Kevin P. Tu

Reliable estimation of biochemical parameters from C3 leaf photosynthesis–intercellular carbon dioxide response curves

The Farquhar-von Caemmerer-Berry (FvCB) model of photosynthesis is a change-point model and structurally overparameterized for interpreting the response of leaf net assimilation (A) to intercellular CO₂ concentration (Ci). The use of conventional fitting methods may lead not only to incorrect parameters but also several previously unrecognized consequences. For example, the relationships between key parameters may be fixed computationally and certain fits may be produced in which the estimated parameters result in contradictory identification of the limitation states of the data. Here we describe a new approach that is better suited to the FvCB model characteristics. It consists of four main steps: (1) enumeration of all possible distributions of limitation states; (2) fitting the FvCB model to each limitation state distribution by minimizing a distribution-wise cost function that has desirable properties for parameter estimation; (3) identification and correction of inadmissible fits; and (4) selection of the best fit from all possible limitation state distributions. The new approach implemented theoretical parameter resolvability with numerical procedures that maximally use the information content of the data. It was tested with model simulations, sampled A/Ci curves, and chlorophyll fluorescence measurements of different tree species. The new approach is accessible through the automated website leafweb.ornl.gov.

Cellulose δ18O is an index of leaf-to-air vapor pressure difference (VPD) in tropical plants

Cellulose in plants contains oxygen that derives in most cases from precipitation. Because the stable oxygen isotope composition, δ18O, of precipitation is associated with environmental conditions, cellulose δ18O should be as well. However, plant physiological models using δ18O suggest that cellulose δ18O is influenced by a complex mix of both climatic and physiological drivers. This influence complicates the interpretation of cellulose δ18O values in a paleo-context. Here, we combined empirical data analyses with mechanistic model simulations to i) quantify the impacts that the primary climatic drivers humidity (ea) and air temperature (Tair) have on cellulose δ18O values in different tropical ecosystems and ii) determine which environmental signal is dominating cellulose δ18O values. Our results revealed that ea and Tair equally influence cellulose δ18O values and that distinguishing which of these factors dominates the δ18O values of cellulose cannot be accomplished in the absence of additional environmental information. However, the individual impacts of ea and Tair on the δ18O values of cellulose can be integrated into a single index of plant-experienced atmospheric vapor demand: the leaf-to-air vapor pressure difference (VPD). We found a robust relationship between VPD and cellulose δ18O values in both empirical and modeled data in all ecosystems that we investigated. Our analysis revealed therefore that δ18O values in plant cellulose can be used as a proxy for VPD in tropical ecosystems. As VPD is an essential variable that determines the biogeochemical dynamics of ecosystems, our study has applications in ecological-, climate-, or forensic-sciences.

Physiological consequences of height-related morphological variation in Sequoia sempervirens foliage

This study examined relationships between foliar morphology and gas exchange characteristics as they vary with height within and among crowns of Sequoia sempervirens D. Don trees ranging from 29 to 113 m in height. Shoot mass:area (SMA) ratio increased with height and was less responsive to changes in light availability as height increased, suggesting a transition from light to water relations as the primary determinant of morphology with increasing height. Mass-based rates of maximum photosynthesis (A(max,m)), standardized photosynthesis (A(std,m)) and internal CO(2) conductance (g(i,m)) decreased with height and SMA, while the light compensation point, light saturation point, and mass and area-based rates of dark respiration (R(m)) increased with height and SMA. Among foliage from different heights, much of the variation in standardized photosynthesis was explained by variation in g(i,) consistent with increasing limitation of photosynthesis by internal conductance in foliage with higher SMA. The syndrome of lower internal and stomatal conductance to CO(2) and higher respiration may contribute to reductions in upper crown growth efficiency with increasing height in S. sempervirens trees.

Partitioning Ecosystem Respiration Using Stable Carbon Isotope Analyses of CO 2

Recent studies have shown that natural abundance stable carbon isotopes can be used to partition measurements of net ecosystem CO 2 exchange into gross fluxes of photosynthesis and respiration. This chapter explores the related application of the natural abundance stable carbon isotope ratio of respired CO 2 to partitioning components of ecosystem respiration. Although, the exploitation of the natural variation in stable isotope abundance in respired CO 2 holds the most promise for widespread application among existing measurement networks (e.g., BASIN, FLUXNET, SIBAE) and for comparative studies focused on ecosystem C balance, the reliance on natural variation may also have limitations from the standpoint that such variation is typically small. Recent evidence suggests, however, that variation can be large and significant such that measurable differences among respiration signatures can allow partitioning of ecosystem respiration and therefore inform ecosystem C-balance studies.

On Uncertainty in Global Terrestrial Evapotranspiration Estimates from Choice of Input Forcing Datasets

AbstractEvapotranspiration ET is a critical water, energy, and climate variable, and recent work has been published comparing different global products. These comparisons have been difficult to interpret, however, because in most studies the evapotranspiration products were derived from models forced by different input data. Some studies have analyzed the uncertainty in regional evapotranspiration estimates from choice of forcings. Still others have analyzed how multiple models vary with choice of net radiation forcing data. However, no analysis has been conducted to determine the uncertainty in global evapotranspiration estimates attributable to each class of input forcing datasets. Here, one of these models [Priestly–Taylor JPL (PT-JPL)] is run with 19 different combinations of forcing data. These data include three net radiation products (SRB, CERES, and ISCCP), three meteorological datasets [CRU, Atmospheric Infrared Sounder (AIRS) Aqua, and MERRA], and three vegetation index products [MODIS; Global I...

Spatial variation in photosynthetic CO2 carbon and oxygen isotope discrimination along leaves of the monocot triticale (Triticum x Secale) relates to mesophyll conductance and the Péclet effect

Carbon and oxygen isotope discrimination of CO(2) during photosynthesis (Δ(13)C(obs) and Δ(18)O(obs)) were measured along a monocot leaf, triticale (Triticum × Secale). Both Δ(13)C(obs) and Δ(18)O(obs) increased towards the leaf tip. While this was expected for Δ(18)O(obs) , because of progressive enrichment of leaf water associated with the Péclet effect, the result was surprising for Δ(13) C(obs). To explore parameters determining this pattern, we measured activities of key photosynthetic enzymes [ribulose bis-phosphate carboxylase-oxygenase (Rubisco), phosphoenolpyruvate carboxylase (PEPC) and carbonic anhydrase) as well as maximum carboxylation and electron transport rates (V(cmax) and J(max)) along the leaf. Patterns in leaf internal anatomy along the leaf were also quantified. Mesophyll conductance (g(m)) is known to have a strong influence on Δ(13)C(obs) , so we used three commonly used estimation methods to quantify variation in g(m) along the leaf. Variation in Δ(13)C(obs) was correlated with g(m) and chloroplast surface area facing the intercellular air space, but unrelated to photosynthetic enzyme activity. The observed variation could cause errors at higher scales if the appropriate portion of a leaf is not chosen for leaf-level measurements and model parameterization. Our study shows that one-third of the way from the base of the leaf represents the most appropriate portion to enclose in the leaf chamber.

Isotopic composition of transpiration and rates of change in leaf water isotopologue storage in response to environmental variables

During daylight hours, the isotope composition of leaf water generally approximates steady-state leaf water isotope enrichment model predictions. However, until very recently there was little direct confirmation that isotopic steady-state (ISS) transpiration in fact exists. Using isotope ratio infrared spectroscopy (IRIS) and leaf gas exchange systems we evaluated the isotope composition of transpiration and the rate of change in leaf water isotopologue storage (isostorage) when leaves were exposed to variable environments. In doing so, we developed a method for controlling the absolute humidity entering the gas exchange cuvette for a wide range of concentrations without changing the isotope composition of water vapour. The measurement system allowed estimation of (18)O enrichment both at the evaporation site and for bulk leaf water, in the steady state and the non-steady state. We show that non-steady-state effects dominate the transpiration isoflux even when leaves are at physiological steady state. Our results suggest that a variable environment likely prevents ISS transpiration from being achieved and that this effect may be exacerbated by lengthy leaf water turnover times due to high leaf water contents.

Understanding the Stable Isotope Composition of Biosphere‐Atmosphere CO2 Exchange

Stable isotopes of atmospheric carbon dioxide (CO2) contain a wealth of information regarding biosphere-atmosphere interactions. The carbon isotope ratio of CO2 (δ13C) reflects the terrestrial carbon cycle including processes of photosynthesis, respiration, and decomposition. The oxygen isotope ratio (δ18O) reflects terrestrial carbon and water coupling due to CO2-H2O oxygen exchange. Isotopic CO2 measurements, in combination with ecosystem-isotopic exchange models, allow for the quantification of patterns and mechanisms regulating terrestrial carbon and water cycles, as well as for hypothesis development, data interpretation, and forecasting. Isotopic measurements and models have evolved significantly over the past two decades, resulting in organizations that promote model-measurement networks, e.g., the U.S. National Science Foundations Biosphere-Atmosphere Stable Isotope Network, the European Stable Isotopes in Biosphere-Atmosphere Exchange Network, and the U.S. National Environmental Observatory Network.

Latent Heat Flux and Canopy Conductance Based on Penman-Monteith, Priestley-Taylor Equation, and Bouchet's Complementary Hypothesis

A novel method is presented to analytically resolve the terrestrial latent heat flux (lE) and conductances (boundary layer gB and surface gS) using net radiation (RN), ground heat flux (G), air temperature (Ta), and relative humidity (RH). This method consists of set of equations where the two unknown internal state variables (gB and gS) were expressed in terms of the known core variables, combining diffusion equations, the Penman‐Monteith equation, the Priestley‐Taylor equation, and Bouchet’s complementary hypothesis. Estimated lE is validated with the independent eddy covariance lE observations over Soil Moisture Experiment 2002 (SMEX-02); the Global Energy and Water Cycle Experiment (GEWEX) Continental-Scale International Project (GCIP) selected sites from FLUXNET and tropics eddy flux, representing four climate zones (tropics, subtropics, temperate, and cold); and multiple biomes. The authors find a RMSE of 23.8‐ 54.6 W m 22 for hourly lEover SMEX-02 and GCIP and 23.8‐29.0 W m 22 for monthlylEover the FLUXNET and tropics. Observational and modeled evidence in the reduction in annual evaporation (E) pattern on the order of 33% from 1999 to 2006 was found in central Amazonia. Retrieved gS responded to vapor pressure deficit, measured lE, and gross photosynthesis in a theoretically robust behavior. However, the current scheme [Penman‐Monteith‐Bouchet‐Lhomme (PMBL)] showed some overestimation of lE in limited soil moisture regimes. PMBL provides similar results when compared with another Priestley‐Taylor‐based lE estimation approach [Priestley‐Taylor‐Jet Propulsion Laboratory (PT-JPL)] but with the advantage of having the conductances analytically recovered.

Stable isotopes as indicators, tracers and recorders of ecological change : synthesis and outlook

Publisher Summary The chapter presented here explores a diversity of ways in which stable isotopes inform present and future changes in a wide variety of ecological systems. The recent advances in contemporary research are highlighted, focusing on the knowledge and methodologies that have been developed during the past two decades. These approaches cover a wide range of time and space scales and levels of biological organization. As summarized by Ehleringer and Dawson, stable isotopes are powerful tools in ecological studies for tracing, recording, sourcing, and integrating different ecological parameters. Further, as noted by Dawson and Siegwolf, with environmental changes occurring at unprecedented rates in the earths history, stable isotope methods and applications will undoubtedly serve a critical role in documenting the nature and magnitude of change, as well as highlighting solutions for mitigating ecological impacts that threaten the future of all organisms. This chapter provides a synthesis of the major findings, knowledge gaps, and outlooks for future research on the use of stable isotopes as indicators, tracers, and recorders of ecological change.