Drought is one of the main abiotic stresses that cause crop yield loss. Improving crop yield under drought stress is a major goal of crop breeding, as it is critical to food security. The mechanism of plant drought resistance has been well studied, and diverse drought resistance genes have been identified in recent years, but transferring this knowledge from the laboratory to field production remains a significant challenge. Recently, some new strategies have become research frontiers owing to their advantages of low cost, convenience, strong field operability, and/or environmental friendliness. Exogenous plant growth regulator (PGR) treatment and microbe-based plant biotechnology have been used to effectively improve crop drought tolerance and preserve yield under drought stress. However, our understanding of the mechanisms by which PGRs regulate plant drought resistance and of plant-microbiome interactions under drought is still incomplete. In this review, we summarize these two strategies reported in recent studies, focusing on the mechanisms by which these exogenous treatments regulate crop drought resistance. Finally, future challenges and directions in crop drought resistance breeding are discussed.

Background: Drought threatens the food supply of the world population. Dissecting the dynamic responses of plants to drought will be beneficial for breeding drought-tolerant crops, as the genetic controls of these responses remain largely unknown. Results: Here we develop a high-throughput multiple optical phenotyping system to noninvasively phenotype 368 maize genotypes with or without drought stress over a course of 98 days, and collected multiple optical images, including color camera scanning, hyperspectral imaging, and X-ray computed tomography images. We develop high-throughput analysis pipelines to extract image-based traits (i-traits). Of these i-traits, 10,080 were effective and heritable indicators of maize external and internal drought responses. An i-trait-based genome-wide association study reveals 4322 significant locus-trait associations, representing 1529 quantitative trait loci (QTLs) and 2318 candidate genes, many that co-localize with previously reported maize drought responsive QTLs. Expression QTL (eQTL) analysis uncovers many local and distant regulatory variants that control the expression of the candidate genes. We use genetic mutation analysis to validate two new genes, ZmcPGM2 and ZmFAB1A, which regulate i-traits and drought tolerance. Moreover, the value of the candidate genes as drought-tolerant genetic markers is revealed by genome selection analysis, and 15 i-traits are identified as potential markers for maize drought tolerance breeding. Conclusion: Our study demonstrates that combining high-throughput multiple optical phenotyping and GWAS is a novel and effective approach to dissect the genetic architecture of complex traits and clone drought-tolerance associated genes.