Calcium oxalate crystals in the stem of Sida hermaphrodita (L.) Rusby (Malvaceae)

$11$/(6

81,9(56,7$7,60$5,$(&85,(6.à2'2:6.$

/8%/,1±32/21,$

92//;,; 6(&7,2&  _{'HSDUWPHQWRI3ODQW$QDWRP\DQG&\WRORJ\0DULD&XULH6NáRGRZVND8QLYHUVLW\} $NDGHPLFND/XEOLQ3RODQG _{'HSDUWPHQWRI=RRORJ\0DULD&XULH6NáRGRZVND8QLYHUVLW\$NDGHPLFND/XEOLQ} 3RODQGHPDLOHZDV]F]XND#SRF]WDXPFVOXEOLQSO

$*$7$/(6=&=8.



_{(:$6=&=8.$}



_{.,1*$67$1,6à$:(.}



_

,/21$0$=85.,(:&=



_{$11$.$635=<.}



&DOFLXPR[DODWHFU\VWDOVLQWKHVWHPRI6LGDKHUPDSKURGLWD

/5XVE\0DOYDFHDH

.U\V]WDá\V]F]DZLDQXZDSQLDZáRG\G]H6LGDKHUPDSKURGLWD/5XVE\0DOYDFHDH 6800$5< 2EVHUYDWLRQVRIFDOFLXPR[DODWHFU\VWDOVRIWKHVWHPRIDQHQHUJHWLFSODQW6KHUPDSKURGLWD /5XVE\IURPWKH0DOYDFHDHIDPLO\ZHUHSHUIRUPHGXVLQJ/0',&DQG&/60PLFURVFRSHV 7KHWUDQVYHUVDODQGORQJLWXGLQDOVHFWLRQVVKRZHGWKHSUHVHQFHRIFDOFLXPR[DODWHFU\VWDOVLQWKH SDUHQFK\PDOWLVVXHGLVWULEXWHGLQYDULRXVOD\HUVRIWKHVWHP7KHFU\VWDOVRFFXUUHGRQO\LQWKHIRUP RIGUXVHV,QWKHLQQHUPRVWSDUWRIWKHVWHPLHLQWKHSLWKWKHFDOFLXPR[DODWHFU\VWDOVRFFXUUHG VLQJO\LQLQGLYLGXDOFHOOV,QWKHSDUHQFK\PDFHOOVVHSDUDWLQJVFOHUHQFK\PD¿EUHVDQGDGMDFHQWWRWKH [\OHPWKHFU\VWDOVZHUHREVHUYHGLQGLYLGXDOO\LQVLQJOHFHOOVEXWWKHFHOOVFRQWDLQLQJGUXVHVIRUPHG URZVFRQVLVWLQJRIHYHQVHYHUDOFHOOV7KHFRUWH[FRQWDLQHGWKHGLIIHUHQWVL]HGUXVHVVFDWWHUHGUDQ GRPO\ZLWKLQWKHFHOOV'UXVHVGLIIHULQVKDSHDQGVL]HEXWWKH\GRQRWSURWUXGHEH\RQGWKHFHOOVDO WKRXJKWKH\YHU\RIWHQ¿OOWKHPFRPSOHWHO\7KHIXQFWLRQVRIFDOFLXPR[DODWHFU\VWDOVDUHGLVFXVVHG .H\ZRUGVFDOFLXPR[DODWHFU\VWDOV0DOYDFHDH6LGDKHUPDSKURGLWDVWHP 675(6=&=(1,( 3U]\XĪ\FLXPLNURVNRSXĞZLHWOQHJR]NRQWUDVWHP1RPDUVNLHJRLNRQIRNDOQHJRSU]HSURZD G]RQRREVHUZDFMHNU\V]WDáyZV]F]DZLDQXZDSQLDZáRG\G]HHQHUJHW\F]QHMURĞOLQ\6LGDKHUPDSK URGLWD/5XVE\]URG]LQ\0DOYDFHDH3U]HNURMHSRSU]HF]QHRUD]SRGáXĪQHZ\ND]Dá\REHFQRĞü NU\V]WDáyZV]F]DZLDQXZDSQLDZWNDQFHSDUHQFK\PDW\F]QHM]QDMGXMąFHMVLĊZUyĪQ\FKZDUVWZDFK áRG\JL.U\V]WDá\Z\VWĊSRZDá\Z\áąF]QLHZSRVWDFLGUX]yZ:QDMJáĊEV]HMF]ĊĞFLáRG\JLWMZUG]H

UMCS



QLXNU\V]WDá\V]F]DZLDQXZDSQLDZ\VWĊSRZDá\SRMHG\QF]RZSRV]F]HJyOQ\FKNRPyUNDFK:NR PyUNDFK SDUHQFK\PDW\F]Q\FK RGG]LHODMąF\FK ZáyNQD VNOHUHQFK\PDW\F]QH L SU]\OHJDMąF\FK GR NV\OHPXREVHUZRZDQRSRMHG\QF]HNU\V]WDá\ZSRV]F]HJyOQ\FKNRPyUNDFKDNRPyUNL]DZLHUDMą FHGUX]\IRUPRZDá\U]ĊG\VNáDGDMąFHVLĊ]NLONXNRPyUHN.RUD]DZLHUDáDUyĪQHMZLHONRĞFLGUX]\ UR]PLHV]F]RQHSU]\SDGNRZRZNRPyUNDFK'UX]\UyĪQLąFHVLĊNV]WDáWHPLZLHONRĞFLąF]ĊVWRZ\ SHáQLDMąFDáNRZLFLHNRPyUNĊDOHQLHZ\VWDMąSR]DQLą:SUDF\G\VNXWRZDQHVąIXQNFMHNU\V]WDáyZ V]F]DZLDQXZDSQLD 6áRZDNOXF]RZH6LGDKHUPDSKURGLWD0DOYDFHDHNU\V]WDá\V]F]DZLDQXZDSQLDáRG\JD ,1752'8&7,21

7KH JHQXV 6LGD LQFOXGHV DERXW  VSHFLHV RI DQQXDO SODQWV SHUHQQLDOV DQG VKUXEV 2QH RI WKH VSHFLHV EHORQJLQJ WR WKLV JHQXV WKH 9LUJLQLD PDOORZ 6LGD KHUPDSKURGLWD / 5XVE\ ZDVLQWURGXFHGLQ3RODQGLQWKH¿IWLHVRIWKHODVWFHQWXU\7KH¿UVWLQLWLDWHGVWXGLHVZHUHIRFXVHGRQ 6KHUPDSKURGLWDDVD¿EUHSODQW6XEVHTXHQWVWXGLHVH[SDQGHGRXUXQGHUVWDQGLQJRIWKHYDOXHRI WKHXVHRIWKLVVSHFLHV7KH\GHPRQVWUDWHGWKHSRVVLELOLW\RIXVLQJ6LGDDVDIRGGHUSODQWDVZHOODV IRUKRQH\SURGXFWLRQELRUHPHGLDWLRQUHVWRUDWLRQDQGDVDSKDUPDFHXWLFDOUDZPDWHULDO,WDOVRKDV DQXPEHURIIHDWXUHVHQVXULQJSRWHQWLDOXVHDVSXOSLQSDSHULQGXVWU\$SDUWIURPDOOWKHVHSURSRV DOVRIWKHXVHRIWKH9LUJLQLDPDOORZLWFDUULHVDJUHDWSURPLVHWREHDYDOXDEOHHQHUJ\SODQW &DOFLXPVDOWVDUHDFRPPRQO\RFFXUULQJFRPSRQHQWRISODQWFHOOV7KLVHOHPHQWLVSUHVHQWLQ WKHLRQLFRUERXQGIRUP,QPRVWSODQWVFDOFLXPLVGHSRVLWHGLQWKHIRUPRIFDOFLXPVDOWVXVXDOO\DV LQVROXEOHFDOFLXPR[DODWHFU\VWDOV,WRFFXUVDVDPRQRRUGLK\GUDWHVDOWDQGUDUHO\DVDWULK\GUDWH 7KHPRQRK\GUDWHIRUPLVPRUHVWDEOHDQGWKHPRVWFRPPRQLQSODQWV&DOFLXPR[DODWHKDVEHHQ LGHQWL¿HGLQPRUHWKDQSODQWIDPLOLHVLQFOXGLQJ0DOYDFHDH7KHFU\VWDOVDUHIRUPHGLQ DOPRVWDOOSDUWVRIWKHSODQWÀRZHUVOHDYHVVWHPDQGURRWV7KHDPRXQW RIDFFXPXODWHGR[DODWHVLVGHSHQGHQWRQWKHDQDWRPLFDOSDUWRIWKHSODQW7KHODUJHVWDPRXQWVFDQEH IRXQGLQORZHUOHDYHVORZHUTXDQWLWLHVLQXSSHUOHDYHVVWHPVDQGVHHGVDQGWKHVPDOOHVWDPRXQWV DUHSUHVHQWLQWKHURRWV([FHVVFDOFLXPDFFXPXODWHVSUHGRPLQDQWO\LQYDFXROHVDQGFLVWHUQVRIWKH HQGRSODVPLFUHWLFXOXPEXWDOVRLQWKHFHOOZDOO&DOFLXPR[DODWHFU\VWDOVPD\EHFRQVLGHUDEO\VPDOO HUWKDQWKHFHOOVFRQWDLQLQJWKHP¿OOWKHFHOOVFRPSOHWHO\RUFDXVHSHUIRUDWLRQVRIWKHLUZDOOV

&HOOV LQ ZKLFK FU\VWDOV DUH IRUPHG PD\ EH DUUDQJHG UDQGRPO\ ZLWKLQ WKH WLVVXH 6SHFLDO L]HG VHFUHWRU\ FHOOV VRFDOOHG LGLREODVWV RIWHQ KDYH D QXPEHU RI GLVWLQJXLVKLQJ FKDUDFWHULVWLFV $PRQJRWKHUVWKH\FDQFRQWDLQDQLQFUHDVHGFHOOQXFOHXVPRGL¿HGSODVWLGVDKLJKO\GHYHORSHG(5 DQG*ROJLDSSDUDWXVLQFUHDVHGOHYHOVRIU51$DQGDXQLTXHFRPSRVLWLRQRIWKHYDFXROHFRQWHQW 7KHF\WRSODVPRIWKHVHFHOOVLVULFKLQYDULRXVRUJDQHOOHVPHPEUDQHVDQGYHVLFOHV7KHFHOOZDOOLV DOVRPRGL¿HGEHFDXVHLWLVLUUHJXODUDQGPXFKWKLQQHULQVRPHDUHDV0XFXVDFRPSRQHQWRIWKH F\WRSODVPVZHOOVKHQFHSHUIRUDWLRQDQGHMHFWLRQRIWKHFU\VWDOVRXWVLGHWKHFHOORFFXUV 0$7(5,$/6$1'0(7+2'6  7KHPDWHULDOFRPSULVHGVWHPVRI6KHUPDSKURGLWD/5XVE\7KHFPWKLFNVWHPIUDJPHQWV ZHUHWDNHQDWDKHLJKWRIPDQG¿[HGLQDPL[WXUHRIHWKDQRODQGDFHWLFDFLG&DUQR\¶V¿[D WLYH7KHXVHGPL[WXUHZDVFRPSRVHGRIHWKDQRODQGDFHWLFDFLGPL[HGDWDUDWLRRI7KHQ WKH PDWHULDO ZDV GHK\GUDWHG LQ LQFUHDVLQJ FRQFHQWUDWLRQV RI HWKDQRO VROXWLRQV7KH PDWHULDO ZDV VRDNHGLQDPL[WXUHRIDQK\GURXVHWKDQRODQGEHQ]HQH7KH¿QDOVWDJHRIWKHSURFHGXUHFRQVLVWHG RIHPEHGGLQJWKHPDWHULDOLQSDUDI¿QIRUDIHZGD\VXSWRRQHZHHN7KHREWDLQHGSDUDI¿QEORFNV ZHUHFXWRQDURWDU\PLFURWRPHLQWRPLFURQWKLFNVHFWLRQV$IWHUUHPRYLQJWKHSDUDI¿QWKHVHF WLRQVPRXQWHGRQWKHJODVVVOLGHVDQGVWDLQHGZLWKVDIUDQLQDQGIDVWJUHHQ $/(6=&=8.(6=&=8.$.67$1,6à$:(./0$=85.,(:&=$.$635=<.

UMCS



7KHPDWHULDO¿[HGLQ&DUQR\¶VPL[WXUHZLWKRXWHPEHGGLQJLQSDUDI¿QZDVFXWRQWKHVOHGJH PLFURWRPH7KHVHFWLRQVREWDLQHGZHUHVWDLQHGZLWKVDIUDQLQDQGIDVWJUHHQ7KHSUHSDUDWLRQVZHUH REVHUYHG XQGHU OLJKW PLFURVFRSH /0 DQG GLIIHUHQWLDO LQWHUIHUHQFH FRQWUDVW PLFURVFRSH ',& )RUWKHREVHUYDWLRQVXQGHUFRQIRFDOODVHUVFDQQLQJPLFURVFRSH&/60WKHVHFWLRQVZHUHVWDLQHG ZLWKHRVLQ<ZDWHUVROXWLRQ 7KHLPDJHVZHUHWDNHQXVLQJ1LNRQ&RROSL[DQG2O\PSXV'3FDPHUDV 5(68/76$1'',6&866,21



&DOFLXPR[DODWHFU\VWDOVDUHFKDUDFWHUL]HGE\DJUHDWGLYHUVLW\RIPRUSKRO

RJ\7KH\WDNHVXFKGLIIHUHQWVKDSHVWKDWWKH\FDQFRQVWLWXWHDXVHIXOGLDJQRVWLF

IHDWXUHLQWKHWD[RQRP\LQPDQ\XQUHODWHGJURXSVRISODQWV&DOFLXPR[DODWHPRVW

FRPPRQO\RFFXUVLQWKHIRUPRIFU\VWDOVIRUPLQJGLIIHUHQWVL]HDQGVSDWLDOVKDSH

JURXSV 7KH EDVLF PRUSKRORJLFDO VWUXFWXUHV FDQ EH FRQVLGHUHG DV FU\VWDO VDQG

UDSKLGHVVW\ORLGVSULVPVDQGFU\VWDOOLQHGUXVHV6W\ORLGVDUHODUJHVL]HGSLOODU

OLNHFU\VWDOV7KH\XVXDOO\RFFXUVLQJO\DQGKDYHEOXQWRUVKDUSHQGV7KH\DUH

YHU\FRPPRQLQWKHFHOOVDQGWLVVXHVRIWKHUHSUHVHQWDWLYHVRIWKHIDPLO\,ULGD

FHDH5DSKLGHVZKLFKDUHFKDUDFWHULVWLFRIPRQRFRWVWDNHWKHIRUPRIFU\VWDO

OLQHQHHGOHVZKLFKDUHXVXDOO\JDWKHUHGLQVLQJOHEXQGOHV7\SLFDOEXQGOHVUHDFK

DFRQVLGHUDEOHVL]HDQGWKH\PD\EHJURRYHG3ULVPDWLFFU\VWDOVDUHVKRUWLUUHJX

ODUSULVPV'UXVHVDUHVSKHULFDOVWHOODUFOXVWHUVDQGSRO\PRUSKLFDJJUHJDWHVRI

FU\VWDOV7KLVIRUPRIFDOFLXPR[DODWHLVRIWHQIRXQGLQGLFRWV'UXVHVDUHYHU\

FRPPRQLQ\RXQJVKRRWVLQWKHUD\VRIWKHFRUHRIPDQ\WUHHV&U\VWDOOLQHVDQGLV

WLQ\FU\VWDOVORRVHO\VFDWWHUHGLQFHOOV7KLVIRUPRIFDOFLXPR[DODWHFDQEHIRXQG

LQSODQWVRIWKH6RODQDFDHDHIDPLO\VXFKDVWKHSRWDWR6RODQXPWXEHURVXPRU

'HDGO\1LJKWVKDGH$WURSDEHOODGRQQD

&DOFLXP R[DODWH FU\VWDOV ZHUH REVHUYHG LQ WKH VWHP RI WKH HQHUJHWLF SODQW

6LGDKHUPDSKURGLWD)LJVDQG7KHVWHPRI6KHUPDSKURGLWDZDVVHFWLRQHG

WUDQVYHUVHO\DWDKHLJKWRIPHWUHV7KHIROORZLQJOD\HUVZHUHFOHDUO\UHFRJQL]

DEOHLQWKHVWHPWKHHSLGHUPLVSULPDU\FRUWH[[\OHPDQGSLWK)LJ,QWKH

LQQHUPRVWSRUWLRQRIWKHVWHPWKHSLWKLVFRPSRVHGRIGLIIHUHQWVL]HSDUHQFK\PDO

FHOOVZKLFKYHU\RIWHQFRQWDLQFDOFLXPR[DODWHFU\VWDOV)LJVDQG,QWKLV

SDUWRIWKH6KHUPDSKURGLWDVWHPWKHFDOFLXPR[DODWHFU\VWDOVRFFXUUHGVLQJO\

LQLQGLYLGXDOFHOOV,QWXUQLQWKHSDUHQFK\PDFHOOVVHSDUDWLQJVFOHUHQFK\PD¿

EUHVWKHFU\VWDOVZHUHDOVRREVHUYHGLQGLYLGXDOO\LQVLQJOHFHOOVEXWLQWKLVFDVH

WKHFHOOVFRQWDLQLQJGUXVHVIRUPHGURZVFRQVLVWLQJRIHYHQVHYHUDOFHOOV)LJ

,QWKHGHVFULEHGWLVVXHLWLVSRVVLEOHWRQRWLFHH[WUHPHO\UHJXODUDUUDQJHPHQWRI

FHOOV FRQWDLQLQJ WKH FDOFLXP R[DODWH FU\VWDOV 2Q WKH RWKHU KDQG LQ WKH FRUWH[

GLIIHUHQWVL]H GUXVHV ZHUH VFDWWHUHG UDQGRPO\ ZLWKLQ WKLV SDUHQFK\PDO WLVVXH

)LJ 7KH ORQJLWXGLQDO VHFWLRQ RI WKH VWHP RI 6 KHUPDSKURGLWD VKRZHG WKH

SUHVHQFHRIGUXVHVLQWKHVHULHVRISDUHQFK\PDFHOOVDGMDFHQWWRWKHWUDFKHDU\HOH

&$/&,802;$/$7(&5<67$/6,17+(67(02)6,'$+(50$3+52',7$/586%<0$/9$&($(



7DE, )LJ6LGDKHUPDSKURGLWD/5XVE\3ODQWVJURZLQJLQDQH[SHULPHQWDO¿HOGRIWKH8QLYHUVLW\ RI/LIH6FLHQFHVLQ/XEOLQ )LJ)UDJPHQWRIWKHJUHHQVWHPRI6KHUPDSKURGLWDZLWKDWKLFNQHVVRIFP )LJV±&URVVVHFWLRQRIWKHVWHPRI6KHUPDSKURGLWD7KH¿JXUHVKRZVWKUHHW\SLFDOOD\HUV HSLGHUPLV(SZLWKWKHFXWLFOH&XFRUWH[&VFOHUHQFK\PD¿EUHV6FDQGYDVFXODUWLVVXHZLWK ÀXRUHVFHQW [\OHP ; &/60 )LJV  DQG  VKRZ GLIIXVH SDUHQFK\PD 3 DQG FOXVWHU FU\VWDOV RIFDOFLXPR[DODWH')LJ/0)LJ',&6FDOHEDU —P

$/(6=&=8.(6=&=8.$.67$1,6à$:(./0$=85.,(:&=$.$635=<.



7DE,,

7KHVWHPRI6KHUPDSKURGLWD

)LJ  7DQJHQWLDO VHFWLRQ )UDJPHQWV RI WKH VWHP VKRZLQJ VFOHUHQFK\PD ¿EUHV 6F DQG SDUHQFK\PDFHOOV3ZLWKGUXVHV'6WDLQLQJZLWKVDIUDQLQDQGIDVWJUHHQ',&6FDOHEDU —P )LJ&URVVVHFWLRQ7KHDQDWRPLFDOVWUXFWXUHRIDIUDJPHQWRIWKHFRUWH[7KH¿JXUHVKRZV YDULRXVVL]HVRISDUHQFK\PDFHOOV3FRQWDLQLQJPXOWLIDULRXVJURXSVRIFDOFLXPR[DODWHFHOOV' 6WDLQLQJZLWKVDIUDQLQDQGIDVWJUHHQ',&6FDOHEDU —P

)LJ  7KH VWHP RI 6 KHUPDSKURGLWD ± ORQJLWXGLQDO VHFWLRQ 7KH IUDJPHQW VKRZV [\OHP HOHPHQWVZLWKYLVLEOHKHOLFDOWKLFNHQLQJV;DQGSDUHQFK\PDFHOOV3ZLWKVHULHVRIGUXVHV' 7KHHORQJDWHGSDUHQFK\PDFHOOVRQWKHOHIWIURPWKHURZFRQWDLQLQJGUXVHVVWDUWHGWRGLIIHUHQWLDWH LQWRDQRWKHUNLQGRIWLVVXH6WDLQLQJZLWKVDIUDQLQDQGIDVWJUHHQ/06FDOHEDU —P

)LJV  DQG  /RQJLWXGLQDO VHFWLRQ 'UXVHV LQ WKH FHOOV DUUDQJHG LQ URZV DQG LQ WKH QHLJKERXULQJFHOOVDIWHUPLWRWLFFHOOGLYLVLRQ6WDLQLQJZLWKVDIUDQLQDQGIDVWJUHHQ/0)LJDQG ',&)LJ6FDOHEDU —P

)LJ  7KH VWHP RI 6 KHUPDSKURGLWD ± FURVV VHFWLRQ )UDJPHQW VKRZV SDUHQFK\PD FHOOV 3ZLWKDURZRIFHOOVFRQWDLQLQJGUXVHV'6WDLQLQJZLWKVDIUDQLQDQGIDVWJUHHQ/06FDOHEDU —P

&$/&,802;$/$7(&5<67$/6,17+(67(02)6,'$+(50$3+52',7$/586%<0$/9$&($(



PHQWVRIWKH[\OHP)LJ,QWKLVSDUHQFK\PDOSDUWRIWKHVWHPFDOFLXPR[DODWH

FU\VWDOVZHUHVSRUDGLFDOO\SUHVHQWLQWKHFHOOVDUUDQJHGLQWZRURZVRIFHOOV)LJV

DQG7KHFURVVVHFWLRQRIWKHGHVFULEHGSDUHQFK\PDOWLVVXHUHYHDOHGORQJ

URZVRIFHOOVFRQWDLQLQJGUXVHVVPDOOHUWKDQQHLJKERXULQJFHOOV)LJ

2EVHUYDWLRQV RI WKH FURVVVHFWLRQ RI WKH 6 KHUPDSKURGLWD VWHPV LQ

GLFDWHG WKH SUHVHQFH RI FDOFLXP R[DODWH FU\VWDOV LQ WKH SDUHQFK\PDO WLV

VXH 7KH FU\VWDOV WDNH WKH IRUP RI GUXVHV ZKLFK DUH LUUHJXODU FOXPSV DQG DJ

JUHJDWHV RI VPDOOHU FU\VWDOV 'UXVHV DUH SUHVHQW LQ YDULRXV OD\HUV RI WKH

6 KHUPDSKURGLWD VWHP 7KH\ DUH SUHVHQW LQ D FUXPE VWUXFWXUH DVVRFLDWHG ZLWK

WKH YDVFXODU EXQGOHV DV ZHOO DV LQ FHOOV RI WKH SDUHQFK\PDO FRUWH[ 0RVW RI

WKH GUXVHV DUH SUHVHQW LQ JURXSV RI FHOOV DUUDQJHG OLQHDUO\ DORQJ WKH [\OHP ¿

EUHV'UXVHVGLIIHULQVKDSHDQGVL]HEXWWKH\GRQRWSURWUXGHEH\RQGWKHFHOOV

DOWKRXJK WKH\ YHU\ RIWHQ ¿OO WKHP FRPSOHWHO\ +RZHYHU WKH FU\VWDOV LQ WKH

6KHUPDSKURGLWDVWHPDUHXVXDOO\ORFDWHGLQFHOOVZLWKVOLJKWO\VPDOOHUGLPHQ

VLRQV$VPHQWLRQHGDERYHGUXVHVFDQ¿OOWKHHQWLUHFHOOEXWWKHLUVL]HVDUHFOHDUO\

GLIIHUHQWLDWHG,QWKH6KHUPDSKURGLWDVWHPVPDOOFU\VWDOVDERXWPLFURQVDQG

WKRVHZLWKDFRQVLGHUDEOHVL]HRI±PLFURQVZHUHREVHUYHG,QRUGHUWRGH

VFULEHVSHFLDOIHDWXUHVDQGWRVSHFLI\WKHSUHFLVHW\SHDQGORFDOL]DWLRQRIFDOFLXP

R[DODWHFU\VWDOVIXUWKHUVWXGLHVDUHUHTXLUHG,QWKHFHOOVRIWKH6KHUPDSKURGLWD

VWHPFU\VWDOVRIFDOFLXPR[DODWHZHUHRQO\REVHUYHGLQWKHIRUPRIGUXVHVDQG

QHYHULQWKHRWKHUIRUPVRIFDOFLXPR[DODWH

7KH VL]H ORFDWLRQ DQG RWKHU SURSHUWLHV RI WKH FDOFLXP R[DODWH FU\VWDOV LQ

SODQWVDUHWKHUHVXOWRISK\VLFDOFKHPLFDODQGELRORJLFDOIDFWRUVVXFKDVWHPSHUD

WXUHOLJKWLQWHQVLW\SUHVVXUHS+DQGLRQFRQFHQWUDWLRQ&U\VWDOVIRUPIURP

HQGRJHQRXVO\V\QWKHVL]HGR[DOLFDFLGDQGFDOFLXPFKDUJHGIURPWKHHQYLURQPHQW

6LQFHWKHPDLQVRXUFHRIR[DOLFDFLGLVDVFRUELFDFLGPDQ\VWXGLHVKDYHVKRZQ

WKDWWKHIRUPDWLRQRI&D2[FDOFLXPR[DODWHFU\VWDOVLVUHODWHGWRWKHPHFKDQLVP

IRUUHJXODWLQJWKHOHYHORIIUHHFDOFLXPLQFHOOV,WKDVEHHQVKRZQWKDWWKHUHLV

DUHODWLRQVKLSEHWZHHQWKHOHYHORIIUHHFDOFLXPDQGWKHDPRXQWDQGVL]HRIFU\V

WDOV$VWKHDPRXQWRIPHWDOLRQVLQWKHFHOOLQFUHDVHVWKHQXPEHURIJHQHUDWHG

FU\VWDOVDOVRLQFUHDVHV+RZHYHUWKHSUHVHQFHRIFDOFLXPR[DODWHLWVPRUSKRORJL

FDOIRUPDQGWKHGLVWULEXWLRQLVFRQWUROOHGJHQHWLFDOO\7KHIRUPDWLRQRIFU\VWDOV

LVQRWDVLPSOHSURFHVVRIVDOWSUHFLSLWDWLRQ7KHIRUPDWLRQRIWKHVHVWUXFWXUHVLV

DFRPELQDWLRQRIJHQHWLFDQGHQYLURQPHQWDOIDFWRUV

7KHSUHYDOHQFHDQGGLYHUVLW\RIFDOFLXPR[DODWHRFFXUULQJLQSODQWVWHQGWR

GH¿QHLWVELRORJLFDOUROH7KHVHVDOWVPD\SOD\DQLPSRUWDQWUROHLQPDQ\YLWDO

IXQFWLRQVRIWKHFHOO7KH\DUHDUHVHUYRLURIH[FHVVFDOFLXPLQWKHFHOODQGDUHLQ

YROYHGLQWKHUHJXODWLRQRILWVOHYHO&D2[FU\VWDOVSOD\DGHIHQVLYHIXQFWLRQLQGH

WHUULQJDQLPDOVWKDWIHHGRQSODQWV7KHFKDUDFWHULVWLFVKDUSIUDJPHQWVRIFU\VWDOV

DUHDQHIIHFWLYHSDVVLYHGHIHQFHDJDLQVWSUH\LQJLQVHFWV5HVHDUFKFRQGXFWHG

$/(6=&=8.(6=&=8.$.67$1,6à$:(./0$=85.,(:&=$.$635=<.

UMCS



RQ WKH VWHP RI 3LVWLD VWUDWLRWHV SURYLGHV LQIRUPDWLRQ RQ DFWLYH GHIHQFH DJDLQVW

KHUELYRUHVDQGDOOHORSDWKLFDFWLYLW\&DOFLXPR[DODWHFU\VWDOVDOVRDFWDVKHOS

HUVLQGHWR[LI\LQJKHDY\PHWDOV([SHULPHQWDODQDO\VLVRIWKHFKHPLFDOFRPSRVL

WLRQRIWKHVHVWUXFWXUHVSUHVHQWLQWKHFHOOVRI0DOYDSDUYLÀRUDSURYHGWKDWWKH

FU\VWDOVFRQWDLQGRSDQWLRQVRIDOXPLQLXPFDGPLXPDQGOHDG7KHLQYHVWLJDWLRQV

DOVRVKRZHGWKDWWKHSUHVHQFHRIKHDY\PHWDOVDIIHFWVWKHELRORJLFDOPHFKDQLVP

RIWKHIRUPDWLRQRIWKHFU\VWDOV%DVHGRQH[SHULPHQWVLQYHVWLJDWLQJWKHSUHVHQFH

RIFU\VWDOVLQWKHOHDIWLVVXHVRI3KDVHROXVYXOJDULVLWZDVK\SRWKHVL]HGWKDWWKH

HPHUJHQFHRIKHDY\PHWDOVUHGXFHVWKHIRUPDWLRQRIWKHVHVWUXFWXUHVLQFHOOV

&DOFLXPR[DODWHFU\VWDOVPD\SRVVLEO\KDYHDQLPSDFWRQSKRWRV\QWKHVLV7KLVLV

GXHWRWKHVSDWLDOVWUXFWXUHDQGWKHSRVVLELOLW\RIUHÀHFWLRQRIOLJKWLQGLIIHUHQWGL

UHFWLRQV7KLVFDQUHVXOWLQORZOLJKWOHYHOVUHDFKLQJWKHGHHSHVWXQGHUO\LQJFHOOV

5HVHDUFK FRQFHUQLQJ GUXVHV ORFDWHG LQ WKH SDOLVDGH SDUHQFK\PD RI 3HSHURPLD

JODEHOODOHDIFRQ¿UPHGWKLVK\SRWKHVLV,WXQGRXEWHGO\UHVXOWVIURPWKHVWXGLHV

VKRZLQJWKDWWKHUHLVDUHODWLRQVKLSEHWZHHQWKHSRVLWLRQDQGWKHQXPEHURIFU\V

WDOVDQGWKHLQWHQVLW\RIOLJKW7KHGXUDEOHFU\VWDOVWUXFWXUHRI&D2[UHLQIRUFHV

FHOOZDOOV7KLVPD\EHHYLGHQFHGE\WKHW\SLFDOORFDWLRQRIWKHVHVWUXFWXUHVLQWKH

FHOOVRIWKHHSLGHUPLVRIGLIIHUHQWSODQWVVWUHQJWKHQLQJWKHFHOOVIRUPLQJWKHYDV

FXODUOD\HUFRYHULQJWKHVHHG7KHFDOFLXPR[DODWHFU\VWDOVLQWKHVWHPRI6LGD

KHUPDSKURGLWDKDYHVLPLODUIXQFWLRQVOLNHLQWKHRWKHUJURXSVRISODQWV9LUJLQLD

PDOORZKDVDVLJQL¿FDQWKHLJKWVRWKHSUHVHQFHRIFU\VWDOVKDVDQLPSRUWDQWUROH

IRUVWUHQJWKHQLQJRIWKHVWHP

5()(5(1&(6  %RUNRZVND+6W\N%ĝOD]RZLHFSHQV\OZDĔVNL6LGDKHUPDSKURGLWD5XVE\8SUDZD LZ\NRU]\VWDQLH:$5/XEOLQ  %UXEDNHU&/+RUQHU+7'HYHORSPHQWRIHSLGHUPDOFU\VWDOVLQOHDÀHWVRI6W\ORVDQWKHV JXLDQHQVLV/HJXPLQRVDH3DSLOLRQRLGHDH&DQDGLDO-RXUQDORI%RWDQ\±  )DKHHG)0D]HQ$$EG(OPRKVHQ63K\VLRORJLFDODQGXOWUDVWUXFWXUDOVWXGLHVRQFDO FLXPR[DODWHFU\VWDOIRUPDWLRQLQVRPHSODQWV7XUNLVK-RXUQDORI%RWDQ\±  )UDQFHVFKL9 5 +RUQHU +7 -U  &DOFLXP 2[DODWH &U\VWDOV LQ 3ODQWV7KH %RWDQLFDO

5HYLHZ±

 )UDQFHVFKL951DNDWD3$&DOFLXP2[DODWHLQ3ODQWV)RUPDWLRQDQG)XQFWLRQ$Q QXDO5HYLHZRI3ODQW%LRORJ\±

 .DWD\DPD + )XMLED\DVKL< 1DJDRND 6 6XJLPXUD<  &HOO ZDOO VKHDWK VXUURXQGLQJ FDOFLXPR[DODWHFU\VWDOVLQPXOEHUU\LGLREODVWV3URWRSODVPD±  .DOHPEDVD6:LĞQLHZVND%:Sá\ZGDZHND]RWXQDSORQELRPDV\ĞOD]RZFDSHQV\O ZDĔVNLHJR6LGDKHUPDSKURGLWD5XVE\RUD]]DZDUWRĞüZQLHMPDNURHOHPHQWyZ$FWD$JUR SK\VLFD±  .RVWPDQ7$)UDQFHVFKL95&HOODQGFDOFLXPR[DODWHFU\VWDOJURZWKLVFRRUGLQDWHGWR DFKLHYHKLJKFDSDFLW\FDOFLXPUHJXODWLRQLQSODQWV3URWRSODVPD± &$/&,802;$/$7(&5<67$/6,17+(67(02)6,'$+(50$3+52',7$/586%<0$/9$&($(

UMCS



 .XR+XDQJ // .X 06% )UDQFHVFKL9 5  &RUUHODWLRQV EHWZHHQ FDOFLXP R[DODWH FU\VWDOVDQGSKRWRV\QWKHWLFDFWLYLWLHVLQSDOLVDGHFHOOVRIVKDGHDGDSWHG3HSHURPLDJODEHOOD %RWDQLFDO6WXGLHV  0HULF&&DOFLXP2[DODWH&U\VWDOVLQ6RPH6SHFLHVRIWKH7ULEH,QXOHDH$VWHUDFHDH $FWD%LRORJLFD&UDFRYLHQVLD±  0RODQR)ORUHV%+HUELYRU\DQG&DOFLXP&RQFHQWUDWLRQV$IIHFW&DOFLXP2[DODWH&U\V WDO)RUPDWLRQLQ/HDYHVRI6LGD0DOYDFHDH$QQDOVRI%RWDQ\±  1DNDWD3$$GYDQFHVLQRXUXQGHUVWDQGLQJRIFDOFLXPR[DODWHFU\VWDOIRUPDWLRQDQG IXQFWLRQLQSODQWV3ODQW6FLHQFH±  3DWWDU39-D\DUDM03KDUPDFRJQRVWLFDQGSK\WRFKHPLFDOLQYHVWLJDWLRQRI6LGDFRUGL IROLD/$WKUHDWHQHGPHGLFLQDOKHUE,QWHUQDWLRQDO-RXUQDORI3KDUPDF\DQG3KDUPDFHXWLFDO 6FLHQFHV±  9RON*0/\QFK+ROP9-.RVWPDQ7$*RVV/-)UDQFHVFKL957KH5ROHRI 'UXVHDQG5DSKLGH&DOFLXP2[DODWH&U\VWDOVLQ7LVVXH&DOFLXP5HJXODWLRQLQ3LVWLDVWUDWLRWHV /HDYHV3ODQW%LRORJ\±  :HEE0$&HOO0HGLDWHG&U\VWDOL]DWLRQRI&DOFLXP2[DODWHLQ3ODQWV7KH3ODQW&HOO ±

 =LQGOHU)UDQN ( +|QRZ 5 +HVVH$  &DOFLXP DQG R[DODWH FRQWHQW RI WKH OHDYHV RI 3KDVHROXVYXOJDULVDWGLIIHUHQWFDOFLXPVXSSO\LQUHODWLRQWRFDOFLXPR[DODWHFU\VWDOIRUPDWLRQ -RXUQDORI3ODQW3K\VLRORJ\±

$/(6=&=8.(6=&=8.$.67$1,6à$:(./0$=85.,(:&=$.$635=<.

UMCS