Winter leaf reddening in 'evergreen' species

Leaf reddening during autumn in senescing, deciduous tree species has received widespread attention from the public and in the scientific literature, whereas leaf reddening in evergreen species during winter remains largely ignored. Winter reddening can be observed in evergreen herbs, shrubs, vines and trees in Mediterranean, temperate, alpine, and arctic regions, and can persist for several months before dissipating with springtime warming. Yet, little is known about the functional significance of this colour change, or why it occurs in some species but not others. Here, the biochemistry, physiology and ecology associated with winter leaf reddening are reviewed, with special focus on its possible adaptive function. Photoprotection is currently the favoured hypothesis for winter reddening, but alternative explanations have scarcely been explored. Intraspecific reddening generally increases with sunlight incidence, and may also accompany photosynthetic inferiority in photosynthetically 'weak' (e.g. low-nitrogen) individuals. Red leaves tend to show symptoms of shade acclimation relative to green, consistent with a photoprotective function. However, winter-red and winter-green species often cohabitate the same high-light environments, and exhibit similar photosynthetic capacities. The factors dictating interspecific winter leaf colouration therefore remain unclear. Additional outstanding questions and future directions are also highlighted, and possible alternative functions of winter reddening discussed.