Microorganisms are the most abundant lifeform on Earth, mediating global fluxes of matter and energy. Over the past decade, high-throughput molecular techniques generating multiomic sequence information (DNA, mRNA, and protein) have transformed our perception of this microcosmos, conceptually linking microorganisms at the individual, population, and community levels to a wide range of ecosystem functions and services. Here, we develop a biogeochemical model that describes metabolic coupling along the redox gradient in Saanich Inlet-a seasonally anoxic fjord with biogeochemistry analogous to oxygen minimum zones (OMZs). The model reproduces measured biogeochemical process rates as well as DNA, mRNA, and protein concentration profiles across the redox gradient. Simulations make predictions about the role of ubiquitous OMZ microorganisms in mediating carbon, nitrogen, and sulfur cycling. For example, nitrite "leakage" during incomplete sulfide-driven denitrification by SUP05 Gammaproteobacteria is predicted to support inorganic carbon fixation and intense nitrogen loss via anaerobic ammonium oxidation. This coupling creates a metabolic niche for nitrous oxide reduction that completes denitrification by currently unidentified community members. These results quantitatively improve previous conceptual models describing microbial metabolic networks in OMZs. Beyond OMZ-specific predictions, model results indicate that geochemical fluxes are robust indicators of microbial community structure and reciprocally, that gene abundances and geochemical conditions largely determine gene expression patterns. The integration of real observational data, including geochemical profiles and process rate measurements as well as metagenomic, metatranscriptomic and metaproteomic sequence data, into a biogeochemical model, as shown here, enables holistic insight into the microbial metabolic network driving nutrient and energy flow at ecosystem scales.M icrobial communities catalyze Earth's biogeochemical cycles through metabolic pathways that couple fluxes of matter and energy to biological growth (1). These pathways are encoded in evolving genes that, over time, spread across microbial lineages and today shape the conditions for life on Earth. High-throughput sequencing and mass-spectrometry platforms are generating multiomic sequence information (DNA, mRNA, and protein) that is transforming our perception of this microcosmos, but the vast majority of environmental sequencing studies lack a mechanistic link to geochemical processes. At the same time, mathematical models are increasingly used to describe local-and global-scale biogeochemical processes or predict future changes in global elemental cycling and climate (2, 3). Although these models typically incorporate the catalytic properties of cells, they fail to integrate the information flow from DNA to mRNA, proteins, and process rates as described by the central dogma of molecular biology (4). Hence, a mechanistic framework integrating multiomic data with geochemical information has...