Environmental regulation and disruption of shell biomineralization in bivalves and gastropods: a mechanistic review

Molluscan shells are increasingly threatened by a complex array of environmental stressors—climate-related drivers, changes in seawater chemistry, contaminants, and biotic cues. This review provides a mechanistic synthesis of how these stressors disrupt the biomineralization process across life stages. We develop a conditional hierarchical response framework from the reviewed evidence. The framework links ionoregulatory and acid-base disturbances with molecular, microstructural, and shell-level responses, while allowing for compensatory, nonlinear, species-specific, and life-stage-specific outcomes. This molecular disruption manifests as defects in calcium carbonate (CaCO3) crystalline microstructure, which may in turn compromise shell macro-scale physical properties (strength, hardness, growth). We synthesize combined and context-dependent effects of co-occurring stressors and distinguish formally tested interactions from qualitative comparisons of combined treatments. Future research must leverage integrated multi-omics, advanced in vivo imaging, and multi-stressor experimental designs to unravel candidate molecular and physiological thresholds and predict adaptive potential. This framework may help identify testable mechanisms, evidence gaps, and context-dependent vulnerabilities relevant to conservation.