L-Ascorbic acid metabolism and regulation in fruit crops

Abstract

L-Ascorbic acid (AsA) is more commonly known as vitamin C and is an indispensable compound for human health. As a major antioxidant, AsA not only maintains redox balance and resists biological and abiotic stress but also regulates plant growth, induces flowering, and delays senescence through complex signal transduction networks. However, AsA content varies greatly in horticultural crops, especially in fruit crops. The AsA content of the highest species is approximately 1,800 times higher than that of the lowest species. There have been significant advancements in the understanding of AsA accumulation in the past 20 years. The most noteworthy accomplishment was the identification of the critical rate-limiting genes for the 2 major AsA synthesis pathways (L-galactose pathway and D-galacturonic acid pathway) in fruit crops. The rate-limiting genes of the former are GMP, GME, GGP, and GPP, and the rate-limiting gene of the latter is GalUR. Moreover, APX, MDHAR, and DHAR are also regarded as key genes in degradation and regeneration pathways. Interestingly, some of these key genes are sensitive to environmental factors, such as GGP being induced by light. The efficiency of enhancing AsA content is high by editing upstream open reading frames (uORF) of the key genes and constructing multi-gene expression vectors. In summary, the AsA metabolism has been well understood in fruit crops, but the transport mechanism of AsA and the synergistic improvement of AsA and other traits is less known, which will be the focus of AsA research in fruit crops.

Figure 1.

AsA content in various horticultural crops (A) and accumulation patterns (B–D). All the data and figures were collected from related studies (Ye 2011; Huang 2013) and reports as well as the Web of Science, China National Knowledge Infrastructure, and public websites. Panels B–D represent different AsA accumulation patterns. The data used to draw schematic diagram were obtained from kiwifruit (A. eriantha), Chinese jujube (cv “Mazao”), and chestnut rose (cv “Guinong 5”) (Huang 2013; Lu et al. 2022; Liu et al. 2022b).

Figure 2.

Long-distance transport and metabolism of AsA in plant organelles. AsA was accumulated into phloem and transported to root tips, shoots, and floral organs, but generally not to mature leaves. At the cytological level, AsA is synthesized in mitochondria and then enters the cytoplasm for transport to various organelles. In addition, AsA and DHA can also be transported outside the cell membrane by simple diffusion or transport proteins, such as the Cytb. After AsA functions in the chloroplast, DHA is produced and transported into the cytoplasm. Elements were modified from FigDraw (https://www.figdraw.com/static/index.html).

Figure 3.

Proposed pathways for AsA metabolism. L-galactose pathway (I), D-galacturonic acid pathway (II), inositol pathway (III), L-gulose pathway (IV), degradation and cyclic regeneration pathway (V) of AsA metabolism, common in the synthetic pathway (VI). PME: Methylesterase; PMI: Mannose-6-phosphate isomerase; PMM: Phosphomannose mutase; GulDH: L-gulose-1,4-lactyl dehydrogenase; GR: Glutathione reductase.

Figure 4.

Reported TFs and environmental regulatory networks for AsA accumulation. Green arrows indicate promotion of expression or AsA accumulation, red arrows indicate inhibition of expression or AsA accumulation. A) D-fructose-6P, (B) D-mannose-6P, (C) D-mannose-1P, (D) GDP-D-mannose, (E) GDP-L-galactose, (F) L-galactose1-P, (G) L-galactose, (H) L-galactono-1,4-lactone, (I) monodehydroascorbate, (J) dehydroascorbate. AMR: AsA mannose pathway regulator.