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Background  RACK1 is a versatile scaffold protein in mammals, regulating diverse developmental processes. Unlike in non-plant organisms where RACK1 is encoded by a single gene, Arabidopsis genome contains three  RACK1  homologous genes, designated as  RACK1A, RACK1B  and  RACK1C , respectively. Previous studies indicated that the loss-of-function alleles of  RACK1A  displayed multiple defects in plant development. However, the functions of  RACK1B  and  RACK1C  remain elusive. Further, the relationships between three  RACK1  homologous genes are unknown.    Results  We isolated mutant alleles with loss-of-function mutations in  RACK1B  and  RACK1C , and examined the impact of these mutations on plant development. We found that unlike in  RACK1A , loss-of-function mutations in  RACK1B  or  RACK1C  do not confer apparent defects in plant development, including rosette leaf production and root development. Analyses of  rack1a ,  rack1b  and  rack1c  double and triple mutants, however, revealed that  rack1b  and  rack1c  can enhance the rack1a mutant's developmental defects, and an extreme developmental defect and lethality were observed in rack1a rack1b rack1c triple mutant. Complementation studies indicated that RACK1B and RACK1C are in principle functionally equivalent to  RACK1A . Gene expression studies indicated that three  RACK1  genes display similar expression patterns but are expressed at different levels. Further,  RACK1  genes positively regulate each other's expression.    Conclusion  These results suggested that  RACK1  genes are critical regulators of plant development and that  RACK1  genes function in an unequally redundant manner. Both the difference in  RACK1  gene expression level and the cross-regulation are likely the molecular determinants of their unequal genetic redundancy.

RACK1 genes regulate plant development with unequal genetic redundancy in Arabidopsis