Impact of interspecific competition and drought on the allocation of new assimilates in trees

In trees, the interplay between reduced carbon assimilation and the inability to transport carbohydrates to the sites of demand under drought might be one of the mechanisms leading to carbon starvation. However, we largely lack knowledge on how drought effects on new assimilate allocation differ between species with different drought sensitivities and how these effects are modified by interspecific competition. We assessed the fate of ¹³C labelled assimilates in above‐ and belowground plant organs and in root/rhizosphere respired CO₂ in saplings of drought‐tolerant Norway maple (Acer platanoides) and drought‐sensitive European beech (Fagus sylvatica) exposed to moderate drought, either in mono‐ or mixed culture. While drought reduced stomatal conductance and photosynthesis rates in both species, both maintained assimilate transport belowground. Beech even allocated more new assimilate to the roots under moderate drought compared to non‐limited water supply conditions, and this pattern was even more pronounced under interspecific competition. Even though maple was a superior competitor compared to beech under non‐limited soil water conditions, as indicated by the changes in above‐ and belowground biomass of both species in the interspecific competition treatments, we can state that beech was still able to efficiently allocate new assimilate belowground under combined drought and interspecific competition. This might be seen as a strategy to maintain root osmotic potential and to prioritise root functioning. Our results thus show that beech tolerates moderate drought stress plus competition without losing its ability to supply belowground tissues. It remains to be explored in future work if this strategy is also valid during long‐term drought exposure.     

Changes in nutrient concentrations of leaves and roots in response to global change factors

Global change impacts on biogeochemical cycles have been widely studied, but our understanding of whether the responses of plant elemental composition to global change drivers differ between above‐ and belowground plant organs remains incomplete. We conducted a meta‐analysis of 201 reports including 1,687 observations of studies that have analyzed simultaneously N and P concentrations changes in leaves and roots in the same plants in response to drought, elevated [CO₂], and N and P fertilization around the world, and contrasted the results within those obtained with a general database (838 reports and 14,772 observations) that analyzed the changes in N and P concentrations in leaves and/or roots of plants submitted to the commented global change drivers. At global level, elevated [CO₂] decreased N concentrations in leaves and roots and decreased N:P ratio in roots but no in leaves, but was not related to P concentration changes. However, the response differed among vegetation types. In temperate forests, elevated [CO₂] was related with lower N concentrations in leaves but not in roots, whereas in crops, the contrary patterns were observed. Elevated [CO₂] decreased N concentrations in leaves and roots in tundra plants, whereas not clear relationships were observed in temperate grasslands. However, when elevated [CO₂] and N fertilization coincided, leaves had lower N concentrations, whereas root had higher N concentrations suggesting that more nutrients will be allocated to roots to improve uptake of the soil resources not directly provided by the global change drivers. N fertilization and drought increased foliar and root N concentrations while the effects on P concentrations were less clear. The changes in N and P allocation to leaves and root, especially those occurring in opposite direction between them have the capacity to differentially affect above‐ and belowground ecosystem functions, such as litter mineralization and above‐ and belowground food webs.     

Growth and photosynthetic responses in Jatropha curcas L. seedlings of different provenances to watering regimes

Seedlings from four provenances of Jatropha curcas were subjected to 80, 50, and 30% of soil field capacity in potted experiments in order to study their responses to water availability. Our results showed that with the decline of soil water availability, plant growth, biomass accumulation, net photosynthetic rate, stomatal conductance (g_s), and transpiration rate (E) decreased, whereas leaf carbon isotope composition (δ¹³C), leaf pigment contents, and stomatal limitation value increased, while maximal quantum yield of PSII photochemistry was not affected. Our findings proved that stomatal limitation to photosynthesis dominated in J. curcas under low water availability. The increase of δ¹³C should be attributed to the decrease in g_s and E under the lowest water supply. J. curcas could adapt to low water availability by adjusting its plant size, stomata closure, reduction of E, increasing δ¹³C, and leaf pigment contents. Moreover, effects of provenance and the interaction with the watering regime were detected in growth and many physiological parameters. The provenance from xeric habitats showed stronger plasticity in the plant size than that from other provenances under drought. The variations may be used as criteria for variety/provenance selection and improvement of J. curcas performance.     

Variation in functional leaf traits among beech provenances during a Spanish summer reflects the differences in their origin

We assessed the response of 11-year-old saplings from six beech provenances growing in a common-garden trial at the southwestern range limit. Provenances from distinct climatic regions across the European beech distribution were selected. The local Spanish provenance appeared well suited to the site conditions, maintaining high rates of assimilation even in midsummer, but so did the provenance of southern continental origin, from Gotze-Delchev, Bulgaria. Those provenances from cooler sites in central Europe, a continental mountain climate in the Czech Republic and a continental range-edge site in eastern Poland, along with a German provenance of mild maritime origin, had good physiological functionality in early summer but reduced carbon assimilation (A _area) and apparent soil?Cleaf hydraulic conductivity (K _L) in midsummer. The northern maritime provenance from Sweden demonstrated severely-reduced photosynthetic capacity. These groupings of provenances according to their photosynthetic performance, stable carbon isotope composition (??¹³C; a proxy for water-use efficiency) and leaf water potential under marginal conditions, during late summer in the trial, suggest that they have divergent strategies for water use. The research highlights large intraspecific differences among beech provenances of distinct origin and strategies which are expected to modify their response to drought, requiring future genetic studies to explicitly determine the basis of this ecophysiological differentiation.     

Tree ring isotopic composition, radial increment and height growth reveal provenance-specific reactions of Douglas-fir towards environmental parameters

In the search of timber species being tolerant towards summer droughts, which are expected to be more frequent in future, Douglas-fir is often discussed as a potential alternative for spruce in Central Europe. To assess physiological and growth reactions of Douglas-fir provenances towards climate- and weather-related environmental conditions we took advantage of a provenance trial with three sites in south-western Germany located along an elevation gradient. We examined six different provenances of Douglas-fir from North America for oxygen (δ¹⁸O) and carbon (δ¹³C) stable isotope composition in tree rings as well as for radial increment for a 7 year period and long-term height growth. Our results show that different Douglas-fir provenances clearly vary in their drought sensitivity at the driest and warmest site in the valley as shown by the radial growth decline in the extreme dry and hot year 2003. The growth decline in the provenances Pamelia Creek, Cameron Lake, Duncan Paldi and Conrad Creek could be clearly attributed to a reduction in stomatal conductance as assessed by the relations between δ¹⁸O and δ¹³C in the tree rings. These responses were not related to the long-term average climate at the places of origin of the provenances and the provenances with the lowest long-term (height) growth potential were the ones least affected in radial increment by the extreme drought of 2003. When selecting suitable Douglas-fir provenances, which are adapted to the climatic conditions projected for the future we thus might need to take into account the trade-off between the adaptation to extreme drought periods and the long-term growth performance. Site-specific evaluations of the probability of extreme drought events are thus needed to select the appropriate provenances.     

Biomass distribution of two subalpine dwarf‐shrubs in relation to soil moisture and nutrient content

Question: Do soil water content and/or soil nitrogen (N) content and/or soil phosphorus (P) content affect the biomass of Vaccinium myrtillus and V. vitis‐idaea in a sub‐alpine heath? Location: Dolomites, northern Italy, 1800 m a.s.l. Methods: We determined above‐ground and below‐ground biomass of the shrubs at three sites, each on a different substrate type. At each site, we determined soil N‐ and P‐contents. We also determined leaf water potential (Psi;₁), N‐ and P‐concentrations in plant tissues and litter, as well as δ¹³C and δ¹⁵N in mature leaves. Results: V. myrtillus biomass was highest at the silicate site, V. vitis‐idaea biomass was highest at the carbonate site. Both shrubs had low biomass at the peat site, possibly due to a toxic effect of waterlogging in wet soils. For both species, pre‐dawn Psi;₁ indicated optimal hydration and midday Psi;₁ did not show any sign of water stress. Water use efficiency (WUE) did not differ among sites for any species. Whole‐plant nutrient concentrations showed that, with increasing biomass, N was diluted in V. myrtillus tissues while P was diluted in V. vitis‐idaea tissues. Foliar N‐concentration was higher overall for V. myrtillus. Foliar P‐concentration in V. myrtillus peaked at the silicate site. Foliar N : P ratios suggested that V. myrtillus was primarily P‐limited and V. vitis‐idaea primarily N‐limited. Conclusions: Water content affected the distribution of the two shrubs in a similar way, higher P‐availability in the soil enhanced V. myrtillus rather than V. vitis‐idaea.     

Relationships of growth,stable carbon isotope composition and anatomical properties of leaf and xylem in seven mulberry cultivars: a hint towards drought tolerance

• The fast growth of mulberry depends on high water consumption, but considerable variations in drought tolerance exist across different cultivars. Physiological and anatomical mechanisms are important to plant survival under drought. However, few research efforts have been made to reveal the relationships of these two aspects in relation to drought tolerance.
• In this study, growth rates, leaf functional physiology and anatomical characteristics of leaf and xylem of 1‐year‐old saplings of seven mulberry cultivars at a common garden were compared. Their relationships were also explored.
• Growth, leaf physiology and anatomy were significantly different among the tested cultivars. Foliar stable carbon isotope composition (δ¹³C) was negatively correlated with growth rates, and closely related to several leaf and xylem anatomical traits. Particularly, leaf thickness, predicted hydraulic conductivity and vessel element length jointly contributed 77% of the variability in δ¹³C. Cultivar Wupu had small stomata, intermediate leaf thickness, the smallest hydraulically weighted vessel diameter and highest vessel number, and higher δ¹³C; Yunguo1 had high abaxial stomatal density, low specific leaf area, moderate hydraulic conductivity and δ¹³C; these are beneficial features to reduce leaf water loss and drought‐induced xylem embolism in arid areas. Cultivar Liaolu11 had contrasting physiological and anatomical traits compared with the previous two cultivars, suggesting that it might be sensitive to drought.
• Our findings indicate that growth and δ¹³C are closely associated with both leaf and xylem anatomical characteristics in mulberry, which provides fundamental information to assist evaluation of drought tolerance in mulberry cultivars and in other woody trees.

     

Drought response of five conifer species under contrasting water availability suggests high vulnerability of Norway spruce and European larch

The ability of tree species to cope with anticipated decrease in water availability is still poorly understood. We evaluated the potential of Norway spruce, Scots pine, European larch, black pine, and Douglas‐fir to withstand drought in a drier future climate by analyzing their past growth and physiological responses at a xeric and a mesic site in Central Europe using dendroecological methods. Earlywood, latewood, and total ring width, as well as the δ¹³C and δ¹⁸O in early‐ and latewood were measured and statistically related to a multiscalar soil water deficit index from 1961 to 2009. At the xeric site, δ¹³C values of all species were strongly linked to water deficits that lasted longer than 11 months, indicating a long‐term cumulative effect on the carbon pool. Trees at the xeric site were particularly sensitive to soil water recharge in the preceding autumn and early spring. The native species European larch and Norway spruce, growing close to their dry distribution limit at the xeric site, were found to be the most vulnerable species to soil water deficits. At the mesic site, summer water availability was critical for all species, whereas water availability prior to the growing season was less important. Trees at the mesic were more vulnerable to water deficits of shorter duration than the xeric site. We conclude that if summers become drier, trees growing on mesic sites will undergo significant growth reductions, whereas at their dry distribution limit in the Alps, tree growth of the highly sensitive spruce and larch may collapse, likely inducing dieback and compromising the provision of ecosystem services. However, the magnitude of these changes will be mediated strongly by soil water recharge in winter and thus water availability at the beginning of the growing season.     

Cover crop root contributions to soil carbon in a no‐till corn bioenergy cropping system

Crop residues are potential biofuel feedstocks, but residue removal may reduce soil carbon (C). The inclusion of a cover crop in a corn bioenergy system could provide additional biomass, mitigating the negative effects of residue removal by adding to stable soil C pools. In a no‐till continuous corn bioenergy system in the northern US Corn Belt, we used ¹³CO₂ pulse labeling to trace plant C from a winter rye (Secale cereale) cover crop into different soil C pools for 2 years following rye cover crop termination. Corn stover left as residue (30% of total stover) contributed 66, corn roots 57, rye shoots 61, rye roots 50, and rye rhizodeposits 25 g C m^?2 to soil. Five months following cover crop termination, belowground cover crop inputs were three times more likely to remain in soil C pools than were aboveground inputs, and much of the root‐derived C was in mineral‐associated soil fractions. After 2 years, both above‐ and belowground inputs had declined substantially, indicating that the majority of both root and shoot inputs are eventually mineralized. Our results underscore the importance of cover crop roots vs. shoots and the importance of cover crop rhizodeposition (33% of total belowground cover crop C inputs) as a source of soil C. However, the eventual loss of most cover crop C from these soils indicates that cover crops will likely need to be included every year in rotations to accumulate soil C.     

不同种源栓皮栎幼苗叶片水分关系和保护酶活性对干旱胁迫的响应

 为了鉴别不同种源的栓皮栎（Quercus variabilis）耐旱性,对4个种源的3年生盆栽幼苗进行了控制条件下的土壤干旱胁迫实验,系统测定了超氧化物歧化酶(SOD)和过氧化氢酶(CAT)活性、MDA（丙二醛）含量、膜透性,以及叶片P-V曲线导出的水分关系参数（Ψπsat、Ψπtlp、WSDtlp、εmax）。结果表明：在土壤干旱胁迫下,种源4（黄龙）和种源3 (秦岭北坡) 抗氧化能力较强, 种源2 (伏牛山)居中, 种源1（巴山）的表现最差。干旱胁迫使各种源 MDA的含量及膜透性均有不同程度的提高,其中种源4的MDA含量及膜透性比较平稳, 种源1变化剧烈。在干旱胁迫下,各种源渗透调节和保持膨压的能力均有提高,但存在差异,以种源4和种源3表现较好,种源2居中,种源1较差。应用Fuzzy方法,对不同种源的叶片水分关系参数、保护酶活性和膜透性在干旱胁迫中的动态进行综合评判,认为不同种源的耐旱性强度次序为：种源4>种源3>种源2>种源1。其中种源4与种源1和种源2差异显著(p<0.05)。     

Carbon cost of plant nitrogen acquisition: global carbon cycle impact from an improved plant nitrogen cycle in the Community Land Model

Plants typically expend a significant portion of their available carbon (C) on nutrient acquisition – C that could otherwise support growth. However, given that most global terrestrial biosphere models (TBMs) do not include the C cost of nutrient acquisition, these models fail to represent current and future constraints to the land C sink. Here, we integrated a plant productivity‐optimized nutrient acquisition model – the Fixation and Uptake of Nitrogen Model – into one of the most widely used TBMs, the Community Land Model. Global plant nitrogen (N) uptake is dynamically simulated in the coupled model based on the C costs of N acquisition from mycorrhizal roots, nonmycorrhizal roots, N‐fixing microbes, and retranslocation (from senescing leaves). We find that at the global scale, plants spend 2.4 Pg C yr^?1 to acquire 1.0 Pg N yr^?1, and that the C cost of N acquisition leads to a downregulation of global net primary production (NPP) by 13%. Mycorrhizal uptake represented the dominant pathway by which N is acquired, accounting for ~66% of the N uptake by plants. Notably, roots associating with arbuscular mycorrhizal (AM) fungi – generally considered for their role in phosphorus (P) acquisition – are estimated to be the primary source of global plant N uptake owing to the dominance of AM‐associated plants in mid‐ and low‐latitude biomes. Overall, our coupled model improves the representations of NPP downregulation globally and generates spatially explicit patterns of belowground C allocation, soil N uptake, and N retranslocation at the global scale. Such model improvements are critical for predicting how plant responses to altered N availability (owing to N deposition, rising atmospheric CO₂, and warming temperatures) may impact the land C sink.     

Fagus sylvatica L. provenances maintain different leaf metabolic profiles and functional response

Most temperate forest tree species will suffer important environmental changes as result of the climate change. Adaptiveness to local conditions could change at different sites in the future. In this context, the study of intra-specific variability is important to clarify the singularity of different local populations. Phenotypic differentiation between three beech provenances covering a wide latitudinal range (Spain/ES, Germany/DE and Sweden/SE), was studied in a greenhouse experiment. Non-target leaf metabolite profiles and ecophysiological response was analyzed in well-watered and water stressed seedlings. There was a provenance-specific pattern in the relative concentrations of some leaf metabolites regardless watering treatment. The DE and SE from the center and north of the distribution area of the species showed a clear differentiation from the ES provenance in the relative concentration of some metabolites. Thus the ES provenance from the south maintained larger relative concentration of some organic and amino acids (e.g. fumaric and succinic acids or valine and isoleucine), and in some secondary metabolites (e.g. kaempferol, caffeic and ferulic acids). The ecophysiological response to mild water stress was similar among the three provenances as a consequence of the moderate water stress applied to seedlings, although leaf N isotope composition (δ¹⁵N) and leaf C:N ratio were higher and lower respectively in DE than in the other two provenances. This would suggest potential differences in the capacity to uptake and post-process nitrogen according to provenance. An important focus of the study was to address for the first time inter-provenance leaf metabolic diversity in beech from a non-targeted metabolic profiling approach that allowed differentiation of the three studied provenances.     

Provenance selection and site conditions determine growth performance of pedunculate oak

Forests of the future need to cope with adverse climatic conditions, in particular drought, to ensure forest stability and productivity. Given the usually long rotation period of forests, forest managers need to select appropriate, i.e. productive and climate-change resilient tree species and/or provenances, to lower tree-mortality risks and sustain current wood production rates at the end of the 21^st century. A frequent means of assessing which provenances of a given species are adapted to anticipated climate conditions is common garden experiments, where trees from different provenances are planted under similar climate conditions. However, in this context soil conditions also play an important role, since they govern how climate translates into plant-available water and hence plant’s ability to cope with extreme drought events. Here, we examine the effects of soil conditions on pedunculate oak (Quercus robur L.), by studying 10 different Dutch oak provenances that were planted in three provenance trials on different soil types in 1988 in the Netherlands. Using dendroecological methods, we quantified provenance-specific productivity and assessed provenance- and site-specific growth patterns. Our results indicated clear differences in productivity among provenances as well as soil-type specific growth patterns. Consequently, our study highlights the importance of incorporating soil characteristics when evaluating the growth performance of provenances within common garden experiments.     

Carbon (δ13C) and nitrogen (δ15N) stable isotope composition in plant and soil in Southern Patagonia's native forests

Stable isotope natural abundance measurements integrate across several biogeochemical processes in ecosystem N and C dynamics. Here, we report trends in natural isotope abundance (δ¹³C and δ¹⁵N in plant and soil) along a climosequence of 33 Nothofagus forest stands located within Patagonia, Southern Argentina. We measured 28 different abiotic variables (both climatic variables and soil properties) to characterize environmental conditions at each of the 33 sites. Foliar δ¹³C values ranged from ?35.4‰ to ?27.7‰, and correlated positively with foliar δ¹⁵N values, ranging from ?3.7‰ to 5.2‰. Soil δ¹³C and δ¹⁵N values reflected the isotopic trends of the foliar tissues and ranged from ?29.8‰ to ?25.3‰, and ?4.8‰ to 6.4‰, respectively, with no significant differences between Nothofagus species (Nothofagus pumilio, Nothofagus antarctica, Nothofagus betuloides). Principal component analysis and multiple regressions suggested that mainly water availability variables (mean annual precipitation), but not soil properties, explained between 42% and 79% of the variations in foliar and soil δ¹³C and δ¹⁵N natural abundance, which declined with increased moisture supply. We conclude that a decline in water use efficiency at wetter sites promotes both the depletion of heavy C and N isotopes in soil and plant biomass. Soil δ¹³C values were higher than those of the plant tissues and this difference increased as annual precipitation increased. No such differences were apparent when δ¹⁵N values in soil and plant were compared, which indicates that climatic differences contributed more to the overall C balance than to the overall N balance in these forest ecosystems.     

Foliar δ15N is affected by foliar nitrogen uptake, soil nitrogen, and mycorrhizae along a nitrogen deposition gradient

Foliar nitrogen isotope (δ¹⁵N) composition patterns have been linked to soil N, mycorrhizal fractionation, and within-plant fractionations. However, few studies have examined the potential importance of the direct foliar uptake of gaseous reactive N on foliar δ¹⁵N. Using an experimental set-up in which the rate of mycorrhizal infection was reduced using a fungicide, we examined the influence of mycorrhizae on foliar δ¹⁵N in potted red maple (Acer rubrum) seedlings along a regional N deposition gradient in New York State. Mycorrhizal associations altered foliar δ¹⁵N values in red maple seedlings from 0.06 to 0.74 ‰ across sites. At the same sites, we explored the predictive roles of direct foliar N uptake, soil δ¹⁵N, and mycorrhizae on foliar δ¹⁵N in adult stands of A. rubrum, American beech (Fagus grandifolia), black birch (Betula lenta), and red oak (Quercus rubra). Multiple regression analysis indicated that ambient atmospheric nitrogen dioxide (NO₂) concentration explained 0, 69, 23, and 45 % of the variation in foliar δ¹⁵N in American beech, red maple, red oak, and black birch, respectively, after accounting for the influence of soil δ¹⁵N. There was no correlation between foliar δ¹³C and foliar %N with increasing atmospheric NO₂ concentration in most species. Our findings suggest that total canopy uptake, and likely direct foliar N uptake, of pollution-derived atmospheric N deposition may significantly impact foliar δ¹⁵N in several dominant species occurring in temperate forest ecosystems.     

Does geographic origin dictate ecological strategies in Acacia senegal (L.) Willd.? Evidence from carbon and nitrogen stable isotopes

Background and aims

Acacia senegal, a leguminous dryland tree, is economically and ecologically important to sub-Saharan Africa. Water-use efficiency (WUE) and biological nitrogen fixation (BNF) are fundamental to plant productivity and survival. We quantify provenance differences in WUE, BNF, photosynthesis, biomass and gum arabic production from A. senegal assessing genetic improvement potential.

Methods

Using stable isotope ratios, we determined WUE (δ¹³C) and BNF (δ¹⁵N) from provenances of mature A. senegal in field-trials (Senegal), sampling leaves at the beginning (wet) and end (dry) of the rainy season. Seedling provenance trials (UK) determined photosynthesis, and biomass and δ¹³C in relation to water table. Environmental data were characterised for all provenances at their sites of origin.

Results

Provenances differed in both δ¹³C and δ¹⁵N. Gum yield declined with increasing WUE. Virtually no BNF was detected during the dry season and seedlings and mature trees may have different WUE strategies. Wind speed and soil characteristics at provenance origin were correlated with isotope composition and gum production.

Conclusion

Provenance differences suggest that selection for desirable traits, e.g., increased gum production, may be possible. As ecological strategies relate to native locality, the environmental conditions at plantation site and provenance origin are important in assessing selection criteria.     

Symbiotic nitrogen fixation in eight Acacia senegal provenances in dryland clays of the Blue Nile Sudan estimated by the 15N natural abundance method

The symbiotic biological N₂fixation by Acacia senegal was estimated using the ¹⁵N natural abundance (δ ¹⁵N) procedure on eight provenances collected from different environments and soil types grown in a clay soil in the Blue Nile region, Sudan. Balanites aegyptiaca (a non-legume) was used as a non-N₂-fixing reference plant to allow ¹⁵N-based estimates of the proportion of the Acacia N derived from atmospheric N₂ (N_dfa) to be calculated. Results show variation in leaf δ ¹⁵N between A. senegal and the reference plant and among years. The relative δ ¹⁵N values (‰) were higher in B. aegyptiaca than in the N₂-fixing acacia provenances. Provenances originally collected from clay soils fixed little N in the first year, but the amount fixed increased as the trees aged. All provenances showed a decrease in δ ¹⁵N with age. The N_dfa varied between 24% (Mazmoom provenance) and 61% (Rahad provenance) 4 years after planting. There was no significant difference in δ ¹⁵N between provenance groups based on soil type or rainfall at original growing site. The amount of N_dfa increased significantly with age in all provenances. The above-ground contribution of fixed N to foliage growth in a 4-year-old A. senegal was highest in the Rahad sand–soil provenance (46.7 kg N ha⁻¹) and lowest in the Mazmoom clay-soil provenance (28.7 kg N ha⁻¹). Our study represents the first use of the δ ¹⁵N method for estimating the N input by A. senegal to the clay plain soils of the gum belt in the Sudan.     

Forest understory plant and soil microbial response to an experimentally induced drought and heat‐pulse event: the importance of maintaining the continuum

Drought duration and intensity are expected to increase with global climate change. How changes in water availability and temperature affect the combined plant–soil–microorganism response remains uncertain. We excavated soil monoliths from a beech (Fagus sylvatica L.) forest, thus keeping the understory plant–microbe communities intact, imposed an extreme climate event, consisting of drought and/or a single heat‐pulse event, and followed microbial community dynamics over a time period of 28 days. During the treatment, we labeled the canopy with ¹³CO₂ with the goal of (i) determining the strength of plant–microbe carbon linkages under control, drought, heat and heat–drought treatments and (ii) characterizing microbial groups that are tightly linked to the plant–soil carbon continuum based on ¹³C‐labeled PLFAs. Additionally, we used 16S rRNA sequencing of bacteria from the Ah horizon to determine the short‐term changes in the active microbial community. The treatments did not sever within‐plant transport over the experiment, and carbon sinks belowground were still active. Based on the relative distribution of labeled carbon to roots and microbial PLFAs, we determined that soil microbes appear to have a stronger carbon sink strength during environmental stress. High‐throughput sequencing of the 16S rRNA revealed multiple trajectories in microbial community shifts within the different treatments. Heat in combination with drought had a clear negative effect on microbial diversity and resulted in a distinct shift in the microbial community structure that also corresponded to the lowest level of label found in the PLFAs. Hence, the strongest changes in microbial abundances occurred in the heat–drought treatment where plants were most severely affected. Our study suggests that many of the shifts in the microbial communities that we might expect from extreme environmental stress will result from the plant–soil–microbial dynamics rather than from direct effects of drought and heat on soil microbes alone.     

Interactions between plant growth and soil nutrient cycling under elevated CO2: a meta-analysis

free air carbon dioxide enrichment (FACE) and open top chamber (OTC) studies are valuable tools for evaluating the impact of elevated atmospheric CO₂ on nutrient cycling in terrestrial ecosystems. Using meta‐analytic techniques, we summarized the results of 117 studies on plant biomass production, soil organic matter dynamics and biological N₂ fixation in FACE and OTC experiments. The objective of the analysis was to determine whether elevated CO₂ alters nutrient cycling between plants and soil and if so, what the implications are for soil carbon (C) sequestration. Elevated CO₂ stimulated gross N immobilization by 22%, whereas gross and net N mineralization rates remained unaffected. In addition, the soil C : N ratio and microbial N contents increased under elevated CO₂ by 3.8% and 5.8%, respectively. Microbial C contents and soil respiration increased by 7.1% and 17.7%, respectively. Despite the stimulation of microbial activity, soil C input still caused soil C contents to increase by 1.2% yr^?1. Namely, elevated CO₂ stimulated overall above‐ and belowground plant biomass by 21.5% and 28.3%, respectively, thereby outweighing the increase in CO₂ respiration. In addition, when comparing experiments under both low and high N availability, soil C contents (+2.2% yr^?1) and above‐ and belowground plant growth (+20.1% and+33.7%) only increased under elevated CO₂ in experiments receiving the high N treatments. Under low N availability, above‐ and belowground plant growth increased by only 8.8% and 14.6%, and soil C contents did not increase. Nitrogen fixation was stimulated by elevated CO₂ only when additional nutrients were supplied. These results suggest that the main driver of soil C sequestration is soil C input through plant growth, which is strongly controlled by nutrient availability. In unfertilized ecosystems, microbial N immobilization enhances acclimation of plant growth to elevated CO₂ in the long‐term. Therefore, increased soil C input and soil C sequestration under elevated CO₂ can only be sustained in the long‐term when additional nutrients are supplied.     

Effects of temperature and drought manipulations on seedlings of Scots pine provenances

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.