Root development under metal stress in Arabidopsis thaliana requires the H+/cation antiporter CAX4.

* The Arabidopsis vacuolar CAtion eXchangers (CAXs) play a key role in mediating cation influx into the vacuole. In Arabidopsis, there are six CAX genes. However, some members are yet to be characterized fully. * In this study, we show that CAX4 is expressed in the root apex and lateral root primordia, and that expression is increased when Ni(2+) or Mn(2+) levels are elevated or Ca(2+) is depleted. * Transgenic plants expressing increased levels of CAX4 display symptoms consistent with increased sequestration of Ca(2+) and Cd(2+) into the vacuole. When CAX4 is highly expressed in an Arabidopsis cax1 mutant line with weak vacuolar Ca(2+)/H(+) antiport activity, a 29% increase in Ca(2+)/H(+) antiport is measured. A cax4 loss-of-function mutant and CAX4 RNA interference lines display altered root growth in response to Cd(2+), Mn(2+) and auxin. The DR5::GUS auxin reporter detected reduces auxin responses in the cax4 lines. * These results indicate that CAX4 is a cation/H(+) antiporter that plays an important function in root growth under heavy metal stress conditions.