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In contrast, drought decreased growth, photosynthetic pigments, protein and Rubisco. MS was enhanced by PAs applied as a seed primer or foliar spray, as shown by clear reductions in EL %, malondialdehyde (MDA) content and the Na+/K+ ratio as well as reduced ROS markers and elevated CAT (but not GPX) activity. Further study showed that the Mix solution of PAs, applied either during seed priming or as a foliar spray, improved chloroplast ultra-structure, suggesting that improvements in Rubisco and photosynthetic pigments were involved in P