DOE PAGES title logo U.S. Department of Energy
Office of Scientific and Technical Information

Title: The quasi-equilibrium framework revisited: analyzing long-term CO2 enrichment responses in plant–soil models

Abstract

Abstract. Elevated carbon dioxide (CO2) can increase plant growth, but themagnitude of this CO2 fertilization effect is modified by soilnutrient availability. Predicting how nutrient availability affects plantresponses to elevated CO2 is a key consideration for ecosystemmodels, and many modeling groups have moved to, or are moving towards,incorporating nutrient limitation in their models. The choice of assumptionsto represent nutrient cycling processes has a major impact on modelpredictions, but it can be difficult to attribute outcomes to specificassumptions in complex ecosystem simulation models. Here we revisit thequasi-equilibrium analytical framework introduced by Comins andMcMurtrie (1993) and explore the consequences of specific model assumptionsfor ecosystem net primary productivity (NPP). We review the literature applying this framework to plant–soilmodels and then analyze the effect of several new assumptions on predictedplant responses to elevated CO2. Examination of alternativeassumptions for plant nitrogen uptake showed that a linear function of themineral nitrogen pool or a linear function of the mineral nitrogen pool withan additional saturating function of root biomass yield similar CO2responses at longer timescales (>5 years), suggesting that the addedcomplexity may not be needed when these are the timescales of interest. Incontrast, a saturating function of the mineral nitrogen pool with lineardependency on root biomass yields no soil nutrientmore » feedback on thevery-long-term (>500 years), near-equilibrium timescale, meaning that oneshould expect the model to predict a full CO2 fertilization effecton production. Secondly, we show that incorporating a priming effect on slowsoil organic matter decomposition attenuates the nutrient feedback effect onproduction, leading to a strong medium-term (5–50 years) CO2response. Models incorporating this priming effect should thus predict astrong and persistent CO2 fertilization effect over time. Thirdly,we demonstrate that using a “potential NPP” approach to represent nutrientlimitation of growth yields a relatively small CO2 fertilizationeffect across all timescales. Overall, our results highlight thefact that the quasi-equilibriumanalytical framework is effective for evaluating both the consequences andmechanisms through which different model assumptions affect predictions. Tohelp constrain predictions of the future terrestrial carbon sink, werecommend the use of this framework to analyze likely outcomes of new modelassumptions before introducing them to complex model structures.« less

Authors:
 [1]; ORCiD logo [2]; ORCiD logo [3]; ORCiD logo [4]; ORCiD logo [2];  [1];  [1]
  1. Western Sydney Univ., Penrith, NSW (Australia)
  2. Max Planck Institute of Biogeochemistry, Jena (Germany)
  3. Univ. of New South Wales, Sydney, NSW (Australia)
  4. Oak Ridge National Lab. (ORNL), Oak Ridge, TN (United States)
Publication Date:
Research Org.:
Oak Ridge National Lab. (ORNL), Oak Ridge, TN (United States)
Sponsoring Org.:
USDOE Office of Science (SC), Biological and Environmental Research (BER)
OSTI Identifier:
1558493
Grant/Contract Number:  
AC05-00OR22725
Resource Type:
Accepted Manuscript
Journal Name:
Geoscientific Model Development (Online)
Additional Journal Information:
Journal Name: Geoscientific Model Development (Online); Journal Volume: 12; Journal Issue: 5; Journal ID: ISSN 1991-9603
Publisher:
European Geosciences Union
Country of Publication:
United States
Language:
English
Subject:
58 GEOSCIENCES

Citation Formats

Jiang, Mingkai, Zaehle, Sönke, De Kauwe, Martin G., Walker, Anthony P., Caldararu, Silvia, Ellsworth, David S., and Medlyn, Belinda E. The quasi-equilibrium framework revisited: analyzing long-term CO2 enrichment responses in plant–soil models. United States: N. p., 2019. Web. doi:10.5194/gmd-12-2069-2019.
Jiang, Mingkai, Zaehle, Sönke, De Kauwe, Martin G., Walker, Anthony P., Caldararu, Silvia, Ellsworth, David S., & Medlyn, Belinda E. The quasi-equilibrium framework revisited: analyzing long-term CO2 enrichment responses in plant–soil models. United States. https://doi.org/10.5194/gmd-12-2069-2019
Jiang, Mingkai, Zaehle, Sönke, De Kauwe, Martin G., Walker, Anthony P., Caldararu, Silvia, Ellsworth, David S., and Medlyn, Belinda E. Tue . "The quasi-equilibrium framework revisited: analyzing long-term CO2 enrichment responses in plant–soil models". United States. https://doi.org/10.5194/gmd-12-2069-2019. https://www.osti.gov/servlets/purl/1558493.
@article{osti_1558493,
title = {The quasi-equilibrium framework revisited: analyzing long-term CO2 enrichment responses in plant–soil models},
author = {Jiang, Mingkai and Zaehle, Sönke and De Kauwe, Martin G. and Walker, Anthony P. and Caldararu, Silvia and Ellsworth, David S. and Medlyn, Belinda E.},
abstractNote = {Abstract. Elevated carbon dioxide (CO2) can increase plant growth, but themagnitude of this CO2 fertilization effect is modified by soilnutrient availability. Predicting how nutrient availability affects plantresponses to elevated CO2 is a key consideration for ecosystemmodels, and many modeling groups have moved to, or are moving towards,incorporating nutrient limitation in their models. The choice of assumptionsto represent nutrient cycling processes has a major impact on modelpredictions, but it can be difficult to attribute outcomes to specificassumptions in complex ecosystem simulation models. Here we revisit thequasi-equilibrium analytical framework introduced by Comins andMcMurtrie (1993) and explore the consequences of specific model assumptionsfor ecosystem net primary productivity (NPP). We review the literature applying this framework to plant–soilmodels and then analyze the effect of several new assumptions on predictedplant responses to elevated CO2. Examination of alternativeassumptions for plant nitrogen uptake showed that a linear function of themineral nitrogen pool or a linear function of the mineral nitrogen pool withan additional saturating function of root biomass yield similar CO2responses at longer timescales (>5 years), suggesting that the addedcomplexity may not be needed when these are the timescales of interest. Incontrast, a saturating function of the mineral nitrogen pool with lineardependency on root biomass yields no soil nutrient feedback on thevery-long-term (>500 years), near-equilibrium timescale, meaning that oneshould expect the model to predict a full CO2 fertilization effecton production. Secondly, we show that incorporating a priming effect on slowsoil organic matter decomposition attenuates the nutrient feedback effect onproduction, leading to a strong medium-term (5–50 years) CO2response. Models incorporating this priming effect should thus predict astrong and persistent CO2 fertilization effect over time. Thirdly,we demonstrate that using a “potential NPP” approach to represent nutrientlimitation of growth yields a relatively small CO2 fertilizationeffect across all timescales. Overall, our results highlight thefact that the quasi-equilibriumanalytical framework is effective for evaluating both the consequences andmechanisms through which different model assumptions affect predictions. Tohelp constrain predictions of the future terrestrial carbon sink, werecommend the use of this framework to analyze likely outcomes of new modelassumptions before introducing them to complex model structures.},
doi = {10.5194/gmd-12-2069-2019},
journal = {Geoscientific Model Development (Online)},
number = 5,
volume = 12,
place = {United States},
year = {Tue May 28 00:00:00 EDT 2019},
month = {Tue May 28 00:00:00 EDT 2019}
}

Journal Article:
Free Publicly Available Full Text
Publisher's Version of Record

Citation Metrics:
Cited by: 5 works
Citation information provided by
Web of Science

Save / Share:

Works referenced in this record:

Impact of priming on global soil carbon stocks
journal, February 2018

  • Guenet, Bertrand; Camino-Serrano, Marta; Ciais, Philippe
  • Global Change Biology, Vol. 24, Issue 5
  • DOI: 10.1111/gcb.14069

Carbon cost of plant nitrogen acquisition: global carbon cycle impact from an improved plant nitrogen cycle in the Community Land Model
journal, January 2016

  • Shi, Mingjie; Fisher, Joshua B.; Brzostek, Edward R.
  • Global Change Biology, Vol. 22, Issue 3
  • DOI: 10.1111/gcb.13131

Carbon-nitrogen interactions regulate climate-carbon cycle feedbacks: results from an atmosphere-ocean general circulation model
journal, January 2009


Towards a representation of priming on soil carbon decomposition in the global land biosphere model ORCHIDEE (version 1.9.5.2)
journal, January 2016

  • Guenet, Bertrand; Moyano, Fernando Esteban; Peylin, Philippe
  • Geoscientific Model Development, Vol. 9, Issue 2
  • DOI: 10.5194/gmd-9-841-2016

Using ecosystem experiments to improve vegetation models
journal, May 2015

  • Medlyn, Belinda E.; Zaehle, Sönke; De Kauwe, Martin G.
  • Nature Climate Change, Vol. 5, Issue 6
  • DOI: 10.1038/nclimate2621

Decade-long soil nitrogen constraint on the CO2 fertilization of plant biomass
journal, September 2012

  • Reich, Peter B.; Hobbie, Sarah E.
  • Nature Climate Change, Vol. 3, Issue 3
  • DOI: 10.1038/nclimate1694

The temporal response of forest ecosystems to doubled atmospheric CO2 concentration
journal, February 1996


Terrestrial nitrogen cycling in Earth system models revisited
journal, May 2016

  • Stocker, Benjamin D.; Prentice, I. Colin; Cornell, Sarah E.
  • New Phytologist, Vol. 210, Issue 4
  • DOI: 10.1111/nph.13997

Nitrogen feedbacks increase future terrestrial ecosystem carbon uptake in an individual-based dynamic vegetation model
journal, January 2014


Influence of carbon-nitrogen cycle coupling on land model response to CO 2 fertilization and climate variability : INFLUENCE OF CARBON-NITROGEN COUPLING
journal, December 2007

  • Thornton, Peter E.; Lamarque, Jean-François; Rosenbloom, Nan A.
  • Global Biogeochemical Cycles, Vol. 21, Issue 4
  • DOI: 10.1029/2006GB002868

A quantitative assessment of a terrestrial biosphere model's data needs across North American biomes: PEcAn/ED model-data uncertainty analysis
journal, March 2014

  • Dietze, Michael C.; Serbin, Shawn P.; Davidson, Carl
  • Journal of Geophysical Research: Biogeosciences, Vol. 119, Issue 3
  • DOI: 10.1002/2013JG002392

Carbon residence time dominates uncertainty in terrestrial vegetation responses to future climate and atmospheric CO 2
journal, December 2013

  • Friend, Andrew D.; Lucht, Wolfgang; Rademacher, Tim T.
  • Proceedings of the National Academy of Sciences, Vol. 111, Issue 9
  • DOI: 10.1073/pnas.1222477110

A process-based model of nitrogen cycling in forest plantations
journal, October 2005


Modelling Canopy Production. II. From Single-Leaf Photosynthesis Parameters to Daily Canopy Photosynthesis
journal, January 1995


A semi-analytical solution to accelerate spin-up of a coupled carbon and nitrogen land model to steady state
journal, January 2012

  • Xia, J. Y.; Luo, Y. Q.; Wang, Y. -P.
  • Geoscientific Model Development, Vol. 5, Issue 5
  • DOI: 10.5194/gmd-5-1259-2012

A representation of the phosphorus cycle for ORCHIDEE (revision 4520)
journal, January 2017

  • Goll, Daniel S.; Vuichard, Nicolas; Maignan, Fabienne
  • Geoscientific Model Development, Vol. 10, Issue 10
  • DOI: 10.5194/gmd-10-3745-2017

Nutrient availability as the key regulator of global forest carbon balance
journal, April 2014

  • Fernández-Martínez, M.; Vicca, S.; Janssens, I. A.
  • Nature Climate Change, Vol. 4, Issue 6
  • DOI: 10.1038/nclimate2177

Sources of Uncertainty in Modeled Land Carbon Storage within and across Three MIPs: Diagnosis with Three New Techniques
journal, April 2018


Nitrogen Availability Reduces CMIP5 Projections of Twenty-First-Century Land Carbon Uptake
journal, March 2015


The role of stoichiometric flexibility in modelling forest ecosystem responses to nitrogen fertilization
journal, July 2015

  • Meyerholt, Johannes; Zaehle, Sönke
  • New Phytologist, Vol. 208, Issue 4
  • DOI: 10.1111/nph.13547

Variability of projected terrestrial biosphere responses to elevated levels of atmospheric CO 2 due to uncertainty in biological nitrogen fixation
journal, January 2016

  • Meyerholt, Johannes; Zaehle, Sönke; Smith, Matthew J.
  • Biogeosciences, Vol. 13, Issue 5
  • DOI: 10.5194/bg-13-1491-2016

Reconciling the optimal and empirical approaches to modelling stomatal conductance: RECONCILING OPTIMAL AND EMPIRICAL STOMATAL MODELS
journal, January 2011


Consequences of Considering Carbon–Nitrogen Interactions on the Feedbacks between Climate and the Terrestrial Carbon Cycle
journal, August 2008

  • Sokolov, Andrei P.; Kicklighter, David W.; Melillo, Jerry M.
  • Journal of Climate, Vol. 21, Issue 15
  • DOI: 10.1175/2008JCLI2038.1

Soil hydrological properties regulate grassland ecosystem responses to multifactor global change: A modeling analysis
journal, January 2008

  • Weng, Ensheng; Luo, Yiqi
  • Journal of Geophysical Research, Vol. 113, Issue G3
  • DOI: 10.1029/2007JG000539

Temperature acclimation of photosynthesis and respiration: A key uncertainty in the carbon cycle-climate feedback: PHYSIOLOGICAL TEMPERATURE ACCLIMATION
journal, October 2015

  • Lombardozzi, Danica L.; Bonan, Gordon B.; Smith, Nicholas G.
  • Geophysical Research Letters, Vol. 42, Issue 20
  • DOI: 10.1002/2015GL065934

Diagnosing errors in a land surface model (CABLE) in the time and frequency domains
journal, January 2011

  • Wang, Ying Ping; Kowalczyk, Eva; Leuning, Ray
  • Journal of Geophysical Research, Vol. 116, Issue G1
  • DOI: 10.1029/2010JG001385

Uncertainty in the response of terrestrial carbon sink to environmental drivers undermines carbon-climate feedback predictions
journal, July 2017


Reducing uncertainty in projections of terrestrial carbon uptake
journal, April 2017

  • Lovenduski, Nicole S.; Bonan, Gordon B.
  • Environmental Research Letters, Vol. 12, Issue 4
  • DOI: 10.1088/1748-9326/aa66b8

Modelling forest response to increasing CO2 concentration under nutrient-limited conditions
journal, October 1994


A roadmap for improving the representation of photosynthesis in Earth system models
journal, November 2016

  • Rogers, Alistair; Medlyn, Belinda E.; Dukes, Jeffrey S.
  • New Phytologist, Vol. 213, Issue 1
  • DOI: 10.1111/nph.14283

Nutrient limitation reduces land carbon uptake in simulations with a model of combined carbon, nitrogen and phosphorus cycling
journal, January 2012


Growth of mature boreal Norway spruce was not affected by elevated [CO2] and/or air temperature unless nutrient availability was improved
journal, July 2013


Increased forest carbon storage with increased atmospheric CO2 despite nitrogen limitation: a game-theoretic allocation model for trees in competition for nitrogen and light
journal, December 2014

  • Dybzinski, Ray; Farrior, Caroline E.; Pacala, Stephen W.
  • Global Change Biology, Vol. 21, Issue 3
  • DOI: 10.1111/gcb.12783

Representation of vegetation dynamics in the modelling of terrestrial ecosystems: comparing two contrasting approaches within European climate space
journal, November 2001


Carbon–nitrogen interactions in idealized simulations with JSBACH (version 3.10)
journal, January 2017

  • Goll, Daniel S.; Winkler, Alexander J.; Raddatz, Thomas
  • Geoscientific Model Development, Vol. 10, Issue 5
  • DOI: 10.5194/gmd-10-2009-2017

Model structures amplify uncertainty in predicted soil carbon responses to climate change
journal, June 2018


Soil processes dominate the long-term response of forest net primary productivity to increased temperature and atmospheric CO 2 concentration
journal, June 2000

  • Medlyn, Belinda E.; McMurtrie, Ross E.; Dewar, Roderick C.
  • Canadian Journal of Forest Research, Vol. 30, Issue 6
  • DOI: 10.1139/x00-026

Evaluation of 11 terrestrial carbon-nitrogen cycle models against observations from two temperate Free-Air CO 2 Enrichment studies
journal, January 2014

  • Zaehle, Sönke; Medlyn, Belinda E.; De Kauwe, Martin G.
  • New Phytologist, Vol. 202, Issue 3
  • DOI: 10.1111/nph.12697

Predicting long-term carbon sequestration in response to CO 2 enrichment: How and why do current ecosystem models differ?
journal, April 2015

  • Walker, Anthony P.; Zaehle, Sönke; Medlyn, Belinda E.
  • Global Biogeochemical Cycles, Vol. 29, Issue 4
  • DOI: 10.1002/2014GB004995

Interactions between soil and tree roots accelerate long-term soil carbon decomposition
journal, November 2007


Plant root distributions and nitrogen uptake predicted by a hypothesis of optimal root foraging: Optimal Root Foraging
journal, May 2012

  • McMurtrie, Ross E.; Iversen, Colleen M.; Dewar, Roderick C.
  • Ecology and Evolution, Vol. 2, Issue 6
  • DOI: 10.1002/ece3.266

A Model of Multiple-Element Limitation for Acclimating Vegetation
journal, August 1992

  • Rastetter, Edward B.; Shaver, Gaius R.
  • Ecology, Vol. 73, Issue 4
  • DOI: 10.2307/1940666

Representing leaf and root physiological traits in CLM improves global carbon and nitrogen cycling predictions: LEAF AND ROOT TRAITS IN CLM
journal, May 2016

  • Ghimire, Bardan; Riley, William J.; Koven, Charles D.
  • Journal of Advances in Modeling Earth Systems, Vol. 8, Issue 2
  • DOI: 10.1002/2015MS000538

On the validation of models of forest CO2 exchange using eddy covariance data: some perils and pitfalls
journal, July 2005


Rhizosphere processes are quantitatively important components of terrestrial carbon and nutrient cycles
journal, January 2015

  • Finzi, Adrien C.; Abramoff, Rose Z.; Spiller, Kimberly S.
  • Global Change Biology, Vol. 21, Issue 5
  • DOI: 10.1111/gcb.12816

Nitrogen cycling and feedbacks in a global dynamic land model: MODELING THE LAND NITROGEN CYCLE
journal, January 2010

  • Gerber, Stefan; Hedin, Lars O.; Oppenheimer, Michael
  • Global Biogeochemical Cycles, Vol. 24, Issue 1
  • DOI: 10.1029/2008GB003336

Modelling forest‐growth response to increasing CO 2 concentration in relation to various factors affecting nutrient supply
journal, January 1998


Why is plant-growth response to elevated CO2 amplified when water is limiting, but reduced when nitrogen is limiting? A growth-optimisation hypothesis
journal, January 2008

  • McMurtrie, Ross E.; Norby, Richard J.; Medlyn, Belinda E.
  • Functional Plant Biology, Vol. 35, Issue 6
  • DOI: 10.1071/FP08128

Nitrogen limitation on land: how can it occur in Earth system models?
journal, February 2015

  • Thomas, R. Quinn; Brookshire, E. N. Jack; Gerber, Stefan
  • Global Change Biology, Vol. 21, Issue 5
  • DOI: 10.1111/gcb.12813

Microbe-driven turnover offsets mineral-mediated storage of soil carbon under elevated CO2
journal, November 2014

  • Sulman, Benjamin N.; Phillips, Richard P.; Oishi, A. Christopher
  • Nature Climate Change, Vol. 4, Issue 12
  • DOI: 10.1038/nclimate2436

CO2 enhancement of forest productivity constrained by limited nitrogen availability
journal, October 2010

  • Norby, R. J.; Warren, J. M.; Iversen, C. M.
  • Proceedings of the National Academy of Sciences, Vol. 107, Issue 45
  • DOI: 10.1073/pnas.1006463107

RESPONSES OF N-LIMITED ECOSYSTEMS TO INCREASED CO 2 : A BALANCED-NUTRITION, COUPLED-ELEMENT-CYCLES MODEL
journal, May 1997


Analytical model of stemwood growth in relation to nitrogen supply
journal, January 1996


Using models to guide field experiments: a priori predictions for the CO 2 response of a nutrient- and water-limited native Eucalypt woodland
journal, May 2016

  • Medlyn, Belinda E.; De Kauwe, Martin G.; Zaehle, Sönke
  • Global Change Biology, Vol. 22, Issue 8
  • DOI: 10.1111/gcb.13268

Experimental warming altered rates of carbon processes, allocation, and carbon storage in a tallgrass prairie
journal, November 2015


Qualitative Analysis of Insect Outbreak Systems: The Spruce Budworm and Forest
journal, February 1978

  • Ludwig, D.; Jones, D. D.; Holling, C. S.
  • The Journal of Animal Ecology, Vol. 47, Issue 1
  • DOI: 10.2307/3939

Processes regulating progressive nitrogen limitation under elevated carbon dioxide: a meta-analysis
journal, January 2016


Progressive Nitrogen Limitation of Ecosystem Responses to Rising Atmospheric Carbon Dioxide
journal, January 2004


Traceable components of terrestrial carbon storage capacity in biogeochemical models
journal, March 2013

  • Xia, Jianyang; Luo, Yiqi; Wang, Ying-Ping
  • Global Change Biology, Vol. 19, Issue 7
  • DOI: 10.1111/gcb.12172

Controls on terrestrial carbon feedbacks by productivity versus turnover in the CMIP5 Earth System Models
journal, January 2015


Quantifying carbon-nitrogen feedbacks in the Community Land Model (CLM4): CARBON-NITROGEN FEEDBACKS IN CLM4
journal, April 2010

  • Bonan, Gordon B.; Levis, Samuel
  • Geophysical Research Letters, Vol. 37, Issue 7
  • DOI: 10.1029/2010GL042430

Long-Term Response of Nutrient-Limited Forests to CO"2 Enrichment; Equilibrium Behavior of Plant-Soil Models
journal, November 1993

  • Comins, H. N.; McMurtrie, R. E.
  • Ecological Applications, Vol. 3, Issue 4
  • DOI: 10.2307/1942099

Faster Decomposition Under Increased Atmospheric CO2 Limits Soil Carbon Storage
journal, April 2014


Nitrogen feedbacks increase future terrestrial ecosystem carbon uptake in an individual-based dynamic vegetation model
journal, January 2014


Changes in soil water holding capacity and water availability following vegetation restoration on the Chinese Loess Plateau
journal, May 2021


Reconciling the optimal and empirical approaches to modelling stomatal conductance
journal, October 2012