General Overview


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This report is the result of the use of the Python 3.4 package Sympy (for symbolic mathematics), as means to translate published models to a common language. It was created by Holger Metzler (Orcid ID: 0000-0002-8239-1601) on 11/03/2016, and was last modified on lm.

About the model

The model depicted in this document considers soil organic matter decomposition. It was originally described by Schimel & Weintraub (2003).

Abstract

Traditional models of soil organic matter (SOM) decomposition are all based on first order kinetics in which the decomposition rate of a particular C pool is proportional to the size of the pool and a simple decomposition constant (\(dC/dt = kC\)). In fact, SOM decomposition is catalyzed by extracellular enzymes that are produced by microorganisms. We built a simple theoretical model to explore the behavior of the decomposition – microbial growth system when the fundamental kinetic assumption is changed from first order kinetics to exoenzymes catalyzed decomposition (\(dC/dt = KC\times Enzymes\)). An analysis of the enzyme kinetics showed that there must be some mechanism to produce a non-linear response of decomposition rates to enzyme concentration—the most likely is competition for enzyme binding on solid substrates as predicted by Langmuir adsorption isotherm theory. This non-linearity also induces C limitation, regardless of the potential supply of C. The linked C and N version of the model showed that actual polymer breakdown and microbial use of the released monomers can be disconnected, and that it requires relatively little N to maintain the maximal rate of decomposition, regardless of the microbial biomass’ ability to use the breakdown products. In this model, adding a pulse of C to an N limited system increases respiration, while adding N actually decreases respiration (as C is redirected from waste respiration to microbial growth). For many years, researchers have argued that the lack of a respiratory response by soil microbes to added N indicates that they are not N limited. This model suggests that conclusion may be wrong. While total C flow may be limited by the functioning of the exoenzyme system, actual microbial growth may be N limited.

Keywords

differential equations, nonlinear, time invariant

Principles

heterogeneity of speed of decay, internal transformations of organic matter, substrate interactions

Available parameter values

Information on given parameter sets
Abbreviation Description
Set 1 original values from linked model (no nitrogen cycle considered in this model here)

State Variables

The following table contains the available information regarding this section:

Information on State Variables
Name Description Units
\(S\) soil organic carbon \(g\)
\(D\) dissolved organic carbon \(g\)
\(M\) microbial biomass \(g\)
\(E\) exoenzymes \(g\)

Decomposition Rates

The following table contains the available information regarding this section:

Information on Decomposition Rates
Name Description Type Units Values

Set 1
\(k_{d}\) collapsed decomposition constant parameter \(d^{-1}\) \(1\)
\(k_{m}\) microbial maintenance rate parameter \(d^{-1}\) \(0.01\)
\(k_{t}\) rate of microbial biomass that dies in each time interval parameter \(d^{-1}\) \(0.012\)
\(k_{l}\) decay constant for exoenzymes parameter \(d^{-1}\) \(0.05\)

Additional Parameters

The following table contains the available information regarding this section:

Information on Additional Parameters
Name Description Type Values

Set 1
\(SUE\) substrate use efficiency parameter \(0.5\)
\(K_{e}\) fraction of of dissolved organic carbon that is allocated to exoenzymes parameter \(0.05\)
\(K_{r}\) proportion of dead microbial biomass that is available for microbial use parameter \(0.85\)

Respiration Variables

The following table contains the available information regarding this section:

Information on Respiration Variables
Name Description Expressions Values

Set 1
\(R_{e}\) respiration to support enzyme synthesis \(R_{e}=\frac{K_{e}\,D\,\left(1-SUE\right)}{SUE}\) -
\(R_{m}\) respiration to support maintenance energy needs \(R_{m}=k_{m}\,M\) -
\(R_{g}\) respiration to support growth of the microbial biomass (not allowed to be negative) \(R_{g}=\operatorname{Max}\left(\left(D-K_{e}\,D-R_{e}-R_{m}\right)\,\left(1-SUE\right), 0\right)\) -

Components

The following table contains the available information regarding this section:

Information on Components
Name Description Expressions
\(C\) carbon content \(C=\left[\begin{matrix}S\\D\\M\\E\end{matrix}\right]\)
\(T\) transition operator \(T=\left[\begin{matrix}-1 & 0 & 0 & 0\\1 & -1 &\frac{K_{r}\cdot k_{t}}{SUE\cdot k_{m} + k_{t}} & 0\\0 & - K_{e} + SUE & -1 & 0\\0 & K_{e} & 0 & -1\end{matrix}\right]\)
\(N\) decomposition operator \(N=\left[\begin{matrix}\frac{E}{S}\cdot k_{d} & 0 & 0 & 0\\0 & 1 & 0 & 0\\0 & 0 & SUE\cdot k_{m} + k_{t} & 0\\0 & 0 & 0 & k_{l}\end{matrix}\right]\)
\(f_{s}\) the right hand side of the ode \(f_{s}=T\,N\,C\)

Pool model representation

Flux description

Figure 1
Figure 1: Pool model representation

Output fluxes

\(D: D\cdot\left(- SUE + 1\right)\)
\(M: M\cdot\left(- K_{r}\cdot k_{t} + SUE\cdot k_{m} + k_{t}\right)\)
\(E: E\cdot k_{l}\)

Internal fluxes

\(S > D: E\cdot k_{d}\)
\(D > M: D\cdot\left(- K_{e} + SUE\right)\)
\(D > E: D\cdot K_{e}\)
\(M > D: K_{r}\cdot M\cdot k_{t}\)

The right hand side of the ODE

\(\left[\begin{matrix}- E\cdot k_{d}\\- D + E\cdot k_{d} + K_{r}\cdot M\cdot k_{t}\\D\cdot\left(- K_{e} + SUE\right) + M\cdot\left(- SUE\cdot k_{m} - k_{t}\right)\\D\cdot K_{e} - E\cdot k_{l}\end{matrix}\right]\)

The Jacobian (derivative of the ODE w.r.t. state variables)

\(\left[\begin{matrix}0 & 0 & 0 & - k_{d}\\0 & -1 & K_{r}\cdot k_{t} & k_{d}\\0 & - K_{e} + SUE & - SUE\cdot k_{m} - k_{t} & 0\\0 & K_{e} & 0 & - k_{l}\end{matrix}\right]\)

Steady state formulas

\(S = S\)
\(D = 0\)
\(M = 0\)
\(E = 0\)

Steady states (potentially incomplete), according jacobian eigenvalues, damping ratio

Parameter set: Set 1

\(S: S\), \(D: 0.0\), \(M: 0.0\), \(E: 0.0\)

\(\lambda_{1}: 0.000\)
\(\lambda_{2}: 0.010-0.000j\)
\(\rho_{2}: -1.000000\)
\(\lambda_{3}: -0.022+0.000j\)
\(\rho_{3}: 1.000000\)
\(\lambda_{4}: -1.054-0.000j\)
\(\rho_{4}: 1.000000\)

References

Schimel, J. P., & Weintraub, M. N. (2003). The implications of exoenzyme activity on microbial carbon and nitrogen limitation in soil: A theoretical model. Soil Biology and Biochemistry, 35(4), 549–563.