“…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 ; Zhang & Wu, 2013 ; 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, 2011 , 2014 , 2019a , 2019b , 2019c , 2020 ; Zhang et al, 2017 ; Zhang, Yang & Seago, 2018 ).…”