Abstract. Chloroplast and mitochondria, the two organelles with an accepted endosymbiotic origin, have developed multiple translocation pathways to ensure the subcellular allocation of proteins synthesized by cytosolic ribosomes, and to guarantee their assembly into functional complexes in coordination also with organellar-encoded subunits. The evolution of different protein import machineries was thus essential for the development of these two organelles within cells. A general overview of the translocation machineries in chloroplast and mitochondrial membranes involved in targeting and import of nuclearencoded proteins, with special focus on plant cells where the two organelles coexist, is expounded.Keywords. Protein translocation, mitochondria, chloroplasts, thylakoids, dual-targeting, endosymbiotic organelles.
Endosymbiosis and the plant cellIt is widely accepted that mitochondria, found in eukaryotic cells, and plastids, just in plant cells, are organelles of endosymbiotic origin [1]. Two independent endosymbiotic events gave rise to the evolution of these organelles in a plant cell. First, mitochondria evolved from a single endosymbiotic event between a host cell and an aerobic a-proteobacterium. As a consequence, a diversity of heterotrophic eukaryotic lineages emerged [2]. Later, plastids were incorporated from a free-living photosynthetic prokaryote, an ancestor of contemporary cyanobacteria, which was engulfed by a heterotrophic eukaryotic host that already contained mitochondria. The primary endosymbiotic event in plastid evolution gave rise to glaucophyta, rhodophyta and viridiplantae lineages [3]. Subsequent endosymbiotic events in plastid evolution have occurred [4]. With time, the majority of the genetic information from the endosymbionts was lost or transferred to the nucleus of the host cell, though some amount of information is still retained by the organelles, thus functioning as a semiautonomous system [5]. Because of the gene relocation in the host cell, several mechanisms were developed to re-target and efficiently transport the proteins acting in the organelles but now expressed in the host cytosol. Some of these machineries were adapted from bacteria; others show