Cover Page

(1)

The handle

http://hdl.handle.net/1887/133432

holds various files of this Leiden University

dissertation.

Author: Akram, M.N.

(2)

543955-L-bw-Akram 543955-L-bw-Akram 543955-L-bw-Akram 543955-L-bw-Akram Processed on: 11-6-2020 Processed on: 11-6-2020 Processed on: 11-6-2020

Processed on: 11-6-2020 PDF page: 7PDF page: 7PDF page: 7PDF page: 7

(3)

ϴ

,QWURGXFWLRQ

3ODQWVDUHRILPPHQVHLPSRUWDQFHIRUKHDOWKFDUHWKURXJKRXWWKHZRUOG7KH\ DUHQRWRQO\ LPSRUWDQWLQWUDGLWLRQDOPHGLFLQHEXWDOVR PRUHWKDQ RIPRGHUQPHGLFLQHVRULJLQDWH IURPSODQWV 7KHUHDUHRYHUVSHFLHVRIDQJLRVSHUPVLQWKHZRUOG$PRQJWKHVH VSHFLHV$PDU\OOLGDFHDH LVDPRQRFRW\OHGRQIDPLO\IURPDQJLRVSHUPVZKLFKFRQVLVWVRI JHQHUDDQGDERXWVSHFLHVNarcissus LLVDJHQXVRIWKLVIDPLO\ZKLFKFRQVLVWVRI  VSHFLHV > @ 7KLV Narcissus / JHQXV RI SHUHQQLDO JHRSK\WHV LV QDWLYH WR WKH 0HGLWHUUDQHDQUHJLRQ>@,W LVDYHU\SRSXODUJDUGHQSODQWLQ(XURSH(FRQRPLFDOO\LWLV DOVRDQLPSRUWDQWSODQWZLWKODUJHQXPEHURIVSHFLHVK\EULGV DQGYDULHWLHVLQFXOWLYDWLRQ 8QLWHG .LQJGRP 8. DQG 7KH 1HWKHUODQGV DUH WRS FXOWLYDWRUV DQG SURGXFHUV RI NarcissusUHVSHFWLYHO\+LVWRULFDOO\ Narcissus EXOEVKDYHEHHQXQGHUFXOWLYDWLRQ LQ7KH 1HWKHUODQGVVLQFHWKHWK FHQWXU\ZKLOH Narcissus hispanicus KDVEHHQFXOWLYDWHGLQWKH 8.IRUPRUHWKDQ \HDUV7KRXJK WKHLPSRUWDQFH RINarcissus DVDIORULFXOWXUDOFURS KDVHVFDODWHG VLQFHWKHODWHWKFHQWXU\>@$FRQVXPHU¶VVXUYH\LQUHSRUWHG WKDWGDIIRGLOVDFKLHYHGWKHVHFRQGSRVLWLRQEDVHGRQ FRQVXPHUSUHIHUHQFHV DPRQJVWFXW IORZHUV LQ WKH 8. 7KLV SRVLWLRQ LV TXLWH KLJK FRQVLGHULQJ WKH IDFW WKDW WKH\ DUH QRW DYDLODEOHWKURXJKRXWWKH\HDUDQGEHLQJUHODWLYHO\LQH[SHQVLYHLQWKHIORZHUPDUNHW>@ ,Q WHUPV RI SURGXFWLRQ 7KH 1HWKHUODQGV LV DW WKH WRS SRVLWLRQ LQ WKH OLVW RI Narcissus SURGXFHUV(YHU\\HDUDURXQGKHFWDUHVRINarcissus DUH SODQWHGLQ7KH1HWKHUODQGV $FFRUGLQJWRDQHFRQRPLFUHSRUWRI7KH1HWKHUODQGVHDUQDOPRVWPLOOLRQ(XURV LQWRWDOVDOHVIURPIORZHULQJEXOEVZKLFKDOVRLQFOXGHVNarcissus>@

(4)

ϵ

Narcissus VWDUWWREORRPZLWKLQWRZHHNV>@$OONarcissus KDYHVRPHDPRXQWRI

IUDJUDQFH WKRXJK WKHjonquil DQGtazetta W\SHVRINarcissus KDYHWKHVWURQJHVW IUDJUDQFH 2UQDPHQWDOYDOXHRIWKHVHSODQWVLV ZHOONQRZQ RYHUWKHWLPH0RUHRYHUWKH\KDYHDOVR VKRZQKLJK SKDUPDFRORJLFDOLPSRUWDQFHDORQJZLWKRUQDPHQWDO7KLVSKDUPDFRORJLFDO LPSRUWDQFHLVGXHWRWKHSUHVHQFHRIFRPSRXQGVZKLFKDUHSURGXFHGE\WKHSODQWVIRU WKHLURZQGHIHQVHDJDLQVWGLIIHUHQWH[WHUQDOVWUHVVIDFWRUV7KHVHSODQWVSHFLHVVSHFLILF FRPSRXQGVDUHFDOOHGVHFRQGDU\PHWDEROLWHV7KHVHPHWDEROLWHV DUHSURGXFHGORFDOO\LQ D VPDOO TXDQWLW\ E\ WKH SODQWV 6RPH RI WKHVH PHWDEROLWHV DUH SURGXFHG FRQVWLWXWLYHO\ ZKLOH RWKHUDUHRQO\SURGXFHGDIWHUVRPHW\SH RIVWUHVVHJ HQYLURQPHQWDO KHUELYRU\RU LQWHUDFWLRQZLWKPLFURRUJDQLVPV Narcissus SODQWV DUH XVHGLQKHUEDOPHGLFLQHV GXHWRWKH SUHVHQFHRIWKHVHPHWDEROLWHV 'ZHFN>@ KDVUHYLHZHGWKHXVHRINarcissus LQKHUEDO PHGLFLQHVDQGIRRG+HKDVDOVRPHQWLRQHGVRPHSRLVRQRXVHIIHFWV RINarcissus SODQWV LQJHQHUDORQKXPDQKHDOWK6LQFHHDUOLHU WLPHV KHUEDOPHGLFLQH Narcissus SODQWVDUHXVHG DVKHUEDOPHGLFLQHWRWUHDWGLIIHUHQWGLVHDVHVVXFKDVFDQFHUEURQFKLWLVZKRRSLQJFRXJK DQG EDOGQHVV Narcissus ZDV DOVR XVHG DV D FRQWUDFHSWLYH PHGLFLQH LQ HDUOLHU WLPHV $FFRUGLQJWR WKHOLWHUDWXUH LWLVFOHDUWKDWNarcissus FRQWDLQVPDQ\ELRDFWLYHFRPSRXQGV ZKLFK DUH EHQHILFLDO IRU KXPDQ KHDOWK 7KHVH FRPSRXQGV LQFOXGH D ODUJH QXPEHU RI DONDORLGVIODYRQRLGVFDURWHQRLGVOHFWLQV DQGYDULRXVYRODWLOHV

(5)

ϭϬ

F

(6)

ϭϭ

F

Figure 1.2. Chemical structures of different Narcissus alkaloids [9].

)URP DOO WKHVH ELRORJLFDOO\ DFWLYH DONDORLGV JDODQWKDPLQH LV RQH RI WKH PRVW VWXGLHG FRPSRXQGV 7KLVLVGXHWRWKHUHDVRQWKDWLWLVDQLQKLELWRURIDFHW\OFKROLQHVWHUDVH HQ]\PH DQGFDQDIIHFWQHXURFHOOGHJHQHUDWLRQ 7KDW¶VZK\LW FDQEHREWDLQHGIURPWKHPDUNHWLQ WKHIRUP RI EURPLGHVDOW DVDPHGLFLQHWRWUHDW$O]KHLPHU¶VGLVHDVH

(7)

ϭϮ

&DURWHQRLGV LV DQRWKHU JURXS RI FRPSRXQGV ZKLFK DUH PDLQO\ SURGXFHG E\ VSHFLILF RUJDQV 7KHVH RUJDQV DUH PRVWO\ LQYROYHG LQ SKRWRV\QWKHVLV 7KHVH SLJPHQWV RFFXU QDWXUDOO\DQG PDMRUO\FRQWULEXWH IRUFRORULQIORZHUOHDYHV IUXLWV DQGYHJHWDEOHV7KHVH FRPSRXQGVDOVRSHUIRUP WZRYHU\LPSRUWDQWIXQFWLRQVGXULQJWKHSKRWRV\QWKHWLFSURFHVV LQ SODQWV )LUVWO\ WKH\ DUH XVHG IRU OLJKW KDUYHVWLQJ DQG VHFRQGO\ WKH\ KHOS LQ WKH SURWHFWLRQRI WKHSKRWRV\QWKHWLFSDUWVRIWKHSODQWIURPSKRWRR[LGDWLRQ>@0RUHRYHU WKH\FDQ DOVR IXQFWLRQDV WKHSUHFXUVRUVRIVLJQDOLQJPROHFXOHV 7KXV QRWRQO\ WKH\FDQ LQIOXHQFH WKHGHYHORSPHQWRISODQWVEXWDOVRLQIOXHQFHWKHUHVSRQVHVWRELRWLFDQGDELRWLF VWUHVV %\ SHUIRUPLQJ DOO WKHVH IXQFWLRQV FDURWHQRLGV IDFLOLWDWH SKRWRPRUSKRJHQHVLV QRQSKRWRFKHPLFDOTXHQFKLQJOLSLGSHUR[LGDWLRQDQG KHOSLQDWWUDFWLQJSROOLQDWRUV> @

/HFWLQV FDQ EH GHILQHG DV FDUERK\GUDWHELQGLQJ SURWHLQV ZKLFK UHFRJQL]H DQG ELQG VSHFLILFDOO\ DQG UHYHUVLEO\ WR FHUWDLQ PRQR RU ROLJRVDFFKDULGHV 7KH FKDUDFWHULVWLF RI WKHVHOHFWLQELQGLQJLVWKDWWKH\ELQG ZLWKRXWDOWHULQJWKHVWUXFWXUHRIWKHERQGLQJOLJDQG >@7KHUHDUHVHYHQOHFWLQIDPLOLHVDQGNarcissus OHFWLQVDUHW\SLFDOUHSUHVHQWDWLYHVRI PRQRFRWPDQQRVHELQGLQJOHFWLQVNarcissus OHFWLQVZHUHILUVWGHVFULEHGLQE\9DQ 'DPPH7KH\KDYHGLIIHUHQWELRORJLFDODFWLYLWLHVZKLFKLQFOXGHDJJOXWLQDWLRQLQUDEELW DQWKUDFLWHVSUHYHQWLRQDJDLQVWWKHUDELHVYLUXVDQGDIIHFWWKHUXEHOODYLUXVPXOWLSOLFDWLRQ 6RPH RI WKH Narcissus OHFWLQV KDYH VKRZQ DFWLYLW\ DJDLQVW KXPDQ LPPXQRGHILFLHQF\ YLUXV>@ LQin-vitro VWXGLHV>@'LIIHUHQWWR[LFDFWLYLWLHVRIWKHVHOHFWLQVKDYHDOVR EHHQWHVWHG

*HQHVDQGHQ]\PHVRINarcissus DUHDOVRIRXQGWREHRIJUHDWLPSRUWDQFHQRWRQO\IRU NarcissusLWVHOIEXWDOVRIRUWKHRWKHUFURSVZKHQWKH\ZHUHRYHUH[SUHVVHGLQWKRVHFURSV $JRRGH[DPSOHRIDNarcissus JHQHRYHUH[SUHVVLRQLVWKHJROGHQULFHZKHUHDJHQHRI Narcissus FDURWHQRLG SDWKZD\ ZDV RYHUH[SUHVVHG WRJHWKHU ZLWK DQRWKHU JHQH IURP EDFWHULDWRRYHUFRPHWKHGHILFLHQF\RIYLWDPLQ$

(8)

ϭϯ

T

Table 1.1. List of genes isolated from Narcissus and expressed in other plants.

(Q]\PH *HQH%DQN DFFHVVLRQQXPEHU ,VRODWHGIURP 7KHSODQWV H[SUHVVHG 5HIHUHQFH 3K\WRHQH V\QWKDVH ; '4 N. pseudonarcissus N. tazetta Oryza sativa DQWLVHQVHH[S LQN. tazetta >@ /\FRSHQH± F\FODVH *4 N. pseudonarcissus Solanum lycopersicum >@ /\FRSHQHȕ

F\FODVH ; N. pseudonarcissus Oryza sativa > @

ȕFDURWHQH

K\GUR[\ODVH $- N. pseudonarcissus >@

2WKHUWKDQWKLVVHYHUDO JHQHVKDYHEHHQLVRODWHGIURPNarcissus DQGXVHGIRUGLIIHUHQW SXUSRVHVLQRWKHUFURSV$IHZRIWKHPDUHPHQWLRQHGLQ7DEOH%HVLGHVDOOWKHXVHV RISXUHVHFRQGDU\PHWDEROLWHV FUXGH Narcissus H[WUDFWVDUHDOVRXVHGLQSHUIXPHU\,WDOO SURYHVWKDWQRZ DGD\V Narcissus LVQRWRQO\ FRQVLGHUHG DQRUQDPHQWDOFURS EXWDOVRD PHGLFLQDOFURS RQLQGXVWULDOVFDOH GXHWRLWVYDOXHWRSURGXFH XVHIXOFRPSRXQGV

7KHVLV $LP

(9)

ϭϰ

7KHVLV2XWOLQH

(10)

ϭϱ

5HIHUHQFHV

$'$6 Narcissus %XOE 3URGXFWLRQ %RRNOHW D $OQZLFN 0$)) 3XEOLFDWLRQV

0HHURZ$:DQG'$6QLMPDQ$PDU\OOLGDFHDHLQThe Families and Genera of Vascular Plants..XELW]NL(GLWRU6SULQJHU%HUOLQS

%DUUHWW6&+DQG/'+DUGHU7KH(YROXWLRQRI3RO\PRUSKLF6H[XDO6\VWHPV LQ'DIIRGLOVNarcissus1HZ3K\WRORJLVWS

'RRUHQERV - 1RWHV RQ WKH +LVWRU\ RI %XOE %UHHGLQJ LQ WKH 1HWKHUODQGV (XSK\WLFDS

g]]DPEDN 0( ( =H\EHNR÷OX g 7XQFD\ 6 %DúHU * +DVSRODW DQG $ 2OJXQ $ 6XUYH\ RQ &XW )ORZHU 3UHIHUHQFHV DQG ([SHFWDWLRQV $FWD +RUWLFXOWXUDHS

%HQVFKRS05.DPHQHWVN\0/H1DUG+2NXERDQG$'H+HUWRJK7KH *OREDO)ORZHU%XOE,QGXVWU\3URGXFWLRQ 8WLOL]DWLRQ5HVHDUFKLQHorticultural Reviews-RKQ:LOH\ 6RQV,QFS

+DQNV *5 &RPPHUFLDO 3URGXFWLRQ RI Narcissus %XOEV LQ Narcissus and Daffodil: The Genus Narcissus*5+DQNV(GLWRU7D\ORUDQG)UDQFLV 1HZ<RUNS

'ZHFN$&7KH)RONORUHRINarcissusLQNarcissus and Daffodil: The Genus Narcissus*5+DQNV(GLWRU7D\ORUDQG)UDQFLV1HZ<RUNS %DVWLGD-DQG)9LODGRPDW$ONDORLGVRINarcissusLQ Narcissus and Daffodil:

The Genus Narcissus*5+DQNV(GLWRU7D\ORUDQG)UDQFLV1HZ<RUN S

%DVWLGD-5/DYLOODDQG)9LODGRPDW&KHPLFDODQG%LRORJLFDO$VSHFWVRI Narcissus$ONDORLGVLQThe Alkaloids(OVHYLHU

(11)

ϭϲ

dLWR÷OX*07DQNHUDQG%*PúHO$QWLLQIODPPDWRU\(IIHFWVRI /\FRULQH DQG+DHPDQWKLGLQH3K\WRWKHUDS\5HVHDUFKS

:HQLJHU % / ,WDOLDQR -3 %HFN - %DVWLGD 6 %HUJRxRQ & &RGLQD $ /REVWHLQDQG5$QWRQ&\WRWR[LF$FWLYLW\RI$PDU\OOLGDFHDH$ONDORLGV3ODQWD 0HGS

*KRVDO6.66DLQLDQG65D]GDQCrinum $ONDORLGV7KHLU&KHPLVWU\DQG %LRORJ\3K\WRFKHPLVWU\S

-LQ = $PDU\OOLGDFHDH DQG 6FHOHWLXP $ONDORLGV 1DWXUDO 3URGXFW 5HSRUWV S

'HQJ / 3 'DL $ &LUR ') 6PHH + 'MDEDOODK DQG 6 6KXPDQ ,GHQWLILFDWLRQ RI 1RYHO $QWLSR[YLUDO $JHQWV 0LWR[DQWURQH ,QKLELWV 9DFFLQLD 9LUXV 5HSOLFDWLRQ E\ %ORFNLQJ 9LULRQ $VVHPEO\ -RXUQDO RI 9LURORJ\ S

+ZDQJ <& --+ &KX 3/ <DQJ : &KHQ DQG 09 <DWHV 5DSLG ,GHQWLILFDWLRQRI,QKLELWRUV7KDW,QWHUIHUHZLWK3ROLRYLUXV5HSOLFDWLRQ8VLQJD &HOO%DVHG$VVD\$QWLYLUDO5HVHDUFKS

)HQQHOO &: DQG 9- 6WDGHQ &ULQXP 6SHFLHV LQ 7UDGLWLRQDO DQG 0RGHUQ 0HGLFLQH-RXUQDORI(WKQRSKDUPDFRORJ\S

0F1XOW\---1DLU-%DVWLGD63DQGH\DQG&*ULIILQ6WUXFWXUH$FWLYLW\ 6WXGLHV RQ WKH /\FRULQH 3KDUPDFRSKRUH $ 3RWHQW ,QGXFHU RI $SRSWRVLV LQ +XPDQ/HXNHPLD&HOOV3K\WRFKHPLVWU\S

'HO*LXGLFH/'50DVVDUGR33RQWLHULDQG.:ROI,QWHUDFWLRQEHWZHHQ <HDVW0LWRFKRQGULDODQG1XFOHDU*HQRPHV1XOO$OOHOHVRI5WJ*HQHV$IIHFW 5HVLVWDQFH WR WKH $ONDORLG /\FRULQH LQ 5KR 3HWLWHV RI Saccharomyces Cerevisiae*HQHS

(YLGHQWH $ 2 $UULJRQL 5 /LVR * &DODEUHVH DQG * 5DQGD]]R )XUWKHU ([SHULPHQWVRQ6WUXFWXUH$FWLYLW\5HODWLRQVKLSVDPRQJWKH/\FRULQH$ONDORLGV 3K\WRFKHPLVWU\S

(12)

ϭϳ

%DVWLGD - & &RGLQD ) 9LODGRPDW 0 5XELUDOWD -& 4XLULRQ DQG % :HQLJHU1DUFLVVXV$ONDORLGV;Y5RVHULQHIURPNarcissus Pallidulus-RXUQDO RI1DWXUDO3URGXFWVS

%DVWLGD - -0 )HUQiQGH] ) 9LODGRPDW & &RGLQD DQG * GH OD )XHQWH $ONDORLGVIURPNarcissus Tortuosus3K\WRFKHPLVWU\S -LQ = $PDU\OOLGDFHDH DQG 6FHOHWLXP $ONDORLGV 1DWXUDO 3URGXFW 5HSRUWV

S

-LQ = = /L DQG 5 +XDQJ 0XVFDULQH ,PLGD]ROH 2[D]ROH 7KLD]ROH $PDU\OOLGDFHDHDQG6FHOHWLXP$ONDORLGV1DWXUDO3URGXFW5HSRUWV S /HZLV-5$PDU\OOLGDFHDHDQG6FHOHWLXP$ONDORLGV1DWXUDO3URGXFW5HSRUWV S /HZLV-5$PDU\OOLGDFHDHDQG6FHOHWLXP$ONDORLGV 1DWXUDO3URGXFW5HSRUWV S

6WDUN /0 ;) /LQ DQG /$ )OLSSLQ 7RWDO 6\QWKHVLV RI $PDU\OOLGDFHDH 3\UURORSKHQDQWKULGLQLXP $ONDORLGV 9LD WKH =LHJOHUí8OOPDQQ 5HDFWLRQௗ 7RUWXRVLQH&ULDVEHWDLQHDQG8QJHUHPLQH7KH-RXUQDORI 2UJDQLF&KHPLVWU\ S

ZHEOLQN+WWS:ZZ+VX(GX,QWHULRU$VS[",G +WWS/SL2UHJRQVWDWH(GX):)ODYRQRLG+WPO

'HPPLJ$GDPV % DQG :: $GDPV 7KH 5ROH RI ;DQWKRSK\OO &\FOH &DURWHQRLGVLQWKH3URWHFWLRQRI3KRWRV\QWKHVLV7UHQGVLQSODQWVFLHQFH S

+DYDX[ 0 DQG .. 1L\RJL 7KH 9LROD[DQWKLQ &\FOH 3URWHFWV 3ODQWV IURP 3KRWRR[LGDWLYH 'DPDJH E\ 0RUH 7KDQ 2QH 0HFKDQLVP 3URFHHGLQJV RI WKH 1DWLRQDO$FDGHP\RI6FLHQFHVS

(13)

ϭϴ

3RJVRQ %- .. 1L\RJL 2 %M|UNPDQ DQG ' 'HOOD3HQQD $OWHUHG ;DQWKRSK\OO &RPSRVLWLRQV $GYHUVHO\ $IIHFW &KORURSK\OO $FFXPXODWLRQ DQG 1RQSKRWRFKHPLFDO 4XHQFKLQJ LQ $UDELGRSVLV 0XWDQWV 3URFHHGLQJV RI WKH 1DWLRQDO$FDGHP\RI6FLHQFHVS

'DPPH (-09 :- 3HXPDQV $ %DUUH DQG 3 5RXJp 3ODQW /HFWLQV $ &RPSRVLWHRI6HYHUDO'LVWLQFW)DPLOLHVRI6WUXFWXUDOO\DQG(YROXWLRQDU\5HODWHG 3URWHLQV ZLWK 'LYHUVH %LRORJLFDO 5ROHV &ULWLFDO 5HYLHZV LQ 3ODQW 6FLHQFHV S

7HQFKHYD - , <DPEROLHY DQG = =KLYNRYD -RXUQDO 2I &KURPDWRJUDSK\ S± 

%DO]DULQL - &) 3HUQR ' 6FKROV DQG ( 'H &OHUFT $FWLYLW\ RI $F\FOLF 1XFOHRVLGH3KRVSKRQDWH$QDORJXHVDJDLQVW+XPDQ,PPXQRGHILFLHQF\9LUXVLQ 0RQRF\WH0DFURSKDJHVDQG3HULSKHUDO%ORRG/\PSKRF\WHV%LRFKHPLFDODQG %LRSK\VLFDO5HVHDUFK&RPPXQLFDWLRQVS :HLOHU%(+&6FKU|GHU96WHIDQRYLFK'6WHZDUW-06)RUUHVW/% $OOHQ%-%RZGHQ0+.UHXWHU59RWKDQG:(*0OOHU6XOSKRHYHUQDQ D3RO\DQLRQLF3RO\VDFFKDULGHDQGWKHNarcissus /HFWLQ3RWHQWO\,QKLELW+XPDQ ,PPXQRGHILFLHQF\ 9LUXV ,QIHFWLRQ E\ %LQGLQJ WR 9LUDO (QYHORSH 3URWHLQ -RXUQDORI*HQHUDO9LURORJ\S

%H\HU36$O%DELOL;<H3/XFFD36FKDXE5:HOVFKDQG,3RWU\NXV *ROGHQ5LFH,QWURGXFLQJWKH^%HWD`&DURWHQH%LRV\QWKHVLV3DWKZD\LQWR5LFH (QGRVSHUPE\ *HQHWLF (QJLQHHULQJ WR 'HIHDW 9LWDPLQD'HILFLHQF\ -1XWU S6

%XUNKDUGW 3 0 6FKOHG] DQG , 3RWU\NXV 7UDQVJHQLF 5LFH 2U\]D 6DWLYD (QGRVSHUP ([SUHVVLQJ 'DIIRGLO 1DUFLVVXV 3VHXGRQDUFLVVXV 3K\WRHQH 6\QWKDVH $FFXPXODWHV 3K\WRHQH D .H\ ,QWHUPHGLDWH RI 3URYLWDPLQ D %LRV\QWKHVLV3ODQW-S

(14)

ϭϵ

/X*4=RX'*XR;=KXDQJ;<X;;LDQJDQG-&DR$JUREDFWHULXP 7XPHIDFLHQV 0HGLDWHG 7UDQVIRUPDWLRQ RINarcissus Tazzeta 9DU &KLQHQVLV 3ODQW&HOO5HSRUWVS

$SHO : DQG 5 %RFN (QKDQFHPHQW RI &DURWHQRLG %LRV\QWKHVLV LQ 7UDQVSODVWRPLF 7RPDWRHV E\ ,QGXFHG /\FRSHQHWR3URYLWDPLQ D &RQYHUVLRQ 3ODQW3K\VLRORJ\S

$O%DELOL6(+REHLNDDQG3%H\HU$&GQD(QFRGLQJ/\FRSHQH&\FODVH $FFHVVLRQ1R;IURPNarcissus Pseudonarcissus /3UJ3ODQW 3K\VLRORJ\S

(15)