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AtKEAs, homologs of bacterial KefB/KefC, are predicted to encode K + /H +  antiporters in Arabidopsis. The  AtKEA  family contains six genes forming two subgroups in the cladogram:  AtKEA1-3  and  AtKEA4-6 . AtKEA1 and AtKEA2 have a long N-terminal domain; the full-length AtKEA1 was inactive in yeast. The transport activity was analyzed by expressing the  AtKEA  genes in yeast mutants lacking multiple ion carriers. AtKEAs conferred resistance to high K +  and hygromycin B but not to salt and Li +  stress.  AtKEAs  expressed in both the shoot and root of Arabidopsis. The expression of  AtKEA1 ,  -3  and  -4  was enhanced under low K +  stress, whereas  AtKEA2  and  AtKEA5  were induced by sorbitol and ABA treatments. However, osmotic induction of  AtKEA2  and  AtKEA5  was not observed in  aba2-3  mutants, suggesting an ABA regulated mechanism for their osmotic response.  AtKEAs ’ expression may not be regulated by the SOS pathway since their expression was not affected in  sos  mutants. The GFP tagging analysis showed that AtKEAs distributed diversely in yeast. The Golgi localization of AtKEA3 was demonstrated by both the stably transformed seedlings and the transient expression in protoplasts. Overall, AtKEAs expressed and localized diversely, and may play roles in K +  homeostasis and osmotic adjustment in Arabidopsis.

A Novel AtKEA Gene Family, Homolog of Bacterial K+/H+ Antiporters, Plays Potential Roles in K+ Homeostasis and Osmotic Adjustment in Arabidopsis