Microbial dormancy improves development and experimental validation of ecosystem model Gangsheng Wang1,2, Sindhu Jagadamma1,2, Melanie A. Mayes1,2, Christopher W. Schadt2,3, J. Megan Steinweg2,3,4, Lianhong Gu1,2, Wilfred M. Post1,2 1 Environmental Sciences Division, Oak Ridge National Laboratory, Oak Ridge, TN 37831, USA 2 Climate Change Science Institute, Oak Ridge National Laboratory, Oak Ridge, TN 37831, USA 3 Biosciences Division, Oak Ridge National Laboratory, Oak Ridge, TN 37831, USA 4 Currently at Biological Sciences Department, University of Wisconsin-Baraboo/Sauk County, Baraboo, WI 53913, USA Supplementary Information Correspondence: Gangsheng Wang Bldg 4500N, F129-S, MS-6301 Oak Ridge National Laboratory Oak Ridge, TN 37831-6301 Tel: (865)576-6685; Fax: (865)574-9501 Email: wangg@ornl.gov 1 MEND model Fig. 1 in the main text Diagram of the Microbial-ENzyme-mediated Decomposition (MEND) model. Soil organic carbon pools include: particulate organic carbon (POC) (e.g., lignocellulose-like compounds and starch-like compounds, denoted by P1 and P2, respectively), mineral-associated organic carbon (MOC, denoted by M), dissolved organic carbon (DOC, D), adsorbed phase of DOC (QOC, Q), active microbial biomass (BA), dormant microbial biomass (BD), POC degraded enzymes (e.g., EP1 and EP2 that break down P1 and P2, respectively), and MOC-degraded enzymes (EM). IP1, IP2, and ID are external inputs to the pools of P1, P2, and D, respectively. Transformation fluxes include: (1) POC1 (P1) decomposition (denoted by the flux F1 in equations in the Supplementary Information), (2) POC2 (P2) decomposition (F2), (3) MOC (M) decomposition (F3), (4, 5) adsorption (F4) and desorption (F5) between DOC (D) and QOC (Q), (6) DOC (D) uptake by BA (F6), (7,8) dormancy (F7) and reactivation (F8) between BA and BD, (9, 10) BA growth respiration (F9) and maintenance respiration (F10), (11) BD maintenance respiration (F11), (12) BA mortality (F12), (13) synthesis of EP1 (F13,EP1), EP2 (F13,EP2), and EM (F13,EM), and (14) turnover of enzymes (F14,EP1, F14,EP2, and F14,EM). 2 The dynamics of each soil carbon pool (see Fig. 1) in the Microbial-ENzyme-mediated Decomposition (MEND) model can be described as dP1 I P1 (1 g D ) F12 F1 dt (S1) dP2 I P 2 F2 dt (S2) dM (1 f D ) F1 F2 F3 dt (S3) dQ F4 F5 dt (S4) dD I D f D F1 F2 g D F12 F3 F14, EP1 F14, EP 2 F14, EM F6 ( F4 F5 ) dt (S5) dBA F6 ( F7 F8 ) F9 F10 F12 F13, EP1 F13, EP 2 F13, EM dt (S6) dBD ( F7 F8 ) F11 dt (S7) dEP1 F13, EP1 F14, EP1 dt (S8) dEP2 F13, EP 2 F14, EP 2 dt (S9) dEM F13, EM F14, EM dt (S10) dCO2 F9 F10 F11 dt (S11) d ( P1 P2 M Q D BA BD EP1 EP2 EM ) I P1 I P 2 I D F9 F10 F11 dt (S12) where the state variables (C pools) are described in Table S1 and Fig. 1; Eq. S11 indicates the total heterotrophic respiration flux and Eq. S12 expresses the overall mass balance of the system. The transformation fluxes (Fig. 1) are elucidated by Eqs. S13–S26 in Table S1. 3 Table S1 Component fluxes in the MEND model (parameters are described in Table 1 in the main text) Flux description Particulate organic carbon (POC) pool 1 (P1) decomposition (F1) Equation F1 VP1 EP1 P1 K P1 P1 (S13) POC pool 2 (P2) decomposition F2 VP 2 EP2 P2 K P 2 P2 (S14) F3 VM EM M KM M (S15) Mineral-associated organic carbon (MOC, M) decomposition Adsorption (F4) and desorption (F5) between dissolved organic carbon (DOC, D) and adsorbed DOC (QOC, Q) DOC (D) uptake by microbes Dormancy (F7) and reactivation (F8) between active (MBA) and dormant (MBD) microbial biomass (BA and BD) MBA (BA) growth respiration (F9) and maintenance respiration (F10) F4 K ads 1 Q / Qmax D F5 K des (Q / Qmax ) F6 1 VD mR D BA YG KD D F7 1 D /( K D D) mR BA F8 D /( K D D) mR BD 1 V BA D F9 1 D YG K D D 1 m BA D F10 1 R YG KD D Synthesis of enzymes for P1 (EP1, F13,EP1), enzymes for P2 (EP2, F13,EP2), and enzymes for M (EM, , F13,EM) F11 mR BD F12 (1 pEP pEM ) mR BA F13,EP1 = P1 / (P1 + P2 )× pEP × mR × BA F13,EP2 = P2 / (P1 + P2 )× pEP × mR × BA Turnover of enzymes (EP1, EP2, EM) F14,EP1 rE EP1 MBD (BD) maintenance respiration MBA (BA) mortality (S16) (S17) (S18) (S19) (S20) (S21) (S22) (S23) (S24) (S25) F13, EM pEM mR BA F14,EP 2 rE EP2 (S26) F14, EM rE EM Notes: Italic symbols like Fi represent component fluxes in equations. Italic symbols P1, P2, M, Q, D, BA, BD, EP1, EP2, and EM are state variables (soil carbon pools) in equations. 4 Table 1 in the main text Microbial-ENzyme-mediated Decomposition (MEND) model parameters Parameter Description Apriori range Eq Units VP1 Maximum specific decomposition rate for P1 (0.1, 3) S13 mg C mg−1 C h−1 VP2 Maximum specific decomposition rate for P2 (1, 50) S14 mg C mg−1 C h−1 KP1 Half-saturation constant for P1 decomposition (1, 100) S13 mg C g−1 soil KP2 Half-saturation constant for P2 decomposition (1, 100) S14 mg C g−1 soil VM Maximum specific decomposition rate for M (0.05, 2) S15 mg C mg−1 C h−1 KM Half-saturation constant for M decomposition (10, 1000) S15 mg C g−1 soil VD Maximum specific uptake rate of D for growth (0.0001, 0.5) S18 mg C mg−1 C h−1 KD Half-saturation constant for uptake of D (0.05, 0.5) S18 mg C g−1 soil mR Specific maintenance rate of BA = VD ∙ α /(1− α) S18 mg C mg−1 C h−1 α = mR /( VD + mR) (0.01, 0.5) β Ratio of dormant maintenance rate to mR 0.001 S23 — YG True growth yield (0.1, 0.9) S18 — fD Fraction of decomposed P1 and P2 allocated to D (0.1, 1) S3 — gD Fraction of dead BA allocated to D (0.01, 1) S1 — pEP Fraction of mR for production of EP1 and EP2 (0.001, 0.1) S25 — pEM Fraction of mR for production of EM (0.001, 0.1) S25 — rE Turnover rate of EP1, EP2, and EM (0.0001, 0.01) S26 mg C mg−1 C h−1 Qmax Maximum DOC sorption capacity (0.5, 5) S16 mg C g−1 soil KBA Binding affinity (1, 16) Kdes Desorption rate (0.0001, 0.1) Kads Specific adsorption rate = Kdes ∙ KBA LCF0 r0 — (mg C g−1 soil) −1 S17 mg C g−1 soil h−1 S16 mg C mg−1 C h−1 Initial fraction of P1 = P1/(P1+P2) (0.1, 0.95) — Initial active fraction = BA/(BA+BD) (0.01, 1) — Notes: The column “Eq” lists the major equation # (see Supplementary Information) in which each parameter is used. 5 Figure S1 Comparison between observed (Obs) and simulated (Sim) microbial biomass carbon (MBC) in three soils (Gelisol, Andisol, Ultisol) with addition of Glucose or Starch. MEND and MEND_wod denote the MEND model with and without dormancy, respectively. The error bars are standard deviations estimated by the Critical Objective Function Index (COFI) method. Sample sizes are 2559, 2672, and 2460 for the three soils, respectively. 6 7 8 Figure S2 Boxplots of MEND model parameter values derived by the Critical Objective Function Index (COFI) method. Parameters are described in Table 1. The bottom and top of the box denote the 25th and 75th percentile (the lower and upper quartiles), respectively. The band near the middle of the box is the 50th percentile (median). The small square in the box is the mean value. The ends of the whiskers represent the lowest datum within 1.5 IQR (interquartile range) of the lower quartile and the highest datum within 1.5 IQR of the upper quartile. The data beyond the two ends of the whiskers might be considered outliers. Different letters above the boxes indicate the parameter samples originate from significantly different distribution at a significance level of 0.05 according to the Kruskal-Wallis (KW) test. Sample sizes are 2346–2672 for the four soils. 9 10 11 12 13 14 15 16 17 18