Aluminum toxicity in a tropical montane forest ecosystem in southern Ecuador Inauguraldissertation der Philosophisch-naturwissenschaftlichen Fakult¨at der Universit¨at Bern vorgelegt von Agnes Rehmus von Deutschland Leiter der Arbeit: Prof. Dr. W. Wilcke Geographisches Institut, Universit¨at Bern Original document saved on the web server of the University Library of Berne. This work is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License. To see the license go to http://creativecommons.org/licenses/by-nc-nd/4.0/ or write to Creative Commons, PO Box 1866, Mountain View, CA 94042, USA. Copyright Notice This work is licensed under a Creative Commons Attribution-NonCommercial- NoDerivatives 4.0 International License. To see the license go to http://creativecommons.org/licenses/by-nc-nd/4.0/. You are free: Share – copy and redistribute the material in any medium or format. 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Aluminum toxicity in a tropical montane forest ecosystem in southern Ecuador Inauguraldissertation der Philosophisch-naturwissenschaftlichen Fakult¨at der Universit¨at Bern vorgelegt von Agnes Rehmus von Deutschland Leiter der Arbeit: Prof. Dr. W. Wilcke Geographisches Institut, Universit¨at Bern Von der Philosophisch-naturwissenschaftlichen Fakult¨at angenommen. Bern, 23.02.2015 Der Dekan: Prof. Dr. G. Colangelo In Andenken an meinen Vater. Contents Contents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I Danksagung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . VI Zusammenfassung . . . . . . . . . . . . . . . . . . . . . . . . . . . . VII List of figures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . XII List of tables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . XVI List of abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . XVII 1 Summarizing overview 1 1.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1.2 Materials and Methods . . . . . . . . . . . . . . . . . . . . . . 6 1.2.1 The hydroponic experiment . . . . . . . . . . . . . . . 6 1.2.2 Aluminum in the south Ecuadorian montane forest ecosys- tem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 1.3 Results and discussion . . . . . . . . . . . . . . . . . . . . . . 18 1.3.1 Aluminumtoxicitytotropicalmontaneforesttreeseedlings in southern Ecuador: response of biomass and plant mor- phology to elevated Al concentrations (Chapter 2) . . . 18 1.3.2 Aluminumtoxicitytotropicalmontaneforesttreeseedlings insouthernEcuador:responseofnutrientstatustoelevated Al concentrations (Chapter 3) . . . . . . . . . . . . . . 19 1.3.3 Response of Al concentrations and speciation in soil so- lution to Ca amendment in a tropical montane forest in southern Ecuador (Chapter 4) . . . . . . . . . . . . . . 20 I 1.3.4 Aluminum cycling in a tropical montane forest ecosystem in southern Ecuador (Chapter 5) . . . . . . . . . . . . 20 1.3.5 Can we expect Al toxicity in the tropical montane forests of south Ecuador? . . . . . . . . . . . . . . . . . . . . . 21 1.3.6 Error discussion . . . . . . . . . . . . . . . . . . . . . . 25 1.4 General conclusions . . . . . . . . . . . . . . . . . . . . . . . . 29 1.5 Author contributions . . . . . . . . . . . . . . . . . . . . . . . 30 1.6 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 2 Aluminum toxicity to tropical montane forest tree seedlings in southern Ecuador: response of biomass and plant morphology to elevated Al concentrations 39 2.1 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 2.2 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 2.3 Materials & Methods . . . . . . . . . . . . . . . . . . . . . . . 43 2.3.1 Experimental Design . . . . . . . . . . . . . . . . . . . 43 2.3.2 Chemical analyses . . . . . . . . . . . . . . . . . . . . 45 2.3.3 Statistical analyses . . . . . . . . . . . . . . . . . . . . 46 2.4 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 2.4.1 Principal component analysis of plant properties . . . . 47 2.4.2 Dose-response curves . . . . . . . . . . . . . . . . . . . 52 2.4.3 Chemical plant properties . . . . . . . . . . . . . . . . 54 2.4.4 Organic layer leachate experiment . . . . . . . . . . . . 56 2.5 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58 2.5.1 Aluminum effects on plant biomass and morphology . . 58 2.5.2 Sensitivity to Al exposure . . . . . . . . . . . . . . . . 61 2.5.3 Hoagland nutrient solution versus organic layer leachate 62 2.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 2.7 Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . 63 2.8 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 II 3 Aluminum toxicity to tropical montane forest tree seedlings in southern Ecuador: response of nutrient status to elevated Al concentrations 69 3.1 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 3.2 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 3.3 Materials & Methods . . . . . . . . . . . . . . . . . . . . . . . 72 3.3.1 Experimental Design . . . . . . . . . . . . . . . . . . . 72 3.3.2 Chemical analyses . . . . . . . . . . . . . . . . . . . . 74 3.3.3 Statistical analyses . . . . . . . . . . . . . . . . . . . . 74 3.4 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 3.4.1 Concentrations of macronutrients in plant tissue . . . . 75 3.4.2 Concentrations of micronutrients, beneficial mineral ele- ments, and Al in plant tissue . . . . . . . . . . . . . . 77 3.4.3 Relationship between nutrient status and biomass produc- tion . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79 3.5 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81 3.5.1 Reduced shoot biomass . . . . . . . . . . . . . . . . . . 81 3.5.2 Stimulated root growth at 300 µM Al . . . . . . . . . . 84 3.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85 3.7 Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . 86 3.8 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 4 Response of Al concentrations and speciation in soil solution to Ca amendment in a tropical montane forest in southern Ecuador 91 4.1 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 4.2 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 4.3 Materials & Methods . . . . . . . . . . . . . . . . . . . . . . . 95 4.3.1 Study site and sampling procedures . . . . . . . . . . . 95 4.3.2 Chemical analyses . . . . . . . . . . . . . . . . . . . . 96 4.3.3 Donnan membrane technique . . . . . . . . . . . . . . 97 4.3.4 Speciation modeling and statistics . . . . . . . . . . . . 98 4.4 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 III 4.5 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105 4.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108 4.7 Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . 109 4.8 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 5 Aluminum cycling in a tropical montane forest ecosystem in southern Ecuador 115 5.1 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 5.2 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 5.3 Materials & Methods . . . . . . . . . . . . . . . . . . . . . . . 118 5.3.1 Study site and sampling procedures . . . . . . . . . . . 118 5.3.2 Chemical analyses . . . . . . . . . . . . . . . . . . . . 121 5.3.3 Calculations and statistical evaluation . . . . . . . . . 122 5.4 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123 5.4.1 Al concentrations and fluxes . . . . . . . . . . . . . . . 123 5.4.2 Drivers of Al concentrations . . . . . . . . . . . . . . . 126 5.4.3 Toxicity indicators . . . . . . . . . . . . . . . . . . . . 129 5.5 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130 5.5.1 Al fluxes . . . . . . . . . . . . . . . . . . . . . . . . . . 130 5.5.2 Drivers of Al concentrations . . . . . . . . . . . . . . . 135 5.5.3 Toxicity indicators . . . . . . . . . . . . . . . . . . . . 136 5.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138 5.7 Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . 139 5.8 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139 A Appendix 147 Erkl¨arung 150 IV
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