Underestimated ecosystem carbon turnover time and sequestration under the steady state assumption: a perspective from long‐term data assimilation

It is critical to accurately estimate carbon (C) turnover time as it dominates the uncertainty in ecosystem C sinks and their response to future climate change. In the absence of direct observations of ecosystem C losses, C turnover times are commonly estimated under the steady state assumption (SSA), which has been applied across a large range of temporal and spatial scales including many at which the validity of the assumption is likely to be violated. However, the errors associated with improperly applying SSA to estimate C turnover time and its covariance with climate as well as ecosystem C sequestrations have yet to be fully quantified. Here, we developed a novel model-data fusion framework and systematically analyzed the SSA-induced biases using time-series data collected from 10 permanent forest plots in the eastern China monsoon region. The results showed that (a) the SSA significantly underestimated mean turnover times (MTTs) by 29%, thereby leading to a 4.83-fold underestimation of the net ecosystem productivity (NEP) in these forest ecosystems, a major C sink globally; (b) the SSA-induced bias in MTT and NEP correlates negatively with forest age, which provides a significant caveat for applying the SSA to young-aged ecosystems; and (c) the sensitivity of MTT to temperature and precipitation was 22% and 42% lower, respectively, under the SSA. Thus, under the expected climate change, spatiotemporal changes in MTT are likely to be underestimated, thereby resulting in large errors in the variability of predicted global NEP. With the development of observation technology and the accumulation of spatiotemporal data, we suggest estimating MTTs at the disequilibrium state via long-term data assimilation, thereby effectively reducing the uncertainty in ecosystem C sequestration estimations and providing a better understanding of regional or global C cycle dynamics and C-climate feedback

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

Endophyte-mediated effects on the growth and physiology of Achnatherum sibiricum are conditional on both N and P availability.

The interaction of endophyte-grass associations are conditional on nitrogen (N) availability, but the reported responses of these associations to N are inconsistent. We hypothesized that this inconsistency is caused, at least in part, by phosphorus (P) availability. In this experiment, we compared the performance of endophyte-infected (EI) and endophyte-free (EF) Achnatherum sibiricum subjected to four treatments comprising a factorial combination of two levels of N (N+ vs. N-, i.e. N supply vs. N deficiency) and two levels of P (P+ vs. P-, i.e. P supply vs. P deficiency) availability. The results showed that A. sibiricum-Neotyphodium associations were conditional on both N and P availability, but more conditional on N than P. Under N+P- conditions, endophyte infection significantly improved acid phosphatase activity of EI plants, such that the biomass of EI plants was not affected by P deficiency (i.e. similar growth to N+P+ conditions), and resulted in more biomass in EI than EF plants. Under N-P+ conditions, biomass of both EI and EF decreased compared with N+P+; however, EI biomass decreased slowly by decreasing leaf N concentration more rapidly but allocating higher fractions of N to photosynthetic machinery compared with EF plants. This change of N allocation not only improved photosynthetic ability of EI plants but also significantly increased their biomass. Under N-P- conditions, EI plants allocated higher fractions of N to photosynthesis and had greater P concentrations in roots, but there was no significant difference in biomass between EI and EF plants. Our results support the hypothesis that endophyte-grass interactions are dependent on both N and P availability. However, we did not find a clear cost of endophyte infection in A. sibiricum

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

Effect of Endophyte infection and clipping treatment on resistance and tolerance of Achnatherum sibiricum

It is well-documented that endophytes can enhance the resistance of agronomical grasses, such as tall fescue and perennial ryegrass to herbivory. For native grasses, however, the related reports are limited, and the conclusions are variable. Achnatherum sibiricum is a grass native to the Inner Mongolian steppe. This grass is highly infected by endophytes but does not produce detectable endophyte-related alkaloids known under normal conditions. In this study, the contributions of endophytes to the resistance of A. sibiricum to Locusta migratoria were studied. We found that locusts preferred EF (endophyte-free) plants to EI (endophyte-infected) plants, and the weight of locusts fed on EI plants was significantly lower than those fed on EF plants. Hence, endophyte infection significantly enhanced the resistance of the host to L. migratoria. Endophyte infection significantly decreased the concentration of soluble sugar and amino acids while significantly increasing the concentration of total phenolic content, and these metabolites may contribute to herbivore resistance of the host. The clipping treatment further strengthened the locust resistance advantage of EI over EF plants. After clipping, the weight of the locusts fed on EI plants significantly decreased compared with those fed on unclipped plants, whereas the weight of the locusts fed on EF plants increased significantly. The results suggested that endophyte infection could increase herbivore resistance while decreasing the tolerance of the host grass by mechanisms apart from endophyte-conferred alkaloid defense

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

Abstract 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

Both evenness and dominant species identity have effects on litter decomposition

Abstract 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

Comparative Research on Metabolites of Different Species of <i>Epichloë</i> Endophytes and Their Host <i>Achnatherum sibiricum</i>

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

Three-way ANOVA for photosynthetic parameters, N allocation and acid phosphatase activity of endophyte-infected (EI) or uninfected (EF) ramets of <i>Achnatherum sibiricum</i>.

<p>Note. N_A, total leaf nitrogen content; P_max, maximum net photosynthetic rate; PNUE, photosynthetic nitrogen use efficiency; P_T the fraction of leaf nitrogen allocated to all components of the photosynthetic machinery.</p