); [EMAIL_ADDRESS] (D.C.);  
Yingdong Fan  
1 College of Horticulture and Landscape Architecture, Northeast Agricultural University, Harbin 150030, China; [EMAIL_ADDRESS] (W.Z. ); [EMAIL_ADDRESS] (I. ); [EMAIL_ADDRESS] (Z.H. ); [EMAIL_ADDRESS] (Y.F. ); [EMAIL_ADDRESS] (J.D. ); [EMAIL_ADDRESS] (C.L. ); [EMAIL_ADDRESS] (T.Y. ); [EMAIL_ADDRESS] (D.C.);  
Yingdong Fan  
1 College of Horticulture and Landscape Architecture, Northeast Agricultural University, Harbin 150030, China; [EMAIL_ADDRESS] (W.Z. ); [EMAIL_ADDRESS] (I. ); [EMAIL_ADDRESS] (Z.H. ); [EMAIL_ADDRESS] (Y.F. ); [EMAIL_ADDRESS] (J.D. ); [EMAIL_ADDRESS] (C.L. ); [EMAIL_ADDRESS] (T.Y. ); [EMAIL_ADDRESS] (D.C.);  
[PERSON]  
1 College of Horticulture and Landscape Architecture, Northeast Agricultural University, Harbin 150030, China; [EMAIL_ADDRESS] (W.Z. ); [EMAIL_ADDRESS] (I. ); [EMAIL_ADDRESS] (Z.H. ); [EMAIL_ADDRESS] (Z.H. ); [EMAIL_ADDRESS] (Y.F. ); [EMAIL_ADDRESS] (J.D. ); [EMAIL_ADDRESS] (C.L. ); [EMAIL_ADDRESS] (T.Y. ); [EMAIL_ADDRESS] (D.C.);  
[PERSON]  
1 College of Horticulture and Landscape Architecture, Northeast Agricultural University, Harbin 150030, China; [EMAIL_ADDRESS] (W.Z. ); [EMAIL_ADDRESS] (I. ); [EMAIL_ADDRESS] (Z.H. ); [EMAIL_ADDRESS] (Z.H. ); [EMAIL_ADDRESS] (Y.F. ); [EMAIL_ADDRESS] (J.D. ); [EMAIL_ADDRESS] (C.L. ); [EMAIL_ADDRESS] (T.Y. ); [EMAIL_ADDRESS] (D.C.);  
[PERSON] and [PERSON]  
1 College of Horticulture and Landscape Architecture, Northeast Agricultural University, Harbin 150030, China; [EMAIL_ADDRESS] (W.Z. ); [EMAIL_ADDRESS] (I. ); [EMAIL_ADDRESS] (Z.H. ); [EMAIL_ADDRESS] (Y.F. ); [EMAIL_ADDRESS] (J.D. ); [EMAIL_ADDRESS] (C.L. ); [EMAIL_ADDRESS] (T.Y. ); daidiche@neanu.edu.cnT triterpenenoid saponins are glycosides consisting of sapogenins combined with sugars. Most triterpenoids discovered so far are tetracyclic and pentacyclic triterpenoids, with the most common structure being the dammarane-type and oleanane-type [7]. Tetracyclic triterpenes are widely distributed in nature; ginensosides and gibberellin are tetracyclic triterpenoids [8]. Pentacyclic saponins are more commonly observed in Chinese herbs such as _Glycyrrhiza uralensis_, _Bupleurum chinense_, and _Polygala tenuifolia_[9]. The saponins are mainly oleanane-type pentacyclic triterpenoids in _P. grandiflora_, of which the main sapogenins is oleanolic acid. Plants mainly derive triterpenoids from the cytosolic mevalonic acid pathway (MVA), with a smaller contribution from the plastidial methylerythirtl phosphate pathway (MEP) [10]. Farnesyl pyrophosphate (FPP) is derived from the precursor isopentenyl diphosphate (IPP) through a series of enzymatic reactions. With the assistance of key enzymes such as squalene epoxidase (SE) and farnesyl-diphosphate synthase (FPPS), Farnesyl-PP is transformed into 2,3-oxidosqualene. Subsequently, 2,3-oxidosqualene is cyclized by 2,3-oxidosqualene cyclase (OSC) to yield \(\beta\)-amyrin [11; 12; 13]. Further, \(\beta\)-amyrin is modified by cytochrome P450 to form oleanolic acid [14]. Cytochrome P450 is a class of heme-based B-type cytochrome superfamily protein that extensively participates in the biosynthesis of specialized metabolites (e.g., terpenes, plant hormones, and alkaloids) in plants [15]. The main enzymes involved in catalyzing the synthesis of oleanolic acid from \(\beta\)-amyrin are from the CY7P716 family of the cytochrome P450 superfamily [16]. McCYP716A12 is the first identified enzyme involved in the synthesis of pentacyclic triterpene saponins, which catalyzes the C-28 oxidation of \(\beta\)-amyrin into oleanolic acid [17]. With the development of sequencing technology, numerous CYP716 enzymes involved in catalyzing oleanolic acid formation have been identified and validated [18; 19; 20]. In _Kalopanax septemblous_ and _Clenopodium quinota_, KosCYP16A94, CqCYP16A78, and CqCYP716A79 catalyze the C-28 oxidation of \(\beta\)-amyrin into oleanolic acid [21; 22]. Meanwhile, five CYP716 enzymes (including PgCYP716A140, PgCYP716A141, PgCYP716D58, PgCYP716S7, and PgCYP716S6) were identified and validated in _P.