Beauty is more than epidermis deep: How cell division and expansion sculpt the leaf spongy mesophyll

As the main location of photosynthesis, leaf mesophyll cells are one of the most abundant and essential cell types on earth. Forming the bulk of the internal tissues of the leaf, their size, shape, and patterns of interconnectivity define the internal structure and surface area of the leaf, which in turn determines the efficiency of light capture and carbon fixation. Understanding how these cellular traits are controlled and translated into tissue- and organ-scale traits, and how they influence photosynthetic performance will be key to our ability to improve crop plants in the face of a changing climate. In contrast to the extensive literature on the anatomical and physiological aspects of mesophyll function, our understanding of the cell-level morphogenetic processes underpinning mesophyll cell growth and differentiation is scant. In this review, we focus on how cell division, expansion, and separation are coordinated to create the intricate architecture of the spongy mesophyll.