“…Given the evergreen nature of their needles, conifers must employ mechanisms to protect their photosynthetic apparatus during severe winters and recover their capacity for photosynthesis during the subsequent spring (Yamazaki et al, 2003;Zarter et al, 2006aZarter et al, , 2006b). Several photoprotective mechanisms have been invoked as preventing photodamage of the photosynthetic apparatus in overwintering evergreens, including (i) enhanced cyclic electron transport around photosystem I (Huner et al, 1988;Ivanov et al, 2001;Ö quist and Huner, 2003), (ii) degradation of key photosystem II proteins (D1 and the oxygen evolving complex) to inhibit superoxide formation (Ottander et al, 1995;Adams et al, , 2006Zarter et al, 2006aZarter et al, , 2006bZarter et al, , 2006c, and (iii) sustained engagement of the carotenoids zeaxanthin and antheraxanthin in continuously high levels of photoprotective energy dissipation, resulting in sustained low PSII efficiency, during the winter Demmig-Adams, 1994, 1995;Adams et al, , 2002Adams et al, , 2006Verhoeven et al, 1996Verhoeven et al, , 1998Adams and Barker, 1998). It has also been reported that, in zeaxanthin-free isolated major light-harvesting complexes (LHCII), neoxanthin Ilioaia et al, 2011;Zubik et al, 2011) and lutein (Ilioaia et al, 2011;Jahns and Holzwarth, 2012;Wahadoszamen et al, 2012) are able to quench fluorescence and thus facilitate 238 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH ᭧ 2013 Regents of the University of Colorado 1523-0430/6 $7.00 thermal energy dissipation (Ilioaia et al, 2011;Ruban et al, 2012).…”