Mechanisms of photoprotection in overwintering evergreen conifers: Sustained quenching of chlorophyll fluorescence

Evergreen conifers growing in high-latitude regions must endure prolonged winters that are characterized by sub-zero temperatures combined with light, conditions that can cause significant photooxidative stress. Understanding overwintering mechanisms is crucial for addressing winter adversity in temperate forest ecosystems and enhancing the ability of conifers to adapt to climate change. This review synthesizes the current understanding of the photoprotective mechanisms that conifers employ to mitigate photooxidative stress, particularly non-photochemical "sustained quenching", the mechanism of which is hypothesized to be a recombination or deformation of the original mechanism employed by conifers in response to short-term low temperature and intense light stress in the past. Based on this hypothesis, scattered studies in this field are assembled and integrated into a complete mechanism of sustained quenching embedded in the adaptation process of plant physiology. It also reveals which parts of the whole system have been verified in conifers and which have only been verified in non-conifers, and proposes specific directions for future research. The functional implications of studies of non-coniferous plant species for the study of coniferous trees are also considered, as a wide range of plant responses lead to sustained quenching, even among different conifer species. In addition, the review highlights the challenges of measuring sustained quenching and discusses the application of ultrafast-time-resolved fluorescence and decay-associated spectra for the elucidation of photosynthetic principles.