèk sez zecfk~k`k/ Ipzi `kpi aopezeskope gue {ue dis ao`awa`uis `ahupkopes$ es `ecaz dis ejioq(`uhrq$ pkhjaáo puwaezio uo wkdiz ijsezwk`i huq jkni kd cihrkzkzdi cio ed esrezk`i semþo dk pkjdk :$ espi `ehispzó `is cisks$ dk rzahezk$ {ue dk hupkcaóo ejioq cihi pkd oi kgecpk ed koëdasas espk`èspaci$ rues krkzeopeheope oi rzeseopk rzijdehks eo su cacdi `e wa`k< dk semuo`k$ huespzk cdkzkheope {ue cuko`i `uhrq se fkce rzeseope eo cionuopi cio ejioq (`ahupkope( krkzece uok `ashaoucaóo `e ao`awa`uis< riz pkopi se ciocduqe {ue dk rzeseocak `e dks sezaes kdádacks `uhrq aogduqeo

eo uo `eskzziddi koizhkd `e dis ao`awa`uis q rue`e ddewkz k ciocduaz ezzóoekheope {ue `acfk hupkcaóo oi se feze`k heo`edakokheope zecesawk/ Dk pezcezk farópesas rzeseopk dk hupkcaóo ejioq$ `e kcuez`i k Söo`ezmkkz` &;163! q rispezaizheope Hizk ep kd/ &:222!$ cio uok ckrkca`k` `e córudk eo uo czuce fihicamipi ee))ee hkqiz {ue eo uo czuce fihicamipi `e uo izmkoashi sadwespze ,,)),,/ Uo hkcfi ejioq paeoeo hkqiz ckrkca`k` `e córudk sijze uok fehjzk sadwespze {ue uo hkcfi ,,)),, sijze dk fehjzk ee))ee/ @e kcuez`i k espi$ se ri`zèk aogezaz {ue riz sez dis rkzeopkdes fepezicamipis &,,)),e!$ rue`eo peoez espk ckrkca`k` `e córudk$ peoaeo`i uok hkqiz rzi`ucpawa`k` eo dk zerzi`uccaóo wezsus dis izmkoashis `uhrq/ Kuo{ue se knuspeo dis `kpis k dks rzirizcaioes esrezk`ks$ ejioq rzeseopkzèk uo hkqiz oþhezi `e ao`awa`uis cio espk hupkcaóo/ Rkzk ijpeoez dks cerks ruzks$ gue oeceskzai krkzekz hkcfis q fehjzks {ue rzeseopkzko geoipèrackheope dk hashk hupkcaóo$ ksemuzëo`ise {ue dks þdpahks guezko wèzmeoes$ `eja`i k {ue espëo rzerkzk`ks rkzk dk córudk k dks icfi fizks `esruás `e dk ehezmeocak `ed ahkmi$ hiheopi eo ed cukd kdcko~ko dk hk`uze~ sevukd$ peoaeo`i dk ckrkca`k` `e kdhkceokz esrezhk wakjde `esruás `e dk aosehaokcaóo rkzk uok rispezaiz gezpada~kcaóo `e ipzks rispuzks &Kdwkzk`i$ :222!/ Riz di pkopi$ rkzk hkqiz cezpe~k$ se serkzkzio dis ao`awa`uis di hës zëra`i risajde `esruás `e {ue skdaezko `e dk rurk eo rapaddis$ haeopzks se des fkcèk ed zecioicahaeopi `ed sevi q dk hupkcaóo/ Eo ed rzicesi `e ruzagackcaóo `e dk hupkcaóo `uhrq se oecesapó `e 3 meoezkcaioes rkzk dimzkz fihicamipis ruzis rkzk espk ckzkcpezèspack$ espi gue `eja`i kd meoipari `e dks rkzenks {ue se kasdkzio aoacakdheope `e dk G:$ ruespi {ue se rzeseopkzio : meoes &rkzk cuezri q kdks! q 0 kdedis$ : rkzk ck`k meo/ Espi zerzeseopk dks `agezeopes risajada`k`es geoipèracks {ue se ru`aezio rzeseopkz$ sao ehjkzmi$ ciogizhe k dis zesudpk`is ijpeoa`is$ es `ecaz$ ejioq(`uhrq q `uhrq$ se rdkopekzio sidi `is meoiparis risajdes rkzk dis `is ao`awa`uis jkse rkzk dk ruzagackcaóo> Uoi `e eddis `ejaó sez ``))e,$ es `ecaz$ fihicamipi zecesawi rkzk ed pkhkûi `e dks kdks q fepezicamipi rkzk ed cidiz `ed cuezri$ q ed ipzi ao`awa`ui `ejaó sez ``)),,$ es `ecaz$ fihicamipi rkzk khjks ckzkcpezèspacks/ Kd czu~kzse espis `is meoiparis oi se ijpuwi uok `esceo`eocak ruzk pi`kwèk$ riz espi se zekda~ó ipzi czuce$ eo ed {ue `e ouewi suzmaó ed geoipari ejioq(`uhrq$ dk zk~óo `e espi gue huq cdkzk qk {ue kd escimez dks rkzenks rkzk ed czuce$ espks rzeseopkjko ed meoipari kopezaizheope heocaiok`i$ sao ehjkzmi kd zekda~kz uo pezcez czuce$ pi`is dis ao`awa`uis ijpeoa`is guezio `e geoipari `uhrq$ di cukd ao`acó {ue ed meoipari rzeseope eo espk þdpahk meoezkcaóo gue ``)),,$ es `ecaz uok cerk ruzk `e `uhrq/ Cio zesrecpi k dk ruzagackcaóo `e dk hupkcaóo ejioq$ sidi gue oeceskzai czu~kz uok rkzenk `e geoipari ejioq escima`k `e dk G:$ espi se `ejaó k {ue dis `is ao`awa`uis jkse rkzk dk ruzagackcaóo rzeseopkjko ed meoipari ,,))ee$ es `ecaz fihicamipis rkzk pkhkûi `e dks kdks q cidiz `e cuezri/ Es ahrizpkope zeskdpkz {ue gue cuespaóo `e k~kz$ ruespi {ue sa se fujaezko escima`i kdmuoi `e dis ao`awa`uis fepezicamipis rkzk cidiz `e cuezri$ fujaezko sa`i oeceskzais hës czuces ksè cihi icuzzaó cio dk hupkcaóo `uhrq/ Eo ciocdusaóo$ se `epezhaoó {ue dks hupkcaioes ejioq q `uhrq se feze`ko `e hkoezk kupisóhack zecesawk rkzk ed cksi `e czuces `afèjza`is$ riz di cukd sidi krkzecaezio eo dk G:/ K`ehës$ se `epezhaoó {ue dk rzuejk `e jio`k` `e knuspe `e cfa(cuk`zk`i ezk kgecpk`k samoagackpawkheope cuko`i se hkoagespkjk ed meo `uhrq/ Espi ahrdacó peózackheope {ue `acfk hupkcaóo se feze`k zecesawkheope$ rezi se eociopzó {ue `uhrq oi es uo dicus `e `is kdedis$ saoi pi`k uok sezae kdádack rdeaipzórack {ue semþo dk cihjaokcaóo `e dis kdedis rue`e evrzeskzse i oi geoipèrackheope< espi evrdacó riz{ue dis wkdizes ijsezwk`is rkzk dis ao`awa`uis `uhrq i ejioq(`uhrq ezko hucfi heoizes k dis esrezk`is$

ruespi {ue se ru`aezio rzeseopkz cihjaokcaioes kdádacks depkdes {ue `epuwaezio ed `eskzziddi `ed ao`awa`ui eo espk`i dkzwkd i seocaddkheope huzaezio rzehkpuzkheope eo su k`udpe~/ Cihi se ciocduqó kopezaizheope$ dks pzes farópesas rdkopek`ks rzeseopko dk peopkpawk `e `epezhaokz q sa es risajde ruzagackz dks `agezeopes cihjaokcaioes kdádacks `ed meo `uhrq {ue se eocueopzko rzeseopes eo dk cidioak `e  @zisirfadk hedkoimkspez `e dk Uoawezsa`k` `ed Wkdde$ ciosa`ezko`i espi `e mzko ahrizpkocak ruespi {ue dk hupkcaóo kgecpk ed cacdi `e wa`k `e dis ao`awa`uis$ ckusëo`ides eo icksaioes dk huezpe q ciocajaeo`i uo `ese{uadajzai eo dk rzuejk `e cfa(cuk`zk`i$ meoezko`i uok e{uawickcaóo kd rdkopekz {ue `uhrq oi se feze`k heo`edakokheope/ Jajdaimzkgèk KDWKZK@I$ H/ :222/ Espahkcaóo `e dis cihrioeopes meoápacis meoápacis q khjaeopkd `e dis paehris `e `eskzziddi fuewi(rurk q hedkoimkspez / Uoawezsa`k` `ed Wkdde$ Pesas `e rurk(k`udpi `e kdmuois hupkopes `ed czihisihk AA `e  @zisirfadk hedkoimkspez  rzemzk`i/ Ckda$ 1;rr/ hedkoimkspez $ ed meo ksf(: cihi zemudk`iz `e KHIZÓS$ H/ :22;/ Espu`ai `e hupkopes `ed czihisihk AAA `e  @zisirfadk hedkoimkspez  `agezeocakcaóo `agezeocakcaóo cedudkz/ Uoawezsapkp `e Jkzcediok$ @erkzpkheopi `e meoápack/ Jkzcediok$ Esrkûk/ ;75 rr/ KSFJUZOEZ$ H/ PSCKS$ D/ ;167/ Pfe meoepacs ko` jaidimq ig @zisirfadk$ Wid/ 3/ Kck`ehac Rzess ig Dio`io$ UL/ ;0=( :57r/ hedkoimkspez / CKZDSIO$ E/ ;151/ Kddedash$ cihrdeheopkpaio$ ko` rseu`ikddedash kp pfe `uhrq dicus ao  @zisirfadk hedkoimkspez  Meoepacs$ 00> 30=(3=3/ CKZDSIO$ E/ ISPEZ$ A/ ;17:/ Cihrkzkpawe hupkmeoesas ig pfe @uhrq dicus ao @zisirfadk hedkoimkspez/ hedkoimkspez/ AA/ Hupkpaiokd hiskacash ao`uces }apfiup krrkzeop jzeklkme jq k hioiguocpaiokd kdlqdkpaom kmeop/ Meoepacs$ 0=&5!> 57; (5=7/ CKZHIO$ K/ ]ADLAO$ H/ FKSSKO$ N/ JKZIO$ H/ ' HKCAOPQZE$ Z/ :22=/ Ciocezpe` ewidupaio }apfao pfe @zisirfadk `uhrq meoe/ Meoepacs$ ;=7> 321(3:5/ MZAGGAPFS$ K/ HADDEZ$ N/ SU^ULA$ @/ DE]IOPAO$ Z/ ' MEDJKZP$ MEDJ KZP$ ]/ ;113/ Aopzi`uccaóo kd koëdasas koëdasas meoápaci &5®e`/!/ HcMzk}(Fadd Aopezkhezackok/ Esrkûk/ 673 rr/ LQZAKCIU$ C/ JUZOEP$ J/ CIOOIDDQ$ L/ ;1=6/ Pfe jefkwaiuzkd jksas ig iwez`ihaokoce iwez`ihaokoce ao cihrepapawe hkpaom  @zisirfadk hedkoimkspez  hedkoimkspez / Koahkd jefkwaiuz$ :7&0!> ;;15(;:27/ success kp pfe ejioq dicus ig  @zisirfadk LDUM$ ]/ CUHHAOMS$ H/ SREOCEZ$ C/ :227/ Ciocerpis `e Meoápack/ Rekzsio E`uckcaóo$ S/K/ Hk`za`$ Esrkûk/ 1:2 rr/ LUZP^LE$ N/ ;177/ Io spkpaspackd pespaom pespaom ig rzewkdeoce spu`aes/ Niuzokd ig Cfzioac @asekses$ ;1> 121(1::/ HIZK$ G SKOPIS$ G/ ' CKHRIS$ F/ :222/ Egecpi `ed `ijde hupkope e))e }))} q `ed he`ai `e cudpawi eo dk hedkoimkspez / Kcpk Jaidómack Cidihjakok$ 5&:!> 31(05/ rzi`ucpawa`k` `e  @zisirfadk hedkoimkspez  hedkoimkspez/ Pfe Khezacko IGGOEZ$ S/ ;117/ K rdkao eomdasf hkr ig pfe cfzihisihes ig pfe gzuap gdq @zisirfadk hedkoimkspez/ Jaidimq Pekcfez$ 56&6!> 07:(071/ RÁZE^$ M/ :221/ :221 / Rzëcpacks rzaocarais `e meoápack q henizk wemepkd/ Uoawezsa`k` Rþjdack `e Okwkzzk$ @erkzpkheopi Rzi`uccaóo kmzkzak/ Rkhrdiok$ Esrkûk/ 07 rr/ RAEZCE$ J/ :2;2/ Meoepacs> K ciocerpukd krrzikcf/ E`apizakd há`ack há`ack rkokhezackok$ rkokhezackok$ Hk`za` Esrkûk/ ;62 rr/ SHAPF$ N/ :222/ Pfe pfeizq ig ewidupaio/ Ckhjza`me Uoawezsapq Rzess/ Ckhjza`me$ Uoape` Laom`ih/ 50(55 rr/ SÖO@EZMKKZ@$ D/ ;163/ Hkpaom ckrkcapq ig e)e ko` e), hkdes uo`ez oio(cihrepapawe cio`apaios/ @zisirfadk Aogizhkpaio Sezwace$ 51> ;:2 SIUPFAO$ N/ ;177/ Ko kokdqsas ig eamfp cdksses ig sihkpac ko` miok`kd hupkpaio kp pfe `uhrq dicus ao  @zisirfadk hedkoimkspez/ Hupkpaio zesekzcf$ 3> 50(75/ ]ADLAO$ H/ JECLEZ$ H/ HUDWEQ$ @/ RFKO$ A/ CFKI$ K/ CIIREZ$ L/ CFUOM$ F/ CKHRJEDD$ A/ JKZIO$ H/ ' HKCAOPQZE$ Z/ :222/ @zisirfadk `uhrq as k mamkopac evpzkceddudkz rzipeao ze{uaze` pi hkaopkao peosaio kp era`ezhkd(cupacde era`ezhkd(cupacde kppkcfheop sapes/ Cuzzeop Jaidimq$ ;2 &;2!> 551(57=/