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Forests store the largest terrestrial pools of carbon (C), helping to stabilize the global climate system, yet are threatened by climate change (CC) and associated air pollution (AP, highlighting ozone (O3) and nitrogen oxides (NOx)). We adopt the perspective that CC-AP drivers and physiological impacts are universal, resulting in consistent stress responses of forest ecosystems across zonobiomes. Evidence supporting this viewpoint is presented from the literature on ecosystem gross/net primary productivity and water cycling. Responses to CC-AP are compared across evergreen/deciduous foliage types, discussing implications of nutrition and resource turnover at tree and ecosystem scales. The availability of data is extremely uneven across zonobiomes, yet unifying patterns of ecosystem response are discernable. Ecosystem warming results in trade-offs between respiration and biomass production, affecting high elevation forests more than in the lowland tropics and low-elevation temperate zone. Resilience to drought is modulated by tree size and species richness. Elevated O3 tends to counteract stimulation by elevated carbon dioxide (CO2). Biotic stress and genomic structure ultimately determine ecosystem responsiveness. Aggrading early- rather than mature late-successional communities respond to CO2 enhancement, whereas O3 affects North American and Eurasian tree species consistently under free-air fumigation. Insect herbivory is exacerbated by CC-AP in biome-specific ways. Rhizosphere responses reflect similar stand-level nutritional dynamics across zonobiomes, but are modulated by differences in tree-soil nutrient cycling between deciduous and evergreen systems, and natural versus anthropogenic nitrogen (N) oversupply. The hypothesis of consistency of forest responses to interacting CC-AP is supported by currently available data, establishing the precedent for a global network of long-term coordinated research sites across zonobiomes to simultaneously advance both bottom-up (e.g., mechanistic) and top-down (systems-level) understanding. This global, synthetic approach is needed because high biological plasticity and physiographic variation across individual ecosystems currently limit development of predictive models of forest responses to CC-AP. Integrated research on C and nutrient cycling, O3-vegetation interactions and water relations must target mechanisms' ecosystem responsiveness. Worldwide case studies must be subject to biostatistical exploration to elucidate overarching response patterns and synthesize the resulting empirical data through advanced modelling, in order to provide regionally coherent, yet globally integrated information in support of internationally coordinated decision-making and policy development., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

European beech forests growing on marginal calcareous soils have been proposed to be vulnerable to decreased soil water availability. This could result in a large-scale loss of ecological services and economical value in a changing climate. In order to evaluate the potential consequences of this drought-sensitivity, we investigated potential species range shifts for European beech forests on calcareous soil in the 21st century by statistical species range distribution modelling for present day and projected future climate conditions. We found a dramatic decline by 78% until 2080. Still the physiological or biogeochemical mechanisms underlying the drought sensitivity of European beech are largely unknown. Drought sensitivity of beech is commonly attributed to plant physiological constraints. Furthermore, it has also been proposed that reduced soil water availability could promote nitrogen (N) limitation of European beech due to impaired microbial N cycling in soil, but this hypothesis has not yet been tested. Hence we investigated the influence of simulated climate change (increased temperatures, reduced soil water availability) on soil gross microbial N turnover and plant N uptake in the beech-soil interface of a typical mountainous beech forest stocking on calcareous soil in SW Germany. For this purpose, triple 15N isotope labelling of intact beech seedling-soil-microbe systems was combined with a space-for-time climate change experiment. We found that nitrate was the dominant N source for beech natural regeneration. Reduced soil water content caused a persistent decline of ammonia oxidizing bacteria and therefore, a massive attenuation of gross nitrification rates and nitrate availability in the soil. Consequently, nitrate and total N uptake of beech seedlings were strongly reduced so that impaired growth of beech seedlings was observed already after one year of exposure to simulated climatic change. We conclude that the N cycle in this ecosystem and here specifically nitrification is vulnerable to reduced water availability, which can directly lead to nutritional limitations of beech seedlings. This tight link between reduced water availability, drought stress for nitrifiers, decreased gross nitrification rates and nitrate availability and finally nitrate uptake by beech seedlings could represent the Achilles' heel for beech under climate change stresses.

Isoprene emissions from poplar (Populus spp.) plantations can influence atmospheric chemistry and regional climate. These emissions respond strongly to temperature, [CO2], and drought, but the superimposed effect of these three climate change factors are, for the most part, unknown. Performing predicted climate change scenario simulations (periodic and chronic heat and drought spells [HDSs] applied under elevated [CO2]), we analyzed volatile organic compound emissions, photosynthetic performance, leaf growth, and overall carbon (C) gain of poplar genotypes emitting (IE) and nonemitting (NE) isoprene. We aimed (1) to evaluate the proposed beneficial effect of isoprene emission on plant stress mitigation and recovery capacity and (2) to estimate the cumulative net C gain under the projected future climate. During HDSs, the chloroplastidic electron transport rate of NE plants became impaired, while IE plants maintained high values similar to unstressed controls. During recovery from HDS episodes, IE plants reached higher daily net CO2 assimilation rates compared with NE genotypes. Irrespective of the genotype, plants undergoing chronic HDSs showed the lowest cumulative C gain. Under control conditions simulating ambient [CO2], the C gain was lower in the IE plants than in the NE plants. In summary, the data on the overall C gain and plant growth suggest that the beneficial function of isoprene emission in poplar might be of minor importance to mitigate predicted short-term climate extremes under elevated [CO2]. Moreover, we demonstrate that an analysis of the canopy-scale dynamics of isoprene emission and photosynthetic performance under multiple stresses is essential to understand the overall performance under proposed future conditions., (© 2015 American Society of Plant Biologists. All Rights Reserved.)

According to climate change scenarios, the seasonal course of temperature will change in most regions of the world, raising the question of how this will influence seasonal nitrogen (N) storage in deciduous trees. The key to this question is a detailed understanding of the underlying regulatory mechanisms, which was addressed in this study by analysing (i) the effects of low temperatures (13-1 °C) on bark storage protein (BSP) transcription, BSP and total protein accumulation and amino acid metabolism; (ii) the effects of interactions between low temperatures and photoperiod on these processes; and (iii) the regulatory role of amino acids in the bark. For this purpose, we exposed grey poplar trees (Populus × canescens) to three different treatments of changing photoperiod at constant temperature, changing temperature at constant photoperiod, and both changing photoperiod and temperature. Under a shortened photoperiod, a substantial increase of BSP transcripts was observed that was correlated with the accumulation of bark proteins, indicating a metabolic shift to promote long-term N storage. Irrespective of the applied photoperiod, exposure to low temperatures (5 or 1 °C) caused a strong increase of BSP transcripts, which was not paralled by significant increases of BSP and total bark proteins. We conclude that the interaction between effects of photoperiod and temperature is dependent on the carbon status of the trees, and reflects a metabolic adjustment of reduced carbon consumption for BSP synthesis. These results demonstrate the differential temperature sensitivity of processes involved in seasonal N storage, implying vulnerability to changing environmental conditions., (© 2012 German Botanical Society and The Royal Botanical Society of the Netherlands.)

Forest ecosystems with low soil nitrogen (N) availability are characterized by direct competition for this growth-limiting resource between several players, i.e. various components of vegetation, such as old-growth trees, natural regeneration and understorey species, mycorrhizal fungi, free-living fungi and bacteria. With the increase in frequency and intensity of extreme climate events predicted in current climate change scenarios, also competition for N between plants and/or soil microorganisms will be affected. In this review, we summarize the present understanding of ecosystem N cycling in N-limited forests and its interaction with extreme climate events, such as heat, drought and flooding. More specifically, the impacts of environmental stresses on microbial release and consumption of bioavailable N, N uptake and competition between plants, as well as plant and microbial uptake are presented. Furthermore, the consequences of drying-wetting cycles on N cycling are discussed. Additionally, we highlight the current methodological difficulties that limit present understanding of N cycling in forest ecosystems and the need for interdisciplinary studies.

Plants are known to respond to warming temperatures. Few studies, however, have included the temperature experienced by the parent plant in the experimental design, in spite of the importance of this factor for population dynamics.We investigated the phenological and growth responses of seedlings of two key temperate tree species (Fagus sylvatica and Quercus robur) to spatiotemporal temperature variation during the reproductive period (parental generation) and experimental warming of the offspring. To this end, we sampled oak and beech seedlings of different ages (1–5 years) from isolated mother trees and planted the seedlings in a common garden.Warming of the seedlings advanced bud burst in both species. In oak seedlings, higher temperatures experienced by mother trees during the reproductive period delayed bud burst in control conditions, but advanced bud burst in heated seedlings. In beech seedlings, bud burst timing advanced both with increasing temperatures during the reproductive period of the parents and with experimental warming of the seedlings. Relative diameter growth was enhanced in control oak seedlings but decreased with warming when the mother plant experienced higher temperatures during the reproductive period.Overall, oak displayed more plastic responses to temperatures than beech. Our results emphasise that temperature during the reproductive period can be a potential determinant of tree responses to climate change. [ABSTRACT FROM AUTHOR]

Bamboos are one of the fastest growing plants on Earth, and are widely considered to have high ability to capture and sequester atmospheric carbon, and consequently to mitigate climate change. We tested this hypothesis by measuring carbon dioxide ( CO2) emissions from bamboo culms and comparing them with their biomass sequestration potential. We analysed diurnal effluxes from Bambusa vulgaris culm surface and gas mixtures inside hollow sections of various bamboos using gas chromatography. Corresponding variations in gas pressure inside the bamboo section and culm surface temperature were measured. SEM micrographs of rhizome and bud portions of bamboo culms were also recorded. We found very high CO2 effluxes from culm surface, nodes and buds of bamboos. Positive gas pressure and very high concentrations of CO2 were observed inside hollow sections of bamboos. The CO2 effluxes observed from bamboos were very high compared to their carbon sequestration potential. Our measurements suggest that bamboos are net emitters of CO2 during their lifespan. [ABSTRACT FROM AUTHOR]

Prolonged summer droughts constitute to a major risk for the cultivation of beech in Central Europe. Therefore the identification of beech ecotypes that cope with growth conditions expected in future is highly desirable for forest practitioners. In a pot experiment under controlled conditions, the influence of longterm water deprivation on growth, root/shoot ratio, nitrogen (N)- and carbon (C) assimilation of seedlings from 6 European beech ( Fagus sylvatica ) provenances, originating from Central Europe (Germany), the Balkan Peninsula (Croatia), and south-east Europe (Bulgaria, Greece) were examined. Genetic diversity and relationships between the provenances were analysed by molecular markers such as nuclear EST microsatellites (EST-SSRs), SNPs and chloroplast microsatellites. Genetic diversity within provenances was high and highlighted close relationships between plants from Greece, Croatia and Germany, whereas beech from Bavaria (Germany) seemed to be admixed with genotypes from Bulgaria. Significant changes in fine root δ 13 C and C/N ratio as well as the intrinsic Water Use Efficiency (iWUE) demonstrate a better adaptability to future environmental conditions of beech ecotypes genetically related to the Greek beech population. [ABSTRACT FROM AUTHOR]

Rising temperatures and more frequent and severe climatic extremes as a consequence of climate change are expected to affect growth and distribution of tree species that are adapted to current local conditions. Species distribution models predict a considerable loss of habitats for Pinus sylvestris. These models do not consider possible intraspecific differences in response to drought and warming that could buffer those impacts. We tested 10 European provenances of P. sylvestris, from the southwestern to the central European part of the species distribution, for their response to warming and to drought using a factorial design. In this common-garden experiment the air surrounding plants was heated directly to prevent excessive soil heating, and drought manipulation, using a rain-out shelter, permitted almost natural radiation, including high light stress. Plant responses were assessed as changes in phenology, growth increment and biomass allocation. Seedlings of P. sylvestris revealed a plastic response to drought by increased taproot length and root-shoot ratios. Strongest phenotypic plasticity of root growth was found for southwestern provenances, indicating a specific drought adaptation at the cost of overall low growth of aboveground structures even under non-drought conditions. Warming had a minor effect on growth but advanced phenological development and had a contrasting effect on bud biomass and diameter increment, depending on water availability. The intraspecific variation of P. sylvestris provenances could buffer climate change impacts, although additional factors such as the adaptation to other climatic extremes have to be considered before assisted migration could become a management option. [ABSTRACT FROM AUTHOR]

Facilitation, reduced competition or increased competition can arise in mixed stands and become essential to the performance of these stands when compared to pure stands. Facilitation and over-yielding are widely held to prevail on poor sites, whereas neutral interactions or competition, leading to under-yielding of mixed versus pure stands, can occur on fertile sites. While previous studies have focused on the spatial variation of mixing effects, we examine the temporal variation of facilitation and competition and its effect on growth. The study is based on tree ring measurement on cores from increment borings from 559 trees of Norway spruce ( Picea abies [ L.] Karst.), European beech ( Fagus sylvatica [ L.]) and sessile oak ( Quercus petraea ( Matt.) Liebl.) in southern Germany, half of which were in pure stands and half in adjacent mixed stands. Mean basal area growth indices were calculated from tree ring measurements for pure and mixed stands for every species and site. The temporal variation, with positive correlations between species-specific growth indices during periods of low growth and neutral or negative correlations during periods of high growth, is more distinct in mixed than in neighbouring pure stands. We provide evidence that years with low growth trigger over-yielding of trees in mixed as opposed to pure stands, while years with high growth lead to under-yielding. We discuss the relevance of the results in terms of advancing our understanding and modelling of mixed stands, extension of the stress gradient hypothesis, and the performance of mixed versus pure stands in the face of climate change. [ABSTRACT FROM AUTHOR]

Climate-tolerant tree species and/or provenances have to be selected to ensure the high productivity of managed forests in Central Europe under the prognosticated climate changes. For this purpose, we studied the responses of saplings from three oak species ( i.e. Quercus robur, Q. petraea and Q. pubescens) and provenances of different climatic origin ( i.e. low or high rainfall, low or high temperature habitats) with regard to leaf nitrogen (N) composition as a measure of N nutrition. Saplings were grown in model ecosystems on either calcareous or acidic soil and subjected to one of four treatments (control, drought, air warming or a combination of drought and air warming). Across species, oak N metabolism responded to the influence of drought and/or air warming with an increase in leaf amino acid N concentration at the expense of structural N. Moreover, provenances or species from drier habitats were more tolerant to the climate conditions applied, as indicated by an increase in amino acid N (comparing species) or soluble protein N (comparing provenances within a species). Furthermore, amino acid N concentrations of oak leaves were significantly higher on calcareous compared to acidic soil. From these results, it can be concluded that seeds from provenances or species originating from drier habitats and - if available - from calcareous soil types may provide a superior seed source for future forest establishment. [ABSTRACT FROM AUTHOR]

Being tolerant to heat and drought, oaks are promising candidates for future forestry in view of climate change in Central Europe. Air warming is expected to increase, and drought decrease soil N availability and thus N supply to trees. Here, we conducted a model ecosystem experiment, in which mixed stands of young oaks ( Quercus robur, Q. petraea and Q. pubescens) were grown on two different soils and subjected to four climate treatments during three growing seasons: air warming by 1-2 °C, drought periods (average precipitation reduction of 43-60%), a combination of these two treatments, and a control. In contrast to our hypotheses, neither air warming nor drought significantly affected N availability, whereas total amounts, vertical distribution and availability of soil N showed substantial differences between the two soils. While air warming had no effect on tree growth and N accumulation, the drought treatment reduced tree growth and increased, or tended to increase, N accumulation in the reduced biomass, indicating that growth was not limited by N. Furthermore, 15 N-labelling revealed that this accumulation was associated with an increased uptake of nitrate. On the basis of our results, climate change effects on N dynamics are expected to be less important in oak stands than reduced soil water availability. [ABSTRACT FROM AUTHOR]

Many forest ecosystems have evolved at sites with growth-limiting nitrogen (N) availability, low N input from external sources and high ecosystem internal cycling of N. By contrast, many poplar species are frequent constituents of floodplain forests where they are exposed to a significant ecosystem external supply of N, mainly nitrate, in the moving water table. Therefore, nitrate is much more important for N nutrition of these poplar species than for many other tree species. We summarise current knowledge of nitrate uptake and its regulation by tree internal signals, as well as acquisition of ammonium and organic N from the soil. Unlike herbaceous plants, N nutrition of trees is sustained by seasonal, tree internal cycling. Recent advances in the understanding of seasonal storage and mobilisation in poplar bark and regulation of these processes by temperature and daylength are addressed. To explore consequences of global climate change on N nutrition of poplar trees, responses of N uptake and metabolism to increased atmospheric CO2 and O3 concentrations, increased air and soil temperatures, drought and salt stress are highlighted. [ABSTRACT FROM AUTHOR]

Owing to its wide European distribution and drought sensitivity, beech (Fagus sylvatica L.) has captured the attention of researchers in the context of climate change. Contrary to central European beech ecosystems, little is known about the nitrogen (N) balance of beech at its south-eastern European distribution limit. Here we studied the seasonal variation of key N-related parameters during the period 2003-2004, and tested whether climatic differences between the years affected the N balance of beech in north-western Greece. The warmer and drier climate of 2003 was characterised by lower soil N availability, compared to 2004. The pool of total soluble non-protein nitrogen (TSNN) was more sensitive than total N to interannual differences in growth conditions, as reflected particularly in the TSNN concentrations, and in the composition of phloem and xylem sap in early spring. In 2003, the N demands of expanding leaves were covered mainly by stored N remobilisation, as indicated by elevated TSNN levels in phloem and xylem sap, and by increased loading of arginine and glutamine, major N-storage compounds, in these N-exchange and transport pools. By contrast, the more favourable growth conditions of 2004 supported new growth also by current root N uptake, as indicated by increased asparagine levels in phloem and xylem sap. [ABSTRACT FROM AUTHOR]

Pedunculate oak (Quercus robur L.) was germinated and grown at ambient CO2 concentration and 650 pmol mol' CO2 in the presence and absence of the ectomycorrhizal fungus Laccaria laccata for a total of 22 weeks under non- limiting nutrient conditions. Sulphate uptake, xylem loading and exudation were analysed in excised roots. Despite a relatively high affinity for sulphate (KM = 1.6 mmol m-3), the rates of sulphate uptake by excised lateral roots of mycorrhizal oak trees were low as compared to herbaceous plants. Rates of sulphate uptake were similar in mycorrhizal and non-mycorrhizal roots and were not affected by growth of the trees at elevated CO2. However, the total uptake of sulphate per plant was enhanced by elevated CO2 and further enhanced by elevated CO2 and mycorrhization. Sulphate uptake seemed to be closely cor- related with biomass accumulation under the conditions applied. The percentage of the sulphate taken up by mycorrhizal oak roots that was loaded into the xylem was an order of magnitude lower than previously observed for herbaceous plants. The rate of xylem loading was enhanced by mycorrhization and, in roots of mycorrhizal trees only, by growth at elevated CO2. On a whole-plant basis this increase in xylem loading could only partially be explained by the increased growth of the trees. Elevated CO2 and mycorrhization appeared to increase greatly the sulphate supply of the shoot at the level of xylem loading. For all treatments, calculated rates of sulphate exudation were significantly lower than the corresponding rates of xylem loading of sulphate. Radiolabelled sulphate loaded into the xylem therefore seems to be readily diluted by unlabelled sulphate during xylem transport. Allocation of reduced sulphur from oak leaves was studied by flap-feeding radiolabelled GSH to mature oak leaves. The rate of export of radioactivity from the fed leaves was 4.5 times higher in mycorrhizal oak trees grown at elevated CO2 than in those grown at ambient CO2. Export of radiolabel proceeded almost exclusively in a basipetal direction to the roots. From these experiments it can be concluded that, in mycorrhizal oak trees grown at elevated CO2, the transport of sulphate to the shoot is increased at the level of xylem loading to enable increased sulphate reduction in the leaves. Increased sulphate reduction seems to be required for the enhanced allocation of reduced sulphur to the roots which is observed in trees grown at elevated CO2. These changes in sulphate and reduced sulphur allocation may be a prerequisite for the positive effect of elevated CO2 on growth of oak trees previously observed. [ABSTRACT FROM AUTHOR]