Plant responses to heterogeneous salinity: growth of the halophyte Atriplex nummularia is determined by the root-weighted mean salinity of the root zone

Soil salinity is generally spatially heterogeneous, but our understanding of halophyte physiology under such conditions is limited. The growth and physiology of the dicotyledonous halophyte Atriplex nummularia was evaluated in split-root experiments to test whether growth is determined by: (i) the lowest; (ii) the highest; or (iii) the mean salinity of the root zone. In two experiments, plants were grown with uniform salinities or horizontally heterogeneous salinities (10450mM NaCl in the low-salt side and 670mM in the high-salt side, or 10mM NaCl in the low-salt side and 5001500mM in the high-salt side). The combined data showed that growth and gas exchange parameters responded most closely to the root-weighted mean salinity rather than to the lowest, mean, or highest salinity in the root zone. In contrast, midday shoot water potentials were determined by the lowest salinity in the root zone, consistent with most water being taken from the least negative water potential source. With uniform salinity, maximum shoot growth was at 120230mM NaCl; similar to 90% of maximum growth occurred at 10mM and 450mM NaCl. Exposure of part of the roots to 1500mM NaCl resulted in an enhanced (+40%) root growth on the low-salt side, which lowered root-weighted mean salinity and enabled the maintenance of shoot growth. Atriplex nummularia grew even with extreme salinity in part of the roots, as long as the root-weighted mean salinity of the root zone was within the 10450mM range.