ROXY1 and ROXY2, two Arabidopsis glutaredoxin genes, are required for anther development

Glutaredoxins (GRXs) are small oxidoreductases that are involved in various cellular processes and play a crucial role in responses to oxidative stress. Three GRX subgroups exist in plants, and GRXs with active sites of the CPYC and CGFS types are common to pro- and eukaryotes. In contrast, GRXs with the CC type motif have so far only been identified in land plants. Here, we report that the two CC-type GRXs ROXY1 and ROXY2 together control anther development in Arabidopsis thaliana. Single roxy1 and roxy2 mutants are fertile and produce normal anthers. However, roxy1 roxy2 double mutants are sterile and do not produce pollen. Strikingly, abaxial and adaxial anther lobe differentiation are differently affected, with early lobe differentiation being defective in the adaxial lobes, whereas later steps during pollen mother cell differentiation are disrupted in the abaxial lobes. Expression studies show that ROXY1 and ROXY2 are expressed with overlapping patterns during anther development. Lack of ROXY1 and ROXY2 function affects a large number of anther genes at the transcriptional level. Genetic and RT-PCR data imply that ROXY1/2 function downstream of the early-acting anther gene SPOROCYTELESS/NOZZLE and upstream of DYSFUNCTIONAL TAPETUM1, controlling tapetum development. Mutagenesis of a conserved glutathione-binding glycine in the ROXY1 protein indicates that CC-type GRXs need to interact with glutathione to catalyze essential biosynthetic reactions. Analysis of these two novel anther genes indicates that redox regulation, as well as participating in plant stress defense mechanisms, might play a major role in the control of male gametogenesis.