Physiological and morphological adaptations of the fruit tree Ziziphus rotundifolia in response to progressive drought stress

The physiological basis of drought resistance in Ziziphus rotundifolia Lamk., which is an important, multipurpose fruit tree of the northwest Indian and zone, was investigated in a greenhouse experiment. Three irrigation regimes were imposed over a 34-day period: an irrigation treatment, a gradual drought stress treatment (50% of water supplied in the irrigation treatment) and a rapid drought stress treatment (no irrigation). Changes in gas exchange, water relations, carbon isotope composition and solute concentrations of leaves, stems and roots were determined. The differential rate of stress development in the two drought treatments did not result in markedly different physiological responses, but merely affected the time at which they were expressed. The initial response to decreasing soil water content was reduced stomatal conductance, effectively maintaining predawn leaf water potential (Psi,(leaf)), controlling water loss and increasing intrinsic water-use efficiency, while optimizing carbon gain during drought. Carbon isotope composition (delta C-13) of leaf tissue sap provided a more sensitive indicator of changes in short-term water-use efficiency than delta C-13 of bulk leaf tissue. As drought developed, osmotic potential at full turgor decreased and total solute concentrations increased in leaves, indicating osmotic adjustment. Decreases in leaf starch concentrations and concomitant increases in hexose sugars and sucrose suggested a shift in carbon partitioning in favor of soluble carbohydrates. In severely drought-stressed leaves, high leaf nitrate reductase activities were paralleled by increases in proline concentration, suggesting an osmoprotective role for proline. As water deficit increased, carbon was remobilized from leaves and preferentially redistributed to stems and roots, and leaves were shed, resulting in reduced whole-plant transpiration and enforced dormancy. Thus, Z. rotundifolia showed a range of responses to different drought intensities indicating a high degree of plasticity in response to water deficits.