Approaches and Perspectives towards Molecular Breeding of Orchids

Fure-Chyi Chen, G. Mangai asthuri, !i-"ung #sai, "ian-$hi %uang, &en-'i 'ee, (oy !uan-%ung 'uo, )u-Feng Chiang, !a-%uei Chen, #een-Chi Cheng, Man*u M. George +ational Pingtung ,niversity of )cience - #echnology, #aiwan

A.stract
Orchids are the largest family of flowering plants and consist of more than 800 genera and 25,000 species of which many are of commercial importance. Phalaenopsis and Oncidium are two major and popular orchids grown for commercial production. The commodity of these orchids includes potted plant and cut flower production. No elty is the dri ing force in ornamental plant industry. The no el forms !i.e." ariant culti ars or new flower colors, shapes, plant and inflorescence architecture, longer shelf life, fragrance modifications, disease and pest resistance are partly achie ed through con entional se#ual hy$ridi%ation and can $e amended $y ad ancement and applications of gene transfer or genetic transformation mediated either $y Agrobacterium tumefaciens or particle $om$ardment for the de elopment of culti ars with no el or aesthetic properties as traditional $reeding process aimed at genetic modification is always limited $y long reproducti e cycle and cross incompati$ility. This article summari%es personal iews a$out Phalaenopsis and Oncidium research, the current state of art towards molecular $reeding of orchids with reference to color mutants, flower color manipulation, control mechanism of flowering etc., and future perspecti es on molecular $reeding of economic orchids species.

/ntroduction
Orchidaceae is one of the largest families of flowering plants, which consists of more than 800 genera and 25,000 species of which many are of commercial importance. Phalaenopsis and onicidium are the two major and popular orchids grown for commercial production as cut flower and potted plants. Potted orchid plants ha e $een the prime importance for agro industries world wide in recent years !&ries$ach, 2002".

Traditional $reeding methods !i.e. continuous crossing and selection" ha e pa ed the way for the $reeders to create new arieties that ha e desira$le traits i%. color, shape, fragrance, plant architecture, ase life and resistance to disease and pests $ut this 'ind is the limited gene pool of any single species. (olecular approaches ha e opened a new way for the genetic transformation of plants to create no el traits, such as flower color modulations, floral formation, plant and inflorescence architecture, fragrance etc. Potential application of this technology for orchid impro ement is $eing pursued in many institutions. Phalaenopsis and onicidium orchids are $eing systematically produced as high alue cash crops in Taiwan. The creation of mutant culti ar has $ecome an important factor in the mar'et for commercial orchid production. No elty is the dri ing force in ornamental plant industry. )ence molecular genetic modification can $e adopted in addition to traditional $reeding for the emergence of plants with no el aesthetic properties. Transgenic plants are produced ia Agrobacterium*mediated and other direct +N, transfer methods li'e P-&, electroporation, microprojectile $om$ardment, and microinjection. .n orchids protocorms from germinated seeds or protocorms*li'e*$odies !P/0s" deri ed from shoot tip or lea es are the most easily o$tained materials that are capa$le of regenerating plants. The routine transformation procedure for orchids ia either Agrobacterium*mediated or microprojectile $om$ardment for introducing genes with horticultural and economically important traits, such as irus disease resistance, is $eing started. 1e eral studies on successful transformation of Phalaenopsis, Oncidium, Cymbidium, and Dendrobium ha e $een reported. )owe er, long period of selection and regeneration, and low reco ery of transgenic plants ha e hindered the efficient transformation of these recalcitrant orchid species. .n this report, we try to ela$orate the potential application of genes with alua$le horticultural traits reported in other crop plants in orchids. ,pproaches and strategies for o$taining important genes are also discussed.

#argets for 0olecular flower .reeding of orchids

Flower color 0odification 2lower color of higher plants is de to the production of pigments, including fla onoids, carotenoids, and $etalains !3hristinet et al., 12234 Tre%%ini 4 5r6d, 78879 :in'el*1hirley, 1225". 2lo onoids contri$ute wide spectrum of colors in plants, including red, $lue, yellow and purple pigments. 1i# su$groupds of the fla onoids are widespread in plants, including chalcones, fla ones, fla onols, fla andiols,

anthocyanins, and condensed tannins !or proanthocyanidins"! :in'el*1hirley, 1225". The $iosynthesis of anthocyanin pigments and fla onol copigments made flowers showy and function to recruit pollinators and seed dispersers. +irect modification of anthocyanin production has $een achie ed in se eral plant species, such as petunia and carnation !(ol et al., 7888". :ildtype carnation was modified to produce $lue colors after introducing a heterologous fla onoid ;<,5<*hydro#ylase gene !(ol et al., 78889 2u'ui et al., 200;". Transgenic carnation with selected color shades has $een introduced in =apan mar'et. This points the possi$ility of color modification through gene technology in the non*food flower crops such as orchids. The $lue petals of the morning glory ! Ipomoea tricolor c . )ea enly 0lue" change from purplish red to $lue during flower opening. The color shift was due to an increase of the acuole p) from >.> to ?.? in the cells of epidermal layers !@amaguchi et al., 20079 @oshida et al. 7885". The increase of acuolar p) in the petals during flower opening was due to acti e transport of NaA andBor CA from the cytosol into acuoles $y the NaAB)A e#changer !N)D7". .mmunochemical and Northern blot analyses have confirmed the correlation of the petal color change with the N)D7 !@amaguchi et al., 20079 @oshida et al., 2005". The manipulation of color shift to $lue may ha e the potential application in phalaenopsis color modification using the Na AB)A e#changer from morning glory, Doritis pulcherrima !$lue flowers", or other plant species using genetic transformation technology. .nterestingly, the metal ion moly$denum !(o" seEuestration in the plant acuoles was shown to $ind with anthocyanins to ma'e the petals $lue !)ale et al., 2007". This may also open an alternati e choice for transgenic $lue phalaenopsis. 2la onoid $iosynthetic gene e#pression is regulated $y the cooperation of F2F; (@0 and $asic heli#*loop*heli# !$)/)" transcription factors, which acti ate anthocyanin $iosynthesis !)ernande% et al., 200G". The (@0 transcription factors from Arabidopsis, Gerbera, petunia and tomato ha e $een shown to function in transgenic plants producing anthocyanins in petals as well as other tissues !0ore it% et al., 20009 -lomaa et al., 200;9 (athews et al., 200;9 (ol et al., 78889 Famsay et al., 200;". ,NT)O3@,N.N7 !,N7" of petunia is a $)/) transcription factor reEuired for the synthesis of anthocyanin pigments !1pelt et al., 2002". Transgenic plants e#pressing ,N7 can acti ate the e#pression of the dfr, gene encoding the en%yme dihydrofla onol G*reductase !1pelt et al., 2000". 1imilarly, o ere#pression of the tomato (@0 transcription factor ANT in to$acco and tomato up*regulated the e#pression of anthocyanin $iosynthesis genes in transgenic plants !(athews et al., 200;". ,n alternati e approach for enhancing flower color is $y the manipulation of the fla onoid* $inding protein. Petunia ,N8 is a glutathione !*transferase responsi$le for the

seEuestration of anthocyanins into acuoles to ma'e deeper flower color !(ueller et al., 2000". The mai%e "#$ gene also encodes glutathione !*transferase which is responsi$le for the anthocyanin accumulation in the acuoles !(arrs et al., 7885". b%$ mutants led to the accumulation of anthocyanins in the cytoplasm where they were o#idi%ed into $rown color !,lfenito et al., 7888". ,n unusual flower pigment, $etalains, has recently $een studied in some detail. The genes related to $etalain $iosynthesis ha e $een cloned. One of them, DODA, encoding G,5*e#tradiol dio#ygenase, was isolated and characteri%ed from Portulaca grandiflora at molecular le el. The gene could function in a mutant white Portulaca petal to complement the production of yellow pigments in the $om$arded cells !3hristinet et al., 1223". This finding may ha e a potential application of the DODA gene in color modification for orchids. (any yellow*flower plants accumulate carotenoids $y carotenoid*$inding protein located in the chromoplasts !Hishne ets'y et al., 7888a". , chromoplast*specific carotenoid*associated protein !3)F3" was identified and cloned from cucum$er corolla !Hainstein et al., 788G9 Hishne ets'y et al., 788>". The promoter of the 3)F3 gene was fused to the reporter gene and $om$arded into cucum$er petals and other tissues, the result showed uniEue e#pression of the 3)F3 promoter in the cucum$er petal !Hishne ets'y et al., 7888$". Three carotenoid*associated protein, or plastid lipid* associated protein !P,P", were also isolated from "rassica rapa, with the P,P2 most a$undant in the yellow petals !Cim et al., 2007". This result prompted us to clone the homologous gene, fibrillin, from the yellow*flowered Oncidium. , full length fi$rillin c+N, has $een cloned from the la$ellum !lip" of the Oncidium flowers. To create white or pale yellow flowers of Oncidium, the FN, interference strategy will $e used to transform Oncidium P/0s with the e#pectation of 'noc'out of the carotenoid accumulation in flowers to ma'e them white. , light colored orchid cut flower may ha e the potential mar'et in =apan. /nflorescence architecture .nflorescence $ranching is an important trait in Oncidium cut flowers. ,t least > $ranches in a flower stal' are regarded as top grades for domestic and e#port mar'ets. +uring the hot and humid summer time in su$tropical areas li'e Taiwan, the inflorescence $ranching is reduced to minimal degree, with pro$a$ly only one single $ranchless main stem present. +uring winter to spring time, the inflorescence $ranching is enhanced. The control mechanism for this inflorescence architecture in Ondicium orchid is currently un'nown. O$ser ations and e#periments from other plant species ha e contri$uted to our

understanding of the mechanisms of shoot $ranching. Through the study of mutants in shoot $ranching of Arabidopsis and pea, se eral genes were implied to play a 'ey role for the shoot $ranching phenotype !(orris et al., 2007". .n pea, reduced cyto'inin content in root #ylem sap was o$ser ed in some $ranching mutants $ut not in others. , no el graft*transmissi$le signal in ol ed in the control of shoot $ranching was proposed !2oo et al., 2007, 20059 Morris et al., 20079 Turn$ull et al., 2002". , mo$ile $ranch* inhi$iting signal may ha e $een produced in the wildtype plant, and could $e transmitted through grafting to the $ranching mutant ma&' of Arabidopsis !1orefan et al., 200;". The gene encodes an au#in*induci$le polyene dio#ygenase family protein !1orefan et al., 200;". , carotenoid clea age dio#ygenase gene ! Dad (PhCCD) 6 was identified in petunia shoot and root. The 33+ en%yme regulates the $iosynthesis a graft* transmissi$le compound that inhi$its $ranching. .n the dad* mutant, the Dad (PhCCD) was mutated and reduced the mo$ile signal so that more shoot $ranching was o$ser ed !1nowden et al., 2005". The graft*transmissi$le $ranch* inhi$iting hormones were shown to $e related to the plastidic dio#ygenase that clea ed multiple carotenoids in ara$idopsis !0oo'er et al., 200G9 1chwart% et al., 200G". ,nother gene, +A,$, controlling shoot lateral $ranching has $een cloned and shown to $e an 2* $o# leucine*rich repeat family of proteins !1tirn$erg et al., 2002". 3urrently, a 33+ homolog from Oncidium &ower Famsey has $een cloned from the flower $uds in this la$ !3hiang, 1. 2., @. =. Tsai 4 2. 3. 3hen, unpu$lished". -#pression le el of the 33+ in the flower stem as well as other tissues of the Oncidium plants will $e determined to chec' its relation with inflorescence $ranching. Transgenic approach with 33+ 'noc'out $y FN, interference strategy is $eing tested in order to o$tain free*$ranching Oncidum cut flowers. ,s there is se eral color mutants of the Oncidium &ower Famsey, namely the wildtype, the al$ino clone I:hite =adeJ, and the orange mutant I1un'istJ, they are alua$le plant materials for molecular genetic studies regarding to the carotenoid pigments changes due to their uniform genetic $ac'ground. They ary in carotenoid contents in the flowers. The mutants will offer us the opportunity to e#amine the relationship of carotenoid $iosynthesis and accumulation, cloning and comparison of the corresponding genes, and su$seEuent transgenic studies to elucidate their functions in flower pigmentation of Oncidium orchids. Fragrance 0anipulation 3onsumers are always allured $y orchids with scents. (any orchid species emit fragrance during the process of flowering. (any Phalaenopsis species, such as violacea, bellina, e-uestris, amboinensis, leuddemaniana, schilleriana, and parishii, will emanate mi#tures of olatile components to attract insects and small animals,

pro$a$ly for the purpose of pollination $y these ectors, or as a defense mechanism !da 1il a et al., 78889 +udare a et al., 200G9 Caiser, 788;9 Nishida et al., 200G9 Pichers'y 4 &ershen%on, 2002". The main components in the P. violacea !bellina" are linalool and geraniol !Caiser, 788;". .t is rather difficult to isolate genes corresponding to the target component from Phalaenopsis flowers. The gene encoding linalool synthase has $een isolated and characteri%ed from other plant species, such as Artemisia, Clar.ia, $asil !3se'e et al., 78889 +udare a et al., 788>9 .ijima et al., 200G$9 =ia et al., 7888". , geraniol synthase gene has also $een isolated from the $asil !.ijima et al., 200Ga". The linalool synthase gene of the 3lar'ia has $een introduced into tomato genome. Transgenic tomato fruit produced more !*linalool and 8*hydro#ylinalool in ripening fruits !/ewinsohn et al., 2007". 1imilarly, three monoterpene synthase genes fro0 le0on have .een introduced into to.acco with increased fragrance in to.acco flowers 7/uc'er et al., 200G". These results suggest us the potential use of heterologous genes for introducing into orchid genomes to ma'e the orchid flowers more fragrant. Fecently, we ha e cloned a linalool synthase*li'e gene from c+N, su$tracted li$rary of the Phalaenopsis flowers. /t will .e characteri8ed at 0olecular as well as .ioche0ical levels to confir0 its role in scent production of Phalaenopsis. Flowering control 2lowering $eha ior aries in different orchids. 1ome species responds to photoperiod and some to growth regulators !&oh4 ,rditti, 7885". .n Phalaenopsis, a four*wee' of night temperature at 75*20 and day temperature at 25 will induce the spi'ing of inflorescence !/ee 4 /in, 788G9 1a'anishi et al., 7880". /ight intensity during the low temperature induction period may affect the spi'ing and Euality of the phalaenopsis plants !Conow 4 :ang, 20079 1a'a'i$ara et al., 78859 1ugiyama et al., 2007". .t has $ecome a standard practice to control flowering of the Phalaenopsis plants in the orchid nurseries worldwide. There are two aspects of flowering control. One is how to control flowering in schedule production. The other one is how to pre ent spi'ing of the mature plants in order to e#port to foreign growers for forcing. To pre ent or delay spi'ing of mature phalaenopsis plants, growers usually raise greenhouse temperature up to 28*;0 during the growing period. )owe er, many hy$rids still may spi'e at some degree during the high temperature egetati e growth stage. .n order to understand the mechanism of inflorescence induction and su$seEuent floral meristem de elopment, it is necessary to isolate genes up* or down*regulated during the flowering induction period. Fegulation of gene e#pression has $een intensi ely studied in the model plant, Arabidopsis !1impson et al., 7888" and many monocot species. The results indicated

conser ation of similar genes among different plant species !)e 4 ,masino, 2005". One of the 'ey genes related to phase change from egetati e to reproducti e stage is the /0A12 !/12" gene. :e ha e recently cloned the /12 gene from a Phalaenopsis hy$rid. .ts function in ol ed in flowering will $e studied $y transformation of the gene into an ,ra$idopsis mutant with defect in /12. The promoter of the Phalaenopsis /12 will also $e studied to elucidate its role in flower induction and plant de elopment.

Orchid #ransfor0ation #echnology

&enetic transformation of se eral orchids either through Agrobacterium or particle $om$ardment*mediated techniEues has $een reported, including 3ym$idium, +endro$ium, Oncidium and Phalaenopsis !,n%ai et al., 788>9 0elarmino 4 (ii, 20009 3hai et al., 788G, 20029 Cnapp et al., 20009 Cuehnle 4 1ugii, 78829 /iau et al., 200;a, $9 (en et al., 200;a, $9 @ang et al., 78889 @u et al., 2007". ,s regeneration protocol for each orchid species aries, to achie ing genetic transformation of the target orchid hy$rids reEuires a relia$le and efficient regeneration system while a oiding somaclonal ariation in the transgenic plants. +espite of many reports on tissue culture and su$seEuent plant regeneration, the protocol may not wor' during the anti$iotic selection procedure due to their sensiti ity to the chemicals, and needs to $e optimi%ed for each species.

Perspectives
Orchids ha e $ecome daily consumption produce of most de eloped or de eloping countries. 3on entional $reeding wor's ha e contri$uted to the a alanche new hy$rids e ery year. 0iotechnology or molecular $iology techniEues will play an important role for the impro ement of commercial traits of orchid hy$rids when com$ined with tissue culture system. .solation and characteri%ation of genes regulating plant growth and de elopment of orchid species will help us understand their function. :e optimistically foresee the orchid hy$rids with no el traits through molecular $reeding in the near future.

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1ci. ;K272*27?. (ol =, 3ornish -, (ason =, Coes F. 7888. No el colored flowers. 3urr. Opin. 0iotechnol. 70K788*207. (orris 1-, Turn$ull 3&, (urfet .3, 0e eridge 3,. 2007. (utational analysis of $ranching in pea. - idence that 6ms and 6ms7 regulate the same no el signal. Plant Physiol. 72>K7205*727;. (ueller /,, &oodman 3+, 1ilady F,, :al$ot H. 2000. ,N8, a petunia glutathione 1* transferase reEuired for anthocyanin seEuestration, is a fla onoid*$inding protein. Plant Physiol. 72;K75>7*75?0. Nishida F, Tan C*), :ee 1*/, )ee ,C*:, Toong @*3. 200G. Phenylpropanoids in the fragrance of the fruit fly orchid, "ulbophyllum cheiri, and their relationship to the pollinator, "actrocera papayae5 0iochem. 1ys. -col. ;2K2G5*252. Pichers'y -, &ershen%on =. 2002. The formation and function of plant olatilesK perfumes for pollinator attraction and defense. 3urr. Opin. Plant 0iol. 5K2;?L2G;. Famsay N,, :al'er ,F, (ooney (, &ray =3. 200;. Two $asic*heli#*loop*heli# genes !+2C* '8 and G/3" from Arabidopsis can acti ate anthocyanin $iosynthesis in a white*flowered +atthiola incana mutant. Plant (ol. 0iol. 52!;"K>?8*>88. 1a'anishi @, .manishi ), .shida &.. 7880. -ffect of temperature on growth and flowering of Phalaenopsis amabilis. 0ull. Mni . Osa'a Pref. 1er. 0 ;2K 7L8. 1a'a'i$ara T, (inami ), Cawase C. 7885. -ffects of light intensity in cooling treatment after flower inducti e growing period on flowering of Phalaenopsis. 0ull. -#p. 2arm Cyoto Mni . >K;?LG7. 1chwart% 1), Pin D, /oewen (. 200G. The $iochemical characteri%ation of two carotenoid clea age en%ymes from Arabidopsis indicates that a carotenoid*deri ed compound inhi$its lateral $ranching. =. 0iol. 3hem. 2?8KG>8G0LG>8G5. 1impson &&, &endall ,F, +ean 3. 7888. :hen to switch to flowering. ,nnu. Fe . 3ell +e . 0iol. 75K 578 550. 1orefan C, 0oo'er =, )aurognQ C, &oussot (, 0ain$ridge C, 2oo -, 3hatfield 1, :ard 1, 0e eridge 3, Fameau 3, /eyser O. 200;. +A,' and 6+! are orthologous dio#ygenase*li'e genes that regulate shoot $ranching in Arabidopsis and pea. &enes +e . 7?K7G>8*7G?G. 1nowden C3, 1im'in ,=, =anssen 0=, Templeton CF, /oucas )(, 1imons =/, Carunairetnam 1, &lea e ,P, 3lar' +&, Clee )=. 2005. The Decreased apical dominance (Petunia hybrida CA6OT0NOID C/0A9AG0 DIO,2G0NA!0) gene affects $ranch production and plays a role in leaf senescence, root growth, and flower de elopment. Plant 3ell 7?K?G>*?58. 1pelt 3, Puattrocchio 2, (ol =N, Coes F. 2000. anthocyanin of petunia encodes a

$asic heli#*loop*heli# protein that directly acti ates transcription of structural anthocyanin genes. Plant 3ell 72K7>78*7>;2. 1pelt 3, Puattrocchio 2, (ol =, Coes F. 2002. ,NT)O3@,N.N7 of petunia controls pig0ent synthesis, vacuolar p%, and seed coat develop0ent .y genetically distinct 0echanis0s. Plant 3ell 7GK 2727*27;5. 1tirn$erg P, an +e 1ande C, /eyser )(. 2002. +A, and +A,$ control shoot lateral $ranching in Arabidopsis. +e elopment 728K77;7*77G7. 1ugiyama @, Cu$ota 1, Coshio'a (. 2007. +elaying anthesis $y dar' treatment in Phalaenopsis. =. =pn. 1oc. )ort. 1ci. ?0K2>G*2>>. Tre%%ini &2, 5r6d =P. 7887. Two $etalains from Portulaca grandiflora. Phytochemistry ;0K788?*7888. Turn$ull 3&, 0oo'er =P, /eyser )(. 2002. (icrografting techniEues for testing long* distance signalling in Arabidopsis. Plant =. ;2K255*2>2. Hainstein ,, )ale y ,), 1mirra ., Hishne ets'y (. 788G. 3hromoplast $iogenesis in Cucumis sativus corollas !rapid effect of gi$$erellin ,; on the accumulation of a chromoplast*specific carotenoid*associated protein". Plant Physiol. 70GK;27*;2>. Hishne ets'y (, O adis (, .t%ha'i ), /e y (, /i$al*:e'sler @, ,dam 5, Hainstein ,. 788>. (olecular cloning of a carotenoid*associated protein from Cucumis sativus corollasK homologous genes in ol ed in carotenoid seEuestration in chromoplasts. Plant =. 70K7777*7778. Hishne ets'y (, O adis (, Hainstein ,. 7888a. 3arotenoid seEuestration in plantsK the role of carotenoid*associated proteins. Trends Plant 1ci. GK2;2*2;5. Hishne ets'y (, O adis (, 5u'er ,, Hainstein ,. 7888$. (olecular mechanisms underlying carotenogenesis in the chromoplastK multile el regulation of carotenoid* associated genes. Plant =. 20KG2;*G;7. :in'el*1hirley B. 1225. 2la onoid $iosynthesis. , colorful model for genetics, $iochemistry, cell $iology, and $iotechnology. Plant Physiol. 72>KG85*G8;. @amaguchi T, 2u'ada*Tana'a 1, .naga'i @, 1aito N, @one'ura*1a'a'i$ara C, Tana'a @, Cusumi T, .ida 1. 2007. &enes encoding the acuolar Na AB)A e#changer and flower coloration. Plant 3ell Physiol. G2K G57LG>7. @ang =, /ee )=, 1hin +), Oh 1C, 1eon =), Pae' C@, )an C). 7888. &enetic transformation of Cymbidium orchid $y particle $om$ardment. Plant 3ell Fep. 78K 8?8*88G. @oshida C, Condo T, O'a%a'i @, Catou C. 7885. 3ause of $lue petal colour. Nature ;?;K 287. @oshida C, Cawachi (, (ori (, (aeshima (, Condo (, Nishimura (, Condo T. 2005. The involve0ent of tonoplast proton pu0ps and +a >7 >69%> e=changers in

the change of petal color during flower opening of 0orning glory, Ipomoea tricolor cv. %eavenly Blue. Plant 3ell Physiol. G>K G0?*G75. @u ), @ang 1), &oh 3=. 2007. Agrobacterium*mediated transformation of a Dendrobium orchid with the class 7 .no& gene DO: . Plant 3ell Fep. 20K;07*;05.

G. Mangai asthuri Man*u M. George

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 !(orris et al., 2007"K{($"# {wUGKW@I9,2(8 l { : & ? !2oo et al., 2007, 20059 Morris et al., 20079 Turn$ull et al., 2002"#%JKLM&#5.{2(I l9,P{ ma&' !1orefan et al., 200;"ma&' [V` I # 4 polyene dio#ygenase * + ( !1orefan et al., 200;" s 8 r s m o dio#ygenase X[V !Dad (PhCCD) 6 33+ XGH9{2(#3N dad* Dad (PhCCD) !1nowden et al., 2005" JKLM 9_{ ?7 HI plastidic dio#ygenase o carotenoids !0oo'er et al., 200G9 1chwart% et al., 200G"&l { [ V+A,$ ? R a 2*$o# w leucine * + ( !1tirn$erg et al., 2002" pO s &ower Famsey Ra { 33+ [ V !3hiang, 1. 2., @. =. Tsai 4 2. 3. 3hen, "yzs1 33+ @UQ8{.:k3 FN,i 33+ { .B+IJ &ower r Famsey no+ I J !I:hite =adeJ" I J !I1un'istJ"IJQz carotenoid f99F K$Hwx{f9&12 QzN&mno wUyzNrmno#3_`:Ra 9:[VP`awxK[VN&3 HA??@lnoN} A ? no > violacea bellina e-uestris amboinensis leuddemaniana schilleriana parishii A . 2 ( E 52 T !da 1il a et al., 78889 +udare a et al., 200G9 Caiser, 788;9 Nishida et al., 200G9 Pichers'y 4 &ershen%on, 2002" P. violacea !bellina" ?@3H linalool geraniol !Caiser, 788;"s&{' Clar.ia U3{[VH9MI2T Artemisia ? !linalool" 3[V ? Ra 9: L. k 3 !3se'e et al., 78889 +udare a et al., 788>9 .ijima et al., 200G$9 =ia et al., 7888" !geraniol" 3 [ V?Ra !.ijima et al., 200Ga"T Clar.ia linalool 3[V? `a NO [V & `a kNO5 O & # [ o !*linalool 8* hydro#ylinalool !/ewinsohn et al., 2007" 9 { R I R a ; &

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$etalain pigment $iosynthesis in Portulaca grandiflora. Plant Physiol. 7;GK2>5*2?G. da 1il a M2, 0or$a -/, 1emir =, (arsaioli ,=. 7888. , simple solid injection de ice for the analyses of "ulbophyllum !Orchidaceae" olatiles. Phytochemistry 50Ks ;7*;G. +udare a N, 3se'e /, 0lanc H(, Pichers'y -. 788>. Evolution of floral scent in Clar.iaK no el patterns of 1*linalool synthase gene e#pression in the C. breweri flower. Plant 3ell 8K77;?*77G8. +udare a N, Pichers'y -, &ershen%on =. 200G. 0iochemistry of plant volatiles. Plant Physiol. 7;5K788;*7802. -lomaa P, Mimari ,, (ehto (, ,l$ert H,, /aitinen F,, Teeri T). 200;. ,cti ation of anthocyanin $iosynthesis in Gerbera hybrida !,steraceae" suggests conser ed protein*protein and protein*promoter interactions $etween the anciently di erged monocots and eudicots. Plant Physiol. 7;;K78;7*78G2. 2oo -, Turn$ull 3&, 0e eridge 3,. 2007. /ong*distance signaling and the control of $ranching in the rms mutant of pea. Plant Physiol. 72>!7"K20;*208. 2oo -, 0ullier -, &oussot (, 2oucher 2, Fameau 3, 0e eridge 3,. 2005. The .ranching gene RAMOSUS1 0ediates interactions a0ong two novel signals and au=in in pea. Plant 3ell 7?!2"KG>G*G?G. 2u'ui @, Tana'a @, Cusumi T, .washita T, Nomoto C. 200;. , rationale for the shift in colour towards $lue in transgenic carnation flowers e#pressing the fla onoid ;N,5N* hydro#ylase gene. Phytochemistry >;!7"K75*2;. &oh 3=, ,rditti =. 7885. Orchidaceae. .nK )ale y ,) !ed", )and$oo' of 2lowering. 3F3 Press, 0oca Faton, 2la., pp;08L;;>. &ries$ach F=. 2002. +e elopment of Phalaenopsis orchids for the mass*mar'et. p. G58L G>5. .nK =. =anic' and ,. :hip'ey !eds.", Trends in New 3rops and New Mses. ,1)1 Press, ,le#andria, H,. )ale C/, (c&rath 1P, /om$i -, 1tac' 1(, Terry N, Pic'ering .=, &eorge &N, Pilon* 1mits -,. 2007. (oly$denum seEuestration in "rassica species. , role for anthocyaninsO Plant Physiol. 72>K7;87*G02. )e @, ,masino F(. 2005. Fole of chromatin modification in flowering*time control. Trends Plant 1ci. 70K;0*;5. )ernande% =(, )eine &2, .rani N&, 2eller ,, Cim (&, (atulni' T, 3handler H/, &rotewold -. 200G. +ifferent mechanisms participate in the F*dependent acti ity of the F2F; (@0 transcription factor 37. =. 0iol. 3hem. 2?8!G>"KG8205*G827;. .ijima @, &ang +F, 2ridman -, /ewinsohn -, Pichers'y -. 200Ga. 3haracteri%ation of geraniol synthase from the peltate glands of sweet $asil. Plant Physiol 7;GK ;?0L;?8. .ijima@, +a ido ich*Fi'anati F, 2ridman -, &ang +F, 0ar -, /ewinsohn -, Pichers'y -. 200G$. The $iochemical and molecular $asis for the di ergent patterns in the

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