Phenotypic responses to lifelong hypoxia in cichlids
Rutjes, Hendrikus Antonius
Citation
Rutjes, H. A. (2006, October 24). Phenotypic responses to lifelong hypoxia in cichlids.
Retrieved from https://hdl.handle.net/1887/4925
Version:
Corrected Publisher’s Version
License:
Licence agreement concerning inclusion of doctoral thesis in the
Institutional Repository of the University of Leiden
Downloaded from:
https://hdl.handle.net/1887/4925
3KHQRW\SLFUHVSRQVHVWROLIHORQJK\SR[LDLQFLFKOLGV &KDSWHU
$%675$&7
'UDPDWLFLQFUHDVHLQJLOOVXUIDFHRIDK\SR[LDUDLVHG/DNH9LFWRULDFLFKOLG
,1752'8&7,21
,QWKHODVWFHQWXU\RFFXUUHQFHRIFKURQLF K\SR[LD FDXVHG E\ KXPDQLQGXFHG SHUWXUEDWLRQV EHFDPH LQFUHDVLQJO\ FRPPRQLQZDWHUERGLHVZRUOGZLGH:X 5HGG\7KLVLVDOVRWKHFDVH LQ /DNH9LFWRULD ZKHUH GXULQJ WKH SDVW GHFDGHV 2 FRQFHQWUDWLRQV GHFUHDVHG FRQVLGHUDEO\ GXH WR HXWURSKLFDWLRQ 2FKXPEDDQG.LEDUDD.DXIPDQ +HFN\HWDO:DQLQNHWDO :LWWH HW DO ,Q DW D PGHHS VDPSOLQJ VWDWLRQ LQ WKH 0ZDQ]D *XOI K\SR[LF FRQGLWLRQV PJ / ZHUH SUHVHQW QHDU WKH ERWWRP
IRU VHYHUDO GD\V RQO\ GXULQJ WKH ORQJ UDLQ\ VHDVRQ YDQ 2LMHQ HW DO +RZHYHU GXULQJ WKH UDLQ\ VHDVRQ LQ DW WKH VDPH VDPSOLQJ VWDWLRQ PRUH VHYHUH K\SR[LD PJ / ZDV
SUHVHQW IRU VHYHUDO PRQWKV 0RUHRYHU K\SR[LFZDWHUOD\HUVUHDFKHGKLJKHULQWR WKHZDWHUFROXPQ:DQLQNHWDO 6WXGLHV RQ VHYHUDO VSHFLHV RI /DNH 9LFWRULD FLFKOLGV VKRZHG WKDW QRQ DFFOLPDWHGDGXOW¿VKZKLFKDUHJUDGXDOO\ H[SRVHG WR DLU VDWXUDWLRQ FD PJ / ZLWKLQ D IHZ KRXUV VXUYLYH
OHVVWKDQRQHGD\5XWMHVXQSXEOLVKHG 7KLV LV QRUPDO IRU PRVW WHOHRVWV 9DQ GHQ 7KLOODUW DQG 9DQ :DDUGH %URRGVRIFLFKOLGVWKDWZHUHH[SRVHGWR LQFUHDVLQJOHYHOVRIK\SR[LDRYHUVHYHUDO ZHHNV FRXOG KRZHYHU EH UDLVHG XQGHU DLU VDWXUDWLRQ $6 7KH ¿VK JUHZ ZHOO DQG HYHQ UHSURGXFHG &KDSWHUV DQG 2QH DGDSWLYH UHVSRQVH IRXQG LQ K\SR[LDUDLVHG +5 ¿VK ZDV D
3KHQRW\SLFUHVSRQVHVWROLIHORQJK\SR[LDLQFLFKOLGV &KDSWHU
IDYRXUDEOH WR LQFUHDVH WKH JLOO VXUIDFH DUHDZLWKRXWLQFUHDVLQJZDWHUUHVLVWDQFH RIWKHJLOOVZKLFKZRXOGUHVXOWLQKLJKHU FRVWRIYHQWLODWLRQ+XJKHVXVHGD PRGHOWKDWUHSUHVHQWHGWKHJLOO¿ODPHQWV DQG VHFRQGDU\ ODPHOODH DV D VHULHV RI UHFWDQJXODU FKDQQHOV DQG SUHGLFWHG HIIHFWV RI FKDQJHV LQ GLPHQVLRQDO SDUDPHWHUV RI WKH JLOOV RQ ZDWHU ÀRZ ,QFUHDVHV LQ JLOO VXUIDFH DUHD WKDW KDG WKH OHDVW HIIHFW RQ WKH WRWDO ÀRZ ZHUH DQ LQFUHDVH LQ WRWDO ¿ODPHQW QXPEHU DQ LQFUHDVH LQ ¿ODPHQW OHQJWK DQG DQ LQFUHDVH LQ KHLJKW RI WKH VHFRQGDU\ ODPHOODH ,Q FRQWUDVW DQ LQFUHDVH RI WKH UHVSLUDWRU\ VXUIDFH E\ KLJKHU IUHTXHQF\ RI WKH VHFRQGDU\ ODPHOODH RU ORQJHU VHFRQGDU\ ODPHOODH ZRXOG FDXVH UHODWLYHO\ ODUJH LQFUHDVHV LQ UHVLVWDQFH DQGZRXOGWKHUHIRUHEHXQIDYRXUDEOH
,W LV ZHOO NQRZQ WKDW WKHUH LV D YDVW LQWHUVSHFL¿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¿FDWLRQ RFFXUHG DQG
2FRQFHQWUDWLRQVRIPJOKDYHEHHQ
REVHUYHG LQ WKH KDELWDW RI + LULV 9DQ 2LMHQHWDO7KHWRWDOJLOODUHDRI WKLVVSHFLHVLVWLPHVODUJHUWKDQWKDW RI + KLDWXV PDLQO\ GXH WR ORQJHU DQG PRUHSULPDU\¿ODPHQWV+RRJHUKRXGHW DO
'UDPDWLFLQFUHDVHLQJLOOVXUIDFHRIDK\SR[LDUDLVHG/DNH9LFWRULDFLFKOLG
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¿VK ZHUH UDLVHG LQ OLWUH DTXDULD LQ WKH VDPH FOLPDWH URRP 7KXV JHQHWLF GLYHUVLW\ ZDV VPDOO DQG WUHDWPHQW RI WKH ¿VK SULRU WR WKH H[SHULPHQW ZDV WKH VDPH :KHQWKH¿VKZHUHPRQWKVROGWKH DLUVDWXUDWLRQ$6OHYHORIWKHZDWHURI WKH +5 JURXS ZDV ORZHUHG VWHSZLVH WR $6LQIRXUZHHNV7KHZDWHULQWKH DTXDULXP RI WKH QRUPR[LD UDLVHG JURXS ZDV NHSW XQGHU DLU VDWXUDWLRQ &LUFXODWLQJ ZDWHU ZDV PDGH K\SR[LF LQ DYDFXXPHTXLOLEUDWLRQFROXPQLQZKLFK GLVVROYHGDLULQWKHZDWHUHTXLOLEUDWHGDW N3D7KH/DTXDULDZHUHFRYHUHG ZLWKDPHWDOSODWHWKDWZDVSODFHGDIHZ FHQWLPHWUHV EHORZ WKH ZDWHU VXUIDFH WR UHVWULFW2H[FKDQJHZLWKWKHVXUIDFHDQG WRSUHYHQWDTXDWLFVXUIDFHUHVSLUDWLRQRI WKH ¿VK 7KH $6OHYHO ZDV FRQVWDQWO\ PRQLWRUHG $', ELRFRQWUROOHU $SSOLNRQ HTXLSSHG ZLWK SRODURJUDSKLF R[\JHQVHQVRUVDQGDGMXVWHGWR$6 YLD D VROHQRLG YDOYH LQ OLQH ZLWK DQ DLU VWRQH7KH¿VKZHUHNHSWDWDWHPSHUDWXUH RI&DQGDOLJKWGDUNUHJLPHRI KUV7KH\ZHUHJLYHQDGLYHUVHGLHWRI ÀDNH IRRG IUR]HQ PLGJH ODUYDH IUR]HQ ]RRSODQNWRQDQGDPL[WXUHRISXOYHUL]HG VKULPSVPXVVHOVDQGÀDNHIRRG%HIRUH DGXOWKRRGVXUYLYDODSSUR[LPDWHG LQERWKJURXSV
5HVSLURPHWU\([SHULPHQWV
3KHQRW\SLFUHVSRQVHVWROLIHORQJK\SR[LDLQFLFKOLGV &KDSWHU
)LJXUH 0HDVXUHV WKDW ZHUH WDNHQ ZLWK OLJKW PLFURVFRS\ DQG VFDQQLQJ HOHFWURQ
'UDPDWLFLQFUHDVHLQJLOOVXUIDFHRIDK\SR[LDUDLVHG/DNH9LFWRULDFLFKOLG
UHVSLURPHWU\SURFHGXUHXQGHUQRUPR[LD WKH\ GLHG ZKHQ WKH\ ZHUH VXEMHFWHG WR WKHHQWLUHSURWRFROLQWKHÀRZFKDPEHUV RI/XQGHU$6RUGXULQJUHFRYHU\ ,Q FRQWUDVW RWKHU KDSORFKURPLQHV RI VLPLODU VL]H VXUYLYHG $ SRVVLEOH H[SODQDWLRQ LV WKH DGGLWLRQDO VWUHVV GXH WR FRQ¿QHPHQW RI + S\UUKRFHSKDOXV ZKLFK LV QRUPDOO\ SHODJLF LQ D YHU\ VPDOOVSDFH:KHQXVLQJÀRZFKDPEHUV RI / + S\UUKRFHSKDOXV GLG VXUYLYH WKH ZKROH SURWRFRO XQGHU K\SR[LD 7R KDYH HQRXJK ELRPDVV ZH FKRVH WR GR WZRJURXSH[SHULPHQWVZLWKWKH/ÀRZ FKDPEHUVXVLQJHDFKWLPHWKHFRPSOHWH 15DQG+5JURXSUHVSHFWLYHO\DQG DQLPDOV 7KH 2 FRQVXPSWLRQ SHU ¿VK ZDV FRUUHFWHG IRU ZHLJKW GLIIHUHQFHV XVLQJ D PRGL¿FDWLRQ RI WKH IRUPXOD XVHG E\ 9DQ GHQ 7KLOODUW DQG .HVEHNH 0 0 : : « :Q
:KHUH P LV WKH PHWDEROLF UDWH RI D ¿VK RI JUDPV 0 LV WKH PHDVXUHG PHWDEROLF UDWH DQG : LV WKH ZHLJKW RI HDFK ¿VK LQ WKH H[SHULPHQW $IWHU UHVSLURPHWU\WKHDQLPDOVZHUHNLOOHGZLWK DQ RYHUGRVH RI DQDHVWKHWLF PJ /
WULFDLQHPHWKDQHVXOIRQDWH )LQTXHO DQG VWRUHG LQ IRUPDOGHK\GH EXIIHUHG ZLWK %RUD[ DW URRP WHPSHUDWXUH$IWHU PRQWKVWKHDQLPDOVZHUHWUDQVIHUUHG WRHWKDQROIRUIXUWKHUVWRUDJH
0HDVXUHPHQWV
2I VL[ 15 DQG VHYHQ +5 ¿VK WKH JLOO DUFKHV RQ WKH ULJKW VLGH RI WKH KHDG ZHUH UHPRYHG VHSDUDWHO\ XQGHU D GLVVHFWLRQ PLFURVFRSH DQG VWRUHG LQ DOFRKRO IRU IXUWKHU SURFHVVLQJ :LWK D GLJLWDO FDPHUD 1LNRQ &RROSL[ WKDW ZDV PRXQWHG RQ D GLVVHFWLRQ PLFURVFRSH SKRWRJUDSKV ZHUH PDGH RI WKH ODWHUDO KHPLEUDQFKV RI WKH IRXU JLOO DUFKHV WRJHWKHU ZLWK D SODVWLF EDU RI D NQRZQ VL]H IRU FDOLEUDWLRQ )URP WKHVH SKRWRJUDSKVWKHQXPEHURI¿ODPHQWVZDV FRXQWHGDQGWKHOHQJWKRI¿ODPHQWVZDV PHDVXUHGXVLQJWKHSURJUDP,PDJHWRRO 9 7H[DV 8QLYHUVLW\ +HDOWK 6FLHQFH &HQWUH 7R GHWHUPLQH WKH DYHUDJH ¿ODPHQWOHQJWKSHUKHPLEUDQFKDWOHDVW ¿ODPHQWV ZHUH PHDVXUHG RI HDFK KHPLEUDQFK 7KH ¿ODPHQWV WKDW ZHUH PHDVXUHG ZHUH GHWHUPLQHG E\ GLYLGLQJ WKH WRWDO QXPEHU SHU KHPLEUDQFK E\ DQGURXQGLQJLWRIIWRWKHQHDUHVWLQWHJHU EHORZ 7KXV UHQGHULQJ ¿ODPHQWV WREHPHDVXUHGWKDWZHUHHTXDOO\GLYLGHG RYHUWKHZKROHKHPLEUDQFK&RPSDULVRQ RIWKHDYHUDJH¿ODPHQWOHQJWKRIVHYHUDO KHPLEUDQFKV EDVHG RQ ¿ODPHQWV ZLWKDYHUDJHVRIWKHVDPHKHPLEUDQFKV EDVHG RQ PHDVXUHPHQW RI DOO ¿ODPHQWV VKRZHGGLIIHUHQFHVRI
3KHQRW\SLFUHVSRQVHVWROLIHORQJK\SR[LDLQFLFKOLGV &KDSWHU HTXDOQXPEHURI¿ODPHQWV$WWKHVDPH SRVLWLRQV¿ODPHQWVZHUHWKHQGLVVHFWHG DQG WKH GLPHQVLRQV RI WKH VHFRQGDU\ ODPHOODHDQGRIWKHFKDQQHOVLQEHWZHHQ ZHUHPHDVXUHG $V WKH PD[LPXP PDJQL¿FDWLRQ RI WKH GLVVHFWLRQ PLFURVFRSH ; GLG QRW DOORZ GHWDLOHG PHDVXUHPHQWV RI WKH GLPHQVLRQVRIWKHVHFRQGDU\ODPHOODHD 6FDQQLQJ (OHFWURQ 0LFURVFRSH 6(0 ZDV XVHG 7KHVH DOORZHG XV WR PDNH VKDUS LPDJHV RI D ¿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
7R PDNH WKH ¿ODPHQWV VWLIIHU WKH\ ZHUH WUHDWHG ZLWK D RVPLXP WHWUR[LGH VROXWLRQ LQ SKRVSKDWH EXIIHU DIWHU GLVVHFWLRQ $IWHUZDUGV WKH DOFRKRODQGZDWHULQWKH¿ODPHQWVZHUH UHSODFHGLQDVHULHVRILQFUHDVLQJDFHWRQH
FRQFHQWUDWLRQV OHDYLQJ WKHP IRU FULWLFDO SRLQW GU\LQJ 7KH DFHWRQH ZDV UHSODFHGE\&2%DOWHF&3'FULWLFDO SRLQWGU\HU6XEVHTXHQWO\WKH&2ZDV HYDSRUDWHG OHDYLQJ WKH GHK\GUDWHG ¿ODPHQWV7KH¿ODPHQWVZHUHWKHQJOXHG RQ VWXEV ZLWK VLOYHU JOXH DQG VSXWWHU FRDWHG ZLWK JROG 3RODURQ (TXLSPHQW /WG 6(0 FRDWLQJ XQLW ( $ VFDQQLQJ HOHFWURQ PLFURVFRSH -HRO -60ZDVXVHGIRUSKRWRJUDSKLQJ WKH¿ODPHQWV
'UDPDWLFLQFUHDVHLQJLOOVXUIDFHRIDK\SR[LDUDLVHG/DNH9LFWRULDFLFKOLG
ZHUH PHDVXUHG LQ DW OHDVW VHFRQGDU\ ODPHOODH RQ HDFK VLGH RI WKH ¿ODPHQW $YHUDJHVL]HVSHU¿ODPHQWHJSRVLWLRQ ZHUH XVHG IRU IXUWKHU FDOFXODWLRQV 7KH WRWDO UHVSLUDWRU\ VXUIDFH WRWDO VXUIDFH DUHD RI WKH VHFRQGDU\ ODPHOODH RI D FRPSOHWHKHPLEUDQFKZDVFDOFXODWHGDV 6XUIDFHDUHD )/$
:KHUH)LVWRWDO¿ODPHQWOHQJWKQXPEHU RI ¿ODPHQWV WLPHV DYHUDJH OHQJWK / LV ODPHOODU IUHTXHQF\ $ LV DUHD SHU VHFRQGDU\ ODPHOOD 7KH WRWDO ¿ODPHQW OHQJWK ZDV PXOWLSOLHG E\ WZR VLQFH ODPHOODHDUHIRXQGRQERWKVLGHVRIHDFK ¿ODPHQW7KHDUHDSHUVHFRQGDU\ODPHOOD ZDV PXOWLSOLHG E\ WZR VLQFH ERWK VLGHV RI HDFK ODPHOOD DUH XVHG IRU R[\JHQ H[FKDQJH
6WDWLVWLFV
$OOGDWDZHUHDQDO\VHGZLWKWKHVRIWZDUH SURJUDP 6366 9 6366 ,QF &KLFDJR,OIRU:LQGRZV)RU$QDO\VLV RI &RYDULDQFH WKH GDWD ZHUH OLQHDULVHG E\ OQWUDQVIRUPDWLRQ :LWK $1&29$ GLIIHUHQFHVEHWZHHQ15DQG+5¿VKZHUH LQYHVWLJDWHG ZKLOH XVLQJ WKH SRVLWLRQ DW WKHKHPLEUDQFKDWRGVHH)LJXUHDV D IDFWRU DQG ¿VK ZHLJKW DV D FRIDFWRU 3DUDPHWHU HVWLPDWHV ZHUH XVHG WR EDFN FDOFXODWHWKHVL]HVRIWKHSDUDPHWHUVWKDW ZHUH PHDVXUHG $OO SDUDPHWHUV ZHUH FRUUHFWHG DORQJ FRPPRQ UHJUHVVLRQ OLQHV WR WKDW RI D ¿VK RI WKH FRPPRQ PHDQZHLJKWRIDOOH[SHULPHQWDODQLPDOV XVHG JUDPV %RWK WKH UDZ GDWD DV
ZHOO DV WKH UHVLGXDOV IURP WKH YDULDQFH DQDO\VLVZHUHQRUPDOO\GLVWULEXWHG 5(68/76 5HVSLURPHWU\ 'XULQJUHVSLURPHWU\H[SHULPHQWVZLWKWKH 15JURXSRQHDQLPDOGLHGDIWHUKRXULQ WKHUHFRYHU\SHULRG7KHH[SHULPHQWZDV WHUPLQDWHGDWWKDWPRPHQW2IWKHGDWD PLQLPXP SHUFHQWLOH PHGLDQ SHUFHQWLOHDQGPD[LPXPSHUFHQWLOH 2 FRQVXPSWLRQ ZDV FDOFXODWHG DV GHVFULEHGLQ&KDSWHU
3KHQRW\SLFUHVSRQVHVWROLIHORQJK\SR[LDLQFLFKOLGV &KDSWHU $6 WKH 2 FRQVXPSWLRQ RI WKH 15 JURXS DW WKH DQG SHUFHQWLOH ZDV EHWZHHQDQGPJ2KJDW $6DQGEHWZHHQDQGPJ2 KJDW$6PHDQLQJDUHGXFWLRQ LQ2FRQVXPSWLRQE\DERXW )LODPHQWV 7KH ZHLJKW RI 15 ¿VK ZDV J J VWDQGDUG GHYLDWLRQ 7KDW RI +5 ¿VK ZDVJJVWDQGDUGGHYLDWLRQ7KH ZHLJKWRI15¿VKZDVVLJQL¿FDQWO\ODUJHU WKDQRI+5VLEOLQJVWWHVWS $OO SDUDPHWHUV ZHUH VL]H GHSHQGHQW 6LQFH
WKHOHQJWKDQGZHLJKWRIWKH¿VKKDUGO\ RYHUODSSHG EHWZHHQ JURXSV DQG VLQFH VL]HGHSHQGHQF\FUHDWHVDODUJHVWDQGDUG GHYLDWLRQLQWKHPHDQYDOXHV$1&29$ ZDVXVHGWRFRUUHFWPHDVXUHPHQWYDOXHV DORQJ D FRPPRQ PHDQ UHJUHVVLRQ OLQH WR D ¿VK RI D FRPPRQ PHDQ ZHLJKW JUDPV 7KH QRUPDOLVHG GDWD DUH SUHVHQWHGLQ7DEOH
7KH QXPEHU RI ¿ODPHQWV ZDV QRW VLJQL¿FDQWO\ GLIIHUHQW EHWZHHQ 15 DQG +5 ¿VK $1&29$ S 7KH ¿ODPHQW OHQJWK RI +5 DQLPDOV ZDV ODUJHU WKDQ WKDW RI 15 VLEOLQJV
'UDPDWLFLQFUHDVHLQJLOOVXUIDFHRIDK\SR[LDUDLVHG/DNH9LFWRULDFLFKOLG
$1&29$ S 7DEOH )LODPHQW OHQJWKYDULHGEHWZHHQWKHIRXUSRVLWLRQV )LJXUH DQG ZDV GHSHQGHQW RQ WKH ZHLJKWRIWKH¿VK7KH¿ODPHQWZLGWKRI +5DQLPDOVZDVRQDYHUDJHODUJHU WKDQRI15VLEOLQJV$1&29$S 7DEOH )URP FDXGDO SRVLWLRQ D WR URVWUDO SRVLWLRQ G RQ WKH KHPLEUDQFK WKH ¿ODPHQW ZLGWK LQFUHDVHG ZLWK UHVSHFWLYHO\DQG 6HFRQGDU\ODPHOODH
7KH VHFRQGDU\ ODPHOODH GHQVLW\ DV PHDVXUHG RYHU DOO IRXU ODWHUDO KHPLEUDQFKV ZDV QHJDWLYHO\ FRUUHODWHG ZLWK WKH ¿VK ZHLJKW 6SHDUPDQ FRUUHODWLRQFRHI¿FLHQW S
DVZHOODVZLWKWKHOHQJWKRIWKH¿ODPHQW 6SHDUPDQ FRUUHODWLRQ FRHI¿FLHQW S 7KHUH ZDV QR VLJQL¿FDQW HIIHFW RI ¿ODPHQW OHQJWK RQ DQ\RIWKHRWKHUPHDVXUHPHQWVWKDWZHUH WDNHQ7KHGHQVLW\RIVHFRQGDU\ODPHOODH RYHU DOO IRXU KHPLEUDQFKV ZDV ORZHULQ+5WKDQLQ15VLEOLQJV:KHQ PHDVXUHG RYHU WKH WKLUG JLOO DUFK RQO\ WKHGLIIHUHQFHZDV
3KHQRW\SLFUHVSRQVHVWROLIHORQJK\SR[LDLQFLFKOLGV &KDSWHU
7KHKHLJKWRIWKHVHFRQGDU\ODPHOODH ZDV RQ DYHUDJH ODUJHU LQ +5 ¿VK S 7KH LQFUHDVH IRXQG LQ +5 ¿VK YDULHG FRQVLGHUDEO\ RYHU WKH IRXU SRVLWLRQV HJ DQG
7KH GLVWDQFH EHWZHHQ VHFRQGDU\ ODPHOODH ZDV QRW VLJQL¿FDQWO\ GLIIHUHQW EHWZHHQ15DQG+5+S\UUKRFHSKDOXV S ,W ZDV QHJDWLYHO\ FRUUHODWHG ZLWKWKHZHLJKWDQGYDULHGRYHUWKHIRXU SRVLWLRQV7KHWKLFNQHVVRIWKHVHFRQGDU\ ODPHOODZDVRQDYHUDJHODUJHULQ +5 + S\UUKRFHSKDOXV S $Q $1&29$ RQ WKH PHDVXUHPHQWV DW DOO IRXUSRVLWLRQVSHUKHPLEUDQFKDQGXVLQJ ERWK ¿ODPHQW OHQJWK DQG ZHLJKW RI WKH ¿VK DV D FRYDULDEOH VKRZHG WKDW WKH GHQVLW\ RI WKH VHFRQGDU\ ODPHOODH ZDV VLJQL¿FDQWO\ORZHUE\DERXWRQHODPHOOD SHUPPS 7DEOH *LOOVXUIDFH :LWK $1&29$ ZH WHVWHG IRU D GLIIHUHQFHLQWRWDOJLOOVXUIDFHDUHDRIWKH WKLUGJLOODUFKEHWZHHQ15DQG+5¿VK XVLQJ ZHLJKW DV D FRYDULDEOH 7KH JLOO VXUIDFHDUHDRIJLOODUFKZDVGHSHQGHQW RQZHLJKWDQGVLJQL¿FDQWO\ODUJHULQ+5 WKDQLQ15¿VK$1&29$S DQG (VWLPDWHVZHUHPPIRU15 DQGPPIRU+5¿VKPHDQLQJWKDW WKHJLOODUHDRIJLOODUFKRI+5¿VKZDV ODUJHU WKDQ WKDW RI 15 VLEOLQJV $SDUWIURPWKHVL]HGLIIHUHQFHWKHJLOOV RI+5¿VKZHUHPXFKGDUNHUWKDQWKRVH RI 15 VLEOLQJV ,Q IUHVK QRQSUHVHUYHG DQLPDOVGLVVHFWHGJLOOVRI+5¿VKZHUH
FRORXUHGGHHSO\UHGZKLOHGLVVHFWHGJLOOV RI15VLEOLQJVZHUHYHU\SDOH
',6&866,21
([SHULPHQWDOGHVLJQ
'UDPDWLFLQFUHDVHLQJLOOVXUIDFHRIDK\SR[LDUDLVHG/DNH9LFWRULDFLFKOLG
UHSOLFDWLRQV ZKHQ FRQVLGHULQJ RQO\ WKH HIIHFWV RI OLIHORQJ K\SR[LD LUUHVSHFWLYH RI VSHFLHV GLIIHUHQFHV )RXUWK ZH IRUPXODWHG D SUHGLFWLRQ DERXW WKH GLUHFWLRQ RI WKH GLIIHUHQFHV EHWZHHQ 15DQG+5VLEOLQJVDQGWKHGLIIHUHQFHV IRXQGFRUUHVSRQGZLWKRXUH[SHFWDWLRQV
5HVSLURPHWU\
7KH REMHFWLYH RI SHUIRUPLQJ UHVSLURPHWU\ H[SHULPHQWV ZDV WR VKRZ WKDW +5 + S\UUKRFHSKDOXV ZHUH DEOH WR UHPDLQ DW WKH VDPH URXWLQH 2 FRQVXPSWLRQ OHYHO DW $6 DV WKH 15VLEOLQJVDW$67KH15JURXS WKDW ZDV H[SRVHG WR K\SR[LD VKRZHG D FRQVLGHUDEOH GHSUHVVLRQ RI PHWDEROLF UDWH DW $6 2QH DQLPDO HYHQ GLHG DIWHU K\SR[LD H[SRVXUH LQGLFDWLQJ WKDW 15 ¿VK DUH RQO\ WHPSRUDULO\ DEOH WR FRSH ZLWK VXFK FRQGLWLRQV ,Q FRQWUDVW WKH URXWLQH 2 FRQVXPSWLRQ RI +5 ¿VK DW$6ZDVHYHQKLJKHUWKDQWKDWRI 15 ¿VK DW $67KLV LQGLFDWHV WKDW SKHQRW\SLFUHVSRQVHVRFFXUUHGHQVXULQJ PDLQWHQDQFHRIURXWLQH2FRQVXPSWLRQ DW$66LPLODUSKHQRW\SLFUHVSRQVHV ZHUH VHHQ LQ WKUHH RWKHU VSHFLHV RI FLFKOLGV&KDSWHUVDQGVKRZLQJWKDW HDUO\H[SRVXUHWRK\SR[LDLQGXFHVVWURQJ DGDSWLYHUHVSRQVHVLQFLFKOLGV (IIHFWVRIFKDQJHGJLOOGLPHQVLRQV 7KH GLI¿FXOW\ LQ LQWHUSUHWLQJ WKH IXQFWLRQDOLW\RIWKHREVHUYHGGLIIHUHQFHV LQJLOOGLPHQVLRQVLVWKHODFNRIDJRRG WKHRUHWLFDO EDVLV ,Q WKH SDVW VHYHUDO DXWKRUV KDYH PDGH DWWHPSWV WR PRGHO
ÀRZ RI EORRG DQG ZDWHU WKURXJK JLOOV +XJKHV-RQHVHWDO6PLWK DQG-RKQVRQ0DWVXGDDQG6DNDL +RZHYHU VLQFH WKHUH DUH PDQ\ SDUDPHWHUV GHWHUPLQLQJ WKH G\QDPLFV LQ 2 GLIIXVLRQ DQG ÀRZ RI EORRG DQG ZDWHU WKURXJK WKH JLOOV HJ )LJXUH D FRPSUHKHQVLYH PRGHO LV VWLOO QRW DYDLODEOH%DVHGRQWZRPRGHOVZHWULHG WRDVVHVVZKDWWKHHIIHFWVRIWKHREVHUYHG DQDWRPLFDO FKDQJHV LQ JLOO GLPHQVLRQ ZHUH RQ JDV H[FKDQJH )RU HDFK LW LV DVVXPHG WKDW WKH RWKHU SDUDPHWHUV UHPDLQFRQVWDQWXQOHVVVWDWHGRWKHUZLVH $W $6 2 H[WUDFWLRQ LV SHUIRUPHG DW FRQGLWLRQV WKDW SRVH YHU\ GLIIHUHQW GHPDQGV RQ WKH JLOOV WKDQ DW QRUPR[LD $V +5 ¿VK DW $6 KDG WKHVDPHURXWLQH2FRQVXPSWLRQDV15 VLEOLQJVKDGDW$6WKH\ZRXOGKDYH QHHGHG DERXW DQ HLJKWIROG LQFUHDVH LQ ZDWHUYHQWLODWLRQWRVXSSO\WKHJLOOVZLWK HQRXJK2$QDVVXPSWLRQLVWKDWWKH2 H[WUDFWLRQ HI¿FLHQF\ ZDV MXVW DV KLJK 7KXV WKH PRUSKRORJ\ RI WKH JLOOV PXVW KDYH DOORZHG IRU D JUHDWHU ZDWHU ÀRZ ZKLFKLVGHSHQGHQWRQWKHUHVLVWDQFHRI WKH JLOOV /DPLQDU QRQWXUEXOHQW ÀRZ RIÀXLGVWKURXJKVPDOOFLUFXODUFKDQQHOV LV GHVFULEHG E\ WKH +DJHQ3RLVXLOOH HTXDWLRQ
T SSʌU
ȘO
:KHUHTÀRZLQPOSRUHVHFLV
3KHQRW\SLFUHVSRQVHVWROLIHORQJK\SR[LDLQFLFKOLGV &KDSWHU
S WKH GLDPHWHU RI WKH FDQDO U WKH YLVFRVLW\RIWKHÀXLGȘDQGWKHOHQJWK RI WKH FDQDO O 6XI¿FLHQW 2 H[FKDQJH LV GHSHQGHQW RQ RWKHU IDFWRUV WKDQ WKH PHUHDYDLODELOLW\RI2LQWKHJLOOVDORQH )LJXUH )LFN¶V ¿UVW ODZ RI GLIIXVLRQ GHVFULEHVWKHJDVH[FKDQJHSURFHVV
-QHW '$ǻ32ǻȤ
$FFRUGLQJ WR )LFN¶V ODZ WKH WRWDO 2 ÀX[IURPZDWHUWREORRG-QHWLVGLUHFWO\ SURSRUWLRQDOWRWKHVXUIDFHDUHD$WKH GLIIHUHQFHLQSDUWLDO2SUHVVXUHǻ32
)LJXUH 7KH PRVW LPSRUWDQW SDUDPHWHUV WKDW DIIHFW JDV H[FKDQJH UDWH LQ WKH JLOOV *DV
'UDPDWLFLQFUHDVHLQJLOOVXUIDFHRIDK\SR[LDUDLVHG/DNH9LFWRULDFLFKOLG
DQGWKHLQYHUVHRIWKHGLIIXVLRQGLVWDQFH ǻȤ
:KHQ DSSO\LQJ WKH WZR HTXDWLRQV DERYHWR¿VKJLOOVLWEHFRPHVFOHDUWKDW FHUWDLQ DOWHUDWLRQV LQ GLPHQVLRQV RI WKH JLOOVSOD\DFHQWUDOUROHLQLQFUHDVLQJERWK JDVH[FKDQJHDQGWKHZDWHUÀRZWKURXJK WKHJLOOV)LJXUH,QWKHVHFWLRQEHORZ ZH GLVFXVV WKH HIIHFWV RI WKH REVHUYHG GLIIHUHQFHV LQ JLOO GLPHQVLRQV RQ ZDWHU UHVLVWDQFHDQGJDVH[FKDQJH
*LOOUHVLVWDQFHDQGZDWHUÀRZ
:KHQ DSSO\LQJ WKH +DJHQ3RLVHXLOOH HTXDWLRQ WR ZDWHU ÀRZ WKURXJK JLOOV ÀRZUDWHFDQEHHQODUJHGE\LQFUHDVLQJ WKHSUHVVXUHGLIIHUHQFHRUGHFUHDVLQJWKH UHVLVWDQFH RI WKH UHVSLUDWRU\ FKDQQHOV 7KH ODWWHU FDQ EH UHDOLVHG E\ FKDQJLQJ WKHGLPHQVLRQVRIWKHJLOOVHJLQFUHDVLQJ WKH FURVVVHFWLRQDO DUHD GHFUHDVLQJ WKH OHQJWK DQG LQFUHDVLQJ WKH QXPEHU RI UHVSLUDWRU\FKDQQHOV
6LQFH VHFRQGDU\ ODPHOOD KHLJKW ZDV LQFUHDVHG E\ 7DEOH WKH FURVV VHFWLRQDODUHDRIHDFKUHVSLUDWRU\FKDQQHO ZDV ODUJHU ZKLFK UHGXFHG UHVLVWDQFH RI HDFK FKDQQHO 7KLV FKDQJH PD\ VHHP VPDOO EXW DFFRUGLQJ WR WKH +DJHQ 3RLVHXLOOH HTXDWLRQ WKH SRVLWLYH HIIHFW RQWKHÀRZLVH[SRQHQWLDOUHVXOWLQJLQD LQFUHDVHLQÀRZSHUSRUH+RZHYHU WKH +DJHQ3RLVHXLOOH HTXDWLRQ FDQ RQO\ EH XVHG VHPLTXDQWLWDWLYHO\ LQ WKLV FDVH EHFDXVH WKH UHVSLUDWRU\ FKDQQHOV DUH QRW WXEHV EXW UHVHPEOH D UHFWDQJXODU VKDSH $FFRUGLQJ WR 0RUWHQVHQ HW DO WKLV VKDSH GLIIHUHQFH KDV D
VOLJKW HIIHFW RQ UHVLVWDQFH GXH WR WKH ODUJHUFLUFXPIHUHQFHDQGDOWHUVWKHÀRZ UHJLPH WKURXJK D FKDQQHO $FFRUGLQJ WR%HQGLEDQG7DEHOLQJWKHÀRZ WKURXJK D WXEH ZLWK D FLUFXODU FURVV VHFWLRQ DQG D UHFWDQJXODU FURVVVHFWLRQ UHVHPEOH HDFK RWKHU FORVHO\ DV ORQJ DV WKH UDWLR EHWZHHQ KHLJKW DQG ZLGWK LV VPDOOHUWKDQ,QWKLVVWXG\WKHUDWLR ZDV DERXW $OWKRXJK WKH VHFRQGDU\ ODPHOOD WKLFNQHVV LQFUHDVHG E\ D VLJQL¿FDQW GHFUHDVH RI WKH UHVSLUDWRU\ FKDQQHO ZLGWK GLG QRW RFFXU VLQFH WKH IUHTXHQF\ RI VHFRQGDU\ ODPHOODH GHFUHDVHGDVZHOO7DEOH,QWKLVZD\ D GHFUHDVH RI WKH FURVVVHFWLRQDO DUHD RI WKH UHVSLUDWRU\ FKDQQHOV WKDW ZRXOG GUDVWLFDOO\ LQFUHDVH WKHLU UHVLVWDQFH ZDV DYRLGHG
7KH +5 ¿VK KDG ORQJHU VHFRQGDU\ ODPHOODH WKDQ 15 VLEOLQJV WKHUHE\ QRW RQO\ LQFUHDVLQJ WKH JLOO VL]H EXW DOVR LQFUHDVLQJ WKH UHVLVWDQFH DQG QHJDWLYHO\ DIIHFWLQJ WKH ZDWHU ÀRZ E\ WKH VDPH DPRXQW ,QWHUHVWLQJO\ WKLV PRUH RU OHVV FRPSHQVDWHV IRU WKH LQFUHDVHGÀRZUDWHWKDWZDVUHDOLVHGE\ WKH LQFUHDVHG KHLJKW RI WKH VHFRQGDU\ ODPHOODH)LJXUH
3KHQRW\SLFUHVSRQVHVWROLIHORQJK\SR[LDLQFLFKOLGV &KDSWHU
7KH WRWDO UHGXFWLRQ RI UHVLVWDQFH RI WKHJLOOVVHHPVWREHUHDOLVHGE\PDLQO\ WKH LQFUHDVHG QXPEHU RI UHVSLUDWRU\ FKDQQHOVDQGQRWVRPXFKE\WKHFKDQJHG VKDSH7KHFRQFRPLWDQWLQFUHDVHLQZDWHU ÀRZGRHVQRWQHDUO\UHVHPEOHWKHHLJKW IROG GLIIHUHQFH LQ ZDWHU ÀRZ QHFHVVDU\ WR VXSSO\ +5 ¿VK ZLWK HQRXJK 2 7KHUHIRUH LQ +5 ¿VK XQGRXEWHGO\ WKH SUHVVXUH GLIIHUHQFH RYHU WKH JLOOV PXVW KDYH LQFUHDVHG DQG ZDWHU ÀRZ VSHHG ZHUHFRQVLGHUDEO\ODUJHUWRDOORZIRUWKH UHTXLUHGLQFUHDVHLQWRWDOÀRZUDWH 2[\JHQGLIIXVLRQ
,Q+5¿VKDW$6WKH2ÀX[-QHW ZDVWKHVDPHDVLQ15¿VKDW$6 +RZHYHUǻ32 ZDV UHGXFHG HLJKWIROG LQ+5¿VK$ORJLFDOSUHGLFWLRQZRXOGEH WKDW+5¿VKPDLQWDLQHGWKHVDPH2ÀRZ WKURXJK DQ LQFUHDVH RI WKH VXUIDFH DUHD DQGDGHFUHDVHRIWKHGLIIXVLRQGLVWDQFH ,Q WKH +5 ¿VK IURP WKH SUHVHQW VWXG\ LQGHHGDQLQFUHDVHLQUHVSLUDWRU\VXUIDFH ZDVUHDOLVHGLQVHYHUDOZD\V)LUVW ORQJHU SULPDU\ ¿ODPHQWV UHVXOWHG LQ DQ LQFUHDVHLQWKHWRWDOQXPEHURIVHFRQGDU\ ODPHOODH6HFRQGWKHVHFRQGDU\ODPHOODH KDGDODUJHUVXUIDFHFDXVHGE\DQ LQFUHDVHLQERWKKHLJKWDVZHOODVOHQJWK 7KH WRWDO UHVSLUDWRU\ VXUIDFH RI WKH +5 + S\UUKRFHSKDOXV IURP WKLV VWXG\ ZDV WKXVHQODUJHGE\DVPXFKDV8QGHU QRUPDO FRQGLWLRQV PXFK RI WKH DUWHULDO EORRG LV VKXQWHG WKURXJK WKH PDUJLQDO YHVVHOV RI WKH JLOO ¿ODPHQWV DQG 15 ¿VK SHUIXVH RQO\ D IUDFWLRQ RI WKH WRWDO DPRXQW RI VHFRQGDU\ ODPHOODH GXULQJ
URXWLQH DFWLYLW\ 5DQGDOO 7KXV WKHGLIIHUHQFHLQWKHUHVSLUDWRU\VXUIDFH WKDWZDVDFWXDOO\XVHGEHWZHHQ15DQG +5+S\UUKRFHSKDOXVLVSUREDEO\ODUJHU WKDQ
'UDPDWLFLQFUHDVHLQJLOOVXUIDFHRIDK\SR[LDUDLVHG/DNH9LFWRULDFLFKOLG
(I¿FLHQF\DQGUHVSLUDWRU\FRVW
$OWKRXJK 2 H[FKDQJH LV LQFUHDVHG E\ DQHQODUJHGUHVSLUDWRU\VXUIDFHWKHZD\ WKLVLVUHDOLVHGKDVDODUJHHIIHFWRQJDV H[FKDQJH HI¿FLHQF\ DQG WKHUHIRUH WKH FRVW RI UHVSLUDWLRQ 6PLWK DQG -RKQVRQ VKRZHGWKDW2VDWXUDWLRQRIWKH EORRG GRHV QRW FKDQJH PXFK DW D ZLGH UDQJHRIZDWHUÀRZVSHHGVDQGVHFRQGDU\ ODPHOOD VKDSHV EXW WKH SHUFHQWDJH RI R[\JHQWKDWLVH[WUDFWHGIURPWKHZDWHU LV HDVLO\ UHGXFHG E\ D KLJK ZDWHU ÀRZ VSHHG)RUPRVW¿VKVSHFLHVKLJKZDWHU ÀRZ VSHHGV WKURXJK WKH UHVSLUDWRU\ FKDQQHOVWKDWUHVXOWLQORZ2H[WUDFWLRQ HI¿FLHQF\ PXVW EH DYRLGHG )RU EXFFDO YHQWLODWRUV ZKLFK QHHG WR JHQHUDWH D YHQWLODWRU\ZDWHUÀRZE\DFWLYHSXPSLQJ PRYHPHQW D UHODWLYHO\ KLJK SURSRUWLRQ RI WKH WRWDO HQHUJ\ EXGJHW LV VSHQW RQ YHQWLODWLRQZKLFKPDNHVKLJKHI¿FLHQF\ DQG ORZ JLOO UHVLVWDQFH QHFHVVDU\ 7KH FRVWRIYHQWLODWLRQUHODWLYHWRWKHWRWDO2 FRQVXPSWLRQ LV LQ 2UHRFKURPLV QLORWLFXV)HUQDQGHVDQG5DQWLQ LQWHQFK7LQFDWLQFD6FKXPDQQ DQG3LLSHUDQGLQWURXW 2QFRUK\QFKXV P\NLVV QRW VZLPPLQJ )HUQDQGH]DQG5DQWLQ,QFRQWUDVW WKHWXQD.DWVXZRPLVSHODPLVZKLFKLVD UDPYHQWLODWRUXVHVRQO\RILWVWRWDO HQHUJ\H[SHQGLWXUHIRUUHVSLUDWLRQ0XLU DQG+XJKHV 8QGHU WKH GHPDQGV WKDW FRVW RI UHVSLUDWLRQPXVWEHORZWKH2H[WUDFWLRQ HI¿FLHQF\ PXVW UHPDLQ KLJK 7KH H[WUDFWLRQ HI¿FLHQF\ LV GHSHQGHQW RQ WKHDPRXQWRIWLPHWKDWZDWHUUHPDLQVLQ FRQWDFWZLWKWKHJLOOV$VVKRZQLQ)LJXUH ZDWHUÀRZUDWHVPXVWKDYHLQFUHDVHG FRQVLGHUDEO\ LQ +5 ¿VK UHVXOWLQJ LQ D UHGXFHG H[WUDFWLRQ HI¿FLHQF\ FDXVHG E\ D VKRUWHU WLPH GXULQJ ZKLFK ZDWHU DQGJLOOVDUHLQFRQWDFWDQG2H[FKDQJH RFFXUV +RZHYHU VHFRQGDU\ ODPHOODH ZHUHORQJHUUHVXOWLQJLQDQHTXDO LQFUHDVHLQWKHFRQWDFWWLPH,QVXPPDU\ WKH ORQJHU VHFRQGDU\ ODPHOODH UHVXOW LQ ODUJHUUHVLVWDQFHWRZDWHUEXWDOVRLQDQ LPSURYHG2H[WUDFWLRQHI¿FLHQF\
3KHQRW\SLFUHVSRQVHVWROLIHORQJK\SR[LDLQFLFKOLGV &KDSWHU
D ODUJHU FDUGLDFRXWSXW 3LORW VWXGLHV RQ WLODSLDDQG+DSORFKURPLV$VWDWRWLODSLD SLFHDWXV VKRZHG WKDW WKLV LV LQGHHG WKH FDVHLQ+5¿VK
&RPSDULVRQZLWKRWKHUVSHFLHV
&KDSPDQ HW DO SHUIRUPHG D VSOLWEURRG H[SHULPHQW ZLWK WKH FLFKOLG 3VHXGRFUHQLODEUXV PXOWLFRORU YLFWRULDH WKDW ZHUH ODERUDWRU\UDLVHG DW QRUPR[LD PJ / RU K\SR[LD PJ / $GGLWLRQDOO\ WKH\ FRPSDUHG WZR SRSXODWLRQV FDXJKW LQ WKH ZLOG LQ D QRUPR[LF DQG K\SR[LF KDELWDW ,Q WKH ODERUDWRU\JURXSWKHUHVSLUDWRU\VXUIDFH RIWKH+5¿VKZDVFDOFXODWHGE\VHWWLQJ 15 JURXSV DW ODUJHU WKDQ WKDW RI 15 VLEOLQJV PDLQO\ FDXVHG E\ ORQJHU¿ODPHQWV7KHUHVSLUDWRU\VXUIDFH RI¿VKFDXJKWLQWKHK\SR[LFKDELWDWZDV ODUJHU WKDQ WKDW RI ¿VK FDXJKW LQ WKH QRUPR[LF KDELWDW7KLV ZDV UHDOLVHG WKURXJK ORQJHU ¿ODPHQWV DQG ODUJHU VHFRQGDU\ ODPHOODH &KDSPDQ HW DO DUJXHG WKDW WKH GLVFUHSDQF\ LQ WKH K\SR[LD UHODWHG GLIIHUHQFH EHWZHHQ ¿VKFDXJKWIURPWKHZLOGDQGWKHVSOLW EURRG H[SHULPHQW FRXOG EH DWWULEXWHG WR LQKHULWHG FKDQJHV LQ DGGLWLRQ WR SKHQRW\SLF SODVWLFLW\ LQ ¿VK IURP WKH ZLOGDOORZLQJIRUODUJHUGLIIHUHQFHVDQG GLIIHUHQWUHVSRQVHV ,Q FRPSDULVRQ LQ RXU VWXG\ WKH UHVSLUDWRU\VXUIDFHRIWKHWKLUGJLOODUFKRI +5DQLPDOVZDVODUJHUWKDQWKDWRI 15¿VK7KLVZDVUHDOLVHGWURXJKORQJHU ¿ODPHQWVDQGODUJHUVHFRQGDU\ODPHOODH ,UUHVSHFWLYHRIWKHGLIIHUHQFHVLQK\SR[LD H[SRVXUH SKHQRW\SLF UHVSRQVHV DORQH
'UDPDWLFLQFUHDVHLQJLOOVXUIDFHRIDK\SR[LDUDLVHG/DNH9LFWRULDFLFKOLG
GHHSHUOLYLQJ + LULV KDG D WLPHV JUHDWHU JLOO VXUIDFH +RRJHUKRXG HW DO 7KH GLIIHUHQFH LQ WKH GHQVLW\ RI VHFRQGDU\ODPHOODHIDOOVZLWKLQWKHUDQJH VHHQLQ15DQG+5+S\UUKRFHSKDOXV +RZHYHUWKHGLIIHUHQFHLQWKHQXPEHURI ¿ODPHQWVEHWZHHQ+LULVDQG+KLDWXV ZDV FRQVLGHUDEO\ ODUJHU WKDQ WKH UDQJH WKDWZDVVHHQLQ+S\UUKRFHSKDOXV
3ODVWLFLW\DQGLWVHFRORJLFDO FRQVHTXHQFHV