Exogenous Substances Used to Relieve Plants from Drought Stress and Their Associated Underlying Mechanisms

Drought stress (DS) is one of the abiotic stresses that plants encounter commonly in nature, which affects their life, reduces agricultural output, and prevents crops from growing in certain areas. To enhance plant tolerance against DS, abundant exogenous substances (ESs) have been attempted and proven to be effective in helping plants relieve DS. Understanding the effect of each ES on alleviation of plant DS and mechanisms involved in the DS relieving process has become a research focus and hotspot that has drawn much attention in the field of botany, agronomy, and ecology. With an extensive and comprehensive review and summary of hundred publications, this paper groups various ESs based on their individual effects on alleviating plant/crop DS with details of the underlying mechanisms involved in the DS-relieving process of: (1) synthesizing more osmotic adjustment substances; (2) improving antioxidant pathways; (3) promoting photosynthesis; (4) improving plant nutritional status; and (5) regulating phytohormones. Moreover, a detailed discussion and perspective are given in terms of how to meet the challenges imposed by erratic and severe droughts in the agrosystem through using promising and effective ESs in the right way and at the right time.