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Root exudates are well-known 'labile' sources of soil carbon that can prime microbial activity. Recent investigations suggest that the stability of labile carbon inputs in soil mostly depends upon the physical, chemical, and biological properties of the surroundings. Here, we propose that, in some ecosystems, such as forests and grasslands, root exudates can function as a source of soil organic carbon (SOC) that can be stabilized through various mechanisms leading to long-term sequestration. Increasing soil carbon sequestration is important for capturing atmospheric CO2 and combating climate change issues. Thus, there is an urgent need to preserve existing ecosystems and to adopt strategies such as afforestation, reforestation, and establishment of artificial grasslands to foster carbon sequestration through higher root exudate inputs in the soil.

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. info:eu-repo/semantics/acceptedVersion

Challenges to food security under conditions of global change are forcing us to increase global crop production. Focussing on belowground plant traits, especially root exudation, has great promise to meet this challenge. Root exudation is the release of a vast array of compounds into the soil. These exudates are involved in many biotic and abiotic interactions. Wild relatives of crops provide a large potential source of information and genetic material and have desirable traits that could be incorporated into modern breeding programs. However, root exudates are currently underexploited. Here, we highlight how the traits of root exudates of crop wild relatives could be used to improve agricultural output and reduce environmental impacts, particularly by decreasing our dependence on pesticides and fertilisers.

Global climate changes such as prolonged duration and intensity of drought can lead to adverse ecological consequences in forests. Currently little is known about soil microbial community responses to such drought regimes in tropical forests. In this study, we examined the resistance and resilience of topsoil prokaryotic communities to a prolongation of the dry season in terms of diversity, community structure and co-occurrence patterns in a French Guianan tropical forest. Through excluding rainfall during and after the dry season, a simulated prolongation of the dry season by five months was compared to controls. Our results show that prokaryotic communities increasingly diverged from controls with the progression of rain exclusion. Furthermore, prolonged drought significantly affected microbial co-occurrence networks. However, both the composition and co-occurrence networks of soil prokaryotic communities immediately ceased to differ from controls when precipitation throughfall returned. This study thus suggests modest resistance but high resilience of microbial communities to a prolonged drought in tropical rainforest soils.

Differences in the elemental composition of plants, mainly C, N, and P, have been shown to be related to differences in their nutritional status, and their morphological and functional traits. The relationship between morphological traits and micronutrients and trace elements, however, has been much less studied. Additionally, in bryophytes, research devoted to investigating these relationships is still very scarce. Here, we analysed 80 samples from 29 aquatic and semi-aquatic (hygrophytic) moss species living in Mediterranean springs to investigate the relationship between moss nutrient concentrations and their micro- and macroscopic morphological traits and growth forms. We found that, across species, the elemental concentration of mosses was more tightly linked to macroscopic traits than to microscopic traits. Growth forms could also be successfully explained by the concentration of elements in mosses. Apart from macronutrients and their stoichiometric ratios (C:N, C:P, and N:P), micronutrients and trace elements were also important variables predicting moss morphological traits and growth forms. Additionally, our results showed that microscopic traits were well related to macroscopic traits. Overall, our results clearly indicate that the elemental composition of mosses can be used to infer their morphological traits, and that elements other than macronutrients should be taken into account to achieve a good representation of their morphological and, potentially, functional traits when comparing the elemental composition across species.

Sea spray aerosol (SSA) is responsible for the large-scale transfer of particles from the sea to the land, leading to significant deposition of a range of ions, predominantly Na+, K+, Mg2+, Ca2+, and Cl−. Up to now, there has been little research into the effects of SSA on spring water chemistry. Therefore, we sampled 303 semi-natural springs across Catalonia (NE Iberian Peninsula) and analysed the concentrations of 20 different ions and elements, and determined the impact of SSA (using distance to the coast as a proxy) as well as climate, lithology and human disturbances. We found that distance to the coast had a clear effect on the water chemical composition of springs, while accounting for potentially confounding factors such as anthropogenic water pollution (nitrate, NO3−), differences in lithology and annual rainfall. Our results showed that springs located closer to the coast had higher Cl−, SO42−, Na+, Mg2+, K+ and Ca2+ concentrations than those of springs located further away. Precipitation was generally negatively correlated with the concentration of almost all elements analysed. The concentration of NO3− increased with distance to the coast, concurrently with farming activities, located mainly inland in the study area. These results demonstrate that SSA has an important effect on the groundwater of coastal zones, up to a distance of around 70 km from the coastline. This analysis reveals the main natural and human processes that influence spring water chemistry in this Mediterranean region, information that could be helpful in similar regions for ecological studies, water quality policies, and for the improvement of predictions in the current context of global change.

Ecological stoichiometry and studies of biogeochemical niches have mainly focused on plankton and vascular plants, but the phenotypically closest modern relatives of early plants, bryophytes, have been largely neglected. We analysed C:N:P stoichiometries and elemental compositions (K, Na, Mg, Ca, S, Fe) of 35 widely distributed bryophyte species inhabiting springs. We estimated bryophyte C:N:P ratios and their biogeochemical niches, investigated how elementomes respond to the environment and determined whether they tend to diverge more for coexisting than non-coexisting individuals and species. The median C:N:P was 145:8:1, intermediate between Redfield's ratio for marine plankton and those for vascular plants. Biogeochemical niches were differentiated amongst species and were phylogenetically conserved. Differences in individual and species-specific elementomes increased with coexistence between species. Our results provide an evolutionary bridge between the ecological stoichiometries of algae and vascular plants and suggest that differences in elementomes could be used to understand community assemblages and functional diversity.

Nutrient economy and plant nutrient concentrations have been suggested as important selective pressures of reproductive traits in vascular plants. However, evidence supporting that the reproduction of bryophytes and their reproductive traits have also been conditioned by nutrient availability is lacking. We here analysed the nutrient (N, P, K, and Fe) concentration of 35 aquatic and semi-aquatic bryophytes and the chemistry of the spring water in which they lived to determine whether bryophyte nutrient concentrations were correlated with the frequency of sexual reproduction, main reproductive mode (sexual vs. asexual), dioicy, and spore diameter, while controlling for climate and nutrient concentrations in the spring water. Water nutrient concentrations, except for K, had a small but positive influence on bryophyte nutrient concentrations. P-rich and K-poor species produced sporophytes more often and were more likely to reproduce sexually and be monoicous. Spore diameter was smaller in P- and Fe-rich mosses but larger in N-rich mosses living in humid climates. P-rich species thus produced more and smaller spores and were able to propagate more extensively. Overall, our results suggest that nutrient economy, especially P economy, has played an important role in shaping the reproductive traits of bryophytes.

Background: Rhizodeposition is the release of organic compounds from plant roots into soil. Positive relationships between rhizodeposition and soil microbial biomass are commonly observed. Rhizodeposition may be disrupted by increasing drought however the effects of water stress on this process are not sufficiently understood. Scope: We aimed to provide a synthesis of the current knowledge of drought impacts on rhizodeposition. The current scarcity of well-defined studies hinders a quantitative meta-analysis, but we are able to identify the main effects of water stress on this process and how changes in the severity of drought may produce different responses. We then give an overview of the links between rhizodeposition and microbial communities, and describe how drought may disrupt these interactions. Conclusions: Overall, moderate drought appears to increase rhizodeposition per gram of plant, but under extreme drought rhizodeposition is more variable. Concurrent decreases in plant biomass may lessen the total amount of rhizodeposits entering the soil. Effects on rhizodeposition may be strongly species-dependant therefore impacts on soil communities may also vary, either driving subsequent changes or conferring resilience in the plant community. Advances in the study of rhizodeposition are needed to allow a deeper understanding of this plant-soil interaction and how it will respond to drought.

The abstract is available here: https://uscholar.univie.ac.at/o:1603267

Bryophytes are some of the most sensitive biological indicators of environmental change. Springs have a significant presence of bryophytes and so are ideal habitats for studying their relationship with the environment. We tested whether bryophyte assemblages can be explained with macro-, meso- and micro-ecological variables (i.e. seasonal climate, altitude, water pH and conductivity) sampling bryophytes from 198 semi-natural springs distributed along montane regions in the north-eastern Iberian Peninsula. We tested the influence of environmental variables on bryophyte assemblages in springs using sparse Partial Least Squares. Our results show that variability in bryophyte assemblages is explained by seasonal climate (temperature and precipitation from winter, spring, summer and autumn and temperature and precipitation seasonality), altitude and water conductivity. The results obtained by the present study will be useful for predicting bryophyte diversity in springs using simple and easy to obtain variables such as climate, water pH and conductivity.

Root exudation is an important input of carbon into soils and affects plant and soil communities, but little is known about the effect of climatic factors such as drought on exudation, and its ability to recover. We studied the impact of increasing drought on root exudation and its subsequent recovery in the Mediterranean tree species Quercus ilex L. in a greenhouse study by measuring the amount of total organic carbon in exudates. The amount of exudation per unit root area increased with drought duration and was 21% higher under the most extreme drought scenario compared with the non-droughted control. The amount of root exudation did not differ between the treatments following 6 weeks of re-watering, indicating a strong capacity for recovery in this species. We concluded that drought could affect the amount of root exudation, which could in turn have a large impact on microbial activity in the rhizosphere, and alter these microbial communities, at least in the short term. This tree species may be able to return to normal levels of root exudation after a drought event, but long-term exudate-mediated impacts on Mediterranean forest soils may be an unforeseen effect of drought.

Anthropogenic activities and intensive farming arc causing nitrate pollution in groundwater bodies. These aquifers are drained by springs which, in the Mediterranean region, act as refugia for preserving biodiversity of species that need continuous water. Some springs are also used for drinking water for wild animals, livestock and humans, so if their water quality is compromised it can become a threat to public health. However, the impact of nitrate pollution on these biotic communities remains unknown. We sampled 338 assemblages of aquatic and semi-aquatic bryophytes (i.e., hygrophytic mosses and liverworts) growing in springs in a gradient of water conductivity, nitrate concentration and climate and distributed across the north-east of the Iberian Peninsula to investigate the impact of nitrate pollution on the diversity of bryophytes and moss functional traits in Mediterranean springs. Based on previous literature suggesting that increased nitrogen load decreases biodiversity in grasslands and freshwater ecosystems, we hypothesised that water nitrate pollution in springs decreases bryophyte diversity at the local and regional scales. Our results indicated that, at the local scale (spring), nitrate pollution reduced the number and the likelihood of finding a rare species in springs. Rare species were found in 4% of the springs with nitrate above 50 mg but in 32% of the springs with nitrate below 50 mg L-1. Moss, liv- erwort and overall bryophyte diversity were not directly affected by nitrate at the local scale but nitrate consistently decreased diversity of mosses, liverworts and rare bryophyte species at the regional scale. We also found that warmer and drier springs presented fewer bryophyte species. Our results show that the combination of nitrate pollution, increasing temperature and drought could severely threaten bryophyte diversity Mediterranean springs. Our results indicate that the absence of rare bryophytes could be used as a bioindicator of nitrate pollution in springs. (C) 2018 Elsevier B.V. All rights reserved.

The drivers of global change, such as increasing drought and nutrient deposition, are affecting soils and their microbial communities in many different habitats, but how these factors interact remains unclear. Quercus ilex and Pinus sylvestris are two important tree species in Mediterranean montane areas that respond differently to drought, which may be associated with the soils in which they grow. We measured soil respiration and physiologically profiled microbial communities to test the impact of drought and subsequent recovery on soil function and diversity for these two species. We also tested whether the addition of nitrogen and phosphorus modified these effects. Drought was the stronger driver of changes to the soil communities, decreasing diversity (Shannon index), and evenness for both species and decreasing soil respiration for Q. ilex when N was added. Soil respiration for P. sylvestris during the drought period was positively affected by N addition but was not affected by water stress. P addition during the drought period did not affect soil respiration for either tree species but did interact with soil-water content to affect community evenness for P. sylvestris. The two species also differed following the recovery from drought. Soil respiration for Q. ilex recovered fully after the drought treatment ended but decreased for P. sylvestris, whereas the soil community was more resilient for P. sylvestris than Q. ilex. Nutrient addition did not affect respiration or community composition or diversity during the recovery period. Soil respiration was generally weakly positively correlated with soil diversity. We demonstrate that short-term water stress and nutrient addition can have variable effects on the soil communities associated with different tree species and that the compositions of the communities can become uncoupled from soil respiration. Overall, we show that drought may be a stronger driver of changes to soil communities than nitrogen or phosphorus deposition.

1. Climate change is a world-wide threat to biodiversity and ecosystem structure, functioning and services. To understand the underlying drivers and mechanisms, and to predict the consequences for nature and people, we urgently need better understanding of the direction and magnitude of climate change impacts across the soil–plant–atmosphere continuum. An increasing number of climate change studies are creating new opportunities for meaningful and high-quality generalizations and improved process understanding. However, significant challenges exist related to data availability and/or compatibility across studies, compromising opportunities for data re-use, synthesis and upscaling. Many of these challenges relate to a lack of an established ‘best practice’ for measuring key impacts and responses. This restrains our current understanding of complex processes and mechanisms in terrestrial ecosystems related to climate change. 2. To overcome these challenges, we collected best-practice methods emerging from major ecological research networks and experiments, as synthesized by 115 experts from across a wide range of scientific disciplines. Our handbook contains guidance on the selection of response variables for different purposes, protocols for standardized measurements of 66 such response variables and advice on data management. Specifically, we recommend a minimum subset of variables that should be collected in all climate change studies to allow data re-use and synthesis, and give guidance on additional variables critical for different types of synthesis and upscaling. The goal of this community effort is to facilitate awareness of the importance and broader application of standardized methods to promote data re-use, availability, compatibility and transparency. We envision improved research practices that will increase returns on investments in individual research projects, facilitate second-order research outputs and create opportunities for collaboration across scientific communities. Ultimately, this should significantly improve the quality and impact of the science, which is required to fulfil society's best practice, coordinated experiments, data management and documentation, ecosystem, experimental macroecology, methodology, open science, vegetation This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. © 2019 The Authors. Methods in Ecology and Evolution published by John Wiley & Sons Ltd on behalf of British Ecological Society.

Mosses are amongst the oldest and simplest plants, they can be found almost everywhere in the world, and they condition the structure and function of many ecosystems. Their sensitivity to environmental changes makes them very interesting subjects of study in ecology, and understanding them can provide insights into the evolutionary history of plants. However, the study of moss traits and their relationship with their environment is far behind that of vascular plants. We sampled 303 assemblages of aquatic and semi‐aquatic (hygrophytic) mosses growing in semi‐natural springs distributed around the northeast of the Iberian Peninsula to study how moss traits vary depending on their evolutionary history, climate and water chemistry. To do so, we analysed 30 moss species and 17 traits using phylogenetic comparative methods and an extended RLQ analysis, accounting for spatial and phylogenetic information. We hypothesized that there is a sclerophylly continuum in mosses living across a gradient of high and low water conductivity springs that may mimic sclerophylly in vascular plants that live in stressful environments. Results indicated that life‐forms and, especially, morphological traits were well preserved phylogenetically and responsive to water chemistry and climate. That combined with spatial autocorrelation in environmental variables resulted in a clustered distribution of phylogenetically closely related mosses in space. Mosses living in springs with a warm and dry climate that discharge hard water mainly presented species with needle‐like leaves, were denser, and had lower water absorption capacity. The opposite was found in cold, humid and soft water springs. Synthesis. Our results suggest that climate and water chemistry are main determinants of traits of hygrophytic mosses and of species distributions. We found evidence of a potential sclerophylly continuum in moss traits, which we hypothesize may be mainly related to physical and physiological constraints produced by water chemistry. Our findings describe moss sclerophylly in a gradient of water conductivity similar to that found in vascular plants with water availability and temperature. Further experimental studies will be required to confirm the observations found in this study. A free Plain Language Summary can be found within the Supporting Information of this article.

We synthesise the results of a large programme of plant ecological research to investigate the selective pressures driving crop domestication and the origins of agriculture in western Asia. We explore this primarily through a series of experiments, comparing the ecological characteristics of: (1) domesticated cereal and pulse species with their wild progenitors and (2) the wild progenitor species with other west Asian grasses and legumes that did not become domesticated during the emergence of agriculture. In particular, we consider the balance between deliberate human selection and unintended consequences of human actions in driving the domestication process. Taken together, our results provide the first empirical evidence to suggest that ecological processes, and unintended selection due to competition between growing plants within anthropogenic environments, may have played a more significant part in the emergence of agriculture than previously supposed. Such human-plant co-evolutionary mechanisms would render unnecessary the search for ‘push’ or ‘pull’ factors, dependent on deliberate human invention to solve a problem or to satisfy a need, as prime movers to explain why hunter-gatherers switched to an agricultural way of life.

Agraïments: Chao Zhang gratefully acknowledges the support from the Chinese Scholarship Council. The photochemical reflectance index (PRI) and red-edge region of the spectrum are known to be sensitive to plant physiological processes, and through measurement of these optical signals it is possible to use non-invasive remote sensing to monitor the plant photosynthetic status in response to environmental stresses such as drought. We conducted a greenhouse experiment using Quercus ilex, a Mediterranean evergreen oak species, to investigate the links between leaf-level PRI and the red-edge based reflectance ratio (R690/R630) with CO2 assimilation rates (A), and photochemical efficiency (FV/FM and Yield) in response to a gradient of mild to extreme drought treatments (nine progressively enhanced drought levels) and corresponding recovery. PRI and R690/R630 both decreased under enhanced drought stress, and had significant correlations with A, FV/FM and Yield. The differential values between recovery and drought treatments of PRI (DPRIrecovery) and R690/R630 (DR690/R630recovery) increased with the enhanced drought levels, and significantly correlated with the increases of DArecovery, DFV/FMrecovery and DYieldrecovery. We concluded that both PRI and R690/R630 were not only sensitive to enhanced drought stresses, but also highly sensitive to photosynthetic recovery. Our study makes important progress for remotely monitoring the effect of drought and recovery on photosynthetic regulation using the simple physiological indices of PRI and R690/R630.

Polar regions are predicted to undergo large increases in winter temperature and an increased frequency of freeze–thaw cycles, which can cause ice layers in the snow pack and ice encasement of vegetation. Early or late winter timing of ice encasement could, however, modify the extent of damage caused to plants. To determine impacts of the date of ice encasement, a novel field experiment was established in sub‐arctic Sweden, with icing events simulated in January and March 2008 and 2009. In the subsequent summers, reproduction, phenology, growth and mortality, as well as physiological indicators of leaf damage were measured in the three dominant dwarf shrubs: Vaccinium uliginosum, Vaccinium vitis‐idaea and Empetrum nigrum. It was hypothesised that January icing would be more damaging compared to March icing due to the longer duration of ice encasement. Following 2 years of icing, E. nigrum berry production was 83% lower in January‐iced plots compared to controls, and V. vitis‐idaea electrolyte leakage was increased by 69%. Conversely, electrolyte leakage of E. nigrum was 25% lower and leaf emergence of V. vitis‐idaea commenced 11 days earlier in March‐iced plots compared to control plots in 2009. There was no effect of icing on any of the other parameters measured, indicating that overall these study species have moderate to high tolerance to ice encasement. Even much longer exposure under the January icing treatment does not clearly increase damage.

During the origin of agriculture in the Fertile Crescent, the broad spectrum of wild plant species exploited by hunter‐gatherers narrowed dramatically. The mechanisms responsible for this specialization and the associated domestication of plants are intensely debated. We investigated why some species were domesticated rather than others, and which traits they shared. We tested whether the progenitors of cereal and pulse crops, grown individually, produced a higher yield and less chaff than other wild grasses and legumes, thereby maximizing the return per seed planted and minimizing processing time. We compared harvest traits of species originating from the Fertile Crescent, including those for which there is archaeological evidence of deliberate collection. Unexpectedly, wild crop progenitors in both families had neither higher grain yield nor, in grasses, less chaff, although they did have larger seeds. Moreover, small‐seeded grasses actually returned a higher yield relative to the mass of seeds sown. However, cereal progenitors had threefold fewer seeds per plant, representing a major difference in how seeds are packaged on plants. These data suggest that there was no intrinsic yield advantage to adopting large‐seeded progenitor species as crops. Explaining why Neolithic agriculture was founded on these species, therefore, remains an important unresolved challenge.

Arctic plants can experience prolonged periods of ice encasement in winter, leading to hypoxia which may cause damage to plants and reduce subsequent summer growth. Further, high CO 2 concentrations, which can occur when plants respire within ice, may cause additional injury, as has been shown to occur in crops. The tolerance of arctic plants to these within-ice atmospheric conditions has previously remained unknown, yet this knowledge is essential for understanding icing impacts on plant community structure and productivity – especially given that icing events are predicted to increase in frequency as a result of climate change. Using a unique field chamber experiment, this study quantified the responses of three widespread sub-arctic dwarf shrubs, Empetrum nigrum and Vaccinium vitis-idaea (both evergreen) and V. myrtillus (deciduous), to 14 days hypoxia and high CO 2 in winter in a factorial field experiment in northern Sweden. Growth, phenology and mortality responses were used to quantify impacts in the following spring supported by electrolyte leakage and chlorophyll fluorescence to assess leaf damage in the two evergreens. Overall, all species showed high resistance to 14 days of hypoxia with no indications of damage. Increased CO 2 did lead to 33% lower bud dormancy in E. nigrum and 70% greater shoot mortality in V. vitis-idaea indicating that these species might be negatively affected by increased occurrences of ice encasement, through elevated CO 2 levels. Despite these responses, effects of high CO 2 were rare overall. Given that the impacts of hypoxia and high CO 2 were considerably less than reported for species in temperate habitats, this suggests a moderate to high degree of tolerance to short periods of icing among these species. If these results apply to longer periods of ice encasement, increasing frequency of icing may only have some species specific impacts without being a major environmental threat to arctic dwarf shrubs.