skip to main content
OSTI.GOV title logo U.S. Department of Energy
Office of Scientific and Technical Information

Title: Aligning theoretical and empirical representations of soil carbon-to-nitrogen stoichiometry with process-based terrestrial biogeochemistry models

Journal Article · · Soil Biology and Biochemistry
ORCiD logo [1];  [2];  [3];  [4];  [5];  [6];  [2];  [5];  [7];  [8];  [9];  [5];  [10]
  1. University of Colorado, Boulder, CO (United States); University of Michigan, Ann Arbor, MI (United States)
  2. University of Montana, Missoula, MT (United States)
  3. West Virginia University, Morgantown, WV (United States)
  4. Lawrence Livermore National Laboratory (LLNL), Livermore, CA (United States)
  5. University of New Hampshire, Durham, NH (United States)
  6. Colorado State University, Fort Collins, CO (United States)
  7. Carbon Direct, New York, NY (United States)
  8. USDA Forest Service, Boise, ID (United States). Rocky Mountain Research Station
  9. University of Michigan, Ann Arbor, MI (United States); University of Minnesota, St. Paul, MN (United States)
  10. University of Colorado, Boulder, CO (United States); National Center for Atmospheric Research (NCAR), Boulder, CO (United States)

Soil carbon-nitrogen (C:N) stoichiometry acts as a control over decomposition and soil organic matter formation and loss, making it a key soil property for understanding ecosystem dynamics and projected ecosystems responses to global environmental change. However, the controls of soil C:N and how they respond to increasing pressures from global change agents are not fully understood. The “foundational” controls on soil C:N, namely plant and microbial C:N, have been used to predict soil C:N, but fail to accurately simulate all ecosystems and may be insufficient for predictions under global environmental change. Here we present an “emerging” representation of controls of soil C:N that includes plant-microbe-mineral feedbacks that have been shown to regulate soil C:N. We argue that including representation of these emerging drivers in process-based terrestrial biogeochemistry models, which include biological N fixation, mycorrhizae, priming, root exudation of organic acids, and mineralogy (including soil texture, mineral composition, and aggregation), will improve mechanistic representation of soil C:N and associated processes. Such improvements will produce models that will better simulate a variety of ecological states and predict soil C:N when global changes modify plant-microbe-mineral interactions. Here, we align our empirical understanding of controls of soil C:N with those controls represented in models, identifying contexts where emerging drivers might be particularly important to represent (e.g., priming and root exudation in nutrient-limited conditions) and areas of future work. Additionally, we show that implementing emerging drivers of soil C:N results in different simulated outcomes at steady state and in response to elevated atmospheric CO2. Our review and preliminary simulations support the need to incorporate emerging drivers of soil C:N into process-based terrestrial biogeochemistry models, allowing for both theoretical exploration of mechanisms and potentially more accurate predictions of land biogeochemical responses to global change.

Research Organization:
Lawrence Livermore National Laboratory (LLNL), Livermore, CA (United States)
Sponsoring Organization:
USDOE National Nuclear Security Administration (NNSA); United States Department of Agriculture (USDA); National Science Foundation (NSF)
Grant/Contract Number:
AC52-07NA27344; 2020-67019-31395; 1926413; DBI-2021898; DEB-1831944; 2031253; 2031238; 2022070; DEB-1754126
OSTI ID:
2293650
Report Number(s):
LLNL-JRNL-852812; 1080356
Journal Information:
Soil Biology and Biochemistry, Vol. 189, Issue N/A; ISSN 0038-0717
Publisher:
ElsevierCopyright Statement
Country of Publication:
United States
Language:
English

References (183)

Mineralogical impact on organic nitrogen across a long-term soil chronosequence (0.3–4100kyr) journal April 2010
Carbon cost of plant nitrogen acquisition: global carbon cycle impact from an improved plant nitrogen cycle in the Community Land Model journal January 2016
Multiple models and experiments underscore large uncertainty in soil carbon dynamics journal October 2018
Nutrient acquisition strategies augment growth in tropical N 2 ‐fixing trees in nutrient‐poor soil and under elevated CO 2 journal February 2019
Increasing effects of chronic nutrient enrichment on plant diversity loss and ecosystem productivity over time journal January 2021
Towards a representation of priming on soil carbon decomposition in the global land biosphere model ORCHIDEE (version 1.9.5.2) journal January 2016
Nitrogen‐induced acidification, not N‐nutrient, dominates suppressive N effects on arbuscular mycorrhizal fungi journal September 2020
Measured soil organic matter fractions can be related to pools in the RothC model journal June 2007
Effects of increasing ultraviolet B radiation on decomposition and soil organic matter dynamics: a synthesis and modelling study journal June 1994
Global meta-analysis of wood decomposition rates: a role for trait variation among tree species? journal January 2009
Organo–organic and organo–mineral interfaces in soil at the nanometer scale journal November 2020
Mineral protection of soil carbon counteracted by root exudates journal March 2015
A global model of carbon, nitrogen and phosphorus cycles for the terrestrial biosphere journal January 2010
Microbial contribution to post-fire tundra ecosystem recovery over the 21st century journal February 2022
Evaluating nitrogen cycling in terrestrial biosphere models: a disconnect between the carbon and nitrogen cycles journal August 2023
Modelling the influence of ectomycorrhizal decomposition on plant nutrition and soil carbon sequestration in boreal forest ecosystems journal October 2016
Feedbacks between plant N demand and rhizosphere priming depend on type of mycorrhizal association journal July 2017
Forest response to elevated CO2 is conserved across a broad range of productivity journal December 2005
Flexible Foliar Stoichiometry Reduces the Magnitude of the Global Land Carbon Sink journal November 2023
Mycorrhizal types differ in ecophysiology and alter plant nutrition and soil processes journal May 2019
Positive priming effect explained by microbial nitrogen mining and stoichiometric decomposition at different stages journal December 2022
Global negative effects of nitrogen deposition on soil microbes journal March 2018
Phosphorus and carbon in soil particle size fractions: A synthesis journal January 2020
Global Carbon Sequestration Is Highly Sensitive to Model‐Based Formulations of Nitrogen Fixation journal January 2020
Soil carbon cycling proxies: Understanding their critical role in predicting climate change feedbacks journal November 2017
Microbial carbon use efficiency and priming of soil organic matter mineralization by glucose additions in boreal forest soils with different C:N ratios journal April 2022
Modelling soil carbon and nitrogen dynamics using measurable and conceptual soil organic matter pools in APSIM journal March 2014
Root carbon inputs to the rhizosphere stimulate extracellular enzyme activity and increase nitrogen availability in temperate forest soils journal December 2012
Depth trends of soil organic matter C:N and 15N natural abundance controlled by association with minerals journal November 2017
Conceptualizing soil organic matter into particulate and mineral‐associated forms to address global change in the 21st century journal November 2019
Evaluating litter decomposition in earth system models with long-term litterbag experiments: an example using the Community Land Model version 4 (CLM4) journal October 2012
Fungal Traits Important for Soil Aggregation journal January 2020
Altered microbial community structure and organic matter composition under elevated CO2 and N fertilization in the duke forest: OM COMPOSITION UNDER FACE AND FERTILIZATION journal June 2010
Arbuscular Mycorrhizal Fungi Increase Organic Carbon Decomposition Under Elevated CO2 journal August 2012
Improving understanding of soil organic matter dynamics by triangulating theories, measurements, and models journal July 2018
Legumes are different: Leaf nitrogen, photosynthesis, and water use efficiency journal March 2016
Minerals in the rhizosphere: overlooked mediators of soil nitrogen availability to plants and microbes journal June 2018
Variability of projected terrestrial biosphere responses to elevated levels of atmospheric CO 2 due to uncertainty in biological nitrogen fixation journal January 2016
Climate, carbon content, and soil texture control the independent formation and persistence of particulate and mineral-associated organic matter in soil journal April 2020
A global meta-analysis on freeze-thaw effects on soil carbon and phosphorus cycling journal August 2021
Stoichiometric imbalances between terrestrial decomposer communities and their resources: mechanisms and implications of microbial adaptations to their resources journal January 2014
Diverse Mycorrhizal Associations Enhance Terrestrial C Storage in a Global Model journal April 2019
Carbon limitation overrides acidification in mediating soil microbial activity to nitrogen enrichment in a temperate grassland journal August 2021
Linking microbial activity and soil organic matter transformations in forest soils under elevated CO2 journal February 2005
Simulation of the effects of photodecay on long‐term litter decay using DayCent journal December 2016
Increases in the flux of carbon belowground stimulate nitrogen uptake and sustain the long-term enhancement of forest productivity under elevated CO2: C fluxes belowground and long-term FACE productivity journal February 2011
Addition of inorganic phosphorus to soil leads to desorption of organic compounds and thus to increased soil respiration journal March 2019
Beyond Static Benchmarking: Using Experimental Manipulations to Evaluate Land Model Assumptions journal October 2019
A mechanistic treatment of the dominant soil nitrogen cycling processes: Model development, testing, and application: NITROGEN CYCLE MODELING journal April 2008
Global-Scale Similarities in Nitrogen Release Patterns During Long-Term Decomposition journal January 2007
Chronic nitrogen additions suppress decomposition and sequester soil carbon in temperate forests journal July 2014
Poorly crystalline mineral phases protect organic matter in acid subsoil horizons journal February 2005
Impacts of nutrient addition on soil carbon and nitrogen stoichiometry and stability in globally-distributed grasslands journal May 2022
Responses of C:N stoichiometry in plants, soil, and microorganisms to nitrogen addition journal September 2020
A global analysis of soil microbial biomass carbon, nitrogen and phosphorus in terrestrial ecosystems: Global soil microbial biomass C, N and P journal December 2012
Land-Use Intensity Effects on Soil Organic Carbon Accumulation Rates and Mechanisms journal April 2007
Beyond clay: towards an improved set of variables for predicting soil organic matter content journal February 2018
Ecological controls on net ecosystem productivity of a seasonally dry annual grassland under current and future climates: Modelling with ecosys journal January 2012
Enhanced root exudation induces microbial feedbacks to N cycling in a pine forest under long-term CO2 fumigation: Rhizosphere feedbacks in CO2-enriched forests journal December 2010
Interactions among plants, bacteria, and fungi reduce extracellular enzyme activities under long-term N fertilization journal February 2018
Chronic nitrogen enrichment affects the structure and function of the soil microbial community in temperate hardwood and pine forests journal July 2004
Strong mineralogic control of soil organic matter composition in response to nutrient addition across diverse grassland sites journal September 2020
The Millennial model: in search of measurable pools and transformations for modeling soil carbon in the new century journal December 2017
Biological stoichiometry from genes to ecosystems journal November 2000
Assessing the effect of elevated carbon dioxide on soil carbon: a comparison of four meta-analyses journal August 2009
Arctic Soil Governs Whether Climate Change Drives Global Losses or Gains in Soil Carbon journal December 2019
Stoichiometrically coupled carbon and nitrogen cycling in the MIcrobial-MIneral Carbon Stabilization model version 1.0 (MIMICS-CN v1.0) journal January 2020
C:N:P stoichiometry in soil: is there a “Redfield ratio” for the microbial biomass? journal July 2007
Elevated CO 2 reduces the nitrogen concentration of plant tissues journal January 1998
Influence of earthworm activity on aggregate-associated carbon and nitrogen dynamics differs with agroecosystem management journal May 2007
Ecosystem model spin-up: Estimating steady state conditions in a coupled terrestrial carbon and nitrogen cycle model journal November 2005
Stoichiometric response of nitrogen-fixing and non-fixing dicots to manipulations of CO2, nitrogen, and diversity journal November 2006
Microbe-driven turnover offsets mineral-mediated storage of soil carbon under elevated CO2 journal November 2014
Asymmetric responses of terrestrial C:N:P stoichiometry to precipitation change journal June 2021
Mycorrhiza-mediated competition between plants and decomposers drives soil carbon storage journal January 2014
Nitrogen Mineralization: Challenges of a Changing Paradigm journal March 2004
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
The priming effect of organic matter: a question of microbial competition? journal June 2003
Implementation of mycorrhizal mechanisms into soil carbon model improves the prediction of long-term processes of plant litter decomposition journal March 2022
The Community Land Model Version 5: Description of New Features, Benchmarking, and Impact of Forcing Uncertainty journal December 2019
Nitrogen and Lignin Control of Hardwood Leaf Litter Decomposition Dynamics journal June 1982
AggModel: A soil organic matter model with measurable pools for use in incubation studies journal August 2013
Ecological controls on N 2 O emission in surface litter and near-surface soil of a managed grassland: modelling and measurements journal January 2016
Testing the environmental controls of microbial nitrogen-mining induced by semi-continuous labile carbon additions in the subarctic journal March 2022
Global patterns of plant and microbial biomass in response to CO2 fumigation journal April 2023
Decreasing of methanogenic activity in paddy fields via lowering ponding water temperature: A modeling investigation journal August 2014
The path less taken: Long-term N additions slow leaf litter decomposition and favor the physical transfer pathway of soil organic matter formation journal March 2022
Patterns and controls of foliar nutrient stoichiometry and flexibility across United States forests journal January 2023
Soil aggregate stability related to soil density, root length, and mycorrhiza using site-specific Alnus incana and Melanogaster variegatus s.l. journal August 2013
Biological Invasion byMyrica fayaAlters Ecosystem Development in Hawaii journal November 1987
Resource stoichiometry and the biogeochemical consequences of nitrogen deposition in a mixed deciduous forest journal December 2016
Impact of ecohydrological fluctuations on iron-redox cycling journal June 2019
Bacterial and Fungal Contributions to Carbon Sequestration in Agroecosystems journal January 2006
Controls over leaf litter decomposition in wet tropical forests journal December 2009
Effects of three global change drivers on terrestrial C:N:P stoichiometry: a global synthesis journal February 2017
Soil aggregation, aggregate stability, organic carbon and nitrogen in different soil aggregate fractions under forest and shrub vegetation on the Loess Plateau, China journal June 2010
Nitrogen cycling in CMIP6 land surface models: progress and limitations journal January 2020
Aggregate Structure and Carbon, Nitrogen, and Phosphorus in Native and Cultivated Soils1 journal January 1986
Past, Present, and Future Atmospheric Nitrogen Deposition journal May 2016
On the fate of anthropogenic nitrogen journal December 2008
Temperature and pH mediate stoichiometric constraints of organically derived soil nutrients journal November 2021
Simulating ectomycorrhiza in boreal forests: implementing ectomycorrhizal fungi model MYCOFON in CoupModel (v5) journal January 2018
Terrestrial C:N stoichiometry in response to elevated CO2 and N addition: a synthesis of two meta-analyses journal February 2011
Soil enzymes as indicators of soil function: A step toward greater realism in microbial ecological modeling journal December 2021
Soil C and N availability determine the priming effect: microbial N mining and stoichiometric decomposition theories journal April 2014
Isotopic evidence for oligotrophication of terrestrial ecosystems journal October 2018
The Microbial Efficiency-Matrix Stabilization (MEMS) framework integrates plant litter decomposition with soil organic matter stabilization: do labile plant inputs form stable soil organic matter? journal February 2013
Direct evidence for microbial-derived soil organic matter formation and its ecophysiological controls journal November 2016
Evidence, causes, and consequences of declining nitrogen availability in terrestrial ecosystems journal April 2022
Ecosystem feedbacks and cascade processes: understanding their role in the responses of Arctic and alpine ecosystems to environmental change journal May 2009
Mycorrhizal association as a primary control of the CO 2 fertilization effect journal June 2016
Submicron structures provide preferential spots for carbon and nitrogen sequestration in soils journal January 2014
Elevated CO 2 shifts soil microbial communities from K ‐ to r ‐strategists journal March 2021
Effects of nitrogen deposition and insect herbivory on patterns of ecosystem‐level carbon and nitrogen dynamics: results from the CENTURY model journal May 2004
Advancing the mechanistic understanding of the priming effect on soil organic matter mineralisation journal March 2022
Simulated rhizosphere deposits induce microbial N‐mining that may accelerate shrubification in the subarctic journal June 2020
Pathways of soil organic matter formation from above and belowground inputs in a Sorghum bicolor bioenergy crop journal February 2019
Functional role of microarthropods in soil aggregation journal March 2015
The mycorrhizal-associated nutrient economy: a new framework for predicting carbon-nutrient couplings in temperate forests journal April 2013
The Role of Plants in the Effects of Global Change on Nutrient Availability and Stoichiometry in the Plant-Soil System journal October 2012
Nitrogen-rich microbial products provide new organo-mineral associations for the stabilization of soil organic matter journal December 2017
Fire frequency drives decadal changes in soil carbon and nitrogen and ecosystem productivity journal December 2017
Sporadic P limitation constrains microbial growth and facilitates SOM accumulation in the stoichiometrically coupled, acclimating microbe–plant–soil model journal February 2022
Carbon:Nitrogen:Phosphorus Stoichiometry in Fungi: A Meta-Analysis journal July 2017
Constraining Carbon and Nutrient Flows in Soil With Ecological Stoichiometry journal October 2019
A unifying framework for dinitrogen fixation in the terrestrial biosphere journal June 2008
Elevated CO2 increases root exudation from loblolly pine (Pinus taeda) seedlings as an N-mediated response journal October 2009
Soil macroaggregate turnover and microaggregate formation: a mechanism for C sequestration under no-tillage agriculture journal December 2000
A trade-off between plant and soil carbon storage under elevated CO2 journal March 2021
Soil food web properties explain ecosystem services across European land use systems journal August 2013
C:N ratios of bulk soils and particle-size fractions: Global trends and major drivers journal November 2022
Modeling ecosystem-scale carbon dynamics in soil: The microbial dimension journal March 2023
Nitrogen input enhances microbial carbon use efficiency by altering plant–microbe–mineral interactions journal June 2022
Assessing microbial residues in soil as a potential carbon sink and moderator of carbon use efficiency journal November 2020
Dual, differential isotope labeling shows the preferential movement of labile plant constituents into mineral-bonded soil organic matter journal March 2016
SOIL CARBON SEQUESTRATION AND TURNOVER IN A PINE FOREST AFTER SIX YEARS OF ATMOSPHERIC CO 2 ENRICHMENT journal July 2005
In Search of a Binding Agent: Nano-Scale Evidence of Preferential Carbon Associations with Poorly-Crystalline Mineral Phases in Physically-Stable, Clay-Sized Aggregates journal May 2018
Leguminous plants significantly increase soil nitrogen cycling across global climates and ecosystem types journal May 2023
SOM fractionation methods: Relevance to functional pools and to stabilization mechanisms journal September 2007
The contentious nature of soil organic matter journal November 2015
Exploring the Role of Cryptic Nitrogen Fixers in Terrestrial Ecosystems: A Frontier in Nitrogen Cycling Research journal November 2022
Soil carbon storage informed by particulate and mineral-associated organic matter journal November 2019
Effects of model structural uncertainty on carbon cycle projections: biological nitrogen fixation as a case study journal April 2015
Evaluation of 11 terrestrial carbon-nitrogen cycle models against observations from two temperate Free-Air CO 2 Enrichment studies journal January 2014
Integrating microbial physiology and physio-chemical principles in soils with the MIcrobial-MIneral Carbon Stabilization (MIMICS) model journal January 2014
Biological Invasion by Myrica Faya in Hawai'i: Plant Demography, Nitrogen Fixation, Ecosystem Effects journal February 1989
Interactions between plant growth and soil nutrient cycling under elevated CO 2 : a meta-analysis journal November 2006
Functional Ecology of Free-Living Nitrogen Fixation: A Contemporary Perspective journal December 2011
Simulating measurable ecosystem carbon and nitrogen dynamics with the mechanistically defined MEMS 2.0 model journal May 2021
Patterns in decomposition rates among photosynthetic organisms: the importance of detritus C:N:P content journal July 1993
Expansion of high-latitude deciduous forests driven by interactions between climate warming and fire journal August 2019
Effects of soil Nutrient Availability on Investment in Acquisition of n and p in Hawaiian rain Forests journal April 2001
Responses of a tundra system to warming using SCAMPS: a stoichiometrically coupled, acclimating microbe–plant–soil model journal February 2014
Nitrogen limitation on land: how can it occur in Earth system models? journal February 2015
Effects of forest management on soil C and N storage: meta analysis journal January 2001
Multi‐element regulation of the tropical forest carbon cycle
  • Townsend, Alan R.; Cleveland, Cory C.; Houlton, Benjamin Z.
  • Frontiers in Ecology and the Environment, Vol. 9, Issue 1 https://doi.org/10.1890/100047
journal February 2011
Nitrogen limitation of decomposition and decay: How can it occur? journal December 2017
Priming effect and microbial diversity in ecosystem functioning and response to global change: a modeling approach using the SYMPHONY model journal February 2014
Carbon cost of plant nitrogen acquisition: A mechanistic, globally applicable model of plant nitrogen uptake, retranslocation, and fixation: CARBON COST OF PLANT N ACQUISITION journal March 2010
The Stoichiometry of Consumer-Driven Nutrient Recycling: Theory, Observations, and Consequences journal April 1999
Sorption of dissolved organic nitrogen by acid subsoil horizons and individual mineral phases journal September 2000
Observations and modeling of biomass and soil organic matter dynamics for the grassland biome worldwide journal December 1993
A novel representation of biological nitrogen fixation and competitive dynamics between nitrogen-fixing and non-fixing plants in a land model (GFDL LM4.1-BNF) journal July 2021
The C:N:P:S stoichiometry of soil organic matter journal September 2016
Global stocks and capacity of mineral-associated soil organic carbon journal July 2022
Adsorption of proteins and nucleic acids on clay minerals and their interactions: A review journal August 2013
Carbon structure and enzyme activities in alpine and forest ecosystems journal November 2007
Hysteretic temperature sensitivity of wetland CH4 fluxes explained by substrate availability and microbial activity journal January 2020
A holistic framework integrating plant-microbe-mineral regulation of soil bioavailable nitrogen journal May 2021
Biological nitrogen fixation: rates, patterns and ecological controls in terrestrial ecosystems journal July 2013
21st‐century biogeochemical modeling: Challenges for Century‐based models and where do we go from here? journal August 2020
The Global Distribution of Biological Nitrogen Fixation in Terrestrial Natural Ecosystems journal March 2020
Microbial and abiotic controls on mineral-associated organic matter in soil profiles along an ecosystem gradient journal July 2019
The role of phosphorus dynamics in tropical forests – a modeling study using CLM-CNP journal January 2014
Improved global-scale predictions of soil carbon stocks with Millennial Version 2 journal January 2022
Unifying soil organic matter formation and persistence frameworks: the MEMS model journal January 2019
Combination of energy limitation and sorption capacity explains 14C depth gradients journal September 2020
Divergent controls of soil organic carbon between observations and process-based models journal July 2021
SoDaH: the SOils DAta Harmonization database, an open-source synthesis of soil data from research networks, version 1.0 journal January 2021
Effects of elevated CO2 on the C:N stoichiometry of plants, soils, and microorganisms in terrestrial ecosystems journal June 2021
Altered ecosystem carbon and nitrogen cycles by plant invasion: a meta-analysis journal February 2008
Carbon cycle confidence and uncertainty: Exploring variation among soil biogeochemical models journal November 2017