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Metabolomes, as chemical phenotypes of organisms, are likely not only shaped by the environment but also by common ancestry. If this is the case, we expect that closely related species of pines will tend to reach similar metabolomic solutions to the same environmental stressors. We examined the metabolomes of two sympatric subspecies of Pinus sylvestris in Sierra Nevada (southern Iberian Peninsula), in summer and winter and exposed to folivory by the pine processionary moth. The overall metabolomes differed between the subspecies but both tended to respond more similarly to folivory. The metabolomes of the subspecies were more dissimilar in summer than in winter, and iberica trees had higher concentrations of metabolites directly related to drought stress. Our results are consistent with the notion that certain plant metabolic responses associated with folivory have been phylogenetically conserved. The larger divergence between subspecies metabolomes in summer is likely due to the warmer and drier conditions that the northern iberica subspecies experience in Sierra Nevada. Our results provide crucial insights into how iberica populations would respond to the predicted conditions of climate change under an increased defoliation in the Mediterranean Basin.

SUMMARY: Anthropogenic global change is driving an increase in the frequency and intensity of drought and flood events, along with associated imbalances and limitation of several soil nutrients. In the context of an increasing human population, these impacts represent a global‐scale challenge for biodiversity conservation and sustainable crop production to ensure food security. Plants have evolved strategies to enhance uptake of soil nutrients under environmental stress conditions; for example, symbioses with fungi (mycorrhization) in the rhizosphere and the release of exudates from roots. Although crop cultivation is managed for the effects of limited availability of nitrogen (N) and phosphorus (P), there is increasing evidence for limitation of plant growth and fitness because of the low availability of other soil nutrients such as the metals potassium (K), calcium (Ca), magnesium (Mg), and iron (Fe), which may become increasingly limiting for plant productivity under global change. The roles of mycorrhizas and plant exudates on N and P uptake have been studied intensively; however, our understanding of the effects on metal nutrients is less clear and still inconsistent. Here, we review the literature on the role of mycorrhizas and root exudates in plant uptake of key nutrients (N, P, K, Ca, Mg, and Fe) in the context of potential nutrient deficiencies in crop and non‐crop terrestrial ecosystems, and identify knowledge gaps for future research to improve nutrient‐uptake capacity in food crop plants. Significance Statement: We review and synthesize the literature on the role of mycorrhizas and root exudates in plant uptake of these key nutrients (N, P, K, Ca, Mg, and Fe) in the context of potential nutrient deficiencies in crop and non‐crop terrestrial ecosystems, and identify knowledge gaps for future research to improve nutrient‐uptake capacity in food crop plants. [ABSTRACT FROM AUTHOR]

Soil enzymes are crucial in mediating ecosystems' responses to environmental drivers, so that the comprehension of their sensitivity to drivers of global change can help make predictions of future scenarios and design tailored interventions of biomanipulation. Drivers of global change usually act in combination of two or more, and indirect effects of one driver acting through modification of another one often occur, yet most of both manipulative and meta‐analysis studies available tend to focus on the direct effect of one single driver on the activity of specific soil enzymes. One of the biggest challenges is, therefore, represented by the difficulty in assessing the interactions between different drivers, due to the complexity of disentangling the single direct effects from the indirect and combined ones. In this review, after elucidating the general mechanisms of soil enzyme production and activity regulation, we display the state‐of‐the‐art knowledge on direct, indirect and combined effects of the main drivers of global change on soil enzyme activities, identify gaps in knowledge and challenges from research, plus we analyse how this can reverberate in the future of biomanipulation techniques for the improvement of ecosystem services. We conclude that qualitative but not quantitative outcomes can be predicted for some interactions such as warming + drought or warming + CO2, while for other ones, the results are controversial: future basic research will have to center on this holistic approach. A general trend toward the overall increase of soil enzyme activities and acceleration of biogeochemical cycles will persist, until an inflection will be caused by factors such as future shifts in microbial communities and changes in carbon use efficiency. Applied research will develop toward the refinement of "in situ" analytical systems for the study of soil enzyme activities and the support of bioengineering for the better tailoring of interventions of biomanipulation. [ABSTRACT FROM AUTHOR]

Calligonum mongolicum is a phreatophyte playing an important role in sand dune fixation, but little is known about its responses to drought and P fertilization. In the present study, we performed a pot experiment to investigate the effects of P fertilization under drought or well-watered conditions on multiple morpho-physio-biochemical attributes of C. mongolicum seedlings. Drought stress leads to a higher production of hydrogen peroxide (H2O2) and malondialdehyde (MDA), leading to impaired growth and metabolism. However, C. mongolicum exhibited effective drought tolerance strategies, including a higher accumulation of soluble sugars, starch, soluble protein, proline, and significantly higheractivities of peroxidase (POD) and catalase (CAT) enzymes. P fertilization increased the productivity of drought-stressed seedlings by increasing their growth, assimilative shoots relative water content, photosynthetic pigments, osmolytes accumulation, mineral nutrition, N assimilation, and reduced lipid peroxidation. Our findings suggest the presence of soil high P depletion and C. mongolicum high P requirements during the initial growth stage. Thus, P can be utilized as a fertilizer to enhance the growth and productivity of Calligonum vegetation and to reduce the fragility of the hyper-arid desert of Taklamakan in the context of future climate change. [ABSTRACT FROM AUTHOR]

More efficient use of soil resources, such as nitrogen (N) and phosphorus (P), can improve plant community resistance and resilience against drought in arid and semi-arid lands. Intercropping of legume and non-legumes can be an effective practice for enhancing P mineralization uptake, and plant nutrient status. However, it remains unclear how intercropping systems using desert plant species impact soil-plant P fractions and how they affect N and water uptake capacity. Alhagi sparsifolia (a legume) and Karelinia caspia (a non-legume) are dominant plant species in the Taklamakan Desert in Xinjiang Province, China. However, there is a lack of knowledge of whether these species, when intercropped, can trigger synergistic processes and mechanisms that drive more efficient use of soil resources. Thus, in a field experiment over two years, we investigated the impact of monoculture and intercropping of these plant species on soil-plant P fractions and soil-plant nutrients. Both plant species' foliar nutrient (N, P, and K) concentrations were higher under monoculture than intercropping (except K in K. caspia). Nucleic acid P was higher in the monoculture plots of A. sparsifolia, consistent with higher soil labile P, while metabolic P was higher in monoculture K. caspia, associated with higher soil moderately labile Pi. However, both species had a higher residual P percentage in the intercropping system. Soils from monoculture and intercropped plots contained similar microbial biomass carbon (MBC), but lower microbial biomass N:microbial biomass phosphorus (MBN:MBP) ratio associated with reduced N-acetylglucosaminidase (NAG) activity in the intercropped soils. This, together with the high MBC:MBN ratio in intercropping and the lack of apparent general effects of intercropping on MBC:MBP, strongly suggest that intercropping improved microbe N- but not P-use efficiency. Interestingly, while EC and SWC were higher in the soil of the K. caspia monoculture plots, EC was significantly lower in the intercropped plots. Plants obtained better foliar nutrition and soil P mineralization in monocultures than in intercropping systems. The possible positive implications of intercropping for reducing soil salinization and improving soil water uptake and microbial N-use efficiency could have advantages in the long term and its utilization should be explored further in future studies. [ABSTRACT FROM AUTHOR]

Soil phosphatase enzymes are produced by plant roots and microorganisms and play a key role in the cycling of phosphorus (P), an often‐limiting element in terrestrial ecosystems. The production of these enzymes in soil is the most important biological strategy for acquiring phosphate ions from organic molecules. Previous works showed how soil potential phosphatase activity is mainly driven by climatic conditions and soil nitrogen (N) and carbon. Nonetheless, future trends of the activity of these enzymes under global change remain little known. We investigated the influence of some of the main drivers of change on soil phosphatase activity using a meta‐analysis of results from 97 published studies. Our database included a compilation of N and P fertilization experiments, manipulation experiments with increased atmospheric CO2 concentration, warming, and drought, and studies comparing invaded and non‐invaded ecosystems. Our results indicate that N fertilization leads to higher phosphatase activity, whereas P fertilization has the opposite effect. The rise of atmospheric CO2 levels or the arrival of invasive species also exhibits positive response ratios on the activity of soil phosphatases. However, the occurrence of recurrent drought episodes decreases the activity of soil phosphatases. Our analysis did not reveal statistically significant effects of warming on soil phosphatase activity. In general, soil enzymatic changes in the reviewed experiments depended on the initial nutrient and water status of the ecosystems. The observed patterns evidence that future soil phosphatase activity will not only depend on present‐day soil conditions but also on potential compensations or amplifications among the different drivers of global change. The responses of soil phosphatases to the global change drivers reported in this study and the consideration of cost–benefit approaches based on the connection of the P and N cycle will be useful for a better estimation of phosphatase production in carbon (C)–N–P models. [ABSTRACT FROM AUTHOR]

The increase in aridity, mainly by decreases in precipitation but also by higher temperatures, is likely the main threat to the diversity and survival of Mediterranean forests. Changes in land use, including the abandonment of extensive crop activities, mainly in mountains and remote areas, and the increases in human settlements and demand for more resources with the resulting fragmentation of the landscape, hinder the establishment of appropriate management tools to protect Mediterranean forests and their provision of services and biodiversity. Experiments and observations indicate that if changes in climate, land use and other components of global change, such as pollution and overexploitation of resources, continue, the resilience of many forests will likely be exceeded, altering their structure and function and changing, mostly decreasing, their capacity to continue to provide their current services. A consistent assessment of the impacts of the changes, however, remains elusive due to the difficulty of obtaining simultaneous and complete data for all scales of the impacts in the same forests, areas and regions. We review the impacts of climate change and other components of global change and their interactions on the terrestrial forests of Mediterranean regions, with special attention to their impacts on ecosystem services. Management tools for counteracting the negative effects of global change on Mediterranean ecosystem- services are finally discussed. Keywords: climate change; drought; global change; mediterranean forests; ecosystem services; resilience; management; migration; extinction; diversity; communities; CO2; plant invasion [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Plants in natural environments are increasingly being subjected to a combination of abiotic stresses, such as drought and warming, in many regions. The effects of each stress and the combination of stresses on the functioning of shoots and roots have been studied extensively, but little is known about the simultaneous metabolome responses of the different organs of the plant to different stresses acting at once., We studied the shift in metabolism and elemental composition of shoots and roots of two perennial grasses, Holcus lanatus and Alopecurus pratensis, in response to simultaneous drought and warming., These species responded differently to individual and simultaneous stresses. These responses were even opposite in roots and shoots. In plants exposed to simultaneous drought and warming, terpenes, catechin and indole acetic acid accumulated in shoots, whereas amino acids, quinic acid, nitrogenous bases, the osmoprotectants choline and glycine betaine, and elements involved in growth (nitrogen, phosphorus and potassium) accumulated in roots. Under drought, warming further increased the allocation of primary metabolic activity to roots and changed the composition of secondary metabolites in shoots., These results highlight the plasticity of plant metabolomes and stoichiometry, and the different complementary responses of shoots and roots to complex environmental conditions. [ABSTRACT FROM AUTHOR]

Aim Potassium ( K) is the second most abundant nutrient in plant photosynthetic tissues after nitrogen ( N). Thousands of physiological and metabolic studies in recent decades have established the fundamental role of K in plant function, especially in water-use efficiency and economy, and yet macroecological studies have mostly overlooked this nutrient. Methods We have reviewed available studies on the content, stoichiometry and roles of K in the soil-plant system and in terrestrial ecosystems. We have also reviewed the impacts of global change drivers on K content, stoichiometry and roles. Conclusions The current literature indicates that K, at a global level, is as limiting as N and phosphorus ( P) for plant productivity in terrestrial ecosystems. Some degree of K limitation has been seen in up to 70% of all studied terrestrial ecosystems. However, in some areas atmospheric K deposition from human activities is greater than that from natural sources. We are far from understanding the K fluxes between the atmosphere and land, and the role of anthropogenic activities in these fluxes. The increasing aridity expected in wide areas of the world makes K more critical through its role in water-use efficiency. N deposition exerts a strong impact on the ecosystem K cycle, decreasing K availability and increasing K limitation. Plant invasive success is enhanced by higher soil K availability, especially in environments without strong abiotic stresses. The impacts of other drivers of global change, such as increasing atmospheric CO2 or changes in land use, remain to be elucidated. Current models of the responses of ecosystems and carbon storage to projected global climatic and atmospheric changes are now starting to consider N and P, but they should also consider K, mostly in arid and semi-arid ecosystems. [ABSTRACT FROM AUTHOR]

At the molecular level, folivory activity on plants has mainly been related to the foliar concentrations of nitrogen (N) and/or particular metabolites., We studied the responses of different nutrients and the whole metabolome of Quercus ilex to seasonal changes and to moderate field experimental conditions of drought, and how this drought may affect folivory activity, using stoichiometric and metabolomic techniques., Foliar potassium ( K) concentrations increased in summer and consequently led to higher foliar K : phosphorus ( P) and lower carbon ( C) : K and N : K ratios. Foliar N : P ratios were not lowest in spring as expected by the growth rate hypothesis. Trees exposed to moderate drought presented higher concentrations of total sugars and phenolics and these trees also experienced more severe folivory attack., The foliar increases in K, sugars and antioxidant concentrations in summer, the driest Mediterranean season, indicated enhanced osmoprotection under natural drought conditions. Trees under moderate drought also presented higher concentrations of sugars and phenolics; a plant response to avoid water loss. These shifts in metabolism produced an indirect relationship between increased drought and folivory activity. [ABSTRACT FROM AUTHOR]

ABSTRACT Hydraulic lift, water movement from deep to upper soil layers by roots, is a widespread process in temperate and semi-arid environments. It can contribute 17-81% of total water transpired and favour the uptake of nutrients available mainly from soil organic matter decomposition (e.g. N). Downward siphoning, water movement from upper to deep soil layers, can represent 10-60% of total transpired water, favouring the uptake of nutrients supplied mainly from the leaching of bedrock minerals (e.g. P and K). These vertical water movements also can affect the N : P ratio of runoff waters when, in the case of hydraulic lift, they open the possibility for a given pulse of water to circulate multiple times across the N-rich upper soil layers. Plants, thus, affect the stoichiometry of nutrients in soils and groundwater not only through the physical protection of the soil and through the water uptake but also through water redistribution. Soil water redistribution can also play an outstanding role in the ecosystem responses to global change drivers. The increase in soil patchiness in current and future arid lands modifies runoff fluxes, hydraulic lift and downward siphoning, allowing plants to dispose of higher water and nutrient availabilities. The higher use of hydraulic lift and/or downward siphoning by alien species is a possible cause of alien plant success. Further mechanistic and quantitative research is thus warranted to discern the plant role in water and nutrient cycling and in the responses to global change. Copyright © 2014 John Wiley & Sons, Ltd. [ABSTRACT FROM AUTHOR]

We review the evidence of how organisms and populations are currently responding to climate change through phenotypic plasticity, genotypic evolution, changes in distribution and, in some cases, local extinction. Organisms alter their gene expression and metabolism to increase the concentrations of several antistress compounds and to change their physiology, phenology, growth and reproduction in response to climate change. Rapid adaptation and microevolution occur at the population level. Together with these phenotypic and genotypic adaptations, the movement of organisms and the turnover of populations can lead to migration toward habitats with better conditions unless hindered by barriers. Both migration and local extinction of populations have occurred. However, many unknowns for all these processes remain. The roles of phenotypic plasticity and genotypic evolution and their possible trade-offs and links with population structure warrant further research. The application of omic techniques to ecological studies will greatly favor this research. It remains poorly understood how climate change will result in asymmetrical responses of species and how it will interact with other increasing global impacts, such as N eutrophication, changes in environmental N : P ratios and species invasion, among many others. The biogeochemical and biophysical feedbacks on climate of all these changes in vegetation are also poorly understood. We here review the evidence of responses to climate change and discuss the perspectives for increasing our knowledge of the interactions between climate change and life. [ABSTRACT FROM AUTHOR]

Abstract: This study evaluated the change induced by the year season and by experimentally induced drought on foliar element stoichiometry of the predominant woody species (Quercus ilex and Erica multiflora) in two Mediterranean ecosystems, a forest and a shrubland. This study is based in two long-term (11 yr) field experiments that simulated drought throughout the annual cycle. The effects of experimental droughts were significant but weaker than the changes produced by ontogeny and seasonality. Leaf N and P concentrations were higher in spring (the main growing season) in E. multiflora and, in Q. ilex in autumn (a period of additional growth). Leaf N:P ratios were lower in spring. In Q. ilex, the highest leaf K concentrations and leaf K:P ratios, and the lowest leaf C:K and N:K ratios, occurred in summer, the season when water stress was greatest. In E. multiflora, leaf K concentrations and K:P ratios were highest, and leaf C:K and N:K ratios were lowest in the plants from the drought-treated plots. The plant capacity to change K concentrations in response to seasonality and to drought is at least as great as the capacity to change N and P concentrations. The results underscore the importance of K and its stoichiometry relative to C, N and P in dry environments. These results indicate first, that N:P ratio shifts are not uniquely related to growth rate in Mediterranean plants but also to drought, and second, that there is a need to take into account K in ecological stoichiometry studies of terrestrial plants. [Copyright &y& Elsevier]

We investigated the effects of warming and drought on C and N concentrations, nitrogen use efficiency (NUE), and C and N accumulation in different ecosystem compartments. We conducted a 6-year (1999–2005) field experiment to simulate the climate conditions projected by IPCC models for the coming decades in a Mediterranean shrubland. We studied the two dominant species, Globularia alypum and Erica multiflora, and an N-fixing species, Dorycnium pentaphyllum, also abundant in this shrubland. Warming (1 °C) decreased N leaf concentrations by 25% and increased N stem concentrations by 40% in G. alypum. Although warming changed the available ammonium in soil in some seasons, it did not increase total soil N contents. Drought (19% average reduction in soil moisture) decreased leaf N concentrations in the two dominant shrub species, E. multiflora and G. alypum by 16% and 19%, respectively, and increased stem N concentrations by 56% and 40%, respectively. Neither warming nor drought changed the leaf N concentrations in the N-fixing species D. pentaphyllum, although warming increased stem N concentration by 9%. In G. alypum, the increase of stem N concentrations contributed to the observed increase of N accumulation in stem biomass in drought treatments with respect to control plots (8 kg N ha−1). Neither warming nor drought changed NUE in the period 1999–2005. Warming increased soil organic C relative to drought. The effects of warming and drought on C and N concentrations, on N accumulation and on leaf/stem N distribution were not the result of dilution or concentration effects produced by changes in biomass accumulation. Other factors such as the changes in soil N availability, photosynthetic capacity, and plant internal C and N remobilization must be involved. These changes which differed depending on the species and the plant tissue show that the climate change projected for the coming decades will have significant effects on the C and N cycle and stoichiometry, with probable implications for ecosystem structure and function, such as changes in plant–herbivore relationships, decomposition rates or community species composition. [ABSTRACT FROM AUTHOR]

Forest ecosystems in the Mediterranean Basin are mostly situated in the north of the Basin (mesic). In the most southern and dry areas, the forest can only exist where topography and/or altitude favor a sufficient availability of water to sustain forest biomass. We have conducted a thorough review of recent literature (2000–2021) that clearly indicates large direct and indirect impacts of increasing drought conditions on the forests of the Mediterranean Basin, their changes in surface and distribution areas, and the main impacts they have suffered. We have focused on the main trends that emerge from the current literature and have highlighted the main threatens and management solution for the maintenance of these forests. The results clearly indicate large direct and indirect impacts of increasing drought conditions on the forests of the Mediterranean Basin. These increasing drought conditions together with over-exploitation, pest expansion, fire and soil degradation, are synergistically driving to forest regression and dieback in several areas of this Mediterranean Basin. These environmental changes have triggered responses in tree morphology, physiology, growth, reproduction, and mortality. We identified at least seven causes of the changes in the last three decades that have led to the current situation and that can provide clues for projecting the future of these forests: (i) The direct effect of increased aridity due to more frequent and prolonged droughts, which has driven Mediterranean forest communities to the limit of their capacity to respond to drought and escape to wetter sites, (ii) the indirect effects of drought, mainly by the spread of pests and fires, (iii) the direct and indirect effects of anthropogenic activity associated with general environmental degradation, including soil degradation and the impacts of fire, species invasion and pollution, (iv) human pressure and intense management of water resources, (v) agricultural land abandonment in the northern Mediterranean Basin without adequate management of new forests, (vi) very high pressure on forested areas of northern Africa coupled with the demographic enhancement, the expansion of crops and higher livestock pressure, and the more intense and overexploitation of water resources uses on the remaining forested areas, and (vii) scarcity and inequality of human management and policies, depending on the national and/or regional governments and agencies, being unable to counteract the previous changes. We identified appropriate measures of management intervention, using the most adequate techniques and processes to counteract these impacts and thus to conserve the health, service capacity, and biodiversity of Mediterranean forests. Future policies should, moreover, promote research to improve our knowledge of the mechanisms of, and the effects on, nutrient and carbon plant-soil status concurrent with the impacts of aridity and leaching due to the effects of current changes. Finally, we acknowledge the difficulty to obtain an accurate quantification of the impacts of increasing aridity rise that warrants an urgent investment in more focused research to further develop future tools in order to counteract the negative effects of climate change on Mediterranean forests. [ABSTRACT FROM AUTHOR]

Long-term drought impacts soil microbial responses and enzymatic activity, affecting carbon budget and nutrient cycling in terrestrial ecosystems. We examined a Mediterranean holm oak forest subjected to 16 years of drought treatment to understand the effects on soil organic matter decomposition, nutrient cycling, and enzymatic activity. We compared potential and biomass-related (normalized to microbial biomass) soil enzyme activity with measurements taken 10 years earlier. Relationships between potential enzyme activity, soil moisture, temperature, nutrient availability, and plant/microbial activity were explored. The prolonged drought led to decreased potential activities of all enzymes, especially acid phosphatase, protease, and urease. However, biomass-related activities of protease, urease, and phosphatase were unaffected. Interestingly, biomass-related beta-glucosidase activity increased during dry seasons, indicating a functional adaptation for carbon acquisition during extreme dry conditions. The negative impact of drought on potential enzyme activity intensified over time, particularly during summer when drought intensity increased. Soil water availability, microbial biomass, and nutrient availability strongly influenced potential enzyme activity. Long-term drought and summer aridity led to increased substrate accumulation in the soil. However, no significant changes were observed in biomass-related activities of nitrogen and phosphorus-acquiring enzymes. This lack of change is likely attributed to a decrease in the absolute potential enzyme activity capacity, caused by a reduction in microbial biomass in drought-affected plots, thereby favoring substrate accumulation. Specific functional adaptations were observed, including increased carbon acquisition by soil microbes during extreme summer drought. Long-term water scarcity in water-limited ecosystems diminishes the system's capacity to acquire resources through enzyme production, impacting mineralization and nutrient dynamics. Future climate scenarios may entail reduced ecosystem-level mineralization, carbon and nitrogen shifts from plants to soil, compromising plant control over nutrients, and increasing the risk of resource loss through leaching and erosion. • Long-term drought reduces microbial biomass and potential enzyme activity. • Drought does not affect mass-specific enzymatic activities in general. • Drought alters carbon acquisition strategies in summer. • Negative impact of drought on enzyme activity intensifies over time. • Drought reduces system's capacity to acquire resources through enzymes. [ABSTRACT FROM AUTHOR]

• There is a clear biogeochemical niche (BN) discrimination among pine populations. • Pine populations may shrink the BN range with increasing aridity and decreasing soil nutrient availability. • Tissue stoichiometric diversity is negatively related to the reproductive traits of pine trees at the population levels. Elemental stoichiometry has recently been integrated with trait diversity metrics to explore the effects of relative balance of multiple elements on environmental fitness of species from the perspective of biogeochemical niche (BN). Nevertheless, there is relatively little knowledge about the variability of intraspecific BN in response to environmental changes as well as how BN flexibility is related to the ecophysiological traits of species. Here we detected the BN shifts of Pinus sylvestris var. mongolica populations along an environmental gradient in northeastern China, with differences in aridity severity and soil nutrient availability. Meanwhile, we also assessed whether the BN characteristics observed are associated with the growth and reproduction of pine trees at the population level. The results indicate that the elemental and stoichiometric compositions of pine trees differed significantly, implying a BN discrimination among distinct populations. The intra-population individual stoichiometric diversity reduced with increasing aridity and decreasing soil nutrient availability, indicating the contraction of BN with increasing environmental stress. Additionally, the results also revealed a generally negative correlation between the tissue stoichiometric diversity and the reproductive traits of pine trees among different sets of populations. For example, we found pine stoichiometric richness and divergence were negatively related to cone weight and volume at the population level. The current work thus provides empirical evidence of a highly contractive BN space formed by a small set of optimal elemental phenotypes appears as a resource of species populations to respond to more stressed environments. The intraspecific BN variabilities caused by individual elemental differences are probably related to the functional performance of species. [ABSTRACT FROM AUTHOR]