Plastid-derived Type II fatty acid biosynthetic enzymes in chromists

Fatty acid biosynthesis is a critical process for living organisms, but the evolution of the enzymes involved in this pathway is poorly understood. Animals and fungi use a Type I fatty acid synthase (FAS), a large multifunctional protein found in the cytosol. Bacteria use a Type II complex, where each enzymatic domain is a discrete polypeptide. In plants, fatty acid biosynthesis takes place in the plastid, and utilises a Type II enzyme complex. Recently, the apicomplexan parasites Plasmodium and Toxoplasma have been shown to contain the plastid-targeted Type II FAS. To investigate the distribution of this pathway, we have characterised two Type II enzymes, FabD and FabI, in three other eukaryotes with plastids derived from red algal endosymbionts: cryptomonads, heterokonts, and haptophytes. Collectively, these are referred to as chromists, and are thought to be related to apicomplexa and their relatives. Phylogenies of these enzymes show that the plastid Type II FAS enzymes are found in all groups studied, which most likely means that they originated from the red algal endosymbiont at the outset of the secondary endosymbiosis of their plastids. In addition, although plastid fab D genes are clearly related to one another, they are not related to cyanobacterial homologues, as would be expected. On the other hand, the strongly supported plastid fab I clade is related to cyanobacteria, and contains genes from chlamydiales.