A Model Study of the Effects of Sulfide-Oxidizing Bacteria
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View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Helsingin yliopiston digitaalinen arkisto Boreal environment research 16: 167–184 © 2011 issn 1239-6095 (print) issn 1797-2469 (online) helsinki 30 June 2011 a model study of the effects of sulfide-oxidizing bacteria (Beggiatoa spp.) on phosphorus retention processes in hypoxic sediments: implications for phosphorus management in the Baltic sea sepehr shakeri Yekta* and lars rahm Department of Thematic Studies — Water and Environmental Studies, Linköping University, SE-581 83 Linköping, Sweden (corresponding author’s e-mail: [email protected]) Received 14 June 2010, accepted 23 Nov. 2010 (Editor in charge of this article: Heikki Seppä) Yekta, s. s. & rahm, l. 2011: a model study of the effects of sulfide-oxidizing bacteria (Beggiatoa spp.) on phosphorus retention processes in hypoxic sediments: implications for phosphorus management in the Baltic sea. Boreal Env. Res. 16: 167–184. Ongoing eutrophication increases phosphorus storage in surficial sediments of the Baltic Sea which can then be released during hypoxic/anoxic events. Such sediments are suit- able habitats for sulfide-oxidizing bacteria,Beggiatoa spp. The objective of this paper is to investigate the effects of these bacteria on the P retention processes in hypoxic sediments using a diagenetic model. This model simulates interactions of the processes controlling P mobility in the sediments with redox reactions from the Beggiatoa metabolism. Modeling results demonstrate that P retention capability is limited when dissolved iron is mineralized as iron sulfides in the sediments. In this regard, sulfide consumption by Beggiatoa spp. potentially decreases the rate of iron sulfide formation and consequently increases the P retention capability in local-scale sediment. Introduction during the middle of the 1990s, mild windy winters led to an increased mixing of the water Surficial sediments of the Baltic Sea contain column, less hypoxic/anoxic bottoms, and a amounts of phosphorus (P) comparable to many drastic drop in P concentration in the water mass years of riverine loads (Conley et al. 2002). (Gustafsson and Stigebrandt 2007). This inspired This P source can be released during hypoxic/ the development of new remedy processes by anoxic events and in a longer perspective may improving the oxygen conditions in the bottom fuel nitrogen (N)-fixing cyanobacteria blooms layers and thereby increasing P retention in the and render remedy actions based on reduction sediments based on increased concentrations of of N loads less successful (Cloern 2001). In Fe(III) oxides (Mortimer 1941). Knowledge of fact, the portion of hypoxic/anoxic sediments the processes at the sediment water interface increased not only relatively but also quantita- regulating the hypoxic/anoxic sediments is cru- tively during the recent decades in the Baltic cial in these attempts. Sea. This increase seems to be a global phe- Hypoxic/anoxic bottoms eliminate higher nomenon (Diaz and Rosenberg 2008). However, orders of life but make the new sediment sur- 168 Yekta & Rahm • Boreal env. res. Vol. 16 face a suitable habitat for sulfide-oxidizing Dissolution of iron oxide sheets to Fe2+ in the bacteria, Beggiatoa spp. (Rosenberg and Diaz top layers of the sediments caused by shifts from 1993). Beggiatoa cells build aggregates in sedi- oxic to hypoxic/anoxic conditions in bottom ments rich in organic matter with a gradient water, changes the sorption capability of the sed- of sulfide across the sediment water interface iments and leads to mobilization of iron-bound (Mußmann et al. 2003). These bacteria are che- phosphate which can be transferred to overlying moautotrophs, gaining metabolic energy by water columns (Conley et al. 2002). oxidizing hydrogen sulfide (Jørgensen 1982). To understand the effects of Beggiatoa mats Widespread Beggiatoa mats have been observed on P retention processes, we developed a dia- in the Stockholm Archipelago (Rosenberg and genetic model for reactive diffusive transport Diaz 1993), in anoxic sediments of the Gulf of of substances to simulate major biogeochemical Finland (Vallius 2006), and in huge areas of the processes in the sediments like those in the Baltic eastern Gotland basin (P. Hall, Gothenburg Uni- Sea with both Beggiatoa-free and Beggiatoa- verity, pers. comm.). The Danish Limfjorden, the covered sediment surfaces. The Beggiatoa-free German Wadden Sea and Eckenförde Bay are sediment model couples biotic processes such other examples where large populations of these as organic matter degradation pathways with bacteria have been observed (Mußmann et al. inorganic processes including iron oxide precipi- 2003, Preisler et al. 2007). tation/dissolution, iron sulfide precipitation, and Beggiatoa spp. is well-known for protecting P adsorption/desorption. Organic matter degra- overlying water from toxic sulfide diffused from dation pathways are the main contributors to the sulfidic layers of the sediments. Beggiatoa redox disequilibrium in the sediments (Tromp metabolism including their physiology and enzy- et al. 1995), while inorganic processes are the mology have been studied in detail (e.g. Jør- major chemical pathways controlling mobility gensen 1977, Hagen and Nelson 1997, Mußmann of P in marine sediments (Jensen et al. 1995, et al. 2003, Sayama et al. 2005, Hinck et al. Carman and Rahm 1997, Gunnars et al. 2002, 2007, Kamp et al. 2008). Furthermore, the role of Conley et al. 2002, Lehtoranta and Pitkänen these bacteria in the cycling of elements has been 2003, Lehtoranta et al. 2008). In the next step, the topic of numerous studies. As an example, we added microbially mediated redox reactions Fenchel and Bernard (1995) investigated major from the Beggiatoa metabolism to the model in microbial processes such as nitrification/denitrifi- order to simulate their interactions with sedi- cation and cycling of N in a Beggiatoa inhabited mentary processes. The development and appli- sediment. Preisler et al. (2007) studied sulfide cation of diagenetic models for the investigation oxidization by Beggiatoa mats and its interac- of sedimentary processes can be found in e.g. tions with manganese and iron in local scale sedi- Berner (1964), Boudreau (1997), and Boudreau ment. Most of the previous studies emphasized and Jørgensen (2001). the importance of these bacteria in the cycling The current model simulates the key P reten- of elements in marine environments, but to our tion processes in natural sediments based on knowledge, none of the earlier studies investi- observations and field measurements of water gated the impact of Beggiatoa spp. on P retention and sediments of the Baltic Sea. Through mode- processes in their growth medium. ling, we do not intend to reproduce the observed Phosphorus (P) retention processes in the data but to provide a realistic numerical rep- sediments of the Baltic Sea have been amply resentation of the sedimentary processes. The researched. For instance, Brügmann et al. goal of the study is to investigate our conceptual (1992), Sternbeck and Sohlenius (1997), Hille understanding of the impact of Beggiatoa mats et al. (2005), and Leipe et al. (2008) have all on the P retention processes in sediments by contributed to knowledge in this area. Previous comparing two conventional diagenetic models studies showed that mobility of P in the sedi- simulating typical Baltic Sea hypoxic sediments ment–porewater medium is chiefly regulated by which differ only in the presence or absence of adsorption of phosphate on iron oxyhydroxides. Beggiatoa mat. Boreal env. res. Vol. 16 • Effects of Beggiatoa spp. on phosphorus in sediments 169 Beggiatoa filaments at Material and methods Mixed water the sediment surface We developed a 1D diagenetic model to simulate Metabolic activities DBL the vertical distribution of solute and solid com- – 2+ 2+ z Porewater: O2, NO3 , Mn , Fe , 2– + 3– pounds in local scale sediment. In doing this, SO4 , NH4 , H2S, PO4 we treated sedimentary solute and solid species Organic and inorganic redox (Fig. 1) as dependent variables and considered reactions, adsorption/desorption, their mass conservation equations in an arbitrary precipitation/dissolution sediment system. We included chemical reaction Sediment Organic and mineral particles, pathways in conservation equations as sources organic carbon, Fe(OH)3, MnO2, FeS or sinks to simulate dynamic interactions of dis- Fig. 1. model conceptualization and incorporated bio- solved and solid compounds. Boudreau and Jør- chemical processes in the modeled sediment column. z gensen (2001) discussed derivation of mass con- represents sediment depth. servation equations for sedimentary substances and biogeochemical processes in detail. 1997) in order to compare simulated and meas- ured concentration profiles and adjusted some Model conceptualization model parameters to approach the measured pro- files. This provided us with a numerical tool for Model conceptualization including incorporated further investigations of the P retention processes biochemical reactions is illustrated in Fig. 1. The as well as the effects of Beggiatoa mats on them. model simulates an arbitrary vertical column of a sediment–porewater system with a given geometry. The present model simulates the dis- Physical model properties tribution of dissolved and solid compounds from the mixed water body downwards across the The physical properties of the model are listed diffusive boundary layer (DBL) and into the in Table 1. We fixed