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.

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.

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.

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.