Artemisia annua is a medicinal plant rich in terpenes and flavonoids with useful biological activities such as antioxidant, anticancer, and antimalarial activities. The transcriptional regulation of flavonoid biosynthesis in A. annua has not been well-studied. In this study, we identified a YABBY family transcription factor, AaYABBY5, as a positive regulator of anthocyanin and total flavonoid contents in A. annua. AaYABBY5 was selected based on its similar expression pattern to the phenylalanine ammonia lyase (PAL), chalcone synthase (CHS), chalcone isomerase (CHI), and flavonol synthase (FLS) genes. A transient dual-luciferase assay in Nicotiana bethamiana with the AaYABBY5 effector showed a significant increase in the activity of the downstream LUC gene, with reporters AaPAL, AaCHS, AaCHI, and AaUFGT. The yeast one-hybrid system further confirmed the direct activation of these promoters by AaYABBY5. Gene expression analysis of stably transformed AaYABBY5 overexpression, AaYABBY5 antisense, and control plants revealed a significant increase in the expression of AaPAL, AaCHS, AaCHI, AaFLS, AaFSII, AaLDOX, and AaUFGT in AaYABBY5 overexpression plants. Moreover, their total flavonoid content and anthocyanin content were also found to increase. AaYABBY5 antisense plants showed a significant decrease in the expression of flavonoid biosynthetic genes, as well as a decrease in anthocyanin and total flavonoid contents. In addition, phenotypic analysis revealed deep purple-pigmented stems, an increase in the leaf lamina size, and higher trichome densities in AaYABBY5 overexpression plants. Together, these data proved that AaYABBY5 is a positive regulator of flavonoid biosynthesis in A. annua. Our study provides candidate transcription factors for the improvement of flavonoid concentrations in A. annua and can be further extended to elucidate its mechanism of regulating trichome development.

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The beneficial endophytic microorganisms have received significant attention in agriculture because of their exceptional capabilities to facilitate functions like nutrient enrichment, water status, and stress tolerance (biotic and abiotic). This review signifies the molecular mechanisms to better understand the Piriformospora indica-mediated plants improvement or protection for sustainable agriculture. P. indica, an endophytic fungus, belonging to the order Sebacinales (Basidiomycota), is versatile in building mutualistic associations with a variety of plants including pteridophytes, bryophytes, gymnosperms, and angiosperms. P. indica has enormous potential to manipulate the hormonal pathway such as the production of indole-3-acetic acid which in turn increases root proliferation and subsequently improves plant nutrient acquisition. P. indica also enhances components of the antioxidant system and expression of stress-related genes which induce plant stress tolerance under adverse environmental conditions. P. indica has tremendous potential for crop improvement because of its multi-dimensional functions such as plant growth promotion, immunomodulatory effect, biofertilizer, obviates biotic (pathogens) and abiotic (metal toxicity, water stress, soil structure, salt, and pH) stresses, phytoremediator, and bio-herbicide. Considering the above points, herein, we reviewed the physiological and molecular mechanisms underlying P. indica-mediated plants improvement or protection under diverse agricultural environment. The first part of the review focuses on the symbiotic association of P. indica with special reference to biotic and abiotic stress tolerance and host plant root colonization mechanisms, respectively. Emphasis is given to the expression level of essential genes involved in the processes that induce changes at the cellular level. The last half emphasizes critical aspects related to the seed germination, plant yield, and nutrients acquisition.

Aim of the current study was to investigate the effect of exogenously inoculated root endophytic fungus, Piriformospora indica , on molecular, biochemical, morphological and physiological parameters of Artemisia annua L. treated with different concentrations (0, 50, 100 and 150 μmol/L) of arsenic (As) stress. As was significantly accumulated in the roots than shoots of P . indica -inoculated plants. As accumulation and immobilization in the roots is directly associated with the successful fungal colonization that restricts most of As as compared to the aerial parts. A total of 4.1, 11.2 and 25.6 mg/kg dry weight of As was accumulated in the roots of inoculated plants supplemented with 50, 100 and 150 μmol/L of As, respectively as shown by atomic absorption spectroscopy. P . indica showed significant tolerance in vitro to As toxicity even at high concentration. Furthermore, flavonoids, artemisinin and overall biomass were significantly increased in inoculated-stressed plants. Superoxide dismutase and peroxidase activities were increased 1.6 and 1.2 fold, respectively under 150 μmol/L stress in P . indica -colonized plants. Similar trend was followed by ascorbate peroxidase, catalase and glutathione reductase. Like that, phenolic acid and phenolic compounds showed a significant increase in colonized plants as compared to their respective control/un-colonize stressed plants. The real-time PCR revealed that transcriptional levels of artemisinin biosynthesis genes, isoprenoids, terpenes, flavonoids biosynthetic pathway genes and signal molecules were prominently enhanced in inoculated stressed plants than un-inoculated stressed plants.

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