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Background  HMA4 transporters are involved in the transport and binding of divalent heavy metals (Cd, Zn, Pb [lead] and Co [cobalt]). In general, as efflux pumps, HMA4 transporters can increase the heavy metal tolerance of yeast and  Escherichia coli . Additional research has shown that the C-terminus of HMA4 contains a heavy metal-binding domain and that heterologous expression of a portion of peptides from this C-terminal domain in yeast provides a high level of Cd tolerance and Cd hyperaccumulation.    Results  We cloned  BjHMA4  from  Brassica juncea , and quantitative real-time PCR analysis revealed that  BjHMA4  was upregulated by Zn and Cd in the roots, stems and leaves. Overexpression of  BjHMA4  dramatically affects Zn/Cd distribution in rice and wheat seedlings. Interestingly, BjHMA4 contains a repeat region named BjHMA4R within the C-terminal region; this repeat region is not far from the last transmembrane domain. We further characterized the detailed function of BjHMA4R via yeast and  E. coli  experiments. Notably, BjHMA4R greatly and specifically improved Cd tolerance, and BjHMA4R transformants both grew on solid media that contained 500 μM CdCl 2  and presented improved Cd accumulation (approximately twice that of wild-type [WT] strains). Additionally, visualization via fluorescence microscopy indicated that BjHMA4R clearly localizes in the cytosol of yeast. Overall, these findings suggest that BjHMA4R specifically improves Cd tolerance and Cd accumulation in yeast by specifically binding Cd 2+  in the cytosol under low heavy metal concentrations. Moreover, similar results in  E. coli  experiments corroborate this postulation.    Conclusion  BjHMA4R can specifically bind Cd 2+  in the cytosol, thereby substantially and specifically improving Cd tolerance and accumulation under low heavy metal concentrations.    Electronic supplementary material  The online version of this article (10.1186/s12870-019-1674-5) contains supplementary material, which is available to authorized users.

bmc plant biology

A repeat region from the Brassica juncea HMA4 gene BjHMA4R is specifically involved in Cd2+ binding in the cytosol under low heavy metal concentrations