“…The leaves have an endodermis, a sclerenchyma layer, and a cuticle as apoplastic barriers (Ribeiro, Santos & Moraes, 2007;De los Ángeles Lagoria et al, 2018;Mahley et al, 2018;Palacios-Rios et al, 2019). Moreover, the morphologies of sun and shade plants are obviously different (Boardman, 1977;Nasrulhaq-Boyce & Mohamed, 1987;Givnish, 1988;Santos-Silva, Mastroberti & De Araujo Mariath, 2011), the leaves of shade plants are large, with large cells, few mesophyll tissues, and decreased stomatal and vein densities (Ribeiro, Santos & Moraes, 2007;Neira et al, 2017;Dematteis et al, 2019;Shah et al, 2019;Baer, Wheeler & Pittermann, 2020). In ferns, gymnosperms, and angiosperms, the endodermis or circumendodermal band and the exodermis act as barriers, which restrict water-solute exchange, reduce oxygen loss after submersion, and terrestrial environments (Ogura, 1972;Lersten, 1997;Enstone, Peterson & Ma, 2003;Hernandez-Hernandez et al, 2012;Geldner, 2013;Xiang et al, 2019;Yang et al, 2011Yang et al, , 2014Yang et al, , 2019aYang et al, , 2019bYang et al, , 2019cYang et al, , 2020Zhang et al, 2017;Zhang, Yang & Seago, 2018).…”