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Transcriptional adapter ADA2 regulates yield and end‐use quality through liquid–liquid phase separation in wheat

Xiaobang Zhang1, Ruijie Zhang1, Yuhan Zhang1, Tao Yang1, Longfei Wu1, Yuan Cao2,3, Jun Xiao2,3,4, Jigang Li5, Xin Gao6, Xinyou Cao6, Jie Liu1, Mingming Xin1, Yuan Zong1, Zhongfu Ni1, Qixin Sun1 and Yingyin Yao1*   

  1. 1. State Key Laboratory of High‐Efficiency Production of Wheat–Maize Double Cropping, Frontiers Science Center for Molecular Design Breeding (MOE), China Agricultural University, Beijing 100193, China
    2. Laboratory of Advanced Breeding Technologies, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China
    3. University of Chinese Academy of Sciences, Beijing 100049, China
    4. CAS‐JIC Centre of Excellence for Plant and Microbial Science (CEPAMS), Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China
    5. State Key Laboratory of Plant Environmental Resilience, College of Biological Sciences, Frontiers Science Center for Molecular Design Breeding (MOE), China Agricultural University, Beijing 100193, China
    6. National Engineering Research Center of Wheat and Maize, State Key Laboratory of Wheat Improvement, Key Laboratory of Wheat Biology and Genetic Improvement in North Yellow & Huai River Valley, Shandong Provincial Technology Innovation Center for Wheat, Crop Research Institute, Shandong Academy of Agricultural Sciences, Jinan 250100, China
    *Correspondence: Yingyin Yao (yingyin@cau.edu.cn)
  • Received:2025-11-19 Accepted:2026-02-26 Online:2026-03-16

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Abstract: Wheat (Triticum aestivum L.) yield and quality are critically influenced by starch and seed storage protein (SSP) content. In this study, we demonstrate that the transcriptional adapter ADA2 physically interacts with histone acetyltransferase GENERAL CONTROL NONDEREPRESSIBLE 5 (GCN5) and regulates starch synthesis and SSP content in wheat grains. In ada2 mutants, reduced H3K9ac enrichment was observed in the promoter regions of key starch synthesis genes and the SSP regulator NAC019 during endosperm development. This reduction leads to lower expression levels, resulting in decreased starch content, smaller grains, reduced yield, and poor gluten quality. We also found that ADA2 contains an intrinsically disordered region 2 (IDR2) that undergoes liquid–liquid phase separation (LLPS) and forms nuclear condensates. In vitro assays, the LLPS of ADA2 is affected by GCN5 through direct interaction with IDR2, and the ratio of their concentrations determines the behavior of phase‐separated condensates and HAT activity. High levels of GCN5 can dissolve ADA2 aggregates, while excessive ADA2 recruits and sequesters GCN5 into co‐aggregated droplets, with lower HAT activity. This potential dynamic regulatory mechanism may facilitate the efficient promotion of transcription by the ADA2–GCN5 complex in wheat. Finally, we identified an elite haplotype of ADA2‐B Hap2, which is significantly associated with grain size and weight, highlighting its potential as a candidate gene for genetic improvement of wheat yield.

Key words: ADA2, GCN5, LLPS, quality, seed storage protein, starch, wheat (Triticum aestivum L.), yield

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