Carbon-Degrading Enzyme Activities Stimulated by Increased ... - Plos

&DUERQ'HJUDGLQJ(Q]\PH$FWLYLWLHV6WLPXODWHGE\ ,QFUHDVHG1XWULHQW$YDLODELOLW\LQ$UFWLF7XQGUD6RLOV $NLKLUR.R\DPD 0DWWKHZ':DOOHQVWHLQ5RGQH\76LPSVRQ-RKQ&0RRUH 1DWXUDO5HVRXUFH(FRORJ\/DERUDWRU\&RORUDGR6WDWH8QLYHUVLW\)W&ROOLQV&RORUDGR8QLWHG6WDWHVRI$PHULFD 'HSDUWPHQWRI%LRORJ\&RORUDGR6WDWH 8QLYHUVLW\ )W &ROOLQV &RORUDGR 8QLWHG 6WDWHV RI $PHULFD 'HSDUWPHQW RI (FRV\VWHP 6FLHQFH DQG 6XVWDLQDELOLW\ &RORUDGR 6WDWH 8QLYHUVLW\ )W &ROOLQV &RORUDGR8QLWHG6WDWHVRI$PHULFD

$EVWUDFW &OLPDWHLQGXFHGZDUPLQJRIWKH$UFWLFWXQGUDLVH[SHFWHGWRLQFUHDVHQXWULHQWDYDLODELOLW\WRVRLOPLFUREHVZKLFKLQ WXUQ PD\ DFFHOHUDWH VRLO RUJDQLF PDWWHU 620 GHFRPSRVLWLRQ :H LQFUHDVHG QXWULHQW DYDLODELOLW\ YLD IHUWLOL]DWLRQ WR LQYHVWLJDWHWKHPLFURELDOUHVSRQVHYLDVRLOHQ]\PHDFWLYLWLHV6SHFLILFDOO\ZHPHDVXUHGSRWHQWLDODFWLYLWLHVRIVHYHQ HQ]\PHVDW IRXUWHPSHUDWXUHV LQ WKUHH VRLO SURILOHVRUJDQLFRUJDQLFPLQHUDOLQWHUIDFHDQGPLQHUDO IURPXQWUHDWHG QDWLYHVRLOVDQGIURPVRLOVZKLFKKDGEHHQIHUWLOL]HGZLWKQLWURJHQ1 DQGSKRVSKRUXV3 VLQFH\HDUV DQG VL[ \HDUV )HUWLOL]HG SORWV ZLWKLQ WKH VLWH UHFHLYHG DQQXDO DGGLWLRQV RI J 1ഃPഃ\HDU DQG J 3ഃPഃ\HDU :LWKLQ WKH VLWH WZR IHUWLOL]HU UHJLPHV ZHUH HVWDEOLVKHG ± RQH LQ ZKLFK SORWV UHFHLYHG J 1ഃPഃ\HDU DQG J 3ഃPഃ\HDU DQG RQH LQ ZKLFK SORWV UHFHLYHG J 1ഃPഃ\HDU DQG J 3ഃPഃ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

(PDLODNLKLURNR\DPD#FRORVWDWHHGX

,QWURGXFWLRQ

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

3/2621(_ZZZSORVRQHRUJ

2FWREHU_9ROXPH_,VVXH_H

(Q]\PH$FWLYLWLHVLQ$UFWLF7XQGUD6RLOV

0DWHULDOVDQG0HWKRGV

7DEOH/LVWRIHQ]\PHVDVVD\HGLQWKLVVWXG\

6WXG\VLWHDQGVDPSOHFROOHFWLRQ (Q]\PH

$EEUHYLDWLRQ

ȕ*OXFRVLGDVH

%*

5HOHDVHVJOXFRVHIURPFHOOXORVH

&HOORELRK\GURODVH

&%

5HOHDVHVGLVDFFKDULGHVIURPFHOOXORVH

;\ORVLGDVH

;
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


6RLOSURSHUW\DQDO\VLV :H DQDO\]HG VRLO ZDWHU FRQWHQW 62& DQG WRWDO 1 FRQWHQWV IRUDOOVDPSOHV6RLOZDWHUFRQWHQWVZHUHGHWHUPLQHGE\GU\LQJ VRLOVDPSOHVDW&IRUKRXUV7RPHDVXUH62&DQGWRWDO 1FRQWHQWVVDPSOHVZHUHILUVWGULHGRXWDW&DQGJURXQG ILQHO\ XVLQJ D %ULQNPDQQ 5HWVFK PLOO +DDQ *HUPDQ\ 2UJDQLF & DQG WRWDO 1 FRQWHQWV RI WKH JURXQG VDPSOHV ZHUH TXDQWLILHG XVLQJ D /(&2 7UX6SHF /HFR &RUSRUDWLRQ 6W -RVHSK0LFKLJDQ86$

(Q]\PHDVVD\V :H TXDQWLILHG SRWHQWLDO DFWLYLWLHV RI VHYHQ K\GURO\WLF HQ]\PHV 7DEOH IRU HDFK VDPSOH XVLQJ IOXRURPHWULF WHFKQLTXHV >@ PRGLILHG IROORZLQJ 6WHLQZHJ HW DO >@ :H PHDVXUHG WKH DFWLYLWLHV RI IRXU HQ]\PHV K\GURO\]LQJ &ULFK VXEVWUDWHV %* &% ;@7KLVHTXDWLRQFDQEHH[SUHVVHG LQWKHIROORZLQJHTXDWLRQ

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Â/ RI &% DQG 3+26 IRU WKHRUJDQLFDQGLQWHUIDFHVRLOVDQGPROÂ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lQQHGRUI 6ZLW]HUODQG ZLWK D VHW RI ZDYHOHQJWK DW DQG QP IRU H[FLWDWLRQ DQG HPLVVLRQ UHVSHFWLYHO\ 5HDGLQJV RI WKH IOXRUHVFHQW DFWLYLWLHV IURP VWDQGDUGV ZHUH XVHG WR FDOFXODWH SRWHQWLDOHQ]\PHDFWLYLWLHVIRUHDFKVDPSOH $FWLYDWLRQ HQHUJ\ ZDV FDOFXODWHG XVLQJ SRWHQWLDO HQ]\PH DFWLYLWLHV DVVD\HG DW IRXU GLIIHUHQW WHPSHUDWXUHV IRU HDFK HQ]\PHDQGVDPSOH7HPSHUDWXUHVDQGDFWLYLW\YDOXHVZHUHILW LQWRWKH$UUKHQLXVHTXDWLRQ>@

lnK =

1 +lnA T

3ORWWLQJOQ.DJDLQVW7(DFDQEHFDOFXODWHGIURPWKHVORSH RI WKH OLQHDU UHJUHVVLRQ >@ $FWLYDWLRQ HQHUJ\ GRHV QRW KDYH HQHUJHWLFPHDQLQJEXWUDWKHUJDXJHVWHPSHUDWXUHGHSHQGHQFH RI HQ]\PH DFWLYLWLHV >@ $FWLYDWLRQ HQHUJ\ GHVFULEHV KRZ HQ]\PHDFWLYLW\LQFUHDVHVZLWKWHPSHUDWXUH>@WKXVSRVLWLYHO\ FRUUHODWHVZLWK4FDOFXODWHGIURPWKHVDPHGDWDVHW

6WDWLVWLFDODQDO\VHV :HXVHGDPL[HGHIIHFW$129$IRUVWDWLVWLFDODQDO\VHVZLWK WKH VLWHV LH WKH DQG VLWHV DQG IHUWLOL]DWLRQ OHYHOV LH FRQWURO ORZ DQG KLJK DV IL[HG HIIHFWV DQG EORFNV DV D UDQGRP HIIHFW $OO FRPSXWDWLRQV ZHUH FDUULHG RXW XVLQJ WKH SDFNDJH QOPH LQ 5 >@ $ VLJQLILFDQFH OHYHO RI S ZDV XVHGWRDVVHVVVWDWLVWLFDOVLJQLILFDQFHEHFDXVHRIWKHUHODWLYHO\ VPDOO VDPSOH VL]HV LQ WKLV VWXG\ DQG DOO SYDOXHV DUH IRU WZR VLGHGFRQILGHQFHLQWHUYDOV

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


7DEOH 6XPPDU\ RI SYDOXHV UHVXOWLQJ IURP PL[HGHIIHFW PRGHO DQDO\VHV IRU SRWHQWLDO HQ]\PH DFWLYLWLHV DVVD\HG DW &

,QGHSHQGHQW YDULDEOHV

%*

&%

;
@ DQG DQ LQYHUVH UHODWLRQVKLS EHWZHHQ SKRVSKDWDVH DFWLYLW\ DQG HQYLURQPHQWDO 3 DYDLODELOLW\ LV JHQHUDOO\ DFFHSWHG >@ 7KHVH UHVSRQVHV DUH FRQVLVWHQW ZLWK D UHVRXUFH HFRQRPLFVWKHRU\IRUHQ]\PHSURGXFWLRQE\VRLOPLFUREHV>@ ,Q FRQWUDVW WR &GHJUDGLQJ HQ]\PHV DQG SKRVSKDWDVH WKH IHUWLOL]DWLRQWUHDWPHQWVGLGQRWDIIHFWWZRHQ]\PHVWKDWGHJUDGH

7DEOH 6XPPDU\ RI SYDOXHV UHVXOWLQJ IURP PL[HGHIIHFW PRGHODQDO\VHVIRUVWRLFKLRPHWU\DVVD\HGDW&

,QGHSHQGHQWYDULDEOHV

&1

%*1

&3

%*3

13

)HUW

3URILOH

6LWH

)HUWî3URILOH

)HUWî6LWH

3URILOHî6LWH

)HUWî3URILOHî6LWH

& 1 DQG 3 UHSUHVHQW SRWHQWLDO HQ]\PH DFWLYLWLHV RI %*&%;@

6RLOSURSHUWLHVDQGSRWHQWLDOHQ]\PHDFWLYLWLHVDOWHUHG E\WKHIHUWLOL]DWLRQWUHDWPHQWV 2XUUHVXOWVVXSSRUWRXUILUVWK\SRWKHVLVWKDWIHUWLOL]DWLRQZRXOG LQFUHDVH SRWHQWLDO HQ]\PH DFWLYLWLHV WKDW GHJUDGH &ULFK VXEVWUDWHV)LJXUH$SSHQGL[6 7KLVUHVXOWVXJJHVWVWKDW SURGXFWLRQ RI WKHVH HQ]\PHV E\ VRLO PLFUREHV LV QXWULHQW OLPLWHGHVSHFLDOO\E\1>@'XULQJWKHJURZLQJVHDVRQQHW 1LPPRELOL]DWLRQ LV RIWHQ REVHUYHG LQ $UFWLF WXQGUD VRLOV >@ :DOOHQVWHLQ HW DO >@ REVHUYHG WKDW PRVW




)LJXUH$FWLYDWLRQHQHUJ\RISRWHQWLDOHQ]\PHDFWLYLWLHV3RWHQWLDOHQ]\PHDFWLYLWLHVPHDVXUHGDWIRXUWHPSHUDWXUHV DQG& ZHUHXVHGWRFDOFXODWHDFWLYDWLRQHQHUJ\IRUWKHWKUHHVRLOSURILOHVFROOHFWHGIURPWKHWZRVLWHV GRLMRXUQDOSRQHJ




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


7DEOH 6XPPDU\ RI SYDOXHV UHVXOWLQJ IURP PL[HGHIIHFW PRGHO DQDO\VHV IRU DFWLYDWLRQ HQHUJ\ DV LQGHSHQGHQW YDULDEOH

,QGHSHQGHQW YDULDEOHV

%*

&%

;
@

$FWLYDWLRQ(QHUJ\ :H GLG QRW ILQG VLJQLILFDQW IHUWLOL]DWLRQ HIIHFWV RQ DFWLYDWLRQ HQHUJ\LQDQ\RIWKHVHYHQHQ]\PHVDVVD\HG)LJXUH7DEOH 2XUILQGLQJRIQRFKDQJHLQDFWLYDWLRQHQHUJ\LQGLFDWHVWKDW FRPSRVLWLRQV RI LVRHQ]\PHV ZHUH QRW VLJQLILFDQWO\ DOWHUHG E\ WKH IHUWLOL]DWLRQ WUHDWPHQWV $ SRRO RI HQ]\PHV WKDW EUHDNV GRZQDSRO\PHULQDQDWXUDOHQYLURQPHQWFRQVLVWVRIDQXPEHU RI LVRIRUPV >@ 7KHVH LVRHQ]\PHV FDQ KDYH GLIIHUHQW DFWLYDWLRQ HQHUJ\ >@ WKXV D FKDQJH LQ WKH FRPSRVLWLRQ RI LVRHQ]\PHV ZRXOG OHDG WR FKDQJH LQ DFWLYDWLRQ HQHUJ\ 2XU ILQGLQJ LV VXUSULVLQJ JLYHQ WKDW IHUWLOL]DWLRQ WUHDWPHQWV RIWHQ FKDQJH FRPPXQLW\ FRPSRVLWLRQV RI VRLO PLFUREHV HJ >@ EXW VHH ZKLFK SURGXFH GLIIHUHQW LVRHQ]\PHV >@:DOOHQVWHLQHWDO>@UHSRUWHGWHPSHUDWXUHVHQVLWLYLW\ PHDVXUHG DV 4 FKDQJHG DORQJ D JURZLQJ VHDVRQ IRU VL[ HQ]\PHV LQ VRLOV FROOHFWHG IURP WKH VDPH HFRV\VWHP DV WKLV VWXG\ VXJJHVWLQJ FRPSRVLWLRQV RI LVRHQ]\PHV FKDQJHG RYHU WKH VDPSOLQJ WLPH 6WRQH HW DO >@ REVHUYHG QR FKDQJH LQ WHPSHUDWXUH VHQVLWLYLW\ RI SRWHQWLDO HQ]\PH DFWLYLWLHV LQ WHPSHUDWHIRUHVWVRLOVZKHQ1ZDVDPHQGHG $FWLYDWLRQ HQHUJLHV IRU PRVW RI WKH HQ]\PHV LQ WKH PLQHUDO VRLOV ZHUH ORZHU WKDQ WKRVH LQ WKH RUJDQLF RU LQWHUIDFH VRLOV )LJXUH7DEOH 2QHSRWHQWLDOH[SODQDWLRQIRUWKHGLIIHUHQFH LV FRPSRVLWLRQV RI LVRHQ]\PHV SURGXFHG LQ GLIIHUHQW VRLO SURILOHV)RULQVWDQFHRUJDQLFVRLOVZHUHVXEMHFWWRWHPSHUDWXUH LQFUHDVH GXULQJ D JURZLQJ VHDVRQ ZKHUHDV WKH PLQHUDO VRLOV ZH FROOHFWHG ZHUH NHSW UHODWLYHO\ FROG LQ WKH GHHSHU SURILOHV DERYH SHUPDIURVW 6XFK WHPSHUDWXUH GLIIHUHQFH LQ VRLO SURILOHV ZKHUH HQ]\PHV ZHUH SURGXFHG FRXOG FRQWULEXWH WR NLQHWLF FKDUDFWHULVWLFV RI HQ]\PHV WKH DFWLYDWLRQ HQHUJ\ RI SV\FKURSKLOLF HQ]\PHV LV ORZHU WKDQ WKDW RI PHVRSKLOLF HQ]\PHV >@ $QRWKHU SRWHQWLDO H[SODQDWLRQ LV VWDELOL]DWLRQ RI HQ]\PHV(Q]\PHVFDQEHVWDELOL]HGZLWKKXPLFDFLGV>@ WDQQLF DFLGV >@ DQG FOD\ PLQHUDOV >@ LQFUHDVLQJ DFWLYDWLRQ HQHUJ\ FRPSDUHG WR IUHH HQ]\PHV >@ +XPLF DQG



WDQQLF DFLGV LQ RUJDQLF DQG LQWHUIDFH VRLOV PLJKW FRQWULEXWH WR KLJKHU DFWLYDWLRQ HQHUJ\ WKDQ WKH PLQHUDO VRLOV 3V\FKURSKLOLF PLFUREHV SURGXFH PRUH H[RSRO\VDFFKDULGHV WKDQ PHVRSKLOLF PLFUREHV >@ ([RSRO\VDFFKDULGHV FDQ DOVR DIIHFW DFWLYDWLRQ HQHUJ\ DV H[RSRO\VDFFKDULGHV VWDELOL]H H[WUDFHOOXODU HQ]\PHV DQGSUHYHQWHQ]\PHGLIIXVLRQ>@

7,) $SSHQGL[ 6 3RWHQWLDO HQ]\PH DFWLYLWLHV LQ WKH PLQHUDO VRLOVDVVD\HGDWDQG& 7,) $SSHQGL[66XPPDU\RISYDOXHVUHVXOWLQJIURPPL[HG HIIHFW PRGHO DQDO\VHV IRU SRWHQWLDO HQ]\PH DFWLYLWLHV DVVHVVHG DW DQG & )HUW DQG 3URILOH UHSUHVHQW IHUWLOL]DWLRQ DQG VRLO SURILOH UHVSHFWLYHO\ SYDOXHV HTXDO WR RU OHVVWKDQDUHVKRZQEROG 3')

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

$SSHQGL[66XPPDU\RISYDOXHVUHVXOWLQJIURPPL[HG HIIHFW PRGHO DQDO\VHV IRU VWRLFKLRPHWU\ DVVHVVHG DW DQG & )HUW DQG 3URILOH UHSUHVHQW IHUWLOL]DWLRQ DQG VRLO SURILOH UHVSHFWLYHO\ SYDOXHV HTXDO WR RU OHVV WKDQ DUH VKRZQEROG 3') $SSHQGL[66WRLFKLRPHWU\RISRWHQWLDOHQ]\PHDFWLYLWLHV LQWKHRUJDQLFVRLOVDVVD\HGDWDQG& 7,) $SSHQGL[66WRLFKLRPHWU\RISRWHQWLDOHQ]\PHDFWLYLWLHV LQWKHLQWHUIDFHVRLOVDVVD\HGDWDQG& 7,) $SSHQGL[66WRLFKLRPHWU\RISRWHQWLDOHQ]\PHDFWLYLWLHV LQWKHPLQHUDOVRLOVDVVD\HGDQG& 7,)

6XSSRUWLQJ,QIRUPDWLRQ $SSHQGL[ 6 3RWHQWLDO HQ]\PH DFWLYLWLHV LQ WKH RUJDQLF VRLOV DVVD\HG DW DQG & 6WDWLVWLFDOO\ VLJQLILFDQW HIIHFWV IRXQG E\ PL[HGHIIHFW $129$ DUH VKRZQ LQ SDQHOV ) IHUWLOL]DWLRQ WUHDWPHQW FRQWURO ORZ DQG KLJK IHUWLOL]DWLRQV 6 VLWH LH ORQJ DQG VKRUWWHUP IHUWLOL]DWLRQ VLWHV DQG )î6 LQWHUDFWLRQEHWZHHQ)DQG66\PEROVLQGLFDWHS S

S 7,)

$FNQRZOHGJHPHQWV :H WKDQN *DLXV 5 6KDYHU /DXUD *RXJK DQG -HQQLH 5 0F/DUHQIRUFRRUGLQDWLQJILHOGZRUNDQG*UHJ6HOE\DQG-DVRQ /HYHUWRQ IRU VDPSOLQJ VRLOV /RJLVWLF VXSSRUW ZDV SURYLGHG E\ 7RROLN)LHOG6WDWLRQ8QLYHUVLW\RI$ODVND)DLUEDQNV86$

$XWKRU&RQWULEXWLRQV $SSHQGL[ 6 3RWHQWLDO HQ]\PH DFWLYLWLHV LQ WKH LQWHUIDFH VRLOV DVVD\HG DW DQG & 6WDWLVWLFDOO\ VLJQLILFDQW HIIHFWVIRXQGE\PL[HGHIIHFW$129$DUHVKRZQLQSDQHOV6 VLWH LH ORQJ DQG VKRUWWHUP IHUWLOL]DWLRQ VLWHV 6\PEROV LQGLFDWHS

&RQFHLYHG DQG GHVLJQHG WKH H[SHULPHQWV -&0 0': 3HUIRUPHG WKH H[SHULPHQWV $. 576 $QDO\]HG WKH GDWD $. &RQWULEXWHG UHDJHQWVPDWHULDOVDQDO\VLV WRROV $. 0': 576 -&0:URWHWKHPDQXVFULSW$.0':576-&0

5HIHUHQFHV 0DFN0&6FKXXU($*%UHW+DUWH066KDYHU*5&KDSLQ)6 (FRV\VWHP FDUERQ VWRUDJH LQ $UFWLF WXQGUD UHGXFHG E\ ORQJWHUP QXWULHQW IHUWLOL]DWLRQ 1DWXUH GRLQDWXUH 3XE0HG +DQVHQ - 6DWR 0 5XHG\ 5 /R . /HD ': HW DO *OREDO WHPSHUDWXUH FKDQJH 3URF 1DWO $FDG 6FL 8B6B$ GRLSQDV3XE0HG 6FKXXU($*9RJHO-*&UXPPHU.*/HH+6LFNPDQ-2HWDO 7KH HIIHFW RI SHUPDIURVW WKDZ RQ ROG FDUERQ UHOHDVH DQG QHW FDUERQ H[FKDQJHIURPWXQGUD1DWXUHGRLQDWXUH 3XE0HG

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




2HFKHO :& +DVWLQJV 6- 9RXUOUWLV * -HQNLQV 0 5LHFKHUV * HW DO 5HFHQW FKDQJH RI $UFWLF WXQGUD HFRV\VWHPV IURP D QHW FDUERQ GLR[LGHVLQNWRDVRXUFH1DWXUHGRLD .LUVFKEDXP 08) :LOO FKDQJHV LQ VRLO RUJDQLF FDUERQ DFW DV D SRVLWLYHRUQHJDWLYHIHHGEDFNRQJOREDOZDUPLQJ"%LRJHRFKHPLVWU\ GRL$ .QRUU : 3UHQWLFH ,& +RXVH -, +ROODQG ($ /RQJWHUP VHQVLWLYLW\ RI VRLO FDUERQ WXUQRYHU WR ZDUPLQJ 1DWXUH ± GRLQDWXUH3XE0HG +REELH6(1DGHOKRIIHU.-+|JEHUJ3 $V\QWKHVLV7KHUROHRI QXWULHQWV DV FRQVWUDLQWV RQ FDUERQ EDODQFHV LQ ERUHDO DQG $UFWLF UHJLRQV3ODQW6RLOGRL$ &KDSLQ )6 ,,, 6KDYHU *5 *LEOLQ $( 1DGHOKRIIHU .- /DXQGUH -$ 5HVSRQVHV RI $UFWLF WXQGUD WR H[SHULPHQWDO DQG REVHUYHG FKDQJHVLQFOLPDWH(FRORJ\GRL 6WXUP 0 +ROPJUHQ - 0F)DGGHQ -3 /LVWRQ *( &KDSLQ )6 HW DO 6QRZ±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


H[SHULPHQWDO GDWD DQG D PRGHO (FRORJ\ GRL >&(27&6@&2 *RXJK/5DPVH\($-RKQVRQ'5 3ODQW±KHUELYRUHLQWHUDFWLRQV LQ $ODVNDQ $UFWLF WXQGUD FKDQJH ZLWK VRLO QXWULHQW DYDLODELOLW\ 2LNRV GRLM[ 6DL\D&RUN .5 6LQVDEDXJK 5/ =DN '5 7KH HIIHFWV RI ORQJ WHUP QLWURJHQ GHSRVLWLRQ RQ H[WUDFHOOXODU HQ]\PH DFWLYLW\ LQ DQ $FHU VDFFKDUXP IRUHVW VRLO 6RLO %LRO %LRFKHP GRL 6 6WHLQZHJ-0'XNHV-6:DOOHQVWHLQ0' 0RGHOLQJWKHHIIHFWVRI WHPSHUDWXUH DQG PRLVWXUH RQ VRLO HQ]\PH DFWLYLW\ /LQNLQJ ODERUDWRU\ DVVD\V WR FRQWLQXRXV ILHOG GDWD 6RLO %LRO %LRFKHP GRL MVRLOELR $UUKHQLXV 6 hEHU GLH 5HDNWLRQVJHVFKZLQGLJNHLW EHL GHU ,QYHUVLRQ YRQ 5RKU]XFNHU GXUFK 6l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|P - HW DO /RQJWHUP LPSDFW RI IHUWLOL]DWLRQ RQ DFWLYLW\ DQG FRPSRVLWLRQ RI


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