Chloroplast
Primary endosymbiosis

Somewhere between 1 and 2 billion years ago, a free-living cyanobacterium entered an early eukaryotic cell, either as food or as an internal parasite, but managed to escape the phagocytic vacuole it was contained in. The two innermost lipid-bilayer membranes that surround all chloroplasts correspond to the outer and inner membranes of the ancestral cyanobacterium's gram negative cell wall, and not the phagosomal membrane from the host, which was probably lost. The new cellular resident quickly became an advantage, providing food for the eukaryotic host, which allowed it to live within it. Over time, the cyanobacterium was assimilated, and many of its genes were lost or transferred to the nucleus of the host. From genomes that probably originally contained over 3000 genes only about 130 genes remain in the chloroplasts of contemporary plants. Some of its proteins were then synthesized in the cytoplasm of the host cell, and imported back into the chloroplast (formerly the cyanobacterium). Separately, somewhere about 90–140 million years ago, it happened again and led to the amoeboid Paulinella chromatophora.

This event is called endosymbiosis, or "cell living inside another cell with a mutual benefit for both". The external cell is commonly referred to as the host while the internal cell is called the endosymbiont.

Chloroplasts are believed to have arisen after mitochondria, since all eukaryotes contain mitochondria, but not all have chloroplasts. This is called serial endosymbiosis —an early eukaryote engulfing the mitochondrion ancestor, and some descendants of it then engulfing the chloroplast ancestor, creating a cell with both chloroplasts and mitochondria.

Whether or not primary chloroplasts came from a single endosymbiotic event, or many independent engulfments across various eukaryotic lineages, has long been debated. It is now generally held that organisms with primary chloroplasts share a single ancestor that took in a cyanobacterium 600–2000 million years ago. It has been proposed this the closest living relative of this bacterium is Gloeomargarita lithophora. The exception is the amoeboid Paulinella chromatophora, which descends from an ancestor that took in a Prochlorococcus cyanobacterium 90–500 million years ago.

These chloroplasts, which can be traced back directly to a cyanobacterial ancestor, are known as primary plastids (" plastid " in this context means almost the same thing as chloroplast). All primary chloroplasts belong to one of four chloroplast lineages—the glaucophyte chloroplast lineage, the amoeboid Paulinella chromatophora lineage, the rhodophyte (red algal) chloroplast lineage, or the chloroplastidan (green) chloroplast lineage. The rhodophyte and chloroplastidan lineages are the largest, with chloroplastidan (green) being the one that contains the land plants.
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