“…Consequently, a precise and 4 integrative landscape of seed development is necessary and useful for both fundamental and 4applied studies on the mechanisms underpinning crop yield and quality. Previously, from 4 viewpoint of crop physiology, we proposed a practical staging system with three phases: 4 embryo morphogenesis, endosperm filling, and seed maturation(An et al, 2020). Here, we 4 update this staging system by integrating of molecular, physiological, and anatomical 4 evidence in the current study as well as from the literature.…”