The AcBBX4/6–AcHY5 Transcription Complex Mediates Flavonoid Biosynthesis during Pineapple Fruit Ripening

led to reduced flavonoid accumulation, whereas overexpression enhanced it. Coordinated manipulation of all three factors produced the most pronounced phenotypic effects on flavonoid content. Collectively, our results define the AcBBXs-AcHY5 complex as a core transcriptional module governing flavonoid biosynthesis during pineapple fruit ripening.