RNA editing is a process of modifying nucleotide bases, resulting in a chemically altered RNA molecule (Ye et al., 2020). In many plants, site-specific editing of RNA transcripts is carried out by pentatricopeptide repeat (PPR) proteins (Okuda et al., 2009). PPR proteins are broadly classified into P type (canonical 35 amino acid motif) and PLS type (P motif, long (L) and S (short) motifs) subfamilies based on their domain structure (Cheng et al., 2016). PLS subfamily of proteins are further classified into various subgroups (PLS (example S type), E1, E2, E+ and DYW) (Cheng et al., 2016). PLS type DYW subgroup proteins are involved in the site-specific binding and editing of RNA (Schallenberg-Rüdinger et al., 2013). To date, more than thousand editing sites have been identified in club moss and hornworts (Lenz and Knoop, 2013), and it has been determined that the cytidine deaminase like conserved C terminal DYW domain in DYW subgroup is responsible for the RNA editing activity (Lenz and Knoop, 2013). While most plant PPR editing proteins are known to edit cytidine to uridine (C to U), very recently reverse editing (uridine to cytidine) has been discovered to occur in ferns, lycophytes, and hornworts (Lenz and Knoop, 2013). To date, there is no experimental structural information for the DYW:KP domain, and the structural architecture, flexibility, and dynamics of the intact PPR-DYW proteins remain unsolved. Therefore, through structural studies we aim to determine the three-dimensional structures of S-type PPR proteins, and DYW-domain and DYW:KP domain containing proteins, to understand the molecular mechanism of action. Furthermore, we also aim to understand the details of RNA binding specificity in S type proteins and how it impacts the conformation of the protein. In order to obtain the required information, wild-type and designer proteins will be expressed and purified, and subjected to appropriate structural methods including computational prediction, X-ray crystallography, cryogenic transmission electronic microscopy (cryo EM) and small angle x-ray scattering (SAXS). RNA-binding and editing assays will be used to evaluate activity of target proteins and rationally designed mutants.