title: Differential nutrient limitation of soil microbial biomass and metabolic quotients (qCO2): is there a biological stoichiometry of soil microbes?
abstract: BACKGROUND
  Variation in microbial metabolism poses one of the greatest current uncertainties in models of global carbon cycling, and is particularly poorly understood in soils. Biological Stoichiometry theory describes biochemical mechanisms linking metabolic rates with variation in the elemental composition of cells and organisms, and has been widely observed in animals, plants, and plankton. However, this theory has not been widely tested in microbes, which are considered to have fixed ratios of major elements in soils.
  
  
  RESULTS
  To determine whether Biological Stoichiometry underlies patterns of soil microbial metabolism, we compiled published data on microbial biomass carbon (C), nitrogen (N), and phosphorus (P) pools in soils spanning the global range of climate, vegetation, and land use types. We compared element ratios in microbial biomass pools to the metabolic quotient qCO2 (respiration per unit biomass), where soil C mineralization was simultaneously measured in controlled incubations. Although microbial C, N, and P stoichiometry appeared to follow somewhat constrained allometric relationships at the global scale, we found significant variation in the C∶N∶P ratios of soil microbes across land use and habitat types, and size-dependent scaling of microbial C∶N and C∶P (but not N∶P) ratios. Microbial stoichiometry and metabolic quotients were also weakly correlated as suggested by Biological Stoichiometry theory. Importantly, we found that while soil microbial biomass appeared constrained by soil N availability, microbial metabolic rates (qCO2) were most strongly associated with inorganic P availability.
  
  
  CONCLUSIONS
  Our findings appear consistent with the model of cellular metabolism described by Biological Stoichiometry theory, where biomass is limited by N needed to build proteins, but rates of protein synthesis are limited by the high P demands of ribosomes. Incorporation of these physiological processes may improve models of carbon cycling and understanding of the effects of nutrient availability on soil C turnover across terrestrial and wetland habitats.
authors.affiliation: Duke University Wetland Center, Nicholas School of the Environment, Duke University, Durham, North Carolina, United States of America. wyatt.hartman@duke.edu
authors.forename: Wyatt H
authors.initials: WH
authors.lastname: Hartman
authors.forename: Curtis J
authors.initials: CJ
authors.lastname: Richardson
mesh_terms: D018533:Biomass; D002244:Carbon / Q000032:analysis / Q000378:metabolism; D057486:Carbon Cycle; D002245:Carbon Dioxide / Q000378:metabolism; D017753:Ecosystem; D008954:Models, Biological; D009584:Nitrogen / Q000032:analysis / Q000378:metabolism; D010758:Phosphorus / Q000032:analysis / Q000378:metabolism; D012987:Soil / Q000737:chemistry; D012988:Soil Microbiology
chemical_list: D012987:Soil; D002245:Carbon Dioxide; D010758:Phosphorus; D002244:Carbon; D009584:Nitrogen