Chloroplast protein translocation complexes and their regulation

Chloroplasts, refined through more than a billion years of evolution in plants and algae, act as highly efficient and resilient converters of solar energy. Additionally, these organelles function as complex anabolic factories, synthesizing a wide array of primary and secondary metabolites. The functionality of chloroplasts is dependent on the involvement of more than 3,000 proteins, the majority of which are encoded by the nuclear genome. These nucleus-encoded proteins must cross the chloroplast double lipid membrane to become functional. This translocation process is facilitated by the translocons at the outer and inner envelope membranes of chloroplasts (the outer chloroplast [TOC] and the inner chloroplast [TIC] complexes, respectively) and is driven by an energy-providing motor. Despite decades of research, the composition of these complexes remains highly controversial, especially regarding the TIC and motor components. However, recent studies have provided valuable insight into the TOC/TIC complexes, while also raising new questions about their mechanisms. In this review, we explore the latest advancements in understanding the structure and function of these complexes. Additionally, we briefly examine the processes of protein quality control, retrograde signaling, and discuss promising directions for future research in this field.