Fumigant efficacy of 1,8-cineole and eugenol on the pupal stage of
Tribolium castaneum
(Herbst)
(Insecta:
Coleoptera:
Tenebrionidae)
Fumigantna učinkovitost 1,8-cineola i eugenola na stadij kukuljice
kestenjastog brašnara
Tribolium castaneum
(Herbst) (Insecta:
Coleoptera: Tenebrionidae)
Liška, A., Rozman, V., Brmež, M., Rebekić, A., Lucić, P.
Poljoprivreda/Agriculture
ISSN: 1848-8080 (Online)
ISSN: 1330-7142 (Print)
DOI:
http://dx.doi.org/10.18047/poljo.21.2.4
'2,SROMR
)80,*$17()),&$&<2)&,1(2/($1'(8*(12/21
7+(383$/67$*(2)
75,%2/,80&$67$1(80
+(5%67
,16(&7$&2/(237(5$7(1(%5,21,'$(
/LåND$5R]PDQ9%UPHç05HEHNLþ$/XFLþ3
2ULJLQDOVFLHQWLILFSDSHU
,]YRUQL]QDQVWYHQLĀODQDN
6800$5<
7KHIXPLJDQWHIILFDF\RIFLQHROHDQGHXJHQROFRPSRXQGVZDVWHVWHGRQWKHSXSDO VWDJHRIWKHUHGIORXUEHHWOH7ULEROLXPFDVWDQHXP+HUEVW(IIHFWVRIWKRVHFRPSRXQGV ZHUHGHWHUPLQHGDVPRUWDOLW\DVZHOODVLPSDFWRQGHYHORSPHQWRIWUHDWHGPDOHDQG IHPDOHSXSDH$OVRGLIIHUHQFHVLQVHQVLWLYLW\WRWKHWHVWHGFRPSRXQGVEHWZHHQVH[HV RI7FDVWDQHXPSXSDHZHUHDVVHVVHG&RPSRXQGVZHUHWHVWHGDWGRVHVUDWH DQGPO/YRODW&DQGUKGXULQJK&RPSRXQGVWR[LFLW\ZDV
H[SUHVVHGLQWZRDFWLRQPRGHVOHWKDODQGDVDQLPSHGLPHQWWRQRUPDOPHWDPRUSKRVLV RI SXSDH LQWR DGXOW VWDJH IRUPLQJ ´DGXOWRLGVµ DQG GHIRUPHG DGXOWV DW ERWK VH[HV 0DOHSXSDHZHUHJHQHUDOO\PRUHVHQVLWLYHWRERWKWHVWHGFRPSRXQGV2YHUDOOEHWWHUHIIL FLHQF\ZDVSHUIRUPHGZLWKFLQHROHZKLOHHXJHQROKDGQRWDFFRPSOLVKHGSURPLVLQJ HIIHFWRQWKHWHVWHGSXSDH
.H\ZRUGVIXPLJDWLRQFLQHROHHXJHQRO7ULEROLXPFDVWDQHXPSXSDH
,1752'8&7,21
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
7KH UHDVRQV RI SKRVSKLQH UHVWULFWLRQ WR EHFRPH FRPSOHWH UHSODFHPHQW RI PHWK\O EURPLGH DUH H[SR VXUH WLPH DQG WHPSHUDWXUH 'XFRP DV ZHOO DV
LQFUHDVLQJ UHVLVWDQW SRSXODWLRQV DPRQJ VWRUDJH SHVWV 0DKURRIDQG+DJVWUXP
7KHRWKHUSRVVLEOHDOWHUQDWLYHLVVXOSKXU\OIOXRULGH +RZHYHU WKHUH DUH DOVR OLPLWDWLRQV ILUVW DQG IRUHPRVW ORZHU DFWLYLW\ DW WKH LQVHFW HJJ VWDJH DQG SVRFLGV $WKDQDVVLRXHWDOLQFOXGLQJSUREOHPVRIUHVLGXHV LQIRRG'XFRP$OOWKHWKLQJVFRQVLGHUHGWKHUH LVDQXUJHQWQHHGIRUQHZFRJQLWLRQRISHVWFRQWUROPHW KRGVZKLFKZRXOGEHDGHTXDWHO\HIILFLHQWDQGVLPLODUO\ UHGXFHDGYHUVHHIIHFWRIV\QWKHWLFVXEVWDQFHVLQVWRUHG SURGXFWVSURWHFWLRQ%HVLGHVWKHUHLVDVWURQJLQWHUHVW LQ WKH UHVHDUFK LQWR SODQW SURGXFWV HIILFLHQF\ DSURSRV RI HVVHQWLDO RLOV DQG WKHLU FRPSRXQGV IRU VWRUHG SHVW FRQWURO %RWDQLFDO LQVHFWLFLGHV VKRZ GLIIHUHQW HIIHFWV RQSHVWVZKLFKFRXOGGLPLQLVKFURVVUHVLVWDQFH,VPDQ PRUHRYHU WKH\ DUH UHOHYDQW SDUW RI LQWHJUDWHG SURWHFWLRQZLWKLQRUJDQLFSURGXFWLRQ(EDGROODKL ,W KDV EHHQ IRXQG RXW WKDW HVVHQWLDO RLOV DQGRU WKHLU FRPSRXQGVKDYHGLIIHUHQWHIIHFWRQSHVWVOLNHZLVHFRQ
$/LåNDHWDO)80,*$17()),&$&<2)&,1(2/($1'(8*(12/217+(383$/67$*(
WDFW UHSHOOHQW DQWLIHHGDQW DQG IXPLJDQW RU LQKLELWRU\ WR HJJV OD\LQJ $UDEL HW DO %HQ]L HW DO %HVLGHV WKH\ KDYH QHXURWR[LF HIIHFW LQGXFLQJ V\PS WRPV VLPLODU WR WKRVH SURGXFHG E\ RUJDQRSKRVSKDWHV DQGFDUEDPDWHEDVHGLQVHFWLFLGHV,VPDQ
0RQRWHUSHQRLGFLQHROHKDYHDSURPLVLQJSRWHQ WLDOIRUXVHDVDIXPLJDQWLQGXFLQJYHU\JRRGUHVXOWVLQ WKHFRQWURORIVWRUHGSURGXFWVLQVHFWV/HHHWDO $XWKRUV*XRHWDOUHSRUWHGWKDWFLQHROHDORQJ ZLWK VRPH RWKHU HVVHQWLDO RLO FRPSRQHQWV RI(WOLQJHUD \XQQDQHQVLV UKL]RPHV H[KLELWHG VWURQJHU FRQWDFW WR[L FLW\WKDQɴFDU\RSK\OOHQHDJDLQVW7FDVWDQHXPDGXOWV 6LQFHLQRXUSUHYLRXVVWXG\/LåNDHWDOFLQH ROH ZDV IDU WKH PRVW HIIHFWLYH FRPSRXQG IROORZHG E\ FDPSKRUDQGHXJHQRODJDLQVW7FDVWDQHXPDGXOWVZH VHOHFWHGFLQHROHDQGHXJHQROIRUWHVWLQJWKHLUHIILFD F\RQWKHLPPDWXUHVWDJHRI7FDVWDQHXP
:LWK WKLV H[SHULPHQW D IXPLJDQW HIILFDF\ RI FLQHROHDQGHXJHQROFRPSRXQGVZHUHWHVWHGDJDLQVW WKHSXSDOVWDJHRIWKHUHGIORXUEHHWOH$OVRGLIIHUHQFHV LQVHQVLWLYLW\EHWZHHQVH[HVRI7FDVWDQHXPSXSDHWR WKHWHVWHGFRPSRXQGVZHUHGHWHUPLQHG
0$7(5,$/$1'0(7+2'6 (VVHQWLDORLOFRPSRXQGV
8VHGFRPSRXQGVFLQHROH*&DQGHXJHQRO *&ZHUHSXUFKDVHGIURPWKHSURGXFHUVÅ6LJPD $OGULFK´*HUPDQ\DQGÅ)OXND´6*6ZLW]HUODQG ,QVHFWVUHDULQJ
7HVWLQVHFWVRI7FDVWDQHXPVSHFLHVZHUHUHDUHG XQGHU WKH FRQWUROOHG FRQGLWLRQV DW & UHODWLYHKXPLGLW\UKLQGDUNQHVVRQDPL[WXUHRIKDUG ZKHDWIORXUDQGGU\\HDVWDWWKHUDWLR/LXDQG+R 6H[LQJSXSDH/\RQZDVSHUIRUPHGZLWK VWHUHR]RRP ORXSH ZLWK GLJLWDO FDPHUD DQG VRIWZDUH 2O\PSXV6=;ZKHQWKHSXSDHZHUHGD\VROG )XPLJDQWWR[LFLW\RIFLQHROHDQGHXJHQRO
)XPLJDWLRQZDVSHUIRUPHGLQWKHJODVVMDUVRI PO YROXPH ILOOHG ZLWK ZKHDW JUDLQ RFFXS\LQJ RI WKHMDUVYROXPH7ZHQW\VH[HGSXSDHZHUHSXWLQWRWKH VLON PHVK FDJHV DQG DOWRJHWKHU SODFHG LQ MDUV LQ IRXU UHSHWLWLRQV&RPSRXQGVFLQHROHDQGHXJHQROZHUH DSSOLHGZLWK´.DUWHOOµPLFURSLSHWWHRQILOWHUSDSHUDWWDF KHGWRWKHOLGVRIWKHJODVVFRQWDLQHUVWLJKWO\VHDOHGDQG NHSWXQGHUFRQWUROOHGFRQGLWLRQVDW&UK LQGDUNQHVVIRUKRXUV7KHFRPSRXQGVZHUHWHVWHGDW GRVHUDWHVDQGPO/YRO7KHFRQWURO
ZDVFDUULHGRXWXQGHUWKHVDPHFRQGLWLRQVRQO\ZLWKRXW RLODSSOLFDWLRQ,QRUGHUWRDVVHVVIXPLJDQWHIILFDF\RIWKH WHVWHGFRPSRXQGVRQWKHSXSDOVWDJHLQVHFWPRUWDOLW\ DQG JURZWK DFWLYLW\ KDYH EHHQ GHWHUPLQHG IROORZLQJ WKH PRGLILHG VFDOH DFFRUGLQJ WR 0DQGDYD %\ WKDW VFDOH D QXPEHU RI GHDG XQLWV LQ WKH SXSDO VWDJH GHYHORSPHQWDOVWDJHDQXPEHURIDGXOWRLGVGHYH ORSPHQWDO VWDJH OLYH DQG GHDG GHIRUPHG DGXOWV GHYHORSPHQWDOVWDJHGHYHORSHGIURPWKHWUHDWHGVXU
YLYHGSXSDHDQGDQXPEHURIQRUPDOO\GHYHORSHGDGXOWV GHYHORSPHQWDO VWDJH ZLWKRXW GHIRUPDWLRQV ZHUH GHILQHG,QWHUPHGLDWHIRUPRU´DGXOWRLGµKDVDFKDUDFWH ULVWLFDSSHDUDQFHZLWKWKHIURQWSDUWRIWKHERG\OLNHZLVH DGXOWDQGLVSLJPHQWHGZLWKZLGHVSUHDGIRUHZLQJVDQG KLQGZLQJVLIGHYHORSHGZKLOHDEGRPHQKDVDW\SLFDO DSSHDUDQFHRISXSDHDQGLVQRQSLJPHQWHG
'DWDDQDO\VLV
)XPLJDQW HIILFDF\ GDWD ZHUH SURFHVVHG E\ VWD WLVWLFDODQDO\VLVV\VWHP6$667$76RIWZDUH ,Q RUGHU WR GHWHUPLQH GLVWULEXWLRQ GHYLDWLRQV D .ROPRJRURY6PLUQRY WHVW ZDV XVHG LQ PRGXOH 6$6 ,QWHUDFWLYH'DWD$QDO\VLV2QHZD\DQDO\VLVRIYDULDQFH RI WKH WHVWHG YDULDEOHV ZDV VXEMHFWHG WR 6$6 $QDO\VW PRGXOH DQG D SURFHGXUH $129$ ZDV XVHG 7XNH\·V 6WXGHQWL]HG5DQJH+6'WHVWZDVXVHGWRGHWHFWGLIIH UHQFHVEHWZHHQPHDQVDWWKHVLJQLILFDQFHOHYHO 5(68/76
&RPSRXQGWR[LFLW\RQPDOHVH[SXSDH
&RPSRXQGWR[LFLW\RQPDOHVH[SXSDHZDVH[SUH VVHGLQWZRPRGHV)LUVWOHWKDOIRUWKHWUHDWHGSXSDH LQFRQVHTXHQFHWKHLUIXUWKHUGHYHORSPHQWZDVGLVDEOHG 7KHSHUFHQWDJHRIGHDGSXSDHDWWKHVWDJHZDVUDQ JHGIURPWRGHSHQGLQJRQWKHFRPSRXQG DQGGRVHV7DEOH$VDQRWKHUPRGHRIDFWLRQLQWHU IHUHQFHWRQRUPDOPHWDPRUSKRVLVRISXSDHLQWRDGXOWV ZDV GHWHFWHG IRUPLQJ ´DGXOWRLGµ XQLWV DQG GHIRUPHG DGXOWVGHYHORSHGIURPWUHDWHGVXUYLYHGSXSDH7KHSHU FHQWDJHRI´DGXOWRLGVµDWWKHVWDJHZDVUDQJHGIURP WRGHDGZKLOHSHUFHQWDJHRIGHIRUPHG XQLWV DW WKH VWDJH ZDV UDQJHG IURP WR OLYHUHVSHFWLYHO\WRGHDGLQUHJDUGWR WKH FRPSRXQG DQG GRVH 7KHVH GHIRUPLWLHV ZHUH OHVV RU PRUH H[SUHVVHG RQ HO\WUD GHYHORSHG DGXOWV :LWK LQFUHDVLQJFLQHROHGRVHIURPWRDQG PO/YROSHUFHQWDJHRIQRUPDOO\GHYHORSHGOLYHDGXOWV
VWDJHZKLFKGHYHORSHGIURPVXUYLYHGWUHDWHGSXSDH KDVEHHQVLJQLILFDQWO\GHFUHDVHGDQG ) GI 3$VLJQLILFDQWHIILFDF\ RIHXJHQROLQUHJDUGWRWKHFRQWUROZDVGHWHFWHGRQO\DW WKHKLJKHVWGRVHH[SUHVVHGZLWKVLJQLILFDQWO\KLJKHUSHU FHQWDJHRIGHDGXQLWVDWWKHVWDJH6LQFHWKLVSHUFHQ WDJHZDVUHODWLYHO\ORZEXWVLJQLILFDQWPRUWDOLW\ RIWUHDWHGSXSDHKDVQRWEHHQUHDFKHGDQGSHUFHQWDJH RIQRUPDOO\GHYHORSHG OLYHDGXOWV KDV QRWEHHQ VLJQLIL FDQWO\GLIIHUHQWIURPWKHFRQWUROLWFRXOGEHFRQFOXGHG WKDWHXJHQROLQWKLVH[SHULPHQWLVQRWHIIHFWLYHDJDLQVW
7DEOH)XPLJDQWHIILFDF\RIFLQHROHDQGHXJHQROWRWKH7ULEROLXPFDVWDQHXP+HUEVWIHPDOHVH[SXSDHVFDOH DFFRUGLQJWR0DQGDYD 7DEOLFD)XPLJDQWQDXĀLQNRYLWRVWFLQHRODLHXJHQRODQDçHQVNHNXNXOMLFHYUVWH7ULEROLXPFDVWDQHXP+HUEVWVNDOD SUHPD0DQGDYL 2LO 8OMH 'RVH PO/ 'R]D PO/YRO
8QLWVRI7ULEROLXPFDVWDQHXPƂE\JURZWKVWDJH
æHQVNHMHGLQNH7ULEROLXPFDVWDQHXPSUHPDVWDGLMXUDVWD *URZWKVWDJH6WDGLMUDVWD 6WDJH 6WDGLM 6WDJH 6WDGLM 6WDJH 6WDGLM 6WDJH 6WDGLM 'HDG 8JLQXOL 0HDQ6( /LYH æLYL 0HDQ6( 'HDG 8JLQXOL 0HDQ6( /LYH æLYL 0HDQ6( 'HDG 8JLQXOL 0HDQ6( /LYH æLYL 0HDQ6( 'HDG 8JLQXOL 0HDQ6( FLQHROH FLQHRO
ED D D D E E D
ED D D D ED E D
D D D D D F D
E D D D E D D
(XJHQRO
(XJHQRO
D D D D D E D
ED D D D E ED D
E D D D D ED D
E D D D E D D
0HDQVLQWKHVDPHFROXPQIROORZHGE\WKHVDPHOHWWHUVLQVXSHUVFULSWDUHQRWVLJQLILFDQWO\GLIIHUHQW3FRPSDULVRQLVIRUHDFKGRVHE\FROXPQV²3URVMHĀQH YULMHGQRVWLXNRORQDPDVLVWLPVORYRPQLVXVWDWLVWLĀNL]QDĀDMQH3XVSRUHGEH]DVYDNXGR]XVXSUHPDNRORQDPD
$OOXQLWVDWWKHVWDJHUHSUHVHQWGHDGSXSDHRQO\²6YHMHGLQNHXID]LSUHGVWDYOMDMXPUWYHNXNXOMLFH
7DEOH)XPLJDQWHIILFDF\RIFLQHROHDQGHXJHQROWRWKH7ULEROLXPFDVWDQHXP+HUEVWPDOHVH[SXSDHVFDOH DFFRUGLQJWR0DQGDYD 7DEOLFD)XPLJDQWQDXĀLQNRYLWRVWFLQHRODLHXJHQRODQDPXåNHNXNXOMLFHYUVWH7ULEROLXPFDVWDQHXP+HUEVWVNDOD SUHPD0DQGDYL 2LO 8OMH 'RVH PO/ 'R]D PO/YRO
8QLWVRI7ULEROLXPFDVWDQHXPƃE\JURZWKVWDJHV
0XåNHMHGLQNH7ULEROLXPFDVWDQHXPSUHPDVWDGLMXUDVWD *URZWKVWDJH6WDGLMUDVWD 6WDJH 6WDGLM 6WDJH 6WDGLM 6WDJH 6WDGLM 6WDJH 6WDGLM 'HDG 8JLQXOL 0HDQ6( /LYH æLYL 0HDQ6( 'HDG 8JLQXOL 0HDQ6( /LYH æLYL 0HDQ6( 'HDG 8JLQXOL 0HDQ6( /LYH æLYL 0HDQ6( 'HDG 8JLQXOL 0HDQ6( FLQHROH FLQHRO
D D E E D E D
D D E D D F D
D D D E D F D
E D E E D D D
(XJHQRO
(XJHQRO
D D D D ED D D
D D ED D ED D D
D D E D D D D
D D E D E D D
0HDQVLQWKHVDPHFROXPQIROORZHGE\WKHVDPHOHWWHUVLQVXSHUVFULSWDUHQRWVLJQLILFDQWO\GLIIHUHQW3FRPSDULVRQLVIRUHDFKGRVHE\FROXPQV²3URVMHĀQH YULMHGQRVWLXNRORQDPDVLVWLPVORYRPQLVXVWDWLVWLĀNL]QDĀDMQH3XVSRUHGEH]DVYDNXGR]XVXSUHPDNRORQDPD
$OOXQLWVDWWKHVWDJHUHSUHVHQWGHDGSXSDHRQO\²6YHMHGLQNHXID]LSUHGVWDYOMDMXPUWYHNXNXOMLFH
&RPSRXQGWR[LFLW\RQIHPDOHVH[SXSDH
7 FDVWDQHXP IHPDOH SXSDH KDYH VKRZQ GLIIH UHQW VHQVLWLYLW\ GHSHQGLQJ RQ WKH DSSOLHG FRPSRXQG DQG GRVH ,Q FRPSDULVRQ RI WKH UHVXOWV 7XNH\·V WHVW
ɲ DPRQJ FRPSRXQGV DQG FRQWURO ZLWKRXW RLO DSSOLFDWLRQ 7DEOH LW LV QRWLFHDEOH WKDW FLQHROH VKRZHG VLJQLILFDQW HIILFLHQF\ IRU IHPDOH SXSDH FRQ
WURO ZLWK VLJQLILFDQWO\ PRUH GHDG XQLWV DW WKH VWDJH DW GRVH RI PO /YRO ) GI
3 DORQJ ZLWK VLJQLILFDQWO\ ORZHU SHUFHQWDJH RIQRUPDOO\GHYHORSHGOLYHDGXOWVXQLWVDWWKHVWDJH ) GI 3ট ,QFUHDVLQJ GRVHV IURP WRPO/YROKDGQRLQIOXHQFHRQVLJQLILFDQWLQFUHDVH
$/LåNDHWDO)80,*$17()),&$&<2)&,1(2/($1'(8*(12/217+(383$/67$*(
&RPSDULVRQRIFRPSRXQGVHIILFDF\EHWZHHQSXSDH VH[
$IWHUWKHIXPLJDWLRQWUHDWPHQW7FDVWDQHXPSXSDH VKRZHG GLIIHUHQW VHQVLWLYLW\ WR WKH WHVWHG FRPSRXQGV GHSHQGLQJ RQ VH[ :LWK FLQHROH VLJQLILFDQW GLIIH UHQFHV LQ VHQVLWLYLW\ EHWZHHQ VH[HV ZHUH REVHUYHG DW KLJKHUGRVHV7DEOH7KXVDWWKHGRVHRIPO/
YROWKHUHZHUHPRUHGHIRUPHGOLYHXQLWVRIPDOHVH[DW WKHVWDJHWKDQIHPDOHVUHVSHFWLYHO\ ) GI 3 $GGLWLRQDOO\ DW WKH VDPH GRVH RI FLQHROH IRU PDOH VH[ WKHUH ZDV D VLJQLIL
FDQWO\ ORZHU SHUFHQWDJH RI OLYH DGXOWV GHYHORSHG IURP WKHWUHDWHGVXUYLYHGSXSDHOLYHPDOHXQLWVDW WKHVWDJHUHVSHFWLYHO\OLYHIHPDOHXQLWVDW WKH VWDJH ) GI 3 DFFRUGLQJO\ SRLQWLQJWRWKHEHWWHUVXUYLYDORIIHPDOHSXSDHLQJHQH UDO0DOHVH[ZDVDWFLQHROHGRVHRIPO/YRO
PRUH VHQVLWLYH ZKLFK LV QRWLFHDEOH WKURXJK D KLJKHU SHUFHQWDJHRI´DGXOWRLGVµDWWKHVWDJHPDOH XQLWV UHVSHFWLYHO\ IHPDOH XQLWV )
GI 3
7DEOH)XPLJDQWHIILFDF\RIFLQHROHEHWZHHQ7ULEROLXPFDVWDQHXP+HUEVWSXSDHVH[VFDOHDFFRUGLQJWR 0DQGDYD
7DEOLFD)XPLJDQWQDXĀLQNRYLWRVWFLQHRODLHXJHQRODL]PHāXVSRORYDNXNXOMLFD7ULEROLXPFDVWDQHXP+HUEVWVNDOD SUHPD0DQGDYL
6H[
6SRO
8QLWVRI7ULEROLXPFDVWDQHXPE\JURZWKVWDJH
-HGLQNH7ULEROLXPFDVWDQHXPSUHPDVWDGLMXUDVWD
*URZWKVWDJH6WDGLMUDVWD
6WDJH
6WDGLM 6WDJH6WDGLM 6WDJH6WDGLM 6WDJH6WDGLM
'HDG
8JLQXOL /LYHæLYL 8JLQXOL'HDG /LYHæLYL 8JLQXOL'HDG /LYHæLYL 8JLQXOL'HDG
'RVHPO/YRO'R]DPO/YRO
0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6(
ƃ D D D D D D D
Ƃ D D D D D D D
'RVHPO/YRO'R]DPO/YRO
0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6(
ƃ D D D D D E D
Ƃ D D D E D D D
'RVHPO/YRO'R]DPO/YRO
0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6(
ƃ D D D D D D D
Ƃ D D E D D D D
0HDQVLQWKHVDPHFROXPQIROORZHGE\WKHVDPHOHWWHUVLQVXSHUVFULSWDUHQRWVLJQLILFDQWO\GLIIHUHQW3FRPSDULVRQLVIRUHDFKGRVHE\FROXPQV²3URVMHĀQH YULMHGQRVWLXNRORQDPDVLVWLPVORYRPQLVXVWDWLVWLĀNL]QDĀDMQH3XVSRUHGEH]DVYDNXGR]XVXSUHPDNRORQDPD
$OOXQLWVDWWKHVWDJHUHSUHVHQWGHDGSXSDHRQO\²6YHMHGLQNHXID]LSUHGVWDYOMDMXPUWYHNXNXOMLFH
'LIIHUHQFHLQHIILFDF\GHSHQGLQJRQSXSDHVH[ZDV OHVVH[SUHVVHGLQWKHWUHDWPHQWZLWKHXJHQRO7DEOH 2QO\DWWKHORZHVWGRVHPO/YROPDOHVH[ZDV
PRUH VHQVLWLYH ZKLFK ZDV H[SUHVVHG DV VLJQLILFDQWO\ KLJKHUSHUFHQWDJHRIGHDG´DGXOWRLGVµZLWKUHJDUGWRWKH IHPDOHVH[UHVSHFWLYHO\) GI 3 $WKLJKHUGRVHVRIHXJHQRODQG PO/YROSXSDHRIERWKVH[HVZHUHHTXDOO\VHQVLWLYH
7DEOH)XPLJDQWHIILFDF\RIHXJHQROEHWZHHQ7ULEROLXPFDVWDQHXP+HUEVWSXSDHVH[VFDOHDFFRUGLQJWR 0DQGDYD
7DEOLFD)XPLJDQWQDXĀLQNRYLWRVWHXJHQRODQDVSRORYHNXNXOMLFD7ULEROLXPFDVWDQHXP+HUEVWVNDODSUHPD0DQGDYL
6H[
6SRO
8QLWVRI7ULEROLXPFDVWDQHXPE\JURZWKVWDJH
-HGLQNH7ULEROLXPFDVWDQHXPSUHPDVWDGLMXUDVWD
*URZWKVWDJH6WDGLMUDVWD
6WDJH
6WDGLM 6WDJH6WDGLM 6WDJH6WDGLM 6WDJH6WDGLM
'HDG
8JLQXOL /LYHæLYL 8JLQXOL'HDG /LYHæLYL 8JLQXOL'HDG /LYHæLYL 8JLQXOL'HDG
'RVHPO/YRO'R]DPO/YRO
0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6(
ƃ D D D D D D D
Ƃ D D E D D D D
'RVHPO/YRO'R]DPO/YRO
0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6(
ƃ D D D D D D D
Ƃ D D D D D D D
'RVHPO/YRO'R]DPO/YRO
0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6( 0HDQ6(
ƃ D D D D D D D
Ƃ D D D D D D D
0HDQVLQWKHVDPHFROXPQIROORZHGE\WKHVDPHOHWWHUVLQVXSHUVFULSWDUHQRWVLJQLILFDQWO\GLIIHUHQW3FRPSDULVRQLVIRUHDFKGRVHE\FROXPQV $OOXQLWVDWWKHVWDJHUHSUHVHQWGHDGSXSDHRQO\
',6&866,21
7KH IXPLJDQW HIIHFWLYHQHVV RI FLQHROH DQG HXJHQRODJDLQVW7FDVWDQHXPSXSDHZDVH[SUHVVHGLQ WZRDFWLRQPRGHVOHWKDODQGDVDQLPSHGLPHQWWRQRU PDOPHWDPRUSKRVLVRISXSDHLQWRDGXOWVWDJHIRUPLQJ ´DGXOWRLGVµ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åND HW DO DW WKH VDPH GRVH PO /YROFLQHROHKDGDVLJQLILFDQWKLJKHUOHWKDOHIIHFW
$/LåNDHWDO)80,*$17()),&$&<2)&,1(2/($1'(8*(12/217+(383$/67$*( &21&/86,21 7KHIDFWWKDWWKHWHVWHGPRQRWHUSHQRLGVIURPRXU UHVHDUFKKDYHDJURZWKUHJXODWRUVIHDWXUHVKRXOGQRWEH QHJOHFWHG+RZHYHULWFRXOGUHSUHVHQWDEDVHIRUIXUWKHU LQYHVWLJDWLRQLQWHUPVRIFRPELQDWLRQRIDIHZHVVHQWLDO RLOV· FRPSRXQGV ZLWK GLIIHUHQW DFWLYLW\ OHWKDO DQG DV D JURZWK UHJXODWRU DOO DLPLQJ WR KDYH PRUH HIIHFWLYH SHVWFRQWURORUPRUHRYHUDSRVVLEOHV\QHUJLVPDPRQJ FRPSRXQGV
$&.12:/('(*0(17
7KLVVWXG\KDVEHHQFRQGXFWHGZLWKLQWKHVFLHQWLILF SURMHFW´%LRDFWLYLW\RIHVVHQWLDORLOFRPSRQHQWVLQVWRUHG JUDLQ SURWHFWLRQµ SURYLGHG E\ WKH 0LQLVWU\RI6FLHQFHDQG6SRUWVRI5HSXEOLFRI&URDWLD 5()(5(1&(6
$UDEL ) 0RKDUUDPLSRXU 6 6HILGNRQ ) &KHPLFDOFRPSRVLWLRQDQGLQVHFWLFLGDODFWLYLW\RIHVVHQ WLDO RLO IURP3HURYVNLD DEURWDQRLGHV/DPLDFHDH DJD LQVW6LWRSKLOXV RU\]DH &ROHRSWHUD &XUFXOLRQLGDHDQG 7ULEROLXPFDVWDQHXP &ROHRSWHUD7HQHEULRQLGDH,QW-7URS,QVHFW6FL
GRLKWWSG[GRLRUJ6 $WKDQDVVLRX &* 3KLOOLSV 7: $LNLQV 0- +DVDQ
00 7KURQH -( (IIHFWLYHQHVV RI VXOSKXU\O IOXRULGHIRUFRQWURORIGLIIHUHQWOLIHVWDJHVRIVWRUHGSUR GXFWSVRFLGV3VRFRSWHUD-(FRQ(QWRPRO
GRLKWWSG[GRLRUJ(&
%HQ]L96WHIDQD]]L1)HUUHUR$$%LRORJLFDO DFWLYLW\RIHVVHQWLDORLOVIURPOHDYHVDQG IUXLWV RI SHSSHU WUHH 6FKLQXV PROOH/ WR FRQWURO ULFH ZHHYLO
6LWRSKLOXVRU\]DH/&KLO-$JU5HV
%RZHUV:6-XYHQLOHKRUPRQHDFWLYLW\RIDUR PDWLFWHUSHQRLGHWKHUV6FLHQFH GRLKWWSG[GRLRUJVFLHQFH 'XFRP 3 0HWK\O EURPLGH DOWHUQDWLYHV ,Q
1DYDUUR 6 %DQNV +- -D\DV '6 %HOO &+ 1R\HV 57 )HUL]OL $* (PHNFL 0 ,VLNEHU $$ $ODJXVXQGDUDP . (GV 3URFHHGLQJV RI WKH WK ,QWHUQDWLRQDO&RQIHUHQFHRQ&RQWUROOHG$WPRVSKHUHDQG )XPLJDWLRQ LQ 6WRUHG 3URGXFW $QWDO\D 7XUNH\ 2FWREHU7XUNH\SS
GRLKWWSG[GRLRUJVSU
(EDGROODKL $ (VVHQWLDO RLOV LVRODWHG IURP 0\UWDFHDH IDPLO\ DV QDWXUDO LQVHFWLFLGHV $QQX 5HY 5HVHDUFKLQ%LRO
)DWKSRXU + 1RRUL $ =HLQDOL % (IIHFWV RI D MXYHQRLG S\ULSUR[\IHQ RQ UHSURGXFWLYH RUJDQ GHYHORSPHQW DQG UHSURGXFWLRQ LQ *HUPDQ FRFNURDFK 'LFW\RSWHUD %ODWWHOOLGDH ,UDQLDQ - 6FL 7HFKQRO 7UDQVDFWLRQ$
*XR 66 <RX &; /LQDJ -< =KDQJ :- *HQJ =) :DQJ &) 'X 66 /HL 1 &KHPLFDO &RPSRVLWLRQ DQG %LRDFWLYLWLHV RI WKH (VVHQWLDO 2LO IURP (WOLQJHUD \XQQDQHQVLV DJDLQVW 7ZR 6WRUHG 3URGXFW ,QVHFWV0ROHFXOHV
GRLKWWSG[GRLRUJPROHFXOHV
,VPDQ 0% 3ODQW HVVHQWLDO RLOV IRU SHVW DQG GLVHDVHPDQDJHPHQW&URS3URWHFW² ,VPDQ0%3URVSHFWLYHERWDQLFDOLQVHFWLFLGHV
IRUULFKHUDQGSRRUHU3HVW0DQDJHPHQW6FL GRLKWWSG[GRLRUJSV
/HH % +R $QQLV 3& 7XPDDOLL ) 7KH SRWHQ WLDO RI FLQHROH DV D IXPLJDQW IRU VWRUHG ZKHDW 3URFHHGLQJV RI WKH $XVWUDOLDQ 3RVWKDUYHVW 7HFKQLFDO &RQIHUHQFH&DQEHUUD²-XQH&6,526WRUHG *UDLQ5HVHDUFK/DERUDWRU\&DQEHUUD
/LåND$5R]PDQ9.DOLQRYLþ,,YH]Lþ0%DOLĀHYLþ 5 &RQWDFW DQG IXPLJDQW DFWLYLW\ RI FLQ HROHHXJHQRODQGFDPSKRUDJDLQVW7ULEROLXPFDVWDQHXP +HUEVW ,Q &DUYDOKR 02 )LHOGV 3* $GOHU &6 $UWKXU )+ $WKDQDVVLRX &* &DPSEHOO -) )OHXUDW /HVVDUG ) )OLQQ 3: +RGJHV 5- ,VLNEHU $$ 1DYDUUR 6 1R\HV 57 5LXGDYHWV - 6LQKD .. 7KRUSH*57LPOLFN%+7UHPDWHUUD3:KLWH1'* (GV 3URFHHGLQJV RI WKH WK ,QWHUQDWLRQDO :RUNLQJ &RQIHUHQFH RQ 6WRUHG 3URGXFW 3URWHFWLRQ (VWRULO 3RUWXJDO-XQHWR-XO\SS
/LåND $ 5R]PDQ9 .DOLQRYLþ , (āHG $ 0XVWDĀ 63HUKRĀ%%LRDFWLYLW\RIFLQHROHDJDLQVW UHG IORXU EHHWOH7ULEROLXP FDVWDQHXP+HUEVW SXSDH 3ROMRSULYUHGD$JULFXOWXUH
/LX =/ +R 6+ %LRDFWLYLW\ RI WKH HVVHQWLDO RLOH[WUDFWHGIURP(YRGLDUXWDHFDUSD+RRNIHW7KRPDV DJDLQVW WKH JUDLQ VWRUDJH LQVHFWV6LWRSKLOXV ]HDPDLV 0RWVFK DQG7ULEROLXP FDVWDQHXP +HUEVW - 6WRUHG 3URG5HV GRLKWWSG[GRLRUJ6; /\RQ:)&RQIXVHGDQG5HGIORXUEHHWOHV2KLR 6WDWH8QLYHUVLW\([WHQWLRQ)DFW6KHHW+<* 0DKURRI50+DJVWUXP':3HVWPDQDJHPHQW LQWKHIRRGLQGXVWU\6WHZDUW3RVWKDUYHVW5HYLHZ 0DQGDYD 1% &5& +DQGERRN RI 1DWXUDO
3HVWLFLGHV0HWKRGHV,Q7KHRU\3UDFWLFHDQG'HWHFWLRQ &5&3UHVV,QF%RFD5DWRQ)/
0DVRQ /- *UDLQ LQVHFW IDFW VKHHW (: UHG DQG FRQIXVHG IORXU EHHWOHV7ULEROLXP FDVWDQHXP %KVWDQG7ULEROLXPFXQIXVXP'XYDO3XUGXH8QLYHUVLW\ 'HSDUWPHQWRI(QWRPRORJ\KWWSH[WHQVLRQHQWPSXUGXH HGXSXEOLFDWLRQV(SGI$FFHVVHG1RYHPEHU 0RQWUHDO 3URWRFRO RQ 6XEVWDQFHV WKDW 'HSOHWH WKH
2]RQHOD\HU81(35HSRUWRIWKH0HWK\O%URPLGH 7HFKQLFDO 2SWLRQV &RPPLWWHH SXUVXDPLW WR $UWLFOH RI WKH0RQWUHDO3URWRFRO'HFLVLRQ,9
6DQWRV-&)DURQL/5$6RXVD$+*XHGHV51& )XPLJDQWWR[LFLW\RIDOO\OLVRWKLRF\DQDWHWRSRSX ODWLRQV RI WKH UHG IORXU EHHWOH7ULEROLXP FDVWDQHXP -6WRUHG3URG5HV
GRLKWWSG[GRLRUJMMVSU 6$6 &RS\ULJKW F E\ 6$6,QVWLWXW ,QF
&DU\1&86$/LFHQVHGWR32/-235,95('1,)$.8/7(7 26,-(.756LWH
6WDPRSRXORV'&'DPRV3.DUDJLDQLGRX* %LRDFWLYLW\ RI ILYH PRQRWHUSHQRLG YDSRXUV WR7ULEROLXP FRQIXVXPGX9DO&ROHRSWHUD7HQHEULRQLGDH-6WRUHG 3URG5HV
)80,*$171$8ÿ,1.29,7267&,1(2/$,(8*(12/$1$67$',-.8.8/-,&(
.(67(1-$672*%5$ä1$5$
75,%2/,80&$67$1(80
+(5%67
,16(&7$&2/(237(5$7(1(%5,21,'$(
6$æ(7$.
)XPLJDQWQDXĀLQNRYLWRVWNRPSRQHQDWDFLQHRODLHXJHQRODWHVWLUDQDMHQDVWDGLMNXNXOMLFHNHVWHQMDVWRJDEUDåQDUD 7ULEROLXPFDVWDQHXP+HUEVW2GUHāHQMHPRUWDOLWHWLDNWLYQRVWUDVWDNXNXOMLFDREDVSRODNDRLUD]OLNHXRVMHWOMLYRVWL L]PHāXVSRORYDNXNXOMLFD7FDVWDQHXPQDLVSLWLYDQHNRPSRQHQWH.RPSRQHQWHVXWHVWLUDQHXNRQFHQWUDFLMH LPOYROXNRQWUROLUDQLPXYMHWLPDQD&UY]XWDPLWLMHNRPVDWL7RNVLĀQRVWNRPSRQHQDWD
ELOD MH L]UDçHQD NUR] GYD QDĀLQD GMHORYDQMD OHWDOQR WH NDR RPHWDQMH QRUPDOQH SUHREUD]EH NXNXOMLFH X RGUDVOL VWDGLM IRUPLUDMXþL ÅDGXOWRLG´ L GHIRUPLUDQH RGUDVOH MHGLQNH NRG RED VSROD .XNXOMLFH PXåNRJD VSROD RSþHQLWR VX VHQ]LELOQLMH QD REMH WHVWLUDQH NRPSRQHQWH 6YHXNXSQR EROMH MH GMHORYDQMH RVWYDUHQR V FLQHRORP GRN V HXJHQRORPQLMHSRVWLJQXW]DGRYROMDYDMXþLXĀLQDNQDWHVWLUDQHNXNXOMLFH
.OMXĀQHULMHĀLIXPLJDFLMDFLQHROHXJHQRO7ULEROLXPFDVWDQHXPNXNXOMLFH
5HFHLYHGRQ0D\DFFHSWHGRQ1RYHPEHU3ULPOMHQRVYLEQMDSULKYDþHQRVWXGHQRJD