Morphological and physiological responses of 14 macadamia rootstocks to drought stress and a comprehensive evaluation of drought resistance

Planting macadamia in karst rocky desertification areas has significant ecological and economic benefits. However, the production of macadamia in karst area are facing serious drought challenges due to the poor water holding capacity of soil, and less and uneven precipitation resulting with the rising of temperature in the world. Selecting and breeding drought-resistant rootstocks is one of the main measures to solve this problem in macadamia nut production in karst area. In this study, the morphological and physiological performance of half-sib family offspring of 14 macadamia cultivars were investigated under severe drought conditions and their drought resistance was evaluated comprehensively in order to select the most suitable macadamia rootstocks. Results revealed that the leaves of all rootstocks were damaged after drought stress. Among the tested rootstocks, the leaves of QA5 were damaged the most severely, and the leaves of the whole plants were chlorotic, followed by C11, A4, and NY2, while the symptoms in leaves of O.C and H2 was less obvious after drought stress for 30 d. At the same time, the O.C and H2 had the lowest drought injury index (DII), which was 0.40, while the QA5 had the highest DII value (0.82). The physiological indices, including malonaldehyde (MDA), superoxide dismutase (SOD), catalase (CAT), peroxidase (POD), ascorbate peroxidase (APX), soluble protein (SP), soluble sugar (SS), proline (Pro) and relative electrolyte leakage (REL) increased (Drought tolerance coefficient, DTC > 1) after drought stress, while relative water content of leaves (L-RWC), chlorophyll (Chl), relative water content of roots (R-RWC) and root activity (RA) decreased (DTC < 1) in all cultivars. Except SP, SS and Pro, there were significant differences in other indexes among different rootstocks. The 13 physiological indices were transformed into seven independent, comprehensive indices through principal component analysis (PCA), whose cumulative contribution ratio reached 82.905%. The 14 rootstocks were divided into 3 categories based on the comprehensive evaluation value (D). The O.C and H2 were clustered into the drought-tolerant group, whose D values were 0.743 and 0.669, respectively. The NY3, 333, 788, NY116, NY1, NY2 were belonged to the intermediate drought tolerance group, the D values were 0.530, 0.508, 0.497, 0.465, 0.457 and 0.441 respectively. Lastly, A16, A4, C11. GR1, 695 and QA5 were clustered into low drought tolerance group with the lowest D values of 0.396, 0.391, 0.360, 0.360, 0.355, 0.324, respectively. The result of stepwise regression analysis showed that MDA, POD, APX, SS, REL, R-RWC and RA were the most significant physiological and biochemical indices that played a significant role in drought tolerance, and could be used as the primary evaluation and identification indicators of drought tolerance of macadamia. These results indicated that the drought-resistant rootstocks of macadamia could improve their drought resistance by stimulating the cluster root development, increasing antioxidant enzyme activity and raising the content of osmotic regulatory substances under drought stress. Our study not only provided alternative rootstocks for macadamia drought-resistant cultivation, but also provided the theoretical basis for the drought-resistant rootstock breeding of macadamia by expounding the physiological mechanism of drought-resistance.