Effects of temperature and light on germination and early seedling development of the pine pink orchid (Bletia purpurea)psbi_

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1 s_s_nner Plnt Speies Biology (12) 27, oi: /j x NOTES AND COMMENTS Effets of temperture n light on germintion n erly seeling evelopment of the pine pink orhi (Bleti purpure)psi_ TIMOTHY R. JOHNSON n MICHAEL E. KANE Plnt Restortion, Conservtion n Propgtion Biotehnology Lortory, University of Flori, Deprtment of Environmentl Hortiulture, Ginesville, Flori , USA Astrt Orhi see physiology is poorly unerstoo phenomenon owing to n emphsis on proution n the hllenges ssoite with propgting orhis from minute see. We investigte the role of simulte south Flori tempertures n illumintion (rk n 12 h photoperio) in regulting germintion n seeling evelopment using symioti see germintion ssys of Bleti purpure. Our ojetives were to etermine whether in situ germintion is limite y sesonl tempertures n to etermine whether temperture lters responses to illumintion. Bleti purpure sees were le to germinte to > 9% uner ll tretments. The gretest germintion fter 3 weeks ws oserve t 29/19 C uner ontinul rkness n t 25 C uner rk n illuminte onitions. The slowest germintion ws oserve t simulte winter tempertures (22/11 C). Illumintion initilly inhiite germintion n evelopment, ut resulte in equl or greter evelopment y week six. Germintion uner 22/11 C ws strongly inhiite y illumintion, initing n intertion etween temperture n light sensing systems. Keywors: ioti effets, symioti germintion, illumintion, Orhiee, see. Reeive 1 Jnury 11; revision reeive 18 April 11; epte 16 My 11 Introution The importne of temperture in promoting n inhiiting see germintion is well oumente n responses re highly vrile mong speies. Het stress plys role in elying germintion in some speies (Kepzynsk et l. 6; Norsworthy & Oliveir 6) n long exposures to high tempertures n inue seonry ormny (Nsimento et l. ). For other speies, wrm strtifition reues ormny or promotes germintion (Leon & Owen 3; Turner et l. 6). Col strtifition (Bskin & Bskin 1; Moyo et l. 9; Hn & Long 1) n osillting tempertures (Bskin & Bskin 1, 3) hve lso een shown to rek ormny n/or promote germintion. Temperture n lso hve profoun effets on the light sensitivity of sees (Hilton 1984) n my ply n Corresponene: Timothy R. Johnson Emil: timjohn@ufl.eu importnt role in regulting sesonl germintion responses (Heshel et l. 7). Flututing tempertures n lso overome fr-re light-inue inhiition of germintion (Benvenuti et l. 1; Hon & Ktoh 7). This response likely funtions s n eologilly importnt gp-sensing mehnism y whih sees sense openings in the vegettive nopy (Hon & Ktoh 7); re soil surfes re expete to e less insulte from temperture flututions thn pth of soil overe with ense vegettion. As sees tht re isperse to lowompetition sfe site (e.g. re soil) my still e expose to fr-re rih-filtere light from tree nopies, flututing tempertures my e more relile gp initor. Sesonl regultion of germintion timing my lso e influene y onitionl ormny. Conitionlly ormnt sees re le to germinte uner some tempertures, ut not uner the wiest possile rnge of tempertures (Bskin & Bskin 1), resulting in enhne germintion in some sesons or simulte sesonl tempertures (Kettenring & Gltowitsh 7). 11 The Authors Journl ompiltion 11 The Soiety for the Stuy of Speies Biology

2 TEMPERATURE AND ORCHID SEED GERMINATION 175 Although the Orhiee is the lrgest plnt fmily n ounts for lrge proportion of the ioiversity of some eosystems, little is known out the role sesonl temperture plys in regulting orhi see germintion. The germintion-promoting effets of pre-hilling on orhi sees hve een well oumente (Rsmussen 1992; Miyoshi & Mii 1998; Shimur & Ko 5; Kuth et l. 11), initing tht germintion timing is t lest prtilly regulte y physiologil ormny. In ition, flututing tempertures were foun to stimulte germintion of Clopogon tuerosus vr. tuerosus more thn onstnt temperture (Kuth et l. 11). Suh investigtions hve importnt implitions for the propgtion of speies for oth ex situ n in situ onservtion progrms n s mens of reveling something out the poorly unerstoo nture of orhi see germintion in situ. Although propgtion is key gol of mny ex situ orhi onservtion progrms, n unerstning of orhi see physiology is rrely use to guie propgtion prties. Few reports ress the effets of temperture on orhi see germintion, ut rther fous on the effets of ulture mei, plnt growth regultors n other mei itives on germintion n seeling proution. This emphsis on proution hs resulte in few physiologil investigtions of how germintion is regulte in situ. In situ see iting stuies hve een use to exmine fungl speifiity, fungl istriution, see nk longevity n time to germintion (Msuhr & Ktsuy 1994; MKenrik et l. ; MKenrik et l. 2; Whighm et l. 6; Hollik et l. 7), ut the mny onfouning ftors n ourse timesles mke suh stuies less thn iel for unerstning how temperture ffets germintion. To stuy the effets n intertions of temperture n illumintion on orhi see germintion, we rrie out n symioti see germintion experiment using the pine pink orhi, Bleti purpure, from Flori s moel orgnism. Bleti purpure is terrestril speies foun in the new worl tropis n sutropis (Sos & Díz- Dums 1997; Plestin & Sos 2). In North Ameri, the speies is foun throughout Mexio, ut is restrite to Flori in the USA where it is liste s stte thretene (Coile & Grln 3). Bleti purpure oupies severl ifferent hitts in Flori n throughout its rnge. It n e foun in ry or mesi hitts, in or long sru lns or pinelns, on floting logs or stumps in ypress swmps n long highly isture lke eges n liffs (Correll 1978; Dressler 1993; Willims & Allen 1998; Brown 5). The ojetives of the present stuy were to etermine the effet of simulte South Flori sesonl tempertures on B. purpure see germintion n erly seeling evelopment n to etermine whether osillting tempertures promote germintion uner symioti ulture onitions. An itionl ojetive ws to etermine whether there ws n intertion etween the effets of temperture n illumintion (ontinul rkness ompre with 12 h photoperio) on germintion n evelopment. Mterils n methos Five unehise, nerly mture B. purpure psules were ollete from popultion of plnts t the Flori Pnther Ntionl Willife Refuge (FPNWR; Collier County) Burn Unit 6 on 23 My 7. The psules were store over Drierite esint t 22 2 C for 3 weeks until the psules were thoroughly ry n/or ehise. Sees were remove from the psules, poole n homogenize n then ple into ol storge t -1 C over Drierite (W.A. Hmmon, Xeni, OH, USA) esint in the University of Flori Orhi See Bnk. The sees were surfe sterilize for 6 s in solution of 6% Clorox leh : 1% ethnol : istille eionize () wter (1:1:), then rinse three times in sterile wter. The sees were then sown onto 9. m imeter Petri pltes ontining 25 ml sterile Vin n Went Moifie Orhi Mei (V895; PhytoTehnology Lortories, Shwnee Mission, KS, USA; Vin & Went 1949) mene with 1 g/l tivte hrol (Dutr et l. 8). Approximtely 3 5 sees (41 12; verge stnr evition [SD]) were sown onto eh plte. Pltes were sele with single lyer of NesoFilm n rnomly ssigne temperture n illumintion tretment. Petri pltes were ulture in growth hmers uner one of five ifferent temperture regimes. Four of these tretments were osillting 12/12 h (y/night) temperture regimes esigne to pproximte sesonl y/night temperture flututions in entrl Flori. The experimentl tempertures teste were 22/11, 29/19, 33/24 n 27/15 C (representing winter, spring, summer n utumn temperture extremes in south Flori, respetively), s well s onstnt temperture tretment of 25 C. Constnt, or ner onstnt, tempertures in the rnge of 25 C re often use for orhi see germintion stuies (see for exmple Peroz-Mnrique & Min- Gutierrez 6; Stewrt & Kne 6; Ymzki & Miyoshi 6; Luzer et l. 7; Dutr et l. 9; Stewrt & Kne 1). Sees were lso trete with either ontinuous rkness or 12/12 h light/rk (L/D) photoperio t these vrious tempertures. Illumintion ws provie y ool white fluoresent lights t ~5 mmol/m 2 /s 1 ). A ompletely rnomize esign (CRD) ws use for this experiment with four replite pltes for ll tretments. The experiment ws repete one (n = 8). Sees n seelings were oserve fter 3 weeks n 6 weeks of ulture. At these times, t were ollete on germintion n susequent lef evelopment (seelings rehing one Plnt Speies Biology 27, The Authors Journl ompiltion 11 The Soiety for the Stuy of Speies Biology

3 176 T. R. JOHNSON AND M. E. KANE lef, two lef n three lef stges). Sees were ounte s germinte one the test ws rke y the imie emryo. Perentge germintion n the perentges of seelings in eh lef stge were rsine trnsforme prior to sttistil nlysis. An nov n lest squre (LS) men seprtion t =.5 ws rrie out using PROC MIXED in SAS v 8.2 (SAS Institute 2). Results Sees exhiite high germintion perentges (nerly 1%) within 6 weeks, regrless of the tretment. However, low tempertures n illumintion elye oth germintion n evelopment (Figs. 1,2). At week three, sees ulture uner illumintion exhiite signifintly lower germintion thn sees in rkness (Tle 1; Fig. 1). Interestingly, there ws no signifint ifferene etween illumintion tretments t onstnt 25 C or 33/24 C. Temperture n the intertion etween the min effets of temperture n illumintion lso h signifint effet on germintion t week three (Tle 1; Germintion (%) Germintion (%) Germintion week 6 f ef e 25/25 22/11 27/15 29/19 33/24 Temperture (L/D C) ef week 3 e Fig. 1 Effet of temperture n illumintion on germintion fter 3 weeks n 6 weeks., rk;, 12/12 h (light/rk; L/D) photoperio. Error rs represent the stnr error. Within eh grph, rs with the sme letter re not signifintly ifferent t =.5. Fig. 1), with the lowest germintion oserve t simulte winter tempertures (22/11 C). After 6 weeks of ulture, germintion ws not signifintly ffete y illumintion, temperture or the intertion etween these min effets (Tle 1). After 3 weeks of ulture no seelings h proue more thn one lef in the rk tretments, wheres smll perentge of sees ulture in light t 25, 29/19 n 33/24 C proue seon lef ( 1.5%; Fig. 2). At this time the perentge of seelings prouing one n two leves ws signifintly ffete y illumintion (Tle 1), with pronoune evelopmentl lg in the proution of the first lef oserve for rk-trete sees t 22/11 C. First lef proution ws enhne in light with inresing temperture tretments, lthough signifint ifferenes were not etete etween the 25 C n 33/24 C tretments. After 6 weeks of ulture, temperture, illumintion n the intertion etween temperture n illumintion h signifint effets on the perentge of seelings in ll three lef stges. Culturing seelings in the rk t 22/11 C resulte in signifintly more seelings eveloping one lef n signifintly fewer seelings eveloping two leves thn other rk tretments. Among the rk tretments, the gretest perentge of seelings eveloping two leves ws oserve in sees ulture t 33/24 C. No signifint ifferenes in the perentge of seelings eveloping three leves were etete in the rk tretments. Culturing sees in light t 22/11 C resulte in signifintly more seelings eveloping one lef n signifintly fewer eveloping two leves thn other light tretments, s ws oserve mong the rk tretments. Higher osillting tempertures resulte in greter proportion of seelings trnsitioning from the one lef stge to the two n three lef stges. However, the gretest perentge of seelings eveloping three leves ws oserve t onstnt 25 C. Disussion Bleti purpure sees germinte n unerwent lef ifferentition uner wie rnge of temperture regimes in oth rkness n light. Germintion ws slower when sees were ulture t lower tempertures, lthough finl germintion ws not ffete. This initil ely in germintion n evelopment t the lower tempertures ws greter in light thn in rk. The lg ws gretest t the lowest teste temperture regime n my serve n eologil funtion y elying germintion n evelopment if sees re ner the soil surfe when frost is likely. The ility of B. purpure to germinte mximlly over wie rnge of simulte sesonl tempertures my e ue to its persistene in tropil n sutropil origins. In this environment the growing seson is not limite y prolonge sesons of freezing winter tempertures n 11 The Authors Plnt Speies Biology 27, Journl ompiltion 11 The Soiety for the Stuy of Speies Biology

4 TEMPERATURE AND ORCHID SEED GERMINATION 177 Drk Light 1 8 One lef week 6 Two leves Three leves Drk Drk week Perentge in eh evelopmentl stge week 3 week /25 22/11 27/15 29/19 33/24 25/25 22/11 27/15 29/19 33/24 Temperture (y/night C) Temperture (y/night C) Fig. 2 Effet of temperture n illumintion on seeling evelopment fter 3 weeks n 6 weeks of ulture. Error rs represent the stnr error. Within eh grph, rs from the sme evelopmentl lss with the sme letter re not signifintly ifferent t =.5. Tle 1 nov results for the effets of temperture n illumintion on germintion n seeling evelopment Prmeter Effet.f. Week 3 Week 6 F P F P Germintion Illumintion < Temperture < Intertion < First lef stge Illumintion < <.1 Temperture < <.1 Intertion < <.1 Seon lef stge Illumintion < <.1 Temperture < <.1 Intertion < <.1 Thir lef stge Illumintion <.1 Temperture <.1 Intertion <.1 F-vlues in ol re signifint t P =.5 frost risk is typilly fleeting. As the risk of ftlity y freezing is reltively low, it my e tht there is little ptive vntge to relying on temperture ues for germintion, prtiulrly if other ftors re more importnt for suessful germintion. Further exmintion of the roles tht soil wter potentil, light quntity n qulity, n sesonl effets on myorrhizl fungl tivity my ply will eluite when germintion is most Plnt Speies Biology 27, The Authors Journl ompiltion 11 The Soiety for the Stuy of Speies Biology

5 178 T. R. JOHNSON AND M. E. KANE likely to our in nture, llowing mngers to voi ommon mngement prties tht woul hrm elite seelings (i.e. mowing n urning) in orer to promote estlishment of this stte thretene speies. Germintion n evelopment in light n rk ws more rpi t onstnt 25 C ompre with osillting tempertures, exept in light t 33/24 C. This is in ontrst to wht hs een reporte for other orhis (Kuth et l. 11) n mny non-orhis (Thompson & Grimes 1983; Proert et l. 1986), in whih flututing tempertures hve een shown to enhne germintion. However, Proert et l. (1985) foun vriility in the egree to whih Dtylis glomert sees from ifferent regions in Europe respone to flututing tempertures n onlue tht germintion ws enhne y flututing tempertures in essions with stronger initil ormny, wheres germintion of less ormnt essions ws not signifintly ffete y flututing tempertures. Similrly, in multi-region stuy of Clopogon tuerosus vr. tuerosus popultions in the USA, Kuth et l. (11) foun tht popultions tht require longer hilling requirements to reh mximum germintion lso enefite most from flututing temperture regimes. This onlusion inites tht flututing tempertures re more importnt to speies with physiologil ormny. In the sene of physiologil ormny, suh s with B. purpure, flututing tempertures my expose sees to tempertures ove or elow n optiml temperture, resulting in elys to germintion n evelopment. Simulte winter tempertures were foun to ely germintion n evelopment more thn higher tempertures. Bleti purpure popultions t the FPNWR re t the northern extent of their rnge n my e etter pte for wrmer tempertures. Alterntively, elye germintion t the lowest teste temperture my e n pttion to ely germintion uring winter when frosts re possile. The stronger inhiition of germintion n evelopment uner illuminte onitions oul e meite y hnges in the tivity of ifferent temperture-epenent phytohromes (Heshel et l. 7). Delye germintion in light t 27/15 C n 29/19 C oul e regulte similrly. A ol-inue negtive photolsti response my protet sees from germinting n eveloping t the soil surfe when tempertures re low n frost is more likely. Aknowlegments Funing for this projet ws provie y the US Fish n Willife Servie. The uthors wish to thnk Dr. Phil Kuth for helpful omments in the preprtion of the mnusript. Referenes Bskin C. C. & Bskin J. M. (1) Sees: Eology, Biogeogrphy, n Evolution of Dormny n Germintion. Aemi Press, Sn Diego. Bskin C. C. & Bskin J. M. (3) When reking see ormny is prolem: try move-long experiment. Ntive Plnts Journl 4: Benvenuti S., Mhi M. & Miele S. (1) Light, temperture n uril epth effets on Rumex otusifolius see germintion n emergene. Wee Reserh 41: Brown P. M. (5) Wil Orhis of Flori. University Press of Flori, Ginesville. Coile N. C. & Grln M. A. (3) Notes on Flori s enngere n thretene plnts. Flori Deprtment of Agriulture n Consumer Servies, Division of Plnt Inustry, Ginesville. Correll D. S. (1978) Ntive Orhis of North Ameri North of Mexio. Stnfor University Press, Stnfor. Dressler R. L. (1993) Fiel Guie to the Orhis of Cost Ri n Pnm. Comstok Pulishing Assoites, Ith. Dutr D., Johnson T. R., Kuth P. J., Stewrt S. L., Kne M. E. & Rihrson L. (8) Asymioti see germintion, in vitro seeling evelopment, n greenhouse limtiztion of the thretene terrestril orhi Bleti purpure. Plnt Cell, Tissue n Orgn Culture 94: Dutr D., Kne M. E. & Rihrson L. (9) Asymioti see germintion n in vitro seeling evelopment of Cyrtopoium punttum: propgtion protool for n enngere Flori ntive orhi. Plnt Cell, Tissue n Orgn Culture 96: Hn C. Y. & Long C. L. (1) Dormny, germintion n storge of Mgnoli ingrt sees. See Siene n Tehnology 38: Heshel M. S., Sely J., Butler C., Whitelm G. C., Shrrok R. A. & Donohue K. (7) A new role for phytohromes in temperture-epenent germintion. New Phytologist 174: Hilton J. R. (1984) The influene of temperture n moisture sttus on the photoinhiition of see germintion in Bromus sterilis L. y the fr-re soring form of phytohrome. New Phytologist 97: Hollik P., MCom J. A. & Dixon K. W. (7) Introution, growth n persistene in situ of orhi myorrhizl fungi. Austrlin Journl of Botny 55: Hon Y. & Ktoh K. (7) Strit requirement of flututing tempertures s relile gp signl in Piris hierioies vr. jponi see germintion. Plnt Eology 193: Kuth P. J., Kne M. E. & Venrme W. A. (11) Comprtive in vitro germintion eology of Clopogon tuerosus vr. tuerosus (Orhiee) ross its geogrphi rnge. In Vitro Cellulr & Developmentl Biology Plnt 47: Kepzynsk E., Piekn-Grohl J. & Kepzynski J. (6) Hormonl regultion of tomto see germintion t suproptiml temperture. At Physiologie Plntrum 28: Kettenring K. M. & Gltowitsh S. M. (7) Temperture requirements for ormny rek n see germintion vry gretly mong 14 wetln Crex speies. Aquti Botny 87: 9 2. Luzer D., Renut S., St-Arnu M. & Bre D. (7) In vitro symioti germintion, protoorm evelopment, n 11 The Authors Plnt Speies Biology 27, Journl ompiltion 11 The Soiety for the Stuy of Speies Biology

6 TEMPERATURE AND ORCHID SEED GERMINATION 179 plntlet limtiztion of Apletrum hyemle (Muhl. ex Will.) Torr. (Orhiee). Journl of the Torrey Botnil Soiety 134: Leon R. G. & Owen M. D. K. (3) Regultion of wee see ormny through light n temperture intertions. Wee Siene 51: Msuhr G. & Ktsuy K. (1994) In situ n in vitro speifiity etween Rhizotoni spp. n Spirnthes sinensis (Persoon) Ames. vr. moen (M. Bieerstein) Hr (Orhiee). New Phytologist 127: MKenrik S. L., Leke J. R., Tylor D. L. & Re D. J. () Symioti germintion n evelopment of myoheterotrophi plnts in nture: ontogeny of Corllorhiz trifi n hrteriztion of its myorrhizl fungi. New Phytologist 145: MKenrik S. L., Leke J. R., Tylor D. L. & Re D. J. (2) Symioti germintion n evelopment of the myoheterotrophi orhi Neotti nius-vis in nture n its requirement for lolly istriute Sein spp. New Phytologist 154: Miyoshi K. & Mii M. (1998) Stimultory effets of soium n lium hypohlorite, pre-hilling n ytokinins on the germintion of Cypripeium mrnthos see in vitro. Physiologi Plntrum 12: Moyo M., Kulkrni M. G., Finnie J. F. & vn Sten J. (9) After-ripening, light onitions, n ol strtifition influene germintion of Mrul [Slerory irre (A. Rih.) Hohst. susp. ffr (Son.) Kokwro] sees. HortSiene 44: Nsimento W. M., Cntliffe D. J. & Huer D. J. () Thermotolerne in lettue sees: ssoition with ethylene n eno--mnnnse. Journl of the Amerin Soiety for Hortiulturl Siene 125: Norsworthy J. K. & Oliveir M. J. (6) Siklepo (Senn otusifoli) germintion n emergene s ffete y environmentl ftors n seeing epth. Wee Siene 54: Plestin R. A. & Sos V. (2) Morphologil vrition in popultions of Bleti purpure (Orhiee) n esription of the new speies B. ripri. Brittoni 54: Peroz-Mnrique J. & Min-Gutierrez Y. (6) Asymioti germintion of Oontoglossum gloriosum Rh.F. (Orhiee) uner in vitro onitions. In Vitro Cellulr & Developmentl Biology Plnt 42: Proert R. J., Smith R. D. & Birh P. (1985) Germintion responses to light n lternting tempertures in Europen popultions of Dtylis glomert L. I. Vriility in reltion to origin. New Phytologist 99: Proert R. J., Smith R. D. & Birh P. (1986) Germintion responses to light n lternting tempertures in Europen popultions of Dtylis glomert L. V. The priniple omponents of the lternting temperture requirement. New Phytologist 12: Rsmussen H. N. (1992) See ormny ptterns in Epiptis plustris (Orhiee), requirements for germintion n estlishment of myorrhiz. Physiologi Plntrum 86: SAS Institute (2) SAS E. SAS Institute, Cry. Shimur H. & Ko Y. (5) Enhne symioti see germintion of Cypripeium mrnthos vr. reunense following inoultion fter ol tretment. Physiologi Plntrum 123: Sos V. & Díz-Dums M. (1997) Orhis from the Greter Antilles I: new speies of Bleti. Brittoni 49: Stewrt S. L. & Kne M. E. (6) Asymioti see germintion n in vitro seeling evelopment of Henri mroertitis (Orhiee), rre Flori terrestril orhi. Plnt Cell, Tissue n Orgn Culture 86: Stewrt S. L. & Kne M. E. (1) Effet of rohyrte soure on the in vitro symioti see germintion of terrestril orhi Henri mroertitis. Journl of Plnt Nutrition 33: Thompson K. & Grimes J. P. (1983) A omprtive stuy of germintion responses to iurnlly-flututing tempertures. Journl of Applie Eology : Turner S. R., Merritt D. J., Riley E. C., Commner L. E., Bskin J. M., Bskin C. C. & Dixon K. W. (6) Eophysiology of see ormny in the Austrlin enemi speies Anthorpus preissii (Dsypogonee). Annls of Botny 98: Vin E. F. & Went F. W. (1949) Some ph hnges in nutrient solutions. Botnil Gzette 11: Whighm D. F., O Neill J. P., Rsmussen H. N., Clwell B. A. & MCormik M. K. (6) See longevity in terrestril orhis Potentil for persistent in situ see nks. Biologil Conservtion 129: Willims L. O. & Allen P. H. (1998) Orhis of Pnm. Missouri Botnil Gren Press, St Louis. Ymzki J. & Miyoshi K. (6) In vitro symioti germintion of immture see n formtion of protoorm y Cephlnther flt (Orhiee). Annls of Botny 98: Plnt Speies Biology 27, The Authors Journl ompiltion 11 The Soiety for the Stuy of Speies Biology

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