Shoot formation is accompanied by active cell proliferation and expansion, requiring that metabolic state adapts to developmental control. Despite the importance of such metabolic reprogramming, it remains unclear how development and metabolism are integrated. Here, we show that disruption of ANGUSTIFOLIA3 orthologs (PpAN3s) compromises gametophore shoot formation in the moss Physcomitrium patens due to defective cell proliferation and expansion. Trans-omics analysis reveals that the downstream activity of PpAN3 is linked to arginine metabolism. Elevating arginine level by chemical treatment leads to stunted gametophores and causes Ppan3 mutant-like transcriptional changes in the wild-type plant. Furthermore, ectopic expression of AtAN3 from Arabidopsis thaliana ameliorates the defective arginine metabolism and promotes gametophore formation in Ppan3 mutants. Together, these findings indicate that arginine metabolism is a key pathway associated with gametophore formation and provide evolutionary insights into the establishment of the shoot system in land plants through the integration of developmental and metabolic processes.

芽的形成伴随着活跃的细胞增殖和扩张，需要新陈代谢状态适应发育控制。尽管这种代谢重编程很重要，但仍不清楚如何将发育与代谢整合在一起。在这里，我们显示破坏ANGUSTIFOLIA3直系同源物（PpAN3 s）会损害苔藓Physcomitrium patens的配子体芽形成，这是由于细胞增殖和扩增缺陷所致。转基因组学分析表明，PpAN3的下游活性与精氨酸代谢有关。通过化学处理提高精氨酸水平会导致发育迟缓，并导致Ppan3野生型植物中的突变体样转录变化。此外，来自拟南芥的AtAN3的异位表达改善了精氨酸的新陈代谢，并促进了Ppan3突变体中配子体的形成。总之，这些发现表明精氨酸代谢是与配子体形成相关的关键途径，并通过发育和代谢过程的整合为陆地植物芽系统的建立提供了进化见识。