Review: Regulated polyhydroxyalkanoate granule biology and polymer phenotype in mangrove-associated bacteria
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Abstract. Darmawan MT, Alviansyah MH, Aditya RD, Fadhila NH, Safika NJ, Salma O, Nugraheni R, Hidayati RK, Nadhira S, Setyawan AD. 2025. Review: Regulated polyhydroxyalkanoate granule biology and polymer phenotype in mangrove-associated bacteria. Cell Biol Dev 9: 129-157. Polyhydroxyalkanoates (PHAs) are biodegradable biopolymers whose molecular weight, composition, and dispersity strongly determine material performance and industrial applicability. Despite extensive efforts to optimize culture conditions and genetic constructs, reproducible control of PHA polymer phenotype remains challenging. This review reinterprets PHA accumulation from a cell-biological perspective by positioning PHA granules as regulated intracellular organelles rather than passive repositories of excess carbon. Synthesizing evidence from mechanistic studies, culture-based experiments, and gene-centric analyses, the review demonstrates that polymer phenotype-including molecular-weight distribution, structural heterogeneity, and batch-to-batch reproducibility-emerges as a physiological output shaped by regulatory gating and granule state. PHA accumulation is shown to be an active and reversible storage strategy embedded within global stress-response networks, rather than a simple consequence of metabolic overflow. Granule-associated proteins, spatial organization, and turnover dynamics act as critical intermediaries translating regulatory state into material properties. The review further highlights that widespread reliance on production-oriented metrics, such as polymer yield or percentage of cell dry weight, obscures intracellular processes that govern polymer history and quality, contributing to inconsistencies across studies. Mangrove-associated bacteria are used as a biologically informative stress-context model to illustrate how conserved intracellular storage mechanisms operate under recurring physiological constraints, without invoking environmental determinism. Building on these insights, a regulatory-granule-phenotype framework is proposed that reframes reproducibility as a biological property dependent on granule-state stability rather than a purely technical issue. By foregrounding granule-level control, this synthesis provides a coherent foundation for integrating cellular regulation with polymer phenotype and outlines experimental directions for achieving more predictable and interpretable PHA production.
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