Photosynthesis, essential for plant productivity and global food security, is vulnerable to compound abiotic stresses like high light, drought, heat, and salinity. These disrupt Photosystem II (PSII) and Photosystem I (PSI), causing efficiency losses and yield declines. We reframe vulnerabilities via architectural asymmetries (rapid D1 turnover in PSII vs. slow Fe–S repair in PSI) and multifaceted protective mechanisms, including non-photochemical quenching (NPQ) subtypes (qE, qT, qZ, qH, qI), cyclic electron transport (CEF), and photosynthetic control (PhotCon). Mapping "ROS geography" emphasizes acceptor side over reduction (Mehler-driven Fe–S damage) and donor side imbalances (¹O₂-mediated P700 oxidation), with metrics like Y(NA), Y(ND), and EPR for phenotyping. Using cryo-EM and genetic models, we link mechanisms to assays (e.g., NPQ relaxation kinetics) to resolve debates, such as Y(ND)'s sufficiency for PhotCon. Based on evolutionary principles, we propose a hierarchical prevention framework from sensing to proteostasis and advocate synthetic plastid engineering with digital twins and optogenetic CEF/NPQ switches. This empowers development of antifragile crops adapting to climate variability, enhancing food security.
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