Beruflich Dokumente
Kultur Dokumente
DOI:
10.1073/pnas.1703375114
Published: 12/09/2017
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Submission PDF
GROWTH RATES -OST POPULATIONS CONVERGE ON THE SAME GROWTH RATE XJMM CF DPNNPO JO BRVBUJD FOWJSPONFOUT
XJUI DPNCJOBUJPOT BOE
BY THE END OF THE EVOLUTION EXPERIMENT (OWEVER POPULATIONS
JOUFOTJUJFT PG ESJWFST IBWJOH TJHOJGJDBOU SFHJPOBM WBSJBUJPO
TP JU
ADAPT MORE WHEN THESE DOMINANT DRIVERS OCCUR IN THE PRESENCE
JT WJUBM UIBU XF VOEFSTUBOE UIF KPJOU DPOUSJCVUJPOT PG UIF JEFOUJUZ
OF OTHER DRIVERS 4HIS IS DUE TO AN INCREASE IN THE INTENSITY OF
OVNCFS
BOE JOUFOTJUZ PG ESJWFST UP USBJU FWPMVUJPO JO QSJNBSZ
SELECTION IN ENVIRONMENTS WITH MORE DRIVERS WHICH ARE MORE
QSPEVDFST 5XP OPOFYDMVTJWF NFDIBOJTNT DPVME DBVTF OBUVSBM TF
LIKELY TO INCLUDE DOMINANT DRIVERS #ONCURRENTLY MANY OF THE TRAIT
MFDUJPO UP BDU EJGGFSFOUMZ JO DBTFT PG DPNQMFY NVMUJESJWFS
WFSTVT
CHANGES THAT OCCUR DURING THE INITIAL SHORT
TERM RESPONSE TO BOTH
TJNQMF TJOHMF ESJWFS
FOWJSPONFOUBM DIBOHF 'JSTU
JG UIF OVNCFS
SINGLE AND MULTIDRIVER ENVIRONMENTAL CHANGE REVERT AFTER ABOUT
PG JOEFQFOEFOU USBJUT VOEFS TFMFDUJPO JODSFBTFT BT UIF OVNCFS
GENERATIONS OF EVOLUTION )N FUTURE AQUATIC ENVIRONMENTS
PG ESJWFST JO UIF FOWJSPONFOU JODSFBTFT
QMFJPUSPQJD JOUFSBDUJPOT
POPULATIONS WILL ENCOUNTER DIFFERING COMBINATIONS OF DRIVERS AND
DPVME MJNJU BEBQUBUJPO JO DPNQMFY FOWJSPONFOUT NPSF UIBO JO
INTENSITIES OF SELECTION WHICH WILL ALTER THE ADAPTIVE POTENTIAL OF
TJNQMF POFT
4FDPOE
JG TFMFDUJPO JOUFOTJUZ JODSFBTFT BT UIF
PRIMARY PRODUCERS !CCURATELY GAUGING THE INTENSITY OF SELECTION
OVNCFS PG ESJWFST JODSFBTFT
UIFO UIF QSPCBCJMJUZ PG QPQVMBUJPO
ON KEY PRIMARY PRODUCERS WILL HELP IN PREDICTING POPULATION SIZE
FYUJODUJPO JODSFBTFT XJUI UIF OVNCFS PG ESJWFST
CVU TVSWJWJOH
AND TRAIT EVOLUTION AT THE BASE OF AQUATIC FOOD WEBS
QPQVMBUJPOT XJMM BEBQU NPSF BOE NPSF SBQJEMZ JO FOWJSPONFOUT
XJUI NPSF ESJWFST 5IFTF UXP NFDIBOJTNT IBWF EJGGFSFOU FDPMPH
MULTIPLE ENVIRONMENTAL DRIVERS | ADAPTATION | MICROBIAL EVOLUTION | JDBM DPOTFRVFODFT *G EJGGFSFODFT JO FWPMVUJPOBSZ SFTQPOTFT BSF
#HLAMYDOMONAS NBJOMZ EVF UP EJGGFSFODFT JO QMFJPUSPQZ VOEFS NVMUJESJWFS DIBOHF
)NTRODUCTION
" HSPXJOH CPEZ PG FWJEFODF GSPN FYQFSJNFOUT TIPXT UIBU GVOD 3IGNIlCANCE
UJPOBM USBJUT JO BRVBUJD QSJNBSZ QSPEVDFST DBO CF BMUFSFE CZ FWPMV
UJPO JO UIF GBDF PG HMPCBM DIBOHF
XIJDI IBT UP EBUF CFFO FYQMPSFE /UR UNDERSTANDING OF HOW PRIMARY PRODUCERS AT THE BASE OF
JO UFSNT PG ESJWFST TVDI BT UFNQFSBUVSF
PS $0 MFWFMT
AQUATIC ECOSYSTEMS RESPOND TO COMPLEX ENVIRONMENTAL CHANGE
5IFTF TUVEJFT JOWFTUJHBUF UIF SFTQPOTFT PG QSJNBSZ QSPEVDFST UP CURRENTLY DEPENDS ON STUDIES USING FEW ENVIRONMENTAL DRIVERS
TJOHMF BTQFDUT PG HMPCBM DIBOHF
BOE UIF SFTVMUT BSF PGUFO OR SCENARIOS WHERE DRIVERS COVARY (OWEVER WE LACK A GEN
ERAL UNDERSTANDING OF EVOLUTION IN MULTIDRIVER ENVIRONMENTS
VTFE UP VOEFSTUBOE IPX DIBOHFT UP UIF CJPMPHJDBM DPNQPOFOU PG 7E EVOLVE A MICROBIAL PRIMARY PRODUCER IN NINETY
SIX DIF
OVUSJFOU DZDMJOH
JODMVEJOH BJSXBUFS DBSCPO FYDIBOHF
XJMM CF JN FERENT MULTIDRIVER ENVIRONMENTS AND lND THAT EVOLUTIONARY
QBDUFE
3FDFOU TIPSUUFSN TUVEJFT TIPX UIBU JO NVMUJESJWFS RESPONSES IN GROWTH ARE LARGELY DRIVEN BY A FEW DRIVERS BUT
FOWJSPONFOUT
UIF NBKPSJUZ PG UIF PSHBOJTNBM SFTQPOTF JT PGUFO THAT THE INTENSITY OF SELECTION IS ON AVERAGE HIGHER IN MUL
TIDRIVER ENVIRONMENTS &UNCTIONAL TRAITS CELL SIZE CHLOROPHYLL
FYQMBJOFE CZ POF PS UXP ESJWFST
8F DBMM UIFTF EPNJOBOU CONTENT OFTEN REVERT TO ANCESTRAL VALUES DURING ADAPTATION
ESJWFST
5IJT TUVEZ JOWFTUJHBUFT IPX UIF FWPMVUJPOBSZ SFTQPOTFT IN MULTIDRIVER ENVIRONMENTS 4HIS EXPANDS THE FRAMEWORK FOR
UP EPNJOBOU FOWJSPONFOUBM ESJWFST EFQFOET PO UIF PUIFS ESJWFST UNDERSTANDING HOW MICROBIAL PRIMARY PRODUCERS EVOLVE UNDER
QSFTFOU 5IJT IFMQT MJOL QBUUFSOT PG DPNQMFY NVMUJESJWFS
FOWJ GLOBAL CHANGE AND THE POTENTIAL RAMIlCATIONS FOR THEIR FUNC
TION IN AQUATIC ECOSYSTEMS
SPONFOUBM DIBOHF UP UIF FWPMVUJPOBSZ QPUFOUJBM PG QSJNBSZ QSP
EVDFST
BOE JOGPSNT UIF EFTJHO PG GVUVSF FYQFSJNFOUT 8F EP 2ESERVED FOR 0UBLICATION &OOTNOTES
UIJT VTJOH BO FWPMVUJPO FYQFSJNFOU UIBU EJTFOUBOHMFT UIF FGGFDUT
PG ESJWFS OVNCFS BOE JEFOUJUZ PO USBJU FWPMVUJPO JO NVMUJESJWFS
FOWJSPONFOUT JO B TJOHMF DFMMFE BMHB
" TNBMM OVNCFS PG FYQFSJNFOUT IBWF JOWFTUJHBUFE UIF QMBTUJD
BOE FWPMVUJPOBSZ SFTQPOTFT UP NVMUJQMF ESJWFST UIVT GBS
–
5IFTF TUVEJFT DPOTJTUFOUMZ TIPX UIBU CPUI QMBTUJD BOE FWPMVUJPOBSZ
SFTQPOTFT UP QBJST PG ESJWFST EJGGFS GSPN SFTQPOTFT UP FJUIFS TJOHMF
ESJWFS *O UIF TIPSUUFSN
UIF FGGFDU PG NVMUJESJWFS FOWJSPONFOUT
PO QMBTUJD SFTQPOTFT DIBOHFT JO QIFOPUZQF JO SFTQPOTF UP BO
FOWJSPONFOUBM DVF UIBU EPFT OPU SFRVJSF DIBOHF JO UIF HFOFUJD
Submission PDF
QIZMM BVUPGMVPSFTDFODF QFS DFMM WPMVNF JO FWPMWFE QPQVMBUJPOT QSPEVDFST NBZ FWPMWF VOEFS EJGGFSFOU DMJNBUF DIBOHF DMVTUFST
JO UIFJS PXO SFHJNFT EJE OPU EJGGFS GSPN UIBU PG FWPMWFE DPOUSPM
UP HSPVQ ESJWFST CBTFE PO UIFJS FGGFDUT PO HSPXUI QPTJUJWF
OFHBUJWF
OFVUSBM
GPS EJGGFSFOU UBYB XIFO BTTFTTJOH XIFUIFS PS
QPQVMBUJPOT JO UIF DPOUSPM FOWJSPONFOU ± µN NFBO
OPU XF FYQFDU DMJNBUF DIBOHF DMVTUFST UP ESJWF FWPMVUJPO
± 4%
5IJT JT DPOTJTUFOU XJUI QPQVMBUJPOT IBWJOH BEBQUFE UP UIFJS
FOWJSPONFOUT
BT UIFZ OP MPOHFS TIPX B TUBOEBSE TJHO PG TUSFTT 0VS BQQSPBDI DPNQMFNFOUT TDFOBSJPCBTFE TUVEJFT 'PS FY
–
BNQMF
#PZE FU BM
NPEFMFE SFHJPOBM DIBOHFT UP NVMUJESJWFS
SFHJNFT
BOE VTFE NFBTVSFE TIJGUT JO QIZUPQMBOLUPO QIZTJPMPHZ
$ISCUSSION
UP NBLF RVBMJUBUJWF QSFEJDUJPOT BCPVU TIJGUT JO CJPHFPHSBQIZ
5P VOEFSTUBOE IPX UIF FWPMVUJPOBSZ SFTQPOTF PG QSJNBSZ QSPEVD 5IFZ EFUBJM UIF SFTQPOTFT PG DPDDPMJUIPQIPSFT DBMDJGJFST
BOE
FST EFQFOET PO UIF JEFOUJUZ BOE OVNCFS PG ESJWFST JO NVMUJESJWFS EJBUPNT TJMJDJGJFST
JO UXP IJHI MBUJUVEF PDFBO QSPWJODFT UP TIJGUT
FOWJSPONFOUT
XF FWPMWFE NJDSPCJBM QPQVMBUJPOT JO OJOFUZTJY JO UFNQFSBUVSF
$0
QIPUPTZOUIFUJDBMMZ BDUJWF SBEJBUJPO 1"3
VOJRVF TJOHMF BOE NVMUJESJWFS FOWJSPONFOUT #PUI UIF BCTPMVUF JSPO
TJMJDBUF
OJUSBUF BOE QIPTQIBUF
BT XFMM BT JOUFSBDUJPOT CF
HSPXUI SBUF PG UIF FWPMWFE QPQVMBUJPOT BOE UIF EJSFDU SFTQPOTF UP UXFFO ESJWFS QBJST #BTFE PO QMBTUJD SFTQPOTFT
UIFZ TVHHFTU UIBU
TFMFDUJPO UIF BNPVOU PG FWPMVUJPO OFFEFE UP SFBDI UIBU HSPXUI FMFWBUFE UFNQFSBUVSF JT MJLFMZ UP DBVTF B QPMFXBSE TIJGU JO DPD
SBUF
XFSF FYQMBJOFE CZ UIF QSFTFODF PG B GFX EPNJOBOU FOWJ DPMJUIPQIPSFT
BOE IJHI 1"3
MPX QIPTQIBUF
BOE MPX TJMJDBUF
SPONFOUBM ESJWFST 4VSQSJTJOHMZ
UIF NVMUJESJWFS DPOUFYU JO XIJDI BSF MJLFMZ UP GBWPS DPDDPMJUIPQIPSFT PWFS EJBUPNT " EFDSFBTF
EPNJOBOU ESJWFST PDDVSSFE IBE MJUUMF FGGFDU PO UIF HSPXUI SBUF JO DBMDJGJDBUJPO JT QSFEJDUFE GPS DPDDPMJUIPQIPSFT BOE B EFDSFBTF
PG FWPMWFE QPQVMBUJPOT PO BWFSBHF )PXFWFS
QPQVMBUJPOT IBE B JO TJMJDJGJDBUJPO GPS EJBUPNT 0VS TUVEZ TVHHFTUT UIBU JO BEEJUJPO
MBSHFS EJSFDU SFTQPOTF UP TFMFDUJPO JO FOWJSPONFOUT UIBU DPOUBJOFE TFMFDUJPO UP JODSFBTF CPUI DBMDJGJDBUJPO BOE TJMJDJGJDBUJPO DPVME
NPSF ESJWFST
JOEJDBUJOH UIBU QSJNBSZ QSPEVDFST FWPMWF NPSF JO ESJWF FWPMVUJPO JO NVMUJESJWFS FOWJSPONFOUT 8F BMTP FYQFDU UIBU
SFTQPOTF UP EPNJOBOU ESJWFST XIFO UIFZ PDDVS JO NVMUJESJWFS FWPMVUJPO SFWFSTF TPNF PG UIF USBJU DIBOHF QSFEJDUFE CBTFE PO
FOWJSPONFOUT UIBO XIFO UIFZ PDDVS TJOHMZ 5IJT JT MBSHFMZ CFDBVTF UIF #PZE FU BM
"EBQUBUJPO DPVME BMTP SFTVMU JO QPTJUJWF
PO BWFSBHF
TFMFDUJPO JT TUSPOHFS JO NVMUJESJWFS FOWJSPONFOUT BOE OFHBUJWF
DIBOHFT JO HSPXUI SBUFT FWFOUVBMMZ BUUFOVBUJOH
5IVT
QPQVMBUJPOT FWPMWJOH JO NVMUJESJWFS FOWJSPONFOUT BEBQU CBTFE PO EBUB GSPN TJOHMFESJWFS FWPMVUJPO FYQFSJNFOUT
NPSF
CVU BSSJWF BU UIF TBNF GJOBM HSPXUI SBUFT BT QPQVMBUJPOT 5IJT JMMVTUSBUFT IPX XF DBO VTF BO VOEFSTUBOEJOH PG IPX OBUVSBM
FWPMWJOH JO TJOHMF ESJWFS FOWJSPONFOUT XJUI UIF TBNF EPNJOBOU TFMFDUJPO BDUT UP SFGJOF QSFEJDUJPOT PG USBJU DIBOHF JO NVMUJESJWFS
ESJWFST FOWJSPONFOUT
8F XFSF JOJUJBMMZ TVSQSJTFE CZ UIFTF SFTVMUT 8F IBE IZQPUI %FTQJUF TPNF QPQVMBUJPOT IBWJOH B MBSHF EJSFDU SFTQPOTF UP
FTJ[FE UIBU QMFJPUSPQJD DPOTUSBJOUT XPVME CF NPSF JNQPSUBOU JO TFMFDUJPO
FWPMVUJPO SFWFSTFE QMBTUJD DIBOHFT JO TFWFSBM USBJUT 5IJT
QPQVMBUJPOT FWPMWJOH JO FOWJSPONFOUT XJUI NPSF ESJWFST CFDBVTF TVHHFTUT UIBU BT QPQVMBUJPOT BEBQU UP NVMUJESJWFS FOWJSPONFOUT
NPSF USBJUT XPVME CF VOEFS TFMFDUJPO
*OTUFBE
UIF SFTQPOTF UIF GVODUJPO PG FWPMWJOH HSPVQT NBZ DIBOHF MFTT UIBO FYQFDUFE
UP TFMFDUJPO JODSFBTFT XJUI UIF OVNCFS PG FOWJSPONFOUBM ESJWFST CBTFE PO QMBTUJD SFTQPOTFT UP NVMUJESJWFS FOWJSPONFOUT QMBTUJD
"CPWF B UISFTIPME OVNCFS PG ESJWFST TFWFO
JO UIJT FYQFSJNFOU
SFTQPOTFT BSF DJSDMFT JO 'JH
1PQVMBUJPO HSPXUI SBUFT
IPXFWFS
SBQJE BEBQUBUJPO XBT OPU QPTTJCMF
BOE QPQVMBUJPOT XFOU FYUJODU BSF OPU SFTUPSFE UP DPOUSPM WBMVFT JO BMM SFHJNFT 4JODF UIF QPQV
5IFTF SFTVMUT TVHHFTU UIBU UIF OVNCFS PG USBJUT VOEFS TFMFDUJPO MBUJPOT JO UIJT FYQFSJNFOU BSF QSPQBHBUFE CZ CBUDI DVMUVSF BOE
EPFT OPU TDBMF XJUI UIF OVNCFS PG FOWJSPONFOUBM ESJWFST 5IJT XFSF OPU BMMPXFE UP SFBDI DBSSZJOH DBQBDJUZ
PWFSBMM TFMFDUJPO XBT
NBLFT TFOTF HJWFO UIBU FWPMVUJPOBSZ SFTQPOTFT BSF ESJWFO CZ B GPS JODSFBTFE JO DFMM EJWJTJPO SBUFT
5IF MPXFS HSPXUI SBUFT
GFX EPNJOBOU ESJWFST $0
MPX QIPTQIBUF
UFNQFSBUVSF
BOE JO TPNF SFHJNFT NBZ CF UIF SFTVMU PG QIZTJPMPHJDBM DPOTUSBJOUT
IFSCJDJEF
TP UIBU UIF USBJUT VOEFS TFMFDUJPO NBZ CF NPSF PS MFTT TJODF SFTPVSDFT DBO MJNJU HSPXUI SBUFT FWFO BGUFS BEBQUBUJPO
DPOTUBOU PWFS UIF SFHJNFT DPOUBJOJOH B HJWFO EPNJOBOU ESJWFS 'PS FYBNQMF
HSPXUI SBUFT BSF MPXFS JO MPX QIPTQIBUF SFHJNFT
"T UIF OVNCFS PG ESJWFST JODSFBTFT
EPNJOBOU ESJWFST BSF NPSF UIBO JO QIPTQIBUF SFQMFUF POFT SFHBSEMFTT PG ESJWFS OVNCFS 'JH
MJLFMZ UP CF QSFTFOU 5IF JEFOUJUZ PG EPNJOBOU ESJWFST JO FBDI 4
%FTQJUF UIJT
UIF QSPQPSUJPO PG DFMMT TIPXJOH TJHOT PG TUSFTT
SFHJNF QBSUJBMMZ FYQMBJOT UIF TNBMM EJGGFSFODFT JO UIF SFTQPOTF UP DIMPSPQIZMM OFHBUJWF BOEPS TNBMM DFMMT
JO FWPMWFE QPQVMBUJPOT
TFMFDUJPO JO NVMUJESJWFS FOWJSPONFOUT 5IF PWFSSJEJOH FGGFDU PG JO UIF MPX QIPTQIBUF BOE MPXOVUSJFOU FOWJSPONFOUT SFUVSOFE
UIF EPNJOBOU ESJWFST JT DPOTJTUFOU XJUI UIF BDDMJNBUJPO SFTQPOTF UP DPOUSPM WBMVFT 'JH
*O DPOUSBTU
QPQVMBUJPOT FWPMWFE JO
5BCMF
XJUI PUIFS TUVEJFT UIBU EFNPOTUSBUFE UIBU JO QPPSRVBMJUZ FOWJSPO 4
5BCMF 4
5IF BODFTUSBM QPQVMBUJPO XBT TQMJU JOUP QPQ
NFOUT QPQVMBUJPOT JOWFTU JO NBJOUBJOJOH DFMM DPOEJUJPO
*O VMBUJPOT 'JHVSF
5BCMF 4
" TJOHMF GPVOEFS DPMPOZ FOTVSFT
$ SFJOIBSEUJJ
DFMM TJ[F JT SFMBUFE UP QPQVMBUJPO HSPXUI UISPVHI JUT UIBU QPQVMBUJPO FWPMVUJPO VTFT EF OPWP WBSJBUJPO &OWJSPONFOUBM
FGGFDU PO DFMM EJWJTJPO
BT B DSJUJDBM TJ[F NVTU CF SFBDIFE CFGPSF DIBOHFT PDDVSSFE JO POF TUFQ BU UIF CFHJOOJOH PG UIF FYQFSJNFOU
DFMMT EJWJEF –
5IJT USFOE PG QIFOPUZQJD SFWFSTJPO EVSJOH DPOUSPM MFWFM JO CSBDLFUT
JODSFBTFE $0 UP QQN QQN
BEBQUJWF FWPMVUJPO IBT BMTP CFFO TFFO JO IJHI $0 FOWJSPONFOUT UFNQFSBUVSF UP P $ P $
EFDSFBTFE Q) UP Q)
MJHIU
JOEJDBUJOH UIBU TPNF QIFOPUZQJD SFWFSTJPO NBZ CF B DPNNPO MFWFMT UP μNPM N T μNPM N T
SFEVDFE QIPTQIBUF
PVUDPNF PG FWPMVUJPO
UP N. N.
OVUSJFOU EFQMFUJPO UP OV
8F EJE OPU FYBNJOF UIF NPMFDVMBS CBTJT PG USBJU SFWFSTJPO USJFOUT 5BCMF 4
µ. PG BUSB[JOF OP IFSCJDJEF
3FHJNFT
IFSF
CVU PGGFS UXP OPOFYDMVTJWF FYQMBOBUJPOT 'JSTU
PVS SFTVMUT XJUI 67 XFSF EPTFE XJUI ,+N 67# UXJDF B XFFL 5BCMF
BSF DPOTJTUFOU XJUI DPNQFOTBUPSZ NVUBUJPOT BGGFDUJOH USBJU FWPMV 4
1PQVMBUJPOT FWPMWFE JO FOWJSPONFOUT XJUI BU MFBTU POF ESJWFS
UJPO JO NVMUJESJWFS FOWJSPONFOUT 'PMMPXJOH B MBSHF ESPQ JO QPQV BSF NVMUJESJWFSFWPMWFE QPQVMBUJPOT 1PQVMBUJPOT FWPMWFE JO UIF
Submission PDF
MBUJPO GJUOFTT
UIF GJSTU CFOFGJDJBM NVUBUJPO GJYFE PGUFO IBT B MBSHF DPOUSPM FOWJSPONFOU BSF DPOUSPM QPQVMBUJPOT "MM QPQVMBUJPOT
FGGFDU
BOE DBO DIBOHF TFWFSBM USBJUT TJNVMUBOFPVTMZ 5IJT NBZ XFSF QSPQBHBUFE CZ CBUDI DVMUVSF 4FF 4* "QQFOEJY GPS EFUBJMT
CF GPMMPXFE CZ DPNQFOTBUPSZ NVUBUJPOT UIBU JODSFBTF GJUOFTT CZ
"TTBZT PG QPQVMBUJPO HSPXUI SBUFT
SFWFSTJOH UIF FGGFDU PG UIF JOJUJBM NVUBUJPO PO USBJUT XIFSF DIBOHF
"O BDDMJNBUJPO QFSJPE PG POF USBOTGFS DZDMF XBT VTFE GPS BMM
XBT OPU BEBQUJWF –
4FDPOE
IFSJUBCMF FQJHFOFUJD NVUBUJPOT
BTTBZT TFF 4*
PS USBOTHFOFSBUJPOBM QMBTUJDJUZ DBO DPOUSJCVUF UP FBSMZ BEBQUBUJPO
5IF BWFSBHF SBUF PG DFMM EJWJTJPO QFS EBZ XBT DBMDVMBUFE PWFS
CVU FWFOUVBMMZ CF SFQMBDFE CZ HFOFUJD NVUBUJPOT –
6O B TJOHMF CBUDI DVMUVSFMFOHUI UJNF VTJOH FRVBUJPO
4FF 4*
EFSTUBOEJOH IPX USBJU SFWFSTJPO JT MJOLFE UP BEBQUBUJPO QSFTFOUT BO
"QQFOEJY GPS EFUBJMT
PQQPSUVOJUZ UP JNQSPWF PVS QSFEJDUJPOT PG GVODUJPOBM USBJU WBMVFT
JO QSJNBSZ QSPEVDFST JO BRVBUJD FOWJSPONFOUT
-BCPSBUPSZ FWPMVUJPO FYQFSJNFOUT VTF TJNQMJGJFE FOWJSPO
NFOUT BOE QPQVMBUJPOT UP HBJO JOTJHIUT JOUP UIF GVOEBNFOUBM
BDUJPO PG OBUVSBM TFMFDUJPO "QQMZJOH UIF SFTVMUT PG UIJT FYQFSJNFOU
()
UP OBUVSBM QIZUPQMBOLUPO QPQVMBUJPO SFRVJSFT UBLJOH JOUP BDDPVOU
IPX QPQVMBUJPO TJ[F BOE EJWFSTJUZ BNPOH PUIFS GBDUPST
TVDI BT /U JT UIF DFMM EFOTJUZ DFMMT NM
BU UJNF U IPVST
BOE / JT UIF DFMM
SFDPNCJOBUJPO SBUFT
BGGFDU BEBQUBUJPO
1SFWJPVT XPSL IBT EFOTJUZ BU UJNF U
TIPXO UIBU IJHIFS TUBOEJOH HFOFUJD WBSJBUJPO DBO BMMPX BEBQUJOH 5IF EJSFDU SFTQPOTF UP TFMFDUJPO XBT NFBTVSFE CZ DPNQBSJOH
QPQVMBUJPOT UP FWPMWF GBTUFS –
4JNJMBSMZ
SFDPNCJOBUJPO UIF HSPXUI PG B NVMUJESJWFSFWPMWFE QPQVMBUJPO BOE B DPOUSPM
BOE QMBTUJDJUZ
DBO CPUI TQFFE VQ BEBQUBUJPO
5IF QPQVMBUJPO JO UIF TBNF NVMUJESJWFS SFHJNF 'JH
5IF EJSFDU
QPXFS PG TJNQMJGJFE MBCPSBUPSZ FYQFSJNFOUT MJFT JO QSPWJEJOH JO SFTQPOTF UP TFMFDUJPO XBT DBMDVMBUFE VTJOH FRVBUJPO
BOE TDBMFE
TJHIU JOUP IPX OBUVSBM TFMFDUJPO XPSLT BQQMZJOH UIFTF JOTJHIUT SF UP UIF OVNCFS PG EJWJTJPOT E
PG UIF DPOUSPM QPQVMBUJPO JO UIF
RVJSFT BDDPVOUJOH GPS PUIFS QSPDFTTFT UIBU DBO NPEJGZ FWPMVUJPO
SFMFWBOU SFHJNF 4FF 4* "QQFOEJY GPS EFUBJMT
BOE GPS UIF TQFDJGJD CJPMPHZ PG XJME QPQVMBUJPOT
#ONCLUSIONS
()
8F TIPX UIBU QPQVMBUJPOT BEBQU NPSF JO SFTQPOTF UP EPNJOBOU
ESJWFST XIFO UIPTF ESJWFST PDDVS JO B NVMUJESJWFS DPOUFYU
VOUJM & JT UIF OVNCFS PG EJWJTJPOT E
PG NVMUJESJWFSFWPMWFE QPQVMB
FOWJSPONFOUT EFUFSJPSBUF FOPVHI UP DBVTF FYUJODUJPOT "MPOHTJEF UJPOT JO UIFJS SFHJNFT
BOE $ JT UIF OVNCFS PG EJWJTJPOT E
PG
UIJT
BEBQUBUJPO DBO SFTVMU JO UIF SFWFSTJPO PG UIF JOJUJBM DIBOHFT FWPMWFE DPOUSPM QPQVMBUJPOT JO UIF TBNF SFHJNFT
UP USBJU WBMVFT JO NVMUJESJWFS FOWJSPONFOUT 8IJMF XF FYQFDU UIBU 'MPX DZUPNFUSJD BOBMZTJT PG QIZTJPMPHJDBM QBSBNFUFST
UIF SFTVMU PG FWPMVUJPO CFJOH ESJWFO QSJNBSJMZ CZ B GFX EPNJOBOU "O BDDMJNBUJPO QFSJPE PG POF USBOTGFS DZDMF XBT VTFE GPS
ESJWFST JO NVMUJESJWFS FOWJSPONFOUT UP CF HFOFSBM
UIF JEFOUJUZ BMM BTTBZT
BT BCPWF " '"$4 $"/50 XBT VTFE UP EFUFSNJOF
PG EPNJOBOU ESJWFST XJMM CF PSHBOJTN BOE DPOUFYU EFQFOEFOU
SFE BVUPGMVPSFTDFODF DIMPSPQIZMM B BOE C
FWFOU OVNCFS DFMM
TVDI UIBU B WBSJFUZ PG BQQSPBDIFT MBSHF GBDUPSJBM FYQFSJNFOUT MJLF EFOTJUZ
BOE GPSXBSE TDBUUFS DFMM TJ[F
4FF 4* "QQFOEJY
UIJT POF
TDFOBSJPCBTFE NPEFMT BOE FYQFSJNFOUT
UBYBTQFDJGJD 4UBUJTUJDBM BOBMZTJT
QIZTJPMPHJDBM BOE FWPMVUJPO TUVEJFT
BSF OFFEFE UP VOEFSTUBOE 5IF FGGFDUT PG J
ESJWFS JEFOUJUZ BOE JJ
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