Flavonoids are an important class of secondary metabolites widely found in plants, contributing to plant growth and development and having prominent applications in food and medicine. The biosynthesis of flavonoids has long been the focus of intense research in plant biology. Flavonoids are derived from the phenylpropanoid metabolic pathway, and have a basic structure that comprises a C15 benzene ring structure of C6-C3-C6. Over recent decades, a considerable number of studies have been directed at elucidating the mechanisms involved in flavonoid biosynthesis in plants. In this review, we systematically summarize the flavonoid biosynthetic pathway. We further assemble an exhaustive map of flavonoid biosynthesis in plants comprising eight branches (stilbene, aurone, flavone, isoflavone, flavonol, phlobaphene, proanthocyanidin, and anthocyanin biosynthesis) and four important intermediate metabolites (chalcone, flavanone, dihydroflavonol, and leucoanthocyanidin). This review affords a comprehensive overview of the current knowledge regarding flavonoid biosynthesis, and provides the theoretical basis for further elucidating the pathways involved in the biosynthesis of flavonoids, which will aid in better understanding their functions and potential uses.

The accumulation of various pigments leads to the formation of different flower colors in plants. However, the regulation mechanism of yellow flower formation and flower color differences between Camellia nitidssima C.W.Chi (CN) and its hybrids C. ‘Zhenghuangqi’ (ZHQ), C. ‘Huangxuanlv’ (HXL), and C. ‘Xinshiji’ (XSJ), remains largely unknown. Here, we showed that the content of two flavonols, quercetin-7-O-glucoside (Qu7G) and quercetin-3-O-glucoside (Qu3G), was positively correlated with the yellow degree of petals in CN and its three hybrids. Additionally, we performed a comparative transcriptomic analysis of petals of the four yellow camellia plants, which revealed 322 common upregulated and 866 common downregulated DEGs (differentially expressed genes) in the CN vs. ZHQ, CN vs. HXL, and CN vs. XSJ comparison groups. Their regulatory pathway analysis showed that flavonol biosynthesis genes (FLSs and GTs) and transcriptional regulatory genes MYBs were all expressed higher in CN than its three hybrids, which corresponded to differences in the flavonol content among the four yellow camellias. Further, two ethylene synthesis genes (ACSs, ACO) and three ethylene signaling genes (EIN2s, EIN3, ERFs) were all upregulated in the yellow petals of CN. In conclusion, the expression of flavonol-related genes and flavonols (Qu7G and Qu3G) accumulation could play a key role in the formation of yellow flowers in camellia, and the ethylene pathway might be involved in the regulation of yellow flower formation of camellias. This work describes the possible regulatory pathway of yellow camellia, thereby laying a foundation for future verification of genes linked to flower coloring and the breeding of yellow camellia.

Camellia nitidissima is a woody plant with high ornamental value, and its golden-yellow flowers are rich in a variety of bioactive substances, especially flavonoids, that are beneficial to human health. Chalcone isomerases (CHIs) are key enzymes in the flavonoid biosynthesis pathway; however, there is a scarcity of information regarding the CHI family genes of C. nitidissima. In this study, seven CHI genes of C. nitidissima were identified and divided into three subfamilies by phylogenetic analysis. The results of multiple sequence alignment revealed that, unlike CnCHI1/5/6/7, CnCHI2/3/4 are bona fide CHIs that contain all the active site and critical catalytic residues. Analysis of the expression patterns of CnCHIs and the total flavonoid content of the flowers at different developmental stages revealed that CnCHI4 might play an essential role in the flavonoid biosynthesis pathway of C. nitidissima. CnCHI4 overexpression significantly increased flavonoid production in Nicotiana tabacum and C. nitidissima. The results of the dual-luciferase reporter assay and yeast one-hybrid system revealed that CnMYB7 was the key transcription factor that governed the transcription of CnCHI4. The study provides a comprehensive understanding of the CHI family genes of C. nitidissima and performed a preliminary analysis of their functions and regulatory mechanisms.