Physiological and Metabolomics Analyses Reveal the Roles of Fulvic Acid in Enhancing the Production of Astaxanthin and Lipids in Haematococcus pluvialis under Abiotic Stress Conditions

In this study, it was found that fulvic acid (FA) enhanced the contents of astaxanthin and lipids in Haematococcus pluvialis under high light and nitrogen starvation conditions by 2- and 1.2-fold, respectively. Meanwhile, the carbohydrate and chlorophyll contents were decreased by FA induction, whereas the levels of reactive oxygen species (ROS) and glutathione (GSH) as well as the expression of astaxanthin and lipid biosynthetic genes were increased. To further explore the interrelation between FA and the biosynthesis of astaxanthin and lipids, a metabolomics analysis of H. pluvialis by combined FA and abiotic stress exposure was conducted by using LC-MS/MS. The contents of some cytoprotective metabolites and signal molecules, including n-maltose, succinate, malic acid, melatonin (MT), and some amino acids, were increased under FA induction and abiotic stress conditions. These metabolites are intermediates in the TCA cycle and Calvin cycle, providing more precursors for the synthesis of astaxanthin and lipids. Moreover, the signal molecules might contribute to enhancing the abiotic stress tolerance. This study provided new insights into the regulatory mechanism of FA on astaxanthin and lipid accumulation in H. pluvialis.