Abstract
Ferns are the most primitive of all vascular plants. One of the characteristics distinguishing them from flowering plants is its triterpene metabolism. Most cyclic triterpenes in ferns are hydrocarbons derived from the direct cyclization of squalene by squalene cyclases (SCs). Both ferns and more complex plants share sterols and biosynthetic enzymes, such as cycloartenol synthases (CASs). Polystichum belongs to Dryopteridaceae, and is one of the most species-rich of all fern genera. Several Polystichum ferns in Japan are classified as one of three possible chemotypes, based on their triterpene profiles. In this study, we describe the molecular cloning and functional characterization of cDNAs encoding a SC (PPH) and a CAS (PPX) from the type species Polystichum polyblepharum. Heterologous expression in Pichia pastoris revealed that PPH and PPX are hydroxyhopane synthase and CAS, respectively. By using the PPH and PPX sequences, we successfully isolated SC- and CAS-encoding cDNAs from six Polystichum ferns. Phylogenetic analysis, based on SCs and oxidosqualene cyclase sequences, suggested that the Polystichum subclade in the fern SC and CAS clades reflects the chemotype—but not the molecular phylogeny constructed using plastid molecular markers. These results show a possible relation between triterpenes and their biosynthetic enzymes in Polystichum.
Highlights
Ferns are the most primitive of all vascular plants
Because ferns lie between bacteria and flowering plants on the evolutionary scale, they produce triterpenes that are similar to those produced by bacteria and flowering plants
A DNA fragment encoding the putative squalene cyclases (SCs) was obtained by reverse-transcription PCR (RT-PCR) by using the degenerate primers
Summary
Ferns are the most primitive of all vascular plants. They produce specialized characteristic metabolites that are not found in flowering plants [1]. These include triterpenoids and phloroglucinols with various biological activities [2,3,4]. Most ferns produce triterpene hydrocarbons by the cyclization of squalene, a C30 acyclic substrate (1). At the gene/enzyme level, the sterol pathway might be conserved between ferns and flowering plants. The study of ferns might be useful in investigating the molecular evolution of triterpene biosynthesis from bacterial to flowering plants
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