Strong influence of the littoral zone on sedimentary lipid biomarkers in a meromictic lake.

Planktonic sulfur bacteria growing in zones of photic zone euxinia (PZE) are important primary producers in stratified, sulfur-rich environments. The potential for export and burial of microbial biomass from anoxic photic zones remains relatively understudied, despite being of fundamental importance to interpreting the geologic record of bulk total organic carbon (TOC) and individual lipid biomarkers. Here we report the relative concentrations and carbon isotope ratios of lipid biomarkers from the water column and sediments of meromictic Mahoney Lake. The data show that organic matter in the central basin sediments is indistinguishable from material at the lake shoreline in both its lipid and carbon isotopic compositions. However, this material is not consistent with either the lipid profile or carbon isotope composition of biomass obtained directly from the region of PZE. Due to the strong density stratification and the intensive carbon and sulfur recycling pathways in the water column, there appears to be minimal direct export of the sulfur-oxidizing planktonic community to depth. The results instead suggest that basinal sediments are sourced via the littoral environment, a system that integrates an indigenous shoreline microbial community, the degraded remains of laterally rafted biomass from the PZE community, and detrital remains of terrigenous higher plants. Material from the lake margins appears to travel downslope, traverse the strong density gradient, and become deposited in the deep basin; its final composition may be largely heterotrophic in origin. This suggests an important role for clastic and/or authigenic minerals in aiding the burial of terrigenous and mat-derived organic matter in euxinic systems. Downslope or mineral-aided transport of anoxygenic, photoautotrophic microbial mats may have been a significant sedimentation process in early Earth history.

Coupled reductive and oxidative sulfur cycling in the phototrophic plate of a meromictic lake.

Mahoney Lake represents an extreme meromictic model system and is a valuable site for examining the organisms and processes that sustain photic zone euxinia (PZE). A single population of purple sulfur bacteria (PSB) living in a dense phototrophic plate in the chemocline is responsible for most of the primary production in Mahoney Lake. Here, we present metagenomic data from this phototrophic plate--including the genome of the major PSB, as obtained from both a highly enriched culture and from the metagenomic data--as well as evidence for multiple other taxa that contribute to the oxidative sulfur cycle and to sulfate reduction. The planktonic PSB is a member of the Chromatiaceae, here renamed Thiohalocapsa sp. strain ML1. It produces the carotenoid okenone, yet its closest relatives are benthic PSB isolates, a finding that may complicate the use of okenone (okenane) as a biomarker for ancient PZE. Favorable thermodynamics for non-phototrophic sulfide oxidation and sulfate reduction reactions also occur in the plate, and a suite of organisms capable of oxidizing and reducing sulfur is apparent in the metagenome. Fluctuating supplies of both reduced carbon and reduced sulfur to the chemocline may partly account for the diversity of both autotrophic and heterotrophic species. Collectively, the data demonstrate the physiological potential for maintaining complex sulfur and carbon cycles in an anoxic water column, driven by the input of exogenous organic matter. This is consistent with suggestions that high levels of oxygenic primary production maintain episodes of PZE in Earth's history and that such communities should support a diversity of sulfur cycle reactions.