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Climate change as projected for Central Europe will lead to prolonged periods of summer drought and enhanced air temperature. Thus, forest management practices are required to take into account how species performance is adapted to cope with these climate changes. Oak trees may play a major role in future forests because of their relative drought-tolerance compared with other species like beech. Therefore, this study investigated the stress responses (i.e., anti-oxidants, free amino acids) in the leaves of three widely distributed oak species in Central Europe (i.e., Quercus robur L., Q. petraea [Matt.] Libel., Q. pubescens Willd.) to drought, air warming and the combination of drought plus air warming under controlled conditions after periods of spring drought, a short rewetting and summer drought. We quantified foliar levels of thiols, ascorbate, and free amino compounds in Q robur, Q. petraea and Q. pubescens. Our study showed that oak saplings had increased levels of γ-glutamylcysteine and total glutathione and proline with drought and air warming. Foliar ascorbate, glutathione disulfide and dehydroascorbic acid levels were not affected. The comparison of stress responses to drought and/or air warming between the three species showed higher foliar thiol levels in Q. robur and Q. pubescens compared with Q. petraea. For total and reduced ascorbic acid and γ-aminobutyric acid, the highest levels were found in Q. robur. In conclusion, our study showed that foliar anti-oxidant and free amino acid levels were significantly affected by drought plus air warming; however, this effect was species-dependent with the drought-tolerant species of Q. pubescens having the highest reactive oxygen species scavenging capacity among three tested oak species. Furthermore, stress responses as shown by increased levels of foliar anti-oxidants and free amino acids differ between calcareous and acidic soil indicating that the capacities of anti-oxidative defense and osmotic stress adjustment developed better on calcareous compared with acidic soil; however, this effect was metabolite- as well as species-specific.

The paradigm that permafrost-affected soils show restricted mineral nitrogen (N) cycling in favor of organic N compounds is based on the observation that net N mineralization rates in these cold climates are negligible. However, we find here that this perception is wrong. By synthesizing published data on N cycling in the plant-soil-microbe system of permafrost ecosystems we show that gross ammonification and nitrification rates in active layers were of similar magnitude and showed a similar dependence on soil organic carbon (SOC) and total nitrogen (TN) concentrations as observed in temperate and tropical systems. Moreover, high protein depolymerization rates and only marginal effects of C:N stoichiometry on gross N turnover provided little evidence for N limitation. Instead, the rather short period when soils are not frozen is the single main factor limiting N turnover. High gross rates of mineral N cycling are thus facilitated by released protection of organic matter in active layers with nitrification gaining particular importance in N-rich soils, such as organic soils without vegetation. Our finding that permafrost-affected soils show vigorous N cycling activity is confirmed by the rich functional microbial community which can be found both in active and permafrost layers. The high rates of N cycling and soil N availability are supported by biological N fixation, while atmospheric N deposition in the Arctic still is marginal except for fire-affected areas. In line with high soil mineral N production, recent plant physiological research indicates a higher importance of mineral plant N nutrition than previously thought. Our synthesis shows that mineral N production and turnover rates in active layers of permafrost-affected soils do not generally differ from those observed in temperate or tropical soils. We therefore suggest to adjust the permafrost N cycle paradigm, assigning a generally important role to mineral N cycling. This new paradigm suggests larger permafrost N climate feedbacks than assumed previously.

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