Both evenness and dominant species identity have effects on litter decomposition.

Exploring how interactions between species evenness and dominant species identity affect litter decomposition processes is vital to understanding the relationship between biodiversity and ecosystem functioning in the context of global changes. We carried out a 127-day litter decomposition experiment under controlled conditions, with interactions of four species evenness types (high, medium, low and single species) and three dominant species identity (Leymus chinensis, Serratula centauroides, Artemisia capillaris). After collecting the remaining litter, we estimated how evenness and dominant species identity affected litter mass loss rate, carbon (C) loss rate, nitrogen (N) loss rate and remaining litter C/N directly or indirectly, and assessed relative mixture effects (RMEs) on litter mass loss. The main results are shown as follows. (1) By generalized linear models, litter mass loss rate was significantly affected by evenness after 69-day decomposition; N loss rate was affected by dominant species identity after 69-day decomposition, with treatment dominated by Serratula centauroides being at least 9.26% higher than that dominated by any of other species; and remaining litter C/N was affected by the interactions between evenness and dominant species identity after 30-, 69- and 127-day decomposition. (2) Twenty-three out of 27 RMEs were additive, and dominant species identity showed a significant effect on RMEs after 127-day decomposition. (3) By confirmatory path analyses, litter mass loss rate was affected by dominant species identity directly after 127-day decomposition, and by both species evenness and dominant species identity indirectly which was mediated by initial litter functional dispersion (FDis) after 30- and 69-day decomposition; remaining litter C/N was affected by evenness indirectly which was mediated by initial litter FDis after 127-day decomposition. These findings highlight the importance of evenness and dominant species identity on litter decomposition. The study provides insights into communities during retrogressive successions in semi-arid grasslands in the context of global changes.

Epichloë Endophyte Enhanced Insect Resistance of Host Grass Leymus Chinensis by Affecting Volatile Organic Compound Emissions.

In plant-herbivore interactions, plant volatile organic compounds (VOCs) play an important role in anti-herbivore defense. Grasses and Epichloë endophytes often form defensive mutualistic symbioses. Most Epichloë species produce alkaloids to protect hosts from herbivores, but there is no strong evidence that endophytes can affect the insect resistance of their hosts by altering VOC emissions. In this study, a native dominant grass, sheepgrass (Leymus chinensis), and its herbivore, oriental migratory locust (Locusta migratoria), were used as experimental materials. We studied the effect of endophyte-associated VOC emissions on the insect resistance of L. chinensis. The results showed that endophyte infection enhanced insect resistance of the host, and locusts preferred the odor of endophyte-free (EF) leaves to that of endophyte-infected (EI) leaves. We determined the VOC profile of L. chinensis using gas chromatography-mass spectrometry (GC-MS), and found that endophyte infection decreased the pentadecane (an alkane) emission from uneaten plants, and increased the nonanal (an aldehyde) emission from eaten plants. The olfactory response experiment showed that locusts were attracted by high concentration of pentadecane, while repelled by high concentration of nonanal, indicating that Epichloë endophytes may increase locust resistance of L. chinensis by decreasing pentadecane while increasing nonanal emission. Our results suggest that endophytes can induce VOC-mediated defense in hosts in addition to producing alkaloids, contributing to a better understanding the endophyte-plant-herbivore interactions.

Intraspecific trait variation and adaptability of Stipa krylovii: Insight from a common garden experiment with two soil moisture treatments.

Understanding patterns of intraspecific trait variation can help us understand plant adaptability to environmental changes. To explore the underlying adaptation mechanisms of zonal plant species, we selected seven populations of Stipa krylovii, a dominant species in the Inner Mongolia Steppe of China, and evaluated the effects of phenotypic plasticity and genetic differentiation, the effects of climate variables on population trait differentiation, and traits coordinated patterns under each soil moisture treatment. We selected seeds from seven populations of S. krylovii in the Inner Mongolia Steppe, China, and carried out a soil moisture (2) × population origin (7) common garden experiment at Tianjin City, China, and measured ten plant traits of S. krylovii. General linear analyses were used to analyze how soil moisture and population origin affected each trait variation, Mantel tests were used to analyze population trait differentiation-geographic distance (or climatic difference) relationships, regression analyses were used to evaluate trait-climatic variable relationships, and plant trait networks (PTNs) were used to evaluate traits coordinated patterns. Both soil moisture and population origin showed significant effects on most of traits. Aboveground biomass, root-shoot ratio, leaf width, specific leaf area, and leaf nitrogen (N) content were significantly correlated with climate variables under the control condition. Specific leaf area and leaf N content were significantly correlated with climate variables under the drought condition. By PTNs, the hub trait(s) was plant height under the control condition and were aboveground biomass, root length, and specific leaf area under the drought condition. This study indicates that both phenotypic plasticity and genetic differentiation can significantly affect the adaptability of S. krylovii. In addition, soil moisture treatments show significant effects on trait-climate relationships and traits coordinated patterns. These findings provide new insights into the adaptive mechanisms of zonal species in the semiarid grassland region.

Endophytic infection increases the belowground over-yielding effects of the host grass community mainly by increasing the complementary effects.

Introduction: Increases in plant species diversity may increase the community diversity effect and produce community over-yielding. Epichloë endophytes, as symbiotic microorganisms, are also capable of regulating plant communities, but their effects on community diversity effects are often overlooked. Methods: In this experiment, we investigated the effects of endophytes on the diversity effects of host plant community biomass by constructing artificial communities with 1-species monocultures and 2- and 4-species mixtures of endophyte-infected (E+) and endophyte-free (E-) Achnatherum sibiricum and three common plants in its native habitat, which were potted in live and sterilized soil. Results and discussion: The results showed that endophyte infection significantly increased the belowground biomass and abundance of Cleistogenes squarrosa, marginally significantly increased the abundance of Stipa grandis and significantly increased the community diversity (evenness) of the 4-species mixtures. Endophyte infection also significantly increased the over-yielding effects on belowground biomass of the 4-species mixtures in the live soil, and the increase in diversity effects on belowground biomass was mainly due to the endophyte significantly increasing the complementary effects on belowground biomass. The effects of soil microorganisms on the diversity effects on belowground biomass of the 4-species mixtures were mainly derived from their influences on the complementary effects. The effects of endophytes and soil microorganisms on the diversity effects on belowground biomass of the 4-species communities were independent, and both contributed similarly to the complementary effects on belowground biomass. The finding that endophyte infection promotes belowground over-yielding in live soil at higher levels of species diversity suggests that endophytes may be one of the factors contributing to the positive relationship between species diversity and productivity and explains the stable co-existence of endophyte-infected Achnatherum sibiricum with a variety of plants in the Inner Mongolian grasslands.

The relative importance of drought stress and neighbor richness on plant-plant interactions shifts over a short time.

Exploring how plant-plant interactions between species and their neighbors vary with biotic and abiotic factors is vital to understanding community assembly mechanisms in the context of global changes. In this study, using a dominant species (Leymus chinensis (Trin.) Tzvel.) in the semiarid Inner Mongolia steppe as the target species and ten other species as neighbors, we carried out a microcosm experiment to evaluate how drought stress, neighbor richness and season affected the relative neighbor effect (Cint) (the ability to reduce growth of target species). The factor of season interactively affected the effect of drought stress or neighbor richness on Cint. In the summer, drought stress decreased Cint directly and indirectly by decreasing SLA hierarchical distance and neighbor biomass. In the subsequent spring, drought stress increased Cint, and neighbor richness increased Cint directly and indirectly by increasing neighbor community functional dispersion (FDis) and neighbor biomass. Specifically, SLA hierarchical distance was positively associated with neighbor biomass, while height hierarchical distance was negatively associated with neighbor biomass in both seasons, which increased Cint. These findings show that the relative importance of drought stress and neighbor richness on Cint shifted over seasons, which provides strong empirical evidence of how plant-plant interactions respond to changes in biotic and abiotic factors in the semiarid Inner Mongolia steppe over a short-term time. Furthermore, this study provides novel insight into community assembly mechanisms in the context of climatic aridity and biodiversity loss in semiarid regions.

Epichloë Endophyte Infection Changes the Root Endosphere Microbial Community Composition of Leymus Chinensis Under Both Potted and Field Growth Conditions.

Epichloë endophytes can not only affect the growth and resistance of the host plant but also change the biotic and abiotic properties of the soil where the host is situated. Here, we used endophyte-infected (EI) and endophyte-free (EF) Leymus chinensis as plant materials, to study the microbial diversity and composition in the host root endosphere and rhizosphere soil under both pot and field conditions. The results showed that endophyte infection did not affect the diversity of either bacteria or fungi in the root zone. There were significant differences in both bacterial and fungal communities between the root endosphere and the rhizosphere, and between the field and the pot, while endophytes only affected root endosphere microbial communities. The bacterial families affected by endophyte infection changed from 29.07% under field conditions to 40% under pot conditions. In contrast, the fungal families affected by endophyte infection were maintained at nearly 50% under both field and pot conditions. That is to say, bacterial communities in the root endosphere were more strongly affected by environmental conditions, and in comparison, the fungal communities were more strongly affected by species specificity. Endophytes significantly affected the fungal community composition of the host root endosphere in both potted and field plants, only the effect was more obvious in potted plants. Endophyte infection increased the abundance of three fungal families (Thelebolaceae, Herpotrichiellaceae and Trimorphomycetaceae) under both field and potted conditions. In potted plants, endophytes also altered the dominant fungi from pathogenic Pleosporales to saprophytic Chaetomiaceae. Endophyte infection increased the relative abundance of arbuscular mycorrhizal fungi and saprophytic fungi, especially under potted conditions.Overall, endophytes significantly affected the fungal community composition of the host root endosphere in both potted and field plants. Endophytes had a greater impact on root endosphere microorganisms than the rhizosphere, a greater impact on fungal communities than bacteria, and a greater impact on root endosphere microorganisms under potted conditions than at field sites.

Comparative Research on Metabolites of Different Species of Epichloë Endophytes and Their Host Achnatherum sibiricum.

Achnatherum sibiricum can be infected by two species of fungal endophytes, Epichloë gansuensis (Eg) and Epichloë sibirica (Es). In this study, the metabolites of Eg, Es, and their infected plants were studied by GC−MS analysis. The results showed that the metabolic profiles of Eg and Es were similar in general, and only six differential metabolites were detected. The direct effect of endophyte infection on the metabolites in A. sibiricum was that endophyte-infected plants could produce mannitol, which was not present in uninfected plants. Epichloë infection indirectly caused an increase in the soluble sugars in A. sibiricum related to growth and metabolites related to the defense against pathogens and herbivores, such as α-tocopherol, α-linolenic acid and aromatic amino acids. Epichloë infection could regulate galactose metabolism, starch and sucrose metabolism, tyrosine metabolism and phenylalanine metabolism of host grass. In addition, there was a significant positive correlation in the metabolite contents between the endophyte and the host.

Infection by Endophytic Epichloë sibirica Was Associated with Activation of Defense Hormone Signal Transduction Pathways and Enhanced Pathogen Resistance in the Grass Achnatherum sibiricum.

Epichloë endophytes can improve the resistance of host grasses to pathogenic fungi, but the underlying mechanisms remain largely unknown. Here, we used phytohormone quantifications, gene expression analysis, and pathogenicity experiments to investigate the effect of Epichloë sibirica on the resistance of Achnatherum sibiricum to Curvularia lunata pathogens. Comparison of gene expression patterns between endophyte-infected and endophyte-free leaves revealed that endophyte infection was associated with significant induction of 1,758 and 765 differentially expressed genes in the host before and after pathogen inoculation, respectively. Functional analysis of the differentially expressed genes suggested that endophyte infection could activate the constitutive resistance of the host by increasing photosynthesis, enhancing the ability to scavenge reactive oxygen species, and actively regulating the expression of genes with function related to disease resistance. We found that endophyte infection was associated with induction of the expression of genes involved in the biosynthesis pathways of jasmonic acid, ethylene, and pipecolic acid and amplified the defense response of the jasmonic acid/ethylene co-regulated EIN/ERF1 transduction pathway and Pip-mediated TGA transduction pathway. Phytohormone quantifications showed that endophyte infection was associated with significant accumulation of jasmonic acid, ethylene, and pipecolic acid after pathogen inoculation. Exogenous phytohormone treatments confirmed that the disease index of plants was negatively related to both jasmonic acid and ethylene concentrations. Our results demonstrate that endophyte infection can not only improve the constitutive resistance of the host to phytopathogens before pathogen inoculation but also be associated with enhanced systemic resistance of the host to necrotrophs after C. lunata inoculation.

Intraspecific more than interspecific diversity plays an important role on Inner Mongolia grassland ecosystem functions: A microcosm experiment.

Biodiversity changes in terrestrial communities continue in the context of global changes. However, the interactive effects of the changes in diversity at inter- and intraspecific levels as well as cascading effects from plant to soil microorganisms on ecosystem functioning under climate changes remains largely unexplored. Using grassland species in the semi-arid Inner Mongolia Steppe, we conducted a microcosm experiment to assess how drought treatment (non-drought and drought conditions), species diversity (2, 4, and 7 species) and genotypic diversity of the dominant species Leymus chinensis (1, 3, and 6 genotypes) affected ecosystem functions directly or indirectly via regulating plant community functional structure [community-weighted mean (CWM) and functional dispersion (FDis)] and soil microbial diversity (Shannon-Wiener index). Drought treatment, species and genotypic diversity significantly and interactively affected soil N, P cycle and soil multifunctionality as well as soil microbial diversity. Drought treatment significantly affected biomass, soil C cycle, CWM and soil microbial diversity. Species diversity significantly affected soil N cycle, CWM and FDis, and genotypic diversity significantly affected all soil functions and soil microbial diversity. CWM regulated the responses of all ecosystem functions except soil N cycle to the changes in soil moisture and species diversity, which supports the mass ratio hypothesis. The cascading effect from genotypic diversity to soil microbial diversity was significant on belowground biomass but not on any of the other ecosystem functions observed in this study. These findings highlight the importance of genotypic diversity of the dominant species L. chinensis in affecting belowground ecosystem functioning as well as soil microbial diversity, which should not be ignored for grassland protection and management. This study provides further insights into biodiversity and ecosystem functioning mechanisms in semi-arid grasslands in the context of global climate changes.

Soil moisture and species richness interactively affect multiple ecosystem functions in a microcosm experiment of simulated shrub encroached grasslands.

Understanding relationships between biodiversity and ecosystem functions (BEF) and the way in which ecosystem functions respond to changing climatic conditions or community composition is useful for predicting ecological consequences of global changes. However, how soil moisture condition, plant species richness interactively affect ecosystem functions in shrub-encroached grasslands is poorly understood. We conducted a soil moisture × species richness microcosm experiment using semi-arid grassland species with a N-fixing shrub Caragana microphylla Lam. as the dominant species to evaluate how soil moisture and plant species richness affected ecosystem functions directly or indirectly via regulating community functional structure, such as community-weighted mean values (CWM) and functional dispersion (FDis). Soil moisture and species richness interactively affected soil functions (soil C-, N-, P cycles and soil multifunctionality), with greater species richness buffering the adverse effects of soil drought. Soil moisture and species richness showed opposite effects on FDis but similar effects on CWM. FDis mediated the indirect effect of soil moisture and species richness on ecosystem functions, while CWM only mediated the indirect effect of soil moisture. More specifically, both soil moisture and plant species richness were negatively associated with soil P cycle, and the CWMPC1 contributed by traits related to resource-conservative strategy was positively associated with soil N cycle. Species richness showed a positive direct effect on total shoot biomass, which was mainly contributed by the complementarity effect of neighbor species richness. This study provides strong empirical support of how biomass and soil nutrient cycles respond to the changes of soil moisture and plant species richness in C. microphylla shrub-encroached grasslands, and insights on the mechanisms underlying the interactive effects of soil condition and community species composition on multiple ecosystem functions in N-fixing shrub encroached grasslands in semi-arid grassland regions.

A direct comparison of the effects and mechanisms between species richness and genotype richness in a dominant species on multiple ecosystem functions.

Both species (interspecific) richness and genotype (intraspecific) richness of dominant species have significant effects on ecosystem functioning directly or indirectly by regulating plant community functional structure. However, the similarities and differences of the effects between inter- and intraspecific levels are poorly understood. In this study, we selected the main species in the semi-arid Eurasian typical steppe as study objects and simultaneously carried out a species richness experiment and a genotype richness experiment of Stipa grandis which is one of the dominant species in this region. We investigated how plants at each of the two richness levels affected multiple ecosystem functions (biomass, soil C, N and P cycles) directly and indirectly by regulating community functional structure, including community-weighted mean trait values (CWM) and functional dispersion (FDis). Both species richness and genotype richness showed significant direct effects on soil P cycle, and FDis significantly mediated the responses of aboveground biomass and soil N cycle to the changes of species richness and the response of belowground biomass to the changes of genotype richness in S. grandis. CWM showed significant effects on biomass in the species richness experiment and soil nutrient cycles in the genotype richness experiment, independently of the levels of plant richness. These findings provide experimental insights of intraspecific richness effects into the relationships between biodiversity and ecosystem functioning, and highlight the importance of conserving the intraspecific diversity of dominant species in the semi-arid steppe regions.

The Inhibitory Effect of Endophyte-Infected Tall Fescue on White Clover Can Be Alleviated by Glomus mosseae Instead of Rhizobia.

In artificial ecosystems, mixed planting of gramineous and leguminous plants can have obvious advantages and is very common. Due to their improved growth performances and stress tolerance, endophyte-infected grasses are considered to be ideal plant species for grasslands. However, endophytic fungi can inhibit the growth of neighboring nonhost leguminous plants. In this study, we chose endophyte-infected and endophyte-free tall fescue (Lolium arundinaceum Darbyshire ex. Schreb.) and clover (Trifolium repens) as the experimental materials to explore whether arbuscular mycorrhizal fungi and rhizobium can alleviate the inhibitory effect of endophyte infection on clover. The results showed that endophytic fungi significantly reduced clover biomass. Arbuscular mycorrhizal fungi inoculation significantly increased the biomass of clover in both endophyte-infected tall fescue/clover and endophyte-free tall fescue/clover systems but the beneficial contribution of arbuscular mycorrhizal fungi was more obvious in the endophyte-infected tall fescue/clover system. Rhizobia inoculation could alleviate the detrimental effect of tall fescue on the growth of clover but did not alleviate the detrimental effect of endophyte infection on the growth of clover.

Endophytic Fungi Activated Similar Defense Strategies of Achnatherum sibiricum Host to Different Trophic Types of Pathogens.

It is well documented that Epichloë endophytes can enhance the resistance of grasses to herbivory. However, reports on resistance to pathogenic fungi are limited, and their conclusions are variable. In this study, we chose pathogenic fungi with different trophic types, namely, the biotrophic pathogen Erysiphales species and the necrotrophic pathogen Curvularia lunata, to test the effects of Epichloë on the pathogen resistance of Achnatherum sibiricum. The results showed that, compared to Erysiphales species, C. lunata caused a higher degree of damage and lower photochemical efficiency (Fv/Fm) in endophyte-free (E-) leaves. Endophytes significantly alleviated the damage caused by these two pathogens. The leaf damaged area and Fv/Fm of endophyte-infected (E+) leaves were similar between the two pathogen treatments, indicating that the beneficial effects of endophytes were more significant when hosts were exposed to C. lunata than when they were exposed to Erysiphales species. We found that A. sibiricum initiated jasmonic acid (JA)-related pathways to resist C. lunata but salicylic acid (SA)-related pathways to resist Erysiphales species. Endophytic fungi had no effect on the content of SA but increased the content of JA and total phenolic compounds, which suggest that endophyte infection might enhance the resistance of A. sibiricum to these two different trophic types of pathogens through similar pathways.

Arbuscular mycorrhizal fungus identity modulates growth effects of endophyte-infected grasses on neighboring plants.

Endophytes of grasses have been reported to affect the colonization by arbuscular mycorrhizal fungi (AMF) not only of their dual host plant but also of neighboring non-endophyte-infected plants. However, studies investigating the impact of AMF identity on the effects of endophyte-infected grasses on neighboring plants are rare. In this study, we investigated the influence of Leymus chinensis litter type (NL, no litter; E-, endophyte-free litter; E-E+, half E+ and half E- litter; E+, endophyte-infected litter) on Stipa krylovii growth with different AMF species (Claroideoglomus etunicatum, CE; Funneliformis mosseae, FM; Claroideoglomus claroideum, CC; Rhizophagus intraradices, RI). The results showed that the root biomass of S. krylovii tended to decrease with the increase of E+ litter in the mycorrhiza-free treatment. With AMF inoculation, the effects of E+ litter on the AMF colonization rate and root biomass of S. krylovii varied with AMF species. Structural equation modeling (SEM) showed E+ litter could modulate the growth of S. krylovii indirectly via changes in AMF colonization rate, but this effect was related to AMF species.

Removal of Soil Microbes Alters Interspecific Competitiveness of Epichloë Endophyte-Infected over Endophyte-Free Leymus chinensis.

Epichloë endophytes may not only affect the growth and resistances of host grasses, but may also affect soil environment including soil microbes. Can Epichloë endophyte-mediated modification of soil microbes affect the competitive ability of host grasses? In this study, we tested whether Epichloë endophytes and soil microbes alter intraspecific competition between Epichloë endophyte-colonized (EI) and endophyte-free (EF) Leymus chinensis and interspecific competition between L. chinensis and Stipa krylovii. The results demonstrated that Epichloë endophyte colonization significantly enhanced the intraspecific competitive ability of L. chinensis and that this beneficial effect was not affected by soil microbes. Under interspecific competition, however, significant interactions between Epichloë endophytes and soil microbes were observed. The effect of Epichloë endophytes on interspecific competitiveness of the host changed from positive to neutral with soil microbe removal. Here higher mycorrhizal colonization rates probably contributed to interspecific competitive advantages of EI over EF L. chinensis. Our result suggests that Epichloë endophytes can influence the competitive ability of the host through plant soil feedbacks from the currently competing plant species.

Corrigendum: Endophytic Fungi Activated Similar Defense Strategies of Achnatherum sibiricum Host to Different Trophic Types of Pathogens.

[This corrects the article DOI: 10.3389/fmicb.2020.01607.].

Tripartite Interactions Between Endophytic Fungi, Arbuscular Mycorrhizal Fungi, and Leymus chinensis.

Grasses often establish multiple simultaneous symbiotic associations with endophytic fungi and arbuscular mycorrhizal fungi (AMF). Many studies have examined pair-wise interactions between plants and endophytic fungi or between plants and AMF, overlooking the interplays among multiple endosymbionts and their combined impacts on hosts. Here, we examined both the way in which each symbiont affects the other symbionts and the tripartite interactions between leaf endophytic fungi, AMF, and Leymus chinensis. As for AMF, different species (Glomus etunicatum, GE; Glomus mosseae, GM; Glomus claroideum, GC; and Glomus intraradices, GI) and AMF richness (no AMF, single AMF taxa, double AMF mixtures, triple AMF mixtures, and all four together) were considered. Our results showed that significant interactions were observed between endophytes and AMF, with endophytes interacting antagonistically with GM but synergistically with GI. No definitive interactions were observed between the endophytes and GE or GC. Additionally, the concentration of endophytes in the leaf sheath was positively correlated with the concentration of AMF in the roots under low AMF richness. The shoot biomass of L. chinensis was positively related to both endophyte concentration and AMF concentration, with only endophytes contributing to shoot biomass more than AMF. Endophytes and AMF increased shoot growth by contributing to phosphorus uptake. The interactive effects of endophytes and AMF on host growth were affected by the identity of AMF species. The beneficial effect of the endophytes decreased in response to GM but increased in response to GI. However, no influences were observed with other GC and GE. In addition, endophyte presence can alter the response of host plants to AMF richness. When leaf endophytes were absent, shoot biomass increased with higher AMF richness, only the influence of AMF species identity outweighed that of AMF richness. However, when leaf endophytes were present, no significant association was observed between AMF richness and shoot biomass. AMF species identity rather than AMF richness promoted shoot growth. The results of this study demonstrate that the outcomes of interspecific symbiotic interactions are very complex and vary with partner identity such that the effects of simultaneous symbioses cannot be generalized and highlight the need for studies to evaluate fitness response of all three species, as the interactive effects may not be the same for each partner.

Community assembly mechanisms and succession processes significantly differ among treatments during the restoration of Stipa grandis - Leymus chinensis communities.

Understanding community assembly mechanisms is helpful to predict community dynamics. To explore which community assembly mechanism(s) drive(s) the grassland restoration in semi-arid region, we investigated the relationships between plant trait and species relative abundance (SRA), and estimated community functional diversity indices for each community under different treatments (enclosure, grazing and mowing treatment) in a restoration region of Stipa grandis - Leymus chinensis communities in the northern China from 2010 to 2012. There was a high fraction of significant relationships between trait value and SRA, suggesting that niche theory structured the grassland restoration in this region. The functional richness was higher and the functional divergence was lower in the enclosure community than that in the grazing or mowing community, and significantly positive plant height - SRA relationship was found in the enclosure community. These findings demonstrated that limiting similarity based on niche theory was more important in structuring the enclosure community and that environmental filtering based on niche theory played a more important role in driving the grazing or mowing community. Only the factor of year significantly affected the functional evenness (FEve), and the lowest FEve in 2011 implied that the relatively lower precipitation could enhance the effect of limiting similarity on community assembly in the semi-arid grassland.

Litter species diversity is more important than genotypic diversity of dominant grass species Stipa grandis in influencing litter decomposition in a bare field.

Studies have indicated that plant litter diversity can affect litter decomposition at both species diversity and genotypic diversity level within a species. However, the essence and relative importance of these two diversity levels on litter decomposition remain unknown. Here, two independent one-factor experiments, litter species diversity and litter genotypic diversity of the dominant species-Stipa grandis, were carried out to explore the effects of initial litter quality, litter composition and diversity on decomposition of mass, nitrogen (N), carbon (C) and phosphorus (P) simultaneously. The results showed that: (1) there were significant relationships between the initial litter N, C/N, lignin/N and the decomposition rate of N, between the initial litter P, N/P and the decomposition rate of P, and the litter composition significantly influenced litter mass, N, C and P remaining in both litter species and genotypic diversity experiments; and (2) litter species diversity significantly affected litter mass, N, C and P remaining, and non-additive relative mixture effects were mainly contributed by synergistic effects especially in 6-species mixtures; however, similar patterns were not found in litter genotypic diversity experiment. The present results emphasized that initial litter quality played the most important role in influencing litter decomposition of mass N, C and P, and suggested that litter species mixtures rather than litter genotypic mixtures of a dominant species could favor nutrient cycling in ecosystem of the semi-arid Inner Mongolia Steppe of China.

Endophyte Infection and Methyl Jasmonate Treatment Increased the Resistance of Achnatherum sibiricum to Insect Herbivores Independently.

Alkaloids are usually thought to be responsible for protecting endophyte-infected (EI) grasses from their herbivores. For EI grasses that produce few alkaloids, can endophyte infection enhance their resistance to herbivores? Related studies are limited. In the Inner Mongolian steppe, Achnatherum sibiricum is highly infected by Epichloë endophytes, but produces few alkaloids. Locusts are the common insect herbivores of grasses. In this study, A. sibiricum was used as plant material. Methyl jasmonate (MJ, when applied exogenously, can induce responses similar to herbivore damage) treatment was performed. The effects of endophyte infection and MJ treatment on the resistance of A. sibiricum to Locusta migratoria were studied. We found that locusts preferred EF (endophyte-free) plants to EI plants in both choice and no-choice feeding experiments. Endophyte infection enhanced the resistance of A. sibiricum to locusts. Endophyte infection decreased soluble sugar concentrations, while it increased the total phenolic content and phenylalanine ammonia lyase (PAL) activity, which may contribute to the resistance of A. sibiricum to locusts. There was an interaction effect between MJ treatment and endophyte infection on the growth of the host. MJ treatment was a negative regulator of the plant growth-promoting effects of endophyte infection. There was no interaction effect between MJ treatment and endophyte infection on the defense characteristics of the host. In groups not exposed to locusts, MJ treatment and endophyte infection had a similar effect in decreasing the soluble sugar content, while increasing the total phenolic content and the PAL activity. In groups exposed to locusts, the effect of MJ treatment on the above characteristics disappeared, while the effect of endophyte infection became more obvious. All of these results suggest that even for endophytes producing few alkaloids, they could still increase the resistance of native grasses to insect herbivores. Furthermore, endophyte infection might mediate the defense responses of the host, independent of jasmonic acid (JA) pathways.