Growth and survival in a changing environment: Combined effects of moderate hypoxia and low pH on juvenile bivalve Macoma balthica

Abstract

Baltic Sea species live in a complex, variable environment characterized by highly fluctuating hydrology, including large seasonal and diel pH variations. For decades, oxygen deficiency caused by anthropogenic eutrophication has affected the Baltic Sea, and large areas of the seafloor are permanently hypoxic resulting in severely degraded benthic communities. Species living in this system are thus potentially tolerant and adapted to this fluctuating environment, but also vulnerable as illustrated by high mortality of benthic species as a result of hypoxia. In the future, the frequency and extent of regularly co-occurring low oxygen and low pH levels will likely increase with on-going climate change. A key species in the Baltic Sea soft-bottom communities, the bivalve Macoma balthica (L.), experiences such conditions throughout its life-cycle, and therefore serves as a good model organism for studying the combined effects of oxygen and pH conditions. To study the response of M. balthica to multiple changes occurring in the benthic environment, we conducted an experiment to investigate the survival and shell growth of newly settled juveniles simultaneously exposed to two pH levels (7.85 and 7.35) and two oxygen levels (8.5 and 3.0mg/l) for 29days in a fully factorial design. Survival was high in all treatments (>60%), but significantly higher in the two low oxygen treatments (>70%). Although positive growth was observed in all treatments, pH and oxygen as well as their interaction significantly affected relative growth. The highest growth was observed in the “low O2/high pH” treatment, which was 2.4 times higher than in both treatments with high oxygen. Although the mechanism for these differences remains unknown, hypoxia-induced metabolic depression likely plays a role. Our results highlight the need to know more about the occurrence and performance of benthic species regularly exposed to changing conditions, and of the range and conditions encountered in situ. With the predicted future negative changes in oxygen availability as well as pH, the adaptive responses of benthic species to multiple stressors will be critical in understanding ecosystem dynamics in the face of change.