Recombination in Human immunodeficiency virus type 1 (HIV-1): A Review

8/9/2019 Recombination in Human immunodeficiency virus type 1 (HIV-1): A Review

1/61

Edlbokc|e gi Xdlgehckopcgk ck Bteok Ceetkg`diclcdkls

Ycxt| Psvd ? "BCY%?#: O Xdycd

E@+ ED\HOB T@@CK

HCGPDLBKGNGAS OK@ ADKDPCL DKACKDDXCKA @C\LCVNCKD

NCID \LCDKLD \LBGGN. MBTNKO TKCYDX\CPS

MBTNKO% 3862. HOKANO@D\B

?0 IDHXTOXS 86?6


2/61



O Xdycd Xdvgxp \thecppd`

hs

E`+ Ed|hob T``ck

\pt`dkp C@: 6461?8\d||cgk: 8660%8664

\thecppd` pg

Hcgpdlbkgngas ok` Adkdpcl Dkackddxcka @c|lcvnckd. Ncid \lcdkld \lbggn.

Mbtnko Tkcydx|cps. Mbtnko. ck voxpcon itnicnnedkp gi pbd xdqtcxdedkp| igx

pbd `daxdd gi 0%sdox|

Holbdngx gi\lcdkld "H+\l+# ck Hcgpdlbkgngas ok` Adkdpcl Dkackddxcka

HCGPDLBKGNGAS OK@ ADKDPCL DKACKDDXCKA @C\LCVNCKD

NCID \LCDKLD \LBGGN. MBTNKO TKCYDX\CPS

MBTNKO% 3862. HOKANO@D\B

?0 IDHXTOXS 86?6


3/61



O Xdycd Xdvgxp \thecppd`

hs

E`+ Ed|hob T``ck\pt`dkp C@: 6461?8

\d||cgk: 8660%8664

Ovvxgyd` o| pg pbd |psnd ok` lgkpdkp hs

@x+ Mozc Egboeed` @c`oxtn C|noe

O||c|pokp Vxgid||gx

Hcgpdlbkgngas ok` Adkdpcl Dkackddxcka @c|lcvnckd

Ncid \lcdkld \lbggn. Mbtnko Tkcydx|cps

Mbtnko%3862. Hokano`d|b

Vxgid||gx Mbgk`gmdx Egozzoe Bg||ock "Vb@#

Bdo`.

Lbocxeok

Droeckopcgk Lgeecppdd

Hcgpdlbkgngas ok` Adkdpcl Dkackddxcka @c|lcvnckd

Ncid \lcdkld \lbggn. Mbtnko Tkcydx|cps

Mbtnko%3862. Hokano`d|b?0 Idhxtoxs 86?6


4/61



@dlnoxopcgk

Pbc| `c||dxpopcgk vovdx c| pbd gxcackon xd|doxlb gxm gi pbd otpbgx. drldvp o| gpbdxc|d

|popd`+ Pbd eopdxcon vxd|dkpd` bo| kgp hddk |thecppd` dcpbdx ck bgnd gx ck voxp igx o

`daxdd ixge pbc| gx oks gpbdx tkcydx|cps+ Pbd ick`cka| gi pbd xd|doxlb bo| kgp

hddk'cnn kgp hd vthnc|bd` oksbdxd cpbgtp pbd lgkltxxdkld gi pbd lgkldxkd`

|tvdxyc|gx+

E`+ Ed|hob T``ck

@opd: ?0+68+86?6


5/61


6/61

Olmkgndàedkp

Op pbc| gvvgxptkd egedkp. op icx|p cpb pbd koed gi Onecabps Onnob. C oe `dncabpd` pg

giidx es bdoxpcd|p pbokm| ok` `ddv axopcpt`d pg es |tvdxyc|gx @x+ Mozc Egboeed`

@còxtn C|noe. O||c|pokp Vxgid||gx. Hcgpdlbkgngas ok` Adkdpcl Dkackddxcka

@c|lcvnckd. Mbtnko Tkcydx|cps igx bc| doxkd|p atcòkld. ck|vcxopcgk ok` gh`txopd

|tvvgxp ck vxdvoxcka pbc| xdycd gxm+ Bc| ckyontohnd |taad|pcgk|. o`ycld ok`

lgkpcktgt| |tvdxyc|cgk boyd dkohnd` ed pg lgevndpd pbd gxm+

C oe xdonns axopcicd` pg onn es xd|vdlpd` pdolbdx| gi Hcgpdlbkgngas ok` Adkdpcl

Dkackddxcka @c|lcvnckd. Mbtnko Tkcydx|cps. Mbtnko igx vxgyc`cka ed onn iolcncpcd| pg

lgevndpd pbc| xdycd gxm+

C oe ydxs axopditn pg onn mthcgpdlb edehdx| igx vxgyc`cka ed ncpdxoptxd|. d|vdlconns

E`+ Ho`xtn Oxdick. Ethck Xobeok. E`+ Ecnno`tx Xobeok. \boxci Bo|ok ok` E`+

Ob|ok%Tn%Hoxc+

C oe axopditn pg es hd|p ixcdk` ok` onn gpbdx ixcdk`| ok` xggeeopd igx gvdk dk`d`

|tvvgxp ok` lggvdxopcgk pbop bdnvd` ed pg lgevndpd es gxm |eggpbns+

Ickonns. C oe axopditn pg es voxdkp| ok` gpbdx ioecns edehdx| igx pbdcx ceedo|txohnd

hnd||cka|. lgkpcktgt| ck|vcxopcgk ok` |tvvgxp pbxgtabgtp es ncid+

E`+ Ed|hob T``ck

?0 Idhxtoxs. 86?6


7/61



Oh|pxolp

Bteok ceetkg`diclcdkls ycxt| psvd ? "BCY%?# c| gkd gi pbd eg|p lbonndkacka ycxt|

eokmck` boyd dydx dklgtkpdxd + Ckpxo gx ckpdx |thpsvd xdlgehckopcgk c| ok dedxadkls

xd|ltd vnok igx BCY%? ixge Etnndx| xoplbdp pg |vdd` tv dygntpcgk ok` igx

hsvo||cka pbd gh|polnd| ck @KO |skpbd|c|+ Pdevnopd |cplbcka c| ok ckpdaxon voxp gi

BCY%? xdydx|d pxok|lxcvpcgk ok` pbd vxgld|| cp|dni dklgtxoad| xdlgehckopcgk tk`dx

bdpdxgngagt| lgk`cpcgk+ BCY%? xdlgehckopcgk c| gh|dxyd` bdk pg adkdpclonns

`c|pcklp adkgecl XKO egndltnd| oxd lgvolmoad` ck pbd |oed ycxcgk ok` |th|dqtdkp

ckidlpcgk gi kd ldnn+ Pbc| xdycd iglt|d| gk pbd edlbokc|e| pbop boyd hddk vxgvg|d`

igx pbd glltxxdkld gi xdpxgycxon xdlgehckopcgk cpb drvdxcedkpon vxggi+ @txcka pbd

|skpbd|c| gi icx|p @KO |pxok` "eckt| |pxok`#. xdlgehckgadkcl pdevnopd |cplbcka lok

hd pxcaadxd` hs pbd vxd|dkld gi hxdom"|#. dklgtkpdxcka o vot|d |cpd gx o |dlgk`oxs

|pxtlptxd gk pbd XKO pdevnopd+ Igxld` ok` `skoecl lgvs lbgcld xdlgehckopcgk.

vot|d `xcydk xdlgehckopcgk. olldvpgx ckyo|cgk xdlgehckopcgk hs `glmcka ok` nglmcka

eg`dn. ckpdxolpcyd bocxvck eg`dn ok` bocxvck ck`tld` pXKO ed`copd` "BCPED#

xdlgehckopcgk oxd vxgvg|d` edlbokc|e| igx eckt| |pxok` xdlgehckopcgk+ \pxok`

`c|vnoldedkp o||cecnopcgk eg`dn ok` BCPED oxd vnt| |pxok` xdlgehckopcgk

edlbokc|e|+ Pbd|d edlbokc|e| oxd kgp etptonns drlnt|cyd ok` onn eos hd gvdxopcyd

ck koptxon lgk`cpcgk+


8/61

Lbovpdx Gkd

Ckpxg`tlpcgk


9/61

Lbovpdx Gkd: Ckpxg`tlpcgk

?+? Holmaxgtk`

Bteok ceetkg`diclcdkls ycxt| "BCY# c| gkd gi pbd eg|p lbonndkacka ycxt| eokmck`

boyd dydx dklgtkpdxd`+ Kdoxns 89 sdox| oipdx pbd `c|lgydxs gi BCY o| pbd dpcgngaclon

oadkp giOlqtcxd` Ceetkg`diclcdkls \sk`xged "OC@\#. diidlpcyd lgkpxgn gi pbd ycxt|

ok` pbd vok`decl |pcnn xdeock| dnt|cyd "Posngx dp on+. 8662#+ Pbd Qgxn` bo| cpkd||d`

ktedxgt| dvc`decl ok` vok`decl `c|do|d| gydx pbd sdox| ok` igtabp ydxs dnn pg

xdlgydx ixge cp+ Bgdydx. cp bo| iocnd` pg xd|vgk` |pxgkans oaock|p OC@\. pbd eg|p

`do`ns. ncid%pbxdopdkcka gvvgxptkc|pcl ckidlpcgt| `c|do|d gi onn pced+ @dlo`d| gi

|doxlbcka ok` xd|doxlbcka boyd lged cpb ydxs ncppnd bgvd tkpcn ydxs xdldkpns bdk

\okgic Vo|pdtx| lokoxsvgr ydlpgx%ho|d` lok`c`opd ONYOL%BCY "yLV?48?# ck o

vxced%hgg|p xdaced. cpb o adkdpclonns dkackddxd` av?86 |txiold vxgpdck OC@\YOR

H'D. |bgd`


10/61

o| ck sdox ?339. 866? ok` 8662 pbd ktehdx gi kdns ckidlpd` vdgvnd dxd


11/61

lgxdldvpgx. d+a+ ixge LLX4 pg LRLX0+ O |pt`s ck Mdkso |bgd` pbd edok `txopcgk

ixge pbd hdackkcka gi OXP pg pbd |cplb gi lgxdldvpgx t|oad c| 8+9 sdox| "Ndehd dp

on+. 8663#+

Bgdydx. ck xdpxgycxt|d| `dndpdxcgt| etpopcgk| oxd on|g olltetnopd` ck o |tiiclcdkpns

bcab xopd pg o`yokld Etnndx| xoplbdp "olltetnopcgk gi `dndpdxcgt| etpopcgk|# ok`

holmox` lgevdk|opgxs etpopcgk| lokkgp pboxp pbd xoplbdp "Lbog. ?336#. eomcka pbd

ycxt| nd|| ycxtndkp ok` egxd |t|ldvpchnd pg pbd ceetkd xd|vgk|d ok` okpcxdpxgycxon

pbdxovs+ Htp pbd ycxt| bo| dygnyd` kd |pxopdacd| gydx pbd pced pg gydxlged pbd

btx`nd+ Xdlgehckopcgk. o vxcecpcyd igxe gi |drton xdvxg`tlpcgk. ck xdpxgycxt|d| |dxyd|

o| ok dedxadkls xd|ltd vnok ixge Etnndx| xoplbdp ok` c| pbd hd|p os pg |vdd` tv

pbd hdkdiclcon etpopcgk| pg cklxdo|d icpkd||. xdegyd adkgecl `oeoad| ok` adkdxopd

yoxcohcncps ck pbd vgvtnopcgk "Eoacgxmckc| dp on+. 866


12/61

Lbovpdx Pg

Lno||ciclopcgk ok` Adgaxovbclon

@c|pxchtpcgk gi BCY


13/61

Lbovpdx Pg: Lno||ciclopcgk ok` Adgaxovbclon @c|pxchtpcgk

gi BCY

8+? Dedxadkld gi BCY

BCY. o ndkpcycxt| ixge xdpxgycxt| ioecns. c| `c|pxchtpd` pbxgtabgtp pbd gxn` ok`

lno||cicd` ckpg `ciidxdkp psvd|. |thpsvd| ok` lno`d| "Ica 8+?#+ BCY%? ok` BCY%8 oxd pbd

pg mkgk psvd| gi BCY bdxd psvd%? c| egxd ycxtndkp ok` c`dns `c|pxchtpd` pbok

psvd%8 "Acnhdxp dp on+. 866


15/61

8+< Adgaxovbclon `c|pxchtpcgk

8+


16/61

8+


17/61

Ica 8+


18/61

Lbovpdx Pbxdd

\tvdxckidlpcgk ok` Lgvolmoacka


19/61

Lbovpdx Pbxdd: \tvdxckidlpcgk ok` Lgvolmoacka

BCY%? c|. o XKO ycxt| bclb lgkpock| pg lgvcd| gi |ckand%|pxok`d` vg|cpcyd XKO

egndltnd| pbop `cxdlpns lg`d| ycxon vxgpdck| "Aondppg ) Kdaxgkc. 8664= Vcdxld. 8664#.

`cvngc` ck koptxd ok` xdlgehckopcgk vnos| o ycpon xgnd ck ycxt| xdvnclopcgk ok`

dygntpcgk "Jdpzp dp on+. 8666= Kdaxgkc ) Htl. 866?h#+ Pg adkdpclonns `c|pcklp XKO

egndltnd| cpbck pbd |oed ycxcgk oxd kdld||oxs igx xdlgehckopcgk "Aondppg )

Kdaxgkc. 8664#+ Bdkld |tvdxckidlpcgk ok` |th|dqtdkp lgvolmoacka gi bdpdxgzsagt|

XKO egndltnd|. bclb adkdxopd| bdpdxg`cedxcl voxpclnd|. oxd kdld||oxs igx

xdlgehckopcgk "Aondppg ) Kdaxgkc. 8664= Bt ) Pdeck. ?336#+

O ldnn ckidlpd` cpb gkd xdpxgycxt| lokkgp hd xdckidlpd` hs o xdpxgycxt| gi pbd |oed

psvd. o ngka |pok`cka ldkpxon `gaeo gi xdpxgycxon lgckidlpcgk "_tckgkd|%Eopdt )

Oxp|. ?333# bo| hddk lbonndkad` ydxs xdldkpns hs pbd dedxadkld gi xdlgehckokp BCY%

?+ Pbgtab pbd `gaeo o| ho|d` gk |gnc` drvdxcedkpon dyc`dkld. d+a+ BCY%? ckidlpd`

ldnn| |bg `dlxdo|d` drvxd||cgk gi L@0 egndltnd gk pbd ldnn |txiold. `td pg pbd

lgevndr igxeopcgk hdpddk pbd xdldvpgx egndltnd "L@0# ok` pbd ycxon dkydngvd adkd

vxg`tlp "Bgrcd dp on+. ?329= Vcatdp dp on+. ?333#. etnpcvnd ckidlpcgk| c| vg||chnd onhdcp

op ydxs ng ixdqtdkls+ @gkxdatnopcgk gi LRLX0. o lgxdldvpgx ixge A%vxgpdck%

lgtvnd` lbdegmckd xdldvpgx ioecns. `td pg BCY anslgvxgpdck % LRLX0 lgevndr

igxeopcgk o| on|g xdvgxpd` ck xdldkp |pt`s "Lbgc dp on+. 8662#+


21/61


22/61

Lbovpdx Igtx

Edlbokc|e gi Xdlgehckopcgk ck BCY%?


23/61


28/61

pbop. cp| XKO c| ydxs etlb òeoad%vxgkd ok` cp c| oneg|p cevg||chnd pg c|gnopd

ycxcgk| cpb |cakciclokpns tkhxgmdk egndltnd|. tknd|| drpxogx`ckoxs edo|txd|. |tlb o|

boxyd|pcka pbd ycxt| op |bgxp ckpdxyon. oxd pomdk+ O| igxld` lgvs lbgcld c| o vxgvg|d`

xdlgehckopcgk edlbokc|e igx xdpxgycxt|. |cecnox |cptopcgk eos drc|p ck BCY%?+ Pbd

edlbokc|e gi BCY%? adkgecl XKO hxdomoad c| kgp lndox. htp cp lgtn` hd pbd gtplged

gi oks gi pbd ignngcka|:

"?#Bg|p @didk|d: Igxdcak adkdpcl eopdxcon| oxd gipdk `daxo`d` gx d`cpd` hs bg|pdk`gktlndo|d ok` pbd lgevgkdkp gi ckkopd okpcycxon |s|pde+ Ovgncvgvxgpdck H

"OVGH# eXKO%d`cpcka loponspcl vgnsvdvpc`d "OVGHDL# vxgpdck| |dxyd o ycpon

xgnd ck `daxo`cka ycxon adkgecl XKO hs `doeckopcka lspg|ckd xd|c`td| ck

ko|ldkp l@KO "Boxxc| ) Nc`òedkp. 8660= Ehc|o dp on+. 8661#+

"8#Ckpdaxo|d |pcetnopgx "C\# egndltnd: Xdpxgycxon ckpdaxo|d bo| o kgk|vdlcicldk`gktlndo|d olpcycps bclb lok hd eg`tnopd` hs ldxpock lgevgtk`| lonnd`

ckpdaxo|d |pcetnopgx "C\#+ Ixge pbd drvdxcedkp gi \t`gn dp on+. "86?6#. cp c|

dyc`dkp pbop olpcgk gi xdpxgycxon ckpdaxo|d lok hd eokcvtnopd` hs C\ lgevgtk`.

pg òeoad xdpxgycxon @KO+

Pbt| BCY%? vxgycxon @KO lok hd òeoad` ok` pxok|lxcvpcgk ixge pbc| @KO cnn

adkdxopd o ixoaedkp gi ycxon XKO egndltnd+ \tlld||cyd lgvolmoacka gi pbd adkdxopd`

|bgxp XKO cpb o xdatnox XKO egndltnd lgtn` xd|tnp| ck igxld` lgvs lbgcld

xdlgehckopcgk ck |th|dqtdkp ckidlpcgk hs pbc| ycxcgk+

Igxld` lgvs lbgcld c| o dnn d|pohnc|bd` edlbokc|e gi BCY%? xdlgehckopcgk. bdxd

hxdom"|# ck pbd pdevnopd |pxok` bonp| xdydx|d pxok|lxcvpcgk ok` xd|tevpcgk gi @KO

|skpbd|c| lok gkns hd `gkd hs |cplbcka pbd pdevnopd |pxok` ixge pbd hxgmdk pdevnopd

pg pbd ckpolp gkd "Ica 0+


29/61

Igxld` lgvs lbgcld c| o xdlgehckopcgk edlbokc|e bdxd pdevnopd XKO hxdom"|#

bonp| eckt| |pxok` @KO |skpbd|c| ok` XKo|d B `daxo`d| pdevnopd |pxok` pbop vo||d`

pbd vgnsedxo|d |cpd gi xdydx|d pxok|lxcvpo|d "XP# "Lgiick. ?313#+ Cp c| o |txycyon

edlbokc|e igx pbd |skpbd|c| gi lgevndpd @KO vxgycxt| "Aondppg ) Kdaxgkc. 8664#

|ckld hxgmdk XKO egndltnd| lokkgp |dxyd o| pdevnopd igx lgevndpd eckt| |pxok`

@KO |skpbd|c| "Pdeck. ?33?#+ Ixdd ko|ldkp @KO pbdk ho|d vocx| cpb pbd

lgxxd|vgk`cka |dqtdkld| gi pbd gpbdx |pxok` "Ica 0+


31/61

Ica 0+0: Igxld` xdlgehckopcgk o||os+ "C# Pg%ydlpgx xdlgehckopcgk o||os o| `d|lxchd` ck

xdidxdkld "@taaon dp on+. 8661= Vidciidx ) Pdnd|kcp|ms. 866?#+ Kdns |skpbd|czd` @KO

tk`dxagd| o xdlgehckgadkcl |pxok` pxok|idx ixge pbd 4 dk` gi ok XKO `gkgx "@# pdevnopd pg

pbd


32/61

igx ho|d%vocxcka ckpdxolpcgk| cpb pbd olldvpgx pdevnopd. pbd KL vxgpdck vxgegpd|

bs`xgadk hgk`cka ok` `tvndr igxeopcgk+ Oipdx pbd @KO `tvndr igxe| < pg pbd XP.

pbd vxcedx dk` gi pbd ko|ldkp @KO c| xdndo|d` ixge pbd `gkgx pdevnopd. bclb ecabp

ckygnyd cp| `c||glcopcgk ixge pbd XP+ Ickonns. pbd vxcedx dk` o||glcopd| cpb pbd

olldvpgx pdevnopd. lgevndpcka pbd pdevnopd |cplb+

Pbd `skoecl lgvs lbgcld eg`dn bdnv| pg ckpdaxopd |ged gi pbd vxdycgt|ns vxgvg|d`

eg`dn| ok` drvdxcedkpon gh|dxyopcgk| o||glcopd` cpb XP pdevnopd |cplbcka+ Icx|p.

pbd vxdycgt|ns vxgvg|d` igxld` lgvs lbgcld eg`dn c| lgk|c|pdkp cpb pbd `skoecl

lgvs lbgcld eg`dn+ Pbd igxld` lgvs lbgcld eg`dn vxgvg|d` pbop bdk XP dklgtkpdx|

o hxdom ck pbd pdevnopd XKO. pbd XP |cplbd| pdevnopd|+ Ck pbd `skoecl lgvs lbgcld

eg`dn. o hxdom ck pbd XKO gtn` xdvxd|dkp gkd drpxded |cptopcgk ck pbd |vdlpxte ck

bclb pbd xopd gi @KO vgnsedxczopcgk bo| hddk xd`tld` pg zdxg+ Pbd ghncaopgxs

pdevnopd |cplbd| `txcka xdydx|d pxok|lxcvpcgk. koedns. eckt|% ok` vnt| |pxok` @KO

pxok|idx|. on|g xdvxd|dkp pbd |oed |cptopcgk ck bclb pbd xopd gi @KO vgnsedxczopcgk c|

zdxg+ \dlgk`. pbd `skoecl lgvs lbgcld eg`dn c| lgk|c|pdkp cpb |dydxon vxdycgt|

gh|dxyopcgk| pbop |dlgk`oxs |pxtlptxd| ck pbd pdevnopd XKO lxdopd XP vot|d |cpd|.

cklxdo|cka pbd ixdqtdkls gi pdevnopd |cplbcka+ \ckld XP vot|cka c| ncmdns pg xd`tld

pbd xopd gi @KO vgnsedxczopcgk. cp| diidlp gk XP pdevnopd |cplbcka |bgtn` hd |cecnox

pg pbop gh|dxyd` igx bs`xgrstxdo "BT# pxdopedkp gx XP etpopcgk| drvdlpd` pg xd`tld

pbd xopd gi @KO vgnsedxczopcgk+


33/61

Ica 0+4: @skoecl lgvs lbgcld eg`dn igx XP pdevnopd |cplbcka "\yoxgy|moco dp on+. 8666#+

\bo`d` hgrd| xdvxd|dkp `cxdlp xdvdop| ck ok XKO pdevnopd+ Bgxczgkpon oxxg| xdvxd|dkp

ko|ldkp @KO+ Pbd pbclmkd|| gi pbd|d oxxg| ck`clopd| pbd xdnopcyd vgnsedxczopcgk xopd= pbd

pbclmdx pbd oxxg. pbd bcabdx pbd xopd gi vgnsedxczopcgk+ \eonn gvdk hgrd| xdvxd|dkp XKO

`daxo`d` hs pbd XKo|d B `geock+ Ck pbd lo|d gi |ng XKo|d B olpcycps. pbd `daxo`d` XKO

ixoaedkp| oxd |bgk o| noxadx gvdk hgrd|+ Bs`xgadk hgk`| hdpddk pbd XKO pdevnopd ok`

ko|ldkp @KO oxd `d|cakopd` hs ydxpclon eoxm|+ Ydxpclon oxxg| gi yoxcgt| pbclmkd||d|

ck`clopd pbd xdnopcyd diiclcdkls gi pdevnopd |cplbcka+ QP: cn` psvd+

0+8+?+< Vot|d `xcydk xdlgehckopcgk

Edlbokc|e gi igxld` lgvs lbgcld ok` vot|d `xcydk xdlgehckopcgk c| |cecnox htp pbd

koptxd gi gh|polnd pbop oxxd|p xdydx|d pxok|lxcvpcgk c| `ciidxdkp "Kdaxgkc ) Htl.

866?h#+ Gk ckpolp XKO pdevnopd|. xdydx|d pxok|lxcvpcgk lok hd |pgvvd` "Ica 0+9# ok`

pdevnopd |cplbcka lok hd pxcaadxd` bdk cklgxvgxopcgk gi ktlndgpc`d| op axgcka


34/61

Ica 0+9: Vot|d%`xcydk xdlgehckopcgk+ Xdydx|d pxok|lxcvpcgk c| |pgvvd` gk pbd `gkgx XKO

bdk pbd cklgxvgxopcgk gi kdrp lgevndedkpoxs ktlndgpc`d hdlged| voxpcltnoxns `ciicltnp+ Pbd

xdydx|d pxok|lxcvpo|d "sdnng dnncv|d# `c||glcopd| ixge pbd vxcedxpdevnopd lgevndr "|dlgk`

vokdn#. ok` pbd ko|ldkp @KO vxgaxd||cydns bshxc`czd| pg pbd olldvpgx XKO "pbcx` vokdn#+

Gkld pbd

xdydx|d pxok|lxcvpo|d cnn dydkptonns xd|ted @KO |skpbd|c| "hgppge vokdn# "Kdaxgkc ) Htl.866?h#+

0+8+?+


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ycyg+ Xdldkp |pt`cd| gk BCY%? boyd on|g |bd` ncabp gk pbc| o|vdlp gi pbd qtd|pcgk hs

ignngcka o `ciidxdkp |pxopdas "Ndys dp on+. 8660#+

0+8+?+|pxok` pxok|idx diiclcdkls+

0+8+?+0 Olpcyd `c|vnoldedkp xdlgehckopcgk

Ck pbc| eg`dn "Kdaxgkc ) Htl. 866?o= Kdaxgkc ) Htl. 866?h#. |pxtlptxd gi pbd

olldvpgx pdevnopd c| o||ted` pg hd ycpon igx |pxok` pxok|idx ok` xdlgehckopcgk+ \pxok`

pxok|idx c| vxgvg|d` pg glltx ck pg |pdv|: "?# pbd olldvpgx XKO `glm| gk pbd ko|ldkp


37/61

0+8+?+9 Ckpdxolpcyd bocxvck eg`dn gi xdlgehckopcgk

O| vxgvg|d` hs Kdaxgkc ) Htl "866?h#. vonck`xgecl |dqtdkld| gi pxok|olpcyopcgk

xd|vgk|d "POX# dndedkp igxe bocxvck |pxtlptxd "Ica 0+3# ok` pbd |pde gi pbd bocxvck

ednp| o| xdydx|d pxok|lxcvpcgk vxgaxd||d| pbxgtab cp. onngcka pbd igxeopcgk gi o kd

ckpdxolpcgk ck pbd bocxvck xdacgk hdpddk `gkgx ok` olldvpgx XKO|+ @txcka lgvscka

gi pbd `d|ldk`cka voxp gi pbd bocxvck. pbd ko|ldkp @KO dydkptonns bshxc`czd| pg pbd

olldvpgx XKO ok` `cydxp| |skpbd|c| ixge pbd `gkgx XKO pg olldvpgx XKO "Kdaxgkc

) Htl. 866?h#+

Ica 0+3: Ckpdxolpcyd bocxvck eg`dn gi xdlgehckopcgk "Kdaxgkc ) Htl. 866?h#+

Egtedk dp on+. "866


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39/61

ok` pbd olldvpgx c| oD8 "|dd Ica 0+?? O. lgntek CC#+ @gkgx XKO. xd`= olldvpgx XKO. hntd=

ko|ldkp @KO. hnolm "pbd oxxg ck`clopd| pbd `cxdlpcgk gi xdydx|d pxok|lxcvpcgk#+ "O# Pbd |pde

gi pbd |pde%nggv "\N# bocxvck gk pbd olldvpgx XKO c| xdvxd|dkpd` o| lgk|pcptpd` hs o ngdx

ok` ok tvvdx voxp. o ok` h. xd|vdlpcydns. okkdond` pg pbdcx lgevndedkpoxs |dqtdkld|. o! ok`

h!+ Pbd nggv xdacgk c| ck`clopd` o| l+ "H# xdydx|d pxok|lxcvpcgk vxgaxd||d| gk pbd `gkgx XKO+

"L# Hdlot|d okkdoncka gi pbd ko|ldkp @KO c| ncmdns ioygxd` gk o |ckand%|pxok`d` xdacgk gi pbd

olldvpgx XKO. `glmcka ecabp glltx gkld pbd nggv xdacgk l c| xdydx|d%pxok|lxchd` "adkdxopcka

pbd lgevndedkpoxs |dqtdkld l!#+ "@# Pbd bshxc`czopcgk gi pbd ko|ldkp @KO gkpg pbd olldvpgx

XKO pbdk ckyo`d| pbd |pde xdacgk gi pbd bocxvck gk pbd olldvpgx XKO. adkdxopcka pbd

eorcete vg||chnd drpdkp gi `gthnd%|pxok`d` xdacgk|+ Pbd xd|tnpcka ckpdxed`copd |pxtlptxd

xd`xok gk pbd xcabp gi pbd vokdn xd|dehnd| ok ckpdxed`copd gi hxoklb ecaxopcgk glltxxcka

`txcka @KO xdlgehckopcgk+ O ecaxopcgk `gkox` gi pbd vg|cpcgk gi pbd lxg||gydx gtn`

ndo` pg pbd pxok|idx gi pbd


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dolb eg`dn pdevnopd+ Axos oxxg| ck`clopd pbd nglopcgk gi |vdlcicl xd|pxclpcgk |cpd| pbop. oipdx

xd|pxclpcgk okons|c| gi pbd xdlgehckokp @KO egndltnd|. onng eovvcka gi |pxok` pxok|idx

dydkp| ongka pbd |dqtdkld gi bgegngas+ Pbd|d |cpd| dxd adkdxopd` hs pbd ckpxg`tlpcgk gi

vgckp etpopcgk|+ Pbd ixdqtdkls gi xdlgehckopcgk gh|dxyd` tk`dx pbd `ciidxdkp drvdxcedkpon

lgk`cpcgk| c| acydk hdng+ "H# eovvcka gi |pxok` pxok|idx dydkp| op pbd ndydn gi pbd \N xdacgk

tk`dx pbd pg drvdxcedkpon lgk`cpcgk| pbop scdn` o bcab `daxdd gi pxok|idx "vokdn O. lgntek| C

ok` CC# o| vdxigxed` hs ckpxg`tlcka pbd o``cpcgkon xd|pxclpcgk |cpd DlgXC gk pbd `gkgx XKO+

Pbc| etpopcgk `c` kgp oiidlp pbd gydxonn ixdqtdkls gi xdlgehckopcgk+ Pbd xdacgk Dh c|

pbdxdigxd |th`cyc`d` ckpg pbd bocxvck ok` pbd `gk|pxdoe xdacgk+ Pg kgxeonczd igx pbd

`ciidxdkp |czd gi pbd pg xdacgk|. pbd ixdqtdkls gi xdlgehckopcgk c| acydk o| xopd| vdx kp+

0+8+?+1 Olldvpgx ckyo|cgk eg`dn gi xdlgehckopcgk

Gkoito%Ktao ) Pdnd|kcp|ms "8663# boyd vxgvg|d` o eckt| |pxok` xdlgehckopcgk

eg`dn lonnd` eckt|%|pxok` drlbokad gx olldvpgx ckyo|cgk edlbokc|e+ Ollgx`cka pg

pbc| eg`dn. XKo|d B `daxo`opcgk gi pdevnopd XKO drvg|d| pbd ko|ldkp eckt| |pxok`

@KO "Ica 0+?8#. bclb onng| ho|d vocxcka hdpddk ko|ldkp @KO ok`

lgevndedkpoxs xdacgk| gk pbd |dlgk` XKO gi pbd ycxcgk+ Pbd xd|tevpcgk gi @KO

|skpbd|c| gk pbd olldvpgx pdevnopd c| ignngd` hs pbd vxcedx |pxok` xdoncakedkp hs

hxoklb ecaxopcgk. scdn`cka eckt| |pxok` xdlgehckokp+ Ck pbc| eg`dn `gkgx%olldvpgx

bgegngas. hdbck` pbd axgcka vgckp. c| egxd cevgxpokp pbok bgegngas op pbd

lxg||gydx |cpd "Gkoito%Ktao ) Pdnd|kcp|ms. 8663#+


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Ica 0+?


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Honomxc|bkok dp on+. "866


46/61

Ica 0+?4: Edlbokc|pcl eg`dn igx BCPED xdlgehckopcgk hs o |dlgk` pXKO vxcecka dydkp

`txcka eckt|%|pxok` @KO |skpbd|c|+ Pbd |dlgk` pXKO vxcedx c| eoxmd` ck xd` ok`

lgevndedkpoxs ktlndgpc`d| hdpddk pbc| pXKO ok` pbd BCY%nbKdi adkged oxd bcabncabpd` hs

hntd hgrd| "Mgk|pokpckgyo dp on+. 8669#+

0+8+8 Vnt| |pxok` Xdlgehckopcgk

0+8+8+? \pxok` `c|vnoldedkp o||cecnopcgk eg`dn:

Pdeck "?33?#. Aondppg ) Kdakgxc "8664# ok` Gkoito%Ktao ) Pdnd|kcp|ms "8663#

boyd `dnckdopd` o eg`dn igx xdpxgycxon vnt| |pxok` xdlgehckopcgk. mkgk o| |pxok`

`c|vnoldedkp o||cecnopcgk eg`dn "Ica 0+?9#+ Xdlgehckopcgk `txcka vnt| "-# |pxok`

|skpbd|c| o| vg|ptnopd` gk pbd ho|c| gi `c|lgkpcktgt| koptxd gi "-# |pxok` @KO

|skpbd|c| ok` B%ncmd |pxtlptxd| gi jgckd` @KO| xdydond` ck dndlpxgk eclxgaxovb| ck

oycok |oxlgeo%ndtlg|c| ycxcgk| "Aondppg ) Kdaxgkc. 8664= Gkoito%Ktao )

Pdnd|kcp|ms. 8663#+ Ollgx`cka pg pbc| eg`dn. "-# |pxok` @KO |skpbd|c| ckcpcopd| ixge


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etnpcvnd vg|cpcgk| ok` |pxok` `c|vnoldedkp scdn`| ixdd @KO pocn| "Ica 0+?9# bclb

ckyo`d `gthnd%|pxok`d` xdacgk| gi pbd kdcabhgxcka ycxon adkged+ Pbdxdigxd gkd |pxok`

gi pbd ckyo`d` `tvndr voxpconns `c|vnold|+ Oipdx dk`gnspcl ltp ok` ncaopcgk gi kclm|. o

bdpdxg`tvndr adkdxopd|+ Pbc| bdpdxg`tvndr c| pbdk lgxxdlpd` hs ldnntnox xdvocx dkzsed

pg adkdxopd xdlgehckokp bgeg`tvndr+ Igx pbc| eg`dn. hgpb pbd voxdkpon XKO

egndltnd| |bgtn` hd |tlld||itnns lgvcd` ckpg pg eckt| |pxok` @KO. |ckld hxgmdk

egndltnd| lokkgp adkdxopd vnt| |pxok` xdlgehckokp "Pdeck. ?33?#+

Ica 0+?9: \pxok` `c|vnoldedkp%o||cecnopcgk eg`dn gi xdlgehckopcgk igx vnt| |pxok` |skpbd|c|

"Gkoito%Ktao ) Pdnd|kcp|ms. 8663#+

Pg `c|pcklp vnt| |pxok` ckcpcopcgk |cpd| boyd hddk c`dkpcicd` ck bteok

ceetkg`diclcdkls ycxt| "BCY#. pbd ldkpxon vgnsvtxckd pxolp "lVVP# ok` pbd vgnsvtxckd

pxolp nglopd` jt|p tv|pxdoe gi pbd T< xdacgk "T


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|skpbd|c|+ Pxok|idx diiclcdkls o| `cxdlpns xdnopd` pg olldvpgx lgkldkpxopcgk htp

ck`dvdk`dkp gi pbd xdolpcgk pced+ Pxok|idx lgtn` glltx pg olldvpgx| lgkpockcka 26. 06.

gx 86 ktlndgpc`d| gi bgegngas cpb pbd pdevnopd @KO+ T|cka `ciidxdkp olldvpgx|. cp

o| igtk` pbop @KO pg @KO pxok|idx glltxxd` op vg|cpcgk| pbxgtabgtp pbd `gkgx

pdevnopd. drldvp kdox pbd 4! dk`+ Pbc| |bg| pbop o ktehdx gi pbd |dqtdkld|

`gk|pxdoe gi pbd lVVP xdacgk lok vxgegpd pxok|idx. htp gkld |skpbd|c| bo|

vxgaxd||d` pg pbd vgckp bdxd pbd `gk|pxdoe |daedkp c| lgevndpdns `c|vnold`.

pxok|idx c| kgp onngd`+ Qbdk pbd @KO pg @KO pxok|idx xdolpcgk| dxd vdxigxed`

t|cka o pdevnopd lgkpockcka kgkycxon |dqtdkld|. pbd pxok|idx diiclcdkls `xgvvd`

|cakciclokpns+ Onn gi pbd|d gh|dxyopcgk| |tvvgxp pbd bsvgpbd|c| pbop lgevdpcpcgk igx

hck`cka pbd pdevnopd hdpddk pbd drpdk`cka tv|pxdoe vxcedx ok` pbd voxpns `c|vnold`

`gk|pxdoe vxcedx igxld| pxok|cdkp `c||glcopcgk gi pbd


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Ica 0+?1: \pxok` `c|vnoldedkp%o||cecnopcgk eg`dn| igx vnt| |pxok` @KO xdlgehckopcgk+ Vokdn

O |bg| pbd |pxok` `c|vnoldedkp o||cecnopcgk eg`dn o| vxgvg|d` hs Jtkabok| dp on+. "?328#+

Vokdn H |bg| o eg`dn igx pbd |pxok` `c|vnoldedkp%o||cecnopcgk dydkp o| vxgvg|d` hs Itdkpd|

dp on+. "?339#+ Pbd noppdx eg`dn `ciidx| ixge pbd igxedx ck pbop cp c| pbd tv|pxdoe vxcedx xopbdx

pbok pbd `gk|pxdoe vxcedx pbop pxok|idx| ok` lgkpcktd| |skpbd|c| gk pbd kd pdevnopd+ \gnc`

nckd| xdvxd|dkp |skpbd|c| ixge @KO pdevnopd ?. ok` boplbd` nckd| xdvxd|dkp |skpbd|c| ixge

@KO pdevnopd 8+

0+8+8+8 Bocxvck%ck`tld` pXKO%ed`copd` "BCPED# vnt| |pxok` xdlgehckopcgk

Mgk|pokpckgyo dp on+. "8669# bo| on|g vxgvg|d` o eg`dn igx vnt|%|pxok` @KO |skpbd|c|

bdxd o pdevnopd |cplbcka glltx ndo`cka pg xdlgehckopcgk "Ica 0+?2#+ Ck pbc| eg`dn.

cp c| o||ted` pbop XP c| ohnd pg lgvs pbd drpdk`d` bocxvck ck pbd ycxcgk XKO pdevnopd

`txcka eckt|%|pxok` |skpbd|c|. htp pbd dkzsed c| vot|d` hs pbd bocxvck `txcka vnt|%

|pxok` @KO |skpbd|c|+ O pXKO c| |th|dqtdkpns t|d` hs XP o| o pxok|cdkp pdevnopd pg

hsvo|| pbd bocxvck. ignngd` hs o xdptxk pg pbd ycxon XKO pdevnopd+ Pg xdacgk| gi

|dqtdkld lgevndedkpoxcps hdpddk pbd pXKO ok` pbd ko|ldkp ycxon @KO iolcncpopd


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52/61

Lbovpdx Icyd: \teeoxs ) Lgklnt|cgk

Pdevnopd |cplbcka c| ok ckpdaxon voxp gi BCY%? xdydx|d pxok|lxcvpcgk ok` pbd vxgld||

cp|dni dklgtxoad| xdlgehckopcgk tk`dx bdpdxgngagt| lgk`cpcgk+ BCY%? xdlgehckopcgk

c| gh|dxyd` bdk pg adkdpclonns `c|pcklp adkgecl XKO egndltnd| oxd lgvolmoad` ck

pbd |oed ycxcgk ok` |th|dqtdkp ckidlpcgk gi kd ldnn+ @txcka pbd |skpbd|c| gi icx|p

@KO |pxok` "eckt| |pxok`#. xdlgehckgadkcl pdevnopd |cplbcka lok hd pxcaadxd` hs

pbd vxd|dkld gi hxdom"|#. dklgtkpdxcka o vot|d |cpd gx o |dlgk`oxs |pxtlptxd gk pbd

XKO pdevnopd+ Igxld` ok` `skoecl lgvs lbgcld xdlgehckopcgk. vot|d `xcydk

xdlgehckopcgk. olldvpgx ckyo|cgk xdlgehckopcgk hs `glmcka ok` nglmcka eg`dn.

ckpdxolpcyd bocxvck eg`dn ok` bocxvck ck`tld` pXKO ed`copd` "BCPED#

xdlgehckopcgk oxd vxgvg|d` edlbokc|e| igx eckt| |pxok` @KO. bcnd |pxok`

`c|vnoldedkp o||cecnopcgk ok` BCPED eg`dn oxd vxgvg|d` igx vnt| |pxok`

xdlgehckopcgk+ Pbd|d edlbokc|e| oxd kgp etptonns drlnt|cyd ok` onn eos hd

gvdxopcyd ck koptxon lgk`cpcgk+ O lgevxdbdk|cyd vxd|dkpopcgk gi onn pbd ohgyd%

`c|lt||d` edlbokc|e| lgtn` hd acydk o| pbd ignngcka "Ica 4+?#+

Eg|p xdpxgycxon xdlgehckopcgk glltx| `txcka eckt|%|pxok` |skpbd|c| ok` xdncd| gk ho|d

vocxcka hdpddk olldvpgx pdevnopd |dqtdkld| `gk|pxdoe gi xdlgehckopcgk lxg||gydx

|cpd| ok` vxcedx |pxok` |dqtdkld| tkeo|md` hs XKo|d B olpcycps+ O eckgx igxe gi

xdlgehckopcgk lok atc`d ixdd vxcedx |pxok` pdxeckc pg onpdxkopd olldvpgx|+ @txcka pbd

igxld` lgvs lbgcld xdlgehckopcgk pbd ndydn gi ckpdaxcps gi pbd ycxon XKO op pbd

egedkp gi xdydx|d pxok|lxcvpcgk c| kgp mkgk ok` cp c| pbdxdigxd `ciicltnp pg d|pceopd

pbd ixdqtdkls gi igxld` lgvs%lbgcld ck ycyg+ Bgdydx. |pxok` pxok|idx glltx| ck ycpxg

op bcab xopd| ck pbd oh|dkld gi eokcid|p hxdom| gk pbd XKO "ohgtp 8%4 r ?6%0

vdx

ktlndgpc`d#+ \g onpbgtab igxld` lgvs lbgcld c| lxtlcon igx xd|ltd |skpbd|c|. cp|

lgkpxchtpcgk pg adkdpcl yoxcohcncps ecabp kgp hd vxd`geckokp. drldvp bdk ycxon

adkged| oxd lgk|c`dxohns `oeoad`+


53/61

Vot|d |cpd| oxd ckpxck|cl pg pbd XKO |dqtdkld ok` ok o|seedpxs c| xdqtcxd` hdpddk

pbd `gkgx ok` pbd olldvpgx pdevnopd|. hdlot|d pbd gh|polnd pg xdydx|d pxok|lxcvpcgk

et|p hd oh|dkp gk pbd olldvpgx XKO+ \g pxok|idxxcka pbd ko|ldkp @KO gk pbd

olldvpgx pdevnopd cnn kgp |gnyd pbd vxghnde gi xd|pgxcka @KO |skpbd|c|. tknd|| pbc|

|seedpxs c| hxgmdk+ Bgdydx. cp c| `ciicltnp pg ceoackd pbop onn pbd|d xdqtcxdedkp| oxd

xdltxxdkpns itnicnnd` ongka pbd adkgecl XKO+ Xopbdx. vot|d%`xcydk xdlgehckopcgk

ovvncd| pg c|gnopd` lo|d| ok` cp c| kgp o vxdyocncka edlbokc|e+ Pbc| ycd c| |tvvgxpd`

hs pbd vggx lgxxdnopcgk hdpddk xdlgehckopcgk bgp|vgp| ok` pbd vot|d |cpd| gh|dxyd`

ck ycpxg gk ngka XKO|+

Ica 4+?: @coaxoeeopcl vxd|dkpopcgk gi vtpopcyd BCY%? xdlgehckopcgk edlbokc|e+

BCY%?

Xdlgehckopcgk

Eckt| \pxok`

xdlgehckopcgk

Lgvs Lbgcld

Igxld` Lgvs

lbgcld

@skoecl LgvsLbgcld

Vot|d `xcydk

Vxceoxs

\dqtdkld

lgevndrcps

Tkioygxohnd

|txxgtk`ckalgk`cpcgk

XKO |pxtlptxd

`xcydk

xdlgehckopcgk

Olldvpgx

ckyo|cgk eg`dn

CkpdxolpcydBocxvck eg`dn

BCPED

Vnt| \pxok`

Xdlgehckopcgk

\pxok`

@c|vnoldedkp

O||cecnopcgk

Eg`dn

BCPED


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55/61

Lbovpdx \cr

Xdidxdkld


56/61

Lbovpdx \cz: Xdidxdkld

ON @BOBDXC.K+\+. VBCNNCV.V+\+. ABOZOQC.O+. ONC.J+. HDDHC.D+. JOHONNOB.\+O+ ) XCZYC.P+O+

"8663#+ Lxg||%volmoacka gi adkdpclonns `c|pcklp egt|d ok` vxceopd xdpxgycxon XKO|+Xdpxgycxgngas9. 99+

OXCDK.M+M+. YOKBOE.A+ ) OXP\.D+J+ "8661#+ C| BCY%? dygnycka pg o nd|| ycxtndkp igxe ck bteok|7

Kop Xdy Eclxghcgn4. ?0?%4?+

[email protected]+O+. AOG.S+. AONDPPG.X+. [email protected]+. OKPBGKS.X+E+. ACOLGEGKC.Y+. OHXDBO.E+.

@D\PDIOKG. J+ J+. KDAXGKC. E+ ) OXP\. D+ J+ "8669#+ Ckintdkld gi |dqtdkld c`dkpcps ok`tkcqtd hxdomvgckp| gk pbd ixdqtdkls gi ckpdx|thpsvd BCY%? xdlgehckopcgk+ Xdpxgycxgngas


58/61

JDPZP.O+D+. ST.B+. MNOXEOKK.A+J+. XGK.S+. VXD\PGK.H+@+ ) @GTABDXPS.J+V+ "8666#+ Bcab

xopd gi xdlgehckopcgk pbxgtabgtp pbd bteok ceetkg`diclcdkls ycxt| psvd ? adkged+ J Ycxgn

10. ?8


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EHC\O. J+ N+. HOXX. X+. PBGEO\. J+ O+. YOK@DAXOOII. K+. @GXQDCNDX. C+ J+. \YOXGY\MOCO. D+ \+.

HXGQK.Q+N+. EOK\MS.N+E+. AGXDNCLM.X+J+. BOXXC\.X+\+. DKADNEOK.O+ ) VOPBOM.

Y+ M+ "8661#+ Bteok ceetkg`diclcdkls ycxt| psvd ? l@KO| vxg`tld` ck pbd vxd|dkld giOVGHDL


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YOK @DX MTSN.O+L+ ) LGXKDNC\\DK.E+ "8661#+ C`dkpciscka BCY%? `ton ckidlpcgk|+ Xdpxgycxgngas0.

91+

[email protected]+ ) QCNNCOE\GK.L+ "8663#+ BCY egndltnox dvc`decgngas: pxok|ec||cgk ok` o`ovpopcgk

pg bteok vgvtnopcgk|+ Ltxx Gvck BCY OC@\0. 801%48+

QT.Q+. HNTEHDXA.H+E+. IOS.V+J+ ) HOEHOXO.X+O+ "?334#+ \pxok` pxok|idx ed`copd` hs bteokceetkg`diclcdkls ycxt| xdydx|d pxok|lxcvpo|d ck ycpxg c| vxgegpd` hs vot|cka ok` xd|tnp| ck

ec|cklgxvgxopcgk+J Hcgn Lbde816.

Recombination in Human immunodeficiency virus type 1 (HIV-1): A Review

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Transcript of Recombination in Human immunodeficiency virus type 1 (HIV-1): A Review