International Journal of Horticulture, 2025, Vol.15, No.5, 234-241 http://hortherbpublisher.com/index.php/ijh 238 texture and sweetness of the fruit at the same time. Antioxidant metabolism genes like ascorbic acid biosynthesis and circulating genes help in fruit ripening to achieve this dynamic balance (Hua et al., 2018). 5.4 Integrated regulatory network of softening, sugar accumulation, and antioxidant metabolism Yellow-skinned Pitaya fruit ripening is a coordinated regulatory network that coordinates fruit softening, sugar buildup, and antioxidant metabolism. The network is regulated by complicated interplays of transcription factors, phytohormones and metabolic routes. For example, HpWRKY3 and Dof transcription factors regulate sugar metabolism genes, while ethylene and ABA signaling routes are central to the overall maturation process (Xu et al., 2021). Furthermore, antioxidant metabolic processes also help control oxidative stress for stability of fruit quality during fruit ripening. Reveal of such a system-thinking regulatory mechanism could enable finding of the fruit ripening's genetic and biochemical mechanism and providing future strategies to improving fruit quality and extending shelf life (Durán-Soria et al., 2020; Li et al., 2023) (Figure 2). Figure 2 General patterns of sugar accumulation conduction and its crosstalk with hormones (Adopted from Durán-Soria et al., 2020) 6 Research Progress in Pitaya Ripening Regulation 6.1 Functional analysis and validation of ripening-related genes Over the past few years, some of the key transcription factors participating in the regulation of bird's nest fructose accumulation and thereby fruit ripening have been identified by scientists. For example, the WRKY transcription factor HpWRKY3 induces the expression of sucrose metabolism-related genes HpINV2 and HpSuSy1, which results in fruit sugar accumulation. Similarly, Dof transcription factors HpDof1.7 and HpDof5.4 were discovered to induce the expression of sugar metabolism genes, leading to the accumulation of glucose and fructose (Vicente et al., 2007). These studies all show that transcription factors play a very significant role in the genetic control of Pitaya fruit ripening. 6.2 Application of omics technologies in fruit ripening studies Omics technology, especially transcriptomics, is especially significant in demonstrating genes involved in fructose metabolism in bird's nest. For example, in transcriptome analysis, a number of sugar and organic acid metabolism genes were identified through analysis, resulting in new findings regarding fruit development processes (Xu et al., 2024). In addition, omics technology also offers the potential to genome-wide screen transcription factors such as R2R3-MYB family proteins regulating fruit ripening through mechanisms such as regulating beet elutin biosynthesis. Coupling with multiomics approaches can further be capable of disclosing the complex network of Pitaya ripening control (Adaskaveg and Blanco‐Ulate, 2023).
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