Plant plastids: from evolutionary origins to functional specialization and organelle interactions

Plastids are highly diverse organelles that play critical roles in supporting many forms of life on Earth. Among them, chloroplasts house the machinery for photosynthesis, providing phototrophic capabilities to eukaryotes such as plants, algae, and photosynthetic protists. The functions of plastids are indispensable for the survival and development of life, and they are widely recognized as endosymbiotic organelles with a single origin. They exhibit morphological diversity, tissue specificity, and the ability to adapt to specific cellular functions. Despite this level of understanding, significant questions remain unanswered, such as how genetic material from the endosymbiont was transferred and integrated into the host nucleus, the timeline for the full integration of the endosymbiont into the host cell, and the processes by which plastids specialized and adapted to various cell types. While plastids have unique features and specialized roles, they are neither autonomous nor physically isolated. Instead, they interact with other sub-cellular compartments through yet-to-be-characterized membrane domains or specialized structures. This review explores the origin and evolution of plastids, their protein-import machinery, compartmentalization, and interactions with other cellular compartments, and highlights key unanswered questions in these areas.The origins, genome evolution, protein-import machinery, and communication mechanisms of plastids within cells reveal how their internal organization and connections with other compartments support essential life processes beyond photosynthesis.