You are on page 1of 5

$KQVGEJPQNQI[CPF$KQRTQEGUU'PIKPGGTKPI

&1+U

Improved Production of Rifamycin SV by


Amycolatopsis mediterranei MM2 by the Feeding
Method of Carbon Source
=
e~É=gççåÖ=pÜáåI=`Ü~åÖJgççå=háãI=~åÇ=pìåÖ=_~É=háãG=
&GRCTVOGPVQH%JGOKECNCPF$KQNQIKECN'PIKPGGTKPICPF'4+)[GQPIUCPI0CVKQPCN7PKXGTUKV[,KPLW-QTGC =
#DUVTCEV 6JG RWTRQUG QH VJKU TGUGCTEJ YCU VQ UVWF[ VJG GPJCPEGOGPV QH TKHCO[EKP 58
458  [KGNF CEEQTFKPI VQ VJG HGGFKPI
OGVJQFQHCECTDQPUQWTEGHQT^ãóÅçä~íçéëáë=ãÉÇáíÉêê~åÉá//458RTQFWEGFFWTKPIHGTOGPVCVKQPFTQRRGFUJCTRN[
CHVGTTGCEJKPICOCZKOWOCPFKVYCUHQWPFVJCVVJKUVTGPFQH458RTQFWEVKQPTGUWNVGFHTQOUKOWNVCPGQWURTQFWEVKQP
CPFFGITCFCVKQPQH4586QTGFWEG458FGITCFCVKQPFWTKPIKPEWDCVKQPVJGGHHGEVQHECTDQPUQWTEGQPEGNNITQYVJ
YCU KPXGUVKICVGF $CUGF QPVJGTGUWNVUINWEQUG YCUCDGVVGTECTDQPUQWTEGVJCP IN[EGTQNHQTEGNNITQYVJCNVJQWIJ
IN[EGTQNYCUDGVVGTVJCPINWEQUGHQT458RTQFWEVKQPCUTGRQTVGFKPQWTRTGXKQWUUVWF[6QEQPHKTOVJKU458[KGNFCPF
FT[EGNNYGKIJV
&%9 YGTGOGCUWTGFCEEQTFKPIVQVJGHGGFKPIOGVJQFQHECTDQPUQWTEGWUKPIC.UK\GDKQTGCEVQT
4GUWNVU UJQYGF VJCV KPKVKCN EGNN ITQYVJ TCVG CPF 458 [KGNF YGTG UKIPKHKECPVN[ KPETGCUGF TGICTFNGUU QH VJG CFFKVKQP
OGVJQFQHINWEQUGKPVQIN[EGTQNRTQXKFGFINWEQUGYCURTGUGPVKPVJGKPKVKCNUVCIGQHEGNNITQYVJl-5$$

hÉóïçêÇëW=êáÑ~ãóÅáå=psI=#O[EQNCVQRUKUOGFKVGTTCPGKjjOI=ÑÉêãÉåí~íáçåI=Å~êÄçå=ëçìêÅÉI=ÑÉÉÇáåÖ=ãÉíÜçÇ=
=
=
=
=
+0641&7%6+10 RTQZKOCVGN[HQNFYJKEJYCUUKIPKHKECPVN[JKIJGTVJCPVJG
 `HQNFTGRQTVGFGNUGYJGTG=?6JKUJKIJUGNGEVKXKV[KPFK
4KHCO[EKP58
458 DGNQPIUVQVJGCPUCO[EKPITQWRCPF ECVGUVJCV#OGFKVGTTCPGK//RTQFWEGUOCKPN[458'XGP
KUWUGFGZVGPUKXGN[VQVTGCVVWDGTEWNQUKUNGRTQU[CPFUGXGTCN VJQWIJ VJG 458 [KGNFU TGRQTVGF RTGXKQWUN[ YGTG ` VKOGU
QVJGT O[EQDCEVGTKCN KPHGEVKQPU =? #NUQ 458 KU C MG[ KP JKIJGTVJCPVJCVQHQWTOCZKOWO[KGNF=?YGDGNKGXGVJKU
VGTOGFKCVG VJCV ECP DG U[PVJGVKECNN[ EQPXGTVGF KPVQ TKHCO JKIJ UGNGEVKXKV[ ECP EQORGPUCVG HQT VJG NQY [KGNF VQ UQOG
RKEKP TKHCDWVKP CPF TKHCRGPVKPG YJKEJ JCXG OQTG RQYGTHWN FGITGG 5KPEG UGRCTCVKQP QH 458 CPF 4$ CPF EQPXGTUKQP
CEVKXKVKGUVJCP 458=?6JGKPFWUVTKCNUVTCKPEWTTGPVN[WUGF HTQO4$VQ458CTGPQVPGGFGFKH#OGFKVGTTCPGK//KU
KP VJG EQOOGTEKCN HGTOGPVCVKQP RTQEGUU RTQFWEGU QPN[ TKHC WUGFOCP[CFXCPVCIGUOC[GZKUVHQTEQOOGTEKCNRTQFWEVKQP
O[EKP$YJKEJJCUOWEJNGUUCPVKDKQVKECEVKXKV[VJCP458 WUKPIVJKUUVTCKP
6JGTGHQTG KV YQWNF DG GEQPQOKECNN[ CFXCPVCIGQWU VQ FG +PQWTRTGXKQWUUVWF[=?YGQDUGTXGFVJCV458EQPEGP
XGNQRCRTQEGUUWUKPICUVTCKPVJCVECPRTQFWEGOCKPN[458 VTCVKQPTGCEJGFCOCZKOWOCVFC[UQHKPEWDCVKQPCPFVJGP
6JG RTQFWEVKQP [KGNF QH 458 JCU DGGP KORTQXGF D[ VJG FTQRRGFUJCTRN[+VYCUVJQWIJVVJCVVJGFGITCFCVKQPQH458
OWVCVKQPCPFUGNGEVKQPQHUVTCKPU=?VJGCFFKVKQPQHQTICPKE RTQFWEGF FWTKPI HGTOGPVCVKQP OKIJV DG TGNCVGF VQ VJG NQPI
PKVTQIGP EQORQWPFU =? VJG RTGVTGCVOGPV QH UQNKF UWDUVTCVG KPEWDCVKQPRGTKQFVJWUVQTGFWEGVJGKPEWDCVKQPRGTKQFVJG
YKVJ FKNWVG *%N =? VJG WUG QH C HGFDCVEJ EWNVWTG =? CPF KPKVKCNEGNNITQYVJTCVGUJQWNFDGKPETGCUGF6JGOCKPQDLGE
VJGKOOQDKNK\CVKQPQHVJGEGNNUQPINCUUYQQN=?4GEGPVN[ VKXG QH VJKU UVWF[ YCU VQ HKPF ECTDQP UQWTEGU VJCV KPETGCUG
YGEQPFWEVGFUVCVKUVKECNQRVKOK\CVKQPQHOGFKCHQT458RTQ KPKVKCNEGNNITQYVJTCVGCPFHGGFKPIOGVJQFUVJCVKORTQXGVJG
FWEVKQPD[#O[EQNCVQRUKUOGFKVGTTCPGK//WUKPI2NCEMGVV RTQFWEVKQP[KGNFQH458
$WTOCPCPF%GPVTCN%QORQUKVGGZRGTKOGPVCNFGUKIPU=?#V 
VJGQRVKOWORQKPVVJGGZRGTKOGPVCN[KGNFYCU± 
OI.CPF VJG UGNGEVKXKV[QH 458QXGTTKHCO[EKP$YCU CR /#6'4+#.5#0&/'6*1&5
 
%QTTGURQPFKPICWVJQT /KETQQTICPKUOCPF+PQEWNWO2TGRCTCVKQP
6GN   (CZ  
GOCKNUDAMKO"IPWCEMT #OGFKVGTTCPGK//YCUQDVCKPGFEQWTVGU[QH2TQHGUUQT

G


G
*)(NQUU
7PKXGTUKV[QH9CUJKPIVQP5GCVVNG75# +VYCU 
ITQYPHQTJQP;/)OGFKWOOCKPVCKPGFKPIN[E 
GTQNCPFUVQTGFCV−°%6JG;/)OGFKWOEQPVCKPGF
KP 
I. [GCUVGZVTCEVOCNVGZVTCEVINWEQUGCPFVTCEG 
GNGOGPVU6JGVTCEGGNGOGPVUWUGFYGTGCFFGFCV
XX VQ 
VJG HQNNQYKPI EQPEGPVTCVKQPU
KP I.  (G51ŧ*1  
<P51ŧ*1  /P51ŧ*1  %W51ŧ*1  
%Q%Nŧ*1RWTG*51 
 
(NCUM(GTOGPVCVKQP 
 
6JG UGGF EWNVWTG YCU RTGRCTGF D[ KPQEWNCVKPI  μ. QH 
IN[EGTQN UVQEM KPVQ  O. QHINWEQUG[GCUV GZVTCEV OGFKWO 
EQPVCKPKPI
KPI. INWEQUG[GCUVGZVTCEV%C%1 (KI %QPEGPVTCVKQPRTQHKNGUQHTKHCO[EKP58&%9CPFTGUKF
6JGRTQFWEVKQPOGFKC
O.KPO.HNCUMQTO.KP WCN IN[EGTQN QDVCKPGF WUKPI RTGXKQWUN[ FGVGTOKPGF QRVK
O. HNCUM  YGTG KPQEWNCVGF D[ CFFKPI  QT  O. QH  OWOEQPFKVKQPU=?
FC[QNF UGGF EWNVWTG #NN EWNVWTGU YGTG KPEWDCVGF CV °% 
CPFTGXOKPKPCUJCMKPIKPEWDCVQT
,GKQ6GEJ-QTGC   
 FT[ EGNN YGKIJV
&%9  # O. UCORNG QH EWNVWTG DTQVJ
.$KQTGCEVQT(GTOGPVCVKQP  VCMGPHTQOVJGHNCUMYCUOKZGFYKVJO.QH0*%NVQ
 FKUUQNXGVJGTGOCKPKPI%C%1#HVGTGCEJQHVJTGGEQPUGEW
6JG UGGF EWNVWTG YCU RTGRCTGF D[ KPQEWNCVKPI  μ. QH VKXGEGPVTKHWICVKQPUHQTOKPCVTGXOKPCPFYCUJGU
IN[EGTQN UVQEM KPVQ  O. QH VJG OGFKWO RTGXKQWUN[ FG YKVJ FKUVKNNGF YCVGT VJG UWRGTPCVCPV YCU FKUECTFGF CPF VJG
UETKDGF6YQ.RTQFWEVKQPOGFKWOKP.UVKTTGFDKQTGCEVQT EGNNRGNNGVYCUFTKGFCV°%HQTJ 

$KQ6TQP-QTGC YCUKPQEWNCVGFYKVJO.QHFC[QNF 
UGGF EWNVWTG 6JG QRGTCVKPI EQPFKVKQPU YGTG  TGXOKP 
°% R*  CPF CGTCVKQP  XXO 6JG R* YCU EQPVTQNNGF 4'57.65#0&&+5%755+10
WUKPI00C1*CPF0*51UQNWVKQPUCPF&1EQPEGP 
VTCVKQP YCU PQV EQPVTQNNGF DGECWUG KV YCU OCKPVCKPGF CV &GITCFCVKQPQH458
ITGCVGT VJCP  FWTKPI VJG GPVKTG KPEWDCVKQP RGTKQF 6JG 
OGFKWO EQPVCKPGF
KP I.  ECTDQP UQWTEG
INWEQUG CPFQT (KI  UJQYU VJG EQPEGPVTCVKQP RTQHKNGU HQT 458 &%9
IN[EGTQN [GCUVGZVTCEV#NUQI.QH2')
RQN[ CPFTGUKFWCNIN[EGTQNQDVCKPGFCVQWTRTGXKQWUN[FGVGTOKPGF
GVJ[NGPGIN[EQN YCUWUGFVQRTGXGPVHQCOHQTOCVKQP QRVKOWO EQPFKVKQP KP C O. HNCUM 7PFGT VJGUG EQPFK
6TCEG GNGOGPVU YGTG CFFGF CV 
XX  QH OGFKWO VQ CNN VKQPUVJGEQPEGPVTCVKQPUKPOGFKWOYGTGIN[EGTQNI.
HGTOGPVCVKQP GZRGTKOGPVU #NN HGTOGPVCVKQP TGUWNVU CTG RTG [GCUVGZVTCEVI.%C%1I.CPFVTCEGGNGOGPVU
UGPVGFCUVJGCXGTCIGQHVYQGZRGTKOGPVUYKVJUVCPFCTFFG
XX  6JG OCZKOWO EQPEGPVTCVKQP QH 458 TGCEJGF CR
XKCVKQPU     RTQZKOCVGN[  OI. CV  FC[U OCKPVCKPGF HQT  OQTG
 FC[UCPFVJGPUJCTRN[FGETGCUGFVQCDQWVJCNHQHVJGOCZK
#PCN[UKU OWOEQPEGPVTCVKQPCVFC[U6JGUCOGVTGPFQHTKHCO[EKP
 RTQFWEVKQPYCUQDUGTXGFD[-TKUJPCGVCN=? 
6JG EQPEGPVTCVKQPU QH INWEQUG CPF IN[EGTQN YGTG OGCU (KIUJQYUVJGFGITCFCVKQPGZRGTKOGPVQHUVCPFCTF458
WTGFD[*2.%VJG*2.%U[UVGOEQPUKUVGFQH5RGEVTC5[U CV VJG KPEWDCVKQP VGORGTCVWTG °% #U UJQYP KP (KI 
VGO 2 RWOR 5JQFGZ 4+ FGVGEVQT CPF $KQ4CF 458 FGETGCUGF UNKIJVN[ WPVKN  FC[U CPF VJGP FGETGCUGF
*2:*EQNWOP
EQPFKVKQPUO.OKP°%/ UJCTRN[6JGTGHQTGVJGOCKPVGPCPEGQH458RTQFWEGFDGHQTG
*51  4KHCO[EKP 58 CPF $ YGTG OGCUWTGF D[ C *2.% CPFCHVGTFC[UCVOCZKOWONGXGNHQT`FC[UCPFVJGP
U[UVGOVJCVEQPUKUVGFQHC.CD#NNKCPEG2RWOR
#RRNG VJGUJCTRFGETGCUGUJQYPKP(KIYCUCVVTKDWVGFVQFGITCFC
8CNNG[ /0 75#  YKVJ C 5RGEVTC  78 FGVGEVQT UGV CV VKQPQH458KVUGNHPQVWVKNK\CVKQPD[EGNNUCUCECTDQPUQWTEGU
 PO # TGXGTUGFRJCUG 2JGPQOGPGZ .WPC % EQNWOP 5KPEG IN[EGTQN EQPEGPVTCVKQP EQPVKPWQWUN[ FGETGCUGF FWTKPI

RCTVKENGUK\GμOOOªEO YCUWUGFCVTQQOVGO VJGNGXGNQHHRGTKQFQH458RTQFWEVKQPCPF458EQPEGPVTC


RGTCVWTG YKVJ C OQDKNG RJCUG QH  / COOQPKWO HQT VKQPKUFGVGTOKPGFD[VJGFKHHGTGPEGDGVYGGPRTQFWEVKQPCPF
OCVGCEGVQPKVTKNG =
XX ? 6JG 2GCM5KORNG %JTQOC FGITCFCVKQP458KUEQPVKPWQWUN[RTQFWEGFCPFFGEQORQUGF
VQITCRJ[ &CVC 5[UVGO
54+ +PUVTWOGPVU 6QTTCPEG %# 5WEJ UKOWNVCPGQWU RTQFWEVKQP CPF FGEQORQUKVKQP YCU CNUQ
75#  YCU WUGF HQT FCVC CESWKUKVKQP CPF CPCN[UKU #NN UVCP TGRQTVGF KP VJG RTQFWEVKQP QH βNCEVCO CPVKDKQVKEU =?
FCTF EJGOKECNU YGTG RWTEJCUGF HTQO 5KIOC %JGOKECN %Q 6JGTGHQTGVQTGFWEGVJGFGITCFCVKQPQH458RTQFWEGFFWT

5V.QKU/175# CPFTKHCO[EKP$UVCPFCTFYCUQDVCKPGF KPI KPEWDCVKQP KV KU KORQTVCPV VQ KPETGCUG VJG KPKVKCN ITQYVJ
EQWTVGU[QH2TQHGUUQT%JWP-[W-KO
+PLG7PKXGTUKV[-KO TCVGYJKEJKPVWTPFGETGCUGUVJGKPEWDCVKQPRGTKQF6JGFGI
JCG-QTGC  TCFCVKQPTCVGQH458CV°%YCUOWEJJKIJGTVJCPVJCVQH
6JG EGNN EQPEGPVTCVKQP YCU FGVGTOKPGF D[ OGCUWTKPI VJG 458CV°%UQVJGFGITCFCVKQPUGGOUVQDGXGT[UGPUKVKXG 
$KQVGEJPQN$KQRTQEGUU'PI=

G
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
(KI &GITCFCVKQP QH RWTG TKHCO[EKP 58 CV FKHHGTGPV VGORGTC (KI'HHGEVQHIN[EGTQNHGGFKPIVKOGQPTKHCO[EKP58RTQFWEVKQP
VWTGU
°%  
 
 I.YCUCRRTQZKOCVGN[VKOGUVJCVYKVJIN[EGTQNI.
 QHINWEQUGYCUUGNGEVGFHQTHWTVJGTUGGFEWNVWTGRTGRCTCVKQP 
 5KPEG INWEQUG YCU OQTG GHHGEVKXG HQT EGNN ITQYVJ VJCP
 IN[EGTQN CPF IN[EGTQN YCU OQTG GHHGEVKXG HQT 458 RTQFWE
 VKQPVJCPINWEQUGYGEQPFWEVGFCRTGNKOKPCT[GZRGTKOGPVVQ
 FGVGTOKPGVJGQRVKOWOVKOGVQCFFIN[EGTQNYJGPUVCTVKPIC
 EWNVWTGYKVJINWEQUGHQNNQYGFD[IN[EGTQNCFFKVKQPKPC
 O.HNCUM
(KI +PKVKCNN[I.QHINWEQUGYCUCFFGFCPF
 VJGPI.QHIN[EGTQNYCUCFFGFCHVGTQTFC[U6JG
 VQVCNECTDQPEQPEGPVTCVKQPWUGFYCUVJGUCOGCUVJGQRVKOWO
 IN[EGTQNEQPEGPVTCVKQPFGVGTOKPGFKPQWTRTGXKQWUUVWF[=?
 #HVGTFC[UVJGCOQWPVQHINWEQUGTGOCKPKPIYCUCDQWV
 I. VJG INWEQUG YCU EQORNGVGN[ EQPUWOGF CHVGT ` FC[U
 9JGPIN[EGTQNYCUCFFGFCVFC[UVJG458[KGNFYCU
 OI.YJKEJYCUCDQWVJKIJGTVJCPVJG[KGNFQDVCKPGF
 YJGPQPN[IN[EGTQNYCUWUGFCUVJGECTDQPUQWTEGCUUJQYP
 KP (KI  9JGP IN[EGTQN YCU CFFGF CV  CPF  FC[U 458
(KI 'HHGEVQHIN[EGTQNQTINWEQUGEQPEGPVTCVKQPQPEGNNITQYVJ [KGNFU FGETGCUGF UKIPKHKECPVN[ 6JKU EQWNF DG FWG VQ VJG
QH^K=ãÉÇáíÉêê~åÉá// UJQTVCIGQHECTDQPUQWTEGDGECWUGIN[EGTQNYCUCFFGFCVVJG
 UVCIGYJGPINWEQUGYCUCNOQUVGZJCWUVGF   
 
VQVGORGTCVWTG   (GGFKPI/GVJQF
 
'HHGEVQH%CTDQP5QWTEG  5KPEGINWEQUGYCUCDGVVGTECTDQPUQWTEGHQTEGNNITQYVJ
 VJCP IN[EGTQN VJG HGGFKPI OGVJQF VJCV KPETGCUGU KPKVKCN EGNN
6Q HKPF C ECTDQP UQWTEG VJCV UVKOWNCVGU VJG KPKVKCN EGNN ITQYVJTCVGKP.DKQTGCEVQTYCUKPXGUVKICVGF(KTUVCEQP
ITQYVJTCVGVJGGHHGEVQHECTDQPUQWTEGQPEGNNITQYVJYCU VTQNGZRGTKOGPVKPYJKEJQPN[IN[EGTQNYCUWUGFCUCECTDQP
KPXGUVKICVGF )GPGTCNN[ CP GHHGEVKXG ECTDQP UQWTEG HQT EGNN UQWTEG YCU ECTTKGF QWV CV VJG RTGXKQWUN[ FGVGTOKPGF QRVK
ITQYVJ KU FKHHGTGPV HTQO VJCV HQT OGVCDQNKVG RTQFWEVKQP OWO EQPFKVKQP #U UJQYP KP (KI  VJG JKIJGUV 458 CPF
=?+PQWTRTGXKQWUTGUGCTEJ=?IN[EGTQNYCUOQTGGH &%9YJKEJYGTGQDUGTXGFCVCRRTQZKOCVGN[CPFFC[U
HGEVKXG VJCP INWEQUG CV KPETGCUKPI 458 [KGNF *QYGXGT KV YGTG  OI. CPF  I. TGURGEVKXGN[ 5KPEG 458 CPF
YCU VJQWIJV VJCV INWEQUG YQWNF DG OQTG GHHGEVKXG HQT EGNN &%9 XCNWGU YGTG OQPQVQPQWUN[ CRRTQCEJGF VJGKT OCZK
ITQYVJ VJCP IN[EGTQN (KI  UJQYU VJG GHHGEV QH INWEQUG QT OWO XCNWGU VJG CXGTCIG OCZKOWO XCNWGU UJQYP KP ITCRJ
IN[EGTQNQPEGNNITQYVJKPCO.HNCUM;GCUVGZVTCEVCPF YGTGUGNGEVGFHQTEQORCTKUQP 
%C%1YGTGCPFI.TGURGEVKXGN[#UIN[EGTQNEQPEGP 6Q CFF INWEQUG KPVQ VJG EWNVWTG OGFKWO VYQ FKHHGTGPV
VTCVKQP YCU KPETGCUGF &%9 EQPVKPWQWUN[ KPETGCUGF DWV CU HGGFKPIOGVJQFUYGTGFGXKUGF
 UKOWNVCPGQWUCFFKVKQPQH
INWEQUG EQPEGPVTCVKQP YCU KPETGCUGF &%9 UJCTRN[ KP INWEQUGCPFIN[EGTQN
 INWEQUGCFFKVKQPHKTUVHQNNQYGFD[
ETGCUGF CPF VJGP FGETGCUGF CHVGT  I. 6JG FGETGCUG QH CFFKVKQPQHIN[EGTQNCHVGTFC[U+PVJGHKTUVOGVJQFCNOQUV
&%9CDQXGI.QHINWEQUGYQWNFDGFWGVQKPJKDKVKQPQH VJGUCOGCOQWPVQHINWEQUG
I. CPFIN[EGTQN
I. 
EGNNITQYVJD[INWEQUG5KPEG&%9XCNWGYKVJINWEQUGCV YGTGUKOWNVCPGQWUN[CFFGFVQVJGTGCEVQTKPUVGCFQHI.


G
 
 
 
 
 
 
 
 
 
 
 
 
(KI %QPEGPVTCVKQPRTQHKNGUQHTKHCO[EKP58&%9CPFTGUKF (KI %QPEGPVTCVKQPRTQHKNGUQHTKHCO[EKP58&%9CPFTGUKF
WCN IN[EGTQN KP C . HGTOGPVGT
ECTDQP UQWTEG  I. WCN INWEQUG CPF IN[EGTQN KP C . HGTOGPVGT
ECTDQP
IN[EGTQN  UQWTEGI.INWEQUGCFFGFKPKVKCNN[CPFI.IN[E
 GTQNCFFGFCHVGTFC[U 
 
 
 YCUCNOQUVVJGUCOG6JKUKURTQDCDN[TGNCVGFVQVJGTGUKFWCN
 IN[EGTQN COQWPV CV VJG NCVGT UVCIG QH VJG KPEWDCVKQP +P QWT
 RTGXKQWUUVWF[=?KVYCUKNNWUVTCVGFVJCVIN[EGTQNYCUOQTG
 GHHGEVKXG VJCP INWEQUG HQT 458 RTQFWEVKQP 5KPEG VJG IN[E
 GTQN EQPEGPVTCVKQP CV NCVGT UVCIG KP VJG UGEQPF OGVJQF YCU
 OCKPVCKPGFCVTGNCVKXGN[JKIJGTNGXGNUVJCPKPVJGHKTUVOGVJQF
 KVKUVJQWIJVVJCVVJG458[KGNFGSWKXCNGPVVQVJCVQHVJGHKTUV
 OGVJQF YCU RTQFWEGF CV  NQYGT &%9 VJCP VJCV QH VJG
 HKTUV OGVJQF 6JGTGHQTG KPKVKCN EGNN ITQYVJ TCVG CPF 458
 [KGNFECPDGUKIPKHKECPVN[KPETGCUGFTGICTFNGUUQHVJGCFFKVKQP
 OGVJQFQHINWEQUGKPVQIN[EGTQNRTQXKFGFINWEQUGKURTGUGPV
(KI %QPEGPVTCVKQPRTQHKNGUQHTKHCO[EKP58&%9CPFTGUKF FWTKPIVJGKPKVKCNUVCIG 
WCN INWEQUG CPF IN[EGTQN KP C . HGTOGPVGT
ECTDQP 9JGP VJG GHHGEV QH INWEQUG QP EGNN ITQYVJ YCU KPXGUVK
UQWTEGI.INWEQUG I.IN[EGTQNCFFGFUKOWN ICVGF CU UJQYP KP (KI  VJG OCZKOWO EGNN EQPEGPVTCVKQP
VCPGQWUN[  YCUQDUGTXGFCVI.QHINWEQUG6JGTGHQTGVJKUEQPEGPVTC
 VKQP YCU UGNGEVGF HQT KPKVKCN UVCTVWR INWEQUG EQPEGPVTCVKQP
 *QYGXGTVJGTGYQWNFDGUQOGKPVGTCEVKQPUDGVYGGPINWEQUG
QHIN[EGTQN(KIUJQYUVJCVCVFC[UVJGJKIJGUV458CPF CPF IN[EGTQN 6Q HKPF VJKU KPVGTCEVKQP VJG CRRNKECVKQP QH TG
&%9 XCNWGU YGTG  OI. CPF  I. TGURGEVKXGN[ URQPUGUWTHCEGOGVJQFVQHKPFVJGQRVKOCNTCVKQQHIN[EGTQNVQ
9JGPVJGUGXCNWGUYGTGEQORCTGFYKVJVJGXCNWGUQHEQPVTQN INWEQUG KP EWNVWTG OGFKWO YQWNF DG PGGFGF #FFKVKQPCNN[
458[KGNFCPF&%9YGTGKPETGCUGFD[CPFTGURGE VJG FGXGNQROGPV QH UWKVCDNG HGGFKPI UVTCVGI[ HQT HGFDCVEJ
VKXGN[5KPEGVJGKPKVKCNITQYVJTCVGYCUUKIPKHKECPVN[JKIJGT EWNVKXCVKQPCNUQEQWNFHWTVJGTKORTQXGVJGRTQFWEVKXKV[
VJCP VJCV QH EQPVTQN VJG KPEWDCVKQP VKOG VQ OCZKOWO 458 
EQPEGPVTCVKQPYCUCNUQFGETGCUGFD[FC[U6JWUINWEQUGKP 
IN[EGTQN UVKOWNCVGU KPKVKCN EGNN ITQYVJ CPF EQPUGSWGPVN[ KP 4GEGKXGF(GDTWCT[CEEGRVGF#RTKN
ETGCUGU458[KGNF#NUQINWEQUGCPFIN[EGTQNCRRGCTVQDG 
EQPUWOGF UKOWNVCPGQWUN[ D[ EGNNU DGECWUG VJG FKHHGTGPEG 
DGVYGGPINWEQUGCPFIN[EGTQNEQPEGPVTCVKQPUCVCP[KPEWDC 4'('4'0%'5
VKQPVKOGFQGUPQVEJCPIGOWEJYKVJVKOG   
+P VJG UGEQPF OGVJQF  I. QH INWEQUG YCU CFFGF KPK  (NQUU * ) CPF 69 ;W
  4KHCO[EKPOQFG QH
VKCNN[CPFVJGPI.QHIN[EGTQNYCUCFFGFCHVGTFC[U CEVKQP TGUKUVCPEG CPF DKQU[PVJGUKU %JGO 4GX 
6JG JKIJGUV 458 CPF &%9 YJKEJ YGTG QDUGTXGF CV CR 
RTQZKOCVGN[  CPF  FC[U YGTG  OI. CPF  I.  5CPHKNKRRQ#%&GNNC$TWPC./CTUKNK'/QTXKNNQ
TGURGEVKXGN[
(KI 6JGKPKVKCNITQYVJTCVGWUKPIOGVJQF % 4 2CUSWCNWEEK ) 5EJKQRRCECUUK CPF & 7PIJGTK
YCUNQYGTVJCPVJCVWUKPIOGVJQFDWVJKIJGTVJCPVJCVQH
 $KQNQIKECNCEVKXKV[QHCPGYENCUUQHTKHCO[EKPU
EQPVTQN #NUQ VJG KPEWDCVKQP VKOG HQT OCZKOWO &%9 YCU 5RKTQRKRGTKF[NTKHCO[EKPU,#PVKDKQV
CDQWV  FC[ NQPIGT VJCP VJCV QH EQPVTQN 6JKU ECP DG GZ  /WTCNK-TKUJPC25)8GPMCVGUYCTNWCPF.84CQ
RNCKPGFD[VJGNCIVKOGPGGFGFHQTEGNNUVQCEENKOCVG VQVJG
 2TQFWEVKQPQHTKHCO[EKP58WUKPIOWVCPVUVTCKPU
UWDUVTCVG VTCPUKVKQP HTQO INWEQUG VQ IN[EGTQN %QORCTGF VQ QH #O[EQNCVQRUKU OGFKVGTTCPGK /6%% 9QTNF , /K
OGVJQF&%9YCUFGETGCUGFD[CDQWVDWV458[KGNF  ETQDKQN$KQVGEJPQN
$KQVGEJPQN$KQRTQEGUU'PI=

G
 /WTCNK -TKUJPC 2 5 ) 8GPMCVGUYCTNW CPF . WUKPI #O[EQNCVQRUKU OGFKVGTTCPGK 9QTNF , /KETQDKQN
8GPMCVGUYCT 4CQ
  'HHGEV QH WTCEKN QP TKHCO[EKP $KQVGEJPQN
58RTQFWEVKQPD[#O[EQNCVQRUKUOGFKVGTTCPGK/84  &GUJRCPFG # & - ) $CJGVK CPF 0 4 %JCVVGTLGG
.GVV#RRN/KETQDKQN
  &GITCFCVKQP QH βNCEVCO CPVKDKQVKEU %WTT 5EK
 -TKUJPC25/)8GPMCVGUYCTNW#2CPFG[CPF. 
84CQ
 $KQU[PVJGUKUQHTKHCO[EKP58D[#O[EQ  /C[GT#(CPF9&&GEMYGT
 5KOWNVCPGQWU
NCVQRUKU OGFKVGTTCPGK /6%% KP UQNKF EWNVWTGU RTQFWEVKQPCPFFGEQORQUKVKQPQHENCXWNCPKECEKFFWTKPI
$KQVGEJPQN#RRN$KQEJGO 5VTGRVQO[EGUENCXWNKIGTWUEWNVKXCVKQPU#RRN/KETQDKQN
 'N'PUJCU[*#7+$GUJC[#+'N&KYCP[*/ $KQVGEJPQN
1OCT # ) 'N-JQN[ CPF 4 'N0CLCT
  +O  %JQY;4.K,9W5/2WCJ590GY93
RTQXGOGPV QH TKHGO[EKPU RTQFWEVKQP D[ #O[EQNCVQRUKU %JKC(.KG6//4CJOCPCPF9,%JQK
 
OGFKVGTTCPGK KP DCVEJ CPF HGFDCVEJ EWNVWTGU #EVC /K /QFGNKPI CPF QRVKOK\CVKQP QH OGVJCPQN CU C EQUQNXGPV
ETQDKQN2QN KPCOQZKEKNNKPU[PVJGUKUCPFKVUCFXCPVCIGQXGTGVJ[NGPG
 #DW5JCF[/4#+'N&KYCP[/#(CTKFCPF* IN[EQN$KQVGEJPQN$KQRTQEGUU'PI
# 'N'PUJCU[
  5VWFKGU QH TKHCO[EKP RTQFWEVKQP  6CQ , . : & 9CPI ; . 5JGP CPF & < 9GK
D[ #O[EQNCVQRUKU OGFKVGTTCPGK EGNNU KOOQDKNK\GF QP
  5VTCVGI[ HQT VJG KORTQXGOGPV QH RTQFKIKQUKP
INCUUYQQN,$CUKE/KETQDKQN RTQFWEVKQP D[ C 5GTTCVKC OCTEGUEGPU OWVCPV VJTQWIJ
 5JKP*,%,-KOCPF5$-KO
 1RVKOK\C HGFDCVEJHGTOGPVCVKQP9QTNF,/KETQDKQN$KQVGEJPQN
VKQPQHEWNVWTGOGFKWOHQTTKHCO[EKP58RTQFWEVKQPD[ 
#O[EQNCVQRUKU OGFKVGTTCPGK // WUKPI UVCVKUVKECN FG  5JK ( < :W CPF 2 %JGP
  1RVKOK\CVKQP QH γ
UKIPU$KQVGEJPQN$KQRTQEGUU'PI RQN[INWVCOKECEKFRTQFWEVKQPD[$CEKNNWUUWDVKNKU<,7
 -TKUJPC 2 5 / ) 8GPMCVGUYCTNW CPF . 8 4CQ WUKPICUWTHCEGTGURQPUGOGVJQFQNQI[$KQVGEJPQN$KQ

 5VWFKGUQPHGTOGPVCVKXGRTQFWEVKQPQHTKHCO[EKP  RTQEGUU'PI

You might also like