Setaphyte VERY-LONG-CHAIN FATTY ACYL DESATURASES Impact Glycerolipid and Sphingolipid Metabolism.

Desaturases in plants are diverse. They vary in localization, source of reducing power, and substrate preference, accepting glycerolipids, long-chain bases, acyl-CoAs, and acyl-ACPs, in varying states of (un)saturation and chain length. Their products are incorporated into membrane glycerolipids, sphingolipids, or storage lipids. We previously characterized a desaturase from Physcomitrium patens that predominantly affects the monounsaturation of very-long-chain fatty acyl (VLCFA) moieties of sphingolipids, naming this desaturase SPHINGOLIPID FATTY ACYL DESATURASE (SFD). Among embryophytes, candidate SFDs were only identified in setaphytes, including one paralog in P. patens and an ortholog in Marchantia polymorpha. Here, we characterize the P. patens paralog, and clarify via mutant analysis that SFDs affect not only sphingolipid metabolism, but also glycerolipid metabolism. We express both paralogs, as well as the candidate gene from M. polymorpha, in Saccharomyces cerevisiae, and show they desaturate VLCFAs incorporated into sphingolipids, triacylglycerols, and acyl-CoAs. The simplest explanation is that "SFDs" likely accept an acyl-CoA, rather than a sphingolipid substrate as initially proposed. We suggest renaming these desaturases VERY-LONG-CHAIN FATTY ACYL DESATURASES (VFADs). The physiological functions of VFADs and functionally similar enzymes from other plant systems are discussed, as are the challenges with classifying desaturases.