AaSPATULA synergistically coordinates glandular trichome initiation and vegetative growth to maximize whole-plant artemisinin yield in Artemisia annua

Artemisinin, the frontline antimalarial sesquiterpene lactone derived from Artemisia annua, is exclusively biosynthesized and stored in glandular secretory trichomes (GSTs). Despite significant progress in elucidating its biosynthetic pathway and key regulatory transcription factors (TFs), critical gaps remain in understanding how hormone signaling integrates GST development with artemisinin biosynthesis and resolves the trade-off between trichome density, metabolic output, and vegetative growth for whole-plant yield optimization. We discuss implications for metabolic engineering and propose future directions, including multi-omics-guided multi-target editing of bHLH hubs, to achieve sustainable high-yield A. annua varieties.