Elevated extracellular pH during early shell formation in the blue mussel Mytilus edulis

Marine calcifiers are amongst the most vulnerable organisms to ocean acidification (OA). However, limited studies investigate the mechanisms underlying their hindered performance under OA stress. Working with larval stages of the blue mussel, Mytilus edulis, we use microsensors to study the pH and calcium conditions necessary for shell deposition. Using 45-48 hour, D-veliger stages, we discover alkaline conditions with respect to ambient seawater pH by 0.28 pH units and higher calcium concentrations (by 0.54mM) in the extra pallial space beneath the growing shell that likely promotes the rapid synthesis of the first shell. We further use enzyme assays in combination with immuno-stainings of sodium-potassium ATPase (NKA) and proton ATPase (VHA) to provide information on the major ion regulatory pathways that enable transport of calcium carbonate required for shell formation and pH homeostasis. We also use the juvenile stages of M. edulis to understand how extracellular pH regulation close to the shell formation site will be influenced by OA stress. This allows us to describe the pH dependency of early shell formation and to begin to develop a model of the ion regulatory network that facilitates biomineralisation in the organism. The results are discussed in the context of environmental change and consequences for mollusc developmental success.