Plastid origin: 110 years since MereschkowskyPlastids are eukaryotic metabolic compartments responsible for photosynthesis [1] and a variety of metabolic functions including the biosynthesis of amino acids [2], nucleotides [3], lipids [4], and cofactors [5]. The importance of plastids to photosynthetic lineages of eukaryotes cannot be overstated. In 1905, Mereschkowsky proposed a fully articulated version of endosymbiotic theory positing that plastids originated from cyanobacteria that came to reside as symbionts in eukaryotic cells [6,7]. The theory did not make its way into mainstream biological thinking until it was revived in a synthesis by Margulis [8,9] that also incorporated Wallin's [10] -and Paul Portier's (published in French, cited in Sapp [11]) -ideas about endosymbiotic theory for mitochondrial origin, yet adorned by Margulis' own suggestion of a spirochaete origin of flagella. The spirochaete story never took hold, leaving endosymbiotic theory with three main players: the plastid, the mitochondrion, and its host, which is now understood to be an archaeon [12]. Well into the 1970s, resistance to the concept of endosymbiosis for the origin of plastids (and mitochondria) was stiff [13][14][15].With the availability of protein and DNA sequences, of which Mereschkowsky knew nothing, strong evidence had accumulated by the early 1980s that plastids originated through endosymbiosis from cyanobacteria, rather than autogenously [16]. However, there have been recurrent suggestions, starting with Mereschkowsky [6] and tracing into the 1970's [17], that there were several independent origins of plastids from cyanobacteria. Today it is widely, but not universally [18,19], accepted that plastids had a single origin. The strongest evidence for that view is that the protein import machinery, a good marker for endosymbiotic events [20], in the three lineages of Archaeplastida -eukaryotes with primary plastids [21] -consists of homologous host-originated components [22][23][24], which would not be the case had plastids in those lineages arisen from independent cyanobacterial symbioses.Genomics has enriched our understanding of plastid origin. We now know that the plastid genome has undergone extreme reduction while leaving its imprints in the nuclear genome through endosymbiotic gene transfer [25,26] and that plastids have spread across eukaryotes through symbiosis and become secondary and tertiary plastids [27]. Genomic data also supports the case for a single plastid origin. Despite some concerns about incomplete sampling and phylogenetic artifacts [18], recent analyses from different groups, incorporating improved phylogenetic algorithms and sampling of cyanobacteria, provide two lines of evidence, in addition to * Corresponding author. Email: bill@hhu.deHandling Editor: Andrzej Bodył INVITED REVIEW Acta Soc Bot Pol 83(4): 281-289 DOI: 10.5586/asbp.2014.045 Received: 2014-11-20 Accepted: 2014-12-04 Published electronically: 2014 Plastid origin: who, when and why?Chuan Ku, Mayo Roettger, Verena Zimorski, Shijulal...