C02-Opname Bij CAM Planten Bromelia's, Phalaenopsis, Kalanchoe En Andere

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C02-Opname Bij CAM Planten Bromelia's, Phalaenopsis, Kalanchoe En Andere PRAKTIJ KDNDERZDEK PLANT & DMGEVING C02-opname bijCA Mplante n Bromelia's, Phalaenopsis, Kalanchoe enander e Literatuurstudie M.G.Warmenhove n Tj. Blacquière Praktijkonderzoek Plant &Omgevin g B.V. Sector Glastuinbouw September 2001 Publicatienummer 255 WAB E N I N G E N r Inhoudsopgave pagina 1 SAMENVATTING 5 2 INLEIDING 7 3 METHODE 9 4 CRASSULACEANACI DMETABOLIS M(CAM ) 11 4.1 WATi sCAM ? 11 4.1.1 Welke vormen vanfotosynthes e zijner ? 11 4.1.2 CAM - fotosynthese nader bekeken 13 4.2 INVLOEDOMGEVINGSFACTORE N 16 4.2.1 C02 16 4.2.2 Temperatuur 18 4.2.3 Licht 19 4.2.4 Water(stress)/ zoutstress 20 4.2.5 Diverse invloeden 21 4.3 WELKEPLANTE NZIJ NCAM ? 22 4.4 WELKE METHODENZIJ NE RO MNIEUW ESOORTE NE NCULTIVAR ST ESCREENE NO PEVENTUEL ECA M - FOTOSYNTHESE 23 5 DISCUSSIE 25 6 LITERATUUR " 27 1 Samenvatting Indi t rapport wordt ingegaan op devolgend e vragen: 1)wa t isCA M2 )welk e soorten/cultivars zijnCA Me n 31ho eku nj e hetCA Mmechanism e aantonen.Voo r deopnam eva nC0 2 (viad e huidmondjes) zijni nhe t plantenrijk een drietal mechanisme aanwezig,C 3 -,C 4 -e nCA M-fotosynthese . BijC 3-fotosyntheseword t C02 door Rubisco direct aanee nC 5suikergebonde nwaarn a C3suikersworde n gevormd. Rubisco bindt echter ook vaak- per ongeluk - zuurstof (fotorespiratie) waardoor energie verloren gaat. Inwarmer e klimaten zal defotorespirati e toenemen eni s het dus belangrijk dat de opname van C02 wordt aangepast. Door het C02 specifieke enzym PEP-carboxylaseword t geen zuurstof gebonden.C 4-fotosynthesebind t C02 met PEP-carboxylase waarna hetvi a eentransportmolecuu l (meestal malaat) naar cellen vand e vaatbundelschede wordt getransporteerd. Hierword t deC0 2 weer afgesplitst omverde r via hetC 3 mechanisme (Rubisco) suiker te vormen.Voo r defotosynthes e istoetredin g van C02 in hetweefse l noodzakelijk. Daartoe heeft de plant huidmondjes. Echter door de huidmondjes verliest de planttegelij k veelwate r (verdamping). Dit dilemma wordt nog urgenter bijwate r tekort. Eenoplossin gvoo r dit dilemma is het CAM: C02 wordt door dehuidmondje s op eenveili g moment opgenomen: ind e nacht. Dani s hetkoe l endonker , waardoor het water verlies gering is. Overdag zijn dehuidmondje s dan dicht. Evenals bij de C4- fotosynthese bindt, bij CAM- fotosynthese , het enzym PEP-carboxylase de C02. Hetmalaa t wordt dannaa r devacuol e getransporteerd engedurend e denach t opgeslagen. Overdagword t het malaat uit devacuol e getransporteerd enC0 2 afgesplitst omverde r het C3-mechanismete volgen- daarbij is licht nodig.D e fixatie van C02 is dus bij C4-fotosyntheseruimtelij k gescheiden enbi j CAM -fotosynthes e in detij d gescheiden. CAM-cyclinge nCA M -idlin g zijnvariante nva n CAM -fotosynthes e waarbij C02 wordt hergebruikt. Daarnaast zijn erfacultatiev e CAM- planten ,afhankelij k vand eomstandighede n vertonen deze C3-,C 4- of CAM - fotosynthese. Wanneer deC0 2 concentratie wordtverhoog d kanoo k bij sommige CAM- plante nd e C02 opname toenemen (vaak meer biomassa). Deextr a opname wordt vooral gerealiseerd bij de overgangva ndonke r naar licht enlich t naar donker waar deCA M- fotosynthes e wordt overgenomen door C3-fotosyntheseo f omgekeerd. Hetverhoge n vand e C02 concentratie kanhe ttijdsti p waarop de huidmondjes opengaa n vervroegen. Optimale CAM- fotosynthes e vindt plaats bij eenwarm e dagtemperatuur enee nkoel e nachttemperatuur. Extreme dagtemperaturen zullen de nachtelijke C02 opname sterk reduceren. Hetverhoge nva nd elichtintensitei t kanresultere n inee nafnam e vand e C02 opname.Verde r verhogenva n de lichtintensiteit zal de opname overdag stoppen. Droogtestress geeft op denduu r dezelfde effecten. Welkfotosynthesepa d wordt gebruikt door de plant is sterk afhankelijk van de beschikbaarheid vanwater , zon of schaduw end e temperatuur. Perfamili e is aangegevenwelk e plantenvolgen s deliteratuu r CAM -o f C3-fotosynthese bedrijven. Het bepalen vand e malaat concentratie of totaal titreerbaar zuur ind e plant zijnnaas t hetvolge n vand e C02 uitwisseling gedurende de dag demees t simpele methode omCA M- fotosynthes e vastte stellen. Inleiding CAMplante n nemennie t overdag maar "s nachts C02 op. Dit mechanisme komt o.a. voor bij Bromelia's, Phalaenopsis enKalancho ë Plantenzij nechte r soms niet obligaat CAM-plant,o f kunnenva n systeem wisselen afhankelijk van omstandigheden,e n insommig e families of geslachten is de ene soort wel CAM- plant, maar eenander e niet. Toegespitst per soort liggen er devolgend evragen : Bromelia's: -welk e soorten/cultivars zijn CAM,e ni nhoeverr e -ho e kan dat eenvoudig gescreend worden,oo k voor nieuwe soorten encultivar s Phalaenopsis: -i s Phalaenopsisstrik tCAM ? Doel: Het doelva n deze literatuurstudie omaa nd e handva nverschene n publicaties inzicht te krijgenin : Wati sCA M Eeninventarisati e vanwelk e soorten/cultivars zijnCA M Methode om CAMmechanism e aant e tonen Daarnaast zalgekeke n worden of op devolgend e vragen een antwoord kanworde ngegeven : Hoeveel licht overdag isvoldoend e om de C02 opnamecapaciteit 's nachtste benutten,e n omgekeerd? Hoe lang magd e dagworde nverleng d met bijv. assimilatiebelichting zonder dat de nachtelijke C02 opname ind ekne lkomt ? Wat stuurt de huidmondjesopening: dag/nacht ritme of gaanz e alvoo r donker weer open? Methode Voor de litteratuurstudie zijnd evolgend e bestandengeraadpleegd : • Agralin (Agrarisch Literatuur Informatiesysteem Nederland) • C.A.B. (Commonwealth Agricultural Bureaux) Voor het zoeken zijn detrefwoorde n C02, CAMe nd e combinatie C02 * CAMgebuikt . Vervolgens isd e geschikte literatuur gescreend enopgevraagd . Met behulpva nwa t hoofdstukken uit leerboeken enee n monografie over CAMi s daarna eenraamwer k gemaakt waarin degeleze n literatuur wordt gepast: • CAMalgemeen :wa t is het, hoewerk t het. • Welke omgevingsfactoren hebben invloed op het al dannie t gebruiken vanhe t CAMpa ddoo r bepaalde planten. • Welke soorten uit het sierteeltsortiment zijn(mogelijk )CAM-plant . • Welke methoden zijn er, liefst eenvoudige, om nieuwe soorten encultivar s te screenen op eventuele CAM-fotosynthese Crassulacean Acid Metabolism (CAM) 4.1 Wati s CAM? 4.1.1 Welkevorme n vanfotosynthes e zijner ? Het omzetten vanenergi e uit zonlicht naar chemische energie is voornamelijk voorbehouden aangroen e planten. Door deopnam e vankoolzuurga s enwate r worden m.b.v.va nd e zonne-energie suiker enzuursto f gevormd. Deoverallreacti e van defotosynthes e isal s volgt: koolzuurgas + water + energie =>suike r + zuurstof Deabsorpti e vanzonne-energi e vindt plaats ind e pigmenten. Bijhoger e planten gaat het hier om chlorofyllen encarotenoïden . Door deabsorpti e vanlich t komt een ketenva nelektronentranspor t opgang. Tijdens het elektronentransport worden energierijke verbindingen gevormd alsAT Pe n NADPH. Hetfotosynthes e proces kanworde n onderverdeeld intwe e type reacties: 1) Processen die lichtafhankelijke zijn(lichtreacties ) zoals het splitsen vanwate r end evormin gva n energierijke verbindingen ATPe n NADPH. 2) Processen die niet lichtafhankelijk zijn(donkerreacties ) zoals de reductie van C02 end evormin gva n suikers. Hiervoor worden de energierijke verbindingen ATPe n NADPHgebruikt . Koolzuurgas (C02)word t via dehuidmondje s ingevangen.D emanie r waarop C02word t opgenomene n verder verwerkt tot suikers via drie verschillende padenverlopen ; C3; C4-,e nCA M- fotosynthese .D e belangrijkste factoren die kan bepalenwel k van depade ngebruik t wordt is de beschikbaarheid vanwate r enC0 2. C3-fotosynthese InC 3-fotosyntheseword t C02 ind e mesofylcellen (bevinden zich direct achter de huidmondjes) door het enzym Rubisco gebonden aanC 5-suike r (Ribulose 1,5- bisphosphate ) tot een instabiele C6verbindin gdi e direct uiteen valt intwe e C3stukje s (PGA- 3-Phosphoglycerate)waarui t het C3-suike r Triose wordt gevormd. Rubisco bindt ook vaak- per ongeluk - zuurstof (fotorespiratie). Door devee l hogere zuurstof concentratie (20%)da n koolzuurgas (0,035%) komt dit nogal eensvoor . Het bindenva nzuursto f resulteert nietto t de inbouw vankoolsto f engeef t dus energieverlies. C3-fotosynthese , opnamee n verwerking,vind t overdag plaats. Met name inee nklimaa t met hoog licht, waar deverhoudin g CCyC^ noglage r is,za l fotorespiratie toenemen. Verder zijn inee n klimaat met hoog licht de belangenva n defotosynthes e (huidmondjes open voor een goede uitwisseling vanC0 2 met deomgeving ) tegenstrijdig met die vand e waterbalans (huidmondjes dicht om uitdroging te voorkomen). Omdez e problemen opte vangen hebben plantenee n tweetal mechanismen ontwikkeld: C4-fotosynthese Doel:fotorespirati e vermijden. InC 4-fotosynthes e wordt C02 ind emesofylcelle n door het enzym PEP-carboxylase gebonden aanee n drager molecuul (fosfo-enolpyruvaat- PEP )waarn a malaat of asparaat (C4-zuren)ontstaat . Het C4-molecuul malaat/asparaa t wordt getransporteerd naar devaatbundelsched e (een krans met cellen rondd e vaatbundels waarin Rubisco zit). Ind evaatbundelsched e wordt de C02 weer afgesplitst enn ugefixeer d met behulp van Rubiscoal s bij C3-fotosynthese . Daar PEP-carboxylase C02 specifiek isword t er geenzuursto f ingebouwd. Deplaat s vanfixati e van C02 isdu s ruimtelijk gescheiden vand e plaatsva nverwerking . Opd e plaats vanverwerkin g wordt de C02 -concentratie kunstmatig hooggehouden . 11 Donker Licht Atmospheric C0 opname Malaatgeef t C0 af. Opnieuw 2 2 Huidmondjes enfixati e fixatie m.b.v. Rubisco dicht F/guur1 - Crassalusean acid metabolism inbla d(Tai z et. all, 1998. (98)) Tabel1 Algemene karakteristieken Cr C4-,e nCA M- fotosynthese (77, 93, 98) Kenmerk
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