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Plants cultivated on the Qinghai-Tibetan Plateau grow in an extremely cold environment and thus are exposed to cold stress. To assess the metabolic processes during cold exposure of Tibetan hulless barley (Hordeum distichon L.), metabolic analyses were conducted on one tolerant (XiLa) and one sensitive (ZangQing) cultivar exposed to six temperatures (24 °C, 12 °C, 5 °C, 0 °C, -5 °C, -8 °C) for 24 h. In total, 770 metabolites were identified, including amino acids and derivatives, carbohydrates, flavonoids, lipids, nucleotides and derivatives, and phenolamides. In principal component analysis, the samples were clearly grouped according to the cultivar, suggesting that the two cultivars have differential responses to cold stress. In cold-grown plants, eight metabolites, including monoacylglycerol (MAG, 18:2), MAG (18:3), deoxyadenosine, 6-methylmercaptopurine, and coniferin, were significantly altered in XiLa, but not in ZangQing when compared to the levels in control plants, and thus, these compounds can be considered as potential biomarkers of exposure to cold stress in hulless barley. Furthermore, differentially altered metabolites between seedlings exposed to -8 °C and those maintained at 24 °C were significantly enriched in glutathione metabolism. The findings of this study will be useful for the development of cultivars with cold stress tolerance., Competing Interests: Declaration of competing interest The authors declare that there is no conflict of interests., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Tibetan hulless barley is widely grown in the extreme environmental conditions of the Qinghai-Tibet Plateau which is characterized by cold, high salinity, and drought. Osmotic stress always occurs simultaneously with drought and its tolerance is a vital part of drought tolerance. The diversity of metabolites leading to osmotic stress tolerance was characterized using widely-targeted metabolomics in tolerant (XL) and sensitive (D) accessions submitted to polyethylene glycol. XL regulated a more diverse set of metabolites than D, which may promote the establishment of a robust system to cope with the stress in XL. Compounds belonging to the group of flavonoids, amino acids, and glycerophospholipids constitute the core metabolome responsive to the stress, despite the tolerance levels. Moreover, 8 h appeared to be a critical time point for stress endurance involving a high accumulation of key metabolites from the class of nucleotide and its derivative which provide the ultimate energy source for the synthesis of functional carbohydrates, lipids, peptides, and secondary metabolites in XL. This intrinsic metabolic adjustment helped XL to efficiently alleviate the stress at the later stages. A total of 22 diverse compounds were constantly and exclusively regulated in XL, representing novel stress tolerance biomarkers which may help improving stress tolerance, especially drought, in hulless barley.

Powdery mildew is a fungal disease that represents a ubiquitous threat to crop plants. Transcriptomic and metabolomic analyses were used to identify molecular and physiological changes in Tibetan hulless barley in response to powdery mildew. There were 3418 genes and 405 metabolites differentially expressed between the complete resistance cultivar G7 and the sensitive cultivar Z13. Weighted gene coexpression network analysis was carried out, and the differentially expressed genes were enriched in five and four major network modules in G7 and Z13, respectively. Further analyses showed that phytohormones, photosynthesis, phenylpropanoid biosynthesis, and flavonoid biosynthesis pathways were altered during Qingke-Blumeria graminis (DC.) f.sp. hordei (Bgh) interaction. Comparative analyses showed a correspondence between gene expression and metabolite profiles, and the activated defenses resulted in changes of metabolites involved in plant defense response, such as phytohormones, lipids, flavone and flavonoids, phenolamides, and phenylpropanoids. This study enabled the identification of Bgh responsive genes and provided new insights into the dynamic physiological changes that occur in Qingke during response to powdery mildew. These findings greatly improve our understanding of the mechanisms of induced defense response in Qingke and will provide new clues for the development of resistant Tibetan hulless barley varieties.