Abstract
A growing number of T2/S-RNases are being discovered in plant genomes. Members of this protein family have a variety of known functions, but the vast majority are still uncharacterized. We present data and analyses of phylogenetic relationships among T2/S-RNases, and pay special attention to the group that contains the female component of the most widespread system of self-incompatibility in flowering plants. The returned emphasis on the initially identified component of this mechanism yields important conjectures about its evolutionary context. First, we find that the clade involved in self-rejection (class III) is found exclusively in core eudicots, while the remaining clades contain members from other vascular plants. Second, certain features, such as intron patterns, isoelectric point, and conserved amino acid regions, help differentiate S-RNases, which are necessary for expression of self-incompatibility, from other T2/S-RNase family members. Third, we devise and present a set of approaches to clarify new S-RNase candidates from existing genome assemblies. We use genomic features to identify putative functional and relictual S-loci in genomes of plants with unknown mechanisms of self-incompatibility. The widespread occurrence of possible relicts suggests that the loss of functional self-incompatibility may leave traces long after the fact, and that this manner of molecular fossil-like data could be an important source of information about the history and distribution of both RNase-based and other mechanisms of self-incompatibility. Finally, we release a public resource intended to aid the search for S-locus RNases, and help provide increasingly detailed information about their taxonomic distribution.
Highlights
One half of all flowering plant species strictly enforce outcrossing
We recover three distinct clades of T2/S-type ribonuclease (S-RNase) (Fig. 1 and Fig. S1), which mirror the previously described three ‘classes’ (e.g., Igić & Kohn, 2001; Steinbachs & Holsinger, 2002). These clades are present in the consensus and maximum credibility trees derived from MrBayes posterior tree sets, as well as the best-scoring RAxML maximum likelihood trees, inferred using both nucleotide and amino acid alignments and substitution models
Several well-characterized T2/S-RNases belong to class I, such as Arabidopsis thaliana RNS1 and RNS3 (Bariola et al, 1994; Bariola, MacIntosh & Green, 1999; Hillwig et al, 2008; Hillwig et al, 2011; LeBrasseur et al, 2002; Nishimura et al, 2014), Nicotiana glutinosa RNase NW and RNase NT (Kariu et al, 1998; Hino, Kawano & Kimura, 2002; Kawano et al, 2006; Kurata et al, 2002), and Solanum lycopersicum RNase LE and RNase LX (Groß, Wasternack & Köck, 2004; Jost et al, 1991; Köck et al, 2004; Köck, Stenzel & Zimmer, 2006; Lers et al, 1998; Lers et al, 2006; Löffler et al, 1992; Nürnberger et al, 1990; Tanaka et al, 2000)
Summary
One half of all flowering plant species strictly enforce outcrossing. A great variety of such mechanisms fall under a single umbrella term—self-incompatibility (SI) systems. Despite their documented presence across angiosperms, it remains unclear how, if at all, the many interacting components of such systems are related. Dozens of independently evolved molecular mechanisms appear to cause SI, but only a few are genetically characterized or studied in great detail How to cite this article Ramanauskas and Igić (2017), The evolutionary history of plant T2/S-type ribonucleases.
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