We review observational, experimental, and model results on how plants respond to extreme climatic conditions induced by changing climatic variability. Distinguishing between impacts of changing mean climatic conditions and changing climatic variability on terrestrial ecosystems is generally underrated in current studies. The goals of our review are thus (1) to identify plant processes that are vulnerable to changes in the variability of climatic variables rather than to changes in their mean, and (2) to depict/evaluate available study designs to quantify responses of plants to changing climatic variability. We find that phenology is largely affected by changing mean climate but also that impacts of climatic variability are much less studied, although potentially damaging. We note that plant water relations seem to be very vulnerable to extremes driven by changes in temperature and precipitation and that heat-waves and flooding have stronger impacts on physiological processes than changing mean climate. Moreover, interacting phenological and physiological processes are likely to further complicate plant responses to changing climatic variability. Phenological and physiological processes and their interactions culminate in even more sophisticated responses to changing mean climate and climatic variability at the species and community level. Generally, observational studies are well suited to study plant responses to changing mean climate, but less suitable to gain a mechanistic understanding of plant responses to climatic variability. Experiments seem best suited to simulate extreme events. In models, temporal resolution and model structure are crucial to capture plant responses to changing climatic variability. We highlight that a combination of experimental, observational, and/or modeling studies have the potential to overcome important caveats of the respective individual approaches.

Provenance-specific growth responses to experimentally applied drought and air warming were studied in saplings of three European oak species: Quercus robur, Quercus petraea and Quercus pubescens. Four provenances of each species were grown in large open-top chambers and subjected to four climates: control, periodic drought, air warming or their combination in 3 subsequent years. Overall growth responses were found among species and provenances, with drought reducing shoot height growth and stem diameter growth and air warming stimulating shoot height growth but reducing stem diameter growth and root length growth. Differential growth responses in shoots, stems and roots resulted in altered allometric growth relations. Root length growth to shoot height growth increased in response to drought but decreased in response to air warming. Stem diameter growth to shoot height growth decreased in response to air warming. The growth responses in shoots and stems were highly variable among provenances indicating provenance-specific sensitivity to drought and air warming, but this response variability did not reflect local adaptation to climate conditions of provenance origin. Shoot height growth was found to be more sensitive to drought in provenances from northern latitudes than in provenances from southern latitudes, suggesting that genetic factors related to the postglacial immigration history of European oaks might have interfered with selective pressure at provenance origins.

Global climate change is expected to increase annual temperatures and decrease summer precipitation in Central Europe. Little is known of how forests respond to the interaction of these climate factors and if their responses depend on soil conditions. In a 3-year lysimeter experiment, we investigated the growth response of young mixed oak stands, on either acidic or calcareous soil, to soil water regime, air-warming and drought treatments corresponding to an intermediate climate change scenario. The air-warming and drought treatments were applied separately as well as in combination. The air-warming treatment had no effect on soil water availability, evapotranspiration or stand biomass. Decreased evapotranspiration from the drought-exposed stands led to significantly higher air and soil temperatures, which were attributed to impaired transpirational cooling. Water limitation significantly reduced the stand foliage, shoot and root biomass as droughts were severe, as shown in low leaf water potentials. Additional air warming did not enhance the drought effects on evapotranspiration and biomass, although more negative leaf water potentials were observed. After re-watering, evapotranspiration increased within a few days to pre-drought levels. Stands not subjected to the drought treatment produced significantly less biomass on the calcareous soil than on the acidic soil, probably due to P or Mn limitation. There was no difference in biomass and water regime between the two soils under drought conditions, indicating that nutrient availability was governed by water availability under these conditions. The results demonstrate that young oak stands can cope with severe drought and therefore can be considered for future forestry.

The quantities and spatial distribution of nutrients in savanna ecosystems are affected by many factors, of which fire, herbivory and symbiotic N 2fixation are particularly important. We measured soil nitrogen (N) pools and the relative abundance of N and phosphorus (P) in herbaceous vegetation in five vegetation types in a humid savanna in Tanzania. We also performed a factorial fertilization experiment to investigate which nutrients most limit herbaceous production. N pools in the top 10 cm of soil were low at sites where fires were frequent, and higher in areas with woody legume encroachment, or high herbivore excretion. Biomass production was co-limited by N and P at sites that were frequently burnt or heavily grazed by native herbivores. In contrast, aboveground production was limited by N in areas receiving large amounts of excreta from livestock. N 2 -fixation by woody legumes did not lead to P-limitation, but did increase the availability of N relative to P. We conclude that the effects of fire, herbivory and N 2fixation upon soil N pools and N:P-stoichiometry in savanna ecosystems are, to a large extent, predictable.Electronic supplementary material: The online version of this article (

Oaks are commonly considered as drought- and heat-tolerant trees that might benefit from a warmer and drier climate. Their tolerance to drought has been frequently studied in the past, whereas studies dealing with elevated temperature or its combination with drought are very limited in number. In this study we investigated seasonal photosynthetic patterns in three European oak species (Quercus robur, Q. petraea, Q. pubescens) exposed in lysimeter-based open-top chambers (OTC) to elevated daytime temperature, drought and their combination. Stomatal and non-stomatal traits of photosynthesis were followed over an entire growing season and related to changes in daytime temperature, soil moisture and pre-dawn leaf water potential (Ψ(PD) ). Elevated daytime temperature enhanced net photosynthesis (P(N) ) in a season-dependent manner, with higher mid-summer rates than in controls exposed to ambient temperature. Drought imposed in early and mid-summer reduced the soil moisture content and caused a gradual decline in Ψ(PD) , stomatal conductance (g(S) ) and P(N) . Drought effects on Ψ(PD) and P(N) were exacerbated when drought was combined with elevated daytime temperature. In general, P(N) tended to be more affected by low soil moisture content or low Ψ(PD) in Q. robur than in Q. petraea and Q. pubescens. Non-stomatal limitations may have contributed to the drought-induced decline of P(N) in Q. robur, as indicated by a down-regulation of PSII photochemistry (F(V) /F(M) ) and decreased chlorophyll content. Taken together, our findings show that European oaks may benefit from elevated temperature, but detrimental effects can be expected when elevated temperature occurs simultaneously with drought.

Tree water relations and their dependence on microclimate and soil moisture were studied over several months in young oaks (Quercus robur L.) subjected in large lysimeter-based open top chambers to environments with a controlled soil water supply. Automated single point dendrometers and the recently developed leaf patch clamp pressure (LPCP) probe were used for monitoring water-related stem radius variations (ΔW) and turgor-dependent leaf patch pressures (Pp). Both parameters showed distinct diurnal patterns with sharp negative and positive peaking of ΔW and Pp, respectively, after solar noon and recovery to initial levels in the evening. During the day, varying solar radiation was responsible for short time fluctuations of Pp in the range of minutes to hours reflecting feedback regulation of leaf turgor by sunlight driven stomatal movements. At longer timescales, i.e. days to months, atmospheric vapour pressure deficit (VPD) and soil water content (SWC) were the main determinants of ΔW and Pp. Daily minimum and maximum values of ΔW and Pp decreased and increased, respectively, with increasing VPD or decreasing SWC and recovery of ΔW and Pp in the evening was impeded by low SWC. In well-watered oaks, daily positive peaking of Pp preceded daily negative peaking of ΔW; these time lags gradually increased with increasing soil drought, suggesting hydraulic uncoupling of stem and leaves.

Climate change is expected to increase temperature and decrease summer precipitation in Central Europe. Little is known about how warming and drought will affect phenological patterns of oaks, which are considered to possess excellent adaptability to these climatic changes. Here, we investigated bud burst and intra-annual shoot growth of Quercus robur, Q. petraea and Q. pubescens grown on two different forest soils and exposed to air warming and drought. Phenological development was assessed over the course of three growing seasons. Warming advanced bud burst by 1–3 days °C−1 and led to an earlier start of intra-annual shoot growth. Despite this phenological shift, total time span of annual growth and shoot biomass were not affected. Drought changed the frequency and intensity of intra-annual shoot growth and advanced bud burst in the subsequent spring of a severe summer drought by 1–2 days. After re-wetting, shoot growth recovered within a few days, demonstrating the superior drought tolerance of this tree genus. Our findings show that phenological patterns of oaks are modified by warming and drought but also suggest that ontogenetic factors and/or limitations of water and nutrients counteract warming effects on the biomass and the entire span of annual shoot growth.

Background and aims Oaks are considered to be drought-and thermo-tolerant tree species. Nevertheless, species and provenances may differ in their ecological requirements. We hypothesised that (i) provenances from xeric sites are better adapted to drought than those from more humid sites, (ii) oaks direct root growth towards resource-rich layers, and (iii) air-warming promotes root growth. Methods To test different provenances of Quercus robur, Q. petraea and Q. pubescens, we conducted a model ecosystem experiment with young trees, grown on acidic and calcareous soil, subjected to drought, air warming, the combination of warming and drought, and a control. Results The results were only in partial agreement with the first hypothesis. As expected the provenances originating from drier sites produced more biomass than those from more humid sites under drought conditions. Surprisingly, however, they reacted more sensitive to water limiting conditions, as they produced also substantially more biomass under well-watered conditions. The drought treatment reduced root mass substantially in the upper soil. In agreement with the second hypothesis this led to a shift in the centre of root mass to lower depth, where water was still more available than closer to the soil surface. In contrast to the third hypothesis, the air-warming treatment, which was very mild however compared to climate change scenarios, had no significant effects on root growth. Conclusions Given that the provenances from drier sites showed more biomass loss at water limiting conditions than provenances from more humid sites, it remains questionable whether provenances from drier sites are better suited for a future climate. Keywords Climate change. Fine root. Root mass distribution. Root:shoot ratio. Quercus robur. Quercus petraea. Quercus pubescens