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The leafy head characteristic is a special phenotype of Chinese cabbage resulting from artificial selection during domestication and breeding.  BREVIS RADIX  ( BRX ) has been suggested to control root elongation, shoot growth, and tiller angle in Arabidopsis and rice. In  Brassica rapa , three  BrBRX  homoeologs have been identified, but only  BrBRX.1  and  BrBRX.2  were found to be under selection in leaf-heading accessions, indicating their functional diversification in leafy head formation. Here, we show that these three  BrBRX  genes belong to a plant-specific  BRX  gene family but that they have significantly diverged from other  BRX-like  members on the basis of different phylogenetic classifications, motif compositions and expression patterns. Moreover, although the expression of these three  BrBRX  genes differed, compared with  BrBRX.3 ,  BrBRX.1,  and  BrBRX.2  displayed similar expression patterns. Arabidopsis mutant complementation studies showed that only  BrBRX.1  could rescue the  brx  root phenotype, whereas  BrBRX.2  and  BrBRX.3  could not. However, overexpression of each of the three  BrBRX  genes in Arabidopsis resulted in similar pleiotropic leaf phenotypes, including epinastic leaf morphology, with an increase in leaf number and leaf petiole length and a reduction in leaf angle. These leaf traits are associated with leafy head formation. Further testing of a SNP (T/C) in  BrBRX.2  confirmed that this allele in the heading accessions was strongly associated with the leaf-heading trait of  B. rapa . Our results revealed that all three  BrBRX  genes may be involved in the leaf-heading trait, but they may have functionally diverged on the basis of their differential expression.

Divergence of three BRX homoeologs in Brassica rapa and its effect on leaf morphology