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In times of unprecedented climate change, ecological restoration efforts have a strong focus on forests for the purpose of carbon sequestration. Grasslands, in contrast, remain relatively neglected in global restoration policies. Concurrently, we are in the midst of a biodiversity crisis—it is estimated that 1 million species are globally threatened with extinction. Here, we present analyses from central Europe and southern Brazil that show that the majority of our endangered plant species are in fact found in open ecosystems. Using Germany as an example, we show that we could reduce plant extinction risk by up to 82% if we restore open, grassy ecosystems. This also holds true for southern Brazil, where grassland species constitute the single largest share of endangered species, but where grassy ecosystems continue to be systematically neglected by restoration policies. We further expand on our biodiversity argument to include the role that grassland restoration can play in mitigating climate change. We posit that ramping up grassland restoration efforts may not only be our best bet to bend the curve of biodiversity loss, but it will also make a critical contribution to the resilience of ecosystems in the dynamic decades to come. It is time for grassland restoration to receive higher priority in global restoration efforts and policy.

Less attention has been given to soil enzymes that contribute to beneficial rhizosphere interactions in intercropping systems. Therefore, we performed a field experiment by growing faba bean, lupine, and maize in mono and mixed cultures in a moderately fertile soil. We measured shoot biomass and the kinetic parameters (maximal velocity (Vmax) and Michaelis-constant (Km)) of three key enzymes in the rhizosphere: Leucine-aminopeptidase (LAP), β-1,4-N-acetylglucosaminidase (NAG), and phosphomonoesterase (PHO). Faba bean benefitted in mixed cultures by greater shoot biomass production with both maize and lupine compared to its expected biomass in monoculture. Next, LAP and NAG kinetic parameters were less responsive to mono and mixed cultures across the crop species. In contrast, both the Vmax and Km values of PHO increased in the faba bean rhizosphere when grown in mixed cultures with maize and lupine. A positive relative interaction index for shoot P and N uptake for faba bean showed its net facilitative interactions in the mixed cultures. Overall, these results suggest that over-productivity in intercropping is crop-specific and the positive intercropping effects could be modulated by P availability. We argue that the enzyme activities involved in nutrient cycling should be incorporated in further research.

Aims: Although different plant foraging responses to the two macronutrients nitrogen (N) and phosphorus (P) are well researched, the effect of timing of fertilizer application on root system architecture (RSA) remains largely unknown. We, therefore, aimed to understand how RSA of Hordeum vulgare L. responds to timing of N and P application. Methods: Plants were grown in rhizoboxes for 38 days in nutrient-poor soil and watered with nutrient solution, lacking either N or P, with the absent nutrient applied once either 2/3/4 weeks after sowing. Positive controls were continuously receiving N and P and a negative control receiving both N and P only after 3 weeks. We tracked root growth over time, measured plant biomass and nutrient uptake. Results: Late N application strongly reduced total root biomass and visible root length compared with continuous NP and late P application. Root mass fractions (total root biomass/total plant biomass) remained similar over all treatments, but relative allocation (% of total root biomass) was higher in lower depth with late N application. Shoot P concentrations remained relatively stable, but the plants receiving P later had higher N concentrations. Conclusions: Late N application had overall more negative effects on early plant growth compared with late P. We propose that future studies under field conditions should try to disentangle the effect of timing from the nutrient availability on RSA responses and hence ultimately plant performance.

In Brazil, the country with the highest plant species richness in the world, biodiverse savannas and grasslands – i.e., grassy ecosystems, which occupy 27% of the country – have historically been neglected in conservation and scientific treatments. Reasons for this neglect include misconceptions about the characteristics and dynamics of these ecosystems, as well as inconsistent or regionally restricted terminology that impeded a more adequate communication about Brazil's savannas and grasslands, both within the country and internationally. Toward improved communication and recognition of Brazil's diversity of ecosystems, we present the key drivers that control the main types of grassy ecosystems across Brazil (including in regions of the country where forests dominate). In doing so, we synthesize the main features of each grassy ecosystem in terms of physiognomy and ecological dynamics (e.g., relationships with herbivores and fire). We propose a terminology both for major grassland regions and for regionally relevant vegetation physiognomies. We also discuss terms associated with human land management and restoration of grassy ecosystems. Finally, we suggest key research needs to advance our understanding of the ecology and conservation values of Brazil's grassy ecosystems. We expect that a common and shared terminology and understanding, as proposed here, will stimulate more integrative research that will be fundamental to developing improved conservation and restoration strategies.

Soil legacies play an important role for the creation of priority effects. However, we still poorly understand to what extent the metabolome found in the soil solution of a plant community is conditioned by its species composition and whether soil chemical legacies affect subsequent species during assembly. To test these hypotheses, we collected soil solutions from forb or grass communities and evaluated how the metabolome of these soil solutions affected the growth, biomass allocation, and functional traits of a forb (Dianthus deltoides) and a grass species (Festuca rubra).Results showed that the metabolomes found in the soil solutions of forb and grass communities differed in composition and chemical diversity. While soil chemical legacies did not have any effect onF. rubra,root foraging byD. deltoidesdecreased when plants received the soil solution from a grass or a forb community. Structural equation modelling showed that reduced soil exploration byD. deltoidesarose via either a root growth-dependent pathway (forb metabolome) or a root trait-dependent pathway (grass metabolome). Reduced root foraging was not connected to a decrease in total N uptake. Our findings reveal that soil chemical legacies can create belowground priority effects by affecting root foraging in later arriving plants.

The order of arrival of plant species during assembly can affect the structure and functioning of grassland communities. These so-called priority effects have been extensively studied aboveground, but we still do not know how they affect the vertical distribution of roots in the soil and the rooting depth of plant communities. To test this hypothesis, we manipulated the order of arrival of three plant functional groups (forbs, grasses and legumes) in a rhizobox experiment. Priority effects were created by sowing one functional group 10 days before the other two. Rhizoboxes in which all functional groups were sown simultaneously were used as controls. During the experiment, the mean rooting depth of plant communities was monitored using image analysis and a new methodological approach using deep learning (RootPainter) for root segmentation. At harvest, we measured aboveground (community and species level) and belowground (community level) biomass, and assessed the vertical distribution of the root biomass in different soil layers. At the community level, all scenarios where one functional group was sown before the other two had similar shoot and root productivity. At the species level, two forbs (Achillea millefolium and Centaurea jacea) benefited from arriving early, and one legume (Trifolium pratense) had a disadvantage when it was sown after the grasses. Priority effect treatments also affected the vertical distribution of roots. When grasses were sown first, plant communities rooted more shallowly than when forbs or legumes were sown first,. In addition, roots moved down the soil profile 24% more slowly in grasses-first communities. Our results highlight that plant functional group order of arrival in grassland communities can affect the vertical distribution of roots in the soil and this may have implications for species coexistence.

Restoration ecology commonly seeks to re-establish species of interest in degraded habitats. Despite a rich understanding of how succession influences re-establishment, there are several outstanding questions that remain unaddressed: are short-term abundances sufficient to determine long-term re-establishment success, and what factors contribute to unpredictable restorations outcomes? In other words, when restoration fails, is it because the restored habitat is substandard, because of strong competition with invasive species, or alternatively due to changing environmental conditions that would equally impact established populations? Here, we re-purpose tools developed from modern coexistence theory to address these questions, and apply them to an effort to restore the endangered Contra Costa goldfields (Lasthenia conjugens) in constructed ("restored") California vernal pools. Using 16 years of data, we construct a population model of L. conjugens, a species of conservation concern due primarily to habitat loss and invasion of exotic grasses. We show that initial, short-term appearances of restoration success from population abundances is misleading, as year-to-year fluctuations cause long-term population growth rates to fall below zero. The failure of constructed pools is driven by lower maximum growth rates compared with reference ("natural") pools, coupled with a stronger negative sensitivity to annual fluctuations in abiotic conditions that yield decreased maximum growth rates. Nonetheless, our modeling shows that fluctuations in competition (mainly with exotic grasses) benefit L. conjugens through periods of competitive release, especially in constructed pools of intermediate pool depth. We therefore show how reductions in invasives and seed addition in pools of particular depths could change the outcome of restoration for L. conjugens. By applying a largely theoretical framework to the urgent goal of ecological restoration, our study provides a blueprint for predicting restoration success, and identifies future actions to reverse species loss. Funding Agencies|Air Force Center for Environmental Excellence; US Fish and Wildlife Service; CH2M Hill; NSF LTREB grants [DEB-0744520, DEB-1257385]; USDA NIFA Predoctoral Fellowship [2021-67034-35111]; Swedish Research Council (Vetenskapsradet) [2017-05245]; Spanish Ministry of Economy and Competitiveness (MINECO); European Social Fund through the Ramon y Cajal Program [RYC2017-23666]

Plant residues with larger carbon (C) to nitrogen (N) ratios can stimulate microbial growth and thereby protect soil nutrients from leaching. In poorly fertilized soil, excessive immobilization may limit nutrient availability and thus plant growth. Little is known about the impact of a shallow straw incorporation on soil microbial regulation of top-dressing fertilizer nutrients and spring crop establishment. We aimed to evaluate if wheat straw in combination with mineral fertilizer has more positive effects on plant performance than mineral fertilization alone and if this relates to changes of the extractable C:N:P ratio and microbial activity close to the roots. In order to conduct small-scale sampling with minimal disturbance during growth of spring barley (Hordeum vulgare L.), we developed rhizotrons with resealable ports. Rhizotrons were filled with loamy-sandy soil and fertilized with an equivalent of 150 kg N and 80 kg P ha−1. Half of the rhizotrons received the top dressing together with 4500 kg wheat straw-C ha−1. Throughout a 90-day greenhouse experiment, we analyzed soil C:N:P dynamics, and carbon dioxide (CO2) and nitrous oxide (N2O) emission, together with microbial biomass, selected bacterial genes (abundance), and transcripts (activity) in bulk and root-affected soil at multiple times. We focused on nitrifiers and denitrifiers and linked our data to barley growth. Interactions between straw and roots caused shifts towards larger C:P and C:N ratios in root-affected soil. These shifts were associated with increased 16S rRNA transcripts and denitrifier activities. Straw increased microbial biomass by 124% in the topsoil and at the same time increased root biomass by 125% and number of tillers by 80%. We concluded that microbial activation at the root-straw interface may positively feed back on soil nutrient regulation and plant performance. Further research has to evaluate if plant roots actively prime mining of previously immobilized nutrients in the straw detritusphere or if effects of pathogen suppression and growth promotion are dominating.

Aim Intercropping often leads to improved productivity of individual species compared to monocultures. We have practically little knowledge of facilitation effects in different intercropping systems and their importance in creating soil legacies that can indirectly affect the succeeding crop in a crop rotation through plant-soil feedback (PSF) effects. Methods To test this, we used a two-phased field experiment where we combined intercropping and crop rotation. During intercropping, we grew maize, faba bean, and lupine in monocultures or two-species crop combinations. The following season, we grew winter barley on the soil previously used for intercropping to test PSF effects under field conditions. Results We found evidence for facilitative effects on aboveground biomass production that were species-specific with faba bean and maize biomass benefitting when intercropped compared to their expected biomasses in monocultures. Lupine, in contrast, performed best in monocultures. After the intercropping phase, total soil mineral nitrogen was higher in legume monocultures creating soil legacies but this did not affect soil microbial parameters and barley biomass production in the follow-up rotation phase. Conclusions We found support for species-specific positive and negative interactions in intercropping. Our results also demonstrated that soil legacies play no significant role under moderately high nutrient environments.

The exotic South African ragwort (Senecio inaequidens DC.) rapidly spread across Central Europe after its introduction, but we still do not know to what extent its timing of arrival in a plant community (i.e. before or after natives) and the composition of the native community being invaded affect (1) its capacity to invade a European grassland, (2) the performance of the native species, and (3) the direction and strength of priority effects. In a greenhouse experiment, we manipulated the timing of arrival of the exotic species (Senecio) and the composition of the native community to test the influence of these factors on the productivity and N content of exotic and native species. We also investigated if the plant species origin (native or exotic) and the native community composition affected the benefit of arriving early and the cost of arriving late in the community. The establishment success of Senecio strongly depended on its timing of arrival in a grassland community. Senecio benefited more from arriving early than did the natives. The presence of legumes in the community did not favour invasion by Senecio. When natives arrived later than Senecio, however, priority effects were weaker when legumes were part of the native community. Our results showed that inhibitory priority effects created by natives can lower the risk of invasion by Senecio. An early arrival of this species at a site with low native species abundance is a scenario that could favour invasion.

Biodiversity-ecosystem functioning experiments have shown that plant species and functional group richness are important drivers of grassland productivity, but the impact that plant order of arrival (i.e. priority effects) has on grassland overyielding and its drivers (complementarity and dominance effects) has been overlooked so far. Using species-specific plant biomass data collected in mixture and monoculture plots of a grassland field experiment (Julich Priority Effect experiment) that manipulated the order of arrival of three plant functional groups (forbs, grasses and legumes), we quantified net biodiversity effects (overyielding) as well as complementarity and dominance effects in mixtures one and 2 years after sowing. In this experiment, priority effects were created by sowing one functional group 6 weeks before the two others. First, we tested whether plant order of arrival affected overyielding, complementarity and dominance effects. Second, we investigated whether the magnitude of net biodiversity, complementarity and dominance effects was dependent on the strength and direction of priority effects. We found that complementarity and dominance effects were affected by plant order of arrival during community assembly. In addition, we found that moving from negative to positive priority effects increased grassland overyielding, mainly via increased complementarity effects. These results highlight the need to combine biodiversity and assembly approaches in future ecosystem functioning research, as this will increase the predictive power of community ecology in conservation and ecological restoration. A free Plain Language Summary can be found within the Supporting Information of this article.

The United Nations (UN) recently declared 2021 to 2030 the Decade on Ecosystem Restoration. Against this background, we review recent social-ecological systems research and summarize key themes that could help to improve ecosystem restoration in dynamic social contexts. The themes relate to resilience and adaptability, ecosystem stewardship and navigation of change, relational values, the coevolution of human and ecological systems, long-range social-ecological connections, and leverage points for transformation. We recommend two cross-cutting new research foci; namely: (i) post hoc cross-sectional assessments of social-ecological restoration projects; and (ii) transdisciplinary social-ecological ‘living labs’ that accompany new restoration projects as they unfold. With global agendas increasingly taking a social-ecological perspective, the recasting of ecosystem restoration as a social-ecological endeavor offers exciting new opportunities for both research and practice.

A 2-year outdoor mesocosm experiment was carried out to determine the effects of high C amendments (HCAs; wheat straw and sawdust) compared to a control with no addition of HCAs (no-HCA) and 2 different crop rotation systems (spring barley/winter barley and faba bean/winter barley) on soil bacterial communities using a molecular barcoding approach. Samples were analyzed after pre-crop harvest (T1) and harvest of winter barley (T2). Our data demonstrate a clear drop in bacterial diversity after winter barley harvest in the no-HCA and wheat straw treatment compared to the pre-crops. Sawdust application had a stabilizing effect on bacterial diversity compared to the pre-crops and induced an increase in carbon (C) stocks in soil which were however negatively correlated with yields. Main responders in the no-HCA and wheat straw treatment compared to the pre-crops were bacteria of the phyla Actinobacteria and Bacteroidetes which were enriched and bacteria belonging to Firmicutes, Gemmatimonadetes, Proteobacteria, and Gemmatimonadaceae which were depleted. Overall differences between wheat straw–amended and no-HCA control samples were small and included single ASVs from various phyla. In sawdust-amended samples, only a shift of some Proteobacteria families was observed compared to the no-HCA control. Overall, pre-crop plant species had small influence on the observed response pattern of the soil microbiome towards the amendments and was only visible for wheat straw.

The prevailing nature-based solution to tackle climate change is tree planting. However, there is growing evidence that it has serious contraindications in many regions. The main shortcoming of global tree planting is its awareness disparity to alternative ecosystem types, mainly grasslands. Grasslands, where they constitute the natural vegetation, may support higher biodiversity and a safer, soil-locked carbon stock than plantations and other forests. We suggest replacing “tree planting” by “restore native vegetation.” This improved action terminology reduces the risks of inappropriate afforestation and, by diversifying target ecosystem types, does not reduce but increases potential land area for nature-based climate mitigation.

Earth is home to over 350,000 vascular plant species that differ in their traits in innumerable ways. A key challenge is to predict how natural or anthropogenically driven changes in the identity, abundance and diversity of co-occurring plant species drive important ecosystem-level properties such as biomass production or carbon storage. Here, we analyse the extent to which 42 different ecosystem properties can be predicted by 41 plant traits in 78 experimentally manipulated grassland plots over 10 years. Despite the unprecedented number of traits analysed, the average percentage of variation in ecosystem properties jointly explained was only moderate (32.6%) within individual years, and even much lower (12.7%) across years. Most other studies linking ecosystem properties to plant traits analysed no more than six traits and, when including only six traits in our analysis, the average percentage of variation explained in across-year levels of ecosystem properties dropped to 4.8%. Furthermore, we found on average only 12.2% overlap in significant predictors among ecosystem properties, indicating that a small set of key traits able to explain multiple ecosystem properties does not exist. Our results therefore suggest that there are specific limits to the extent to which traits per se can predict the long-term functional consequences of biodiversity change, so that data on additional drivers, such as interacting abiotic factors, may be required to improve predictions of ecosystem property levels.

Priority effects refer to the order or timing of species arrival, including how species that arrive early at a site either positively or negatively affect establishment, growth, or reproduction of species that arrive later. Despite the clear implications of priority effects for ecological restoration, there have been no reviews of how and where priority effects have been studied and the extent to which findings can be applied to restoration. Here, we systematically review the literature on priority effects by (1) synthesizing information from papers that compared simultaneous and nonsimultaneous planting or sowing; (2) discussing the mechanisms through which priority effects operate, (3) considering how these mechanisms might be manipulated to achieve restoration goals; and (4) highlighting future research needed to improve the use of priority effects in restoration. In a term‐targeted search, we found 43 studies that experimentally manipulated the order of arrival of different species. Overall, these concluded that even small delays in arrival time, as opposed to simultaneous arrival of species, can promote differences in subsequent community composition as well as ecosystem functions. There were very few studies on the long‐term stability of these priority effects, and the majority were conducted in temperate grasslands. Our findings suggest that creating alternative vegetation states via priority treatments is a promising avenue for restoration. However, for the concept to be best operationalized for restoration, we need research in more ecosystems that are priorities for restoration, and treatments that are followed over extended time periods. Priority effects refer to the order or timing of species arrival, including how species that arrive early at a site either positively or negatively affect establishment, growth, or reproduction of species that arrive later. Despite the clear implications of priority effects for ecological restoration, there have been no reviews of how and where priority effects have been studied and the extentto which findings can be applied to restoration. Here, we systematically review the literature on priority effects by (1) synthesizing information from papers that compared simultaneous and nonsimultaneous planting or sowing; (2) discussing the mechanisms through which priority effects operate, (3) considering how these mechanisms might be manipulated to achieve restoration goals; and (4) highlighting future research needed to improve the use of priority effects in restoration. In a termtargeted search, we found 43 studies that experimentally manipulated the order of arrival of different species. Overall, these concluded that even small delays in arrival time, as opposed to simultaneous arrival of species, can promote differences in subsequent community composition as well as ecosystem functions. There were very few studies on the long-term stability of these priority effects, and the majority were conducted in temperate grasslands. Our findings suggest that creating alternative vegetation states via priority treatments is a promising avenue for restoration. However, for the concept to be best operationalized for restoration, we need research in more ecosystems that are priorities for restoration, and treatments that are followed over extended time periods.

Aims: Root system responses to the limitation of either nitrogen (N) or phosphorus (P) are well documented, but how the early root system responds to (co-) limitation of one (N or P) or both in a stoichiometric framework is not well-known. In addition, how intraspecific competition alters plant responses to N:P stoichiometry is understudied. Therefore, we aimed to investigate the effects of N:P stoichiometry and competition on root system responses and overall plant performance. Methods: Plants (Hordeum vulgare L.) were grown in rhizoboxes for 24 days in the presence or absence of competition (three vs. one plant per rhizobox), and fertilized with different combinations of N:P (low N + low P, low N + high P, high N + low P, and high N + high P). Results: Shoot biomass was highest when both N and P were provided in high amounts. In competition, shoot biomass decreased on average by 22%. Total root biomass (per plant) was not affected by N:P stoichiometry and competition but differences were observed in specific root length and root biomass allocation across soil depths. Specific root length depended on the identity of limiting nutrient (N or P) and competition. Plants had higher proportion of root biomass in deeper soil layers under N limitation, while a greater proportion of root biomass was found at the top soil layers under P limitation. Conclusions: With low N and P availability during early growth, higher investments in root system development can significantly trade off with aboveground productivity, and strong intraspecific competition can further strengthen such effects. Aims Root system responses to the limitation of either nitrogen (N) or phosphorus (P) are well documented, but how the early root system responds to (co-) limitation of one (N or P) or both in a stoichiometric framework is not well-known. In addition, how intraspecific competition alters plant responses to N:P stoichiometry is understudied. Therefore, we aimed to investigate the effects of N:P stoichiometry and competition on root system responses and overall plant performance.Methods Plants (Hordeum vulgare L.) were grown in rhizoboxes for 24 days in the presence or absence of competition (three vs. one plant per rhizobox), andfertilized with different combinations of N:P (low N + low P, low N + high P, high N + low P, and high N + high P).Results Shoot biomass was highest when both N and P were provided in high amounts. In competition, shoot biomass decreased on average by 22%. Total root biomass(per plant) was not affected by N:P stoichiometry and competition but differences were observed in specific root length and root biomass allocation across soil depths. Specific root length depended on the identity of limiting nutrient (N or P) and competition. Plants had higher proportion of root biomass in deeper soil layers under N limitation, while a greater proportion of rootbiomass was found at the top soil layers under P limitation.Conclusions With low N and P availability during early growth, higher investments in root system development can significantly trade off with aboveground productivity,and strong intraspecific competition can further strengthen such effects.

Interactions between plants can be both positive and negative, denoting facilitation and competition. Although facilitative effects of having legume neighbours (focus on yield productivity) are well studied, a better mechanistic understanding of how legumes interact with non-legumes in terms of root distribution is needed. We tested the effects of neighbour identity, its spatial location, as well as the effects of plant order of arrival on above and belowground traits and root distribution. We performed a rhizotron experiment (4 weeks duration) in which we grew maize alone, with only a legume, only another grass, or with both species and tracked roots of the plant species using green and red fluorescent markers. Maize grew differently when it had a neighbour, with reduced development when growing with wheat compared to alone. Growing with a legume generally equated to the same outcome as not having a neighbour. Roots grew towards the legume species and away from the wheat. Order of arrival affected aboveground traits to a certain extent, but its effects on maize roots were dependent on spatial location. Our study provides evidence of facilitation, showing the importance of the identity of the neighbours, together with their spatial location, and how order of arrival can modulate the outcome of these initial interactions.

Bastin et al.’s estimate (Reports, 5 July 2019, p. 76) that tree planting for climate change mitigation could sequester 205 gigatonnes of carbon is approximately five times too large. Their analysis inflated soil organic carbon gains, failed to safeguard against warming from trees at high latitudes and elevations, and considered afforestation of savannas, grasslands, and shrublands to be restoration., Funding: Supported by the Texas A&M Sid Kyle Global Savanna Research Initiative (T.W.B.); Swiss National Science Foundation (20FI20_173691) (N.B.); Centre National pour la Recherche Scientifique CNRS PICS 2018-2020 (RESIGRASS) (E.B.); CNPq (Brazil, 303179/2016-3) (G.D.); CNPq (Brazil) (G.W.F.); CNPq (Brazil, 303988/2018-5) (A.F.); NASA award NNX17AK14G (F.F.); NSF award 1354943 (W.A.H.); Fundação de Amparo à Pesquisa do Estado de Minas Gerais (Brazil, 2016/13232-5) (S.L.S.); the Office of the Royal Society (IC170015) (C.E.R.L.); CNPq (Brazil, 310345/2018-9) (G.E.O.); the Spanish Government (FIROTIC, PGC2018-096569-B-I00) (J.G.P.); the National Research Foundation (ACCESS, 114695) (N.S.); CNPq (Brazil, 303568/2017-8) (F.A.O.S.); NSF awards 1342703 and 1926431 (C.J.S. and D.M.G.); NSF award EAR-1253713 (C.A.E.S.); Deutsche Forschungsgemeinschaft grant 5579 POEM (V.M.T.); and USDA-NIFA Sustainable Agricultural Systems Grant 12726253 (J.W.V.)

During biogas production anaerobic digestion of plant material produces a nutrient-rich residue called digestate. The application of the nutrients present in the digestate should improve soil fertility, particularly in nutrient poor soils, and thus crop yield, promoting the closure of the nutrient cycle. This study evaluated the effect of digestate application on the germination and early stages of plant development since these are the first steps to be considered when studying the benefits on plant growth in low fertility substrates. A greenhouse experiment was conducted to evaluate the effects of three substrates of different texture and fertility (field loam, field sand, sand), as well as type and amount of fertilizer (pure maize digestate vs. inorganic nitrogen/phosphorus/potassium (NPK) fertilizer) on both germination and early plant performance of maize (Zea mays L. subsp. mays). While digestate and NPK fertilizer applications had no significant effect on germination in the two field soils, digestate applications significantly decreased the germination rate in sand (36&ndash, 82% reduction) due to an increase of surface water repellency. In contrast, for aboveground biomass yield, the most positive fertilization effects of digestate application were found on sand (up to 3.5 times the biomass of the unamended control) followed by field sand (1.5 times), compared to no effect for field loam. Our findings suggest that digestate application have positive fertilization effects in low-fertility substrates, similar to NPK, even though digestate application may have a negative impact on the permeability in sandy substrates that could interfere with germination.

The South African ragwort (Senecio inaequidens DC.) is one of the fastest exotic plant invaders in Central Europe but, despite its large distribution area, it is still not commonly found in European grasslands. In order to better understand the mechanisms behind invasion resistance of grassland communities to S. inaequidens, we determined (1) to what extent the timing of arrival of S. inaequidens in the community affected its invasiveness as well as the performance of the native species, and (2) how the direction and strength of priority effects were affected by the composition of the native community being invaded, particularly with regard to the presence of N2-fixing species (legumes). In a greenhouse experiment, we manipulated the timing of arrival of the exotic species in the community and the composition of the native background community to test the influence of these factors on the productivity and N content of exotic and native species. Using a set of interaction indices, we also investigated if the plant species origin (native or exotic) and the native community composition affected the benefit of arriving early and the cost of arriving late (i.e., priority effects) in the community. We showed that both exotic and native species created inhibitory priority effects for late-arriving species. The establishment success of S. inaequidens strongly depended on its timing of arrival in a grassland community. On average, S. inaequidens benefited more from arriving early than the natives. We did not find any evidence to support that the presence of legumes in the background community would favour invasion by S. inaequidens. When natives arrived later than S. inaequidens, however, priority effects were weaker when legumes were present in the native community. Synthesis: we showed that priority effects created by natives can lower the risk of invasion by S. inaequidens and are an important mechanism to explain why this exotic species is not commonly found in European grasslands yet. Our results suggest that an early arrival of this species at a site with low native species abundance (e.g., following a disturbance) is a scenario that could favour invasion by S. inaequidens.

Tephrosia apollinea (Leguminosae) is a desert legume shrub from UAE deserts and can fix atmospheric nitrogen through Rhizobium symbiosis. Uniform seedlings of T. apollinea were hydroponically grown in beakers (100 ml) covered with sink strainers in a growth chamber and subjected to different levels of polyethylene glycol (PEG) (0%, 5%, 10%, and 15%) treatment. Potential PEG-induced osmotic stress was evaluated through several plasticity traits such as plant biomass, root nodulation and viability of nodules, and stable isotopes of carbon (δ13C) and nitrogen (δ15N). Osmotic stress decreased plant fresh (FW) and dry biomass (DW), with the largest reduction being found at 15% PEG. Osmotic stress significantly decreased δ13C in both roots and leaves, but the reduction was more prominent in the roots. Conversely, osmotic stress significantly increased δ15N in both roots and leaves. We identified four different strains of nitrogen fixing bacteria (Rhizobium leguminosarum, R. radiobacter, R. meliloti and Bradyrhizobium sp.) from root nodules forming a symbiotic association with T. apollinea. The overall results indicate that carbon allocation, root nodule activity, and carbon and nitrogen isotope signatures are useful functional attributes to measure in relation to adaptive survival of T. apollinea in arid deserts. In addition, the combination of ecophysiological and morphological plasticity in T. apollinea suggest it as an interesting biomass source and nurse plant in nutrient-poor soils of desert environments.

Earth is home to over 350,000 vascular plant species1 that differ in their traits in innumerable ways. Yet, a handful of functional traits can help explaining major differences among species in photosynthetic rate, growth rate, reproductive output and other aspects of plant performance2–6. A key challenge, coined “the Holy Grail” in ecology, is to upscale this understanding in order to predict how natural or anthropogenically driven changes in the identity and diversity of co-occurring plant species drive the functioning of ecosystems7, 8. Here, we analyze the extent to which 42 different ecosystem functions can be predicted by 41 plant traits in 78 experimentally manipulated grassland plots over 10 years. Despite the unprecedented number of traits analyzed, the average percentage of variation in ecosystem functioning that they jointly explained was only moderate (32.6%) within individual years, and even much lower (12.7%) across years. Most other studies linking ecosystem functioning to plant traits analyzed no more than six traits, and when including either only six random or the six most frequently studied traits in our analysis, the average percentage of explained variation in across-year ecosystem functioning dropped to 4.8%. Furthermore, different ecosystem functions were driven by different traits, with on average only 12.2% overlap in significant predictors. Thus, we did not find evidence for the existence of a small set of key traits able to explain variation in multiple ecosystem functions across years. Our results therefore suggest that there are strong limits in the extent to which we can predict the long-term functional consequences of the ongoing, rapid changes in the composition and diversity of plant communities that humanity is currently facing.

Biodiversity-ecosystem functioning (BEF) research grew rapidly following concerns that biodiversity loss would negatively affect ecosystem functions and the ecosystem services they underpin. However, despite evidence that biodiversity strongly affects ecosystem functioning, the influence of BEF research upon policy and the management of ‘real-world’ ecosystems, i.e., semi-natural habitats and agroecosystems, has been limited. Here, we address this issue by classifying BEF research into three clusters based on the degree of human control over species composition and the spatial scale, in terms of grain, of the study, and discussing how the research of each cluster is best suited to inform particular fields of ecosystem management. Research in the first cluster, small-grain highly controlled studies, is best able to provide general insights into mechanisms and to inform the management of species-poor and highly managed systems such as croplands, plantations, and the restoration of heavily degraded ecosystems. Research from the second cluster, small-grain observational studies, and species removal and addition studies, may allow for direct predictions of the impacts of species loss in specific semi-natural ecosystems. Research in the third cluster, large-grain uncontrolled studies, may best inform landscape-scale management and national-scale policy. We discuss barriers to transfer within each cluster and suggest how new research and knowledge exchange mechanisms may overcome these challenges. To meet the potential for BEF research to address global challenges, we recommend transdisciplinary research that goes beyond these current clusters and considers the social-ecological context of the ecosystems in which BEF knowledge is generated. This requires recognizing the social and economic value of biodiversity for ecosystem services at scales, and in units, that matter to land managers and policy makers.

Synthetic fertilizer production is associated with a high environmental footprint, as compounds typically dissolve rapidly leaching emissions to the atmosphere or surface waters. We tested two recovered nutrients with slower release patterns, as promising alternatives for synthetic fertilizers: struvite and a commercially available organic fertilizer. Using these fertilizers as nitrogen source, we conducted a rhizotron experiment to test their effect on plant performance and nutrient recovery in juvenile tomato plants. Plant performance was significantly improved when organic fertilizer was provided, promoting higher shoot biomass. Since the microbial community influences plant nitrogen availability, we characterized the root-associated microbial community structure and functionality. Analyses revealed distinct root microbial community structure when different fertilizers were supplied. However, plant presence significantly increased the similarity of the microbial community over time, regardless of fertilization. Additionally, the presence of the plant significantly reduced the potential ammonia oxidation rates, implying a possible role of the rhizosheath microbiome or nitrification inhibition by the plant. Our results indicate that nitrifying community members are impacted by the type of fertilizer used, while tomato plants influenced the potential ammonia-oxidizing activity of nitrogen-related rhizospheric microbial communities. These novel insights on interactions between recovered fertilizers, plant and associated microbes can contribute to develop sustainable crop production systems. Synthetic fertilizer production is associated with a high environmental footprint, as compounds typically dissolve rapidly leaching emissions to the atmosphere or surface waters. We tested two recovered nutrients with slower release patterns, as promising alternatives for synthetic fertilizers: struvite and a commercially available organic fertilizer. Using these fertilizers as nitrogen source, we conducted a rhizotron experiment to test their effect on plant performance and nutrient recovery in juvenile tomato plants. Plant performance was significantly improved when organic fertilizer was provided, promoting higher shoot biomass. Since the microbial community influences plant nitrogen availability, we characterized the root-associated microbial community structure and functionality. Analyses revealed distinct root microbial community structure when different fertilizers were supplied. However, plant presence significantly increased the similarity of the microbial community over time, regardless of fertilization. Additionally, the presence of the plant significantly reduced the potential ammonia oxidation rates, implying a possible role of the rhizosheath microbiome or nitrification inhibition by the plant. Our results indicate that nitrifying community members are impacted by the type of fertilizer used, while tomato plants influenced the potential ammonia-oxidizing activity of nitrogen-related rhizospheric microbial communities. These novel insights on interactions between recovered fertilizers, plant and associated microbes can contribute to develop sustainable crop production systems.

1. Plant species that arrive first in the system can affect assembly (priority effects). However, effects of order of arrival of different plant functional groups (PFGs) on root development have not yet been investigated under field conditions. 2. We measured standing and fine root length density in the first and third year of a grassland field experiment. We wanted to know if manipulating PFG order of arrival would affect root development, and if priority effects are modulated by soil type. 3. Sowing legumes first created a priority effect that was found in the first and third year, with a lower standing root length density in this treatment, even though the plant community composition was different in each of the studied years. Fine root length density was not affected by order of arrival, but changed according to the soil type.4. Synthesis. We found strong evidence that sowing legumes first created a priority effect below-ground that was found in the first and third year of this field experiment, even though the functional group dominance was different in each of the studied years.

Spatial variation in species composition (‘beta diversity’) is comprised of two components, spatial turnover and nestedness. Disentangling the drivers of each component allows a refined evaluation of the community assembly mechanisms that generate regional diversity patterns. This study examines how environmental sorting as well as past and present dispersal processes regulate spatial turnover and nestedness of plant species in human-transformed grassland landscapes, and whether the relative importance of these drivers varies among two landscapes with contrasting regional soil productivity. We found that nestedness always resulted from differences in historical connectivity to the characteristic species pool of pre-transformed calcareous grassland landscapes, irrespective of regional productivity. This was probably due to dispersal limitation of numerous species from this rich species pool under current landscape configuration and management, the absence of which cannot be compensated for by contemporary immigration rates in modern landscapes. By contrast, spatial turnover was driven by opposing mechanisms in nutrient-rich vs. nutrient-poor landscapes. In the landscape with low soil fertility due to phosphorous (P) limitation, spatial turnover was generated by variation in past and present dispersal processes. In the fertile landscape, species turnover was driven by environmental factors and increased mainly with differences in P and nitrate supply. The findings are consistent with theoretical considerations and empirical evidence from small-scale experiments that often find a reduced importance of dispersal at higher soil resource supply, showing that this pattern also manifests at the landscape scale. In summary, our study indicates that contrasting assembly mechanisms generate spatial compositional turnover in landscapes with contrasting soil conditions, which has significant implications for conservation and habitat restoration. It also demonstrates that detailed reconstructions of the land-use history of individual sites and their surroundings are critical for understanding beta diversity in human-transformed landscapes.

We have detailed knowledge from controlled environment studies on the influence of root temperature on plant performance, growth and morphology. However, in all studies root temperature was kept spatially uniform, which motivated us to test whether a vertical gradient in soil temperature affected development and biomass production. Roots of barley seedlings were exposed to three uniform temperature treatments (10, 15 or 20°C) or to a vertical gradient (20-10°C from top to bottom). Substantial differences in plant performance, biomass production and root architecture occurred in the 30-day-old plants. Shoot and root biomass of plants exposed to vertical temperature gradient increased by 144 respectively, 297%, compared with plants grown at uniform root temperature of 20°C. Additionally the root system was concentrated in the upper 10cm of the soil substrate (98% of total root biomass) in contrast to plants grown at uniform soil temperature of 20°C (86% of total root biomass). N and C concentrations in plant roots grown in the gradient were significantly lower than under uniform growth conditions. These results are important for the transferability of 'normal' greenhouse experiments where generally soil temperature is not controlled or monitored and open a new path to better understand and experimentally assess root-shoot interactions.

The effects of flooding on rhizospheric organic acid concentrations of three abundant flooding tolerant plant species (Alternanthera philoxeroides Mart., Arundinella anomala Steud., Salix variegata Franch.) from the water fluctuation zone of the Three Gorges Reservoir (TGR, Yangtze River) were investigated. Soil solution samples of eight low molecular weight organic acids were obtained from rhizotrons using micro suction cups during 3 weeks of waterlogging, after 6 weeks flooding and after a 1 week recovery. To estimate the contribution of water temperature and microbial community, plants in sterile glass bead substrate and original Yangtze sediment were submerged in laboratory at +10°, +20° and +30°C. Waterlogged plants did seldom express a significantly different pattern of rhizospheric organic acid (OA) composition compared to control plants. Flooding caused no burst of organic acid concentration in soil solution: All species express a silencing strategy. Average OA levels were higher in A. anomala rhizosphere than in the other two species, but increased again after resurfacing in all species. Temperature had a stronger influence in sediment than in sterile setup. In contrast to field measurements, succinate, malate and citrate were detected in the sterile setup. Microbial contribution appeared to have great influence on increasing OA occurrence.

Increasing productivity of mixtures as compared to monocultures has been reported from numerous experimental studies, but so far the variable contribution of individual species to higher mixture productivity in biodiversity experiments is not well understood. To address this issue, we quantified the productivity of 60 species in monocultures and mixtures of varying species richness (2, 4, 8, 16, 60) and functional group number and composition (1, 2, 3, 4; legumes, grasses, small herbs, tall herbs) and tested how species properties are related to species performance in mixtures in the third year after sowing. We analysed monoculture biomass, plant biomass from separately grown plant individuals (=estimate of plant growth rates), and the monoculture resource-use characteristics canopy height and structure (leaf area index) as indicators for light acquisition, and soil nitrate concentration (=estimate of depletion of plant available nitrogen) and biomass:N ratios (=estimate of biomass produced per unit plant N) as indicators for nitrogen acquisition and use. High monoculture productivity was related to different combinations of resource-use characteristics. The biomass of a species and its proportional contribution to mixture biomass correlated positively with species relative yields, suggesting that highly productive mixture species were most important for an overyielding of mixtures. Although monoculture biomass was a significant predictor for species performance in mixtures except for grasses, a combination of monoculture biomass, plant growth rates and resource-use traits associated with nutrient and light acquisition explained non-legume species performance best. Legume performance was best associated with their monoculture biomass and traits associated with light acquisition. In spite of the fact that high species performance in mixtures was associated with a species’ competitive ability as represented by monoculture productivity, growth rates and resource-use traits, our results suggest that species uniqueness in resource acquisition strategies increases the chance for niche differentiation among overyielding species.

Highly variable effects of legumes have been observed in biodiversity experiments, but little is known about plant diversity effects on N2 fixation of legume species. We used the 15N natural abundance method in a non-fertilized regularly mown 6-year biodiversity experiment (Jena Experiment) to quantify N2 fixation of 12 legume species. The proportion of legume N derived from the atmosphere (%Ndfa) differed significantly among legume species. %Ndfa values were lower in 2004 after setting-up the experiment (73 ± 20) than in the later years (2006: 80 ± 16; 2008: 78 ± 12). Increasing species richness had positive effects on %Ndfa in 2004 and 2006, but not in 2008. High biomass production of legumes in 2004 and 2006 declined to lower levels in 2008. In 2006, legume positioning within the canopy best explained variation in %Ndfa values indicating a lower reliance of tall legumes on N2 fixation. In 2008, larger %Ndfa values of legumes were related to lower leaf P concentrations suggesting that the availability of phosphorus limited growth of legumes. In summary, diversity effects on N2 fixation depend on legume species identity, their ability to compete for soil nutrients and light and may vary temporally in response to changing resource availability.

Under elevated atmospheric CO2 concentrations, soil carbon (C) inputs are typically enhanced, suggesting larger soil C sequestration potential. However, soil C losses also increase and progressive nitrogen (N) limitation to plant growth may reduce the CO2 effect on soil C inputs with time. We compiled a data set from 131 manipulation experiments, and used meta-analysis to test the hypotheses that: (1) elevated atmospheric CO2 stimulates soil C inputs more than C losses, resulting in increasing soil C stocks; and (2) that these responses are modulated by N. Our results confirm that elevated CO2 induces a C allocation shift towards below-ground biomass compartments. However, the increased soil C inputs were offset by increased heterotrophic respiration (Rh), such that soil C content was not affected by elevated CO2. Soil N concentration strongly interacted with CO2 fumigation: the effect of elevated CO2 on fine root biomass and –production and on microbial activity increased with increasing soil N concentration, while the effect on soil C content decreased with increasing soil N concentration. These results suggest that both plant growth and microbial activity responses to elevated CO2 are modulated by N availability, and that it is essential to account for soil N concentration in C cycling analyses.

Developing and strengthening a more mutualistic relationship between the science of restoration ecology and the practice of ecological restoration has been a central but elusive goal of SERI since its inaugural meeting in 1989. We surveyed the delegates to the 2009 SERI World Conference to learn more about their perceptions of and ideas for improving restoration science, practice, and scientist/practitioner relationships. The respondents' assessments of restoration practice were less optimistic than their assessments of restoration science. Only 26% believed that scientist/practitioner relationships were “generally mutually beneficial and supportive of each other,” and the “science–practice gap” was the second and third most frequently cited category of factors limiting the science and practice of restoration, respectively (“insufficient funding” was first in both cases). Although few faulted practitioners for ignoring available science, many criticized scientists for ignoring the pressing needs of practitioners and/or failing to effectively communicate their work to nonscientists. Most of the suggestions for bridging the gap between restoration science and practice focused on (1) developing the necessary political support for more funding of restoration science, practice, and outreach; and (2) creating alternative research paradigms to both facilitate on-the-ground projects and promote more mutualistic exchanges between scientists and practitioners. We suggest that one way to implement these recommendations is to create a “Restoration Extension Service” modeled after the United States Department of Agriculture's Cooperative Extension Service. We also recommend more events that bring together a fuller spectrum of restoration scientists, practitioners, and relevant stakeholders.

Summary Unravelling the factors determining the allocation of carbon to various plant organs is one of the great challenges of modern plant biology. Studying allocation under close to natural conditions requires non-invasive methods, which are now becoming available for measuring plants on a par with those developed for humans. By combining magnetic resonance imaging (MRI) and positron emission tomography (PET), we investigated three contrasting root/shoot systems growing in sand or soil, with respect to their structures, transport routes and the translocation dynamics of recently fixed photoassimilates labelled with the short-lived radioactive carbon isotope 11C. Storage organs of sugar beet (Beta vulgaris) and radish plants (Raphanus sativus) were assessed using MRI, providing images of the internal structures of the organs with high spatial resolution, and while species-specific transport sectoralities, properties of assimilate allocation and unloading characteristics were measured using PET. Growth and carbon allocation within complex root systems were monitored in maize plants (Zea mays), and the results may be used to identify factors affecting root growth in natural substrates or in competition with roots of other plants. MRI–PET co-registration opens the door for non-invasive analysis of plant structures and transport processes that may change in response to genomic, developmental or environmental challenges. It is our aim to make the methods applicable for quantitative analyses of plant traits in phenotyping as well as in understanding the dynamics of key processes that are essential to plant performance.