11
Xk`a~ Ozzg F`b`x HAI ?:"?::6!}::?>>::9?:090  ?>= A^IFIBOG ZOZ@^ ]ibfg` bpmg`axih` zagyea~zki}e f`baxyzibf ib zagyzgaih vk`ox vixk xk` Iggpeibo Fagh`bFox` o}}oy @hpo~h Olkpbas ¶ Mko~g`} Bimag`x ¶ Dob Hsa~ol ^`m`is`h4 ?; Oz~ig >::; ! Omm`zx`h4 >8 Oz~ig >::9 © Xk` Opxka~+}) >::9" Xki} o~ximg` i} zpngi}k`h vixk az`b omm`}} ox ]z~ibf`~gibl"mae On}x~omx ]ibfg` bpmg`axih` zagyea~zki}e} +]BZ}) o~` ibhi}z`b}ong` ib }pmk ozzgimoxiab} o} o}}amioxiab eozzibf obh mab}x~pmxiab ac kifkh`b}ixy f`b`xim eoz}" X k`}` ozzgi moxiab} p}poggy ~`qpi~` f`baxyzibf ac xkap}obh} ac ]BZ} ib o go~f` bpen`~ ac ibhisihpog}" Ogxkapfk o bpen`~ ac ]BZ f`baxyzibf o}}oy} o~` osoigong`' ea}x ac xk`e o~` h`}ifb`h ca~ ]BZ f`baxyzibf ib hizgaih ibhisihpog}" K`~`' v` h`eab}x~ox` xkox xk` Iggpeibo Fagh`bFox` o}}oy mapgh n` p}`h ca~ ]BZ f`baxyzibf ac kaeatyfap} x`x~ozgaih obh k`{o zgaih vk`ox gib`}" F`baxyzibf ~`omxiab} mapgh n` mo~~i`h apx hi~`mxgy ab f`baeim HBO vixkapx xk` b`m`}}ixy ac z~` gieibo~y ZM^ oezgiVmoxiab" O xaxog ac 0= x`x~ozgaih obh =; k`{ozgaih kaeatyfap} vk`ox gib`} v`~` f`baxyz`h ox 93 ]BZ gami" Xk` f`baxyzibf `~~a~ ~ox` `}xieox`h ocx`~ ~ `easog ac gavqpogixy hoxo vo} : obh ?% ca~ x`x~ozgaih obh k`{o zgaih vk`ox' ~`}z`mxis`gy" H`s`gaz`h ]BZ f`baxyzibf o}}oy} v`~` }kavb xa n` p}`cpg ca~ f`baxyzibf vk`ox mpgxiso~}" Xki} }xphy h`eab}x~ox`h xkox xk` Fagh`bFox` o}}oy i} o s`~y `Ymi`bx xaag ca~ kifkxk~apfkzpx f`baxyzibf ac zagy zgaih vk`ox' az`bibf b`v za}}inigixi`} ca~ xk` obogy}i} ac f`b`xim so~ioxiab ib vk`ox obh hi}}`mxiab ac f`b`xim no}i} ac maezg`{ x~oix} p}ibf o}}amioxiab eozzibf ozz~aomk" Ibx~ahpmxiab ]ibfg` bpmg`axih` zagyea~zki}e +]BZ) i} o pniqpixap} xyz` ac f`b`xim so~ioxiab ib `plo~yaxim f`bae`}" ]BZ} o~` ih` oggy }pix`h ca~ xk` mab}x~pmxiab ac kifk~`}agpxiab f`b`xim eoz}' ibs`}xifoxiab ac zazpgoxiab `sagpxiabo~y ki}xa~y obh hi}mas`~y ac eo~l`~”x~oix o}}amioxiab} ib o}}amioxiab eoz zibf `{z`~ie`bx} +Ibx`~boxiabog KozEoz Mab}a~xipe >::6< Tko a ` x og" >::6< O~obtobo `x og" >::0)" Ib o}}amioxiab eoz zibf' o h`b}` }`x ac ]BZ eo~l`~} mas`~ibf xk` `bxi~` f`bae` i} b``h`h ca~ Vbhibf o mo}pog epxoxiab a~ ob ]BZ xkox i} ib giblof` hi}`qpigin~ipe vixk xk` mo}pog epxoxiab +O~ob tobo `x og" >::0< Cgibx Fo~mio `x og" >::0)" Ibmgp}iab ac o go~f` bpen`~ ac ibhisihpog} ib ob o}}amioxiab eozzibf }xphy i} m~iximog xa kos` }pYmi`bx zav`~ ac o}}amioxiab x`}x +O~obtobo `x og" >::0< Cgibx Fo~mio `x og" >::0)" Mab}x~pm xiab ac kifkh`b}ixy f`b`xim eoz} ~`qpi~`} f`baxyzibf go~f` z~af`by ox o go~f` bpen`~ ac gami +X~affia `x og" >::6< Ibx`~boxiabog KozEoz Mab}a~xipe >::6)" Xk`}` ozzgimo xiab}' xk`~`ca~`' ~`qpi~` f`baxyzibf zgoxca~e} mozong` ac z~ahpmibf epgxigamp} f`baxyz`} ib o go~f` zob`g ac ibhi sihpog}" O bpen`~ ac f`baxyzibf x`mkbagafi`} o~` mp~~`bxgy osoigong` xkox }oxi}cy xk`}` ~`qpi~`e`bx}" Xk` ~`m`bxgy h`s`gaz`h f`baxyzibf zgoxca~e Iggpeibo N`ohO~~oy mae nib`h vixk xk` Fagh`bFox` o}}oy i} ong` xa f`baxyz` pz xa ?'0=3 zagyea~zkim }ix`} ib =;8 ibhisihpog} ib o }ibfg` ~`om xiab +Agizko bx `x og" >::>)" Eag`mpgo~ Ibs`~}iab Z~an` +EIZ) x`mkbagafy aS `~} ?>':::zg`{ ]BZ f`baxyzibf o}}oy Maeepbimox`h ny O" ]mkpgeob" @g`mx~abim }pzzg`e`bxo~y eox`~iog Xk` abgib` s`~}iab ac xki} o~ximg` +hai4?:"?::6!}::?>>::9?:090 ) mabxoib} }pzzg`e`bxo~y eox`~iog' vkimk i} osoigong` xa opxka~it`h p}`~}" @" Olkpbas ¶ D" Hsa~ol H`zo~xe`bx ac Zgobx ]mi`bm`}' Pbis`~}ixy ac Mogica~bio' Ab` ]ki`gh} Os`"' Hosi}' MO 903?3' P]O M" Bimag`x F`bae` obh Niae`himog ]mi`bm`} Comigixy' HBO X`mkbagafi`} Ma~` Comigixy' Pbis`~}ixy ac Mogica~bio' 80? K`ogxk ]mi`bm`} H~is`' Hosi}' MO 903?3' P]O Z~`}`bx Ohh~`}}4 @" Olkpbas +#) H`zo~xe`bx ac Zgobx Zoxkagafy' Xk~amlea~xab Zgobx ]mi`bm`} M`bx`~' Lob}o} ]xox` Pbis`~}ixy' Eobkoxxob' L] 330:3' P]O `eoig4 `olkpbasJl}p"`hp

Single Nucleotide Polymorphism Genotyping in Polyploid Wheat

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Page 1: Single Nucleotide Polymorphism Genotyping in Polyploid Wheat

8/14/2019 Single Nucleotide Polymorphism Genotyping in Polyploid Wheat

http://slidepdf.com/reader/full/single-nucleotide-polymorphism-genotyping-in-polyploid-wheat 1/11

Xk`a~ Ozzg F`b`x

HAI ?:"?::6!}::?>>::9?:090

 ?>=

A^IFIBOG ZOZ@^

]ibfg` bpmg`axih` zagyea~zki}e f`baxyzibf ib zagyzgaih vk`ox

vixk xk` Iggpeibo Fagh`bFox` o}}oy

@hpo~h Olkpbas ¶ Mko~g`} Bimag`x ¶ Dob Hsa~ol

^`m`is`h4 ?; Oz~ig >::; ! Omm`zx`h4 >8 Oz~ig >::9

© Xk` Opxka~+}) >::9" Xki} o~ximg` i} zpngi}k`h vixk az`b omm`}} ox ]z~ibf`~gibl"mae

On}x~omx ]ibfg` bpmg`axih` zagyea~zki}e} +]BZ}) o~`

ibhi}z`b}ong` ib }pmk ozzgimoxiab} o} o}}amioxiab eozzibfobh mab}x~pmxiab ac kifkh`b}ixy f`b`xim eoz}" Xk`}` ozzgi

moxiab} p}poggy ~`qpi~` f`baxyzibf ac xkap}obh} ac ]BZ} ib

o go~f` bpen`~ ac ibhisihpog}" Ogxkapfk o bpen`~ ac ]BZ

f`baxyzibf o}}oy} o~` osoigong`' ea}x ac xk`e o~` h`}ifb`h

ca~ ]BZ f`baxyzibf ib hizgaih ibhisihpog}" K`~`' v`

h`eab}x~ox` xkox xk` Iggpeibo Fagh`bFox` o}}oy mapgh n` p}`h

ca~ ]BZ f`baxyzibf ac kaeatyfap} x`x~ozgaih obh k`{o

zgaih vk`ox gib`}" F`baxyzibf ~`omxiab} mapgh n` mo~~i`h

apx hi~`mxgy ab f`baeim HBO vixkapx xk` b`m`}}ixy ac z~`

gieibo~y ZM^ oezgiVmoxiab" O xaxog ac 0= x`x~ozgaih obh =;

k`{ozgaih kaeatyfap} vk`ox gib`} v`~` f`baxyz`h ox 93

]BZ gami" Xk` f`baxyzibf ̀ ~~a~ ~ox` ̀ }xieox`h ocx`~ ~`easog

ac gavqpogixy hoxo vo} : obh ?% ca~ x`x~ozgaih obh k`{o

zgaih vk`ox' ~`}z`mxis`gy" H`s`gaz`h ]BZ f`baxyzibf

o}}oy} v`~` }kavb xa n` p}`cpg ca~ f`baxyzibf vk`ox mpgxiso~}"

Xki} }xphy h`eab}x~ox`h xkox xk` Fagh`bFox` o}}oy i} o

s`~y `Ymi`bx xaag ca~ kifkxk~apfkzpx f`baxyzibf ac zagyzgaih vk`ox' az`bibf b`v za}}inigixi`} ca~ xk` obogy}i} ac 

f`b`xim so~ioxiab ib vk`ox obh hi}}`mxiab ac f`b`xim no}i} ac 

maezg`{ x~oix} p}ibf o}}amioxiab eozzibf ozz~aomk"

Ibx~ahpmxiab

]ibfg` bpmg`axih` zagyea~zki}e +]BZ) i} o pniqpixap} xyz`

ac f`b`xim so~ioxiab ib `plo~yaxim f`bae`}" ]BZ} o~` ih`

oggy }pix`h ca~ xk` mab}x~pmxiab ac kifk~`}agpxiab f`b`xim

eoz}' ibs`}xifoxiab ac zazpgoxiab `sagpxiabo~y ki}xa~y obh

hi}mas`~y ac eo~l`~”x~oix o}}amioxiab} ib o}}amioxiab eoz

zibf `{z`~ie`bx} +Ibx`~boxiabog KozEoz Mab}a~xipe >::6<

Tkoa x̀ og" >::6< O~obtobo ̀ x og" >::0)" Ib o}}amioxiab eoz

zibf' o h`b}` }`x ac ]BZ eo~l`~} mas`~ibf xk` `bxi~`

f`bae` i} b``h`h ca~ Vbhibf o mo}pog epxoxiab a~ ob ]BZ

xkox i} ib giblof` hi}`qpigin~ipe vixk xk` mo}pog epxoxiab

+O~obtobo `x og" >::0< CgibxFo~mio `x og" >::0)" Ibmgp}iab

ac o go~f` bpen`~ ac ibhisihpog} ib ob o}}amioxiab eozzibf

}xphy i} m~iximog xa kos` }pYmi`bx zav`~ ac o}}amioxiab x`}x

+O~obtobo `x og" >::0< CgibxFo~mio `x og" >::0)" Mab}x~pm

xiab ac kifkh`b}ixy f`b`xim eoz} ~`qpi~`} f`baxyzibf go~f`

z~af`by ox o go~f` bpen`~ ac gami +X~affia `x og" >::6<

Ibx`~boxiabog KozEoz Mab}a~xipe >::6)" Xk`}` ozzgimo

xiab}' xk`~`ca~`' ~`qpi~` f`baxyzibf zgoxca~e} mozong` ac 

z~ahpmibf epgxigamp} f`baxyz`} ib o go~f` zob`g ac ibhi

sihpog}" O bpen`~ ac f`baxyzibf x`mkbagafi`} o~` mp~~`bxgy

osoigong` xkox }oxi}cy xk`}` ~`qpi~`e`bx}" Xk` ~`m`bxgy

h`s`gaz`h f`baxyzibf zgoxca~e Iggpeibo N`ohO~~oy mae

nib`h vixk xk` Fagh`bFox` o}}oy i} ong` xa f`baxyz` pz xa

?'0=3 zagyea~zkim }ix`} ib =;8 ibhisihpog} ib o }ibfg` ~`om

xiab +Agizkobx `x og" >::>)" Eag`mpgo~ Ibs`~}iab Z~an`

+EIZ) x`mkbagafy aS ̀ ~} ?>':::zg`{ ]BZ f`baxyzibf o}}oy

Maeepbimox`h ny O" ]mkpgeob"

@g`mx~abim }pzzg`e`bxo~y eox`~iog Xk` abgib` s`~}iab ac xki}

o~ximg` +hai4?:"?::6!}::?>>::9?:090) mabxoib} }pzzg`e`bxo~y

eox`~iog' vkimk i} osoigong` xa opxka~it`h p}`~}"

@" Olkpbas ¶ D" Hsa~ol 

H`zo~xe`bx ac Zgobx ]mi`bm`}' Pbis`~}ixy ac Mogica~bio'

Ab` ]ki`gh} Os`"' Hosi}' MO 903?3' P]O

M" Bimag`x

F`bae` obh Niae`himog ]mi`bm`} Comigixy'

HBO X`mkbagafi`} Ma~` Comigixy' Pbis`~}ixy ac Mogica~bio'

80? K`ogxk ]mi`bm`} H~is`' Hosi}' MO 903?3' P]O

Z~`}`bx Ohh~`}}4

@" Olkpbas +#)

H`zo~xe`bx ac Zgobx Zoxkagafy'

Xk~amlea~xab Zgobx ]mi`bm`} M`bx`~'

Lob}o} ]xox` Pbis`~}ixy' Eobkoxxob' L] 330:3' P]O

`eoig4 `olkpbasJl}p"`hp

Page 2: Single Nucleotide Polymorphism Genotyping in Polyploid Wheat

8/14/2019 Single Nucleotide Polymorphism Genotyping in Polyploid Wheat

http://slidepdf.com/reader/full/single-nucleotide-polymorphism-genotyping-in-polyploid-wheat 2/11

Xk`a~ Ozzg F`b`x

 ?>=

ca~ f`baxyzibf ac o gieix`h bpen`~ ac ibhisihpog} +Ko~h`b

nag `x og" >::0)" Ob az`b}ap~m`' e`hipe xk~apfkzpx ]BZ

f`baxyzibf zgoxca~e ko} ~`m`bxgy n``b h`s`gaz`h ca~ f`ba

xyzibf xkap}obh} ac ibhisihpog} ox x`b} xa kpbh~`h} ac ]BZ

gami +Eomhabogh `x og" >::0)" Ogg xk`}` x`mkbagafi`} o~`

no}`h ab agifa gifoxiab o}}oy +AGO) +Agizkobx `x og" >::>)"

Xk` Iggpeibo Fagh`bFox` o}}oy pxigit`} xva ogg`g`}z`

miVm agifabpmg`axih`} +O]A}) xa hi}m~ieibox` n`xv``b xk`ogg`gim }xox`} ox ob ]BZ gamp} +Cob `x og" >::=)" Xk` hi}m~ie

iboxibf bpmg`axih`} o~` gamox`h ox xk` = `bh} ac xk` zoi~ ac 

O]A}" Xk` xki~h z~ie`~' ~`c`~~`h xa o} xk` gamp} }z`miVm agi

fabpmg`axih` +G]A)' obb`og} havb}x~`oe ac O]A" Xki}

z~ie`~ mabxoib} ob ohh~`}} }`qp`bm` +IggpeiMah`) pbiqp`

ca~ `omk ]BZ }ix`" Ocx`~ obb`ogibf' xk` O]A vixk o =`bh

no}` maezg`e`bxo~y xa xk` no}` ox xk` ]BZ }ix` i} `{x`bh`h

ny zagye`~o}` obh gifox`h xa xk` ma~~`}zabhibf G]A ca~e

ibf o ZM^ x`ezgox`" Xk` `{x`b}iab obh gifoxiab ~`omxiab i}

z`~ca~e`h ox epgxizg` +pz xa ?'0=3) ]BZ gami }iepgxo

b`ap}gy" Xk` x`ezgox`} o~` xk`b ZM^ oezgiV`h p}ibf o }`x

ac xk~`` z~ie`~} maeeab xa ogg ]BZ }ix`}" Xva ac xk`e o~`

gon`g`h vixk `ixk`~ My= a~ My0 [pa~`}m`bx hy` obh `omk

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obb`og} xa o maeeab }`qp`bm` ib xk` G]A}" ZM^ z~ahpmx}

o~` obb`og`h xa ob o~~oy maeza}`h ac n`oh} mo~~yibf agifa

bpmg`axih`} maezg`e`bxo~y xa xk` IggpeiMah` }`qp`bm`} ib

xk` G]A} xkox o~` p}`h ca~ ~`mafbixiab ac `omk ]BZ }ix`"

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p}`h xa h`x`~eib` xk` ogg`gim }xox` ox ob ]BZ }ix`" @qpog

ibx`b}ixy ac My=!My0 }ifbog +~oxia ?4?) ma~~`}zabh} xa xk`

k`x`~atyfap} }xox`< ?4: a~ :4? }ifbog ~oxia} ma~~`}zabh xa

ab` ac xk` ogx`~boxis` kaeatyfap} }xox`} ox ob ]BZ }ix`"

Xk` omxpog f`baxyz` moggibf i} z`~ca~e`h ocx`~ mgp}x`~ibf ac 

[pa~`}m`bm` ibx`b}ixy sogp`} xolibf ibxa ommapbx so~ioxiab

ib hoxo magg`mx`h c~ae ibhisihpog n`oh} obh os`~ofibf obh

~`easibf apxgi`~}" Pz xa xk~`` f`baxyz` mgp}x`~} mapgh n`

anxoib`h ca~ `omk ]BZ }ix` ib o hizgaih a~fobi}e"

Ea}x ac xk` kifkxk~apfkzpx ]BZ f`baxyzibf zgoxca~e}

obh xk` ]BZ hoxo obogy}i} ogfa~ixke} v`~` h`s`gaz`h ca~

f`b`~oxibf obh obogytibf hizgaih f`baxyz` hoxo" F`baxyz

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}z`mi`}' i} maezgimox`h ny xk` z~`}`bm` ac xva a~ ea~` f`b`

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p}ibf z~ie`~} }z`miVm xa ab` a~ obaxk`~ mazy ac xk` hpzgi

mox`h gamp} +Moghv`gg `x og" >::8)" Kav`s`~' xki} ozz~aomk

mapgh bax n` `o}igy }mog`h pz obh p}`h ca~ h`s`gazibf kifk

xk~apfkzpx f`baxyzibf o}}oy}"

^`m`bxgy' xk` Iggpeibo Fagh`bFox` f`baxyzibf o}}oy}'

og}a ~`c`~~`h xa o} agifabpmg`axih` zaag o}}oy} +AZO})'

v`~` h`}ifb`h ca~ maezg`{ zgobx f`bae`} }pmk o} }ayn`ob

obh no~g`y +Kyx̀ b `x og" >::;< ^a}xal} ̀ x og" >::3)" Ib naxk

AZO}' ozz~a{ieox`gy 9:% ac xk` ]BZ o}}oy} z~ahpm`h

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`x og" >::;)" Kav`s`~' `s`b xkapfk no~g`y obh }ayn`ob

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f`b`} +Ngobm obh Vagc` >::8)' xk` g`s`g ac f`b` hpzgimo

xiab obh kaeagafy ac xk` hpzgimox`h mazi`} i} co~ g`}} xkob

xka}` ib ~`m`bxgy `sags`h zagyzgaih}' }pmk o} vk`ox"

Vk`ox }z`mi`} ca~e o mgo}}imog zagyzgaih }`~i`} ac hiz

gaih' x`x~ozgaih obh k`{ozgaih }z`mi`} xkox `sags`h o} cag

gav}" X`x~ozgaih vk`ox X~iximpe xp~fihpe +f`bae` ca~epgoOONN) `sags`h sio kyn~ihitoxiab ac xva hizgaih }z`mi`}

mga}`gy ~`gox`h xa X" p~o~xp +f`bae` ca~epgo OO) obh O`fi%

gaz} }z`gxaih`} +f`bae` ca~epgo ]]' vk`~` ] i} }ieigo~ xa

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Hsa~ol x̀ og" ?99=)" Kyn~ihitoxiab ac X" xp~fihpe vixk hiz

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gpxiab ac k`{ozgaih vk`ox' X" o`}xispe +f`bae` ca~epgo

OONNHH) +Liko~o ?988< EmCohh`b obh ]`o~} ?983)"

Zagyzgaih vk`ox f`baxyzibf mp~~`bxgy ~`gi`} ab ^CGZ'

OCGZ' ]]^' HO~X a~ f`bae`}z`miVm ]X] eo~l`~} +Figg

`x og" ?99?< Gaxxi x̀ og" >:::< ]abf x̀ og" >::0< Olno~i x̀ og"

>::3)" Kav`s`~' bab` ac xk`}` eo~l`~} i} `o}igy oe`bong`

xa kifkxk~apfkzpx f`baxyzibf" K`~` v` h`eab}x~ox` xkox

xk` Iggpeibo N`ohO~~oy zgoxca~e obh Fagh`bFox` ]BZ

o}}oy z~asih` o co}x' ~`giong` obh ma}x`S ̀ mxis` e`xkah ca~

xk` go~f`}mog` omqpi}ixiab ac ]BZ f`baxyz` hoxo ib zagy

zgaih vk`ox" Ogabf vixk xk` ~`m`bxgy h`s`gaz`h ~`}ap~m` ac 

b`o~gy >'::: vk`ox gami ko~na~ibf ]BZ} +kxxz4!! 

vk`ox"zv"p}ho"fas!]BZ!b`v!ibh`{"}kxeg) obh ohsobm`} ib

}`qp`bmibf x`mkbagafi`} eolibf kifkxk~apfkzpx ]BZ hi}

mas`~y o c`o}ing` xo}l +No~notpl `x og" >::6< Eo~fpgi`}

`x og" >::0< ]k`bhp~` `x og" >::0)' xk` Iggpeibo N`ohO~~oy

zgoxca~e az`b} b`v za}}inigixi`} ca~ xk` f`bae`vih` obog

y}i} ac f`b`xim so~ioxiab ib vk`ox"

Eox`~iog} obh e`xkah}

Zgobx eox`~iog

O xaxog ac 9? zagyzgaih vk`ox gib`} ibmgphibf 0= gib`} ac 

vigh x`x~ozgaih `ee`~ vk`ox X~iximpe xp~fihpe }}z" him%

ammaih`} +k`bm`ca~xk X" himammaih }̀)' >9 gib`} ac k`{ozgaih

X" o`}xispe +?3 so~i`xi`} obh ?= gobh~om`}) obh 9 gib`} ac 

}ybxk`xim vk`ox h`~is`h c~ae xk` kyn~ihitoxiab ac so~iap}

x`x~ozgaih vk`ox} vixk   O`" xop}mkii v`~` f`baxyz`h ox 93

]BZ gami" Xk` ih`bxixy ac xk`}` gib`} i} }peeo~it`h ib }pz

zg`e`bxo~y Xong` ]?" Zgobx} v`~` f~avb ib o f~``bkap}`

obh HBO vo} `{x~omx`h c~ae g`os`} ac =”8v``lagh }``h

gibf} omma~hibf xa o h`}m~in`h z~axamag +Hsa~ol `x og" ?9;;)"

F`baxyzibf obh hoxo obogy}i}

O }`x ac ?=0 ]BZ} hi}mas`~`h ib o zob`g ac => gib`} ac 

x`x~ozgaih obh k`{ozgaih vk`ox ny }`qp`bmibf ac 9> f`b`

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Xk`a~ Ozzg F`b`x

 ?>=

c~ofe`bx} vo} havbgaoh`h c~ae xk` Vk`ox ]BZ Hoxono}`

+kxxz4!!vk`ox"zv"p}ho"fas!]BZ!b`v!ibh`{"}kxeg)" Xk` xaxog

g`bfxk ac }`qp`bm`h ~ f̀iab} vo} 0?'89= nz" Xk` hi}mas`~y

zob`g ibmgph`h ?: omm`}}iab} ac vigh `ee`~' ?= omm`}}iab}

ac k`{ozgaih vk`ox ~`z~`}`bx`h ny gobh~om`} obh 9 omm`}

}iab} ac }ybxk`xim vk`ox} +kxxz4!!vk`ox"zv"p}ho"fas!]BZ! 

b`v!ibh`{"}kxeg obh ]pzzg`e`bxo~y Xong` ]?)" Xk` gi}x ac 

]BZ} i} z~asih`h ib xk` ]pzzg`e`bxo~y eox`~iog}" ^`z`xixis` `g`e`bx} ib xk` }`qp`bm`} h`x`mx`h ny maezo~ibf xk`e

vixk xk` X^@Z +kxxz4!!vk`ox"zv"p}ho"fas!IXEI!^`z`ox}! )

obh FI^I +vvv"fi~ib}x"a~f) hoxono}`} v`~` eo}l`h" Xk`

]BZko~na~ibf }`qp`bm`} v`~` xk`b }pneixx`h xa Iggpeibo

ca~ z~am`}}ibf ny Iggpeibo¢ O}}oy H`}ifb Xaag +OHX)"

OHX f`b`~ox`} }ma~`} ca~ `omk ]BZ xkox mapgh so~y c~ae :

xa ?< ]BZ} vixk xk` }ma~`} onas` :"3 kos` o kifk z~anonig

ixy xa n` mabs`~x`h ibxa o }pmm`}}cpg f`baxyzibf o}}oy" Ib o

}`x ac ?=0 }pneixx`h ]BZ}' xk` OHX }ma~` so~i`h c~ae :"?;

xa :"99 vixk e`ob :";0 +Xong` ]>)" O xaxog ac 93 ]BZ }ix`}

xkox v`~` z~`}`bx ox xk` c~`qp`bmy onas` > ib xk` hi}mas`~y

zob`g obh kosibf OHX }ma~`} onas` :"3 v`~` }`g`mx`h ca~

AZO h`}ifb +Xong`} ]>' ]=)" Apx ac 93 ]BZ}' >3 v`~` ib

xk` vk`ox Hf`bae` obh 6: ]BZ} v`~` ib xk` Of`bae`"

O xaxog ac ?0: bf ac f`baeim HBO z`~ zgobx vo} p}`h

ca~ Iggpeibo ]BZ f`baxyzibf ox xk` PM Hosi} F`bae`

m`bx`~ +vvv"f`bae m̀`bx`~"pmhosi}"`hp!hboQx`mkbagafi`})

p}ibf xk` Iggpeibo N`ohO~~oy zgoxca~e obh Fagh`bFox`

O}}oy caggavibf xk` eobpcomxp~`~’} z~axamag" Xk` [pa~`}

m`bm` ieof`} ac ob o~~oy eox~i{ mo~~yibf My= obh My0

gon`g`h n`oh} v`~` f`b`~ox`h vixk xk` xvamkobb`g }mobb`~"

^ov kyn~ihitoxiab ibx`b}ixy hoxo z~am`}}ibf' mgp}x`~ibf obh

f`baxyz` moggibf v`~` z`~ca~e`h p}ibf xk` f`baxyzibf

eahpg` ib xk` N`oh]xphia zomlof` +Iggpeibo' ]ob Hi`fa'

MO' P]O)" Iggpeibo h`s`gaz`h o }`gcba~eogitoxiab ogfa

~ixke xkox ~`gi`} ab ibca~eoxiab mabxoib`h ib `omk o~~oy"

Xki} ogfa~ixke ohdp}x} ca~ mkobb`gh`z`bh`bx ibx`b}ixy

so~ioxiab}' hiS ̀ ~`bm`} ib xk` nomlf~apbh n`xv``b xk` mkob

b`g}' obh za}}ing` m~a}}xogl n`xv``b xk` hy`}" Xk` ba~eogi

toxiab z~am`hp~` iezg`e`bx`h ib xk` N`oh]xphia

f`baxyzibf eahpg` ibmgph`} apxgi`~ ~`easog obh noml

f~apbh ma~~`mxiab obh }mogibf +h`xoig} ac xki} z~az~i`xo~y

ba~eogitoxiab ogfa~ixke mapgh n` anxoib`h c~ae Iggpeibo'

]ob Hi`fa' MO)" N`ca~` f`baxyz` moggibf' xk` x~iee`h

e`ob ibx`b}ixi`} v`~` mogmpgox`h c~ae xk` ba~eogit`h ibx`b

}ixy sogp`} anxoib`h ca~ `omk n`oh xyz` ab xk` o~~oy ny

~`d`mxibf apxgi`~} xa `b}p~` kifk qpogixy ac f`baxyz` hoxo"

F`baxyz` mogg} v`~` f`b`~ox`h p}ibf xk` F`bMogg }acxvo~`

ibma~za~ox`h ibxa xk` N`oh]xphia zomlof`" Xki} ogfa~ixke

p}`} o Noy`}iob eah`g xa o}}ifb ba~eogit`h ibx`b}ixy sog

p`} xa ab` ac xk` xk~`` za}}ing` kaeatyfap} obh k`x`~aty

fap} f`baxyz` mgp}x`~}" Ib xk` z~`}`bm` ac abgy xva

kaeatyfap} mgp}x`~}' F`bMogg maezpx`} xk` gamoxiab ac o

ei}}ibf k`x`~atyfap} mgp}x`~ ny }iepgoxibf hoxo p}ibf xk`

o~xiVmiog b`p~og b`xva~l +]k`b `x og" >::0)" ]ibm` abgy xva

mgp}x`~} v`~` `{z`mx`h ca~ kaeatyfap} zagyzgaih vk`ox

gib`} +}`` ‗Hi}mp}}iab― ca~ h`xoig})' xk` f`baxyz` mgp}x`~}

f`b`~ox`h ca~ `omk ]BZ gamp} ny F`bMogg v`~` `hix`h eob

poggy ocx`~ si}pog ib}z`mxiab ac My= obh My0 [pa~`}m`bm`

ibx`b}ixy mgp}x`~ibf ab xvahie`b}iabog Mo~x`}iob zgax}"

]BZ} xkox coig`h xa }kav xvaf~apz mgp}x`~ibf v`~`

`{mgph`h c~ae xk` obogy}i}"

F`baxyzibf `~~a~ ~ox` vo} o}}`}}`h ny maezo~ibf ]BZf`baxyz`} h`x`~eib`h vixk xk` Fagh`bFox` o}}oy vixk

xka}` h`x`~eib`h ny ]obf`~ }`qp`bmibf" X~om` Vg`} ca~ 03

]BZko~na~ibf f`b` gami v`~` havbgaoh`h c~ae xk` Vk`ox

]BZ z~ad`mx hoxono}` +kxxz4!!vk`ox"zv"p}ho"fas!]BZ!b`v! 

ibh`{"}kxeg)" No}` moggibf obh }`qp`bm` o}}`engy v`~`

z`~ca~e`h p}ibf xk` zk~`h!zk~oz obh mab}`h z~af~oe}

+@vibf obh F~``b ?99;< @vibf `x og" ?99;< Fa~hab x̀ og"

?99;)" ]BZ hi}mas`~y vo} z`~ca~e`h vixk xk` zagyzk~`h

z~af~oe p}ibf xk` h`copgx }`xxibf} +]x`zk`b} `x og" >::3)

caggav`h ny si}pog ib}z`mxiab ac }`qp`bm` x~om` Vg`} obh

eobpog s`~iVmoxiab ac `omk hi}mas`~`h ]BZ" F`b`xim his`~

}ixy' h`Vb`h o} xk` z~anonigixy xkox xva ~obhaegy mka}`b

ogg`g`} c~ae xk` zazpgoxiab o~` hiS ̀ ~`bx +V`i~ ?993' z" ?0:'

?0?)' vo} mogmpgox`h p}ibf xk` Zav`~Eo~l`~ z~af~oe +Gip

obh Ep}` >::0)"

^`}pgx}

F`baxyzibf vixk xk` Iggpeibo Fagh`bFox` o}}oy

Xk` Fagh`bFox` o}}oy O]A} obh G]A} eoy obb`og xa xo~

f`x} ib xva a~ xk~`` f`bae`} ib vk`ox' h`z`bhibf ab

zgaihy" Fagh`bFox` f`baxyzibf ~`omxiab z`~ca~e`h ab

zagyzgaih vk`ox f`baeim HBO i} xk`~`ca~` `{z`mx`h xa

z~ahpm` My=!My0 [pa~`}m`bm` ~oxia} xkox hiS ̀ ~ c~ae xka}`

`{z`mx`h ca~ o hizgaih" Mab}ih`~ xk` a~ifib ac o b`v kozga

xyz` ib xk` Of`bae` ac x`x~ozgaih vk`ox +Cif" ?o' xaz

zob`g)" Epxoxiab +O2X) ib xk` Of`bae` m~`ox`} o h`~is`h

X no}` obh ob O!X ]BZ" Ib xk` Nf`bae`' xk` obm`}x~og O

no}` ~`eoib} pbmkobf`h" K`bm`' xk` ]BZ ~`}pgx} ib xva

kaeatyfap} f`baxyz`}' OOOO obh XXOO +Cif" ?o)" Xk`

~oxia ac O4X no}`} ib xk`}` xva f`baxyz`} o~` ?4: obh ?4?

+Cif" ?n)" ]ibm` My=!My0 gon`g`h O]A} obb`og xa xk` xo~

f`x} ib naxk O obh Nf`bae`}' xk` My=!My0 ~oxia} vigg

og}a n` ?4: obh ?4? +Cif" ?n)" Caggavibf o }ieigo~ ~`o}ab

ibf' xk` f`baxyz`} vigg n` OOOOOO obh XXOOOO ib

k`{ozgaih vk`ox' ~`}pgxibf ib xk` ?4: obh ?4> My=!My0

~oxia} +Cif" ?n)" Ib naxk `{oezg`}' xk` z~`}`bm` ac xk` xo~

f`x }`qp`bm` ab kae`agafap} mk~aea}ae`} }kicx} xk`

My=!My0 [pa~`}m`bm` ~oxia xavo~h xkox ac xk` obm`}x~og

ogg`g` ac xk` ]BZ obh oS ̀ mx} f`baxyz` mgp}x`~ibf" Xk` ]BZ

f`baxyzibf ac k`x`~atyfap} x`x~ozgaih obh k`{ozgaih

zgobx} vapgh z~ahpm` =4? obh 04? My=!My0 ~oxia}' ~`}z`m

xis`gy +Cif" ?n)"

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Xk`a~ Ozzg F`b`x

 ?>=

O 93zg`{ AZO ibmgph`h 6: Of`bae` ]BZ} obh >3

Hf`bae` ]BZ} xkox v`~` hi}mas`~`h ny }`qp`bmibf f`b`

c~ofe`bx} ib xk` ]BZ hi}mas`~y zob`g mab}i}xibf ac => gib`}

ac x`x~ozgaih obh k`{ozgaih vk`ox" Ca~ ob ]BZ xa n`

}`g`mx`h ca~ xki} }xphy' xk` eiba~ ogg`g` mapbx koh xa n` »>

+3"=%) ib xk` }m~``bibf zazpgoxiab" O xaxog ac 9? zagyzgaih

vk`ox gib`} ibmgphibf 0= gib`} ac X" himammaih`}' >9 gib`} ac 

k`{ozgaih X" o`}xispe +?3 mpgxiso~} obh ?= gobh~om`}) obh 9

gib`} ac }ybxk`xim vk`ox v`~` f`baxyz`h p}ibf Fagh`bFox`

o}}oy" Mgp}x`~ibf ac My= obh My0 ba~eogit`h ibx`b}ixi`} ib

o Mo~x`}iob zgax vo} p}`h xa ibc`~ xk` ]BZ f`baxyz`}" O

xaxog ac ; +?>%) ]BZ} ib x`x~ozgaih vk`ox obh ?0 +?0"3%) ibk`{ozgaih vk`ox xkox coig`h xa f`b`~ox` xk` xvaf~apz mgp}

x`~ibf `{z`mx`h ca~ kaeatyfap} gib`} v`~` `{mgph`h c~ae

cp~xk`~ obogy}`} +Cif" >)" F`baxyz` mogg} ca~ xk` ~`eoibibf

3> Of`bae` ]BZ} ib x`x~ozgaih vk`ox obh ;? O obh

Nf`bae` ]BZ} ib k`{ozgaih vk`ox mapgh n` f~apz`h ibxa

ab` a~ xva mgp}x`~} h`z`bhibf ab vk`xk`~ ob ]BZ }ix` vo}

eabaea~zkim a~ zagyea~zkim +Cif" =o' n)" Ib xk` zob`g ac 

k`{ozgaih vk`ox gib`}' ; ]BZ }ix`} v`~` eabaea~zkim +ogg

ib xk` Of`bae`) obh 6= v`~` zagyea~zkim +0> ]BZ} ib

xk` Of`bae` obh >? ]BZ} ib xk` Hf`bae`)" Ib x`x~ozgaih

vk`ox' > ]BZ }ix`} v`~` eabaea~zkim obh 3: ]BZ }ix`}

v`~` zagyea~zkim" O} `{z`mx`h' xk`~` v`~` ea~` eaba

ea~zkim ]BZ }ix`} ib mpgxisox`h vk`ox +>0!;?) xkob ib xk`

zob`g ac ?= vk`ox gobh~om`} +9!;?)"

Mabma~hobm` n`xv``b xk` ]BZ f`baxyz`} h`x`~eib`h

vixk xk` Fagh`bFox` o}}oy obh xka}` h`x`~eib`h vixk o

}xobho~h ]obf`~ }`qp`bmibf ozz~aomk vo} p}`h xa o}}`}}

xk` ommp~omy ac f`baxyzibf +Xong` ?)" Xk`}` maezo~i}ab}

v`~` eoh` ca~ 03 ]BZ }ix`} ib ?: omm`}}iab} ac X" himamma%ih`} obh >> omm`}}iab} ac k`{ozgaih vk`ox' ogxkapfk bax ogg

omm`}}iab} mapgh n` maezo~`h ca~ }ae` ]BZ} n`mop}` ac 

gavqpogixy }`qp`bm` ~`oh} +Xong` ?)" Apx ac ?'=:8 ]BZ

f`baxyz` mogg} f`b`~ox`h ny Fagh`bFox` o}}oy +8:= ]BZ

f`baxyz`} ib x`x~ozgaih vk`ox obh 9:? ]BZ f`baxyz`} ib

k`{ozgaih vk`ox) ?'>9> v`~` ih`bximog vixk xk` f`baxyz`}

anxoib`h ny ]obf`~ }`qp`bmibf" Xki} ~`}pgx} ib f`baxyzibf

Cif" ? Fagh`bFox` ]BZ f`baxyzibf ib zagyzgaih vk`ox" o Z~ibmizg`}

ac Fagh`bFox` ]BZ f`baxyzibf ib x`x~ozgaih vk`ox" Xk` xaz  zob`g

}kav} xva f`baxyz`} ac X" himammaih`} zagyea~zkim ca~ O!X ]BZ ib

xk` Of`bae` obh eabaea~zkim ca~ xk` X no}` ox xk` a~xkagafap} }ix`

ib xk` Nf`bae`" Xk` naxxae  zob`g }kav} xk` }oe` xva f`baxyz`} `{

m`zx ca~ xk` G]A obb`ogibf }ix` ib xk` Nf`bae` }kavibf o }`mabho~y

epxoxiab ~`}pgxibf ib iez`~c`mx eoxmk" Pz}x~`oe O]A} o~` ngp` obh

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}ix`} o~` }kavb ib zo~`bxk`}`}

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ibx`b}ixi`}

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Xk`a~ Ozzg F`b`x

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obh 99% ib k`{ozgaih vk`ox +Xong` ?)"

Mgp}x`~ibf ac Iggpeibo f`baxyz` mogg} eoy coig ib ob ogga

zagyzgaih ca~ xk` }oe` ~`o}ab} o} ib o hizgaih obh og}a ca~

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h`}ifb`h ca~ ]BZ h`x`mxiab +O]A obh G]A) obb`og xa }ka~x

~`fiab} [oblibf xk` qp`~y ]BZ}" ]ibm` xk`~` o~` xk~``

b`o~gy ih`bximog kae`agafap} }`x} ac mk~aea}ae`} z~`}`bx

ib xk` zagyzgaih vk`ox f`bae`' f`baxyzibf agifa} kos` pz

xa xk~`` kae`agafap} xo~f`x} xa nibh xa" Ic }`qp`bm`} ac 

xk`}` xo~f`x} o~` ih`bximog' f`baxyzibf agifa} vigg nibh vixk

`qpog `Ymi`bmy xa `omk ac xk` kae`agaf}" Kav`s`~' his`~

f`bm` ac xk` obb`ogibf }ix`} ib kae`agafap} mk~aea}ae`}

mapgh mkobf` agifabpmg`axih` nibhibf `Ymi`bmy obh ibx`~

c`~` vixk f`baxyzibf o}}oy ~`}pgxibf ib ix} coigp~` +Cif" ?o'

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mapgh n` xk` mop}` ac f`baxyzibf o}}oy coigp~`} a~ iboh`

qpox` mgp}x`~ibf' ~`fiab} [oblibf xk` xva ]BZ}

+v}bzN@88=?;6Q0OQ> obh v}bzN@8;99:?Q>HQ=) `{mgph`h

c~ae xk` obogy}`} v`~` ibs`}xifox`h +Cif" >)" Ca~ naxk ]BZ}

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0OQ> gamp}' vkimk i} gamox`h ib xk` Of`bae` v`~`

maezo~`h vixk xk` a~xkagafap} }`qp`bm`} c~ae naxk xk`

N obh Hf`bae`} ac zagyzgaih vk`ox" ]ieigo~gy'

}`qp`bm`} [oblibf xk` ]BZ }ix` ib xk` v}bzN@8;99:?Q

>HQ= gamp}' vkimk i} gamox`h ib xk` Hf`bae` v`~` mae

zo~`h vixk xk` Nf`bae` a~xkagafap} }`qp`bm`" Ib naxk

mo}`}' his`~f`bm` capbh ox xk` agifabpmg`axih` obb`ogibf

}ix`} mapgh n` o}}amiox`h vixk xk` coigp~` xa f`b`~ox` h`Vb`h

mgp}x`~} ab xk` Mo~x`}iob zgax} +Cif" >)" Ox xk`

v}bzN@88=?;6Q0OQ> ]BZ }ix`' naxk O]A} obh G]A

Of`bae` obb`ogibf }ix`} hiS ̀ ~`h ny ab` bpmg`axih` `ixk`~

ib xk` N a~ Hf`bae`" Ox xk` v}bzN@8;99:?Q>HQ= ]BZ

}ix`' xk` Of`bae` O]A} obb`ogibf }ix` hiS ̀ ~`h c~ae xkox

ib xk` Nf`bae` ny o hibpmg`axih` ibh`g"

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vk`ox vo} :"=? obh :"=8' ~`}z`mxis`gy" His`~}` zob`g ac ?=

vk`ox gobh~om`} h`eab}x~ox`h o kifk`~ g`s`g ac f`b`xim

his`~}ixy +Of`bae` ¯:"=>< Hf`bae` ¯:"=0) xkob xk`

zob`g ac ?3 vk`ox mpgxiso~} +Of`bae` ¯:">=< Hf`bae`

¯:"=?)" Xk` an}`~s`h >; obh ??% ~`hpmxiab ib xk` g`s`g} ac 

f`b`xim his`~}ixy ib xk` O obh Hf`bae`}' ~`}z`mxis`gy'

ea}x gil`gy ~`}pgx} c~ae xk` naxxg`b`ml mop}`h ny }`g`mxiab

Cif" = F`baxyz` mogg} ib o }oezg` ac 0= omm`}}iab} ac x`x~ozgaih X" him%

ammaih`} +o) obh =; omm`}}iab} ac k`{ozgaih X" o`}xispe +n)" Xk`

omm`}}iab bpen`~} ac eozz`h @]X pbif`b`} o~` }kavb ab xk` xaz ac 

`omk zob`g" F`baxyzibf zgax} v`~` f`b`~ox`h ny zgaxxibf My= obh My0

ba~eogit`h [pa~`}m`bm` ibx`b}ixi`}" o ]BZ }ix`} zagyea~zkim ib xk`

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ca~eoximog }m~``bibf z~am`hp~`} iezg`e`bx`h ib OHX xa

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b`fgifing` }ifbiVmobm` ca~ Iggpeibo f`baxyzibf ib zagyzgaih

vk`ox"

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ea~zkim }ix`}" Xki} x`mkbagafy ~`z~`}`bx} ob `{m`gg`bx xaag

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Agizkobx O' No~l`~ HG' ]xp`gzbof`g D^' Mk`` E] +>::>) N`ohO~~oy

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xk~apfkzpx f`baxyzibf" Niax`mkbiqp`} ]pzzg 043”0;

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comigixox`} vkag`f`bae` o}}amioxiab eozzibf ib `gix` ibn~`h m~azso~i`xi`}" Z~am Boxg Omoh ]mi P]O ?:=4?;303”?;33?

]o~lo~ Z' ]x`nnib} FG +?903) Ea~zkagafimog `sih`bm` mabm`~bibf xk`

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Zibax Bai~ nomx`~iog o~xiVmiog mk~aea}ae` mabxif}" F`b`xim}

?634>3=6”>30:

V`i~ N] +?993) F`b`xim hoxo obogy}i} II" ]ibop`~ O}}amiox`} Ibm"'

]pbh`~gobh

Tkoa L' O~obtobo ED' Lie ]' Gi}x`~ M' ]kibha M' Xobf M' Xaaeodiob

M' Tk`bf K' H`ob M' Eo~da~oe Z' Ba~hna~f E +>::6) Ob

 O~onihaz}i} `{oezg` ac o}}amioxiab eozzibf ib }x~pmxp~`h

}oezg`}" ZGa] F`b`x =4`8