Certain foraminiferal species are abundant within the chemocline of marine sediments. Ultrastructurally, most of these species possess numerous peroxisomes complexed with the endoplasmic reticulum; mitochondria are often interspersed among these complexes. In the Santa Barbara Basin, pore-water bathing Foraminifera and co-occurring sulfur-oxidizing microbial mats had micromolar levels of hydrogen peroxide, a reactive oxygen species that can be detrimental to biological membranes. Experimental results indicate that adenosine triphosphate concentrations are significantly higher in Foraminifera incubated in 16 μM H 2 O 2 than in specimens incubated in the absence of H 2 O 2 . New ultrastructural and experimental observations, together with published results, lead us to propose that foraminiferans can utilize oxygen derived from the breakdown of environmentally and metabolically produced H 2 O 2 . Such a capability could explain foraminiferal adaptation to certain chemically inhospitable environments; it would also force us to reassess the role of protists in biogeochemistry, especially with respect to hydrogen and iron. The ecology of these protists also appears to be tightly linked to the sulfur cycle. Finally, given that some Foraminifera bearing peroxisome-endoplasmic reticulum complexes belong to evolutionarily basal groups, an early acquisition of the capability to use environmental H 2 O 2 could have facilitated diversification of foraminiferans during the Neoproterozoic.
KeywordsAnoxia; Beggiatoa; Blake Ridge; cold seep; hydrogen peroxide; hydrogen sulfide; microbial mat; Neoproterozoic; Santa Barbara Basin; Soledad BasinThe anoxic-oxic interface and associated reduction-oxidation (redox) zone are sites at which a great variety of biogeochemical reactions occur (e.g. Brune, Frenzel, and Cypionka 2000;Hulth et al. 2005;Preisler et al. 2007). In organic-rich sedimentary settings where the overlying waters are depleted of oxygen, redox gradients are particularly steep (e.g. Røy, Huettel, and Jørgensen 2005), and hydrogen sulfide can often reach considerable concentrations within one centimeter of the aerated sediment surface (e.g. Bernhard, (Tapley, Buettner, and Shick 1999). ROS are also formed as byproducts of normal metabolic reactions in most organisms (reviewed in Genestra 2007). Prolonged ROS exposure can seriously damage many biological molecules such as proteins, lipids, and nucleic acids, but cells normally have protective mechanisms by which to minimize the effect of such oxidative stress (e.g. Abele et al. 1998;Genestra 2007;Lesser 2006;Sigler et al. 1999). Because H 2 O 2 is an uncharged molecule, it is less reactive than most ROS and can cross biological membranes via diffusion and/or through channel proteins such as aquaporins ( This conversion typically is mediated by catalase, a metalloenzyme containing four porphyrin heme (iron) groups that react with hydrogen peroxide, which often appears as a crystalline core in peroxisomes (e.g. Masters and Crane 1995).Foraminifera (rhizarian protis...