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Month Aug. Sept. Day 26 28 2 4 9 11 16 18 23 Topic Introduction The ecosystem concept Climate/soils Soils II Ecosystem energy balance Water cycling Carbon GPP/NPP NEP C,M&M 1 2 3 4 4 5 6 6 Atmospheric Carbon Dioxide Tons of CO2-C emitted per person per year Anthropogenic C Emissions: Land Use Change Carbon Emissions from Tropical Deforestation 2000-2006 1.5 Pg C y-1 1.60 Africa 1.40 Latin America 1.20 S. & SE Asia 1.00 SUM (16% total emissions) 0.80 0.60 0.40 0.20 Houghton, unpublished 2000 1990 1980 1970 1960 1950 1940 1930 1920 1910 1900 1890 1880 1870 1860 0.00 1850 Pg C yr-1 1.80 Carbon Biosphere has large effect on atmospheric CO2 concentrations Photosynthesis is the primary energy base for all terrestrial ecosystems Net Primary Productivity (NPP) = Gross Primary Productivity (GPP) – RespirationPlant Plant growth = Photosynthesis - Respiration Ecosystem Carbon Cycling Net Ecosystem Production (NEP) = GPP - REcosys Recosys= Respiration of plants, animals, and soil microbes, plus other C transfers from plants Positive NEP; GPP > Recosys; ecosystem is removing C from the atmosphere = C sink Negative NEP; GPP < Recosys; ecosystem is releasing C to the atmosphere = C source Political Implications of Ecosystem Carbon Cycling Kyoto Protocol to the UN Framework Convention on Climate Change: international treaty on greenhouse gas emissions (incld. CO2) Negotiated 1997 Came into force 2004 (Russia and the “55% clause”) Now 161 counties, ~61% emissions EU (Annex 1): ratified protocol, submit annual gas inventory; reduce emissions to 5% below 1990, effectively 15% below projected rates for 2008 Developing countries (Non-annex 1) can choose to reduce emissions and sell reductions to Annex 1 as carbon credits (“cap and trade”) Review of C3 Photosynthesis In the chloroplast, light reactions transform light energy into chemical energy Photosynthetically active radiation (PAR) absorbed by chlorophyll (APAR) between 400-700 nm Electrons derived from oxidation of H2O to O2 used to make high-energy ATP, NADPH Review of C3 Photosynthesis Dark reaction use chemical energy (ATP, NADPH) to fuel carbon fixation reaction (Calvin cycle). CO2 is used to carboxylate ribulose bisphosphate (RuBP) with the enzyme ribulose bisphosphate carboxylase/oxygenase (Rubisco) Rubisco accounts for ≈ 25% of leaf Nitrogen Other Ps enzymes account for ≈ 25% of leaf N Carboxylation of RuBP produces 2, 3-C sugars Review of C3 Photosynthesis Rubisco can also oxygenate RuBP, ultimately losing CO2 through photorespiration 20-40% of energy from Ps immediately lost via photorespiration Depends on relative concentrations of O2 and CO2 in leaf, temperature Net Ps: difference between tot. Ps and photorespiration=GPP C4, CAM use same biochemistry, but different timing and morphology to affect O2/CO2 ratio in leaf Limitation Plants adjust components of Ps so physical and biochemical processes co-limit C fixation Plants adjust resource acquisition to maximize capture of the most limiting resource • CO2 • Water Acclimation versus adaptation! • Light • Nitrogen • [Temperature-modulator] CO2 response curve of photosynthesis • Diffusion limitation affected by stomata • Biochemical limitation affected by light/enzymes • Plants equalize physical and biochemical limitations Inherent tradeoff between CO2 gain and H2O loss Water availability • Limits full use of light in arid conditions due to tradeoff with CO2 • Water Use Efficiency (WUE) Lo H2O ecosystems Lo stomatal conductance Lo leaf area, deciduousness Steep leaf angels Reflective leaves Small leaves C4, CAM Deep roots Light response curve of photosynthesis • Light limitation slope (quantum yield) constant for C3 • Light saturation limited by investment in enzymes (N) • Photoinhibition results from damaged membranes Light variation in ecosystems Lecture ended here Light environment • …for individual leaves, it is determined by total leaf area above • Penetration of light into the canopy depends on leaf angles and distribution in space • Acclimation and adaptation to light environment • Light Use Efficiency (LUE) Leaf nitrogen determines Ps capacity Rubisco accounts for ≈ 25% of leaf Nitrogen Other Ps enzymes account for ≈ 25% of leaf N Nitrogen availability • Inherent tradeoff between traits that favor high Ps and traits that favor retention of N • Easier for a plant to retain N then to acquire from the environment Low N ecosystem Increased leaf toughness Increased structural C Thick leaves (high weight/area) Increased leaf life span Increased defense (herbivory) Low Ps (per unit weight) High N ecosystem Decreased leaf toughness Decreased structural C Thin leaves (low weight/area) Decreased leaf life span High Ps (per unit weight) Temperature extremes • Plants minimize effects of lo T by increasing Ps capacity (increasing leaf N) • Minimize effects of hi T by • hi stomatal conductance (Hi H2O) • dissected, reflective, deciduous leaves (Lo H2O)