Effect of plant root symbionts on performance of native woody species in competition with an invasive grass in multispecies microcosms

The majority of terrestrial plants form mutualistic associations with arbuscular mycorrhizal fungi (AMF) and rhizobia (i.e., nitrogen‐fixing bacteria). Understanding these associations has important implications for ecological theory and for restoration practice. Here, we tested whether the presence of AMF and rhizobia influences the performance of native woody plants invaded by a non‐native grass in experimental microcosms. We planted eight plant species (i.e., Acacia acuminata, A. microbotrya, Eucalyptus loxophleba subsp. loxophleba, E. astringens, Calothamnus quadrifidus, Callistemon phoeniceus, Hakea lissocarpha and H. prostrata) in microcosms of field‐conditioned soil with and without addition of AMF and rhizobia in a fully factorial experimental design. After seedling establishment, we seeded half the microcosms with an invasive grass Bromus diandrus. We measured shoot and root biomass of native plants and Bromus, and on roots, the percentage colonization by AMF, number of rhizobia‐forming nodules and number of proteaceous root clusters. We found no effect of plant root symbionts or Bromus addition on performance of myrtaceous, and as predicted, proteaceous species as they rely little or not at all on AMF and rhizobia. Soil treatments with AMF and rhizobia had a strong positive effect (i.e., larger biomass) on native legumes (A. microbotrya and A. acuminata). However, the beneficial effect of root symbionts on legumes became negative (i.e., lower biomass and less nodules) if Bromus was present, especially for one legume, i.e., A. acuminata, suggesting a disruptive effect of the invader on the mutualism. We also found a stimulating effect of Bromus on root nodule production in A. microbotrya and AMF colonization in A. acuminata which could be indicative of legumes’ increased resource acquisition requirement, i.e., for nitrogen and phosphorus, respectively, in response to the Bromus addition. We have demonstrated the importance of measuring belowground effects because the aboveground effects gave limited indication of the effects occurring belowground.     

Plant–fungal symbiosis affects litter decomposition during primary succession

Microbial symbionts of plants can affect decomposition by altering the quality or quantity of host plant tissue (substrate) or the micro‐environment where decomposition occurs (conditioning). In C₃ grasses, foliar fungal endophytes (Clavicipitaceae) can increase plant resistance to drought and/or produce alkaloids that reduce herbivory – effects that may also influence host litter composition and subsequent litter decomposition. We studied the effect of the endophyte Epichloë sp. on litter decomposition in the Great Lakes dunes (USA) using a reciprocal design altering endophyte presence/absence in both American beachgrass Ammophila breviligulata substrate (litter bags) and its conditioning of the decomposition microenvironment. Symbiont treatments were crossed with rain‐out shelters that altered growing season precipitation. The first year of decomposition, senesced leaf substrate from A. breviligulata with Epichloë decomposed 21% faster than endophyte‐free substrate. By the third year, conditioning by live symbiotic plants reduced cumulative decomposition by 33% compared to plots planted with endophyte‐free plants. Of the traits we examined – litter quantity, C:N ratio, mineral composition, fungal colonization, and carbon chemistry – increased litter quantity via greater tiller production was the primary trait shift associated with endophyte symbiosis. Epichloë in A. breviligulata litter also altered litter nitrogen decomposition dynamics, as evidenced by lower nitrogen and protein content in decomposed tissue from plants that hosted the endophyte. Differences in initial litter quality and subsequent colonization by saprotrophic fungi were ruled out as key drivers. Altered precipitation had negligible effects on decomposing processes in the dunes. Grass–Epichloë symbiosis altered nutrient cycling through increasing the rate of litter decomposition when present in the litter and through reducing litter decomposition by conditioning the decomposition microenvironment. Epichloë are widespread symbionts of grasses. Thus, their effects on decomposition could be an important, but often overlooked, driver of nutrient cycling in grass‐dominated ecosystems.     

Insights into the history of the legume‐betaproteobacterial symbiosis

The interaction between legumes and rhizobia has been well studied in the context of a mutualistic, nitrogen‐fixing symbiosis. The fitness of legumes, including important agricultural crops, is enhanced by the plants’ ability to develop symbiotic associations with certain soil bacteria that fix atmospheric nitrogen into a utilizable form, namely, ammonia, via a chemical reaction that only bacteria and archaea can perform. Of the bacteria, members of the alpha subclass of the protebacteria are the best‐known nitrogen‐fixing symbionts of legumes. Recently, members of the beta subclass of the proteobacteria that induce nitrogen‐fixing nodules on legume roots in a species‐specific manner have been identified. In this issue, Bontemps et al. reveal that not only are these newly identified rhizobia novel in shifting the paradigm of our understanding of legume symbiosis, but also, based on symbiotic gene phylogenies, have a history that is both ancient and stable. Expanding our understanding of novel plant growth promoting rhizobia will be a valuable resource for incorporating alternative strategies of nitrogen fixation for enhancing plant growth.     

Competition for alfalfa nodulation under metal stress by the metal‐tolerant strain Ochrobactrum cytisi Azn6.2

Legume plants, in association with rhizobia, are gaining increasing interest for heavy metal rhizoremediation. This symbiotic interaction combines the advantages of rhizoremediation and soil nitrogen enrichment. In metal polluted soils, Ochrobactrum cytisi can elicit non‐fixing nodules on legumes, including Medicago sativa. Nodulation kinetics was much slower when M. sativa plants were inoculated with O. cytisi Azn6.2 compared with the natural symbiont Ensifer meliloti 1021 and nodules were ineffective in nitrogen fixation. A competition experiment was performed using alfalfa grown on heavy metals, and co‐inoculated with equal amounts of the metal‐sensitive E. meliloti 1021 and the metal‐resistant O. cytisi Azn6.2. When plants were inoculated in non‐polluted substrates, all nodules were formed by E. meliloti 1021. Nevertheless, under increasing metal concentrations, the number of nodules occupied by O. cytisi grew. At the highest metal concentration, all nodules were elicited by O. cytisi, suggesting that the resistant species can take the place of the natural symbiont. This fact has important ecological and environmental implications when proposing legume–rhizobia symbioses for rhizoremediation and highlights the need of selecting highly resistant rhizobia in order to be competitive in polluted soils.     

Balancing multiple mutualists: asymmetric interactions among plants, arbuscular mycorrhizal fungi, and fungal endophytes

Most organisms engage in beneficial interactions with other species; however, little is known regarding how individuals balance the competing demands of multiple mutualisms. Here we examine three-way interactions among a widespread grass, Schedonorus phoenix , a protective fungal endophyte aboveground, Neotyphodium coenophialum , and nutritional symbionts (arbuscular mycorrhizal fungi) belowground. In a greenhouse experiment, we manipulated the presence/absence of both fungi and applied a fertilizer treatment to individual plants. Endophyte presence in host plants strongly reduced mycorrhizal colonization of roots. Additionally, for plants with the endophyte, the density of endophyte hyphae was negatively correlated with mycorrhizal colonization, suggesting a novel role for endophyte abundance in the interaction between the symbionts. Endophyte presence increased plant biomass, and there was a positive correlation between endophyte hyphal density and plant biomass. The effects of mutualists were asymmetric: mycorrhizal fungi treatments had no significant impact on the endophyte and negligible effects on plant biomass. Fertilization affected all three species – increasing plant biomass and endophyte density, but diminishing mycorrhizal colonization. Mechanisms driving negative effects of endophytes on mycorrhizae may include inhibition via endophyte alkaloids, altered nutritional requirements of the host plant, and/or temporal and spatial priority effects in the interactions among plants and multiple symbionts.     

Potential for endophyte symbiosis to increase resistance of the native grass <Emphasis Type="Italic">Poa alsodes</Emphasis> to invasion by the non-native grass <Emphasis Type="Italic">Microstegium vimineum</Emphasis>

Microbial symbionts can improve the competitive ability and stress tolerance of plant hosts and thus may enhance native plant resistance against invaders. We investigated whether symbiosis between a native grass, Poa alsodes, and a fungal endophyte (Neotyphodium sp.) improved the grass’s ability to compete against Microstegium vimineum (Japanese stiltgrass), a common invader in the eastern USA. We challenged naturally endophyte-symbiotic and experimentally endophyte-free P. alsodes plants with the invader. In the first experiment, we manipulated symbiosis and water availability to test for context-dependency in symbiont benefits. In the second experiment, we manipulated symbiosis and M. vimineum diversity (the number of invader populations), since greater intraspecific diversity is expected to improve invasion success and might alter the efficacy of symbiosis in invasion resistance. In both experiments, presence of the endophyte reduced the per plant biomass of M. vimineum and increased P. alsodes biomass. We found no evidence that benefits of the symbiont depended on water availability, and population-level diversity had a minor influence on M. vimineum: inflorescence number showed a parabolic relationship with increasing numbers of M. vimineum populations. Overall, symbiosis in the native grass had stronger effects on invader growth than either water availability or invader genetic diversity. Our results suggest that endophyte symbioses in native plants can increase host performance against an invader, although this conclusion needs confirmation in more complex field settings where other important factors, such as herbivores and fluctuating abiotic conditions, come into play.     

Interspecific conflict and the evolution of ineffective rhizobia

Microbial symbionts exhibit broad genotypic variation in their fitness effects on hosts, leaving hosts vulnerable to costly partnerships. Interspecific conflict and partner‐maladaptation are frameworks to explain this variation, with different implications for mutualism stability. We investigated the mutualist service of nitrogen fixation in a metapopulation of root‐nodule forming Bradyrhizobium symbionts in Acmispon hosts. We uncovered Bradyrhizobium genotypes that provide negligible mutualist services to hosts and had superior in planta fitness during clonal infections, consistent with cheater strains that destabilise mutualisms. Interspecific conflict was also confirmed at the metapopulation level – by a significant negative association between the fitness benefits provided by Bradyrhizobium genotypes and their local genotype frequencies – indicating that selection favours cheating rhizobia. Legumes have mechanisms to defend against rhizobia that fail to fix sufficient nitrogen, but these data support predictions that rhizobia can subvert plant defenses and evolve to exploit hosts.     

禾-豆混播草地中土壤水分与种间关系研究进展

从植物形态变化和生理反应两方面探讨了禾草和豆科牧草在干旱环境下所采取的不同生存策略,阐述了禾-豆之间存在的水分竞争关系以及豆科植物的水力提升作用对水分资源高效利用的意义,分析了干旱对豆科植物固氮和转氮能力的影响.提出应探讨适于我国干旱区域的深根系豆科牧草与禾草的共存机制,认为揭示水分因子对地上竞争和地下竞争以及对两种竞争间反馈关系的影响是研究其共存机制的一个重要方面.     

MALDI mass spectrometry‐assisted molecular imaging of metabolites during nitrogen fixation in the Medicago truncatula–Sinorhizobium meliloti symbiosis

Symbiotic associations between leguminous plants and nitrogen‐fixing rhizobia culminate in the formation of specialized organs called root nodules, in which the rhizobia fix atmospheric nitrogen and transfer it to the plant. Efficient biological nitrogen fixation depends on metabolites produced by and exchanged between both partners. The Medicago truncatula–Sinorhizobium meliloti association is an excellent model for dissecting this nitrogen‐fixing symbiosis because of the availability of genetic information for both symbiotic partners. Here, we employed a powerful imaging technique – matrix‐assisted laser desorption/ionization (MALDI)/mass spectrometric imaging (MSI) – to study metabolite distribution in roots and root nodules of M. truncatula during nitrogen fixation. The combination of an efficient, novel MALDI matrix [1,8–bis(dimethyl‐amino) naphthalene, DMAN] with a conventional matrix 2,5–dihydroxybenzoic acid (DHB) allowed detection of a large array of organic acids, amino acids, sugars, lipids, flavonoids and their conjugates with improved coverage. Ion density maps of representative metabolites are presented and correlated with the nitrogen fixation process. We demonstrate differences in metabolite distribution between roots and nodules, and also between fixing and non‐fixing nodules produced by plant and bacterial mutants. Our study highlights the benefits of using MSI for detecting differences in metabolite distributions in plant biology.     

Interrelated effects of mycorrhiza and free‐living nitrogen fixers cascade up to aboveground herbivores

Aboveground plant performance is strongly influenced by belowground microorganisms, some of which are pathogenic and have negative effects, while others, such as nitrogen‐fixing bacteria and arbuscular mycorrhizal fungi, usually have positive effects. Recent research revealed that belowground interactions between plants and functionally distinct groups of microorganisms cascade up to aboveground plant associates such as herbivores and their natural enemies. However, while functionally distinct belowground microorganisms commonly co‐occur in the rhizosphere, their combined effects, and relative contributions, respectively, on performance of aboveground plant‐associated organisms are virtually unexplored. Here, we scrutinized and disentangled the effects of free‐living nitrogen‐fixing (diazotrophic) bacteria Azotobacter chroococcum (DB) and arbuscular mycorrhizal fungi Glomus mosseae (AMF) on host plant choice and reproduction of the herbivorous two‐spotted spider mite Tetranychus urticae on common bean plants Phaseolus vulgaris. Additionally, we assessed plant growth, and AMF and DB occurrence and density as affected by each other. Both AMF alone and DB alone increased spider mite reproduction to similar levels, as compared to the control, and exerted additive effects under co‐occurrence. These effects were similarly apparent in host plant choice, that is, the mites preferred leaves from plants with both AMF and DB to plants with AMF or DB to plants grown without AMF and DB. DB, which also act as AMF helper bacteria, enhanced root colonization by AMF, whereas AMF did not affect DB abundance. AMF but not DB increased growth of reproductive plant tissue and seed production, respectively. Both AMF and DB increased the biomass of vegetative aboveground plant tissue. Our study breaks new ground in multitrophic belowground–aboveground research by providing first insights into the fitness implications of plant‐mediated interactions between interrelated belowground fungi–bacteria and aboveground herbivores.     

Origins of cheating and loss of symbiosis in wild Bradyrhizobium

Rhizobial bacteria nodulate legume roots and fix nitrogen in exchange for photosynthates. These symbionts are infectiously acquired from the environment and in such cases selection models predict evolutionary spread of uncooperative mutants. Uncooperative rhizobia – including nonfixing and non‐nodulating strains – appear common in agriculture, yet their population biology and origins remain unknown in natural soils. Here, a phylogenetically broad sample of 62 wild‐collected rhizobial isolates was experimentally inoculated onto Lotus strigosus to assess their nodulation ability and effects on host growth. A cheater strain was discovered that proliferated in host tissue while offering no benefit; its fitness was superior to that of beneficial strains. Phylogenetic reconstruction of Bradyrhizobium rDNA and transmissible symbiosis‐island loci suggest that the cheater evolved via symbiotic gene transfer. Many strains were also identified that failed to nodulate L. strigosus, and it appears that nodulation ability on this host has been recurrently lost in the symbiont population. This is the first study to reveal the adaptive nature of rhizobial cheating and to trace the evolutionary origins of uncooperative rhizobial mutants.     

Three symbionts involved in interspecific plant-soil feedback: epichloid endophytes and mycorrhizal fungi affect the performance of rhizobia-legume symbiosis

Aims

Plants interact by modifying soil conditions in plant-soil feedback processes. Foliar endophytes of grasses exert multiple effects on host rhizosphere with potential consequences on plant-soil feedback. Here, we hypothesize that the grass-endophyte symbiosis impairs soil symbiotic potential, and in turn influences legume performance and nitrogen acquisition.

Methods

Soil was conditioned in pots, growing Lolium multiflorum with or without the fungal endophyte Epichloë and with or without arbuscular mycorrhizal fungi (AMF). Then, Trifolium repens grew in all types of conditioned soils with high or low rhizobia availability.

Results

Endophyte soil conditioning reduced AMF spores number and rhizobial nodules (?27 % and ?38 %, respectively). Seedling survival was lower in endophyte-conditioned soil and higher in mycorrhizal soils (?27 % and +24 %, respectively). High rhizobia-availability allowed greater growth and nitrogen acquisition, independent of soil conditioning. Low rhizobia-availability allowed both effects only in endophyte-conditioned soil.

Conclusion

Endophyte-induced changes in soil (i) hindered symbiotic potential by reducing AMF spore availability or rhizobia nodulation, (ii) impaired legume survival irrespective of belowground symbionts presence, but (iii) mimicked rhizobia effects, enhancing growth and nitrogen fixation in poorly nodulated plants. Our results show that shoot and root symbionts can be interactively involved in interspecific plant-soil feedback.

     

Diversity and abundance of macro‐invertebrates on abandoned cattle kraals in a semi‐arid savanna

Abandoned cattle (Bos taurus) kraals are sources of habitat heterogeneity in dystrophic semi‐arid African savannas with a strong positive effect on soil nutrients and plant productivity. However, little is known regarding how macro‐invertebrate assemblages vary between abandoned kraals and the surrounding savanna matrix. We tested whether herbaceous biomass and basal and aerial covers and soil nutrients have an effect on aboveground and belowground macro‐invertebrate assemblages. Twelve abandoned kraals were contrasted with their paired control plots for soil characteristics, herbaceous productivity, and macro‐invertebrate assemblages in Save Valley Conservancy, Zimbabwe. Abandoned kraals had significantly higher concentrations of soil nitrogen (N), phosphorus (P), potassium (K), and calcium (Ca) as well as herbaceous biomass and basal and aerial covers than control plots. Both aboveground and belowground macro‐invertebrate species richness were higher on abandoned kraals. However, only belowground macro‐invertebrate diversity (Shannon H′ and Hill number 1) was significantly higher on abandoned kraals. Soil nutrients and herbaceous productivity had positive and significant correlations with the dominant taxa (Coleoptera, Hymenoptera, Hemiptera, Isoptera, and Myriapoda) on abandoned kraals. These results add to the growing body of evidence that abandoned kraals exert significant effects on savanna spatial heterogeneity years later, with implications on ecosystem processes and functioning.     

不同水分处理下双接种丛枝菌根真菌和根瘤菌对疏叶骆驼刺生长及氮素转移的影响

以疏叶骆驼刺为研究对象,设定3个水分梯度正常水分(土壤相对含水量(70±5)%)、干旱胁迫(土壤相对含水量(20±5)%)和复水处理(干旱胁迫60天后恢复至正常水分)与四组接种处理(单接种丛枝菌根真菌(AMF)、单接种根瘤菌、双接种AMF+根瘤菌和不接种),分析不同水分条件下双接种丛枝菌根真菌和根瘤菌对疏叶骆驼刺的生长以及供、受体疏叶骆驼刺之间氮素转移的影响。结果表明,正常水分处理时,双接种疏叶骆驼刺的AMF侵染率、地上生物量、地下生物量、总生物量以及氮含量均要高于单接种处理;根瘤数量、最大荧光(Fm)、初始荧光(Fo)、最大光化学效率(Fv/Fm)与单接种处理之间无差异;在遭遇干旱胁迫时,双接种疏叶骆驼刺的AMF侵染率、总生物量、Fv/Fm均小于单接种处理;地上生物量、地下生物量、根瘤数、Fm、Fo以及氮含量与单接种之间无差异。复水后,双接种疏叶骆驼刺的地上生物量、地下生物量、总生物量、根瘤数均优于单接种;AMF侵染率、氮含量低于单接种;Fm、Fo、Fv/Fm均与单接种之间无差异。在氮素转移方面,正常水分时,双接种与单接种的氮素转移率无差异,在遭遇干旱胁迫时,双接种疏叶骆驼刺的氮素转...     

Consequences of simultaneous interactions of fungal endophytes and arbuscular mycorrhizal fungi with a shared host grass

Plants simultaneously associate with multiple microbial symbionts throughout their lifetimes. To address the question of whether the effects of simultaneous symbionts are contingent on the specific identities, we conducted a greenhouse experiment manipulating the presence and identities of arbuscular mycorrhizal fungi (AMF) and fungal endophytes on the shared host grass Elymus hystrix. Each plant host was inoculated with one of two AMF species having varying effects on host growth, or a sterile soil control. Further, we used naturally occurring endophyte‐infected (E+) and uninfected (E–) individuals from two populations of the endophyte Epichloë elymi that varied in their interaction with E. hystrix. We then measured responses of plants, AMF, and fungal endophytes. Overall, we found that the combined effects of AMF and fungal endophytes on plant growth were additive, reflecting the mutualistic quality of each symbiont independently interacting with host plants. However, fungal endophyte infection differentially altered hyphal colonization of the two AMF species and the identity of the coinfecting AMF species affected fungal endophyte fitness traits. The results of this study demonstrate that the outcome of interspecific symbiotic interactions varies with partner identity such that the effects of simultaneous symbioses can not be generalized.     

Earthworm and belowground competition effects on plant productivity in a plant diversity gradient

Diversity is one major factor driving plant productivity in temperate grasslands. Although decomposers like earthworms are known to affect plant productivity, interacting effects of plant diversity and earthworms on plant productivity have been neglected in field studies. We investigated in the field the effects of earthworms on plant productivity, their interaction with plant species and functional group richness, and their effects on belowground plant competition. In the framework of the Jena Experiment we determined plant community productivity (in 2004 and 2007) and performance of two phytometer plant species [Centaurea jacea (herb) and Lolium perenne (grass); in 2007 and 2008] in a plant species (from one to 16) and functional group richness gradient (from one to four). We sampled earthworm subplots and subplots with decreased earthworm density and reduced aboveground competition of phytometer plants by removing the shoot biomass of the resident plant community. Earthworms increased total plant community productivity (+11%), legume shoot biomass (+35%) and shoot biomass of the phytometer C. jacea (+21%). Further, phytometer performance decreased, i.e. belowground competition increased, with increasing plant species and functional group richness. Although single plant functional groups benefited from higher earthworm numbers, the effects did not vary with plant species and functional group richness. The present study indicates that earthworms indeed affect the productivity of semi-natural grasslands irrespective of the diversity of the plant community. Belowground competition increased with increasing plant species diversity. However, belowground competition was modified by earthworms as reflected by increased productivity of the phytometer C. jacea. Moreover, particularly legumes benefited from earthworm presence. Considering also previous studies, we suggest that earthworms and legumes form a loose mutualistic relationship affecting essential ecosystem functions in temperate grasslands, in particular decomposition and plant productivity. Further, earthworms likely alter competitive interactions among plants and the structure of plant communities by beneficially affecting certain plant functional groups.     

Cascading effects of belowground predators on plant communities are density‐dependent

Soil food webs comprise a multitude of trophic interactions that can affect the composition and productivity of plant communities. Belowground predators feeding on microbial grazers like Collembola could decelerate nutrient mineralization by reducing microbial turnover in the soil, which in turn could negatively influence plant growth. However, empirical evidences for the ecological significance of belowground predators on nutrient cycling and plant communities are scarce. Here, we manipulated predator density (Hypoaspis aculeifer: predatory mite) with equal densities of three Collembola species as a prey in four functionally dissimilar plant communities in experimental microcosms: grass monoculture (Poa pratensis), herb monoculture (Rumex acetosa), legume monoculture (Trifolium pratense), and all three species as a mixed plant community. Density manipulation of predators allowed us to test for density‐mediated effects of belowground predators on Collembola and lower trophic groups. We hypothesized that predator density will reduce Collembola population causing a decrease in nutrient mineralization and hence detrimentally affect plant growth. First, we found a density‐dependent population change in predators, that is, an increase in low‐density treatments, but a decrease in high‐density treatments. Second, prey suppression was lower at high predator density, which caused a shift in the soil microbial community by increasing the fungal: bacterial biomass ratio, and an increase of nitrification rates, particularly in legume monocultures. Despite the increase in nutrient mineralization, legume monocultures performed worse at high predator density. Further, individual grass shoot biomass decreased in monocultures, while it increased in mixed plant communities with increasing predator density, which coincided with elevated soil N uptake by grasses. As a consequence, high predator density significantly increased plant complementarity effects indicating a decrease in interspecific plant competition. These results highlight that belowground predators can relax interspecific plant competition by increasing nutrient mineralization through their density‐dependent cascading effects on detritivore and soil microbial communities.     

The role of legumes as a component of biodiversity in a cross-European study of grassland biomass nitrogen

To investigate how plant diversity loss affects nitrogen accumulation in above‐ground plant biomass and how consistent patterns are across sites of different climatic and soil conditions, we varied the number of plant species and functional groups (grasses, herbs and legumes) in experimental grassland communities across seven European experimental sites (Switzerland, Germany, Ireland, United Kingdom (Silwood Park), Portugal, Sweden and Greece). Nitrogen pools were significantly affected by both plant diversity and community composition. Two years after sowing, nitrogen pools in Germany and Switzerland strongly increased in the presence of legumes. Legume effects on nitrogen pools were less pronounced at the Swedish, Irish and Portuguese site. In Greece and UK there were no legume effects. Nitrogen concentration in total above‐ground biomass was quite invariable at 1.66±0.03% across all sites and diversity treatments. Thus, the presence of legumes had a positive effect on nitrogen pools by significantly increasing above‐ground biomass, i.e. by increases in vegetation quantity rather than quality. At the German site with the strongest legume effect on nitrogen pools and biomass, nitrogen that was fixed symbiotically by legumes was transferred to the other plant functional groups (grasses and herbs) but varied depending on the particular legume species fixing N and the non‐legume species taking it up. Nitrogen‐fixation by legumes therefore appeared to be one of the major functional traits of species that influenced nitrogen accumulation and biomass production, although effects varied among sites and legume species. This study demonstrates that the consequences of species loss on the nitrogen budget of plant communities may be more severe if legume species are lost. However, our data indicate that legume species differ in their N₂ fixation. Therefore, loss of an efficient N₂‐fixer (Trifolium in our study) may have a greater influence on the ecosystem function than loss of a less efficient species (Lotus in our study). Furthermore, there is indication that P availability in the soil facilitates the legume effect on biomass production and biomass nitrogen accumulation.     

Shifting grassland plant community structure drives positive interactive effects of warming and diversity on aboveground net primary productivity

Ecosystems worldwide are increasingly impacted by multiple drivers of environmental change, including climate warming and loss of biodiversity. We show, using a long‐term factorial experiment, that plant diversity loss alters the effects of warming on productivity. Aboveground primary productivity was increased by both high plant diversity and warming, and, in concert, warming (≈1.5 °C average above and belowground warming over the growing season) and diversity caused a greater than additive increase in aboveground productivity. The aboveground warming effects increased over time, particularly at higher levels of diversity, perhaps because of warming‐induced increases in legume and C4 bunch grass abundances, and facilitative feedbacks of these species on productivity. Moreover, higher plant diversity was associated with the amelioration of warming‐induced environmental conditions. This led to cooler temperatures, decreased vapor pressure deficit, and increased surface soil moisture in higher diversity communities. Root biomass (0–30 cm) was likewise consistently greater at higher plant diversity and was greater with warming in monocultures and at intermediate diversity, but at high diversity warming had no detectable effect. This may be because warming increased the abundance of legumes, which have lower root : shoot ratios than the other types of plants. In addition, legumes increase soil nitrogen (N) supply, which could make N less limiting to other species and potentially decrease their investment in roots. The negative warming × diversity interaction on root mass led to an overall negative interactive effect of these two global change factors on the sum of above and belowground biomass, and thus likely on total plant carbon stores. In total, plant diversity increased the effect of warming on aboveground net productivity and moderated the effect on root mass. These divergent effects suggest that warming and changes in plant diversity are likely to have both interactive and divergent impacts on various aspects of ecosystem functioning.     

Endophyte species influence the biomass production of the native grass Achnatherum sibiricum (L.) Keng under high nitrogen availability

Research on the interaction of endophytes and native grasses normally takes infection status into account, but less often considers the species of endophyte involved in the interaction. Here, we examined the effect of endophyte infection, endophyte species, nitrogen availability, and plant maternal genotype on the performance of a wild grass, Achnatherum sibiricum. Six different Epichloë‐infected maternal lines of A. sibiricum were used in the study; three lines harbored Epichloë gansuensis (Eg), while three lines harbored Epichloë sibirica (Es). These endophytes are vertically transmitted, while Eg also occasionally produces stromata on host tillers. We experimentally removed the endophyte from some ramets of the six lines, with the infected (E+) and uninfected (E?) plants grown under varying levels of nitrogen availability. Eg hosts produced more aboveground biomass than Es hosts only under high nitrogen supply. Endophyte species did not show any influence on the maximum net photosynthetic rate (P_max), photosynthetic nitrogen use efficiency, or total phenolics of A. sibiricum under all nitrogen conditions. However, the plant maternal genotype did influence the P_max and shoot biomass of A. sibiricum. Our results show that endophyte species influenced the shoot biomass of A. sibiricum, and this effect was dependent on nitrogen supply. As with most coevolutionary interactions, A. sibiricum that harbored Eg and Es may show pronounced geographic variation in natural habitats with increased nitrogen deposition. In addition, stroma‐bearing endophyte (Eg) provides positive effects (e.g., higher biomass production) to A. sibiricum plants during the vegetative growth stage.