“…Leaves located in upper and outer parts of a tree crown have larger photosynthetic capacity and higher amounts of nitrogen (N) stored in photosynthetic proteins, such as Rubisco and chlorophyll-binding polypeptides, compared with leaves in the lower part or interior of the crown ( Leuning et al , 1991 ; Camm, 1993 ; Ackerly, 1999 ; Millard and Grelet, 2010 ; Bresinsky et al , 2013 ). The foliage density required to maximize C uptake is commonly far exceeded in tree crowns, suggesting different roles for lower-crown foliage other than contributing to the net gain in C assimilation ( Thomas and Sadras, 2001 ; Hirose, 2005 ; Körner, 2018 ). Shaded leaves have been recognized to serve as nutrient and C stores for later resorption by younger leaves, and as a buffer in the case of foliage loss, especially in evergreen conifers ( Nambiar and Fife, 1987 ; Thomas and Sadras, 2001 ; Millard and Grelet, 2010 ; Hirose, 2012 ).…”