Table 3.
Transcription factors involved in artemisinin biosynthesis pathway.
| Name | Function | Expression Mode | Fold Change of Artemisinin Content | Reference |
|---|---|---|---|---|
| WRKY TF family | ||||
| AaWRKY1 | AaWRKY1 activates the expression of ADS, CYP71AV1, DBR2 genes. | Overexpression | 1.30–2.00 | [68] |
| AaWRKY4 | AaWRKY4 prompts the expression of ADS, CYP71AV1, DBR2, ALDH1 genes. | Overexpression | 1.35–1.50 | [69] |
| AaWRKY9 | AaWRKY9 positively regulates the expression of AaDBR2 and AaGSW1 genes. | Overexpression | 1.60–2.20 | [70] |
| Down-regulation | 0.55–0.65 | |||
| AaWRKY17 | AaWRKY17 activates the transcription of ADS, PR5 and NHL10 genes and enhances the resistance of A. annua to the pathogen P. syringae. | Overexpression | 1.50–1.87 | [71] |
| Down-regulation | 0.79–0.85 | |||
| AaGSW1 | AaGSW1 is a GST-specific TF, and positively regulates the expression of CYP71AV1 and AaORA genes. | Overexpression | 1.55–2.00 | [72] |
| AaGSW2 | As a GST-specific TF, AaGSW2 positively regulates the initiation of GST. | Overexpression | 2.00 | [74] |
| Down-regulation | 0.54 | |||
| bHLH TF family | ||||
| AabHLH1 | AabHLH1 activates transcription of ADS and CYP71AV1 genes and responds to JA induction. | Overexpression | 1.30 | [77] |
| Down-regulation | 0.60 | |||
| AaMYC2 | AaMYC2 prompts the expression of CYP71AV1 and DBR2 genes and responses to JA induction. | Overexpression | 1.24 | [78] |
| Down-regulation | 0.55 | |||
| AabHLH112 | AabHLH112 promotes artemisinin biosynthesis by activating TF AaERF1. | Overexpression | 1.70 | [79] |
| AabHLH2 | AabHLH2 and AabHLH3 suppress the expression of ADS and CYP71AV1 genes by antagonizing AaMYC2, and negatively regulate artemisinin biosynthesis. | Overexpression | 0.34–0.73 | [81] |
| Down-regulation | 1.42–1.87 | |||
| AabHLH3 | Overexpression | 0.39–0.79 | ||
| Down-regulation | 1.35–1.60 | |||
| AP2/ERF TF family | ||||
| AaERF1 | AaERF1 and AaERF2 activate the transcription of ADS and CYP71AV1 genes and respond to JA signaling. | Overexpression | 1.19–1.67 | [80] |
| AaERF2 | Overexpression | 1.24–1.51 | ||
| AaORA | AaORA is a trichome-specific TF, which promotes the expression of ADS, CYP71AV1, DBR2 and AaERF1 genes, and improves the resistance of A. annua to the pathogen B. cinerea. | Overexpression | 1.40–1.53 | [82] |
| Down-regulation | 0.64–0.52 | |||
| AaTAR1 | AaTAR1 activates the transcription of ADS and CYP71AV1 genes and regulates trichome development. | Overexpression | 1.22–1.38 | [83] |
| Down-regulation | 0.36–0.61 | |||
| bZIP TF family | ||||
| AabZIP1 | AabZIP1 activates the transcription of ADS and CYP71AV1 genes through ABA signaling, indirectly promotes the expression of AaDBR2 and AaALDH1 genes through AaMYC2 and improves the drought resistance of A. annua. | Overexpression | 1.40–1.60 | [84,85] |
| AabZIP9 | AabZIP9 activates the expression of ADS gene and positively regulates artemisinin biosynthesis. | Overexpression | 1.23–1.67 | [86] |
| AaHY5 | AaHY5 positively regulates light-induced artemisinin biosynthesis through interacting with TF AaGSW1. | Overexpression | 2.00 | [87] |
| Down-regulation | 0.50 | |||
| AaTGA6 | AaTGA6 takes part in artemisinin biosynthesis regulated by SA signaling and promotes artemisinin biosynthesis by activating the TF AaERF1. | Overexpression | 1.90–2.20 | [88] |
| Down-regulation | 0.40–0.80 | |||
| MYB TF family | ||||
| AaMYB1 | AaMYB1 participates in the biosynthesis of artemisinin and GA and regulates the trichome development. | Overexpression | 2.00 | [89] |
| AaMIXTA1 | AaMIXTA1 is a GST-specific TF, which regulates GST formation and cuticle biosynthesis. | Overexpression | 2.00 | [90] |
| Down-regulation | 0.75 | |||
| AaTAR2 | AaTAR2 positively regulates GST development, as well as artemisinin and flavonoid biosynthesis. | Overexpression | ~1.38–1.54 1 | [91] |
| Down-regulation | ~0.57–0.84 1 | |||
| AaMYB16 (AaMIXTA-like 2) | AaMYB5 and AaMYB16 show the antagonism and regulate the development of GST by competitively binding to the AaHD1 promoter to form AaHD1-AaMYB5 or AaHD1-AaMYB16 complexes. | Overexpression | 1.34–1.56 | [92] |
| Down-regulation | 0.73–0.81 | |||
| AaMYB5 | Overexpression | 0.73–0.85 | ||
| Down-regulation | 1.45–1.84 | |||
| AaMYB17 | AaMYB17 is a GST-specific TF and positively regulates artemisinin biosynthesis. | Overexpression | 1.88 | [93] |
| Down-regulation | 0.75 | |||
| AaMYB15 | As a GST-specific TF, AaMYB15 negatively regulates the biosynthesis of artemisinin by inhibiting the expression of AaORA gene. | Overexpression | ~0.60–0.74 1 | [94] |
| Down-regulation | ~1.52 1 | |||
| AaTLR1 | AaTLR1 and AaTLR2 interact with AaWOX1 to form the AaTLR1-AaWOX1-AaTLR2 complex to negatively regulate GST formation and artemisinin biosynthesis. | Overexpression | 0.51–0.88 | [95] |
| Down-regulation | 1.32–1.84 | |||
| AaTLR2 | Overexpression | 0.57–0.81 | ||
| O t her TF family | ||||
| AaZFP1 | As a C2H2-type TF, AsZFP1 activates the transcription of AaIPPI1 gene and prompts artemisinin biosynthesis. | Overexpression | 1.60 | [96] |
| AaSPL9 | AaSPL9 positively regulates glandular trichomes initiation and artemisinin biosynthesis by activating TF AaHD1. | Overexpression | 1.33–1.60 | [97] |
| AaSEP4 | AaSEP4, as a MADS TF, promotes artemisinin biosynthesis by directly activating TF AsGSW1. | Overexpression | 1.19–1.72 | [98] |
1 The change fold was calculated according to the data in the references.