Abstract

Abstract. Several land biogeochemical models used for studying carbon–climate feedbacks have begun explicitly representing microbial dynamics. However, to our knowledge, there has been no theoretical work on how to achieve a consistent scaling of the complex biogeochemical reactions from microbial individuals to populations, communities, and interactions with plants and mineral soils. We focus here on developing a mathematical formulation of the substrate–consumer relationships for consumer-mediated redox reactions of the form A + BE→ products, where products could be, e.g., microbial biomass or bioproducts. Under the quasi-steady-state approximation, these substrate–consumer relationships can be formulated as the computationally difficult full equilibrium chemistry problem or approximated analytically with the dual Monod (DM) or synthesizing unit (SU) kinetics. We find that DM kinetics is scaling inconsistently for reaction networks because (1) substrate limitations are not considered, (2) contradictory assumptions are made regarding the substrate processing rate when transitioning from single- to multi-substrate redox reactions, and (3) the product generation rate cannot be scaled from one to multiple substrates. In contrast, SU kinetics consistently scales the product generation rate from one to multiple substrates but predicts unrealistic results as consumer abundances reach large values with respect to their substrates. We attribute this deficit to SU's failure to incorporate substrate limitation in its derivation. To address these issues, we propose SUPECA (SU plus the equilibrium chemistry approximation – ECA) kinetics, which consistently imposes substrate and consumer mass balance constraints. We show that SUPECA kinetics satisfies the partition principle, i.e., scaling invariance across a network of an arbitrary number of reactions (e.g., as in Newton's law of motion and Dalton's law of partial pressures). We tested SUPECA kinetics with the equilibrium chemistry solution for some simple problems and found SUPECA outperformed SU kinetics. As an example application, we show that a steady-state SUPECA-based approach predicted an aerobic soil respiration moisture response function that agreed well with laboratory observations. We conclude that, as an extension to SU and ECA kinetics, SUPECA provides a robust mathematical representation of complex soil substrate–consumer interactions and can be applied to improve Earth system model (ESM) land models.

Highlights

  • Near-surface soils hold more than twice the carbon in the current atmosphere; a small change in land carbon dynamics can imply significant feedbacks to the ongoing climate warming (Ciais et al, 2013)

  • We find that dual Monod (DM) kinetics is scaling inconsistently for reaction networks because (1) substrate limitations are not considered, (2) contradictory assumptions are made regarding the substrate processing rate when transitioning from single- to multi-substrate redox reactions, and (3) the product generation rate cannot be scaled from one to multiple substrates

  • We address the above process-scaling question by first presenting the step-by-step derivation of DM kinetics and synthesizing unit (SU) kinetics from the equilibrium chemistry (EC) formulation of the redox reaction kinetics can be viewed as a direct application of chemical kinetics that the reaction rate of substrates A and B with consumer E is determined by the product of A and B’s binding probability to E (which in Monod form is [A] (KA + [A]) for substrate

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Summary

Introduction

Near-surface soils hold more than twice the carbon in the current atmosphere; a small change in land carbon dynamics can imply significant feedbacks to the ongoing climate warming (Ciais et al, 2013). To r-K scaling (e.g., Tilman, 1982; Litchman and Klausmeier, 2008), single-substrate Monod kinetics requires the product-generation rate to be faster than the dissociation rate of consumer–substrate complexes This contrasting requirement on the relative magnitudes of parameters, as we will show later, implies that DM kinetics cannot achieve consistent scaling of substrate–consumer interactions for generic biogeochemical modeling. We define a kinetic formulation to have consistent process scaling when the formulated substrate–consumer relationship (1) can seamlessly transition from a single substrate– consumer pair to a network of many substrate–consumer pairs without changing its mathematical form ( known as the partition principle as in Newton’s second law of motion; Feynman et al, 1963) and (2) does not predict any singularity over the range of substrate and consumer concentrations ( known as the non-singular principle; e.g., Schnell and Maini, 2000; Tang, 2015). We discuss how one can apply SUPECA kinetics to traitbased modeling approaches in various biogeochemical systems (e.g., Bouskill et al, 2012; Follows et al, 2007; Litchman and Klausmeier, 2008)

Governing equations
Dual Monod kinetics and synthesizing unit kinetics
Dual Monod kinetics
Synthesizing unit kinetics
SUPECA kinetics
SUPECA kinetics for a network of reactions
Numerical accuracy of SUPECA kinetics
Example application to modeling aerobic heterotrophic respiration
Findings
Conclusions
Full Text
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