1958; Larsen, 1965; Barbour and Racine, 1967; Fox and. Thompson, 1971; Grime .... hydrated for 2 h on soaked paper in Petri dishes before their transfer to the moist ... radicle will appear is above the black band at the edge of the white area.
4'5'#4%*#46+%.'
#PKPUVTWOGPVCNUGVWRHQTUGGFIGTOKPCVKQPUVWFKGUYKVJ VGORGTCVWTG EQPVTQN CPF CWVQOCVKE KOCIG TGEQTFKPI 2GFTQ%06GKZGKTC ,QUÃ#%QGNJQ0GVQ*WODGTVQ4QEJCCPF,WTCE[/FG1NKXGKTC %GPVTQFG2GUSWKUCFQ4KQFG,CPGKTQ(WPFCÁºQ/QMKVK1MCFC4WC+VCDCKCPC)TCLCÕ4KQFG,CPGKTQ4,$TCUKN %QTTGURQPFKPICWVJQT HOQRSTKQ"EROQQTIDT
4GEGKXGF #RTKN 4GVWTPGFHQTTGXKUKQP ,WPG #EEGRVGF ,WN[
6JKURCRGTFGUETKDGUCPGYKPUVTWOGPVCNOGVJQFQNQI[RTQRQUGFHQTVJGUVWF[QHUGGFIGTOKPCVKQPYJKEJECPDGCRRNKGF VQ IGTOKPCVKQP RJ[UKQNQI[ CITKEWNVWTG CPF HQTGUV TGUVQTCVKQP 6JKU KPUVTWOGPV RGTHQTOU VJG TGEQTFKPI QH KOCIGU QH IGTOKPCVKPIUGGFUVJTQWIJVJGWUGQHCHNCVDGFUECPPGTEQWRNGFVQCEQORWVGTYJGTGCURGEKHKERTQITCOEQOOCPFUVJG ECRVWTGQHVJGUGKOCIGUCVRTGFGVGTOKPGFVKOGKPVGTXCNU6JGUGKOCIGUCTGCTEJKXGFHQTRQUVGTKQTCPCN[UKU6JGFGXKEG CNNQYUVJGEQNNGEVKQPQHFCVCCPFVJGGNCDQTCVKQPQHIGTOKPCVKQPEWTXGUYKVJJKIJCEEWTCE[(WTVJGTOQTGVJGFGXKEG RGTOKVUVJGUVWF[QHUGGFIGTOKPCVKQPQXGTCDTQCFTCPIGQHVGORGTCVWTGUCUYGNNCUVJGGCU[CRRNKECVKQPQHEJGOKECN CIGPVUVQVJGUGGFUKPQTFGTVQUVWF[VJGKTKPHNWGPEGQPVJGIGTOKPCVKQPRTQEGUU.CEVWECUCVKXCUGGFUYGTGWUGFKPVJKU UVWF[CUCPGZRGTKOGPVCNOQFGN)GTOKPCVKQPEWTXGUYGTGRNQVVGFCVFKHHGTGPVVGORGTCVWTGUKPQTFGTVQGXCNWCVGVJG FGXKEGHQTGZRGTKOGPVCNYQTM6JKURCRGTCNUQRTGUGPVUCOGVJQFQNQI[QHUVCVKUVKECNCPCN[UKUVJCVUWRRQTVUVJGTGNKCDKNKV[ QHVJKUGSWKROGPV -G[YQTFU FKIKVCNKOCIG.CEVWECUCVKXCUECPPGT5KGIGNŏUVGUVVJGTOCNITCFKGPVVCDNGU
&KURQUKVKXQRCTCGUVWFQUFGIGTOKPCÁºQEQOEQPVTQNGFGVGORGTCVWTCGTGIKUVTQCWVQO¶VKEQFGKOCIGO 'UVGCTVKIQ FGUETGXGWOCPQXCOGVQFQNQIKCKPUVTWOGPVCNRCTCQGUVWFQFGIGTOKPCÁºQFGUGOGPVGUCRNKE¶XGN´HKUKQNQIKCFCIGTOK PCÁºQCITKEWNVWTCGCTGUVCWTCÁºQFGHNQTGUVCU'UVGFKURQUKVKXQTGIKUVTCKOCIGPUFGUGOGPVGUGOIGTOKPCÁºQOGFKCPVG QWUQFGWOőUECPPGTŒCEQRNCFQCWOEQORWVCFQTQPFGWORTQITCOCGURGEÈHKEQEQOCPFCCECRVWTCFGUUCUKOCIGPU RCTC KPVGTXCNQU FG VGORQ RTÃFGVGTOKPCFQU 'UVCU KOCIGPU UºQ CTSWKXCFCU RCTC RQUVGTKQT CP¶NKUG 'UVG FKURQUKVKXQ RGTOKVGCEQNGÁºQFGFCFQURCTCCGNCDQTCÁºQFGEWTXCUFGIGTOKPCÁºQFGOQFQRCTVKEWNCTOGPVGCEWTCFQGOSWGC IGTOKPCÁºQFCUUGOGPVGURQFGUGTTGCNK\CFCRCTCWOCNCTICHCKZCFGVGORGTCVWTC CNÃOFKUUQCDTGUGCRQUUKDKNKFCFGFG CRNKECÁºQH¶EKNFGCIGPVGUSWÈOKEQUPCUUGOGPVGUPQUGPVKFQFGGUVWFCTCKPHNWÄPEKCFGUUGUCIGPVGUPQRTQEGUUQFG IGTOKPCÁºQ 5GOGPVGU FG .CEVWEC UCVKXC HQTCO WUCFCU EQOQ OQFGNQ GZRGTKOGPVCN (QTCO GUVCDGNGEKFCU EWTXCU FG IGTOKPCÁºQ RCTC FKHGTGPVGU VGORGTCVWTCU PQ UGPVKFQ FG CXCNKCT PQUUQ FKURQUKVKXQ GO VTCDCNJQ GHGVKXQ 'UVG CTVKIQ CRTGUGPVCVCODÃOWOCOGVQFQNQIKCFGCP¶NKUGGUVCVÈUVKECSWGICTCPVGCEQPHKCDKNKFCFGFQFKURQUKVKXQQTCFGUETKVQ 2CNCXTCUEJCXG KOCIGOFKIKVCN.CEVWECUCVKXCOGUCFGITCFKGPVGVÃTOKEQőUECPPGTŒVGUVGFG5KGIGN
+0641&7%6+10 6JG OGVJQFU EWTTGPVN[ WUGF KP VJG UVWF[ QH UGGF IGTOKPCVKQPWPFGTEQPVTQNNGFEQPFKVKQPUYKVJTGICTFVQ TGUGCTEJKPDQVCP[CPFRNCPVRJ[UKQNQI[CUYGNNCUVJG GXCNWCVKQP QH UGGF RQVGPVKCN HQT HQTGUV TGEQXGT[ CPFCITKEWNVWTGKPKVUDTQCFGTUGPUGCTGDCUGFQPFKTGEV
XKUWCNQDUGTXCVKQP6JKUHTGSWGPVN[TGSWKTGUVJGWUGQHC OCIPKH[KPI INCUU QT C UVGTGQOKETQUEQRG CPF VJG OCPKRWNCVKQP QH UGGFU VQ OGCUWTG KPFKXKFWCNN[ RCTCOGVGTU UWEJ CU UGGF CPF UGGFNKPI UK\G +P UVCPFCTF RTQEGFWTGUUGGFUCTGRNCEGFQPIGTOKPCVKQPVTC[UYJKEJ CTG WUWCNN[ MGRV KP ENKOCVK\GF ECDKPGVU CV RTGXKQWUN[
$TC\,2NCPV2J[UKQN
2%0 6'+:'+4# GV CN
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
$TC\,2NCPV2J[UKQN
VJGURGEKCNK\GFNKVGTCVWTG &GNNŏ#SWKNC CPFUQOG KPUVTWOGPVUWUGCUCPKOCIGECRVWTGTGEQTFGTCHNCVDGF UECPPGTQTCJKIJTGUQNWVKQPXKFGQECOGTCEQWRNGFVQC EQORWVGT6JGOCLQTKV[QHVJGUGKPUVTWOGPVUCTGFGXKUGF VQ WUG FKIKVK\GF KOCIGU HQT VJG CPCN[UKU QH EGTVCKP EJCTCEVGTKUVKEUUWEJCUTCFKENGRTQVTWUKQPUGGFNKPIUK\G UGGFUJCRGUGGFEQNQTCPFUGGFFGHGEVU5QOGQHVJGUG KPUVTWOGPVUWUGCHNCVDGFUECPPGTVQECRVWTGKOCIGUQH UGGFUUQYPQPUWDUVTCVGUNC[GTGFCVVJGDQVVQOQH2GVTK FKUJGU )GPGXG CPF -GUVGT QT C XKFGQ ECOGTC
&GNNŏ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
#0 +05647/'06#. 5'6 72 (14 5''& )'4/+0#6+10 567&+'5
CRRTCKUCN QH KVU RQVGPVKCN WUG KP VJG UVWF[ QH UGGF IGTOKPCVKQP .CEVWEC UCVKXC UGGFU YGTG UQYP KP VJG GZRGTKOGPVCNFGXKEGCPFUWDOKVVGFVQCDTQCFVGORGTCVWTG TCPIG CPF VJG IGTOKPCVKQP DGJCXKQT QH VJG YJQNG RQRWNCVKQP TGEQTFGF CNQPI VJG RTQEGUU CU FGUETKDGF CDQXG 9KVJ VJKU RTQEGFWTG VJG HQNNQYKPI KUUWGU YGTG EJGEMGF K GZRGTKOGPVCNTGRTQFWEKDKNKV[QHVJGU[UVGO UWRRQTVGF D[ C UVTKPIGPV EQPVTQN QH VGORGTCVWTG KK IGTOKPCVKQP JQOQIGPGKV[ QXGT VJG FKHHGTGPV TGIKQPU QP VJG IGTOKPCVKQP UWRRQTV UWTHCEG QT DGVYGGP FKHHGTGPV IGTOKPCVKQPDNQEMU CPF KKK ECRCEKV[QHVJKUU[UVGOVQ RTQFWEG C NCTIG XQNWOG QH IGTOKPCVKQP FCVC CV RTQITCOOGF VKOG KPVGTXCNU VJGTGD[ CNNQYKPI VJG EQPUVTWEVKQPQHIGTOKPCVKQPEWTXGUYKVJITGCVCEEWTCE[ CPFTGNKCDKNKV[
UWRRQTV6JKURCRGTYCURTKPVGFYKVJKPFGNKDNGKPMKPQTFGT VQ HQTO CP CTTC[ QH XGTVKECN VJKEM DNCEM DCTU ETQUUGF RGTRGPFKEWNCTN[D[VJKPNKPGU (KIWTG TGUWNVKPIKP UOCNNEGNNU'CEJEGNNJCUVJGTGHQTGVJGHQTOQHCYJKVG USWCTG YKVJ C DNCEM DCPF CV GCEJ UKFG# UKPING UGGF KU RQUKVKQPGFQPGCEJYJKVGCTGCUQVJCVVJGRQKPVYJGTGVJG TCFKENGYKNNCRRGCTKUCDQXGVJGDNCEMDCPFCVVJGGFIGQH VJG YJKVG CTGC 1PEG TCFKENG RTQVTWUKQP QEEWTU KVU RCNG [GNNQY KOCIG EQPVTCUVU UJCTRN[ CICKPUV VJG DNCEM DCPF 6JGUGGFUDGHQTGIGTOKPCVKQPQPVJGOQKUVRCRGTYGTG MGRV HQT J CV % KP CP CVVGORV VQ KORTQXG U[PEJTQPK\CVKQP VJTQWIJ C EQNF UVTCVKHKECVKQP RTQEGUU
9KNNKCOCPF8KXKCP ;COCIWEJKGVCN +P V J K U RCRGT VJG GXGPV EQPUKFGTGF CU KPFKECVKXG QH IGTOKPCVKQP KU TCFKENG RTQVTWUKQP CPF GNQPICVKQP VJG DQVCPKECNETKVGTKQP
/#6'4+#.#0&/'6*1&5
'SWKROGPV FGUETKRVKQP 6JG U[UVGO KU GUUGPVKCNN[ EQORQUGFQHHQWTOQFWNGU UGG(KIWTG UVOQFWNGŌ IGTOKPCVKQP EJCODGT # PF OQFWNG Ō IGTOKPCVKQP DNQEMU VJGTOQUVCVKECNN[EQPVTQNNGF YKVJCOGVCNNKERNCVG WUGF CU VJG UWRRQTV DGF HQT VJG OQKUV RCRGT YJKEJ TGEGKXGUVJGUGGFU $ TFOQFWNGŌVTCPURCTGPVYKPFQY YKVJ VJGTOCN KPUWNCVKQP RTQRGTVKGU VJTQWIJ YJKEJ VJG UGGFIGTOKPCVKQPKOCIGU TCFKENGRTQVTWUKQPU CTGRKEMGF WR YJKNG VJG UGGFU CTG OCKPVCKPGF KP C EQPFKVKQP QH EQPUVCPV VGORGTCVWTG CPF JWOKFKV[ % VJ OQFWNG Ō C
5GGFU .CEVWECUCVKXCEX)TCPF4CRKFUUGGFU URGEKCNN[ HWTPKUJGF HTGG QH HWPIKEKFG YGTG WUGF RWTEJCUGF HTQO (GNVTKP 5VCVG QH 4KQ )TCPFG FQ 5WN $TC\KN
YYYUGOGPVGUHGNVTKPEQODT 5GGF)GTOKPCVKQP 5GGFUUGNGEVGFYKVJYGKIJVUDGVYGGP CPF OI YKVJ RTGEKUKQP QH v OI YGTG RTG J[FTCVGFHQTJQPUQCMGFRCRGTKP2GVTKFKUJGUDGHQTG VJGKTVTCPUHGTVQVJGOQKUVRCRGTWUGFQPVJGIGTOKPCVKQP
(KIWTG6QRXKGYQHVJGIGTOKPCVKQPEJCODGT1DUGTXGVJGTGIWNCTOCVTKZFKUVTKDWVKQPQHUGGFUQPVJGCTTC[QHYJKVG EGNNUCPFVJGRTQVTWFKPITCFKENGUQXGTVJGCFLCEGPVDNCEMDCPFU
$TC\,2NCPV2J[UKQN
2%0 6'+:'+4# GV CN
D
C1
C
C3 B3 B1 B2
B4
B
A1 A
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
# 6JGIGTOKPCVKQPUWRRQTVU DNQEMU HWPEVKQPCUJGCV EQPFWEVQTUJQOQIGPK\KPIVJGJGCVFKUVTKDWVKQPCNQPIVJG YJQNG UWTHCEG YJGTG VJG UGGFU CTG RNCEGF 6JG VGORGTCVWTG EQPVTQN KP VJGUG CNWOKPWO DNQEMU $ KU QDVCKPGF CU C EQPUGSWGPEG QH VJG VJGTOCN GSWKNKDTKWO CVVCKPGF VJTQWIJ VJG KPVGTCEVKQP QH C EQNF CPF C JQV
$TC\,2NCPV2J[UKQN
UQWTEG 6JG EQNF UQWTEG KU VJG YCVGT EQOKPI HTQO C VJGTOQUVCVKE DCVJ EKTEWNCVKPI KPVQ VJG IGTOKPCVKQP DQZ VJTQWIJUVCKPNGUUUVGGNVWDGU # 6JKUDQZKUHKNNGFYKVJ YCVGTKPQTFGTVQGUVCDNKUJVJGTOCNEQPVCEVDGVYGGPVJG DNQEMU CPF VJG EQNF UQWTEG 6JG JQV UQWTEG $ KU CP GNGEVTQPKECNN[ EQPVTQNNGF JGCVGT TGUKUVQT 6JG HKPCN VGORGTCVWTG CFLWUVOGPV KU QDVCKPGF VJTQWIJ VJG RQYGT EQPVTQN QH VJG JGCVKPI TGUKUVCPEG KP HWPEVKQP QH C FKHHGTGPEG KP VGORGTCVWTG DGVYGGP VJG CNWOKPWO DNQEM CPF VJG VGORGTCVWTG CFLWUVGF D[ VJG QRGTCVQT 6JKU FKHHGTGPEG YCU OQPKVQTGF D[ C %1'. VGORGTCVWTG EQPVTQNNGTOQFGNYKVJ2+&NQQREQPVTQNTGEGKXKPIKVU CPCNQIKEXQNVCIGUKIPCNKPRWVHTQOCV[RG,VJGTOQEQWRNG
$ CPFUGPFKPICPCPCNQIKEEWTTGPVUKIPCNQWVRWV HTQO VQO# VQC.QVKRJCUGCPINGV[RGEQPVTQNNGTOQFGN $%YJKEJEQPVTQNUVJGFGNKXGT[QHRQYGT 6JGUGEQPFOQFWNGKUEQORTKUGFGUUGPVKCNN[QHCUGV QH IGTOKPCVKQP DNQEMU CPF CP GSWCN PWODGT QH IGTOKPCVKQPUWRRQTVDGFU6JGIGTOKPCVKQPUWRRQTVDGFU
$ CTGOCFGQHVJKPUVCKPNGUUUVGGNUJGGVUVJCVHKVQXGT VJGDNQEMUCPFCTGJGNFHKTON[D[UETGYUVQVJGDQF[QH VJG DNQEMU 6JG UWRRQTV DGFU JCXG VYQ TGUGTXQKTU CNQPIUKFGEQPVCKPKPIYCVGTQTCP[FGUKTGFUQNWVKQPHQT VJGOCKPVGPCPEGQHVJGJWOKFKV[NGXGNKPVJGOQKUVRCRGT QTGXGPVWCNN[VQTGEGKXGUQOGEJGOKECNFKUUQNXGFKPVJG IGTOKPCVKQPYCVGT 6JG VJKTF OQFWNG KU EQPUVTWEVGF YKVJ C FQWDNG YCNN VTCPURCTGPVINCUUYKPFQYVQHCEKNKVCVGKOCIGECRVWTGHTQO YKVJKPVJGIGTOKPCVKQPEJCODGTDGUKFGUHWPEVKQPKPICUC JGCV KPUWNCVQT CPF CXQKFKPI YCVGT EQPFGPUCVKQP 6JKU OQFWNGKUEQPUVTWEVGFD[VJGKPUGTVKQPQHCEQTMICUMGV
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
#0 +05647/'06#. 5'6 72 (14 5''& )'4/+0#6+10 567&+'5
FTQRNGVU KP VJG YKPFQYU VJCV ECP QEEWT CV JKIJGT VGORGTCVWTGU 6JG HQWTVJ OQFWNG EQPUKUVU QH C HNCVDGF UECPPGT
)GPKWU OQFGN %QNQT 2CIG'2 Ō QRVKECN TGUQNWVKQP FRK 6JG UECPPGT KU RQUKVKQPGF FKTGEVN[ QXGT VJG VJKTF OQFWNGUQVJCVKVUUECPPKPIYKPFQYHCEGNKGURCTCNNGNVQ VJG UGGF UWRRQTVU 6JKU KU QPN[ RQUUKDNG KH VJG UECPPGT WPKVKURNCEGFKPCPWRUKFGFQYPRQUKVKQP6QFQUQVJG UWRRQTVKPI TQFU YGTG CFCRVGF VQ HKV VJG UECPPGT WPKV JQTK\QPVCNN[QXGTVJGIGTOKPCVKQPEJCODGT 6JGEQORWVGTEQWRNGFVQVJGHQWTVJOQFWNGYCUWUGF VQECRVWTGCPFUVQTGVJGKOCIGUTGEGKXGFCVTGIWNCTVKOG KPVGTXCNU HTQO VJG HNCVDGF UECPPGT +OCIG ECRVWTG CWVQOCVKQPKURGTHQTOGFD[VJGVCUMUEJGFWNGUQHVYCTG PCOGN[5EJGFWNGT2TQHGUUKQPCNHQT9KPFQYU8GTUKQP
YYYURNKPVGTYCTGEQO QT CP[ QVJGT RTQITCO URGEKHKECNN[FGXGNQRGFHQTVJKURWTRQUG +OCIG TGEQTF 6JG ECRVWTGF KOCIGU YGTG RTQEGUUGF D[ VJGRTQITCO2CIG+OCIG 2TQNCD6GEJ%Q.VF CPFVJG TGEQTFGF KOCIGU YGTG CTEJKXGF YKVJ VJG6CIIGF +OCIG (KNG(QTOCV VKH CVCTGUQNWVKQPQHFRK DNCEMCPFYJKVG KOCIGU %QORWVGT EQPHKIWTCVKQP 2GPVKWO ++ +PVGN /*\ 4#//$EQPVCKPKPIC)$JCTFFKUM 5VCVKUVKEU CPF ITCRJKECN CPCN[UKU 6JG UQHVYCTG $+1'56#6 #[TGUGVCN YCUWUGFHQTUVCVKUVKECN CPCN[UKU6JG'ZEGNUQHVYCTGYCUCRRNKGFVQQRVKOK\GVJG IGTOKPCVGFUGGFEQWPVKPIRTQEGFWTG)TCRJKECNRNQVVKPI CPFCPCN[UKUYGTGECTTKGFQWVYKVJVJGCKFQH1TKIKP
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
WUGFVQKFGPVKH[VJGEJTQPQNQIKECNQTFGTQHIGTOKPCVKQP GXGPVU QDUGTXGF 6JGUG PWODGTU CTG KPVGIGT PWODGTU UVCTVKPIHTQOYJKEJEQTTGURQPFUVQCPGXGPVVJCVYCU QDUGTXGFKPVJGKOCIGTGIKUVGTGFCVVJGVKOGYJGPVJGHKTUV UECPPKPIYCURGTHQTOGF6JGKPFGZPWODGTKPETGCUGUKP WPKVCT[UVGRUEQTTGURQPFKPIVQVJGUGSWGPEGPWODGTQH GCEJ UECP 6JWU KPFGZ PWODGT TGRTGUGPVU C IGTOK PCVKQPGXGPVQDUGTXGFKPVJGKOCIGTGIKUVGTGFCVVJGVKOG QHVJGUGEQPFKOCIGEQNNGEVKQPCPFUQQP(QTGZCORNGKH C IGTOKPCVGF UGGF CRRGCTU QP VJG OQPKVQT UETGGP CV C FGVGTOKPGF RQUKVKQP KP VJG KOCIG EQNNGEVGF CV VJG HKTUV QDUGTXCVKQP VKOG VJG PWODGT KU TGEQTFGF KP VJG EGNN EQTTGURQPFKPIVQVJGRQUKVKQPQHVJKUUGGFKPVJGRTKPVGF URTGCFUJGGV%QORNGVKPIVJKUEQWPVKPIHQTCNNVJGVKOGU
RKEVWTGU VJG FCVC HTQO VJG TGUWNVKPI URTGCFUJGGV CTG TGRTQFWEGF QP CP 'ZEGN URTGCFUJGGV MGGRKPI VJG UCOG NC[QWV EQTTGURQPFGPEG DGVYGGP URTGCFUJGGV EGNNU CPF RQUKVKQPUQPVJGIGTOKPCVKQPRNCVGCUUJQYPKP(KIWTG 9KVJKPVJGUEQRGQHUVCVKUVKECNCPCN[UKUQHVJGTGUWNVU VJGTG YCU C PGGF VQ GXCNWCVG CXGTCIG CPF FKURGTUKQP XCNWGU HQT IGTOKPCVKQP EQWPVKPI YKVJ VKOG CNQPI VJG IGTOKPCVKQP EWTXG (QT VJKU VJG TGUWNVKPI URTGCFUJGGV YCU UWDFKXKFGF KPVQ UKZ TGIKQPU GCEJ TGIKQP YKVJ UGGFU VJGUJCRGQHVJGTGIKQPUKUTGRTGUGPVGFKP(KIWTG NCDGN# (QTGCEJIGTOKPCVKQPVKOGVJGECNEWNCVKQPQHVJG CTKVJOGVKE OGCP CPF VJG 5& YGTG RGTHQTOGF CPF VJGUG XCNWGU WUGF VQ TGRTGUGPV VJG CXGTCIG CPF FKURGTUKQP GUVKOCVGUTGURGEVKXGN[ (QT VJG GNCDQTCVKQP QH IGTOKPCVKQP EWTXGU VJG EQWPVKPIEQTTGURQPFKPIVQCURGEKHKEECRVWTGFKOCIGYCU NQECVGF YKVJKP GCEJ QH VJG UKZ TGIKQPU QP VJG DNQEM 6CMKPIKPCEEQWPVVJGEQWPVKPIFCVCQHCNNDNQEMUHQTCNN KPFGZGUIGTOKPCVKQPEQWPVKPIYCURNQVVGFCICKPUVVKOG VQQDVCKPVJGIGTOKPCVKQPEWTXG 6QXGTKH[RQUUKDNGFKHHGTGPEGUDGVYGGPVJGVYQDNQEMU CPF COQPI VJG FKHHGTGPV TGIKQPU QH GCEJ DNQEM C UVCVKUVKECNVTGCVOGPVYCUCRRNKGFKPQTFGTVQCUUWTGVJCV VJG HCEVQTU YJKEJ CTG TGURQPUKDNG HQT IGTOKPCVKQP QT JCXGCP[GHHGEVQPKV CTGRGTHGEVN[FKUVTKDWVGFQXGTVJG YJQNGUGGFRQRWNCVKQP+PQVJGTYQTFUVJGU[UVGOECPDG EJGEMGFVJKUYC[HQTVJGETGCVKQPQHTGIKQPCNCTVKHCEVUKP VJGIGTOKPCVKQPFCVC 6QXGTKH[VJGFCVCJQOQIGPGKV[COQPIVJGFKHHGTGPV TGIKQPUQHGCEJDNQEMVJGFCVCDNQEMUTGRTQFWEGFQPVJG 'ZEGNURTGCFUJGGVYGTGUWDFKXKFGFKPVQTGIKQPUYKVJ GKIJV EGNNU RGT TGIKQP C V[RKECN NC[QWV QH VJG TGIKQP KU
$TC\,2NCPV2J[UKQN
2%0 6'+:'+4# GV CN
$TC\,2NCPV2J[UKQN
Row
Column
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
EQPUKFGTKPIUGSWGPEGUCNQPITQYUQTCNQPIEQNWOPUCU UVCVGF DGHQTG CPF VJQUG KPFGZGU DGNQPIKPI VQ VJG VKOG TCPM6QVJGUGUGVUQHPWODGTU5KGIGNŏUTCPFQOPGUUVGUV YCU CRRNKGF KP QTFGT VQ GXCNWCVG JQY VJG OGCP IGTOKPCVKQPVKOGXCNWGUCTGFKUVTKDWVGFQXGTVJGDNQEMŏU UWTHCEGYJGVJGTKPCTCPFQOQTKPCFGVGTOKPKUVKEHCUJKQP 5KGIGNŏUTCPFQOPGUUVGUVOC[DGDGUVCRRNKGFVQCDKPCT[ TCPMUGSWGPEG HQTGZCORNGYKVJŏUCPFŏU 6JGTGUWNV QHVJGVGUVEQTTGURQPFUVQCPGXCNWCVKQPRCTCOGVGTHQTVJG TCPFQOFKUVTKDWVKQPQHŏUCPFŏUKPVJGUGSWGPEGYJGTG VJGTG YGTG 0 XCNWGU CPF 0 XCNWGU KP VJG DKPCT[ UGSWGPEG 6JG RTCEVKECN CRRNKECVKQP QH VJKU VGUV VQ VJG FCVCRTQFWEGFKPQWTGZRGTKOGPVUYCURQUUKDNGVJTQWIJ VJGWUGQHVJGHQNNQYKPIRTQEGFWTG CVVTKDWVKPIVJGXCNWG VQVKOGTCPMXCNWGUGSWCNQTUOCNNGTVJCPCPFXCNWGVQ VKOGTCPMXCNWGUJKIJGTVJCPKVYCURQUUKDNGVQETGCVGC UGSWGPEGQHDKPCT[KPFGZGUEQTTGNCVGFYKVJVJGTGIKQPU
UGGőENCUUKHKECVKQPŒKP6CDNG 6JGRTQEGFWTGFGUETKDGF CDQXGIGPGTCVGFCUGSWGPEGQHTCPMKPFGZGUCPF TCPMKPFGZGU 0 CPF0 5GCTEJKPIYKVJKPVJKU UGSWGPEGHQTUWDITQWRUEQPUVKVWVGFD[TGRGVKVKQPQHQPG QHVJGTCPMKPFGZGU QT KVYCURQUUKDNGVQEQORWVGVJG
B
1
A
(KIWTG4GRTGUGPVCVKQPQHVJG'ZEGNURTGCFUJGGVYJGTG VJGIGTOKPCVKQPGXGPVUCTGPWOGTKECNN[RTGUGPVGF#CPF $TGRTGUGPVUVJGTGIKQPETGCVGFVQRGTHQTOVJGUVCVKUVKECN CPCN[UKU UGG/CVGTKCNCPF/GVJQFUKVGO 6JGPWODGT KP VJKU URTGCFUJGGV YJKEJ KU KPEQORNGVG CPF QPN[ RTGUGPVGFJGTGCUCPGZCORNG KPFKECVGUVJGRNCEGYJGTG VJGHKTUVGXGPVYCUTGIKUVGTGF VJGHKTUVIGTOKPCVKQP 6CDNG 5KGIGNŏU VGUV FCVC CU CRRNKGF VQ IGTOKPCVKQP GXGPVU UGG/CVGTKCNCPF/GVJQFUKVGO )GTOKPCVKQP$NQEM 4QYQTFGTGFTGIKQPU%QNWOPQTFGTGFTGIKQPU )GTOKPCVKQP)GTOKPCVKQP 4GIKQP6KOG%NCUUKHKECVKQP4GIKQP6KOG%NCUUKHKECVKQP TCPMTCPM
#PCN[UKU2CTCOGVGTU#PCN[UKU2CTCOGVGTU 004004
#0 +05647/'06#. 5'6 72 (14 5''& )'4/+0#6+10 567&+'5
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
A
Cumulative germination (seed number)
50 40
3.4ºC 5.9ºC 8.4ºC 10.8ºC 13.3ºC 15.8ºC 18.3ºC
30 20 10 0
(seed number)
0
2000
4000
6000
8000
10000 12000
B
45 40 35 30 25 20 15 10 5 0
19.9ºC 21.7ºC 25.8ºC 28.3ºC 29.9ºC 32.5ºC 34.1ºC 36.6ºC 0
500 1000 1500 2000 2500 3000 3500 4000 Time (min)
(KIWTG )GTOKPCVKQP EWTXGU QH .CEVWEC UCVKXC UGGFU YJGTGGCEJRQKPVTGRTGUGPVUVJGOGCPXCNWGQHIGTOKPCVKQP FCVC HQT # CPF $ DNQEMU 'TTQTU HQT GCEJ RQKPV CTG TGRTGUGPVGF D[ VJG UOCNN XGTVKECN DCTU +P # UGGF IGTOKPCVKQPCV%%%%%% CPF % +P $ UGGF IGTOKPCVKQP CV % % %%%%%CPF%
KPFKECVGVJGUKOKNCTKV[QHIGTOKPCVKQPDGJCXKQTQPVJGUG VYQ DNQEMU UQ VJCV VJGUG FCVC UGVU CTG RTCEVKECNN[ KPFKUVKPIWKUJCDNG 6JKU KU VTWG PQV QPN[ HQT GCEJ VGORGTCVWTGDWVCNUQHQTGCEJVKOGCUKPFKECVGFD[VJG GZEGNNGPVCITGGOGPVDGVYGGPRCKTGFFCVCEQTTGURQPFKPI VQDQVJDNQEMU+PQVJGTYQTFUVJGIGTOKPCVKQPDGJCXKQT KURTCEVKECNN[VJGUCOGHQTVJGUGVYQDNQEMU %QPUKFGTKPIVJGGXCNWCVKQPQH5KGIGNŏUVGUVVJGXCNWG 4YCUEQORWVGFCUUVCVGFDGHQTGHQTGCEJVGORGTCVWTG HQT GXGT[ TQY CPF EQNWOP HQT VJG VYQ DNQEMU 6JG 4 XCNWGUKPFKECVGVJGFGITGGQHTCPFQOPGUU#EEQTFKPIVQ 5KGIGNŏUVGUVHQTCFKUVTKDWVKQPYJGTG0 CPF0 VJG GZRGEVGF 4 XCNWGU HQT C TCPFQO UGSWGPEG OWUV DG DGVYGGPCPF+P(KIWTGCRNQVKUUJQYPQHVJG4 XCNWGUEQORWVGFYKVJVJGFCVCHTQOCNNVJGGZRGTKOGPVU CXCKNCDNGRGTHQTOGFQXGTCDTQCFTCPIGQHVGORGTCVWTGU CPFEQPUKFGTKPIRQUKVKQPCNUGSWGPEKPIQHXCNWGUDQVJKP VJG TQY UGPUG CPF EQNWOP UGPUG 6JG FCUJGF NKPGU TGRTGUGPV VJG XCNWGU QH GZRGEVCVKQP NKOKVU OGPVKQPGF CDQXG +V KU GXKFGPV VJCV VJG GZRGTKOGPVCN 4 XCNWGU QDVCKPGF YGTG YGNN ITQWRGF CTQWPF VJG OQUV RTQDCDNG XCNWG QH EQORTKUGF OQUVN[ YKVJKP VJG GZRGEVCVKQP NKOKVUYKVJVJGGZEGRVKQPQHVYQRQKPVU 6JKU CUUGTVKQP YCU CNUQ VGUVGF WUKPI CPQVJGT UVCVKUVKECNVGUV%QPUKFGTKPIVJGHQTOGTRTQRQUGFFKXKUKQP QH VJG GZRGTKOGPVCN FCVC KP UKZ TGIKQPU VJG FCVC UGV EQTTGURQPFKPIVQVJGUGTGIKQPUQPVJG'ZEGNURTGCFUJGGV HQT DQVJ IGTOKPCVKQP DNQEMU YGTG UWDOKVVGF VQ C PQPRCTCOGVTKEXCTKCPEGCPCN[UKUVJG-TWUMCN9CNNKU Cumulative germination (seed number)
4XCNWGYJKEJOWUVTGRTGUGPVVJGVQVCNPWODGTQHVJGUG UWDITQWRU9KVJVJKUXCNWGVJGCRRNKECVKQPQHVJG5KGIGN ŏU VGUVYCURQUUKDNG CUSWQVGFD[5CPVCPCCPF4CPCN
Block A 3.4ºC Block B 19.9ºC 50
Block A Block B 36.6ºC
Block A Block B
40 30 20 10 0 0
2000
4000
6000
8000
10000 12000
Time (min)
(KIWTG)GTOKPCVKQPEWTXGUTGNCVKXGVQDNQEM#CPFDNQEM $HQTIGTOKPCVKQPVGORGTCVWTGUQH%%CPF%
$TC\,2NCPV2J[UKQN
Value R
2%0 6'+:'+4# GV CN
Temperature (ºC)
(KIWTG 'ZRGTKOGPVCN XCNWGU QH 4 5KGIGNŏU VGUV HQT IGTOKPCVKQPQPDNQEM#CPFDNQEM$CPCN[\GFKPTQYUQT EQNWOPU HQT FKHHGTGPV VGORGTCVWTGU 6JG FCUJGF NKPGU FGVGTOKPGVJGWRRGTCPFVJGNQYGTNKOKVUQHTGNKCDKNKV[HQT4 XCNWGU VGUV #[TGUGVCN YJGTGGXGT[TGIKQPQPGCEJDNQEM YCU OCVEJGF YKVJ VJG P TGOCKPKPI TGIKQPU 6JKU VGUV YCUCRRNKGFVQVJGFCVCEQTTGURQPFKPIVQ%CPF%
FGRKEVGFKP(KIWTG &WGVQVJGUOCNNPWODGTQHUGGFU IGTOKPCVKPICV%VJKUVGUVEQWNFPQVDGCRRNKGFKPC TGNKCDNG YC[ VQ VJG FCVC EQORWVGF CV VJKU VGORGTCVWTG 6JKU VGUV RTQFWEGF HQT VJG FCVC QDVCKPGF CV % CPF %-TWUMCN9CNNKURTQDCDKNKV[XCNWGUQHCPF TGURGEVKXGN[ KPFKECVKPI C JKIJN[ JQOQIGPGQWU DGJCXKQTQHVJGYJQNGUGVQHIGTOKPCVKQPFCVCHQTCNNVJG TGIKQPUQPGCEJDNQEMCPFVJGUCOGDGJCXKQTQEEWTTKPI DGVYGGP VJG VYQ DNQEMU 6JGUG TGUWNVU QH 5KGIGNŏU VGUV CPFVJG-TWUMCN9CNNKUVGUVEKVGFCDQXGCTGKPXGT[IQQF CITGGOGPV YKVJ VJG GZRGEVGF TCPFQO PCVWTG QH IGTOKPCVKQPFCVCFKUVTKDWVKQP
&+5%755+10 6JGKPUVTWOGPVCNU[UVGORTGUGPVGFJGTGHQTVJGUVWF[ QHUGGFIGTOKPCVKQPTGRTGUGPVUCPQXGNFGXKEGCPFCSWKVG GHHKEKGPV VQQN HQT VJKU RWTRQUG 6JG FGUETKRVKQP QH VJKU FGXKEGCPFKVUCUUQEKCVGFOGVJQFQNQI[ENGCTN[KPFKECVGU KVU XGTUCVKNKV[ CPF KVU UGXGTCN CFXCPVCIGU KP EQORCTKUQP YKVJVJGEWTTGPVOGVJQFUQHUGGFIGTOKPCVKQPCPCN[UKUCU RQKPVGF QWV KP VJG +PVTQFWEVKQP 6JGUG CFXCPVCIGU CTG DCUGFOCKPN[QPKVUJKIJNGXGNQHCWVQOCVKQPYJGTGVJG KOCIG ECRVWTG D[ VJG KOCIG UGPUQT KU RGTHQTOGF KP C
$TC\,2NCPV2J[UKQN
RTGEKUGN[EQPVTQNNGFOCPPGTCVUEJGFWNGFVKOGKPVGTXCNU #U C EQPUGSWGPEG VJG RTQEGFWTG KPEQTRQTCVGU C UGV QH RTCEVKECN CFXCPVCIGU K VJG TGEQTFKPI CPF UVQTCIG QH IGTOKPCVKQPFCVCKUUWEJVJCVVJGTGVTQCEVKXGCPCN[UKUQH VJG FCVC KU RQUUKDNG VJKU DGKPI RGTHQTOGF D[ FKTGEV XKUWCNK\CVKQPQPVJGOQPKVQTUETGGPQTKPCFKIKVK\GFHQTO D[CEQORWVGTRTQITCO CPKORQUUKDNGVCUMYJGPWUKPI EQPXGPVKQPCNOGVJQFUDCUGFQPXKUWCNQDUGTXCVKQP KK VJGRQUUKDKNKV[QHKOCIGECRVWTGCVCP[RTGXKQWUN[EJQUGP KPUVCPVCPFCVENQUGTVKOGKPVGTXCNUVJWUTGUWNVKPIKPVJG OKPKOK\CVKQPQHICRUDGVYGGPGZRGTKOGPVCNRQKPVUKPVJG IGTOKPCVKQP EWTXGU CU YGNN CU UJCTRGT CPF JKIJN[ TGRTQFWEKDNG EWTXG RTQHKNGU KP EQPVTCUV VQ VJG EWTXGU EQPUVTWEVGF D[ EQPXGPVKQPCN OGVJQFU CPF KKK VJG RQUUKDKNKV[QHGNCDQTCVKPICRTQITCOVQFGVGTOKPGVJGVKOG KPVGTXCNDGVYGGPEQPUGEWVKXGKOCIGECRVWTGUVQQRVKOK\G VJG WUG QH VJG UECPPGT VJGTGD[ CXQKFKPI WPPGEGUUCT[ KOCIGECRVWTGUYKVJVJGHWTVJGTRQUUKDKNKV[QHEQPVTQNNKPI VJGQDUGTXCVKQPYKPFQYYKFVJ#FFKVKQPCNN[FWTKPIVJG RGTKQFYJGPVJGHCUVGUVIGTOKPCVKQPTCVGKUGZRGEVGFVJG VKOGDGVYGGPKOCIGECRVWTGUOC[DGTGFWEGFVQCPGZVGPV VJCV KU RTCEVKECNN[ KORQUUKDNG YKVJ FKTGEV XKUWCN QDUGTXCVKQP (WTVJGTOQTG VJG JKIJGT HKFGNKV[ KP VJG CPCN[UKUQHVJGUGGFRQRWNCVKQPCUCYJQNGTGUWNVUKPVJG EQPUVTWEVKQPQHEWTXGUTKEJGTKPKPHQTOCVKQPCPFITGCVGT KP CEEWTCE[ $GUKFGU VJGUG CPCN[VKE CFXCPVCIGU VJGTG KU CPQRGTCVKQPCNCFXCPVCIGYQTVJ[QHPQVG VJGRQUUKDKNKV[ QH SWKEM CPF GCU[ YCVGT UWRRN[ TGRNGPKUJOGPV YKVJQWV ITGCVFKUVWTDCPEGUQHVJGGPXKTQPOGPVQHVJGGZRGTKOGPVU 6JKURTQEGFWTGCNUQRGTOKVUVJGCFOKPKUVTCVKQPQHUGXGTCN EJGOKECNUVQVJGUGGFUKPENWFKPIRJ[VQJQTOQPGUYJGP VJGTGCEVKQPUGNKEKVGFD[VJGUGCIGPVUECPDGQDUGTXGFKP TGCNVKOG 9KVJ TGICTF VQ VGORGTCVWTG EQPVTQN VJG GSWKROGPV FGUETKDGFJGTGECTTKGUQWVVJKUVCUMKPFKXKFWCNN[KPGCEJ IGTOKPCVKQPDNQEMVJTQWIJVJGWUGQHUGRCTCVGGNGEVTQPKE VJGTOQUVCVU 6JKU MKPF QH VGORGTCVWTG EQPVTQN CNNQYU ITGCV XGTUCVKNKV[ KP VJG CUUKIPOGPV QH RTGFGVGTOKPGF VGORGTCVWTGU VQ GCEJ DNQEM 6JKU CNUQ RGTOKVU VJG KPETGCUGQTFGETGCUGQHVJGVGORGTCVWTGQHCNNDNQEMUQTC RCTVKEWNCTDNQEMFWTKPICPGZRGTKOGPV6JGUGHGCVWTGUCTG RCTVKEWNCTN[CFXCPVCIGQWUHQTTGUGCTEJKPVJGTOQDKQNQI[ 9JGPWUKPIUECPPGTUEQTTGURQPFKPIVQRCRGTUK\G# KPVJGCRRCTCVWUJGTGFGUETKDGFVYQIGTOKPCVKQPDNQEMU RGTUECPPGTOC[DGWUGF9QTMKPIYKVJHQTGZCORNG DNQEMU CV VJG UCOG VKOG YKVJ UGVU QH FKHHGTGPV
#0 +05647/'06#. 5'6 72 (14 5''& )'4/+0#6+10 567&+'5
EQPFKVKQPUCVNGCUVUECPPGTUOWUVDGWUGF6QQDXKCVG VJKU SWGUVKQP URGEKCN UECPPGTU EQWNF DG FGXGNQRGF HQT VJKU RWTRQUG ECRCDNG QH UECPPKPI ITGCVGT NGPIVJU CPF EQPUGSWGPVN[ VJG NGPIVJ QH VJG IGTOKPCVKQP EJCODGT EQWNFDGGZVGPFGFCEEQTFKPIN[VQJQNFCITGCVGTPWODGT QHIGTOKPCVKQPDNQEMU.CTIGTUECPPGTUWUKPIVJGUGVWR RTQRQUGF JGTG EQWNF CEEQOOQFCVG UGGFU QH OGFKWO QT ITGCVGTUK\G # OKPQT NKOKVCVKQP QH VJKU GSWKROGPV VJCV YKNN DG QXGTEQOG KP C OQFGN PQY WPFGT EQPUVTWEVKQP KU VJG FKHHKEWNV[KPXKUWCNK\CVKQPQHVJGUGGFRQRWNCVKQPDGECWUGQH YCVGTEQPFGPUCVKQPCVVGORGTCVWTGUCDQXG% 5QOG HGCVWTGU QH VJKU CRRCTCVWU VJCV FGUGTXG URGEKCN GORJCUKU CTG KVU TGNKCDKNKV[ CPF VJG EQPUKUVGPE[ QH VJG TGUWNVU 6JKU YCU FGOQPUVTCVGF VJTQWIJ VJG ECTGHWN CRRNKECVKQP QH UVCVKUVKECN OGVJQFU QH CPCN[UKU VQ VJG EQORWVGF FCVC CUUWTKPI VJG CDUGPEG QH CTVKHCEVU YJKEJ EQWNFKPFWEGGTTQTUKPFCVCKPVGTRTGVCVKQPU (WTVJGT CURGEVU QH VJKU CRRCTCVWU CPF KVU CUUQEKCVGF OGVJQFQNQI[ CTG YQTVJ[ QH EQPUKFGTCVKQP 4CFKENG RTQVTWUKQPCUQDUGTXGFJGTGECPDGCPCFXCPVCIGQWUVQQN HQT VJG GXCNWCVKQP QH UGGF XKIQT VCMKPI KPVQ CEEQWPV VJG DQVCPKECNETKVGTKQPHQTVJKUKORQTVCPVRCTCOGVGTCPFVJKUECP DGEQORNGOGPVGFYKVJVJGOGCUWTGOGPVQHTCFKENGITQYVJ TCVG 5WEJ EQORWVGTCKFGF KOCIG GXCNWCVKQPU JCXG DGGP RTQRQUGFCPFRGTHQTOGFD[UGXGTCNCWVJQTU *QHHOCUVGTGV CN 1CMNG[GVCN 6GKZGKTCGVCN &WVVCPF )GPGXG YJQRQKPVQWVVJGCDUGPEGQHUWDLGEVKXKV[CPF UJQTVGT YQTM VKOG YJGP EQORCTGF YKVJ VJG UGXGTCN EQPXGPVKQPCN OGVJQFU QH XKUWCN QDUGTXCVKQP %QORWVGT CKFGFKOCIGCUUGUUOGPVECPDGECTTKGFQWVKPCPGCUKGTHCUV CPFTGNKCDNGYC[D[VJGKPUVTWOGPVCPFOGVJQFQNQI[JGTG RTGUGPVGF #EMPQYNGFIOGPVU 6JG CWVJQTU YKUJ VQ CEMPQYNGFIG (GNVTKP+ORQTVCFQTCFG5GOGPVGU(CTTQWRKNJC4KQ)TCPFG FQ5WNHQTVJGMKPFIKHVQHVJG.CEVWECUCVKXCUGGFUUGNGEVGF CPFHTGGQHHWPIKEKFGU
4'('4'0%'5 #[TGU/#[TGU/,#[TGU&.5CPVQU#5 $KQ'UVCV #RNKECÁÐGU'UVCVÈUVKECUPCUTGCUFCU%KÄPEKCU$K QNÎIKECUG/ÃFKECU VJGF5QEKGFCFG%KXKN/COWKT¶ /%6+ORTGPUC1HKEKCNFQ'UVCFQFQ2CT¶$GNÃO $CTDQWT /# 4CEKPG %* %QPUVTWEVKQP CPF RGTHQTOCPEG QH C VGORGTCVWTGITCFKGPV DCT CPF
EJCODGT'EQNQI[ &GNNŏ#SWKNC#XCP'EM,9XCPFGT*GKLFGP)9#0 6JG CRRNKECVKQP QH KOCIG CPCN[UKU KP OQPKVQTKPI VJG KODKDKVKQP RTQEGUU QH YJKVG ECDDCIG $TCUUKEC QNGTCEGC. UGGFU5GGF5EK4GU &GNNŏ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
#P CWVQOCVGF U[UVGO HQT XKIQT VGUVKPI VJTGG FC[QNFUQ[DGCPUGGFNKPIU5GGF5EK6GEJPQN .CDQWTKCW .) # VJGTOCN ITCFKGPV DNQEM HQT IGTOKPCVKQPGZRGTKOGPVU4GX$TCU$KQN .CDQWTKCW.)%CXCNECPVK4$ 7ODNQEQFGITCFKGP VG VÃTOKEQ FG H¶EKN EQPUVTWÁºQ RCTC GZRGTKOGPVQU VGTOQDKQNÎIKEQU4GX$TCU(KUKQN8GI .CTUGP #. 7UG QH VJGTOQITCFKGPV RNCVG HQT UVWF[KPI VGORGTCVWTG GHHGEVU QP UGGF IGTOKPCVKQP 2TQE+PV5GGF6GUV#UUQE /E.CWIJNKP 0$ $QYGU )4 6JQOCU #) &[EM ($ .KPFUC[6/9KUG4( #PGYFGUKIPHQTCUGGF IGTOKPCVQT YKVJ KPFGRGPFGPVN[ VGORGTCVWTG EQPVTQNNGFEGNNU9GGF4GU 1CMNG[--GUVGT56)GPGXG4. %QORWVGTCKFGF FKIKVCNKOCIGCPCN[UKUQHUGGFNKPIUK\GCPFITQYVJTCVG HQT CUUGUUKPI UGGF XKIQT KP +ORCVKGPU 5GGF 5EK 6GEJPQN
$TC\,2NCPV2J[UKQN
2%0 6'+:'+4# GV CN
5CPVCPC)&4CPCN/# #P¶NKUGFC)GTOKPCÁºQ7O 'PHQSWG 'UVCVÈUVKEQ 'F (WPFCÁºQ 7PKXGTUKFCFG FG $TCUÈNKC$TCUÈNKC 6GKZGKTC'(%ÈEGTQ5/0GVQ&& #P¶NKUGFGKOCIGPU FKIKVCKUFGRN¸PVWNCURCTCCXCNKCÁºQFQXKIQTFGUGOGPVGU FGOKNJQ4GX$TCU5GO 9KNNKCO,88KXKCP-6 5VWFKGUQHVJGOGEJCPKUOQH
$TC\,2NCPV2J[UKQN
GPJCPEGOGPV QH RJ[VQEJTQOG FGRGPFGPV NGVVWEG UGGF IGTOKPCVKQPD[RTGEJKNNKPI2NCPV2J[UKQN ;COCIWEJK;1ICYC/-WYCJCTC#*COCFC#-CO[KC; ;COCIWEJK 5 #EVKXCVKQP QH IKDDGTGNNKP DKQU[PVJGUKUCPFTGURQPUGRCVJYC[UD[NQYVGORGTCVWTG FWTKPI KPJKDKVKQP QH #TCDFQRUKU VJCNKCPC UGGFU 2NCPV %GNN
