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Formation of lithified micritic laminae in modern marine stromatolites
This study investigated the formation of lithified micritic laminae in modern marine stromatolites in the Bahamas through biogeochemical and microbial analyses. The research found that cyanobacterial photosynthesis, sulfate reduction by bacteria, and anaerobic sulfide oxidation cause calcium carbonate precipitation and formation of lithified layers, while aerobic respiration and aerobic sulfide oxidation cause calcium carbonate dissolution. Specifically, layers with the highest biomass and rates of sulfate reduction and sulfide oxidation correlated with lithified micritic horizons in the stromatolites. The study concludes that sulfur cycling driven by these microbial processes is responsible for lamination and early lithification in the Bahamian stromatolites.
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Formation of lithified micritic laminae in modern marine stromatolites
- 1. GEOL533 - Carbonatesand Evaporites Assignment 2 - Paper Presentation Formation of lithified micritic laminae in modern marine stromatolites (Bahamas): The role of sulfur cycling Omar Atef Radwan PhD Student – Geosciences Dept.
- 2. Visscher, P. T.,Reid, R. P., Bebout, B. M., Hoeft, S. E., Macintyre, I. G., & Thompson, J. A. (1998). Formation of lithified micritic laminae in modern marine stromatolites (Bahamas): the role of sulfur cycling. American Mineralogist, 83(11), 1482-1493. 2 Reid, R. P., Visscher, P. T., Decho, A. W., Stolz, J. F., Bebout, B. M., Dupraz, C., ... & Steppe, T. F. (2000). The role of microbes in accretion, lamination and early lithification of modern marine stromatolites. Nature, 406(6799), 989-992. Cited by 156 documents Cited by 396 documents
- 3. Outline • Previously • Background •Literature review • Research Gap • Objective • Approach • Outcomes • Implications • Summary 3
- 4.
- 5. Background • Stromatolites: laminated sedimentarystructures produced by the activities of benthic microbial mats • Earth’s oldest macrofossils • dominate the fossil record for 85% of Earth’s history • record the interactions of biological and geological processes throughout the 3.5 billion year history of life on Earth Reid et al., 2000 5 James & Jones, 2015
- 6. Background • Photosynthesis of cyanobacteriahas been linked to CaCO3 precipitation • Aerobic respiration by heterotrophic bacteria results in CaCO3 dissolution 6
- 7. Background 7 • Reduction ofsulfate yields HCO3 - and HS- and results in CaCO3 precipitation • The sulfide is oxidized in chemolithotrophic respiration, either with O2 (aerobic) or with NO3 (anaerobic)
- 8. Literature review • discoveredin the early 1980s • (Dravis 1983). • found at numerous locations • (Dill et al. 1986; Reid and Browne 1991; Reid et al., 1995). • modern analogs of ancient stromatolitic microbial communities? • Guerrero Negro, Mexico (Canfield and Des Marais 1991) • Solar Lake, Sinai (Jørgensen and Cohen 1977, Krumbein et al. 1977) • Texel, The Netherlands (Visscher and Van Gemerden 1993) Visscher et al., 1998 8
- 9. Research Gap • Althoughmost researchers agree that, “microbial mats and their associated sediments must be lithified early in order to be preserved in the record as stromatolites” • the proposed mechanisms and precise timing of early lithification have been "vigorously debated" 9
- 10. Objective to assess therelative importance of • Photosynthesis • aerobic respiration • sulfate reduction • sulfide oxidation in stromatolite formation. 10
- 11. Approach Visscher et al.,1998 11 • Highborne Cay was chosen as the study site because stromatolites at this locality exhibit exceptionally well- developed lamination. Early observations: • the mat is a prokaryotic community dominated by the cyanobacteria Schizothrix sp and Solentia Sp.
- 12. Approach Bergman et al.,2010 Visscher et al., 1998 Map showing stromatolite locations on the margins of Exuma Sound 1. Highborne Cay, study site for this paper 2. Schooner Cays (Dravis 1983) 3. Lee Stocking Island (Dill et al. 1986) 4. Stocking Island (Reid and Browne 1991; Macintyre et al. 1996; Steneck et al. 1997) Unnumbered sites (Reid et al. 1995) Map of Highborne Cay, showing the location of the stromatolites (xxxx) in a fringing reef along the eastern margin. 12
- 13. Approach Measurement of biogeochemicalgradients Determination of the activities of the metabolic reactions To examine the role of different functional groups of bacteria in the formation of lithified micritic horizons in Exuma stromatolites • oxygenic phototrophic bacteria (Eq. 1) • aerobic heterotrophic microbes (Eq. 2) • sulfate-reducing bacteria (Eq. 3) • sulfide-oxidizing bacteria (Eq. 4) 13
- 14. • a rangeof geological, microbial and chemical analyses • O2, sulphide, pH needle electrodes • Physicochemical indices • light, scanning electron, transmission electron, and scanning laser confocal • microstructural features • epifluorescence microscopy • microbial populations Approach Visscher et al., 1998 14
- 15. • Optical microscopyshows that the mat is a prokaryotic community dominated by the cyanobacteria Schizothrix sp. • Schizothrix filaments are: • abundant in Layers 1 • common in Layers 3 and 5 • scarce in Layers 2 and 4 • Endolithic cyanobacteria are also abundant in Layer 3 Outcomes 15 Reid et al., 2000
- 16. • A petrographicthin section shows that: • Layers 3 and 5 are micritic horizons with the characteristic features of lithified layers in Exuma stromatolites. • Thin micrite crusts overlie micritized grains. 16 Outcomes Visscher et al., 1998
- 17. Outcomes Biogeochemical gradients • Concentrationprofiles of O2 and HS- in the stromatolite mat showed distinct diel fluctuations. Depth distribution of O2 (filled box), HS- (filled diamond) and pH (filled circle) 17Visscher et al., 1998
- 18. Outcomes • Depth distributionof Oxygen • 6:00am: 2 mm • 1:00pm: 4–5 mm • 6:00pm: 3 mm • 2:00am: 0.75–1 mm 18Visscher et al., 1998
- 19. Outcomes • Depth distributionof Sulfide • 6:00am: 2.5 mm • 1:00pm: 7 mm • 6:00pm: 3.5 mm • 2:00am: 1 mm 19Visscher et al., 1998
- 20. Outcomes 20Visscher et al.,1998 • Gradients of pH showed less diel variability than the O2 and HS- profiles.
- 21. Outcomes • Rate ofphotosynthesis • maximum primary production of O2 at 12:30 p.m. at the interface of Layers 1 and 2 21Visscher et al., 1998
- 22. Outcomes • Aerobic respirationrate • maximum rates of aerobic respiration occurred in early afternoon, reaching values of O2 just below the interface of Layers 1 and 2 22Visscher et al., 1998
- 23. Outcomes • Sulfate reductionrates • during the day and in the night show that the highest rates are in Layer 3, regardless of whether O2 was present or absent 23 Visscher et al., 1998 • The rate of HS- oxidation • difficult to measure because of formation of many different reaction products. Some of these products (e.g., S2O3 -2 ) are rapidly reduced back to HS- by sulfate-reducing bacteria. However, based on the location of the oxycline, the rate of HS- oxidation is expected to peak in Layer 3.
- 24. Outcomes 24 • distinct variationsin the abundance of sulfate-reducing and sulfide- oxidizing bacteria in individual layers of the stromatolite mat: • Maximum population densities of sulfate-reducing bacteria (8·105 cells/cm3) and sulfide- oxidizing bacteria (2·104 cells/cm3) are found within lithified Layer 3. • Indeed, sulfate-reducing bacteria were two orders of magnitude more abundant in Layer 3 than in any other layer.
- 25. Implications • Lithified micritichorizons are correlated with layers of high biomass. • Microbial processes (biomass activities) within these stratified mats produce distinctive patterns of lithification as follows: (1) Photosynthesis and respiration (2) Sulfate reduction (3) Sulfide oxidation 25
- 26. Implications 26 PS = photosynthesis AR= aerobic respiration SR = sulfate reduction ASO = aerobic sulfide oxidation NSO = denitrifying sulfide oxidation (anaerobic respiration) Arrow lengths indicate depth zones over which the respective processes are active Arrow widths indicate the relative importance of the processes light arrows are associated with CaCO3 dissolution Dark arrows are indicative of CaCO3 precipitation Visscher et al., 1998
- 27. Implications (1) Photosynthesis and respiration highin Layers 1 (high biomass) and 2 (low biomass), causing precipitation and dissolution of CaCO3. results in little or no net lithification in these layers, depending on the amount of organic carbon produced by photosynthesis that is used for aerobic respiration. 27 Visscher et al., 1998
- 28. Visscher et al.,1998 Implications (2) Sulfate reduction high in Layer 3 (high biomass), where it causes CaCO3 precipitation. results in a lithified layer in which carbonate sand grains are cemented together by micritic precipitates. The depth to the top of Layer 3 is controlled by: the amount of photosynthetic production of {CH2O} by cyanobacteria in Layer 1 the amount of consumption of {CH2O} by sulfate-reducing bacteria in Layer 3. 28
- 29. Visscher et al.,1998 Implications (3) Sulfide oxidation at the oxic-anoxic interface at the top of Layer 3 may have a twofold effect: coupled processes of dissolution and precipitation associated with aerobic and anaerobic HS- oxidation may result in: etching and truncation of previously microbored grains the precipitation of hard, micritic crusts; these crusts resemble micritic laminae found in ancient stromatolites. 29
- 30. Summary • This isthe first study to define a specific set of mechanisms that link lamination in marine stromatolites to a dynamic balance between sedimentation, a succession of prokaryotic communities and early lithification. • The findings indicate a close correlation between dynamic sulfur cycling and the formation of lithified micritic laminae in stromatolite-forming microbial mats in the Exuma Cays. 30
- 31. Summary • Cyanobacterial photosynthesis,sulfate reduction, and anaerobic sulfide oxidation in stromatolitic mats at Highborne Cay are responsible for CaCO3 precipitation, whereas aerobic respiration and aerobic sulfide oxidation cause CaCO3 dissolution. • photosynthesis coupled to sulfate reduction and sulfide oxidation is more important than photosynthesis coupled to aerobic respiration in the formation of lithified micritic laminae in Highborne Cay stromatolites. 31
- 32. References • Bergman, K.L., Westphal, H., Janson, X., Poiriez, A., & Eberli, G. P. (2010). Controlling parameters on facies geometries of the Bahamas, an isolated carbonate platform environment. In Carbonate Depositional Systems: Assessing Dimensions and Controlling Parameters (pp. 5-80). Springer Netherlands. • Reid, R. P., Visscher, P. T., Decho, A. W., Stolz, J. F., Bebout, B. M., Dupraz, C., ... & Steppe, T. F. (2000). The role of microbes in accretion, lamination and early lithification of modern marine stromatolites. Nature, 406(6799), 989-992. • Visscher, P. T., Reid, R. P., Bebout, B. M., Hoeft, S. E., Macintyre, I. G., & Thompson, J. A. (1998). Formation of lithified micritic laminae in modern marine stromatolites (Bahamas): the role of sulfur cycling. American Mineralogist, 83(11), 1482-1493. 32
- 33.
- 34.
- 35.
- 36. • Chemolithotrophic • Oxycline •microautoradiography 36
Editor's Notes
- #6 Before the Cambrian diversification of life, laminated carbonate build-ups called stromatolites were widespread in shallow marine seas. These ancient structures are generally thought to be microbial in origin. Little is known about stromatolite formation, especially the relative roles of microbial and environmental factors in stromatolite accretion.
- #7 Microbial processes are well known to cause precipitation and dissolution of CaCO3.
- #9 Modern stromatolites forming in open ocean water of normal marine salinity were first discovered in the Schooner Cays, on the east margin of Exuma Sound, Bahamas (Fig. 1), in the early 1980s (Dravis 1983). Since then, they have been found at numerous locations throughout the Exuma Cays, on the west margin of Exuma Sound (Fig. 1; Dill et al. 1986; Reid and Browne 1991; Reid et al. 1995). Modern microbial mats such as those found in Guerrero Negro, Mexico (Canfield and Des Marais 1991), Solar Lake, Sinai (Jørgensen and Cohen 1977; Krumbein et al. 1977), Sabkha Gavish, Sinai (Krumbein et al. 1979), and Texel, The Netherlands (Visscher and Van Gemerden 1993), are commonly viewed as modern analogs of ancient stromatolitic microbial communities (Ward et al. 1989; Des Marais 1990). However, an important difference between these mats and ancient stromatolites is that the former are unlithified, whereas ancient stromatolites formed as actively mineralizing structures. Although the mats at Solar Lake (Lyons et al. 1984), Sabkha Gavish (Krumbein et al. 1979; Gavish et al. 1985), and Guerro Negro (J. Farmer, personal communication) contain diffuse precipitates of CaCO3, lithified micritic crusts, the characteristic features of ancient stromatolites, do not form in these mats. This raises the question why benthic microbial mats building ancient stromatolites and stromatolites in the Exuma Cays form lithified micritic laminae, whereas others do not.
- #12 Some previous studies have focused on textural differences between Exuma and ancient stromatolites, suggesting that the modern forms, which are sandy, are inappropriate analogs for ancient stromatolites, which are typically micritic (e.g., Awramik and Riding 1988; Riding et al. 1991; Riding 1994). These studies, however, overlooked a fundamental similarity between the modern and ancient structures: Lamination in both reflects periodic formation of lithified micritic horizons. Moreover, the micritic crusts in Exuma stromatolites are remarkably similar in thickness to micritic laminae in many ancient stromatolites (e.g., Walter 1983; Bertrand Sarfati 1976; Monty and Mas 1981). Exuma stromatolites thus offer a unique opportunity to investigate the formation of lithified micritic laminae in stromatolites forming in a modern marine environment. Initial observations indicating that lithified micritic laminae in Exuma stromatolites form within microbial mats at the surface of these structures led to the present study, which examines microbial processes involved in this lithification process. Solentia
- #13 Bahamian system LBB = Little Bahama Bank GBB = Great Bahama Bank AC = Acklins AI = Andros Island BI = Berry Islands BM= Bimini CC = Cat Cay CI = Cat Island CR = Crooked Island CSB = Cay Sal Bank ELI = Eleuthera Island EI = Exuma Islands GAI = Great Abaco Island GBI = Great Bahama Island IN = Great and Little Inagua JC = Joulters Cays LI = Long Island MA = Mayaguana MO = Mouchoir NP = New Providence SS = San Salvador Oceanographic features BBE = BlakeBahama Escarpment ES = Exuma Sound FS = Florida Straits NEPC = Northeast Providence Channel NWPC = Northwest Providence Channel OBC = Old Bahama Channel SC = Santaren Channel TOTO = Tongue of the Ocean WP = Windward Passage
- #15 Mat communities and microstructural features were identified using a variety of microscope techniques (light, scanning electron, transmission electron, and scanning laser confocal29) and microbial populations were enumerated using epifluorescence microscopy counts This study combined a range of geological, microbial and chemical analyses. An extensive field program was conducted during January and June 1997, and March and August 1998. Physicochemical indices of stromatolite mats were determinedin situ, primarily with O2, sulphide, pH needle electrodes (0.8 mm outer diameter)9,
- #16 Cycling between communities, indicated by large arrows, is a response to intermittent sedimentation (see text). a, b, Pioneer community: filamentous cyanobacteria (arrows) bind carbonate sand grains. c–e , Bacterial biofilm community: a continuous sheet of amorphous exopolymer (arrows, c, d) with abundant heterotrophic bacteria (Fig. 3) forms uppermost surface; aragonite needles precipitate within this surface film (e). f, g, Climax community: a surface biofilm overlies filamentous cyanobacteria and endolith-infested grains, which appear grey and are fused (arrow, f). Precipitation in tunnels that cross between grains leads to welding (g). a, c, f, Petrographic thin sections, plane polarized light; cyanobacteria are stained with methylene blue. b, d, e, g, Scanning electron microscope images. Scale bars: a, b, c,f, 100 m; d, 50 m; e, 5 m; g, 10 m
- #18 Measurements were taken on June 14–15, 1997. Ambient light intensity and was 30 mE/ m2/s (6:00 a.m.), 1760 mE/m2/s (1:00 p.m.), 210 mE/m2/s (6:00 p.m.), and 0 mE/m2/s (2:00 a.m.); the pH of ambient seawater is 7.9.
- #21 Gradients of pH within the stromatolite mat (Fig. 4) showed less diel variability than the O2 and HS- profiles. Surface values of pH ;7.5 were slightly lower than ambient sea water, with a pH of 7.9 From late afternoon until early morning, pH values gradually increased with depth mat pH progressively increased to values of about 8–8.2 at 5 mm depth The lower pH values at the surface of the stromatolite may be associated with production of organic acids through excretion by phototrophs during photorespiration and formation of H2SO4 from HS- oxidation A major departure from this trend of gradual increase of pH with depth was observed in early afternoon. At this time, a peak in pH of 8.7 occurred just below the depth of maximum O2 concentration and coinciding with the top of lithified Layer 3. During this early afternoon period, pH values at depths of 1 mm to 8 mm were higher than the pH of ambient sea water. High rates of photosynthetic CO2 fixation and perhaps sulfate reduction (i.e., removal of H2SO4) could have contributed to this rise in pH. pH values at 3 mm, which is the top of Layer 3, ranged from 7.85 at night to 8.75 during the early afternoon.
- #23 showed patterns of daytime variability that were similar to those seen in the O2 and photosynthesis profiles:
- #27 Schematic representation showing the spatial and temporal distribution of microbial processes resulting in precipitation and dissolution of carbonate (see Eqs. 1–4)