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Universität Bayreuth BayCEER – Labor für Isotopen-Biogeochemie SS 2007 Modul: Stabile Isotope in der Biogeochemie Vorlesung Stabile Isotope in der Ökologie Lehrperson: Gerhard Gebauer Termin: Do. 8 - 10 Uhr Raum: S 37. Stabile Isotope in der Ökologie.
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Universität Bayreuth BayCEER – Labor für Isotopen-Biogeochemie SS 2007 Modul: Stabile Isotope in der Biogeochemie Vorlesung Stabile Isotope in der Ökologie Lehrperson: Gerhard Gebauer Termin: Do. 8 - 10 Uhr Raum: S 37
Stabile Isotope in der Ökologie • Inhalt der Vorlesung (II): • Stickstoffisotope: • N-Quellen für Pflanzen (substanzspezifische und räumliche Variation im • Isotopenverhältnis, Wirkung anthropogener Einflüsse) • Aufnahme bodenbürtiger N-Quellen (Ammonium, Nitrat, organ. N) • Rolle der Pilze im N-Kreislauf • mykoheterotrophe Pflanzen • Aufnahme atmosphärischer N-Depositionen und Wirkung auf den N-Haushalt • Nitrataustrag aus Ökosystemen • Lachgasproduktion und -konsumption • Symbiontische N2-Fixierung • N-Ernährung insektenfangender Pflanzen • Isotopenverschiebung bei Nahrungsketten
Mykorrhizaformen und beteiligte Symbiosepartner St = gebündelte Myzelstränge APH = äussere Pilzhülle HN = Hartig´sches Netz IN = Interzelluläres Mycelnetz IK = Intrazelluläres Mycelknäuel V = Pilzvesikel A = Arbuskel SP = Spore Nach Gianinazzi und Gianinazzi-Pearson (1988) aus: Larcher: Ökophysiologie der Pflanzen, 6. Auflage, 2001
From: Schulze, Chapin and Gebauer (1994) Oecologia 100: 406-412
From: Michelsen, Quarmby, Sleep and Jonasson (1998) Oecologia 115: 406-418
Frequencies From: Gebauer and Dietrich (1993) Isotopenpraxis Environ. Health Stud. 29: 35-44
Fichtelgebirge Sept. 1989 Frequencies From: Gebauer and Dietrich (1993) Isotopenpraxis Environ. Health Stud. 29: 35-44
From: Schmidt and Gleixner (1997) In: Griffiths (ed) Stable Isotopes: Integration of Biological, Ecological and Geochemical Processes
Russula ochroleuca Laccaria laccata Cystoderma amianthinum Trametes versicolor + Calocera viscosa
Strobilurus esculentus Micromphale perforans
Relation between d15 N values of fungal fruit bodies and of the substrates for their mycelium
Partial mycoheterotrophy – A novel nutritional mode of forest ground vegetation
The forest ground: a light-limited site for plant growth • Light availability in Central European forests at the ground related to the insolation above the canopy: • Pine forest ca. 20 % • Oak forest ca. 10 % • Beech forest ca. 1 – 5 %
Ectomycorrhiza of forest trees Fungal hyphae Fungal partners Hardly cultivable basidiomycetae and ascomycetae, (obligatory ectomycorrhizal fungi) Fungal partners Easily cultivable, saprotrophic basidiomycetae of the Rhizoctonia group Fungal hyphae Endomycorrhiza of orchids
Benefit of the mycorrhiza for plants and fungi Ectomycorrhiza of forest trees CO2 Photosynthesis Carbohydrates ECM fungi Tree Mineral nutrients Water Exoenzymes N assimilation Soil Nitrate, ammonium CO2 Endomycorrhiza of green orchids Photosynthesis ? ? ? Green orchid Saprotrophic fungi Mineral nutrients Water (Carbohydrates) N assimilation Exoenzymes Soil Nitrate, ammonium
Monotropa hypopitys Examples of mycoheterotrophic plants from European forests Neottia nidus-avis
Relative enrichment in heavy isotopes (eMH-R) of mycoheterotrophic orchids (MHO), monotropoids (MHM) and pyroloids (MHP)
Are green orchids really autotrophic? Cephalanthera damasonium Cephalanthera rubra
Are green orchids really autotrophic? Epipactis atrorubens Platanthera chlorantha
Data from: Gebauer & Meyer (2003) New Phytol. 160: 209-223 Bidartondo, Burghardt, Gebauer, Bruns & Read (2004) Proc. R. Soc. Lond. B 271: 1799-1806 Non-orchids with ectomycorrhiza Non-orchids with arbuscular mycorrhiza Green orchids(group 1) Green orchids (group 2) Mycoheterotrophic orchids
Data from: Bidartondo, Burghardt, Gebauer, Bruns & Read (2004) Proc. R. Soc. Lond. B 271: 1799-1806
CO2 Photosynthesis Organ. C Organ. N Mineral nutrients Water Mineral nutrients Water N assimilation Nitrate ammonium Transition from autotrophic to mycoheterotrophic nutrition CO2 Photosynthesis ECM fungus A Carbohydrates Saprotrophic fungi ECM fungus B Orchids Trees Mineral nutrients Water Exoenzymes ECM fungus C Soil N assimilation Exoenzymes Nitrate ammonium Soil
Cephalanthera damasonium Albino varieties among green orchids Albino variety From: Julou, Burghardt, Gebauer, Berveiller, Damesin & Selosse (2005) New Phytol. 166: 639-653 Green individuum
Non-orchids with ectomycorrhiza Non-orchids with arbuscular mycorrhiza Green orchids with ectomycorrhiza Albino variety of green orchids Ectomycorrhizal fungi From: Julou, Burghardt, Gebauer, Berveiller, Damesin & Selosse (2005) New Phytol. 166: 639-653
Are green pyroloids also partially mycoheterotrophic? Chimaphila umbellata Orthilia secunda
Data from: Zimmer, Hynson, Gebauer, Allen, Allen & Read (2007) New Phytol. 175: 166-175 Forest sites with different light climate (NE Bavaria, Germany) Reference plants with ectomycorrhiza Reference plants with arbuscular mycorrhiza Green Pyroleae Mycoheterotrophic orchids or Pyroleae
Source B Source A Contribution of source B to plant total C or N [%] Quantifying mycoheterotrophic nutrition
Data from: Gebauer (2005) Rundgespräche der Kommission für Ökologie, Vol. 30, pp 55-67 Zimmer, Hynson, Gebauer, Allen, Allen & Read (2007) New Phytol. 175: 166-175
Conclusions • Partial mycoheterotrophy is a hitherto unknown nutritional • mode of putatively autotrophic forest ground vegetation. • Partial mycoheterotrophy occurs at least in two important • plant families. • Partial mycoheterotrophy is associated with a switch of • fungal partners, that provides a link between ground • vegetation, ectomycorrhizal fungi and forest trees. • C gain from partial mycoheterotrophy is related to plant • species and to light climate. It allows the plants to move • into the deepest shade of forests without competition. • Combining stable isotope data with molecular biological • techniques provides essentially new perspectives in • research on mycorrhizas.