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1. Objectives This project combines automated in situ observations of the isotopologues of CO2 with root observations, novel experimental manipulations of belowground processes, and isotope-enabled ecosystem modeling to investigate mechanisms of below- vs. aboveground carbon sequestration at the Harvard Forest Environmental Measurements Site (EMS). The proposed objectives, which have now been largely accomplished, include: A. Partitioning of net ecosystem CO2 exchange (NEE) into photosynthesis and respiration using long-term continuous observations of the isotopic composition of NEE, and analysis of their dynamics ; B. Investigation of the influence of vegetation phenology on the timing and magnitude of carbon allocated belowground using measurements of root growth and indices of belowground autotrophic vs. heterotrophic respiration (via trenched plots and isotope measurements); C. Testing whether plant allocation of carbon belowground stimulates the microbial decomposition of soil organic matter, using in situ rhizosphere simulation experiments wherein realistic quantities of artificial isotopically-labeled exudates are released into the soil; and D. Synthesis and interpretation of the above data using the Ecosystem Demography Model 2 (ED2). 2. Highlights Accomplishments: • Our isotopic eddy flux record has completed its 5th full year and has been used to independently estimate ecosystem-scale respiration and photosynthesis. • Soil surface chamber isotopic fluxmore » measurements were carried out during three growing seasons, in conjunction with a trenching manipulation. Key findings to date (listed by objective): A. Partitioning of Net Ecosystem Exchange: 1. Ecosystem respiration is lower during the day than at night—the first robust evidence of the inhibition of leaf respiration by light (the “Kok effect”) at the ecosystem scale. 2. Because it neglects the Kok effect, the standard NEE partitioning approach overestimates ecosystem photosynthesis (by ~25%) and daytime respiration (by ~100%) in the first half of the growing season at our site, and portrays ecosystem photosynthetic light-use efficiency as declining when in fact it is stable until autumnal senescence. B. Vegetation Phenology and belowground allocation: Findings: 1. Autotrophic respiration (Ra) showed a seasonal pattern, peaking in mid-summer when trees were most active. 2. The effective age of the substrate for belowground respiration is less than 2 weeks. 3. Above and belowground phenology are more synchronous in deciduous hardwood stands than evergreen hemlock stands. 4. The decline in root respiration rates in the fall is related to temperature rather than acclimation of root respiration or substrate limitations. Methodological Issues: 5. The isotopic signatures of autotrophic and heterotrophic respiration are too similar for isotopic partitioning of belowground respiration into these two components at our site—in keeping with the recent findings of Bowling et al. (2015) in a subalpine conifer forest. 6. Artifacts of the trenching method, such as changes in soil moisture and increased carbon substrate from the newly severed roots, are significant and need to be quantified when determining daily to annual estimates of autotrophic and heterotrophic respiration. C. Effects of simulated exudates on priming of microbial decomposition: The stoichiometry of root exudates influences both the amount and the mechanism by which priming occurs. At low C:N, SOC loss is caused by an increase in microbial efficiency. At high C:N, SOC loss is caused by an increase in microbial biomass. D. Modeling with the Ecosystem Demography Model (ED2): 1. Incorporation of 13C tracking to create an isotopically-enabled Ecosystem Demography v2 model (ED2) 2. State-of-the-art parameter optimization methodology developed for improving ED2 model predictions and parameters. 3. Significantly improved model predictions of growth- and maintenance-related carbon fluxes and 13C fluxes« less

Partitioning CO<sub>2</sub> fluxes with isotopologue measurements and modeling to understand mechanisms of forest carbon sequestration