Estimating Forest Productivity Using Carbon Isotopes in Tree Rings at Harvard Forest 1992-2010

doi:10.6073/pasta/5a75f8c54be833bc3b615975e0668b60 Estimating Forest Productivity Using Carbon Isotopes in Tree Rings at Harvard Forest 1992-2010 Soumaya Belmecheri https://orcid.org/0000-0003-1258-2741 R. Stockton Maxwell https://orcid.org/0000-0002-4811-5929 Kenneth Davies Alan Taylor 2023 English

We investigated relationships between tree-ring δ13C and growth, and flux tower estimates of gross primary productivity (GPP) at Harvard Forest from 1992 to 2010. Seasonal variations of derived photosynthetic isotope discrimination (Δ13C) and leaf intercellular CO2 concentration (ci) showed significant increasing trends for the dominant deciduous and coniferous species. Δ13C was positively correlated to growing-season GPP and is primarily controlled by precipitation and soil moisture indicating that site conditions maintained high stomatal conductance under increasing atmospheric CO2 levels. Increasing Δ13C over the 1992-2010 period is attributed to increasing annual and summer water availability identified at Harvard Forest and across the region. Higher Δ13C is coincident with an enhancement in growth and ecosystem-level net carbon uptake. This work suggests that tree-ring δ13C could serve as a measure of forest GPP and be used to improve the calibration and predictive skill of ecosystem and carbon cycle models.

carbon cellulose climate change photosynthesis stable isotopes tree physiology tree rings LTER controlled vocabulary organic matter disturbance LTER core area Harvard Forest HFR LTER USA HFR default

This dataset is released to the public under Creative Commons CC0 1.0 (No Rights Reserved). Please keep the dataset creators informed of any plans to use the dataset. Consultation with the original investigators is strongly encouraged. Publications and data products that make use of the dataset should include proper acknowledgement.

Creative Commons Zero v1.0 Universal https://spdx.org/licenses/CC0-1.0.html CC0-1.0 https://harvardforest.fas.harvard.edu/exist/apps/datasets/showData.html?id=hf235 Prospect Hill Tract (Harvard Forest). Coordinates based on WGS84 datum. -72.20 -72.17 +42.55 +42.53 280 420 meter 1992 2010 genus Quercus species rubra red oak genus Tsuga species canadensis hemlock complete Information Manager Harvard Forest

324 North Main Street Petersham MA 01366 USA

(978) 724-3302 hf-im@lists.fas.harvard.edu Harvard Forest

324 North Main Street Petersham MA 01366 USA

(978) 724-3302 (978) 724-3595 https://harvardforest.fas.harvard.edu

Ring Width Chronology and Basal area increment To identify interannual variability in tree growth and tree-ring δ13C, we cored Q. rubra and T. canadensis with a 5-mm increment borer at 1.37 m stem height (Table 1). Two cores were taken from each tree to build ring-width chronologies. Additionally, we cored five and four trees respectively of each species at the same height with a 12-mm increment borer. The large cores were used to analyze the δ13C in the LW of the tree rings. A sample of four trees is sufficient to achieve good precision of the δ13C sample mean (McCarroll and Loader; 2004; Leavitt, 2008). Ring-width chronologies were developed for each species using standard dendrochronological techniques (Speer, 2010). For the site, Q. rubra mean age was 97 years (range 71-115 yrs) and T. canadensis mean age was 145 years (range 89-221 yrs). For isotope cores, Q. rubra mean age was 103 years (range 87-113 yrs) and T. canadensis mean age was 142 years (range 93-188 yrs). All ages were estimated from inner ring dates. Basal area increment (BAI) was calculated for each tree and species using the dplR package in R (Bunn, 2008). We used the “outside in” function to convert raw ring-width measurements to BAI based on the diameter of the tree and the width of each ring moving towards the pith of the tree. The method assumes a circular growth pattern. BAI was used instead of ring width as a surrogate of radial growth and carbon gain because it drepresents more accurately tree annual biomass increment without the need for standardization (Biondi and Qeadan 2008). A mean BAI value was computed for each species at an annual resolution, averaged over all the trees cored for both carbon isotope analyses and ring-width chronologies (i.e., 16 trees/42 cores for Q. rubra and 22 trees/55 cores for T. canadensis).

Stable istope analyses We analyzed the LW portion of each tree ring from individual trees over the 1992-2010 period. The samples were milled using an ultra-centrifugation mill (Qiagen TissueLyserII) and α-cellulose was extracted from each wood sample following the Soxhlet method elaborated by Green (1963) and modified by Leavitt and Danzer (1993). δ13C ratios were measured on the CO2 produced by α-cellulose combustion in a Costech elemental analyzer (EA) coupled with a Thermo Delta V-IRMS. The sample accuracy was determined to be within 0.08‰ (1 standard deviation calculated from the average difference between measured and true internal standard, n = 13). The isotopic value is expressed in the delta (δ) notation relative to the VPDB (‰ VPDB).

Data Analysis We used δ13C to determine carbon isotope discrimination (Δ) by the plant against atmospheric δ13C, and variation in plant iWUE. The Δ describes the isotopic difference between the δ13C of air (δ13Cair) and the plant (δ13Cplant) and results from the preferential use of 12C over 13C during photosynthesis. Δ is calculated using: Δ ‰ = (δ13Cair - δ13Cplant)/(1 + δ13Cplant/1000) [1] Records of δ13Cair were obtained from Mauna Loa from 1992 to 2002, and from aircraft measurements collected in Worcester, Massachusetts at 500 m above ground and available from 2003-2010 (White and Vaughn 2011). The model of Farquhar, O’Leary, and Berry 1982 describes isotopic Δ for C3 plants via: Δ ‰ = a + (b - a)ci/ca [2] where a is the fractionation during CO2 diffusion through the stomata (4.4‰: O'Leary, 1981); b is the fractionation by RuBP carboxylase (27‰: Farquhar and Richards, 1984); and ci and ca, are the leaf intercellular space and ambient CO2 concentrations (μmol. Mol-1), respectively. We used the data of atmospheric CO2 concentration (ca) measured at 29 m height on the eddy-covariance tower to calculate ci from Equation [2]. The fractionations due to diffusion and carboxylation are constant but additive; therefore, the δ13C records variations in ci as regulated by two main processes: stomatal conductance (ca/ci) and photosynthetic assimilation rate (A). The iWUE is the ratio of the net photosynthetic assimilation rate (A) and water vapor conductance (gH2O) and is described by Ehleringer and Cerling (1995) as: iWUE = A/gH2O = ca - ci(1/1.6) [3] We used summer (June, July and August) values of δ13Cair and ca to calculate Δ, and to reconstruct ci and iWUE for the period corresponding to LW formation.

Harvard Forest Long-Term Ecological Research Harvard Forest

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(978) 724-3302 (978) 724-3595 https://harvardforest.fas.harvard.edu https://ror.org/059cpzx98 pointOfContact The Harvard Forest Long-Term Ecological Research (LTER) program examines ecological dynamics in the New England region resulting from natural disturbances, environmental change, and human impacts. National Science Foundation LTER grants: DEB-8811764, DEB-9411975, DEB-0080592, DEB-0620443, DEB-1237491, DEB-1832210. hf235-01-carbon-isotope.csv carbon isotope data hf235-01-carbon-isotope.csv 2985 631bc517f2925851a83c0f496161df19 1 \r\n column , https://harvardforest.fas.harvard.edu/data/p23/hf235/hf235-01-carbon-isotope.csv year year YYYY 1 year NA missing value d13c.tsca delta-13 C of isotope analysis of Tsuga canadensis (TSCA) measured from four trees/cores (parts per thousand) dimensionless 0.00000001 real NA missing value d13c.quru delta-13 C of Quercus rubra (QURU) measured from five trees (parts per thousand) dimensionless 0.00000001 real NA missing value cid.tsca carbon isotope discrimination by the plant against atmospheric delta-13 C for Tsuga Canadensis (parts per thousand) dimensionless 0.00000001 real NA missing value cid.quru carbon isotope discrimination by the plant against atmospheric delta-13 C for Quercus rubra (parts per thousand) dimensionless 0.00000001 real NA missing value ci.tsca intercellular space CO2 concentration for Tsuga canadensis (ppm) dimensionless 0.00000001 real NA missing value ci.quru intercellular space CO2 concentration for Quercus rubra (ppm) dimensionless 0.00000001 real NA missing value iwue.tsca intrinsic water use efficiency for Tsuga canadensis (µmol/mol) dimensionless 0.0000001 real NA missing value iwue.quru intrinsic water use efficiency for Quercus rubra (µmol/mol) dimensionless 0.00000001 real NA missing value cica.tsca stomatal conductance for Tsuga canadensis (ci/ca) dimensionless 0.000000001 real NA missing value cica.quru stomatal conductance for Quercus rubra (ci/ca) dimensionless 0.000000001 real NA missing value bai.tsca basal area increment for all sampled trees of Tsuga canadensis centimeterSquaredPerYear 0.00000001 real NA missing value bai.quru basal area increment for all sampled trees of Quercus rubra centimeterSquaredPerYear 0.00000001 real NA missing value 19