Downregulation of PSI regulates photosynthesis in early successional tree species. Evidence from a field survey across European forests

Photosynthesis rates and electron transport are key physiological traits that distinguish the successional status of plants (early successional – ES, and late successional – LS) in a community. ES plants can respond quickly to sudden changes in sunlight radiation exposure and display a greater photosynthetic plasticity, with respect to the LS ones, thanks to their ability to regulate the photosynthesis. Photosynthesis regulation is connected to the efficiency of the two photosystems (PSI and PSII). PSI plays an important role in the photosynthetic machinery functioning to respond against sudden changes of metabolism in response to heterogeneous environmental conditions. It can be downregulated under high pressure of reductants, to maintain the balance between the reductant pressure and the metabolic demand. The efficiency of PSII and of PSI can be estimated, respectively, by the maximum quantum yield of photosystem II (FV/FM) and the contribution of the thermal phase in the chlorophyll fluorescence emission (expressed by the parameter ΔVIP, that quantify the I-P phase in the fluorescence emission transient). Both parameters are evaluated in dark-adapted samples and can be measured by prompt fluorescence technique. Here, we analyzed a large dataset from a European-wide survey on mature forests, considering four representative forest tree species growing at six sampling sites distributed along a latitudinal gradient. Two early successional (Pinus sylvestris and Betula pendula) and two late successional (Picea abies and Fagus sylvatica) species were considered. The correlations between FV/FM and I-P phase were mostly positive for different plant species at each sampling site for FV/FM values < 0.82 (assumed as optimal threshold for this parameter), but not for FV/FM values > 0.82. In this last case, the correlations were mostly negative for early successional species, and not significant for late successional species. Foliar nitrogen concentration plays a relevant role for the PSI regulation in early successional species: trees with high nitrogen content have higher I-P phase values and negative correlations between FV/FM and I-P phase for FV/FM > 0.82. An opposite pattern was found in tree species with low foliar nitrogen level. We conclude that the late successional tree species show a substantial regulation and stability of the photosynthetic machinery and photosystem stoichiometry, whereas early successional species have more dynamic behavior of PSI. Early successional species can modulate the photosynthetic efficiency and are able both to up-regulate and down-regulate the PSI concentration and activity in relation to quantum yield capacity and leaf nitrogen content.