Survival, ATP pool, and ultrastructural characterization of benthic foraminifera from Drammensfjord (Norway): response to anoxia

Abstract

While much evidence indicates that certain benthic foraminifera are facultative anaerobes, little is known regarding the physiologic response of foraminifera to anoxia. In order to assess their response, specimens of four foraminiferal species, collected from a typically dysoxic area of Drammensfjord, Norway (45 m water depth), were incubated in seawater purged with nitrogen. Over a time course of > 3 weeks, the specimens were extracted for adenosine triphosphate (ATP) in a nitrogen-flushed glove bag to assess their survival and ATP reserve under such conditions. For comparative purposes, similar extractions were done on conspecifics one week after their collection from the seafloor, as well as on other conspecifics, obtained from the same site, incubated in aerated conditions. The survival rates of nitrogen-treated Adercotryma glomeratum, Psammosphaera bowmanni, and Stainforthia fusiformis were not significantly lower than those of the control specimens. However, the ATP concentrations of nitrogen-incubated A. glomeratum and S. fusiformis were significantly lower than those of their aerated conspecifics, while there was no significant difference between the [ATP] of P. bowmanni from the two treatments. Both the survival rate and the ATP concentrations of nitrogen-incubated Bulimina marginata were significantly lower than those of control specimens. The ultrastructure of B. marginata and S. fusiformis incubated in N 2 for 18 days were compared with those of specimens fixed within 15 minutes of collection. For both species, the specimens that survived the experimental treatment had ultrastructures indistinguishable from those fixed just after field collection. However, the ultrastructure of B. marginata differed from that of S. fusiformis in that it lacked the numerous peroxisome-endoplasmic reticulum (ER) complexes and what appeared to be algal chloroplasts observed in S. fusiformis. Copious arrays of paracrystals were observed in both species from the experimental treatment as well as the shipboard-fixed specimens, suggesting that neither population had extensive pseudopodial networks. When considered in combination, our results indicate that the four species respond to and survive anoxia differently, with responses including dormancy and, as yet unidentified, anaerobic metabolic pathways.