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The key enzymatic components of the PCO N-degron pathway (PCO, ATE, and PRT6) likely evolved in the last common ancestor of eukaryotes. Cys2-ERFs arose independently multiple times in land plants, with ERFVIIs evolving O2-dependent degradability in vascular plants. In bryophytes, Cys2-ERF proteins belong to groups X, VII (mosses), and IV (liverworts). In ferns, the extended N-terminal sequence of ERFVIIs (the CMVII motif) shows greater variability compared with spermatophytes and lycophytes. ERFVII-like proteins are also found in liverworts, although these proteins lack the Cys residue at the second position. The recruitment of HRPE and GCC-boxcis-elementsin promoters of hypoxia-regulated genes is a feature specific to tracheophytes.
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